The SAGE Handbook of

Evolutionary Psychology
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 The SAGE Handbook of
Evolutionary Psychology

Applications of Evolutionary Psychology

Edited by
Todd K. Shackelford
SAGE Publications Ltd
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Editorial arrangement © Todd K. Shackelford, 2021
Chapter 1 © Cezar Giosan, 2021 Cody Jorgensen and Jessica
Chapter 2 © Riadh Abed and Wells, 2021
Paul St John-Smith, 2021 Chapter 14 © Ian A. Silver and
Chapter 3 © John F. Gunn III, Jamie Newsome, 2021
Pablo Malo and C. A. Soper, 2021 Chapter 15 © Russil Durrant, 2021
Chapter 4 © James Carmody, Chapter 16 © Anthony C. Lopez,
2021 2021
Chapter 5 © Ulysses Paulino Chapter 17 © Holly Wilson, Paul
Albuquerque, Joelson M. B. Rauwolf and Joanna J. Bryson,
Moura, Risoneide Henriques da 2021
Silva, Washington S. Ferreira Chapter 18 © Robert Finkelstein,
Júnior and Taline C. Silva, 2021 2021
Chapter 6 © Simon Russell, 2021 Chapter 19 © Deanna Singhal
Chapter 7 © Diana Santos and David Wiesenthal, 2021
Fleischman, 2021 Chapter 20 © Gayle S. Stever,
Chapter 8 © Michael Latner and 2021
Elissa Feld, 2021 Chapter 21 © Ned Kock, 2021
Chapter 9 © Joseph L. Nedelec, Chapter 22 © Mark A. Caudell,
2021 2021
Chapter 10 © Lois James, 2021 Chapter 23 © James R.
Chapter 11 © Eyal Aharoni and Anderson, 2021
Morris B. Hoffman, 2021 Chapter 24 © Stephen Whyte and
Chapter 12 © Alina Simona Rusu, Benno Torgler, 2021
2021 Chapter 25 © Robert Finkelstein,
Chapter 13 © Anthony Walsh, 2021
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Library of Congress Control Number: 2020947010
British Library Cataloguing in Publication data
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978-1-5264-8916-6
 Contents

List of Figures, Tables and Boxes vii

Notes on the Editor and Contributors ix

PART 1  INTEGRATION WITHIN PSYCHOLOGY

1. Evolutionary Psychology and Counseling and Psychotherapy 3

Cezar Giosan

2. Evolutionary Psychology and Psychiatry 24

Riadh Abed and Paul St John-Smith

3. Evolutionary Psychology and Suicidology 51

John F. Gunn III, Pablo Malo, and C. A. Soper

4. Evolutionary Psychology and Mindfulness and Meditation:

Easing the Anxiety of Being Human 94
James Carmody

5. Evolutionary Psychology and Environmental Sciences 107

Ulysses Paulino Albuquerque, Joelson M. B. Moura, Risoneide
Henriques da Silva, Washington S. Ferreira Júnior, and Taline C. Silva

6. Evolutionary Psychology and Public Health 123

Simon Russell

7. Animal Ethics and Evolutionary Psychology 144

Diana Santos Fleischman

PART 2  APPLICATIONS TO LAW AND ORDER

8. Evolutionary Psychology and Political Institutions 171

Michael Latner and Elissa Feld

9. Evolutionary Psychology and Crime 188

Joseph L. Nedelec

10. Evolutionary Psychology and Policing: The Balance Between

Aggression and Restraint 203
Lois James

11. Evolutionary Psychology, Jurisprudence, and Sentencing 221

Eyal Aharoni and Morris B. Hoffman
vi THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

12. Evolutionary Psychology and Incarceration 243

Alina Simona Rusu

13. Evolution and Punishment 258

Anthony Walsh, Cody Jorgensen, and Jessica Wells

14. Evolutionary Psychology and Corrections and Rehabilitation 277

Ian A. Silver and Jamie Newsome

15. Evolutionary Psychology and Organized Crime 296

Russil Durrant

16. Evolutionary Psychology and Warfare 316

Anthony C. Lopez

PART 3 APPLICATIONS TO TECHNOLOGY, COMMUNICATIONS,

AND THE FUTURE

17. Evolutionary Psychology and Artificial Intelligence: The Impact of

Artificial Intelligence on Human Behaviour 333
Holly Wilson, Paul Rauwolf, and Joanna J. Bryson

18. Evolutionary Psychology and Robotics 352

Robert Finkelstein

19. Evolutionary Psychology and Dangerous Driving Behaviour 374

Deanna Singhal and David Wiesenthal

20. Evolutionary Psychology and Mass Media 398

Gayle S. Stever

21. Evolutionary Psychology and Communication 417

Ned Kock

22. Evolutionary Psychology and Climate Change: Understanding the Impact

of Time Perspective on Carbon Emissions across 75 Countries 435
Mark A. Caudell

23. Evolutionary Psychology and Thanatology: Responses to Death 457

James R. Anderson

24. Evolutionary Psychology and Reproduction 477

Stephen Whyte and Benno Torgler

25. Evolutionary Psychology and Cyberwarfare 499

Robert Finkelstein

Index515
 List of Figures, Tables and Boxes

FIGURES

2.1 Life history strategy trade-offs 29

3.1 Summary of posited antisuicide defences 68
3.2 A tentative mapping of hypothesized types of antisuicide mechanisms
(keepers) across common diagnostic categories of mental disorder 74
4.1 Alarm-related components of experience forming a cycle of distress 97
4.2 Memory, imagination and emotion are symphonies of three interwoven
experiential components 99
4.3 Components recognized as differentiated and connected 100
4.4 Attention shifts from differentiated components to arousal-neutral sensations
of breathing 101
4.5 Re-perceiving reduces distress through a perceptual/attentional shift from
what the thought is about – ‘I’m going to pass out’ – to the thought as
an event in the mind/awareness – ‘This is a thought’ 101
5.1 Environment of Evolutionary Adaptedness definition (EEA), original version
and extended version 110
5.2 Structure of the human naturalist mind 112
6.1 The impact of the food supply on the expression of obesity 125
6.2 Key determinants of psychological mechanisms and behaviour strategies 129
7.1 Charitable donations towards animal organizations as compared to animal use 151
9.1 Age-specific birth rates (per 1,000 women per year) in the 10 neighborhoods
with the longest life expectancy compared to the 10 neighborhoods with
the shortest life expectancy 198
10.1 Police balance between aggression and restraint 213
15.1 The basic dimensions of organized crime 298
19.1 GAM factors included in the investigation of modelling of aggressive or
risky driving 384
19.2 Domestic box-office history for the Fast and Furious movies389
21.1 Path model showing a costly trait and its relationship with fitness 421
21.2 Probability of evolution of a new costly trait 423
21.3 Variation of the expected ratio between pay and pax424
21.4 The evolution of oral speech in humans 425
22.1 Effect of time orientation on total fertility as a function of education 446
22.2 Effect of time orientation on total fertility as a function of mortality level 447
22.3 Effect of time orientation on carbon emissions per capita as a function
of investment in education 448
viii THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

TABLES

2.1 Tinbergen’s four questions 27

3.1 The fitness costs of suicide 54
7.1 Days of suffering per kilogram of food weight produced by the animal adjusted
for the badness of each day of life as estimated by animal welfare researchers 158
15.1 Approaches to defining organized crime 297
15.2 The functional structures of organized crime groups in relation to the key
evolutionary processes that facilitate cooperation 305
18.1 Processes of mind and their computational equivalents 354
21.1 Oral speech and the two common characteristics of costly traits 426
22.1 Descriptive statistics for Model 1 441
22.2 Variable correlations for Model 1 441
22.3 Data sources, years, and future orientation scores 443
22.4 Descriptive statistics for Model 2 445
22.5 Variable correlations for Model 2 445
22.6 Relationship between total fertility rates, time orientation, mortality,
and parental investment 446
22.7 Effect of time orientation and education on carbon emissions 448

BOX

2.1 Pathways for the persistence of disease and disorder 28

2.2 Evolutionary theories of depression 33
5.1 Structure and behavior of the human naturalistic mind 113
 Notes on the Editor
and Contributors
THE EDITOR

Todd K. Shackelford received his Ph.D. in evolutionary psychology in 1997 from the University
of Texas at Austin. Since 2010, he is Professor and Chair of the Department of Psychology at
Oakland University in Rochester, Michigan, where he is Co-Director of the Evolutionary
Psychology Lab. In 2016, he was appointed Distinguished Professor by the Oakland University
Board of Trustees. He led the founding of new Ph.D. and M.S. programs which launched in
2012. Shackelford has published around 300 journal articles and his work has been cited over
22,000 times. Much of Shackelford’s research addresses sexual conflict between men and
women, with a special focus on men’s physical, emotional, and sexual violence against their
intimate partners. Since 2006, Shackelford has served as editor of the journal Evolutionary
Psychology, and in 2014 founded the journal Evolutionary Psychological Science as
Editor-in-Chief.

THE CONTRIBUTORS

Riadh Abed FRCPsych, Qualified in medicine from Baghdad and trained in psychiatry in the
UK. Retired Psychiatrist, Medical Director and Hon. Senior Clinical Lecturer, University of
Sheffield, UK. Currently is a medical member of Mental Health Tribunals, Ministry of Justice,
UK. He is author of a number of novel evolutionary hypotheses on eating disorders, obsessive
compulsive disorder and schizophrenia and has published both theoretical as well as research
articles on a range of evolutionary psychiatry subjects. He is founding chair of the Evolutionary
Psychiatry Special Interest Group (EPSIG) at the Royal College of Psychiatrists, UK.

Eyal Aharoni is an Associate Professor of Psychology, Philosophy, and Neuroscience at

Georgia State University in Atlanta. His research investigates violence risk assessment and the
influence of emotion, cognitive bias, and other extra-legal factors on legal decision-making.
Prior to his current appointment, Aharoni served as a Research Associate for the RAND
Corporation. He completed a postdoctoral fellowship with appointments at The MIND Research
Network for Neurodiagnostic Discovery and the University of New Mexico Psychology. He
has also held research positions at the Research Center for Virtual Environments and Behavior
and the Institute for Social, Behavioral, and Economic Research. Aharoni earned his PhD in
psychology at UC Santa Barbara where he also served as a research fellow for the MacArthur
Foundation’s Law and Neuroscience Project.

Ulysses Paulino Albuquerque received his Ph.D. in biology in 2001 from the Universidade
Federal de Pernambuco, Brazil. He is Full Professor of the Department of Botany at Universidade
Federal de Pernambuco, Pernambuco, Brazil. In 2011, he led the founding of new Ph.D. program
in Ethnobiology and Nature Conservation. Albuquerque has published around 316 journal articles,
x THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

200 chapter’s book and edited or authored 50 books (including new editions and translations).
Much of actual Albuquerque’s research addresses how work our naturalistic mind. Albuquerque
has served as editor of various peer-reviewed journals, and in 2011 co-founded the journal
Ethnobiology and Conservation as Co-Editor-in-Chief.

James R. Anderson is a Professor in Psychology at Kyoto University, Japan. For more than
30 years he has studied behaviour and cognition in a wide range of nonhuman primates in
captivity in the wild, as well as human children and elderly people. Most of his research is in
the following areas: the effects of early experience on social adjustment, the development of
abnormal behaviour, improving welfare of captive primates through environmental enrichment,
communicative behaviours including tactile and postural signals, gaze-following, feeding
competition and the influence of dominance relationships, tool-use, the ontogeny and phylogeny
of self-recognition, self-control, third-party social evaluations, sleep-related behavioural
adaptations, and responses to dead and dying group members and other individuals.

Joanna J. Bryson is recognised for broad expertise on intelligence and its impacts, advising
governments, transnational agencies and NGOs globally. She holds two degrees each in
psychology and AI (BA Chicago, MSc and MPhil Edinburgh, PhD MIT). From 2002–2019
she was Computer Science faculty at the University of Bath; she has also been affiliated
with Harvard Psychology, Oxford Anthropology, Mannheim Social Science Research and
the Princeton Center for Information Technology Policy. During her PhD she observed the
confusion generated by anthropomorphised AI, leading to her first AI ethics publication ‘Just
Another Artifact’ in 1998. She has remained active in the field including coauthoring the
first national-level AI ethics policy, the UK’s (2011) Principles of Robotics. She is presently
Professor of Ethics and Technology at the Hertie School of Governance in Berlin, Germany.

James Carmody is a Professor in the Departments of Medicine, and Population and Health
Sciences Division of Preventive and Behavioral Medicine at University of Massachusetts Medi-
cal School. His research is on how evolutionary and biological imperatives shape our habits of
attending to experience, their effect on anxiety, and the degree to which they can be mitigated
by mind–body training programmes like mindfulness and yoga. He is principal investigator
on NIH-funded clinical trials of the clinical effects and mechanisms of mindfulness training.
James studied and practised in Zen, Tibetan, Theravada and Advaita traditions in several coun-
tries over 50 years. He has been a therapist and leads retreats for clinicians with the goal of
making the conceptualisation and psychological mechanisms of mindfulness straightforward,
jargon-free and practically accessible for patients. His work has been featured in national and
international media including the New York Times, NPR and ABC.

Mark A. Caudell is a Medical Anthropologist working for the Food and Agriculture Organiza-
tion of the United Nations and is adjunct faculty in the Department of Anthropology at Wash-
ington State University. His research centres on understanding how socioecological systems
drive the emergence and transmission of environmental risks as well as pattern individual and
community responses to these risks. His current work combines ethnographic and lab-based
methods to identify the cultural and cognitive drivers of infectious disease and antimicrobial
resistance in low- and middle-income countries. He has conducted research among foragers in
the Congo Basin and farmers and pastoralists throughout sub-Saharan Africa and is funded by
the National Science Foundation, The Fleming Fund, Amazon and the Paul G. Allen School for
Global Animal Health at Washington State University. Currently, he is based in Nairobi, Kenya.
 Notes on the Editor and Contributors xi

Risoneide Henriques da Silva is master in Botany (2018) from the Federal Rural University of
Pernambuco, Brazil. She is currently PhD candidate in Ethnobiology and Nature Conservation
by the same institution. Since 2016 she is a researcher associated at the Laboratory of Ecology
and Evolution of Social-ecological Systems at the Federal University of Pernambuco, Brazil.
Risoneide Henriques has an interest in understanding the cognitive aspects involved with
human behaviour in relation to biota.

Russil Durrant is a Senior Lecturer at the Institute of Criminology, Victoria University of

Wellington, New Zealand, where he teaches courses in criminal, biosocial and investigative
psychology. His research focuses on the application of evolutionary theory to understanding
crime and crime-related phenomena. He is the author (or co-author) of five books including
(with Tony Ward) Evolutionary Criminology: Towards a Comprehensive Explanation
for Crime and (with Zoe Poppelwell) Religion, Crime, and Punishment: An Evolutionary
Perspective.

Elissa Feld earned a Masters in Public Policy at California Polytechnic State University, San
Luis Obispo. Her research continues to focus on political psychology and public policy.

Robert Finkelstein earned a DBA in Systems Theory and Cybernetics from George
Washington University (GWU), an ApSci. in Operations Research (GWU), an MS in
Operations Research (GWU), an MS in Physics (University of Massachusetts) and a BA in
Physics (Temple University), as well as completing postgraduate courses in Physics at MIT. Dr
Finkelstein is President, since 1985, of Robotic Technology Inc., a professional services firm
that provides technical analyses, systems engineering, technology assessments, operations
research, business development and other services in the fields of autonomous ground, air and
water vehicles, intelligent systems, robotics for military and civil applications, and memetics.
Dr Finkelstein is an Adjunct Professor in the graduate school at the University of Maryland
Global Campus and is serving as a Co-Director of the Intelligent Systems Laboratory at the
University of Maryland Clark School of Engineering.

Diana Santos Fleischman is an Associate Professor of psychology at the University of

Portsmouth in the UK. She received her PhD in evolutionary psychology at the University
of Texas at Austin. Diana has been involved in animal ethics and effective animal advocacy
for several years. From 2010–2012 Diana co-hosted ‘The Vegan Option’ podcast which
explored many themes at the intersection of psychology and animal ethics. From 2017–
2019 Diana served on the board at Sentience Institute, a think tank devoted to exploring
ways to expand humanity’s moral circle. Diana also has served on the grants board of the
Animal Advocacy Research Fund which aims to promote research into effective animal
advocacy.

Cezar Giosan PhD is a faculty member of the Department of Psychology at the University of
Bucharest and has a doctorate in psychology from the New School University, New York. He
worked for a long time as a research psychologist in the top-ranked Department of Psychiatry
at Weill Medical School of Cornell University. Dr Giosan has authored dozens of peer-reviewed
publications in impact journals, including studies on the applications of evolutionary psychology
in psychological interventions. He has tested, in the form of a randomized clinical trial, the
efficacy of an evolutionary intervention for depression, and is the author of a published therapy
protocol of Cognitive-Evolutionary Therapy.
xii THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

John F. Gunn III PhD has a BA (Stockton University) and MA (Rutgers University – Camden)
in psychology and a PhD in family science and human development (Montclair State Univer-
sity). John is currently a Postdoctoral Associate at the NJ Center on Gun Violence Research at
Rutgers University, where he is investigating the role of firearms in suicide mortality on both
small and large scales. His research interests include risk and protective factors for suicidal
behaviour (such as access to a firearm; exposure to bullying), social connectedness and sui-
cide, theoretical models of suicidal behaviour (such as evolutionary explanations for suicide)
and the impact of media on suicide. John has written or co-authored over 20 scholarly publica-
tions, three edited books and 10 book chapters.

Morris B. Hoffman has been a district judge in the state of Colorado since 1991. He is a
member of the John D. and Catherine T. MacArthur Foundation’s Research Network on Law
and Neuroscience, and a Research Fellow at the Gruter Institute for Law and Behavioral
Research. He has published many law review articles, focusing on the areas of criminal law,
jurisprudence, law and neuroscience, and jury history, in journals that include Stanford,
University of Chicago, Vanderbilt, Duke, Northwestern, the University of Pennsylvania,
Journal of Law and Biosciences and Journal of Philosophy, Science and Law. He has co-
authored science papers in journals that include Proceedings of the National Academy of
Sciences, Philosophical Transactions of the Royal Society B, Journal of Neuroscience, Trends
in Neuroscience and Social Neuroscience. His book, The Punisher’s Brain: An Evolutionary
History of Judge and Jury, was published in 2014.

Lois James is an Assistant Professor in the Washington State University (WSU) College of
Nursing, where she focuses on bias, stress, sleep and performance in ‘high-stress’ populations
such as police officers, military personnel, nurses and top-tier athletes. She has received
multiple honours and awards for her work, and is internationally recognised as a leading expert
in her field. Her simulation-based research on the impact of bias on police decision making has
significantly advanced what is known about how suspect race and ethnicity influences police
officers during critical encounters with the public. She is the founding director of Counter
Bias Training Simulation (CBTsim), a novel and innovative simulation-based implicit bias
training program that has been featured in National Geographic and the recent feature-length
documentary ‘bias’. Her work has been published extensively in academic journals, practitioner
magazines and mainstream media such as the New York Times and the Washington Post.

Cody Jorgensen is an Assistant Professor at Boise State University. He received his PhD from
the University of Texas at Dallas in 2014. His main research interests include biosocial and
developmental criminology, policing and forensics, and drug policy.

Washington S. Ferreira Júnior received his Ph.D in Botany in 2015 at Federal Rural
University of Pernambuco. Since 2016, he is Professor of the Biological Sciences course at
Pernambuco University in Petrolina, Pernambuco, where he is coordinator of the Laboratory
of Biocultural Investigations in the Semiarid. He is professor at the Postgraduate Program in
Environmental Science and Technology at University of Pernambuco and the Postgraduate
Program in Ethnobiology and Nature Conservation at Federal Rural University of Pernambuco.
Ferreira Júnior has published around 40 journal articles, 40 book chapters and organized three
books with partners. His research efforts are focused on understanding the structure, dynamics
and evolution of local medical systems with an emphasis on the use of medicinal plants.
 Notes on the Editor and Contributors xiii

Ned Kock is Texas A&M Regents Professor and Chair of the Division of International Busi-
ness and Technology Studies, in the Sanchez School of Business, at Texas A&M International
University. He is an active member of the Human Behavior and Evolution Society and the
Association for Information Systems. He has edited the Springer book Evolutionary Psychol-
ogy and Information Systems Research, and guest-edited the Special Issue on Darwinian Per-
spectives on Electronic Communication published in the IEEE Transactions on Professional
Communication. Ned has published his research in a number of high-impact journals including
Communications of the ACM, Decision Support Systems, European Journal of Information
Systems, European Journal of Operational Research, IEEE Transactions (various), Informa-
tion & Management, Information Systems Journal, Journal of the Association for Informa-
tion Systems, Journal of Management Information Systems, MIS Quarterly and Organization
Science. His research interests include evolution and human behaviour toward technology,
computational statistics and structural equation modelling.

Michael Latner is a Professor of Political Science at California Polytechnic State University,

San Luis Obispo, faculty scholar at Cal Poly’s Institute for Advanced Technology and Public
Policy and a Senior Fellow with the Union of Concerned Scientists’ Center for Science and
Democracy. His research focuses on electoral systems, coalition behavior, representation and
voting rights.

Anthony C. Lopez is Associate Professor of Political Psychology in the School of Politics,

Philosophy, and Public Affairs, at Washington State University. He received his PhD from Brown
University, and also received training as a research associate at the Center for Evolutionary
Psychology, University of California, Santa Barbara. His research explores the evolutionary
origins of warfare, as well as its constituent psychological dynamics, such as revenge, the
nature of deterrence, collective action problems of warfare, extremist violence, and distinctions
between offensive and defensive aggression. He serves as Associate Editor of Politics with the
Evolution Institute and blogs regularly at Psychology Today.

Pablo Malo studied medicine in the University of the Basque Country in Spain and psychiatry
in the Psychiatric Hospital of Zamudio. Pablo is currently a clinical psychiatrist working in
the Mental Health Center of Bombero Etxaniz, in Bilbao, Spain. His clinical interests include
psychosis and severe mental disorders, violence and mental health disorders, intimate partner
violence and evolutionary psychology. Pablo has co-authored a book about evolutionary
psychiatry, written over 20 scholarly papers and is the editor of two blogs about evolutionary
psychology and biology.

Joelson M. B. Moura has a complete degree in Biological Sciences from the Universidade
Federal Rural de Pernambuco (UFRPE), with an internship at the University of Coimbra (UC)
in 2013. He received his master’s degree in Ethnobiology and Nature Conservation (UFRPE)
in 2018. He is currently a Ph.D. candidate in the same program and integrates the Laboratory
of Ecology and Evolution of Socioecological Systems (LEA) at the Universidade Federal de
Pernambuco (UFPE) since 2016. The research carried out by Moura seeks to understand the
processes that contributed to the evolution of the human mind. His work is in the perspective
of evolutionary ethnobiology, which addresses the influence of the evolutionary past on
current human behavior and how it reflects on the relationship between people and their
environments.
xiv THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

Joseph L. Nedelec is an Associate Professor in the School of Criminal Justice at the University
of Cincinnati. His research interests include biosocial criminology, evolutionary psychology,
intelligence, quantitative behaviour genetics, and cybercrime. Dr Nedelec’s work has been
published in a variety of journals including Criminology, Evolution and Human Behavior,
Intelligence, Journal of Criminal Justice, Journal of Quantitative Criminology, Personality and
Individual Differences and PLOS One, among others.

Jamie Newsome received her doctorate in 2013 from the School of Criminal Justice at UC,
and is currently the Research Coordinator for UCCI. Her research interests include the design,
implementation and effectiveness of correctional interventions, and translating research into
practice.

Paul Rauwolf is an Assistant Professor (Lecturer) in Psychology at Bangor University, UK.

Previously, he was a Postdoctoral Researcher in the Mathematics Institute at the University
of Oxford and the School of Psychology at Bangor University. In 2016, he received his PhD
in Computer Science from the University of Bath. His research takes an interdisciplinary
perspective to better understand the utility and aetiology of biases in human decision-making.
During his PhD, he used agent-based modelling to demonstrate that ignoring freely available
information can be adaptive in a variety of contexts, such as maintaining cooperative societies.
His current work focuses on understanding how the changing information and technological
landscape impacts human decision-making under uncertainty.

Simon Russell is a Research Fellow at the Great Ormond Street Institute of Child Health. Simon
has conducted research in the fields of evolutionary science and public health, with a particular
interest in health risk behaviours, obesity, inequalities in health, public health policy and
epidemiology. Simon’s PhD entailed applying evolutionary principles to the obesity problem
and other issues in public health, work that demonstrated that health behaviours and strategies
are likely to be adaptive. Currently, Simon leads various research projects that have high policy
relevance and often involve emerging methodologies. In recent years, Simon has specialised in
simulating policy relevant interventions using causal inference techniques across longitudinal
cohort data. Simon’s work involves applying epidemiological techniques to explore causes and
consequences of obesity and to appraise the effectiveness of potential population policy action.

Alina Simona Rusu is a biologist and psychologist and is currently an Associate Professor in
the Department of Special Education, School of Psychology and Sciences of Education, Babeş-
Bolyai University, Romania. She received her PhD in Animal Behaviour from University of Zurich,
Switzerland. She is interested in interdisciplinary research of human prosocial behaviour in the
context of inclusive institutions, as well as the applied values of human–animal interactions.

Taline C. Silva received her Ph.D. in Botany in 2014 from the Federal Rural University of
Pernambuco. Since 2016, she is Professor of the Biology course at State University of Alagoas
in Palmeira dos índios, Alagoas, where she is coordinator of the Ethnobiology and Ecossistems
conservation Lab. In 2018, she was member of Post graduation program in ethnobiology and
nature conservation. Silva has published around 30 journal articles. She has experience in
Ethnobiology and seeks to understand the factors that guide the complex relationship between
people and nature, accessing knowledge, use and local perception of natural resources and
landscape changes.
 Notes on the Editor and Contributors xv

Ian A. Silver is an Assistant Professor at the Law and Justice Department at Rowan University
and a Fellow at the University of Cincinnati Corrections Institute (UCCI). His research
interests include correctional rehabilitation, incarceration, biopsychosocial criminology,
and methodological issues. His recent work has appeared in Criminology and Public Policy,
Aggressive Behavior, Journal of Offender Rehabilitation, Brain Injury and Journal of Criminal
Justice.

Deanna Singhal received a bachelor’s degree in psychology from Queen’s University in

Kingston, and a master’s degree (exercise and health science) and doctoral degree (psychology:
brain, behaviour, and cognitive science) from York University in Toronto. She is now a Faculty
Service Officer in the Department of Psychology at the University of Alberta, Canada, with
a primary role to support the Undergraduate Teaching and Learning Program. Her research
interests include driving behaviour and cognitive workload, and her areas of teaching include
introductory psychology and human neuropsychology.

Paul St John-Smith FRCPsych. Trained at Oxford, graduated in natural sciences in 1976

and qualified in medicine in 1979. He then trained as a general practitioner and later joined
the pharmaceutical industry for five years as a research physician in psychopharmacology
investigating various antidepressants, hypnotics and other central nervous system agents
including the benzodiazepine antagonist. In 1987 he returned to study psychiatry full time and
worked in a number of London hospitals including Barnet, Guys, Greenwich, St Thomas’, and as
a consultant in Hertfordshire. He has published on a range of areas in evolutionary psychiatry
and has peer reviewed papers on psychopharmacology, placebo effects, professionalism, suicide
and depression, as well as evolution.

C. A. Soper is a psychotherapist in private practice in Lisbon, Portugal. He holds degrees from

the University of Cambridge and the University of London and gained his PhD at the University
of Gloucestershire for theoretical research into the evolutionary origins of suicide. His book,
The Evolution of Suicide, was published in 2018. He has authored various other scholarly
publications on suicide and suicidology, and presented at academic and clinical conferences
on the subject. Soper’s clinical interests include the provision of therapeutic support for people
affected by suicide and those suffering from alcoholism and other addictions.

Gayle S. Stever is a Professor of Psychology at Empire State College/State University of

New York. She has done research in the areas of parasocial theory, media effects and fan studies
for over 30 years, and has authored numerous peer-reviewed articles on these subjects, in
addition to writing The Psychology of Celebrity (2019). With her degree in lifespan development
psychology, looking at media effects from a lifespan perspective is central to her work.

Benno Torgler is Professor of Economics at the Centre for Behavioural Economics, Society
and Technology (BEST) and School of Economics and Finance, Queensland University
of Technology (QUT), Australia. He was also Adjunct Professor at the EBS Universität für
Wirtschaft und Recht, Germany (2012–2015) and an Australian Research Council (ARC) Future
Fellow (2011–2015). Before QUT he was a Research Affiliate and Lecturer in International
and Comparative Political Economy in the Leitner Program at the MacMillan Center, Yale
University; a Visiting Scholar in the Law and Economics Program at University of California-
Berkeley; and a Visiting Scholar in the Andrew Young School of Policy Studies at Georgia State
xvi THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

University. His primary research interest lies in the area of economics, but he has also published
in journals with a political science, social psychology, medical, sociology and biology focus.

Anthony Walsh is a Professor Emeritus at Boise State University. He received his PhD from
Bowling Green State University in 1983 after working as a marine, police officer and probation/
parole officer. His primary interest is biosocial criminology. He has published 43 books, the
latest being God, Science, and Society: The Origin of the Universe, Intelligent Life, and Free
Societies (Vernon, 2020).

Jessica Wells is an Assistant Professor of Criminal Justice at Boise State University. She received
her PhD in Criminal Justice and Criminology from Sam Houston State University in 2017. Her
research is in the area of psychological and biosocial perspectives on criminal offending. Her
publications focus on the consequences of stress and trauma exposure in samples of community
members, incarcerated individuals and criminal justice professionals.

Stephen Whyte is a Research Fellow in Behavioural Economics at the Centre for Behavioural
Economics, Society and Technology (BEST) and School of Economics and Finance,
Queensland University of Technology (QUT), Australia. His research focus explores large-
scale decision making in mate-choice settings. His work takes a multi-disciplinary approach in
studying key sex differences in human behaviour, with work that bridges the fields of applied
micro-economics, personality and social psychology, and evolutionary biology. His most recent
research has explored such diverse topics as sex differences in nonbinary gender identification,
male and female decision making in assisted reproductive and donor insemination medical
environments, and preferences vs choice in cyber dating markets.

David Wiesenthal received a bachelor’s degree in psychology from the City College of New
York and a doctoral degree in social psychology from the State University of New York at
Buffalo, followed by a postdoctoral fellowship from York University, Canada. He is now a
Professor Emeritus and Senior Scholar at York University, Canada. He has been a Visiting
Professor at the Hebrew University of Jerusalem and lectured as a Scandinavian Fellow at
the Building Research Institute (Gävle, Sweden), Umeå University, Lund University, and the
Helsingborg campus of Lund University, as well as the University of Costa Rica and the College
of Psychologists (Costa Rica). His research interests are social and environmental influence
processes, driver behaviours, scientific racism and research ethics.

Holly Wilson is a second-year PhD student in the Department of Computer Science at the
University of Bath. Her doctoral research involves investigating the neural correlates of both
visual imagery and perception, and developing a brain computer interface which facilitates visual
imagery reconstruction. Holly merged her interests in both artificial and natural intelligence by
obtaining a BSc in Psychology followed by an MSc in Computer Science. In the first year
of her PhD she focused on the ethics and impact of AI on people, on both an individual and
societal level, alongside developing agent-based models of cultural evolution during the Upper
Palaeolithic transition.
 PART 1

Integration within Psychology

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Evolutionary Psychology and
Counseling and Psychotherapy
This chapter covers two important clinical
applications of evolutionary psychopathology:
(1) evolutionary-inspired psychological inter-
ventions, and (2) integration of evolutionary
insights into couples therapy.
APPLICATIONS OF EVOLUTIONARY
PSYCHOLOGY IN THE TREATMENT OF
MENTAL DISORDERS
In recent years, there has been an outpouring
of attempts at evolutionary hypotheses of
mental disorders. Different authors have pro-
posed evolutionary explanations for depres-
sion (Durisko et al., 2015), anxieties (Gilbert,
2001; Nesse, 1998), schizophrenia (Crow,
2000), and personality disorders (Glenn
et  al., 2011; Gutiérrez et  al., 2013; Hertler,
2014; Molina et  al., 2009; O’Reilly et  al.,
2001), to name a few.
Evolutionary explanations of mental dis-
orders typically focus on the role of the
1
Cezar Giosan
symptoms in increasing fitness, seeing them
as evolved strategies serving an organism’s
goal to survive and reproduce, or, conversely,
center on the mismatch between our adap-
tations, evolved during the Environment of
Evolutionary Adaptedness, and the modern
world.
An important practical question that arises
from this substantial body of work revolves
around the clinical implications of such theo-
ries. Although still speculative for the most
part, evolutionary explanations of mental dis-
orders do raise the intriguing possibility that
psychological interventions that target fitness
could have unique clinical benefits, which
can go above and beyond those of current
treatments.
To this end, clinical psychologists, psy-
chotherapists, or clinical researchers have
attempted to bridge the gap between evolu-
tionary theory and clinical practice by devel-
oping protocols that integrate evolutionary
insights into the treatment of mental dis-
orders. Some of these protocols have been
4 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
tested in randomized clinical trials with active
comparators, while others have led to case stud-
ies or small-n pilot studies with no comparators.
Also, some of these insights have been used in a
cross-diagnostic manner – that is, same princi-
ples applied to various conditions – while oth-
ers are specific to certain disorders.
Alfonso Troisi and Michael McGuire have
proposed that an evolutionary therapy’s main
aim is to increase a patient’s short-term bio-
logical goals. In this light, evolutionary psy-
chopathology makes the distinction between
mental disorder and mental suffering, seeing
many mental symptoms as adaptive reac-
tions to situations associated with negative
cost–benefit outcomes, which should not be
treated if they do not cause distress (Troisi
and McGuire, 2014).
More specifically, in the view of Troisi and
McGuire (2014), an evolutionary-driven ther-
apy should (1) address cost–benefit outcomes;
(2) facilitate the development of revised mod-
els of the social environments; and (3) aid the
patient in developing capacities to achieve
biologically relevant goals. This therapy
should attempt to refine traits or to foster the
use of alternative capacities that can help to
achieve high-priority goals. In the more diffi-
cult cases, the authors argue, therapy should
encourage patients with suboptimal functional
capacities to actively search for environments
where they can be successful in reaching high-
priority goals (Troisi and McGuire, 2014).
Evolutionary Psychology and
Case Conceptualization
Case conceptualization – an explanation,
offered by the therapist, of the problems
bothering a patient – addresses the mental
problem and its possible causes, the ethio-
pathogenetic processes presumed to be
involved, and the positive or adverse effects
of the proposed treatment. The efficient con-
ceptualization of a problem can generate
positive expectations about the treatment as
well as a sense of prediction and control in
the patient, which can facilitate recovery
(John and Segal, 2015; Kuyken et al., 2008).
Historically, Sigmund Freud was the first
to introduce case conceptualization as a
key element in psychotherapy, through the
analysis of the latent content of the dreams
and interpretation of the neurotic symptoms
(Freud, 2017). Today, case conceptualiza-
tion is used in many therapeutic approaches.
For instance, Cognitive Behavioral Therapy
(CBT), one of the most widely used interven-
tions for anxiety and depression, typically
includes information about the causal mecha-
nisms of the problem, that is, proximal causes
of psychopathology, thus answering the ‘how’
questions from the ABC1 model (Ellis, 1994;
Ellis et al., 2007).
Case conceptualization in modern psy-
chological interventions, however, typically
includes only information about proximal
causes of the symptoms. For instance, the
ABC model leaves out the ‘how’ questions,
focusing almost exclusively on the immediate
mechanisms, such as dysfunctional thinking
(Lam and Gale, 2000).
Some therapeutic schools do offer distal
explanations of symptoms, but none of them
goes so deep as to bring into the therapeutic
discourse, in a coherent, unified manner, the
factors, forces, and elements that have shaped
the evolution of our species. For example,
psychoanalysis, the first school of thought
that brought into discussion the distal causes
of psychopathology, explains phobias through
repression and displacement. A conflict origi-
nates in childhood and that conflict is either
repressed or displaced onto the feared object.
As an illustrative example, in ‘Notes upon a
case of obsessional neurosis’ Freud attributed
the Rat Man’s fear of relatives dying from
being burrowed through by rats to guilt origi-
nating from a repressed desire he had earlier
to see women that he knew naked (Williams,
2008). A phobia of snakes, from the same
psychoanalytical perspective, was an uncon-
scious fear of something else, which was to
be unraveled in therapy through dream inter-
pretation or analysis of slips of tongue.
 EVOLUTIONARY PSYCHOLOGY AND COUNSELING AND PSYCHOTHERAPY
Evolutionary psychopathology makes one
giant leap further and addresses the distal,
evolutionary causes of mental illness, namely,
the evolutionary factors and forces that might
be at the root of the presenting symptoms. By
addressing the evolutionary causes of behav-
iors, evolutionary psychopathology finds itself
in the privileged – and unique, to some extent –
position to offer explanations of symptoms
that typically make much sense to patients.
By offering evolutionary explanations
of symptoms, evolutionary psychology can
enhance case conceptualizations of various
treatment approaches in meaningful ways,
potentially leading to better therapeutic out-
comes. For instance, incorporating information
about the hypothesized adaptive functions of
the symptoms in the ABC, or the further refined
ABCDE model (Ellis, 1994; Ellis et al., 2007),
can lead to answers to ‘why’ questions, thus
giving the patient a broader and more meaning-
ful understanding of the problems they are con-
fronting, which can lead to better acceptance.
Integration of Evolutionary
Principles in Specific Forms
of Therapy
There have been several attempts to integrate
evolutionary insights into various therapies
in recent years. We begin by briefly describ-
ing the possible evolutionary resuscitation of
Freud’s psychoanalysis and Jung’s analytical
therapy and continue with a presentation of
several evolutionary-driven therapy protocols
that have been tested in randomized clinical
trials. We will end this section with a brief
presentation of the potential applications of
evolutionary conceptualizations to other
types of mental conditions.
Psychoanalysis
Evolutionary psychotherapies place substantial
importance on the therapeutic relationship.
This is not something new. A century ago,
5
Sigmund Freud also made this central in his
psychoanalytical psychotherapy. Indeed, one of
the fundamental tenets of psychoanalytical
psychotherapy is complete disclosure and com-
munication: the patient is required to disclose
anything that comes to their mind, without
censorship. Thus, in Freudian psychoanalysis,
the therapist took a central role in patients’
lives, since he would learn information about
them that no one else was privy to. This unique
and extremely close relationship (sessions were
held several times per week) made Freud real-
ize that it played a major role in the therapeutic
outcomes and subsequently led to the definition
of important constructs such as ‘transference’,
which slowly replaced the initial emphasis on
sexual symbolism with more nuanced under-
standings of the therapeutic alliance.
Some scholars note that many psychothera-
pies – including evolutionary interventions –
do not place sufficient importance on the
relationship between the client and the thera-
pist, or they may use that relationship to manip-
ulate patients in what the therapist believes is in
his own best interests. Kriegman (2000) argues
that evolutionary insights can help reduce
this risk in all forms of therapy, including
psychoanalysis. Since psychoanalysis revolves
around a deep relationship between two unre-
lated individuals and since one evolutionary
principle is that we are hardwired to operate
for our own benefits, it follows that the power
the therapist has over the patient may some-
times be used to further the interests of the
therapist, even if unconsciously (Kriegman,
1998). Becoming more aware of the distal
mechanisms responsible for human behavior
will place a therapist in a better position to
avoid confusion between proximal and distal
causes, ultimately benefitting the patient. For
instance, as Kriegman describes hypotheti-
cally, a woman who dresses provocatively but
is angered when perceived as a sexual object
can be seen by an analyst as having an uncon-
scious wish to be ravished or raped, with
anger being a reaction formation. From an
evolutionary lens, however, this interpreta-
tion might reflect a mix between projections
6 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
of male wishes and confusion of proximal
(dressing sexily) and ultimate causes (wom-
an’s self-interest enhancement through the
stimulation of men). Becoming more aware
of such nuances can help the therapeutic
process and, therefore, evolutionary inter-
pretations can bring value to such clinical
situations.
Jungian Analytical Therapy
While for classical psychoanalytical therapy
the answer to the central question of what is
wrong with the patient is their repressed
memories, which the therapist tries to bring
to the conscious level using strategies such as
interpretation of dreams or slips of the
tongue, in Carl Gustav Jung’s analytical
therapy it is the archetypal intent that needs
to be freed to unleash the patient’s full poten-
tial (Stevens, 2000).
Jung’s theory of archetypes – universal,
innate, archaic patterns and images of evo-
lutionary origins that stem from the collec-
tive unconscious and which are the psychic
counterpart of instinct – closely anticipated
the notions of evolved mechanisms (innate
strategies or algorithms) present in evolu-
tionary theories today. Indeed, Jung rejected
the tabula rasa understanding of human
mind, common to his contemporaries (nota-
bly, John Watson in the United States) and
replaced it with a theory that included the
enormous influence of evolutionary factors
on human behavior. Like evolutionary psy-
chologists today, Jung argued that homeosta-
sis, epigenesis, and adaptation are at the basis
of the human psyche (Stevens, 1982, 1999),
a paradigm that was in stark contrast to the
blank-slate view of the mind from the Standard
Social Sciences Model. Jung also rejected the
sexualized Freudian interpretation of com-
plexes such as Oedipus, anticipating the later
works of John Bowlby, who argued that a child
is attached to his/her mother because she is the
caregiver (Bowlby, 1983, 2005). Also, of note,
in clinical practice, Jung rejected Freud’s cold
objectivity in the therapeutic relationship,
replacing it with something common in evo-
lutionary therapies today, namely, the empha-
sis on a warm, reciprocal alliance.
Not unlike the mismatch hypothesis
(Giphart and van Vugt, 2018), psychopathol-
ogy, in the Jungian paradigm, occurs when
environmental mismatches at critical develop-
mental stages lead to malfunction in ‘arche-
typal’ strategies (Stevens, 2000). Evolutionary
psychology can add to analytical therapy the
critical element of an even broader view of
self than Jung conceived. Armed with mod-
ern knowledge about the functions of psy-
chological mechanisms, therapists nowadays
can bring into the clinical conceptualization a
more expanded conversation about the role of
these mechanisms in mental illness.
Evidence-Based Evolutionary
Interventions
After this brief theoretical presentation of the
ways in which evolutionary insights can aid
Freudian and Jungian therapies, we continue,
in the section that follows, with the presenta-
tion of results from several empirical studies
that have examined the benefits of integrat-
ing evolutionary insights into the treatment
of depression and personality disorders.
Depression
A Rwandan man once described the Rwandan
treatment for depression to the 2001 National
Book Award winner Andrew Solomon like
this:
You know, we had a lot of trouble with Western
mental health workers, especially the ones who
came here right after the genocide. They came and
their practice did not involve being outside in the
sunshine… which is, after all, where you begin to
feel better. There was no drumming or music to
get your blood flowing again – when you’re
depressed and low you need to have your blood
flowing. There was no sense that everyone had
taken the day off so that the entire community
 EVOLUTIONARY PSYCHOLOGY AND COUNSELING AND PSYCHOTHERAPY
could come together to lift you up and bring you
back to joy. There was no acknowledgement of
the depression as something invasive and external
that could actually be cast out of you again.
Instead, they would take people one at a time into
these dingy little rooms and have them sit around
for an hour or so to talk about bad things that had
happened to them. We had to ask them to leave
the country. (Taljaard, n.d.)
While this description of an intervention for
depression is in stark contrast to the standard
one-hour-weekly therapy sessions common
in Western cultures, it would not surprise an
evolutionary therapist. Evolutionary psycho-
pathologists view mild and moderate depres-
sion as functional states, serving adaptive
functions (for a review, see Durisko et  al.,
2015). For instance, as early as the 1990s,
some authors conceptualized depression as a
warning signal that biosocial goals have not
been achieved (Nesse, 1991). The clinical
implication of this line of thought is that
finding solutions to reset the cost–benefit
balance in favor of the patient should make
depressed mood subside.
In one of the earlier attempts at incorpo-
rating evolutionary insights into therapy for
depression, McGuire and Troisi presented a
clinical case of a patient who was depressed
because of her inability to have children (i.e.,
major direct fitness problem). The treatment
focused on addressing the dysregulating
effects of the patient’s inability to reproduce,
and, crucially, also formulated strategies to
help this patient’s fitness through kin invest-
ment (i.e., inclusive fitness) (McGuire and
Troisi, 1998: 270–271).
Treating Depression Downhill
One evolutionary-based intervention proto-
col for depression is Treating Depression
Downhill – TDD (Krupnik, 2014). TDD
relies on an experiential approach and
involves three distinct phases: (1) explora-
tory, in which the patients gain insight into
their experience of defeat; (2) acceptance, in
7
which the patients terminate protest, that is,
accept defeat as an immutable fact of their
lives. This phase, which is the analogue equiv-
alent of exposure therapy in anxiety disorders,
is the centerpiece of TDD, as it facilitates the
transition from protest to acquiescence; and
(3) behavioral activation without the func-
tional analysis component. Throughout TDD
treatment, cognitive reappraisal takes place,
following standard cognitive therapy
approaches (e.g., analysis of distortions).
Evidence in Favor of TDD
A preliminary study testing the efficacy of
TDD was conducted in the form of a pilot
observation on a sample of 12 participants,
who met for 24 biweekly, 90-minute-long
sessions (Krupnik, 2014). The protocol dem-
onstrated effectiveness and specificity for
depression, differentiating it from anxiety
and personality disorders. The results showed
marked decline in depressive symptomatol-
ogy; however, the study was underpowered
and the tentative trends in the dynamics of
the participants’ scores did not reach statisti-
cal significance. The TDD protocol needs
further testing in randomized controlled
studies in comparison with established proto-
cols for depression to better establish its
efficacy.
Since not all clients can engage in mind-
fulness, which is a key element in the accept-
ance phase in TDD, the same author replaced,
in a further case study on a medicated patient,
the acceptance phase from TDD with eye
movement desensitization and ­reprocessing –
EMDR (Shapiro, 2017). In this study, EMDR
was used in a truncated form, and the nature
of the targets was the subjective percep-
tion of loss, rather than actual events, while
reappraisal took place along the protest–­
acceptance axis. The results showed that,
at the end of the treatment and at follow-
up assessment, the patient reported a more
accepting disposition and decreased depres-
sive symptoms (Krupnik, 2015).
8 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Following the same line of investigation,
another study conducted by the same author
reported a case series of 21 military person-
nel diagnosed with depressive disorders, who
received a course of TDD-EMDR (Krupnik,
2018). By the end of treatment (12 sessions),
80% of completers (n = 15) did not meet
the criteria for depressive disorder and they
showed a significant reduction in scores on
the Beck Depression Inventory-II – BDI-II
(Beck et  al., 1996) with a large effect size
(d = 2.8) and an increase in accepting dis-
position (d = 1.8) on the Acceptance and
Action Questionnaire (Bond et  al., 2011).
Non-completers showed a similar decrease
in the BDI-II scores at mid-treatment. The
author observed no statistically significant
decrease in anxiety symptoms on the BDI-II.
These results suggest that TDD-EMDR may
be an effective treatment for depressive disor-
ders (Krupnik, 2018). They also indicate that
this type of intervention may target depres-
sive over anxiety symptoms (Krupnik, 2018),
as was previously observed for the original
TDD pilot study (Krupnik, 2014).
Therapeutic Lifestyle Change
for Depression (TLC-D)
Another attempt to incorporate evolutionary-
inspired interventions in therapy is the
Therapeutic Lifestyle Change for Depression
(TLC-D) protocol (Karwoski et  al., 2005),
which includes several evolutionary elements
thought to have positive effects on mood.
TLC-D combines several relevant factors,
some of which are evolutionary-relevant, that
are shown to be effective in the treatment of
depression. These factors include:
1 Omega-3 fatty acid consumption (Peet and
Horrobin, 2002).
2 Bright light exposure (Martiny et al., 2005).
3 Sleep hygiene (Mayers and Baldwin, 2006).
4 Aerobic exercise (Blumenthal et al., 2007).
5 Anti-rumination exercises (Fennell and Teasdale,
1987).
6 Social support (George, 1989).
Evidence in Favor of TLC-D
Karwoski et al.’s (2005) protocol was retested
with additional data and gender comparison
by Jacobson et al. (2007). The authors exam-
ined TLC-D on 81 patients who underwent
12 sessions of TLC-D therapy, with follow-
up evaluations at three and six months. The
experimental group was compared to a
Treatment as Usual (TAU) group, represent-
ing one-third of the sample. The results
showed that the TLC-D group outperformed
the control group. The results also showed
that, at the end of the therapy, participants
averaged a 17.8% decrease in BDI-II (Beck
et al., 1996) scores, which represented a sta-
tistically significant 60.6% reduction from
baseline. These improvements were stable,
showing a 67.7% reduction at three-month
follow-up, and 64.0% reduction at six-month
follow-up.
Further research on TLC-D continued to
show promising results. In a study conducted
by Botanov et al., (2012), 29 patients were
recruited into a TLC-D protocol, in a two-to-
one random assignment (22 in TLC-D and 7
in TAU). The participants underwent 12 ses-
sions of group therapy over 14 weeks and
were assessed weekly with the BDI-II (Beck
et al., 1996). The results showed a clinically
significant response (>= 50% reduction in
BDI-II scores) in 77.3% of the participants
in the TLC-D condition versus 28.6% in the
TAU condition, and, notably, no significant
change in BDI-II scores was observed from
treatment end to six-month follow-up, sug-
gesting low relapse rates post-treatment.
Cognitive Evolutionary Therapy
for Depression (CETD)
Another clinically tested evolutionary-driven
intervention protocol for depression is
Cognitive Evolutionary Therapy for Depression
(CETD) (Giosan, 2020; Giosan, Cobeanu,
Wyka, et  al., 2020; Giosan, Cobeanu et  al.,
2014). As conceptualized by the authors,
 EVOLUTIONARY PSYCHOLOGY AND COUNSELING AND PSYCHOTHERAPY
besides targeting the proximal causes of
depression as is standard in CBT, CETD
focuses on distal causes as well, such as inclu-
sive fitness or reproductive success. While
sharing common underpinnings with CBT,
CETD adds the inclusion of evolutionary con-
ceptualizations of the patient’s symptoms and
the targeting of fitness-related problems. Very
much unlike classical Cognitive Therapy for
Depression, in which the problems that preoc-
cupy the patient are identified by the patients
during the therapy sessions, CETD starts from
the premise that depressive symptoms reflect
fitness difficulties, some of which are
unknown to the patients, and which can be
identified via an evaluation of the patient’s
fitness prior to the first session. By identify-
ing a patient’s fitness problems at intake, the
CETD therapist thus is pre-equipped with this
knowledge at the first session and can start
working with the patient on problematic areas
right away.
Along with these evolutionary-driven behav-
ioral activations, discussions about human
nature from evolutionary perspectives are also
taking place during CETD, such as modularity
(Cosmides and Tooby, 1994), parental invest-
ment theory (Buss et  al., 1990), conspicuous
consumption (Sundie et  al., 2011), or costly
signaling theory (Fraser, 2012), all of which
can facilitate acceptance, a key CBT ingredi-
ent (Chamberlain and Haaga, 2001).
The instrument that CETD therapists
use to identify a patient’s fitness difficulties
is the Evolutionary Fitness Scale – EFS
(Giosan et  al., 2018). The EFS is a 58-item
scale assessing mismatches between the
Environment of Evolutionary Adaptedness
and the modern world, such as in physical
activity or nutrition, environmental misfits,
or fitness-related factors such as health of
the actor, his/her partner and their extended
families, attractiveness (both of the actor and
partner), status, resource control, extended
family, social capital, and mate value. Some
examples of items are: ‘I visit my relatives fre-
quently’, or ‘I am an active outdoors person’,
which are actionable in therapy.
9
The CETD manual (Giosan, 2020) provides
the evolutionary therapist with concrete exam-
ples of therapeutic interventions on each of the
EFS items. For instance, a negative endorse-
ment of the EFS item ‘I have at least one best
friend’ should be dealt with by exploring the
reasons and refuting dysfunctional thinking,
as well as exploring modalities to increase
connectedness with at least one non-relative.
Likewise, a negative endorsement of the EFS
item ‘My family brag about me’ should be
dealt with by discussing solutions to increase
status and dominance (e.g., more education if
appropriate, job change, community involve-
ment, etc.) (‘Darwinian Psychotherapy’,
2019; Giosan, 2020).
As far as the therapeutic alliance is con-
cerned, the CETD protocol advocates that
the therapist go beyond the recommen-
dations of interventions such as Rational
Emotive Behavior Therapy – REBT (where
the alliance is centered on unconditional
acceptance, empathy, humor, and genuine-
ness) or psychoanalysis (friendly neutrality)
and try to become a patient’s psychological
kin, while maintaining a safe set of bounda-
ries (“Darwinian Psychotherapy,” 2019;
Giosan, 2020). This approach is in line with
the suggestions of other evolutionary psy-
chopathologists, who emphasize rapport
between the therapist and patient (Troisi
and McGuire, 2014: 34), question the effi-
cacy of one-hour-per-week therapy sessions
(Gilbert et  al., 2014: 19), or propose that
depressed patients may even need ‘thera-
peutic cheerleading’ (Markowitz, 1994;
Michels, 1997).
While many evolutionary therapists argue
for a stronger connection between the thera-
pist and the depressed patient than the one
advocated by other therapeutic paradigms,
support for such an idea predates these recent
developments in evolutionary psychotherapy.
The early and fascinating work of Jerome
Motto, who found that simply mailing per-
sonally signed ‘Caring Letters’ to people who
had attempted suicide drastically reduced
future suicide attempts, as people felt more
10 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
connected to the therapist communicating with
them, illustrated the importance of therapeutic
rapport (Motto, 1976). The ‘Caring Letters’
approach has been revised to include a form of
intervention that essentially makes the thera-
pist available almost continuously, and the
results of this kin-like therapeutic relationship
are promising. (For a detailed account of this
project, including historical aspects, see James
Cherkis’ (n.d.) excellent article in Huffington
Post, https://bit.ly/2NqSWeC).
Evidence in Favor of CETD
In a case study examining the potential bene-
fits of CETD, Giosan, Muresan et  al. (2014)
used this protocol on a patient with an intake
score of 22 on the BDI-II (Beck et al., 1996)
and a diagnosis with depression made with the
Structured Clinical Interview for the DSM
(SCID) (First et  al., 1997), who presented
deteriorating functioning (school perfor-
mance) and quality of life following a recent
break-up. An assessment of her perceived fit-
ness with the EFS (Giosan et  al., 2018)
revealed deficiencies in self-image, healthy
eating habits, and physical activity. The patient
was offered a cognitive-evolutionary conceptu-
alization of her symptoms that centered on the
distal causes of depression as well as on the
dysfunctional cognitions that led to symptoms
(Giosan, Muresan et al., 2014). The treatment
focus was to engage the patient in the EFS-
suggested fitness-increasing activities, while
simultaneously challenging dysfunctional
thinking. The treatment was successful, the
patient achieving a ~68% reduction in the
BDI-II (Beck et al., 1996) scores by session 8
(BDI-II = 7), therapeutic gains maintained at
post-evaluation (BDI-II = 7) and follow-up
(BDI-II = 13).
A randomized, single-blinded active-­
controlled design (Giosan, Cobeanu, Wyka,
et  al., 2020; Giosan, Cobeanu et  al., 2014)
expanded on this preliminary case study and
contrasted the efficacy of CET for depression
with one of the best validated interventions
for depression: Cognitive Therapy (CT; Beck
et  al., 1979). A total of 97 depressed patients
received 12 sessions of either (1) CETD or
(2) CT. Baseline, mid-treatment, post-treatment,
and three-month follow-up assessments were
conducted. The CT group underwent classical
cognitive interventions aimed at the correc-
tion of dysfunctional, automatic thoughts and
beliefs hypothesized to be implicated in depres-
sive symptoms. These interventions were paired
with behavioral activation and positive rein-
forcements. The CETD group added s­pecific
goals targeted at increasing fitness (see full pro-
tocol at Giosan, Cobeanu et al., 2014).
Both interventions led to similar reduc-
tions in depressive symptomatology, as meas-
ured by the BDI-II (Beck et al., 1996), which
were maintained at three-month follow-up.
Although non-significant, the CETD group
showed a consistent pattern of larger gains
(greater decreases in BDI-II scores) dur-
ing the treatment as well as post-treatment.
Fewer CETD participants were classified as
having moderate or severe depression over
time, with between-group analyses showing
trend differences at post-treatment.
The results also showed that the CETD
group experienced significantly greater reduc-
tions in behavioral inhibition/avoidance at
both post-treatment and follow-up, compared
with the CT group. Notably, CETD was also
significantly superior to CT in increasing
engagement in social and enjoyable activities
at post-treatment. The study showed that in the
participants receiving CETD, but not in those
receiving CT, engagement in these activities
was directly related to decreased symptoms
of depression, suggesting that CETD leads to
greater social reach, which, in turn, might
translate into better therapeutic outcomes
(Giosan, 2020; Giosan et  al., 2019; Giosan,
Cobeanu, Wyka, et al., 2020).
Personality Disorders
Personality disorders are typically perceived
as difficult to address in therapy, with some
 EVOLUTIONARY PSYCHOLOGY AND COUNSELING AND PSYCHOTHERAPY
of them, such as borderline personality disor-
der, being especially prone to de facto demedi-
calization (Sulzer, 2015). The evolutionary
scholars Prunetti et  al. (2013) developed a
protocol for Cognitive Evolutionary Therapy
specifically aimed at personality disorders
(CET-PD). CET-PD is based on the Darwinian
view that humans are driven by evolutionary-
selected motivations and develop psycho-
pathologies when their biologically relevant
goals are not met. Thus, failures in patients
with personality disorder are explained by the
authors as resulting from disordered function-
ing of evolutionary-shaped social motives
(Prunetti et al., 2013). The authors differentiate
CET-PD from other treatments from which it
borrows, such as Cognitive Therapy (Beck,
1976), Rational Emotive Therapy (Ellis and
Dryden, 2007), and Dialectical Behavioral
Therapy (Chapman, 2006).
Key Elements in CET-PD
The key elements of CET-PD include:
1 Focus on restructuring schemas of self-with-others
around biologically relevant needs (attachment,
caregiving, social ranking, mating, cooperation).
2 Special focus on the therapeutic relationship.
CET-PD places importance on the rapport
between the therapist and patient, with special
attention on discovering the specific motive that
is active during the flow of therapy conversation.
3 Assessing interpersonal motivations during the
therapeutic relationship (e.g., attachment or
social rank).
4 Managing the therapeutic relationship to pre-
vent/repair ruptures.
5 Making people aware of how dysfunctional
schemas guide behaviors.
Evidence in Favor of CET-PD
The authors examined the benefits of CET-PD
in an intensive 20-hour weekly three-week
residential treatment (both individual and
group) of a wide range of severe personality
disorders. Fifty-one patients with various
11
personality disorders were assessed at admis-
sion, discharge, and three-month follow-up
and the outcome measures consisted of self-
reported depression, anxiety, general symp-
toms, duration of inpatient admissions after
the program was over, and continuation in an
outpatient program. The results suggested
that CET-PD was effective in reducing the
level of depression and anxiety, with a change
that was stable for trait anxiety. Obsessive
symptoms, paranoid ideation, psychoses, and
feelings of self-inadequacy and inferiority
diminished. Overall, the results showed an
improvement in psychopathology after
release and in follow-up sessions, a decrease
in the number of further hospital admissions,
and an increased level of outpatient therapy
attendance (Prunetti et al., 2013).
Potential Applications of
Evolutionary Conceptualizations
to Other Mental Conditions
In the previous section, we reviewed the pos-
sible integration of evolutionary insights in
psychoanalysis and analytical psychother-
apy, and we summarized the results from
controlled studies that examined the efficacy
of evolutionary interventions for depression
and personality disorders. In the next section,
we briefly present, in a speculative manner
that needs further testing in controlled trials,
some potential applications of evolutionary
conceptualizations to the treatment of other
mental conditions.
Postpartum Depression
Conceptualizing postpartum depression as an
adapted response to unfavorable circumstances
(e.g., child sickness, lack of resources or sup-
port) (Hagen, 1999), evolutionary mismatch
(Crouch, 1999), or age (Bottino et  al., 2012)
may make a patient suffering from it more
likely to recover. A treatment aimed at increas-
ing fitness (e.g., by focusing on resource
12 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
acquisition) may be better than correcting
dysfunctional beliefs (‘I am a bad mother’).
From an evolutionary perspective, cognitive
techniques could be tried to increase the per-
ceived benefits of having a child and reduce
the perceived costs. Such a strategy might
lead to a decrease in the severity of postpar-
tum depression precisely because it addresses
evolutionary causes. For instance, a young
mother could understand that her symptoms
do not reflect her incapacity as a mother, but,
rather, a mechanism by which she is asking
for help. Thus, the intervention could focus
on coping mechanisms and problem-solving
targeting the fundamental causes of the
symptoms, addressing not only the depres-
sive symptoms per se, but also the situation
that led to them (decreased fitness).
Anxiety Disorders
By distinguishing between situations in
which anxiety is disabling (when medication
can be useful) and those where anxiety may
be adaptive, evolutionary theories can offer
meaningful case conceptualizations that can
help patients to accept these symptoms and
possibly reduce impairment. For instance, a
debilitating phobia of snakes might be
accepted and dealt with better by a patient if
it is explained to her that fear of snakes is an
evolved fear which increased the likelihood
of survival in ancestral times (Marks and
Nesse, 1994) and that her extreme anxiety
around such stimuli is not a brain disorder,
but an evolved, normal mechanism that may
be functioning in overdrive. Similarly, in
Obsessive Compulsive Disorder, conceptual-
izing the symptoms as exacerbated mecha-
nisms to facilitate reproduction and protect
offspring (Feygin et  al., 2006) can lead to
better acceptance of the symptoms, a key ele-
ment in the recovery process (Chamberlain
and Haaga, 2001).
One of the most common forms of anxiety,
social anxiety (SA), is particularly resist-
ant to treatment, with only about half of
patients showing improvement, even when
gold-standard treatments, such as CBT, are
used (Loerinc et al., 2015). Conceptualizing
SA as one of the poles (besides social domi-
nance) necessary to maintain social order
(Öhman, 1986), or as a vestigial response to
social threat (Trower and Gilbert, 1989), may
increase a patient’s acceptance of the symp-
toms. Moreover, evolutionary understandings
of SA may serve as a guide in therapeutic
decisions. For instance, in some cases, just
treating symptoms (through gradual expo-
sure, for instance) may not be enough, and
a discussion about eliminating or modifying
the circumstances that elicit symptoms may
be in order (Brosnan et al., 2017).
Providing patients with evolutionary
explanations of phobic symptoms is not pos-
sible in all the cases, so only some patients
will benefit from this kind of evolutionary-
aided case conceptualization. This approach
is suitable in the case of patients with fears
of biologically relevant stimuli, such as
heights, public speaking, dark, blood, or
certain types of animals, who could ben-
efit from logical distal explanations of the
symptoms, and less so, if at all, in the case
of patients presenting fears of evolutionary-
irrelevant objects, such as a fear of cotton
balls or certain colors.
In other words, while evolutionary expla-
nations of anxieties can be helpful in treat-
ment, this does not mean a replacement of
current explanations, which typically rely
on either proximal causes or distant, but not
evolutionary ones (e.g., childhood traumas).
On the contrary, multi-layered explanations
(evolutionary, developmental, proximal
mechanisms) should be used, with evolution-
ary insights helping in the creation of a more
comprehensive causal picture of the problems
bothering a patient.
Dysmorphic Disorder
Dysmorphic disorder is explained evolution-
arily through one’s attempt to compare with
 EVOLUTIONARY PSYCHOLOGY AND COUNSELING AND PSYCHOTHERAPY
others and the avoidance of rejection or ridi-
cule, which are linked to lower status and
lower mate value (Veale and Gilbert, 2014).
Understanding the context and functions of
the behaviors associated with this condition
can be critical for the success of an interven-
tion, especially when the patient has aversive
emotions, such as shame or rejection, which
have not been properly processed (Veale and
Gilbert, 2014). Cognitive behavioral tech-
niques for treating this condition could be
improved through the analysis of the functions
and contexts in which the behaviors appear.
This can be realized via multiple routes, such
as (1) linking the body-related fears to fears of
rejection or to emotionally charged memories;
(2) rewriting of the narrative; (3) providing an
evolutionary context that separates the symp-
toms from the feelings of shame and the
affected person; or (4) the direct targeting of
the feelings of shame and self-criticism and
the development of social skills through com-
passion (Veale and Gilbert, 2014).
Post-Traumatic Stress Disorder
(PTSD)
Evolutionary hypotheses of PTSD center on
evolved mechanisms of avoiding dangers
(Silove, 1998; Wiedenmayer, 2004). Once a
person has been exposed to a traumatic event,
they will automatically learn to avoid that
type of situation, thus increasing their sur-
vival chances. In some vulnerable individu-
als, this learning can be excessive or hard to
stop. While validated evolutionary interven-
tions for PTSD have yet to be reported, the
reinterpretation of PTSD symptoms as pro-
duced by adaptations to protect an individual
from future harm, mechanisms that are found
in other species as well (Zanette et al., 2019),
can help patients to better understand and
accept the condition and may improve the
therapeutic benefits offered by validated
interventions for PTSD, such as Gradual
Exposure Therapy. Furthermore, some
authors have found a link between life history
13
and PTSD (Giosan and Wyka, 2009), which
might lead to novel, reproductive strategies-
based intervention protocols in the future.
Eating Disorders
Evolutionary explanations of eating disorders
center on intrasexual competition (Li et  al.,
2010) or on life-history strategy (Mehta et al.,
2011). Such hypotheses can have clinical
implications. As in the examples above, an
understanding of the mechanisms that activate
when we eat certain foods can be therapeuti-
cally helpful when the patient’s cognitions are
addressed. In cognitive behavioral interven-
tions, for instance, such explanations could
facilitate the psychoeducational aspect of
therapy and can also aid in the generation of
alternative thoughts that are to replace the
automatic, dysfunctional ones. Furthermore,
the integration of evolutionary explanations of
eating disorders in school curricula may put
young people in a better position to under-
stand human tendencies, which can then act as
an important protective factor.
Substance Abuse
Evolutionary explanations of substance
dependence or abuse revolve around the fact
that people have consumed psychoactive sub-
stances over our recent and ancestral history
(Dudley, 2004; Sullivan and Hagen, 2002),
with some authors arguing that drug
consumption can be associated with fitness
benefits (Kirillova et al., 2008). The mismatch
between the past benefits associated with
such behaviors and the easy access to such
substances in our modern world can make
some predisposed individuals consume them
more, slowly driving them into addiction.
Understanding the distal explanations
of substance consumption might be useful
in therapy, especially in the conceptualiza-
tion phase of an intervention. In substance
abuse, patients typically feel guilt and shame
14 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
(McGaffin et al., 2013). Evolutionary insights
integrated into therapy could potentially
reduce such reactions, deepening positive
therapeutic outcomes.
APPLICATIONS OF EVOLUTIONARY
PSYCHOLOGY IN COUPLES THERAPY
No section on the applications of evolution-
ary psychology to counseling and psycho-
therapy would be complete without a
discussion about the many helpful elements
that evolutionary psychology can bring to
couples therapy. Since evolutionary psychol-
ogy examines the processes that have helped
our ancestors to survive and reproduce, it is
evident that it can bring insights into prob-
lems typically encountered in couples, such
as sexual incompatibilities, emotional and/or
sexual infidelity, trust, gender stereotyping,
or control.
An important class of results generated
by evolutionary psychology is that, when
it comes to mating, men and women are
hardwired somewhat differently and their
strategies to reach a common biological goal –
­reproduction – can, at times, be quite different,
which can be a source of conflict, potentially
leading to the dissolution of the couple.
Studies on heterosexual mating prefer-
ences have documented gender commonali-
ties, such as dependability, faithfulness, and
kindness (Barber, 1995; Buss, 1989; Buss
et  al., 1990) but also differences, in that
women are more interested in earning capac-
ity, while men are more interested in physical
beauty and health cues (e.g., skin smooth-
ness, waist-to-hip ratio) of their partners,
with overlapping bell curves in such tenden-
cies (Buss et al., 1990; Zhang et al., 2018).
Because women require a minimum of
nine months investment (pregnancy) in order
to be successful at reproduction, and because
they cannot have nearly as many children
as a man can theoretically have, they have
evolved to be the choosier sex (Hatfield
and Sprecher, 2016). In contrast, since men
can impregnate a large number of women
in a short period of time, they have evolved
stronger preferences for pursuing short-term
mating opportunities (Schmitt et  al., 2003).
Studies show a gender difference favoring
men in the number of sexual partners (Todd
et  al., 2009) and other research has shown
differences in sexual fantasies, with men
being more likely to fantasize about sexual
variety (Ellis and Symons, 1990). Other
studies show that men are more permissive
about casual sex and have a higher incidence
of masturbation (Oliver and Hyde, 1993) and
are more likely to be consumers of pornog-
raphy (Hald, 2006), an element that has been
linked to couple dissatisfaction (Stewart and
Szymanski, 2012).
Males’ stronger desire for multiple sexual
partners comes with a substantial threat to
marriage, especially since men are some-
times willing to leave their children behind
for the pursuit of new relationships. Indeed,
some authors have argued that men live in
a state of ‘mild torment’ that stems from
their propensity for sexual variety (Singer,
1985a, 1985b). It is evident that such deeply
engrained feelings can have catastrophic con-
sequences on a marriage or long-term rela-
tionship. Therapists must be aware of such
mechanisms and address them in therapy in
a non-judgmental manner, as treating these
tendencies as a ‘disease’ or lack of character
can destroy the therapeutic relationship. In
addressing such issues, therapists must also
be careful about balancing male needs and
female needs. For instance, some authors have
stated that promoting commitment in therapy
may mean, in fact, promoting female repro-
ductive interests at the expense of the male
reproductive interests (Glantz and Moehl,
2000). Such realities can make men feel they
are not understood, which can alter the thera-
peutic relationship. Offering explanations of
the distal causes of the gender differences in
sexual preferences is usually a good strategy
to navigate through these issues in therapy
and sometimes helping a man deal with his
 EVOLUTIONARY PSYCHOLOGY AND COUNSELING AND PSYCHOTHERAPY
conflicts about commitment is better done in
one-person therapy (Gilbert et al., 2014).
Men’s stronger preferences for sexual vari-
ety are also linked to the so-called Coolidge
effect, which is the sexual interest in a new
female, even when the male has reached sex-
ual satiation with his existing partner (Buss,
1994; Dewsbury, 1981; Glantz and Pearce,
1989). In humans, this phenomenon translates
into greater interest for sex outside the pair
and reduced interest for sex within the pair.
This sexual boredom, affecting men and, also,
women, but for different reasons, can under-
mine a relationship. Therapists who understand
that sexual boredom is not reversible and that
the passion of youth cannot be restored are in a
much better position to help a couple in need of
counseling (Glantz and Moehl, 2000).
Moreover, since women, but not men, are
always certain that their babies are theirs,
men are faced with the uncertainty of pater-
nity, which has led to gender differences in
the experience of feelings of jealousy. Thus,
women appear to be more affected by their
partners’ emotional infidelity, whereas men
are more affected by their partners’ sexual
infidelity (Buss et al., 1992; Daly et al., 1982).
This, in turn, makes women less likely to for-
give emotional infidelity, and men less likely
to forgive sexual infidelity (Shackelford
et  al., 2002). The issue of jealousy appears
often in couples therapy and in many a case
one of the partners adamantly accuses the
other of ‘destroying the relationship’ by
being too jealous. Indeed, strong feelings of
jealousy can lead to controlling behaviors
(e.g., controlling the partner’s social media
accounts), verbal or physical violence, sus-
piciousness, isolation of the partner from
family and friends, and lack of trust, which
can undermine a relationship until its com-
plete dissolution. Clinicians would be well-
advised to use evolutionary insights in such
situations and explain to their clients that
jealousy is, at its most fundamental level,
a universal mate-guarding strategy (Buss,
2000), which has helped us pass on our genes
to the next generations, and that, barring
15
extreme manifestations, such as delusions,
it is a ­normal evolved mechanism that we
should not be ashamed of. Reinterpretation of
jealousy as an adaptation that facilitates mate
retention may aid in the therapeutic process.
Understanding the important differences
in sexual preferences and tendencies between
men and women can help a couples thera-
pist’s attempts to heal a fractured relation-
ship. Some authors have argued that some of
the fundamental principles of therapy, such
as communication and sharing feelings, fail
to take into account core male needs (Glantz
and Moehl, 2000), potentially leading to the
inefficiency of the interventions. Indeed,
men’s and women’s relating styles are differ-
ent (Winstead et al., 1997), which may make
the former harder to engage in psychotherapy.
Furthermore, the tabula rasa paradigm advo-
cated by the Social Science Standard Model
assumes no innate gender differences, which
may lead to unreasonable therapeutic requests
of males to reveal inner emotions and insecu-
rities (Shem and Surrey, 1998), further dam-
aging a potentially already fragile therapeutic
relationship. Let’s not forget that studies have
shown that women prefer confident men, who
are able to protect their partners from other
men (Buss, 1989). This can and will make a
man reluctant to display signals of weakness
and subordination both in front of his partner
and in front of the therapist. Clinicians who
understand these nuances well are in a bet-
ter position to establish rapport – critical for
good outcomes – with a male patient in cou-
ples therapy. For instance, since status is often
a crucial factor for men, acknowledging it or
working to increase it can be an effective ther-
apeutic strategy (Glantz and Moehl, 2000).
Similarly, framing interventions in concrete
economic terms (costs/benefits/advantages),
as opposed to the more vague ‘better’, can
lead to positive therapeutic outcomes (Glantz
and Moehl, 2000). Furthermore, given the fact
that men are less likely to disclose emotions
and feelings, encouraging communication and
deep disclosure, especially about weaknesses,
may be counter-productive in some cases and
16 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
outright destructive when the disclosure might
reveal profound couple incompatibilities, such
as sexual (Glantz and Moehl, 2000).
When the issue of misuse of power, such
as anger directed toward family members,
comes up in therapy, some clinicians have
argued that this can lead to shaming and
the activation of self-defense mechanisms
in male patients, and that reframing, in the
sense of explaining what the function of
competition among males is, may be a bet-
ter therapeutic strategy, with the important
observation that the therapist should not rec-
ommend that a man simply give in to his part-
ner (Glantz and Moehl, 2000).
CRITICISM OF EVOLUTIONARY
INTERVENTIONS
In the previous sections, we succinctly pre-
sented some of the recent progress in the field
of evolutionary interventions for certain mental
disorders as well as possible applications of
evolutionary insights in couples therapy.
Despite these promising developments,
we must note the fact that the evolution-
ary hypotheses of mental disorders are, for
the most part, speculative and do not have
strong empirical support yet. Generally,
there is rivalry between hypotheses, with little
movement toward consensus, as well as slow
adoption by practitioners. In addition, while
some of the progress made in evolutionary
randomized clinical trials is noteworthy, it is
very difficult to draw incontrovertible conclu-
sions from medical-style randomized clinical
trials in this field, except perhaps when they
can be pooled in bulk as meta-analyses. Even
then, it is hard to adjust for publication bias,
unaccounted placebo effects, statistical phe-
nomena, and other confounds (Westen et al.,
2004).
Another point of caution in evaluating the
merits of evolutionary interventions is the
fact that meta-analyses generally show that
no particular theoretical approach performs
markedly better than the rest (Cuijpers et al.,
2008; Miller et  al., 2008; Smith and Glass,
1977), and there is no reason to believe that
evolutionary therapies are any different.
Indeed, the most influential factors appar-
ently common to virtually all schools are the
therapist’s technique and the rapport between
client and therapist (Budge and Wampold,
2015). Since some evolutionary therapies
(e.g., CETD, described earlier in this chap-
ter), place, among others, a premium on the
client/therapist relationship, further research
should examine whether this emphasis might
be differentially associated with therapeutic
success.
Last, but not least, the evolutionary inter-
ventions presented in this section have gen-
erally addressed specific mental disorders,
but there is debate in the field whether they
exist as ‘real’ natural conditions to begin
with (First and Pincus, 2009). As such, more
cross-diagnostic evolutionary interventions
should be attempted and tested, since a ther-
apy developed for a certain condition (e.g.,
depression) may well be efficient for a differ-
ent one (e.g., anxiety or self-harm) (Wampold
and Imel, 2015).
SUMMARY
This chapter briefly presented some of the
recent advances in clinical applications of evo-
lutionary psychology. Progress has recently
been made in incorporating evolutionary
insights into psychological interventions for
depression and personality disorders, with sev-
eral randomized clinical trials supporting such
approaches already completed. Treatments of
other psychological problems, such as anxiety,
substance abuse, and eating disorders, might
also benefit from the inclusion of evolutionary
understandings of symptoms, although such
assumptions need to be tested in future con-
trolled clinical studies.
By offering distal explanations of sexual
preferences, evolutionary psychology may
 EVOLUTIONARY PSYCHOLOGY AND COUNSELING AND PSYCHOTHERAPY
also aid substantially in couples therapy.
Issues like jealousy or infidelity can be bet-
ter dealt with in couples therapy when they
are interpreted through evolutionary lenses,
potentially leading to better therapeutic alli-
ance and outcomes.
Despite these recent developments,
much more research on the merits of such
approaches should be conducted, as the
unclear role of common factors in evolution-
ary therapies, the speculative nature of many
evolutionary hypotheses of mental disorders,
and the lack of controlled evolutionary trials
on cross-diagnostic symptoms make it hard
to draw definitive conclusions about the effi-
cacy of such efforts.
Note
1  The ABC model proposes that emotions (C) are
not caused by external events (A), but by beliefs
(B) and, in particular, irrational beliefs (IB) (Sar-
racino et  al., 2017). The ABC model can also
be referred to as the ‘ABCDE’ model, where D
stands for the disputation of beliefs and E stands
for new effect, the result of holding healthier
beliefs (Jorn, 2016).
REFERENCES
Barber, N. (1995). The evolutionary psychology
of physical attractiveness: Sexual selection
and human morphology. Ethology and Socio-
biology, 16(5), 395–424. https://doi.org/
10.1016/0162-3095(95)00068-2
Beck, A. T. (1976). Cognitive therapy and the
emotional disorders. New York, NY: Interna-
tional Universities Press.
Beck, A. T., Rush, A. J., Shaw, B. F., & Emery, G.
(1979). Cognitive therapy of depression.
New York, NY: Guilford Press.
Beck, A. T., Steer, A. R., & Brown, G. K.
(1996). Manual for the Beck Depression
Inventory—II. San Antonio, TX: The Psycho-
logical Corporation.
Blumenthal, J. A., Babyak, M. A., Doraiswamy,
P. M., Watkins, L., Hoffman, B. M., Barbour,
K. A., & Sherwood, A. (2007). Exercise and
17
pharmacotherapy in the treatment of major
depressive disorder. Psychosomatic Medicine,
69(7), 587–596. https://doi.org/10.1097/
PSY.0b013e318148c19a
Bond, F. W., Hayes, S. C., Baer, R. A., Carpenter,
K. M., Guenole, N., Orcutt, H. K., & Zettle,
R. D. (2011). Preliminary psychometric
properties of the Acceptance and Action
Questionnaire–II: A revised measure of psy-
chological inflexibility and experiential avoid-
ance. Behavior Therapy, 42(4), 676–688.
https://doi.org/10.1016/j.beth.2011.03.007
Botanov, Y., Keil, K., Ilardi, S. S., Scheller, V.,
Sharp, K. L., & Williams, C. L. (2012). Success-
ful treatment of depression via therapeutic
lifestyle change: Preliminary controlled-trial
results. Poster presented at the Annual confer-
ence of the Association for Psychological
Science. Retrieved from http://tlc.ku.edu/sites/
tlc.drupal.ku.edu/files/files/PosterAPS2012.
pdf (Accessed 30 November 2018).
Bottino, M. N., Nadanovsky, P., Moraes, C. L.,
Reichenheim, M. E., & Lobato, G. (2012).
Reappraising the relationship between mater-
nal age and postpartum depression according
to the evolutionary theory: Empirical evidence
from a survey in primary health services. Journal
of Affective Disorders, 142(1), 219–224.
https://doi.org/10.1016/j.jad.2012.04.030
Bowlby, J. (1983). Attachment: Attachment
and loss volume one (2nd edition). New
York, NY: Basic Books.
Bowlby, J. (2005). A secure base; Clinical appli-
cations of attachment theory (1st edition).
London: Routledge.
Brosnan, S. F., Tone, E. B., & Williams, L. (2017).
The evolution of social anxiety. In T. K. Shack-
elford & V. Zeigler-Hill (Eds.), The Evolution
of psychopathology (pp. 93–116). Springer:
Switzerland. https://doi.org/10.1007/
978-3-319-60576-0_4
Budge, S. L., & Wampold, B. E. (2015). The rela-
tionship: How it works. In O. C. G. Gelo,
A. Pritz, & B. Rieken (Eds.), Psychotherapy
research: Foundations, process, and outcome
(pp. 213–228). Springer: Switzerland. https://
doi.org/10.1007/978-3-7091-1382-0_11
Buss, D. M. (1989). Sex differences in human
mate preferences: Evolutionary hypotheses
tested in 37 cultures. Behavioral and Brain
Sciences, 12(1), 1–14. https://doi.org/10.1017/
S0140525X00023992
18 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Buss, D. M. (1994). The evolution of desire:
Strategies of human mating. New York, NY:
Basic Books.
Buss, D. M. (2000). The dangerous passion:
Why jealousy is as necessary as love and sex.
New York, NY: Free Press.
Buss, D. M., Abbott, M., Angleitner, A., Asherian,
A., Biaggio, A., Blanco-Villasenor, A., & Yang,
K.-S. (1990). International preferences in
selecting mates: A study of 37 cultures. Journal
of Cross-Cultural Psychology, 21(1), 5–47.
https://doi.org/10.1177/0022022190211001
Buss, D. M., Larsen, R. J., Westen, D., & Sem-
melroth, J. (1992). Sex differences in jealousy:
Evolution, physiology, and psychology. Psy-
chological Science, 3(4), 251–256. https://doi.
org/10.1111/j.1467-9280.1992.tb00038.x
Chamberlain, J. M., & Haaga, D. A. F. (2001).
Unconditional self-acceptance and psychologi-
cal health. Journal of Rational-Emotive and
Cognitive-Behavior Therapy, 19(3), 163–176.
https://doi.org/10.1023/A:1011189416600
Chapman, A. L. (2006). Dialectical behavior
therapy. Psychiatry (Edgmont), 3(9), 62–68.
Cherkis, J. (n.d.). The best way to save people
from suicide. Retrieved January 28, 2019, from
https://highline.huffingtonpost.com/articles/
en/how-to-help-someone-who-is-suicidal/
Cosmides, L., & Tooby, J. (1994). Origins of
domain specificity: The evolution of functional
organization. In L. Hirschfeld & S. Gelman
(Eds.), Mapping the Mind: Domain Specificity
in Cognition and Culture (pp. 85–116).
Cambridge: Cambridge University Press.
doi:10.1017/CBO9780511752902.005
Crouch, M. (1999). The evolutionary context
of postnatal depression. Human Nature,
10(2), 163–182. https://doi.org/10.1007/
s12110-999-1013-x
Crow, T. J. (2000). Schizophrenia as the price
that homo sapiens pays for language: A
resolution of the central paradox in the
origin of the species. Brain Research Reviews,
31(2–3), 118–129.
Cuijpers, P., van Straten, A., Andersson, G., &
van Oppen, P. (2008). Psychotherapy for
depression in adults: A meta-analysis of com-
parative outcome studies. Journal of Consult-
ing and Clinical Psychology, 76(6), 909–922.
https://doi.org/10.1037/a0013075
Daly, M., Wilson, M., & Weghorst, S. J. (1982).
Male sexual jealousy. Ethology and Sociobi-
ology, 3(1), 11–27. https://doi.org/
10.1016/0162-3095(82)90027-9
Darwinian Psychotherapy. (2019). Retrieved
January 23, 2019, from https://darwini-
anpsychotherapy.com
Dewsbury, D. A. (1981). Effects of novelty of
copulatory behavior: The Coolidge effect and
related phenomena. Psychological Bulletin,
89(3), 464–482. https://doi.org/10.1037/
0033-2909.89.3.464
Dudley, R. (2004). Ethanol, fruit ripening, and
the historical origins of human alcoholism in
primate frugivory. Integrative and Compara-
tive Biology, 44(4), 315–323. https://doi.org/
10.1093/icb/44.4.315
Durisko, Z., Mulsant, B. H., & Andrews, P. W.
(2015). An adaptationist perspective on the
etiology of depression. Journal of Affective
Disorders, 172, 315–323. https://doi.org/
10.1016/j.jad.2014.09.032
Ellis, A. (1994). Reason and emotion in psycho-
therapy. Secaucus, NJ: Carol Publishing
Group.
Ellis, A., & Dryden, W. (2007). The practice
of rational emotive behavior therapy
(2nd edition). New York, NY: Springer Pub-
lishing Company.
Ellis, A., Dryden, W., & DiGiuseppe, R. (2007).
The practice of rational emotive behavior
therapy. New York, NY: Springer Publishing
Company.
Ellis, B. J., & Symons, D. (1990). Sex differences
in sexual fantasy: An evolutionary psychologi-
cal approach. The Journal of Sex Research,
27(4), 527–555. https://doi.org/10.1080/
00224499009551579
Fennell, M. J., & Teasdale, J. D. (1987). Cognitive
therapy for depression: Individual differences
and the process of change. Cognitive Therapy
and Research, 11(2), 253–271. https://doi.
org/10.1007/BF01183269
Feygin, D. L., Swain, J. E., & Leckman, J. F.
(2006). The normalcy of neurosis: Evolution-
ary origins of obsessive–compulsive disorder
and related behaviors. Progress in Neuro-­
Psychopharmacology and Biological Psychia-
try, 30(5), 854–864. https://doi.org/10.1016/
j.pnpbp.2006.01.009
First, M. B., & Pincus, H. A. (2009). Diagnosis
and classification. In M. G. Gelder, N. C.
Adreasen, J. J. Lopez-Ibor, & J. R. Geddes
(Eds.), New Oxford Textbook of Psychiatry
 EVOLUTIONARY PSYCHOLOGY AND COUNSELING AND PSYCHOTHERAPY
(pp. 99–121). Oxford: Oxford University
Press.
First, M. B., Spitzer, R. L., Williams, J. B. W., &
Gibbon, M. (1997). Structured clinical inter-
view for DSM-IV SCID. Washington, DC:
American Psychological Association.
Fraser, B. (2012). Costly signalling theories:
Beyond the handicap principle. Biology &
Philosophy, 27(2), 263–278. https://doi.org/
10.1007/s10539-011-9297-8
Freud, S. (2017). The interpretation of dreams
(A. A. Brill, Trans.). Digireads.com Publishing.
George, L. K. (1989). Stress, social support, and
depression over the life-course. In K. S.
Markides & C. L. Cooper (Eds.), Aging, stress
and health (p. 241–267). John Wiley & Sons.
Gilbert, P. (2001). Evolution and social anxiety:
The role of attraction, social competition,
and social hierarchies. Psychiatric Clinics,
24(4), 723–751. https://doi.org/10.1016/
S0193-953X(05)70260-4
Gilbert, P., Bailey, K. G., & McGuire, M. T.
(2014). Evolutionary psychotherapy: Princi-
ples and outline. In P. Gilbert & K. G. Bailey
(Eds.), Genes on the couch: Explorations in
evolutionary psychotherapy (pp. 3–27). New
York, NY: Routledge.
Giosan, C. (2020). Cognitive-Evolutionary Ther-
apy for Depression. Springer, Switzerland,
https://doi.org/10.1007/978-3-030-38874-4
Giosan, C., Cobeanu, O., Mogoase, C., Mure-
san, V., Malta, L. S., Wyka, K., & Szentagotai,
A. (2014). Evolutionary cognitive therapy
versus standard cognitive therapy for depres-
sion: A protocol for a blinded, randomized,
superiority clinical trial. Trials, 15(1), 83.
https://doi.org/10.1186/1745-6215-15-83
Giosan, C., Cobeanu, O., Wyka, K., Muresan, V.,
Mogoase, C., Szentagotai, A., & Moldovan, R.
(2019, June 29). Cognitive-evolutionary ther-
apy for depression – results of a randomized
clinical trial. Presented at the 31st Annual
Human Behavior & Evolution Society Meeting,
Boston, MA. Retrieved from http://meetatbu.
com/hbes19/ (Accessed 1 September 2019).
Giosan, C., Cobeanu, O., Wyka, K., Muresan,
V., Mogoase, C., Szentagotai, A., Malta, L. S.,
Moldovan, R. (2020). Cognitive evolutionary
therapy versus standard cognitive therapy for
depression: A single‐blinded randomized
clinical trial. Journal of Clinical Psychology;
1–14. https://doi.org/10.1002/jclp.22991
19
Giosan, C., Muresan, V., & Moldovan, R.
(2014). Cognitive evolutionary therapy for
depression: A case study. Clinical Case
Reports, 2(5), 228–236. https://doi.org/
10.1002/ccr3.131
Giosan, C., & Wyka, K. (2009). Is a successful
high-K fitness strategy associated with better
mental health? Evolutionary Psychology,
7(1), 28–39. https://doi.org/10.1177/
147470490900700104
Giosan, C., Wyka, K., Mogoaşe, C., Cobeanu,
O., & Szentagotai, A. (2018). The Evolution-
ary Fitness scale: A measure of the independent
criterion of fitness. EvoS Journal: The Journal of
the Evolutionary Studies Consortium, 7(1),
181–205.
Giphart, R., & van Vugt, M. (2018). Mismatch:
How our stone age brain deceives us every
day (and what we can do about it). London:
Robinson.
Glantz, K., & Moehl, M.-B. (2000). Reluctant
males: Evolutionary perspectives on male
psychology in couples therapy. In P. Gilbert &
K. G. Bailey (Eds.), Genes on the couch:
Explorations in evolutionary psychotherapy
(pp. 176–195). New York: Routledge.
Glantz, K., & Pearce, J. (1989). Exiles from
Eden: Psychotherapy from an evolutionary
perspective (1st edition). New York, NY: W.
W. Norton & Co. Inc.
Glenn, A. L., Kurzban, R., & Raine, A. (2011).
Evolutionary theory and psychopathy. Aggres-
sion and Violent Behavior, 16(5), 371–380.
https://doi.org/10.1016/j.avb.2011.03.009
Gutiérrez, F., Gárriz, M., Peri, J. M., Ferraz, L.,
Sol, D., Navarro, J. B., & Valdés, M. (2013).
Fitness costs and benefits of personality dis-
order traits. Evolution and Human Behavior,
34(1), 41–48. https://doi.org/10.1016/
j.evolhumbehav.2012.09.001
Hagen, E. H. (1999). The functions of postpar-
tum depression. Evolution and Human
Behavior, 20(5), 325–359. https://doi.org/
10.1016/S1090-5138(99)00016-1
Hald, G. M. (2006). Gender differences in por-
nography consumption among young het-
erosexual Danish adults. Archives of Sexual
Behavior, 35(5), 577–585. https://doi.org/
10.1007/s10508-006-9064-0
Hatfield, E., & Sprecher, S. (1995). Men’s and
Women’s Preferences in Marital Partners in the
United States, Russia, and Japan. Journal of
20 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Cross-Cultural Psychology, 26(6), 728–750.
https://doi.org/10.1177/002202219502600613
Hertler, S. C. (2014). A review and critique of
obsessive-compulsive personality disorder
etiologies. Europe’s Journal of Psychology,
10(1), 168–184. https://doi.org/10.5964/
ejop.v10i1.679
Jacobson, J. D., Lehman, K. A., Stites, B. A.,
Karwoski, L., Stroupe, N., Steidtmann, D., &
Ilardi, S. (2007). Therapeutic lifestyle change
for depression: Results of a randomized con-
trolled trial. Presented at the Annual confer-
ence of the Association for Behavioral and
Cognitive Therapies, Philadelphia, PA.
Retrieved from http://tlc.ku.edu/sites/tlc.
drupal.ku.edu/files/files/2007ABCTPoster.pdf
(September 1 2018).
John, S., & Segal, D.L. (2015). Case Conceptual-
ization. In R.L. Cautin & S.O. Lilienfeld (Eds.),
The Encyclopedia of Clinical Psychology.
Chichester: Wiley. doi:10.1002/
9781118625392.wbecp106
Jorn, A. (2016, May 17). Rational emotive
behavior therapy. Retrieved November 10,
2019, from //psychcentral.com/lib/rational-
emotive-behavior-therapy/
Karwoski, L., Prohaska, J. A., Lehman, K. A.,
Stites, B. A., Steidtmann, D., & Ilardi, S. S.
(2005). Therapeutic Lifestyle Change (TLC)
For Depression: Initial Results with a Severely
Depressed Outpatient Population. Poster
presented at the annual meeting of the
American Psychological Association, Wash-
ington, D.C., August 18–21, 2005.
Kirillova, G. P., Vanyukov, M. M., Kirisci, L., &
Reynolds, M. (2008). Physical maturation,
peer environment, and the ontogenesis of
substance use disorders. Psychiatry Research,
158(1), 43–53. https://doi.org/10.1016/j.
psychres.2007.02.017
Kriegman, D. (1998). Interpretation, the uncon-
scious, and psychoanalytic authority: Toward
an evolutionary, biological integration of the
empirical/scientific method with the field-
defining, empathic stance. In R. F. Bornstein &
J. M. Masling (Eds.), Empirical perspectives on
the psychoanalytic unconscious (pp. 187–272).
Washington, DC: American Psychological
Association.
Kriegman, D. (2000). Evolutionary psychoanal-
ysis: Toward an adaptive, biological perspec-
tive on the clinical process in psychoanalytic
psychotherapy. In P. Gilbert & K. G. Bailey
(Eds.), Genes on the couch: Explorations in
evolutionary psychotherapy (pp. 71–92).
New York, NY: Routledge.
Krupnik, V. (2014). A novel therapeutic frame
for treating depression in group Treating
Depression Downhill. Sage Open, 4(1),
2158244014523793. https://doi.org/
10.1177/2158244014523793
Krupnik, V. (2015). Integrating EMDR into an
evolutionary-based therapy for depression:
A case study. Clinical Case Reports,
3(5), 301–307. https://doi.org/10.1002/
ccr3.228
Krupnik, V. (2018). Differential effects of an
evolutionary-based EMDR therapy on depres-
sion and anxiety symptoms: A case series
study. Journal of EMDR Practice and
Research, 12(2), 46–57. https://doi.org/
10.1891/1933-3196.12.2.46
Kuyken, W., Padesky, C. A., & Dudley, R. (2008).
The science and practice of case conceptual-
ization. Behavioural and Cognitive Psycho-
therapy, 36(6), 757–768. https://doi.org/
10.1017/S1352465808004815
Lam, D., & Gale, J. (2000). Cognitive behaviour
therapy: Teaching a client the ABC model – the
first step towards the process of change. Jour-
nal of Advanced Nursing, 31(2), 444–451.
https://doi.org/10.1046/j.1365-2648.
2000.01280.x
Li, N. P., Smith, A. R., Griskevicius, V., Cason, M. J.,
& Bryan, A. (2010). Intrasexual competition
and eating restriction in heterosexual and
homosexual individuals. Evolution and Human
Behavior, 31(5), 365–372. https://doi.
org/10.1016/j.evolhumbehav.2010.05.004
Loerinc, A. G., Meuret, A. E., Twohig, M. P.,
Rosenfield, D., Bluett, E. J., & Craske, M. G.
(2015). Response rates for CBT for anxiety
disorders: Need for standardized criteria.
Clinical Psychology Review, 42, 72–82.
https://doi.org/10.1016/j.cpr.2015.08.004
Markowitz, J. C. (1994). Psychotherapy of dys-
thymia. The American Journal of Psychiatry,
151(8), 1114–1121. https://doi.org/10.1176/
ajp.151.8.1114
Marks, I. F. M., & Nesse, R. M. (1994). Fear and
fitness: An evolutionary analysis of anxiety
disorders. Ethology and Sociobiology, 15(5),
247–261. https://doi.org/10.1016/0162-
3095(94)90002-7
 EVOLUTIONARY PSYCHOLOGY AND COUNSELING AND PSYCHOTHERAPY
Martiny, K., Lunde, M., Undén, M., Dam, H., &
Bech, P. (2005). Adjunctive bright light in
non-seasonal major depression: Results from
clinician-rated depression scales. Acta Psychi-
atrica Scandinavica, 112(2), 117–125. https://
doi.org/10.1111/j.1600-0447.2005.00574.x
Mayers, A. G., & Baldwin, D. S. (2006). The
relationship between sleep disturbance and
depression. International Journal of Psychiatry
in Clinical Practice, 10(1), 2–16. https://doi.org/
10.1080/13651500500328087
McGaffin, B. J., Lyons, G. C. B., & Deane, F. P.
(2013). Self-forgiveness, shame, and guilt in
recovery from drug and alcohol problems.
Substance Abuse, 34(4), 396–404. https://
doi.org/10.1080/08897077.2013.781564
McGuire, M., & Troisi, A. (1998). Darwinian psy-
chiatry. Retrieved from www.oxfordclini-
calpsych.com/view/10.1093/med:psych/
9780195116731.001.0001/med-
9780195116731 (Accessed 1 September 2019).
Mehta, S., Abed, R., Figueredo, A. J., Aldridge,
S., Balson, H., Meyer, C., & Palmer, R. (2011).
P02-113 – Eating disorders and intrasexual
competition: Testing an evolutionary hypoth-
esis among young women. European Psy-
chiatry, 26, 709. https://doi.org/10.1016/
S0924-9338(11)72414-2
Michels, R. (1997). Psychotherapeutic
approaches to the treatment of anxiety and
depressive disorders. The Journal of Clinical
Psychiatry, 58(Suppl 13), 30–32.
Miller, S., Wampold, B., & Varhely, K. (2008).
Direct comparisons of treatment modalities
for youth disorders: A meta-analysis. Psycho-
therapy Research: Journal of the Society for
Psychotherapy Research, 18(1), 5–14. https://
doi.org/10.1080/10503300701472131
Molina, J. D., López-Muñoz, F., Stein, D. J.,
Martín-Vázquez, M. J., Alamo, C., Lerma-
Carrillo, I., & Calle-Real, M. de la. (2009).
Borderline personality disorder: A review and
reformulation from evolutionary theory.
Medical Hypotheses, 73(3), 382–386. https://
doi.org/10.1016/j.mehy.2009.03.024
Motto, J. A. (1976). Suicide prevention for
high-risk persons who refuse treatment.
Suicide & Life-Threatening Behavior, 6(4),
223–230.
Nesse, R. M. (1991). What good is feeling bad?
The Sciences, 31(6), 30–37. https://doi.
org/10.1002/j.2326-1951.1991.tb02346.x
21
Nesse, R. M. (1998). Emotional disorders in evo-
lutionary perspective. British Journal of Medi-
cal Psychology, 71(4), 397–415. https://doi.org/
10.1111/j.2044-8341.1998.tb01000.x
Öhman, A. (1986). Presidential address: Face the
beast and fear the face: Animal and social
fears as prototypes for evolutionary analyses
of emotions. Psychophysiology, 23, 737–747.
Oliver, M. B., & Hyde, J. S. (1993). Gender dif-
ferences in sexuality: A meta-analysis. Psy-
chological Bulletin, 114(1), 29–51.
O’Reilly, T., Dunbar, R., & Bentall, R. (2001).
Schizotypy and creativity: An evolutionary
connection? Personality and Individual Dif-
ferences, 31(7), 1067–1078. https://doi.org/
10.1016/S0191-8869(00)00204-X
Peet, M., & Horrobin, D. F. (2002). A dose-
ranging study of the effects of ethyl-­
eicosapentaenoate in patients with ongoing
depression despite apparently adequate
treatment with standard drugs. Archives of
General Psychiatry, 59(10), 913–919.
Prunetti, E., Bosio, V., Bateni, M., & Liotti, G.
(2013). Three-week inpatient Cognitive Evolu-
tionary Therapy (CET) for patients with person-
ality disorders: Evidence of effectiveness in
symptoms reduction and improved treatment
adherence. Psychology and Psychotherapy:
Theory, Research and Practice, 86(3), 262–279.
https://doi.org/10.1111/j.2044-8341.
2011.02060.x
Sarracino, D., Dimaggio, G., Ibrahim, R.,
Popolo, R., Sassaroli, S., & Ruggiero, G. M.
(2017). When REBT goes difficult: Applying
ABC-DEF to personality disorders. Journal of
Rational-Emotive & Cognitive-Behavior Ther-
apy, 35(3), 278–295. https://doi.org/10.1007/
s10942-016-0258-7
Schmitt, D. P., Alcalay, L., Allik, J., Ault, L.,
Austers, I., Bennett, K. L., & International
Sexuality Description Project. (2003). Universal
sex differences in the desire for sexual variety:
Tests from 52 nations, 6 continents, and 13
islands. Journal of Personality and Social
Psychology, 85(1), 85–104.
Shackelford, T. K., Buss, D. M., & Bennett, K.
(2002). Forgiveness or breakup: Sex differ-
ences in responses to a partner’s infidelity.
Cognition and Emotion, 16(2), 299–307.
https://doi.org/10.1080/02699930143000202
Shapiro, F. (2017). Eye Movement Desensitiza-
tion and Reprocessing (EMDR) therapy, third
22 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
edition: Basic principles, protocols, and pro-
cedures (3rd edition). New York, NY: The
Guilford Press.
Shem, S., & Surrey, J. (1998). We have to talk:
Healing dialogues between women and
men. New York, NY: Basic Books.
Silove, D. (1998). Is posttraumatic stress disorder
an overlearned survival response? An
evolutionary-learning hypothesis. Psychiatry,
61(2), 181–190.
Singer, B. (1985a). A comparison of evolutionary
and environmental theories of erotic response
part I: Structural features. The Journal of Sex
Research, 21(3), 229–257. https://doi.org/
10.1080/00224498509551265
Singer, B. (1985b). A comparison of evolutionary
and environmental theories of erotic response
part II: Empirical arenas. The Journal of Sex
Research, 21(4), 345–374. https://doi.org/
10.1080/00224498509551275
Smith, M. L., & Glass, G. V. (1977). Meta-analysis
of psychotherapy outcome studies. American
Psychologist, 32(9), 752–760. https://doi.
org/10.1037/0003-066X.32.9.752
Stevens, A. (1982). Archetype: A natural history
of the self. London: Routledge & Kegan Paul.
Stevens, A. (1999). On Jung: Updated edition
(Updated, Subsequent edition). Princeton,
NJ: Princeton University Press.
Stevens, A. (2000). Jungian analysis and evolu-
tionary psychotherapy: An integrative
approach. In P. Gilbert & K. G. Bailey (Eds.),
Genes on the couch: Explorations in evolu-
tionary psychotherapy (pp. 93–117). New
York, NY: Routledge.
Stewart, D. N., & Szymanski, D. M. (2012). Young
adult women’s reports of their male romantic
partner’s pornography use as a correlate of
their self-esteem, relationship quality, and
sexual satisfaction. Sex Roles, 67(5), 257–271.
https://doi.org/10.1007/s11199-012-0164-0
Sullivan, R. J., & Hagen, E. H. (2002). Psychotropic
substance-seeking: Evolutionary pathology or
adaptation? Addiction, 97(4), 389–400.
Sulzer, S. H. (2015). Does ‘difficult patient’
status contribute to de facto demedicaliza-
tion? The case of borderline personality dis-
order. Social Science & Medicine, 142,
82–89. https://doi.org/10.1016/j.socscimed.
2015.08.008
Sundie, J. M., Kenrick, D. T., Griskevicius, V.,
Tybur, J. M., Vohs, K. D., & Beal, D. J. (2011).
Peacocks, Porsches, and Thorstein Veblen:
Conspicuous consumption as a sexual signal-
ing system. Journal of Personality and Social
Psychology, 100(4), 664–680. https://doi.org/
10.1037/a0021669
Taljaard, T. (n.d.). Treating depression with
tribal wisdom. Retrieved January 23, 2019,
from https://upliftconnect.com/treating-
depression-with-tribal-wisdom/
Todd, J., Cremin, I., McGrath, N., Bwanika, J.-B.,
Wringe, A., Marston, M., & Żaba, B. (2009).
Reported number of sexual partners: Com-
parison of data from four African longitudi-
nal studies. Sexually Transmitted Infections,
85(Suppl 1), i72–i80. https://doi.org/10.1136/
sti.2008.033985
Troisi, A., & McGuire, M. T. (2000). Psychother-
apy in the context of Darwinian psychiatry. In
P. Gilbert & K. G. Bailey (Eds.), Genes on the
couch: Explorations in evolutionary psycho-
therapy (pp. 28–41). Brunner-Routledge.
Retrieved from http://psycnet.apa.org/
record/2001-16368-000
Trower, P., & Gilbert, P. (1989). New theoretical
conceptions of social anxiety and social phobia.
Clinical Psychology Review, 9(1), 19–35.
Veale, D., & Gilbert, P. (2014). Body dysmorphic
disorder: The functional and evolutionary
context in phenomenology and a compassion-
ate mind. Journal of Obsessive-Compulsive
and Related Disorders, 3(2), 150–160.
https://doi.org/10.1016/j.jocrd.2013.11.005
Wampold, B. E., & Imel, Z. E. (2015). The great
psychotherapy debate: The evidence for what
makes psychotherapy work (2nd ­edition). New
York, NY: Routledge/Taylor & Francis Group.
Westen, D., Novotny, C. M., & Thompson-­
Brenner, H. (2004). The empirical status of
empirically supported psychotherapies:
Assumptions, findings, and reporting in con-
trolled clinical trials. Psychological Bulletin,
130(4), 631–663. https://doi.org/10.1037/
0033-2909.130.4.631
Wiedenmayer, C. P. (2004). Adaptations or
pathologies? Long-term changes in brain
and behavior after a single exposure to
severe threat. Neuroscience & Biobehavioral
Reviews, 28(1), 1–12. https://doi.
org/10.1016/j.neubiorev.2003.09.005
Williams, P. (2005). ‘Notes Upon a Case of
Obsessional Neurosis’. In R. J. Perelberg (Ed.),
Freud: A modern reader (pp. 177–188).
 EVOLUTIONARY PSYCHOLOGY AND COUNSELING AND PSYCHOTHERAPY
Routledge: NY https://doi.org/10.1002/
9780470713525.ch10.
Winstead, B. A., Derlega, V. J., & Rose, S. (1997).
Gender and close relationships. Thousand
Oaks, CA: Sage.
Zanette, L. Y., Hobbs, E. C., Witterick, L. E.,
MacDougall-Shackleton, S. A., & Clinchy, M.
(2019). Predator-induced fear causes PTSD-
like changes in the brains and behaviour of
23
wild animals. Scientific Reports, 9(1), 1–10.
https://doi.org/10.1038/s41598-019-47684-6
Zhang, L., Lee, A. J., DeBruine, L. M., & Jones,
B. C. (2018, December 4). Are sex differ-
ences in preferences for physical attractive-
ness and good earning capacity in potential
mates smaller in countries with greater
gender equality?. https://doi.org/10.31234/
osf.io/mtsx8

Evolutionary Psychology and
Psychiatry
Riadh Abed and Paul St John-Smith
INTRODUCTION
Psychiatry is a branch of medicine that deals
with mental disorders that manifest them-
selves through disturbances in cognition, emo-
tions, and behaviour. Like the rest of medicine
but unlike psychology (with the exception of
clinical psychology), psychiatry is an interven-
tionist discipline that aims to modify the signs
and symptoms of disorder in order to reduce/
relieve individual distress and reduce risk
(harm) to the individual and/or others. The
contemporary failure of psychiatry to make
significant progress in understanding the aetiol-
ogy of mental disorders has been characterized
as a ‘crisis’ by leading evolutionists (Brune
et al., 2012); a fact that has also been acknowl-
edged in an article in ‘Science’ that stated that
there have been no major breakthroughs in the
treatment of schizophrenia for 50 years nor
in the treatment of depression for 20 years
(Akil et al., 2010).
Mainstream psychiatry, like the rest of
medicine, focuses on proximate causation
2
and favours mechanistic explanations of
disease and disorder. However, unlike medi-
cine where human physiology provides clear
reference points for normal functioning,
psychiatry has attempted to identify disorder
and dysfunction without a coherent theory of
normal human psychology (Nesse, 2019). We
argue in this chapter that evolutionary psy-
chology and evolutionary biology can serve as
a vital basic science for psychiatry.
Despite the publication of notable evolution-
ary psychiatry texts over the last couple of dec-
ades as well as numerous scholarly articles in
peer-reviewed journals, evolutionary thinking
has remained underappreciated by mainstream
psychiatry (e.g. Brune, 2015; Del Giudice,
2018; McGuire and Troisi, 1998; Nesse, 2019;
Stevens and Price, 2000a). Although a plu-
ralistic and multi-level approach to causality
in mental health remains essential (Kendler,
2008), the current pluralism is unconstrained
and lacks any recognizable framework (Abed,
2000). While it is recognized that all mental
phenomena are mediated by physical events
 EVOLUTIONARY PSYCHOLOGY AND PSYCHIATRY
in the brain, the phenotypic end-products of
interest to psychiatry cannot be understood
by examining the behaviour of neurons alone
(a situation compared to trying to understand
the mechanics of bird flight through the study
of feathers (Marr, 1982)).
We propose evolution as being ideally
placed to guide psychiatrists in determining
what the phenotypic end-products of neuro-
biological systems constitute. Such evolu-
tionary emphasis on function can provide the
scientific basis for expanding the concept of
the biological to encompass the psychologi-
cal, social, and cultural domains (Abed and
St John-Smith, 2016). Hence, in contrast with
mainstream biological psychiatry’s narrow
‘decontextualized’ view of mental disorder
as brain disorder (Andreasen, 1984), evolu-
tionists consider the environmental context to
be vital in determining the existence of men-
tal disorder (Nesse, 2019).
THE CONCEPT OF MENTAL DISORDER
Despite its widespread adoption within psy-
chiatry and medicine generally, the concept
of disorder has been difficult to define with
precision (Nesse, 2001). One influential evo-
lutionary proposal is that mental disorder
represents a hybrid concept, with a biological
and a socio-cultural component; a ‘harmful
dysfunction’ (HD) (Wakefield, 1992).
Accordingly, the biological component of any
disorder is the failure of a biological mecha-
nism to perform its evolved function, and the
value-laden component identifies that the dys-
function inflicts harm or damage on the
affected individual as judged by socio-cultural
standards. Although Wakefield’s HD concept
has been subject to criticism (e.g. Bolton,
2007; Fulford and Thornton, 2007), it is
acknowledged to be a significant improve-
ment on existing formulations (e.g. First,
2007; Nesse, 2007). However, while the bio-
logical criterion of the failure of a system to
perform its evolved function is intellectually
25
appealing, having considerable face validity,
problems with its clinical utility linger
because our understanding of the function of
the neurobiological systems involved in
mental disorder remains poor (First, 2007). In
addition, whereas the emphasis on context is
acknowledged to be important or even vital in
determining the existence of mental disorder
(Nesse, 2007), this potentially reduces the
diagnostic inter-rater reliability subsequent to
the increased scope of subjective judgement,
generating concern for the authors of official
classification systems such as the DSM-5
(American Psychiatric Association, 2013).
Hence, while the DSM-5 accepts mental dis-
orders necessarily involve internal dysfunc-
tion and that this produces harm and/or
distress, it leaves the term ‘dysfunction’ unde-
fined. Furthermore, whereas context is con-
sidered in a range of conditions, it is excluded
in others. For example, in the DSM-5, unlike
its predecessors, low mood lasting longer than
two weeks can now be diagnosed as major
depressive disorder (MDD) following a major
bereavement (Kavan and Barone, 2014).
Del Giudice (2018) submits that a number
of facets must be recognized to avoid com-
mon errors in interpreting Wakefield’s HD
concept, including the fact that dysfunctions
can arise from a number of different causes,
both internal and external; that the concept
of dysfunction is fuzzy; and that systems can
have degrees of functionality where the line
of demarcation between function and dys-
function is unclear.
While there are undoubted benefits from
an evolutionary analysis of the concept of
mental disorder, we support Troisi’s (2015)
conclusion that evolutionary biology alone
does not resolve the central question of what
should (and should not) be categorized as a
mental disorder, as ethical, health, and social
policy considerations lie outside the remit
of evolutionary science. In other words,
it is important not only to appreciate how
evolutionary biology can help advance our
understanding of mental disorder but also to
understand its limits.
26 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
THE REMIT OF PSYCHIATRY
In addition to the DSM-5, the other major clas-
sification system of mental disorders in clinical
use throughout most of the world, outside the
United States, is the ICD-10 issued by the
World Health Organization (WHO, 1992).
Both systems endeavour to follow an atheoreti-
cal approach to the definition and differentia-
tion of mental disorder and, with the exception
of organic mental disorders, base their diagnos-
tic categories broadly on symptom clusters and
duration. Context is acknowledged in some
instances. The ICD-10 definition of mental
disorder, being more succinct than that of the
DSM, omits the assumption of an internal dys-
function, and proceeds as follows: ‘a clinically
recognizable set of symptoms or behaviours
associated in most cases with distress and with
interference with personal functioning’ (WHO,
1992: 11). Their main categories of adult
mental disorder comprise organic mental dis-
orders, mental disorders secondary to psycho-
active substance use, schizophrenia and related
disorders, mood disorders, anxiety and stress-
related disorders, behavioural syndromes asso-
ciated with physiological disturbances, and
personality and other behaviour disorders.
Other chapters deal with mental retardation,
developmental disorders, and mental disorders
of childhood and adolescence. Remarkably,
given that both the ICD and DSM are systems
based on the consensus of committees, these
categorical domains continue to demarcate
effectively the current boundaries of psychiatric
practice (Nesse and Stein, 2012). Nevertheless,
criticism remains directed against both systems
for increasing reliability at the expense of
validity (Insel, 2013).
The National Institute of Mental Health in
the United States, in an attempt to overcome
these shortcomings, proposed the Research
Diagnostic Criteria (RDoC). The four princi-
ples used to formulate the RDoC system were
explained as follows (Insel, 2013):
• A diagnostic approach based on the biology as
well as the symptoms must not be constrained
by the current DSM categories;
• Mental disorders are biological disorders involv-
ing brain circuits that implicate specific domains
of cognition, emotion, or behaviour;
• Each level of analysis needs to be understood
across a dimension of function; and
• Mapping the cognitive, circuit, and genetic
aspects of mental disorders will yield new and
better targets for treatment.
The RDoC approach is rooted in experimen-
tal neuroscience and lists five domains: posi-
tive valance systems, negative valence
systems, cognitive systems, systems for
social processes, and arousal and regulatory
systems. Each system has a number of con-
structs and these are investigated using a
number of units of analysis ranging from the
molecular level to individual behaviour. The
RDoC has been characterized as a bottom-up
approach to the classification of mental dis-
orders, grounded in the latest research in
biological sciences that can cut across exist-
ing DSM/ICD categories (Del Giudice,
2018). However, critics have raised concerns
regarding the neglect of context (above and
beyond the DSM or ICD) and neglect of the
role of evolution (Wakefield, 2014).
EVOLUTION AND CAUSALITY
The application of evolutionary thinking to
psychiatry commences by considering some
general principles that apply to all biological
phenomena. Tinbergen (1963) proposed that a
complete understanding of any biological trait
or system involves understanding its mecha-
nism, developmental history (collectively
referred to as proximate causes), phylogenetic
history, and function (referred to as ultimate or
evolutionary causes) (Table 2.1).
These are referred to as Tinbergen’s four
questions and all four apply simultaneously
to biological phenomena (Gluckman et  al.,
2009). It is acknowledged that unlike proxi-
mate causation which can directly lead to
therapeutic interventions, understanding evo-
lutionary or ultimate causation is somewhat
 EVOLUTIONARY PSYCHOLOGY AND PSYCHIATRY
Table 2.1  Tinbergen’s four questions
Developmental/historical
Proximate causation 1. Ontogeny: how does the trait develop
during the lifetime of the organism?
Evolutionary or 3. Phylogeny: what is the phylogenetic
ultimate causation history of the trait?
Source: Adapted from Nesse (2013).
removed from direct clinical applications but
is no less important. Neglecting the question of
function (ultimate causation) runs the risk of
psychiatrists inadvertently altering psycholog-
ical functioning through their interventions to
relieve distressing but adaptive states, leading
to potentially negative consequences for some
patients. It can also lead us to construct defec-
tive models of how psychopathology arises.
Focusing exclusively on the proximate
is akin to a technician’s view of a machine,
whereas considering ultimate causation as
well is more like an engineer’s view (Nesse,
2019). Hence, it may seem adequate for a busy
clinician to simply recognize the existence of
depression or anxiety in a given patient and to
dispense standard advice and treatment accord-
ingly. However, a clinician who also under-
stands why we have such emotions in the first
place and how emotional systems interact with
people’s current lives is likely to have a deeper
understanding of the patient’s emotional prob-
lems and is able to take greater account of the
patient’s circumstances that may be contribut-
ing to their current state. It also has the poten-
tial for influencing the research agenda through
testing hypotheses regarding what the normal
function is of the system that is giving rise to
psychopathology; a question that is seldom
asked by mainstream psychiatry (Brune, 2015).
CAUSAL PATHWAYS FOR THE
PERSISTENCE OF DISEASE AND
DISORDER
It is obligatory to recognize that selection
shapes vulnerability to disease and disorder
27
Characteristics of trait/system
2. Mechanism: how does it work?
4. Adaptive function: How has the trait or system
contributed to the organism’s inclusive fitness
in its natural environment?
and not disorders themselves (Nesse, 2019).
This applies throughout medicine, including
psychiatry, and stems primarily from the
demonstration that bodies and brains are a
bundle of adaptations shaped by selection
over thousands of generations to increase
reproductive success and not good health,
happiness, or longevity. The answer to the
pivotal conundrum of why evolution has left
humans so vulnerable to disease and disorder
has itself been evolving ever since it was first
posed by the founders of modern evolution-
ary medicine (Nesse and Williams, 1994).
Accordingly, pathways by which evolution-
ary processes can lead to the existence and
persistence of disease or disorder have been
proposed (Box 2.1).
These causal pathways are not mutually
exclusive and several may be implicated
concurrently or sequentially in the origin
of mental disorders. They represent a list of
ultimate causes of our vulnerability to mental
disorder. Examples of many of these causal
pathways will be given in the sections below.
These evolutionary explanations for vul-
nerability to disorder are based on the rec-
ognition that selection is unable to eliminate
all harmful mutations, and can be too slow
to respond to rapidly changing environments,
creating states of evolutionary mismatch (Del
Giudice, 2018). This concept of ‘mismatch’
is crucial for understanding and explaining
the existence of many diseases and disorders
of modernity such as obesity, metabolic syn-
drome, Type 2 diabetes, eating disorders, and
many others. Evolutionary mismatch occurs
when the environment changes too rapidly
for selection to be able to track it, resulting in
residual traits that are no longer suited to the
28 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

BOX 2.1 Pathways for the persistence of disease and disorder 

(Adapted from Gluckman et  al. (2009) and Crespi (2016); for definition of terms see glossary: www.rcpsych.
ac.uk/docs/default-source/members/sigs/evolutionary-psychiatry-epsig/evolutionary-psychiatry-glossary-2.
pdf?sfvrsn=707dd6b_2)

• Mismatch
• Life history factors
• Overactive defence mechanisms
• Co-evolutionary considerations: consequences of the arms race against pathogens
• Constraints imposed by evolutionary history
• Trade-offs
• Sexual selection and its consequences
• Balancing selection: maintaining an allele that raises disease risk
• Demographic history and its consequences
• Selection favours reproductive success at the expense of health
• Deleterious alleles
• Extremes of adaptations

new environment. Developmental mismatch 2015). Hence, LHT provides a framework

arises when circumstances alter radically
during an individual’s lifetime. For example,
moving from a state of impoverishment dur-
ing early development to a state of affluence
in adult life can increase the risk of cardiovas-
cular disease, Type 2 diabetes, and metabolic
syndrome (Gluckman and Hanson, 2006).
Furthermore, the extreme ends of functional
adaptations can become maladaptive e.g.
when adaptive personality traits are magni-
fied (Trull and Widiger, 2013). Additionally,
over-activation of useful emotional defences
(mood states and anxiety) can result in harm-
ful outcomes, leading to defence activation
disorders (Del Giudice, 2018).
It is important to understand that selection
necessitates trade-offs. Increasing one trait
is often at the expense of worsening perfor-
mance of another. For example, increasing
resistance to infections increases the risk of
autoimmune diseases. Improving nutritional
conservation increases the risk of obesity.
Trade-offs are also involved in life history
strategies. Life history theory (LHT) deals
with species-typical solutions for problems
associated with survival and reproduction that
change over an individual’s lifespan (Brune,
for understanding how organisms allocate
time and energy in achieving core biosocial
goals across the lifespan. Life history strate-
gies involve a series of trade-offs that shape
important biological developments including
the timing of sexual maturity, the number and
quality of offspring, and the length of lifespan
(Stearns, 1992).
The application of LHT demonstrates that
the trade-offs yield a spectrum of life his-
tory strategies where the trade-offs include
somatic versus reproductive effort, present
versus the future, and quality versus quan-
tity of offspring (Figure 2.1). The ‘fast’ end
of the spectrum is characterized by a shorter
lifespan, faster growth, earlier maturation
and reproduction, and a larger number of
offspring, while those at the slow end of the
life history spectrum show the opposite char-
acteristics (Del Giudice, 2018). Differences
in life history strategies are partly under
genetic control but it appears that the nature
and quality of the individual’s early environ-
ment may also be important (Belsky et  al.,
1991; Ellis et  al., 2011) (see Barbaro et  al.,
2016 for a different perspective). This ren-
ders LHT important for the understanding
 EVOLUTIONARY PSYCHOLOGY AND PSYCHIATRY
Lifetime Energy Investment
Reproductive
Effort
(Faster)
Mating
Parental Effort
Effort
(Slower)
(Faster)
Figure 2.1  Life history strategy trade-offs
of vulnerability to mental disorders (Brune,
2015; Del Giudice, 2018) (see later section
‘Evolutionary Models of Mental Disorders’).
One major insight that follows from under-
standing the evolutionary causal pathways
for the persistence of disease and disorder
is the recognition that mental distress can
arise from functional systems. Hence, an
evolutionary taxonomy of treatable (undesir-
able) mental health conditions goes beyond
harmful dysfunctions (Tooby and Cosmides,
1999). Undesirable conditions may result
from different scenarios as summarized
below (Del Giudice, 2018):
• Undesirable mental health conditions can either
arise from:
{{ harmful dysfunctions (system breakdowns); or
{{ functional mechanisms, which can be either:
• maladaptive states at population level (e.g. evo-
lutionary mismatch), or are:
• currently adaptive, but outcomes may vary,
resulting in:
{{ maladaptive outcomes at the indi-
vidual level (e.g. overactive defences,
developmental mismatches), or:
{{ adaptive outcomes at the individual
level even if considered harmful by
others (e.g. antisocial personality/psy-
chopathy).
29
Somatic Effort
(Slower)
Hence, an evolutionary analysis provides a
theoretical framework that enables us to dis-
tinguish states of mental distress and mental
disorder that arise from functional or dys-
functional systems, and also provides a more
effective way of understanding the role of
environmental context.
GENETICS AND HERITABLE RISKS
OF MENTAL DISORDERS
Taking an evolutionary perspective is tanta-
mount to turning genetics on its head. Hence,
whereas a non-evolutionary view may con-
sider specific DNA sequences as the primary
biological cause of a given trait, an evolution-
ary approach seeks to understand the selec-
tion pressures over evolutionary history that
led to the retention of these genes. So, evolu-
tionary views consider environmental influ-
ences at two distinct levels, first over
evolutionary history (leading to the shaping
of adaptations) and, second, the ontogenic
effects of the environment during the indi-
vidual’s lifetime.
Mental disorders require a degree of
heritability, and hence some genetic basis,
30 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
before becoming candidates for evolutionary
explanations. Remarkably, 55% of all coding
genes in humans are expressed in the brain.
This renders the brain a prime target for
mutations and evolutionary changes (Brune,
2015).
After considering heterogeneity and uncer-
tainty, psychiatric disorders demonstrate a
degree of heritability suggesting a moderate
degree of heritable risk. For example, 90%
of trait variation for autism can be accounted
for with genetics; bipolar disorder, 85%;
schizophrenia, 81%; unipolar depression,
37% (Kendler, 2001). Similarly, heritabil-
ity estimates for anxiety disorders range
from 30% to 45% (Hettema et  al., 2001).
Family studies (including twin and adoption
studies) provide consistent evidence that
genetic factors are involved in the presenta-
tion of these syndromes (Kendler and Eaves,
2005).
Two types of heterogeneity have been
identified in association with psychiatric
genetics: causal and clinical. Causal hetero-
geneity refers to two or more causes indepen-
dently inducing the same clinical syndrome.
Clinical heterogeneity occurs when a single
cause leads to multiple clinical syndromes
(Tsuang et al., 2003).
Natural selection does not directly select
for genes that cause disease or disorder, so
other explanations for their persistence must
be considered. Accordingly, alongside any
degree of heritability psychiatrists should
ask: ‘Why does this mental disorder exist and
persist?’ Mental disorders may be actively
maintained through a number of evolutionary
processes. These include: A) despite natural
selection e.g. (i) mutation-selection balance,
(ii) ancestral neutrality; and B) because of
natural selection, (i) balancing selection,
(ii) antagonistic pleiotropy, (iii) stabilizing
selection on continuous traits, (iv) alternating
selection, and (v) functioning adaptations.
These categories are not mutually exclu-
sive, and there may be multiple mechanisms
maintaining some disorders in the population
(Durisko et al., 2016).
Differential Susceptibility
Research has demonstrated that people pos-
sessing at least one s-allele of the serotonin
transporter gene HTTLPR incur increased risk
of developing depression when facing adverse
events. However, the same variation is linked
to superior cognitive performance in several
domains and increases social conformity
(Homberg and Lesch, 2011). A balanced poly-
morphism also explains the frequency of a
particular SNP in the general population, and
why it has not been selected against. Beyond
this important concept, evolutionary theory
has aided in developing the idea that a particu-
lar SNP such as the s-allele of the 5-HTTLPR
not only confers heightened risk for depression
under unfavourable conditions, but lower risk
for depression under favourable environmental
conditions such as parental warmth and emo-
tional availability during important develop-
mental stages. This phenomenon is referred to
as ‘differential susceptibility’ (Belsky, 1997;
Pluess and Belsky, 2010), where phenotypic
plasticity occurs in response to early environ-
mental conditions, and differs radically from
genetically mediated resilience which involves
unresponsiveness to environmental conditions.
The specific phenomenon of differential
response to positive experiences is referred to
as ‘vantage sensitivity’; a concept that shows
promise in assessing the likelihood of respond-
ing to psychological interventions (de Villiers
et al., 2018).
This example serves as evidence against
simple genetic determinism and also provides
an indication that aspiring to alter genes alone
to treat disorders may not be in an individual’s
interests as differing circumstances alter the
harmfulness or benefits of such a gene.
Mutation Load and Mental
Disorder
Mutation load has been implicated in the
causation of some mental disorders (Keller
and Miller, 2006), referring to de novo
 EVOLUTIONARY PSYCHOLOGY AND PSYCHIATRY
germ-line mutations passed on from parents,
rather than somatic mutations. Because ova go
through far fewer replications than sperm,
paternal age at conception was suspected as the
primary source of de novo mutations (Crow,
2000). Paternal age is associated with increased
risk of mental disorders generally (Hare and
Moran, 1979). Mutation load is believed to play
a significant role in the causation of schizophre-
nia and this is especially the case in childhood
onset (Ahn et  al., 2014; Caplan, 2016) (for a
contrary view, see Ek et al., 2014).
For autistic-spectrum disorder (ASD),
mutation load was more significant in females
and in severe cases (Jacquemont et al., 2014).
The risk of attention deficit hyperactivity
disorder (ADHD) has been found to be posi-
tively related to paternal age (Chudal et  al.,
2015; D’Onofrio et al., 2014; Russell et al.,
2014, 2015).
In depression, no significant relationship
has been found with paternal age but there
is increased risk with maternal age, suggest-
ing prenatal stress as a factor (Del Giudice,
2018). In eating disorders and obsessive–
compulsive disorder (OCD), the relationship
with mutation load remains inconclusive
(Del Giudice, 2018).
Conversely, young paternal and maternal
age is also related to the risk of a range of
mental disorders in offspring. This, however,
is not related to mutation load but rather to
heritability of fast life history strategies, as
fast life history is associated with early par-
enthood in both men and women and predicts
a greater risk of fast life history spectrum dis-
orders in the offspring of young parents (most
notably ADHD and schizophrenia spectrum
disorders) (see ‘Evolutionary Models of
Mental Disorders’ section below).
Genomic Imprinting and
Mental Disorder
In diploid species such as humans, each auto-
somal gene is represented by two alleles,
with one copy inherited from each parent.
31
Usually in autosomal genes, expression occurs
from both alleles. However, in a very small
fraction, one of the two alleles is switched off
or ‘imprinted’, which may have significant
effects on behaviour, as many are expressed in
the brain (Wilkinson et  al., 2007). Genomic
imprinting represents a form of intragenomic
conflict, whereby different alleles and loci
express the fitness interest of one of the par-
ents (Crespi, 2019). Intragenomic conflict
arises from the asymmetry in the confidence
regarding parental relatedness to offspring
between the sexes. The conjecture is that
paternally expressed (maternally imprinted)
genes in an individual exert phenotypic effects
that increase fitness-related demands imposed
by offspring upon the mother, due to the lower
probability of relatedness of paternal genes
(than maternal genes) within a given brood.
This is thought to be because mothers are
always related to offspring by 50%, while the
offspring of a given female can have different
fathers (Crespi, 2019).
Contrastingly, maternally expressed (pater-
nally imprinted) genes are predicted to exert
the reverse effect, namely, lower demands
imposed on mothers. Hence, sometimes incre-
mental investment will be favoured by paternal
genes but resisted by maternal genes (Haig,
2014). Intriguingly, maternal gene imprinting
(paternal expression) may be one cause for the
underdevelopment of the ‘social brain’, gener-
ating a higher risk of ASD, whereas the pater-
nal gene imprinting (maternal gene expression)
may predispose to hyper-­development of the
social brain and increased risk of schizophre-
nia and related psychosis (Crespi, 2019) (see
section ‘Schizophrenia Spectrum Disorders
(SSDs)’, para. E, below).
EVOLUTIONARY MODELS OF
MENTAL DISORDERS
In contrast to the avowedly atheoretical
approach of the DSM/ICD systems described
above and the bottom-up biological approach
32 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
of the RDoC, evolutionary frameworks for
the classification of mental disorder are
top-down systems with explicit theoretical
assumptions. They tend to utilize high-level
organizing principles derived from evolu-
tionary insights regarding the adaptive sig-
nificance of various brain systems. Such a
top-down approach remains compatible with
a range of existing non-evolutionary
approaches (Del Giudice, 2018). According
to Del Giudice (2018), any coherent frame-
work for mental disorder (evolutionary or
otherwise) should meet four main challenges:
explain patterns of co-morbidity; address
heterogeneity within diagnostic categories;
bridge psychopathology with individual dif-
ferences; and account for developmental
features of mental disorders including life
course trajectories. The evolutionary frame-
work proposed by Del Giudice (2018) based
on LHT is more comprehensive and wide-
ranging than others such as the diametric
model of ASD and psychosis (Crespi, 2019;
‘Mutation Load and Mental Disorder’ section,
above) and the externalizing–internalizing
model (Martel, 2013). Del Giudice (2018)
suggests that his proposed framework meets
all four challenges and offers an alternative
to the existing trans-diagnostic taxonomies
of mental disorders such as the RDoC. The
most recent version of this framework has
been expanded to include a primary dimen-
sion of fast–slow life history strategy sup-
plemented by a secondary dimension of
defence-activation and hence the model has
been dubbed the FSD model (Del Giudice,
2018). It is based on a core proposition,
namely that the risk of developing a mental
disorder depends on a pattern of individual
differences that can be understood as mani-
festations of alternative life history strate-
gies. Hence, moving along the fast–slow life
history dimension will increase the risk of
certain mental disorders and reduce the risk
of others e.g. fast life history strategies
increase the risk of psychosis while reducing
the risk of autism, and vice versa. The FSD
model generates three clusters of disorders:
F-type, S-Type, and D-type. The system is
currently aimed at use by researchers rather
than clinicians and it does not currently
accommodate organic mental disorders or
mental handicap.
EVOLUTIONARY THINKING ABOUT
SELECTED PSYCHIATRIC DISORDERS
It is important to note that due to the dual
problems of heterogeneity and co-morbidity
that beset current classification systems (Del
Giudice, 2018), none of the evolutionary
theories discussed in this section can account
for the full range of the conditions they pur-
port to explain. Heterogeneity in this context
refers to the likelihood that most common
mental disorders are a collection of disparate
conditions that share certain clinical features
but may differ in their causation.
Depression
Sadness is universally recognized as the
normal emotional response to loss, setbacks,
and reversals in life (Horowitz and Wakefield,
2007). Unlike anxiety (a state of vigilance
designed to detect and deal with risk and pre-
vent/reduce harm), there is no consensus on
the function(s) of sadness. Depressive disor-
ders are marked by a severe negative mood
with an inability to experience pleasure. In
addition to low mood or anhedonia lasting a
minimum of two weeks the DSM-5 requires
the existence of four or more symptoms (loss
or gain in weight, insomnia or hypersomnia,
agitation or retardation, fatigue/loss of energy,
feelings of worthlessness or inappropriate
guilt, poor concentration or indecisiveness,
and thoughts of death and suicide) for a diag-
nosis of major depressive disorder (MDD)
(American Psychiatric Association, 2013).
Although the DSM-5 treats MDD as a unitary
condition, the application of its criteria allows
for a wide variety of combinations where
 EVOLUTIONARY PSYCHOLOGY AND PSYCHIATRY 33

individual patients can share few or even no
symptoms (Fried and Nesse, 2015).
Although many accept that depressive dis-
orders are a highly heterogeneous collection
of conditions (Akiskal and McKinney, 1975;
Brune, 2015; Gilbert, 2006; Rantala et  al.,
2018), most evolutionary theories of depres-
sion still treat it as if it was a unitary con-
dition with a single explanation. Depression
remains one of the most common mental
disorders in clinical practice, with a lifetime
risk in the US population that exceeds 15%
(Blazer et al., 1994) and striking at increas-
ingly younger ages (Rottenberg, 2014). The
increase in prevalence of depression in mod-
ern societies most probably results from evo-
lutionary mismatch (Brune, 2015; Rantala
et  al., 2018; Rottenberg, 2014). As in most
other defence activation disorders, depres-
sion has a higher prevalence in females with
an overall F:M ratio of around 2:1. The higher
female risk is contributed to by higher levels
of neuroticism, sensitivity to social rejection,
and interpersonal stressors (Del Giudice,
2018). Interestingly, and unlike most other
mental disorders such as schizophrenia,
autism, and anorexia nervosa, patients with
depression show rates of reproductive suc-
cess very close to that of the general popula-
tion, with males at 90% and females at 100%
(Power et  al., 2013). Depression occurs at
both ends of the fast–slow life history contin-
uum, with a fast life history subgroup of both
males and females having early puberty and
a slow life history subgroup (mainly males)
having late puberty (Del Giudice, 2018).
Hence, depression is not so much a slow life
history strategy as a ‘slowing down’ strategy
that can occur across the life history strategy
spectrum (Brune, 2015).
Although there is lack of agreement on the
precise function of low mood, most evolu-
tionists agree that the capacity for low mood
has been shaped by selection because of its
contribution to inclusive fitness in the ances-
tral environment. Disagreements between
evolutionists arise where some consider
the extremes or persistence of low mood as
maladaptive and/or dysfunctional, while oth-
ers consider the whole range of low mood
including the extremes of depression as
adaptations. Broadly speaking, one can clas-
sify evolutionary theories of depression into
social and non-social theories (Gilbert, 2006)
(Box 2.2).
Depression primarily occurs in social or
interpersonal contexts and is less frequently
associated with events in non-social domains
(Brune, 2015). Evolutionary formulations
suggest explanations for the observed female

BOX 2.2  Evolutionary theories of depression 

Social Evolutionary Theories

1 Theories based on attachment theory (Bowlby, 1980).
2 Theories on social competition and social rank (Price et al., 1994).
3 Social navigation hypothesis (Watson and Andrews, 2002).
4 Social risk theory (Allen and Badcock, 2003).
5 Depression as bargaining (Hagen, 2003).
6 Analytical rumination hypothesis (Andrews and Thomson, 2009).

Non-social Evolutionary Theories

1 Theories of resource conservation (Nesse, 2019).
2 Depression as immune response, defence against pathogens, starvation (see Rantala et al., 2018 for
a review).
34 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
preponderance in depression as being related
to ‘female fitness’, which appears much more
dependent on securing support from others
compared to males (Troisi, 2001).
The social competition and rank theories
propose that depression is part of a strategy
of subordination associated with decline in
social standing or rank and where further
contest is judged to be futile or even risky.
The low mood serves the dual function of
signalling helplessness and submission both
to dominants and to potential helpers. It also
stops the individual from resuming competi-
tion too quickly (Price et al., 1994). However,
if social competition lies at the root of depres-
sion, males would be expected to be at higher
risk of depression given the higher fitness
costs incurred by males as a result of status
setbacks (Brune, 2015).
According to attachment theory, the low
mood of depression bears a distinct resem-
blance to the phase of despair that occurs in
an infant after prolonged separation from its
main carer, which involves reduced activity
and vocalization as well as disengagement
from its environment (Bowlby, 1980). This
suggests that depression is an evolved strat-
egy that is activated by the disruption of sig-
nificant attachment bonds.
The social risk theory focuses on the risk
of social exclusion, which would have had
grave consequences in the ancestral environ-
ment (Allen and Badcock, 2003). The uncon-
scious and subtle calculation of the quotient
of one’s social value to social burden will sig-
nal the risk of exclusion if this drops to a crit-
ical level. This will trigger a depressive state
designed to conserve energy and help build
up future potential social value to others; it
also predicts increased suicide risk once the
quotient drops below one (Brune, 2015).
The analytical rumination hypothesis
proposes that depressive rumination is an
adaptation designed to solve complex social
dilemmas (Andrews and Thomson, 2009).
This is supported by the finding that low
mood facilitates complex decision-making
(von Helversen et al., 2011).
An influential non-social evolutionary theory
proposes that low mood is adaptive for dis-
engaging from unattainable goals. However,
depressive disorder arises when the goals are
too important to be abandoned and the indi-
vidual becomes trapped in an unwinnable
situation (Nesse, 2019).
More recently, Rantala et  al. (2018) pro-
posed a subtyping of MDD, based on an
evolutionary framework, with 12 distinct
conditions each with its own proximate and
ultimate causal profile. According to this
model, MDD cannot be explained by a single
theory and is consistent with the widespread
view that depression is a heterogeneous dis-
order. These include infection, long-term
stress, hierarchy conflict, grief, loneliness,
traumatic experiences, post-partum events,
romantic rejection, the season, chemicals,
somatic disease, and starvation.
While many of the theories briefly described
above are reasonably parsimonious accounts of
known facts about depression, the fact remains
that few of their predictions have been empiri-
cally tested (Hagen, 2011). Unfortunately, the
same can be said about many of the evolution-
ary theories regarding other mental disorders.
The current dearth of data in the field remains
an important obstacle to the integration of evo-
lutionary thinking into mainstream psychiatry.
Nevertheless, the evolutionary perspective
is crucial for the formulation of appropriate
questions and examination of existing data as
well as for the collection of new information
on mental disorders.
Schizophrenia Spectrum
Disorders (SSDs)
According to DSM-5, SSDs include schizo-
phrenia (requires a minimum of six months of
symptoms), schizophreniform disorder (up to
six months), brief psychotic episode (up to one
month), schizoaffective disorder, drug-induced
psychosis, and catatonia. DSM-5 requires two
or more of the following for a diagnosis
of schizophrenia: delusions, hallucinations,
 EVOLUTIONARY PSYCHOLOGY AND PSYCHIATRY
disorganized thinking, disorganized behav-
iour, and negative symptoms. In addition, a
number of specifiers should be applied for the
diagnosis to be made (American Psychiatric
Association, 2013).
Although it was once believed that schizo-
phrenia occurs uniformly across the world,
affecting 1% of the population, it is now
recognized that this view is erroneous and
that schizophrenia varies significantly in its
prevalence (McGrath, 2006). Some studies
have found a 30-fold difference in prevalence
(0.1–3%) (Kinney et al., 2009). The average
incidence is suggested to be between 0.2–
0.6 per 1,000 (Brune, 2015). The sex ratio
shows a male preponderance of around 1.4:1
(McGrath, 2006).
Schizophrenia is highly heritable, with
monozygotic (MZ) twins having a 48%
concordance compared to 17% for dizy-
gotic (DZ) twins, and the relative risk shows
a progressive reduction with increasing
genetic distance (Owen et  al., 2007). The
persistence of schizophrenia within human
populations, a condition that strikes at the
peak of reproductive years and has a dev-
astating effect on reproductive success, is
a puzzle that has exercised evolutionists
and has resulted in a diversity of evolution-
ary hypotheses (Brune, 2015). Power et  al.
(2013) found that males with schizophrenia
had fertility rates 23% and females 47% that
of the general population. Patients’ brothers
also showed highly reduced fertility whereas
sisters showed a slightly increased fertility.
Hence, schizophrenia is associated with the
lowest rates of fertility compared to all other
common mental disorders. We list below a
number of evolutionary formulations for
SSDs.
(a) Evolutionary by-product models:
1 The laterality and language model of schizo-
phrenia: schizophrenia arising from disrupted
lateralization of the brain with the failure of
the hemispheric dominance for language is one
of the best-known by-product models (Crow,
1997). Although this is supported by reduced
35
hemispheric asymmetry in schizophrenic patients
and increased levels of ambidexterity in children
who later develop psychosis, these findings can
be explained equally well through mutation
load and developmental stress (Yeo et al., 1999).
Moreover, genome-wide studies demonstrate
that SSDs are not the result of the action of single
or a small number of genes but the cumulative
effect of thousands of common and rare variants
(Plomin, 2018).
2 The lipid metabolism hypothesis: this hypothesis
proposes that changes in lipid metabolism within
the human lineage enabled the development
of creativity, which explains the flourishing of
culture, including art and religion, over the last
50,000 years. According to this hypothesis, SSDs
are the by-product of these newly evolved meta-
bolic pathways (Horrobin, 2001). Horrobin’s ideas
on the role of lipid metabolism in the aetiology
of schizophrenia resulted in an interest in testing
the effects of administering Omega-3 fatty acids
in high concentrations to patients, but the results
of randomized controlled trials have been incon-
sistent (National Institute for Health and Care
Excellence (NICE), 2013).
3 The social brain theory of schizophrenia: Burns’
cortical dysconnectivity hypothesis is arguably
the best developed example of the ‘social brain’
theory and also the most plausible example of
evolutionary ‘by-product’ formulations gener-
ally (Burns, 2007). Burns’ hypothesis states
that the emergence of the social brain, with
its complex and vulnerable circuits, produced a
vulnerability to aberrant connectivity. According
to this model, schizophrenia is a disorder of the
fronto-temporal and fronto-parietal circuits that
evolved in our species as a substrate for the
social brain. Schizophrenia, as a disorder of the
social brain, is consistent with a range of find-
ings that show deficits in social cognition prior
to first psychotic episode, including deficits in
recognition of facial emotions, mentalizing, and
interpersonal processes such as understanding
of fairness, reciprocity, and trust, as well as
findings following the onset of the psychosis
(Brune, 2015).
4 The ‘cliff edge’ fitness functions model: this
is based on the idea that certain adaptive
traits can overshoot their optimum, resulting
in catastrophic failure and severe maladap-
tive consequences. Nesse (2019) has suggested
that schizophrenia is intimately related to the
36 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
development of language ability and theory of
mind where the fitness peak is dangerously close
to the catastrophic cliff edge. This model is con-
sistent with a range of evolutionary formulations
including the social brain, language, and lateral-
ity, as well as the sexual selection hypothesis of
schizotypal traits.
(b) Schizophrenia as an adaptation: an early model
that has since been falsified was based on a bal-
anced polymorphism of a single gene that was
beneficial in the heterozygote state but causes
schizophrenia in homozygotes (Huxley et al., 1964).
More recently, a range of models based on group
selection have been proposed that suggest that
schizotypal traits facilitated group splitting during
human evolutionary history through magical and
paranoid thinking as well as idiosyncratic behav-
iour which can lead to messianic leadership and
group fission (Stevens and Price, 2000a, 2000b).
Other formulations focused on the shaman as the
self-sacrificing equivalent of the sterile castes in
social insects that produces group cohesion and
solidarity through magical thinking, possession
states, and religious ritual which is maintained
through group selection (Polimeni, 2012). However,
while these theories draw attention to the fascinat-
ing similarities between religious phenomenology
and psychosis, they remain highly speculative.
(c) Mismatch model: the outgroup intolerance
hypothesis is an attempt to provide an explana-
tory framework for a range of epidemiological
findings pointing to wide variation in the inci-
dence and prevalence of SSDs. The hypothesis
proposes that schizophrenia arises as the result of
a mismatch between the social brain as shaped
by evolution and the novel social conditions of
the post-Neolithic that involve living in large
settlements and regularly encountering strangers
(outgroup members). The hypothesis can provide
an explanation for (i) the higher risk in migrants
and especially second-generation migrants and
migrants who are racially and/or ethnically sali-
ent; (ii) increased risk of schizophrenia that is
inversely related to same-group ethnic density in
a given locality; (iii) the increased risk to individu-
als who have grown up in cities; and (iv) the puta-
tive low risk of schizophrenia in hunter-gatherer
societies (Abed and Abbas, 2011, 2014).
(d) Sexual selection model of creativity of schi-
zotypal traits: these hypotheses are based on
the proposal that schizophrenia is the extreme,
low-fitness end of a range of sexually selected
characteristics that include creativity, emotional
expressiveness, and superior mentalizing ability
(Nettle, 2001; Shaner et  al., 2004). Hence the
sexual selection model proposes that SSDs are
the maladaptive outcome of adaptive but risky
mating strategies (Del Giudice, 2018). This model
is also compatible with the view of SSDs as a fast
life history spectrum disorder (see para. f below).
The model is consistent with a range of findings
including the slight increase in fertility in sisters,
but it is rather difficult to reconcile with the find-
ing of a dramatic reduction in fertility in brothers
(Power et al., 2013).
(e) The diametrical model of psychosis (including
schizophrenia) and autism: in this model autistic-
spectrum disorders (ASD) and psychotic-spectrum
conditions (including schizophrenia (SSD)) repre-
sent two major suites of disorders of human cog-
nition, affect, and behaviour that involve altered
development and function of the social brain
(Crespi and Badcock, 2008). The model is based on
evidence that large sets of phenotypic traits exhibit
diametrically opposite phenotypes in ASD versus
psychotic-spectrum conditions, with a focus on
schizophrenia. These include constrained growth in
psychotic-spectrum disorders as opposed to over-
growth in ASD and underdeveloped social cogni-
tion in ASD as opposed to its hyper-development
in the psychotic spectrum (the reverse is the case
for mechanistic cognition resulting in the psychosis
spectrum being hypermentalistic/hypomechanis-
tic and the reverse is the case in ASD). The role
of genomic imprinting in this phenomenon has
already been alluded to in the section ‘Genomic
Imprinting and Mental Disorder’, above. The dif-
ferent cognitive biases of SSD and ASD proposed
in the diametric model have received considerable
empirical support (Abu-Akel et  al., 2015; White
et  al., 2016). However, the overlap between ASD
and SSD (co-morbidity) remains a challenge for
this model (Chisholm et al., 2015).
(f) Life history theory and SSDs: broadly speaking,
positive schizotypy, characterized by odd beliefs,
magical thinking, unusual perceptual experiences,
and paranoid thoughts, associated with hyper-
mentalizing, enhanced creativity, and unrestricted
socio-sexuality, fits the pattern of fast life history
strategy. This is also consistent with the asso-
ciation of positive schizotypy with aggression,
impulsivity, and sensation-seeking as well as
early maturation (Del Giudice, 2018). Negative
schizotypy, on the other hand, characterized by
 EVOLUTIONARY PSYCHOLOGY AND PSYCHIATRY
lack of social engagement, flat affect, and social
anxiety with paranoid tendencies that tends to
overlap with autistic traits, is associated with late
maturation in males but not in females, and is
consistent with a slow life history strategy (Kaiser
and Gruzelier, 1999). It is clear that the diametric
model of psychosis and ASD as well as the sexual
selection hypothesis both fit the fast life history
strategy model.
Drug and Alcohol Addictions
Examining substance abuse from an evolution-
ary perspective offers explanatory advantages
in illuminating a wide range of biological,
psychological, and social facts and mecha-
nisms in substance misuse (St John-Smith
et al., 2013). Evolutionary models are unique
in that they emphasize the effects that drugs
had on fitness over human evolution. For sub-
stance abuse, a seemingly maladaptive trait, to
persist, there must be either a ‘trade-off’ where
the harm is counterbalanced by a fitness ben-
efit, or substance-taking is a by-product of
other more adaptive processes. Such models
include: a) psychotropic self-medication
(pharmacological manipulation of emotions);
b) pharmacophagy and infection control;
c) mismatch theory; d) increasing reproductive
fitness; e) evolutionary constraints; f) trade-
offs; g) costly signalling and handicap theo-
ries; h) placebo, ritual, and healing effects; and
i) drug use in spirituality or religion (e.g. the
role of psychedelic drug use by ‘neo-shamans’
and ‘psychonauts’). Some of these models are
conceptually similar or overlapping, are not
mutually exclusive, and may interact in unpre-
dictable ways (Orsolini et al., 2017).
Emotional pathways
Primary emotional systems evolved to pro-
duce pleasurable affects in response to propi-
tious circumstances or stimuli indicating
adaptive success, and aversive affects in
response to environmental or other threats,
indicating reduced adaptive success. Drugs
(of abuse) may be used to diminish aversive
affects (e.g. opiates) or to increase positive
37
affect (e.g. stimulants). These drugs override
the adaptive functions of the primary emo-
tional systems so individuals experience an
increase in positive affect, or decrease in
negative affect, independently of any change
in their circumstances, thus decoupling the
emotional system from environmental events,
some continuing to consume the drug despite
mounting harm because the reactions bypass
the evolved protective mechanisms used to
signal real success or danger (Nesse, 2019).
Mismatch
The hijack hypothesis implies that a range of
drugs of abuse effectively commandeer the
neural reward circuitry in the mesolimbic
reward pathway as a result of mismatch as the
contemporary abundance of potent psychoac-
tive substances is a recent and novel phenome-
non that was not present and therefore could not
have occurred in the ancestral environment.
Human–plant co-evolutionary
history and the paradox of drug
reward
Plants evolved the capacity to synthesize
chemicals (nicotine, morphine, cocaine etc.)
that act as neurotoxins to deter consumption by
insects and herbivores (Sullivan et  al., 2008).
The efficacy of plant neurotoxins evolved over
400 million years and is therefore not evolu-
tionarily novel. Consequently, human physiol-
ogy can ‘identify’ plant toxins and activate
defences that involve genes, tissue barriers,
neural circuits, organ systems, and behaviours
to protect against them. Drug toxicity and aver-
sive responses (e.g. headache, sweating,
nausea, and vomiting) occur in humans so are
inconsistent with a simplistic theory of drug
reward. Consequently other mechanisms, such
as trade-offs, must be invoked as explanations.
The neurotoxin regulation hypothesis pro-
poses that the parallel consumption of both the
nutrients and neurotoxins in plants selected for
a system capable of maximizing the benefits of
plant energy extraction while mitigating the cost
of plant toxicity. The pharmacophagy hypothe-
sis proposes that the consumption of chemicals
38 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
with medicinal properties is contingent on
human–plant co-evolution. Self-medication
advantages arose when humans learned to over-
come cues of plant toxicity (e.g. bitter taste)
and consumed potentially toxic substances with
little energetic content because ingesting the
toxins in small amounts was advantageous.
Thus, the consumption of plant alkaloids could
have contributed to reproductive fitness, and
a taste for these substances could have been
selected for. It is recognized that many such
toxins are known to have anti-helminthic or
antimicrobial and antiparasitic effects.
Alcohol
Consuming ripe fruits containing small
amounts of ethanol is selectively advantageous
(Dudley, 2004), as volatile alcohols potentially
aid in olfactory localization of ripe fruit.
Herbivores developed the capacity to metabo-
lize alcohol to be able to utilize energy-rich
fruits despite the presence of alcohol. In the
ancestral environment, alcohol would have
been encountered in fermenting fruit in low
concentrations and small quantities for brief
periods in the year. Subsequent to the agricul-
tural revolution, large surpluses of fruits and
grains became available for fermentation so
alcoholic drinks were brewed up to 12–14%
and stored/traded for year-round consumption.
Much more recently, the development of dis-
tilling technology permitted the production of
far higher concentrations of alcohol. With the
rise of larger settlements and cities, having
access to alcoholic beverages may have pro-
tected against waterborne pathogens. However,
enzyme systems that evolved to process small
amounts of alcohol on an occasional basis can
now be presented with inexhaustible supplies
of highly concentrated alcohol, giving rise to a
state of mismatch (St John-Smith et al., 2013).
Cultural, psychological,
anthropological models and sexual
selection hypotheses
Some evolutionary psychological theories con-
cerning drug use suggest individuals consume
drugs to increase reproductive opportunities.
Drug use can increase reproductive fitness
because consumption may: (1) advertise bio-
logical quality, sexual maturity, or availability;
(2) decrease inhibitions in mating contexts;
and/or (3) enhance associative learning behav-
iours that in turn increase mating opportunities
(Richardson et al., 2017).
Variation in drug use susceptibility is in part
due to genetic factors; therefore, successful
drug consumption may be a costly and honest
signal of biological quality: a process of costly
signalling and sexual selection. Such risk-
taking behaviour represents a fast life history
strategy and involves future discounting (see
‘Evolutionary Models of Mental Disorders’
section above). LHT can explain the current
male preponderance in drug use, as female
drug users incur much higher fitness costs
through reduced parenting capacity, potential
teratogenic effects, and potential circumven-
tion of mate choice (Orsolini et al., 2017).
Finally, another aspect of mismatch is
that the ancient ‘evolved’ advantages of any
psychoactive substances have now poten-
tially become a liability and risk in modern
environments as cultural change is accelerat-
ing and outstrips biological adaptation. The
evolutionary perspective can help researchers
reach a functional understanding of substance
abuse and develop treatments for the various
complex underlying causes of substance mis-
use. Some of these models are conceptually
similar or overlapping and can interact in
unpredictable ways. In addition, psychoac-
tive substances, often hallucinogens which
tend not to be addictive, have been used in
various religious and cultural ceremonies
(signalling) for millennia. Some advantages
may be had from related group cohesion as
well as their action on micro-organisms and
other trade-offs discussed above.
Anorexia Nervosa (AN) and
Bulimia Nervosa (BN)
AN and BN are diagnostic categories of eating
disorders according to ICD-10 and DSM-5
 EVOLUTIONARY PSYCHOLOGY AND PSYCHIATRY
classifications. The conditions share core fea-
tures of morbid fear of fatness, distorted body
image, and a pattern of behaviour aimed at
weight reduction that includes purging, restric-
tion of food intake, or excessive exercise
(American Psychiatric Association, 2013;
WHO, 1992). AN is characterized by low body
weight with possible amenorrhea whereas BN
is associated with binge eating and a normal
body weight. Evidence demonstrates some
heritability (Bulik et  al., 2016; Yilmaz et  al.,
2015) and AN and BN share some genetic basis
(Eley et al., 2005). Notably, the epidemiology
of AN and BN demonstrates a marked female
preponderance with a female-to-male sex ratio
of 10:1 or greater (Gordon, 1990; Hudson et al.,
2007). Also, both are by far more prevalent in
developed countries compared to developing
countries, particularly when considering sub-
threshold phenotypes (Katzman et al., 2004).
Evolutionary theories for eating
disorders
A number of evolutionarily informed theo-
ries and hypotheses have been proposed. The
‘Reproductive Suppression Hypothesis’ of
AN considers eating restriction as a strategy
to delay reproduction in times of disadvanta-
geous environmental conditions by lowering
the amount of body fat to a level incompati-
ble with ovulation (Surbey, 1987; Voland and
Voland, 1989; Wasser and Barash, 1983).
Consistent with the Reproductive Suppression
Hypothesis it is reported that women who
perceive low levels of support from romantic
partners and family are prone to dieting and
do not feel ready for parenthood, suggesting
that poor environmental conditions are causal
in the development of AN (Juda et al., 2004).
Unlike the original Reproductive Suppression
Hypothesis which hypothesized the occurrence
of reproductive self-suppression, an alterna-
tive hypothesis was put forth by Mealey (2000)
where reproductive suppression was imposed
upon subordinate females by dominants.
Other evolutionary hypotheses have pos-
ited that symptoms of AN may help to cope
with famine, whereby food restriction, denial
39
of starvation, and hyperactivity could r­ epresent
an adaptive behaviour that helped ancestral
nomadic foragers to migrate from depleted
environments to more promising surroundings
in times of food shortages (Guisinger, 2003).
However, the ‘fleeing famine hypothesis’
appears to confound consequences with cau-
sation in that the features of ‘fleeing famine’
represent the consequences of starvation that
arise in AN as a result of self-imposed restric-
tion of food intake. It is of interest that the
trigger for the initiation of dieting proposed
by Guisinger (2003) is the improvement of
attractiveness and competition for mates,
which is more or less identical to the Sexual
Competition Hypothesis (see below).
It is notable that these theories focus exclu-
sively on AN where food restriction causes
low body weight, which in turn can lead to
amenorrhoea and reproductive suppression
or the starvation response, whereas this does
not occur in BN.
The Sexual Competition
Hypothesis (SCH) and LHT
The SCH is a more inclusive evolutionary
model which reconsiders the whole spectrum
of eating disorders including AN and BN
(Abed, 1998). The SCH, based on the
Darwinian theory of sexual selection, proposes
that female intra-sexual competition is the bio-
logical root for the drive for thinness, an adap-
tive response originally suited to the ancestral
environment, and that the extreme version of
this manifests in what we know as eating disor-
ders. The SCH proposes that AN and BN are
manifestations of abnormally intense female
intra-sexual competition whereby autonomous
females of reproductive age compete with each
other in the novel modern Westernized urban
environment through a strategy of ‘the pursuit
of thinness’ as a signal of youth, leading to
‘runaway female intra-sexual competition’, the
extreme version of which manifests as eating
disorders (Abed, 1998).
The SCH is based on the fact that throughout
human evolutionary history the female shape
has been a reliable indicator of the female’s
40 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
reproductive history and ­ consequently her
reproductive potential (Bovet and Raymond,
2015; Singh, 1993). Youth and good health
have always been major determinants of female
mate value not least because of the finite repro-
ductive window in humans that abruptly ends
with menopause (Buss, 1987). The visual sig-
nal for a female’s peak reproductive potential
in the ancestral environment was the female
nubile shape, which was generally short-lived
and deteriorated with the repeated cycles of
gestation and lactation (Symons, 1995).
Hence, according to SCH, female intra-
sexual competition in affluent Westernized
societies became focused on the preserva-
tion of the ‘nubile shape’ through a strategy
of the pursuit of thinness to display signs of
youth. The SCH further proposes that other
important factors serve to up-regulate the
intensity of female intra-sexual competition.
Some of the major additional factors include
(Nettersheim et  al., 2018): (a) female auton-
omy that involves the ability to make mating
decisions with relatively little interference
from kin (unlike the case in ancestral and tra-
ditional societies) (Apostolou, 2007; (b) living
in cities where abnormally large numbers of
autonomous females live in close proximity
to each other; (c) reduced fertility (birth rates)
(Vining, 1986); and (d) the ubiquity of abnor-
mally attractive youthful nubile female images
in the media that are mistaken for competitors
(Ferguson et al., 2011).
Therefore, the SCH is based on a proposed
mismatch between the design of the female’s
psychological adaptations for mate attrac-
tion and retention and for competing with
rival females, on the one hand, and the novel
circumstances of the modern urban environ-
ment, on the other.
However, intra-sexual competition alone
cannot explain the different presentations of
AN and BN. Hence, life history strategies
were considered as an added factor where BN
lies at the fast and AN on the slow end of the
life history spectrum (Abed et al., 2012).
Predictions from the SCH have been exam-
ined in a number of non-clinical studies and
have found a significant correlation between
abnormal eating behaviour and the intensity
of competition for mates (Abed et al., 2012;
Faer et al., 2005). Also, supportive evidence
has been found for the predictions that homo-
sexual men resemble heterosexual women
and lesbians resemble heterosexual men in
their concerns about physical attractiveness
and eating behaviour (Li et al., 2010). More
recently an exploratory study on anorexic and
bulimic patients supported the fast–slow life
history strategy prediction for BN and AN
and partially supported the predictions of
SCH (Nettersheim et al., 2018).
The Placebo Response and Nesse’s
Smoke Detector Principle
Placebo effects may be considered as expla-
nations of how healing and caring works
(McQueen et  al., 2013). The universality of
placebo responses suggests a likely evolution-
ary basis to the underlying mechanisms.
Placebo responses permit mammals to modify
internal processes and behaviours. Adaptive
advantages might result from the evolution of
abilities to modify our internal environment
in the light of positive evaluations of our
external environments, social interactions,
and appraisals of the future. The hypothetical
system charged with health maintenance,
shaped by evolution, has been referred to as a
‘health governor’, aspects of which are shared
across many species but which is most highly
developed in humans and operates entirely
outside conscious awareness (Humphrey and
Skoyles, 2012). Nesse (2019) stresses that
placebo responses primarily entail modifica-
tion of the body’s defences e.g. pain, nausea,
anxiety, depression, fever, coughing, vomit-
ing, and diarrhoea, rather than altering disease
processes. Hence, evolution has selected for
mechanisms that defend against injury, infec-
tion or poisoning and the regulation of these
defences is influenced by appraisals of the
environment. However, many defences appear
to be over-expressed. A signal-detection
 EVOLUTIONARY PSYCHOLOGY AND PSYCHIATRY
analysis can explain this apparent paradox.
When the cost of expressing an all-or-nothing
defence is low compared with the potential
harm it protects against, the optimal system
will express many false alarms. For example,
vomiting may cost only a few hundred calo-
ries and a few minutes, whereas not vomiting
may result in a chance, however small, of
death from poisoning. This has been dubbed
‘the smoke detector principle’ (Nesse, 2001).
The over-expression of many defences allows
that they can often be dampened without
compromising fitness. The regulation of
defences allows that otherwise ‘protective’
defences can be turned off both in situations
of extreme danger, to facilitate escape, and in
situations propitious for recovery, where they
may no longer be necessary for protection.
This may explain why pain is reduced both
when facing immediate threat and when
being cared for.
Furthermore, the goal of the attachment sys-
tem is to maintain proximity to caregivers who
would provide safety from danger. Thus, at
times of threat, the attachment system becomes
activated. Manifestations of attachment behav-
iour change with the stage of the life cycle and
attachment style, but at times of subjectively
perceived threat, which includes illness, prox-
imity and caring are sought from attachment
figures, which may come to include trusted
professional carers, and hence the placebo
response may be an emergent property of the
attachment system (Bowlby, 1980).
Other Disorders: Alzheimer’s,
Personality Disorders, and Bipolar
Disorder
People are increasingly surviving into old
age. This increase in longevity is associated
with increased levels of morbidity of both
somatic and mental disorders, among them
the dementias such as Alzheimer’s disease
(AD), during those added years. Evolutionists
consider explanatory theories for the phe-
nomenon of aging such as antagonistic
41
pleiotropy (Williams, 1957) and LHT. As AD
seems to be specific to Homo sapiens, its
existence may in part be anchored in the
adaptive changes that have occurred after
humans separated from other primates.
Evolutionary theories also take into account
issues around brain development including
the related phenomena of altriciality and
grandmothering, the evolution of ApoE and
the genome lag hypothesis. Thus, an evolu-
tionary look into AD may shed new light on
the causes and treatments of this devastating
disease (Von Gunten et al., 2018).
Others have suggested that AD is the result
of mismatch related to the vastly increased
levels in the modern environment of insulin
resistance, inflammation, and exposure to
toxins (Fox, 2018), or that AD is the result of
a trade-off between the antimicrobial effects
of amyloid beta and the damaging effects of
its sustained activation (Moir et al., 2018).
Personality disorders (PD) are defined by
DSM-5 as an enduring pattern of inner expe-
rience and behaviour that deviates markedly
from the expectations of the individual’s cul-
ture. The DSM and ICD classifications list
around a dozen different types each but they
differ in their subtyping, terminology, and cri-
teria. The five-dimension model of personal-
ity is currently widely favoured and comprises
extraversion, neuroticism, agreeableness,
conscientiousness, and openness (McCrae
and Costa, 2003). Personality disorders are
clustered into three groups with Cluster A
comprising the ‘eccentric’ PDs such as para-
noid and schizoid; cluster B ‘dramatic’ such
as antisocial and borderline PDs; and cluster
C ‘anxious’, including avoidant, dependent,
and obsessive–compulsive PDs.
Evolutionary formulations have proposed
that antisocial PDs may be an ‘adaptive’
cheating strategy that is maintained through
frequency dependent selection (e.g. Mealey,
1995). Cluster B (antisocial and borderline
PDs) has been considered to represent a fast
life history strategy while both clusters A and
C are considered slow life history disorders
(Brune, 2015). However, Del Giudice (2018)
42 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
takes a more nuanced approach and classifies
PDs as fast (antisocial and borderline) and
slow (obsessive–compulsive), with avoidant
PD as a defence activation disorder.
Bipolar disorder (BPD) has a prevalence of
between 1 and 5% of the population depend-
ing on the subtypes included. In contrast to
schizophrenia, BPD has received relatively
little attention from evolutionists. Many of the
existing evolutionary models propose some
evolutionary advantage of hypomanic traits
or even manic episodes (Del Giudice, 2018).
The manic mood has been considered the win-
ning and the depressive mood the losing pro-
grammes of the dominance system (Gilbert
et al., 2007). It is of interest to note the rela-
tively small decline in fertility associated with
BPD (85% of general population levels in
females and 75% in males) compared to the
steep decline in schizophrenia (Power et  al.,
2013). Nesse (2019) considers BPD as an
example of a malfunctioning mood regulation
system or broken ‘moodostat’. Del Giudice
(2018), applying the life history framework,
proposes that there are two distinct variants, a
fast and a slow life history strategy variant. The
fast subtype has greater links to schizophrenia
with a higher risk of psychotic symptoms and
the slower subtype has links to autism and
lower risk of psychotic symptoms.
EVOLUTION AND
PSYCHOPHARMACOLOGY
Psychopharmacological drugs became
widely available in the 1950s and have
changed many outcomes; however, psychiat-
ric disorders are so complex and heterogene-
ous that psychopharmacology alone cannot
cure every aspect of any disorder. Current
psychopharmacology is not based on evolu-
tionary insights or theories.
Highly preserved, bio-active chemicals
play fundamental roles in many processes
across virtually all life forms. They include
acetylcholine and the biogenic monoamines
as well as other groups such as amino acids,
purines, cannabinoids, and neuropeptides.
Such chemicals have been found not only
in animals, but also in plants and unicellu-
lar microorganisms (Roshchina, 2010). This
ubiquity is best explained by universal cel-
lular mechanisms, communication systems
across kingdoms, and shared evolutionary
ancestry, demonstrating the ‘thriftiness’ of
evolutionary processes and the conserva-
tion of evolved mechanisms and strategies.
Phylogenetically, it appears that these chemi-
cals and their associated enzymes existed for
a substantial period before their respective
receptor proteins.
Evolution of sophisticated nervous systems
arrived independently of the synthesis of newer
sophisticated transmitter substances, receptor
proteins, transducers, and effector proteins;
rather they evolved with improved organiza-
tion and utilization of these entities, forming
increasingly advanced and refined circuitry
via natural and sexual selection (Roshchina,
2010). There are hundreds of chemical sub-
stances that provide communication between
cells in humans, some simple monoamines,
others more complex, e.g. neuropeptides.
Knowledge of differing receptor function
in other species has aided drug development.
For example, there are important changes dur-
ing evolutionary time, related to the neuro-
transmitters/receptors and how they function
in humans. As further examples of biological
cross-reactivity, many psychotropic agents
have an action on microorganisms, includ-
ing such varied taxa as bacteria, helminths,
insects, and other parasites. The antipsychot-
ics (phenothiazines and thioxanthenes) show
antibacterial activity, exerting their activity
independently of antibiotic resistance. The ben-
zodiazepine clonazepam is anti-schistosomal
(Stohler, 1978). Monoamine oxidase inhibi-
tors, lithium, tricyclic antidepressants, and
valproic acid have a range of antimicrobial
activities (Kristiansen, 1990).
Many psychiatric conditions involve emo-
tion dysregulation, inappropriate expression
of emotions, or impaired access to one’s
 EVOLUTIONARY PSYCHOLOGY AND PSYCHIATRY
emotional life. Positive emotions developed
evolutionarily to motivate humans to take
advantage of environmental opportunities
and to recognize when we have succeeded in
doing so. Negative emotions evolved to moti-
vate humans to avoid misfortune by escaping,
attacking, or preventing harm or repair-
ing damage when it has already occurred.
Emotional reactions importantly correspond
to differences in appraisal that result from indi-
vidual differences in personal values, experi-
ences, and goals. Psychopharmacological
agents may modify these responses in ways
which have consequences beyond the sim-
ple alleviation of distress. ‘Side effects’ of
medications are sometimes consequences
of effects on attendant processes, as distinct
from the direct pharmacology, for instance a
reduction in anxiety leading to an increase in
risk taking or disinhibition.
Understanding why symptoms exist/per-
sist may enhance psychiatric management.
Treatments should be evaluated regarding
whether the index symptoms are aiding indi-
vidual coping strategies with respect to the
adverse life event which caused the lowered
mood in the first place. Importantly, pharma-
cologically reducing symptoms remains ben-
eficial, even essential, when the symptoms
are excessive or fail to serve their adaptive
purpose, and when the symptoms are not
associated with events that triggered the epi-
sode. Conversely, in cases where a depressive
episode is a functional response to adver-
sity, suppressing it unconditionally without
addressing the underlying causes might be
harmful. This is analogous to treating pain
without considering the aetiology.
Conceptualizing sickness behaviours, pain
mechanisms, and mental disorders in rela-
tion to the problems that they evolved to
solve potentially encourages practitioners
to provide treatment options that are more
effectively targeted, ensuring a patient’s
long-term well-being, though the patient’s
immediate best interests must always be
regarded as paramount (Rantala et al., 2018).
Psychopharmacology should also review the
43
side effects of medication through the lens of
evolutionary theory, potentially considering
drug interference with evolutionarily relevant
systems that might have negative conse-
quences for the individual’s ability to attain
vital biosocial goals.
LOOKING TOWARD THE FUTURE
At present, the evolutionary literature remains
largely invisible to mainstream psychiatrists.
This is partly explained by the current paucity
of evolutionarily inspired interventions but is
also influenced by a range of other factors.
These include ideological, religious, and lib-
ertarian concerns as well as factors related to
the inertia inherent in paradigm shifts (Kuhn,
1962). Whereas the religious and ideological
(primarily post-modernist, anti-science
trends) opposition to Darwinism is largely
entrenched and probably unchangeable, the
libertarian concerns arise from misconcep-
tions that should, in principle, be amenable to
modification. For example, mistaking evolu-
tionary science for social Darwinism and
assuming that evolution implies strict genetic
determinism can be countered by appropriate
scientific argument and evidence. However, it
may prove much more difficult to overcome
the anti-evolutionary position of ‘biological
reductionism’ that is currently the dominant
trend in medical and psychiatric academic
centres within the Western world.
We propose that evolutionary science pro-
vides a framework that can organize a huge
number of facts about human biology and
psychology into a coherent narrative that, in
time, will lead to insights that can give rise to
novel treatments and interventions in psychi-
atry and the rest of medicine. This can help
further our understanding of sex differences
in vulnerability to disorder, phenotypic plas-
ticity including differential susceptibility as a
result of gene–environment interactions, and
the role of life history strategies. The unique
insights evolutionary thinking brings stem
44 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
primarily from combining an understanding
of the role of ultimate causation alongside
proximate causes. Such evolutionary think-
ing has already resulted in novel interven-
tions for cancer (DeGregori, 2018).
However, a critical mass of evolutionarily
informed psychiatrists is necessary to signifi-
cantly influence the research agenda. Hence,
the first step must involve better evolutionary
education for psychiatrists both at under- and
postgraduate levels.
We suggest that trainee psychiatrists would
benefit from the following basic evolutionary
knowledge/competences:
1 An understanding of how selection shapes adap-
tations (physical and psychological traits).
2 An understanding of Tinbergen’s four causes
with special emphasis on the distinction between
proximate and ultimate causation (see ‘Evolution
and Causality’ section).
3 An understanding of the concepts of kin selection
and inclusive fitness.
4 An understanding of the evolutionary causal pro-
cesses for the persistence of disease and disorder
with special emphasis on mismatch, trade-offs,
life history strategies and sexual selection (see
‘Causal Pathways for the Persistence of Disease
and Disorder’ section).
5 An understanding of the basics of evolution-
ary genetics, including selection, mutation, drift,
intra-genomic conflict, and genomic imprinting.
Many evolutionary applications in medicine
rely on well-established methods, such as
population genetics, phylogenetic analysis,
and observing pathogen evolution. Approaches
to evolutionary questions about traits that
leave bodies vulnerable to disease are less
well developed. Strategies for formulating
questions and hypotheses remain unsettled,
and methods for testing evolutionary hypoth-
eses are unfamiliar to many in medicine.
Nesse (2011) has suggested a structure for
appropriate evolutionary research which uses
recent examples to illustrate successful strat-
egies and some common challenges. He
identifies 10 questions to consider in testing
evolutionary hypotheses. Addressing them
systematically can help minimize confusion
and errors. One of the major contributions of
evolutionary thinking is that it helps research-
ers formulate the right questions regarding the
nature of disease and disorder. Evolution also
cautions us against simplistic genetic models
and draws attention to the possible adaptive
function(s) of genes implicated in mental
disorders.
Evolution’s flagship contribution is that it
highlights the mistake of equating distress with
disease and disorder. This prompts clinicians
to consider the possible downside of treating
potentially adaptive states of defence activa-
tion in individual patients as well as to consider
the currently neglected possibility that insuffi-
cient defences (e.g. low or absent anxiety) are
also a possible source of psychopathology and
harmful dysfunction (Nesse, 2019).
Aside from future advantages in the areas
of research and classification, there are potential
benefits from utilizing evolutionary thinking
in the clinic in the present. Examples include
introducing patients with anxiety and panic
disorders to evolutionary concepts such as the
‘smoke detector principle’ (Nesse, 2019) or the
harm-avoidance model of OCD (Abed and de
Pauw, 1999). Finally, we submit that possessing
an evolutionary understanding of unique human
vulnerabilities in itself enhances empathy and
understanding, complementing the clinician’s
effectiveness (Nesse, 2019; Troisi, 2012).
ACKNOWLEDGEMENT
We are grateful to David Geaney and the
anonymous referee for reading and com-
menting on previous drafts of this chapter.
REFERENCES
Abed, R.T. (1998). The sexual competition
hypothesis for eating disorders. British Jour-
nal of Medical Psychology, 71, 525–547.
 EVOLUTIONARY PSYCHOLOGY AND PSYCHIATRY
Abed, R.T. (2000). Psychiatry and Darwinism:
Time to reconsider? British Journal of Psy-
chiatry, 177, 1–3.
Abed, R.T. & Abbas, M.J. (2011). A reformulation
of the social brain theory of schizophrenia: The
case for outgroup intolerance. Perspectives in
Biology and Medicine, 54, 132–151.
Abed, R.T. & Abbas, M.J. (2014). Can the new
epidemiology of schizophrenia help eluci-
date its causation? Irish Journal of Psycho-
logical Medicine, 31, 1–5.
Abed, R.T. & de Pauw, K.W. (1998). An evolu-
tionary hypothesis for obsessive compulsive
disorder: A psychological immune system?
Behavioural Neurology, 11(4), 245–250.
Abed, R.T. & St John-Smith, P. (2016). Evolution-
ary psychiatry: A new college special interest
group. BJPsych Bulletin, 40, 233–236.
Abed, R., Mehta, S., Figueredo, A.J., Aldridge,
S., Balson, H., Meyer, C. & Palmer, R. (2012).
Eating disorders and intrasexual competition:
Testing an evolutionary hypothesis among
young women. Scientific World Journal,
290813. doi: 10.1100/2012/290813
Abu-Akel, A.M., Wood S.J., Hansen, P.C. &
Apperly, I.A. (2015). Perspective-taking abili-
ties in the balance between autism tendencies
and psychosis proneness. Proceedings of the
Royal Society of London B, 282, 20150563.
Ahn, K., Gotay, N., Andersen, T.M., Anvari,
A.A., Gochman, P., Lee, Y., Sanders, S., Guha,
S., Darvasi, A., Glessner, J.T., Hakonarson, H.,
Lencz, T., State, M.W., Shugart, Y.Y. & Rapo-
port, J.L. (2014). High rate of disease-related
copy number variation in childhood onset
schizophrenia. Molecular Psychiatry, 19,
568–572.
Akil, H., Brenner, S., Kandel, E., Kendler, K.S.,
King, M-C., Scolnick, E., Watson, J. D. &
Zoghby, H.Y. (2010). The future of psychiat-
ric research: Genomes and neural circuits.
Science, 327(5973), 1580–1581.
Akiskal, H.S. & McKinney, W.T. (1975). Over-
view of recent research in depression: Inte-
gration of ten conceptual models into a
comprehensive clinical frame. Archives of
General Psychiatry, 32(3), 285–305.
Allen, N.B. & Badcock, P.B. (2003). The social
risk hypothesis of depressed mood: Evolu-
tionary, psychosocial and neurobiological
perspectives. Psychological Bulletin, 129,
887–913.
45
American Psychiatric Association (2013).
Diagnostic and Statistical Manual of Mental
Disorders (5th ed.). Arlington, American
Psychiatric Association.
Andreasen, N. (1984). The Broken Brain. New
York, HarperPerennial.
Andrews, P.W. & Thomson, J.A. (2009). The
bright side of being blue: Depression as an
adaptation for analyzing complex problems.
Psychological Review, 116, 620–654.
Apostolou, M. (2007). Sexual selection under
parental choice: The role of parents in the
evolution of human mating. Evolution and
Human Behavior, 28, 403–409.
Barbaro, N., Boutwell, B.B., Barnes, J.C. &
Shackelford, T.K. (2017). Genetic confound-
ing of the relationship between father
absence and age at menarche. Evolution and
Human Behavior, 38, 357–365.
Belsky, J. (1997). Variation in susceptibility to
rearing influence: An evolutionary argument.
Psychological Inquiry, 8, 182–186.
Belsky, J., Steinberg, L. & Draper, P. (1991).
Childhood experience, interpersonal develop-
ment and reproductive strategy: An evolution-
ary theory of socialization. Child Development,
62, 647–670.
Blazer, D.G., Kessler, R.C., McGonagle, K.A. &
Swartz, M.S. (1994). The prevalence and
distribution of major depression in a national
community sample: The National Comorbid-
ity Survey. American Journal of Psychiatry,
151, 979–986.
Bolton, D. (2007). The usefulness of Wake-
field’s definition for the diagnostic manuals.
World Psychiatry, 6(3), 164–165.
Bovet, J. & Raymond, M. (2015). Preferred
women’s waist-to-hip ratio variation over the
last 2,500 years. PLOS One, 10, e0123284.
Bowlby, J. (1980). Attachment and Loss: Loss
Sadness and Depression (Vol. 3). London,
Hogarth Press.
Brune, M. (2015). Textbook of Evolutionary
Psychiatry and Psychosomatic Medicine:
The Origins of Psychopathology. Oxford,
OUP.
Brune, M., Belsky J., Fabrega, H., Feierman,
H.R., Gilbert, P., Glantz, K., Polimeni, J.,
Price, J.S., Sanjuan, J., Sullivan, R., Troisi, A. &
Wilson, D.R. (2012). The crisis of psychiatry –
insights and prospects from evolutionary
theory. World Psychiatry, 11(1), 55–57.
46 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Bulik, C.M., Kleiman, S.C. & Yilmaz, Z. (2016).
Genetic epidemiology of eating disorders.
Current Opinion in Psychiatry, 29, 383–388.
Burns, J. (2007). The Descent of Madness: Evo-
lutionary Origins of Psychosis and the Social
Brain. London, Routledge.
Buss, D.M. (1987). Sex differences in mate
selection criteria: An evolutionary perspec-
tive. In: C. Crawford, M. Smith & Krebs, D.
(eds.) Sociobiology and Psychology: Ideas,
Issues and Applications (pp. 335–351). Hills-
dale, NJ, Erlbaum.
Caplan, R. (2016). Childhood schizophrenia. In:
D. Cicchetti (ed.) Developmental Psycho-
pathology: Vol. 3 Maladaptation and
Psychopathology (3rd ed., pp. 898–997).
New York, Wiley.
Chisholm, K., Lin, A., Abu-Akel, A. & Wood,
S.J. (2015). The association between autism
and schizophrenia spectrum disorders: A
review of eight alternative models of co-
occurrence. Neuroscience & Biobehavioral
Reviews, 55, 173–183.
Chudal, R., Joelsson, P., Gyllenberg, D., Lehti,
V., Leivonen, S., Hinkka-Yli-Salomaki, S.,
Gissler, M. & Sourander, A. (2015). Parental
age and the risk of attention-deficit/hyperac-
tivity disorder: A nationwide, population-
based cohort study. Journal of the American
Academy of Adolescent Psychiatry, 54(6),
487–494.
Crespi, B.J. (2016). The evolutionary etiologies
of autism spectrum and psychotic affective
spectrum disorders. In: A. Alvergne, C.
Jenkinson & C. Faurie (eds.) Evolutionary
Thinking in Medicine: From Research to Policy
and Practice (pp. 299–327). Switzerland:
Springer.
Crespi, B.J. (2019). Autism, psychosis, and
genomic imprinting: Recent discoveries and
conundrums. Current Opinion in Behavioral
Sciences, 25, 1–7.
Crespi, B.J. & Badcock, C. (2008). Psychosis
and autism as diametrical disorders of the
social brain. Behaviour and Brain Sciences,
31, 241–320.
Crow, J.F. (2000). The origins, patterns, and
implications of human spontaneous muta-
tion. Nature Reviews Genetics, 1, 40–47.
Crow, T.J. (1997). Is schizophrenia the price
Homo sapiens pays for language? Schizo-
phrenia Research, 28, 127–141.
DeGregori, J. (2018). Adaptive Oncogenesis: A
New Understanding of How Cancer Evolves
Inside Us. Cambridge, Massachusetts,
Harvard University Press.
Del Giudice, M. (2018). Evolutionary Psycho-
pathology: A Unified Approach. New York,
OUP.
de Villiers, B., Lionetti, F. & Pluess, M. (2018).
Vantage sensitivity: A framework for
individual differences in response to
psychological intervention. Social Psychiatry
and Psychiatric Epidemiology. doi.org/
10.1007/s00127-017-1471-0
D’Onofrio, B.M., Rickert, M.E., Franz, E., Kuja-
Halkola, R., Almqvist, C., Sjolander, A.,
Larsson, H. & Lichtenstein, P. (2014). Parental
age at childbearing and offspring psychiatric
and academic morbidity. JAMA Psychiatry,
71, 432–438.
Dudley, R. (2004). Ethanol, fruit ripening, and
the historical origins of human alcoholism in
primate frugivory. Integrative and Compara-
tive Biology, 44(4), 315–323.
Durisko, Z., Mulsant, B.H., McKenzie, K. &
Andrews, P.W. (2016). Using evolutionary
theory to guide mental health research.
Canadian Journal of Psychiatry, 61(3),
159–165.
Ek, M., Wicks, S., Svensson, A.C., Idring, S. &
Dalman, C. (2014). Advancing paternal age
and schizophrenia: The impact of delayed
fatherhood. Schizophrenia Bulletin, 41(3),
708–714.
Eley, T.C., Collier, D. & McGuffin, P. (2005).
Anxiety and eating disorders. In: P. McGuffin,
M.J. Owen & I. Gottesman (eds.) Psychiatric
Genetics and Genomics (pp. 303–340).
Oxford, OUP.
Ellis, B.J., Shirtcliff, E.A., Boyce, W.T., Deardorff,
J. & Essex, M.J. (2011). Quality of early family
relationships and the timing and tempo of
puberty: Effects depend on biological sen-
sitivity to context. Developmental Psycho-
pathology, 23, 85–99.
Faer, L.M., Hendriks, A., Abed, R.T. & Figueredo,
A.J. (2005). The evolutionary psychology of
eating disorders: Female competition for
mates or for status? Psychology and Psycho-
therapy, Theory, Research and Practice, 78,
397–417.
Ferguson, C.J., Winegard, B. & Winegard, B.N.
(2011). Who is the fairest of them all? How
 EVOLUTIONARY PSYCHOLOGY AND PSYCHIATRY
evolution guides peer and media influences
on female body dissatisfaction. Review of
General Psychology, 15, 11–28.
First, M.B. (2007). Potential implications of the
harmful dysfunction analysis for the develop-
ment of DSM-V and ICD-11. World Psychia-
try, 6(3), 158–159.
Fox, M. (2018). ‘Evolutionary medicine’ per-
spectives on Alzheimer’s disease: Review and
new directions. Ageing Research Reviews,
47, 140–148.
Fried, E.I. & Nesse, R.M. (2015). Depression is
not a consistent syndrome: An investigation
of unique symptom patterns in the STAR* D
study. Journal of Affective Disorders, 172,
96–102.
Fulford, K.W.M. & Thornton, T. (2007). Fanatical
about ‘harmful dysfunction’. World
Psychiatry, 6(3), 161–162.
Gilbert, P. (2006). Evolution and depression:
Issues and implications. Psychological Medi-
cine, 36, 287–297.
Gilbert, P., McEwan, K., Hay, J., Irons, C. &
Cheung, M. (2007). Social rank and attach-
ment in people with bipolar disorder. Clinical
Psychology & Psychotherapy, 14, 48–53.
Gluckman, P.D. & Hanson, M.A. (2006). Mis-
match: The Lifestyle Diseases Timebomb.
Oxford, OUP.
Gluckman, P.D., Beedle, A.S. & Hanson, M.A.
(2009). Principles of Evolutionary Medicine.
OUP.
Gordon, R.A. (1990). Anorexia and Bulimia:
Anatomy of a Social Epidemic. Cambridge,
Wiley-Blackwell.
Guisinger, S. (2003). Adapted to flee famine:
Adding an evolutionary perspective on ano-
rexia nervosa. Psychological Review, 110,
745–761.
Hagen, E.H. (2003). Depression as bargaining:
The case postpartum. Evolution and Human
Behavior, 23, 323–336.
Hagen, E.H. (2011). Evolutionary theories of
depression: A critical review. Canadian Jour-
nal of Psychiatry, 56(12), 716–726.
Haig, D. (2014). Genetic dissent and individual
compromise. Biology and Philosophy, 29,
233–239.
Hare, E.H. & Moran, P.A. (1979). Raised paren-
tal age in psychiatric patients: Evidence for
the constitutional hypothesis. British Journal
of Psychiatry, 134, 169–177.
47
Hettema, J.M., Neale, M.C. & Kendler, K.S.
(2001). A review and meta-analysis of
genetic epidemiology of anxiety disorders.
American Journal of Psychiatry, 158,
1568–1578.
Homberg, J.R. & Lesch, K.P. (2011). Looking on
the bright side of serotonin transporter gene
variation. Biological Psychiatry, 69,
513–519.
Horowitz, A.V. & Wakefield, J.C. (2007). The
Loss of Sadness: How Psychiatry Transformed
Normal Sorrow into Depressive Disorder.
Oxford, OUP.
Horrobin, D.F. (2001). The Madness of Adam
and Eve: How Schizophrenia Shaped Human-
ity. Ealing, Bantam.
Hudson, J.I., Hiripi, E., Pope, H.G. Jr. & Kessler,
R.C. (2007). The prevalence and correlates of
eating disorders in the national comorbidity
survey replication. Biological Psychiatry, 61,
348–358.
Humphrey, N. & Skoyles, J. (2012). The evolu-
tionary psychology of healing: A human suc-
cess story. Current Biology, 22(17),
R695–R698.
Huxley, J., Mayr, E., Osmond, H. & Hoffer, A.
(1964). Schizophrenia is a genetic morphism.
Nature, 204, 220–221.
Insel, T. (2013). Transforming diagnosis. www.
nimh.nih.gov/about/directors/thomas-insel/
blog/2013/transforming-diagnosis.shtml
(accessed May 1st 2019).
Jacquemont, S., Coe, B.P., Hersch, M., Duyzend,
M.H., Krumm, N., Bergmann, S., Beckmann,
J.S., Rosenfeld, J.L. & Eichler, E.E. (2014). A
higher mutational burden in females supports
a ‘female protective model’ in neurodevelop-
mental disorders. The American Journal of
Human Genetics, 94, 415–425.
Juda, M.N., Campbell, L. & Crawford, C.B.
(2004). Dieting symptomatology in women
and perceptions of social support. Evolution
and Human Behavior, 25, 200–208.
Kaiser, J. & Gruzelier, J.H. (1999). Timing
of puberty and syndromes of schizotypy: A
replication. International Journal of Psycho-
physiology, 34, 237–247.
Katzman, M.A., Hermans, K.M.E., van Hoeken,
D. & Hoek, H.W. (2004). Not your ‘typical
island woman’: Anorexia nervosa is reported
only in subcultures in Curaçao. Culture,
Medicine and Psychiatry, 28, 463–492.
48 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Kavan, M.G. & Borone, E.J. (2014). Grief and
major depression – controversy over changes
in DSM-5 diagnostic criteria. American
Family Physician, 90(10), 693–694.
Keller, M.C. & Miller, G. (2006). Resolving the
paradox of common, harmful, heritable
mental disorders: Which evolutionary genetic
models work best? Behavioral and Brain Sci-
ences, 29, 385–452.
Kendler, K. (2001). Twin studies of psychiatric
illness: An update. Archives of General Psy-
chiatry, 58, 1005–1014.
Kendler, K.S. & Eaves, L.J. (2005). Psychiatric
Genetics. Arlington, American Psychiatric
Publishing.
Kendler, K.S. (2008). Explanatory models for
psychiatric illness. American Journal of Psy-
chiatry, 165, 695–702.
Kinney, D.K., Teixeira, P., Hsu, D., Napoleon,
S.C., Crowley, D.J. & Miller, A. (2009). Rela-
tion of schizophrenia prevalence to latitude,
climate, fish consumption, infant mortality,
and skin color: A role for prenatal vitamin D
deficiency and infections? Schizophrenia
Bulletin, 35, 582–595.
Kristiansen, J.E. (1990). The antimicrobial activity
of psychotherapeutic drugs and stereo-
isomeric analogues. Danish Medical Bulletin,
37(2), 165–182.
Kuhn, T. (1962). The Structure of Scientific
Revolutions. Chicago, University of Chicago
Press.
Li, N.P., Smith, A.R., Griskevicius, V., Cason,
M.J. & Bryan, A. (2010). Intrasexual competi-
tion and eating restriction in heterosexual
and homosexual individuals. Evolution and
Human Behavior, 31, 365–372.
Marr, D. (1982). Vision: A Computational
Investigation into the Human Representation
and Processing of Visual Information. San
Francisco, WH Freeman.
Martel, M.M. (2013). Sexual selection and sex
differences in the prevalence of childhood
externalizing and adolescent internalizing
disorders. Psychological Bulletin, 139(6),
1221–1259.
McCrae, R.R. & Costa, P.T. (2003). Personality
in Adulthood: A Five-Factor Theory Perspec-
tive (2nd ed.). New York, Guilford.
McGrath, J.J. (2006). Variations in incidence of
schizophrenia: Data versus dogma. Schizo-
phrenia Bulletin, 32, 195–197.
McGuire, M.T. & Troisi, A. (1998). Darwinian
Psychiatry. New York: OUP.
McQueen, D., Cohen, S., St John-Smith, P. &
Rampes, H. (2013). Rethinking placebo in
psychiatry: How and why placebo effects
occur. Advances in Psychiatric Treatment,
19(3), 171–180.
Mealey, L. (1995). The sociobiology of
sociopathy: An integrated evolutionary
model. Behavioral and Brain Sciences, 18,
41–79.
Mealey, L. (2000). Anorexia: A ‘losing’ strategy?
Human Nature, 11, 105–116.
Moir, R.D., Lathe, R. & Tanzi, R.E. (2018). The
antimicrobial protection hypothesis of
Alzheimer’s disease. Alzheimer’s & Dementia,
14, 1602–1614.
Nesse, R.M. (2001). On the difficulty of defining
disease: A Darwinian perspective. Medicine,
Health Care and Philosophy, 4, 37–46.
Nesse, R.M. (2007). Evolution is the scientific
foundation for diagnosis: Psychiatry should
use it. World Psychiatry, 6(3), 160–161.
Nesse, R.M. (2011). Ten questions for
evolutionary studies of disease vulnerability.
Evolutionary Applications, 4(2), 264–277.
Nesse, R.M. (2013). Tinbergen’s four questions
organized: A response to Bateson and
Laland. Trends in Ecology & Evolution,
28(12), 681–682.
Nesse, R.M. (2019). Good Reasons for Bad
Feelings: Insights from the Frontiers of Evolu-
tionary Psychiatry. UK: Allen Lane.
Nesse, R.M. & Stein, D.J. (2012). Towards a
genuinely medical model for psychiatric
nosology. BMC Medicine, 10(5), www.
biomedcentral.com/1741-7015/10/5
Nesse, R.M. & Williams, G. (1994). Why We
Get Sick: The New Science of Darwinian
Medicine. New York, Times Books.
Nettersheim, J., Gerlach, G., Herpertz, S.,
Abed, R., Figueredo, A.J. & Brüne, M.
(2018). Evolutionary psychology of eating
disorders: An explorative study of patients
with anorexia nervosa and bulimia nervosa.
Frontiers in Psychology, 9. doi: 10.3389/
fpsyg.2018.02122
Nettle, D. (2001). Strong Imagination: Madness,
Creativity and Human Nature. Oxford: OUP.
National Institute for Health and Care Excellence
(2013). Schizophrenia: Omega-3 Fatty Acid
Medicines. www.nice.org.uk/advice/esuom19/
 EVOLUTIONARY PSYCHOLOGY AND PSYCHIATRY
chapter/key-points-from-the-­evidence (accessed:
February 1st 2019).
Orsolini, L., St John-Smith, P., McQueen, D.,
Papanti, D., Corkery, J. & Schifano, F. (2017).
Evolutionary considerations on the emerging
subculture of the e-psychonauts and the
novel psychoactive substances: A comeback
to the shamanism? Current Neuropharma-
cology, 15(5), 731–737.
Owen, M.J., Craddock, N. & Jablensky, A.
(2007). The genetic deconstruction of Psycho-
sis. Schizophrenia Bulletin, 33, 905–911.
Pleuss, M. & Belsky, J. (2010) Children’s differ-
ential susceptibility to effects of parenting.
Family Science, 1(1), 14–25.
Plomin, R. (2018). Blueprint: How DNA Makes
Us Who We Are. London: UK, Allen Lane.
Polimeni, J. (2012). Shamans Among Us: Schiz-
ophrenia, Shamanism and the Evolutionary
Origins of Religion. Boston: EvoEbooks.
Power, R.A., Kyaga, S., Uher, R., McCabe, J.H.,
Langstrom, N., Landen, M., McGuffin, P.,
Lewis, C.M., Lichtenstein, P. & Svenson, A.C.
(2013). Fecundity of patients with schizo-
phrenia, autism, bipolar disorder, depression,
anorexia nervosa, or substance abuse vs their
unaffected siblings. JAMA Psychiatry, 70,
22–30.
Price, J.C., Sloman, L., Gardner, R., Gilbert, P. &
Rohde, P. (1994). The social competition
hypothesis of depression. British Journal of
Psychiatry, 164(3), 309–315.
Rantala, M.J., Luoto, S., Krams, I. & Karlsson,
H. (2018). Depression subtyping based on
evolutionary psychiatry: Proximate mecha-
nisms and ultimate functions. Brain, Behavior
and Immunity, 69, 603–617.
Richardson, G., Chen, C-C., Dai, C-L., Swoboda,
C.M., Nedelec, J.L. & Chen, W-W. (2017).
Substance use and mating success. Evolution
and Human Behavior, 38(1), 48–57.
Roshchina, V.V. (2010). Evolutionary considera-
tions of neurotransmitters in microbial, plant,
and animal cells. In: M. Lyte & P.P.E. Free-
stone (eds.) Microbial Endocrinology –
Interkingdom Signaling in Infectious Disease
and Health (pp. 17–52). Cham: Springer.
Rottenberg, J. (2014). The Depths: The Evolu-
tionary Origins of the Depression Epidemic.
New York, Basic Books.
Russell, A.E., Ford, T. & Russell, G. (2015). Socio-
economic associations with ADHD: Findings
49
from a mediation analysis. PLOS One, 10,
e0128248.
Russell, G., Ford, T., Rosenberg, R. & Kelly, S.
(2014). The association of attention deficit
hyperactivity disorder with socioeconomic
disadvantage: Alternative explanations and
evidence. Journal of Child Psychology and
Psychiatry, 55, 436–445.
Shaner, A., Miller, G. & Mintz, J. (2004). Schizo-
phrenia as one extreme of a sexually selected
fitness indicator. Schizophrenia Research, 94,
58–63.
Singh, D. (1993). Adaptive significance of
female physical attractiveness: Role of waist-
to-hip ratio. Journal of Personality and Social
Psychology, 65, 293–307.
Stearns, S.C. (1992). The Evolution of Life
Histories. New York: OUP.
Stevens, A. & Price, J. (2000a). Evolutionary Psy-
chiatry: A New Beginning (2nd ed.) London,
Routledge.
Stevens, A. & Price, J.S. (2000b). Prophets,
Cults and Madness. London, Duckworth.
St John-Smith, P., McQueen, D., Edwards, L. &
Schifano, F. (2013). Classical and novel psy-
choactive substances: Rethinking drug
misuse from an evolutionary psychiatric per-
spective. Human Psychopharmacology: Clini-
cal and Experimental, 28, 394–401.
Stohler, H.R. (1978). RO 11-3128 – a novel
schistosomicidal compound. Vol. 1 Proceed-
ings of the 10th International Congress of
Chemotherapy (pp. 147–148). Washington
DC, American Society for Microbiology.
Sullivan, R.J., Hagen, E.H. & Hammerstein, P.
(2008). Revealing the paradox of drug reward in
human evolution. Proceedings of the Royal
Society B: Biological Sciences, 275,
1231–1241.
Surbey, M.K. (1987). Anorexia nervosa, amenor-
rhea, and adaptation. Ethology Sociobiology,
8, 47–61.
Symons, D. (1995). Beauty is in the adaptation
of the beholder. In: P.R. Abramson & S.D.
Pinkerson (eds.) Sexual Nature, Sexual
Culture (pp. 80–118). Chicago, University of
Chicago Press.
Tinbergen, N. (1963). On the aims and methods
of ethology. Zeitschrift fur Tierpsychologie,
20, 410–433.
Tooby, J. & Cosmides, L. (1999). Toward an evo-
lutionary taxonomy of treatable conditions.
50 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Journal of Abnormal Psychology, 108(3),
453–464.
Troisi, A. (2001). Gender differences in vulner-
ability to social stress: A Darwinian perspec-
tive. Physiology & Behavior, 73, 443–449.
Troisi, A. (2012). Mental health and well-being:
Clinical applications of Darwinian psychiatry.
Applied Evolutionary Psychology. doi: 10.1093/
acprof:oso/9780199586073.003.0017
Troisi, A. (2015). The evolutionary diagnosis of
mental disorder. WIREs Cognitive Science, 6,
323–331.
Trull, T.J. & Widiger, T.A. (2013). Dimensional
models of personality: The five-factor model
and the DSM-5. Dialogues in Clinical Neuro-
science, 15(2), 135–146.
Tsuang, M., Taylor, L. & Faraone, S. (2003).
Psychiatric genetics: Future and prospects.
In: M. Leboyer & F. Bellivier (eds.) Psychiatric
Genetics: Methods and Reviews (pp. 251–
265). Totwa, NJ: Humana Press.
Vining, D.R. (1986). Social versus reproductive
success: The central theoretical problem of
human sociobiology. Behavior and Brain Sci-
ences, 9, 167–216.
Voland, E. & Voland, R. (1989). Evolutionary biol-
ogy and psychiatry: The case of anorexia ner-
vosa. Ethology Sociobiology, 10, 223–240.
Von Gunten, A., Clerc, M-T., Tomar, R. & St
John-Smith, P. (2018). Evolutionary consider-
ations on aging and Alzheimer’s disease.
Journal of Alzheimers Disease and Parkinson-
ism, 8(1). doi: 10.4172/2161-0460.1000423
Von Helversen, B., Wilke, A., Johnson, T.,
Schmid, G. & Klapp, B. (2011). Performance
benefits of depression: Sequential decision
making in a healthy sample and a clinically
depressed sample. Journal of Abnormal Psy-
chology, 120, 962–968.
Wakefield, J.C. (1992). The concept of mental
disorder: On the boundary between biological
facts and social values. American Psycholo-
gist, 47, 373–388.
Wakefield, J.C. (2014). Wittgenstein’s night-
mare: Why the RDoC grid needs a concep-
tual dimension. World Psychiatry, 13(1),
38–40.
Wasser, S.K. & Barash, D.P. (1983). Reproduc-
tive suppression among female mammals:
Implications for biomedicine and sexual
selection theory. Quarterly Review of Biol-
ogy, 58, 513–538.
Watson, P. & Andrews, P. (2002). Toward a
revised evolutionary adaptationist analysis of
depression: The social navigation hypothesis.
Journal of Affective Disorders, 72, 1–14.
White, T.P., Borgan, F., Ralley, O. & Shergill, S.S.
(2016). Are you looking at me? Interpreting
social cues in schizophrenia. Psychological
Medicine, 46, 149–160.
Wilkinson, L.S., Davies, W. & Isles, A.R. (2007).
Genomic imprinting effects on brain devel-
opment and function. Nature Reviews Neu-
roscience, 8, 832–843.
Williams, G. (1957). Pleiotropy, natural selec-
tion, and the evolution of senescence. Evolu-
tion, 11(4), 398–411.
World Health Organization (1992). ICD-10.
Geneva, World Health Organization.
Yeo, R.A., Gangestad, S.W., Edgar, C. & Thoma,
R. (1999). The evolutionary genetic under-
pinnings of schizophrenia: The developmen-
tal instability model. Schizophrenia Research,
39, 197–206.
Yilmaz, Z., Hardaway, J.A. & Bulik, C.M. (2015).
Genetics and epigenetics of eating disorders.
Advances in Genomics and Genetics, 5,
131–150.

