Thursday, April 11, 2019

Review of Robert L. Trivers' Memoir, Wild Life (by Clara B. Jones, 2016)

Wild Life: Adventures of an Evolutionary Biologist
Robert L. Trivers
Biosocial Research
New Brunswick, NJ
2015
225 pages
$9.58 (Paperback and Kindle)

Reviewed by Clara B. Jones, Asheville, NC, USA (February, 2016; slightly revised, 4/11/2019)

To dispense with preliminaries, I have interacted with Robert L. “Bob” Trivers intermittently since I was in graduate school but have never been in his “inner circle.” I requested a complimentary copy of Wild Life from Robert, telling him that I intended to review it for a journal that was, at the time, undetermined. Robert was enthusiastic about the idea and sent the memoir. I have reviewed a number of books for ISBE Newsletter so it was logical for me to consider it as the venue for publication, particularly, since I speculated that many readers would be interested. I appreciate Andreas Svensson for giving me this opportunity.

In addition to several other biologists (e.g., Louise Emmons, Dan Janzen, Russ Lande, Nancy Moran, Steve Stearns, Mary Jane West-Eberhard, Don E. Wilson, EO Wilson), I consider Trivers to be a National Treasure. Against numerous odds, he has risen to the top of his field, having published papers that are fundamental contributions to Social Biology, generators of immeasurable bodies of systematic and informal research and publications, as well as, continuing streams of productive thought—among his peers, other scientists, educators, students, journalists, and the general public. Many persons in and outside the scientific community will purchase a copy of Wild Life hoping, even, expecting, to find inside their names or mention of one of their vivid memories with Robert. Most of these men and women will be crushed, and I can think of four or five who will never recover. Robert's new book should have included a detailed index, not only for reference purposes, but, also, to prevent time wasted by some who would otherwise rifle through the book in search of evidence that Trivers considers them worthy of mention.

In the literary world, a memoir belongs to the genre, Creative Nonfiction. Wild Life, however, for the most part, appears to be uncontrived, a convincing example of a brilliant, though, controversial, figure telling it as he truly sees it. As the saying goes, “What you see is what you get.”, though Robert is clear to point out that he is unable to measure the full extent to which deception and self-deception influenced this project. After reading Wild Life, I was left with several messages, one being that its author has written a memoir hoping to determine the arc of his legacy. With the exception of Chapters 14 and 15, he crafts his story, as he has presented all of his ideas, with clinical, even, scapular, precision. There is a sense in which the dominant theme of the book can be summarized by a single observation expressed on page 186 when speaking of W.D. Hamilton: “I thought of Bill as perhaps the greatest evolutionary theorist since Darwin. Certainly, where social theory based on natural selection is concerned, he was our deepest and most original thinker.” Many would place E.O. Wilson in second place. However, in Wild Life, as well as, elsewhere, Robert seems of the opinion that Wilson is greatly overrated (an opinion with which I disagree). Though he does not say so explicitly, Trivers likely considers himself to hold second rank after Hamilton, a self-assessment that many would challenge if only because of the author's non-normative record of behavior. Nonetheless, as Robert told a reporter at Rutgers University in 2014, “I don’t want to sound immodest, but I am one of the greatest social theorists in evolutionary biology alive, period.”

Wild Life, dedicated to Robert's teacher, William H. Drury, Jr., comprises fifteen chapters, seven of which, in addition to numerous additional remembrances, concern his life and research in Jamaica. He reports, with obvious pride, that all of his five children are “American/Jamaican,” referring to himself as an “out-breeder” created by his peripatetic experiences as the son of a career diplomat. For decades, I was an obsessive reader of the print-version of The New York Times. One day in 1987, scanning the obituary page, I noticed the name, Howard Trivers [sic, no middle initial], and, because, at that time, I had received second-hand reports of Robert's life from one of his close friends, I determined from relevant details that the deceased was the evolutionary biologist's father. Among other prominent roles in the U.S. Department of State, Howard Trivers had been Director of its Office of Research and Analysis. It was impossible not to note that the diplomat's obituary included no personal information—no mention of parents, a wife, or other family members. I had been told that Robert had a very contentious relationship with his father, in part, because of the latter's profession. Robert has only mentioned his father to me once, in passing; thus, I was not totally surprised to find virtually no reference to his family, including his mother, in Wild Life. For this reason, combined with the, sometimes shocking, even, disturbing, particulars of his life, Robert would provide a Freudian scholar with copious material and room for speculation (e.g., Oedipal failure?). In addition to passing mention of his “family of origin” and his children, the Preface expresses the author's opinions about the conventional lives of scientists (“This kind of life never appealed to me.”), and the remaining few paragraphs introduce the reader to themes that follow.

