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Home , Bret Weinstein, David Buss, Margo Wilson, Mark Flinn, Martin Daly (professor), Robert Trivers

and Human Behavior 40 (2019) 133–139

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Evolution and Human Behavior

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Obituary: The nine lives of Richard D. Alexander T ⁎ Kyle Summersa, , David Lahtib, Stanton Braudec, Beverly Strassmannd, Joan Strassmannc a Department of , East Carolina University, Greenville, NC 27858, b Department of Biology, Queens College, City University of New York, Flushing, NY, 11367, United States c Department of Biology, University in St. Louis, St. Louis, MO 63130, United States d Department of Anthropology, , 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 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 . 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 “” or “evolutionary ” 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: , George C. Williams, , and Mildred Dickemann. In the front, William D. Hamilton, , and Richard D. Alexander. Three other people who profoundly influenced Alexander's thinking are , William Irons, and . 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 disparate lines of logic and evidence and introduced new ones to construct what would eventually become a comprehensive explanation of human behavior and life history that was based on an unwavering emphasis on the power of . His approach fully respected genes, development, and environment as interacting partners. The theory eventually incorporated plasticity and learning while regarding behaviors as evolving traits like any others. Most impressively, it managed to integrate a vast array of human social traits – with special reference to those that were unique or rare among animals– into a compelling explanatory structure that was internally consistent, mutually reinforcing, and empirically sound. He continued to develop these ideas throughout the rest of his life. One theme of Alexander's work dealt with the suppression of conflicts within groups. This work was strongly influenced by Egbert Leigh's (1971, 1977) publications on the evolution of Mendelian segregation and “fair meiosis”, and the concept of the “parliament of genes” evolved to suppress rogue driving elements in the genome. Alexander and Borgia (1978) made novel contributions to this topic, and he was able to see the relevance of this theory to the evolution of “reproductive leveling mechanisms” in a general sense, and particularly to human groups and polities. Alexander was also influenced by reading the moral philosophers, particularly Rawls' (1971) famous theory of justice, in which impartial justice is developed from the assumption of a “veil of ignorance” by those who write the laws. Alexander was able to unite Fig. 5. Cover image of the edited volume: Biology of the Naked Mole Rat. these disparate intellectual threads to develop a general theory of cooperation through suppression of intragroup conflict (Alexander, related question: if there were a eusocial vertebrate, what would it be 1987). This theory has been formalized and further developed, like? He speculated that such a vertebrate would have certain char- then applied to many areas of evolutionary biology, such as sperm acteristics and suggested a hypothetical subterranean rodent, living in competition (Frank, 2013). Alexander's theory of cooperation through the African savannah (associated with patchily distributed plants with suppression of within-group conflict is a major part of our understanding large storage root storage organs), which fed on these tubers and suf- of cooperation, and especially human cooperation (Frank, 2013). fered predation by snakes. Each trait followed from a deep under- Another key contribution of Alexander's work, related to his ideas standing of life history constraints and tradeoffs. Underground burrows concerning the suppression of conflict in groups, was the recognition of would allow expansion for large colonies; roots would allow low risk the importance of intergroup aggression and competition in human foraging; snakes would require self-sacrifice to defend a colony. After evolution, a topic originally broached by Darwin (1871). Alexander listening to Alexander present this hypothetical eusocial vertebrate at a proposed that hominids became so ecologically dominant that compe- seminar at Northern Arizona University, mammologist Terry Vaughn tition with other groups of humans became a crucial factor influencing pointed out that it sounded very much like the naked mole-rat and, survival and reproductive success. Starting in the late 1960's, Alexander soon after, Alexander went to Kenya to collect a live colony for ob- (e.g. Alexander & Tinkle, 1968) argued that intergroup aggression was servation. This led to an explosion of scientific research on this odd- likely to have favored the evolution of complex human cooperation and looking mammal, culminating in the 1991 volume, The Biology of the extreme intelligence, traits that are far more advanced in humans than Naked Mole Rat (Sherman, Jarvis, & Alexander, 1991). Like the 1845 in other primates. This became known as the “Balance of Power” hy- discovery