Showing posts with label Hamilton's Rule. Show all posts
Showing posts with label Hamilton's Rule. Show all posts

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










Wednesday, March 13, 2019

When to exhibit Social Behavior [Cooperation, Altruism]? Clara B. Jones [Graph, Legend]



Fig. 1. When should an Actor or a Recipient exhibit Social* Behavior [Cooperation, Altruism]? Clara B. Jones, 3/13/2019

1. This [idealized] graph**** depicts & predicts when an [idealized] organism--an Actor or a Recipient--should give/exhibit Cooperation or Altruism, defined, formally, as those interactions during which the Recipient benefits [in relative reproductive units].
2. X-axis: Time [T], Energy [E] investment in Social Behavior [reflects organism's T/E budget for a given interaction].
3. Y-axis: Costs [C, in reproductive units to Actor or Recipient] or Benefits [B, in reproductive units to Actor or Recipient] to Inclusive Fitness** [IF= selfish reproductive units (usually, & clasically,  offspring, though, offspring might be delayed, such as when transaction in wealth or other goods--see "Altruism" below) + reproductive units of kin]. r, coefficient of relatedness, is incorporated in measure/estimate of IF [how to measure?].
4. C rise or reach asymptote over T. Costs, for Actor or Recipient, or both, may prevent Cooperation or Altruism between kin when, for example, Cooperation or Altruism increases Competition [for limiting resources such as food, mates, burrows], increasing [for ego or ego's offspring] likelihoods of  dispersal, "competitive exclusion" [demand for ~identical limiting resources], death, etc. Cooperation and Altruism are expected, by definition, to be responses for the avoidance or neutralization of Competition.
5. B rise, then decrease over T.
6. "x" represents the optimal point at which an Actor or a Recipient should exhibit Cooperation [both Actor and Recipient gain reproductive units] or Altruism. [Actor loses, Recipient gains reproductive units]. "x" is the extension to the X-axis of the vertical, black line connecting the points with the greatest separation between the C and B curves--where B > C. "x" is the idealized point maximizing inclusive fitness benefits.
7. The shaded area represents the zone around "x" for which Actor's benefits from Social Behavior are highest. Ideally, the broken vertical lines extend vertically & downward from those points on either side of "x" on the B curve at which the B curve first begins to descend. The shaded area represents the zone of maximal benefit to Inclusive Fitness of A or B, in this case, maximum benefit from an act of Cooperation or Altruism ["IF maximizing"].
8. Actor & Recipient curves and values will usually be asymmetric, by definition [possibly, even, for clones (e.g., identical twins)]. In other words, zones of maximal benefit from social behavior should differ for Actor and Recipient. Overlaying Actor's & Recipient's curves will determine whether there is overlap between Actor & Recipient zones of greatest benefit to ego from Cooperation or Altruism. If there is overlap, the zone of mutual benefit is defined, so that Actor & Recipient should exhibit Cooperation [+, +] within these parameters [of T, E]. If there is no overlap, then conflict arises [between Actor and Recipient &, possibly, kin of one or both].
9. Similarly, an Actor should exhibit Altruism [-, +] to a Recipient when the long-term B of such action[s] outweigh the short-term C [determined by hard-wired Hamiltonian algorithm?].
10. If B from Social Behavior are minimal or non-existent [e.g., low T/E "budget;" little or no overlap between Actor-Recipient zones of greatest benefit], then interactants should adopt an alternate behavior [Alternative Reproductive Behavior--tactic or strategy], such as fight***, avoidance, flight, force, coercion, persuasion, manipulation, exploitation, coexistence--or some combination of these.
11. Group-living organisms experience relatively ongoing "decisions" about how to behave relative to conspecifics; however, it is important to keep in mind that "fitness budgets" change over T and that behavior is condition- [context-] dependent.
12. Ceteris paribus, and on average, curves, B & C, as well as, shapes of zones of maximum benefits for Males [time-minimizers] and Females [energy-maximizers] should differ. Curves, also, are expected to be Age-, Class-, & Role-dependent.

