History of Evolutionary Thought Since 1930: February 2019

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History of Evolutionary Thought Since 1930
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DOI: 10.1093/obo/9780199941728-0118
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History of Evolutionary Thought Since 1930

Roberta Millstein, Michael Dietrich, Robert A. Skipper
LAST MODIFIED: 27 FEBRUARY 2019

DOI: 10.1093/OBO/9780199941728-0118
Introduction
Evolutionary biology underwent several significant transformations in the period after 1930. While mid-century was dominated by
the evolutionary synthesis and the professionalization of evolutionary biology, the second half of the 20th century saw
evolutionary research diversifying and the domain of evolutionary phenomena expanded, especially in response to the rise of
molecular biology. While what happened in 1950 may seem ancient to a contemporary evolutionary biologist, to a historian of
biology the last half of the 20th century is recent history. Where the literature on Darwin could fill a bibliography by itself, the
history of modern evolutionary biology is neither as extensive nor as comprehensive. Some topics such as the evolutionary
synthesis and molecular evolution have a rich historical literature. Others such as cultural evolution or quantitative genetics still
await further historical analysis: not because they are undeserving, but because they are just now becoming ripe for historical
investigation. For the purposes of this article, we selected significant pieces of historical and some philosophical scholarship that
address major developments in the history of evolutionary biology. We did not select so-called classic papers by evolutionary
biologists or review papers written by biologists. Instead our focus was on evolutionary biology as it has been contextualized and
discussed by historians who have often been seeking to address a range of non-biological issues regarding the nature and
practice of science, and how different aspects of evolutionary biology reflect the time and place of their development.
General Overviews
Only a few works of historical analysis are either comprehensive in their scope or develop themes that are central to all of
evolutionary biology after 1930. Smocovitis 1996 offers an important overview of the history of the evolutionary synthesis period
at mid-century and its continuing impact. Bowler 2009 and Gayon 1998 both begin in the Darwinian era and extend into the
mid-20th century. The collections from Ruse and Travis 2009 and Ruse 2013 cover a wider array of topics in evolutionary biology
from Darwin to the present with entries by historians, philosophers, and biologists.
Bowler, Peter. 2009. Evolution: The history of an idea. Berkeley: Univ. of California Press.
First published in 1989, this new edition provides an accessible overview of evolutionary thought from the 19th century to the end
of the 20th.
Gayon, Jean. 1998. Darwinism’s struggle for survival: Heredity and the hypothesis of natural selection. Cambridge, UK:
Cambridge Univ. Press.
Gayon provides a focused narrative that traces the history of selectionist explanations from Darwin through the Mendelian-
Biometrician controversies to the evolutionary synthesis and the challenge of the neutral theory of molecular evolution.
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Ruse, Michael, ed. 2013. The Cambridge encyclopedia of Darwin and evolutionary thought. Cambridge, UK: Cambridge
Univ. Press.
This historically structured volume provides a detailed and informative survey of the history of evolutionary thought from ancient to
modern times.
Ruse, Michael, and Joe Travis, eds. 2009. Evolution: The first four billion years. Cambridge, MA: Harvard Univ. Press.
This comprehensive introduction to evolutionary biology includes essays from historians, philosophers, and biologists on the
many different dimensions of the study of evolutionary biology and its wider impact.
Smocovitis, V. B. 1996. Unifying biology: The evolutionary synthesis and evolutionary biology. Princeton, NJ: Princeton
Univ. Press.
The evolutionary synthesis continues to be an enigmatic historical entity. Unifying Biology offers an important analysis of the
multifaceted nature of the synthesis and its impact on modern evolutionary thought.
Journals
There are a number of journals that publish articles addressing the history of evolutionary biology. The Journal of the History of
Biology focuses almost exclusively on the history of biology, while History and Philosophy of the Life Sciences and Studies in
History and Philosophy of Science Part C regularly include articles written from a more integrated philosophical and historical, and
sometimes sociological perspective. Several history of science journals will occasionally include articles on the history of
evolutionary thought including Isis, Historical Studies in the Natural Sciences, Centaurus, the British Journal for the History of
Science, and the more interdisciplinary Perspectives on Science.
British Journal for the History of Science.

Published quarterly, this journal is sponsored by the British Society for the History of Science. Its scope includes the entire history
of science.
Centaurus.
Published quarterly, this journal is the official journal of the European Society for the History of Science. Its scope includes the
entire history of science.
History and Philosophy of the Life Sciences.

Published quarterly, this interdisciplinary journal offers articles from historical, philosophical, and interdisciplinary perspectives. It
is sponsored by the Naples Zoological Station.
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Historical Studies in the Natural Sciences.

Published five times per year, this journal includes articles covering all aspects of the history of science.
Isis.
Published quarterly, Isis is the official journal of the History of Science Society. Its scope includes the entire history of science.
Journal of the History of Biology.

Published quarterly, JHB concentrates on the history of 19th-, 20th-, and 21st-century biology.
Perspectives on Science.
Published quarterly, this journal publishes interdisciplinary science studies articles that combine perspectives from history,
philosophy, and sociology.
Studies in History and Philosophy of Science Part C: Studies in History and Philosophy of Biological and Biomedical
Sciences.
Published quarterly, Studies C includes a mix of articles focusing on the history, philosophy, and sociology of biology.
Topic Areas
We have organized this article by topic areas with evolutionary biology. The entries within each topic area address significant
contribution to the history of that particular topic. Our topic list is not exhaustive and neither are the titles within each topic. In
constructing this list, we selected topics that we believe have been the subject of extensive historical discussion.
Chance and Contingency
Chance has been recognized as a part of evolutionary biology since Darwin. Ramsey and Pence’s anthology contains a number
of essays detailing various understandings of the role that chance and contingency play in evolution. The history of the idea that
mutations and genetic variations are chance occurrences is covered by Merlin 2010. Millstein 2011 surveys the different
meanings of chance in evolutionary biology and develops an account of what the different meanings have in common. Thinking
about contingency is typically traced to Stephen Jay Gould’s idea of “replaying the tape of life” with different results on each
replay. This idea was developed more fully by Beatty 1995 and framed in terms of historicity by Desjardins 2011. (See also Drift
entry for discussions of chance in evolution).
Beatty, John. 1995. The evolutionary contingency thesis. In Concepts, theories, and rationality in the biological
sciences. Edited by Gereon Wolters and James G. Lennox. The Second Pittsburgh-Konstanz Colloquium in the
Philosophy of Science. Pittsburgh: Univ. of Pittsburgh Press.
Beatty characterizes Gould’s contingency thesis into weak and strong forms and shows how both forms follow from the nature of
the evolutionary process. As a result of this inevitable contingency, Beatty argues, there are no laws in biology; instead, there is
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widespread pluralism and arguments over the relative significance of different generalizations.
Desjardins, Eric. 2011. Historicity and experimental evolution. Biology and Philosophy 26.3: 339–364.