of the planet Neptune by Adams and Leverrier, Alexander pothesis: “The general hypothesis that I support to account for the explored evolutionary space by extrapolation, with a deep under- maintenance and elaboration of group-living and complex sociality in standing of the forces that shape social behavior, and was able to pre- humans… derives from a theme attributable to Darwin (1871) and dict eusociality in a taxon where it had not yet been directly observed Keith (1949), and developed by a succession of more recent authors… It (Fig. 5). includes group-against-group, within species competition as a central driving force, leading to balance-of-power races with a positive feed- back upon cooperative abilities and social complexity. It implies that 2. the only plausible way to account for the striking departure of humans from their predecessors and all other species with respect to mental and Exactly one hundred years after Darwin published his great work on social attributes is to assume that humans uniquely became their own The Descent of Man (1871), Alexander published his first paper on the principle hostile force of nature” (Alexander, 1989). The importance of same topic, entitled The Search for an Evolutionary Philosophy of Man intergroup competition in the evolution cooperation and intelligence (Alexander, 1971), sketching a quest that would occupy most of his has become a major theme in the scientific literature, thanks in large attention for the remainder of his career. Because of that paper, and part to Alexander's contributions. Robert Trivers' monumental first publication The Evolution of Reciprocal Alexander's ideas about intelligence were also strongly influenced (Trivers, 1971), the year 1971 can reasonably be hailed as by Humphrey's theory on the paramount importance of social

135 K. Summers et al. Evolution and Human Behavior 40 (2019) 133–139 interactions in the evolution of intelligence (Humphrey, 1976). But monogamy”, where harsh environmental conditions make it impossible Alexander recognized that the evolution of extreme intelligence in the for males to monopolize more than a single female. human lineage depended on the multilevel nature of human interac- Of course, human history reveals episodes and societies character- tions within hierarchical groups (and groups within groups), particu- ized by extreme inequality in reproductive opportunity (Alexander, larly in the context of intense intergroup competition. This led to open- 1979). Nevertheless, Alexander thought that, for the majority of hu- ended coevolutionary arms races of strategizing and counter-strate- mans that have lived across the course of human history, most have had gizing (requiring continuously expanding cognitive capacities), both to some expectation of opportunities to mate and produce their own off- succeed in intergroup competition with other groups of strategizing spring. He further suggested (e.g. Alexander, 1987) that trends towards humans, and to successfully negotiate advancement in the increasingly neolocal nuclear human families and associated large-scale patterns of complex multilevel hierarchies comprising human groups (from small reproductive opportunity-leveling in modern times have been propelled multifamily groups, to nomadic bands, to semi-permanent groups, to by the continued (and enhanced) importance of inter-group competi- villages, to village coalitions, to tribes, and on up to nation states). By tion and aggression (on a broad scale), and consequent pressure for combining his ideas about the effects of intergroup aggression with rulers and elites to concede reproductive opportunities to those lower in those on the key importance of intra-group social interactions on the the hierarchy. Subsequent comparative work has supported that claim, evolution of intelligence, he was able to provide a cohesive and com- and has brought to light additional factors that are likely to be involved. prehensive explanatory framework that led to many new insights into Another important contribution was Alexander's development of the the evolution of human cognitive capacities (Alexander, 1989, 1990a, theory of indirect reciprocity. In 1971, Robert Trivers published his 1990b). Alexander realized that these evolving cognitive capacities classic paper (Trivers, 1971) on “”, which elaborated provide the key to understanding our most cherished abilities and tra- how reciprocal exchange of benefits could increase the fitness of both ditions, including the evolution of language and “scenario-building” parties in an interaction, given certain assumptions, such as the re- (the intellectual ability to imagine situations, analyzing them “in the ciprocally asymmetric value of exchanged goods and services (often mind's eye”) and the development of culture, religion, art and science. created by specialization in complex societies), and repeated interac- In parallel with these ideas, Alexander developed increasingly so- tions between the same individuals, with indefinite end points. This phisticated ideas about the evolution of human family structure and theory profoundly influenced Alexander's thinking (and that of many other uniquely human traits. He began from a comparative perspective, others), but he realized