*After Hamilton (1964), "Social Behavior" is defined as Cooperation [both Actor and Recipient gain reproductive units] or Altruism [Actor forfeits reproductive units; Recipient gains reproductive units]
**The term "inclusive fitness" can be confusing since, technically, "fitness" is the frequency of an allele in a population averaged across the reproductive success of all individuals in the population bearing the allele, leading one to inquire how important it is to measure individual variation of traits [since individual variation will be subsumed in the measure of an allele's "fitness"].
***As pointed out by Geoff Parker [1974], aggression should not usually be a first resort because the C of damaging behavior generally outweigh the B.
****The same graphical/conceptual framework could be used for Coexistence if axes modified so that X-axis= savings in T, E to Mean fitness of allele 1 [or 2]; Y-axis= B, C from T, E savings to fitness [Mean frequency] of allele 1 [or 2] in a population. In the case of Coexistence, a Community Ecology process, the population would be escaping Competitive Exclusion. [See, also, my 2014 Springer Brief, Ch 2, for discussion of Coexistence in relation to Hamilton's Rule.]

Acknowledgments: Thanks to my son, Luke [M. Luke Jones] for stimulating discussion and for drawing the figure. Further, via e-mail, I asked Andrew Bourke [Univ. of E. Anglia] to comment on this blogpost. On 4/11/2019, I received the following reply [n.b.: my original blogpost above, including, the graph, have NOT been revised subsequent to Bourke's comments]:

1.     Perhaps the X axis needs clarifying a little – after all, time and energy seem to have most relevance in this context insofar as they are proxies for fitness, so then there would be some relationship between the metrics on the X axis and on the Y axis, and how these interact might then require consideration.
2.     It would be good to have concrete, empirical examples of when the scheme in the figure needs invoking, i.e. showing how considering the time/energy investment dimension in the manner advocated in the figure enlarges our understanding of the occurrence of social behaviours.  Personally I find it hard to think of things in the abstract all the time, so even some hypothetical biological examples/applications might be useful.
3.     I think your footnote 8 makes a good point, i.e. that actor and recipient interests need not overlap even for altruistic and cooperative interactions.  This point was explored theoretically in a fairly old paper by Yamamura and Higashi (1992)*****.  Their work does not seem to have been followed up empirically to any great extent, perhaps because the usual assumption that one of the parties has most control over the interaction (in which case conflict is minimised – e.g. an insect larva may often have little choice but to accept the level of care it’s given) holds.



References
Hamilton WD (1964) The genetical evolution of social behaviour. J Theor Biol 7: 1-52.
Parker, G (1974) Assessment strategy and the evolution of fighting behavior. J Theor Biol 47: 223-243.
*****Yamamura N, Higashi M (1992) An evolutionary theory of conflict-resolution between relatives: altruism, manipulation, compromise. Evolution 46: 1236-1239.

http://www.kurims.kyoto-u.ac.jp/~kyodo/kokyuroku/contents/pdf/0827-14.pdf



Sunday, October 21, 2012

Clara B. Jones: Brief CV


CLARA B. JONES, Independent Researcher [DOB: 8/12/1943-]: Brief CV [1970s-present]: h-index, 22; i-10 index, 43 as of April 2024 ...

Cell: 828-279-4429
Twitter [Social Biology, Behavioral Ecology]: http://twitter.com/cbjones1943 

Training, Research, Employment [selected]:

Cornell University Ph.D. Biopsychology 1978 [Dissertation Advisors: William C. Dilger: birds, Ethology; Ruth E. Buskirk: spiders, primates, Behavioral Ecology]

Harvard University Postdoctoral Fellow in Population Genetics 1981-1982 [Richard C. Lewontin]

Independent Researcher [including field research in Latin America 1973-2007 (plants; especially, animals): Costa Rica, Panama, Belize, Mexico, Colombia (Colombian Amazon, Rio Negro, squirrel monkey, Saimiri sciurius; Isla San Andres, Colombia--fish, blenny, Entomacrodus nigricans)]

Community Conservation, Inc., USA, Associate 1997-2007 [Rob Horwich]

Organization for Tropical Studies [OTS], Course # 1973-2, Costa Rica; San Andres Island, Colombia

Max Planck Institute for Behavioral Physiology; Seewiesen, Bavaria 1981 [Irenaus Eibl-Eibesfeldt]

American Museum of Natural History, NYC, Visiting Scientist, Mammals 1985-1986: Pleistocene forest refuges, Africa, Primates [Sydney Anderson]

Rutgers University, NJ, Institute of Animal Behavior, Visiting Faculty 1991-1996

Universidad Veracruzana, Veracruz, Mexico, Visiting Scientist 1996 [Ernesto Rodriguez-Luna]