Based on an analysis of long-term experimental evolution research, Desjardins offers criteria and conditions for the historicity of
evolution. In doing so he explains the relationship between historicity and historical contingency in evolution.
Merlin, Francesca. 2010. Evolutionary chance mutation: A defense of the modern synthesis’ consensus view.
Philosophy and Theory in Biology 2:e103.
Merlin argues for a modified version of “evolutionary chance” defended by Millstein 2011 and introduces a number of important
distinctions to help sort through the various ways in which a chance mutation can occur. The enhanced concept is used to
illuminate mutator mechanisms and other recent findings; understood properly, Merlin argues, these new findings say nothing to
contradict the synthesis view of what constitutes a chance mutation.
Millstein, Roberta L. 2011. Chances and causes in evolutionary biology: How many chances become one chance. In
Causality in the sciences. Edited by P. McKay Illari, F. Russo, and J. Williamson, 425–444. Oxford: Oxford Univ. Press.
Millstein characterizes seven meanings of “chance” in evolutionary biology and argues that there is a unified concept of chance
underlying these seven, dubbed the Unified Chance Concept (UCC). The chapter argues that each is characterized by which
causes are considered, ignored, or prohibited, and thus that chance in evolutionary biology can only be understood through
understanding causes at work.
Ramsey, Grant, and Charles H. Pence, eds. 2016. Chance in evolution. Chicago: Univ. of Chicago Press.
This multi-authored volume edited by Ramsey and Pence explores chance in evolution from philosophical, historical, religious,
and biological perspectives, providing the reader with analyses of the various meanings of chance from the past through the
present and examinations of how those meanings have been applied (rightly or wrongly, well or badly) to evolutionary
phenomena. The reader is also given insight into the empirical debates over the presence of chance in evolution and the authors’
views on where those debates stand today.
Drift
The topic of random genetic drift has been controversial almost from the outset; Provine 1986 documents the legendary
disagreements between Sewall Wright and R. A. Fisher over drift. Beatty 1992 surveys the history and changing meanings of the
term, as does Plutynski 2007. Provine 1986 and Beatty 1987 focus on the changing views concerning drift of two biologists in
particular (Theodosius Dobzhansky and Wright, respectively), whereas Dietrich 2006 explores the use of drift in molecular
evolution. Beatty 1984 shows the challenges of distinguishing drift from selection. Millstein, et al. 2009 shows how our
understanding of drift should derive from the phenomena that biologists such as Wright and Fisher sought to model. Millstein
2017 connects key figures in the development of the drift concept to contemporary debates more broadly.
Beatty, John. 1984. Chance and natural selection. Philosophy of Science 51:183–211.
Beatty traces the use of chance in evolutionary theory from Darwin through Fisher, Dobzhansky, and Wright, through the neutral
theory: argues that it is often difficult to distinguish chance (and drift in particular) from natural selection.
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Beatty, John. 1987. Dobzhansky and drift: Facts, values, and chance in evolutionary biology. In The probabilistic
revolution. Vol. 2. Edited by L. Kruger, G. Gigerenzer, and M. Morgan, 271–311. Cambridge, MA: MIT Press.
Beatty traces Dobzhansky’s changing views on the importance of drift, and his shift at mid-century toward a more panselectionist
viewpoint.
Beatty, John. 1992. Random drift. In Keywords in evolutionary biology. Edited by E. F. Keller and Elizabeth Lloyd,
273–281. Cambridge, MA: Harvard Univ. Press.
This is a historical overview of the concept of drift, beginning with the first origins of the idea with the Hagedoorns and Gulick.
Dietrich, Michael. 2006a. Three perspectives on neutrality and drift in molecular evolution. Philosophy of Science
73:666–677.
Dietrich analyzes three uses of neutrality and drift in molecular evolution. In the first, neutrality is assumed as a simplest case for
modeling. In the second and third, concepts of drift and neutrality are developed within the context of population genetics testing
and the development and application of the molecular clock.
Millstein, Roberta L. 2017. Genetic drift. In The Stanford encyclopedia of philosophy. Edited by Edward N. Zalta.
Stanford, CA: Stanford Univ. Press.
Millstein traces the concept of genetic drift from John Gulick and A. C. and A. L. Hagedoorn to R.A. Fisher and Sewall Wright to
Motoo Kimura and Tomoko Ohta to contemporary debates in biology and philosophy of biology.
Millstein, Roberta, Robert Skipper Jr., and Michael Dietrich. 2009. (Mis)interpreting mathematical models: Drift as a
physical process. Philosophy & Theory in Biology 1.
Millstein, Skipper, and Dietrich argue that drift cannot be understood solely through an inspection of the mathematical models;
rather, one must understand the phenomenon—an indiscriminate sampling process—which biologists such as Fisher and Wright
sought to model.
Plutynski, Anya. 2007. Drift: A historical and conceptual overview. Biological Theory 2.2: 156–167.
In this historical survey, Plutynski claims that drift is a heterogeneous concept that collectively refers to a variety of different ways
in which chance, broadly understood, affects evolving populations over time. She maintains that over the history of the use of this
concept, the term “drift” has been used to refer to a variety of outcomes and causes.
Provine, William. 1986. Sewall Wright and evolutionary biology. Chicago: Univ. of Chicago.
Provine traces the history of evolutionary theory, paying special attention to Sewall Wright’s influence on it. This biography
touches on numerous topics in evolutionary genetics, natural selection, evolutionary ecology, and drift, especially in relation to the
debates between Wright and R. A. Fisher.
Ecological Genetics
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E. B. Ford, considered the founder of the Oxford School of Ecological Genetics and the field of ecological genetics more
generally, emphasized that ecological genetics incorporates evolutionary fieldwork attentive to the ecology of habitats as well as
laboratory genetics. Ecological genetics tends to be selectionist and adaptationist in orientation. In Cain and Provine 1992, Cain
gives a firsthand account of this approach, while Provine provides historical context; see also Provine 1986. Millstein 2008 and
Millstein 2009 examine what Provine has termed “The Great Snail Debate” over the role of selection and drift in the evolution of
the land snail Cepaea nemoralis, focusing on the debates that Lamotte had with Cain and Sheppard in the 1950s. Rudge 2005a
and Rudge 2005b focus on another classic ecological genetics study of the peppered moth, Biston betularia.
Cain, A. J., and William Provine. 1992. Genes and ecology in history. In Genes in ecology. Edited by R. J. Berry, T. J.
Crawford, and G. M. Hewitt, 3–28. Oxford: Blackwell Scientific.
This essay, although technically co-authored, contains sections written by Cain and sections written by Provine. Cain provides a
firsthand account of his experiences and perspective on ecological genetics, while Provine argues for the rise of the selectionist
hypothesis as null hypothesis in the 1950s and 1960s, only to be followed by the neutral theory as a null hypothesis.