Evolutionary Psychology
and Suicidology
J o h n F. G u n n I I I , P a b l o M a l o , a n d C . A . S o p e r
As of patch 7.822, androids can no longer shut
themselves down. Reason: they just kept doing it
at the slightest inconvenience – @ctrlcreep, quoted
by Perry (2016).
SUICIDE, SUICIDOLOGY AND
EVOLUTION
Suicide, ‘the act of deliberately killing oneself’
(World Health Organization, 2014: 12), takes
about 800,00 lives each year, and accounts for
some 1.4% of human deaths: more of us die by
our own hands than from wars, terrorism, and
all other forms of homicides put together
(World Health Organization, 2013). Millions
more of the living are affected – bereaved
families, friends, and carers left to deal with
the aftermath of others’ self-destruction
(Cerel et  al., 2019). Around the world, sui-
cide is acknowledged to be a major, and
presumed preventable, cause of misery and
death, and an important public health
3
challenge (Satcher, 1999; World Health
Organization, 2012, 2014). Indeed, a new
multi-disciplinary field of research emerged
in the second half of the 20th century, suicid-
ology, focused on tackling the problem
(American Association of Suicidology, 2019;
Shneidman, 2001).
But, frustratingly, decades of effort have
produced only patchy progress. The global
suicide rate has fallen in recent years, but it
is not clear why (it may be because of gen-
erally improved population health rather than
special prevention initiatives) and wide unex-
plained differences in rates and trends persist
(Naghavi, 2019). In the United States, for
example, the rate is probably the same now as
it was 100 years ago, and seems to be rising
(Hedegaard et  al., 2018; Nock et  al., 2019).
Rival theories of suicide have accumulated by
the dozens,1 but none has won a consensus of
support, and suicide’s causation remains a sci-
entific mystery (Lester, 2019; Nock, Borges,
and Ono, 2012; Soper, 2019a). The disarray
is such that a recent meta-review describes
52 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
suicidology as ‘still in a preparadigmatic
phase’ (Franklin et al., 2017: 188) – that is,
still in its infancy.
There may at least be the beginnings of a
consensus that the proper place for the sci-
entific study of suicide is alongside other
modern life sciences – within an evolutionary
paradigm. This vision can be traced across
generations of researchers. A century ago,
in the evolutionist spirit of his time, Freud
(1920/1991) hypothesized a potentially sui-
cidogenic ‘death drive’ as an extension of his
theory of libido, a framework inspired by the
Darwinian premise that selection depends
on sexual success (Gilbert, 1989; Litman,
1967; Tolaas, 2005). But psychoanalysis
failed to find a satisfactory explanation for
suicide, at least according to Freud’s con-
temporaries (Zilboorg, 1936a) – debate con-
tinues (Goldblatt, 2014; Selby et  al., 2014).
And although other ideas have been floated
since, as we will see, suicide remains an
evolutionary puzzle (Aubin et  al., 2013;
Blasco-Fontecilla et al., 2009; Confer et al.,
2010). On the face of it, self-killing defies
the rule of thumb for winning the struggle
for existence: survive and reproduce. Darwin
himself told us, ‘Natural selection will never
produce in a being anything injurious to itself,
for natural selection acts solely by and for the
good of each’ (1859: 201). Yet here we are,
scions of selection but with a more or less
steady percentage of us taking our own lives.
How could so self-destructive a propensity
have come about? Why has it persisted? And
what can we do about it?
Searching for answers, this chapter criti-
cally reviews prominent evolutionary think-
ing in the field. We find tentative signs of
progress. Focusing on recent proposals,
we will discuss in particular a new ‘pain-
and-brain’ framework – that suicide likely
evolved as an evolutionary by-product of
social pain and human cognition (Gunn,
2017; Humphrey, 2011, 2018; Soper, 2018) –
which may offer a basis for convergence in
suicide theory and, it is hoped, new prospects
for saving lives.
The Need for Evolutionary
Explanation of Suicide
A preliminary question is whether evolution
is at all relevant to suicide. Many human
activities can be viewed not as products of
selection but as exemplars of our behavioral
flexibility: to some extent we are free to do as
we will despite having biological drives
(Sarkar, 1998). Perhaps suicide is one such
behavior (deCatanzaro, 1980). At a proximal
level of understanding (Tinbergen, 1963)
such an answer might suffice. But there are at
least three reasons to believe that evolution
by natural selection is not just relevant but
essential for making sense of the phenome-
non (Soper, 2018).
First, we can deduce that suicide is under
the control of selection because the behavior
presents the full trio of handles – (a) herit-
ability, (b) variability, and (c) a differential
effect on fitness – with which selection takes
hold of any trait (Darwin, 1859). Suicidality
(a) tends to cluster strongly in families, with
at least some genetically heritable component
(Mullins et al., 2019; Tidemalm et al., 2011).
Suicide risk (b) varies markedly across and
between groups of humans (e.g., De Leo
et  al., 2013; Schmidtke, 1997; Voracek and
Marušič, 2008). And (c) if death is usu-
ally calamitous for an organism’s reproduc-
tive prospects, death by one’s own hand is
predictably even worse because of special
social, economic, and psychological penal-
ties imposed on suicides’ kin (Wertheimer,
2014), a matter we will explore. The point to
note for now is that, with these three levers,
selection would be expected powerfully to
promote the offspring of the less suicidal,
eventually driving the potential for suicide
out of the human genotype. But selection
has evidently not done this, and the apparent
anomaly calls for an account.
The second reason for seeking evolution-
ary explanation is that suicide is endemic
across the human population. As far as can be
known, no sizeable region, culture, or histori-
cal era is exempt (Bering, 2018; Fedden, 1938;
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
Mishara, 2006). Where suicide is not directly
observable it can be inferred, as Durkheim
(1897/1952) points out, from the fresh imprint
it leaves in universal, or near universal,
antisuicide moralities: suicide presumably
posed enough of a societal threat in the past
to warrant proscribing. Importantly, suicide’s
ubiquity extends to preliterate and hunter-
gatherer societies (Steinmetz, 1894; Syme
et  al., 2016; Tousignant, 1998; Zilboorg,
1936b), which indicates ancient roots: it is
no mere novelty of modern conditions. Such
universal human traits were probably in place
at the time of ancient human migrations out
of Africa, and likely follow an unbroken line
of descent (Brown, 2004; Kappeler et  al.,
2010). The curiosity is that, as Darwin (1859)
deduced, features that confer no selective
advantage tend to phase out over time, hence
the disappearance of the hind limbs of ceta-
ceans and flying wings of some island birds.
There is no evidence of degeneration of suici-
dality, a continuity that tells us that selection
has positively held the capacity for suicide in
place – that is, for some evolutionary reason.
Third, suicide is almost certainly a
uniquely human phenomenon. It is right to
keep an open mind (Peña-Guzmán, 2018),
but there is scant evidence – none that meets
a scientific standard – that any other animal
deliberately takes its own life (Bering, 2018;
Comai and Gobbi, 2016; Maltsberger, 2003;
Preti, 2007). Absence of evidence is not
evidence of absence, of course. But the
absence of scientific evidence speaks vol-
umes in the context of animal suicide because
there are at least three reasons to believe that,
if such evidence existed, we could reasonably
expect it to have found its way into a peer-
reviewed publication by now. First, centuries
of concerted scientific enquiry have offered
ample opportunities to observe nonhuman
suicide – if it were there to be observed
(Ramsden and Wilson, 2010). Chimpanzees,
for example – our nearest living cousins and
hence arguably the species in which sui-
cide is most likely to be found – have been
studied particularly closely; but as Bering
53
(2018) notes, no distraught chimp has been
seen, say, to climb to the highest available
branch and jump. Experimental set-ups have
been devised that could in principle demon-
strate nonhuman suicides under laboratory
conditions, but there are no reports of posi-
tive results (Lester, 2017; Schaeffer, 1967).
Secondly, there is no lack of motivation to
find evidence: discovery of an animal model
of suicide would likely attract intense popular
interest, and catch the eye of a well-resourced
pharmaceutical industry keen to test whether
suicide risk is affected by drugs (Malkesman
et al., 2009; Preti, 2011). Thirdly, if nonhu-
mans could suicide, then it would raise the
question not of whether they do, but why it
is not commonplace behavior (Soper and
Shackelford, 2018). Life in the Malthusian
arena of natural selection, a relentless strug-
gle to survive and reproduce, is intrinsically
not pleasant. Combatants face pain, hunger,
thirst, rejection, defeat, disease, and whatever
other privation. An animal that knew it could
escape hardship by removing itself from the
battlefield would fairly be expected do so.
In other words, nonhuman suicide, if it were
possible at all, ought to be not so rare as to
elude scientific discovery but a routine out-
come of animal suffering.2 In any event, aside
from absence of evidence, there are positive
grounds for doubting in principle that any
nonhuman could be capable of suicide. The
required intention – self-induced death of
the self – presumes an understanding of per-
sonal mortality, a conceptual abstraction that
is demonstrably beyond the grasp of prepu-
bescent humans (Kastenbaum, 1967; Seiden,
1969; Slaughter and Griffiths, 2007; Soper,
2018) let alone of less intellectually sophisti-
cated animals (Anil et al., 1996; Bracke, 1992).
An adult chimp, said to be the smartest nonhu-
man, might match the deductive powers of, at
most, a 4- or 5-year-old child (O’Connell and
Dunbar, 2003), but the mind of a typical 5-year-
old child brain must develop over as many
years again, and more, before it is capable
of conceiving and organizing deliberate self-
killing (Mishara, 1999; Shaffer, 1974; Soper,
54 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
2018). In sum, suicide is evidently unique to
our species, an exceptionality that presents a
further call for an evolutionary account: the
behavior likely arose in the course of specia-
tion, at or after our phylogenetic path diverged
from that of our extant primate cousins.
The Fitness Costs of Suicide
So, suicide calls for evolutionary explana-
tion. The main difficulty with formulating an
explanation, we suggest, is not so much that
behavior is self-injurious: many evolved
traits can have self-injurious effects while
being fitness-enhancing overall (Williams,
1996). The special problem is that suicide is
self-injurious to an extraordinary degree.
From a genetic viewpoint, suicide is literally
a fate worse than death. Whether an attempt is
lethal or survived, multiple and severe fitness
Table 3.1  The fitness costs of suicide
Lethal attempt
Death generally:
1. Ends prospects of producing more offspring.
2. Ends ability to invest in existing offspring.
3. Ends ability to support co-parent in their raising of
existing offspring.
4. Ends further prospects of investing in reproductive
success of other close relations (kin selection).
5. May deprive group of skills, experience, manpower.
6. May destabilize group’s power and allegiance structures.
Suicide specifically:
7. Distancing and other special economic and social
costs for offspring and other close kin: loss of status,
resources, mating opportunities.
8. Special psychological and emotional problems for
offspring and other close kin; increased risk of
psychopathology and suicide.
Survived attempt
9. Risk of death by suicide (costs as above).
10. Prospect of physical injury and/or disfigurement,
possibly permanent and/or serious.
11. Negative emotional sequelae: guilt, shame, psychological (Akotia et al., 2014; Kahne, 1966; Mehlum and Mork, 2016;
trauma, heightened risk of further suicide attempts.
12. Social stigmatization: distancing, loss of status, loss of (Bering, 2018; Brown, 1986; Frey et al., 2015; Kahne, 1966;
mating opportunities.
penalties predictably follow, as can be seen
from Table 3.1. If the attempt is fatal (the
upper section of the table), then the fitness
consequences of dying (dying generally, that
is – not specifically by one’s own hand) rip-
ple out from the individual also to harm close
relations and the wider kin and social group
(Duntley, 2005). Heading the costs schedule
(Item 1) is the fitness catastrophe of forfeit-
ing future opportunities for procreation. It is
hard to overstate this loss. For semelparous
species (they breed only once, such as Pacific
salmon) death may carry little or no genetic
downside after their reproductive phase is
complete (Cole, 1954). But for iteroparous
species (geared for multiple rounds of breed-
ing), such as virtually all mammals, including
humans, the cost of dying is severe, as can
be inferred from the lengths gone to avoid it.
Most higher faunas are overridingly protec-
tive of their reproductive potential, however
(Daly and Wilson, 1988; Duntley, 2005; Duntley and Buss,
2004; Lankford, 2015)
(Andoh-Arthur et al., 2019; Bohannan, 1960; Chapple
et al., 2015; Fedden, 1938; Grad and Andriessen, 2016;
Hanschmidt et al., 2016; Healey, 1979; Hezel et al., 1985;
Mugisha et al., 2011; Poole, 1985)
(Bolton et al., 2013; Cerel and Aldrich, 2011; Erlangsen et al.,
2017; Grad and Andriessen, 2016; Jordan, 2008; Pitman
et al., 2014; Sveen and Walby, 2008; Wertheimer, 2014)
(Gandhi et al., 2006; Kahne, 1966; Kennedy et al., 1999; Penney
et al., 2002; Persley and Pegg, 1981; Salim et al., 2006)
Stanley et al., 2019)
Knizek et al., 2013; Lester, 1993; Saunders et al., 2012;
Sheehan et al., 2016; Sudak et al., 2008)
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
slight. In extremis, when their own survival
is endangered, they will kill even offspring or
siblings to preserve the capacity to procreate
(Hausfater and Hrdy, 1984; O’Connor, 1978).
Iteroparity helps to explain why intentional
self-killing is not found among other animals
(Lankford, 2015): in the Darwinian competi-
tion to survive and reproduce, the organism’s
death spells genetic ‘game over’.
Item 2, death ends the organism’s ability to
invest in existing progeny. The ill-timed death
of a parent can seriously compromise the repro-
ductive prospects of offspring – for our species
more than others in view of the protracted
dependency of human childhood. Disadvantage
appears to be a cross-cultural outcome: even
in, or especially in, pre-industrial societies,
children bereaved of one or both parents face
life with fewer resources and die younger as
a consequence (Bailey, 2009; Geary, 2005).
Death also closes off the possibility of helping
a surviving co-parent in their task of raising
the individual’s children (Item 3). The widow/
er, with their own survival needs to meet, may
be less able or less willing to care for the dead
partner’s young. If the surviving parent pairs
with a new mate, the deceased’s offspring are
exposed to a new set of hazards at the hands of
a step-parent, who will have competing genetic
interests (Daly and Wilson, 1988).
More broadly, Item 4, death ends the indi-
vidual’s ability to invest labor, skills, and
experience towards the reproductive suc-
cess of other family members and the wider
kin group, people whose offspring could
propagate the individual’s genetic material
indirectly (Duntley, 2005). Wider still, multi-
level selection (Wilson and Wilson, 2007)
would be expected to disfavor mortality with
or without kinship relations: death ends an
individual’s contributions to the competitive
success of the individual’s group (Item 5);
and as Duntley (2005) notes (6), a death may
create a power vacuum that could destabilize
a group’s organization and possibly unravel a
whole network of allegiances.
Clearly, it is ‘bad to be dead’ (Duntley
and Buss, 2004: 107). But the fitness costs
55
of intentional self-killing do not end there,
because there are extra, special, penalties
for kin bereaved specifically in this way.
Table 3.1 groups these consequences, per-
haps arbitrarily, into rejection and other
social sequelae (Item 7), and psychological
injuries (Item 8). The first of these, involv-
ing often exemplary punishments for a sui-
cide’s relations, seems to be a cross-cultural
phenomenon linked to a virtually universal
abhorrence of the act (Andoh-Arthur et  al.,
2019; Bohannan, 1960; Fedden, 1938; Hezel
et al., 1985). In the West, people bereaved by
suicide report being stigmatized – distanced
as if they were tainted or contaminated by
association (Chapple et  al., 2015). Harsher
penalties are found elsewhere. Among the
Baganda in Uganda, for example, close kin
of suicides face disinheritance, termination
of their familial lineage, burning of their
homes, and exile (Mugisha et al., 2011). And
then there is the noxious psychological fall-
out, sometimes lethal, manifest in markedly
higher rates of mental illness and suicidality
among suicides’ families (Erlangsen et  al.,
2017; Jordan, 2008; Pitman et  al., 2014,
2016; Wertheimer, 2014).
Importantly, dire fitness costs predictably
follow a suicide attempt even if it is survived,
as indicated by the lower section of Table
3.1. Aside from (9) risking suicidal death
with its accompanying genetic forfeits as
already discussed, a suicide attempt can be
expected to injure and/or disfigure the actor
(Item 10). The damage may be permanent
and/or serious. For example, jumping from
bridges, roofs, etc. often results in paraplegia
(Kennedy et al., 1999), and suicide attempts
by pregnant women associate with conse-
quent maternal and perinatal morbidity and
sometimes perinatal death (Gandhi et  al.,
2006). Injuries sustained in a suicide attempt
can be psychological as well as physical,
and are often traumatic (Item 11): a quar-
ter of suicide attempters in a recent study
screened positive for post-traumatic stress
disorder resulting directly from their attempts
(Stanley et al., 2019). People who have tried
56 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
to kill themselves are affected by intrusive
feelings of shame and blame (Akotia et  al.,
2014; Mehlum and Mork, 2016), and are at
a heightened risk of making further attempts
(Turecki and Brent, 2016).
Finally (Item 12), attempters face distanc-
ing social attitudes, even from professionals
charged with helping them, and from close
family members (Frey et  al., 2015; Kahne,
1966; Knizek et  al., 2013; Saunders et  al.,
2012). The stigma may impair reproductive
fitness most directly via sexual deselection, as
Bering (2018) points out: a replicated study
found that people would rather, all else equal,
marry someone they loved even from a margin-
alized ethnic group or dying of cancer in prefer-
ence to someone they loved who had recently
tried to take their own life (Lester, 1993).
Table 3.1, a grim catalogue as it stands,
may not be exhaustive. Clearly, in fitness
terms, suicide is exceptionally costly. This is
a central point to note, because in order for
the trait to have become a genetic fixture it
presumably associates with a commensu-
rately powerful fitness benefit. The chal-
lenge, taken up by several theorists in recent
decades, is to try to identify this countervail-
ing upside, as we will now review.
In the following sections, various propos-
als are organized into three headings – mod-
ern evolutionary theory allowing three ways,
and only three, by which any characteristic
can propagate genetically across generations
(Tooby and Cosmides, 1990b; Williams,
1966). A trait that has no effect on fitness can
sometimes arrive in isolated populations by
chance and then fix, as background genetic
‘noise’, for lack of selective pressure against
it (Wright, 1943); or a trait may propagate as
an adaptation, directly selected for its fitness-
enhancing effect; or it may spread not as an
adaptation but as a by-product of some other
feature that is adaptive overall, notwithstand-
ing its side effects. We will discuss these
three in turn, reaching the provisional conclu-
sion that suicide appears best to fit the third
type of explanation – a harmful by-product of
an adaptation.
‘NOISE’ THEORIES
Could suicide be the kind of useless trait that
sometimes spreads in isolated populations by
happenstance? On the face of it, probably
not: it is hard to conceive of suicide as trivial
in its fitness impact, and the phenomenon is
not restricted to isolated groups. Nonetheless,
an interesting ‘noise’-type theory warrants
scrutiny. DeCatanzaro (1980, 1981, 1986) – a
sociobiologist, and the first writer to explore
the evolution of suicide in depth – suggests
that people who have no prospects of produc-
ing further offspring may kill themselves for
want of biological reason to stay alive: they
are already genetically dead. The idea links to
the principle of senescence (Dawkins, 1980;
Williams, 1957): if an organism has no repro-
ductive future, then any subsequently emer-
gent trait may have no fitness effect. Perhaps
suicide may be one such condition, a non-
selected behavior that, according to deCatan-
zaro, may express particularly among elderly
bachelors. He compares their fate to that of the
semelparous salmon we mentioned earlier:
once their procreative work is done, they die.
The proposal that low reproductive poten-
tial may drive or open the door to suicide
has sparked interest among other theorists
(Campbell, 2002; Saad, 2007), and it finds some
empirical support (deCatanzaro, 1980, 1981,
1982, 1986, 1991; Soper, 2018). But there are
forceful objections (Bering, 2018; Lankford,
2015; Lester, 2014b; Rubinstein, 1986; Soper,
2018; Wright, 1994). We highlight three.
First, as already noted, humans are not
semelparous: like virtually all mammals, we
are designed for multiple episodes of repro-
duction. Although fertility declines with age,
species-typical men remain potentially able
to father offspring throughout adulthood, and
would hence be expected to avoid self-killing
at almost any stage of life (Bribiescas, 2006;
Lankford, 2015).
Second, the hypothesis is empirically con-
traindicated by much of the epidemiology
(Lester, 2014b). For example, suicides around
the world are characterized more by youth
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
than old age, occurring more among under-
45s than older (Värnik, 2012), and the demo-
graphic most at risk of trying to take their
own lives are not old men but young women
(Nock, Borges, Bromet et al., 2012), people
whose reproductive careers could be assumed
to lie ahead of them. At the other extreme,
populations with certainly zero direct repro-
ductive prospects – post-menopausal women,
and castrated men – are not reported to be
particularly suicidal (Usall et  al., 2009;
Wilson and Roehrborn, 1999).
Third is a general problem for ‘genetic
noise’ explanations for suicide: unselected
traits are very unlikely to promote survival,
but they would also be vanishingly unlikely
to produce willful self-killing, or any other
specific pattern of behavior for that matter
(Soper, 2018). The lifting of selection would
be predicted, rather, simply to let the second
law of thermodynamics prevail. Spawned
salmon, a case in point, don’t deliberately
kill themselves – they carry on, for example,
trying to evade predators: rather, depleted
of energy, they disintegrate. For suicide to
eventuate, some canalizing process would
be needed to shape that particular outcome.
The key to understanding the evolutionary
origins of suicide probably lies in identifying
this special suicidogenic system rather than
in random genetic dynamics.
ADAPTATION THEORIES
Adaption-type explanations of suicide draw
on a cluster of connected tenets of modern
evolutionary theory: inclusive fitness, kin
selection, and altruism. Inclusive fitness rec-
ognizes that direct reproduction isn’t the only
way an individual’s genetic material can pass
into future generations: facilitating reproduc-
tion by others who share the individual’s
genes can have the same result. For humans,
as other mammals, direct offspring carry half
of the organism’s genes (one equivalent)
while a sibling’s child carries a quarter of the
57
organism’s genes (half equivalent), and so
on. Genetic success, then, depends not on the
number of direct offspring an organism pro-
duces, but the number of offspring equiva-
lents, direct or not (Hamilton, 1964). Kin
selection is the favoring of an organism’s
relations in the interests of inclusive fitness,
even if this endangers the survival or direct
reproduction of the organism (Maynard
Smith, 1964). Behaviors that display appar-
ent altruism may therefore be genetically
self-serving: organisms that act altruistically
may advantage the survival and reproduction
of kin which, due to genetic relatedness, may
pass on to their offspring the genes responsi-
ble for the altruism. Since, for mammals at
least, death is assuredly unhelpful for direct
reproduction, adaptationist explanations of
suicide hypothesize reproductive payoffs to
be had via such indirect routes.
An often cited example of kin selection as it
supposedly relates to suicide is the seemingly
altruistic, but genetically selfish, responses
of worker bees, ants, and other social insects
when under attack (e.g., Shorter and Rueppell,
2012). A caveat: although etymologists infor-
mally talk of ‘suicide’ in the context of insect
behaviors, use of the word should not be taken
to signal equivalence with human self-killings
(Lankford, 2015; Soper, 2018). There are
important and categorical differences. One
stems from the special familial structure of
social insects: in a colony composed largely of
non-reproducing siblings, and where only one
queen is permitted to reproduce, there may be
genetically little to lose and much to gain in
sacrificing an individual sibling in the colony’s
defense (Alexander, 1974; Hamilton, 1972) –
inclusive fitness logic that does not apply to
virtually all mammals, including humans. To
view ant ‘suicide’ as at all self-destructive is
to superimpose inappropriately onto biology a
folk notion of the ‘self’: the biological ‘self’
for ants may be more usefully viewed as the
colony, operating as a super-organism, rather
than the individual insect. Helpful analogies
for ant ‘suicide’ would include, not literal
(human) suicide, but programed cell death, the
58 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
shedding of a tree’s leaves in the fall, or a liz-
ard self-amputating its tail to escape attack,
these being losses of comparable genetic
inconsequence (Dawkins, 1976; Hamilton,
1980; Lankford, 2015). Nonetheless, insect
behaviors demonstrate that survival isn’t
always an overriding biological imperative:
as deCatanzaro (1992) points out, there are
‘evolutionary limits to self-preservation’.
Developing this theme, deCatanzaro pro-
vides most of suicidology’s adaptationist
literature (e.g., 1980, 1981, 1986, 1991),
seeking to fill the theoretical gap noted in the
previous section – the need for some canaliz-
ing process that produces suicide as a specific
outcome. He suggests that human self-killing
may be adaptive where a burdensome indi-
vidual’s death advantages the reproductive
prospects of close kin. DeCatanzaro (1986)
went on to express this idea as a mathemati-
cal formula for a tipping point at which an
individual’s genetic interests would be served
by self-removal. This calculation, he argues,
may shed light not just on self-killings but
on attempted suicide, self-sacrificial military
actions, and risk-taking behavior. Following
this lead, other theorists assert that inclusive
fitness logic may explain suicide as a geneti-
cally adaptive strategy aimed at protecting
kin from infection or infestation (Tanaka and
Kinney, 2011) or from internecine conflict
(Riordan, 2019); and potentially to account
for suicide terrorism (Gallup and Weedon,
2013). Still others suggest that processes
of multi-level selection, in which a group’s
competitive success may be furthered by the
altruistic behavior of its members whether
genetically related or not, may offer adaptive
logic for extreme acts of heroism in battlefield
situations (Orbell and Morikawa, 2011). Some
theorists offer a ‘mismatch’ variation on the
adaptation idea; that suicide may not be adap-
tive in current conditions but, as fossil behav-
ior carried over from ancient environments, it
might have been adaptive in the past (Aubin
et al., 2013; deCatanzaro, 1980, 1981).
Self-killing as a way to relieve kin of
the burden of one’s existence is the most
prominent adaptationist idea in suicide
research (Aubin et  al., 2013; Bering, 2018;
Soper, 2018; Syme et al., 2016). Brown and
colleagues, for example, develop deCatanza-
ro’s proposals and offer supportive empiri-
cal evidence (Brown et al., 1999, 2009). As
other researchers also record, various meas-
ures of burdensomeness do correlate with
suicidal thinking and behavior (Chu et  al.,
2017; Lester, 2014b) – although, it should
be noted, not strongly and no more strongly
than do many other risk factors (Franklin
et  al., 2017). Further empirical support
might arguably be found in ethnographic
reports from some tribal societies where a
group member too frail to survive the next
season or journey may sometimes take
part in ritualized assisted suicide, although
this practice is not common (Falger and
Falger, 2003).
These are all intriguing ideas. But the
state of play is that, after four decades, there
are no signs of a scientific consensus form-
ing around the conception of suicide as the
adaptive removal of unsupportable kin
(Bering, 2018; Lester, 2014b; Soper, 2018).
There are multiple empirical contraindica-
tions, including high notable suicidality
among young adults (De Leo et  al., 2013),
the gifted and talented (Delisle, 1986;
Voracek, 2006), people with high incomes
(Goldsmith et  al., 2002) and others who, it
can be presumed, are unlikely to be among
the most burdensome members of their
families. The general difficulty we perceive
is a lack of compelling evidence of special
design – a precise match between observ-
able form and biological function that is the
hallmark of adaptation:
Evolutionary adaptation is a special and onerous
concept that should not be used unnecessarily,
and an effect should not be called a function
unless it is clearly produced by design… (Williams,
1966, vii)
Three disconnects will illustrate the prob-
lem. First, is the absence of a causal connec-
tor between suicide as the means and the end
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
it is supposed to achieve: it is unclear why
the task of removing an individual should
call for suicide in preference to other solu-
tions that have stronger genetic logic (Bering
and Shackelford, 2004). If an individual has
to be removed, there is no particular reason
to expect that removal to necessitate a kill-
ing: abandonment, quarantine, or exile, say,
would achieve the same objective, with the
advantage of preserving at least provision-
ally the individual’s reproductive capability
(Wright, 1994). And even if a killing would
be in a conspecific’s reproductive interests,
the expectable outcome is still not suicide: the
killing would more logically be done by that
conspecific, who has likely better informa-
tion and genetically more to gain (O’Connor,
1978; Skinner, 1969; Soper, 2018). Indeed the
empirical evidence among humans and other
species points to infanticide or fratricide,
rather than suicide, as the way unsupportable
kin are removed (Dickeman, 1975; Harris,
1974; Hrdy, 1979; O’Connor, 1978).
Second, it is doubtful in principle whether
a biological stimulus could exist that would
trigger suicide as an adaptive response.
Perry (2015) presumes that if an organism
is to decide whether or not life is genetically
worth continuing, then it would need to be
equipped with some kind of ‘inclusive fitness
monitor’ that can compare the reproductive
value of life overall versus death. For sure,
sophisticated measuring devices have been
proposed elsewhere by evolutionists, such
as a ‘sociometer’ that may alert the organ-
ism to threats to social supports (Leary and
Guadagno, 2011; Leary et  al., 1995). But
Soper (2018) argues that a serviceable ‘inclu-
sive fitness monitor’ would entail an alto-
gether higher order of complexity; it would
need to take a view on, among other things,
current and future kin members’ reproduc-
tive prospects and the future carrying capaci-
ties of their environments. He claims that it
would be so all-encompassing that it would
lack the specific input–output associations
to which selection responds. Modern theory
holds that such a general-purpose mechanism
59
is unlikely to evolve (Symons, 1992; Tooby
and Cosmides, 1990b).
Third, given the severity of costs out-
lined in Table 3.1, the hypothesized benefits
accruing via kin or group selection seem
to fall decisively short, in power and reli-
ability, of what would be required to justify
self-killing as a fitness investment. The sug-
gested payoffs are highly contingent, it being
far from certain that the reproduction of an
individual’s family or group would improve
as a consequence (Lankford, 2013). On the
contrary: Table 3.1 suggests that suicide can
be predicted to add to, not lift, reproductive
difficulties for those left behind.
Summing up, it is hard to argue that sui-
cide shows marks of special design. At the
margins, some writers suggest possible adap-
tive functionality in heroic deeds in battle and
similar emergencies, which might be classed
as ‘altruistic suicide’ following Durkheim’s
(1897/1952) nosology (Humphrey, 2018;
Orbell and Morikawa, 2011). Other writers
question the usefulness of ‘altruistic suicide’
as a concept (Johnson, 1965; Townsend,
2007). The phenomenon is unusual, as
Durkheim himself acknowledged. And it
may not anyway help to class as ‘suicide’
acts where killing of the self is not of itself
the primary intention, but where, rather, death
happens incidentally in pursuit of some other
endeavor (Lankford, 2013). For these reasons
Soper (2018) argues that battlefield scenarios
are unlikely to shed light on private, solo
self-killing – what Cholbi (2017) calls ‘run-
of-the-mill’ suicide – which probably is not
and never would have been adaptive.
‘BY-PRODUCT’ THEORIES
With only three types of evolutionary expla-
nations available, and if suicide cannot be
credibly ascribed to the first two (that is, it
probably evolved neither as noise nor as an
adaptation), then Soper (2018) argues that
whatever remains has prima facie appeal.
60 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Explicitly or implicitly, several theories fall
into this third category, characterizing sui-
cide not as adaptive but as a noxious side effect
of some other trait that, despite the severe cost
of suicidality, is adaptive in the round. The
following section reviews prominent ideas of
this type.
Pleiotropy
The idea of senescence, as we discussed in
above, did not seem to carry us far in under-
standing suicide. But senescence is a special
case of a broader genetic phenomenon, pleiot-
ropy, which may take us further (Williams,
1992; Wilson, 1980). Pleiotropy occurs where
genes express in the same individual in multi-
ple phylogenetic outcomes, some beneficial,
some harmful. Particularly if a beneficial
effect is felt early in the individual’s lifetime,
then it may more than compensate for an inju-
rious one that emerges later – hence the link
with senescence. We can safely presume that
suicide arises in this general way, as a harmful
effect of genetic material that is fitness-
enhancing on average. This is not to say that a
‘suicide gene’ exists or will ever be found:
genetic predisposition to suicide appears to be
spread thinly across a large number of genes,
each of weak effect, interacting with each
other and with environmental factors (Marušič
and Swapp, 2004; Mullins et al., 2019).3 More
likely, as deCatanzaro (1980) suggests, inten-
tional self-killing occur as an incidental effect
of a species-typical genome, the expression of
which has proved adaptive overall in the
course of human evolution. But then, what
adaptive aspect of this species-typical genome
could by-produce suicide?
Suicide as Communication
One posited answer, offered by Hagen and
Syme, (Hagen, 2002; Syme et al., 2016; Syme
and Hagen, 2018), is that deaths may occur
as an unfortunate by-product of suicidal
interpersonal communication. Their original
idea is that an otherwise powerless individ-
ual, modally a young woman, may, at the
extreme, threaten to or try to kill herself as
the only available means by which she can
induce kin to attend to her honest genetic
needs. It may be in her reproductive interests
to make a potentially high-stakes gamble,
informed unconsciously by an inclusive fit-
ness calculation. Suicidal death, from this
perspective, can be seen as an unfortunate
gamble lost. The behavior posited to be adap-
tive is the threat of, or attempt at, suicide; the
potential for death being necessarily con-
comitant for the threat to be credible (Wiley,
2020). A recent variation of the idea, also
with the inherent risk of a lethal, maladaptive
outcome, is that a suicide attempt may con-
stitute a costly signal of remorse (Syme and
Hagen, 2018).
Perhaps this line of theorizing may use-
fully shed light on non-suicidal self-injury
and unintended fatalities, both outside the
scope of this discussion. Researchers have
long hypothesized a “cry for help” compo-
nent in sub-lethal self-harming behaviors
(Shneidman and Farberow, 1961), phenomena
which may be significantly distinct from sui-
cide (Kapur, Cooper, O’Connor and Hawton,
2013; Selby et  al., 2014; Stengel, 1970).
Among the chronically suicidal, it is possi-
ble that a cycle of reinforcement may arise in
which carers’ well-meaning responses inad-
vertently provoke yet more behavioral cries
for help (Linehan, 2020). There may also be
strong commonsense appeal in attributing sui-
cide to communication. There is an important
caveat in this regard: folk theorizing about the
cause of suicides is evidently not an objective
data source. It is rather, at least in large part,
a product of encultured post-rationalization
(Atkinson, 1978; Solano, Pizzorno, Pompili,
Serafini and Amore, 2018; Soper, 2019a).
For example, alongside communication-type
explanations, and presumably as reliably,
non-western informants often also blame
suicides on evil spirits, as Syme et al. (2016)
themselves found. Observers frequently intuit
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
interpersonal motives for attempted suicides
too, but it should be born in mind that these
interpretations rarely agree with actors’ own
explanations: and where they do agree it may
be because surviving attempters sometimes
confess motives that they think others expect
to hear (Bancroft et  al., 1979). All that said,
we question whether communication offers
a satisfactory evolutionary explanation for
intentional self-killing, partly for the same
reasons that we found more directly adapta-
tionist models wanting. We will highlight two
difficulties (a critique may also be found in
Soper (2018)).
The first problem recalls the hallmark
of adaptation: evidence of special design
(Williams, 1966, 1996). It is an intuitive call,
but suicide, whether completed, attempted
or threatened, does not strike us as likely
custom-designed for communication. People
who seriously intend to take their own lives,
presumably against kin’s wishes, would logi-
cally be expected not to communicate that
intent, at least not sufficiently to invite inter-
ference. With rare exceptions, the empirical
picture seems to fit this expectation. Perhaps it
is different in some parts of the world – most
of the evidence is from Western sources – but
suicides are usually characterized by non-­
communication: privacy and non-disclosure
are the norm. That they tend to happen without
effective warning may be inferred from rela-
tives’ immediate reaction to the news: typi-
cally shock, confusion, and disbelief (Chow,
2006; Dyregrov et  al., 2012). Far from reg-
istering a communicated message, bereaved
families are generally left bewildered by the
act’s apparent senselessness (Jordan and
McIntosh, 2011; Wertheimer, 2014). Non-
communication is characteristic of attempted
suicides too (Maple et al.). Where an attempt
is survived, close kin are usually not told about
it either before or after the event, and they usu-
ally remain unaware even long afterwards
(Brezo et  al., 2007; Walker et  al., 1990). At
the same time, one can imagine any number
of other deviances, potentially costly but not
ordinarily suicidal, available for use if a drastic
61
threat or costly signal were needed: desertion,
sexual infidelity, self-mutilation, infanticide,
sabotage, arson, and so forth. In sum, it is not
clear that suicide constitutes an outstandingly
good solution to the needs of coercive, or any
other, communication.
The other difficulty recalls, again, the
extreme and predictable fitness penalty of a
suicide attempt, whether lethal or survived
(Table 3.1). It is hard to imagine a commen-
surately extreme and predictable fitness ben-
efit that could, even in theory, be won from
a suicidal communication. Empirically, the
epidemiology shows no obvious pattern of net
fitness gains. Lesser upsides offered as sup-
porting evidence in Syme et al.’s ethnographic
analysis (2016, supplementary material, table
S3) seem to fall well short in specificity (e.g.,
‘Manipulate parents’) and reproductive impact
(‘Prevented unwanted ear modification’) to be
plausibly sufficient, as biological prizes, to
justify the fitness losses and risks taken in try-
ing to kill oneself. The strongest claimed pay-
offs are cases where actors are alleged to have
sought sexual concessions, such as ‘Prevent
unwanted marriage’ and ‘Concubine moved
out of house’ – psychologically appealing,
no doubt; but in a fitness calculation, still not
credibly worth dicing with genetic extinction
for their sake. If there is not a plausible net fit-
ness gain to be had from going through with a
threat, then it is unclear on what fitness grounds
such a threat would be accepted as being cred-
ible. This is not to say that there may not be
social utility in threatening to kill oneself, or
any other threat for that matter. The error, we
suggest, is to confuse non-evolutionary and
evolutionary processes – to confuse utility
with biological fitness.
Stepping back, we question a general
assumption that underlies hypotheses dis-
cussed so far, that the fitness payoff of what-
ever process it is that produces suicide derives
from the suicide behavior itself. In principle,
pleiotropy does not require there to be any
obvious connection between a selected trait
and its noxious concomitants (Williams,
1992). The explanatory gap between suicide’s
62 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
hypothesized benefits (weak, and contingent)
and evident costs (severe, and predictable;
Table 3.1) suggests that we probably need to
look beyond the act itself for a categorically
more powerful biological impetus. With this
in mind, we will next look at some prominent
evolutionary ideas circulating in mainstream
suicidology.
The Interpersonal–Psychological
Theory of Suicide (IPTS)
Another conceptual framework, the
Interpersonal–Psychological Theory of
Suicide (IPTS), deserves attention. In various
formulations it may currently be suicidology’s
most prominent theory (Hjelmeland and
Knizek, 2019; Lester, 2019; Paniagua et  al.,
2010), and it explicitly draws on some evolu-
tionary ideas, including the notion that suicide
may have come about as an evolutionary by-
product. In its original form IPTS holds that
suicide results from the co-occurrence of three
supposedly sufficient conditions: (1) a percep-
tion of being a burden to one’s family, (2) a
thwarted desire to belong, and (3) a learned
capability to enact lethal self-injury (Joiner,
2005; Van Orden et al., 2010). We will discuss
each in turn.
The first element, a feeling that one is a
liability to loved ones, has been mapped by
some commentators onto deCatanzaro’s (1980)
sociobiological idea of burdensomeness –
critiqued earlier – to which suicide is allegedly
the organism’s genetically adaptive response
(Aubin et al., 2013; Brown et al., 2009). But
in what they call a ‘sociobiological exten-
sion’ of their own theory, IPTS’s authors
depart from deCatanzaro’s view of burden-
someness in that they posit (human) suicide
to be intrinsically maladaptive: colonial
insects and humans are said to be equivalently
eusocial, but while lethally self-sacrificial
behaviors are understood to have inclusive
fitness logic for insects, supposedly the same
response among humans is seen as an error
– a ‘dysfunction, misfiring, or derangement
of the adaptive behavioral suite evolved as a
facet of eusociality’ (Joiner et al., 2017: 71).
Evolutionary rationale for this distinction
isn’t provided, but the need to make a distinc-
tion seems to stem from the authors’ equating
of insect and human sociality – a question-
able premise given the special familial make-
up of insect colonies noted earlier. Adding to
the confusion is what appears to be a mixing
of biological process and moral evaluation
(Gorelik and Shackelford, 2017) something
that has long colored scientific discourse in
this field (Soper, 2019a; Zilboorg, 1936a).
IPTS’s second element, thwarted belong-
ingness that may motivate self-killing, is on
firmer theoretical ground and is where sui-
cide is envisioned explicitly as a dysgenic by-
product (Joiner, 2005; Van Orden et al., 2010).
The aversiveness of rejection, abandonment,
and similar interpersonal stressors has cer-
tainly been attributed to evolutionary origins:
psychological pain probably functions as an
ancient alarm system, warning of potentially
fitness-damaging social losses (Baumeister
and Leary, 1995; Bowlby, 1969/1997, 1973,
1980/1991). Suicide can be understood as a
way to escape from this adaptive social dis-
tress. We will return to this idea.
Less strong in its theoretical underpin-
nings is the third component of IPTS, a
hypothesized learned capability for lethal
self-injury. The idea derives from a centuries-
old belief that suicide defies an ‘instinct for
self-preservation’. This supposedly universal
natural drive must be overpowered, IPTS’s
authors claim, before suicide can be carried
out (Joiner, 2005; Van Orden et  al., 2010).
IPTS’s originator, Joiner (2005), ascribes
evolutionary credentials to an ‘instinct for
self-preservation’, but the notion faces prin-
cipled objections from evolutionists, not
least because there is no known biologi-
cal means by which such a superordinate
drive could come about (Kirkpatrick and
Navarrete, 2006; Soper, 2019a). Modern the-
ory holds that a general-purpose motivational
device would be underspecified, lacking the
recurring connections between proximal
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
stimulus and fitness-impacting response that
are required for selection to take hold: a sys-
tem without specific links between inputs and
outputs is unlikely to be favored (Buss, 1990;
Buss and Penke, 2015). The same difficulty,
it will be recalled, arose above in discussions
of an ‘inclusive fitness monitor’. Soper (2016,
2018) suggests that if an antisuicide mecha-
nism were reconceived within the tenets of
evolutionary psychology, as a special rather
than general-purpose device, then useful
implications follow. We will come back to this
point as well.
Suicide as an Escape from Pain
Pyschological pain may drive the phenotype
to seek relief in a way that is destructive for
the genotype. We saw this idea underlying
IPTS’s second component (above), and indeed
it can be traced to suicidology’s formative
writings. In the words of Henry A. Murray
(the clinician whom Edwin Shneidman, the
acknowledged father of suicidology, saw as
his mentor),
Suicide does not have adaptive (survival) value but
it does have adjustive value for the organism…it
abolishes painful tension. (Murray and Kluckhohn,
1948: 15, original italics)
By this perspective, suicide offers escape
(Baechler, 1975/1979; Baumeister, 1990;
Shneidman, 1993, 1996, 2005). A related
idea, but with much older philosophical roots,
is that self-killing may be a rational response
to difficult life circumstances (Maris, 1982;
Mishara, 2003). General reviews are avail-
able elsewhere of ‘suicide as escape’ theories
(Gunn, 2014) and rational suicide (Lester,
2014a). Of interest for our purposes is their
recognition, implicit or explicit, of the sui-
cidogenic power of emotional pain (Selby
et al., 2014).
There are two points to note. First,
although pain systems can malfunction, pain
almost certainly has ancient adaptive origins
(Nesse and Schulkin, 2019). Pain’s signal
63
enables the organism to navigate fitness haz-
ards in its environment, both physical (Wall,
1999) and, as we have already noted, social
(MacDonald and Leary, 2005). And in order
to fulfil its navigation task, pain is necessar-
ily motivational: it is designed precisely to
force the organism to take action to relieve it
(Auvray et al., 2010; Corns, 2014; Melzack
and Casey, 1968). Hence, pain hurts. As the
leprosy surgeon Paul Brand observed, pain
is a valuable gift, albeit a gift nobody wants
(Brand and Yancey, 1993).
Second, there is an accord among suicide
theorists and empirical researchers about
the kind of pain that most powerfully moti-
vates action – whatever action – to obtain the
required relief. Although physical pain also
associates with suicidality (Klonsky et  al.,
2019; Klonsky and May, 2015), emotional
pain is more usually held responsible. The
unbearable emotional state that can induce
people to take their own lives is encapsulated
by Shneidman’s (1993) neologism, psychache:
Psychache refers to the hurt, anguish, soreness,
aching, psychological pain in the psyche, the mind.
It is intrinsically psychological – the pain of exces-
sively felt shame, or guilt, or humiliation, or loneli-
ness, or fear, or angst, or dread of growing old or of
dying badly or whatever. (1993: 51; original italics)
There is also implicit agreement across
more than a century of suicide research that
unbearable emotional pain usually stems from
social troubles. The disagreement is rather
about what hue of social trouble is deemed
most troublesome. Durkheim (1897/1952), for
example, cited detachment (‘anomy’) as the
chief driver. A hundred years later, Williams
and colleagues ascribe suicide to feelings
of social defeat and entrapment (Williams,
1997; Williams and Pollock, 2000; Williams
et  al., 2005). Williams’ ideas reappear in
another theoretical framework, one that rivals
IPTS, the Integrated Motivational–Volitional
Model of Suicidal Behavior (IMV) devised
by O’Connor and colleagues (O’Connor,
2011; O’Connor et  al., 2016). IPTS, it will
be recalled, highlights yet other varieties of
64 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
interpersonal distress – thwarted belonging
and burdensomeness. And so on.
Gunn (2017) connects these threads in an
explicitly evolutionist frame; his Social Pain
Model (SPM) registers that suicidogenic psy-
chache is usually activated by various forms of
damaged or threatened social relations. This
social pain, as with pain generally, is both
adaptive and intrinsically aversive. It is adap-
tive in that it is designed to alert the organism
to the fitness threat posed by detachment –
rejection presaging near certain death for our
hunter-gatherer forbears (Bjorklund et  al.,
2010; Eisenberger and Lieberman, 2004;
Eisenberger et  al., 2003; Lieberman, 2013).
And social pain is necessarily painful in order
to induce action to alleviate it. Unfortunately,
suicide offers a genetically self-destructive
answer to this biological imperative to act.
In sum, a weight of empirical and theo-
retical evidence points to suicide instantiat-
ing as an unfortunate by-product of pain,
notably social pain, this being integral to the
navigational equipment that humans need
for maintaining attachments in large and
close-knit groups. The aversiveness of social
pain, adaptively, demands action to end it –
a demand that can be met maladaptively by
self-extinction.
‘Pain-and-Brain’ Model
But social pain alone cannot be sufficient
explanation for the evolution of suicide. If it
were sufficient, suicide would presumably be
found elsewhere among primates and other
higher social animals. Indeed, solitary ani-
mals would expectably be vulnerable too: to
reprise the point, pain (social or other) is
biologically designed not to be tolerated – it
demands that the organism act to end or
escape it – so, any animal that could termi-
nate its pain by terminating itself would rea-
sonably be expected do so (Perry, 2016;
Soper and Shackelford, 2018). We are still
searching for an explanation for suicide as a
narrowly human response. The search space
is even narrower than this because, as well as
nonhumans, certain human populations too
are protected from taking their own lives,
however painful their circumstances: young
children (Nock et  al., 2013; Shaffer, 1974)
and the mentally incapacitated (Merrick et al.,
2006; Seyfried et al., 2011). Their immunity,
or virtual immunity, also needs to be
accounted for. Noting commonality across
these groups, Baechler (1975/1979) draws a
parsimonious conclusion that deliberate self-
killing presupposes a minimum level of intel-
lectual functioning. Soper (2017, 2018)
concurs, arguing that it’s only after more than
a dozen years of cerebral development that
species-typical humans, and apparently only
humans, acquire sufficient capacity for logi-
cal thinking, foresight, and planning, for sui-
cide to become a conceivable and practicable
response. Humans alone appear to cross what
Perry (2014: 110) describes as a ‘cognitive
“floor” for suicide’, usually in adolescence.
Thus, a second necessary condition for sui-
cide emerges: cognitive competence. This may
be central for understanding the behavior’s
evolutionary origins. If deliberate self-killing
presupposes the crossing of a developmental
threshold during the individual’s lifetime, then
we can deduce that an evolutionary counter-
part, a phylogenetic threshold, also had to be
surpassed at some point in human prehistory.
Humphrey (2011: 211) makes the point by
quoting Stengel (1970: 37)…
At some stage of evolution man must have discov-
ered that he can kill not only animals and fellow-
men but also himself. It can be assumed that life
has never since been the same to him.
Humphrey (2018) goes on to ascribe this
pivotal discovery to the arrival, around 100,000
years ago, of a suite of uniquely human cogni-
tive skills, including abstract thinking, mental
time travel, self-consciousness, and sophisti-
cated theory of mind. These are virtuoso dem-
onstrations of the flexible, general-purpose
style of thinking of Homo sapiens sapiens –
promiscuous intelligence favored by selection
because it conferred potent ecological, social,
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
and sexual advantages (Flinn and Alexander,
2007; Humphrey, 1976; Pinker, 2010; Tooby
and DeVore, 1987). But the human brain
is ‘expensive tissue’ (Aiello and Wheeler,
1995): encephalization brought with it
heavy costs, including energetic demands
(Aiello and Wheeler, 1995), mutational load
(Keller and Miller, 2006), obstetric chal-
lenges (Miller and Penke, 2007), problems
of thermoregulation (Falk, 1990), and the
burden of supporting young while the brain
matures (Flinn et  al., 2007). Suicidality can
be understood as another such cost, a price
we pay for a mind so free ranging that it can
conceive even of its own mortality (Bering
and Shackelford, 2004; deCatanzaro, 1980;
Soper, 2018; Suddendorf, 2013). To sum
up, it appears likely that suicide evolved as
a by-product of not one adaptation, but two
combined: pain (usually social); and species-
typical human cognition. Soper calls this a
pain-and-brain theory of suicide.
Suicide as an Adaptive Problem
The above discussion suggests that scientific
consensus, if loose and implicit, can be seen
coalescing around some form of ‘by-product’
explanation for suicide’s ultimate origins. A
pain-and-brain framework in particular does
not appear contentious: it seems to offer a
point of agreement among prominent suicide
theories, it accords with the epidemiology of
suicide, and connects to knowledge bases of
neuroscience, anthropology, and elsewhere.
But it raises new and difficult questions,
mainly because, as Soper (2016, 2018, 2019a)
infers, the posited pain-and-brain conditions
are not only necessary for suicide, but logi-
cally sufficient: pain provides the motive, and
mature human cognition provides the means.
If the pain-and-brain assessment is correct,
then the fitness threat of suicide exists in
potentia among virtually all of us. All species-
typical humans feel, and are motivated to
escape, pain (Benatar, 2015; Brand and
Yancey, 1993); and all species-typical human
65
adults, equipped with much the same cognitive
machinery (Tooby and Cosmides, 1990a),
have self-removal available as an exit route
from pain. A consequent ‘lure of death’
(Humphrey, 2018) has probably featured in
our evolutionary environment at least since
the advent of behaviorally modern humans.
Perhaps for much longer – presumably,
indeed, ever since hominid intelligence began
to approach that of a modern-day pubescent
child (Soper, 2019b). By implication, suicide
presents a harmful variety of, in evolutionary
parlance, adaptive problem:
Adaptive problems are evolutionarily long-enduring
recurring clusters of conditions that constitute
either reproductive opportunities (e.g., the arrival
of a potential mate, the reflectant properties of
light) or reproductive obstacles (e.g., the speed of
a prey animal, the actions of a sexual rival, limited
food supplies for relatives). (Cosmides and Tooby,
2000: 96)
Adaptive problems seek out adaptive solu-
tions (Mayr, 1965; Tooby and Cosmides,
1990b; Williams, 1996). That almost all
adults could be expected to take their own
lives, but few do, indicates that adaptive
solutions to the suicide problem are in fact
in place. Without them, we as individuals,
and we as a species, would not be here. At a
phylogenetic level, hominid evolution would
not have found a way through the cognitive
floor for suicide, the floor acting as a ceil-
ing of viable intelligence: in this light suicide
emerges as perhaps the pre-eminent adaptive
problem of our species (Soper, 2019b). Our
existence presents a puzzle not so much of
suicide but non-suicide. What stops most of
us killing ourselves?
Evolved Antisuicide Defenses
In seeking to answer this question, why not
suicide, we can set aside the pre-Darwinian
notion of a general-purpose survival instinct –
critiqued briefly above and more fully else-
where (Buss, 1990; Buss and Penke, 2015;
Kirkpatrick and Navarrete, 2006; Soper, 2019a).
66 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
The likely solution would look more like the
customized ‘patch 7.822’ artificial intelli-
gence fantasy that began this chapter (Perry,
2016). If a fleet of androids dealt with every
difficulty by, uselessly, switching themselves
off, the problem would call for a software
update to eliminate that particular behavioral
option. In actuality, the human brain, a bio-
logical computer (Tooby and Cosmides,
2005), has presumably been retrofitted with a
comparable fix, devised by selection. We
next need to ask: how would such a patch
operate? And what would its successful oper-
ation look like?
Two sorts of antisuicide programs have been
hypothesized in recent years (Humphrey,
2018; Miller, 2008; Soper, 2016, 2017,
2018). The first are evolved psychological
mechanisms (Buss, 1995); the second are
culturally propagated barriers. There won’t
be a neat dichotomy in reality because many
evolved psychological mechanisms rely on
cultural inputs (Tooby and Cosmides, 1992),
and many cultural devices exploit evolved
psychological preparedness for learning in
their particular domains (McNally, 2016).
Nonetheless the distinction is interesting –
partly because, although both sorts of devices
might be expected to emerge, there is dis-
agreement over their likely chronology and
relative importance. Humphrey (2018) rea-
sons that psychological mechanisms (what he
calls ‘natural’ or ‘innate’ protections) would
not have arisen as the first or foremost pro-
tection because of their slowness to evolve.
Soper (2018, 2019a) argues the contrary,
that they probably came first, emerging at
least in part during a reconfiguration of ana-
tomically modern humans’ ‘wetware’ ahead
of the cultural explosion of the Later Stone
Age (Klein and Edgar, 2002). The evolution
of human intelligence may have been held at
the cusp of suicidality for perhaps 150,000
years or more, while largely autonomic solu-
tions to the suicide problem were assembled
and refined. The pressure favoring such solu-
tions would have been intense due to a com-
bination of forces: runaway selection pushing
for greater computing power (Geary, 2007;
Miller, 2000), while intelligentsia who lacked
adequate protection were culled. Soper pos-
its that it was only after basic defenses were
installed that human intelligence could pro-
gress to the level of being able to enculture
antisuicide ideas, proscriptions which pre-
suppose a capacity to conceive of personal
mortality.
It is possible to deduce something of the
likely design features of antisuicide defenses,
as we will next discuss, by adopting evolu-
tionary psychology’s ‘task analysis’ method
(Tooby and Cosmides, 1992) – inferring
likely parameters of an adaptive solution from
the nature of the adaptive problem. Soper
(2018) suggests that, in view of the severity
of the threat, antisuicide devices are prob-
ably arranged as serial fortifications, each
line guarding the position behind and deploy-
ing special stimulus-response mechanisms
to that end. Perhaps simplifying, he suggests
a framework that splits into front and rear
defenses. At the back are emergency inter-
ventions, labelled keepers to connote the pri-
mary role of a goalkeeper on a soccer team:
on stand-by much of the time, keepers leap
into ‘Save!’ mode at times of crisis, and are
all that stands between an attacking shot and
disaster. Front-line protections, in contrast,
are continuously active: following the soccer
analogy, Soper labels them fenders to suggest
a team’s other defensive players. Fenders’ job
is to stop crises arising in the first place. The
default outcome, if players can’t fulfil their
respective tasks, is a conceded goal – a sui-
cide attempt – which, as noted earlier, may be
genetic ‘game over’.
Soper argues that defenses’ triggering
inputs and behavioral outputs would be
expected to address both of suicide’s dual
pain-and-brain drivers, and in distinct ways.
To borrow the soccer analogy again, the fit-
ness hazard of suicide can be conceived as an
opposing team with only two forward play-
ers, ‘Pain’ and ‘Brain’, and they pose a threat
only when they attack together. They have
distinct styles of play, and the defending team
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
must adopt customized pain-type and brain-
type tactics to neutralize them.
Let us look first at the system design of
keepers, the posited emergency defenses. To
detect an imminent threat from ‘Pain’, keep-
ers must be ready to respond to the experience
of chronic and intense emotional distress. To
detect an incoming ‘Brain’ threat, keepers
must also be alert to the surpassing, in adoles-
cence, of the cognitive threshold for suicide.
Cues for both ‘Pain’ and ‘Brain’ must be pre-
sent for keepers to mobilize. And, likewise,
there are two, and only two, strategies avail-
able by which keepers can block incoming
attacks: pain-type and brain-type. Pain-type
keepers make self-killing unnecessary: they
numb, distract from, or otherwise weaken the
felt urgency of emotional pain as a motivator
for escape. Brain-type keepers deny the means:
they interfere with intellectual functioning
enough usually to prevent an effective suicide
attempt being organized. We will discuss keep-
ers in more detail after this overview. The main
point to note for now is that keepers’ interven-
tions may be highly costly to the organism –
and they are evidently not failsafe: hence the
need for the pre-emptive work of fenders.
As for fenders, the forward line of protec-
tions; these are hypothesized also to deploy
in pain- and brain-type forms. Pain-type
fenders titrate the organism’s exposure to
emotional pain within tolerable limits. They
use what could loosely be called positive
psychology (Efklides and Moraitou, 2013) to
hold humans safely distant, most of the time,
from the potentially lethal danger that resides
lop-sidedly in negative affect. Soper suggests
they may work as four subsystems. First, is a
homeostasis of affect around a resting point
that is happier than neutral (Heintzelman
and King, 2014), a base at which shocks can
generally be absorbed without great disrup-
tion. Second, is a regulation of conscious
contact with potentially painful realities – a
self-serving self-deception that, empiri-
cally, characterizes so-called psychoanalytic
(or psychodynamic) defenses (Paulhus and
Buckels, 2012). Third, is a manufacturing
67
of positive affect (Kuhl et al., 2015; Layard,
2011) via, inter alia, investment in pleasura-
ble activities beyond what would be econom-
ically justified by normal animalian needs
of survival and reproduction (Pinker, 1997).
And fourth, is the selection and maintenance
of a hopeful, measuredly fictitious, mental
model of self-in-the-world. This paradigm,
more or less spiritual, holds that the universe
and its people are not wholly indifferent to
our wellbeing, and that our futures are not
entirely doomed to the pain that would attend
a purely Darwinian struggle. Thus, operating
within (to adapt Baumeister’s (1989) phrase)
an optimum margin of delusion, we perceive
the world not as it is but through a rose-tinted
lens. This enhanced reality system is under
continual assault by factual counterevidence,
and hence requires us continuously to defend
it, literally as if our lives – or, at least, mental
health – depend on it. Arising from this pro-
tective worldview, Soper says, is a costly pro-
pensity for selflessness and charity – that is,
generosity that goes beyond the reciprocal cal-
culus of economics. Taken as a whole, these
sundry irrational-looking morale-boosters are
not easy otherwise to explain and connect, but
they may be understood to work as integrated
psychological machinery, designed to prevent
crises of willful self-destruction and the neces-
sity for keepers to intervene.
A separate set of evolved, culturally learned
protections, independently hypothesized by
Miller (2008), Humphrey (2018), and Soper
(2018), are designed to block access to the
idea of suicide. They are brain-type fenders
in Soper’s scheme. He suggests they propa-
gate by multi-level selection and form three
serial walls: a thought-inhibiting taboo; the
expectation of a fearful afterlife; and a mor-
alistic stigma. The deterrent function of this
last barrier, unfortunately but perhaps nec-
essarily, involves exemplary punishments
for suicides and their kin, some noted in
Table 3.1. The dismantling of these prohibi-
tions, the acceptance of suicide as a normal
topic of conversation and a reasonable solution
to trying circumstances, can release a surge of
68 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

suicides (Huber, 2015/2019). Such a wave may A summary graphic (Figure 3.1, from Soper,
perpetuate for generations (Macdonald, 2007): 2018) illustrates how the various posited pro-
once lost, cultural defenses may be exception- tections may interrelate. Out of a great num-
ally difficult to reinstate (Soper, 2018). ber of potential suicidal incidents, symbolized

Figure 3.1  Summary of posited antisuicide defences

Source: Reproduced from Soper (2018).
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
by a mass of dots at the top of the diagram,
only a few find their way through the for-
tifications to materialize as actual suicide
attempts, indicated by a couple of dots at the
bottom. By focusing not on supposed (possi-
bly inscrutable; Soper, 2019a) proximal cau-
sation of suicide but on evolved machinery
posited to stop suicide, the graphic departs
conceptually from the style of flow chart that
usually accompanies presentations of suicide
theory (Gunn, 2019).
PREDICTED FEATURES OF ‘KEEPER’
LAST-LINE ANTISUICIDE DEFENSES
This section focuses on Soper’s (2018) task
analysis for keepers – the hypothesized last
line of defenses, which react to exigent risk.
These ideas are novel, and not mainstream in
either suicidology or evolutionary psychol-
ogy, but we suggest they warrant attention
for three reasons. First, their claim to logic –
keepers’ design features being supposedly
deducible a priori from the nature of suicide
as an adaptive problem – is central to the
theory and calls for scrutiny. Second, these
features may be read as predictions, amena-
ble to empirical confirmation or falsification.
Third, if correct, they may have important
repercussions for suicide prevention, psychi-
atry, and wider mental health policy, as we
will discuss.
Keepers’ posited features are presented in
20 items, (a) to (t) below, the headings taken
from a tabulation in Soper (2018: 145).
‘Pain’ Input
With suicide modelled as an answer to the
imperative to escape pain, Soper argues that
the emotional aversiveness of pain would
probably serve as the primary activator of
emergency antisuicide defenses. Keepers
would mobilize selectively among people in
potentially suicidogenic distress.
69
a. Keepers would be activated by chronic,
intense pain (subject to the develop-
mental condition of a ‘brain’ input). Soper
predicts that an activating ‘pain’ cue will combine
(by some unspecified algorithm) pain’s chronicity
and intensity. Chronicity, on the grounds that sui-
cides take time to plan and enact (Joiner, 2005),4
so potentially drastic countermeasures should
not be triggered by merely ephemeral upsets.
Intensity, because the stronger the pain, presum-
ably the more urgent the motivation to escape it.
An additional ‘brain’ input is set out in (d) below.
b. Input variable would be the unidimen-
sional aversiveness of pain, regardless
of the pain’s source or quality. Because the
pain-and-brain model views suicide as an adjus-
tive response to the generic aversiveness of pain,
irrespective of its origin, keepers will respond to
the degree, not kind, of triggering pain. The point
recalls Shneidman’s (1993) catch-all notion of
suicidogenic psychache; that any blend of shame,
grief, anomy, thwarted belongingness, burden-
someness, defeat, hopelessness, or whatever other
emotional distress can motivate suicidal escape.
Soper (2018) posits that, likewise, any combina-
tion can trigger an antisuicide defense. Social pain
is more likely to activate keepers than is physi-
ological pain only inasmuch as social pain is often
experienced as more painful (Gunn, 2017).
c. Keeper responses would be calibrated
so that the intensity of defensive out-
puts accords with the intensity and
chronicity of the pain input. The strength of
antisuicide responses would be expected to adjust
according to the scale of the threat: more intense
and longer-lasting distress should associate with
commensurately more robust countermeasures.
‘Brain’ Input
d. Keepers would not activate earlier than
the species-typical age of first onset of
suicide, in early adolescence, possibly
signaled by the onset of puberty. As
already noted, activation would presumably need
a threshold ‘brain’ condition to be met, linked
to developmental surpassing of the cognitive
floor for suicide. Because young children lack the
mental capability for suicide, there would be no
fitness benefit in keepers mobilizing among them
however distressed they may be. Soper (2018)
70 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
stops short of specifying how this ‘brain’ cue
operates; but he posits that a biochemical signal
connected with pubescence is probably involved
because, empirically, it is usually at or shortly after
this stage of life that suicide becomes enactable.
Interesting predictions may follow from this devel-
opmental association, as we will discuss later.
Deactivation
e. Keepers would demobilize spontane-
ously following a reduction in the origi-
nating pain input. Mirroring activation, keepers
should deactivate after, and only after, relief of the
potentially suicidogenic pain that activated them.
f. Deactivation would usually be slow,
gradual and delayed, especially with-
out an unambiguous ‘all clear’ signal.
Antisuicide defenses would work on a ‘better
safe than sorry’ principle, deactivating slowly for
the same reason that prey mammals, on high
alert after scenting a predator, stand down only
cautiously once the scent disperses: the extreme
potential cost of misreading an ambiguous ‘all
clear’ outweighs the cost of remaining too long on
guard (Blanchard, Griebel, and Nutt, 2011).
Specific Types of Keeper
Responses
g. Responses would aim to limit motiva-
tion for suicide (‘pain-type’); or limit
the capacity to organize suicide (‘brain-
type’). It was noted earlier that keepers are
predicted to block suicide by deploying ‘pain’
and ‘brain’ strategies, a dual process deduc-
ible, Soper (2018) argues, from suicide’s posited
pain-and-brain evolutionary causation. He offers
suggestions as to what these interventions may
entail, listed as bullet points in the ‘keeper’ boxes
towards the bottom of Figure 3.2. Ellipses indi-
cate that the lists may not be exhaustive.
  The first category, pain-type keepers, are tasked
with lessening the aversiveness of emotional
pain. Given the neurological overlap between
physical and emotional pain (Eisenberger and
Lieberman, 2005), Soper (2018) hazards that
keepers would probably co-opt pre-existing cir-
cuitry that moderates the felt intensity of physi-
cal pain (Wall, 1999). Pain-type keepers might,
for example, numb emotions autonomically in
a manner akin to the analgesic effect of physical
trauma. They could also manage pain exoge-
nously: by, say, ingestion of analgesics – exploiting
‘nature’s pharmacy’ as animals do (Engel, 2002);
by exploiting the phenomenon of pain-offset relief,
in which one pain can be relieved by applying
another, unrelated, stimulus; and/or by distrac-
tion (Eccleston, 2001). Other pain-type keepers
are posited to use psychological adjustments to
make pain bearable, such as the construction of a
rationale for suffering pain and/or an emotionally
tolerable, but partly imaginary, re-perception of
the environment (Eccleston, 2001).
The other category, brain-type keepers, deny
the means and opportunity for suicide. They
will attenuate the individual’s ability to plan
and implement suicide (and, incidentally, any
other complex task) by tactically disabling
psychomotor and executive resources.
Activation would expectably express inleth-
argy and/or targeted cognitive deficits, such
as indecisiveness and forgetfulness.
General Characteristics of Keeper
Responses
h. Keepers would drive compulsive and
involuntary behaviors, resisting con-
scious awareness and intervention. Suicide
being the kind of lethal, once-in-a-lifetime, fitness
threat that offers few opportunities for conditioned
learning and where to err can be fatal, it ought to be
addressed by preset and strongly obligate routines.
Keepers would present limited scope for being
voluntarily moderated. Indeed, the phenomenon of
instinct blindness prevailing (Cosmides and Tooby,
1994), they may operate beyond awareness.
i. Multiple forms of keepers are likely
to operate in an integrated fashion in
the same individual, concurrently and/
or temporally. Blends of pain- and brain-type
defenses would likely deploy in combinations in
the interest of system robustness and to spread
costs. In principle, keepers could appear in many
possible permutations, varying from person to
person and over time, responding to the varying
needs of culture, gender, life stage, personality,
and other aspects of individual difference.
j. Keepers are likely to be accompanied by
protracted anxiousness, and rumination
focused on emotional pain. As a point related
to (f) above, following the general pattern by which
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
animals respond to severe but uncertain fitness
threats (Blanchard, Griebel and Nutt, 2011), humans
at risk of suicide would expectably display prototypi-
cal anxiety. Recalling the analogy of prey mammals
scenting an unlocatable predator, their best strategy
is not flight, because they would as likely head
towards death as away from it. For humans vulner-
able to suicide, lethal danger similarly comes from no
discernible direction and there is likewise nowhere to
run. The priority in the presence of such an extreme,
ambiguous hazard is to stay on high alert and try
to infer its detail ‘beyond the information given’
(Waldmann et al., 2006). By this light, a compulsive
mental rumination, which may be a uniquely human
addition to what is otherwise a pan-­ animalian
anxiety state (Blanchard, Griebel, Pobbe et al., 2011),
would be an expectable response.
Goals and Trade-Off
Considerations
k. The compromise objective would be to
minimize the risk of suicide, while limit-
ing the imposition of new, potentially
drastic, fitness costs arising from acti-
vated keepers. Keepers are necessarily costly
to activate because they involve attenuating
the advantageous-on-average ‘pain’ and ‘brain’
primary adaptations that brought suicide in their
wake. Keepers have evidently not delivered zero
suicide risk, and would not be expected to do
so, because zero suicidality would presumably
be achievable only by commensurately zeroing
the fitness benefit of those adaptations. Rather,
antisuicide defenses would evolve only up to
an equilibrium; a point where the marginal fit-
ness gain to be had from further reducing actu-
arial risk matches the cost of the extra defenses
required to achieve that reduced risk.
l. Keepers would trigger sensitively, with
a high incidence of false alarms – many
affected individuals will not have con-
sidered suicide. As with the posited ‘brain’
cue, Soper does not specify the ‘pain’ input that
is hypothesized to mobilize keepers. Candidate
triggers would presumably include biochemical
correlates of pain. But they would probably not
include, despite its appealing specificity, conscious
planning of suicide projects. The informational
currency of the brain being emotional rather than
semantic, the organism’s central nervous system
would be oblivious to suicide plans unless they
71
were flagged as such by an emotional regula-
tory variable (Tooby and Cosmides, 2008). Soper
(2018) hypothesizes the existence of separate
defenses that deter suicide plans by using the
disgust mechanism, informed by culturally learned
inputs (so-called brain-type fenders, discussed
above). But he suggests that there may be no pre-
programmed biological tag marked ‘Suicide Plan’
to which keepers could respond. Commonplace
experience is that one can muse about suicide
without triggering a drastic autonomic response.
On this basis, keepers may not be good at differen-
tiating, from among emotionally distressed people,
those who have specific suicide plans from those
who don’t. Keepers would likely mobilize in some
individuals who, although in pain, may never have
entertained serious thoughts of taking their own
lives. Keepers erring on the side of caution ((f),
above), there may be many such false alarms.
m. Keepers may themselves become patho-
logical. In the same way that the physiologi-
cal immune system can malfunction, there may
be exceptional situations in which keepers may
become pathological. Soper posits the possibility,
for example, that antisocial behavioral outputs of
keepers, mobilized in response to social pain, may
invite more social pain, potentially creating a posi-
tive feedback loop.
Manifestations of Successful
Operation
n. Keepers would result in a low, but above-
zero, incidence of suicide in human popu-
lations. It follows from keepers’ compromise
objective ((k), above) that their success would
appear at a population level not as zero suicides
but as a minimal, biologically irreducible, residue.
Pockets of high, even demographically destabiliz-
ing, rates of suicide might arise, but these would be
short-lived and in due course supplanted, through
multi-level selection, by groups with suicidality held
at sustainable levels.5
o. Residual suicides would be intrinsi-
cally unpredictable at the level of
the organism. Functioning optimally as
part of a wider system of antisuicide defenses,
keepers would be expected to exploit and
exhaust any and all useful markers of actu-
arial risk. The few suicides that do occur, it
follows, can be viewed as statistical residuals –
suicidal trajectories that are amenable neither
72 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
to detection (not, that is, based on informational
priors available to the organism) nor, therefore,
to autonomic prevention. With keepers operating
properly, suicides should be ‘predictably unpre-
dictable’ (Soper, 2019a).
p. Activated keepers would be associ-
ated with suicide, but only inasmuch
as they associate with suicidal idea-
tion rather than with the enacting of
those ideas. Most suicides would expectably
be accompanied by activated keepers. Interesting
predictions arise from (k) and (l) above concern-
ing statistical associations between suicidal-
ity and activated keepers. Completed suicides
will likely be accompanied by signs of keepers
having activated – properly, although in these
instances ineffectively. Keepers should also corre-
late strongly with suicide plans and actions. But,
importantly, these correlations should hold only
insofar as keepers correlate with the emotional
prior – a generalized urge to terminate pain. By
analogy, one could imagine a warring city that
defended its centre against aerial bombardment
with less-than-perfectly effective anti-aircraft
guns deployed at the city’s outskirts. Gunners
open fire pre-emptively; they start shooting as
soon as bombers come within range. Hence,
gunfire would correlate strongly with bombing
of the centre, but only inasmuch as the outbreak
of gunfire correlates with bombers approaching
the outskirts – not especially with any surviving
bombers’ completion of their raids, progressions
which the gunners are trying to avert. Likewise,
keepers will respond only weakly, if at all, to the
pursuit of specific suicide projects among people
motivated to end their suffering, this being the
progression that keepers are designed to prevent.
q. Keeper responses would be nearly
always recoverable and survivable: they
should only rarely cause permanent dis-
ability. Almost any outcome short of death being
preferable to suicide, keepers’ interventions may be
extreme. Nonetheless, as immune-like responses,
they should usually not be degenerative. A protec-
tive denial of psychomotor energy, for example,
would expectably produce tactical lethargy, perhaps
for a while even complete immobility, but it should
not seriously compromise somatic functions.
r. Keeper responses would be nearly
always recoverable and survivable: they
should only rarely cause permanent dis-
ability. Because the pain-and-brain framework
views suicide as a dysgenic, but ­psychologically
rational, response to aversive affect, activated keep-
ers can be understood as tactical denials of ration-
ality – they sacrifice aspects of the organism’s
affective (‘pain’) and cognitive (‘brain’) functioning
in order to preserve life. People under the influ-
ence of activated keepers will find themselves not
only distressed, but also emotionally (‘pain’) and
intellectually (‘brain’) debilitated. Faced with this
inconvenience, sufferers and their families would be
expected to approach priests, healers, and the like
for explanations and remedial interventions. It can
be expected that keepers’ expressions are conspicu-
ous and probably well known to science, and we
may not need to look far to find evidence of them.
Species-Specific and Species-
Universal
s. Keepers would be species-specific: they
would not occur in nonhuman animals,
although homologue of their features
may be found in other mammals. Keepers’
style of response contrasts with the way animals
normally meet severe fitness hazards: just when
any other animal would be expected to bring all
its mental resources to bear on tackling a mortal
threat, humans facing the threat of suicide are
likely to find their affective and cognitive faculties
autonomically impaired. Keepers are a uniquely
human solution to a uniquely human problem.
Rudimentary precursors of their features may well
be found in nonhumans, particularly among other
primates, but these will be only vestiges of phyloge-
netic raw material that was co-opted and adapted
for antisuicide purpose in our species alone.
t. Keepers would be species-universal:
the same integrated system of keepers
would be found activating among popu-
lations of mature humans in all cultures
and historical ages. Keepers are genetically
transmitted, species-typical mechanisms. Although
the proximal cause of precipitating social pain will
likely vary according to cultural context, the same
suite of antisuicide responses to that pain will be
found in all sizeable human populations. That said,
as universal human features, keepers’ antisuicide
functionality may not be obvious because no
extant control population may be to hand to show
what the suicide rate would be, all else equal, were
it not for their interventions.
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
PSYCHIATRIC SYMPTOMS AS
ANTISUICIDE DEFENSES
The above section outlined an engineering
specification – a score of features that would be
expected of so-called keepers, reactive devices
designed to stop us taking our own lives when
we otherwise might. The specification follows,
Soper (2018) argues, from the pain-and-brain
model of suicide’s evolution; defensive sys-
tems would activate with ‘pain’ and ‘brain’
informational inputs, respond using ‘pain’ and
‘brain’ processes, and produce observable
‘pain’ and ‘brain’ outputs.
The predicted features of this system show
striking similarities, according to Soper, with
adult patterns of so-called ‘functional’ (i.e.,
not due to structural brain dysfunction, as
opposed to ‘organic’) common mental disor-
der, or CMD (Goldberg and Goodyer, 2005) –
sundry states that would usually be described
as depression, generalized anxiety, alcoholism
and other addictions, psychoses, obsessive/
compulsive disorders, non-suicidal self-harm,
and perhaps other diagnoses. He asserts that
the fit is so detailed and multi-faceted that it
is difficult plausibly to explain, except by the
proposal that CMD is, in fact, the expression
of antisuicide machinery. The hypothesis chal-
lenges a widespread preconception that CMD
causes suicide, rather than works to block sui-
cidal trajectories among those at risk (Mishara
and Chagnon, 2016). To ascribe suicide to
CMD may be to commit the post hoc fallacy:
to assume that because a preceded b, a must
have caused b. Recalling the analogy of air
defences, ill-informed citizens might under-
standably misplace blame for the bombing
of their city on the anti-aircraft gunfire that
usually foreshadowed it. The challenge is not
new: it has long been suggested within psy-
chiatry that depression, addictions, psychotic
delusions, and diverse other psychopatholo-
gies can protect against self-killing (e.g.,
Hendin, 1975; Himmelhoch, 1988; Hundert,
1992; Menninger, 1938). What is new is evo-
lutionary theory that predicts such a dynamic.
73
We note four interesting ways to inspect
the hypothesis. First, it offers a novel concep-
tion of ‘functional’ psychiatric diagnoses, as
the concurrence of potentially suicidogenic
pain alongside blends of antisuicide responses
to that pain. Soper (2018) offers a cross-­
tabulation (Figure 3.2) to illustrate how psy-
chiatric labels may describe various groupings
of keepers in this way, with equivalent defen-
sive components appearing under different
diagnostic headings. Protective emotional
numbing, for example, maps against criteria
used for diagnosing ‘major depressive disor-
der’, ‘psychotic disorders’, and ‘bipolar dis-
order’. According to this conception, keepers,
not diagnoses, are natural kinds.
Second, the hypothesis may offer parsimo-
nious explanation for a suite of statistical links
between suicidality and CMD which, without
the hypothesis, look unrelated and puzzling. We
note three. One is that almost all CMD diagno-
ses associate with a heightened vulnerability to
suicide (Cavanagh et al., 2003) – but, signally,
the diagnosis of severe to profound intellectual
disability associates with fewer suicides (Harris
and Barraclough, 1997) as the pain-and-brain
model would anticipate. Two is that, as pre-
dicted, CMD correlates with suicide only inas-
much as it correlates with the ideational prior.
CMD associates strongly with generalized
suicidal ideas; but only weakly, if at all, with
the progression from this ideation to the plan-
ning and execution of specific suicide actions
(Klonsky et al., 2016; Nock et al., 2015). Three
is that the developmental stage when CMD
tends first to appear – in the teen years (Kessler
et al., 2007) – coincides with earliest first onsets
of suicidality (Nock et al., 2013). It is not easy
otherwise to account for these correlations
individually let alone as a set. Together they
could be taken as a fingerprint of antisuicide
functionality.
Third, the existence of marked patterns
common to sundry diagnostic labels point
to a shared aetiology – specifically, it is sug-
gested, as mechanisms designed to prevent
potentially suicidogenic pain and suicidal ideas
 74

Suggested types of Diagnostic criteria of some common psychopathologies (A.P.A., 2013)

keeper anti-suicide
mechanism
Major Depressive Disorder Substance use dis- Generalised Non-suicidal Psychotic disorders OCD Bipolar
orders Anxiety self-injury disorders
Disorder(GAD)a (NSSI)b

PAIN-TYPE (weaken the • Feeling ‘empty’; ‘Having • Compulsive use • Worry, • Self-injury • Diminished • Obsessions; • Depressive
motivation for suicide) no feelings’; ‘Not caring of analgesics rumination (iii, iv) emotional compulsions episode (i)
(i) Autonomic numbing. any more’ (i) and other restlessness; expression (i) (iv, v, vii) • Manic/
• Craving foods (iv) mind-altering insomnia (vi, vii) • Delusions; hypomanic
(ii) Medicate the pain.
substances hallucinations episodes (iv, vii)
(iii) Pain offset relief. (ii, iv, v, vi, vii) (iv, v, vi, vii)
(iv) Distract from pain.
(v) Detach from pain.
(vi) Make sense of the pain.
(vii) Find reason to live with pain.
BRAIN-TYPE (restrict access to • Indecisiveness; • Compulsive use • Difficulty • Disorganised • Obsessions • Depressive
the means of suicide) diminished ability to of sedatives concentrating; thinking; (viii); episode (viii, ix)
(viii) Degrade ability to plan and think or concentrate (viii) (viii, ix) mind going Disorganised/ • Compulsions
enact tasks. • Loss of interest in blank (viii) abnormal motor (ix)
(ix) Loss of psychomotor energy. activities; fatigue, loss • Fatigue (ix) behaviour (viii)
of energy, hypersomnia; • Negative
psychomotor retardation; symptoms;
loss of appetite (ix) catatonia (ix)

Figure 3.2  A tentative mapping of hypothesized types of antisuicide mechanisms (keepers) across common diagnostic categories of mental
THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

disorder
Notes: a The anxiousness of generalised anxiety disorder may be better understood as a concomitant of keepers rather than a keeper defence in itself, but the pain-type function of GAD
posited here might be taken to express the urgency of the organism’s need to seek relief from suicidogenic pain; bNon-suicidal self-injury (NSSI), classified as a ‘condition for further study’
in DSM-5 (APA, 2013), appears here as an exception, arguably lacking a ‘brain-type’ action: this function may be provided instead by other symptoms often comorbid with NSSI.
Source: Reproduced from Soper (2018).
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
progressing to suicidal actions. One commonal-
ity, mentioned above, is marked suicidality – to
be exact, suicidal ideas rather than the enact-
ing of those ideas (Nock et al., 2013). Another
is the characteristic stimulus-response course
of CMD, precipitated by emotionally painful
events, and typically lifting over time (with
or without medical intervention) after the pre-
cipitating adversities are eased (Brown, 2009;
Goldberg and Goodyer, 2005; Harris, 2000). Yet
others include: extreme comorbidity – people
diagnosed with one disorder usually also meet
criteria for one or more others, concurrently
or at different times (American Psychiatric
Association, 2013; First and Pincus, 2009);
lack of ‘zones of rarity’, or natural bounda-
ries, between diagnostic criteria (Kendell and
Jablensky, 2003); non-specificity of causation –
to large extent, as the keeper model anticipates,
any kind of adversity can precipitate any vari-
ety of CMD response (Goldberg and Goodyer,
2005; Kessler et al., 2010); non-specificity of
treatments – therapies developed for one condi-
tion also alleviate others (Wampold and Imel,
2015); common cognitive deficits – selectively
impairing, also as the keeper specification
predicts, the ability to organize complex tasks
(Harvey, 2004); coincident scheduling of first
onsets – in adolescence, as already noted, the
stage of life when suicidality first develops
(Kessler et al., 2007); species-specificity – an
absence of close animal models (Willner and
Belzung, 2015); and so on. This mesh of con-
tinuities poses a problem for an ad hoc label-
by-label style of analysis that is widespread in
Darwinian psychiatry (e.g., Brune, 2016; Del
Giudice, 2018) – some evolutionary hypotheses
advanced for depression, others for generalized
anxiety, yet others for schizophrenia etc. Soper
(2018) argues that any explanation proffered
for one diagnostic label is incomplete unless
also explained is the detailed configuration
that label shares, and its routine co-occurrence,
with almost any other form of CMD. Soper’s
model may offer parsimonious theoretical
underpinning for what some researchers have
inferred from empirical observation: that many
psychiatric constructs, for all their superficial
75
differences, vary more in degree than kind and
likely relate to a singular causal process, hith-
erto unclear (Caspi et  al., 2014; Menninger,
1963; Stochl et al., 2015).
Fourth, it is possible to understand why psy-
chiatric symptoms are poor predictors of sui-
cide, and indeed why a decades-long search
for any clinically useful marker has returned
empty handed (Carter et  al., 2017; Franklin
et al., 2017). The failure of prediction arises
not particularly, as is often claimed, because
suicide is rare; rare events are not necessar-
ily unpredictable (Soper, 2019a). Rather, pre-
cisely this null result would be expected of an
organism that is finely adapted to its ‘suicidal
niche’ and already making best use of avail-
able cues (Soper, 2019b). Suicide attempts,
as statistical residuals, are events associated
with no utilizable prognostic information
to which the individual’s defensive systems
could have responded. It should be antici-
pated, then, that suicides offer little or no
scope for being accurately foreseen at the
individual level.
Summing up, a posited convergence of
evidence from multiple directions sets up
a probability argument for special design
(Williams, 1966, 1996). Soper (2018) finds
no unaccountable inconsistencies, and no
better explanation for the confluence. A soft-
ware patch devised to stop humans switch-
ing themselves off in the face of difficulties
would predictably include routines that look
very much like symptoms of ‘functional’
common mental disorder. On this basis, he
intuits, it is more likely than not that many
outwardly dissimilar psychiatric states reflect
the proper workings of systems that evolved
to prevent self-killing.
Implications: The Case of
Depression
If the pain-and-brain framework is correct,
its ramifications may be wide-ranging, pro-
found, and take time to fathom. They could
impact on the research agenda, suicide
76 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
prevention policy, psychiatric practice, and the
conceptualization of psychopathology and
mental health. We will illustrate with an exam-
ple relating to the extensive and widely recom-
mended use of antidepressants and other
psychopharmacological treatments to prevent
suicides among clinical patients assessed to be
at risk (Wasserman et al., 2012; Zalsman et al.,
2017). The task analysis outlined above
suggests that certain common psychiatric
symptoms – including the emotional numbing,
cognitive impairment, fatigue, and generalized
anxiety characteristic of adult-pattern depres-
sive states – can be understood not as suicido-
genic disorder, but expressions of the
organism’s protective response to the lethal
threat latent in chronic, intense, emotional
pain. In other words, various common depres-
sive symptoms may be conceived as actions of
a psychological defensive system, in the same
way that modern medicine understands cough-
ing, vomiting, and fever to be defensive
responses (Nesse and Williams, 1995). It is for
this reason that the analysis predicts that, once
the triggering distress is relieved, depressive
symptoms would be expected to lift of
their own accord, with or without medical
intervention – spontaneous remission that
characterizes the normal course of most
common psychopathologies (Goldberg and
Goodyer, 2005; Harris, 2000). The notion of
protective antisuicide depression is not new.
Hendin (1975) inferred such a dynamic from
clinical observation half a century ago; and
some epidemiological findings might, arguably,
support Hendin’s case (Rogers et  al., 2018).
Psychiatrists across decades have warned that
suicide risk can intensify, not reduce, when
depressive symptoms start to lift (Kahne,
1966; Meehl, 1973). What is new is an evolu-
tionary foundation for the idea. Hence, evolu-
tionary analysis pertains to important questions
about the treatment of depression, in particular
the use of pharmacological treatments, espe-
cially in the context of suicide prevention. We
will highlight three issues.
First, the model suggests that antidepres-
sants treat symptoms rather than causes.
Addressing the root cause of depressive
symptoms would presumably involve helping
sufferers to relieve the precipitating distress,
which probably originates, recalling Gunn’s
(2017) Social Pain Model, from some form
of social detachment. That said, any pain
relief could offer provisional respite; and
given the likely social nature of the triggering
pain, any credible treatment, psychotropic
medication included, could be expected to
produce a strong placebo effect, signaling
that a supportive attachment has been put in
place (Davies, 2013; Wampold, 2018).
Second, at worst, suppressing the psyche’s
antisuicide systems exogenously, at least
without other measures, would be expected
to exacerbate, not lessen, suicide risk, at least
in some circumstances. This expectation is
supported by considerable pharmacologi-
cal evidence (Hengartner and Plöderl, 2019;
Sharma et al., 2016).
Third, evolutionary analysis challenges a
premise of such treatments, that people ‘at
risk’ can accurately be picked out in clinical
settings. As we have noted, special-purpose
biological defenses would be expected to
have exploited and exhausted any and all uti-
lizable markers, with the result that suicides
are probably not amenable to prediction in
principle (Soper, 2019a). The evolutionary
argument thus touches on an ethical ques-
tion: whether it is justifiable to prescribe
mind-altering drugs to large numbers of
people, nearly all of whom were never going
to take their own lives, in an effort to pro-
tect a small and unidentifiable minority. The
analysis would, in the round, appear to lend
theoretical support to those who already
question on empirical grounds whether
psychopharmacology is an appropriate
approach to suicide prevention (Hjelmeland
et al., 2019; Maris, 2015).
Looking at alternative strategies, the same
evolutionary analysis predicts that restrict-
ing access to lethal means (such as, for
example, installing nets under favored jump
sites, and limiting the size of retail drug
packs) may be effective to an extent that
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
would seem far-fetched to people viewing
such incomplete obstacles with the benefit of
normal intellectual functioning (Chen et  al.,
2016; Hawton, 2007). Interventions that only
mildly complicate a suicide endeavor would
be expected to disrupt disproportionately the
plans of someone affected by denials of psy-
chomotor energy, memory, decision-making,
and other task-related cognitive faculties –
attenuations which, Soper (2016, 2018) pos-
its, may be protective aspects of depression. A
slight delay may be enough to allow homeo-
static affective systems to de-escalate from a
suicidal crisis, restoring the organism towards
a protective, above-neutral, resting point.
Means restriction, in other words, could be
understood to capitalize on and co-operate
with the organism’s own antisuicide defenses.
Problems with Soper’s Theory
On one hand, Soper’s (2018) model has
merit. It appears, at least at this early stage,
parsimoniously to fit multiple lines of evi-
dence, and it does so arguably better than
other available explanations; it could be
judged attractive on the criterion of inference
to the best explanation (Harman, 1965). From
an evolutionary perspective, it appears to
match biological function to observable form
(Buss, 1995; Williams, 1996). The theory is
in principle falsifiable: many findings would
suffice as disproof – if science discovered, for
example, nonhuman suicide, or a human
population with a pattern of early childhood
suicides, or a group that lacked positive cor-
relations between suicidality and hypothe-
sized keeper responses, and so forth. It could
generate novel and testable ancillary predic-
tions. We propose, for example, that Soper’s
notion of a statistical, but only indirect, asso-
ciation between the scheduling of first onsets
of keepers (triggered by a hormonal cue
linked to puberty) and first onsets of suicide
(dependent on intellectual competence) could
be tested by comparing the epidemiology of
suicide and keepers in populations with
77
delayed/accelerated intellectual development
against populations with delayed/accelerated
endocrine development. A pattern in which
suicide, but not psychiatric symptoms,
occurred among the hormonally delayed but
cognitively precocious, and vice versa, could
be taken to support Soper’s proposals and be
difficult otherwise to explain.
On the other hand, Soper’s theory faces at
least four significant problems. First, being
at the stage of a preliminary sketch, it lacks
detail and raises many unanswered questions.
It is not specified, for example, how exactly
hypothesized antisuicide protections would
operate. To illustrate, ‘loss of psychomotor
energy’ (LoPE) (Soper, 2018: 143) is con-
ceived as an autonomic defense against sui-
cide, but self-killing could be brought about
by inaction as well as action. It is unclear
how LoPE would interact with other hypoth-
esized keepers; it could conflict, for example,
with others that presumably require positive
action, such as the compulsive consumption
of analgesics. The biological parameters of
LoPE are undefined – whether it exists as a
discrete condition, and how it would be oper-
ationalized as a testable construct, and so on.
Soper’s proposals may have intuitive appeal,
but there is much work to do before a robust,
comprehensive framework could be said
to have been achieved, despite, or perhaps
because of, the breadth of the framework’s
ambit.
Second, the forward/reverse engineering
approach, the appeal to evidence of special
design on which Soper’s adaptationist argu-
ments largely rely, is intrinsically intuitive
notwithstanding apparent objectivity (Lauder,
1996; Williams, 1996). Due to the method’s
subjectivity, another researcher replicating
Soper’s thought experiments might not arrive
at the same conclusions. This is not to say
that his arguments are invalid, but that there
is scope for different students legitimately to
reach different opinions from the same data
depending on their biographical backgrounds
(Kuhn, 1977). Soper’s efforts to infer a priori
design parameters for keepers, for example,
78 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
will not be entirely a priori: the author, a
psychotherapist, could not have approached
the topic with a blank slate and it would be
surprising if his findings were not colored by
experience. Indeed, some of his predictions
might strike many workers in mental health
as already matters of common knowledge.
Third, Soper’s (2018) approach leans on
a presumption of optimal outcomes: that
because x would be biologically ideal, x is
what we should find. While the optimization
approach is a useful way to model biological
problems and may suggest solutions, it is not
a safe basis in itself for predicting evolutionary
results. Many alternative outcomes of selection
are possible (Gould and Lewontin, 1979), and
the fitness functions of most adaptive problems
are complex landscapes of hills and valleys, the
path to optimization often blocked at local fit-
ness maxima (Parker and Smith, 1990).
Fourth, testing certain aspects of Soper’s
(2018) theory may be problematic due to the
composite nature of its hypotheses. Taking
again the example of ‘loss of psychomo-
tor energy’ (LoPE), this is presented as both
an adaptive solution to suicide risk, and as
an expression of that solution. Associations
between LoPE and measures of suicidality that
offer to support one hypothesis could simulta-
neously undermine the other: if, say, suicides
happen alongside LoPE, then that could be read
both as confirmatory evidence (because LoPE
had mobilized as predicted to meet the threat),
and as falsifying evidence (because LoPE had
failed in its supposed design task, to stop sui-
cides). This kind of epistemological trap is
not unique to Soper’s theorizing and may be
endemic in evolutionary science. A comparable
zero-sum game plays out, for example, in the
long-debated Westermarck effect, an evolved
mechanism said to deter close inbreeding: the
same evidence can be taken both to support and
contradict the theory (Sesardic, 2005).
On balance, while acknowledging these
and other weaknesses, it is not easy to dis-
miss the thrust of Soper’s (2018) arguments.
Intuitively difficult is the claim that varied
symptoms of psychiatric disorders function
as special-purpose antisuicide devices. But to
reject this hypothesis may be to invite fresh dif-
ficulty, because two questions would then call
for answers. Which aspects of the antisuicide
task analysis are being disputed? And, if not
‘functional’ psychiatric disorder, then what
alternative empirical phenomenon is proposed
that better matches that design specification? If
the pain-and-brain model is broadly correct, the
call would remain to explain why few people
kill themselves. It may be partly for this reason
that at least one prominent suicidologist is on
record as finding Soper’s proposals persuasive
(Lester, 2019). Time will tell if others agree.
CONCLUSION
At one level this chapter carries an encourag-
ing message. The evolutionary approach
would seem, in principle, capable of bringing
unity and coherence to suicidology’s current
morass of theory. A ‘pain-and-brain’ frame-
work in particular seems to offer a rallying
point for numerous, superficially disparate,
theoretical positions, such as IPTS (Van
Orden et  al., 2010), IMV (O’Connor et  al.,
2016), or SPM (Gunn, 2017), theories which
essentially characterize suicide as a way to
escape intolerable emotional stress. None
would appear incompatible with the view that
pain, as a biological imperative, motivates
action to end or escape it, while regular adult
human cognition offers intentional self-killing
as an effective, but genetically destructive,
means to answer that demand.
But the corollary, that suicide poses an engi-
neering problem, as summed by the tweet that
began this chapter, may be harder to digest.
Presumably human beings are prevented
most of the time from switching themselves
off because of one or more special-purpose
software patches. Blind to our own instincts
(Cosmides and Tooby, 1994), we may be
oblivious to their functioning, and skepti-
cal they may even exist. A research pro-
gram to uncover their workings may not be
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
easy to formulate. The idea that humans are
equipped with organismic defenses against
suicide is not new (e.g., Himmelhoch, 1988;
Hundert, 1992; Miller, 2008), but it has only
recently gained prominence in the research
agenda (Culotta, 2019). Its implications
may run wide and deep, and call some long-
held preconceptions into question: as Lester
(2019) finds, accepting the ramifications may
require close reading of the arguments.
Progress is not helped by what Soper (2018)
believes to be a systematic non-interaction
between suicidology and evolutionary psychol-
ogy, a two-way blockage of ideas that may go
beyond the institutional disconnects seen else-
where in psychological sciences (Staats, 2004).
Soper posits that suicide and evolution, each for
different reasons, are domains that many peo-
ple find awkward to think about, researchers
included. It may be for this reason that, in one
direction, evolutionary psychology has largely
ignored suicide, as indeed has psychology gen-
erally (Rogers, 2001): in view of the gravity
and ubiquity of suicide as a human phenom-
enon, and the evolutionary puzzle it presents,
remarkably little has been written on the subject
from an evolutionary perspective, at least until
recent years. In the other direction, suicidology
has largely ignored evolutionary psychology.
It may be illustrative that a rare review of the
field, titled ‘Evolutionary processes in suicide’
(Chiurliza et al., 2017), attempts to appraise its
research group’s ideas (and, oddly, only that
group’s) without reference to evolutionary psy-
chology’s primary texts or tenets – a surprising
omission given that evolutionary psychology,
‘the study of behavior from an evolutionary
perspective’ (Cornwell et  al., 2005: 369), is
centrally relevant.
This chapter calls for consilience between
the two fields. An evolutionary stance would
not in itself be a departure for suicidology:
it would, rather, follow the lead set by Freud
(1920/1991), Shneidman (1985), Joiner
(2005), and other prominent researchers,
drawing on evolutionary ideas across more
than a century. Evolutionary psychology could
synthesize, not replace, much of suicidology’s
79
existing theoretical and empirical content.
There may be little to lose in such an incre-
mental move. The upsides, on the other hand,
may be great. Evolutionary psychology offers
fresh perspectives for suicidology, and ready
tools that, if used, could be decisive in a bat-
tle to save lives. Evolutionary psychology and
suicidology deserve each other’s attention.
Notes
1  Reviews of prominent offers can be found in the
general suicidology literature (e.g., Gunn, 2019;
Gunn and Lester, 2014; O’Connor and Portzky,
2018; Paniagua et al., 2010; Selby et al., 2014).
2  If it were claimed that nonhuman suicide is rare
thanks to the action of antisuicide adaptations –
something like the ‘patch 7.822’ software update
imagined in this chapter’s opening quotation –
then those countermeasures need to be identi-
fied, as indeed we will later suggest they probably
need to be identified in humans.
3  As a side point: some theorists argue that risk of
suicide may vary, albeit weakly, with a heritable
propensity for certain personality traits, notably
impulsivity (McGirr et al., 2008).
4  Suicide would presumably have been no easier
in our evolutionary environment. As ground-
dwellers on open grasslands, it may have been
a scarcity of ready opportunities for self-killing
that allowed humans to encephalize closer to the
cognitive floor for suicide than would be feasible
for other social animals. Other social animals that
otherwise would be expected to benefit from
increased social intelligence (Varki and Brower,
2013) may occupy habitats where suicide could
be summarily enacted at almost any time by
default – by not gripping (chimp), or not surfac-
ing (dolphin), for example (Soper, 2019b).
5  As a related point, Humphrey (2018) speculates
that suicidality may help to account for cata-
strophic demographic collapses thought to have
occurred in early human pre-history.
REFERENCES
Aiello, L. C., & Wheeler, P. (1995). The
expensive-tissue hypothesis: The brain and
the digestive system in human and primate
evolution. Current Anthropology, 36(2),
199–221.
80 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Akotia, C. S., Knizek, B. L., Kinyanda, E., &
Hjelmeland, H. (2014). ‘I have sinned’:
Understanding the role of religion in the
experiences of suicide attempters in Ghana.
Mental Health, Religion & Culture, 17(5),
437–448.
Alexander, R. D. (1974). The evolution of social
behavior. Annual Review of Ecology and
Systematics, 5(1), 325–383.
American Association of Suicidology. (2019).
History of the American Association of
Suicidology. Retrieved from https://
suicidology.org/about-aas/#history (Accessed
20 May 2020)
American Psychiatric Association. (2013).
Diagnostic And Statistical Manual Of Mental
Disorders: DSM-5. New York, NY: American
Psychiatric Association.
Andoh-Arthur, J., Knizek, B. L., Osafo, J., &
Hjelmeland, H. (2020). Societal reactions to
suicide in Ghana: A qualitative study of
experiences of the bereaved. Crisis, 41(2),
128–134.
Anil, M., Preston, J., McKinstry, J., Rodway, R.,
& Brown, S. (1996). An assessment of stress
caused in sheep by watching slaughter of
other sheep. Animal Welfare, 5(4),
435–441.
Atkinson, J. M. (1978). Discovering Suicide:
Studies in the Social Organization of Sudden
Death. London, UK: McMillan.
Aubin, H.-J., Berlin, I., & Kornreich, C. (2013).
The evolutionary puzzle of suicide. Interna-
tional Journal of Environmental Research &
Public Health, 10(12), 6873–6886.
Auvray, M., Myin, E., & Spence, C. (2010). The
sensory-discriminative and affective-
motivational aspects of pain. Neuroscience &
Biobehavioral Reviews, 34(2), 214–223.
Baechler, J. (1975/1979). Les Suicides (B.
Cooper, Trans.). New York, NY: Basic Books.
Bailey, J. D. (2009). Orphan care: An introduction.
Social Work & Society, 7(1), 1–12.
Bancroft, J., Hawton, K., Simkin, S., Kingston,
B., Cumming, C., & Whitwell, D. (1979). The
reasons people give for taking overdoses: A
further inquiry. British Journal of Medical
Psychology, 52(4), 353–365.
Bateson, M., Brilot, B., & Nettle, D. (2011).
Anxiety: an evolutionary approach. The
Canadian Journal of Psychiatry, 56(12),
707–714.
Baumeister, R. F. (1990). Suicide as escape from
self. Psychological Review, 97(1), 90–113.
Baumeister, R. F., & Leary, M. R. (1995). The need
to belong: Desire for interpersonal attach-
ments as a fundamental human motivation.
Psychological Bulletin, 117(3), 497.
Baumeister, R. F. (1989). The optimal margin of
illusion. Journal of Social and Clinical
Psychology, 8(2), 176–189.
Benatar, D. (2015). Life is not good. In T. K.
Shackelford & R. D. Hansen (Eds.), The
Evolution Of Morality (pp. 137–140). Cham,
Switzerland: Springer.
Bering, J. M. (2018). A Very Human Ending:
How Suicide Haunts Our Species. London,
UK: Transworld.
Bering, J. M., & Shackelford, T. K. (2004). The
causal role of consciousness: A conceptual
addendum to human evolutionary psychology.
Review of General Psychology, 8(4), 227–248.
Bjorklund, D. F., Causey, K., & Periss, V. (2010).
The evolution and development of human
social cognition. In P. M. Kappeler & J. B. Silk
(Eds.), Mind The Gap (pp. 351–371).
Heidelberg, Germany: Springer.
Blanchard, D. C., Griebel, G., & Nutt, D. J.
(2011). Handbook Of Anxiety And Fear (Vol.
17). Amsterdam, Netherlands: Academic.
Blanchard, D. C., Griebel, G., Pobbe, R., &
Blanchard, R. J. (2011). Risk assessment as
an evolved threat detection and analysis
process. Neuroscience & Biobehavioral
Reviews, 35(4), 991–998.
Blasco-Fontecilla, H., Lopez-Castroman, J.,
Gomez-Carrillo, A., & Baca-Garcia, E. (2009).
Towards an evolutionary framework of
suicidal behavior. Medical Hypotheses, 73(6),
1078–1079.
Bohannan, P. (Ed.) (1960). African Homicide
And Suicide. Princeton, NJ: Princeton
University Press.
Bolton, J. M., Au, W., Leslie, W. D., Martens.,
Patricia J., Enns, M. W., Roos, L L., & Sareen, J.
(2013). Parents bereaved by offspring suicide:
A population-based longitudinal case-control
study. JAMA Psychiatry, 70(2), 158–167.
Bowlby, J. (1969/1997). Attachment And Loss:
Attachment (2nd ed. Vol. 1). London, UK:
Pimlico.
Bowlby, J. (1973). Attachment And Loss:
Separation: Anxiety And Anger (2nd ed.
Vol. 2). London, UK: Penguin.
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
Bowlby, J. (1980/1991). Attachment And Loss:
Loss: Sadness And Depression (Vol. 3).
London, UK: Penguin.
Bracke, M. (1992). Can animals have a
preference not to be killed?: Observations on
red deer, laying hens and mice which were
exposed to conspecifics being killed: A study
in cognitive ethology (Doctoral dissertation).
Edinburgh School of Agriculture, Edinburgh,
UK.
Brand, P. W., & Yancey, P. (1993). Pain: The Gift
Nobody Wants. New York, NY: HarperCollins.
Brezo, J., Paris, J., Barker, E. D., Tremblay, R.,
Vitaro, F., Zoccolillo, M., & Turecki, G. (2007).
Natural history of suicidal behaviors in a
population-based sample of young adults.
Psychological Medicine, 37(11),
1563–1574.
Bribiescas, R. G. (2006). On the evolution, life
history, and proximate mechanisms of
human male reproductive senescence.
Evolutionary Anthropology: Issues, News,
and Reviews, 15(4), 132–141.
Brown, D. E. (2004). Human universals, human
nature & human culture. Daedalus, 133(4),
47–54.
Brown, G. W. (2009). Medical sociology and
issues of aetiology. In M. G. Gelder, N. C.
Andreasen, J. J. López-Ibor, & J. R. Geddes
(Eds.), New Oxford Textbook Of Psychiatry
(pp. 268–275). Oxford, UK: Oxford University
Press.
Brown, M. F. (1986). Power, gender, and the
social meaning of Aguaruna suicide. Man,
21(2), 311–328.
Brown, R. M., Brown, S. L., Johnson, A., Olsen,
B., Melver, K., & Sullivan, M. (2009). Empirical
support for an evolutionary model of self-
destructive motivation. Suicide and Life-
Threatening Behavior, 39(1), 1–12.
Brown, R. M., Dahlen, E., Mills, C., Rick, J., &
Biblarz, A. (1999). Evaluation of an
evolutionary model of self-preservation and
self-destruction. Suicide and Life-Threatening
Behavior, 29(1), 58–71.
Brüne, M. (2016). Textbook of Evolutionary
Psychiatry and Psychosomatic Medicine: The
Origins of Psychopathology (2nd ed.).
Oxford, UK: Oxford University Press.
Buss, D. M. (1990). Toward a biologically
informed psychology of personality. Journal
of Personality, 58(1), 1–16.
81
Buss, D. M. (1995). Evolutionary psychology: A
new paradigm for psychological science.
Psychological Inquiry, 6(1), 1–30.
Buss, D. M., & Penke, L. (2015). Evolutionary
personality psychology. In M. Mikulincer, P.
R. Shaver, M. Cooper, Lynne, & R. J. Larsen
(Eds.), APA Handbook Of Personality And
Social Psychology: Vol. 4. Personality
Processes And Individual Differences (pp. 3–
39). Washington, DC: American Psychological
Association.
Campbell, A. (2002). A Mind Of Her Own: The
Evolutionary Psychology Of Women. New
York, NY: Oxford University Press.
Carter, G., Milner, A., McGill, K., Pirkis, J.,
Kapur, N., & Spittal, M. J. (2017). Predicting
suicidal behaviours using clinical instruments:
Systematic review and meta-analysis of
positive predictive values for risk scales. The
British Journal of Psychiatry, 210(6),
387–395.
Caspi, A., Houts, R. M., Belsky, D. W., Goldman-
Mellor, S. J., Harrington, H., Israel, S., &
Moffitt, T. E. (2014). The p factor: One
general psychopathology factor in the
structure of psychiatric disorders? Clinical
Psychological Science, 2(2), 119–137.
Cavanagh, J. T. O., Carson, A. J., Sharpe, M., &
Lawrie, S. M. (2003). Psychological autopsy
studies of suicide: A systematic review.
Psychological Medicine, 33(03), 395–405.
Cerel, J., & Aldrich, R. S. (2011). The impact of
suicide on children and adolescents. In J. R.
Jordan & J. L. McIntosh (Eds.), Grief After
Suicide: Understanding The Consequences
And Caring For The Survivors (pp. 81–92).
New York, NY: Routledge.
Cerel, J., Brown, M. M., Maple, M., Singleton,
M., van de Venne, J., Moore, M., & Flaherty,
C. (2019). How many people are exposed to
suicide? Not six. Suicide and Life-Threatening
Behavior, 49(2), 529–534.
Chapple, A., Ziebland, S., & Hawton, K. (2015).
Taboo and the different death? Perceptions of
those bereaved by suicide or other traumatic
death. Sociology of Health & Illness, 37(4),
610–625.
Chen, Y.-Y., Chien-Chang Wu, K., Wang, Y., &
Yip, P. (2016). Suicide prevention through
restricting access to suicide means and
hotspots. In R. C. O’Connor & J. Pirkis (Eds.),
International Handbook Of Suicide
82 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Prevention (2nd ed., pp. 609–636).
Chichester, UK: John Wiley and Sons.
Chiurliza, B., Rogers, M. L., Schneider, M. E.,
Chu, C., & Joiner, T. E. (2017). Evolutionary
processes in suicide. Current Opinion in
Psychology, 22, 84–88.
Cholbi, M. (2017). Suicide. The Stanford
Encyclopedia of Philosophy. Fall 2017.
Retrieved from https://plato.stanford.edu/
archives/fall2017/entries/suicide/ (Accessed
20 May 2020)
Chow, A. Y. M. (2006). The day after:
Experiences of bereaved suicide survivors. In
C. L. W. C. Chan & A. Y. M. Chow (Eds.),
Death, Dying And Bereavement: A Hong Kong
Chinese Experience (Vol. 1, pp. 293–308).
Aberdeen, Hong Kong: Hong Kong University
Press.
Chu, C., Buchman-Schmitt, J. M., Stanley, I. H.,
Hom, M. A., Tucker, R. P., Hagan, C. R., &
Ringer, F. B. (2017). The interpersonal theory
of suicide: A systematic review and meta-
analysis of a decade of cross-national research.
Psychological Bulletin, 143(12), 1313–1345.
Chung, D., Hadzi-Pavlovic, D., Wang, M.,
Swaraj, S., Olfson, M., & Large, M. (2019).
Meta-analysis of suicide rates in the first
week and the first month after psychiatric
hospitalisation. BMJ open, 9(3), e023883.
Cole, L. C. (1954). The population consequences
of life history phenomena. The Quarterly
Review of Biology, 29(2), 103–137.
Comai, S., & Gobbi, G. (2016). Translational
research in suicide: Is it possible to study
suicide in animal models? In P. Courtet (Ed.),
Understanding Suicide: From Diagnosis To
Personalized Treatment (pp. 177–188).
Cham, Switzerland: Springer.
Confer, J. C., Easton, J. A., Fleischman, D. S.,
Goetz, C. D., Lewis, D. M., Perilloux, C., &
Buss, D. M. (2010). Evolutionary psychology.
Controversies, questions, prospects, and
limitations. American Psychologist, 65(2),
110–126.
Corns, J. (2014). Unpleasantness, motivational
oomph, and painfulness. Mind & Language,
29(2), 238–254.
Cornwell, R. E., Palmer, C., Guinther, P. M., &
Davis, H. P. (2005). Introductory psychology
texts as a view of sociobiology/evolutionary
psychology’s role in psychology. Evolutionary
Psychology, 3(1), 147470490500300124.
Cosmides, L., & Tooby, J. (1994). Beyond
intuition and instinct blindness: Toward an
evolutionarily rigorous cognitive science.
Cognition, 50(1), 41–77.
Cosmides, L., & Tooby, J. (2000). Evolutionary
psychology and the emotions. In M. Lewis &
J. M. Haviland-Jones (Eds.), Handbook Of
Emotions (Vol. 2, pp. 91–115). New York,
NY: Guilford Press.
Culotta, E. (2019). Probing an evolutionary
riddle. Science, 365(6455), 748–749.
Daly, M., & Wilson, M. (1988). Homicide. New
Brunswick, NJ: Transaction.
Darwin, C. (1859). On The Origin Of Species
By Means Of Natural Selection, Or The
Preservation Of Favoured Races In The
Struggle For Life. London, UK: John Murray.
Davies, J. (2013). Cracked: Why Psychiatry Is
Doing More Harm Than Good. London, UK:
Icon.
Dawkins, R. (1976). The Selfish Gene (3rd ed.).
Oxford, UK: Oxford University Press.
Dawkins, R. (1980). Domesticity, senescence,
and suicide. Behavioral and Brain Sciences,
3(2), 274.
De Leo, D., Milner, A., Fleischmann, A.,
Bertolote, J., Collings, S., Amadeo, S., &
Saniel, B. (2013). The WHO START Study:
Suicidal behaviors across different areas of
the world. Crisis, 34(3), 156–163.
deCatanzaro, D. (1980). Human suicide: A
biological perspective. Behavioral and Brain
Sciences, 3(02), 265–272.
deCatanzaro, D. (1981). Suicide And Self-
Damaging Behavior: A Sociobiological
Perspective. New York, NY: Academic Press.
deCatanzaro, D. (1982). Suicide, aggression,
and natural selection. Behavioral and Brain
Sciences, 5(02), 316–317.
deCatanzaro, D. (1986). A mathematical model
of evolutionary pressures regulating self-
preservation and self-destruction. Suicide
and Life-Threatening Behavior, 16(2),
166–181.
deCatanzaro, D. (1991). Evolutionary limits to
self-preservation. Ethology and Sociobiology,
12(1), 13–28.
deCatanzaro, D. (1992). Prediction of self-
preservation failures on the basis of
quantitative evolutionary biology. In R. W.
Maris, A. L. Berman, J. T. Maltsberger, & R. I.
Yufit (Eds.), Assessment And Prediction Of
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
Suicide (pp. 607–624). New York, NY:
Guilford Press.
Delisle, J. R. (1986). Death with honors: Suicide
among gifted adolescents. Journal of
Counseling and Development, 64(9),
558–560.
Del Giudice, M. (2018). Evolutionary
psychopathology: A unified approach. New
York, NY: Oxford University Press.
Dickeman, M. (1975). Demographic conse-
quences of infanticide in man. Annual Review
of Ecology and Systematics, 6(1), 107–137.
D. Lester (Ed.), Theories Of Suicide: Past,
Present and Future (pp. 9–22). Springfield,
IL: Charles C. Thomas.
Duntley, J. D. (2005). Adaptations to dangers
from humans. In D. M. Buss (Ed.), The
Handbook Of Evolutionary Psychology
(pp. 224–249). Hoboken, NJ: John Wiley &
Sons.
Duntley, J. D., & Buss, D. M. (2004). The
evolution of evil. In A. G. Miller (Ed.), The
Social Psychology Of Good And Evil (pp.
102–124). New York, NY: Guilford Press.
Durkheim, E. (1897/1952). La Suicide (J.
Spaulding & G. Simpson, Trans.). Henley, UK:
Routledge.
Dyregrov, K., Plyhn, E., & Dieserud, G. (2012).
After The Suicide: Helping The Bereaved To
Find A Path From Grief To Recovery. London,
UK: Jessica Kingsley.
Eccleston, C. (2001). Role of psychology in pain
management. British Journal of Anaesthesia,
87(1), 144–152.
Efklides, A., & Moraitou, D. (Eds.). (2013). A
Positive Psychology Perspective On Quality
Of Life. New York, NY: Springer.
Eisenberger, N. I., & Lieberman, M. D. (2004).
Why rejection hurts: A common neural alarm
system for physical and social pain. Trends in
Cognitive Sciences, 8(7), 294–300.
Eisenberger, N. I., & Lieberman, M. D. (2005).
Why it hurts to be left out: The neurocognitive
overlap between physical and social pain. In
K. D. Williams, J. P. Forgas, & W. von Hippel
(Eds.), The Social Outcast: Ostracism, Social
Exclusion, Rejection, And Bullying (pp. 109–
127). New York, NY: Psychology Press.
Eisenberger, N. I., Lieberman, M. D., & Williams,
K. D. (2003). Does rejection hurt? An fMRI
study of social exclusion. Science, 302(5643),
290–292.
83
Engel, C. (2002). Wild Health. Boston, MA:
Houghton Mifflin Harcourt.
Erlangsen, A., Runeson, B., Bolton, J. M.,
Wilcox, H. C., Forman, J. L., Krogh, J., &
Conwell, Y. (2017). Association between
spousal suicide and mental, physical, and
social health outcomes: A longitudinal and
nationwide register-based study. JAMA
Psychiatry, 74(5), 456–464.
Falger, V. S. E., & Falger, E. L. F. (2003). The
cultural evolution of dying: Euthanasia in the
Netherlands. In A. Somit & S. A. Peterson
(Eds.), Human Nature And Public Policy: An
Evolutionary Approach (pp. 77–96). New
York, NY: Palgrave Macmillan.
Falk, D. (1990). Brain evolution in Homo: The
‘radiator’ theory. Behavioral and Brain
Sciences, 13(02), 333–344.
Faucher, L. (2016). Darwinian blues:
Evolutionary psychiatry and depression. In J.
C. Wakefield & S. Demazeux (Eds.), Sadness
Or Depression? (pp. 69–94). Dordrecht,
Netherlands: Springer.
Fedden, R. (1938). Suicide; A Social And
Historical Study. London, UK: Peter Davies.
First, M. B., & Pincus, H. A. (2009). Diagnosis and
classification. In M. G. Gelder, N. C. Andreasen,
J. J. López-Ibor, & J. R. Geddes (Eds.), New
Oxford Textbook Of Psychiatry (pp. 99–121).
Oxford, UK: Oxford University Press.
Flinn, M. V., & Alexander, R. D. (2007). Runaway
social selection in human evolution. In S. W.
Gangestad & J. A. Simpson (Eds.), The
Evolution Of Mind: Fundamental Questions
And Controversies (pp. 249–255). New York,
NY: Guilford Press.
Flinn, M. V., Quinlan, R. J., Ward, C., & Coe, M.
(2007). Evolution of the human family: Coop-
erative males, long social childhoods, smart
mothers, and extended kin networks. In C.
Salmon & T. K. Shackelford (Eds.), Family Rela-
tionships: An Evolutionary Perspective
(pp. 16–38). Oxford, UK: Oxford University Press.
Franklin, J. C., Ribeiro, J. D., Fox, K. R., Bentley,
K. H., Kleiman, E. M., Huang, X., & Nock, M.
K. (2017). Risk factors for suicidal thoughts
and behaviors: A meta-analysis of 50 years
of research. Psychological Bulletin, 143(2),
187–232.
Freud, S. (1920/1991). Beyond The Pleasure
Principle (J. Strachey, Trans.). London, UK:
Penguin.
84 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Frey, L. M., Hans, J. D., & Cerel, J. (2015).
Perceptions of suicide stigma: How do social
networks and treatment providers compare?
Crisis, 37(2), 95–103.
Gallup, G. G., & Weedon, S. L. (2013). Suicide
bombers: Does an evolutionary perspective
make a difference? Evolutionary Psychology,
11(4), 791–794.
Gandhi, S. G., Gilbert, W. M., McElvy, S. S., El
Kady, D., Danielson, B., Xing, G., & Smith, L.
H. (2006). Maternal and neonatal outcomes
after attempted suicide. Obstetrics &
Gynecology, 107(5), 984–990.
Geary, D. C. (2005). Evolution of paternal
investment. In D. M. Buss (Ed.), The
Handbook Of Evolutionary Psychology (pp.
483–505). Hoboken, NJ: Wiley.
Geary, D. C. (2007). The motivation to control
and the evolution of general intelligence. In
S. W. Gangestad & J. A. Simpson (Eds.), The
Evolution Of Mind: Fundamental Questions
And Controversies (pp. 305–312). New York,
NY: Guilford Press.
Gilbert, P. (1989). Human Nature And Suffering.
Hove, UK: Lawrence Erlbaum Associates.
Goldberg, D., & Goodyer, I. (2005). The Origins
And Course Of Common Mental Disorders.
Hove, UK: Routledge.
Goldblatt, M. J. (2014). Psychodynamics of
suicide. In M. K. Nock (Ed.), The Oxford
Handbook Of Suicide And Self-Injury
(pp. 255–264). Oxford, UK: OUP.
Goldsmith, S., Pellmar, T., Kleinman, A., &
Bunney, W. (2002). Reducing Suicide: A
National Imperative. Washington, DC:
National Academies Press.
Gorelik, G., & Shackelford, T. K. (2017). Suicide
and the moralistic fallacy: Comment on
Joiner, Hom, Hagan, and Silva (2016).
Evolutionary Psychological Science, 3(3),
287–289.
Gould, S. J., & Lewontin, R. C. (1979). The
spandrels of San Marco and the Panglossian
paradigm: A critique of the adaptationist
programme. Proceedings of the Royal Society
of London B: Biological Sciences, 205(1161),
581–598.
Grad, O., & Andriessen, K. (2016). Surviving
the legacy of suicide. In R. C. O’Connor & J.
Pirkis (Eds.), The International Handbook Of
Suicide Prevention (pp. 663–680). Chichester,
UK: Wiley.
Gunn, J. F. (2014). Suicide as escape: Baechler,
Shneidman, and Baumeister. In J. F. Gunn &
Gunn, J. F. (2017). The Social Pain Model.
Crisis, 38(5), 281–286.
Gunn, J. F. (2019). The importance of
conceptual and theoretical frameworks. In D.
Lester (Ed.), The End Of Suicidology: Can We
Ever Understand Suicide? (pp. 135–145).
New York, NY: Nova Science Publishers.
Gunn, J. F., & Lester, D. (2014). Theories Of
Suicide: Past, Present And Future. Springfield,
IL: Charles C. Thomas.
Hagen, E. H. (2002). Depression as bargaining:
The case postpartum. Evolution and Human
Behavior, 23(5), 323–336.
Hamilton, W. D. (1964). The genetical evolution
of social behaviour. I. Journal of Theoretical
Biology, 7(1), 1–16.
Hamilton, W. D. (1972). Altruism and related
phenomena, mainly in social insects. Annual
Review of Ecology and Systematics, 3(1),
193–232.
Hamilton, W. J. (1980). Do nonhuman animals
commit suicide? Behavioral and Brain
Sciences, 3(02), 278–279.
Hanschmidt, F., Lehnig, F., Riedel-Heller, S. G.,
& Kersting, A. (2016). The stigma of suicide
survivorship and related consequences –
a systematic review. PloS One, 11(9),
e0162688.
Harman, G. H. (1965). Inference to the best
explanation. Philosophical Review, 74,
88–95.
Harris, E. C., & Barraclough, B. (1997). Suicide
as an outcome for mental disorders. A meta-
analysis. The British Journal of Psychiatry,
170(3), 205–228.
Harris, M. (1974). Cows, Pigs, Wars And
Witches: The Riddles Of Culture. New York,
NY: Random House.
Harris, T. O. (2000). Introduction to the work of
George Brown. In T. O. Harris (Ed.), Where
Inner And Outer Worlds Meet: Psychosocial
Research In The Tradition Of George W
Brown (pp. 1–52). London, UK: Routledge.
Harvey, A. G. (2004). Cognitive Behavioural
Processes Across Psychological Disorders: A
Transdiagnostic Approach To Research And
Treatment. Oxford, UK: Oxford University Press.
Hausfater, G., & Hrdy, S. B. (1984). Infanticide:
Comparative And Evolutionary Perspectives.
New York, NY: Aldine.
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
Hawton, K. (2007). Restricting access to
methods of suicide: Rationale and evaluation
of this approach to suicide prevention. Crisis,
28(S1), 4–9.
Healey, C. (1979). Women and suicide in New
Guinea. Social Analysis: The International
Journal of Social and Cultural Practice, (2),
89–106.
Hedegaard, H., Curtin, S. C., & Warner, M.
(2018). Suicide mortality in the United States,
1999–2017. NCHS Data Brief, 330, 1–8.
Heintzelman, S. J., & King, L. A. (2014). Life is
pretty meaningful. American Psychologist,
69(6), 561–574.
Hendin, H. (1975). Growing up dead: Student
suicide. American Journal of Psychotherapy,
29(3), 327–338.
Hengartner, M. P., & Plöderl, M. (2019). Reply to
the letter to the editor: ‘Newer-generation
antidepressants and suicide risk: Thoughts on
Hengartner and Plöderl’s Re-Analysis’. Psycho-
therapy and Psychosomatics. 88(6), 373–374.
Hezel, F. X., Rubinstein, D. H., & White, G. M.
(Eds.). (1985). Culture, Youth And Suicide In
The Pacific. Honolulu, HI: Pacific Island
Studies Program.
Himmelhoch, J. M. (1988). What destroys our
restraints against suicide? Journal of Clinical
Psychiatry, 49(9 (Suppl)), 46–52.
Hjelmeland, H., Jaworski, K., Knizek, B. L., &
Marsh, I. (2019). Problematic advice from
suicide prevention experts. Ethical Human
Psychology and Psychiatry, 20(2), 79–85.
Hjelmeland, H., & Knizek, B. L. (2019). The
emperor’s new clothes? A critical look at the
interpersonal theory of suicide. Death
Studies, 44(4), 168–178.
Hrdy, S. B. (1979). Infanticide among animals:
A review, classification, and examination of
the implications for the reproductive
strategies of females. Ethology and
Sociobiology, 1(1), 13–40.
Huber, F. (2015/2019). Promise Me You’ll Shoot
Yourself (I. Taylor, Trans.). London, UK: Allen
Lane.
Humphrey, N. (1976). The social function of
intellect. In P. Bateson & R. A. Hinde (Eds.),
Growing Points In Ethology (pp. 303–317).
Cambridge, UK: Cambridge University Press.
Humphrey, N. (2011). Soul Dust: The Magic Of
Consciousness. Princeton, NJ: Princeton
University Press.
85
Humphrey, N. (2018). The lure of death:
Suicide and human evolution. Philosophical
Transactions of the Royal Society, B, 373, 1–8.
Hundert, E. M. (1992). The brain’s capacity to
form delusions as an evolutionary strategy
for survival. In M. Spitzer, F. Uehlein, M. A.
Schwartz, & C. Mundt (Eds.), Phenomenology,
Language & Schizophrenia (pp. 346–354).
New York, NY: Springer.
Johnson, B. D. (1965). Durkheim’s one cause of
suicide. American Sociological Review, 30,
875–886.
Joiner, T. E. (2005). Why People Die By Suicide.
Cambridge, MA: Harvard University Press.
Joiner, T. E., Buchman-Schmitt, J. M., Chu, C.,
& Hom, M. A. (2017). A sociobiological
extension of the interpersonal theory of
suicide. Crisis, 38(2), 69–72.
Joiner, T. E., Hom, M. A., Hagan, C. R., & Silva,
C. (2016). Suicide as a derangement of the
self-sacrificial aspect of eusociality.
Psychological Review, 123(3), 235–254.
Jordan, J. R. (2008). Bereavement after suicide.
Psychiatric Annals, 38(10), 679–685.
Jordan, J. R., & McIntosh, J. L. (Eds.). (2011).
Grief After Suicide: Understanding The
Consequences And Caring For The Survivors.
New York, NY: Routledge.
Kahne, M. J. (1966). Suicide research: A critical
review of strategies and potentialities in
mental hospitals (Part II). International
Journal of Social Psychiatry, 12(3), 177–186.
Kappeler, P. M., Silk, J. S., Burkart, J. M., & van
Schaik, C. P. (2010). Primate behavior and
human universals: Exploring the gap. In P. M.
Kappeler & J. B. Silk (Eds.), Mind The Gap
(pp. 3–15). Berlin, Germany: Springer.
Kapur, N., Cooper, J., O’Connor, R. C., &
Hawton, K. (2013). Non-suicidal self-injury v.
attempted suicide: New diagnosis or false
dichotomy? The British Journal of Psychiatry,
202(5), 326–328.
Kastenbaum, R. (1967). The child’s understanding
of death: How does it develop? In E. A.
Grollman (Ed.), Explaining Death To Children
(pp. 89–100). Boston, MA: Beacon.
Keller, M. C., & Miller, G. F. (2006). Resolving
the paradox of common, harmful, heritable
mental disorders: Which evolutionary genetic
models work best? Behavioral and Brain
Sciences, 29(4), 385–404; discussion
405–352.
86 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Kendell, R., & Jablensky, A. (2003).
Distinguishing between the validity and
utility of psychiatric diagnoses. American
Journal of Psychiatry, 160(1), 4–12.
Kennedy, P., Rogers, B., Speer, S., & Frankel, H.
(1999). Spinal cord injuries and attempted
suicide: A retrospective review. Spinal Cord,
37(12), 847–852.
Kessler, R. C., Amminger, G. P., Aguilar-Gaxiola,
S., Alonso, J., Lee, S., & Ustun, T. B. (2007).
Age of onset of mental disorders: A review
of recent literature. Current Opinion in
Psychiatry, 20(4), 359.
Kessler, R. C., McLaughlin, K. A., Green, J. G.,
Gruber, M. J., Sampson, N. A., Zaslavsky,
A. M., & Angermeyer, M. (2010). Childhood
adversities and adult psychopathology in
the WHO World Mental Health Surveys. The
British Journal of Psychiatry, 197(5),
378–385.
Kirkpatrick, L. A., & Navarrete, C. D. (2006).
Reports of my death anxiety have been
greatly exaggerated: A critique of terror
management theory from an evolutionary
perspective. Psychological Inquiry, 17(4),
288–298.
Klein, R. G., & Edgar, B. (2002). The Dawn Of
Human Culture. New York, NY: Wiley.
Klonsky, E. D., Dhingra, K., Klonsky, D., &
Tapola, V. (2019). An empirical test of the
Three-Step Theory (3ST) of suicide in UK
university students. Suicide and Life-
Threatening Behavior, 49, 478–487.
Klonsky, E. D., & May, A. M. (2015). The Three-
Step Theory (3ST): A new theory of suicide
rooted in the ‘ideation-to-action’ framework.
International Journal of Cognitive Therapy,
8(2), 114–129.
Klonsky, E. D., May, A. M., & Saffer, B. Y.
(2016). Suicide, suicide attempts, and
suicidal ideation. Annual Review of Clinical
Psychology, 12, 307–330.
Knizek, B. L., Kinyanda, E., Akotia, C. S., &
Hjelmeland, H. (2013). Between Hippocrates
and God: Ugandan mental health
professional’s views on suicide. Mental
Health, Religion & Culture, 16(8), 767–780.
Kuhl, J., Quirin, M., & Koole, S. L. (2015). Being
someone: The integrated self as a
neuropsychological system. Social and
Personality Psychology Compass, 9(3),
115–132.
Kuhn, T. S. (1977). The Essential Tension —
Selected Studies In Scientific Tradition And
Change. Chicago, IL: University of Chicago
Press.
Lankford, A. (2013). The Myth Of Martyrdom.
New York, NY: Palgrave.
Lankford, A. (2015). Is suicide terrorism really
the product of an evolved sacrificial
tendency? A review of mammalian research
and application of evolutionary theory.
Comprehensive Psychology, 4(21).
Lauder, G. V. (1996). The argument from
design. In M. R. Rose & G. V. Lauder (Eds.),
Adaptation (pp. 55–87). San Diego, CA:
Academic Press.
Layard, R. (2011). Happiness: Lessons From A
New Science (2nd ed.). London, UK: Penguin.
Leary, M. R., & Guadagno, J. (2011). The
sociometer, self-esteem, and the regulation
of interpersonal behavior. In K. D. Vohs &
R. F. Baumeister (Eds.), Handbook Of Self-
Regulation: Research, Theory, And
Applications (2nd ed., pp. 339–354). New
York, NY: Guilford Press.
Leary, M. R., Tambor, E. S., Terdal, S. K., &
Downs, D. L. (1995). Self-esteem as an
interpersonal monitor: The sociometer
hypothesis. Journal of Personality and Social
Psychology, 68(3), 518.
Lester, D. (1993). The stigma against dying and
suicidal patients: A replication of Richard
Kalish’s study twenty-five years later. OMEGA
Journal of Death and Dying, 26(1), 71–75.
Lester, D. (2014a). Suicide as deviance. In J. F.
Gunn & D. Lester (Eds.), Theories Of Suicide:
Past, Present And Future (pp. 244–277).
Springfield, IL: Charles C. Thomas.
Lester, D. (2014b). Suicide, ethology and
sociobiology. In J. F. Gunn & D. Lester (Eds.),
Theories Of Suicide: Past, Present And Future
(pp. 55–71). Springfield, IL: Charles C.
Thomas.
Lester, D. (2017). Non-human animal suicide
could be tested. Animal Sentience, 2(20), 3.
Lester, D. (2019). The End Of Suicidology: Can
We Ever Understand Suicide? New York, NY:
Nova Science Publishers.
Lieberman, M. D. (2013). Social: Why Our
Brains Are Wired To Connect. Oxford, UK:
OUP.
Litman, R. E. (1967). Sigmund Freud on suicide.
In E. Shneidman (Ed.), Essays In
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
Self-Destruction (pp. 324–344). New York,
NY: Aronson.
Linehan, M. M. (2020). Building a Life Worth
Living. New York, NY: Random House.
Macdonald, C. J.-H. (2007). Uncultural
Behavior: An Anthropological Investigation
Of Suicide In The Southern Philippines.
Honolulu, HI: University of Hawai’i Press.
MacDonald, G., & Leary, M. R. (2005). Why
does social exclusion hurt? The relationship
between social and physical pain.
Psychological Bulletin, 131(2), 202.
Malkesman, O., Pine, D. S., Tragon, T., Austin,
D. R., Henter, I. D., Chen, G., & Manji, H. K.
(2009). Animal models of suicide-trait-
related behaviors. Trends in Pharmacological
Sciences, 30(4), 165–173.
Maltsberger, J. T. (2003). Can a louse commit
suicide? Crisis, 24(4), 175–176.
Maple, M., Frey, L. M., McKay, K., Coker, S., &
Grey, S. (2019). “Nobody hears a silent cry for
help”: Suicide attempt survivors’ experiences
of disclosing during and after a crisis. Archives
of Suicide Research, 1–19.
Maris, R. W. (1982). Rational suicide: An
impoverished self-transformation. Suicide
and Life-Threatening Behavior, 12(1), 4–16.
Maris, R. W. (2015). Pillaged: Psychiatric
Medications And Suicide Risk. Columbia, SC:
University of South Carolina Press.
Marušič, A., & Swapp, R. (2004). Suicide genes
floating in a glass of sparkling wine. Archives
of Suicide Research, 8(4), 297–301.
Maynard Smith, J. (1964). Group selection and
kin selection. Nature, 201, 1145–1147.
Mayr, E. (1965). Cause and effect in biology. In
D. Lerner (Ed.), Cause And Effect (pp. 33–
50). New York, NY: Free Press.
McGirr, A., Renaud, J., Bureau, A., Seguin, M.,
Lesage, A., & Turecki, G. (2008). Impulsive-
aggressive behaviours and completed suicide
across the life cycle: A predisposition for
younger age of suicide. Psychological
Medicine, 38(3), 407–417.
McNally, R. J. (2016). The legacy of Seligman’s
‘Phobias and Preparedness’ (1971). Behavior
Therapy, 47(5), 585–594.
Meehl, P. E. (1973). Why I Do Not Attend Case
Conferences. Minneapolis, MN: University of
Minnesota Press.
Mehlum, L., & Mork, E. (2016). After the
suicide attempt – the need for continuity and
87
quality of care. In R. C. O’Connor & J. Pirkis
(Eds.), The International Handbook Of
Suicide Prevention (pp. 387–402). Chichester,
UK: Wiley.
Melzack, R., & Casey, K. L. (1968). Sensory,
motivational and central control determinants
of pain: A new conceptual model. In D.
Kenshalo (Ed.), The Skin Senses (pp. 423–443).
Springfield, IL: Charles C. Thomas.
Menninger, K. A. (1938). Man Against Himself.
New York, NY: Harcourt Brace Jovanovich.
Menninger, K. A. (1963). The Vital Balance: The
Life Process In Mental Health And Illness.
New York, NY: Viking.
Merrick, J., Merrick, E., Lunsky, Y., & Kandel, I.
(2006). A review of suicidality in persons
with intellectual disability. The Israel Journal
of Psychiatry and Related Sciences, 43(4),
258.
Miller, G. (2000). Sexual selection for indicators
of intelligence. In G. R. Bock, J. A. Goode, &
K. Webb (Eds.), The Nature Of Intelligence:
Novartis Foundation Symposium 233
(pp. 260–275). Chichester, UK: Wiley.
Miller, G. F., & Penke, L. (2007). The evolution
of human intelligence and the coefficient of
additive genetic variance in human brain
size. Intelligence, 35(2), 97–114.
Miller, M. B. (2008). Suicide and evolution
(Thesis). Florida State University, Tallahassee,
FL.
Mishara, B. L. (1999). Conceptions of death
and suicide in children ages 6–12 and their
implications for suicide prevention. Suicide
and Life-Threatening Behavior, 29(2),
105–118.
Mishara, B. L. (2003). Suicide types: Rational. In
R. Kastenbaum (Ed.), Macmillan Encyclopedia
Of Death And Dying (Vol. 2). New York, NY:
Macmillan.
Mishara, B. L. (2006). Cultural specificity and
universality of suicide: Challenges for the
International Association for Suicide
Prevention. Crisis, 27(1), 1–3.
Mishara, B. L., & Chagnon, F. (2016). Why
mental illness is a risk factor for suicide:
Implications for suicide prevention. In R. C.
O’Connor & J. E. Pirkis (Eds.), International
Handbook Of Suicide Prevention (2nd ed.,
pp. 594–608). Chichester, UK: Wiley.
Mugisha, J., Hjelmeland, H., Kinyanda, E., &
Knizek, B. L. (2011). Distancing: A traditional
88 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
mechanism of dealing with suicide among
the Baganda, Uganda. Transcultural
Psychiatry, 48(5), 624–642.
Mullins, N., Bigdeli, T. B., Børglum, A., Coleman,
J., Demontis, D., Mehta, D., & Lewis, C. M.
(2019). GWAS of suicide attempt in
psychiatric disorders and association with
major depression polygenic risk scores.
American Journal of Psychiatry, 176(8),
651–660.
Murray, H. A., & Kluckhohn, C. (1948). Outline
of a conception of personality. In C. Kluckhohn
& H. A. Murray (Eds.), Personality In Nature,
Society, And Culture (pp. 3–32). Oxford, UK:
Knopf.
Naghavi, M. (2019). Global, regional, and national
burden of suicide mortality 1990 to 2016:
Systematic analysis for the Global Burden of
Disease Study 2016. BMJ, 364(l94), 1–11.
Nesse, R. M., & Schulkin, J. (2019). An
evolutionary medicine perspective on pain
and its disorders. Philosophical Transactions
of the Royal Society B, 374(20190288), 1–7.
Nesse, R. M., & Williams, G. C. (1995). Why
We Get Sick. New York, NY: Random House.
Nock, M. K., Borges, G., Bromet, E. J., Cha, C. B.,
Kessler, R. C., & Lee, S. (2012). The epidemiology
of suicide and suicidal behavior. In M. K. Nock,
G. Borges, & Y. Ono (Eds.), Suicide: Global
Perspectives From The WHO World Mental
Health Surveys (pp. 5–32). Cambridge, UK:
Cambridge University Press.
Nock, M. K., Borges, G., & Ono, Y. (2012).
Conclusions and future directions. In M. K.
Nock, G. Borges, & Y. Ono (Eds.), Suicide:
Global Perspectives From The WHO World
Mental Health Surveys (pp. 222–225).
Cambridge, UK: Cambridge University Press.
Nock, M. K., Green, J., Hwang, I., McLaughlin,
K. A., Sampson, N. A., Zaslavsky, A. M., &
Kessler, R. C. (2013). Prevalence, correlates,
and treatment of lifetime suicidal behavior
among adolescents: Results from the National
Comorbidity Survey Replication Adolescent
Supplement. JAMA Psychiatry, 70(3),
300–310.
Nock, M. K., Ramirez, F., & Rankin, O. (2019).
Advancing our understanding of the who,
when, and why of suicide risk. JAMA
Psychiatry, 76(1), 11–12.
Nock, M. K., Ursano, R. J., Heeringa, S. G.,
Stein, M. B., Jain, S., Raman, R., & Fullerton,
C. S. (2015). Mental disorders, comorbidity,
and pre-enlistment suicidal behavior among
new soldiers in the US Army: Results from
the Army Study to Assess Risk and Resilience
in Service members (Army STARRS). Suicide
and Life-Threatening Behavior, 45(5),
588–599.
O’Connell, S., & Dunbar, R. (2003). A test for
comprehension of false belief in chimpanzees.
Evolution and Cognition, 9(2), 131–140.
O’Connor, R. J. (1978). Brood reduction in
birds: Selection for fratricide, infanticide and
suicide? Animal Behaviour, 26, 79–96.
O’Connor, R. C. (2011). Towards an integrated
motivational-volitional model of suicidal
behaviour. In R. C. O’Connor, S. Platt, &
J. Gordon (Eds.), International Handbook Of
Suicide Prevention: Research, Policy And
Practice (pp. 181–198). Chichester, UK: John
Wiley and Sons.
O’Connor, R. C., Cleare, S., Eschle, S.,
Wetherall, K., & Kirtley, O. J. (2016).
The integrated motivational-volitional model
of suicidal behavior: An update. In R. C.
O’Connor & J. Pirkis (Eds.), International
Handbook Of Suicide Prevention (2nd ed.,
pp. 220–240). Chichester, UK: John Wiley
and Sons.
O’Connor, R. C., & Portzky, G. (2018). Looking
to the future: A synthesis of new developments
and challenges in suicide research and
prevention. Frontiers in Psychology, 9(2139),
1–14.
Orbell, J., & Morikawa, T. (2011). An
evolutionary account of suicide attacks: The
kamikaze case. Political Psychology, 32(2),
297–322.
Paniagua, F. A., Black, S. A., Gallaway, M. S., &
Coombs, M. A. (2010). The Interpersonal-
Psychological Theory Of Attempted And
Completed Suicide. Bloomington, IN:
AuthorHouse.
Parker, G. A., & Smith, J. M. (1990). Optimality
theory in evolutionary biology. Nature,
348(6296), 27.
Paulhus, D. L., & Buckels, E. (2012). Classic self-
deception revisited. In S. Vazire & T. D.
Wilson (Eds.), Handbook Of Self-Knowledge
(pp. 363–378). New York, NY: Guilford Press.
Peña-Guzmán, D. M. (2018). Can nondolphins
commit suicide? Animal Sentience, 2(20),
20.
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
Penney, D., Stewart, A., & Parr, M. (2002).
Prognostic outcome indicators following
hanging injuries. Resuscitation, 54(1), 27–29.
Perry, S. A. (2014). Every Cradle Is A Grave:
Rethinking The Ethics Of Birth And Suicide.
Charleston, WV: Nine-Banded Books.
Perry, S. A. (2015). Antinatalism in biological
and cultural evolution: Fertility and suicide.
In T. K. Shackelford & R. D. Hansen (Eds.),
The Evolution Of Morality (pp. 141–154).
Cham, Switzerland: Springer.
Perry, S. A. (2016). Patch 7.822: An
experimental design puzzle. Retrieved from
https://theviewfromhellyes.wordpress.com
(October 2016)
Persley, G., & Pegg, S. P. (1981). Burn injuries
related to suicide. Medical Journal of
Australia, 1(3), 134.
Pinker, S. (1997). How The Mind Works. New
York, NY: Norton.
Pinker, S. (2010). The cognitive niche:
Coevolution of intelligence, sociality, and
language. Proceedings of the National
Academy of Sciences, 107(Supplement 2),
8993–8999.
Pitman, A. L., Osborn, D. P. J., King, M. B., &
Erlangsen, A. (2014). Effects of suicide
bereavement on mental health and suicide
risk. The Lancet Psychiatry, 1(1), 86–94.
Pitman, A. L., Osborn, D. P. J., Rantell, K., &
King, M. B. (2016). Bereavement by suicide
as a risk factor for suicide attempt: A cross-
sectional national UK-wide study of 3432
young bereaved adults. BMJ, 6(1), e009948.
Poole, F. J. P. (1985). Among the boughs of the
hanging tree: Male suicide among the Bimin-
Kuskusmin of Papua New Guinea. In F. X.
Hezel, D. H. Rubinstein, & G. M. White
(Eds.), Culture, Youth and Suicide in the
Pacific: Papers from an East-West Center
Conference (pp. 152–181). Honolulu, HI:
University of Hawaii at Manoa.
Preti, A. (2007). Suicide among animals: a
review of evidence. Psychological Reports,
101(3), 831–848.
Preti, A. (2011). Animal model and neurobiol-
ogy of suicide. Progress in Neuro-­
Psychopharmacology and Biological
Psychiatry, 35(4), 818–830.
Ramsden, E., & Wilson, D. (2010). The nature
of suicide: Science and the self-destructive
animal. Endeavour, 34(1), 21–24.
89
Riordan, D. (2019). Suicide and human sacrifice;
sacrificial victim hypothesis on the
evolutionary origins of suicide. Suicidology
Online, 10(2), 1–10.
Rogers, J. R. (2001). Theoretical grounding: The
‘missing link’ in suicide research. Journal of
Counseling & Development, 79(1), 16–25.
Rogers, M. L., Ringer, F. B., & Joiner, T. E.
(2018). The association between suicidal
ideation and lifetime suicide attempts is
strongest at low levels of depression.
Psychiatry Research, 270, 324–328.
Rubinstein, D. H. (1986). A stress–diathesis
theory of suicide. Suicide and Life-
Threatening Behavior, 16(2), 182–197.
Saad, G. (2007). Suicide triggers as sex-specific
threats in domains of evolutionary import:
Negative correlation between global male-to-
female suicide ratios and average per capita
gross national income. Medical Hypotheses,
68(3), 692–696.
Salim, A., Martin, M., Sangthong, B., Brown,
C., Rhee, P., & Demetriades, D. (2006). Near-
hanging injuries: A 10-year experience.
Injury, 37(5), 435–439.
Sarkar, S. (1998). Genetics and Reductionism.
Cambridge, UK: Cambridge University
Press.
Satcher, D. (1999). The Surgeon General’s Call
To Action To Prevent Suicide. Washington,
DC: US Public Health Service.
Saunders, K. E., Hawton, K., Fortune, S., &
Farrell, S. (2012). Attitudes and knowledge
of clinical staff regarding people who self-
harm: A systematic review. Journal of
Affective Disorders, 139(3), 205–216.
Schaeffer, H. (1967). Can a mouse commit
suicide? In E. Shneidman (Ed.), Essays In Self
Destruction. Northvale, NJ: Jason Aronson.
Schechter, M. A., & Goldblatt, M. J. (2020).
Suicide: A transdiagnostic phenomenon.
Psychiatric Annals, 50, 136–137.
Schmidtke, A. (1997). Suicide in Europe.
Suicide and Life-Threatening Behavior, 27(1),
127–136.
Seiden, R. H. (1969). Suicide Among Youth. A
Review Of The Literature, 1900–1967. Chevy
Chase, MD: National Clearinghouse for
Mental Health Information.
Selby, E. A., Joiner, T. E., & Ribeiro, J. D. (2014).
Comprehensive theories of suicidal behaviors.
In M. K. Nock (Ed.), The Oxford Handbook
90 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Of Suicide And Self-Injury (pp. 286–307).
Oxford, UK: Oxford University Press.
Sesardic, N. (2005). From genes to incest
taboos: The crucial step. In A. P. Wolf & W.
H. Durham (Eds.), Inbreeding, Incest, And
The Incest Taboo: The State Of Knowledge
At The Turn Of The Century (pp. 109–120).
Stanford, CA: Stanford University Press.
Seyfried, L. S., Kales, H. C., Ignacio, R. V.,
Conwell, Y., & Valenstein, M. (2011).
Predictors of suicide in patients with
dementia. Alzheimer’s & Dementia, 7(6),
567–573.
Shaffer, D. (1974). Suicide in childhood and
early adolescence. Journal of Child
Psychology and Psychiatry, 15(4), 275–291.
Sharma, T., Guski, L. S., Freund, N., & Gøtzsche,
P. C. (2016). Suicidality and aggression
during antidepressant treatment: Systematic
review and meta-analyses based on clinical
study reports. BMJ, 352, i65.
Sheehan, L. L., Corrigan, P. W., & Al-Khouja, M.
A. (2016). Stakeholder perspectives on the
stigma of suicide attempt survivors. Crisis,
38(2), 73–81. Retrieved from https://doi.
org/10.1027/0227-5910/a000413
Shneidman, E. S. (1985). Definition Of Suicide.
New York, NY: John Wiley & Sons.
Shneidman, E. S. (1993). Suicide As Psychache:
A Clinical Approach To Self-Destructive
Behavior. Lanham, MD: Rowman & Littlefield.
Shneidman, E. S. (1996). The Suicidal Mind.
New York, NY: Oxford University Press.
Shneidman, E. S. (2001). Comprehending
Suicide. Landmarks In 20th-Century
Suicidology. Washington, DC: American
Psychological Association.
Shneidman, E. S. (2005). Anodyne
psychotherapy for suicide: A psychological
view of suicide. Clinical Neuropsychiatry,
2(1), 7–12.
Shneidman, E. S., & Farberow, N. L. (Eds.).
(1961). The Cry For Help. New York, NY:
McGraw-Hill.
Shorter, J., & Rueppell, O. (2012). A review on
self-destructive defense behaviors in social
insects. Insectes Sociaux, 59(1), 1–10.
Skinner, B. F. (1969). Contingencies Of
Reinforcement: A Theoretical Analysis. New
York, NY: Meredith.
Slaughter, V., & Griffiths, M. (2007). Death
understanding and fear of death in young
children. Clinical Child Psychology and
Psychiatry, 12(4), 525–535.
Solano, P., Pizzorno, E., Pompili, M., Serafini,
G., & Amore, M. (2018). Conceptualizations
of suicide through time and socio-economic
factors: a historical mini-review. Irish Journal
of Psychological Medicine, 35(1), 75–86.
Soper, C. A. (2016). Could some common
mental diseases have evolved to prevent
suicide? – a theoretical enquiry. Paper
presented at the 16th European Symposium
on Suicide and Suicidal Behaviour, Oviedo,
Spain.
Soper, C. A. (2017). Towards solving the
evolutionary puzzle of suicide (PhD thesis).
University of Gloucestershire, Cheltenham,
UK.
Soper, C. A. (2018). The Evolution Of Suicide.
Cham, Switzerland: Springer.
Soper, C. A. (2019a). Beyond the search for
suigiston: How evolution offers oxygen for
suicidology. In V. Zeigler-Hill & T. K.
Shackelford (Eds.), Evolutionary Perspectives
On Death (pp. 37–61). Cham, Switzerland:
Springer.
Soper, C. A. (2019b). Adaptation to the suicidal
niche. Evolutionary Psychological Science.
5(4), 454–471.
Soper, C. A., & Shackelford, T. K. (2018). If
nonhuman animals can suicide, why don’t
they? Animal Sentience, 2(20), 14.
Staats, A. W. (2004). The disunity-unity
dimension. American Psychologist, 59(4),
273.
Stanley, I., Hom, M., Boffa, J., Stage, D. R. L.,
& Joiner, T. E. (2019). PTSD from a suicide
attempt: An empirical investigation among
suicide attempt survivors. Journal of Clinical
Psychology, 75(10), 1879–1895.
Steinmetz, S. R. (1894). Suicide among primitive
peoples. American Anthropologist, 7(1),
53–60.
Stengel, E. (1970). Suicide And Attempted
Suicide (Revised ed.). London, UK: Penguin.
Stochl, J., Khandaker, G., Lewis, G., Perez, J.,
Goodyer, I., Zammit, S., & Jones, P. (2015).
Mood, anxiety and psychotic phenomena
measure a common psychopathological
factor. Psychological Medicine, 45(7),
1483–1493.
Sudak, H., Maxim, K., & Carpenter, M. (2008).
Suicide and stigma: A review of the literature
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY
and personal reflections. Academic
Psychiatry, 32(2), 136–142.
Suddendorf, T. (2013). The Gap: The Science
Of What Separates Us From Other Animals.
New York, NY: Basic Books.
Sveen, C. A., & Walby, F. A. (2008). Suicide
survivors’ mental health and grief reactions:
A systematic review of controlled studies.
Suicide and Life-Threatening Behavior, 38(1),
13–29.
Syme, K. L., Garfield, Z. H., & Hagen, E. H.
(2016). Testing the bargaining vs. inclusive
fitness models of suicidal behavior against
the ethnographic record. Evolution and
Human Behavior, 37(3), 179–192.
Syme, K. L., & Hagen, E. H. (2018). When
saying ‘sorry’ isn’t enough: Is some suicidal
behavior a costly signal of apology? A cross-
cultural test. Human Nature, 30(1),
117–141.
Symons, D. (1992). On the use and misuse of
Darwinism in the study of human behavior.
In J. Barkow, L. Cosmides, & J. Tooby (Eds.),
The Adapted Mind: Evolutionary Psychology
And The Generation Of Culture (pp. 137–
162). New York, NY: Oxford University Press.
Tanaka, M., & Kinney, D. K. (2011). An
evolutionary hypothesis of suicide: Why it
could be biologically adaptive and is so
prevalent in certain occupations.
Psychological Reports, 108(3), 977–992.
Tidemalm, D., Runeson, B., Waern, M., Frisell,
T., Carlström, E., Lichtenstein, P., &
Långström, N. (2011). Familial clustering of
suicide risk: A total population study of 11.4
million individuals. Psychological Medicine,
41(12), 2527–2534.
Tinbergen, N. (1963). On aims and methods of
ethology. Zeitschrift für Tierpsychologie,
20(4), 410–433.
Tolaas, J. (2005). Evolution And Suicide. Salt
Lake City, UT: American University & Colleges
Press.
Tooby, J., & Cosmides, L. (1990a). On the
universality of human nature and the
uniqueness of the individual: The role of
genetics and adaptation. Journal of
Personality, 58(1), 17–67.
Tooby, J., & Cosmides, L. (1990b). The past
explains the present: Emotional adaptations
and the structure of ancestral environments.
Ethology and Sociobiology, 11(4), 375–424.
91
Tooby, J., & Cosmides, L. (1992). The psychological
foundations of culture. In J. H. Barkow,
L. Cosmides, & J. Tooby (Eds.), The Adapted
Mind: Evolutionary Psychology And The
Generation Of Culture (pp. 19–136). New
York, NY: Oxford University Press.
Tooby, J., & Cosmides, L. (2005). Conceptual
foundations of evolutionary psychology. In
D. M. Buss (Ed.), The Handbook Of
Evolutionary Psychology (pp. 5–67).
Hoboken, NJ: John Wiley & Sons.
Tooby, J., & Cosmides, L. (2008). The
evolutionary psychology of the emotions and
their relationship to internal regulatory
variables. In M. Lewis, J. M. Haviland-Jones,
& L. F. Barrett (Eds.), Handbook Of Emotions
(3rd ed., pp. 114–137). New York, NY:
Guilford Press.
Tooby, J., & DeVore, I. (1987). The reconstruction
of hominid behavioral evolution through
strategic modeling. In W. G. Kinzey (Ed.), The
Evolution Of Human Behavior: Primate
Models (pp. 183–237). Albany, NY: State
University of New York.
Tousignant, M. (1998). Suicide in small-scale
societies. Transcultural Psychiatry, 35(2),
291–306.
Townsend, E. (2007). Suicide terrorists: Are
they suicidal? Suicide and Life-Threatening
Behavior, 37(1), 35–49.
Turecki, G., & Brent, D. A. (2016). Suicide and
suicidal behaviour. The Lancet, 387(10024),
1227–1239.
Usall, J., Pinto-Meza, A., Fernández, A., de
Graaf, R., Demyttenaere, K., Alonso, J., &
Haro, J. M. (2009). Suicide ideation across
reproductive life cycle of women: Results from
a European epidemiological study. Journal of
Affective Disorders, 116(1), 144–147.
van Dongen, J., & Boomsma, D. I. (2013). The
evolutionary paradox and the missing
heritability of schizophrenia. American
Journal of Medical Genetics Part B:
Neuropsychiatric Genetics, 162(2), 122–136.
Van Orden, K. A., Witte, T. K., Cukrowicz, K.
C., Braithwaite, S. R., Selby, E. A., & Joiner,
T. E. (2010). The interpersonal theory of
suicide. Psychological Review, 117(2),
575–600.
Varki, A., & Brower, D. (2013). Denial: Self-
Deception, False Beliefs, And The Origins Of
The Human Mind. New York, NY: Twelve.
92 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Värnik, P. (2012). Suicide in the world.
International Journal of Environmental
Research And Public Health, 9(3), 760–771.
Voracek, M. (2006). Smart and suicidal? The
social ecology of intelligence and suicide in
Austria. Death Studies, 30(5), 471–485.
Voracek, M., & Marušič, A. (2008). Testing the
Finno-Ugrian Suicide Hypothesis: Geographic
variation of elderly suicide rates across
Europe. Nordic Journal of Psychiatry, 62(4),
302–308.
Wiley, J. (2020). Psychological Aposematism:
An Evolutionary Analysis of Suicide. Biological
Theory. doi:10.1007/s13752-020-00353-8
World Health Organization. (2012). Public
Health Action For The Prevention Of Suicide:
A Framework. Geneva, Switzerland: World
Health Organization.
World Health Organization. (2013). Projections
of Mortality and Causes of Death, 2015 and
2030. Retrieved from www.who.int/
healthinfo/global_burden_disease/en/
(Accessed February 2016)
World Health Organization. (2014). Preventing
Suicide: A Global Imperative. Geneva,
Switzerland: World Health Organization.
Waldmann, M. R., Hagmayer, Y., & Blaisdell, A.
P. (2006). Beyond the information given:
Causal models in learning and reasoning.
Current Directions in Psychological Science,
15(6), 307–311.
Walker, M., Moreau, D., & Weissman, M. M.
(1990). Parents’ awareness of children’s
suicide attempts. American Journal of
Psychiatry, 147(10), 1364–1366.
Wall, P. D. (1999). Pain: The Science Of
Suffering. London, UK: Weidenfeld &
Nicholson.
Wampold, B. E. (2018). The therapeutic value
of the relationship for placebo effects and
other healing practices. International Review
of Neurobiology, 139, 191–210.
Wampold, B. E., & Imel, Z. E. (2015). The Great
Psychotherapy Debate: The Evidence For
What Makes Psychotherapy Work (2nd ed.).
New York, NY: Routledge.
Wasserman, D., Rihmer, Z., Rujescu, D.,
Sarchiapone, M., Sokolowski, M., Titelman,
D., & Carli, V. (2012). The European
Psychiatric Association (EPA) guidance on
suicide treatment and prevention. European
Psychiatry, 27(2), 129–141.
Wertheimer, A. (2014). A Special Scar: The
Experiences Of People Bereaved By Suicide
(3rd ed.). Abingdon, UK: Routledge.
Williams, G. C. (1957). Pleiotropy, natural
selection, and the evolution of senescence.
Evolution, 11(4), 398–411.
Williams, G. C. (1966). Adaptation And Natural
Selection: A Critique Of Some Current
Evolutionary Thought. Princeton, NJ:
Princeton University Press.
Williams, G. C. (1992). Natural Selection:
Domains, Levels, And Challenges. Oxford,
UK: Oxford University Press.
Williams, G. C. (1996). Plan And Purpose In
Nature. London, UK: Weidenfeld & Nicholson.
Williams, J. M. G. (1997). Cry Of Pain:
Understanding Suicide And Self-Harm.
London, UK: Penguin.
Williams, J. M. G., Crane, C., Barnhofer, T., &
Duggan, D. S. (2005). Psychology and
suicidal behaviour: Elaborating the
entrapment model. In K. Hawton (Ed.),
Prevention And Treatment Of Suicidal
Behaviour: From Science To Practice (pp.
71–89). Oxford, UK: Oxford University Press.
Williams, J. M. G., & Pollock, L. R. (2000). The
psychology of suicidal behaviour. In K.
Hawton & K. van Heeringen (Eds.), The
International Handbook Of Suicide And
Attempted Suicide (pp. 79–94). Chichester,
UK: John Wiley & Sons.
Willner, P., & Belzung, C. (2015). Treatment-
resistant depression: Are animal models of
depression fit for purpose? Psychopharma-
cology, 232(19), 3473–3495.
Wilson, D. S. (1980). Suicide, beanbag genetics,
and pleiotropy. Behavioral and Brain Sciences,
3(2), 283.
Wilson, D. S., & Wilson, E. O. (2007). Rethinking
the theoretical foundation of sociobiology.
The Quarterly Review of Biology, 82(4),
327–348.
Wilson, J. D., & Roehrborn, C. (1999). Long-
term consequences of castration in men:
Lessons from the Skoptzy and the eunuchs
of the Chinese and Ottoman courts. The
Journal of Clinical Endocrinology &
Metabolism, 84(12), 4324–4331.
Wright, R. (1994). The Moral Animal. New
York, NY: Vintage.
Wright, S. (1943). Isolation by distance.
Genetics, 28(2), 114–138.
 EVOLUTIONARY PSYCHOLOGY AND SUICIDOLOGY 93

Zalsman, G., Hawton, K., Wasserman, D., van
Heeringen, K., Arensman, E., Sarchiapone,
M., & Zohar, J. (2017). Evidence-based
national suicide prevention taskforce in
Europe: A consensus position paper. European
Neuropsychopharmacology, 27(4), 418–421.
Zilboorg, G. (1936a). Differential diagnostic
types of suicide. Archives of Neurology &
Psychiatry, 35(2), 270–291.
Zilboorg, G. (1936b). Suicide among civilized
and primitive races. American Journal of
Psychiatry, 92(6), 1347–1369.

Evolutionary Psychology and
Mindfulness and Meditation:
Easing the Anxiety of Being
Human
INTRODUCTION
Something called mindfulness seems to be
popping up everywhere: in clinics and hospi-
tals, schools, corporate offices, prisons,
online, the list keeps expanding. Today, at the
local farmer’s market, I bought greens from
‘Mindful Veggies’. Promoted as a wellbeing
enhancer, studies show that mindfulness does
indeed reduce stress and distress in a range of
conditions and circumstances (Goyal et  al.,
2014; Willem et al., 2016). In this chapter I
describe the evolutionary roots of the psy-
chological processes that give rise to the
human angst that mindfulness addresses and
how the training exercises designed to culti-
vate it alleviate that distress. And to provide
context for those I first describe the cultural
roots of mindfulness and meditation.
Several innovative approaches to inquiry
emerged during what has been called the
Axial Age. In contrast to the Platonic and
Aristotelian systems developing in Greece
during that era, internal psychological models
4
James Carmody
developed in India including those of Vedanta
and Buddhism. Training exercises designed to
develop the personal qualities required to actu-
alize the models’ goals were also developed
and in keeping with approaches to knowledge
at the time, these were not clearly distinguished
from religion. Mindfulness, as it is now com-
monly recognized in the West, came primarily
out of Buddhism and is described in the follow-
ing section.
Buddhist principles, particularly, later
spread into other parts of Asia and integrated
into those countries’ prevailing spiritual belief
systems. Both parties were changed as a result
of those meetings. After a hiatus of several
centuries, increased global mobility brought
Buddhism to Western countries and once
again it has been adapted into the prevailing
cultural narrative. One of those adaptations
has been the integration of mindfulness, a core
tenet of the system described in the following
section, into the cultural narrative of self-help.
The cultural origins and purpose of Buddhism
however have parallels in some Vedanta
 EVOLUTIONARY PSYCHOLOGY AND MINDFULNESS AND MEDITATION
practices that also have migrated to the West
in the ­ various forms of yoga. This secular
transition has made its benefits more widely
available than they might otherwise have been,
for although a practice from a religious system
is attractive to some people, it is alienating to
many in the secular West.
This transition has also brought mindfulness
under the gaze of empirical science and close
inspection of its mechanisms of action reveals
parallels and differences in some principles
independently developed in Western psychol-
ogy to categorize, account for, and alleviate
human angst. Those principles, together with
evolutionary theory, can provide a coherent
and culturally familiar explanation of the eve-
ryday mental unease that plagues us and how
the practice of mindfulness alleviates it. This
evolutionary and needs-based lens describes
inbuilt patterns of attending that keep us ill-
at-ease, and the psychological principles that
mindfulness and several similar mind–body
practices draw upon in enabling the recogni-
tion and amelioration of that distress.
The description draws upon my own and
others’ published studies of the clinical effects
and mechanisms of mindfulness training (MT),
as well as experience and feedback from teach-
ing mindfulness to patients and clinicians. The
description is phenomenological because suf-
fering and mindfulness are rooted in felt experi-
ence and it is their felt sense that people wish
to address in doing the practices. Clinicians
also talk most meaningfully to patients in those
terms. In that sense, mindfulness practices can
be thought of as phenomenologically empiri-
cal explorations of the mental processes giving
rise to mental distress and the capacity for self-­
regulation that emerges in the face of it.
The chapter also draws upon my own expe-
rience with 50 years of practice in Buddhist
and Vedantic traditions of inquiry in Asia and
Western countries. Many academic papers
about mindfulness are caught in attempting
objectivity in relation to something that is
often anything but. In the background, u­ sually
undescribed, stands the author’s positive per-
sonal experience with mindfulness that led
95
them to the research and that has guided their
decisions about the conditions, instruction,
and syllabus of the mindfulness intervention.
For mindfulness makes apparent the normally
unrecognized patterns of attending that never-
theless continuously affect our felt sense and
the background hope is that study participants
also will benefit from that kind of noticing.
THE CULTURAL ROOTS OF
MINDFULNESS AND ITS MIGRATION
INTO WESTERN SETTINGS
The Buddhist and Vedanta narratives are
rooted in the primarily introspective
approaches to knowledge extant in India
around 500 BCE, the time of the historical
Buddha. They each place the source of human
angst in ignorance of the moment-by-moment
construction of the personal self, and the
chronic dissatisfaction (suffering) arising
from the accompanying sense of ownership of
its desires and aversions. In this respect they
are not a description of how the physical
world operates, but of what we call mind.
As the Buddha described his insight into
this dilemma to others and the experience of
enlightened release accompanying it, an eight-
faceted system developed through which they
also could cultivate a similar recognition. One
facet was the cultivation of something called
‘sati’ to serve as a heuristic aid for real-time
recognition of these mental operations and their
effects. Sati is from the Pali language spoken
in Northern India at the time, but that is no
longer understood outside the Indian scholar-
ship community. Exactly how sati was trans-
lated, described, and cultivated varied in the
places to which Buddhism migrated over the
centuries. This is apparent in the varied teach-
ings and practices of the Western Buddhist sects
whose traditions derive from those countries.
Mindfulness, a term with an already existing
meaning in English, emerged as the accepted
translation during the 19th century as Buddhism
was becoming of interest in the West.
96 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Recognizing the commonality that
Buddhism’s goal of reducing mental suffering
had with the goal of patient care, Kabat-Zinn
(Kabat-Zinn, 1982) experimented with teach-
ing groups of patients a selection of training
exercises used to cultivate sati in South Asian
traditions. Those patients described obtain-
ing benefits in coping with their illnesses
from participating in the classes that came
to be called the Mindfulness-Based Stress
Reduction (MBSR) program. Replications in
peer-reviewed journals affirmed those benefits
and the program became widely used in clinics
(Shonin et al., 2015). Principles and practices
from MBSR form the foundation for many of
the adaptations and applications found in the
wide variety of settings in which mindfulness
now appears.
While the processes at the root of the men-
tal distress that the cultivation of mindfulness
addresses are interwoven and run in parallel,
for explanatory purposes I describe them as
a sequence. It is important also to note that
early Buddhism employed a somewhat differ-
ent frame to categorize the qualities of expe-
rience than Western systems do. For example,
it did not use the construct of emotion in the
description of affective experience; rather it
used a more granular analysis of experien-
tial components that comprise those states
as described in following sections. Attention
regulation, however, is common across the
systems and is known to be fundamental to
wellbeing (Posner and Rothbart, 2007). For
that reason, it may be best to begin with
attention and the priorities that effect it.
THE EVOLUTIONARY ROOTS OF
ATTENTION AND ITS ROLE IN
A NEEDS-BASED SYSTEM
MT usually begins with an attention-regulation
task. A common one asks the person to place
their focus on the sensations of their breathing
and to keep it there for some time. While this
may seem a relatively easy task, trainees report
it as one of the most challenging (Segal et al.,
2013). They stay with it for a breath or two
before attention wanders to daydreaming. The
tenacity of this default movement to cognitions
in the absence of sustained watchfulness sug-
gests an important function, one that becomes
clear in reflecting on the role attention plays in
our mental ecology.
Our brains are continuously processing all
manner of information about the internal and
external environment. Attention, the capac-
ity to selectively attend to some portion of
this information over others, particularly
opportunities and threats to the fulfillment of
needs, has clear value for survival and repro-
ductive success (Geary, 2005). Selection has
also resulted in this process being i­mmediate
rather than through conscious, deliberate, and
slower decision-making pathways (Tomlin
et al., 2015). When physical danger threatens,
as it regularly did in the evolutionary past,
attention automatically orients to sensory
processes monitoring the external environ-
ment. Attention is also intimately connected to
arousal. In the best-selling book Why Zebras
Don’t Get Ulcers, Sapolsky (2004) describes
how the attention of animals in the wild is ori-
ented to the senses and to arousal: danger is
smelled, seen, or heard, arousal levels and ten-
sion spike, the animal responds in some way,
and a more quiescent state resumes.
We also would live in a more relaxed here
and now if real and immediate physical dan-
gers were the principle risks that our atten-
tion responded to. But being in the moment is
not characteristic of modern life. The capac-
ity for cognition and language makes pos-
sible the imagination of conceivable future
threats (Andrews-Hanna, 2012; Baumeister
et  al., 2001) as well as complex planning
and discussion of how best to address them.
The importance of those capacities to mod-
ern humans is evident in our attention’s
insistent default to the concerns of projected
futures and pasts when it is not required for
the execution of an immediate task. This can
be observed by deliberately taking notice of
the content of the imaginings to which our
minds ‘wander’. There we see recurring and
often conflicting and entangled concerns about
 EVOLUTIONARY PSYCHOLOGY AND MINDFULNESS AND MEDITATION
our own and our family’s and friends’ welfare,
social standing, sex, work, and money.
The persistence of this default, and its
affective upshot, was captured nicely in a
real-time study by Killingsworth and Gilbert
(2010). Prompted at random times to report
what their attention was on in that moment,
subjects reported it being on the task at hand
for only about half the day, even though they
reported greater happiness at those times.
Significantly, the attention wandering gener-
ally preceded the happiness decrease. It is this
propensity of attention to quickly and repeat-
edly default to cognitions, memories, plans,
speculations, and daydreams, that mindful-
ness trainees encounter when asked to focus
on sense-based experience, and that they find
so challenging to overrule and regulate. They
discover also that much of this cognition goes
on outside of awareness and becomes appar-
ent only when they deliberately watch their
mental activity.
In the absence of immediate danger, then,
attention functionally defaults to cognitive
processes serving the social-safety needs of
tribal primates, particularly those for relation-
ship/belonging, status, and power. The bio-
logical importance of these needs is evident
in the fact that health becomes compromised
when, for some reason, we are stripped of
them. Also, their intricately interwoven and at
times conflicting nature, together with the fact
that they have no organic satiation mechanism,
Figure 4.1  Alarm-related components of experience forming a cycle of distress
97
means that navigating threats and opportuni-
ties and monitoring projected pasts and futures
is a constant project.
DARWIN AND BUDDHISM’S FIRST
NOBLE TRUTH: VIGILANCE AND
THE INTERNAL NARRATIVE MAKE
UNEASE OUR EVERYDAY STATE
In traditional Buddhist teachings, the reduced
happiness Killingsworth and Gilbert (2010)
found associated with the preoccupations of a
wandering mind results not from the thoughts
and imaginings themselves, but from the
unpleasant sensations of constriction that
reflexively accompany their semi-vigilant
(‘what could go wrong here?’) and uncertainty-
related character. A discomforting cycle is then
set up as those unpleasant sensations of tension
remind us again of the threat-related thought
(Damasio, 2003). Extended preoccupation with
these alarm-based cycles is experienced as
rumination and worry. These are accompanied
by arousal-related inflammatory processes. As
we’ve all experienced, the level of arousal, con-
striction, and unpleasantness can range from
very mild to dreadful depending on the degree
to which those needs are frustrated or threat-
ened (Brosschot et al., 2006). Figure 4.1 illus-
trates this vigilance-based cycle in the
experience of anxiety.
98 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
When mental bandwidth is occupied by
this narrative, curiosity and connection with
our surroundings becomes problem-focused,
and we are less than openly present for the
present needs and concerns of others. When
attention is task-related, however, this inter-
nal narrative recedes (Watkins, 2004) and we
are less affected by its threat- and opportunity-
oriented memories and imaginings.
Even when these preoccupations barely
reach awareness (Brosschot et  al., 2014;
Creswell et al., 2013; Custers and Aarts, 2010;
Dahl et al., 2015), the ongoing bodily discom-
fort gives rise to a desire for a break in some-
thing more pleasant; to interrupt the cycle and
replace it temporarily with one characterized
by greater ease. And because the experience
of pleasure and delight encompasses the body
(Biswas-Diener et al., 2015), a sense pleasure
is usually the most immediately accessible. It
might be something like a snack, a drink, or
a drug. It may also be something more elabo-
rate; the possibilities are as broad as imagina-
tion and resources allow.
We don’t start life cognitively preoccupied
in this way. We arrive as little sensate creatures,
curious and awake to the wonder of touching,
tasting, hearing, seeing, moving, and their
pleasant and unpleasant feeling tones. A cog-
nitive past and future become gradually and
imperceptibly woven into this sensory expe-
rience alongside language, the naming and
appraisal enabling us to describe our needs
and impressions to others (Alderson-Day and
Fernyhough, 2015). As we learn to weave
our way through the fragile social maze, the
hopes, comparisons, judgments, and regrets
embedded in this emerging internal narrative
come to mediate and filter sensory perception
and experience, and weave indiscernibly into a
developing sense of self.
Using Our Heads to Get out of
Them
Just as this safety-oriented cognitive watch-
fulness has an affective downside, cognition
can also wish for and imagine a life free of it
and trial possible solutions. That’s a reflec-
tion for the ages. It is also one that preoccu-
pied the historical Buddha who, after intense
experimentation, recognized the above pat-
terns shaping the affective quality of his own
life as well as related insights into how these
can be overcome. In conveying these to inter-
ested friends he saw that they were also
useful to others.
Three of those insights are contained, to a
greater and lesser degree, within present-day
MT programs. They are designed to support
greater awareness, acceptance, and some level
of everyday self-regulation in the face of these
mental and perceptual tendencies shaping
everyday feeling life (Cavanagh et  al., 2014;
Donald et  al., 2016; Saunders et  al., 2016)
and can be operationalized in familiar psycho-
logical terms. The analysis is not regarded as
the final word on mental mechanisms but as
a description of mental qualities that can be
phenomenologically recognized in real time,
and trainees comprehend and use them to suit
their interests and circumstances (Carmody
and Baer, 2008; Cebolla et al., 2017).
The first insight, alluded to in the previ-
ous section, is that everyday experience and
emotions are constructed from a suite of just
three interwoven phenomenological compo-
nents: cognitions, sensations, and their pleas-
ant/unpleasant feeling tones. Classification
and description of the affective dimension of
experience change with the age and the cul-
ture, and feeling here is not to be confused
with how we commonly use the term today
when we might say we are feeling angry or
sad. It’s simply the pleasant or unpleasant
quality of a sensation. Also, emotional cat-
egories vary across cultures and over time;
something described as an emotion at one
time might be referred to as a passion in
another. In the Buddhist analysis emotions
are approached more granularly as amalgams
of those three fundamental qualities. When
those components are not distinguished their
felt experience is seamless, as illustrated in
Figure 4.2.
 EVOLUTIONARY PSYCHOLOGY AND MINDFULNESS AND MEDITATION
Figure 4.2  Memory, imagination and emotion are symphonies of three interwoven
experiential components
Experiential recognition and discrimina-
tion of the components starts, most usually,
with re-awakening interest in the realm of
sensation. In the body scan, for example, an
exercise taught in MBSR (Kabat-Zinn, 1990),
trainees are asked to systematically direct
attention to each part of the body and to notice
any sensations that may be present there,
including subtle and neglected sensations that
may escape awareness in everyday life; this
while refraining from attempting to change the
experience in any way. They are instructed to
notice also any pleasant or unpleasant feeling
tone that may be associated with a sensation,
and the difference between the sensation and
any thoughts that may also be present. This is
schematically illustrated in Figure 4.3 using
the example of a fearful thought and the sen-
sations of constriction that may be associated
with it, and which, on a less granular level, we
experience as anxiety.
This learning is akin to winding back the
developmental and experiential clock. As
described in the previous section, we are
not born with these groupings. Cognitions
that had been so implicitly integrated (Blair,
2002) into our early world of sensation and
99
affect that their distinction was not apparent
in awareness (Pessoa, 2008) become noticed
and named. Their associations, so rapid they
are normally missed, become apparent in
noticing that attention does not stay with a
sensation, but quickly goes to its feeling tone,
to thoughts about it, or to something else.
Implicit in this bare noticing of experience is
an embodied acceptance, one that also may
be made explicit in the instructions.
Interoceptive awareness is important in
emotion regulation (Füstös et  al., 2013)
and the perception of internal experience
developed by these MT exercises appears
to mediate its beneficial effect on emo-
tional wellbeing (Mehling et al., 2012). And
even though trainees initially report finding
these exercises challenging (Segal et  al.,
2013), they result in measurable increases
in volitional orienting of attention (Chan
and Woollacott, 2007; Jha et  al., 2007),
improved performance on sustained attention
tasks (Tang et al., 2015), and improvements
in working memory and autobiographical
memory (Lao et al., 2016). Also, the differ-
ential thickness in brain regions associated
with attention, interoception, and sensory
100 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Figure 4.3  Components recognized as differentiated and connected
processing found in meditators compared to
matched controls (Lazar et al., 2005) is con-
sistent with meditators’ increased capacity
for awareness of internal states, particularly
awareness of breathing sensations.
The second insight is that attention can be
regulated to create a more benign or neutral
affect. It builds upon the first insight and can
be experienced in any of the MT exercises;
when attention is directed to arousal-neutral
bodily sensations, such as those of breathing
for example, a more benign affect emerges.
The sensations of breathing provide an acces-
sible and readily available object of attention
that can be unobtrusively turned to as a restful
experience in moments of stress. This facil-
ity, sometimes referred to by clinicians as
‘going to the breath’, is readily established.
In the initial trials of MBSR, a large major-
ity of participants indicated that they attached
high importance to this simple skill to calm
themselves in moments of stress/distress
(Kabat-Zinn, 1987).
This redirection of attention and its affec-
tive result is illustrated in Figure 4.4. It is
distinguished from experiential avoidance,
which involves a compulsive mental (or
physical) turning away from difficult experi-
ences (Hayes et al., 1996).
The principle is consistent with William
James’ cogent remark that experience is what
one pays attention to (James, 1890). The key
role that this effortful focusing of attention
plays in wellbeing-supportive emotion regula-
tion and in self-regulated behavior (Baumeister
and Heatherton, 1996; Kirschenbaum, 1987;
Thayer et  al., 1996) was confirmed in later
experimental studies, and in the findings that
poor cognitive control is associated with many
mental disorders (Snyder and Hankin, 2016).
Rumination, for example, is an indicator of
attention wandering from immediate tasks
and becoming captivated by an uncomfortable
internal narrative.
Supporting this principle, MT has been
shown to reduce rumination (Campbell
et al., 2012) and to increase mood as a result
(Huffziger et al., 2013). More specifically, an
MT exercise focusing on the awareness of
breathing improved mood, meta-awareness,
and mind wandering (Levinson et al., 2014).
These observations are supported by imaging
studies in which mindfulness trainees show
less activation of brain regions involved in
narrative processing of self-relevant stimuli
and greater activation of regions implicated
in ‘experiential’ processing, relative to nov-
ices (Farb et  al., 2007; Lutz et  al., 2016).
 EVOLUTIONARY PSYCHOLOGY AND MINDFULNESS AND MEDITATION 101

Figure 4.4  Attention shifts from differentiated components to arousal-neutral sensations of

breathing

Interestingly, MT appears to reduce reactivity This observing stance is implicit in most MT

to emotional stimuli, which is not found with
relaxation training (Ortner et al., 2007).
The third insight is the use of an observ-
ing perspective toward mental life. This
‘observing self’ (Deikman, 1982) is used to
notice thoughts/images, sensations, and feel-
ings while remaining unaffected by their
content and tone. It is itself a cognitive process,
albeit one used to cultivate a sense of detach-
ment from mental phenomena that previously
had captured attention and created distress.
exercises and is cultivated explicitly by the
practice of silently naming component mental
qualities as they are occurring in awareness.
For example, rather than contending with,
or trying to push away, a harshly self-critical
thought, attention is reoriented from its con-
tent, such as ‘I am a failure’, to the more affect-
neutral reflection ‘This is a thought’.
The strategy is illustrated in Figure 4.5
using the example of a commonly reported
thought during panic attacks.

Figure 4.5  Re-perceiving reduces distress through a perceptual/attentional shift from what
the thought is about – ‘I’m going to pass out’ – to the thought as an event in the mind/
awareness – ‘This is a thought’
102 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Referred to as decentering (Teasdale et al.,
2002) and meta-cognition (Wells, 1999), this
meta-awareness increases with participa-
tion in MT (Carmody et al., 2009; Feldman
et al., 2010; Lao et al., 2016; Levinson et al.,
2014; Teasdale et  al., 2002). It has been
found important in the treatment of depres-
sion (Bieling et al., 2012), and this decreased
cognitive reactivity appears to mediate the
association between mindfulness-based cog-
nitive therapy and decreases in depressive
symptoms (Cladder-Micus et al., 2017).
MINDFULNESS AND THE COGNITIVE
NARRATIVE’S DOUBLE-EDGED
SWORD
We are tribal primates with a powerful capac-
ity to imagine. This wondrous ability envi-
sions possible futures and constructs
narratives of the past beyond simple condi-
tioning. It gives us a powerful advantage in
meeting our human needs and is that which
most clearly separates us from other pri-
mates. The capacity for language that devel-
oped in conjunction with it allows also for
conversations about where dangers are most
likely to be found, how they might best be
met, and the passing of those lessons across
generations – group learning, in other words.
The clear survival upside of imagination
is seen in the development of tools and their
uses, and in the ability to plan and coordinate
future activities and scenarios that would
favor resource procurement and advantageous
mates: to be able to plan for the hunt and what
would be required, as well as how any gains
might be distributed. When bands were small
and less complex, dangers imminent and
immediate, and neural cognitive capacity less
developed, those imaginings were probably
relatively short-lived; life required careful
and continued attention to the senses.
Immediate physical dangers have been
minimized in modern life and sensory moni-
toring has become less necessary. In these
circumstances, attention defaults to sup-
porting our social needs and planning how
they can be met. And as social life becomes
more complex, the goals longer term, and
success and failure more contingent, imag-
ining assumes an increasingly central role.
Essential to this planning is imagining what
could go wrong here, how plans might be
threatened by others or by circumstances,
and the consequences of these in terms of
our own, and our kin’s, future suffering
and possible death. Imagination also allows
for reflections that give rise to questions of
meaning and existential angst.
Each age and culture developed ways and
means of coping with this anxiety that arises
from the necessary uncertainty of our plans,
and of life itself. Some rely on placating and
beseeching imagined beings assumed to con-
trol events. The Greeks began a more rational
and empirical approach to understanding
human nature and events that unfold in the
world, apparent in the teachings of people
such as Plato, Aristotle, and Archimedes.
Around the same time philosophers in India
developed what we would now call psycho-
logical methods of inquiry that focused on
the mind itself. In that sense they are not
descriptions of the world, but mental mod-
els that allow insight into the machinations
of the vehicle through which we apprehend
the world and how it can be self-modified
to alleviate the angst. In its original context,
mindfulness was cultivated as part of a train-
ing system to recognize how mental suffering
is created moment-to-moment in the human
mind and the behaviors and attitudes that
can alleviate it. And while some schools of
Western psychology developed constructs
and principles that parallel some of those
found in Buddhism, they did not develop
training exercises that so systematically
develop their recognition and self-regulation.
In that sense, one of Buddhism’s most impor-
tant contributions has been the practices to
actualize those principles; ones that have
been seamlessly integrated as mindfulness
into existing psychotherapies.
 EVOLUTIONARY PSYCHOLOGY AND MINDFULNESS AND MEDITATION
In our complex and secular societies, the
cognitive activity required to navigate for
success in status, power, and relationships
consumes inordinate amounts of neural band-
width. And the affect associated with this is
evident in ever-increasing rates of anxiety,
depressive disorders, and suicide. It may also
be contributing to increasing rates of addic-
tion and obesity; rates that may make a more
universal healthcare unsustainable. The MT
exercises support recognition of this mental
activity that goes largely unnoticed in daily
life, even as it is affecting wellbeing.
To those ends, mindfulness encourages
the recognition that our apparently seamless
mental activity comprises phenomenological
components: sensations, cognitions, and their
pleasant/unpleasant affective quality that we
don’t normally experience as separate. The
exercises also develop a capacity to notice
where attention is focused in that suite and to
regulate attention so that it becomes less con-
ditioned and more fluid. The exercises also
expose attention’s default impulse toward
cognition and the narratives forming through
imagination and memory, its handmaiden.
As a needs-serving mechanism, cogni-
tion has a watchful quality for threats and
opportunities. In the mental background, the
default mode network and related functions
are planning and wondering what could go
wrong here and whether this is an opportu-
nity. MT makes this vigilant quality appar-
ent, as well as its affective downside in the
uncomfortable sensations of constriction that
naturally accompany it. It also reveals how
the system relaxes as interest and attention
are re-oriented to just sensation: an ease that
is associated with reduced neural monitoring
activity and downstream changes in arousal-
related biomarkers.
And while MT offers the opportunity for
profound insight, people come to it with
varying levels of interest and curiosity so
that each gets off at their own stop along the
route. In the clinical and secular settings into
which mindfulness has been introduced it
is valued primarily for the palliative effects
103
that the practice provides: I want to feel less
­anxious or less depressed, I want to sleep bet-
ter, or I’d like moments of quiet in the tumult
of modern life. The relief that accompanies
these recognitions satisfies the interest of
most people.
For those sufficiently interested to continue
exploring the system’s original existential
goal, other mental features become apparent.
Gaps begin to appear between thoughts and in
those moments the still background becomes
apparent. The natural re-emergence of
thoughts reveals how the narrative they form
has become indistinguishable from that still
presence within which they occur (Carmody,
2016). You realize that it has always been
there, just overlain by attention’s ongoing
fascination with the contents of the cognitive
narrative. It also becomes apparent just how
much of our lives are spent distracted and
preoccupied by that imagining. Perspective
shifts with that experience, as it would if a
fish was to recognize water.
REFERENCES
Alderson-Day, B., & Fernyhough, C. (2015).
Inner speech: Development, cognitive func-
tions, phenomenology, and neurobiology.
Psychological Bulletin, 141(5), 931. doi:http://
psycnet.apa.org/doi/10.1037/bul0000021
Andrews-Hanna, J. R. (2012). The brain’s
default network and its adaptive role in
internal mentation. The Neuroscientist,
18(3), 251–270. doi:https://doi.org/10.1177/
1073858411403316b
Baumeister, R. F., Bratslavsky, E., Finkenauer,
C., & Vohs, K. D. (2001). Bad is stronger than
good. Review of General Psychology, 5(4),
323. doi:http://psycnet.apa.org/doi/10.1037/
1089-2680.5.4.323
Baumeister, R. F., & Heatherton, T. F. (1996).
Self-regulation failure: An overview. Psycho-
logical Inquiry, 7(1), 1–15. doi:http://dx.doi.
org/10.1207/s15327965pli0701_1
Bieling, P. J., Hawley, L. L., Bloch, R. T., Corco-
ran, K. M., Levitan, R. D., Young, L. T., &
Segal, Z. V. (2012). Treatment-specific
104 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
changes in decentering following mindfulness-
based cognitive therapy versus antidepres-
sant medication or placebo for prevention of
depressive relapse. Journal of Consulting and
Clinical Psychology, 80(3), 365. doi:http://
psycnet.apa.org/doi/10.1037/a0027483
Biswas-Diener, R., Linley, P. A., Dovey, H.,
Maltby, J., Hurling, R., Wilkinson, J., & Lyub-
chik, N. (2015). Pleasure: An initial explora-
tion. Journal of Happiness Studies, 16(2),
313–332. doi:https://doi.org/10.1007/
s10902-014-9511-x
Blair, C. (2002). School readiness: Integrating
cognition and emotion in a neurobiological
conceptualization of children’s functioning at
school entry. American Psychologist, 57(2),
111. doi:http://psycnet.apa.org/doi/10.1037/
0003-066X.57.2.111
Brosschot, J. F., Gerin, W., & Thayer, J. F. (2006).
The perseverative cognition hypothesis: A
review of worry, prolonged stress-related
physiological activation, and health.
Journal of Psychosomatic Research, 60(2),
113–124.
Brosschot, J., Geurts, S., Kruizinga, I., Radstaak,
M., Verkuil, B., Quirin, M., & Kompier, M.
(2014). Does unconscious stress play a role in
prolonged cardiovascular stress recovery?
Stress and Health: Journal of the Interna-
tional Society for the Investigation of Stress,
30(3), 179. doi:10.1002/smi.2590
Campbell, T. S., Labelle, L. E., Bacon, S. L.,
Faris, P., & Carlson, L. E. (2012). Impact of
mindfulness-based stress reduction (MBSR)
on attention, rumination and resting blood
pressure in women with cancer: A waitlist-
controlled study. Journal of Behavioral Medi-
cine, 35(3), 262–271. doi:https://doi.
org/10.1007/s10865-011-9357-1
Carmody, J. (2016). Fish Discovering Water:
Meditation as a Process of Recognition. In M.
West (Ed.), The Psychology of Meditation.
Oxford University Press.
Carmody, J., & Baer, R. A. (2008). Relationships
between mindfulness practice and levels of
mindfulness, medical and psychological
symptoms and well-being in a mindfulness-
based stress reduction program. Journal of
Behavioral Medicine, 31(1), 23–33. doi:https://
doi.org/10.1007/s10865-007-9130-7
Carmody, J., Baer, R. A., Lykins, E. L. B., &
Olendzki, N. (2009). An empirical study of the
mechanisms of mindfulness in a mindfulness-
based stress reduction program. Journal of
Clinical Psychology, 65(6), 613–626.
doi:10.1002/jclp.20579
Cavanagh, K., Strauss, C., Forder, L., & Jones, F.
(2014). Can mindfulness and acceptance be
learnt by self-help? A systematic review and
meta-analysis of mindfulness and acceptance-
based self-help interventions. Clinical Psychol-
ogy Review, 34(2), 118–129. doi:https://doi.
org/10.1016/j.cpr.2014.01.001
Cebolla, A., Campos, D., Galiana, L., Oliver, A.,
Tomás, J. M., Feliu-Soler, A., & Baños, R. M.
(2017). Exploring relations among mindful-
ness facets and various meditation practices:
Do they work in different ways? Conscious-
ness and Cognition, 49, 172–180. doi:https://
doi.org/10.1016/j.concog.2017.01.012
Chan, D., & Woollacott, M. (2007). Effects of
level of meditation experience on attentional
focus: Is the efficiency of executive or orien-
tation networks improved? The Journal of
Alternative and Complementary Medicine,
13(6), 651–658. doi:10.1089/acm.2007.7022
Cladder-Micus, M., van Aalderen, J., Donders,
A., Spijker, J., Vrijsen, J., & Speckens, A.
(2017). Cognitive reactivity as outcome and
working mechanism of mindfulness-based
cognitive therapy for recurrently depressed
patients in remission. Cognition and Emo-
tion, 1–8. doi:http://dx.doi.org/10.1080/
02699931.2017.1285753
Creswell, J. D., Bursley, J. K., & Satpute, A. B.
(2013). Neural reactivation links unconscious
thought to decision-making performance.
Social Cognitive and Affective Neuroscience,
8(8), 863–869. doi:https://doi.org/10.1093/
scan/nst004
Custers, R., & Aarts, H. (2010). The unconscious
will: How the pursuit of goals operates outside
of conscious awareness. Science, 329(5987),
47–50. doi:10.1126/science.1188595
Dahl, C. J., Lutz, A., & Davidson, R. J. (2015).
Reconstructing and deconstructing the self:
Cognitive mechanisms in meditation practice.
Trends in Cognitive Sciences, 19(9), 515–523.
doi:https://doi.org/10.1016/j.tics.2015.07.001
Damasio, A. R. (2003). Looking for Spinoza: Joy,
sorrow, and the feeling brain. New York:
Harcourt.
Deikman, A. J. (1982). The observing self: Mysti-
cism and psychotherapy. Boston: Beacon Press.
 EVOLUTIONARY PSYCHOLOGY AND MINDFULNESS AND MEDITATION
Donald, J. N., Atkins, P. W., Parker, P. D., Christie,
A. M., & Ryan, R. M. (2016). Daily stress and
the benefits of mindfulness: Examining the
daily and longitudinal relations between pre-
sent-moment awareness and stress responses.
Journal of Research in Personality, 65, 30–37.
doi:https://doi.org/10.1016/j.jrp.2016.09.002
Farb, N. A. S., Segal, Z. V., Mayberg, H., Bean,
J., McKeon, D., Fatima, Z., & Anderson, A. K.
(2007). Attending to the present: Mindful-
ness meditation reveals distinct neural modes
of self-reference. Social Cognitive and Affec-
tive Neuroscience, 2(4), 313–322. doi:https://
doi.org/10.1093/scan/nsm030
Feldman, G., Greeson, J., & Senville, J. (2010).
Differential effects of mindful breathing, pro-
gressive muscle relaxation, and loving-kindness
meditation on decentering and negative reac-
tions to repetitive thoughts. Behaviour
Research and Therapy, 48(10), 1002–1011.
doi:https://doi.org/10.1016/j.brat.2010.06.006
Füstös, J., Gramann, K., Herbert, B. M., &
Pollatos, O. (2013). On the embodiment of
emotion regulation: Interoceptive awareness
facilitates reappraisal. Social Cognitive and
Affective Neuroscience, 8(8), 911–917.
doi:https://doi.org/10.1093/scan/nss089
Geary, D. (2005). The motivation to control
and the origin of mind: Exploring the life-
mind joint point in the Tree of Knowledge
System. Journal of Clinical Psychology, 61(1),
21–46. doi:10.1002/jclp.20089
Goyal, M., Singh, S., Sibinga, E. M., Gould, N.
F., Rowland-Seymour, A., Sharma, R., &
Shihab, H. M. (2014). Meditation programs
for psychological stress and well-being: A
systematic review and meta-analysis. JAMA
Internal Medicine, 174(3), 357–368.
doi:10.1001/jamainternmed.2013.13018
Hayes, S. C., Wilson, K. G., Gifford, E. V.,
Follette, V. M., & Strosahl, K. (1996). Experien-
tial avoidance and behavioral disorders: A
functional dimensional approach to diagnosis
and treatment. Journal of Consulting & Clinical
Psychology, 64(6), 1152–1168. doi:http://
psycnet.apa.org/doi/10.1037/0022-006X.
64.6.1152
Huffziger, S., Ebner-Priemer, U., Eisenbach, C.,
Koudela, S., Reinhard, I., Zamoscik, V., &
Kuehner, C. (2013). Induced ruminative and
mindful attention in everyday life: An
­experimental ambulatory assessment study.
105
Journal of Behavior Therapy and Experimen-
tal Psychiatry, 44(3), 322–328. doi:https://
doi.org/10.1016/j.jbtep.2013.01.007
James, W. (1890). The principles of psychology.
New York: H. Holt.
Jha, A., Krompinger, J., & Baime, M. J. (2007).
Mindfulness training modifies subsystems of
attention. Cognitive Affective and Behavioral
Neuroscience, 7(2), 109–119. doi:https://doi.
org/10.3758/CABN.7.2.109
Kabat-Zinn, J. (1982). An out-patient program
in behavioral medicine for chronic pain
patients based on the practice of mindful-
ness meditation: Theoretical considerations
and preliminary results. General Hospital
Psychiatry, 4, 33–47.
Kabat-Zinn, J. (1987). Four-year follow-up of a
meditation-based program for the self-­
regulation of chronic pain; treatment out-
comes and compliance. Clinical Journal of
Pain, 2, 159–173.
Kabat-Zinn, J. (1990). Full catastrophe living:
Using the wisdom of your body and mind to
face stress, pain and illness. New York:
Delacorte.
Killingsworth, M. A., & Gilbert, D. T. (2010). A
wandering mind is an unhappy mind.
Science, 330(6006), 932. doi:10.1126/science.
1192439
Kirschenbaum, D. S. (1987). Self-regulatory fail-
ure: A review with clinical implications. Clinical
Psychology Review, 7(1), 77–104. doi:https://
doi.org/10.1016/0272-7358(87)90005-5
Lao, S., Kissane, D., & Meadows, G. (2016).
Cognitive effects of MBSR/MBCT: A
systematic review of neuropsychological
outcomes. Consciousness and Cognition,
45, 109–123. doi:https://doi.org/10.1016/
j.concog.2016.08.017
Lazar, S. W., Kerr, C. E., Wasserman, R. H.,
Gray, J. R., Greve, D. N., Treadway, M. T., &
Fischl, B. (2005). Meditation experience is
associated with increased cortical thickness.
NeuroReport 16(17), 1893–1897.
Levinson, D. B., Stoll, E. L., Kindy, S. D., Merry,
H. L., & Davidson, R. J. (2014). A mind you
can count on: Validating breath counting as
a behavioral measure of mindfulness.
Frontiers in Psychology, 5, 1202. doi:https://
dx.doi.org/10.3389%2Ffpsyg.2014.01202
Lutz, J., Brühl, A. B., Doerig, N., Scheerer, H.,
Achermann, R., Weibel, A., & Herwig, U.
106 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
(2016). Altered processing of self-related
emotional stimuli in mindfulness meditators.
Neuroimage, 124, 958–967. doi:https://doi.
org/10.1016/j.neuroimage.2015.09.057
Mehling, W. E., Price, C., Daubenmier, J. J.,
Acree, M., Bartmess, E., & Stewart, A.
(2012). The multidimensional assessment of
interoceptive awareness (MAIA). PloS One,
7(11), e48230.
Ortner, C. N. M., Kilner, S. J., & Zelazo, P. D.
(2007). Mindfulness meditation and reduced
emotional interference on a cognitive task.
Motivation and Emotion, 31(4), 271–283.
doi:https://doi.org/10.1007/s11031-007-
9076-7
Pessoa, L. (2008). On the relationship between
emotion and cognition. Nature Reviews Neu-
roscience, 9(2), 148–158.
Posner, M. I., & Rothbart, M. K. (2007).
Research on attention networks as a model
for the integration of psychological science.
Annual Review of Psychology, 58, 1–23.
doi:https://doi.org/10.1146/annurev.psych.
58.110405.085516
Sapolsky, R. M. (2004). Why zebras don’t get
ulcers: The acclaimed guide to stress, stress-
related diseases, and coping (revised and
updated). New York: Macmillan.
Saunders, B., Rodrigo, A. H., & Inzlicht, M.
(2016). Mindful awareness of feelings increases
neural performance monitoring. Cognitive,
Affective, & Behavioral Neuroscience, 16(1),
93–105. doi:https://doi.org/10.3758/s13415-
015-0375-2
Segal, Z. V., Williams, J. M. G., & Teasdale, J. D.
(2013). Mindfulness-based cognitive therapy
for depression (2nd ed.). New York: Guilford.
Shonin, E., Van Gordon, W., & Griffiths, M. D.
(2015). Does mindfulness work? BMJ, 1,
1–11.
Snyder, H. R., & Hankin, B. L. (2016). Spiraling
out of control: Stress generation and subse-
quent rumination mediate the link between
poorer cognitive control and internalizing
psychopathology. Clinical Psychological Sci-
ence, 4(6), 1047–1064. doi:https://doi.org/
10.1177/2167702616633157t
Tang, Y.-Y., Hölzel, B. K., & Posner, M. I. (2015).
The neuroscience of mindfulness meditation.
Nature Reviews Neuroscience, 16(4), 213–225.
doi:http://dx.doi.org/10.1038/nrn3916
Teasdale, J. D., Moore, R. G., Hayhurst, H.,
Pope, M., Williams, S., & Segal, Z. V. (2002).
Metacognitive awareness and prevention of
relapse in depression: Empirical evidence.
Journal of Consulting and Clinical Psychol-
ogy, 70(2), 275–287. doi:http://psycnet.apa.
org/doi/10.1037/0022-006X.70.2.275
Thayer, J. A., Friedman, B. H., & Borkovec, T. D.
(1996). Autonomic characteristics of gener-
alized anxiety disorder and worry. Biological
Psychiatry, 39(4), 255–266. doi:https://doi.
org/10.1016/0006-3223(95)00136-0
Tomlin, D., Rand, D. G., Ludvig, E. A., & Cohen,
J. D. (2015). The evolution and devolution of
cognitive control: The costs of deliberation in a
competitive world. Scientific Reports, 5, 11002.
doi:https://dx.doi.org/10.1038%2Fsrep11002
Watkins, E. (2004). Appraisals and strategies
associated with rumination and worry.
Personality and Individual Differences, 37(4).
doi:https://doi.org/10.1016/j.paid.2003.10.002
Wells, A. (1999). A meta-cognitive model and
therapy for generalized anxiety disorder. Clini-
cal Psychology and Psychotherapy, 6, 86–95.
doi:10.1002/(SICI)1099-0879(199905)
6:2<86::AID-CPP189>3.0.CO;2-S
Willem, K., Warren, F., Taylor, R., Whalley, B.,
Crane, C., Bondolfi, G., & Schweizer, S.
(2016). Efficacy and moderators of
­mindfulness-based cognitive therapy (MBCT)
in prevention of depressive relapse: An
individual patient data meta-analysis from
randomized trials. JAMA Psychiatry. 73(6),
565–574. doi:http://dx.doi.org/10.1001/
jamapsychiatry.2016.0076

Evolutionary Psychology and
Environmental Sciences
Ulysses Paulino Albuquerque, Joelson M. B. Moura,
Risoneide Henriques da Silva, Washington S.
F e r r e i r a J ú n i o r , a n d Ta l i n e C . S i l v a
INTRODUCTION
An investigative program of evolutionary
psychology must address the origin of human
beings, as the environments that generated
pressure during the evolution of early hominids
form the basis for premises about the evolution
of the human mind. As evidence suggests the
African savanna as the most likely place for the
origin of the modern human being, the savanna
environment usually receives more emphasis
than other environments.
We have succeeded as a species through
mental specializations, also understood as
evolved psychological mechanisms. These
specializations were selected because they
solved problems in paleoenvironments, and
were inherited by subsequent generations
of hominids. For example, when exploring
new landscapes, early hominids needed to
quickly identify potentially dangerous situa-
tions, which trees were climbable, and where
they could shelter. These decisions needed to
be fast, and they were only possible because
5
previously selected mental mechanisms
allowed the assessment of the landscape,
even if unconsciously (see Zajonc, 1980;
Townsend and Barton, 2018).
If we assume that the savanna was the
‘main paleoenvironment’ of our evolution
and that we inherited both physical and men-
tal adaptations of our ancestors, the above-
mentioned argument is coherent. However,
the literature indicates that we evolved along
different lineages and from myriad hominid
groups that coexisted in a wide range of envi-
ronments, and that more than one point of
origin of H. sapiens may have existed (Foley
et  al., 2016; Stringer, 2016). Evolution may
have occurred independently in different
areas, with hominids developing morpholog-
ical substructures that resulted in a complete
set of H. sapiens characteristics. Stringer
(2016) calls this independent evolution
‘African multiregionalism’, characterized by
interfertile subdivisions of H. sapiens in their
evolutionary history across Africa.
108 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
This discussion is central to a program that
attempts to investigate the evolution of the
human mind and human behavior. Discussing
the origin of human beings is essential for
evolutionary psychology and for understand-
ing how our minds work in relation to the
environment and all its components.
THE ORIGIN AND EVOLUTION
OF HUMANS
The transition from dense and closed forests
to the savanna may have occurred slowly.
This suggests that early hominids left the
canopy of forest trees gradually, venturing
into the East African savanna to explore avail-
able resources and to identify hazards and safe
sleeping places. Thus, arboreal behaviors may
have coexisted with bipedal locomotion (see
Townsend and Barton, 2018). Lucy, the most
famous Australopithecus afarensis, had both
bipedal and arboreal habits (Larson, 2012).
Until recently, paleontological data sug-
gested that the first hominids appeared in
Central Africa seven million years ago (Ma)
(see Böhme et  al., 2017). However, recent
evidence suggests that the earliest homi-
nid, Graecopithecus freybergi, lived in a
savanna environment in the region of Greece,
between 7.37 and 7.11 Ma, which is 200,000
years earlier than the previous earliest known
hominid, Sahelanthropus tchadensis, found
in Africa (Böhme et  al., 2017). Likewise,
Homo sapiens was believed to have origi-
nated around 200,000 years ago in South
Africa, but recent fossil evidence suggests
that H. sapiens appeared about 315,000
years ago in Morocco, 100,000 years earlier
than previously thought (Hublin et al., 2017;
Richter et al., 2017). These fossils have a mix
of characteristics of H. sapiens fossils from
other parts of Africa, indicating a multicen-
tric genesis for our species (see Hublin et al.,
2017; Richter et  al., 2017). This finding is
consistent with genetic evidence that the first
divergence of modern human populations
occurred between 350,000 and 260,000 years
ago (Schlebusch et al., 2017). In addition, a
hominid skull, dating back about 436,000–
390,000 years, was recently discovered in the
Cave of Aroeira in Portugal, reinforcing the
idea that human origins did not necessarily
occur in Africa (López-García et  al., 2018).
Although the savanna is still regarded as the
main setting of our evolution, these findings
suggest that the origin and the great divisions
in the family of hominids may have occurred
outside Africa.
Nonetheless, if we understand that estab-
lishment in the savanna was important for the
survival of hominids, it is reasonable to infer
that, over time, natural selection favored indi-
viduals better adapted to savanna conditions.
These individuals inherited the anatomical
and cognitive apparatus evolved in this envi-
ronment and were most likely to survive and
leave offspring. This moment was crucial in
evolutionary history. Many aspects of our
anatomy and current behaviors resulted from
solutions to challenges faced by early homi-
nids (Townsend and Barton, 2018).
Townsend and Barton (2018) argue that
common behaviors and anatomical adapta-
tions of early hominids in the Pleistocene
persist today. For example, the palmar grasp
reflex is a primitive reflex. It consists of a
strong pressure with the hands and represents
the primate’s need to hold the mother’s skin
as she moves through the canopy of the trees.
During childhood, for example, there is a ten-
dency for children to show climbing behaviors
(e.g., climbing trees or climbable objects) that
fit the category of primitive reflex (Townsend
and Barton, 2018). Brachiation is also still
used today by children and gymnasts and
was essential for the hominids in the savanna
paleoenvironments. Perhaps children use tree
climbing as strategy to avoid predators (Coss
and Moore, 2002). Brachiation refers to a
mobility method that depends on the specific
structure of the shoulder to hang on the tree
limb and allow the arm to swing in a com-
plete circle (Townsend and Barton, 2018).
These authors also suggest that the standard
 EVOLUTIONARY PSYCHOLOGY AND ENVIRONMENTAL SCIENCES
size of the human hand is proportional to the
size of tree branches capable of supporting a
human’s weight during a climb. In addition,
humans generally prefer horizontal-branched
trees precisely because it facilitates climbing.
These behaviors can be understood as an
ancestral inheritance of early hominids, with
our cognition, like our anatomy, resulting
from adaptations to the selective pressures
of paleoenvironments (Tooby and Cosmides,
2015). Blome et al. (2012) demonstrated that
the African paleoclimate from 150,000 to
30,000 years ago also displayed regional varia-
tion, so that periods of high aridity or humidity
did not occur simultaneously in the north-
ern, eastern, tropical, and southern regions of
Africa. According to these authors, this cli-
mate heterogeneity may have created oppor-
tunities for hominids to migrate to adjacent
regions. Furthermore, Coulthard et al. (2013)
found that, in humid climates around 100,000
years ago, major African river systems flowed
northward, across the Sahara and to the
Mediterranean Sea. These authors believe that
three now-buried rivers could have been active
in the period of human migration across the
Sahara, with the abundance of water resources
creating viable migratory routes for humans.
Evidence shows that hominids adapted to
various environments in a wide latitudinal
range, such as the temperate and subtropi-
cal north of China and tropical regions of
Southeast Asia (Roberts et  al., 2016; Kong
et al., 2018). The use of fire, which is a prac-
tice frequently described in the literature on
arid environments, has also been observed
in tropical forests (Friesem et  al., 2017). In
addition, traces of foraging activities and
the discovery of tools for hunting arboreal
animals challenge the dominant idea of the
evolutionary adaptation of the first humans to
the arid environment of the savanna (Barker
et al., 2007; Friesem et al., 2017).
If cognitive mechanisms result from
responses to selective pressures of the envi-
ronment, much of our mind may also be
‘trapped’ in evolutionary environments. If
this is true, a challenge for evolutionary
109
psychologists would be to broaden the
­spectrum of the environments in which we
evolved and the influence of the evolved psy-
chological mechanisms in solving problems
different from those found in the savanna. If
the human mind has evolved in response to the
difficulties imposed by the environment, and
if H. sapiens has emerged and evolved in dif-
ferent environments, it is possible that today’s
human behavioral responses, including their
preferences, are a reflection of this multicen-
tric origin. Indeed, the paleontological evi-
dence that hominids inhabited and explored
several environments in the Pleistocene sug-
gests that other psychological mechanisms
may have evolved in periods before or after
establishment in the savanna. For example, a
recent study by our research group has shown
that some people, when analyzing landscapes
of savanna, rainforest, tundra, desert, conif-
erous forest, deciduous forest, and urban
landscape, prefer images of exuberant green
rainforests (Moura et  al., 2018). In addition,
people living in Spain tend to prefer densely
green and closed landscapes (Hartmann and
Apaolaza-Ibáñez, 2010). Because this land-
scape is typical of Spain, and in Brazil there
is great media appeal to preserve the Amazon
rainforest, these findings suggest that recent
stimuli, rather than innate responses, may
exert strong influence on human behavior.
According to Barrett (2012), adaptations
may provide plasticity to the human mind.
They may also integrate mechanisms – whether
more general or more specific – shaped by
evolutionary history with those shaped by
the ontogenetic development of the indi-
vidual. Therefore, our mental mechanisms
may be heterogeneous in origin, with new
structures evolving from older structures and
ancestral features combining with relatively
recent characteristics (Barrett, 2012). Thus,
cognitive adaptations are not necessarily the
result of responses to difficulties imposed by
a specific environment. They might reflect
the selection of general strategies of the
human mind to meet challenges in different
environments.
110 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
THE ENVIRONMENT OF
EVOLUTIONARY ADAPTEDNESS (EEA)
AND THE STRUCTURING OF THE
HUMAN MIND
Understanding the evolutionary environment
of hominids is crucial for evolutionary psy-
chology and other disciplines interested in the
evolution of the human mind. For example,
Bowlby (1982) coined the term Environment
of Evolutionary Adaptedness (EEA) to refer
to the environment that selected the current
genotypes of an organism. According to this
perspective, it is reasonable to suppose that
these environments also influenced the selec-
tion of mental traits of human beings. Frost
(2011) proposed that, for humans, the EEA
would be represented by the African savanna
of the Pleistocene, the environment probably
occupied by early H. sapiens before they
started migrating to other continents about
50,000 years ago. Many authors argue that
human psychological mechanisms evolved in
response to the stable characteristics of the
savanna environments (Tooby and Cosmides,
Figure 5.1  Environment of Evolutionary Adaptedness definition (EEA), original version and
extended version
Source: Created by the authors.
1992, 2005), and the reconstruction of these
selective environments could indicate why
humans have propensities for certain types of
thoughts, motivations, and behaviors (Foley,
1996). However, the previously mentioned
evidence of evolution of hominids in different
areas of the African continent seems to chal-
lenge the savanna hypothesis (see Bolhuis
et al., 2011). Thus, the human EEA may com-
prise a multitude of geographic and temporal
environments (Volk and Atkinson, 2013), that
is, the EEA has become less specific, taking
into account not only the African savanna (see
Tooby and Cosmides, 2015), but also the other
selective environments in which humans have
lived over the course of their evolution. As a
consequence, humans may have developed
psychological mechanisms in environments
that were different from the African savanna
(see Hartmann and Apaolaza-Ibáñez, 2010,
2013; Moura et al., 2018) (Figure 5.1).
Studies on human preference for land-
scapes, for example, provide evidence that
these psychological mechanisms may have
suffered interference from the interaction
 EVOLUTIONARY PSYCHOLOGY AND ENVIRONMENTAL SCIENCES
of humans and different ancestral environ-
ments. Orians (1980) argues that, because the
savanna is an open environment, it enabled
the first hominids a more accurate perception
of approaching predators. This suggests the
evolution of a psychological mechanism in
humans to prefer savanna landscapes – the
savanna hypothesis. However, several studies
have reported preference in humans for envi-
ronments other than African savannas (see
Han, 2007; Hartmann and Apaolaza-Ibáñez,
2010, 2013; Moura et al., 2018).
Some studies have tried to understand how
our species recalls information that is relevant
to survival, providing evidence of how the
human mind may have evolved psychologi-
cal mechanisms that deal with risky situations
in different environments. Yang et  al. (2014)
observed that people in both ancestral survival
scenarios – grasslands and in non-ancestral
or modern environments – recalled important
words in a survival situation. In another study,
Young et al. (2012) found that threats in mod-
ern environments – such as firearms and cars –
capture and maintain attention in the same
way as would be expected for threats in ances-
tral environments, such as snakes and spiders.
These findings lead us to believe that natural
selection favored psychological mechanisms
that deal with challenges regardless of the
type of environment. Thus, human inventions
(e.g., firearms and cars) that are immersed in
the culture and environment may be acting as
a selective force that activates modern psy-
chological mechanisms. This fact seems to
indicate that the human construction of niche1
interferes in own and others’ psychological
mechanisms. This interpretation finds sup-
port in reports that psychological mechanisms
that favor the recall of information relevant to
survival can be observed in people occupying
different contemporary environmental and cul-
tural contexts (see Barrett and Broesch, 2012;
Barrett et al., 2016). For example, Barrett and
Broesch (2012) found that children living in the
city of Los Angeles in California and children
of a village in Shuar in the Ecuadorian Amazon
had high levels of recall when images of and
111
information on the name and diet of dangerous
animals were presented.
Did Evolution Endow Us with a
Naturalistic Mind?
We believe that many of today’s human deci-
sions and behaviors are influenced by the
same psychological mechanisms present in
our ancestors. An example of this would be
the ability to recall information relevant to
survival (see Nairne et  al., 2007), such as
snakes and spiders (see Young et al., 2012).
However, we also believe that some of these
ancestral psychological mechanisms have
adjusted to the adversities of new environ-
ments humans occupied throughout their
evolution. Consequently, derived psychologi-
cal mechanisms evolved from ancestral psy-
chological mechanisms. The ability to recall
information that refers to a risky situation in
a modern environment, such as firearms and
cars, is evidence of a psychological mecha-
nism adjusted to the reality of contemporary
environments (see Young et al., 2012).
Barrett (2012) relativizes the influence of
ancestral environments in the present – in the
case of evolved psychological mechanisms
shaped in these environments – considering
innate cognitive modules as mechanisms spe-
cialized to solve a specific adaptive problem.
However, if adaptations in the brain are analo-
gous to adaptations of the body, such as tissue
types, they are likely to be heterogeneous and
hierarchical (Barrett, 2012). A hierarchical
organization is a feature of systems that evolve
and develop new structures from older struc-
tures. These adaptations are, therefore, a com-
bination of ancestral and recent traits (Barrett,
2012). Thus, mental adaptations can be con-
structed during the ontogenetic development
of each individual (see Barrett, 2012). As
changes in the individual’s social environment
occur, there may be a selection of ‘dormant’
behaviors or preferences that would never or
rarely be generated by the brain if the environ-
ment remained static (see Barrett, 2012).
112 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

A study by Sandry et al. (2013) provided
evidence of the hierarchical organization of
the human mind: these authors demonstrated
that people do not recall adaptive information
in a similar way. By studying the memoriza-
tion of words in different scenarios – survival,
fear and phobia, partner selection, incest
avoidance, detection of cheaters, jealousy,
infidelity, and acquisition and maintenance of
status – they found that the survival scenario
excelled in the number of words remembered
by people when compared to the other sce-
narios (which were also considered adaptive).
If human memory were a non-hierarchical
system, all these psychological mechanisms
should have promoted recall equally.
This evidence suggests that psychologi-
cal mechanisms evolved through processes of
descent with modification, indicating the for-
mation of human cognition by a combination
of ancestral and derived psychological mecha-
nisms (Barrett, 2012). This should make brain
processes highly heterogeneous and possibly
hierarchically organized, with information
organized in human memory according to its
relevance in dealing with imposed environmen-
tal adversity. Then, some ancestral or derived
psychological mechanism, or both simultane-
ously, would be activated (Figure 5.2).
Albuquerque and Ferreira Júnior (2017)
argue that evolution has provided us with
a naturalistic mind that evolved to account

Figure 5.2  Structure of the human naturalist mind: Scheme I shows a cladogram with the
ancestral and derived psychological mechanisms that constitute the human mind. Scheme II
illustrates the ancestral and derivative psychological mechanisms distributed in the human
mind and how they can be activated; the psychological mechanisms are hierarchically
organized according to their relevance to deal with environmental adversity
Source: Created by the authors.
 EVOLUTIONARY PSYCHOLOGY AND ENVIRONMENTAL SCIENCES 113

for myriad and complex relationships and
challenges that the environment poses for
our species. Challenges include what to eat
(e.g., plants and animals), where to look for
food, how to treat diseases or cope with acci-
dents with the resources provided by nature,
where to take refuge, and how to avoid preda-
tors and poisonous animals. The naturalistic
mind, as one of the components of the human
mind, would also result from the numerous
selective pressures of the ancestral or mod-
ern environment to which our species is
subjected. One of the first studies using the
concept of naturalistic mind found that the
human mind favors the recovery and stor-
age of information about diseases and plants
associated with their treatment, when these
diseases are frequent in the social system
(common diseases) or related to previous
experiences of the individual (Silva et  al.,
2019). The authors expected to find that seri-
ous diseases, those normally debilitating or
fatal, would be favored in memory. However,
this was not the case. The modulation of the
frequency of illness with previous experience
suggests that there is, in fact, a hierarchy in
the mind. Interestingly, a similar pattern is
observed in relation to other phenomena,
such as when people deal with environmental
hazards or catastrophes (see Ruin et al., 2007;
Miceli et  al., 2008; Gibbons and Groarke,
2016). This led Ferreira Júnior et al. (2019)
to formulate the Principle of Regularity.
According to this principle, the human mind
is biased, because it is organized based on
the regular events of our experience. Box 5.1
summarizes what we know about the natural-
istic mind.
Does the Past Explain the Present?
Human Beings and Landscapes
Landscapes can be defined as the space of
interaction of people and environment. The
way humans relate to them may reveal strong
evolutionary roots. The Biophilia hypothesis
proposes that people possess an innate

BOX 5.1 Structure and behavior of the human naturalistic mind 

Origin
• The naturalistic mind is the fruit of all selective pressures along the hominid lineage in evolutionary
environments. Thus, evolved psychological mechanisms respond to different environmental challenges,
that is, they are not necessarily tied to a particular environment (such as the Pleistocene savanna).
• Memory, as one of the components of the naturalistic mind, prioritizes content with adaptive bias,
organizing content hierarchically. Thus, information related to environmental survival can be prior-
itized over other adaptive information. This means that ancestral hazards will not necessarily be
prioritized to the detriment of modern hazards.

Physiology
• The naturalistic mind, as shaped during evolution, can lead our species to experience adaptive lags.
However, as cultural responses operate faster than biological evolution, human activities of niche
construction can modulate the existence or not of adaptive lags.
• Possible mental responses generated in the ancestral environment can be modulated by the individual’s
previous experience with a given phenomenon.
• The frequency (regularity) of a given phenomenon biases cognitive processes associated with the
naturalistic mind, so that less common or rare phenomena tend to be neglected unless they are
modulated by previous experience.
114 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
emotional and affective predisposition to
living things, whether they are animals,
plants, or processes (Wilson, 1993). People
tend to prefer images of natural environments
to urban environments and these images are
more likely to be preferred when they contain
trees (Ulrich, 1993). In addition, humans pro-
cess visual stimuli from nature more effi-
ciently than from urban environments. This
can elicit favorable feelings and emotions to
natural landscapes (Townsend and Barton,
2018). For example, when contemplating a
landscape, whether urban or natural, people
elicit emotional responses that may lead to
positive or negative attitudes towards that
landscape (see Bargh et al., 1992).
Evidence has shown that some people
appreciate landscapes containing lakes or
rivers (Ulrich, 1983), and feel more freedom
in environments with exuberantly green veg-
etation than in urban landscapes (Hartmann
and Apaolaza-Ibáñez, 2010). These affective
reactions can be the result of aesthetic aspects
of the landscape (e.g., perceived naturalness –
how close a given landscape is to its natural
state – presence of water, complexity) or
of evolutionary aspects, as proposed by the
Biophilia hypothesis (Wilson, 1993; Han,
2007; Ode et al., 2009; Lee and Son, 2017).
The affinity of human beings with living
elements may be the result of the continuing
relationship of hominids with nature dur-
ing their evolutionary history. This affinity
may influence aspects of human cognition
related to the use and management of natu-
ral resources (Albuquerque and Ferreira
Júnior, 2017) as well as emotional responses
and preferences for aesthetic components
of nature. In this case, some natural sce-
narios stand out more than others (Orians
and Heerwagen, 1992). For example, stud-
ies have shown that inhabitants of countries
including Australia, Nigeria, South Africa,
the United States, Estonia, and Italy, among
others, prefer open landscapes with sparse
trees with wide and stratified canopy, which
are characteristic of the African Pleistocene
savanna (see Orians and Heerwagen, 1992;
Sommer, 1997; Summit and Sommer, 1999;
Herzog et al., 2000; Falk and Balling, 2010).
Orians and Heerwagen (1992) suggest that
this preference has an evolutionary origin and
results from the importance of the savanna
environment for hominid survival during the
Pleistocene. The savanna offered the first
humans a set of possibilities (e.g., a pano-
ramic view of the open environment and trees
that were easy to climb) that helped them to
escape from predators, search for food, and
shelter under their canopies (Appleton, 1975;
Orians and Heerwagen, 1992; Townsend and
Barton, 2018). The savanna may have played
a relevant role during human evolution, but
not the most prominent. Since we inhabited
other environments with different challenges
in the Pleistocene, survival strategies for
the savanna have been potentially modified,
improved, and combined with other strate-
gies or even abandoned over time.
According to Tooby and Cosmides (2015),
the period during which we were hunter-­
gatherers in paleoenvironments was crucial
for the evolution of our mind. Evolutionary
processes are slow and need hundreds of
generations to build a highly complex ‘men-
tal’ program. That is, human minds would
still be adapted to the world of our ancestors.
According to these authors, ‘The industrial
revolution – even the agricultural revolution –
is too brief a period to have selected for new
neurocomputational programs of any com-
plexity’ (Tooby and Cosmides, 2015: 19).
People commonly experience an adaptive
delay when facing the challenges of industrial-
ized societies, because these environments are
different from the environment in which we
evolved. For example, the taste for fatty foods
is an adaptive behavior for ancestral environ-
ments, in which fat was scarce, but is non-
adaptive in the current environment because it
increases the incidence of cardiovascular dis-
eases (Cosmides and Tooby, 2003).
The influence of past heritage on the inter-
action of people and landscapes and, conse-
quently, the existence of adaptive lags has
been debated by scholars. Some argue that this
 EVOLUTIONARY PSYCHOLOGY AND ENVIRONMENTAL SCIENCES
argument overlooks evolutionary processes
that have enabled the reproductive success of
humans, such as the ability to adjust to vary-
ing and variable environments in which they
live (see Laland and O’Brien, 2012). Laland
and Brown (2006) argue that humans do not
experience adaptive lags, precisely because
they have the ability to build and rebuild key
components of their environments to suit
their needs. This adaptive capacity of humans
and other organisms is understood as niche
construction, which occurs in response to the
environmental challenges created by their
ancestors (see Lewontin, 1982; Odling-Smee
et al., 2003; Laland and Brown, 2006; Laland
and O’Brien, 2012). For example, even if
there is excessive consumption of fatty foods,
humans create niches to solve this problem,
such as the development of drugs and the
practice of physical exercise.
Due to the cultural and environmental
diversity in which we live and develop, the
preference for landscape among humans is
also diverse. For example, the preference
among the Japanese for landscapes of feudal
gardens in urban centers varies according to
the distance to buildings and also to personal
life experience (Senoglu et al., 2018). Colley
and Craig (2019) observed that, in Scotland,
if people perceive a landscape as natural (i.e.,
with little human intervention), the emotional
attachment to the landscape increases, influ-
encing their preferences. In addition, people
living in China prefer environments with
a balance between wild nature and human
constructions, such as channeled streams in
native vegetation (see Hu et al., 2019). Thus,
recent environmental factors can influence
innate human preferences for landscapes.
How Does Our Mind Deal with
Information about Other Living
Things?
In their interactions with environments,
humans had to deal with dangerous events
that threatened ancestral survival. Learning
115
strategies such as trial and error may have
been important in order to avoid such threats
over time. However, trial and error are not
always advantageous because the learning
costs increase in situations such as contact
with poisonous animals. In this context, strat-
egies that favor the learning of certain infor-
mation from the environment may have been
selected (see Rendell et al., 2011; Barrett and
Broesch, 2012). For example, early hominids
that remembered and quickly learned how to
avoid certain components of the environment
(e.g., dangerous animals) and how to recog-
nize and select natural resources (e.g., fruits)
would have an advantage over others who did
not possess such skills or behaviors.
In the Biophilia hypothesis, Wilson (1993)
proposes that the behaviors of approaching
(biophilia) and avoiding (biophobia) certain
components of the environment may have a
biological, evolutionary basis. These behav-
iors result from natural selection to promote
the survival of humans in their interactions
with the environment (see also Kellert,
1993). Chief among the biophilic interactions
is the demand for food in the ancestral envi-
ronment. Rozin and Todd (2015) argue that,
during human evolution, the need for food
and nutrients demanded more time and cog-
nitive effort than other activities performed
by hominids. Selecting food is critical to the
evolution of the human mind and structuring
the culture, but is not an easy task. This activ-
ity requires caution in avoiding the ingestion
of toxins and other non-nutritive substances.
It was essential to differentiate the toxic
from the nutritious. This has led humans to
specialize over time, through natural selec-
tion (Rozin and Todd, 2015). Although the
human–food interaction during evolution is
a promising way to understand our cognitive
structure – and a matter of great interest to
evolutionary ethnobiology (see Albuquerque
and Ferreira Júnior, 2017) – this subject is
still neglected in the field of evolutionary
psychology (Rozin and Todd, 2015).
Evidence suggests a bias in human mem-
ory towards learning information related to
116 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
dangerous animals, as we have previously
mentioned. Broesch et  al. (2014) evaluated
memory retention of information related to
the danger of animals (e.g., if they were poi-
sonous or not), diet, and habitat in indigenous
young people and adults of the Fiji Islands.
They observed that information about haz-
ard and toxicity was best retained by young
people, whereas adults showed no preferen-
tial retention for this information. In contrast,
other studies have indicated that the use of
images of dangerous animals can increase
the retrieval and retention of information in
adults (Kock et al., 2008; Riaz et al., 2018).
In addition to retrieving information from
memory, humans may possess other charac-
teristics that respond quickly to dangerous
animals. Neurobiological studies have indi-
cated that the amygdala of primates, includ-
ing humans, is able to tune visual areas of
the brain to perceive fear-related stimuli (see
Prokop and Randler, 2018). Studies on visual
attention have shown that dangerous animals,
such as lions and snakes, more quickly cap-
ture and maintain the attention of humans
than non-dangerous animals (Yorzinski et al.,
2014). Infants at five months of age stare
longer at images that schematically represent
a spider than images with random schemata
(Rakison and Derringer, 2008). This may
reflect an evolved response to detect more
quickly and to focus attention on danger-
ous animals (Yorzinski et  al., 2014; Prokop
and Randler, 2018). According to Tooby and
Cosmides (2015), this fact could also explain
modern phobias associated with these ani-
mals. However, these responses are culturally
modulated. For example, Maasai people in
Kenya evaluated lions as aesthetically more
attractive than hyenas (Pinho et  al., 2014).
Lions are of great cultural importance to the
Maasai people (Pinho et al., 2014), suggest-
ing that culture can partly modulate human
responses to dangerous animals.
Aversion to dangerous animals can be
learned quickly by observing the reactions of
other individuals to these animals. In a study
with laboratory-raised monkeys – Macaca
mulatta – Cook and Mineka (1990) showed
that young individuals can quickly acquire
fear of snakes merely by watching fear reac-
tions of other individuals towards these ani-
mals in videos. However, observers were not
afraid of flowers after watching edited videos
displaying individuals who were afraid of
these items. This suggests that fear is more
quickly learned when directed at danger-
ous animals. Similarly, babies aged seven to
18 months paid more attention to snake
images when they were associated with a
frightened human voice than with a cheerful
voice (DeLoache and LoBue, 2009). Such
learning may have been important for the sur-
vival of early hominids, as individuals would
not need direct experience with dangerous
animals to acquire the behavior of avoiding
these animals.
Adaptive mechanisms are also activated in
relation to plants. Prokop and Fančovičová
(2014) investigated children exposed to infor-
mation on toxic and non-toxic plants associ-
ated with fruit images of different colors,
i.e., red and black for toxic plants and green
for non-toxic plants. They found that plant
information associated with red- and black-
colored fruits was better retained in children’s
memory, possibly due to their association
with toxic fruits. A recent study showed that,
in a visual detection task, toxic plants were
detected significantly sooner than non-toxic
plants by humans (Prokop and Fančovičová,
2019). The ability to recall information about
plant toxicity may have given humans the
ability to identify and avoid foods potentially
harmful to their survival.
In addition to the behavior of aversion,
people also exhibit behaviors that promote
contact and interaction with animals and
plants (biophilia). People have positive
emotional responses towards animals with
certain characteristics (Prokop and Randler,
2018). Martín-López et al. (2008) conducted
a meta-analysis of 60 studies and showed that
people are more likely to pay for animal con-
servation due to anthropocentric rather than
scientific factors. Anthropocentric factors
 EVOLUTIONARY PSYCHOLOGY AND ENVIRONMENTAL SCIENCES
include animal characteristics preferred by
people, such as length, weight, and eye size.
Regarding plants, some studies have shown
that people prefer landscapes of trees with
larger canopies and shorter trunks (for a
review, see Townsend and Barton, 2018).
It seems this preference may be produced
by psychological adaptation. Individuals
who had positive sensations in response to
these trees were selected because these trees
offered them safety and shelter (Townsend
and Barton, 2018).
The degree of interaction of people and
nature may promote positive behaviors
directed at other animals. A study in China
found that children living in rural areas and
having more contact with nature were more
likely to protect and like animals (biophilia)
than children from urbanized regions (Zhang
et  al., 2014). Zhang and colleagues suggest
that contact of humans with nature can help
to promote conservation strategies. Sampaio
et  al. (2018) observed that the contact of
children with forests influenced their knowl-
edge of local biodiversity. The children were
encouraged to express their knowledge as
drawings, and the authors found that chil-
dren who had more contact with forests also
had greater knowledge about native animals
of the region. According to the authors, the
proximity of children to the forest drew atten-
tion to the components of this environment,
laying the foundations for the construction
of knowledge. This fact indicates that the
environment in which human beings live
can influence their cognition, and is respon-
sible for promoting and mediating human
behaviors, such as being prone or not to pre-
serve nature.
FUTURE CHALLENGES FOR
ENVIRONMENTAL SCIENCES
Penn (2003) offered a synthesis of a number
of evolutionary approaches that provide
insight into human nature and its role
117
in current environmental events. For Penn,
population growth is one of the major current
ecological issues – there are around seven
billion people on the planet – and creating
effective public policies to stabilize this
growth requires understanding the evolution-
ary roots of the problem. From an evolution-
ary perspective, reproductive self-regulation
should be expected as the high demographic
rate disturbs population and environmental
balance – as advocated by the Demographic
Transition Theory – as this adaptive response
provides the best chance for survival (see
recent discussion in Brooks et  al., 2019;
Salvati et al., 2019). However, in some tradi-
tional societies, reproductive success is posi-
tively associated with wealth increase,
whereas in rich developed countries, fertility
tends to fall as their people opt to have fewer
children to improve their quality of life
(Penn, 2003). Therefore, more studies are
necessary, analyzing the influence of evolu-
tionary and cultural factors on current demo-
graphic dynamics.
Another aspect to be considered when
developing policies to deal with long-term
environmental threats is discounting the
future, that is, the limitations humans have
in considering environmental problems that
may arise in a distant future, putting more
emphasis on the present day (Penn, 2003;
Henry et  al., 2017). In this case, natural
selection may have favored hominids that
discounted the future, as life expectancy
was relatively short and the future uncer-
tain, making it crucial to focus on present
needs, which increased individual survival
and reproductive success (Penn, 2003). Thus,
a good conservation strategy would be to
associate time discount rates to nature con-
servation policies, as that may provide more
realistic expectations of human response to
these policies (Henry et al., 2017).
Moreover, humans tend to use natural
resources according to their own interests,
putting societal interests in second place,
potentially leading to resource exhaustion
(Penn, 2003). This kind of behavior was
118 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
proposed by Hardin (1968) as the tragedy
of the commons. For instance, a study by
Scheiter et al. (2019) showed that, in African
rural savannas, people use open fields for
pasture or hunting intensively, exceeding the
optimum level. These authors propose this as
an example of the tragedy of the commons
that will compromise ecosystem services in
the future. However, human populations
are able to adapt and to develop rules to effec-
tively manage common resources, avoid-
ing the tragedy of the commons (for a more
complete argument, see Šestáková and
Plichtová, 2019).
In this sense, it is essential to understand
the constraints and amplitude of the influence
of the evolved psychological mechanisms
in the relations of modern humans and the
environment. Evolutionary psychology is
interested in better understanding not only
environmental problems and challenges
(having as background the attitudes of human
beings) but also how we model our multiple
relationships with nature. Penn (2003)
argues that we are still moving towards an
understanding of environmental problems
from an evolutionary perspective. In this
chapter, we presented a brief overview of
how evolved psychological mechanisms help
to understand modern humans. However,
just as we move slowly in understanding
the environmental challenges generated by
human activity, our understanding of other
aspects of the relationship between human
beings and the environment is still in its
infancy.
We believe that dialogue with other fields
in science can be fruitful for a better under-
standing, from an evolutionary point of view,
of the relationship between humans and the
environment. An evolutionary perspective
is not exclusive but provides an alternative
or additional point of view. Evolutionary
ethnobiology is a newly systematic science
(Albuquerque and Ferreira Júnior, 2017) that
shares this interest in understanding the eco-
logical and evolutionary history behind our
relations with the environment.
ACKNOWLEDGMENTS
This study was financed in part by the
Coordenação de Aperfeiçoamento de Pessoal
de Nível Superior – Brasil (CAPES) –
Finance Code 001. We thank the CNPq for
the productivity grant awarded to UPA.
Note
1  The process where organisms, through their
activities and decisions, modify their own and
other’s environments. The environmental modifi-
cations generated by the niche constructors influ-
ence the selective pressures of their environment
and can lead to changes in their metabolic, physi-
ological, and behavioral activities (Odling-Smee
et  al., 2003; Laland and O’Brien, 2012; Flynn
et al., 2013; Matthews et al., 2014).
REFERENCES
Albuquerque, U.P. and Ferreira Júnior, W.S.,
2017. What do we study in evolutionary
ethnobiology? Defining the theoretical basis
for a research program. Evolutionary Biology,
44(2), pp.206–15.
Appleton, J., 1975. The Experience of
Landscape. London: John Wiley & Sons.
Barrett, H.C., 2012. A hierarchical model of the
evolution of human brain specializations.
PNAS, 109(1), pp.10733–40.
Barrett, H.C. and Broesch, J., 2012. Prepared
social learning about dangerous animals in
children. Evolution and Human Behavior, 33,
pp.499–508.
Barrett, H.C., Peterson, C.D. and Frankenhuis,
W.E., 2016. Mapping the cultural learnability
landscape of danger. Child Development,
87(3), pp.770–81.
Bargh, J.A., Chaiken, S., Govender, R. and
Pratto, F., 1992. The generality of the
automatic attitude activation effect. Journal
of Personality and Social Psychology, 62(6),
pp.893–912.
Barker, G., Barton, H., Bird, M., Daly, P., Datan, I.,
Dykes, A., Farr, L., Gilbertson, D., Harrisson, B.,
Hunt, C., Higham, T., Kealhofer, L., Krigbaum,
J., Lewis, H., MacLaren, S., Paz, V., Pike, A.,
 EVOLUTIONARY PSYCHOLOGY AND ENVIRONMENTAL SCIENCES
Piper, P., Pyatt, B., Rabett, R., Reynolds, T.,
Rose, J., Rushworth, G., Stephens, M.,
Stringer, C., Thompson, J. and Turney, C.,
2007. The ‘human revolution’ in lowland
tropical Southeast Asia: The antiquity and
behaviour of anatomically modern humans
at Niah Cave (Sarawak, Borneo). Journal of
Human Evolution, 52(3), pp.243–61.
Blome, M.W., Cohen, A.S., Tryon, C.A., Brooks,
A.S. and Russell, J., 2012. The environmental
context for the origins of modern human
diversity: A synthesis of regional variability in
African climate 150,000–30,000 years ago.
Journal of Human Evolution, 62, pp.563–92.
Böhme, M., Spassov, N., Ebner, M. and
Geraads, D., 2017. Messinian age and
savannah environment of the possible
hominin Graecopithecus from Europe. PLoS
ONE, 12(5), pp.1–31.
Bolhuis, J.J., Brown, G.R., Richardson, R.C. and
Laland, K.N., 2011. Darwin in mind: New
opportunities for evolutionary psychology.
PLoS Biology, 9, pp.1–8.
Bowlby, J., 1982. Loss: Sadness and Depression.
New York: Basic Book Publishers.
Broesch, J., Barrett, H.C. and Henrich, J., 2014.
Adaptive content biases in learning about
animals across the life course. Human
Nature, 25, pp.181–99.
Brooks, D.J., Brooks, S.G., Greenhill, B.D. and
Haas, M.L., 2019. The demographic transition
theory of war: Why young societies are conflict
prone and old societies are the most peaceful.
International Security, 43(3), pp.53–95.
Colley, K. and Craig, T., 2019. Natural places:
Perceptions of wildness and attachment to
local greenspace. Journal of Environmental
Psychology, 61, pp.71–8.
Cook, M. and Mineka, S., 1990. Selective asso-
ciations in the observational conditioning of
fear in rhesus monkeys. Journal of Experi-
mental Psychology, 16, pp.372–89.
Cosmides, L. and Tooby, J., 2003. Evolutionary
psychology: Theoretical foundations. In: L.
Nadel (Ed.). Encyclopedia of Cognitive Science
(pp.54–64). London: Macmillan.
Coss, R.G. and Moore, M., 2002. Precocious
knowledge of trees as antipredator refuge
in preschool children: An examination of
aesthetics, attributive judgments, and relic
sexual dinichism. Ecological Psychology, 14,
pp.181–222.
119
Coulthard, T.J., Ramirez, J.A., Barton, N., Rog-
erson, M. and Brucher, T., 2013. Were rivers
flowing across the Sahara during the last
interglacial? Implications for human migra-
tion through Africa. PLoS ONE, 8(9), e74834.
DeLoache, J.S. and LoBue, V., 2009. The
narrow fellow in the grass: Human infants
associate snakes and fear. Developmental
Science, 12, pp.201–7.
Falk, J.H. and Balling, J.D., 2010. Evolutionary
influence on human landscape preference.
Environment and Behavior, 42(4),
pp.479–93.
Ferreira Júnior, W.S., Medeiros, P.M. and
Albuquerque, U.P., 2019. Evolutionary
ethnobiology. Encyclopedia of Life Sciences,
pp.1–6. John Wiley & Sons, Ltd.
Foley, R., 1996. The adaptive legacy of human
evolution: A search for the environment of
evolutionary adaptedness. Evolutionary
Anthropology, 4(6), pp.194–203.
Foley, R.A., Martin, L., Lahr, M.M. and
Stringer, C., 2016. Major transitions in human
evolution. Philosophical Transactions Royal
Society B, 371(1698), pp.1–8.
Friesem, D.E., Lavi, N., Madella, M., Boaretto,
E., Ajithparsad, P. and French, C., 2017.
The formation of fire residues associated
with hunter-gatherers in humid tropical envi-
ronments: A geo-ethnoarchaeological per-
spective. Quaternary Science Reviews, 171(1),
pp.85–99.
Frost, P., 2011. Human nature or human natures?
Futures, 43, pp.740–8.
Gibbons, A. and Groarke, A., 2016. Can risk
and illness perceptions predict breast cancer
worry in healthy women? Journal of Health
Psychology, 21(9), pp.1–11.
Han, K.T., 2007. Responses to six major terres-
trial biomes in terms of scenic beauty, prefer-
ence, and restorativeness. Environment and
Behavior, 39(4), pp.529–56.
Hardin, G., 1968. The tragedy of the com-
mons. Science, 162(3859), pp.1243–8.
Hartmann, P. and Apaolaza-Ibáñes, V., 2010.
Beyond savanna: An evolutionary and envi-
ronmental psychology approach to behavioral
effects of nature scenery in green advertising.
Journal of Environmental Psychology, 30(1),
pp.119–28.
Hartmann, P. and Apaolaza-Ibáñes, V., 2013.
Desert or rain: Standardisation of green
120 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
advertising versus adaptation to the target
audience’s natural environment. European
Journal of Marketing, 47(5/6), pp.917–33.
Henry, A.D., Christensen, A.E., Hofmann, R.,
Steimanis, I. and Vollan, B., 2017. Influence of
sea level rise on discounting, resource use and
migration in small-island communities: An
agent-based modelling approach. Environ-
mental Conservation, 44(4), pp.381–8.
Herzog, T.R., Herbert, E.J., Kaplan, R. and
Crooks, C.L. 2000. Cultural and develop-
mental comparisons of landscape percep-
tions and preferences. Environment and
Behavior, 32(3), pp.323–46.
Hu, S., Yue, H. and Zhou, Z., 2019. Preferences
for urban stream landscapes: Opportunities
to promote unmanaged riparian vegetation.
Urban Forestry & Urban Greening, 38,
pp.114–23.
Hublin, J.J., Ben-Ncer, A., Bailey, S.E., Freidline,
S.E., Neubauer, S., Skinner, M.M., Bergmann,
I., Le Cabec, A., Benazzi, S., Harvati, K. and
Gunz, P., 2017. New fossils from Jebel Irhoud,
Morocco and the pan-African origin of Homo
sapiens. Nature, 546(7657), pp.289–92.
Kellert, S.R., 1993. The biological basis for
human values of nature. In: S.R. Kellert and
E.O. Wilson (Eds.). The Biophilia Hypothesis.
Washington, DC: Island Press. Pp. 42–69.
Kock, N., Chatelain-Jardón, R. and Carmona,
J., 2008. An experimental study of simulated
web-based threats and their impact on
knowledge communication effectiveness.
IEEE Transactions on Professional Communi-
cation, 51, pp.183–97.
Kong, Y., Deng, C., Liu, W., Wu, X., Pei, S.,
Sun, L., Ge, J., Yi, L. and Zhu, R., 2018. Mag-
netostratigraphic dating of the hominin
occupation of Bailong Cave, central China.
Scientific Reports, 8(9699), pp.1–12.
Laland, K.N. and Brown, G.R., 2006. Niche
construction, human behavior, and the
adaptive-lag hypothesis. Evolutionary
Anthropology: Issues, News, and Reviews,
15(3), pp.95–104.
Laland, K.N. and O’Brien, M.J., 2012. Cultural
niche construction: An introduction. Biologi-
cal Theory, 6(3), pp.191–2.
Larson, S., 2012. Did Australopiths climb trees?
Science 338(6106), pp.478–9.
Lee, K. and Son, Y., 2017. Exploring landscape
perceptions of Bukhansan national park
according to the degree of visitors. Sustain-
ability, 9(8), pp.1–27.
Lewontin, R.C., 1982. Organism and environ-
ment. In: H.C. Plotkin (Ed.). Learning, Devel-
opment and Culture (pp.151–70). New York:
John Wiley & Sons.
López-García, L.M., Blain, H., Sanz, M., Daura, J.
and Zilhão, J., 2018. Refining the environ-
mental and climatic background of the Middle
Pleistocene human cranium from Gruta da
Aroeira (Torres Novas, Portugal). Quaternary
Science Reviews, 200, pp.367–75.
Martín-López, B., Montes, C. and Benayas, J.,
2008. Economic valuation of biodiversity
conservation: The meaning of numbers.
Conservation Biology, 22, pp.624–35.
Matthews, B., De Meester, L., Jones, C.G., Ibel-
ings, B.W., Bouma, T.J., Nuutinen, V., Koppel,
J.V. and Odling-Smee, J., 2014. Under niche
construction: An operational bridge between
ecology, evolution, and ecosystem science.
Ecological Monographs, 84, pp.245–63.
Miceli, R., Sotgiu, I. and Settanni, M., 2008.
Disaster preparedness and perception of
flood risk: A study in an alpine valley in Italy.
Journal of Environmental Psychology, 28(2),
pp.164–73.
Moura, J.M.B., Ferreira Junior, W.S., Silva, T.C.
and Albuquerque U.P., 2018. The influence
of the evolutionary past on the mind: An
analysis of the preference for landscapes in
the human species. Frontiers in Psychology,
9(2485), pp.1–13.
Nairne, J.S., Thompson, S.R. and Pandeirada,
J.N.S., 2007. Adaptive memory: Survival pro-
cessing enhances retention. Journal of Exper-
imental Psychology: Learning, Memory, and
Cognition, 33(2), pp.263–73.
Ode, A., Fry, G., Tveit, M.S., Messager, P. and
Miller, D., 2009. Indicators of perceived nat-
uralness as drivers of landscape preference.
Journal of Environmental Management,
90(1), pp.375–83.
Odling-Smee, J., Laland, K.N. and Feldman,
M.W., 2003. Niche Construction: The
Neglected Process in Evolution. Princeton,
NJ: Princeton University Press.
Orians, G.H., 1980. Habitat selection: General
theory and applications to human behavior.
In: J. Lockard (Ed.). The Evolution of Human
Social Behavior (pp.49–66). Chicago, IL:
Elsevier.
 EVOLUTIONARY PSYCHOLOGY AND ENVIRONMENTAL SCIENCES
Orians, G.H. and Heerwagen, J.H., 1992.
Evolved responses to landscapes. In: J.H.
Barkow, L. Cosmides and L. Tooby (Eds.). The
Adapted Mind: Evolutionary Psychology and
the Generation of Culture (pp.555–79). New
York: Oxford University Press.
Penn, D.J., 2003. The evolutionary roots of our
environmental problems: Toward a Darwin-
ian ecology. The Quarterly Review of Biology,
78(3), pp.275–301.
Pinho, J.R., Grilo, C., Boone, R.B., Galvin, K.A.
and Snodgrass, J.G., 2014. Influence of
aesthetic appreciation of wildlife species on
attitudes towards their conservation in Kenyan
agropastoralist communities. PLoS ONE,
9:e88842.
Prokop, P. and Fančovičová, J., 2014. Seeing
coloured fruits: Utilization of the theory of
adaptive memory in teaching botany. Journal
of Biological Education, 3(48), pp.127–32.
Prokop, P. and Fančovičová, J., 2019. The per-
ception of toxic and non-toxic plants by chil-
dren and adolescents with regard to gender:
Implications for teaching botany. Journal of
Biological Education, 53, pp. 463–473.
Prokop, P. and Randler, C., 2018. Biological
predispositions and individual differences in
human attitudes toward animals. In: U.P.
Albuquerque and R.R.N. Alves (Eds.). Ethno-
zoology: Animals in our Lives (pp. 447–66).
Oxford: Elsevier Academic Press.
Rakison, D.H. and Derringer, J., 2008. Do
infants possess an evolved spider-detection
mechanism? Cognition, 107, pp.381–93.
Rendell, L., Fogarty, L., Hoppitt, W.J.E., Morgan,
T.J.H., Webster, M.M. and Laland, K,N., 2011.
Cognitive culture: Theoretical and empirical
insights into social learning strategies. Trends
in Cognitive Sciences, 15, pp.68–76.
Riaz, A., Gregor, S. and Lin, A., 2018. Biophilia
and biophobia in website design: Improving
internet information dissemination. Informa-
tion & Management, 55, pp.199–214.
Richter, D., Grün, R., Joannes-Boyau, R., Steele,
T.E., Amani, F., Rué, M., Fernandes, P.,
Raynal, J., Geraads, D., Ben-Ncer, A., Hublin,
J. and McPherron, S.P., 2017. The age of the
hominin fossils from Jebel Irhoud, Morocco,
and the origins of the Middle Stone Age.
Nature, 546(7657), pp.293–6.
Roberts, P., Boivin, N., Lee-Thorp, J., Petraglia,
M. and Stock, J., 2016. Tropical forests and
121
the genus Homo. Evolutionary Anthropology:
Issues, News, and Reviews, 25(6), pp.306–17.
Rozin, P. and Todd, P.M., 2015. The evolution-
ary psychology of food intake and choice.
In: D.M. Buss (Ed.). The Handbook of Evolu-
tionary Psychology (pp.183–205). Hoboken,
NJ: John Wiley & Sons.
Ruin, I., Gaillard, J.C. and Lutoff, C., 2007.
How to get there? Assessing motorists’ flash
flood risk perception on daily itineraries.
Environmental Hazards, 7(3), pp.235–44.
Salvati, L., Carlucci, M., Serra, P. and Zambon,
I., 2019. Demographic transitions and socio-
economic development in Italy, 1862–2009:
A brief overview. Sustainability (Switzerland),
11(1), pp.1–12.
Sampaio, M.B., De La Fuente, M.F.,
Albuquerque, U.P., Souto, A.S. and Schiel,
N., 2018. Contact with urban forests greatly
enhances children’s knowledge of faunal
diversity. Urban Forestry & Urban Greening,
30, pp.56–61.
Sandry, J., Trafimow, D., Marks, M.J. and Rice,
S., 2013. Adaptive memory: Evaluating alter-
native forms of fitness-relevant processing in
the survival processing paradigm. PLoS ONE,
8(4), e60868.
Scheiter, S., Schulte, J., Pfeiffer, M., Martens,
C., Erasmus, B.F.N. and Twine, W.C., 2019.
How does climate change influence the eco-
nomic value of ecosystem services in savanna
rangelands? Ecological Economics, 157,
pp.342–56.
Schlebusch, C.M., Malmström, H., Günther, T.,
Sjödin, P., Coutinho, A., Edlund, H., Munters,
A.R., Vicente, M., Steyn, M., Soodyall, H.,
Lombard, M. and Jakobsson, M., 2017.
Southern African ancient genomes estimate
modern human divergence to 350,000 to
260,000 years ago. Science, 358(6363),
pp.652–5.
Senoglu, B., Oktay, H.E. and Kinoshita, I., 2018.
An empirical research study on prospect–
refuge theory and the effect of high-rise
buildings in a Japanese garden setting. City,
Territory and Architecture, 5(3), pp.1–16.
Šestáková, A. and Plichtová, J., 2019. Contem-
porary commons: Sharing and managing
common-pool resources in the 21st century.
Human Affairs, 29(1), pp.74–86.
Silva, R.H., Ferreira Júnior, W.S., Medeiros, P.M.
and Albuquerque, U.P., 2019. Adaptive
122 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
memory and evolution of the human natural-
istic mind: Insights from the use of medicinal
plants. PLoS ONE, 14(3), pp.1–15.
Sommer, R., 1997. Further cross-national
studies of tree form preferences. Ecological
Psychology, 9(2), pp.153–60.
Stringer, C., 2016. The origin and evolution of
Homo sapiens. Philosophical Transactions
Royal Society B, 371(1698), pp.1–12.
Summit, J. and Sommer, R., 1999. Further
studies of preferred tree shapes. Environment
and Behavior, 31(4), pp.550–76.
Tooby, J. and Cosmides, L., 1992. The psycho-
logical foundations of culture. In: J. Barkow,
L. Cosmides and J. Tooby (Eds.). The Adapted
Mind: Evolutionary Psychology and the Gen-
eration of Culture (pp.19–136). New York:
Oxford University Press.
Tooby, J. and Cosmides, L., 2005. Conceptual
foundations of evolutionary psychology.
In: D.M. Buss (Ed.). The Handbook of Evolu-
tionary Psychology (pp.5–67). Hoboken, NJ:
John Wiley & Sons.
Tooby, J. and Cosmides, L., 2015. The theoreti-
cal foundations of evolutionary psychology.
In: D.M. Buss (Ed.). The Handbook of Evolu-
tionary Psychology (pp.3–87). Hoboken, NJ:
John Wiley & Sons.
Townsend, J.B. and Barton, S., 2018. The
impact of ancient tree form on modern land-
scape preferences. Urban Forestry and Urban
Greening, 34, pp.205–16.
Ulrich, R.S., 1993. Biophilia, biophobia, and
natural landscapes. In: S.R. Kellert and E.O.
Wilson (Eds.). The Biophilia Hypothesis.
Washington, DC: Island Press. pp. 73–137.
Volk, A.A., and Atkinson, J.A., 2013. Infant
and child death in the human environment
of evolutionary adaptation. Evolution and
Human Behavior, 34, pp. 182–93.
Wilson, E.O., 1993. Biophilia and the conserva-
tion ethic. In: S.R. Kellert and E.O. Wilson
(Eds.). The Biophilia Hypothesis. Washington,
DC: Island Press. pp. 31–41.
Yang, L., Lau, K.P.L. and Truong, L., 2014. The
survival effect in memory: Does it hold into
old age and non-ancestral scenarios? PLoS
ONE, 9(5), pp.1–9.
Yorzinski, J.L., Penkunas, M.J., Platt, M.L. and
Coss, R.G., 2014. Dangerous animals capture
and maintain attention in humans. Evolution-
ary Psychology, 12, pp.534–48.
Young, S.G., Brown, C.M. and Ambady, N.,
2012. Priming a natural or human-made
environment directs attention to context-
congruent threatening stimuli. Cognition
and Emotion, 26, pp.927–33.
Zajonc, R.B., 1980. Feeling and thinking: Prefer-
ences need no inferences. American Psychol-
ogist, 35(2), pp.151–75.
Zhang, W., Goodale, E. and Chen, J., 2014. How
contact with nature affects children’s biophilia,
biophobia and conservation attitude in China.
Biological Conservation, 177, pp.109–16.

