Festschrifts + Commentary

Evolution & Human Behavior special issue
Summers-Crespi volume
HBES Award
Obituaries & Memorials

Two main compilations have been published that are aimed at honoring and advancing RDA’s thought and work: a 2005 special issue of Evolution and Human Behavior, and a 2013 volume entitled Human Social Evolution edited by Kyle Summers and Bernard Crespi.

Special Issue of Evolution & Human Behavior

In 2005 the journal of the Human Behavior and Evolution Society released its largest issue to date, a collection of papers by students and friends of RDA on aspects of their own research that were inspired by his ideas. Introducing the issue is an overview by William Irons on the ways in which RDA was advancing our understanding of the evolution of human behavior and psychology. Below are the contents of that issue, each article of which can be downloaded by clicking on the reference.

[PDF] Irons, W. 2005. How has evolution shaped human behavior? Richard Alexander’s contribution to an important question. Evolution and Human Behavior 26: 1-9.

[PDF] Flinn, M. V., Geary, D. C. & Ward, C. V. 2005. Ecological dominance, social competition, and coalitionary arms races: Why humans evolved extraordinary intelligence. Evolution and Human Behavior 26: 10-46.

[PDF] Lahti, D. C. & Weinstein, B. S. 2005. The better angels of our nature: group stability and the evolution of moral tension. Evolution and Human Behavior 26: 47-63.

[PDF] Low, B. S. 2005. Women’s lives there, here, then, now: a review of women’s ecological and demographic constraints cross-culturally. Evolution and Human Behavior 26: 64-87.

[PDF] Nesse, R. M. 2005. Natural selection and the regulation of defenses – A signal detection analysis of the smoke detector principle. Evolution and Human Behavior 26: 88-105.

[PDF] Summers, K. 2005. The evolutionary ecology of despotism. Evolution and Human Behavior 26: 106-135.

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Summers & Crespi volume: Human Social Evolution: The Foundational Works of Richard D. Alexander

In 2013, biologists Kyle Summers (East Carolina University) and Bernard Crespi (Simon Fraser University), both of them erstwhile students and longtime friends of RDA, published an ingenious collection.  Fifteen topics on which RDA made important contributions (4 of which lay the foundations for an understanding of human social evolution and 11 of which deal explicitly with it), are each represented by one classic RDA paper or lengthy excerpt.  As RDA had a penchant for publishing ideas opportunistically without much thought to how easily people might access them, such a compendium is valuable just for its own sake. But what makes this volume amazing is that Summers and Crespi assembled colleagues and former students of RDA who have professional interests in these same areas. Each of these scientists has contributed a chapter designed to be paired with one of those RDA pieces. In these introductions or companion pieces, the authors describe the contemporary state of the field in the light of RDA’s work, drawing clear lines between his ideas and the important questions of today’s science.

This book can be purchased from Amazon (here), and from Oxford University Press (here).

Table of Contents:

Introduction – Kyle Summers and Bernard Crespi

Part I – General Foundations

Chapter 1: Insect Behavior & Evolution
Introduction by Mary Jane West-Eberhard, Smithsonian Tropical Research Institute.
Excerpt from Alexander, R. D. 1969. Comparative animal behavior and systematics. In: Systematic Biology. Proceedings of the International Conference on Systematics (Ann Arbor, Michigan, July 1967). National Academy of Sciences Publication 1962: 494-517.

Chapter 2: Cooperation
Introduction by Steven Frank, University of California at Irvine.
Excerpt from: Alexander, R.D. 1986. The Biology of Moral Systems. New York, Aldine Press.

Chapter 3: Eusociality in Naked Mole Rats
Introduction by Paul Sherman, Cornell University.
Excerpt from Alexander, R.D., Noonan, K.M. and Crespi, B.J. 1991. The Evolution of Eusociality. In P. Sherman, J. Jarvis and R.D. Alexander (eds). The Biology of the Naked Mole Rat: 3-44. Princeton, NJ: Princeton University Press.

Chapter 4: Parent-offspring Conflict and Manipulation
Introduction by David Queller, Washington University.
Excerpt from Alexander, R.D. 1974. The evolution of social behavior. Annual Review of Ecology and Systematics 5:325-383.