Millstein, Roberta L. 2008. Distinguishing drift and selection empirically: ‘The Great Snail Debate’ of the 1950s. Journal
of the History of Biology 41:339–367.
Analyzes the evidence provided in the dispute over Cepaea nemoralis between Cain and Sheppard on the one hand and Lamotte
on the other, arguing that Lamotte did in fact provide good evidence for a significant role for drift in addition to that played by
selection.
Millstein, Roberta L. 2009. Concepts of drift and selection in ‘The Great Snail Debate’ of the 1950s and early 1960s. In
Descended from Darwin: Insights into American evolutionary studies, 1925–1950. Edited by J. Cain and M. Ruse,
271–298. Philadelphia: American Philosophical Association Press.
Millstein shows how some of the dispute over Cepaea nemoralis between Lamotte and the team of Cain and Sheppard lay in
understandings of drift that were sometimes ambiguous and unclear, particularly with respect to the connection between drift and
nonadaptiveness.
Rudge, David. 2005a. The beauty of Kettlewell’s classic experimental demonstration of natural selection. BioScience
55.4: 369–375.
Rudge shows that we see Kettlewell’s classic industrial melanism experiments as “beautiful” because of the simplified way they
are often portrayed, with the complex details left out. The more complex story, however, does not cast doubt on the claim that
Kettlewell demonstrated natural selection (especially in light of subsequent confirmations) or justify their removal from textbooks.
Rudge, David. 2005b. Did Kettlewell commit fraud? Re-examining the evidence. Public Understanding of Science 14.3:
249–268.
Did Kettlewell commit fraud in the classic studies of industrial melanism, as Judith Hooper has charged? Through a critical
examination of Hooper’s evidence, Rudge argues persuasively that he did not.
Evolutionary Developmental Biology
Although a relatively recent field, evolutionary developmental biology has attracted the attention of a number of historians seeking
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historical precedents for the intersection of evolution and development. Amundson 2005 offers the most comprehensive single
narrative of the effort to map the changing intersection of evolution and development. The essays in Laubichler and Maienschein
2007 provide more detailed analyses of relevant historical figures, such as Haeckel, Goldschmidt, and Bateson, as well as essays
on important areas of research such as morphology and developmental genetics. Love and Raff 2003 provides a broad historical
account that informs contemporary evo-devo. Davis, et al. 2009 explores the assimilation of developmental biology into
evolutionary biology through the study of homeotic mutations. Gilbert 2000 traces Waddington’s influence on the Evolutionary
Synthesis, while Gilbert 2003 casts a wider net for precursors to contemporary evo-devo. Griesemer 2013 discusses an
integrated approach drawing in multiple fields that have informed evo-devo research done by David Wake.
Amundson, Ronald. 2005. The changing role of the embryo in evolutionary thought: Roots of evo-devo. Cambridge, UK:
Amundson explores the history of the relationship between evolution and development through the lens of evo-devo. Amundson
argues that extant histories were written in a way that vindicates neo-Darwinism and sets development aside.
Davis, Gregory, Michael Dietrich, and David Jacobs. 2009. Homeotic mutants and the assimilation of developmental
genetics into the evolutionary synthesis. In Descended from Darwin: Insights into American evolutionary studies,
1900–1970. Edited by Joe Cain and Michael Ruse, 133–154. Philadelphia, PA: American Philosophical Association Press.
Davis, Dietrich, and Jacobs argue that developmental biology was not left out of the evolutionary synthesis. Focusing on research
on homeotic mutants, they describe how developmental genetics was assimilated into evolutionary biology.
Gilbert, Scott. 2000. Diachronic biology meets evo-devo: C. H. Waddington’s approach to evolutionary developmental
biology. American Zoologist 40:729–737.
An important analysis of Conrad Waddington’s historical impact on the integration of evolution, development, and genetics at mid-
century. Gilbert discusses Waddington’s influential contributions, such as canalization and the epigenetic landscape.
Gilbert, Scott. 2003. The morphogenesis of evolutionary developmental biology. International Journal of Developmental
Biology 47:467–477.
In this essay Gilbert traces the historical impact of evolutionary morphology and genetics on the integration of evo-devo but
argues that ecology and medicine must also be appreciated as important influences on contemporary practice.
Griesemer, J. 2013. Integration of approaches in David Wake’s model-taxon research platform for evolutionary
morphology. Studies in History and Philosophy of the Biological and Biomedical Sciences 44:525–536.
Griesemer examines the work of David Wake, describing Wake’s “integration of approaches” as an intersection of fields such as
comparative anatomy, functional morphology, systematics, and evolutionary biology, helping to define the emerging field of
evolutionary developmental biology.
Laubichler, Manfred, and Jane Maienschein, eds. 2007. From embryology to evo-devo: A history of developmental
evolution. Cambridge, MA: MIT Press.
An important set of essays on the interplay between evolution and development ranging from 19th-century studies of body plans
to contemporary evo-devo based on molecular foundations.
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Love, Allan, and R. A. Raff. 2003. Knowing your ancestors: Themes in the history of evo-devo. Evolution & Development
5:327–330.
Love and Raff argue for a broader historical understanding of developmental biology that would in turn inform a broader
conception of contemporary practice in evo-devo.
Evolution of Sex
The evolution of sexual reproduction has been the subject of many different biological explanations. Fehr 2006 explores the
plurality of models use to explain the origin and maintenance of sexual reproduction. Mooney 1995 provides a detailed history
and analysis of many of the genetic models from the mid-20th century, while Segestråle 2013 focuses on the work of W. D.
Hamilton and his broader perspective in terms of host parasite relationships. Ridley 1995 offers a popular survey of the wide
range of research on this topic.
Fehr, Carla. 2006. Explanations of the evolution of sex: A plurality of local mechanisms. In Scientific pluralism,
Minnesota studies in the philosophy of science. Edited by Stephen H. Kellert, Helen E. Longino, C. Kenneth Waters,
167–189. Minneapolis: Univ. of Minnesota Press.
Fehr characterizes three explanations for the evolution of sex: Red Queen, Muller’s Ratchet, and DNA Repair. She argues for a
pluralistic approach to understanding the evolution of sex, with different processes existing at different levels of organization.
Mooney, Susan. 1995. H. J. Muller and R. A. Fisher on the evolutionary significance of sex, Journal of the History of
Biology 28:133–149.
Mooney provides a history of the origins and development of theories of the evolution of sex. Along the way, she clarifies and
organizes the host of questions concerning the evolution of sex.
Ridley, Matt. 1995. The Red Queen: Sex and the evolution of human nature. New York: Penguin.
Ridley surveys theoretical work in evolutionary biology, anthropology, ecology, sociology, and molecular biology. Explores the
question, “Why did sex evolve?”
Segestråle, Ullica. 2013. Nature’s oracle: The life and work of W. D. Hamilton. Oxford: Oxford Univ. Press.