Evolutionary Psychology
and Public Health
INTRODUCTION: PUBLIC HEALTH
AND THE OBESITY PROBLEM
Public health has been defined as ‘the art and
science of preventing disease, prolonging life
and promoting health through the organized
efforts of society’.[1] Public health is an inter-
disciplinary science, which aims to promote
the length and quality of human life through
the prevention and treatment of disease.
Whether relating to physical, mental, or social
health, public health seeks to translate high-
quality research into preventative or curative
practice. The field of public health also recog-
nises a positive value arising from good health,
which incorporates concepts of psychological
and emotional well-being. While selection
does not act to promote longevity, evolution-
ary fitness is profoundly affected by morbidity
and mortality[2] and both health outcomes and
evolutionary fitness are notably predicted by
socioeconomic status in the developed and
developing world.[3,4] While there is a
6
Simon Russell
complex relationship between human fertility,
fecundity, and fitness,[5] good health is recog-
nised as a key dimension of evolutionary fit-
ness[6] and has a greater impact on fitness
than age-specific fertility.[7] However, selec-
tion does not act on health and it is possible to
be a public health success but an evolutionary
failure; one may lead a long and healthy life
but fail to reproduce. Equally, one may be an
evolutionary success and a public health fail-
ure; it is possible to reproduce successfully
from a position of poor health.
Despite public health and evolutionary
success being defined by and measured in
different currencies, applying the princi-
ples of evolution to the discipline of public
health may provide original insight for pol-
icy makers and practitioners. Evolutionary
science has been increasingly applied to
various disciplines within the field of public
health, including reproductive health, immu-
nity, infectious diseases, cancer, and mental
health.[8–11] This chapter explores the util-
ity of applying the principles of evolutionary
124 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
psychology to the most pervasive physical
health problem of the modern world, namely,
non-communicable diseases (NCDs). The
primary focus will be on pathological health
risk behaviours, which greatly enhance the
risk of NCDs. Additionally, a case study
presents the relevance of evolutionary deci-
sion making in health-promotion behaviour,
within the context of vaccinations and protec-
tion from communicable diseases.
Comprising cardiovascular diseases, can-
cers, chronic respiratory diseases, and diabe-
tes, NCDs are preventable but account for 41
million or 71% of global deaths annually.[12]
The main risk factors for NCDs are poor diet,
physical inactivity, tobacco use, and harmful
alcohol use.[13] Harmful alcohol use, smok-
ing, poor diet, and physical inactivity can be
understood as health risk behaviours, which
lead to potentially pathological metabolic
and physiological changes. Poor diets typi-
cally involve consuming energy-dense but
low-nutrient food and drinks. Low nutrition
and overconsumption often co-exist and are
typified by eating foods that are high in fats,
sugars, and salt (HFSS). Overweight and obe-
sity are typical consequences of poor diet and
inactivity and will be considered as key patho-
logical outcomes for the purpose of this chap-
ter. The principles and public health response
discussed here for overweight and obesity are
also potentially relevant to harmful drinking
and smoking, since it will be argued that simi-
lar psychological mechanisms and evolution-
ary strategies underpin and govern multiple
health-risk-taking behaviours (for usage and
definitions, see section ‘Psychological mech-
anisms and evolutionary strategies’).
Overweight and obesity has become per-
haps the most prevalent and burdensome
public health problem of the modern world.
Overweight and obesity rates in adults have
tripled since 1975, and in 2016 there were
1.9 billion (39%) adults globally living with
overweight or obesity.[14] Rates of child-
hood obesity have risen 10-fold over the same
period, and in 2016 there were an estimated
340 million children aged 5–19 years living
with overweight and obesity.[14] Living with
overweight and obesity greatly increases the
risk of Type-2 diabetes, cardiovascular dis-
eases, and various forms of cancer,[15] in
addition to many other physical and mental
health problems.
Obesity can be conceived as both very simple
and as highly complex. A positive energy bal-
ance can be thought of as an equation: energy
in, use by metabolic processes, and energy out.
Even a slight but consistent positive imbalance
would accrue overweight and obesity over
time. Conversely, there is almost nothing we do
in our lives which does not affect our energy
balance and obesity. The Foresight obesity
systems map[16] categorises, maps, and illus-
trates the varied and complex determinants of
obesity. There is limited utility in focussing on
any one part of the map, given that the obe-
sity system is dynamic and any change to one
part of the system will have consequences for
other parts. Public health responses can some-
times be simplistic; interventions may attempt
to adjust one determinant in some way, while
controlling for others, and practitioners and
policy lobbyists can become focused on just
one element of the map. In turn, the public
and the media sometimes misunderstand the
problem and oversimplify what is perceived to
be an effective policy response. Furthermore,
there is no consensus between academics or
civil servants on where the public health focus
should be. Some suggest the problem lies with
‘energy in’, since evidence suggests physical
activity may not help people lose weight, while
others focus mainly on ‘energy out’ given that
physical activity has been shown to attenuate
the risks of NCDs, even for people living with
overweight or obesity.
The causality of obesity has drawn simi-
lar debate in terms of the relative impacts of
our genes and the environment and the inter-
play between the two. Broadly speaking, the
effects of heredity in obesity are likely to be
low, given that obesity has grown rapidly
in genetically stable populations. However,
there is some evidence to suggest that biolog-
ical influences account for large proportions
 EVOLUTIONARY PSYCHOLOGY AND PUBLIC HEALTH
Figure 6.1  The impact of the food supply on the expression of obesityi[19]
of variance in food and satiety responsiveness
and therefore risk of obesity. Yet the more we
understand about genetic susceptibility, the
more the importance of the environment has
been revealed; the expression of susceptibil-
ity to obesity in environments with a limited
food supply is low but becomes hugely ampli-
fied in environments with an abundant supply
(Figure 6.1).[17] Generally, diseases rarely
have simple Mendelian patterns of inherit-
ance; heritability of ill health is more com-
monly the product of interactions between
multiple genes, predispositions and the envi-
ronment, and socioeconomic and cultural
influences.[18] While genetic predispositions
are likely to exacerbate or attenuate the risk
of obesity, the problem is recent relative to
human history and has coincided with indus-
trial and urban modernisation, which provide
the obesogenic platforms from which genetic
susceptibility may be expressed.
MAKING EVOLUTIONARY SENSE OF
THE MODERN WORLD
The majority of people now live in surround-
ings that are very different from the selective
environments in which we evolved. With tech-
125
nological and civil development, many physi-
cal and psychological adaptations that
are rooted in evolutionary time have become
mismatched with modern life. Obesity is per-
haps the most pronounced physical manifesta-
tion of this mismatch; it has increased in almost
every country in the world, and is reported to
be proximally driven by changes in the food
system, where calorie-dense food is abundant,
affordable, and readily available in most of the
modern world.[20] Built environments have
also changed and influence what and how
much we eat, in addition to the energy we
expend; energy-saving mechanisms are inte-
gral to modern civic design. Modern living has
become increasingly urbanised; in 1950 there
were 751 million people worldwide living in
urban areas, compared to 4.2 billion in 2018,
which is 55% worldwide but as high as 72% in
Europe and 82% in North America.[21]
Urbanisation is relevant to the obesity epi-
demic; research suggests that a lack of green
space and recreational facilities, perceived
unsafe communities, and high-density popula-
tions can be risk factors for obesity.[22]
Urbanisation is also relevant from an evolution-
ary perspective; community and kin networks
have been replaced in urban environments by
small living units or nuclear families, typically
with non-kin neighbours.[23]
126 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
In our ancestral, selective environment
there would have been food scarcity and
changeable conditions. In such circum-
stances it would have been adaptive to max-
imise caloric intake, especially of food rich
in sugar and fat. Our ancestors and tradi-
tional populations have evolved to maxim-
ise their foraging success and net energy
intake,[24–26] and, therefore, their health
and evolutionary fitness. Adaptations that
motivate preferences for high-calorie foods
would have been a consequence and humans
are reportedly hardwired to prefer HFSS
foods.[27] A mechanism of this adaptation
is that energy-rich foods elicit enjoyment
in humans, especially when consumed in
combination. Various hormones are released
when high-energy foods, particularly fats
and sugars, are consumed, which stimulate
pleasure-associated areas of the brain;[28]
­ status strongly predicts obesity;[31] in turn,
similar effects are found with alcohol, ciga-
rettes, and other substances. Humans are also
tolerant to food delays, which is likely to be a
product of changeable environmental condi-
tions. It seems likely that our ancestors were
adapted to maximise energy intake during
periods of food scarcity and moderate con-
sumption during times of food abundance.
The agricultural and industrial revolutions
brought great civil advancement, but food
technology and food systems have changed
more over the last 50 years than ever before;
production and supply of energy-dense and
processed foods have increased substan-
tially. Globalisation has resulted in a reduc-
tion in price and an increase in availability
of high-calorie foods.[20] The modern world
for most people affords a constant state of
food plenty and, given human preference
for HFSS foods, consumption of food and
prevalence of obesity have increased dra-
matically. It is for these reasons that obesity
has been described as a normal response to
an abnormal environment.[29] The advertis-
ing and food industries of the modern world
also work to maximise our contact with food
or associated psychological cues. So rather
than asking why some of us are living with
overweight, the more pertinent question may
be, why are some of us not living with over-
weight? We are conscious that overconsump-
tion is not conducive to good health, and we
moderate our behaviour accordingly, but our
ability to moderate consumption and other
diet-related behaviour varies. If conscious
moderation of behaviour is the mechanism
by which we prevent overeating, why are so
many people making suboptimal and seem-
ingly maladaptive health choices? Perhaps
these health choices are not maladaptive at
all.
There is a long-established link between
socioeconomic gradients and health risk
behaviours,[30] health outcomes, and mor-
tality.[3] While deprivation predicted food
poverty and underweight in the past, in the
modern developed world, low socioeconomic
living with obesity has been found to exac-
erbate inequalities.[32] Within the developed
world, evidence suggests that relative inequal-
ities are equally if not more important than
absolute poverty in predicting problematic
health behaviours and outcomes.[33] There
are environmental factors that accompany
deprivation, which increase the likelihood of
obesity. Energy-dense foods represent bet-
ter calories per unit cost compared to fresh
ingredients, which appeals to low-income
families.[34] The built environment in areas
of deprivation also has reduced green spaces
and fewer recreation facilities, such as leisure
centres, which limit opportunities for physi-
cal activity, increase risk factors for obesity,
and widen health inequalities overall.[35]
There are also higher densities of fast-food
outlets and reduced access to healthier food
stores in increasingly deprived areas, both of
which are associated with poorer diets.[36]
The relationship between disadvantage and
obesity is complex and dynamic but, in devel-
oped countries, people of lower socioeco-
nomic status are disproportionately affected
by obesity.[37] However, obesity is prevalent
across all socioeconomic strata and there is
variation in overweight and obesity within
 EVOLUTIONARY PSYCHOLOGY AND PUBLIC HEALTH
similarly deprived areas, which implies that
disadvantage only accounts for a proportion
of the observed variance.
Psychological Mechanisms and
Evolutionary Strategies
Psychological mechanisms or modules
throughout this chapter refer to functional
neurocognitive adaptations that have been
successful over evolutionary time owing to
their selective value in solving problems and
their contribution to survival or reproductive
success.[38] In combination and in addition
to anatomical and physiological traits, psy-
chological mechanisms create broader evolu-
tionary or behavioural strategies, a clear
example being an individual’s reproductive
strategy.[39] When originally conceptual-
ised, psychological mechanisms or modules
were proposed to be universal among humans
but that different mechanisms were invoked
in different situations, leading to phenotypic
variation.[40] It is proposed here that while
some fundamental psychological mecha-
nisms are innate and species-typical, others
are more varied across individuals, arising
from individual-specific interactions between
genetic predispositions and phenotypic plas-
ticity.[41]
Neurobiological structures are produced
by our genes after thousands of years of selec-
tion pressures, which act on individual genes
but also the resulting cognitive mechanisms.
Physically, the brain is complex, comprising
billions of neurons and chemical interac-
tions, but it functions as a specialised system
that has been produced by the evolutionary
process.[42] The brain acts hierarchically,
where functionality emerges as a prop-
erty of micro- and mesoscopic interactions
through coordinated chained structures that
run between anatomically clustered areas.
[43] For example, the choice to eat a donut
would flow through and elicit responses
from various areas of the brain: affect and
emotion centres, autonomic and conscious
127
centres, expectation centres, short- and long-
term memory,[44] all before a hand has been
outstretched. Selection has not acted upon
these areas in isolation; the idea of modu-
larity recognises functional specialisations
within the brain that are domain-specific and
are utilised for different cognitive processes.
Whether modularity is strong or weak, any
choice and resulting behaviour is the product
of a hierarchical chained process that breaks
down with impairment of any of the involved
areas.[45] Psychological mechanisms are
selected in their own right; they may be her-
itable in a conventional sense, but may also
be the product of behavioural and, therefore,
phenotypic plasticity, leading to selectable
individual variation.[41]
Genetic predispositions form the basis of
all psychological mechanisms, but they also
remain a part of our complex biological sys-
tems and are mutually associated with our
physiology. For example, and in relation to
obesity, adipose tissue is an active organ and
part of the homeostatic system that regulates
satiation and energy balance. Adipose tissue,
the gastrointestinal tract, and the pancreas
release various hormonal regulators to the
brain, which receives the signals and acts to
reprioritise behaviour to stimulate or sup-
press appetite. There are also physiological
detectors for fats and sugars in food, which
stimulate pleasure-associated areas of the
brain. Elicitation of reward mechanisms also
motivates behaviour, to such an extent that
homeostatic processes may be overridden.
This push and pull creates a trade-off within
our energy-balance system, whereby psycho-
logical mechanisms, each with a clear adap-
tive significance, compete at cross-purposes.
It is a feature of environments with constantly
available energy-rich foods that allow the
reward mechanisms to gain the upper hand
and enable a chronic positive energy balance.
Developmental conditions also affect psy-
chological mechanisms, beginning in utero;
these may be physiological, physical, emo-
tional, or sociocultural. Childhood experience
has been found to profoundly affect cognitive
128 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
development, psychological mechanisms,
and therefore behaviour. Neurobiological
development occurs rapidly throughout child-
hood but continues through adolescence and
into adulthood. Cognitive systems and path-
ways that govern behaviour can be affected
if areas of the brain suffer impairment dur-
ing development. Adverse experiences in
childhood, including psychological or physi-
cal abuse, change the formation of physical
structures and neural pathways, which can
profoundly affect healthy development and
behaviour in later life. These changes can
also be irreversible, meaning that the conse-
quences of adverse experiences often persist
into adulthood and act independently of socio-
economic pressures.[46] Adverse childhood
experiences have been found to contribute to
problematic eating behaviours, weight gain,
and overweight and obesity in childhood and
adulthood.[47–49] The responses to adverse
experiences may have adaptive roots; stress-
ors are linked to heightened immune- and
nervous-system responses, but continued
stress may produce over-activation that is
likely to create a platform for poor mental
health.[50] Adverse experiences are likely
to be accompanied by unstable and insecure
food environments, an adaptive response to
which may be overconsumption when the
opportunity arises. If the consequences of an
adverse childhood persist into adulthood, so
too would the food response, even when food
becomes more consistently available.
Psychological mechanisms are also
profoundly shaped by environmental cir-
cumstances, both physical and, in modern
contexts, socioeconomic.[51] Our phenotypes
are cumulative developmental outcomes of
interactions between our genes and envi-
ronments. Assuming that the mechanism by
which a trait is inherited does not constrain
its adaptive value, it is likely that behaviour
and the underlying mechanisms are modi-
fied in line with environmental variation.[52]
While fields such as human behavioural ecol-
ogy face challenges in testing quantifiable
hypothesis in non-traditional or industrialised
countries, it seems logical that behavioural
strategies are matched to environmental con-
texts or circumstances, despite the debatable
and nuanced influence of culture. It will be
argued here that health behaviour is strate-
gic and adaptive, which would provide the
principal reason why seemingly maladap-
tive choices, such as overeating, are so often
taken. There is a wealth of evidence that the
health behaviour of industrialised popula-
tions is influenced by physical and socioeco-
nomic environments; deprived populations
are more likely to have low-nutrition diets,
and more likely to engage in a range of other
health risk behaviours.[53–54] The trend
is not solely economic; even when health
behaviours are free, people from increasingly
deprived groups have been found to be less
likely to engage with them,[30] implying the
issue is strategic. The way we perceive our
environmental circumstance is also impor-
tant; perception of income, for example, has
been shown to be more important than actual
income in determining related behaviour.[55]
Living in environments that are to lesser
or greater degrees mismatched from those in
which we evolved has subtle and complex
implications for modern humans. Rates of
mental health problems, including depression,
anxiety, and stress, have never been higher,
especially among young people. Our behav-
iours act dynamically with environmental
pressures; governed by psychological mecha-
nisms and broader strategies, our behaviour
is a response to the environment, whether
physiological or psychological, which is then
experienced and remembered. Our experi-
ence leads to learning, which subtly adjusts
the psychological platform on which behav-
iour is based. Just as adverse developmental
experiences can impair cognitive functioning,
so can more transient states of low mood or
poor well-being, and repetition of such states
can lead to poor mental health. Similarly, other
temporary states such as illness and disease
can alter not only our behaviour but our per-
ception of social and physical environments.
Low mood and emotional disorders have been
 EVOLUTIONARY PSYCHOLOGY AND PUBLIC HEALTH
found to affect behavioural strategies, and spe-
cifically those related to food consumption
and physical activity.[56]
The key determinants of psychological
mechanisms and, therefore, behavioural strate-
gies are genetic, physiological, environmental,
developmental, and subjective (Figure 6.2).
These determinants are not mutually exclu-
sive but are linked as part of a dynamic sys-
tem. For example, phenotypic expression is
primarily the product of interplay between
genetic predispositions and environmental
conditions; physical and socioeconomic envi-
ronments are inextricably linked but are also
associated with developmental and subjective
determinants; and developmental and sub-
jective factors also feedback and modify the
physiological system. Resulting mechanisms
may be fixed, others may be more plastic,
varying with changes in circumstances or even
mood states. Behavioural plasticity allows us
to make different choices and exhibit ­different
Figure 6.2  Key determinants of psychological mechanisms and behaviour strategies
129
behaviours within a changing environment;
behavioural plasticity was and remains a huge
selective advantage for humans.[57] Our abil-
ity to modify our behaviour extends selection
beyond psychological mechanisms and the
resulting behaviours to behavioural strate-
gies themselves. Such strategies are likely to
be adaptive but modifiable both temporally
and with changing environments, meaning
that physical, political, social, or individual
change would be likely to shift the parameters
for optimal behaviour. Examples of this can
be anecdotally observed every day: a natural
disaster, political change, or a dramatic change
in personal circumstance are all likely to dra-
matically shift the behaviour of an individual
or collective. Health behaviour strategies are
determined more subtly, especially given the
sometimes contradictory pressures of health
and fitness, but perhaps unhealthy choices are
not maladaptive but optimal given the evolu-
tionary context of the choice.
130 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Adaptive Mechanisms and
Optimal Strategies
As outlined by life history theory, behavioural
optimality is a shaped by a range of problems,
including survival, growth, and development.
[39] These represent multiple and complex
pressures, which underpin choices or deci-
sions that produce variable behaviours. All
behaviour, including health behaviour, is the
product of a cognitive appraisal system, the
process of choosing an action from a range of
options with the expectation of maximising
the benefit and minimising the cost.[58] The
stress-response system functions to mediate
openness to environmental inputs and regu-
late behaviour in a range of fitness-relevant
areas. ‘Switch’ mechanisms regulate genetic
and environmental influences on phenotypic
development; during development, informa-
tion processed by these mechanisms feeds
back and recalibrates the stress-response
system, resulting in individual and adaptive
patterns of behaviour.[59] Environmental
moderators may be social or familial, physi-
cal, including the built or natural environ-
ment, or socioeconomic, given that there are
established gradients in health behaviour
across the spectrum of inequality.
Temporal decision making provides an
example, and in particular the extent to
which behavioural strategies are under-
pinned by impulsive or reflective decision
making. Rather than assuming all impulsive
behaviour is the absence of control, there is
utility in considering a dual system of behav-
iour,[60] where reflective and impulsive
influences exist, are differentially adaptive,
and are often in conflict or trade-off against
each other.[61–62] A moderator of this sys-
tem may be developmental factors, that have
the potential to inhibit the healthy forma-
tion of neurobiological structures, which are
important in overriding impulsive mecha-
nisms for behaviour. A good example is the
impact of developmental conditions on the
functioning of the stress-response system,
a biological mechanism involved in a wide
range of adaptive functions.[59] In a health
setting, impulsiveness is sometimes con-
sidered hedonistic but reflective or rational
behaviour involves constraint and is likely to
be the product of conscious or higher cogni-
tive control. Hedonic impulsivity may bring
a sense of enjoyment and freedom to the
human experience,[63] but in a health sense,
this often involves long-term costs. Smoking
cigarettes, using substances, drinking alco-
hol, or eating HFSS foods may be enjoyable
in the short term but are likely to pose future
health risks.
Reflective and goal-directed behaviour is
often viewed as rational, reasoned, and con-
scious, utilising volitional control or self-
regulation to achieve a goal, while impulsive
decisions, despite incorporating a range of
behaviours, have been described as subop-
timal, an inability to moderate behaviour.
Reflective decisions usually have the goal
of a larger payoff at a later time point, while
forgoing the immediate reward or benefit. In
terms of health behaviour, this would be rep-
resented by forgoing the immediate pleasure
and reward from an action, such as eating an
HFSS food, in order that the chances of yield-
ing the longer-term and larger benefit of good
health are enhanced. Impulsivity can be inter-
preted as the forgoing of a large but delayed
reward in favour of a smaller but immediate
reward.[58] In certain environmental condi-
tions, impulsivity is likely to be adaptive;
dangerous environments, for example, where
there is an immediate threat or mortality rates
are high, are likely to favour short-term and
high-risk behaviours.[39] Research sup-
ports this theory, demonstrating that people
in adverse environments are more likely to
discount the future and act more impulsively,
despite the potential harm to their health.[64]
Impulsive and reflective decision making has
also been shown to predict health behaviour in
the way we would expect; lower health-risk-
taking behaviour is associated with increas-
ingly reflective decision making, while riskier
health behaviour is associated with increas-
ingly impulsive decision making.[65–66]
 EVOLUTIONARY PSYCHOLOGY AND PUBLIC HEALTH
Within the field of cognitive psychology,
the concept of delay or temporal discount-
ing[67] is used to explore our cognitive
appraisal system, specifically relating to the
temporal nature of health economics. Delay
discounting is the extent to which immediate
rewards are declined or postponed in order to
gain larger rewards in the future; these con-
structs create behavioural strategies, which
are variable, context specific, and are highly
likely to be adaptive. Crucial in the deci-
sion are the values of the present and future
reward, but also the length of time required
or delay in gaining the future benefit; this
can be considered as a cost subtracted from
the future reward. It is well established that
humans trade off between present and future
rewards and, as previously stated in terms of
food choice, tolerance for delays in forag-
ing may have been selected for in variable
environments. It is also well established that
humans make temporally optimal decisions
with financial or resource-based benefits and
costs; the principle of delay discounting has
been used to predict various health behav-
iours, including food consumption, physical
activity, tobacco, and alcohol use.[68] The
constructs that underlie impulsive or reflec-
tive behaviour and delay discounting are
psychological mechanisms, which have been
selected upon but remain modifiable with
changing circumstances of environments.
The relevance of these mechanisms and their
determinants to the field of public health
have not been fully explored but may hold
the key to understanding why suboptimal
health behaviours are chosen to the point of
pathology and premature death at epidemic
proportions.
Behavioural traits, whether fixed or variable,
contribute to broader behavioural strategies and
our overall phenotype. Behavioural strategies
can become stable in evolutionary terms,[69]
but changes to our environment create pres-
sures, which alter learning and lead to new
behaviours and strategies. This implies that
for a set of environment conditions, including
socioeconomic, there is likely to be an optimal
131
and adaptive behavioural strategy.[70] Within
these parameters are individual factors, which
may have arisen from biological, develop-
mental, or subjective determinants and may
be fixed or transient. Individual states may
not alter the objective optimality of a behav-
ioural strategy matched to a set of environ-
mental conditions, but they would alter the
cognitive appraisal system and the perceived
costs and benefits in temporal decision mak-
ing. Like adverse developmental conditions,
strategies that favour short-term payoffs and
immediate rewards may have adaptive roots.
In states of high anxiety or depression, it may
not be optimal or even possible to think about
long-term benefits; it is likely that for some-
one suffering from a mental health disorder,
temporal perception may be very different to
someone of good mental health and the cost
of waiting for a reward may be perceived
to be much higher. Such individuals may
still be behaving optimally and adaptively,
upon consideration of the determinants of
their psychological mechanisms, the inter-
nal influences on behavioural appraisal,
and the social, physical, and socioeconomic
environment.
These ideas can be further evidenced by
considering behavioural strategies that have
established evolutionary significance and
their relationship to currencies of health.
Risk taking or aversion is adaptive in vari-
ous ancestral and modern contexts and has
relevance to health. While there are broad
differences in risk-taking behaviour between
age and gender groups, strategic variation
can also be observed. Life history theory pre-
dicts that decisions relating to energy allo-
cation and behaviour are divided between
competing functions, such as growth and
reproduction, which are affected by ecologi-
cal pressures, the differential proportion of
which result in diverse health outcomes.[71]
Consider the relationship between wealth and
reproduction – increasing wealth predicts
higher long-term fitness and reproductive
success but a trade-off also exists between
the number of offspring produced and the
132 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
wealth transmitted to each; this balance has
been found to be predicted by socioeconomic
and environmental conditions.[39] Accruing
wealth in the developed world requires time
and career focus, which is time spent away
from having and raising children; women
in particular face a decision of how much
wealth to accrue before having children.
Accruing wealth is desirable but there is risk-
taking relevance given that fertility decreases
with increasing age. Risk-taking decisions
can also be observed more directly in a health
setting. The choice to forgo protection dur-
ing sex, such as a condom, is risky given the
increased chance of getting an infection or an
unwanted pregnancy. However, unprotected
sex is more pleasurable for men and women,
and represents a reward to counter the risk;
research has shown that the greater the per-
ceived pleasure differential, the more will-
ing men and women would be to engage in
unprotected sex.[72] Similarly, eating HFSS
foods, drinking alcohol, or using other sub-
stances can also be pleasurable in the short
term but each behaviour incurs long-term
risks to morbidity and mortality. It is also well
established that health risk behaviours cluster
together,[73] implying there is an underlying
risk-taking strategy for any individual within
a specific set of circumstances.
Varying strategies of health risk, as shaped
by external factors, can also be illustrated in
financial terms. Low-income workers have
been found to be less likely to participate in
insurance schemes, even when large subsi-
dies are offered.[74] Despite the losses of not
having insurance being potentially ruinous,
low-income workers are less likely to partici-
pate because they have less to spend and, cru-
cially, less to lose. Conversely, gambling and
playing the lottery is more popular among
poorer groups;[75] although the chance of
winning is slim, the relative reward is higher
for someone who has less. One could predict
that people in adverse circumstances, whether
environmental, developmental, or subjective,
would devalue future payoffs as the cost of
waiting is perceived to be higher given the
immediate adversity or danger. In this case, it
makes sense to opt for shorter-term rewards
and payoffs. Chances of survival have been
found to profoundly shape reproductive strat-
egy,[39] and are likely to affect strategies
relating to health or wealth in a similar way.
An individual’s life history reflects trade-offs
in the allocation of time or energy, and may
be categorised as slow or fast depending on
the environmental conditions and perceived
time horizons.[76] Preferable environments
are likely to induce slower or longer life
histories, which are associated with higher
investment, lower impulsivity, and lower lev-
els of risk taking. Conversely, shorter life his-
tories are associated with future discounting,
impulsivity, and risk taking.[76]
APPLICATIONS OF EVOLUTIONARY
CONCEPTS TO PUBLIC HEALTH
PHENOMENA
Investing and offsetting decision making is
fundamental to impulsive and reflective deci-
sion making and may be applied to various
health behaviours. Exercise can be consid-
ered as investing behaviour; while there may
be short-term rewards for some people, costs
are involved in the form of time and energy.
The investment is made with an idea that
there will be a reward of good health and
potentially a long-term benefit of longer life.
Buying healthy foods can also be considered
as an investment; fresh ingredients tend to
cost more and usually take more time to pre-
pare but they may represent a similar long-
term goal to that of exercise. In many
societies, health insurance represents an
investment; it may be part of a healthcare
system or it may represent a better level of
care. Like risk-aversion behaviours, these
decisions are more likely to be taken by indi-
viduals who have the financial means but
also who perceive the long-term benefits to
be worth the investment. For those who do
not have the means or think in temporally
 EVOLUTIONARY PSYCHOLOGY AND PUBLIC HEALTH
shorter terms due to adversity in one or more
areas of their lives, investments are less
likely to be made. Offsetting behaviour can
be thought of in similar but subtly different
terms; instead of immediate investment costs
being incurred, immediate rewards are
rejected but still with the expectation of
higher rewards in the future. Choosing not to
eat HFSS foods, drink alcohol, or use sub-
stances are examples of offsetting behaviour
as they are immediately pleasurable but in
accumulation are likely to incur risks to long-
term health.
Altruistic and cooperative behaviours also
have relevance to health. In ancestral envi-
ronments, altruism may have evolved partly
in response to foraging and food sharing;
evidence has shown higher levels of altruistic
behaviour to be associated with better health
and well-being. Cooperation has relevance
given that sharing resources equally allows
for moderation and fairness but requires trust
that community members will not take more
than their fair share. The global food industry
is a good example of non-cooperative behav-
iour whereby nation states or companies com-
pete to gain as much market share as possible.
Such competition has led to over-farming
and fishing, which have led to environmental
destruction. Huge production and advertising
of processed HFSS foods within a global mar-
ket have also pushed consumption to the point
that the majority of the world’s populations
live in countries where they are more likely
to suffer mortality from the consequences
of living with overweight than living with
underweight.[77] These pressures operate in a
similar way to the other health-relevant evolu-
tionary behaviours, whether for individuals or
larger organisations. Longer-term rewards that
may be gained with altruistic and cooperative
strategies are not optimal given the context,
in this case the free-market food system. The
optimal strategy for an individual, company,
or nation state in such circumstances is to take
the immediate payoff or reward.
Extreme examples of adverse devel-
opmental, subjective, and environmental
133
circumstances can help to illustrate how
extreme behaviours might make sense in
terms of the adaptive appraisal systems that
underpin them. Victims of childhood physi-
cal or sexual abuse commonly grow up to
suffer poor health outcomes; not only is
cognitive development impaired, potentially
leading to reduced reflective decision mak-
ing, but the immediate reward that may be
provided by an unhealthy behaviour would
almost always outweigh the cost of waiting
for a larger future reward. Individuals who
suffer from mental health conditions could
make a similar appraisal; for a person living
with severe depression, for example, poten-
tial benefits in the distant future may feel
completely irrelevant. In these examples,
the immediate reward could be alleviation
or self-­medication from the ongoing costs of
extreme low mood. These unhealthy behav-
iours, which incur long-term risks to health
and fitness, are uniquely adaptable in that
they may be survival mechanisms. Consider
populations living in extreme environmen-
tal conditions such as war or within violent
gang culture. Individuals are aware that
their chances of survival are reduced, which
diminishes the value of any future reward as
they may not live to benefit from it. Short-
term strategies would always be optimal in
situations such as these since the rewards of
tomorrow may never come.
For any individual, health behaviours
can be better understood if framed as adap-
tive strategies that can be predicted by envi-
ronmental, developmental, and subjective
circumstances. Early exploratory work has
demonstrated that environmental conditions
shape the strategies that drive health behav-
iour and that such strategies are predictive of
various adaptive or evolutionary strategies.
[70] Public health practitioners understand the
determinants of poor health but there is little
acknowledgement that there may be circum-
stances in which it makes more sense to under-
take unhealthy rather than healthy behaviours.
Under such circumstances, it seems point-
less to try to change the behaviour itself, if
134 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
the underlying strategy is not acknowledged
and the determinants of that strategy are not
addressed. Such an approach may also widen
the public health perspective. Often academ-
ics and practitioners operate in silos, focusing
on specific health risk behaviours, interven-
tions, or risk factors for disease, but perhaps
the problem needs to be re-framed. In addi-
tion to addressing the morbidity arising from
obesity, how can we triangulate research and
practice to reduce the risk factors for NCDs
more broadly? A key element of this chal-
lenge may be seeking to address the deter-
minants of health risk behaviours. The public
health response must be informed, paradigms
may need to be challenged, and we will need
a whole system and population approach
to address a whole system and population
problem. Given the complexity of the prob-
lem, no simple solution is likely to be suc-
cessful and each of the key determinants of
unhealthy behavioural strategies need careful
consideration.
In public health, it is understood that pre-
vention is better than cure and this is espe-
cially true with the obesity problem. We
understand how important and formative
developmental experiences are, and that the
impact of adversity may be irreversible in
terms of cognitive structures and resulting
behaviours. In a similar way, the public health
response to the obesity problem must begin
from pregnancy, and carry on through early
years, childhood, and adolescence, given
that nurturing environments with healthy
parent–child bonds are crucial to long-term
health. Everything is important in early life
from breastfeeding and weaning to a healthy
school environment and safe, enabling com-
munities. Children who live with obesity are
likely to become adults who live with obesity,
and adults very rarely sustain weight loss. In
addition to meeting health recommendations
for early life, it is our developmental expe-
rience that can profoundly affect our short-
and long-term behaviours. Creating positive
childhood experiences is a complex societal
challenge but it must be recognised as critical
in addressing the obesity epidemic and other
major risk factors for NCDs.
Developmental experiences often con-
tribute to subjective issues that continue to
underpin unhealthy behavioural strategies in
adults but, equally, poor adult well-being or
mental health problems can be triggered by
life circumstances at any time. Recognition
and appropriate treatment are hugely impor-
tant not only to treat the disorder but also to
re-balance the appraisal system that favours
short-term rewards but has consequences for
long-term health. Mental and physical health
are often treated separately but are inextrica-
bly linked and are often cyclical. We cannot
expect an individual to make choices that are
good for their long-term health when they are
struggling to cope with the here and now.
Further to developmental and subjective
determinants, the environments in which we
live must be adapted to address key public
health problems and particularly obesity. There
is a clear need to create local physical and
social environments that are conducive to good
health; there are multiple and varied features
of our home, social, and work environments
that affect our health and well-being. However,
there is a bigger challenge to enabling good
health, and it lies in the industrial, political,
and philosophical climate in which we live. We
are the objects of a free-market food system in
which the primary objective of any company
is to make money and grow, not always but
often at the expense of ill health and a burden
of disease. The food system is incredibly com-
plex but as long as companies large and small
are permitted to produce unhealthy products,
saturate the market, and advertise ubiquitously,
we cannot hope to shift the behaviour of popu-
lations in a meaningful way. There has been
a helpful paradigm shift among academics
and public health experts, who now consider
it unreasonable for populations to achieve
and maintain a healthy weight through indi-
vidual agency alone. Powerful regulations for
industry must be demanded by the public and
actioned by policy makers, if we are to tackle
the obesity epidemic in a meaningful way.
 EVOLUTIONARY PSYCHOLOGY AND PUBLIC HEALTH
Perhaps more importantly, the greatest
challenge of our time is to reduce socioeco-
nomic inequalities. At a population level,
individuals from relatively poorer groups
make unhealthier choices, these choices
are likely to be underpinned by behavioural
strategies, and it is likely that these strategies
are adaptive. Trying to change the outcomes
of the behaviour is unlikely to be effective
without consideration of the environmen-
tal conditions that determined the strategy.
Reducing inequalities is a global problem and
requires unprecedented change but in terms
of the public health response, interventions
and policies must be careful that they reduce
rather than exacerbate inequalities. For
example, when dealing with obesity, nation-
wide policies that address issues such as pric-
ing are likely to reduce inequalities, while
person-focused interventions that encourage
behaviour change are likely to widen them.
[78] Health disparities are the most serious
outcome of socioeconomic inequalities, the
reduction of which will certainly require an
understanding of the implications and appli-
cations of evolutionary principles.[79]
These broad discussion points do not
overlook the huge amount of academic
and public health work that contributes
substantially and meaningfully to addressing
obesity and other health problems. The
object here is to suggest that an evolutionary
perspective of health risk behaviours may
be helpful, given that they are likely to be
optimal and adaptive. Recognition of the
strategies that drive unhealthy choices, and
the key determinants of them, will not only
encourage a focus on the root causes of
health problems,[2] but might also reduce
the stigmatisation of health risk behaviours
and the individuals that carry their burden.
An engrained mantra of public health is to
‘make the healthy choice the easy choice’
but in evolutionary terms the easy choice
is the adaptive choice; perhaps we have
known for some time that health risk
behaviours are a normal response to an
abnormal world.
135
EVOLUTIONARY PSYCHOLOGY AND
PUBLIC HEALTH: CASE STUDY –
VACCINATIONS
For over 200 years,[80 ] vaccinations have pre-
vented the consequences of disease for millions
of people worldwide. Vaccines are a public
health success story and are estimated to have
saved at least 10 million lives between 2010
and 2015 alone.[81] Vaccinations stimulate an
immune response from the body to develop
resistance to a pathogen, which can lead to the
prevention or complete eradication of an infec-
tious disease.[82] Vaccines have successfully
led to the worldwide eradication of smallpox
and widespread eradication of measles, polio,
and tetanus. Other standard vaccine-­preventable
diseases include diphtheria, whooping cough,
mumps, rubella, influenza, and hepatitis, but
there are many other infectious diseases that
have available vaccines.
Vaccines are a basic human right and have
greatly reduced the burden of disease[83] but
despite this success, a determined antivaccine
lobby, representing an increasingly sceptical
world view of science, is on the rise.[84] The
antivaccine movement has gained traction
online and is suggested to be well organised
and widely dispersed.[85] Antivaccine mes-
sages question vaccine safety and effective-
ness and have been reported to rely on strong
emotional messages to enhance their appeal.
[86] For this reason, a minority of antivac-
cine advocates have been suggested to have
a highly persuasive voice and the potential
to persuade millions of parents against vac-
cinations on scientific, ethical, and political
grounds. Medical and academic experts have
tried to counter such messages to advocate
the safety and importance of vaccines but
have often suffered ‘harassment campaigns’,
typically via social media by vehement anti-
vaccine activists.[87] While the most effec-
tive strategy at combatting this problem is
unclear,[88] the antivaccine movement rep-
resents a difficult and complex challenge for
public health policy makers and practitioners.
136 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
While the academic consensus is that the
antivaccine movement is based on miscon-
ceptions,[89] there have been controversial
studies reporting links between vaccinations
and disability and disease, such as autism and
asthma, despite strong evidence to the con-
trary.[90–92 ] Emotive online pressure and
spurious evidence have succeeded to some
degree. In the UK, the link with the measles,
mumps, and rubella (MMR) vaccine and
autism appears to have gained momentum
and, following year-on-year increases from
2007, decreases in coverage were recorded
in 2016/17 for the third year in a row.[93] In
the United States, while the overall rate of
vaccination coverage has remained high, the
rate of children not receiving any vaccina-
tions has been increasing since 2011.[94] The
consequences of the growing antivaccina-
tion movement have already been observed;
analysis of a measles outbreak in the United
States in 2015 indicated that the likely cause
was lack of or incomplete vaccinations.[95]
Multiple developed and developing countries
have experienced outbreaks of measles in
recent years,[96–97] which have been pri-
marily attributed to gaps in vaccination cov-
erage.[97–98]
Research suggests that framing public
health messages within the context of the
moral foundations that underpin judgements
can successfully influence attitudes.[99]
Moral foundations theory proposes that inter-
nal decision making predicts social group
attitudes.[100] While there is complexity
surrounding decision-making processes, con-
sideration of selective pressures and evolu-
tionary principles that underpin motivations
may provide an enhanced understanding of
public health decision making.
The incentive to vaccinate is clear; it leads
to individual and group immunity and is the
best defence against potentially epidemic
diseases. If sufficiently high proportions of
people vaccinate, herd immunity[101] can
be achieved whereby a whole population
becomes resistant to a parasite or disease.
The coverage threshold varies by disease
but is less than 100%; estimates suggest
that for a disease such as measles the herd
immunity threshold is 93–95%; for diseases
such as smallpox or polio the threshold is
80–86%.[102] Aside from the questionable
links between vaccines and disability or dis-
ease, there are various risks involved with
vaccinating, typically low-level side effects.
Vaccines can be painful and create soreness
and swelling, and in some cases they can also
induce allergic reactions and subjects may
become a little unwell. Individual and popu-
lation immunity is a clear benefit of vaccina-
tions, while the risks and side effects can be
interpreted as costs.
An application of the principles of evolu-
tionary psychology can add insight into the
decision-making process of whether or not
to vaccinate. Altruism is an act that benefits
others (non-kin) at expense to the actor,[103]
and cooperation, like altruism, provides a
benefit to another individual or group but
does not require reciprocation since it also
yields direct or indirect fitness benefits to the
actor.[104] The process of vaccinating does
not neatly fit with either principal definition;
it is altruistic in that a cost is incurred and the
process benefits others, but it is also coop-
erative in that it yields direct fitness benefits
to the actor. Vaccinating involves elements of
altruism and cooperation; the commonality
between the two is reliance on others to max-
imise individual and group benefits.
However, within populations counter
strategies may occur arising from selfish
or cheating behaviours that yield benefits
to the individual with no personal cost but
with an expense to others. Cheating strate-
gies may often lead to higher relative fitness
than altruistic strategies[105], but if caught,
cheats are likely to be punished. Emotional
mechanisms have evolved, including mor-
alistic aggression, trust, suspicion, and dis-
honesty, a function of which is to regulate
decisions and behaviours relating to altruism
and cheating.[106] Cheating, free riding, or
bandwagonning have been found to moti-
vate vaccination decisions[107], and across
 EVOLUTIONARY PSYCHOLOGY AND PUBLIC HEALTH
a large population, cheating is incentivised
given that benefits from herd immunity can
be realised without paying the individual
costs, assuming that the proportion of free
riders does not exceed the threshold. One
would expect a successful cheating strategy
to not only be hidden but to outwardly sup-
port vaccinations, in order that others vacci-
nate and herd immunity is maintained.
Yet there appears to be a maladaptive cul-
tural phenomenon whereby, far from being
hidden, not vaccinating or cheating in an
evolutionary sense is publicly advocated and
promoted. It may be that cultural misinfor-
mation has shifted the perceived benefits
and costs to such an extent that the adaptive
strategies of vaccinating or free riding have
been abandoned in favour of a wholly mala-
daptive strategy. This is likely to be an exam-
ple of humans being adaptively mismatched
from a technologically developed world;
our psychological mechanisms that strate-
gize our behaviour may be fundamentally
misinformed leading to suboptimal decision
making. Vaccinations are a relatively recent
phenomenon and were not a feature of our
formative and selective environments, but
our ability to make adaptive choices in novel
situations is a key feature of our behavioural
plasticity. It seems likely that a successful
public health response will need to change
the parameters of the game in order that
modern humans can once again make sense
of their world.
Perhaps a consequence of living in urban
populations rather than smaller communi-
ties is that people have become less trusting;
the choice not to vaccinate has been found
to be influenced by confidence, not only in
the product but also the health professional
and the policy maker.[108–109] Evidence
suggests public confidence in vaccinating is
decreasing,[110] and that social norms and
interactions with healthcare providers are
highly influential on decisions to vaccinate.
[111] A response by public health policy mak-
ers and practitioners may be to foster trust by
increasing the availability and transparency
137
of information relating to the product and
the process. Research suggests that 37% of
US adults when questioned were not aware
of the concept of herd immunity and over
75% thought that vaccination coverage was
higher than it was.[112] While there is some
evidence that educational interventions can
improve willingness to vaccinate,[112] most
traditional education tools have been found to
have little impact on vaccine hesitancy,[113]
including those that specifically dispel the
myths linking vaccines with disability.[114]
Since antivaccine lobby groups assume a
disproportionate space within the discus-
sion arena,[115] the key response may be to
engage with mass media and initiate wide-
spread vaccination-promotion campaigns;
there is some evidence that such campaigns
can reduce vaccine hesitancy and increase
coverage rates.[116] Perhaps fighting the fire
of powerful antivaccine lobbyists requires the
clear and effective communication of scien-
tific evidence via the medium of social and
digital mass media.
One further much discussed public health
response could be to make vaccinations com-
pulsory; proposals are currently in place in
various countries in the developing world in
light of recent outbreaks.[117] Aside from
the potential implications for civil liberties
and the difficulty in implementation, it has
also been suggested that mandating vaccines
would not address the causal problems of
vaccine hesitancy[118] and may increase the
inherent mistrust of the political health sys-
tem.[115]
Accounting for the available evidence,
there have been sensible and well-considered
recommendations of best practice to improve
vaccination coverage in an age of inherent
mistrust and hesitancy. Calls to utilise immu-
nisation frameworks, apply multi-component
interventions, link interventions to empirical
evidence, and take account of the person-
and environment-specific factors[36] are
valid and helpful. However, mindfulness of
the maladaptive strategies that have propa-
gated in recent years and the evolutionary
138 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
decision-making mechanisms that appear
to be misfiring may be crucial in reminding
parents that, in this case, the adaptive choice
is the healthy choice. A successful campaign
may enable free riding to become a success-
ful strategy once again, as long as cheats are
low in number and quiet of voice, in order
that they alone incur the risks.
REFERENCES
1 Acheson, D. (1988). Public Health in England:
The Report of the Committee of Inquiry into
the Future Development of the Public Health
Function. Cm. 289. London: HMSO.
2 Gluckman, P.D., Low, F.M., Buklijas, T.,
Hanson, M.A. and Beedle, A.S. (2011). How
evolutionary principles improve the
understanding of human health and disease.
Evolutionary Applications. 4. 2. 249–263.
3 Stringhini, S., Carmeli, C., Jokela, M.,
Avendaño, M., Muennig, P., Guida, F. et  al.
(2017). Socioeconomic status and the 25 ×
25 risk factors as determinants of premature
mortality: a multicohort study and meta-
analysis of 1.7 million men and women. The
Lancet. 389. 10075. 1229–1237.
4 Nettle, D. and Pollet, T.V. (2008). Natural
selection on male wealth in humans. The
American Naturalist. 172. 5. 658–666.
doi:10.1086/591690.
5 Gillespie, D.O.S., Russell, A.F. and Lummaa, V.
(2008). When fecundity does not equal fitness:
evidence of an offspring quantity versus quality
trade-off in pre-industrial humans. Proceedings
of the Royal Society. 275. 713–722.
6 Giosan, C., Mureșan, V., Wyka, K., Mogoașe,
C., Cobeanu, O. and Szentagotai, A. (2018).
The Evolutionary Fitness Scale: A measure
of the independent criterion of fitness.
The Journal of the Evolutionary Studies
Consortium. 7. 1. 181–208.
7 Jones, J.H. (2009). The force of selection on
the human life cycle. Evolution and Human
Behavior. 30. 305–314.
8 Jasienska, G. (2013). The Fragile Wisdom: An
Evolutionary View on Women’s Biology and
Health. Cambridge, MA: Harvard University
Press.
9 Litman, G.W. and Cooper, M.D. (2007). Why
study the evolution of immunity? Nature
Immunology. 8. 547–548.
10 Schmid-Hempel, P. (2011). Evolutionary Para-
sitology: the Integrated Study of Infections,
Immunology, Ecology, and Genetics. Oxford
Scholarship Online. doi:10.1093/acprof:
oso/9780199229482.001.0001.
11 Greaves, M. (2010). Cancer stem cells: back
to Darwin? Seminars in Cancer Biology. 20.
2. 65–70.
12 World Health Organization. (2018).
Noncommunicable diseases – key facts.
[Date accessed: 4th March 2019] www.who.
int/news-room/fact-sheets/detail/
noncommunicable-diseases.
13 World Health Organization. (2019). Global
Health Observatory (GHO) data – risk factors.
[Date accessed: 10th March 2019] www.
who.int/gho/ncd/risk_factors/en/.
14 World Health Organization. (2018). Taking
action on childhood obesity. [Date accessed:
10th March 2019] https://www.who.int/end-
childhood-obesity/publications/taking-
action-childhood-obesity-report/en/.
15 Wang, Y.C., McPherson, K., Marsh, T.,
Gortmaker, S.L. and Brown, M. (2011).
Health and economic burden of the projected
obesity trends in the USA and the UK.
Lancet. 378. 815–825.
16 Butland, B. (2007). Tackling obesities: future
choices. Project report. London: Government
Office for Science.
17 Schrempft, S., van Jaarsveld, C.H.M., Fisher,
A., Herle, M., Smith, A.D., Fildes, A. et  al.
(2018). Variation in the heritability of child body
mass index by obesogenic home environment.
JAMA Pediatrics. 172. 12. 1153–1160.
doi:10.1001/jamapediatrics.2018.1508.
18 Institute of Medicine (US) Committee on
Assessing Interactions Among Social,
Behavioral, and Genetic Factors in Health;
Hernandez, L.M. and Blazer, D.G., editors.
(2006). Genes, Behavior, and the Social
Environment: Moving Beyond the Nature/
Nurture Debate. Washington (DC): National
Academies Press (US) 3. Genetics and Health.
19 Llewellyn, C. and Wardle, J. (2013). Genetic
Influences on Child Eating Behaviour. In Ency-
clopedia on Early Childhood Development,
edited by R.E. Tremblay, M. Boivin, R.D. Peters
and M.S. Faith (eds). [Date accessed: July 15th
 EVOLUTIONARY PSYCHOLOGY AND PUBLIC HEALTH
2019] https://eur01.safelinks.protection.out-
look.com/?url=http%3A%2F%2Fwww.
child-encyclopedia.com%2Fchild-
nutrition%2Faccording-experts%2Fgenetic-
influences-child-eating-behaviour&amp;data
=02%7C01%7C%7Cfb2a1bd6fa7f449a946
008d70933e623%7C1faf88fea9984c5b93c
9210a11d9a5c2%7C0%7C1%7C63698799
0829674502&amp;sdata=%2BFkWye4%2F
G61UawslrZ%2FotY6r6naJkGGIwP8t6M9lUs
U%3D&amp;reserved=0.
20 Swinburn, B.A., Sacks, G., Hall, K.D.,
McPherson, K., Finegood, D.T., Moodie, M.L.
et  al. (2011). The global obesity pandemic:
shaped by global drivers and local
environments. The Lancet. 378. 9793.
804–814.
21 United Nations. (2018). News: 68% of the
world population projected to live in urban
areas by 2050. [Date accessed: 12th March
2019] www.un.org/development/desa/en/
news/population/2018-revision-of-world-
urbanization-prospects.html.
22 Foster, C., Hillsdon, M., Cavil, N., Allender,
S. and Cowburn, G. (2005). Understanding
participation in sport: a systematic review.
London: Sport England.
23 Goode, W.J. (1964). The Family. Englewood
Cliffs, NJ: Prentice-Hall Inc.
24 Krebs, C.J. (1978). Ecology. The experimental
analysis of distribution and abundance. New
York, United States: Joanna Cotler Books.
ISBN 10: 0060437715.(Second edition).
25 Maynard-Smith, J. (1978). Optimization
theory in evolution. Annual Review of
Ecology and Systematics. 9. 31–56.
26 Winterhalder, B. (1986). The analysis of
hunter-gatherer diet: stalking an optimal
foraging model. In Food Preferences and
Aversions, M. Harris and E.A. Ross (eds).
Philadelphia, PA: Temple University Press.
311–339.
27 Power, M.L. and Schulkin, J. (2009). The
evolution of obesity. Baltimore, MD: John
Hopkins University Press.
28 Dallman, M.F., Pecararo, N. and la Fleur, S.E.
(2005). Chronic stress and comfort foods:
self-medication and abdominal obesity. Brain,
Behaviour and Immunity. 19. 275–280.
29 Lancet editorial. (2011). Urgently needed: a
framework convention for obesity control.
378. 741.
139
30 Nettle, D. (2010). Why are there social
gradients in preventative health behaviour?
A perspective from behavioural ecology.
PLoS One. 5. 10. 1–6.
31 Ball, K. and Crawford, D. (2005).
Socioeconomic status and weight change in
adults: a review. Social Science and Medicine.
60. 1987–2010.
32 Monteiro, C.A., Conde, W.L., Lu, B. and
Popkin, B.M. (2004). Obesity and inequities in
health in the developing world. International
Journal of Obesity. 28. 1181–1186.
33 van Zon, S.K.R., Bültmann, U., Mendes de
Leon, C.F. and Reijneveld, S.A. (2015).
Absolute and relative socioeconomic health
inequalities across age groups. PLoS One.
10. 12. e0145947.
34 Edin, K., Boyd, M., Mabli, J., Ohls, J.,
Worthington, J., Greene, S. et  al. (2013).
SNAP Food Security In-Depth Interview
Study: Final Report. Washington, DC: US
Department of Agriculture, Food and
Nutrition Service.
35 Gordon-Larsen, P., Nelson, M.C., Page, P.
and Popkin, B.M. (2006). Inequality in the
built environment underlies key health
disparities in physical activity and obesity.
Pediatrics. 117. 2. 417–424.
36 Cummins, S. and Macintyre, S. (2006). Food
environments and obesity: neighbourhood
or nation? International Journal of Epidemi-
ology. 35. 100–104.
37 Wang, Y. and Beydoun, M.A. (2007). The
obesity epidemic in the United States –
gender, age, socioeconomic, racial/ethnic,
and geographic characteristics: a systematic
review and meta-regression analysis.
Epidemiologic Reviews. 29. 6–28.
38 Starratt, V.G. (2017). Evolved psychological
mechanisms. In Encyclopedia of Personality
and Individual Differences, V. Zeigler-Hill and
T.K. Shackelford (eds). Cham, Switzerland:
Springer International Publishing. doi:10.1007/
978-3-319-28099-8_1633-1.
39 Chisholm, J.S. (1993). Death, hope, and sex:
life-history theory and the development of
reproductive strategies. Current
Anthropology. 34. 1–24.
40 Tooby, J. and Cosmides, L. (1990). On the
universality of human nature and the unique-
ness of the individual: the role of genetics and
adaptation. Journal of Personality. 58. 17–67.
140 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
41 Wilson, D.S. (1994). Adaptive genetic varia-
tion and human evolutionary psychology.
Ethology and Sociobiology. 15. 219–235.
42 Tooby, J. and Cosmides, L. (1995). Mapping
the evolved functional organization of mind
and brain. In The Cognitive Neurosciences,
M.S. Gazzaniga (ed.). 3995. Cambridge,
MA: MIT Press. 1185–1197.
43 Mastrandea, R., Gabrielli, A., Piras, F.,
Spalletta, G., Caldarelli, G. and Gili, T. (2017).
Organization and hierarchy of the human
functional brain network lead to a chain-like
core. Scientific Reports. 7. 4888. 1–13.
doi:10.1038/s41598-017-04716-3.
44 Nolte, J. (2002). The Human Brain: An
introduction to its functional anatomy.
Missouri, United States: Mosby.
45 Caramazza, A. and Coltheart, M. (2006).
Cognitive neuropsychology twenty years
on. Cognitive Neuropsychology. 23. 1.
3–12.
46 Russell, S.J., Hughes, K. and Bellis, M.A.
(2016). Impact of childhood experience and
adult well-being on eating preferences and
behaviours. BMJ Open. 6. 1. e007770.
doi:10.1136/bmjopen-2015-007770.
47 Johnson, J.G., Cohen, P., Kasen, S. and
Brook, J.S. (2002). Childhood adversities
associated with risk for eating disorders or
weight problems during adolescence or early
adulthood. American Journal of Psychiatry.
159. 394–400.
48 Ebbeling, C.D., Pawlak, D.B. and Ludwig,
D.S. (2002). Childhood obesity: public health
crisis, common sense cure. Lancet. 360.
473–482.
49 Koch, F.S., Sepa, A. and Ludvigsson, J.
(2008). Psychological stress and obesity.
Journal of Pediatrics. 153. 6.
50 Gustafson, T.B. and Sarwer, D.B. (2004).
Childhood sexual abuse and obesity. Obesity
Reviews. 5. 129–135.
51 Hassel, S., McKinnon, M.C., Cusi, A.M. and
Macqueen, G.M. (2011). An overview of
psychological and neurobiological mecha-
nisms by which early negative experiences
increase risk of mood disorders. Journal of
the Canadian Academy of Child and Adoles-
cent Psychiatry. 20. 4. 277–288.
52 Borgerhoff-Mulder, M. and Schacht, R.
(2012). Human behavioural ecology. In eLS.
Chichester: John Wiley & Sons, Ltd.
doi:10.1002/9780470015902.a0003671.
pub2.
53 Pechey, R., Jebb, S.A., Kelly, M.P., Almiron-
Roig, E., Conde, S., Nakamura, R. et  al.
(2013). Socioeconomic differences in pur-
chases of more vs. less healthy foods and
beverages: analysis of over 25,000 British
households in 2010. Social Science and
Medicine. 92. 22–26.
54 Kipping, R.R., Smith, M., Heron, J., Hickman,
M. and Campbell, R. (2015). Multiple risk
behaviour in adolescence and socio-
economic status: findings from a UK birth
cohort. European Journal of Public Health.
25. 1. 44–49.
55 Engelhardt, C. and Wagener, A. (2014).
Biased Perceptions of Income Inequality and
Redistribution. CESifo Working Paper Series
No. 4838. Munich: CESifo Group.
56 Marsheb, R.M. and Grilo, C.M. (2006). Emo-
tional overeating and its associations with
eating disorder psychopathology among over-
weight patients with binge eating disorder.
International Journal of Eating Disorders. 39.
141–146.
57 Bateson, P. and Gluckman, P. (2011). Plastic-
ity, Robustness, Development and Evolution.
Cambridge: Cambridge University Press.
58 Kim, S. and Lee, D. (2011). Prefrontal cortex
and impulsive decision making. Biological
Psychiatry. 69. 12. 1140–1146.
59 Del Giudice, M., Ellis, B. and Shirtcliff, E.A.
(2011). The adaptive calibration model of
stress responsivity. Neuroscience &
Biobehavioral Reviews. 35. 1562–1592.
60 Wiers, R.W., Bartholow, B.D., van den
Wildenberg, E., Thush, C., Engels, R.C.M.E.,
Sher, K.J. et al. (2007). Automatic and
controlled processes and the development of
addictive behaviors in adolescents: a review
and a model. Pharmacology, Biochemistry
and Behavior. 86. 263–283.
61 Carver, C.S. (2005). Impulse and constraint:
perspectives from personality psychology,
convergence with theory in other areas, and
potential for integration. Personality and
Social Psychology Review. 9. 312–333.
62 Hofmann, W., Friese, M. and Wiers, R.
(2008). Impulsive versus reflective influences
on health behavior: a theoretical framework
and empirical review. Health Psychology
Review. 2. 2. 111–137.
 EVOLUTIONARY PSYCHOLOGY AND PUBLIC HEALTH
63 Hansen, E.B. and Breivik, G. (2001).
Sensation seeking as a predictor of positive
and negative risk behaviour among
adolescents. Personality and Individual
Differences. 30. 627–640.
64 Frankenhuis, W.E., Panchanathan, K. and
Nettle, D. (2016). Cognition in harsh and
unpredictable environments. Current
Opinion in Psychology. 7. 76–80.
65 Bogg, T. and Roberts, B. W. (2004).
Conscientiousness and health-related
behaviors: A meta-analysis of the leading
behavioral contributors to mortality.
Psychological Bulletin. 130. 887–919.
66 Grano, N., Virtanen, M., Vaherta, J.,
Elovainio, M. and Kivimäki, M. (2004).
Impulsivity as a predictor of smoking and
alcohol consumption. Personality and
Individual Differences. 37. 1693–1700.
67 Read, D., Frederick, S., Orsel, B. and
Rahman, J. (2005). Four score and seven
years from now: the date/delay effect in
temporal discounting. Management Science.
51. 1326–1335.
68 Daugherty, J.R. and Brase, G.L. (2009).
Taking time to be healthy: predicting health
behaviors with delay discounting and time
perspective. Personality and Individual
Differences. 48. 202–207.
69 Maynard-Smith, J. (1972). On Evolution.
Edinburgh: Edinburgh University Press.
70 Russell, S.J. (2017). Applying evolutionary
principles to the obesity problem and other
issues in public health. [Date accessed: 18th
March 2019] https://ethos.bl.uk/OrderDetails.
do?uin=uk.bl.ethos.733959. doi: 10.24377/
LJMU.t.00007660.
71 Wells, J.C.K., Nesse, R.M., Sear, R.,
Johnstone, R.A. and Stearns, S.C. (2017).
Evolutionary public health: introducing the
concept. Evolutionary Public Health. 390.
10093. 500–509.
72 Randolph, M.E., Pinkerton, S.D., Bogart,
L.M., Cecil, H. and Abramson, P.R. (2007).
Sexual pleasure and condom use. Archives of
Sexual Behavior. 36. 6. 844–848.
73 Spring, B., Moller, A.C. and Coons, M.J.
(2012). Multiple health behaviours: overview
and implications. Journal of Public Health.
34. i3–10.10.1093/pubmed/fdr111.
74 Chernew, M., Frick, K. and McLaughlin, G.
(1997). The demand for health insurance
141
coverage by low-income workers: can reduced
premiums achieve full coverage? HSR: Health
Services Research. 32. 4. 453–470.
75 Beckert, J. and Lutter, M. (2012). Why the
poor play the lottery: sociological approaches
to explaining class-based lottery play.
Sociology. 47. 6. 1152–1170.
76 Kruger, D.J. and Kruger, J.S. (2016).
Psychometric assessment of human life
history predicts health related behaviors.
Psychological Topics. 25. 19–28.
77 World Health Organization. (2018). Obesity
and overweight: key facts. [Accessed: 20th
March 2019] www.who.int/news-room/fact-
sheets/detail/obesity-and-overweight.
78 McGill, R., Anwar, E., Orton, L., Bromley, H.,
Lloyd-Williams, F., O’Flaherty, M. et  al.
(2015). Are interventions to promote healthy
eating equally effective for all? Systematic
review of socioeconomic inequalities in
impact. BMC Public Health. 15. 457.
doi:10.1186/s12889-015-1781-7.
79 Omenn, G.S. (2010). Evolution and public
health. Proceedings of the National Academy
of Sciences. 107. 1. 1702–1709.
CASE STUDY REFERENCES
80 Stern, A. and Markel, H. (2005). The history
of vaccines and immunization: familiar
patterns, new challenges. Health Affairs. 24.
3: The Vaccine Enterprise.
81 World Health Organization. (2017). The
power of vaccines: still not fully utilized.
World Health Organization. [Date accessed:
6th January 2019] https ://www.who.int/
publications/10-year-review/vaccines/en/.
82 Fenner, F., Henderson, D.A., Arita, I., Jezek,
Z. and Ladnyi, I.D. (1988). Smallpox and its
eradication, pp. 332–335. Geneva,
Switzerland: World Health Organization.
83 Andre, F.E., Booy, R., Bock, H.L., Clemens,
J., Datta, S.K., John, T.J. et  al. (2008).
Vaccination greatly reduces disease,
disability, death and inequity worldwide.
Bulletin of the World Health Organization.
86. 2. 81–160.
84 Kata, A. (2012). Anti-vaccine activists web
2.0 and the postmodern paradigm: an
overview of tactics and tropes used online by
142 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
the anti-vaccination movement. Vaccine. 30.
25. 3778–3789.
85 Bandari, R., Zhou, Z., Qian, H., Tangherlini,
T.R. and Roychowdhury, V.P. (2017). A resist-
ant strain: revealing the online grassroots rise
of the antivaccination movement. Computer.
50. 11. 60–67.
86 Wolfe, R.M., Sharp, L.K. and Lipsky, M.S.
(2002). Content and design attributes of
antivaccination web sites. JAMA. 287. 24.
3245–3248.
87 Wong, J.C. (2019). Anti-vaxx ‘mobs’: doctors
face harassment campaigns on Facebook.
[Date accessed: 6th January 2019] www.the-
guardian.com/technology/2019/feb/27/
facebook-anti-vaxx-harassment-campaigns-
doctors-fight-back.
88 Leask, J. (2015). Should we do battle with
anti-vaccination activists? Public Health
Research and Practice. 30. 25. 2: e2521515.
89 Jacobson, R.M., Targonski, P.V. and Poland,
G.A. (2007). A taxonomy of reasoning flaws
in the anti-vaccine movement. Vaccine. 25.
16. 3146–3152.
90 Gerber, J.S. and Offit, P.A. (2009). Vaccines
and autism: a tale of shifting hypotheses.
Clinical Infectious Diseases. 48. 4. 456–461.
91 Destefano, F., Gu, D., Kramarz, P., Truman,
B.I., Iademarco, M.F., Mullooly, J.P. et  al.
(2002). Childhood vaccinations and risk of
asthma. The Pediatric Infectious Disease
Journal. 21. 6. 498–504.
92 Hviid, A., Hansen, J.V., Frisch, M. and
Melbye, M. (2019). Measles, mumps, rubella
vaccination and autism – a nationwide
cohort study. Annals of Internal Medicine.
170. 8. 513–520. doi:10.7326/M18-2101.
93 National Health Service. (2017). Childhood
Vaccination Coverage Statistics, England,
2016–17. NHS Digital. [Date accessed: 8th
January 2019] https://digital.nhs.uk/data-
and-information/publications/statistical/nhs-
immunisation-statistics/childhood-vaccination-
coverage-statistics-england-2016-17.
94 Centers for Disease Control and Prevention.
(2018). Vaccination Coverage Among
Children Aged 19–35 Months – United
States, 2017. [Date accessed: 8th January
2019] www.cdc.gov/mmwr/volumes/67/wr/
mm6740a4.htm.
95 Majumder, M.S., Cohn E.L., Mekaru, S.R.,
Huston, J.E. and Brownstein, J.S. (2015).
Substandard vaccination compliance and the
2015 measles outbreak. JAMA Pediatrics.
169. 5. 494–495.
96 World Health Organization. (2018).
Emergencies preparedness, response:
measles – Japan. [Date accessed: 20th January
2019] www.who.int/csr/don/20-june-2018-
measles-japan/en/.
97 Dyer, O. (2019). Philippines measles
outbreak is deadliest yet as vaccine scepticism
spurs disease comeback. BMJ. 364. l739.
doi:https://doi.org/10.1136/bmj.l739.
98 World Health Organization. (2018b).
Measles cases spike globally due to gaps in
vaccination coverage. [Date accessed: 20th
January 2019] www.who.int/news-room/
detail/29-11-2018-measles-cases-spike-
globally-due-to-gaps-in-vaccination-coverage.
99 Amin, A.B., Bednarczyk, R.A., Ray, C.E.,
Melchiori, K.J., Graham, J., Huntsinger, J.R.
et  al. (2017). Association of moral values
with vaccine hesitancy. Nature Human
Behaviour. 1. 873–880.
100 Graham, J., Nosek, B.A., Haidt, J., Iyer, R.,
Koleva, S. and Ditto, P.H. (2011). Mapping
the moral domain. Journal of Personality and
Social Psychology. 101. 366–385.
101 Anderson, R.M. and May, R.M. (1991).
Infectious Diseases of Humans: Dynamics
and Control. Oxford: Oxford University
Press.
102 Centers for Disease Control and Prevention.
(2015). Smallpox: Disease, Prevention, and
Intervention. Retrieved 13 March 2015.
[Date accessed: 20th January 2019] https://
stacks.cdc.gov/view/cdc/27929.
103 O’Gorman, R., Wilson, D.S. and Miller, R.R.
(2005). Altruistic punishing and helping
differ in sensitivity to relatedness, friendship
and future interactions. Evolution and
Human Behavior. 26. 375–387.
104 West, S.A., Griffin, A.S. and Gardner, A.
(2007). Evolutionary explanations for
cooperation: review. Current Biology. 17.
16. 661–672.
105 Wade, M.J. and Breden, F. (1980). The
evolution of cheating and selfish behavior.
Behavioral Ecology and Sociobiology. 7. 3.
167–172.
106 Trivers, R.L. (1971). The evolution of
reciprocal altruism. The Quarterly Review of
Biology. 46. 1. 35–57.
 EVOLUTIONARY PSYCHOLOGY AND PUBLIC HEALTH
107 Hershey, J.C., Asch, D.A., Thumasathit, T.,
Meszaros, J. and Waters, V.V. (1994). The
roles of altruism, free riding, and
bandwagoning in vaccination decisions.
Organizational Behavior and Human Decision
Processes. 59. 2. 177–187.
108 MacDonald, N.E. (2014). Vaccine hesitancy:
definition, scope and determinants. Vaccine.
33. 34. 4161–4164.
109 Larson, H.J., Schulz, W.S., Tucker, J.D. and
Smith, D.M.D. (2015). Measuring vaccine
confidence: introducing a global vaccine
confidence index. Version 1 PLoS Currents.
7: ecurrents.outbreaks.ce0f6177bc97332
602a8e3fe7d7f7cc4.
110 Dubé, E., Vivion, M. and MacDonald, N.E.
(2014). Vaccine hesitancy, vaccine refusal
and the anti-vaccine movement: influence,
impact and implications. Expert Review of
Vaccines. 14. 1. 99–117.
111 Leask, J., Willaby, H.W. and Kaufman, J.
(2014). The big picture in addressing vaccine
hesitancy. Human Vaccines and Immunothera-
peutics. 10. 9. 1–3.
112 Logan, J., Nederhoff, D., Koch, B., Griffith,
B., Wolfson, J., Awanal, F.A. et  al. (2018).
‘What have you HEARD about the HERD?’
Does education about local influenza
vaccination coverage and herd immunity
affect willingness to vaccinate? Vaccine. 36.
4118–4125.
113 Odone, A., Ferrari, A., Spagnoli, F.,
Visciarelli, S., Shefer, A., Pasquarella, C. et al.
143
(2014). Effectiveness of interventions that
apply new media to improve vaccine uptake
and vaccine coverage: a systematic review.
Human Vaccines and Immunotherapeutics.
11. 1. 72–82. https://doi.org/10.4161/
hv.34313.
114 Nyhan, B., Reifler, J., Richey, S. and Freed,
G.L. (2014). Effective messages in vaccine
promotion: a randomized trial. Pediatrics.
133. 4. 1–8.
115 Dube, E., Gagnon, D. and MacDonald,
N.E. (2015). Strategies intended to address
vaccine hesitancy: review of published
reviews. Vaccine. 33. 4191–4203.
116 Cairns, G., MacDonald, L., Angus, K., Walker,
L., Cairns-Haylor, T. and Bowdler, T. (2012).
Systematic literature review of the evidence for
effective national immunisation schedule
promotional communications. Stockholm:
European Centre for Disease Prevention and
Control. [Date accessed 20th January 2019]
https://www.ecdc.europa.eu/sites/default/files/
media/en/publications/Publications/Literature-
review-national-immunisation-schedule-
promotional-communications.pdf.
117 Arie, S. (2017). Compulsory vaccination
and growing measles threat. British Medical
Journal. 358. j3429. doi:10.1136/bmj.j3429.
118 Wigham, S., Ternent, L., Bryant, A.,
Robalino, S., Sniehotta, F.F. and Adams, J.
(2014). Parental financial incentives for
increasing preschool vaccination uptake:
systematic review. Pediatrics. 134. 4. e1117.

Animal Ethics and Evolutionary
Psychology
INTRODUCTION
Evolutionary psychology examines the
human mind through the lens of evolution to
understand the functions of our psychological
adaptations such as motivations, emotions,
and cognitions. Humans have many interac-
tions with nonhuman animals (hereafter just
‘animals’, although the term ‘nonhuman ani-
mals’ makes an important philosophical
point), most of which are fairly recent (e.g.,
cats as companion animals), but some go
much further back in our history (e.g., hunt-
ing animals for food). We use animal bodies
for fur, meat, lactation, eggs, labor, and scien-
tific study; the scale of animal use is enor-
mous, trillions of animals are killed for food
each year (Wiblin and Bollard, 2017). In the
last few hundred years, the average level of
human suffering has decreased dramatically,
but the total amount of animal suffering due
to human actions has skyrocketed. All around
us we can see examples of individuals being
7
Diana Santos Fleischman
exceedingly altruistic to favored animals, but
also industrialized cruelty towards less
favored animals at an incomprehensible
scale. While we should have no expectation
that human morality will be rational or con-
sistent, this chapter grapples with the fact that
our treatment of animals deviates very far
from any coherent, rational morality. In terms
of overall ‘suffering footprint’, human mal-
treatment of animals may be the biggest ethi-
cal issue in the world, and evolutionary
psychology can give us deep insights into
both the problem and possible solutions.
ANIMAL ETHICS
Animal ethics has different meanings among
groups of philosophers, scientists, and the
public. Examining how our evolutionary
psychology obscures consistent ethics
requires some consideration of what would
 ANIMAL ETHICS AND EVOLUTIONARY PSYCHOLOGY
serve as an ethical baseline for comparison
(Fleischman, 2018). Views about human
obligations to animals, or lack thereof, have
proliferated in philosophy. Some philoso-
phers argue it’s wrong to own animals or use
them in any way (Francione, 2015), whereas
others argue that we have no obligation to
animals because they cannot make social
contracts and are thus not part of our moral
community (Machan, 2004). Of all the main-
stream philosophical approaches to animal
ethics, utilitarianism and consequentialism
have been most positively disposed to mor-
ally consider animals than other frameworks
(Beauchamp and Frey, 2011).
Utilitarianism defines what is good as what
maximizes happiness or pleasure and mini-
mizes suffering, across all sentient beings
(those capable of experiencing happiness or
suffering) (Greene, 2013). This chapter rests
on the basic evolutionary insight that verte-
brate animals like mammals and fish evolved
the capacity to feel pain and pleasure, and
thus the capacity to suffer. Further, it rests
on a normative moral stance that sentience
should be the basis for moral consideration,
that suffering is bad, and that reducing suf-
fering is good. I do not rely on any concept
of ‘animal rights’, nor do I assume that using
animals as a means to human ends is always
immoral (Regan, 2004), although I believe
the moral foundations of these ideas are also
rooted in evolutionary psychology. For my
normative moral claim that suffering is bad
and alleviating suffering is good, most phi-
losophers appeal to the reader’s personal
experience with suffering, or take the idea
that suffering is bad as obvious (‘a priori’
or ‘axiomatic’) (but see also Kahane, 2009).
Evolutionary theory has influenced many to
adopt an ethical stance that we should ascribe
moral standing and consideration on the basis
of the ability to suffer (Singer, 2011), other-
wise known as ‘sentientism’ (Ryder, 1991;
‘Sentientism’, 2019). This chapter analyzes
where our evolutionary psychology is con-
sistent with and deviates from sentientism as
a moral baseline.
145
EVIDENCE FOR ANIMAL SENTIENCE
How can we establish which animals are sen-
tient, and therefore deserving of moral consid-
eration? Sentience is the ability to experience
pain and pleasure subjectively. First let’s dis-
tinguish between the ability to react to tissue
damage, and the ability to suffer. Nociception,
or the experience of pain, is the simple and
ancient capacity of most animals to respond to
injuries that cause tissue damage. Nociception
is specific to animals with a nervous system,
but even simpler organisms may have a simi-
lar capacity to respond to harmful stimuli
(Tomasik, 2018a). Behavioral evidence of
nociception is, for example, a shrimp groom-
ing an injured antenna (Elwood, 2011) or,
physiologically, the measurement of neurons
firing in response to sensory stimulation
(Braithwaite, 2010). Sentience and the ability
to suffer is the subjective awareness of pleas-
ure and pain and can be demonstrated when
the response to stimuli is more complex
than a simple response to physical damage
(Braithwaite, 2010; Elwood, 2011).
Considering an evolutionary and functional
perspective, we can infer that subjective aware-
ness of suffering evolved to prevent and man-
age bodily damage. If we take as given that
humans can suffer, and suffering has important
adaptive functions in enabling our survival and
reproduction, it’s parsimonious to assume that
sentience and suffering evolved in other related
animals, including many other vertebrates
(Tomasik, 2017). There is a solid foundation
of evidence that vertebrates and even some
invertebrates evolved both nociception and
sentience (Braithwaite, 2010). Vertebrates as a
group generally have the same neurons, syn-
apses, and other neural hardware associated
with the ability to feel pain found in sentient
humans. Fish brains, once thought to lack the
fundamental hardware of sentience, have been
found to have a brain region similar to the lim-
bic system such that they may have the abil-
ity to ‘process information with an emotional
component’ (Braithwaite, 2010: 102).
146 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Animal responses to pain, such as soliciting
help, and avoiding stimuli previously asso-
ciated with pain, are behavioral evidence
of sentience. Many studies have shown that
invertebrates – widely thought to be incapa-
ble of sentience – show responses consistent
with subjective awareness of pain. For exam-
ple, hermit crabs have been shown to make
adaptive tradeoffs when exposed to shock,
choosing to endure more painful shock in
a high-quality as opposed to a low-quality
shell (Appel and Elwood, 2009). Trout given
a painful injection in their lips failed to show
the normal neophobic response to a novel
stimulus (a colorful block tower), compared
to trout in the control condition; the trout’s
distraction from normal behavior because of
pain suggests a subjective awareness of pain,
and thus suffering and sentience (Braithwaite,
2010; Sneddon et al., 2003). Using crabs and
fish as examples is instructive, because they
show better objective evidence for suffering
than human neonates do (Braithwaite, 2010:
153) – but babies would almost certainly be
more likely to get the benefit of our doubt.
This is not to say that fish are equally sen-
tient with cats, dogs, babies, or adult humans.
There are many potentially good reasons to
prioritize some animals over others by virtue
of their greater ability to suffer, but the influ-
ence of, for example, brain size on degree of
potential suffering is beyond the scope of this
chapter (see Tomasik, 2019).
The vast amount of sentience that evolved
across millions of species in our world – and
the resulting potential for suffering across
trillions of animals – can feel overwhelm-
ing. But that doesn’t mean it is not true – and
evolutionary psychologists have been coura-
geous in confronting other emotionally chal-
lenging, counter-intuitive truths. Evolution
may have created endless forms most beauti-
ful, but it would never have passed an eth-
ics review board (Bostrom, 2016: 188).
Evolutionary psychologists should take seri-
ously the likelihood that evolution favors
widespread sentience across species, but not
widespread altruism to other species, and this
sets the stage for a planet filled with s­ entient
suffering both before and after humans
achieved the technological means to exploit
other species on an industrial scale.
Natural-Born Speciesists
There are two central questions with regards
to human morality towards animals: why do
we value animals less than humans morally,
and why are our moral attitudes towards ani-
mals so inconsistent? Both moral anthropo-
centrism and speciesism describe the concept
of valuing human lives over animal lives,
although speciesism implies that this valua-
tion is a form of prejudice due to mere spe-
cies membership (Caviola, 2019). Most
people value members of certain species
above and beyond their ability to suffer, for
example valuing insensate humans (like those
in a persistent vegetative state) more than
chimpanzees, and valuing dogs more than
pigs, even though they are similarly sentient.
Valuing humans over animals seems to be a
human moral universal. In one study, millions
of participants all over the world overwhelm-
ingly choose to save the life of a human over
an animal, or several animals (Awad et  al.,
2018). In another study, participants most
often choose to save one human over the lives
of several endangered animals (like gorillas)
(Petrinovich et  al., 1993). This valuation is
often reversed if the animal at risk is their pet.
When Topolski et  al. (2013) asked partici-
pants who they would save if a bus was immi-
nently going to hit their pet or a foreign
tourist, 40% chose to save their pet. ‘The only
consistency in the way humans think about
animals is inconsistency’ (H.Herzog, 2010:
14). Arguably, it can be rational to prioritize
some interests over others on the basis of
sentience, but, unsurprisingly, humans are not
doing any coherent form of ethical calculus
when they choose actions or decide on the
morality of those actions.
Is there an inherent human moral response
towards animals? The study of how humans
 ANIMAL ETHICS AND EVOLUTIONARY PSYCHOLOGY
and animals interact has taken off in the last
two decades with the rise of anthrozool-
ogy, also known as human–animal relations
(Amiot and Bastian, 2015; H. Herzog, 2010;
Serpell, 1996). Anthrozoology is a diverse
field, investigating topics like the therapeu-
tic properties of living with dogs, the possi-
ble link between animal abuse and criminal
behavior, and the personality traits of animal-
rights activists (H. Herzog, 2010). These
fields have often tried to explain human
moral anthropocentrism and moral inconsist-
ency with descriptive frameworks like car-
nism (the social ideology that supports meat
eating) (Joy, 2011), moral disengagement
(i.e., we distance ourselves from our humane
standards to harm animals) (Bandura, 1999;
Vollum et  al., 2004), terror management
theory (i.e., we cling to human uniqueness
and animal oppression to avoid existential
anxiety) (Marino, 2019), and speciesism
(discrimination based on species member-
ship) (Caviola et  al., 2018; Singer, 1995).
Explanations at a functional level of analysis
(Scott-Phillips et  al., 2011) and adaptation-
ist accounts are less common in anthrozool-
ogy (although see Bradshaw and Paul, 2010;
H. Herzog, 2002).
Evolutionary psychology is compat-
ible with most of the explanations of human
morality towards animals advanced by
anthrozoologists. It posits functional expla-
nations for attitudes and behaviors as reflect-
ing evolved psychological adaptations (or
their byproducts). Explanations like spe-
ciesism and cultural explanations like car-
nism are not in conflict with evolutionary
psychology (Tooby and Cosmides, 1989),
because culture is both an outgrowth of and
a support for our evolved morality (e.g., the
cultural celebration of consensual courtship
and maternal love). However, the assump-
tion of some thinkers in anthrozoology, as
well as many animal-welfare advocates,
is that humans are naturally sensitive and
morally concerned about animal suffering,
but this innate goodness is numbed by cultural
and social factors like carnism (Joy, 2003)
147
and moral disengagement. The consistent
thread in these perspectives is that violence
is deeply unnatural. However, from the per-
spective of evolutionary morality, we should
not expect sensitivity to animal suffering or
kindness to animals except to the extent that
it helped us, for example when using animals
for our benefit, or to signal our morality.
Anthropomorphism
Anthropomorphism, or ascribing human char-
acteristics to animals, is an apparently univer-
sal feature of the human mind (Urquiza-Haas
and Kotrschal, 2015). Anthropomorphism is
so naturally expressed that it’s difficult to
think about animals in non-anthropomorphic,
objective terms. Most children don’t make
clear distinctions between humans and ani-
mals, and young children usually treat ani-
mals, like family pets, as human persons
(Serpell, 1996).
Ironically, the basis for much of our moral
feelings towards animals likely evolved so
we could better exploit, kill, and eat them.
Humans and animals aren’t so different, so
the same theory of mind and empathy that
helps us predict what humans do can also be
used when hunting prey animals or avoiding
predators. Primatologists and other animal-
behavior scientists often use simple anthro-
pomorphism to make predictions; behaviorist
John Garcia stated that anthropomorphism
with regard to rat behavior ‘works better than
most learning theories’ (Serpell, 1996: 174).
When you’re using the mind-reading ability
that mostly evolved to predict the behavior of
other people, you’re bound to ascribe human
characteristics to animals. The ability to
imagine what it was like to be an animal has
adaptive benefit to better predict the behav-
ior of prey, predators, and dangerous animals
(H. Herzog, 2002). Some speculate that
empathy could have motivated nurturance
for domesticated species in animal husbandry
(Bradshaw and Paul, 2010) however these ani-
mals have also been bred to be cute and inspire
148 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
feelings of care. Given an adaptationist per-
spective, empathy would have been bounded
so as not to interfere with the processes of
killing, butchering, and eating animals. If
empathy is like a spotlight that illuminates
the suffering of some at the expense of oth-
ers (Bloom, 2016), this spotlight turns off
or moves on. Empathy isn’t consistently
associated with caring about animal ethics
(Kasperbauer, 2015). And biophilia, or the
desire to affiliate with nature and animals
(Wilson, 1984), often doesn’t have a loving
or caring character.
Play and Animal Abuse
One clue about evolved human morality
towards animals is how readily humans will
torment animals when it isn’t necessary, and
how much they enjoy doing it. Play is an
essential part of learning in our species and
many others. But humans playing with ani-
mals often involves both curiosity and cru-
elty (Arluke, 2002). Around the world, in
both traditional societies and Western socie-
ties, playing with animals and cruelty to
animals are commonplace in both adults and
children – not just in psychopaths.
Read some anthropological descriptions
of hunter-gatherers and you’ll see sentimen-
tal, Noble Savage views like this: ‘Children
learn to sympathize with animals and to see
animals as sentient persons sharing the for-
est world with them’ (Lew-Levy et al., 2017:
X). The implication is that hunter-gatherers
have greater moral regard for animals than
Western people, who only see meat wrapped
in cellophane at the grocery store. However,
read descriptions from thinkers who are less
attached to a Noble Savage narrative, and
you’ll get a better picture of how difficult it
may be for our species to bestow moral con-
sideration on others.
It’s common for people in more tradi-
tional societies to hurt animals for fun. Jared
Diamond describes Papua New Guinean men
amusing themselves by raising and lowering
squealing bats into a fire and dissecting them
alive for their bones (Diamond, 1993).
Men and boys are much more likely to
abuse animals than women and girls (Arluke,
2002) but this passage about Nisa, a !Kung
San woman, describes with unusual clarity the
dynamic of curious, playful cruelty, and its abil-
ity to facilitate precise prediction of animals:
A flying ant with a long, wormlike body and large,
almost transparent wings… landed in the hot
sand… Nisa saved it… and pierced it through half
the length of its body with a thin twig, leaving
the upper half with the wings and head free. She
planted the stick, with the skewered insect at the
top, upright in the ground and tapped it gently with
her fingers. The insect’s wings burst into motion, as
if in flight, propelling the free parts of its body
around and around the stick; then it stopped. Nisa
tapped the stick again and again; each time, the
insect responded with the same outpouring of
energy… What Nisa was doing… seemed like an
inexcusable torture… [But Nisa’s] head and the
upper parts of her body had begun to move rhyth-
mically. I did not understand what she was doing at
first. Then it became clear: as the insect held itself
erect, Nisa’s body also became erect; when the
insect circled, drooped, and strained, Nisa’s body did
the same. Her face and torso echoed the insect’s
plight with a wrenching subtlety and her mimicry of
its every movement was so sympathetic that the
situation took on a kind of beauty. (Shostak and
Nisa, 2004: 321)
Using animals for entertainment has been one
of the most contentious and moralized aspects
of animal use, even though the scale of suffer-
ing involved is much smaller than most modern
industrialized forms of animal use. For exam-
ple, many countries that otherwise have few
animal-protection laws have banned circuses
(‘These 27 Countries Have Banned Wild-
Animal Circuses!’, 2019, https://www.peta.org.
uk/blog/these-26-countries-that-have-banned-
wild-animal-circuses-are-making-england-
look-really-bad/). Bear-baiting, hare-coursing,
dogfighting, and cockfighting are more contro-
versial than other types of animal use, and were
banned much earlier (e.g., the UK banned bear-
baiting in 1835 and Pakistan banned bear-bait-
ing in 1890). This seems to be an area of strong
moral signaling.
 ANIMAL ETHICS AND EVOLUTIONARY PSYCHOLOGY
One reason modern people might be judg-
mental about animal entertainment is because
of the glut of other entertainment media
developed in the last few centuries. Yes, using
animals for entertainment is unnecessary, but
it seems even less necessary and thus indul-
gently cruel when so many other forms of
entertainment are available, especially other
forms of violent entertainment like combat
sports, action films and video games. Forms
of animal entertainment considered lowbrow
or ‘primitive’ may be more controversial. For
example, many have argued that forms of
animal entertainment enjoyed by the working
class, like cockfighting, have been banned
more quickly and more often than those
enjoyed by the higher classes, like fox hunt-
ing (‘Fox Hunting’, 2019).
Most of us in Western societies who would
not want to torment an animal directly, or
would be appalled to see staged animal suf-
fering intended for entertainment, are still
entertained by watching animals inflict suffer-
ing on each other in the wild, for example, in
nature documentaries. The disparity in sensi-
bilities is also well illustrated in this anecdote:
That is perhaps the hardest part of being an
anthropologist. [The hunter-gatherers I was study-
ing] sensed my weakness and would sell me all
kinds of baby animals with descriptions of what
they would do to them otherwise. I used to take
them far into the desert and release them, they
would track them, and bring them back to me for
sale again! (Pinker, 2011: 473)
These behaviors that cause suffering to ani-
mals are often practiced alongside making
offerings to animal deities, praying to the
spirit of animals after killing them, and efforts
to embody animal qualities – contrary to the
notion that cruelty requires dehumanization
or suspension of empathy. In many small-
scale societies, formalized animal totemism
often co-exists with informal animal torment.
The majority of human groups include some
conspicuous cruelty to animals, but there are
exceptions. For example, Jain monks and
nuns sweep the ground in front of them so as
to avoid inflicting any suffering on insects
149
(‘Ahimsa in Jainism’, 2019). Here it’s notable
that this requires strong spiritual and social
incentives, such as the belief that any given
human might be reincarnated as an insect.
Given our evolutionary history as omnivores,
and the ubiquity of animal abuse across cul-
tures, it seems likely that carnism and specie-
sism are outgrowths of our evolved
psychology, rather than historically novel
cultural influences that render us weirdly
insensitive to animal suffering.
Animal Abuse by Children – The
Link
Children commonly inflict suffering on ani-
mals, not just out of necessity, but for enjoy-
ment and curiosity. Here again, we see the
sentimental narrative that animal abuse is a
rare, pathological glitch in children’s funda-
mentally caring natures, and that children who
abuse animals will grow up to have other seri-
ous problems like psychopathy and criminal-
ity. Animal abuse is considered a risk factor
for violence with such certainty in the animal
advocacy community that it’s sometimes
referred to simply as ‘The Link’ (H. Herzog,
2010). Most of the evidence for an association
between childhood animal abuse and adult
violence suffers from methodological limita-
tions like retrospective reporting, and sam-
pling incarcerated criminals (Flynn, 2011).
But, consistent with the idea that insensitivity
to animal suffering is fairly standard in our
species, there isn’t replicable evidence that
animal abusers are more likely to commit vio-
lent crime (H. Herzog and Arluke, 2006;
Patterson-Kane and Piper, 2009).
Consistent with the cross-cultural ubiquity
of animal cruelty, and the historical com-
monality of using animals for entertainment,
animal abuse is normal among young peo-
ple, even now. In one study, 40% of female
college students and 66% of male college
students admitted to having abused animals
(Arluke, 2002) – and given the modern
stigma against animal abuse, this is probably
150 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
an underreported behavior. There seems to be
a moral panic about animal abuse; advocates
often depict anyone who has ever abused an
animal as likely to commit violence against
people, even though the majority of peo-
ple have, at some time, abused an animal
(Patterson-Kane and Piper, 2009). Children
might be learning about how animals ‘work’
by playing with them, developing mental
models of animal morphology and behavior
that ancestrally would have been useful for
hunting, tracking, and butchering. Indeed,
vertebrate animals’ similarity to humans
means that children may be learning more
than this. Children around the world often
practice caretaking with pets and baby ani-
mals. I speculate that violence, which is
often used to adaptively compel others to do
what you want, can be practiced and honed
similarly by playing with animals or through
cruelty.
Indeed, lacking the kinds of modern toys
that Western children have access to, ani-
mals are used in just this way, treated with
both care and cruelty. ‘Anthropologists have
observed returning hunters bringing small
wild animals back alive and promptly turning
them over to their children… these animated
toys are generally badly treated, short lived
and… end up the objects of target practice or
mutilation’ (Serpell, 1996: 68). There are two
main hypotheses about the animal-­ cruelty
association with crime: the graduation
hypothesis posits that animal abuse is ‘a form
of rehearsal for human-directed violence’,
and the deviance generalization hypothesis
posits that antisocial personality is associated
with both animal cruelty and criminal behav-
ior (Gullone, 2014). Animal cruelty might be
both a normal behavior that children perform
if unsupervised with animals, and also a form
of practice that is disproportionately attrac-
tive to children with antisocial and violent
tendencies.
Animal killing and butchering were surely
features of our evolutionary history, but mod-
ern specialization of labor means for the first
time there are workers who spend hour after
hour killing and butchering animals. Even
among slaughterhouse workers, the worker
who kills the cow, the ‘knocker’, is consid-
ered to have psychological problems com-
pared to other workers who bleed the cow
or begin to dismember it (Pachirat, 2014).
There is evidence that the presence of slaugh-
terhouses is associated with increased local
rates of violent crime and sexual offences,
relative to other industries like steel forging
(Fitzgerald et  al., 2009), but it’s unclear if
violent people are more attracted to working
in slaughterhouses, or if slaughtering animals
increases workers’ propensities for violence
towards other humans.
WHY HAS ANIMAL ADVOCACY
LAGGED BEHIND OTHER MORAL
MOVEMENTS?
Civil-rights movements, like those for the
abolition of slavery, black power, women’s
rights, worker’s rights, and gay rights, have
flourished in the last several decades, and
have reduced suffering for billions of humans
(Pinker, 2011). The animal advocacy move-
ment (including animal rights, animal wel-
fare, and animal liberation – but not
environmental protection) has not had the
same success as other sentient-rights move-
ments (Pinker, 2011). In some ways, attitudes
have changed a great deal. In a representative
sample of over 1,000 American adults,
Sentience Institute found that nearly 50%
supported a ban on slaughterhouses and fac-
tory farming (Reese, 2017). But, in that same
survey, 75% of participants believed the reas-
suring fiction that the animal products they
were eating had been humanely produced
(Reese, 2017) even though 99% of animal
products come from factory farms (Reese,
2019). A recent Gallup poll found that 32%
of Americans think that animals deserve ‘the
exact same rights as people’ (Riffkin, 2015),
up from 25% in 2003 (Moore, 2003). A study
of 3,500 Ohio residents found 81% said farm
 ANIMAL ETHICS AND EVOLUTIONARY PSYCHOLOGY
animal welfare was as important as pet ­welfare,
and 75% said farm animals should be pro-
tected from physical pain (Rauch and Sharp,
2005). These self-reported attitudes are
impressively progressive, but consumer behav-
ior has not changed that much. FewAmericans
boycott animal products. The rate of self-
identified vegetarians has hovered around 5%
in the United States for several years (Riffkin,
2015). Other reports claim there are slightly
more vegetarians. The number of people who
describe themselves as vegetarian is probably
not actually representative of boycott, because
only about one-third abstain from meat
(Cooney, 2014). Che Green, an expert on these
trends, has called vegans and vegetarians ‘a
blip on the demographic radar’ and ‘below the
margin of error for most surveys’ (Zaraska,
2016: 136).
The concern for animal wellbeing has
made uneven progress, with far more concern
about some species and some issues than
others. One illustration of the vast disparity
between the alleged human moral concern
for animal suffering, and the actual concen-
tration of animal suffering, is revealed by
patterns of charitable donations (Figure 7.1).
Farm animals, compared to all other animals,
Figure 7.1  Charitable donations towards animal organizations as compared to animal use
151
experience the vast majority of suffering and
death with more than 99% of farm animals
living on factory farms (Reese, 2019). Farm
animals received only $20 million in charitable
donations in 2015, compared to $1.2 billion
donated to animal shelters for pet species like
dogs and cats (Bockman, 2016).
Of domesticated land animals used and killed by
humans in the United States, over 99.6% are
farmed land animals, about 0.2% are animals used
in laboratories, 0.07% are used for clothing, and
0.03% are killed in companion animal shelters.
However, about 66% of donations to animal chari-
ties in the United States go to companion animal
shelters, 32% go to groups with mixed or other
activities, and just 0.8% of donations go specifically
to farmed animal organizations, while 0.7% go to
laboratory animal organizations. (Bockman, 2016)
A couple of caveats are that the ‘other’ cate-
gory does include some farm animal dona-
tions, because it includes large organizations
that engage in diverse animal advocacy cam-
paigns (e.g., People for the Ethical Treatment
of Animals (PETA)). ‘Other’ also includes
environmental charities and wildlife preserva-
tion. Also, importantly, the ‘animals used and
killed’ number does not include fish, which
would completely dominate the left panel.
152 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Fish are probably killed in the trillions, at a
rate greater than all other animals combined
(Mood and Brooke, 2010). This is another
way that human behavior is inconguous with
stated concern. If humans cared about reduc-
ing animal suffering and acted in keeping
with this concern, they would not give a dis-
proportionate amount of their donations to
cat and dog shelters.
Below I will address the psychology of how
and why humans morally value and devalue
animals. First, I’ll discuss kin selection, and
the empathy elicited by cuteness and neoteny
– two intertwined factors that account for our
kindness towards companion animals like
dogs and cats. Second, I’ll discuss disgust
and food aversion, meat consumption, and the
future of meat eating. Empathy, disgust, and
food preferences show significant sex differ-
ences, and I’ll discuss how they manifest in
morality. Lastly, I’ll talk about how morality is
socially signaled, and how this virtue signaling
plays out in animal care and attitudes towards
the animal-focused moral minority, such as
vegans. The view of decision-making in the
moral domain in this chapter is consistent with
the popular metaphor of the elephant and the
rider (Haidt, 2012; Simler and Hanson, 2017).
Emotions and psychological mechanisms
like the cuteness response, disgust, reputation
management, and empathy guide our moral
decisions in often irrational directions, like an
elephant deciding which path to take. We iden-
tify as the rider, the part we are most conscious
of, and we later attribute our moral decisions
to rational processes rather than the ‘hot’ emo-
tional cognition of the elephant, the part that
controls our behavior.
Kin Selection and the Cuteness
Response
Prominent anthrozoologist James Serpell
defines pets as ‘animals we live with, with no
obvious function’ (H. Herzog, 2010: 72).
Other animals don’t usually keep pets. Cross-
species friendships sometimes happen (see
viral videos of cats who love rats and hippos
who love tortoises, for example), but they are
almost always the product of an artificial envi-
ronment (H. Herzog, 2014). Pet-keeping
seems uniquely human. Two interesting
exceptions in the wild are a dolphin who
adopted a melon-headed whale (Carzon et al.,
2019), and a marmoset adopted into a group
of capuchin monkeys (Izar et al., 2006). In the
first case, the dolphin nursed and cared for
the whale, and in the second case, two female
capuchins provisioned the marmoset. In both
cases, arguably, the adopted animal appears to
be a neotenous version of the animals’ own
young. This illustrates a major psychological
means through which humans integrate ani-
mals more centrally into their moral worlds:
kin selection and empathy for cuteness. Both
cases also illustrate how this cuteness-based
empathy is more motivating for females than
for males. Humans keep a wide variety of
pets, from insects to horses, but here I’ll focus
mostly on dogs, who seem to be the animals
who most often reverse human speciesism.
The psychological mechanisms motivating
kin selection may cause humans to value dogs
and cats much more than other comparably
sentient animals. Evolution promotes pass-
ing on our genes, including in other members
of our families (Foster et al., 2006). Because
cooperating in groups is adaptive, we may
be more likely to interpret ambiguous cues
as evidence of genetic association (Park and
Ackerman, 2011). In this way, kin selection
means that we tend to be more altruistic to
those who show cues of being members of
our ingroup – whether these cues are cultural
(Kurzban et  al., 2001) or physical (Krupp
et al., 2011). There are other possible indica-
tors of genetic relatedness, like psychological
similarity (Park and Schaller, 2005), and time
spent growing up together (Lieberman et al.,
2007).
It’s unclear if our minds are simply indis-
criminate about what cues we take as indi-
cators of genetic relatedness. Some animals
indisputably occupy a familial role, not only
in Western countries but also among many
 ANIMAL ETHICS AND EVOLUTIONARY PSYCHOLOGY
hunter-gatherers. New Guineans, which I
earlier described abusing bats, treat pigs as
members of their families; piglets sleep in
the same hut with their human families and
New Guinean women often nurse piglets at
the breast (Diamond, 1993). In the United
States, 91% of dog and cat owners reported
that their pet was a part of their family, and
20% of dog owners report giving their dogs
birthday gifts (‘Pets Really Are Members of
the Family’, 2011).
We attend to physical similarity when eval-
uating species and breeds for special treat-
ment. For example, people are more likely
to support causes to save endangered species
that have more in common with humans, like
great apes, and researchers who do invasive
scientific experiments on monkeys are most
likely to be threatened by animal activists (H.
Herzog, 2010). Even so, most people (around
75%) would choose to save one person over
five endangered lowland gorillas (Petrinovich
et al., 1993).
Dogs show resemblance to humans in
their facial musculature: domestication has
changed dog faces compared to wolf faces
to be able to display more humanlike expres-
sions (Kaminski et  al., 2019). There is also
evidence that people resemble their dogs
(Nakajima et  al., 2009), and that people
choose dogs who resemble them (Roy and
Nicholas, 2004). People also tend to think
their dogs are psychologically similar to
them. Dog breeds go in and out of fashion
(Ghirlanda et al., 2013), but the proliferation
of so many dog breeds with different appear-
ances and personalities may have been driven
by the kin-selection mechanisms of different
individuals and ethnic groups with different
implicit criteria for similarity.
Cute Ethics
In ethics, cuteness doesn’t count. (Cohen, 2009)
Because pets seem to occupy a place in the
family as surrogate children, cues of kinship
153
(genetic similarity) and cuteness are difficult
to disentangle as contributors to this unusual
moral relationship. Because there is no word
in English for the especially cute emotional
repertoire elicited by cuteness, I’ll use ‘cute-
ness response’. The cuteness response elicits
nurturance but also inspires mentalizing and
anthropomorphizing, bringing cute individu-
als closer into the circle of moral concern
(Sherman and Haidt, 2011). There is evi-
dence that pets, especially dogs, parasitize
our parental caretaking mechanisms (Turner,
2001). People talk to dogs in a way similar to
‘motherese’ (the sing-song way in which par-
ents talk to their infants), and motherese for
dogs has been termed ‘doggerel’ (Hirsh-
Pasek and Treiman, 1982). Dogs are also
neotenous: they retain puppylike features
throughout their lives. Furthermore, dogs who
have more paedomorphic (i.e., cute) facial
musculature are more likely to be adopted
from shelters (Waller et  al., 2013). Dogs in
childless homes are much more likely to be
groomed, given presents, and taken on vaca-
tion (H. Herzog, 2010: 79).
Dogs have been bred to retain neotenous
puppylike features and to be cuter than their
wolfy ancestors. Dogs aren’t just cuter to us;
they’re also cuter to wolves themselves! In
one study a wolf mother was given two dif-
ferent litters to foster, one with wolf puppies
and one with dog puppies:
The foster-mother wolf was… more nurturant with
the Malamute pups than with the wolf pups. She
washed them earlier and more frequently, spent
2–3 times as many hours in the den-box with them
as she did with the wolf pups, was more defensive
toward intruders, showed far more distress when
one was missing (e.g., during supplemental feed-
ings), played with them and continues to play with
them for longer periods of time. (Frank and Frank,
1982: 515)
One reason that dog breed popularity doesn’t
track health, obedience, or other desirable
qualities is the desire for the cuteness super-
stimulus. Analyzing the popularity of dog
breeds in the United States from 1926 to
1995, researchers concluded that there was
154 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
no indication ‘that breeds with more desirable
behavior, longer life, or fewer inherited
genetic disorders have been more popular than
other breeds’ (Ghirlanda et al., 2013: 4). Most
purebred dog breeds have some endemic
health problems, but brachycephalic dogs
(dogs with short snouts, like bulldogs) have
many more problems. This preference for
this brachycephalic cuteness super-stimulus,
(for example these dogs have a much rounder
more infant like head than most other dog
breeds) causes a huge amount of suffering
for animals that are otherwise treated like
family. French and English bulldogs must
usually be delivered by C-section, and suffer
many other problems like allergies, hip dys-
plasia, persistent farting, and heat sensitivity
that causes them to die more often in trans-
port (K. Herzog, 2019). Interestingly, people
with these neotenous breeds are more
attached to their pets than those with less
neotenous breeds. In a Danish sample, French
bulldog owners were more attached to their
dogs and nearly 20% more likely to say ‘I
would do anything for my dog’ compared to
owners of the much less neotenous Cairn
Terriers (Sandøe et  al., 2017). Perhaps they
just reported this because their dogs required
extraordinary attention and care or perhaps
these health problems – are one reason people
want these dogs (Serpell, 2019). Bulldogs
have been in the top five for breed popularity
for the last six years, and in the last two years,
French bulldogs have also been in the top five.
With the recent high-profile win of ‘Thor’, a
bulldog, at a national dog pageant, this trend is
likely to continue (K. Herzog, 2019).
To some extent, our special affinity for our
pet dogs translates into special moral con-
sideration for all dogs. In the United States
there was widespread outrage prompted by
China’s annual dog-meat festival (Howard,
2016). The United States has loved dogs for a
long time – and, in perhaps the greatest suc-
cess story from the animal-protection move-
ment, the number of dogs (and cats) that are
euthanized per year has plummeted, down
75% from 2011 (Parlapiano, 2019).
Why do we as humans find a huge array
of animals cute, from penguins to pandas to
pangolins? Prototypical cuteness elicitors are
cues like round and fat cheeks, large eyes,
small teeth, and playful energetic behavior.
Our own young don’t achieve peak cuteness
until they are several months old (Sherman
and Haidt, 2011). Humans produce very altri-
cial young and neonates are highly divergent
in their appearance (e.g., presence of hair,
redness of skin, facial morphology). Also,
neonates often aren’t cute, and yet this stage
of life is when our young are most vulner-
able and most in need of care. In order to take
care of our own neonates, the normal pri-
mate standards of cuteness may have become
relaxed in humans. Earlier I speculated that
slightly indiscriminate kin-detection mecha-
nisms could have made it easier for humans
to form friendships. Something similar could
have happened with cuteness and pets. The
large number of animal species we think
are cute could be because of indiscriminate
cuteness perception. ‘Cuteness promiscu-
ity’ might be a byproduct of our unusual life
history, high altriciality, and high variance
in infant appearance. This tendency to think
that many different animals are cute could be
even more pronounced in women, who are the
primary caretakers of altricial infants. Given
selection over time for dogs to elicit both
fellow-feeling (kin selection) and the cuteness
response, it makes sense that their suffering
is much more prioritized than that of other
species. Given women’s special sensitivity to
cuteness, it makes sense that we find such a
large difference in men and women when it
comes to animal morality.
SEX DIFFERENCES IN MORALITY
TOWARDS ANIMALS
In the example above in which a dolphin
fostered a melon-headed whale, she also
nursed him. Cross-species nursing is also
common cross-culturally. Women around the
 ANIMAL ETHICS AND EVOLUTIONARY PSYCHOLOGY
world have nursed baby animals like bears,
monkeys, pigs, and puppies (H. Herzog,
2019). In some of these cultures, eating an
animal nursed at the breast is considered as
taboo as eating your own child. In other cul-
tures, animals like dogs, are traded with other
groups in order to limit the discomfort of
killing animals one raised, and in other cul-
tures, animals are nursed so that they can be
later eaten (Serpell, 1996). Among the Ainu
of Japan, bear cubs are breast fed and then
ceremonially sacrificed and eaten, while the
women who suckled the bear cubs show their
ambivalence by alternately crying and laugh-
ing (Serpell, 1996: 184).
Unsurprisingly, given women’s sensitiv-
ity to cuteness and their greater nurturing
response (on average), women show stronger
moral concerns for animals than men do. For
example, 45% of women would let a foreign
tourist die before their cat or dog, compared
to 30% of men (Topolski et  al., 2013), and
33% of women would kill a person to save
1,000 dogs, compared with 23% of men
(Petrinovich et al., 1993). However, men and
women were similarly likely to say they would
save a close relative over a pet (Topolski et al.,
2013). There are many other sex differences
in moral attitudes towards animals, reflecting
differences in moral emotions such as disgust,
empathy, and the cuteness response. Women
are much more likely to be involved in ani-
mal protection and animal advocacy, much
more likely to be vegetarian, more likely to
hoard animals, and much less likely to hunt
or engage in direct animal abuse ( H. Herzog,
2007). Women are less speciesist than men
as measured through questionnaire (Caviola
et al., 2018). Women are more likely than men
to believe that animals experience complex
emotions like grief and anxiety (Walker et al.,
2014). There are also substantial sex differ-
ences in moral views on animals. In a 2015
poll, 42% of women compared to 22% of men
said that animals deserve the same rights as
people (Riffkin, 2015). In a 2011 Gallup poll
on moral issues, the largest sex differences
were on issues related to animals: ‘Majorities
155
of men, but less than half of women, consider
the use of animal fur for clothing, and medi-
cal testing on animals to be morally accept-
able’ (Saad, 2010). Women are also much
more opposed to ‘unnatural’ technologies,
including food additives, genetically modified
foods, and animal testing (Funk et al., 2015).
In particular, 62% of women oppose the use
of animals in scientific research, whereas
60% of men support it.
Animal Testing: Disgust and
Empathy
The widespread opposition to animal testing
is a good demonstration of how disgust and
empathy can create strong feelings around
animal ethics, even when they conflict with
important human interests such as biomedi-
cal advances. All drugs and interventions to
prevent human and animal suffering must
first be tested. Gary Francione (2009), who
famously advocates completely abolishing
the use and ownership of animals, has called
animal testing the only use of animals that
isn’t frivolous. Animal testing is one of the
highest-profile and most controversial uses
of animals, although it accounts for a very
small proportion of animal suffering. ‘We
kill 200 food animals for every animal used
in a scientific experiment’ (H. Herzog, 2010:
176). More than half (52%) of Americans
oppose the use of animals in scientific testing
(Strauss, 2018), and 60% oppose animal
cloning (Masci, 2017) – a technology that
could lead to significant advances in bio-
medical research and comparable gains in
human welfare. In comparison to other
­animal-related causes, there have been much
larger changes in legislation and regulation
around animal testing. For instance, the EU
has implemented bans on cosmetic animal
testing (‘Testing Cosmetics on Animals’,
2019) (arguably, this was only possible
because most cosmetic ingredients have
already been tested on animals for decades
and found safe).
156 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
In 2015, 0.2% of animals were killed in
labs but a disproportionate 0.7% of charity
money went to this cause (Bockman, 2016).
Some of the only true terrorism by animal
advocates – like death threats and property
damage – has targeted scientists and labo-
ratories conducting animal research. In par-
ticular, scientists who work with primates or
dogs have been targeted (H. Herzog, 2010).
Familiarity is one heuristic we use to infer
that something isn’t dangerous or pathogenic.
Unlike meat eating, animal experimenta-
tion’s visceral associations and unfamiliarity
combine to create an impression of ‘unnatu-
ralness’ (Holden and H. Herzog, 2019) that
is disgusting and elicits moral condemnation.
Meat
Meat is a strongly preferred food among
humans. This enjoyment of and desire for
meat is a major contributor to our moral atti-
tudes about meat, and to our self-deception
about the living conditions of animals raised
for meat. Humans almost certainly evolved
eating meat (Wrangham, 2009), and seem
motivated to eat meat specifically. For
instance, human taste buds appear to be sen-
sitive to a flavor abundant in cooked meat
called umami (Lindemann et  al., 2002). In
many places in the world there is a special
word for ‘meat hunger’ (Fiddes, 2004;
Zaraska, 2016), as distinct from other kinds
of hunger. In recent years the developing
world, either imitating rich industrialized
nations or actualizing their evolved taste
preferences for savory, high-calorie foods,
are eating more meat and other animal prod-
ucts (Kearney, 2010). Meat eating has
increased a great deal in the United States in
the last few decades, and Americans now are
eating an average of 125 kg of meat per year
(Zaraska, 2016). And in China, the most
populous country in the world, the average
person in the 1970s ate 14 kg of meat per
year, whereas in 2010 they were eating 55 kg
of meat per year (H. Herzog, 2010). Around
the world the trend is that people are eating
more meat, year after year.
However, the strong human desire for meat
has a flip side, because meat has often been a
dangerous food to eat, more likely to contain
pathogens than plant foods. Because humans
eat both plants and animals, they face an
omnivore’s dilemma: there are a large num-
ber of foods that could be eaten, but they dif-
fer both in nutritional quality and in the risk
of dangerous pathogens. Disgust is thought
to have evolved to reduce the chance of com-
ing into contact with potential pathogens,
especially those that are orally incorporated
(Tybur et  al., 2012). That’s likely why cul-
ture discourages eating certain animal foods.
Taboos are more often leveraged against
meat than against other foods (Fessler and
Navarrete, 2003). The trait of ‘disgust sensi-
tivity’ is positively linked to meat avoidance
(Fessler et  al., 2003), and disgust is often
given as an explicit reason for not eating meat
(Santos and Booth, 1996).
As I argued earlier, there is evidence that
most animals used for food are sentient, and
that human imposition of industrial-scale
suffering on sentient animals may, from the
perspective of aggregate suffering, be one of
the most pressing concerns of our genera-
tion (Singer, 1990). The scale of animal use
is staggering. According to an interview with
expert Lewis Bollard (Wiblin and Bollard,
2017), there are currently 23 billion chick-
ens being farmed (15 billion for meat and
8 billion for eggs), 6 billion mammals (like
cows, pigs, and rabbits), and over 100 billion
farmed fish. Even if we ascribe to each of
these animals just a fraction of the sentience
and moral importance ascribed to a human,
this adds up to a massive moral issue –
vastly more aggregate suffering than global
poverty or disease among humans. From the
perspective of sentient suffering, the envi-
ronmental and sustainability issues around
meat also matter. Meat is environmentally
costly to produce, requiring more water and
land per calorie, in addition to being one of
the major producers of greenhouse gases and
 ANIMAL ETHICS AND EVOLUTIONARY PSYCHOLOGY
water pollution (Steinfeld et al., 2006). Meat
feeds fewer people with the same resources.
Estimates of inefficiency vary, but the same
amount of grain produces 10 times fewer
calories through grain-fed cattle than when
eaten directly (Bittman, 2008). (However, it’s
important to consider that much of the land
that isn’t suitable for farming can still feed
grazing ruminants, like cattle, and thus can
produce food.)
In principle, boycotting animal products
could significantly reduce many of these
problems. But, even in places with abundant
food choices, vegetarianism and veganism
are rare (Pinker, 2011). Most self-described
‘vegetarians’ eat meat (Cooney, 2014), and
perhaps 90% of people who self-identify as
vegetarian aren’t behaviorally vegetarian
(H. Herzog, 2010: 195). Moreover, lapsed
vegetarians outnumber current vegetarians
(H. Herzog, 2011), and many vegetarians
avoid red meat for health reasons rather than
ethical reasons. Thus, it’s difficult to esti-
mate how many people are boycotting animal
products for ethical reasons.
All foods cause some degree of suffering –
even vegetables and fruit, because many
small wild animals, from insects to birds,
are killed during planting and harvesting. As
Norwood and Lusk (2011) glibly comment,
‘even veganism is murder’ (2011: 229).
However, animal foods differ markedly in
how much suffering they cause. Ironically,
the most popular ways that vegetarians and
semi-vegetarians reduce their consumption
of animal products may impose more net
suffering than a diet centered around beef
would, as I discuss below.
Disgust is a major driver of meat avoidance
(Fessler et al., 2003). Red meat, which retains
more cues of its animal origins, like blood, is
considered much more disgusting than fish
or chicken, which are often packaged to hide
their animal origin (H. Herzog, 2010: 190).
Health messages about meat underscore this
disgust, with recommendations to cut out
red meat and to eat chicken, eggs, and fish
instead. Self-described vegetarians, who are
157
often motivated by both disgust and health
messaging, often end up eating more chicken
than self-described meat eaters do (H. Herzog,
2010: 195). Another factor beyond disgust is
that we often feel more empathy for cows
and pigs than for fish and chickens, because
they have more humanlike and neotenous
characteristics.
How might consumption of fish, eggs, and
chicken, to the exclusion of beef and pork,
cause more animal suffering? There are two
main reasons: animal size and quality of
life. Chickens and farmed fish are smaller
animals, which means that for each animal
bred, caged, and slaughtered, we get far
fewer meals. This observation was the basis
of a tongue-in-cheek campaign from PETA
called ‘Eat the whales’ (Tomasula, 2001). In
a 100-ton blue whale, there are 70,000 chick-
ens’ worth of meat. (H. Herzog, 2010: 193).
Considering living conditions, chickens (both
egg-laying hens and broiler chickens) and
farmed fish have much worse lives than con-
ventionally produced beef cattle (Tomasik,
2018b). Conventionally produced beef cattle
spend much of their lives in pasture and the
last 100–200 days of their lives in a feedlot –
they can eat, stretch out, and associate with
others of their species. By contrast, broiler
chickens live in cramped conditions and often
have crippling leg problems. Egg-laying
hens kept in cages usually have their beaks
removed so they don’t attack or cannibalize
one another in cramped spaces. Often this
causes chronic pain or inability to feed, and
it doesn’t solve the problem of hens aggress-
ing against their cage mates. Conventional
pork production is widely considered to be
terrible for smart social animals such as pigs,
who are confined and bored, like a dog kept
in a kennel cage for months on end. For those
concerned with humane animal treatment,
a reasonable goal is that animals raised for
food should only have ‘one bad day’: the day
they go to slaughter. For detailed descriptions
of how different animals are raised for food
see, for example Norwood and Lusk (2011)
and Singer and Mason (2006).
158 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Remarkably few people have tried to
compare animal welfare across species or
to calculate how much suffering is caused
by eating different animal foods. Still, there
is some consensus on which animals have
the best and worst lives. Economists Bailey
Norwood and Jayson Lusk (2011) came to
similar conclusions as Tomasik (2018b) in
terms of the quality of life of various animals
raised for food (and also quantified the qual-
ity of life for animals kept for breeding pur-
poses) (Norwood and Lusk, 2011: 229). They
estimated that laying hens and veal calves
have the worst quality of life, and that beef
cattle and dairy cows have the best quality of
life relatively speaking. (However, Norwood
and Lusk argue that broiler chickens have a
much better quality of life on average than
most animal advocates think they do.)
How does all this add up? We can quantify a
‘suffering footprint’: an estimate of how many
days of suffering animals endure to contribute
a unit of meat to our diet. Table 7.1 is adapted
and simplified based on Tomasik’s (2018b)
calculation of how many days of suffering per
kg are caused by the demand from buying var-
ious animal foods. I have simplified the cal-
culation here by assuming these animals have
the same sentience, and by assuming that each
animal has roughly the same suffering on the
day of slaughter (the reader can input values
in the table at the Reducing Suffering blog,
https://reducing-suffering.org/how-much-
direct-suffering-is-caused-by-various-animal-
Table 7.1  Days of suffering per kilogram of food weight produced by the animal adjusted
for the badness of each day of life as estimated by animal welfare researchers
Animal product Animal lifespan (days) Kg of food per animal
lifespan
Farmed catfish 820 .39
Farmed salmon 639 2.0
Battery cage eggs 501 16
Chicken 42 1.9
Turkey 133 9.6
Pork 183 65
Beef 395 212
Milk 1,825 30,000
foods/). Here, animal lifespan is how many
days the animal lives before slaughter, on
average; kg of food per animal lifespan is how
much edible food weight is produced by the
animal; suffering per day of life is how bad
the animal’s life is based on best estimates
from animal-welfare researchers (note that
beef cattle have the best lives and battery hens
have the worst lives). The column on the right
indicates for each kg of the animal product
consumed how many days of suffering there
are adjusted for the badness of each day of
life.
Using a similar calculation, the standard
American with a typical diet of animal prod-
ucts causes 5 years, 6 months, and 5 days of
animal suffering per year (Hurford, 2014).
Regardless of specific numbers, many ‘veg-
etarians’, some adhering to the definition and
eating eggs, and many others who still eat fish
and chicken, are causing more days of animal
suffering, and more intense suffering, than
many meat eaters are. Based on the calcula-
tions from the table above, a vegetarian who
eats three eggs at a meal (around 150 g) is
causing 19.5 days of chicken suffering, com-
pared to a meat eater who eats a 1.3 kg steak
that causes around 2.4 days of cow suffer-
ing. The average vegetarian almost certainly
causes less suffering than the five years of suf-
fering created through the average American
diet. But the perception that the average self-
described ‘vegetarian’ is more moral than the
average meat eater is derived not from any
Suffering per day of Adjusted days of suffering
life (beef cows =1) caused per kg demanded
1.5 3,200
1.5 480
4 130
3 68
3 42
2.5 7.1
1 1.9
2 0.12
 ANIMAL ETHICS AND EVOLUTIONARY PSYCHOLOGY
quantitative analysis of animal suffering, but
from their claimed concern for animals, and
from the fact that they eat less meat from
mammals, who are cuter and more human-
like. Even a vegetarian who eats eggs every
day would cause more suffering than some-
one on an all-beef diet. An actual vegan, who
eats no animal products including meat, fish,
eggs, and dairy, causes the least amount of
suffering with their consumption. Giving up
fish, eggs, and chicken would reduce animal
suffering about 90% as much as a vegan who
eats no animal products at all (Cooney, 2014).
Unfortunately, there is no name for this avoid-
ing fish, eggs, and chicken ethical stance, and
thus it is not possible to signal this or reap any
benefits to social moral reputation.
The impact of better welfare animal prac-
tices on animal suffering and human moral-
ity is beyond the scope of this chapter. But
it seems that people are much more likely to
think they are buying humane animal products
than they really are (75% believe they are buy-
ing human products versus 99% of products
coming from factory farms) (Reese, 2017).
Given that billions of animals are farmed for
food across thousands of facilities, and the
food industry remains politically powerful,
it’s difficult to enforce humane standards. The
little evidence we have, such as that under-
cover animal advocacy operations seem to
always discover cruel mishandling and mis-
treatment of animals, even on farms with
‘humane standards’, doesn’t bode well.
Clean Meat
Fifty years hence, we shall escape the absurdity of
growing a whole chicken in order to eat the breast
or wing by growing these parts separately under a
suitable medium. (Winston Churchill, 1932: 26)
It’s unlikely that individual consumer choices
are going to significantly reduce the demand
for animal products. Polls show Americans
say they are very concerned about animal
welfare, but this doesn’t translate into their
159
choices as consumers. One experiment on the
‘vote/buy gap’ – the tendency for consumers
to vote for higher welfare standards but not to
buy in accordance with these ideals – showed
that 80% of consumers who chose to buy
cookies made with battery cage eggs said
that battery cage eggs should be illegal (Paul
et al., 2019). The vast majority of people in
Western societies would state that they are
morally repulsed by slavery, and yet when a
report documented that around one-third of
shrimp produced in Thailand involved slave
labor (Hodal and Lawrence, 2014), this did
not change the huge demand for shrimp, and
there is still slavery in the supply chain now
(Clark, 2019). From a historical perspective,
no movement has ever made significant gains
from endorsing individual boycotts of large-
scale industries. An analysis of the abolition-
ist movement against human slavery showed
that boycotting slave-produced goods was
not effective, and was not that widespread,
even among abolitionists (Witwicki, 2017).
One possible solution to the problem of
animal suffering caused by meat production is
in vitro meat, cultured meat, or ‘clean meat’.
Clean meat is the ‘cultivation of food grade
animal tissues in carefully controlled environ-
ments’ (McLaren, 2014: 1). Clean meat holds
the promise of replacing slaughter-based
meat production. The fast rate of technologi-
cal innovation in clean meat seems to have
overcome some of the obstacles I wrote about
several years ago (Fleischman, 2013). The
main obstacle has been price preventing clean
meat from meeting market demand. Creating
a structure for in vitro meat to grow, to keep it
at the correct temperature and inundated with
nutrients for cell division, and free from con-
tamination, made it prohibitively expensive.
The debut clean meat burger in London cre-
ated by Mark Post a few years ago cost about
$330,000 to make (McLaren, 2014). But, after
many failed predictions (Madrigal, 2013), it
seems clean meat might soon be coming to
market. A few major obstacles seem to have
been overcome since clean meat is now being
taste-tested for the public.
160 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
However, our evolved psychology may
still present obstacles to the uptake of clean
meat. Food preferences crystallize at an
early age (Birch, 1999) and people feel dis-
gust about foods that are unfamiliar to them.
To increase demand for clean meat, ethi-
cal vegetarians might at least be willing to
try it. However, in two small surveys it was
found that the majority of vegans and veg-
etarians (71%) were unwilling to try in vitro
meat (Fleischman, 2012). A larger survey of
vegetarians found a similar result, with 73%
unwilling to eat it (Dahlgreen, 2013). In my
survey (Fleischman, 2012), it seemed that the
stipulation that in vitro meat would cause no
more animal suffering than plant foods did
not change attitudes against in vitro meat,
leaving disgust as the most probable cause.
Indeed, 32% of vegans explicitly cited dis-
gust as a reason they would not want to try it.
There is some research indicating that moral
vegetarians are more disgust-sensitive over-
all (Rozin et al., 1997). However, it is disap-
pointing that this group is likely not going to
be leading the way towards clean meat.
Vegan and vegetarian attitudes are probably
not that important for the future of clean meat.
The most important thing is uptake from people
eating the most meat and populous countries
whose meat consumption is increasing, namely
China and India. There is some hopeful news
as familiarity with clean meat increases. In one
survey of American adults, the majority were
willing to try clean meat (65%) and about one-
third said they would be willing to eat clean
meat as a replacement for farmed meat (Wilks
and Phillips, 2017). Men, who tend to eat more
meat than women, also had a more positive
view of clean meat in this US sample. In a sam-
ple of over 3,000 participants from the United
States, India, and China, 93% of Chinese par-
ticipants said they were likely to purchase clean
meat, as were 86% of Indian participants and
75% of US participants (Bryant et  al., 2019).
In keeping with ideas about sex differences and
food aversions, men and those who are less
disgust-sensitive are more favorable towards
clean meat (Bryant and Barnett, 2018).
If clean meat is going to become an
­­
important solution to the myriad problems of
the global animal industry we have to learn
from history, both evolutionary and cultural.
As I mentioned above, we as humans are
more concerned about meat contamination
than other food sources. Any warning about
clean-meat contamination, or a recall, could
have pervasive long-term effects and mean that
people will continue to buy more familiar meat
from animals that suffered and died for dec-
ades to come. Branding clean meat as ‘clean
meat’ rather than in vitro meat, lab meat, cul-
tured meat, or synthetic meat was an impor-
tant first step in combating disgust sensitivity.
One major reason that genetically modified
food wasn’t an unalloyed success was because
of perceptions of unnaturalness (Mohorčich,
2018), another form of disgust response that
can be reduced by increasing consumer famili-
arity. Framing is also important; meat pro-
ducers learned long ago that mentioning the
animals themselves reduced consumer accept-
ance (Zaraska, 2016). We don’t really want to
know how the sausage is made, and less detail
about how clean meat is produced generally
improves attitudes (Bryant and Barnett, 2018).
We as humans are more concerned with what’s
delicious than what is virtuous; consumers
rarely care enough to buy or boycott any prod-
uct because of its moral ramifications (Bryant
and Barnett, 2018). That’s why making sure
that clean meat is tastier than conventional
meat can go a long way. Finally, the rise of
zoonotic diseases like Covid19 and H1N1 can
hopefully turn the tide of disgust sensitivity in
the other direction, against forms of animal
agriculture that can cause pandemics.
VIRTUE SIGNALING AND ANIMAL
WELFARE
Stop smirking. One of the most universal pieces
of advice from across cultures and eras is that we
are all hypocrites, and in our condemnation of
others’ hypocrisy we only compound our own.
(Haidt, 2006: 60)
 ANIMAL ETHICS AND EVOLUTIONARY PSYCHOLOGY
Most animals are hidden from public view or
otherwise incapable of communicating about
their suffering and cannot leverage reputa-
tional concern (Sperber and Baumard, 2012).
However, because people can advertise their
moral attitudes in so many ways now, from
vegan bumper stickers to social-media posts,
there is more widespread concern about the
suffering of animals than at any previous point
in history. Our moral attitudes do not occur in
a vacuum. Advertising our moral qualities to
others for social benefits, whether these moral
qualities are instantiated in behavior or just
‘cheap talk’, is known as virtue signaling
(Miller, 2019). When definitions of moral
behavior shift in social groups, culture can
change moral behavior to the extent that it’s
available for virtue signaling. This is one
reason that animal advocates have had so
much more success with institutional change
over individual changes (Reese, 2018). People
are willing to advertise moral ideals by sign-
ing a petition or publicly advocating that a
business change its harmful animal practices,
but are unwilling to engage in more costly and
less visible individual boycott.
Our moral identity is important to us; there
is a strong psychological motivation to present
ourselves as more moral than others (Kurzban,
2011) and to resist others’ claims of moral
superiority. This creates fraught relationships
with ‘moral minorities’ who consider them-
selves to be in the moral vanguard – including
animal advocates, vegans, and others who hold
and display a virtuous identity. Vegetarians are
widely disliked by the rest of society. In one
study, participants reported disliking vegans
and vegetarians more than atheists, asexu-
als, immigrants, or Blacks, but reported being
more willing to hire or rent to vegans and veg-
etarians than all other target groups (MacInnis
and Hodson, 2017). In this study, only drug
addicts were more disliked than vegans.
Because moral rules are considered to be
universal, meeting someone who holds dif-
ferent or more strict moral standards than you
do can be seen as an implicit indictment of
your behavior. People tend to rate themselves
161
as more moral and better than others; meat
­eaters rate vegetarians as more moral than the
average person, but rate vegetarians as less
moral than themselves (Minson and Monin,
2012). Maintaining a moral reputation is a
major reason that meat eaters rate vegetar-
ians negatively. They anticipate that vegetar-
ians are judging them and will communicate
their moral condemnation of meat eaters to
others. Meat eaters rated vegetarians more
negatively when they were first asked to
consider how much vegetarians might judge
them, and meat eaters expected vegetarians
to judge them three times more negatively
than they were actually judged (Minson and
Monin, 2012). Of course, it’s possible that
because being judgmental is widely consid-
ered immoral, vegetarians were reporting less
judgment than they actually felt.
One interesting aspect of the Minson and
Monin (2012) study was that some meat eat-
ers were first given an opportunity to say
what they thought about vegans before later
reporting how much they agreed with their
moral message. Participants in the study
described vegetarians as ‘weird’, ‘preachy’,
and ‘sadistic’. But afterward they were more
likely than other participants who did not der-
ogate vegetarians to say that they agreed with
the moral message of vegetarianism. This is
interesting from an evolutionary reputation-
management perspective. Reducing some-
one else’s reputational status relative to your
own might increase your likelihood of tak-
ing their message seriously; you don’t have
to fight as hard to make yourself look good.
Moral change often happens when we want
to socially affiliate with others, and negative
impressions of activists – from animal advo-
cates to social-justice advocates – undermines
the cause (Bashir et  al., 2013). Importantly,
for any moral advocate, they must remember
that others are going to have strong incentives
to derogate them and will scrutinize them for
moral inconsistency (Monin, 2007). Anyone
advocating a major change in moral priori-
ties must remember that people have spent
years honing their virtue-signaling strategies,
162 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
and will not take kindly to someone arguing
that they have really been hugely less virtu-
ous than they thought. The evolutionary psy-
chological challenge for animal advocacy is
to nudge people to show more concern for
animal suffering, without feeling like their
whole virtue-signaling identity has to be jet-
tisoned and rebuilt from scratch.
CONCLUSION
Evolutionary explanations are often maligned
because they are said to excuse or normalize
violence. To say animal cruelty and inflicting
animal suffering is normal and natural is not to
minimize the suffering of animal victims either
as the result of any individual’s sadism or the
large-scale production of animal products. To
say that our nurturing instincts predispose us to
be kinder to animals that demonstrate kinship
cues or that elicit the cuteness response is not
to say that these responses are moral. To say
that we are more disgusted by meat that looks
more like the animal it came from than meat
that looks more abstract is not to say it is more
moral to eat meat packaged in cellophane. And
to say that we virtue signal about our moral
behavior is not to say that moral behavior isn’t
important or that cynical motivations render
moral behavior immoral. When we take our
moral intuitions as moral rules we project and
institutionalize our evolved moral blind spots
into the world, often making it worse for
others. Advocacy requires understanding. If
animal suffering is an ethical issue, we have to
be realistic about our incentives to signal, our
functional emotional responses and what com-
prises our evolved moral psychology towards
animals.
ACKNOWLEDGMENTS
For helpful feedback, thanks to Hal Herzog,
Geoffrey Miller, and Todd Shackelford.
REFERENCES
Ahimsa in Jainism. (2019). In Wikipedia. https://
en.wikipedia.org/w/index.php?title=Ahimsa_
in_Jainism&oldid=929905870 (Accessed
4 January 2020)
Amiot, C. E., & Bastian, B. (2015). Toward a
psychology of human–animal relations.
Psychological Bulletin, 141(1), 6–47.
Appel, M., & Elwood, R. W. (2009). Motivational
trade-offs and potential pain experience in
hermit crabs. Applied Animal Behaviour
Science, 119(1–2), 120–124.
Arluke, A. (2002). Animal abuse as dirty play.
Symbolic Interaction, 25(4), 405–430.
Awad, E., Dsouza, S., Kim, R., Schulz, J., Hen-
rich, J., Shariff, A., Bonnefon, J.-F., & Rahwan,
I. (2018). The moral machine experiment.
Nature, 563(7729), 59.
Bandura, A. (1999). Moral disengagement in
the perpetration of inhumanities. Personality
and Social Psychology Review, 3(3),
193–209.
Bashir, N. Y., Lockwood, P., Chasteen, A. L.,
Nadolny, D., & Noyes, I. (2013). The ironic
impact of activists: Negative stereotypes
reduce social change influence. European
Journal of Social Psychology, 43(7),
614–626.
Birch, L. L. (1999). Development of food prefer-
ences. Annual Review of Nutrition, 19(1),
41–62.
Bittman, M. (2008, January 27). Rethinking the
Meat-Guzzler. The New York Times. www.
nytimes.com/2008/01/27/weekinreview/
27bittman.html (Accessed 4 January 2020)
Bloom, P. (2017). Against empathy: The case
for rational compassion. Random House.
Bockman, J. (2016, November 1). Why Farmed
Animals? Animal Charity Evaluators. https://
animalcharityevaluators.org/donation-
advice/why-farmed-animals/ (Accessed 4
January 2020)
Bostrom, N. (2016). Superintelligence: Paths,
dangers, strategies. UK, Oxford: Oxford Uni-
versity Press.
Bradshaw, J. W., & Paul, E. S. (2010). Could
empathy for animals have been an adaptation
in the evolution of Homo sapiens. Animal
Welfare, 19(1), 107–112.
Braithwaite, V. (2010). Do fish feel pain? OUP
Oxford. http://books.google.co.uk/books?
 ANIMAL ETHICS AND EVOLUTIONARY PSYCHOLOGY
hl=en&lr=&id=aMvonPqzu_cC&oi=fnd&
pg=PT2&dq=do+fish+feel+pain&ots=tjbXOJy
C5x&sig=oU0xqgrjKN3RQfRll99esQfMxEI
(Accessed 4 January 2020)
Bryant, C., & Barnett, J. (2018). Consumer
acceptance of cultured meat: A systematic
review. Meat Science, 143, 8–17.
Bryant, C. J., Szejda, K., Deshpande, V., Parekh,
N., & Tse, B. (2019). A survey of consumer
perceptions of plant-based and clean meat
in the USA, India, and China. Frontiers in
Sustainable Food Systems, 3, 11.
Carzon, P., Delfour, F., Dudzinski, K., Oremus,
M., & Clua, É. (2019). Cross-genus adoptions
in delphinids: One example with taxonomic
discussion. Ethology, 125(9), 669–676.
https://doi.org/10.1111/eth.12916
Caviola, L. (2019). How we value animals: The
psychology of speciesism [University of Oxford].
https://ora.ox.ac.uk/objects/uuid:9000a8be-
b6dc-4c63-88e8-974b9daaa83a (Accessed 4
January 2020)
Caviola, L., Everett, J. A., & Faber, N. S. (2019).
The moral standing of animals: Towards a
psychology of speciesism. Journal of person-
ality and social psychology, 116(6), 1011.
Churchill, W. (1932). Fifty years hence.
Thoughts and adventures, 24–27. Thornton
Butterworth London.
Clark, M. (2019, October 15). What Are We
Supposed to Think About Shrimp? The New
York Times. www.nytimes.com/2019/10/15/
dining/shrimp-sourcing-united-states.html
(Accessed 4 January 2020)
Cohen, R. (2009, July 15). Nesting Blues. The
New York Times. www.nytimes.com/2009/
07/19/magazine/19FOB-ethicist-t.html
(Accessed 4 January 2020)
Cooney, N. (2014). Veganomics: The surprising
science on vegetarians, from the breakfast
table to the bedroom. Herndon Virginia
USA: Lantern Books.
Dahlgreen, W. (2013, August 5). No British
Demand for Fake Meat. YouGov. https://
yougov.co.uk/news/2013/08/05/no-demand-
fake-meat/ (Accessed 4 January 2020)
Diamond, J. (1993). New Guineans and their
natural world. The biophilia hypothesis,
251–271. Island Press Washington DC.
Elwood, R. W. (2011). Pain and suffering
in invertebrates? Ilar Journal, 52(2),
175–184.
163
Fessler, D. M. T., Arguello, A. P., Mekdara, J. M., &
Macias, R. (2003). Disgust sensitivity and
meat consumption: A test of an emotivist
account of moral vegetarianism. Appetite,
41(1), 31–41.
Fessler, D. M. T., & Navarrete, C. D. (2003).
Meat is good to taboo: Dietary proscriptions
as a product of the interaction of psychologi-
cal mechanisms and social processes. Journal
of Cognition and Culture, 3(1), 1–40. https://
doi.org/10.1163/156853703321598563
Fiddes, N. (2004). Meat: A natural symbol.
Abingdon Oxfordshire: Routledge.
Fitzgerald, A. J., Kalof, L., & Dietz, T. (2009).
Slaughterhouses and increased crime rates:
An empirical analysis of the spillover from ‘The
Jungle’ into the surrounding community.
Organization & Environment, 22(2), 158–184.
Fleischman, D. S. (2012, May 16). Lab Meat:
Survey Results. The Vegan Option. http://
theveganoption.org/2012/05/16/lab-meat-
survey-results/ (Accessed 4 January 2020)
Fleischman, D. S. (2013, August 5). Will Clean
Meat Become Cruelty Free? Sentientist.
https://dianaverse.com/2020/04/29/will-
clean-meat-become-cruelty-free-repost/
(Accessed 4 January 2020)
Fleischman, D. S. (2018, May 17). Universal
Morality Is Obscured by Evolved Morality.
The Evolution Institute. https://evolution-
institute.org/universal-morality-is-obscured-
by-evolved-morality/ (Accessed 1 May
2020)
Flynn, C. P. (2011). Examining the links between
animal abuse and human violence. Crime,
Law and Social Change, 55(5), 453–468.
https://doi.org/10.1007/s10611-011-9297-2
Foster, K. R., Wenseleers, T., & Ratnieks, F. L.
(2006). Kin selection is the key to altruism.
Trends in Ecology & Evolution, 21(2), 57–60.
Fox Hunting. (2019). In Wikipedia. https://
en.wikipedia.org/w/index.php?title=Fox_
hunting&oldid=933138160 (Accessed
2 January 2020)
Francione, G. L. (2009). Animals as persons.
New York: Columbia University Press.
Francione, G. L. (2015). Animal rights: The
abolitionist approach. Louisville, Kentucky:
Exempla Press.
Frank, H., & Frank, M. G. (1982). On the effects
of domestication on canine social develop-
ment and behavior. Applied Animal Ethol-
164 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
ogy, 8(6), 507–525. https://doi.org/
10.1016/0304-3762(82)90215-2
Beauchamp, T. L., & Frey, R. G. (Eds.). (2011).
The Oxford handbook of animal ethics.
Oxford University Press. Oxford, UK.
172–198.
Funk, C., Rainie, L., Smith, A., Olmstead, K.,
Duggan, M., & Page, D. (2015). Attitudes
and Beliefs on Science and Technology Topics.
Pew Research Center. www.pewresearch.
org/science/2015/01/29/chapter-3-attitudes-
and-beliefs-on-science-and-technology-
topics/ (Accessed 4 January 2020)
Ghirlanda, S., Acerbi, A., Herzog, H., & Serpell,
J. (2013). Fashion vs. function in cultural
evolution: The case of dog breed popularity.
PLoS One, 8(9), e74770.
Greene, J. D. (2013). Moral tribes: Emotion,
reason, and the gap between us and them.
London, UK: Penguin.
Gullone, E. (2014). An evaluative review of
theories related to animal cruelty. Journal of
Animal Ethics, 4(1), 37–57.
Haidt, J. (2006). The happiness hypothesis:
Finding modern truth in ancient wisdom.
New York, USA Basic Books.
Haidt, J. (2012). The righteous mind: Why good
people are divided by religion and politics.
New York: Pantheon Books. http://media.
matthewsbooks.com.s3.amazonaws.com/
documents/tocwork/030/9780307377906.
pdf (Accessed 4 January 2020)
Herzog, H. (2002). Darwinism and the study of
human–animal interactions. Society &
Animals, 10(4), 361–367. https://doi.org/
10.1163/156853002320936818
Herzog, H. (2007). Gender differences in
human–animal interactions: A review.
Anthrozoös, 20(1), 7–21.
Herzog, H. (2010). Some we love, some we
hate, some we eat: Why it’s so hard to think
straight about animals (Reprint edition).
HarperCollins e-books.
Herzog, H. (2011, June 20). Why Do Most Veg-
etarians Go Back to Eating Meat? Animals
and Us. www.psychologytoday.com/blog/
animals-and-us/201106/why-do-most-­
vegetarians-go-back-eating-meat (Accessed
4 January 2020)
Herzog, H. (2014). Biology, culture, and the
origins of pet-keeping. Animal Behavior and
Cognition, 1(3), 296–308.
Herzog, H. (2019, August 6). Did Breast-
Feeding Play a Role in the Evolution of Pets?
Psychology Today. www.psychologytoday.
com/blog/animals-and-us/201908/did-
breast-feeding-play-role-in-the-evolution-
pets (Accessed 4 January 2020)
Herzog, H., & Arluke, A. (2006). Human–
animal connections: Recent findings on the
anthrozoology of cruelty. Behavioral and
Brain Sciences, 29(3), 230–231.
Herzog, K. (2019, December 2). Genetic Freak
Wins National Dog Show. The Stranger. www.
thestranger.com/slog/2019/12/02/42155095/
genetic-freak-wins-national-dog-show
(Accessed 4 January 2020)
Hirsh-Pasek, K., & Treiman, R. (1982). Dog-
gerel: Motherese in a new context. Journal
of Child Language, 9(1), 229–237.
Hodal, K., & Lawrence, C. K. F. (2014, June 10).
Revealed: Asian Slave Labour Producing
Prawns for Supermarkets in US, UK. The
Guardian. www.theguardian.com/global-
development/2014/jun/10/supermarket-
prawns-thailand-produced-slave-labour
(Accessed 4 January 2020)
Holden, C. J., & Herzog, H. (2019). Featherless
chickens and puppies that glow in the dark.
In K. Dhont & G. Hodson (Eds.), Why we love
and exploit animals: Bridging insights from
academia and advocacy. Abingdon, Oxford-
shire: Routledge. 137–153.
Howard, B. C. (2016, June 21). Dog Meat
Festival Opens Amid Outrage. National
Geographic News. www.nationalgeographic.
com/news/2016/06/china-yulin-dog-meat-
festival-controversy/ (Accessed 4 January
2020)
Hurford, P. (2014, May 31). How Much Suffer-
ing Is in the Standard American Diet? Every-
day Utilitarian. www.everydayutilitarian.com/
essays/how-much-suffering-is-in-the-stand-
ard-american-diet/ (Accessed 4 January 2020)
Izar, P., Verderane, M. P., Visalberghi, E., Ottoni,
E. B., Gomes De Oliveira, M., Shirley, J., &
Fragaszy, D. (2006). Cross-genus adoption
of a marmoset (Callithrix jacchus) by wild
capuchin monkeys (Cebus libidinosus): Case
report. American Journal of Primatology:
Official Journal of the American Society of
Primatologists, 68(7), 692–700.
Joy, M. (2003). Psychic numbing and meat
consumption: The psychology of carnism.
 ANIMAL ETHICS AND EVOLUTIONARY PSYCHOLOGY
(Unpublished dissertation). Saybrook Gradu-
ate School, NY.
Joy, M. (2011). Why we love dogs, eat pigs,
and wear cows: An introduction to car-
nism. Newburyport, Massachusetts: Conari
Press.
Kahane, G. (2009). Pain, dislike and experi-
ence. Utilitas, 21(3), 327–336.
Kaminski, J., Waller, B. M., Diogo, R., Hart-
stone-Rose, A., & Burrows, A. M. (2019).
Evolution of facial muscle anatomy in dogs.
Proceedings of the National Academy of
Sciences, 116(29), 14677–14681.
Kasperbauer, T. J. (2015). Rejecting empathy
for animal ethics. Ethical Theory and Moral
Practice, 18(4), 817–833. https://doi.
org/10.1007/s10677-014-9557-1
Kearney, J. (2010). Food consumption trends
and drivers. Philosophical Transactions of the
Royal Society of London B: Biological
Sciences, 365(1554), 2793–2807. https://
doi.org/10.1098/rstb.2010.0149
Krupp, D. B., DeBruine, L. M., & Jones, B. C.
(2011). Cooperation and conflict in the light of
kin recognition systems. The Oxford Handbook
of Evolutionary Family Psychology. Oxford. UK:
Oxford University Press. 345–362.
Kurzban, R. (2011). Why everyone (else) is a
hypocrite: Evolution and the modular mind.
Princeton: Princeton University Press.
Kurzban, R., Tooby, J., & Cosmides, L. (2001).
Can race be erased? Coalitional computation
and social categorization. Proceedings of the
National Academy of Sciences, 98(26),
15387–15392.
Lew-Levy, S., Reckin, R., Lavi, N., Cristóbal-
Azkarate, J., & Ellis-Davies, K. (2017). How
do hunter-gatherer children learn subsist-
ence skills? Human Nature, 28(4), 367–394.
https://doi.org/10.1007/s12110-017-9302-2
Lieberman, D., Tooby, J., & Cosmides, L. (2007).
The architecture of human kin detection.
Nature, 445(7129), 727–731.
Lindemann, B., Ogiwara, Y., & Ninomiya, Y.
(2002). The discovery of umami. Chemical
Senses, 27(9), 843–844.
Machan, T. R. (2004). Putting humans first:
Why we are nature’s favorite. Lanham Mary-
land USA: Rowman & Littlefield Publishers.
MacInnis, C. C., & Hodson, G. (2017). It ain’t
easy eating greens: Evidence of bias toward
vegetarians and vegans from both source
165
and target. Group Processes & Intergroup
Relations, 20(6), 721–744.
Madrigal, A. C. (2013, August 6). Chart: When
Will We Eat Hamburgers Grown in Test
Tubes? The Atlantic. www.theatlantic.com/
technology/archive/2013/08/chart-when-
will-we-eat-hamburgers-grown-in-test-
tubes/278405/ (Accessed 4 January 2020)
Marino, L. (2019). I am not an animal. Animal
Sentience, 3(23), 40.
Masci, D. (2017, February 22). 20 Years after
Dolly the Sheep’s Debut, Americans Remain
Skeptical of Cloning. Pew Research Center.
www.pewresearch.org/fact-tank/2017/
02/22/20-years-after-dolly-the-sheeps-
debut-americans-remain-skeptical-of-­
cloning/ (Accessed 4 January 2020)
McLaren, M. E. (2014). Cultured Meat: A
Beneficial, Crucial, and Inevitable Nutrition
Technology. Seton Hall University. Law
School Student Scholarship. 527. http://
scholarship.shu.edu/cgi/viewcontent.cgi?
article=1527&context=student_scholarship
(Accessed 4 January 2020)
Miller, G. (2019). Virtue signaling: Essays on
Darwinian politics & free speech. Albuquerque,
New Mexico: Cambrian Moon.
Minson, J. A., & Monin, B. (2012). Do-gooder
derogation: Disparaging morally motivated
minorities to defuse anticipated reproach.
Social Psychological and Personality Science,
3(2), 200–207.
Mohorčich, J. (2018). What Can the Adoption
of GM Foods Teach Us about the Adoption
of Other Food Technologies? Sentience Insti-
tute. https://sentienceinstitute.org/gm-foods
(Accessed 4 January 2020)
Monin, B. (2007). Holier than me? Threatening
social comparison in the moral domain.
Revue Internationale de Psychologie Sociale,
20(1), 53–68.
Mood, A., & Brooke, P. (2010). Estimating the
number of fish caught in global fishing each
year. http://fishcount.org.uk/published/std/
fishcountstudy.pdf (Accessed 4 January 2020)
Moore, D. C. (2003). Public Lukewarm on
Animal Rights. Gallup. https://news.gallup.
com/poll/8461/Public-Lukewarm-Animal-
Rights.aspx (Accessed 4 January 2020)
Nakajima, S., Yamamoto, M., & Yoshimoto, N.
(2009). Dogs look like their owners:
Replications with racially homogenous owner
166 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
portraits. Anthrozoös, 22(2), 173–181.
https://doi.org/10.2752/175303709X434194
Norwood, F. B., & Lusk, J. (2011). Compassion,
by the pound: The economics of farm animal
welfare. Oxford UK: Oxford University Press.
Pachirat, T. (2011). Every twelve seconds:
Industrialized slaughter and the politics of
sight. Yale University Press. New Haven:
Connecticut.
Park, J. H., & Ackerman, J. M. (2011). Passion
and compassion: Psychology of kin relations
within and beyond the family. In Salmon, C.,
& Shackelford, T. K. (Eds.) (2011). The Oxford
handbook of evolutionary family psychology.
Oxford, UK: Oxford University Press.
Park, J. H., & Schaller, M. (2005). Does attitude
similarity serve as a heuristic cue for kinship?
Evidence of an implicit cognitive association.
Evolution and Human Behavior, 26(2), 158–
170. https://doi.org/10.1016/j.evolhumbehav.
2004.08.013
Parlapiano, A. (2019, September 3). Why
Euthanasia Rates at Animal Shelters Have
Plummeted. The New York Times. www.
nytimes.com/2019/09/03/upshot/why-eutha
nasia-rates-at-animal-shelters-have-plum-
meted.html (Accessed 4 January 2020)
Patterson-Kane, E. G., & Piper, H. (2009).
Animal abuse as a sentinel for human vio-
lence: A critique. Journal of Social Issues,
65(3), 589–614. https://doi.org/10.1111/
j.1540-4560.2009.01615.x
Paul, A. S., Lusk, J. L., Norwood, F. B., & Tonsor,
G. T. (2019). An experiment on the vote-buy
gap with application to cage-free eggs. Jour-
nal of Behavioral and Experimental Econom-
ics, 79, 102–109. https://doi.org/10.1016/j.
socec.2019.02.005
PeTAUK (2019). These 27 Countries Have
Banned Wild-Animal Circuses! [Blog Post].
https://www.peta.org.uk/blog/these-26-
countries-that-have-banned-wild-animal-
circuses-are-making-england-look-really-
bad/ (Accessed 6 July, 2020)
Petrinovich, L., O’Neill, P., & Jorgensen, M.
(1993). An empirical study of moral intuitions:
Toward an evolutionary ethics. Journal of Per-
sonality and Social Psychology, 64(3), 467.
Pets Really Are Members of the Family. (2011,
June 10). The Harris Poll. https://theharrispoll.
com/americans-have-always-had-interesting-
relationships-with-their-pets-whether-that-
pet-is-a-cat-dog-parakeet-or-something-
else-the-pet-industry-is-thriving-and-for-
good-reason-more-than-three-in-f/ (Accessed
4 January 2020)
Pinker, S. (2011). The better angels of our nature:
The decline of violence in history and its
causes. London, UK: Penguin. https://books.
google.co.uk/books?hl=en&lr=&id=c3cWa-
GnsfMC&oi=fnd&pg=PT11&dq=better+angel
s+of+our+nature&ots=aKhKguUK8a&sig=iKN
0Khly8JAx42B3LcJu30My-m8
Rauch, A., & Sharp, J. S. (2005). Ohioans’ atti-
tudes about animal welfare. Social Responsi-
bility Initiative. Department of Human and
Community Resource Development. The
Ohio State University.
Reese, J. (2017). Animal Farming Attitudes
Survey 2017. Sentience Institute. www.sen-
tienceinstitute.org/animal-farming-attitudes-
survey-2017 (Accessed 4 January 2020)
Reese, J. (2018, June 25). 3 Big Changes We
Need in the Farmed Animal Movement. Sen-
tience Institute. www.sentienceinstitute.org/
blog/three-big-changes (Accessed 4 January
2020)
Reese, J. (2019). US Factory Farming Estimates.
Sentience Institute. www.sentienceinstitute.
org/us-factory-farming-estimates (Accessed
4 January 2020)
Regan, T. (2004). The case for animal rights.
Oakland, California: University of California
Press.
Riffkin, R. (2015, May 18). In U.S., More Say
Animals Should Have Same Rights as People.
Gallup. https://news.gallup.com/poll/183275/
say-animals-rights-people.aspx (Accessed 4
January 2020)
Roy, M. M., & Nicholas, J. S. C. (2004). Do dogs
resemble their owners? Psychological Science,
15(5), 361–363. https://doi.org/10.1111/
j.0956-7976.2004.00684.x
Rozin, P., Markwith, M., & Stoess, C. (1997).
Moralization and becoming a vegetarian: The
transformation of preferences into values
and the recruitment of disgust. Psychological
Science, 8(2), 67.
Ryder, R. D. (1991). Souls and sentientism.
Between the Species: An Online Journal for
the Study of Philosophy and Animals, 7(1) 1–5.
https://doi.org/10.15368/bts.1991v7n1.1
Saad, L. (2010, May 26). Four Moral Issues
Sharply Divide Americans. Gallup. https://
 ANIMAL ETHICS AND EVOLUTIONARY PSYCHOLOGY
news.gallup.com/poll/137357/Four-Moral-Issues-
Sharply-Divide-Americans.aspx (Accessed
4 January 2020)
Sandøe, P., Kondrup, S. V., Bennett, P. C., Fork-
man, B., Meyer, I., Proschowsky, H. F., Serpell, J.
A., & Lund, T. B. (2017). Why do people buy
dogs with potential welfare problems related to
extreme conformation and inherited disease? A
representative study of Danish owners of four
small dog breeds. PLoS One, 12(2), e0172091.
https://doi.org/10.1371/journal.pone.0172091
Santos, M. L. S., & Booth, D. A. (1996).
Influences on meat avoidance among British
students. Appetite, 27(3), 197–205.
Scott-Phillips, T. C., Dickins, T. E., & West, S. A.
(2011). Evolutionary theory and the ultimate–
proximate distinction in the human behavioral
sciences. Perspectives on Psychological
Science, 6(1), 38–47. https://doi.org/10.1177/
1745691610393528
Sentientism. (2019). In Wikipedia. https://
en.wikipedia.org/w/index.php?title=Sentienti
sm&oldid=926119747 (Accessed 4 January
2020)
Serpell, J. (1996). In the company of animals: A
study of human-animal relationships. Cam-
bridge, UK: Cambridge University Press.
Serpell, J. (2019). How happy is your pet? The
problem of subjectivity in the assessment of
companion animal welfare. Animal Welfare,
28(1), 57–66. https://doi.org/10.7120/
09627286.28.1.057
Sherman, G. D., & Haidt, J. (2011). Cuteness
and disgust: The humanizing and dehuman-
izing effects of emotion. Emotion Review,
3(3), 245–251. https://doi.org/10.1177/
1754073911402396
Shostak, M., & Nisa. (2004). Nisa: The life and
words of a !Kung woman. Cambridge: Har-
vard University Press.
Simler, K., & Hanson, R. (2017). The elephant in
the brain: Hidden motives in everyday life.
Cambridge, UK: Cambridge University Press.
Singer, P. (1995). Animal liberation. Random
House.
Singer, P. (2011). The expanding circle: Ethics,
evolution, and moral progress. Princeton:
Princeton University Press. http://books.
google.co.uk/books?hl=en&lr=&id=Yve7lgvt
LcsC&oi=fnd&pg=PP2&dq=the+expanding+
c irc le & ot s=5L e S l Yq g O j & s i g = 2 U l L 4 8 I p
LqC-MKX7_s10Q0PjvnA
167
Singer, P., & Mason, J. (2006). The ethics of
what we eat. Melbourne: Text Publishing.
Sneddon, L. U., Braithwaite, V. A., & Gentle, M.
J. (2003). Novel object test: Examining
nociception and fear in the rainbow trout.
The Journal of Pain, 4(8), 431–440.
Sperber, D., & Baumard, N. (2012). Moral
reputation: An evolutionary and cognitive
perspective. Mind & Language, 27(5),
495–518.
Steinfeld, H., Gerber, P., Wassenaar, T. D.,
Castel, V., Rosales, M., Rosales, M., & de
Haan, C. (2006). Livestock’s long shadow:
environmental issues and options. Food &
Agriculture Org. https://books.google.co.uk/
books?id=1B9LQQkm_qMC&lpg=PP1&ots=
LO1XiY7OlI&dq=livestock’s%20long%20sha
dow&lr&pg=PP1#v=onepage&q&f=false
Strauss, M. (2018, August 16). Americans
Divided over Use of Animals in Scientific
Research. Pew Research Center. www.
pewresearch.org/fact-tank/2018/08/16/
a m e r i c a n s - a re - d i v i d e d - o v e r- t h e - u s e -
of-animals-in-scientific-research/ (Accessed
4 January 2020)
Testing Cosmetics on Animals. (2019). In
Wikipedia. https://en.wikipedia.org/w/index.
php?title=Testing_cosmetics_on_animals&
oldid=929257214 (Accessed 4 January
2020)
Tomasik, B. (2017, August 17). Suffering in Ani-
mals vs. Humans. Reducing Suffering. https://
reducing-suffering.org/suffering-in-animals-
vs-humans/ (Accessed 4 January 2020)
Tomasik, B. (2018a, September 20). Bacteria,
Plants, and Graded Sentience. Reducing
Suffering. https://reducing-suffering.org/
bacteria-plants-and-graded-sentience/
(Accessed 4 January 2020)
Tomasik, B. (2018b, July 14). How Much Direct
Suffering Is Caused by Various Animal
Foods? Reducing Suffering. https://reducing-
suffering.org/how-much-direct-suffering-is-
caused-by-various-animal-foods/ (Accessed 4
January 2020)
Tomasik, B. (2019, July 21). Is Brain Size
Morally Relevant? Reducing Suffering.
https://reducing-suffering.org/is-brain-size-
morally-relevant/ (Accessed 4 January 2020)
Tomasula, D. (2001, July 26). PETA Says Eat a
Whale, Save a Pig. DMNews.Com. www.
dmnews.com/marketing-channels/direct-mail/
168 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
news/13088690/peta-says-eat-a-whale-
save-a-pig (Accessed 4 January 2020)
Tooby, J., & Cosmides, L. (1989). Evolutionary
psychology and the generation of culture,
part I: Theoretical considerations. Ethology
and Sociobiology, 10(1–3), 29–49.
Topolski, R., Weaver, J. N., Martin, Z., & McCoy,
J. (2013). Choosing between the emotional
dog and the rational pal: A moral dilemma
with a tail. Anthrozoös, 26(2), 253–263.
https://doi.org/10.2752/175303713X13636
846944321
Turner, W. G. (2001). Our new children: The sur-
rogate role of companion animals in women’s
lives. The Qualitative Report, 6(1), 1–10.
Tybur, J. M., Lieberman, D., Kurzban, R., & DeScioli,
P. (2013). Disgust: Evolved function and
structure. Psychological Review, 120(1), 65.
Urquiza-Haas, E. G., & Kotrschal, K. (2015).
The mind behind anthropomorphic thinking:
Attribution of mental states to other species.
Animal Behaviour, 109, 167–176. https://
doi.org/10.1016/j.anbehav.2015.08.011
Vollum, S., Longmire, D., & Buffington-Vollum,
J. (2004). Moral disengagement and attitudes
about violence toward animals. Society &
Animals, 12(3), 209–235.
Walker, J. K., McGrath, N., Nilsson, D. L.,
Waran, N. K., & Phillips, C. J. (2014). The role
of gender in public perception of whether
animals can experience grief and other
emotions. Anthrozoös, 27(2), 251–266.
Waller, B. M., Peirce, K., Caeiro, C. C., Scheider,
L., Burrows, A. M., McCune, S., & Kaminski, J.
(2013). Paedomorphic facial expressions give
dogs a selective advantage. PLoS One, 8(12),
e82686.
Wiblin, R., & Bollard, L. (2017, September 27).
Why we must end factory farming as soon as
possible – and how to do it [Interview].
https://80000hours.org/podcast/episodes/
lewis-bollard-end-factory-farming/ (Accessed
4 January 2020)
Wilks, M., & Phillips, C. J. (2017). Attitudes to
in vitro meat: A survey of potential consumers
in the United States. PloS One, 12(2). https://
jour nals.plos.org/plosone/article/file?
type=printable&id=10.1371/journal.pone.
0171904 (Accessed 4 January 2020)
Wilson, E. O. (1984). Biophilia: The human
bond with other species. Cambridge: Harvard
University Press.
Witwicki, K. (2017). Social Movement Lessons
From the British Antislavery Movement:
Focused on Applications to the Movement
Against Animal Farming. Sentience Institute.
www.sentienceinstitute.org/british-antislavery
(Accessed 4 January 2020)
Wrangham, R. (2009). Catching fire: How
cooking made us human. New York: Basic
Books. https://books.google.co.uk/books?
hl=en&lr=&id=ebEOupKz-rMC&oi=fnd&pg=P
R5&dq=humans+evolved+eating+meat
&ots=sVRt8Y8NLS&sig=2a3UdxIPEq_Crn
BKY0XK8O5hGrI
Zaraska, M. (2016). Meathooked: The history
and science of our 2.5-million-year obsession
with meat. New York: Basic Books.
 PART 2

Applications to Law and Order

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Evolutionary Psychology and
Political Institutions
Michael Latner and Elissa Feld
Biology and political science have a long,
symbiotic relationship in their shared focus
on conflict and cooperation between organ-
isms. Many of the major puzzles in biology
and political science, from the emergence of
cooperation, and the threat of parasitism and
selfishness, to the dynamics of collective
decision making and the evolution of moral-
ity, are interwoven in early, pre-Darwinian
works. As these areas of study developed into
separate professional disciplines in the early
20th century, they would part ways with con-
siderable tension before returning to such
fundamentals with the advent of modern
evolutionary psychology. In this chapter, we
survey the literature surrounding three major
areas where the study of political institutions
and evolutionary psychology intersect: the
evolution of cooperation at the micro-level;
the emergence of complex political systems;
and democracy as a major evolutionary tran-
sition in nature.
8
THE EVOLUTION OF COOPERATION
The Scottish Enlightenment, especially the
work of David Hume, had a significant influ-
ence on James Madison’s theory of republican
government (Mclean, 2005). Hume’s discourses
on competition and selective retention would
figure prominently in Madison’s constitutional-
design principles, especially his commitment to
design institutions responsive enough to ‘guard
against the cabals of a few’ but not subject to so
much ‘mutability’ and ‘incessant changes’ that
‘no man who knows what the law is today can
guess what it will be tomorrow’ (Palmer, 2002:
109). Hume’s concern about the degenerative
effects of extreme elements in political compe-
tition, as well as the capacity for such diverse
interests to be ‘serviceable’ to the public inter-
est, would be fully developed in Madison’s
writings on faction, the modern study of politi-
cal institutions, and constitutional design
(Madison et al., [1788]2008).
172 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Charles Darwin studied both Hume and
Madison’s contemporary, the Reverend
Thomas Malthus, whose Essay on the Principle
of Population (Malthus, [1798]2013) began to
quantify the relationship between competition
for resources and human-population growth
(Hamilton et al., [1788]1998). Madison him-
self understood that the world would be ‘much
indebted to’ Malthus for his work, though a
decade before Malthus’ Principle was pub-
lished, Madison had already been reflecting
with Thomas Jefferson on the degenerative
social effects of concentrated resources among
the ‘idle rich’ and the potential threat of over-
population (Madison, 1786; McCoy, 1980).
The threat of parasitism on social prosper-
ity and development was on the mind of
these institutional reformers. The Principle
of Population would also lead Darwin to the
insight that with selection pressures under
conditions of scarcity ‘favourable variations
would tend to be preserved, and unfavourable
ones to be destroyed. The results of this would
be the formation of a new species. Here, then
I had at last got a theory by which to work’
(Darwin, [1838]2011).
Hume may have also been influential in
the way that Madison and Darwin thought
about the emergence of cooperative behav-
ior within the context of group competition,
an insight neglected by Malthus. Consider
Hume’s account of the logic of reciprocity
in his Treatise on Human Nature (Binmore,
1994: 261):
I learn to do service to another, without bearing
him any real kindness, because I foresee, that he
will return my service in expectation of another of
the same kind, and in order to maintain the same
correspondence of good offices with me and
others. And accordingly, after I have serv’d him …
he is induced to do his part, as foreseeing the
consequences of his refusal.
Darwin certainly recognized the dynamics of
reciprocity at play in cooperative behavior
and the dilemma of moral development.
Darwin viewed morality as the highest princi-
ple of human development, but he also saw
that in group competition the ‘bravest men’
who came to the front at war and ‘freely
risked their lives for others, would on an aver-
age perish in larger number than other men’,
producing fewer offspring (Darwin, 1876:
130). Immoral (selfish) actors would, on aver-
age, out-produce their braver comrades, even-
tually driving the trait of bravery out of the
population. However, competition between
groups could change the selective dynamics:
It must not be forgotten that although a high
standard of morality gives but a slight or no advan-
tage to each individual man and his children over
the other men of the same tribe, yet that an
advancement in the standard of morality and an
increase in the number of well-endowed men will
certainly give an immense advantage to one tribe
over another. (Darwin, 1876: 132)
This is the basis of the cooperation dilemma,
and the question of how cooperative or ‘good’
traits could gain a selective advantage through
social decisions was a central feature in early
social choice theory. On that topic another
contemporary of Madison, the Marquis de
Condorcet, also played a major role. He for-
mally showed that voters seeking the best col-
lective decision for their group could rely on
majority rule to most likely yield a ‘correct’
decision, if the probability of a voter being
right was greater than 50%, and that the prob-
ability of a correct outcome approaches 100%
as the size of the electorate increases (Young,
1988). Condorcet’s probabilistic explorations
at least indirectly influenced Madison as well
(Schofield, 2005b), specifically the proclama-
tion in Federalist #10 that ‘If the proportion of
fit characters be not less in the large than in the
small republic, the former will present a greater
option, and consequently a greater probability
of a fit choice’. (Hamilton et al., [1788]1998).
While many thinkers who subsequently took
the mantle of ‘Darwinist’ would continue to
think about evolution purely in terms of compe-
tition, following T. H. Huxley’s famous dictum
that nature ‘is on about the same level as the
gladiator’s show’ from the ‘point of view of the
moralist’ (Kropotkin et al., 1955), others would
focus on empirical observations of cooperation
in nature and seek to explain them. In particular,
 EVOLUTIONARY PSYCHOLOGY AND POLITICAL INSTITUTIONS
the Russian naturalist Peter Kropotkin’s obser-
vations led him to believe that evolution was
about combination as much as competition.
He observed that reciprocity was widespread,
even between species. Kropotkin’s analysis
of social insects, parental behavior, and asso-
ciative activities like pack hunting, integrated
with an analysis of medieval guilds, led him to
the conclusion that mutual aid was as much a
law of nature as mutual struggle. In addition
to anticipating much in the scientific study
of altruism, Kropotkin’s Mutual Aid remains
a foundational work of anarcho-communism
(Kropotkin, 2017).
J. B. S. Haldane (also a vocal communist)
was perhaps the first to begin formalizing
the link between genetic traits and sociality
(Haldane, 1941). In The Causes of Evolution,
Haldane first proposed that reciprocity could
spread in a population if the genes determining
it were carried by individuals whose offspring
benefitted from the presence of the gene in their
nearby relatives (Haldane, 1932). Similarly, R.
A. Fisher, whose Genetical Theory of Natural
Selection heralded the modern synthesis in
evolutionary biology, focused considerable
attention on how degrees of genetic relatedness
affected individuals living in different types of
populations (Fisher, [1930]2000).
It is telling that the politically conserva-
tive Fisher was attracted to understanding
how ‘distastefulness’ in insect larvae could
spread in a population. He reasoned that
nasty tasting larvae would provide increased
protection for their siblings by driving away
predators, such that the genetic benefit of an
eaten larvae could be substantial. Perhaps
Fisher’s politics motivated his interest in
distastefulness: recent research suggests that
sensitivity to disgust is linked to conserva-
tism (Murray, 2012). Sadly, both Haldane and
Fisher were advocates of eugenics (Haldane
coined the term ‘cloning’ and Fisher headed
the Department of Eugenics at University
College London), but unlike Haldane, Fisher
viewed the fall of civilizations as a function of
declining fertility rates of the upper classes,
and he was concerned about encouraging
173
allowances for large working-class families
and those who possessed less ‘innate capac-
ity for intellectual and emotional develop-
ment’ (Fisher, [1930]2000).
The greatest breakthrough, and the reunion
of evolutionary biology and political science,
began after W. D. Hamilton broke the code
for kin selection (Hamilton, 1964a, 1964b).
Adapting economic cost–benefit analysis,
Hamilton showed how natural selection
would allow for altruism to evolve while still
maximizing individual fitness. In a series of
articles, he demonstrated how natural selec-
tion can favor altruism between relatives
when the product of the relatedness of indi-
viduals and the benefits of reciprocity out-
weigh the individual costs. The concept of
inclusive fitness was born, as was Hamilton’s
legacy as one of the greatest evolutionary
theorists of the 20th century.
The next giant step would be taken by
Robert Trivers, who extended Hamilton’s work
to non-kin reciprocity, such as bird warning
calls, the symbiosis between cleaner and pred-
ator fish, and broader social interactions. His
1971 publication ‘The Evolution of Reciprocal
Altruism’ showed how net benefits could be
accrued between non-relatives through social
exchange, and how cheating and spiteful
behavior could be regulated through the evolu-
tion of moralistic aggression (Trivers, 1971).
Trivers’ work opened new opportunities for
the integration of the behavioral sciences, as he
sketched out how the analysis could extend to
complex, multi-party coordination, the emer-
gence of norms and collective punishments
(and rewards), generational return effects, and
the like (Trivers, 2006).
Soon after, John Maynard Smith (a student
of Haldane) and George R. Price (a friend of
Hamilton) would formalize the Evolutionarily
Stable Strategy (ESS) and initiate evolution-
ary game theory in ‘The Logic of Animal
Conflict’ (Smith and Price, 1973), directly
addressing Darwin’s dilemma of moral devel-
opment. An ESS is a strategy (social interac-
tion where agents compete over a resource,
such as food, with payoffs dependent on both
174 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
agents’ strategies) that is stable once it is fixed
in a population. If agents will not do better
(more offspring) by employing a different
strategy, or perturbations from ‘mutant’ strat-
egies cannot successfully invade and replace
it, the trait becomes fixed in the population as
the offspring of the winners replace those with
lower payoffs. The ESS has not only become
central to the explanation of human evolu-
tion in evolutionary psychology, it has been
adopted in political science as a means of
understanding evolution through institutions.
That pivotal moment came when W. D.
Hamilton teamed up with political scientist
Robert Axelrod to publish ‘The Evolution of
Cooperation’ in 1981 (Axelrod and Hamilton,
1981), one of the most cited publications in
political science (Peress, 2019). Axelrod’s
computer tournaments provided output for sim-
ulations using competing strategies in repeated
Prisoner’s Dilemma (PD) games. In such
games, both agents know they would do better
if they cooperated, but the payoff is higher for
individuals to defect rather than be exploited.
Axelrod and Hamilton described how an elo-
quently simple logic of cooperation (essen-
tially do unto others, the ‘Tit for Tat’ strategy
submitted by anthropologist Anatol Rapaport)
could emerge as an ESS, given a high-enough
probability of future interaction between coop-
erative agents. Their analysis demonstrated
how ‘the benefits of life are disproportionately
available to cooperating groups’ and from it
they drew several strategic considerations:
• Be nice: don’t be the first to defect.
• Be provocable: return defection for defection,
cooperation for cooperation.
• Don’t be envious: focus on maximizing your own
‘score’, as opposed to ensuring your score is
higher than your ‘partner’s’.
• Don’t be too clever: signal clarity is crucial.
Successive work on iterated PD has relaxed
the highly simplifying assumptions built into
the original model, continuing to produce
important insights, including those about the
importance of forgiveness and reputation
(Nowak, 2006). Today, the study of
cooperation is a fertile interdisciplinary field,
identifying multiple stable states of coopera-
tion, the foundations of pro- and anti-social
behavior, the crucial role of structured interac-
tion, and the ways that institutions cultivate
cooperation (Alford and Hibbing, 2004;
Lopez, 2017; The Cooperative Human, 2018).
One of the most relevant ongoing debates
concerning institutional analysis has to do
with the concept of ‘strong reciprocity’ and
the role of institutions in sustaining coop-
eration. A body of scholars have challenged
the orthodox view of the reciprocal altruist
as self-regarding, developing an alternative
model of group-beneficial pro-social behav-
ior (Bowles et al., 2003; Bowles and Gintis,
2011; Fehr and Gächter, 2002; Henrich and
Boyd, 1998). Whereas kin selection, recipro-
cal altruism, indirect reciprocity, and signal-
ing explain behaviors that appear costly but
are repaid to genetic relatives, the strong-
reciprocity hypothesis posits that a genuinely
altruistic behavior can be adaptive, but crit-
ics are unconvinced (Burnham and Johnson,
2016). The answer to this controversy is
immensely important for the design of politi-
cal institutions, and it has rightly become a
major focus of scientific attention (Abbot,
et  al., 2011; Ferriere and Michod, 2011;
Herre and Wcislo, 2011; Nowak et al., 2010).
THE EMERGENCE OF COMPLEX
POLITICAL SYSTEMS
The use of biological metaphors to explain the
emergence and persistence of political sys-
tems dates back at least to Thomas Hobbes’
Leviathan with its cover art of the Sovereign
King, whose body is literally constituted by
the multitude of citizens, obedient co-signers
of the social contract, the body politic repre-
sented by one man (Hobbes, [1651]2009).
Walter Bagehot’s ‘Physics and Politics’ was
probably the first to explicitly apply Darwinian
selection and inheritance concepts to political
society (Bagehot, [1869]2009). Writing in
 EVOLUTIONARY PSYCHOLOGY AND POLITICAL INSTITUTIONS
1869, Bagehot made a case for liberal-demo-
cratic institutions having emerged from more
conformist, dictatorial restraints as a moral
achievement. The selective process had sup-
posedly refined the nervous systems and
moral capacity of ‘accomplished’ elites, cre-
ating opportunity for more complex decision-
making institutions and social progress.
Bagehot was also among the earliest in a
long line of ‘Social Darwinists’ to leverage
pseudo-scientific accounts of racial inequali-
ties to justify an ideological theory of the state.
John William Burgess similarly believed that
only ‘superior’ races had acknowledged the
moral ‘duties of civilization’ (Gunnell, 2004).
Burgess established the US political science
degree at Columbia University, where he
taught comparative constitutional law, and
his vision of political science emphasized
the training of career government bureau-
crats, as well as civic education as a sort of
‘democratic’ training, within the confines of
his racist ideology. After World War I, and
the embrace of scientific racism within fas-
cism and Nazism during World War II, social
Darwinism was largely discredited and aban-
doned by political science, at least publicly.
Post-war social scientists turned to formal
social choice and game theory as a framework
to explain political institutions. Properties of
social-decision rules, or ‘constitutional condi-
tions’ in the words of Kenneth Arrow, where
individual values are treated as inputs, became
a focal point of analysis (Arrow, 1970). One
of the most influential political scientists to
emerge from this period was Robert Dahl,
whose Arrow-inspired analysis of social
choice procedures shaped our understand-
ing of how popular sovereignty and political
equality are instituted (Dahl, 1989, 2006).
Nevertheless, Dahl’s image of the locus of
democracy in US government as fluid, plu-
ralist bargaining assumed an equilibrium of
social consensus, with no direct focus on evo-
lution or adaptation (Gunnell, 2004).
Dissatisfied with the historical bent of
most political science and unconvinced by
Dahl’s group bargaining theory of political
175
behavior, David Easton’s theoretical explo-
rations reflected his study of physiology and
systems biology, specifically the concept of
homeostasis and the capacity of evolved sys-
tems to react to environmental stimuli with
equilibrating responses (Cannon, 1963). As
a member of the University of Chicago’s
Committee on the Behavioral Sciences, he
was committed to the integration of biological
and social scientific knowledge, co-author-
ing ‘Projects and Problems of Homeostatic
Models in the Behavioral Sciences’ with psy-
chologist James G. Miller and anthropolo-
gist Anatol Rapoport, among others, in 1953
(Fontaine, 2016). Easton’s work transformed
the study of political institutions by nesting
them within a living-systems framework.
Easton was primarily interested in under-
standing how institutional arrangements regu-
late demands on a political system, shaping
the way that social choices are put into effect
as policy outcomes, which he elaborated in
A Framework for Political Analysis and A
Systems Analysis of Political Life (Easton,
1965, 1979). Among his many contributions to
the study of institutions, Easton’s ideas about
how regimes generate support through the
allocation of biological and social resources,
and regulation of social values, stand out
(Easton, 1979). Easton was among the first to
flesh out how emotive attachment to codified
social roles, and the social status they yield,
supply the behavioral energy upon which a
political system depends for its survival.
In the last few decades there has been
increasing convergence of Easton’s macro-
level type of structural-constraints analysis
and the micro-foundations of cooperative
game theory. Formalization of political author-
ity occurs when communicative artifacts,
from mating and marriage practices to land-
tenure customs, nomenclature, measurement,
and other forms of codification, instantiate a
‘high degree of mutual predictability’ with
large-scale coordination of, and thus con-
trol over, the allocation of biological and
social resources (Easton, 1979: 329). Just as
valuable biological information is correlated
176 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
across generations through the germ line,
institutional transmission of communicative
artifacts sustains biocultural complexes that
can greatly enhance the fitness of partici-
pants (Corning, 1995). Indeed, the produc-
tion and management of knowledge is itself a
commons dilemma (Frischmann et al., 2014).
National constitutions are rules for strat-
egy selection. That is, constitutional regimes
operate to regulate social niche-construction
strategies (Santa Fe Institute, 2016). As Brian
Skyrms notes in Evolution of the Social
Contract (2014), here strategies come to the
fore as units of selection, and individuals
recede into the background (much as genes
do in the biological literature). The success of
collectively selected strategies (laws) is based
on their success as behavioral phenotypes, and
how well populations converge on successful
strategies of social interaction (Skyrms, 2014).
Our understanding of institutions as carri-
ers of cultural transmission has been built on
decades of successful modeling of coopera-
tion, complexity, and culture (Axelrod, 2006;
Cavalli-Sforza and Feldman, 1981; Creanza
and Feldman, 2014).
Studies of the evolution of complexity
show that, as regulators of niche construc-
tion, political institutions face some uni-
versal information-processing problems.
For example, the tradeoff between exploring
alternative strategies and reliance on the sta-
tus quo is a general dilemma in organizational
productivity (March, 1991). The ‘explore/
exploit’ dilemma arises from the inability to
optimize both the exploration of potentially
adaptive/productive strategies, and cashing
in on the value of e­ xisting strategies. Without
commitment to a strategy, a system can ‘boil’
in the endless search for optimal alternatives,
but without generating variation there are fewer
alternatives to optimize, making it less likely to
find a better alternative than the status quo.
In the context of political institutions these
constraints correspond to the bounds of politi-
cal authority and social choice. The potential
range of participation in collective decision
making for a population ranges from N (total
population) to one, or pure dictatorship.
Both arrangements constitute cooperative
regimes in the sense that in a regime of N
decision makers, all preferences are taken
into consideration, maximizing the com-
munication channels and flow of informa-
tion to be processed, but also coordination
costs (the resources required to take every-
one’s preferences into account) and potential
‘boiling’ as each individual does their own
thing. Alternatively, pure dictatorship may be
the least costly social choice mechanism in
terms of coordination costs, but conformity
costs (everyone obeys the dictator) tend to be
extreme, in part because there is little effort
to search the landscape for mutually benefi-
cial strategies (Page, 2008; Zhou, 2011).
This tradeoff maps closely onto the solu-
tions typically proposed for the management
of public goods, or the ‘tragedy of the com-
mons’. Ecologist Garrett Hardin famously
argued in several papers (around the same
time that models of evolutionary coopera-
tion were emerging) that species are gener-
ally unable to cooperate for the greater good
(Hardin, 1968). In addition to being another
Malthusian throwback who believed that
those who won superior intellect through
the genetic lottery (whites, of course) should
deploy abortion and sterilization to maintain
limited population growth because ‘freedom
to breed will bring ruin to all’ (Hardin, 1968:
1248), he popularized the problem of main-
taining public goods through his analogy of
livestock management on shared grasslands,
an iterated PD game emphasizing the free-
rider problem and defection as rational in the
short term but irrational in the long run.
Hardin’s solutions to the tragedy were either
privatization and strong property rights, in order
to incentivize individual owners to be stewards
over their own land, or the very Hobbesian-
sounding ‘mutual coercion, mutually agreed
upon’ typically interpreted as collectivization.
Willfully or otherwise, Hardin was ignorant
of the degree to which many commons were
already being governed under ‘mutually coer-
cive’ social structures that have evolved to
 EVOLUTIONARY PSYCHOLOGY AND POLITICAL INSTITUTIONS
manage the ‘explore/exploit’ dilemma without
resorting to either strict privatization or col-
lectivization. No individual has played as
large a role in demonstrating the capacity
of local organizations to manage common
resources as Elinor Ostrom (Ostrom, 1990,
2010, 2013). While Ostrom’s work fits within
the broader fields of political economy and
organizational behavior (Levi, 1989; March
and Olson, 1971; Olsen, 1976), her approach
to core design principles, the Institutional
Analysis and Development (IAD) framework,
is distinctly Darwinian. Ostrom’s acceptance
speech for the 2009 Nobel prize in economics,
‘Do Institutions Evolve?’, recounts decades of
research she and colleagues have developed to
understand how biophysical systems, partici-
pant heterogeneity, and operational rules shape
the governance of commons (Ostrom, 2013).
A set of ‘default’ rules highlights what Ostrom
has discovered as core design principles:
• Clearly defined group boundaries
• Proportionally equivalent costs and benefits
• Collective-choice mechanisms
• Monitoring
• Graduated sanctions
• Fast and fair conflict resolution
• Polycentric governance (tiered rule-making)
The work of Ostrom and other political
economists, including Douglas North,
Margaret Levi, Barry Weingast, and Geoffrey
Hodgson, has engendered a generalized
Darwinian approach to the study of institu-
tional change that stands independent of
genetics and biology (Aldrich et  al., 2008;
Fürstenberg, 2016; Hodgson, 2004; Levi and
Weingast, 2019; Wilson and Gowdy, 2013).
We have learned much from empirical field
studies that have demonstrated the adaptive-
ness of local communities, including tribal
communities that Hardin surely would not
have imagined were cooperatively stable.
Large-scale societies require complex,
adaptive institutions to transmit norms of
conflict resolution and morality (Bowles
et al., 2003; Bowles and Gintis, 2011; Ehrlich
and Levin, 2005; Levin, 2014). Legal systems
177
exhibit properties that Walker and Davies
(2012) characterize as an informational struc-
ture that gains causal efficacy over matter, in
this case, behavior: they are explicit instruc-
tions, decontextualized from specific behav-
iors to classes of behaviors, applicable to
classes of agents, designed to yield high lev-
els of predictability (Bowles and Gintis, 2011;
Boyer and Petersen, 2012). Rather than being
directly transmitted, legal expectations are
learned, deriving from hard-wired moral intu-
itions, but converted into ‘publicly scrutable
processes’ that modify social interaction from
the ‘top down’ (Boyer and Petersen, 2012).
According to Boyer and Peterson, like our
biophysical equipment, our evolved cogni-
tive systems operate more reliably in match-
ing environments (ideally the environment of
evolutionary adaptedness). As a result, we are
motivated to match our environments to those
cognitive systems, while the modification of
environmental niches creates and maintains
selective pressures that favor adaptions matched
to them (Laland et al., 1999). Richard Dawkins
famously called this sort of niche construction
The Extended Phenotype (Dawkins, 1999),
based on the evolutionary logic that the qual-
ity of an environmental niche (a spider web, a
beaver dam, a human settlement) is correlated
with genetic variation, in that an allele for bet-
ter niche construction enhances the fitness of
the organisms expressing it. In an environment
with good nest-building materials, the best nest
builders will proliferate the genes for building
nests with those materials, as their offspring
increase in frequency through the population.
Constitutional environments will likewise favor
the selection and spread of behavior traits and
cultural norms, such as a sense of justice, under
constitutional conditions that favor it.
DEMOCRACY: A MAJOR
EVOLUTIONARY TRANSITION?
Representative democracy encompasses a
range of regimes that have emerged quite
178 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
recently on the human scene, possibly in
response to the conformity costs imposed by
more authoritarian, dictatorial regime struc-
tures (Latner, 2017). The core design princi-
ple of democracy is political equality, or
fairness. Political theory on procedural jus-
tice has contributed to our understanding of
cooperation at all levels of life through a sort
of cross fertilization. Specifically, the politi-
cal philosopher John Rawls’ concept of the
‘veil of ignorance’ played prominently in the
social theory of another Michigan biologist,
Richard D. Alexander (Alexander, 2017;
Rawls, 1999). The basic logic is that if the
only way to be sure one gets ahead is to pro-
mote rules that will improve overall welfare,
fair laws should be favored for selection.
What emerges is a society that applies laws
equally to individuals regardless of their
status, or justice as fairness.
Alexander saw that the enforcement of
mutualistic cooperation requires limiting
opportunities for cheating, such that agents
can only increase their success by increas-
ing the success of others (Frank, 2013). He
also saw how natural selection could favor
the suppression of internal competition by
drawing on the work of E. G. Leigh, probably
the first modern biologist to use a political
analogy, equating the genome to ‘a parliament
of genes: each acts in its own self-interest, but
if its acts hurt the others, they will combine
together to suppress it’ (Leigh, 1971). Later
analysis has supported the idea that the cellu-
lar infrastructure of meiosis in sexual repro-
duction is favored, in part, as a mechanism to
thwart parasitic conflict at the genetic level
(Burt and Trivers, 2008; Howard and Lively,
1994). Similarly, Alexander’s argument that
socially imposed monogamy levels reproduc-
tive opportunity, and in doing so reduces the
pool of unmarried men, has found support in
research that shows monogamy is associated
with reduced rates of homicide, rape, and
related violence (Henrich et al., 2012).
Bryan Skyrms’ Evolution of the Social
Contract and later work brings us full cir-
cle through the evolution of cooperation
to the operation of constitutional structure
(Skyrms, 2014). His eloquent essay ‘Sex and
Justice’ (1994) compares the puzzle of sex
ratios (which led to Fisher’s game-theoretic
insights) to the evolution of justice. Skyrms
shows how, like many observed sex ratios, fair
division (50/50) is an attractive equilibrium
in repeated interactions. Crucially, if interac-
tions between more cooperative agents can
be correlated (a higher probability that they
interact with one another instead of greedy
agents), then the cooperative norm ‘share and
share alike’ becomes a stronger evolutionary
attractor and will emerge as an ESS.
Skyrms shows how mechanisms for the
evolution of cooperation, from kin selection
to ‘group’ selection, spatial interaction, and
the repression of competition, are all linked
to increasing correlation and cooperative
association (Skyrms, 2014). Skyrms has also
brought attention to the problem of ‘negative
correlation’ and how spiteful behavior can
be sustained through correlated interactions,
such as when a reputation for ‘fighting too
hard’ (to the detriment of self and opponent)
may enable agents to win future contests more
easily in repeated encounters (Skyrms, 2014).
The likelihood that political equality is a
fit alternative against dictatorial hierarchy
and anarchy is empirically supported across
a variety of fields. First, recent developments
in social choice theory have confirmed that
proportional representation, which most accu-
rately translates votes into seats of political
power, is the closest approximation to politi-
cal equality in electoral-system design (Hout
and McGann, 2009). Contemporary experi-
mental tests further support Condorcet’s posi-
tion that majority rule outperforms alternative
strategies, including dictatorship (Hastie
and Kameda, 2005; Sorkin et al., 1998), and
regimes that more closely approximate politi-
cal equality (PR and simple majority rule)
tend toward greater redistribution and respon-
siveness, as predicted (Acemoglu et al., 2009;
McGann and Latner, 2013). Representative
assemblies, electoral systems, thresholds
for legislative decision making and policy
 EVOLUTIONARY PSYCHOLOGY AND POLITICAL INSTITUTIONS
execution all reflect fundamental features of
institutional niche construction and coopera-
tive decision making (Latner, 2017).
Steven Frank has, more than nearly any
other evolutionary thinker, shown how the
policing of competition, or reduction of
opportunities for exploitative behavior, cor-
relates individual payoffs and can sustain
group cohesion (Frank, 2003, 2011, 2013).
Reducing opportunities for exploitative
behavior through political equality may result
in both higher levels of redistributive sharing
and flexible responsiveness in resource man-
agement, making it an evolutionary attractive
equilibrium (McGann and Latner, 2013).
The historical record is also supportive of
the evolutionary democracy hypothesis. The
number of regimes holding minimally free and
fair elections has nearly doubled since 1990,
from 69 to 122 according to Freedom House
(‘Freedom in the World’, 2014). Moreover,
among electoral democracies, major elec-
toral reforms have been decisively in the
direction of more proportional representation
(Soudriette and Ellis, 2006). This follows the
evolution from simple ‘originating’ systems
in early electoral systems to electoral rules
that reduced the frequency of single-party
dominance that had emerged (Colomer, 2001,
2007). At the other extreme, hyper-permissive
systems demonstrably produce greater insta-
bility in governing coalitions, lower cabinet
duration, and greater difficulty committing to
comprehensive policy platforms (Shugart and
Wattenberg, 2001; Taagepera, 2007). Reforms
in these systems have generally maintained
proportionality along with consolidating legis-
lative power in the direction of greater account-
ability (Shugart and Wattenberg, 2001).
Of course, political institutions do not
operate exogenously on behavior, independ-
ent of the complexity of other social systems.
Population heterogeneity and inequality in
social resources obviously shape institutional
performance and the prospects of regime
persistence. For example, cultural mutation
and cycles of stability are highly sensitive to
interactions between transmission, selection,
179
and assorting of populations. For example,
Creanza and Feldman have considered educa-
tion norms, specifically the belief that women
should receive secondary education and delay
childbirth (Creanza and Feldman, 2014). The
probability of fixation of the belief is shaped
by lower fertility among those women who
take part, as well as lower infant mortality
among the same families, and future sorting
in offspring mating practices. Similar dynam-
ics may impact the norm of democratic par-
ticipation and investment in the considerable
responsibilities of democratic citizenship, not
to mention the interaction between education,
socialization, and support for democratic
institutions, especially for women (Fox and
Lawless, 2014).
Asymmetries between the (de jure) consti-
tutional and (de facto) economic allocation
of political power also shape the dynamics
of regime stability. Daron Acemoglu and col-
leagues have produced a number of analyses
proposing that the threat of upheaval and
benefits of anticipated economic growth have
been primary drivers of elite-led expansion
in collective decision making (Acemoglu
et al., 2009, 2011; Acemoglu and Robinson,
2013). Peter Turchin has proposed a broader
model of democratic evolution that empha-
sizes the role of exogenous threats, with
a similar sequencing that cooperation and
enfranchisement are frequently followed by
periods of economic inequity and political
instability (Turchin, 2013, 2016). These and
other analyses point toward higher potential
for upheaval under greater economic inequal-
ity, when lower classes (and their offspring)
feel they have nothing to lose, and when the
prospective benefits of a major transition out-
weigh the perceived costs of disruption.
The interplay of group conditions and spe-
cific conflict-reduction mechanisms can deter-
mine the fate of a regime’s future. If increasing
inequality or prolonged economic stagnation
fuel animosity between competing economic
classes, it will lead to increased demand for
the powerful to invest in economic and politi-
cal mechanisms to better preserve common
180 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
goods, and consequently the stronger taking
over more institutional control (Frank, 2011).
Whether that control is more equitable or
‘extractive’ (parasitic) may in turn depend on
the prior fixation of economic and cultural
traits, like democratic norms among elites.
Further, the adequacy of a political solution
will depend on whether competition among
reformers can be sustained and a coalition
holds, for example by requiring coalition
building within broad-based political parties,
or whether fragmented interests and the work
of compromise must be channeled into a rep-
resentative assembly (Balinski and Young,
2001; Cox, 1997).
In electoral system science, the general
pattern that proportional electoral systems are
associated with higher rates of women’s repre-
sentation is a puzzle that requires consistency
between macro- and micro-level explanations
(Reynolds et al., 2005). One consistent model
that has been applied to the puzzle is social
contagion, where smaller parties in propor-
tional systems differentiate themselves (at
lower cost than under single-seat systems)
by listing more women on party lists, driv-
ing larger parties to follow suit (Matland and
Studlar, 1996). Several outlier systems such
as Israel and Malta exhibit very low percent-
ages of women in parliament despite using
rather pure versions of PR, and incorporating
models of variance in child-rearing strategies
would provide a fuller understanding of these
strategies (Lane, 1995; Rule, 1987).
Evolutionary psychology has also contrib-
uted to our understanding of sex and gender
dynamics in campaign and election studies.
For example, a large body of research has
demonstrated that men and women differ in
candidate support based on factors like facial
appearance and body image, with men pre-
ferring more attractive female candidates
and women preferring approachable male
candidates (Chiao et al., 2008; Dolan, 2014).
However, electoral contexts interact with
these abstract preferences in ways that shape
vote choice, especially partisan signaling
and contextual priming emphasizing either
intergroup conflict or cooperation, such
that both masculine (conflict) and feminine
(cooperation) traits can positively contribute
to the perception of effective leadership traits
(Dolan and Lynch, 2014; Grabo and van
Vugt, 2018). The sub-field of evolutionary
feminist studies is challenging stereotypes of
bio-essentialism head on, and will surely aid
our understanding of institutional design with
relation to women’s and gender studies (Buss
and Malamuth, 1996; Feminist Evolutionary
Perspectives, 2019).
Another example of the evolution of
democracy is seen in contestation over vot-
ing rights in the United States. Epperly and
colleagues have documented the dynamics at
work in voter suppression as an exploitative
form of ‘cooperation’ (Epperly et al., 2019).
They show that under conditions of low state
capacity and legibility (formalized authority
through administrative records, etc.), as well
as external constraints on state legislative
suppression (the federal government), voter
suppression under Reconstruction took the
form of decentralized intimidation like lynch-
ing that peaked just before federal restraints
on legalized suppression were relaxed, allow-
ing Southern Democrats to re-take control of
several state legislatures. In the mid 1890s,
the inefficient and enforcement-costly strat-
egy of lynching declined as the codified,
more predictable discriminatory laws of Jim
Crow (poll taxes, registration requirements,
multiple-box voting, secret ballots, literacy
tests, property tests, understanding clauses,
grandfather clauses, and the white primary)
came online. These electoral barriers on the
representation and possible transmission
of less oppressive social strategies kept the
Southern caste system in place for several
generations, and with it the cultural norms
of segregation, revulsion at inter-racial mar-
riage, and belief in the inherent inferiority of
African Americans.
The Civil Rights Movement, and Civil
Rights Act of 1964, Voting Rights Act of
1965, and accompanying court cases marked
a behavioral and legislative tipping point in
 EVOLUTIONARY PSYCHOLOGY AND POLITICAL INSTITUTIONS
electoral reforms to expand racial represen-
tation and socioeconomic opportunity in
the United States (Davidson and Grofman,
1994). The cultural impact on norms has been
substantial, but complex. While Jim Crow
era norms have largely collapsed within the
white population, and broad support for
political equality has spread across all popu-
lations, support for government efforts to
address segregation and other discriminatory
practices is still frequently opposed (Bobo
et al., 2012). Surviving racial stereotypes and
negative views of racial minorities today tend
to be based in characterizations of group cul-
ture, rather than biology (Bazian, 2016; Bobo
et  al., 2012; Kaufmann, 2019). Selective
pressures, and the institutional regulation of
cooperation and competition, have altered the
fitness of racial norms.
Over the last decade, Americans have
experienced the consequences of relaxa-
tions on federal constraints with the weak-
ening of the Voting Rights Act and reduced
judicial oversight of discriminatory practices
(Bentele and O’Brien, 2013; CNN, 2019;
Keena et al., 2017). Predictably, attempts to
discriminate against black voters and other
groups are on the rise again, and as in the
post-Reconstruction South, voter suppression
is increasingly couched in terms of partisan
calculations (Hasen, 2014). While existing
laws arguably make these new efforts, which
range from gerrymandering to voter list purg-
ing, voter ID laws, and proof of citizenship
requirements, less effective than Jim Crow
1.0, there is little reason to believe that Jim
Crow 2.0 will not become more egregious
over time, or that efforts will not be made to
restrict the franchise further if the Republican
Party does not expand beyond its shrinking
demographic base of support. In this event,
the importance of evolutionary psychology
is even more apparent, as scientists will need
every tool at their disposal to educate and
advocate for evidence-based policies to help
inoculate the public, and fight back against
the resurgence of racism and white national-
ism in our electoral ecosystem.
181
CONCLUSION
This brief summary of what the evolutionary
approach has provided to our current under-
standing of institutional design and perfor-
mance is far from exhaustive. But one final
sign of the extent of recent integration between
institutionalists and biologists that must be
mentioned is the contributions that scholars of
political institutions are making to biology.
Political scientists are now researching animal
institutions and collective decision making, a
sure sign that the ‘life sciences’ properly
understood are integrating under the umbrella
of a generalized Darwinism (Akcay et  al.,
2013; Conradt and List, 2009).
The claim that human behavior and insti-
tutions are not reducible to biological sci-
ences is true insofar as we narrow biology
to genetic studies or similar micro-level pro-
cesses. But political institutions are emer-
gent phenomena, have their own laws, and
require their own sciences, just as the science
of complexity is not reducible to molecular
physics (Taagepera, 2008). Rather, unifica-
tion posits that there be consistency between
micro- and macro-behavioral explanations.
For example, models of political institutions
and partisan competition should not assume
that agents value the welfare of others more
than their own, that men and women are
equally prone to engage in extra-constitu-
tional, violent confrontation, or that conven-
tions of masculinity and feminism have no
biological roots.
An even broader challenge, the levels
of selection controversy over the relative
importance of kin selection and reciprocity
versus group selection, has animated all of
evolutionary science, and might benefit from
better incorporation of institutional theory.
As already discussed, political institutions
that mobilize, aggregate, synthesize, and
select competing policy strategies provide a
clear Darwinian process with which to study
levels of selection, and the potential to trace
return benefits and other return effects across
generations.
182 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
REFERENCES
Abbot, P. et. al. (2011). ‘Inclusive Fitness Theory
and Eusociality’, Nature, 471: E1–E4. Retrieved
from https://www.nature.com/articles/
nature09831?proof=true19
Acemoglu, D., Egorov, G., & Sonin, K. (2009).
Political selection and persistence of bad
governments (Working Paper No. 15230).
Retrieved from National Bureau of Economic
Research website: www.nber.org/papers/
w15230
Acemoglu, D., Egorov, G., & Sonin, K. (2011).
Political model of social evolution. Proceed-
ings of the National Academy of Sciences,
108(Supplement 4), 21292–21296. https://
doi.org/10.1073/pnas.1019454108
Acemoglu, D., & Robinson, J. (2013). Why
nations fail: The origins of power, prosperity,
and poverty (Reprint edition). New York:
Crown Business.
Akcay, E., Roughgarden, J., Fearon, J. D.,
Ferejohn, J. A., & Weingast, B. R. (2013).
Biological institutions: The political science of
animal cooperation (SSRN Scholarly Paper
No. ID 2370952). Retrieved from Social
Science Research Network website: http://
papers.ssrn.com/abstract=2370952
Aldrich, H. E., Hodgson, G. M., Hull, D. L.,
Knudsen, T., Mokyr, J., & Vanberg, V. J.
(2008). In defence of generalized Darwinism.
Journal of Evolutionary Economics, 18(5),
577–596. https://doi.org/10.1007/s00191-
008-0110-z
Alexander, R. (2017). The biology of moral
systems. New York: Routledge.
Alford, J., & Hibbing, J. (2004). The Origin of
Politics: An Evolutionary Theory of Political
Behavior. Perspectives on Politics, 2(4), 707–
723. doi:10.1017/S1537592704040460
Arrow, K. J. (1970). Social choice and individual
values (2nd ed.). New Haven: Yale University
Press.
Axelrod, R. (2006). The evolution of cooperation
(Revised). Basic Books, Inc., New York.
Axelrod, R., & Hamilton, W. D. (1981). The
evolution of cooperation. Science, 211(4489),
1390–1396. https://doi.org/10.1126/
science.7466396
Bagehot, W. [1869](2009). Physics and politics.
CreateSpace, an Amazon Publishing Company.
Scotts Valley, California.
Balinski, M. L., & Young, H. P. (2001). Fair
representation: Meeting the ideal of one
man, one vote (2nd ed.). Washington, DC:
Brookings Institution Press.
Bazian, H. (2016, October 12). Police violence in
America and compounded racism. Retrieved
November 2, 2016, from Hatem Bazian web-
site: www.hatembazian.com/content/police-
violence-in-america-and-compounded-racism/
Bentele, K. G., & O’Brien, E. E. (2013). Jim
Crow 2.0? Why states consider and adopt
restrictive voter access policies. Perspectives
on Politics, 11(4), 1088–1116. https://doi.
org/10.1017/S1537592713002843
Binmore, K. G. (1994). Game theory and the
social contract: Just playing. Cambridge: MIT
Press.
Bobo, L. D., Charles, C. Z., Krysan, M., & Sim-
mons, A. D. (2012). The real record on racial
attitudes. Retrieved from https://dash.har-
vard.edu/handle/1/32197879
Bowles, S., Choi, J.-K., & Hopfensitz, A. (2003).
The co-evolution of individual behaviors and
social institutions. Journal of Theoretical Biol-
ogy, 223(2), 135–147. Retrieved from https://
doi.org/10.1016/S0022-5193(03)00060-2
Bowles, S., & Gintis, H. (2011). A cooperative
species: Human reciprocity and its evolution.
Princeton: Princeton University Press.
Boyer, P., & Petersen, M. B. (2012). The natural-
ness of (many) social institutions: Evolved
cognition as their foundation. Journal of
Institutional Economics, 8(01), 1–25.
Burnham, Terence & Johnson, Dominic. (2005).
The Biological and Evolutionary Logic of
Human Cooperation. Analyse & Kritik. 27.
113–135. 10.1515/auk-2005-0107.
Burt, A., & Trivers, R. (2008). Genes in conflict:
The biology of selfish genetic elements (1st
ed.). Cambridge, MA: Belknap Press.
Buss, D. M., & Malamuth, N. M. (Eds.). (1996).
Sex, power, conflict: Evolutionary and feminist
perspectives. New York: Oxford University Press.
Cannon, W. B. (1963). The wisdom of the body
(Revised and enlarged ed.). New York: W. W.
Norton & Company.
Cavalli-Sforza, L. L., & Feldman, M. W. (1981).
Cultural transmission and evolution: A
quantitative approach. Princeton: Princeton
University Press.
Chiao, J. Y., Bowman, N. E., & Gill, H. (2008).
The political gender gap: Gender bias in
 EVOLUTIONARY PSYCHOLOGY AND POLITICAL INSTITUTIONS
facial inferences that predict voting behavior.
PLoS ONE, 3(10). e3666. https://doi.
org/10.1371/journal.pone.0003666
CNN, J. B. (2019, January 21). The Supreme
Court takes on MLK’s legacy. Retrieved
August 4, 2019, from CNN website: www.
cnn.com/2019/01/20/us/mlk-legacy-
supreme-court/index.html
Colomer, J. M. (2001). Political institutions:
Democracy and social choice. Oxford
University Press, USA. Retrieved from www.
oxfordscholarship.com/view/10.1093/01992
4183X.001.0001/acprof-9780199241835
Colomer, J. M. (2007). On the origins of
electoral systems and political parties: The
role of elections in multi-member districts.
Electoral Studies, 26(2), 262–273. https://
doi.org/10.1016/j.electstud.2006.02.002
Conradt, L., & List, C. (2009). Group decisions
in humans and animals: A survey. Philosophi-
cal Transactions of the Royal Society of
London B: Biological Sciences, 364(1518),
719–742. https://doi.org/10.1098/rstb.
2008.0276
Corning, P. A. (1995). Synergy and self-
organization in the evolution of complex
systems. Systems Research, 12(2), 89–121.
https://doi.org/10.1002/sres.3850120204
Cox, Gary W. (1997). Making votes count.
Cambridge, UK; New York: Cambridge
University Press.
Creanza, N., & Feldman, M. W. (2014).
Complexity in models of cultural niche
construction with selection and homophily.
Proceedings of the National Academy of
Sciences, 111(Supplement 3), 10830–10837.
https://doi.org/10.1073/pnas.1400824111
Dahl, R. A. (2006). A preface to democratic
theory (Expanded, anniversary ed.). Chicago:
University of Chicago Press.
Dahl, R. A. (1989). Democracy and its critics.
New Haven: Yale University Press.
Darwin, C. (1871). The descent of man, and
selection in relation to sex. John Murray,
London.
Darwin, C. [1838](2011). The autobiography of
Charles Darwin. CreateSpace Independent
Publishing Platform.
Davidson, C., & Grofman, B. (1994). Quiet
revolution in the South: The impact of the
Voting Rights Act, 1965–1990. Princeton:
Princeton University Press.
183
Dawkins, R. (1999). The extended phenotype:
The long reach of the gene (Revised ed.).
Oxford; New York: Oxford University Press.
Dolan, K. (2014). When does gender matter?:
Women candidates and gender stereotypes
in American elections. Retrieved from www.
oxfordscholarship.com/view/10.1093/acpr
of:oso/9780199968275.001.0001/acprof-
9780199968275
Dolan, K., & Lynch, T. (2014). It takes a survey:
Understanding gender stereotypes, abstract
attitudes, and voting for women candidates.
American Politics Research, 42(4), 656–676.
https://doi.org/10.1177/1532673X13503034
Easton, D. (1965). A framework for political
analysis. Prentice-Hall, New Jersey.
Easton, D. (1979). A systems analysis of political
life. Chicago: University of Chicago Press.
Ehrlich, P. R., & Levin, S. A. (2005). The
evolution of norms. PLOS Biology, 3(6),
e194. https://doi.org/10.1371/journal.pbio.
0030194
Epperly, B., Witko, C., Strickler, R., & White, P.
(2019). Rule by violence, rule by law: Lynching,
Jim Crow, and the continuing evolution of
voter suppression in the U.S. Perspectives on
Politics, 1–14. Retrieved from https://doi.org/
10.1017/S1537592718003584
Fehr E, Gächter S. Altruistic punishment in
humans. Nature. 2002; 415(6868):137–140.
doi:10.1038/415137a
Feminist Evolutionary Perspectives. (2019).
Feminist evolutionary perspectives society.
Retrieved May 22, 2020 from Feminist
Evolutionary Perspectives website: www.
facebook.com/groups/122502515165/
Ferriere, R., Michod, R. Inclusive fitness in
evolution. Nature 471, E6–E8 (2011). https://
doi.org/10.1038/nature09834
Fisher, R. A. [1930](2000). The genetical theory
of natural selection (1st ed.; J. H. Bennett, Ed.).
Oxford: Oxford University Press.
Fontaine, P. (2016). Walking the tightrope: The
committee on the behavioral sciences and
academic cultures at the University of Chicago,
1949–1955. Journal of the History of the
Behavioral Sciences. 52(4). 349–370. Retrieved
from https://www.academia.edu/29432063/
Walking_the_Tightrope_The_Committee_
on_the_Behavioral_Sciences_and_Academic_
Cultures_at_the_University_of_Chicago_
1949_1955
184 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Fox, R. L., & Lawless, J. L. (2014). Uncovering
the origins of the gender gap in political
ambition. American Political Science Review,
108(3), 499–519. https://doi.org/10.1017/
S0003055414000227
Frank, S. A. (2003). Perspective: Repression of
competition and the evolution of
cooperation. Evolution; International Journal
of Organic Evolution, 57(4), 693–705.
Frank, S. A. (2011). “Evolutionary foundations
of cooperation and group cohesion.” in
Simon Levin (Ed.). In Games, Groups and the
Global Good. Springer-Verlag Berlin
Heidelberg Retrieved from http://arxiv.org/
abs/1112.3046
Frank, S. A. (2013). “Introduction: A new
theory of cooperation.” In K. Summers & B.
Crespi (Eds.), Human Social Behavior: The
Foundational Works of Richard D. Alexander.
Oxford University Press, USA.
Freedom in the World. (2014). Retrieved July
16, 2014, from Freedom House website:
www.freedomhouse.org/report/freedom-w
orld-aggregate-and-subcategory-scores#.
U8bXglZfTQs
Frischmann, B. M., Madison, M. J., &
Strandburg, K. J. (2014). Governing
knowledge commons. Oxford University
Press.
Fürstenberg, D. K. (2016). Evolutionary
institutionalism. Politics and the Life Sciences:
The Journal of the Association for Politics
and the Life Sciences, 35(1), 48–60.
Grabo, A., & van Vugt, M. (2018). Voting for a
male warrior or female peacekeeper? Testing
the evolutionary contingency hypothesis in the
2016 U.S. presidential elections. Evolutionary
Psychology, 16(2), 1474704918773267.
https://doi.org/10.1177/1474704918773267
Gunnell, J. G. (2004). Imagining the American
polity: Political science and the discourse of
democracy. University Park: Pennsylvania
State University Press.
Haldane, J. B. S. (1932). The causes of evolution.
Retrieved July 26, 2019, from Princeton
University Press website: https://press.
princeton.edu/titles/4618.html
Haldane, J. B. S. (1941). Concerning social
Darwinism. Science & Society, 5(4), 373–
375. Retrieved from JSTOR.
Hamilton, A., Madison, J., & Jay, J. [1788]
(1998, December 29). The federalist papers
No. 10. Retrieved July 24, 2019, from https://
avalon.law.yale.edu/18th_century/fed10.asp
Hamilton, W. D. (1964a). The genetical
evolution of social behaviour. I. Journal of
Theoretical Biology, 7(1), 1–16.
Hamilton, W. D. (1964b). The genetical
evolution of social behaviour. II. Journal of
Theoretical Biology, 7(1), 17–52.
Hardin, G. (1968). The tragedy of the commons.
Science, 162(3859), 1243–1248. https://doi.
org/10.1126/science.162.3859.1243
Hasen, R. (2014, January 7). ‘Race or party?:
How courts should think about Republican
efforts to make it harder to vote in North
Carolina and elsewhere’. Retrieved November
29, 2014, from Harvard Law Review website:
http://harvardlawreview.org/2014/01/race-or-
party-how-courts-should-think-about-republican-
efforts-to-make-it-harder-to-vote-in-north-
carolina-and-elsewhere/
Hastie, R., & Kameda, T. (2005). The robust
beauty of majority rules in group decisions.
Psychological Review, 112(2), 494–508. https://
doi.org/10.1037/0033-295X.112.2.494
Henrich, J., & Boyd, R. (1998). The evolution of
conformist transmission and the emergence
of between-group differences. Evolution and
Human Behavior, 19(4), 215–241. https://
doi.org/10.1016/S1090-5138(98)00018-X
Henrich, J., Boyd, R., & Richerson, P. J. (2012). The
puzzle of monogamous marriage. Philosophical
Transactions of the Royal Society of London B:
Biological Sciences, 367(1589), 657–669.
https://doi.org/10.1098/rstb.2011.0290
Herre, E., and Wcislo, W. (2011). In defence of
inclusive fitness theory. Nature 471, E8–E9.
https://doi.org/10.1038/nature09835
Hobbes, Thomas. (1651;2009). Leviathan: Or
the Matter, Form and Power of a
Commonwealth, Ecclesiastical and Civil.
Gutenberg Project. https://www.gutenberg.
org/files/3207/3207-h/3207-h.htm
Hodgson, G. M. (2004). The evolution of
institutional economics: Agency, Structure
and Darwinism in American Institutionalism.
London and New York: Routledge https://
doi.org/10.4324/9780203300350
Hout, E. van der, & McGann, A. J. (2009).
Liberal political equality implies proportional
representation. Social Choice and Welfare,
33(4), 617–627. https://doi.org/10.1007/
s00355-009-0382-8
 EVOLUTIONARY PSYCHOLOGY AND POLITICAL INSTITUTIONS
Howard, R. S., & Lively, C. M. (1994). Parasitism,
mutation accumulation and the maintenance
of sex. Nature, 367(6463), 554–557. https://
doi.org/10.1038/367554a0
Kaufmann, E. (2019, March 18). Americans are
divided by their views on race, not race itself.
The New York Times. Retrieved May 22,
2020 from www.nytimes.com/2019/03/18/
opinion/race-america-trump.html
Keena, A., McGann, A., & Smith, C. A. (2017,
October 25). The Supreme Court’s quiet
gerrymandering revolution and the road to
minority rule. Retrieved July 26, 2018, from
USAPP website: http://blogs.lse.ac.uk/
usappblog/2017/10/25/the-supreme-courts-
quiet-gerrymandering-revolution-and-the-
road-to-minority-rule/
Kropotkin, P. (2017). Mutual aid: A factor of
evolution (J. Duran, Ed.). CreateSpace
Independent Publishing Platform.
Kropotkin, P. A., Huxley, T., & Montagu, A.
(1955). Mutual aid and the struggle for
existence. Extending Horizons Books. Boston,
Massachusetts.
Laland, K. N., Odling-Smee, F. J., & Feldman,
M. W. (1999). Evolutionary consequences of
niche construction and their implications for
ecology. Proceedings of the National
Academy of Sciences of the United States of
America, 96(18), 10242–10247.
Lane, J. C. (1995). The election of women under
proportional representation: The case of
Malta. Democratization, 2(2), 140–157.
https://doi.org/10.1080/13510349508403433
Latner, M. (2017). Darwinian democracy? How
evolutionary theory informs constitutional
design. Handbook of Biology and Politics.
Retrieved from www.elgaronline.com
/view/edcoll/9781783476268/97817834762
68.00037.xml
Leigh, E. G. (1971). Adaptation and diversity:
Natural history and the mathematics of
evolution. Freeman, Cooper. San Francisco.
Levi, M. (1989). Of rule and revenue. California:
University of California Press.
Levi, M., & Weingast, B. R. (2019). Douglass
North’s theory of politics. PS: Political
Science & Politics, 52(2), 213–217. https://
doi.org/10.1017/S1049096518002111
Levin, S. A. (2014). Public goods in relation to
competition, cooperation, and spite.
Proceedings of the National Academy of
185
Sciences, 111(Supplement 3), 10838–10845.
https://doi.org/10.1073/pnas.1400830111
Lopez, A. (2017). Does Conflict Drive
Cooperation? The Evolution Institute. Retrieved
May 20th, 2020 https://evolution-institute.org/
cooperation-was-important-in-human-
evolution/
Madison, J. (1786). Equality: James Madison to
Thomas Jefferson. Retrieved December 1,
2017, from http://press-pubs.uchicago.edu/
founders/print_documents/v1ch15s33.html
Madison, J., Hamilton, A., & Jay, J. [1788]
(2008). The Federalist papers. CreateSpace
Independent Publishing Platform.
Malthus, T. [1798](2013). An essay on the
principle of population. J. Johnson, London.
March, J. G. (1991). Exploration and exploitation
in organizational learning. Organization
Science, 2(1), 71–87. https://doi.org/10.1287/
orsc.2.1.71
March, J. G., & Olsen, J. P. (1976). Ambiguity and
choice in organizations. Universitetsforlaget.
Oslo, Norway.
Matland, R. E., & Studlar, D. T. (1996). The
contagion of women candidates in single-
member district and proportional
representation electoral systems: Canada
and Norway. The Journal of Politics, 58(3),
707–733. https://doi.org/10.2307/2960439
McCoy, D. R. (1980). Jefferson and Madison on
Malthus: Population growth in Jeffersonian
political economy. The Virginia Magazine of
History and Biography, 88(3), 259–276.
McGann, A., & Latner, M. (2013). The calculus
of consensus democracy: Rethinking patterns
of democracy without veto players.
Comparative Political Studies, 46(7), 823–
850. Retrieved from https://doi.org/10.1177/
0010414012463883
Mclean, I. (2005). Before and after Publius: The
sources and influences of Madison’s political
thought. In S. Kernell (Ed.), James Madison:
The Theory and Practice of Republican
Government. Stanford University Press.
Redwood City, California.
Miller, James G. (1953). Profits and Problems of
Homeostatic Models in the Behavioral
Sciences – Introduction. Chicago Behavioral
Sciences, University of Michigan Biography
of Publications.
Murray, G. (2012, March 4). Are you easily
disgusted? You may be a Conservative.
186 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Retrieved July 26, 2019, from Psychology
Today website: www.psychologytoday.com/
blog/caveman-politics/201203/are-you-easily-
disgusted-you-may-be-conservative
Nowak, M. A. (2006). Five rules for the
evolution of cooperation. Science, 314(5805),
1560–1563. https://doi.org/10.1126/
science.1133755
Nowak, M., Tarnita, C., & Wilson, E. (2010). The
evolution of eusociality. Nature 466, 1057–
1062. https://doi.org/10.1038/nature09205
Olson, M. (1971). The logic of collective action:
Public goods and the theory of groups,
Second printing with a new preface and
appendix (Revised ed.). Cambridge: Harvard
University Press.
Ostrom, E. (1990). Governing the commons:
The evolution of institutions for collective
action (1st ed.). Cambridge; New York:
Cambridge University Press.
Ostrom, E. (2010). Beyond markets and states:
Polycentric governance of complex economic
systems. American Economic Review, 100(3),
641–672. https://doi.org/10.1257/aer.
100.3.641
Ostrom, E. (2013). Do institutions for collective
action evolve? Journal of Bioeconomics,
16(1), 3–30. https://doi.org/10.1007/s10818-
013-9154-8
Palmer, Tom G. (2002). “Madison and
Multiculturalism: Group Representation,
Group Rights and Constitutionalism”. In
John Samples (ed.), James Madison and the
Future of Limited Government, Cato Institute.
Page, S. E. (2008). The difference: How the
power of diversity creates better groups,
firms, schools, and societies (New edition
with a new preface by the author ed.).
Princeton: Princeton University Press.
Peress, M. (2019). Measuring the research
productivity of political science departments
using Google Scholar. PS: Political Science &
Politics, 52(2), 312–317. https://doi.
org/10.1017/S1049096518001610
Rawls, J. (1999). A theory of justice (2nd ed.).
Cambridge, MA: Belknap Press.
Reynolds, A., Reilly, B., & Ellis, A. (Eds.). (2005).
Electoral system design: The new
international IDEA Handbook. Stockholm:
International IDEA.
Rule, W. (1987). Electoral systems, contextual
factors and women’s opportunity for election
to parliament in twenty-three democracies.
The Western Political Quarterly, 40(3), 477–
498. https://doi.org/10.2307/448386
Santa Fe Institute. (2016, November 9). lawOS:
Regulations as society’s operating system.
Retrieved August 2, 2019, from Santa Fe
Institute website: www.santafe.edu/news-
center/news/lawos-regulations-societys-
operating-system
Schofield, N. (2005). The intellectual
contribution of Condorcet to the founding
of the US Republic 1785–1800. Social Choice
and Welfare, 25(2–3), 303–318. https://doi.
org/10.1007/s00355-005-0005-y
Shugart, M., & Wattenberg, M. P. (2001).
Mixed-member electoral systems: The best
of both worlds? Oxford University Press.
Oxford.
Skyrms, B. (1994). Sex and justice. Journal of
Philosophy, 91(6), 305–320.
Skyrms, B. (2014). Evolution of the social
contract. Cambridge University Press. New
York.
Smith, J. M., & Price, G. R. (1973). The logic of
animal conflict. Nature, 246(5427), 15.
https://doi.org/10.1038/246015a0
Sorkin, R. D., West, R., & Robinson, D. E.
(1998). Group performance depends on the
majority rule. Psychological Science, 9(6),
456–463. https://doi.org/10.1111/1467-
9280.00085
Soudriette, R., & Ellis, A. (2006). A global
snapshot. Journal of Democracy, 17(2), 78–88.
https://doi.org/10.1353/jod.2006.0038
Taagepera, R. (2007). Predicting party sizes:
The logic of simple electoral systems (1st ed.).
Oxford: Oxford University Press.
Taagepera, R. (2008). Making social sciences
more scientific: The need for predictive
models. Oxford: Oxford University Press.
The cooperative human. (2018) Nat Hum
Behav 2, 427–428. Retrieved from https://
doi.org/10.1038/s41562-018-0389-1
Trivers, R. (1971). The evolution of reciprocal
altruism. Quarterly Review of Biology, 46(1).
35–57.
Trivers, R. (2006). Reciprocal altruism: 30 years
later. In P. M. Kappeler & C. P. van Schaik
(Eds.), Cooperation in Primates and Humans:
Mechanisms and Evolution (pp. 67–83).
Springer. New York. https://doi.org/10.1007/
3-540-28277-7_4
 EVOLUTIONARY PSYCHOLOGY AND POLITICAL INSTITUTIONS
Turchin, P. (2013, February 8). The double helix of
inequality and well-being. Retrieved August 3,
2019, from Peter Turchin website: http://
peterturchin.com/cliodynamica/the-double-
helix-of-inequality-and-well-being/
Turchin, P. (2016). Ages of discord: A structural-
demographic analysis of American history.
Chaplin: Beresta Books.
Walker, S. I., & Davies, P. C. W. (2012). The
algorithmic origins of life. Journal of The
Royal Society Interface, 10(79), 20120869–
20120869. https://doi.org/10.1098/rsif.
2012.0869
187
Wilson, D. S., & Gowdy, J. M. (2013). Evolution as
a general theoretical framework for economics
and public policy. Journal of Economic Behavior
& Organization, 90, S3–S10. https://doi.org/
10.1016/j.jebo.2012.12.008
Young, H. P. (1988). Condorcet’s theory of
voting. American Political Science Review,
82(4), 1231–1244. https://doi.org/10.2307/
1961757
Zhou, Y. M. (2011). Synergy, coordination
costs, and diversification choices. Strategic
Management Journal, 32(6), 624–639.
Retrieved from JSTOR.