Part II – Human Social Evolution

Chapter 5: Biology and Culture
Introduction by Mark Flinn, University of Missouri
Excerpt from Alexander, R.D. Evolution and culture. In Evolutionary Biology and Human Social Behavior: an Anthropological Perspective. N. Chagnon and W.G. Irons (eds). Pp 59-78. North Scituate, MA: Duxbury Press.

Chapter 6: Intergroup Competition and Within-group Cooperation
Introduction by Bobbi Low, University of Michigan.
Excerpt from Alexander, R.D. and Tinkle, D.W. 1968. Review of On Aggression by Konrad Lorenz and The Territorial Imperative by Robert Ardrey. Bioscience 18: 245-248.

Chapter 7: Kinship, Parental Care, and Human Societies
Introduction by Beverly Strassmann, University of Michigan.
Excerpt from Alexander, R.D. and Noonan, K.M. 1979. Concealment of Ovulation, parental care, and human social evolution. In N.A. Chagnon and W.G. Irons (eds). Evolutionary Biology and Human Social Behavior: An Anthropological Perspective. 436-453. North Scituate, MA: Duxbury Press.

Chapter 8: Human Childhood
Introduction by Paul Turke, University of Michigan.
Altriciality: Why are human babies helpless? In Alexander, R.D. 1990. How Did Humans Evolve? Reflections on a Uniquely Unique Species. Univ. Mich. Zool. Special
Publication 1: 1-38.

Chapter 9: Indirect Reciprocity
Introduction by Karl Sigmund, University of Vienna.
Excerpt from Alexander, R.D. 1986. The Biology of Moral Systems. New York, Aldine Press.

Chapter 10: The Evolution of Intelligence
Introduction by Robin Dunbar, Oxford University.
Alexander, R.D. Evolution of the Human Psyche 1989. In P. Mellars and C. Stringer (eds). The Human Revolution. Behavioral and Biological Perspectives on the Origins of Modern Humans: pp. 455-513. Princeton, NJ: Princeton University Press.

Chapter 11: Evolution of Morality
Introduction by David Lahti, City University of New York.
Alexander, R.D. Biology and the Moral Paradoxes. Journal of Biological Structures 5:389-395.

Chapter 12: Evolution and Humor
Introduction by Stanton Braude, Washington University.
Alexander, R.D. Ostracism and Indirect Reciprocity: The Reproductive Significance of Humor. 1986. Ethology and Sociobiology 7: 253-270.

Chapter 13: Ecological Constraints and Human Cooperation
Introduction by Laura Betzig, University of Michigan.
Excerpt from Alexander, R.D., Noonan, K.M. and Crespi, B.J. 1991. The Evolution of Eusociality. In P. Sherman, J. Jarvis and R.D. Alexander (eds). The Biology of the Naked Mole Rat: pp. 3-44. Princeton, NJ: Princeton University Press.

Chapter 14: Evolution and Religion
Introduction by William Irons, Northwestern University.
Religion, Evolution and the Quest for Global Harmony – Original essay by RDA for this volume.

Chapter 15: Evolution and the Arts
Introduction by Kyle Summers, East Carolina University & Bernard Crespi, Simon Fraser University.
Excerpt from Alexander, R.D. 2003. Evolutionary Selection and the Nature of Humanity. Chapter 15. In: V. Hosle and Ch. Illies (eds). Darwinism and Philosophy. University of Notre Dame Press.

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HBES Distinguished Scientific Award for Lifetime Contribution to Evolution and Human Behavior

In 2008, the Human Behavior and Evolution Society awarded RDA with this lifetime achievement award at their annual meeting, in Kyoto, Japan.  Here is the announcement of this award, including a brief explanation of RDA’s contributions to the field, as printed in the Winter 2008 issue of the HBES Newsletter.

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Obituaries & Memorials

RDA passed away peacefully on August 20, 2018, at 88 years old.  Here is a transcript of his obituary (from the Dignity Memorial website), which was published in the Ann Arbor News on August 23, 2018.

A professional remembrance of his life and work was published in the official journal of the Human Behavior & Evolution Society, which he co-founded:

[PDF] Kyle Summers, David Lahti, Stanton Braude, Beverly Strassmann, and Joan Strassmann. 2019. Obituary: The nine lives of Richard D. Alexander. Evolution and Human Behavior 40:133-139.

A Celebration of Life Memorial was held on April 27, 2019, attended by his family, friends, former students, and colleagues. Here is the announcement.