A biography of English biologist W. D. Hamilton. Includes a detailed history of his work on the evolution of sex as well as his work
on inclusive fitness and cooperation in nature.
Evolutionary Ecology and Population Biology
These areas attempt to show how our theories and fieldwork can use integrated models of evolutionary and ecological processes.
Anderson 2013, Dritschilo 2008, and Odenbaugh 2013 concentrate on the work of particular scientists (David Lack, Daniel
Simberloff, and Robert MacArthur, respectively). Collins 1986 focuses on work in the 1950s and 1960s. Mitman 1992 and
Odenbaugh 2006 both examine collaborative studies, with Mitman focusing on work at the University of Chicago and Odenbaugh
analyzing the efforts of Levins and colleagues. Hagen 1992 explores ecosystem ecology and its split from evolutionary biology. A
short history of evolutionary ecology since the 1960s is provided in Futuyma 2016.
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Anderson, Ted. 2013. The life of David Lack: The father of evolutionary ecology. Oxford: Oxford Univ. Press.
A biography of ornithologist David Lack with extensive discussion of his work on Darwin’s finches and the evolutionary ecology of
bird populations.
Collins, James P. 1986. Evolutionary ecology and the use of natural selection in ecological theory. Journal of the History
of Biology 19:257–288.
Collins gives a historical overview that explores two related questions: (1) how did ecologists use evolutionary theory before the
late 1950s? And (2) in what sense was there a rapprochement between ecology and evolutionary biology during the 1950s and
1960s?
Dritschilo, William. 2008. Bringing statistical methods to community and evolutionary ecology: Daniel S. Simberloff. In
Rebels, mavericks, and heretics in biology. Edited by Oren Harman and Michael R. Dietrich, 356–371. New Haven, CT:
Yale Univ. Press.
A history of Simberloff’s research on island biogeography and his tenets for experimentation in ecology.
Futuyma, Douglas J. 2016. The evolution of evolutionary ecology. Israel Journal of Ecology & Evolution 59.4: 172–180.
A historical review of major developments in evolutionary ecology since its origins in the 1960s.
Hagen, Joel. 1992. An entangled bank: The origins of ecosystem ecology. New Brunswick, NJ: Rutgers Univ. Press.
Hagen documents the split between ecosystem ecology and evolutionary biology.
Mitman, Greg. 1992. The state of nature: Ecology, community, and social thought, 1900–1950. Chicago: Univ. of Chicago
Press.
Mitman traces how ecology and evolutionary thought were intertwined in the work of an influential group of ecologists at the
University of Chicago that included Allee and Emerson.
Odenbaugh, J. 2006. The strategy of “The strategy of model building in population biology.” Biology and Philosophy
21.5: 607–621.
Odenbaugh analyzes Levins’s classic essay in light of the collaboration between Levins, Robert MacArthur, Richard Lewontin, E.
O. Wilson, and others as a response to systems ecology.
Odenbaugh, Jay. 2013. Searching for patterns, hunting for causes: Robert MacArthur, the mathematical naturalist. In
Outsider scientists: Routes to innovation in biology. Edited by Oren Harman and Michael R. Dietrich, 181–198. Chicago:
Univ. of Chicago Press.
Odenbaugh explores MacArthur’s influence on ecology paying special attention to the ways in which MacArthur’s mathematical
expertise allowed him to introduce new approaches to the field.
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Evolutionary Genetics
Evolutionary genetics aims to understand the genetic basis of evolutionary change: that is, the effects that modes of selection,
drift, mutation, and migration have on genetic variation. Dietrich 2006 provides a brief and broad overview of the history of
evolutionary genetics. Beatty 1987 situates the debate between the classical and balance explanations of genetic variation in the
sociopolitical context of the time. Carlson 1981 provides a comprehensive biography of Muller’s contributions to evolutionary
genetics. Lewontin 1981 traces Dobzhansky’s contributions to the field. Crow 1987 argues that the Muller and Dobzhansky
disagreed about less than is commonly understood. Paul 1987 describes the origins and reception of Muller’s idea of genetic load
and genetic death. Sarkar 1992 offers a set of essays on the impact of Wright, Fisher, and Haldane on the history of evolutionary
and population genetics.
Beatty, John. 1987. Weighing the risks: Stalemate in the classical/balance controversy. Journal of the History of Biology
20:289–319.
Beatty traces the history of the classical/balance controversy in evolutionary genetics. He argues that while the issues of
underdetermination of theories by evidence is a central issue in the debate, the empirical problems cannot be properly
understood without also considering the sociopolitical considerations involved.
Carlson, Elof. 1981. Genes, radiation, and society: The life and work of H. J. Muller. Ithaca, NY: Cornell Univ. Press.
This is a scientific biography of the geneticist H. J. Muller that explores the period in genetics from the 1900s to the 1950s.
Muller’s work on genetic loads is considered in detail.
Crow, James. 1987. Muller, Dobzhansky, and overdominance. Journal of the History of Biology 20:351–380.
Crow analyzes the classical-balance controversy as a disagreement between Dobzhansky and Muller over the extent to which
genetic variation depends on overdominance.
Dietrich, Michael. 2006. From Mendel to molecules: A brief history of evolutionary genetics. In Evolutionary genetics:
Concepts and case studies. Edited by Charles W. Fox and Jason B. Wolf, 3–13. Oxford: Oxford Univ. Press.
Dietrich provides a historical overview of major controversies in evolutionary genetics, including the controversies between the
Mendelians and biometricians, Fisher and Wright, the importance of selection in evolution, the classical/balance approaches, and
the neutralist/selectionist approaches.
Lewontin, Richard. 1981. The scientific work of Theodosius Dobzhansky. In Dobzhansky’s genetics of natural
populations I-XLIII. Edited by R. C. Lewontin, J. A. Moore, W. B. Provine, and B. Wallace, 93–115. New York: Columbia
Univ. Press.
Lewontin critically discusses Dobzhansky’s contributions to evolutionary genetics, particularly the Genetics of Natural Populations
series.
Paul, Diane. 1987. ‘Our load of mutations’ revisited. Journal of the History of Biology 20:321–335.
Paul traces the controversy between Muller and Dobzhansky over the nature of the variation visible to natural selection. She
argues that their views are closer than they appear at first glance.
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Sarkar, Sahotra, ed. 1992. The founders of evolutionary genetics: A centenary reappraisal. Dordrecht, The Netherlands:
Kluwer Academic.
This is a collection of essays aimed at reassessing the biological work of Fisher, Haldane, Muller, and Wright at the (rough)
centenary of their birth (1889–1892).