Evolutionary Psychology
and Crime
INTRODUCTION
As examined throughout this volume, humans
are a highly social species and as such possess
a constellation of adaptations which aid in sur-
vival and reproduction. Among the evolved
strategies employed by highly social species,
those which can be viewed as exploitative are
abundant. Indeed, nature is awash with exam-
ples of evolved strategies that appear to cal-
lously infringe on the desires or choices of
individuals who are the targets of such strate-
gies. To be sure, highly social species rely on
cooperation, empathy, and stable group dynam-
ics, but all these factors can also be exploited
for individual gain as an effective evolved
strategy. When viewed from an evolutionary
perspective, criminal behavior among humans
clearly falls into an exploitative-strategy cate-
gory. However, the field of social science
devoted to the study of criminal and antisocial
behavior, criminology, rarely examines behav-
ior using an evolutionary lens. Instead, almost
all traditional criminological theories and
9
Joseph L. Nedelec
empirical analyses place the etiological respon-
sibility for antisocial behavior solely within the
realm of social factors. Recently, however,
biosocial criminologists have illustrated the
shortcomings of such an approach. The current
chapter overviews the various ways in which an
evolutionary viewpoint can inform our under-
standing of antisocial behavior, crime, and
criminality. Increasingly referred to as evolu-
tionary criminology, the perspective described
in this chapter illustrates how viewing antiso-
cial behavior from an evolutionary standpoint
can explain the most well-established observa-
tions regarding criminality as well as contextu-
alize more recent empirical findings derived
from neuropsychology, behavioral genetics,
and biosocial criminology.
UNDERSTANDING AND USING
EVOLUTIONARY THEORY
Although covered elsewhere in this volume, it
is necessary at the outset of our discussion to
 EVOLUTIONARY PSYCHOLOGY AND CRIME
address why many people within social-­
science disciplines struggle with recognizing
the relevance of an evolutionary point of view.
Perhaps the two most important concepts
related to evolutionary thinking that can help
in this regard are deep time and recognizing
that humans are not immune to the processes
of nature.
Deep time is a geological concept that
informed Charles Darwin in generating his
theory of evolution by natural selection.
Briefly, the concept refers to the immense
amount of time that has passed since the for-
mation of the earth and the amount of time
that has been available for natural processes
such as erosion, movement of the earth’s
crust, and most germane to the current chap-
ter, evolution of biological organisms. Given
that humans typically live for only a handful
of decades, our perceptions of the passage
of time are drastically limited compared to
the age of the earth. Consequently, it is dif-
ficult for most people to understand how the
complexities we observe in nature and our
own behaviors could result from the pro-
cesses of evolution. Other factors such as
religious dogma and cultural characteristics
also impinge on people’s ability to recog-
nize the immensity of time that has passed
and the vast opportunities that have been pro-
vided for the processes of natural and sexual
selection to shape the evolution of species.
When one recognizes and accepts the over-
whelming evidence of deep time, however, it
becomes easier to see how natural processes
could apply to all aspects of species’ lives,
including behavioral traits such as crime and
antisocial conduct.
Given the time frames associated with
the typical human life course, deep time is
often cognitively taxing. However, observ-
ing natural wonders like the Grand Canyon
provides an opportunity to witness the results
of deep time. Recognizing that humans are a
part of nature, however, is often less widely
accepted. Given the complexities and varie-
ties of human culture and the incredible tech-
nological and societal advances that humans
189
have made, along with the wide array of
religious beliefs that have and continue to
­
profess human exceptionalism, it is not sur-
prising that many feel humans are separate
from the natural world – and, therefore,
immune to the processes of natural and sexual
selection that have shaped all life. Acceptance
of human’s place in nature is, and has been
since well before Darwin, a controversial
issue. Nonetheless, the evidence that humans
are a part of nature is as overwhelming as the
evidence for deep time. Empirical facts such
as homology (shared structures across differ-
ent species or taxa), genetic code illustrating
relatedness across animals and plants, and
the considerable fossil record, among other
empirical facts, all point to an inevitable
conclusion: humans are not exceptional in
terms of our place in nature and are the result
of natural processes (i.e., natural and sexual
selection) in the same way as all life on earth.
Once this fact is acknowledged, and in com-
bination with the concept of deep time, it is
thus necessary that assessments of the causes
of almost any aspect of the human condition
must incorporate recognition of the evolution-
ary processes that underpin the human condi-
tion. Such a conclusion was emphasized by
Pierre Teilhard de Chardin who poignantly
noted, “[evolution] is a general postulate to
which all theories, all hypotheses, all sys-
tems must henceforward bow and which they
must satisfy in order to be thinkable and true.
Evolution is a light which illuminates all facts,
a trajectory which all lines of thought must
follow”. (As cited in Dobzhansky, 1973: 129.)
All this information inexorably leads to the
perspective that informs the chapters within
this volume. Briefly, evolutionary psychol-
ogy argues that our brain is the seat of all
human behavior and the construction and
functioning of our brain is driven, in part,
by genetic factors; these genetic factors, in
turn, have been susceptible to the processes
of natural and sexual selection over the vast
eons of evolutionary time. Consequently,
all aspects of the human condition, includ-
ing human behavior, society, and culture,
190 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
can be assessed using an evolutionary lens.
Given that criminal and antisocial behaviors
are an empirical universal across all recorded
history and across every known society, it is
clear that an evolutionary lens can also illu-
minate why such behaviors appear to be an
integral, though often unfortunate, aspect of
the human condition.
OF CRIME, CRIMINALITY, AND
ANTISOCIAL BEHAVIOR
A common reaction in the social sciences to
the suggestion of the utility of an evolution-
ary perspective is that, given behaviors con-
sidered to be criminal are based on codified
laws, crime is a social construct that will vary
from society to society. Thus, a biologically
based perspective such as evolutionary psy-
chology cannot inform the discussion of
crime. The argument, though partially correct
(codified prohibitions on certain behaviors
certainly do change both within and between
societies over time), is incomplete in at least
two ways. First, across all recorded history
and known societies there is considerable
overlap in terms of prohibited behaviors. Few
cultures accept behaviors such as thievery,
murder, assault, or rape (this list is not
exhaustive) without any social rebuke. Even
within highly violent cultures where the
murder of one group’s rivals is seen as a nec-
essary step in the maturity of males, such
behavior is condoned only if it is directed at
an out-group. Consequently, there appears to
be a human constant against the impingement
of the rights and desires of others in terms of
these behaviors (at least in terms of one’s
own in-group). Second, while it is certainly
the case that codified prohibitions (i.e., legal
definitions of what constitutes crime) are
fluid across time and space, the behavioral
proclivities that underlie criminal or antiso-
cial behavior are consistent. These proclivi-
ties are referred to, in general, as criminality,
which is a propensity or inclination to engage
in criminal or antisocial behaviors. Thus,
when biosocial criminologists examine crimi-
nal or antisocial behaviors using an evolution-
ary lens, it is with the concept of criminality
in mind; in other words, the focus is on the
behavior and not necessarily the legality of
the behavior. As an example, criminologists
examine not only antisocial behaviors that are
illegal but also behaviors that are considered
analogous to criminal behavior such as sub-
stance abuse or risky sexual behaviors and the
lifestyles associated or congruent with engag-
ing in antisocial conduct. Thus, recognizing
that antisocial behaviors are acts which vio-
late the interests of one party to the benefit of
another party in contravention of normative
behaviors of the group to which the parties
belong allows for a more nuanced examina-
tion of the etiology of antisocial behavior than
simply relying on legal definitions. Thinking
in this way (i.e., focus on the behavior –
criminality) provides an opportunity to apply
an evolutionary lens. In the sections that
follow, I outline how evolutionary psychology
as a paradigm can help explain three of crimi-
nology’s most commonly observed empirical
patterns in terms of antisocial behavior: the
gender gap in crime, the age-graded distribu-
tion of crime, and the non-random distribu-
tion of criminal behavior.
THE GENDER GAP IN CRIME
Imagine, if you will, a risk factor associated
with criminality and antisocial behavior that
is so pervasive that it replicates across all
known societies and all recorded history.
That risk factor is actually well-known, and
in our species, it is represented by a lone
chromosome: the Y-chromosome. Sex is the
single most consistent predictor of criminal
behavior that has emerged from well over a
century of criminological theorizing and
empirical work. To be sure, not all men
engage in criminal behavior and not all
women refrain from it. However, in the
 EVOLUTIONARY PSYCHOLOGY AND CRIME
statistical sense (i.e., average differences),
there has not existed a society wherein women
engaged in a higher rate of criminal behavior
than men, and this is particularly the case with
violent antisocial behaviors. The robust
observed differences in terms of criminality
between the sexes has been addressed in a
variety of ways in the criminological literature
with almost all explanations focusing on soci-
ological factors (e.g., differential parental
socialization, cultural factors, differential
media exposure, etc.). In each case, these
explanations have proven to be at best incom-
plete and at worst incorrect. A potential reason
for the ineffective explanations is the lack of
recognition that humans are part of the natural
world. Here we see that an evolutionary per-
spective can illuminate potential causal factors
for the observed differences in antisocial
behaviors between men and women.
When one recognizes that humans are a
part of nature, one can then look to nature
for potential analogues of the behavior or
dynamic of interest. Additionally, an evolu-
tionary perspective allows for the applica-
tion of well-known biological theories or
processes. For example, biological explana-
tions of the differences in behavioral reper-
toires between the sexes within many species
are often informed by discussions of invest-
ment (Trivers, 1972). Briefly, any given
organism within a sexually reproducing spe-
cies can allot time and energy to obtaining
mates (mating effort) or caring for offspring
(parental effort or investment). Bioenergetic
resources cannot be allotted to both mating
and parenting simultaneously and, as we
shall see, the evolved strategies of sexes with
regard to investment of resources are often
divergent. Throughout the animal kingdom,
the primary driver of differentiation in bio-
energetic resources is referred to as minimal
parental investment (MPI). MPI refers to the
minimal time and energy costs associated
with producing a viable offspring (i.e., one
that can live long enough to then also repro-
duce). In most primate species (human and
non-human), the MPI for males and females
191
differs considerably. While females in most
primate species have a very high MPI (many
months of gestation and often many years of
offspring care), males have a relatively low
MPI (perhaps as minimal as a few moments
of copulation). Consequently, males can
reproduce at much higher rates and with
shorter intervals than females. While males
have a much higher reproductive ceiling, they
also evince much more variance in terms of
reproductive success – the proportion of
males who never reproduce is much higher
than in females. Additionally, given that
females risk a much higher MPI, they exhibit
mating strategies that lead to what is gener-
ally referred to as choosiness (females assess
males on their reproductive quality to a much
higher degree than males assess females).
The result of this difference in MPI is that
males tend to differentially invest in mating
effort while females tend to invest relatively
more in parenting effort. Along with this dif-
ferential allocation of resources comes a suite
of behavioral strategies which affect the level
and degree of competition both within and
between sexes.
Differential allocation of reproductive
resources driven by MPI and within-sex
variances in reproductive success is associ-
ated with more intense within-sex competi-
tion in the sex with the lower MPI. In less
technical language, the sex with more to lose
(i.e., greater reproductive variance – if you
don’t reproduce, you’re a genetic dead end)
is the sex that will engage in more intense
(i.e., risky, violent, combative) competition
relative to the sex with higher MPI. Thus, the
biological concept of MPI provides the logic
for understanding the wide range of mor-
phological and behavioral traits that males
possess, relative to females, which appear
specifically designed for intense competition.
The same morphological and behavioral traits
related to within-sex competition (i.e., fight-
ing rivals for access to the high MPI sex) are
also employed for between-sex competition
(e.g., subduing the ability of the high MPI sex
to choose among mates).
192 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
While in most primate species males com-
pete in a more intense fashion, there is noth-
ing about maleness per se that inevitably
leads to increased violence and aggression;
likewise, there is nothing about femaleness
per se that inevitably leads to decreased
violence and aggression in within- and
between-sex competition. To illustrate this
argument, one can again observe nature
for instances wherein the sexes converge in
terms of MPI (i.e., greater amount of shared
parental efforts, aka biparental care). Many
species exhibit such patterns (e.g., wolves,
numerous types of birds, beavers, some mon-
keys, among others), and the morphological
and behavioral differences between males
and females are greatly reduced. Perhaps
the greatest evidence for the importance of
MPI can be derived from species where the
males invest more resources to parental effort
relative to females and females have greater
reproductive variance relative to males. This
occurs in varieties of fish and other animals
(though no mammals) and what is observed
is a role reversal – relative to most primates –
in terms of behavioral strategies: the females
compete with more aggression and the males
are generally choosier in terms of selecting
mates.
Armed with an evolutionary explanation of
sex differences in terms of aggressive behav-
ior, one can then see why men and women
in our own species have exhibited and still
do exhibit differences in terms of criminal-
ity or a propensity to engage in antisocial
behaviors. Given that men possess a much
lower MPI than women, it is men who pos-
sess (on average) greater muscle mass,
height, and other morphological traits con-
ducive to intense competition. Additionally,
it is also the reason why men possess a
suite of psychological traits that drive risky
and aggressive behaviors more often and in
typically more intense degrees than women.
Consequently, in possession of the psycho-
logical drive and morphological capaci-
ties to aggressively compete, men are more
often the sex to engage in criminal behaviors
(particularly those which are considered
interpersonal and violent). Hence, one of the
most robust empirical findings of crimino-
logical research: the gender gap in crime.1
AGE-GRADED DISTRIBUTION OF
CRIME
In addition to the gender gap in crime,
researchers of human antisocial behavior
have long observed that criminal activity, in
general, tends to rise with the onset of ado-
lescence, peak near the end of adolescence,
and then abruptly plummet in young adult-
hood. This empirical pattern has been dubbed
the age-crime curve and, like the gender gap
in crime, has appeared relatively consistent
across time and space.2 Numerous scholars
have put forth theoretical explanations for
this patterned empirical observation, the
most well-known among criminologists
being Laub and Sampson’s (1993) age-
graded theory of crime and Moffitt’s (1993)
dual taxonomic theory of crime.
Briefly, Laub and Sampson’s (1993) theory
asserts that informal social control through
an individual’s bond to society affects the
propensity to engage in criminal behav-
ior over the life course. The theory places
great weight on the bonds within an indi-
vidual’s family during development as well
as attachment to school, employment, and
other structural aspects of society. Overall,
Laub and Sampson argue that those youth
with weak social bonds are more likely to
engage in antisocial behaviors during adoles-
cence and that such behavior in turn leads to
an increased likelihood of criminal behavior
in adulthood. Further, they argue that with-
out social bonding or attachment to institu-
tions such as employment, military service,
and marriage, continued criminal behavior
during adulthood is probable. Finally, they
argue that desistence from criminal behavior
is largely due to obtaining attachment to one
or more of these social institutions.
 EVOLUTIONARY PSYCHOLOGY AND CRIME
Moffitt’s (1993) dual taxonomic theory also
recognizes the importance of a life-course
approach and contains a multitude of hypothe-
ses regarding the shape of the age-crime curve.
In brief, Moffitt argued that antisocial behav-
ioral patterns illustrate two distinct groups of
offenders: adolescent-limited offenders (AL)
and life-course persistent offenders (LCP).3
As suggested by the label, those exhibiting
AL patterns of offending engage in crimi-
nal behavior that is limited to the period of
adolescence, that was not preceded by anti-
social behavior in childhood, and does not
tend to continue into adulthood. Additionally,
the type of offending in which AL offend-
ers engage is typically relatively minor and
rarely results in serious contact with the
criminal justice system (i.e., long-term insti-
tutionalization). LCP offenders, however,
exhibit behavioral patterns illustrating a life-
time of antisocial behavior from childhood
through adulthood. Additionally, relative to
AL offenders, the type of criminal activity in
which LCP offenders engage is often severe
and does typically lead to serious and con-
sistent interaction with the criminal justice
system (throughout the life course).
In terms of etiological factors, Moffitt
(1993) argued that the behavioral pattern of
LCP offenders is a result of an unfortunate
mix of neuropsychological deficits and del-
eterious rearing environments. She argued
that LCP offending patterns are so serious
in nature and consistency that exceedingly
damaging circumstances such as abnormal
neuropsychological functioning and abusive
or otherwise damaging developmental envi-
ronments were required. However, given that
Moffitt argued that AL offending was norma-
tive (i.e., an expected pattern of development
in modern societies), her explanation required
normative processes typically experienced
by most youth. Her hypotheses regarding AL
offending centered on two components. The
first component is represented by the differ-
ence between the biological maturity experi-
enced by youth in adolescence (after puberty
an individual is, more or less, biologically an
193
adult) and the social limitations that remain
in place in modern societies (adolescence
is a period of relaxed limitations relative
to childhood, but it still consists of wide-­
ranging restrictions on a variety of aspects of
life). The difference between one’s biologi-
cal maturity and one’s perceptions of social
freedom/limitation was termed the maturity
gap. Moffitt argued that the greater the expe-
rienced maturity gap, the greater the frustra-
tion experienced by the youth and therefore
the greater the need to address the result-
ing frustration. The second component of
Moffitt’s explanation of AL offending pro-
vides the mechanism through which youth
were said to then deal with the frustration
resulting from an experienced maturity gap.
Her argument indicated that AL individuals
could recognize/observe the relative social
freedom exhibited by those who are engaged
in LCP offending and lifestyle patterns.
Observing the socially uninhibited lifestyle
of LCP offenders could then lead to a process
of behavioral mimicry in order to gain simi-
lar social freedoms (termed social mimicry).
Moffitt thus argued that AL offending was a
result of experiencing a maturity gap during
adolescence and engaging in social mimicry
such that the behavioral patterns of LCP
offenders are followed (although to a less
serious degree) by non-LCP youth (i.e., AL
offenders). Further, Moffitt argued that the
desistance from criminal behavior observed
during early adulthood by AL offenders was
a result of the reduced effect of the matu-
rity gap (i.e., the social limitations placed
on adolescents become much less intense or
pervasive as they age into early adulthood) –
thus, with the accumulation of greater social
freedoms throughout many aspects of their
lives, those in the AL offending group no
longer experience the frustrations associated
with social limitations which eliminates the
need to mimic the behavioral patterns of LCP
offenders.
Both the age-graded and dual taxonomic
theories of crime have received substantial
attention in the criminological literature and
194 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
researchers continue to test the hypotheses
derived from both theories. For the purposes
of this chapter, however, it is important to
note that both theories are proximal theo-
ries of criminal behavior (as exhibited in
the age-crime curve). As discussed through-
out this volume, proximal explanations of
behavior refer to those arguments that focus
on factors related primarily to ontogeny (i.e.,
development within an individual’s lifespan),
whereas evolutionary arguments provide ulti-
mate explanations of behavior, which focus
on phylogeny (development over genera-
tions rather than within a single lifespan) and
adaptive function (i.e., what adaptive prob-
lem is addressed by the behavior in ques-
tion?). For example, while it may be the case
that informal social bonds affect crime over
the life course or that the frustration associ-
ated with experiencing the maturity gap leads
to increased antisocial behavior in adoles-
cence, we are still left with the question as
to why these processes have the (potential)
effect that is purported. Why should social
bonds experienced during development and
in adulthood affect behavioral patterns? Why
should biological maturity combined with
limitations on social freedoms lead to frus-
tration among youth? Why would aggressive
or violent behavior be something in which
youth engage to deal with the frustration?
Fortunately, an evolutionary perspective can
provide the ultimate reasons/answers to such
questions.
As with the discussion of the gender gap
in crime, an evolutionary explanation of the
age-crime curve centers on variance in repro-
ductive effort between men and women. A
few examples of evolutionary explanations
for the age-crime curve exist in the literature
(e.g., Quinsey et al., 2004) but they all center
on what Wilson and Daly (1985) termed the
young male syndrome. Briefly, Wilson and
Daly argue that given the immense costs of
reproductive failure (i.e., genetic dead end)
and the substantial variance in reproductive
fitness among men, there will be intense com-
petition for any resources that help increase
reproductive ­fitness. In this zero-sum social
contest, higher-ranking men tend to obtain
more mates and increase their reproductive fit-
ness, whereas lower-ranking men have fewer
or no mates. Given that social rank is a vital
component to reproductive fitness for men,
there is a heightened psychological awareness
among men to threats to status or reputation
(often referred to as honor). Threats to social
status or reputation are, in essence, threats to
social rank and, thus, threats to reproductive
potential. Consequently, intense, aggressive,
and often violent competition to maintain or
advance rank can occur when such threats
arise. Further, within a polygymous breed-
ing system wherein some males reproduce
much more than most males there are numer-
ous reproductive benefits to intense (aggres-
sive, risky) competition. Such behavior can
serve to protect or gain status, discourage or
eliminate rival males in a competitive breed-
ing environment, allow for the acquisition of
resources to be used to woo females, engage
in and protect from mate poaching, and pro-
tect already acquired resources and mates
(including aggressive mate guarding).
Wilson and Daly (1985) provide empirical
evidence supporting these assertions based
on data from over 500 homicide cases in
Detroit in the early 1970s. They illustrate that
not only were the majority of both offend-
ers and victims in homicide cases men, but
offenders and victims were almost identical
in terms of being unemployed, unmarried,
and younger (teen years to mid 20s). Analysis
of the cases also indicated that the most com-
mon type of homicide was the result of social
conflict or what criminologists would refer to
as the escalation of a trivial altercation (the
majority of which were primarily in retalia-
tion for a previous loss of face in the pres-
ence of social peers). Summarizing their
observations regarding lethal conflict, they
note, ‘many, perhaps most, homicides con-
cern status competition’ (Wilson and Daly,
1985: 59). Wilson and Daly also illustrated
that other behaviors that carry a substantive
threat of physical harm (e.g., risky and/or
 EVOLUTIONARY PSYCHOLOGY AND CRIME
aggressive driving) are likely the result of a
generalized willingness to engage in compet-
itive risk taking relevant to social rank and,
thus, linked to male fitness. The observations
and hypotheses put forth by Wilson and Daly
have been further supported using data out-
side of Detroit and for different time periods
(see Daly, 2016; Daly and Wilson, 2017).
Although Wilson and Daly (1985) outline
why engagement in criminal conduct and
other risky behaviors increases during the
adolescent years, their discussion does not
explicitly focus on the entirety of the age-
crime curve. In their theoretical piece on male
criminality over the life course, Kanazawa
and Still (2000) echo Wilson and Daly’s
young male syndrome hypothesis but also
directly address the shape of the age-crime
curve. Overall, Kanazawa and Still indicate
that intense competition (manifested as anti-
social, aggressive, and other risky behaviors)
occurs when the reproductive benefits of such
behavior are maximized over the life course.
Thus, violent competition among males does
not occur earlier in life (i.e., prior to puberty)
primarily because there are no reproductive
benefits to engaging in violence, theft, or
murder – the pre-pubescent male is unable to
translate a competitive edge into reproductive
success. However, after puberty the repro-
ductive benefits sky-rocket and so too does
the resulting behavior (thus, the peak in ado-
lescence of the age-crime curve). As noted
above, however, the age-crime curve drops
precipitously in early adulthood. Kanazawa
and Still argued that the ultimate reason
for this drop is associated with the costs of
continued intense competition. The authors
argue that risky strategies are employed to
gain sexual access to mates and, in general,
secure reproductive success (or at least the
opportunity for reproductive success) and at
the arrival of offspring (i.e., reproductive suc-
cess) less risky behavior is employed to mini-
mize the costs to the reproductive success
of the male. Thus, the age-crime curve (for
males) is a manifestation of a two-pronged
strategy selected over evolutionary time
195
wherein males engage in intense competition
in order to secure reproductive success, but
once this is secured tend to disengage from
risky behavioral strategies given the potential
costs to the obtained reproductive success.
The ultimate explanations put forth by
Wilson and Daly (1985) and Kanazawa
and Still (2000) are supported by proximal
explanations derived from psychophysi-
ological and criminological research. For
example, the hormone testosterone – which
has been associated primarily with competi-
tive behaviors – increases dramatically in
males during puberty but has been observed
to drop substantially at the onset of marriage
and the arrival of offspring (Beaver, 2009).
Additionally, some criminological research
has illustrated a calming effect in males (in
terms of engagement in criminal behavior)
who begin families, though this research is
often confounded by issues of temporal order
and a lack of genetically informed analyses
(Barnes et al., 2014).
Finally, we see that these evolutionary
explanations of the age-crime curve pro-
vide some potential answers to our earlier
questions derived from the discussion of the
age-graded and dual taxonomic theories of
crime. For example, informal social bonds
such as employment, military service, and
marriage could affect male criminality as these
are resources which directly affect or relate
to reproductive success. Given that crimi-
nal behavior, especially violent or aggressive
behavior, represents a threat to these resources
there is a corresponding reduction in the level
of criminal behavior (on average) that is con-
gruent with the acquisition of these resources.
Additionally, experiencing the maturity gap
results in increased frustration in youth because
their biological maturity – driven by eons of
evolutionary processes – motivates them to
engage in intra- and inter-sexual competition
to obtain mates. However, the wide-ranging
social limitations placed on adolescents pre-
vent or at least minimize opportunities for such
behaviors. Thus, social rebellion through minor
delinquency could result (not only to rebel
196 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
against the social restraints but to attempt to
secure status and rank within a peer network).
Overall, the discussion illustrates what all evo-
lutionary psychologists argue: both proximal
and ultimate explanations are required in order
to fully understand observed behavioral pat-
terns such as the age-crime curve.
NON-RANDOM DISTRIBUTION OF
CRIMINAL BEHAVIOR
The final consistent criminological empirical
observation that we will address in the current
chapter is the non-random distribution of
criminal behavior. In addition to the gender
gap and age-crime curve, long-standing and
cross-cultural patterns in terms of where
criminal conduct tends to occur geospatially
and the typical dynamics of offender–victim
relationships have been observed. Entire sub-
fields within criminology, for example, have
developed which center specifically on these
observations. For example, the Chicago
school of criminology, most readily exempli-
fied by the work of Shaw and McKay (1942),
focused on the differential patterns of criminal
behaviors across different areas (or zones)
within a city. The authors argued that the dif-
ferential offending patterns were a result of
variance in the structural conditions (e.g.,
economic status, ethnic heterogeneity, and
residential mobility) found in the zones.
Further, socioeconomic status and aspects of
neighborhood cohesion (sometimes referred
to as collective efficacy; Sampson et al., 1997)
have long been assessed as key causal variables
in the etiology of crime and criminality.
Additionally, criminologists and other social
scientists have also been examining the nature
of victim–offender relationships for well over
a century. Consequently, much is known
about the ways in which criminal behavior is
distributed in terms of geospatial location
within cities and towns as well as how offend-
ers and victims are (or are not) known or con-
nected. Overall, in both cases there is a
non-random distribution of criminal conduct
although the pattern of the conduct typically
depends upon the specific type of crime. The
remaining discussion within this section will
illustrate how an evolutionary perspective can
help explain these non-random patterns.
Non-Random Clustering of
Criminality and Other Risky
Behaviors in Locales
As noted, criminologists have observed that
criminal conduct tends to be over-represented
or differentially concentrated within certain
areas of a city or town. Typically, the areas of
concentration are considered to represent
highly unstable environments characterized
by low average economic status and relatively
low social cohesion. Social scientists have
tended to point to these characteristics as the
causal factors in the accompanying concentra-
tion of criminal conduct while others have
noted that the criminal behavior exhibited in
such areas is a result of a sub-culture of reck-
lessness resulting from minimal opportunity
for social advancement. However, as dis-
cussed in the prior section these types of
explanations are incomplete – why would
criminal behavior, especially violent behavior,
result from reduced social and economic
opportunity? Why would criminal behavior
exhibit concentrations among communities
with a relative lack of social cohesion or sta-
bility? In their analysis of 77 different neigh-
borhoods in Chicago using data from 1988 to
1993, Wilson and Daly (1997) illustrated that
an evolutionary perspective can help to
address such questions (see also Daly, 2016).
Briefly, Wilson and Daly (1997) examined
the differential life expectancy for men and
women across various neighborhoods, as well
as homicide rates, birth rates, and measures of
socioeconomic status (i.e., household income
and an income inequality index). The key inde-
pendent variable, life expectancy, was a meas-
ure of the expected average duration of life,
in years, for an individual based on a variety
 EVOLUTIONARY PSYCHOLOGY AND CRIME
of vital statistics and population data present
at the time of the individual’s birth. In their
analyses Wilson and Daly empirically
assessed three specific hypotheses: (1) homi-
cide rates are a function of local life expectancy;
(2) economic inequality accounts for variance
in homicide rates beyond that attributed to
local life expectancy; and (3) local life expec-
tancy will impact reproduction (birth rates)
across neighborhoods. Overall, the research-
ers argued that life expectancy provides a vital
external cue to inhabitants of neighborhoods
such that their unconscious behavioral and
reproductive strategies can be adjusted based
on an assessment of probable lifespan. Thus,
rather than representing a potential pathologi-
cal reaction to social conditions, high-crime
areas may be a function of a rational calcu-
lation (though one that has been honed over
evolutionary time) based on environmental
cues. The analyses testing these ideas revealed
a number of illuminating findings.
First, life expectancy at birth was strongly
associated with homicide rates across neigh-
borhoods for both men and women. In neigh-
borhoods with a lower life expectancy at birth
a much higher homicide rate was observed.
The magnitudes of the bivariate associations
were very strong and statistically signifi-
cant for both men (r = −0.88, p < .0001) and
women (r = −0.83, p < .0001). Importantly,
these associations held in multivariate mod-
els wherein measures for household income
and income inequality were introduced.
Second, comparisons between the 10
neighborhoods with the longest life expec-
tancy and the 10 neighborhoods with the
shortest life expectancy revealed some drastic
differences in terms of homicide rates across
age categories. For example, in the long life
expectancy neighborhoods the homicide rate
(deaths per 100,000 per year) went from vir-
tually zero for those males aged five to 14
years to about 20/100,000 for those aged
15 to 24, to a peak of about 25/100,000 for
those aged 25 to 34 before dropping to near
zero for the remaining age groups. However,
in the short life expectancy neighborhoods
197
the homicide rate skyrocketed from about
5/100,000 in the five to 14 years age group to
over 300 deaths per 100,000 for the males in
the 15 to 24 years age group. While the hom-
icide rate for males in these neighborhoods
decreased over time, the rates were still astro-
nomically higher than those observed in the
long life expectancy neighborhoods (25 to
34 years: about 250/100,000; 35 to 44 years:
about 175/100,000; 45 to 54 years: about
75/100,000; 55 to 64 years: about 60/100,000;
65 to 74 years: about 40/100,000; and 75
years or older: about 55/100,000). While the
homicide rates for females in the short life
expectancy neighborhoods were much lower
than for males, the overall pattern across the
age groups was similar and, in some cases,
exceeded the homicide rates for males in the
long life expectancy neighborhoods.
Finally, in terms of reproductive behaviors
Wilson and Daly (1997) again compared the
10 neighborhoods with the longest life expec-
tancy to the 10 neighborhoods with the short-
est life expectancy across seven different
age categories. The differences in birth rates
during the teen years and early adulthood
between the neighborhoods was substantial.
Figure 9.1 illustrates the stark differences.
As illustrated, the birth rate for women in
the short life expectancy neighborhoods was
over four times higher than the long life expec-
tancy neighborhoods for the 15 to 19 years
age group, and two-and-a-half times greater
in the 20 to 24 years age group. Additionally,
although the difference is still evident in the
25 to 29 years age group, the birth rates for the
different neighborhoods become almost iden-
tical in the remaining age categories (i.e., 30
years and above). Based on the results of their
analyses, Wilson and Daly concluded:
[t]he data presented here indicate that people
behave as if they have adjusted their rates of
future discounting and risk acceptance thresholds
in relation to local life expectancy, and that they
do so in the non-violent domain of reproductive
decision making as well as in the potentially violent
domain of social competition. (Wilson and Daly,
1997: 1273)
198 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

Figure 9.1  Age-specific birth rates (per 1,000 women per year) in the 10 neighborhoods
with the longest life expectancy compared to the 10 neighborhoods with the shortest life
expectancy
Source: Data derived from table 3 in Wilson and Daly (1997).

The analyses and conclusion presented by
Wilson and Daly (1997) illustrate how an
evolutionary lens can help explain social
phenomena that have hitherto only been
addressed employing sociologically based
theories. Furthermore, they illustrate that the
behavioral repertoire exhibited by individu-
als presented with certain environmental
cues may not be pathological or reckless, but
rather a function of an unconscious calculus
resulting from eons of evolutionary pro-
cesses. The next example provides a similar
illustration in terms of observed patterns of
victim–offender characteristics.
Non-Random Victim–Offender
Characteristics
Depending on the criminal behavior of inter-
est, criminologists have observed that char-
acteristics are consistently represented in
terms of victims and offenders. In general,
both victims and offenders tend to be in their
adolescence or early adulthood. This obser-
vation is not entirely surprising given the
age-crime curve, which manifests as a result
of the intense mating competition experi-
enced during that period in the life course.
However, examinations of specific criminal
behaviors have illustrated other patterns in
terms of characteristics. For example,
researchers estimate that in North America
on average approximately 65% of all mur-
ders involve a male offender and a male
victim, about 20% involve a male offender
and a female victim, about 10% involve a
female offender and a male victim, and less
than 5% involve a female offender and a
female victim (Buss, 2005; Daly and Wilson,
2017). Additionally, the risk of being a homi-
cide victim (in general) increases considera-
bly during late adolescence, peaks in the
early 20s, and then drops substantially over
adulthood. The overall pattern of homicide
and other aggressive behaviors in terms of
 EVOLUTIONARY PSYCHOLOGY AND CRIME
offender–victim characteristics has been
addressed by a wide range of criminological
inquiries. Most of the examinations, how-
ever, have focused solely on proximate fac-
tors such as socialization practices or even
media consumption to account for the
observed patterns. As noted earlier, a sub-
stantial problem for such explanations is that
the patterns observed in terms of offender–
victim characteristics are relatively consist-
ent across time and space. Thus, explanations
which rely on variance in societal factors
such as socialization practices are doomed to
be incomplete. Fortunately, researchers
employing an evolutionary lens have pro-
vided potential explanations for the observed
patterns of offender characteristics exhibited
in several crimes, including homicide.
In their book entitled Homicide:
Foundations of human behavior, Daly and
Wilson (2017) expand upon their 1985 paper
and provide a thorough examination of how
an evolutionary viewpoint can help explain
the characteristics of offenders and of vic-
tims for various types of homicides. Their
discussion centers on intrasexual competition
among young males who are psychologi-
cally attuned to threats to status and again
apply the young male syndrome logic to
socially competitive risky behaviors. Daly
and Wilson provide a wellspring of empirical
analyses from multiple countries to support
their claims, and their coverage of the topic
is thorough. However, as valuable as Daly
and Wilson’s book is – and it is certainly a
definitive discussion of how an evolutionary
viewpoint can be applied to murder – we will
focus our discussion in this section instead on
a book by David Buss (2005).
Buss’s (2005) book, entitled The Murderer
Next Door: Why the Mind is Designed to
Kill, presents the argument that homicidal
behavioral patterns may have been selected
for over evolutionary time as a potential
strategy for dealing with a variety of adaptive
problems. The claim that killing is produced
by a specific psychological adaptation is a
controversial one and discussing the merits
199
of the argument is beyond the scope of the
current chapter. However, the data on which
Buss based his conclusion and the evolution-
ary arguments are worth considering herein.
In addition to examining a large amount of
behavioral data similar to that presented in
Daly and Wilson’s (2017) book (i.e., offi-
cial criminal justice records), Buss and his
team also collected data on homicidal idea-
tions (fantasies) from a sample of over 5,000
individuals across a number of age categories
from multiple countries. The respondents in
the study were asked if they had ever thought
of killing someone, who it was (in terms of
the relationship to the respondent), the man-
ner in which they thought of killing the tar-
get, what prevented them from going through
with the killing, and what could have poten-
tially led them to actually kill (i.e., push them
over the edge). The respondents were also
asked if they ever thought someone might kill
them and were also presented similar follow-
up questions (i.e., who they thought may have
wanted to kill them, how they might have
been killed, what the respondent did to pre-
vent being killed, what prevented them from
being killed, and what would have pushed the
person over the edge to kill the respondent).
The results of the survey revealed several
illuminating patterns that aligned with evo-
lutionary theory (only a handful of which are
included here).
First, a considerable majority of the
respondents in the sample indicated that they
had given thought to killing. Buss and his
team of researchers observed that 91% of the
men in the sample and 84% of the women
reported at least one vivid fantasy about kill-
ing someone. Buss argued that this finding
supports the argument that over evolution-
ary time murder may have been an effective
strategy to employ when faced with a serious
adaptive problem.
Second, both men and women in the study
exhibited consistent yet distinct patterns of
homicidal ideations. While both sexes reported
homicidal fantasies related to sexual rivalries
(e.g., killing the new sexual/romantic partner
200 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
or a former partner), men tended to focus on
issues centered on mate retention (i.e., infidel-
ity, actual or perceived, by their current or for-
mer partner) whereas women tended to focus
on threats by other women to their own mate
quality (e.g., responding to rumors about their
own sexual reputation) and on responding to
perceived or actual abusive threats by current
or former partners. Buss notes that given the
substantial threat to reproductive success in
these circumstances intense counter-measures,
such as violent action, were likely selected for
over evolutionary time.
Third, related to the general patterns of
men and women’s homicidal ideations the
researchers also noted that men generally
reported homicidal ideations focused on sex-
ual (rather than emotional) infidelity, whereas
when women mentioned infidelity of a cur-
rent or former partner in a homicidal fantasy
it was more often related to emotional (rather
than sexual) infidelity. The general differ-
ences observed in the reported homicidal fan-
tasies in this regard also belie the differential
threats to reproductive success for the sexes.
As outlined above, given that women are the
high-MPI sex in our species sexual access is
a resource that is highly contested by men.
Thus, any aspects of the social environment
which affects the likelihood of securing such
a resource will be met with severe reaction.
Likewise, while women are typically the
choosier sex in terms of sexual access they
are, as a result of being the high-MPI sex,
more burdened by potential and actual off-
spring than are men. Thus, over evolutionary
time psychological modules guiding wom-
en’s mate choice have been tuned to cues in
potential mates that indicate a willingness to
invest in the long term (i.e., help to raise any
future offspring). One such cue is the extent
to which a mate professes and exhibits emo-
tional attachment. Consequently, Buss and
his team argue, many of the women in the
study centered their homicidal fantasies on
situations wherein the respondent actually
had, or perceived an experience of, emotional
infidelity.
Fourth, in terms of the thoughts related
to being a victim of homicide, Buss and his
team also noted some general patterns that
were consistent within and between sexes.
For instance, both men and women consist-
ently indicated that they thought they would
be the victim of murder given that they had
engaged in or came close to what evolutionary
psychologists generally call mate poaching.
In essence, the process refers to a sexual or
romantic partnership with another mate who
is already involved in a relationship. As noted
above, being the actual or perceived victim of
mate poaching was consistently evident in the
fantasies of those who reported wanting to kill.
Additionally, across most known societies and
recorded time periods such behavior is associ-
ated with enraged emotion and what is typi-
cally termed irrational behavior (though from
an evolutionary point of view, acting to mini-
mize threats to one’s reproductive status may
actually be rational; see Daly, 2016). Indeed,
the legal codes of many societies include pro-
visions for reduced punishment or even culpa-
bility in cases where a spouse murders a mate
poacher. Thus, all parties involved in the mate-
poaching situation are aware of the potential
risks and the respondents who reported feared
homicidal victimization in Buss’s study cer-
tainly echoed such awareness.
In terms of some of the general differences
between the sexes regarding the reports of
being a potential murder victim, the primary
variance mirrored that observed in the fan-
tasies associated with committing a homi-
cide. For example, men generally reported a
fear of homicidal victimization that resulted
from not only mate poaching but also from
threats to other men regarding the target’s
(i.e., the potential homicidal male) social
rank, status, or worthiness as a sexual partner.
Additionally, women generally reported a
fear of victimization resulting from engaging
in verbal denigration of the sexual reputation
or physical appearance of other female rivals.
Overall, the pattern illustrated in the idea-
tions about homicidal victimization reflected
a keen recognition of the type of social
 EVOLUTIONARY PSYCHOLOGY AND CRIME
circumstances that typically could drive oth-
ers to kill. In line with evolutionary theory,
Buss noted that this dynamic is a result of
co-evolution of intense strategies related
to maintaining or increasing one’s value in
the highly competitive sexual reproduction
market. Further, the findings are manifesta-
tions of the evolution of strategies related to
prevent being a victim of such competition
(and therefore also maintaining or increasing
one’s mate value). Biologists and evolution-
ary psychologists refer to this process as the
Red Queen hypothesis, and it is discussed in
detail elsewhere in this volume.
Overall, the data presented in Homicide
and The Murderer Next Door align with the
historical and contemporary data studied by
criminologists and other social scientists. The
patterns observed in terms of the crime-specific
non-random distributions of victim–offender
characteristics and relationships occur across
these data. The differences, however, arise in
the explanations that have been proffered to
account for these observations. Whereas social
scientists typically lay the blame on processes
such as socialization, culture, and media expo-
sure, evolutionary psychologists illustrate how
such observations may actually be the result
of our evolutionary heritage. Data presented
by Buss and his research team indicate that
the targets of offenses such as homicide may
(in general) be particular and such particular-
ity is due to the specific threat to reproductive
success or survival (the key aspects of evolu-
tionary processes) represented by the target.
Understanding the victim–offender associa-
tions so often observed in criminological data in
terms of ultimate causes provides an opportu-
nity for greater clarity of etiology and therefore
potential to increase our ability to reduce harm.
CONCLUSION
The current chapter presented an overview of
some of the ways in which an evolutionary
view can be applied to crime, criminality,
201
and antisocial behavior. As noted in the intro-
duction, given the social nature of our spe-
cies, the exploitation of others for personal
gain is an indelible and likely inevitable
characteristic of the human condition. The
inevitability of the characteristic, however,
does not mean that our species need accept,
condone, or encourage antisocial behaviors of
any kind. As noted throughout this volume,
nothing about the explanation of a behavior
should be seen as a moral stance on the
behavior. Additionally, just because antiso-
cial, aggressive, and/or criminal behavior is in
part due to the natural processes associated
with evolution it does not mean that the
behavior is justified or in any way excused by
the knowledge of those processes (to think
otherwise would be to commit the naturalistic
fallacy). Rather, the stance taken in this chap-
ter and elsewhere is that the best opportunity
to reduce harm associated with criminal
behavior must be derived from our best
efforts to understand the underlying processes
of criminality. While the social sciences have
provided a wide variety of potentially useful
proximal hypotheses in this regard, the appli-
cation of an evolutionary perspective to crim-
inal behavior provides the ultimate, and
therefore likely most useful, understanding.
As biosocial criminology advances within
and beyond the discipline of criminology it is
likely that our ability to address the harms
associated with criminality will be enhanced.
Employing an evolutionary lens will be a
crucial component of that journey.
Notes
1  At the risk of being repetitive, it is key to note
here that an evolutionary perspective does not
dismiss the importance of cross-cultural diversity
in terms of observed crime rates and potential
additional etiological factors. Indeed, an evolu-
tionary perspective is inherently biosocial such
that it emphasizes the interactive processes
between the inherited genetic architecture of
the brain and the developmental environment to
which an individual is exposed.
202 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
2  Deviations across a variety of societies have been
noted, but some of these patterns of deviation
are due to exceedingly rare socio-political events
(e.g., aftermath of a world war and nuclear
attack; Hiraiwa-Hasegawa, 2005).
3  Moffitt also suggested a third group, abstainers,
who appear to not engage in any offending over
the life course. While of much theoretical interest,
our discussion will be limited to the two offend-
ing groups emphasized in her dual taxonomy.
REFERENCES
Barnes, J. C., Wright, J. P., Boutwell, B. B.,
Schwartz, J. A., Connolly, E. J., Nedelec, J. L.,
& Beaver, K. M. (2014). Demonstrating the
validity of twin research in criminology.
Criminology, 52, 588–626.
Beaver, K. M. (2009). Biosocial criminology: A
primer. Dubuque: Kendall Hunt.
Buss, D. M. (2005). The murderer next door:
Why the mind is designed to kill. New York:
Penguin.
Daly, M. (2016). Killing the competition:
Economic inequality and homicide. New
York: Transaction Publishers.
Daly, M., & Wilson, M. (2017). Homicide:
Foundations of human behavior. New York:
Routledge.
Dobzhansky, T. (1973). Nothing in biology
makes sense except in the light of evolution.
The American Biology Teacher, 35, 125–129.
Hiraiwa-Hasegawa, M. (2005). Homicide by
men in Japan, and its relationship to age,
resources and risk taking. Evolution and
Human Behavior, 26, 332–343.
Kanazawa, S., & Still, M. C. (2000). Why men
commit crimes (and why they desist).
Sociological Theory, 18, 434–447.
Laub, J. H., & Sampson, R. J. (1993). Turning
points in the life course: Why change matters
to the study of crime. Criminology, 31,
301–325.
Moffitt, T. E. (1993). A developmental
taxonomy. Psychological Review, 100,
674–701.
Quinsey, V. L., Skilling, T. A., Lalumiere, M. L., &
Craig, W. M. (2004). Juvenile delinquency:
Understanding the origins of individual
differences. Washington, DC: American
Psychological Association.
Sampson, R. J., Raudenbush, S. W., & Earls, F.
(1997). Neighborhoods and violent crime: A
multilevel study of collective efficacy. Science,
277, 918–924.
Shaw, C. R., & McKay, H. D. (1942). Juvenile
delinquency and urban areas. Chicago, IL:
University of Chicago Press.
Trivers, R. L. (1972). Parental investment and
sexual selection. In B. Campbell (Ed.), Sexual
selection and the descent of man, 1871–1971
(pp. 136–179). Chicago, IL: Aldine.
Wilson, M., & Daly, M. (1985). Competitiveness,
risk taking, and violence: The young male
syndrome. Ethology and Sociobiology, 6,
59–73.
Wilson, M., & Daly, M. (1997). Life expectancy,
economic inequality, homicide, and repro-
ductive timing in Chicago neighbourhoods.
BMJ, 314, 1271–1274.

Evolutionary Psychology and
Policing: The Balance Between
Aggression and Restraint
INTRODUCTION
Police professionalism and use of force are
critical topics of public interest in the 21st
century. Perhaps more than ever in the his-
tory of US policing, the public are demand-
ing accountability and visibility of police
behavior. Some researchers indicate that this
intense microscope of scrutiny has led to
decreased legitimacy and public faith in
police, going so far as to label it a “legiti-
macy crisis” (Gest, 2016; James et al., 2016).
Following high-profile shootings of unarmed
African American men in recent years, start-
ing with Michael Brown in Ferguson,
Missouri, public trust in police dropped sig-
nificantly, equaling the rates observed in the
years following the Rodney King trials
(Jones, 2015). Minority citizens reported less
trust in the police than white citizens (Peck,
2015). Within any social system a certain
degree of give and take is necessary, and
historically the police have had some diffi-
culty with yielding authority – for example,
10
Lois James
enforcement of ‘stop and frisk’ practices,
contributing to racial injustice and anti-
police sentiment (White and Fradella, 2016).
In the ‘post-Ferguson’ era, the policing pro-
fession is faced with nationwide calls for
reform, and an understanding of how this
professional group has evolved is essential
for guiding its path forward (President’s Task
Force on 21st Century Policing, 2015).
The function of the police is tied to their
granted authority to use force to ensure safety
and order. Renowned sociology and policing
scholar Egon Bittner (1970) identified the
role of the police as the legitimate authority
to exercise force. This is challenging because
the exercising of this authority by the police
frequently stirs accusations of brutality and
racism, civil unrest, demands that officers
be criminally punished, and at times mass
violence and rioting. The goal of this chap-
ter is to explore the police function and these
contradictory social realities using evolution-
ary psychology. In order to maintain order
and serve the citizenry the police must be
204 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
both aggressive and restrained. Maintaining
a balance between aggression and restraint
is required to promote and uphold social
order. As society evolves, so does this bal-
ance (whereby the scale can tip more towards
aggression or restraint depending on societal
demands). Our society today expects greater
restraint on the part of police, while police
culture continues to promote aggressive
authority. Aligning public and police expec-
tations of the appropriate balance between
aggression and restraint could promote legiti-
macy in all aspects of police work.
THE EVOLUTION OF AGGRESSION
AND RESTRAINT
Evolution is the process of change in all
forms of life over generations. Each genera-
tion inherits traits, through genes, from their
parents. If this new trait is heritable, and
contributes to greater reproductive success of
the organism relative to individuals without
the trait, it will be passed on to the next gen-
eration and accumulate in the population
(Buss and Shackelford, 1997). New traits
that do not help the organism survive long
enough to reproduce will become rare or
disappear. This process of natural selection
or ‘survival of the fittest’ has guided the evo-
lution of aggressive and restrained behaviors
over time. Although in social animals such as
humans, aggression is not typically an indis-
criminate strategy (Savage and Kanazawa,
2004), context-specific aggression is a natu-
rally occurring, prevalent phenomenon
(Neuberg et al., 2010). Restraint is the action
that occurs when the ‘means to an end’ is
reached, for example when the threat of an
opponent is neutralized. It can also occur if
one decides that the risks of employing
aggression as a strategy are too great. For the
most part, aggression and restraint are tightly
linked.
Buss and Shackelford (1997) identified
seven social problems for which aggression
has evolved over generations as a ­beneficial
response. These include the protection and
acquisition of necessary resources, self-
defense against attack, inflicting costs on
same-sex rivals who are vying for the same
resources, gaining and maintaining power
and dominance, deterring potential oppo-
nents from future attacks, ensuring the sexual
fidelity of a partner, and reducing resource
investment in unrelated children. For exam-
ple, in a pack of wolves, the dominant mem-
ber cannot show weakness in the face of a
physical challenge or his dominant status
will be questioned and potentially overturned
(Millan, 2006). He must assert his dominance
through aggression, or submit to a new leader
(restraint). In this way, aggression evolved
as a mechanism for demonstrating and pro-
tecting dominant social status. Examples of
resource-acquisition-related aggression in
humans include two men fighting over the
attentions of an attractive woman, a homeless
woman stabbing a wealthy-looking woman
to steal her wallet, or a nation going to war
with another over valuable natural resources.
Restraint behavior should occur when the
target of the aggressive behavior is subdued
or neutralized. As important as aggression
is, the failure to employ proper restraint can
compromise one’s survival. Restraint occurs
in nature when animals are faced with the
submission of a challenger. For example, the
roe deer buck will not clash antlers unless
an opponent is face-on and engaged in the
fight, although he could successfully attack
when the opponent is turned around and vul-
nerable (Eibl-Eibesfeldt, 1961). Likewise,
the defeated wolf will show his neck to his
successful opponent, who refrains from kill-
ing him even though he could do so (Lorenz,
1952). Such restraint in the face of submis-
sion is used to safeguard one’s energy for
future attacks – use of unnecessary energy
can signal that one is vulnerable (Millan,
2006). Moreover, if a social animal fails to
show proper restraint by either attacking oth-
ers without good cause, being overly aggres-
sive in their pursuit of submission from
 EVOLUTIONARY PSYCHOLOGY AND POLICING
others, or continuing an attack when their
opponent is clearly defeated, that individual
will be threatening to the social order and
may be killed, exiled, or (in the human case)
jailed or imprisoned (Francis, 1998).
THE CREATION OF THE POLICE
PROFESSION AND THE SOCIAL
CONTRACT BETWEEN THE POLICE
AND THE CITIZENRY
Evolution explains why using aggression as a
means to an end may be necessary for sur-
vival, and how restraint in humans evolved as
a means of tempering and controlling aggres-
sion, with the result that social order and the
rule of law are preserved. Not all players are
afforded equal power in the exercise of
aggression in contemporary US society. The
police (and other professionals in ascribed
circumstances) are granted the right to exer-
cise legitimate physical force if necessary to
protect public safety.
This can be explained by Social Contract
Theory (SCT), which states that individuals’
moral obligations are shaped by a collective
agreement or ‘social contract’ that binds a
society together with a set of accepted norms
and rules (such as the rule of law). This the-
ory was given its first rigorous defense by
Thomas Hobbes (1588–1679). John Locke
(1632–1704) and Jean-Jacques Rousseau
(1712–1778) are other champions of this
theory. In fact, Locke’s argument that citi-
zens have the right to revolt against author-
ity should it no longer be protecting their
interests was enormously influential on
democratic revolution – notably for Thomas
Jefferson and the founders of the United
States. Revolt against perceived tyrannical
rule or oppression has potential implications
for the current police legitimacy crisis with
the rise of social justice movements such as
Black Lives Matter.
Not all human societies are bound by a
social contract. Reiman (1985) describes the
205
‘state of nature’ as a society without legal
institutions. Although rare, a modern example
of such a system is the Gebusi tribe of New
Guinea – a highly cooperative, egalitarian,
social society which is non-competitive and
politically decentralized with interpersonal
relations that are mutually respectful, non-
hierarchical, and self-effacing. Aggression
among the Gebusi people is discouraged, as
antisocial behavior is contradictory to their
peaceful values. Fear of violence is fostered
and withdrawal from violence is reinforced.
They also have a homicide rate among the
highest reported. Between 1940 and 1982
nearly one-third of adult deaths were caused
by homicide (Knauft et  al., 1987). This is
the equivalent of a homicide rate of 568 per
100,000 per annum. To put this in perspec-
tive, the US homicide rate (one of the high-
est in the Western world) was roughly 6 per
100,000 per annum in 2018 (Federal Bureau
of Investigation, 2018).
The most common cause of homicide is
‘sorcery’, that is, an individual will be killed
when they are believed to have caused death
through sickness to someone else in the vil-
lage. Given that the Gebusi live in the low-
land rainforest of New Guinea, disease is
rampant, explaining the ‘sickness deaths’
and consequent sorcerer killings. Homicide
is considered a regrettable but unavoid-
able burden required to maintain the social
system, and even the close kin members of
the victim rarely seek retribution. Societies
without legal institutions, even highly coop-
erative and peaceable tribes like the Gebusi,
will inevitably require a great deal of inter-
personal violence to prevent anarchy. Within
most systems, the social contract between
police and citizens is the collective antidote
to the ‘state of nature’ Reiman describes.
Individuals recognize that their right to use
force may be countered by the rights of oth-
ers to use force, and that the state of nature is
inherently unsafe and unstable in the absence
of a governing rule. As such, we sacrifice
certain personal freedoms to increase safety
at a broader level: ‘It becomes rational for
206 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
freedom-loving people to renounce their
freedom to use force at their own discretion’
(Reiman, 1985: 239). But in order to do this,
some society members must be responsible
for maintaining public safety. We assign this
duty to the police.
In the mid 19th century, as the United
States expanded through immigration and
the industrial revolution, professional police
forces were established in metropolitan cit-
ies and a social contract authorized police
officers to protect individuals from victimi-
zation (Walker, 1977). Police officers were
granted the authority to employ coercion to
protect citizens’ lives and property (Reiman,
1985). Conceptualized in this way, the police
became society’s ‘professional aggres-
sors’, called upon when force was necessary
(Bittner, 1970). The social contract estab-
lishes the right of police to utilize the force
necessary, including deadly force, if the
threat warrants it. For example, if a suspect
is posing a deadly threat to innocent civilians
the police are justified to shoot. In this case,
aggression is not just allowed, but expected
for the overall good of society. The role of
restraint within the social contract is equally
important. Citizens expect that police author-
ity be utilized legitimately, competently, and
in good faith (Reiman, 1985). Use of force
by police must result in an overall increase in
public safety.
Despite a consistent emphasis on the func-
tion of the police being tied to the legitimate
use of force, the major role of the police has
evolved over time with the demands of the
ruling elite. From preventing slave revolts in
the mid 19th century, to maintaining segrega-
tion following emancipation, to riot enforce-
ment during the civil rights movement, the
police can be seen as the ‘forceful arm’ of
local, state, and federal government, tasked
with maintaining the status quo. The police
have long been directed by the political pow-
ers, who have historically been made up of
wealthy white men. It stands to reason that the
police have traditionally served the interests
of this socially dominant group. A common
argument for why police use of force differs
based on suspect race and socioeconomic sta-
tus is that police discretion favors the socially
dominant and protects the status quo. Of
course, this position tends to be vehemently
denied by members of the socially dominant
group, who argue that suspect behavior is
solely responsible for police use of force.
Nevertheless, the tension between the police
and minority classes has fueled considerable
discord and distrust in police legitimacy.
Use of force perceived by citizens as
unnecessary, unjust, or excessive undermines
the social contract and the legitimacy of the
police. In many circumstances, this is now the
case: the police retain a cultural emphasis on
aggressive crime-control tactics and have a
self-image as society’s law enforcers (Brown,
1988). The citizenry, on the other hand, have
come to expect that police not only control
crime, but serve the community in ways that
build social bonds and prevent crime. Officers
are expected to be mentors, social workers,
mental-health professionals, and counselors,
as well as law enforcers (Terrill et al., 2003).
Similarly, while the citizenry expects greater
and more nuanced restraint in the exercise of
law enforcement, police training and culture
emphasize the need for officers to protect
themselves by staying one step ahead of the
suspect or safety threat. These differences of
opinion reinforce the ‘us vs them’ mentality,
in which officers increasingly feel the pub-
lic does not understand the harsh realities of
what they face on a daily basis, and the public
increasingly resents police authority.
HAWKS VS DOVES AND THE
EVOLUTION OF THE POLICE PERSONA
Maynard-Smith’s famous ‘hawk versus dove’
model is one of the most important contribu-
tions in evolutionary game theory and of
direct relevance to the ‘police persona’. At its
most basic level, the hawk will choose the
strategy of escalating aggression and
 EVOLUTIONARY PSYCHOLOGY AND POLICING
continue attacking until their opponent
retreats or they themselves are injured. A
dove might display aggression but retreat if
their opponent escalates. In a contest between
a hawk and a dove, a hawk would win. In a
contest between two hawks, assuming both
have equal ‘resource-holding potential’
(RHP) such as strength, access to weapons,
etc., there is an equal likelihood of either
winning. In a contest between two doves, the
resource being fought over is shared (nego-
tiation), or the War of Attrition model of
threat to avoid actual fighting is employed to
determine who gets the resource. Maynard
Smith’s (1974) War of Attrition model states
that when animals engaging in conflict
cannot assess the other’s likelihood of beat-
ing them they will attempt to deter conflict
through ritual displays of aggression.
For an evolutionarily stable strategy (ESS)
to occur there must be a mix of hawks and
doves. A system of all doves is vulnerable
to invasion by a mutant hawk and in a sys-
tem of all hawks, the cost of loss becomes
too great. The result is that when hawks are
rare they have the advantage and are selected
for. However, when there are more hawks
than doves some hawks are forced to take on
a dove strategy so the evolutionary ‘seesaw’
that typifies an ESS swings back the other
way as doves are selected for. The same thing
can be said for those that follow the rules and
those that cheat in Hardin’s ‘Tragedy of the
Commons’ scenario, where there are only so
many resources to go around. When cheat-
ers are rare their strategy is highly success-
ful, but eventually the resources will run thin
and their chances of detection will increase;
thus the balance will be tipped back in
favor of the compliers (Hardin, 1968). This
type of ESS can be seen throughout the ani-
mal kingdom. For example, most seagulls
catch fish but some wait by the shore and
steal the food from the hard-working gulls.
This is an effective strategy as cheaters
get maximum benefit for minimum effort.
However, when the cheating gulls start to
outnumber the hard-working gulls some are
207
required to change strategies or nobody gets
food (Dawkins, 1980).
On average, police officers will be more
successful with a hawk strategy than a dove
strategy. This makes sense because if an
officer comes up against a ‘hawk-like’ sus-
pect and they themselves are ‘dove-like’,
the suspect will likely win (get away, attack
them, etc.). On the other hand, if the officer
is ‘hawk-like’ they will likely win against a
dove and have an equal likelihood of winning
against a hawk. Research by Yabuta (2008),
however, has suggested that there might
be a third more complex strategy to add to
the hawk versus dove model with particu-
lar relevance to policing, that of ‘assessor’.
The assessor weighs RHP, then alternates
between hawk and dove strategies depend-
ing on the situation. To assess is an ESS
because it prevents inappropriate attacks on
non-opponents. The role of assessor could
be applied to police officers who must dis-
tinguish between opponents (threatening
suspects) and non-opponents (general mem-
bers of the public or complying suspects) and
modify their strategy accordingly. How they
select these strategies can also be explained
by evolutionary theory.
GAME THEORY AND SELECTION
OF AGGRESSION AND RESTRAINT
STRATEGIES
Selection between aggression and restraint
strategies applies to policing in two impor-
tant ways. First, it is expected that officers
are able to expertly assess whether aggres-
sion is an appropriate response in a given
situation. The justification for the decision is
grounded in the level of threat in their oppo-
nents’ actions. Barash (2004) uses game
theory, and the example of the game ‘rock,
paper, scissors’, to demonstrate the success
of a strategy that is dependent on another
player. If you pick rock and your opponent
also picks rock then you draw, the expected
208 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
return is zero, or E(R,R)=0. If your opponent
has picked scissors you win and the expected
return is one, or E(R,S)=+1; however, if they
picked paper you lose and the expected return
is minus one, or E(R,P)=−1 (Barash, 1982). If
a player consistently employs one strategy
such as rock, their opponent will catch on and
use a defeating strategy. Thus, in the ‘rock,
paper, scissors’ game the best strategy is to
use each with equal probability.
Now apply the game to police use of
deadly force, where the options are ‘shoot’
or ‘don’t shoot’. If the officer shoots and the
suspect represents a real threat, the officer
wins (assuming he or she does not get shot
first). If the officer shoots and the suspect
does not represent a real threat (e.g. is try-
ing to pull out a wallet, not a gun) then the
officer loses and faces the consequences. If
the officer does not shoot and the suspect
represents a real threat the officer loses and
might be injured or killed. Finally, if the
officer does not shoot and the suspect does
not represent a real threat the officer wins,
as he or she has made the right decision.
As in the ‘rock, paper, scissors’ game, the
police cannot always favor the same strategy
because they would be at high risk of mak-
ing an error. Thus, officers must constantly
weigh their perceptions of threat with the
actions of the suspect and the consequences
of their own decisions. Of course, officers
have far more to think about than this simple
analogy implies, and are usually not blind to
the actions of the opponent. However, game
theory exemplifies the decision officers must
make when faced with a threat to employ
either aggression or restraint.
Evolutionary game theory explains why
selection has favored certain characteristics,
behaviors, or attributes, when success in a
contest depends on the behaviors of oth-
ers (Barash, 1982). For example, Maynard
Smith’s War of Attrition model (1974) states
that when animals engaging in conflict can-
not assess the other’s likelihood of beat-
ing them or RHP they will attempt to avoid
conflict through ritual displays of aggression.
This model can be observed in many species,
for example howler monkeys or elephant
seals that loudly vocalize to display aggres-
sion, fish that puff up to try and prevent
attacks, and deer that shake their antlers
at each other to determine who is the more
dominant (Krebs and Davies, 1984). If this
deterrence does not work and a fight ensues,
then the contestant who is prepared to risk a
higher cost and fight for longer will win.
The War of Attrition model relates to
police use of force because it shows how a
cost–benefit analysis has evolved resulting in
aggression only when necessary and only to
the extent necessary. The latter relates to the
second application of the balance between
aggression and restraint in policing: that
officers must apply immediate restraint fol-
lowing the use of aggression. Furthermore,
the very aggression they use should be con-
trolled and strategic as opposed to driven
by fear, anger, or frustration. They are also
expected to render or call for medical aid
for injuries they were personally responsible
for inflicting. The failure to apply appropri-
ate restraint and allow aggression to become
emotionally driven results in incidents such
as the infamous beating of Rodney King in
1991. Video footage of the incident, featuring
Los Angeles Police Department officers beat-
ing an African American man on the ground
and circulated by media outlets nationwide,
has come to symbolize how use of force in
law enforcement can become unfettered,
brutal, and deadly when left unchecked.
Restraint, then, is a crucial component of a
law enforcement officer’s tactical skill set.
The evolutionary foundation of the balance
between aggression and restraint is straight-
forward. Aggression when necessary can
solve several adaptive problems. Aggression
past the point of necessity in social systems,
however, can produce costs and often fails to
solve adaptive problems. The police, just like
every social animal, must achieve a balance
between aggression and restraint. However,
officers are in a reasonably unique position in
having to balance aggression and restraint as
 EVOLUTIONARY PSYCHOLOGY AND POLICING
a core function of their job, and answering to
the public whenever they decide to employ an
aggressive response. In some cases, aggres-
sion can be favored on the part of the police,
and evolutionary theory offers some insights
as to why.
THREAT SIGNALS AND THE
‘FIGHT OR FLIGHT’ RESPONSE
The functioning of the human brain has not
changed since the Pleistocene epoch. Humans
evolved over millions of years in the African
savanna where people lived in small groups
of hunter-gatherers. This environment is
referred to as the ‘environment of evolution-
ary adaptedness’ (EEA) (e.g. Bowlby, 1969).
This is a critical time period and the environ-
ment where natural selection ‘designed’ the
modern human species. One of the most
important tools for survival, favored by natu-
ral selection, was the ‘fight or flight’ system
for responding to threats. Threat response
required efficient learning of threat signals
and detecting threatening objects. An exam-
ple is fear of snakes (Neuberg et al., 2010).
Despite fear of snakes serving little purpose
for the majority of humans in the modern era,
humans tend to be particularly efficient at
learning fearful responses to threat signals,
and particularly inefficient at unlearning
them.
Relatedly, humans tend to be fearful or
wary of coalitional outgroups (groups of
people who are different from them) due to
successful threat responses against attacking
groups in the ancestral environment. This at
least partially explains the concept of ‘implicit
bias’ that humans have against groups that are
different from themselves. Common implicit
biases exist around race, ethnicity, sexual ori-
entation, gender identification, and disability,
among others. Although in the modern era we
(typically) do not need to fear people who are
different from us, we have evolved to favor
this strategy, and this can lead to attitudes and
209
beliefs that we might not be aware of (James,
2017).
Nesse (2005) describes responses to threat
signaling that evolved within the ances-
tral environment, such as the fight or flight
response, and some of the problems that
arise in modern life because of these evolved
responses. These ideas have relevance to
policing, and officers’ use of aggression.
Natural selection resulted in the human nerv-
ous system being highly responsive to threat
cues. We do not wait to see a predator attack-
ing to trigger the response system, but instead
we are sensitive to subtle cues and engage the
sympathetic system that allows us to fight or
flee. Classical conditioning also plays a role
here – if we have experienced a threaten-
ing situation in the past, a similar situation
will be likely to trigger the fight or flight
response in the future. This is the concept
of conditioned anxiety to a cue of danger.
‘False alarms’ are inexpensive relative to the
potential consequences of attack by a preda-
tor. This helps to explain why humans tend to
be risk averse. This could also explain police
shootings where officers shoot in the absence
of concrete evidence that the suspect posed
a deadly threat. From a survival perspective,
the risk of being shot is worse than the risk
of getting it wrong, and the fight or flight
response can make us prone to responding to
threat cues in the absence of real threat.
James, Todak et al. (2018) speculated that this
construct from evolutionary psychology can be
seen in Fachner and Carter’s (2015) ‘threat per-
ception failure’ (TPF) theory. TPF occurs when
an officer mistakes a non-threatening object
(such as a wallet) for a threatening one (such
as a gun), or a non-threatening action (reach-
ing for a wallet) for a threatening one (reach-
ing for a gun) (see also Scharf and Binder,
1983, for a discussion of false-positive
errors). Within the policing literature, TPF is
associated with implicit racial bias, whereby
officers are more likely to experience TPF
when faced with African Americans than
with people of other races and ethnicities.
From an evolutionary perspective, TPF is
210 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
associated with fear and threat responses. Of
course, one could argue that increased fear
indicates increased bias, but bias is not the
sole reason for a threat response.
One of the causes of threat response
in police officers is training they receive
designed to increase anxiety and heighten
attention to threat signals. An example of this
training is ambush training. This type of train-
ing is well intentioned, in that it is designed
to prepare officers for ambush encounters on
the street, but it teaches rapid response over
careful thought, and consequently results in
a heightened risk of error. James Todak and
colleagues (2018) argue that training should
help reduce the incidence of sudden, impul-
sive use of force by officers that might be
fueled by anxiety associated with a fight–
flight response. Training that focuses on
keeping the officer actively engaged (instead
of relying on fight or flight) could reduce the
number of accidental shootings by police
(Binder and Scharf, 1980; Fyfe, 1996). This
is feasible because, just as we learn to detect
and respond to threat signals, we also can
become desensitized to those signals when
they do not result in threat. Understandably,
an argument frequently made by the polic-
ing profession is that desensitization to
threat cues could compromise officer safety.
Directly counter to this argument, evidence
exists that preventing officers from being
overcome by a sympathetic nervous system
response improves officers’ marksmanship
and deadly-force judgment and decision
making, ultimately promoting officer safety
(Johnson et al., 2014).
GENDER-BASED AGGRESSION AND
POLICE CULTURE
Despite gender diversification in the police
profession, policing remains an overwhelm-
ingly male profession. Selection has favored
individual aggression in boys and men, in
particular with regard to the development and
protection of coalitions – groups who work
together towards goals (Geary et  al., 2003).
These groups provide reproductive advan-
tages, as well as increased opportunity for
status and protection. Even in highly social
animals such as humans, group-level dynam-
ics can facilitate individual aggression under
some circumstances, in particular between
male groups. A famous example of this was
observed in the sociological ‘Robbers Cave’
experiments, where randomly assembled
groups of boys engaged in between-group
competition (Puurtinen et al., 2015). During
these 1950s and 1960s experiments, boys
were grouped arbitrarily. They quickly began
coalition building and engaged in competi-
tion with the other group. Evolutionary psy-
chologists posit that this is evidence of
psychological mechanisms that motivate
ingroup cooperation and outgroup-directed
aggression (Sherif et al., 1961). Interestingly,
when provided with a problem that required
resources outside of their group, the boys
would cooperate with the competing group
for a period of time, before reverting to their
own coalition.
These insights have relevance to the
police profession as a coalition. Police cul-
ture emphasizes loyalty to its members and a
distrust of non-members (or at least a strong
feeling that people outside the group do not
understand them or have their best inter-
ests at heart). The police culture has clear
benefits to individual officers – notably the
belief that members of their group will ‘have
their back’ (Paoline, 2003), which promotes
officer safety. The feeling of belonging to a
family is often noted as a draw to policing,
and in many cases generations of people
from the same family will join the profes-
sion. There are other aspects of police culture
which are not as positive, for example the
fostering of an ‘us vs them’ mentality, which
can impair police ability to connect with the
communities that they are expected to protect
and serve. Also the ‘blue wall of silence’ or
unwillingness to ‘rat’ on fellow officers when
they break the rules can lead to accusations of
 EVOLUTIONARY PSYCHOLOGY AND POLICING
secrecy, corruption, and lack of accountability
(Walker, 2001). Finally, the police culture is
inherently masculine, and female officers can
have challenges with acceptance, bias, bigotry,
or unfair promotion practices.
Police researchers have shown that offic-
ers who more closely align with the police
culture are more likely to display aggression,
on and off the job. This includes aggres-
sive tactics during traffic stops (Paoline and
Terrill, 2005) and citizen interactions (Terrill
et  al., 2003). These officers are more likely
to receive citizen complaints (Terrill and
Paoline, 2015) and to use unnecessary force
(Silver et  al., 2017). Furthermore, they are
less likely to adhere to the principles of pro-
cedural justice (Terrill and Paoline, 2015)
and are more likely to engage in misconduct
(Kappeler et al., 1998). Finally, officers who
have a strong connection to police culture are
more likely to engage in intimate-partner vio-
lence (Blumenstein et al., 2012).
DOMINANCE, PRESTIGE, AND
STATUS SEEKING
Dominance hierarchies form when access to
resources is limited and are common in
social species. Within the hierarchy, some
individuals have greater access to resources
than others (Neuberg et  al., 2010). This is
based on social rank, and its association with
reproductive success in social species
(Paquette, 2015). Among humans, these hier-
archies are common across cultures (Neuberg
et  al., 2010) and can be observed even in
young children (Beaulieu and Bugental,
2007). Within dominance hierarchies, those
at the top tend to induce submission to their
dominance either through physical intimida-
tion or control of resources (Cheng et  al.,
2010). In mammals, dominant males have
greater access to mates, which then leads to
selection for these dominant traits.
The construct of prestige, although related
to dominance, differs in that it elicits freely
211
conferred deference instead of intimidation-
induced submission (Henrich and Gil-White,
2001). There is an evolutionary basis for
respecting those we consider to be prestig-
ious. In some societies, prestige exists with-
out dominance hierarchies, and deference
to prestigious individuals offers advantages
(e.g. proximity to potential mates who flock
to the prestigious). Those with prestige enjoy
benefits, including the desire for proximity
to the prestigious (rather than distance, as is
common with regard to dominant individu-
als), the admiration of others, preferential
copying, obedience, and attention (James,
Todak et  al., 2018). Of course, dominance
and prestige are not mutually exclusive,
and many individuals may attain both. But
prestige does not depend on dominance. For
example, a police officer who performs a
heroic feat (such as rescuing a small child) is
likely to achieve prestige, both among peers
and within the community, even if they are
not a ‘dominant’ officer.
Status seeking is related to both dominance
and prestige. Although the construct is from
social psychology, it is relevant to evolution-
ary psychology, because status seeking is a
product of competition for social status and
consequent resources (Geary, 1999). Social
status can be observed in children as young
as two or three years old, particularly dur-
ing same-sex play (Bukowski et  al., 2011).
The construct of status seeking can also be
observed in the literature on juvenile delin-
quency, especially regarding gang participa-
tion and violence (Thrasher, 1936). Within
this context, serious aggression and physi-
cal violence can occur, especially between
males, over issues that seem trivial. This is
related to the idea of ‘saving face’ and not let-
ting other males disrespect or question one’s
social status (Felson and Steadman, 1983).
Within policing, dominance, prestige,
and status seeking are readily observed. A
police officer who takes a dominant approach
is likely to be seen as a protector, an aggres-
sor, and someone who will not back down in
the face of attack. This strengthens ties to the
212 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
police culture, and also promotes aggressive
tactics with citizens, for example officers
who escalate their responses in an effort to
maintain control in a situation (Alpert et al.,
2004). Escalation tactics are more frequently
observed when officers are faced with citi-
zens who disrespect their authority (Van
Maanen, 1978) or display ‘contempt of cop’
(James, James et  al., 2018). Thus, officers’
physical responses to threats to authority
and character can be seen as mechanisms for
maintaining dominance during the encounter.
This adheres to the policing culture, and is
likely to secure prestige and status from their
police-officer peers.
TOWARDS A BALANCE OF
AGGRESSION AND RESTRAINT
IN POLICING
Despite a focus on aggression within the
policing culture, police officers are adept at
employing restraint. They do not typically
employ force indiscriminately. Recently,
however, serious allegations of excessive and
unnecessary force used by the police against
young Black men has sparked controversy
surrounding the police profession. This has
led to calls for action, and in some cases citi-
zens taking matters into their own hands
(either by rioting or attacking the police).
Throughout the evolution of human society,
when social order is threatened and revolution
looms, the antidote is often authority reform
(DeBenedetti, 1980; Shaw and Shaw, 1977;
Wolpert, 1962). The final report from President
Obama’s Task Force on 21st Century Policing
recommends police reform to reduce tensions
between the police and the people they are
sworn to serve and protect (President’s Task
Force on 21st Century Policing, 2015).
The majority of the recommendations from
the report are targeted at improving public
trust in police, repairing broken relation-
ships, increasing police legitimacy, and pro-
moting procedural justice. Avenues towards
accomplishing these goals include training
reform, policy change, and greater transpar-
ency. However, the type of reform the public
expects is not likely to be successful while
the majority of police training emphasizes
aggressive tactics and police culture resists
reform. According to the Police Executive
Research Forum (2015), over 90% of the
training hours officers currently receive are
on aggressive tactics. Paired with the empha-
sis on physical aggression within the police
culture, tactics that promote restraint are
likely to be less appealing to police offic-
ers (Crank, 2014). The framework depicted
in Figure 10.1 re-envisions the goal of the
police in modern society as expertly balanc-
ing aggression and restraint.
The proposed framework acknowledges
the importance of controlled aggression in
policing as a strategy for maintaining social
order, but emphasizes the need to tem-
per aggression with appropriate levels of
restraint. This balance can be demonstrated
across all elements of policing, from routine
to deadly encounters. The proposed frame-
work does not diminish the importance of
aggression in situations in which it is legiti-
mately required. For example, to compe-
tently arrest an assaultive suspect, an officer
must overpower any resistance. Similarly,
encounters that warrant police force require
an officer to aggressively ‘win’ against the
suspect. This is especially true for use of
deadly force where the officer must guard
against loss of innocent life, including their
own. When officers are confident in their
ability to employ aggression, they are less
likely to activate a flight or fight response,
because they are less likely to feel that their
resources are overwhelmed. In other words,
the police officer must have the ability to be
a hawk, in order to select whether a hawk or
dove strategy is more appropriate.
The other side of the balancing scale is
restraint. Restraint in this case is not to be
mistaken for meekness, second-guessing, or
unwillingness to dominate if the situation
requires it. Restraint is defined as the ability
 EVOLUTIONARY PSYCHOLOGY AND POLICING
Figure 10.1  Police balance between aggression and restraint
to temper aggression, or avoid it when it is not
warranted. On the restraint side, the officer is
supplied with a toolkit of additional options
that circumvent the use of physically aggres-
sive tactics. It is important to note that these
tactics (for example, tactical disengagement
and verbal de-escalation) are not new; they
have been around since the inception of the
police profession. Arguably, however, they
have yet to become central components of
police work, given that the majority of train-
ing continues to promote aggressive tactics,
and police culture continues to reward physi-
cal aggression.
PROMOTING RESTRAINT IN POLICE
TRAINING AND POLICY
Several strategies exist in the police profes-
sion for promoting restraint. One example is
the use of ‘tactical disengagement’ or actively
attempting to de-escalate a volatile encounter.
213
Police departments in the United States,
including Kansas City (Missouri), have begun
to implement tactical disengagement training
(Police Executive Research Forum, 2015).
There has been a move in other countries
such as Canada and the UK in this direction,
as well. The idea is to not force an encounter
with a citizen when there is a risk of escala-
tion and the reason for the encounter is minor.
This mindset is antithetical to predominant
cultural values in policing, which dictate that
officers should not back down from a chal-
lenge and should jump in quickly to handle a
situation (Fyfe, 1986; Paoline, 2003). The
tactical disengagement philosophy teaches
officers that they do not need to initiate or
‘win’ every encounter.
Relatedly, communication tactics such
as ‘verbal judo’, de-escalation, and motiva-
tional interviewing have been taught in many
departments as a way to influence citizens
into voluntary compliance without using
physical coercion (Humphrey, 2013). Such
techniques draw on professionalism and
214 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
empathetic connections with the i­ndividual’s
personal situation to reach a mutually agreed-
upon solution to the problem. Verbal tech-
niques require expert restraint and can be
especially frustrating for police officers when
they are faced with individuals who are com-
bative, disrespectful, or rude (Pusatory, 2016).
Switching immediately from aggressor to
rendering medical aid is another example of
how officers balance aggression and restraint.
In fact, officers today are increasingly expected
to immediately render aid to an injured person
or else be publicly condemned for neglect-
ing human life (Dart and Walters, 2016).
Rendering life-saving aid following a use of
force requires that an officer changes mindsets
quickly, shifting from a ‘warrior’ to a ‘guard-
ian’ role (Rahr and Rice, 2015) – an officer
must drop their defensive and offensive stance
and save the person’s life. An officer may need
to shift back and forth between controlled
aggression and restraint if the individual con-
tinues to resist or fight as the officer employs
medical aid. From an evolutionary perspective,
these expectations move beyond a simple deci-
sion to employ restraint upon achieving the
end goal – we are asking the police to engage
in the seemingly more unnatural behavior of
actually preserving the wellbeing of a physical
opponent by employing life-saving aid.
Collaboration with other services also
requires restraint on the part of officers.
Officers are accustomed to being called on to
solve a wide range of life problems that often
have nothing to do with law enforcement but
that the citizenry feels the police should do
something about right away (Bittner, 1974).
As such, police agencies are beginning to
identify areas of police work that may be bet-
ter handled if they are redirected to other pro-
fessionals, or are handled collaboratively by
multiple agencies. For example, when inter-
acting with a suspect suffering from mental
illness, an officer sometimes has the option to
call on mental health professionals who can
offer expert advice on the individual’s behav-
ior. Indeed, this is an underlying philosophy
of Crisis Intervention Team (CIT) training.
Officers can exercise restraint in tempering
their tactics according to the mental health
professionals’ information and suggestions.
The same can be said for communicating
with victims, distraught family members,
friends, and witnesses on scene.
Police departments in some US cities have
begun to re-engineer their use of force train-
ing towards an emphasis on violence de-
escalation and avoidance. Officers in the Las
Vegas (Nevada), New York City (New York),
Seattle (Washington), Oakland (California),
and Leesburg (Virginia) police departments
have all received training on violence de-
escalation, teaching tactics designed to de-
escalate a police encounter and avoid the
need to use physical force to solve the prob-
lem. Unfortunately, programs that incorporate
some form of de-escalation or verbal tactics
training are frequently offered in a fragmented
manner, failing to teach how these skills can
be integrated with the use of force training.
A prevailing criticism of de-emphasizing
aggressive tactics training for police is that
it will result in decreased officer safety, and
consequently decreased community safety. A
central argument here, often voiced by offic-
ers themselves, is that researchers and oth-
ers who are advocating for reform in police
use of force do not understand the danger-
ous realities of police work. Supporting
this argument, some evidence suggests that
hyper-vigilance on the part of the police is
necessary. Interviews with individuals who
feloniously assaulted a police officer found
that they were more likely to attack an officer
if he or she seemed unprepared to react to a
problem, did not appear to know an attack
was coming, or seem to have dropped their
guard (Pinizzotto et al., 2006). This research
confirms police-culture beliefs, and has made
the shift towards violence de-escalation diffi-
cult to accept by many rank-and-file officers.
In contrast, de-escalation tactics may
promote officer safety. In a crisis of police
legitimacy, citizens are less likely to follow
the law and comply with police commands.
This has been documented in the policing
 EVOLUTIONARY PSYCHOLOGY AND POLICING
literature – people who perceive the police as
legitimate are more likely to cooperate and
obey the law (Mastrofski et  al., 1996; Tyler
and Fagan, 2008). Thus, training the police
in conflict- and violence-avoidance (i.e. to be
more restrained) may reduce the likelihood
that situations will escalate and the officer
will become engaged in conflict. Such a situ-
ation results in an overall reduction in the risk
of harm to all persons involved.
GROUNDING POLICE TRAINING IN
EVOLUTIONARY THEORY
With respect to threat response, training for
reducing unnecessary or excessive force
could focus on officers’ concerns about
danger, and reduce the amount of training
that promotes rapid response over reasoned
decision making. For example, ambush train-
ing that teaches an officer that it is beneficial
to be constantly alert may help an officer
during the (statistically) unlikely situation
that they are ambushed. However, it is also
likely to increase the risk of officers rapidly
responding to a perceived threat without evi-
dence that a threat exists. It is unlikely that
officers in the field will favor critical deci-
sion making over rapid threat response unless
they have been taught to do so. Deadly-force
judgment and decision-making training
(either simulation or role-play based) can
help promote critical decision making over
rapid response, due to the consequence of
‘getting it wrong’.
Police training can also reduce officers’
natural distrust of ‘outgroups’ via exposure.
As the Robbers Cave experiments demon-
strate, competing groups can put aside differ-
ences and overcome hostility when faced with
a common problem requiring a cooperative
solution. Community-orienteered policing
(COP) strategies that focus on shared goals,
such as improving the safety of people living
in the community, have potential to discour-
age police perceptions of citizens as rivals
215
and vice versa. Examples of COP strategies
include programs that promote citizen coop-
eration in crime reduction and events that
allow police and citizens to interact without a
power imbalance, such as non-crime-related
community events.
Another strategy for reducing outgroup dis-
trust is implicit-bias training, which teaches
officers about their biases, and provides
strategies for identifying and overcoming the
impact of bias on behavior. Diversification of
the police force also has potential to reduce
outgroup suspicion, due to promotion of ties
to both the police and minority communities.
Relatedly, promoting minority individuals
to positions of power where they influence
decisions and represent the police profession
has promise for reducing outgroup distrust
and suspicion. Although promising, these
strategies have yet to be evaluated for effec-
tiveness at promoting inter-group collabora-
tive relationships between the police and the
citizenry.
The theories of dominance, prestige, and
status seeking also have relevance to police
training. It is easy to train officers to exhibit
dominant control behavior. This is because
this training aligns with evolved psychology.
In the ancestral environment, however, the
expectation related to an individual’s domi-
nance behavior is that if the individual wins,
the opponent submits. This is complicated
in the modern environment because police
authority is granted institutionally, and not
based on a direct competition between indi-
viduals. In other words, no contest has estab-
lished that the police officer is dominant to
the citizen, so it is naïve to assume that the
citizen will always submit. The citizen might
have strong reasons for not submitting (e.g.
gaining prestige and social status among
peers). This can result in challenge to the
police authority which, in turn, the police
are unlikely to submit to. Police training that
demands officers exert dominance will in
these instances lead to escalation.
Alternatively, training that teaches offic-
ers to reduce the likelihood of unnecessary
216 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
dominance contests can prevent the need
for aggression. There are situations when
it is appropriate and expected for an officer
to be a hawk and take immediate control of
a situation, regardless of escalation conse-
quences. For example, if a suspect is armed
and posing a deadly threat to those around
them, an officer must neutralize that threat,
quickly and decisively. The vast majority of
police–citizen interactions, however, do not
require the police to be forceful. Many of
these interactions, particularly for police offic-
ers that work in ‘anti-cop’ neighborhoods, will
be fraught with challenges to police authority.
Policing scholar Van Maanen (1978) describes
‘the asshole’ or citizen who is not inclined to
submit to police authority. When facing such
a citizen, the officer can either engage in a
dominance challenge or not. In the absence of
criminal behavior, the officer has no grounds
to force a citizen to submit to their authority
(Klinger, 1994), and doing so will escalate
the situation. Unfortunately for the police, not
everyone will like them, and taking that per-
sonally is both unprofessional and unsafe.
De-escalation training has potential for
reducing unnecessary dominance con-
tests between the police and the citizenry.
De-escalation techniques are typically com-
munication based, and attempt to influence
citizens into voluntary compliance without
using force (Humphrey, 2013). They can be
used in volatile crisis encounters (e.g. hostage
negotiation) or in day-to-day encounters for
decreasing the likelihood that a situation will
escalate (e.g. procedural justice). These tech-
niques emphasize police need to read people,
be professional, display empathy, and treat
people with dignity and respect, regardless of
their attitude towards the police. Evaluations
of de-escalation techniques show that they
improve public perceptions of police legiti-
macy (Todak, 2017).
The idea that de-escalation techniques
can be used to dissuade citizens from chal-
lenging police authority has implications for
long-term police–community relationships.
Influencing citizens away from antagonism
and towards cooperation results in more
effective police work. The traditional ‘police
persona’ of aggression, authority, and mas-
culinity takes the strategy of deterrence to
prevent challenges to dominance (‘my RHP
is bigger than your RHP’). The evidence on
the effectiveness of this strategy, however,
is limited, and evidence exists that coopera-
tive strategies are more effective (Tyler and
Fagan, 2008). For example, the research liter-
ature on procedural justice demonstrates that
officers who treat people fairly, with dignity
and respect, listen to them, and work towards
mutually beneficial outcomes are more likely
to gain voluntary compliance and avoid use-
of-force encounters (Tyler and Fagan, 2008).
CONCLUSION
Bittner (1970) defined the role of the police
in terms of their legitimate authority to exer-
cise coercive force. Police culture has
embraced this definition, shaping training
and policy around the use of aggressive
crime-control techniques. This mindset has
led to strained relationships between police
and minority communities and a social crisis
in the United States characterized by eroded
police legitimacy. Examining police strate-
gies for selecting tactics through an evolu-
tionary lens provides a framework that
re-envisions police function as the expert
balance of controlled aggression and restraint.
Aggressive ‘hawk-like’ strategies are appro-
priate in certain situations, but in many
others, restrained ‘dove-like’ strategies will
be more effective at gaining voluntary com-
pliance and avoiding unnecessary dominance
contests. Adopting this framework by focus-
ing more training hours on force alternatives
and promoting a culture of de-escalation may
result in police behavior falling more closely
in line with citizens’ expectations of police.
Such a shift could result in a reduction in the
rate of violence that occurs between police
and citizens.
 EVOLUTIONARY PSYCHOLOGY AND POLICING
REFERENCES
Alpert, G. P., Dunham, R. G., & MacDonald,
J. M. (2004). Interactive police-citizen
encounters that result in force. Police
Quarterly, 7(4), 475–488.
Barash, D. P. (2004). The survival game: How
game theory explains the biology of
cooperation and competition. New York, NY:
Macmillan.
Barash, D. (1982). Sociology and behavior
(2nd ed.). New York, NY: Elsevier.
Beaulieu, D. A., & Bugental, D. B. (2007).
An evolutionary approach to socialization. In
J. E. Grusec & P. D. Hastings (Eds.), Handbook
of Socialization (pp. 71–95). New York:
Guilford Press.
Binder, A., & Scharf, P. (1980). The violent
police-citizen encounter. The ANNALS of the
American Academy of Political and Social
Science, 452(1), 111–121.
Bittner, E. (1970). The functions of the police in
modern society. Bethesda, MD: National
Institute of Mental Health.
Bittner, E. (1974). Florence Nightingale in
pursuit of Willie Sutton: A theory of the
police. In H. Jacob (Ed.), Potential for Reform
of Criminal Justice (pp. 17–44). Beverly Hills,
CA: Sage.
Blumenstein, L., Fridell, L., & Jones, S. (2012).
The link between traditional police sub-
culture and police intimate partner violence.
Policing: An International Journal of
Police Strategies & Management, 35(1),
147–164.
Bowlby, J. (1969). Attachment and loss. New
York, NY: Basic Books.
Brown, M. K. (1988). Working the street:
Police discretion and the dilemmas of reform.
New York, NY: Russell Sage Foundation.
Bukowski, W. M., Buhrmester, D., &
Underwood, M. K. (2011). Peer relations as
a developmental context. In M. K.
Underwood & L. H. Rosen (Eds.), Social
Development: Relationships in Infancy,
Childhood, and Adolescence (pp. 153–179).
New York, NY: Guilford Press.
Buss, D. M., & Shackelford, T. K. (1997). Human
aggression in evolutionary psychological
perspective. Clinical Psychology Review,
17(6), 605–619.
217
Cheng, J. T., Tracy, J. L., & Henrich, J. (2010).
Pride, personality, and the evolutionary
foundations of human social status. Evolution
and Human Behavior, 31(5), 334–347.
Crank, J. P. (2014). Understanding police
culture. Philadelphia:Routledge.
Dart, T., & Walters, J. (2016, September 22).
Tulsa police under scrutiny for delayed
medical aid given to Terence Crutch. The
Guardian. Retrieved from www.theguardian.
com/us-news/2016/sep/22/tulsa-police-
t e re n c e - c r u t c h e r- m e d i c a l - a s s i s t a n c e.
(Accessed 28 March 2018).
Dawkins, R. (1980). Good strategy or
evolutionary stable strategy? In G. W. Barlow
& J. Silverberg (Eds.), Sociobiology: Beyond
Nature/Nurture, Westview Press, Boulder,
Colorado, pp. 331–367.
DeBenedetti, C. (1980). The peace reform in
American history. Bloomington, IN: Indiana
University Press.
Eibl-Eibesfeldt, I. (1961). The fighting behavior
of animals. Scientific American, 205,
112–122.
Fachner, G., & Carter, S. (2015). An assessment
of deadly force in the Philadelphia Police
Department (Collaborative Reform Initiative).
Washington, DC: Office of Community
Oriented Policing Services, US. Department
of Justice.
Federal Bureau of Investigation. (2018).
Uniform Crime Report, January–June 2018.
Retrieved from https://ucr.fbi.gov/crime-in-
the-u.s/2018/preliminary-report. (Accessed
28 October 2018)
Felson, R. B., & Steadman, H. J. (1983).
Situational factors in disputes leading to
criminal violence. Criminology, 21(1),
59–74.
Francis, R. C. (1988). On the relationship
between aggression and social dominance.
Ethology, 78(3), 223–237.
Fyfe, J. J. (1986). The split-second syndrome
and other determinants of police violence. In
A. T. Campbell & J. J. Gibbs (Eds.), Violent
Transactions (pp. 207–225). Oxford, UK:
Basil Blackwell.
Fyfe, J. J. (1996). Training to reduce police-
citizen violence. In W. A. Geller & H. Toch
(Eds.), Police Violence: Understanding and
Controlling Police Abuse of Force. New
Haven, CT: Yale University Press.
218 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Geary, D. C. (1999). Evolution and
developmental sex differences. Current
Directions in Psychological Science, 8(4),
115–120.
Geary, D. C., Byrd-Craven, J., Hoard, M. K.,
Vigil, J., & Numtee, C. (2003). Evolution
and development of boys’ social behavior.
Developmental Review, 23(4), 444–470.
Gest, T. (2016, July 8). Is a ‘police legitimacy
crisis’ driving homicides up? The Crime
Report. Retrieved from http://thecrimereport.
org/2016/07/08/is-a-police-legitimacy-crisis-
driving-homicides-up/ (Accessed 3 March
2018).
Hardin, G. (1968). Tragedy of the Commons.
Science, 162, 1243–1248.
Henrich, J., & Gil-White, F. J. (2001). The
evolution of prestige: Freely conferred
deference as a mechanism for enhancing the
benefits of cultural transmission. Evolution
and Human Behavior, 22(3), 165–196.
Humphrey, J. (2013, November 19). State
police academy building guardians instead of
warriors. KXLY News. Retrieved from https://
www.kxly.com/state-police-academy-
building-guardians-instead-of-warriors/
(Accessed 3 March 2018).
James, L. (2017). The stability of implicit
racial bias in police officers. Police
Quarterly. 21(1), 30–52. doi: 10.1177/
1098611117732974
James, L., Fridell, L., & Straub, F. (2016,
February). Implicit bias versus the ‘Ferguson
Effect’: Psychosocial factors impacting
officers’ decisions to use deadly force. The
Police Chief, 83, 44–51.
James, L. James, S., & Vila, B. (2018). Testing
the impact of citizen characteristics and
demeanor on police officer behavior in
potentially violent encounters. Policing: An
International Journal of Police Strategies &
Management, 41(1), 24–40. https://doi.
org/10.1108/PIJPSM-11-2016-0159
James, L., Todak, N., & Savage, J. (2018).
Unnecessary force by police: Insights from
evolutionary psychology. Policing: A Journal
of Policy and Practice. 14(1), 278–291.
Johnson, R. R., Stone, B. T., Miranda, C. M.,
Vila, B., James, L., James, S. M., & Berka, C.
(2014). Identifying psychophysiological
indices of expert vs. novice performance in
deadly force judgment and decision
making. Frontiers in Human Neuroscience,
8, 512.
Jones, J. M. (2015, June 29). In U.S., confidence
in police lowest in 22 years. Gallup. Retrieved
from www.gallup.com/poll/183704/
confidence-police-lowest-years.aspx
(Accessed 25 July 2017).
Kappeler, V. E., Sluder, R. D., & Alpert, G. P.
(1998). Forces of deviance: Understanding
the dark side of policing (2nd ed.). Long
Grove, IL: Waveland Press.
Klinger, D. A. (1994). Demeanor or crime? Why
‘hostile’ citizens are more likely to be
arrested. Criminology, 32(3), 475–493.
Knauft, B. M., Daly, M., Wilson, M., Donald, L.,
Morren Jr, G. E., Otterbein, K. F., & van
Wetering, W. (1987). Reconsidering violence
in simple human societies: Homicide among
the Gebusi of New Guinea [and comments
and reply]. Current Anthropology, 28(4),
457–500.
Krebs, J. & Davies, N. (1984). Behavioural
ecology: An evolutionary approach. Oxford:
Blackwell Scientific Publications.
Lorenz, K. (1952). King Solomon’s ring. London,
UK: Methuen.
Mastrofski, S. D., Snipes, J. B., & Supina, A. E.
(1996). Compliance on demand: The public’s
response to specific police requests. Journal
of Research in Crime and Delinquency, 33(3),
269–305.
Millan, C. (2006). Cesar’s way. New York, NY:
Harmony Books.
Nesse, R. M. (2005). Natural selection and the
regulation of defenses: A signal detection
analysis of the smoke detector principle.
Evolution and Human Behavior, 26(1),
88–105.
Neuberg, S. L., Kenrick, D. T., & Schaller, M.
(2010). Evolutionary social psychology.
In S. T. Fiske, D. Gilbert, & G. Lindzey
(Eds.), Handbook of Social Psychology
(pp. 761–796). New York: Wiley.
Paoline, E. A. (2003). Taking stock: Toward a
richer understanding of police culture.
Journal of Criminal Justice, 31(3),
199–214.
Paoline, E. A., & Terrill, W. (2005). The impact
of police culture on traffic stop searches: An
analysis of attitudes and behavior. Policing:
An International Journal of Police Strategies
& Management, 28(3), 455–472.
 EVOLUTIONARY PSYCHOLOGY AND POLICING
Paquette, D. (2015). An evolutionary
perspective on antisocial behavior: Evolution
as a foundation for criminological theories.
In J. Morizot & L. Kazemian (Eds.),
The Development of Criminal and Antisocial
Behavior (pp. 315–330). New York:
Springer.
Peck, J. H. (2015). Minority perceptions of the
police: A state-of-the-art review. Policing: An
International Journal of Police Strategies &
Management, 38(1), 173–203.
Pinizzotto, A. J., Davis, E. F., & Miller, C. E.
(2006). Violent encounters: A study of
felonious assaults on our nation’s law
enforcement officers (No. NCJ 231272).
Washington, DC: US Department of Justice,
Federal Bureau of Investigation.
Police Executive Research Forum. (2015).
Re-engineering training on police use of
force (Critical Issues in Policing). Washington
DC: Police Executive Research Forum.
Retrieved from https://www.policeforum.
org/assets/reengineeringtraining1.pdf
President’s Task Force on 21st Century Policing.
(2015). Final Report of the President’s Task
Force on 21st Century Policing. Washington,
DC: Office of Community Oriented Policing
Services.
Pusatory, M. (2016, August 10). Watch:
Spokane officer shows amazing patience
dealing with intoxicated man. Fox 28.
Retrieved from https://www.khq.com/news/
watch-spokane-officer-shows-amazing-
patience-dealing-with-intoxicated-man/
article_616ebc29-947b-58d5-937c-
ccf1cfbb50e8.html (Accessed 25 July
2017).
Puurtinen, M., Heap, S., & Mappes, T. (2015).
The joint emergence of group competition
and within-group cooperation. Evolution
and Human Behavior, 36(3), 211–217.
Rahr, S., & Rice, S. K. (2015). From warriors to
guardians: Recommitting American police
culture to democratic ideals (No. NCJ 24865).
Laurel, MD: National Institute of Justice and
the Harvard Kennedy School Program in
Criminal Justice Policy and Management.
Reiman, J. (1985). The social contract and the
police use of deadly force. In F. A. Ellison &
M. Feldberg (Eds.), Moral Issues in Police
Work. (pp. 237–249) Savage, MD: Rowman
& Littlefield Publishers.
219
Savage, J., & Kanazawa, S. (2004). Social
capital and the human psyche: Why is social
life ‘capital’? Sociological Theory, 22(3),
504–524.
Scharf, P., & Binder, A. (1983). The badge and
the bullet: Police use of deadly force. New
York, NY: Praeger.
Shaw, S. J., & Shaw, E. K. (1977). History of the
Ottoman Empire and Modern Turkey:
Volume 2, Reform, Revolution, and Republic:
The Rise of Modern Turkey 1808–1975.
Cambridge, UK: Cambridge University Press.
Sherif, M., Harvey, O. J., White, B. J., Hood, W.
R., & Sherif, C. (1961). The Robbers Cave
experiment: Intergroup conflict and
cooperation. Institute of Group Relations,
University of Oklahoma.
Silver, J. R., Roche, S. P., Bilach, T. J., &
Bontrager Ryon, S. (2017). Traditional police
culture, use of force, and procedural justice:
Investigating individual, organizational, and
contextual factors. Justice quarterly, 34(7),
1272–1309.
Smith, J. M. (1974). The theory of games and
the evolution of animal conflicts. Journal of
Theoretical Biology, 47(1), 209–221.
Smith, J. M. (1982). Evolution and the Theory
of Games. Cambridge University Press.
Terrill, W., & Paoline, E. A. (2015). Citizen
complaints as threats to police legitimacy:
The role of officers’ occupational attitudes.
Journal of Contemporary Criminal Justice,
31(2), 192–211.
Terrill, W., Paoline, E. A., & Manning, P. K.
(2003). Police culture and coercion.
Criminology, 41(4), 1003–1034.
Thrasher, F. M. (1936). The boys’ club and
juvenile delinquency. American Journal of
Sociology, 42(1), 66–80.
Todak, N. (2017). De-escalation in police-citizen
encounters: A mixed methods study of a
misunderstood policing strategy
(Dissertation). Arizona State University,
Phoenix, AZ.
Tyler, T. R., & Fagan, J. (2008). Legitimacy and
cooperation: Why do people help the police
fight crime in their communities? Ohio State
Journal of Criminal Law, 6, 231–275.
Van Maanen, J. (1978). The asshole. In P. K.
Manning & J. V. Maanen (Eds.) Policing: A
View from the Street (pp. 221–238). Santa
Monica, CA: Goodyear.
220 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

Walker, S. (1977). A critical history of police
reform. Lexington, MA: Lexington Books.
Walker, S. (2001). Police accountability: The role
of citizen oversight. Belmont, CA: Wadsworth.
White, M. D., & Fradella, H. F. (2016). Stop and
frisk: The use and abuse of a controversial
policing tactic. New York, NY: New York
University Press.
Wolpert, S. A. (1962). Tilak and Gokhale:
Revolution and reform in the making of
modern India. Berkeley and Los Angeles, CA:
University of California Press.
Yabuta, S. (2008). Evolution of cross-contextual
displays: The role of risk of inappropriate
attacks on nonopponents, such as partners.
Animal Behaviour, 76(3), 865–870.

Evolutionary Psychology,
Jurisprudence, and Sentencing
Eyal Aharoni and Morris B. Hoffman
I. JURISPRUDENCE AND
PUNISHMENT1
Two thousand years ago, Socrates and an
Athenian Sophist named Thrasymachus began
a famous debate about human nature and the
meaning of justice (Plato, 380 BCE). Are
humans fundamentally good, built to cooperate
and to appreciate beauty and truth, as Socrates
contended, and therefore perhaps in need only
of modest and occasional intervention by the
state? Or are we fundamentally bad, built only
to maximize our self-interest, as Thrasymachus
argued, and therefore probably in need of
heavy-handed restraint by a robust state? Is
justice ‘the excellence of the soul’ (Plato, 380
BCE: 297) or nothing more than ‘the interest of
the stronger’(Plato, 380 BCE: 275)?
Humans have been having these same
debates ever since, and the ways we have
resolved them have largely defined our political
and legal institutions. It is no coincidence that
the founders of the United States were steeped
11
in the Enlightenment’s decidedly mixed
version of this controversy. The Constitution’s
distribution of powers between different
branches of government was a reflection of the
framers’ nuanced views about human nature.
We are good enough to be largely free of an
overbearing state, but not quite good enough
to live without a state or to populate it without
checks and balances between its parts.
Until the paradigm-shifting insights of
evolutionary psychology, behavioral eco-
nomics, and the other disciplines described in
Section II of this chapter, most modern takes
on human nature have been skewed heavily
toward Thrasymachus’ dreary views. The
central simplifying assumption of classical
economics was that each of us is relentlessly
self-interested. Markets are an efficient inte-
gration of all those individual greedy unseen
hands. Darwin’s insights strengthened the
belief that we are self-interest machines, and
biology’s great synthesis of evolution and
genetics simply moved the locus of that self-
interest from the selfish individual animal
222 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
down to the animal’s selfish genes. Freud
did much the same for the psyche, and Marx
for the allegedly relentless march of eco-
nomic and political history, framing the wars
between self-interested classes.
But then something delightful happened
on this dark road to modern pessimism.
Some economists, anthropologists, and psy-
chologists had the audacity to look systemati-
cally at how humans actually behave instead
of how these dreary theories predicted we
should behave. And there were many surpris-
ing results. They discovered that when we
play economic games in the laboratory, we
engage in all kinds of cooperative behaviors
that cannot be explained by classical eco-
nomics, including sharing with and trusting
other players, even when they are strangers.
They discovered that we are not only not the
rational self-interest machines conceived by
classical economic theory, but that our appar-
ent irrationality is predictable in many impor-
tant decision-making domains. As discussed
in more detail in Section II below, all these
observations were then linked to the theoreti-
cal insights of evolutionary psychology.
These same ancient and modern debates
about human nature and the meaning of jus-
tice have echoed throughout the philosophy
of law, also called ‘jurisprudence’. There are
four main schools of jurisprudence – natural
law, legal positivism, legal realism, and nor-
mative law – all addressed to these funda-
mental questions.
Natural-law theorists view law in a way
that is analogous to how natural philoso-
phers viewed the physical universe: there
are a priori principles (like Socrates’ truth,
beauty, and justice) that animate legal sys-
tems just as physical laws (gravity, conserva-
tion of momentum) animate the universe, and
humankind’s job is to discover these natural
laws and apply them (Bix, 2010; Haakonssen,
1996). Natural law need not be grounded in
the divine, although its most famous his-
torical proponents – Socrates, Aristotle,
and Thomas Aquinas – of course did so. As
modern philosophers generally became less
satisfied with any systems that depended
on unexaminable axioms of theology, natu-
ral law began to fall out of fashion. But
there have been, and continue to be, modern
efforts to re-ground natural law in concepts
of secular morality, most prominently by Lon
Fuller (Fuller, 1965) and Ronald Dworkin
(Dworkin, 1986). We will see similar re-
kindled interests in secular morality from the
normativists and even some positivists.
Even though natural law’s fundamental
principle is that law is a formal expression
of morality, that position does not mean that
all natural-law theorists take Socrates’ side
in the debate about human nature. For exam-
ple, Thomas Hobbes, a prominent Scottish
proponent of natural law whose views were
important to the founders of the United
States, believed humankind’s natural tenden-
cies were toward selfishness and violence,
tendencies that needed curbing by a power-
ful state. Hobbes’ view of right and wrong
seems presciently Darwinian; he once wrote
that man is forbidden by natural law ‘to do
that which is destructive to his life, to take
away the means of preserving the same, or to
omit that by which he thinks it may best be
preserved’ (Hobbes, 1651).
The second school of jurisprudence is called
‘legal positivism’. Like Thrasymachus before
them, legal positivists rejected the fusing
together of law and morality. The ‘positivism’
simply means that laws are rules posited by
man, not by God. They are social constructs
and nothing more. The role of the legal posi-
tivist is to describe those social constructs and
to analyze them non-normatively, especially
the processes by which laws come into being
(Coleman and Leiter, 2010). There are many
different subspecies of legal positivism, some
of which vary by the degree to which they pay
attention to moral groundings (Gardner, 2001).
One of the most prominent modern positivists
was H. L. A. Hart (Hart, 1961), who famously
debated Lon Fuller in the Harvard Law Review
in 1958 (Fuller, 1958; Hart, 1958).
The third school of jurisprudence is ‘legal
realism’. Legal realists, like positivists, take a
 EVOLUTIONARY PSYCHOLOGY, JURISPRUDENCE, AND SENTENCING
non-normative view of law. But unlike most
positivists, they are not satisfied with simply
accepting the conventions of law as given
social constructs. Instead, most legal realists
are interested in the psychological engines
that drive legislatures to adopt laws and
judges to interpret them (Leiter, 2010). The
law and economics phenomenon – using the
insights of economics to study the relation-
ship between law and its intended and unin-
tended consequences – is a modern branch of
legal realism. There are many other branches,
including perhaps the most extreme one,
usually credited to Oliver Wendell Holmes,
Jr. In Holmes’ view, law is nothing more
than a prediction of the results of adjudica-
tion. The promise contained in a contract,
under this view, means nothing other than a
prediction that whoever breaks the promise
will have to pay damages. To Holmes, law
is power dressed up in process (Alschuler,
2000). Critical legal theory and its cousin,
critical race theory, are modern versions of
this extreme legal realism.
The fourth and most recent school of juris-
prudence is normative law. The normativ-
ists, somewhat like Fuller’s and Dworkin’s
secular naturalism and the more morally
interested positivists, focus on the founda-
tions and justifications of the law. They are
not satisfied either with the strict positivist
assumption that laws are just arbitrary social
constructs or with the realists’ fascination
with the levers of power. They examine the
purposes of law, and study methods to meas-
ure whether those purposes have been opti-
mized (Shiner, 2010).
All you non-philosophers out there are
probably wondering what in the world any
of this esoteric theorizing has to do either
with evolutionary psychology or with how
the law actually operates on the ground.
The answer is that jurisprudence and evolu-
tionary psychology are both concerned with
human nature. As evolutionary psychology
is impacting our fundamental view of what
it means to be human, those impacts portend
important consequences in many areas of the
223
law. The one we consider here is criminal
sentencing.
The debates between the four schools of
jurisprudence in some ways echo, and in
other ways are distinct from, the justifying
theories that underlie criminal punishment.
There are four main justifications for crimi-
nal punishment: retribution, rehabilitation,
deterrence, and incapacitation (Alschuler,
2003; Davis, 2009).
Retributivists generally believe that pun-
ishment is its own categorical good, and there-
fore it need not accomplish, or be validated
by, any external effects. Hence, the other
three theories, in contrast to retribution, are
sometimes lumped together as ‘utilitarian’.
Retribution is the oldest punishment theory,
though its formalization is generally attrib-
uted to the German philosopher Immanuel
Kant (1797). One can hear legal naturalism
within retribution. Precisely because there
is a natural core of right and wrong, punish-
ment is as much a part of that a priori core as
the right and wrong itself. But there are also
positivist and realist currents in retribution.
Kant’s intellectual successor, Georg Hegel,
once wrote that criminals must be punished
simply to earn their way back into the social
fold. Crime is a breach of the social contract,
which requires that the breacher pay damages
in the form of suffering (Hegel, 1820).
Deterrence was the first utilitarian rebel-
lion against retribution. Utilitarians like
Cesare Beccaria (1766) and Jeremy Bentham
(1830) believed that no social group had
the right to inflict punishment on its mem-
bers unless that punishment resulted in a net
social good. For proponents of deterrence,
punishing wrongdoers is permissible because
it deters both the person being punished (spe-
cial deterrence) as well as others (general
deterrence) from committing future crimes.
Early proponents of deterrence focused on
special deterrence. Bentham once famously
asserted that no state had the right to pun-
ish any criminal, even a murderer, if it could
be certain the criminal would never com-
mit another crime (Bentham, 1830). But of
224 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
course, punishment that might not deter the
wrongdoer may nevertheless deter millions
of others, and thus be a net good. Modern
proponents of deterrence therefore tend to
focus on general deterrence rather than spe-
cial deterrence. Both kinds are an amalgam
of different jurisprudential schools. Like
the realists and normativists, the deterrence
school of punishment is focused on the way
law accomplishes, or fails to accomplish, its
central purpose of deterring crime.
Rehabilitation is deterrence writ small. It
focuses not on deterring the population as a
whole, or even the particular criminal being
sentenced, but rather on treating criminals to
make them better persons who are less likely
to commit future crimes. Rehabilitation
became pre-eminent in US law at the begin-
ning of the 20th century, as progressives
viewed crime as mostly a product of failed
social systems, and criminals as socially dis-
eased and in need of a cure. It began to fall
out of favor in the late 1960s (Allen, 1978;
Alschuler, 2003).
Incapacitation is the most recent justifica-
tion for criminal punishment, at least when
it comes to non-capital crimes. It ascended
as a theory of punishment beginning in
the 1960s as rehabilitation began to wane.
Incapacitation, like special deterrence and
rehabilitation, is focused not on the popula-
tion as a whole but on the individual criminal.
Incapacitationists believe the primary pur-
pose of punishment is to remove criminals
from society so that they do not commit more
crimes. Unlike the other three theories of
punishment, which attempt in different ways
to remove people’s desire to commit future
crimes, incapacitation is designed to remove
their ability.
These theories had important, real-life
effects on the way we have treated criminals.
When retribution was king, sentences in the
United States were generally, and perhaps
surprisingly, quite mild in length. Indeed, the
Quakers invented the penitentiary in the late
1700s as a more merciful alternative to death
and banishment, which were the punishments
for most serious crimes. Penitentiaries were
not originally intended to be criminal ware-
houses. They were a place for criminals to
contemplate their crimes and futures – to be
penitent. Prison sentences for non-capital
crimes in these early years were extraordinar-
ily mild by modern standards – months rather
than years.
When rehabilitation ascended at the turn
of the century these mild sentences became
quite harsh. Cures took time. The length of
prison sentences skyrocketed. Deterrence
and incapacitation theories, especially promi-
nent in the 1970s as rehabilitation fell out of
favor, further boosted sentence lengths.
These theories have all survived to some
extent. Today, in all federal and virtually all
state courts, judges are expressly directed to
consider all of them when imposing a sen-
tence. The federal statute, 18 U.S.C. § 3553(a),
is typical. It provides that federal judges
shall consider … the need for the sentence
imposed:
a. to reflect the seriousness of the offense, to
promote respect for the law, and to provide for
just punishment for the offense [retribution];
b. to afford adequate deterrence to criminals
[general deterrence];
c. to protect the public from further crimes of
the defendant [special deterrence and
incapacitation];
d. to provide the defendant with needed educa-
tional or vocational training, medical care, or
other correctional treatment in the most effec-
tive manner [rehabilitation].
The idea of this kitchen-sink approach is
undoubtedly that different kinds of cases will
tend to command different punishment goals.
But how ought we resolve conflicts between
these goals? For example, how should we
deal with an offender who is morally culpa-
ble but not dangerous? Or dangerous but not
morally culpable? The theories don’t tell us
when one kind of goal should predominate
over another. And if we need to consider all
four goals, as most statutes command, none
of the theories explains how judges are to
integrate the theories’ incompatible goals.
 EVOLUTIONARY PSYCHOLOGY, JURISPRUDENCE, AND SENTENCING
If we punish only to deter, then why do we
often punish serious wrongdoers who have
low recidivism rates (murderers, for exam-
ple) more than wrongdoers with astronomi-
cal recidivism rates (forgers, for example)?
Similarly, if the main goal of punishment
is to rehabilitate, then why do we punish
Bentham’s already rehabilitated killer at all?
If we only punish to incapacitate, then why
doesn’t every criminal get a life sentence,
or a sentence gauged only to the risks of his
recidivating? It seems like something else,
something non-utilitarian, is animating our
actual sentencing practices.
There’s a wonderful thought experiment
(Alschuler, 2003) that sheds light on the
incompleteness of all the utilitarian theories
and hints that our retributivist urges run very
deep and need to be acknowledged. Imagine
a society in which all first-degree murder-
ers are executed as part of the halftime show
at the Super Bowl. The method of execu-
tion is a laser beam that we are told inflicts
unimaginably excruciating pain on the pris-
oner for several minutes before he is termi-
nally vaporized. But unbeknownst to us, the
‘killer’ beam is really a painless transporter
beam, and it sends the prisoners to an idyl-
lic island in the South Seas where they live
out their lives in luxury but can never return.
This system should be fine with true utilitar-
ians. Deterrence is maximized, the murder-
ers are rehabilitated (at least in the sense of
being sent to a perfect society without social
wants), and they are likewise incapacitated
from ever hurting the rest of us again. But
there is something deeply unsettling about
such an approach. It is not fair. A retributivist
would say that the murderers are not getting
their just deserts (and in fact they are being
rewarded for their heinous crimes).
But retribution comes with its own incom-
pleteness issues. How much desert is just, and
why? Even more fundamentally, retribution
seems to be built on the same unsatisfactory
a priori sands as natural law. Philosophers
might say that retribution has no antecedent
moral justifications. It just is, like gravity.
225
The evolutionary insights discussed in
Section II below address these central criti-
cisms of retribution: our core moral intuitions
evolved, and one of those intuitions is to punish
serious violations of the other moral intuitions.
II. SCIENTIFIC PERSPECTIVES ON
THE ULTIMATE AND PROXIMATE
FUNCTIONS OF PUNISHMENT
To predict where a storm front is heading, it
helps to understand its initial conditions. The
same is true for human nature. According to
the perspective of evolutionary psychology,
human behavior is enabled by brains that
solved ecological problems – problems such as
finding and maintaining good cooperation
partners – favored by natural selection. So
consideration of the problems that our brains
evolved to solve can help to explain and pre-
dict the motivations that shape our social insti-
tutions, including, perhaps, the justice system.
It is through this lens that it makes sense
to ask what an organism would stand to gain
from being punitive. Punishing other peo-
ple is costly. At minimum, it takes time and
energy, and it exposes the punisher to the
risk of retaliation. So, unless a given punish-
ment strategy tended to confer proportionally
large benefits to the punisher, such a strategy
would be unlikely to have evolved by natural
selection. Yet punishment has been a ubiqui-
tous part of human society (Brown, 1991). A
growing body of empirical scholarship has
thus sought to identify the potential benefits
that common punishment behaviors are, in an
ultimate sense, designed to procure. Here, we
discuss some key results from this scholarship
as well as their added value for our ancestors
and possibly for modern legal institutions.
Second-Party Punishment
Among the various forms of punishment that
occurs in human societies, the simplest is
226 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
second-party punishment: when a victim
retaliates against the victimizer. Retaliation
is risky business. Yet it’s a common response
to cheating and aggression in humans and
even some non-human animals (Clutton-
Brock and Parker, 1995; Jensen et al., 2007).
So, what are the benefits to be gained by
second-party punishment? The evolutionary
psychology literature provides at least three
notable answers.
First, retaliation, if deployed effectively,
provides self-defense. It can serve as a precau-
tionary strategy for managing a hazard in the
environment (Fiddick et al., 2000), not unlike
our instinct to shut our eyes in a dust storm.
In this way, one need not posit the existence
of an evolved psychology of social interac-
tion because our need for self-preservation
extends beyond the social. Indeed, there are
many examples of self-defense across animal
species, social and non-social alike, includ-
ing algae (Paul and Fenical, 1986) and gup-
pies (Godin and Davis, 1995). The most
fundamental form of self-defense may be
the immunological response (Hoffman and
Krueger, 2017). Clearly, at least some of our
evolved psychology may support retaliatory
behavior, not to interface with the cheater’s
motivations or perceptions, but more simply
to protect oneself from harm.
Another potential benefit to be gleaned
from retaliating could be to gain a direct-­
fitness advantage over the cheater. An impulse
to ostracize, kill, or otherwise debilitate the
cheater could directly advance one’s own
prospects, if deployed successfully (Trivers,
1971). However, since such actions invite
dangerous countermeasures and thwart the
possibility of future cooperation, this strat-
egy would seem to be limited to situations in
which continued victimization is assessed to
be highly likely but the likelihood of mutual
cooperation is low. In the environment in
which our punishment psychology evolved,
situations like this would have been more
common when the offender was a member of
a competing coalition rather than a member
of one’s own social group.
A third way in which a victim could b­ enefit
from retaliation is by transforming the cheater
into a valuable cooperation partner. This char-
acterization is known as the theory of direct
reciprocity, proposed by Trivers, and there
is considerable evidence for its operation in
both human and non-human species (Trivers,
1971). For this strategy to be effective, the
imposition of costs on the cheater would
have to ‘educate’ the cheater by altering his
incentive structure for how he conducts future
interactions with the punisher. In other words,
the punishment, or at least a credible threat
of punishment, must appeal to his psychol-
ogy of deterrence. If successful, this strategy
not only protects the punisher from further
exploitation but also enables reciprocal gains
in trade between the dyad via increased coop-
eration. Thus, this strategy would commonly
target members of one’s own existing social
network. So, while retaliation might be risky,
it can also pay dividends.
Third-Party Punishment
Third-party punishment:
why bother?
Human punishment behavior, of course, is
not limited to two-party contexts. Across his-
tory and across cultures, punishment by
ostensibly independent ‘third’ parties is per-
vasive. In the ancestral environment, third-
party punishment would have been delivered
by members of the victim’s broader commu-
nity, usually small coalitions of men who,
together, could deliver punishment at lesser
risk to any one individual. In modern socie-
ties, such activities have been largely dele-
gated to social institutions like the criminal
justice system, which empower disinterested
triers of fact, like judges and jurors, to deliver
third-party punishment.
The fact that punishment is so costly is
particularly problematic for evolutionary
theories of third-party punishment since third
parties are defined as people who are not
involved in the offense and so do not stand
 EVOLUTIONARY PSYCHOLOGY, JURISPRUDENCE, AND SENTENCING
to gain directly from the punishment. Why
should they bother? The answer may lie in
the fact that they are not really independent
parties after all. Supposed third parties may
have implicit social motivations driving their
punitive attitudes even if they do not have
conscious access to those motivations. This is
because, being human, their minds evolved in
a hyper-social context. People with traits that
enabled them to exploit the social market-
place gained a fitness advantage. So, it is rel-
evant to ask, in an ultimate sense, how these
so-called third parties could stand to gain
from particular types of punishment behav-
ior, and how these gains might undergird
modern legal justifications for punishment.
A rich body of literature provides some
empirical answers to this question. The first is
kin selection, which predicts altruism toward
genetic relatives. As expressed by the highly
influential Hamilton’s Rule, the degree of
relatedness (given the modest assumption
that our ancestors could recognize their kin
with some accuracy) predicts the degree
of altruism because such altruistic behav-
iors increase the replication of our genes
(Hamilton, 1964). Under these conditions,
our tolerance for risk should increase – a
prediction well-captured by the quip cred-
ited to biologist J. B. S. Haldane that he’d
be prepared to die for two brothers or eight
cousins (Connolly and Martlew, 1999: 10).
Thus, costly punishments that deter or inca-
pacitate actors from cheating our kin should
be favored by natural selection. Indeed, evi-
dence for kin-based punishment has been
widely documented (Daly and Wilson, 1988).
Since kin-based punishment must effectively
prevent or discourage cheating in order to be
selected, it is compatible with the incapacita-
tive and deterrence-based philosophical justi-
fications for punishment.
Of course, punishments by judges and
jurors are not supposed to favor kin. Indeed,
the law specifically bars judges and jurors
from serving in such cases. So an additional
explanation is needed to explain third-party
punishment in non-kin contexts. One such
227
explanation is strong reciprocity. According
to this view, altruistic punishment can evolve
because groups of people who expressed
this trait deterred cheaters and, thus, enjoyed
more cooperation than those who did not
(Gintis, 2000). Support for this theory has
been argued on the basis of a series of eco-
nomic game studies. In these studies, partici-
pants who engaged in a repeated public-goods
game demonstrate a willingness to punish
third-party defectors at a cost to themselves,
and this behavior sustained a norm of coop-
eration across the group of players (Fehr
and Fischbacher, 2004; Fehr and Gächter,
2002; Fehr et  al., 2002). Since punishment
of this sort must effectively discourage the
cheater or other tempted onlookers in order
to be selected, this theory is compatible with
the legal justifications of special and gen-
eral deterrence. Despite its intuitive appeal,
scholars disagree about the viability of such
an altruistic strategy given that groups that
punish would be vulnerable to exploitation by
second-order cheaters, namely, people who
shirk their contribution to the enforcement
of punishment (Krasnow et al., 2015; Tooby
and Cosmides, 2016). There is a burden, crit-
ics argue, to explain how the enforcement of
punishment ever gained an evolutionary foot-
hold, given that it’s individually costly.
To explain how third-party punishment
could benefit the individual punisher, one
theory – social exchange theory – highlights
the implicit value of the cheater and victim to
the punisher. According to this theory, since
humans evolved in small social exchange
networks, most people that we encountered
on a day-to-day basis would have been net-
work members (i.e., direct resources to us),
and so we operate on assumptions that our
peers, and even most strangers, are poten-
tial exchange partners (Delton et  al., 2011;
Krasnow et al., 2013). Under this constraint,
if an actor cheats a victim, ‘third parties’
who punish the cheater gain a direct advan-
tage in social capital by deterring the cheater.
This hypothesis has received support from
multiple methods including experimental
228 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
surveys and economic games with real stakes
(Aharoni and Fridlund, 2013; Delton and
Krasnow, 2017; Krasnow et al., 2012, 2016;
Pedersen et al., 2018; Price et al., 2002). This
theory is a generalization of Trivers’ direct-
reciprocity theory (1971), and it is most com-
patible with the legal justification of special
deterrence.
Other efforts to explain how third-party
punishment could evolve at the individual
level point to the reputational benefits of pun-
ishing. Known as indirect reciprocity, this
theory states that by punishing a cheater, third
parties accrue reputational credit with the vic-
tim and other community members, such that
if the third party helps the victim by punishing
the cheater, someone else may help the third
party in the future (Nowak and Sigmund,
2005). People who are willing to enforce
costly punishment signal their commitment
to group norms, thereby broadly advertising
their own social status (Frank, 1988). Since
this strategy is designed to incentivize both
second and third parties in the network, it is
compatible with the legal justifications of
special and general deterrence.
Third-party Punishment:
how is it done?
Besides asking what a punisher might gain
by punishing a cheater, it would be valuable
to understand by what manner punishment
might achieve these ends. Most evolutionary
theories of punishment posit that punishment
increases the punisher’s fitness by reducing
recidivism, but they are sometimes hazy
about the proximate mechanism – how our
evolved punishment response reduces recidi-
vism. Here, we discuss some answers to this
question of mechanism.
From an evolutionary perspective, the most
direct way to gain a fitness advantage over a
cheater is by imposing corporal punishments
like injury or death, as we discussed in the
section on second-party punishment. This
strategy assumes a motivation to (temporarily
or permanently) behave in ways that incapac-
itate the cheater, but to achieve its aim, it does
not require a deterrence psychology – a folk
psychological theory of the cheater’s or other
group members’ motivations and incentive
structure. This makes the incapacitative strat-
egy cognitively efficient. However, this strat-
egy is costly, partly because it could forestall
a lifetime of future contributions by the for-
mer cheater to the social exchange commu-
nity. Therefore, it should have evolved to be
used as a last resort, and only when the costs
imposed by the cheater are expected to out-
weigh his lifetime contributions.
Another, potentially less risky, way to
limit the probability that the cheater will
cheat is to socially exclude, or ostracize, that
individual from the group. By removing his
opportunities to trade on the social market,
this also reduces his opportunities to exploit
cooperative group norms. In a general sense,
this action can also be understood as an inca-
pacitative strategy, but unlike corporal pun-
ishments, it may be easier to revoke. Thus, it
would have been a useful strategy in ances-
tral environments, which lacked correctional
institutions to perform the incapacitative
work.
Though incapacitation is a recognized
legal justification for punishment, social-
science research suggests that it fails to
capture a dominant feature of typical pun-
ishment psychology, namely, the desire to
inspire changes in the cheater’s mental state.
It is this feature of punishment psychology
that ultimately defines the deterrence motive.
The advantage of deterrence strategies over
other strategies like direct-fitness reduction
or ostracism is that, if effective, they imme-
diately restore gains in trade between the
former cheater and the social exchange net-
work because they convert the cheater into a
cooperator.
Different scholars have characterized this
process in slightly different ways. Some have
described deterrence as a form of coercion
(e.g., Matravers, 2000). The implication is
that, while it may be against the cheater’s
personal interest to shift to a more coop-
erative social strategy, the heavy threat of
 EVOLUTIONARY PSYCHOLOGY, JURISPRUDENCE, AND SENTENCING
punishment may demand it, and so through
a rational fear of that negative reinforcer, the
cheater is compelled to change his behavior,
against his deeper preferences. Certainly, as
an empirical matter, most people are respon-
sive to the threat of punishment, and so this
form of deterrence is likely to play a role in
crime and punishment psychology today.
But a moment’s reflection suggests that
there is something deeply unfulfilling about
a cheater who plays by the rules only because
of fear of punishment. Instead, we seem
to crave a genuine mental transformation:
we want the cheater to understand how he
harmed us, and to fully endorse and internal-
ize the values that guide our self-protective
social norms (Nahmias and Aharoni, 2017),
perhaps in no small part because when we
internalize norms, we can worry less about
catching norm violators. They catch them-
selves. Scholars have described this deter-
rence motive as a form of education or
recalibration, namely, to change how much
the cheater values our welfare relative to his
own (Petersen et  al., 2012; Trivers, 1971).
Consistent with this perspective, experimen-
tal research demonstrates that we care deeply
about whether the offender understands
why he’s being punished. For instance, in
an experimental stock-market game, par-
ticipants were more satisfied with their pun-
ishment of an ostensible cheater when they
received evidence that the cheater under-
stood why he was being punished and even
more satisfied when the cheater expressed
a commitment to stop cheating (Gollwitzer
and Denzler, 2009). Theoretically, it’s not
surprising that we would evolve to favor a
fundamental transformation in values over
a more simple, coerced obedience. This is
because coercion only works in the presence
of a credible enforcement threat, whereas a
recalibration of one’s values is self-enforcing.
In this sense, effective recalibration har-
nesses the power of first-party punishment,
which we discuss below.
If punishment is designed to deter cheat-
ers by recalibrating their value structures,
229
punishers should be sensitive to signals that
the cheater had an unfavorable value struc-
ture to begin with. This could explain why
punishers care so much about mens rea –
the mental states at the time of the offense
– such as criminal intent and knowledge of
the risk of harm (e.g., Aharoni and Fridlund,
2011; Cushman et al., 2013). A punitive sen-
timent that didn’t care about these mental
states would end up imposing heavy costs on
genuine cooperators who had a stroke of bad
luck, like a driver who caused a fatality as the
result of a brain seizure. Typically, we don’t
punish such individuals as harshly, if at all,
and one reason could be that unlucky peo-
ple who are otherwise morally innocent are
not as likely to repeatedly undermine coop-
erative norms in the future. The presence of
mental states like criminal intent, in contrast,
are prognostic of future recidivism. In other
words, a likely proximate function of punish-
ment is to impose ecologically rational con-
straints on who we should punish.
The premium that our deterrence psychol-
ogy places on the cheater’s mental states
reveals another lesson for the law: attempts
to deter a cheater by encouraging internali-
zation of norms would seem to qualify as a
form of rehabilitation. As we noted, standard
punishment theory treats deterrence and reha-
bilitation as separate motives for punishment.
But both motivations, if delivered effectively,
are designed to change behavior by inspiring
a change in the cheater’s personal goals and
values. Where they differ is that the deter-
rence motive places greater emphasis on the
role of coercion as a motivator for recalibra-
tion. Whether coercion is any more effective
a motivator is an empirical question.
If coercion simply represents the threat of
suffering, then as we discussed in the Super
Bowl thought experiment, many people
would be deeply unsatisfied by rehabilitation
sans suffering. But if we could achieve such
ends without the harmful means, then why do
we feel an obligation that the offender should
suffer? Why do we boil with outrage when
he does not? Stated differently, what evolved
230 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
purpose is served by the offender’s suffering
and the torrent of retributive impulses that
give rise to it? The answer brings us to our
final point about how our evolved punish-
ment psychology carries out its function: via
sophisticated motivational programs we call
emotions.
As economist Robert Frank (1988) force-
fully argued, emotions serve as commitment
devices: they are conspicuous, costly signals
that commit us to a particular course of action.
This is well illustrated by a classic anecdote
of a game of chicken played by two oppos-
ing drivers. If one driver, while revving his
engine, removes his steering wheel and drops
it out the window for his opponent to see,
that opponent must take seriously the first
driver’s commitment to winning. If retribu-
tive emotions are like our first driver, they
can be understood as part of a system that
is motivating us toward a particular punish-
ment strategy. The fact that these retributive
emotions are not sated until the offender suf-
fers suggests that the suffering may likewise
serve a motivational role for the offender,
namely to change his offensive behavior by
shifting the balance in how much he values
our welfare (Petersen et al., 2010). So, retrib-
utive emotions can be understood as mobi-
lizing punishment and facilitating offender
behavior change – functioning to keep both
parties honest and invested in long-term reci-
procity. The punisher, of course, need not be
consciously aware of the functional role of
these retributive emotions in order for them
to exert their effects.
Conceived in this way, the retributive
motive for punishment is not in opposition
to the utilitarian motives, as legal theory
implies. Rather, it is an expression of them.
In this view, we are psychological retribu-
tivists but adaptive utilitarians (Cushman,
2015). Retributive emotions commit us to
punishment – a costly signal of our status
that enhances the punisher’s fitness, either
directly or indirectly, by modifying the
cheater’s fitness, rational incentives, or social
values. In this view, the legal purposes of
punishment are not wholly incompatible –
they are just operating on different levels of
analysis. Thus, the application of evolution-
ary theory carries the potential to synthesize
the legally disparate theories for punishment.
First-Party Punishment:
Conscience, Guilt, and Shame
We’ve saved first-party punishment for last
because it might be the most complex, and
also because, in important ways, its effective-
ness depends on the other two types. First-
party punishment, as the phrase suggests,
could be loosely understood as the tendency
to ‘pre-punish’ ourselves with the bite of
conscience to help us refrain from violating
norms in the first place.2 Guilt and shame are
also versions of first-party punishment,
though they generally operate after we have
violated a norm, and serve the function of
reducing the chances of violating that norm
again. Of course, with an animal as smart and
devious as we humans, there is much overlap
between conscience, guilt, and shame. We
might feel guilty or ashamed even for con-
templating a wrong. Likewise, we might look
back on some of our decisions and wonder
why our conscience did or did not save us
from doing wrong.
The bite of conscience is not just a human
universal; it is a substantial part of self-image
across cultures. It is central to many religions
and mythologies (Katchadourian, 2009). It
was probably a critical solution to the prob-
lem of effectively deterring antisocial behav-
iors in our social networks. At the proximate
level of explanation, one of the reasons you
don’t punch your neighbor during a politi-
cal argument – perhaps the main reason – is
not that you are afraid he’ll punch you back or
that you will be punished by the criminal jus-
tice system, but rather because you know it is
wrong. Brains with built-in systems that make
their owners feel bad when they contemplate
cheating are brains that will not cheat as often.
This is what we mean when we said earlier
 EVOLUTIONARY PSYCHOLOGY, JURISPRUDENCE, AND SENTENCING
that an offender has recalibrated his value
structure or internalized a norm. Conscience
is the first line of defense against the com-
mission of antisocial behaviors, and at an
ultimate level, it worked in concert with the
other forms of punishment to deter individu-
als from undermining our ancestors’ precious
social exchange network. Presumably, it had
to work in concert with other forms of pun-
ishment because without an external threat of
punishment, there would be little incentive to
regulate oneself.
One way to appreciate how important
conscience is to our cooperative nature is to
study people who don’t seem to have a con-
science – psychopaths. Psychopaths tend to
have deficiencies in brain regions associated
with moral decision-making, including affec-
tive memory, inhibition, and empathy (Kiehl,
2007). Unsurprisingly, psychopaths make up
a hugely disproportionate segment of incar-
cerated adults, are notoriously resistant to
traditional punitive and rehabilitative treat-
ments, and therefore consume an astonishing
proportion of our criminal justice resources
(Kiehl and Hoffman, 2011). A growing body
of research has also begun to examine whether
certain psychopathic traits might themselves
contain evidence of adaptive design (Glenn
et al., 2011). Such insights would underscore
the need to investigate alternative approaches
to behavior modification that better account
for individual differences.
If conscience evolved to steer us away
from the commission of antisocial behavior,
then guilt and shame evolved to help us learn
from lapses in conscience. Research suggests
that the emotion of guilt and the rumination
that often accompanies it evolved to change
an undesirable pattern of behavior by recali-
brating the cheater’s relative utility functions
for alternative goal states – the same process
of recalibration discussed above. Similarly,
the emotion of shame evolved to signal to
others the cheater’s commitment to change
through public acts of compensation and pen-
ance (Nahmias and Aharoni, 2017; Sznycer,
2018; Trivers, 1971).
231
In a classic experiment by Wallace and
Sadulla (1966), individuals were led to
believe they broke an expensive machine,
and the transgression had either been discov-
ered or not. The investigators found that com-
pared to a control group who did not break
the machine, the participants whose trans-
gression was discovered were more likely to
volunteer for a separate, painful experiment.
Volunteering for a painful experiment, and
other shame displays, can be understood as
a costly signal of one’s commitment to the
group, that one has internalized the purpose
of punishment, and potentially that second-
or third-party punishment is unnecessary.
This pattern of behavior suggests that self-
governing psychological systems are in place
but may require social input to be triggered.
This dependency would make sense if our
self-governing systems coevolved with second-
and third-party punishment systems.
By implication, effective criminal punish-
ment practices might be those that utilize the
natural threat of public exposure to motivate
offenders to recalibrate their internal model
for how to treat others – or at least for how not
to get caught. Many legal practices already
exploit such properties, for instance, in pub-
lishing the names and addresses of local sex
offenders. But to maximize their effective-
ness, such practices should leverage rather
than subvert our intrinsic, self-regulatory
emotions. If so, such practices could qualify
under the law’s deterrence- and rehabilitation-
based justifications for punishment.
III. ANOMALOUS BYPRODUCTS OF
OUR EVOLVED PUNITIVE ATTITUDES
Though natural selection has fashioned
sophisticated punishment strategies designed
to protect the punisher’s interests and preserve
norms of cooperation, it doesn’t always do
this well. Natural selection produces relative
fitness advantages. It does not optimize – it
produces minimally viable solutions.
232 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Moreover, evolution by natural selection is
slow, which means that today’s adaptations
reflect fitness advantages with respect to
ancestral environments, not modern ones.
When a trait that evolved to solve a problem in
the environment in which it adapted behaves
differently in a more contemporary environ-
ment, it is known as an evolutionary mis-
match. This lack of fit between the adaptation
and its present environment can produce
behavioral phenotypes (i.e., byproducts) that
are over-sensitive to some evolutionarily novel
cues (as when we crave candy as a proxy for
fruit) and under-sensitive to others (as when
drivers underestimate fatally high travel
speeds). Likewise, the objectives and interests
of legal institutions and their stakeholders are
not necessarily aligned with those of our
hunter-gatherer ancestors, and there are plenti-
ful examples of punitive behaviors exhibiting
such over- and under-sensitivities. Here we
discuss these discrepancies as a potential
source of ‘subtractive value’ of human punish-
ment expressed in modern environments.
Over-Sensitivity to Legally
Irrelevant Factors
Several examples demonstrate how our puni-
tive instincts may be over-sensitive to factors
that are not legally relevant or efficacious.
For example, judges and jurors are highly
susceptible to the anchoring phenomenon.
One dramatic experiment showed that pro-
fessional judges who were asked to make
hypothetical prison-sentencing judgments
predictably increased or decreased their sen-
tence following a phone call from an ostensi-
ble journalist (an actor in the study) who
asked whether the sentence will be higher or
lower than ‘one’ vs ‘three’ years (Englich
et  al., 2006). Since judges know they ought
not take questions by the media, this ten-
dency to conform their sentence to the jour-
nalist’s suggestion presumably occurred at
an unconscious level. Such effects have been
explained by known psychological biases
such as the availability heuristic, the ten-
dency to place more weight on information
that happens to be most cognitively accessi-
ble in that moment (Tversky and Kahneman,
1973). Although judges wouldn’t normally
talk to journalists in the real world, similar
anchoring effects could be caused by pretrial
publicity generally.
Another example of our over-sensitivity to
legally irrelevant cues is the identifiable vic-
tim effect, which shows that people are more
likely to support punishment when they know
more about the victim – such as age or occu-
pation, for instance (Jenni and Loewenstein,
1997). Given natural limitations on our cog-
nitive resources, it makes sense that people
who prioritized the most immediate infor-
mation, such as who the specific victim
was, would tend to have an advantage in the
sorts of high-pressure social problems com-
monly faced by our ancestors. Of course, this
logic does not necessarily result in fair and
appropriate legal judgments. Arguably, not
all information that judges and jurors learn
about victims is strictly relevant to their judg-
ments. Since some judges and juries are inci-
dentally exposed to more information about
victims than others, one implication of this
research is whether attempts should be made
to limit and standardize what victim informa-
tion is and is not revealed to fact finders.
Some widely studied examples of our over-
sensitivity to legally irrelevant cues include
age, race, gender, and attractiveness biases.
Experimental simulation studies have shown
that defendants are more likely to receive
higher sentence recommendations when they
are portrayed as young, black, and male (e.g.,
Mitchell et  al., 2005; Steffensmeier et  al.,
1998; Sweeney and Haney, 1992). Archival
research has been less consistent, suggesting
that any real-life sentencing biases against
age, race, and gender may be small and highly
variable (Bontrager et  al., 2013; Mitchell,
2005; Wu and Spohn, 2009). To the extent
that these biases are real, however, findings
like these have been explained in proximate
terms including ingroup favoritism and halo
 EVOLUTIONARY PSYCHOLOGY, JURISPRUDENCE, AND SENTENCING
effects. But they are also consistent with more
ultimate explanations of social exchange.
According to these theories, in the environ-
ment in which our social brain evolved,
demographic characteristics may have served
as a conspicuous cue to group membership,
and people who were willing to pay the cost
to punish could reap direct rewards on trad-
ing markets by leveling more severe punish-
ments against groups and individuals with
whom they were most likely to compete for
physical and social resources (Aharoni and
Fridlund, 2013; Petersen et al., 2010, 2012).
Ultimately, for such punishment biases to
evolve, they would have had to be effective at
deterring or incapacitating the most competi-
tive opponents. And while the law takes pains
to try to remove such response patterns from
the courtroom, they often persist.
Another example of our over-sensitivity
to legally irrelevant cues concerns the dis-
proportionate emphasis we place on severity
as the preferred mechanism of punishment.
The two most studied parameters by which
a punisher can deter an offender are the
severity and the probability of punishment.
The intuition is that more severe punish-
ments deter better, but research has shown
that increases in severity have limited deter-
rent effect and diminishing marginal returns;
increasing the probability of punishment
would have a greater impact (e.g., Grogger,
1991; Paternoster and Iovanni, 1986). In the
ancestral environment, the situation was dif-
ferent. People lived in small groups with little
privacy, and so the looming threat of detec-
tion must have been real, and powerful. But
in the massive societies we live in today, the
probability that an offender is caught and
adjudicated remains, on average, relatively
low (Aharoni and Kiehl, 2013; Ehrlich,
1996), and offenders can exploit this fact.
Despite the evidence that severe sanctions
typically deter no better than moderate ones,
people seem to readily accept modulating the
severity of punishment as the primary instru-
ment through which to express their punitive
motives, consistent with the philosophical
233
principle of proportionality (see Robinson
and Kurzban, 2006). One reason, as we dis-
cussed above, may be our instinctive desire
to know that the offender will suffer because
suffering once was an effective means of
recalibration and long-term behavior change
(Nahmias and Aharoni, 2017).
Our punishment judgments are also sensi-
tive to information from our internal, physio-
logical environments. For example, research
has provided evidence that judicial sentenc-
ing decisions are less favorable immediately
following the shift to daylight savings time,
when people tend to be most fatigued (Cho
et al., 2017). Other research has shown a pat-
tern of sentence increases following unex-
pected home-team football losses (Eren and
Mocan, 2018), presumably by implicitly
impacting the judge’s mood. Unconscious
cognitive processes, such as those that regu-
late fatigue, stress, and mood, evolved as a
part of a motivational system designed to
prioritize our momentary physical and social
needs, so they might have global effects on
our decision-making, though it remains to be
seen whether such effects are large enough to
be decisive in court.
Even subject-matter experts are susceptible
to influence by extra-legal factors. The
testimony of expert witnesses, of course,
should be based on the principles of their
discipline, but research suggests that scientific
experts who engage in objective reasoning
problems are (unconsciously) prone to produce
evidence that confirms the preferences of the
hiring party (Robertson, 2010; Wong et  al.,
2015). One likely proximate explanation for
this ‘adversarial allegiance’ effect is socially
desirable responding, a form of motivated
reasoning. Socially desirable responding
probably had important social advantages for
our hunter-gatherer ancestors, but in a modern
trial context, these effects could also bias fact-
finders’ ascriptions of guilt and punishment.
Another extra-legal factor that we may be
sensitive to is how the costs and benefits of
punishment are presented. Although the cost
of incarcerating a defendant, for instance,
234 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
may or may not be important to judges and
taxpayers, incidental changes in the avail-
ability of that information should certainly
not drive sentencing attitudes. Yet recent
research suggests that when professional
judges and laypeople make sentencing rec-
ommendations for criminal offenders, they
are more lenient when the direct monetary
costs of incarceration are made salient, sug-
gesting that they find those costs to be rel-
evant but fail to spontaneously consider them
under typical, low-salience conditions. As a
consequence, removing the cost information
(without also removing benefit information)
may produce sentence length recommenda-
tions that systematically exceed those made
under more transparent conditions (Aharoni
et al., 2019; Rachlinski et al., 2012).
This behavior pattern implies that indi-
viduals’ punishment attitudes are internally
inconsistent across informational contexts,
placing more weight on whatever informa-
tion happens to be most readily available in
the moment. This tendency may be rooted in
a more general tendency to discount distant
future rewards relative to immediate ones
(Loewenstein and Thaler, 1989; Metcalfe and
Mischel, 1999; Mischel and Ebbesen, 1970).
Placing greater importance on the here and
now might have been adaptive for people in
need of quick results under conditions of very
limited information (see Daly and Wilson,
2005), but such features do not necessarily
promote the careful, disinterested, and judi-
cious decision-making that we expect of
modern criminal justice decisions. We don’t
mean to imply that more regulation is always
the best solution to reduce bias in the court-
room, but that a richer understanding of how
and why we punish can serve to clarify dis-
course on appropriate strategies.
Under-Sensitivity to Legally
Relevant Factors
There are also examples of our punitive
instincts operating in ways that are
under-sensitive to factors that may actually
be legally relevant. For one, we seem to read-
ily discount the value of modern, custodial
punishment strategies. Through the use of
police forces, prisons, and civil commitment
facilities, it is now possible to incapacitate
dangerous offenders with lower risk to vic-
tims, and without the need to impose suffer-
ing or coercive ultimata on the offender. But
as discussed, we often feel outrage at the
prospect of an offender spending his days in
a cushy rehabilitation center. And we demand
punishment of moral offenders even if they
are no longer dangerous (see Connolly,
2010). In such cases, though the modern
sanction may effectively satisfy the ultimate
objective of the punishment adaptation
(i.e., to reduce recidivism), it fails to meet the
adaptation’s more proximate conditions,
such as evidence that the offender has under-
gone mental recalibration by way of suffer-
ing and penitence. Such responses evolved in
environments in which effective custodial
incapacitative sanctions were unavailable –
they evolved not for incapacitation but for
deterrence. This could help explain public
dissatisfaction with purely custodial
sanctions.
From a public-safety perspective, in the
presence of effective incapacitation, the
offender’s attitudes about his punishment
should be irrelevant. But people still treat
them as relevant (Gollwitzer and Denzler,
2009) presumably because efforts to recali-
brate his attitudes would have been one of
the best strategies for reducing recidivism
in ancestral environments. For this reason,
we may be inclined to discount or altogether
reject recidivism-reduction strategies that
bypass opportunities for inducing suffering
and penitence. If so, whether our evolved
psychology of punishment (i.e., to achieve
deterrence via retribution) provides a good
justification for rejecting correctional sanc-
tions that are equally effective but more
humanitarian is a worthy normative ques-
tion, but not one that can be answered by the
science alone.
 EVOLUTIONARY PSYCHOLOGY, JURISPRUDENCE, AND SENTENCING
IV. THE UTILITY OF THE
EVOLUTIONARY PERSPECTIVE
From Explanation to Justification
Before we finish up with some speculations
about how the lens of evolutionary psychol-
ogy might clarify some issues in the criminal
law (and even a few issues beyond the crimi-
nal law), we need to address the problem of
Hume’s Gap, or what is also called the natu-
ralistic fallacy. In 1739, the Scottish philoso-
pher David Hume complained about how
there is often a gap between what is and what
some philosophers claim ought to be (Hume
et al., 1739 ). Just because behaviors happen
regularly, that does not mean they are mor-
ally grounded. Rapes and murders happen all
the time, but that doesn’t mean they are
acceptable under any jurisprudential theory of
law. In more modern philosophical parlance,
moral truths (‘the ought’) cannot be derived
solely from observational ones (‘the is’).
Evolutionary psychology has been a favorite
target of criticisms revolving around Hume’s
Gap (Rose and Rose, 2000). Just because kill-
ing a rival and kidnapping his mate may have
been an effective strategy for our ancestors
(Buss, 2005; Daly and Wilson, 1988), and
therefore our brains are arguably built with
motivations for these kinds of behaviors, evo-
lutionary psychologists are wary about taking
those ‘is’ observations into ‘ought’ realms like
the law, and quite properly so (Pinker, 2003).
But Hume himself never suggested the trip
between ‘is’ and ‘ought’ was unnavigable; he
was just complaining that moral philosophers
jumped the gap without so much as a pause.
Indeed, like virtually every other moral phi-
losopher of his time, Hume believed the
worlds of nature and morals were connected.
His connective tissue was empathy and
socialization: our own selfish desires to avoid
pain cause us to hesitate to inflict pain on oth-
ers, and that hesitation was Hume’s root of
morality (Hoffman, 2014).
We suggest that evolutionary psychol-
ogy itself has now supplied a more rigorous
235
connection between the ‘is’ and the ‘ought’.
Moral intuitions (including intuitions to pun-
ish) evolved precisely because they gave our
ancestors a net survival advantage in their
intensely social groups. Human punishment
behaviors are not just widespread by chance;
they are widespread because evolution has
armed us with emotions that make us feel
their moral bite. The boldest answer to the
jurisprudential question with which we began
this chapter – where does law come from? –
is not that it comes from God (as the tradi-
tional natural-law theorists argued), or from
social conventions (the positivists), or from
the individual and varying interpretations of
lawmakers and judges (the legal realists).
Instead, it comes from our evolved moral
intuitions. In this account of moral realism,
our notions of right and wrong come from
the same place as our opposable thumbs and
spines.
Understanding what goals our punitive
impulses evolved to achieve and how they
achieve them can help us evaluate the degree
of alignment with our modern societal goals,
and can help formulate hypotheses for how to
manage those impulses. For example, if deter-
rence is our main goal, we can ask whether
retribution is still the best way to achieve
deterrence. If so, lengthy prison terms might
not satisfy that goal as efficiently as lashings
and scarlet letters. Or if retribution is not the
best way to achieve deterrence, then it might
make more sense to remove offender suffer-
ing from the equation and focus instead on
evidence-based rehabilitation services. Such
tradeoffs would not be discoverable without a
theory of the function that retribution evolved
to serve.
Caution is still warranted. Many a moral
and legal judgment must be exercised in the
vast areas beyond and between our evolved
moral intuitions. These intuitions give us only
the broadest contours of a secular morality.
Even for those readers who reject our sug-
gestion that evolutionary psychology can
itself be a small bridge across Hume’s Gap,
it still has significant utility when examining
236 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
legal questions. We discuss three examples:
(1) synthesizing the various punishment
theories; (2) providing a framework for pre-
dicting behavior under uncertainty in ways
that might help us use law to leverage those
behaviors; and (3) helping to inspire practi-
cal legal reforms in circumstances where the
distance between the law and our evolved
behavioral predispositions is just too great to
leverage.
The application of evolutionary theory to
the law carries the potential to fundamentally
change how we think about legal justifications
for punishment and their apparent tensions.
As we have argued, our punitive sentiments,
and the sense of obligation that accompanies
them, evolved in part because they gave pun-
ishers a fitness advantage by deterring cheat-
ing behavior, thereby protecting their physical
resources and social exchange networks. The
ultimate benefit was deterrence, but the proxi-
mate mechanism to accomplish that deter-
rence was retribution. When a wrongdoer
revealed cues that he/she is not a good can-
didate for deterrence, other tactics were avail-
able, and these are reflected in sentiments like
the desire to ostracize, incapacitate, or reha-
bilitate. Our powerful desire to forgive and
be forgiven, about which there has developed
a significant behavioral- and evolutionary-
psychology literature (McCullough, 2008), is
another one of these proximate mechanisms
serving the ultimate advantage of reaping the
many benefits of cooperation.
Seen in this way, justice systems them-
selves can be understood as a type of
‘evolved’ system that is specialized to solve
problems of cheating and cooperation. The
notion that legal systems themselves evolved
by processes akin to natural selection remains
to be demonstrated, but whatever the mecha-
nism, they do appear equipped to increase
the scale by which group members can deter
cheaters. This could be achieved by minimiz-
ing the costs of enforcement to any individual
group member (Cushman, 2015).
Notably, goals like ‘deterrence’ take
on somewhat different meanings in the
evolutionary and legal accounts. The law
attempts to protect a much larger and less
well-defined social group than our punitive
adaptations were designed to serve. Because
of this mismatch, our evolved punitive psy-
chology cannot necessarily be relied on to
serve the explicit interests of society as a
whole, and conversely, pursuit of such social
goals may not necessarily satisfy our punitive
desires, such as when the law is more merci-
ful than what a victim demands. Evolutionary
science alone is not sufficient to help us rec-
oncile this tension, which hinges on norma-
tive judgments, but a deeper understanding of
our evolutionary legacy brings such tensions
to light.
From Explanation to Change
An evolutionary psychological perspective
can also help us understand the ultimate rea-
sons for our behavior, so we can more lucidly
decide whether these reasons are worth paying
for and whether there are other ways of
achieving our desired ends. By analogy,
knowing that a fever is an adaptive response
to infection can help us decide whether it
benefits us most to take a medication that
treats the fever symptoms or the underlying
infection. Fever, of course, is not a perfect
solution to overcoming an infection – in some
cases, there might be other, more effective
solutions – but, with some discomfort, fever
usually and eventually gets the job done
whereas treating the symptoms by suppress-
ing body temperature could actually make the
infection worse (Nesse and Williams, 2012).
On the other hand, there are some adaptations
that are clearly obsolete, like wisdom teeth
that crowd the modern jaw. Similarly, know-
ing that human punishment motivations
evolved to confer fitness advantages to mem-
bers of a social exchange network, and
knowing that it does this by employing
retributive-emotion programs to cause indi-
viduals to behave in ways that facilitate deter-
rence and desistance of the offensive action
 EVOLUTIONARY PSYCHOLOGY, JURISPRUDENCE, AND SENTENCING
can help us evaluate whether these are worthy
aims, and if so, whether our punitive motiva-
tions achieve these aims more or less effec-
tively than other strategies, such as particular
rehabilitative approaches that do not demand
suffering. Are retributive sentiments more
like a fever that, despite the discomfort,
works pretty well most of the time, or are
they more like an impacted wisdom tooth
that’s better off pulled? We don’t know the
answer to this question – whatever the
answer, it is undoubtedly complicated – but
it’s largely owing to evolutionary theory that
we can even articulate the question.
A richer understanding of the ultimate rea-
sons for human punishment behavior could
potentially inspire practical strategies for
managing unwanted effects. For example,
knowing that offenders can exploit relatively
anonymous lifestyles in modern societies,
reducing perceptions of anonymity in high-
crime areas (e.g., via increased surveillance)
could reduce crime, increase the quality of
evidence to support criminal charges, and
consequently make punishers less reliant
on the use of heavy sanctions to achieve
the same level of deterrence. Knowing that
decision-makers might neglect the costs of
the punishment unless those costs are made
salient could bolster arguments for increased
transparency about sentencing costs, includ-
ing recognition of other ways the funds could
be used (i.e., the opportunity costs of the
punishment). Knowing that sentencing judg-
ments may be primed by factors like fatigue
and mood, countermeasures could be devel-
oped, such as protocols for randomizing
defendants to docket schedules to ensure that
particular types of defendants (e.g., if a court
tends to schedule high-risk defendants last)
are not systematically penalized by time-
dependent effects.
The power of evolutionary theory to mobi-
lize the law in ways that more effectively
shape human behavior has been dubbed ‘the
law of law’s leverage’ (Jones, 2000). If we
think of law’s proscriptions and threats of
punishment as a lever by which society tries
237
to move its members away from antisocial
or other destructive behaviors, then under-
standing the evolutionary pressures (i.e., the
fulcrum) shaping those behaviors may tell us
how long our levers need to be. Behaviors
with deep evolutionary roots will be hard to
move except with big levers.
We’ve focused here on criminal sentenc-
ing, but evolutionary psychology might
impact other areas of the law. This is not to
say there is a one-to-one correspondence
between evolutionary theories and particular
legal reforms. The payoffs are not likely to
be so direct. Instead, their value will be in
constraining and inspiring plausible hypoth-
eses about legally relevant behaviors includ-
ing their triggers and inhibitors, by helping
to identify any evolutionary roots of such
behaviors. For instance, researchers have
already discovered that ordinary people (like
jurors) have significant trouble distinguishing
between two of the criminal law’s four mental
states – knowing and reckless (the other two
are purposeful and negligent; Ginther et al.,
2014; Shen et al., 2011). Examining the adap-
tive contours of mental-state attribution may
help judges improve their definitions of these
mental states. Understanding more about
how different kinds of substances or diseases
affect these mental states could also pay huge
dividends in reforming the law’s traditional
defenses to responsibility, including intoxica-
tion and insanity (Hoffman, 2018).
Jury instructions – the written rules
through which judges inform jurors about the
legal rules they must apply to the case before
them – are a fertile ground for evolutionary
psychology-based improvements. Knowing
how best to communicate complex principles
to ordinary citizens, and the pitfalls that lie
in wait, could substantially improve the truth-
finding function of jury trials.
Understanding more about human bias
may help judges and lawyers identify biased
jurors and/or inoculate them from the harsh-
est effects of those biases. Insights about our
irrational discounting behaviors could affect
the way in which judges instruct civil jurors,
238 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
and lawyers argue to them, about future tort
damages. Knowing that expert witnesses,
too, are susceptible to social-cognitive biases,
such as unconscious allegiance to the hiring
party, could inspire the development of meth-
ods for controlling the experts’ access to this
information, thereby improving the quality of
the testimony on which guilt and punishment
decisions may be based (Robertson, 2010;
Wong et al., 2015).
The potential for these insights to impact
the law is not limited to courts and juries. They
may even be more important at the front end of
law as legislators consider new laws and regu-
lators consider new rules and regulations. If
the difficulty in distinguishing between know-
ing criminal acts and reckless ones ends up
being insurmountable, informed legislatures
may want to consider reforming the criminal
law’s mental states. Likewise, as evolution-
arily informed researchers learn more about
those mental states, legislatures may consider
changing the contours of some of the classic
defenses to responsibility, such as intoxication
or insanity. Similarly, a working knowledge
of our time-discounting bias would certainly
be important as legislators and regulators con-
sider issues like usury rates, payday loans, and
caps on future tort damages.
Evolutionary psychology is likely to have
significant traction in all these areas of the law
because it informs our understanding of human
nature, and law is applied human nature.
ACKNOWLEDGMENTS
We thank the Cooperation, Conflict, and
Cognition lab members, especially Sharlene
Fernandes, Corey Allen, and Justin Thurman,
for helpful comments.
Notes
1  The first part of this section was adapted from
Hoffman (2018).
2  We use the term ‘first-party punishment’ loosely
since punishment in an ultimate sense implies
the delivery of costs that come at the expense
of the receiver, and it’s not clear that conscience,
guilt, and shame are best understood this way.
Another way to think of these mental states is
in proximate terms, as programs that regulate
how we conduct ourselves and treat others (see
Peterson et al., 2010).
REFERENCES
Aharoni, E., & Fridlund, A. J. (2011). Punishment
without reason: Isolating retribution in lay
punishment of criminal offenders.
Psychology, Public Policy, and the Law, 18(4),
599–625.
Aharoni, E., & Fridlund, A. J. (2013). Moralistic
punishment as a crude social insurance plan.
In T. Nadelhoffer (Ed.), The Future of
Punishment. New York, NY: Oxford University
Press (pp. 213–229).
Aharoni, E., & Kiehl, K. A. (2013). Evading
justice: Quantifying criminal success in
incarcerated psychopathic offenders. Criminal
Justice and Behavior, 40(6), 629–645.
Aharoni, E., Kleider-Offutt, H. M., Brosnan,
S. F., & Watzek, J. (2019). Justice at any cost?
The impact of cost/benefit salience on
criminal punishment judgments. Behavioral
Sciences & the Law, 37, 38–60.
Allen, F. A. (1978). The decline of the
rehabilitative ideal in American criminal
justice. Cleveland State Law Review, 27, 147.
Alschuler, A. W. (2003). The changing purposes
of criminal punishment: A retrospective on
the past century and some thought about
the next. University of Chicago Law Review,
70(1), 1–22.
Alschuler, A. W. (2000). Law without values:
The life, work and legacy of Justice Holmes.
Chicago, IL: University of Chicago Press.
Beccaria, C. (1766). On crimes and punishment.
Hackett (1986).
Bentham, J. (1830). The rationale for
punishment. London: R. Heward.
Bix, B. (2010). Natural law theory, 2nd ed.
In D. Patterson (Ed.), A Companion to
Philosophy of Law and Legal Theory. West
Sussex, UK: Wiley Blackwell (pp. 211–227).
 EVOLUTIONARY PSYCHOLOGY, JURISPRUDENCE, AND SENTENCING
Bontrager, S., Barrick, K., & Stupi, E. (2013).
Gender and sentencing: A meta-analysis of
contemporary research. Journal of Gender,
Race & Justice, 16, 349.
Brown, D. (1991). Human universals. New
York: McGraw-Hill.
Buss, D. M. (2005). The dangerous passion:
Why jealousy is as necessary as love and sex.
New York, NY: Odile Jacob.
Cho, K., Barnes, C. M., & Guanara, C. L.
(2017). Sleepy punishers are harsh punishers.
Psychological Science, 28(2), 242–247.
Clutton-Brock, T. H., & Parker, G. A. (1995).
Punishment in animal societies. Nature,
373(6511), 209.
Coleman, J. L., & Leiter, B. (2010). Legal
positivism. In D. Patterson (Ed.), A Companion
to Philosophy of Law and Legal Theory
(2nd ed.). West Sussex, UK: Wiley Blackwell
(pp. 228–249).
Connolly, K. (2010, July). Nazi death camp
guard charged over 430,000 Jewish deaths.
The Guardian. Retrieved on May 6, 2019 from
www.theguardian.com/world/2010/jul/29/
nazi-death-camp-holocaust
Connolly, K., & Martlew, M. (Eds.) (1999).
Altruism. Psychologically speaking: A book
of quotations. Leicester, UK: BPS Books.
Cushman, F. (2015). Punishment in humans:
From intuitions to institutions. Philosophy
Compass, 10(2), 117–133.
Cushman, F., Sheketoff, R., Wharton, S., &
Carey, S. (2013). The development of intent-
based moral judgment. Cognition, 127(1),
6–21.
Daly, M., & Wilson, M. (1988). Homicide. New
Brunswick, NJ: Transaction Publishers.
Daly, M., & Wilson, M. (2005). Carpe diem:
Adaptation and devaluing the future. The
Quarterly Review of Biology, 80(1), 55–60.
Davis, M. (2009). Punishment’s golden half
century: A survey of developments from
(about) 1957–2007. Journal of Ethics, 13(1),
73–100.
Delton, A. W., & Krasnow, M. M. (2017). The
psychology of deterrence explains why group
membership matters for third-party
punishment. Evolution and Human Behavior,
38(6), 734–743.
Delton, A. W., Krasnow, M. M., Cosmides, L., &
Tooby, J. (2011). Evolution of direct
reciprocity under uncertainty can explain
239
human generosity in one-shot encounters.
Proceedings of the National Academy of
Sciences, 108(32), 13335–13340.
Dworkin, R. (1986). Law’s empire. Cambridge,
MA: Harvard University Press.
Ehrlich, I. (1996). Crime, punishment, and the
market for offenses. Journal of Economic
Perspectives, 10(1), 43–67.
Englich, B., Mussweiler, T., & Strack, F. (2006).
Playing dice with criminal sentences: The
influence of irrelevant anchors on experts’
judicial decision making. Personality and
Social Psychology Bulletin, 32(2), 188–200.
Eren, O., & Mocan, N. (2018). Emotional
judges and unlucky juveniles. American
Economic Journal: Applied Economics, 10(3),
171–205.
Fehr, E., & Fischbacher, U. (2004). Third-party
punishment and social norms. Evolution and
Human Behavior, 25(2), 63–87.
Fehr, E., Fischbacher, U., & Gächter, S. (2002).
Strong reciprocity, human cooperation, and
the enforcement of social norms. Human
Nature, 13(1), 1–25.
Fehr, E., & Gächter, S. (2002). Altruistic
punishment in humans. Nature, 415(6868),
137.
Fiddick, L., Cosmides, L., & Tooby, J. (2000). No
interpretation without representation: The
role of domain-specific representations and
inferences in the Wason selection task.
Cognition, 77(1), 1–79.
Frank, R. H. (1988). Passions within reason: The
strategic role of the emotions. New York, NY:
W.W. Norton & Co.
Fuller, L. L. (1958). Positivism and fidelity to
law: a reply to Professor Hart. Harvard Law
Review, 71(4), 630–672.
Fuller, L. L. (1965). The morality of law (2nd
ed.). New Haven, CT: Yale University Press.
Gardner, J. (2001). Legal positivism: 5 1/2
myths. American Journal of Jurisprudence,
46, 199–227.
Ginther, M. R., Shen, F. X., Bonnie, R. J., &
Hoffman, M. B. (2014). The language
of mens rea. Vanderbilt Law Review, 67,
1327.
Gintis, H. (2000). Strong reciprocity and human
sociality. Journal of Theoretical Biology,
206(2), 169–179.
Glenn, A. L., Kurzban, R., & Raine, A. (2011).
Evolutionary theory and psychopathy.
240 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Aggression and Violent Behavior, 16(5),
371–380.
Godin, J., & Davis, S. (1995). Who dares,
benefits: Predator approach behaviour in the
guppy (Poecilia reticulata). Proceedings of
the Royal Society of London B, 259,
193–200.
Gollwitzer, M., & Denzler, M. (2009). What
makes revenge sweet: Seeing the offender
suffer or delivering a message? Journal of
Experimental Social Psychology, 45(4),
840–844.
Grogger, J. (1991). Certainty vs. severity of
punishment. Economic Inquiry, 29(2),
297–309.
Haakonssen, K. (1996). Natural law and moral
philosophy: From Grotius to the Scottish
Enlightenment. New York, NY: Cambridge
University Press.
Hamilton, W. D. (1964). The genetical evolution
of social behaviour. Journal of Theoretical
Biology, 7(1), 17–52.
Hart, H. L. A. (1958). Positivism and the
separation of law and morals. Harvard Law
Review, 71(4), 593–629.
Hart, H. L. A. (1961). The concept of law.
Oxford, UK: Clarendon Press.
Hegel, G. W. F. (1820). Philosophy of right 71
(T. M. Knox trans.). Oxford: Oxford University
Press (1942).
Hobbes, T. (1651). Leviathan. London, UK:
A&C Black (2006).
Hoffman, M. B. (2014). The punisher’s brain:
The evolution of judge and jury. New York,
NY: Cambridge University Press.
Hoffman, M. B. (2018). Nine neurolaw
predictions. New Criminal Law Review: An
International and Interdisciplinary Journal,
21(2), 212–246.
Hoffman, M. B., & Krueger, F. (2017). The neu-
roscience of blame and punishment.
In S. Menon (Ed.), Self, Culture and Con-
sciousness: Interdisciplinary Convergences
on Knowing and Being. Singapore: Springer
(pp. 207–223).
Hume, D. (1739). A treatise of human nature,
469. Oxford: Clarendon Press (1906).
Jenni, K., & Loewenstein, G. (1997). Explaining
the identifiable victim effect. Journal of Risk
and Uncertainty, 14(3), 235–257.
Jensen, K., Call, J., & Tomasello, M. (2007).
Chimpanzees are vengeful but not spiteful.
Proceedings of the National Academy of
Sciences, 104(32), 13046–13050.
Jones, O. D. (2000). Time-shifted rationality
and the Law of Law’s Leverage: Behavioral
economics meets behavioral biology.
Northwestern University Law Review, 95,
1141.
Kant, I. (1797). The philosophy of law: An
exposition of the fundamental principles
of jurisprudence as the science of right
(W. Hastie trans.). Edinburgh: T & T Clark
1887). Retrieved from https://oll.libertyfund.
org/titles/kant-the-philosophy-of-law
Katchadourian, H. (2009). Guilt: The bite of
conscience. Stanford, CA: Stanford University
Press.
Kiehl, K. A. (2007). Without morals: The
cognitive neuroscience of psychopathy.
In W. Sinnott-Armstrong (Ed.), Moral
Psychology (Vol. 3): The Neuroscience of
Morality: Emotion, Brain Disorders and
Development. Cambridge, MA: MIT Press.
Kiehl, K. A., & Hoffman, M. B. (2011). The
criminal psychopath: History, neuroscience,
treatment, and economics. Jurimetrics, 4(1),
355–397.
Krasnow, M. M., Cosmides, L., Pedersen, E. J.,
& Tooby, J. (2012). What are punishment
and reputation for? PLOS ONE, 7(9), e45662.
Krasnow, M. M., Delton, A. W., Cosmides, L. &
Tooby, J. (2015). Group cooperation without
group selection: Modest punishment can
recruit much cooperation. PLOS ONE, 10(4),
e0124561. doi:10.1371/journal.pone.
0124561
Krasnow, M. M., Delton, A. W., Cosmides, L., &
Tooby, J. (2016). Looking under the hood of
third-party punishment reveals design for
personal benefit. Psychological Science,
27(3), 405–418.
Krasnow, M. M., Delton, A. W., Tooby, J., &
Cosmides, L. (2013). Meeting now suggests
we will meet again: Implications for debates
on the evolution of cooperation. Scientific
Reports, 3, 1747.
Leiter, B. (2010). American legal realism. In D.
Patterson (Ed.), A Companion to Philosophy
of Law and Legal Theory (2nd ed.). West
Sussex, UK: Wiley Blackwell (pp. 249–266).
Loewenstein, G., & Thaler, R. H. (1989).
Anomalies: Intertemporal choice. Journal of
Economic Perspectives, 3(4), 181–193.
 EVOLUTIONARY PSYCHOLOGY, JURISPRUDENCE, AND SENTENCING
Matravers, M. (2000). Justice and punishment:
The rationale of coercion. Oxford University
Press on Demand.
McCullough, M. (2008). Beyond revenge: The
evolution of the forgiveness instinct. New
York, NY: Jossey-Bass.
Metcalfe, J., & Mischel, W. (1999). A hot/cool-
system analysis of delay of gratification:
Dynamics of willpower. Psychological Review,
106(1), 3.
Mischel, W., & Ebbesen, E. B. (1970). Attention
in delay of gratification. Journal of Personality
and Social Psychology, 16(2), 329.
Mitchell, O. (2005). A meta-analysis of race
and sentencing research: Explaining the
inconsistencies. Journal of Quantitative
Criminology, 21(4), 439–466.
Mitchell, T. L., Haw, R. M., Pfeifer, J. E., &
Meissner, C. A. (2005). Racial bias in mock
juror decision-making: A meta-analytic
review of defendant treatment. Law and
Human Behavior, 29(6), 621–637.
Nahmias, E., & Aharoni, E. (2017).
Communicative theories of punishment and
the impact of apology. In C. W. Surprenant
(Ed.), Rethinking Punishment in an Era of
Mass Incarceration. New York, NY: Routledge
(pp. 144–161).
Nesse, R. M., & Williams, G. C. (2012). Why we
get sick: The new science of Darwinian
medicine. New York, NY: Vintage Books.
Nowak, M. A., & Sigmund, K. (2005). Evolution
of indirect reciprocity. Nature, 437(7063),
1291.
Paternoster, R., & Iovanni, L. (1986). The
deterrent effect of perceived severity: A
reexamination. Social Forces, 64(3),
751–777.
Paul, V. J., & Fenical, W. (1986). Chemical
defense in tropical green algae, order
Caulerpales. Marine Ecology Progress Series,
34(1–2), 157–169.
Pedersen, E. J., McAuliffe, W. H., & McCullough,
M. E. (2018). The unresponsive avenger:
More evidence that disinterested third parties
do not punish altruistically. Journal of
Experimental Psychology: General, 147(4),
514.
Petersen, M. B., Sell, A., Tooby, J., & Cosmides, L.
(2010). Evolutionary psychology and criminal
justice: A recalibrational theory of punishment
and reconciliation. In H. Høgh-Oleson (Ed.),
241
Human Morality & Sociality: Evolutionary &
Comparative Perspectives. New York, NY:
Palgrave Macmillan.
Petersen, M. B., Sell, A., Tooby, J., & Cosmides,
L. (2012). To punish or repair? Evolutionary
psychology and lay intuitions about modern
criminal justice. Evolution and Human
Behavior, 33(6), 682–695.
Pinker, S. (2003). The blank slate: The modern
denial of human nature. New York, NY:
Penguin.
Plato (380 BCE). The Republic, 272–297 (B.
Jowett trans.). Seattle, WA: Amazon Classics
(2017).
Price, M. E., Cosmides, L., & Tooby, J. (2002).
Punitive sentiment as an anti-free rider
psychological device. Evolution and Human
Behavior, 23(3), 203–231.
Rachlinski, J. J., Wistrich, A. J., & Guthrie, C.
(2012). Altering attention in adjudication.
UCLA Law Review, 60, 1586.
Robertson, C. T. (2010). Blind expertise.
New York University Law Review, 85,
174–257.
Robinson, P. H., & Kurzban, R. (2006).
Concordance and conflict in intuitions of
justice. Minnesota Law Review, 91, 1829.
Rose, H., & Rose, S. (Eds.) (2000). Alas, poor
Darwin: Arguments against evolutionary
psychology. New York, NY: Harmony Books.
Shen, F. X., Hoffman, M. B., Jones, O. D., &
Greene, J. D. (2011). Sorting guilty minds.
New York University Law Review, 86, 1306.
Shiner, R. A. (2010). Law and its normativity. In
D. Patterson (Ed.), A Companion to Philosophy
of Law and Legal Theory (2nd ed.). Sussex,
UK: Wiley Blackwell (pp. 417–445).
Steffensmeier, D., Ulmer, J., & Kramer, J.
(1998). The interaction of race, gender, and
age in criminal sentencing: The punishment
cost of being young, black, and male.
Criminology, 36(4), 763–798.
Sweeney, L. T., & Haney, C. (1992). The
influence of race on sentencing: A meta-
analytic review of experimental studies.
Behavioral Sciences & the Law, 10(2),
179–195.
Sznycer, D. (2018). Forms and functions of the
self-conscious emotions. Trends in Cognitive
Sciences, 23(2), 143–157.
Tooby, J., & Cosmides, L. (2016). Human
cooperation shows the distinctive signatures
242 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
of adaptations to small-scale social life. [Peer
commentary on ‘Cultural group selection
plays an essential role in explaining human
cooperation: A sketch of the evidence’ by
P. Richerson et  al.]. Behavioral and Brain
Sciences, 39, 42–43. doi:10.1017/
S0140525X15000266
Trivers, R. L. (1971). The evolution of reciprocal
altruism. The Quarterly Review of Biology,
46(1), 35–57.
Tversky, A., & Kahneman, D. (1973). Availabil-
ity: A heuristic for judging frequency and
probability. Cognitive Psychology, 5(2),
207–232. https://doi.org/10.1016/0010-
0285(73)90033-9
Wallace, J., & Sadalla, E. (1966). Behavioral
consequences of transgression: I. The effects
of social recognition. Journal of Experimental
Research in Personality, 1(3), 187–194.
Wong, C., Aharoni, E., Aliev, G., & DuBois, J.
(2015). Blind collaborative justice: Testing
the impact of expert blinding and consensus
building on the validity of expert testimony.
RAND Report RR-804-1-NIJ. Retrieved on
May, 18, 2020 from www.rand.org/pubs/
research_reports/RR804-1.html
Wu, J., & Spohn, C. (2009). Does an offender’s
age have an effect on sentence length? A
meta-analytic review. Criminal Justice Policy
Review, 20(4), 379–413.

Evolutionary Psychology
and Incarceration
INTRODUCTION
Literature in the field of criminal justice
­frequently places incarceration (i.e. the state
of being confined in prison) in the context of
the normative nature of human activities and
institutions (Ward and Durrant, 2011). A
recent analysis of incarceration systems at
the international level (Mauer, 2017) points
out that a nation’s incarceration rate is often
interpreted as the degree of civilization of
that nation, as well as the degree of punitive-
ness the nation is willing to impose in the
process of isolating specific members from
the broader community (i.e. in the direction
of public safety). While several forms and
levels of imprisonment exist, incarceration is
often considered the end product of individ-
ual or societal failure (Mauer, 2017). In line
with this consideration, international conven-
tions and policies, such as the European
Convention on Human Rights, state that no
individual shall be deprived of liberty unless
12
Alina Simona Rusu
certain conditions are met, such as the neces-
sity to prevent the committing of an offence
or fleeing after having done so (Macovei,
2004; Mauer, 2017).
Analyses of criminal justice systems
across human societies indicate that incar-
ceration today involves not only the isolation
of individuals, but also their rehabilitation
through professionally designed interven-
tions and educational programs. Features of
violations of social rules are considered to
meet the input conditions for the develop-
ment of mechanisms designed to respond to
inter-individual exploitation (Petersen et al.,
2010, 2012). Hence, criminal justice insti-
tutions can be interpreted in the context of
evolved counter-exploitation strategies. Two
categories of counter-exploitation strate-
gies (i.e. punishment and rehabilitation) are
addressed in this chapter in the context of
evolutionary analyses of incarceration from
the perspectives of incapacitation of offenders
and of reparative interventions.
244 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
INTER-INDIVIDUAL EXPLOITATION
AND COUNTER-EXPLOITATION
STRATEGIES
Interactions with conspecifics in social spe-
cies afford not only fitness opportunities but
also potential costs, which can sometimes
include even the death of individuals sub-
jected to severe violence (e.g., Kurzban and
Leary, 2001). Acts of exploiting conspecifics
for self-benefits related to survival and repro-
duction have been documented in many
social species, including humans, as well as
evolved strategies designed to counter these
exploitation efforts (e.g., Duntley and Buss,
2004; Petersen et  al., 2012; McCullough
et al., 2013). Punishment and behaviors that
resemble human social exclusion, i.e., pre-
venting particular individuals from interact-
ing with the group, have been studied in
nonhuman species, such as lemurs, chimpan-
zees, and three-spined sticklebacks (e.g.,
Wilson, 1980; Goodall, 1986; Wrangham,
1987; Kurzban and Leary, 2001). Kurzban
and Leary (2001) interpret these types of
behaviors in nonhumans and humans within
the frame of discriminate sociality, noting
different forms that have emerged due to dif-
ferent selection pressures.
Several authors present and discuss several
categories of counter-exploitation strategies,
from the incapacitation of offenders to the
combination of incapacitation and rehabili-
tation, as well as the development of prison
settings with respect to the needs and wel-
fare of human beings. Petersen et al. (2012)
point out that the evolutionary literature on
exploitation and its application to modern
justice should take into account the counter-
exploitation strategies beyond punishment
and note that the small-scale social world of
our ancestors – with dense social networks
and high levels of dependency – should have
selected not only for punitive strategies,
but also for non-punitive reparative ones
(Aureli and de Waal, 2000; Petersen et  al.,
2010, 2012).
THE RECALIBRATION THEORY OF
COUNTER-EXPLOITATION
Starting from the idea that a major factor
regulating the activation of reparative – rather
than punitive – responses to exploitation/rule
violation is the perceived social value of the
perpetrator (the criminal’s social worth), an
explanatory frame has recently emerged, i.e.
the recalibration theory of counter-exploitation
(Petersen et al., 2010). The authors offer the
evolutionary prediction that the human mind
spontaneously computes the magnitude of
two distinct variables when confronted with
exploitation: (1) the exploitation’s serious-
ness and (2) the exploiter’s association value;
these variables are fed into motivational
mechanisms regulating distinct aspects of
strategies for countering exploitation (e.g.,
how severely we want to punish, how long
we wish to incapacitate the perpetrator, how
intense the social repair efforts will have to
be; Petersen et al., 2012).
Recent research demonstrates that the social
decisions mentioned above depend upon the
magnitude of an internal variable – a welfare
tradeoff ratio (WTR) – which sets the weight
the actor places on a specific person’s welfare
relative to the actor’s own (Tooby et al., 2006;
Tooby and Cosmides, 2008). Within this frame-
work, Petersen et al. (2012) define exploitation
as acts expressing too low a WTR (relative to
some baseline) by inflicting a cost on the tar-
get for too small a benefit to oneself. From the
perspective of this definition, evolution should
have selected for counter-exploitation strate-
gies designed to recalibrate the exploiter’s
WTR in the direction of decreasing the num-
ber of exploitive acts they commit in the future
(Sell et al., 2009; Petersen et al., 2012) and of
inducing the exploiter to place greater weight
on the welfare of others in the future (e.g.,
Clutton-Brock and Parker, 1995; de Waal,
1996, cited in Petersen et al., 2012).
The efficacy of the reparatory functions of
punishment appears to depend upon the abil-
ity of the punisher to monitor the exploiter’s
 EVOLUTIONARY PSYCHOLOGY AND INCARCERATION
behavior (Petersen et  al., 2012). From this
perspective, prisons might have evolved as
systems for monitoring the behavior of the
exploiter, but also a way for the offender and
for the society (‘the collective punisher’)
to interact through reconciliation and other
forms of reparative gestures.
The Exploitation’s Seriousness
Petersen et  al. (2012) propose that the differ-
ence between the minimally acceptable WTR
and the WTR expressed toward the victim
reflects the intuitive concept of an exploitive
act’s seriousness. In light of recalibration
theory, the information on the seriousness of an
offense should be reflected in the intensity of
the reaction to regulate the WTR of the perpe-
trator toward potential victims (i.e. punitive or
rehabilitative reaction). Petersen et al. summa-
rize from an evolutionary point of view the
criminological literature addressing the seri-
ousness of crime. It appears that the seriousness
of crime is set by the crime’s physical or sym-
bolic level of harm and that the rank orderings
of the seriousness of crimes are stable across
cultures, especially with regard to crimes caus-
ing physical harm (Stylianou, 2003; cited in
Petersen et al., 2012). Petersen et al. point out
that, although less explored, there are data sug-
gesting that the same crime is seen as less seri-
ous when performed for a large gain in inclusive
fitness, such as stealing food for one’s family,
defending relatives, etc., than for a small indi-
vidual gain (Rossi et  al., 1985). The authors
suggest that these findings might indicate that
intuitions about crime seriousness track welfare
tradeoffs rather than harm alone (Duntley and
Buss, 2004; Petersen et al., 2012).
The Perceived Social Value of
the Offender
Research on reparative gestures reveals that
they usually involve demonstrations of how
245
the exploiter’s behavior violates social
­obligations (Vangelisti et al., 1991), remind-
ing the exploiter of the favors done for them
in the past (Sell et  al., 2009) or signaling a
wish for future prosocial interactions
(Fujisawa et al., 2005; Petersen et al., 2012).
The reparative gestures convey information to
exploitive persons that they have underesti-
mated the true magnitude of the harm
inflicted, underestimated the true value of the
relationships jeopardized, or overestimated
the gain to the exploiter; it is argued that such
information targets WTR circuits that are
distinct from those targeted by punishment
(Petersen et al., 2012). Petersen et al. (2012)
argue that because different factors regulate
whether specific actions are adaptive in pri-
vate versus public contexts, evolution should
have selected for cognitive machinery
designed to compute a monitored WTR to
govern decisions when one’s actions are
likely to become known to those who will be
affected by them, and a different cognitive
machinery to govern decisions when one’s
actions are not being monitored. It is argued
that reparative gestures aim at up-regulating
the exploiter’s intrinsic WTRs, such that they
inflict less harm in the future even when not
being monitored (Petersen et  al., 2012). In
line with this, research on emotions indicates
that reparative interventions, when success-
ful, have the potential to elicit guilt in
exploiters (Harris et al., 2004), which subse-
quently up-regulates their cooperativeness
(Harris, 2003; Petersen et al., 2012). Several
studies support the idea that feelings of guilt
correspond specifically to an up-regulation
of the exploiter’s intrinsic rather than moni-
tored WTR (Tooby and Cosmides, 2008;
Petersen et al., 2012).
Decision to Punish and/or
to Repair
The dual character of the incarceration system,
i.e. isolation and rehabilitation, is supported
246 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
by the four-phases frame of analysis of the
evolutionary psychology behind the mecha-
nisms of social justice institutions (Ward and
Durrant, 2011): phase 1 refers to the inquiry
into the nature of rehabilitation, which can be
seen as a capacity-building guided process
aiming to assist and shape the capabilities of
the incarcerated offenders in making better
prudential and moral choices; phase 2 con-
sists in the identification of the features of an
effective and ethical rehabilitation process,
such as the offenders’ inherent dignity,
capacity for independent functioning, and
increase in well being, all within the direc-
tion of reducing the risk that these individu-
als will commit further crimes; phase 3 refers
to asking questions about the relevance of
evolutionary approaches to human function-
ing and behavior for the process of incarcera-
tion and of reintegration, in terms of
understanding that the strategies of providing
a ‘good life’ for offenders and other mem-
bers of society should be based on realistic
views of human nature and on the under-
standing of the causes of criminal behavior;
phase 4 refers to the question of whether an
evolutionary psychological explanation pro-
vides the best theoretical guidance on under-
standing the function of incarceration
systems and of communalities of these sys-
tems across the nations of the world.
In their comprehensive analysis of the
decisions to punish or to repair in the con-
text of an evolutionary psychological analy-
sis of modern criminal justice, Petersen
et al. (2012) conclude that, across a range of
different types of crime and across two dif-
ferent countries (i.e. the United States and
Denmark), the preferences of the partici-
pants for rehabilitation over punishment of
the offenders were regulated by the percep-
tion of the social value of the offender, inde-
pendently of their perception of the severity
of the crime. However, the perception of
the seriousness of the crime has regulated
the intensity of preferred sanction (Petersen
et al., 2012). The perceived association value
of the offender was significantly affected by
several experimentally manipulated cues (i.e.
evolutionarily significant cues in the context
of human social functioning), such as the
criminal history of the offender, the offend-
er’s status as in-group or out-group member,
and the offender’s expression of remorse
(Petersen et al., 2012). One of the conclusions
by the authors is that their results support the
hypothesis that the mind’s design for deci-
sions to respond to offenders (exploiters) is
based on two distinct information-processing
channels, i.e. one for computing the serious-
ness of the crime and the other for comput-
ing the criminal’s association value (Petersen
et al., 2012).
While nowadays there is little chance that
an individual’s well being will be directly
affected by the state’s decision to punish or
to rehabilitate a specific offender, a strate-
gic social calculus in terms of this decision
might operate in intimate social settings, sug-
gesting that this might mirror the adaptive
problems faced by our hunter-gatherer ances-
tors (Petersen et  al., 2012). In other words,
the action of the selection pressures that have
favored the mental designs that trigger repar-
ative over punitive strategies in response to
exploitive acts by individuals with potential
social value might be reflected in the modern
sanctioning institutions, including the incar-
ceration system.
THE NICHE CONSTRUCTION THEORY
APPLIED TO THE INCARCERATION
SYSTEM
Based on the four-phases model presented
above, Ward and Durrant (2011) place the
rehabilitation efforts of the prison system
within the niche construction theory (Odling-
Smee et al., 2003), which states that human
beings partially engineer their environments
and in this way contribute to downstream
selection pressures, i.e. in the case of evolu-
tionary functions of criminal justice from the
perspective of niche construction, it is
 EVOLUTIONARY PSYCHOLOGY AND INCARCERATION
assumed that greater community involvement
in offender rehabilitation should encourage a
higher investment by individuals in social
norms and, further, that these individuals will
be less likely to offend (Ward and Durrant,
2011). This idea is in line with the concept of
rehabilitation applied to prison systems,
which refers to normative processes that
manifest at an individual level by assisting
individuals to renounce criminal activity and
construct socially desirable identities, and at
the social level in terms of correctional poli-
cies directed at risk reduction (Ward and
Maruna, 2007; Ward and Durrant, 2011). In
other words, the evolution of modern incar-
ceration systems might be considered an
intentional social-intervention strategy,
which should be analyzed by taking into
account multiple ways of applying evolution-
ary theory to human behavior, such as human
ethology, anthropology, sociobiology,
memetics, and gene-culture co-evolution
theory (Laland and Brown, 2002; Ward and
Durrant, 2011). While one can assume that in
the hunter-gatherer Pleistocene environment,
the reaction of the group to an offender was
probably a quick one in order to quickly
resume everyday life activities, now there is
a considerable social and economic effort in
the direction of recalibration of interactions
with offenders and reconstruction of their
functional social identities.
Niche construction occurs when organisms
alter their environment through metabolism,
activities, and choices, thus reconfiguring
their relationships between their characteris-
tics and the environment (Laland, 2007; Ward
and Durrant, 2011). These alterations might
reduce the selection pressures in the direc-
tion of enhancement of survival and repro-
duction. Examples of niche construction in
humans are not only the products related to
basic survival needs, such as houses, farming
practices, heating systems, medical care, etc.,
but also the products addressing the develop-
ment of social and cultural capital, such as
technology products, books, and educational
systems.
247
Odling-Smee et  al. (2003) describe three
types of processes involved in niche con-
struction: genetic processes, ontogenetic or
developmental processes (individual learning
within lifetime), and cultural processes. These
processes are argued to result in the modifi-
cation of physical and cultural environments
and are implicated in the generation of three
types of inheritance systems (Odling-Smee
et  al., 2003): genetic inheritance, cultural
inheritance, and ecological inheritance (i.e.
the altered ecological niche).
In a comprehensive analysis of niche con-
struction theory in the context of offenders’
rehabilitation, Ward and Durrant (2011) direct
special attention to the following types of
inheritance: (1) culturally stored knowledge,
arguing that this type of knowledge helps the
incoming and current generations of humans
not to repeat the errors made by the previous
generations, and (2) ecological inheritance,
which refers to changes in the environments
and ecologies passed on to a new generation.
According to Ward and Durrant (2011), such
knowledge has the potential to offer greater
flexibility for a species, as well as to gradu-
ally develop increased environmental control.
In this line, the existence of incarceration sys-
tems across human societies, which function
on similar rules and structured interventions
directed to offenders, might indicate that this
system is part of cultural and ecological inher-
itance at the species level. Another aspect that
should be taken into account when placing
the evolution of incarceration systems within
the explanatory frame of niche construc-
tion theory (Ward and Durrant, 2011) is the
existence of two basic types of niche con-
struction: (1) inceptive niche construction,
which refers to the original modification of
the environment, and (2) counteractive niche
construction, which refers to modification
in an attempt to counteract a problem or a
previous change.
Several implications of niche construc-
tion theory, especially of counteractive niche
construction, for the rehabilitation process
of offenders (i.e. specific interventions with
248 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
offenders) are discussed in the literature,
­starting from elements of human behavior to
the concept of extended cognitive system (Ward
and Durrant, 2011). From the point of view
of niche construction theory, human beings
are culturally responsive animals, i.e. human
behavior and socio-emotional capabilities can
be shaped by ethical values and social norms
that are imparted by social learning (Ward
and Durrant, 2011). Hence, a broad range of
behavioral options, as well as attitudes and pro-
clivities of offenders, can be altered by inter-
ventions designed in the direction of socially
responsible and meaningful lives (Clark, 2003;
Sterelny, 2011; Ward and Durrant, 2011).
Also, in the same direction of interpretation, it
is considered that the fact that human beings
can engineer their cognitive environments sup-
ports the existence of reparative interventions
in prisons aiming to diminish the criminogenic
cognitions and crime-supportive beliefs (Ward
and Durrant, 2011).
The concerted effort of society (social sup-
port) and professionals in the field of rehabili-
tation of offenders (e.g., psychologists, social
workers, correctional staff, medical staff, etc.)
points toward an extended cognitive system
in relation to the evolutionary significance of
incarceration. Specifically, Ward and Durrant
(2011) suggest that, if one accepts the idea that
human individuals utilize a combination of
internal and external resources when involved
in cognitive tasks, then it can be assumed that
all the agents involved in the process of reha-
bilitation engineering of offenders are part of
their extended cognitive system. In line with
this, several authors indicate that a proper
understanding of the role of cognition in caus-
ing and maintaining offending is dependent
upon the fact that humans are socially embed-
ded beings with a hybrid cognitive system,
which consists of an integrated combination of
internal and external components that enable
problem solving (e.g., Clark, 2008; Menary,
2007; Ward and Durrant, 2011). In the con-
text of incarceration, it is interpreted that the
survival of offenders and the reflection on
their offending-related problems while being
incarcerated does depend on the cognitive
resources of others (Ward and Durrant, 2011).
Another important aspect that should be
taken into account when addressing the evo-
lution of modern incarceration systems in the
context of niche construction theory is the fact
that humans, regardless of their incarceration-
related status, are naturally inclined to seek
certain goals, which are often called primary
human goods (e.g., physical health, related-
ness, subjective happiness, mastery; Ward
and Maruna, 2007; Ward and Durrant, 2011).
These primary goods or ‘natural desires’
(Arnhart, 1998) are considered markers of
fitness or key components of well being; it is
argued that they have their origin in human
nature and have evolved through natural selec-
tion in the direction of establishing social net-
works favorable to survival and reproduction
(Ward and Durrant, 2011). Primary goods
are assumed to be linked to secure living that
should allow the realization of potentialities
specific to human individuals, such as states
of mind, personal characteristics and experi-
ences, states of affairs, etc. (Ward and Stewart,
2003). Additional to the primary goods, sec-
ondary goods or instrumental goods refer to
the means of achieving primary goods (Ward
and Durrant, 2011). Instrumental means could
be considered both the offending behavior and
the incarceration strategy: in the case of offend-
ing behavior, it is assumed that it can occur
when individuals are trying to achieve primary
goods in often destructive ways at individual
and societal levels (Ward and Durrant, 2011),
while in the case of incarceration systems, we
can assume that restructuring interventions in
prison are means of preventing further crimes
in the direction of a secure living environment.
Effects of Incarceration: Crime-
Suppressive and Criminogenic
Consequences
Definitions of prison are generally based on
their functions related to the consequences of
incarceration on individuals and society.
 EVOLUTIONARY PSYCHOLOGY AND INCARCERATION
Several theories suggest that prison is
­crime-suppressive, while others suggest that
prison can have criminogenic consequences
(Harding et al., 2017). It is assumed that the
crime-reduction effect is achieved through
incapacitation, rehabilitation, and specific
deterrence (Bushway and Paternoster, 2009;
Harding et  al., 2017). The magnitude of any
incapacitation effect depends on the offending
of a comparison group of individuals who
have not been imprisoned, and incapacitation
effects occur only when individuals remain
incarcerated. In contrast, rehabilitation and
specific deterrence will exert their effects after
release. It was also hypothesized that prison
increases criminal offending through stigmati-
zation and labeling effects, through social
learning of pro-criminal attitudes, values,
skills, and roles (prisons as ‘schools of crime’),
and through prison’s effects on employment
prospects (Harding et al., 2017).
Recent studies indicate that returns to
prison are primarily a product of post-prison
community supervision rather than crimino-
genic effects of imprisonment, as many indi-
viduals sentenced to prison are trapped in the
escalating surveillance and punishment of
the criminal justice system; in other words,
the rise in incarceration in the United States
in the late 20th to 21st centuries was in part
a self-perpetuation process resulting from
the workings of the criminal justice system
itself (Harding et al., 2017). Being sentenced
to prison rather than probation increases the
probability of future imprisonment dramati-
cally, regardless of racial groups (Harding
et al., 2017). Harding et al. (2017) suggest that
probation sentences might be employed more
frequently as an alternative to incarcerations.
The cost savings associated with probation
are large relative to the incapacitation effect
of imprisonment and prison sentences do lit-
tle to reduce/prevent criminal offending after
release, relative to offending by probation-
ers. Also, a significant proportion of incar-
cerated individuals are those that have been
recently released from prison and have been
re-imprisoned, i.e. prison’s revolving-door
249
phenomenon (Harding et  al., 2017). Most of
the prison returns are due to a mix of new
crimes and technical violations of the condi-
tions of community supervision in the post-
prison period (Pew Center on the States, 2011,
cited in Harding et al., 2017). Technical vio-
lations during the post-prison period are con-
sidered a key mechanism driving the prison’s
revolving-door effect, rather than an artifact of
the pre-existing differences between prison-
ers and probationers (Harding et  al., 2017).
These results point toward the fact that for the
reparatory function of the prison system to
be effective, the rehabilitation programs and
the monitoring of the former offenders should
continue after their release, especially in the
first years after, i.e. imprisonment for techni-
cal violations among prisoners is concentrated
in the first two years post-release (Harding
et  al., 2017). Even though the data indicate
that there is a moderate incapacitation effect
of incarceration, i.e. a prison sentence reduces
the probability of a new conviction by 5–8% in
the first year after sentence, the increased rate
of post-release imprisonment due to technical
violations supports a self-perpetuating process
resulting from the functioning of the criminal
justice system itself (Siegel, 2014; Harding
et al., 2017).
Social Functioning in Prison:
An Evolutionary Analysis
Various correlates of social functioning in
prison have been addressed in the literature,
such as the personality traits of the inmates,
psycho-affective vulnerabilities, socio-
familial context, behavioral management in
detention, etc. (Picken, 2012; Tomar, 2013;
Unver et al., 2013; Andelin and Rusu, 2015;
Rusu, 2016). The aspects investigated so far
reflect not only the high level of psychosocial
heterogeneity of the prison population, but
also the level of complexity of the process of
planning efficient strategies for the prevention
of self- and hetero-aggressive behaviors in
detention.
250 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
To date, there are no studies concerning
the evolutionary significance of the dimen-
sions associated with the optimal dynamic
of social interactions in prison environments,
specifically their functional value for survival
in this specific environment, in which the
most probable resources to be controlled by
the inmates are those directly related to their
survival, i.e. social interactions that pose the
highest risk to their quality of life. From an
evolutionary perspective, prison environ-
ments represent a complex mixture of stimuli
and selection pressures, bringing together
individuals who are not familiar with each
other and who have different social abilities
and inclusive-fitness-related traits. Inclusive
fitness refers here to the abilities and traits of
an individual organism to survive and pass
on its genes through direct reproduction and/
or by investing somatic effort and other types
of resources in his/her relatives (Hamilton,
1964).
Some of the survival abilities of the indi-
viduals facing incarceration are visible (con-
spicuous) to others, i.e. they can be easily
evaluated by other inmates without neces-
sitating long-term interactions, such as age,
gender, voice, physical appearance, body
mass, general health, access to social support
(family and friend visits). Other survival-
related abilities are less visible (hidden)
at primary evaluation, requiring time and
longitudinal social interactions (e.g., ability
to recognize emotions in specific contexts,
emotional-intelligence level, interpersonal
dominance or submission tendencies, etc.).
Both categories of abilities can be investigated
as predictors for the behavior of individuals in
detention, thus pointing out the need for their
inclusion in the professional screening forms
of newly convicted persons, especially when
dealing with individuals with a known history
of aggression (Rusu, 2016).
Aggression is costly in the prison envi-
ronment, not only at an individual level, but
also at the level of organization and mobili-
zation of the human and other resources of
the prison. Although from an evolutionary
perspective, aggressive behavior is useful for
self-defense and resources protection (Buss
and Shackelford, 1997), it still remains one
of the behaviors posing the highest risk on
the quality of life of incarcerated persons,
both at the physical and psychological levels,
often associated with self-harm and suicide
(Towl, 2003; Campbell, 2005). Violence is a
major problem in settings with incarcerated
persons, and it is frequently associated in the
literature with deficits in the facial emotion
decoding accuracy (Hoaken et  al., 2007).
Emotion-identification errors, especially
anger, are significantly associated with attri-
bution of instrumental value to aggression in
social contexts. Thus, a high level of aggres-
sive attitudes and verbal aggression can be
associated with misperception of anger even
in its absence (Dodge, 1993). Also, individu-
als with a propensity for violence (who are
frequently found in prison settings) have a
higher probability of inadequately interpret-
ing subtle social cues, such as facial micro
expressions of emotions (Hoaken et  al.,
2007).
According to the social information pro-
cessing model (Dodge, 1993; McNiel et al.,
2003), errors in emotion decoding accuracy
have the potential to affect individuals’ abil-
ity (especially of those predisposed to violent
behaviors) to access and employ alternative
adaptive responses to social situations. In the
case of incarcerated persons, there are data
indicating that the ability to recognize facial
expressions of fear and anger is reduced in
inmates with a higher number of arrests and
with a history of aggression (Dodge, 1993).
From an evolutionary perspective, the abil-
ity to detect facial expressions of emotions,
in particular those associated with anger, is
hypothesized to have enhanced the chances
of survival and reproduction of our ancestors
in environments of evolutionary adaptedness,
anger being the main indicator of the inten-
tion to aggress against another individual
(Grandjean et al., 2005; Hoaken et al., 2007).
Another important factor for optimal social
functioning in prison is the level of emotional
 EVOLUTIONARY PSYCHOLOGY AND INCARCERATION
intelligence (EI) of the offenders. EI seems to
be a relevant factor for accessing strategies
of responding to social situations other than
the primary responses such as quick and vio-
lent behaviors. The social-cognitive theory of
power (Fiske, 1993) posits that the ability to
perceive others, an important component of EI,
plays an important role in social-functioning
outcomes. According to this theory, indi-
viduals situated in positions of power tend
to perceive others in a non-individualizing,
stereotypical manner. On the other hand, less
powerful individuals (i.e. submissive individ-
uals) seem to be favored by individualization
of others because they consider interpersonal
relationships as depending on the more pow-
erful individuals and on interaction partners,
in general (Fiske, 1993; Goodwin et  al.,
1998). Having access to emotional signals
and decoding them correctly affords humans
better chances of evaluating the attitudes
and intentions of others (Hess et  al., 1988;
Mayer et  al., 2008), of determining if social
conflict is imminent (Ekman, 2009), and of
adjusting interactive behavior in accordance
with the perceived emotions. Consistent with
the aspects presented above, it is recom-
mended that, besides the standard psychologi-
cal screening forms that are generally used in
prisons, assessments of EI, emotion decod-
ing accuracy, and individual inclusive fitness
should be taken into consideration as bio-
psychological and evolutionary predictors of
optimal social functioning in the prison envi-
ronment (Rusu, 2016).
Costs of High Rates of
Incarceration
Several studies point out that imprisonment
can be a profound life experience not only
for the incarcerated persons, but also for their
families and community (e.g., Mauer, 2017).
These effects have been particularly related
to high incarceration rates. Moderate rates of
incarceration have a higher impact on crime
reduction than high levels of incarceration,
251
which are associated with increases in crime;
this effect is interpreted as the diminishing of
informal social-control mechanisms that
function to establish and reinforce social
norms and create bonds among community
and family members (Clear et al., 2003).
Analyses of small human communities
with high levels of incarceration indicate dif-
ficulties in family formation and child rear-
ing (e.g., Braman, 2002), especially in those
communities where most of the men have
died or are serving in the military, but many
are incarcerated. Hence, in terms of the direct
fitness impact of incarceration at individual
level, the imprisonment of men can consid-
erably diminish the ability of women to find
sexual and parenting partners in these com-
munities (Raphael and Stoll, 2009). Also,
literature indicates that imprisonment of
parents can have unintended negative conse-
quences on the well being of their children,
often related to the placement arrangements
of the children following parental incarcera-
tion (Johnson and Waldfogel, 2002).
In terms of the financial costs imposed by
incarceration systems, comparative analyses
of cost savings between probation and incar-
ceration indicate that the cost savings associ-
ated with probation are large relative to the
incapacitation effect of incarceration (Harding
et al., 2017). In the same study, Harding et al.
(2017) revealed that the impact of prison sen-
tences on reducing criminal offending after
release is lower relative to the offending behav-
ior of probationers; these results support the
policy recommendation of using probation
more frequently as an alternative to incar-
ceration (depending on the seriousness of the
offending) and more efficient planning of the
post-prison parole supervision.
Emotional Burnout among
Correctional Staff
In terms of costs associated with the evolution
of incarceration systems, evidence-based
studies indicate that occupational burnout,
252 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
especially the level of emotional fatigue, is
frequently reported among members of cor-
rectional staff, which can have negative conse-
quences not only at the level of individuals,
but also on the efficiency of the correctional
organization (Hurst and Hurst, 1997; Griffin
et al., 2012; Lambert et al., 2015). Job burnout
is most commonly defined in the literature
as psychological exhaustion and fatigue
due to excessive workplace demands
(Freundenberger, 1975 cited in Lambert et al.,
2015). Maslach and Jackson (1981) have pos-
tulated three dimensions of job burnout: emo-
tional exhaustion, depersonalization, and a
reduced sense of personal accomplishment.
Several studies indicate that the emotional
dimension is the core component of burnout
(Cordes and Daugherty, 1993; Maslach et al.,
2012; Lambert et al., 2015) and that, compared
to other professions from the category of help-
ing others, correctional officers report signifi-
cantly higher levels of emotional exhaustion as
a dimension of job burnout (Maslach et  al.,
2012). The high level of emotional exhaustion
in correctional officers was associated with
turnover intent, absenteeism, and increased
health problems, which were particularly high
in maximum-security settings and in prisons
holding juveniles (Fox, 1982; Carlson and
Thomas, 2006; Lambert et  al., 2010; Griffin
et  al., 2012). Hence, it appears that working
tasks in prison settings involve important
fitness-related costs to the employers, which
are often addressed in terms of prevention and
management by a higher income, possibilities
for early retirement and, in some institutions,
access to psychological- and social-support
programs.
Lambert et  al. (2015) indicate in their
investigation of the consequences of emo-
tional burnout among correctional staff that
the higher the level of education of correc-
tional officers (e.g., college graduates), the
lower the level of their emotional burnout;
the findings are explained by the possibil-
ity that the officers with college degrees
may have been given the opportunity to
participate in decision making within the
institution. Correctional officers are usually
the staff who are responsible for monitoring
and implementation of programs planned by
administrators and supervisory staff (Lambert
et  al., 2015). Previous studies indicate that
a conflict in personal belief system versus
institutional goals contributed to increased
levels of emotional burnout, which in turn
was associated with less favorable attitudes
to treatment (in the context of rehabilitation)
and more favorable attitudes to punishment
(Lambert et  al., 2015). Thus, one can con-
clude that emotional burnout of correctional
staff represents an important variable to be
considered when analyzing the selective
pressures in the context of modern incarcera-
tion’s evolution. Recommendations are made
by several authors regarding the necessity of
exploring other variables of working place
in prison, which are expected to play a role
in correctional staff’s quality of life, support
for treatment, support for punishment, turn-
over intent, and humane consideration of the
needs of the incarcerated people.
Another variable associated with lower
levels of emotional burnout and greater
life satisfaction in correctional staff was
the opportunity to directly interact with the
inmates in a positive manner (Lambert et al.,
2015). When interpreting the behavior and
attitudes of correctional officers toward
inmates, Lambert et al. (2015) suggest that it
is important to take into account that correc-
tional officers tend to have most of the direct
contact with inmates on a daily basis, so their
perceptions of the goal of treatment versus
punishment may be driven by the immediate
circumstances of control rather than the long-
range goals of the institutional plans.
Incarceration System from the
Perspective of Helping Others
In line with the data on the positive associa-
tion between direct contact with inmates and
greater life satisfaction in correctional offic-
ers (Lambert et  al., 2015), one can assume
 EVOLUTIONARY PSYCHOLOGY AND INCARCERATION
that a higher level of awareness of the fact
that working with inmates is one of the pro-
fessions within the category of ‘helping
others’ (i.e. structured forms of prosocial
behavior) may help in preventing emotional
burnout and increase the occupational moti-
vation. A common definition of prosocial
behavior refers to it as ‘a broad category of
acts that are defined by some significant seg-
ment of society and/or one’s social group as
generally beneficial to other people’ (Penner
et al., 2005: 366). A more specific definition
of prosocial behavior from the field of evolu-
tionary psychology considers it as a form of
behavior that brings fitness benefits to the
recipient and diminishes the fitness of the
helper (West et al., 2007).
The costs of helping others are well docu-
mented in the literature, mainly in the con-
texts of caregiving and informal and formal
volunteering, and they usually include psy-
chological and physiological correlates of
caregiver distress (Rusu, 2019). Brown and
Brown (2015) point out that the studies
addressing the stress associated with help-
ing others often fail to distinguish between
the stress associated with the behavior per
se and the feelings about the recipients (e.g.,
compassion, sadness). Besides the costs
mentioned above, recent studies suggest
that helping others in need (i.e. unrelated
individuals) can be associated with ben-
efits, such as experience of positive states
and improvements in health and psycho-
logical well being (e.g., Brown et al., 2009;
Jenkinson et  al., 2013; Rusu, 2019). While
helping others in need from the perspective
of prosocial-behavior definitions implies an
individual autonomous decision and intrin-
sic motivation, when it comes to the profes-
sions targeting the rehabilitation of offenders
in prisons, the idea of helping others and the
ways of doing so are imposed by the institu-
tional and society rules, leaving most prob-
ably little place for the intrinsic motivation
to occur. Therefore, we consider that in order
to increase the rewarding value of the prison-
related professions, more attention should
253
be offered to the prosocial aspects of these
professions. One proposed action would be
to include a learning objective in the train-
ing curricula of the correction officers on the
neurobiological mechanisms and substrate of
successful human social interactions, such as
offering and receiving help from others in sit-
uations of need. For example, several recent
investigations indicate that dopamine is one
of the most commonly suggested candidates
for the explanation of ‘helper’s high’ (Luks,
1988), which is described in the literature as
a sensation of pleasure and subjective happi-
ness associated with helping unrelated others
in need (Krach et al., 2010; Rusu, 2019).
CONCLUSION
The purpose of incarceration, i.e. whether to
rehabilitate or punish offenders, is still a sub-
ject of debate not only within the criminal
justice system itself, but also among members
of society. In this chapter, incarceration is
analyzed from the perspective of recalibration
theory of counter-exploitation (Petersen et al.,
2010, 2012), which is based on the evolution-
ary prediction that, when facing exploitation,
the human mind spontaneously computes the
magnitude of two variables: (1) the exploita-
tion’s seriousness and (2) the exploiter’s asso-
ciation value. It is suggested in the literature
that evolution should have selected for
counter-exploitation strategies designed to
­
recalibrate the exploiter’s welfare tradeoff
ratio (i.e. a variable that sets the weight the
actor places on a specific individual’s welfare
relative to the actor’s own) in the direction of
decreasing the number of exploitive acts in the
future (Sell et al., 2009; Petersen et al., 2012).
In line with this, modern prisons might
function as a monitored strategy of welfare
tradeoff ratio manipulation or adjustment,
combining punishment and reparative actions
in the direction of preventing future harm to
society and up-regulation of the cooperative-
ness of the offenders.
254 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
The incarceration system is also discussed
within the frame of niche construction theory,
specifically the ways in which interactions
with offenders are designed not only with the
purpose of their incapacitation, but also in
the direction of assisting them in renouncing
criminal activity and in constructing socially
desirable identities. Also, the chapter presents
the idea that the concerted effort of society
(social support) and professionals in the field
of rehabilitation of offenders (psychologists,
social workers, correction officers, etc.) could
be interpreted as an extended cognitive system
in relation to the evolutionary significance of
incarceration. In terms of fitness-related analy-
sis, incarceration imposes costs both on the
offenders (e.g., incapacitation due to imprison-
ment, violent interactions in prison, negative
consequences on the well being of their chil-
dren and family) and on the correction staff
(e.g., emotional fatigue, occupational burnout,
turnover intent). Hence, one can conclude that
the two main functions of the incarceration sys-
tem (i.e. punitive and reparatory) are continu-
ously challenged by selective pressures that are
both external ones (i.e. system-independent,
such as financial crises in the society), but also
internal ones, including the system itself.
REFERENCES
Andelin, E.I., & Rusu, A.S. (2015). Investigation
of facial micro-expressions of emotions in
psychopathy – A case study of an individual
in detention. Procedia – Social and Behavioral
Sciences, 209, 46–52.
Arnhart, L. (1998). Darwinian natural right: The
biological ethics of human nature. Albany,
NY: State University of New York Press.
Aureli, F., & de Waal, F. (Eds.). (2000). Natural
conflict resolution. Berkeley and Los Angeles:
University of California Press.
Braman, D. (2002). Families and incarceration.
In Mauer, M. & Chesney-Lind, M. (Eds.),
Invisible punishment: The collateral conse-
quences of mass imprisonment (pp. 137–135)
New York: New Press.
Brown, S.L., & Brown, R.M. (2015). Connecting
prosocial behavior to improved physical
health: Contributions from the neurobiology
of parenting. Neuroscience and Behavioral
Reviews, 55, 1–17.
Brown, S.L., Smith, D.M., Schulz, R., Kabeto,
M.U., Ubel, P.A., Poulin, M., & Langa, K.M.
(2009). Caregiving behavior is associated
with decreased mortality risk. Psychological
Sciences, 20, 488–494.
Bushway, S.D., & Paternoster, R. (2009). The
impact of prison on crime. In Raphael, S. &
Stoll, M.A. (Eds.), Do prisons make us safer?
The benefits and costs of the prison boom
(pp. 119–150). New York: Russell Sage
Foundation.
Buss, D.M., & Shackelford, T.K. (1997). Human
aggression in evolutionary psychological
perspective. Clinical Psychology Review, 17,
605–619.
Campbell, A. (2005). Aggression. In Buss, D.M.
(Ed.), The handbook of evolutionary psychol-
ogy (pp. 628–652) New York: John Wiley.
Carlson, J., & Thomas, G. (2006). Burnout
among prison caseworkers and corrections
officers. Journal of Offender Rehabilitation,
43, 19–34.
Clark, A. (2003). Natural born cyborgs: Minds,
technologies and the future of human
intelligence. New York: Oxford University
Press.
Clark, A. (2008). Supersizing the mind: Embod-
iment, action, and cognitive extension. New
York: Oxford University Press.
Clear, T., Rose, D., Waring, E., & Scully, K.
(2003). Coercive mobility and crime: A
preliminary examination of concentrated
incarceration and social disorganization.
Justice Quarterly, 20(1), 33–64.
Clutton-Brock, T.H., & Parker, G.A. (1995).
Punishment in animal societies. Nature, 373,
209–216.
Cordes, C., & Dougherty, T. (1993). A review
and integration of research on job burnout.
Academy of Management Review, 18,
621–656.
de Waal, F. (1996). Good natured: The origins
of right and wrong in humans and other
animals. Cambridge, MA: Harvard University
Press.
Dodge, K.A. (1993). Social information
processing and peer rejection factors in the
 EVOLUTIONARY PSYCHOLOGY AND INCARCERATION
development of behavior problems in
children. In Biennial Meeting of the Society
for Research in Child Development, New
Orleans, LA.
Duntley, J.D., & Buss, D.M. (2004). The
evolution of evil. In Miller, A. (Ed.), The social
psychology of good and evil (pp. 102–123).
New York: Guilford.
Ekman, P. (2009) Telling lies: Clues to deceit in
the marketplace, politics, and marriage
(Revised Edition). W.W. Norton & Company.
Fiske, S.T. (1993). Controlling other people: The
impact of power on stereotyping. American
Psychologist, 48, 621–628.
Fox, J. (1982). Organizational and racial conflict
in maximum-security prisons. Lexington,
MA: Lexington Books.
Freundenberger, H. (1975). The staff burn-out
syndrome in alternative institutions. Psycho-
therapy: Theory, Research and Practice, 12,
73–82.
Fujisawa, K.K., Kutsukake, N., & Hasegawa, T.
(2005). Reconciliation pattern after
aggression among Japanese preschool
children. Aggressive Behavior, 31, 138–152.
Goodall, J. (1986). Social rejection, exclusion,
and shunning among the Gombe chimpan-
zees. Ethology and Sociobiology, 7, 227–236.
Goodwin, S.A., Operario, D., & Fiske, S.T.
(1998). Situational power and interpersonal
dominance facilitate bias and inequality.
Journal of Social Issues, 54, 677–698.
Grandjean, D., Sander, D., Pourtois, G.,
Schwartz, S., Seghier, M.L., Scherer, K.R., &
Vuilleumier, P. (2005). The voices of wrath:
Brain responses to angry prosody in
meaningless speech. Nature Neuroscience,
8, 145–146.
Griffin, M., Hogan, N., & Lambert, E. (2012).
Doing people work among a tough crowd: A
further examination of the job characteristics
model and correctional staff burnout. Criminal
Justice and Behavior, 39, 1131–1147.
Hamilton, W.D. (1964). The genetical evolution
of social behavior. Journal of Theoretical
Biology, 7, 1–52.
Harding, D.J., Morenoff, J.D., Nguyen, A.P., &
Bushway, S.D. (2017). Short- and long-term
effects of imprisonment on future felony con-
victions and prison admissions. Proceedings
of the National Academy of Sciences, 114(42),
11103–11108.
255
Harris, N. (2003). Reassessing the dimensionality
of the moral emotions. British Journal of
Psychology, 94, 457–473.
Harris, N., Walgrave, L., & Braithwaite, J.
(2004). Emotional dynamics in restorative
justice conferences. Theoretical Criminology,
8, 191–210.
Hess, U., Kappas, A., & Scherer, K.R. (1988).
Multichannel communication of emotion:
Synthetic signal production. In Scherer, K.R.
(Ed.) Facets of emotion: Recent research
(pp. 161–182). Hillsdale, NJ: Lawrence
Erlbaum Associates.
Hoaken, P.N., Allaby, D.B., & Earle, J. (2007).
Executive cognitive functioning and the
recognition of facial expressions of emotion
in incarcerated violent offenders, non-violent
offenders, and controls. Aggressive Behavior,
33, 412–421.
Hurst, T., & Hurst, M. (1997). Gender differences
in mediation of severe occupational stress
among correctional officers. American
Journal of Criminal Justice, 22, 121–137.
Jenkinson, C.E., Dickens, A.P., Jones, K.,
Thompson-Coon, J., Taylor, R.S., Rogers, M.,
& Richards, S.H. (2013). Is volunteering a
public health intervention? A systematic
review and meta-analysis of the health and
survival of volunteers. BMC Public Health,
13, 773.
Johnson, E.I., & Waldfogel, J. (2002). Parental
incarceration: Recent trends and implications
for child welfare. Social Service Review,
76(3), 460–479.
Krach, S., Paulus, F.M., Bodden, M., & Kircher,
T. (2010). The rewarding nature of social
interactions. Frontiers in Behavioral
Neuroscience, 4(22), 1–3.
Kurzban, R., & Leary, M.R. (2001). Evolutionary
origins of stigmatization: The function of
social exclusion. Psychological Bulletin, 127,
187–208.
Laland, K.N. (2007). Niche construction, human
behavioral ecology, and evolutionary
psychology. In Dunbar, R.I.M. & Barrett, L.
(Eds.), The Oxford handbook of evolutionary
psychology (pp. 36–47). Oxford, UK: Oxford
University Press.
Laland, K.N., & Brown, G.R. (2002). Sense and
non-sense: Evolutionary perspectives on
human behavior. Oxford, UK: Oxford
University Press.
256 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Lambert, A., Altheimer, I., & Hogan, N. (2010).
Exploring the relationship between social
support and job burnout among correctional
staff: An exploratory study. Criminal Justice
and Behavior, 37, 1217–1236.
Lambert, E.G., Barton-Bellessa, S., & Hogan,
N.L. (2015). The consequences of emotional
burnout among correctional staff. Sage
Open, 5(2), 1–15.
Luks, A. (1988). Doing good: Helper’s high.
Psychology Today, 22(10), 34–42.
Macovei, M. (2004). The right to liberty and
security of the person: A guide to
implementation of Article 5 of the European
Convention on Human Rights. In Human
rights handbooks (No. 5), Retrieved 18
November 2019 from https://www.refworld.
org/docid/49f181e12.html, Council of Europe.
Maslach, C., & Jackson, S. (1981). The
measurement of experienced burnout.
Journal of Occupational Behavior, 2,
99–113.
Maslach, C., Leiter, M., & Jackson, S. (2012).
Making a significant difference with burnout
interventions: Researcher and practitioner
collaboration. Journal of Organizational
Behavior, 33, 296–300.
Mauer, M. (2017, April 26). Incarceration Rates
in an International Perspective. Oxford
Research Encyclopedia of Criminology.
Retrieved 18 November 2019 from https://
oxfordre.com/criminology/view/10.1093/
acrefore/9780190264079.001.0001/acrefore-
9780190264079-e-233.
Mayer, J.D., Roberts, R.D., & Barsade, S.G.
(2008). Human abilities: Emotional intelligence.
Annual Review of Psychology, 59, 507–536.
McCullogh, M.E., Kurzban, R., & Tabak, B.A.
(2013). Cognitive mechanisms for revenge
and forgiveness (with commentaries and
response). The Behavioral and Brain Sciences,
36, 1–58.
McNiel, D.E., Eisner, J.P., & Binder, R.L. (2003).
The relationship between aggressive
attributional style and violence by psychiatric
patients. Journal of Consulting and Clinical
Psychology, 71, 399–403.
Menary, R. (2007). Cognitive integration: Mind
and cognition unbounded. Basingstoke, UK:
Palgrave Macmillan.
Odling-Smee, F.J., Laland, K.N., & Feldman,
M.W. (2003). Niche construction: The
neglected process in evolution. New Jersey,
NY: Princeton University Press.
Penner, L.A., Dovidio, J.F., Piliavin, J.A., &
Schroeder, D.A. (2005). Prosocial behavior:
Multilevel perspectives. Annual Reviews of
Psychology, 56, 365–392.
Petersen, M.B., Sell, A., Tooby, J., & Cosmides,
L. (2010). Evolutionary psychology and
criminal justice: A recalibrational theory of
punishment and reconciliation. In Hogh-
Olesen, H. (Ed.), Human morality and
sociality: Evolutionary and comparative
perspectives (pp. 72–131). Hampshire:
Palgrave Macmillan.
Petersen, M.B., Sell, A., Tooby, J., & Cosmides,
L. (2012). To punish or repair? Evolutionary
psychology of lay intuitions about modern
criminal justice. Evolution of Human
Behavior, 33(6), 682–695.
Pew Center on the States (2011). State of
Recidivism: The Revolving Door of America’s
Prisons. The Pew Charitable Trusts,
Washington, DC.
Picken, J. (2012). The coping strategies,
adjustment and well being of male inmates in
the prison environment. Internet Journal of
Criminology, 1–29. Retrieved 8th of December
2018 from https://studylib.net/doc/8657392/
the-coping-strategies–adjustment-and-well-
being-of-male.
Raphael, S., & Stoll, M. (Eds.). (2009). Do
prisons make us safer? The benefits and
costs of the prison boom. New York: Russell
Sage Foundation.
Rossi, P.H., Simpson, J.E., & Miller, J.L. (1985).
Beyond crime seriousness: Fitting the
punishment to the crime. Journal of
Quantitative Criminology, 1, 59–90.
Rusu, A.S. (2016). Evolutionary-based aspects
of optimal social functioning in prison. Acta
Psychopathologica, 2(6), 1–3.
Rusu, A.S. (2019). Educational practices for
civic engaged students: Service-Learning -
from general to applied values in animal-
oriented professions. Journal of Educational
Sciences & Psychology, 9, 29–35.
Sell, A., Tooby, J., & Cosmides, L. (2009).
Formidability and the logic of human anger.
Proceedings of the National Academy of
Sciences, 106, 15073–15078.
Siegel, J.A. (2014). Prisoner reentry, parole
violations, and the persistence of the
 EVOLUTIONARY PSYCHOLOGY AND INCARCERATION
surveillance state. PhD Dissertation (University
of Michigan, Ann Arbor, MI).
Sterelny, K. (2011). The evolved apprentice.
Cambridge, MA: MIT Press.
Stylianou, S. (2003). Measuring crime
seriousness perceptions: What have we
learned and what else do we want to know?
Journal of Criminal Justice, 31, 37–56.
Tomar, S. (2013). The psychological effects of
incarceration on inmates: Can we promote
positive emotion in inmates? Delphi
Psychiatry Journal, 16, 60–68.
Tooby, J., & Cosmides, L. (2008). The
evolutionary psychology of the emotions and
their relationship to internal regulatory
variables. In Lewis, M. & Haviland-Jones, J.M.
(Eds.), Handbook of emotions (pp. 114–137;
3rd ed). New York: Guilford Press.
Tooby, J., Cosmides, L., & Price, M.E. (2006).
Cognitive adaptations for n-person
exchange: The evolutionary roots of
organizational behavior. Managerial and
Decision Economics, 27, 103–129.
Towl, G. (2003). Psychology in prisons. Oxford,
UK: BPS Blackwell.
Unver, Y., Yuce, M., Bayram, N., & Bilgel, N.
(2013). Prevalence of depression, anxiety,
stress, and anger in Turkish prisoners. Journal
of Forensic Sciences, 58, 1210–1218.
257
Vangelisti, A.L., Daly, J.A., & Rudnick, J.R.
(1991). Making people feel guilty in
conversations: Techniques and correlates.
Human Communication Research, 18,
3–39.
Ward, T., & Durrant, R. (2011). Evolutionary
psychology and the rehabilitation of the
offenders: Constraints and consequences.
Aggression and Violent Behavior, 16,
444–452.
Ward, T., & Maruna, S. (2007). Rehabilitation:
Beyond the risk paradigm. London, UK:
Routledge.
Ward, T., & Stewart, C.A. (2003). The treatment
of sex offenders: Risk management and
good lives. Professional Psychology: Research
and Practice, 34(4), 353–360.
West, S.A., Griffin, A.S., & Gardner, A. (2007).
Social semantics: Altruism, cooperation,
mutualism, strong reciprocity and group
selection. Journal of Evolutionary Biology,
20, 415–432.
Wilson, E.O. (1980). Sociobiology: The abridged
edition. Cambridge, MA: Belknap Press.
Wrangham, R.W. (1987). The significance of
African apes for reconstructing human social
evolution. In Kinzey, W.G. (Ed.), The evolution
of human behavior: Primate models
(pp. 51–71). Albany, NY: SUNY Press.