Jackson (MS) State University, Department of Psychology, Visiting Scientist 2002 [Sheree Watson]


National Evolutionary Synthesis Center [NESCent], Visiting Scholar 2005, 2006

Additional Coursework: Environmental Sciences [M.A. Program, Montclair State (NJ) University, not completed: Harbans Singh]; GIS [M.A. Program, U-MD College Park, not completed: Derek Thompson]

Doctoral Committee: Biological Psychology: [Ethology: William C. Dilger, birds (Department of Neurobiology & Behavior); Behavioral Ecology: Ruth E. Buskirk, spiders, primates (Department of Neurobiology & Behavior); Social Psychology: Stephen C. Jones, humans (Department of Psychology)]

Current Research Interests: Animal Behavior; Behavioral Ecology [cf. John Hurrell Crook, 1964]; Social Biology [especially, Social Evolution: Major Transitions Approach, especially, Mammals, including, Humans]; Thermal Biology [Metabolic Theory]; Hystricognaths; Bathyergidae [African mole-rats]; Evolution of Interdependence; Evolution of Cooperation; Evolution of Division-of-Labor; General Principles, especially, Hamilton's Rule [rb - c>0 -----> rb>c]
Taxa studied: Tropical Plants [Botany Mentor, Harlan Banks, Cornell University]: Fieldwork  Pithecellobium saman, Andira inermis; bracken fern [Dennstaedtiaceae spp.]; Tabebuia neocrysantha [Bignoniaceae]; xaté [Chamaedorea spp.]; Animals, Fieldwork Published: Fish [Entomacrodus nigricans]; Mammals [including, humans]; particularly, howler monkeys, Alouatta spp. [Alouatta palliata, 3 sub-spp.; also, A. pigra, A. caraya]; Fieldwork  Unpublished: spiders, vultures, dung beetles; Laboratory Unpublished: albino [Norway] rats; Madagascar cockroaches; planaria; Fieldwork  Unpublished: vultures; scorpions; dung beetles
Publications: > 100, including, scientific articles and book chapters; 9 books [including, five conventionally-published books (2 of these edited volumes--one of these with co-author & the other singly-authored); two self-published books; one self-published monograph; one self-published blogpost--self-published texts available at lulu dot com]; 2 special Issues [1 issue comprised of 2 issues]; book reviews; technical reports; newsletter; and newspaper articles


Primary influences: Sydney Anderson, Harlan Banks, Irwin Bernstein, Andrew Bourke, Jack Bradbury, Bernie Crespi, William C. Dilger, Irenaus Eibl-Eibesfeldt, John F. Eisenberg, Stephen T.  Emlen, "Griff" Ewer, Steven A. Frank, Masao Kawai, Harry Levin, Richard C. Lewontin, Jasper Loftus-Hills, Martin Moynihan, Gene E. Robinson, M.E.P. Seligman, Norman J. Scott, Jr., Robert L. Trivers, Sandy Vehrencamp, Frederick O. Waage, Stuart A. West, Mary Jane West-Eberhard



Books


1.      Jones CB [ed] [2003] Sexual selection and reproductive competition in primates: new perspectives and directions. American Society of Primatologists, Norman, OK


2.      Jones CB [2005] Behavioral flexibility in primates: causes and consequences. Springer, New York


3.      Hager R Jones CB [eds] [2009] Reproductive skew in vertebrates: proximate and ultimate causes. Cambridge University Press, New York


4.      Jones CB [2012] Robustness, plasticity, and evolvability in mammals: a thermal niche approach. Springer, New York
5. Jones CB [2014] The evolution of mammalian sociality in an ecological perspective. Springer Brief, Springer, New York
6. Jones CB [2020] Female mantled howler monkey (Alouatta palliata
palliata: Primates, Atelidae) life-history strategies—a “major transitions” 
approach to mammalian social evolution. Lulu dot com. [self-published]
7. Jones CB [2021] A mechanistic approach to studying mammalian populations. lulu dot com. [self-published] ... book highlights "social parasitism" & includes a simple mathematical model, pp 44-46:
https://www.lulu.com/shop/clara-b-jones/a-mechanistic-approach-to-studying-mammalian-populations/paperback/product-8dnw7q.html?q=clara+b.+jones&page=1&pageSize=4