Evolutionary Synthesis
The evolutionary synthesis is the period in the history of evolutionary biology from the 1920s to the 1950s. The traditional view is
that the synthesis starts with the bringing together of natural selection and Mendelian genetics; develops with field studies,
experiments, and museum-based analyses aimed at demonstrating evolution in Mendelian populations; and ends with the
dominance of natural selection as the main explanatory mechanism for most evolutionary patterns. Cain 2009 and Cain 1993
argue that there is much more to the evolutionary synthesis period than is in the traditional view. Smocovitis 1996 presents the
synthesis as of a piece with the broader movement of unifying knowledge. Provine 1988 argues that the synthesis was a
constriction of acceptable topics of research; Gould 1983 similarly describes the narrowed interest in selection as the hardening
of the synthesis. Smocovitis 2006 and Mayr and Provine 1980 explore the ways in which various fields in biology participated in
the synthesis period. Dietrich 1995 traces the influences that Goldschmidt’s controversial views had on the synthesis.
Cain, Joe. 1993. Common problems and cooperative solutions: Organizational activities in evolutionary studies,
1936–1947. Isis 84.1: 1–25.
Cain documents the ways in which relevant organizational activities had influence on the development of evolutionary genetics.
Cain, Joe. 2009. Rethinking the synthesis period in evolutionary studies. Journal of the History of Biology 42:621–648.
Cain proposes that historians abandon the idea of the evolutionary synthesis. There was much more to evolutionary studies in the
1920s and 1930s than is suggested in our commonplace narratives of this object in history.
Dietrich, Michael. 1995. Richard Goldschmidt’s ‘Heresies’ and the evolutionary synthesis. Journal of the History of
Biology 28:431–461.
Dietrich considers the controversial macroevolutionary views of German biologist Richard Goldschmidt and argues that opposition
to Goldschmidt’s saltational theory of evolution helped unify advocates of the evolutionary synthesis.
Gould, Stephen Jay. 1983. The hardening of the modern synthesis. In Dimensions of Darwinism. Edited by M. Grene,
71–93. Cambridge, UK: Cambridge Univ. Press.
Gould argues that during the 1940s, the synthesis “hardened” from a pluralistic state to an insistence that cumulative natural
selection leading to adaptation was the primary mechanism of evolutionary change.
Mayr, Ernst, and Provine, William, eds. 1980. The evolutionary synthesis. Cambridge, MA: Harvard Univ. Press.
Organized by Ernst Mayr, this collection of solicited contributions offers a number of different disciplinary perspectives on the
evolutionary synthesis and its origins.
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Provine, William. 1988. Progress in evolution and the meaning of life. In Evolutionary progress. Edited by M. Nitecki,
49–74. Chicago: Univ. of Chicago Press.
Provine argues that the search for a single synthetic theory of neo-Darwinian evolution is misguided and that synthesis should be
replaced by the idea of constriction. Architects of the synthesis did not agree on a common theory but agreed on what should be
eliminated from modern evolution and so constricted evolutionary theory.
Smocovitis, V. B. 1996. Unifying biology: The evolutionary synthesis and evolutionary biology. Princeton, NJ: Princeton
Univ. Press.
Smocovitis reconstructs the history of the evolutionary synthesis of Darwinian natural selection and Mendelian genetics during the
1930s–1940s as part and parcel of the contemporaneous global philosophical movement toward the unity of knowledge.
Smocovitis, V. B. 2006. Keeping Up with Dobzhansky: G. Ledyard Stebbins, plant evolution, and the evolutionary
synthesis. History and Philosophy of the Life Sciences 28:11–50.
Smocovitis extends the history of the synthesis to include the important contributions on plant evolution articulated by G. Ledyard
Stebbins.
Human Evolution
The study of human evolution concerns the processes of human origins and evolution as well as its products. Bowler 1986
provides a broad historical account of theories of human evolution. Delisle 2006 shows that the debates in paleoanthropology all
stem from the general theory of evolution by natural selection. Goodrum 2009 sets out a historiography of paleoanthropology.
Sommer 2008 traces the rise of molecular anthropology. De Vos 2009 discusses the receptions of the discoveries of Neanderthal
Man, Homo erectus, and Homo floresiensis. Gannett 2001 and Kaplan and Winther 2013 explore the concept of race in studies of
human evolution.
Bowler, Peter. 1986. Theories of human evolution: A century of debate, 1844–1944. Baltimore, MD: Johns Hopkins Univ.
Press.
Bowler explores the history of alternative theories of human evolution, discussing their import against contemporaneous fossil
evidence, cultural evolution, and the sociopolitical climate.
Delisle, Richard. 2006. Debating humankind’s place in nature, 1860–2000: The nature of paleoanthropology. Upper
Saddle River, NJ: Prentice Hall.
Delisle presents a comprehensive history of debates in paleoanthropology, showing that all stem from one general theory:
evolution by natural selection.
de Vos, John. 2009. Receiving an ancestor in the phylogenetic tree. Journal of the History of Biology 42:361–379.
De Vos compares the reception of the discoveries of Neanderthal Man, Homo erectus, and Homo floresiensis.
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Gannett, Lisa. 2001. Racism and human genome diversity research: The ethical limits of ‘population thinking.’
Philosophy of Science 63:S479–S492.
Gannett rejects the commonplace view that in the 1950s population genetics banished scientific racism by banishing the term
“race” in favor of the term “population.” Instead, Gannett argues, races were reconceptualized as populations and a “populational”
concept of race was substituted for a “typological” one.
Goodrum, Matthew. 2009. The history of human origins research and its place in the history of science: Research
problems and historiography. History of Science 47:337–357.
Goodrum presents a historiography of paleoanthropology and other sciences that explore human origins. In so doing, he identifies
new areas of promising inquiry and related historical methods.
Kaplan, J. M., and R. G. Winther. 2013. Prisoners of abstraction? The theory and measure of genetic variation, and the
very concept of ‘race.’ Biological Theory 7:401–412.
Kaplan and Winther characterize three approaches that have been taken to the study of human genetic variation: genetic
diversity, genetic differentiation, and genetic heterozygosity. They argue that while these approaches may seem to reach different
conclusions, they are in fact all legitimate approaches focusing on different aspects of genetic variation, with the choice of
approach being a matter of convention.
Sommer, Marianne. 2008. History in the gene: Negotiations between molecular and organismal anthropology. Journal for
the History of Biology 41:473–528.
Sommer explores the rise of molecular anthropology through the 1960s–1980s, paying special attention to the forces influencing
the anthropological gene/genome to become the primary evidential arbiter.
Molecular Evolution
Molecular evolution emerged as a new hybrid field in the 1960s. Dietrich 1994 focuses on the development of the neutral theory
of molecular evolution, which gave a greater role to drift. Provine 1990 focuses on the work of Motoo Kimura and his path to the
neutral theory. Suarez and Barahona 1996 shifts the focus from theory to experiment in a careful analysis of the role of
technological innovations that allowed greater access to molecular information. Dietrich 1998 and Hagen 1999 contextualize the
rise of molecular evolution within the wider struggle between naturalists and experimentalists and organismal and molecular
biologists over the nature of biology. Strasser 2010 builds on a similar distinction to examine the role of the first sequence
database. Morgan 1998 offers a careful history of the early days of the molecular clock. More material on the clock and a range of
other topics are available at the Perspectives on Molecular Evolution website, including contributions by historians and biologists
active in the field.
