Evolution and Human Behavior 40 (2019) 133–139 Contents lists available at ScienceDirect Evolution and Human Behavior journal homepage: www.elsevier.com/locate/ens Obituary: The nine lives of Richard D. Alexander T ⁎ Kyle Summersa, , David Lahtib, Stanton Braudec, Beverly Strassmannd, Joan Strassmannc a Department of Biology, East Carolina University, Greenville, NC 27858, United States b Department of Biology, Queens College, City University of New York, Flushing, NY, 11367, United States c Department of Biology, Washington University in St. Louis, St. Louis, MO 63130, United States d Department of Anthropology, University of Michigan, Ann Arbor, MI 48104, United States Richard D. Alexander, T.H. Hubbell Distinguished University carver. In addition to his professional scientific writings, he was a Professor of Biology at the University of Michigan, Curator of Insects at prolific author of many kinds of books, including children's stories, the Museum of Zoology there, a member of the National Academy of biographical texts and practical guides, especially on horse training Sciences, and a pioneer in the study of the evolutionary basis of human (Fig. 2). behavior, died on the 20th of August 2018 at age 88. Alexander is survived by his wife of 68 years Lorraine Kearnes Alexander; his brother 1. Evolutionary entomology Noel (Donna); his daughters Susan (Sarita) and Nancy; his grand- children Morgan, Lydia, Lincoln, and Winona; and his great-grandson Alexander began his professional career in the bosom of entomology, Ezekiel; several nieces and nephews; and “young” Tom Pyle who lived with a firm focus on the inter-related fields of acoustic communication, with Alexander's family for years while growing up (Fig. 1). systematics and speciation. It was his expertise in those fields that Alexander's life is an iconic American success story. He rose from equipped him to become a major contributor to our understanding of the humble beginnings, having been born on November 18th, 1929 and evolution of cooperation in general and a leading thinker on the evolu- raised on a small, single-family farm in rural Illinois, without electricity tion of the social behavior of that “uniquely unique” species – humans. or indoor plumbing. Although his early schooling took place in a one His early studies of insect behavior convinced him that behavior and room school house, and he had no thought of attending college, his communication are clearly evolved phenomena, underlain by genes as good grades and keen intellect enabled him to attend Blackburn College certainly as morphology is, an insight he came to partly because of the and then Illinois State Normal University, obtaining his bachelor's de- utility of behavior in distinguishing species. This understanding led him gree in 1950. He served in the army during the Korean War, stationed at to realize the importance of behavior in adaptive evolution, leading Fort Knox, then went on complete a PhD in Entomology at Ohio State naturally to further insights into social evolution. The communication University in 1956. As a graduate student he pioneered the use of new systems of crickets had consequences for diversification and speciation, acoustic recording technology, developed in WWII, to study insect be- and the social behavior of insects like wasps that benefitted offspring, as havior, revolutionizing the study of acoustic communication and spe- well as more distant kin, via kin selection and mutualism – were all ciation in the process. In 1957, he was hired by the University of comprehensible in terms of Darwinian selection and likewise applicable Michigan, where he spent his career. At Michigan Alexander became a to humans (Fig. 3). national leader in evolutionary biology. His early career awards include From beginnings in evolutionary entomology, Richard Alexander the Newcomb Cleveland Prize (1961) from the American Association developed multiple, ground-breaking theories concerning the evolution for the Advancement of Science for his paper “The role of behavioral and development of key human social traits, including monogamy, ju- study in cricket classification” and the Daniel Giraud Eliot medal (1971) venile helplessness (altriciality), parental and alloparental care, incest from the National Academy of Sciences for “outstanding fundamental and cousin-marriage, cooperation in increasingly large social groups work on the systematics, evolution, and behavior of crickets.” and the associated problems of warfare, deceit and self-deception, The “nine lives” in the title of this obituary does not refer to a language and scenario-building, music and the arts, humor, religion, particular fondness for cats on Alexander's part (in fact, he was partial and even science as a human endeavor (Fig. 