RDA at his farm with one of his (hundreds of) carved walking sticks, 2017


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Articles Showcasing RDA’s Work:

[PDF] 1963. Singing crickets and periodical cicadas. Research News 14(5), 6 pp. Office of Research Administration, University of Michigan, Ann Arbor, MI.

[PDF] 1974. Evolution and behavior. Research News 25(5), 18 pp. Office of Research Administration, University of Michigan, Ann Arbor, MI.

[PDF] 1978. Evolutionary biology called key to old riddle. Ann Arbor News, November 23, p.A-6.

[PDF] Stanton Braude. 1997. The predictive power of evolutionary biology and the discovery of the naked mole-rat. Reports of the National Center for Science Education 17(4):12-15. An unpublished addendum that tabulates all of the relevant predictions is here.

Podcasts Showcasing RDA’s Ideas:

Evolution Can’t Explain The Arts, Can It? Kyle Summers discusses the ideas of Richard Alexander, what evolutionary theory has to tell us about the arts, and the evolutionary ecology of despotism. Biological Realist Podcast, Michael McConkey. 2016. [Link to audio file MP3 (50MB) or online at YouTube].

Humans: Our Own Hostile Force Of Nature? Mark Flinn discusses the contributions of Richard Alexander to the evolutionary theory of human sociality.  Biological Realist Podcast, Michael McConkey. 2016. [Audio file MP3 (46MB) or online at YouTube].


RDA did not often publish abstracts of presentations for which there are no accompanying publications with him as an author.  No attempt has been made to find published abstracts, and in most cases they are probably not of lasting interest.

[PDF] Dunning, D. Covalt & Richard D. Alexander. 1971. Courtship in Magicicada septendecim. Ecological Society of America 52 (1):57-58.  This work became the basis of a publication, part of the continuing story of speciation and maintenance of reproductive isolation in periodical cicadas that was originally established by RDA and colleagues:  Dunning, D. Covalt, John A. Byers, & C. Dwight Zanger. 1979. Courtship in two species of periodical cicadas, Magicicada septendecim and Magicicada cassini. Animal Behaviour 27:1073-1090 [PDF]. In the Acknowledgements to this paper the following text appears: “We thank Dr. Richard D. Alexander, with whom the senior author worked on Brood X in Michigan in 1970. Dr. Alexander was most generous of time and research space for a novice cicada person and provided an exciting introduction to work on these animals. He also read and criticized an early draft of this paper.”

[PDF] Alexander. R. D. 1972. Reproductive competition and the evolution of sociality. American Society of Zoologists 12:648.  This is RDA’s first published elaboration of his parental manipulation theory, which would form a large part of his 1974 papers, “The evolution of social behavior”, Annual Review of Ecology and Systematics 5: 352-383 [PDF]; and “Evolution of sterile castes in the social insects: Kin selection or parental manipulation?”, Florida Entomologist 57(1): 32 [PDF]. From a historical perspective a striking feature of this abstract is that, as can be seen in the digital scan, it immediately follows Robert Trivers’ (characteristically brief) abstract on the same topic, entitled “Mother-offspring conflict”. Trivers’ work would eventually be developed into one of the most highly cited articles in the evolutionary study of behavior: Robert Trivers. 1974. Parent-offspring conflict. American Zoologist 14:249-264 [PDF]. For decades following this conference, Trivers would refer to his difference with RDA on this topic, and insist that RDA had neglected the offspring’s interests in representing the conflict unilaterally as parental manipulation. However, RDA’s ARES paper notes that neither Trivers nor anyone else has determined who wins in a parent-offspring conflict scenario, and proposes that “genes which reduce the mother’s reproduction by causing competitive interactions among embryos in the uterus cannot spread regardless of their advantage to the embryos possessing them”. Apparently neither the authors nor anyone else has resolved this theoretical disagreement since these two back-to-back abstracts first appeared.

[PDF] Alexander, R. D. 1976. The evolution of social behavior. In The Bicentennial: Learn From The Past – Learn For The Future. Proceedings of the Twelfth Annual Southeastern Michigan Junior Science And Humanities Symposium. February 21-22, 1976. Pp. 3-4.

A pastel of RDA whose origin is uncertain– perhaps created by an undergraduate student from Japan. Courtesy of Nancy Alexander.

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