Dietrich, Michael. 1994. The origins of the neutral theory of molecular evolution. Journal of the History of Biology
27:21–59.
Dietrich argues that the neutral theory was not a simple continuation of the classical-balance controversy. Biochemists provided
important new sources of evidence for neutrality and drift.
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Dietrich, Michael. 1998. Paradox and persuasion: Negotiating the place of molecular evolution within evolutionary
biology. Journal of the History of Biology 31:85–111.
Dietrich places the emergence of molecular evolution within the context of a struggle for authority between organismal and
molecular biologists by examining the debate over the differences between rates of molecular and morphological evolution.
Hagen, Joel. 1999. Naturalists, molecular biologists, and the challenges of molecular evolution. Journal of the History of
Biology 32:321–341.
Hagen describes the tensions between molecular and organismal biologists in the history of molecular evolution focusing on the
development of molecular systematics.
Morgan, Gregory. 1998. Emile Zuckerkandl, Linus Pauling, and the molecular evolutionary clock, 1959–1965. Journal of
Morgan describes the discovery of the molecular clock in the 1960s and its impact on the new field of molecular evolution.
Perspectives on Molecular Evolution.

This website offers a range of documents, profiles, commentaries, and video clips on the history of molecular evolution.
Provine, William. 1990. The neutral theory of molecular evolution in historical perspective. In Population biology of
genes and molecules. Edited by Naoyuki Takahata and James Crow, 17–31. Tokyo: Baifukan.
Provine describes Kimura’s approach to the neutral theory and its initial confusion with earlier debates over drift.
Strasser, Bruno. 2010. Collecting, comparing, and computing sequences: The making of Margaret O. Dayhoff’s Atlas of
protein sequence and structure, 1954–1965. Journal of the History of Biology 43:623–660.
Strasser describes Dayhoff’s efforts to create the first database of molecular sequences for comparative analysis. He argues that
this effort was an extension of earlier ways of knowing.
Suarez, Edna, and Anna Barahona. 1996. The experimental roots of the neutral theory of molecular evolution, History
and Philosophy of the Life Sciences 18:55–81.
Suarez and Barahona focus on the important role of technological advances in the history of molecular evolution, which allowed
for increasing experimental access to measures of genetic variation.
Natural Selection
In second half of the 20th century and the early part of the 21st, biologists and philosophers sought to clarify the theory of natural
selection and to identify methodologies for detecting it. Sober 1984, Hodge 1987, and Okasha 2006 offer historically informed
philosophical analyses of natural selection. Borrello 2010 reveals the history of group selection; Harman 2010 focuses on the
evolution of altruism. Shavit 2004 offers an explanation of the persistence of debates over group selection.
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Borrello, Mark. 2010. Evolutionary restraints: The contentious history of group selection. Chicago: Univ. of Chicago
Press.
Borrello traces the history of debates over group selection, focusing in particular on group selection advocate Wynne-Edwards
and his disagreements with Ernst Mayr, G. C. Williams, and Richard Dawkins.
Harman, Oren. 2010. The price of altruism: George price and the search for the origins of kindness. New York: W. W.
Norton.
This is a scientific biography of George Price centered around his struggle to understand how evolution by natural selection could
allow an organism to sacrifice itself for the good of other organisms, that is, biological altruism.
Hodge, M. J. S. 1987. Natural selection as a causal, empirical, and probabilistic theory. In The probabilistic revolution.
Vol. 2. Edited by L. Krüger, G. Gigerenzer, and M. Morgan, 233–270. Cambridge, MA: The MIT Press.
Hodge describes natural selection as a Darwinian vera causa but importantly also as a probabilistic cause, distinguishing it from
drift, which Hodge describes as fortuitous differential reproduction.
Okasha, Samir. 2006. Evolution and the levels of selection. Oxford: Oxford Univ. Press.
Okasha provides a comprehensive critical analysis of the levels of selection debate in evolutionary biology. The centerpiece of the
book is an analysis of the price equation and multivariate regression in the context of multilevel selection.
Shavit, Ayelet. 2004. Shifting values partly explain the debate over group selection. Studies in History and Philosophy of
Science Part C: Studies in History and Philosophy of Biological and Biomedical Sciences 35.4: 697–720.
Shavit argues that history of social and political uses of “group” and “group selection” can explain, at least in part, some of the
empirical deficiencies of the debate as well as its persistence.
Sober, Elliott. 1984. The nature of selection: Evolutionary theory in philosophical focus. Chicago: Univ. of Chicago
Press.
Read by philosophers and biologists alike, this book aims to provide a systematic account of the theory of natural selection that
clarifies ambiguities within the theory. It is notable for its account of the Hardy-Weinberg Law as a “zero-force” state, with other
evolutionary processes such as natural selection acting as forces.
Paleobiology and Macroevolution
Paleobiology combines natural history biology with paleontology to understand life’s evolutionary history. Macroevolution is the
study of evolutionary processes above the species level, often on a geological time scale. Sepkoski 2012 and Sepkoski and Ruse
2009 provide comprehensive overviews of paleobiology. Cain 2009 examines Simpson’s influence on paleontology. Millstein 2000
analyzes models of macroevolution in relation to drift. Sapp traces the history of the theory of symbiosis.
Cain, Joe. 2009. Ritual patricide: Why Stephen Jay Gould assassinated George Gaylord Simpson. In The paleobiological
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revolution: Essays on the growth of modern paleontology. Edited by David Sepkoski and Michael Ruse, 346–363.
Chicago: Univ. of Chicago Press.
Cain examines the way Gould led the transition away from Simpson’s phyletic gradualism to Gould’s gradualism interrupted by
catastrophe or punctuated equilibrium.
Millstein, Roberta L. 2000. Chance and macroevolution. Philosophy of Science 67.4: 603–624.
This paper offers an analysis of the stochastic macroevolutionary models of Stephen Jay Gould, David Raup, Jack Sepkoski, and
colleagues, comparing them to models of random drift.
Sapp, Jan. 1994. Evolution by association: A history of symbiosis. Oxford: Oxford Univ. Press.
Sapp presents a comprehensive history of the theory of symbiosis from its origins in the late 19th century to its current status as a
major theory of the origins of eukaryotes.
Sepkoski, David. 2012. Rereading the fossil record: The growth of paleobiology as an evolutionary discipline. Chicago:
Univ. of Chicago Press.
Sepkoski documents the efforts of paleontologists in the second half of the 20th century to develop their own distinctive
evolutionary theories and models.
Sepkoski, David, and Michael Ruse, eds. 2009. The paleobiological revolution: Essays on the growth of modern
paleontology. Chicago: Univ. of Chicago Press.
