1 Obituary: The Nine Lives of Richard D. Alexander 2 3 Richard D. Alexander, T.H. Hubbell Distinguished 4 University Professor of Biology at the University of 5 Michigan, Curator of Insects at the Museum of 6 Zoology there, a member of the National 7 Academy of Sciences, and a pioneer in the 8 study of the evolutionary basis of human 9 behavior, died on the 20th of August, 2018 at age 10 88. Alexander is survived by his wife of 68 years 11 Lorraine Kearnes Alexander; his brother Noel 12 (Donna); his daughters Susan (Sarita) and Nancy; 13 (Photo by Mark O’Brien) his grandchildren Morgan, Lydia, Lincoln, and 14 Winona; and his great-grandson Ezekiel; several nieces and nephews; and “young" 15 Tom Pyle who lived with Alexander’s family for years while growing up. 16 17 Alexander’s life is an iconic American success story. He rose from humble beginnings, 18 having been born on November 18th, 1929 and raised on a small, single-family farm in 19 rural Illinois, without electricity or indoor plumbing. Although his early schooling took 20 place in a one room school house, and he had no thought of attending college, his good 21 grades and keen intellect enabled him to attend Blackburn College and then Illinois 22 State Normal University, obtaining his bachelor’s degree in 1950. He served in the army 23 during the Korean War, stationed at Fort Knox, then went on complete a PhD in 1 24 Entomology at Ohio State University in 1956. As a graduate student he pioneered the 25 use of new acoustic recording technology developed in WWII, to study insect behavior, 26 revolutionizing the study of acoustic communication and speciation in the process. In 27 1957, he was hired by the University of Michigan, where he spent his career. At 28 Michigan Alexander became a national leader in evolutionary biology. His early career 29 awards include the Newcomb Cleveland Prize (1961) from the American Association for 30 the Advancement of Science for his paper "The role of behavioral study in cricket 31 classification” and the Daniel Giraud Eliot medal (1971) from the National Academy of 32 Sciences for “outstanding fundamental work on the systematics, evolution, and behavior 33 of crickets." 34 35 Richard Alexander and friend in 2015 (photo by David Lahti) 2 36 The "nine lives" in the title of this obituary does not refer to a particular fondness for cats 37 on Alexander’s part (in fact, he was partial to horses), or to some series of near death 38 experiences (although he had some of those). Rather, it refers to the many different 39 interests that Alexander pursued during his lifetime. Growing up, he worked as a farmer, 40 rancher and horse trainer, activities he continued throughout his life. He was also a 41 talented poet, songwriter, musician, and wood-carver. In addition to his professional 42 scientific writings, he was a prolific author of many kinds of books, including children’s 43 stories, biographical texts and practical guides, especially on horse training. 44 45 Evolutionary Entomology 46 Alexander began his professional career in the bosom of 47 entomology, with a firm focus on the inter-related fields of 48 acoustic communication, systematics and speciation. It 49 was his expertise in those fields that equipped him to 50 become a major contributor to our understanding of the 51 (cricket by R.D. Alexander) evolution of cooperation in general and a leading thinker on 52 the evolution of the social behavior of that “uniquely unique” species -- humans. His 53 early studies of insect behavior convinced him that behavior and communication are 54 clearly evolved phenomena, underlain by genes as certainly as morphology is, an 55 insight he came to partly because of the utility of behavior in distinguishing species. 56 This understanding led him to realize the importance of behavior in adaptive evolution, 57 leading naturally to further insights into social evolution. The communication systems of 58 crickets had consequences for diversification and speciation, and the social behavior of 3 59 insects like wasps that benefitted offspring, as well as more distant kin, via kin selection 60 and mutualism – were all comprehensible in terms of Darwinian selection and likewise 61 applicable to humans. 62 63 A meeting in 1987 of several of the thinkers who were responsible for the modern unification of 64 evolutionary and behavioral science. Left to right, in the back: David Buss, George C. Williams, Martin 65 Daly, and Mildred Dickemann. In the front, William D. Hamilton, Napoleon Chagnon, and Richard D. 66 Alexander. Three other people who profoundly influenced Alexander’s thinking are Robert Trivers, William 67 Irons, and Margo Wilson. Photo from richarddalexander.com maintained by David Lahti. 68 69 From beginnings in evolutionary entomology, Richard Alexander developed multiple, 70 ground-breaking theories concerning the evolution and development of key human 71 social traits, including monogamy, juvenile helplessness (altriciality), parental and 72 alloparental care, incest and cousin-marriage, cooperation in increasingly large social 4 73 groups and the associated problems of warfare, deceit and self-deception, language 74 and scenario-building, music and the arts, humor, religion, and even science as a 75 human endeavor. 