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. Because of that paper, and
142 Robert Trivers' monumental first publication "The Evolution of Reciprocal Altruism"
143 (Trivers 1971), the year 1971 can reasonably be hailed as inaugurating the modern
144 evolutionary study of human behavior. Alexander developed a new, synthetic approach
145 to understanding human psychology, rooted firmly in the fresh understanding of
146 adaptation that had emerged in the mid-1960s thanks especially to inclusive fitness
147 theory as developed by William D. Hamilton and subsequently applied and enriched by
148 George C. Williams, Robert Trivers and John Maynard Smith. With such a foundation,
149 along with a thorough study of ethnographic works, the behavioral sciences, and the
150 lifestyles of animals, Alexander drew together disparate lines of logic and evidence and
151 introduced new ones to construct what would eventually become a comprehensive
152 explanation of human behavior and life history that was based on an unwavering
153 emphasis on the power of natural selection. His approach fully respected genes,
154 development, and environment as interacting partners. The theory eventually
155 incorporated plasticity and learning while regarding behaviors as evolving traits like any
156 others. Most impressively, it managed to integrate a vast array of human social traits --
157 with special reference to those that were unique or rare among animals-- into a
158 compelling explanatory structure that was internally consistent, mutually reinforcing, and
159 empirically sound. He continued to develop these ideas throughout the rest of his life.
160 One theme of Alexander’s work dealt with the suppression of conflicts within groups.
161 This work was strongly influenced by Egbert Leigh’s (1971, 1977) publications on the
162 evolution of Mendelian segregation and “fair meiosis”, and the concept of the
163 “parliament of genes” evolved to suppress rogue driving elements in the genome.
8 164 Alexander and Borgia (1978) made novel contributions to this topic, and he was able to
165 see the relevance of this theory to the evolution of “reproductive leveling mechanisms”
166 in a general sense, and particularly to human groups and polities. Alexander was also
167 influenced by reading the moral philosophers, particularly Rawls’ (1971) famous theory
168 of justice, in which impartial justice is developed from the assumption of a “veil of
169 ignorance” by those who write the laws. Alexander was able to unite these disparate
170 intellectual threads to develop a general theory of cooperation through suppression of
171 intragroup conflict (Alexander, 1987). This theory has been formalized and further
172 developed, then applied to many areas of evolutionary biology, such as sperm
173 competition (Frank 2013). Alexander’s theory of cooperation through suppression of
174 within-group conflict is a major part of our understanding of cooperation, and especially
175 human cooperation (Frank 2013).
176
177 Another key contribution of Alexander’s work, related to his ideas concerning the
178 suppression-of-conflict in groups, was the recognition of the importance of intergroup
179 aggression and competition in human evolution, a topic originally broached by Darwin
180 (1871). Alexander proposed that hominids became so ecologically dominant that
181 competition with other groups of humans became a crucial factor influencing survival
182 and reproductive success. Starting in the late 1960’s, Alexander (e.g. Alexander and
183 Tinkle 1968) argued that intergroup aggression was likely to have favored the evolution
184 of complex human cooperation and extreme intelligence, traits that are far more
185 advanced in humans than in other primates. This became known as the “Balance of
186 Power” hypothesis: “The general hypothesis that I support to account for the
9 187 maintenance and elaboration of group-living and complex sociality in humans… derives
188 from a theme attributable to Darwin (1871) and Keith (1949), and developed by a
189 succession of more recent authors… It includes group-against-group, within species
190 competition as a central driving force, leading to balance-of-power races with a positive
191 feedback upon cooperative abilities and social complexity. It implies that the only
192 plausible way to account for the striking departure of humans from their predecessors
193 and all other species with respect to mental and social attributes is to assume that
194 humans uniquely became their own principle hostile force of nature” (Alexander 1989).
195 The importance of intergroup competition in the evolution cooperation and intelligence
196 has become a major theme in the scientific literature, thanks in large part to Alexander’s
197 contributions.
