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The Oxford Handbook of Human Mating
 OX F O R D L I B R A RY O F P S YC H O LO G Y

The Oxford Handbook

of Human Mating
Edited by
David M. Buss
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CONTENTS

List of Contributors ix

​Introduction to Human Mating Strategies 1

David M. Buss

Part I • Theoretical Perspectives

1. Writing Trivers’s 1972 Theory of Parental Investment and
Sexual Selection 11
Robert Trivers
2. Sexual Selection and the Animal’s Mating Mind 16
Michael J. Ryan
3. The Interface of Sexual Selection, Conflict, and Evolutionary
Psychology: Emerging Core Themes 33
Jennifer C. Perry and Tracey Chapman
4. Extensions of Sexual Strategies Theory across Peoples, Cultures,
and Ecologies 66
David P. Schmitt

Part II • Attraction and Mate Selection

5. The Early Stages of Mate Selection 121
Norman P. Li and Bryan K. C. Choy
6. Computational Models of Mate Choice 154
Daniel Conroy-​Beam
7. ​The Logic of Physical Attractiveness: What People Find Attractive,
When, and Why 178
David M. G. Lewis, Kortnee C. Evans, and Laith Al-​Shawaf
8. Incest Avoidance Adaptations: Evolved Function and Proximate
Mechanisms 206
Debra Lieberman and Carlton Patrick
9. Parental Influence over Mate Choice 222
Menelaos Apostolou
10. Beyond Sex: Reproductive Strategies as a Function of Local Sex
Ratio Variation 240
Ryan Schacht and Caroline Uggla
11. Mating Strategies in Sexually Egalitarian Cultures 262
Leif Edward Ottesen Kennair, Trond Viggo Grøntvedt, Andrea M. Kessler,
Steven W. Gangestad, and Mons Bendixen
12. Mating Strategy Variation by Sexual Orientation 286
David A. Frederick, Jenna C. Alley, Scott Semenyna, and Justin R. Garcia

Part III • Mate Competition

13. Contest Competition for Mates and the Evolution of Human Males 317
David Puts, David Carrier, and Alan R. Rogers
14. An Evolutionary Review of Female Intrasexual Competition 378
Maryanne L. Fisher and Jaimie Arona Krems
15. Mate Competition between the Sexes: Evidence from Two Non-​Western
Cultures 404
Scott W. Semenyna, Francisco R. Gómez Jiménez, and Paul L. Vasey

Part IV • Pair-​Bonded Relationship Dynamics

16. Evolving Connections: Attachment and Human Mating Strategies 427
Gurit E. Birnbaum
17. Archaeology of love: A Review of the Ethnographic Exploration of Love
around the World 447
William Jankoviak and Alex J. Nelson
18. Marital and Sexual Satisfaction 466
Andrea L. Meltzer
19. Sex Differences in Jealousy: The State of the Theory 483
John E. Edlund, Brad J. Sagarin, and Kristyn M. Kinner
20. Mate Guarding 502
Valerie G. Starratt and Todd K. Shackelford

Part V • Sexual Conflict in Mating

21. Deception in Human Mating 515
Gayle Brewer
22. Marriage and Monogamy in Cross-​Cultural Perspective 531
Brooke A. Scelza
23. Sperm Competition 555
Valerie G. Starratt and Todd K. Shackelford
24. Violent Mates 566
Joshua D. Duntley

vi C o n ten ts
25. The Dark Triad Traits and Mating Psychology 590
Peter K. Jonason and Vlad Burtaverde
26. Sexual Harassment 606
Kingsley R. Browne
27. Sexual Coercion 629
Joseph A. Camilleri
28. Women’s Avoidance of Sexual Assault 648
Rachel M. James, Melissa M. McDonald, and Viviana Weekes-​Shackelford

Part VI • Mating and Endocrinology

29. Hormones and Human Mating 667
James R. Roney
30. Understanding Women’s Estrus and Extended Sexuality: The Dual
Sexuality Framework 700
Steven W. Gangestad, Tran Dinh, Lauren Lesko, and Martie G. Haselton
31. Ovulatory Cycle Effects and Hormonal Influences on Women’s Mating
Psychology 739
Julia Stern and Lars Penke
32. The Impact of Hormonal Contraceptives on Women’s Sexual and
Mating Psychology 756
Katja Cunningham and Sarah E. Hill

Part VII • Mating in the Modern World

33. Mating in the Digital Age 777
Helen E. Fisher and Justin R. Garcia
34. Popular Culture and Human Mating: Artifacts of Desire 796
Catherine A. Salmon and Rebecca L. Burch
35. Evolutionary Mismatch and Human Mating: Understanding the Mating Mind
in the Modern World 814
Cari D. Goetz

Index 833

Conte nts vii

 L I S T O F CO N T R I B U TO R S
Jenna C. Alley, Ph.D.
University of California, Los Angeles
Laith Al-​Shawaf, Ph.D.
University of Colorado
Menelaos Apostolou, Ph.D.
University of Nicosia
Mons Bendixen, Ph.D.
Norwegian University of Science and
Technology (NTNU)
Gurit E. Birnbaum, Ph.D.
Reichman University (IDC Herzliya)
Gayle Brewer, Ph.D., Ed.D.
University of Liverpool
Kingsley R. Browne, M.A., J.D.
Wayne State University
Rebecca L. Burch, Ph.D.
State University of New York at
Oswego
Vlad Burtaverde, Ph.D.
University of Bucharest
David M. Buss, Ph.D.
University of Texas, Austin
Joseph A. Camilleri, Ph.D.
Westfield State University
David Carrier
University of Utah
Tracey Chapman, Ph.D.
University of East Anglia
Bryan K. C. Choy, M.Phil.
Singapore Management University
Daniel Conroy-​Beam, Ph.D.
University of California, Santa
Barbara
Katja Cunningham, M.S.
Texas Christian University
Tran Dinh, M.A.
University of New Mexico
Joshua D. Duntley, Ph.D.
Stockton University
John E. Edlund, Ph.D.
Rochester Institute of Technology
Kortnee C. Evans, B.Sc.
Murdoch University
Helen E. Fisher, Ph.D.
The Kinsey Institute
Maryanne L. Fisher, Ph.D.
Saint Mary’s University
David A. Frederick, Ph.D.
Chapman University
Steven W. Gangestad, Ph.D.
University of New Mexico,
Albuquerque
Justin R. Garcia, Ph.D.
Indiana University
Cari D. Goetz, Ph.D.
California State University San
Bernardino
Francisco R. Gómez Jiménez, Ph.D.
University of Toronto Mississauga
Trond Viggo Grøntvedt, Ph.D.
Norwegian University of Science
and Technology (NTNU)
Martie G. Haselton, Ph.D.
University of California, Los Angeles
Sarah E. Hill, Ph.D.
Texas Christian University
Rachel M. James, M.S.
Oakland University
William Jankowiak, Ph.D.
University of Nevada, Las Vegas
Peter K. Jonason, Ph.D.
University of Padua and Cardinal
Stefan Wyszynski University in
Warsaw
Leif Edward Ottesen Kennair, Ph.D.
Norwegian University of Science
and Technology (NTNU)
Andrea M. Kessler, M.Sc.
Norwegian University of Science
and Technology (NTNU)
Kristyn M. Kinner, B.A.
Rochester Institute of Technology
Jaimie Arona Krems, Ph.D.
Oklahoma State University
Lauren Lesko
University of California, Los Angeles
David M. G. Lewis, Ph.D.
Murdoch University
Norman P. Li, Ph.D., M.B.A., M.A., B.A.
Singapore Management University
Debra Lieberman, Ph.D.
University of Miami
Melissa M. McDonald, Ph.D.
Oakland University
Andrea L. Meltzer, Ph.D.
Florida State University
Alex J. Nelson, Ph.D.
Appalachian State University
Carlton Patrick, J.D., Ph.D.
University of Central Florida
Lars Penke, Prof., Dr.
Georg August University Göettingen
Jennifer C. Perry, Ph.D., M.Sc., B.Sc.
University of East Anglia
x L i s t of C on tr ibutor s
David Puts, Ph.D.
Pennsylvania State University
Alan R. Rogers, Ph.D.
University of Utah
James R. Roney, Ph.D.
University of California, Santa
Barbara
Michael J. Ryan, Ph.D.
University of Texas
Brad J. Sagarin, Ph.D.
Northern Illinois University
Catherine A. Salmon, Ph.D.
University of Redlands
Brooke A. Scelza, Ph.D.
UCLA
Ryan Schacht, Ph.D.
East Carolina University
David P. Schmitt, Ph.D.
Brunel University London
Scott W. Semenyna, Ph.D.
Stetson University
Todd K. Shackelford, Ph.D.
Oakland University
Valerie G. Starratt, Ph.D.
Nova Southeastern University
Julia Stern, Ph.D.
University of Bremen
Caroline Uggla, Ph.D.
Stockholm University
Paul L. Vasey, Ph.D.
University of Lethbridge
Robert Trivers, Ph.D.
Rutgers University
Viviana Weekes-​Shackelford, Ph.D.
Oakland University
Introduction to Human Mating Strategies