Containing essays from illustrious historians of biology and biologists, this book covers major innovations in paleobiology, recent
paleontology and its historical and conceptual significance, and recent paleobiology.
Population Genetics
In population genetics, evolution is defined as any change in the frequency of alleles within a population from one generation to
the next. As such, its aim is to explain the evolution of populations in terms of the dynamics of gene frequency change. In the
latter part of the 20th century, controversies over the field’s foundations deepened. Provine 2001 provides a general history of the
origins of theoretical population genetics via its architects, Haldane, Fisher, and Wright. Dietrich and Skipper 2012 explores
Wright’s adaptive landscape metaphor, whereas Plutynski 2006 explores Fisher’s Fundamental Theorem of Natural Selection.
Skipper 2002 and Skipper 2009 analyze the persistent controversy between Fisher and Wright. Hodge 1992 provides a
historiography of population genetics via the philosophical commitments of Fisher and Wright. Sarkar 2007 discusses Haldane’s
contributions to population genetics. Slatkin and Veuille 2002 collects essays concerning the influence of Malécot on population
genetics.
Dietrich, M. R., and R. A. Skipper. 2012. A shifting terrain: A brief history of the adaptive landscape. In The adaptive
landscape in evolutionary biology. Edited by E. Svensson and R. A. Calsbeek, 3–15. Oxford: Oxford Univ. Press.
Dietrich and Skipper discuss the origins, development, and extension of Sewall Wright’s adaptive landscape diagram.
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Hodge, M. J. S. 1992. Biology and philosophy (including ideology): A study of Fisher and Wright. In The founders of
evolutionary genetics: A centenary reappraisal. Edited by S. Sarkar, 231–293. Dordrecht, The Netherlands: Kluwer
Academic.
Hodge explores the philosophical views of Fisher and Wright as a way of offering a historiography of population genetics that
explores three historical topics: Darwin as mechanist, the evolution revolution, and the probabilistic revolution in evolutionary
genetics.
Plutynski, Anya. 2006. What was Fisher’s Fundamental Theorem and what was it for? Studies in History and Philosophy
of Science C: Studies in History and Philosophy of Biology 37:59–82.
Plutynski revisits the Price-Ewens interpretation of Fisher’s Fundamental Theorem of Natural Selection, paying special attention
to its role in Fisher’s work on population genetics.
Provine, William. 2001. The origins of theoretical population genetics. Chicago: Univ. of Chicago Press.
Provine elaborates a history of the origins of theoretical population genetics through its founders, R. A. Fisher, Sewall Wright, and
J. B S. Haldane.
Sarkar, Sahotra. 2007. Haldane and the emergence of modern evolutionary theory. In Philosophy of biology. Edited by M.
Matthen and C. Stephens, 49–86. Amsterdam, The Netherlands: Elsevier.
Sarkar elaborates J. B. S. Haldane’s role at the origin of theoretical population genetics.
Skipper, Robert A., Jr. 2002. The persistence of the R.A. Fisher-Sewall Wright controversy. Biology and Philosophy
17:341–367.
Skipper critical analyzes a debate in the late 1990s between biologists Jerry Coyne and Michael Wade over the scope and
application of Fisher’s and Wright’s evolutionary theories.
Skipper, Robert A., Jr. 2009. Revisiting the Fisher-Wright controversy. In Descended from Darwin: Insights into American
evolutionary studies, 1925–1950. Edited by J. Cain and M. Ruse, 299–322. Philadelphia: American Philosophical
Association Press.
Skipper extends the extant historical analysis of the Fisher-Wright controversy from the 1950s to the early 2000s. Special
attention is paid to the debates over the scope of Fisher’s and Wright’s evolutionary theories, theories of dominance, and role of
drift in evolution.
Slatkin, Montgomery, and Michel Veuille, eds. 2002. Modern developments in theoretical population genetics: The legacy
of Gustav Malécot. Oxford: Oxford Univ. Press.
This is an edited collection of essays written by biologists exploring the contributions to population genetics inspired by Gustav
Malécot.
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Sexual Selection
With the publication of Robert Trivers’s Bateman-inspired account of natural selection, it seemed for a time as though the “good
genes” account of sexual selection on males, with its promiscuous males and coy females, would hold sway. However, later
developments would broaden sexual selection considerably to include a variety of assumptions and models. Cronin 1991 offers
an account of the canonical view of sexual selection and its historical roots. Hrdy 2006 as well as Snyder and Gowaty 2007
reexamine and challenge the canon. Milam 2011 fills in major gaps in the usual histories of sexual selection.
Birkhead, Tim. 2000. Promiscuity: An evolutionary history of sperm competition. Cambridge, MA: Harvard Univ. Press.
Birkhead provides a comprehensive account of the evolution of sexual behaviors across a broad range of species, including
monogamy versus polygamy, testicle size, ejaculatory volume versus promiscuity in females, and so on.
Cronin, Helena. 1991. The ant and the peacock: Altruism and sexual selection from Darwin to today. Cambridge, UK:
In this book, covering the topic of altruism in evolution as well as sexual selection, Cronin explores contemporary debates over
sexual selection through the lens of the debate between Darwin and Wallace.
Hrdy, Sara. 2006. Empathy, polyandry, and the myth of the coy female. In Conceptual issues in evolutionary biology. 3d
ed. Edited by E. Sober, 131–159. Cambridge, MA: The MIT Press.
Hrdy reviews the work of Bateman and Trivers, arguing that it has been inappropriately extrapolated to primates and showing how
subsequent work in primatology demands new types of sexual selection explanations. In particular, she argues that sexual
selection often operates on females, not just males.
Milam, Erika L. 2011. Looking for a few good males: Female choice in evolutionary biology. Baltimore: Johns Hopkins
Univ. Press.
Milam’s research shows that studies of sexual selection did not end in the early 20th century only to be reborn in the late 20th,
revealing the sexual selection research that occurred throughout the entire 20th century.
Snyder, B. F., and P. A. Gowaty. 2007. A reappraisal of Bateman’s classic study of intrasexual selection. Evolution 61.11:
2457–2468.
Snyder and Gowaty defend the thesis that Bateman’s classic 1948 study of sexual selection in Drosophila melanogaster, while
groundbreaking, was methodologically flawed. They argue that problems included the elimination of genetic variance, sampling
biases, miscalculations of fitness variances, statistical pseudo-replication, and selective presentation of data.
Sociobiology
E. O. Wilson inaugurated modern sociobiology in the 1970s as an attempt to extend evolutionary biology to the social, cultural,
and behavioral phenomena. Kaye 1986 provides a history of sociobiology from its roots in social Darwinism. Jumonville 2002
discusses the cultural debates stemming from sociobiology. Li Jianhui and Fan 2003 traces the spread of sociobiology in China.
Segestråle 2013 chronicles the contributions of Hamilton to sociobiology and evolutionary biology more generally. Gibson 2013
discusses Wilson’s recent abandonment of kin selection theory.