that it was insufficient to explain the kind of evaluating evidence regarding the ancestral state of hominid family large scale levels of cooperation seen in human groups and societies structure and then trying to infer what factors could have driven the involving multitudes of participants and interactions. Expanding on broad swath of uniquely human traits, including not only such com- Trivers' concept of “generalized reciprocity”, Alexander developed the monly discussed phenomena as consciousness, foresight, tool-use, lan- theory of indirect reciprocity, which posited that in contrast to direct guage and symbolic thought, but also less often discussed traits such as reciprocity “in indirect reciprocity, the return is expected from someone monogamy, paternal care, extended childhood, relative hairlessness, other than the recipient of the beneficence. The return can come from and concealed ovulation, among many others. essentially any individual or collection of individuals in the group” Alexander and Noonan (1979) realized that distinctive traits like (Alexander, 1987). Alexander began writing about indirect reciprocity concealed ovulation were likely to have played a key role in the evo- in the 1970's, but his ideas were largely overlooked as theoreticians lution of the family centered patterns of cooperation in human socie- focused on the mechanisms and implications of direct reciprocity. ties, in which every human retains an interest in having the opportunity However, in the 1990's, theoreticians began exploring the mechanisms to reproduce. They had the critical perception that concealed ovulation and implications of Alexander's concept of indirect reciprocity in and related traits were adaptations that enhanced the stability and earnest, and this led to confirmation of the potential importance of this permanence of human pair bonds, allowing for the increased invest- mechanism in the evolution of wide-scale cooperation in large societies ment required to produce successful offspring in an increasingly com- (reviewed by Sigmund, 2013). The key role of indirect reciprocity is petitive social environment. They argued that females could develop now a central theme of both theoretical and empirical work on the and maintain consortships with specific males in multimale/multi- evolution of cooperation in large human societies. female groups, in part through the evolution of concealed ovulation (favoring long-term association by the male to ensure paternity). These 3. Culture relationships then provided high confidence of paternity – and a direct payoff to males investing in (their own) offspring. They benefitted fe- Multifarious lines of reasoning were combined by Alexander into a males through investment by their male partner, including protection comprehensive, yet subtle and nuanced view of evolution and human from aggression or infanticide by other males. Alexander thought that behavior. One of the challenges in reading Alexander's work is precisely the extreme altriciality of human infants and children must have the heavily integrated nature of its framework; one can read some piece evolved in conjunction with the prodigious learning capacities required of his on human ecological dominance, or our mating strategies and to be a successful adult in an increasingly complex social world. He marriage, or concealment of ovulation, or lifelong parenting, extensive argued that this complexity was likely to be positively reinforcing in a and differential nepotism, direct and indirect reciprocity, in-group/out- social runaway process (Alexander, 1990a). group dynamics, extended juvenile period, our mind as a social tool, Alexander and Noonan (1979) originally proposed that dominant scenario-building, consciousness, self-deception, learning, culture, males would be the first to form pair bonds, but later work indicated morality, law, political structures, humor, the arts, or religion… but that this hypothesis was more likely to apply to subordinate males, understanding the function and relevance of any one of these in given that dominant males have higher incentives to pursue polygynous Alexander's thought requires an understanding of many others. The flip mating strategies rather than investing in parental care (reviewed in side of this, of course, is that as more of these elements are understood, Strassmann, 2013). Yet, why would dominant males not break up ex- the more that a big picture begins to emerge, and the more easily ad- clusive associations between subordinate males and females? As the ditional pieces fit into place. importance of inter-group competition increased in the human lineage, Alexander was careful to consider and include the complex social the value of subordinates to dominants within groups also increased, and cognitive abilities of humans in the development of this theories. placing constraints on the levels of despotism and reproductive skew He recognized the importance of developmental, behavioral, and cul- that dominants could afford to impose (Alexander, 1979). Alexander tural plasticity in human behavior and especially, the role of the en- (1979) coined the term “Socially Imposed Monogamy” (SIM) to capture vironment during human development. One of Alexander's key con- this dynamic, which he contrasted to “ecologically imposed tributions was his early appreciation of how people could behave