David M. Buss

Abstract
In sexually reproducing species, mating success is a non-​negotiable requirement for
evolutionary fitness. Consequently, selection has created a rich array of adaptations that
are products of a long and unbroken line of human ancestors, each of whom succeeded in
the complex game of mating. This Handbook showcases the current state of knowledge
about those adaptations. These include mate preferences, tactics of attraction, forms of
mate competition, tactics for dealing with sexual conflict, modes of mate retention, mate
switching strategies, and many more. Chapters on the endocrinology of mating adaptations
provide state-​of-​the-​art knowledge about some of the key biological drivers. Chapters
on mating in the modern world highlight key ways in which mating adaptations, forged
over millions of years in environments long gone, get expressed in modern environments,
sometimes creating evolutionary mismatches. This Handbook is not the final word on
human mating strategies. Rather, it gives readers and researchers an impressive foundation
of what is known and unknown, and importantly, a roadmap for future discoveries about
what may be the most complex evolved psychology humans possess.

Key Words: human mating strategies, sexual selection, mate preferences, mate
competition, evolutionary mismatches

The scientific study of human mating strategies is one of the major success stories of
evolutionary psychology. The explosion of evolutionarily anchored theories and thou-
sands of empirical studies of human mating is unique among the social sciences in at least
two respects—​the volume of hypothesis generation and the cumulative quality of the
scientific empirical testing. This makes good theoretical sense. In all sexually reproducing
species, evolutionary processes must all pass through the rigorous filters of successful mat-
ing. Survival is not enough. Each living human comes from a long and literally unbroken
chain of ancestors, each of whom succeeded in selecting a mate, attracting a mate, being
reciprocally chosen by that mate, navigating the complexities of sexual intercourse suf-
ficient for conception, having a conceptus who survived the nine-​month hurdles to be
born, and after birth survived to reproductive age to begin the process anew.
Passing through these successive hurdles had to happen not just a dozen times but
millions upon millions of iterations going back through the distant mists of human evo-
lutionary history, our primate lineage, our mammalian lineage, and the origin of sexual
reproduction itself more than a billion years ago. If any one of our ancestors had failed in
any of these tasks, the chain would be irreparably broken and we would not be alive to read
these pages. In this important sense, we are all evolutionary success stories. As descendants
of this unimaginably long line of forbears, each us carries with us the finely honed adapta-
tions that led to our ancestors’ success. The current Handbook of Human Mating provides
an up-​to-​date summary of the current state of the science of human mating—​the modern
theories, hypotheses, predictions, and empirical findings relevant to each.

Sexual Selection Theory

Sexual selection theory (Darwin, 1871) provides the most important theoretical framework
for understanding the mating adaptations of sexually reproducing species (e.g., Andersson,
2019). Sexual selection centers on the evolution of adaptations not due to survival success but
rather due to mating success. Darwin identified two causal processes by which mating success
could be attained—​intrasexual competition and preferential mate choice. The classic example
of intrasexual competition is two male stags locking horns in combat. Winners in these physi-
cal contests gained preferential sexual access to females, and so passed on genes for attributes
that led to their success such as greater strength, agility, speed, and perhaps fearlessness and
bellicosity. The attributes of losers bit the evolutionary dust because they failed in this brutal
game of mating. Contest competition turns out to have been an extremely important sexually
selective process in humans (Puts et al., this volume).
The second process of sexual selection articulated by Darwin identified the evolution of
mating success through preferential mate choice. If members of one sex preferred certain
qualities in the other sex in mating contexts, then those who possessed the desired quali-
ties were more frequently chosen. Over evolutionary time, if there exists some degree of
consensus about the qualities desired and those qualities are partly heritable, then the
desired qualities evolve or increase in frequency.
Although Darwin envisioned intrasexual competition as pertaining primarily to males
and preferential mate choice pertaining primarily to females, when it comes to humans,
both processes apply to both sexes, as the chapters in this Handbook document. Moreover,
the modes of intrasexual competition in humans involve not just physical contests,
although it’s clear that these have been important. Rather, arenas of intrasexual competi-
tion include tactics for navigating status hierarchies and the use of language to influence
social perceptions of one’s own mating desirability and to impugn the reputation of intra-
sexual rivals (Buss & Dedden, 1990; Fisher & Krems, this volume).
Sexual selection theory itself has witnessed important theoretical developments since
Darwin’s 1871 articulation of it (see Ryan, this volume). Trivers (1972), for example, fur-
nished a major theoretical advance by identifying the relative parental investment of each
sex as a key determinant of which sex does the competing and which sex does the choos-
ing (see Trivers, this volume, on the personal context in which he developed his theory).
Sexual selection theory has also expanded to include processes such as sperm competition
(Starratt & Shackelford, this volume) and sexual coercion (see Camilleri, this volume).
The chapters in this volume showcase the “state of the art” of sexually selected human
mating strategies and the underlying psychological adaptations that give rise to them.

2 Davi d M. Buss
Sexual Conflict Theory
Another critical theoretical development since Darwin’s time has been sexual conflict
theory (Parker, 2006; Perry & Chapman, this volume). When the genetic interests of
females and males diverge, sexual conflict will ensue. There exist many domains of sexual
conflict in humans, zones in which the optimal mating strategy from a female perspec-
tive differs from the optimal mating strategy from a male perspective. It is sometimes
in a male’s best interest to initiate sex sooner, or with less investment, compared to the
optimal interests of the female. These differing fitness interests create sexually antagonistic
arms races very much analogous to those that occur between predators and prey. Each sex
evolves adaptations to influence the other to be closer to its own optimum, which creates
counteradaptations or defenses in the other to resist that influence and to manipulate the
other sex to closer to its own optimum. This form of sexually antagonist evolution is often
perpetual.
The many chapters on sexual conflict in this volume highlight the theoretical utility
of sexual conflict theory in guiding researchers to discoveries that were entirely unknown
prior to this theoretical development. These include predictable forms of deception in
mating (Brewer, this volume), infidelity and jealousy (Scelza, this volume), adaptations
for sperm competition (Starratt & Shackelford, this volume), intimate partner violence
(Duntley, this volume), various forms of sexual coercion (Brown, this volume; Camilleri,
this volume) as well as women’s defenses against sexual coercion (James et al., this volume).