Chapter 1 is revealing to all students of Trivers' work. He documents his early precocity in mathematics, particularly, The Calculus. I consider myself an amateur student of the thinking strategies of famous scientists. Many population ecologists, for example, think like physicists, characterizing genes and other events as mass flowing in space and time. Ecosystem ecologists, on the other hand, are likely to think, not only spatially, but at multiple scales at once. Robert's mind is that of a quantitative modeler expertly identifying and manipulating decision rules. A limitation but, even, more, a strength, of his mostly verbal theories is that they bypass complexity to identify fundamental principles of natural selection for mechanisms and functions of general import. Trivers reveals that Drury taught him to begin with interesting questions about human behavior, and Robert has exhibited a remarkable ability to choose topics basic to non-human, as well as, human, Social Biology, always cognizant of intra- and inter-individual conflict (-of-interest) and of genetics operating at the level of individuals, constrained by Hamilton's Rule. While I imagine there is intuition and art involved in Trivers' generative processes, I think his successes can be attributed, primarily, to his cogent choice of topics and his ability to write with laser clarity, as close as verbal models can come to mathematical ones. In my humble opinion, Robert is a better verbal modeler than Charles Darwin.

Chapters 2 and 3 describe early experiences conducting fieldwork and the impact of Drury and Ernst Mayr, respectively, on the development of the memoirist's early thinking, publications, and career. Robert convincingly communicates the extent to which he honors these men, and he shares with candor the pivotal role they played in the formation of some of his most important ideas. Trivers has reason to be confident that his acknowledgments of others' inputs will not reduce his reputation and that, though some critics may attempt to diminish his project, particularly, posthumously, Robert's theories are not subject to claims that they are derivative. Chapter 4 is titled, “I Become a Lizard Man in Jamaica,” describing how he became “a green lizard freak” after accompanying Ernest Williams there as a research assistant. Some readers will be offended by both men's evaluation of primatologists and, by implication, Anthropology as a field (but, see Chapter 6). In another chapter he denigrates Psychology, and it is clear throughout the memoir that Robert's opinion about what constitutes a Science is a narrow one.

Wild Life is peppered with interesting facts about lizards and other animals, as well as, snapshots of geography and human nature. Chapters 5, 7, 8, 9, and 10 recount a variety of experiences in Jamaica, some of them life-threatening, as well as, significant friendships, mostly with men. The only women who receive a lingering nod in the memoir are “mother-in-law,” “Miss Nini,” and her daughter, Robert's “wife,” Lorna, mother of four of his children. Even though these chapters, and a few other accounts, document Robert's capacity for deep, sincere, and reciprocated feelings, he sometimes refers to friends using clinical, seemingly detached, language.

Chapter 11 is an interesting one in which the author reminisces about his relationship to Huey Newton and the Black Panther Party, and Chapter 12 continues descriptions of his entanglements with intra-specific conflict, including, incarceration. Robert seems not to have learned a litany from Behavioral Ecology that the costs of aggression (or, spite) usually outweigh its benefits (see the self-analysis in Chapter 15, and Parker 1974). Chapter 13, titled, “Vignettes of Famous Evolutionary Biologists,” suggests that, in addition to Drury, Mayr, and Williams, Richard Dawkins and W.D. Hamilton, effectively, complete Trivers' list of illustrious figures in the field who have influenced him and his ideas, and in this chapter, Robert includes a few sarcastic paragraphs about Stephen J. Gould that I consider gratuitous and unnecessary.