136 K. Summers et al. Evolution and Human Behavior 40 (2019) 133–139 adaptively without having specific genes “for” particular behaviors. practice what we preach? We claim the equal value of all people re- This understanding involved his realization that genetic variants under gardless of their relation to us, we promulgate the Golden Rule, we strong natural selection would tend to go to fixation, and yet could teach our kids to be good regardless of the possibility of being caught… work through plastic developmental and psychological mechanisms to and yet nobody truly acts as though we believe these things. We still endow their owners with the adaptive flexibility to do the right thing privilege our own groups, we place ourselves and our people far above for reproduction in the various environmental circumstances en- others in our practical estimations, and we are inordinately concerned countered. As Alexander (1990b) wrote: “Adaptation is not restricted to for reputation and how our actions appear to observers. Thus a work- situations in which genes program specifically for particular behavioral able evolutionary theory of morality has to explain not only the ideals alternatives: natural selection of alternative alleles may also yield we tend to espouse but also our typical failure to abide by them. abilities and tendencies to engage in conditional strategies, to assess A brief synopsis of Alexander's theory of morality (drawing on the costs and benefits in directly or indirectly reproductive terms. Inhu- insights discussed above) is as follows: early humans achieved ecolo- mans, such cost-benefit assessments may be conducted entirely through gical dominance, to such an extent that the most significant selective mental scenario-building, or even through absorbing and judging the agents impinging on their survival and reproduction were other hu- mental scenarios of others, without either admission or cognizance of mans. As humans lived in social groups, competition would manifest the reproductive significance of the assessment.” itself not only within a group but between groups. This would further Alexander's views of human behavior gave him a unique perspective intensify our dependence on groups, and create a ratcheting effect on the evolution of the human psyche. Well before disagreements un- where traits contributory to group cohesion would be increasingly fa- folded between evolutionary psychologists, evolutionary anthro- vored. In this situation, individuals could benefit themselves and their pologists and gene-culture coevolutionists, Alexander had developed kin reproductively in two different ways: directly by competing with deep insights into the extreme flexibility of the human mind and the others within their groups, and indirectly by helping others in their implications for . Alexander saw human cognitive groups, which in turn would strengthen the group in competition with capacities, including intelligence, consciousness, foresight, empathy, other groups. Thus we evolved to cooperate in order to compete, both theory of mind, moral reasoning, language, abilities such as scenario- within and between social groups. The result was the primacy of in- building and story-telling, and cultural legacies such as art and religion, dividual social reputation, and a heavily scenario-building, socially as stemming from continual arms races of social competition intuitive, and strategizing human personality. Each individual would (Alexander, 2006; Flinn & Alexander, 2007). He emphasized the in- balance the direct and indirect means of garnering benefits. Each in- terplay of cooperation and competition between individuals and dividual would also benefit more by the cooperation of others thanby groups. The cross-generational, cumulative effects of human re- cooperating oneself, resulting in a few powerful items entering into the productive striving, as collected and codified in cultural traditions and human strategic toolbox: tendencies to over-encourage beneficence in trends, have perennially created new environments in which human others, to overestimate one's own level of commitment, to look for strategizing and social competition has unfolded. violations in others and conceal them in oneself, and in general to Alexander appreciated the likely importance of “Darwinian algo- dispense kindness strategically depending on the expected returns to rithms” (psychological mechanisms designed to accomplish adaptive oneself and one's kin. Moreover, these objectives could often be un- ends), in biasing human perception and behavior in (historically) dertaken with greater efficiency if the motivating factors were con- adaptive directions, as appreciated by evolutionary psychologists cealed from the consciousness not only of others but of the agents (Alexander 1990b). But his keen awareness of the value of novelty and themselves. All of these strategies and others presuppose conflicts of creativity in the context of social arms races led him to be cautious interest among genetically different individuals. Moral systems are es- about proposing limits to the power of the human