Sexual Strategies Theory

Social scientists, including psychologists, typically receive no training in evolutionary
biology. Consequently, most were unaware of these key theoretical developments in evo-
lutionary theory and their potential applicability to human mating. In the last half of the
1980s and early 1990s, this started to change. Cunningham (1986) launched experimen-
tal tests of the evolution of female beauty from an evolutionary perspective. Kenrick et al.
published a paper documenting that social dominance increased the attractiveness of men
as mates but did not increase the attractiveness of women (Sadalla et al., 1987). And Buss
(1989) published the first large-​scale cross-​cultural tests of evolutionary hypotheses about
sex differences in human mate preferences.
The first major attempt to develop a more comprehensive evolution-​based psychological
theory of human mating strategies came under the rubric “sexual strategies theory” (Buss
& Schmitt, 1993). This theory, containing nine key premises and twenty-​two empirical
predictions, proposed that humans have evolved a menu of mating strategies, not a single
mating strategy. It provided a task analysis of the different adaptive problems that women
and men recurrently face in mating, cross-​cut with the temporal dimension of short-​term
mating (e.g., casual sex and brief affairs) and long-​term mating (e.g., marriage and other
temporally enduring mating relationships). It proposed that women as well as men engage
in short-​term mating, but the adaptive challenges they face when pursuing this strategy

In troduc tion to Human Mating Strate gie s 3

differ profoundly. The reproductive benefits to men of pursuing a short-​term mating strat-
egy are fairly straightforward, since men’s reproductive success is most heavily limited by
the number of fertile sex partners they can attract. Consequently, sexual strategies theory
predicted that men would desire a larger number of partners than women do, let less time
elapse before seeking sexual intercourse, and possess a host of other psychological design
features that would promote the success of a short-​term mating strategy (e.g., the male
sexual overperception bias, first predicted by Buss, 1994).
Although women generally cannot (and never could) increase their direct reproduc-
tive success through short-​term mating since adding additional sex partners would
rarely have resulted in more offspring, they could use short-​term mating to obtain key
adaptive benefits. According to sexual strategies theory, these might include (1) imme-
diate access to resources, (2) assessing several men through trial runs to determine
which would be a good long-​term mate, (3) cultivating a backup mate for insurance
should something go wrong with her long-​term relationship, (4) using short-​term mat-
ing as a tactic for “trading up” to a higher mate value partner, and perhaps (5) obtain-
ing higher-​quality genes (e.g., genes for good health) from an affair partner than she
could obtain from her regular partner. In the intervening years since the publication
of sexual strategies theory in 1993, a tremendous volume of research has focused on
testing hypotheses and predictions based on these theoretical benefits (e.g., Greiling &
Buss, 2000, on women’s perceptions of the benefits of short-​term mating; Buss et al.,
2017 on the mate-​switching hypothesis; Gangestad et al., this volume, on the good
genes hypothesis).
Sexual strategies theory also provided a task analysis of the adaptive challenges
each sex faces when pursuing a long-​term mating strategy. Women, for example, were
hypothesized to face challenges such as (1) identifying men who are able to invest in
them and their offspring, (2) identifying men who are willing to invest in them and
their offspring, (3) identifying men able and willing to protect them (e.g., from sexual
aggression from other men), (4) assessing men who give cues to long-​term commit-
ment to them, and (5) assessing men who give cues to “good dad” qualities linked with
parenting skills. Men, while facing some of these adaptive challenges in selecting their
mates for long-​term mating such as commitment and good parenting, also had to solve
additional sex-​differentiated ones: (1) choosing women likely to be sexually faithful,
thus solving the problem of paternity uncertainty (stemming from the fact that fertiliza-
tion occurs internally within women), and (2) identifying women of high reproductive
value (i.e., future reproductive capacity).
Since the 1993 publication of sexual strategies theory, a large body of empirical evi-
dence has accumulated that supports many of the specific predictions generated by it (see
Buss & Schmitt, 2019, for a review). Moreover, sexual strategies theory has been extended
in new directions to include testing hypotheses across peoples, cultures, and ecologies (see
Schmitt, this volume).

4 Davi d M. Buss
 In this Handbook, multiple chapters showcase important theoretical developments and
novel empirical extensions of the scientific understanding of human mating strategies.
In the section “Attraction and Mate Selection,’ Li and his colleagues outline the logic
and evidence of strategies in the early stages of mate selection. Conroy-​Beam provides a
novel computational model of mate selection that yields a sophisticated way of viewing
the multidimensional process of mate selection. Lewis and colleagues provide a “state
of the science” summary of theory and research on physical attractiveness using a cue-​
based approach. Lieberman and Patrick discuss theory and research on incest avoidance
adaptations. Apostolou presents arguments and evidence for the importance of parents in
influencing, and in some cases selecting, the mates of their daughters and sons. Schacht
and Uggla highlight the importance of sex ratio, which surely varied tremendously across
cultures and over time, on the mating strategies people pursue.
Kennair and colleagues discuss mating strategies in sexually egalitarian cultures such
as Norway. Although some theories, such as traditional sex role theory, predict that sex
differences should diminish or vanish in sexually egalitarian cultures, the data do not sup-
port those predictions. Indeed, some sex differences become larger, not smaller, in sexually
egalitarian cultures. The concluding chapter in this section, by Frederick and colleagues,
summarizes what is known about mating strategies as a function of individual differences
in sexual orientation such as gay men, lesbian women, and bisexual individuals.

Pair-​Bonded Relationship Dynamics

Once mate selection and mate attraction processes have been successfully implemented,
complex dynamics occur within those who form long-​term pair-​bonded mating rela-
tionships. Birnbaum examines the role of attachment within pair-​bonds. Jankoviak and
colleague articulate the importance of love as universal phenomenon within long-​term
mateships. Meltzer explores the determinants of marital and sexual satisfaction within
relationships. Edlund and colleagues examine the importance of jealousy within mate-
ships, which some have argued is as central as love in human mating (Symons, 1979).
Jealousy is an emotion often tied to mate guarding, the subject of the final chapter in this
section, written by Starratt and Shackelford.

The Endocrinology of Human Mating

All mammals have complex endocrine systems that influence, and are influenced by,
mating, and humans are no exception. These are sex-​differentiated to an important extent.
Reproductive-​age women who are not on hormonal contraceptives, for example, experi-
ence predictable endocrinological changes as they progress through their ovulatory cycle.
Roney kicks off this section by providing a broad overview of the state of the science on
hormones and human mating. However, there is currently scientific controversy about
the effects of the ovulation cycle on women’s mating strategies. Gangestad and colleagues
argue that women possess a “dual mating strategy,” suggesting that women have not “lost

In troduc tion to Human Mating Strate gie s 5

estrus” as many have argued, and moreover show preference and other mating strategy
shifts depending on whether the odds of impregnation are high or low. Sterns and Penke,
in contrast, argue that the cycle-​phase shifts are more ephemeral or absent, according to
large-​scale attempts at replication. Resolution of this ongoing debate must await further
research within this vibrant and cutting-​edge area of the evolution of human mating.
Concluding this section is a lively chapter by Cunningham and Hill who summarize the
evidence on the effects of modern hormonal contraceptives on women’s sexuality and
mating psychology.

Mating in the Modern World

The final section of the Handbook explores mating in the modern world, which has
changed dramatically from the environment in which humans spent 99 percent of their
evolutionary history. Helen Fisher and Justin Garcia explore mate choice in the digital
age, with the dramatic surge in online dating and mating. Salmon and Burch examine
popular culture in the modern world, including pornography, romance novels, and lit-
erature. Concluding this section is a chapter by Goetz, who outlines a number of key
“mismatches” between the mating environments in which humans evolved and those they
now inhabit.
I hope this Handbook provides a key reference for researchers and budding scientists
who devote their efforts to uncovering the complexities of human mating. Biologist
Theodosius Dobzhansky famously noted that “nothing in biology makes sense except in
light of evolution.” It may not be too far a stretch to say that “nothing in human behavior
makes sense except in light of human mating.”

Acknowledgements
This Handbook owes a great debt to Patrick Durkee, who offered suggestions through-
out its creation and provided valuable feedback on a handful of chapters. Thanks also go
to Joan Bossert, the editor at Oxford University Press who believed in the importance
of this Handbook, and to Martin Baum, editor at Oxford University Press, who helped
marshall the Handbook to completion and publication. Mostly I wish to thank the several
dozen authors who wrote sterling chapters for this Handbook.

References
Andersson, M. (2019). Sexual selection. Princeton University Press. (Original work published 1994)
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. (2016). The evolution of desire: Strategies of human mating. Basic Books. (Original work pub-
lished 1994)
Buss, D. M., & Dedden, L. A. (1990). Derogation of competitors. Journal of Social and Personal Relationships,
7(3), 395–​422.
Buss, D. M., Goetz, C., Duntley, J. D., Asao, K., & Conroy-​Beam, D. (2017). The mate switching hypothesis.
Personality and Individual Differences, 104, 143–​149.