76 77 Alexander was instrumental in founding the multi-disciplinary Human Behavior and 78 Evolution Program at the University of Michigan, alumni of which founded the Human 79 Behavior and Evolution Society. Long before "sociobiology" or “evolutionary 80 psychology” were widely discussed, Alexander had already written many profoundly 81 influential works on evolution and social behavior. His review article entitled “The 82 Evolution of Social Behavior”, published in 1974 in the “Annual Review of Ecology and 83 Systematics,” has been cited over 4000 times. He continued to publish significant 84 contributions on the evolution of social behavior on a regular basis throughout the rest 85 of his career. A volume in his honor celebrating and summarizing many of these 86 contributions was published in 2013 (Human Social Evolution: The Foundational Works 87 of Richard D. Alexander (K. Summers and B. Crespi, eds), 2013, Oxford University 88 Press). 89 90 Alexander’s (1974) review of the evolution of social behavior presented a 91 comprehensive theoretical framework for understanding eusociality and social systems 92 in insects, birds, and mammals. The 58 page synthetic review laid out hypotheses that 93 a generation of biologists and anthropologists have spent their lives testing. When 94 asked why it was so frequently cited, Alexander offered an uncharacteristically self- 5 95 deprecating answer: ‘because I got so many things wrong and every hot-shot, young 96 scientist wants to make his reputation by showing me up’. 97 98 One criticism of the review led Alexander to one of his best known discoveries, 99 eusociality – societies with sterile workers -- in the naked mole-rat: why had eusociality 100 evolved many times among the hymenoptera and only once in all other insects? 101 William Hamilton had recently suggested that kin selection might help answer the 102 question since hymenoptera (ants, bees, wasps) are haplodiploid which means sisters 103 are more closely related to each other than they are to brothers or their own offspring, 104 something that favors sisters becoming workers and rearing more sisters instead of 105 their own young. It turned out that the factors driving the evolution of sterile workers are 106 more complicated than simple genetic relatedness, but the insight of this finding is still 107 important. In his 1974 review, Alexander argued that parental care (not asymmetric 108 relatedness) was the essential ancestral character in the evolution of insect eusociality. 109 Unlike Hamilton’s haplodiploid hypothesis, Alexander’s included the Isoptera (termites), 110 a diploid group that evolved from ancestral wood roaches with parental care. One 111 criticism of Alexander’s parental care hypothesis was that there are many other taxa 112 with parental care, particularly within vertebrates, where eusociality has failed to 113 emerge. 114 115 Alexander took this criticism seriously and explored it by posing the related question: if 116 there were a eusocial vertebrate, what would it be like? He speculated that such a 117 vertebrate would have certain characteristics and suggested a hypothetical 6 118 subterranean rodent, living in the African savannah (associated with patchily distributed 119 plants with large storage root storage organs), which fed on these tubers and suffered 120 predation by snakes. Each trait followed from a deep understanding of life history 121 constraints and tradeoffs. Underground burrows would allow expansion for large 122 colonies; roots would allow low risk foraging; snakes would require self-sacrifice to 123 defend a colony. After listening to Alexander present this hypothetical eusocial 124 vertebrate at a seminar at Northern Arizona University, mammologist Terry Vaughn 125 pointed out that it sounded very much like the naked mole-rat and, soon after, 126 Alexander went to Kenya to collect a live colony for observation. This led to an 127 explosion of scientific research on this odd-looking 128 mammal, culminating in the 1990 volume, “The Biology 129 of the Naked Mole Rat” (Sherman, Jarvis and Alexander 130 1990). Like the 1845 discovery of the planet Neptune by 131 Adams and Leverrier, Alexander explored evolutionary 132 space by extrapolation, with a deep understanding of the 133 forces that shape social behavior, and was able to 134 predict eusociality in a taxon where it had not yet been 135 directly observed. 136 137 Human Evolution 138 Exactly one hundred years after Darwin published his great work on The Descent of 139 Man (1871), Alexander published his first paper on the same topic, entitled "The Search 140 for an Evolutionary Philosophy of Man" (Alexander 1971), sketching a quest that would 7 141 occupy most of his attention for the remainder of his career.