mind to adapt to sentially the means by which these conflicts are identified and resolved. novel circumstances. His deep understanding of the pervasive influence Reception to these ideas was (and has continued to be) uneven, for of natural selection on human conscious and unconscious thought and several reasons, some of which remain relevant even for those who emotion (and attendant strategies of deceit and self-deception) led him understand evolution and selection. First, the ideas are intellectually to be skeptical concerning cultural evolution as a process independent demanding, and require substantial investment in order for one to un- of organic evolution, untethered from the adaptive strategies of human derstand them and avoid hyperbole, simplification, or an imbalance of beings (Alexander, 1979). Instead, Alexander viewed cultural evolution components. Often commentators have taken the easier route of as intimately and inextricably linked to individual reproductive lumping Alexander's work with facile treatments of these problems, that striving. pay mere lip service to kin selection, reciprocal altruism, and other subjects whose complicated nature is discussed and documented by 4. Morality Alexander. As Alexander was developing and publishing his ideas, es- sentially no rival well-developed evolutionary theory of morality ex- The faculty and institution of morality was a chief and overriding isted that was placed fully into the context of human life history, and interest during all of these pursuits in Alexander's quest to understand that took seriously the real moral experience of people, warts and all. humanity. Morality was the human trait he contemplated most ex- Many people, including those who depict humans as as fundamentally tensively, and after Darwinism and Human Affairs (1979) was the sole good, prefer to overlook those warts rather than attempt to explain human affair he chose to investigate further in a book of itsown, The them. Many are discomforted by the picture of a strategizing human Biology of Moral Systems (1987). In considering humans as a product of who has not thoroughly abandoned the organismal tendency towards natural selection, Alexander found in moral exhortations to selfless selfishness. Many are additionally disgusted at the idea of self-decep- beneficence a fascinating professional challenge along the same linesas tion. To this, Alexander might first offer the olive branch that self- that posed to Darwin by the presence of sterile castes in social insects. ishness and self-deception are used in an evolutionary sense, and on Alexander started from a hypothesis that would be uncontroversial to that meaning neither deception nor selfishness are expected always to evolutionary biologists with respect to the traits of any other organism, be conscious. Strangely enough, “self-deception” actually rescues the but got him maligned and misunderstood by many when he suggested it possibility of authentic moral action by separating evolved objectives of morality: that it is “reproductively selfish”. Even human moral atti- from consciousness. But Alexander would never let that molasses go tudes and behaviors that seem so other-regarding, so selfless, tended to down without the medicine, for the objection to evolutionary self- return benefits to the actor. The theory would also have to address some ishness is usually symptomatic of a general human trait that frustrates very peculiar features of humans– things Alexander called the moral efforts like Alexander's to understand ourselves: our natural resistance paradoxes, the convolutedness of morality. For instance, why do we not to honest self-study. We tend to avoid being confronted with our typical

137 K. Summers et al. Evolution and Human Behavior 40 (2019) 133–139 attitudes and actions and their reproductive functions. Alexander in- large set of interacting hypotheses, consistent within themselves and sisted that we must investigate and accept whatever we find to underlie among each other, and able to explain many aspects of the human our thoughts and deeds, both the noble and the disgusting, as this condition. His excitement in discovery and his tireless critical thinking, knowledge is our only hope if we want to improve ourselves and the sustained over a period of so many years, was shared openly with his world. students and we all benefitted from it. One wrote, speaking for all, that Both the depth and iconoclasm in Alexander's ideas on morality can we were lucky to have a seat at the table. be illustrated in one of his favorite stories. Theodore H. Hubbell, a Alexander was a great teacher; his students have dispersed across friend and colleague at the UMMZ, was troubled by the contention that academia and beyond. Over his long and distinguished career he human behavior is by and large self-interested. Once he thought he had mentored many graduate and undergraduate students, including a good counter-example, and approached Alexander: “Dick, this Kenneth C. Shaw of Iowa State University, Mary Jane West Eberhard of morning on my way to work I saw a caterpillar moving across the the Smithsonian Tropical Research Institute, Daniel Otte of the sidewalk. I knew it had fallen off its host plant, so I picked it upand