198
199 Alexander’s ideas about intelligence were also strongly influenced by Humphrey’s
200 theory on the paramount importance of social interactions in the evolution of intelligence
201 (Humphrey 1976). But Alexander recognized that the evolution of extreme intelligence
202 in the human lineage depended on the multilevel nature of human interactions within
203 hierarchical groups (and groups within groups), particularly in the context of intense
204 intergroup competition. This led to open-ended coevolutionary arms races of
205 strategizing and counter-strategizing (requiring continuously expanding cognitive
206 capacities), both to succeed in intergroup competition with other groups of strategizing
207 humans, and to successfully negotiate advancement in the increasingly complex
208 multilevel hierarchies comprising human groups (from small multifamily groups, to
209 nomadic bands, to semi-permanent groups, to villages, to village coalitions, to tribes,
10 210 and on up to nation states). By combining his ideas about the effects of intergroup
211 aggression with those on the key importance of intra-group social interactions on the
212 evolution of intelligence, he was able to provide a cohesive and comprehensive
213 explanatory framework that led to many new insights into the evolution of human
214 cognitive capacities (Alexander 1989, 1990). Alexander realized that these evolving
215 cognitive capacities provide the key to understanding our most cherished abilities and
216 traditions, including the evolution of language and “scenario-building” (the intellectual
217 ability to imagine situations, analyzing them “in the mind’s eye”) and the development of
218 culture, religion, art and science.
219
220 In parallel with these ideas, Alexander developed increasingly sophisticated ideas about
221 the evolution of human family structure and other uniquely human traits. He began from
222 a comparative perspective, evaluating evidence regarding the ancestral state of hominid
223 family structure and then trying to infer what factors could have driven the broad swath
224 of uniquely human traits, including not only such commonly discussed phenomena as
225 consciousness, foresight, tool-use, language and symbolic thought, but also less often
226 discussed traits such as monogamy, paternal care, extended childhood, relative
227 hairlessness, and concealed ovulation, among many others.
228
229 Alexander and Noonan (1979) realized that distinctive traits like concealed ovulation
230 were likely to have played a key role in the evolution of the family centered patterns of
231 cooperation in human societies, in which every human retains an interest in having the
232 opportunity to reproduce. They had the critical perception that concealed ovulation and
11 233 related traits were adaptations that enhanced the stability and permanence of human
234 pair bonds, allowing for the increased investment required to produce successful
235 offspring in an increasingly competitive social environment. They argued that females
236 could develop and maintain consortships with specific males in multimale/multifemale
237 groups, in part through the evolution of concealed ovulation (favoring long-term
238 association by the male to ensure paternity). These relationships then provided high
239 confidence of paternity – and a direct payoff to males investing in (their own) offspring.
240 They benefitted females through investment by their male partner, including protection
241 from aggression or infanticide by other males. Alexander thought that the extreme
242 altriciality of human infants and children must have evolved in conjunction with the
243 prodigious learning capacities required to be a successful adult in an increasingly
244 complex social world. He argued that this complexity was likely to be positively
245 reinforcing in a social runaway process (Alexander 1990a)
246
247 Alexander and Noonan (1979) originally proposed that dominant males would be the
248 first to form pair bonds, but later work indicated that this hypothesis was more likely to
249 apply to subordinate males, given that dominant males have higher incentives to pursue
250 polygynous mating strategies rather than investing in parental care (reviewed in
251 Strassmann 2013). Yet, why would dominant males not break up exclusive associations
252 between subordinate males and females? As the importance of inter-group competition
253 increased in the human lineage, the value of subordinates to dominants within groups
254 also increased, placing constraints on the levels of despotism and reproductive skew
255 that dominants could afford to impose (Alexander 1979). Alexander (1979) coined the
12 256 term “Socially Imposed Monogamy” (SIM) to capture this dynamic, which he contrasted
257 to “ecologically imposed monogamy”, where harsh environmental conditions make it
258 impossible for males to monopolize more than a single female.