6 Davi d M. Buss
Buss, D. M., & Schmitt, D. P. (1993). Sexual strategies theory: An evolutionary perspective on human mating.
Psychological Review, 100(2), 204–​232.
Buss, D. M., & Schmitt, D. P. (2019). Mate preferences and their behavioral manifestations. Annual review of
psychology, 70, 77–​110.
Cunningham, M. R. (1986). Measuring the physical in physical attractiveness: quasi-​experiments on the socio-
biology of female facial beauty. Journal of personality and social psychology, 50(5), 925–​935.
Darwin, C. (1871). The descent of man and selection in relation to sex. Murray.
Greiling, H., & Buss, D. M. (2000). Women’s sexual strategies: The hidden dimension of extra-​pair mating.
Personality and individual Differences, 28(5), 929–​963.
Parker, G. A. (2006). Sexual conflict over mating and fertilization: An overview. Philosophical Transactions of the
Royal Society B: Biological Sciences, 361(1466), 235–​259.
Sadalla, E. K., Kenrick, D. T., & Vershure, B. (1987). Dominance and heterosexual attraction. Journal of per-
sonality and social psychology, 52(4), 730–​738.
Symons, D. (1979). The evolution of human sexuality. Oxford University Press.
Trivers, R. L. (1972). Parental investment and sexual selection. In B. Campbell (Ed.), Sexual selection and the
descent of man (pp. 136–​179). Aldine. David M. Buss, August 9, 2021

In troduc tion to Human Mating Strate gie s 7

 PART
I
Theoretical
Perspectives
  riting Trivers’s 1972 Theory of
W
C H A P T E R

1 Parental Investment and Sexual

Selection

Robert Trivers

Abstract
A brief summary of the most wide-ranging paper of mine, covering the evolution of
sex differences in all species. Especially novel is the application to differential mortality
by sex, which varies widely across species and higher taxa. The paper provided a
framework, anchored in sexual selection and parental investment, for discovering and
explaining many sex differences in human mating strategies. The paper has become
one of the most widely cited publications in evolutionary biology, accruing more than
17,000 scientific citations as of the year 2022. It is also one of the most widely cited
papers in the mushrooming field of evolutionary psychology.

Key Words: Robert Trivers, parental investment, sexual selection, sex differences, mating

I have described in print and in detail (Trivers 2002) how I came to write this chapter
so I will not go into great detail now. There were three parts: pigeon behavior (Drury), key
logic (Mayr, Bateman, 1948), and also valuable sublogic (Williams). Were pigeon couples
really monogamous with no internal conflict (ornithology); nonsense (Drury, personal
communication, date); what was the underlying logic (parental investment (Bateman,
1948; Mayr) and sublogic (Williams, 1966).
The first was to see that the conflict-​free monogamous bird family was a fantasy of bird
scientists, supported by neither evidence nor logic. All you had to do was watch an actual
bird couple—​up close and personal—​and you could see the truth. Why would a male
force his partner (for life) into sleeping on the sloping roof every night instead of the gut-
ter, where all the other pigeons slept including himself? Does it have anything to do with
the fact that in the gutter next to her would be another “monogamous” male sitting close
to his partner? It was Bill Drury who taught me this entire approach. And Bill did not joke
around. When I invented a patently nonfunctional explanation for the function of a gull
trait we were observing—​side by side with binoculars—​he told me “Never assume the ani-
mal you are studying is as stupid as the one studying him.” I was that stupid animal. Here
is a more extensive description of Bill Drury and what he meant to me (Trivers 2002):
“Go Thou to the Pigeon”
After I had spent a year or so watching herring gulls and other sea birds with Bill Drury,
I wanted to start a project on a species of my own, a species that I could study on land.
I believe I suggested the lesser marsh wren, so to speak—​that is, a species whose social
behavior and ecology had not yet been studied, though something would be known about
a closely related species. Drury immediately batted down that idea. He said it would take
me eighteen months to find the species on a regular basis and another eighteen months to
acclimate individuals sufficiently to me to permit detailed behavioral observations. That it
had not yet been studied, he said, might better be taken as a warning than as an invitation.
He suggested I go in the other direction—​study the pigeon, he said. They were everywhere
in Cambridge and too ugly (due in part to earlier domestication) and common to attract
any ornithologist since a monograph by Whitman (1919). The variability in feather pat-
terns that helped make them ugly also made individuals easy to identify, so behavioral
observations of known individuals could begin right away without the need to capture and
handle the birds. As it turned out, at the North Cambridge third-​floor apartment where I
was living, there were in fact pigeons that roosted on the roof of the house next door that
could provide a steady stream of behavioral observations right through the night!
What soon became clear in this monogamous species was that males were sexually
much more insecure than were females, and males acted to deprive their mates of what
they would be perfectly happy to indulge in themselves, that is, an extra-​pair copulation.
For example, the group outside my window began with four pigeons—​two mated couples.
They slept next to each other in the gutter of the roof of the house next door. They often
settled on the roof any time after four o’clock in the afternoon. When spending the night
together as a foursome, the two males, although they were the more aggressive sex, always
sat next to each other with each one’s mate on the outside. By sitting next to each other,
the males could ensure that each one was sitting between his mate and the other male.
Then, for a period of several days, a new male arrived and was regularly attacked by each
of the two resident males and driven off. Finally, after four or five days of persistence, such a
male might still be sleeping twenty yards down the gutter from the other four pigeons, and
subject to attack without notice. The very day he arrived with his own mate, however, the
distance to the other birds was cut in half, suggesting that male concern about male visitors
might be associated with some sexual threat or increased chance that his mate would indulge
in an extra-​pair copulation. More striking still, when the third couple managed to join the
other two, it was no longer possible for each male to sit between his own mate and all other
males. What happened then was that the outermost males kept their mates on the outside,
thus sitting between their mate and the other two males, but the innermost male forced his
female onto the sloping roof in front of them, rather than allow her to sit between him and
his neighbor to the right! The female was not happy with this situation and would return
to the more comfortable (and warmer) gutter, only to be forced back onto the sloping roof.
Sometimes she would wait for him to fall asleep and would slip down beside him unnoticed,

12 Ro b ert Tr iv er s
but I would soon hear roo-​koo-​kooing out my bathroom window and would rush to see
her pushed back onto the roof. This, for me, was a surprising observation because it put the
lie to the notion, so common in ornithology and evolutionary thinking at the time, that
the monogamous relationship was one without internal conflict. Here was a male willing to
force his own mate, mother of his offspring-​to-​be, up onto the sloping roof all night long
because of his sexual insecurities. This suggested relatively strong selection pressures.
Whitman (1919) reported a sex difference in behavior upon viewing the partner in adul-
tery that I thought was instructive along these same lines. Whitman said that when a male
pigeon saw his female about to begin copulating with another male, he flew straight at the
second male, attempting to knock him off her; that is, he interrupted the copulation as soon
as possible. By contrast, a female seeing her own mate involved in the same behavior would
not attempt to stop the copulation but would intervene immediately afterward, separate the
couple, and act to keep the other female away from her mate. What was going on here? The
obvious answer was suggested by the relative investment of the two sexes in the offspring,
certainly at the time of copulation. The male’s investment at copulation is trivial, or relatively
minor, but the female’s investment may be associated with a year’s worth of reproductive
effort. Thus, males chosen as extra-​pair partners by females enjoy the possibility of a large
immediate benefit (paternity of offspring who will be reared by the female with the help of
another male) and similarly inflict a large cost on the “cuckolded” or genetically displaced
male. These large potential selective effects would explain both a male’s eagerness to indulge
in such extra-​pair copulations and his anxiety that his own mate might act similarly!”