Academy of Natural Sciences in Philadelphia, Mitchell Weiss of Rutgers replaced it on the proper plant species. Was that not an act of pure University, Jon Waage of Brown University, Ann Pace of the Bell altruism?” To this Alexander replied, “It may have been, until you told Museum of Natural History at the University of Minnesota, Harry Power me about it.” of , Paul W. Sherman of Cornell University, John L. Despite Alexander's evolutionary analysis of morality, and despite Hoogland of the Appalachian Environmental Laboratory at the his consistent refusal to resort to what he called “the transcendent”, University of Maryland, Richard D. Howard of Purdue University, Alexander never exhibited any signs of retreating into the moral cyni- Marianne N. Feaver of North Carolina State University, Gerald Borgia of cism or nihilism that is often associated with “” (a the University of Maryland, Katherine M. Noonan of Albany, , term he avoided). In fact, he struggled with increasing fervor to solve David Foltz of Louisiana State University, Cynthia Kagarise Sherman of what he called the world's most terrible puzzle: our “unholy capacity Ithaca, New York, Joan Strassmann of Washington University, Nancy and willingness to turn close-knit groups into fighting machines” Moran of the University of Texas, Marlene Zuk of the University of (Alexander, 2014, p. 58), a tendency he well knew was a product of the Minnesota, David Queller of Washington University, Alex Mintzer of mechanism that had fashioned our moral psychology in the first place. Texas A&M University, Bernard J. Crespi of Simon Fraser University, Whereas in the 1980s and 1990s he would end his papers on morality Richard Connor of the University of Massachusetts at Dartmouth, with the general idea that understanding ourselves and our biases is Beverly Strassmann of the University of Michigan, Steven Frank of the half the battle, by the 2010s his primary goal was to nail specific so- University of California at Irvine, Stanton Braude of Washington lutions to the “reciprocating echoes of intra-group amity and inter- University, Eileen Lacey of the University of California at Berkeley, Kyle group enmity” (Alexander, 2014, p. 56). When he sat down to write his Summers of East Carolina University, Andrew F. Richards of the City final professional papers in his 80s, he focused not on evolutionary College of New York, John Pepper of the National Institutes of Health, explanation as an end in itself, but rather insofar as it might contribute Deborah Cizek of Boulder Colorado, David Marshall and John Cooley of to “global harmony” (2013) and “the future of human society” (2014). the University of Connecticut, Anna Bess Sorin of the University of He considered the central condition for the survival of humanity to be Memphis, and Heather Heying of the Aspen Center for the dissolution of the evolved concept of “the other” as it manifests as a Human Development, Daniel Kruger of the University of Michigan, and motivator of hatred towards the outgroup. Hence, Alexander (2013, p. David Lahti of the City University of New York. Alexander also men- 417) wrote: “The greatest difficulty in seeking global harmony may tored many postdoctoral scholars, including James Lloyd of the derive from human groups targeting one another. Humans alone – University of Florida, Jasper J. Loftus-Hills of the University of among all the world's species – plot, plan, and organize massive con- Michigan, Daniel Otte of the Philadelphia Academy of Natural Sciences, flicts to defeat or displace similarly organized and cooperative members Paul Turke of Turke and Tomashow Pediatrics, Laura Betzig of the of their own species. Can we learn to use the current consciousness of University of Michigan, Mark V. Flinn of Baylor University, Wendy our human background to adjust team efforts of all kinds so that hon- Orent of Emory University, Andrew F. Richards of the City University of esty, fairness, and negotiation can increase and lead us toward global New York, Rachel Smolker of the University of Vermont. harmony?” We end this obituary with an excerpt from a poem (“Heroes are For Alexander the keys to this project would be the universalization works of art”) by Alexander (2011, p. 52). He did not write this with of morality, the reduction of aggression and hyper-competitiveness, reference to himself, but we think it is appropriate. temperance with respect to views of morality and God, the conversion of despotic governments that treat their citizens as property, and the Heroes are for all times, all ages radical extrapolation of kinship and expansion of social reciprocity. He they help us to know what we issued various concrete proposals in this vein, for instance that team hadn't even known we wanted to know, sports could soften and redirect outgroup animosity. He also wondered to facilitate all that we discover whether God might be translated into a naturalistic concept based on a can be discovered, all that universal social group or extended kinship in the context of our system can be absorbed into ourselves. of morality. Nevertheless, he repeatedly expressed towards the end of his life (e.g. Alexander, 2011) that the biggest regret of his professional Acknowledgements career was that he had