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Gibson, Abraham. 2013. Edward O. Wilson and the organicist tradition. Journal of the History of Biology 46:599–630.
Gibson discusses Edward O. Wilson’s recent decision to abandon kin selection theory.
Jumonville, Neil. 2002. The cultural politics of the sociobiology debate. Journal of the History of Biology 35:569–593.
Jumonville argues that the sociobiology debates of the last quarter of the 20th century is best understood in the context of a
struggle between two main ideologies: liberal universalism and identity politics.
Kaye, H. L. 1986. The social meaning of modern biology: From social Darwinism to sociobiology. New Haven, CT: Yale
Univ. Press.
Kaye provides a historical and philosophical analysis of the implication of biological evolution on our understanding of human
nature and how we are to live. Kay argues that in spite of the weaknesses of the cultural theories, they have been historically
attractive.
Li Jianhui, Li, and Hong Fan. 2003. Science as ideology: The rejection and reception of sociobiology in China. Journal of
Li Jianhui and Fan trace the spread of sociobiology in China.
Segestråle, Ullica. 2013. Nature’s oracle. The life and work of W.D. Hamilton. Oxford: Oxford Univ. Press.
Segestråle offers a scientific biography of W. D. Hamilton. She portrays Hamilton as a voraciously curious iconoclast whose
abilities produced groundbreaking work in evolutionary genetics, including genic selectionism and the evolution of sex.
Species and Speciation
Speciation is the process or processes by which a lineage, or species, splits into two. Defining “species” and understanding the
mechanisms of speciation are controversial topics in biology. Wilkins 2011 is a comprehensive historical and philosophical
analysis of species from Antiquity. Haffer 2007 is a biography of Mayr that explores his contributions to debates about species.
Laporte 1994 traces Simpson’s contributions to species and speciation. Schloegel 1999 analyzes Sonneborn’s work on species in
protozoa. Provine 1989 discusses founder effect speciation. McOuat 2001 discusses the notion of species as natural kinds.
Haffer, Jürgen. 2007. Ornithology, evolution, and philosophy: The life and science of Ernst Mayr 1904–2005. Berlin:
Springer.
This is the first comprehensive scientific biography of Ernst Mayr.
Laporte, Léo F. 1994. Simpson on species. Journal of the History of Biology 27.1: 141–159.
LaPorte traces Simpson’s participation in the evolutionary synthesis via his focus on his development of the concept of species.
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McOuat, Gordon. 2001. From cutting nature at its joints to measuring it: Species, new kinds and new kinds of people in
biology. Studies in History and Philosophy of Science 32:613–645.
McOuat explores the question of species as natural kinds —and questioning views on this concept—during the transition from the
19th- to 20th-century biology.
Plutynski, Anya. 2018. Speciation post synthesis: 1960–2000. Journal of the History of Biology.
Plutynski characterizes the history of work on speciation since the mid-20th century as a history of divergence and diversification.
It covers both theoretical advances and empirical insights into how different lineages, with different genetics and ecological
conditions, are shaped by very different modes of diversification.
Provine, William. 1989. Founder effects and genetic revolutions in microevolution and speciation: A historical
perspective. In Genetics, speciation, and the founder principle. Edited by L. V. Giddings, K. Kaneshiro, and W. Anderson,
43–78. Oxford: Oxford Univ. Press.
Provine critically analyzes the historical biological work on founder effect speciation.
Schloegel, Judy Johns. 1999. From anomaly to unification: Tracy Sonneborn and the species problem in protozoa,
1954—1957. Journal of the History of Biology 32:93–132.
Examines the critique of the biological species concept advanced by protozoan geneticist Tracy Sonneborn at the 1955 AAAS
symposium on “the species problem,” published subsequently in 1957.
Wilkins, John S. 2011. Species: The history of the idea. Berkeley: Univ. of California Press.
Wilkins provides a comprehensive historical and conceptual analysis of the concept of species from Plato to the present.
Systematics
John Lindley, in 1830, was the first to define “systematics,” though his term was “systematic botany.” Generally, systematics is the
comparative study and classification of groupings of organisms at the species level or higher over their evolutionary histories.
Cain 2004 traces the origins of the Society of Systematic Zoology. Buhs 2000 explores the history of ant systematics in the early
20th century, while Strasser 2010 looks at systematic practices during the same period. Hagen 2003 and Suárez-Díaz 2013
discuss the rise of statistical techniques and molecular data in systematics. The Hamilton 2013 collection of essays explores the
origins and development of phylogenetic systematics. Sapp 2009 traces the history of microbial evolution against the background
of the Tree of Life. Hull 1988 offers a picture of scientific change as evolutionary change, using systematic biology as a case
study.
Buhs, Joshua. 2000. Building on bedrock: William Steel Creighton and the reformation of ant systematics, 1925–1970.
Journal of the History of Biology 33:27–70.
Ideas about the natural world are intertwined with the personalities, practices, and the workplaces of scientists.
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Cain, Joe. 2004. Launching the society of systematic zoology in 1947. In Milestones in systematics. Edited by D.
Williams and P. Forey, 19–48. London: CRC Press.
Cain supplants Richard Blackwelder’s history of the formation of the Society of Systematic Zoology by contextualizing it within
postwar reconstruction.
Hagen, Joel. 2003. The statistical frame of mind in systematic biology from quantitative zoology to biometry. Journal of
Hagen traces the rise in the use of statistics in systematics, that is, the statistical frame of mind, after World War II.
Hamilton, Andrew, ed. 2013. The evolution of phylogenetic systematics. Berkeley: Univ. of California Press.
This is a collection of essays by historians, philosophers, and biologists on the rise of phylogenetic systematics. Its aim is to place
systematics in a broader context of the history of the biological sciences.
Hull, David. 1988. Science as a process: An evolutionary account of the social and conceptual development of science.
Chicago: Univ. of Chicago Press.
Hull develops an account of scientific change using the principles of evolution. In particular, Hull argues that the success of
science can be explained via competition and cooperation.
Sapp, Jan. 2009. The new foundations of evolution: On the Tree of Life. Oxford: Oxford Univ. Press.
Sapp presents a history of microbial evolution, arguing that evolution at the microbial level involves processes that classical
evolutionists eschew.
Strasser, Bruno. 2010. Laboratories, museums, and the comparative perspective: Alan A. Boyden’s serological
taxonomy, 1925–1962. Historical Studies in the Natural Sciences 40.2: 149–182.
Strasser argues that contra to the view that the practices of natural history have been replaced by experimentation, natural history
and experimental practices have blended together.
Suárez-Díaz, Edna. 2013. The long and winding road of molecular data in phylogenetic analysis. Journal of the History of
Biology 47.2: 443–478
Suárez-Díaz explores the development of molecular techniques in phylogenetic analysis, concluding that a proper history comes
from understanding the broader role of molecular techniques across a variety of biological sciences.
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