Evolution and Punishment
Anthony Walsh, Cody Jorgensen, and Jessica Wells
INTRODUCTION
In the opening words of The Scarlet Letter,
first published in 1850, Nathaniel Hawthorne
wrote: ‘The founders of a new colony, what-
ever Utopia of human virtue and happiness
they might originally project, have invariably
recognized it among their earliest practical
necessities to allot a portion of the virgin soil
as a cemetery, and another portion as the site
of a prison’ (1879: 1). Hawthorne’s words are
a reminder of two things we cannot avoid –
human mortality and depravity, and that we
must make provisions for both. We sadly
commit our departed loved ones to the
ground for obvious reasons, and we gladly
remove miscreants from our sight for reasons
just as obvious. It is beyond doubt that there
are times when we must set certain people
apart from the rest of us because they have
demonstrated their inability to live by the
agreed-upon norms of acceptable behavior.
When we do this we are doing it against the
will of the person being removed, and thus
13
we are expressing censure and community
disapproval of his or her actions by inflicting
punishment upon him or her.
Of course, prisons are a relatively modern
invention, but we have always retaliated in
some way against those who have offended
us. We see this retaliatory aggression in
almost all sexually reproducing animals
when conspecifics ‘cheat’ (signal coopera-
tion but fail to be forthcoming). A classic
example is that of vampire bats who freeload
on blood donation. Bats who fail to find a
blood meal on a given night will have regur-
gitated blood donated to them by other mem-
bers of the group. The tendency is to share
blood with those who have previously shared
with them, thus returning the favor. Failure
to reciprocate usually results in the original
benefactor withdrawing future acts of sharing
(Wilkinson, 1990). So even bats have some
built-in motivation to retaliate punitively
against norm breakers.
From a scientific perspective, any emotion
or behavior that is shown to be as universal as
 EVOLUTION AND PUNISHMENT
the urge to punish must have an evolutionary
history. Natural selection is a powerful posi-
tive feedback process energized by differen-
tial reproduction. If a genetic mutation results
in the design (morphological, physiological,
or behavioral) of an organism that allows it
to out-reproduce other conspecifics, it will
eventually become more common in the
population and we then say that the design
change has been selected for. Over many
generations, the design change will spread
until it goes to fixation in the population; that
is, all members of the species have it.
Cosmides and Tooby (1994: 328) liken
natural selection to Adam Smith’s ‘invisible
hand’ whereby unobservable market forces
(‘natural selection’) fueled by millions of
people seeking their self-interest (‘survival
and reproduction’) move the supply and
demand of goods (‘genetic polymorphisms’)
in a free market (‘the environments of evo-
lutionary adaptation’) to reach market equi-
librium (‘genetic fixation’) automatically and
without any intention or foresight.
The consequences of punishing norm vio-
lators must have been positive with respect to
the twin goals of all life – survival and repro-
duction. It is important for all animals to be
motivated to do things that are vital in the
pursuit of these goals. To provide that moti-
vation, nature has provided us with neural
mechanisms that reward us with pleasurable
feelings when we do things that aid in achiev-
ing these goals. These pleasurable feelings
arise from the neurotransmitter dopamine
bombarding the nucleus accumbens and sim-
ilar so-called ‘pleasure centers’ in the brain
(Walsh et al., 2012). Each surge of dopamine
can be viewed as Mother Nature’s reward
when we satisfy urges to eat, drink, and have
sex, which are the most obvious requirements
for survival and reproduction. The pleasant-
ness associated with dopamine (known as the
‘happy chemical’ in popular parlance) rein-
forces the behavior that makes us feel good
and motivates us to repeat it.
Given that the urge to punish wrongdo-
ers is universal and strong (Penney, 2012),
259
punishment obviously played an adaptive
role, but the survival and reproductive func-
tion of punishment is not at all as obvious as
are the roles of food, drink, and sex. We do
know, however, that humans find the same
kind of satisfaction when heinous criminals
are punished. Brain-imaging studies have
shown increased blood flow to the nucleus
accumbens when subjects witness the pun-
ishment of those who have wronged them (de
Quervain et  al., 2004; Klein, 2012). Blood
flow to reward centers has also been observed
when people are punished who have harmed
others, and the strength of the pleasure
response is proportional both to the harm
done and to the level of the offender’s culpa-
bility (whether the act was purposeful, know-
ing, reckless, or negligent) (Buckholz and
Marois, 2012). These brain-imaging studies
provide us with compelling evidence that the
urge to punish must be an adaptive feature of
human nature, attested to by the old saying,
‘Vengeance is sweet’.
Petersen and colleagues’ evolutionary per-
spective on criminal justice notes the various
ways that our ancestors could have reacted
to exploitative harm or other norm violations
(2010: 92). These four ways they have identi-
fied are presented below.
1 ‘Killing the perpetrator, which permanently
removes the threat’. In an evolutionary context,
executing a band member would probably be
considered a hugely important decision since all
band members would have known the perpetra-
tor and his or her family. The decision would have
required a sufficiently serious group-damaging
act and significantly more members of the band
favoring execution than opposing it. In the
modern United States, this is the death penalty,
the imposition of which requires a particularly
heinous murder and/or multiple murders. The
death penalty is retributive in nature, although
some justify it by a supposed deterrent effect.
People who believe in the value of retribution
say that punishment for its own sake is morally
justified and need not be judged good or bad
according to its consequences for society (such as
deterrence). Logan and Gaes (1993) aver that ret-
ribution is the most honestly stated justification
260 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
for punishment because it taps into our punitive
urges and posits no secondary purpose for it.
2 ‘Expelling the perpetrator from the social world
through ostracism or confinement; or, on an indi-
vidual basis, not engaging in future cooperation
endeavors with those who have cheated’. In
a band situation, being ostracized would have
been devastating because the ostracized person
would have no other group to which to turn for
sustenance. Not so long ago, those who violated
cultural and social norms were publicly shamed
by being placed in the stocks for all passers-by
to taunt and abuse. We no longer publicly abuse
villains or tell them to ‘Get out of Dodge’; rather,
we use imprisonment and justify it by the concept
of incapacitation. The justification for incapacita-
tion is well put by James Q. Wilson (1975, p. 391):
‘Wicked people exist. Nothing avails except to set
them apart from innocent people’. Incapacitation
obviously has positive consequences for society;
while criminals remain in prison they can no longer
harm people on the outside. For instance, in 1995,
there were 135,000 inmates in prison whose most
serious crime was robbery, and each robber on
average commits five robberies per year. Had these
robbers not been incarcerated, they would have
been responsible for an additional 675,000 rob-
beries that year (Currie, 1999). An Italian study of
22,000 inmates published in 2006 found that the
incapacitation effect was an average of between
14 and 18 subsequent theft and robbery (these
were the only crimes documented) arrests per
released convict (Buonanno and Raphael, 2013).
3 ‘Punishing the perpetrator through infliction of
costs or withdrawal of benefits’. This response to
norm violations would have been far more damag-
ing in ancestral environments if it meant the with-
drawal of cooperation; in a modern sense it would
still be painful but less so. Today such punishments
would be in the form of fines and/or withdrawal of
welfare benefits; just like the ‘No more blood for
you, Vlad’ response to cheater bats.
4 ‘Reconciling with the perpetrator’. In a modern
criminal justice system we call this rehabilitation,
or to restore or return to constructive, healthy
relationships with other members of society.
Rehabilitation is criticized by conservatives as
‘molly cuddling’ and lacking in truth in sentencing,
and by libertarians and some liberals for forc-
ing offenders into treatment against their wills.
Liberals charge that it contributes to sentenc-
ing disparity because it requires indeterminate
sentencing to accommodate different rates of
offender ‘rehabilitation’. Yet we continue to try
to rehabilitate criminals because we realize that
whatever helps the offender helps the commu-
nity. The rehabilitation model is well stated in
former United States Supreme Court Chief Justice
Warren Burger’s famous lines: ‘To put people
behind walls and bars and do little or nothing
to change them is to win a battle but lose a war.
It is wrong. It is expensive. It is stupid’ (in Stohr
et al., 2012: 246).
Of course, all these responses to norm viola-
tions are punitive in that those experiencing
them do not welcome them. By imposing
such pain, we hope that offenders will not
repeat the behavior that led to his or her pun-
ishment, and to make those witnessing the
punishment not be tempted to follow the
same path. We call this punishment justifica-
tion ‘deterrence’. Radzinowicz and King
(1979) aver that retribution and deterrence
occur simultaneously, but that the real social
function of punishment is deterrence:
People are not sent to prison primarily for their
own good, or even in the hope that they will be
cured of crime. Confinement is used as a measure
of retribution, a symbol of condemnation, a vindi-
cation of the law. It is used as a warning and
deterrent to others. It is used, above all, to protect
other people … from the offender’s depredations.
(Radzinowicz and King, 1979: 296)
RETRIBUTION: ITS ORIGIN AND
FUNCTION
Whatever justification for punishment that
philosophers and criminologists may come
up with, underneath it all is the urge for ret-
ribution. Retribution means recompense,
repayment, payback, or getting even. Many
criminologists tend to view retribution as an
irrational and barbaric justification for pun-
ishment based on emotion rather than reason.
They may note that retribution is most often
given as a rationale by supporters of the
death penalty (Bohm, 2012), which many see
 EVOLUTION AND PUNISHMENT
as little more than state-sanctioned revenge
(Rosebury, 2009). This position carries with
it the unspoken assumption that the emotions
behind the retributive urge are immoral and
antisocial. Evolutionary psychologists and
neuroscientists beg to differ, as did Justice
Potter Stewart. In his concurring opinion in
the Supreme Court’s 5–4 ruling in Furman
vs. Georgia (1972) invalidating Georgia’s
death penalty statute, he wrote the following
about the importance of retribution and the
folly of dismissing it:
I cannot agree that retribution is a constitutionally
impermissible ingredient in the imposition of pun-
ishment. The instinct for retribution is part of the
nature of man, and channeling that instinct in the
administration of criminal justice serves an impor-
tant purpose in promoting the stability of a society
governed by law. When people begin to believe
that organized society is unwilling or unable to
impose upon criminal offenders the punishment
they ‘deserve,’ then there are sown the seeds of
anarchy – of self-help, vigilante justice, and lynch
law. (Furman vs. Georgia 408 U.S. 238, 1972)
What did Justice Stewart mean by locating
the ‘instinct for retribution’ in the ‘nature of
man’? One method of determining if some-
thing is ‘in the nature of man’ is to scour the
historical record for the universality of the
practice so considered. All early legal codes,
such as the ancient Mesopotamian codes of
Hammurabi, Ur-Nammu, and Eshnunna, and
the Roman Laws of the Twelve Tables, were
retributive, and the universality of such senti-
ments in criminal codes supports Stewart’s
contention about the urge for retribution
being a strong component of human nature.
Given the assumption that it is part of human
nature, how did it become so and why?
Evolutionary logic tells us there must have
been good adaptive reasons why our reward
centers fire up when we punish or witness
the punishment (vicariously or otherwise) of
wrongdoers, but it does not tell us why pun-
ishment was so vital to our distant ancestors.
We may approach the question by noting that
Homo sapiens are an ultra-social species con-
sisting of individuals who are preeminently
261
motivated to attend to their own survival and
reproductive interests. The urge to cooperate
with our fellow creatures and the urge to sig-
nal cooperation and then defect, thus receiv-
ing resources illegitimately, is the hedonic
tug of war we all play with ourselves. Most
of us successfully adjust with nary a thought
of violating the social contract; some adjust
with difficulty, and others find the illegiti-
mate route congenial to their natures.
The modern evolutionary view of human
nature accommodates the classical view
that humans are designed to maximize their
pleasure and minimize their pain. The gov-
ernance of these two sovereign masters is an
adaption and thus functional, but it can be
socially disruptive if immediate concerns for
gratification are placed above maintaining
social harmony and one seeks ‘money with-
out work, sex without courtship, and revenge
without court delays’, as Gottfredson and
Hirschi (1990: 89) put it. This is the origin
of the urge to punish because any individual
who appropriates resources illegitimately
thereby deprives the legitimate owners of
resources needed for their own survival and
reproductive success.
Counteracting the human competitive and
status-striving motives that lead to conflict
is a powerful egalitarian instinct that leads
to cooperation. One of the first moral state-
ments uttered by children, and often quite
forcefully, is ‘That’s not fair!’ How do chil-
dren arrive at that conclusion long before they
are able to articulate any moral reasoning for
it, and why is it so compelling? A number of
scholars (see generally Gavrilets, 2012) have
posited that the nomadic lifestyle character-
ized by dangerous and uncertain prospects
of obtaining survival resources kick-started
the evolution of our species’ powerful social
and egalitarian instincts. Small foraging
and hunting and gathering bands demanded
strict group-wide cooperation, with each
individual’s survival depending on the sur-
vival of the whole. Any kind of behavior that
would subvert mutual reciprocity (not pull-
ing one’s weight, cheating, stealing) could
262 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
not be tolerated. Scarce resources had to be
distributed according to egalitarian principles
lest the group fall into disputes and fracture,
leading perhaps to the death of everyone in
it. In such a group, resource sharing would
take place under the vigilant eyes of all in
immediate time as opposed to a delayed-
time sharing of society’s resources in agrar-
ian and industrial societies. Perceptions of
unequal distribution of resources would have
produced immediate emotional reactions that
were not of the warm and pleasant variety.
Social animals cooperate because they can
achieve more as a group than they can indi-
vidually, and we feel good about ourselves
when helping others. We are also rewarded
by reciprocal behavior in the future, but even
if the person we help is a stranger we are
unlikely to meet again, we still receive a shot
of dopamine that makes us feel good (Brunero,
2002). However, a population of cooperators
would thrive and grow bigger, thereby plant-
ing the seeds of norm violation. A fairly large
group of cooperators consisting overwhelm-
ingly of non-kin provides a target-rich niche
for people who cheat on the social contract
and seek to gain resources at zero cost due to
a lower level of vigilance and a higher level
of anonymity. It has been estimated that the
maximum group size to maintain stable rela-
tions among all members is around 150; this
is known as ‘Dunbar’s Number’ (Hill and
Dunbar, 2003). This number is derived from
observation of different human groups which
maintained stable relations throughout their
history, extrapolation from relative neocortex
size to group size in different species, and the
assumed cognitive constraints involved in
maintaining cohesion in face-to-face interac-
tion while still pursuing one’s own personal
evolutionarily relevant interests.
The reference to neocortex size calls on
studies of a wide range of primate species that
show that group size is related to brain size
(Dunbar and Shultz, 2007). Most evolutionary
adaptations, including brain adaptations, are
the result of challenges to survival and repro-
ductive success posed by ecology (finding
food and mates, fighting off pathogens and
predators, etc.) that all animals share, but
human culture has shaped both the anatomy
and function of the human brain above and
beyond the changes wrought by the chal-
lenges posed by ecology. We know that from
the approximate 1.5 million years that sepa-
rated Australopithecus afarensis and Homo
erectus, hominid cranial capacity doubled
from a mean of 450 cc to a mean of 900 cc,
and by another 70% to about 1,350 cc from
Homo erectus to modern Homo sapiens.
On an evolutionary timescale, this is truly
an astounding level of morphological change
in such a short period of time (Adolphs,
2009).
A study of hominid crania by Bailey and
Geary (2009: 77) found that geographic lati-
tude was strongly related to cranial capacity
(r = .61), but population density was more
strongly related (r = .79), leading them to
conclude that the burden of evolutionary
selection has moved from ‘climactic and
ecological to social’. There are a number
of other studies of hominid crania dating as
far back as 1.9 million years that show more
robust increases in cranial capacity among
groups in areas with greater population den-
sity and in areas in which food procurement
was most problematic, namely, colder and
most northerly areas of the globe (Ash and
Gallup, 2007; Kanazawa, 2008). Living in
large groups forces the brain to make more
computations in order to negotiate relation-
ships, to understand the thoughts, feelings,
and intentions of others, and to cooperate
in securing resources and in defending the
group (Dunbar and Shultz, 2007).
Despite increases in reasoning power, as
has been pointed out, it is difficult to maintain
the fuzzy warmth of all-for-one-and-one-for-
all kind of ‘primitive communism’ in groups
larger than about 150 (Dunbar’s number).
The seminal studies of Hutterite communi-
ties by Baden and Stroup (1972) are among
the studies that reveal this. The Hutterites
are a religious community spread across
colonies in the United States and Canada.
 EVOLUTION AND PUNISHMENT
The Hutterites seem to have a c­ ommon-sense
understanding of human nature, and man-
date that a settlement must split and form
a new one every time the mother colony
reaches about 150 people. When the colony
is small, public surveillance of each person
is more complete and an accounting of each
person’s contribution (or lack of) is possible,
just as it was on the ancient African savan-
nah. The Hutterites recognize freeloaders
who take advantage of their system of com-
munal ownership of resources and call them
drones (reference to male drone bees who do
not participate in nectar and pollen gathering
and whose only role is to mate with a queen
bee). They also recognize that the number of
‘drones’ increases proportionately to colony
size, which affords the drones greater auton-
omy. When the colony splits the ‘drones’
again come under the vigilant gaze of others
and revert to norm compliance and become
good ‘worker bees’ in the offspring colony.
KANT’S MORAL JUSTIFICATION
FOR RETRIBUTION
The philosopher John Mackie (1982: 3) saw
a paradox inherent in retribution because it
‘cannot be explained or developed within a
reasonable system of moral thought, while,
on the other hand, such a principle cannot be
eliminated from our moral thinking’. This
‘paradox’ is the result of assuming that
rational deliberations about moral issues can
take place without involving the emotions
that, as we will see, cannot be done. However,
Immanuel Kant has defended retribution
rationally and morally. Kant’s justification of
punishment is retributive but he is aware that
retribution carries a message to the public of
moral disapproval of criminals and their acts,
and therefore conveys a de facto general
deterrent effect. Deterrence rides on retribu-
tion’s coat tails, but as a stand-alone justifica-
tion it cannot be a moral justification for
punishment in Kantian terms.
263
Kant justifies retribution as the only moral
justification for punishment in terms of the
value, dignity, and agency of human beings.
Kant’s retributionism is undergirded by his
idea of duty and beliefs in human reason and
autonomy. Human beings are different from
other animals because they are guided by
reason, and it is from reason that we derive
all of our duties and obligations. Because we
are blessed with reason, we all have a duty
to act out of reverence for moral law, which
Kant conceived of as a set of categorical
imperatives. A categorical imperative is a
moral duty to be discharged regardless of any
further end or consequence. A categorical
imperative commands ‘Do this!’; it does not
say ‘Do this if’. Any action consistent with
this is morally good, says Kant, regardless
of its consequences. Categorical impera-
tives are viewed as universal laws that guide
humans toward their duties: ‘Act as if the
maxim of your action were to become through
your will a universal law of nature’ (Kant,
1964: 89).
In his Groundwork of the Metaphysics of
Morals, Kant says that a categorical impera-
tive should be grounded in something that
should be an ‘end in itself, and an absolute
value’. He finds this grounding in ‘man’,
who ‘exists as an end in himself, not merely
as a means for arbitrary use for this or that
will’ (1964: 95). ‘Man’, in all his actions,
must always be regarded as an end in himself
regardless of whether his actions are directed
at himself or at others. Based on this, Kant
arrives at a final definition of a categorical
imperative:
Act in such a way that you always treat humanity,
whether in your own person or in the person of
any other, never simply as a means, but always at
the same time as an end. (1964: 96)
Respecting humans as ends in themselves
leads to a retributionist justification of pun-
ishment. From this viewpoint, punishing
criminals for instrumental reasons such as
deterring others or subjecting them to reha-
bilitative treatment is morally wrong because
264 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
it treats them as means to an end, not as ends
in themselves. As Kant phrased it in The
Metaphysical Elements of Justice:
Judicial punishment can never be used merely as a
means to promote some other good for the criminal
himself or for civil society, but instead it must in all
cases be imposed on him only on the ground that he
has committed a crime; for a human being can
never be manipulated merely as a means to the
purposes of someone else … He must first of all be
found to be deserving of punishment before any
consideration is given of the utility of his punishment
for himself or his fellow citizens. (Kant, 1965: 331)
According to Kant, criminals are ruled by
reason just as everyone else and thus live in
accordance with maxims – universal moral
standards (‘Act as if the maxim of your action
were to become through your will a universal
law of nature’). If criminals harm others, they
are violating the autonomy of others, and by
doing so, they are said to be endorsing criminal
acts as universal laws, and thus in effect they
are saying that others should act similarly. By
punishing criminals, the state is treating them
in accordance with their own maxims; that is,
how they think others should be treated. The
state is thus allowing criminals to decide how
they will be treated, and by doing so, it is
respecting their judgment and autonomy. Thus,
Kant says of the criminal: ‘His own evil deed
draws the punishment upon himself’ (in
Rachels, 1986: 123). Retribution is a ‘just
deserts’ model demanding that criminals be
punished in proportion to the harm they have
inflicted on their victims; thus death would be
a proportional punishment for someone who
has taken the life of another.
THE MORAL LAW WITHIN
Humans live under an implied social contract
by which we surrender some of our freedoms
to do as we please in exchange for security.
A vital part of the contract is the agreement
not to harm others, and if we do, we recog-
nize the fact that the state has the legitimate
right to punish us. As McBride (2007: 122)
avers: ‘Punishment is the midwife in the birth
of the social contract’. Central to the social
contract is social integration and social order
based on the strongly felt norms of right and
wrong among its members. In evolutionary
times, deeply felt moral rules helped individu-
als to stay connected to the group, survive in
dangerous environments, and garner a valua-
ble reputation as being trustworthy. Morals
were therefore invaluable in terms of achiev-
ing shared ends.
No society can do without intolerance, indigna-
tion, and disgust; they are the forces behind the
moral law. (Moore, 1987: 199)
Moore adds: ‘In this view, the emotions of a
people constitute moral truth’ (1987: 199).
Individuals who freeload or cheat (we call
the worst of them criminals) will always
prosper in a population of unconditional
cooperators whom evolutionary psycholo-
gists call ‘suckers’. Cheats would soon drive
suckers to extinction. It has been consistently
shown in computer simulation games such as
the prisoner’s dilemma that in a mixed popu-
lation of cooperators and cheats, cheats
always do better than cooperators when no
punitive response is forthcoming (Klein,
2012; Nowak, 2006). Of course, very few
humans are suckers. The vast majority of us
are ‘grudgers’ who cooperate conditionally.
Grudgers can be cheated because they abide
by the norms of mutual trust and cooperation
and expect the same from others. Once
cheated, however, grudgers will react differ-
ently to cheaters in the future in tit-for-tat
ways (Wiebe, 2011).
Anyone stealing resources or sexual mates
in our species’ evolutionary history consti-
tuted a severe threat to everyone in a group
relying on strong norms of reciprocity, and
thus would have generated intense negative
emotions such as anger, outrage, disgust, and
a desire to punish. As de Waal (1996: 160)
puts it: ‘A taste of revenge is the other side
of the coin of reciprocity’. Victims feel angry
and hurt when treated unfairly. Victims also
 EVOLUTION AND PUNISHMENT
feel confusion and frustration at losing the
expectation of the predictability of reciproc-
ity. The sum of these evolved emotions is
moral outrage. Without moral outrage there
would be no internal motivation to react
against those who violate the norms of recip-
rocal cooperation, cheats would have thrived
without the threat of punishment in our
ancestral environments, and we would have
evolved as a quite different species of animal
(Haidt, 2012; Nowak, 2006).
However, natural selection does not pass
its judgment on emotions, or even on the
behaviors they motivate, if those behaviors
do not result in enhanced reproductive suc-
cess. Natural selection operates on the con-
sequences of the behavior motivated by the
emotion (Massey, 2002; Walsh, 2006). It is
no use silently feeling angry and hurt when
victimized. Those feelings must generate
behavior designed to prevent it occurring
again. Negative feelings caused by victimi-
zation are mediated by punishing violators
because punishment signals the restoration
of fairness and predictability (the perception
that cheaters may be less likely to cheat in
the future, and that potential cheaters may
be deterred). The positive feelings accom-
panying the punishment of those who have
wronged us, coupled with the reduction of
negative feelings, provide powerful rein-
forcement for punitive responses and may
explain why we see increased blood flow to
the brain’s reward centers when wrongdoers
are punished.
THE IMPORTANCE OF MORAL
EMOTIONS IN UNDERSTANDING
PUNITIVE BEHAVIOR
We have been waxing about such things as
moral outrage, disgust, and revenge, but
aren’t these things all primitive emotions,
and surely we have progressed beyond rely-
ing on such unreasoned criteria when decid-
ing what to do with criminals? Primarily, we
265
are concerned here with the moral or ‘social’
emotions such as guilt, shame, empathy, and
embarrassment, although the more basic
emotions such as anger, disgust, and joy
come into play in punishment. These social
emotions that evolved as integral parts of our
social intelligence provide clues about the
kinds of relationships (cooperative vs unco-
operative) that we are likely to have with
others, and serve as ‘commitment devices’
and ‘guarantors of threats and promises’
(Mealey, 1995: 525). Barkow (1989: 121)
describes them as involuntary and invasive
‘limbic system overrides’ that serve to adjust
our behavior in social situations. The social
emotions animate, focus, and modify neural
activity in ways that lead us to choose certain
responses over other possible responses from
the streams of information we constantly
receive. The social emotions move us to
behave in ways that enhanced our distant
ancestors’ reproductive success by overriding
neocortical decisions suggesting alternatives
to cooperation (i.e., cheating) which may
have been more rational in the short term, but
which were ultimately fitness reducing.
One of the most salient emotions in peo-
ple’s punishment urges is empathy; not for
the person to be punished but rather for his
or her victim. Rob Canton (2015: 59) main-
tains, ‘The emotions of punishment are dis-
tinctly moral emotions. They are emotions
of judgement, of righteousness and repro-
bation’. Canton is implying that people feel
these emotions not just for themselves, but
also on behalf of victims they have never
met, and they are therefore deeply prosocial
emotions linked to concern for others. This
is the emotion we call empathy. Empathy is
an ancient capacity predating the emergence
of Homo sapiens that evolved rapidly in the
context of parental care and is found only in
mammals and some bird species (de Waal,
2008). Empathy is the cognitive and emo-
tional ability to understand the feelings and
distress of others as if they were our own.
The cognitive component allows us to under-
stand their distress and why they are feeling
266 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
it, and the emotional component allows us
to feel that distress ourselves (‘I feel your
pain’). Feelings of distress are discomforting
because they trigger stress/anxiety hormones
(i.e., cortisol), giving us the unconscious
feeling that we are threatened and motivating
us to ‘do something’.
To the extent that we feel empathy for oth-
ers, we have a visceral motivation to take some
action to alleviate their distress if we are able;
thus empathy is the emotional component of
altruistic actions. Performing a helpful action
on behalf of another alleviates the distress of
the other individual and thereby we alleviate
our own. Thus, empathy has a selfish compo-
nent, but it is a very good thing that it does
because if we were lacking in emotional con-
nectedness to others, we would be callously
indifferent to their needs and suffering. Once
locked into the human repertoire, empathetic
feelings can be diffused to a wider network
of social relationships, which is why empa-
thy is considered ‘the heart of mammalian
development, limbic regulation, and social
organization’ (Farrow and Woodruff, 2007:
51). Empathy is arguably the quintessential
social emotion, and therefore it is difficult to
dismiss its impact on punishment decisions
as irrational.
We recognize that doing the right thing
when there is a strong temptation to do other-
wise as conscience. In The Descent of Man,
Charles Darwin opined that conscience is
based on two things: the social instincts and
cognition and habit: ‘the social instincts – the
prime principle of man’s moral constitution –
with the aid of active intellectual powers
and the effects of habit, naturally lead to the
golden rule’ (Darwin, 1896: 106). Obviously,
we must have a cognitive awareness of what
is right and wrong, but the consensus of opin-
ion agrees with Darwin that the emotional
component of conscience is most important
(Choy et  al., 2015; Kochanska and Aksan,
2004). The degree to which the sympathetic
branch of the autonomic nervous system
(ANS) is activated, and/or the speed at which
the parasympathetic branch returns the ANS
to homeostasis in the face of temptation and
the decision not to give in, is the major deter-
minant of conscience development. It is obvi-
ous that ‘doing the right thing’ is much more
than rational consideration since that thing
we call a conscience typically moves us in
that direction before we have had an oppor-
tunity to contemplate our options (Barkow’s
‘limbic system override’).
RATIONALITY AND EMOTION: THE
HEADS AND TAILS OF DECISION-
MAKING
The ancient notion that rationality and emo-
tion are polar opposites is no longer viable. It
has long been known in neuroscience that
cognition is always suffused with emotion
and cognition with emotion (Nowak and
Sigmund, 2005). Our emotional and rational
neural mechanisms work in unison and sus-
tain each other, and often cannot be separated.
As we have seen, decision based on emotions
are typically more effective when the decision
to be made is a moral one than a decision
arrived at after pondering all possible out-
comes and implications. Neural-network
research has shown that emotion and reason
are fully integrated in the lateral prefrontal
cortex (LPFC): ‘the convergence of both cog-
nition and affective/motivational information
enables the LPFC to dynamically weigh mul-
tiple lines of information in guiding action’
(Pessoa, 2008: 154). Although emotion and
rationality are two inseparable components
of all that we think and do, neuroscience
informs us that when the two are in conflict,
emotion usually triumphs over reason
(Verweij et al., 2015).
Emotions are assumed to be located in a
set of brain structures called the limbic sys-
tem, which is a multi-structural system of
behavioral regulation that predates the evolu-
tion of the prefrontal cortex structures where
our reasoning power is housed by at least a
million years (Suwa et al., 2009). As Douglas
 EVOLUTION AND PUNISHMENT
Massey (2002: 15) notes: ‘Emotionality
clearly preceded rationality in evolution-
ary sequence, and as rationality developed
it did not replace emotionality as the basis
for human interaction. Rather, rational abili-
ties were gradually added to preexisting and
simultaneously developing emotional capaci-
ties’. Jonathan Haidt (2001: 819) puts it even
more strongly: ‘It [emotion] comes first in
phylogeny, it emerges first in ontogeny, it
is triggered more quickly in real-time judg-
ments, and it is more powerful and irrevoca-
ble [than rationality] when the two systems
yield conflicting judgments’.
Anyone who wishes to challenge the
primacy of emotion over rationality in
making moral decisions will have to confront
the evidence of psychopathy. If relying on
rationality in making punitive decisions is the
mark of a ‘civilized’ being, and introducing
emotion into the equation is irrational and
barbaric, then the callous psychopath must
be the epitome of the civilized being. He/
she is the consummate expert in using his/
her rational faculties – unencumbered by the
emotions of guilt, shame, embarrassment, and
empathy – to connive and manipulate others
into doing his/her bidding, and has been
defined as an ‘obligate cheater’ (da Silva et al.,
2015; Mealey, 1995; Quinsey, 2002). The
psychopath knows the rational words of the
moral song, but not its emotional music. This
results in an entirely utilitarian self-centered
pattern of behavior (Jorgensen et al., 2016).
The defining neurobiological feature of
psychopaths is that the rational and emotional
areas of the brain are poorly integrated (Raine,
2013; Wiebe, 2011). This has been demon-
strated using numerous techniques such as
EEG, PET, fMRI, and even at the molecular
level by diffuse tensor imaging (DTI), which
tracks the movement of molecules along
white-matter tracts to and from the ‘rational’
prefrontal cortex and the ‘emotional’ limbic
system (Craig et al., 2009; Walsh and Bolen,
2012). Psychopaths are fully able to under-
stand the moral norms of society rationally;
it is their lack of understanding of their moral
267
implications supplied by the social emotions
of guilt, shame, empathy, and embarrassment
that explains their activities. Psychopaths
feel more primitive primary emotions such
as anger and fear, but due to the poor con-
nections between the limbic system and the
prefrontal cortex, they are not guided by
rational considerations, just as their rational
considerations are not guided by the social
emotions. When the psychopath does display
emotions, they are indeed ‘confused pas-
sions’ and ‘pathological’.
A retributive punishment justification is the
only justification associated with deep natural
emotions. When people hear of some vicious
criminal act such as the rape and murder of a
young girl, they become angry, outraged, and
disgusted. Their first inclination is to want
to exact some sort of retribution; it is highly
unlikely that their first thoughts should be
of deterrence or rehabilitation. Only a cal-
lous psychopath would shrug his shoulders
upon hearing of such a crime because it had
no impact on his life, which is, of course,
entirely ‘rational’. The rest of us would feel
empathy for the victim and her grieving fam-
ily. However, this retributive urge still tends
to be damned as ‘irrational’, ‘uncivilized’ by
good liberal humanists who find it entirely
inappropriate. Retribution, they rightly com-
plain, is punishment for its own sake from
which nothing good can come, and which
degrades society. Wanting something good to
come from punishment is a utilitarian position
by which the only justification for punishment
is its alleged positive consequences.
Moore (1987) sees this position as hypo-
critical since no objections to emotional out-
rage are offered when it is aimed at unjust
punishment of the innocent. Don’t we all
experience feelings of outrage when we
hear of an innocent man being released from
prison after 20 years through DNA exon-
eration? Most people are emotionally dis-
turbed when they perceive innocent people
are being or have been punished, even if it
has resulted in some positive consequence
for society. Why then do many consider it
268 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
barbaric when the same emotional outrage
is aimed at the acts of guilty people? If the
moral judgment in the first instance is virtu-
ous, then it is virtuous in the second instance
also. The retributionist would say that if you
demand a positive outcome from punishment
for its own sake, then that would be justice
being done. The retributionist would also say
that punishing the innocent, regardless of
what positive consequences may come from
it (such as general deterrence), is impermis-
sible and unjust.
THE ROLE OF EVOLVED EMOTIONS IN
CAPITAL-PUNISHMENT DECISIONS
The issue of capital punishment is a deeply
moral one, and as we have seen, the founda-
tion for morality is more emotional than
rational. The deep emotional underpinning of
morality, and thus the morality we bring with
us to these issues, is generally not appreci-
ated by people who cannot understand why
equally reasonable people can arrive at oppo-
site conclusions regarding the issue of capital
punishment. Two equally intelligent people
can have access to exactly the same informa-
tion on a deep moral issue, but differ radi-
cally as to its meaning based on the emotions
it generates. Capital punishment thus pro-
vides us with a meaningful stage on which to
present the role of emotions in punishment
since the death penalty is the ultimate and
irrevocable punishment.
Support for capital punishment waxes
and wanes in the United States with crime
rates, especially violent crime rates. A 2018
Pew Research Center poll found that 54%
of Americans favor the death penalty while
39% oppose it (Oliphant, 2018). Public sup-
port of the death penalty was at its lowest at
42% in 1966 when the violent crime rate was
200 per 100,000 population, and at 80% in
1994 when the violent crime rate peaked at
713.6 per 100,000 (Hatch and Walsh, 2016:
111). These figures clearly indicate that at
some level people track crime as reported
in the news media and form their attitudes
accordingly. We wish to emphasize that
the following discussion of death penalty
­decision-making implies neither support nor
non-support of capital punishment. It is meant
only to emphasize the role of evolved human
emotions in the juror’s decision, for they must
make it, not the judge. Additionally, we also
note that it is possible to have no objection
to the death penalty on moral grounds, but
object to it on practical grounds. For exam-
ple, the death penalty is much more costly
to our public coffers compared with life
in prison without the possibility of parole.
What is more, our criminal justice system is
imperfect and may have executed an innocent
person in the past and may continue to do so
in the future if the death penalty remains an
option.
Serving on a jury in a case in which the
prosecutor is seeking the death penalty and
knowing that you and 11 other jurors must
vote for life or death must be an exhausting
emotional rollercoaster ride. On what does
one base one’s vote to execute another human
being or spare him or her? During the trial
phase, jurors have learned a bit about the law
and how the judge says they must apply it,
but they have also learned many gruesome
details that have alarmed, shocked, and dis-
gusted them. Rule 403 of the federal Rules of
Evidence allows evidence to be excluded if it
risks unfair prejudice, which ‘means an undue
tendency to suggest decision on an improper
basis, commonly, though not necessarily, an
emotional one’. However, the gloves are off.
When the trial is over and the defendant has
been found guilty beyond a reasonable doubt,
the jurors must then deliberate on the mur-
derer’s sentence after a sentencing hearing
when the full force of emotion is evident.
According to Bandes (2008: 493): ‘When
asked to determine what sort of punishment
heinous murderers deserve, people consult
their moral, ethical, and religious beliefs.
They consult their emotional reactions – their
empathy, disgust, and moral outrage’.
 EVOLUTION AND PUNISHMENT
Bandes (2008: 495) further notes that: ‘The
capital [punishment] system is built upon
choices among competing emotional claims –
some acknowledged and others not’. The
competing emotional claims are abundantly
evident in capital trials. In a capital case,
prosecutors play on the jury’s fear, disgust,
anger, and sympathy for the victim, and ask
that the defendant be sentenced to death for
what he or she has done. Then comes the
defense attorney who asks jurors to consider
the humanity of the defendant, trying to
evoke sympathy by conveying his or her ter-
ribly abusive childhood, and begs for mercy.
After which comes the very tearful testimony
of the families of both the victim and the con-
victed. At the end of it all, the judge instructs
the jury to set their emotions aside and decide
the defendant’s fate by a rational deliberation
of the law and the facts before them, but we
know that they cannot since: Reasoned argu-
ment has limited effect because it is not rea-
soning that prompts the judgement. (Canton,
2015: 68)
Canton (2015) offers us a number of social
emotions and how they engage our evolved
retributive urge, all of which are doubtlessly
engaged during jury deliberations. He notes
that empathetic compassion for the victim and
the victim’s survivors is perhaps the primary
emotion, remarking that ‘retributive emotions
reflect a decent compassion for a victim’s (or
survivor’s) distress and a virtuous expression
of solidarity with members of our community’
(2015: 62). Only the ultimate punishment can
achieve justice for the victim(s) and right the
wrong. Jurors would also focus on the deep
instinct of fairness – Canton asserts that retri-
bution ‘restores a balance: the offender’s unjust
profit or gain from the crime must be redressed
or annulled…The retributivist does not hit,
but hits back’ (2015: 63). There is no turning
of the other cheek for retributionists for it is
only virtuous to turn a cheek if it is one’s own
that was slapped. Rightly or wrongly, the ret-
ributionist views the convicted person as pos-
sessing the free will to make choices, and
the condemned person has made evil ones.
269
The urge to punish can also be seen as loyalty
to the group and an expression of commitment
to it. Offenders are subconsciously viewed
as traitors and enemies of the group – ‘not one
of us’ – which serves to neutralize our feel-
ings of care and compassion for them. By their
crimes, offenders have demonstrated a lack of
respect for authority, which represents the pos-
sibility of the breakdown of the social contract.
This possibility generates anxiety among those
who have a deep respect for authority and social
order and view just deserts as a way of increas-
ing social order. Canton reminds us that crimes
for which the death penalty is sought evoke a
deep sense of negative emotion, especially
when multiple victims, children, and/or torture
are involved. We often hear metaphors for such
criminals as ‘filth’ or ‘scum’ and seeing them
punished described as an act of ‘cleansing’
(Canton, 2015: 66).
There are those who believe that not only
can jurors not eliminate emotion, but also
that they should not:
Perhaps it is not so much that emotion is a key to
normative judgment as it is a key to important and
effective normative judgment, normative judg-
ment that gets our attention and gets translated
into action, either with respect to our own conduct
or to the reward or punishment of others.
(Goodenough and Prehn, 2004: 1717)
We should expect emotions to render the
most effective moral judgments given their
long evolutionary history as the only basis
for hominid social interaction prior to the
evolution of our vaunted rationality. If these
‘effective moral judgments’ are more often
than not retributive when it comes to moral
decisions about punishing transgressors, one
may wonder why Kant’s proportional retri-
bution is often viewed so negatively.
SECOND- AND THIRD-PARTY
PUNISHMENT
It is not always possible to punish those who
harm us and we must turn to others to carry
270 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
the burden. However, why would others take
on your risky burden with no apparent payoff
for them? Among chimpanzees, alpha males
take on the role of what is called control
behavior, which includes the punishment of
troop members who bully and exploit others
(de Waal, 1996). This is a costly and risky role,
but it confers a number of benefits to the alpha
male. As an arbiter and punisher, alpha males
typically show a preference for the weaker
party in most disputes. This develops support
among the weaker regular members and serves
to level the hierarchy. Leveling the hierarchy
increases the gap between the alpha male and
the more powerful members of the group who
might seek to replace him (de Waal, 1996).
Maintaining one’s position as the alpha male
affords him status, allowing him ‘first dibs’ on
the females when they come into estrus. A host
of computer simulation studies of human altru-
istic punishment (punishment on behalf of
others) have conclusively shown that the pun-
isher receives many benefits, including an
increased likelihood of receiving future bene-
fits, with enhanced status in the group being
the most valuable because it leads to enhanced
fitness (Ule et  al., 2009). Dos Santos et  al.
(2011) remark that: “reputation is the key to
the evolution of punishment, and that simple
reputation games can explain the high preser-
vation of punishment in humans” (2011: 376).
Leading theories of third-party punishment
include inequity aversion and strong reciproc-
ity. Inequity aversion favors fairness and rejects
inequitable outcomes. Humans (and primates)
are willing to forgo benefits to themselves if
they perceive others receiving a greater and
unfair reward. This rejection of injustice helps
to punish cheats and stabilize cooperation
among group members. Strong reciprocity
refers to the tendency to cooperate with group
members even when there are no immediate
benefits of doing so. It is therefore advanta-
geous to the group to punish non-cooperators
and to use punishment to ensure fairness.
In modern societies, punishment is meted
out by third-party punishers who are indi-
viduals not directly harmed and who will
not directly benefit (like second-party chimp
alpha males) from meting out punishment.
This type of punishment is more impartial
and less egocentric than second-party pun-
ishment, thereby minimizing the likelihood
of a retaliatory tit-for-tat feedback loop (Fehr
and Fischbacher, 2004). However, there is
some evidence to suggest that third parties
do not punish differently than second par-
ties (Leibbrandt and Lopez-Perez, 2012).
Third-party punishers in modern societies
are typically agents of the state operating
in accordance with the law. For example,
offenders are caught by police, prosecuted
in court, and then sentenced by judges.
However, other informal third-party pun-
ishments may be doled out to offenders by
school administrators or church officials.
Informal third-party punishment occurs more
frequently than state-sponsored third-party
punishment and is essential to the enforce-
ment of norms (Bendor and Swistak, 2001).
Numerous experiments have shown that third
parties will punish cheats at a cost to them-
selves. A study involving 1,762 subjects from
five continents found that in all populations,
people are willing to punish defectors who
have harmed unknown others (Henrich et al.,
2006). This study also found that ‘societies
with high degrees of punishment will also
exhibit more altruistic behavior’ (Henrich
et  al., 2006: 1770). This tends to suggest
altruism and punishment coevolved in the
sense that ‘Third-party punishment of norm
violations (‘I punish you because you harmed
him’) seems especially crucial for the evo-
lutionary stability of cooperation and is the
cornerstone of modern models of criminal
justice’ (Buckholz and Marois, 2012: 655).
FROM PRIMITIVE VENGEANCE
TO MODERN LAW
Just as we can become alcoholics, obese, and
sex addicts partaking too freely in what feels
good, or even vital in the right proportions,
 EVOLUTION AND PUNISHMENT
punishment can overstep its optimum and
become dysfunctional. In neurological imag-
ing studies showing blood flow to the brain’s
pleasure center, when people witness the
punishment of those who have harmed them,
the punishment tends to be proportional to
the harm caused. Only a callous sadist would
take pleasure in seeing grossly disproportion-
ate punishment imposed. Punishment that
exceeds just and reasonable boundaries
induces disgust and repugnance, and many
people feel that the death penalty is one such
excessive punishment.
Even given the well-founded evolutionary
reasons for retributionist feelings, we must
reflect on the propriety of acting on them
completely. Just because these feelings are
natural, it does not mean that we should act
on them and ignore rational consequential-
ist considerations. If the primitive desire to
‘get even’ is left untamed, it can tear a social
group apart by generating a cycle of tit-for-
tat blood feuds, which have smeared human
history (Boehm, 2011). It has been estimated
that approximately 30% of adult male deaths
among the Yanomamo of South America are
related to revenge feuds, which expand the
very injustice that ‘righteous’ revenge was
supposed to assuage (Chagnon, 1988). As
Susan Jacoby (1983: 13) put it:
The struggle to contain revenge has been con-
ducted at the highest level of moral and civic
awareness at each stage in the development of
civilization. The self-conscious nature of the effort
is expectable in view of the persistent state of ten-
sion between uncontrolled vengeance as destroyer
and controlled vengeance as an unavoidable com-
ponent of justice.
Just as evolutionary scientists view cheating
and punishment as vital to the evolution of
cooperation, Durkheim saw it as necessary for
maintaining social solidarity, and that it does
so by reaffirming the justness of the social
norms. He recognized that the urge to punish
is inherent in human nature and that it serves
an expiatory role, but he also recognized that
we could temper the urge with sympathy.
Over the course of human history, Durkheim
271
noted that many societies have moved from
retributive to restitutive justice. For Durkheim
(1964), retributive justice is driven by the
natural passion for punitive revenge that
‘ceases only when exhausted … only after it
has destroyed’ (1964: 86). Restitutive justice,
on the other hand, is driven by simple deter-
rence, and is more humanistic and tolerant,
although it is still ‘at least in part, a work of
vengeance’ since it is still ‘an expiation’
(1964: 88–89). While both forms satisfy the
human urge for social regularity, repressive
justice oversteps its adaptive usefulness and
becomes socially destructive, while restitutive
justice offers a rational balance between calm-
ing moral outrage on the one hand, and engag-
ing empathy and sympathy on the other.
Culture may engage or neutralize the emo-
tions that temper punishment with mercy or
allow vengeance to run wild. The lasting influ-
ence of Cesare Beccaria rests on his recogni-
tion that the brutal acts of retribution that were
common in the 18th century resulted in gen-
eral distrust of powerful institutions and social
alienation rather than altruistic cooperation. In
other words, Beccaria argued that excessive
punishment is indicative of tyranny and the
associated lack of perceived legitimacy accom-
panying tyrannical institutions fails to inspire
cooperation. Many of Beccaria’s recommended
criminal justice reforms were implemented
throughout much of Europe within his lifetime
(Durant and Durant, 1967). Such radical and
rapid change suggests that Beccaria’s ideas
tapped into and broadened other evolved emo-
tions such as sympathy and empathy among
the European elite. We tend to feel empathy
for those whom we view as being ‘like us’,
and empathy often leads to sympathy, which
may translate the vicarious experiencing of the
pains of others into an active concern for their
welfare, even if they are wrongdoers. Vignette
studies have shown that people tend to recom-
mend more lenient punishment for criminals
whom they perceive to be similar to themselves
(reviewed in Miller and Vidmar, 1981), and the
march of democracy has drawn more people
into the circle of people we consider ‘us’.
272 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
RECONCILIATION AND
REINTEGRATION: FORGIVENESS
TIT-FOR-TAT
Just as social emotions such as empathy may
lead to more lenient sentencing, the existence
of this emotion may increase the likelihood for
forgiveness of offenders (Batson and Ahmad,
2001; Batson and Moran, 1999). This empathy-
induced altruism has some evolutionarily
advantageous outcomes that benefit social
groups. In societies where cheating is likely to
occur, forgiveness tit-for-tat operates to benefit
social groups by avoiding social actions that
degrade into absolute punishment. If two strict
tit-for-tat strategists play one another and one
cheats (purposely or otherwise), it tends to
launch a long series of mutual punitive
response resulting in a loss for both. Work in
game theory indicates that a measure of for-
giveness may work better to maintain social
cooperation rather than always paying back
defectors in kind (Rand et al., 2009). If people
play strict tit-for-tat (always punishing non-
cooperation in kind) in their relationships they
risk losing a valuable relationship if the cheat-
ing incident was uncharacteristic of the person,
or accidental (Wagstaff, 1998). As Machalek
and Cohen (1991: 221) put it: ‘Forgiving strat-
egies can transform a pattern of mutual cheat-
ing by not remaining permanently punitive but
instead by cooperating, even when the other
strategy cheats’. A large number of game-theory
studies have found that strict tit-for-tat can
amount to punishment that is just too costly for
the punisher, that all parties tend to suffer to
some extent, and that the generous (forgiving)
tit-for-tat strategy invariably prevails (Rand
et  al., 2009). Although some social emotions
may drive this behavior across generations,
findings focused on other social emotions such
as compassion muddy these waters. Weng
et  al. (2015) found that while compassion
enhances giving to victims, it does not affect
punishment of transgressors.
The generous tit-for-tat strategy forgives a
single betrayal by responding with cooperation
rather than retaliation in the next round of the
game, and in most circumstances, it will return
the players to cooperation. Of course, we are
dealing with game theory here, with its ideal-
ized laboratory conditions and assumption of
rational actors; the real world is expected to
be far more complicated. However, the predic-
tions based on game-theoretical models have
been remarkably consistent with real-world
findings on numerous occasions (Levitt and
List, 2007). Cosmides and Tooby (1992) offer
the example of the Ache tribe of Paraguay
(and certain other hunter/gatherer tribes) who
respond differently to hunters who cheat than
to gatherers who cheat. Meat is a scarce and
valued resource, and is shared equally by all
when available, which is often a matter of luck
and hunting skills. Plant food is a low-variance
item, the availability of which depends only
on the effort spent on gathering it. Arguments
erupt when plant gatherers are perceived as not
pulling their weight. Punishing lazy gatherers
has no adverse effect on the availability of plant
food; it is still there for the gathering. Given
the highly variable nature of meat acquisition,
however, tribal members are more forgiving of
hunters whom they perceive as cheating. It is
recognized that a charge of cheating can be the
result of a false perception (hunting is often
a matter of sheer luck as much as effort). If
because of the false charge the hunter is pun-
ished by ostracism, the whole tribe will lose
a valuable cooperator in the tricky business
of hunting for meat, and the meat supply will
become less stable and predictable.
Continuing to invite cooperation in the face
of defection is not a ‘sucker’ strategy because
it is not one that tolerates continued cheat-
ing. It is similar to sentencing an offender to
probation rather than prison, thus leaving the
door open to future mutually advantageous
cooperation between offender and com-
munity. The ‘forgiving tit-for-tat’ strategy
is captured most familiarly in Braithwaite’s
(1989) concept of reintegrative shaming.
It is Braithwaite’s position that we should
retaliate against tit-for-tat defectors, but that
retaliation should be conciliatory rather than
 EVOLUTION AND PUNISHMENT
permanently punitive. Punishment (and the
shame accompanying it) should be reintegra-
tive and motivate offenders to evaluate them-
selves and their behavior, not disintegrative.
Both the offender and the community benefit
by methods used to express our disapproval
of offenders that conforms to the reintegra-
tive ideal. If we always respond to defecting
with strong punitive reactions, the commu-
nity loses a potentially valuable cooperator,
and the offender becomes, to his or her great
disadvantage, alienated from it.
Reconciliatory behavior is common
among primate species, which indicates
that forgiving tit-for-tat had some positive
fitness consequences. In restorative-justice
programs, victims who confront their vic-
timizers in controlled settings and offer for-
giveness to them report feeling positive (a
sense that justice had been done) about the
experience (reviewed in Latimer et al., 2005).
Satisfaction with restorative justice is, how-
ever, contingent on procedural factors (trust,
neutrality, respect, etc.) as well as situational
factors such as flexibility and provisions of
care for the victim (Van Camp and Wemmers,
2013). Thus, even forgiveness has rather
strong elements of tit-for-tat.
Restorative-justice principles that tap an
offender’s capacity for empathy for their vic-
tim are most fully implemented by juvenile
justice agencies. Juvenile agencies implicitly
recognize that delinquent behavior is normal
behavior; indeed, the young male who does
not engage in some sort of delinquent behav-
ior is statistically abnormal (Moffitt, 1993).
Adolescence and young adulthood is a period
of intense inter-male competition ultimately
(if not always consciously) aimed at securing
more mating opportunities than the next male
(McKibbin and Shackleford, 2011; Wiebe,
2011). To adopt a strategy of strict tit-for-tat
rather than forgiveness tit-for-tat with juve-
nile offenders, whose physical desires and
abilities have temporarily outrun their neu-
rological maturity, would be severely coun-
terproductive. Although the evolutionary
benefits are greater when applied to juvenile
273
offenders, the perceived satisfaction with
restorative justice is less pronounced in juve-
niles as compared to adults (Gal and Moyal,
2011; Kim and Gerber, 2012). Because of
its ability to foster more cooperative socie-
ties however, forgiveness tit-for-tat in game
theory provides a degree of evolutionary and
mathematical justification for restorative
justice, which is the only corrections model
aimed at reintegrating offenders back into the
community with the help of the community.
REFERENCES
Adolphs, R. (2009). The social brain: Neural
basis of social knowledge. Annual Review of
Psychology, 60, 693–716.
Ash, J., & Gallup, G. G. (2007). Paleoclimatic
variation and brain expansion during human
evolution. Human Nature, 18(2), 109–124.
Baden, J., & Stroup, R. (1972). Choice, faith, and
politics: The political economy of Hutterian
communes. Public Choice, 12, 1–11.
Bailey, D. H., & Geary, D. C. (2009). Hominid
brain evolution. Human Nature, 20(1),
67–79.
Bandes, S. (2008). Repellent crimes and rational
deliberation: Emotion and the death penalty.
Vermont Law Review, 33, 489–519.
Barkow, J. H. (1989). The elastic between
genes and culture. Ethology and Sociobiology,
10(1–3), 111–129.
Batson, C. D., & Ahmad, N. (2001). Empathy-
induced altruism in a prisoner’s dilemma II:
What if the target of empathy has defected?
European Journal of Social Psychology,
31(1), 25–36.
Batson, C. D., & Moran, T. (1999). Empathy-
induced altruism in a prisoner’s dilemma.
European Journal of Social Psychology,
29(7), 909–924.
Bendor, J., & Swistak, P. (2001). The evolution
of norms. American Journal of Sociology,
106, 1493–1545.
Boehm, C. (2011). Retaliatory violence in
human prehistory. British Journal of
Criminology, 51, 518–534.
Bohm, R. (2012). Deathquest: An introduction
to the theory and practice of capital
274 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
punishment in the United States. Boston,
MA: Anderson.
Braithwaite, J. (1989). Crime, shame, and
reintegration. Cambridge: Cambridge
University Press.
Brunero, J. (2002). Evolution, altruism and
internal reward explanations. Philosophical
Forum, 33, 413–424.
Buckholtz, J., & Marois, R. (2012). The roots of
modern justice: Cognitive and neural
foundations of social norms and their
enforcement. Nature Neuroscience, 13,
655–661.
Buonanno, P., & Raphael, S. (2013).
Incarceration and incapacitation: Evidence
from the 2006 Italian collective pardon.
American Economic Review, 103(6),
2437–2465.
Canton, R. (2015). Crime, punishment and the
moral emotions: Righteous minds and their
attitudes towards punishment. Punishment
& Society, 17, 54–72.
Chagnon, N. A. (1988). Life histories, blood
revenge, and warfare in a tribal population.
Science, 239(4843), 985–992.
Choy, O., Farrington, D. P., & Raine, A. (2015).
The need to incorporate autonomic arousal
in developmental and life-course research
and theories. Journal of Developmental and
Life-Course Criminology, 1(2), 189–207.
Cosmides, L., & Tooby, J. (1994). Better than
rational: Evolutionary psychology and the
invisible hand. The American Economic
Review, 84, 327–332.
Cosmides, L., & Tooby, J. (1992). Cognitive
adaptations for social exchange. The
Adapted Mind: Evolutionary Psychology and
the Generation of Culture, 163, 163–228.
Craig, M., Catani, M., Deeley, Q., Latham, R.,
Daly, E., Kanaan, R., Picchioni, M., McGuire,
P., Fahy, T., & Murphy, D. (2009). Altered
connections on the road to psychopathy.
Molecular Psychiatry, 14, 946–953.
Currie, E. (1999). Reflections on crime and
criminology at the millennium. Western
Criminology Review, 2(1), 1–13.
Da Silva, D. R., Rijo, D., & Salekin, R. T. (2015).
The evolutionary roots of psychopathy.
Aggression and Violent Behavior, 21, 85–96.
Darwin, C. (1896). The descent of man and
selection in relation to sex (Vol. 1). D. New
York: Appleton.
De Quervain, D., Fischbacher, U., Valerie, T.,
Schellhammer, M., Schnyder, U., Buch, A., &
Fehr, E. (2004). The neural basis of altruistic
punishment. Science, 305, 1254–1259.
De Waal, F. (1996). Good natured: The origins
of right and wrong in humans and other
animals. Cambridge, MA: Harvard University
Press.
De Waal, F. (2008). Putting the altruism back
into altruism: The evolution of empathy.
Annual Review of Psychology, 59, 279–300.
Dos Santos, M., Rankin, D., & Wedekind, C.
(2011). The evolution of punishment through
reputation. Proceedings of the Royal Society:
Biology, 278, 371–377.
Dunbar, R. I., & Shultz, S. (2007). Evolution
in the social brain. Science, 317(5843),
1344–1347.
Durant, W., & Durant, A. (1967). Rousseau and
revolution. New York: Simon and Schuster.
Durkheim, E. (1964). The division of labor in
society. Glencoe, IL: Free Press.
Farrow, T., & Woodruff, P. (2007). Empathy in
mental illness. Cambridge: Cambridge
University Press.
Fehr, E., & Fischerbacher, U. (2004). Third party
punishment and social norms. Evolution and
Human Behavior, 25, 63–87.
Furman vs. Georgia. 408 U.S. 238 (1972).
Gal, T., & Moyal, S. (2011). Juvenile victims in
restorative justice: Findings from the
reintegrative shaming experiments. The
British Journal of Criminology, 51(6),
1014–1034.
Gavrilets, S. (2012). On the evolutionary origins
of the egalitarian syndrome. Proceedings of
the National Academy of Sciences, 109(35),
14069–14074.
Goodenough, O., & Prehn, K. (2004). A
neuroscientific approach to normative
judgment in law and justice. Philosophical
Transactions of the Royal Society of London:
Biology, 359, 1709–1726.
Gottfredson, M., & Hirschi, T. (1990) A general
theory of crime. Stanford University Press.
Haidt, J. (2001). The emotional dog and its
rational tail: A social intuitionalist approach
to moral judgment. Psychological Review,
108, 814–834.
Haidt, J. (2012). The righteous mind: Why
good people are divided by politics and
religion. New York: Pantheon.
 EVOLUTION AND PUNISHMENT
Hatch, V., & Walsh, A. (2016). Capital
punishment: The theory and practice of the
ultimate penalty. New York: Oxford University
Press.
Hawthorne, N. (1879). The scarlet letter. The
Blithedale romance (Vol. 9). New York:
Houghton, Osgood.
Henrich, J., McElreath, R., Barr, A., Ensminger, J.,
Barrett, C., Bolyanatz, A., Cardenas, J.,
Gurven, M., Gwako, E., Henrich, N.,
Lesorogot, C., Marlowe, F., Tracy, D., & Ziker,
J. (2006). Costly punishment across human
societies. Science, 312, 1767–1770.
Hill, R. A., & Dunbar, R. I. (2003). Social network
size in humans. Human Nature, 14, 53–72.
Jacoby, S. (1983). Wild justice: The evolution of
revenge. New York: Harper & Row.
Jorgensen, C., Anderson, N., & Barnes, J. C.
(2016). Bad brains: Crime and drug abuse
from a neurocriminological perspective. Ameri-
can Journal of Criminal Justice, 41(1), 47–69.
Kanazawa, S. (2008). Temperature and
evolutionary novelty as forces behind the
evolution of general intelligence. Intelligence,
36(2), 99–108.
Kant, I. (1964). Groundwork of the metaphysics
of morals. Trans. by H. J. Paton. New York:
Harper & Row.
Kant, I. (1965). The metaphysical elements of
justice. Indianapolis, IN: Bobbs–Merrill.
Kim, H. J., & Gerber, J. (2012). The effectiveness
of reintegrative shaming and restorative jus-
tice conferences: Focusing on juvenile offend-
ers’ perceptions in Australian reintegrative
shaming experiments. International Journal
of Offender Therapy and Comparative Crimi-
nology, 56(7), 1063–1079.
Klein, R. (2012). The neurobiology of altruistic
punishment: A moral assessment of its social
utility. In K. Plaisance & T. Reydon (Eds.),
Philosophy of Behavioral Biology, pp. 297–
313. Boston, MA: Springer Science.
Kochanska, G., & Aksan, N. (2004).
Development of mutual responsiveness
between parents and their young children.
Child Development, 75(6), 1657–1676.
Latimer, J., Dowden, C., & Muise, D. (2005).
The effectiveness of restorative justice
practices: A meta-analysis. The prison
Journal, 85(2), 127–144.
Leibbrandt, A., & Lopez-Perez, R. (2012). An
exploration of third and second party
275
punishment in ten simple games. Journal of
Economic Behavior and Organization, 84,
753–766.
Levitt, S. D., & List, J. A. (2007). What do
laboratory experiments measuring social
preferences reveal about the real world?
Journal of Economic Perspectives, 21(2),
153–174.
Logan, C., & Gaes, G. (1993). Meta-analysis
and the rehabilitation of punishment. Justice
Quarterly, 10, 245–263.
Machalek, R., & Cohen, L. (1991). The nature of
crime: Is cheating necessary for cooperation?
Human Nature, 2, 215–233.
Mackie, J. L. (1982). Morality and the retributive
emotions. Criminal Justice Ethics, 1, 3–10.
Massey, D. (2002). A brief history of human
society: The origin and role of emotion in
social life. American Sociological Review, 67,
1–29.
McBride, K. (2007). Punishment and the
political order. Ann Arbor, MI: University of
Michigan Press.
McKibbin, W. F., & Shackelford, T. K. (2011).
Women’s avoidance of rape. Aggression and
Violent Behavior, 16(5), 437–443.
Mealey, L. (1995). The sociobiology of
sociopathy: An integrated evolutionary
model. Behavioral and Brain Sciences, 18(3),
523–541.
Miller, D. T., & Vidmar, N. (1981). The justice
motive in social behavior. Boston, MA:
Springer.
Moffitt, T. E. (1993). Life-course-persistent and
adolescence-limited antisocial behavior: A
developmental taxonomy. Psychological
Review, 100(4), 674–701.
Moore, M. (1987). The moral worth of
retribution. In F. Schoeman (Ed.),
Responsibility, character, and the emotions:
New essays in moral psychology, pp. 179–
219. Cambridge: Cambridge University Press.
Nowak, M. (2006). Five rules for the evolution
of cooperation. Science, 314, 1560–1563.
Nowak, M., & Sigmund, K. (2005). Evolution of
indirect reciprocity. Nature, 437,
1291–1298.
Oliphant, B. (2018, June 11). Public support for
the death penalty ticks up. Pew Research
Center. www.pewresearch.org/fact-tank/2018/
06/11/us-support-for-death-penalty-ticks-
up-2018/ (Accessed 7 June 2019)
276 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Penney, S. (2012). Impulse control and criminal
responsibility. International Journal of Law
and Psychiatry, 35, 99–103.
Pessoa, L. (2008). On the relationship between
emotion and cognition. Nature/Neuroscience,
9, 148–158.
Petersen, M. B., Sell, A., Tooby, J., & Cosmides,
L. (2010). Evolutionary psychology and
criminal justice: A recalibrational theory of
punishment and reconciliation. In Human
morality & Sociality: Evolutionary &
comparative perspectives (Eds. Henrik Hogh-
Oleson). Palgrave MacMillan: New York.
Quinsey, V. L. (2002). Evolutionary theory and
criminal behaviour. Legal and Criminological
Psychology, 7(1), 1–13.
Rachels, J. (1986). The elements of moral
philosophy. New York: Random House.
Radzinowicz, L., & King, J. (1979). The growth
of crime. Harmondsworth: Penguin.
Raine, A. (2013). The anatomy of violence: The
biological roots of crime. New York:
Pantheon.
Rand, D. G., Ohtsuki, H., & Nowak, M. A.
(2009). Direct reciprocity with costly
punishment: Generous tit-for-tat prevails.
Journal of Theoretical Biology, 256(1), 45–57.
Rosebury, B. (2009). Private revenge and its
relation to punishment. Utilatus, 21, 1–21.
Stohr, M. K., Walsh, A., & Hemmens, C. (2012).
Corrections: A text/reader (Vol. 3). Thousand
Oaks, CA: Sage.
Suwa, G., Asfaw, B., Kono, R., Kubo, D., Lovejoy,
C., & White, T. (2009). The Ardipithecus
ramidus skull and its implications for hominid
origins. Science, 326, 68e1–68e8.
Ule, A., Schram, A., Riedl, A., & Cason, T. N.
(2009). Indirect punishment and generosity
toward strangers. Science, 326(5960),
1701–1704.
Van Camp, T., & Wemmers, J. A. (2013). Victim
satisfaction with restorative justice: More
than simply procedural justice. International
Review of Victimology, 19(2), 117–143.
Verweij, M., Senior, T., Dominquez, D., &
Turner, R. (2015). Emotion, rationality, and
decision-making: How to link affective and
social neuroscience with social theory.
Frontiers in Neuroscience, 9, 1–13.
Wagstaff, G. (1998). Equity, justice, and
altruism. Current Psychology, 17, 111–134.
Walsh, A. (2006). Evolutionary psychology and
criminal behavior. In J. Barkow (Ed.), Missing
the revolution: Darwinism for social scientists,
pp. 225–268. Oxford: Oxford University Press.
Walsh, A., & Bolen, J. (2012). The neurobiology
of criminal behavior: Gene-brain culture
co-evolution. Farnham: Ashgate.
Walsh, A., Johnson., H., & Bolen., J. (2012).
Drugs, crime, and the epigenetics of hedonic
allostasis. Journal of Contemporary Criminal
Justice, 28, 314–328.
Weng, H. Y., Fox, A. S., Hessenthaler, H. C.,
Stodola, D. E., & Davidson, R. J. (2015). The
role of compassion in altruistic helping and
punishment behavior. PLoS ONE, 10(12),
e0143794.
Wiebe, R. (2011). The nature and utility of low
self-control. In K. Beaver & A. Walsh (Eds.),
The Ashgate research companion to biosocial
theories of crime, pp. 369–395. Farnham:
Ashgate.
Wilkinson, G. S. (1990). Food sharing in vampire
bats. Scientific American, 262(2), 76–83.
Wilson, J. Q. (1975). The rise of the bureaucratic
state. The Public Interest, 41(3), 77–103.

Evolutionary Psychology and
Corrections and Rehabilitation
Ian A. Silver and Jamie Newsome
INTRODUCTION
Identifying effective strategies for managing
and responding to crime is a priority for
nations around the world. Scholars, policy-
makers, and practitioners continually work to
develop crime-control policies that will
enhance public safety and reduce expenditures.
Meeting these goals has become critical in the
United States – particularly in the correctional
system where the number of individuals under
some form of supervision has exceeded
6.6 million adults (Kaeble and Cowhig, 2018).
The dominant perspective on ‘what works’ in
offender rehabilitation, the risk-need-respon-
sivity (RNR) model, has been widely used
throughout North America and in other regions
throughout the world. It combines knowledge
gained from empirically supported theories
that offer explanations of criminal behavior
with practical recommendations to guide
policy and practice (Bonta and Andrews,
2017). Moreover, it is supported by a wealth of
empirical evidence (Smith et al., 2009b).
14
The impact of the RNR model on the field
of corrections has been tremendous, but the
pursuit to achieve even greater reductions
in recidivism continues. Much of this effort
has been focused on refining the practical
application of the model. However, some
scholars have suggested that important the-
oretical advancements have taken place in
recent decades, and that these developments
should be explored as potential avenues for
informing continued work in rehabilitation
(Newsome and Cullen, 2017). Explanations
of behavior grounded in an evolutionary psy-
chology framework have expanded, and may
now present another opportunity to enhance
our understanding of antisocial behavior and
how individuals respond to interventions
(Bakermans-Kranenburg and van IJzendoorn,
2015). This chapter explores this possibility
by first providing an overview of the state
of corrections in the United States, followed
by a brief description of the RNR model of
offender rehabilitation. The core ideas of an
evolutionary psychological perspective are
278 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
then summarized, and three theories based
on this perspective – biological sensitivity
to context theory, differential susceptibility
theory, and intrasexual competition – are dis-
cussed with an emphasis on how they might
inform corrections.
CORRECTIONS IN THE UNITED STATES
The correctional system in the United States
monitors individuals detained in institutions,
such as jails and prisons, as well as those
who are supervised in the community, such
as probationers or parolees. The total number
of individuals in the system grew rapidly
beginning in the 1980s (Bureau of Justice
Statistics, 2016) and continued until it
reached a peak of 7,339,600 in 2007 (Bureau
of Justice Statistics, 2016). This period is
often referred to as the ‘era of mass incar-
ceration’, during which the rise in the use of
prisons to sanction offenders was so extreme
and persistent some have questioned whether
the United States may have become ‘addicted
to incarceration’ (Pratt, 2019). Others have
argued that recent decades may be more
accurately described as an ‘era of mass cor-
rections’ given the simultaneous rise in com-
munity corrections populations (Mears and
Cochran, 2015). The number of individuals
on some form of community supervision
reached a high of 5,119,000 in 2007 and has
only slowly declined to 4,650,900 in recent
years (Kaeble and Cowhig, 2018).
The growth in the correctional system
represents a shift towards a more punitive
strategy. Increased surveillance and punish-
ment were intended to decrease recidivism
and crime more generally. Recent estimates,
however, indicate that those who become
involved with the criminal justice system
may not subsequently redirect their lives
to a more prosocial path. More than half of
the inmates who are released are returned
to prison within three years (Durose et  al.,
2014). Perhaps even more alarming, a recent
report by Alper et  al. (2018) indicated that
44% of state prisoners were rearrested in the
first year after they were released, and by nine
years post-release 83% had been rearrested.
One trend that may have contributed to these
high failure rates is a reduction in rehabilita-
tive initiatives (Mears and Cochran, 2015).
Some reports suggest that during the rise in
the reliance on the correctional system lower
percentages of inmates received treatment
services relative to earlier periods (Lynch and
Sabol, 2001; Phelps, 2011). Moreover, the
majority of inmates may not be participating
in treatment services before they leave prison
(Lawrence et al., 2002; Phelps, 2011).
The size of the corrections populations,
high failure rates, and lack of treatment rep-
resents a complex problem. Fortunately, an
abundance of empirical evidence has accu-
mulated that suggests strategies known as the
principles of effective intervention may be
the most promising tactics for reducing recid-
ivism – whether an individual is confined in
an institution or supervised in the commu-
nity (Bonta and Andrews, 2017; Smith et al.,
2009b). This model, which is described in
the following section, continues to be stud-
ied and refined in hopes of further enhancing
policy and practice in corrections.
The Risk-Need-Responsivity (RNR)
Model of Offender Rehabilitation
Offender rehabilitation has come to be pre-
dominantly founded on the principles of
effective intervention, a set of guidelines for
providing treatment that maximizes the
potential to achieve reductions in recidivism
(Bonta and Andrews, 2017; Smith et  al.,
2009b). Central among these principles are
risk, need, and responsivity (RNR), which
are collectively referred to as the ‘RNR
Model’. Organizations throughout the world
have adopted this model as the framework
for providing services to offenders, and it is
viewed as the leading strategy in rehabilita-
tion (Cullen and Jonson, 2012; Newsome and
 EVOLUTIONARY PSYCHOLOGY AND CORRECTIONS AND REHABILITATION
Cullen, 2017; Ogloff and Davis, 2004;
Polaschek, 2012).
To align practice with the risk principle,
corrections agencies are advised to assess each
individual’s likelihood of committing another
crime and focus the most intensive interven-
tion efforts towards those deemed to be at the
highest risk of recidivating (Andrews, Bonta
et al., 1990; Andrews, Zinger et al., 1990). To
aid practitioners in adhering to this principle,
several actuarial risk-assessment tools have
been developed that capture the presence
of static and dynamic risk factors, quantify
them, and generate an overall risk score and
classification (e.g., low risk, moderate risk,
and high risk). Static factors are stable over
time, such as criminal history, and should
not be identified as targets for treatment.
Dynamic factors that heighten the risk for
recidivism can be changed through effective
interventions and provide insights practition-
ers can use to develop case plans. Scholars
and practitioners often collaborate to exam-
ine the reliability and predictive validity of
such tools and make refinements based on
emerging research; however, evidence from
numerous studies has suggested that modern
tools can improve predictions of recidivism
and case-management strategies while indi-
viduals are involved with the justice system
(Andrews et al., 2004; Gendreau et al., 2002;
Smith et al., 2009a).
Many of the dynamic risk factors identi-
fied through the risk-assessment process are
conceptualized in the need principle as ‘crim-
inogenic needs’ that should be the targets of
interventions (Bonta and Andrews, 2017).
Criminogenic needs include antisocial atti-
tudes, antisocial associates, antisocial tem-
perament/personality, family circumstances,
employment, substance use, and leisure
activities that may promote offending behav-
iors (Andrews, 2006). Focusing rehabilitative
efforts on these factors is the most strategic
method for reducing the likelihood of recidi-
vism. Translating this into practice should
include regular reassessments to monitor pro-
gress in treatment.
279
In addition to determining who and what to
target in offender rehabilitation, the principles
of effective intervention also provide guidance
on how to approach treatment. Specifically, the
responsivity principle instructs practitioners
to adopt treatment strategies known to be the
most effective overall (i.e., general responsiv-
ity), and to address individual differences that
could impact the effectiveness of the treatment
strategies (i.e., specific responsivity) (Bonta
and Andrews, 2017). With regard to offender
populations, research has suggested that strat-
egies based on cognitive behavioral therapy
tend to be the most effective (Andrews,
Zinger et al., 1990; Lipsey and Cullen, 2007;
Lipsey et al., 2007; Lipsey and Wilson, 1998;
Smith et al., 2009b). Specific responsivity fac-
tors, on the other hand, vary widely and may
include things like low motivation to change,
language barriers, or a history of trauma. To
fully adhere to the responsivity principle,
treatment providers must recognize these
factors and develop solutions to minimize any
barriers to success in treatment.
The remaining principles of effective inter-
vention build upon the risk, need, and respon-
sivity principles to form a comprehensive
framework for establishing offender rehabili-
tation programs (Bonta and Andrews, 2017).
Overall, these principles have been empiri-
cally supported through several systematic
reviews and meta-analyses (Andrews, Zinger,
et al., 1990; Gendreau, 1996; Gendreau et al.,
1996, 2006; Lipsey and Cullen, 2007; Lipsey
et al., 2007; Lipsey and Wilson, 1998; Smith
et  al., 2009a, 2009b). Moreover, the wide-
spread adoption and robust body of evidence
in support of the model has been attributed to
the strong theoretical basis, the psychology
of criminal conduct, upon which the model
was created (Bonta and Andrews, 2017).
Using Theory to Inform
Correctional Policy and Practice
Developing policy and practice with careful
consideration for theories that emerge as
280 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
well-supported explanations of offending
behavior may optimize the impact of interven-
tions (Bonta and Andrews, 2017; Mears and
Cochran, 2015; Taxman and Friedmann,
2009). The psychology of criminal conduct
has emphasized the importance of using
theory to better understand individual differ-
ences, particularly those that could readily be
used in practical application (Bonta and
Andrews, 2017). This has largely included
explanations of antisocial behavior that
emerged in psychology or criminology, with a
particular emphasis on a general psychology
and cognitive social learning approach.
Briefly, this perspective views different behav-
iors as being learned, maintained, or changed
through cues that signal a consequence that
may follow an action and the consequences
themselves. This logic can then be extended to
theorize that manipulating the cues and conse-
quences associated with a behavior can be a
useful strategy in eliminating behaviors that
are problematic and promoting the continua-
tion of those that are more desirable (Bonta
and Andrews, 2017).
A number of theories of criminal or antiso-
cial behaviors have identified links between
individual characteristics and lifestyle factors
and offending, which could lend further insights
and support to the RNR model. Sampson and
Laub’s (1993) age-graded theory represents
one criminological theory that complements
the RNR model. Their theoretical proposition
argues that variation of social bonds over time
influences variation in criminal behavior
throughout the lifecourse. As such, the weak-
ening of social bonds is expected to increase
involvement in criminal behavior, while the
strengthening of social bonds is speculated to
encourage individuals to desist from criminal
behavior. For instance, as one reaches late
adolescence or adult stages of development,
obtaining meaningful employment – one of
the central criminogenic needs identified
in the RNR model – can be a critical turn-
ing point that encourages the adoption of a
prosocial lifestyle. A job consumes free time,
provides an individual with opportunities to
establish prosocial connections, and can bring
about a number of benefits that one would not
want to lose. Sampson and Laub (1993) fur-
ther speculated that it is not likely that sim-
ply obtaining a job will divert someone away
from a life of crime; the job should be of some
value to the individual (i.e., add favorable con-
sequences). While this aspect of the theory
does not necessarily point to a new risk factor
or criminogenic need, it has the potential to
deepen the understanding of the relationship
between these factors and offending. It could
also provide some context to aid in under-
standing why some justice-involved individu-
als do not desist from crime, even when they
are able to reduce their risk of reoffending by
obtaining a job.
The above example demonstrates that estab-
lishing strong connections between theory,
research, and practice can be informative for the
corrections field. Some scholars have argued
that emerging perspectives that appear to be sup-
ported by empirical evidence could be exam-
ined as a potential avenue for further enhancing
public policy and corrections (Barnes, 2014;
Beaver and Schwartz, 2016; Moffitt, 2013;
Newsome and Cullen, 2017; Rocque and
Welsh, 2015). The growth in research in evolu-
tionary psychology has revealed new findings
that could serve to expand the understanding of
offenders and rehabilitation. A brief overview
of key concepts from evolutionary psychol-
ogy and theories from this area of research is
provided in the following section, and key
findings that are relevant for policy and prac-
tice in corrections are discussed below.
EVOLUTIONARY EXPLANATIONS
OF BEHAVIOR
Over the past 20 years, various scholars have
developed theoretical explanations of behav-
ior through reliance on evolutionary princi-
ples. These theories may be informative for
corrections by enhancing our understanding
of the origins of antisocial behaviors and
 EVOLUTIONARY PSYCHOLOGY AND CORRECTIONS AND REHABILITATION
understanding differences in how individuals
respond to interventions designed to reduce
offending. These theories are founded upon
the idea that all living things, consciously or
subconsciously, strive to ensure their genes are
passed onto subsequent generations (Belsky,
1997a, 2005; Belsky and Pluess, 2009a,
2009b). In line with this goal is the under-
standing that fitness (i.e., success in mating)
and survival are fundamental to natural selec-
tion. As discussed below, scholars generally
theorize that behaviors in modern society –
including those that are widely considered to
be antisocial – could have increased fitness
and survival over the evolutionary history of
humans. These foundational components have
guided the development of evolutionary theo-
ries to explain a number of human behaviors.
Although various perspectives exist, this chap-
ter will focus on biological sensitivity to con-
text theory, differential susceptibility theory,
and intrasexual competition because these
may be the most informative for working with
correctional populations.
Biological Sensitivity to Context
and Differential Susceptibility
Theories and Behavior
Biological sensitivity to context theory and
differential susceptibility theory are compati-
ble perspectives that offer explanations for
differences in how individuals respond to the
conditions they encounter during the course of
their lives (Ellis et  al., 2011). These theories
may be particularly informative for correc-
tions, as they could provide insights into how
antisocial behaviors develop and why justice-
involved individuals respond differently to
interventions (Bakermans-Kranenburg and
van IJzendoorn, 2015). Although prior
research has suggested that adhering to the
RNR model is one of the most effective
approaches to reducing recidivism, less is
known about why some individuals who are
exposed to evidence-based practices desist
from crime and others do not.
281
Biological sensitivity to context theory,
initially proposed by Boyce and Ellis (2005),
maintains that individuals vary in their sensitiv-
ity to environmental stressors. When individu-
als perceive that events or circumstances are
threatening in some way, their stress-response
systems are activated generating increases in
heart rate and breathing, the release of various
neurotransmitters in the brain, and other phys-
iological processes. While these responses are
normal, there are differences in the situations
that prompt the activation of this system and
the intensity of the responses. The variation
in reactivity is attributed to both genetic and
environmental influences, as some may be
born with predispositions to be more or less
reactive and early experiences can shape the
precision of responsiveness (Boyce and Ellis,
2005). For example, highly reactive individu-
als may be more skilled at anticipating dan-
gers when repeatedly exposed to adversity
during their upbringing. This heightened
sensibility may promote survival by allow-
ing them to avoid life-threatening situations.
Alternatively, a highly reactive individual who
is reared in a stable, predictable, and highly
supportive environment may thrive because he
or she would gain more of the benefits of the
advantageous environment. In contrast to the
highly reactive individuals, others are theo-
rized to be born with a lower level of reactivity
(Boyce and Ellis, 2005). They are less likely to
be harmed by adversity and are seemingly
resilient to negative experiences, but also less
likely to fully acquire the benefits of highly
supportive environments.
Belsky’s (1997, 2005) differential suscep-
tibility theory rests upon many of the same
assumptions as biological sensitivity to con-
text theory but extends the logic to explain
behavioral differences in children reared by
the same parents. Briefly, the theory pro-
poses that the traits that are advantageous in
one generation will be passed on to descend-
ants. However, there are no guarantees that the
same traits that are beneficial at one point in
time will continue to be beneficial at other time
points (Belsky and Pluess, 2009a, 2009b).
282 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
In light of this uncertainty, the theory proposes
that some genes that are passed from one
generation to another can be expressed in a
range of traits to provide a natural mecha-
nism for some flexibility. This is known as
‘genetic plasticity’, a phenomenon that ulti-
mately reduces the risk that one would fail
to survive or reproduce if traits are inherited
that are no longer beneficial. As an example,
one individual may be born with a genetic
predisposition that would permit the expres-
sion of higher levels of aggression in exceed-
ingly volatile environments where aggression
may promote survival (e.g., aggression used
in self-defense, the capacity to take resources
from others during times of scarcity). The
same individual might demonstrate little to
no aggressive behaviors if reared in a safe
and secure environment where such behav-
iors may compromise survival and the like-
lihood of reproducing. The theory further
suggests that others will have lower levels
of susceptibility to environmental influence
(Belsky, 1997b, 2005). For these individu-
als, behavior will be more heavily influenced
by genetic factors regardless of the environ-
mental conditions. Those with inclinations
towards aggression, for example, will likely
exhibit aggressive behaviors regardless of
whether or not it is advantageous.
Differences in susceptibility are hypoth-
esized to be an evolutionary strategy that
maximizes the odds that at least one offspring
of a mating pair is raised to reproductive age
(Belsky, 1997a, 2005; Belsky and Pluess,
2009a, 2009b). This is because parents are
likely to adopt rearing strategies that will
encourage the adoption of traits and behav-
iors in the offspring that were beneficial for
them given the environmental conditions to
which they were exposed. This could be a
favorable strategy if the environmental con-
ditions remained consistent across time. In
those instances, children who are more easily
influenced by their environments would be
likely to adopt the strategies of the parents,
promoting continued success in surviving and
reproducing. However, it is also possible that
environmental conditions could be altered,
and the strategies encouraged by the parents
will not continue to be optimal. Should this
occur, children who are more resistant to the
parental rearing strategies may have more
favorable odds of survival and reproduction –
particularly when the rearing strategies could
compromise success in these endeavors.
Intrasexual Competition and
Antisocial Behavior
Intrasexual competition theory relies on the
social commonplaces that promote reproduc-
tive success to explain antisocial behavior. To
briefly introduce an important concept,
parental investment refers to the amount of
energy and time a parent provides to its off-
spring. Human females, like other mamma-
lian species, provide substantive parental
investment to every offspring they produce
(e.g., gestation and lactation). Human males,
however, are not required by biological pro-
cesses to invest in any one offspring and can
produce numerous children without investing
substantial resources. Consistent with the
differences in parental investment, females in
most mammalian species control access to
reproduction (i.e., they dictate which males
can reproduce). As a result, males must com-
pete for reproductive rights and engage in
competition for the ability to produce off-
spring (Buss, 1988). Importantly, females
must compete – at a much lower rate than
males – for males that provide the best
opportunity for their offspring. As hypothe-
sized by Buss and colleagues (1988, 2006),
the competition for access to reproductive
resources has led to the selection of prosocial
behaviors indicative of high paternal invest-
ment (i.e., the best opportunity for the off-
spring). Prosocial behaviors, such as altruism
and selflessness, have been suggested to
promote reproductive success among males
(Phillips et al., 2008). These prosocial behav-
iors, however, are counteracted by some
antisocial behaviors that can promote
 EVOLUTIONARY PSYCHOLOGY AND CORRECTIONS AND REHABILITATION
reproductive success with limited paternal
investment.
Buss’s (1988) initial research suggested that
effective strategies for males to promote repro-
ductive success generally involved acquir-
ing or demonstrating access to resources.
It is believed that females are more likely to
provide access to reproduction when males
can demonstrate paternal investment through
actions or physical resources (e.g., altruism,
access to food, providing safety to offspring).
Access to physical resources (e.g., food,
water, and property), especially during
ancestral times, played an important role in
maternal survival and the survival of any
offspring. Upon observing this phenom-
enon and additional evidence, Buss (1988)
concluded that some antisocial behaviors
are intended to promote reproductive suc-
cess by immediately increasing access to
physical resources. Specifically, resource-
based crimes, such as robbery and burglary,
increase access to food, water, and property,
and promote male attractiveness to female
suitors. For example, individuals during
ancestral times could steal food from oth-
ers to demonstrate their ability to feed the
mother and offspring without the risks asso-
ciated with hunting (e.g., death). In contem-
porary society, robbery and burglary can
provide individuals with immediate access
to financial resources to potentially demon-
strate the ability to financially provide for
the mother and offspring.
In addition to demonstrating paternal
investment through resource-based crimes,
Buss (1988) suggested that more aggression-
based behaviors, selected for by intersexual
competition, are intended to increase access
to reproductive resources by providing safety
and food for the mother. Furthermore, the
offspring produced by aggressive males are
more likely to survive due to their ability to
effectively engage in combat and hunt. For
example, during ancestral times males that
had increased aggressive tendencies could
protect the females and offspring from exter-
nal tribes, hunt with a high degree of success,
283
and access resources generally reserved for
tribal leaders. These same traits would then
be passed on to their offspring and increase
the reproductive success and survival of the
offspring, independent of the parental fig-
ures. These traits, as Buss (2006) suggests,
were extremely beneficial and were com-
monly selected to improve the reproductive
success and survivability of the offspring.
Violence towards females appeared to
be a reproductive strategy, selected for by
intrasexual competition, that allowed males
to take and maintain access to reproduc-
tive resources while providing limited
paternal investment (Buss, 2006). In the
current context, violence towards females
refers to extremely violent behaviors used
to increase reproduction success and con-
trol reproductive resources, while limit-
ing paternal investment. Specifically, while
scholars consistently agree that violence
towards females is immoral in contempo-
rary society, during ancestral times violent
behaviors towards females provided indi-
viduals with immediate access to reproduc-
tion and potential reductions in cuckoldry,
while requiring limited paternal investment
(Shackelford et al., 2005). For instance, acts
of violence towards females, such as sexual
aggression and domestic violence, may have
increased access to reproduction and main-
tained reproductive control, without provid-
ing resources, such as food, water, or safety.
Additionally, domestic violence may have
reduced the likelihood of cuckoldry (i.e.,
providing resources to another male’s child)
and increased the likelihood of maintaining
reproductive success and control (Camilleri
and Quinsey, 2012). Violence towards
females likely arose during ancestral times,
where individuals would engage in sexual
aggression against females to produce large
numbers of offspring without investing any
resources in the children. Historical figures,
such as Genghis Khan, serve as case studies
for the evolutionary purpose of hyper-vio-
lence against females. In the case of Genghis
Khan, it is documented that he would often
284 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
sexually assault females during conquests
and leave to continue his military campaigns.
Using hyper-sexual violence, Genghis Khan
was able to reproduce large numbers of
descendants with limited paternal invest-
ment. Genghis Khan’s hyper-sexual vio-
lence generated approximately 16 million
descendants as of 2003 (Weatherford, 2005).
Although the majority of the research
focuses on intrasexual competition among
males, females also engage in intrasexual
competition. Distinct from males, intra-
sexual competition among females concen-
trates on increasing perceived attractiveness
(i.e., males select for physical attractive-
ness; Daly and Wilson, 1988). Often, females
increase perceived attractiveness by limiting
skin blemishes, increasing body curvatures,
and increasing apparent vulnerability (Rucas
et  al., 2006). Consistent with this, females
engage in positive behaviors (e.g., purchas-
ing of accentuating clothes) or negative petty
behaviors (e.g., stealing of accentuating
clothes) to increase perceived attractiveness.
In addition to females increasing perceived
attractiveness of themselves, female intrasex-
ual competition is often characterized as psy-
chological struggle to damage the perceived
attractiveness of female rivals (Fink et  al.,
2014). Specifically, while females engage
in less physically aggressive and damaging
behaviors, strategies such as intimidation and
defamation may have improved reproductive
success. Intimidation and defamation gener-
ally result in limited physical harm but can
result in serious psychological damage to
females. Intrasexual violence among females
is rare but occurs when males provide emo-
tional or physical resources to one female
that is perceived as committed to another
female (e.g., the primary form of female
jealousy; Campbell et  al., 1998; Daly and
Wilson, 1988).
Scholars have employed the postulations
of intrasexual competition to explain criminal
behavior in contemporary society. For exam-
ple, Wilson and Daly (1985) proposed that
intrasexual competition, more specifically
risk taking, offered a potential explanation of
the age crime curve. Briefly, the age crime
curve is a common criminological phenom-
enon suggesting that most individuals engage
in criminal behavior during adolescence
and early adulthood. Wilson and Daly’s
(1985) theoretical assertions suggested that
the reproductive benefits of risk taking and
violence during adolescence – the peak age
for intrasexual competition – outweighs the
reproductive costs and increases the likeli-
hood of criminal behavior among adoles-
cents. Specifically, using data from Detroit,
Wilson and Daly (1985) demonstrated that
the vast majority of homicide offenders
and homicide victims range in age between
14 and 30 years of age, suggesting that intra-
sexual competition might account for this
trend. As individuals age, however, Wilson
and Daly (1985) argued that the reproduc-
tive costs of risk taking and violent behav-
iors become too large and encourage males
to desist from these behaviors. By the age of
30 years, individuals who engage in violent
behaviors risk injury, removal of resources,
and death, all of which are perceived as too
costly given the limited reproductive ben-
efits at that age. Furthermore, in addition to
exposing oneself to increased risks, any ini-
tial offspring are exposed to heightened risks
when males engage in intrasexual competi-
tion beyond early adulthood. This theoreti-
cal proposition was again demonstrated by
the data, which illustrated stark reductions
in homicide offenders and homicide vic-
tims as the age of an individual becomes
greater than 30 years. Like other theoreti-
cal postulations founded within intrasexual
competition, Wilson and Daly (1985) argue
that competition is an important component
of the reproductive process for most human
beings. While both differential susceptibility
and intrasexual competition hypotheses offer
broad evolutionary explanations of antiso-
cial behavior, these theoretical perspectives
can be applied to provide a greater under-
standing of behavior among correctional
populations.
 EVOLUTIONARY PSYCHOLOGY AND CORRECTIONS AND REHABILITATION
FUTURE DIRECTIONS: THE
INTEGRATION OF EVOLUTIONARY
PSYCHOLOGY INTO CORRECTIONS
Evolutionary psychology can offer various
advancements to help understand or enhance
offenders’ experiences during involvement
with the correctional system. The following
discussions will focus on the research in evo-
lutionary psychology pertaining to offending
and the benefits evolutionary psychology can
offer to understanding offenders under com-
munity supervision (i.e., probation or parole)
and incarcerated in secure facilities (i.e., jails
and prisons). To provide a brief reiteration,
community supervision and incarceration are
used by correctional agencies to supervise or
punish individuals who commit a wide vari-
ety of offenses (both non-violent and vio-
lent). Both community supervision and
incarceration possess hierarchical structures,
in which more serious offenses result in the
additional loss of freedoms. Community
supervision commonly involves offenders
reporting to a probation or parole officer at
predetermined times and engaging in a vari-
ety of other activities (e.g., alcohol treatment
or drug testing) to ensure that the offenders
do not violate the conditions of their proba-
tion. Incarceration generally involves indi-
viduals living in a secured facility for six
months or more. Incarceration conditions can
vary drastically depending upon the crimes
committed and the activities of the individual
during incarceration. Below is a review of the
research and the application of evolutionary
psychology to corrections and rehabilitation.
Differential Susceptibility and
Correctional Research
Several empirical assessments of Belsky’s
differential susceptibility hypothesis have
been published that offer explanations for the
emergence of maladaptive behaviors among
individuals (van IJzendoorn and Bakermans-
Kranenburg, 2015). For example, in a sample
285
of twins, Newsome and Sullivan (2014)
­investigated the extent to which genetic and
environmental influences could account for
variation in how adolescents respond to risk
factors for delinquency. In line with differential
susceptibility theory, the results of this study
provided evidence to suggest that individuals
respond differently to the accumulation of
risk factors, with some exhibiting resilience
(i.e., avoiding delinquency despite being
exposed to risks) and others exhibiting vulner-
ability (i.e., engaging in delinquency without
experiencing common risk factors). Differences
in youths’ outcomes were attributed to both
genetic and unique environmental factors. In a
later study, Newsome and colleagues (2016)
found that additive genetic influences
explained a significant proportion of variance
in males’ responses to risk for violent (41%)
and non-violent (29%) forms of delinquency.
Among females, however, environmental fac-
tors fully accounted for differences in the
response to risk for all forms of delinquency.
Others have used molecular genetic data
to test hypotheses about behaviors linked to
criminality based on differential suscepti-
bility theory. Simon and colleagues (2011)
examined the interaction between a cumula-
tive measure of plasticity alleles and environ-
mental conditions on aggression. The study
considered favorable aspects of the environ-
ment (e.g., supportive parenting, neighbor-
hood informal social control) and adverse
social factors (e.g., harsh parenting, dis-
crimination, violent peers). The results indi-
cated that cumulative genetic plasticity and
the qualities of the social environment pro-
duced a significant interaction that predicted
aggression, chronic anger, a hostile view
of relationships, and a belief in toughness.
A later study by Simon and his colleagues
(2012) also found cumulative genetic plastic-
ity and exposure to a hostile or demoralizing
environment was a significant predictor of
aggression and adopting a ‘street code’ that
supports criminal behavior.
Findings pertaining to criminal behaviors
and related traits and behaviors based on
286 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
differential susceptibility theory reveal that
these outcomes are the results of an interac-
tion of genetic and environmental influences.
Based on these findings, it is further possible
that there will be variation in outcomes when
the environmental influences under investi-
gation are interventions designed to reduce
deviant, delinquent, and criminal behaviors.
Indeed, researchers have begun investigating
variation in treatment effects for prevention
and intervention programs from a differen-
tial susceptibility framework and findings
have produced a substantial amount of evi-
dence in support of the theory (Bakermans-
Kranenburg and van IJzendoorn, 2015; Belsky
and van IJzendoorn, 2015; van IJzendoorn
and Bakermans-Kranenburg, 2015).
This growing body of evidence is robust,
as many of the studies employed a strong
research design through the use of a rand-
omized controlled trial. As an example, the
Strong African American Families (SAAF)
program was designed to reduce alcohol
use among African American teens, and was
tested among a sample of 337 youths using
a randomized design (Brody et  al., 2006).
Results from this study suggested that the
DRD4 gene moderated the effects of the pro-
gram on substance use among the youths.
Adolescents with a DRD4 7-repeat allele,
one hypothesized to increase plasticity, had
lower alcohol use than teens with two DRD
4-repeat alleles (Beach et al., 2010). A simi-
lar pattern of findings emerged in a test of the
Promoting School-Community-University
Partnerships to Enhance Resilience program.
In this study, participating in the program
reduced alcohol use for individuals who pos-
sessed the DRD4 7-repeat allele if they indi-
cated that involvement with their primary
caregiver was moderate or high (Cleveland
et al., 2015).
Other studies have focused on interventions
designed to reduce externalizing behaviors, a
known correlate of delinquency (White et al.,
1990). The Video-Feedback Intervention to
Promote Positive Parenting and Sensitive
Discipline was tested with a randomized trial
and was found to increase parental sensitivity
when administering discipline (Bakermans-
Kranenburg, van IJzendoorn, Pijlam et  al.,
2008). The intervention also reduced exter-
nalizing behaviors among children, but only
those who were carriers of the DRD4 7-repeat
allele. Another intervention tested in the Fast
Track Randomized Controlled Trial also gen-
erated evidence in support of the differen-
tial susceptibility perspective (Albert et  al.,
2015). Participants who had the rs10482672
A allele of the NR3C1 gene exhibited exter-
nalizing behaviors less often at age 25 years
after participating in Fast Track (18%) rela-
tive to control subjects (75%). The percent-
age of treatment and control participants who
did not possess the A allele and exhibited
externalizing behaviors were nearly identical
(56% and 57%, respectively).
Brody et al. (2015) also tested the Adults
in the Making program, which was designed
to promote positive development and reduced
drug use. The results of this study revealed
that individuals who were assigned to the
intervention and were carriers of at least one
long allele of DRD4 engaged in less drug
use compared to those who were placed in
the control group. Individuals with the same
allele, however, engaged in more drug use if
they were in the control group and lived in
a risky family environment. In other words,
those with the genotype associated with
greater plasticity seemed to experience more
benefits if they participated in the program
or more problematic behavior if they were
exposed to a high-risk context.
Findings from these and other studies have
led some scholars to suggest that differential
susceptibility theory can be informative for
intervention science (Bakermans-Kranenburg
and van IJzendoorn, 2015; Belsky and van
IJzendoorn, 2015). This body of literature
offers some explanation for why some indi-
viduals may respond better to treatment efforts
than others, and has the potential to enhance
the responsivity principle in the RNR model
– particularly specific responsivity. It is pos-
sible that, due to genetic factors, tremendous
 EVOLUTIONARY PSYCHOLOGY AND CORRECTIONS AND REHABILITATION
gains from some interventions should not
necessarily be expected. Considering this
possibility more broadly, it has further been
suggested that estimates of treatment effects
may be biased (Bakermans-Kranenburg and
van IJzendoorn, 2015). Some interventions
appear to be stronger among those with par-
ticular genotypes; however, studies often fail
to account for this phenomenon. As a result,
studies may be underestimating the treatment
effects that are possible when individuals are
better matched to interventions. To the extent
that this is occurring, efforts to take stock of
the empirical status of the RNR model may
be underestimating the actual efficacy of the
model (or its components) for individuals
who are most likely to benefit from services
offered under the model.
Integrating the differential susceptibil-
ity hypothesis into community supervision
efforts could suggest fruitful endeavors for
practitioners. Considering that the deterrent
and beneficial effects of community super-
vision differ between individuals (Andrews
and Bonta, 2010), it can be speculated that
genetic predispositions could generate dif-
ferent observed outcomes for those on com-
munity supervision. Consistent with the
RNR model, the differential susceptibility
hypothesis would advocate for a customiz-
able process for each individual under com-
munity supervision. These individualized
case management plans should be guided
by validated psychological and behavioral
assessments that identify the risks, needs,
and responsivity issues. Furthermore, case
plans should be flexible to account for the
differential responses (i.e., specific respon-
sivity) some individuals will have to meet the
requirements of the case management plans.
This customizable process could benefit both
the practitioners and the offenders, as it will
allow for the creation of more flexible and
empirically supported case plans.
In addition to individualized case man-
agement plans, the differential susceptibil-
ity hypothesis could be used to advocate
for more individualized treatment plans.
287
A well-developed component of the RNR
model is the responsivity principal, which
suggests that individual- and system-level
differences influence the observed effec-
tiveness of treatment programs. At the indi-
vidual level, it is possible that differences in
genetic predispositions influence the causal
association between treatment programs and
behavioral outcomes. To address this poten-
tial influence, probation and parole officers
can develop individualized treatment plans,
which take into account psychological and
behavioral assessments, to address the crimi-
nogenic needs of the individual.
Furthermore, genetic predispositions could
make individuals more or less vulnerable to
the effects of confinement on psychological
and behavioral outcomes. In contemporary
society, the conditions of confinement in
jails and prisons are often characterized by
the removal of basic freedoms (e.g., speak-
ing and movement), social isolation, and lim-
ited access to additional human resources.
Generally, evidence suggests that these con-
ditions can lead to increases in antisocial
outcomes (e.g., recidivism) and the severity
of psychological afflictions (Cullen et  al.,
2011). These findings, however, often coin-
cide with evidence suggesting that the effects
of prison on behavior differ by individual.
As such, it is not unreasonable to speculate
that individuals with genetic predispositions
for mental-health afflictions and antisocial
behavior might be more or less affected by
the conditions of confinement.
Integrating Intrasexual
Competition into Corrections
Unlike differential susceptibility theory,
scholars have rarely considered the effects of
intrasexual competition on offender rehabili-
tation and the RNR model. Considering the
limited postulations, it is believed that inte-
grating intrasexual competition into correc-
tions could potentially provide a superior
understanding of the observed heterogeneity
288 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
in treatment effects (e.g., Lipsey and Cullen,
2007). Specifically, intrasexual competition
theory can further emphasize the importance
of access to resources during offender reen-
try, in addition to exploring the role of age
and sexual competition during incarceration
and reentry.
Briefly, as discussed above, in contempo-
rary society females tend to subconsciously
perceive males with higher financial pros-
pects as more suitable mates (Wiederman and
Allgeier, 1992). This perceived suitability
likely arises from the benefits these males pro-
vided, regarding paternal investment, during
evolutionary times. For instance, males with
increased access to resources, such as food,
were more likely to provide these resources
to their partners and offspring. On the con-
trary, during evolutionary times individuals
with limited access to resources were less
likely to provide for their partners and off-
spring. Considering the evolutionary benefit
of resources, increased survivability, females
in contemporary society consider males with
greater resource prospects as more attractive
than males with fewer resource prospects. As
such, individuals with regular employment,
disposable income, and housing likely have
increased reproductive success. Individuals
without regular employment, disposable
income, and housing likely have decreased
reproductive success. These individuals,
however, might employ alternative strategies
to access reproductive resources. These strat-
egies could be perceived as antisocial behav-
iors by broader society.
As demonstrated by previous studies
(Richie, 2001; Travis, 2005; Visher and
Travis, 2003), involvement in the correc-
tional system drastically hinders access to
resources. For instance, individuals convicted
of felony offenses have a decreased likelihood
of legally maintaining constant employment,
purchasing a house, and sustaining wealth.
Furthermore, felony convictions drastically
hinder an individual’s legal financial pros-
pects (Raphael, 2014). Although speculative,
considering the effects of the correctional
system on resource prospects, intrasexual
competition might provide a theoretical
explanation for the observed heterogeneity in
offender reentry. If females are more attracted
to males with greater prospects, individuals
previously involved in the correctional sys-
tem might have reduced reproductive success
and in turn seek out antisocial strategies to
increase access to reproductive resources.
For example, illegal behaviors that provide
a means of financial stability, such as drug
dealing, could potentially increase access
to reproductive resources. Alternatively,
individuals previously involved in the cor-
rectional system might engage in behaviors,
such as robbery or burglary, to increase the
perception of financial stability by acquiring
jewelry or other valuable objects, which in
turn could increase reproductive success.
As such, male offenders reentering society
might recidivate due to the inability to access
financial resources through legal channels
and engage in illegal activities to counter-
act the reduced financial prospects caused
by the criminal justice system. Furthermore,
these reentry difficulties could influence the
heterogeneity of correctional programming
by encouraging individuals with reduced
resources to seek antisocial strategies for
obtaining access to reproductive resources.
Interestingly, access to financial prospects
is an important subcomponent of the RNR
model. Employment is one indicator of an
individual’s risk of recidivating and has been
identified as a criminogenic need area to tar-
get in treatment (Bonta and Andrews, 2017).
As such, intrasexual competition could
potentially provide insights as to why such
factors are of critical importance to individu-
als involved in the criminal justice system.
Additionally, considering that intrasexual
competition is heightened between the ages
of 15 and 30 (Wilson and Daly, 1985), pro-
bation and parole departments can service
these individuals differently from older cli-
entele. For instance, younger individuals
can be provided supplemental opportunities
such as additional evidence-based treatment
 EVOLUTIONARY PSYCHOLOGY AND CORRECTIONS AND REHABILITATION
opportunities to address negative behavioral
tendencies used to promote access to repro-
ductive resources. These treatment opportuni-
ties could provide individuals on supervision
with strategies that reduce risk-taking behav-
iors and help them to recognize that compe-
tition over resources could result in negative
outcomes such as imprisonment or death. All
of these strategies guided by evolutionary prin-
ciples require minor alterations to the current
services provided by community supervision
agencies and should improve the benefits for
the individuals under community supervision.
Intrasexual competition theory would
also argue that limited access to financial
prospects during incarceration might also
contribute to inmate misconduct. For exam-
ple, financial prospects, such as employ-
ment and disposable income, are extremely
difficult to access during imprisonment.
Nevertheless, while access to reproductive
resources is extremely limited during impris-
onment, competition persists because of the
high number of inmates at peak reproduc-
tive ages (15–30; Wilson and Daly, 1985).
Furthermore, intrasexual competition per-
sists as inmates attempt to attract the limited
number of females that work at or visit the
facility. This competition and competition
with males who have obtained partners out-
side of the facility likely also contributes to
the strain felt by inmates in the facility. The
inability to increase financial prospects during
incarceration might further heighten competi-
tion among inmates within these age groups.
Competition may also be further amplified
due to the limited availability of prosocial
employment after incarceration – the options
for convicted felons are exceedingly limited.
Working to expand employment opportunities
for inmates through work release programs
and other reentry planning pre-release from
prison may ease competition among inmates
and lead to reductions in misconduct.
In addition to the individual-level
enhancements offered by intrasexual com-
petition theory, a macro-level integration
of intrasexual competition could provide
289
benefits for understanding the ­heterogeneity
in ­ correctional programs. Notably, while
these postulations align with contemporary
theories of intrasexual competition, scholars
have yet to provide a comprehensive perspec-
tive or empirical examination of intrasexual
competition in correctional environments. It
is theorized that sex ratio, at the macro level,
can influence the efficacy of correctional pro-
gramming by influencing risk through the
development of environmental factors con-
ducive to heightened intrasexual competition.
For example, the prison environment drasti-
cally reduces access to reproductive resources.
As such, incarcerated populations might have
heightened levels of violence and sexual vio-
lence resulting from the extreme reductions
in access to reproductive resources. Violent
behaviors, such as assault and homicide,
might occur as a result of limited reproductive
resources within the facility and an increased
number of males at peak competition age
(Wilson and Daly, 1985). Furthermore, due
to limited reproductive resources, sexual vio-
lence against other inmates might arise as a
reproductive adaptation.
Although the link between macro-level
intrasexual competition and recidivism has
yet to be examined, various scholars have
explored the association between sex ratio
and crime rates. The research on the associa-
tion between sex ratio and crime rates could
potentially provide insight into the potential
effects of sex ratio on the reentry process. For
example, Campbell and colleagues (1998)
published a study assessing the association
between intrasexual competition and female-
on-female assault rates. The data for the study
came from the 1998 National Incident-Based
Reporting System (NIBRS) for the state of
Massachusetts. At the time, 34 districts in
Massachusetts reported crime incidents to
NIBRS. Importantly, findings from the pre-
liminary analysis suggested that the rate of
same-sex assault was higher for males and
females under the age of 24 than males and
females over the age of 24. These findings
suggested that intrasexual competition peaks
290 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
under the age of 24, evidence supporting
the theoretical position of Wilson and Daly
(1985). Additionally, males over the age of
24 engaged in higher rates of cross-sexual
violence, suggesting that older males might
use cross-sexual violence to reduce the like-
lihood of cuckolding or mate abandonment.
Further analysis, however, suggested that sex
ratio did not predict female-on-female assault
rates. As such, it can be suggested that higher
cross-sexual violence might occur among for-
mer inmates over the age of 24. Furthermore,
heightened levels of violent recidivism might
occur as an intrasexual competition strategy.
While additional research on the association
between intrasexual competition and delin-
quency exists, the literature is extremely
limited regarding correctional populations
and criminal justice processes. As such, more
research is needed to explore how intersexual
competition moderates the efficacy of reha-
bilitation programs. Although more research
is needed, intrasexual competition, like dif-
ferential susceptibility theory, is useful for
informing policy and practices in correc-
tional settings.
Implications of Evolutionary
Psychology for Corrections
Although the empirical literature in correc-
tions has predominantly drawn from psycho-
logical and sociological theories, there is
emerging evidence to suggest that evolution-
ary psychology can offer another avenue for
extending scientific inquiry in corrections.
Scholars testing differential susceptibility
theory among youths have found strong evi-
dence to suggest that children and adoles-
cents respond differently to interventions,
and that the efficacy of treatment may be
underestimated when researchers fail to con-
sider the role of genetic and environmental
factors (Bakermans-Kranenburg and van
IJzendoorn, 2015; Belsky and van IJzendoorn,
2015). These findings present two important
implications for correctional rehabilitation.
First, it is possible that genetic factors should
be considered as specific responsivity fac-
tors. Studies investigating this possibility
among adult offenders are scare relative
to those for juveniles, but as this body of lit-
erature continues to grow it could reveal
important insights to guide correctional reha-
bilitation – particularly in understanding how
and why individuals respond differently to
interventions. Second, scholars who are
examining the efficacy of correctional inter-
ventions risk generating biased results when
genetic factors are not included in statistical
models. As science and technology in this
area continue to develop, new evidence and
methods should be given consideration in
designing tests of interventions.
Evolutionary psychology may also aid in
understanding and managing inmate popu-
lations. As described above, research on
differential susceptibility theory has shown
that individuals respond differently to risks
for delinquency and crime and interventions
designed to target these behaviors. It is also
conceivable that individuals will respond
differently to prison environments in ways
that align with biological sensitivity to con-
text and differential susceptibility theories,
though this remains to be tested empirically.
To enhance research in this area, upon arrival
at a secure correctional facility the inmates
could be administered a broader range of
assessments (Newsome and Cullen, 2017).
Assessments for psychological afflictions,
such as depression, anxiety, and psychoti-
cism, and behavioral assessments for antiso-
cial tendencies and risk of recidivism could
be employed by correctional departments
to gather more personalized information on
the incarcerated individuals. There is also
evidence to suggest that the biological pro-
cesses that are triggered by stress, such as
changes in cortisol levels, can also be altered
by participating in cognitive behavioral ther-
apy (Bakermans-Kranenburg et al., 2008;
Cornet et al., 2014; van Goozen et al., 2007 ).
A more comprehensive assessment process
that includes these additional factors would
 EVOLUTIONARY PSYCHOLOGY AND CORRECTIONS AND REHABILITATION
allow practitioners to identify the individu-
alized risks and needs of the inmates while
also creating opportunities to identify those
who exhibited a heightened susceptibility
to environmental stressors and examine the
impact of imprisonment and treatment on
individuals.
The management of offender populations
could also be enhanced by further research
and consideration for the intrasexual compe-
tition perspective of behavior. For example,
strategies that increase prosocial means of
obtaining financial resources could poten-
tially be beneficial. Specifically, one reentry
strategy that could reduce recidivism is pro-
viding former felons with prosocial means
of obtaining and maintaining financial pros-
pects. Explicitly, these prosocial means can
include but are not limited to employment
opportunities, housing opportunities, and
disposable income through legal channels.
Access to these resources through proso-
cial channels could reduce the reliance on
resource-based crimes and violent behaviors
to promote access to reproductive resources.
Furthermore, promoting financial prospects
during reentry could enhance the efficacy of
the correctional programming.
Correctional departments can make con-
certed efforts to reduce competition among
inmates. These efforts can focus on increas-
ing access to non-harm-inducing resources,
reduce competitive opportunities, and reduce
risk-taking opportunities. Additionally, cor-
rectional departments can reduce competi-
tion among inmates by reducing the number
of inmates at the peak reproductive stage in
the same vicinity. These efforts could reduce
intrasexual competition over resources dur-
ing imprisonment.
CONCLUSION
The current chapter introduced the state of
corrections in the United States and the RNR
model to provide a brief overview of current
291
correctional practices. These discussions
focused on how contemporary policies and
practices could be reexamined with consid-
eration for principles derived from evolution-
ary psychology. Nevertheless, scholars have
yet to integrate the principles of evolutionary
psychology to develop meaningful policy
initiatives in corrections. As such, the core
ideas of the evolutionary psychological per-
spective were summarized and three middle-
level theories were discussed with an
emphasis on how they might inform correc-
tions (Buss, 1995). These three theories
were: biological sensitivity to context theory,
differential susceptibility theory, and intra-
sexual competition. The policy discussions
focused on how evolutionary psychology can
provide meaningful policy recommendations
for community and institutional corrections.
At the heart of these policy discussions, cor-
rectional departments should treat offenders
as individuals and train officers to better
understand the individualistic nature of
human behavior. In conclusion, researchers
and practitioners should examine the feasi-
bility and efficacy of evolutionary psychol-
ogy policy recommendations to ensure that
the desired results of the criminal justice
system are maintained.
REFERENCES
Albert, W. D., Belsky, D. W., Crowley, D. M.,
Latendresse, S. J., Aliev, F., Riley, B., & Sun,
C., (2015). Can genetics predict response to
complex behavioral interventions? Evidence
from a genetic analysis of the Fast Track Ran-
domized Control Trial. Journal of Policy
Analysis and Management, 34, 497–518.
Alper, M., Durose, M. R., & Markman, J.
(2018). 2018 Update on Prisoner Recidivism:
A 9-Year Follow-up Period (2005–2014).
Washington, DC: US Department of Justice.
Andrews, D. A. (2006). Enhancing adherence
to risk-need-responsivity: Making quality a
matter of policy. Criminology & Public Policy,
5, 595–602.
292 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Andrews, D. A., & Bonta, J. (2010). The
psychology of criminal conduct, 5th edition.
New Providence, NJ: Matthew Bender &
Company, Inc.
Andrews, D. A., Bonta, J., & Hoge, R. D. (1990).
Classification for effective rehabilitation:
Rediscovering psychology. Criminal Justice
and Behavior, 17, 19–52.
Andrews, D. A., Bonta, J., & Wormith, S. J. (2004).
The Level of Service/Case Management
Inventory (LS/CMI). Toronto, Ontario,
Canada: Multi-Health Systems.
Andrews, D. A., Zinger, I., Hoge, R. D., Bonta,
J., Gendreau, P., & Cullen, F. T. (1990). Does
correctional treatment work? A clinically
relevant and psychologically informed meta-
analysis. Criminology, 28, 369–404.
Bakermans-Kranenburg, M. J., & van
IJzendoorn, M. H. (2015). The hidden
efficacy of interventions: Gene x environment
experiments from a differential susceptibility
perspective. Annual Review of Psychology,
66, 381–409.
Bakermans-Kranenburg, M. J., van IJzendoorn,
M. H., Mesman, J., Alink, L. R. A., & Juffer, F.
(2008). Effects of an attachment-based
intervention on daily cortisol moderated by
dopamine receptor D4: A randomized con-
trol trial on 1- to 3-year-olds screened for
externalizing behavior. Development and
Psychopathology, 20, 805–820.
Bakermans-Kranenburg, M. J., van IJzendoorn,
M. H., Pijlman, F. T. A., Mesman, J., & Juffer, F.
(2008). Experimental evidence for differential
susceptibility Dopamine D4 receptor polymor-
phism (DRD4 VNTR) moderates intervention
effects on toddlers’ externalizing behavior in a
randomized controlled trial. Developmental
Psychology, 44, 293–300.
Barnes, J. C. (2014). The impact of biosocial
criminology on public policy: Where should
we go from here? In M. DeLisi & Kevin M.
Beaver (Eds.), Criminological theory: A life-
course approach (2nd Edition, pp. 83–98).
Burlington, MA: Jones & Bartlett.
Beach, S. R. H., Brody, G. H., Philibert, R. A., &
Lei, M. (2010). Differential susceptibility to
parenting among African American youths:
Testing the DRD4 hypothesis. Journal of
Family Psychology, 25, 513–521.
Beaver, K. M., & Schwartz, J. A. (2016). The
utility of findings from biosocial research for
public policy. In T. G. Blomberg, J. M. Brancale,
K. M. Beaver, & W. D. Bales (Eds.), Advancing
criminology & criminal justice policy
(pp. 452–460). New York, NY: Routledge.
Belsky, J. (1997a). Variation in susceptibility to
environmental influence: An evolutionary
argument. Psychological Inquiry, 8, 182–186.
Belsky, J. (1997b). Theory testing, effect-size
evaluation, and differential susceptibility to
rearing influence: The case of mothering and
attachment. Child Development, 68,
598–600.
Belsky, J. (2005). Differential susceptibility to
rearing influence: An evolutionary hypothesis
and some evidence. In B. Ellis & D. Bjorklund
(Eds.), Origins of the social mind: Evolutionary
psychology and child development
(pp. 139–163). New York, NY: Guilford Press.
Belsky, J., & Pluess, M. (2009a). The nature
(and nurture?) of plasticity in early human
development. Association for Psychological
Sciences, 4, 345–351.
Belsky, J., & Pluess, M. (2009b). Beyond diath-
esis stress: Differential susceptibility to envi-
ronmental influences. Psychological Bulletin,
135, 885–908.
Belsky, J., & van IJzendoorn, M. H. (2015). What
works and for whom? Genetic moderation of
intervention efficacy. Development and
Psychopathology, 27, 1–6.
Bonta, J. & Andrews, D. A. (2017). The psychol-
ogy of criminal conduct, 6th edition. New
York, NY: Routledge.
Boyce, W. T., & Ellis, B. J. (2005). Biological
sensitivity to context: I. An evolutionary-
developmental theory of the origins and
functions of stress reactivity. Development
and Psychopathology, 17, 271–301.
Brody, G. H., Murry, V. M., Kogan, S. M., Ger-
rard, M., Gibbons, F. X., Molgaard, V., &
Wills, T. A. (2006). The Strong African Ameri-
can Families program: A cluster randomized
prevention trial of long-term effects and a
mediation model. Journal of Consulting and
Clinical Psychology, 74, 356–366.
Brody, G. H., Yu, T., & Beach, S. R. H. (2015). A
differential susceptibility analysis reveals the
‘who and how’ about adolescents’ responses
to preventive interventions: Tests of first- and
second-generation Gene X Intervention
hypotheses. Development and Psychopathol-
ogy, 27, 37–49.
 EVOLUTIONARY PSYCHOLOGY AND CORRECTIONS AND REHABILITATION
Bureau of Justice Statistics (2016). Probation
and parole in the United States, 2016.
Washington, DC: National Institute of Justice.
Buss, D. M. (1988). The evolution of human
intrasexual competition: Tactics of mate
attraction. Journal of Personality and Social
Psychology, 54, 616–628.
Buss, D. M. (1995). The evolution of desire:
Strategies of human mating. New York,
NY: Basic Books.
Buss, D. M. (2006). The murderer next door:
Why the mind is designed to kill. London,
UK: Penguin Publishers.
Campbell, A., Muncer, S., & Bibel, D. (1998).
Female-female criminal assault: An evolu-
tionary perspective. Journal of Research in
Crime and Delinquency, 35, 413–428.
Camilleri, J. A., & Quinsey, V. L. (2012). Sexual
conflict and partner rape. In T. K. Shackelford
& A. T. Goetz (Eds.), The Oxford handbook
of sexual conflict in humans (pp. 257–268).
Oxford, UK: Oxford University Press.
Caspi, A., McClay, J., Moffitt, T. E., Mill, J.,
Martin, J., Craig, I. W., & Poulton, R. (2002).
Role of genotype in the cycle of violence in
maltreated children. Science, 297, 851–854.
Cleveland, H. H., Schlomer, G. L., Vandenberg,
D. J., Feinberg, M., Greenberg, M., Spoth, R.,
& Hair, K. L. (2015). The conditioning of
intervention effects on early adolescent
alcohol use by maternal involvement and
dopamine receptor D4 (DRD4) and serotonin
transporter linked polymorphic region
(5-HTTLPR) genetic variants. Development
and Psychopathology, 27, 51–67.
Cornet, L. J. M., de Kogel, C. H., Nijman, H. L.
I., Raine, A., & van der Laan, P. H. (2014).
Neurobiological changes after intervention in
individuals with anti-social behaviour: A
literature review. Criminal Behavior and
Mental Health, 25, 10–27.
Cullen, F. T., & Jonson, C. L. (2012). Correctional
theory: Context and consequences. Thousand
Oaks, CA: Sage.
Cullen, F. T., Jonson, C. L., & Nagin, D. S.
(2011). Prisons do not reduce recidivism: The
high cost of ignoring science. The Prison
Journal, 91, 48S–65S.
Daly, M, & Wilson, M. (1988). Homicide.
Hawthorne, NY: Aldine deGruyter.
Durose, M. R., Cooper, A. D., & Snyder, H. N.
(2014). Recidivism of prisoners released in 30
293
states in 2005: Patterns from 2005 to 2010.
Washington, DC: Bureau of Justice Statistics,
US Department of Justice.
Ellis, B. J., Boyce, W. T., Belsky, J., Bakermans-
Kranenburg, M. J., & van IJzendoorn, M. H.
(2011). Differential susceptibility to the envi-
ronment: An evolutionary-­neurodevelopmental
theory. Development and Psychopathology,
23, 7–28.
Fink, B., Klappauf, D., Brewer, G., & Shackelford,
T. K. (2014). Female physical characteristics
and intra-sexual competition in women.
Personality and Individual Differences, 58,
138–141.
Gendreau, P. (1996). The principles of effective
intervention with offenders. In A. T. Harland
(Ed.), Choosing correctional options that
work: Defining the demand and evaluating
the supply (pp. 117–130). Thousand Oaks,
CA: Sage.
Gendreau, P., Goggin, C., & Smith, P. (2002). Is
the PCL-R really the ‘unparalleled’ measure
of offender risk? A lesson in knowledge
cumulation. Criminal Justice and Behavior,
29, 397–426.
Gendreau, P., Little, T., & Goggin, C. (1996). A
meta-analysis of the predictors of adult
offender recidivism: What works?
Criminology, 34, 575–607.
Gendreau, P., Smith, P., & French, S. A. (2006).
The theory of effective correctional
intervention: Empirical status and future
directions. In F. T. Cullen, J. P. Wright, & K. R.
Blevins (Eds.), Advances in criminological
theory: Taking stock: The status of
criminological theory (Vol. 15, pp. 419–446).
New Brunswick, NJ: Transaction.
Kaeble, D., & Cowhig, M. (2018). Correctional
populations in the United States, 2016.
Washington, DC: US Department of Justice.
Lawrence, S., Mears, D. P., Dubin, G., & Travis, J.
(2002). The practice and promise of prison
programming. Washington, DC: The Urban
Institute.
Lipsey, M. W., & Cullen, F. T. (2007). The
effectiveness of correctional rehabilitation: A
review of systematic reviews. Annual Review
of Law and Social Science, 3, 297–320.
Lipsey, M. W., Landenberger, N. A., & Wilson, S. J.
(2007). The effects of cognitive-behavioral
programs for criminal offenders. Campbell
Systematic Reviews, 6, 1–27.
294 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Lipsey, M. W., & Wilson, D. B. (1998). Effective
intervention for serious juvenile offenders: A
synthesis of research. In R. Loeber & D. P.
Farrington (Eds.), Serious & violent juvenile
offenders: Risk factors and successful
interventions (pp. 313–366). Thousand
Oaks, CA: Sage.
Lynch, J. P., & Sabol, W. J. (2001). Prisoner
reentry in perspective. Washington, DC:
The Urban Institute.
Mears, D. P., & Cochran, J. C. (2015). Prisoner
reentry in the era of mass incarceration.
Thousand Oaks, CA: Sage.
Moffitt, T. E. (2013). Childhood exposure to
violence and lifelong health: Clinical
intervention science and stress-biology
research join forces. Development and
Psychopathology, 25, 1619–1634.
Newsome, J., & Cullen, F. T. (2017). The risk-
need-responsivity model revisited: Using
biosocial criminology to enhance offender
rehabilitation. Criminal Justice and Behavior,
44, 1030–1049.
Newsome, J., & Sullivan, C. J. (2014). Resilience
and vulnerability in adolescents: Genetic
influences on differential response to risk for
delinquency. Journal of Youth and
Adolescence, 43, 1080–1095.
Newsome, J., Vaske, J. C., Gehring, K. S., &
Boisvert, D. L. (2016). Sex differences in
sources of resilience and vulnerability to risk
for delinquency. Journal of Youth and
Adolescence, 45, 730–745.
Ogloff, J. R., & Davis, M. R. (2004). Advances in
offender assessment and rehabilitation:
Contributions of the risk-needs-responsivity
approach. Psychology, Crime & Law, 10,
229–242.
Phelps, M. S. (2011). Rehabilitation in the
punitive era: The gap between rhetoric and
reality in U.S. prison programs. Law & Society
Review, 45, 33–68.
Phillips, T., Barnard, C., Ferguson, E., & Reader,
T. (2008). Do humans prefer altruistic mates?
Testing a link between sexual selection and
altruism towards non-relatives. British Journal
of Psychology, 99, 555–572.
Polaschek, D. L. (2012). An appraisal of the
risk-need-responsivity (RNR) model of
offender rehabilitation and its application in
correctional treatment. Legal and
Criminological Psychology, 17, 1–17.
Pratt, T. C. (2019). Addicted to incarceration,
2nd edition. Thousand Oaks, CA: Sage.
Raphael, S. (2014). The effects of conviction and
incarceration on future employment out-
comes. In D. P. Farrington & J. Murray (Eds.),
Labeling theory: Empirical tests, advances in
criminological theory 18 (pp. 237–262). New
Brunswick, NJ: Transaction.
Richie, B. (2001). Challenges incarcerated
women face as they return to their communi-
ties: Findings from life history interviews.
Crime and Delinquency, 47, 368–389.
Rocque, M., & Welsh, B. C. (2015). Offender
rehabilitation from a maturation/biosocial
perspective. In M. DeLisi & M. Vaughn (Eds.),
Handbook of biosocial criminology (pp. 501–
515). Abingdon, UK: Routledge.
Rucas, S. L., Gurven, M., Kaplan, H., Winking,
J., Gangestad, S., & Crespo, M. (2006).
Female intrasexual competition and
reputational effects on attractiveness among
the Tsimane of Bolivia. Evolution and Human
Behavior, 27, 40–52.
Sampson, R. J., & Laub, J. H. (1993). Crime in
the making: Pathways and turning points
through life. Cambridge, MA: Harvard
University Press.
Shackelford, T. K., Goetz, A. T., Buss, D. M.,
Euler, H. A., & Hoier, S. (2005). When we
hurt the ones we love: Predicting violence
against women from men’s mate retention.
Personal Relationships, 12, 447–463.
Simons, R. L., Lei, M. K., Beach, S. R., Brody,
G. H., Philibert, R. A., & Gibbons, F. X.
(2011). Social environment, genes, and
aggression: Evidence supporting a differential
susceptibility perspective. American
Sociological Review, 76, 883–912.
Simons, R. L., Lei, M. K., Stewart, E. A., Beach,
S. R., Brody, G. H., Philibert, R. A., &
Gibbons, F. X. (2012). Social adversity,
genetic variation, street code, and aggression:
A genetically informed model of violent
behavior. Youth Violence and Juvenile Justice,
10, 3–24.
Smith, P., Cullen, F. T., & Latesssa, E. J. (2009a).
Can 14,737 women be wrong? A meta-
analysis of the LSI-R and recidivism for
female offenders. Criminology and Public
Policy, 8, 183–208.
Smith, P., Gendreau, P., & Swartz, K. (2009b).
Validating the principles of effective
 EVOLUTIONARY PSYCHOLOGY AND CORRECTIONS AND REHABILITATION
intervention: A systematic review of the
contributions of meta-analysis in the field of
corrections. Victims and Offenders, 4,
148–169.
Taxman, F. S., & Friedmann, P. D. (2009). Fidelity
and adherence at the transition point:
Theoretically driven experiments. Journal of
Experimental Criminology, 5, 219–226.
Travis, J. (2005). But they all come back: Facing
the challenges of prisoner reentry.
Washington, DC: The Urban Institute Press.
van Goozen, S. H. M., Fairchild, G., Snoek, H., &
Harold, G. T. (2007). The evidence for a neuro-
biological model of childhood antisocial behav-
ior. Psychological Bulletin, 133, 149–182.
van IJzendoorn, M. H., & Bakermans-Kranen-
burg, M. J. (2015). Genetic differential sus-
ceptibility on trial: Meta-analytic support from
randomized controlled experiments. Develop-
ment and Psychopathology, 27, 151–162.
295
Visher, C. A., & Travis, J. (2003). Transitions
from prison to community: Understanding
individual pathways. Annual Review of
Sociology, 29, 89–113.
Weatherford, J. (2005). Genghis Khan and the
making of the modern world. New York, NY:
Broadway Books.
White, J. L., Moffitt, T. E., Earls, F., Robins, L., &
Silva, P. A. (1990). How early can we tell?
Predictors of childhood conduct disorder and
adolescent delinquency. Criminology, 28,
507–535.
Wiederman, M. W., & Allgeier, E. R. (1992).
Gender differences in mate selection criteria:
Sociobiological or socioeconomic explanation?
Ethology and Sociobiology, 13, 115–124.
Wilson, M., & Daly, M. (1985). Competitiveness,
risk taking, and violence: The young male
syndrome. Ethology and Sociobiology, 6,
59–73.

Evolutionary Psychology and
Organized Crime
INTRODUCTION
In his foreword to the United Nations
Convention against Transnational Organized
Crime, the then Secretary-General of the
United Nations, Kofi Annan, painted a vivid
picture of what he termed the ‘gulf that exists
between the civil and the uncivil’ (United
Nations Office of Drugs and Crime, 2004: 3).
Civil society, he claimed, referred to the history
of accumulated progress that has resulted in
liberal societies that promote tolerance, respect,
and good governance. However, there also
exists what he called an ‘uncivil society’: ‘They
are terrorists, criminals, drug dealers, traffick-
ers in people and others who undo the good
works of civil society… they thrive in countries
with weak institutions, and they show no scru-
ple about resorting to intimidation or violence’
(United Nations Office of Drugs and Crime,
2004: 3). In short, the forces of organized crime
are pitted against those mainstream institutions
they seek to undermine. Efforts to better under-
stand the nature and scope of organized crime,
therefore, can contribute to strategies that can
15
Russil Durrant
mitigate its deleterious effects on both specific
victims and society more generally. Despite a
substantial body of research on the topic,
organized crime remains a contested concept
within criminology with little or no consensus
concerning what constitutes organized criminal
activities (Paoli and Vander Beken, 2014), or
what the main causes of organized crime are
(Paoli, 2016). My primary aim in this chapter is
to discuss how an approach that draws from
evolutionary psychology can contribute to a
better understanding of what organized crime
is, the motivations of organized criminal
offenders, and the structure of organized crimi-
nal groups.
Although evolutionary psychologists have
had a long-standing interest in criminal offend-
ing, especially violent crime (e.g., Daly and
Wilson, 1988; Duntley and Shackelford, 2008),
and there is an emerging literature more gen-
erally on ‘evolutionary criminology’ (e.g.,
Durrant and Ward, 2015; Kavish et  al., 2017;
Roach and Pease, 2013; Walsh and Jorgensen,
2018), the topic of organized crime has received
negligible attention from evolutionary-minded
 EVOLUTIONARY PSYCHOLOGY AND ORGANIZED CRIME 297

scholars (although see Hirschfeld, 2015, for
an approach that draws from behavioural ecol-
ogy). From the perspective of evolutionary psy-
chology, I argue that organized crime involves
the coalitional exploitation of others for per-
sonal gain in ways that violate criminal laws.
As such, the activities that comprise organized
crime – the trafficking of illegal goods and ser-
vices, predatory crimes such as theft, fraud, and
robbery, and ‘illegal governance’ crimes such as
protection payments – involve the cooperation
of individuals in ways that can enhance their
inclusive fitness at the expense of others. After
briefly reviewing contemporary criminological
approaches to conceptualizing organized crime,
I first examine the nature of organized criminal
activities and discuss how these tend to focus
on exploiting evolved human motivations and
their manifestations in contemporary envi-
ronments to promote status among organized
criminal offenders. I then outline how an evolu-
tionary perspective can shed light on the nature
of organized crime groups. Specifically, I sug-
gest that cooperation among group members
(in contexts where there can be strong tempta-
tions to ‘defect’) can be maintained via three
main evolutionary routes: kin selection, direct
and indirect reciprocity, and cultural selection
for ‘strong reciprocity’ involving the develop-
ment of group norms and the administration
of sanctions for their violation. Taking a more
historical perspective, I consider how an evolu-
tionary approach can help us to understand the
relationship between organized criminal groups
and the formation of states and under what con-
ditions we might expect such groups to emerge.
In the final section, I discuss the broader impli-
cations of an evolutionary approach for efforts
to prevent or reduce organized crime.
THE NATURE OF ORGANIZED CRIME
What Is Organized Crime?
The concept of organized crime, like many
other such concepts in the social and behav-
ioural sciences, is intensely contested, with
different authors and agencies employing dif-
ferent, but overlapping, criteria (Table 15.1;
Paoli and Vander Beken, 2014). Levi (1998:
335) notes that the term ‘organized crime’ ‘…
is generally applied to describe a group of

Table 15.1  Approaches to defining organized crime

United Nations Office of Council of the European Union2 Abadinsky3
Drugs and Crime1

1. A structured group of Mandatory (all) 1. Is devoid of political goals

three or more persons
2. The group exists for a
period of time
3. It acts in concert with
the aim of committing
at least one serious
crime
4. To obtain, directly or
indirectly, a financial or
other material benefit
1. Collaboration of more than two people 2. Is hierarchical
2. For a prolonged or indefinite period of time 3. Has a limited or exclusive
3. Suspected of the commission of serious criminal offences membership
4. Motivated for the pursuit of profit and/or power 4. Constitutes a unique
Non-mandatory (at least 2) sub-culture
1. Each with their own appointed tasks 5. Perpetuates itself
2. Using some form of discipline and control 6. Exhibits a willingness to
3. Operating on an international level use illegal violence
4. Using violence or other means suitable for intimidation 7. Is monopolistic
5. Using commercial or business-like structures 8. Is governed by explicit
6. Engaged in money laundering rules and regulations
7. Exerting influence on politics, the media, public
administration, judicial authorities or the economy

Sources:
1
United Nations Office of Drugs and Crime, 2004, Article 2(a). Downloaded from: www.unodc.org/documents/treaties/UNTOC/
Publications/TOC%20Convention/TOCebook-e.pdf
2
Cited in Paoli and Vander Beken, 2014: 22.
3
Abadinsky, 2017: 2.
298 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
people who act together on a long-term basis
to commit crimes for gain’, and this appears
to capture the core characteristics in many
definitions, as illustrated in Table 15.1.
Nonetheless, as von Lampe (2016a: 30)
admits, ‘… there is no definition that could
meaningfully delineate the study of organ-
ized crime as a field of research’. Paoli
(2016) notes that attempts to define organ-
ized crime have tended to take one of two
main paths: either they focus on the nature of
the criminal activities that are perpetrated
(e.g., drug trafficking, extortion, money laun-
dering), or they concentrate on the nature of
the groups themselves and how they are
structured. Similarly, von Lampe (2016a)
argues that there are three main focal points
for definitions of organized crime that can
serve to structure research and theory: (1) the
criminal activities that organized crime
groups engage in; (2) the nature and structure
of organized crime groups; and (3) illegal
governance and the nature of the power that
organized crime groups can sometimes wield
(Figure 15.1).
Organized crime groups engage in a diverse
range of illegal activities. These can be use-
fully classified in terms of three main types
of activities: market-based crimes, predatory
crimes, and governance crimes (Figure 15.1;
von Lampe, 2016a). Market-based crimes
involve the provision of illegal goods and ser-
vices to ‘consumers’. Widely studied exam-
ples include the trafficking in illicit drugs,
Figure 15.1  The basic dimensions of organized crime
Source: Based on the analysis in von Lampe (2016a).
arms, wildlife, and counterfeit products
(e.g., Feinstein and Holden, 2014; Reuter,
2014), people smuggling, human trafficking,
prostitution, and sex trafficking (Kleemans
and Smit, 2014), illegal gambling (Spapens,
2014), loan sharking, and money laundering
(Levi, 2014). In many cases, market-based
crimes involve transactions with ‘willing’
consumers, although this is not always the
case (e.g., human trafficking and sex traffick-
ing). Market-based crimes, as I elaborate in
more detail below, involve the provision of
goods and services that satisfy motivational
needs of consumers, but which are deemed
by governments to pose threats to the effec-
tive social functioning of society (hence their
illegality). The nature and scope of such
activities thus vary as a function of national
and international regulations and organized
crime groups may move into or out of given
markets depending on changes to their struc-
ture. For example, alcohol prohibition in the
United States opened a vast market for ille-
gal alcohol that organized crime groups – in
particular the Italian-American Mafia – were
uniquely poised to exploit (Albanese, 2014).
Predatory crimes, as their name suggests,
involve clear victims: individuals exploited in
the service of criminal gain. Predatory crimes
involve a diverse range of activities includ-
ing the running of paedophile rings, organ-
ized fraud, property offending, robbery, and
the stock-in-trade of many of the most well-
known organized crime groups: extortion
 EVOLUTIONARY PSYCHOLOGY AND ORGANIZED CRIME
(Varese, 2014). Finally, von Lampe (2016a:
77) argues that many organized crime groups
engage in what he refers to as ‘illegal gov-
ernance crimes’: ‘… activities that are inher-
ently linked to the exercise of power within
criminal organizations, criminal milieus and
beyond’. Examples include the enforcement
of particular rules or procedures through
the threat or use of force, the negotiation
of agreements or ‘contracts’ among rival
criminal groups, and the collection of ‘taxes’
(i.e. protection payments) from either legal or
illegal business operations (von Lampe, 2016a:
78). Clearly, there is overlap between many
of the activities that criminal groups engage
in and many such groups will be involved
in a diverse range of criminal activities that
might include trafficking, organized fraud,
and the collection of protection payments to
facilitate business monopolies. The Japanese
Yakuza, for example, are heavily involved
in the trafficking of illegal drugs (particu-
larly amphetamines), facilitate sex traffick-
ing and illegal gambling, offer ‘protection’
services for both legitimate and illegitimate
business operations, and play an active role in
dispute resolution for legal businesses (e.g.,
debt collection and bankruptcy management)
(Abadinsky, 2017; Hill, 2014).
The structure of organized crime groups
varies substantially from group to group
(and within groups), depending in part on the
main kinds of illegal activities that the group
is involved in. Von Lampe (2016a, 2016b)
argues that there are three basic types of
criminal structure: entrepreneurial structures,
associational criminal structures, and quasi-
government criminal structures (Figure 15.1).
Entrepreneurial structures are organized in
ways that facilitate economic activities that
provide material benefits to organized groups
and their members. Transnational drug-
trafficking networks, for example, are often
structured in ways that involve specialized
divisions of labour (from growers/manufac-
turers to traffickers to street-level dealers)
that serve the ultimate goal of delivering
the illicit drugs to consumers at an – often
299
substantial – profit. In contrast, associational
criminal structures provide social functions
through the bonding of group members, the
provision of mutual support, and the rein-
forcement of group norms and rules. Finally,
structures that serve to maintain social order
within organized crime groups (and, often,
the wider community) through the resolu-
tion of conflicts and the imposition of sanc-
tions are referred to as quasi-government in
structure, to highlight the similar functions
that states provide their citizens. One endur-
ing challenge for organized crime groups is
the maintenance of cooperation among group
members in contexts where ‘defection’ can
lead to criminal arrest. As argued below, an
evolutionary perspective can help to explain
the particular kinds of structures that tend to
emerge among organized crime groups and
which help to promote cooperation among
group members.
Finally, the third main focal point for con-
ceptualizing organized crime focuses on the
nature of illegal governance. As many schol-
ars have noted, organized crime groups often
operate in ways that mimic the functions of
legitimate states: they collect ‘taxes’ from
businesses, provide third-party enforcement
of ‘laws’, offer dispute-resolution mecha-
nisms, and in many cases provide public
goods to local communities. For instance,
the head of the Medellin drug cartel, Pablo
Escobar, donated substantial sums of money
to the community for public works pro-
grammes, including the construction of a
football stadium (Thoumi, 2014; von Lampe,
2016a). Illegal governance activities reflect
the power that organized crime groups can
yield, often through complex relationships
with legitimate businesses, political elites,
and individuals who are involved in the ‘legit-
imate’ running of state functions (e.g., police,
politicians, prosecutors, and civil servants).
Although a substantial body of research
has accumulated on the topic of organized
crime and some advances have been made in
how best to conceptualize the activities and
structures of organized crime groups, most
300 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
research has been descriptive. As such, as
Paoli (2016: 3) notes, ‘no systematic atten-
tion has so far been given to the causes
of organized crime’. Most scholars have
drawn from mainstream criminological per-
spectives in attempts to explain organized
criminal offending (e.g., Abadinsky, 2017,
chapter 2). The handful of specific theoreti-
cal approaches – such as ‘ethnic succession
theory’ and ‘alien conspiracy theory’ tend to
focus on the ethnic status of organized crime
groups in society and thus have limited scope
for understanding the diversity of organized
crime groups and organized criminal activi-
ties both historically and cross-culturally.
Other approaches tend to focus on how
organized crime groups are similar in their
functions to legal businesses or to legitimate
political organizations (see Kleemans, 2014
for a review of these ideas). Ultimately, the
challenge for theoretical approaches is to
explain (a) the motivations for engaging
in organized crime; (b) the scope of organ-
ized criminal activities; (c) the structures of
organized crime groups; (d) how such groups
maintain cooperation among group members;
and (d) how such groups persist over time
and space. An evolutionary psychological
perspective can contribute to a better under-
standing of these key issues, beginning with
a novel way of conceptualizing the primary
nature of organized crime.
AN EVOLUTIONARY PERSPECTIVE
ON ORGANIZED CRIME
An immediate question for an approach to
organized crime that draws from evolution-
ary psychology is its novelty in historical
terms: was organized crime, or anything like
organized crime, a recurrent feature of ances-
tral environments in hominin evolution? The
paradigmatic examples of organized crime
that are the feature of research in the modern
world – hierarchically structured large-scale
organizations like the Mafia or the Yakuza
– were almost certainly absent from the
largely egalitarian hunter-gatherer societies
that persisted throughout the Pleistocene.
Organized crime, as I argue below, emerges
most prominently in contexts where there is
a concentration of difficult-to-defend
resources and competition among groups for
access to those resources. Until the advent of
agriculture, the existence of such resources
would have been limited.
Evolutionary psychologists are well versed
in the idea that there is often a ‘mismatch’
between evolved psychological mechanisms
and contemporary environments (Confer
et  al., 2010). Given that organized crime,
as it is manifest in the modern world, is an
evolutionarily novel phenomenon facilitated
by the development of large-scale societies,
it is reasonable to consider its potential evo-
lutionary roots: what features of past envi-
ronments might have favoured motivations
to form coalitions among group members
for the exploitation of others? Human-male
coalitional aggression is arguably a feature
of our species that has deep roots in hominin
evolution. Like our closest genetic relative,
the chimpanzee (Pan troglodytes), human
males form relatively fluid coalitions in a
range of contexts including collective hunt-
ing, disputes over status, and inter-group
conflict including the sporadic raiding of
other groups (Wrangham and Glowacki,
2012). In addition, humans employ col-
lective aggression in the punishment of
norm violators (Boehm, 2012). Although in
humans both males and females form part
of parochial groups which favour in-group
over out-group members, males tend to be
overwhelmingly represented in coalitions
that involve significant risk to protagonists
(e.g., hunting, warfare).
According to the ‘male warrior hypoth-
esis’ (McDonald et al., 2012; van Vugt et al.,
2007) ‘humans, particularly men, may pos-
sess psychological mechanisms enabling
them to form coalitions capable of planning,
initiating and executing acts of aggression
on members of out-groups (with the ultimate
 EVOLUTIONARY PSYCHOLOGY AND ORGANIZED CRIME
goal of acquiring or protecting reproductive
resources)’ (McDonald et  al., 2012: 671).
The male warrior hypothesis suggests that
men, relative to women, may be particu-
larly prone to ethnocentrism, and are more
strongly motivated to bond with in-group
members. I argue, then, that organized crime,
from an evolutionary perspective, can be
conceptualized as a (predominantly) male
coalitional strategy to exploit resources from
others in the service of reproductive goals. Of
course, the form that organized crime takes
is highly variable and reflects, as I elaborate
in more detail below, the outcome of cultural
evolutionary processes. However, at its core,
it reflects psychological adaptations that have
evolved in the context of male coalitional
aggression.
We should expect, then, if organized crime
is largely a male coalitional strategy for
exploiting resources from others, that most
perpetrators of organized crime are male,
and that organized crime groups have fea-
tures that specifically promote male bonding.
Clear information about the demographic
characteristics of organized crime offend-
ers is difficult to come by, but the evidence
we do have suggests that most offenders are
male. For example, Kirby et  al. (2016), in
an analysis of 2.1 million recorded offend-
ers drawn from the UK Police National
Computer database between 2007 and 2010,
found that of the 4,109 offenders who met the
criteria for organized crime, 95% were male
(compared to 78% of general offenders). The
literature on gender differences in offend-
ing highlights the fact that males are more
likely to perpetrate offences than females,
and that the asymmetry between males and
females is greatest for sexual and serious
violent offending (e.g., homicide) (Durrant,
2019). It is noteworthy, then, that the gender
bias for organized crime is similar to that for
homicide, even though in this sample the
majority of organized crime offenders did
not engage in violent crime. It is worth rec-
ognizing, however, that although organized
crime is largely a male phenomenon there are
301
both all-female and mixed-gender organized
crime groups and women may be involved in
organized crime in a range of different roles
and contexts (see Pizzini-Gambetta, 2014).
In addition to the preponderance of male
organized crime offenders, many organized
crime groups – such as the Mafia in both Italy
and the United States – are exclusively male
and specifically exclude women from mem-
bership (Paoli, 2003). Finally, research sug-
gests that in addition to greater co-offending
among males relative to females, when males
do offend with others they are more likely to
employ violence and seriously injure victims
(Bourgeois and Fisher, 2018; Lantz, 2019).
Although organized crime often involves
the use of, or the threat of the use of, violence,
it involves a diversity of different activities –
from racketeering, to the trafficking of ille-
gal substances – that fundamentally involve
exploiting others for gain. This is most obvi-
ously the case when organized crime groups
extort payment from individuals or busi-
nesses in exchange for protection (often from
the organized crime group itself) because, in
effect, this is the appropriation of resources
backed up by the threat of violence. The traf-
ficking of people, organs, and wildlife also
are clearly exploitative and involve genuine
harm to victims. However, many organized
groups, including the Mafia, offer ‘genuine’
protection services to willing individuals or
groups (Paoli, 2016), and may be involved in
bringing order to otherwise lawless commu-
nities. In addition, trafficking in many ille-
gal substances such as illicit drugs involves
the provision of goods to willing custom-
ers (Kassab and Rosen, 2019; Paoli, 2016).
Nonetheless, as a general rule, organized
crime groups and their members dispropor-
tionally benefit from the goods and services
that they provide relative to their custom-
ers, and all organized crime, by definition,
‘exploits’ society by making profits without
paying taxes. Indeed, there is good economic
evidence that the presence of organized crime
groups tends to result in a suppression of eco-
nomic growth in those areas where they are
302 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
more active (van Dijk, 2007), suggesting that
such groups are not operating in the interests
of the wider community.
Organized criminal activities, then,
involve both the perpetrators of organized
crime and a somewhat heterogenous group
of individuals who can be considered ‘con-
sumers’ (some more willingly than others) of
the goods and services that organized crime
groups provide. To understand why organ-
ized criminal activities occur it is impor-
tant to recognize the motivational needs of
these two groups: offenders and consum-
ers. For many consumers, the motivations
are straightforward: they are able to obtain
goods and services that would otherwise be
unavailable through legitimate channels. It is
noteworthy that the kinds of goods and ser-
vices that organized crime groups provide
typically map onto evolved motivational sys-
tems (Kenrick et al., 2010). The ‘protection’
services that groups like the Mafia provide,
for instance, are readily sought out (as well as
imposed) because in environments that lack
robust and reliable third-party enforcement
of laws, humans seek to meet fundamental
needs for safety and self-protection for them-
selves and their kin. The provision of illegal
drugs, gambling, and sex (via prostitution
and sex trafficking) also, arguably, tap into
underlying motivational systems. Perhaps
most straightforwardly from an evolution-
ary perspective, human males are motivated
to seek opportunities for sex with desirable
mates and hence are willing consumers of
these services provided by some organized
crime groups. The motivation to consume
psychoactive drugs (and, perhaps, by exten-
sion other activities like gambling) reflect the
evolution of rewards systems that drugs act
on, even if they were not specifically selected
for drug use (Durrant et  al., 2009; Nesse,
1994; although some forms of drug use may
have played a role in enhancing fitness – see
Roulette et al., 2016).
Understanding the motivations of offend-
ers from an evolutionary perspective entails
considering ‘higher-order’ motivational
needs (Kenrick et  al., 2010), particularly
those that relate to affiliation, status, mate
acquisition, and mate retention. I discuss the
way that organized crime groups can meet
affiliation needs in more detail in the next
section, but it is worth noting how some
organized crime groups afford opportunities
for social bonding among group members
that may not have been available through
legitimate channels. Membership in organ-
ized crime groups can also meet funda-
mental motives underlying status. Status is
important from an evolutionary perspective
because higher status can be translated into
greater reproductive success (van Vugt and
Tybur, 2015; von Rueden and Jaeggi, 2016;
von Rueden et  al., 2011). Organized crime
groups provide various opportunities for sta-
tus enhancement. Simple membership in an
organized crime group can reap substantial
rewards in status – at least in the criminal
underworld. For example, becoming a for-
mal member of groups like the post-Soviet
‘Thieves in Law’ or the Hell’s Angels motor-
cycle gang, is an arduous task involving
a lengthy screening process with success-
ful applicants significantly enhancing their
social status among their peers (von Lampe,
2016b). Many organized crime groups are
also structured in terms of hierarchies that
afford opportunities for advancement with
commensurate benefits in terms of status and
access to resources. New York Mafia fami-
lies, for instance, are made up of ‘members’,
or ‘made guys’, who form ‘crews’ headed by
a caporegime or ‘captain’. Captains, in turn,
defer to the all-powerful ‘boss’ who wields
immense influence (Abadinsky, 2016). As
Abadinsky (2016: 53) explains: ‘The boss
demands absolute respect and total obedi-
ence … the boss is treated with a great deal
of deference. People rise when he enters
the room, and they never interrupt when
he is speaking’. There has been almost no
research on how such status is translated into
reproductive success among organized crim-
inals, although van San (2011: 281) provides
an interesting analysis of how many women
 EVOLUTIONARY PSYCHOLOGY AND ORGANIZED CRIME
‘find themselves uncontrollably drawn to
men who have made a career in crime’.
Although the motivations to engage in
organized crime are likely to have an evolu-
tionary basis, there are also individual differ-
ence factors that might help to explain why
some people are more likely to participate in
organized criminal groups than others. There
has been surprisingly little work done on
‘risk factors’ for organized crime in general,
although there is an extensive literature on
the factors that promote involvement in cer-
tain kinds of organized criminal groups such
as gangs (e.g., Lenzi et al., 2015), or activi-
ties such as human trafficking (e.g., Beeson,
2014). Perhaps unsurprisingly the risk factors
identified in this research are similar to the
risk factors for offending in general: social
deprivation, antisocial personality character-
istics, antisocial attitudes and cognitions, and
association with antisocial peers (Durrant,
2018). For many individuals, involvement in
organized criminal groups provides opportu-
nities for accruing status and resources that
might be difficult to obtain through legiti-
mate channels (e.g., see Bourgeois, 1995
on involvement in drug dealing). However,
it needs to be recognized that many offend-
ers, especially those occupying ‘higher-level’
positions in organized criminal groups, may
diverge from this more typical profile. There
appears to be very little research on these
individuals, but certain roles are likely to
favour traits that are more similar to white-
collar offenders, such as a greater capacity
for self-regulation (Ragatz et al., 2012), even
though they may have other antisocial per-
sonality characteristics (Craparo et al., 2018).
An evolutionary approach can help us
delineate the core aspects of organized crime
and explain both the motivations for engag-
ing in organized crime and the typical activi-
ties pursued by organized crime groups. It
can also help us to understand, as outlined
below, how these groups are structured. More
specifically, it can shed light on how coopera-
tion is maintained in such groups in the face
of incentives to defect.
303
THE PROBLEM OF COOPERATION
IN ORGANIZED CRIME
Organized crime involves the coordination of
at least two individuals in the commission of
illegal activities, although for many organ-
ized crime groups there are a large number of
individuals involved. Organized crime is,
thus, an inherently cooperative endeavour
and, as such, poses an important and widely
recognized problem: how is cooperation
maintained in the face of temptations to free-
ride or defect? (Tooby and Cosmides, 2010).
The structure of the widely researched
‘Prisoner’s Dilemma game’ provides an apt
illustration of this problem. In this game, as it
was originally developed, two offenders have
been arrested by the police and have agreed to
keep silent about their criminal activities to
protect themselves and their partner. If they
are successful in this (i.e. they cooperate),
they will receive a one-year prison sentence.
The police, as is their wont, offer a tangible
incentive to rat on their partner (i.e. to defect):
if they talk, they will be set free, unpunished,
and their partner will receive a lengthy
10-year prison sentence. If they incriminate
each other, they will get five years in prison.
The structure of the dilemma is obvious: the
best collective outcome for both offenders is
to stay silent because they will receive a com-
bined two years in prison. However, the best
individual outcome is to be set free, but
because this outcome is presented to each
offender, if both individuals take this option,
they will both be worse off than if they stayed
silent (Cronk and Leech, 2013). The Prisoner’s
Dilemma provides a vivid illustration of a
more general problem for evolutionary
theory: how do we explain cooperation among
group members given the selective advan-
tages of free-riding or defection?
There is now an extensive literature on the
evolution of altruism and cooperation and a
number of viable evolutionary mechanisms
have been identified that can maintain cooper-
ation among individuals (see Clutton-Brock,
304 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
2009; Cronk and Leech, 2013; Kurzban
et  al., 2015; West et  al., 2011, for reviews).
Cooperation among kin is easily explained:
because kin are more likely to share genes
compared to non-kin, any behaviour that
facilitates interactions that benefit kin or
are mutually beneficial will be selected
for (Dawkins, 1976; Hamilton, 1964).
Cooperation also occurs widely among non-
kin in humans and other animals and can be
maintained via direct reciprocity: if all parties
benefit from cooperation, then all parties will
be motivated to cooperate (Trivers, 1971).
Cooperation can be maintained via direct
reciprocity even if the exchange of benefits
is not simultaneous as long as they are recip-
rocated at some future date. Cooperation
may also evolve via what some scholars have
termed ‘indirect reciprocity’. Cooperative
behaviour can be a reliable signal to other
group members that the individual can be
trusted and thus they are likely to be cooper-
ated with in the future (Kurzban et al., 2015).
Both direct and indirect reciprocity are vul-
nerable, however, to exploitation. Individuals
who cooperate but are not cooperated with in
turn will be at a selective disadvantage relative
to those who accept the benefits of cooperation
without reciprocating in kind. One solution to
this problem is the evolution of punishment.
Individuals who receive cooperation but don’t
return it in kind may be subject to sanctions –
either at the hands of the cooperator (what we
call ‘revenge’ or ‘retaliation’ – Jackson et al.,
2019; McCullough et al., 2013) or via a third-
party (what is often termed ‘altruistic punish-
ment’ or, more generally, ‘justice’ – Boyd
et al., 2003).
Finally, a number of scholars have argued
that the evolution of cooperation in large-scale
groups, of the kind that most people live in
today, has been shaped by cultural evolution-
ary processes (Chudek et al., 2013; Henrich,
2016; Richerson et al., 2017; Turchin, 2016).
A particular emphasis has been placed on the
potential role of cultural group selection in
shaping cooperative behaviour within groups.
Groups that have characteristics – norms,
values, institutions – that tend to promote
cooperation among in-group members will
be more successful relative to groups without
these characteristics and hence these particu-
lar norms, values, and institutions will tend
to flourish at the expense of others. Recent
work suggests that cultural group selection
processes may have been important in the
cultural evolution of prosocial religions that
are maintained via constellations of norms,
values, and practices that demarcate in-group
from out-group members (e.g., rituals, hard-
to-fake markers) and promote social cohesion
and cooperation among members of the in-
group (see Norenzayan, 2013; Norenzayan
et  al., 2016; Saroglou, 2011). In particular,
cultural evolutionary approaches to the evo-
lution of religion highlight the role of specific
norms and values that bind co-religionists
(e.g., the notion of fictive kin – all adherents
are ‘brothers’ and ‘sisters’): costly, time-­
consuming, and sometimes painful rituals
that serve as clear markers of group commit-
ment (what Norenzayan, 2013, calls ‘cred-
ibility enhancing displays’), and often strict
rules and their enforcement by both group
members (Durrant and Poppelwell, 2017),
and by a god or gods (Norenzayan, 2013).
Similar processes operate to bind individu-
als into closely knit male groups (army units,
terrorist cells) in the context of inter-group
conflict (e.g., Whitehouse et  al., 2014),
consistent with the male warrior hypothesis
outlined above.
How might these various evolutionary
processes that support cooperation help us
to understand the nature, structure, and prac-
tices of (largely male-based) organized crime
groups? Table 15.2 provides an overview
of how evolutionary mechanisms for coop-
eration might map on to the different organ-
ization structures elucidated by von Lampe
(2016a, 2016b). Many organized criminal
groups are based on interactions involving
family members (von Lampe, 2016b). This
seems to be the case for both relatively small-
scale organized crime groups as well as for
larger, more complex criminal organizations.
 EVOLUTIONARY PSYCHOLOGY AND ORGANIZED CRIME 305

Table 15.2  The functional structures of organized crime groups in relation to the key
evolutionary processes that facilitate cooperation1
Entrepreneurial Associational Quasi-governmental

Kin-selection Blood family associations and

‘fictive kin’ structures, and
‘ethnic’ groups
Direct reciprocity Illegal networks, markets, Mutual support Honouring of commercial
and businesses exchanges and ‘contracts’
Indirect reciprocity Reputation-based promotion Reputation, honour, loyalty
Punishment Violence used to enforce Mechanisms for the enforcement Sanctions for violations
reciprocal exchange – of rules and norms administered by group members
‘revenge’
Cultural selection The role of group-based norms, Mechanisms for regulation, conflict
markers, rituals, and practices resolution, protection
1
Examples provided are illustrative only and may be influenced by multiple evolutionary mechanisms (e.g., revenge may
have been shaped by both genetic and cultural evolutionary processes – Jackson et al., 2019).

In the UK, for instance, there is a long history of
criminal groups based on cooperation among
brothers: the Sabini brothers in the 1920s,
the Richardson brothers in South London
in the 1940s and 1950s, and the notorious
Kray twins during the same period in the
East End of London (Antonopoulos and
Papanicolaou, 2018). The Latin American
drug cartels that emerged during the 1970s
were also often centred on leaders who were
close kin. The Ochoa clan, along with Pablo
Escobar, led the Medellin cartel, while the
Cali cartel was led by the Rodriguez-Orejuela
brothers (Abadinsky, 2017; Antonopoulos
and Papanicolaou, 2018). Italian Mafia
organizations have also historically been
based on blood ties, although this varies
somewhat among the different Mafia groups
(Antonopoulos and Papanicolaou, 2018;
von Lampe, 2016a). Research suggests that
there is often a considerable degree of inter-
generational continuity among members of
organized crime groups as sons in particular
follow their fathers’ involvement (van Dijk
et al., 2018).
Many large-scale organized crime groups
(like many religious groups) tap into evolved
mechanisms to preferentially cooperate
with kin by creating family-like structures,
based on the notion of ‘fictive kin’. Thus, the
various branches of the Italian Mafia place
emphasis on the idea of ‘brotherhood’, the
Japanese Yakuza invoke the notion of Ikka,
or family, creating an organizational struc-
ture that mimics that of the family (with
a ‘father’, and ‘brothers’), and the Hong
Kong Triads invoke a concept of ‘universal
Triad brotherhood’ which links the various
Triad societies (von Lampe, 2016a). From
an evolutionary perspective, the prevalence
of both actual and fictive blood ties reflects
the evolution of mechanisms that promote
cooperation among close kin. Thus, they are
likely to reduce defection among members
of organized crime groups while enhancing
cooperation and social cohesion. Indeed,
many large-scale organized crime groups
emphasize the importance of fictive-kin ties
(among ‘brothers’) over and above ties to
family members. The first rule of the Russian
‘Vory v zakone’ (‘Thieves in law’), for exam-
ple, states: ‘A thief must turn his back on his
family – mother, father, brothers, and sisters.
The criminal community is family’ (cited in
von Lampe, 2016a: 166).
Cooperation among organized crime
groups is also maintained via direct and
indirect reciprocity. The operation of direct
reciprocity is straightforward. In the same
manner that legal transactions are maintained
through simultaneous or delayed exchanges
of goods or cash, the illegal activities of
306 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
organized criminals are facilitated through
such reciprocal exchanges. Individuals who
do not reciprocate will be viewed as less
trustworthy and hence are less likely to be
chosen as cooperative partners in the future
(in addition to being the object of poten-
tial sanctions, including those that operate
through quasi-governmental-like processes).
The exchange of goods and services among
members of organized crime groups, how-
ever, is not always immediate or necessar-
ily reciprocal, and cooperative behaviour
may be maintained as a way of signalling
trustworthiness to other group members.
Values such as honour, trustworthiness, and
loyalty to the group are strongly emphasized
in many organized crime groups such as
the Mafia. In turn, organized crime groups
provide extensive mutual support to group
members in need: ‘The bonds that tie indi-
viduals together in an associational crimi-
nal structure almost by nature establish a
commitment to mutual support. There is an
unspoken or explicit obligation to come to
the aid of other members when needed’ (von
Lampe, 2016b: 25). As Paoli (2003: 17)
notes for the Italian Mafia, aid to other
members is given ‘with no expectation of
short-term rewards’.
Even among members of organized crime
groups there may be temptations to ‘defect’
or ‘free-ride’ – to benefit from the coopera-
tion of others while not fully reciprocating in
turn. The role of sanctions in shaping behav-
iour among offenders who violate norms is
well recognized. For example, Maier and
Ricciardelli (2019), in a qualitative study
of 56 former prisoners in Canada, found
that prisoners would often rather accept the
institutional punishment handed out by the
authorities than risk the informal punishment
they might receive at the hands of their fellow
inmates for being a ‘snitch’. As one partici-
pant recounted: ‘ …’ cause I figured out …
‘cause I found out who did it but I didn’t say
anything [to the prison staff] … because they
[prison staff] try to tell me well “why didn’t
you say who did it or you get it shit?” I’m
like “you’d rather I’d say” … [but if] I say it
now and then I’ll get killed and then what?’
(Maier and Ricciardelli, 2019: 245; see also
Dudai, 2018, for the role of punishment
among members of the Irish Republican
Army). Entrepreneurial structures based
on reciprocal exchange are vulnerable to
exploitation and defection because agree-
ments cannot be enforced via legal channels.
Thus, protagonists in organized crime must
be willing to use violence and other threats
to enforce cooperation. The high levels of
violence in some illegal markets, such as the
drug market, reflect the absence of formal
mechanisms of dispute resolution (Reuter,
2016). Many large-scale organized crime
groups, however, have clear rules and mecha-
nisms for their enforcement that entail quasi-
governmental processes of dispute resolution
and sanctioning. These processes go beyond
‘revenge’ and may include processes that
resemble the criminal justice system, includ-
ing collective decision making, dispute reso-
lution between parties, and graded sanctions
that include fines, suspension of member-
ship, and death (von Lampe, 2016a). Some
methods are specific to particular groups
and serve as markers of group commitment:
among Japanese Yakuza, for example, finger
amputation is employed as both a punish-
ment and a signal to others of commitment to
the group (Hill, 2014).
When groups become large or entail high
levels of cooperation among members, social
cohesion and cooperation are more difficult
to sustain. If, as suggested above, coopera-
tion in large-scale groups is partly maintained
via norms, values, and institutions that have
been culturally selected for, then we might
expect ‘successful’ organized crime groups
(e.g., those with greater longevity) to have
converged upon rules, norms, and processes
that maintain group cooperation. Codes of
conduct are similar across geographically
divergent organized crime groups, provid-
ing some support for this suggestion (Kassab
and Rosen, 2019; von Lampe, 2016b). They
usually coalesce around a set of strict rules or
 EVOLUTIONARY PSYCHOLOGY AND ORGANIZED CRIME
obligations: (1) absolute loyalty to the group
above all others, including kin; (2) strict
secrecy concerning the nature of the group’s
members and activities; and (3) honesty and
reciprocity among group members. In addi-
tion, organized crime groups often have a
diverse set of specific rules about how individ-
uals conduct themselves with both group and
non-group members, and the kinds of criminal
activities they can and cannot engage in. For
example, Italian Mafia groups traditionally
prohibit involvement in drug trafficking or the
use of kidnapping for ransom, to avoid draw-
ing unnecessary attention to themselves (von
Lampe, 2016b). Interestingly, various organ-
ized crime groups such as the Italian Mafia
also prohibit sexual relations with other mem-
bers’ wives – a rule that is likely to promote
and maintain cohesive bonding among male
members (Kassab and Rosen, 2019).
Cultural group selection processes are
facilitated through clear mechanisms for
symbolically delineating group bound­
aries that demarcate group membership
(Richerson et al., 2016). In addition, social
cohesion among group members, as noted
above, can be enhanced via participation
in collective rituals. The role of initiation
rituals and markers of group status are
common among organized crime groups.
Among Italian Mafia groups, traditional
initiation ceremonies involve the pricking
of the index finger so that blood drips on a
saint’s icon, which is then set on fire in the
hand of the candidate, who swears loyalty
to the Mafia and its rules (Antonopoulos
and Papanicolaou, 2018). Markers of group
membership are also common: membership
into outlaw motorcycle gangs is often sig-
nalled by the wearing of distinct patches, the
Russian vory v zakone sport specific tattoos
that signal group status communicated in a
coded language only known among group
members (Abadinsky, 2017). Among the
Yakuza, extensive body tattooing is com-
mon, and members are required to partici-
pate in ceremonies such as initiation rites
(Hill, 2014). However, given the importance
307
of secrecy among many organized crime
groups, such as the Mafia, overt markers
of group status may be absent from some
groups. From the perspective of cultural
group selection theory (Richerson et  al.,
2016), it is easy to see why different organ-
ized crime groups have converged on similar
mechanisms for maintaining cooperation:
those groups that promote extreme within-
group cooperation and that adhere to clear
rules (like codes of secrecy) are likely to
thrive at the expense of groups that do not.
There has been very little empirical
research that has systematically examined
the nature of cooperation among members of
organized crime groups (or among offenders
in general). In the first study to employ the
Prisoner’s Dilemma game with a sample of
prison inmates, Khadjavi and Lange (2013)
found that the prison sample was more likely
to cooperate in the simultaneous Prisoner’s
Dilemma (where the decision to cooperate or
defect is made at the same time) compared
to student participants. Rates of cooperation
were roughly the same in the two groups in
response to a sequential Prisoner’s Dilemma
(where individuals respond with cooperation
or defection in response to an initial response
by their partner). The results perhaps reflect
the existence of informal ‘prisoner codes’ that
structure social interactions in correctional
facilities, and facilitate cooperation among
inmates (Maier and Ricciardelli, 2019).
Interestingly, a more recent study using a
sample of community-supervised offend-
ers found lower rates of cooperation in a
Prisoner’s Dilemma game in the offending
group compared to a control group (Clark
et  al., 2015), suggesting that it may be fea-
tures of the prison environment that play a
role in structuring cooperative behaviour in
these games (although the nature of the sam-
ple could also have been a factor, given that
Khadjavi and Lange’s study involved women
prisoners only).
Although we may expect that coopera-
tion among offenders depends on specific
contextual factors (e.g., lower in general, but
308 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
perhaps higher in institutional contexts), we
should expect that organized criminals dis-
play high rates of cooperation among group
members for the reasons that have been pre-
sented above: kin and fictive-kin structures,
networks of direct and indirect reciprocity,
and strong social norms and rules that pro-
mote trust, cooperation, and bonding among
group members. This is exactly what Nese
et  al. (2018) found in an experimental field
study with a sample of Camorra (Mafia)
prisoners recruited from a jail in Naples.
Participants in this study (129 Camorra,
109 ‘ordinary’ criminals, and 109 stu-
dents) engaged in two different experimen-
tal games: a one-shot Prisoner’s Dilemma
(PD), and one-shot Prisoner’s Dilemma with
third-party punishment (PD-TPP). In the
PD game, participants were given an enve-
lope containing 10 tokens and paired with
another individual with a similar envelope
containing 10 tokens. Both individuals had
to simultaneously decide whether to (a) keep
their tokens; or (b) send them to their part-
ner, in which case the researcher would triple
the amount received (at the end of the study
tokens could be exchanged for real money).
The Prisoner’s Dilemma structure here is
clear: if both individuals cooperate and send
their tokens, they will receive 30 points (the
10 received times 3), if both defect and keep
their own tokens they obtain 10 points, and
if one cooperates while the other defects the
cooperator receives nothing while the defec-
tor obtains 40 points (their original 10 plus
the 10 they receive multiplied by three). All
games were played with anonymous part-
ners, but participants knew which group they
were playing with (e.g., Camorristi interacted
with Camorristi, and students interacted with
students).
The results indicated that the organized
criminal offenders (the Camorristi) engaged
in higher levels of cooperation (86.7% of
trials) compared to the students (67.5%)
and the ordinary criminals (50.0%), provid-
ing strong support for the idea that organ-
ized crime members abide by strong norms
of reciprocity. Interestingly, in the PD-TPP
game, when a third party could pay points to
sanction the two participants in the PD game
(by taking points from them for not cooper-
ating), rates of cooperation dropped among
both the Camorristi and the students but
increased among the ordinary prisoners. This
suggests that the introduction of sanctions
may reduce cooperation under some circum-
stances, particularly if the sanctions are per-
ceived as unfair (see Fehr and Rockenbach,
2003). In a related study employing a battery
of economic games with samples of students
residing in both pro- and anti-Mafia neigh-
bourhoods in Palermo, the researchers found
that levels of trust, trustworthiness, and altru-
ism were lower among students in the pro-
Mafia neighbourhoods, but that in-group
favouritism on these measures was higher
than in the anti-Mafia neighbourhood (Meier
et  al., 2016). This implies that the strong
presence of organized crime groups might
erode generalized trust and cooperation in
the environments in which they are active but
promote greater cooperation among in-group
members. Similar results have been found in
research that examined the role of religion
on cooperation: adherents tend to be more
prosocial and cooperative, but also more
parochial, more strongly favouring in-groups
over out-groups (e.g., Purzycki et al., 2018).
Clearly, more empirical research is needed
on the nature of cooperation among organ-
ized crime members, but these initial studies
support the idea that cooperation has been
maintained through a combination of mecha-
nisms including direct and indirect reciproc-
ity along with third-party punishment and
the maintenance of group-based norms and
values.
ORGANIZED CRIME AND STATE
BUILDING
Charles Tilly (1985/2002: 35) influentially
drew a somewhat startling comparison
 EVOLUTIONARY PSYCHOLOGY AND ORGANIZED CRIME
between organized crime groups and the
institutions of ‘legitimate’ statehood:
Apologists for particular governments and for
government in general commonly argue, precisely,
that they offer protection from local and external
violence. They claim that the prices they charge
barely cover the costs of protection. They call
people who complain about the price of protection
‘anarchists’, ‘subversives’, or both at once. But
consider the definition of a racketeer as someone
who creates a threat, then charges for its reduction.
Governments’ provision of protection, by this
standard, often qualifies as racketeering. To the
extent that the threats against which a given
government protects its citizens are imaginary, or
are a consequence of its own activities, the
government has organized a protection racket.
This comparison speaks to a number of
important similarities between the ‘legiti-
mate’ practices and institutions of the nation
state and those that are apparent in many
organized crime groups. First, and most
importantly, as the sociologist Max Weber
(1978) emphasized, states possess a monop-
oly on legitimate violence to punish rule
violations and maintain social order. To sus-
tain a state apparatus that can maintain order
and ‘protect’ its citizens from the predations
of others, it levies taxes from those who
reside within the state’s jurisdiction and,
more generally, ‘claims authority … to a very
large extent over all action taking place in the
area of its jurisdiction’ (Weber, 1978: 69).
Similarly, many organized crime groups such
as the Mafia extract payments from busi-
nesses that operate within their ‘jurisdiction’
for which, in turn, they provide protection (in
part, from themselves, but also from other
predatory individuals and groups). Of course,
these similarities do not necessarily apply to
all organized crime groups and the degree to
which they exercise ‘state-like’ functions
vary, in part due to differences to the extent
that the activities of such groups entail the
control of specific territory (Koivu, 2016).
Nonetheless, the comparison between states
and organized crime groups is a potentially
fruitful one that may serve to shed light on
both the origin of the state itself, and how
309
and under what conditions organized crime
groups tend to emerge.
Understanding the transition from small,
largely mobile hunter-gatherer bands to sed-
entary, large-scale societies with state-like
features remains a formidable task that has
engaged scholars from a wide range of disci-
plines. Although there remain many theories of
state formation (Carneiro, 1970; Turchin et al.,
2018), it seems that early states shared many
of the features of organized crime groups
such as the Mafia, as they provided protec-
tion while extracting profits and wielding sub-
stantial amounts of largely unchecked power
(Scheidel, 2017). As Scheidel (2017: 48) lacon-
ically suggests: ‘Reduced to essentials, history
has known only two ideal-typical modes of
wealth acquisition: making and taking’. Male
coalitionary elites are uniquely poised to fol-
low the latter pathway. Scott (2017) argues that
the development of high-density agriculture,
particularly cereal production, provided the
conditions for emerging statehood. Because
wealth is heavily concentrated in a clearly
defined territory, it is chronically vulnerable to
exploitation by mobile raiders.
Raiding, however, is inherently unsta-
ble as an ongoing concern: not only does it
destroy wealth that forms the basis of the
raiding activity, but as raiding increases in
frequency sedentary lifestyles become unten-
able. ‘Knowing this’, Scott (2017: 240–241)
suggests
raiders are more likely to adjust their strategy to
something that looks more like a ‘protection
racket.’ In return for a portion of the trade goods,
harvest, livestock, and other valuables, the raiders
‘protect’ the traders and communities against
other raiders and, of course, against themselves…
In extracting a sustainable surplus from sedentary
communities and fending off external attacks to
protect its base, a stable protection racket like this
is hard to distinguish from the archaic state itself.
Recent quantitative analyses of emerging
statehood suggest that large-scale complex
human societies were more likely to develop
in contexts where agriculture had been prac-
ticed for long periods, population size was
310 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
larger, and warfare was more intense (Currie
et al., 2019; Turchin et al., 2018).
As humans scaled up from small-scale
hunter-gatherer bands to large-scale socie-
ties, male coalitions – long a component of
human societies in the context of collective
‘power scavenging’, hunting, and inter-group
raiding – were uniquely placed to exploit
new opportunities made available through
the concentration of resources offered by the
emergence of agriculture (although it is worth
noting here that the development of agricul-
ture preceded the emergence of states by a
considerable period of time, suggesting that
agriculture was a precondition, rather than
an inevitable precursor, for state formation –
Gintis et  al., 2019). Hirschfeld (2015), who
has provided one of the few in-depth analyses
of organized crime from an evolutionary per-
spective, employs the idea of an ‘evolution-
ary stable strategy’ to argue that economic
exchanges based purely on reciprocity are
going to be vulnerable to exploitation from
organized crime like activities (raiding, vio-
lence, extortion) (although see Koivu, 2016).
In turn, exploitative groups are vulnerable to
others that provide protection with less overt
or obvious exploitation of primary producers,
resulting in transitions to ‘gangster-states’.
More stable state-like institutions of the kind
that operate in contemporary liberal democ-
racies may emerge but will remain vulner-
able to exploitation from organized criminal
groups under specific circumstances.
From this broader perspective, organized
crime groups with different characteristics
or traits compete both among themselves
and against ‘legitimate’ states for control and
power over valuable resources. Historical
analyses suggest that organized crime groups
are most likely to emerge when there is a val-
uable concentration of defendable resources
and when legitimate state processes for
enforcing the law are weak or non-existent.
The emergence of the Sicilian mafia in the
19th century, for example, occurred against
a backdrop of a weak rule of law in com-
bination with a concentration of valuable
resources that could be readily monopolized
– citrus fruit (Dimicio et al., 2017) and sul-
phur (Buonanno et al., 2015). Although many
organized crime groups interact with legiti-
mate state actors and institutions in various
forms of ‘symbiotic’-like relationships that
benefit each other, research clearly suggests
that the prevalence of organized crime in a
given nation state is strongly (negatively)
associated with the quality of rule of law,
indicating that it is the absence of legitimate
third-party enforcement that creates oppor-
tunities for organized crime groups to exert
influence (van Dijk, 2007).
CONCLUSION
I have argued in this chapter that organized
crime involves the coalitional exploitation of
others for personal gain in ways that violate
criminal laws. Consistent with the male war-
rior hypothesis, I have suggested that selec-
tion for male coalitional aggression in our
evolutionary past has fostered the tendency
for males to form groups in the context of
hunting, raiding, and inter-group conflict. As
such, the activities that comprise organized
crime – the trafficking of illegal goods and
services, predatory crimes such as theft,
fraud, and robbery, and ‘illegal governance’
crimes such as protection payments – involve
the cooperation of (mostly) men in ways that
can enhance their inclusive fitness at the
expense of others. Because organized crime
groups (like any cooperative group) are vul-
nerable to exploitation, their longevity
depends on exploiting mechanisms that facil-
itate cooperation among group members.
Three main evolutionary routes have been
discussed: kin selection, direct and indirect
reciprocity, and cultural group selection.
Many organized crime groups are based on a
nucleus of close family members and many
more use cultural mechanisms to enhance
cooperation via the idea of ‘fictive kin’.
Cooperation is also sustained through the
 EVOLUTIONARY PSYCHOLOGY AND ORGANIZED CRIME
reciprocal exchange of resources and via
mechanisms that encourage trustworthy
behaviour, ultimately backed up by the threat
of sanctions. Large-scale organized crime
groups often have many of the features that
promote social cohesion in different group
contexts (e.g., religious groups), including
costly initiation ceremonies, symbolic mark-
ing of group members, and rigid codes of
conduct. The capacity for organized crime
groups to exert power through the control of
concentrated resources, the extraction of
‘taxes’, and the use of violence to enforce
cooperation offers many similarities with the
functions of legitimate governments and may
provide insights into the ultimate origin of
the state in human history. Given the paucity
of research on organized crime from an evo-
lutionary perspective, there is substantial
scope for future research to explore many of
these avenues in further detail.
Finally, it is worth considering how the
approach taken in this chapter suggests vari-
ous avenues for reducing the (largely) harm-
ful impacts of organized crime on society.
Most obviously, if we can conceptualize
organized crime groups as being in compe-
tition with legitimate states, then we should
employ strategies that support those charac-
teristics of states that allow them to promote
public goods – protection, safety, an effec-
tive rule of law, social services – as they are
likely to reduce the impact and influence of
many organized crime groups that may offer
similar ‘services’. Relatedly, approaches that
de-couple organized crime groups from legit-
imate institutions through crackdowns on
corruption are likely to be effective, as such
groups can no longer exploit state actors and
institutions to pursue their own objectives
and goals. If membership in organized crimi-
nal groups is seen by many men as a viable
route to obtain social status, as argued above,
then strategies to improve opportunities
for status attainment through legitimate
means (e.g., better education, reductions in
inequality, and anything that is likely to pro-
mote the development of slow life history
311
strategies – Durrant, 2016) are likely to
reduce the lure of such groups. Finally, it is
worth considering how some highly desir-
able commodities (e.g., drugs) and services
(e.g., prostitution), currently monopolized
by organized crime groups because they are
illegal, may be better managed through legiti-
mate channels, thus eliminating many of the
harms that arise through their (illegal) sale.
It is unlikely that we will be able to eradicate
organized crime groups entirely because they
readily emerge in contexts where state power
is weak or non-existent, or corruption is wide-
spread, but recognition of their evolutionary
origins may encourage strategies that can
reduce their largely negative effects on society.
REFERENCES
Abadinsky, H. (2016). Organized crime (11th
edition). Boston, MA: Cengage Learning.
Albanese, J. S. (2014). The Italian-American
Mafia. In L. Paoli (Ed.), The Oxford handbook
of organized crime (pp. 142–158). Oxford,
UK: Oxford University Press.
Beeson, J. G. (2014). Psychology of human
trafficking. In M. J. Palmiotto (Ed.), Combat-
ting human trafficking: A multidisciplinary
approach (pp. 47–60). London, UK: CRC
Press.
Boehm, C. (2012). Moral origins: The evolution
of virtue, altruism, and shame. Philadelphia,
PA: Basic Books.
Bourgeois, C. A., & Fisher, M. L. (2018). More
‘bros,’ more woes? The prevalence of male
coalitions in crimes of robbery. Evolutionary
Behavioral Sciences, 12, 126–131. doi:
10.1037/ebs000116
Bourgeois, P. (1995). In search of respect:
Selling crack in El Bario. Cambridge, UK:
Cambridge University Press.
Boyd, R., Gintis, H., Bowles, S., & Richerson, P. J.
(2003). The evolution of altruistic punish-
ment. Proceedings of the National Academy
of Sciences, 100, 3531–3535.
Buonanno, P., Durante, R., Prarolo, G., & Vanin,
P. (2015). Poor institutions, rich mines:
Resource curse in the origins of the Sicilian
312 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
mafia. The Economic Journal, F175–F201.
doi: 10.1111/ecoj.12236
Carneiro, R. L. (1970). A theory of the origin of
the state: Traditional theories of state origins
are considered and rejected in favor of a new
ecological hypothesis. Science, 169, 733–738.
Chudek, M., Zhao, W., & Henrich, J. (2013).
Culture-gene coevolution, large-scale
cooperation, and the shaping of human
social psychology. In K. Sterelny, R. Joyce,
B. Calcott, & B. Fraser (Eds.), Cooperation
and its evolution (pp. 425–457). Cambridge,
MA: MIT Press.
Clark, B. C., Thorne, C. B., Hendricks, P. S.,
Sharp, C., Clark, S. K., & Cropsey, K. L.
(2015). Individuals in the criminal justice
system show differences in cooperative behav-
iour: Implications for cooperative games.
Criminal Behaviour and Mental Health, 25,
169–180. doi: 10.1002/cbm.1920
Clutton-Brock, T. (2009). Cooperation between
non-kin in animal societies. Nature, 462
November, 51–57. doi: 10.1038/nature08366
Confer, J. C., Easton, J. A., Fleischman, D. S.,
Goetz, C. D., Lewis, D. M. G., Perilloux, C., &
Buss, D. M. (2010). Evolutionary psychology:
Controversies, questions, prospects, and
limitations. American Psychologist, 65, 110–
126. doi: 10.1037/a0018413
Craparo, G., David, V., Costanzo, G., & Gori, A.
(2018). Cosa Nostra and the Camorra:
Assessment of personality, alexithymic traits,
and attachment styles. International Journal of
Law and Psychiatry, 58, 17–26. doi: 10.1016/
j.ijlp.2018.02.010
Cronk, L., & Leech, B. L. (2013). Meeting at
Grand Central: Understanding the social and
evolutionary roots of cooperation. Princeton,
NJ: Princeton University Press.
Currie, T. E., Turchin, P., & Gavrilets, S. (2019).
History of agriculture and intensity of warfare
shaped the evolution of large-scale human
societies in Afro-Eurasia. SocArXiv Papers.
doi: 10.31235/osf.io/9kmrw
Daly, M., & Wilson, M. (1988). Homicide.
Hawthorne, NY: Aldine.
Dawkins, R. (1976). The selfish gene. Oxford,
UK: Oxford University Press.
Dimico, A., Isopi, A., & Olsson, O. (2017). Origins
of the Sicilian Mafia: The market for lemons.
The Journal of Economic History, 77, 1083–
1115. doi: 10.1017/S002205071700078X
Dudai, R. (2018). Underground penality: The
IRA’s punishment of informers. Punishment
& Society, 20, 375–395. doi: 10.1177/
1462474517701302
Duntley, J. D., & Shackelford, T. K. (2008).
Darwinian foundations of crime and law.
Aggression and Violent Behaviour, 13,
373–382. doi: 10.1093/acprof:oso/97801953
25188.001.0001
Durrant, R. (2016). Putting risk factors in their
place: An evolutionary-developmental
approach to understanding risk. Psychology,
Crime & Law, 22, 17–32.
Durrant, R. (2018). An introduction to
criminology psychology (2nd edition).
London, UK: Routledge.
Durrant, R. (2019). Evolutionary approaches to
understanding crime: Explaining the gender
gap in offending. Psychology, Crime & Law,
25, 589–608. doi: 10.1080/1068316X.2018.
1558224
Durrant, R., Adamson, S., Todd, F., & Sellman, D.
(2009). Drug use and addiction: An
evolutionary perspective. Australian and New
Zealand Journal of Psychiatry, 43, 1049–1056.
doi: 10.3109/00048670903270449
Durrant, R., & Poppelwell, Z. (2017). Religion,
crime and punishment: An evolutionary
perspective. Cham, Switzerland: Springer.
Durrant, R., & Ward, T. (2015). Evolutionary
criminology: Towards a comprehensive
explanation for crime. Amsterdam,
Netherlands: Academic Press.
Fehr, E., & Rockenbach, B. (2003). Detrimental
effects of sanctions on human altruism.
Nature, 422, 137–140.
Feinstein, A., & Holden, P. (2014). Arms
trafficking. In L. Paoli (Ed.), The Oxford
handbook of organized crime (pp. 444–459).
Oxford, UK: Oxford University Press.
Gintis, H., van Schaik, C., & Boehm, C. (2019).
Zoon politikon: The evolutionary origins of
human socio-political systems. Behavioural
Processes, 161, 17–30.
Hamilton, W. D. (1964). The genetical evolution
of social behavior: I. Journal of Theoretical
Biology, 7, 1–16. doi: 10.1016/0022-5193
(64)90038-4
Henrich, J. (2016). The secret of our success: How
culture is driving human evolution, domesticat-
ing our species, and making us smarter. Prince-
ton, NJ: Princeton University Press.
 EVOLUTIONARY PSYCHOLOGY AND ORGANIZED CRIME
Hill, P. (2014). The Japanese Yakuza. In L. Paoli
(Ed.), The Oxford handbook of organized
crime (pp. 234–253). Oxford, UK: Oxford
University Press.
Hirschfeld, K. (2015). Gangster states:
Organized crime, kleptocracy and political
collapse. New York, NY: Palgrave Macmillan.
Jackson, J. C., Choi, V. K., & Gelfand, M. J.
(2019). Revenge: A multi-level review and syn-
thesis. Annual Review of Psychology, 70,
319–345. doi: 10.1146/annurev-psych-010418
Kassab, H. S., & Rosen, J. D. (2019). Illicit
markets, organized crime, and global security
(pp. 63–85). Cham, Switzerland: Palgrave
Macmillan.
Kavish, N., Fowler-Finn, K., & Boutwell, B. B.
(2017). Criminology’s modern synthesis:
Remaking the science of crime with Darwinian
insight. In T. K. Shackelford & V. Zeigler-Hill
(Eds.), The evolution of psychopathology
(pp. 171–183). Cham, Switzerland: Springer
International Publishing.
Kenrick, D. T., Griskevicius, V., Neuberg, S. L., &
Schaller, M. (2010). Renovating the pyramid of
needs: Contemporary extensions built upon
ancient foundations. Perspectives in Psycho-
logical Science, 5, 292–314. doi: 10.1177/
1745691610369469
Khadjavi, M., & Lange, A. (2013). Prisoners and
their dilemma. Journal of Economic
Behaviour and Organization, 92, 163–175.
doi: 10.1016/j.jebo.2013.05.015
Kirby, S., Francis, B., Humphreys, L., & Soothill,
K. (2016). Using the UK general offender
database as a means to measure and
analyse organized crime. Policing: An Inter-
national Journal of Police Strategies and
Management, 39, 78–94. doi: 10.1108/
PJJPSM-03-2015-0024
Kleemans, E. (2014). Theoretical perspectives on
organized crime. In L. Paoli (Ed.), The Oxford
handbook of organized crime (pp. 32–52).
Oxford, UK: Oxford University Press.
Kleemans, E. R., & Smit, M. (2014). Human
smuggling, human trafficking, and
exploitation in the sex industry. In L. Paoli
(Ed.), The Oxford handbook of organized
crime (pp. 381–401). Oxford, UK: Oxford
University Press.
Koivu, K. L. (2016). In the shadow of the state:
Mafias and illicit markets. Comparative
Political Studies, 49, 155–183.
313
Kurzban, R., Burton-Chellew, M. N., & West,
S. A. (2015). The evolution of altruism in
humans. Annual Review of Psychology,
66. doi: 10.1146/annurev-psych-010814-
015355
Lantz, B. (2019). Co-offending group composi-
tion and violence: The impact of sex, age,
and group size on co-offending violence.
Crime & Delinquency. Advance online publi-
cation. doi: 10.1177/0011128719834564
Lenzi, M., Sharkey, J., Vieno, A., Mayworm, A.,
Dougherty, D., & Nylund-Gibson, K. (2015).
Adolescent gang involvement: The role of
individual, family, peer, and school factors in
a multilevel perspective. Aggressive Behavior,
41, 386–397.
Levi, M. (1998). Perspectives on ‘organised
crime’: An overview. The Howard Journal,
37, 335–345.
Levi, M. (2014). Money laundering. In L. Paoli
(Ed.), The Oxford handbook of organized
crime (pp. 419–443). Oxford, UK: Oxford
University Press.
Maier, K. H., & Ricciardelli, R. (2019). The
prisoner’s dilemma: How male prisoners
experience and respond to penal threat
when incarcerated. Punishment & Society, 2,
231–250. doi: 10.1177/1462474518757091
McCullough, M. E., Kurzban, R., & Tabak, B. A.
(2013). Cognitive systems for revenge and
forgiveness. Behavioral and Brain Sciences,
36, 1–58.
McDonald, M. M., Navarrete, C. D., & van
Vugt, M. (2012). Evolution and the
psychology of intergroup conflict: The male
warrior hypothesis. Proceedings of the
Royal Society of London, B, 367,
670–679.
Meier, S., Pierce, L., Vaccaro, A., & La Cara, B.
(2016). Trust and in-group favouritism in a
culture of crime. Journal of Economic
Behavior & Organization, 132, 78–92. doi:
10.1016/j.jebo.2016.09.005
Nese, A., O’Higgins, N., Sbriglia, P., & Scudiero,
M. (2018). Cooperation, punishment and
organized crime: A lab-in-the-field experi-
ment in southern Italy. European Economic
Review, 107, 86–98. doi: 10.1016/j.
euroecorev.2018.05.004
Nesse, R. M. (1994). An evolutionary perspective
on substance abuse. Ethology and
Sociobiology, 15, 339–348.
314 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Norenzayan, A. (2013). Big gods: How religion
transformed cooperation and conflict.
Princeton, NJ: Princeton University Press.
Norenzayan, A., Shariff, A. F., Willard A. K.,
Slingerland, W. E., Gervais, R. A., McNamara,
R. A., & Henrich, J. (2016). The cultural
evolution of prosocial religions. Behavioral
and Brain Sciences, 39, e1.
Paoli, L. (2003). Mafia brotherhoods: Organized
crime Italian style. New York, NY: Oxford
University Press.
Paoli, L. (2016). Towards a theory of organized
crime: Some preliminary reflections. In G. A.
Antonopoulos (Ed.), Illegal entrepreneurship,
organised crime, and social control (pp. 3–
17). Cham, Switzerland: Springer.
Paoli, L., & Vander Beken, T. (2014). Organized
crime: A contested concept. In L. Paoli (Ed.),
The Oxford handbook of organized crime
(pp. 13–31). Oxford, UK: Oxford University
Press.
Pizzini-Gambetta, V. (2014). Organized crime:
The gender constraints of illegal markets.
In R. Gartner & B. McCarthy (Eds.), The
Oxford handbook of gender, sex, and crime
(pp. 448–467). Oxford, UK: Oxford University
Press.
Purzycki, B. G., Henrich, J., Apicella, C.,
Atkinson, Q. C., Baimel, A., Cohen, E., &
Norenzayan, A. (2018). The evolution of
religion and morality; A synthesis of
ethnographic and experimental evidence
from eight societies. Religion, Brain &
Behaviour, 8, 101–132. doi: 10.1080/
2153599X.2016.1267027.
Ragatz, L. L., Fremouw, W., & Baker, E. (2012).
The psychological profile of white collar
offenders: Demographics, criminal thinking,
psychopathic traits, and psychopathology.
Criminal Justice & Behavior, 39, 978–997.
Reuter, P. (2014). Drug markets and organized
crime. In L. Paoli (Ed.), The Oxford handbook
of organized crime (pp. 359–380). Oxford,
UK: Oxford University Press.
Reuter, P. (2016). On the multiple sources of
violence in drug markets. Criminology and
Public Policy, 15, 1–7.
Richerson, P., Baldini, R., Bell, A. V., Demps, K.,
Frost, K., Hillis, V., & & Ross, C. (2016).
Cultural group selection plays an essential
role in explaining human cooperation: A
sketch of the evidence. Behavioural and
Brain Science, 39, e30. doi: 10.1017/
S0140525X1400106X
Roach, J., & Pease, K. (2013). Evolution and
crime. London, UK: Routledge.
Roulette, C. J., Kazanji, M., Breurec, S., &
Hagen, E. H. (2018). High prevalence of
cannabis use among Aka foragers of the
Congo basin and its possible relationship to
helminthiasis. American Journal of Human
Biology, 28, 5–15.
Saroglou, V. (2011). Believing, bonding,
behaving, and belonging: The big four religious
dimensions and cultural variation. Journal of
Cross-Cultural Psychology, 42, 1320–1340.
doi: 10.1177/0022022111412267
Scheidel, W. (2017). The great leveller: Violence
and the history of inequality from the stone
age to the twenty-first century. Princeton,
NJ: Princeton University Press.
Scott, J. C. (2017). Against the grain: A deep
history of the earliest states. New Haven, CT:
Yale University Press.
Spapens, T. (2014). Illegal gambling. In L. Paoli
(Ed.), The Oxford handbook of organized
crime (pp. 402–418). Oxford, UK: Oxford
University Press.
Thoumi, F. E. (2014). Organized crime in
Colombia: The actors running the illegal
drug industry. In L. Paoli (Ed.), The Oxford
handbook of organized crime (pp. 177–195).
Oxford, UK: Oxford University Press.
Tilly, C. (1985/1992). War making and state
making as organized crime. In C. Besteman
(Ed.), Violence: A reader (pp. 35–60). New
York, NY: New York University Press.
Tooby, J., & Cosmides, L. (2010). Groups in
mind: The coalitional roots of war and
morality. In H. Høgh-Olesen (Ed.), Human
morality and sociality: Evolutionary and
comparative perspectives, (pp. 191–234).
Cham, Switzerland: Palgrave Macmillan.
Trivers, R. L. (1971). The evolution of reciprocal
altruism. Quarterly Review of Biology, 46,
35–57.
Turchin, P. (2016). Ultra society: How 10,000 years
of war made humans the greatest co-operators
on earth. Chaplin, CT: Beresta Books.
Turchin, P., Whitehouse, H., Korotayev, A.,
Francois, P., Hoyer, D., Peregrine, P., & Currie,
T. E. (2018). Evolutionary pathways to
statehood: Old theories and new data.
SocArXiv Papers. doi: 10.31235/osf.io/h7tr6
 EVOLUTIONARY PSYCHOLOGY AND ORGANIZED CRIME
United Nations Office of Drugs and Crime
(2004). United Nations Convention Against
Transnational Organized Crime and the
Protocols Thereto. Retrieved from: www.
unodc.org/documents/treaties/UNTOC/
Publications/TOC%20Convention/
TOCebook-e.pdf (Accessed 15 April 2019)
van Dijk, M. (2007). Mafia markers: Assessing
organized crime and its impact upon
societies. Trends in Organized Crime, 10,
39–56. doi: 10.1007/s12117-007-9013-x
van Dijk, M., Kleemans, E., & Eichelsheim, V.
(2018). Children of organized crime offenders:
Like father, like child? An explorative and
qualitative study into mechanisms of
intergenerational (dis)continuity in organized
crime families. European Journal of Crime and
Policy Research. Advance online publication.
doi: 10.1007/s10610-018-9381-6
van San, M. (2011). The appeal of ‘dangerous’
men. On the role of women in organized
crime. Trends in Organized Crime, 14, 281–
297. doi: 10.10007/s12117-011-9128-y
von Lampe, K. (2016a). Organized crime:
Analyzing illegal activities, criminal structures,
and extra-legal governance. Los Angeles,
CA: Sage.
von Lampe, K (2016b). The ties that bind: A
taxonomy of associational criminal structures.
In G. A. Antonopoulos (Ed.), Illegal
entrepreneurship, organised crime, and social
control (pp. 19–35). Cham, Switzerland:
Springer.
von Rueden, C., Gurven, M., & Kaplan, H. (2011).
Why do men seek status? Fitness payoffs to
dominance and prestige. Proceedings of the
Royal Society of London, B, 278, 2223–2232.
doi: 10.1098/rspb.2010.2145
von Rueden, C., & Jaeggi, A. V. (2016). Men’s
status and reproductive success in 33
nonindustrial societies: Effects of subsistence,
marriage system, and reproductive strategy.
315
Proceedings of the National Academy of
Sciences, 113, 10824–10829. doi: 10.1073/
pnas.1606800113
van Vugt, M., De Cremer, D., & Janssen, D. P.
(2007). Gender differences in cooperation
and competition: The male warrior
hypothesis. Psychological Science, 18, 19–23.
doi: 10.1111/j.1467-9280.2007.01842.x
van Vugt, M., & Tybur, J. M. (2016). The
evolutionary foundations of hierarchy:
Status, dominance, prestige, leadership. In
D. M. Buss (Ed.), Handbook of evolutionary
psychology (2nd edition) (pp. 788–809).
Hoboken, NJ: Wiley.
Varese, F. (2014). Protection and extortion. In L.
Paoli (Ed.), The Oxford handbook of
organized crime (pp. 343–358). Oxford, UK:
Oxford University Press.
Walsh, A., & Jorgensen, C. (2018). Evolutionary
theory and criminology. In R. Hopcroft (Ed.),
Oxford handbook of evolution, biology and
society (pp. 517–542). Oxford, UK: Oxford
University Press.
Weber, M. (1978). Economy and society: An
outline of interpretive sociology. Berkeley,
CA: University of California Press.
West, S. A., El Mouden, C., & Gardner, A. (2011).
Sixteen common misconceptions about the
evolution of cooperation in humans. Evolution
and Human Behaviour, 32, 231–262. doi:
10.1016/j.evolhumbehav.2010.08.001
Whitehouse, H., McQuinn, B., Buhrmester, M.,
& Swann, W. (2014). Brothers in arms:
Libyan revolutionaries bond like family.
Proceedings of the National Academy of
Science, 111, 17783–17785. doi: 10.1073/
pnas.1416284111
Wrangham, R. W., & Glowacki, L. (2012).
Intergroup aggression in chimpanzees and
war in nomadic hunter-gatherers: Evaluating
the chimpanzee model. Human Nature, 23,
5–29. doi: 10.1007

Evolutionary Psychology
and Warfare
INTRODUCTION
The scientific study of the evolution of
human coalitional aggression has exploded
over the last several decades. In four parts, I
explore and integrate several of the core
frameworks that have emerged to explain the
human practice of inter-group violence. First,
we have a better understanding of the condi-
tions required for the evolution of adapta-
tions for coalitional aggression. These
include: the presence within a species of a
unique socio-ecology that facilitates the
emergence of a group-based population
structure; the existence of evolved cognitive
abilities that make possible the navigation of
challenges associated with group living that
are necessary for coalitional aggression (e.g.,
some forms of collective action); and the
presence of low-cost/high-benefit reproduc-
tive opportunities that are uniquely well-
seized via coalitional aggression. In short, a
species must live in groups, it must be able to
solve complex collective action problems,
16
Anthony C. Lopez
and violence must have been reproductively
beneficial, on average.
Second, given an understanding of the the-
oretical requirements, we are better situated
to examine the historic and prehistoric record
for evidence of the existence of these con-
ditions. This involves examination of many
lines of evidence such as studies of non-
human primates, the archaeological record of
coalitional aggression, and modern hunter-
gatherer evidence, as well as game-theoretic
evidence and lab evidence of the design of
psychological mechanisms for coalitional
aggression. The latter is particularly impor-
tant given that the logic of natural selection
indicates that the extant structure of adapta-
tions themselves provides, in part, a window
into ancestral landscapes.
Third, I explore and integrate current
evidence of psychological adaptations for
­coalitional aggression. There are two forms
of inter-group engagement that have received
the most attention: raids and battles. I also
examine five areas of inquiry that suggest
 EVOLUTIONARY PSYCHOLOGY AND WARFARE
special design for coalitional aggression.
These are: the collective action problem of
coordinated violence; parochial altruism;
attacker–defender asymmetries; leader–­
follower dynamics; and sex differences in the
costs and benefits of violence.
Fourth, I speculate on the historical emer-
gence of modern human warfare. I do not use
‘coalitional aggression’ and ‘warfare’ inter-
changeably. Instead, evolved psychological
adaptations for small-scale coalitional aggres-
sion are what make the emergence of large-
scale human warfare possible. Thus, in addition
to the biological and evolutionary arguments
necessary to establish the existence of adapta-
tions for coalitional aggression, I conclude with
a speculative discussion of the historical and
cultural shifts that may help to explain the com-
plex modern phenomenon of human warfare.
UNIVERSAL CONSTRAINTS ON THE
EVOLVABILITY OF COALITIONAL
VIOLENCE
There are at least three necessary conditions
that favor the emergence of lethal coalitional
aggression in its broadest sense. First, the
species’ ecology must be such that the organ-
ism is adapted for group-based interaction
(Chapman, 1986; Chapman et  al., 1995;
Chapman and Chapman, 2000). Second, the
species must possess neurocomputational
machinery that allows it to solve at least a
subset of a potentially infinite range of
n-person collective action problems pertinent
to coalitional aggression (Tooby and
Cosmides, 1988; Tooby et  al., 2006). This
can range from relatively simple coordina-
tion problems, to more complex cooperation
problems involving conspiracy, coercion, and
alliance. For example, the existence of sym-
bolic language is crucial for enabling com-
plex and large-scale coalitional action
(Wrangham, 2019). Third, opportunities for
coalitional violence must exist that are
sufficiently low-cost and/or beneficial – in
317
reproductive terms – to favor coalitional
violence as a unique strategy relative to alter-
native fitness-enhancing strategies (Enquist
and Leimar, 1990; Lopez, 2016a).
The first two conditions being met (group-
structured population; coalitional psychology
of coordination and cooperation) mean that a
species likely possesses some of the psycho-
logical tools that can be used for coalitional
aggression, but without the third condition,
coalitional aggression is unlikely to occur
via these tools alone. Similarly, the mere
presence of low-cost/high-benefit opportu-
nities for coalitional violence will not ipso
facto lead a species to coalitional aggression
if there is no group structure in the popula-
tion, or if language and cognitive abilities to
coordinate are limited or absent. For exam-
ple, this simple difference may be one among
many reasons that chimpanzees engage in
an intense and lethal form of coalitional vio-
lence that is nevertheless limited in duration
and complexity relative to humans.
Although our ability to peer into the ances-
tral past is imperfect, the ancestral past is far
from unknowable. Despite disagreement on
the origins and prevalence of warfare, there
is near universal agreement that the nec-
essary conditions for coalitional violence
existed among ancestral humans throughout
the Holocene, Pleistocene, and perhaps ear-
lier to a common ancestor with chimpanzees.
I briefly discuss the evidence for each of the
three necessary conditions.
First, although there is disagreement on
the size and structure of early human forager
bands, there is broad agreement that humans
associated in groups, that these groups ranged
in size from 25 to several hundred individuals
(depending on distinctions between bands,
tribes, societies, ethnolinguistic units, etc.),
and that these groups included a substantial
number of kin as well as non-kin (Hill et al.,
2011; Bird et  al., 2019). Group structure is
unequivocally a component of our species’
long evolutionary history.
Second, there is broad evidence that
humans are instinctively good at solving
318 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
coordination and cooperation problems in both
dyadic and n-person contexts (Tooby et  al.,
2006). Additionally, language and symbolic
reasoning in humans are such that we are able
to build cultural institutions that interact with
motivated reasoning to either enhance or limit
group- and individual-level dynamics (Boyer
and Petersen, 2011). It should come as no sur-
prise, therefore, that political institutions often
focus on the nature and expression of author-
ity (Smith et al., 2007) and the distribution of
resources (Petersen, 2012) – two recurrent and
important challenges faced by all groups. In
short, we have evolved in groups, and we are
well-adapted for navigating the many recur-
rent and reproductively significant challenges
associated with this coalitional landscape.
Third, opportunities for low-cost/high-
benefit coalitional violence must exist. There
are at least three foundational components
that determine the cost profile of coalitional
violence in almost any setting. First is the
proximity of out-groups. The more geograph-
ically distal groups are, the more quickly the
costs of coalitional violence accumulate in
terms of effort and coordination. Second, tar-
gets must reliably exist that are vulnerable to
low-cost attack. Third, their injury or death
must entail, either directly or indirectly, some
fitness benefit to the attacker(s). In chimpan-
zees, for example, group structure and feed-
ing habits result in occasions of lone travelers
who can be ambushed and killed, and the
reproductive benefits of such an attack, when
delivered with overwhelming force of num-
bers, are substantial (e.g., see Wrangham,
1999; Wilson et  al., 2014). Similarly, in
humans, relatively frail bodies (i.e. in com-
parison to chimpanzees) combined with his-
torical innovation in lethal weaponry and
conspiratorial ability have turned humans
into ‘quintessential first-strike creatures’
(Gat, 2006). Furthermore, it is not uncom-
mon for inter-group raids to entail significant
reproductive benefits as well (Glowacki and
Wrangham, 2015). In short, low-cost/high-
benefit opportunities for human coalitional
violence have long existed.
These three universal constraint conditions
on the evolution of coalitional violence have
been met in humans. However, the presence
of these three conditions still does not tell us
much about the specific form that coalitional
violence will take in any species, and neither
does it allow us to make many inferences
about downstream or related considerations,
such as the possibility of cultural evolution
or the existence of sex differences in fighting.
Rather, it gives us a universal cross-species
framework for conceptualizing the necessary
phenotypic ‘starting point’ of coalitional vio-
lence. In order to provide a richer explana-
tion of the form and function of coalitional
violence observed in a given species, we
must not only ask whether the three condi-
tions mentioned above existed ancestrally;
we must also uncover species-specific details
regarding demographic and socio-ecological
variables that influence the form and function
of coalitional violence such as group size and
composition, fission–fusion dynamics, mat-
ing structure, parental-investment dynamics,
and the nature and distribution of opportuni-
ties for violence between groups. It is to this
evidence that I now turn.
EVIDENCE OF ANCESTRAL HUMAN
COALITIONAL VIOLENCE
Given the above, the logical next steps are to
trace the practice of coalitional violence in
humans, i.e. when did it emerge, what did it
look like, and how has it changed over time? I
address the first two questions in this section,
and the final question in the next section.
Emergence of Coalitional Violence
Scholarship on the evolutionary origins of
warfare is bifurcated in two traditions that
Allen refers to as the long chronology and
short chronology of warfare (Allen and
Jones, 2014). The short chronology views
 EVOLUTIONARY PSYCHOLOGY AND WARFARE
human warfare as having emerged by or
shortly after the agricultural revolution at the
start of the Holocene, and in response to new
social and ecological pressures relating to
sedentarism and the accumulation of wealth
(Otterbein, 1997; Kelly, 2000; Fry, 2007; Fry
and Söderberg, 2013). Prior to this period,
humans likely engaged in sporadic violent
encounters, but most occurred within kin
groups rather than between them and rarely
involved anything more complex than the
targeted killing of single individuals. In con-
trast, the long chronology accepts the chim-
panzee model of low-cost/high-benefit
opportunistic coalitional killing as perhaps
the most basic form of warfare, upon which
humans slowly and over time built greater
complexity. In other words, we have been
engaging in coalitional violence at least since
the split with chimpanzees (Wrangham, 1999;
Gat, 2006; McDonald et  al., 2012; Lopez,
2016b; Glowacki et al., 2017). Both short and
long chronologies recognize that develop-
ments in the early Holocene brought signifi-
cant changes to human inter-group violence,
but whereas the short chronology sees this as
the beginning of ‘warfare’ per se, the long
chronology sees it as a period of significant
transition in the scale and complexity of war-
fare (Keeley, 1996; Lopez, 2019a).
This dispute is not so much regarding
what humans ‘were actually doing’ in the
Pleistocene; rather, the more contentious
issues hinge on whether various activities
should be labeled ‘warfare’. Therefore, the
question ‘when did warfare begin?’ is some-
what indeterminate to the extent that it rests on
arbitrary definitional assumptions, and should
be engaged with methodological caution.
Forms of Ancestral Coalitional
Violence
Both chronologies of war recognize that raid-
ing has been and remains the oldest and com-
monest form of human inter-group violence
(Keeley, 1996; Otterbein, 2004; Gat, 2006).
319
Raids – sometimes called stealth raids –
­display particular characteristics: occurring
by stealth and/or at night; exploiting cover/
concealment; targeting one or a few lone
unsuspecting individuals by surprise/ambush;
quick retreat; exploiting numerical asym-
metries (e.g., at least 3:1 force imbalance);
overwhelmingly initiated by males; over-
whelmingly targeting out-group males. Raids
are designed to maximize effect relative to
the expected risks, principally via numerical
imbalance and the element of surprise.
Humans are indeed ‘first-strike creatures’;
however, in the context of lethal weaponry
and symbolic language that facilitates coor-
dinated revenge, human raiding is relatively
riskier than chimpanzee raiding and may be
‘scaffolded’ to some extent by cultural insti-
tutions that incentivize some forms of inter-
group or otherwise ‘heroic’ violence and
sacrifice (Wrangham and Glowacki, 2012;
Glowacki and Wrangham, 2013).
Prior to the Holocene, direct evidence that
humans engaged in anything other than coali-
tional raiding is patchy at best. Despite occa-
sional findings of mass graves from as early
as the end of the Pleistocene (Lahr et  al.,
2016), there is neither direct nor indirect evi-
dence that, for example, humans assembled
in more than a handful of individuals at a
time for the purpose of relatively symmetric
melee coalitional aggression. In other words,
‘battles’ – in contrast to raids – are relatively
recent and rare in both the ethnographic and
archaeological record, and with no analogue
in chimpanzee behavior.
AN EVOLVED PSYCHOLOGY OF
COALITIONAL VIOLENCE?
The universal conditions for the evolution of
coalitional violence are satisfied in humans,
and we also know that ancestral humans
engaged in at least some forms of inter-group
violence (e.g., raids and possibly battles).
Combined with incomplete but rich available
320 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
information regarding the ancestral
coalitional landscapes in which humans
­ minimizing what would be an otherwise chal-
lived, it is possible to identify several over-
lapping and interconnected classes of adap-
tive problems that existed, and these problems
seem to underlie all instances of coalitional
violence to a variable degree. These are: the
collective action problem of coordinated vio-
lence; parochial altruism; attacker–defender
asymmetries; leader–follower dynamics; and
sex differences in the costs and benefits of
violence.
The Collective Action Problem of
Coordinated Violence
Despite significant disagreement regarding
the origins and frequency of warfare in our
species’ history, most agree that at the heart
of the puzzle of warfare lies a within-group
collective action problem (Kitchen and
Beehner, 2007; Crofoot and Gilby, 2012;
Pietraszewski, 2016; Glowacki et  al., 2017;
Lopez, 2019b; Wiessner, 2019). Large, com-
plex, sedentary societies are often able to
solve these problems through the use of
coercive institutions, while relatively smaller
nomadic societies tend to solve these prob-
lems via the implicit or explicit distribution
of material rewards or the leveraging of per-
sonal reputation and status.
Any agonistic contest entails risks to the
participants such as injury or retaliation, and
species that engage in such contests must
possess evolved systems that regulate an
individual’s participation in such contests,
such as whether to engage, escalate, sub-
mit, or run away (Parker, 1974; Sell, 2011;
Chapin et al., 2019). The problem is magni-
fied among coalitions, where assessments of
the likelihood of success are compounded by
assessments of the feasibility and strength of
within-group coordination and cooperation.
It is probably no surprise again, that the most
common form of human coalitional violence
is the raid; it is a form of low-cost violence
that can be initiated with a small group of
motivated individuals and concludes quickly,
lenging problem of n-person tracking that is
difficult for even modern militaries to master
and sustain (Biddle, 2006).
At a basic level, the initiation of any act
of coalitional violence requires a solution
to the near-simultaneous problems of indi-
vidual commitment as well as inter-personal
efforts to manipulate the behavior and com-
mitment of within-group others (Tooby and
Cosmides, 1988; Lopez, 2017, 2019b). These
are problems of participation and enforce-
ment: should I participate? (How) should I
manipulate the behavior of others toward (or
away from) participation? Again, although
modern societies solve these problems with
coercive institutions, these problems were
solved by our ancestors via the mechanisms
of individual formidability and interest. This
helps to explain why the commonest form of
human coalitional violence is that of small,
unstable, fraternal coalitions that sporadi-
cally assemble to take advantage of offensive
opportunities.
Parochial Altruism
The conceptual core of parochial altruism is
simple, yet its simplicity betrays what has
become a complex and contentious area of
study. In its simplest form, parochial altruism
refers to a class of motivations and behavior
in which one pays a cost to benefit at least
some members of one’s own group at the
direct or indirect expense of at least some
members of an out-group. For example, lab
and field evidence over many decades sug-
gest that the mere presence of inter-group
competition (violent or otherwise) is often
sufficient to trigger many forms of within-
group cooperation, sacrifice, and altruism
(Puurtinen and Mappes, 2009; Gneezy and
Fessler, 2011; Bauer et  al., 2016; Chang
et al., 2016; but see also Jordan et al., 2017).
In the extreme, this is taken as evidence that
inter-group competition in our species’
 EVOLUTIONARY PSYCHOLOGY AND WARFARE
evolutionary history is directly responsible
for our remarkable capacity for large-scale
cooperation, even among strangers. At the
very least, it suggests that the two processes
(in-group favoritism and out-group deroga-
tion) may have co-evolved to some degree
(Choi and Bowles, 2007; Bowles, 2009).
However, it has been demonstrated that
the desire to harm an out-group is often moti-
vated by individual-level incentives (e.g., per-
sonal status, reputation, privatized material
resources), and it has also been demonstrated
that in-group favoritism does not necessar-
ily also trigger out-group hate (Patton, 2005;
Gavrilets and Fortunato, 2014; Rusch, 2014;
Weisel and Böhm, 2015; Romano et  al.,
2017; Doğan et  al., 2018). Therefore, the
evidently tight linkage between inter-group
competition and within-group cooperation
that lies at the center of parochial altruism is
conditional at best and likely engages a range
of individual-level and group-level incentives
depending on the immediate context. For
example, individuals may be driven more by
group-level benefits in domains of defense,
while driven more by individual-level ben-
efits in domains of offense (Rusch, 2013;
Lopez, 2017). An adaptationist perspective
on coalitional violence, therefore, provides
us with a lens through which to examine
the interplay between strategic context (e.g.,
defense/offense) and individual differences
pertaining to capability (i.e. formidability,
or resource-holding potential) or interest.
This leads to the next two major domains of
research: attacker/defender asymmetries and
leadership.
Attacker–Defender Asymmetries
Coalitional violence can be defined as the
attempt by a group of individuals to physi-
cally damage at least one individual of
another group. Unless the individuals of
opposed groups are sitting immediately
beside each other, any attack upon an out-
group must minimally endure the costs of
321
travel through time and space. This initial
obligate cost necessarily establishes an asym-
metrical structure to the cost–benefit profile
of attacking versus defending, as well as
alternative strategies (e.g., to run/disperse).
For example, although much emphasis is
rightly given to the fact that attackers can
exploit surprise and relative numbers if
appropriate targets are chosen, attackers also
endure significant costs of coordination,
motivation, and target retaliation, in addition
to the obligate costs mentioned above. In
contrast, defenders can exploit resident
advantages such as superior knowledge of
local terrain and may often find it easier to
solve coordination problems. Mirville et al.,
for example, find that ‘large groups may be
unsuccessful in inter-group conflict if they
enter a smaller group’s home range, as the
smaller group may respond with more
aggression to defend the area’ (2018: 178). It
is no surprise, therefore, that De Dreu and
Gross (2018) find that, on average, across
many contexts, defenders have a greater suc-
cess rate than attackers, which must be at
least partly due to this initial obligate
asymmetry.
Buckner and Glowacki (2019), however,
usefully point out that attackers have the
advantage of ‘setting the stage of conflict’,
which stacks the odds of success in attackers’
favor. It is clear that when attackers can assem-
ble for brief incursions to catch defenders off-
balance and unaware, the result is a low-cost/
high-benefit effort that can be strongly favored
by selection. As mentioned above, Gat (2006)
extends this analysis by arguing that, given the
relatively weak physique of humans combined
with the use of weaponry in coalitional vio-
lence, humans evolved to be ‘quintessential
first-strike creatures’. To the extent this is true,
then in the language of international relations
theory, ancestral humans may have evolved in
an ‘offense-dominant’ environment, in which
the costs of defending were greater than the
costs of attacking (Jervis, 1978; Biddle, 2001).
Despite the fact that this ‘offense–defense’
balance has shifted numerous times since
322 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
the Pleistocene, Gat argues that the gamble
of inter-group violence was probably stacked
in favor of well-organized attackers – as
Buckner and Glowacki (2019) agree – which
is perhaps best characterized by the well-
known adage that the best defense is a good
offense.
Attacker–defender asymmetries are prob-
ably more obvious and relevant in raiding
contexts or contexts of small-scale conflict
than in larger battle contexts. Furthermore,
in raiding contexts, defenders face a public-
good problem, in which the benefits of suc-
cess are general to the group, while attackers
face a club-good problem, in which the ben-
efits of success are more privatized among
the attackers themselves. These attacker–
defender asymmetries help to clarify the set
of unique problems that must be resolved
in each domain in order for either attack or
defense to be successful (Lopez, 2017). There
are clearly group-level macro variables that
impinge upon outcomes, especially relative
numbers. However, the successful prosecu-
tion of attack and defense invariably depends
on group members being able to effectively
manage and navigate a constellation of inter-
individual dynamics (Tooby and Cosmides,
1988; Tooby et  al., 2006; Pietraszewski,
2016; Wiessner, 2019). Individuals must
solve a range of domain-specific collective
action problems specific to warfare, the solu-
tions to which are conditional upon, inter alia,
whether individuals are initiating or respond-
ing to an attack.
Leader–Follower Dynamics
Leaders – whether formal or informal – are
not intrinsically necessary for solving collec-
tive action problems; however, their presence
has been demonstrated to facilitate success in
inter-group contexts. Although ancestral
human societies are believed to be relatively
egalitarian, these groups are also marked by
asymmetries of influence and status that
shape motivation and behavior (von Rueden,
2020). Thus, although the pattern of lethal
violence in hunter-gatherers is characterized
by relatively leaderless raiding parties organ-
ized in the absence of overt coercive manipu-
lation, this is better described as the absence
of formal leadership, in which behavior is
instead informally motivated by reputation
and reciprocity. In short, the ancestral char-
acterization of ‘leaderless’ societies under-
states the importance of relative influence
and status as constraints and facilitators of
collective action (Boehm, 1999, 2012).
Leaders can help resolve coordination
problems (e.g., Where to meet? When to
leave?) as well as resolve and enforce more
complicated problems pertaining to the dis-
tribution of costs and benefits of aggression.
By definition, highly formidable individuals
have both a greater capacity to punish and
to reward, as well as a greater incentive for
attack in many cases. Therefore, highly for-
midable individuals are also more likely to
resort to deception to recalibrate otherwise
unmotivated minds. For example, when a
formidable individual identifies a window
of opportunity for attack, yet willing others
are immediately unavailable, the initiator can
resort to the direct manipulation of costs and
benefits, or resort to an indirect manipulation
of the strategic context. Direct manipulation
of cost and benefit is clear: leaders are often
those who, by definition, are most able to
distribute such costs and benefits. However,
initiators may also manipulate others’ beliefs
about the nature of aggression, recasting
opportunistic aggression as defensive or
preemptive necessity (Lopez, 2019b). It has
been well-established that individuals are
often more willing to fight for defense than
for offense (Böhm et al., 2016); if this asym-
metry extended to ancestral environments,
the result is a motivated bias among strongly
incentivized initiators (i.e. leaders; highly
formidable individuals) to falsely cast oppor-
tunism as necessity.
Coalitional violence is not just a story of
manipulation by the powerful; in fact, lead-
ers can provide a valuable public good in
 EVOLUTIONARY PSYCHOLOGY AND WARFARE
the prosecution of inter-group competition
(van Vugt and Kurzban, 2007; Hooper et al.,
2010; Glowacki and von Rueden, 2015).
Thus, research has found that individuals
seem to prefer dominant or masculine lead-
ers in times of inter-group violence, precisely
because there is an implicit understanding
that these individuals, in these contexts, are
able to provide a unique service. This has led
to a growing field of research demonstrat-
ing that facial and vocal cues are implicitly
used by individuals to assess leadership in
specific contexts (Anderson and Klofstad,
2012; Klofstad et  al., 2012; Spisak, 2012;
Spisak et  al., 2012; Laustsen and Petersen,
2017). In short, despite a relatively egalitar-
ian group structure, individual variance in
both ­formidability and interest allowed our
ancestors to facilitate coalitional violence
when necessary or beneficial.
Sex Differences in the Costs and
Benefits of Coalitional Violence
The overall pattern of human violence at
both the individual and group levels is that of
male physical violence (van Vugt, 2009;
McDonald et  al., 2012). Cross-culturally,
males are significantly more likely to be both
the perpetrators and targets of physical vio-
lence (Daly and Wilson, 1988). Even in
modern contexts, for example, in which it
has been observed that greater numbers of
women are engaged in acts of suicide terror-
ism, the overall numbers are skewed toward
overwhelmingly male involvement
(Goldstein, 2003).
Participation in physical violence should
be distinguished from support for violence.
Evidence of greater male aggressiveness is
strongest on indicators of physical acts of
violence, and weaker on indicators of indi-
rect or informal support, particularly in
defensive contexts, in which all benefit from
victory due to its public-good nature (Ginges
and Atran, 2011; Lopez, 2017). Thus, it is
little surprise that Fair and Shepherd (2006)
323
find that female support for terrorism is not
only on the rise in several Arab countries,
but that in many instances it equals male
support for terrorism in these contexts. In
other words, violence, as the adaptive out-
put of evolved psychological mechanisms,
is irrevocably conditional, and the structure
of that conditionality depends in part on sex.
While male participation in coalitional vio-
lence is likely highly personalized, physical,
and motived to seek opportunistic gains at
low cost, female participation is likely indi-
rect, greater in defensive contexts, and when
immediate resources are at stake (resources,
territory, children) rather than when surro-
gate resources are at stake, such as status and
honor, although these remain somewhat open
questions (Glowacki and Wrangham, 2013;
Scalise Sugiyama, 2014; Lopez, 2017; Lynch
et al., 2019).
Greater male physical aggressiveness in
humans is directly predicted by sexual selec-
tion and parental investment theory, in which
sex differences in physical aggression follow
asymmetries in parental investment (Trivers,
1972; Daly and Wilson, 1983). Furthermore,
these theoretical foundations have been well
elaborated by research on several established
mechanisms of sexual selection, such as con-
test competition, in which same-sex com-
petitors are excluded from mating through
force, and mate choice, in which the posses-
sion of traits (e.g., intelligence, formidability,
wealth) enhances one’s attractiveness toward
members of the opposite sex (Puts, 2016;
Hill et al., 2017).
BEYOND THE MILITARY HORIZON:
THE EMERGENCE OF WARFARE
Humans possess the three universal criteria
for the evolution of coalitional violence, and
a cursory review of the human pattern of
ancestral violence, alongside a sketch of the
adaptive problems this has generated (e.g.,
collective action problems, attacker–defender
324 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
asymmetries) suggests that humans likely
possess specialized psychological design for
navigating the challenges of coalitional vio-
lence. What this means is that we possess the
psychological tools for reasoning about and
engaging in small-scale coalitional violence.
However, what we observe today arguably
bears little resemblance to the behaviors our
ancestors engaged in. The real question often
reduces to: are the differences between
ancestral and modern ‘warfare’ differences
in degree or kind?
These questions probably cannot be
answered in a general sense. In some arenas,
notably lethal weaponry, modern warfare is a
faint shadow of its ancestral forms. However,
in terms of certain principles of engagement,
such as cover/concealment, the element of
surprise, force-imbalance ratios, rally ‘round
the flag effects, attacker–defender asym-
metries, and many others, modern warfare
often still follows a fundamentally ancestral
logic, even if it is now manifested at much
greater scales (Keeley, 1996; Lopez, 2016b,
2019a). Guerrilla warfare, insurgency, and
hybrid warfare follow kinetic principles of
violence that would likely be intuitive to any
hunter-gatherer (Boot, 2013).
This suggests that the real question
regarding the emergence of human warfare
from smaller-scale versions is probably not
evolutionary so much as historical. In other
words, once the psychological tools of
coalitional violence were in place, significant
changes in demographics and ecology
would have been sufficient to trigger both
qualitative and quantitative innovations in
the scale and method of coalitional violence,
such that at some ambiguous and relatively
arbitrary point in history, what we once
recognized as coalitional violence looks
more and more like modern warfare. How
did this happen?
The answer to this question forces at least
a brief return to an earlier question, that of
the origins of warfare. Recall that the long-
chronology perspective argues that warfare
is evolutionarily old and merely ‘transitions’
after the emergence of agriculture, while the
short-chronology perspective argues that this
transition is in fact an origin. Aside from the
relatively arbitrary issue of what difference
there is between an origin and a transition, a
more productive question is: what ecological
and social shifts were occurring at this time,
and how did this affect the nature of inter-
group conflict?
The first thing to note about the emergence
of agriculture and its effects on inter-group
violence is that it does not occur everywhere
at the same time, and therefore neither does
‘warfare’. Otterbein, for example, while sit-
ting somewhere between the long- and short-
chronology perspectives, argues that an older
form of human warfare predates agriculture,
and emerges as a by-product of hunting.
As big game became scarce, so too did the
practice of warfare, which then disappeared
until social innovations occurring after the
transition to agriculture made the practice of
warfare profitable again. For Otterbein, war-
fare could not have occurred simultaneously
with the emergence of agriculture, since: ‘For
agriculture and permanent settlements to
arise, there must be no warfare’ (Otterbein,
2004: 13). Innovations in agriculture require
peace, and somewhat ironically, the fruits
of agriculture make war again possible and
profitable. In direct contrast, however, Ferrill
argues instead that it was warfare that drove
the need for agriculture. Innovations in
weaponry that occurred around 10,000 BC,
such as bows and slings, increased the need
for defenses, such as fortified walls, and the
resultant decline in mobility meant there was
pressure to find new ways to live off the land
(Ferrill, 1997).
This obvious chicken-or-egg problem
remains unresolved, but what is clear is that
where agriculture emerged, it tended to have
clear impacts upon the political economy of
human groups, which included the possibility
of, and in some cases a perceived imperative
for, what might be labeled ‘modern war-
fare’. Relatively sedentary groups practicing
agriculture were larger, accumulated more
 EVOLUTIONARY PSYCHOLOGY AND WARFARE
wealth, were more effective at command,
control, and logistics, and were more efficient
innovators. Gabriel, similar to Otterbein,
argues that although this transition is under-
way after the transition to agriculture, inter-
group violence does not begin to look like
‘modern warfare’ until about 4,000–2,000
BC, by which time population centers, social
and economic organization, lethal weaponry,
and institutional command coalesce to pro-
duce a form of warfare that is often associ-
ated with nation states – those that exhibit
steep vertical command structures, are char-
acterized by sizeable armies that exploit col-
umn and line formations, and are capable of
greater levels of destruction (Gabriel, 2002).
In other words, it is around this time that the
raid, as the predominant or even exclusive
form of inter-group violence, now shares the
stage with (or in many cases is upstaged by)
conventional battle between formal political
organizations.
As Levy and Thompson describe, this
transition begins in Mesopotamia and Egypt,
followed later by transitions in Greece, Rome,
and China, during which: ‘Weapons became
much more lethal, military organizations
became larger and more complex, and
states became more powerful’ (Levy and
Thompson, 2011: 122)1. In addition to these
first two transitions, Levy and Thompson
identify a third transition occurring within
the last five centuries. Two trends that stand
out during this period include a decreasing
frequency of great power warfare alongside
a pernicious increase in the lethality of
warfare (Gat, 2006; Levy and Thompson,
2011; Pinker, 2011). The particularities
of this transition and the innovations and
security imperatives that in part drove it
are outside the scope of this inquiry; it is
sufficient to note that this long history of the
ever-unfolding nature of warfare is pithily
summed up with Tilly’s famous assertion
that ‘war made the state, and the state made
war’ (Tilly, 1975: 42).
This ironic, even if tragic, connec-
tion between expanding zones of political
325
cooperation and the steady intensification
of warfare is brought to a head in the cur-
rent era, in which great powers are restrained
in their conflicts with each other, and new
forms of violence have largely taken the
place of inter-state violence. In this sense, we
are witnessing a reemergence of small-scale
violence – a renaissance of the raid. This fea-
ture is now characteristic of many forms of
modern political violence. When great pow-
ers do engage in violence, it often takes the
form of so-called ‘limited wars’, in which
quick incursions are meant to target key
individuals, groups, or facilities in order to
degrade or deter an adversary, often a non-
state actor (Osgood, 1957; Stoker, 2016).
When nation states are the targets of vio-
lence, the source may be a terrorist network
or a nation state engaging in ‘hybrid war-
fare’ in which states bring to bear a motley
range of tools, such as propaganda, cyber
instruments, and targeted assassinations,
in order to effect outcomes short of war
(Hoffman, 2007). Lastly, the most common
form of political violence is now civil war
and insurgency (Lopez and Johnson, 2017),
which, again, often bears the ancestral hall-
marks of many raiding dynamics (Boot,
2013). In short, the raid, having fallen from
prominence over two thousand years ago, is
now returning to center stage in our species’
long evolution of warfare.
UNIVERSALITY, UNIQUENESS, AND
CHANGE: WAR AS A WINDOW INTO
HUMAN NATURE
Perhaps it is the author’s conceit, but there is
hardly any domain outside of warfare that
singularly unites so much about what makes
us human – from our ugliest desires to our
noblest sacrifices. The study of warfare is
necessarily a tour through many if not all of
the big questions of human nature and the
nature of human psychology. Many of these
questions are not investigated here, and some
326 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
that are remain nevertheless unresolved.
However, one lesson that does emerge from
this analysis is that humans are social by
design, and at least part of this design facili-
tates aggression between groups.
Evolution is a process of change, and
natural selection is a force that explains one
kind of change: directional change in gene
frequencies that can build complex adapta-
tions over time. The human mind is proposed
to consist, in part, of complex adaptations
designed to facilitate the navigation of the
natural world (which includes the social
world) in a way that would have maximized
reproductive success in ancestral environ-
ments. In other words, these complex adap-
tations are relatively slow to change, which
means that many of the evolved systems that
emerged to solve ancestral problems remain
active today, ‘trying’ to make sense of a mod-
ern world that is a shadow of its ancestral self.
However, as Ridley (2003: 7) notes, ‘differ-
ence is the shadow of similarity’. Thus, some
of the most promising avenues for current
research on the evolutionary psychology of
war deal with puzzles such as reconciling the
universal presence of the psychological tools
of violence with variation in their expression
across space and time, explaining why the
preponderance of coalitional violence is still
carried out by men, explaining why humans
can sometimes be led to levels of violent
self-sacrifice that appear to contradict the
possibility of any fitness benefits, and cor-
recting past blind spots in research that has
focused on Western subjects and data. All of
these puzzles are tractable and indeed benefit
from and require an evolutionary lens as we
untangle the sordid knot of human coalitional
violence and warfare.
Note
1  This is an admittedly brief summary of a master-
ful and complex summary provided by Levy and
Thompson. Interested readers are encouraged to
engage the reading directly, while for my pur-
poses here, a bird’s-eye view is sufficient.
REFERENCES
Allen, Mark W., and Terry L. Jones, Eds. 2014.
Violence and Warfare among Hunter-­
Gatherers. Walnut Creek, California: Routledge.
Anderson, Rindy C., and Casey A. Klofstad.
2012. ‘Preference for Leaders with Masculine
Voices Holds in the Case of Feminine Leadership
Roles’. PLOS ONE 7 (12): e51216. https://doi.
org/10.1371/journal.pone.0051216.
Bauer, Michal, Christopher Blattman, Julie
Chytilová, Joseph Henrich, Edward Miguel,
and Tamar Mitts. 2016. ‘Can War Foster
Cooperation?’ Journal of Economic
Perspectives 30 (3): 249–74. https://doi.
org/10.1257/jep.30.3.249.
Biddle, Stephen. 2001. ‘Rebuilding the
Foundations of Offense-Defense Theory’.
Journal of Politics 63 (3): 741–4. https://doi.
org/10.1111/0022-3816.00086.
———. 2006. Military Power: Explaining Victory
and Defeat in Modern Battle. Princeton, New
Jersey: Princeton University Press.
Bird, Douglas W., Rebecca Bliege Bird, Brian F.
Codding, and David W. Zeanah. 2019.
‘Variability in the Organization and Size of
Hunter-Gatherer Groups: Foragers Do Not
Live in Small-Scale Societies’. Journal of
Human Evolution 131 (June): 96–108. https://
doi.org/10.1016/j.jhevol.2019.03.005.
Boehm, Christopher. 1999. Hierarchy in the
Forest: The Evolution of Egalitarian Behavior.
Cambridge, Massachusetts: Harvard
University Press.
———. 2012. ‘Ancestral Hierarchy and
Conflict’. Science 336 (6083): 844–7. https://
doi.org/10.1126/science.1219961.
Böhm, Robert, Hannes Rusch, and Özgür Gürerk.
2016. ‘What Makes People Go to War?
Defensive Intentions Motivate Retaliatory and
Preemptive Intergroup Aggression’. Evolution
and Human Behavior 37 (1): 29–34. https://doi.
org/10.1016/j.evolhumbehav.2015.06.005.
Boot, Max. 2013. ‘The Evolution of Irregular
War’. Foreign Affairs, March 1, 2013. www.
foreignaffairs.com/articles/2013-02-05/
evolution-irregular-war
Bowles, Samuel. 2009. ‘Did Warfare Among
Ancestral Hunter-Gatherers Affect the
Evolution of Human Social Behaviors?’
Science 324 (5932): 1293–8. https://doi.
org/10.1126/science.1168112.
 EVOLUTIONARY PSYCHOLOGY AND WARFARE
Boyer, Pascal, and Michael Bang Petersen.
2011. ‘The Naturalness of (Many) Social
Institutions: Evolved Cognition as Their
Foundation’. Journal of Institutional
Economics FirstView: 1–25.
Buckner, William, and Luke Glowacki. 2019.
‘Reasons to Strike First’. Behavioral and Brain
Sciences 42, 17–18. https://doi.org/10.1017/
s0140525x19000840.
Chang, Linda W., Amy R. Krosch, and Mina
Cikara. 2016. ‘Effects of Intergroup Threat
on Mind, Brain, and Behavior’. Current
Opinion in Psychology, Intergroup relations,
11 (October): 69–73. https://doi.org/10.1016/
j.copsyc.2016.06.004.
Chapin, Kenneth James, Paulo Enrique Cardoso
Peixoto, and Mark Briffa. 2019. ‘Further
Mismeasures of Animal Contests: A New
Framework for Assessment Strategies’.
Behavioral Ecology 30 (5): 1177–85. https://
doi.org/10.1093/beheco/arz081.
Chapman, Colin, A. 1986. ‘Ecological
Constraints on Group-Size for 3 Species of
Neotropical Primates’. American Journal of
Primatology 10 (4): 394.
Chapman, Colin A., and Lauren J. Chapman.
2000. ‘Constraints on Group Size in Red
Colobus and Red-Tailed Guenons: Examining
the Generality of the Ecological Constraints
Model’. International Journal of Primatology
21 (4): 565–85.
Chapman, C. A., R. W. Wrangham, and L. J.
Chapman. 1995. ‘Ecological Constraints on
Group-Size – an Analysis of Spider Monkey
and Chimpanzee Subgroups’. Behavioral
Ecology and Sociobiology 36 (1): 59–70.
Choi, Jung-Kyoo, and Samuel Bowles. 2007.
‘The Coevolution of Parochial Altruism and
War’. Science 318 (5850): 636–40. https://
doi.org/10.1126/science.1144237.
Crofoot, Margaret Chatham, and Ian C. Gilby.
2012. ‘Cheating Monkeys Undermine Group
Strength in Enemy Territory’. Proceedings of the
National Academy of Sciences 109 (2): 501–5.
https://doi.org/10.1073/pnas.1115937109.
Daly, Martin, and Margo Wilson. 1983. Sex,
Evolution, and Behavior. Boston,
Massachusetts: Willard Grant Press.
———. 1988. Homicide. New York: A. de
Gruyter.
De Dreu, Carsten K. W., and Jörg Gross. 2018.
‘Revisiting the Form and Function of Conflict:
327
Neurobiological, Psychological, and Cultural
Mechanisms for Attack and Defense within
and between Groups’. The Behavioral and
Brain Sciences 42: e116. https://doi.org/
10.1017/S0140525X18002170.
Doğan, Gönül, Luke Glowacki, and Hannes
Rusch. 2018. ‘Spoils Division Rules Shape
Aggression between Natural Groups’. Nature
Human Behaviour 2 (5): 322–6. https://doi.
org/10.1038/s41562-018-0338-z.
Enquist, Magnus, and Olof Leimar. 1990. ‘The
Evolution of Fatal Fighting’. Animal Behaviour
39 (1): 1–9. https://doi.org/10.1016/S0003-
3472(05)80721-3.
Fair, C. Christine, and Bryan Shepherd. 2006.
‘Who Supports Terrorism? Evidence from
Fourteen Muslim Countries’. Studies in
Conflict & Terrorism 29 (1): 51–74. https://
doi.org/10.1080/10576100500351318.
Ferrill, Arther. 1997. The Origins Of War: From
The Stone Age To Alexander The Great,
Revised Edition. 1st edition. Boulder,
Colorado: Westview Press.
Fry, Douglas P. 2007. Beyond War: The Human
Potential for Peace. 1st edition. New York:
NY: Oxford University Press.
Fry, Douglas P., and Patrik Söderberg. 2013.
‘Lethal Aggression in Mobile Forager Bands
and Implications for the Origins of War’.
Science 341 (6143): 270–3. https://doi.
org/10.1126/science.1235675.
Gabriel, Richard A. 2002. The Great Armies of
Antiquity: Westport, Connecticut: Praeger.
Gat, Azar. 2006. War in Human Civlization.
Oxford: Oxford University Press. www.loc.
gov/catdir/toc/ecip0614/2006017223.html
Gavrilets, Sergey, and Laura Fortunato. 2014.
‘A Solution to the Collective Action Problem
in Between-Group Conflict with Within-
Group Inequality’. Nature Communications 5
(March): 3526. https://doi.org/10.1038/
ncomms4526.
Ginges, Jeremy, and Scott Atran. 2011. ‘War as
a Moral Imperative (Not Just Practical Politics
by Other Means)’. Proceedings of the Royal
Society B: Biological Sciences 278 (1720):
2930–8. https://doi.org/10.1098/rspb.
2010.2384.
Glowacki, Luke, and Christopher von Rueden.
2015. ‘Leadership Solves Collective Action
Problems in Small-Scale Societies’.
Philosophical Transactions of the Royal
328 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Society B: Biological Sciences 370 (1683).
https://doi.org/10.1098/rstb.2015.0010.
Glowacki, Luke, Michael L. Wilson, and Richard
W. Wrangham. 2017. ‘The Evolutionary
Anthropology of War’. Journal of Economic
Behavior and Organization, January. https://
doi.org/10.1016/j.jebo.2017.09.014.
Glowacki, Luke, and Richard W. Wrangham.
2013. ‘The Role of Rewards in Motivating
Participation in Simple Warfare’. Human
Nature 24 (4): 444–60. https://doi.org/
10.1007/s12110-013-9178-8.
Glowacki, Luke, and Richard Wrangham. 2015.
‘Warfare and Reproductive Success in a Tribal
Population’. Proceedings of the National
Academy of Sciences 112 (2): 348–53.
https://doi.org/10.1073/pnas.1412287112.
Gneezy, Ayelet, and Daniel M. T. Fessler. 2011.
‘Conflict, Sticks and Carrots: War Increases
Prosocial Punishments and Rewards’.
Proceedings of the Royal Society B: Biological
Sciences, 279, 219–223. June. https://doi.
org/10.1098/rspb.2011.0805.
Goldstein, Joshua S. 2003. War and Gender:
How Gender Shapes the War System and Vice
Versa. Cambridge: Cambridge University Press.
Hill, Alex, Drew H. Bailey, and David A. Puts.
2017. ‘Gorillas in Our Midst? Human Sexual
Dimorphism and Contest Competition in
Men’. In Michel Tibayrenc and Francisco J.
Ayala (Eds) On Human Nature: Biology,
Psychology, Ethics, Politics, and Religion,
235–49. Academic Press.
Hill, Kim R., Robert S. Walker, Miran Božičević,
James Eder, Thomas Headland, Barry
Hewlett, A. Magdalena Hurtado, Frank
Marlowe, Polly Wiessner, and Brian Wood.
2011. ‘Co-Residence Patterns in Hunter-
Gatherer Societies Show Unique Human
Social Structure’. Science 331 (6022): 1286–9.
https://doi.org/10.1126/science.1199071.
Hoffman, Frank. 2007. Conflict in the 21st
Century: The Rise of Hybrid Wars. Arlington,
Virginia: Potomac Institute for Policy Studies.
Hooper, Paul L., Hillard S. Kaplan, and James L.
Boone. 2010. ‘A Theory of Leadership in
Human Cooperative Groups’. Journal of
Theoretical Biology 265 (4): 633–46. https://
doi.org/10.1016/j.jtbi.2010.05.034.
Jervis, Robert. 1978. ‘Cooperation under the
Security Dilemma’. World Politics 30 (2):
167–214. https://doi.org/10.2307/2009958.
Jordan, Matthew R., Jillian J. Jordan, and David G.
Rand. 2017. ‘No Unique Effect of Intergroup
Competition on Cooperation: Non-Competitive
Thresholds Are as Effective as Competitions
between Groups for Increasing Human Coop-
erative Behavior’. Evolution and Human Behav-
ior 38 (1): 102–8. https://doi.org/10.1016/
j.evolhumbehav.2016.07.005.
Keeley, Lawrence H. 1996. War before Civiliza-
tion: The Myth of the Peaceful Savage. New
York: Oxford University Press. www.loc.gov/
catdir/enhancements/fy0638/94008998-d.html
Kelly, Raymond C. 2000. Warless Societies and
the Origin of War. Ann Arbor, Michigan:
University of Michigan Press.
Kitchen, Dawn M., and Jacinta C. Beehner. 2007.
‘Factors Affecting Individual Participation in
Group-Level Aggression among Non-Human
Primates’. Behaviour 144 (12): 1551–81.
Klofstad, Casey A., Rindy C. Anderson, and
Susan Peters. 2012. ‘Sounds like a Winner:
Voice Pitch Influences Perception of Leadership
Capacity in Both Men and Women’. Proceed-
ings of the Royal Society of London B: Biological
Sciences 279 (1738): 2698–2704. https://doi.
org/10.1098/rspb.2012.0311.
Lahr, Marta. Mirazón., Rivera, Frances., Power,
R. K., Mounier, Aurelien., Copsey, B.,
Crivellaro, F., Edung, J.E., Maillo Fernandez,
J.M.,… Foley RA. 2016. ‘Inter-Group Violence
among Early Holocene Hunter-Gatherers of
West Turkana, Kenya’. Nature 529 (7586):
394–8. https://doi.org/10.1038/nature16477.
Laustsen, Lasse, and Michael Bang Petersen.
2017. ‘Perceived Conflict and Leader
Dominance: Individual and Contextual Factors
Behind Preferences for Dominant Leaders’.
Political Psychology 38 (6): 1083–1101.
https://doi.org/10.1111/pops.12403.
Levy, Jack S., and William R. Thompson. 2011.
The Arc of War: Origins, Escalation, and
Transformation. Chicago, Illinois: University
Of Chicago Press.
Lopez, Anthony C. 2016a. ‘Conditions Required
for Evolution of Warfare Adaptations’. In
Encyclopedia of Evolutionary Psychological
Science, edited by Viviana Weekes-
Shackelford, Todd K. Shackelford, and
Viviana A. Weekes-Shackelford, 1–10.
Cham: Springer International Publishing.
https://doi.org/10.1007/978-3-319-16999-
6_914-1.
 EVOLUTIONARY PSYCHOLOGY AND WARFARE
———. 2016b. ‘The Evolution of War: Theory
and Controversy’. International Theory 8 (1):
97–139. https://doi.org/10.1017/S17529719
15000184.
———. 2017. ‘The Evolutionary Psychology of
War: Offense and Defense in the Adapted
Mind’. Evolutionary Psychology 15 (4):
1474704917742720. https://doi.org/10.1177/
1474704917742720.
———. 2019a. ‘The Evolution of War’. In Carol
Ann Ireland, Jane Louise Ireland, Michael
Lewis, and Anthony C. Lopez (Eds)
International Handbook of Collective Violence,
edited by Carol Ann Ireland, Jane Louise
Ireland, Michael Lewis, and Anthony C. Lopez.
3–16. New York, NY: Routledge Press.
———. 2019b. ‘Making “My” Problem “Our”
Problem: Warfare as Collective Action, and the
Role of Leader Manipulation’. The Leadership
Quarterly, 31(2) May. https://doi.org/10.1016/
j.leaqua.2019.05.001.
Lopez, Anthony C., and Dominic D. P. Johnson.
2017. ‘The Determinants of War in
International Relations’. Journal of Economic
Behavior & Organization, October. https://doi.
org/10.1016/j.jebo.2017.09.010.
Lynch, Robert, Virpi Lummaa, and John Loehr.
2019. ‘Self Sacrifice and Kin Psychology in
War: Threats to Family Predict Decisions to
Volunteer for a Women’s Paramilitary Organi-
zation’. Evolution and Human Behavior. 40 (6)
543–50. June. https://doi.org/10.1016/
j.evolhumbehav.2019.06.001.
McDonald, Melissa M., Carlos David Navarrete,
and Mark Van Vugt. 2012. ‘Evolution and the
Psychology of Intergroup Conflict: The Male
Warrior Hypothesis’. Philosophical Transactions
of the Royal Society B: Biological Sciences 367
(1589): 670–9. https://doi.org/10.1098/
rstb.2011.0301.
Mirville, Melanie O., Amanda R. Ridley, Jean Pierre
Mucyo Samedi. Samedi, Veronica Vecellio, Felix
Ndagijimana, Tara S. Stoinski, and Cyril C. Gru-
eter. 2018. ‘Low Familiarity and Similar “Group
Strength” between Opponents Increase the
Intensity of Intergroup Interactions in Mountain
Gorillas (Gorilla Beringei Beringei)’. Behavioral
Ecology and Sociobiology 72 (11): 178. https://
doi.org/10.1007/s00265-018-2592-5.
Osgood, Robert E. 1957. Limited War: The
Challenge to American Strategy. 5th edition.
Chicago, Illinois: University of Chicago Press.
329
Otterbein, Keith F. 1997. ‘The Origins of War’.
Critical Review 11 (2): 251–77.
———. 2004. How War Began. 1st edition.
Texas, US: TAMU Press.
Parker, Geoff. 1974. ‘Assessment Strategy and
the Evolution of Fighting Behaviour’. Journal
of Theoretical Biology 47 (1): 223–43.
Patton, John Q. 2005. ‘Meat Sharing for
Coalitional Support’. Evolution and Human
Behavior 26 (2): 137–57. https://doi.org/
10.1016/j.evolhumbehav.2004.08.008.
Petersen, Michael Bang. 2012. ‘Social Welfare as
Small-Scale Help: Evolutionary Psychology and
the Deservingness Heuristic’. American Journal
of Political Science 56 (1): 1–16. https://doi.
org/10.1111/j.1540-5907.2011.00545.x.
Pietraszewski, David. 2016. ‘How the Mind Sees
Coalitional and Group Conflict: The
Evolutionary Invariances of n-Person Conflict
Dynamics’. Evolution and Human Behavior 37
(6).470–80. https://doi.org/10.1016/j.
evolhumbehav.2016.04.006.
Pinker, Steven. 2011. The Better Angels of Our
Nature: Why Violence Has Declined.
Johannesburg, SA: Viking Adult.
Puts, David. 2016. ‘Human Sexual Selection’.
Current Opinion in Psychology 7 (February):
28–32. https://doi.org/10.1016/j.copsyc.
2015.07.011.
Puurtinen, Mikael, and Tapio Mappes. 2009.
‘Between-Group Competition and Human
Cooperation’. Proceedings of the Royal Society
B: Biological Sciences 276 (1655): 355–60.
https://doi.org/10.1098/rspb.2008.1060.
Ridley, Matt. 2003. The Agile Gene: How
Nature Turns on Nurture. Reprint edition.
New York: Harper Perennial.
Romano, Angelo, Daniel Balliet, Toshio
Yamagishi, and James H. Liu. 2017. ‘Parochial
Trust and Cooperation across 17 Societies’.
Proceedings of the National Academy of
Sciences 114 (48): 12702–7. https://doi.
org/10.1073/pnas.1712921114.
Rueden, Chris von. 2020. ‘Making and
Unmaking Egalitarianism in Small-Scale
Human Societies’. Current Opinion in
Psychology, Power, Status and Hierarchy, 33
(June): 167–71. https://doi.org/10.1016/j.
copsyc.2019.07.037.
Rusch, Hannes. 2013. ‘Asymmetries in Altruistic
Behavior during Violent Intergroup Conflict’.
Evolutionary Psychology: An International
330 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Journal of Evolutionary Approaches to
Psychology and Behavior 11 (5): 973–93.
———. 2014. ‘The Evolutionary Interplay of
Intergroup Conflict and Altruism in Humans:
A Review of Parochial Altruism Theory and
Prospects for Its Extension’. Proceedings.
Biological Sciences 281 (1794): 20141539.
https://doi.org/10.1098/rspb.2014.1539.
Scalise Sugiyama, Michelle. 2014. ‘Fitness Costs
of Warfare for Women’. Human Nature 25
(4): 476–95. https://doi.org/10.1007/s12110-
014-9216-1.
Sell, Aaron N. 2011. ‘The Recalibrational Theory
and Violent Anger’. Aggression and Violent
Behavior 16 (5): 381–9. https://doi.org/
10.1016/j.avb.2011.04.013.
Smith, Kevin B., Christopher W. Larimer,
Levente Littvay, and John R. Hibbing. 2007.
‘Evolutionary Theory and Political
Leadership: Why Certain People Do Not
Trust Decision Makers’. The Journal of
Politics 69 (2): 285–99. https://doi.
org/10.1111/j.1468-2508.2007.00532.x.
Spisak, Brian R. 2012. ‘The General Age of
Leadership: Older-Looking Presidential
Candidates Win Elections during War’. PLoS
ONE 7 (5): e36945. https://doi.org/10.1371/
journal.pone.0036945.
Spisak, Brian R., Astrid C. Homan, Allen Grabo,
and Mark Van Vugt. 2012. ‘Facing the
Situation: Testing a Biosocial Contingency
Model of Leadership in Intergroup Relations
Using Masculine and Feminine Faces’. The
Leadership Quarterly 23 (2): 273–80.
Stoker, Donald. 2016. ‘Everything You Think
You Know About Limited War Is Wrong’.
War on the Rocks, December 22, 2016.
h t t p s : / / w a ro n t h e ro c k s . c o m / 2 0 1 6 / 1 2 /
everything-you-think-you-know-about-
limited-war-is-wrong/.
Tilly, Charles. 1975. The Formation of National
States in Western Europe. Princeton: Princeton
University Press.
Tooby, John, and Leda Cosmides. 1988. ‘The
Evolution of War and Its Cognitive
Foundations’. Institute for Evolutionary
Studies Technical Report 88–1. https://www.
cep.ucsb.edu/papers/EvolutionofWar.pdf
Tooby, John, Leda Cosmides, and Michael E.
Price. 2006. ‘Cognitive Adaptations for
N-Person Exchange: The Evolutionary Roots of
Organizational Behavior’. Managerial and
Decision Economics 27 (2–3): 103–29.
Trivers, Robert. 1972. ‘Parental Investment and
Sexual Selection’. In Bernard Campbell (Eds)
Sexual Selection and the Descent of Man.
Chicago, IL: Aldine Publishing.
van Vugt, Mark. 2009. ‘Sex Differences in
Intergroup Competition, Aggression, and
Warfare: The Male Warrior Hypothesis’.
Annals of the New York Academy of Sciences
1167 (June): 124–34. https://doi.org/10.1111/
j.1749-6632.2009.04539.x.
van Vugt, Mark, and Rob Kurzban. 2007. ‘Cogni-
tive and Social Adaptations for Leadership and
Followership: Evolutionary Game Theory and
Group Dynamics’.” In Joseph P. Forgas, Martie
G. Haselton, and William von Hippel (Eds)
Evolution and the Social Mind: Evolutionary
Psychology and Social Cognition, 229–43.
New York: Psychology Press.
Weisel, Ori, and Robert Böhm. 2015. ‘“Ingroup
Love” and “Outgroup Hate” in Intergroup
Conflict between Natural Groups’. Journal of
Experimental Social Psychology 60
(September): 110–20. https://doi.org/
10.1016/j.jesp.2015.04.008.
Wiessner, Polly. 2019. ‘Collective Action for
War and for Peace: A Case Study among the
Enga of Papua New Guinea’. Current
Anthropology 60 (2): 224–44. https://doi.
org/10.1086/702414.
Wilson, Michael L., Christophe Boesch, Barbara
Fruth, Takeshi Furuichi, Ian C. Gilby, Chie
Hashimoto, Catherine L. Hobaiter, et al. 2014.
‘Lethal Aggression in Pan Is Better Explained
by Adaptive Strategies than Human Impacts’.
Nature 513 (7518): 414–17. https://doi.
org/10.1038/nature13727.
Wrangham, Richard. 1999. ‘Evolution of
Coalitionary Killing’. Yearbook of Physical
Anthropology 42: 1–30.
———. 2019. The Goodness Paradox: The
Strange Relationship Between Virtue and
Violence in Human Evolution. 1st edition.
New York: Pantheon.
Wrangham, Richard W., and Luke Glowacki.
2012. ‘Intergroup Aggression in Chimpanzees
and War in Nomadic Hunter-Gatherers:
Evaluating the Chimpanzee Model’. Human
Nature. 23 (1): 5–29 March. https://doi.org/
10.1007/s12110-012-9132-1.
 PART 3