The final two chapters are intimate in nature, Chapter 14 is titled, “Ambivalence About Jamaica,” describing the unsettling and increasing rates of violence there. Some readers will find the final chapter haunting since Robert candidly assesses personal failings and outlines his burial plans. I was surprised that he had nothing to say about politics or the state of the world (e.g., climate change, income inequality, racism, biodiversity loss, terrorism), if only to inform the reader about how his opinions and values might have changed since his Black Panther days (“One man's terrorist is another man's freedom fighter.”, as the CIA used to say). His life is not as dark as it may appear, however, since Robert has recently outlined several proposals for future research, specifically, speciation processes (with Koos Boomsma); evolutionary heterogamety; natural selection of honor killings; evolutionary dynamics of homosexuality; and, human evolutionary genetics.

It would be remiss and unrealistic not to provide some academic critique of Trivers' work, and I consider it appropriate to suggest a couple of limitations*. Since I was introduced to his publications in the 1970s by Behavioral Ecologists at Cornell, it has concerned me that, though "inclusive-fitness maximizing" and sexual selection are fundamental to Robert's writings, his body of work fails to reflect the importance of Evolutionary Ecology, particularly, life-history evolution (e.g., Stearns 1976), as well as, evolution in changing environments about which there is a significant literature pre-dating Robert's first publication (e.g., Levins 1968, Lewontin 1957). Though Evolutionary Ecology is relatively recent as a systematic discipline, Robert is aware of Population Genetics and G x E interactions, an operation receiving limited treatment in his publications (e.g., How do “selfish” genes and social traits behave when conditions vary or along gradients? When and under what conditions is social behavior situation-dependent? ...flexible?). As important as his neglect of evolution in heterogeneous regimes [Population Ecology] & of life-history evolution, Robert, in addition, fails to address Behavioral Ecology, especially, the fundamental & "classic" work of John Hurrell Crook (1964: group structure in response to limiting resource dispersion [distribution and abundance] in Time & Space). All of these concerns relate to the "condition-dependence"/"situation-dependence" of organismic responses & adaptation, or, lack thereof, in the context of "local adaptation."

On the other hand, Robert's “feel” for statistical thinking is that of an expert, and I would expect more treatment of variation and deviations from central tendencies in his canon. Of particular import, is that, regardless of having produced seminal papers in his early career, as well as, an interesting book on sociality (Trivers 1985), Robert has shown little interest in the EVOLUTION of sociality [as, Major Transitions Approach: West et al. 2015], per se, as per transitions from solitary breeding, to breeding in groups [including, group formation & group maintenance], to Presocial state, to Subsocial state, to Cooperation [as per Hamilton '64], & to Complex Sociality (reproductive division-of-labor; specialization as per Eusociality variously defined); see, Wilson 1971's schema]. Furthermore, treatment of social evolution in comparative perspective [within & between genera, families, orders, classes] is lacking in Robert's oeuvre.

On the other hand, one way that Trivers' work achieves elegance, is by simplifying complex phenomena; thus, environmentally-focused, realistic theories might not have been as successful, productive, or fundamental. In a sense, the author's insights are primitive in the deep, honorific sense that the word is employed to describe some treatments in pure mathematics. Nonetheless, all theoretical work of import is subject to vetting by subsequent theory, models, experiments and other empirical tests. Already, a few researchers have modified certain details of Trivers' theory of sex ratio selection, and his ideas about parental investment and sexual selection have been challenged by some feminist biologists. Related to any discussion of Robert's legacy, in Wild Life, the memoirist states how important the appreciation of conflict (genetic and whole organism, intra- and inter-individual) has been to his success. In my opinion, this observation confirms Trivers' understanding that asymmetries produce differential “fitness optima” and that differential (asymmetric) phenotypes are exposed to environments upon which selection may act. I strongly recommend Wild Life to all who are interested in Ecology and Evolutionary Biology, not only, for its explication of “wild” experiences, but, also, for insights into how a stunning mind works.

References

Crook JH (1964) The evolution of social organization and visual communication in the weaver birds (Ploceinae). Behaviour Supplement #10: 1-201.

Levins R (1968) Evolution in changing environments: some theoretical explorations (No. 2). Princeton University Press.

Lewontin RC (1957) The adaptation of populations to varying environments. Cold Spring Harbor Symp Quant Biol 22: 395-408.