259
260 Of course, human history reveals episodes and societies characterized by extreme
261 inequality in reproductive opportunity (Alexander, 1979). Nevertheless, Alexander
262 thought that, for the majority of humans that have lived across the course of human
263 history, most have had some expectation of opportunities to mate and produce their
264 own offspring. He further suggested (e.g. Alexander 1987) that trends toward neolocal
265 nuclear human families and associated large-scale patterns of reproductive opportunity-
266 leveling in modern times have been propelled by the continued (and enhanced)
267 importance of inter-group competition and aggression (on a broad scale), and
268 consequent pressure for rulers and elites to concede reproductive opportunities to those
269 lower in the hierarchy. Subsequent comparative work has supported that claim, and has
270 brought to light additional factors that are likely to be involved.
271
272 Another important contribution was Alexander’s development of the theory of indirect
273 reciprocity. In 1971, Robert Trivers published his classic paper (Trivers 1971) on
274 “reciprocal altruism”, which elaborated how reciprocal exchange of benefits could
275 increase the fitness of both parties in an interaction, given certain assumptions, such as
276 the reciprocally asymmetric value of exchanged goods and services (often created by
277 specialization in complex societies), and repeated interactions between the same
278 individuals, with indefinite end points. This theory profoundly influenced Alexander’s
13 279 thinking (and that of many others), but he realized that it was insufficient to explain the
280 kind of large scale levels of cooperation seen in human groups and societies involving
281 multitudes of participants and interactions. Expanding on Trivers’ concept of
282 “generalized reciprocity”, Alexander developed the theory of indirect reciprocity, which
283 posited that in contrast to direct reciprocity “in indirect reciprocity, the return is expected
284 from someone other than the recipient of the beneficence. The return can come from
285 essentially any individual or collection of individuals in the group” (Alexander 1987).
286 Alexander began writing about indirect reciprocity in the 1970’s, but his ideas were
287 largely overlooked as theoreticians focused on the mechanisms and implications of
288 direct reciprocity. However, in the 1990’s, theoreticians began exploring the
289 mechanisms and implications of Alexander’s concept of indirect reciprocity in earnest,
290 and this led to confirmation of the potential importance of this mechanism in the
291 evolution of wide-scale cooperation in large societies (reviewed by Sigmund 2013). The
292 key role of indirect reciprocity is now a central theme of both theoretical and empirical
293 work on the evolution of cooperation in large human societies.
294
295 Culture
296 Multifarious lines of reasoning were combined by Alexander into a comprehensive, yet
297 subtle and nuanced view of evolution and human behavior. One of the challenges in
298 reading Alexander's work is precisely the heavily integrated nature of its framework; one
299 can read some piece of his on human ecological dominance, or our mating strategies
300 and marriage, or concealment of ovulation, or lifelong parenting, extensive and
301 differential nepotism, direct and indirect reciprocity, in-group/out-group dynamics,
14 302 extended juvenile period, our mind as a social tool, scenario-building, consciousness,
303 self-deception, learning, culture, morality, law, political structures, humor, the arts, or
304 religion... but understanding the function and relevance of any one of these in
305 Alexander's thought requires an understanding of many others. The flip side of this, of
306 course, is that as more of these elements are understood, the more that a big picture
307 begins to emerge, and the more easily additional pieces fit into place.