“Bateman ’48 in Heredity”

Were it not for Ernst Mayr I would never have heard of “Bateman ’48 in Heredity.” I
was taking a reading course from him on genetics and having done no reading on genetics
I told him pigeon stories. “Have you read ‘Bateman ’48 in Heredity’?” (Never heard of it.)
“Go read it; it has the key to what you are talking about.”
Three weeks later I returned and still not having done any work on genetics, I launched
more pigeon stories at him, when he briskly cut me off: “Have you YET read ‘Bateman
’48 in Heredity’?’ No, I had not. “I will not continue this conversation until you have,”
Mayr said, turning away from me. That was how Ernst Mayr made sure I read the key
paper—​Bateman 1948 in Heredity!
And read it I did. That evening I xeroxed the paper when Harvard did not charge a
graduate student for doing so.
And Bateman was the key. It had variance in reproductive success as a function of dif-
ferences in relative parental investment; the less investment in offspring, the higher vari-
ance in their number. The variance data were unambiguous: 4 percent of females had 0
reproductive success; the rest varied by their ability to produce eggs. Fully 21 percent of
males had no surviving offspring. This was because of low male parental investment. The
cost of sperm was trivial compared to the cost of eggs. So, males were in competition with

Writing Trivers’s 1972 Theory of Parental Investment and Sexual Selection 13

each other and some did better. They either outmuscled their brothers or they were more
attractive to females.
It is interesting that no one, but no one, cited “Bateman ’48 in Heredity.” The paper was
completely overlooked. Ironically, “Bateman ’49,” a trivial paper on Drosophila genetics,
was cited numerous times. For me, “Bateman ’48 in Heredity” immediately opened up a
brand-​new world unexplored by others—​the connection between variance in reproduc-
tive success and relative parental investment. The latter was defined by the tendency for
parental investment to inhibit future parental investment. Eggs vs. sperm. All I had to do
was fill in the details.
George Williams had a strong influence on my thinking, of which I became conscious
only after I had finished writing the chapter. I was about to teach a course at Harvard in
1971 on natural selection and social behavior in the anthropology department under Irv
DeVore. Since I was preparing for the course, I reread Williams (1966) and was astonished
to see how much of my own chapter was borrowed from his book. Sex role reversed spe-
cies. Very simple. Males invest more than females. So, there’s a sex role reversal based on
sex reversal in parental investment. Differential mortality by sex. Which sex dies faster
under what conditions? He did not solve the over-​age-​17 problem, but he raised it. There
were other examples.

Citations
Parental investment and sexual selection has been cited more than 17,000 times. This
is extremely high for a theoretical paper, one of the top 5 of 30 million. It is not a direct
measure of importance or quality. For example, the most important paper published in all
of 20th-​century biology was Watson and Crick (1953), which gave the structure of DNA,
the double helix. It has been cited half as often as reciprocal altruism which has been
cited some 13,000 times. Sometimes I like to joke that of course Watson and Crick is less
important than reciprocal altruism, but is it really less than half as important as reciprocal
altruism, as the data would actually suggest?
Why is a paper cited so often? On the one hand, parental investment and sexual selec-
tion give you a general theory for the evolution of sex differences in all species. They
therefore apply wherever there are two sexes. And they link together data from all different
species.
Perhaps more important than a general theory of sex differences, parental investment
and sexual selection defined parental investment. And it was different than parental care,
it was different than cost of parental care. It was parental investment and an analogy
between growth of populations and growth of money. You invest money to get back more
money in the future. You invest in current offspring in order to get more offspring in the
future. Once you define a variable, it is natural to get cited whenever it is first used.

14 Ro b ert Tr iv er s
The Writing of the Chapter
I wrote the chapter in several stages.
I began by rereading the entire literature start to finish. I had assembled about 70
articles over two or three years. Some principles became obvious when the whole literature
was read, while only suspected from reading the literature one by one. I was supposed to
be focused on amphibians and reptiles and decided to concentrate on differential mortal-
ity by sex as the only topic they might contribute to. Alas, data from nature were useless
since they measured not adult sex ratio but ease of capture—​males disperse more widely,
so they encounter traps more frequently and are less timid so they enter them more often.
I therefore expanded my subject to include all organisms.
This stage took about seven months, reading the entire available literature and organiz-
ing it into subcategories.
I then decided to embed the entire literature simultaneously in my brain “without fear
or favor” (e.g., bias due to time of acquisition). To do so, I mixed all the papers up at ran-
dom and then spent the next three days speed-​reading the whole lot.
At this point I was ready to write the paper. I spent about nine weeks writing the chap-
ter itself. Emended and improved since then, but what you see is what you got.

References
Bateman, A. J. (1948). Intra-​sexual selection in Drosophila. Heredity, 2(3), 349–​368.
Trivers, R. L. (1971). The evolution of reciprocal altruism. Quarterly Review of Biology, 46(1), 35–​57.
Trivers, R. L. (1972). Parental investment and sexual selection. In B. Campbell (Ed.), Sexual selection and the
descent of man (pp. 136–​179). Aldine.
Trivers, R. (2002). Natural selection and social theory: Selected papers of Robert Trivers. Oxford University Press.
Watson, J. D., & Crick, F. (1953). A structure for deoxyribose nucleic acid. Nature 171(3), 737–​738.
Whitman, C. O. (1919). Posthumous works: The behaviour of pigeons. 3 vols. H. A. Carr, Ed. Carnegie Institution
of Washington.
Williams, G. C. (1966). Adaptation and natural selection: A critique of some current evolutionary thought (Vol.
75). Princeton University Press.

Writing Trivers’s 1972 Theory of Parental Investment and Sexual Selection 15

 S exual Selection and the
C H A P T E R

2 Animal’s Mating Mind

Michael J. Ryan

Abstract
Moving from Darwin’s theory of natural selection, I briefly review his theory of sexual
selection and focus on his idea of female mate choice and the sexual aesthetics that
motivate it. I review some early hypotheses on the adaptive significance of mate
choice. Going forward, I then introduce studies that attempt to uncover the underlying
mechanisms that define a female’s sexual aesthetics. Much of evolutionary psychology
has been derived from basic notions in evolutionary biology, especially in mate choice.
Here, I turn the tables and review studies of animal mate preferences that have been
inspired by studies of humans in the fields of psychophysics, behavioral economics, and
neuroaesthetics.

Key Words: sexual selection, mate choice, psychophysics, sexual beauty, irrational choice

Sexual Selection Theory

It has been 150 years since the publication of The Descent of Man and Selection in
Relation to Sex (Darwin 1871), and a bit longer since Charles Darwin exclaimed, “. . . the
sight of a feather in a peacock’s tail, whenever I gaze at it, makes me sick!” (Darwin 1860).
Darwin already had one important theory, natural selection, under his belt when he
turned his attention to what initially seemed to be a contradictory theory, sexual selection.
Darwin had an aversive reaction to the peacock’s tail because the mere existence of this
ornament seemed to run counter to predictions of natural selection, the evolution of traits
that enhance survivorship. The peacock’s tail is not an isolated example. Elaborate traits
that seem to compromise survivorship are found throughout the animal kingdom among
most sexually reproducing taxa, and these ornaments are perceived in all sensory modali-
ties. These types of traits include birdsong, cricket and frog calls, firefly flashes, brilliant
colors of many fish, and the musty odors of many mammals. These traits share other attri-
butes: they are involved in mating behavior and they are sexually dimorphic, often being
more extensively developed in males than in females.
In retrospect, Darwin’s sexual selection hypothesis does not seem complex. Like natural
selection, evolution by sexual selection requires variation, selection, and inheritance. The
difference is in the second step. Natural selection favors those variants with higher survi-
vorship while sexual selection favors those variants with greater mating success. Darwin
suggested that sexual selection favored two types of traits: those that enhanced an indi-
vidual’s access to members of the other sex and traits that made courters more attractive
to members of the opposite sex, the choosers; in short, armaments and ornaments. An
important caveat is that there is substantial variation in the animal kingdom, and there
are cases in which females exhibit the more elaborate courtship traits and other species,
such as our own, in which males and females both exhibit these types of traits (Rosenthal
2017). Nevertheless, Darwin posited that in most animal mating systems, males are com-
peting for either access to or the attention of females.
Male-​male competition was not a controversial idea perhaps for several reasons (Richards
2017; Ryan 2021). Males battling for access to females did not seem all that different from
activities favored by natural selection and conformed to Tennyson’s epitaph that nature
was “red in tooth and claw.” The idea of mate choice, however, rankled many of Darwin’s
supporters, especially Alfred Russel Wallace. Many of Darwin’s contemporaries had a hard
time empowering female animals as the agents who decide who gets to mate, and they
also found Darwin’s explanation of why females have these preferences, that they were
grounded in the female’s sexual aesthetics, as rather wanting. Darwin’s theory laid mostly
fallow for about a century (Richards 2017; Ryan 2021).
There was a resurgence of interest in sexual selection by mate choice one hundred
years later that resulted in large part from Campbell’s edited book Sexual Selection and
the Descent of Man, 1871-​1971 (Campbell 1972), and especially Trivers’s (1972) chapter
on parental investment and sexual selection. Trivers was influenced by the early work
of Bateman (1948), who showed that in fruit flies the number of matings had a larger
effect on male reproductive success than on female reproductive success. Trivers pointed
out that this discrepancy arises from greater female investment in her gametes than male
investment in his gametes. This gives rise to a sexual conflict of interest (Arnqvist and
Rowe 2005) in which males should be under selection to mate often and females should
be under selection to mate carefully. A point sometimes overlooked by critics in the social
sciences is that this formulation firmly planted females in the driver’s seat; they were the
deciders. But when Darwin moved on to discuss humans, he returned to a more Victorian
view of the sexes (Richards 2017).
Bateman’s principle, as it is now called, has been productively criticized on a number
of fronts (Gowaty et al. 2012; Tang-​Martínez 2016). A recent meta-​analysis of a large
number of mating systems, however, upheld Bateman’s and Trivers’s prediction of these
sex differences in mating roles (Janicke et al, 2016). In addition, Borgerhoff Mulder and
Ross (2019) critically and carefully applied Bateman’s principle to their analyses of twenty
years of data on marriage systems of the Pimbwe in East Africa.
Darwin’s theory of sexual selection by mate choice has attracted more attention and
consternation than his theory about male competition for access to females. It was much