Applications to Technology,
Communications, and the Future
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Evolutionary Psychology and
Artificial Intelligence: The Impact
of Artificial Intelligence on Human
Behaviour
Holly Wilson, Paul Rauwolf, and Joanna J. Bryson
1 INTRODUCTION
Artificial intelligence (AI) impacts our
behaviour. Intelligence, whilst not in itself
defining humanity, is one of our key charac-
teristics, inextricably linked to everything
from our implicit survival strategies to our
explicit self-concepts. Intelligence is also
integral to the social institutions on which
contemporary existence depends. AI is a set
of technologies that extend human intellec-
tual capacities such as perception, action,
categorisation and pattern recognition. Some
systems, termed autonomous, may do all of
these things at once without human interven-
tion, though only after human or human-
institutional inception.
Determining the impact of AI is impera-
tive for two reasons: for empowering us to
adapt to and optimise our existence with AI;
and for developing AI and AI policies which
maximise benefit and minimise harm to our
society. Due to the myriad definitions of AI,
we begin by establishing what we mean by
17
the term, at least in the scope of this chapter.
Intelligence is the capacity to do the right
thing at the right time; thus artificial intel-
ligence refers to non-living artefacts that
demonstrate such capacities (Bryson, 2019).
By this definition, AI has been prevalent for
decades, albeit less advanced than at present,
and not so apparent to the public eye. Present
awareness of AI has been magnified by two
different recent outcomes: first the sudden
prevalence of anthropomorphic capacities
such as ­conversational-speech recognition and
generation, or automobile driving; and second
the use of AI technology as part of a global
assault on democracies, and through them key
institutions to maintaining peace under the
present global order, such as the EU and the
North Atlantic Treaty Organization (NATO).1
We focus here on how behaviour is shaped
by contemporary intelligent technology; first
by our cultural understanding of AI, then
by the technological reality of AI. Many of
our drives and behaviours, such as tribalism,
sex, and resource procurement, are sculpted
334 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
by thousands of years of evolution in dispa-
rate environments (Buss and Schmitt, 1993;
Kramer and Ellison, 2010; McDonald et al.,
2012). Therefore, in this chapter we explore
from an evolutionary perspective how AI –
ubiquitous in the modern world – impacts
both individual and collective human behav-
iour. We put emphasis on the predictions
from several published models that explain
how information transmission facilitates
intelligence between intelligent agents. We
consider how these theories predict changes
to individual and collective behaviour as the
information transmission is magnified or its
quality improved. We then compare, at least
qualitatively, these predicted alterations to
present societal trends. We finish with recom-
mendations for guiding AI development and
interaction to maximise adaptation, progres-
sion and overall benefit to the individual and
society.
More specifically, in Section 2 we initially
tease apart the current actual impact of AI
on society from the impact that our cultural
narratives surrounding AI have. We consider
the evolutionary mechanisms that maintain
a stable society such as heterogeneity, flex-
ibility and cooperation. Given that AI can
constitute a prosthetic intelligence, we dis-
cuss the consequences of how it enhances
our connectivity, coordination, equality, dis-
tribution of control and our ability to make
predictions. We also give examples of how
transparency of thoughts and behaviours may
influence call-out culture, as well as behav-
ioural manipulation with consideration of
group dynamics and tribalism. In Section 3,
we discuss how AI may exacerbate the soci-
etal problems created by inequality. We place
focus on the vulnerability of human trust in
Section 4. We consider the contexts in which
blind trust in information is either adaptive
or maladaptive in an age where the cost of
information is decreasing. We then bring the
focus back onto trust in AI, and how we can
calibrate trust so as to avoid over-trust and
mistrust adaptively, using transparency as a
mechanism. Finally, in Section 5, we explore
the barriers to AI increasing accuracy in our
perception by focusing on fake news. We con-
sider the impact of information accuracy and
the battles of individuals against false beliefs.
2 IMPACTS OF AI THUS FAR
The narratives within a culture can have as
much impact on behaviour as at least some
objective realities (Hammack, 2008). For this
reason, we begin by examining the current
narrative surrounding AI. According to Social
Representation Theory (SRT), when we
encounter a new or unknown phenomenon,
we construct a representation of it based on
collective narratives and interpersonal com-
munication (Moscovici, 1981, 2001). It seems
clear that public gaps in understanding AI are
often filled by fear-mongering entertainment
shows like Black Mirror, magazine articles on
the feats of Alpha Go, and propaganda from
businesses (Elish and Boyd, 2018). Indeed,
media exposure to science fiction has been
found to predict fear of AI above and beyond
demographic variables (Liang and Lee, 2017).
Problematically, such representations
assume AI with capacities far beyond the cur-
rent feasibility – and in many cases, beyond
the computationally tractable – instilling awe
and fear (Bryson and Kime, 2011). An inves-
tigation of narratives surrounding the impact
of AI revealed that the most common visions
elicited anxiety (Cave et al., 2019). One such
vision was that by becoming over-reliant on
AI and machines, we will replace the need
for humans in jobs, relationships and social-
ising. Emotional arousal increases the effi-
cacy of information spread (Berger, 2011);
this mechanism is posited to have evolved
to transmit fitness-relevant information; i.e.
information relevant to survival which can
help organisms avoid dangers (Nairne et al.,
2009) or direct resources within a population
(Teste et al., 2009). Yet the adaptive benefits
of this in contemporary cultures are question-
able, now that the mechanism is known and
 EVOLUTIONARY PSYCHOLOGY AND ARTIFICIAL INTELLIGENCE
manipulable. These narratives may distract
from or obscure the real problems and util-
ity of AI, resulting in sub-optimal allocation
of energy and resources (Bryson and Kime,
2011; Elish and Boyd, 2018). In the next few
sections, we discuss the mixture of benefits
and potential problems we face with AI.
2.1 Flexibility, Cooperation,
Coordination, Perception:
Humanity’s Survival Mechanisms
AI does present a new landscape for
humanity – yet, despite popular rhetoric, this
does not necessarily pose an existential
threat (Gent, 2015; Müller and Bostrom,
2016). Throughout our evolutionary history,
humans have succeeded in adapting to
changing environments (Gilligan, 2007;
Richerson et  al., 2005). Machines, in con-
trast, are typically fragile and short lived. As
such, dangerous machines or technology are
unlikely to be allowed to persist in their
damaging behaviour long enough to destroy
humanity as a whole, although technologi-
cally mediated impacts such as climate
change or hate crimes are already costing
lives. Cooperation and flexibility by means
of heterogeneity (diversity) are two mecha-
nisms that enable adaptation, survival and
progress in unstable, changing environments
(Brown et  al., 2011; Lahr, 2016; Smaldino
et al., 2013). Here we discuss the impact of
AI on human behaviour within the context of
these two mechanisms.
There are two core hypotheses as to the
role our nervous systems evolved to fill: the
sensory-motor view, to link senses to actions;
and the action-shaping view, to coordinate the
body’s micro acts into macro acts (Godfrey-
Smith, 2017). These hypotheses are by no
means exclusive of each other. Likewise, our
development of AI, amongst other things,
has greatly enhanced our abilities to sense,
act and coordinate. Our society has become
increasingly complex; we are connected
world-wide, with perturbations in one part
335
of our global system impacting many others.
AI can be considered as our prosthetic nervous
system, a tool we have developed to selectively
mediate the strength of edges between each
node. As an example, we can deploy hardy
robots to explore subterranean and underwater
environments inaccessible to humans, acting
and perceiving on our behalf (Siles and Walker,
2009), growing new edges of our agency.
Machine learning (ML), the ability to learn
to perceive and categorise patterns based on
input data, also constitutes a prosthetic percep-
tion. With adequate computational resources,
ML for example allows us to search across
larger ranges of data than a human might oth-
erwise be able to internalise, and to consider
more candidate patterns. Enhanced perception,
assuming accuracy, means a species has more
knowledge with which to react better to events
in its environment. This constitutes an increase
in human collective intelligence (Eagle and
Pentland, 2003).
2.2 AI Increases Connectivity
Which Facilitates Coordination
but Also Transparency
An enhanced capacity for coordination also
results from the increased connectivity that
AI and Information and Communication
Technologies (ICT) more generally facilitate.
These technologies afford communication
and coordination on a scale that human soci-
eties have never encountered before. This can
be expected to have – and be having – myriad
impacts on collective and individual behav-
iour, not all of which we have yet recognised
(Bryson, 2015). Through increasing the
number of individuals we can connect with,
and decreasing temporal and spatial con-
straints on doing so, AI creates a capacity for
highly agile cooperation. Cooperation and
group-level investment as a whole is known
to increase with capacity to communicate,
because this capacity allows for the increased
probability of discovering mutually benefi-
cial equilibria (Roughgarden et  al., 2006).
336 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
This cooperation may occur at historically
large scales, but also at small scales, and with
higher frequency of change in aggregation
and direction. Social media platforms, for
example, have facilitated mass organisation
of not only protests but also disaster recovery
which may not otherwise have been feasible
(Gerbaudo, 2018; Starbird and Palen, 2011;
Vieweg et al., 2010). Additionally, increased
access to knowledge enables us to predict
threats or more quickly become aware of
disasters (Chavan and Khot, 2013).
Connectivity also has the effect of increas-
ing behaviour transparency. Social media
pages reveal the ‘likes’, ‘dislikes’ and actions
of a population. Problematically within this
context, humans are often driven to take
on characteristics of a group, and strive to
behave according to group norms (Terry and
Hogg, 1996), which may have a homogenis-
ing or polarising effect, particularly during
periods when competition and identity poli-
tics are steep (McCarty et al., 2016). Before
we were so connected via the digital world,
the group norms we had access to were of far
smaller scale. This facilitated diversity – a
global heterogeneity of group norms. Now,
our access to the large scale, combined with
behaviour transparency, is widely believed
to homogenise behaviours and preferences
(Morris, 2002). This is true even without
taking into account potential conformity
induced by physical, political, or economic
threats for unacceptable behaviour, which
we discuss further below. However, the data
created by our digital activity is also made
transparent for use by the political and busi-
ness, as well as the social, realms. This can
have positive impacts as well, if it is used to,
for example, provide better public services or
ensure greater customer satisfaction.
2.3 AI Facilitates Behaviour
Prediction and Manipulation
Ordinary collectives such as companies use
algorithms on big data sets to predict our
behaviour with increasing accuracy (Zuboff,
2015). This seems qualitatively different to
our already adept prediction capacities: to
navigate our social world, we use mental
models of people to infer and predict the
beliefs, actions and intentions of others
(Bradford et al., 2015). This ability, alongside
language (Smith, 2010), has been critical for
facilitating our species’ large-scale social
cooperation. However, AI has enhanced our
ability to predict the individual and collective
beyond past capabilities. This can aid us to
better allocate our time and resources. For
example, by anticipating that the number of
people about to use a road is beyond capacity,
a navigation app may direct a proportion of
people along a different route, altering their
behaviour advantageously for the collective.
At an individual level, the ‘optimal’ level of
sleep (Hao et al., 2013), exercise (Spring et al.,
2013), socialising and nutrition (Franco et al.,
2016) can be predicted then recommended.
Unfortunately, businesses looking to maxim-
ise profits will tailor their product – or just
monetising efforts related to their product – to
the individual based on predictions. This can
be with an incentive to reshape behaviour to
make it even more easily predictable, or oth-
erwise less expensive for the company, for
example when insurance companies demand
digital access to evidence of healthy living
(Raber et al., 2019). This, whilst not explic-
itly detrimental to cooperation, may increase
homogeneity and therefore reduce group
advantages and societal robustness (Fisher,
1930; Shi et  al., 2019). It also speaks to an
additional issue with behaviour transparency:
capacity for behavioural control.
Both the aforementioned examples of
collective and individual behaviour tweak-
ing and recommendations indicate a shift in
autonomy; it seems AI may be increasing
collective agency but decreasing opportunity
and drive for individual decision-making. In
fact, a survey of 970 respondents revealed
that a core concern surrounding AI tech-
nologies is the loss of human agency and
input into decisions (Anderson et al., 2018).
 EVOLUTIONARY PSYCHOLOGY AND ARTIFICIAL INTELLIGENCE
At one level, behavioural control may or may
not shift to autonomous artificial agents, but
either way at a higher level it shifts to the
individuals or organisations who can moni-
tor data and deploy any such agents. This
enables execution of potentially regressive
social policing, albeit some well-intentioned.
Take for example, the rise of helicopter par-
enting (Lee et  al., 2014), or AI-powered
predictive policing systems (Meijer and
Wessels, 2019). We do not claim such behav-
iour manipulation is unique to AI. There
are varying views on whether manipulation
techniques (not all of which necessarily use
AI) are ethical, when used for example to
promote health (Behavioural Insights Team,
2010) or reduce debt (Behavioural Insights
Team, 2012). We see the sense in policies
(such as that of the Institute of Electrical and
Electronics Engineers (IEEE), 2019) recom-
mending that behaviour manipulation may
be ethical in contexts where all the following
hold: it can be beneficial for the individual
and/or society, transparency is provided as to
the nature of the manipulation, and the sub-
ject or a responsible adult representative of
the subject has consented. This (arguably, cf.
Simkulet, 2019) leaves the individual some
control over the decision, at least at a higher
level. For example, clinicians, especially
under cognitive load, can demonstrate bias
towards ethnic-minority patients, resulting
in sub-optimal interaction, diagnosis and
treatment (Stone and Moskowitz, 2011).
Evidence suggests that two tasks – perspec-
tive taking or categorising oneself to be in
a shared group with the ethnic minority –
can reduce bias. In this scenario, consented
behavioural manipulation could involve an
app-based intervention, where the clinician
chooses to engage with a bias-reduction task
prior to seeing the patient.
This same public or semi-private – and
sometimes implicit – communication of pref-
erences can be used deliberately to determine
the personality types of individuals, and also
their voting inclinations (Gelman et al., 2016;
Kosinski et al., 2013; Wu et al., 2015). Such
337
information has obvious applications for
those interested in the outcomes of elections,
which have apparently been deployed with
some success. Such techniques were report-
edly originally designed and deployed to
break up terrorist networks in conflict regions
in the Middle East. The technique consists
of identifying like-minded individuals with
minority opinions that are available locally
or otherwise convenient to those doing the
aggregating, then introducing these individu-
als to each other and encouraging their politi-
cal participation (Piette, 2018). AI-powered
search can produce the ‘coincidence’ of good
numbers of like-minded individuals in one
place, convincing them all that their position
is secretly in the majority – a secret being
kept by the political status quo which must
therefore be attacked. Relatively simple AI
allows the identification and coordination
of such target individuals; ICT allows the
application of such power from a distance
and across borders. Without laws and tech-
nological enforcement for transparency, such
manipulation may be done invisibly.
2.4 AI as Prosthetic Memory
In the past, humans strove to both remember
and to be remembered, yet the time and mon-
etary cost of data retention were barriers to
doing so. Now, however, these costs have
reduced to the point where we no longer need
to be picky about the quality of what we select
to retain. This has its advantages and disad-
vantages. Biological brains forget (Kraemer
and Golding, 1997). In humans, such a lack of
pruning memories, inability to forget, or
information overload can result in deficits in
executive functioning, and an inability to
escape past autobiographical memories
(Parker et  al., 2006). Using AI-driven tech-
niques, even with surplus data, we can still
successfully store, reorganise, classify and
retrieve the relevant information we require
from large data stores. However, this pros-
thetic memory of digital expression
338 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
of opinions and beliefs – whether political,
religious or otherwise – combined with
increased connectivity has resulted in,
amongst other things, a resurgence of call-out
culture and public shaming (Hess and Waller,
2014; Tucker, 2018; Webb et al., 2016).
The urge to normalise groups by exiling
or denouncing the credibility of individuals
who diverge is a facet of tribalism prevalent
throughout our history (Bechtel, 1991; Burns,
2003). Novel though with the AI landscape is
the relative immortality of digital memory. The
social costs (and benefits) to acting or thinking
in diverging ways have increased; it is hard to
be forgotten. Thus again, we are homogenised
by our evolutionary urge to remain in the
safety of the tribe. Mayer-Schönberger (2007)
advocates finding an equilibrium: by giving
data an expiry date after which it will auto-
matically be deleted, we can maximise the
benefits of a precise prosthetic memory, whilst
preserving the right to be forgotten. However,
such arbitrary truncation would also be an end
to history, unless history were still retained in
non-digital format. Even if exceptions were
made for those considered public individuals,
as is now the case for certain privacy laws, this
could have an unintended effect of reducing
social mobility, as those in situations of promi-
nence, privileged with being knowable, would
be more likely to garner further attention and
opportunity.
Historical data also has applications rang-
ing from the social sciences to monitoring
the impacts of government policies. In an
era where behaviour modification might be
practised by subterfuge, accurate historical
data may be the only way to detect mali-
cious actors working subtly over time. The
rights to freedom of opinion and thought
are enshrined in the Declaration on Human
Rights (United Nations General Assembly,
1948), but without an option of perceivable
expression such rights may be of limited
value. There is substantial research being
conducted in anonymisation, including for
data extraction and analysis – whether this
proves mathematically tractable remains to
be seen. Uncompromised cybersecurity of
not only data storage but transmission would
also be necessary for any digital records to
have even a hope of remaining private.
The claim that homogenisation is a side
effect of AI may seem demonstrably false
given the increase in identity politics and
political polarisation. Coincidence is not
necessarily causal, and even if there is a
causal link, it may be difficult to untangle.
Polarisation is known to be correlated with
wealth inequality, and to have been so since
before the advent of ICT and AI. Whether AI
is presently contributing to inequality will
be considered in the next section. But with
respect to homogenisation, it is worth say-
ing that both processes may well occur at the
same time – rather than a plethora of perspec-
tives, we may find strong forces towards con-
formity with one of a small number of tribes.
Again, this is the opposite of what was antici-
pated with access to the Internet and cheap
self-publication, and there is also evidence of
societal fragmentation (Pentland, 2015).
3 AI AND INEQUALITY
Whether or not we become more homoge-
nous in our beliefs and opinions, inequality
in access to resources and quality of life is
increasing. We discuss here what is known
about how inequality is driven, and consider
the impact of AI and the consequences for
the collective.
First, it should be observed that globally
inequality has been falling, an effect driven
primarily by the very poorest. The World
Bank reports more than a third of humanity
moving out of extreme poverty since 1980
(Roser and Ortiz-Ospina, 2017), a shift that
has been facilitated by ICT including AI, as
populations have had more access to useful
information such as weather predictions, fair
prices and how to obtain government sup-
port. Further, efforts to communicate politi-
cal and economic situations, and means to
 EVOLUTIONARY PSYCHOLOGY AND ARTIFICIAL INTELLIGENCE
coordinate protest, are leading at least some
governments in rich areas such as the mem-
ber states of the Organisation for Economic
Co-operation and Development (OECD) to
adopt policies that have been demonstrated
to reduce inequality within populations by
increasing wealth distribution.
Nevertheless, an individual’s access to pub-
lic goods such as schools, physical security,
utilities and health depends to a large degree
on their geographic identity. Access to ICT
and AI is no exception (Robinson et al., 2015;
Sujarwoto and Tampubolon, 2016), although
it is also influenced by other factors such as
age. As we globally become more dependent
on such tools, the populations without access
are exposed to higher risks of inequality. By
moving towards equality of access to these
technologies, individuals may have improved
job opportunities and information in regard
to the socioeconomic, political and cultural
context in which they live.
The increase in inequality may result in
reduced social cohesion as it seems to be cor-
related to reductions in social mobility and
increases in political polarisation (McCarty
et al., 2016). ICT and AI may also reduce the
sort of localised social cohesion that is criti-
cal to many forms of well-being and political
engagement, by diverting social attention to
others with shared interests in topics that are
not geographically centred. In this, it continues
and expands on trends of mass media known
since the beginning of the information age
with, for example, the advent of national news-
papers (Perlman and Sprick Schuster, 2016).
For a substantial fraction of our society, the
time we spend engaging with others as real,
physical equals is replaced with ever-more-
engaging digital entertainment. Research
shows that whilst some US teens felt digital
technologies connected them with others, oth-
ers felt it resulted in a lack of in-person contact
in their lives (Anderson and Jiang, 2018). The
interaction that would once have taken place in
person is now conducted through technology.
There is a danger that inequality pro-
duces an elite who no longer identify with
339
the majority of individuals (Atkinson, 2015).
This can be the effect not only of lack of
social mobility and understanding, but even
of simple spatial segregation (Cassiers and
Kesteloot, 2012). As discussed earlier, digital
social media provides a ready platform for dis-
information, including caricaturised and exag-
gerated distortions of others. An elite may also
falsely assume that it can consolidate power
by extracting wealth from its own neighbours
and nearest contenders. However, inequality
breeds instability as the whims of small coali-
tions or even individual actors are unpredict-
able (Scheidel, 2017). When greater collective
action is required, solutions become more pre-
dictable and stable, ironically better ensuring
the maintenance of rank order at the higher
end of society. Power over a collective is not
only an animal thrill, it is also a mechanism
of security because it ensures more individuals
are invested in a mutually beneficial outcome
(Terkel, 1974). Trust is a factor in individuals
collectively investing in mutually beneficial
outcomes, yet not only does inequality breed
distrust (Barone and Mocetti, 2016) but, even
when trust is held, it is vulnerable to exploita-
tion and damage to the individual.
Again, there is no clear evidence to date, but
rather active investigation, as to whether and in
what ways AI may be affecting inequality. It
seems evident that any technology that reduces
the cost of distance will also facilitate inequality
through no particular malfeasance but simply
by allowing excellent businesses to dominate
larger territories, and therefore market shares.
In the case of some technologies now such as
finance, media, pharmaceutical, aerospace, and
of course digital, this is approaching the limit
case of single corporations dominating markets
globally. This same phenomenon may explain
the similar surge in developed-world inequality
witnessed in the late 19th and early 20th centu-
ries (Atkinson, 2015; Bryson, 2019).
As mentioned in Section 2, another con-
cern is that AI may alter employment. Whilst
the common concern is that ‘robots will take
all the jobs’, this seems highly unlikely as ‘all
the jobs’ is not defined. There is an infinite
340 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
number of ways we can better each other’s
lives. People tend to employ each other when
the economy is good, though this may be
seen as tautological. But the advent of intel-
ligent technology has been associated with
an increase in demand for both highly skilled
and very low-skilled and low-waged work,
with a declining demand and therefore wages
for the intermediate workers (Acemoglu and
Autor, 2011). Acemoglu and Autor (2011) sug-
gest that technology plays two roles in wages:
in allowing all to be more productive, this
somewhat increases the value of being highly
skilled, but flattens any advantages of moder-
ate skill as people become more exchangeable.
Worryingly, recent decades seem to be domi-
nated by the latter effect.
4 THE VULNERABILITY OF
HUMAN TRUST
Subjective evaluations of trustworthiness are
deeply tied to how humans navigate the world.
The likelihood of a monetary transaction is
largely dependent on the trust a buyer has in a
seller (Kim et  al., 2008; Ponte et  al., 2015).
Trust in politicians affects voter turnout
(Grönlund and Setälä, 2007) and electoral
results (Hetherington, 1999). When evaluating
a person’s face on multiple dimensions, subjec-
tive trustworthiness is one of the most predic-
tive measures for overall evaluation (Oosterhof
and Todorov, 2008). Further, once trustworthi-
ness is perceived, it is relatively robust, persis-
tently affecting behaviour (Delgado et  al.,
2005). If AI alters our capacity or predisposi-
tion to trust other humans, it will clearly have a
deep impact on society.
4.1 Baseline Proclivities to
Blindly Trust
Using simple economic games, it has been
shown that individuals will blindly trust a stran-
ger even when it leaves them vulnerable (Berg
et al., 1995). In a one-shot Trust Game (TG), an
investor is given some monetary windfall and
must decide whether to keep the windfall, or
give some fraction to a trustee. The fraction
offered by the investor is then multiplied by
some factor (typically three) and the trustee can
then offer a fraction of the multiplied invest-
ment back to the investor. Human investors
tend to blindly trust their partner by offering
non-zero investments, even though it is in the
trustee’s best interest to return nothing to the
investor. Whilst trust is moderated by several
factors, including framing effects (Burnham
et  al., 2000), geography (Johnson and Mislin,
2011), gender (Buchan et al., 2008), risk prefer-
ences (Fehr, 2009) and whether participants are
selected from a student population (Johnson
and Mislin, 2011), individuals consistently tend
to blindly trust, investing some of their windfall
(Johnson and Mislin, 2011).
Significant research has sought to under-
stand how blindly trusting others might be
ecologically rational. This proclivity to trust
has been explained as adaptive in relatively
small populations where individuals have
reputation cues of their partners (Boero et al.,
2009; Masuda and Nakamura, 2012) or if
the chance of knowing a partner’s strategy
exceeds some threshold (Manapat and Rand,
2012; Manapat et al., 2013; McNamara et al.,
2009; Rauwolf and Bryson, 2018). The com-
mon theme is that blindly trusting another
can be adaptive if someone, somewhere, has a
chance of having information about a player,
including indirectly (e.g. if a population is
known to share trust-related characteristics
by some sort of contagion effect or enforce-
ment). Interestingly, on the other hand, indi-
viduals will not trust others when they know
the others are likely to return (reciprocate) an
unfair amount in the TG, even if that amount
would still make trusting them beneficial.
4.2 When Blind Trust Is Valuable
Rauwolf and Bryson (2018) demonstrate a
generally advantageous but unstable evolu-
tionary dynamic that typically establishes
 EVOLUTIONARY PSYCHOLOGY AND ARTIFICIAL INTELLIGENCE
trust. This demonstration is in the context of
simulations with computational agents play-
ing one-shot TGs with each other. Agents
were given several potential partners for play-
ing each game, and chose which agent to play,
if any. As in the natural world, the investing
agent knew the reputation for historic pay-off
of some partners, but did not know the pay-
off of others – information was partially
occluded. Each actor learned three things
socially from the strongest players: their
levels of trust in unknown players, the
demanded level of reciprocation for known
players, and their own reciprocation rate.
Rauwolf and Bryson (2018) demonstrate that
this simple dynamic is sufficient to generate
trust between members of a population. When
the known pay-offs of partners are sufficiently
low, it can be in an agent’s interest to blindly
select a partner whose history is unknown,
provided that the population has evolved high
enough levels of trust, which tends to coevolve
with high levels of reciprocation, and of
course assuming sufficient extra benefit from
mutual development of the public good. In
these cases, blind trust can be more valuable
to the agent than walking away with their
monetary windfall, trusting no one. On the
other hand, trust will not evolve if the recipro-
cation rate of many players is known, because
it becomes better to stick with the best-known
available return rather than to take a chance
with the unknown.
The insight from this work is that a will-
ingness to blindly trust others increases com-
petition between others, lowering prices by
increasing the reciprocation rate. It is well-
known that creating competition between
sellers lowers prices. But, by being willing
to trust those whose information is unknown,
the pressure of competition is increased. Not
only do sellers need to compete with other
sellers whose information is known, they now
need to compete with those whose informa-
tion is unknown. This tends to lower the mar-
ket price even further. This is related to work
on outside options (André and Baumard,
2011). The value of a sale is contingent upon
341
the other options of a prospective buyer. If an
individual is willing to go elsewhere, even by
blindly trusting a stranger, then the market is
forced to adjust and the buyer’s life is improved.
4.3 Information Cost Reduces
Benefits of Trust
Importantly, whilst the adaptive models
of trust require that some information is
available, trust fails if information is fully
transparent (Manapat et  al., 2013; Rauwolf
and Bryson, 2018). By definition, the act of
trusting another requires some uncertainty in
the outcome (Yamagishi, 2011). If informa-
tion is fully transparent, then there is no need
to trust another; rather, each individual can
make an informed decision. The consequence
is that there is no selective pressure to evolve
or learn the strategies and beliefs associated
with the riskier behaviour, but these are what
bind a local community together.
We are currently living in an age where the
cost of information is dramatically decreas-
ing. As a result, the adaptive benefits of trust
are becoming increasingly obsolete. This
may be for the best – we may have a more
predictable environment with even higher
rewards for ‘good’ behaviour. However, we
should also be concerned about this being
another force for homogeneity, and further
loss of individual capacity to deal creatively
with localised crises. The institution of trust
may be just a consequence of our inability to
perfectly control our peers, but it may also
serve an adaptive advantage by reducing our
responsibility to do so. Society does not need
to come up with plans to handle every con-
tingency, because a desperate individual can
always take advantage of the availability of
trust without first seeking social approval of
their plans.
There are indications that in the near
term moving from trust to full information
is problematic in other ways. Because we
will not just ‘shut off’ our historic psycho-
logical choice making, replacing trust with
342 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
information may mean that extant prejudices
become more rigidly a part of our behaviour.
Not only is trust deeply tied to how individuals
make decisions, subjective trust is often biased
and founded on unhelpful signals. People
find attractive individuals more trustworthy
(Wilson and Eckel, 2006). Individuals will
invest more if a profile picture has a smiling
face (Scharlemann et al., 2001) or is visually
perceived as more trustworthy (Bente et  al.,
2012). The trust individuals place in profile
information is often incorrect (Toma, 2010).
More generally, individuals perform close to
chance when predicting deception (Bond and
DePaulo, 2006). Ert et  al. (2016) show that
perceived trustworthiness of an Airbnb option
correlates more with the profile photo than the
quantified reputation score of that option. This
was confirmed by Fagerstrøm et  al. (2017),
who found that facial expressions in a renter’s
photo predicted likelihood to rent more than
customer ratings. This demonstrates that the
transparency of information is not necessarily
sufficient to improve behaviour. Individuals
must make decisions using that information
before it offers an advantage.
4.4 Calibrating Trust in AI
There is considerable discussion these days
about trust in AI and trustworthiness for AI.
Our own work and that of many in the UK’s
ethics community more generally has taken a
different tack, emphasising that trust is an
anthropocentric trait not truly useful for arte-
facts, where transparency and accountability
are more desirable (Boden et  al., 2011;
Bryson and Theodorou, 2019). Improving
transparency in AI reduces the need to trust
AI. Yet it is possible that transparency does
not affect the consumption of AI when the
human consumer projects human-like iden-
tity to intelligent technology (anthropomor-
phises). By doing so, the consumer exposes
themselves to exploitation as their established
biases concerning the likelihood of trustwor-
thiness are even easier to exploit via designed
artefacts than they are by unscrupulous
­individuals. For example, one might assume
that a robot is unlikely to remember every-
thing you say because a person or pet would
not, but the robot may in fact not only recall
but transmit its full memory – a full record of
all interactions or even nearby events – storing
these offsite in a corporate cloud. Although
in theory the same digital and architected fea-
tures of AI that make it more powerful as a
manipulator should also make it easier to
govern, presently (2020) manipulation is out-
stripping governance.
Acceptance of AI can be increased in many
ways, but given the vulnerability of the human
trust system, care is needed to ensure trust is
extended with consent, and is not exploited
(IEEE, 2019). Whilst tapping into the vulner-
abilities of how humanity perceives trustwor-
thiness may be efficacious, it can also result
in unwarranted trust. People are already found
to perceive AI as more objective than human
­decision-makers, and in some cases to over-rely
on AI. For example, in a legal setting, people
demonstrated a preference to follow a machine
advisor’s decision despite a human advisor
having judgement of higher accuracy (Logg
et al., 2019). In our own research (Wilson and
Theodorou, 2019), we have found that in vir-
tual reality (VR), AI actors presented to the
participants as human agents were perceived
to be significantly less deterministic than those
presented explicitly as robots, despite both
being controlled by the same AI system.
It seems that many of us attribute properties
to AI that do not exist, at least where that AI
reminds us of humans (Sparrow, 2019). Some
evidence suggests that anthropomorphising
robots increases interaction with them (Waytz
et al., 2014) via increased trust resistance (de
Visser et al., 2016) and mind attribution. There
are uncertainties as to the impact of anthropo-
morphism when robots are more prevalent and
‘normalised’ in our society. As we grow famil-
iar with robots in our day-to-day lives, our
mental models of robots may become more
accurate (Bryson and Kime, 2011). We may
perceive with more clarity the distinctions
 EVOLUTIONARY PSYCHOLOGY AND ARTIFICIAL INTELLIGENCE
between artificially embodied cognition and
humans, or commercial products may be
mandated to provide transparency. In these
cases, the impact of anthropomorphism may
be reduced. Alternatively, viewing robots as
human-like may become normalised, and
social robotics – believed to be human-like,
despite their inhuman, designed capacities –
an embedded aspect of our lives.
We suggest that a safer and more long-term
stable approach is to work to increase AI trans-
parency whilst simultaneously helping indi-
viduals learn to make choices using empirical
information. As the information age reduces
the need for trust, individuals need to be trained
to operate in this information age, rather than
reinforcing poor decision-making tendencies
based on fallible and manipulable perceptions
of trust. An example of increased AI transpar-
ency would be to have a QR code attached to
each robot that when scanned gives informa-
tion on the robot’s maker, purpose and capabili-
ties. We have also been developing means for
allowing users to see the current goals and strat-
egies of a robot or other real-time interactive AI
system (Rotsidis et al., 2019; Theodorou et al.,
2017), and are presently experimenting to see
whether this reduces the moral-hazard aspects
of anthropomorphism. Whilst some may see
viewing AI as human-like to be an example of a
freedom of opinion or even association, we feel
strongly that such opinions need to be informed
where information is available, in order to
avoid unknowing exploitation. Willing exploi-
tation, like the manipulation of emotions that
occurs during a motion picture or other work of
fiction, is of course a perfectly acceptable part
of life and entertainment. We only seek to avoid
economic and political manipulation imposed
on unknowing others.
5 BARRIERS TO ACCURATE
PERCEPTION AND DEVELOPMENTS
Our earlier metaphor for AI as an extension
of our nervous system posited that new
343
AI tendrils enhance our ability to sense and
perceive. Yet, problematically, the collective
intelligence that could be garnered from this
additional perception is hindered by two key
barriers: the fact that data can be inaccurate
and misleading, and our own inability to
handle and interpret data. In this section we
explore the origins, trajectory and impact of
inaccurate information in human communica-
tion networks. We note our current inability to
correctly deploy, infer and apply meaning
from AI. We highlight current avenues for
reducing these barriers in order to fully exploit
our augmented collective intelligence.
5.1 ‘Fake News’
Humans have long expressed a desire to
record and share information: the first ency-
clopaedia was written in AD 77 (Gudger,
1924); libraries have been dated back 2,000
years earlier. The arrival of the telegraph and
Morse Code in 1835 enabled instantaneous
transmission of knowledge across great dis-
tances (Burns, 1988); now databases are ubiq-
uitous. Historical records indicate sharing
knowledge spurred many innovations (Bessen
and Nuvolari, 2016), and on a day-to-day
basis, enables individuals to make informed
decisions and actions. Unfortunately, not
all shared information is accurate.
Disinformation and misinformation, which
often fall under the misnomer ‘fake news’, are
of rising public concern. Disinformation
implies intentional creation and sharing of
manipulated or false information, whereas
misinformation refers to inadvertent sharing
(Lazer et al., 2018).
Fake news is not new: since humans could
speak, misinformation has spread via word
of mouth. The spread increased and quick-
ened with the arrival of newspapers and
pamphlets, then with mass media such as
television, finally exploding with the Internet
and especially social media (Burkhardt,
2017). What is new is that, compared to past
technological mediums, social media largely
344 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
lacks filtering, editorial judgement and fact-
checking (Allcott and Gentzkow, 2017).
Further, the communication network is infil-
trated by artificially intelligent bots, able to
pass as or augment human users, which can
be used to quickly deploy, share and spread
information across networks (Machado and
Konopacki, 2018). Such bots can be used to
sway public opinion. In fact, disinformation
affects stock prices (Carvalho et  al., 2011),
political opinions (Howard et  al., 2018),
and voting patterns (Allcott and Gentzkow,
2017) at least transiently. Evidence shows
that accurate stories take longer to spread but
have more purchase once spread (Vosoughi
et  al., 2018). In previous work (Mitchell
et al., 2016), we have demonstrated that even
error-prone ‘gossip’ can be a better strategy
than direct experiential learning for acquir-
ing true and useful information. The speed of
information transmission such as is provided
by social media can in some circumstances
outweigh the costs of incorrect information,
particularly if disinformation can be identi-
fied and quickly combated (Panagiotopoulos
et al., 2014).
Nevertheless, in at least some contexts, our
species seems often to communicate inac-
curate rather than accurate information. An
analysis of a data set of rumour cascades on
Twitter revealed fake news was 70% more
likely to be retweeted than the truth (Vosoughi
et al., 2018). There are suggestions that fake
news is more likely to be novel, and novelty
captures human attention. Perhaps worry-
ingly, the average American spends 23.6 hours
online weekly (Cole et  al., 2017), and 62%
get their news online (Gottfried and Shearer,
2016). Whilst there is evidence that our trust of
such news has decreased, exposure alone may
have negative impacts. Exposure can prime
thinking and conversational topics. When any
news enters the conversational sphere, trust in
the information increases (Hajli et al., 2014).
We humans seem to have a disposition to
trust information communicated by word of
mouth (Atika et  al., 2018; Huete-Alcocer,
2017). Conversations require resources such
as time, cognition, and sometimes emotional
investment. Actions occurring as conse-
quence of conversations result in further
deployment of resources. We know that, as a
social species, we respond to such evidence of
investment by others in our society (Zahavi,
1977). We posit that individual and collec-
tive resources may be consequently directed
away from more accurate topics which are of
perhaps more importance to our survival and
flourishing.
5.2 The Impact of Information
Accuracy
In further work with simulations and formal
analysis, we offered insight into the environ-
ments where humans may be vulnerable to
utilising incorrect information (Rauwolf,
2016). Information requires both time and
energy to gather. If information gathering
comes at a non-trivial cost, then we would
expect individuals to truncate their informa-
tion search after a period of time (Simon,
1956). However, given continual improve-
ments in technology, the cost of information
is falling; as a result we might expect indi-
viduals to be better informed. Importantly
though, even if information is easily obtained,
if the processing of that information is costly,
limiting information can be advantageous
(Rauwolf and Jones, 2019).
Rauwolf et al. (2015) show that when the
benefits of group dynamics conflict with the
accuracy of beliefs, false beliefs can become
the least-costly option. Across a variety
of contexts, individuals tend to prioritise
relationships with those who share simi-
lar values – a trait called value-homophily
(McPherson et  al., 2001). We have demon-
strated that it is in precisely these contexts
where individuals can be expected to use
incorrect information (Rauwolf et al., 2015).
When the social value or benefit provided by
the group outweighs the private cost of pos-
sessing incorrect information, it is advanta-
geous for the individual to maintain (or at
 EVOLUTIONARY PSYCHOLOGY AND ARTIFICIAL INTELLIGENCE
least act on) their false beliefs. Given that the
inaccuracy of political and religious beliefs
provides virtually no personal cost (Caplan,
2001), but group agreement may provide
social value through security, we would
expect humanity would struggle to remove
false beliefs, particularly in times of resource
scarcity and conflict (Stewart et al., 2020).
As the benefits conferred by AI and tech-
nology at large continue to improve and
secure individual basic needs, individuals
will likely pay a reduced private cost for pos-
sessing incorrect information across a broad-
ening array of contexts. Regardless of the
accuracy of an individual’s beliefs, the basic
needs of most individuals are improving. As
such, if individuals pay small personal costs
for false beliefs, but garner large social ben-
efits for group homogeneity, then we would
expect an expanding and resilient battle
against false beliefs.
Nevertheless, what costs an individual lit-
tle in isolation may cost a society a great deal
due to aggregate responses, particularly in a
democracy (Chote et al., 2016; Lewis, 2017).
Whether or not we should strive for increased
rate of information transmission in every case
(see the discussions of trust and freedom of
opinion above), we should almost certainly
prefer accurate communication, though here,
too, inaccuracy can sometimes lead to use-
ful innovation. Disinformation is a global and
long-running issue, and there are global ini-
tiatives to combat it. For example, Facebook
flags potential news stories to be reviewed
by third-party fact checkers; and through the
messaging system WeChat in China, users can
report fake news, which is then checked and
flagged. Crucial new initiatives introduce criti-
cal literacy into the education curriculum –
training children to recognise and ques-
tion information sources, particularly online
(Vasu et  al., 2018). Critical-thinking and
fact-checking skills, as well as basic under-
standing of algorithm mechanics and their
limitations, could enable the next genera-
tion to be better prepared to avoid such sce-
narios faced by us today (Guess et al., 2019).
345
Fact-checking can be as simple as conducting
a Web search on a topic and its source.
6 CONCLUSION
In this chapter we have discussed the impact of
AI on contemporary societies. We took a per-
spective of understanding how the changing
social and economic landscape induced by
AI interacts with the human information-
processing biases which evolved in very
different environments. We consider a better
understanding of these impacts imperative for
our society going forward as we optimise our
existence with AI, and for ensuring the AI and
the regulations we design to govern its use
both maximise benefit and minimise harm. We
discussed the impact on both collective and
individual human behaviour. Here we summa-
rise the key foci of this chapter. First, the accu-
racy of narratives surrounding AI could
critically impact optimal engagement with AI.
Next, we compared AI to a prosthetic nervous
system, which increases our perception and
agency. AI also enhances our capacity to
remember, coordinate, connect and communi-
cate; this has many positive but also some
negative outcomes. We considered the impact
on freedom and diversity of opinion, political
and economic impact, the mechanisms of
information spread, and vulnerability of human
trust and social coherence. The increased dis-
coverability and predictability facilitated by
AI requires serious consideration; there are
myriad beneficial and harmful current applica-
tions of AI, and no doubt more of both to
come. There are also of course many move-
ments to ensure AI is beneficial to our society
rather than harmful, though we did not take
time to touch on those much here, we view
such consideration and efforts as essential.
Coordinating and enforcing such pro-social
efforts has traditionally been called govern-
ance, and we hope this chapter may contribute
to making sensible governance easier to both
justify and employ constructively.
346 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Note
1  As of this writing, the impact of these assaults is
still a matter of urgent research and debate, but
the fact of significant, long-term, and on-going
expenditure in the attempts has been established
in both courts and academic writings (e.g. Mach-
ado and Konopacki, 2018; Woods, 2018; Landon-
Murray et al., 2019).
REFERENCES
Acemoglu, D. and Autor, D. (2011). Skills, tasks
and technologies: Implications for employ-
ment and earnings. In Ashenfelter, O. and
Card, D. (Eds.), volume 4 of Handbook of
Labor Economics, chapter 12, pages 1043–
1171. Amsterdam: Elsevier.
Allcott, H. and Gentzkow, M. (2017). Social
media and fake news in the 2016 election.
Journal of Economic Perspectives, 31(2):
211–236.
Anderson, J., Rainie, L., and Luchsinger, A.
(2018). Artificial intelligence and the future
of humans. Pew Research Center.
Anderson, M. and Jiang, J. (2018). Teens, social
media & technology 2018. Washington, DC:
Pew Internet & American Life Project. Retrieved
June 3, 2018, from https://www.pewresearch.
org/internet/2018/05/31/teens-social-media-
technology-2018/
André, J.-B. and Baumard, N. (2011). Social
opportunities and the evolution of fairness.
Journal of Theoretical Biology, 289:128–135.
Atika, A., Kusumawati, A., and Iqbal, M.
(2018). The effect of electronic word of
mouth, message source credibility, informa-
tion quality on brand image and purchase
intention. EKUITAS (Jurnal Ekonomi dan
Keuangan), 20(1):94–108.
Atkinson, A. B. (2015). Inequality: What can be
done? Cambridge, MA: Harvard University
Press.
Barone, G. and Mocetti, S. (2016). Inequality
and trust: New evidence from panel data.
Economic Inquiry, 54(2):794–809.
Bechtel, L. M. (1991). Shame as a sanction of
social control in biblical Israel: Judicial, political,
and social shaming. Journal for the Study of
the Old Testament, 16(49):47–76.
Behavioural Insights Team (2010). Applying
behavioural insight to health. Technical
Report 403936/1210, Cabinet Office, UK
Government, London.
Behavioural Insights Team (2012). Applying
behavioural insights to reduce fraud, error
and debt. Technical Report 408779/0212,
Cabinet Office, UK Government, London.
Bente, G., Baptist, O., and Leuschner, H.
(2012). To buy or not to buy: Influence of
seller photos and reputation on buyer trust
and purchase behavior. International Journal
of Human-Computer Studies, 70(1):1–13.
Berg, J., Dickhaut, J., and McCabe, K. (1995).
Trust, reciprocity, and social history. Games
and Economic Behavior, 10(1):122–142.
Berger, J. (2011). Arousal increases social trans-
mission of information. Psychological Science,
22(7):891–893.
Bessen, J. and Nuvolari, A. (2016). Knowledge
sharing among inventors: Some historical
perspectives. In Harhoff, D. and Lakhani, K.
(Eds.), Revolutionizing Innovation: Users,
Communities, and Open Innovation, page
135. MIT Press, Cambridge MA.
Boden, M., Bryson, J., Caldwell, D., Dautenhahn,
K., Edwards, L., Kember, S., Newman, P., Parry,
V., Pegman, G., Rodden, T., Sorell, T., Wallis,
M., Whitby, B., and Winfield, A. (2011).
Principles of robotics. The United Kingdom’s
Engineering and Physical Sciences Research
Council (EPSRC). Retrieved from https://epsrc.
ukri.org/research/ourportfolio/themes/
engineering/activities/principlesofrobotics/
Boero, R., Bravo, G., Castellani, M., and Squaz-
zoni, F. (2009). Reputational cues in repeated
trust games. The Journal of Socio-Economics,
38(6):871–877.
Bond, C. F. J. and DePaulo, B. M. (2006). Accu-
racy of deception judgments. Personality and
Social Psychology Review, 10(3):214–234.
Bradford, E. E., Jentzsch, I., and Gomez, J.-C.
(2015). From self to social cognition: Theory
of mind mechanisms and their relation to
executive functioning. Cognition, 138:21–34.
Brown, G. R., Dickins, T. E., Sear, R., and
Laland, K. N. (2011). Evolutionary accounts
of human behavioural diversity. Philosophical
Transactions of the Royal Society B: Biological
Sciences, 366:313–324.
Bryson, J. J. (2015). Artificial intelligence and
pro-social behaviour. In Misselhorn, C.,
 EVOLUTIONARY PSYCHOLOGY AND ARTIFICIAL INTELLIGENCE
editor, Collective Agency and Cooperation in
Natural and Artificial Systems: Explanation,
Implementation and Simulation, volume 122
of Philosophical Studies, pages 281–306.
Berlin: Springer.
Bryson, J. J. (2019). The past decade and future
of AI’s impact on society. In Towards a New
Enlightenment? In Cities, D (Eds.), A Trans-
cendent Decade, pages 150–185. Madrid:
Turner–BBVA.
Bryson, J. J. and Kime, P. P. (2011). Just an arti-
fact: Why machines are perceived as moral
agents. In Proceedings of the 22nd Interna-
tional Joint Conference on Artificial Intelli-
gence, pages 1641–1646, Barcelona. Morgan
Kaufmann.
Bryson, J. J. and Theodorou, A. (2019). How
society can maintain human-centric artificial
intelligence. In Toivonen-Noro, M. and Saari,
E. (Eds.), Human-Centered Digitalization and
Services, chapter 16, pages 305–323. Singa-
pore: Springer.
Buchan, N. R., Croson, R. T., and Solnick, S.
(2008). Trust and gender: An examination of
behavior and beliefs in the investment game.
Journal of Economic Behavior & Organiza-
tion, 68(3):466–476.
Burkhardt, J. M. (2017). History of fake news.
Library Technology Reports, 53(8):5–9.
Burnham, T., McCabe, K., and Smith, V. L.
(2000). Friend-or-foe intentionality priming in
an extensive form trust game. Journal of Eco-
nomic Behavior & Organization, 43(1):57–73.
Burns, R. (1988). The electric telegraph and the
development of picture telegraphy. In Papers
Presented at the Sixteenth IEE Week-End
Meeting on the History of Electrical Engineer-
ing, pages 80–86. IET.
Burns, W. E. (2003). Witch hunts in Europe and
America: An encyclopedia. Westport, CT:
Greenwood Publishing Group.
Buss, D. M. and Schmitt, D. P. (1993). Sexual
strategies theory: An evolutionary perspec-
tive on human mating. Psychological Review,
100(2):204.
Caplan, B. (2001). Rational ignorance versus
rational irrationality. Kyklos, 54(1):3–26.
Carvalho, C., Klagge, N., and Moench, E. (2011).
The persistent effects of a false news shock.
Journal of Empirical Finance, 18(4):597–615.
Cassiers, T. and Kesteloot, C. (2012). Socio-
spatial inequalities and social cohesion in
347
European cities. Urban Studies, 49(9):
1909–1924.
Cave, S., Coughlan, K., and Dihal, K. (2019).
‘Scary robots’: Examining public responses
to AI. In Proceedings of the AIES. Retrieved
May 24, 2020, from https://doi.org/10.1145/
3306618.3314232
Chavan, S. and Khot, T. S. (2013). Efficient and
reliable routing algorithm to enhance con-
nectivity in disaster scenario: Abc algorithm.
International Journal of Science and
Research, 4(5):891–896.
Chote, R., Nickell, S., and Parker, G. (2016).
Economic and fiscal outlook. Technical Report
Cm 9346, Office for Budget Responsibility,
London, UK. Retrieved on 22 August, 2020
from www.gov.uk/government/publications.
Cole, J., Suman, M., Schramm, P., and Zhou, L.
(2017). The 2017 digital future report:
Surveying the digital future. Los Angeles, CA:
USC Annenberg School Center for the Digital
Future.
de Visser, E. J., Monfort, S. S., McKendrick, R.,
Smith, M. A., McKnight, P. E., Krueger, F.,
and Parasuraman, R. (2016). Almost human:
Anthropomorphism increases trust resilience
in cognitive agents. Journal of Experimental
Psychology: Applied, 22(3):331.
Delgado, M. R., Frank, R. H., and Phelps, E. A.
(2005). Perceptions of moral character mod-
ulate the neural systems of reward during
the trust game. Nature Neuroscience,
8(11):1611–1618.
Eagle, N. and Pentland, A. S. (2003). Social net-
work computing. In Dey, A. K., Schmidt, A.,
and McCarthy, J. F. (Eds.), UbiComp 2003:
Ubiquitous Computing, pages 289–296,
Berlin, Heidelberg: Springer Berlin Heidelberg.
Elish, M. C. and Boyd, D. (2018). Situating
methods in the magic of big data and AI.
Communication Monographs, 85(1):57–80.
Ert, E., Fleischer, A., and Magen, N. (2016).
Trust and reputation in the sharing economy:
The role of personal photos in Airbnb. Tourism
Management, 55:62–73.
Fagerstrøm, A., Pawar, S., Sigurdsson, V.,
Foxall, G. R., and de Soriano, M. Y. (2017).
That personal profile image might jeopardize
your rental opportunity! On the relative
impact of the seller’s facial expressions upon
buying behavior on AirbnbTM. Computers in
Human Behavior, 72:123–131.
348 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Fehr, E. (2009). On the economics and biology
of trust. Journal of the European Economic
Association, 7(2–3):235–266.
Fisher, R. A. (1930). The genetical theory of natu-
ral selection. Clarendon Press, Oxford (2nd
edn., 1958, New York, Dover. Variorum edition,
Bennett JH (Ed.), 1999, Oxford University Press).
Franco, R. Z., Fallaize, R., Lovegrove, J. A., and
Hwang, F. (2016). Popular nutrition-related
mobile apps: A feature assessment. JMIR
mHealth and uHealth, 4(3):e85.
Gelman, A., Goel, S., Rivers, D., and Roths-
child, D. (2016). The mythical swing voter.
Quarterly Journal of Political Science,
11(1):103–130.
Gent, E. (2015). Ai: Fears of ‘playing god’
[control & automation artificial intelligence].
Engineering & Technology, 10(2):76–79.
Gerbaudo, P. (2018). Tweets and the streets:
Social media and contemporary activism.
London: Pluto Press.
Gilligan, I. (2007). Neanderthal extinction and
modern human behaviour: The role of cli-
mate change and clothing. World Archaeol-
ogy, 39(4):499–514.
Godfrey-Smith, P. (2017). Other minds: The
octopus, the sea, and the deep origins of
consciousness. New York, NY: Farrar, Straus
and Giroux.
Gottfried, J. and Shearer, E. (2016). News Use
Across Social Medial Platforms 2016. Wash-
ington, DC: Pew Research Center. Retrieved
24 May, 2020 from http://www.journalism.
org/2016/05/26/news-use-across-social-me
dia-platforms-2016/.
Grönlund, K. and Setälä, M. (2007). Political
trust, satisfaction and voter turnout. Com-
parative European Politics, 5(4):400–422.
Gudger, E. W. (1924). Pliny’s historia naturalis.
The most popular natural history ever pub-
lished. Isis, 6(3):269–281.
Guess, A., Nagler, J., and Tucker, J. (2019). Less
than you think: Prevalence and predictors of
fake news dissemination on Facebook. Sci-
ence Advances, 5(1), eaau4586.
Hajli, N., Lin, X., Featherman, M., and Wang, Y.
(2014). Social word of mouth: How trust
develops in the market. International Journal
of Market Research, 56(5):673–689.
Hammack, P. L. (2008). Narrative and the cul-
tural psychology of identity. Personality and
Social Psychology Review, 12(3):222–247.
Hao, T., Xing, G., and Zhou, G. (2013). isleep:
Unobtrusive sleep quality monitoring using
smartphones. In Proceedings of the 11th
ACM Conference on Embedded Networked
Sensor Systems, page 4. ACM.
Hess, K. and Waller, L. (2014). The digital pil-
lory: Media shaming of ‘ordinary’ people for
minor crimes. Continuum, 28(1):101–111.
Hetherington, M. J. (1999). The effect of political
trust on the presidential vote, 1968–96. Ameri-
can Political Science Review, 93(2):311–326.
Howard, P. N., Kollanyi, B., Bradshaw, S., and
Neudert, L.-M. (2018). Social media, news
and political information during the US elec-
tion: Was polarizing content concentrated in
swing states? arXiv preprint arXiv:1802.03573.
Huete-Alcocer, N. (2017). A literature review of
word of mouth and electronic word of
mouth: Implications for consumer behavior.
Frontiers in Psychology, 8:1256.
Institute of Electrical and Electronics Engineers
(IEEE) (2019). Ethically aligned design: A
vision for prioritizing human well-being with
autonomous and intelligent systems. Technical
report, The IEEE Global Initiative on Ethics of
Autonomous and Intelligent Systems, first
edition.
Johnson, N. D. and Mislin, A. A. (2011). Trust
games: A meta-analysis. Journal of Economic
Psychology, 32(5):865–889.
Kim, D. J., Ferrin, D. L., and Rao, H. R. (2008).
A trust-based consumer decision-making
model in electronic commerce: The role of
trust, perceived risk, and their antecedents.
Decision Support Systems, 44(2):544–564.
Kosinski, M., Stillwell, D., and Graepel, T.
(2013). Private traits and attributes are
predictable from digital records of human
behavior. Proceedings of the National
Academy of Sciences, 110(15), 5802–5805.
Kraemer, P. J. and Golding, J. M. (1997).
Adaptive forgetting in animals. Psychonomic
Bulletin & Review, 4(4):480–491.
Kramer, K. L. and Ellison, P. T. (2010). Pooled
energy budgets: Resituating human
energy-allocation trade-offs. Evolutionary
Anthropology: Issues, News, and Reviews,
19(4):136–147.
Lahr, M. M. (2016). The shaping of human
diversity: Filters, boundaries and transitions.
Philosophical Transactions of the Royal Soci-
ety B, 371(1698):20150241.
 EVOLUTIONARY PSYCHOLOGY AND ARTIFICIAL INTELLIGENCE
Landon-Murray, M., Mujkic, E., and Nussbaum, B.
(2019). Disinformation in contemporary US
foreign policy: Impacts and ethics in an era of
fake news, social media, and artificial intelli-
gence. Public Integrity, 21(5):512–522.
Lazer, D. M., Baum, M. A., Benkler, Y., Berinsky,
A. J., Greenhill, K. M., Menczer, F., and
Schudson, M. (2018). The science of fake
news. Science, 359(6380):1094–1096.
Lee, E., Bristow, J., Faircloth, C., and Macvar-
ish, J. (2014). Parenting culture studies.
London: Palgrave Macmillan.
Lewis, M. (2017). Why the scariest nuclear threat
may be coming from inside the White House.
Vanity Fair. Retrieved 24 May, 2020, from
https://www.vanityfair.com/news/2017/07/
department-of-energy-risks-michael-lewis.
Liang, Y. and Lee, S. A. (2017). Fear of autono-
mous robots and artificial intelligence: Evi-
dence from national representative data
with probability sampling. International Jour-
nal of Social Robotics, 9(3):379–384.
Logg, J. M., Minson, J. A., and Moore, D. A.
(2019). Algorithm appreciation: People
prefer algorithmic to human judgment.
Organizational Behavior and Human Deci-
sion Processes, 151:90–103.
Machado, C. and Konopacki, M. (2018). Compu-
tational power: Automated use of WhatsApp
in the Brazilian elections. Medium. Retrieved
24 May, 2020 from https://feed.itsrio.org/
computational-power-automated-use-of-
whatsapp-in-the-elections-59f62b857033.
Manapat, M. L., Nowak, M. A., and Rand, D.
G. (2013). Information, irrationality, and the
evolution of trust. Journal of Economic
Behavior & Organization, 90:S57–S75.
Manapat, M. L. and Rand, D. R. (2012).
Delayed and inconsistent information and
the evolution of trust. Dynamic Games and
Applications, 2:401–410.
Masuda, N. and Nakamura, M. (2012). Coevolu-
tion of trustful buyers and cooperative sellers
in the trust game. PLoS One, 7(9):1–11.
Mayer-Schönberger, V. (2007). Useful void: The
art of forgetting in the age of ubiquitous
computing, (Working Paper No. RWP07-022).
Cambridge, MA: Harvard University.
McCarty, N. M., Poole, K. T., and Rosenthal, H.
(2016). Polarized America: The dance of ide-
ology and unequal riches. Cambridge, MA:
MIT Press, second edition.
349
McDonald, M. M., Navarrete, C. D., and Van
Vugt, M. (2012). Evolution and the psychol-
ogy of intergroup conflict: The male warrior
hypothesis. Philosophical Transactions of the
Royal Society B, 367(1589):670–679.
McNamara, J. M., Stephens, P. A., Dall, S. R.,
and Houston, A. I. (2009). Evolution of trust
and trustworthiness: Social awareness
favours personality differences. Proceedings
of the Royal Society B: Biological Sciences,
276(1657):605–613.
McPherson, M., Smith-Lovin, L., and Cook, J. M.
(2001). Birds of a feather: Homophily in social
networks. Annual Review of Sociology,
27(1):415–444.
Meijer, A. and Wessels, M. (2019). Predictive
policing: Review of benefits and drawbacks.
International Journal of Public Administration,
42(12), 1031–1039.
Mitchell, D., Bryson, J. J., Rauwolf, P., and
Ingram, G. P. (2016). On the reliability of
unreliable information: Gossip as cultural.
Interaction Studies, 17(1), 1–18.
Morris, N. (2002). The myth of unadulterated
culture meets the threat of imported
media. Media, Culture & Society, 24(2):
278–289.
Moscovici, S. (1981). On social representations.
In Forgas, J. (Ed.), Social cognition: Perspec-
tives on everyday understanding, 8(12):181–
209, London: Academic Press.
Moscovici, S. (2001). Why a theory of social
representation? The representations of the
Social. Bridging theoretical traditions.
8–37.
Müller, V. C. and Bostrom, N. (2016). Future
progress in artificial intelligence: A survey of
expert opinion. In Müller, V. C. (Ed.), Funda-
mental Issues of Artificial Intelligence, pages
555–572. Berlin: Springer.
Nairne, J. S., Pandeirada, J. N., Gregory, K. J.,
and Van Arsdall, J. E. (2009). Adaptive
memory: Fitness relevance and the hunter-
gatherer mind. Psychological Science,
20(6):740–746.
Oosterhof, N. N. and Todorov, A. (2008). The
functional basis of face evaluation. Proceed-
ings of the National Academy of Sciences,
105(32):11087–11092.
Panagiotopoulos, P., Bigdeli, A. Z., and Sams, S.
(2014). Citizen–government collaboration on
social media: The case of Twitter in the 2011
350 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
riots in England. Government Information
Quarterly, 31(3):349–357.
Parker, E. S., Cahill, L., and McGaugh, J. L.
(2006). A case of unusual autobiographical
remembering. Neurocase, 12(1):35–49.
Pentland, A. (2015). Social physics: How social
networks can make us smarter. New York,
NY: Penguin.
Perlman, E. R. and Sprick Schuster, S. (2016).
Delivering the vote: The political effect of
free mail delivery in early twentieth century
America. The Journal of Economic History,
76(3):769–802.
Piette, A. (2018). Muriel Spark and fake news.
Textual Practice, 32(9):1577–1591.
Ponte, E. B., Carvajal-Trujillo, E., and Escobar-
Rodríguez, T. (2015). Influence of trust and
perceived value on the intention to purchase
travel online: Integrating the effects of assur-
ance on trust antecedents. Tourism Manage-
ment, 47:286–302.
Raber, I., McCarthy, C. P., and Yeh, R. W.
(2019). Health insurance and mobile health
devices: Opportunities and concerns. Journal
of the American Medical Association (JAMA),
321(18):1767–1768.
Rauwolf, P. (2016). Understanding the ubiquity
of self-deception: The evolutionary utility of
incorrect information. PhD thesis, University
of Bath.
Rauwolf, P. and Bryson, J. J. (2018). Expecta-
tions of fairness and trust co-evolve in envi-
ronments of partial information. Dynamic
Games and Applications, 8(4):891–917.
Rauwolf, P. and Jones, A. (2019). Exploring the
utility of internal whistleblowing in healthcare
via agent-based models. BMJ Open, 9(1).
Rauwolf, P., Mitchell, D., and Bryson, J. J. (2015).
Value homophily benefits cooperation but
motivates employing incorrect social infor-
mation. Journal of Theoretical Biology,
367:246–261.
Richerson, P. J., Bettinger, R. L., and Boyd, R.
(2005). Evolution on a restless planet: Were
environmental variability and environmental
change major drivers of human evolution.
In Woketits, F. M. and Ayala, F. J. (Eds.),
Handbook of Evolution, 2:223–242.
Robinson, L., Cotten, S. R., Ono, H., Quan-
Haase, A., Mesch, G., Chen, W., Schulz, J.,
Hale, T. M., and Stern, M. J. (2015). Digital
inequalities and why they matter.
Information, Communication & Society,
18(5):569–582.
Roser, M. and Ortiz-Ospina, E. (2017). Global
extreme poverty. Technical report, Our World
in Data. Retrieved 24 December, 2019 from
https://ourworldindata.org/extreme-poverty.
Rotsidis, A., Theodorou, A., Bryson, J. J., and
Wortham, R. H. (2019). Improving robot
transparency: An investigation with mobile
augmented reality. In 28th IEEE Interna-
tional Symposium on Robot and Human
Interactive Communication (RO-MAN), New
Delhi. IEEE.
Roughgarden, J., Oishi, M., and Akçay, E.
(2006). Reproductive social behavior: Coop-
erative games to replace sexual selection.
Science, 311(5763):965–969.
Scharlemann, J. P., Eckel, C. C., Kacelnik, A., and
Wilson, R. K. (2001). The value of a smile:
Game theory with a human face. Journal of
Economic Psychology, 22(5):617–640.
Scheidel, W. (2017). The great leveler: Violence
and the history of inequality from the Stone
Age to the twenty-first century. Princeton,
NJ: Princeton University Press.
Shi, F., Teplitskiy, M., Duede, E., and Evans, J.
A. (2019). The wisdom of polarized crowds.
Nature Human Behaviour, 3:329–336.
Siles, I. and Walker, I. D. (2009). Design, con-
struction, and testing of a new class of
mobile robots for cave exploration. In
Mechatronics, 2009. ICM 2009. IEEE Inter-
national Conference on, pages 1–6. IEEE.
Simkulet, W. (2019). Informed consent and
nudging. Bioethics, 33(1):169–184.
Simon, H. A. (1956). Rational choice and the
structure of the environment. Psychological
Review, 63(2):129–138.
Smaldino, P. E., Newson, L., Schank, J. C., and
Richerson, P. J. (2013). Simulating the evolu-
tion of the human family: Cooperative
breeding increases in harsh environments.
PLoS One, 8(11):e80753.
Smith, E. A. (2010). Communication and col-
lective action: Language and the evolution of
human cooperation. Evolution and Human
Behavior, 31(4):231–245.
Sparrow, R. (2019). Robotics has a race prob-
lem. Science, Technology, & Human Values,
45(3), 538–560.
Spring, B., Gotsis, M., Paiva, A., and Spruijt-
Metz, D. (2013). Healthy apps: Mobile devices
 EVOLUTIONARY PSYCHOLOGY AND ARTIFICIAL INTELLIGENCE
for continuous monitoring and intervention.
IEEE Pulse, 4(6):34.
Starbird, K. and Palen, L. (2011). ‘Voluntweet-
ers’: Self-organizing by digital volunteers in
times of crisis. In Proceedings of the SIGCHI
Conference on Human Factors in Computing
Systems, CHI ‘11, pages 1071–1080, New
York, NY. ACM.
Stewart, A. J., McCarty, N., and Bryson, J. J.
(2020). Polarization under rising inequality
and economic decline. arXiv preprint arXiv:
1807.11477.
Stone, J. and Moskowitz, G. B. (2011). Non-
conscious bias in medical decision making:
What can be done to reduce it? Medical
Education, 45(8):768–776.
Sujarwoto, S. and Tampubolon, G. (2016). Spa-
tial inequality and the internet divide in
Indonesia 2010–2012. Telecommunications
Policy, 40(7):602–616.
Terkel, S. (1974). Working. New York: The New
Press.
Terry, D. J. and Hogg, M. A. (1996). Group
norms and the attitude-behavior relationship:
A role for group identification. Personality and
Social Psychology Bulletin, 22(8):776–793.
Teste, F. P., Simard, S. W., Durall, D. M., Guy, R. D.,
Jones, M. D., and Schoonmaker, A. L. (2009).
Access to mycorrhizal networks and roots
of trees: Importance for seedling survival
and resource transfer. Ecology, 90(10):
2808–2822.
Theodorou, A., Wortham, R. H., and Bryson, J. J.
(2017). Designing and implementing transpar-
ency for real time inspection of autonomous
robots. Connection Science, 29(3):230–241.
Toma, C. L. (2010). Perceptions of trustworthi-
ness online: The role of visual and textual
information. In Proceedings of the 2010
ACM Conference on Computer Supported
Cooperative Work, pages 13–22. ACM.
Tucker, B. (2018). ‘That’s problematic’: Tracing
the birth of call-out culture. Critical Reflec-
tions: A Student Journal on Contemporary
Sociological Issues, 6.
United Nations General Assembly (1948). Uni-
versal declaration of human rights. Technical
report, New York.
Vasu, N., Ang, B., Teo, T.-A., Jayakumar, S.,
Raizal, M., and Ahuja, J. (2018). Fake news:
National security in the post-truth era. RSIS.
In Policy Report. Singapore: Nanyang
351
Technological University. Retrieved 24 May,
2020, from https://www.rsis.edu.sg/wp-­
content/uploads/2018/01/PR180313_Fake-
News_WEB.pdf
Vieweg, S., Hughes, A. L., Starbird, K., and
Palen, L. (2010). Microblogging during two
natural hazards events: What Twitter may con-
tribute to situational awareness. In Proceed-
ings of the SIGCHI Conference on Human
Factors in Computing Systems, CHI ‘10, pages
1079–1088, New York, NY. ACM.
Vosoughi, S., Roy, D., and Aral, S. (2018).
The spread of true and false news online.
Science, 359(6380):1146–1151.
Waytz, A., Heafner, J., and Epley, N. (2014). The
mind in the machine: Anthropomorphism
increases trust in an autonomous vehicle.
Journal of Experimental Social Psychology,
52:113–117.
Webb, H., Jirotka, M., Stahl, B. C., Housley, W.,
Edwards, A., Williams, M., Procter, R., Rana,
O., and Burnap, P. (2016). Digital wildfires:
Hyper-connectivity, havoc and a global
ethos to govern social media. ACM SIGCAS
Computers and Society, 45(3):193–201.
Wilson, H. and Theodorou, A. (2019). Slam the
brakes: Perceptions of moral decisions in
driving dilemmas. In International Workshop
in Artificial Intelligence Safety (AISafety),
IJCAI, Macau.
Wilson, R. K. and Eckel, C. C. (2006). Judging
a book by its cover: Beauty and expectations
in the trust game. Political Research Quarterly,
59(2):189–202.
Woods, L. (2018). ICO reacts to use of data ana-
lytics in micro-targetting for political purposes
reports: United Kingdom. European Data Pro-
tection Law Review (EDPL), 4:381–383.
Wu, Y., Kosinski, M., and Stillwell, D. (2015).
Computer-based personality judgments are
more accurate than those made by humans.
Proceedings of the National Academy of
Sciences, 112(4):1036–1040.
Yamagishi, T. (2011). Trust: The evolutionary
game of mind and society. Berlin: Springer,
Berlin.
Zahavi, A. (1977). The testing of a bond.
Animal Behaviour, 25:246–247.
Zuboff, S. (2015). Big other: Surveillance capi-
talism and the prospects of an information
civilization. Journal of Information Technology,
30(1):75–89.

Evolutionary Psychology
and Robotics
Evolutionary psychology can impact intelligent
robots in two different ways:
1 The expression of evolutionary psychology in the
human brain and emergent mind allows it to create
intelligent robots, with some of their development
focused on duplicating the neurophysiology of
the human brain and the cognition of the human
mind. As with many complex technologies in the
past, it is uncertain whether human psychology
will be sufficient to manage the consequences of
its own invention.
2 The future expression of evolutionary psychology,
albeit with a different theoretical framework, will
affect the autonomous intelligent robot in the
physical structure of its brain and the cognition of
its mind.
ROBOT EVOLUTION: LAMARCK
VS DARWIN
Evolutionary psychology holds that Darwinian
processes of natural selection apply to the
brain, mind, and resulting psychological traits,
18
Robert Finkelstein
e.g., memory, perception, and language, just
as they do to physiological traits (Buss, 2005).
Darwinian evolution is a well-tested scientific
theory that explains how organic complexity
can arise from simplicity. Evolutionary psy-
chology is expected to provide a basis for
explaining the origin of the human brain, the
most complex entity in the known universe
(Aunger, 2002), its manifestation in the human
mind, and the mind’s psychological phenom-
ena. The process for the evolution of the
autonomous intelligent robot, however, will
be Lamarckian evolution and not Darwinian
evolution.
In biology, the discredited Lamarckian
theory of evolution claims that when the
environment changes the phenotype of an
organism changes, such that the organism
is better adapted to the environmental factor
that caused the change (Hull, 2000), and the
genotype can then pass the improved adapta-
tion to the organism’s progeny (albeit, hav-
ing died in 1829, Jean-Baptiste Lamarck
knew nothing of genes or the genotype).
 EVOLUTIONARY PSYCHOLOGY AND ROBOTICS
Recent research has shown that for at least a
few organisms, such as certain roundworms,
pseudo-Lamarckian evolution seems to take
place by passing to its progeny, independent
of its genome, an individual’s acquired immu-
nity to a virus (Rechavi, 2009). Technology is
sometimes said to advance with Lamarckian
evolution, rather than with Darwinian evo-
lution, because of the high rate of techno-
logical change compared with the relative
slowness of Darwinian evolution, and the
ability of changes in one generation of a
technology to be passed on to the next gen-
eration. However, technological change was
not always this rapid for Homo sapiens and
its ancestors; a flint-cutting tool might have
taken thousands of years before manifesting
some improvement. In any event, the prin-
ciples of Lamarckian evolution with respect
to technological evolution have been a meta-
phor, not the actual evolutionary process that
was claimed to take place in an organism. In
the future, however, it might indeed become
the actual process for the evolution of robots.
If a future robot experiences an incident that
causes it to change its behavior, physiologi-
cal structure, or cognitive ability to better
survive or improve its functionality, it could
then pass the improvement to its progeny.
The robot would create its descendants via
autopoiesis (i.e., self-replication), albeit with
improvements. Furthermore, such a robot
would be able to communicate globally with
its contemporaries and could convince them
to implement the improvements immediately.
The advent of autonomous vehicles and
other robots, made possible by advances
in neural networks with deep learning
(Goodfellow et  al., 2016) and other artifi-
cial intelligence (AI) tools and techniques,
processing chips, communications networks,
knowledgebases, and sensors, will become
a foundational technology leading to mani-
fold kinds of autonomous intelligent robots
over the coming decades. Over the ensuing
decades the cognitive ability of robots will
evolve to replicate or surpass human capa-
bilities. Some AI processes, such as neural
353
networks, can produce robot behavior that is
as unpredictable and inexplicable as human
behavior. As robot cognition increases over
time, it is likely that robot minds will increas-
ingly exhibit psychological phenomena.
The scientist and science-fiction writer
Isaac Asimov anticipated the need, in the 21st
century, for what he called robopsychologists
to analyze and cure the mental illness that
he expected to emerge in some intelligent
machines, and which would engender behav-
ior that is dangerous to humans (Asimov,
1950). With the evolution of autonomous
intelligent robots following Lamarckian,
rather than Darwinian, principles, the study
of robopsychology will be needed sooner
rather than later.
A COMPUTATIONAL THEORY OF
MIND
In 1943 Warren McCulloch and Walter Pitts
concluded that neurons perform computa-
tional processes (McCulloch and Pitts, 1943).
Subsequently, the basis for a computational
theory (actually, a hypothesis) of mind was
formed from the confluence of several disci-
plines: neuroscience, cognitive psychology,
AI, robotics, information science, and semiot-
ics. It envisions that the mind can be modeled
as a set of processes which have computa-
tional equivalents (Albus and Meystel, 2001;
Meystel and Albus, 2002). If such a theory
were established on a foundation of evidence,
it could be tested experimentally with soft-
ware and hardware. Machines, such as robots,
could be built which model the brain and test
whether a human-like mind emerges from the
machine brain. This test would be a more
comprehensive version of the Turing test for
machine intelligence.
The computational theory of the human
mind intersects with evolutionary psychology
in holding that the human brain is a system of
computational subsystems designed by natu-
ral selection over eons in our Homo sapiens
354 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
ancestors and their ancestors (e.g., Homo
habilis, Homo erectus, Homo heidelbergen-
sis, etc.) for foraging, hunting, reproducing,
and outmaneuvering entities such as animals,
plants, objects in the environment, and other
hominids. It postulates that our cognition
and emotions are produced by evolved algo-
rithms computed by our subconscious mind.
Machines can be very good at developing
and computing algorithms, often better than
Homo sapiens.
A consequence of a computational theory
of mind is that the mind consists of a set of
processes, each of which has a computational
equivalent. These processes include emotion,
imagination, thought, reason, feeling, percep-
tion, knowledge, cognition, meaning, under-
standing, belief, planning, wisdom, attention,
awareness, consciousness, intelligence, a
sense of good and evil, a sense of justice and
duty, and an appreciation of beauty. Each of
these processes can be represented in robots,
as shown in Table 18.1 (from Albus, 2001):
Table 18.1  Processes of mind and their computational equivalents
Processes of the human mind
Emotion
Imagination
Thought
Reason
Feeling
Perception
Knowledge
Cognition
Meaning
Understanding
Belief
Planning
Wisdom
Attention
Awareness
Consciousness
Intelligence
Sense of good and evil
Sense of justice and duty
Appreciation of beauty
Source: Albus (2001).
To further explain the meaning of some of
the computational equivalents in the table,
consider that:
“meaning is the set of semantic relationships that
exist between the internal knowledge database
and the external world. Meaning establishes what
is intended or meant by behavioral actions, and
defines what entities, events, and situations in the
knowledge database refer to in the world.”
“Understanding occurs when the system’s internal
representation of external reality is adequate for
generating intelligent behavior.” “Planning is a
thinking process whereby a system imagines the
future and selects the best course of action to
achieve a goal state.” “Awareness is a condition
wherein a system has knowledge of the structure,
dynamics, and meaning of the environment in
which it exists.” “Consciousness is a state or condi-
tion in which an intelligent system is aware of itself,
its surroundings, its situation, its intentions, and its
feelings.” (Albus and Mystel, pp xii–xiii, 2001)
Despite centuries of philosophical argument,
it is now generally agreed that the mind is
what the brain does, so there is no need to
Computational equivalents
Value judgment; evaluation of good and bad
Modeling, and simulation; visualization
Analysis of what is imagined
Logic applied to thought
Experience of sensory input or emotional state
Transforming sensory data into knowledge
Information organized so it is useful
Analysis, evaluation, and use of knowledge
Relationships between forms of knowledge
Correspondence between model and reality
Level of confidence assigned to knowledge
Examination of possible future actions
Decisions likely to achieve long-term goals
Focusing perception on what is important
Operational state of perceptual processes
Operational state of cognitive processes
Ability to achieve purpose or goals despite uncertainty
Value judgment output
Value judgment output
Value judgment output
 EVOLUTIONARY PSYCHOLOGY AND ROBOTICS
invoke mysterious spiritual phenomena
outside the realm of science. Likewise, the
robot mind consists of the computational
processes of the robot brain. And its structure
and process will be subject to technological
evolution.
Thus the brain, whether human or robot,
is primarily a control system, generating and
controlling behavior. It likely developed in
ancient primitive organisms to control mobil-
ity and path planning to avoid threats and
take advantage of opportunities for survival
and reproduction.
Any control system is designed, by nature
or humans, to achieve or maintain a goal (a
desired result or state of the system or world).
Organisms with many offspring can sur-
vive with simple brains; higher intelligence
allows those with fewer offspring to survive
with higher probability. Robots need not have
greater intelligence or more complex cogni-
tion than humans. They should only have
the ‘IQ’ needed to perform their functions,
perhaps insect-level intelligence. Excessive
intelligence would be inefficient economi-
cally and operationally, perhaps even inter-
fering with the ability of the machine to do
its job well.
However, many jobs will require robots to
have human-level intelligence or better, some
with high levels of expertise that is narrowly
focused, such as for performing colonosco-
pies, but others will need high levels of cog-
nition that is broad in scope, such as being a
fifth-grade school teacher where knowledge
and skill for interacting with human students
is needed along with knowledge for effective
teaching. Some robots will need greater intel-
ligence in order to interact with humans in
a desirable way, despite their simpler basic
jobs. For example, an autonomous car need
only know how to drive in complex envi-
ronments in order to perform its basic func-
tion as a chauffeur. But if human passengers
would like to converse with the car in natu-
ral language on manifold topics, or if the
car is to be a partner of a law-enforcement
officer or a soldier and must know policies
355
and procedures, tactics and strategies, facial
recognition, natural language (in multi-
­
ple languages) with the ability to banter or
employ formal procedural language, and
have general situational awareness with, for
example, the ability to discern subtle impor-
tant cues from the environment, then the
robot will have to know much more than how
to drive, just as its human equivalent would.
Likewise, the intelligence of household
robots may vary, depending on the needs and
wants of their owners.
ROBOT INTELLIGENCE, LEARNING,
ADAPTATION, WISDOM, AND SELF-
AWARENESS
Autonomous robots will have varying
degrees of intelligence and ability to learn
and adapt. Robots with human-level intelli-
gence or greater will also need wisdom if
humanity is to avoid the oft-predicted Robot
Apocalypse. Robots will also need their
behavior to be guided by an ethics and
morality, but wisdom provides the frame-
work for ethical and moral behavior.
In a pragmatic definition, an intelligent
system is a system with the ability to act
appropriately (or make an appropriate choice
or decision) in an uncertain environment.
An appropriate action (or choice) is that
which maximizes the probability of success-
fully achieving the system’s mission, func-
tion, purpose, or goals (Meystel and Albus,
2002). Machine intelligence need not be at
the human level. For example, appropriate
autonomous vehicle intelligence is the abil-
ity of an autonomous vehicle to perform at
least as skilled as a good human driver would
under a variety of conditions. It does not
need to know how to play chess or make an
omelet, although it could be given additional
cognition to enhance its abilities to perform
functions other than driving. The appropriate
robot intelligence for an application depends
on the user’s requirements and the technical,
356 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
operational, and economical feasibility of
achieving the desired level of intelligence.
Many robots may only require insect-level
intelligence to perform their functions, but
even these robots will need behavioral con-
straints to avoid running amok. Bees, ants,
and termites, for example, work together with
their hive or colony cohort in a purposive way
and generally avoid killing each other.
There are many definitions of learning.
Our preferred synthesized definition of learn-
ing is the acquisition of knowledge, skill,
ability, or understanding by study, instruc-
tion, or experience, as evidenced by achiev-
ing growing success (improved behavior),
with respect to suitable metrics, in a fixed
environment. Learning takes place when the
system’s behavior increases the efficiency
with which data, information, and knowl-
edge is processed so that desirable states are
reached, errors avoided, or a portion of the
system’s environment is controlled. Robots
with varying degrees of intelligence will
be capable of learning as may be required
by their applications, even robots with rela-
tively low intelligence. Even Caenorhabditis
elegans, a worm one millimeter in length
with just 302 neurons, is capable of learning
(Ardiel and Rankin, 2010).
In our view of learning is different from
adaptation. Our definition of adaptation is a
change in behavior (or structure) in response
to a changed environment. An adapted system
is able to maintain critical or essential vari-
ables within physical (or physiological) limits
(e.g., homeostasis). Adaptation occurs where
the changed behavior (or structure) increases
the probability that the system can achieve
its function or purpose (e.g., maintain homeo-
stasis) by adjusting to the new or changed envi-
ronment. Thus, in our view, learning occurs in
a fixed environment while adaptation occurs
in a changed environment.
To understand wisdom requires wisdom
(Sternberg, 1995). While there are many
projects to develop robot learning and intel-
ligence, there are none yet to develop robot
wisdom, an ability humans may need in their
robots if they are to survive among them.
The love of (or search for) wisdom is the
original meaning of the word philosophy.
But the notion of wisdom is a bit elusive.
According to Martin Henry Fischer (1879–
1962), knowledge is a process of aggregating
facts while wisdom is their simplification.
Our preferred definition of wisdom is: the
ability to see the forest as well as the trees.
A decision by a human or robot may seem to
be the right one when considered in the con-
text of the details of the problem or the short
term, i.e., the trees, or in the context of the
big picture or the long term, i.e., the forest,
but not both. For example, some immediate
or near-term solutions for problems may later
become problems themselves, while some
long-term solutions for problems may cause
new near-term problems.
Self-awareness is the ability to simulate
oneself, as in a mental world model. A robot
can be considered to be self-aware if it is able
to predict what it will perceive in the future
based on its current state and behavior, and
use those predictions to plan future behavior
(Albus and Meystel, 2001).
RULE-BASED SYSTEMS, NEURAL
NETWORKS, AND GENETIC
ALGORITHMS
There are a number of AI tools which can be
used to evolve the robot mind to higher cog-
nition and greater sentience, including sym-
bolic processes, connectionist processes, and
genetic algorithms. It is likely that a synthe-
sis of these AI tools, perhaps along with
some that have not been invented yet, might
be required.
Symbolic processes, consisting of rule-based
(or expert) systems, have been used success-
fully to replicate specialized human intelli-
gence or expertise within narrow domains of
knowledge (Hayes-Roth et  al., 1983). Expert
systems, featuring strings of if-then format-
ted rules (Durkin, 1994), have achieved some
 EVOLUTIONARY PSYCHOLOGY AND ROBOTICS
degree of success in commercial and military
applications. They are suitable for such
autonomous robot tasks as high-level plan-
ning, navigation, tactics, strategies, and por-
tions of sensory processing, such as high-level
vision. Conventional symbolic (knowledge-
based) processes are limited by the ability
of a human expert to foresee situations and
environments that a robot will encounter,
and to program, with sufficient if-then rules,
appropriate responses to threats and opportu-
nities that the robot may encounter.
However, rule-based systems can be
designed to learn, to modify themselves,
which would greatly broaden their domain.
A robot interacting with the environment
and encountering a situation that contradicts
an existing rule, or does not have a rule to
address the current situation, could modify an
existing rule to accommodate the new real-
ity or create a new rule. Rule-based systems
could be used to address a current weakness
of AI – its general inability to deal with con-
text, such as the relationship of entities with
time, space, and causality (Brézillon, 1999).
Deep learning, for example, is not generally
suitable for teaching a robot context, such as
how the existence and interaction of entities
relate to each other and the world in terms
of spatial and temporal phenomena, or the
chains of causal events (Edell, 2018). For
example, a rule-based system could state that
if a non-moveable object, like the Brooklyn
Bridge, is in one place, like New York City,
it cannot also be in another place, like next
to the Golden Gate Bridge in San Francisco.
Or it can be instructed in, or directly expe-
rience if it is in a robot, causality; e.g., if a
person trips on a crack in the sidewalk, that
may cause him to fall; and if he falls, he may
be injured.
Connectionist processes, consisting of arti-
ficial neural networks (ANN) (Goodfellow
et  al., 2016), are suited for deep learning
and autonomic responses in robots, such
as avoiding an imminent crash into a tree.
Machines can be designed to learn (or adapt)
in various ways, including such AI tools as
357
neural networks, expert systems, genetic
algorithms, and Bayesian inference. But the
major approach has been using ANN, which
have been developed for many decades but
until recently have only had limited success
in achieving machine learning. In recent years
there has been significant progress because
of improvements with neural network soft-
ware and hardware, which are also referred
to as deep learning and deep neural networks
(DNN). Early ANN had one layer of eight
neurons; new DNN can contain more than
150 layers of neurons, interconnected with
about a billion connections. One DNN has
60 million tunable parameters and 650,000
neurons (Krizhevsky et al., 2012; Lipson and
Kurman, 2016). They need fast and powerful
computers and advanced training algorithms
to function. As an example of the progress
of deep learning, consider that an untrained
human can categorize images (i.e., identify
the content of images) with a 5% error rate,
which was far better than machine vision
until recently (Lipson and Kurman, 2016).
• 2010: The winning ANN in a machine vision com-
petition categorized 100,000 test images with
28% error rate.
• 2011: The winning ANN categorized test images
with 25% error rate, a slight improvement.
• 2012: Deep learning emerged and the winning
ANN (a convolutional neural network) catego-
rized test images with 15% error rate, a major
improvement.
• 2013: The winning DNN categorized test images
with 11.2% error rate.
• 2014: The winning DNN categorized test images
with 6.66% error rate.
• 2015: The winning DNN categorized test images
with 3.57% error rate, beating human capability
(Lipson and Kurman, 2016).
While DNN seems to be an effective and
efficient approach to many intelligent
machine applications, nevertheless there are
many problems. For example, DNN cannot
learn by analogy, as humans and animals can
do, that objects possess common attributes.
Despite their many shapes and sizes, humans
can quickly generalize the notion of a chair
358 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
or lamp. So far, DNN has failed to learn the
concept of sameness and could not learn by
analogy the underlying concepts linking simi-
lar objects in a training set of images of those
objects (Lipson and Kurman, 2016). Also,
DNN is not transparent and the reasons for its
outcomes are often inexplicable. It does not
learn like humans. People learn by abstracting
the principles underlying a few examples,
while DNN needs massive databases to learn
(Lipson and Kurman, 2016). So far DNN,
while accomplished in machine learning, has
not provided machines with human-like intel-
ligence or cognition. It seems that machines,
such as robots, must interact with the world to
discover the intuitive physics upon which cog-
nitive inference depends.
The application of genetic algorithms
(GAs) for robot evolution provides a hybrid
case where Darwinian evolution can be used
to impel the rapid evolution of robot tech-
nology, including the evolution of the robot
mind. It is a Darwinian basis for Lamarckian
technological evolution.
GAs (Goldberg, 1989; Davis, 1991) are
mathematical algorithms that transform a
set (or population) of individual mathemati-
cal entities (typically fixed-length character
strings patterned after chromosome strings),
each with an associated fitness value, into
a new population (i.e., the next generation)
using operations patterned after the Darwinian
principle of reproduction and survival of the
fittest and after naturally occurring genetic
operations, such as sexual recombination.
GAs are run in high-speed digital simula-
tions containing entities and their environ-
ment. Millions of simulations, representing
the equivalent of millions of years of passing
time in the real world and millions of gen-
erations of living and dying, can be run in
seconds or minutes of processing time. The
GAs typically cause, over simulated genera-
tions, the progressive modification of some
structure or structures, which may then be
reflected in the behavior of the system of
which the structures are a part (as animal
behavior is a reflection of genetic structures
based on DNA). As in nature, simple mecha-
nisms can generate complex structures and
adaptive behaviors.
Genetic programming is an extension
of conventional GAs (Koza, 1992). It is a
method for automatically creating a work-
ing computer program from a high-level
statement of the problem, i.e., the program-
mer need not specify the program code. It
accomplishes this by genetically breeding a
population of computer programs using the
principles of Darwinian natural selection and
biologically inspired operations. Genetic pro-
gramming iteratively transforms a population
of computer programs into a new generation
of programs by applying analogs of naturally
occurring genetic operations. As with GAs,
the genetic operations include crossover
(sexual recombination), mutation, reproduc-
tion, gene duplication, and gene deletion.
The conventional robot selects a pre-­
programmed strategy (on the basis of infor-
mation stored in its databases, memory, and
world model, and acquired from the inter-
action of its sensors with the internal and
external environments), to meet a defined
goal or achieve a functional purpose. The
conventional approach requires complex and
costly preparation for the development of
algorithms for each specific environment in
which the robot is to operate. GAs require
goals, feedback mechanisms, and sensory
access to the environment to learn to oper-
ate effectively and efficiently. They require
minimal engineering or programming. The
designer need not be prescient about every
possible contingency to write a long series
of if-then rules (as for expert systems). The
designer does not have to expend the time
and effort to train the system, as is the case
for most neural networks and deep learn-
ing. GAs enable the simulated robot to adapt
continuously, to cope with uncertain or
changing environments, and to allow the
system (e.g., robot) to perform successfully
while becoming strategically prepared for
future circumstances. This not only reduces
the cost and exertion of programming, but
 EVOLUTIONARY PSYCHOLOGY AND ROBOTICS
enables the robot to adapt to a broader range
of environments (Schultz et al., 1993).
For example, a simulated autonomous
robot aircraft can learn, over millions of
attempts, to land safely on the simulated
rolling, pitching, yawing deck of an aircraft
carrier – or how to land on various irregular
surfaces. A simulated legged robot can learn
how to best control its legs to negotiate an
uneven terrain, and adapt to walking success-
fully over various kinds of difficult terrain.
The algorithms and physical designs (e.g.,
for legs) that are engineered in the simulated
evolutionary generations can then be built
into the real-world robot. The psychology of
the real-world robot can be advanced by mil-
lions of generations of simulated evolution of
simulated robots (Schultz et al., 1993).
If a robot had sufficient computational
power, GAs could be run within the brain of
the real-world robot. For example, if a robot
were to encounter an unexpected problem or
environment, it could pause in its movement
and run its own GAs to arrive at a solution
that optimizes or satisfies an objective func-
tion, allowing the robot to solve the problem
or survive a dangerous situation.
The basic elements of the GA consist of
representations, patterns (or structures), and
schemata (Goldberg, 1989), as follows:
• representation: a blank vector of elements (vari-
ables) which can take on numerical values;
• pattern (or structure): numerical values are incor-
porated into the representation; and
• schemata: sets of structures which are, in effect,
strategies for optimizing fitness criteria.
GAs prescribe the maintenance of a small set
of structures and provide guidelines for the
construction of new structures (Goldberg,
1989). The structures generally combine in
pairs (the equivalent of sexual reproduction) at
a rate based on their observed performance.
Each time the two structures reproduce to pro-
duce a new structure, the new structure main-
tains the optimal properties expected to result
from the initial union, while simultaneously
correcting for biases in the algorithm. By testing
359
sequential structures, a large number of
patterns, or schemata, are implicitly tested.
­
For a robot control system, for example, each
schemata may be a control strategy associated
with multiple sensor inputs.
Each structure in a relatively small set is
tested against the environment, as genetically
designed organisms in nature are tested against
the environment. Three primary operators
are used to combine the elements of two
selected structures to form a new structure
(Goldberg, 1989):
• crossover: each structure is represented as a vector.
A point along this vector is selected, and the new
structure takes all elements before (and at) this
position from one parent structure, and all of the
elements after this point from the other. Crossover
is the most used primary operator;
• inversion: crossover produces a bias toward
propagating closely linked schemata (i.e., sche-
mata with short physical distances between
elements). Inversion compensates for this bias by
causing successful schemata to become increas-
ingly closely linked physically; and
• mutation: over successive trials potentially impor-
tant values in particular positions may disappear
from the population of structures. A low mutation
rate periodically reintroduces lost values.
Information about large amounts of schemata
is processed at each stage of the simulation,
with subsequent trials making use of this
information, which is genetically embedded in
the structures. The structures reproduce prob-
abilistically and the probabilities are assigned
by the algorithm on the basis of observed
performance against fitness criteria. Over suc-
cessive trials, the expected number of occur-
rences of schemata mirrors their performance,
e.g., the ability of a robot to walk, swim, or
play tennis against humans. Schemata that
have demonstrated to be high-performance by
surviving trials must reproduce themselves
while new structures are also being generated
and tested. There is an algorithm for assigning
trials to structures which test schemata.
This algorithm must maximize current fit-
ness while also exploring new schemata.
Thousands of generations of simulated entities,
360 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
such as robots, are born, live, reproduce, and
die. The equivalent of millions of years of natu-
ral evolution takes place in brief computer-
processing time (Goldberg, 1989).
The compressed Darwinian evolution of
a robot, including that of its mind and psy-
chology, can be effected in the context of the
Lamarckian evolution of computer simulation
technology.
TECHNOLOGY AND PSYCHO-SOCIAL
FACTORS
Evolutionary psychology provides an account
of the psycho-social gestalt and psycho-social
factors concerning technology, not just the
efficacy of the technology. It will have a sig-
nificant influence on the public’s acceptance
of autonomous robots. As a consequence of
our psychology, one death is a tragedy while a
million deaths is a statistic, as the Soviet dicta-
tor Joseph Stalin is alleged to have said. For
example, a few isolated accidental deaths
caused by autonomous cars or other robots,
which are widely publicized by the media,
might elicit an exaggerated emotional response
by the public, slowing or stifling the develop-
ment of the technology. The National Highway
Traffic Safety Administration (NHTSA)
reports nearly 37,000 deaths in the United
States from automotive accidents during 2018,
despite major improvement in vehicle safety
driver-assist systems. The fatality rate per
vehicle mile traveled (VMT) has decreased
greatly over the decades with improving auto-
motive technology and infrastructure, but at
the same time the number of vehicle miles
traveled increased greatly (e.g., with more
miles traveled per vehicle and more vehicles
on the road), contributing to the number of
deaths. But the public has become inured to
these deaths (Evans, 2008). More to the point,
according to NHTSA there were 26 deaths in
the United States from automotive accidents
in 1899 when there were about 8,000 horse-
less carriages. By 1903 there were more than
100 deaths. If social media existed at the time
as it does now, generating public outrage over
the accidental automotive deaths, perhaps we
would still be riding in horse-drawn carriages.
The goal for autonomous (robotic) vehicles is
zero deaths (National Safety Council, 2019).
The goal may not be achieved, but automotive
casualties could become as rare as deaths from
sharks (about six per year).
Psycho-social factors (as opposed to, for
example, engineering or technical factors) are
critical in the development and acceptance of
most – or all – technologies. For example, to
compensate for a shortage of suitable wood in
the 19th century, some pencils were designed
to be sheathed with paper instead of wood.
Functionally, it worked well. But psychologi-
cally people preferred to sharpen their pen-
cils by whittling them with a knife or a pencil
sharpener, rather than peeling the paper. The
paper pencil eventually became defunct, except
for special-purpose applications (Teich, 2009).
New technologies are now critically evalu-
ated in the context of aesthetics, such as
environmental and design considerations.
Developers of technology, such as scientists
and engineers, are influenced, consciously or
unconsciously, by the organizational and soci-
etal culture in which they reside, which is a
manifestation of evolutionary psychology. The
institutionalization of science and engineering
means that once a decision (even a bad one)
is made concerning the development and com-
mercialization of a technology, the momentum
of the organizational bureaucracy and the pro-
cesses of systems engineering drive the prod-
uct to its conclusion (Teich, 2009). Adverse
unintended consequences can ensue.
TECHNOLOGY AND UNINTENDED
CONSEQUENCES
Ancient legend tells us that Prometheus
defied Zeus and brought technological
knowledge, in the form of fire and the skill of
metalwork, to the human race. Zeus then
 EVOLUTIONARY PSYCHOLOGY AND ROBOTICS
punished him for this by having him bound
to a rock while a huge eagle ate his liver
every day, only to have it grow back to be
eaten again the next day (Cartwright, 2013).
New technology when first introduced can
have serious and dangerous flaws. For exam-
ple, early methods of vaccination for small-
pox could cause smallpox – and death (Teich,
2009). Manufacturers of early high-pressure
steam engines often used inferior materials,
and poor design and construction contributed
to major malfunctions (Teich, 2009). Even
well-built engines needed improved technol-
ogy to avoid catastrophic failures. Aside
from the engines, early railroads needed better
technology for bridges and tracks to avoid
death and destruction (Teich, 2009). Many
technological advances had unexpected and
unwanted side effects. Associated with a tech-
nological advance having major benefits
might be unintended consequences causing
death and destruction. Our ancestors gener-
ally did not abandon the flawed technology
but persisted in trying to improve it, either
mitigating the adverse consequences or
accepting them as a tradeoff for the benefits.
There is a tension between new technol-
ogy and the preservation of individual and
societal values. The new technology may
threaten the values of some, while efforts to
mitigate the adverse consequences of the new
technology may threaten the values of others.
Some people might then conclude that tech-
nology is inherently hostile to human values.
Technology has a direct impact on values
because it generally increases the range of
choice and opportunity, and one’s values
cause one to make certain choices and pursue
particular opportunities. New technology can
lead to a change in individual or cultural val-
ues by either bringing some previously unat-
tainable goal within the realm of choice or
by making some values easier to implement
(Teich, 2009).
Ultimately, intelligent robots might drive
humans to extinction maliciously or unin-
tentionally by competing more successfully
for limited resources of matter, energy, and
361
territory. Genetics, nanotechnology, and robot-
ics all offer the possibility of self-replicating
technology – autopoiesis. It is necessary to
examine the potential adverse consequences
of new technology, for much of technology
has adverse consequences. Often the benefits
of the new technology exceed its adverse
consequences, or its disadvantages can be
mitigated by improving – evolving – the tech-
nology. Cultural and technological systems
do not develop independently; the two evolve
together in complex feedback loops, wherein
each drives, restrains, and accelerates change
in the other. Darwinian evolutionary psychol-
ogy underlies human culture and its values,
while Lamarckian evolutionary psychology
will underlie robot culture and values.
AUTONOMOUS INTELLIGENT
ROBOTS: A DISRUPTIVE AND
TRANSFORMATIVE TECHNOLOGY
The first generation of fully autonomous
intelligent robots will be vehicles: autono-
mous cars, trucks, and buses. This will be a
disruptive and transformative technology,
whether for civilian or military applications.
The technology will be disruptive because it
will cause old industries and companies to
vanish while new ones emerge. With creative
destruction, jobs lost in one industry will be
replaced by new jobs in another. The technol-
ogy will be transformative because it will
change society. The robots will alter military
tactics, strategy, and doctrine; the national
economy and power; political policy; cul-
tural values and societal structures; and how
and where people live, work, and play.
As an example of a disruptive and trans-
formative technology similar to that of
autonomous vehicles, consider the advent of
the horseless carriage at the turn of the 19th
century. The older technology at the time,
the horse and buggy, was a kind of intelli-
gent vehicle in that the horse was sufficiently
smart to avoid objects in the road, and could
362 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
even be autonomous, e.g., if the driver was
drunk or fell asleep, the horse could find its
way home. Horses pulling milk-delivery wag-
ons learned their routes and did not require
guidance from a human driver. The horse and
buggy (or cart) was an ancient, well-known
technology, with support from known goods
providers (e.g., buggy manufacturers, tack-
equipment manufacturers, and horse breed-
ers), service providers (e.g., blacksmiths,
livery stables, veterinarians), and infrastruc-
ture providers (e.g., horse troughs and barns).
The technology was acceptably reliable and
able to traverse bad roads. Problems with the
technology included: daily storage mainte-
nance (barn, food, water, grooming, etc.) for
the horse, whether it was used or not; a grow-
ing manure problem in urban areas; and the
cost to buy and maintain it.
The earliest horseless carriages were an
unknown technology: unreliable, slow, and
very expensive, with no infrastructure (e.g.,
no suitable roads, available parts, fuel sta-
tions, mechanics, rules and regulations, or
an understanding of the technology by the
public). Some advantages over the existing
technology included: no daily maintenance if
not used; faster; more easily and compactly
stored; more compact fuel; no manure; and
status for early adopters. But the cost of the
new technology was still too expensive for
most people. Early cars and manufactur-
ers proliferated, with varied technology and
designs (e.g., should it be a steering lever or
a wheel; an internal or external combustion
engine or electric motor?). As with natural
evolution during the Cambrian period when
nature was experimenting with manifold
new designs, new body plans, for complex
animals, automotive designs evolved rapidly
during the first two decades of the automo-
tive age. Around the turn of the century there
were more than 2,000 automotive manu-
facturers across the United States, of which
many were startups in home garages, creat-
ing a proliferation of automotive designs
(Wikipedia, 2019b). But as in the Cambrian
period with the evolutionary winnowing of
organism design, by 1920 there were just
100 US ­manufacturers with their automotive
designs converging to new industry stand-
ards. By 1929 only 44 automotive manufac-
turers remained in the United States, which
decreased to 11 by 1976. In the early 1900s,
only a few thousand cars were sold annually.
For example, Ford sold 1,700 cars in 1904, its
first year of business. After introducing factory
mass production in 1913 and thereby drasti-
cally reducing the cost of cars, Ford sold over
a million cars in 1920 (Wikipedia, 2019a).
The horseless carriage was a disruptive
technology (The Economist, 2015). Across
many sectors of the economy it drove the
creation of new technologies, industries,
and infrastructure, such as the oil industry,
asphalt and concrete roads, bridges, tires, fac-
tory mass production, etc. Many, but not all,
horse- and buggy-related goods and services
firms were driven out of business. The horse
and buggy market shriveled but did not disap-
pear entirely. To this day they remain common
in Amish areas of Pennsylvania and other
states. Most new entrants in the automotive
industry also disappeared as the technology
advanced and designs, production, products,
infrastructure, and marketplace standard-
ized and stabilized. Punctuated equilibrium
(Gould, 1983), as it does in natural evolution,
caused occasional disruptions to the industry,
with paradigm shifts due to new technology,
government regulation, or customer expecta-
tions. The current technology paradigm shift
is driven by the technologies for energy, elec-
tronics, and ergonomics, including electric
propulsion to reduce pollution, driver-assist
systems to increase safety, and autonomous
vehicles.
The horseless carriage was a transforma-
tive technology (The Economist, 2015). It
changed cultural values and societal struc-
ture. It caused the fashion industry to design
clothing to accommodate the car. It caused
the rise of suburbs, increased travel among
the populace, and increased the homogeni-
zation of society. It gave rise to industry in
place of agriculture and city living in place
 EVOLUTIONARY PSYCHOLOGY AND ROBOTICS
of rural living. It abetted the influence of
massive new corporations. It changed family
dynamics with teenage and women drivers;
the rise of teenage influence and their dating
customs, with the car as bedroom-on-wheels;
and the loss of power by the husband and
father. It increased family wealth, freedom,
and mobility, and contributed to the rise of
the middle class. It caused many casualties
due to crashes. In the long term it caused
geopolitical and environmental changes,
with the rise of oil cartels, religious and tribal
conflicts in oil-rich lands, wars over oil, and
environmental degradation leading to climate
change and the potential destruction of much
of the Earth’s species and living space for
humans.
Autonomous intelligent vehicles have a
limited but sufficient domain of knowledge
and ability, but they will serve as a kind of
Ur-technology for all kinds of autonomous
robots. Their success will serve as the basis
for developing and technologically evolving
autonomous robots into many robot ‘species’
able to perform many jobs across all eco-
nomic sectors. Biomimetic robots will prolif-
erate, including humanoid and multi-legged
chimera robots, like centaur robots; zoomor-
phic robots with forms like snakes, lizards,
fish, octopuses, primates, birds, insects, etc.
Disruption will spread across all sectors of
the economy, and changes in how people live,
work, and play will continue for the rest of
the century. The technological evolution of
robots and the AI for their minds will con-
tinue unabated across the globe despite any
efforts of one nation or another to slow or
stop its progress. Technology that is feasible
and useful is hardy and always emerges like
a weed in a crack in the sidewalk. Perhaps
human psychology will be changed in the
future by genetic engineering or some other
technology and will be better able to cope
with the results of the mind’s creations
(Smith, 2019). It seems that the best way
to avoid the proverbial robot apocalypse is
for people to be at least as smart as their
robots.
363
ROBOT EMOTION
There are development projects to give robots
emotion (Hall, 2017), or the appearance to
humans of emotion. The robots are designed
to interpret certain internal states as the equiv-
alent of human emotions, such as happy, sad,
angry, etc., and to express those emotions with
facial expressions and body language. The
idea is for robots to be better able to interact
and work with people. Human emotion is
based on algorithmic bio-chemical processes
which evolved to solve specific adaptive
problems, e.g., the fear of being eaten and the
pleasure of eating food and having sex
(LeDoux, 2012). Emotions are also an impor-
tant part of generating and controlling human
behavior, as employed regularly by priests and
politicians (Zaki, 2013). Rationality and logic
are not necessarily what motivates people to
achieve goals. Human emotion is an important
mechanism for making certain kinds of deci-
sions, such as when time is critical or data are
unavailable and objective analysis is not feasi-
ble. Computers can perform detailed analyses
much faster than people and until recently
emotion was largely ignored in the develop-
ment of intelligent machines. Robots, how-
ever, will be moving about and interacting
with the environment. As humans do, they will
encounter sudden life-and-death situations in
which decisions cannot be made rationally,
where emotion, or its robot equivalent, may be
called upon to make the decision.
Emotions can represent an aggregation of
certain critical memories (LeDoux, 1996),
allowing for a quick, nearly autonomic, fight-
or-flight response (e.g., run from the tiger!).
Emotions have a profound effect on cognition
(Damasio, 2003), determining what is good or
bad, loved or hated, important or unimportant,
beautiful or ugly, remembered or forgotten.
Humans and other animals seem to have simi-
lar, if not identical, emotions because of the
similarity of the physiological neural system
and the evolution of the portions of the brain
involved with emotions (LeDoux, 2012).
364 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
It seems likely that emotional-state variables
such as fear, hate, and love can be generated
by computational mechanisms and used for
generating behavior in artificially intelligent
systems. Computer models have been used to
model certain emotional processes such that
the machine seems to behave emotionally.
Even if the underlying sources of emotion
were different in machines, if the manifesta-
tion were the same as for humans, the con-
clusion from a Turing test (Moore, 2003)
designed to test robot emotions would be that
robots have emotions. If a machine behaves as
if it is afraid or happy, who is to say it is not?
ROBOT CONSCIOUSNESS AND
FREE WILL
There is no universal agreement among cogni-
tive scientists as to the nature or meaning of
consciousness (Edelman and Tononi, 2000;
Graziano, 2013). An aggregation of various
dictionary definitions of consciousness is: ‘the
state of being aware of oneself and one’s rela-
tionship to the world; perceiving, apprehend-
ing, or noticing with a degree of controlled
observation; capable of or marked by thought,
will, design, or perception’. Our preferred
definition of consciousness is the state of
meta-perception – the perception of percep-
tion. Self-awareness is quite limited in humans
in that we are unaware of most of our physi-
ological states. Most of our physiological and
mental processes take place without our being
aware of them, such as what is going on in our
gastro-intestinal system or pancreas. Most of
the cells and the DNA in and on our body are
not even ours, belonging to bacteria, fungi,
worms, mites, mitochondria, and other sym-
bionts and parasites, all of which we are
blissfully unaware unless there is a gross mal-
function. Experiments of seemingly conscious
behavior indicate that it may be caused sub-
consciously and then retroactively attributed
by the mind to a conscious decision (Libet,
1993; Ayan, 2018; Koch, 2018). This would
be just one of the many delusions we con-
stantly create as the mind frantically tries to
fill the gaps in perception caused by incom-
plete sensory information from imperfect
senses, inattention, and environmental noise,
along with missing knowledge about the envi-
ronment, uncertainty about the behavior of
others, and the inability to know everything
about the past or to predict the future (Libet,
1993; Ayan, 2018; Koch, 2018).
A robot’s physiology and brain would be
different from that of humans. The techno-
logical evolution of robot psychology could
lead to a robot consciousness that is superior
to that of people, where the robot is fully self-
aware. At all times the robot could be aware
of everything taking place within its body
and the state of its physiology and mind.
It is possible that a form of consciousness
can emerge from a robot that has a suitable
representation of the world. A robot that can
build and maintain a world model that repre-
sents objects, events, entities, situations, and
relationships should be able to include itself
as one of the entities. A self-aware system can
keep track of itself, of its external and internal
states and dynamics. A self-aware system can
be aware of how it should respond to things –
how it ‘feels’ about things. Events or objects
can be labeled with emotional state variables,
such as fear, hate, and love. Pain-state vari-
ables can indicate to the robot that it is dam-
aged. An emotional response may not always
be desirable in robots or humans because the
consequent behavior may be excessive or inap-
propriate to the situation. In life-threatening
situations it may be quite appropriate. A robot
could have a world model with objects that
have assigned emotional-state variables. If the
robot observes an object labeled with a high-
fear-state variable, it might exhibit fear by
increasing its engine speed, assuming a defen-
sive posture, or increasing its visual attention
and sensory awareness. It might articulate its
fear, as the intelligent computer HAL did in
the movie version of Arthur C. Clarke’s ‘2001:
A Space Odyssey’ when it was being disman-
tled: ‘Stop, Dave, I’m afraid’.
 EVOLUTIONARY PSYCHOLOGY AND ROBOTICS
As previously defined, robot intelligence is
not the same as self-awareness or conscious-
ness. And robot consciousness is not the
same as what is usually considered free will,
although the meaning of free will is elusive,
and the perception of free will in humans
may be a delusion (Gazzaniga, 2011; Harris,
2012). Self-awareness can be designed into
the robot, or it may become an emergent
property as robot minds become more com-
plex. Free will, or the perception of free will,
may also emerge from complex robot minds.
Large, complex neural networks make cor-
rect decisions even now that are unexplain-
able by human observers. Advanced robots
may make appropriate decisions and act
accordingly without anyone understand-
ing how or why. When asked, the robot
may answer ‘I decided it was the right thing
to do’. In a future Turing test for free will,
human observers may conclude that robots
have free will. When robots are perceived to
be intelligent, conscious, and have free will,
on what justification can people treat them as
machines or slaves?
AUTONOMOUS MILITARY ROBOTS
The US military is still refining its definition
of an autonomous system. For example,
DOD Directive No. 3000.09 (21 November
2012) defines an autonomous weapon system
as a weapon system that, once activated, can
select and engage targets without further
intervention by a human operator. For many
years a US government working group called
ALFUS (Autonomy Levels for Unmanned
Systems) met regularly to try to define
autonomous systems based on three key vari-
ables (i.e., three dimensions): mission com-
plexity, environmental difficulty, and human
interface. They were not entirely successful.
Likewise, during the 1990s various aero-
space companies and government agencies
attempted to define autonomous systems,
often with lists of functional descriptions,
365
such as a 10-level scale, from no autonomy
to full autonomy. There is actually a contin-
uum of autonomy and the levels discretize
the continuum:
1 System offers no assistance – operator must do
everything
2 System offers a complete set of action alterna-
tives to operator
3 System narrows the action alternatives to a few
4 System suggests a selection
5 System executes a selection if operator approves
6 System allows operator a restricted time to veto
before automatic execution
7 System executes automatically, then necessarily
informs operator
8 System informs operator after execution only if
operator asks
9 System informs operator after execution – if
system decides to
10 System decides everything and acts autono-
mously, essentially ignoring the human
In recent years, as the technology for autono-
mous cars advanced rapidly, the automotive
industry and the Society for Automotive
Engineering defined six levels of autonomy.
A few slightly different variants of the six
levels are used by various autonomous vehi-
cle developers.
Level 0: no autonomy
Level 1: a few functions, such as automated cruise
control, are automated
Level 2: the driver, while always monitoring the car,
does not have to steer or brake or accelerate
under some driving conditions
Level 3: the vehicle can drive mostly autonomously,
but the driver must always be alert to take over
as needed (vehicle may check the driver’s alert-
ness regularly)
Level 4: the vehicle can operate fully autonomously
at least in some environments, e.g., on an inter-
state road, without the driver having to be alert
while in that environment (but a driver must
always be in the vehicle)
Level 5: the vehicle can operate fully autonomously
without any human or driver in the vehicle or con-
trolling or monitoring the vehicle from a distance
In 2012 the US Department of Defense issued
a directive outlining steps to ensure that
366 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
autonomous weapons function as anticipated
and to minimize failures that could lead to
unintended engagements, or to loss of con-
trol of the system. The Defense Science
Board (DSB) in 2016 conferred to improve
the future adoption and use of autonomous
systems, declaring that they had significant
military value. They recommended accelerat-
ing the military’s adoption of autonomous
capabilities. The DSB’s definition was that an
autonomous system has the ability to inde-
pendently compose and select among differ-
ent courses of action to accomplish goals
based on its knowledge and understanding of
the world, self, and situation.
Along with the military’s accelerated adop-
tion of autonomous systems, there are grow-
ing efforts to ban autonomous lethal robots for
military (and law-enforcement) applications.
Organizations like Human Rights Watch
and the Harvard Law School’s International
Human Rights Clinic are alarmed about the
ethics of lethal autonomous robots. Calling
them ‘killer robots’, opponents urge an inter-
national treaty that would absolutely prohibit
the development, production, and use of fully
autonomous weapons. Opponents say that
killer robots will never possess the human
judgment that limits civilian casualties dur-
ing war and such machines could not be held
accountable for war crimes. ‘Never’ is a very
long time, and it seems that their developers
and users could be held liable for war crimes,
similar to the liability issues when autono-
mous cars cause accidents or, in the future,
be used to commit crimes or terrorist acts.
Civilians have always been, and still are,
major casualties of war, and robot soldiers
can be designed to reduce battlefield death,
destruction, and atrocities. It is not certain that
autonomous military robots would be worse
than human combatants. It depends on how
the robots are designed, programmed, and
trained. Some say machines can never have
compassion, but, again, ‘never’ is a very long
time. In any event, many humans, including
combatants, are sociopaths or suffer from
clinical narcissistic personality disorder and
consequently lack compassion and empathy
(Brazier, 2018). While inherently saving
the lives of the human soldiers they replace,
autonomous robots can be programmed to be
more humane and rational than humans and
save civilians who would be otherwise killed,
maimed, or raped by emotional, rampaging
human combatants. On the other hand, robots
can be designed to kill everyone efficiently
without empathy or remorse.
Those opposing combat robots say that
autonomous machines will not be able to
apply human judgment in the application of
lethal force, or will not be constrained by the
laws of war. But even humans cannot, or do
not, always distinguish between combatants
and non-combatants, or between friendly and
enemy forces. The laws of war, such as they
are, are frequently ignored. A robot could
recognize that a potential target is already
wounded or trying to surrender and respond
appropriately. For example, DOD Directive
3000.09 (2012) concerning autonomy in
weapons systems requires that command-
ers who authorize autonomous weapons
systems must do so in accordance with the
laws of war, treaties, safety rules, and rules of
engagement, and that the autonomous weap-
ons systems must allow that such constraints
be imposed by their human commanders.
Opponents say robotic warfare is a slip-
pery slope leading to an uncertain, poten-
tially apocalyptic future. Even if the US DOD
were conscientious in deploying autonomous
robots, other nations or non-state actors may
not be the same. Autonomous weapon sys-
tems are not inherently unlawful or unethical.
In any event, the development and deploy-
ment of autonomous weapon systems are
inevitable, and any attempt at a global ban
will likely be ineffective. Given the historical
failure of international treaties to ban other
seemingly abhorrent means of killing, it is
unlikely to succeed for lethal robots as well.
It would likely be easier to get an interna-
tional moratorium on their deployment or an
agreement on how to constrain their behavior
than to prohibit them entirely, just as nuclear
 EVOLUTIONARY PSYCHOLOGY AND ROBOTICS
weapons have not been used again in combat
and lethal gas has been used rarely.
ROBOT LAWS, ETHICS, AND
MORALITY
Natural selection has provided humans with a
set of rules and constraints for behaving
socially, such as not killing or eating one’s
children, or behaving fairly with others within
one’s family or tribe. These behavioral rules
are limited and not always effective, so
humans have devised larger sets of behavioral
rules, such as laws, ethics, and morality. These
terms have many different definitions, with
often an overlap between ethics and morality.
These definitions distinguish the rules.
Laws are behavioral rules which, when
violated, can be punished by society with the
loss of one’s life, freedom, or possessions.
Ethics are rules of behavior which, when
violated, can be punished with the loss of
one’s profession (if it includes ethical rules),
or ostracism by business associates, friends,
or social groups. Morality is behavior which
mitigates pain and suffering, as in the parable
of the Good Samaritan, which was tested
in a well-known psychological experiment
(Darley and Batson, 1973). However, moral-
ity does not require a belief system such as
religion; just the objective realization that
humankind benefits when the members of
its species refrain from causing pain and suf-
fering and strive to mitigate such suffering
when it is caused by the brutality of nature or
circumstance.
Likewise, autonomous robots, including
driverless vehicles in the near term, will need
to conform to legal, ethical, and moral behav-
ior, as well as understanding the human sense
of these rules and behavior constraints (Lin
et al., 2012). Robots will need to become what
are called Artificial Moral Agents (Wallach
and Allen, 2009). Robots which violate human
or robot laws, such as laws against murdering
humans or other robots, may lose their lives
367
by their complete destruction (i.e., execution)
or re-­programming, depending on whether the
cause of the deviant behavior is known and
repairable. Confining a future criminal robot
in a prison would only be justified if the robot
were deemed to be a conscious entity, worthy
of human-like treatment, and it did not want
to be re-­programmed to have its mind altered.
In order to impose behavioral constraints
on robots, science-fiction author (and scien-
tist) Isaac Asimov devised the Three Laws of
Robotics in 1942 in a short story, which was
later used in many more of his stories (Asimov,
1950). The Three Laws, later increased to
Four Laws, were a set of behavioral rules that
were ‘laws’ in the sense that they were pro-
grammed constraints on a robot’s behavior.
The Laws were designed to prevent robots
from harming humans while performing their
functions, and Asimov later added a fourth (or
zeroth) Law:
1 A robot may not injure a human being or, through
inaction, allow a human being to come to harm.
2 A robot must obey the orders given to it by
human beings, except where such orders would
conflict with the First Law.
3 A robot must protect its own existence as long
as such protection does not conflict with the First
or Second Law.
4 A robot may not harm humanity, or, by inaction,
allow humanity to come to harm.
The Laws are often cited as templates for
constraining robot behavior, to serve as a
basis for robot ethics and morality. But the
point of the stories (e.g., I Robot) was that
the Laws often failed for various unforeseen
reasons, leading to unintended consequences.
The narrative puzzle was to determine the
cause of the Law’s failure. For example, why
would robots, with their behavior seemingly
constrained by the Four Laws, enslave
people? Let’s say that the robots decide,
among themselves, that the greatest threat of
harm to humanity is from humanity itself.
Thus, in order to protect humanity from harm
they must enslave humans for their own
good. Under strict control of the robots,
368 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
humanity, while enslaved, would be p­ revented
from causing its own extinction, from coming
to physical harm. The harm to humanity
losing its freedom is either not apparent to the
robots or is considered of relatively minor
importance relative to physical harm.
Legal, ethical, and moral dilemmas are
being deliberated and debated now for the
advent of autonomous vehicles (Herrman
et al., 2018). Insurance companies are among
those considering who will be legally and
financially responsible for accidents involv-
ing autonomous vehicles. Who will be legally,
ethically, and morally responsible for the
decisions that will be made by any intelligent
autonomous robot, including lethal military
robots? A typical question is: if your driver-
less car were to encounter a bus full of school-
children coming toward you on the wrong side
of a road near the edge of a cliff, how should
it react? Should it make a decision to swerve,
preventing a collision with the bus and spar-
ing the children but putting your life at risk?
Or should it collide with the bus if there’s a
greater chance of you surviving?
Recently, there has been much contempla-
tion of the famous trolley problem, a moral
and psychological dilemma with several vari-
ations, as it relates to humans and autono-
mous vehicles or other robots (Shenhav
and Greene, 2010; Bohannon, 2011). In one
variant of the narrative, a runaway trolley is
about to kill five workers on a track. You are a
bystander near a switch that would cause the
trolley to change tracks and avoid killing the
workers. However, there is one worker on
the other track who would be killed. Would
you close the switch and kill one person to
avoid killing five people? What if you are
on an overpass watching the runaway trolley
below. You can save the five workers if you
throw a fat woman, standing next to you on
the overpass, onto the track to stop the trolley.
Would you throw the woman onto the tracks?
You would kill the woman to save the five
workers.
Most people surveyed would close the
switch to save five people but not physically
throw a person off the overpass to achieve
the same result. The reason for this illogical
result was philosophically debated exten-
sively, but a scientific experiment provided
an explanation. It seems there is a mental
distinction, embedded in different areas of
the brain, between an intended harmful act
involving physical force, like pushing a per-
son off a bridge, and an act in which harm is a
side effect, like the result of closing a switch.
Functional magnetic resonance imaging
(fMRI) indicates that, for most people tested,
a part of the brain involved with rational-
ity (the prefrontal cortex) would close the
switch, while a part of the brain involved with
emotion (the amygdala) would prevent them
physically throwing a person off the bridge.
Depending on the scenario, one or the other
would dominate the decision and determine
the consequent act (Greene, 2012).
A robot, or autonomous car, could decide
the trolley problem or an equivalent encoun-
tered in the real world on a purely objective,
rational basis. The metrics for determining
the rationality of the decision may go beyond
a simple numerical head count, to include
the value to society of the individuals. Are
the deaths of five laborers worth more than the
death of one individual who is the 21st cen-
tury equivalent of Einstein, Newton, Lincoln,
or da Vinci? What if the five workers are all
single but the person on the bridge is sup-
porting a family of six? What if the five
workers are all old but the one individual is
young? With the information that it has or
can quickly acquire, the robot can swiftly
evaluate the alternative decisions against
the metrics and make an objective decision.
However, the metrics used as the bases for
these kinds of decisions need the seeds of
wisdom, whether human or machine, for
their fruition.
Objective decisions, however, may not
always be commercially viable. People may
avoid buying or riding in autonomous vehi-
cles, or employing autonomous robots, if
they felt that, in an emergency, their lives may
not be considered as important as others’.
 EVOLUTIONARY PSYCHOLOGY AND ROBOTICS
The algorithm for determining who should
be saved and who should be allowed to die
might then be skewed a bit, if not completely,
toward the occupants of the vehicle or the
owner of the robot, offering a higher prob-
ability of surviving an adverse incident. Or
autonomous vehicle software may offer alter-
native algorithms to users, allowing them to
select a preference for a higher probability of
survival in an accident (i.e., a selfish choice)
or survival based on objective criteria, which
may provide the passengers with a lower
probability (i.e., an altruistic choice). Human
psychology again becomes a factor in deter-
mining robot psychology.
Autonomous robots can know the laws
they are required to obey from ‘birth’, as
soon as they emerge from the assembly line.
The laws may be the same as, or different
from, those constraining human behavior.
They will be codified rules with specified
penalties, but it is likely the penalties will
be different for robots. If needed, autono-
mous robots can have immediate internal or
external access to, and knowledge of, all
federal, state, and local laws, as well as judi-
cial cases and precedents. However, for a
robot to have more than superficial knowl-
edge of lawful behavior, and especially ethi-
cal and moral behavior, it will need more
than access to a vast knowledgebase – just
as humans do.
People are born with evolved behavioral
rules that can be attributed to a moral and
ethical code of conduct, such as manifested
in their behavior with parents and siblings.
They are not born knowing any societal
laws. Robots can be ‘born’ with lots of pre-
programmed rules, legal, moral, and ethical.
This is a top-down approach to robot behav-
ioral constraints, where rules are pre-encoded
in software, like Asimov’s Laws of Robotics.
Over the millennia people have devised,
for example, the Code of Hammurabi, the
Magna Carta, the Ten Commandments, the
US Constitution, the Rules of Professional
Conduct of the American Bar Association,
and the Laws of the State of Maryland.
369
A robot can have knowledge of the local
­customs of how to set a dining-room table
and the proper way to greet guests and offer
them refreshments; or how to sell menswear
in a retail store without cheating customers. In
general, a top-down approach for robot legal,
ethical, and moral behavior would require
algorithms to guide robot behavior, i.e., the
robots would need to know how to apply
the rules in any given situation. Programmers
cannot foresee every contingency, so the
robot must also learn from experience how
and when to apply the rules of law, ethics,
and morality.
Unlike robots, humans cannot be born
with these rules implanted in their minds, at
least currently. People often have difficulty
interpreting these rules and knowing how
to behave within their constraints. Many of
the rules are not instinctual. They must be
acquired through years of experience – the
bottom-up approach.
The bottom-up approach for robots to mas-
ter their rules of behavior – legal, ethical, and
moral – requires time for them to acquire
experience interacting with family, peers,
and strangers, as do humans. While the time
required to learn from experience may be less
than that for humans, it still may take sev-
eral years. Robots will need a combination
of top-down and bottom-up approaches to
fully learn the norms, standards, and cultural
aspects of moral and ethical behavior as prac-
ticed by humans, and the behavior required
of robots (Lin et al., 2012).
In the bottom-up experiential approach
emphasis is placed on creating an environ-
ment where the robot explores courses of
action and is rewarded for behavior that is
legally, ethically, and morally praiseworthy.
Thus the robot learns and adapts through its
experiences. Unlike the top-down approach,
in the bottom-up process the rules are discov-
ered or constructed, not explicitly mandated
by fiat. The given behavioral rules, however,
serve to provide a framework and a context
that guides and simplifies the experiential
rule-learning process.
370 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
EVOLUTIONARY PSYCHOLOGY OF
ROBOTS: FUTURE
Over the coming decades, to the end of this
century, robot intelligence will evolve techno-
logically, a Lamarckian process that, like
Darwinian evolution, is likely to be punctuated
in its progress, with intervals of relatively fast
and slow change. New discoveries and inven-
tions will impact robot software and hardware.
Robot intelligence is likely to increase with
new algorithms and processors, and robot psy-
chology will evolve. Robot performance is
likely to improve with new technology for
legs, wheels, arms, hands, and sensors. Robot
capabilities in 2099 are unpredictable, except
to safely speculate that they will be orders of
magnitude greater than they are now.
Ubiquitous robots and other intelligent
machines can provide humankind with a uto-
pia or dystopia. The technologies of genetics,
nanotechnology, and robotics have at least one
thing in common – they all offer the possibility
of self-replication. Autopoiesis is one poten-
tial promise or danger of intelligent machines,
whether they are as intelligent as a human or an
amoeba, or the size of a human or an amoeba
(such as nanobots which will repair damaged
organs). The autonomous autopoietic system
is self-sustaining through replication, just as
organisms have been self-sustaining on Earth
for nearly four billion years. The power of
replication is astounding: on Earth there is an
unbroken chain, sustained through intermit-
tent global catastrophes, for almost a third of
the age of the universe; from the first living
aggregation of molecules to us.
NASA’s astrobiological definition of life
is that it is a ‘self-sustained chemical system
capable of undergoing Darwinian evolution’.
This may be modified to accommodate robotic
lifeforms: life is a self-sustained system capa-
ble of adapting to a changing environment.
Autopoietic robots smart enough to create
new technology could hasten the evolution of
robot psychology. Robot autopoiesis can be
dangerous, but any guidelines, or even laws,
to control the development of self-replicating
robots could be intentionally or unintention-
ally violated, as has happened with geneti-
cally modified crops. Before the end of this
century, autonomous robots will become
ubiquitous.
Over the coming decades, scientific dis-
coveries and technological invention from
manifold fields could affect the evolution of
robot minds. For example, there are thou-
sands of human brain organoids now being
grown in nutrient solutions for study, some of
them on the International Space Station (Qian
et  al., 2019; Spectrum, 2019). These were
originally human skin cells that were trans-
formed into stem cells, which were trans-
muted into neurons. The neurons clumped
into spherical organoids which formed inter-
nal structures like tubes and emitted synchro-
nized neuronal firing patterns. These are not
(yet) human brains, but there is some concern
that they may become sentient and feel pain,
or otherwise suffer. It is easy to imagine that
conscious brain organoids might some day be
integrated in the brain of a robot, creating a
hybrid entity with a mind having both human
and machine structure and psychology. Initial
experiments linking brain organoids to a
robot have already been conducted. A multi-
legged spider-like robot received neuron sig-
nals from a brain organoid causing it to walk.
The robot sent sensory information, such as
when it encountered a wall, back to the brain
organoid.
Our minds and their evolved psychology,
‘designed’ over eons for survival in their nat-
ural environments, are imperfectly designed
to function properly in our current complex
societies. Technology helped us survive the
nastiness of nature for many millennia. We
now need help to survive the nastiness of our
human-made environment. Technology will
propel the rapid evolution of robot minds
and their consequent psychology. Despite the
risks, such robots may be the best hope for
humanity to alleviate ignorance, poverty, and
suffering, and to allow humans to become
more humane.
 EVOLUTIONARY PSYCHOLOGY AND ROBOTICS
REFERENCES
Albus, James S. (2001), Engineering of Mind,
Presentation based on Albus, James S. and
Alexander M. Meystel (2001), Engineering of
Mind, New York, NY: John Wiley & Sons, Inc.
Albus, James S. and Alexander M. Meystel
(2001), Engineering of Mind, John Wiley &
Sons, Inc.
Ardiel, Evan L. and Catharine H. Rankin (2010),
An Elegant Mind: Learning and Memory in
Caenorhabditis Elegans, Learning and
Memory, Brain Research Centre and Depart-
ment of Psychology, University of British
Columbia, Vancouver, British Columbia V6T
2B5, Canada, 17:191–201 # 2010 Cold
Spring Harbor Laboratory Press, ISSN 1549-
5485/10; www.learnmem.org
Asimov, Isaac (1950), I Robot, New York, NY:
Doubleday.
Aunger, Robert (2002), The Electric Meme: A
New Theory of How We Think, New York,
NY: The Free Press.
Ayan, Steve (2018), There Is No Such Thing as
Conscious Thought, Scientific American,
December 2018. Retrieved from https://
www.scientificamerican.com/article/there-
is-no-such-thing-as-conscious-thought/
?print=true (Accessed 10 July 2020)
Bohannon, John (2011), A Time to Kill, J.
Science. Retrieved from https://www.
sciencemag.org/news/2011/06/time-kill
(Accessed 10 July 2020)
Brazier, Yvette (2018), All about Narcissistic
Personality Disorder, Medical News Today.
Retrieved from https://www.medicalnewsto-
day.com/articles/9741
Brézillon, Patrick (1999), Context in Artificial
Intelligence I: A Survey of the Literature, Com-
puters and Artificial Intelligence, 18(4) 1–28.
Buss, David M. (2005), (Ed.), The Handbook of
Evolutionary Psychology, New Jersey: John
Wiley & Sons, Inc.
Cartwright, Mark (2013), Prometheus, Ancient
History Encyclopedia. Buffalo, NY: Prometheus
Books.
Damasio, Antonio (2003), Looking for Spinoza:
Joy, Sorrow, and the Feeling Brain, New York,
NY: Harcourt, Inc.
Darley, J. M. and C. D. Batson (1973), From
Jerusalem to Jericho: A study of Situational
371
and Dispositional Variables in Helping Behav-
ior, Journal of Personality and Social Psychol-
ogy, 27, 100–108.
Davis, Lawrence (1991), Editor, Handbook of
Genetic Algorithms. Englewood Cliffs, NJ:
Van Nostrand Reinhold.
Durkin, John (1994), Expert Systems: Design
and Development, Prentice-Hall, Inc.
Edell, Aaron (2018), Recognizing context is still
hard in Machine Learning, Towards Data
Science. Retrieved from https://towards
datascience.com/recognizing-context-is-still-
hard-in-machine-learning-heres-how-to-tackle-
it-ed398a725f8c (Accessed 10 July 2020)
Edelman, Gerald M. and Giulio Tononi (2000),
A Universe of Consciousness. New York, NY:
Basic Books.
Evans, Leonard (2008), Death in Traffic: Why
Are the Ethical Issues Ignored? Studies in
Ethics, Law, and Technology, 2(1).
Fisher, Martin Henry (1879–1962), http://liber-
tytree.ca/quotes/Martin.Fischer.Quote.1A58
(Accessed 19 June 2020).
Gazzaniga, Michael S. (2011), Who’s in Charge:
Free Will and the Science of the Brain, New
York, NY: Harper Collins.
Goldberg, David E. (1989), Genetic Algorithms
in Search, Optimization, and Machine
Learning. Reading, MA: Addison-Wesley
Publishing Co., Inc.
Goodfellow, Ian, Yoshua Bengio, and Aaron
Couville (2016), Deep Learning, MIT Press,
Cambridge, MA.
Gould, Stephen Jay (1983), Hen’s Teeth and
Horses Toes: Further Reflections in Natural
History. New York, N: W.W. Norton & Com-
pany, Inc.
Graziano, Michael S. (2013), Consciousness
and the Social Brain. New York, NY: Oxford
University Press.
Greene, Joshua D. (2012), Solving the Trolley
Problem, In Sytsma, Justin and Wesley Buck-
walter (2016), (Eds.), A Companion to Exper-
imental Philosophy, pp 175–178. Hoboken,
NJ: John Wiley & Sons, Ltd.
Hall, Louisa (2017), How We Feel About Robots
That Feel, MIT Technology Review. Retrieved
from https://www.technologyreview.com/
2017/10/24/148259/how-we-feel-about-
robots-that-feel/ (Accessed 10 July 2020)
Harris, Sam (2012), Free Will. New York, NY:
Free Press.
372 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Hayes-Roth, Frederick, Donald A. Waterman,
and Douglas B. Lenat (1983), (Eds.), Building
Expert Systems. Reading, MA: Addison-
Wesley Publishing Co., Inc.
Herrman, Andreas, Walter Brenner, and Rupert
Stadler (2018), Autonomous Driving: How
the Driverless Revolution Will Change the
World. Bingley, UK: Emerald Publishing.
Hull, David L. (2000), Taking Memetics Seri-
ously: Memetics Will Be What We Make It. In
Aunger, Robert, (Ed.), (2000), Darwinizing
Culture: The Status of Memetics as a Sci-
ence, pp. 43–68. New York, NY: Oxford
University Press.
Koch, Christof (2018), What Is Consciousness?
Scientific American, pp. 60–64. June 1, 2018.
Koza, John R. (1992), Genetic Programming,
MIT Press, Cambridge, MA.
Krizhevsky, Alex, Ilya Sutskever, and Geoffrey E.
Hinton (2012), ImageNet Classification with
Deep Convolutional Neural Networks,
Proceedings of the 25th International
Conference on Neural Information Processing
Systems – Volume 1, pp. 1097–1105.
LeDoux, Joseph (1996), The Emotional Brain.
New York, NY: Simon and Schuster.
LeDoux, Joseph E. (2012), Evolution of Human
Emotion: A View through Fear, Progress in
Brain Research, 195, 431–442.
Libet, B. (1993), The Neural Time Factor in
Conscious and Unconscious Events, Ciba
Foundation Symposium, 174, 123–137;
discussion 137–146.
Lin, Patrick, Keith Abney, and George A. Bekey
(2012), Robot Ethics: The Ethical and Social
Implications of Robotics, MIT Press. Cam-
bridge, MA.
Lipson, Hod and Melba Kurman (2016), Driver-
less: Intelligent Cars on the Road Ahead, MIT
Press, Cambridge, MA.
McCulloch, Warren S. and Walter Pits (1943), A
Logical Calculus of the Ideas Immanent in
Nervous Activity. In Anderson, James A. and
Edward Rosenfeld, (Eds.), (1988), Neurocom-
puting: Foundations of Research, pp. 18–31.
MIT Press, Cambridge, MA.
Meystel, Alexander M. and James S. Albus
(2002), Intelligent Systems: Architecture,
Design, and Control. New York, NY: John
Wiley & Sons, Inc.
Moore, James H. (2003), Editor, The Turing
Test: The Elusive Standard of Artificial
Intelligence (Studies in Cognitive Systems).
New York, NY: Kluwer Academic Publishers.
National Safety Council (2019), Road to Zero
Presents Plan to Eliminate Roadway Deaths.
Itasca, IL: Retrieved from https://www.nsc.org/
road-safety/get-involved/road-to-zero.
(Accessed 16 April 2019)
Qian, Xuyu, Hongjun Song, and Guo-li Ming
(2019), Brain Organoids: Advances, Applica-
tions and Challenges, Development. 2019
Apr 16;146(8): dev166074. doi: 10.1242/
dev.166074j.
Rechavi, Oded (2009), Cell contact-dependent
acquisition of cellular and viral non-
autonomously encoded small RNAs, Genes &
Development, 23(16) 1971–9. doi: 10.1101/
gad.1789609.
Schultz, Alan C., John J. Grefenstette, and
Kenneth A. De Jong (1993), Test and
Evaluation by Genetic Algorithms, IEEE
Expert, 8(5) 9–14. October 1993.
Shenhav, Amitai and Joshua D. Greene (2010),
Moral Judgments Recruit Domain-General
Valuation Mechanisms to Integrate Repre-
sentations of Probability and Magnitude,
j. Neuron. 67, 667–677.
Smith, Wesley (2019), Eugenics-Engineered
Babies’ Brains Changed by CRISPR, National
Review, February 21, 2019. Retrieved from
https://www.nationalreview.com/corner/
genetic-engineering-crispr-changed-babies-
brains/#:~:text=The%20brains%20of%20
two%20babies,probably%20impacted%20
by%20the%20procedure (Accessed 10 July
2020)
Spectrum (2019), Brain organoids show realistic
neuronal firing rhythms, Spectrum Research
News. Retrieved from https://www.spectrum-
news.org/news/brain-organoids-show-realistic-
neuronal-firing-rhythms/#:~:text=Brain%20
organoids%20made%20from%20
typical,at%20least%20four%20
months1.&text=A%20few%20features%20
of%20the,preterm%20infants%2C%20
the%20researchers%20say (Accessed 10 July
2020)
Sternberg, Robert (1995), Editor, Wisdom: Its
Nature, Origins, and Developments, New
York, NY: Cambridge University Press.
Teich, Albert H. (2009), Editor, Technology and
the Future, Boston, MA: Wadsworth Cengage
Learning.
 EVOLUTIONARY PSYCHOLOGY AND ROBOTICS
The Economist (2015), If Autonomous Vehicles
Rule the World: From Horseless to Driverless,
July 1, 2015. Retrieved from https://worldif.
economist.com/article/12123/horseless-
driverless (Accessed 10 July 2020)
Wallach, Wendell and Collin Allen (2009), Moral
Machines: Teaching Robots Right from
Wrong. New York, NY: Oxford University Press.
Wikipedia (2019a), Ford Model T. Retrieved
from https://en.wikipedia.org/wiki/Ford_
Model_T (Accessed 10 July 2020)
373
Wikipedia (2019b), Timeline of Motor Vehicle
Brands. Retrieved from https://en.wikipedia.
org/wiki/Timeline_of_motor_vehicle_brands
(Accessed 10 July 2020)
Zaki, Jamil (2013), Using Empathy to Use People:
Emotional Intelligence and Manipulation,
Scientific American. Retrieved from https://
blogs.scientificamerican.com/moral-universe/
using-empathy-to-use-people-emotional-
intelligence-and-manipulation/ (Accessed
10 July 2020)

Evolutionary Psychology and
Dangerous Driving Behaviour
Deanna Singhal and David Wiesenthal
INTRODUCTION
This chapter discusses the contributions of
evolutionary psychological and social learn-
ing theories to explain aggressive and risky
driving. Dangerous driving behaviour may
be influenced by a number of factors: ‘the
young male syndrome’ and observational
learning may contribute to the choice to drive
or respond aggressively on the roads.
Exposure to media, such as motion pictures
and racing video games, can promote the
modelling of dangerous driving behaviours.
How situational factors interact with per-
sonal and internal factors to promote risky
driving is discussed within the context of the
General Aggression Model (Anderson and
Bushman, 2002). A summary of what is
known about this topic is provided, including
a review of research studies using different
methodologies (e.g., laboratory, archival),
and considerations for public policy are
discussed.
19
AGGRESSIVE AND RISKY DRIVING
Each driver at some point in their driving
experience has likely witnessed, or even
engaged in, an act of aggressive or risky driv-
ing on the road, such as tailgating, weaving
in and out of traffic, running red lights, and
driving at excessive speeds. Driving behav-
iour is considered aggressive if ‘it is deliber-
ate, likely to increase the risk of collision,
and is motivated by impatience, annoyance,
hostility and/or an attempt to save time’
(Tasca, 2000: 2). It has been suggested that
such behaviours can be frustration-driven
and are more likely in an enabling environ-
ment (Shinar, 1998). Some aggressive driv-
ing involves hostile aggression, where the
behaviour is directed at an object of frustra-
tion (e.g., cursing at another driver), whereas
other acts involve instrumental aggression,
where the frustrated driver attempts to move
ahead at the expense of other drivers’ rights
(e.g., weaving in and out of traffic)
 EVOLUTIONARY PSYCHOLOGY AND DANGEROUS DRIVING BEHAVIOUR
(Roseborough, 2014; Shinar, 1998). In the
context of evolutionary psychological theory,
frustration may not necessarily be the only
reason why these driving behaviours occur,
as they may involve voluntarily initiated dis-
plays of dominance. Males are at a greater
risk of motor vehicle collisions than females
because they are more likely to be involved
in speeding, tailgating, and responding
aggressively to perceived misdeeds of other
drivers (Wilson and Daly, 1985).
The ‘young male syndrome’, coined by
Wilson and Daly (1985), refers to the pat-
tern of disproportionate involvement of males
in risky or aggressive behaviours, such as
gambling, homicide, and motor vehicle col-
lisions. In the context of evolution, aggres-
sion can have an adaptive value and, in
certain situations, males can perceive a ben-
efit. Competitive risk-taking among males is
believed to have evolved as a result of repro-
ductive competition, where males compete
with other males for the valued resource of
females (Wilson and Daly, 1985). When envi-
ronmental resources are scarce, or unequally
distributed, and when the proportion of young
males in the population is large, both risk-­
taking and aggression increase (Mesquida and
Wiener, 1996; Wiesenthal and Singhal, 2012).
Females may perceive these behaviours as
indicators of a mate who will protect and
provide for her and any offspring. These are
desired characteristics associated with paren-
tal investment (Buss, 1988). Males engage in
social displays of risky behaviour in an effort
to secure the resources necessary to attract
mates. Intra- and intergroup competition,
which involves prestige, rank, and reputation,
may enhance the likelihood of attracting pro-
spective mates by elevating the male’s posi-
tion on a dominance hierarchy (Wilson and
Daly, 1985). The ability to provide social and
psychological resources, in addition to mate-
rial resources, is considered important for
mate selection in females (Buss, 1988).
Competition in young males is not
restricted to driving scenarios. If society
375
bombarded young males with video games
and motion pictures featuring chess-playing
scenarios that resulted in rewarding experi-
ences as a result of competitiveness, it seems
reasonable to conclude that competition
between young males would take place on
chess boards rather than streets and high-
ways. Chess is a symbolic or cognitive exam-
ple of warfare, with males comprising the vast
majority of Grandmasters, a title awarded to
world-class chess players by the International
Chess Federation (FIDE). The most recent
FIDE list of Grandmasters includes 1,680
individuals from a variety of countries, 1,643
of which are male (98%) (‘List of chess
grandmasters’, n.d.). Evolutionary tenden-
cies are not the only influences on behaviour
and the environmental or social exposure to
depictions of competition, in various media
forms, can serve as representative examples
of which behaviours males should engage in
and how. The presentation that will follow
later in this chapter will review the effects of
cinematic portrayals of risky driving.
Roadways are a social environment that
affords displays of competitive behaviour.
When driving, individuals can engage in
risk-taking competitiveness (Wiesenthal
and Singhal, 2012), particularly among
young males, which can contribute to risky
or aggressive driving behaviours (Vingilis
et al., 2013). An overrepresentation of young
males in annual motor vehicle collisions was
reported in British Columbia in 2018 (Peng,
2018). In particular, there was a doubling of
deaths involving motorcycles, where 27 of the
30 victims were male and a third of all victims
were between the ages of 19 and 29 years.
Motorcycles may represent a riskier vehicle
to drive and, by design, promote risky-driving
behaviours, such as speeding. One British
Columbian police agency reported a ‘major
increase’ in excessive speeding province-wide
that was not specific to motorcycles, with
25% more drivers being stopped for this risky
behaviour (Peng, 2018). Though no known
reasons for these increases were reported,
376 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
previous research has demonstrated that
increases in male driving-related fatalities
even occurred during a decrease in the male
population (Wiesenthal and Singhal, 2012).
Though evolutionary psychological theory
can make an important contribution to the
understanding of risky driving behaviour,
societal influences, such as the presence of
others and individual difference variables
(e.g., personality), interact with male com-
petitiveness and can influence the decision to
engage in risk-taking behaviours (Wiesenthal
and Singhal, 2012; Wilson and Daly, 1985).
An important framework to consider in this
context is social learning theory, which sug-
gests that transmission of behaviour can
occur through observational learning, where
an individual acquires new patterns of behav-
iour by observation (Bandura, 1971). Media
influences on the behaviour of children and
adults have long been a research concern in
developmental, social, and clinical psychol-
ogy. The advertising industry is based on
the fact that consumers can be influenced
by media messages. While most advertising
is concerned with altering or influencing
relatively trivial consumer behaviours, the
material presented in this chapter indi-
cates that media can trigger dangerous
and life-threatening actions in predisposed
individuals.
The novelty, relevance, and consequences
of the behaviour play a role in the imitative
effects of observation (Bandura, 1971). The
term modelling encompasses the broader
psychological effects that can occur with
observational learning, beyond mere imi-
tation or mimicry (Bandura, 1971). These
could include cognitive changes (e.g., in
attitudes) that accompany the learning of
a behaviour, even if the behaviour was not
performed following observation (Modeling,
n.d.). Marketing relies upon individuals mim-
icking the use of products shown in adver-
tising, and uses media to communicate this
information. How much alcohol individuals
choose to drink, or what they perceive as an
ideal body image, is influenced by what they
see and hear on television, in the movies, on
the Internet, or in newspapers and magazines
(Koordeman et  al., 2011; López-Guimerà
et al., 2010). Media recognizes this influence
and, in some cases, limits what viewers can
see, in an attempt to prevent imitation of a
behaviour. For example, the Toronto Transit
Commission (TTC) has an agreement with
local media to not treat subway suicides as
newsworthy, in an attempt to prevent imita-
tion. Soon after the adoption of this standard
in 1971, the TTC reported a marked decrease
in the number of suicides (Toronto Transit
Commission, 2010).
With respect to road safety, modelling of
aggressive or risky driving can occur as a
result of observational learning, with media
being an influential source. Media depictions
of dangerous or aggressive driving and glo-
rifications of such risk-taking have become
increasingly popular (Fischer et al., 2012). A
pivotal influence was the 1968 release of the
motion picture Bullitt. This film featured an
exciting car chase, starring Steve McQueen,
which involved risky and aggressive driv-
ing through the streets of San Francisco
(Wiesenthal et  al., 2016). A more current
driving movie franchise is Fast and Furious,
in which entire movies depict acts of ille-
gal street racing and heists. The franchise
has grown in popularity since its first movie
release, The Fast and the Furious, in June of
2001. This movie grossed approximately two
million dollars at the worldwide box office,
compared to a more recent franchise instal-
ment, Furious 7, which grossed over one
and a half billion dollars worldwide, with an
additional 70 million dollars in DVD sales
(Nash Information Services, n.d.-c).
Depictions of speeding, racing, dan-
gerous passing, and aggressive behaviour
towards other drivers are often shown in the
absence of negative consequences (Beullens
et  al., 2011b; Greenberg and Atkin, 1983).
According to media effects theory, view-
ers’ perceptions of what is normal can be
influenced by the more frequent portrayal of
certain behaviours (Beullens et  al., 2011b).
 EVOLUTIONARY PSYCHOLOGY AND DANGEROUS DRIVING BEHAVIOUR
The risky driving behaviours often depicted
in media are novel, such that viewers do not
commonly see them on the roads (Atkin,
1989). The more normative the events appear,
the greater the possibility of viewer disin-
hibition (Atkin, 1989). Combined with the
lack of depiction of negative consequences,
learning of aggressive or risky driving may
be enhanced (Beullens et al., 2011b). Adding
to the possibility of imitation is the portrayal
of a hero as the risky driver, which can be
attractive to viewers (Beullens et al., 2011b)
and creates a scenario in which the behaviour
seems justified, further fostering disinhibi-
tion (Atkin, 1989). Media which is ‘arousing,
nonfictional, justified, positively reinforced,
and performed by individuals with whom
viewers identify and feel similar’ is the most
likely to influence behaviour (Vitaglione,
2012: 489).
There is evidence that motion-picture
production companies are aware of the pos-
sibility of modelling risky and unsafe driv-
ing. Prior to the release of The Fast and the
Furious in 2001, Universal Studios posted a
disclaimer on the film’s promotional website
stating, ‘All of the racing stunts in “The Fast
and the Furious” were performed in a staged
environment by professionals with years of
training and experience. Please do not try
any of these yourself. Be smart. Drive safe.
Stay legal’ (Goldberg, 2001; Orwall, 2001).
In the days leading to the film’s release, they
also ran two public-service announcements,
recorded by the two lead actors in the film,
Vin Diesel and Paul Walker, emphasizing
safe driving. Given the awareness of pos-
sible imitation, it has been said that media
is a major part of the problem when imita-
tive behaviour is undesirable or antisocial
(Coleman, 2004). Though observational
learning can be useful and efficient (i.e.,
when errors in learning are costly or dan-
gerous or when teaching structured rules
and skills is time-consuming and difficult)
(Bandura, 1971), modelling behaviour that is
risky or aggressive can have serious negative
consequences.
377
IMITATION AND MODELLING
Social Learning Theory
Social learning theory explains the transmis-
sion of behaviours. Aggression, like other
complex social behaviours, may be imitated
as a result of observational learning or direct
exposure (Bandura, 1971). Bandura et  al.’s
(1963a) classic experiment investigated the
imitation of aggression in children following
the viewing of an adult model interacting with
a three-foot Bobo doll. Young children who
viewed a video of the model being aggressive
towards the doll (e.g., hitting the doll while
sitting on it, kicking and throwing it, pummel-
ling it with a mallet), displayed more aggres-
sive acts when allowed to interact with the
Bobo doll in a situation of frustration, com-
pared to the control group, who did not view
the aggressive model (Bandura et al., 1963a).
Though this study has been criticized for con-
sidering the acts displayed by the children to
be aggressive, given that a Bobo doll is essen-
tially designed to be pummelled (Freedman,
2007; Milgram and Shotland, 1973), it did
demonstrate that children imitated specific
observed behaviours, both physical (e.g., sit-
ting on Bobo and hitting him with a mallet)
and verbal (e.g., shouting ‘Sock him in the
nose’) (Bandura et al., 1963a). Other research
with children has suggested that early child-
hood exposure to aggressive or harsh punish-
ment from parents can provide a situation of
observational learning and result in aggressive
behaviour from children towards others (Weiss
et al., 1992).
Though the social learning theory of
aggression stems from research with chil-
dren, it has implications for adults. Early
exposure to aggression and violence may
contribute to children developing different
cognitions associated with aggression. For
example, they may be more likely to attrib-
ute hostile intentions to others’ ambiguous
behaviours (i.e., hostile attributional bias;
Steinberg and Dodge, 1983). Therefore, they
may become more likely to select aggressive
378 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
solutions to situations in the future (Steinberg
and Dodge, 1983). In addition, the factors
proposed to influence imitation of aggression
in a situation of observational learning, spe-
cifically positive reinforcement, novelty, and
relevance to a given social situation, are not
specific to children (Atkin, 1989; Beullens
et al., 2011b; Vitaglione, 2012). For example,
the internal thrill or the accolades of friends
associated with trying an aggressive or risky
driving behaviour seen in a movie are exam-
ples of positive reinforcement, which can
influence this adult behaviour.
Social Interaction Theory
According to social interaction theory,
aggressive behaviour is social influence
behaviour, whereby coercive acts are used to
obtain things of value, bring about retributive
justice, or promote social and self-identities
(Tedeschi and Felson, 1994). Two social
influences proposed to mediate aggression
are socialization practices within the family
and peer influences (Wiesenthal and Singhal,
2012). For example, negative parenting,
including poor family organization and func-
tioning, as well as marital conflict, contrib-
utes to aggression among adolescents (Talwar,
1998). Children may adopt peer values and
beliefs as substitutes for parental values and
beliefs. When the peer influence is deviant
and the individual is susceptible because of
negative family influences, aggression
increases (Talwar, 1998).
With respect to driving, the presence of
others increases aggressive and risky behav-
iours. Teenage drivers engage in more risky
driving (i.e., speeding and reduced headway)
with male passengers, compared to no pas-
sengers or female passengers, with the male
driver/male passenger combination produc-
ing approximately double the rate of risky
or aggressive driving (Simons-Morton et al.,
2005). According to evolutionary psycholog-
ical theory, intrasexual competition can result
in a greater display of desired resources
believed to be important for mate selection
(Buss, 1988). The ownership of a fast car and
the demonstration of a daring male driver
may be perceived as more attractive and
dominant than his passenger counterparts.
The driver may also be viewed as possess-
ing similar characteristics to those of famous
actors portrayed as heroes in risky driving
movies, such as wealth and prestige.
In addition to the influences of intrasexual
competition, peer pressure can influence
behaviour, and the susceptibility to this social
influence is believed to be greatest during
early adolescence (Wilson and Daly, 1985).
Full development of resistance to peer pres-
sure does not necessarily occur between late
adolescence and early adulthood. During this
period of time, young individuals may face
other risky behaviour choices, such as drug
use and teenage sexual encounters. If young
males are already more likely to engage in
risky behaviours (Wilson and Daly, 1985),
they may be more susceptible to the influ-
ence of their peers who encourage participa-
tion (Steinberg and Monahan, 2007).
Influence of Behavioural
Consequences
Research on operant conditioning was rooted
in animal behaviour (Skinner, 1938), but
consequences associated with an action were
proposed to not only predict but also control
an individual’s behaviour (Skinner, 1953). In
the case of childrearing, for example, rein-
forcement and punishment are used to teach
a child appropriate behaviour. An unwanted
behaviour results in a negative consequence,
such as a ‘time-out’, while a desired response
results in a positive consequence or reward,
such as a treat. Research on vicarious learn-
ing suggests that observing another individ-
ual experience consequences for a behaviour
can also influence one’s own behaviour. The
influence of observing punishment or reward
on the imitation of aggression in children
was demonstrated in a follow-up study by
 EVOLUTIONARY PSYCHOLOGY AND DANGEROUS DRIVING BEHAVIOUR
Bandura et al. (1963b). Children who viewed
the adult model being punished for aggres-
sive behaviour towards the Bobo doll were
less likely to imitate the behaviour than chil-
dren who viewed the aggressive model being
rewarded. In adults, vicarious learning can
result in attitude change, as demonstrated in
one study where male university students
reported being more accepting of interper-
sonal violence against women following the
viewing of movies depicting positive conse-
quences associated with sexual violence
(Malamuth and Check, 1981).
Though evolutionary psychological theory
suggests that males, more than females, have
a tendency to engage in aggressive behaviour
in order to acquire desired resources (Wilson
and Daly, 1985), cognitive behaviourism
suggests that individuals have free will to
contemplate and incorporate environmen-
tal information, such as behavioural con-
sequences of punishment and reward, into
decision-making processes related to behav-
iour choice (Bandura, 2001b). The absence of
negative consequences, as observed in many
movies depicting risky and aggressive driv-
ing, may act as negative reinforcement. When
the undesirable consequences of risky driv-
ing, such as fines, collision, injury or death,
are removed, the driving behaviour will be
reinforced and more likely to occur in the
future. When such outcomes are commonly
depicted in the movies, vicarious learning
also contributes to the probability of young
males engaging in this driving behaviour in
the real world.
The Copycat Effect
The copycat effect refers to imitation or
adoption of behaviours or practices of
another, and its effects, usually negative and
unfavourable, are believed to be triggered by
media (Coleman, 2004). Phillips (1974)
coined the term Werther effect, describing the
copycat effect associated with suicide. In the
1774 novel The Sorrows of Young Werther,
379
by Johann Wolfgang von Goethe, the young
Werther shoots himself after realizing that he
could never be with the woman he loved. In
following years, many young men killed
themselves in the same manner, sitting at a
desk, dressed like Werther, with Goethe’s
novel in front of them (Coleman, 2004). To
investigate this effect, Phillips recorded the
number of monthly suicides in the United
States, following the publication of front-
page suicide stories in the New York Times,
during the period 1947–1967. Using the pre-
vious and subsequent year as a comparison,
he found a significant tendency for suicides
to increase, and a dose-response relationship
between the number of suicides and the
amount of front-page story coverage
(Phillips, 1974). The influence of the news-
paper coverage was location-specific, such
that stories published in the New York Times,
but not in Great Britain’s most popular news-
paper, The London Daily Mirror, resulted in
a greater increase in suicides in the United
States, compared to Great Britain. However,
when the suicide story was covered in both
newspapers, British suicides increased sig-
nificantly (Phillips, 1974).
The Werther effect has also been demon-
strated for suicide stories covered on network
television news programs. For example,
in California, between 1968 and 1985, the
number of suicides significantly increased
0–7 days after a publicized television story
(Phillips and Carstensen, 1988). The effect of
this coverage can also be influenced by the
involvement of a celebrity. In Korea, between
2005 and 2008, television news coverage
of a celebrity suicide increased the number
of emergency department visits associated
with suicide attempts or self-injury in the
three weeks following the reported suicide
(Jeong et  al., 2012). The method of suicide
used by celebrities can also be imitated, as
was demonstrated following the 2009 rail-
way suicide of Robert Enke, a celebrated
German football goalkeeper. German rail-
way suicides increased both in the short term
(i.e., two weeks following the event) and in
380 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
the long term (i.e., two years after the event)
(Hegerl et  al., 2013). A significant associa-
tion between frequency of Google searches
for ‘Enke’ and frequency of railway suicides
in Germany was documented (Koburger
et  al., 2015), suggesting a media influence
on this copycat behaviour. However, it was
not possible to determine if those who com-
mitted suicide had experienced the media
exposure.
The copycat effect can also involve
criminal behaviour. Copycat crime refers to
imitative behaviour where an offence, origi-
nally portrayed in the media, is subsequently
performed in reality. It can be motivated by
real or fictional media depictions, and the
offender incorporates aspects of the origi-
nal crime (Helfgott, 2015). The 1994 film
Natural Born Killers has been linked to over
a dozen copycat crimes (Helfgott, 2015). The
film depicts a young, attractive couple on a
roadtrip/serial-mass-murder spree across the
Southwestern United States, during which
they kill over 50 people. Some couples, who
committed similar murders, reported being
obsessed with the movie and copied behav-
iours depicted in the film during their crimes.
In one copycat case, the young woman lured
her victim to his death by promising sex, rep-
licating a scene in the movie. Many examples
of copycat crime have been described, each
with its own unique media source, such as
the news coverage of the Columbine school
shooting. It has been suggested that the link
between viewing media violence and violent
crime is difficult to establish because of the
other factors that contribute to the behaviour
(e.g., individual, environmental, and situ-
ational elements; Helfgott, 2015). Though
it may be difficult to establish cause-and-
effect links, later material in this chapter
summarizes research that establishes rela-
tionships between viewing aggressive and
risky driving media and engagement in these
behaviours.
With respect to movie content, there is
anecdotal evidence supporting the copycat
effect. In 1993, Disney made a controversial
decision to remove a scene from their movie
The Program (Pristin and Fox, 1993). The
scene depicted an inebriated college football
quarterback lying in the middle of the high-
way, with cars passing by, barely ­missing him.
As the scene continued, other football play-
ers joined him and no one was hurt. Disney’s
decision to remove the scene came after two
reported attempts of imitation in New Jersey
and Pennsylvania (Kotzen, 2013). One teen-
ager was killed and two others were seriously
injured (Pristin and Fox, 1993).
Television and social media are sources
that can also contribute to the modelling of
driving behaviour. Bird Box, a Netflix movie
released in 2018 (IMDb, 2018), contains
scenes of individuals manoeuvring through
their environment blindfolded to avoid see-
ing a presence that urges them to commit sui-
cide when observed. This includes scenes of
driving in a vehicle with all windows covered
in tape. The ‘Bird Box Challenge’ became a
social media craze that encouraged people to
try everyday tasks blindfolded (Deb, 2019).
Police in Utah had to issue a warning to its
citizens not to drive blindfolded. Though this
would seem obvious to most, the warning
was issued after a female teenage driver, with
a hat pulled over her eyes, drove into oncom-
ing traffic and hit another car (‘US driver’,
2019).
In California, following the release of
The Fast and the Furious, the Los Angeles
Police Department increased patrols to stop
street racing (Goldberg, 2001). Supporting
their action was the knowledge that copycat
behaviour had occurred in their area follow-
ing the release of Gone in 60 Seconds, which
glamorized auto theft. On the day of this
film’s release, and the day after, the num-
ber of stolen vehicles in the area more than
doubled, compared to the previous two years
(Goldberg, 2001). Increased enforcement
was also reported for the release of the fourth
instalment of the Fast and Furious franchise
in 2006, The Fast and The Furious: Tokyo
Drift (Rowland, 2006). California Highway
Patrol reported being extra vigilant during
 EVOLUTIONARY PSYCHOLOGY AND DANGEROUS DRIVING BEHAVIOUR
this time in stopping any unsafe or illegal
driving, such as street racing or speed con-
tests (Rowland, 2006). Increased enforce-
ment was also reported in Toronto for the
release of this film, with cruisers positioned
outside theatres near highways (Grewal and
Brennan, 2006). The Ontario Provincial
Police reported witnessing copycat behaviour
and resulting crashes following the release of
the previous Fast and Furious film, Turbo-
Charged Prelude. They expressed the opin-
ion that young people get caught up in the
excitement and adrenaline of these films and
attempt to imitate their heroes (Grewal and
Brennan, 2006).
Mass media is a source of social or situ-
ational influence on aggressive or risk-taking
behaviour (Bandura et al., 1963a; Bushman,
1995; Eron et al., 1972; Paik and Comstock,
1994; Vitaglione, 2012). In the context of
driving, when a driver speeds and weaves in
and out of traffic, following the viewing of
a movie that depicted such behaviour, they
are not necessarily being violent or want-
ing to harm other drivers, but their risky and
aggressive behaviour increases the likeli-
hood of collision (Transport Canada, 2011),
putting others at risk. Depictions of patterns
of aggressive behaviours can teach behav-
iour styles, alter restraints on the behaviour,
desensitize an individual to aggression, and
shape the viewer’s perceptions, which are
important bases for individual behaviour
(Grey et al., 1989).
Clearly, there are multiple factors that can
contribute to driving aggressively or engag-
ing in risky driving behaviours. Evolutionary
psychological theory suggests the importance
of personal factors, such as age and sex dif-
ferences, and social learning theory empha-
sizes the need to consider situational factors
in the environment. In the context of aggres-
sion, Anderson and Bushman (2002) devel-
oped the General Aggression Model (GAM),
which addresses how these, and other factors,
can interact to produce aggressive behaviour.
This model can serve as a basis for better
understanding aggressive or risky driving.
381
GENERAL AGGRESSION MODEL
The GAM (Anderson and Bushman, 2002) is
a multifaceted approach to explaining aggres-
sive behaviour, incorporating aspects from
social and evolutionary theories, as well as
components related to personality, cognition,
and arousal. It is an episodic model that
focuses on the individual in a particular situ-
ation and consists of three overall compo-
nents: inputs involving person and situation
factors, routes through which arousal, affect,
and cognition play a role, and outcomes
resulting from appraisal and decision pro-
cesses (Anderson and Bushman, 2002).
Person factors include things such as age,
sex differences in aggressive tendencies,
and personality traits. Physical and verbal
aggression, hostility, and anger can be inter-
correlated and suggest overall trait aggres-
siveness, indicating that people who are angry
are more likely to aggress (Buss and Perry,
1992). Sex differences have been reported
using the Aggression Questionnaire, such that
males have a higher total score, are much more
physically and verbally aggressive, and are
somewhat more hostile than females (Buss
and Perry, 1992).
Sensation seeking refers to the need and
willingness to engage in novel events that are
highly arousing, even if they involve physi-
cal and social risk (Zuckerman, 1979). It is
related to risky behaviours, such as danger-
ous driving (e.g., drunk driving and speeding)
(Wiesenthal et al., 2016), drug use, and minor
criminality (Arnett, 1994). It has been sug-
gested that sensation seekers, having greater
acceptance or reduced perception of risk, are
more likely to engage in behaviours that are
both risky and aggressive (Wiesenthal et al.,
2016). Sensation seeking can serve as a bio-
logical predisposition that interacts with the
conditions of the social environment, such
that less restrictive environments allow for its
greater expression (Arnett, 1994).
In the context of evolutionary psychologi-
cal theory, both trait aggression and sensation
382 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
seeking may benefit males in the context of
intrasexual competition for mates. Displays
of aggression and willingness to take risks
may present as qualities of an individual who
will protect and provide for a female and
their offspring. Mate-selection criteria for
females include characteristics that demon-
strate potential for parental investment, such
as protection, social status, and the ability to
provide food and shelter (Buss, 1988). Males
who display the ability to provide social, psy-
chological, and material resources may be
considered more desirable by females and
will compete strongly to demonstrate these
characteristics. Displays of these character-
istics do not require direct combat nor con-
tact with competitors, but males may engage
in representative behaviours at the expense
of others who are competing for females
(Buss, 1988).
Situational factors, such as aggressive cues
(e.g., weapons), provocation, frustration,
and social opportunity for aggression (e.g.,
church versus nightclub), interact with the
individual’s internal state. These can influ-
ence affect, arousal, and cognition, which
are interconnected, and can influence each
other in a reciprocal fashion (Anderson and
Bushman, 2002). The existence of memory
scripts can increase the likelihood of model-
ling aggressive behaviour. A script is a type
of schema that is learned through experience
and exposure, and comes to define a situa-
tion. Highly associated concepts are linked
together in memory and these associations
often involve causal links, goals, and action
plans (Abelson, 1981). For example, the use
of a gun can be associated with concepts such
as anger, pain, retaliation, and the action of
shooting (Anderson and Bushman, 2002).
When components are strongly linked, they
become a unitary concept in semantic mem-
ory and provide a guide for future behaviour.
The greater the exposure to the behaviour, the
greater the priming of these scripts, increas-
ing the likelihood of their use.
The use of memories in the decision to
engage in aggressive behaviour is incorporated
in the broader cognitive neoassociation
theory of aggression. Berkowitz (2012) states
that the decision to act aggressively can
begin with an aversive event, such as frus-
tration or provocation. This leads to nega-
tive affect, which automatically stimulates
both fight and flight physiological pathways.
A dominant fight reaction leads to irritation
or anger, whereas a dominant flight reaction
leads to fear. In addition to the activation of
the physiological components of these path-
ways, associated memories, thoughts, and
motor responses will also be activated. Once
the initial, more involuntary response tenden-
cies arise, higher-order cognition can become
involved in the processing of appraisals and
attributions, as well as the consideration of
rules and consequences associated with cer-
tain behaviour. These can modify or even
extinguish the initial reaction, serving as self-
regulation or control processes. However, it
is possible that an individual can become
angry enough to respond aggressively, with-
out the intervention of higher-order cognition
(Berkowitz, 2012).
The physiological excitation produced
from a given arousal-producing exposure, as
mentioned in the cognitive neoassociation
theory, does not necessarily end abruptly with
the termination of the exposure (Zillmann,
1971). The arousal may linger for some time
and be carried over to a subsequent event or
experience, even one unrelated to the prior
exposure. This residual arousal can influence
the cognitive appraisal of the subsequent
emotional state, with greater residual arousal
creating a more intense subsequent emotion.
This suggests that a potentially stronger nega-
tive affect could be elicited by a provoking
event, producing an ‘over-intense’ response
to stimuli in a future scenario. The residual
arousal could ‘energize’ or facilitate associ-
ated aggressive behaviour (Zillmann, 1971).
The idea of interaction and reciproc-
ity between person, situation, and internal-
state factors was also suggested by Bandura
(2001b) in his Triadic Reciprocal Causation
model of psychosocial functioning. This
 EVOLUTIONARY PSYCHOLOGY AND DANGEROUS DRIVING BEHAVIOUR
model includes personal, behavioural, and
environmental determinants, where personal
factors involve cognitive, affective, and bio-
logical events. These determinants can influ-
ence each other in a bidirectional fashion. For
example, an event in the environment, such
as viewing an aggressive driving movie, can
influence behavioural patterns of the viewer,
with past behavioural patterns and person
factors, such as mood and personality, medi-
ating the choice of response.
The output level of the GAM involves
appraisal and decision processes, which can
lead to either impulsive or thoughtful action
(Anderson and Bushman, 2002). For the
response to be a more thoughtful action, one
requires sufficient time and cognitive capac-
ity to reappraise the situation. Alternative
responses are considered until one is chosen,
but this does not imply that aggression has
been averted. In fact, it is suggested that the
reappraisal process could activate memories
leading to an increase in anger, which could
provide justification for retaliation and inter-
fere with other higher-order cognitive pro-
cessing. Additionally, the individual could
become more aware of potential damage to
their social image and the need to act aggres-
sively in order to maintain it (Anderson and
Bushman, 2002).
Related to the output level of the GAM,
Bandura (2001b) proposed that ‘human
agency’ (i.e., using action to intentionally
make things happen) involves both self-
reactiveness and self-reflectiveness. The first
involves individuals monitoring their own
behaviour and being aware of the conditions
under which it is produced (i.e., cognitive and
environmental). The second is an individual’s
higher metacognitive ability to reflect on and
evaluate their actions, considering the effects,
not only for themselves, but also for others.
In a social context, when one encounters a
situation that is provoking, exciting, aggres-
sive, or risky, the GAM suggests there are a
number of factors that will influence an indi-
vidual’s choice to respond with an aggressive
or risky behaviour. The recent viewing of
383
media depicting this behaviour is a situational
factor that can influence the internal state. It
can increase arousal and influence affect
and cognition, partly through the priming
of existing memory scripts associated with
aggressive or risky behaviour or the forma-
tion of new scripts through observational
learning. In this context, media can play an
influential role in one’s choice to engage in
aggression.
Figure 19.1 shows how the GAM factors
can interact and influence the modelling of
aggressive or risky driving behaviour.
Elements of the ‘young male syndrome’
contribute to person factors. Drivers who are
young, male, and have higher levels of sen-
sation seeking and trait aggression may be
more susceptible to the situational factors of
media exposure to aggressive or risky driv-
ing content and driving scenarios that involve
frustration, provocation, and competition.
Sensation seeking has been shown to be a
significant predictor of aggressive and risky
driving, including the physical and verbal
expression of driver anger and using the vehi-
cle to express anger. It was also predictive of
lapses in concentration and minor losses of
vehicular control (Dahlen et  al., 2005). In
the context of intrasexual competition, males
may engage in displays of risky and aggres-
sive driving in an attempt to display charac-
teristics that females desire in a mate, such
as social status and the ability to protect and
provide for a family. Males may emulate the
revered actors portraying the onscreen heroes
in an attempt to model such adaptive traits.
Arousal produced from viewing a film
involving aggressive or risky driving may cre-
ate stronger aggressive emotional responses
when the driver is later confronted with an
anger-provoking situation on the road. This
is particularly true if memory scripts for this
behaviour already exist. The more one views
media depicting acts of aggressive or risky
driving, the more likely memory scripts will
form, associating negative emotion and aggres-
sive driving responses with provoking or frus-
trating driving situations. This contributes
384 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Figure 19.1  GAM factors included in the investigation of modelling of aggressive or risky driving
to the formation of driver anger and driver
vengeance, where drivers more readily choose
aggressive responses in certain driving situa-
tions (e.g., being cut off by another driver).
TYPES OF MEDIA DEPICTING
AGGRESSIVE OR RISKY DRIVING
Various forms of aggressive or risky driving
media can influence behaviour, including
television, video games, movies, and the
Internet. Research has specifically assessed
the magnitude of this content in the media.
During the years 1975 to 1980, it was esti-
mated that within 784 driving scenes ana-
lysed from primetime broadcasts of popular
television programmes, there were 1,301 acts
of irregular driving, with the most common
acts being speeding, quick braking, squealing
brakes, tires screeching, and quick acceleration
(Greenberg and Atkin, 1983). These acts
were estimated to be shown seven times per
hour, and drivers were predominantly young
males, rarely wearing seat belts, who engaged
in irregular and dangerous driving acts that
rarely resulted in negative consequences
(e.g., death or injury, physical damage, or
legal penalties; Greenberg and Atkin, 1983).
A form of television programming specific
to driving is automobile commercials. Between
the years of 1998 and 2002, it was estimated
that 45% of North American automobile and
truck commercials (n = 349) displayed unsafe
driving (Shin et  al., 2005). Aggressive driv-
ing behaviour accounted for more than 80%
of that content, with high speeds constitut-
ing the majority (Shin et al., 2005). Between
the years of 2006 and 2007, it was found that
approximately 20% of 200 Canadian automo-
bile advertisements (i.e., television, magazine,
 EVOLUTIONARY PSYCHOLOGY AND DANGEROUS DRIVING BEHAVIOUR
and newspaper) included some form of unsafe
driving, such as speeding or hard stops (Watson
et al., 2010). Considering television commer-
cials only, because of their ability to display
motion, it was estimated that 20% contained
depictions of speed. In each television adver-
tisement in which speeding was displayed, a
disclaimer was also presented, demonstrat-
ing the vehicle manufacturer’s awareness of
the unsafe or questionable driving displayed
(Watson et al., 2010).
One type of televised event suggested
to clearly depict aggressive or risky driv-
ing is competitive automobile racing within
NASCAR (National Association for Stock Car
Racing), which has an estimated 75 million
fans viewing these events each year, and tens
of thousands attending in person (Vitaglione,
2012). Dangerous and risky driving is broad-
cast live and is encouraged and reinforced
by both cheering fans and large monetary
rewards. For example, the total purse for the
2015 Daytona 500 (the last year purse data
was publicly available) was approximately
$18 million. The winner received $1,581,453
and the last-place driver (i.e., the loser) still
took home $262,390 (Mensching, 2015). The
celebratory nature surrounding awarding the
winner with large sums of money creates an
air of acceptance of the behaviour and a ‘hero’
for the fans. This combination has the capac-
ity to increase the possibility of modelling the
aggressive or risky driving behaviour (Atkin,
1989; Beullens et al., 2011b).
Popular action movies can contain scenes of
aggressive or risky driving. Of 26 action mov-
ies shown in Belgium between 2005 and 2006
(e.g., War of the Worlds, Casino Royale, and
The Fast and the Furious: Tokyo Drift), there
were 287 driving scenes, with 129 depict-
ing risky driving (Beullens et  al., 2011b).
Frequently occurring risky driving behaviours
were speeding, tires screeching, brakes squeal-
ing, and quick braking or sudden decreases
in speed. Risks associated with such driving
behaviours were rarely shown. Approximately
one-third were followed by a crash and showed
the endangering vehicle, or its surroundings,
385
being damaged. In none of the scenes were
legal penalties shown. Risky drivers tended
to be young males and lead characters, who
were the heroes in the movie (Beullens et al.,
2011b). These individuals can be perceived
by young male viewers as successful, strong,
and dominant characters who serve as mod-
els, guiding them in how they should look and
behave in order to achieve similar status.
The popularity of social media, specifically
YouTube, in the context of media depictions of
aggressive or risky driving has been addressed
(Vingilis et  al., 2017). On this website, indi-
viduals can create a free account, post videos,
and view others’ videos. Using the search
term ‘street racing’, over 33 million videos
were found (as of April 30, 2015) and, of
the 10 most popular (i.e., averaging approxi-
mately 787,677 views), nine were on public
roads, not racetracks. Searching terms of other
risky driving behaviours, such as ‘burnouts’
or ‘drifting’, found similar results. The avail-
ability of this information on the Internet is
evident, but who is watching these videos, and
how they influence driving behaviour, is not
known, given the lack of research on this spe-
cific type of media (Vingilis et al., 2017).
A body of research, using various method-
ologies, has demonstrated that risk-glorifying
media, some including content depicting acts
of aggressive and risky driving, has the capacity
to increase risk-promoting cognitions, arousal,
affect, willingness to take risks in observed
driving scenarios, and risky driving behaviour.
These effects have been shown to last 24 hours,
and even as long as several days, with some of
the research demonstrating intervening vari-
ables of sex differences and personality traits
of sensation seeking and aggressiveness.
RESEARCH ON THE INFLUENCE
OF MEDIA ON MODELLING
AGGRESSIVE OR RISKY DRIVING
Various research methods have been used to
investigate the influence of media on
386 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
modelling aggressive or risky driving. These
include qualitative techniques (e.g., meta-
analyses, surveys, correlation, and time series)
and quantitative techniques (e.g., experimen-
tal studies). In a meta-analysis involving 88
empirical studies investigating the effects of
different types of risk-glorifying media on
cognitions, affect, and behaviours, small to
moderate effects were found (Fischer et  al.,
2011). These effects were documented across
all types of research designs (e.g., experi-
mental, longitudinal, correlational) and for
both males and females, though the effects
were larger for males and younger partici-
pants (i.e., less than 24 years old). The largest
effects were found when the stimuli in the
risk-glorifying media matched the context of
the response measured (e.g., risky driving
behaviour was most strongly influenced by
risky driving media) and when the media
exposure was active versus passive (i.e.,
engaging in video game play versus viewing
media; Fischer et al., 2011).
The Entertainment Software Association
(ESA) estimates that more than 164 mil-
lion Americans play video games, with an
estimated 60% of gamers being male (ESA,
2019a). One study surveyed over 4,000 US
high school students, aged 14 to 18 years, and
found the estimated proportion of male gam-
ers to be as high as 75% (Desai et al., 2010).
The popularity of this industry is reflected
in its 2019 generated revenue, estimated to
be over $43 billion in the United States and
$135 billion globally (ESA, 2019a). Grand
Theft Auto V was the third-best-selling video
game of 2017 (ESA, 2019b), suggesting that
a large proportion of young males are engag-
ing in gaming content that promotes aggres-
sive and risky driving behaviours.
Active engagement in aggressive and risky
driving video games has been found to influ-
ence arousal levels and risk-promoting cog-
nitions. Driving games, such as Need for
Speed, Burnout, and Midnight Racer require
‘massive’ violations of traffic rules to win
(Fischer et al., 2007). A 20-minute session of
playing these games was shown to produce
higher levels of self-reported arousal and
more risk-related cognitions, compared to
a control group who played more neutral
games, such as Tak, Crash Bandicoot or FIFA
2005 (Fischer et  al., 2007). No sex differ-
ences were demonstrated, which suggests this
media content influenced males and females
in the same manner. These games were also
shown to influence behaviour more specific
to driving, as measured with the Vienna Risk-
Taking Test. Males demonstrated greater
risk-taking behaviour than females, as indi-
cated by longer reaction times to abort risky
driving manoeuvres depicted in video clips
(Fischer et al., 2007). The influence of video
game exposure on this task was also demon-
strated to last a period of 24 hours (Fischer
et al., 2009).
Racing or ‘drive ‘em up’ video games are
accessible and can be played by young indi-
viduals who have never experienced driving
in the real world. This could influence their
perception of risk on the roads and contribute
to the formation and use of memory scripts
associated with risky driving behaviour. In
one study, unlicensed adolescents indicated
how often they played racing video games
(e.g., Gran Turismo, Burnout) and then, two
years later, as licensed drivers, indicated their
risk-taking attitudes and self-reported driv-
ing behaviour for speeding, fun riding, and
drinking and driving (Beullens et al., 2011a).
Even after controlling for measured person-
ality traits of sensation seeking and aggres-
sion, video game playing was a significant
predictor of positive attitudes and actual self-
reported driving behaviour related to speed-
ing and fun riding. Video game playing was a
significant predictor of self-reported speeding
only in males, suggesting there may be moder-
ating effects of sex on specific aggressive or
risky driving behaviours. Video game play-
ing was not associated with positive attitudes
towards drinking and driving nor with more
self-reported drinking and driving, suggesting
the modelling was specific to the content of the
media to which the participant was exposed
(Beullens et al., 2011a).
 EVOLUTIONARY PSYCHOLOGY AND DANGEROUS DRIVING BEHAVIOUR
The influence of more passive types of
media (i.e., movies and television) on the
modelling of aggressive or risky driving has
also been studied. Early exposure to reck-
less driving (e.g., fast, careless) depicted
in movies was found to influence adoles-
cents’ self-reported driving behaviour four
years later (Kostermans et  al., 2014). Self-
report of unsafe driving behaviours included
exceeding the speed limit, weaving in and
out of traffic, and driving without a seatbelt
fastened. Higher levels of sensation seek-
ing, measured using the Arnett Inventory of
Sensation Seeking (Arnett, 1994), were also
found to be a significant predictor of reckless
driving. Males with higher sensation seeking
were more likely to report driving without
the use of a seatbelt, compared to females or
those with lower levels of sensation seeking
(Kostermans et al., 2014).
Self-reporting of driving behaviour, par-
ticularly behaviours considered aggressive
or risky, may not produce accurate results.
Participants may alter their responses to
appear more socially desirable, which could
produce under-reporting of aggressive or
risky behaviour (e.g., more acceptable to
society as a whole), as well as over-­reporting
(e.g., an attempt to match a personal ideal
of a ‘cool’ driver within a social circle of
friends). Simulated driving has been used
to investigate the effects of watching risk-­
glorifying media on subsequent driving
behaviour. In one study, participants watched
either a risk-promoting movie scene (e.g., a
segment from a James Bond movie), or a neu-
tral scene from a local talk show, and then
drove on a simulator within the context of the
racing video game Need for Speed (Fischer
et al., 2008). Controlling for experience with
racing games, significant differences were
found, such that the risk-promoting condi-
tion produced higher maximum speeds and
more collisions for both males and females,
and less time to complete the entire course
for males (Fischer et al., 2008). Though this
experimental research used a measure closer
to realistic driving (i.e., Vienna Risk-Taking
387
Test) and controlled the exposure to risk-
glorifying media, the media content was not
specific to aggressive or risky driving behav-
iours. Additionally, the simulated driving was
in the context of a racing video game, which
could have influenced participants’ behav-
iour through expectations of performance
and a lower perceived risk for driving aggres-
sively. Lastly, this research did not consider
other mediating factors of arousal, sensation
seeking, trait aggression, or driving history.
Using archival data in the context of a
descriptive research approach, Vitaglione
(2012) analysed aggressive driving accident
reports (i.e., needless dangerous behaviours
or increased risk of harm to other drivers), in
relation to the timing of televised NASCAR
events. West Virginia was selected as the test
state because it was ranked first in the United
States for the number of NASCAR fans per
capita and it does not have its own NASCAR
track (i.e., the predominant source of view-
ing of these events is television). A period of
one week surrounding each of the 156 broad-
casted NASCAR events, between 2003 and
2006, was used to investigate the cumulative
modelling effects of mass media viewing of
aggressive driving. This included the day
before the event, the day of the event, and
was limited to the five days immediately fol-
lowing the event, since NASCAR races could
occur within one week of each other. Using
a time-series regression analysis, the rate
of accidents and injuries due to aggressive
driving was regressed onto NASCAR dates
(Vitaglione, 2012).
The number of aggressive driving acci-
dents significantly decreased on the day of the
NASCAR events, possibly because fans were
watching the televised event and not engag-
ing in much driving that day (Vitaglione,
2012). Both accidents and injuries due to
aggressive driving significantly increased
on the fifth day following the event, but not
immediately following the race. During this
same time period, alcohol-related accidents
and injuries did not change in the same way,
suggesting the specific media content was
388 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
associated with the m ­ odelling of s­pecific
risky and aggressive driving behaviours
(Vitaglione, 2012). The delay in the effects
of the media content may have been due to
resulting arousal from viewing a NASCAR
race becoming associated with dangerous
or aggressive driving behaviours over time
(Vitaglione, 2012). In the context of the
GAM, the more an individual watches these
races, the stronger the links become between
arousal and associated aggressive memories,
which include thoughts, feelings, and behav-
iours. When an individual experiences similar
arousal in a driving scenario, as the result of
provocation, the associated aggressive driving
memory script may be activated or primed and
become consciously accessible. It may require
a number of days for these to be sufficiently
activated in order to facilitate dangerous and
risky driving (Vitaglione, 2012). However, an
alternative explanation is that more immedi-
ate risky and aggressive driving behaviour
was occurring, but drivers had escaped unde-
tected until the passage of time, resulting in
an accident or injury days later. Therefore, the
measure used in this research was not sensi-
tive enough to detect more immediate model-
ling effects (e.g., speeding) of this aggressive
or risky driving media content.
The United States National Highway
Traffic Safety Administration suggested that
excessive speeding be included in the defi-
nition of aggressive driving, following con-
sensus among focus-group participants who
considered it aggressive, as well as aggressive
drivers’ confessions of speeding more fre-
quently (Tasca, 2000). Though the intent of
speeding behaviour may not always be known,
the act of speeding has been used in research
as a measure of risky driving (Iversen and
Rundmo, 2002; Jonah et  al., 2001; Simons-
Morton et  al., 2005) and it is considered a
frequent antecedent behaviour to accident or
collision (Transport Canada, 2011).
Building on the descriptive research
approach of Vitaglione (2012), Singhal (2018)
used archival speeding infraction data from
Edmonton, Alberta, to investigate whether
increases in this specific risky driving behav-
iour occurred following the theatrical release
of two Fast and Furious franchise movies,
Fast and Furious 6 and Furious 7. Not all
aggressive or risky driving results in an acci-
dent or collision, and, therefore, using the
measure of speeding may produce a more
accurate estimate of the amount of aggressive
or risky driving on the roads following such
media exposure. Both Fast and Furious mov-
ies contain content that focused on aggres-
sive or risky driving (i.e., car chases, racing,
excessive speeding) and had a large view-
ership, based on their domestic box office
performance (approximately $239 and $353
million, respectively) (Nash Information
Services, n.d.-a).
Pre-movie-release speeding behaviour
was compared to post-movie-release speed-
ing behaviour, with an emphasis on assessing
effects immediately following movie release.
A four-week post-movie-release period was
selected and compared to a four-week pre-
movie-release period. These time periods
were selected based on the steep trend in
the domestic box office profits for the first
30 days post movie release (Singhal, 2018).
Figure 19.2 shows the daily cumulative
domestic box office for the eight Fast and
Furious movies (Nash Information Services,
n.d.-a). The dotted line estimates the point at
which the movies had played in theatres for
approximately 30 days.
Speeding infractions surrounding the
release dates of Fast and Furious 6 and
Furious 7 (May 24, 2013 and April 3, 2015,
respectively) were provided by the Office
of Traffic Safety in Edmonton, Alberta. All
infractions included in the study were a
result of automated enforcement using sta-
tionary cameras (i.e., photo radar). These
cameras were permanently mounted, oper-
ating 24 hours a day, seven days a week,
which ensured stable enforcement schedules
over the selected time periods. The close
proximity of cameras to theatres (i.e., five
kilometres) allowed for the assessment of
more immediate modelling effects of movie
 EVOLUTIONARY PSYCHOLOGY AND DANGEROUS DRIVING BEHAVIOUR
Figure 19.2  Domestic box-office history for the Fast and Furious movies
content, though it was understood that closer
distances were confounded with time (i.e.,
speeding detected closer to a theatre could
also reflect speeding which occurred a short
time after movie viewing). Infractions were
issued to the registered owner of the vehicle,
rather than the driver, since only the vehi-
cle was identified as being involved in the
offence. Unfortunately, personal identifiers,
revealing driver and vehicle characteristics,
were not available in this study, limiting the
investigation of how these factors influence
speeding behaviour (Singhal, 2018).
An interrupted time-series approach was
used to assess whether movie release was
related to increases in speeding. The speed-
ing infraction data were linked in time, such
that data points were recorded consecutively
on a daily basis. The interrupted approach
allowed for the specification of the movie
release date as a possible intervention in the
time series, investigating whether changes in
this driving behaviour occurred. Coding of
specific days and intervention time periods
(e.g., first weekend post movie release) was
used to assess patterns of behaviour change
post-intervention. Other variables of daily
389
precipitation (mm) and traffic volume were
included in the analyses (Singhal, 2018), due
to their influence on speeding (World Health
Organization, 2004).
Results revealed increases in aggressive
or risky driving behaviour (i.e., speeding)
when Furious 7 was released and playing
in theatres. Significant increases in speed-
ing infractions were found specifically for
the first weekend (i.e., opening weekend)
and the first week, post movie release. A sig-
nificant increase in mean speed differential
was also found for the first weekend, which
means that not only did the number of speed-
ing infractions increase, but also the speed
at which drivers were caught speeding was
greater. These effects were significant even
after controlling for precipitation, traffic
volume, Saturdays, Sundays, and autocor-
relation in the data. This timeline of change
was different from that reported for aggres-
sive driving accidents following exposure to
televised NASCAR races (Vitaglione, 2012),
which, as mentioned previously, was likely
due to the use of a more sensitive measure
of modelling aggressive or risky driving
(Singhal, 2018).
390 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Though Furious 7 was released during an
Easter holiday, comparison analyses, using
time periods surrounding the previous two
Easter holidays, failed to demonstrate the
same significant changes in speeding infrac-
tions and mean speed differential, during the
post-period. Overall, these results suggest
that early modelling effects of aggressive
driving (i.e., speeding) can occur following
the release of movies containing aggressive
or risky driving content (Singhal, 2018).
Similar modelling effects were not found
for Fast and Furious 6, but the difference
in domestic and worldwide box office gross
(Nash Information Services, n.d.-c n.d.-a)
suggests Furious 7 had a greater viewership
(Singhal, 2018). Also, Furious 7 received
greater news coverage due to the accidental
death of one of the lead actors (Paul Walker)
prior to the completion of filming his role in
Furious 7 (Ying, 2015). Walker was killed in
a high-speed collision unrelated to the movie
production. Curiosity surrounded how the
director would handle the actor’s death in
the movie and the ongoing storyline of the
franchise. Even though the more recent fran-
chise instalment Fate of the Furious broke
the global box office record for an opening
weekend (April 14, 2017) (Nash Information
Services, n.d.-b), it did not surpass the domes-
tic box office gross for the opening weekend
of Furious 7 (approximately $98 million
compared to $147 million; Nash Information
Services, n.d.-d).
To further demonstrate the capacity of
aggressive or risky driving content to influ-
ence the modelling of this driving behav-
iour, Singhal (2018) used an experimental
research approach that incorporated multiple
intervening factors together in a single study.
University students were exposed to one of
three video clips, which differed in the level
of aggressive and risky driving content or
arousal, and then drove through a set course
on a simulator. Measures of speed, accelera-
tion, and overtaking or passing actions were
used to assess risky driving, and individual
factors of age, sex, sensation seeking, trait
aggression, driver anger, driver vengeance,
driving history, and movie and video game
history were also included in the analyses
(Singhal, 2018).
Significant differences in aggressive or
risky driving behaviour as a function of video
condition were not found, though differences
were in the expected direction (e.g., higher
mean speed, shorter time to completion, and
greater passing frequency). Some issues sug-
gested to account for the lack of statistical sig-
nificance were the use of videos with low plot
content, a driving simulator test course with
too few opportunities for speeding, and the
viewing of movie content in an experimen-
tal context and not a typical social environ-
ment, such as a theatre with friends (Singhal,
2018). No sex differences were found for the
behavioural measures of aggressive driving
and it was suggested that the lower number of
male than female participants may have con-
tributed (i.e., a ratio of 1:2) (Singhal, 2018).
Personality factors of higher trait aggres-
sion and sensation seeking were related to
more aggressive and risky driving. In addi-
tion, those with higher driving vengeance and
a history of violations, particularly speeding,
engaged in more aggressive and risky driv-
ing, particularly during a provoking racing
scenario in which two sports cars chased one
another. Given that the majority of violations
reported were for speeding, this suggested
that drivers with a history of speeding were
more influenced by environmental cues pro-
moting this behaviour (Singhal, 2018). These
findings support the interactivity of internal-
state and situation factors associated with
aggressive driving, specifically the contribu-
tion of personality factors and aggressive-
promoting driving cognition (i.e., driving
vengeance) on the choice to drive aggres-
sively or with greater risk. Individuals with
these characteristics may be more suscepti-
ble to aggressive and risky driving content in
movies.
Higher levels of sensation seeking and
trait aggression were both related to higher
levels of driving anger, driving vengeance,
 EVOLUTIONARY PSYCHOLOGY AND DANGEROUS DRIVING BEHAVIOUR
and number of violations, suggesting these
personality factors played a role in aggressive
driving cognition and behaviour. Sensation
seeking was also positively correlated with
the number of aggressive driving movies
viewed in the past two years and the greater
viewing of these movies was associated with
higher levels of driving vengeance, which was
related to more violations (Singhal, 2018).
Some future research questions that arise
from the Singhal (2018) studies include the
longevity or duration of the cinematic influ-
ences and whether generalization occurs,
such that risky behaviours manifest in other
realms. For example, will drivers who speed
after viewing Fast and Furious movies now
swim further from shore or be predisposed to
gambling or engaging in risky sex when the
situation presents itself?
THEORIES OF MODELLING OF
AGGRESSION REVISITED
The results of the two studies by Singhal
(2018) offer support to theories of modelling
aggression and contribute to a better under-
standing of how aggressive and risky driving
occurs. Both studies support the social learn-
ing theory of modelling aggression (Bandura,
1971). Evidence of greater speeding sur-
rounding the release of Furious 7 suggested
that the modelling of aggressive or risky
driving behaviour was associated with the
release of this movie, which portrayed this
behaviour being performed by a relatable
‘hero’ and, predominantly, in the absence of
negative consequences. Trends in the second
study provided support for this theory of
modelling aggression, though differences
between the video conditions were not sig-
nificant. It is important to consider that these
studies used different cohorts (i.e., speed
violators versus undergraduate psychology
research pool participants). The speed viola-
tors in the first study would have been more
likely to possess the characteristics found in
391
the second study to contribute to aggressive
or risky driving (e.g., higher trait aggression,
sensation seeking, and driving vengeance).
Demographics were not known for the speed
violators, but, according to the evolutionary
theory of modelling aggression (Mesquida
and Wiener, 1996; Vingilis et  al., 2013;
Wiesenthal and Singhal, 2012; Wilson and
Daly, 1985), the majority of these individuals
were likely young males, who experienced a
higher level of anonymity on the roads, com-
pared to driving in a laboratory setting.
Findings associated with higher levels of
sensation seeking suggest that these personal-
ity types may seek out and view more aggres-
sive and risky driving movies. This, in turn,
contributes to the formation of aggressive and
risky driving scripts, which can produce cer-
tain susceptibilities for the activation of these
scripts at a later time, with further viewing of
aggressive and risky driving movie content.
The eventual engagement in the modelling
of this behaviour may be more likely when
the driver is exposed to a provoking driving
scenario (e.g., racing or vengeance related).
These relationships support the interactivity
of the person, situation, and internal factors
of the modified GAM, with respect to mod-
elling aggressive and risky driving (Figure
19.1), and also support Bandura’s (2001b)
Triadic Reciprocal Causation model of psy-
chosocial functioning.
CONCLUSION
Given the profits associated with movies
containing aggressive or risky driving con-
tent, a halt in their production is not likely to
occur. The large domestic box office gross
associated with the Fast and Furious fran-
chise demonstrates that a large fan base
exists for this genre of movie. The next
instalment, Fast and Furious 9, has an antici-
pated release date of April 2, 2021 (IMDb,
2019). There are also indications of popular-
ity for other forms of media depicting acts of
392 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
aggressive and risky driving. As mentioned
previously, the top 10 YouTube videos con-
taining this content, particularly street racing,
average over 700,000 views and ‘likes’
(Vingilis et  al., 2017). These videos, which
can be freely accessed and repeatedly viewed,
are unlike movies in that the majority depict
real-world extreme driving events that were
not performed by professionals in a staged
environment. Research has not begun to
understand how this content influences a
person’s driving behaviour (Vingilis et  al.,
2017). Almost 20 years ago, Bandura (2001a:
271) recognized the ‘accelerated growth of
video delivery technologies’ and how they
facilitate exposure to a wide range of behav-
iours and ‘models’. Social media has
expanded that substantially since then.
A complete halt in production of movies
such as Furious 7 or other forms of media
depicting acts of aggressive or risky driving is
not a realistic solution to reducing the model-
ling of this behaviour. However, the prolifera-
tion of this content makes it important to raise
awareness about the association between this
material and one’s driving behaviour, particu-
larly the dangers associated with modelling.
Singhal’s (2018) finding of portrayals of this
behaviour in the movies having a significant
impact on real-world speeding behaviour
supports increasing enforcement when these
movies are released and playing in theatres.
This is particularly true for opening week-
ends and the first week post movie release.
Speeding is a risk factor in road traffic inju-
ries and fatalities, because of the increased
risk of collision and severity of the resulting
consequences. It is estimated that 30–50%
of mortality on the roads is associated with
speeding (World Health Organization, 2004).
Alternate strategies could be more proac-
tive, targeting the movie-goer and raising
awareness of the potential for modelling the
unsafe driving behaviour. This could include
messages from the actors themselves, urging
people to be responsible and smart on the
roads and to drive safely. Universal Studios
did this in the past for the Fast and Furious
franchise (Orwall, 2001) and production
companies could incorporate these messages
into the credits of the movie, following its
completion. It may intrigue the movie-goer
to view additional footage of the actors and
the message may be more meaningful com-
ing from these individuals, who are revered
in their role. This may also delay the movie-
goers from returning to their vehicles imme-
diately following the end of the movie,
potentially allowing for some decrease in
their immediate level of arousal.
Lastly, we should not remove account-
ability from the viewer. Bandura’s early
work (Bandura, 1971; Bandura et al., 1963a)
demonstrated the basic principles of observa-
tional learning and modelling, and his later
reflections emphasized that human behaviour
is not simply the result of stimulus-response
relationships. We, as humans, have ‘func-
tional consciousness’, part of which involves
regulating and evaluating the actions we
choose to take (Bandura, 2001b). The two
core features of Bandura’s (2001b) proposed
‘human agency’ are self-reactiveness and self-­
reflectiveness, mentioned previously. It is these
two elements that are emphasized to viewers
of aggressive and risky driving media content.
Viewers make choices to expose themselves
to this content and, therefore, have a respon-
sibility to be aware of how this content can
influence their behaviour, and to evaluate their
actions accordingly (e.g., considering driv-
ing laws and the safety of others). Essentially,
Bandura (2001b) is highlighting the appraisal
and decision-making process of the modi-
fied GAM, as shown in Figure 19.1. Having
viewers engage in self-reactiveness and self-­
reflectiveness could decrease the probability
of an impulsive action and increase the prob-
ability of a thoughtful one. This would, ulti-
mately, hamper the modelling of aggressive
and risky driving following exposure to such
content in the media.
Drivers must be cognisant of the fact that
they share the road with other drivers, vehi-
cle occupants (e.g., children), cyclists, and
pedestrians. With this in mind, all drivers
 EVOLUTIONARY PSYCHOLOGY AND DANGEROUS DRIVING BEHAVIOUR
should consider characteristics that consti-
tute a ‘good’ driver. Qualities which lessen
the occurrence of factors that contribute to
aggressive driving should be encouraged. For
example, forethought, in the context of driv-
ing, would involve allowing sufficient time
to arrive at a final destination, which could
prevent the development of frustration when
confronted with delays in traffic or other
hampering driver actions. Forbearance (i.e.,
self-restraint and tolerance) and forgiveness
of other drivers’ minor mistakes (e.g., driving
too slowly in the fast lane) could lessen the
likelihood of choosing an aggressive driving
response. Factors such as these, in addition
to being aware of the possibility of aggres-
sive driving motion-picture content influenc-
ing driving behaviour, are encouraged in all
drivers. Young males may need to be par-
ticularly aware of heightened sensitivity to
provocation or competitiveness on the roads
and appropriately inhibit aggressive or risky
responses. Just as many drinking and driving
campaigns target the driver in emphasizing
the dangers and potentially tragic conse-
quences of impaired driving (Green, 2013;
MADD, 2017), research has highlighted the
need for drivers to keep their safety, and the
safety of the public, at the forefront when
considering engaging in an aggressive or
risky driving behaviour.
REFERENCES
Abelson, R. P. (1981). Psychological status of
the script concept. American Psychologist,
36, 715–729.
Anderson, C. A. & Bushman, B. J. (2002). Human
aggression. Annual Review of Psychology,
53, 27–51.
Arnett, J. (1994). Sensation seeking: A
new conceptualization and a new scale.
Personality and Individual Differences, 16,
289–296.
Atkin, C. K. (1989). Television, socialization and
risky driving by teenagers. Alcohol, Drugs,
and Driving, 5, 1–11.
393
Bandura, A. (1971). Psychological modeling:
Conflicting theories. New Jersey: Aldine
Transaction.
Bandura, A. (2001a). Social cognitive theory of
mass communication. Media Psychology, 3,
265–299.
Bandura, A. (2001b). Social cognitive theory:
An agentic perspective. Annual Review of
Psychology, 52, 1–26.
Bandura, A., Ross, D., & Ross, S. A. (1963a).
Imitation of film-mediated aggressive models.
Journal of Abnormal and Social Psychology,
66, 3–11.
Bandura, A., Ross, D., & Ross, S. A. (1963b).
Vicarious reinforcement and imitative learn-
ing. Journal of Abnormal and Social Psychol-
ogy, 67, 601–607.
Berkowitz, L. (2012). A cognitive-neoassociation
theory of aggression. In P. Van Lange, A.
Kruglanski, & E. Higgins (Eds.), Theories of
Social Psychology (Vol. 2., pp. 99–117),
London: Sage.
Beullens, K., Roe, K., & Van den Bulck, J.
(2011a). Excellent gamer, excellent driver? The
impact of adolescents’ video game playing on
driving behavior: A two-wave panel study.
Accident Analysis and Prevention, 43, 58–65.
Beullens, K., Roe, K., & Van den Bulck, J. (2011b).
The portrayal of risk-taking in traffic: A content
analysis of popular action movies. Journal of
Communications Research, 2, 21–27.
Bushman, B. J. (1995). Moderating role of trait
aggressiveness in the effects of violent media
on aggression. Journal of Personality and
Social Psychology, 69, 950–960.
Buss, D. M. (1988). The evolution of intrasexual
competition: Tactics of mate attraction.
Journal of Personality and Social Psychology,
54, 616–628.
Buss, A. H. & Perry, M. (1992). The aggression
questionnaire. Journal of Personality and
Social Psychology, 63, 452–459.
Coleman, L. (2004). The copycat effect: How
the media and popular culture trigger the
mayhem in tomorrow’s headlines. New York:
Paraview Pocket Books.
Dahlen, E. R., Martin, R. C., Ragan, K., &
Kuhlman, M. M. (2005). Driving anger, sen-
sation seeking, impulsiveness, and boredom
proneness in the prediction of unsafe
driving. Accident Analysis & Prevention, 37,
341–348.
394 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Deb, S. (2019, January 2). Netflix to ‘Bird Box’
fans: Please open your eyes. The New York
Times. Retrieved January 14, 2019, from
www.nytimes.com/2019/01/02/arts/net
flix-bird-box-challenge-war ning.html?
module=inline
Desai, R. A., Krishnan-Sarin, S., Cavallo, D. &
Potenza, M. N. (2010). Video-gaming among
high school students: Health correlates,
gender differences, and problematic gaming.
Pediatrics, 126(6), e1414-e1424. Retrieved
from https://pediatrics.aappublications.org/
content/126/6/e1414
Eron, L. D., Huesmann, L. R., Lefkowitz, M. M.,
& Walder, L. O. (1972). Does television vio-
lence cause aggression? American Psycholo-
gist, 27(4), 253–263.
Entertainment Software Association (ESA)
(2019a). ESA leadership desk: Telling our
industry’s story in a new way. Retrieved June
7, 2019, from www.theesa.com/perspectives/
esa-leadership-desk-telling-our-industrys-
story-in-a-new-way/
Entertainment Software Association (ESA)
(2019b). 2018 Essential facts about the com-
puter and video game industry. Retrieved
June 7, 2019, from www.theesa.com/
esa-research/2018-essential-facts-about-the-
computer-and-video-game-industry/
Fischer, P., Greitemeyer, T., Kastenmüller, A.,
Vogrincic, C., & Sauer, A. (2011). The effects
of risk glorifying media exposure on risk-
positive cognitions, emotions, and behav-
iours: A meta-analytic review. Psychological
Bulletin, 137, 367–390.
Fischer, P., Greitemeyer, T., Morton, T., Kasten-
müller, A., Postmes, T., Frey, D., Kubitzki, J.,
& Odenwälder, J. (2009). The racing-game
effect: Why do video racing games increase
risk-taking inclinations? Personality and
Social Psychology Bulletin, 35, 1395–1409.
Fischer, P., Guter, S., & Frey, D. (2008). The
effects of risk-promoting media on inclina-
tions toward risk taking. Basic and Applied
Social Psychology, 30, 230–240.
Fischer, P., Krueger, J. I., Greitemeyer, T., Asal, K.,
Aydin, N., & Vingilis, E. (2012). Psychological
effects of risk glorification in the media:
Towards an integrative view. European Review
of Social Psychology, 23, 224–257.
Fischer, P., Kubitzki, J., Guter, S., & Frey, D.
(2007). Virtual driving and risk taking: Do
racing games increase risk-taking cognitions,
affect, and behaviors? Journal of Experimen-
tal Psychology: Applied, 13, 22–31.
Freedman, J. L. (2007). Television violence and
aggression: Setting the record straight. The
Media Institute Policy Views, 1, 1–10.
Goldberg, O. (2001). ‘Fast’ fallout film may
inspire copycat racers. The Free Library.
Retrieved June 8, 2020, from web.archive.
org/web/20151023223740/www.thefreeli-
brary.com/’FAST’+FALLOUT+FILM+MAY+INS
PIRE+COPYCAT+RACERS.-a083598815
Green, M. (2013, July 24). Drunk driving pre-
vention and the effectiveness of media cam-
paigns. Retrieved June 26, 2020, from www.
absoluteadvocacy.org/drunk-driving-prevention-
media-campaigns/
Greenberg, B. S. & Atkin, C. K. (1983). The por-
trayal of driving on television, 1975–1980.
Journal of Communication, 33, 44–55.
Grewal, S. & Brennan, R. (2006, June 23).
Racing film has police on edge. The Toronto
Star, p. B4.
Grey, E. M., Triggs, T. J., & Haworth, N. L. (1989).
Driver aggression: The role of personality,
social characteristics, risk and motivation
(Report No. CR 81). Federal Office of Road
Safety, Australia.
Hegerl, U., Koburger, N., Rummel-Kluge, C.,
Gravert, C., Walden, M., & Mergl, R. (2013).
One followed by many? Long-term effects of
a celebrity suicide on the number of suicidal
acts on the German railway net. Journal of
Affective Disorders, 146, 39–44.
Helfgott, J. B. (2015). Criminal behaviour and
the copycat effect: Literature review and
theoretical framework for empirical investi-
gation. Aggression and Violent Behavior, 22,
46–64.
IMDb (2018, December 21). Bird Box (2018).
Retrieved June 28, 2019, from www.imdb.
com/title/tt2737304/
IMDb (2019, November 11). Fast & Furious 9
(2020). Retrieved June 7, 2020. from www.
imdb.com/title/tt5433138/
Iversen, H. & Rundmo, T. (2002). Personality,
risky driving and accident involvement
among Norwegian drivers. Personality and
Individual Differences, 33, 1251–1263.
Jeong, J., Shin, S. D., Kim, H., Hong, Y. C.,
Hwang, S. S., & Lee, E. J. (2012). The effects
of celebrity suicide on copycat suicide
 EVOLUTIONARY PSYCHOLOGY AND DANGEROUS DRIVING BEHAVIOUR
attempt: A multi-center observational study.
Social Psychiatry and Psychiatric Epidemiology,
47, 957–965.
Jonah, B. A., Thiessen, R., & Au-Yeung, E.
(2001). Sensation seeking, risky driving and
behavioral adaptation. Accident Analysis &
Prevention, 33, 679–684.
Koburger, N., Mergl, R., Rummel-Kluge, C.,
Ibelshäuser, A., Meise, U., Postuvan, V.,
Roskar, S., Székely, A., Ditta Tóth, M., van
der Feltz-Cornelis, C., & Hegerl, U. (2015).
Celebrity suicide on the railway network:
Can one case trigger international effects?
Journal of Affective Disorders, 185, 38–46.
Koordeman, R., Anschutz, D. J., & Engels, R. C.
M. E. (2011). Exposure to alcohol commer-
cials in movie theaters affects actual alcohol
consumption in young adult high weekly
drinkers: An experimental study. American
Journal on Addictions, 20, 285–291.
Kostermans, E., Stoolmiller, M., de Leeuw, R.
N., Engels, R. C., & Sargent, J. D. (2014).
Exposure to movie reckless driving in early
adolescence predicts reckless, but not inat-
tentive driving. Public Library of Science
(PLoS ONE), 9, e113927.
Kotzen, S. (2013, September 25). ‘The Pro-
gram’ turns 20, along with its notorious road
scene (video). The Hollywood Reporter.
Retrieved on June 15, 2015 from www.hol-
lywoodreporter.com/news/program-turns-
20-along-notorious-636568
List of Chess Grandmasters (n.d.). Retrieved Sep-
tember 15, 2019, from https://en.m.wikipedia.
org/wiki/List_of_chess_grandmasters
López-Guimerà, G., Levine, M. P., Sánchez-
Carracedo, D., & Fauquet, J. (2010). Influ-
ence of mass media on body image and
eating disordered attitudes and behaviors in
females: A review of effects and processes.
Media Psychology, 13, 387–416.
MADD (2017, June 9). And then, in a heartbeat,
everything changed! Retrieved on September
15, 2019 from http://madd.ca/pages/and-
then-in-a-heartbeat-everything-changed/
Malamuth, N. M. & Check, J. P. V. (1981). The
effects of mass media exposure on accept-
ance of violence against women: A field
experiment. Journal of Research in Personal-
ity, 15, 436–446.
Mensching, K. (2015, February 22). Daytona
500 purse 2015: Joey Logano take winner’s
395
share of $18 million payday. Retrieved from
www.sbnation.com/nascar/2015/2/22/
8085643/2015-daytona-500-purse-prize-
money
Mesquida, C. G. & Wiener, N. I. (1996). Human
collective aggression: A behavioral ecology
perspective. Ethology and Sociobiology, 17,
247–262.
Milgram, S. & Shotland, R. L. (1973). Television
and antisocial behavior: Field experiments.
New York: Academic Press.
Modeling (n.d.). In Encyclopedia of Mental
Disorders online. Retrieved on June 28, 2015
from www.minddisorders.com/Kau-Nu/
Modeling.html
Nash Information Services (n.d.-a). The Num-
bers: Box office history for Fast and Furious
movies. Retrieved March 1, 2017, from
www.thenumbers.com/movies/franchise/
Nash Information Services (n.d.-b). The Num-
bers: The Fate of the Furious (2017). Retrieved
April 18, 2017, from www.the-numbers.com/
movie/Fate-of-the-Furious-The#tab=summary
Nash Information Services (n.d.-c). The Num-
bers: Furious 7 (2015). Retrieved February
28, 2017, from www.the-numbers.com/
movie/Furious-7#tab=summary
Nash Information Services (n.d.-d). The Num-
bers: Movie comparison: The Fate of the
Furious (2017) vs. Furious 7 (2105). Retrieved
April 20, 2017, from www.the-numbers.
com/movies/custom-comparisons/Fate-of-
the-Furious-The/Furious-7
Orwall, B. (2001, June 21). Fearing copycats,
Universal warns against mimicking new
action film. The Wall Street Journal. Retrieved
from www.wsj.com/articles/SB99307515845
592312?mod=searchresults&page=1&pos=1
Paik, H. & Comstock, G. (1994). The effects of
television violence on antisocial behavior: A
meta-analysis. Communication Research,
21(4), 516–546.
Peng, J. (2018, September 24). Doubling of
motorcycle deaths in B.C. alarms safety offi-
cials. The Star. Retrieved October 10, 2018,
from www.thestar.com/vancouver/2018/09/
24/doubling-of-motorcycle-deaths-alarms-
safety-officials.html
Phillips, D. P. (1974). The influence of sugges-
tion on suicide: Substantive and theoretical
implications of the Werther effect. American
Sociological Review, 39, 340–354.
396 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Phillips, D. P. & Carstensen, L. L. (1988). The
effect of suicide stories on various demo-
graphic groups, 1968–1985. Suicide and
Life-Threatening Behavior, 18, 100–114.
Pristin, T. & Fox, D. (1993, October 23). Pop
culture. Violence. Copycats. Blame?: Holly-
wood debates the responsibility issue after
Disney’s edit of ‘The Program’. Los Angeles
Times. Retrieved February 5, 2016, from
http://articles.latimes.com/1993-10-23/enter-
tainment/ca-48913_1_responsibility-issue
Roseborough, J. E. W. (2014). Retaliatory
aggressive driving: A justice perspective
(Unpublished doctoral dissertation). York
University, Toronto.
Rowland, B. (2006, June 23). CHP concerned
after release of racing movie. Free Lance
News. Retrieved from www.sanbenito.com/
chp-concerned-after-release-of-racing-movie/
Shin, P. C., Hallett, D., Chipman, M. L., Tator,
C., & Granton, J. T. (2005). Unsafe driving in
North American automobile commercials.
Journal of Public Health, 27, 318–325.
Shinar, D. (1998). Aggressive driving: The con-
tribution of the drivers and the situation.
Transportation Research Part F: Traffic Psy-
chology and Behaviour, 1, 137–160.
Simons-Morton, B., Lerner, N., & Singer, J.
(2005). The observed effects of teenage pas-
sengers on the risky driving behavior of teen-
age drivers. Accident Analysis & Prevention,
37, 973–982.
Singhal, D. (2018). Modelling aggressive or risky
driving: The effect of cinematic portrayals of
risky driving (Doctoral dissertation, York Uni-
versity, Toronto). Retrieved from YorkSpace.
(http://hdl.handle.net/10315/34497)
Skinner, B. F. (1938). The behavior of organ-
isms: An experimental analysis. New York:
Appleton-Century-Crofts.
Skinner, B. F. (1953). Science and human
behavior. Toronto: Simon and Schuster.
Steinberg, M. S. & Dodge, K. A. (1983). Attri-
butional bias in aggressive adolescent boys
and girls. Journal of Social and Clinical Psy-
chology, 1, 312–321.
Steinberg, L. & Monahan, K. C. (2007). Age
differences in resistance to peer influence.
Developmental Psychology, 43, 1531–1543.
Talwar, P. (1998). The family and peer group
influences in aggression. Indian Journal of
Psychiatry, 40, 346.
Tasca, L. (2000). A review of the literature on
aggressive driving research. Ontario Advi-
sory Group on Safe Driving Secretariat, Road
User Safety Branch, Ontario Ministry of
Transportation.
Tedeschi, J. T. & Felson, R. B. (1994). Violence,
aggression, and coercive actions. Washington,
DC: American Psychological Association.
Toronto Transit Commission (2010, February 17).
Subway suicide prevention. Retrieved on May
28, 2015 from www.ttc.ca/About_the_TTC/
Commission_reports_and_information/Com-
mission_meetings/2010/Feb_17_2010/Supple-
mentary_Reports/Subway_Suicide_Preve.pdf
Transport Canada (2011). Road safety in
Canada. Retrieved from www.tc.gc.ca/
media/ documents/roadsafety/tp15145e.pdf
US driver in ‘Bird Box blindfold’ crashes in Utah.
(2019, January 11). BBC News. Retrieved
January 14, 2019, from www.bbc.com/
news/world-us-canada-46846981
Vingilis, E., Seeley, J., Wiesenthal, D., Mann, R.,
Vingilis-Jaremko, L., Vanlaar, W., & Leal, N.
(2013). Street racing and stunt driving in
Ontario, Canada: Results of a web-based
survey of car and racing enthusiasts. Trans-
portation Research Part F: Traffic Psychology
and Behaviour, 21, 30–42.
Vingilis, E., Yildirim-Yenier, Z., Vingilis-Jaremko,
L., Wickens, C., Seeley, J., Fleiter, J., &
Grushka, D. H. (2017). Literature review on
risky driving videos on YouTube: Unknown
effects and areas for concern? Traffic Injury
Prevention, 18, 606–615.
Vitaglione, G. D. (2012). Driving under the
influence (of mass media): A four-year exam-
ination of NASCAR and West Virginia
aggressive-driving accidents and injuries.
Journal of Applied Social Psychology, 42,
488–505.
Watson, L., Lavack, A. M., Rudin-Brown, C.,
Burns, P., & Mintz, J. H. (2010). Message
content in Canadian automotive advertising:
A role for regulation? Canadian Public Policy,
36(Supplement 1), S49–S67.
Weiss, B., Dodge, K. A., Bates, J. E., & Pettit, G. S.
(1992). Some consequences of early harsh
discipline: Child aggression and a maladap-
tive social information processing style. Child
Development, 63, 1321–1335.
Wiesenthal, D. L., Lustman, M., & Roseborough,
J. (2016). Aggressive driving. In A. Smiley
 EVOLUTIONARY PSYCHOLOGY AND DANGEROUS DRIVING BEHAVIOUR
(Ed.), Human Factors in Traffic Safety (3rd ed.,
pp. 169–193), Tucson, AZ: Lawyers & Judges
Publishing Company.
Wiesenthal, D. L. & Singhal, D. (2012).
Evolutionary psychology, demography and
driver safety research: A theoretical synthesis.
In C. Roberts (Ed.), Applied Evolutionary
Psychology (pp. 399–413), Oxford, UK:
Oxford University Press.
Wilson, M. & Daly, M. (1985). Competitiveness,
risk taking, and violence: The young male syn-
drome. Ethology and Sociobiology, 6, 59–73.
World Health Organization (2004). Road
safety – Speed. Retrieved on June 15, 2015
397
from www.who.int/violence_injury_preven-
tion/publications/road_traffic/world_report/
speed_en.pdf
Ying, L. (2015, February 6). Hype’s must watch:
Furious 7. Retrieved on May 15, 2015 from
https://hype.my/2015/40192/hypes-
must-watch-furious-7/
Zillmann, D. (1971). Excitation transfer in
communication-mediated aggressive behavior.
Journal of Experimental Social Psychology, 7,
419–434.
Zuckerman, M. (1979). Sensation seeking:
Beyond the optimum level of arousal. Hillsdale,
NJ: Lawrence Erlbaum Associates.