Special Issues
1.      Jones CB [ed] [2001] Sampling Neotropical primates: implications for conservation and socioecology. Primate Report 61: 3-71


2.      Jones CB [ed] [2003] Primate dispersal: proximate and ultimate causes and consequences [Part 1]. Primate Report 67: 3-98


3.      Jones CB [ed] [2004]. Primate dispersal: proximate and ultimate causes and consequences [Part 2]. Primate Report 68: 3-95


Other [Selected] Publications

Horwich R et al. [2012] Community conservation. In: Moutinho P (ed), Deforestation around the world [Ch 14], pp 283-318. InTechOpen.com

http://www.communityconservation.org/publications/InTech-Preserving_biodiversity_and_ecosystems_catalyzing_conservation_contagion.pdf

Jones CB [2013] Seasonal tropical forests. In Horwath RW [ed], pp 163-168. Biomes and ecosystems. Ipswich, MA, Salem Press

Jones CB [2013] Sub-tropical forest biome. In Horwath RW [ed], pp 142-148. Biomes and ecosystems. Ipswich, MA, Salem Press

Book Review: Trivers RL [2015], Wild Life, Biosocial Research, NJ; International Society of Behavioral Ecology Newsletter 28-1, Spring/Summer 2016

Book Review: Ebensperger LA & Hayes LD [2016], Sociobiology of Caviomorph Rodents, Wiley-Blackwell; Koenig WD, Dickinson JL [2016], Cooperative Breeding in Vertebrates, CUP; ISBE Newsletter 28-2, Fall/Winter 2016

Book Review: Clutton-Brock T [2017], Mammal Societies, Wiley-Blackwell; ISBE Newsletter 29-1, Spring/Summer 2017

      Book Review: Wilson EO [2018] Genesis

        https://vertebratesocialbehavior.blogspot.com/2019/04/review-of-eo-wilsons-new-book-genesis.html

    NOTE: Jones CB (2022) A Note Concerning Constraints on Speciation and the Monospecific Status of Genus: Homo, Emphasizing Environmental Potential and the Role of Gene Flow Among Nomadic Hunter-Gatherers, Facilitated by Behavioral Flexibility and Phenotypic Diversity, Including, Cultural Innovations; lulu dot com


Videos

1. Terminology in Social Biology (2016) YouTube ~6 min

https://www.youtube.com/watch?v=o0cMyWzB33o&feature=youtu.be

2, Are Humans Co[-]operative Breeders (2016) You Tube ~6 min

https://www.youtube.com/watch?v=Cvz3D3sKlJ8

3. Clara B. Jones reading part of book on Naked Mole-Rats (2018); YouTube

https://www.youtube.com/watch?v=X5xlRHmKB84

    4. Mammal Social Evolution


Current project

Social Evolution: Major Transitions Approach, especially, Mammals [see 1st blogpost of this blog; available in hard copy at lulu dot com] 


Selected Scientific Contributions:

1.      First systematic utilization of “Focal” data-collection technique employing randomized [1 randomly-selected focal subject/d] baseline using physical lab data sheets [10-columns, min x min recording]; all publications for aged and marked Costa Rican Mantled howler Monkeys**, Alouatta palliata palliata, and Riverbanks Zoo Black [now, Black and Gold] howler monkeys, A. caraya; Costa Rican Mantled Howler Monkeys studied in Tropical Dry Forest habitats, Canas, CR in 2 habitats, drier, Deciduous habitat (Group 12) & wetter, Riparian habitat (Group 5) [Behavioral Ecology]; Dissertation research carried out studying 1 species in two habitats, the first or among the first such research designs in Primatology [Behavioral Ecology]

2.       First systematic utilization of Radio-telemetry*** [AVM receiving equipment w hand-held antenna; lab-made transmitter attached to one adult female] in field primate studies; all publications for Costa Rican mantled howler monkeys, A.  p. palliata, in Deciduous Habitat


http://pin.primate.wisc.edu/news/cons/COMBELEN.html



3.  Devised fist systematic qualitative system [visual inspection] to determine estrus stages in howler monkeys [in primates?: 3 stages based on differential tissue color & presence/absence of vaginal secretions [A. p. palliata]





http://link.springer.com/article/10.1007/BF02382013



4.   3rd field translocation experiment (1976) utilizing primates as subjects [published (A. p. palliata]; see, also,  Kawai M(asao) [1960] Primates 2: 181-255 and Sugiyama Y(ukimaru) [1966] Primates 7: 41-72.]*****


http://link.springer.com/article/10.1007/BF02381443


5.       Systematized and implemented “focal-tree” data-collection method (published: Brenesia; blogpost @ vertebratesocialbehavior.blogspot.com)****