Sexual Selec tion and t he Animal’ s Mating Mind 17

easier to explain the evolution of armaments than ornaments. Post-​Trivers, behavioral
ecologists showed that in many cases female mate choice garnered resources that increased
her reproductive success. But the question that seemed to most interest behavioral ecolo-
gists was what kind of information about the male’s genetic quality is communicated to
the choosing females. The good gene’s hypothesis posits that males advertise genes that
enhance survivorship, and females are under selection to choose these males because of
the genetic benefits to their offspring (Hamilton and Zuk 1982; Rowe and Houle 1996;
Zahavi 1975). Theory suggests this hypothesis is logical (Grafen 1990), but there is not
overwhelming empirical support that it is biological. Surely, good genes selection occurs,
but estimates of its relative effect in nature are relatively small (Achorn and Rosenthal
2020; Kirkpatrick and Barton 1997; Møller and Alatalo 1999; Rosenthal 2017).
More recently, research in the realm of sensory ecology, neuroethology, and neurosci-
ence have shown how selection on sensory, neural, and cognitive systems in other domains
can have important influences on what females find sexually beautiful. These studies are
providing important insights into the scaffolding of an animal’s sexual aesthetics. This is
the line of research I follow in the remainder of this chapter.

Sexual Selection and Evolutionary Psychology

Evolutionary biologists have long been interested in human biology, and especially
human behavior. Darwin is a prime example (Darwin 1871, 1872). Much of the current
interest in the evolutionary basis of human behavior probably has its roots in Sociobiology
(Wilson 1975). That book, and especially the last chapter, “Man: From Sociobiology to
Sociology,” both garnered the most criticism from the general audience and also provided
the foundation for evolutionary psychology. Many of the subjects in evolutionary psy-
chology (e.g., altruism, mutualism, war, and sex) address ideas that populated that chapter
(Barkow et al. 1992; Buss 1994; Miller 2011). Thus it seems fair to say that evolutionary
biology writ large has inspired much of what is going on today in evolutionary psychology.
I am going to turn the tables. I am going to review a number of recent avenues of
interest in animal sexual selection and mate choice that are inspired by studies of humans,
especially in the fields of psychophysics, behavioral economics, and neuroaesthetics.

Cognitive Aspects of Mate Choice

A number of researchers working parallel to or collaborating with behavioral ecolo-
gists have delved into the neural underpinnings of female preferences. What is it about
the female’s brain that makes her perceive some traits as more attractive than others? In a
variety of taxa and in all sensory modalities, researchers have uncovered neural biases at
the sensory periphery and neural circuits in the brain that contribute to the female’s sexual
aesthetics (e.g., Endler 1992; Gerhardt and Huber 2002; Greenfield 2002; Wilczynski
and Ryan 2010).

18 M i c hael J. Ryan
Another random document with
no related content on Scribd:
Dynastic, dynasties, 434, 446, 457, 500

East Indies, East Indians, 44, 46, 53, 67, 98, 213, 221, 232, 253,
260, 289, 343, 423, 471, 485, 486, 487, 488, 490, 504, 506
Easter Island, 98
Eclipses, 254
Ecuador, 228, 338
Egypt, Egyptian, 30, 55, 96, 104, 113, 141, 142, 173, 202-204,
211, 223, 232, 244, 253, 255, 258, 259, 262, 265-267, 269,
291, 305, 330, 333, 353, 371, 402, 414, 416-419, 423, 425,
431, 433-435, 438, 440-449, 453-459, 468, 479, 496-500,
503-505.
Eighteen Provinces, 461
Elam, Elamite, 441, 449, 450
Elementary Ideas, 195
Elephant, 152, 174, 350
Ellis Landing mound, 321, 323
Empire, 333, 360, 380, 483
Encyclopædia, 468
Endocrine, 66
Endogamy, 481
Eneolithic, 417, 450
England, English, Englishmen, 67, 76, 78, 84, 134, 213, 400,
401, 407, 408, 417
Environment, 326, 502
Eoanthropus, 23
Eocene, 18
Eolithic, 146-148, 444, 446
Ephthalites, 475
Epicanthic fold, 44
Epi-Palæolithic, 409
Eriocomi, 54
Eskimo, 32, 45, 51, 53, 100, 121, 146, 181, 212, 213, 241, 336,
345, 346, 366, 367, 370, 375, 390, 391, 475
Estrangelo, 291
Estufa, 371
Ether, 479
Ethiopian, 49, 52, 472
Ethnography, 6
Ethnographic province, 295
Ethnology, 6
Etruscans, 209, 211, 217, 248, 249, 251, 278, 423, 438, 451
Eubœa, 274
Eugenics, 7
Euphrates, 203, 441, 447, 448, 451-453, 504
Euplocomi, 54
Eurasian, Eur-Asiatic, 53, 253, 327, 431, 500, 504
Euthycomi, 54
Evolution, 7
Evolutionistic anthropology, 9
Examinations, literary, 468, 470, 487
Exogamy, 232-238, 355-360, 490, 492, 493, 500, 501
Eye Color, 40, 106
Faïence, 447
Family, 232;
linguistic, 88, 194, 345;
names, 487
Far East, 423, 424, 474
Fasting, 364
Fashions, 126, 129, 215
Fertile Crescent, 440, 453
Fetish bundle, 368
Feudal, Feudalism, 125, 425, 448, 469, 471;
Kingdom of Egypt, 446
Fibula, see safety-pin
Fiji, 45
Filipinos, 67
Finland, Finns, 95, 427, 474, 476
Finno-Ugric, 95, 110, 474, 475, 476
First Salmon Rite, 304-316
Firearms, 419, 467, 474, 484, 489
Fire, 140, 169, 176, 395, 426
Fire-drill, 218, 349
Fire-worship, 302, 452
Fish, 183
Flake, 160, 164, 176, 395, 398
Flax, 414, 446
Flemish, Fleming, 105, 111
Flood legends, 200
Florida, 385
Focus (of culture), 189, 356, 377, 426, 431, 437, 440, 467, 472,
473, 476
Folk-lore, 198-202
Folkways, 128
Fonts, 282
Font-de-Gaume, 408
Foramen magnum, 26
Fossil, 137
Fowl, 414, 486, 489-491, 498, 499, 501
France, see French
Franciscan, 333
Frank, 104
French, France, 43, 117, 121, 136, 220, 250, 253, 276, 395,
398, 400, 402, 405, 407, 408, 418, 424, 426, 429, 432, 506
Fricative sounds, 92
Frija, 256
Frontal angle, 33
Fuegian, 469
Fuyu, 470