308
309 Alexander was careful to consider and include the complex social and cognitive abilities
310 of humans in the development of this theories. He recognized the importance of
311 developmental, behavioral, and cultural plasticity in human behavior and especially, the
312 role of the environment during human development. One of Alexander’s key
313 contributions was his early appreciation of how people could behave adaptively without
314 having specific genes “for” particular behaviors. This understanding involved his
315 realization that genetic variants under strong natural selection would tend to go to
316 fixation, and yet could work through plastic developmental and psychological
317 mechanisms to endow their owners with the adaptive flexibility to do the right thing for
318 reproduction in the various environmental circumstances encountered. As Alexander
319 (1990b) wrote: “Adaptation is not restricted to situations in which genes program
320 specifically for particular behavioral alternatives: natural selection of alternative alleles
321 may also yield abilities and tendencies to engage in conditional strategies, to assess
322 costs and benefits in directly or indirectly reproductive terms. In humans, such cost-
323 benefit assessments may be conducted entirely through mental scenario-building, or
15 324 even through absorbing and judging the mental scenarios of others, without either
325 admission or cognizance of the reproductive significance of the assessment.”
326
327 Alexander’s views of human behavior gave him a unique perspective on the evolution of
328 the human psyche. Well before disagreements unfolded between evolutionary
329 psychologists, evolutionary anthropologists and gene-culture coevolutionists, Alexander
330 had developed deep insights into the extreme flexibility of the human mind and the
331 implications for cultural evolution. Alexander saw human cognitive capacities, including
332 intelligence, consciousness, foresight, empathy, theory of mind, moral reasoning,
333 language, abilities such as scenario-building and story-telling, and cultural legacies such
334 as art and religion, as stemming from continual arms races of social competition
335 (Alexander 2006; Flinn & Alexander 2007). He emphasized the interplay of cooperation
336 and competition between individuals and groups. The cross-generational, cumulative
337 effects of human reproductive striving, as collected and codified in cultural traditions and
338 trends, have perennially created new environments in which human strategizing and
339 social competition has unfolded.
340
341 Alexander appreciated the likely importance of “Darwinian algorithms” (psychological
342 mechanisms designed to accomplish adaptive ends), in biasing human perception and
343 behavior in (historically) adaptive directions, as appreciated by evolutionary
344 psychologists. But his keen awareness of the value of novelty and creativity in the
345 context of social arms races led him to be cautious about proposing limits to the power
346 of the human mind to adapt to novel circumstances. His deep understanding of the
16 347 pervasive influence of natural selection on human conscious and unconscious thought
348 and emotion (and attendant strategies of deceit and self-deception) led him to be
349 skeptical concerning cultural evolution as a process independent of organic evolution,
350 untethered from the adaptive strategies of human beings (Alexander 1979). Instead,
351 Alexander viewed cultural evolution as intimately and inextricably linked to individual
352 reproductive striving.
353
354
355 Morality
356 The faculty and institution of morality was a chief and overriding interest during all of
357 these pursuits in Alexander's quest to understand humanity. Morality was the human
358 trait he contemplated most extensively, and after Darwinism and Human Affairs (1979)
359 was the sole human affair he chose to investigate further in a book of its own, The
360 Biology of Moral Systems (1987). In considering humans as a product of natural
361 selection, Alexander found in moral exhortations to selfless beneficence a fascinating
362 professional challenge along the same lines as that posed to Darwin by the presence of
363 sterile castes in social insects. Alexander started from a hypothesis that would be
364 uncontroversial to evolutionary biologists with respect to the traits of any other
365 organism, but got him maligned and misunderstood by many when he suggested it of
366 morality: that it is "reproductively selfish". Even human moral attitudes and behaviors
367 that seem so other-regarding, so selfless, tended to return benefits to the actor. The
368 theory would also have to address some very peculiar features of humans-- things
369 Alexander called the moral paradoxes, the convolutedness of morality. For instance,
17 370 why do we not practice what we preach? We claim the equal value of all people
371 regardless of their relation to us, we promulgate the Golden Rule, we teach our kids to
372 be good regardless of the possibility of being caught... and yet nobody truly acts as
373 though we believe these things. We still privilege our own groups, we place ourselves
374 and our people far above others in our practical estimations, and we are inordinately
375 concerned for reputation and how our actions appear to observers. Thus a workable
376 evolutionary theory of morality has to explain not only the ideals we tend to espouse but
377 also our typical failure to abide by them.