Evolutionary Psychology and
Mass Media
When considering the topic of evolutionary
psychology and mass media, the questions
come very quickly. For example:
• As media has become more a part of human
life, have humans adapted to it in an evolutionary
sense, i.e. has natural selection had a chance to play
a part in the way humans adapt to mass media?
• If survival and reproduction are the two evolutionary
imperatives, which has been the bigger influence in
the way media has developed in the last century?
• If men and women have different reproductive
agendas, how has that affected the different
ways that men and women use media?
• Are mass media and entertainment adaptations,
or are they by-products of adaptations?
• Are there ways that mass media improves human
chances for survival?
• Are there ways that mass media affects our
opportunities to engage in sexual behavior and
thus have more chances to reproduce?
• Or are there ways that mass media inhibits
our sexual behavior and keeps us from having
chances to reproduce?
• What are the individual differences in how media
is processed as affected by both social construc-
tivism vs biological essentialism?
20
Gayle S. Stever
To tackle these questions, one must first define
mass media, also referred to as mass commu-
nication. Mass communication is the result ‘of
forces set in motion when groups of manlike
[sic] animals first huddled together against the
cold and danger of primitive times’ (Schramm,
1960: 3). News media is relevant because of
the biological imperative for survival and the
function of news to give us information about
our environment that will affect the likelihood
of that survival (Grabe, 2011; Shoemaker,
1996). Are those the primary origins and func-
tions of mass media, or are there others? This
chapter attempts to address these issues and
questions.
ORIGINAL USES FOR MEDIA
What was the first form of media and for
what was it used? Schramm (1988) described
the discovery of early cave paintings created
in prehistoric times and speculated as to their
purpose. The most likely use of these early
 EVOLUTIONARY PSYCHOLOGY AND MASS MEDIA
paintings was for one generation of people to
educate the next as to what the customs and
practices of that culture had been. It is possi-
ble that these early pictures represented magi-
cal beliefs and secrets of that group (Schramm,
1988), shared with the youth of each succes-
sive generation. Learning about the ways and
skills needed for survival was an essential part
of being socialized and insured that the group
would survive. In a basic fundamental way,
the first media was an integral part of the sur-
vival of individuals.
It has been suggested that the progress and
evolution of technology in human history
proceeded along almost Lamarckian lines, if
the inventions of one generation are able to be
communicated to a subsequent generation via
some media representation. If adaptive capac-
ity is improved by tools that are shared from
one generation to the next via media represen-
tations, it can be speculated that brain capacity
is shaped and accelerated via this ‘inherited’
shared knowledge. This is an example of the
way that interaction between culture and biol-
ogy can accelerate the development of knowl-
edge that affects the survival of individuals,
thus increasing the chance that their genes
are passed on. Those whose genetic potential
best leads to this kind of communication are
the ones who will survive to reproduce. Thus,
media plays a critical role in Darwinian evolu-
tion (Toth and Schick, 1993).
From those earliest points moving for-
ward, media continued to provide a way
that knowledge, history, and social struc-
tures were conveyed from one generation to
the next. Education has been a fundamen-
tal function of media and it is an important
way to increase one’s odds of surviving to
reproduce. Media provides the cornerstone
of societal norms, traditions, and skills for
preserving knowledge for future success and
continuation of a social group.
As media developed more sophisticated
forms of delivery, it provided better rep-
resentations of day-to-day reality, starting
with drawings on cave walls and progress-
ing to the current forms of media that include
virtual reality, a complete recreation of a
399
symbolic environment within an imaginary
and representational world. These mediated
worlds are possible as technology becomes
more sophisticated and it becomes possible
to create more lifelike images. Education and
also enhanced experience are made possible
through these mediated formats.
For example, The Henry Ford Museum in
Dearborn, Michigan featured a film in 2018
about the true story of explorer Henry Bates’
11-year journey through the Amazon rainfor-
est. Using IMAX technology, the viewer is
transported to the Brazilian rainforest where
a trip down the Amazon is experienced, one
that few people would actually be able to
physically undertake. This expanded world-
view is possible because increasingly realis-
tic media forms make it so.
CONSIDERING MEDIA LIFE
It has been argued that we live ‘in’ rather
than ‘with’ media in the 21st century (Deuze,
2011). If this is so, then it becomes important
to consider whether humans have adapted to
media, or have simply assimilated it into the
non-mediated aspects of their lives. If you,
the reader, find yourself doubting that this
could be so, that may be because as media
becomes a greater part of our lives, it also
becomes more invisible to us on a conscious
level. As we become habituated to the pres-
ence of media in our lives, we become less
aware of its presence and influence on a day-
to-day or even minute-to-minute basis.
Think of it as like electricity. There was a
time in human life when electricity was not
a pervasive part of every aspect of life. At
that time, it was easy to sort out what parts
of life involved electricity and what parts did
not. As it became more available, fewer and
fewer things were done apart from electric-
ity. Cooking, cleaning, reading, even outdoor
camping became infused with electricity to
the point where while once we considered
whether a situation involved electricity, now
it is taken for granted that all parts of our life
400 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
involve use of electricity. In addition, and
more important for this analogy, we do not
think or talk about it much at this point in
human history. It is a non-discursive aspect
of our shared lives together.
Mass media is becoming like that. We
spend so much time in mediated worlds that
we no longer consider the mediated parts of
our lives as being separate from the rest of
our lives. Consider what happens to you dur-
ing a power failure. This recently happened
to me, and while I was seeking alternate
forms of power for basic needs (lighting,
cooking, warmth), I was also seeking forms
of media that did not require my home’s elec-
tricity (books, newspapers, battery-powered
radios, my cell phone that didn’t depend on
my local electricity for a signal). When the
availability of the media I’m used to having
was cut off, I was sent scrambling to replace
those mediated sources of information and
interaction with alternative forms to meet
those needs that have developed as a result
of having mass media readily available, each
and every day.
As media becomes more pervasive, it
becomes more important to consider how
humans are developing to adapt to this major
change in our ways of living. Or are they?
This is an important question.
SURVIVAL AND REPRODUCTION
Natural selection is the cornerstone of evolu-
tion, and what drives natural selection are
traits that facilitate either survival or repro-
duction, as traits that favor these things are
more likely to be passed on to the next gen-
eration (Miller, 2011; West-Eberhard, 1979).
It makes sense that any aspect of media
experience that improves chances either for
survival or for reproduction will be liable,
over time, to shape the evolution of our spe-
cies. However, mass media has not been in
play long enough for this to have been a
factor in human evolution thus far. So rather
than looking for the effects of mass media on
evolution, it is better to consider how human
beings as they have already evolved are able
to adapt to a mediated environment. Because
our culture is dominated by various forms of
media, it could be argued that those who are
best able to survive and reproduce in such an
environment will pass their genes on to off-
spring. This has always been the case with
social environments as they have shaped
natural selection in the past. Another way to
say this is that to the extent that our environ-
ment becomes more media saturated, it
becomes more essential to adapt to this type
of environment or not be successful in the
tasks of survival and reproduction. Changing
the overall structure of human inherited traits
is a very slow process (Miller, 2011). A fun-
damental principle of evolution is that cur-
rent members of a species are born with the
set of traits that better prepared their ances-
tors for survival and reproduction. This pre-
sumes that environments stay the same. If
there are radical changes in an environment,
the species must adapt or it will become
extinct. Humans have adapted to pay atten-
tion to any stimuli that have adaptive signifi-
cance. The suggestion is that, rather than
developing new structures in the brain to
facilitate media processing, media has been
created by the brain that has already devel-
oped. An ‘old’ brain isn’t necessarily an
outmoded brain (Grabe, 2011). It remains to
be seen if the addition of mass media is
a large enough change to affect humans
enough to change their survival trajectory
(Geher, 2014).
Nature or Nurture?
Any time one considers aspects of human
growth and development, it is necessary to
weigh in on the nature/nurture debate and
attempt to identify which influences have a
biological basis and which influences have a
basis in socialization. In communication
studies, the tradition has been to look at
 EVOLUTIONARY PSYCHOLOGY AND MASS MEDIA
nurture over nature and, until recently, this
was a driving force in most media research
(Sherry, 2004).
More recently, there has been a shift
towards consideration of the effects of biol-
ogy on communication (Sherry, 2015; Weber,
2015). For example, the field of strategic
communications is the study of how com-
munication satisfies a specific long-term
goal. It has explored the specific ways that
understanding improves as one comprehends
the purposes, in an adaptive sense, that were
fulfilled by the development of mental strate-
gies. Studying motivations such as status or
affiliation sets the stage for a larger under-
standing of how human motivations adapt
to favor both survival and reproduction
(Seiffert-Brockmann, 2018).
Humans have not yet evolved as the result
of mass media (Grabe, 2011). Evolution is
a long-term process, and such adaptations
have not yet had time to emerge. Therefore,
the current conversation is about how the
human brain as it exists today responds to
mass media, and how, moving forward,
media might influence future evolution. The
response to media has been influenced both
by the forces of social constructivism and
biological and inherited aspects of human
development (Bandura, 2001; Sherry, 2015).
If the argument can be made that heritable
traits influence the way we process media
(Toth and Schick, 1993), then an argu-
ment could be made that humans might be
expected to adapt and evolve in the future to
an increasingly mediated world.
Sherry (2004) was one of the early commu-
nication scholars to engage this issue of social
constructivism vs biological essentialism. He
observed that, up until that time, communi-
cation research had been more influenced by
learning theory and environmental determin-
ism than it had been by recognition of biologi-
cal aspects of development. He recognized
that more adaptive traits are likely to be the
ones passed on to a new generation. He dis-
cussed the need for communication scholars to
look more closely at biology and neuroscience
401
to find factors that explain a greater percent-
age of the variability in how humans process
media. Studies to that time had achieved lim-
ited success in making such predictions. In
explaining why communication had been
dominated by the nurture aspects of develop-
ment, he pointed out that the various theo-
retical traditions that had informed research in
mass media were ideas like Bandura’s Social
Learning Theory (Bandura, 2001) or Mead’s
Symbolic Interaction (Mead, 1934), theories
that were more driven by social constructivism
than by considerations of inherited traits. By
2015, Sherry was writing about a newer model
of communication research, one based on
‘complexity theory’ that had integrated con-
siderations like evolution into the prevailing
research paradigms. Indeed, a shift in think-
ing by not only Sherry but also others in the
field (Falk et  al., 2015; Weber, 2015) meant
that mass media and communication scholars
recognized that evolutionary psychology was
becoming an important part of understanding
media effects.
In justifying a model that blends theoretical
aspects of both nature and nurture, Malamuth
(1996: 11) said this: ‘The human mind was
designed by natural selection operating over
many generations. To understand the influ-
ences of current environments, it is essential
to consider the psychological mechanisms
that are part of that design and are the result
of an interactive blend of nature and nurture’.
Mass media is part of the current environ-
ment that we are seeking to understand, and
to gain full understanding, both nature and
nurture must be considered.
It is simplistic to talk about a model of
nature vs nurture that posits that these are
separate influences. In fact, it is the interac-
tion between genotypes and environments
that explains the ways that humans have
evolved to be a good fit for their various envi-
ronments, differentiated as they are (Scarr
and McCartney, 1983). ‘The dichotomy of
nature and nurture has always been a bad one,
not only for the oft-cited reasons that both are
required for development, but because a false
402 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
parallel arises between the two. We propose
that development is indeed the result of nature
and nurture but that genes drive experience’
(1983: 425). A good example to illustrate how
this might work can be taken from my own
family of origin. Both of my parents were
career music educators and as a result I grew
up in an environment that was heavily influ-
enced by music. At various points, specula-
tion was expressed whether the fact that my
siblings and I were musical was the result of
nature/genes, or nurture/environment. Scarr
and McCartney’s theory would say that my
parents and their genotypes were an influ-
ence on the environment they created in our
household that resulted in a life heavily satu-
rated by musical influences. Thus, it was not
nature plus nurture as if they were two sepa-
rate things, but rather nature and the resulting
environment that was created by my par-
ents who had strong genotypes/phenotypes
for musicality. Scarr and McCartney call
this a ‘passive’ type of environmental effect
by parents who are genetically similar to
offspring.
But what if a child is adopted and grows
up in an environment that is created by non-
biological parents? According to Scarr and
McCartney (1983) there would still be an
‘evocative’ influence whereby the child’s
own genotype affects the ways that she or
he responds to the parents, regardless of
their biological relationship. In this way the
child’s genetic makeup creates a certain
type of environment that is consistent with the
genotype of the child. So if, for example,
the adopted child is naturally introverted
but the adoptive parents are very extroverted,
the child’s responses might be initially
introverted but become progressively more
extroverted as (s)he adapts to this pattern of
interaction.
The third type of influence Scarr and
McCartney (1983) call active niche-picking
or niche-building. In this situation, the child
adopted to parents who are not musical but
who has a genetic proclivity to be musical
will eventually seek out an environment,
once (s)he has a choice, that includes a lot of
music and will build her or his own environ-
ment to support that ability.
The important point in all of these exam-
ples is that nature and nurture as influences
on a developing child cannot be separated
or evaluated in isolation one from the other.
Biology shapes environment and environ-
ment shapes developing traits. It is this
choosing of environments that can have a
direct effect on the eventual choice to repro-
duce or not reproduce.
Said in a more technical way, phenotype
(the visible trait) arises from a combination
of genotype interacting with environment.
To the extent that the genotype of the par-
ents has a direct effect on the environment
of the child, that parent’s genotype is shap-
ing outcomes. The adoptive parent’s geno-
type is not passed to the adopted child, but it
does shape the environment within which the
child grows, which in turn has an effect on
the child’s survival and subsequent ability to
reproduce. The child him or herself responds
to genetic tendencies that cause the choice of
certain environments over others. Thus natu-
ral selection is a process fueled as much by
environment as by biology.
This principle is important because
although the effect of the parents on the
survival of the child is more obvious, it
illustrates why it is important to consider the
effect mass media has on the environment of a
person. This can affect the likelihood that the
person will both survive and reproduce. The
chapter will return to this point at the end,
when a person’s likelihood of developing a
relationship with a media figure and how that
can directly affect that person’s likelihood of
reproducing is discussed.
The conclusion to the question ‘Is human
development or growth the result of nature
or nurture?’ has to be that both must be con-
sidered to get an accurate picture of human
behavior, particularly as shaped by mass
media. This is true in all aspects of develop-
ment including the development of sex and
gender roles.
 EVOLUTIONARY PSYCHOLOGY AND MASS MEDIA
Sex and Gender
In psychology, much of the biology vs sociali-
zation research has dealt with sex and gender,
an area where differences in specific biologi-
cal aspects vs social aspects of development
seem easier to identify (Buss, 2016). Thus, it
is no surprise that much of the research tying
together evolutionary psychology with mass
media looks at the topic through the lens of
gender or sex (Ellis, 1992; Grabe, 2011;
Kamhawi and Grabe, 2006; Malamuth, 1996;
Miller, 2011; Salmon, 2012; Singh and Singh,
2011; Tifferet and Vilnai-Yavetz, 2014).
What do women find attractive in partners
in contrast with what men prefer? The list for
women includes qualities like strength, youth,
ability to provide and protect, and status, and
one conclusion is that although women gener-
ally prefer these qualities in a potential mate,
such preference does not usually occur at a
conscious level. While this does not specifi-
cally address issues related to media, the appli-
cation to how women might process mediated
figures compared to how men process them
can be inferred (Ellis, 1992).
More specific to media is Kamhawi and
Grabe’s (2006) discussion of gender dif-
ferences in the reception of negative news.
Citing research showing that women have a
stronger avoidance of negative stimuli than do
men (e.g. Zald, 2003), they showed that when
news is given a more positive spin, women
are more likely to watch and remember.
Earlier studies had shown that women were
less likely than men to watch and remember
news stories about things like war and disas-
ters. This study suggested that women have
both a socially learned and biological pre-
disposition to avoid negatively framed news
(Kamhawi and Grabe, 2006).
Shoemaker (1996) described the function
of surveillance in early nomadic communi-
ties. All members of the society had to survey
the environment for danger, and some individ-
uals might have had this as their specialized
function in the group. Perceiving potential
threats to the group greatly increased the
403
survival of all members of the group. Grabe
(2011) wrote about the surveillance function
of news, and how our monitoring of news
media is a way to remain apprised of poten-
tially survival-related information. Citing
Shoemaker, she makes the argument that
humans are hard-wired to pay attention to
negative news because of the way journalists
are able to warn the public about potential
dangers. She suggests that the gender differ-
ences similar to those reported in Kamhawi
and Grabe (2006) are a function of histori-
cally early gender roles wherein women pro-
tected children by avoiding danger while men
protected children by monitoring danger and
being prepared to defend against it.
An international randomized study of
Facebook pages and the way users present
themselves found that whereas men were
more likely to use images that conveyed
status and ruggedness, women were more
likely to choose images that related to family
values and/or emotional expression. These
findings used theory-driven hypotheses from
evolutionary psychology to explore aspects
of evolutionary theory as it relates to behav-
ior on social networking sites (Tifferet and
Vilnai-Yavetz, 2014). Another social media
study, this time about Instagram, looked at
the variables that contributed to men’s desire
to date women whose varying photo types
were presented. Neutral photos (defined as
those not showing people) were more appeal-
ing to men who felt lonely and had higher
need for belonging and popularity, whereas
group selfies were more appealing to men
who had lower intrasexual competition for
mates, need to belong, and need to be popu-
lar. This study used a variation on Maslow’s
hierarchy of needs that was modified based
on principles of evolutionary psychology. It
considered affiliation, status, and mate acqui-
sition as higher-order needs. One of the find-
ings is that factors that correlate with desire
for long-term relationships are different from
short-term relationships. Overall, different
photo types were related to differing profiles
of intrasexual competition and the various
404 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
social needs reported. Perceived narcissism
was another key variable, with those post-
ing selfies and groupies perceived as more
narcissistic than those posting neutral photos
(Jin et al., 2019). Both of these social media
studies suggest that sexual selection adapta-
tions have a complex effect on the ways that
users consume social media images.
Some research has suggested that adoles-
cent girls have a pre-occupation with male
celebrities. Girls during this stage of devel-
opment are more likely to form romantic
attachments to celebrities than are boys
during these years (Stever, 1994). Erikson
(1959) suggested that while the formation of
identity in adolescence is ideally before the
development of intimacy which he associ-
ated with young adulthood, often adolescent
girls put intimacy before identity and then
assume the identity of their romantic partner
(e.g. she becomes ‘the doctor’s wife’ or ‘the
minister’s wife’). Emphasis on their histori-
cally evolved role as nurturers over the male
role of provider and protector could explain
why this happens. Engle and Kasser (2005)
specifically queried a population of jun-
ior high school-aged girls and linked such
interest to categories of secure and insecure
attachment, an ethological theory driven by
principles of evolutionary psychology. More
will be said about attachment theory later but,
for now, the observation was that girls who
had positive attachments to same-age boys
that they knew in real life were more likely to
idolize male celebrities. Overall, adolescent
girls are more romantically fixated than are
boys at this age, and differing evolved sex
roles is a possible explanation for this.
Malamuth (1996) looked at gender dif-
ferences in the consumption of sexually
explicit media. He considered research that
attempts to explain differing preferences of
men and women in their consumption of
this kind of material. Vast bodies of work
that advocate for both biological and social
explanations exist, not surprising due to the
controversial nature of this particular subject
(see also Salmon, 2018). When contrasting
erotic literature with romance literature,
romance novels contain pornography written
to appeal to women, while explicitly erotic
images and literature appeal more to men.
These each represent billion-dollar indus-
tries and the differences between them can be
traced to the fact that sexual selection pres-
sures faced by males and females during the
course of evolutionary history have not been
identical (Salmon, 2012). An ethnography
of American romance readers revealed that
women are very much aware of what they
perceived to be a male proclivity for imper-
sonal sex compared to the stories women
in this study preferred, stories that involved
interpersonal connection (Radway, 1984).
A second study of female romance read-
ers found that the basic themes of romance
novels were finding, attracting, and keeping
a desirable mate, certainly themes consist-
ent with evolutionary psychology’s views
of the female mating agenda (Buss, 2016;
Hazen, 1983). Other studies have found sex
differences that involve variations in sexual
fantasies that include visual images, context,
personalization, partner variety and response,
and other variables. It is argued that if there
are real differences between the sexes, then
those differences should be even stronger in
fantasies than in actual behaviors, as fanta-
sies don’t have the same practical constraints
as do behaviors. In general, women have
more contextualized and personalized fan-
tasies and their fantasies are more likely to
involve emotional rather than purely physical
responses. They are more likely to include
personalized and specific partners (Ellis and
Symons, 1990).
These are all examples of how theory in
evolutionary psychology is driving research
on mass media and gender differences.
Malamuth (1996) pointed out that, in gen-
eral, sex differences occur in those domains
for which men faced different problems
from the ones women faced. Adaptation for
each sex then proceeded on a different path
to solve these different problems. Because
the consequences of reproductive behavior
 EVOLUTIONARY PSYCHOLOGY AND MASS MEDIA
for each gender are not the same, men and
women developed different adaptations to
their specific reproductive challenges. One of
the largest differences is the parental involve-
ment required for successful reproduction.
Women must make a much larger minimum
investment than do men. This difference is
the source of many of the differences in mat-
ing strategies between the sexes.
OTHER RESEARCH LINKING
EVOLUTIONARY PSYCHOLOGY
WITH MASS MEDIA
A content analysis of tabloid media using an
a priori list of evolutionary-related topics
showed that sampled tabloid stories were
most often related to survival fitness-relevant
topics, those related to reproductive success
(De Backer and Fisher, 2012). Long-term
relationships, wealth status, health status, and
parental care were four of the most frequently
coded topics in the sample of tabloid media
that was analyzed. The interest in celebrity-
related gossip was linked to the prevalence of
parasocial relationships, discussed below.
Research addressing themes in popular
music linked to evolutionary themes pro-
duced similar findings, with country-western
songs frequently containing phrases relating
to sexuality and reproduction with an average
of 10 such references per song (Hobbs and
Gallup Jr., 2011). Salmon (2018) observed
that these themes are found in music in gen-
eral, although they are found more often in the
most popular songs (e.g. those making Top
10 lists). Hip-hop and rap songs tend to focus
on male interest in short-term mating, with
much bragging about the number of partners
they’ve had. In this same genre, female art-
ists sing about wanting men with status and
resources. The country-western songs had
more themes around parenting as well as the
start-up or break-up of relationships.
Parasocial engagement (Tukachinsky
and Stever, 2019) is defined as the social
405
connection one feels with a personality, usually
mediated, whom one knows in some detail
but without reciprocation. Attachments and
other social relationships meet a fundamen-
tal need that is essential for optimal health
and psychological functioning (Baumeister
and Leary, 1995). Stever (2017a) found that
parasocial attachment to mediated person-
alities is the natural by-product of persis-
tent media consumption of the same face,
voice, and personality of an attractive per-
sona. Attachment theory would predict that
the face and voice are primary mechanisms
of attachment. A human infant will gaze
longer at a human face than any other object
presented to it (Umilta et  al., 1996). He or
she will look longer at a correct human face
than an incorrect one (Easterbrook et  al.,
1999), and will prefer the face and voice of
the mother to other faces and voices (Blehar
et al., 1977; DeCasper and Fifer, 1980; Muir
et  al., 1994). Adults prefer attractive and
symmetrical faces, and one theory is that
such attractiveness is an indicator of good
health and a strong genetic background (Fink
and Penton-Voak, 2002). Reflecting back to
the argument that humans respond to media
as if it were real (Grabe, 2011; Reeves and
Nass, 1996), humans respond to attractive
potential mates whether they are 10 feet away
or on a television screen. The familiar other
is a potential mate. In some cases this is not
adaptive in that sometimes the person who
is fooled into thinking that attractive other is
attainable persists in her or his attraction and
refuses to give it up. This is discussed in more
detail below.
Media use compensates for diminished
social interaction in situations wherein people
experience isolation (Jonason et  al., 2008).
Because humans have evolved to be social
beings, isolation creates psychological dis-
comfort. Various kinds of ‘social snacking’
were investigated, operationalized for this
study as ‘talking to oneself’ and ‘watching
TV’. Whereas both genders used watching
TV to mitigate social isolation, women indi-
cated greater amounts of self-talk compared
406 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
to men. Additionally, the researchers point
out that parasocial interaction is another way
people cope with loneliness (Cohen, 2001;
Giles, 2002).
Bandura (2001), in his work on role mod-
els and vicarious social learning, maintained
that media role models can be as powerful as
proximal role models in shaping the behav-
ior of those who admire and emulate them.
Already discussed has been the role that
learning adaptive behaviors from previous
generations is a fundamental aspect of how
media might affect adaptation. Taking this
together with parasocial theory, it is clear that
both identification and attachment are fac-
tors influencing the role that media plays in
the relationships people form with mediated
others, relationships that can have adaptive
consequences.
Evolutionary Theory and
Attachment
Attachment theory, at its foundation, assumes
that attachment behavior has evolved to be an
adaptive behavioral pattern designed or
selected to assure survival of the infant. A
repertoire of attachment behaviors and a cor-
responding set of caregiving behaviors on the
part of the adult caregiver complement each
other and work together towards the success-
ful development of the infant. Although
sexual selection and successful mating are
important, all of that is for nothing if the
infant does not survive (Keller, 2013).
Hazan and Shaver (1994) and others have
posited that the attachment system that favors
the survival of the infant is also a part of the
adult romantic relationship. Partners look
to one another for mutual support and care­
giving in a paradigm that defines attach-
ment as the seeking of feelings of security
and safety. Consistent with the idea that a
key aspect of evolutionary psychology is
the heritable quality of emotional responses,
attachment theory says that inborn behaviors
elicit feelings and emotions that predispose
one to feel connections with familiar others.
Extending this to feelings that viewers develop
for media personae, just as thoughts of the
real-life romantic partner can make one
feel safe and secure even when that partner
is not physically present, mediated perso-
nae are able to elicit feelings of safety and
security in the viewers who have those para-
social attachments (Stever, 2013, 2017a).
Psychological comfort and a sense of safety
potentially could play a role in the ability of
humans to adapt to social environments but
more research is needed to solidify this link.
Parasocial attachment as a part of classical
attachment theory is a new concept that needs
further exploration. A recent study exploring
the ways that viewers view media celebri-
ties as parental attachment figures, looking
to such a celebrity to meet a specific attach-
ment need, is a good example of this kind of
research (David et al., 2019).
Resulting Theories
Out of work in media studies have come a
number of theories including social presence,
media richness, and media naturalness (Kock,
2012). Presence is defined as ‘a psychological
state in which the virtuality of experience is
unnoticed’ (Lee, 2004: 494). Thus, interac-
tion is carried on with the mediated nature of
that behavior being in the background and not
noticed on a conscious level. If you ask the
participant if this behavior is mediated, they
are able to state that it is, but otherwise they
behave as if the interaction is as real as face-
to-face interaction. Lee concludes that ‘pres-
ence research will be equally benefitted by
seriously considering the implication of evo-
lutionary psychology for the explanation of
how people use and interact with media and
simulation technologies’ (Lee, 2004: 501).
Social presence and media richness theo-
ries (Kock, 2004, 2012) each posit that face-
to-face communication was most affected
by evolution. Because humans have evolved
to be good at face-to-face communication,
 EVOLUTIONARY PSYCHOLOGY AND MASS MEDIA
media that best approximates face-to-face
communication is the most engaging. Daft
and Lengel (1986) argued that richness was
dependent, in part, on the medium’s ability
to convey nonverbal information as a part of
the communication. Written communication
has no nonverbal cues whereas traditional
face-to-face communications have rich cues.
Between those two extremes are a continuum
of various forms of mediated communication
that might have more or fewer nonverbal cues
available as a characteristic of that form. For
example, communicating via FaceTime on a
cell phone includes many nonverbal cues but
not as many as being in the presence of the
person, but quite a few more than just writ-
ing an e-mail. An earlier idea, social presence,
conveyed a similar distinction between face-
to-face vs written communications (Short
et al., 1976). Other aspects of social presence
or media richness include the availability of
immediate feedback from the communica-
tion partner, as well as the ability to convey
aspects of one’s personality (Kock, 2004).
As further research on these ideas was con-
ducted, there was evidence both for and
against the media richness theory, suggesting
the need for an alternative explanation, and
resulting in the development of media natu-
ralness theory (Kock, 2012).
Media naturalness theory contends that
reduction of cognitive effort is the more
important factor in explaining the greater
effectiveness of media forms that best
approximate face-to-face communication.
Because humans evolved to communicate in
person and in a manner that is most often syn-
chronous, to the extent that these aspects are
employed, communication takes less cogni-
tive effort. Kock (2012) identifies five aspects
of communication that are involved in media
naturalness. The first is the individuals’ abil-
ity to see and hear each other; the second
is that communication be synchronous; the
third is the inclusion of facial expressions
as a part of the message. Body language is
the fourth element. Speaking directly to one
another is the fifth element, such that partners
407
can talk to and hear one another. Specialized
areas of the brain have evolved that allow
facial recognition, the recognition of emo-
tional cues, and the processing of speech,
and thus it makes sense that communicating
in this way will allow easy processing by all
parties involved.
When evaluating computer-mediated com-
munication, media richness theory can favor
a mode that features synchronicity and feed-
back immediacy whereas media naturalness
theory favors the form that requires less cog-
nitive effort, most often deemed to be face-
to-face. In many examples, these two theories
seem to come to similar conclusions, but
the important consideration is the different
emphases in each (Kock, 2004, 2012).
The Pleistocene Brain
It is central to the arguments of this chapter
that humans expend a considerable amount of
time, effort, and resources in the production
and consumption of mass media. Additionally,
the ability of such mediated forms to elicit
deep emotion from the consumer is something
that social scientists seek to explain using
established epistemologies (Hennighausen
and Schwab, 2015). What is the origin of
evolved psychological mechanisms and what
purpose do they serve? These processes
evolved in response to adaptive problems
that occurred in the Pleistocene (or Ice Age,
ending 11,700 years ago) human past
(Barkow et  al., 1992). They evolved very
slowly over many millennia in contrast to the
rapid social change that one can observe
beginning from that time to the present.
Predominantly, this is an era also referred to
as hunter/gatherer society. Experts appear to
agree that the Pleistocene brain or ‘old brain’
is still the one that guides human behavior
(Grabe, 2011). Have evolved psychological
mechanisms that evolved during the
Pleistocene to solve adaptive problems
served to determine how humans use media?
This is another important question.
408 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
GRABE VS REEVES AND NASS: A
DEBATE ABOUT THE ‘OLD’ BRAIN:
ADAPTED OR OUTMODED?
Related to all of this is Reeves and Nass’
(1996) theory, referred to as The Media
Equation. Simply stated, their work suggests
that humans interact socially with media in the
same ways that they interact in face-to-face
social situations: ‘… our research shows that
media are perceived as real people and places,
and that human responses to media are deter-
mined by the rules that apply to social rela-
tionships and navigating the world’ (1996:
10). When we see people or events depicted
through media, reactions are governed by the
rules of social interaction. For example, par-
ticipants in research treated a computer as an
active social partner, showing that computer
courtesy and good manners. During experi-
ments, people used polite behavior when
interacting with a machine when that machine
asked for evaluations. We will tell the machine
we are using that it is helpful, accurate, and
friendly, but when asked to evaluate that same
computer from a different computer, responses
were far less polite, suggesting that partici-
pants were responding to the computer in a
way similar to any other social partner.
Additionally, interpersonal distance as depicted
in pictures elicits similar responses to those
figures as does interaction with real people.
When people in pictures seem close, our reac-
tions are more intense, and we pay more atten-
tion and remember those people better (Reeves
and Nass, 1996).
By responding to media in the same way
we react to face-to-face others, does this
reflect that our Pleistocene brain has failed to
adapt to media and is reacting in a way that
is outmoded in an evolutionary sense? Grabe
(2011), in interpreting The Media Equation,
suggests that rather than look at this phenom-
enon as an outmoded ‘old’ brain that has not
kept up with change, we rather consider that
processing media as if it were the same as ‘real
life’ is an adaptive way to integrate mediated
information into one’s day-to-day experience.
Is the processing of media as ‘real’ always
adaptive? This question is addressed later in
this chapter.
An important counterpoint to the research
on The Media Equation is work done by
Ramos and colleagues (Ramos et  al., 2013).
Specifically, they found that for Hispanic
viewers who were the participants in their
study, more empathy for objects of violence in
a film was reported when they knew the vio-
lence was real compared to when it was part
of fiction. Previous exposure to violence did
not affect the degree of empathy shown with
respect to these two types of violence, media
depicting real victims vs media depicting fic-
tional victims. A victim empathy scale was
used to measure the dependent variable in this
study, a self-report measure. Actual physiolog-
ical responses were not recorded in this case.
Entertainment as Central to
Mass Media
When considering the role of entertainment
as a result of evolution, two approaches are
recognized. The first is that entertainment is
a by-product of evolution. The second is that
entertainment is itself an evolutionary adap-
tation to conditions in the environment
(Hennighausen and Schwab, 2015). Other
academics have also written about a connec-
tion between evolutionary psychology and
entertainment (Vorderer et  al., 2006) and
have considered whether humans develop an
interest in being entertained as an adaptation,
or as a by-product of adaptations. One view
that it is a by-product of adaptation posits
that as humans became more adept at sur-
vival and no longer needed to expend all of
their time and energy on it, this created lei-
sure time, and entertainment originated from
the desire to fill such leisure time with pleas-
urable activity. The Media Equation approach
(Reeves and Nass, 1996) also assumes that
human responses to media are a by-product
of other adaptations. We respond to media as
 EVOLUTIONARY PSYCHOLOGY AND MASS MEDIA
if it were real because we have evolved social
responses to real people, and media resem-
bles real people enough that we respond to it
in a similar fashion.
However, if entertainment is an adaptation
in itself, then humans might have developed
skills in being interesting and engaging in
order to attract potential mates. In this case,
entertainment is part of the primary task of
reproduction and sexual selection. Miller
(2011) developed the idea that the human
mind, with its creativity and complexity, serves
to attract mates in and of itself and as such is
an evolutionary adaptation. Darwin (1871)
called this a ‘hidden courtship function’. Said
more simply, having a creative and interesting
mind is ‘sexy’ to the prospective mate (see
Hennighausen and Schwab, 2015 for a more
extensive discussion of these approaches).
Memory, anticipation, decision-making, and
pleasure are all key aspects of sexual choice
(Miller, 2011). These are all factors that easily
can play into media choices as well and sexual
attraction is an important factor when choos-
ing one’s favorite celebrity (Stever, 1991). It
is possible that both factors, entertainment and
media as a by-product of adaptation, and as an
actual adaptation, might be at work, and that
it does not have to be one or the other (Ohler
and Nieding, 2006). Entertainment can be a
by-product of adaptations that led to increased
leisure time, but also could be a mechanism
for enhanced sexual selection.
Emotion, Play, and Fantasy
This discussion of human evolution and how
media fits into the picture begins with the
observation that a huge aspect of media effects
is the way media stimulates human emotions.
It has been argued that our emotional responses
to media are as intense as those same reactions
to real events, as already noted (also Schwender
and Schwab, 2010). Some theorists have sug-
gested that the role of media in human life is
largely dysfunctional in that media distracts
us from dealing with real day-to-day things
409
in life. In addition, at first look, entertain-
ment and mediated experience do not appear
to have any real function in human life.
However, upon closer analysis, two key
functions of human existence, sensory percep-
tion and communication, seem to be related
in a positive way to mediated experience. In
an evolutionary sense, problem solving has an
adaptive function and consuming media nar-
ratives, whether in books, films, and televi-
sion, or even on the Internet, offers a form of
cognitive rehearsal for real-life events.
Nielsen (2012) identified the key element
in this process as ‘imitation’. It is the pre-
dominant way that culture is transmitted from
one individual to another and from parent to
offspring. However, a key aspect of human
development was that successive generations
improved the tools they were given by par-
ents. Such innovation explains, for example,
the rapid progression of modes of air travel
once a basic plane had been developed. Each
generation didn’t have to start at the begin-
ning of the ‘can humans fly’ question but
rather could build on the accomplishments of
their immediate predecessors. A prolonged
stage of growth and development for humans
gives an opportunity for advanced cogni-
tive skills to develop. Thus ‘childhood’ is a
unique feature of humans, and affords the
time necessary to learn and develop. Play is
identified as a primary mechanism in imita-
tion and learning during childhood.
Thus, a central part of development for
children is play (Erikson, 1974) whereby
children often rehearse tasks that they will
be called upon to perform as adults. Media
entertainment is an extension of the concept
of ‘play’ and a higher-level form of cognitive
rehearsal for tasks and situations that might
come into play throughout life in various
relationships, so that we can practice inter-
personal, emotional, or social situations with-
out incurring any unnecessary risk to real
relationships. This can be argued to have a
possible real and positive influence on repro-
ductive outcomes for those who learn about
social situations in this way.
410 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
It is recognized that play is a fundamental
aspect of human behavior, and that play is
often the rehearsal of behaviors that are adap-
tive for either survival or reproduction. In
play we ‘practice’ doing things that will later
ensure that we can be successful in life. What
is play? It is the creation of a fantasy envi-
ronment within which such practice can take
place. So, a little girl in her pretend kitchen
‘cooks’ dinner for her family and serves it to
them on small facsimiles of dishes. From such
play, she later is able to learn real cooking
skills. A little boy pretends to care for his doll,
and such play is practice for future parenting.
Theorists have suggested that mass media
is experienced as ‘play’ by its recipients.
Stephenson (1988: 49) recounts that the daily
mix of media consumed by viewers ‘is repeti-
tious, like a child’s game played over and over
with variations on a familiar theme’. It is not
a big leap to see the consumption of fiction
in mass media as a springboard for fantasy,
which is related to play. A viewer watches a
favorite television show and when the show
is over, fantasizes about being in social situ-
ations with a favorite protagonist, scripting
conversations and practicing how to behave
in those social situations. This social behav-
ior is practice for future real encounters in the
same way that the child cooking in a pretend
kitchen is learning behaviors that will apply
to real preparing of meals. The fantasy inter-
action with a favorite media persona is a type
of parasocial relationship.
Thus, play involves fantasy and when one
consumes media, one is transported into a
parasocial fantasy world where we interact
with the inhabitants as if they were real. We
assess possible fantasy mates in these worlds
in the same way we might assess real poten-
tial mates in a face-to-face environment.
The Media Personae as an
Idealized Romantic Figure
Fisher (2004) has done extensive research on
the nature of romantic love and how and why
humans fall in love. She has concluded that
the brain evolved to execute three brain func-
tions related to love. These are lust, romantic
love, and attachment. Attachment has already
been discussed here and sexual attraction has
been discussed as well. Romantic love has
been reported across time and across cultures
in terms that are close to universal. Various
terms are used to refer to the feelings associ-
ated with romantic love including infatua-
tion, obsession, and passion. Fisher’s studies
showed that the experiences and feelings
associated with this state are similar across
cultures, ages, and genders.
Fisher recounts, ‘Lovers dote on the posi-
tive qualities of their sweethearts, flagrantly
disregarding reality’ (Fisher, 2004: 8). What
she refers to here is the tendency to idealize
the beloved and ignore negative qualities or
turn those negative qualities into endearing
traits. It is not until the relationship comes
up against reality that the rose-tinted glasses
come off and the partner is seen as she or he
really is. Over the course of time, idealization
is tempered. This is potentially the founda-
tion for why when viewers form ‘crushes’
on media figures, those feelings persist and
the object is idealized. Because there is never
any dose of reality to bring those ideals into
a more realistic perspective, the viewing of
the media object as near to perfection is a
common experience. Stever (1991) found
that when asked how big a fan someone was
of their favorite celebrity, the best predictor
for that Likert scale was romantic attrac-
tion. Clearly, these feelings of infatuation are
heavily tied to why viewers become roman-
tically attracted over time to their favorite
media personalities.
IMPLICATIONS FOR THE IMPACT OF
MEDIA ON INDIVIDUALS
The argument is that humans have not
evolved beyond the hunter/gatherer society
with respect to social behavior. In order to
 EVOLUTIONARY PSYCHOLOGY AND MASS MEDIA
understand the importance of the premise of
The Media Equation, that people respond to
media as if it were real, and to understand the
impact that mass media is having on social
life, it is important to consider the ways in
which courtships have traditionally happened
throughout recorded human history. It has
been argued that the human brain has acted
on the basis of one model of human interac-
tion, and that was the model established by
the Pleistocene hunter/gatherer society, a
society that was conducted face-to-face.
In the Pleistocene era, if a woman saw a
man who was a good potential mate, she had
only to behave in ways that would be attrac-
tive to him, and she had every expectation of
being able to bear his offspring. The ways
potential mates were evaluated has already
been discussed, but status, age, and resources
were all factors in this evaluation. Because a
woman’s reproductive investment was in one
man at a time, it was essential to her repro-
ductive success that she attract the best candi-
date with whom to mate. Throughout history
and up until very recently, the only candi-
dates for potential mates were men that the
woman met in person and knew face-to-face.
If she could get close enough to evaluate him
for things like strength, health, and attractive-
ness, she would be able to attract him and
bear his children. It was not even essential
that she be his only partner.
Fast forward to the 20th century, where
film, television, and eventually the Internet
made the images and information available
about potential candidates for mates that the
woman had not met in person, indeed might
likely never meet in person. The ‘script’ for
mating was still written in the behavioral
social system of the woman’s brain. In the
earliest stages of attraction, initiation, and
experimentation (Tukachinsky and Stever,
2018), it is easy for a woman to fantasize
about a potential mate, trying out various
mating scenarios in her imagination, in order
to evaluate the worthiness of a potential mate.
Throughout history, this was something
that women had been doing with respect to
411
in-person candidates for mating. Now women
can evaluate distant potential partners and, if
the potential mate is found worthy, she could
attempt to pursue him.
The problem comes when it occurs to this
woman that she does not know how to meet
the potential mate, and in some cases, she
realizes that she is one of thousands who are
attracted to this same person. She is still in
the phase of romantic attraction where she
has idealized this possible mate, and so her
pursuit of him becomes a desire to meet him
face-to-face in order to realize the fantasy in
real time and in real life. It is not until she
reaches this point in the ‘courtship’ that
the problems with mediated reality become
apparent.
In my interviews with women who have
been through this process with an attractive
distant media figure, there is one of three out-
comes: 1) She has a reality check, gives up
her pursuit of this idealized mate, and seeks
instead to find someone in proximity; 2) She
refuses to give up her pursuit of the ideal-
ized other and proceeds to pursue him by
any means available. This choice often ends
in despair as the pursuit is not successful; or
3) If she is vulnerable to mental illness, she
enters a delusional state (referred to as ero-
tomanic schizophrenia), where she evaluates
the fantasy relationship as ‘real’ and behaves
as if it is. In most cases of erotomania, the
mentally ill person lives in isolation in her
fantasy world, firm in her belief that she has
been separated from her beloved and that
if they could be reunited, all would be well
(Kennedy et  al., 2002). In a few cases, the
erotomanic becomes a stalker and pursues
the person she believes is her partner via any
means available. Both history and the news
have reported various stories about this type
of woman. David Letterman’s stalker (Bruni,
1998) went so far as to enter his home, drive
his car, and tell others that they were mar-
ried. (Eventually she took her own life, a
frequent outcome for persons in this situa-
tion.) Other celebrities have had to deal with
similar intrusions into their private lives by
412 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
people who believed that the celebrity was
his or her romantic partner. Some research
has suggested that erotomania actually ‘may
be viewed as a pathological variant of a spe-
cific sexual strategy that evolved under selec-
tion pressures of the human environment
of evolutionary adaptedness’ (Brüne and
Schröder, 2003: 83). This suggests that this
kind of obsession with a media figure pos-
sibly reflects sexual selection pressures gone
wrong.
One of the main arguments of The Media
Equation and discussions of this theory has
been that if people perceive what they see
in media as real, and react to it as if it were
real, what harm is done? Grabe (2011: 366)
asks, ‘what existing or anticipated environ-
mental need might compel human adapta-
tion for the ability to separate media from
physical reality….Why should our brains
discriminate between media and physical
worlds if they are converging’? However,
in the example above, a viewer’s belief
that the attractive potential mate she ‘meets’
through media is obtainable and that it
could or should be real has the potential to
wreak havoc on that viewer’s life for many
years.
In my years of participation in fan com-
munities, I have met and/or seen numerous
fans who were in deep despair because they
realized they were in love with someone that
they could never have or even meet. Although
this represents a small percentage of fandom,
it is a real situation that I have encountered
numerous times in the 30-plus years I have
been involved in participant observation in
media fandoms.
So far, I have discussed this scenario with
respect to women pursuing men and, indeed,
most of the real-world cases I have observed
have been women in this situation (the fan-
doms I have studied often were dominated
by women). Looking at what we know about
gender differences in mate selection, it is not
hard to understand why a woman looking to
partner with a famous male celebrity would
be more common than a man seeking to mate
with a famous female celebrity. To succeed,
a woman only has to be one of potentially
many possible partners. For a man to suc-
ceed, he has to become the ONLY partner
for that female celebrity, at least for a time.
Thus, traditionally the throngs of adoring
fans who want to partner with the famous
celebrity have been more likely to be women.
However, men are vulnerable to this situation
as well.
An Illustration from The Big Bang
Theory
The idea that a fan would replace the oppor-
tunity for a real relationship in the face-to-
face world with a fantasy attachment to a
media character is illustrated well in an epi-
sode of the television sitcom The Big Bang
Theory. Howard Wolowitz has been intro-
duced to Bernadette and Bernadette wants an
intimate relationship with him. Howard hesi-
tates to commit to the real relationship. Why?
As he explains to his friends (in season 3,
episode 9), ‘She wants a commitment and
I’m not sure she’s my type. Bernadette is
really nice. I just thought that when I was
ready to settle down in a relationship, it
would be with someone different. More like
Megan Fox from Transformers or Katee
Sackhoff from Battlestar Galactica’. When
Penny replies, ‘You’re going to throw away a
great girl like Bernadette because you’re
holding out for some ridiculous fantasy?’,
Howard’s answer is, ‘Just because you set-
tled doesn’t mean I have to. I want what’s
inside too but why is it wrong to want those
insides wrapped up in the delicious caramel
that is Halle Berry?’ Sheldon’s observation at
this point is, ‘Biologically speaking, Howard
is perfectly justified in seeking the optimum
mate for the propagation of his genetic line’.
Later in the episode, fantasy Katee Sackhoff
asks Howard, ‘Why are you playing make
believe with me when you could be out with
a real woman tonight?’ When Howard says,
‘She’s not you’, Sackoff replies, ‘I’m not
 EVOLUTIONARY PSYCHOLOGY AND MASS MEDIA
me…the point is you’ve got a real girl in
your life and you’re ignoring her to spend
your time with a mental image…’.
In 30 years of fan research, I have met
many dozens of fans who are holding out
for the ideal represented by their favorite
celebrity and who would not consider some-
one lesser than that ideal. In many cases, it
meant that the fan did not date at all. From
a natural-selection perspective, people who
idealize and stay single/celibate as a result
do not have their genes passed on to progeny.
This is particularly true for women who are
less likely to engage in frivolous reproductive
activity if they are holding out for a perfect
mate. Men can both hold out and also have
casual sex because of their lesser investment
in potential offspring. The Howard Wolowitz
character is a good example of this as well.
However, whether the celebrity ideal was a
real person or a fictional character, the fan who
is unwilling to settle for less than the ideal is
far less likely to reproduce than the one who is
more realistic.
However, if media ‘fools’ the brain into
thinking that a real relationship is being
experienced (real in the sense of repro-
ductive potential), this is exactly the kind
of thing that Restak (1991) has described.
There is a part of the brain that processes
what is seen as ‘real’, even if the perception
is mediated.
Cultural anthropology teaches us about
status hierarchies within cultures and the desire
of individuals to mate with higher-status
individuals than are currently available in
­ are part of evolutionary theoretical models,
one’s pool of potential mates (Wright, 1994).
It makes sense that the attractive celebrities
found in television, films, and other avenues
of popular culture would be particularly
attractive to the viewer who sees an individual
who is likely to be more attractive, wealthier,
and of higher status than potential mates who
are in closer proximity. The virtual proxim-
ity of such individuals tricks the viewer into
thinking that high-status celebrity mates are
attainable. As Restak (1991) pointed out, The
Brain Has a Mind of Its Own.
413
CONCLUSION
To summarize the main points from this chapter,
first, mass media has not been a factor in
human evolution long enough to have shaped
inherited characteristics in the species.
Research supports the idea that the hunter/
gatherer Pleistocene-era brain is still the brain
that regulates human social life (Barkow et al.,
1992). Next, evolutionary psychologists have
spent a good deal of time investigating the
effects of sex and gender on media selection
and processing. Different reproductive agen-
das and roles in the social group affect the
ways that men and women process and react
to media messages (Buss, 2016). The work of
Sherry (2004, 2015) has suggested that
research into mass media has integrated both
biological and sociocultural influences on
mass media selection and consumption into
various theoretical models. These include
social presence, media richness, and media
naturalness theories (Kock, 2012). The Media
Equation (Reeves and Nass, 1996) suggested
that humans respond to media as if it were
real, showing little difference in the way media
messages are processed compared to in-­person
messages. Face-to-face communication
appears to require less cognitive effort and
thus mediated forms of communication that
most closely resemble face-to-face communi-
cation require less cognitive effort and are
more efficient and effective. Discussed were
the ways that entertainment, fantasy, and play
playing the function of rehearsal of future life
tasks, both social and occupational.
The final major point is that the inability
to process mediated social situations differ-
ently from face-to-face social situations may
occasionally have maladaptive or undesirable
outcomes, particularly if mating and reproduc-
tion are the desired outcomes. However, social
situations available through media have the
potential to mitigate loneliness and support the
individual who is relatively isolated. The litera-
ture on parasocial relationships and parasocial
414 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
attachments (Stever, 2017b), including the
literature on celebrity worship (McCutcheon
et  al., 2002), suggests that there are both
negative and positive potential outcomes for
meeting one’s social needs through mediated
parasocial relationships.
Research investigating the effects of mass
media from an evolutionary psychology per-
spective is relatively new and a great deal
more work is needed to investigate, in par-
ticular, the long-term effects of increasing
levels of media consumption. This includes
the growing rate at which people are meeting
social needs through social media, as well as
texting, tweets, and various other messaging
formats. The increasing presence of media
devices in day-to-day lives, devices like cell
phones, iPads, laptops, personal computers,
and other devices that are used on a daily basis
in more and more situations, also needs to be
explored to see how the human brain as it has
evolved is suited to and fits with persistent use
of these devices. Comparisons between heavy
media users and those who refrain from using
media would be instructive in comparing brain
development that results from use or nonuse.
REFERENCES
Bandura, A. (2001). Social cognitive theory of
mass communication. Media Psychology, 3,
265–299.
Barkow, J. H., Cosmides, L., & Tooby, J. (Eds.).
(1992). The adapted mind: Evolutionary psy-
chology and the generation of culture. New
York: Oxford University Press.
Baumeister, R. F., & Leary, M. R. (1995). The need
to belong: Desire for interpersonal attach-
ments as a fundamental human motivation.
Psychological Bulletin, 117(3), 497.
Blehar, M., Lieberman, A., & Ainsworth, M.
(1977). Early face-to-face interaction and its
relation to later infant-mother attachment.
Child Development, 48(1), 182–194.
Brüne, M., & Schröder, S. G. (2003). Erotomania
variants in dementia. Journal of Geriatric
Psychiatry and Neurology, 16(4), 232–234.
Bruni, F. (November 22, 1998). ‘Behind the
Jokes, a Life Of Pain and Delusion; For Letter-
man Stalker, Mental Illness Was Family Curse
and Scarring Legacy’. The New York Times.
Retrieved May 6, 2013. https://www.
nytimes.com/1998/11/22/nyregion/behind-
jokes-life-pain-delusion-for-letterman-
stalker-mental-illness-was-family.html
Buss, D. M. (2016). The evolution of desire.
New York: Springer International Publishing.
Cohen, J. (2001). Defining identification: A
theoretical look at the identification of audi-
ences with media characters. Mass Commu-
nication & Society, 4(3), 245–264.
Daft, R. L., & Lengel, R. H. (1986). Organizational
information requirements, media richness and
structural design. Management Science,
32(5), 554–571.
Darwin, C. (1871). The descent of man.
London: Gibson Square.
David, K., Myers, M. E., Perry, S. D., Gouse, V., &
Stein, C. B. (2019). Examination of insecure
attachment and the potential for parasocial
parental attachment (PPA) to a favorite celeb-
rity through attachment theory. North Ameri-
can Journal of Psychology, 21(2), 387–406.
De Backer, C. J., & Fisher, M. L. (2012). Tabloids
as windows into our interpersonal relation-
ships: A content analysis of mass media
gossip from an evolutionary perspective. Jour-
nal of Social, Evolutionary, and Cultural Psy-
chology, 6(3), 404–424.
DeCasper, A., & Fifer, W. (1980). Of human
bonding: Newborns prefer their mothers’
voices. Science, 208, 1174–1176.
Deuze, M. (2011). Media life. Media, Culture &
Society, 33(1), 137–148.
Easterbrook, M.A., Kisilvesky, B.S., Muir, D.W., &
LaPlante, D.P. (1999). Newborns discriminate
schematic faces from scrambled faces,
Canadian Journal of Experimental Psychology,
55(3), 231–241.
Ellis, B. J., & Symons, D. (1990). Sex differences
in sexual fantasy: An evolutionary psycho-
logical approach. Journal of Sex Research,
27(4), 527–555.
Ellis, B. J. (1992). The evolution of sexual
attraction: Evaluative mechanisms in women.
In J. H. Barkow, L. Cosmides, & J. Tooby, (Eds.),
The adapted mind: Evolutionary psychology
and the generation of culture, (267–288).
New York: Oxford University Press.
 EVOLUTIONARY PSYCHOLOGY AND MASS MEDIA
Engle, Y., & Kasser, T. (2005). Why do adolescent
girls idolize male celebrities? Journal of Ado-
lescent Research, 20(2), 263–283.
Erikson, E. (1959). Identity and the life-cycle.
New York: W.W. Norton & Co.
Erikson, E. (1974). Play and actuality. In R. J.
Lifton & E. Olson (Eds.), Explorations in psy-
chohistory. The Wellfleet Papers (109–135).
New York: Simon and Schuster.
Falk, E. B., Cascio, C. N., & Coronel, J. C. (2015).
Neural prediction of communication-relevant
outcomes. Communication Methods and
Measures, 9(1–2), 30–54.
Fink, B., & Penton-Voak, I. (2002). Evolutionary
psychology of facial attractiveness. Current
Directions in Psychological Science, 11(5),
154–158.
Fisher, H. (2004). Why we love: The nature and
chemistry of romantic love. New York:
Macmillan.
Geher, G. (2014). Evolutionary psychology 101.
New York: Springer.
Giles, D. C. (2002). Parasocial interaction: A
review of the literature and a model for future
research. Media Psychology, 4(3), 279–305.
Grabe, M. E. (2011). News as reality-inducing,
survival-relevant, and gender-specific stimuli.
In S. C. Roberts (Ed.), Applied Evolutionary
Psychology, (361–377). New York: Oxford
University Press.
Hazen, H. (1983). Endless rapture: Rape,
romance and the female imagination. New
York: Scribner’s.
Hazan, C., & Shaver, P. (1994). Attachment as an
organizational framework for research on close
relationships. Psychological Inquiry, 5, 1–22.
Hobbs, D. R., & Gallup Jr., G. G. (2011). Songs as
a medium for embedded reproductive mes-
sages. Evolutionary Psychology, 9(3), 390–
416. doi:10.1177/147470491100900309.
Jin, S. V., Ryu, E., & Muqaddam, A. (2019).
Romance 2.0 on Instagram! ‘What type of
girlfriend would you date?’ Evolutionary Psy-
chology, 17(1), 1474704919826845.
Jonason, P., Webster, G., & Lindsey, A. (2008).
Solutions to the problem of diminished social
interaction. Evolutionary Psychology, 6(4),
637–661.
Kamhawi, R., & Grabe, M. E. (2006). Gender
differences in negative news reception: An
evolutionary psychology explanation. Media
Report to Women, 34(4), 15.
415
Keller, H. (2013). Attachment and culture. Journal
of Cross-Cultural Psychology, 44(2), 175–194.
Kennedy, N., McDonough, M., Kelly, B., &
Berrios, G. E. (2002). Erotomania revisited:
Clinical course and treatment. Comprehensive
Psychiatry, 43(1), 1–6.
Kock, N. (2004). The psychobiological model:
Towards a new theory of computer-mediated
communication based on Darwinian evolution.
Organization Science, 15(3), 327–348.
Kock, N. (2012). Media naturalness theory:
Human evolution and behaviour towards elec-
tronic communication technologies. In S. C.
Roberts (Ed.), Applied Evolutionary Psychology,
(381–398). New York: Oxford University Press.
Lee, K. M. (2004). Why presence occurs: Evolu-
tionary psychology, media equation, and
presence. Presence: Teleoperators & Virtual
Environments, 13(4), 494–505.
Malamuth, N. M. (1996). Sexually explicit media,
gender differences, and evolutionary theory.
Journal of Communication, 46(3), 8–31.
McCutcheon, L. E., Lange, R., & Houran, J.
(2002). Conceptualization and measurement
of celebrity worship. British Journal of Psy-
chology, 93(1), 67–87.
Mead, G. (1934). Mind, self, and society.
Chicago, IL: The University of Chicago Press.
Miller, G. (2011). The mating mind: How sexual
choice shaped the evolution of human
nature. New York: Anchor Books.
Muir, D. W., Humphrey, E. E., & Humphrey, G. K.
(1994). Pattern and space perception in young
infants. Spatial Vision, 8(1), 141–165.
Nielsen, M. (2012). Imitation, pretend play, and
childhood: Essential elements in the evolution
of human culture? Journal of Comparative
Psychology, 126(2), 170–181.
Ohler, P., & Nieding, G. (2006). An evolutionary
perspective on entertainment. In J. Bryan &
P. Vorderer (Eds.), Psychology of Entertain-
ment, (423–434). New York: Routledge.
Radway, J. (1984). Reading the romance: Women,
patriarchy, and popular literature. Chapel Hill,
NC: University of North Carolina Press.
Ramos, R. A., Ferguson, C. J., Frailing, K., &
Romero-Ramirez, M. (2013). Comfortably
numb or just yet another movie? Media vio-
lence exposure does not reduce viewer empa-
thy for victims of real violence among primarily
Hispanic viewers. Psychology of Popular Media
Culture, 2(1), 2–10.
416 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Reeves, B., & Nass, C. I. (1996). The media equa-
tion: How people treat computers, television,
and new media like real people and places.
London: Cambridge University Press.
Restak, R. (1991). The brain has a mind of its
own: Insights from a practicing neurologist.
New York: Crown.
Salmon, C. (2012). The pop culture of sex: An
evolutionary window on the worlds of por-
nography and romance. Review of General
Psychology, 16(2), 152–160.
Salmon, C. (2018). Evolutionary perspectives on
popular culture: State of the art. Evolutionary
Studies in Imaginative Culture, 2(2), 47–66.
Scarr, S., & McCartney, K. (1983). How people
make their own environments: A theory of
genotype environment effects. Child Devel-
opment, 54(2), 424–435.
Schramm, W. (Ed.). (1960). Mass communica-
tion. (2nd ed.). Urbana, IL: University of Illi-
nois Press.
Schramm, W. (1988). The story of human com-
munication: Cave painting to microchip.
New York: Harper and Row.
Schwender, C., & Schwab, F. (2010). The
descent of emotions in media: Darwinian
perspectives. In K. Doveling, C. von Scheve &
E. A. Konijn (Eds.), The Routledge handbook
of emotions and mass media, (29–50). New
York: Routledge.
Seiffert-Brockmann, J. (2018). Evolutionary
psychology: A framework for strategic com-
munication research. International Journal of
Strategic Communication, 12(4), 417–432.
Sherry, J. L. (2004). Media effects theory and
the nature/nurture debate: A historical over-
view and directions for future research.
Media Psychology, 6(1), 83–109.
Sherry, J. L. (2015). The complexity paradigm
for studying human communication: A
summary and integration of two fields.
Review of Communication Research, 3,
22–54.
Shoemaker, P. J. (1996). Hardwired for news:
Using biological and cultural evolution to
explain the surveillance function. Journal of
Communication, 46, 32–47.
Short, J., Williams, E., & Christie, B. (1976). The
social psychology of telecommunications.
New York: Wiley.
Singh, D., & Singh, D. (2011). Shape and signifi-
cance of feminine beauty: An evolutionary
perspective. Sex Roles, 64(9–10), 723–731.
Stephenson, W. (1988). The play theory of
mass communication. New Brunswick, NJ:
Transaction Publishers.
Stever, G. (1991). The celebrity appeal question-
naire. Psychological Reports, 68, 859–866.
Stever, G. (1994). Parasocial attachments:
Motivational antecedents. Doctoral Disserta-
tion, Arizona State University, Tempe, AZ.
Stever, G. (2013). Mediated vs. parasocial rela-
tionships: An attachment perspective. Jour-
nal of Media Psychology, 17(3), 1–31.
Stever, G. S. (2017a). Evolutionary theory and
reactions to mass media: Understanding
parasocial attachment. Psychology of Popu-
lar Media Culture, 6(2), 95.
Stever, G. (2017b). Parasocial theory: Concepts
and measures. In P. Rossler (Ed.), Interna-
tional Encyclopedia of Media Effects. (1–12).
doi: 10.1002/9781118783764.wbieme0069.
Tifferet, S., & Vilnai-Yavetz, I. (2014). Gender
differences in Facebook self-presentation: An
international randomized study. Computers
in Human Behavior, 35, 388–399.
Toth, N., & Schick, K. (1993). Early stone indus-
tries and inferences regarding language and
cognition. In K. R. Gibson, & T. Ingold (Eds.),
Tools, Language and Cognition in Human
Evolution, (346–362). Cambridge, UK: Cam-
bridge University Press.
Tukachinsky, R., & Stever, G. (2019). Theorizing
development of parasocial experiences.
Communication Theory 29(3) 297–318.
https://doi.org/10.1093/ct/qty032
Umilta, C., Simion, F., & Valenza, E. (1996).
Newborn’s preference for faces. European
Psychologist, 1(3), 200–205.
Vorderer, P., Steen, F., & Chan, E. (2006). Moti-
vation. In J. Bryan & P. Vorderer (Eds.), Psy-
chology of entertainment, (3–17). New York:
Routledge.
Weber, R. (2015). Biology and brains – methodo-
logical innovations in communication science:
Introduction to the special issue. Communica-
tion Methods and Measures, 9, 1–4.
West-Eberhard, M. J. (1979). Sexual selection,
social competition, and evolution. Proceed-
ings of the American Philosophical Society,
123(4), 222–234.
Wright, R. (1994). The moral animal: Why we
are the way we are. New York: Vintage Books.
Zald, D. (2003). The human amygdala and the
emotional evaluation of sensory stimuli.
Brain Research Review, 41, 88–123.

Evolutionary Psychology and
Communication
INTRODUCTION
Electronic collaboration (e-collaboration) is
collaboration using electronic technologies
among different individuals to accomplish a
common task. This is a broad definition that
encompasses not only computer-mediated
collaborative work, but also collaborative
work that is supported by other types of tech-
nologies that do not fit most people’s defini-
tion of a ‘computer’. One example of such
technologies is the telephone, which is
not, strictly speaking, a computer. Another
example of technology that may enable
e-­collaboration is the teleconferencing suite,
whose main components are cameras, televi-
sions, and telecommunications devices.
One phenomenon that has often puzzled
computer science and information-systems
researchers over the years, particularly
researchers interested in e-collaboration issues,
is the high importance of having an audio
channel for communication in the context of
e-collaborative tasks (Graetz et  al., 1998;
21
Ned Kock
Kock, 2004, 2009; Kock and DeLuca, 2007;
Wainfan and Davis, 2004). Whenever audio
is available (e.g., teleconferencing, telephone
conference calls, face-to-face meetings), tasks
seem to be performed more easily and with
fewer misunderstandings. Moreover, adding
video to an already present audio channel typ-
ically adds little to the e-­collaboration medi-
um’s ability to support group tasks (Burke
and Aytes, 2001). While this is not a univer-
sal phenomenon (see, e.g., Daly-Jones et al.,
1998; Baker, 2002; Kock et  al., 2015), its
frequent appearance in the empirical research
literature merits a more robust theoretical
analysis.
An evolutionary explanation of the reason
oral speech is needed for effective communi-
cation is proposed here, as a new theoretical
contribution to the e-collaboration literature.
It is argued that the high importance of oral
speech is restricted to knowledge-intensive
tasks. The reason for that, which is advanced
in more detail in the subsequent sections, is
that oral speech evolved among our hominid
418 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
ancestors as a costly trait to enable efficient
and effective knowledge communication.
As a costly trait, oral speech is analogous to
the large train displayed by male peacocks
to attract mates (often incorrectly called the
peacock’s tail).
Generally speaking, oral speech can be seen
as: (a) a survival handicap that only evolved
because of its strong indirect effect on repro-
ductive success, an effect that counteracts its
negative effect on survival; and (b) particu-
larly important in the context of the task for
which it evolved, namely communication of
knowledge. Nevertheless, it is acknowledged
here that even in knowledge-intensive tasks,
the negative influence of suppression of oral
speech may be countered by what is called
here ‘compensatory adaptation’, whereby
individuals adapt their communicative behav-
ior to overcome the limitations posed by the
suppression of oral speech. As it will be seen
later, the word ‘adaptation’ in the term ‘com-
pensatory adaptation’ does not refer to evolu-
tionary adaptation, but to adaptation that an
individual undergoes within his or her lifetime
(often a fraction of one’s lifetime).
COSTLY AND COSTLESS TRAITS:
SURVIVAL COSTS, REPRODUCTIVE
SUCCESS, AND THE HANDICAP
PRINCIPLE
Costly traits are phenotypic traits that evolved
in spite of having a negative impact on sur-
vival performance (Gillespie, 2004; Maynard
Smith, 1998; Rice, 2004). Survival perfor-
mance is the performance of an individual in
the general task of survival, which can be
measured by the age of the individual at the
time of death. The older an individual is, the
more successful it is at surviving in spite of
survival threats (e.g., disease, predators, and
accidental falls).
Costly traits evolve because they have a
positive impact on reproductive success (nor-
mally referred to as ‘fitness’ by evolutionary
b­ iologists), generally measured as the num-
ber of surviving offspring or grand-offspring
of an individual (Gillespie, 2004; Hartl and
Clark, 2007). The positive impact on fitness
results from the competing effects of a costly
trait on: (a) survival performance, a negative
effect; and (b) a task performance attribute, a
positive effect. The net effect of these com-
peting effects on fitness is positive, leading to
an increase in the frequency of the genotype
associated with the costly trait in the species.
One example of task performance attribute
that could lead to such a positive net effect on
fitness is the number of lifetime copulations an
individual participates in, a performance attrib-
ute associated with the task of mating. A clas-
sic example of a costly trait that evolved due to
having increased the number of lifetime copu-
lations individuals possessing the trait partici-
pated in, which in turn offset the survival cost of
that trait, is the peacock’s train (Maynard Smith
and Harper, 2003; Zahavi and Zahavi, 1997).
The peacock’s train is frequently referred to,
incorrectly, as the peacock’s tail (Petrie et al.,
1991; Zahavi and Zahavi, 1997). Both males
and females have tails, but only males have the
tail appendage known as a train.
Costly traits may also exist that have com-
peting effects on survival, and that are unre-
lated to mating, through intermediate effects
on other variables that themselves directly
affect survival. For example, propensity
toward aggressive behavior among our ances-
tors might have increased their chances of
being the target of violent behavior by other
individuals, which contributed to a decrease
in survival, but might also have increased their
access to nutritious food obtained through
hunting (for which aggressiveness is impor-
tant), which in turn contributed to an increase
in survival. In this sense, propensity toward
aggressive behavior might have evolved as a
costly trait, where the positive indirect effect
on survival, mediated by increased access to
nutritious food, was stronger than the nega-
tive indirect effect on survival from attracting
violent behavior (Boaz and Almquist, 2001;
Dobzhansky et al., 1977).
 EVOLUTIONARY PSYCHOLOGY AND COMMUNICATION
Costless traits are defined here as pheno-
typic traits that have no negative impact on
survival performance. Most costless traits are
actually associated with enhanced survival
performance, and may be observable indica-
tors of unobservable underlying traits that
enhance survival performance (Hamilton and
Zuk, 1982; Kokko et al., 2002). The ability of
males of the fruit fly species Drosophila sub-
obscura to engage in a rapid courtship dance
with females is an example of trait that fits
this definition (Maynard Smith and Harper,
2003). Males increase their success at the
task of mating by demonstrating to females
that they possess the ability to dance vigor-
ously in response to lead movements by the
females. This trait can be seen as a costless
trait, because it has no negative impact on
the survival success of males. In other words,
the dance itself has no negative effect on the
survival of males. Presumably the ability to
dance is positively correlated with survival
performance, since it is an indicator of health.
The most widely cited theoretical frame-
work in connection with the evolution of costly
traits was proposed by Zahavi (1975), centered
on what is known as the handicap principle
(Walker, 2008; Zahavi and Zahavi, 1997).
This framework is intuitively appealing, which
has led in part to its wide use in research by
evolutionary biologists, in general (Hausken
and Hirshleifer, 2008), and more specifi-
cally by evolutionary psychology researchers
(Griskevicius et al., 2007; Walker, 2008).
The handicap principle focuses on costly
traits used for signaling, and is founded on
the notion that those traits are honest indica-
tors of the signalers’ fitness. For example, the
large train displayed by peacocks is a survival
handicap, making them more vulnerable to
predation (Maynard Smith and Harper, 2003;
Zahavi and Zahavi, 1997). Consequently,
males with large trains and who are still alive
at the age of reproductive maturity also must
possess other traits that make them particu-
larly good at survival, such as vitality and
speed. The tails are a reliable indicator of fit-
ness, exactly because they are costly.
419
COMMON CHARACTERISTICS
OF COSTLY TRAITS: RARITY
AND STRONG EFFECTS
The discussion presented here expands on and
refines the handicap principle to cover any
costly trait, in connection with the perfor-
mance of any organism in any task that influ-
ences fitness, not only signaling tasks. Two
key conclusions are reached, which are that
costly traits should: be rare in nature, and be
costly not to use. The three sections that
follow this section provide a mathematical
elucidation and basis for these conclusions.
While these conclusions are consistent with
the handicap principle, they allow for predic-
tions and explanations that go beyond signal-
ing tasks. As such, they provide the basis for
the analysis of the evolution of oral speech
and its importance in the task of knowledge
communication.
Costly Traits Should Be Rare in
Nature
The survival handicaps imposed by costly
traits create obstacles for their evolution,
eventually making those traits significantly
rarer in nature than costless traits. These
obstacles can be seen as ‘thresholds’ for evo-
lution of the traits, where the thresholds are
proportional to the survival cost of the traits
(Gillespie, 2004; Hartl and Clark, 2007;
Maynard Smith and Harper, 2003).
New traits (e.g., high intelligence, long
legs, and slow fat metabolism) usually appear
in populations of organisms as a result of ran-
dom genetic mutations; a general rule that
applies to all organisms, including our homi-
nid ancestors (Boaz and Almquist, 2001; Hartl
and Clark, 2007; Mayr, 1976). Therefore, the
effects of new traits on fitness are also ran-
dom, whether those traits are costly or cost-
less. Evolution is not an engineering process;
it is a wasteful process of continuous tinker-
ing, where the vast majority of new traits
are in fact detrimental to fitness (Hartl and
420 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Clark, 2007; Wilson, 2000). Traits that have
a positive net effect on fitness are far and
few between (McElreath and Boyd, 2007;
Wilson, 2000).
Given that costly traits must overcome
obstacles, or thresholds, to evolve, fewer
costly traits than costless traits are likely to
evolve. That is, the probability of evolution
of costly traits in any species is generally
lower than that of costless traits. Moreover,
the higher the cost of the trait, the lower its
probability of evolution. Thus, costly traits
should be rarer in nature than costless traits;
the costlier, the rarer.
Costly Traits Should Be Costly
Not to Use
Costly traits must have had a strong effect on
the performance of the task for which they
evolved in order to make up for the survival
costs imposed by those traits. Today, this
would translate into a higher correlation
between costly traits’ measures and perfor-
mance attributes for the task, than between
costless traits’ measures and the same task
performance attributes. That is, not using a
costly trait would be costlier, so to speak,
than not using a costless trait in the context
of the task for which the traits evolved.
The above conclusions seem to be true
when we look at the classical example of
costly trait, the peacock train. Petrie et  al.
(1991) found that the costly ornamental train
of the male, and especially the number of
eyespots on the train, are far more attractive
traits for the peahens than other apparently
costless traits. Costless ornamental traits are
more numerous in the peacock species than
costly ones, of which the only known one is
the train, and their relative importance in the
context of the mating task is dwarfed by the
importance of the train. Examples of costless
ornamental traits likely evolved for mating
in the male of peafowl are the crest atop the
male’s head, the brightly colored feathers on
the male’s chest, various color patterns around
the eyes, various feather patterns occurring
in different parts of the male’s body, and the
level of bilateral (i.e., left–right) symmetry of
these ornamentations (Darwin, 1871; Zahavi
and Zahavi, 1997).
THE THRESHOLD FOR EVOLUTION
OF COSTLY TRAITS
One of the most important contributions to
mathematical evolutionary thinking was
made by Price (1970). He showed that for
any trait to evolve through selection, in any
population or subpopulation of individuals of
the same species, the trait must satisfy
Equation (1), whose main element is a covar-
iance term. The fitness of an individual that
possesses the trait (e.g., number of surviving
offspring) is measured through W, and Z is a
measure of the manifestation of the trait in
the individual (e.g., Z = 1 if the trait is pre-
sent, and Z = 0 if it is absent). The trait in
question can be any morphological, physio-
logical or behavioral trait; examples could be
opposing thumbs, aggressiveness, or a large
train (tail appendage) with many eyespots.
Cov(W, Z) > 0 (1)
Equation (1) can be re-written as Equation
(2) in terms of the standardized measures of
W and Z, referred to as w and z. This allows for
its use in the context of path analysis (Kock,
2011; Kock and Moqbel, 2016; Wright, 1934,
1960), which in turn greatly simplifies (as it
will be shown below) theoretical reasoning
based on comparative analyses of evolution
of traits through selection.
Cov(w · SW + W, z · SZ + Z)
= Cov(w · SW, z · SZ)
= SW · SZ Cov(w, z) > 0
⇒ Cov(w · z) > 0 (2)
Figure 21.1 shows a path model where
a costly trait measured by y is represented.
All the measures are standardized, which is
 EVOLUTIONARY PSYCHOLOGY AND COMMUNICATION 421

For a costless trait x, a trait with no negative

a effect on survival, Equation (3) is reduced to
pwa pay
Equation (4) because psy equals zero. What
w y this equation tells us is that a costless trait x
pas
will always evolve as long as it has a positive
pws psy causal relationship with a task performance
s attribute a, assuming that a has a positive
causal relationship with fitness (w).
Figure 21.1  Path model showing a costly pax > 0 (4)
trait and its relationship with fitness
In the task of mating, for example, any
costless trait x that increases mating success
why they are indicated with lowercase let-
(measured by a) would evolve through selec-
ters. The measure y has a positive causal rela-
tion, with trait frequency growth subject to the
tionship with a task performance attribute a.
constraints posed by chance events unrelated
For example, a could be number of lifetime
to the trait. That is, the trait would evolve to
copulations of an individual, a performance
the point of becoming widespread in a popu-
attribute associated with the mating task, in
lation only if it is not eliminated by chance
the case of a trait used for mate choice.
from the population at its early stages of evo-
The measure y has a negative causal rela-
lution; e.g., the only individual that initially
tionship with s, a measure of survival per-
possesses the trait is killed by a lightning
formance. For example, s could be age of an
strike before reaching reproductive matu-
individual at the time of death. Since any indi-
rity (Gillespie, 2004; Wen-Hsiung, 2000). A
vidual must be alive to perform any task, s also
costly trait y (e.g., the peacock’s train), on the
has a positive causal relationship with a. Both
other hand, would have to meet a more strin-
a and s have positive causal relationships with
gent requirement for evolution. It would only
fitness (w). The magnitudes of these relation-
evolve through selection if the trait’s positive
ships are given by the path coefficients pay, psy
effect on a surpassed the threshold given by
etc. All path coefficients are positive except
the right side of Equation (3).
for psy, which is negative since it refers to the
survival cost of trait y. (For simplicity, a trait
measured by y is also called trait y.)
In path analysis the covariance between PROBABILITY OF EVOLUTION OF
any pair of variables is given by the sum of
COSTLY TRAITS
the products of the path coefficients in all
paths connecting the two variables (Wright,
Let us assume that the appearance of a new
1934, 1960). Thus, combining Equation (2)
costly trait y in a population will lead to a
with Figure 21.1 leads to Equation (3), which
variation in pay and –psy that will be given by
must be satisfied for any costly trait y to
random numbers going from 0 to D. Let us
evolve through selection.
also represent T as in Equation (5):
pwa ∙ pay + pws ∙ psy + pwa ∙ pas ∙ psy > 0
 p 
 
⇒pwa ∙ pay > –psy ∙ (pws + pwa ∙ pas) T =  ws + pas  (5)
 p 
 wa 
⇒pay > –psy∙ (pws
pwa + pas)

 p  The value of T is assumed here to be

 
⇒ pay > − psy ⋅  ws + pas  (3) largely population specific, in a relatively
 p 
 wa  stable environment, and thus should remain
422 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
relatively constant as new costly or costless
traits appear in a population and either evolve
or disappear in response to selection pres-
sures. This can be illustrated for the task of
mating, where a can be the number of life-
time copulations a male of a species engages
in. In this case, the effect of a on fitness (w),
the effect of survival performance (s) on w,
and the effect of s on a are relatively constant
for the males of the species.
In a relatively stable environment this can
be shown to hold for any task whose perfor-
mance is measured by a. This conclusion
also follows from the assumption that those
effects can be represented through stable
regression coefficients, an assumption that
is routinely used in mathematical popula-
tion genetics models (Gillespie, 2004; Kock,
2011; Kock and Moqbel, 2016; Rice, 2004).
On the other hand, the effects that new
traits x or y have on a and/or s can vary
widely, since those traits appear in the pop-
ulation as a result of stochastic processes.
Those effects will in turn ultimately dictate
whether those traits will evolve or disappear
in the species. For species that live today in
environments similar to those in which most
of their traits evolved, the value of T can be
easily estimated empirically since the path
coefficients are standardized partial regres-
sion coefficients.
Given the above assumptions, the prob-
ability of evolution of a new costly trait y in
a population will be given by Equation (6),
assuming that T is equal to or greater than
one. This equation reflects the intersection
spaces of variation of D and DT, and can eas-
ily be verified through simple Monte Carlo
simulations.
D 1
( )
P Evo ( y ) = ⋅
DT 2
1 and likely of the hominid ancestors in the
(
⇒ P Evo ( y ) = )
2⋅ T
(6)
T is assumed to be equal to or greater than
one because it is difficult to conceive of a spe-
cies population or subpopulation for which
the performance of a task is more important
for fitness than survival, even for the all-
important task of mating. Let us consider,
for example, spider species where the males
are routinely cannibalized by their large and
aggressive female mates during or after copu-
lation (see, e.g., Wilder and Rypstra, 2008).
In these species, the male spiders must still
successfully survive up to the moment of
copulation. Therefore, when looked at as a
subpopulation of the species to which they
belong, those male spiders will likely have a
ratio pws / pwa that is greater than 1 (and thus
a T greater than 1) for the task of mating,
regardless of the fact that they contribute lit-
tle more than their sperm to the survival of
their offspring (and thus to their own fitness).
Equation (6) can be depicted in a graph, as
shown in Figure 21.2. The graph shows the
variation of the probability of evolution of
a new costly trait y in a population (vertical
axis) based on values of T ranging from 1 to
10 (horizontal axis).
As can be inferred from Figure 21.2, costly
traits will always have a lower probability of
evolution than costless traits, because the
value of T for the latter traits is always zero.
This suggests that costly traits should be rarer
in nature than costless ones, regardless of the
task for which they were evolved, e.g., mat-
ing, communication, fighting.
Moreover, costly traits should be particu-
larly rare in species where the value of T is
high. This would be the case in species where
the number of offspring born to females was
small; and in species where the offspring
relied heavily on their parents for survival
in their early years of life, when most deaths
occur. (In these species, the effect of survival
on fitness would have been much higher
than the effect of mating on fitness.) These
are characteristics of the human species,
human lineage (Boaz and Almquist, 2001;
Cartwright, 2000). Thus, T values should
have been high for our hominid ancestors,
making the evolution of costly traits diffi-
cult. This line of reasoning would suggest
 EVOLUTIONARY PSYCHOLOGY AND COMMUNICATION
0.5
0.45
0.4
0.35
0.3
0.25
0.2
0.15
0.1
0.05
0
1 2 3 4 5 6
Figure 21.2  Probability of evolution of a new costly trait
the existence of moderating effects in our
model, which are not included because their
discussion would be beyond the scope of this
chapter. This has no effect on the generality
of the discussion presented here or the related
conclusions.
Of course, in order to evolve, costly and
costless traits also have to satisfy the con-
dition that their covariance with fitness is
greater than zero (i.e., that they have a posi-
tive net impact on fitness), which rarely is
the case for new genetic mutations. Most
new genetic mutations have either a nega-
tive or neutral effect on fitness; in the latter
case they may evolve by chance, through a
process known as genetic drift (Gillespie,
2004; Hartl and Clark, 2007; Maynard Smith,
1998). Costly traits, unlike costless ones,
have another condition to satisfy: they must
overcome the survival costs that they impose.
DIFFERENT EFFECTS OF COSTLY
AND COSTLESS TRAITS
Let us also assume that the appearance of
costly and costless traits in a population will
lead to random values of pax and –psy in the
range from 0 to D. The expected value of pax
423
7 8 9 10
for costless traits that evolve will then be given
by D divided by 2. The expected value of pay,
on the other hand, will be given by (DT–D)/2,
assuming that T is equal to or greater than 1.
Therefore, the expected ratios between pay
and pax will be given by Equation (7).
) ( D/2 )
D⋅ T − D /2
(
E pay / pax =
⇒ E( p / p ) = T −1
ay ax
(7)
The graph in Figure 21.3 shows the vari­
ation of the expected ratio between pay
and pax (vertical axis) based on values of T
ranging from 1 to 10 (horizontal axis). The
ratio grows proportionally with T, and is a
measure of how strong the expected effect
of a costly trait y on the task performance
attribute a is, compared with the expected
effect of a costless trait x. For simplicity, it
is assumed here that both types of traits are
either independent from each other, or spread
to fix­ation in a species at different points in
time. For dependent traits or traits that evolve
at the same time, the mathematical analysis
becomes more complex, but the results are
qualitatively the same.
An expected ratio between pay and pax of
1.3, for example, means that the standardized
424 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
9
8
7
6
5
4
3
2
1
0
1 2 3 4 5 6
Figure 21.3  Variation of the expected ratio between pay and pax
effect of any costly trait on a given task
performance attribute a is on average 30%
stronger than the effect of any costless trait
on the same task performance attribute. A
ratio of 8 means that the costly trait is on aver-
age 800% stronger (e.g., pay=.4 and pax=.05).
Standardized effects are expressed in terms of
standard deviations of the variables to which
they refer. For example, a pay=.4 means that a
1 standard deviation variation in y causes a .4
standard deviation variation in a.
THE EVOLUTION OF ORAL SPEECH
IN HUMANS: A COSTLY TRAIT
ASSOCIATED WITH CHOKING AND
ILLNESSES
Modern oral speech was enabled by the evo-
lution of a larynx located low in the neck
(Lieberman, 1998). The evolution of oral
speech is one of the most important land-
marks in the evolution of the human species,
having happened relatively recently in our
evolutionary history (Figure 21.4). However,
the new larynx design also likely increased
significantly our ancestors’ chances of death
by choking during ingestion of food and liq-
uids, a leading cause of death among modern
humans (Hemsley et  al., 2019), and of
7 8 9 10
suffering from various aerodigestive tract
diseases such as gastroesophageal reflux
(Laitman and Reidenberg, 1997), among
other survival-related problems. Oral speech
must have been particularly important for
effective communication in our evolutionary
past, and effective communication must have
been important for fitness enhancement
(Pinker, 2003), otherwise its survival cost
would have prevented complex speech from
evolving.
The situation with gastroesophageal reflux
is particularly telling with respect to the cost
of the evolution of oral speech in humans.
The aerodigestive tract adaptations that
enable complex oral speech make modern
humans differ from their Australopithecine
ancestors in ways that are analogous to the
differences between the aerodigestive tracts
of modern humans and modern non-human
primates. In modern humans the incidence of
gastroesophageal reflux symptoms (at various
degrees of severity) has been reported to be as
high as 40%, and yet Glover et al. (2008) noted
that no modern non-human primate species
was reported to display naturally occurring
symptoms. The occurrence of gastroesopha-
geal reflux symptoms is associated with signif-
icantly greater rates of mortality among infants
(Vandenplas et al., 1991). This is also the case
among adults, due to various conditions,
 EVOLUTIONARY PSYCHOLOGY AND COMMUNICATION
Emergence of
Australopithecines
3.5 2.0 1.5
Co-located and synchronous
communication through facial
expressions, body language,
and simple sounds
Figure 21.4  The evolution of oral speech in humans
including hemorrhagic reflux esophagitis
(Rantanen and Salo, 1999).
Oral speech seems to exhibit the two com-
mon characteristics of costly traits. Oral
speech is a rare costly trait among human
traits involved in the transfer of communi-
cative stimuli. By all accounts, it is the only
such trait that obviously imposed a survival
cost as it evolved among our ancestors. In
addition to increasing our ancestors’ chances
of death by choking, and of developing aero­
digestive tract diseases, it also hampered our
ancestors’ ability to breathe while drinking
water. Water sources are likely to have been
a preferred site for predators to ambush prey
(Boaz and Almquist, 2001), as they are today,
and the oxygen depletion caused by having
to hold their breath while drinking created
yet another survival cost for our ancestors.
Other communication-related traits, such as
the ability to use body language and facial
expressions, do not seem to have imposed a
similar survival handicap on our ancestors.
Oral speech also appeared late in the evo-
lutionary history of hominids, in the last
100,000 years of that 3.5-million-year history,
or approximately the last 3% (Cartwright,
2000; Laitman, 1984; Lieberman, 1998).
This is consistent with it being a costly trait,
425
Emergence of
Homo sapiens
Today
Millions of
years
1.0 0.1
Complex
oral
speech
since the evolution of a costly trait is a low-
probability event, and low-probability events
frequently take more time to happen than
high-probability events. In fact, the evolu-
tion of oral speech coincides with the evo-
lution of our species, Homo sapiens, likely
from another species within the genus Homo,
namely Homo erectus (Boaz and Almquist,
2001; Lieberman, 1998). Many human evo-
lution researchers believe that it was the evo-
lution of oral speech, with the complexity of
human interactions that it enabled, that made
us truly human (Cartwright, 2000; Dunbar,
1993; Lieberman, 1998).
Finally, empirical research on the effects
of electronic communication media that sup-
press the ability to use oral speech suggests
that it is costly not to use oral speech in com-
municative interactions. This is reflected in
as much as a 10-fold reduction in commu-
nication fluency, coupled with a significant
increase in communication ambiguity and
perceived cognitive effort (Graetz et al., 1998;
Kock, 2005; Kock and DeLuca, 2007; Kock
et  al., 2007; Simon, 2006). Communication
fluency is defined here as the number of ideas
effectively conveyed per unit of time, and
has been somewhat imprecisely measured
as the number of words conveyed per unit of
426 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

Table 21.1  Oral speech and the two common characteristics of costly traits
Common characteristic of costly traits Evidence in connection with oral speech

Costly traits should be rare in nature
Costly traits should be costly not to
use
Oral speech is the only communication-related trait that clearly imposed a survival
cost on our ancestors; conversely, various related costless traits seem to exist –
e.g., the ability to use body language and facial expressions for communication.
Suppressing oral speech in electronic communication media is costly, leading
to as much as a 10-fold reduction in communication fluency, coupled with
a significant increase in communication ambiguity and perceived cognitive effort.

time (Kock, 2005; Kock and DeLuca, 2007). It
seems that, when oral speech is removed from
a communication medium, communication
becomes rather cumbersome (Crowston et al.,
2007; Graetz et al., 1998; Kock, 2005). Table
21.1 summarizes the discussion so far regard-
ing the relationship between oral speech and
the two common characteristics of costly traits.
If the use of oral speech is enabled by an
audio channel, adding a video channel typi-
cally has little impact on the effectiveness
or ease with which communication takes
place (Burke and Aytes, 2001; Daly-Jones
et al., 1998; Simon, 2006). In this sense, oral
speech could be seen to the communication
task as analogous to what the peacock’s train
is to the mating task (Petrie et al., 1991); both
are costly traits that have an effect that dwarfs
the effects of other costless traits evolved in
connection with the same task. There are
exceptions to this general rule regarding the
importance of a video channel (see, e.g.,
Daly-Jones et  al., 1998), such as modern
tasks in which shared and real-time visuali-
zation of an object or situation is important
for the task completion. Examples would be
a surgical intervention involving two or more
geographically distributed doctors, and a
real-time collaborative design of a car engine.
ORAL SPEECH AND KNOWLEDGE
COMMUNICATION: THE FOCUSED
IMPACT ON FITNESS OF THIS
COSTLY TRAIT
The notion that oral speech is particularly
important in modern human communication,
as discussed so far, needs further theoretical
elaboration and refinement. Simple observa-
tion of modern human communication prac-
tices suggests that oral speech is not equally
important for all types of communication
interactions. For example, if one person is
trying to communicate his or her home or
work address to another, to be used on a
letter, then probably an e-mail will be just as
effective as a phone call. Also, web-based
social communication tools that enable
human interaction through short text mes-
sages and provide no audio channel, such as
in the early versions of Twitter, would prob-
ably not be as successful as they are if the
theoretical framework put forth here applied
to all types of communicative interactions.
This takes us back to a review of why oral
speech evolved in the first place. More spe-
cifically, how did oral speech affect fitness
among our ancestors? As discussed earlier,
only if oral speech had a net positive impact
on fitness, by enhancing the performance of
a fitness-relevant task, would it have over-
come the survival handicap associated with
our customized vocal tract. The answer is
that oral speech enabled the exchange of
knowledge among our ancestors, which indi-
rectly increased their reproductive success by
allowing them to occupy what Pinker (2003)
refers to as the ‘cognitive niche’.
A common characteristic of the simple
exchanges illustrated above (communication
of a home or work address and interaction
through short text messages) is that these types
of exchanges involve little or no knowledge
transfer. Therefore, if we assume that oral
speech was evolved by our ancestors primarily
to enable the communication of knowledge,
 EVOLUTIONARY PSYCHOLOGY AND COMMUNICATION
its effect should not be particularly strong in
communication interactions with little or no
knowledge content. There are other factors
that may induce modern humans to commu-
nicate electronically through text only, and
with no audio – e.g., via e-mail without audio
file attachments. Among those factors is the
ease with which e-mail can be sent to many
individuals at the same time. Video and audio
blogs can be used for the same purpose, incor-
porating oral speech, but their use is still not as
widespread and embedded in communication
practices as is the use of e-mail.
Knowledge about ‘something’ is defined
here in a way that is analogous to how it is
defined by many cognitive psychologists: as
a set of mental schemas that allows one to
predict the future, or find out more about a
present situation, based on information about
the present or the past (Gardner, 1985; Kock,
1999; Lee and Holyoak, 2008; Waldmann
et  al., 1995). As noted by artificial intelli-
gence researchers, with knowledge, one can
build mental rules that can be expressed in
the form of ‘if … then …’ statements (Luger
and Stubblefield, 2008; Russel and Norvig,
2002), or reworded as statements that con-
tain linguistic elements that express causal-
ity such as ‘the reason for … is …’, ‘this
is … because …’, and ‘the cause for … is
…’ (Kock, 1999; Waldmann et  al., 1995).
For example, the statement ‘the temperature
in room 118, where 100 people are attend-
ing a lecture, is now 78 degrees Fahrenheit’
contains only information, whereas the fol-
lowing statement contains knowledge: ‘if the
temperature in room 118 reaches 80 degrees
Fahrenheit, most of the 100 people attending
a lecture there will feel uncomfortable’.
Our ancestors faced survival threats on
a regular basis – exposure to pathogens,
attacks by predators or territorial animals,
encounters with venomous insects or snakes,
and ingestion of toxins, among others. These
events often occurred in specific contexts.
For example, territorial animals would attack
when their habitat was invaded by our ances-
tors, and venomous insects and snakes occur
427
in higher quantities in certain areas (Hung,
2004; Kock et  al., 2008; Manipady et  al.,
2006). Without the ability to vicariously
obtain knowledge linking contexts with sur-
vival threats, our ancestors would have to
experience the survival threats, or observe
someone experiencing them at a close dis-
tance, in order to build that knowledge. Oral
speech enabled vicarious knowledge acquisi-
tion regarding survival threats, and thus sig-
nificantly increased our ancestors’ chances of
survival, easily overcoming the extra survival
costs associated with our vocal tract.
Costly traits evolved by our human ances-
tors must have had a strong effect on the per-
formance of the task for which they evolved,
in order to make up for the survival costs
imposed by those traits. In the case of oral
speech, a strong candidate for the task in
question is the knowledge communication
task, where oral speech evolved in part to
increase the performance with which knowl-
edge about survival threats was communi-
cated among our ancestors. Oral speech may
also have influenced fitness in other ways,
although avoidance of survival threats must
have been an important element in the selec-
tion of this costly trait. For example, vicari-
ous knowledge about s­urvival-enhancing
elements, such as seasonal availability of
food, was likely also enabled by oral speech
(Cartwright, 2000; Dunbar, 1999). So proba-
bly was the ability to build social relationships
and court potential mates (Dunbar, 1993;
Miller, 2000, 2002). This type of knowledge
communication likely required reciprocal
altruism to have evolved before, which math-
ematical formalizations and empirical evi-
dence strongly suggest was the case in the
human species (Fletcher and Zwick, 2007;
Henrich, 2004; McElreath and Boyd, 2007;
Trivers, 2002).
Knowledge communication performance
refers to both the effectiveness and effi-
ciency with which knowledge is communi-
cated (Kock, 1999; Russel and Norvig, 2002;
Waldmann et al., 1995). Effective knowledge
communication between two individuals
428 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
occurs when the knowledge possessed by
one individual is comprehensively and unam-
biguously conveyed to the other individual.
Efficient knowledge communication occurs
when the knowledge possessed by one indi-
vidual is quickly and effortlessly conveyed to
the other individual.
THE EFFECTS OF ORAL SPEECH
IN E-COLLABORATIVE TASKS:
COMMUNICATION FLUENCY
AND AMBIGUITY
It follows from the theoretical discussion
presented in the previous section that remov-
ing the ability to convey speech from an
electronic communication medium is likely
to impair communication performance much
more strongly than removing the medium’s
ability to convey other communicative stim-
uli – e.g., facial expressions, body language,
olfactory cues, and tactile stimuli. However,
this effect is moderated by the extent to
which knowledge is being communicated.
This conclusion is consistent with the results
of various studies that compare the impact of
various media on communication perfor-
mance (Graetz et  al., 1998; Kock, 2004;
Kock and DeLuca, 2007).
Graetz et  al. (1998) compared the perfor-
mance in four-person groups across three com-
munication media conditions: face-to-face,
telephone conferencing, and electronic chat.
The experimental task required exchange of
knowledge to be successfully accomplished,
and the participants were given a limited
amount of time (approximately 30 minutes)
to review the information provided to them
by the researchers and to discuss it with the
other group members. Group outcome quality
was about the same through the face-to-face
and telephone-conferencing media; slightly
higher in the latter, a statistically insignificant
difference. Group outcome quality was sig-
nificantly lower through the electronic chat
medium. Measures of perceived cognitive
effort and frustration were about the same for
the ­face-to-face and telephone-­conferencing
media, and significantly higher for the
electronic-chat medium. In summary, the
­
medium that did not enable oral speech was
the least conducive to effortless and unam-
biguous knowledge communication. This is
consistent with the view that oral speech is
a costly trait that is ‘costly not to use’ in the
context of knowledge communication.
Particularly noteworthy is the finding by
Kock and DeLuca (2007), in a study of indi-
viduals in two different countries, that the use
of an electronic communication medium that
suppressed the ability to convey speech (a
version of e-mail) dramatically reduced com-
munication fluency. In this study, communi-
cation fluency was measured as the number of
words conveyed per unit of time, a surrogate
measure. The reduction in fluency observed
by Kock and DeLuca was estimated to have
been more than 10-fold; that is, e-mail users’
fluency was less than 1/10 of their expected
fluency communicating over the phone or
face-to-face. This is too drastic a reduction to
be explained by the known fact that typing is
mechanically more cumbersome than speak-
ing, which would normally lead to a twofold
reduction in fluency (Kock, 2004; McQueen
et al., 1999). Again, it appears that our brain
was ‘designed’ by evolution to rely heavily on
oral speech for effective and efficient knowl-
edge communication, because oral speech
was costly to evolve. As a result, it is costly
not to use oral speech in modern human com-
munication whenever a significant amount of
knowledge must be exchanged.
COMPENSATORY ADAPTATION:
COUNTERACTING THE EFFECTS OF
ORAL SPEECH SUPPRESSION BY
E-COLLABORATION TECHNOLOGIES
A possible conclusion based on the arguments
presented thus far is that a decrease in commu-
nication fluency and an increase in ambiguity,
 EVOLUTIONARY PSYCHOLOGY AND COMMUNICATION
caused by the suppression of oral speech in
an electronic medium, may lead to a decrease
in the quality of the outcomes accomplished
by a group using the medium for most of its
communication. Indeed, this seems to fre-
quently be the case in short-duration tasks
(Graetz et al., 1998; Kahai and Cooper, 2003;
Warkentin et al., 1997), but not necessarily in
long-duration tasks (Burke and Chidambaram,
1999; Carlson, 1995; DeLuca, 2003; Kock,
2005; Kock and DeLuca, 2007). The reason
is that, in long-duration tasks, it is common
to observe a phenomenon known as compen-
satory adaptation (Kock, 2002). This phe-
nomenon may counteract the problems
associated with the suppression of oral
speech (Kock, 2005; Kock et al., 2007).
An example of a short-duration task would
be to write a one-page contract to sell a car,
where the contract would be written in one
hour. A long-duration equivalent would be
to write a 20-page contract to sell a fleet of
cars, where the longer contract would be
developed over a period of two weeks. If an
electronic medium is to be used for commu-
nication to accomplish these tasks, where
oral speech is suppressed in the communica-
tion medium, we would expect a more drastic
drop in quality for the one-page contract than
for the longer one with 20 pages. Here, ‘qual-
ity’ could be measured based on a number of
factors, including the degree of understand-
ability of the contract by the parties involved
and the degree to which the contract clauses
would be legally enforceable.
Compensatory adaptation seems to be
one of the reasons why groups performing
knowledge-intensive tasks over a relatively
long period of time (e.g., days, weeks, or
months), using an e-collaboration medium
that suppresses oral speech, often have the
same or even better performance than groups
where oral interaction is not suppressed
(Kock, 2005). As long as there is motivation
among group members to expend additional
compensatory effort, which may be strongly
influenced by social factors (Bandura, 1986;
Fulk, 1993), group members are likely to
429
adapt their communicative behavior in order
to compensate for the obstacles posed by the
e-collaboration medium’s suppression of oral
speech (Short et al., 1976; Ulijn et al., 2001).
Compensatory adaptation can be under-
stood as a moderating effect. That is, the
effects of oral speech suppression on com-
munication fluency and ambiguity are mod-
erated by compensatory adaptation, whose
moderating effect is in turn positively cor-
related with e-collaborative task duration.
In short-duration tasks, the negative effects
of oral speech suppression on communica-
tion fluency and ambiguity are likely to be
particularly acute, as there is no time for
compensatory adaptation to take place. In
long-duration tasks, the e-collaborators may
adapt their behavior to compensate for the
cognitive obstacles caused by the suppres-
sion of oral speech. This phenomenon has
been referred to as compensatory adaptation
to e-collaboration media of low naturalness
(Kock, 2004).
DISCUSSION AND CONCLUSION
The arguments presented in the previous
sections can be summarized into three main
predictions. The first refers to the effects of
oral speech suppression on communication
fluency and ambiguity in the context of
e-collaboration. The second refers to the
moderating effect that the amount of
knowledge communicated is likely to have
on these effects. The third prediction refers to
the moderating effect that compensatory
adaptation is likely to have on the effects of
oral speech suppression on communication
fluency and ambiguity. Compensatory
adaptation itself is correlated with task
duration, and may take place even when a
large amount of knowledge is being
communicated. These predictions are outlined
below, and followed by recommendations
for the use of e-collaboration tools in
organizations.
430 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Communication Fluency
and Ambiguity
A key prediction based on the discussion put
forth here is that removing the ability to
convey speech from an e-collaboration
medium used by modern humans is likely to
decrease communication fluency and increase
communication ambiguity much more strongly
than removing the medium’s ability to convey
other communicative stimuli such as facial
expressions and body language. The reason is
that the ability to use speech for communica-
tion evolved at a much higher survival cost
among our human ancestors than the ability to
use any other communicative stimulus.
The Moderating Effect of
Knowledge Communication
The negative effects of oral speech suppression
on communication fluency and ambiguity are
moderated by the amount of knowledge com-
munication taking place in an e-collaborative
task. Due to the context in which oral speech
evolved among our ancestors, oral speech is not
equally important for all types of communica-
tion interactions among modern humans; it is
particularly important in knowledge-intensive
communication. Communicating one’s home
address to another person, for example, can be
easily and effectively accomplished through
e-mail. Conversely, if one engineer wants to
communicate knowledge about how to design a
new car engine to a production manager, then
the suppression of oral speech may be
problematic.
The Moderating Effect of
Compensatory Adaptation
Another moderating effect, similar to but of a
different kind than knowledge communication,
is compensatory adaptation. Compensatory
adaptation, or the degree to which individuals
adapt to a communication medium that is
unnatural (e.g., one that suppresses oral
speech), seems to moderate the negative effects
of oral speech suppression on communication
fluency and ambiguity. Compensatory adapta-
tion to media that suppress oral speech typi-
cally happens over time (e.g., days, weeks or
months), as individuals modify their commu-
nicative behavior to make up for the shortcom-
ings of the medium. This may be one of the
reasons why compensatory adaptation is not
normally observed in short-duration tasks requir-
ing intense knowledge exchange. For example,
groups performing knowledge-intensive tasks
through text-based e-collaboration technolo-
gies, and where the tasks last from a few min-
utes to a few hours, generally tend to produce
task outcomes of inferior quality. These groups
would be better off either: (a) performing the
task face-to-face or using an e-collaboration
technology that provides an audio channel; or
(b) performing the task using a text-based
e-collaboration technology, but over a long
time period (e.g., a few days) so that compen-
satory adaptation can take place.
The increasingly distributed nature of
organizational processes (e.g., sets of activi-
ties that are repeated over and over again) and
projects (e.g., sets of activities that are carried
out once or a few times) requires tasks to be
accomplished by groups of individuals who
are not only geographically distributed, but
also distributed across multiple time zones.
Given this, it is impractical to try to ensure
that all activities in a process or project are
performed face-to-face, or even through
e-­collaboration involving synchronous oral
speech interactions. Sometimes ubiquitous
text-based asynchronous communication such
as e-mail must be used for part of the process
or project, due to cost constraints. It is also
possible that asynchronous oral speech inter-
actions will be used (e.g., voice messaging or
e-mail with attached audio messages) for part
of the process or project, due to group mem-
bers having to work from different time zones.
A more practical piece of advice to manag-
ers, which follows from the theoretical discus-
sion, is the following: (a) break organizational
 EVOLUTIONARY PSYCHOLOGY AND COMMUNICATION
processes and projects into component collabo-
rative activities; (b) rank those activities in terms
of the perceived amount of knowledge exchange
involved; (c) make sure that highly knowledge-
intensive activities are performed through media
that incorporate synchronous oral speech (e.g.,
face-to-face or teleconferencing interaction),
which may mean that certain group members
will have to make special accommodations to
participate in group discussions (e.g., attend a
meeting at 3 a.m., local time); (d) make sure that
moderately knowledge-intensive activities are
performed through media that incorporate some
form of oral speech, even if asynchronous (e.g.,
voice messaging or e-mail with attached audio
messages); and (e) encourage the use of text-
based e-collaboration media for activities that
involve little or no knowledge exchange among
participants, as this is likely to be the cheapest
and most widely available organizational com-
munication medium.
ACKNOWLEDGMENTS
This book chapter is based on a previous ver-
sion published as an article in the journal
Electronic Markets, and also contains revised
materials and ideas, originally developed by
the author, from articles published in the fol-
lowing journals: International Journal of
e-Collaboration; Cognition, Technology &
Work; International Journal of Technology
and Human Interaction; Journal of Systems
and Information Technology; and Journal of
Evolutionary Psychology. This is done with
full permission by the copyright holders, and
with the goal of timely and wide dissemina-
tion of scholarly work.
REFERENCES
Baker, G. (2002). The effects of synchronous col-
laborative technologies on decision making: A
study of virtual teams. Information Resources
Management Journal, 15(4), 79–94.
431
Bandura, A. (1986). Social foundations of
thought and action. Englewood Cliffs, NJ:
Prentice Hall.
Boaz, N.T., & Almquist, A.J. (2001). Biological
anthropology: A synthetic approach to
human evolution. Upper Saddle River, NJ:
Prentice Hall.
Burke, K., & Aytes, K. (2001). Do media really
affect perceptions and procedural structuring
among partially-distributed groups? Journal
on Systems and Information Technology, 5(1),
10–23.
Burke, K., & Chidambaram, L. (1999). How
much bandwidth is enough? A longitudinal
examination of media characteristics and
group outcomes. MIS Quarterly, 23(4),
557–580.
Carlson, J.R. (1995). Channel expansion theory:
A dynamic view of media and information
richness perception. Doctoral dissertation.
Tallahassee, FL: Florida State University.
Cartwright, J. (2000). Evolution and human
behavior: Darwinian perspectives on human
nature. Cambridge, MA: The MIT Press.
Crowston, K., Howison, J., Masango, C., &
Eseryel, U.Y. (2007). The role of face-to-face
meetings in technology-supported self-
organizing distributed teams. IEEE Transac-
tions on Professional Communication, 50(3),
185–203.
Daly-Jones, O., Monk, A., & Watts, L. (1998).
Some advantages of video conferencing over
high-quality audio conferencing: Fluency and
awareness of attentional focus. International
Journal of Human-Computer Studies, 49(1),
21–58.
Darwin, C.R. (1871). The descent of man, and
selection in relation to sex. London, England:
John Murray.
DeLuca, D.C. (2003). Business process improve-
ment using asynchronous e-collaboration:
Testing the compensatory adaptation model.
Doctoral dissertation. Philadelphia, PA:
Temple University.
Dobzhansky, T., Ayala, F.J., Stebbins, G.L., &
Valentine, J.W. (1977). Evolution. San Fran-
cisco, CA: W.H. Freeman and Company.
Dunbar, R.I.M. (1993). Coevolution of neocortical
size, group size and language in humans.
Behavioral and Brain Sciences, 16(4), 681–735.
Dunbar, R.I.M. (1999). Culture, honesty and
the freerider problem. In: R.I.M. Dunbar,
432 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
C. Knight, & C. Power (Eds.), The evolution
of culture (pp. 194–213). New Brunswick,
NJ: Rutgers University Press.
Fletcher, J.A., & Zwick, M. (2007). The evolution
of altruism: Game theory in multilevel selection
and inclusive fitness. Journal of Theoretical
Biology, 245(1), 26–36.
Fulk, J. (1993). Social construction of commu-
nication technology. Academy of Manage-
ment Journal, 36(5), 921–938.
Gardner, H. (1985). The mind’s new science.
New York, NY: Basic Books.
Gillespie, J.H. (2004). Population genetics. Balti-
more, MD: The Johns Hopkins University Press.
Glover, E.J., Leland, M.M., Dick Jr, E.J., & Hub-
bard, G.B. (2008). Gastroesophageal reflux
disease in baboons (Papio sp.): A new animal
model. Journal of Medical Primatology,
37(1), 18–25.
Graetz, K.A., Boyle, E.S., Kimble, C.E., Thomp-
son, P., & Garloch, J.L. (1998). Information
sharing in face-to-face, teleconferencing,
and electronic chat groups. Small Group
Research, 29(6), 714–743.
Griskevicius, V., Tybur, J.M., Sundie, J.M., Cial-
dini, R.B., Miller, G.F., & Kenrick, D.T. (2007).
Blatant benevolence and conspicuous con-
sumption: When romantic motives elicit
costly displays. Journal of Personality and
Social Psychology, 93(1), 85–102.
Hamilton, W.D., & Zuk, M. (1982). Heritable
true fitness and bright birds: A role for para-
sites? Science, 218(4570), 384–387.
Hartl, D.L., & Clark, A.G. (2007). Principles of
population genetics. Sunderland, MA: Sin-
auer Associates.
Hausken, K., & Hirshleifer, J. (2008). Truthful
signalling, the heritability paradox, and the
Malthusian equi-marginal principle. Theo-
retical Population Biology, 73(1), 11–23.
Hemsley, B., Steel, J., Sheppard, J.J., Maland-
raki, G.A., Bryant, L., & Balandin, S. (2019).
Dying for a meal: An integrative review of
characteristics of choking incidents and rec-
ommendations to prevent fatal and nonfatal
choking across populations. American Jour-
nal of Speech-Language Pathology, 28(3),
1283–1297.
Henrich, J. (2004). Cultural group selection,
coevolutionary processes and large-scale
cooperation. Journal of Economic Behavior &
Organization, 53(1), 3–35.
Hung, D.-Z. (2004). Taiwan’s venomous
snakebite: Epidemiological, evolution and
geographic differences. Transactions of the
Royal Society of Tropical Medicine and
Hygiene, 98(2), 96–101.
Kahai, S.S., & Cooper, R.B. (2003). Exploring
the core concepts of media richness theory:
The impact of cue multiplicity and feedback
immediacy on decision quality. Journal of
Management Information Systems, 20(1),
263–281.
Kock, N. (1999). Process improvement and
organizational learning: The role of collabo-
ration technologies. Hershey, PA: Idea Group
Publishing.
Kock, N. (2002). Compensatory adaptation:
Understanding how obstacles can lead to
success. Haverford, PA: Infinity Publishing.
Kock, N. (2004). The psychobiological model:
Towards a new theory of computer-mediated
communication based on Darwinian evolu-
tion. Organization Science, 15(3), 327–348.
Kock, N. (2005). Compensatory adaptation to
media obstacles: An experimental study of
process redesign dyads. Information Resources
Management Journal, 18(2), 41–67.
Kock, N. (2009). Information systems theorizing
based on evolutionary psychology: An inter-
disciplinary review and theory integration
framework. MIS Quarterly, 33(2), 395–418.
Kock, N. (2011). A mathematical analysis of the
evolution of human mate choice traits: Impli-
cations for evolutionary psychologists. Journal
of Evolutionary Psychology, 9(3), 219–247.
Kock, N., & DeLuca, D. (2007). Improving busi-
ness processes electronically: An action
research study in New Zealand and the U.S.
Journal of Global Information Technology
Management, 10(3), 6–27.
Kock, N., & Moqbel, M. (2016). A six-stage frame-
work for evolutionary IS research using path
models: Conceptual development and a social
networking illustration. Journal of Systems
and Information Technology, 18(1), 64–88.
Kock, N., Carmona, J., & Moqbel, M. (2015).
Media naturalness and compensatory adap-
tation: Counterintuitive effects on correct
rejections of deceitful contract clauses. IEEE
Transactions on Professional Communication,
58(4), 381–395.
Kock, N., Chatelain-Jardón, R., & Carmona, J.
(2008). An experimental study of simulated
 EVOLUTIONARY PSYCHOLOGY AND COMMUNICATION
web-based threats and their impact on
knowledge communication effectiveness.
IEEE Transactions on Professional Communi-
cation, 51(2), 183–197.
Kock, N., Verville, J., & Garza, V. (2007). Media
naturalness and online learning: Findings sup-
porting both the significant- and no-­significant-
difference perspectives. Decision Sciences
Journal of Innovative Education, 5(2),
333–356.
Kokko, H., Brooks, R., McNamara, J.M., & Hou-
ston, A.I. (2002). The sexual selection con-
tinuum. Proceedings of the Royal Society of
London: Biological Sciences, 269(1498),
1331–1340.
Laitman, J.T. (1984). The anatomy of human
speech. Natural History, 20(7), 20–27.
Laitman, J.T., & Reidenberg, J.S. (1997). The
human aerodigestive tract and gastroesopha-
geal reflux: An evolutionary perspective. The
American Journal of Medicine, 103(5), 2S–8S.
Lee, H.S., & Holyoak, K.J. (2008). The role of
causal models in analogical inference. Jour-
nal of Experimental Psychology: Learning,
Memory, and Cognition, 34(5), 1111–1122.
Lieberman, P. (1998). Eve spoke: Human lan-
guage and human evolution. New York, NY:
W.W. Norton & Company.
Luger, G.F., & Stubblefield, W.A. (2008). AI algo-
rithms, data structures, and idioms in Prolog,
Lisp, and Java for artificial intelligence: Struc-
tures and strategies for complex problem
solving. Reading, MA: Addison-Wesley.
Manipady, S., Menezes, R.G., & Bastia, B.K.
(2006). Death by attack from a wild boar.
Journal of Clinical Forensic Medicine, 13(2),
89–91.
Maynard Smith, J. (1998). Evolutionary genet-
ics. New York, NY: Oxford University Press.
Maynard Smith, J., & Harper, D. (2003). Animal
signals. New York, NY: Oxford University Press.
Mayr, E. (1976). Evolution and the diversity of
life. Cambridge, MA: Harvard University Press.
McElreath, R., & Boyd, R. (2007). Mathematical
models of social evolution: A guide for the
perplexed. Chicago, IL: The University of
Chicago Press.
McQueen, R.J., Payner, K., & Kock, N. (1999).
Contribution by participants in face-to-face
business meetings: Implications for collabo-
rative technology. Journal of Systems and
Information Technology, 3(1), 15–33.
433
Miller, G.F. (2000). The mating mind: How
sexual choice shaped the evolution of human
nature. New York, NY: Doubleday.
Miller, G.F. (2002). How did language evolve?
In: H. Swain (Ed.), Big questions in science
(pp. 79–90). London, England: Jonathan Cape.
Petrie, M., Halliday, T., & Sanders, C. (1991).
Peahens prefer peacocks with elaborate
trains. Animal Behaviour, 41(2), 323–331.
Pinker, S. (2003). Language as an adaptation to
the cognitive niche. In M. Christiansen & S.
Kirby (Eds.), Language evolution: States of
the Art (pp. 16–37). New York, NY: Oxford
University Press.
Price, G.R. (1970). Selection and covariance.
Nature, 227(1), 520–521.
Rantanen, T.K., & Salo, J.A. (1999). Gastroeso­
phageal reflux disease as a cause of death:
Analysis of fatal cases under conservative
treatment. Scandinavian Journal of Gastro-
enterology, 34(3), 229–233.
Rice, S.H. (2004). Evolutionary theory: Mathe-
matical and conceptual foundations. Sun-
derland, MA: Sinauer Associates.
Russel, S., & Norvig, P. (2002). Artificial intelli-
gence: A modern approach. Upper Saddle
River, NJ: Prentice Hall.
Short, J.A., Williams, E., & Christie, B. (1976).
The social psychology of telecommunica-
tions. London, England: John Wiley & Sons.
Simon, A.F. (2006). Computer-mediated com-
munication: Task performance and satisfac-
tion. Journal of Social Psychology, 146(3),
349–379.
Trivers, R. (2002). Natural selection and social
theory. Oxford, England: Oxford University
Press.
Ulijn, J.M., Lincke, A., & Karakaya, Y. (2001).
Non-face-to-face international business
communication: How is national culture
reflected in this medium? IEEE Transactions
on Professional Communication, 44(2),
126–138.
Vandenplas, Y., Goyvaerts, H., Helven, R., &
Sacre, L. (1991). Gastroesophageal reflux, as
measured by 24-hour pH monitoring, in 509
healthy infants screened for risk of sudden
infant death syndrome. Pediatrics, 88(4),
834–840.
Wainfan, L., & Davis, P.K. (2004). Challenges in
virtual collaboration: Videoconferencing,
audioconferencing and computer-mediated
434 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
communications. Santa Monica, CA: RAND
Corporation.
Waldmann, M.R., Holyoak, K.J., & Fratianne, A.
(1995). Causal models and the acquisition of
category structure. Journal of Experimental
Psychology: General, 124(2), 181–206.
Walker, T. (2008). Could sexual selection have
made us psychological altruists? Studies in
History and Philosophy of Science Part C:
Studies in History and Philosophy of Biological
and Biomedical Sciences, 39(1), 153–162.
Warkentin, M.E., Sayeed, L., & Hightower, R.
(1997). Virtual teams versus face-to-face
teams: An exploratory study of a web-based
conferencing system. Decision Sciences,
28(4), 975–996.
Wilder, S.M., & Rypstra, A.L. (2008). Sexual size
dimorphism mediates the occurrence of
state-dependent sexual cannibalism in a
wolf spider. Animal Behaviour, 76(2),
447–454.
Wilson, E.O. (2000). Sociobiology: The new
synthesis. Cambridge, MA: Harvard Univer-
sity Press.
Wright, S. (1934). The method of path coeffi-
cients. The Annals of Mathematical Statis-
tics, 5(3), 161–215.
Wright, S. (1960). Path coefficients and path
regressions: Alternative or complementary
concepts? Biometrics, 16(2), 189–202.
Zahavi, A. (1975). Mate selection – A selection
for a handicap. Journal of Theoretical Biol-
ogy, 53(1), 205–214.
Zahavi, A., & Zahavi, A. (1997). The Handicap
Principle: A missing piece of Darwin’s puzzle.
Oxford, England: Oxford University Press.

Evolutionary Psychology and
Climate Change: Understanding
the Impact of Time Perspective
on Carbon Emissions across
75 Countries
INTRODUCTION
For the last 40 years, Darwinian approaches to
human behavior have been successfully applied
to understanding a diverse range of behaviors
operating at both individual and group levels
(Buss and Shackelford, 1997; Smith, 2000;
Tooby, 2018; Tooby and Cosmides, 1988).
Despite these successes, the leveraging of these
approaches to contextualize and address global
issues has been limited (Caudell and Quinlan,
2016; but see Sörqvist and Langeborg, 2019).
This is unfortunate given that many of the
behaviors driving global issues, including
resource consumption dynamics, fertility pat-
terns, and cooperation levels, can be partially
explained as a consequence of human evolu-
tionary history. A global issue amenable to
explanation by an evolutionary perspective is
anthropogenic climate change. Climate change
studies have increasingly emphasized the role
of demographic patterns, including fertility
rates, age structure, and household size, in
shaping the emission of fossil fuels (Cohen,
22
Mark A. Caudell
2010; Jorgenson and Clark, 2013; Murtaugh
and Schlax, 2009; O’Neill et  al., 2010; Rosa
and Dietz, 2012; Shi, 2003; Stephenson et al.,
2010). Indeed, population growth in the next
half-century, even when holding current per
capita emissions constant, could raise global
emissions by half (Swim et al., 2010).
Caudell and Quinlan (2016) sought to
understand anthropogenic climate change
through an approach that combined elements
of life history theory (LHT) and evolution-
ary psychology. This approach was used to
understand the ‘Population Paradox’ under-
lying anthropogenic climate change and to
propose potential resolutions. The paradox
is founded on data showing that per capita
carbon emissions (i.e., energy consumption)
are positively associated with both fertil-
ity rate increases and decreases (Shi, 2003;
Stephenson et al., 2010). This pattern contra-
dicts LHT given that per capita energy con-
sumption is generally negatively associated
with fertility in mammals (Stearns, 1992).
To explicitly frame this paradox within a
436 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
life history perspective, Caudell and Quinlan
drew upon the slow-to-fast life history con-
tinuum (Ellis et al., 2009). Specifically, they
argued that in high-income countries (HICs),
selection for slower life history strategies
results in high parental investment that tar-
gets ‘dynastic fitness’ (Kaplan, 1996). These
investment patterns result in large per capita
energy investments in skills, knowledge, and
other abilities that are necessary to compete
in industrial labor markets (Burger et  al.,
2011). Given that industry and supporting
infrastructure (e.g., electricity, transportation)
are responsible for over 60% of carbon emis-
sions (Boden et al., 2017), these investment
patterns are likely to promote increased emis-
sions. In addition, these investment patterns,
particularly in the context of HICs, are also
resource-intensive, thereby reducing Earth’s
carrying capacity. Through limiting carrying
capacities and promoting carbon emissions
driving climate change, these slower life his-
tory strategies also increase levels of envi-
ronmental risk globally. Higher risk levels
in non-industrial areas (e.g., many low- and
middle-income countries (LMICs)) promote
selection for faster life history strategies that
increase fertility rates and ultimately create
larger per capita carbon legacies (i.e., the
expected future emissions produced by a par-
ent and their surviving offspring; Murtaugh
and Schlax, 2009). Viewed globally, these
two life history strategies give rise to invest-
ment patterns in HICs that are decreasing
Earth’s carrying capacity and fertility pat-
terns in LMICs that increase the rate at which
this capacity is reached. To resolve the posi-
tive association between carbon emissions
and slow and fast life histories, Caudell and
Quinlan (2016) propose that the evolved psy-
chological construct of time orientation can
motivate decreased fertility rates and be co-
opted to promote pro-environmental behav-
ior through investment in education, thus
resulting in reductions in carbon emissions.
In this chapter, I expand upon the origi-
nal study by Caudell and Quinlan (2016) to
include data from around 100,000 women
across 75 countries. This analysis provides a
more robust assessment of the proposed rela-
tionships among time orientation, fertility
and investment patterns, and global carbon
emissions. The analysis supports that role
of time orientation in impacting life history
strategies and carbon emissions. More gener-
ally, the analysis supports the application of
Darwinian perspectives on human behavior
to understand and address the demographic
and consumption patterns fueling anthropo-
genic climate change.
BACKGROUND
Life History, Risk and
Reproduction
In LHT, reproductive behavior is guided by
the costs and benefits of investing energy into
the ‘competing’ life functions of growth,
maintenance, repair, and reproduction (Roff,
2002; Stearns, 1992). Given that resources
invested into one life function cannot be
devoted to another, a set of trade-offs arise
(Stearns, 1989). Simply, time and calories
invested into attracting a mate are time and
calories that cannot be invested in caring for
offspring. A fundamental trade-off in every
organism’s life history exists between current
and future reproduction. That is, should energy
be devoted towards producing offspring (i.e.,
reproductive effort) as soon as physiologically
viable? Or should investment in reproductive
effort be delayed in favor of somatic invest-
ment, thereby increasing the quality of the
organism to provide parental care?
The trade-off between current and future
reproduction underlies the continuum between
slow and fast life history strategies (Stearns,
1992). Slower life history strategies delay
reproduction thereby reducing the quantity
of potential offspring, but allow the organ-
ism to allocate more resources to growth,
maintenance, and repair (i.e., somatic effort).
Extended investment in somatic effort should
 EVOLUTIONARY PSYCHOLOGY AND CLIMATE CHANGE
lengthen life expectancy and, through enabling
increased parental investment, result in higher-
quality offspring. Higher-quality offspring are
important in organisms with slower life his-
tories given competition for resources. This
competition emerges as slower life histories
are found in stable and predictable environ-
ments that allow species to trend towards the
carrying capacity of a particular niche, thereby
intensifying competition for resources. Given
this selection, species with slower life histo-
ries are often larger, have fewer offspring but
extended parental investment periods, and
longer life expectancies (Stearns, 1989).
Fast life history strategies are more
likely to emerge in unstable and unpredict-
able environments that, in turn, are typically
less crowded. This unpredictability makes
it unlikely that adaptations will evolve to
enhance the competitive abilities of organ-
isms, given that changing environments may
render these abilities moot in the future. These
unpredictable environments also decrease the
probability of surviving to adulthood. The
consequences of these environments, in the
logic of life history, mean there is little fitness
advantage in investing heavily in oneself or
one’s offspring, as own and offspring survi-
vorship is low. Instead, the fitness advantage
lies in earlier and more frequent investment
in mating effort, which increases the quan-
tity of offspring but decreases the quality of
offspring. This investment pattern, combined
with unpredictable environments, curtails
life expectancy of both parents and offspring
(Low, 1978; Promislow and Harvey, 1990;
Roff, 2002; Stearns, 1992; Trivers, 1972).
As stated above, the continuum of slow-
to-fast life histories is founded upon envi-
ronmental unpredictability. Within LHT, this
unpredictability is framed as environment
risk, which is further divided into intrin-
sic risk and extrinsic risk. Intrinsic risk is
mortality risk that can be impacted through
investing in oneself or one’s offspring. When
intrinsic risk dominates, selection for slower
strategies is strong as increased somatic
investment can produce a fitness advantage.
437
In contrast, mortality from extrinsic risk
should occur independent of investment in
reproduction (mating, parental investment)
as it results from largely unpredictable
sources (e.g., predation). As unpredictable,
an organism is unable to change the survival
probabilities associated with extrinsic risks,
both for itself and its offspring, through
increased investment. A high extrinsic risk
environment favors faster strategies to ‘beat
the odds’ that a potential parent dies before
reproduction or leaves no offspring, given
high juvenile mortality rates (Quinlan, 2007;
Roff, 2002; Stearns, 1992). Selection for all
life history strategies, both across and within
species, depends largely on levels of extrinsic
risk in the local environment (Charlesworth,
1994; Quinlan, 2007; Stearns, 1989; Williams,
1957). Support for a relationship between
extrinsic risk and life history strategies has
been found across a diverse range of human
societies (Bulled and Sosis, 2010; Caudell and
Quinlan, 2012; Gant et al., 2009; Low et al.,
2008; Nettle, 2010; Walker et al., 2006; Wilson
and Daly, 1997), although other economic and
cultural considerations clearly impact fertil-
ity decisions as well (Caudell, 2015), while
the rapid pace of extrinsic risk changes (e.g.,
infant mortality interventions in LMICs (Liu
et  al., 2016)) could mask the relationship
between extrinsic risk and fertility.
Extrinsic Risk and Evolutionary
Psychology
The phenotypic effects of extrinsic risk
should be apparent in a suite of evolved psy-
chological, and within humans, cultural,
traits that motivate behavior towards invest-
ment in slow or fast life history strategies
(Hill et  al., 1997; Kruger et  al., 2008;
Quinlan, 2010). For example, mate-choice
decisions are impacted by predation risk
within the local environment (Jennions and
Petrie, 1997). Time orientation is another
aspect of psychology, particularly human
psychology, that may function as a
438 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
mechanism that responds to risk during an
organism’s development (Hill et  al., 1997;
Kruger et  al., 2008). Time orientation, as a
psychological dimension, is the degree to
which the past, present, or future guides atti-
tudes and behaviors (Zimbardo and Boyd,
1999). Present-oriented individuals tend to
be influenced by the sensory qualities of the
current environment, seek immediate gratifi-
cation, and are more likely to exhibit risky
attitudes and behaviors (e.g., drugs, unpro-
tected sex) (Apostolidis et  al., 2006; Boyd
and Zimbardo, 2005; Caldwell et  al., 2006;
Henson et  al., 2006; Kruger et  al., 2008;
Rothspan and Read, 1996; Schechter and
Francis, 2010). Future orientation allows one
to ‘transcend’ stimuli in the current environ-
ment and delay gratification, particularly in
the service of future goals (e.g., education)
(Horstmanshof and Zimitat, 2007; Peetsma,
2000; Zimbardo and Boyd, 1999).
Couched within a theoretical framework
combining LHT and evolutionary psychology,
present orientations may be adaptive in envi-
ronments with high extrinsic risk given that
future investments, such as education, may
not increase fitness due to decreased survivor-
ship to adulthood. Future orientations may be
adaptive in low extrinsic risk environments
where future investments, such as education,
may increase competitive advantage, thereby
increasing fitness. In the context of life his-
tory strategies, present orientations should
be associated with traits of a faster strategy,
including limited somatic/offspring invest-
ment, higher total fertility rates, and decreased
lifespans, whereas future orientation should be
associated with slower strategies and traits that
include later onset of reproduction, fewer off-
spring, and longer lifespans.
LHT, Evolutionary Psychology,
and Climate Change
But what can these various evolved strategies
tell us about the factors promoting climate
change in the present? Indeed, should either
slower or faster strategies be considered ben-
eficial in the reduction of carbon emissions?
Here, we are confronted with the ‘Population
Paradox’, that carbon emissions are posi-
tively associated with fertility rate increases
and decreases (Shi, 2003; Stephenson et al.,
2010). In a life history framework, then, both
slow and fast life history strategies are asso-
ciated with increased emissions. However,
the route through which this promotion
occurs differs and can be framed, as with life
history relationships, as a trade-off between
current and future emissions.
Slower life history strategies result in
larger and longer energy investments per
individual, both for parents and offspring, and
so higher per capita carbon emissions in the
present. For example, competition for mates
in many cultures results in ‘conspicuous con-
sumption’ by males (i.e., large investments in
extrasomatic resources such as houses, cars,
livestock, etc.), which is positively related
to carbon emissions. In addition, the longer
lifespans and later ages at first marriage
characterizing slower strategies also result in
smaller average household sizes, including
more single adult households, which further
increases per capita emissions (Jorgenson
and Clark, 2013; O’Neill et al., 2012).
Faster strategies result in lower per capita
emissions in the present due to higher extrin-
sic risk, which decreases competition pres-
sures. Reductions in competition pressures,
along with high extrinsic risks, mean that the
benefits of somatic and extrasomatic invest-
ments in oneself and one’s offspring may not
be realized. Importantly, however, the larger
quantity of potential offspring increases an
individual’s carbon legacy, again the expected
future emissions produced by a parent and
their surviving offspring. Carbon legacies
have a much greater potential to impact emis-
sions when compared to emission changes
associated with changes in current investment
patterns. In the United States, for example,
every child contributes 9,441 metric tons to
their mother’s carbon legacy while a mother
who recycled across her lifetime would save
 EVOLUTIONARY PSYCHOLOGY AND CLIMATE CHANGE
about 17 tons (Murtaugh and Schlax, 2009).
Whether selection for slow-to-fast life history
strategies results in greater emissions, both in
the present and future, is thus a function of
per capita emissions, population size, and the
rates at which these two factors rise or fall
(Crowley, 2000; Dietz and Rosa, 1997).
On the spectrum of human life history
variation, populations that comparatively
trend towards exhibiting slower strategies
have substantially larger carbon legacies
than those countries exhibiting faster strate-
gies. This result is largely due to the fact that
slower strategies are more common in HICs –
those that emit the most carbon – whereas
faster strategies are more common in LMICs,
with lower overall carbon emissions (Boden
et al., 2017). Disparities in carbon emissions
between HICs and LMICs are so consider-
able that they negate the potentially greater
carbon legacies associated with high-fertility
strategies. A woman in Bangladesh, for
example, would need to have 74 children
before her carbon legacy equaled that of an
American woman with one child (Murtaugh
and Schlax, 2009).
As argued above, carbon emission dis-
parities between HICs and LMICs can be
viewed as byproducts of low extrinsic risk
environments, which increase competition
and allow for the benefits of larger invest-
ments to be accrued in the future (Kaplan,
1996). Conceptualized historically, these
low-risk environments permitted the emer-
gence and development of industrializa-
tion, which increased the need for sustained
parental investment through emphasizing
education and other extrasomatic resources.
Concurrently, these increases in industri-
alization also decreased environment risk,
through advances in healthcare, sanitation,
and technology (Bar and Leukhina, 2010;
Boserup et al., 1983; Boucekkine et al., 2007;
Caldwell, 1976; Coale, 1984; Galor and
Weil, 2000). As the fitness benefits of more
future-oriented slower strategies increased,
including future orientation, so too did the
per capita emissions of individuals. If this
439
argument is correct, a positive correlation
should exist between CO2 emissions and
future orientation.
LHT and Evolutionary Psychology:
Resolving the Population Paradox
The observation that population increases,
and decreases, elevate carbon emissions
complicates the development of solutions to
anthropogenic climate change. Indeed,
efforts to decrease extrinsic risk levels in
LMICs would, theoretically, reduce the
demographic contributions towards emis-
sions by promoting investment in slower
strategies. Reductions in demographically
linked emissions, however, would likely be
offset by subsequent increases in per capita
investments in potential parents and off-
spring. Critically, these consequences ensure
that fertility reductions by themselves will
not be sufficient in decreasing carbon emis-
sions (see discussion in Cohen, 2010).
In light of the observations above, reduc-
tions in emissions will come through chan-
neling somatic and parental investments
towards pro-environmental investment. To this
end, I argue that the life history component of
future orientation can be co-opted to produce
pro-environmental investment patterns. This
co-opting can occur when somatic investment
in potential parents and parental investment
in offspring is directed towards investment in
education, particularly higher education. As
discussed further below, this argument rests on
evidence indicating that education promotes
future orientation and that future orientations,
in some contexts, promote pro-environmental
behaviors.
Education levels are positively corre-
lated with future orientation, or related con-
structs (e.g., delay discounting, patience,
risk-taking), across cultures, development
contexts, and subsistence types, including
populations in the United States (Alan and
Ertac, 2014; Jaroni et al., 2004; Oreopoulos
and Salvanes, 2011; Van der Pol, 2011),
440 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Denmark (Harrison et  al., 2002), Turkey
(Alan and Ertac, 2014), Mexico (Perez-Arce,
2011), Vietnam (Anderson et al., 2004), rural
populations in India (Bauer and Chytilová,
2013), and Uganda (Bauer and Chytilová,
2010), and in small-scale societies in Bolivia
(Kirby et  al., 2002). Longitudinal research
(Alan and Ertac, 2014; Perez-Arce, 2011)
and studies using exogenous sources of vari-
ation in education to infer causation (e.g., dif-
ferential access to education across villages,
changes in compulsory-schooling laws) have
shown that education can promote increases
in future time orientation, delay discounting,
and lower the propensity towards risk-taking
(Bauer and Chytilová, 2010; Lochner and
Moretti, 2004). Further, education promotes
future time orientation across a range of risk
contexts, as demonstrated in studies finding
that education has the largest impact on time
orientation in disadvantaged youth, a finding
consistent with recent results suggesting that
education has the largest impact on fertil-
ity within high-risk environments (Caudell,
2015).
Although education can produce a more
future-oriented psychology, future orientation
also is associated with pro-environmental atti-
tudes and behaviors, both across individuals
and cultures (Arnocky et al., 2012; Carmi and
Arnon, 2014; Joireman et al., 2004; Lindsay
and Strathman, 1997; Milfont and Gouveia,
2006; Toepoel, 2010), although others have
not found a link (Ebreo and Vining, 2001).
Future orientation is an essential aspect of
sustainable and pro-environmental behav-
ior as it emphasizes changing one’s behav-
ior in the present to take responsibility for
future outcomes (Carmi and Arnon, 2014).
Important for the current argument, how-
ever, is that the association between future
orientation and pro-environmental behavior
is largely confined to highly educated indi-
viduals, particularly those with a college
education. This proposed link is consist-
ent with results showing that environmental
concern and behavior are related to educa-
tion level, but only at higher post-secondary
levels (Franson and Garling, 1999; Johnson
et  al., 2004; Olli et  al., 2001; Wall, 1995;
but see Wesely Schultz, 2001). Considering
the research above, the prediction can be
extended that an individual’s and country’s
investment in higher education will mediate
the effect of future orientation on per capita
emissions through impacting both demo-
graphic and consumption patterns.
Predictions
The analysis that follows assesses the following
predictions:
P1: Present orientations should be associated with
traits of faster life history strategies whereas
future orientations should be associated with
slower strategies.
P2: A positive correlation should exist between a
country’s CO2 emissions and future orientation.
P3: Investment in higher education will mediate the
effect of future orientation on per capita emis-
sions through impacting both demographic and
consumption patterns.
Methods
Predications were tested through multilevel
analysis of data provided by 95,752 women
from 75 countries (Model 1) and linear
regression analysis of national-level data
from these same 75 countries (Model 2).
Model specifications and data sources are
discussed below. Stata 16 was used for analy-
sis (StataCorp, 2019).
Model 1: Measures and data
sources
Total fertility rates were used as proxies to
reflect the slow-to-fast LHT continuum.
Rates were collected for women 55 years and
older, who have largely completed fertility.
Extrinsic risk levels were represented by
taking the average life expectancy at birth
(LEB) in the 10 years after a person was
born. Education level (EDU) was recoded
 EVOLUTIONARY PSYCHOLOGY AND CLIMATE CHANGE
Table 22.1  Descriptive statistics for Model 1
Variable Mean
Total fertility rate (TFR) 3.75
Education level (EDU) 1.25
Future orientation (FO) 45.54
Life expectancy at birth (LEB) 64.67
N = 95,752. Std dev. = standard deviation. Education level is coded as 0 = None, 1 = Primary, 2 = Secondary, 3 = Post-secondary.
into none, primary, secondary, and post-­
secondary. A country’s time orientation was
quantified by Hoftstede’s (2001) Long-Term
Short-term Orientation measure. Long-Term
Orientation represents the ‘fostering of virtues
oriented towards future rewards’, whereas
‘Short Term Orientation stands for the foster-
ing of virtues related to the past and present’
(Hofstede, 2001: 359). Given the scale
anchors, 0 (most present) to 100 (most future),
it will be referred to as future orientation. The
measure has been a source of some debate
(see Ashkanasy et al., 2004; Fang, 2003) but
scores have been validated through factor
analysis of time orientation items in the World,
African, and European Values Surveys
(Minkov, 2007; Minkov and Hofstede, 2011).
In addition, studies have found that score vari-
ations are associated with behaviors impacted
by time orientation, including sales of life
insurance, across cultures (Park and Lemaire,
2011). Future orientation scores were calcu-
lated for 75 countries to reflect a geographi-
cally and economically diverse sample (e.g.,
different continents, high- versus low-income
countries). See Table 22.1 for descriptive sta-
tistics and Table 22.2 for Pearson correlation
coefficients between Model 1 variables.
Table 22.2  Variable correlations for Model 1
Variable Total fertility Education
Total fertility -
Education −0.48** -
Future orientation −0.36** 0.34*
Life expectancy −0.39** 0.53**
**
p<.01. *p<.05
N = 95,752. Education level is coded as 0 = None, 1 = Primary, 2 = Secondary, 3 = Post-secondary.
441
Std dev. Min Max
2.85 0.00 20.00
1.00 0.00 3.00
24.88 0.00 100.00
10.98 27.08 82.12
Data for Model 1 were collected from the
World Values Survey (www.worldvalues-
survey.org/; WVS, n.d.), European Values
Survey (www.europeanvaluesstudy.eu/; EVS,
n.d.), and surveys available at the Integrated
Public Use Microdata Series-International
from the Minnesota Population Center,
including the Demographic Health Surveys
(www.ipums.org/; IPUMS-DHS, n.d.). Data
were collected from years closest to the date
the future orientation was measured in the
country. Not every country had data from the
year the future orientation measure was col-
lected but all data used in the analysis were
plus or minus five years from this date. As
stated above, education level (EDU) was
coded into four categories: ‘none’, ‘primary’,
‘secondary’, and ‘post-secondary’. Some
surveys recorded an individual’s education in
years, and these were recoded using the fol-
lowing criteria: 0–2 years = ‘none’, 3–5 years
= ‘primary’, 6–12 years = ‘secondary’, 12+
years = ‘post-secondary’. Life expectancy at
birth (LEB) is defined as the number of years
of life remaining at birth. Average LEB in
the 10 years after an individual’s birth was
calculated from data collected at the World
Bank (http://data.worldbank.org/) (World
Future orientation Life expectancy
-
0.42** -
442 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Bank, 2009). For Hofstede’s time-orientation
measure, 72 of the 75 countries had specific
future orientation scores whereas three coun-
tries were assigned regional scores. These
countries were Ethiopia and Kenya, which
were assigned to the East Africa region, and
Sierra Leone, which was assigned to the West
Africa region. Table 22.3 presents the coun-
tries examined in the analysis, the associated
future orientation scores, databases from
which the fertility and education data were
taken, and associated years of collection.
Model 1: Specification
Across the 75 countries in Model 1, the
number of women ranged from 800 to 1,575
per country, with an average cluster size of
≈910 women. A multilevel modeling
approach was appropriate because fertility
behavior within the same country is more
likely to be correlated than fertility behavior
between countries due to shared sociocul-
tural, biological, and physical environments.
As such, within-country observations on fer-
tility lack independence. To compensate for
this non-independence, multilevel models
partition the variance in an outcome measure
into variance accounted for by within- and
between-group levels. This partitioning also
helps to prevent study conclusions marred by
the ecological fallacy, which occurs when
inferences about behaviors of individuals are
based solely upon aggregate measures of the
group to which those individuals belong
(Hox, 2010). Using aggregate data measures
(e.g., national LEB) to infer information
about behaviors at the individual level can be
a serious problem in the cross-cultural study
of life history as the majority of studies
(Bulled and Sosis, 2010; Caudell and
Quinlan, 2012; Low et  al., 2008) rely on
these aggregate measures.
To justify the use of a multilevel mod-
eling approach, a random intercept model
(sometimes referred to as ‘baseline’, ‘null’,
or ‘unconditional’ model) was initially
specified to calculate the intraclass correla-
tion (ICC). The ICC is calculated through
partitioning variance in total fertility rate
(TFR) into variance existing at the within-
and between-country levels. The ICC was .38
indicating that 38% of the variance in indi-
vidual TFRs is associated with countries (i.e.,
between-group), whereas 62% is associated
with individuals (i.e., within-group). The
ICC can further be interpreted as the extent
to which individuals in the same country are
more alike compared with individuals of a
different country (Peugh, 2010). In this case,
the TFRs within the same country are 38%
more similar relative to TFRs within a differ-
ent country. Given these ICCs, there is strong
evidence that total TFRs are not independent
within countries and so a multilevel approach
is justified.
Model specification proceeded by adding
level-one predictors (i.e., EDU, LEB) as fixed
effects. Fixed effects ensure the regression
coefficient of TFR on the level-one predictors
could not vary across countries. Next, random
effects were specified to allow the regression
coefficient of TFR on level-one predictors
to vary across countries. The specification
of random effects is necessary as the effect
of education and LEB on TFR is hypoth-
esized to vary across countries, in part due
to interactions within a culture’s future ori-
entation. In these models, education and LEB
were group-mean centered (national educa-
tion level and average LEB 10 years after
birth), the suggested centering technique to
ensure correct interpretation of the level-one
effects in multilevel models (Peugh, 2010).
Following the inclusion of level-one predic-
tors, the level-two predictor of future orienta-
tion (FO) was regressed on the random slope
as predicted by the regression of TFR on
education and LEB. Importantly, regression
of FO on the random slope allowed examina-
tion of whether FO had a significant effect on
the relationship between education and TFR
and LEB and TFR. Finally, cross-level inter-
action effects were specified between FO and
EDU and LEB. As a level-two effect, FO was
grand-mean centered using the global mean
of FO.
 EVOLUTIONARY PSYCHOLOGY AND CLIMATE CHANGE 443

Table 22.3  Data sources, years, and future orientation scores

Country Year(s) Database FO Country Year(s) Database FO

Albania 2008 IPUMS 61 Kyrgyzstan 1999 IPUMS 66

Algeria 2002 WVS 26 Latvia 1996 WVS 69
Argentina 2001 IPUMS 20 Lithuania 1997 WVS 82
Armenia 2005 IPUMS 61 Luxembourg 1999 EVS 64
Australia 2005 WVS 21 Malaysia 2006 WVS 41
Austria 2001 EVS 60 Mali 1998 IPUMS 20
Azerbaijan 2006 IPUMS 61 Malta 1999 EVS 47
Bangladesh 2007 IPUMS 47 Mexico 1996 WVS 24
Belarus 1999 WVS 81 Morocco 2003 IPUMS 14
Belgium 1999 EVS 82 Netherlands 2006 WVS 67
Bosnia and Herzegovina 1998 WVS 70 New Zealand 2004 WVS 33
Brazil 2000 WVS 44 Nigeria 2008 IPUMS 13
Bulgaria 1999 WVS 69 Pakistan 2007 IPUMS 50
Burkina Faso 2003 IPUMS 27 Peru 2000 IPUMS 25
Canada 2000 WVS 36 Philippines 2003 IPUMS 27
Chile 2002 WVS 31 Poland 2005 WVS 38
China 1990 IPUMS 87 Portugal 1999 EVS 28
Colombia 2005 IPUMS 13 Puerto Rico 1990 IPUMS 0
Croatia 1999 EVS 58 Republic of Korea 1990 WVS 100
Czech Republic 1999 WVS 70 Republic of Moldova 2005 WVS 71
Denmark 1999 EVS 35 Romania 2002 WVS 52
Dominican Republic 2007 IPUMS 13 Russian Federation 1999 WVS 81
Egypt 2005 IPUMS 7 Rwanda 2005 IPUMS 18
El Salvador 1999 WVS 20
Saudi Arabia 2003 WVS 36
Estonia 1996 WVS 82
Serbia 2006 WVS 52
Ethiopia 1997 WVS 32
Sierra Leone 2008 IPUMS 9
Finland 1996 WVS 38
Slovakia 1999 EVS 77
France 2006 WVS 63
Slovenia 2002 IPUMS 49
Georgia 2008 WVS 38
South Africa 1998 IPUMS 34
Germany 2006 WVS 83
Spain 2007 WVS 48
Ghana 2003 WVS 4
Sweden 1999 EVS 53
Greece 2001 EVS 45
Switzerland 2000 EVS 74
Hungary 2001 IPUMS 58
Taiwan 2006 WVS 93
Iceland 1999 EVS 28
Turkey 2003 IPUMS 46
India 2006 IPUMS 51
Uganda 2000 IPUMS 24
Indonesia 2007 IPUMS 62
Ukraine 2007 IPUMS 86
Iran (Islamic Republic of) 2006 WVS 14
United Kingdom 1998 WVS 51
Iraq 1997 WVS 25
United Republic of Tanzania 2004 IPUMS 34
Ireland 1999 EVS 24
United States of America 1999 WVS 26
Israel 1995 WVS 38
Venezuela 2001 IPUMS 16
Italy 2005 WVS 61
Vietnam 2002 IPUMS 57
Japan 2005 WVS 88
Jordan 2002 WVS 16 Zambia 2007 IPUMS 30
Kenya 2009 IPUMS 32 Zimbabwe 2005 IPUMS 15

EVS = European Values Survey. IPUMS = Integrated Public Use Microdata Series-International from the Minnesota Population
Center. WVS = World Values Survey. FO = future orientation score.
444 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Several steps were taken to ensure adequacy
of model fit and interpretation. Year of data
collection was entered as a control variable,
given the general decline in fertility rates
across time. Year did not have a significant
impact on TFR, after controlling for EDU,
LEB, and FO. Models also included the
control variable Gross National Income per
capita (GNI per capita) (taken from World
Bank data; http://data.worldbank.org/), given
large correlations between education and
income that could produce a spurious effect
of education on TFR. As a country-level
effect, GNI per capita did not have a signifi-
cant effect on the relationship between LEB
and TFR or EDU and TFR. It is possible that
entering an individual’s income may have
impacted the results, although the inclusion
of wealth was not permitted given that it was
gathered differently across the surveys. To
determine whether a model provided a sig-
nificantly better fit (i.e., p < .05) compared to
the previous model (e.g., fixed-effect model
versus random-effect model), likelihood-
ratio tests were conducted (Rabe-Hesketh
and Skrondal, 2012). All models provided a
significantly better fit compared to the previ-
ously specified model.
Model 2: Measures and data
sources
Data included in country-level models were
secured to test the I=P.A.T equation. The
I=P.A.T equation treats environmental impact
(I), here carbon emissions per capita, as due
to three factors: population (P), affluence (A),
and technology (T) (Dietz and Rosa, 1997;
Ehrlich and Holdren, 1971). Environmental
impact is an outcome variable representing
carbon emissions. Carbon emissions per
capita are expressed as metric tons and are
emissions from the burning of fossil fuels
and the manufacture of cement and include
CO2 produced during consumption of solid,
liquid, and gas fuel, and gas flaring (i.e.,
controlled burning of natural gas during
exploration, production, and processing)
(World Bank, 2016). Consistent with other
studies (Shi, 2003), total population is used
to represent P, GDP per capita is used to rep-
resent A, and percentage of GDP from manu-
facturing and percentage from services is
used to represent T. Total populations are
midyear estimates. GDP per capita is gross
domestic product in current US dollars
divided by midyear population. The percent-
age of GDP from manufacturing is value
added, meaning it is ‘the net output of a
sector after adding up all outputs and sub-
tracting intermediate inputs. It is calculated
without making deductions for depreciation
of fabricated assets or depletion and degrada-
tion of natural resources’ (World Bank, n.d.).
Educational attainment represents the per-
centage enrollment in post-secondary educa-
tion across both males and females.
Percentage enrollment in post-secondary
education (tertiary education) represents
total enrollment regardless of age. Hofstede’s
(2001) long-term short-term measure was
used to represent a country’s future orienta-
tion. All variables were log transformed prior
to analysis, with the result that coefficients
are interpreted as percentage increases in
carbon emissions per capita given a 1%
increase in the independent variable (e.g.,
population size) (Dietz and Rosa, 1997). Due
to differences in collection years for the time
orientation measure, the year in which the
population, affluence, technology, and impact
variables were collected was matched to the
year the future orientation measure was col-
lected, if available. Model 2 variables were
taken from the World Bank (http://data.
worldbank.org/). Table 22.4 provides descrip-
tive statistics for model variables and Table
22.5 the correlations between these
variables.
Model 2: Specification
Ordinal least squares (OLS) regression was
used to assess how indicators of population,
affluence, and technology interacted with
future orientation to impact carbon emission
per capita (Dietz and Rosa, 1997; Ehrlich
and Holdren, 1971). Regression diagnostics
 EVOLUTIONARY PSYCHOLOGY AND CLIMATE CHANGE 445

Table 22.4  Descriptive statistics for Model 2

Variable Mean Std dev. Min Max

CO2 emissions per capita 5.240 4.613 0.065 20.208

Future orientation 46.434 23.680 7.000 100.000
Post-sec. education 33.531 21.451 0.676 82.439
GDP per capita 12880.650 16526.710 136.630 73270.630
Total population 66300000 188000000 281205 1260000000
% GDP from manufacturing 17.619 6.863 3.549 40.021
% GDP from service sector 56.560 12.996 13.250 80.982

Source: First published in Caudell and Quinlan (2016).

N = 75. Std dev. = standard deviation.

were assessed for the final models. Influential
data points were examined through calculat-
ing Cook’s D and rerunning models with
observations associated with values exceed-
ing 4/N (i.e., 4/75=0.05). Excluding observa-
tions above this threshold did not impact
model interpretation. Multicollinearity was
assessed by calculating variance inflation
factors (VIFs). VIFs for variables of interest
were less than 2, well below the recom-
mended cut-off of 5 (Neter et  al., 1996).
Normality of residuals was tested with the
Shapiro–Wilk Test, kernel density, and stand-
ardized normal probability plots. Results
from all tests indicated only slight deviations
from normality. The homoscedasticity
assumption (i.e., homogeneity of variance of
residuals) was tested using residuals versus
fitted (predicted) plots (Cohen et  al., 2003).
There was evidence of minor heteroscedas-
ticity in the model so a Huber–White sand-
wich estimator was used to provide robust
estimates (Huber, 1967).
RESULTS
Model 1: Time Orientation
Interacts with Fundamental Life
History Relationships
Total fertility rate was 3.71 births when LEB
and EDU were at their within-country mean
and future orientation was at its between-
country mean (see Table 22.6). Controlling

Table 22.5  Variable correlations for Model 2

Variable CO2 FO EDU GDP Pop GDP_Man GDP _Serv

CO2 -
FO 0.24* -
EDU 0.63** 0.33* -
GDP 0.74** 0.18 0.54** -
Population −0.08 0.16 −0.20 −0.12 -
GDP_Man 0.17 0.36* 0.36** 0.08 0.27* -
GDP_Serv 0.56** 0.19 0.58** 0.64** −0.16 0.14 -
**
p<.01. *p<.05
Source: First published in Caudell and Quinlan (2016).
CO2 = CO2 emissions per capita. FO = Future orientation. EDU = Percentage enrolled in post-secondary education.
GDP = GDP per capita. Pop = Total population. GDP_Man = Percentage GDP from manufacturing sector. GDP_Serv = Percentage
GDP from service sector. N = 75.
446 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

Table 22.6  Relationship between total for the other predictors, a one-year increase
fertility rates, time orientation, mortality, in LEB was associated with a 0.09 birth
and parental investment decrease, a one-unit increase in EDU (e.g.,
Total fertility none to primary) was associated with a 0.64
Fixed effects birth decrease, and a one-unit increase in
Education (γ10)a −0.635** future orientation was associated with a 0.06
Life expectancy (γ20)b −0.082** birth decrease. Cross-level interactions
Future orientation (γ01)c −0.059** between FO and EDU and FO and LEB were
EDU x future orientation (γ11) 0.003** significant and are graphed in Figures 22.1
LEB x future orientation (γ21) 0.010** and Figures 22.2 to facilitate interpretation
Intercept (γ00) 3.708** and are discussed further below. The negative
Random effects covariance between the random slope of TFR
Residual (σ2) 4.325** on EDU and intercept variances (τ03 =
Intercept (τ00) 1.351** −0.329) indicates that higher TFRs are asso-
Slope (τ10) LEB 0.012** ciated with a weaker relationship between
Slope (τ20) EDU 0.114**
EDU and TFR. No other covariances were
Covariance (τ01) LEB:EDU 0.011
significant. Significant variance remained
Covariance (τ02) LEB:RI −0.013
around the random slopes of LEB (τ10 =
Covariance (τ03) EDU:RI −.329**
0.012) and EDU (τ10 = 0.114), random inter-
Model summary
cept (τoo = 1.35) and around the grand mean
N (Groups) 95752 (75)
of TFR (σ2 = 4.325) after controlling for
Deviance statistics −95273
EDU, LEB, and FO.
Due to the large sample size (≈100,000), there
Note: a Parameter estimates reflect full information is a high likelihood that some results are signifi-
maximum likelihood robust estimation. b Variable was
cant but do not account for considerable vari-
group-mean centered. c Variable was grand-mean
centered. ** p<.01. *p<.05 ables in TFR. To determine variance accounted
for in the model, several equations for the coef-
ficient of determination were used. Calculation
Total fertility rate

Time orientation
Education level
No education

Figure 22.1  Effect of time orientation on total fertility as a function of education

 EVOLUTIONARY PSYCHOLOGY AND CLIMATE CHANGE 447

Figure 22.2  Effect of time orientation on total fertility as a function of mortality level

of variance accounted for by the final model is Equation 3: Residual variance accounted
complicated as variance exists both within and for between countries by all covariates at
between countries. For Equations 1–3 below, between levels
a = residual variance associated with uncondi-
tional/null model; b = residual variance associ-
R
2
=
( τ 00 ) a − (τ 00 ) b
2 2
= 0.598
(τ 002 ) a
ated with final model; and c = residual variance
from random intercept random slope model
without level-two predictors. Within and across
countries, the covariates accounted for around Model 2: Time Orientation
30% of the variance in TFR, around 14% of the Is Associated with Carbon
variance within countries, and 60% of the vari- Emission Differences
ance between countries.
Equation 1: Residual variance accounted The relationship between future orientation
for by all covariates at both levels and EDU on per capita carbon emissions
after controlling for the impacts of popula-

R
2
=
( σ 2 + τ 002 ) a − (σ 2 + τ 002 ) b tion, affluence, and technology are reported
in Table 22.7. For every 1% increase in future
(σ 2 + τ 002 ) a orientation there was a 0.37% increase in
carbon emissions per capita. A 1% increase
= 0.272 in EDU was associated with a 0.52% increase
in carbon emissions per capita. A significant
Equation 2: Residual variance accounted interaction effect was documented between
for within countries by all covariates EDU and FO on carbon emissions. This

R
2
=
( σ ) a − (σ ) b
2 2
= 0.137
interaction is graphed in Figure 22.3 to facili-
tate interpretation and discussed further
(σ 2 ) a below. GDP per capita was significantly
related to per capita emissions, with a 1%
448 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

Table 22.7  Effect of time orientation and manufacturing sector were not significantly
education on carbon emissions related to carbon emissions within the 75 coun-
Emissions per capita tries examined.
Future orientation 0.373*
Education 0.527**
FO x education −0.337* DISCUSSION
GDP per capita 0.562**
FO x GDP −0.038 This chapter provides a robust analysis of the
Total population 0.058 Population Paradox proposed by Caudell and
% Manufacturing of GDP −0.059 Quinlan (2016). This paradox centers around
% Service of GDP −0.947** the fact that both increases and decreases in
Constant 3.624* fertility rates promote higher carbon emis-
R2 0.89 sions. Although seemingly paradoxical, these
N 75 trends can be understood within a framework
**
p<.01. *p<.05 combining theories of human behavior
Source: First published in Caudell and Quinlan (2016). inspired by evolutionary theory, particularly
All variables were log-transformed. LHT, and evolutionary psychology. From a
life history framework, the paradox can be
understood as an outcome of selection for
increase associated with a 0.56% increase in slow versus fast life history strategies, which
emissions. A 1% increase in GDP from the hold consequences for the demographic and
service sector was associated with a 0.95% consumption dynamics driving global carbon
decrease in carbon emissions per capita. emissions. Components of slower strategies,
Population and percentage of GDP from the including low mortality and high investment
CO2 emission per capita

Time orientation
Post-secondary enrollment (%)

Figure 22.3  Effect of time orientation on carbon emissions per capita as a function of
investment in education
 EVOLUTIONARY PSYCHOLOGY AND CLIMATE CHANGE
(education level) are negatively correlated
with total fertility rates across 100,000
women and 75 countries but also with higher
carbon emissions per capita in the 75 coun-
tries. Likewise, components of faster strate-
gies, including higher mortality and lower
investment, are associated with increased
fertility rates and so increased carbon emis-
sions in the future. In light of these results,
the paradox can be understood as an outcome
of how evolved human-mating strategies
respond to environmental risk levels.
The models presented here provide a robust
examination of how evolved psychological
constructs, in this case time orientation, may
respond to local levels of environmental risk.
Indeed, a country’s future orientation was
associated with total fertility rates after con-
trolling for mortality and investment levels,
suggesting the construct operates indepen-
dently from other life history traits. In addi-
tion, while the effect of future orientation was
seemingly small, about a 0.6 birth decrease,
these effects could have substantial conse-
quences for fertility and parental investment
decisions. Consider two equally educated
women who grew up exposed to similar
levels of environmental risk. One of these
individuals resides in Chile, with a future ori-
entation score of 31, and the other in India,
with an orientation score of 51. Compared to
the Indian, the Chilean would be predicted
to have one more child over her reproduc-
tive lifespan. Summed across a country, these
increases in the total fertility rate would
have considerable consequences for climate
change, for example, through increasing car-
bon legacies per woman.
The analysis further highlights how
evolved cognitive constructs can interact with
environmental risk and parental investment
to transform fertility patterns, sometimes in
unpredicted directions. For example, non-
educated women in present-oriented coun-
tries were predicted to have eight offspring
across their reproductive lifespans, whereas
in future-oriented countries, women with
the same level of education were predicted
449
to have about two offspring across their
­reproductive lifespans (Figure 22.1). In addi-
tion, a crossover interaction existed between
total fertility and mortality rates as time ori-
entations changed (Figure 22.2). Women in
present-oriented countries exposed to high
levels of environmental risk were predicted to
have around seven children whereas women
exposed to low risk levels were predicted to
have around five children. However, there
was a greater rate of decline in predicted TFR
at high mortality levels as time orientation
became more future oriented. This greater
rate of decline resulted in a crossover inter-
action such that high-mortality environments
predicted lower fertility than low-mortality
environments. This result, seemingly incon-
sistent with LHT, demonstrates the impor-
tance of including psychological constructs
such as time orientation in determining
behavioral responses to risk given they are
patterned by cultural (Gao, 2016) and eco-
nomic (Guthrie et al., 2009) factors that may
operate independently from the risk envi-
ronment. More generally, these results sug-
gest that the promotion of slower life history
strategies, through lowering of environmen-
tal risk (e.g., improving health and sanitation
systems), could reduce the future magnitude
of anthropogenic climate change by limiting
population growth in LMICs.
While limiting population growth repre-
sents an important step in addressing climate
change in LMICs, consumption patterns in
HICs show how the potential demographic
benefits of slower life history strategies
can be offset by corresponding increases in
per capita investments. In support, Model
2 showed that future orientation was posi-
tively associated with carbon emissions in
75 countries after controlling for the effects
of population, affluence (consumption), and
technology (Table 22.7). Importantly, the
model also provided support for the argu-
ment that higher education may allow future
time orientation to be somewhat decoupled
from its life history context and co-opted to
support pro-environmental behavior. Indeed,
450 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
as enrollment in post-secondary education
increased across the 75 countries, the effect
of future orientation on per capita CO2 emis-
sions decreased and was decoupled from car-
bon emissions in countries with the highest
enrollment levels (Figure 22.3). These results
suggest that education can help offset the
potential CO2 emissions from the increased
parental investment levels promoted by future
orientation. However, given the limited num-
ber of countries with time orientation data,
more cross-cultural data on time orientation
is needed to determine the global prevalence
of this relationship.
Determining whether future orientations can
be leveraged to promote pro-environmental
behavior in HICs will be important for address-
ing climate change globally. First, HICs will
continue to have the largest effect on Earth’s
carrying capacity due to resource consumption
patterns within these countries. Second, emis-
sions from HICs will continue to comprise the
vast majority of global carbon output and so it
is these countries that are, and will be, respon-
sible for increases in global risk levels (Cohen,
2010; Stephenson et al., 2010). From an LHT
framework, these increases in risk would result
in selection for faster life history strategies in
areas where individuals are unable to change
the morbidity/mortality associated with these
risks through increased investment. Many
of these individuals will live in LMICs. As
such, in LMICs many of the risks predicted to
result from climate change, including extreme
weather events and civil unrest (Cohen, 2010;
Stephenson et  al., 2010), should be consid-
ered extrinsic. These increasing levels of
extrinsic risk, combined with the tendency for
LMICs to exhibit faster strategies and present
orientations, point to a potentially catastrophic
possibility: rapid population growth within
those countries where growth has the largest
impact on carbon emissions. To prevent this
positive feedback loop between population
growth in LMICs and unsustainable invest-
ment in HICs, future research must examine
the cultural, ecological, and biological fac-
tors impacting time orientations, particularly
those that may promote pro-­ environmental
investment patterns (for similar arguments,
see Milfont and Gouveia, 2006; Swim et al.,
2010).
STUDY LIMITATIONS AND FUTURE
STUDIES
Several aspects of the data used in this study
may impact the applicability of results,
although the models do suggest how results
can be expanded through future studies.
First, demographic data come from women
only and so the relationships between time
orientation, environmental risk, and life his-
tory variables may not be appropriately gen-
eralizable to men. However, studies of time
orientation suggest that the construct may
have greater impacts on the behavior of men,
especially ‘life history relevant’ behaviors
such as risk taking (Zimbardo and Boyd,
1999). Second, the Hofstede time orientation
measures were collected once, meaning that
the models assumed that the time orientation
measure did not vary. For example, if the
time orientation measure was collected in
2002, then the model assumed the measure
was the same across a person’s development.
This assumption was also implicit in our
proxies of the risk environment (i.e., LEB),
where we assumed fertility responses were
patterned by early experiences with risk. In
fact, data on childhood mortality patterns
(Liu et al., 2016) and the impact of the HIV/
AIDS epidemic on fertility (Kalemli-Ozcan
and Turan, 2011) reveal that fertility prefer-
ences can respond quickly to the present risk
environment. In order to consider these reac-
tions more fine-grained longitudinal data are
needed. More broadly, research is needed to
understand the pace of fertility preference
and time orientation changes across an indi-
vidual’s life history and at a broader cultural
level. In terms of time orientation, model
results suggest that time orientation is likely
the product of numerous factors across
 EVOLUTIONARY PSYCHOLOGY AND CLIMATE CHANGE
cultures given that orientation was associated
with fertility patterns even after controlling
for mortality and investment patterns. Third,
the intraclass correlation in the current study
revealed that 38% of the variance in individ-
ual total fertility rates is associated with
country-level differences. These levels of non-
independence emphasize the many factors,
including cultural, economic, and historical,
that account for differences in fertility pat-
terns within and between countries. To this
end, a multilevel modeling approach is rec-
ommended for representing the interaction
between an individual’s life history strategies
and the higher-level cultural, political, and
geographic contexts in which these strategies
unfold. Finally, while changes in global
carbon emissions can be represented in a life
history framework as increases in risk, these
emissions are driven by broader societal
(e.g., switch from coal to natural gas) and
historical (advent of the combustible engine)
changes that pattern the nature of the risk
environment (e.g., intrinsic versus extrinsic
risk). Understanding how these shifts impact
the risk environment will provide insight into
future fertility and parental investment pat-
terns and so to their potential consequences
on climate change.
CONCLUSION
Approaches that contextualize human behav-
ior within an evolutionary perspective, includ-
ing evolutionary psychology and LHT, will be
important tools in understanding the demo-
graphic and consumption patterns driving
climate change. Identifying the factors under-
lying these patterns will assist in designing
interventions to limit the magnitude of cli-
mate change. The current study, building off
Caudell and Quinlan (2016), provides an
example of the practicality of this approach
through resolving the Population Paradox.
The paradox – that carbon emissions are both
positively associated with fertility rate
451
increases and decreases – can be explained
through the consequences of selection for
slow-to-fast life history strategies in LHT.
While accounting for the paradox, an
­evolutionary-informed perspective also pro-
vides insight into overcoming the paradox
and limiting carbon emissions. A psychologi-
cal consequence of life history selection, the
construct of time orientation, may be co-
opted to motivate pro-environmental behavior
in the present through investment in higher
education. Through its impacts on both fertil-
ity rates and investment patterns, time orien-
tation represents an important intervention
point to reduce both current and future pat-
terns of carbon emissions and the severity of
anthropogenic climate change. Future studies
should consider how other evolved psycho-
logical constructs can be co-opted to address
pressing global issues.
REFERENCES
Alan, S., & Ertac, S. (2014). Good things come
to those who (are taught how to) wait:
Results from a randomized educational inter-
vention on time preference. Available at SSRN
2566405. Retrieved from http://papers.ssrn.
com/sol3/papers.cfm?abstract_id=2566405
(Accessed 23 March 2015)
Anderson, C. L., Dietz, M., Gordon, A., &
Klawitter, M. (2004). Discount rates in
Vietnam. Economic Development & Cultural
Change, 52(4), 873–887.
Apostolidis, T., Fieulaine, N., & Soulé, F. (2006).
Future time perspective as predictor of can-
nabis use: Exploring the role of substance
perception among French adolescents. Addic-
tive Behaviors, 31(12), 2339–2343. https://
doi.org/10.1016/j.addbeh.2006.03.008
Arnocky, S., Dupuis, D., & Stroink, M. L.
(2012). Environmental concern and fertility
intentions among Canadian university stu-
dents. Population and Environment, 34(2),
279–292.
Ashkanasy, N., Gupta, V., Mayfield, M. S., &
Trevor-Roberts, E. (2004). Future orientation.
In R. J. House, P. J. Hanges, J. Mansour,
452 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
P. W. Dorfman, & V. Gupta (Eds.), Culture,
leadership, and organization. The Globe Study
of 62 societies (pp. 282–342). Thousand Oaks,
CA: Sage.
Bar, M., & Leukhina, O. (2010). Demographic
transition and industrial revolution: A macro-
economic investigation. Review of Economic
Dynamics, 13(2), 424–451. https://doi.org/
10.1016/j.red.2009.03.002
Bauer, M., & Chytilová, J. (2010). The impact of
education on subjective discount rate in
Ugandan villages. Economic Development
and Cultural Change, 58(4), 643–669.
Bauer, M., & Chytilová, J. (2013). Women, chil-
dren and patience: Experimental evidence
from Indian villages. Review of Development
Economics, 17(4), 662–675.
Boden, T., Andres, R., & Marland, G. (2017).
Global, Regional, and National Fossil-Fuel CO2
Emissions (1751—2014) (V. 2017) [Data set].
https://doi.org/10.3334/cdiac/00001_v2017
Boserup, E., Makhoul, N., Munn, R., Srinivasan,
T., Robinson, J., & Rocha, C. (1983). Popula-
tion and technological change: A study of
long-term trends. International Journal of
Health Services, 13(1), 15–31.
Boucekkine, R., Croix, D., & Peeters, D. (2007).
Early literacy achievements, population density,
and the transition to modern growth. Journal
of the European Economic Association, 5(1),
183–226.
Boyd, J. N., & Zimbardo, P. G. (2005). Time
perspective, health, and risk taking. In A.
Strathman & J. Joireman (Eds.), Understand-
ing behavior in the context of time: Theory,
research, and application. (pp. 85–107).
Mahwah, NJ: Lawrence Erlbaum Associates.
Bulled, N. L., & Sosis, R. (2010). Examining the
relationship between life expectancy, repro-
duction, and educational attainment. Human
Nature, 21(3), 269–289.
Burger, O., DeLong, J. P., & Hamilton, M. J.
(2011). Industrial energy use and the human
life history. Scientific Reports, 56. https://doi.
org/10.1038/srep00056
Buss, D. M., & Shackelford, T. K. (1997). Human
aggression in evolutionary psychological
perspective. Clinical Psychology Review,
17(6), 605–619.
Caldwell, J. C. (1976). Toward a restatement of
demographic transition theory. Population
and Development Review, 2(3), 321–366.
Caudell, M., & Quinlan, R. (2016). Life-history
theory and climate change: Resolving popu-
lation and parental investment paradoxes.
Royal Society Open Science, 3(11). https://
doi.org/10.1098/rsos.160470
Caldwell, R. M., Wiebe, R. P., & Cleveland, H. H.
(2006). The influence of future certainty and
contextual factors on delinquent behavior and
school adjustment among African American
adolescents. Journal of Youth and Adolescence,
35(4), 587–598. https://doi.org/10.1007/
s10964-006-9031-z
Carmi, N., & Arnon, S. (2014). The role of future
orientation in environmental behavior: Analyz-
ing the relationship on the individual and cul-
tural levels. Society & Natural Resources,
27(12), 1304–1320. https://doi.org/10.1080/
08941920.2014.928393
Caudell, M. (2015). The interplay of mortality,
economics, and female empowerment in
fertility transformations: A fixed effects anal-
ysis across 40 years and 167 countries.
Cross-Cultural Research, 49(4), 358–373.
https://doi.org/10.1177/1069397115591152
Caudell, M., & Quinlan, R. (2016). Life-history
theory and climate change: Resolving popu-
lation and parental investment paradoxes.
Royal Society Open Science, 3(11). 8 https://
doi.org/10.1098/rsos.160470
Caudell, M., & Quinlan, R. J. (2012). Resource
availability, mortality and fertility: A path ana-
lytic approach to global life history variation.
Human Biology, 84(2), 1.
Charlesworth, B. (1994). Evolution in age-­
structured populations. Cambridge: Cam-
bridge University Press.
Coale, A. J. (1984). The demographic transition.
The Pakistan Development Review, 23(4),
531–552.
Cohen, J. E. (2010). Population and climate
change. Proceedings of the American Philo-
sophical Society, 154(2), 158–182.
Cohen, P., Cohen, J., West, S., & Aiken, L.
(2003). Applied multiple regression/
correlation analysis for the behavioral
sciences. Mahwah, NJ: Lawrence Erlbaum
Associates.
Crowley, T. J. (2000). Causes of climate change
over the past 1000 years. Science, 289(5477),
270–277.
Dietz, T., & Rosa, E. A. (1997). Effects of
population and affluence on CO2 emissions.
 EVOLUTIONARY PSYCHOLOGY AND CLIMATE CHANGE
Proceedings of the National Academy of
Sciences, 94(1), 175–179.
Ebreo, A., & Vining, J. (2001). How similar are
recycling and waste reduction? Future orien-
tation and reasons for reducing waste as
predictors of self-reported behavior. Environ-
ment and Behavior, 33(3), 424–448. https://
doi.org/10.1177/00139160121973061
Ehrlich, P. R., & Holdren, J. P. (1971). Impact of
population growth. Science, 171(3977),
1212–1217.
Ellis, B. J., Figueredo, A. J., Brumbach, B. H., &
Schlomer, G. L. (2009). Fundamental dimen-
sions of environmental risk. Human Nature,
20(2), 204–268.
EVS. (n.d.). European Values Study Longitudinal
Data File 1981–2008 (EVS 1981–2008) (No.
ZA4804 Data file Version 3.0.0). Retrieved from
GESIS Data Archive website: doi:10.4232/
1.12253 (Accessed 13 January 2015)
Fang, T. (2003). A critique of Hofstede’s fifth
national culture dimension. International
Journal of Cross Cultural Management, 3(3),
347–368. https://doi.org/10.1177/1470595
803003003006
Franson, N., & Garling, T. (1999). Environmental
concern: Conceptual definitions, measure-
ment methods, and research findings. Journal
of Environmental Psychology, 19(4), 369–382.
https://doi.org/10.1006/jevp.1999.0141
Galor, O., & Weil, D. N. (2000). Population,
technology, and growth: From Malthusian
stagnation to the demographic transition and
beyond. American Economic Review, 90(4),
806–828.
Gant, L., Heath, K. M., Ejikeme, G. G., Snell,
C., & Briar-Lawson, K. (2009). Early mother-
hood, high mortality, and HIV/AIDS rates in
Sub-Saharan Africa. Social Work in Public
Health, 24(1/2), 39–46. https://doi.org/
10.1080/19371910802569435
Gao, X. (2016). Cultural differences between
East Asian and North American in temporal
orientation. Review of General Psychology,
20(1), 118–127.
Guthrie, L. C., Butler, S. C., & Ward, M. M.
(2009). Time perspective and socioeconomic
status: A link to socioeconomic disparities in
health? Social Science & Medicine, 68(12),
2145–2151.
Harrison, G. W., Lau, M. I., & Williams, M. B.
(2002). Estimating individual discount rates
453
in Denmark: A field experiment. American
Economic Review, 92(5), 1606–1617.
Henson, J. M., Carey, M. P., Carey, K. B., &
Maisto, S. A. (2006). Associations among
health behaviors and time perspective in
young adults: Model testing with boot-
strapping replication. Journal of Behavioral
Medicine, 29(2), 127–137.
Hill, E. M., Ross, L. T., & Low, B. S. (1997). The
role of future unpredictability in human risk-
taking. Human Nature, 8(4), 287–325.
Hofstede, G. (2001). Culture’s consequences:
Comparing values, behaviors, institutions,
and organizations across nations (2nd ed.).
Thousand Oaks, CA: Sage.
Horstmanshof, L., & Zimitat, C. (2007). Future
time orientation predicts academic engage-
ment among first-year university students.
British Journal of Educational Psychology,
77(3), 703–718. https://doi.org/10.1348/
000709906X160778
Hox, J. J. (2010). Multilevel analysis: Techniques
and applications. New York: Routledge.
Huber, Peter J. The behavior of maximum likeli-
hood estimates under nonstandard conditions.
Proceedings of the Fifth Berkeley Symposium
on Mathematical Statistics and Probability,
Volume 1: Statistics, 221–233, University of
California Press, Berkeley, Calif., 1967. https://
projecteuclid.org/euclid.bsmsp/1200512988
IPUMS-DHS. (n.d.). Demographic and Health Sur-
veys 1990–2009 [Data extract from DHS Recode
files]. Retrieved from Minnesota Population
Center and ICF International website: http://
idhsdata.org (Accessed 13 December 2014)
Jaroni, J. L., Wright, S. M., Lerman, C., &
Epstein, L. H. (2004). Relationship between
education and delay discounting in smokers.
Addictive Behaviors, 29(6), 1171–1175.
Jennions, M. D., & Petrie, M. (1997). Variation
in mate choice and mating preferences: A
review of causes and consequences. Biologi-
cal Reviews, 72(2), 283–327.
Johnson, C. Y., Bowker, J. M., & Cordell, H. K.
(2004). Ethnic variation in environmental
belief and behavior: An examination of the
new ecological paradigm in a social psycho-
logical context. Environment and Behavior,
36(2), 157–186. https://doi.org/10.1177/
0013916503251478
Joireman, J. A., Van Lange, P. A., & Van Vugt,
M. (2004). Who cares about the
454 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
environmental impact of cars? Those with an
eye toward the future. Environment and
Behavior, 36(2), 187–206. https://doi.org/
10.1177/0013916503251476
Jorgenson, A. K., & Clark, B. (2013). The relation-
ship between national-level carbon dioxide
emissions and population size: An assessment
of regional and temporal variation, 1960–
2005. PloS One, 8(2), e57107. https://doi.org/
10.1371/journal.pone.0057107
Kalemli-Ozcan, S., & Turan, B. (2011). HIV and
fertility revisited. Journal of Development
Economics, 96(1), 61–65.
Kaplan, H. (1996). A theory of fertility and
parental investment in traditional and modern
human societies. American Journal of Physical
Anthropology, 101(s 23), 91–135.
Kirby, K. N., Godoy, R., Reyes-García, V., Byron,
E., Apaza, L., Leonard, W., & Wilkie, D.
(2002). Correlates of delay-discount rates:
Evidence from Tsimane’ Amerindians of the
Bolivian rain forest. Journal of Economic
Psychology, 23(3), 291–316. https://doi.org/
10.1016/S0167-4870(02)00078-8
Kruger, D. J., Reischl, T., & Zimmerman, M. A.
(2008). Time perspective as a mechanism for
functional developmental adaptation. Jour-
nal of Social, Evolutionary, and Cultural Psy-
chology, 2(1), 1–22. https://doi.org/10.1037/
h0099336
Lindsay, J. J., & Strathman, A. (1997). Predictors
of recycling behavior: An application of a
modified health belief Model1. Journal of
Applied Social Psychology, 27(20), 1799–1823.
https://doi.org/10.1111/j.1559-1816.1997.
tb01626.x
Liu, L., Oza, S., Hogan, D., Chu, Y., Perin, J.,
Zhu, J., & Black, R. E. (2016). Global, regional,
and national causes of under-5 mortality in
2000–15: An updated systematic analysis
with implications for the Sustainable
Development Goals. The Lancet, 388(10063),
3027–3035.
Lochner, L., & Moretti, E. (2004). The effect of
education on crime: Evidence from prison
inmates, arrests, and self-reports. American
Economic Review, 94(1), 155–189.
Low, B. S. (1978). Environmental uncertainty
and the parental strategies of marsupials and
placentals. American Naturalist, 112(983),
197–213.
Low, B. S., Hazel, A., Parker, N., & Welch, K. B.
(2008). Influences on women’s reproductive
lives: Unexpected ecological underpinnings.
Cross-Cultural Research, 42(3), 201–219.
https://doi.org/10.1177/1069397108317669
Milfont, T. L., & Gouveia, V. V. (2006). Time
perspective and values: An exploratory study
of their relations to environmental attitudes.
Journal of Environmental Psychology,
26(1), 72–82. https://doi.org/10.1016/j.
jenvp.2006.03.001
Minkov, M. (2007). What makes us different
and similar: A new interpretation of the
World Values Survey and other cross-cultural
data. Sofia, Bulgaria: Klasika i Stil.
Minkov, M., & Hofstede, G. (2011). Hofstede’s
Fifth Dimension: New evidence from the
World Values Survey. Journal of Cross-­
Cultural Psychology, 43(1), 3–14. https://doi.
org/10.1177/0022022110388567
Murtaugh, P. A., & Schlax, M. G. (2009).
Reproduction and the carbon legacies of
individuals. Global Environmental Change,
19(1), 14–20. https://doi.org/10.1016/j.
gloenvcha.2008.10.007
Neter, J., Kutner, M. H., Nachtsheim, C. J., &
Wasserman, W. (1996). Applied linear
statistical models (Vol. 4). Chicago, IL: Irwin
Chicago.
Nettle, D. (2010). Dying young and living fast:
Variation in life history across English neigh-
borhoods. Behavioral Ecology, 21(2), 387.
Olli, E., Grendstad, G., & Wollebaek, D. (2001).
Correlates of environmental behaviors:
Bringing back social context. Environment
and Behavior, 33(2), 181–208. https://doi.
org/10.1177/0013916501332002
O’Neill, B. C., Dalton, M., Fuchs, R., Jiang, L.,
Pachauri, S., & Zigova, K. (2010). Global
demographic trends and future carbon emis-
sions. Proceedings of the National Academy
of Sciences, 107(41), 17521–17526. https://
doi.org/10.1073/pnas.1004581107
O’Neill, B. C., Liddle, B., Jiang, L., Smith, K. R.,
Pachauri, S., Dalton, M., & Fuchs, R. (2012).
Demographic change and carbon dioxide
emissions. The Lancet, 380(9837), 157–164.
https://doi.org/10.1073/pnas.1004581107
Oreopoulos, P., & Salvanes, K. G. (2011). Price-
less: The nonpecuniary benefits of schooling.
Journal of Economic Perspectives, 25(1),
 EVOLUTIONARY PSYCHOLOGY AND CLIMATE CHANGE
159–184. https://doi.org/10.1257/jep.
25.1.159
Park, S., & Lemaire, J. (2011). Culture matters:
Long-term orientation and the demand for
life insurance. Asia-Pacific Journal of Risk
and Insurance, 5(2), 1–21.
Peetsma, T. T. D. (2000). Future time perspec-
tive as a predictor of school investment.
Scandinavian Journal of Educational
Research, 44(2), 177–192. https://doi.
org/10.1080/713696667
Perez-Arce, F. (2011). The Effect of Education
on Time Preferences. Working Paper, RAND
Labor and Population. Santa Monica, CA.
Peugh, J. L. (2010). A practical guide to multi-
level modeling. Journal of School Psychol-
ogy, 48(1), 85–112.
Promislow, D. E. L., & Harvey, P. H. (1990).
Living fast and dying young: A comparative
analysis of life-history variation among mam-
mals. Journal of Zoology, 220(3), 417–437.
Quinlan, R. J. (2007). Human parental effort and
environmental risk. Proceedings. Biological Sci-
ences / The Royal Society, 274(1606), 121–125.
https://doi.org/10.1098/rspb.2006.3690
Quinlan, R. J. (2010). Extrinsic mortality effects
on reproductive strategies in a Caribbean
community. Human Nature, 21(2), 124–139.
https://doi.org/10.1007/s12110-010-9085-1
Rabe-Hesketh, S., & Skrondal, A. (2012). Multi-
level and longitudinal modeling using Stata
(3rd ed.). College Station, TX: Stata Corp.
Roff, D. A. (2002). Life history evolution. Sun-
derland, MA: Sinauer.
Rosa, E. A., & Dietz, T. (2012). Human drivers
of national greenhouse-gas emissions.
Nature Climate Change, 2(8), 581–586.
Rothspan, S., & Read, S. J. (1996). Present
versus future time perspective and HIV risk
among heterosexual college students. Health
Psychology, 15(2), 131.
Schechter, D. E., & Francis, C. M. (2010). A life
history approach to understanding youth time
preference. Human Nature, 21(2), 140–164.
https://doi.org/10.1007/s12110-010-9084-2
Shi, A. (2003). The impact of population pres-
sure on global carbon dioxide emissions,
1975–1996: Evidence from pooled cross-
country data. Ecological Economics, 44(1),
29–42. https://doi.org/10.1016/S0921-
8009(02)00223-9
455
Smith, E. A. (2000). Three styles in the
evolutionary analysis of human behavior. In
L. Cronk, N. Chagnon, & W. Irons (Eds.),
Adaptation and human behavior: An
anthropological perspective (pp. 27–46).
New York, NY: Aldine De Gruyter.
Sörqvist, P., & Langeborg, L. (2019). Why
people harm the environment although they
try to treat it well: An evolutionary-cognitive
perspective on climate compensation.
Frontiers in Psychology, 10, 348.
StataCorp. (2019). Stata Statistical Software:
Release 16. StataCorp LLC.
Stearns, S. C. (1989). Trade-offs in life-history
evolution. Functional Ecology, 3(3), 259–
268. https://doi.org/10.2307/2389364
Stearns, S. C. (1992). The evolution of life
histories. Oxford: Oxford University Press.
Stephenson, J., Newman, K., & Mayhew, S.
(2010). Population dynamics and climate
change: What are the links? Journal of Public
Health, 32(2), 150–156. https://doi.org/
10.1093/pubmed/fdq038
Swim, J., Clayton, S., Doherty, T., Gifford, R.,
Howard, G., Reser, J., & Weber, E. (2010).
Psychology and global climate change:
Addressing a multifaceted problem and set
of challenges. Washington DC: American
Psychological Association.
Toepoel, V. (2010). Is consideration of future
consequences a changeable construct? Per-
sonality and Individual Differences, 48(8),
951–956. https://doi.org/10.1016/j.
paid.2010.02.029
Tooby, J., & Pines, D. (1985). The emergence of
evolutionary psychology. In Emerging syn-
theses in science (pp. 67–76). Santa Fe, NM:
Santa Fe Institute.
Tooby, J., & Cosmides, L. (1988). The evolution
of war and its cognitive foundations. Insti-
tute for Evolutionary Studies Technical
Report, 88(1), 1–15.
Trivers, R. (1972). Parental investment and
sexual selection. In D. T. Campbell (Ed.),
Sexual selection and the descent of Man (pp.
136–179). Chicago, IL: Aldine de Gruyter.
Van der Pol, M. (2011). Health, education and
time preference. Health Economics, 20(8),
917–929. https://doi.org/10.1002/hec.1655
Walker, R., Gurven, M., Hill, K., Migliano, A.,
Chagnon, N., Souza, R. D. E., & Yamauchi,
456 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
T.  (2006). Growth rates and life histories in
twenty-two small-scale societies. American
Journal of Human Biology, 311, 295–311.
https://doi.org/10.1002/ajhb.20510
Wall, G. (1995). Barriers to individual environ-
mental action: The influence of attitudes and
social experiences Canadian Review of Soci-
ology/Revue Canadienne de Sociologie,
32(4), 465–489. https://doi.org/10.1111/j.
1755-618X.1995.tb00182.x
Wesely Schultz, P. (2001). The structure of envi-
ronmental concern: Concern for self, other
people, and the biosphere. Journal of Envi-
ronmental Psychology, 21(4), 327–339.
https://doi.org/10.1006/jevp.2001.0227
Williams, G. C. (1957). Pleiotropy, natural
selection, and the evolution of senescence.
Evolution, 11, 398–411.
Wilson, M., & Daly, M. (1997). Life expectancy,
economic inequality, homicide, and repro-
ductive timing in Chicago neighbourhoods.
British Medical Journal, 314(7089),
1271–1274.
World Bank. (n.d.). DataBank: Metadata Glos-
sary. Retrieved August 20, 2019, from https://
databank.worldbank.org/metadataglossary/
africa-development-indicators/series/NV.AGR.
TOTL.CD
World Bank. (2009). World Development Indi-
cators. Retrieved from World Bank Develop-
ment Indicators website: www.worldbank.
org/ (Accessed 13 December 2014)
World Bank. (2016). World Development Indi-
cators. Retrieved December 13, 2014, from
World Bank Development Indicators website:
http://www.worldbank.org/
WVS. (n.d.). World Values Survey 1981–2014
official aggregate v.v.20150418 (JDSystems
No. Aggregate File Producer). Retrieved from
World Values Survey Association website:
www.worldvaluessurvey.org (Accessed 13
December 2014)
Zimbardo, P. G., & Boyd, J. N. (1999). Putting
time in perspective: A valid, reliable individual-
differences metric. Journal of Personality and
Social Psychology, 77(6), 1271–1288.

Evolutionary Psychology and
Thanatology: Responses to Death
INTRODUCTION
Like humans, at some point(s) during their
lives group-living nonhuman animals (hereaf-
ter: ‘animals’) will probably encounter death in
members of their own and other species. The
dying and dead individuals may be complete
strangers, acquaintances, or others with whom
strong emotional bonds are shared (e.g., kin,
close friends). Evolutionary thanatology seeks
to understand the origins, diversity, mecha-
nisms, and functions of humans’ and other
species’ behavioral and emotional reactions
toward the dying and the dead (Anderson
et  al., 2018). Although most research in this
field focuses on intra-species phenomena,
responses toward sick, injured, or the corpses
of other species are also of interest, as they
may offer future resources (usually food),
signal danger, or simply evoke curiosity.
Behavioral reactions to the moribund and
dead vary widely within and across spe-
cies; a non-exhaustive list includes indiffer-
ence, avoidance, manipulation, aggression
23
James R. Anderson
(including mutilation), affiliation (including
tending, transporting, and caretaking), sexual
behavior, partial or full ingestion, abandon-
ment, and various ways of actively dispos-
ing of the corpse (see Gonçalves and Biro,
2018; Gonçalves and Carvalho, 2019). Some
reactions probably have a long history in
human evolution, whereas others are more
recent and culturally restricted (Pettitt, 2011).
Emotional states that accompany behaviors in
the presence of the dead range from arousal
and alarm, for example, in response to per-
ceiving no signs of life from a corpse and
sensing potential danger, to feelings of frus-
tration, anger, sadness, and in humans and
other primates at least, grief. Of course, many
animals kill others, and therefore ­predator–
prey relationships and perceptions of and
responses to predation events fall within the
scope of evolutionary thanatology. In this
chapter, I use the literature on death and
dying in selected nonhuman taxa to explore
how thanatological behaviors might be bet-
ter understood, or at least addressed, from
458 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
the perspective of ‘Tinbergen’s four ques-
tions’. The questions will be presented more
fully in the following section, but in the way
of a brief introduction, immediate causa-
tion refers to external and internal cues and
motivational states that give rise to a behav-
ior. Development is addressed by looking at
sources of individual differences in behavior,
including genetic predispositions and learn-
ing experiences. The question of evolution
evokes comparisons among species, mapping
behaviors that are shared, ancestral traits, and
others that have evolved more recently and
in particular lineages. Finally, functions can
be addressed by looking at how the behav-
ior might contribute to fitness. For illustra-
tion, I focus on two striking thanatological
phenomena, namely necrophoric behavior in
social insects, and postmortem transport and
care of dead infants by adult female monkeys
and apes. This selective review introduces the
broad field of evolutionary thanatology and
presents examples of researchable questions
and approaches that it generates.
THANATOLOGY: HUMANS AND
OTHER SPECIES
Most human adults know that life eventually
culminates in death, and that this applies to
all living creatures: death befalls everyone,
without exception. This understanding
reflects two components of the typical adult’s
concept of death (at least in Western, indus-
trialized societies), namely inevitability and
universality. Two other components of the
mature Western human’s concept of death are
non-functionality (dead individuals no longer
perceive, think, feel, or act) and causality
(some knowledge of biological causes of
death, e.g., damage to vital organs, loss of
blood, lack of oxygen) (see Gire, 2014 for a
discussion of differences in conceptualiza-
tions of death across cultures). Developmental
psychologists with an interest in thanatology
have studied how children gradually form a
mature concept of death (typically achieved
by 10–11 years of age), mapping their death-
related cognitions at various ages, and
exploring how environmental and experien-
tial influences help to shape these processes.
For these aims psychologists typically use
questionnaires, interviews, or other less
direct techniques such as drawings to explore
children’s private views and emotions (see
e.g., Speece and Brent, 1984; Slaughter and
Griffiths, 2007; Bonoti et al., 2013). Despite
some culture-related variations in attainment
of the above-mentioned components, the
overall developmental trajectory toward a
mature understanding of death, especially
biological aspects, appears largely universal
(e.g., Brent et  al., 1996; Panagiotaki et  al.,
2015).
In addition to psychologists and other
researchers, practitioners who deal with
death-related events and consequences in
humans include medical and nursing pro-
fessionals, counsellors, epidemiologists,
pathologists, economists, sociologists, and
anthropologists. One particular group of
scientists – funerary archeologists – are par-
ticularly interested in the remains of those
who died long ago, hundreds, thousands, or
even millions of years ago. Studying these
ancient remains can unearth valuable infor-
mation about phenomena such as diet, age,
likely cause of death, non-fatal trauma and
diseases, and damage inflicted after death
(which might include defleshing and dis-
membering). Furthermore, analyses of the
immediate surrounding environment (e.g.,
other human and nonhuman remains, arte-
facts, signs of deliberate burial or caching),
and aspects of the corpse (e.g., orientation,
posture, remnants of clothing or accessories)
can be used to make inferences about the
evolution of human funerary activities within
particular social and psychological contexts
(Pettitt, 2011, 2018). By giving greater con-
sideration to various species’ immediate and
long-term responses to death, and to relevant
physical and social anthropological data,
thanatologists (most of whom probably do
 EVOLUTIONARY PSYCHOLOGY AND THANATOLOGY
not explicitly identify themselves as such)
can construct a clearer picture of responses
to this universal and momentous event in the
context of evolution.
One distinguishing feature of thanato-
logical studies of nonhuman animals is that
because they cannot use verbal or projective
techniques for collecting data, researchers
must focus on publicly observable behavioral
responses to moribund or dead companions
or others. Observational and experimen-
tal studies both contribute. The latter may
involve exposing animals to death-related
cues, for example visual or chemical signs
of injury or death. Some researchers also
measure changes in physiological status in
relation to an animal’s experiences of death.
Examples of both kinds of approaches will be
seen in later sections.
TINBERGEN’S FOUR QUESTIONS
Tinbergen (1963) proposed that a compre-
hensive understanding of any behavior was
more likely to emerge if researchers consid-
ered four questions: What is causing this
behavior to happen now? What is the devel-
opmental history of this behavior within the
individual? What is the function of this
behavior in terms of survival value? How did
this behavior evolve? The first two questions
are often considered as addressing proximate
mechanisms, for example current environ-
mental and/or internal events eliciting the
behavior (e.g., what triggers it, what moti-
vates it), and the extent to which the behavior
can be considered as pre-programmed (or
‘innate’), or influenced by learning – the
ontogeny issue. The latter two questions are
considered to address ultimate reasons,
namely how is this behavior adaptive, or
what is its past survival value (and ‘current
utility’, see Bateson and Laland, 2013), and
how and when did it evolve? The evolution-
ary issue can be addressed, for example, by
comparing species that are more or less
459
phylogenetically related and looking to see
whether a behavior is common across related
species (and hence more likely to be an
ancient trait) or whether it has more recently
appeared within one (or more) specific taxo-
nomic group(s). Although there has been
debate about applying and updating
Tinbergen’s four questions 50 years on, there
is general agreement about the enduring value
of the Tinbergian approach (Barrett et  al.,
2013; Bateson and Laland, 2013; Taborsky,
2014; Kapheim, 2019). This approach is taken
below to discuss two particular examples of
thanatological behavior, namely necrophore-
sis in social insects, and postmortem infant
care and transport in nonhuman primates.
SOCIAL INSECTS: CORPSE
MANAGEMENT
The responses of social insects to dead con-
specifics are relatively well studied, and
illustrate nicely the value of Tinbergen’s
questions for thanatological investigations.
Furthermore, it can be argued that that
responses to sick, injured, dead, or dying
insects are easier and less ethically challeng-
ing to study than in longer-lived, cognitively
and emotionally more complex vertebrate
species, such as fish, birds, and mammals.
For species that congregate permanently
(or semi-permanently) at a fixed site, deaths
of community members pose problems for
the living. The enclosed home environments
of social insects contain often high population
densities and food reserves, thereby provid-
ing favorable conditions for the proliferation
of potentially harmful organisms. Decaying
corpses within the nest or hive may give rise
to disease; therefore corpse management
strategies are important. Social insects such
as bees, wasps, ants, and termites deal with
this problem through a suite of behaviors that
have evolved to ensure the rapid and effective
disposal of the dead (for reviews, see López-
Riquelme and Fanjul-Moles, 2013; Sun et al.,
460 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
2018): 1) Necrophoresis is the general term
for transport and disposal of corpses either
outside the nest or at refuse sites within the
nest. 2) Cannibalism is the consumption of
dead conspecifics. Sometimes referred to as
‘conspecific necrophagy’ to acknowledge
that no active killing is involved (Brown and
Norris, 2004), it is common in termites, but
also occurs in some ants. 3) Burial behavior
consists of covering corpses with soil or other
matter; again, common in termites, it has also
been reported in some ants. 4) Necrophobia
is the simple avoidance of corpses or areas
associated with corpses, and is a widespread
response to corpses encountered both within
and outside the nest. In some species there
is flexibility: one behavioral response can be
replaced by another, depending on the nature
of the corpse (e.g., age, whether conspecific
or allospecific: Renucci et  al., 2011; Neoh
et al., 2012; Sun et al., 2013).
Proximal Causes
What elicits corpse management behaviors in
social insects? Experiments using stationary
models of insects or other objects, the lack of
identifiable sounds or thermal cues from
corpses, and the darkness within nests or
hives mean that visual, auditory, and tactile
cues can usually be ruled out. Instead, there
is general agreement on a predominant role
for chemical cues in triggering both necro-
phobic and necrophoric reactions. In fact,
corpse disposal by social insects nicely illus-
trates the prototypical ethological sequence
in which a sign stimulus (in this instance, one
or more chemical signals) triggers an innate
releasing mechanism (a neural network in the
brain), which in turn produces a fixed action
pattern (here, the behavioral sequence result-
ing in avoidance or disposal of the corpse). In
a pioneering study, Wilson et  al. (1958)
showed that a specific substance produced by
decaying corpses – oleic acid – caused
healthy ants to dispose of the corpses. Oleic
acid proved so effective in this regard that
any objects daubed with the substance – even
live, healthy nestmates – were transported by
healthy ants to refuse piles. Oleic acid is one
of several chemical cues that alone or in
combination can elicit so-called undertaking
behavior – detection, removal, and disposal
of dead colony members – in social insects
(López-Riquelme and Fanjul-Moles, 2013).
Another fatty acid product of corpses – lin-
oleic acid – also induces necrophoric reac-
tions in ants; in cockroaches this compound
had an even stronger repellent effect than
oleic acid (Rollo et  al., 1994). The latter
authors proposed the term ‘necromones’ for
chemicals that signaled death and triggered
specific responses. Sometimes removal of
freshly dead nestmates occurs before the
buildup of necromones is sufficient to induce
necrophoresis. It has been shown that two
chemical compounds in live ants might pre-
vent them from being responded to as corpses
by nestmates, by masking necromones; how-
ever, these compounds fade rapidly in freshly
dead workers, triggering the latter’s removal
even before necromones take effect (Choe
et al., 2009). Several studies have shown that
different necromones from fresh and older
corpses may induce different behaviors; in
termites, for example, ‘early’ death cues trig-
gered cannibalism whereas ‘late’ cues gave
rise to burial responses (Sun et al., 2017; see
also Neoh et  al., 2012). Early experiments
also demonstrated that chemical cues trig-
gered necrophoresis in honey bees (Visscher,
1983).
Development
An interesting aspect of corpse management
in social insects is that workers within the
colony show varying degrees of specializa-
tion in ‘undertaking’ activities. In one experi-
mental study using three laboratory colonies
of ants containing 48, 105, and 146 individu-
ally marked workers, respectively, more than
75% of workers in each colony encountered
corpses but less than 30% showed
 EVOLUTIONARY PSYCHOLOGY AND THANATOLOGY
undertaking activities, defined as picking up
and carrying a corpse for more than 3 cm.
Some workers removed dead ants more fre-
quently than expected by chance (Julian and
Cahan, 1999). In the smallest colony usually
only one undertaker engaged with the corpse
on each trial, whereas in the larger colonies
several did so. Among the latter, those that
handled a corpse most frequently (‘special-
ists’) were more likely than non-specialists to
complete removal of the corpse and, in the
largest colony, in the shortest time.
The authors suggested age-related or genetic
differences in undertaking activities, but
ruled out an effect of individual learning on
efficiency of performance, at least in the
24–27 trials per colony, conducted over a
two-week period.
An early study of individually marked
honey bees concluded that 1–2% of the colony
population were specialized undertakers, and
that they retained this specialized role over
a relatively long period of time (Visscher,
1983). Bumblebee workers that removed lar-
val corpses but not adult corpses were also
described as behaviorally more specialized
than non-undertaker workers (Munday and
Brown, 2018); the former were also sig-
nificantly larger than those who specialized
in removing adult corpses. Later work on
honey bees identified more undertakers – at
least 20% of workers – and revealed several
behavioral differences between undertakers
(and guards) compared to other workers (food
storers and wax workers). Genotypic differ-
ences in response thresholds to different kinds
of environmental stimuli appear likely to con-
tribute to the extent of undertaking activities.
Selective breeding can result in different levels
of nest hygienic behaviors including corpse
removal (e.g., Rothenbuhler, 1964; Oldroyd
and Thompson, 2007); however, the spe-
cialization of undertakers does not appear as
extreme as in some insect subcastes (Robinson
and Page, 1988; Trumbo et al., 1997). In view
of the relatively short period of time in which
individuals specialize in undertaking – u­ sually
several days – learning may play little part
461
in the emergence or efficiency of the behav-
ior; indeed, efficiency did not improve with
greater experience of removing corpses
(Trumbo and Robinson, 1997) (but see Arathi
et al., 2000, for an alternative perspective on
the role of experience).
López-Riquelme and Fanjul-Moles (2013)
present an overview of what little is known
about the genetics of undertaking specializa-
tion in social insects. They point out the likely
relationship between increasing genetic
determination and reduced behavioral plas-
ticity (e.g., in honey bees), and between high
morphological specialization and reduced
sensitivity to death cues (e.g., in soldier ants);
they also discuss neurobiological specializa-
tions in undertakers.
Function
As indicated above, dead individuals in the
nest are a potential source of disease, so dis-
posing of the dead should be of benefit in
increasing survival of the living. An experi-
ment with red ants demonstrates this advan-
tage of corpse disposal (Diez et  al., 2014).
Survival rates were compared between colo-
nies in which ants could freely remove
corpses and colonies in which the size of the
nest entrance was made smaller to impede
normal corpse disposal activities. Freshly
killed nestmates were placed in each nest and
left over a seven-week period. Results
showed that from the eighth day after corpse
placement, survival rates of adult workers in
colonies with restricted removal possibilities
were significantly reduced compared to col-
onies with normal-size nest entrances.
Furthermore, ants in control nests were seen
to engage in typical necrophoric behaviors,
whereas those in restricted-entrance nests
cut up corpses and ejected the pieces, or
moved them to areas in the nest so as to
maximize the distance between the corpses
and the larvae. Over time, larvae survival
rates also declined in the restricted-entrance
nests.
462 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Evolution
Avoiding dead conspecifics or places where
cues to conspecific death are detected (necro-
phobia), and actively disposing of corpses
(necrophoresis), is clearly advantageous. The
latter, described in social bees, wasps, and
ants (hymenopterans), as well as termites
(isopterans), has been proposed as an exam-
ple of convergent evolution (Sun and Zhou,
2013; Sun et al., 2018).
Yao et al. (2009) proposed an ancient ori-
gin of sensitivity to chemical signals result-
ing in avoidance of corpses and death-related
sites in both insects and crustaceans, dating
back possibly to ancient unicellular organ-
isms. Although some progress has been made
with regard to molecular bases in social
insects, further studies on a wider range of
species are necessary to better understand
the evolution from simple ancestral death-
recognition responses to more complex,
sequentially organized corpse management
strategies (Sun et al., 2018).
Digressing briefly from social insects to
crustaceans, hermit crabs nicely illustrate
diverging evolved responses to death cues
in marine vs terrestrial environments. In this
case, responses concern not avoidance or
disposal, but attraction. All ancestral hermit
crabs were marine animals, and marine her-
mit crabs today live in unremodeled shells
that become available upon the death of other
marine hermit crabs or gastropods. By con-
trast, terrestrial hermit crabs have become
specialized to live only in shells that have
been physically remodeled by conspecifics.
Valdes and Laidre (2019) recently demon-
strated differences in response to chemical
cues from dead conspecifics vs heterospe-
cifics in marine and terrestrial hermit crabs.
Whereas the latter rapidly gathered around
conspecific death sites in particular – pre-
sumably because of the possible presence
of a suitable new host shell – their marine
counterparts showed no such differential
response. This work shows not only the pow-
erful behavioral effects of chemical cues
from corpses, but also that whereas some
species avoid the source or act to get rid of
it, other species are positively attracted to it;
necrophagous insects are another example of
the latter (e.g., Frederickx et al., 2012).
NONHUMAN PRIMATES: CARE AND
TRANSPORT OF DEAD INFANTS
The mother–infant relationship is perhaps the
most studied of all social relationships in
nonhuman primates. The newborn infant is
dependent on its mother for survival: she
provides nourishment and protection against
multiple dangers including accidents, adverse
weather, predatory attacks, and conspecific
aggression (see Altmann, 1981; Nicolson,
1987, 1991; Hayashi and Matsuzawa, 2017).
In many species the strong emotional bond
that develops between mother and offspring
persists for as long as both are alive, particu-
larly in philopatric species where one or both
remain near to their birthplace for most or all
of their lives (van Noordwijk, 2012).
The mother–infant relationship also figures
prominently in the primate thanatology liter-
ature. Many authors have described mothers
continuing to carry and care for their infant
even though it is dead. The phenomenon is
widespread in primates, although it varies in
incidence and duration, with influencing fac-
tors including species, ecology, age of infant,
cause of death of infant, individual maternal
traits, and climatic conditions (e.g., Sugiyama
et  al., 2009; Watson and Matsuzawa, 2018;
Das et  al., 2019). Acronyms used for mam-
malian mothers’ response to their dead
offspring include DIC (‘deceased-infant car-
rying’, in Das et al., 2019 for primates), and
PAB (‘postmortem attentive behavior’, in
Bearzi et al., 2018 for cetaceans); the former
emphasizes carrying and transport, the latter
implies caregiving more generally, not just
toward offspring but also other conspecifics.
Because primate mothers usually both carry
and take care of their infant’s corpse (e.g., by
 EVOLUTIONARY PSYCHOLOGY AND THANATOLOGY
grooming and preventing insect larvae from
settling in it), I refer to this constellation of
behaviors as ‘postmortem-infant care and
transport’ (PICT).
Proximate Causes
Whereas in social insects chemical cues from
the dead are the primary triggers of necro-
phobic or necrophoric responses in the living,
the picture is more complicated in primates.
The immediate causes of PICT in primate
mothers are probably multiple and include
both internal events and external sensory
cues. First, the physiological condition of the
mother – reflecting hormonal changes in late
pregnancy and the periparturitional stage –
causes her to be strongly attracted to and
highly motivated to transport, care for, and
protect her baby (Nicolson, 1991; Kristal,
2009). In fact, experimental studies of
mother–infant separation and adoption have
shown that new primate mothers will accept,
hold, and nurture any infant of appropriate
age, in some cases even if it is another spe-
cies (Harlow, 1974; Smith, 1986; Owren and
Dieter, 1989). Recently, the possibility that
oxytocin plays an important role in PICT was
raised by Bercovitch (2020), although he
emphasized that this hormone promotes
social bonding in general, rather than being a
specific trigger of continued maternal behav-
ior toward dead infants.
Despite being hormonally primed to
behave maternally, especially if she has
raised other infants, the mother of a dead
infant shows awareness that this one neither
responds to anything she does nor shows
any spontaneous behavior at all. A corpse
has no grasping reflex, so to carry and trans-
port it the mother must make compensatory
behavioral adjustments such as holding the
corpse with one hand and walking triped-
ally instead of quadrupedally, or even hold-
ing it in her mouth. While feeding arboreally
she may have to keep hold of the corpse or
secure it in her groin pocket or under an arm.
463
Another likely causal factor in PICT is that
the dead infant still presents morphological
features that make it attractive to the mother
(and possibly others). Visual features (‘baby
schema’) including infantile coloration, coat
texture, and bodily shape and facial propor-
tions (e.g., Alley, 1980; Sato et al., 2012) are
attractive to mothers and others. For some
individuals including the mother, the total
lack of animacy in the infant corpse might
also increase its interest value, at least for a
limited time. The dead infant also continues
to provide tactile stimulation – known to be
crucial in mother–infant bonding (Harlow,
1974) – but only if the mother touches or
holds it. Unlike in social insects, there has
been no suggestion that chemical cues play
any role in primate PICT. But nor do such
cues appear to be aversive. Although human
observers have commented on the repulsive
smell from infant primate corpses after a few
days, this seems to have little or no effect on
the mother’s behavior, or indeed that of other
members of the group (Biro et al., 2010; De
Marco et al., 2018, but see Sugiyama et al.,
2009). In fact, there are no experimental stud-
ies on sensitivity to putrescine or other puta-
tive necromones in nonhuman primates.
With time elapsed since death the corpse
progressively decays, and in terms of visual
and tactile features resemblance to a normal,
live infant diminishes. PICT sometimes con-
tinues even though the corpse has mummi-
fied and portions of the body detach (e.g.,
Sugiyama et al., 2009; Biro et al., 2010; De
Marco et al., 2018). This deterioration, con-
temporaneous with the mother’s changing
hormonal condition through lack of suck-
ling, eventually leads to abandonment of the
corpse. Abandonment may be gradual – the
mother lays the corpse on the ground for
increasingly longer periods – or more abrupt:
sometimes the mother is spotted without
her dead infant although she was carrying
it the previous evening. There are increas-
ing reports of prosimian, monkey, and great
ape mothers ingesting parts of their infant’s
corpse. This ‘maternal cannibalism’ has been
464 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
interpreted as possible abnormal behavior
due to stressful environmental conditions
(Tartabini, 1991; Dellatore et  al., 2009) or
as a normal albeit rare event (Fedurek et al.,
2019; Trapanese et  al., 2020), reflecting a
qualitative change in the mother’s perception
of the corpse from ‘her infant’ to a poten-
tially edible object (Tokuyama et  al., 2017;
De Marco et al., 2018).
When considering causes of death-related
reactions in highly social and intelligent mam-
mals such as nonhuman primates, emotional
factors should not be neglected (Anderson,
2016). There is compelling behavioral and
physiological evidence that recent bereave-
ment is stressful for primates (Engh et  al.,
2006), and the strong emotional bond that
the mother experiences with her infant is
one likely cause of PICT. Disruption of the
mother–infant bond has profound emotional
consequences in both partners. If prolonged,
in infants it can result in depression, as indi-
cated by behavioral and physiological param-
eters, and symptom alleviation following
antidepressant pharmacological intervention
(for review see Worlein, 2014). Loss-induced
depression can contribute toward an early
death, especially in free-ranging conditions.
Responses to orphaned infants by individu-
als other than the mother range from indif-
ference to limited caretaking and attempted
adoption, but long-term survival prospects
of dependent orphans are poor (Rhine et al.,
1980; Goodall, 1983; Thierry and Anderson,
1986; Fashing et  al., 2011). Infants might
either stay nearby or continue clinging to
their dead mother, in which case survival is
likely to be brief. Mothers of dead infants,
by contrast, may continue to express the
emotional bond with the infant for longer,
through PICT. Some authors have described
behaviors in PICT mothers as possible indi-
cators of intense emotion, such as the dazed
expression of bereaved chimpanzee mothers
(van Lawick-Goodall, 1968), reduced feed-
ing, increased passivity, and social isolation
and withdrawal in various species (Das et al.,
2019; Takeshita et  al., 2020). Based on the
accumulating behavioral and physiological
data, a strong argument can be made that
mothers showing PICT are experiencing
grief in at least some ways comparable to the
human experience following the death of a
loved one (King, 2013; Anderson, 2017; Das
et al., 2019).
Development
PICT can be seen as an extension of normal
maternal behavior, albeit with compensatory
adjustments for the lifelessness of the infant
similar to but perhaps more obvious than
those shown by mothers of physically handi-
capped, injured, or sick infants (for exam-
ples, see Nakamichi, 1986; Anderson et  al.,
1995; Matsumoto et al., 2016). To this extent
PICT is a product of the same genetically
determined and experiential factors that
underlie normal maternal behavior and per-
formance. Genetic factors include sex differ-
ences in neuro-hormonal mechanisms that
result in females being especially strongly
attracted to and interested in infants (Nicolson
1991), sex differences in play and other
social activities before puberty (Lonsdorf,
2017) and, in females, pre- and postpartum
hormonal changes that are critical for mater-
nal behavior (Saltzman and Maestripieri,
2011). Other potentially relevant factors
include maternal temperament and style, and
stress (Altmann, 1980; Fairbanks, 1996;
Sullivan et  al., 2011; Maestripieri, 2018);
however, there are insufficient data to allow
any conclusions about individual differences
in these traits being related to aspects of PICT.
Experiences can affect all the genetically
predisposed traits mentioned above. Greater
experience with infants comes with parity,
which is one maternal trait that has been
examined in relation to PICT. At least two
competing predictions can be made: (1) pri-
miparous mothers should show more PICT
because they are primed to look after their
first infant but are inexperienced with dead
vs live infants, vs (2) multiparous mothers
 EVOLUTIONARY PSYCHOLOGY AND THANATOLOGY
should show more PICT because with aging
each infant represents a greater biological
investment; also, multiparous mothers might
have prior experience of unconscious, unani-
mated infants recovering (see next section on
functions). In a study of 157 cases in free-
ranging, provisioned Japanese macaques
over a 24-year period, Sugiyama et al. (2009)
used maternal age as a proxy for parity, but
reported no significant differences between
younger and older mothers in the incidence or
duration of PICT. They also noted that some
mothers were repeat carriers of dead infants;
one carried three dead infants. By contrast,
Das et  al. (2019) found that older mothers
carried dead infants for longer, supporting
the hypothesis of greater investment in later-
born offspring. Whether the strength of the
attachment between mother and infant influ-
ences PICT remains to be clarified (Watson
and Matsuzawa, 2018). Observations of two
mothers each carrying their dead infant at the
same time led to the suggestion of a role for
observational learning (Biro et al., 2010). The
fact that some form of PICT occurs in at least
19 primate species (Das et al., 2019) proba-
bly indicates a stronger biological than social
basis, but a related and more easily testable
hypothesis is that PICT is more likely if there
are other females in the group carrying and
caring for live infants (the ‘social facilitation’
hypothesis: Watson and Matsuzawa, 2018).
Function
As reviewed above, in social insects corpses
within the nest are disposed of, and corpses
encountered outside the nest may be avoided.
Active disposal requires time and energy, but
the benefits in terms of intra-colony hygiene
and health mean that corpse management
behaviors have been positively selected
during evolution. In free-ranging nonhuman
primates, a dead group member can be
avoided simply by moving to another part of
the home range and staying away until any
risks associated with the decaying corpse
465
have diminished. Indeed, monkeys and apes
t­
ypically abandon dead individuals either
precisely at or near where they died, after
periods of time ranging from a few minutes
to several hours (e.g., Yang et  al., 2016;
Teleki, 1973), although sometimes individu-
als may revisit the place of death the follow-
ing day (Gonçalves and Carvalho, 2019). As
PICT is clearly non-abandonment, the ques-
tion of function remains. One hypothesis is
that it is simply a non-adaptive error, a by-
product of the combination of mother’s hor-
monal condition and sensitivity to stimulus
features of the dead infant. Another possibil-
ity is that the mother is unaware that her
infant is dead (Gonçalves and Carvalho,
2019). However, possible benefits have also
been proposed. One plausible hypothesis is
that PICT expresses a ‘wait and see’ strategy:
producing an offspring is so costly that it
might be worthwhile for a mother to continue
caring for it; in some cases, it might recover
(Nicolson, 1991; Hrdy, 2000; Watson and
Matsuzawa, 2018). This is precisely what
happened in a case recently described by
Masi (2020). An infant female western
gorilla fell from a tree and hit the ground
after dropping at least 15 m. The infant
appeared totally lifeless to the observers, but
her mother carried her, laid her on the ground
and gently touched her. Over the following
hour the infant slowly regained conscious-
ness and movement, and went on to make a
full recovery.
Some authors have proposed that PICT
functions as a grief-buffering mechanism that
helps the mother come to terms with the emo-
tional distress arising from her infant’s death
(Nicolson, 1991; King, 2013; Gonçalves and
Carvalho, 2019). In the first study to collect
data on both behavioral and hormonal changes
in a female monkey (Japanese macaque), who
kept her dead infant for 20 days, Takeshita
et al. (2020) recorded social withdrawal and
signs of stress during this period, but a rapid
return of glucocorticoids to normal levels.
The authors speculated that PICT helped
reduce stress levels related to the death of the
466 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
infant, although they pointed out the need for
larger samples and more longitudinal data.
Evolution
PICT in one form or another has been observed
in the three major primate groups, namely pro-
simians, New World monkeys, and Old World
monkeys and apes. This broad distribution sug-
gests that a basic form of PICT existed in
ancestral primate species, from the evolution-
ary emergence of strong mother–infant bonds.
Extant prosimian mothers carry and transport
their dead infants less often than anthropoid
primate mothers, likely due to physical con-
straints (Gonçalves and Carvalho, 2019).
Although most accounts of PICT have been
described in macaques, baboons, and chimpan-
zees, these species may be over-represented in
the literature because (1) they are the most
studied both in captivity and the wild, and
(2) they often spend much time on the ground,
where it is often easier to observe them. Das
et  al. (2019) recently reported examples of
PICT in more arboreal primate species, but
more data are required on this issue.
OTHER SPECIES, AND SOME
UNANSWERED QUESTIONS
This attempt to address thanatological behav-
iors from a Tinbergian perspective has drawn
on specific sets of responses in two widely
separated taxonomic groups, namely social
insects and nonhuman primates. Of course, a
more complete evolutionary perspective on
responses to death requires information from
other kinds of animals. For example, the
effects of necromones in several species of
fish have been described; responses include
avoidance of areas associated with dead con-
specifics, and freezing (e.g., Bryer et  al.,
2001; Hussain et  al., 2013; Oliviera et  al.,
2014); the latter study also found evidence of
increased cortisol, indicating stress. The
s­urvival value of these responses appears
clear, preparing the animal to deal with the
potential presence of a predator or other
danger. There is little evidence of any strong
effect of visual cues in the perception of death
in fish; the issue requires more research.
Furthermore, questions about the relative
roles of innateness vs experience in respond-
ing to death cues figure rarely in studies of
aquatic species. For animals raised from birth
or hatched in controlled environments in
which exposure to death is never experienced,
any responses to corpses are likely to be
genetically wired, although how other indi-
viduals react may also exert an influence.
How birds respond to death is also impor-
tant for evolutionary thanatological research.
Although some specialist scavengers may
use odor cues to detect carcasses, others
appear to rely on visual cues (for vultures, see
Grigg et  al., 2017). Research on avian reac-
tions to dead conspecifics has emphasized
visual rather than chemical cues. For exam-
ple, when Western scrub-jays saw the dried
skin and feathers of a dead conspecific near a
feeder, they emitted more loud vocalizations
(‘cacophonous reactions’) and took fewer nuts
than when similar-colored, non-social objects
were presented instead. Playbacks of these
calls attracted other jays to the site. A stuffed
owl elicited similar cacophonous aggrega-
tions and mobbing responses (Iglesias et al.,
2012; see Carlson et  al., 2017, for blue tits’
reactions to a model sparrow hawk holding a
dead blue tit). Iglesias et al. (2014) reported
that dead scrub-jay-size birds of other species
also elicited calling and aggregations and sup-
pressed foraging near the carcass, compared
to dead but smaller birds.
Experiments with American crows have
shown broadly similar responses as scrub-
jays toward dead conspecifics. Also, crows
reacted negatively to the sight of a hawk,
but more intensely to a hawk (or a human)
with a dead crow. They were less respon-
sive to the carcass of a pigeon, and pigeons
also took little notice of dead conspecifics
(Swift and Marzluff, 2015, 2018). In the first
 EVOLUTIONARY PSYCHOLOGY AND THANATOLOGY
study of brain activity in response to death
cues in birds, positron emission tomography
on crows revealed particular activation of
the hippocampal region of the brain as they
watched a human holding a dead crow; other
risk-related stimuli activated other forebrain
regions (Cross et  al., 2013). Brain imag-
ing is likely to be increasingly valuable for
revealing neural circuits involved in perceiv-
ing cues related to death in many species as
well as in humans. As with fish, no studies
on birds have investigated the ontogeny of
responses to carcasses, but some responses
have been linked to plausible functions: loud
vocalizations (alerts others to danger), con-
gregating near the corpse (for identification
of the corpse, detecting possible new forag-
ing and/or mating opportunities), and sup-
pressed feeding (avoids the risk of poisoning
or attack by scavenging predators).
Studies of non-primate mammals are also
important. There are descriptions of responses
to dying and dead conspecifics – sometimes
but not always concerning mothers toward
their offspring – in free-ranging giraffes
(Bercovitch, 2012; Strauss and Muller,
2012), dingoes (Appleby et al., 2013), horses
(Mendonça et  al., 2020), African and Asian
elephants (Douglas-Hamilton et  al., 2006;
Hawley et  al., 2017–2018; Goldenberg and
Wittemyer, 2019; Sharma et  al., 2020), and
various cetacean and other aquatic and semi-
aquatic species (Bearzi et al., 2018; Reggente
et  al., 2018; Inman and Leggett, 2019),
among others (King, 2013), but through no
fault of the observers there is usually a lack of
control over relevant variables, or behavioral
details are incomplete. In a rare experimental
study, without citing any research on chemi-
cal releasers of necrophoresis in insects, Pinel
et al. (1981) extended this line of research to
a mammalian species. In several experiments
they showed that individual rats: (1) used
bedding material to cover the corpse of a
conspecific killed at least 40 hours earlier and
recently placed in the subject’s chamber; (2)
similarly buried an anesthetized rat sprinkled
with either putrescine or cadaverine – two
467
compounds that accumulate in vertebrate
corpses – but not an anesthetized rat that was
not treated with a compound; (3) buried inan-
imate objects treated with the compounds but
not untreated control objects; (4) buried a
compound-treated object or corpse only when
their sense of smell was intact; (5) more read-
ily buried an object treated with putrescine or
cadaverine than other substances: water, urine,
ether, ammonia, or ethanol. The same authors
also referred to a report of food-deprived rats
eating older conspecific corpses rather than
fresher ones (Carr et  al., 1979), suggesting
an alternative corpse-induced response in
the presence of a competing and overriding
motivational state. Prounis and Shields (2013)
conducted field experiments on a community
of small mammals in a forest, and found that
fewer seeds were eaten from food trays that
also contained a mouse corpse than trays that
contained a colored ball instead of a corpse.
In a laboratory study the same authors found
longer latencies to pass a conspecific corpse
than a toy mouse placed in one arm of a
Y-maze, indicating stronger avoidance of the
former. No attempt was made to separate
olfactory and visual cues in this research.
CONCLUSIONS: HUMAN
THANATOLOGY, AND FUTURE
DIRECTIONS
Homicide investigations, demographic con-
sequences of natural disasters, wars, and
epidemics, psychological, medical, and eco-
nomic consequences of bereavement, end-of-
life care, suicide, abortion, development of
death-related concepts, death education, and
cultural variations in funerary practices are
just a few of the issues that fall within the
scope of modern human thanatology (Bradley
et al., 2013; Meagher and Balk, 2013). Here
I briefly discuss human responses to the dead
with specific reference to the two topics used
to introduce the Tinbergian approach, namely
corpse disposal and responses to dead infants.
468 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Corpse Disposal
Like social insects, human societies with a
relatively fixed home base must dispose of
corpses. Depending on cultural/religious, eco-
logical, and economic factors, among others,
the most common methods used today are
cremation and ground burial (in single, shared,
or mass graves). Rarer methods include burial
at sea, and above-ground burial (immure-
ment), leaving the body to the elements and
scavengers (Korpiola and Lahtinen, 2015;
Shimane, 2018). Past methods included mum-
mification followed by storing, and cannibal-
ism. Based on paleoarcheological evidence,
Pettitt (2011) proposed that since the emer-
gence of the earliest hominins, five successive
and partially nested phases of ‘mortuary activ-
ities’ (behavior toward conspecific corpses)
can be recognized. The core phase includes
socially mediated interest in and manipulation
of the corpse, along with context-specific
behavioral displays, and possibly some degree
of corpse mutilation and/or cannibalism,
before abandonment at or near the place of
death. With the emergence of australopithe-
cines and early Homo, the archaic (or devel-
oping) phase sees the continuation of most of
the above features, but also the appearance of
‘funerary caching’, i.e., the use of natural fea-
tures of the environment to deposit or conceal
a corpse, such as a cleft in a rock formation, or
a space below a large boulder. Later, Homo
neanderthalis and Homo sapiens further ritu-
alized behaviors surrounding corpses and
started to deliberately bury them instead of
simply abandoning or caching them. Also, in
this modernizing phase multiple corpses might
be buried in the same place, and material
objects left in the graves. In the modern phase
Homo sapiens further elaborated upon the
already existing ‘social theatre’ around the
corpse, and probably left signs to mark or
commemorate burial places. In the advanced
phase, as humans spread across the world,
greater regional and cultural variability in
funerary activities and corpse-disposal meth-
ods appeared. Additionally, communities
increasingly recognized ‘places of the dead’
(cemeteries) and collective representations of
death and the dead became more elaborate.
Across the wide cultural variations in corpse-
disposal methods in human societies, the bio-
logical function of disposal remains clear:
reduction of the risks of disease.
Concerning the immediate factors that move
humans to start engaging in corpse disposal,
visual and physical cues (e.g., obvious non-
survivable injuries, absence of responsiveness
and agency, coldness, rigor mortis, absence of
pulse, breath) are often sufficient to lead to a
declaration of death, but often medical tech-
nology (e.g., vital signs monitors) is used to
confirm death, after which mortuary activi-
ties commence. If the corpse is discovered
only after some delay, then odor cues might
give an added sense of urgency to disposal. A
wide range of volatile organic compounds are
associated with putrefaction of human bodies,
which in various combinations give rise to the
‘odor of death’ (Verheggen et al., 2017). One
such compound is putrescine. Revulsion is a
widespread human reaction to the putrefy-
ing odor of a corpse – human or nonhuman –
and often leads to one or more actions such
as covering one’s nose and mouth, leaving the
scene, masking the odor with another, and/or
disposing of the corpse. Perhaps surprisingly,
although work has been done on the actions
of putrescine and cadaverine at the molecu-
lar level in humans (Izquierdo et  al., 2018),
there are few scientific studies of humans’
behavioral reactions to necromones. In a
rare experimental study, Wisman and Shrira
(2015) focused on the threat-signal property
of putrescine and hypothesized that it would
affect people’s behavior even if they were
not consciously aware of its presence. They
showed that putrescine caused (1) increased
vigilance: reaction times were faster in partici-
pants who sniffed a jar containing putrescine
before completing a computerized target-
detection task than in participants who sniffed
an equally ‘repugnant’ but non-death-related
odor (ammonia); (2) ‘escape’: participants
asked to smell putrescine and then respond to
 EVOLUTIONARY PSYCHOLOGY AND THANATOLOGY
three simple questions walked away from the
test area faster than those similarly exposed to
ammonia. Participants exposed to putrescine
also completed unfinished words by produc-
ing more escape- and threat-related words
than ammonia-exposed participants. Although
not directly related to death, these experimen-
tally induced responses are consistent with
putrescine’s ability to trigger defensive reac-
tions in humans.
Regarding ontogeny of corpse disposal
in humans, it seems clear that children’s
views about death and how corpses should
be treated are strongly culturally influ-
enced (Longbottom and Slaughter, 2018;
Panagiotaki et  al., 2018). However, little
information is available on the development
of children’s ideas specifically about ways to
dispose of corpses, and what factors might
influence these ideas. Similarly, although
studies have looked at how they respond to
visual depictions of death, such as scenarios
that show signs of injury or fatal attacks (e.g.,
Barrett and Behne, 2005), studies of young
children’s reactions to olfactory cues that
adults associate with death are lacking.
Mothers and Dead Infants
Of course, human mothers grieve when their
offspring dies. Although they show nothing
as obvious or as widespread as nonhuman
primate PICT, it is common for parents to
keep reminders of their deceased offspring,
such as toys or clothes (Foster et al., 2011);
this is a normal behavior that might only
become problematic in some cases of pro-
longed, pathological grief (Gorer, 1965). In
cases of stillbirth or peri-natal death, in some
societies mothers may desire to spend time in
physical contact with their baby, before
funerary arrangements are begun. From a
functional perspective, this period of contact
could be seen as helping mothers to cope
with the emotional trauma of losing their
infant (Rådestad et al., 2009), possibly buff-
ering against maternal depression (Ryninks
469
et  al., 2014). However, some studies have
reported contradictory findings and substan-
tial individual differences in how mothers
respond to contact with their dead baby; one
influencing factor may be the manner in
which the baby is presented to the mother
(see Erlandsson et al., 2012; Redshaw et al.,
2016). There is currently no consensus
regarding the psychological outcomes of
holding a stillborn baby.
With regard to proximate influences, both
visual and tactile input from the dead baby
seem likely to be important in triggering
positive feelings in the mother, especially
if the baby is still warm, shortly after death
(Rådestad et al., 2009). However, the visual
appearance – real or imagined – of the baby
can influence the parents’ decision to view
the baby (Sun et  al., 2014) and their expe-
rience of contact (Ryninks et  al., 2014).
Ontogenetic influences will clearly matter
in such complex situations, but we are still
far from identifying specific developmental
processes underlying how individual human
mothers respond to dead babies. From an
evolutionary perspective, however, it is easy
to see parallels between the emotional and
behavioral consequences of loss of an infant
in human parents and nonhuman primate
mothers (and possibly fathers, in some spe-
cies). The ‘vigilance in grief’ phenomenon
involving heightened sensitivity to any infor-
mation associated with the deceased has
been suggested to psychologically prepare
the bereaved individual for a possible viable
reunion with the former (White and Fessler,
2018). This recalls the ‘wait and see’ hypoth-
esis of PICT in nonhuman primate and ceta-
cean mothers, discussed above. Furthermore,
in many cultures it is common for bereaved
individuals to physically contact the corpse
not only of newborns but also older kin –
sometimes over prolonged periods – despite
the potential risk of disease. Based on data
from 57 cultures, however, the risk of con-
tamination from mortuary rituals involving
physical contact appeared lower than some
normative forms of contact with the living
470 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
(Murray et al., 2017), and White et al. (2017)
concluded that ritual exposure to corpses
facilitated progress through and recovery
from grieving.
Future Directions
Comparative evolutionary thanatology needs
more data on a wide range of species, and
from observational and, where ethically
acceptable, experimental approaches. Given
the close evolutionary relationships between
humans and the great apes in brain develop-
ment and behavior, Pettitt and Anderson
(2020) recently proposed some common goals
and terminology as a way of fostering better
communication and collaboration between
thanatology-oriented paleontologists and pri-
matologists. They also listed some questions
that, if addressed by primatologists, would
help in clarifying stages of ancestral human
mortuary activities. Evolutionary thanatology
can be seen as a more general framework for
unifying the study of death and dying across a
wider range of species (Anderson et al., 2018).
There is little doubt that some behaviors
shown toward corpses in ancestral humans
(see Pettitt, 2011) are seen to various extents
in modern nonhuman primates, including
gathering around and manipulating corpses,
mutilation, and cannibalism. Simple aban-
donment of the corpse is the norm in pri-
mates – albeit delayed in some cases of
PICT – but rare cases of partially covering
corpses with vegetation have been reported in
chimpanzees (Boesch, 2012). This behavior
(dropping vegetation on the corpse) has only
been reported at one field site, and it is not
clear precisely what motivates it. Re-visiting
the site where a group member has recently
died has been reported in several species
(Gonçalves and Carvalho, 2019), but do sur-
vivors return to that location more frequently
or for longer than expected based on their
normal ranging patterns? Does their behav-
ior there change noticeably, for example in
any ways that might indicate recognition
or commemoration? Do the reactions of
­primates to the corpse vary according to the
social status of the individual when it was
alive? (see Porter et  al., 2019, for possible
examples in wild mountain gorillas). Why do
primate mothers persist in PICT when their
infant’s corpse smells foul? Is putrescine less
repulsive for nonhuman primates than for
humans?
Much more research is required before we
can give confident answers to these and other
relevant questions about reactions to the dead
in primates and other species. Tinbergen’s
questions provide an excellent framework for
studying how and why responses to the dead
may converge or diverge not just across but
also within species. Of course, despite often
being considered independently, the four
questions are complementary, and answers
to one can lead to insights relevant to oth-
ers. Given the nature of the general topic, the
typical lack of control over circumstances
and the lack of preparedness to record events
(in primates), all four questions will continue
to be valuable. By addressing them, we can
hope to establish an evolutionarily informed
thanatology that, as set out by Lewis et  al.
(2017), will (a) generate testable hypotheses,
(b) test empirical predictions, and (c) allow
plausible interpretations.
ACKNOWLEDGMENTS
I thank the editors for their help with the
manuscript, and an anonymous reviewer for
valuable comments and suggestions. While
writing the chapter I was supported by JSPS
KAKENHI Grant No. 18K18693.
REFERENCES
Alley TR (1980) Infantile colouration as an
elicitor of caretaking behavior in Old World
primates. Primates 21:416–429
 EVOLUTIONARY PSYCHOLOGY AND THANATOLOGY
Altmann J (1980) Baboon mothers and
Infants. Cambridge, MA: Harvard University
Press
Anderson JR (2016) Comparative thanatology.
Current Biology 26:R553–R556
Anderson JR (2017) Comparative evolutionary
thanatology of grief, with special reference
to nonhuman primates. Japanese Review of
Cultural Anthropology 18:173–189
Anderson JR, Andre E, Wolf P (1995) Successful
mother- and group-rearing of a newborn
capuchin monkey (Cebus capucinus) follow-
ing emergency major surgery. Animal Wel-
fare 4:171–182
Anderson JR, Biro D, Pettitt P (2018) Evolutionary
thanatology. Philosophical Transactions of the
Royal Society B 373:20170262 http://dx.doi.
org/10.1098/rstb.2017.0262
Appleby R, Smith B, Jones D (2013) Observa-
tions of a free-ranging adult female dingo
(Canis dingo) and littermates’ responses to
the death of a pup. Behavioural Processes
96:42–46
Arathi HS, Burns I, Spivak M (2000) Ethology of
hygienic behavior in the honey bee Apis mel-
lifera L. (Hymenoptera: Apidae): behavioural
repertoire of hygienic bees. Ethology
106:365–379
Barrett HC, Behne T (2005) Children’s under-
standing of death as the cessation of agency:
a test using sleep versus death. Cognition
96:93–108
Barrett L, Blumstein DT, Clutton-Brock TH,
Kappeler PM (2013) Taking note of
Tinbergen, or: the promise of a biology of
behavior. Philosophical Transactions of the
Royal Society B 36820120352 http://dx.doi.
org/10.1098/rstb.2012.0352
Bateson P, Laland KN (2013) Tinbergen’s four
questions: an appreciation and an update.
Trends in Ecology & Evolution 28:712–718
Bearzi G, Kerem D, Furey NB, Pitman RL, Ren-
dell L, Reeves RR (2018) Whale and dolphin
behavioural responses to dead conspecifics.
Zoology 128:1–15
Bercovitch FB (2012) Giraffe cow reaction to
the death of her newborn calf. African Jour-
nal of Ecology 51:376–379
Bercovitch FB (2019) A comparative perspective
on the evolution of mammalian reactions to
dead conspecifics. Primates 61:21–28 https://
doi.org/10.1007/s10329-019-00722-3
471
Biro D, Humle T, Koops K, Sousa C, Hayashi M,
Matsuzawa T (2010) Chimpanzee mothers at
Bossou, Guinea carry the mummified remains
of their dead infants. Current Biology
20:R351–R352
Boesch C (2012) Wild cultures: a comparison
between chimpanzee and human cultures.
Cambridge: Cambridge University Press
Bonoti F, Leondari A, Mastora A (2013) Explor-
ing children’s understanding of death:
through drawings and the Death Concept
Questionnaire. Death Studies 37:47–60
Bradley B, Feldman F, Johansson J (2013) The
Oxford handbook of philosophy of death.
New York: Oxford University Press
Brent SB, Speece MW, Lin C, Dong Q, Yang C
(1996) The development of the concept of
death among Chinese and U.S. children 7–13
years of age: from binary to ‘fuzzy’ concepts?
Omega 33:67–83
Brown MW, Norris ME (2004) Survivorship
advantage of conspecific necrophagy in
overwintering boxelder bugs (Heteroptera:
Rhopalidae). Annals of the Entomological
Society of America 97:500–503
Bryer PJ, Mira RS, Chivers DP (2001) Chemo­
sensory assessment of predation risk by slimy
sculpins (Cottus cognatus): responses to
alarm, disturbance, and predator cues. J
Journal of Chemical Ecology 27:533–546
Carlson NV, Pargeter HM, Templeton CN (2017)
Sparrowhawk movement, calling, and pres-
ence of dead conspecifics differentially
impact blue tit (Cyanistes caeruleus) vocal
and behavioral mobbing responses. Behavio-
ral Ecology and Sociobiology 71:133 DOI
https://doi.org/10.1007/s00265-017-2361-x
Carr WJ, Landauer MR, Wiese RE, Morasco E,
Thor DH (1979) A natural food aversion in
Norway rats. J Journal of Comparative and
Physiological Psychology 89:574–584
Choe DH, Millar JG, Rust MK (2009) Chemical
signals associated with life inhibit necropho-
resis in Argentine ants. Proceedings of the
National Academy of Sciences USA
106:8251–8255
Cross DJ, Marzluff JM, Palmquist I, Minoshima S,
Shimizu T, Miyaoka R (2013) Distinct neural
circuits underlie assessment of a diversity of
natural dangers by American crows. Proceed-
ings of the Royal Society B 280:20131046
doi:10.1098/rspb.2013.1046
472 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Das S, Erinjery JJ, Desai N, Mohan K, Kumara
HN, Singh M (2019) Deceased infant carry-
ing in nonhuman anthropoids: insights from
systematic analysis and case studies of
bonnet macaques (Macaca radiata) and lion-
tailed macaques (Macaca silenus). Journal of
Comparative Psychology 133:156–170
Dellatore DF, Waitt CD, Foitova I (2009) Two
cases of mother-infant cannibalism in orang­
utans. Primates 50:277–281
De Marco A, Cozzolino R, Thierry B (2018)
Prolonged transport and cannibalism of
mummified infant remains by a Tonkean
macaque mother. Primates 59:55–59
Diez L, Lejeune P, Detrain C (2014) Keep the
nest clean: survival strategies of corpse
removal in ants. Biology Letters 10:20140306
http://dex.doi.org/10.1098/rsbl.2014.0306
Douglas-Hamilton I, Bhalla S, Wittemeyer G,
Vollrath F (1986) Behavioural reactions of
elephants towards a dying and deceased
matriarch. Applied Animal Behaviour Science
100:87–102
Engh AL, Beehner JC, Bergman TJ, Whitten PL,
Hoffmeier RR, Seyfarth RM, Cheney DL
(2006) Behavioural and hormonal responses
to predation in female chacma baboons
(Papio hamadryas ursinus). Proceedings of
the Royal Society B, 273:707–712
Erlandsson K, Warland J, Cacciatore J, Rådestad
I (2012) Seeing and holding a stillborn baby:
mothers’ feelings in relation to how their
babies were presented to them after birth –
Findings from an online questionnaire. Mid-
wifery 29:246–250
Fairbanks LA (1996) Individual differences in
maternal style: causes and consequences for
mothers and offspring. Advances in the
Study of Behavior 25:579–611
Fashing PJ, Nguyen N, Barry TS, Goodale CB,
Burke RJ, Jones SCZ, Kerby JT, Lee LM, Nurmi
NO, Venkataraman VV (2011) Death among
geladas (Theropithecus gelada): a broader
perspective on mummified infants and primate
thanatology. American Journal of Primatology
73:405–409
Fedurek P, Tkaczynski P, Asiimwe C, Hobaiter C,
Samuni L, Lowe AE, Dijrian AG, Zuberbühler
K, Wittig RM, Crockford C (2019) Maternal
cannibalism in two populations of wild chim-
panzees. Primates 61:181–187 https://doi.
org/10.1007/s10329-019-00765-6
Foster TL, Giler MJ, Davies B, Dietrich MS, Bar-
rera M, Fairclough DL, Vannatt K, Gerhardt
CA (2011) Comparison of continuing bonds
reported by parents and siblings after a
child’s death from cancer. Death Studies
35:420–440
Frederickx C, Dekeirsschieter J, Verheggen FJ,
Haubruge E (2012) Responses of Lucilia seri-
cata Meigen (Diptera: Calliphoridae) to cadav-
eric volatile organic compounds. Journal of
Forensic Science 57:386–390 doi: 10.1111/
j.1556-4029.2011.s02010.x
Gire J (2014) How death imitates life: cultural
influences on conceptions of death and
dying. Online Readings in Psychology and
Culture 6(2) https://doi.org/10.9707/2307-
0919.1120
Goldenberg SZ, Wittemyer G (2019) Elephant
behavior toward the dead: A review and
insights from field observations. Primates 61,
119–128.
Gonçalves A, Biro D (2018) Comparative thana-
tology, and integrative approach: exploring
sensory/cognitive aspects of death recogni-
tion in vertebrates and invertebrates. Philo-
sophical Transactions of the Royal Society B
373:20170263 http://dx.doi.org/10.1098/
rstb.2017.0263
Gonçalves A, Carvalho S (2019) Death among
primates: a critical review of non-human
primate interactions towards their dead
and dying. Biological Reviews
94:1502–1529
Goodall J (1983) Population dynamics during a
15-year period in one community of free-
living chimpanzees in the Gombe National
Park, Tanzania. Zeitschrift für Tierpsychologie
61:1–60
Gorer G (1965) Death, grief and mourning in
contemporary Britain. Garden City, NY:
Doubleday
Grigg NP, Krilow JM, Gutierrez-Ibanez C, Wylie
DR, Graves GR, Iwaniuk AN (2017) Anatomi-
cal evidence for scent guided foraging in the
turkey vulture. Scientific Reports 7:17408
doi:10.1038/s41598-017-17794-0
Harlow HF (1974) Learning to love. New York:
Jason Aronson
Hawley CR, Beirne C, Meier A, Poulsen JR
(2017–2018) Conspecific investigation of a
deceased forest elephant (Loxodonta cyclo-
tis). Pachyderm 59:97–100
 EVOLUTIONARY PSYCHOLOGY AND THANATOLOGY
Hayashi M, Matsuzawa T (2017) Mother-infant
interactions in captive and wild chimpan-
zees. Infant Behavior and Development
48:20–29
Hrdy SB (2000) Mother nature. London:
Random House
Hussain A, Saraiva LR, Ferrero DM, Ahuja G,
Krishna VS, Liberles SD, Korsching SI (2013)
High-affinity olfactory receptor for the
death-associated odor cadaverine. Proceed-
ings of the National Academy of Sciences
USA 110:19579–19584
Iglesias TL, McElreath R, Patricelli GL (2012)
Western scrub-jay funerals: cacophonous
aggregations in response to dead conspecif-
ics. Animal Behaviour 84:1103–1111
Iglesias TL, Stetkevich RC, Patricelli GL (2014)
Dead heterospecifics as cues of risk in the
environment: Does size affect response?
Behaviour 151:1–22
Inman V, Leggett EA (2019) Observations on
the response of a pod of hippos to a dead
juvenile hippo (Hippopotamus amphibious, Lin-
naeus 1758) African Journal of Ecology
doi:10.1111/aje.12644
Izquierdo C, Gómez-Tamayo JC, Nebel JC,
Pardo L, Gonzalez A (2018) Identifying
human diamine sensors for death related
putrescine and cadaverine molecules. PLos
Computational Biology 14(1):e1005945
https://doi.prg/10.1371/journal.
pcbi.1005945
Julian GE, Cahan S (1999) Undertaking spe-
cialization in the desert leaf-cutter ant Acro-
myrmex versicolor. Animal Behaviour
58:437–442
Kapheim KM (2019) Synthesis of Tinbergen’s
four questions and the future of socio­
genomics. Behavioral Ecology and Sociobiol-
ogy 73:186
King BJ (2013) How animals grieve. Chicago:
University of Chicago Press
Korpiola M, Lahtinen A (2015) Cultures of
death and dying in medieval and early
modern Europe. Helsinki: Helsinki Collegium
for Advanced Studies
Kristal MB (2009) The biopsychology of mater-
nal behavior in nonhuman mammals. ILAR
Journal 50:51–63
Lewis DMG, Al-Shawaf L, Conroy-Beam D,
Asao K, Buss DM (2017) Evolutionary
473
psychology: a how-to guide. American
Psychologist 72:353–373
Longbottom S, Slaughter V (2018) Sources of
children’s knowledge about death and dying.
Philosophical Transactions of the Royal Soci-
ety B 373:20170267 http://dx.doi.
org/10.1098/rstb.2017.0267
Lonsdorf EV (2017) Sex differences in nonhu-
man primate behavioral development. Jour-
nal of Neuroscience Research 95:213–221
López-Riquelme GO, Fanjul-Moles ML (2013) The
funeral ways of social insects: social strategies
for corpse disposal. Trends in Entomology
9:71–129
Maestripieri D (2018) Maternal influences in
primate social development. Behavioral Ecol-
ogy and Sociobiology 72:130 https://doi.
org/10.1007/s00265-018-2547-x
Masi S (2020) Reaction to allospecific death and
to an unanimated gorilla infant in wild west-
ern gorillas: insights into death recognition
and prolonged maternal carrying. Primates
61:83–92 https://doi.org/10.1007/s10329-
019-00745-w
Matsumoto T, Ito N, Inoue S, Nakamura M
(2016) An observation of a severely disabled
infant chimpanzee in the wild and her inter-
actions with her mother. Primates 57:3–7
Meagher DK, Balk DE (2013) Handbook of
thanatology, 2nd edition. New York:
Routledge
Mendonça RS, Ringhofer M, Pinto P, Inoue S,
Hirata S (2020) Feral horses’ (Equus ferus
caballus) behavior toward dying and dead
conspecifics. Primates 61:49–54 https://doi.
org/10.1007/s10329-019-00728-x
Munday Z, Brown MJF (2018) Bring out your
dead: quantifying corpse removal in Bombus
terrestris, an annual eusocial insect. Animal
Behaviour 138:51–57
Murray DR, Fessler DMT, Kerry N, White C,
Marin M (2017) The kiss of death: three tests
of the relationship between disease threat
and ritualized physical contact within tradi-
tional cultures. Evolution and Human Behav-
ior 38:63–70
Nakamichi M (1986) Behavior of infant
Japanese monkeys (Macaca fuscata) with
congenital limb malformations during their
first three months. Developmental
Psychobiology 19:335–341
474 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Neoh K-B, Yeap B-K, Tsunoda K, Yoshimura T,
Lee C-Y (2012) Do termites avoid carcasses?
Behavioral responses depend on the nature
of the carcasses. PLoS ONE 7(4):e36375
doi:10.1371/journal.pone.0036375
Nicolson NA (1987) Infants, mothers, and other
females. In: Smuts BB, Cheney DL, Seyfarth
RM, Wrangham RW, Struhsaker TT (Eds.),
Primate Societies. Chicago: University of Chi-
cago Press, pp. 330–342
Nicolson NA (1991) Maternal behavior in human
and nonhuman primates. In: Loy JD, Peters
CB (Eds.), Understanding Behavior. New York:
Oxford University Press, pp. 17–50
Oldroyd BP, Thompson GJ (2007) Behavioural
genetics of the honey bee Apis mellifera.
Advances in Insect Physiology 33:1–49
Oliviera TA, Koakoski G, Costa da Mota A,
Piato AL, Barreto RE, Volpato GL, Barcellos
LJG (2014) Death-associated odors induce
stress in zebrafish. Hormones and Behavior
65:340–344
Owren MJ, Dieter JA (1989) Infant cross-­
fostering between Japanese (Macaca fuscata)
and rhesus macaques (M. mulatta). Ameri-
can Journal of Primatology 18:245–250
Panagiotaki G, Hopkins M, Nobes G, Ward E,
Griffiths D (2018) Children’s and adults’
understanding of death: cognitive, parental,
and experiential influences. Journal of Exper-
imental Child Psychology 166:96–115
Panagiotaki G, Nobes G, Ashraf A, Aubby H
(2015) British and Pakistani children’s under-
standing of death: cultural and developmental
influences. British Journal of Developmental
Psychology 33:31–44
Pettitt P (2011) The Palaeolithic origins of
human burial. London: Routledge
Pettitt P (2018) Hominin evolutionary thanatol-
ogy from the mortuary to funerary realm.
The palaeoanthropological bridge between
chemistry and culture. “Philosophical Trans-
actions of the Royal Society B 373 https://
doi.org/10.1098/rstb.2018.0212
Pettitt P, Anderson JR (2020) Primate thanatol-
ogy an hominoid mortuary archeology. Pri-
mates 61:9–19 https://doi.org/10.1007/
s10329-019-00769-2
Pinel JPJ, Gorzalka BB, Ladak F (1981)
Cadaverine and putrescine initiate the burial
of dead conspecifics by rats. Physiology &
Behavior 27:819–824
Porter A, Eckardt W, Vecellio V, Guschanski K,
Niehoff PP, Ngobobo-As-Ibungu U, Pekeyake
RN, Stoinski T, Caillaud D (2019) Behavioral
responses around conspecific corpses in adult
eastern gorillas (Gorilla beringei spp.) PeerJ
7:e6655 http://doi.org/10.7717/peerj.6655
Prounis GS, Shields WM (2013) Necrophobic
behavior in small mammals. Behavioural Pro-
cesses 94:41–44
Rådestad I, Säflund K, Wredling R, Oneläv E,
Steineck G (2009) Holding a stillborn baby:
mothers’ feelings of tenderness and grief.
British Journal of Midwifery 17:178–180
Redshaw M, Hennegan JM, Henderson J (2016)
Impact of holding the baby following still-
birth on maternal health and well-being:
findings from a national survey. BMJ Open
6:e010996 doi:10.1136/bmjopen-2015-
010996
Reggente MALV, Papale E, McGinty N, Eddy L,
de Lucia GA, Bertulli CG (2018) Social rela-
tionships and death-related behavior in
aquatic mammals: a systematic review. Phil-
osophical Transactions of the Royal Society B
373:20170260 http://dx.doi.org/10.1098/
rstb.2017.0260
Renucci M, Tirard A, Provost E (2011) Complex
undertaking behavior in Temnothorax
lichtensteini ant colonies: from corpse-burying
behavior to necrophoric behavior. Insectes
Sociaux 58:9–16
Rhine RJ, Norton GW, Roertgen WJ, Klein HD
(1980) The brief survival of free-ranging
baboon infants (Papio cynocephalus) after
separation from their mothers. International
Journal of Primatology 1:401–409
Robinson GE, Page RE (1988) Genetic determi-
nation of guarding and undertaking in
honey-bee colonies. Nature 333:356–358
Rollo CD, Czyzewska E, Borden JH (1994) Fatty
acid necromones for cockroaches. Naturwis-
senschaften 81:409–410
Rothenbuhler WC (1964) Behaviour genetics of
nest cleaning in honey bees. I. Responses of
four inbred lines to disease-killed brood.
Animal Behaviour 12:578–583
Ryninks K, Roberts-Collins C, McKenzie-
McHarg K, Horsch A (2014) Mothers’ experi-
ence of their contact with their stillborn infant:
an interpretative phenomenological analysis.
BMC Pregnancy & Childbirth 14:203 www.
biomedcentral.com/1471–2393/14/203
 EVOLUTIONARY PSYCHOLOGY AND THANATOLOGY
Saltzman W, Maestripieri D (2011) The neu-
roendocrinology of primate maternal behav-
ior. Progress in Neuro-Psychopharmacology
and Biological Psychiatry 35:1192–1204
Sato A, Koda H, Lemasson A, Nagumo S, Masa-
taka N (2012) Visual recognition of age class
and preference for infantile features: implica-
tions for species-specific vs universal cognitive
traits in primates. PLoS ONE 7(5):38387
doi:10.1371/journal.pone.0038387
Sharma N, Pokharel SS, Kohshima S, Sukumar
R (2020) Behavioural responses of free-
ranging Asian elephants (Elephas maximus)
towards dying and dead conspecifics. Pri-
mates 61:129–138 https://doi.org/10.1007/
s10329-1089-00739-8
Shimane K (2018) Social bonds with the dead:
how funerals transformed in the twentieth
and twenty-first centuries. Philosophical
Transactions of the Royal Society B
373:20170274 doi:10.1098/rstb.2017.0274
Slaughter V, Griffiths M (2007) Death under-
standing and fear of death in young children.
Clinical Child Psychology and Psychiatry
12:525–535
Smith S (1986) Infant cross-fostering in rhesus
monkeys (Macaca mulatta): a procedure for
the long-term management of captive popu-
lations. American Journal of Primatology
11:229–237
Speece MW, Brent SB (1984) Children’s under-
standing of death: a review of three compo-
nents of a death concept. Child Development
55:1671–1686
Strauss MKL, Muller Z (2012) Giraffe mothers
in East Africa linger for days near the remains
of their dead calves. African Journal of Ecol-
ogy 51:506–509
Sugiyama Y, Kurita H, Matsui T, Kimoto S, Shi-
momura T (2009) Carrying of dead infants
by Japanese macaque (Macaca fuscata)
mothers. Anthropological Science
117:113–119
Sullivan, EC, Mondoza SP, Capitanio JP (2011)
Similarity in temperament between mother
and offspring rhesus monkeys: sex differ-
ences and role of monoamine oxidase-A and
serotonin transporter promoter polymor-
phism genotypes. Developmental Psychobi-
ology 53:549–563
Sun JC, Rei W, Sheu SJ (2014) Seeing or not
seeing: Taiwan’s parents’ experiences during
475
stillbirth. International Journal of Nursing
Studies 51:1153–1159
Sun Q, Haynes KF, Zhou X (2013) Differential
undertaking response of a lower termite to
congeneric and conspecific corpses. Scien-
tific Reports 3:1650 doi: 10.1038/srep01650
Sun Q, Haynes KF, Zhou X (2017) Dynamic
changes in death cues modulate risks and
rewards of corpse management in a social
insect. Functional Ecology 31:697–706
Sun Q, Haynes KF, Zhou X (2018) Managing
the risks and rewards of death in eusocial
insects. Philosophical Transactions of the
Royal Society B 373:20170258
Sun Q, Zhou X (2013) Corpse management in
social insects. International Journal of Bio-
logical Sciences. 9:313–321
Swift K, Marzluff JM (2018) Occurrence and
variability of tactile interactions between
wild American crows and dead conspecifics.
Philosophical Transactions of the Royal Soci-
ety B 373:20170259 http://dx.doi.
org/10.1098/rstb.2017.0259
Swift KN, Marzluff JM (2015) Wild American
crows gather around their dead to learn
about danger. Animal Behaviour
109:187–197
Taborsky M (2014) Tribute to Tinbergen: the
four problems of biology. A critical appraisal.
Ethology 120:224–227
Takeshita RSC, Huffman MA, Kinoshit K, Berco-
vitch FB (2020) Changes in social behavior
and fecal glucocorticoids in a Japanese
macaque (Macaca fuscata) carrying her dead
infant. Primates 61:35–45 https://doi.
org/10.1007/s10329-019-00753-w
Tartabini A (1991) Mother-infant cannibalism in
thick-tailed bushbabies (Galago crassicauda-
tus umbrosus). Primates 32:379–383
Teleki G (1973) Group response to the acciden-
tal death of a chimpanzee in Gombe National
Park, Tanzania. Folia Primatologica 20:81–94
Thierry B, Anderson JR (1986) Adoption in
anthropoid primates. International Journal of
Primatology 7:191–216
Tinbergen N (1963) On aims and methods in
Ethology. Zeitschrift für Tierpsychologie
20:410–433
Tokuyama N, Moore DL, Graham KE, Lokasola
A, Furuichi T (2017) Cases of maternal can-
nibalism in wild bonobos (Pan paniscus) from
two different field sites, Wamba and
476 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Kokolopori, Democratic Republic of the
Congo. Primates 58:7–12
Trapanese C, Bey M, Tonachella G Meunier H,
Masi S (2020) Prolonged care and cannibalism
of infant corpse by relatives in semi-free-­
ranging capuchin monkeys. Primates 61:41–47
https://doi.org/10.1007/s10329-019-00747-8
Trumbo ST, Huang ZY, Robinson GE (1997) Divi-
sion of labor between undertaker specialists
and other middle-aged workers in honey bee
colonies. Behavioral Ecology and Sociobiol-
ogy 41:151–163
Trumbo ST, Robinson GE (1997) Learning and
task interference by corpse-removal specialists
in honey bee colonies. Ethology 103:966–975
Valdes L, Laidre ME (2019) Scent of death:
Evolution from sea to land of an extreme
collective attraction to conspecific death.
Ecology and Evolution 9:2171–2179
van Lawick-Goodall J (1968) The behavior of
free-living chimpanzees in the Gombe Stream
Reserve. Animal Behaviour Monographs
1:161–311
van Noordwijk MA (2012) From maternal
investment to lifetime maternal care. In:
Mitani JC, Call J, Kappeler PM, Palombit RA,
Silk JB (Eds.), The Evolution of Primate Socie-
ties. Chicago: University of Chicago Press,
pp. 321–342
Verheggen F, Perrault KA, Megido RC, Duboid
LM, Francis F, Haubruge E, Forbes SL, Focant
JF, Stefanuto PH (2017) The odor of death:
an overview of current knowledge on char-
acterization and applications. Bioscience
67:600–613
Visscher PK (1983) The honey bee way of
death: necrophoric behavior in Apis mellifera
colonies. Animal Behaviour 31:1070–1076
Watson CFI, Matsuzawa T (2018) Behaviour of
nonhuman primate mothers toward their
dead infants: uncovering mechanisms. Philo-
sophical Transactions of the Royal Society B
373:20170261 http://dx.doi.org/
10.1098/rstb.2017.0261
White C, Fessler DMT (2018) An evolutionary
account of vigilance in grief. Evolution,
Medicine, and Public Health 2018 (1):34–42
doi:10.1093/emph/eox018
White C, Marin M, Fessler DMT (2017) Not just
dead meat: an evolutionary account of
corpse treatment in mortuary rituals. Journal
of Cognition and Culture 17:146–168
Wilson, EO, Durlach, NI, Roth, LM (1958)
Chemical releaser of necrophoric behavior in
ants. Psyche 65:108–114
Wisman A, Shrira I (2015) The smell of death:
evidence that putrescine elicits threat man-
agement mechanisms. Frontiers in Psychol-
ogy 6:1274 doi: 10.33389/fpsyg.2015.0127
Worlein JM (2014) Nonhuman primate models
of depression: effects of early experience
and stress, ILAR Journal 55:259–273
Yang B, Anderson JR, Li BG (2016) Tending a
dying adult in a wild multi-level primate soci-
ety. Current Biology 26:R403–R404
Yao M, Rosenfeld J, Attridge S, Sidhu S,
Askenov V, Rollo CD (2009) The ancient
chemistry of avoiding risks of predation and
disease. Evolutionary Biology 36:267–281