4). to horses), or to some series of near death experiences (although he had Alexander was instrumental in founding the multi-disciplinary some of those). Rather, it refers to the many different interests that Human Behavior and Evolution Program at the University of Michigan, Alexander pursued during his lifetime. Growing up, he worked as a alumni of which founded the Human Behavior and Evolution Society. farmer, rancher and horse trainer, activities he continued throughout Long before “sociobiology” or “evolutionary psychology” were widely his life. He was also a talented poet, songwriter, musician, and wood- discussed, Alexander had already written many profoundly influential ⁎ Corresponding author. E-mail address: [email protected] (K. Summers). https://doi.org/10.1016/j.evolhumbehav.2018.12.002 Received 5 December 2018; Accepted 11 December 2018 1090-5138/ © 2018 Elsevier Inc. All rights reserved. K. Summers et al. Evolution and Human Behavior 40 (2019) 133–139 Fig. 1. Richard D. Alexander (photo by Mark O'Brien). Fig. 4. A meeting in 1987 of several of the thinkers who were responsible for the modern unification of evolutionary and behavioral science. Left to right, in the back: David Buss, George C. Williams, Martin Daly, and Mildred Dickemann. In the front, William D. Hamilton, Napoleon Chagnon, and Richard D. Alexander. Three other people who profoundly influenced Alexander's thinking are Robert Trivers, William Irons, and Margo Wilson. Photo from richarddalexander.com maintained by David Lahti. works on evolution and social behavior. His review article entitled The Evolution of Social Behavior, published in 1974 in the Annual Review of Ecology and Systematics, has been cited over 4000 times. He continued to publish significant contributions on the evolution of social behavior on a regular basis throughout the rest of his career. A volume in his honor celebrating and summarizing many of these contributions was published in Summers and Crespi, 2013 (Human Social Evolution: The Foundational Works of Richard D. Alexander (K. Summers and B. Crespi, eds), 2013, Oxford University Press). Alexander's (1974) review of the evolution of social behavior pre- Fig. 2. Richard Alexander and friend in 2015 (photo by David Lahti). sented a comprehensive theoretical framework for understanding eu- sociality and social systems in insects, birds, and mammals. The 58 page synthetic review laid out hypotheses that a generation of biologists and anthropologists have spent their lives testing. When asked why it was so frequently cited, Alexander offered an uncharacteristically self-depre- cating answer: ‘because I got so many things wrong and every hot-shot, young scientist wants to make his reputation by showing me up’. One criticism of the review led Alexander to one of his best known discoveries, eusociality – societies with sterile workers – in the naked mole-rat: why had eusociality evolved many times among the hyme- noptera and only once in all other insects? William Hamilton had re- cently suggested that kin selection might help answer the question since hymenoptera (ants, bees, wasps) are haplodiploid which means sisters are more closely related to each other than they are to brothers or their own offspring, something that favors sisters becoming workers and rearing more sisters instead of their own young. It turned out that the factors driving the evolution of sterile workers are more complicated than simple genetic relatedness, but the insight of this finding is still important. In his 1974 review, Alexander argued that parental care (not asymmetric relatedness) was the essential ancestral character in the evolution of insect eusociality. Unlike Hamilton's haplodiploid hy- pothesis, Alexander's included the Isoptera (termites), a diploid group that evolved from ancestral wood roaches with parental care. One cri- ticism of Alexander's parental care hypothesis was that there are many Fig. 3. Cricket by R.D. Alexander. other taxa with parental care, particularly within vertebrates, where eusociality has failed to emerge. Alexander took this criticism seriously and explored it by posing the 134 K. Summers et al. Evolution and Human Behavior 40 (2019) 133–139 inaugurating the modern evolutionary study of human behavior. Alexander developed a new, synthetic approach to understanding human psychology, rooted firmly in the fresh understanding of adap- tation that had emerged in the mid-1960s thanks especially to inclusive fitness theory as developed by William D. Hamilton and subsequently applied and enriched by George C. Williams, Robert Trivers and John Maynard Smith. With such a foundation, along with a thorough study of ethnographic works, the behavioral sciences, and the lifestyles of ani- mals, Alexander drew together