378 A brief synopsis of Alexander’s theory of morality (drawing on the insights discussed
379 above) is as follows: early humans achieved ecological dominance, to such an extent
380 that the most significant selective agents impinging on their survival and reproduction
381 were other humans. As humans lived in social groups, competition would manifest itself
382 not only within a group but between groups. This would further intensify our
383 dependence on groups, and create a ratcheting effect where traits contributory to group
384 cohesion would be increasingly favored. In this situation, individuals could benefit
385 themselves and their kin reproductively in two different ways: directly by competing with
386 others within their groups, and indirectly by helping others in their groups, which in turn
387 would strengthen the group in competition with other groups. Thus we evolved to
388 cooperate in order to compete, both within and between social groups. The result was
389 the primacy of individual social reputation, and a heavily scenario-building, socially
390 intuitive, and strategizing human personality. Each individual would balance the direct
391 and indirect means of garnering benefits. Each individual would also benefit more by the
392 cooperation of others than by cooperating oneself, resulting in a few powerful items
18 393 entering into the human strategic toolbox: tendencies to over-encourage beneficence in
394 others, to overestimate one's own level of commitment, to look for violations in others
395 and conceal them in oneself, and in general to dispense kindness strategically
396 depending on the expected returns to oneself and one's kin. Moreover, these objectives
397 could often be undertaken with greater efficiency if the motivating factors were
398 concealed from the consciousness not only of others but of the agents themselves. All
399 of these strategies and others presuppose conflicts of interest among genetically
400 different individuals. Moral systems are essentially the means by which these conflicts
401 are identified and resolved.
402 Reception to these ideas was (and has continued to be) uneven, for several reasons,
403 some of which remain relevant even for those who understand evolution and selection.
404 First, the ideas are intellectually demanding, and require substantial investment in order
405 for one to understand them and avoid hyperbole, simplification, or an imbalance of
406 components. Often commentators have taken the easier route of lumping Alexander’s
407 work with facile treatments of these problems, that pay mere lip service to kin selection,
408 reciprocal altruism, and other subjects whose complicated nature is discussed and
409 documented by Alexander. As Alexander was developing and publishing his ideas,
410 essentially no rival well-developed evolutionary theory of morality existed that was
411 placed fully into the context of human life history, and that took seriously the real moral
412 experience of people, warts and all. Many people, including those who depict humans
413 as as fundamentally good, prefer to overlook those warts rather than attempt to explain
414 them. Many are discomforted by the picture of a strategizing human who has not
415 thoroughly abandoned the organismal tendency towards selfishness. Many are
19 416 additionally disgusted at the idea of self-deception. To this, Alexander might first offer
417 the olive branch that selfishness and self-deception are used in an evolutionary sense,
418 and on that meaning neither deception nor selfishness are expected always to be
419 conscious. Strangely enough, "self-deception" actually rescues the possibility of
420 authentic moral action by separating evolved objectives from consciousness. But
421 Alexander would never let that molasses go down without the medicine, for the
422 objection to evolutionary selfishness is usually symptomatic of a general human trait
423 that frustrates efforts like Alexander’s to understand ourselves: our natural resistance to
424 honest self-study. We tend to avoid being confronted with our typical attitudes and
425 actions and their reproductive functions. Alexander insisted that we must investigate
426 and accept whatever we find to underlie our thoughts and deeds, both the noble and the
427 disgusting, as this knowledge is our only hope if we want to improve ourselves and the
428 world.
429 Both the depth and iconoclasm in Alexander's ideas on morality can be illustrated in one
430 of his favorite stories. Theodore H. Hubbell, a friend and colleague at the UMMZ, was
431 troubled by the contention that human behavior is by and large self-interested. Once he
432 thought he had a good counter-example, and approached Alexander: “Dick, this
433 morning on my way to work I saw a caterpillar moving across the sidewalk. I knew it had
434 fallen off its host plant, so I picked it up and replaced it on the proper plant species. Was
435 that not an act of pure altruism?” To this Alexander replied, “It may have been, until you
436 told me about it.”