Parker GA (1974) Assessment strategy and the evolution of fighting behaviour. J Theor Biol 224: 115-126,

Stearns SC (1976) Life-history tactics: a review of the ideas. Quart Rev Biol 51: 3-47.

Trivers RL (1985) Social evolution. Benjamin-Cummings.

West SA, Fisher RM, Gardner RA, Kiels ET (2015) Major evolutionary transitions in individuality. PNAS 112(33): 10112-10119.

Wilson EO (1971) The insect societies. Belknap (Harvard), Cambridge, MA.


*First, no American social biologist can compete with Wilson's expertise as a student of a single social group, ants, in Wilson's case. Second, IMO, The Insect Societies (1971) is the greatest book ever written in Social Biology, indeed, in Animal Behavior and Ethology as a whole.

Third, unlike E.O. Wilson, Trivers is not a synthesizer, though his most heralded papers have general import. Trivers has not communicated much interest in a search for general patterns--within, between, and across taxa. Also, again, in contrast to E.O. Wilson, Trivers' canon pays scant attention to Genotype<----->Phenotype<----->Environment<-----> causes and effects. One seeks, for the most part, in vain, to locate Ecology--abiotic & biotic Environments [however conceptualized]--in Trivers' writings. Furthermore, to my knowledge, Trivers has not emphasized the topics--group-formation, group maintenance, and the environmental conditions facilitating the emergence of sociality [which may or may not follow group-formation]. Especially pertaining to the latter are the topics, competition and differential access to limiting resources, as well as, limiting resource dispersion [distribution and abundance in time and space].

Trivers may be one of the last remaining extreme genetic thinkers. He typically asks a question, then, considers what consequences would obtain given alternate, pairwise combinations of related individuals [parents, full sibs: 1/2; first cousins: .1/8, etc.]. This approach has yielded several fundamental papers; however, Trivers' work does not satisfactorily address variations in inter-individual interactions nor evolution in heterogeneous regimes nor phenotypic plasticity nor the principle that behavior is condition-dependent. In other words, an actor & a recipient, whatever their "r" [coefficient of relationship], will respond relative to "b" [benefits to recipient] and "c" [costs to actor ("donor"--of reproductive units)], rather than, strictly, "r" [Hamilton's Rule: rb - c >0].

Trivers' literal logic based, apparently, on "r" alone, may reveal one unfortunate consequence of the term, "kin selection" that leads many to assume that it is always in ego's favor to exhibit social behavior towards kin. The latter assumption may be an assumption behind Trivers' [very successful and justifiably heralded] publications. Furthermore, as I [and several others before me (thanks to James Marshall and Andrew Bourke for making me aware of this literature)] have suggested, it may be useful to consider the role of competition influencing behavior between actor [donor] and recipient and to question whether Hamilton's Rule adequately incorporates the consequences for actor and recipient and for the expression or non-expression of cooperation or altruism [i.e., "social behavior"] of interindividual competition [for limiting resources, e.g., food, mates, space].

Thus, sometimes, kin may be ego's "worst enemy," a litany of Behavioral Ecology [it may not be beneficial for ego to assist the reproduction of kin; it may be in the interest of ego to assist the reproduction--depending upon environmental regime] where predation is non-random by genotype [where cooperation or altruism toward a relative would increase ego's chances of becoming prey]. But, complicating the matter, in certain conditions, death, however, defined [e.g., self-induced, other-induced], can benefit kin. Clearly, systematic empirical and theoretical studies, in addition to modeling, are needed.

In the final analysis, however, the impressive success of Trivers' verbal models based on "r" may demonstrate the power of Hamilton's Rule to predict a very broad array of the social acts [cooperation, altruism] observed in Nature, including, Human Nature. However, we should not only ask, "What is "r"?, but, also (or rather?), "'r' relative to what?" According to Hamilton's Rule, the effects of "r" are expected to be constrained by the components that comprise "b" & "c".

Clara B. Jones (foucault03@gmail.com)
Asheville, NC, USA (now, Silver Spring, MD, USA)
February, 2016 In International Society of Behavioral Ecology Newsletter