Gables, 502
Gabrielino, 188, 190, 310, 311, 320
Gafsa, 400
Galley Hill, 29, 32
Galton, 83
Gamma, 270, 275
Ganges river, 479, 480
Ganggraeber, 430
Gaul, 105, 305, 425, 465
Gender, sex, 119
Genealogy, 491, 505
Generation, 57
Genetic classification, 88, 103
Genius, 71, 83, 273
Gens, 232
German, Germany, 43, 104, 117, 118, 135, 398, 421, 424, 427,
432, 464, 472
Germanic, 95, 124, 221, 419, 425, 460, 473
Ghost-dance, 334
Gibbon, 13
Gibraltar, 24, 32, 398, 404
Gideon, 456
Gimel, 270
Girls’ Rite, 300-316, 365
Glabella, 31, 33
Glaciation, Glacial period, 18, 23, 149, 350, 444
Glass, 447
Goat, 44, 415, 441, 451, 463, 473
Gold, 373, 374, 416, 421, 479
Gorilla, 13, 21, 22, 26, 27, 32, 64
Goths, Gothic, 104, 220, 251, 284;
architecture, 250
Græco-Bactrian, 484
Grain, 379, 446, 462, 463, 473, 503
Grammar, 482
Great Basin, 236, 296, 336
Greece, Greek, 93, 95, 103, 111, 113, 126, 129, 204, 210, 211,
220, 226, 244, 253, 265, 269, 270-273, 346, 359, 395, 419,
421, 424, 431, 432, 447, 454, 455, 456, 457, 459, 472, 480,
483, 484, 506
Greenland, 150
Gregorian calendar, 377
Grenelle, 30
Grimaldi, 27-29, 34, 48, 155, 157, 344, 395, 404, 497
Grimm’s law, 93
Gros Ventre, 236
Ground painting, 310
Ground stone, 142, 144, 410, 444, 492, 494, 495
Gschnitz, 406
Guatemala, 185, 223, 352, 362
Guiana, 339, 383
Guinea, 502
Gunpowder, 426, 467, 474
Günz, 18, 21, 150
Gypsy, 56

Habit, 275, 283

Hadrian, tomb of, 249
Haeckel, 55
Hafting, 168, 176
Haida, 295, 356
Hairiness, 39, 62
Hair texture, 39, 41, 45, 62
Half-breeds, 81
Half-hitch coiling, 329
Hallstadt, 424, 425, 460
Hamites, Hamitic, 96, 113, 119, 120, 448, 450, 472, 497
Hammock, 361, 381, 382
Hammurabi, 451, 458
Han, 463, 465-468, 470
Harpoon, 165, 167, 348, 349, 389, 390, 396, 406, 408, 411, 426,
428, 429, 492
Haruspicy, 209, 210
Harvey, 125
Hawaii, 69, 73
Hawaiki, 491
Head hunting, 489, 490
Hebrew, Hebrews, 96, 103, 201, 207, 211, 253, 265, 269, 282,
285, 286, 451, 457, 458, 472, 473, 505.
See also Jew
Heddle, 222, 361
Hellenism, Hellenistic, 225, 255, 484
Hellespont, 453
Hemp, 415, 416, 466
Hepatoscopy, see Liver divination
Heraldry, 203
Herd instinct, 59, 128, 277
Heredity, 34, 72, 80, 239
Hermes, 256
Herodotus, 9
Hesi Dance, 309
Hiaksai, 470
Hieratic, 266, 449, 454
Hieroglyphic writing, 266, 443, 449, 454, 456, 458
Himyarites, 287
Hindi, 221
Hindu, 39, 41, 42, 44, 111, 126, 210, 224, 231, 239, 247, 260,
287, 288, 289, 346, 423, 472, 481, 482, 484, 490, 504, 505,
506;
Hinduism, 476, 480, 493.
See also India
Hippopotamus, 152
History, 482
Hittites, 202, 223, 268, 269, 422, 423, 442, 451, 453, 458, 506
Hokan, 121
Homer, 210, 278, 459, 479
Homo Heidelbergensis, 22;
Mousteriensis, 24;
Neandertalensis, 24, 27, 29;
primigenius, 24;
sapiens, 27, 29, 34, 155, 395
Homonyms, 223
Honan, 464, 466
Hongkong, 68, 69
Hopi, 135, 181, 187, 236, 252
Hoplites, 129
Horse, 152, 350, 384, 387, 414, 426, 433, 448, 455, 458, 462,
463, 473, 479, 497, 498
Hottentot, 45, 52, 54, 71, 96, 120, 121, 145, 502
Hour, 207, 225
Hrdlička, 65
Huastec, 135
Huichol, 203
Hun, 462, 465, 475
Hundsteig, 157, 412
Hungary, Hungarian, 69, 87, 95, 110, 424, 431, 432, 474, 476.
See also Magyar
Hupa, 313, 320
Huxley, 55
Hyksos, 104, 446, 448, 451, 458

Iberian, 43, 432, 451

Ibero-insular, 53
Ideograms, ideographic writing, 223, 224, 263, 291, 329, 449
Independent Evolutions, 260.
See also Parallelism
Igorot, 372
Ikhnaton, 448, 455
Iliad, 422
Illyrian, 460
Imitation, 216, 239, 326, 327, 468
Inca, 134, 242, 371, 378, 380, 382
Incorporating languages, 100, 102, 104, 121
India, 44, 46, 52, 53, 95, 96, 105, 202, 204, 210, 211, 223, 251,
258, 269, 287, 290, 353, 371, 406, 419, 423, 426, 452, 454,
459, 462, 463, 466, 467, 469, 472-486, 488-490, 493, 499,
503, 504.
See also Hindu
Indian Ocean, 45, 49
Indic, 135, 452
Indo-Afghan, 53
Indo-Bactrian, 287
Indo-Australian, Indo-Australoid, 44-46, 55, 476, 477, 486, 488
Indo-China, 46, 234, 260, 358, 463, 469, 485, 486, 489
Indo-European, 95, 96, 100, 111, 113, 119, 120, 121, 124, 125,
135, 220, 221, 286, 346, 450-453, 457, 459, 460, 473, 477-
479
Indo-Germanic, 95
Indo-Iranian, Indo-Iranic, 221, 479
Indo-Melanesian, 493
Indo-Oceanic, 478, 489, 502
Indonesia, Indonesian, 44, 53, 487
Indus River, 479, 480
Inflecting languages, 100, 102, 220
Inion, 31
Initiation, 363, 364, 389, 438
Inter-continental distribution of races, 49
Intelligence tests, 75
Interglacial periods, 18, 23, 250, 398
Invention, 58, 142, 166, 167, 168, 176, 179, 182, 185, 186, 191,
197, 216, 239, 264, 268, 269, 271, 273, 286, 311, 327, 328,
 353, 371, 376, 398, 418, 431, 438, 459, 461, 467, 482, 493
Iran, Iranian, Iranic, 95, 450-452, 479
Ireland, 246, 420, 432
Iron, 332, 373, 419, 421, 422, 426, 430, 433, 445, 447, 455,
458, 469, 479, 488, 490, 496, 497, 498, 501, 502, 505
Iron Age, 142, 146, 394, 408, 415, 419-426, 431, 446, 450, 456,
470
Iroquois, 100, 121, 347, 356, 359, 386
Irrationality, 277
Irula, 41, 46, 476, 486
Ishtar, 256
Islam, 253, 332, 455
Israel, 456
Isolating languages, 100, 102, 124
Isolation, 182, 195, 197, 383, 492, 501, 505
Isthmus, 440, 441
Italy, Italian, 76-79, 209-217, 247, 250, 253, 274, 400-402, 404,
417-419, 423, 424, 432, 497, 506
Ivory, 166, 172, 175

Jackal, 348, 429

Jade, 466, 473
Jahveh, 455
Japan, Japanese, 24, 35, 39, 40, 65, 69, 97, 100, 107, 113, 119,
204, 210, 224, 259, 343, 419, 424, 470-472, 475, 485, 504
Java, 19, 258, 289, 489, 491
Javelin, 165
Jersey, 24
Jerusalem, 453
Jesuit, 204
Jesus, 255, 257, 454
Jeu-di, 256
Jew, Jewish, 44, 53, 57, 127, 183, 257, 258, 423, 454, 484.
See also Hebrew
Jim Crow, 58
Jimsonweed, 306-316
Johannesburg, 67
Jumping Dance, 312, 313
Jupiter, 254, 256, 258