437 Despite Alexander's evolutionary analysis of morality, and despite his consistent refusal
438 to resort to what he called "the transcendent", Alexander never exhibited any signs of
20 439 retreating into the moral cynicism or nihilism that is often associated with "evolutionary
440 ethics" (a term he avoided). In fact, he struggled with increasing fervor to solve what he
441 called the world's most terrible puzzle: our "unholy capacity and willingness to turn
442 close-knit groups into fighting machines" (Alexander 2014, p. 58), a tendency he well
443 knew was a product of the mechanism that had fashioned our moral psychology in the
444 first place. Whereas in the 1980s and 1990s he would end his papers on morality with
445 the general idea that understanding ourselves and our biases is half the battle, by the
446 2010s his primary goal was to nail specific solutions to the "reciprocating echoes of
447 intra-group amity and inter-group enmity" (Alexander 2014, p. 56). When he sat down to
448 write his final professional papers in his 80s, he focused not on evolutionary explanation
449 as an end in itself, but rather insofar as it might contribute to "global harmony" (2013)
450 and "the future of human society" (2014). He considered the central condition for the
451 survival of humanity to be the dissolution of the evolved concept of "the other" as it
452 manifests as a motivator of hatred towards the outgroup. Hence, Alexander (2013, p.
453 417) wrote: “The greatest difficulty in seeking global harmony may derive from human
454 groups targeting one another. Humans alone – among all the world’s species – plot,
455 plan, and organize massive conflicts to defeat or displace similarly organized and
456 cooperative members of their own species. Can we learn to use the current
457 consciousness of our human background to adjust team efforts of all kinds so that
458 honesty, fairness, and negotiation can increase and lead us toward global harmony?”
459 For Alexander the keys to this project would be the universalization of morality, the
460 reduction of aggression and hyper-competitiveness, temperance with respect to views
461 of morality and God, the conversion of despotic governments that treat their citizens as
21 462 property, and the radical extrapolation of kinship and expansion of social reciprocity. He
463 issued various concrete proposals in this vein, for instance that team sports could soften
464 and redirect outgroup animosity. He also wondered whether God might be translated
465 into a naturalistic concept based on a universal social group or extended kinship in the
466 context of our system of morality. Nevertheless, he repeatedly expressed towards the
467 end of his life (e.g. Alexander 2011) that the biggest regret of his professional career
468 was that he had not made satisfactory progress towards fostering our peaceful
469 coexistence. Thus, Alexander left us with a consistent plea, a take-home message from
470 over half a century of thought: accept and fully interpret our evolutionary heritage; for
471 our own sake, strive sympathetically and constructively to understand humanity.
472
473 Conclusion
474 Richard Alexander did not develop his major ideas in a linear fashion. Rather, they were
475 all developed interactively in the ferment of his mind over the course of his life. Over
476 those many years, he generated an expansive vision of human behavioral evolution,
477 composed of a large set of interacting hypotheses, consistent within themselves and
478 among each other, and able to explain many aspects of the human condition. His
479 excitement in discovery and his tireless critical thinking, sustained over a period of so
480 many years, was shared openly with his students and we all benefitted from it. One
481 wrote, speaking for all, that we were lucky to have a seat at the table.
482 Alexander was a great teacher; his students have dispersed across academia and
483 beyond. Over his long and distinguished career he mentored many graduate and
484 undergraduate students, including Kenneth C. Shaw of Iowa State University, Mary
22 485 Jane West Eberhard of the Smithsonian Tropical Research Institute, Daniel Otte of the
486 Academy of Natural Sciences in Philadelphia, Mitchell Weiss of Rutgers University,
487 John Waage of Brown University, Ann Pace of the Bell Museum of Natural History at
488 the University of Minnesota, Harry Power of Rutgers University, Paul W. Sherman of
489 Cornell University, John L. Hoogland of the Appalachian Environmental Laboratory at
490 the University of Maryland, Richard D. Howard of Purdue University, Marianne N.