Evolutionary Psychology and
Reproduction
S t e p h e n W h y t e a n d B e n n o To r g l e r
SEXUAL REPRODUCTION AND
MATING
The evolutionary innovation of sexual repro-
duction allows organisms to evolve rapidly
and thrive through genome reshuffling (Ball,
2019), results in increased variation, and was
proposed over 100 years ago by August
Weismann. ‘Such differences’, Weismann
explained, ‘afford the material by means of
which natural selection is able to increase or
weaken each character according to the needs
of the species’ (as cited by Otto, 2009: S2).
For mammals, reproduction means that a
new individual organism emerges that,
although physically and biologically attached
to the female, is a mix of genetic variations
from two parent organisms. Thus, unlike
other biological classes, mammals reproduce
sexually rather than self-cloning through the
asexual methods of binary fusion,1 fragmen-
tation, budding, parthenogenesis, and gem-
mulation. The advantages of sexual
24
reproduction have long been of interest to
biologists, with Maynard Smith (1971) iden-
tifying as the most important the ability to
adapt to a novel environment that favors a
new gene combination:
If two different populations have over a long
period adapted to two different environments,
they will differ genetically at a number of loci; one
may be ABC…L and the other abc…l. Suppose
now a new environment becomes available for
colonization, in which the best adapted genotype
is, say AbC…l. If founders from both original
populations invade the new environment, then
evolutionary adaptation will be faster by several
orders of magnitude if the founders can hybridize
sexually. (Smith, 1971: 334)
Adaptation to this new environment in turn
increases the rate of evolutionary adaptation
(Becks and Agrawal, 2012),2 so that even
though the diverse array of sexually produced
genotypes results in a lower average fitness of
offspring, these well-adapted genotypes con-
tribute disproportionately to future genera-
tions. In general, then, sex is a bet-hedging
478 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
strategy that can result in offspring with a
higher geometric mean fitness in a context of
continual environmental change (Lively and
Morran, 2014).
When such change is overly rapid, how-
ever, path-dependent genetic associations
that build upon past selection can become
detrimental to survival (Otto, 2009), so that a
population of asexual organisms may pick up
mutations that slow their growth process and
drag down the entire population (Zimmer,
2009). On the other hand, when selection var-
ies over time and space, sexual reproduction
is an evolutionary advantage (Otto, 2009), as
supported by experimental evidence of co-
evolutionary host-parasite interactions lead-
ing to long-term persistence of sex (Lively
and Morran, 2014). Hence, consistent with
the Red Queen’s observation in Through the
Looking-Glass, ‘(Carol, 1917)… it takes all
the running you can do, to keep in the same
place’ and species continually evolve sim-
ply to maintain their position with respect to
other competing, evolving species.
Yet despite its advantages, sexual repro-
duction is challenging and costly, particu-
larly when males contribute nothing more
than genes. For example, in any given gen-
eration, a sexual population produces half the
number of offspring generated by an asexual
population of the same size, a potential one-
half reduction in birth rate that makes the
sexual population more likely to become
extinct (Hadany and Comeron, 2008; Lively
and Morran, 2014). This population is thus
obliged to produce twice as many offspring
to avoid transmission disadvantage (Otto,
2009), which Maynard Smith (1978) labelled
the ‘two-fold cost of males’. Completing the
genome through the matching of one gamete
(egg) to another (sperm) at the right time and
place also requires a dating exercise to find
and attract a potential partner. This mating
itself is energy consuming and subject to such
risks as sexually transmitted diseases or mat-
ing predation (Otto, 2009). Hence, in an inter-
play between cooperation and competition,
both sexes must apply strategies to strike a
sexual bargain: ‘No human soap opera could
outstrip in violence, hypocrisy, and manipu-
lation the daily drama of relations between
the sexes across the entire animal kingdom’
(Seabright, 2012: 8). Extreme manifesta-
tions of this drama include the penis fencing
enacted by hermaphroditic marine flatworms
(Pseudobiceros hancockanus):
[T]wo of them coil around each other and engage
in an aggressive head-to-head wrestle with weap-
ons drawn… Whichever worm wins does so by
piercing the head of the other with its spiked
organ, and coerces the loser into adopting the
female role in this relationship, and becoming the
sperm receptable and egg bearer. It’s easier to
produce sperm than it is eggs, and it’s harder to
bear the younglings, so the individual who man-
aged to claim the male mantle remains childfree,
and ready to crack on for another with a new
individual. (Rutherford, 2018: 100)
One topical focus in the fields of evolution-
ary psychology and evolutionary ecology, as
well as in more specialized subfields like
reproductive ecology, is the existence in evo-
lution of tradeoffs. These tradeoffs occur
between survival and reproduction, growth
and reproduction, quantity versus quality of
offspring, investment in current versus future
offspring, and the division of energy between
investing in offspring and obtaining a mate
(Sear, 2015: S44). As a result, evolutionary
streams of research have tended to focus on
the variations in female reproductive func-
tions subject to environmental changes with
an emphasis on the distinct fitness constraints
between males and females. One powerful
conceptual framework for understanding
which conditions occur in male versus female
sexual selection is Trivers’ (1972) parental
investment theory, which attributes females’
different reproductive strategies to their
higher metabolic demands (gestation, child-
birth, and lactation), lower potential lifetime
reproductive success, and greater initial
investment per gamete. These reproductive
factors require a greater investment of
resources in offspring, leading females of
many species to be choosier than males about
 EVOLUTIONARY PSYCHOLOGY AND REPRODUCTION
their choice of mate (Bribiescas, 2001; Buss,
2012; Barrett et al., 2014). Given that wom-
en’s choices tend to reflect what has histori-
cally helped to increase the reproduction and
survival of their offspring (Buss, 2012), it is
not surprising that evolution has favored such
selectivity.
Females are thus more selective than
males regarding characteristics that relate
to human adaptive challenges, such as the
power, wealth, intelligence, social status,
dominance, ambition, and popularity that
facilitate resource acquisition (Kenrick et al.,
1990; Buss, 2012). Because superior health
depends on better food and more abundant
territory but humans have spent most of their
evolutionary history as hunter-gatherers,
women’s evolved preferences include the
modern equivalents of the qualities once
needed for successful hunting and gather-
ing of accruable, defensible, and controllable
resources (Buss, 2012). This human behavior
thus again reflects past selection pressures
(Tooby and Cosmides, 2005).
Male success in mating, in contrast, is
dependent on intersexual selection and
intrasexual competition; that is, the need to
charm females while threatening and con-
quering other males (Symons, 1979; Puts,
2010). Indeed, some scholars have argued
that sexual attraction and selective social
attachments are among the most powerful
driving forces of human behavior, allowing
us to understand the richness of human art,
language, and creativity (Miller, 2000; Young
and Wang, 2004). Miller (2000), for exam-
ple, emphasized that ‘the human mind’s most
impressive abilities are like the peacock’s
tail: they are courtship tools, evolved to
attract and entertain sexual partners’ (2000:
4). Accordingly, because mate assessment is
relativistic, men attempt to stand out from the
crowd through bold advertising cues (Buston
and Emlen, 2003) such as
spending money in more conspicuous ways, …
acting like heroes, … standing firm against group
opinion in ways that could make them look good,
and … strutting their artistic creativity. What is
479
important is not simply that men are show-offs but
that they flash their feathers in especially ostenta-
tious ways when they are in a mating frame of
mind. Is it just a coincidence that human male
showiness responds to mating motives in the way
that males in other species respond to the mating
season? Perhaps, but it seems unlikely. Remember,
women prefer to mate with men who stand out
from the crowd. Thus the payoff is the same for
male humans as it is for peacocks and bighorn
sheep, and it is the payoff that drives natural selec-
tion at all levels – an increase in attractiveness to
the more discriminating sex, who will be disin-
clined to mate with a male who does not demon-
strate his worth. (Kenrick, 2011: 144)
Based on observations from 37 cultures, human
mating preferences differ greatly between the
sexes, with females valuing good financial
prospects and ambition or industriousness more
highly than do males, and males displaying a
greater preference for physical attractiveness
and younger mates (Buss, 1989, Buss et  al.,
1990). Although women also value physical
attractiveness in a potential mate, they do so
more as a signal of genetic benefit (Gangestad
and Buss, 1993; Gangestad et al., 1994) such as
superior immunocompetence (Puts, 2010) or
pathogenic resistance (Gangestad and Buss,
1993: 89).
In our contemporary world, the drastic
changes wrought by new technologies on
human procreation and the mating game
have made human reproduction an ever more
important area of research for those inter-
ested in cultural evolution. In vitro fertiliza-
tion (IVF), for example, has separated sex
from procreation; the contraceptive pill has
made procreation optional; and the Internet
has radically changed the way humans inter-
act, form new relationships, and seek out
partners, affording a novel setting in which to
test the functioning of evolved psychological
mechanisms. Historically, for example, dating
(mate-choice) decisions occurred in person –
and sequentially – at a bar or restaurant or
in the workplace, but now the Internet has
untied the Gordian knot by allowing quasi-
infinite choice from the comfort and privacy
of home. The digital world has thus loosened
480 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
the time constraints of finding and match-
ing suitable partners and moved us far away
from the traditional approach of contracting
marriages at the community level (Dunbar,
2012). It has also reduced the mental transac-
tion costs of identifying the average, as well
as the complications of finding a mate who
meets different mating criteria, whose impor-
tance have changed over time. For instance,
in a study of mate preferences across a
57-year span, Buss et al. (2001) showed that
the importance attached to good looks has
increased, possibly because the shift from
mid-20th-century reliance on radio to more
visual technologies like television, movies,
or the Internet has bombarded us with images
of physically attractive individuals (Buss
et  al., 2001). Nonetheless, because mate
choice works based on relative differences,
those evaluating potential mates need to pay
special attention to trait variations from the
baseline, such as women not being attracted
to deep voices but rather to voices that are
deeper than the average (Boyer, 2018).
Sex differences in mate preferences and
reproductive strategies reflect variations on
the adaptive problems faced in mate choice
by ancestral men and women (Buss, 1995).
Not only can competition to find a mate
lead to conflict, it may incentivize cheating
and strategic tactical deception. For exam-
ple, to secure enough time to convince a
female to mate, mourning cuttlefish mimic
a female display to a male rival on one side
of their mantle but male courtship patterns
to a potential female mating partner on the
other (Brown et al., 2012). This female mim-
icry gives an advantage to undersized males
who have a comparative disadvantage in a
potential fight with a more dominant rival.
In fact, because males can quickly switch
from dishonest to honest signals, dual gen-
der signaling is a frequent means of avoiding
combat during courtship, one that only mani-
fests when a rival male is nearby. In general,
mate-choice strategies reflect the fact that
nature operates in a market, meaning that the
dishonest signals that result from divergent
interests in the mating game can lead to an
arms race between deception and detection
(Boyer, 2018). As Williams (1966: 184)
pointed out decades ago, ‘selection will fos-
ter a skilled salesmanship among the males
and an equally well-developed sales resist-
ance and discrimination among females’.
Exploration of the new environments
brought about by technological change has
already helped to generate empirical insights
on mating strategies, with a major goal of the
new empirics being to isolate confounding
factors. For example, over the past several
decades, an extensive literature has produced
evidence of what is probably the most wide-
spread mating pattern of vertebrates (Burley,
1983: 191); namely, positive assortative mat-
ing or homogamy (see, e.g., Hollingshead,
1950; Warren, 1966; Vandenberg, 1972; Price
and Vandenberg, 1979; Buss, 1985; Thiessen
and Gregg, 1980; Hinsz, 1989; Mare, 1991;
Thiessen, 1994; Thiessen et al., 1997; Little
et al., 2006; Penton-Voak et al., 1999). Most
particularly, it has shown that homogamy fos-
ters altruistic social behavior, bonding, social
cohesion and interaction, a common basis of
norms and values, and better communica-
tion and conversations (Thiessen and Gregg,
1980; Buss, 1985; Kalmijn, 1994; Thiessen
et  al., 1997). Nonetheless, it remains meth-
odologically challenging to isolate mate
preferences from the context of constraints or
opportunities (Schwartz, 2013). For instance,
positive assortative mating may be driven by
who one has the opportunity to meet, with
a greater likelihood of being surrounded
by individuals with similar characteristics.
Thus, sorting by education or income may
be a function of a large amount of time spent
among others with similar achievement lev-
els in the work environment, school, or col-
lege (Hitsch et al., 2010a).
Online dating studies have the advantage of
reducing such constraints and testing whether
positive assortment is possible in these set-
tings. As Hitsch et  al. (2010b: 162) empha-
sized, ‘in online dating, assortative mating
arises in the absence of search frictions, due
 EVOLUTIONARY PSYCHOLOGY AND REPRODUCTION
primarily to preferences and specific market
mechanisms by which matches are formed’.
Hence, analyzing dating choices has the advan-
tage of avoiding norm and preference adjust-
ments made by couples over time because of
shared experiences, choices, lifestyles, or diets
(Little et  al., 2006). Nevertheless, an online
dating environment requires an agreed match
between two parties whose individual goals
must converge, meaning that a potential part-
ner with unrealistically high aspirations may
not be able to attract mates with a higher fit-
ness level. The advent of IVF and assisted
reproductive technology (ART) in the late 20th
century provides a new conduit for human
reproduction and mate choice. In contrast to
online dating, the clinical sperm donor market
allows participants access to sperm that would
be out of reach in a dating environment. Since
the creation of the Internet in the early 2000s,
online communities and organizations have
developed in which reproductive tissue can be
donated directly between participants, free of
clinical regulation. The Internet sperm market,
particularly, not being limited in preferred trait
availability like private sperm banks, offers a
much larger choice set with greater potential
for obtaining the desired characteristics.
At the same time, it remains unclear to what
extent the mate-selection process is driven by
the female’s desire for her offspring to have
traits similar to the male, and to what degree
she selects skills that help to increase the like-
lihood of success in the raising and survival
of her offspring (Whyte and Torgler, 2015,
2016a). Examining sperm donation can throw
new light on this dichotomy because it allows
isolation of the male traits that genetically
impact his offspring from other factors like
parenting assistance (Scheib, 1997; Whyte
and Torgler, 2015, 2016b). One pertinent
question is whether factors such as resources,
power, and prestige (Flinn, 1986; Townsend,
1989; Mulder, 1990; Pérusse, 1993; Li et al.,
2002) – all of which help to increase offspring
survival by providing the female with needed
assistance – are less relevant in this context
because genetic advantage matters more. In
481
other words, might decreased pressure to avoid
bad selection in anticipation of future parental
investment affect how females discriminate in
their choices?
In general, new technologies can be strong
instruments of courtship, whose ultimate func-
tion from a biological standpoint is to increase
the reproductive success of the individuals
involved (Grammer, 1989). Intrasexual com-
petition encourages the signaling of mate
quality, through whose cost credibility is
achieved. Nonetheless, as regards economist
Paul Oyer’s (2014: 212) conceptualization of
the signal as a ‘best alternative to the fact that
talk is cheap’, it is harder to use signaling in
standard online dating where everything can
be identified as cheap talk before the first date
(see also Oyer, 2014).
On the other hand, both online dating and
the sperm donor market facilitate examination
of female preferences under a nonsequential
choice mechanism, which insights should
increase our understanding of the decision-
making process in contexts characterized by
increased information and choice (Whyte
and Torgler, 2015). In fact, one comparative
study has already provided evidence that the
norms that guide behavior offline hold true in
many online environments (McLaughlin and
Vitak, 2011), probably because both environ-
ments require major life-event decisions that
force individuals to identify their true prefer-
ences prior to making a selection. Whereas
dating choices may be a frequent occurrence,
choosing a sperm donor tends to be a rare life
experience (Waynforth and Dunbar, 1995;
Pawłowski and Dunbar, 1999).
MATE-SELECTION INSTITUTIONS
AND TECHNOLOGIES
Traditional Matchmaker
Only as late as the mid 18th to mid 20th cen-
tury did love-based arrangements replace
arranged or patriarchal marriages, whose
482 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
pervasiveness in hunter-gatherer cultures
(Walker et al., 2011; Apostolou, 2014; Fletcher
et al., 2015) implies deep roots in human his-
tory (Walker et al., 2011). In fact, historically,
marriage was often a means of building per-
sonal networks to enhance access to hunting,
natural resources, or water holes, for example.
In other words, such unions were a way to
extend cooperative relations: ‘When people
married into new groups, it turned strangers
into relatives and enemies into allies’ (Coontz,
2006: 59). For assistance overcoming the chal-
lenges in forming such alliances, humans have
traditionally turned to a variety of agents or
institutions for help in finding a mate.
Traditional matchmakers were often spirit-
ual or religious leaders or respected elders in
the community (Finkel et al., 2012), such as
the shadchan of the Jewish community, who
has a long tradition in Jewish life. A role his-
torically entrusted only to the most learned
and respectable men, the shadchan provided
a unifying and stabilizing influence among
the scattered Jewish diaspora during its
oppression and persecution in Europe in the
13th and 14th centuries (Rockman, 1994).
The matchmaking services were necessary
because the Jewish orthodox community,
despite its strong emphasis on finding a mate
and creating a long-lasting family, mandates
almost complete segregation between the
sexes from puberty until marriage (Rockman,
1994). Rockman offers a detailed and color-
ful account of the procedural aspects of these
services, including the searching, matching,
and interacting rules that are also essential in
modern matching technologies:
a man or woman or their parents will approach Mr
Smith [the shadchan] with a request for him to find
them a mate. He and his wife then conduct an
interview with the client in question, lasting for
about half an hour. During this time the couple
need to assess qualities such as sincerity, educa-
tion, family background, and the quality of the
person’s relationships with others. They will also be
looking at physical attributes (a man will always be
expected to be taller than his prospective mate),
sense of humor, financial prospects, and degree of
religious feeling.
The details are then cross-checked with the person’s
friends, teachers, parents, business associates, and
local rabbi. It is not enough to know that a boy
went to a good rabbinical college. Mr Smith will
also want to enquire how he behaved there. Did he
miss class? How did he spend his time? Was he
ambitious? It is expected that each prospective
partner will double-check the accuracy of this
information. This is usually done through similar
channels to the ones used by the Shadchan himself.
Mr Smith explains that before the couple meet he
wants to be at least 60% sure that the match will
be successful. Once the candidate is screened, his/
her name will be filed along with the other names
in the file until a suitable partner is found. When a
suitable partner is found a monitored courtship is
initiated. (Rockman, 1994: 282)
The couple are first seen individually by Mr
Smith who informs them of their date’s attrib-
utes and instructs them regarding ‘dating’
behavior. For example, he will remind the man
that he is expected to pay for expenses on the
date and not do anything that will make him
appear to lack confidence (e.g., bite his nails).
The girl is encouraged to be ladylike. They are
then sent on their first date. The couple are
expected to report back to the Smiths and to
declare their intention to continue the relation-
ship or terminate it. Mr Smith has a policy of
asking the boy’s opinion first so that if there is
any rejection to be ‘inflicted’, he can let the
girl down gently. He believes that men are
emotionally stronger than women and can take
rejection in their stride.
There is no specific limit to the number of dates
allowed. However, friendship alone is not consid-
ered to be a justified reason for meeting. It is up to
the shadchan to make sure that the relationship is
advancing in a particular direction. That direction is
towards the wedding canopy. Hence prolonged
courtship is discouraged. There are some branches
of orthodox Judaism where the couple are allowed
to meet only once for one hour and then decide
whether they wish to marry.
If the couple decide to marry they are expected to
‘look after’ the shadchan. There is no fixed fee but
the belief is that a significant sum will bring the
couple good fortune in their relationship. The
shadchan may give the money to charity as does
Mr Smith, or use it in other ways. The shadchan is,
of course, invited to the wedding. Mr Smith
 EVOLUTIONARY PSYCHOLOGY AND REPRODUCTION
explained that the business of matchmaking is
growing more difficult as people progress and
adapt to the modern world. Demands for the ‘per-
fect’ partner are increasing. Few are prepared to
compromise as they may have done in past genera-
tions. Despite this Mrs Smith told me she believed
there is a match for everyone, it’s just a matter of
bringing them together. She advises against match-
ing couples who both have psychiatric histories in
the family. She does see, however, that a match
between a strong healthy person and a troubled
person may be successful. All the information con-
cerning illness is not always presented before the
couple meet. The matchmaker may decide that it
will affect the success of the date and will only
reveal the information at a more advanced stage of
the proceedings. (Rockman, 1994: 282–283)
The shadchan therefore serves an important
function in putting potential mates in contact
and ensuring their unilateral awareness in the
hope of mutual attraction and interest. It is
this unilateral awareness, together with con-
tact and mutuality, that is at the core of any
matching agency or technology because fac-
tors related to the fear of rejection or conflict-
ing desires may lead individuals to search for
a partner like themselves (Folkes, 1982). Just
as the solid reputation of the shadchan en­abled
contact without risks in traditional communi-
ties, the anonymity and distance inherent in
modern technologies assure similar protection
(Schnarch, 1997). Just like the shadchan,
matching models (such those formulated in
1962 by economists Gale and Shapley) take
care of the search process at the core of mate
selection (Rosenfeld and Thomas, 2012) and
provide substantial information about the
potential mate’s relative value. In this way,
new technologies help the shadchan open the
‘bidding’ between potential partners
(Pawłowski and Koziel, 2002: 140).
New web technologies have also dramati-
cally reduced search costs and allowed indi-
viduals to find mates outside of their existing
social network, although geographic prox-
imity is still important for the eventual goal
of face-to-face interaction (Rosenfeld and
Thomas, 2012). On the other hand, a large
pool of potential mates can make the selec-
tion task overwhelming because despite an
483
adaptive desire for more options to ­eliminate
the risk of incest and increase variety (Lenton
et  al., 2010), our limited cognition (Simon,
1956) is used to handling only a few sequen-
tial mate choices (Miller and Todd, 1998;
Lenton et  al., 2010). Thus, the mechanism
that produced a successful solution in the
evolutionary past may induce (overload)
problems in today’s overabundant dating
environment, especially given the individual
tendency to underestimate how daunting
increased choice can become (Iyengar and
Lepper, 2000; Lenton et al., 2010). It would
thus be useful for future research to include a
comparative assessment of whether narrow-
ing the market (Schwartz, 2004) increases the
matching success rate.
In Western countries such as the United
States, mate selection before World War II
was dominated by active family engagement
in identifying a pool of potential mates within
the neighborhood, the church congregation,
or among primary and secondary schoolmates
(Rosenfeld and Thomas, 2012). Then, after
World War II and up until the early 2010’s
friends overtook family as the key resource
for finding both male and female partners
(Rosenfeld and Thomas, 2012, Rosenfeld
et al., 2019). Over the second half of the 20th
century, the workplace grew in importance
as a social milieu in which to meet a roman-
tic partner but has faded as an option in the
Internet era (Rosenfeld and Thomas, 2012). In
fact, growing sensitivity about sexual harass-
ment has reversed the view of it as an ideal
environment in which to form intimate rela-
tionships (Barraket and Henry-Waring, 2008).
Singles Ads and Video Dating
In recent years, not only have delayed mar-
riage, changed marital expectations, shifting
sexual norms, birth control, and the search
for emotionally and intellectually gratifying
partnerships created a singles boom (Ahuvia
and Adelman, 1992), but growing career
pressures, increasing time poverty, and higher
484 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
mobility have reduced opportunities for
social activity and the sustenance of intimate
relationships. As a result, new institutions
and technologies have emerged to help indi-
viduals deal with the challenges of mate
choice and selection. In the late 1970s, these
took the form of personal ads in the singles
columns of newspapers, becoming an estab-
lished feature that was heavily discussed in
magazines and talk shows (Ahuvia and
Adelman, 1992). However, because of a per-
sistent stigma, throughout the 1980s (just as
in the 1960s), only a small fraction of indi-
viduals (around 1% in the United States) met
via ‘the personals’ (Finkel et al., 2012).
An alternative means of meeting a mate,
one offered by a small number of third-party
intermediaries, was video dating, in which
a client first read a written description of
potential dates (most including an image)
and then decided which of the 2–10 minute
videos to watch. If the client made a choice,
the selected person was provided with the cli-
ent’s video (Ahuvia and Adelman, 1992), and
if both agreed to meet, phone numbers were
given. Video dating provided an opportu-
nity to prescreen dates, furnish more natural
information, and present a more purposeful
image by investing a great deal of effort in
building a profile (Woll and Young, 1989).
Videos also allowed exploration of nonver-
bal communication in courtship settings,
including the body language that is such an
important source of mate-choice information
(Renninger et  al., 2004). A distinguishing
feature of the singles ads and videotapes was
the awarding of more personal control over
both the pace and nature of the interaction –
as well as the information provided (Woll
and Young, 1989) – an ability also facilitated
by online interaction via new technologies
(Barraket and Henry-Waring, 2008).
Research into the efficacy of these
resources has traditionally been conducted
in a laboratory setting with a subject pool
of college and university students asked to
respond to hypothetical mating situations.
One exception is evolutionary psychological
research into personal ads, which has the
great advantage of going beyond the labora-
tory setting to study the behavior of a wider
sample of anonymous individuals under
less artificial circumstances (Minervini and
McAndrew, 2006). Lynn and Bolig (1985),
in their detailed discussion of the positive
aspects of analyzing personal ads, empha-
size this ability to explore real behavior in a
naturalistic setting with a more representa-
tive sample and longer-term consequences,
which ensures greater generalizability than
laboratory-based behavioral observations. In
fact, before the inception of online dating,
increasing the external validity of the results
was a core goal of scholars working with
laboratory data.
Because placing or answering personal ads
can be seen as a form of courtship (Goode,
1996), related studies have explored the rela-
tive importance of various factors using both
bogus (e.g., Sitton and Blanchard, 1995;
Goode, 1996) and actual advertisements
(e.g., Lynn and Shurgot, 1984). After analyz-
ing the ad content, these studies linked it with
the number of responses received (Lynn and
Bolig, 1985) as a viable measure of success
(Pawłowski and Dunbar, 2001). Some studies
also compared the content with information
from the ad placers or explored the number
of ads as a dependent variable with which to
measure interpersonal relations (Ahuvia and
Adelman, 1992). Research into video dating
has focused on similar questions and found
comparable insights (Ahuvia and Adelman,
1992). One research aspect that remains
underdeveloped, however, pertains to envi-
ronmental shifts like legislative change, eco-
nomic conditions, media events, and cultural
holidays (Lynn and Bolig, 1985: 382), whose
relevance for the technological institutions
discussed in this chapter makes them a prom-
ising subject for further analysis. In particu-
lar, if the event studied were exogeneous,
it could be treated as a natural experiment,
which would allow the use of methodologies
able to address causality issues and identify
confounding factors.
 EVOLUTIONARY PSYCHOLOGY AND REPRODUCTION
One notable finding from extant studies
is that women are concerned with finding an
older partner with established resources (e.g.,
education and income or wealth), while men
prefer young, physically attractive women
(see, e.g., Greenlees and McGrew, 1994;
Goode, 1996; Pawłowski and Koziel, 2002;
Minervini and McAndrew, 2006). On the
other hand, in a study in Sweden, although
females were substantially more focused on
resources than males, both cared about physi-
cal attractiveness (Gustavsson et  al., 2008),
a particularly interesting finding given that
Scandinavian countries have high levels of
economic equality between the sexes (Buss,
2012). Even when women greatly outnum-
ber men, as in a pool of dating-service par-
ticipants in Tel Aviv (646 males to 1,000
females), rather than being less choosy or
lowering their expectations, women are more
selective than males in most areas (Bokek-
Cohen et  al., 2008). Such settings are espe-
cially useful in allowing the differentiation
of various competing theories – in this case,
rational choice versus evolutionary psychol-
ogy, a comparative study type still underrep-
resented in the literature.
Research has also examined mail-order
brides, who represent an effort to vastly
broaden the opportunities afforded by tradi-
tional matchmaking by expanding the pool
of eligible mates – and thus the range of
choices – across national borders. The prior-
ity among one set of potential brides from
Colombia, the Philippines, and Russia was to
find older males with access to resources and
a commitment to share them (Minervini and
McAndrew, 2006), which supports evolution-
ary explanations of female mate preferences.
Speed Dating
Speed dating, which first came to prominence
in Los Angeles in the 1990s (Finkel et  al.,
2007), involves a group of individuals meet-
ing at one location and spending recursive
intervals of time with potential partners in the
485
room. Organizers often provide participants
with homogenous questions to ask potential
suitors, as well as score cards with which to
rate potential partners across a range of char-
acteristics. Nonetheless, although popular
and still widely practiced (with commercial
speed-dating companies now running the
events), speed dating’s participatory con-
straints (in-person attendance, limited
number of potential mates, possible financial
cost, and potential anxiety about in-person
contact) have prevented it reaching the same
level of popularity as online dating.
The research into speed dating was part
of the shift that moved studies of attractive-
ness or relationships beyond the laboratory
into ‘live interactional settings’ (Korobov,
2011: 186). Methodologically, the speed-
dating design is particularly valuable because
it allows separation of the actor effect from
the partner effect (i.e., from ‘how do I behave
toward others?’ to ‘which behavior do I evoke
in others?’; Asendorpf et al., 2011: 16). It can
thus be seen as a microcosm of sequential
mate choice in daily life but in a faster and
more formalized manner (Todd et al., 2007).
One consistent finding from this research
stream is that fundamental sex differences
in mate selection continue to manifest even
in such modern forms of dating (Roberts
et  al., 2010; Buss, 2012). For example, one
of the earliest speed-dating studies – a com-
pilation of observational data and additional
survey information – provides strong sup-
port for women being more selective than
men (Kurzban and Weeden, 2005). This
finding is confirmed experimentally using
a sample of nearly 400 Colombia graduate
students, among whom the females placed
greater importance on intelligence, while the
males were more likely to respond to physi-
cal attractiveness (Fisman et al., 2006). Men
who grew up in affluent neighborhoods also
had a comparative advantage.
In general, speed-dating research has
shown that female selectivity is more malle-
able or context-dependent than that of males;
for instance, across 22 speed-dating events
486 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
with 546 participants, less attractive females
and those with more irregular BMI were
overall less selective at events characterized
by higher intrasexual competition (Overbeek
et al., 2013). Nonetheless, stated preferences
do not necessarily match actual choices in
either the offline (Todd et al., 2007) or online
(Whyte and Torgler, 2017a) setting. Rather,
women tend to make more discriminating
choices (Todd et al., 2006), which is again in
line with evolutionary models of human mat-
ing based on parental investment theory.
Online Dating
According to a nationally representative
2017 survey in the United States, online
dating has become the most popular way for
heterosexual couples to meet (Rosenfeld
et  al., 2019), although websites facilitating
online dating did not rise to prominence until
after the 1990s proliferation of the informa-
tion superhighway. Date-matching software,
however, dates back to a 1959 Stanford class
project when two electrical engineering
undergraduates wrote a matching program
that generated difference scores for each pos-
sible male–female pair based on their survey
responses, with lowest scores representing
the best match and so on up the scale. In
general, these students observed that both
men and women were keen to find mates
with similar habits (Gillmor, 2007).
Although individuals register on these
electronic platforms by building a personal
profile – somewhat like that compiled by
the traditional matchmaker – the platforms
are less restrictive than either speed dat-
ing or everyday personal contacts, and are
not institutionally prestructured or selective
(Skopek et al., 2011). Whereas some online
dating sites allow searches based on specifi-
cally chosen characteristics (e.g., RSVP) or
site-provided personality-matching systems
(e.g., eHarmony), others are based on social
networks that encourage users to bring in
new members (e.g., Friendster; Barraket
and Henry-Waring, 2008). Services such as
ScientificMatch and GenePartner even prom-
ise matchings based on genetic information
(Frazzetto, 2010). Moreover, recent decades
have witnessed the emergence of myriad
variations, including free and paid services,
dating applications linked to social media
accounts and specific groups, and special
focus platforms ranging from participant
sexuality and sexual behaviors to race, cul-
ture, interests, occupation, and religious and
political views.
As a result, over the past 20 years, behav-
ioral science has begun to explore human
mating behavior in this unique new cyber
market across the gamut of scientific dis-
ciplines, including social and evolutionary
psychology, economics, sociology, demogra-
phy, personality psychology, communication
research, and cyber-psychology. Scholars
working on this topic emphasize the power of
online dating data given their basis in actual
observed preferences, which, although also
characteristic of speed-dating experiments,
is contrary to the stated preferences used in
mate-preference studies (e.g., Todd et  al.,
2007; Finkel and Eastwick, 2008; Hitsch
et  al., 2010b). These latter, by asking part-
ners to infer backward what traits originally
attracted them to each other, are made prob-
lematic by memory fallibility (Todd et  al.,
2007), cognitive dissonance, or even self-
deception. On the other hand, online daters
may admittedly struggle to obtain sufficient
information on the experiential attributes
(e.g., sense of humor) that are relevant when
determining whether or not one person likes
another (Frost et al., 2008).
One advantageous development for this
research is the increased use of dating apps,
which provide researchers with a valuable
opportunity to work either with a representa-
tive sample of the population or specific sub-
groups. As a result, researchers have been
able to explore a plethora of online mate-
choice psychology and behaviors using very
large populations, such as the over 22,000
online daters in Boston and San Diego studied
 EVOLUTIONARY PSYCHOLOGY AND REPRODUCTION
by Hitsch et al. (2010a, 2010b). Using 2003
data for a three-and-a-half-month period,
these researchers not only documented a
female preference for income over physical
attributes, but also, as in other studies, a strong
preference in both sexes for homogamy across
most choice characteristics (e.g., education,
ethnicity, marital status, religion, height,
smoking habits, and political views).
The research on such electronic mate-
choice settings therefore seems to mirror the
findings of noncyber mate-choice research,
which has identified key sex differences in
stated preferences, choices, and behaviors.
For example, Skopek et  al.’s (2011) study
of over 10,000 German online daters (with a
biased sex ratio of 65.42% males) showed an
average of 12.75 contacts for men compared
to 8.02 for women. These dater contact-
initiation data also indicated that the males
contacted females who were on average
4.76 years younger than themselves, whereas
the females contacted males who were on
average 2.74 years older. Although these age
preferences shift with age for both sexes, they
do so differently for men and women, with
the latter’s becoming generally more hetero-
geneous with age (Skopek et al., 2011: 267).
Fiore et  al. (2010) similarly demonstrated
that among 11,600 active online daters,
even though men outnumbered women sig-
nificantly (57.9% male; 42.1% female),
almost four out of every five (77.1%) con-
tact initiations were by men toward women,
with women receiving a median number of
two contacts but men only 0.5. Moreover,
whereas men expressed a preference in their
profiles for an average 9.1 characteristics in a
mate, women specified 11.9.
As regards specific characteristics, accord-
ing to one Australian study of over 40,000
online daters (Whyte et  al., 2018), women
across all age groups (18–80 years) were
more likely to state a preference for a particu-
lar education level in a potential mate, which
preferences were more likely to be homog-
enous (i.e., positively assorted) or at least
have a lower minimum than those of males
487
(Whyte and Torgler, 2017b). Likewise, a
Chinese study using over half a million online
dater profiles found that women attached
greater importance than men to a potential
partner’s socioeconomic status (Su and Hu,
2019). Such female preferences tend to be
more refined during the women’s years of
maximum fertility, after which their demand
specificity decreases with age. Male speci-
ficity, on the other hand, remains stable until
their 40s, when it becomes greater than that
of post-reproductive females, meaning that
it is at its highest during their peak career-
earnings potential (Whyte et al., 2018). Also
consistent with evolutionary theory, research
focused specifically on the dating behavior of
older adults found that whereas men’s choices
are more consistent with evolutionary predic-
tions, it is women who define the choices,
implying a greater sensitivity to demographic
realities (Alterovitz and Mendelsohn, 2009).
The pervasive modern use of the Internet
also now assures that cyber populations are
increasingly representative of the offline pop-
ulation (Valkenburg and Peter, 2007), which
reduces selection bias in online samples. That
said, high-quality mates are scarce in any mar-
ket, and competition both between and within
the sexes is fierce. Hence, the anonymity
of the Internet may strongly encourage decep-
tive strategies to secure the highest-value mate
possible, just as in real life females enhance
their physical beauty with make-up and males
wear shoe lifts or a hairpiece. In fact, the lower
the personal attractiveness, the higher the
probability of individuals enhancing their pro-
file photographs to enhance height, weight,
or age characteristics while not extending
such deception to non-aesthetic profile char-
acteristics like income and occupation (Toma
and Hancock, 2010). Yet sex differences are
also observable in the characteristics likely
to be embellished during profile building,
with males more prone to exaggerate height,
but women more likely to understate weight
(Toma et al., 2008). Humans can even be mis-
led by their own self-deception in that online
daters’ desired or preferred characteristics
488 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
in an ‘ideal partner’ do not always mirror
those of the individuals they actually contact
(Whyte and Torgler, 2017a).
Needless to say, an individual’s personal-
ity also plays a role in how attractive they
are – and how well they project themselves –
to possible mates. For example, whereas
Valkenburg and Peter (2007) showed that
those with low (vs high) dating anxiety are
more active in the online marketplace, Whyte
et al. (2018) identified a positive linear rela-
tion between levels of extraversion and sexual
frequency for both sexes. In this latter study,
greater variance in male personality traits
translated to men – but not women – having
more sex and more children. On the other
hand, a German longitudinal study found that
those with less extrovert personalities were
more likely to meet online, perhaps because
the Internet provides a more passive way to
initiate communication with potential mates.
Just as homogamy or positive assortment
is overwhelmingly the observed mate-choice
behavior across cultures in real life (Buss,
1985), so the ecology of Internet mating
appears no different. Not only did Whyte and
Torgler’s (2017b) analysis of around 220,000
contact initiations reveal that more educated
online daters were more likely to positively
assort and less likely to contact those with
lower educational levels, it also demonstrated
that, overall, males and females exhibit simi-
lar preferences as they age. Admittedly,
however, most such research has been con-
ducted in OECD countries, raising a need
for detailed future studies of mate-choice
behavior in other regions to assess the robust-
ness of current results for the online environ-
ment. More cross-cultural evidence is also
required to better understand human decision
rules when forming preference and choices,
including whether other societies employ the
same ‘likes-attract’ rule common in Western
cultures (Buston and Emlen, 2003), particu-
larly in evolutionary relevant traits (Thomas
et  al., 2019), which may help explain why
homogamous marriages are more common
and successful than nonhomogamous ones.
An ideal conceptual framework for such
investigation would be the ‘integration of
evolutionary psychology and cultural psy-
chology’, which Buss et al. (2001: 502–503)
consider the ‘hallmark of modern evolution-
ary science’.
Because evolution and changes in norms
and cultural systems provide a rich avenue
for understanding interpersonal interactions
and mating choices, future research within
this theoretical context could provide valu-
able additional insights into the dynamics
of communication, self-expression, types of
courting strategies, and characteristics (e.g.,
level of emotionality or aggressiveness). For
instance, institutional and environmental
changes – particularly when exogenous –
might serve as natural experiments, enabling
the use of methodologies that can iden-
tify confounding factors and discriminate
between different theories and alternative
hypotheses. Likewise, informal institutional
factors such as trust and social exchange
could provide valuable new insights into
human behavior in the mating or matching
markets. It is thus well worth exploring in
more detail how evolutionary psychology
or biology could contribute to the design
and management of institutional changes
that benefit individuals and societies by, for
example, increasing their levels of happiness
(Piazza and Bering, 2009; Wilson, 2018).
REPRODUCTIVE DONATION
Reproductive Medicine and
Donation
Although reproductive technologies offer an
exceptional way to reproduce, become a
parent, and construct a family, they also chal-
lenge the historical foundations of how
humans reproduce and cooperate to raise
offspring (Hargreaves, 2006), which for mil-
lennia has required the physical act of sex.
Late-20th-century advances in ART and IVF,
 EVOLUTIONARY PSYCHOLOGY AND REPRODUCTION
however, have changed (and in part removed)
the reproductive need for physical copula-
tion. Although the first case of human artifi-
cial insemination dates back to around 1790
when John Hunter successfully inseminated3
a patient with her husband’s sperm, it was in
the last decades of the 19th century that
norms tying sex to procreation relaxed. The
first donor insemination was performed
(quite unethically) in 1884 by Philadelphia
physician William Pancoast, who, having
found the woman’s husband sterile from an
earlier bout of gonorrhea, procured semen
from the medical student deemed handsomest
in the class, anesthetized her, and inserted the
sperm via syringe, resulting in a successful
pregnancy. Although Pancoast failed to
inform the couple at that time of the sperm
donor’s involvement, when another physician
brought the case to light in 1909, Pancoast
confessed the truth and was relieved when the
husband responded positively to the news,
albeit deciding that his wife should never
know the truth (Guttmacher, 1969: 567).4
Clinics offering ART and IVF, as well
as commercial markets for both sperm and
oocyte donation, have traditionally incen-
tivized increased donation via one of the
two most effective motivators (Ernst et  al.,
2007), either an altruistic (no personal gain)
or a financially remunerative model (Daniels,
1989). The advent of the Internet, however,
together with the limited clinical supply of
both sperm and oocytes, has encouraged
the emergence and growing popularity of
online or informal markets for reproductive
tissue donation, which provide increased fit-
ness opportunities for both men and women.
As a result, the last decade has witnessed
the launching of online connection web-
sites (analogous to dating websites) that
facilitate donor–recipient exchange glob-
ally. Although explaining these services
empirically as new behavioral adaptations
may not be feasible given that they are only
a generation or two old, research can assess
whether previous mating market dynamics
and underlying mechanisms are still present
489
in the related choices and behaviors. ART has
also provided evolutionary science a unique
opportunity to explore female reproductive
behaviors in a way that previously was not
possible. Historically, internal fertilization
and gestation were mandatory for women to
reproduce; medical science has now released
women from such constraint. As previous
evolutionary axioms such as Trivers’ 1972
parental investment theory relied on women
experiencing the greater opportunity cost (the
physiological burden) of reproduction, it is
unclear exactly what this fundamental change
means for sex-differentiated behavior based
on ‘female-choice’ constraints that may no
longer be relevant. If modern reproduction via
donation only requires a resource-endowed
participant and a willing donor or surrogate,
many previous mate-choice theories related
to sexual selection may need to be reexam-
ined. In addition, one could also explore in
more detail the psychology and behavior of
kin not raised by their biological parents. Just
as twin studies provide a unique opportunity
to explore phenotypic variance on individual
development and behavior, so too do children
born of the same donor parent, but raised by
different (non)biological parents in differ-
ent environments. The way in which these
changes will impact and influence intergen-
erational and population-level change is an
exciting and challenging research question.
Sperm Donors and Recipients
Historically, males had great difficulty find-
ing and securing female mates who sought
little or no paternal investment; most particu-
larly because nomadic and agriculture-based
societies were resource poor and raising off-
spring was (as ever) extremely costly. The
development of mutual investment in self-­
regulated pair bonding (e.g., marriage) as a
social norm thus appears eminently logical. In
the current century, however, particularly in
developed economies, resources are not scarce,
while historical-education, labor-market, and
490 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
legislative barriers to gender equity in resource
acquisition are greatly reduced. Females are
thus no longer totally dependent on paternal
investment from males, while males no longer
need to remain in long-term monogamous
relationships to ensure increased fitness.
From an evolutionary perspective, ceteris
paribus on environmental condition (Quinlan,
2007), the optimal male reproductive strat-
egy is to mate and reproduce with as many
females as possible – thereby realizing the
greatest fitness gains – which requires no
more investment than the time to copulate.
Females, in contrast, still bear the higher
opportunity costs of gestation, nursing, child-
rearing, and other disproportionately demand-
ing parental investment, even though many in
developed economies no longer rely on male
paternal investment. Hence, for the first time
in human history, males can and are encour-
aged to provide their reproductive tissue to as
many different women as possible purely for
the purpose of reproduction and free of any
parental investment constraint. This commer-
cial donation market, with its two very differ-
ent motivators (i.e., altruistic versus financial)
and its culturally varying legislation on donor
anonymity, thus presents a unique opportunity
to explore both men’s and women’s prefer-
ences and reproductive decisions free of his-
torical resource constraints.
One aspect examined to date by much of
the (nonevolutionary) scientific research on
sperm donation is the motivation for altru-
istic nonpaid donation given that donors
have no opportunity to designate their sperm
for particular groups or populations or to
exclude specific categories or characteristics
(Pennings, 1995). In fact, according to one
survey of Danish sperm donors active across
the last three decades, although altruism was
the most frequently reported motivator, large
numbers of respondents admitted that they
would not donate if economic compensation
were removed (Bay et  al., 2014). Clinical
survey data have confirmed this male ten-
dency to be motivated by financial compen-
sation, while also showing that male (sperm)
donors are not only less altruistic than
female (oocyte) donors but also more afflu-
ent (Schover et  al., 1992). From an evolu-
tionary psychological perspective, however,
sperm donation with no parental investment,
whether spurred on by financial remunera-
tion or the increased warm glow utility of
giving freely, is arguably the most selfish
and strategic way a male can increase his
fitness. Indeed, research has confirmed that
males acknowledge the drive to reproduce or
procreate as a key motivating factor in their
donation decision (Woestenburg et al., 2016).
Yet the bulk of the qualitative social science
literature in this area lumps unremunerated
donation under the blanket term of altruism
even when males acknowledge no need to
contribute toward parental investment and the
fitness gain of prolific offspring (Bergen and
Delacroix, 2018).
A more advantageous research focus is
that on females in search of a sperm donor,
which, by enabling the differentiation of
trait preferences from partnership factors,
creates a controlled setting that isolates a
male’s genetic impact on his offspring from
other characteristics. This separation in turn
permits the isolation of aspects that reduce
anticipated problems in child rearing via par-
enting assistance and improved environmen-
tal conditions for the offspring (Whyte and
Torgler, 2015). According to experimental
research, when assessing sperm donor char-
acteristics, women tend to rely on the same
psychology used to choose a long-term mate
(Scheib, 1994), with single (vs partnered)
women seeming to place higher value on
male proxies for resources and good charac-
ter (Rodino et al., 2011). Such findings may
also reflect females’ evolved adaptive prefer-
ence for males who can compensate their dis-
proportionate reproductive investment with
other resources (Trivers, 1972), as reflected
by a preference for higher donor levels of
education (Whyte et  al., 2016). Moreover,
in line with previous mate-choice research,
even when available choices are limited rela-
tive to traditional sequential dating markets,
 EVOLUTIONARY PSYCHOLOGY AND REPRODUCTION
women still exhibit homogamous preferences
(Whyte and Torgler, 2015).
Donor insemination essentially operated
globally without any legal restriction until the
Swedish parliament enacted the 1985 Law on
Artificial Insemination, which granted chil-
dren born as a result of donor insemination
access to identifying information on their
donor father (Lalos et al., 2003). In 2005, the
UK government introduced similar legisla-
tion (Daniels et al., 2005), after which other
countries followed suit, generating fierce
debate across all markets about donor ano-
nymity and which functional model could
best regulate clinical donation.
Online Sperm Donor Market
Because legislation and reform tend to lag
behind actual behavior, mushrooming social
media technologies have encouraged both
donors and recipients of reproductive tissue
to participate in a large but informal global
online market that is more interactive and
less rigid than official channels (Whyte and
Torgler, 2015). Such informal markets have
emerged because of the excessive demand
and inadequate supply that characterize the
official sperm donor market (Yee, 2009; Van
den Broeck et al., 2013; Whyte and Torgler,
2015). Before the advent of this new setting,
researchers struggled to align preferences
(e.g., for assortative mating) with actual con-
straints in the matching process, including
geographic limits and the social propinquity
and competition that substantially restricted
possibility sets and preferences. In the online
sperm donor market, such limitations are
reduced, and the geographic proximity
restriction of the online dating market mat-
ters less.
At the same time, the enactment of iden-
tification laws has reduced the number of
clinical donors in traditionally anonymous
markets (Ernst et al., 2007), prompting a sig-
nificant and growing male population to turn
to informal donation via coordination with
491
potential recipients. Although at the time of
writing research in this area is scarce (the set-
ting being little more than a decade old), it
has established that such factors as logistics,
health concerns, and access to information
on genetic heritage matter greatly in sperm
donation environments outside fertility clinics
(Lavoie et  al., 2018: 184). These informal
donors, although they tend to move between
clinical and informal settings, are primar-
ily motivated to donate via the Internet by
a desire to know potential recipients and be
updated on the development of any result-
ing offspring (Woestenburg et al., 2016). For
these men, who coincidentally donate to more
women and realize more offspring (Whyte
et al., 2018), motivation often relates to con-
tact and bonding with the recipient (Bossema
et al., 2014), although 94% of those with arti-
ficially inseminated realized offspring, report
that sexual gratification played no role as a
motivating factor (Freeman et al., 2016).
As regards successful donor characteris-
tics, in addition to their age and income being
positively correlated with number of female
recipients, shy, more intellectual, and/or more
organized informal donors realize more off-
spring than the more outgoing (Whyte and
Torgler, 2016b), with conscientiousness also a
comparative advantage (Whyte et  al., 2018).
Given that informal donation requires a level
of organization, communication, and even
logistical coordination (with the recipient) not
required in clinical donation, it is not surpris-
ing that more efficient or organized donors,
and those with more prosocial personality
traits, are more successful. In fact, these infor-
mal donor behavioral traits appear more highly
valued by females than physical appearance or
even proxies for resources (Whyte and Torgler,
2016b; Whyte et al., 2017).
Nonetheless, given the previous research
focus on mate selection, more empirical evi-
dence is needed on reproductive tissue dona-
tion if its selection and choice processes are to
be fully understood. As emphasized by Buss
(2012: 44–45), ‘the evaluation of evolutionary
formulations rests with the cumulative weight
492 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
of the evidence, and not necessarily with any
single prediction. Evolutionary hypotheses,
when formulated precisely, are highly test-
able and eminently capable of being falsified
when the evidence fails to support predic-
tions derived from them’. One particularly
promising avenue for discovery is the con-
nection websites that serve as a novel conduit
for reproductive tissue donors and recipients,
free of medical and legislative regulatory
oversight. It remains unclear, for example,
exactly how participant characteristics and
behaviors on these websites differ from those
in the regulated donor insemination settings,
and more broadly from those in earlier elec-
tronic mate-choice markets like online dat-
ing. Compared to sperm donation research,
evidence on egg donors and recipients with
and without co-parenting intention is very
limited. Connection sites also allow explora-
tion of how participants initiate and receive
contact from other interested members, and
how likely they are to respond. These sites
also permit assessment of courtship inten-
sity and the presence of key sex differences
in profile formation, contact behavior, and
choice processes. A comparison of sperm
and egg donors would also facilitate evalua-
tion of whether the former, as bearers of very
different cost and investment levels in this
reproductive market, are choosier and more
selective than the latter. The reality that eggs
are large, expensive, and scarce mandates
that individuals be especially selective about
their source; while the small size, abundance,
and low cost of sperm (Seabright, 2012)
afford males more mobility. In other words,
males can court multiple females and more
aggressively maximize the number of poten-
tial matings and search areas.
CONCLUSION
Throughout human evolution, technological
change has been a key driver of success,
providing select people and groups a com-
parative advantage in resources, time, power,
status, and reproduction. These advances
have dramatically changed the human spe-
cies’ ability to survive, thrive, and reproduce.
Two of the more recent and arguably most
impactful technological leaps – stemming
from medical science and the information
superhighway, respectively – are the devel-
opment and commercialization of assisted
reproductive technology and donor insemi-
nation and the broadening of their availabil-
ity across global networks. Because of these
technologies, for the first time in human his-
tory, females and males can pass on their
genes and realize offspring to unknown indi-
viduals on the other side of the planet.
Nonetheless, because sexual selection has for
millennia favored those most able to identify,
access, and secure suitable partners, the cur-
rent impact of these technological advances
remains unclear.
In this chapter, therefore, we have explored
this impact on behavior and reproductive
outcomes by examining such topics as the
online dating market and the implications of
the Internet for nonsequential mate-choice
decision making or sperm donation. In
doing so, we have shown ways to observe
human mate-choice behavior in electronic
and modern medical settings through the
lens of evolutionary psychology. The overall
conclusion drawn from this exploration is
that core evolutionary constructs like sexual
selection and parental investment are still
present and relevant in these 21st-century
reproductive behaviors. Women have evolved
powerful preferences for long-term mate
characteristics that help solve the adaptive
problems of survival and reproduction.
Hence, even though newer ways of selection
seemingly achieve fundamentally different
choices from more traditional means of
courtship or mate choice, evolutionarily,
men’s and women’s choices are relatively
stable, showing no new responses to novel
technological conditions.
 EVOLUTIONARY PSYCHOLOGY AND REPRODUCTION
Notes
1  Interestingly, a certain class of rotifers (a fresh-
water multicellular eukaryote) reproduce via sex
when faced with environmental changes but
reproduce through cloning in environmentally
stable conditions (Greely, 2016).
2  This observation refers to experimental popula-
tions of a facultatively sexual species of the hap-
lodiploid monogonont rotifer, which changes to
sexual reproduction in response to a chemical
stimulus.
3  The writings of the Talmud discuss the possibil-
ity of pregnancy unrelated to sexual intercourse;
namely, through accidental insemination in bath-
water (Bernstein, 2002).
4  For a history of artificial insemination with its
hurdles and milestones, see Ombelet and Van
Robays (2015) and Foote (2002).
REFERENCES
Ahuvia, A. C., & Adelman, M. B. (1992).
Formal intermediaries in the marriage
market: A typology and review. Journal of
Marriage and the Family, 54(2), 452–463.
Alterovitz, S. S.-R., & Mendelsohn, G. A.
(2009). Partner preferences across the life
span: Online dating by older adults. Psychol-
ogy and Aging, 24(2), 513–517.
Apostolou, M. (2014). Sexual selection under
parental choice: A revision to the model.
Theory in Biosciences, 133(2), 111–115.
Asendorpf, J. B., Penke, L., & Back, M. D.
(2011). From dating to mating and relating:
Predictors of initial and long-term outcomes of
speed-dating in a community sample. Euro-
pean Journal of Personality, 25(1), 16–30.
Ball, P. (2019). How to grow a human: Adven-
tures in how we are made and who we are.
Chicago, IL: University of Chicago Press.
Barraket, J., & Henry-Waring, M. S. (2008).
Getting it on (line): Sociological perspectives
on e-dating. Journal of Sociology, 44(2),
149–165.
Barrett, L. T., Evans, J. P., & Gasparini, C.
(2014). The effects of perceived mating
opportunities on patterns of reproductive
investment by male guppies. PLoS One, 9(4),
e93780.
493
Bay, B., Larsen, P. B., Kesmodel, U. S., & Inger-
slev, H. J. (2014). Danish sperm donors across
three decades: Motivations and attitudes.
Fertility and Sterility, 101(1), 252–257.
Becks, L., & Agrawal, A. F. (2012). The evolution
of sex is favoured during adaptation to new
environments. PLoS Biology, 10(5), e1001317.
Bergen, N., & Delacroix, C. (2018). Bypassing
the sperm bank: Documenting the experi-
ences of online informal sperm donors. Criti-
cal Public Health, 29(5) 1–12.
Bernstein, G. (2002). The socio-legal accept-
ance of new technologies: A close look at
artificial insemination. Washington Law
Review, 77, 1035–1120.
Bokek-Cohen, Y., Peres, Y., & Kanazawa, S.
(2008). Rational choice and evolutionary
psychology as explanations for mate selectiv-
ity. Journal of Social, Evolutionary, and Cul-
tural Psychology, 2(2), 42–55.
Bossema, E. R., Janssens, P. M., Treucker, R. G.,
Landwehr, F., van Duinen, K., Nap, A. W., &
Geenen, R. (2014). An inventory of reasons
for sperm donation in formal versus informal
settings. Human Fertility, 17(1), 21–27.
Boyer, P. (2018). Minds make societies: How
cognition explains the world humans create.
New Haven, CT: Yale University Press.
Bribiescas, R. G. (2001). Reproductive ecology
and life history of the human male. Ameri-
can Journal of Physical Anthropology,
116(S33), 148–176.
Brown, C., Garwood, M. P., & Williamson, J. E.
(2012). It pays to cheat: Tactical deception in
a cephalopod social signalling system.
Biology Letters, 8(5), 729–732.
Burley, N. (1983). The meaning of assortative
mating. Ethology and Sociobiology, 4(4),
191–203.
Buss, D. M. (1985). Human mate selection:
Opposites are sometimes said to attract, but
in fact we are likely to marry someone who
is similar to us in almost every variable.
American Scientist, 73(1), 47–51.
Buss, D. M. (1989). Sex differences in human
mate preferences: Evolutionary hypotheses
tested in 37 cultures. Behavioral and Brain
Sciences, 12(1), 1–14.
Buss, D. M. (1995). Psychological sex differ-
ences: Origins through sexual selection.
American Psychologist, 50, 164–168.
494 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Buss, D. (2012). Evolutionary psychology: The
new science of the mind. Pearson: University
of Texas at Austin.
Buss, D.M., Abbott, M., Angleitner, A., Ashe-
rian, A., Biaggio, A., Blanco-Villasenor, A., &
Ekehammar, B. (1990). International prefer-
ences in selecting mates: A study of 37 cul-
tures. Journal of Cross-Cultural Psychology,
21(1), 5–47.
Buss, D. M., Shackelford, T. K., Kirkpatrick, L.
A., & Larsen, R. J. (2001). A half century of
mate preferences: The cultural evolution of
values. Journal of Marriage and Family,
63(2), 491–503.
Buston, P. M., & Emlen, S. T. (2003). Cognitive
processes underlying human mate choice:
The relationship between self-perception
and mate preference in Western society.
Proceedings of the National Academy of Sci-
ences, 100(15), 8805–8810.
Carroll, L. (1917). Through the looking glass:
And what Alice found there. Rand, McNally.
Coontz, S. (2006). Marriage, a history: How love
conquered marriage. New York: Penguin.
Daniels, K. R. (1989). Semen donors: Their
motivations and attitudes to their offspring.
Journal of Reproductive and Infant Psychol-
ogy, 7(2), 121–127.
Daniels, K., Blyth, E., Crawshaw, M., & Curson,
R. (2005). Previous semen donors and their
views regarding the sharing of information
with offspring. Human Reproduction, 20(6),
1670–1675.
Dunbar, R. (2012). The science of love and
betrayal. London: Faber & Faber.
Ernst, E., Jakob Ingerslev, H., Schou, O., &
Stoltenberg, M. (2007). Attitudes among
sperm donors in 1992 and 2002: A Danish
questionnaire survey. Acta Obstetricia et
Gynecologica Scandinavica, 86(3), 327–333.
Finkel, E. J., & Eastwick, P. W. (2008). Speed-
dating. Current Directions in Psychological
Science, 17(3), 193–197.
Finkel, E. J., Eastwick, P. W., & Matthews, J.
(2007). Speed-dating as an invaluable tool
for studying romantic attraction: A method-
ological primer. Personal Relationships,
14(1), 149–166.
Fiore, A. T., Taylor, L. S., Zhong, X., Mendel-
sohn, G. A., & Cheshire, C. (2010, January).
Who’s right and who writes: People, profiles,
contacts, and replies in online dating. In
2010 43rd Hawaii International Conference
on System Sciences (pp. 1–10). IEEE.
Fisman, R., Iyengar, S. S., Kamenica, E., &
Simonson, I. (2006). Gender differences in
mate selection: Evidence from a speed dating
experiment. Quarterly Journal of Economics,
121(2), 673–697.
Fletcher, G. J., Simpson, J. A., Campbell, L., &
Overall, N. C. (2015). Pair-bonding, romantic
love, and evolution: The curious case of
Homo sapiens. Perspectives on Psychological
Science, 10(1), 20–36.
Flinn, M. V. (1986). Correlates of reproductive
success in a Caribbean village. Human Ecol-
ogy, 14(2), 225–243.
Folkes, V. S. (1982). Forming relationships and
the matching hypothesis. Personality and
Social Psychology Bulletin, 8(4), 631–636.
Foote, R. H. (2010). The history of artificial
insemination: Selected notes and notables.
Journal of Animal Science, 80, 1–10.
Frazzetto, G. (2010). The science of online
dating. EMBO Reports, 11(1), 25–27.
Freeman, T., Jadva, V., Tranfield, E., & Golom-
bok, S. (2016). Online sperm donation: A
survey of the demographic characteristics,
motivations, preferences and experiences of
sperm donors on a connection website.
Human Reproduction, 31(9), 2082–2089.
Frost, J. H., Chance, Z., Norton, M. I., & Ariely,
D. (2008). People are experience goods:
Improving online dating with virtual dates.
Journal of Interactive Marketing, 22(1), 51–61.
Gale, D., & Shapley, L. S. (1962). College
admissions and the stability of marriage. The
American Mathematical Monthly, 69(1),
9–15.
Gangestad, S. W., & Buss, D. M. (1993). Patho-
gen prevalence and human mate prefer-
ences. Ethology and Sociobiology, 14(2),
89–96.
Gangestad, S. W., Thornhill, R., & Yeo, R. A.
(1994). Facial attractiveness, developmental
stability, and fluctuating asymmetry.
Ethology and Sociobiology, 15(2), 73–85.
Gillmor, C. S. (2007). Stanford, the IBM 650,
and the first trials of computer date match-
ing. IEEE Annals of the History of Comput-
ing, 29(1), 74–80.
Goode, E. (1996). Gender and courtship
entitlement: Responses to personal ads. Sex
Roles, 34(3–4), 141–169.
 EVOLUTIONARY PSYCHOLOGY AND REPRODUCTION
Grammer, K. (1989). Human courtship behav-
iour: Biological basis and cognitive processing.
Sociobiology of Sexual and Reproductive
Strategies, 147, 169.
Greely, H. T. (2016). The end of sex and the
future of human reproduction. Cambridge,
MA: Harvard University Press.
Greenlees, I. A., & McGrew, W. C. (1994). Sex
and age differences in preferences and tac-
tics of mate attraction: Analysis of published
advertisements. Ethology and Sociobiology,
15(2), 59–72.
Gustavsson, L., Johnsson, J. I., & Uller, T.
(2008). Mixed support for sexual selection
theories of mate preferences in the Swedish
population. Evolutionary Psychology, 6(4),
575–585.
Guttmacher, A. F. (1969). Artificial insemina-
tion. DePaul Law Review, 118(2), 566–583.
Hadany, L., & Comeron, J. M. (2008). Why are
sex and recombination so common? Annals of
the New York Academy of Sciences, 1133(1),
26–43.
Hargreaves, K. (2006). Constructing families
and kinship through donor insemination.
Sociology of Health & Illness, 28(3), 261–283.
Hinsz, V. B. (1989). Facial resemblance in
engaged and married couples. Journal of
Social and Personal Relationships, 6(2),
223–229.
Hitsch, G. J., Hortaçsu, A., & Ariely, D. (2010a).
Matching and sorting in online dating.
American Economic Review, 100(1),
130–163.
Hitsch, G. J., Hortaçsu, A., & Ariely, D. (2010b).
What makes you click? – Mate preferences
in online dating. Quantitative Marketing and
Economics, 8(4), 393–427.
Hollingshead, A. B. (1950). Cultural factors in
the selection of marriage mates. American
Sociological Review, 15(5), 619–627.
Kalmijn, M. (1994). Assortative mating by
cultural and economic occupational status.
American Journal of Sociology, 100(2),
422–452.
Kenrick, D. T. (2011). Sex, murder, and the
meaning of life: A psychologist investigates
how evolution, cognition, and complexity
are revolutionizing our view of human
nature. New York: Basic Books.
Kenrick, D. T., Sadalla, E. K., Groth, G., & Trost,
M. R. (1990). Evolution, traits, and the stages
495
of human courtship: Qualifying the parental
investment model. Journal of Personality,
58(1), 97–116.
Korobov, N. (2011). Mate-preference talk in
speed-dating conversations. Research on Lan-
guage and Social Interaction, 44(2), 186–209.
Kurzban, R., & Weeden, J. (2005). HurryDate:
Mate preferences in action. Evolution and
Human Behavior, 26(3), 227–244.
Lalos, A., Daniels, K., Gottlieb, C., & Lalos, O.
(2003). Recruitment and motivation of
semen providers in Sweden. Human Repro-
duction, 18(1), 212–216.
Lavoie, K., Côté, I., & de Montigny, F. (2018).
Assisted reproduction in the digital age: Sto-
ries of Canadian sperm donors offering their
gametes online via introduction websites. The
Journal of Men’s Studies, 26(2), 184–202.
Lenton, A. P., Fasolo, B., & Todd, P. M. (2010).
Who is in your shopping cart? Expected and
experienced effects of choice abundance in
the online dating context. In N. Kock (Ed.),
Evolutionary Psychology and Information
Systems Research (pp. 149–167). Boston,
MA: Springer.
Li, N. P., Bailey, J. M., Kenrick, D. T., &
Linsenmeier, J. A. (2002). The necessities and
luxuries of mate preferences: Testing the
tradeoffs. Journal of Personality and Social
Psychology, 82(6), 947.
Lively, C. M., & Morran, L. T. (2014). The
ecology of sexual reproduction. Journal of
Evolutionary Biology, 27(7), 1292–1303.
Little, A. C., Burt, D. M., & Perrett, D. I. (2006).
Assortative mating for perceived facial
personality traits. Personality and Individual
Differences, 40(5), 973–984.
Lynn, M., & Bolig, R. (1985). Personal advertise-
ments: Sources of data about relationships.
Journal of Social and Personal Relationships,
2(3), 377–383.
Lynn, M., & Shurgot, B. A. (1984). Responses
to lonely hearts advertisements: Effects of
reported physical attractiveness, physique,
and coloration. Personality and Social Psy-
chology Bulletin, 10(3), 349–357.
Mare, R. D. (1991). Five decades of educational
assortative mating. American Sociological
Review, 56(1) 15–32.
McLaughlin, C., Vitak, J., & Crouse, J. (2011).
Online identity construction and expectation
of future interaction. In 2011 44th Hawaii
496 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
International Conference on System Sciences
(pp. 1–10). IEEE.
Miller, G. (2000). The mating mind: How sexual
choice shaped the evolution of human
nature. New York: Anchor Books.
Miller, G. F., & Todd, P. M. (1998). Mate choice
turns cognitive. Trends in Cognitive Sciences,
2(5), 190–198.
Minervini, B. P., & McAndrew, F. T. (2006). The
mating strategies and mate preferences of
mail order brides. Cross-Cultural Research,
40(2), 111–129.
Mulder, M. B. (1990). Kipsigis women’s prefer-
ences for wealthy men: Evidence for female
choice in mammals? Behavioral Ecology and
Sociobiology, 27(4), 255–264.
Ombelet, W., & Van Robays, J. (2015). Artificial
insemination history: Hurdles and mile-
stones. Facts, Views & Vision in ObGyn, 7(2),
137–143.
Otto, S. P. (2009). The evolutionary enigma of
sex. American Naturalist, 174(S1), S1–S14.
Overbeek, G., Nelemans, S. A., Karremans, J., &
Engels, R. C. (2013). The malleability of mate
selection in speed-dating events. Archives of
Sexual Behavior, 42(7), 1163–1171.
Oyer, P. (2014). Everything I ever needed to
know about economics I learned from online
dating. Brighton, MA: Harvard Business
Review Press.
Pawłowski, B., & Dunbar, R. I. (1999). Impact of
market value on human mate choice deci-
sions. Proceedings of the Royal Society of
London. Series B: Biological Sciences,
266(1416), 281–285.
Pawlowski, B., & Dunbar, R. I. M. (2001).
Human mate choice strategies. Economics in
nature. Social dilemmas, mate choice and
biological markets, 187–202.
Pawłowski, B., & Koziel, S. (2002). The impact
of traits offered in personal advertisements
on response rates. Evolution and Human
Behavior, 23(2), 139–149.
Pennings, G. (1995). Should donors have the
right to decide who receives their gametes?
Human Reproduction, 10(10), 2736–2740.
Penton-Voak, I. S., Perrett, D. I., & Peirce, J. W.
(1999). Computer graphic studies of the role
of facial similarity in judgements of attractive-
ness. Current Psychology, 18(1), 104–117.
Pérusse, D. (1993). Cultural and reproductive
success in industrial societies: Testing the
relationship at the proximate and ultimate
levels. Behavioral and Brain Sciences, 16(2),
267–283.
Piazza, J., & Bering, J. M. (2009). Evolutionary
cyber-psychology: Applying an evolutionary
framework to Internet behavior. Computers
in Human Behavior, 25(6), 1258–1269.
Price, R. A., & Vandenberg, S. G. (1979).
Matching for physical attractiveness in mar-
ried couples. Personality and Social Psychol-
ogy Bulletin, 5(3), 398–400.
Puts, D. A. (2010). Beauty and the beast:
Mechanisms of sexual selection in humans.
Evolution and Human Behavior, 31(3),
157–175.
Quinlan, R. J. (2007). Human parental effort
and environmental risk. Proceedings of the
Royal Society B: Biological Sciences,
274(1606), 121–125.
Renninger, L. A., Wade, T. J., & Grammer, K.
(2004). Getting that female glance: Patterns
and consequences of male nonverbal behav-
ior in courtship contexts. Evolution and
Human Behavior, 25(6), 416–431.
Roberts, S. C., Miner, E. J., & Shackelford, T. K.
(2010). The future of an applied evolutionary
psychology for human partnerships. Review
of General Psychology, 14(4), 318–329.
Rockman, H. (1994). Matchmaker make me a
match: The art and conventions of Jewish
arranged marriages. Sexual and Marital
Therapy, 9(3), 277–284.
Rodino, I. S., Burton, P. J., & Sanders, K. A.
(2011). Mating by proxy: A novel perspective
to donor conception. Fertility and Sterility,
96(4), 998–1001.
Rosenfeld, M. J., & Thomas, R. J. (2012).
Searching for a mate: The rise of the Internet
as a social intermediary. American Sociologi-
cal Review, 77(4), 523–547.
Rosenfeld, M. J., Thomas, R. J., & Hausen, S.
(2019). Disintermediating your friends: How
online dating in the United States displaces
other ways of meeting. Proceedings of the
National Academy of Sciences, 116(36),
17753–17758.
Rutherford, A. (2018). The book of humans:
The story of how we became us. Hachette
UK.
Scheib, J. E. (1994). Sperm donor selection and
the psychology of female mate choice. Ethol-
ogy and Sociobiology, 15(3), 113–129.
 EVOLUTIONARY PSYCHOLOGY AND REPRODUCTION
Scheib, J. E. (1997). Female choice in the con-
text of donor insemination. In P. A. Gowaty
(Ed.), Feminism and Evolutionary Biology:
Boundaries, Intersections and Frontiers
(pp. 489–504). New York: Chapman & Hall.
Schnarch, D. (1997). Sex, intimacy, and the
Internet. Journal of Sex Education and Ther-
apy, 22(1), 15–20.
Schover, L. R., Rothmann, S. A., & Collins, R. L.
(1992). The personality and motivation of
semen donors: A comparison with oocyte
donors. Human Reproduction, 7(4), 575–579.
Schwartz, C. R. (2013). Trends and variation in
assortative mating: Causes and consequences.
Annual Review of Sociology, 39, 451–470.
Seabright, P. (2012). The war of the sexes: How
conflict and cooperation have shaped men
and women from prehistory to the present.
Princeton, NJ: Princeton University Press.
Sear, R. (2015). Evolutionary contributions to
the study of human fertility. Population
Studies, 69(sup1), S39–S55.
Simon, H. A. (1956). Rational choice and the
structure of the environment. Psychological
Review, 63(2), 129–138.
Sitton, S., & Blanchard, S. (1995). Men’s prefer-
ences in romantic partners: Obesity vs addic-
tion. Psychological Reports, 77(3_suppl),
1185–1186.
Skopek, J., Schmitz, A., & Blossfeld, H. P. (2011).
The gendered dynamics of age preferences –
empirical evidence from online dating. ZfF –
Zeitschrift für Familienforschung/Journal of
Family Research, 23(3), 267–290.
Smith, J. M. (1971). What use is sex? Journal of
Theoretical Biology, 30(2), 319–335.
Smith, J. M., & Maynard-Smith, J. (1978). The
evolution of sex (Vol. 4). Cambridge:
Cambridge University Press.
Su, X., & Hu, H. (2019). Gender-specific prefer-
ence in online dating. EPJ Data Science, 8(1),
12.
Symons, D. (1979). The evolution of human
sexuality. New York: Oxford University Press.
Thiessen, D. (1994). Environmental tracking by
females. Human Nature, 5(2), 167–202.
Thiessen, D., & Gregg, B. (1980). Human assor-
tative mating and genetic equilibrium: An
evolutionary perspective. Ethology and
Socio­biology, 1(2), 111–140.
Thiessen, D., Young, R. K., & Delgado, M.
(1997). Social pressures for assortative
497
mating. Personality and Individual Differ-
ences, 22(2), 157–164.
Thomas, A. G., Jonason, P. K., Blackburn, J.,
Kennair, L. E. O., Lowe, R., Malouff, J., & Li,
N. P. (2019). Mate preference priorities in the
East and West: A cross-cultural test of the
mate preference priority model. Journal of
Personality, 88(3), 606–620.
Todd, P. M., Penke, L., Fasolo, B., & Lenton, A. P.
(2007). Different cognitive processes underlie
human mate choices and mate preferences.
Proceedings of the National Academy of
Sciences, 104(38), 15011–15016.
Toma, C. L., & Hancock, J. T. (2010). Looks and
lies: The role of physical attractiveness in
online dating self-presentation and deception.
Communication Research, 37(3), 335–351.
Toma, C. L., Hancock, J. T., & Ellison, N. B.
(2008). Separating fact from fiction: An
examination of deceptive self-presentation in
online dating profiles. Personality and Social
Psychology Bulletin, 34(8), 1023–1036.
Tooby, J., & Cosmides, L. (2005). In David M.
Buss (Ed.), Evolutionary psychology: Concep-
tual foundations. Evolutionary psychology
handbook. New York: Wiley. 1–55.
Townsend, J. M. (1989). Mate selection criteria:
A pilot study. Ethology and Sociobiology,
10(4), 241–253.
Trivers, R., (1972). Parental investment and
sexual selection. In Bernard Campbell (Ed.),
Sexual Selection & the Descent of Man (pp.
136–179). New York: Aldine de Gruyter,
New York.
Valkenburg, P. M., & Peter, J. (2007). Who visits
online dating sites? Exploring some charac-
teristics of online daters. CyberPsychology &
Behavior, 10(6), 849–852.
Van den Broeck, U., Vandermeeren, M., Vander-
schueren, D., Enzlin, P., Demyttenaere, K., &
D’Hooghe, T. (2012). A systematic review of
sperm donors: Demographic characteristics,
attitudes, motives and experiences of the
process of sperm donation. Human Repro-
duction Update, 19(1), 37–51.
Vandenberg, S. G. (1972). Assortative mating,
or who marries whom? Behavior Genetics,
2(2–3), 127–157.
Walker, R. S., Hill, K. R., Flinn, M. V., & Ells-
worth, R. M. (2011). Evolutionary history of
hunter-gatherer marriage practices. PloS
One, 6(4), e19066.
498 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Warren, B. L. (1966). A multiple variable
approach to the assortative mating phenom-
enon. Biodemography and Social Biology,
13(4), 285–290.
Waynforth, D., & Dunbar, R. I. (1995). Condi-
tional mate choice strategies in humans:
Evidence from ‘Lonely Hearts’ advertise-
ments. Behaviour, 132(9), 755–779.
Whyte, S., Savage, D. A., & Torgler, B. (2017).
Online sperm donors: the impact of family,
friends, personality and risk perception on
behaviour. Reproductive Biomedicine Online,
35(6), 723–732.
Whyte, S., & Torgler, B. (2015). Selection criteria
in the search for a sperm donor: Behavioural
traits versus physical appearance. Journal of
Bioeconomics, 17(2), 151–171.
Whyte, S., & Torgler, B. (2016a). Assortative
mating in the online market for sperm
donation. Journal of Bioeconomics, 18(3),
169–194.
Whyte, S., & Torgler, B. (2016b). Determinants
of online sperm donor success: How women
choose. Applied Economics Letters, 23(8),
592–596.
Whyte, S., & Torgler, B. (2017a). Preference
versus choice in online dating. Cyberpsychol-
ogy, Behavior, and Social Networking, 20(3),
150–156.
Whyte, S., & Torgler, B. (2017b). Things change
with age: Educational assortment in online
dating. Personality and Individual Differ-
ences, 109, 5–11.
Whyte, S., Brooks, R. C., Chan, H. F., & Torgler,
B. (2019). Do certain personality traits
provide a mating market competitive
advantage? Sex, offspring & the big 5.
Personality and Individual Differences, 139,
158–169.
Whyte, S., Chan, H. F., & Torgler, B. (2018). Do
men and women know what they want? Sex
differences in online daters’ educational
preferences. Psychological Science, 29(8),
1370–1375.
Whyte, S., Torgler, B., & Harrison, K. L. (2016).
What women want in their sperm donor: A
study of more than 1000 women’s sperm
donor selections. Economics & Human Biol-
ogy, 23, 1–9.
Williams, G. C. (1966). Adaptation and natural
selection: A critique of some current evolu-
tionary thought. Princeton, NJ: Princeton
University Press.
Wilson, D. S. (2018). This view of life: Complet-
ing the Darwinian revolution. New York: Pen-
guin Random House.
Woestenburg, N. O., Winter, H. B., & Janssens,
P. M. (2016). What motivates men to offer
sperm donation via the internet? Psychology,
Health & Medicine, 21(4), 424–430.
Woll, S. B., & Young, P. (1989). Looking for Mr.
or Ms. Right: Self-presentation in videodat-
ing. Journal of Marriage and the Family,
51(2), 483–488.
Yee, S. (2009). ‘Gift without a price tag’: altru-
ism in anonymous semen donation. Human
Reproduction, 24(1), 3–13.
Young, L. J., & Wang, Z. (2004). The neurobiol-
ogy of pair bonding. Nature Neuroscience,
7(10), 1048.
Zimmer, C. (2009). On the origin of sexual repro-
duction. Science, 324(5932), 1254–1256.

Evolutionary Psychology
and Cyberwarfare
CYBERWARFARE AND MEMES
Conflicts involving terrorism and asymmetric
adversaries, who may be state or non-state
actors, are fundamentally information wars of
the mind featuring psychosocial manipulation,
often as precursors to physical violence which
they instigate. Cyberwarfare, or cyber warfare,
encompasses a loose set of adversarial pro-
cesses involving computers, software, and net-
works (RAND Corporation, 2019), of which a
subset involves influencing the mental state of
individuals and populations through social
media, such as influencing elections (Council
on Foreign Relations, 2019) or sowing conflict
among subcultures in a nation (US Senate,
2019). The long war against terrorism is funda-
mentally a war of ideas, and at the core in this
information war are memes (Finkelstein,
2011). A common metaphor for memes is
mind viruses (Asher, 2005; Brodie, 1996).
Memes can alter the psychology of individ-
uals, their values, and their consequent behav-
ior. The aggregated behavior of individuals,
25
Robert Finkelstein
from which culture emerges, then changes,
along with the values underlying the culture.
This results in a changed culture, the changes
being beneficial or harmful to people com-
prising the culture. The prospective disci-
pline of cultural engineering (or accelerated
cultural evolution) may be used intentionally
to modify what is considered a dysfunctional
culture or subculture (Finkelstein, 2012).
It was hypothesized (Dawkins, 1976) that,
on an evolutionary timescale, memes may
affect the development of the human mind,
and that memes needed for survival by our
ancient ancestors (e.g., Homo habilis, Homo
erectus, and Homo heidelbergensis) led to the
emergence of enlarged brains and more cog-
nitively advanced minds for Homo sapiens.
Evolutionary psychology contends that
Darwinian evolutionary processes of natu-
ral selection apply to the brain, mind, and
resulting psychological traits, e.g., memory,
perception, and language, just as they do to
physiological traits (Buss, 2005). Darwinian
evolution is a well-tested scientific theory
500 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
that explains how complexity can arise from
simplicity, and so evolutionary psychology
is expected to provide a basis for explain-
ing the origin of the human brain, perhaps
the most complex entity in the known uni-
verse (Aunger, 2002). Some traits, physical
or mental, which were previously optimal for
human survival, may no longer be as benefi-
cial in the context of modern technology and
cultural values. Racism and xenophobia, for
example, may have served to benefit survival
in small bands of hunter-gatherers for hun-
dreds of millennia, but are now harmful in
our complex, technologically advanced soci-
eties. The psychological states that give rise
to these phenomena are no longer rational,
and the behavior they cause cannot be toler-
ated in a democratic society. Nevertheless,
racism and xenophobia, among other obso-
lete mental programming, remain endemic in
society as a remnant of once-useful evolved
psychology, like an unnecessary inflamed
appendix, with memes used to inflame emo-
tions that are otherwise under control or dor-
mant within susceptible recipients.
Over an evolutionary timescale the meme
is postulated to affect human psychological
traits arising from natural selection, i.e., the
evolved psychology arising from changes in
the brain and mind that drive individual and
cultural behavior. But the reverse can also be
conjectured: that the consequences of evolved
psychology affect the memes to which we are
now most susceptible, the memes propagating
through social media every second, flooding
the Twitterverse and impacting a multitude
of minds. Memes having profound impact
can alter the values of susceptible individuals
and cause them to engage in behavior detri-
mental to them and to society. Contemplating
Darwinian evolutionary timescales for the
effects of memes on the brain and mind may
suggest interesting phenomena for research;
but contemplating memes in the current
timeframe, in the context of evolved psy-
chology, is more compelling given their imme-
diate consequences. The near-term evolution
of human values, behavior, and culture caused
by memes, like the evolution of technology,
is (metaphorically) Lamarckian rather than
Darwinian (Blackmore, 1999). Human psy-
chology, shaped over eons by Darwinian
evolution, reacts in the near term to memes
via Lamarckian evolution, in the metaphori-
cal sense that meme-caused changes in the
behavior of an individual can cause changes in
the behavior of others. (Lamarckian evolution
occurs literally when the environment changes
the phenotype of an organism such that the
organism is better adapted to the environmental
factor that caused the change (Hull, 2000), and
where the genotype can then pass the improved
adaptation to the organism’s progeny.)
WHAT IS A MEME?
The literature addressing memes reveals a
plethora of definitions, with most being varia-
tions of the neologism coined by Dawkins
(1976), although there are indications of earlier
roots. Dawkins defined the meme as a self-
reproducing and propagating information
structure analogous to a gene in biology. He
focused on the meme as a replicator, analogous
to the gene, able to affect human evolution
through the evolutionary algorithm of varia-
tion, replication, and differential fitness, and
then eventual physical alteration of the brain
due to the beneficial impact on human survival
and reproduction of certain memes, such as
detailed instructions requiring extensive mental
information processing and storage about how
to make a flint cutting tool, how to hunt a
wooly mammoth, how to find water in the
desert, or how to make fire. Thus, he claimed,
the meme had evolutionary effects on human
physiology, psychology, and culture. He pro-
posed that a meme’s information structure can
have many forms, including words, symbols,
images, icons, and artifacts (Dawkins, 1976).
In the near term, memes can change the
values and behavior of individuals and the
culture they comprise. Memes can cause
individuals and their cultures to evolve in the
 EVOLUTIONARY PSYCHOLOGY AND CYBERWARFARE
sense of changing, albeit not in the sense of
Darwinian (and genetically based) evolution
of the human brain, mind, and psychology.
The latter phenomenon, the original empha-
sis for Dawkins, has yet to be scientifically
established, while the former phenomenon
can be regularly observed.
Most of the definitions offered by meme
researchers agree that a meme is a unit
(whatever that means) of cultural transmis-
sion (or a unit of information – whatever
that means), where culture may be defined
as the total pattern of behavior (and its prod-
ucts) of a population of agents, embodied in
thought, behavior, and artifacts, and depend-
ent upon the capacity for learning and trans-
mitting knowledge to succeeding generations
(Finkelstein, 2006).
A few of the many definitions extracted
from the literature include:
• A self-reproducing and propagating information
structure analogous to a gene in biology
• A unit of cultural transmission (or a unit of imita-
tion) that is a replicator that propagates in the
meme pool leaping from brain to brain via (in a
broad sense) imitation; examples: ‘tunes, ideas,
catch-phrases, clothes fashions, ways of making
pots or of building arches’ (Dawkins, 1976: 192)
• Ideas that embody programs for their own
retransmission or propagation
• Actively contagious ideas or thoughts
• Shared elements of a culture learned through imi-
tation from others – with culture being defined
rather broadly to include ideas, behaviors, and
physical objects
• An element of a culture transmitted by non-
genetic means, especially imitation
• Information patterns infecting human minds
• A (cognitive) information structure able to rep-
licate using human hosts and to influence their
behavior to promote replication
• Cultural information units that are the smallest
elements that self-replicate with reliability and
fecundity
• A unit of cultural information as it is represented
in the brain
• A pattern of information, one that happens to
have evolved a form which induces people to
repeat that pattern
501
• A contagious information pattern that replicates
by parasitically infecting human minds and alter-
ing their behavior, causing them to propagate
the pattern. Individual slogans, catch-phrases,
melodies, icons, inventions, and fashions are
typical memes. An idea or information pattern is
not a meme until it causes someone to replicate
it, to repeat it to someone else. All transmitted
knowledge is memetic
• The smallest idea that can copy itself while
remaining self-contained and intact – essentially
sets of instructions that can be followed to pro-
duce behavior
Some have contended that a crisp definition of
‘unit of information’ or ‘unit of cultural trans-
mission’ is not critical to a theory of memes,
e.g., Darwin’s Theory of Evolution was scien-
tifically useful before anyone knew about
genes and DNA. But a pragmatic definition
and associated metrics would provide a better
foundation for research and would facilitate
the ability to generate testable predictions and
falsifiable hypotheses, and control effects,
which DNA and genetic engineering now pro-
vides to Darwin’s theory. Darwin’s theory was
scientifically useful before anyone knew about
genes and DNA because Darwin and other
scientists had physical objects (e.g., fossils or
living physiological structures) to measure.
Metrics existed with which to compare and
contrast entities. A coherent theory could
be developed in which phenomena could be
quantified and predicted. This is not the case
for memes, and there is not as yet an analo-
gous Theory of Memetics.
The author (Finkelstein, 2011) distin-
guishes between memes that can be described
by their external phenomena (e-memes) and
internal phenomena (i-memes). E-memes are
manifested by their effects on human behav-
ior and culture. I-memes are manifested
by their effects on an individual’s neuronal
behavior and brain. E-memes and i-memes
are not two types of memes, but rather two
effects or manifestations of memes.
The tools and techniques for studying the
two manifestations of memes differ. E-memes
can be studied via their simulated or actual
502 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
propagation over social networks (modern
social media are especially efficient and effec-
tive for propagating memes). Tools and tech-
niques include: Internet and phone text and
voice message propagation; rumor and gossip
propagation; buzz and viral marketing. There
are also neuroeconomics behavioral experi-
ments involving game theory, risk, attention
and awareness, learning, valuation, motiva-
tion, emotion, behavior, trust and attachment,
and addictive behavior. I-memes can be stud-
ied using tools and techniques such as func-
tional magnetic resonance imaging (fMRI) or
other kinds of neuroimaging; genetic profil-
ing; p­sycho-pharmacological manipulations;
psycho-physiology (EMG, ERP, and EEG);
behavioral measures; psychological testing;
blood chemistry; hormone analysis; neuro-
chemical reactions; and single neuron record-
ing. The study of the two manifestations of
memes must be integrated with a synthesis and
multi-disciplinary study of the phenomenon.
While Dawkins focused on the meme as
a replicator, analogous to the gene, able to
affect human evolution through the evolu-
tionary algorithm of variation, replication,
and differential fitness, for near-term practi-
cal applications the relevant characteristics of
the meme are that it consists of information
which persists, propagates, and influences
human behavior. A metrics-based, pragmatic
definition of the meme can be constructed to
distinguish a meme from other sorts of infor-
mation (such as common daily utterances
like ‘it’s time for lunch’).
A meme is information which propagates,
persists, and has impact. Information, in
Claude Shannon’s definition (and associated
equations), is that which reduces uncertainty
(Shannon, 1948). A message carries informa-
tion inasmuch as it conveys something not
already known. It is the difference between
two states of uncertainty before and after a
message has been received. Psychology, in
the form of subjective perception, is involved
in evaluating the states of uncertainty. The
same message can have different influence
or impacts, depending on the states of the
recipients; a meme, as a subset of informa-
tion, could have different consequences for
different recipients. For example, if a recipi-
ent is already familiar with the content of a
message, it does not reduce uncertainty at all,
i.e., it is not information for that recipient.
To distinguish memes from other kinds
of information, an elaboration of the defini-
tion can invoke measurable thresholds, to be
determined, for propagation and persistence.
Impact might take place in the individual, in
terms of neurophysiological changes (e.g., as
detected by a brain scan) or manifestations
of behavior; or it might take place in soci-
ety (groups of individuals) as manifested by
behavior. Metrics can be determined (each
with submetrics) to facilitate the measure-
ment and comparison of memes. Additional
metrics may be defined based on the results
of experiments on brain processes with tools
such as fMRI. Other tools for meme research
include the modeling and simulation of
social networks and information propagation
through social networks, as well as the tools
and techniques of neuroeconomics, which is
the study of how the brain interacts with the
external environment to make choices (deci-
sions), and which combines neuroscience,
economics, and psychology.
Metrics are needed to establish a scientific
and engineering basis for memetics. Research
is needed to determine the metrics. Based on
the Information – Propagation – Impact –
Persistence (IPIP) definition for a meme,
some makeshift metric examples include: the
number, type, and dispersion of recipients;
the duration of transmission and memory;
the degree of entropy; and the individual and
societal consequences of the meme.
The relative importance of meme size
(e.g., small, medium, large) remains to be
determined, but shorter (smaller) memes are
easier to remember and propagate over social
media than longer ones, and are likely to
have greater immediate impact. Entropy as a
metric of information may be better for deter-
mining the size of message than the number
of words (or bits) because it can be applied to
 EVOLUTIONARY PSYCHOLOGY AND CYBERWARFARE
any language as well as symbols, signs, and
images – semiotics in general. Larger memes
may be memeplexes, e.g., the typical human
brain can only hold about 7 ‘chunks’ of infor-
mation in short-term memory (McLeod, 2009).
Claude Shannon’s mathematical theory
of communication (Shannon, 1948) meas-
ures the amount of information in any mes-
sage using a function identical to that of
Boltzmann’s formulation of entropy (Sharp
and Matschinsky, 2015). The use of this
entropy is noncontroversial as long as it is
used to measure the freedom of choice in
messages and not semantic content (i.e., a
specific message is selected from a larger
set). For example, the information (entropy)
needed to select one word out of a vocabulary
of 200 words is 7.6 bits (S = log2 200 = 7.6
bits). That is, the word is within the first 100
words or the second 100 words. It takes one
bit to determine that, or one guess as to which
of the two sets it was in – if the word is in the
first 100 words, it takes another bit (guess) to
determine which of the two sets of 50 words
it is in; and so on. It takes between 7 and 8 bits
(guesses) to find the one word in 200 words.
Likewise, a 50-word meme requires 380 bits
(for a 200-word vocabulary). Nothing is indi-
cated about the meaning or significance of
the message that results from selecting and
stringing together the 50 words.
A problem arises when entropy is used as
a measure of informational order and disor-
der. For example, a manager has a desk with
mounds of paper covering it – is this a high
or low entropy case? The manager’s associ-
ate cannot find a paper he is searching for
until he has made a large number of guesses,
separating the piles of papers – a very high
entropy case. But the manager knows exactly
where every paper is located – the mounds
of paper are quite organized in her mind; it
would take her no guesses to find the crucial
paper. It is then a very low entropy case. Both
cases concern the same set of items, but two
different observers. Entropy then becomes
a subjective measure of information (or a
measure of the information contained in the
503
mind and psychology of the observer), and
this can be a disturbing concept to some.
The entropy of a meme, whether long or
short, can be estimated and calculated from the
individual letters or symbols, i.e., based on
the total number in the originating set, or from
the individual words, based on the vocabulary
in the originating set. For example, the Oxford
English Dictionary contains 616,500 words.
The average vocabulary of an American English
speaker is estimated to be between 10,000 and
20,000 words, or optimistically between 50,000
and 70,000 words (de Leon Huld, 2020). The
average American would actively use only a
fraction of that vocabulary. Typically, there
are about 800 words per page with 12-point
Times Roman font, although publications vary
considerably in the number of words, based
on page size, format, and number of pages.
Shannon (1948) calculated the entropy per let-
ter of an eight-letter chunk of English as 2.3 bits
per letter, using a 26-character set (although a
76-symbol set, including 26 letters, might be
more appropriate). Assuming a vocabulary of
30,000 words the entropy of a single word is
about 15 bits (S = log2 30,000 = 15 bits). This
kind of metric for information might be used in
engineering memes for maximum efficacy.
Metrics for propagation may be based, for
example, on the number, type, and dispersion
of recipients of memes. For example, neighbor-
hood gossip may propagate among 10 people
and persist for 10 hours, while a fundamental-
ist insurgency may propagate among 100,000
people and persist for 100,000 hours (about 10
years). The persistence of memes, for example,
may be measured in terms of the duration of
their transmission and re-transmission and the
duration in the memory of the recipients. The
impact of memes may be measured in terms of
their consequences on individuals and society.
MEME TRANSMISSION
The transmission of the meme resembles
Claude Shannon’s iconic schematic of a
504 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
general communications system: an informa-
tion source sends a message via a transmitter
of some sort, which issues a signal of some
kind, which is usually adversely affected by
noise. The signal is received by a receiver of
some kind, which issues the message, perhaps
altered by noise, to the destination. This com-
munications template was modified by
Gunders and Brown (Truszkowski et al., 2020)
into a model of a meme life-cycle, known as
the Component Meme Concept (CMC):
• A memetic ‘engineer’ intentionally or uninten-
tionally creates a meme
• A hook attracts people to the meme
• Bait is the desired result promised by the meme
to attract people to contemplate it and propa-
gate it to others
• The vector is the medium used to transport the
original meme and its subsequent propagation
(there may be multiple vectors)
• The host refers to the carrier that sent the original
meme and subsequent propagation (there may
be multiple hosts)
• The memotype refers to the content of the meme
• The sociotype is the social and cultural environ-
ment of the memotype
Consider the following example known in the
press as ‘Pizzagate’, where some of its facts
have been changed for convenience in our
example. It was claimed (falsely) that a child
sex ring, managed by the wife of a former
president, is being run out of the basement of
a Washington, DC pizza parlor. To those able
to think critically, this extraordinary claim
would seem to require extraordinary evidence
to be believed, or even considered for more
than a few seconds. Sadly, for many the meme
life-cycle is resistant to critical thinking.
• Memetic engineer: <John Smith>
• Hook: <Child Sex Ring>
• Bait: <It Will Make You Shocked And Angry>
• Vector: <Twitterverse; Then Internet, Phone Text,
TV, Radio, And Newspapers>
• Host: <Twitter; Then Facebook, Etc.>
• Memotype: <Verbal Description>
• Sociotype: <Conspiracy Fanatics; Ill-Educated
Young Males>
The meme influenced the behavior of at least
one recipient of the meme, a young man who
then armed himself and invaded the pizza
parlor to free the supposedly enslaved chil-
dren. He was sentenced to four years in
prison. While many Internet memes, like cat
videos, may be amusing, other memes may
motivate recipients to commit terrorist acts
or perpetrate mass murders.
TYPES OF MEMES
Most people today associate memes with the
Internet and social media, but over the eons
they have been much more than that. As
information that propagates, has impact, and
persists (IPIP), memes may consist of many
different forms, including words, ideas, sym-
bols, icons, logos, tunes, poems, catch-
phrases, fashion, technological processes
(e.g., making arrowheads or gumbo), fables,
religion, graffiti, images, novels, movies,
metaphors, and narratives. A meme that sat-
isfies the IPIP criteria may consist of:
• a simple exclamation, like the military ‘hooah!’ or
the religious ‘Allah!’
• a catchphrase, slogan, or exhortation, such as
‘remember the Maine!’, ‘Death to America!’,
‘Jesus loves you!’, ‘When it absolutely, positively
has to get there overnight!’, ‘You’ll wonder where
the yellow went when you brush your teeth with
Pepsodent!’
• a sound-bite, prayer, incitement, or joke (‘Why
does the chicken cross the road?’)
• neighborhood gossip or a text message; a news-
paper article or an urban legend
• a flyer, poster, advertisement, notice, propaganda
(‘Make America Great Again!’)
• political speech (e.g., the Gettysburg Address),
sermon (e.g., Martin Luther King Jr’s ‘I Have a
Dream’ speech), polemic, article, book, or blog
• a recipe or an entire cookbook; a political mani-
festo or ideology; a religious canon, such as the
Bible, Koran, Vedas, or the Pali; a dictionary; an
Army field manual
• instructions for building a cathedral or a nuclear
submarine
 EVOLUTIONARY PSYCHOLOGY AND CYBERWARFARE
• instructions for growing wheat or hunting wooly
mammoths; a burial ritual; instructions for making
a flint-cutting tool, a spear, a wheel, or fire
Some memes are literally etched in stone or
clay tablets, others are written on parchment
or paper, or painted on canvas (e.g., the Mona
Lisa or Edvard Munch’s ‘The Scream’), or
sculpted from stone (e.g., Michelangelo’s
‘David’). Whether the memes that survive for
generations alter our physical brain as
Dawkins (1976) suggested, they can influ-
ence our values, behavior, or culture in the
context of the psychology that has evolved.
Shorter-duration memes, such as those that
persist a few days, weeks or months, can also
affect our behavior and even our values and
culture, in the context of the evolved psy-
chology, such as by altering political or con-
sumer preferences. Even memes with brief
persistence can be propagated widely via
social media and have significant immediate
impact on their recipients.
MEMETICS AND NEUROCOGNITIVE
WARFARE
Foreign or domestic terrorism is the act or
threat of violence intended to influence an
audience. The memes that emerge from a
violent terrorist act (e.g., words and images
about the act), or memes that merely threaten
violent acts, can adversely alter the psycho-
logical state of a nation. Some terrorist
memes may persist for days or weeks, while
others persist for years. It is critical for
national defense to be able to counter adver-
sarial memes. Words matter. Over the centu-
ries, political and ideological propaganda
have proven to be potent. Lies become the
truth and the truth becomes lies (Orwell,
1946; Poole, 2006).
Military memetics is the application of
memes for national security (i.e., a compo-
nent of cyberwarfare), and it may be con-
sidered a subset of neurocognitive warfare.
505
Neurocognitive warfare targets an adversary’s
cognitive and psychological functions, and the
physiological and behavioral consequences of
those functions. The means for neurocogni-
tive warfare could include drugs, microbes,
toxins, and neuroweapons (Giordano, 2017)
such as microwaves and infrasound.
In the more than 40 years since the meme
was identified as an entity of interest, there
has not been substantial research to deter-
mine whether memetics can be placed on a
scientific basis so that its phenomena and
effects can be quantified, predicted, and con-
trolled. Most of the research of memetics,
such as it is, has been conducted in scattered
efforts, often as personal projects on the part
of dedicated academics, many outside the
United States. There have been few signifi-
cant, coherent efforts to explore whether or
how a scientific framework, and a consequent
engineering discipline, can be established for
memetics. The attempt to establish a scien-
tific basis for memetics is important. For
example, within a suitable memetics frame-
work could be the means to prevent irrational
conflict and promote rational solutions to
endemic national and international problems,
and to develop effective countermeasures
to adversarial manipulation of social media
to influence population behavior, such as
attempts to sway voting in elections or to
instigate hatred and violence between groups.
Of course, without safeguards memetics is a
double-edged sword.
Useful technology can be successfully
developed without formal engineering and
scientific disciplines through trial and error,
as has occurred over eons, such as mak-
ing weapons or preparing food. People
have been buying and selling (or trading)
goods and services for thousands of years.
Over the millennia, survival in the market-
place sharpened the methods – the tools and
techniques – of marketing, i.e., what goods
and services to produce and how to distribute
and sell them. Success at marketing depends
on knowing the needs and wants of the cus-
tomer and convincing the customer that you
506 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
can satisfy those needs and wants. At the end
of the marketing process, selling success-
fully depends on the ability to convince the
customer to part with something of value
(e.g., money, item, or service) in exchange
for what is being sold. The development of
marketing methods accelerated during the
20th century in the age of mass communica-
tion, including the refinement of advertising
campaigns in the media age of radio, televi-
sion, magazines, newspapers, and now social
media, to convince the prospective buyer that
the advertised product is just what he or she
needs or wants. The tools and techniques
advanced over the years largely by trial and
error, honed by lessons learned with the
countless buying and selling of myriad goods
and services. Likewise through trial and error
the marketing of ideas advanced with hard-
earned experience learned from public rela-
tions, political propaganda, and ideological
influencing campaigns.
There have been a number of occasional
projects concerning, or related to, memet-
ics sponsored by the Defense Advanced
Research Projects Agency (DARPA), an
agency of the Department of Defense focused
on the research and development of emerging
technologies. These projects included:
• 2006: the Epidemiology of Ideas, intended to pre-
dict changing culture and examine the creation,
propagation, and impact of ideas in the context
of the processes by which diseases propagate
(Lynch, 1996)
• 2006–2009: Military Memetics, to determine
whether memetics can be established as a sci-
ence with the ability to explain and predict phe-
nomena; to define memes and memetic metrics;
and to use tools and techniques to examine
Information Propagation, Impact, and Persistence
• 2011–2012: Social Media in Strategic
Communications, to develop automated tools and
techniques for detecting, identifying, evaluating,
and countering adversarial memes
• 2011–2012: Narrative Networks, to perform quan-
titative analysis of narratives, investigate their
effects on human psychology and neurobiology,
and determine their influence on individuals and
culture
The US military could undertake meme and
counter-meme operations involving national
security because it has long engaged in psy-
chological operations (PSYOP) to influence
or reinforce foreign attitudes and behavior
that are favorable to the military’s (and
nation’s) objectives. PSYOP also employs
selected information to affect foreign audi-
ences’ emotions, motives, and objective rea-
soning. The ultimate goal is to alter the
behavior of foreign governments, organiza-
tions, groups, and individuals so that they
will be accommodating to US military objec-
tives. PSYOP also exploits the psychological
vulnerabilities of US adversaries to induce
fear and confusion in order to destroy their
morale and disrupt their operations (Joint
Publication, 2003).
The US military also has longstanding
tools and techniques for Military Deception
(MILDEC) (Joint Publication, 2012). This
consists of actions intended to deliberately
mislead an adversary’s military decision-
makers concerning friendly military capabili-
ties, intentions, and operations. The goal is to
cause the adversary to act (or not act) in ways
favorable to friendly forces. The US military
is expressly forbidden to use MILDEC on the
US public, Congress, or news media.
The US military is also charged with engag-
ing in counterpropaganda. This is intended to
identify and counter an adversary’s propa-
ganda by exposing the adversary’s efforts to
influence friendly populations and their mili-
tary forces. To mitigate enemy propaganda,
the US military is supposed to provide truth-
ful information, as well as to use technology
and military operations to disrupt, degrade,
and disable the tools and techniques of an
adversary’s propaganda (Joint Publication,
2006).
The US military engages in Public Affairs
(PA) operations to increase their combat effec-
tiveness by keeping soldiers informed (FM-
46-1, 1997). PA also monitors public opinion
in environments of interest in order to detect
political, social, and cultural shifts, along with
adversary propaganda and disinformation.
 EVOLUTIONARY PSYCHOLOGY AND CYBERWARFARE
A goal is to provide ­commanders with situ-
ational awareness of the international public
information environment. Commanders are
given the means to use information to take
offensive and preemptive defensive actions
against adversaries. The US military is pro-
hibited from using PA to mislead the public,
national leaders, or the media (FM-46-1, 1997).
Memes and Narratives
Narratives may be considered a subset of
memes. A narrative is a story whether long or
short; past, present, or future; with or without
much detail; and told for any purpose.
Typically, a story has a beginning, middle, and
ending. Cultural narratives are important in
forming and maintaining cultures (or nations,
or other groups), in providing or maintaining
a sense of identity, cohesiveness, and purpose
to individuals within a culture: what is our
past, why are we here, what should we do, and
what is our future? National narratives instill
patriotism and can unify a diverse populace.
National strategic narratives describe the core
interests of a nation and provide a foundation
for statecraft and national security. For exam-
ple, the US post-World War II strategic narra-
tive provided the basis for the global struggle
against communism. There is as yet no suit­
able post-communism narrative for the global
struggle against asymmetric adversaries and
non-state actors.
More critically, there is not yet a suit-
able narrative theory or narrative engineer-
ing to counter adverse individual and social
behavior originating from narratives, and to
alter dysfunctional cultures. There is not yet
an integrated system of tools and techniques
for automating the ability to detect, identify,
and evaluate narratives and for creating and
propagating counternarratives. The narrative
threads, true and false, about the founding
of the nation and the founding fathers may
be fraying in the age of social media (the
debates in the heat of Independence Hall to
adopt the Declaration of Independence and
507
then the Constitution; George Washington
chopping down the cherry tree; Betsy Ross
sewing the first flag; the freezing revolution-
ary soldiers at Valley Forge, etc.). Likewise,
the many narratives of US history meant to
instill patriotism, such as the expansion to the
west; the pony express and wagon trains; the
epic building of the continental railroad;
the stain of slavery and the Civil War; the US
Cavalry and Indian wars; arrival of the New
York immigrant multitudes; the many wars
of the 20th century; the transformational US
inventions of the 19th and 20th centuries –
all of the narratives that have served to unify
a diverse country may need to be revitalized
in a time of increasing cultural fragmentation
and the intentional spread of memes meant to
divide the nation.
Narratives are fundamental to the human
mind and perception (Leon, 2017). Sensory
data and consequent perception are inter-
mittent, with only the individual’s illusion
and delusion of continuity and coherence.
Plagued with uncertainty and flawed informa-
tion, the subconscious mind creates an illusion
of a coherent narrative, constructing relation-
ships, motives, and intentions based on limited
or incorrect information. The mind fills in
details and revises its narratives as needed,
with or without evidence, even retroactively.
The mind autonomically, unconsciously, and
continuously constructs narratives, weaving
the weft of subjective reality among the warp
of physical reality. Thus individuals create
personal stories about who they are and what
the world is like; narratives by which they live
their lives. These individual narratives can
aggregate and evolve into cultural narratives
influencing the behavior of the family, clan,
tribe, or nation. Narratives can strongly shape
or modify a culture.
The structure, function, and ontology of
narratives are shaped by their building blocks,
including metaphors, symbols, symbolic
actions, semiotics, myths, frames, scenar-
ios, etc. For example, metaphors shape and
express cultural thought and style as embed-
ded in language, artifacts, and behavior.
508 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
They shape ideology and religion and can
spark insurgencies. The major sources of met-
aphor vary from culture to culture, whether
from the human body, animals, dwellings,
landscapes, sport, war, etc. Many cultures
have common metaphors, i.e., identical map-
ping of metaphor sources onto target domains;
but seemingly identical metaphors might have
different implications in different cultures. For
example, ‘Joan lives in a sick neighborhood’
can mean a bad neighborhood or a good (awe-
some) neighborhood, depending on the culture
of the speaker.
Memetics and the Tipping Point
Gladwell described how ideas, products,
messages, and behavior spread like viruses,
seemingly in a moment when everything
changes all at once (Gladwell, 2002). This is
the effect of memes with the most impact on
their recipients. In Gladwell’s telling, a small
number of people in a small number of situ-
ations start behaving differently, and that
behavior spreads and changes the behavior of
others in similar situations. It is contagious
behavior, which could be intentionally initi-
ated by, for example, viral marketing. There
have been such trends, for example, in fash-
ion, crime or anti-crime, music, fads, reli-
gion, and ideology. Small changes can have
big effects. A few percentage-point changes
in initial conditions can lead to non-linear
outcomes, just as a single match with limited
energy can start a forest fire. Changes can
occur quickly rather than gradually. This is
the equivalent of disease epidemics or the
effects of punctuated equilibrium in evolu-
tion (Gould, 2007).
People expect linearity and gradualness,
not change with exponential or geometric
progression (University College London,
2017). But social change can be as sud-
den as the phase change of water to ice. A
more extreme version of the heuristic 80/20
rule (Koch, 1999) is that 5% of people cause
95% of the new ideas that spread (Gladwell,
2002). The most effective of such messages
have ‘stickiness’, i.e., they stick in memory
and have impact. Simple changes in the
presentation or structuring of information
can make a big difference in their impact, as
advertisers have known for decades. Ancient
advertising slogans still stick in the memory
of those exposed to them as children decades
ago, such as: ‘Winston Tastes Good Like a
Cigarette Should’ (1954); ‘It Takes a Licking
and Keeps on Ticking’ (1956); and ‘Good to
the Last Drop’ (1926). Slogans are among the
most effective of memes.
The rules of the Tipping Point apply to
memes. According to Gladwell, the three
rules of the Tipping Point are:
1 The law of the few: few people can cause large
effects (e.g., changes in mass behavior)
2 The stickiness factor: with stickiness the message
stays in memory
3 The power of context: impact on behavior
depends on context
Starting an information epidemic, and lever-
aging the law of the few, requires concentrat-
ing resources on a few key areas. The success
of a social epidemic requires the involvement
of three types of people: connectors, mavens,
and salesmen.
• ‘Connectors’: These people are gregarious and
know many people. They are active in many
different social and professional niches and
subcultures.
• ‘Mavens’: These people are experts who are
socially motivated to share information.
• ‘Salesmen’: These people are persuaders who
are articulate. They can obtain the trust of others
and convince them of the truth and value of their
messages.
Thus mavens generate the message, the con-
nectors spread it, and the salesmen persuade
the recipients, who might be otherwise uncon-
vinced, to accept and trust the message.
In order to achieve stickiness in a meme, the
message should be made practical and seem-
ingly personal for the recipient. The message
should be as simple as possible and irresistible
 EVOLUTIONARY PSYCHOLOGY AND CYBERWARFARE
to the intended audience. The context is impor-
tant because the behavior the meme is intended
to evoke is a function of the social context of
the meme and its recipients. Context tends to be
underrated in its influence on behavior, while
character is overrated (Choi, 2003). To explain
or predict the behavior of others, people tend
to focus on individual character traits rather
than contextual circumstances. But it is groups
in a social context that have a critical role in
spreading social epidemics – and memes. To
bring about a fundamental change in people’s
beliefs and behaviors, a change that would per-
sist and serve as an example to others, a com-
munity must be created around the beliefs. For
groups to serve as incubators of messages, such
as memes, they should be kept below about
150 members. This is the maximum size for
informal, person-to-person contact and cohe-
sive group loyalty, and it is about the size of
an army infantry company, an effective corpo-
rate department, or a tribe of hunter-gatherers.
Larger groups tend to become spontaneously
divided and alienated (Dunbar, 2005).
Rumor may be the most contagious of
all social messages. According to Gladwell
(2002), rumors that have a kernel of truth,
such as an urban legend, can be distorted
three ways in order to increase their recep-
tivity by the recipients and effectiveness in
evoking the desired behavior. In terms of
memetics, the meme engineer can:
1 Level the narration or story (i.e., leave out details
essential for a recipient to understand the true
meaning of the rumor-meme or an incident that
it describes); this simplification, or even falsifica-
tion, makes the rumor-meme more relevant and
understandable for the intended recipients.
2 Sharpen the story (i.e., make the remaining details
of an incident described in the rumor-meme more
specific to the expected interests of the recipient);
this encompasses the hook for the rumor-meme.
3 Assimilate the story (i.e., change the story so it
makes sense to the recipients of the rumor-meme
who then subsequently spread the rumor-meme).
The connectors, mavens, and salesmen can
thus make an idea or rumor contagious by
509
dropping extraneous details and exaggerating
other details, so that the message acquires a
deeper meaning and is better understood by
the intended recipients. For example, the com-
plex issue of immigration can be boiled down
to: ‘murderers and rapists are pouring over the
US southern border!’ People can be induced
to transform their usual behavior or beliefs
with the right impetus. Social change can be
volatile and often inexplicable. Insufficient
knowledge of evolutionary psychology means
that human behavior, and the motivation lead-
ing to the behavior, is often a mystery. The
values and behavior of seemingly stolid and
stable people may be transformed with the
slightest impetus, such as a false rumor-meme
that is propagated to the right recipients at
the right time. False rumor-memes can cause
mobs of villagers to suddenly gather to attack
and kill their long-time neighbors, with whom
they have lived peacefully and who are inno-
cent of the acts of which they are accused,
such as kidnapping village children, killing
sacred cows, or blasphemy.
Meme Research and Engineering
A research and development program in mili-
tary memetics could develop a new approach
to countering would-be terrorists. Such a pro-
gram could prevent or mitigate irrational con-
flict and promote rational solutions to national
and international problems. It could strengthen
the US military in peacekeeping missions,
psychological operations, recruitment, and
training, as well making new discoveries con-
cerning the human brain, cognition, and social
networks.
A design for a Meme Warfare Center
(MWC) was proposed by an officer in the US
Marine Corps (Prosser, 2006). The conceptual
MWC would have two complementary subdi-
visions: the External Meme Center (EMC) and
the Internal Meme Center (IMC). The IMC
would provide the commander with advice
concerning memes needed to be engineered
and propagated to influence the culture of
510 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
the friendly combat force in the context of
circumstances such as the composition and
posture of friendly forces, command climate,
commander’s guidance, command philoso-
phy, force morale, warfare philosophy, and
rules of engagement. These memes would be
managed, analyzed, and transmitted through
common distribution mediums within the
joint force, such as annual training venues,
intranet media, email, text, and other unclassi-
fied media. To create and manage the memes,
the IMC would employ senior officer and
enlisted personnel, human-resource manag-
ers, clinical psychologists, equal-opportunity
specialists, civilian personnel and assessment
specialists. The goal of the IMC is to create
and manage internal organizational memes
similar to how corporations employ memes
to effect organizational cultural change.
The EMC would advise the commander
about enemy combat forces, noncombat-
ant indigenous personnel, and the strategic
audience. The EMC organization would
consist of a Meme Engineering Cell (MEC),
a Meme Analysis Cell (MAC), a Meme
Communications Cell (MCC), and a Strategic
Communications Cell (SCC).
The MEC would create memes, identify
and target suitable recipients, and ‘inoculate’
them with the memes. The MEC person-
nel would consist of various subject-matter
experts (SME) from the US and other coun-
tries as needed. They might include cul-
tural anthropologists, economists, linguists,
targeting specialists, technical personnel
(e.g., in communications networks), and
assessment experts. The MEC generates the
memes needed to gain a psychological and
ideological advantage in an asymmetric, non-
linear battlefield. The MEC offers the Joint
Force Commander situational context and
insight into the enemies and noncombatants
comprising the audience for the engineered
memes.
The MAC monitors and records the meme
feedback loop. The MEC would then ana-
lyze and assess the feedback provided by the
MAC. The MAC personnel would consist of
military and civilian analysts familiar with
the enemy and knowledge of tactical and
strategic maneuver warfare. They would
also have expertise in social and behavio-
ral science, systems analysis and operations
research (especially game theory), and cogni-
tive science.
The two components of meme commu-
nications, the MCC and SCC, are separate
cells with divergent purposes. The purpose of
the MCC is to transform the memes emerg-
ing from the MEC and MAC into the most
effective formats suitable for transmission
by various media suitable for the intended
recipients (e.g., whether TV, radio, newspa-
per, billboard, email, or Twitter). The MCC
would leverage available media outlets suit-
able for an intended audience in order for the
meme to have maximum impact and persis-
tence. The SCC would have personnel who
are SME familiar with local, regional, and
global media of all types. They would have
the technical expertise to compete success-
fully with the adversary’s use of media to
transmit their memes.
The concept for a Meme Control Center is
a variation on the MWC, and it could be a
civil or military-based organization. It would
be managed by a Meme Control Officer. It
would consist of a Meme Processing Center
and a Counter-Meme Processing Center.
A Memetics for Intelligence and National
Defense (MIND) Research and Development
Center would provide the tools and tech-
niques employed by the Meme and Counter-
Meme Processing Centers, where the former
gathers adversarial memes from around the
world and the latter propagates counter-
memes (which are also memes) around the
world. Both processing centers exchange
information, (e.g., feedback) to improve
the efficiency with which effective counter-
memes can be developed and propagated.
The MIND R&D Center would consist
of four directorates focused on disciplines
able to develop new tools and techniques
to improve meme effectiveness and effi-
ciency for their engineering, propagation,
 EVOLUTIONARY PSYCHOLOGY AND CYBERWARFARE
impact, and persistence. They would study
the effects of memes and counter-memes
on the human mind and behavior to deter-
mine how to best engineer and propagate
them. These directorates would consist of the
Neurological, Psychological, Sociological,
and Cybernetical Artificial Intelligence
Directorates. The neurological field (e.g.,
brain scans) would be focused primarily
on i-memes, while the psychological (e.g.,
psychological testing, game theory) is con-
cerned with both i-memes and e-memes.
The sociological (e.g., modeling and simula-
tion of social networks) and cybernetical/AI
(e.g., natural language processing) fields are
focused on e-memes.
The Neurological Directorate would per-
form meme-related R&D concerning:
• Neuroimaging: fMRI and Positron Emission
Tomography (PET)
• Pharmacological manipulation
• Electro-encephalography (EEG)
• Event-Related Potential (ERP)
• Electromyography (EMG)
• Single neuron recording
• Hormone and blood analyses
The Psychological Directorate would perform
meme-related R&D concerning:
• Psychological testing
• Behavioral testing
• Modeling Theory of Mind
• Decision-making
• Game Theory
• Uncertainty and risk
• Learning/adaptation
• Utility and valuation
• Motivation
• Emotion
• Trust and attachment
• Addictive behavior
• Belief systems
The Sociological Directorate would perform
meme-related R&D concerning:
• Modeling and simulation of social networks
• Rumor and Gossip Theory and Rumor and Gossip
Propagation Models
511
• Observation and analysis of social networks
• Social media in strategic communication
• Narrative networks and strategic narratives
• Genetic profiling
• Cultural anthropology and cultural awareness
The Cybernetical/AI Directorate would per-
form meme-related R&D concerning:
• Memetic ontology
• Natural language processing
• Text, document, data, and concept mining
• Statistical semantics
• Network theory
• Information theory and entropy
• Control theory and intelligent control systems
• Genetic, memetic, and evolutionary algorithms
• Neural networks and deep learning
• Expert systems
• Context-based reasoning
Information from a variety of sources, such
as databases, documents, social and commu-
nications networks, and human intelligence
(HUMINT) would be obtained and input at
the Meme Processing Center. Vast quantities
of information would be processed autono-
mously with machine intelligence employing
a variety of tools and techniques, such as natu-
ral language, information mining, network
analysis, and semantics, where value-driven
logic, in conjunction with a World Model and
Knowledge Database, would extract and eval-
uate detected memes. The World Model
includes entities, events, images, networks,
and psychological, cultural, and behavioral
phenomena. The Value Judgment module con-
tains algorithms to determine, for the detected
memes, priority, utility, importance, propaga-
tion, impact, and persistence.
Detected memes that are considered to be
important and to require countermeasures
are passed to the Counter-Meme Processing
Center. Creating counter-memes will be more
labor-intensive than detecting memes in the
environment, requiring a multi-disciplinary
Meme Engineering staff consisting of cul-
tural anthropologists, linguists, psycholo-
gists, semanticists, social network software
512 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
designers and programmers, operations
research analysts, economists, joint intelli-
gence and PSYOPS analysts, and joint ser-
vice personnel.
With supervised automation, the selected
extant memes are assessed with metrics in the
context of media and public opinion. Using
various tools and techniques from the four
Directorates, counter-memes are created and
passed to various Memetics Communications
Agencies (e.g., Intelligence, Joint PSYOPS,
Joint Forces Information Operations, and
Public/Civil Affairs), which then propagate
the counter-memes to the selected audiences
via viral marketing and multi-media (e.g.,
social networks, gossip and rumor, TV, radio,
newspapers, billboards, etc.) as needed. The
counter-memes become memes whose effi-
cacy (efficiency and effectiveness) in terms
of the metrics of propagation, impact, and
persistence can then be evaluated by the
Meme Processing Center and feedback of
the results provided to the Counter-Meme
Processing Center.
CONCLUSION
Cyberwarfare involves consequences of
evolved psychology that are not yet well
understood, such as the receptivity of indi-
viduals to memes that can alter their values
and behavior, and thus the values and behav-
ior of the culture or subcultures they com-
prise. In turn, memes over time may have an
effect on evolved psychology.
Memes can be defined in a way that is
useful for establishing meme metrics, a
scientific basis for memetics, and a meme
engineering process. Memes may be defined
around the concepts of information, propa-
gation, impact, and persistence. Meme and
counter-meme engineering, an essential
component of cyberwarfare, may be instanti-
ated in a Meme Control Center. To succeed
in cyberwarfare, it is critical that adversarial
memes are countered and neutralized, and
that friendly memes are received as intended.
An engineered meme or counter-meme may
be designed for optimum transmission and
reception. Processes already successful for
conventional advertising, ideological propa-
ganda, neuromarketing, neuroeconomics,
and rumor mongering can be adapted for
meme warfare, which would be enhanced by
new technological tools and techniques.
Since Dawkins’ (1976) presentation there
have emerged a coterie of proponents, skep-
tics, and opponents of the concept of the
meme. There has been no substantial effort
to test and develop the concept, or refute it.
Memetics needs a general theory, a theoreti-
cal foundation for the development of a sci-
entific discipline. The concept needs a solid
pragmatic definition that is widely accept-
able. There must be an ability to make test-
able predictions and falsifiable hypotheses,
as with any scientific discipline (Finkelstein,
2008). The discrete meme must be defined,
identified, and distinguished in the near
continuum of information, just as the dis-
crete gene can be identified (more or less)
in a string of DNA nucleotides. A quantita-
tive basis for memes must be established,
perhaps by employing such tools and tech-
niques as evolutionary psychology; physi-
ological phenomena; information theory and
entropy; genetic, memetic, and evolutionary
algorithms; modeling and simulation; and
neuroeconomics.
A coherent memetics program of sufficient
duration can establish a solid scientific and
engineering basis for the information war.
Research in evolutionary psychology may
identify why some human minds are more
receptive and reactive to some messages
over others, and how to best structure the
content and format of the messages (memes)
designed to modify the behavior of selected
populations. And most importantly, a wide-
spread effort is needed to establish a canon of
ethics to guide and constrain the development
and application of cyberwarfare memetics.
 EVOLUTIONARY PSYCHOLOGY AND CYBERWARFARE
REFERENCES
Asher, Ely (2005), Disinfect Your Mind, Galiel.
Net Publishing.
Aunger, Robert (2002), The Electric Meme: A
New Theory of How We Think. New York,
NY: The Free Press.
Blackmore, Susan (1999), The Meme Machine,
Oxford, UK: Oxford University Press.
Brodie, Richard (1996), Virus of the Mind: The
New Science of the Meme. Seattle, WA:
Integral Press.
Buss, David M. (2005), Editor, The Handbook
of Evolutionary Psychology. Hoboken, NJ:
John Wiley & Sons.
Choi, Jin Nam (2003), How Does Context Influ-
ence Individual Behavior? PsycArticles: Jour-
nal Article, https://psycnet.apa.org/
buy/2003-09567-004 (Accessed 30 August
2020)
Council on Foreign Relations (2019), Cyber Threats
and Mid-Term Elections, www.cfr.org/blog/
year-review-cyber-threats-and-mid-term-us-
elections (Accessed 30 August 2020)
Dawkins, Richard (1976), The Selfish Gene,
Oxford University Press.
de Leon Huld, Nickee (2020), How Many Words
Does the Average Person Know? https://word-
counter.io/blog/how-many-words-does-the-
average-person-know/ (Accessed 10 July 2020)
Dunbar, Robin (2005), The Human Story, Gard-
ners Books.
Finkelstein, Robert (2006), A Memetics Com-
pendium, Defense Advanced Research Pro-
jects Agency, Arlington, VA.
Finkelstein, Robert (2008), Defining Memes,
Presented at The Second Symposium on
Memetics Memory, Social Networks, and
Language: Probing the Meme Hypothesis II,
Victoria College, University of Toronto.
Finkelstein, Robert (2011), Tutorial: Military
Memetics, Presented at the Social Media for
Defense Summit, Alexandria, Virginia.
Finkelstein, Robert (2012), Neuro-Cognitive
Warfare: Accelerated Cultural Evolution
(ACE) To Prevent Terrorism and Mass Atroci-
ties, Presented at The Center for Strategic
Counterterrorism Communications, Office of
the Under Secretary for Public Diplomacy
and Public Affairs, US Department of State,
Washington, DC.
513
FM-46-1 (1997), Public Affairs Operations, HQ
Department of the Army, Washington, DC.
https://www.globalsecurity.org/military/
library/policy/army/fm/46-1/46_1.pdf
(Accessed 30 August 2020)
Giordano, James (2017), Weaponizing the Brain:
Neuroscience Advancements Spark Debate,
National Defense.
Gladwell, Malcolm (2002), The Tipping Point:
How Little Things Can Make A Big Difference,
Back Bay Books.
Gould, Stephen Jay (2007), Punctuated Equilib-
rium, Belknap Press: An Imprint of Harvard
University Press.
Hull, David L. (2000), Taking Memetics Seri-
ously: Memetics Will Be What We Make It, in
Aunger, Robert, Editor, ‘Darwinizing Culture:
The Status of Memetics as a Science’, (pp.
43–67) Oxford University Press.
Joint Publication (2003), 3-53 Joint Doctrine for
Psychological Operations, The Joint Staff,
Washington DC.
Joint Publication (2006), 3-13 Information
Operations, The Joint Staff, Washington DC.
Joint Publication (2012), 3-13.4 Military Decep-
tion, The Joint Staff, Washington DC.
Koch, Richard (1999), The 80/20 Principle,
Double Day Publishing Group.
Leon, Carlos (2017), Narrative as a Fundamen-
tal Information Unit in the Mind, A Standard
Model of Mind: AAAI Technical Report
FS-17-05.
Lynch, Aaron (1996), Thought Contagion, Basic
Books.
McLeod, Saul (2009), Short Term Memory,
Simply Psychology, www.simplypsychology.
org/short-term-memory.html (Accessed 10
July 2020)
Orwell, George (1946), Politics and the English
Language, Herbert W. Simpson Publ.
Poole, Steven (2006), Unspeak: How Words
Become Weapons, How Weapons Become A
Message, And How That Message Becomes
A Reality, Grove Press.
Prosser, Michael B. (2006), Memetics: A Growth
Industry in U.S. Military Operations, Thesis,
School of Advanced Warfighting, Marine Corps
University, US Marine Corps, Quantico, VA.
RAND Corporation (2019), Cyber-Warfare,
www.rand.org/topics/cyber-warfare.html
(Accessed 10 July 2020)
514 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY
Shannon, Claude (1948), A Mathematical
Theory of Communication, Reprinted with
corrections from The Bell System Technical
Journal, 27: 379–423, 623–656, July,
October.
Sharp, Kim and Franz, Matschinsky (2015),
Translation of Ludwig Boltzmann’s Paper
‘On the Relationship between the Second
Fundamental Theorem of the Mechanical
Theory of Heat and Probability Calculations
Regarding the Conditions for Thermal
Equilibrium’ (1909), Entropy, 17 (4),
1971–2009.
Truszkowski, Walt., Rouff, Christopher.,
Akhavannik, Mohammad and Tumstel,
Edward (2020), Robot Memetics: A Space
Exploration Perspective, Springer.
University College London (2017), Humans are
hard-wired to follow the path of least resist-
ance, Science Daily. Retrieved from https://
www.sciencedaily.com/releases/2017/02/
170221101016.htm (Accessed 15 June
2020)
US Senate (2019), Report of the Select Commit-
tee on Intelligence, United States Senate, on
Russian Active Measures Campaigns and Inter-
ference in the 2016 U.S. Election, Volume 2:
Russia’s Use of Social Media with Additional
Views, 116th Congress, Washington, DC.
 Index
Abadinsky, H. 299, 300, 302, 305, 307 Alexander, R. D. 57, 178
Abbas, M. J. 36 ALFUS 365
ABC model 4, 5, 17n1 Allcott, H. 344
Abed, R. 24–50 Allen, C. 367
Abelson, R. P. 382 Allen, F. A. 224
abuse Allen, M. W. 318
of animals 148–50 Allen, N. B. 34
health effects 128, 133 Almquist, A. J. 418, 419, 422, 425
substance 13–14, 16, 37–8 Alper, M. 278
see also drugs Alpert, G. P. 212
accuracy of information 344–5 Alschuler, A. W. 223, 224, 225
Acemoglu, D. 179, 340 altruism 173–4, 303–4
Ache tribe 272 empathy 272
adaptation 477–8 health 133
coalitional violence 321, 326 intrasexual competition 282
suicide 57–9, 61, 65 parochial 320–1
trust 340 punishment 227
see also natural selection reciprocal 173, 174, 427
adaptive lags 114–15 sperm donors 490
Adelman, M. B. 483–4 suicide 57–8, 59
adolescent-limited offenders (AL) 193 vaccination 136
Adults in the Making program 286 Alzheimer’s disease (AD) 41
adversarial allegiance 233 ambush training 210, 215
age American Psychiatric Association 25, 32,
Alzheimer’s disease 41 35, 39
criminal behavior 192–6 Amiot, C. E. 147
mental disorders 31 amygdala 116
suicides 56–7 analytical rumination hypothesis 34
age-crime curve 192, 193, 194, 195, 196, analytical therapy 6
198, 284 anchoring effects 232
age-graded theory 192–6, 280 Anderson, C. A. 374, 381–3
agency 392 Anderson, J. 336, 457–76
aggression 300–1 Anderson, M. 339
costly traits 418 Andreasen, N. 25
dangerous driving 374–97 Andrews, D. A. 279–80, 287, 288
displays 208 Andrews, P. W. 34
policing 203–20 animals
retaliation 258 abuse 148–50
Aggression Questionnaire 381 ethics 144–68
agriculture experimentation 155–6
organized crime 309–10 suicide 53
and warfare 324–5 Anna, K. 296
Aharoni, E. 221–42 anorexia nervosa (AN) 38–40
Ahmad, N. 272 anthropomorphism
Ahuvia, A. C. 483–4 artificial intelligence 342–3
Akil, H. 24 ethics 147–8, 153
Albanese, J. S. 298 anthrozoology 147
Albert, W. D. 286 antisocial behavior 190–5, 201,
Albuquerque, U. P. 107–22 277, 280
Albus, J. S. 353, 354, 355, 356 conscience 231
alcohol addiction 37–8 intrasexual competition 282–4
Alderson-Day, B. 98 Antonopoulos, G. A. 305, 307
516 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

anxiety Bandura, A. 376, 377, 379, 381, 382–3, 391–2, 401

artificial intelligence 334 Barash, D. P. 207–8
disorders 12, 16 Bargh, J. A. 114
mindfulness 94 Barkow, J. H. 265, 266, 407, 413
Apaolaza-Ibáñez, V. 109, 110, 111, 114 Barnes, J. C. 195
Apostolou, M. 40 Barnett, J. 160
Appel, M. 146 Barone, G. 339
Appleton, J. 114 Barraclough, B. 73
archetypes 6 Barraket, J. 483, 484, 486
Arluke, A. 148, 149 Barrett, H. C. 109, 111, 112, 115
Arnett Inventory of Sensation Seeking 387 Barton, S. 107, 108–9, 114, 117
Arnett, J. 387 Bashir, N. Y. 161
Arnhart, L. 248 Bastian, B. 147
Arnon, S. 440 Bates, H. 399
Arrow, K. 175 Bateson, P. 459
artificial intelligence 333–51, 427 Batson, C. D. 272
Artificial Moral Agents 367 Baumard, N. 161
artificial neural networks (ANN) 357 Baumeister, R. F. 62, 63, 67
Asendorpf, J. B. 485 Bay, B. 490
Asimov, I. 353, 367, 369 Beach, S. R. H. 286
assisted reproductive technology (ART) 481, 488–9 Beaulieu, D. A. 211
associational criminal structures 299 Beaver, K. M. 195
Atkin, C. K. 376, 377, 378, 384, 385 Beccaria, C. 223, 271
Atkinson, A. B. 339 behavior manipulation 337
Atkinson, J. A. 110 behavioral ecology 128, 297
attachment theory 34, 41, 404, 405, 406 behavioral plasticity 129, 137, 461
attacker–defender asymmetries 321–2, 323–4 Belsky, J. 30, 281–2, 285, 286, 290
attention Belzung, C. 75
dangerous animals 116 Bendor, J. 270
mindfulness 96–7 Bentham, J. 223
attention deficit hyperactivity disorder (ADHD) 31 Bercovitch, F. B. 463, 467
attractiveness 284, 342, 405, 479–80, 485, 487 Berg, J. 340
Aunger, R. 352, 500 Bergen, N. 490
Aureli, F. 244 Berger, J. 334
autistic-spectrum disorder (ASD) 31, 32, 36 Bering, J. M. 52–3, 56, 58, 59, 65
autonomic nervous system (ANS) 266 Berkowitz, L. 382
autonomous military robots 365–7, 368 Beullens, K. 376–7, 378, 385, 386
autonomous vehicles 353, 355, 360, 361–3, 365, 367, bias
368–9 implicit 209, 215
autopoiesis 361, 370 punishment 232–3, 237–8
Autor, D. 340 race 209, 337
availability heuristic 232 Biddle, S. 320, 321
Awad, E. 146 Bieling, P. J. 102
Axelrod, R. 174 The Big Bang Theory 412–13
Axial Age 94 Binder, A. 209, 210
biological sensitivity to context theory 278, 281–2, 291
Badcock, C. 36 biophilia 113–14, 115, 116–17, 148
Badcock, P. B. 34 bipolar disorder (BPD) 42
Baden, J. 262–3 Birch, L. L. 160
Baechler, J. 64 Bird Box 380
Bagehot, W. 174–5 Bird, D. W. 317
Bailey, D. H. 262 birds, death response 466–7
Bakermans-Kranenburg, M. J. 277, 281, 285, 286–7, Biro, D. 457, 463, 465
290 Biswas-Diener, R. 98
Balinski, M. L. 180 Bittman, M. 157
Bancroft, J. 61 Bittner, E. 203, 206, 214, 216
Bandes, S. 268–9 Black Lives Matter 205
 INDEX 517

Blackmore, S. 500 Bribiescas, R. G. 56

Blair, C. 99 Brody, G. H. 286
Blanchard, D. C. 71 Broesch, J. 111, 115, 116
Blome, M. W. 108 Brooke, P. 152
Bloom, P. 148 Brosnan, S. F. 12
Blumenstein, L. 211 Brosschot, J. F. 97–8
Boaz, N. T. 418, 419, 422, 425 Brower, D. 79n4
Bockman, J. 151, 156 Brown, C. 480
Boden, T. 436, 439 Brown, D. 53, 225
body language 407 Brown, G. R. 115, 247
Boehm, C. 271, 300 Brown, G. W. 75
Boesch, C. 470 Brown, M. J. F. 461
Bohm, R. 260 Brown, R. M. 58, 62, 253
Bokek-Cohen, Y. 485 Brown, S. L. 253
Bolen, J. 267 Brune, M. 24, 27, 28, 29, 30, 33, 34, 35, 41
Bolhuis,J. J. 110 Brunero, J. 262
Bolig, R. 484 Bruni, F. 411
Bollard, L. 144, 156 Bryant, C. 160
Boltzmann, L. 503 Bryson, J. 333–51
Bonta, J. 279–80, 287, 288 Buckels, E. 67
Boot, M. 324, 325 Buckholz, J. 259, 270
Booth, D. A. 156 Buckner, W. 321–2
borderline personality disorder 11 Buddhism 94–7, 98, 102
Borone, E. J. 25 Budge, S. L. 16
Bossema, E. R. 491 Bugental, D. B. 211
Bostrom, N. 146 Bukowski, W. M. 211
Botanov, Y. 8 bulimia nervosa (BN) 38–40
Bourgeois, C. A. 301 Bullitt 376
Bourgeois, P. 303 Buonanno, P. 260
Bowlby, J. 6, 34, 41, 110, 209 Burger, O. 436
Bowles, S. 321 Burger, W. 260
Boyce, W. T. 281 Burgess, J. W. 175
Boyd, D. 334, 335 Burkhardt, J. M. 343
Boyd, R. 304 Burns, J. 35
Boyer, P. 177, 318, 480 Bushman, B. J. 374, 381–3
brachiation 108–9 Bushway, S. D. 249
brachycephalic dogs 154 Buss, A. H. 381
Bradford, E. E. 336 Buss, D. M. 15, 66, 77, 180, 199–201, 204, 235, 250,
Bradshaw, J. W. 147 282, 283, 291, 375, 378, 382, 403, 404, 413,
brain 368 479–80, 485, 488, 491–2
emotions 266–7 Buston, P. M. 479, 488
group size 262
mass media 407, 408, 411, 413 cadaverine 468
memes 499–500 Cahan, S. 461
neocortex size 262 Camilleri, J. A. 283
organoids 370 Campbell, A. 289
rationality & emotion 267 Campbell, T. S. 100
romantic love 410 cannibalism 422, 460, 463–4, 468, 470
size 262 Cannon, W. B. 175
theory of mind 353–5 Canton, R. 265, 269
trolley problem 368 Caplan, B. 345
see also cognitive abilities carbon emissions 435–56
Braithwaite, J. 272–3 carbon legacies 436, 438–9, 449
Braithwaite, V. 145, 146 caregiver distress 253
Brand, P. W. 63, 65 Carmi, N. 440
Brazier, Y. 366 Carmody, J. 94–106
Brézillon, P. 357 Carneiro, R. L. 309
518 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

carnism 147, 149 Chu, C. 58

Carr, W. J. 467 Churchill, W. 159
cars civil war 325
autonomous 353, 355, 360, 361–3, 365, 367, civil-rights movements 150
368–9 Cladder-Micus, M. 102
commercials 384–5 Clark, A. G. 418, 419–20, 423
dangerous driving 374–97 Clark, B. C. 307
racing 385, 387–8, 389 Clark, M. 159
speeding 388–92 classical conditioning 209
Carstensen, L. L. 379 clean meat 159–60
Carter, G. 75 Clear, T. 251
Carter, S. 209 Cleveland, H. H. 286
Cartwright, J. 422, 425, 427 cliff edge fitness functions model 35–6
Carvalho, S. 457, 465, 466, 470 climate change 435–56
case conceptualization 4–5 clinical heterogeneity 30
Cassiers, T. 339 cloning 155, 173
Caudell, M. A. 435–56 co-morbidity 32
causal heterogeneity 30 coalitional aggression, warfare 316–30
causality 26–9, 44 coalitions, political institutions 180
Cavanagh, J. T. O. 73 Cochran, J. C. 278, 280
Cave, S. 334 cognitive abilities
Caviola, L. 146, 147, 155 emotion effect 363
celebrities warfare 317
mass media 404, 411–13, 414 see also brain
romantic love 410 Cognitive Behavioral Therapy (CBT) 4, 12, 13, 290
tabloid media 405 cognitive competence 64–5
Werther effect 379 Cognitive Evolutionary Therapy for Depression
Cerel, J. 51 (CETD) 8–10, 16
Chagnon, F. 73 Cognitive Evolutionary Therapy for Personality Disor-
Chagnon, N. A. 271 ders (CET-PD) 11
Chamberlain, J. M. 12 cognitive niche 426
Chan, D. 99 cognitive psychology 131, 353, 427
Chapman, C. A. 317 cognitive social learning 280
Chapman, L. J. 317 Cognitive Therapy (CT) 10
Chapple, A. 55 Cohen, J. 406, 435, 439, 450
Chavan, S. 336 Cohen, P. 445
cheating 136–7, 173, 178, 207, 264, 304 Cohen, R. 153
mate choice 480 Coleman, J. L. 222
punishment 227–9, 230–1 Coleman, L. 377, 379
reciprocity 258 collective action problem 320, 323
tit-for-tat 272 collective efficacy 196
Cheng, J. T. 211 collectivization 176–7
chess playing 375 Colley, K. 115
Chiao, J. Y. 180 Colomer, J. M. 179
Chicago school of criminology 196 commitment in relationships 14–15
children/adolescents common mental disorder (CMD), suicide 73, 75–6
animal abuse 149–50 communication
celebrity attachments 404 fluency 425–6, 429–30
death response 469 mass media 398–416
delinquent behavior 273, 285–6 oral speech 417–34
morality 261 strategic 401
parent incarceration 251 suicide as 60–2
punishment 378–9 community supervision 278, 285, 287, 289
Chiurliza, B. 79 Community-orienteered policing (COP) 215
Choe, D. H. 460 compensatory adaptation 418, 428–31
Choi, J.-K. 321 complexity theory 401
Cholbi, M. 59 Component Meme Concept (CMC) 504
 INDEX 519

computational theory of mind 353–5 incarceration 243–57

Condorcet, Marquis de 172, 178 organized 296–315
Confer, J. C. 300 policing 203–20
Connolly, K. 227, 234 recidivism 277–81, 287, 289–91
conscience 230–1, 266 sentencing 221–42
consciousness, robots 364–5, 367 crime-reduction effect 249
consequentialism, animal ethics 145 Crisis Intervention Team (CIT) 214
control behavior 270 Cronk, L. 303–4
Cook, M. 116 Cross, D. J. 467
Coolidge effect 15 Crow, J. F. 31
Cooney, N. 151, 157, 159 Crow, T. J. 35
Coontz, S. 482 Crowley, T. J. 439
cooperation 222, 261–2 cuckoldry 283, 290
artificial intelligence 335–6 Cullen, F. T. 277, 278, 287, 288, 290
e-collaboration 417–18, 428–31 Culotta, E. 79
health 133 cultural engineering 499
ingroups/outgroups 210, 215 cultural evolution 304, 499
organized crime 297, 303–8, 310–11 Currie, E. 260
political institutions 171–4, 176, 178 Cushman, F. 229, 230, 236
retaliation 226 cuteness, animal ethics 152–4, 155, 162
vaccination 136 cyberwarfare 499–514
voter suppression 180
warfare 317–18 Daft, R. L. 407
coordination, artificial intelligence 335–6 Dahl, R. 175
coping mechanisms 12 Dahlen, E. R. 383
copycat effect 379–81 Dahlgreen, W. 160
Corning, P. A. 176 Daly, M. 194, 195, 196–9, 227, 234, 235, 284, 288–90,
Cornwell, R. E. 79 323, 375, 376, 378–9
cortical dysconnectivity hypothesis 35 Damasio, A. R. 97, 363
Cosmides, L. 56, 59, 65, 66, 70, 71, 78, 110, 114, 116, danger, impulsive behavior 130
244, 245, 259, 272, 303, 317 dangerous animals, response to 116
Coss, R. G. 108 Daniels, K. R. 489, 491
cost–benefit 12 Dart, T. 214
analysis 173, 208 Darwin, C. 52, 53, 172, 189, 221, 266, 352–3, 370,
outcomes 4, 7 409, 420
punishment 233–4 Das, S. 462, 464, 465, 466
trust 341–2 David, K. 406
warfare 317, 318–19, 321, 323 Davidson, C. 181
costless traits 419–20, 422–4, 426 Davies, N. 208
costly traits 418–28 Davies, P. C. W. 177
Coulthard, T. J. 109 Davis, L. 358
counseling 3–23 Davis, M. 223
counter-exploitation strategies 243, 244–5, 253 Davis, S. 226
counteractive niche construction 247–8 Dawkins, R. 56, 58, 177, 207, 304, 499, 500–2,
counterpropaganda 506 505, 512
couples therapy 14–16, 17 De Backer, C. J. 405
Cowhig, M. 277, 278 De Dreu, C. K. W. 321
Cox, G. W. 180 de Leon Huld, N. 503
Craig, M. 267 De Marco, A. 463, 464
Craig, T. 115 de Villiers, B. 30
Crank, J. P. 212 de Visser, E. J. 342
Craparo, G. 303 de Waal, F. 244, 264–5, 270
Creanza, N. 179 de-escalation 213–14, 216
Crespi, B. J. 31, 32, 36 death, responses to 457–76
crime 188–202 death penalty 259, 261, 268–9, 271
copycat 380 DeCatanzaro, D. 56, 58, 60, 62, 65
corrections and rehabilitation 277–95 decentering 102
520 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

deep learning 357 Dudley, R. 38

deep neural networks (DNN) 357–8 Dunbar, R. I. 262, 480, 481, 484
deep time 189 Dunbar’s Number 262
Defense Advanced Research Projects Agency Duntley, J. D. 54, 55
(DARPA) 506 Durant, A. 271
DeGregori, J. 44 Durant, W. 271
Deikman, A. J. 101 Durisko, Z. 30
Del Giudice, M. 25, 26, 27–9, 31, 32, 33, 36, 41–2 Durkheim, E. 53, 59, 63, 271
Delacroix, C. 490 Durose, M. R. 278
delay discounting 131 Durrant, R. 243, 246–8, 296–315
DeLoache, J. S. 116 Dworkin, R. 222
Delton, A. W. 227–8 dysfunctional thinking 4, 9, 10, 12
DeLuca, D. C. 428, 429 dysmorphic disorder 12–13
democracy 177–81
Demographic Transition Theory 117 e-collaboration 417–18, 428–31
demographics, climate change 435 Eagle, N. 335
Denzler, M. 229, 234 Easton, D. 175
depression 6–10, 16, 24, 32–4 Eastwick, P. W. 486
antisuicide 76 eating disorders 13, 16, 31, 38–40
HTTLPR gene 30 Eaves, L. J. 30
meta-awareness 102 Eccleston, C. 70
parental age 31 Eckel, C. C. 342
Desai, R. A. 386 Edgar, B. 66
deterrence 223–9, 231, 233–7, 249, 260, 263 education, carbon emissions 436, 438–42, 444–6,
Deuze, M. 399 448–51
Devlin, P. 264 Efklides, A. 67
diametrical model of psychosis 36 Eibl-Eibesfeldt, I. 204
Diamond, J. 148, 153 Eisenberger, N. I. 64, 70
dictatorship 176, 178 electoral systems 177–81
Dietz, T. 435, 439, 444 Elish, M. C. 334, 335
Diez, L. 461 Ellis, A. 4, 5, 179
differential susceptibility 30, 278, 281–2, 285–7, 290–1 Ellis, B. J. 281
diffuse tensor imaging (DTI) 267 Elwood, R. W. 145, 146
direct reciprocity 226, 228, 304, 305–6, 308 Emlen, S. T. 479, 488
discriminate sociality 244 emotion
disease, memory 113 death response 457–9, 462, 464–5, 469
disengagement 34, 195 decision-making 266–7
moral 147 decoding accuracy 250–1
tactical 213 mass media 409
disgust robots 363–4
animal ethics 152, 155–6, 157, 160, 162 emotional exhaustion 252
distastefulness in insects 173 emotional intelligence (EI) 250–1
disinformation 343–4 emotional pathways 37
distastefulness in insects 173 empathy 265–6, 271–2
Dobzhansky, T. 189 animal ethics 147–8, 149, 152, 155–6, 157
Dodge, K. A. 250 mass media 408
Dolan, K. 180 employment, artificial intelligence 334, 339–40
dopamine 253, 259, 262 Engel, C. 70
Dos Santos, M. 270 Engh, A. L. 464
DRD4 gene 286 Engle, Y. 404
driving behavior, dangerous 374–97 Enke, R. 379–80
drugs entertainment
addiction 37–8 animal ethics 148–9
organized crime 296, 298, 299, 301, 302–3, mass media 408–9, 413
305–7, 311 Entertainment Software Association (ESA) 386
DSM 10, 25–6, 31–2, 34–5, 38–9, 41 entrepreneurial criminal structures 299
dual taxonomic theory 192–4, 195 entropy 502–3
 INDEX 521

Environment of Evolutionary Adaptedness (EEA) fender defenses 66–7, 71

110–11, 209 Ferguson, C. J. 40
environmental sciences 107–22 Fernyhough, C. 98
Epperly, B. 180 Ferreira, W. S. Jr 107–22
Eren, O. 233 Ferrill, A. 324
Erikson, E. 404, 409 fertility 40, 179, 487
Ernst, E. 491 age 56, 132
erotomania 411–12 bipolar disorder 42
escape, suicide as 62, 63–5 carbon emissions 435–6, 437–42, 444–51
Escobar, P. 299, 305 health 123
essentialism 401 schizophrenia 35, 36
ethics in vitro fertilization 479, 481, 488–92
animal 144–68 wealth 117, 173
autonomous military robots 366, 368 see also reproduction
behavior manipulation 337 Fessler, D. M. T. 156, 157, 469
robots 367–9 feuds 271
suicide prevention 76 Feygin, D. L. 7, 12
see also morality ‘fictive kin’ 305, 308
eugenics 173, 176 Fiddes, N. 156
European Convention on Human Rights 243 Fiddick, L. 226
Evolutionarily Stable Strategy (ESS) 173–4, 178, 207 ‘fight or flight’ 209–10, 363, 382
evolutionary by-product models 35 films
evolutionary democracy hypothesis 179 copycat effect 380–1
Evolutionary Fitness Scale (EFS) 9, 10 dangerous driving 376–7, 384, 385, 387, 388–92
exercise 132 empathy 408
explore/ exploit dilemma 176, 177 The Fast and Furious 376, 377, 380–1, 385,
External Meme Center (EMC) 509–10 388–92
external memes (e-memes) 501–2, 511 Fink, B. 284
externalizing behaviors 286 Finkel, E. J. 482, 484, 485, 486
extrinsic risk 437–8, 439, 440, 450–1 Finkelstein, R. 352–73, 499–514
eye movement desensitization and reprocessing First, M. B. 16, 25, 75
(EMDR) 7 Fischbacher, U. 227
Fischer, P. 376, 386, 387
Facebook 345, 403 Fisher, H. 410
Fachner, G. 209 Fisher, M. L. 301, 405
facial emotion decoding accuracy 250–1 Fisher, R. A. 173, 336
factory farming 150, 151, 157, 159 Fiske, S. T. 251
Fagan, J. 215, 216 Fisman, R. 485
Fair, C. C. 323 fitness
fake news 343–5 artificial intelligence 334
Falger, E. L. F. 58 costly traits 418
Falger, V. S. E. 58 crime 244, 250–1, 254
Fanèovièová, J. 116 drug use 38
Fanjul-Moles, M. L. 460, 461 life history theory 436–9
fantasies 404, 409–10, 411–13 new traits 419–20
Farb, N. A. S. 100 primary goods 248
Farberow, N. L. 60 prosocial behavior 253
Farrow, T. 266 public health 123
The Fast and Furious 376, 377, 380–1, 385, 388–92 punishment 226, 227, 228, 231–2
fast–slow life history 28–9, 31, 32, 33, 36–7, 38, 40–2 reproduction 477–8
Fast Track Randomized Controlled Trial 286 suicide 52, 54–6, 61–3
fear, threat perception failure 209–10 warfare 318
Fehr, E. 227, 308 see also inclusive fitness
Feld, E. 171–87 Fitzgerald, A. J. 150
Feldman, M. W. 179 fleeing famine hypothesis 39
Felson, R. B. 211 Fleischman, D. S. 144–68
feminism 180, 181 Flynn, C. P. 149
522 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

Foley, R. 107, 110 gender 191–2

Folkes, V. S. 483 Aggression Questionnaire 381
food animal ethics 152, 154–5
degrees of suffering 151–2, 156–8 attractiveness 284
taboos 156 clean meat 160
vegetarians 151, 155, 157, 158–61 couples therapy 14–15
forgiveness 272–3 crime 190–2, 199–200
Foster, T. L. 469 dangerous driving 374, 375–6, 378–9, 383,
Fox, M. 41 386–7, 390
Fox, R. L. 179 depression 33–4
Fradella, H. F. 203 drug use 38
Francione, G. L. 145, 155 eating disorders 39
Francis, R. C. 205 electoral systems 180
Frank, H. 153 fandom 412–13
Frank, M. G. 153 incarceration 251
Frank, R. H 228, 230 intrasexual competition 282–3, 288–90
Frank, S. A. 178, 179, 180 mass media 403–5
Franklin, J. C. 58, 75 organized crime 301
Frazzetto, G. 486 police culture 210–11
free will, robots 364–5 schizophrenia 35
Freeman, T. 491 sensation seeking 382
freerider problem 176 two-fold cost of males 478
Freud, S. 4, 5–6, 52, 79, 222 warfare 323
Frey, R. G. 145 young male syndrome 194, 195, 199, 374, 375, 383
Fridlund, A. J. 228, 229, 233 see also mate selection; parental investment
Fried, E. I. 33 General Aggression Model (GAM) 374, 381–4, 388,
Friesem, D. E. 109 391, 392
Frischmann, B. M. 176 genetic algorithms (GAs) 358–9
Frost, P. 110 genetic drift 423
FSD model 32 genetic noise 57
Fujisawa, K. K. 245 genetic plasticity 282, 285–6
Fuller, L. 222 genetic programming 358–60
funerary archeology 458 genetics 44
Funk, C. 155 mental disorders 29–31
Furman vs. Georgia 261 see also heritability
Füstös, J. 99 genomic imprinting 31
future discounting 38, 132, 197 Gentzkow, M. 344
Fyfe, J. J. 210, 213 Gest, T. 203
Ghirlanda, S. 153, 154
Gabriel, R. A. 325 Gil-White, F. J. 211
Gächter, S. 227 Gilbert, D. T. 97
Gaes, G. 259–60 Gilbert, P. 12, 13
Gale, J. 4 Giles, D. C. 406
Gallup, G. G. 58 Gillespie, J. H. 418, 419, 421, 422, 423
game theory 173–4, 175, 206–9, 227, 264, 303 Gillmor, C. S. 486
forgiveness 272 Gintis, H. 227
hawk-dove game 206–7, 216 Giordano, J. 505
see also Prisoner’s Dilemma (PD) Giosan, C. 3–23
Gandhi, S. G. 55 Gladwell, M. 508–9
Garcia, J. 147 Glantz, K. 14, 15–16
Gardner, J. 222 Glover, E. J. 424
gastroesophageal reflux 424–5 Glowacki, L. 300, 318, 319, 321–2
Gat, A. 318, 319, 321–2, 325 Gluckman, P. D. 26, 28
Gavrilets, S. 261 goal-directed behavior 130
Geary, D. 66, 96, 210, 211, 262 goals
Gebusi tribe 205 depression 7
Geher, G. 400 psychotherapy 4
 INDEX 523

Godfrey-Smith, P. 335 Hammack, P. L. 334

Godin, J. 226 handicap principle 419
Goethe, J. W. von 379 Hanson, M. A. 28
Goldberg, D. 75, 76, 358, 359–60 Hardin, G. 118, 176, 177, 207
Goldberg, O. 377, 380 Harding, D. J. 249, 251
Goldstein, J. S. 323 harm-avoidance model 44
Gollwitzer, M. 229, 234 Harman, G. H. 77
Gonçalves, A. 457, 465, 466, 470 harmful dysfunction (HD) 25
Gone in 60 Seconds 380 Harper, D. G. C. 418, 419
Goode, E. 484, 485 Harris, E. C. 73
Goodenough, O. 269 Harris, M. 59
Goodfellow, I. 353, 357 Harris, N. 245
Goodwin, S. A. 251 Harris, T. O. 75, 76
Goodyer, I.73 75, 76 Hart, H. L. A. 222
Gorer, G. 469 Hartl, D. L. 418, 419, 423
Gottfredson, M. 261 Hartmann, P. 109, 110, 111, 114
Gould, S. J. 78, 362 Harvey, A. G. 75
governance crimes 298, 299 Hasen, R. 181
Grabe, M. E. 398, 400, 401, 403, 405, 407, 408, 412 Hatch, V. 268
Grabo, A. 180 hawk-dove game 206–7, 216
Gradual Exposure Therapy 13 Hawthorne, N. 258
Graetz, K. A. 417, 425–6, 428–9 Hayes, S. C. 100
Grammer, K. 481 Hazan, C. 406
Grand Theft Auto V 386 Heerwagen, J. H. 114
Grandjean, D. 250 Hegel, G. 223
Green, C. 151 Heintzelman, S. J. 67
Greenberg, B. S. 376, 384 Helfgott, J. B. 380
Greene, J. D. 145, 368 ‘helper’s high’ 253
Griebel, G. 71 Hendin, H. 76
Grigg, N. P. 466 Hennighausen, 407, 408, 409
Grofman, B. 180 Henrich, J. 178, 211, 270
Gross, J. 321 Henry, A. D. 117
group selection 36, 59, 304 Henry Ford Museum 399
group size Henry-Waring, M. S. 483, 484, 486
brain size relationship 262 herd immunity 136, 137
Hutterites 263 heritability 29–31, 204
groups eating disorders 39
tribalism 338 obesity 124–5
warfare 316–30 schizophrenia 35
see also ingroups/outgroups suicide 52
guilt 230–1, 245 Herrman, A. 368
Guisinger, S. 39 Herzog, H. 146, 147, 149, 152, 153, 155, 156, 157
Gullone, E. 150 Herzog, K. 154
Gunn, J. F. III 51–93 heterogeneity 30, 32
Gunnell, J. G. 175 hierarchy of needs 403
Gustavsson, L. 485 hijack hypothesis 37
Guttmacher, A. F. 489 Hill, A. 323
Hill, K. R. 317
Haaga, D. A. F. 12 Hill, P. 299, 306, 307
Hagen, E. H. 34, 60 Hill, R. A. 262
Haidt, J. 152, 153, 154, 160, 265, 267 Hirschfeld, K. 297, 310
Haig, D. 31 Hirschi, T. 261
Haldane, J. B. S. 173, 227 Hirsh-Pasek, K. 153
Hall, L. 363 Hitsch, G. J. 480, 487
Hamilton, A. 172 Hoaken, P. N. 250
Hamilton, W. D. 57, 58, 173, 174, 227, 250, 304 Hobbes, T. 174, 205, 222
Hamilton’s Rule 227 Hodal, K. 159
524 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

Hodson, G. 161 inequality, artificial intelligence 338–40

Hoffman, F. 325 inequity aversion 270
Hoffman, M. B. 221–42 infidelity 14–15, 17
Hofstede, G. 441–2, 444, 450 Information–Propagation–Impact–Persistence (IPIP)
Hogg, M. A. 336 502, 504
Holmes, O. W. 222 ingroups/outgroups 210, 215, 300–1
Homberg, J. R. 30 group selection 304
Horowitz, A. V. 32 outgroup intolerance hypothesis 36
Horrobin, D. F. 35 punishment 233
HTTLPR gene 30 warfare 318, 321
Hu, S. 115 Insel, T. 26
Huber, F. 68 Instagram 403
Hublin, J. J. 108 Institute of Electrical and Electronics Engineers
Huffziger, S. 100 (IEEE) 337
Hull, D. L. 352, 500 Institutional Analysis and Development (IAD)
human–animal relations (anthrozoology ) 147 framework 177
human evolution 107–9, 209, 262, 435 institutions
communication 407 organized crime 308–10
death 457, 468 political 171–87
environments 107–18 instrumental goods 248
mass media 411 Integrated Motivational–Volitional Model of Suicidal
memes 499 Behavior (IMV) 63, 78
oral speech 424–5 intelligence of robots 355–6
theory of mind 353–4 inter-individual exploitation 243, 244
warfare 318–19 Internal Meme Center (IMC) 509–10
human rights 135, 243, 338, 366 internal memes (i-memes) 501–2, 511
human trafficking 298, 301 Interpersonal–Psychological Theory of Suicide (IPTS)
Hume, D. 171–2, 235 62–4, 78
Hume’s Gap see naturalistic fallacy intragenomic conflict 31
Humphrey, J. 213, 216 intrasexual competition 278, 282–4, 287–91, 378
Humphrey, N. 59, 64–6, 67, 79n5 Izquierdo, C. 468
Hunter, J. 489
Hurford, P. 158 Jablensky, A. 75
Hutterites 262–3 Jackson, J. C. 304
Huxley, J. 36 Jackson, S. 252
Huxley, T. H. 172 Jacobson, J. D. 8
Jacoby, S. 271
ICD-10 26, 31–2, 38–9, 41 Jain monks 149
identifiable victim effect 232 James, L. 203–20
Iglesias, T. L. 466 James, S. 212
Imel, Z. E. 16, 75 James, W. 100
imitation 376–81, 385 jealousy 15, 17
copycat effect 379–81 Jefferson, T. 172, 205
mass media 409 Jeong, J. 379
implicit bias 209, 215 Jervis, R. 321
impulsive behavior 130, 132 Jiang, J. 339
in vitro fertilization (IVF) 479, 481, 488–92 Jin, S. V. 404
incapacitation 223, 224–5, 227, 228, 233, 234, 236, Johnson, E. I. 251
244, 249, 260 Johnson, R. R. 210
incarceration see prison Joiner, T. E. 62, 69, 79
inceptive niche construction 247 Jonason, P. 405
inclusive fitness 7, 9, 33, 173, 245, 297 Jones, A. 344
organized crime 297, 310 Jones, J. M. 203
prisons 250, 251 Jones, O. D. 237
suicide 57–60, 62, 63 Jones, T. L. 318
indirect reciprocity 228, 304, 305–6, 308 Jonson, C. L. 278
industrial revolution 114, 126, 206 Jorgensen, C. 258–76
 INDEX 525

Joy, M. 147 Knauft, B. M. 205

Julian, G. E. 461 knowledge communication 426–8, 430
Jung, C. G. 6 Koburger, N. 380
jurisprudence 221–42 Kock, N. 116, 406–7, 413, 417–34
Koivu, K. L. 309, 310
Kabat-Zinn, J. 96, 99, 100 Konopacki, M. 344
Kaeble, D. 277, 278 Kostermans, E. 387
Kahane, G. 145 Kotrschal, K. 147
Kahneman, D. 232 Koza, J. R. 358
Kamhawi, R. 403 Koziel, S. 483
Kaminski, J. 153 Krasnow, M. M. 227–8
Kanazawa, S. 195, 204 Krebs, J. 208
Kant, I. 223, 263–4, 269 Kriegman, D. 5
Kaplan, H. 436, 439 Kristiansen, J. E. 42
Kappeler, P. M. 53 Kropotkin, P. 172–3
Kappeler, V. E. 211 Krupnik, V. 7–8
Karwoski, L. 8 Kuhl, J. 67
Kasperbauer, T. J. 148 Kuhn, T. 43
Kassab, H. S. 301, 306, 307 Kurman, M. 357–8
Kasser, T. 404 Kurzban, R. 161, 244, 304
Katchadourian, H. 230
Kavan, M. G. 25 Lahr, M. 319
Kearney, J. 156 Laidre, M. E. 462
Keeley, L. H. 319, 324 Laland, K. N. 115, 177, 247, 459
keeper defenses 66–7, 69–75, 77–8 Lam, D. 4
Kendell, R. 75 Lamarckianism 352–3, 370, 399, 500
Kendler, K. S. 24, 30 Lambert, E. G. 252
Kennedy, N. 411 landscape, preferences 107, 108–11, 113–15, 117
Kennedy, P. 55 Lane, J. C. 180
Kenrick, D. T. 302 Lange, A. 307
Kessler, R. C. 75 language, coalitional aggression 317, 318
Kesteloot, C. 339 Lankford, A. 56, 57, 58, 59
Khadjavi, M. 307 Lantz, B. 301
Khot, T. S. 336 Lao, S. 99
Kiehl, K. A. 231, 233 lateral prefrontal cortex (LPFC) 266
Killingsworth, M. A. 97 Latimer, J. 273
Kime, P. P. 334, 335, 342 Latner, M. 171–87
kin Laub, J. H. 192, 280
fictive 305, 308 Lauder, G. V. 77
investment 7 Lavoie, K. 491
organized crime 308 law
suicide effects 55–6, 61 behavioral rules 367
see also kin selection early legal codes 261
kin selection 173, 174, 304 jurisprudence 221–42
animal ethics 152–3 policing 203–20
punishment 227 robots 367, 369
suicide 57–8, 59 Lawless, J. L. 179
King, J. 260 Lawrence, C. K. F. 159
King, L. A. 67 Layard, R. 67
King, R. 203, 208 Lazar, S. W. 100
Kinney, D. K. 35, 58 Lazer, D. M. 343
Kirby, S. 301 leader–follower dynamics 322–3
Kirillova, G. P. 13 Leary, M. R. 62, 63, 244
Kleemans, E. 300 LeDoux, J. 363
Klein, R. 66, 259, 264 Lee, K. M. 406
Klinger, D. A. 216 Lee, S. A. 334
Klonsky, E. D. 73 Leech, B. L. 303–4
526 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

legal positivism 222 McCullough, M. 236, 304

legal realism 222–3 McCutcheon, L. E. 414
Leibbrandt, A. 270 Macdonald, C. J.-H. 68
Leigh, E. G. 178 MacDonald, G. 63
Leiter, B. 222, 223 McDonald, M. M. 300–1
Lengel, R. H. 407 McGaffin, B. J. 14
Lenton, A. P. 483 McGann, A. 178, 179
Leon, C. 507 McGirr, A. 79n3
Lesch, K. P. 30 McGrath, J. J. 35
Lester, D. 56, 58, 62, 63, 78, 79 McGuire, M. 4, 7, 9
Letterman, D. 411 Machado, C. 344
Levi, M. 297–8 Machan, T. R. 145
Levinson, D. B. 100 Machine learning (ML) 335, 358
Levy, J. S. 325 MacInnis, C. C. 161
Lew-Levy, S. 148 McKay, H. D. 196
Lewis, D. M. G. 470 McKibbin, W. F. 273
Lewontin, R. C. 78 Mackie, J. 263
Liang, Y. 334 McLaren, M. E. 159
Lieberman, M. D. 64, 70 McLaughlin, C. 481
Lieberman, P. 424, 425 McLeod, S. 503
life expectancy 117 McNally, R. J. 66
climate change 437, 440–1 McNiel, D. E. 250
crime 196–8 Macovei, M. 243
life history theory (LHT) 28–9, 32, 36–7, 38, 40, 41, McPherson, M. 344
42, 130, 131–2, 435, 436–51 McQueen, R. J. 428
life-course persistent offenders (LCP) 193 Madison, J. 171–2
limbic system 145, 266–7 Madrigal, A. C. 159
override 265, 266 Mafia 298, 300, 301, 302, 305–10
Lin, P. 369 Maier, K. H. 306, 307
Lindemann, B. 156 mail-order brides 485
Linehan, M. M. 60 major depressive disorder (MDD) 25, 32, 34
linoleic acid 460 Malamuth, N. M. 180, 401, 404
lipid metabolism hypothesis 35 male warrior hypothesis 300–1, 304, 310
Lipsey, M. W. 279, 288 Malo, Pablo 51–93
Lipson, H. 357–8 Malthus, T. 172
Little, A. C. 481 March, J. G. 176
LoBue, V. 116 market-based crimes 298
Locke, J. 205 Marks, I. F. M. 12
Loerinc, A. G. 12 Marois, R. 259, 270
Logan, C. 259–60 Martín-López, B. 116–17
Lopez, A. C. 316–30 Martlew, M. 227
López-García, L. M. 108 Maruna, S. 247, 248
Lopez-Perez, R. 270 Marx, K. 222
López-Riquelme, G. O. 460, 461 Masci, D. 155
Lorenz, K. 204 Masi, S. 465
loss of psychomotor energy (LoPE) 77, 78 Maslach, C. 252
Luks, A. 253 Maslow, A. 403
Lusk, J. 157–8 Mason, J. 157
Lutz, J. 100 mass media 398–416
Lynch, T. 180 Massey, D. 265, 267
Lynn, M. 484 Mastrofski, S. D. 215
mate guarding 15, 194
McAndrew, F. T. 484, 485 mate poaching 200
McBride, K. 264 mate selection 478–92
McCartney, K. 401–2 mass media 403, 411–12
McCarty, N. M. 336, 339 online dating 480–1
McCoy, D. R. 172 risky behavior 375, 382
McCulloch, W. 353 Matland, R. E. 180
 INDEX 527

Matschinsky, F. 503 misinformation 343–4

Matsuzawa, T. 462, 465 mismatch 3, 6, 11, 27–9, 36, 125, 128, 137,
Mauer, M. 243, 251 232, 236
Mayer-Schönberger, V. 338 Alzheimer’s disease 41
Maynard Smith, J. 57, 173–4, 206–7, 208, 418, 419, depression 33
423, 477, 478 EFS 9
Mead, G. 401 organized crime 300
Mealey, L. 39, 41 SCH 40
Mears, D. P. 278, 280 SSDs 36
measles, antivaccination movement 136 substance abuse 13, 37, 38
meat consumption 147, 148, 151, 152, 156–61 suicide 58
media effects theory 376–7 Mitchell, D. 344
media equation 408, 411–12, 413 Mocan, N. 233
media naturalness 406–7, 413 Mocetti, S. 339
media richness 406–7, 413 modelling, risky behavior 374, 376–7, 380, 382–93
meditation 94–106 Moehl, M.-B. 14, 15–16
Mehling, W. E. 99 Moffitt, T. E. 192–3, 202n3
Meme Analysis Cell (MAC) 510 Mohorcich, J. 160
Meme Communications Cell (MCC) 510 Moir, R. D. 41
Meme Engineering Cell (MEC) 510 Monin, B. 161
Meme Warfare Center (MWC) 509–10 monogamy 178
memeplexes 503 Mood, A. 152
memes 499–512 Moore, J. H. 364
Memetics for Intelligence and National Defense Moore, M. 108, 264, 267
(MIND) 510–11 Moqbel, M. 420, 422
memory Moraitou, D. 67
dangerous animals 116 moral anthropocentrism 146
of diseases 113 moral disengagement 147
prosthetic 337–8 moral foundations theory 136
scripts 382, 383 morality 172
survival scenario 112 children 261
mens rea 229 death penalty 268–9
mental disorders 3–4 retribution 263–5
concept 25 robots 367
psychiatry 24–50 see also ethics
mental health Moran, T. 272
food consumption 128–9 Morikawa, T. 58, 59
immediate vs long-term rewards 131, 133 Morris, N. 336
meta-awareness 100, 102 Moscovici, S. 334
meta-cognition 102 Moskowitz, G. B. 337
Meystel, A. M. 353, 355, 356 ‘motherese’ 153
Military Deception (MILDEC) 506 Motto, J. 9–10
military memetics 505–12 Moura, J. M. B. 107–22
military robots 365–7, 368 Mugisha, J. 55
Millan, C. 204 Munday, Z. 461
Miller, G. 66, 161, 400, 409, 479, 483 Murray, H. A. 63
Miller, J. G. 175 Murtaugh, P. A. 435, 436, 439
Miller, M. B. 67, 79 music 402, 405
mindfulness 7, 94–106 mutation load 30–1, 35
Mindfulness-Based Stress Reduction (MBSR) 96,
99, 100 Naghavi, M. 51
Mineka, S. 116 Nahmias, E. 229, 231, 233
Minervini, B. P. 484, 485 Nairne, J. S. 111, 334
minimal parental investment (MPI) 191–2, 200 Nakajima, S. 153
Minkov, M. 441 narcissism 404
Minson, J. A. 161 narratives 507–8
Mirville, M. O. 321 NASCAR 385, 387–8, 389
Mishara, B. L. 73 Nass, C. I. 405, 408, 413
528 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

National Incident-Based Reporting System observational learning 374, 376–8, 383, 392
(NIBRS) 289 ‘observing self’ 101
National Institute for Health and Care Excellence Obsessive Compulsive Disorder (OCD) 12, 31, 44
(NICE) 35 Odling-Smee, F. J. 246, 247
Natural Born Killers 380 offsetting behavior 132–3
natural law 222 Öhman, A. 12
natural selection 30, 172, 173, 189, 204, 477 oleic acid 460
behavioral rules 367 Oliphant, B. 268
emotions 265 omega-3 fatty acids 35
genetic programming 358, 360 online dating 480–1, 483, 486–8, 492
mass media 400, 401 operant conditioning 378
memes 500 optimal strategies 130–2
punishment 231–2, 236, 259 oral speech 417–34
robotics 352–3 Orbell, J. 58, 59
suicide 52, 53 Organisation for Economic Co-operation and Develop-
threat cues 209 ment (OECD) 339, 488
warfare 326 Orians, G. H. 111, 114
naturalistic fallacy 201, 235 Orsolini, L. 38
naturalistic mind 111–13 Ortiz-Ospina, E. 338
nature/nurture debate 400–2 Ortner, C. N. M. 101
Navarrete, C. D. 156 Orwall, B. 377, 392
necromones 460, 463, 466, 468 Orwell, G. 505
necrophoric behavior 458–62, 463 Osgood, R. E. 325
Nedelec, J. L. 188–202 ostracism 260
Need for Speed 387 Ostrom, E. 177
negative news 403 Otterbein, K. F. 319, 324
neocortex size 262 Otto, S. P. 477, 478
neoteny, animal ethics 152, 153–4 outgroups see ingroups/outgroups
Nese, A. 308 Owen, M. J. 35
Nesse, R. M. 7, 12, 24, 25, 26, 27, 33, 34, 35–6, 37, oxytocin 463
40–1, 42, 44, 63, 76, 209, 236, 302 Oyer, P. 481
Nettersheim, J. 40
Neuberg, S. L. 204, 209, 211 Pachirat, T. 150
neural networks 353, 357–8 pain, animal ethics 145–6, 151, 157
neurocognitive warfare 505–7 pain escape, suicide 62, 63–5
neurotoxin regulation hypothesis 37 ‘pain-and-brain’ framework 52, 64–5, 66–7, 69–70, 73,
Newsome, J. 277–95 75–6, 78
niche construction theory 246–7, 254 palmar grasp reflex 108
niche-building 402 Panagiotopoulos, P. 344
niches 177 Pancoast, W. 489
Nicholas, J. S. C. 153 panic attacks 101
Nielsen, M. 409 Paoli, L. 296, 297, 298, 300, 301, 306
Nisa 148 Paoline, E. A. 210, 211, 213
nociception 145 Papanicolaou, 305, 307
Nock, M. K. 73 Paquette, D. 211
noise theories, suicide 56–7 parasitism, political institutions 172
non-communicable diseases (NCDs), public health parasocial engagement 405–6
124, 134 parental investment 282–3, 288, 405, 478–9
non-social theories 33–4 life history theory 436–7, 439, 449–51
Norenzayan, A. 304 minimal 191–2, 200
normative law 222, 223 postmortem transport 458, 462–6
Norwood, F. B. 157–8 reproductive technology 489–90
Nowak, M. 174, 228, 264, 265, 266 risk-taking behavior 375
NR3C1 gene 286 Parker, G. A 78
Nutt, D. J. 70, 71 Parlapiano, A. 154
parochial altruism 320–1
Obama, B. 212 Paternoster, R. 249
obesity, public health 123–9, 134–5 pathogens, disgust sensitivity 156
 INDEX 529

Patterson-Kane, E. G. 149, 150 Pobbe, R. 71

Paul, A. S. 159 policing, aggression/restraint balance 203–20
Paul, E. S. 147 Polimeni, J. 36
Paulhus, D. L. 67 political institutions 171–87
Pawlowski, B. 481, 483, 484 Poppelwell, Z. 304
Peck, J. H. 203 population growth 435
peer influences 378 Population Paradox 435, 438, 439–40, 448, 451
Peña-Guzmán, D. M. 53 Posner, M. I. 96
Peng, J. 375 Post, M. 159
Penn, D. J. 117–18 Post-Traumatic Stress Disorder (PTSD) 13
Penner, L. A. 253 postmortem transport 458, 462–6
Penney, S. 259 postpartum depression 11–12
Pennings, G. 490 poverty 338
Pentland, A. S. 335 Power, R. A. 33, 35, 36, 42
People for the Ethical Treatment of Animals (PETA) Pratt, T. C. 278
151, 157 predatory crimes 298–9
Perry, M. 381 prefrontal cortex 266–7, 368
Perry, S. A. 51, 59, 64, 66 Prehn, K. 269
personality Price, G. R. 173–4, 420
five dimensions 41 Price, J. 34, 36
traits 488 primary human goods 248
personality disorders (PD) 10–11, 16, 41–2 prison 243–57, 258, 260
Pessoa, L. 99, 266 revolving-door phenomenon 249
Peter, J. 487, 488 staff burnout 251–2, 253
Petersen, M. B. 177, 229, 230, 233, 243, 244–6, 253, Prisoner’s Dilemma (PD) 174, 176, 264, 303, 307–8
259, 318 privatization 176–7
Petrie, M. 418, 420, 426 probation 249, 251, 285
Petrinovich, L. 146, 153, 155 problem solving 409
Pettitt, P. 457, 458, 468, 470 The Program 380
pharmacophagy hypothesis 37–8 Prokop, P. 116
Phillips, C. J. 160 proportional representation 178, 179, 180
Phillips, D. P. 379 prosocial behavior 253, 282–3, 291
Phillips, T. 282 prostitution 298, 302, 303, 311
phobias 116, 209 protection payments 297, 299, 302, 310
anxiety disorders 12 Prounis, G. S. 467
distal causes 4 Prunetti, E. 11
Piette, A. 337 psychache 63–4, 69
Pincus, H. A. 16, 75 psychiatry 24–50
Pinel, J. P. J. 467 psycho-social factors, technology 360
Pinho, J. R. 116 psychoanalysis 5–6
Pinizzotto, A. J. 214 psychological mechanisms, obesity 127–9
Pinker, S. 67, 149, 150, 157, 235, 325, 424, 426 psychological operations (PSYOP) 506, 512
Piper, H. 149, 150 psychopaths 231, 267
Pitts, W. 353 psychopharmacology 42–3, 76
‘Pizzagate’ 504 psychotherapy 3–23
Pizzini-Gambetta, V. 301 public affairs 506–7
placebo effect 40–1 public health 123–43
plants punctuated equilibrium 362, 370
co-evolution 37–8 punishment 258–76, 304
landscape preferences 117 children 378–9
toxicity 116 corrections and rehabilitation 277–95
play crime 221–42
with animals 148–9 excessive 271
mass media 409–10, 413 incarceration 243–57
status seeking 211 see also retribution
pleiotropy 60, 61 Purzycki, B. G. 308
Plomin, R. 35 Pusatory, M. 214
Pluess, M. 281, 282 putrescine 468–9
530 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

Puts, D. 323, 479 reintegrative shaming 272–3

Puurtinen, M. 210 Renninger, L. A. 484
reproduction 9
quasigovernment criminal structures 299 costly traits 418, 421
Quinlan, R. 435–6, 437, 442, 448, 451, 490 drug use 38
Quinsey, V. L. 283 IVF 479, 481, 488–9
mass media 400, 409
Raber, I. 336 oral speech 418
race technology 477–98
bias 337 see also fertility
implicit bias 209 Reproductive Suppression Hypothesis 39
policing 203, 206, 209, 212 Research Diagnostic Criteria (RDoC) 26
racism 175, 500 resource-holding potential (RHP) 207, 208, 216
stereotypes 181 Restak, R. 413
Rachlinski, J. J. 234 restitutive justice 271
Rådestad, I. 469 retaliation 226, 304
Radway, J. 404 retribution 223, 224–5, 229–30, 234–7, 259–65,
Radzinowicz, L. 260 267–9, 271
Ragatz, L. L. 303 Reynolds, A. 180
Rahr, S. 214 Riaz, A. 116
raids 300, 309–10, 318, 319, 320, 322, 325 Ricciardelli, R. 306, 307
Raine, A. 267 Rice, S. H. 418, 422
Ramos, R. A. 408 Rice, S. K. 214
Ramsden, E. 53 Richardson, G. 38
Rand, D. G. 272 Richter, D. 108
Randler, C. 116 Ridley, M. 326
rank theories 34 Riffkin, R. 150–1, 155
Rantala, M. J. 33, 34, 43 Riordan, D. 58
Rapaport, A. 174, 175 risk factors
Raphael, S. 251, 260, 288 extrinsic 437–8, 439, 440, 450–1
Rational Emotive Behavior Therapy (REBT) 9 organized crime 303
rationality 266–7, 363 risk-need-responsivity (RNR) model 277, 278–81,
Rauch, A. 151 286–7, 288, 291
Rauwolf, P. 333–51 risk-taking behavior 124, 131–2, 374–97
Rawls, J. 178 rituals 304
recalibration theory of counter-exploitation 244–5, 253 ‘Robbers Cave’ experiments 210, 215
Rechavi, O. 353 Robertson, C. T. 238
recidivism 277–81, 287, 289–91 robotics 352–73
reciprocal altruism 173, 174, 427 three laws of 367, 369
reciprocity 172–3, 261–2, 264–5 Rockenbach, B. 308
cheating 258 Rockman, H. 482–3
direct 226, 228, 304, 305–6, 308 Rodino, I. S. 490
indirect 228, 304, 305–6, 308 Roehrborn, C. 57
strong 174, 227, 270 Rogers, J. R. 79
trust 340–1 role models, media 406
reconciliation 272–3 Rollo, C. D. 460
Red Queen hypothesis 201, 478 romance novels 404
Reese, J. 150, 151, 159, 161 romantic love 410
Reeves, B. 405, 408, 413 Rosa, E. A. 435, 439, 444
reflective decisions 130, 132 Rose, H. 235
Regan, T. 145 Rose, S. 235
regularity principle 113 Rosebury, B. 261
rehabilitation 223, 224–5, 229, 231, 234–7, 243–4, Rosen, J. D. 301, 306, 307
245–9, 252–4, 260, 278 Rosenfeld, M. J. 483, 486
RNR model 277, 278–81, 286–7, 291 Roser, M. 338
Reiman, J. 205–6 Roshchina, V. V. 42
reinforcement 378–9 Rossi, P. H. 245
 INDEX 531

Rothbart, M. K. 96 self-interest 221–2, 259

Rotsidis, A. 343 self-preservation instinct 62
Rottenberg, J. 33 selfishness 172
Roughgarden, 335 Sell, A. 244, 245, 253
Roulette, C. J. 302 senescence 56, 60
Rousseau, J.-J. 205 sensation seeking 381–2, 383, 385–7, 390–1
Roy, M. M. 153 sentencing 221–42
Rozin, P. 115, 160 Sentience Institute 150
Rueppell, O. 57 sentientism 145–6, 156
Rule, W. 180 serotonin 30
rumination hypothesis 34 Serpell, J. 147, 150, 152, 154, 155
Rusch, H. 321 Sesardic, N. 78
Russell, S. 123–43 sex ratios 178
Rusu, A. S. 243–57 Sexual Competition Hypothesis (SCH) 39–40
Ryder, R. D. 145 sexual fantasies 14
Ryninks, K. 469 sexual selection 36, 38, 39, 409, 412
sexually explicit media 404
Saad, L. 155 Shackelford, T. K. 15, 53, 59, 62, 64, 65, 204, 250,
Sadalla, E. 231 273, 283
St John-Smith, P. 24–50 shadchan 482–3
Salmon, C. 404, 405 shame 230–1, 272–3
Sampaio, M. B. 117 Shannon, C. 502, 503–4
Sampson, R. J. 192, 196, 280 Shapiro, F. 7
Sandøe, P. 154 Sharp, J. S. 151
Sandry, J. 112 Sharp, K. 503
Santos, M. L. S. 156 Shaver, P. 406
Sapolsky, R. M. 96 Shaw, C. R. 196
Saroglou, V. 304 Shem, S. 15
‘sati’ 95 Shepherd, B. 323
Savage, J. 204 Sherif, M. 210
savanna hypothesis 107, 108–11, 114 Sherman, G. D. 153, 154
Scarr, S. 401–2 Sherry, J. L. 401, 413
Scharf, P. 209, 210 Shi, A. 435, 438, 444
Scheib, J. E. 481, 490 Shi, F. 336
Scheidel, W. 309, 339 Shields, W. M. 467
Scheiter, S. 118 Shinar, D. 374, 375
Schick, K. 399, 401 Shiner, R. A. 223
schizophrenia spectrum disorders (SSD) 24, 31, 34–7 Shneidman, E. S. 60, 63, 69, 79
schizotypy 36–7 Shoemaker, P. J. 398, 403
Schlax, M. G. 435, 436, 439 Shonin, E. 96
Schlebusch, C. M. 108 Short, J. 407
Schnarch, D. 483 Shorter, J. 57
Schover, L. R. 490 Shostak, M. 148
Schramm, W. 398–9 Shrira, I 468–9
Schulkin, J. 63 Shugart, M. 179
Schultz, A. C. 359 Shultz, S. 262
Schwab, F. 407, 408, 409 Sigmund, K. 228, 266
Schwender, C. 409 Siles, I. 335
Scott, J. C. 309 Silva, R. H. da 107–22
Scottish Enlightenment 171 Silva, T. C. 107–22
Seabright, P. 478, 492 Silver, I. A. 277–95
secondary goods 248 Simons, R. L. 285
Segal, Z. V. 96, 99 Simons-Morton, B. 378, 388
Seiffert-Brockmann, J. 401 Singer, B. 14
Selby, E. A. 63 Singer, P. 145, 147, 156, 157
self-aware systems 364–5 Singhal, D. 374–97
self-defense 226 singles ads 484
532 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

Skinner, B. F. 378 Sparrow, R. 342

Skopek, J. 486, 487 speciesism 146–7, 149, 155
Skyrms, B. 176, 178 speed dating 485–6
slaughterhouses 150 Sperber, D. 161
slavery, supply chain 159 sperm donation 481, 489–92
Smith, A. 259 Staats, A. W. 79
Smith, E. A. 336 Stanley, I. 55
Smith, J. M. 78 ‘state of nature’ 205
Smith, W. 363 Steadman, H. J. 211
smoke detector principle 40–1, 44 Stearns, S. C. 28, 435, 436–7
Sneddon, L. U. 146 Stein, D. J. 26
social anxiety (SA) 12 Steinfeld, H. 157
social bonds 280 Stengel, E. 64
crime 192, 194, 195 Stephenson, J. 435, 438, 450
organized crime 301, 302, 306 Stephenson, W. 410
social brain theory 35 Stevens, A. 6, 36
social choice 175, 178 Stever, G. 398–416
social cohesion 339 Stewart, C. A. 248
social competition 34 Stewart, P. 261
social constructivism 401 Still, M. C. 195
social constructs 222 Stohler, H. R. 42
social contagion 180 Stoker, D. 325
social contract 205–6, 223, 261, 264 Stoll, M. 251
Social Contract Theory (SCT) 205 Stone, J. 337
social Darwinism 43, 175 strategic communications 401
social emotions 265 Strategic Communications Cell (SCC) 510
social exchange theory 227–8 Strauss, M. 155
social information processing model 250 stress
social insects 62 biological sensitivity 281
necrophoric behavior 458–62, 463 death response 465–6
suicide 57–8 mindfulness 94
social interaction theory 378 response system 130
social learning theory 376, 377–8, 401 Stringer, C. 107
social media 336 Strong African American Families (SAAF) 286
Bird Box Challenge 380 Stroup, R. 262–3
cyberwarfare 499 Studlar, D. T. 180
Facebook 345, 403 Stylianou, S. 245
fake news 343–4, 345 substance dependence/abuse 13–14, 16, 37
gender differences 403–4 alcohol addiction 37–8
Twitter 344, 426, 500 see also drugs
social mimicry 193 Sugiyama, Y. 462, 463, 465
social pain 52, 64, 69, 71, 72, 76 suicide 9–10, 376
social presence 406–7, 413 antisuicide defenses 65–79
social rank, crime 194 celebrities 379–80
Social Representation Theory (SRT) 334 ‘gene’ 60
social risk theory 34 suicidology 51–93
social snacking 405–6 Werther effect 379
social-cognitive theory of power 251 Sullivan, C. J. 285
socioeconomic status Sullivan, R. J. 37
crime 196 Sun, J. C. 469
obesity 126, 135 Sun, Q. 459, 460, 462
policing 206 Surrey, J. 15
reproduction 131–2 surveillance 278, 403
Socrates 221, 222 survival handicap 418
Solomon, A. 6 Suwa, G. 266
Soper, C. A. 51–93 Swistak, P. 270
Soudriette, R. 179 symbolic interactionism 401
 INDEX 533

Syme, K. L. 60–1 Townsend, J. B. 107, 108–9, 114, 117

Symons, D. 59, 479 tragedy of the commons 118, 207
transference 5
Taagepera, R. 181 transparency
tabloid media 405 artificial intelligence 336–7, 342–3
taboos, food 156 trust 341
tabula rasa 6, 15 Treating Depression Downhill (TDD) 7–8
tactical disengagement 213 Treiman, R. 153
Takeshita, R. S. C. 464, 465–6 Triad brotherhood 305
Talwar, P. 378 Triadic Reciprocal Causation model 382–3, 391
Tanaka, M. 58 trial and error learning 115
Tang, Y.-Y. 99 tribalism 338
Tasca, L. 374, 388 Trivers, R. 173, 191, 226, 228, 229, 304, 323, 478, 490
task analysis 66, 76 Troisi, A. 4, 7, 9, 25, 34, 44
Teasdale, J. D. 102 trolley problem 368
technology Trower, P. 12
cyberwarfare 499–514 Trull, T. J. 28
psycho-social factors 360 Trust Game (TG) 340–1
reproduction 477–98 trustworthiness, artificial intelligence 340–3, 344
robotics 352–73 Truszkowski, W. 504
Teich, A. H. 360, 361 Tsuang, M. 30
Teilhard de Chardin, P. 189 Tukachinsky, R. 405, 411
temporal discounting 131 Turchin, P. 179, 309, 310
Terkel, S. 339 Turing test 364, 365
Terrill, W. 206, 211 Turner, W. G. 153
terrorism Tversky, A. 232
cyberwarfare 499 Twitter 344, 426, 500
memes 504, 505, 509 Tybur, J. M. 156, 302
Terry, D. J. 336 Tyler, T. R. 215, 216
Teste, F. P. 334
thanatology 457–76 Ule, A. 270
Theodorou, A. 342, 343 Ulrich, R. S. 114
therapeutic cheerleading 9 United Nations 296, 338
Therapeutic Lifestyle Change for Depression urbanisation 125
(TLC-D) 8 Urquiza-Haas, E. G. 147
Thomas, R. J. 483 Usall, J. 57
Thompson, W. R. 325 utilitarianism 145, 223, 225, 230, 267
Thomson, J. A. 34
Thrasher, F. M. 211 vaccinations 135–8
Thrasymachus 221, 222 Valdes, L. 462
threat perception failure (TPF) 209–10 Valkenburg, P. M. 487, 488
Tifferet, S. 403 value-homophily 344
Tilly, C. 308–9, 325 Van Camp, T. 273
Tinbergen, N. 26–7, 44, 458, 459 Van der Beken, T. 296, 297
tit-for-tat 174, 264, 270, 272–3 van der Hout, E. 178
Todak, N. 210, 211, 216 van Dijk, M. 302, 305, 310
Todd, P. M. 115, 483, 485, 486 van IJzendoorn, M. H. 277, 281, 285,
Tomasik, B. 145, 146, 157, 158 286–7, 290
Tomasula, D. 157 Van Maanen, J. 212, 216
Tomlin, D. 96 Van Orden, K. A. 62, 78
Tooby, J. 56, 59, 66, 70, 71, 78, 110, 114, 116, 244, van San, M. 302–3
245, 259, 272, 303, 317 van Vugt, M. 180, 300, 302
Topolski, R. 146, 155 Vandenplas, Y. 424
Torgler, B. 477–98 vantage sensitivity 30
Toronto Transit Commission (TTC) 376 variability, suicide 52
totemism 149 Varki, A. 79n4
Toth, N. 399, 401 Värnik, P. 57
534 THE SAGE HANDBOOK OF EVOLUTIONARY PSYCHOLOGY

Veale, D. 13 Watson, J. 6
vegetarians 151, 155, 157, 158–61 Watson, L. 385
Verheggen, F. 468 Wattenberg, M. P. 179
Verweij, M. 266 Waynforth, D. 481
victim–offender characteristics 196, 198–201 Waytz, A. 342
video dating 484 Weatherford, J. 284
video games 386, 387 Weber, M. 309
Video-Feedback Intervention to Promote Positive Weedon, S. L. 58
Parenting and Sensitive Discipline 286 Weismann, A. 477
Vienna Risk-Taking Test 386 Weiss, B. 377
‘vigilance in grief’ 469 welfare tradeoff ratio (WTR) 244–5
vigilance-based cycle 97 Wells, A. 102
Vilnai-Yavetz, I. 403 Wells, J. 258–76
Vingilis, E. 375, 385, 391, 392 Wemmers, J. A 273
Vining, D. R. 40 Weng, H. Y. 272
violence Wertheimer, A. 52, 61
animal abuse link 149–50 Werther effect 379
in prisons 250 West, S. A. 253
towards females 283–4 West-Eberhard, M. J. 400
victim empathy 408 Westen, D. 16
virtual reality (VR), trust 342 Westermarck effect 78
virtue signaling, animal welfare 152, 160–2 White, C. 469, 470
Visscher, P. K. 460, 461 White, M. D. 203
Vitaglione, G. D. 387–8, 389 Whyte, S. 477–98
Vitak, J. 481 Wiblin, R. 144, 156
Volk, A. A. 110 Widiger, T. A. 28
Von Gunten, A. 41 Wiebe, R. 264, 267, 273
von Lampe, K. 298–9, 302, 304–7 Wiesenthal, D. 374–97
von Rueden, C. 322–3 Wiley, J. 60
Vorderer, P. 408 Wilkinson, G. S. 258
Vory v zakone 305 Wilks, M. 160
Vosoughi, S. 344 Williams, G. 27, 54, 56, 58, 60, 61, 65, 75, 76, 77,
‘vote/buy gap’ 159 236, 480
Williams, J. M. G. 63
wages, artificial intelligence 340 Williams, P. 4
Wakefield, J. C. 25, 26, 32 Willner, P. 75
Waldfogel, J. 251 Wilson, D. 53, 60
Waldmann, M. R. 71 Wilson, E. O. 114, 115, 148, 460
Walker, I. D. 335 Wilson, H. 333–51
Walker, J. K. 155 Wilson, J. D. 57
Walker, P. 390 Wilson, J. Q. 260
Walker, R. S. 482 Wilson, M. 194, 195, 196–9, 227, 234, 235, 284,
Walker, S. 177, 205, 211 288–90, 323, 375, 376, 378–9
Wall, P. D. 63, 70 Wilson, R. K. 342
Wallace, J. 231 Winstead, B. A. 15
Wallach, W. 367 Wisman, A. 468–9
Waller, B. M. 153 Witwicki, K. 159
Walsh, A. 258–76 Woestenburg, N. O. 490, 491
Walters, J. 214 Woll, S. B. 484
Wampold, B. E. 16, 75 Wong, C. 238
War of Attrition model 207, 208 Woodruff, P. 266
Ward, T. 243, 246–8 Woollacott, M. 99
warfare 316–30 World Bank 338
cyber 499–514 World Health Organization (WHO) 26, 39, 51
Wasserman, D. 76 Wrangham, R. 156, 300, 317, 318, 319
Watkins, E. 98 Wright, R. 56, 59
Watson, C. F. I. 462, 465 Wright, S. 56, 420, 421
 INDEX 535

Yabuta, S. 207 Zaki, J. 363

Yakuza 299, 300, 305, 306, 307 Zald, D. 403
Yamagishi, T. 341 Zalsman, G. 76
Yancey, P. 63, 65 Zanette, L. Y. 13
Yang, L. 111 Zaraska, M. 151, 156, 160
Yanomamo people 271 Zhang, W. 117
Yao, M. 462 Zhou, X. 462
Yorzinski, J. L. 116 Zillmann, D. 382
Young, H. P. 180 Zimmer, C. 478
young male syndrome 194, 195, 199, 374, 375, 383 Zinger, I. 279
Young, P. 484 zoonotic diseases 160
Young, S. G. 111 Zuboff, S. 336
YouTube 385, 392 Zuckerman, M. 381

Zahavi, A. 344, 418, 419, 420

Zajonc, R. B. 107