Kalahari, 501
Kamchadal, 475
Kaph, 274
Kapila, 480
Kappa, 275
Kardouchoi, 452
Karma, 482
Kassites, 451, 455, 458
Katun, 376
Kayak, 391
Kelts, Keltic, 95, 104, 105, 419, 424, 460, 472
Keres, 187
Kesslerloch, 157, 177
Keystone, 247
Kharoshthi, 287
Khmer, 485
Khorasan, 417
Kings, Kingship, 441, 446, 478, 488, 491, 499, 500, 502, 505
Kiowa, 294
Kitchenmiddens, 412, 429, 430, 435
Kiva, 371
Kjökkenmöddings, 429
Klamath, 115
Knossos, 456
Kokorai, 470
Kolarian, 41, 46, 98, 105, 477, 478, 486
Koph, 274, 275
Koppa, 275
Korai, 470
Korea, Korean, 96, 204, 291, 292, 424, 468-471
Koryak, 210, 475
Kossæans, 451
Krapina, 24, 32, 154
Kris, 419
Kuksu Cult, 306-316
Kurds, 53, 452
Kwakiutl, 295, 356
Kyoto, 69
La Ferrassie, 24, 157
Lake-dwellings, 434, 463, 473
La Madeleine, 153, 175.
See also Magdalenian
La Mairie, 176
Lamp, 389, 390, 396
Language, 57
Laotse, 464
La Plata, 338, 339
Lapp, 53, 475, 476
La Quina, 24
La Tène, 424, 425, 460
Latin, Latins, 76, 95, 103-105, 111, 113, 119, 124, 126, 132, 136,
274, 278, 346
Latitude, geographical, 68
Laugerie Basse Haute, 27
Laurel-leaf blade, 162
Law, phonetic, 92;
scientific, 324
Lead, 374
League of Five Nations, 356
Least common multiples, 226
Leaven, 463
Le Moustier, 24, 32, 153-179.
See also Mousterian
Lentils, 414
Ligurians, 459
Limiting conditions, 226, 335
Linguistic family, see family, linguistic
Linnæus, 49
Lintel, 243, 244
Lissotrichi, 54, 55
Lithuanian, 95
Litorina litorea, 428;
period, 428, 430
Littoral, 53
Liver divination, 209, 210, 217, 248, 438
Llama, 342, 350, 361, 380
Lolo, 95, 210, 468, 469
London, 29
Loom, 222, 332, 360-362, 379, 414
Lophocomi, 54
Los Angeles, 310
Lotus, 244
Louisiana, 2
Lourdes, 174
Lucretius, 9
Luiseño, 188, 310, 320
Lunation, 374, 375, 376
Lybians, 472
Lydia, 451, 455

Macedonia, Macedonian, 104, 129, 451, 484

Mackenzie (Yukon) area, 295, 336, 388, 389, 391
Madagascar, 98, 488, 500
Magdalenian, 27, 28, 146, 153-179, 348, 390, 395, 396, 400,
402, 404-408, 411, 412, 427, 433, 496, 502
Magellan, Strait of, 351
Magic, 200, 219, 232, 254, 491, 494
Magic Flight, 198, 201, 218, 391
Magism, 452
Maglemose, 396, 408, 410, 412, 428, 429
Magnetic needle, 467
Magyar, 95, 110, 474.
See also Hungarian
Maidu, 121, 307, 309
Maize, 185, 218, 237, 341, 353, 379, 382, 467, 468, 490
Malay, 52, 54, 488, 489
Malayan, 49, 149
Malayo-Polynesian, 98, 100, 119, 121, 485, 487
Malay Peninsula, 45, 98, 258, 290, 486, 487
Malaysian, Malaysia, 41, 44, 46, 136, 209, 222, 223, 257, 289,
423, 463, 469, 471, 472, 486-490, 500
Mammoth, 151, 152, 174, 175
Manchu, 95, 291, 454, 465, 466, 474, 476
Maneh, 207
Mangyan, 290
Manila, 68, 69, 290
Manioc, 382
Marduk, 256
Marginal areas, cultures, 335, 383, 388, 437, 440, 469, 473,
475, 478, 503
Mars, 254, 255
Marseilles, 424
Mas d’Azil, 157, 406
Masks, 187, 294, 356, 363, 366, 369, 502
Masonry, 370, 371, 380, 385, 418, 426, 427, 430, 447, 498
Mathematics, 482
Matrilinear descent, 232-238, 331, 355-360, 490, 493, 500
Mauer, 22
Mauretanians, 496
Maya, 100, 105, 113, 116, 135, 197, 205, 206, 223, 225, 226,
228, 230-232, 239, 246, 261, 262, 266, 268, 333, 338, 347,
349, 358, 362, 368, 369, 371, 372, 376-378, 438
Measles, 66
Measures, 454
Medes, 452
Medicine-man, see Shaman
Mediterranean (race), 41-43, 55, 77, 82, 457, 505, 506
Mediterranean Sea, area, 43, 49, 81, 120, 222, 250, 287, 335,
398, 402, 404, 419, 425, 430, 435, 454, 474, 476, 485, 499,
504
Megalith, 426, 433, 460, 471, 496, 497
Melanesia, Melanesian, 41, 44, 52, 55, 98, 227, 232, 234, 236,
487, 489, 490, 493, 502, 504
Melanochroid, 53, 55
Mena, 434, 435
Mencius, 464
Menhir, 416
Mentone, 27, 29
Memphis, 70
Mercre-di, 256
Mercury, 254-256
Mesha, 269
Mesolithic, 396, 409, 410
Mesocephalic, 37
Metal Age, 141, 149
Metallurgy, 332, 341, 373
Mesopotamia, 202, 215, 247, 250, 440, 441, 451-453, 473
Mesozoic, 15
Mexico, Mexican, 65, 105, 185, 203, 228, 229, 236, 244, 251,
260, 261, 290, 295, 310, 329, 332, 338, 340, 342, 351, 356,
357, 361, 369, 370, 372, 374, 379, 384, 385, 387, 431, 440,
442, 504
Mexico, Gulf of, 369
Microliths, 406, 407, 428
Micronesia, Micronesian, 98, 487, 490
Middle America, 205, 206, 213, 340-342, 352-358, 361, 363,
367, 368, 370, 371, 373, 381, 383, 385, 388, 391, 440-442
Middle Kingdom, 446, 451
Middle Stone Age, 396, 409
Midianite, 456
Migration, 195, 214, 228, 461, 472, 491
Milan, 250
Milk, 463, 468
Millet, 414, 446, 463, 502
Milton, 115
Mina, 204
Mindel, 18, 150
Mindoro, 290
Ming, 468
Minoan, 423, 456-459, 479, 480
Minos, 456
Minuscules, 281
Minusinsk, 462
Miocene, 18, 148
Mission style, 251
Missing link, 11
Missionaries, 204, 333
Mississippi River, 294, 340, 385;
valley, 386
Mit, 259
Mitanni, 451, 458
Mithra, Mithraism, 258, 259
Miwok, 236, 307
Moab, 269
Mogul, 251, 475
Mohammedan, Mohammedanism, 96, 198, 223, 251, 258, 259,
290, 451, 463, 475, 476, 483, 484, 488-490, 497, 500, 504,
506
Mohave, 188, 190, 236, 311
Moi, 41, 46, 486
Moiety, 232-238, 355, 360, 490, 492, 493, 494
Mon, 485
Mon-Khmer, 98
Money, 480
Mongol, 95, 210, 214, 251, 291, 343, 424, 454, 462, 465, 466,
468, 473, 474, 475, 484, 485, 487
Mongolia, 204, 462, 485
Mongolian, 30, 35, 44, 54, 343, 476, 477, 485
Mongoloid, 39, 41, 42, 44, 46, 49, 52, 53, 55, 62, 343, 475, 486-
488, 504, 505
Monolatry, 455
Monosyllabism, 124
Monotheism, 448, 455
Moor, Moorish, 250, 496
Moravia, 24, 29, 432
Mores, 128
Morocco, Moroccans, 211, 448, 496
Mortillet, Gabriel de, 153
Mosaic law, 184
Mother goddess, 455
Mother tongue, 94, 96
Mound Builders, 212, 373, 386
Mourning Anniversary, 303-316
Mousterian, 23, 25, 45, 153-179, 395, 398, 400, 405, 406, 427,
433, 444
Mouth, 374, 376
Mulatto, 80