not made satisfactory progress towards fostering our peaceful coexistence. Thus, Alexander left us with a consistent plea, The authors gratefully acknowledge valuable advice, discussion and a take-home message from over half a century of thought: accept and commentary from Nancy Alexander, Laura Betzig, Gerald Borgia, fully interpret our evolutionary heritage; for our own sake, strive Napoleon Chagnon, Richard Connor, John Cooley, Bernard Crespi, sympathetically and constructively to understand humanity. Martin Daly, Mary Jane West Eberhard, , Steven Frank, Heather Heying, John Hoogland, William Irons, Daniel Kruger, Eileen 5. Conclusion Lacey, Bobbi Low, David Marshall, David Queller, Andrew Richards, Anna Bess Sorin, Paul Turke and Bret Weinstein. Richard Alexander did not develop his major ideas in a linear fashion. Rather, they were all developed interactively in the ferment of References his mind over the course of his life. Over those many years, he gener- ated an expansive vision of human behavioral evolution, composed of a Alexander, R. D. (1971). The search for an evolutionary philosophy of man. Proceedings of

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the Royal Society of Victoria, Melbourne, 84, 99–120. Darwin, C. (1871). The descent of man and selection in relation to sex. New York, NY: Alexander, R. D. (1974). The evolution of social behavior. Annual Review of Ecology and Appleton. Systematics, 5, 352–383. Flinn, M., & Alexander, R. D. (2007). Runaway in humans. In S. W. Alexander, R. D. (1979). Darwinism and human affairs. Seattle, WA: University of Gangestad (Ed.). The evolution of mind. Fundamental questions and controversies (pp. Washington Press. 249–255). New York: Guilford Press. Alexander, R. D. (1987). The biology of moral systems. Hawthorne, NY: Aldine de Gruyter. Frank, S. (2013). A new theory of cooperation. In K. Summers, & B. Crespi (Eds.). Human Alexander, R. D. (1989). The evolution of the human psyche. In C. Stringer, & P. Mellars social evolution: The foundational works of Richard D. Alexander. Oxford, UK: Oxford (Eds.). The human revolution (pp. 455–513). University of Edinburgh Press. University Press. Alexander, R. D. (1990a). How did humans evolve? reflections on the uniquely unique Humphrey, N. K. (1976). The social function of intellect. In P. P. G. Bateson, & R. A. Hinde species. University of Michigan Museum of Zoology, 1, 1–38 (Special Publication). (Eds.). Growing points in (pp. 303–317). Cambridge, UK: Cambridge Alexander, R. D. (1990b). Epigenetic rules and Darwinian algorithms: The adaptive study University Press. of learning and development. Ethology and Sociobiology, 11, 241–303. Keith, A. (1949). A new theory of human evolution. New York, NY: Philosophy Library. Alexander, R. D. (2006). The challenge of human social behavior. , Leigh, E. G. (1971). Adaptation and diversity. San Francisco, CA: Freeman, Cooper. 4(2), 1–28. Leigh, E. G. (1977). How does selection reconcile individual advantage with the good of Alexander, R. D. (2011). The Mockingbird's River song: Poems, essays, songs, and stories: the group. Proceedings of the National Academy of Sciences of the United States of 1946–2011. Manchester, MI: Woodlane Farm Books. America, 74, 4542–4546. Alexander, R. D. (2013). Religion, evolution and the quest for global harmony – Original Rawls, J. (1971). A theory of justice. Cambridge, MA: Press. essay for this volume. In K. Summers, & B. Crespi (Eds.). Human Social Evolution: The Sherman, P., Jarvis, J., & Alexander, R. D. (1991). The biology of the naked mole rat. Foundational Works of Richard D. Alexander. Oxford, UK: Oxford University Press. Princeton, NJ: Press. Alexander, R. D. (2014). Darwin's challenges and the future of human society. In F. Sigmund, K. (2013). Introduction: The basis of morality, Richard Alexander on indirect Wayman, P. Williamson, & B. Bueno de Mesquita (Eds.). Predicting the future in sci- reciprocity. In K. Summers, & B. Crespi (Eds.). Human social evolution: The founda- ence, economics, and politics (pp. 55–108). Ann Arbor: University of Michigan Press. tional works of Richard D. Alexander. Oxford, UK: Oxford University Press. Alexander, R. D., & Borgia, G. (1978). , altruism, and the levels of orga- Strassmann, B. (2013). Introduction: Concealed ovulation in humans: Further evidence. nization of life. Annual Review of Ecology and Systematics, 9, 449–474. In K. Summers, & B. Crespi (Eds.). Human social evolution: The foundational works of Alexander, R. D., & Noonan, K. M. (1979). Concealment of ovulation, parental care and Richard D. Alexander. Oxford, UK: Oxford University Press. human social evolution. In N. A. Chagnon, & W. G. Irons (Eds.). Evolutionary biology Summers, K., & Crespi, B. (2013). Human social evolution: The foundational works of and human social behavior: An anthropological perspective (pp. 436–453). North Richard D. Alexander. Oxford, UK: Oxford University Press. Scituate, Mass.: Duxbury Press. Trivers, R. (1971). The evolution of reciprocal altruism. The Quarterly Review of Biology, Alexander, R. D., & Tinkle, D. W. (1968). A comparative review (of on Aggression and the 46, 35–57. territorial imperative). Bioscience, 18(3), 245–248.

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