491 Feaver of North Carolina State University, Gerald Borgia of the University of Maryland,
492 Katherine M. Noonan of Albany, California, David Foltz of Louisiana State University,
493 Cynthia Kagarise Sherman of Ithaca, New York, Joan Strassmann of Washington
494 University, Nancy Moran of the University of Texas, Marlene Zuk of the University of
495 Minnesota, David Queller of Washington University, Alex Mintzer of Texas A&M
496 University, Bernard J. Crespi of Simon Fraser University, Richard Connor of the
497 University of Massachusetts at Dartmouth, Beverly Strassmann of the University of
498 Michigan, Steven Frank of the University of California at Irvine, Stanton Braude of
499 Washington University, Eileen Lacey of the University of California at Berkeley, Kyle
500 Summers of East Carolina University, Andrew F. Richards of the City College of New
501 York, John Pepper of the National Institutes of Health, Deborah Chizek of Boulder
502 Colorado, David Marshall and John Cooley of the University of Connecticut, Anna Bess
503 Sorin of the University of Memphis, Bret Weinstein and Heather Heying of the Aspen
504 Center for Human Development, Daniel Kruger of the University of Michigan, and David
505 Lahti of the City University of New York. Alexander also mentored many postdoctoral
506 scholars, including James Lloyd of the University of Florida, Jasper J. Loftus-Hills of the
507 University of Michigan, Daniel Otte of the Philadelphia Academy of Natural Sciences,
23 508 Paul Turke of Turke and Tomashow Pediatrics, Laura Betzig of the University of
509 Michigan, Mark V. Flinn of Baylor University, Andrew F. Richards of the City University
510 of New York, Rachel Smolker of the University of Vermont, and David C. Marshall of the
511 University of Connecticut.
512
513 We end this obituary with an excerpt from a poem (“Heroes are works of art”) by
514 Alexander (2011, p. 52). He did not write this with reference to himself, but we think it is
515 appropriate.
516
517 Heroes are for all times, all ages
518 they help us to know what we
519 hadn’t even known we wanted to know,
520 to facilitate all that we discover
521 can be discovered, all that
522 can be absorbed into ourselves.
523
524 Submitted by Kyle Summers1, David Lahti2, Stanton Braude3, Beverly Strassmann4,
525 Joan Strassmann3.
24 526 1) Department of Biology, East Carolina University, Greenville, NC 27858
527 2) Department of Biology, Queen’s College, City University of New York, Flushing,
528 NY, 11367
529 3) Department of Biology, Washington University in St. Louis, St. Louis, MO, 63130
530 4) Department of Anthropology, University of Michigan, Ann Arbor, MI, 48104
531
532 The authors gratefully acknowledge valuable advice, discussion and commentary from
533 Nancy Alexander, Laura Betzig, Gerald Borgia, Napoleon Chagnon, Richard Connor,
534 John Cooley, Martin Daly, Mary Jane West Eberhard, Mark Flinn, Steven Frank, John
535 Hoogland, William Irons, Daniel Kruger, Eileen Lacey, Bobbi Low, David Marshall,
536 David Queller, Andrew Richards, Anna Bess Sorin and Paul Turke.
537
538 Literature Cited
539
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542
543 Alexander, R.D. 1971. The search for an evolutionary philosophy of man. Proc. Royal
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545
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548
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550 organization of life. Ann. Rev. Ecol. Syst. 9: 449-474.
551
552 Alexander, R.D. and Noonan, K.M. 1979. Concealment of ovulation, parental care and
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554 Anthropological Perspective, N. A. Chagnon and W. G. Irons (eds), North Scituate,
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561 NY.
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564 C. Stringer and P. Mellars (eds), Univ. of Edinburgh Press, pp. 455-513.
565
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