6.       Principal descriptions of "age-reversed” ["age-inversed"] dominance system in 3 Alouatta [howler monkey] species: A. palliata [3 subspecies: 1978 (dissertation), 1980], A. caraya, A. pigra*****; published: https://link.springer.com/article/10.1007/BF02390468 

Bioaccumulation: my new hypothesis for the evolution of the "age-reversed" ["age-inversed"] dominance system [2nd blogpost in this blog]; i think this H could be easily tested in the field ...

7.       Conducted first systematic [field] experimental manipulation of primate herbivore-plant interactions, A. p. palliata [blogpost @ vertebratesocialbehavior.blogspot.com]


8.   Conducted opportunistic field experiments using Costa Rican mantled howler monkeys [blogpost @ vertebratesocialbehavior.blogspot.com]

9.    First [only?] systematic use in primates of “Vehrencamp’s RRS Method” to calculate “relative reproductive success” [RRS] devised by Sandra L. Vehrencamp [University of CA, San Diego, communication, mid-1970s, in Costa Rica]; published in Neotropical Primates

10.   Demonstrated “displacement coalitions” by male and female mantled howler monkeys; published Jones CB 1980, Primates ... these apparently coordinated displacements appeared to be opportunistic rather than collaborations or alliances ...

11. First quantitative modeling of climate time-series "mapped" onto primate population life table to demonstrate "fine-grained" conditions: Jones CB [1997] Life-history patterns of howler monkeys in a time-varying environment. Boletin Primatologico Latinoamericano 6: 1-8


12. Demonstrated correlation between folivority and capacities for colonization [Belizean black howler monkeys, Alouatta pigra] and frugivory and minimal capacities for colonization [Central Americal (Belize) spider monkeys, Ateles geoffroyi] due to even spatiotemporal dispersions of leaves, clumped spatiotemporal dispersions of most fruit species [Jones & Jost 2007, Laboratory Primate Newsletter]******.


13. Preliminary demonstration of "temporal division-of-labor" [TDL] in a primate: Jones CB [1996] Temporal division of labor in a primate: age-dependent foraging behavior. Neotropical Primates 4: 50-53

http://www.primate-sg.org/storage/PDF/NP4.2.pdf

Monograph, Female mantled howler monkey (Alouatta palliata palliata, Primates, Atelidae) life-history strategies--a major transitions approach, 122 pp, lulu.com ... also, PDF available linked to Profile of my Twitter feed, @cbjones1943 [see Abstract below] ...


14. Began to compare social mammals [primates] and social insects in Jones CB 1980 Primates; e.g*


http://www.redalyc.org/pdf/457/45712103.pdf

15. Publications on "behavioral flexibility" & "phenotypic plasticity"

16. Using a verbal model, synthesized Hamilton's Rule, Competition Theory, and Coexistence Theory [Jones CB (2014) Springer, Chapter 2]

17. Probably the 1st to apply a Major Transitions Approach to mammalian social evolution [see 1st blogpost of this blog].
 
18. "Temporal division of labor" ["age polyethism"] in a mammal: Female mantled howler monkey [Alouatta palliata palliata: Primates, Atelidae] life-history strategies--a "Major Transitions" approach to mammalian social evolution [2020] ... available in hard copy at lulu dot com; available in PDF format linked to Profile of my Twitter feed, @cbjones1943

ABSTRACT
Arboreal howler monkeys [Alouatta spp.] are wholly herbivorous [“primary
consumers”]. Following earlier work (1978; 1980) on Costa Rican mantled howler
monkeys, Alouatta palliata palliata, located in Tropical Dry Forest, the dominance system is described whereby young adult females [~5-7 y.o.] are dominant to older females, middle-aged females [~7-10 y.o.] and middle-aged to old females [~10-15 y.o.] are dominant to old females [~15+ y.o.]. Importantly, the dominance system is characterized by "temporal division-of-labor" ["age polyethism"] 
whereby adult females, specialized for "social foraging"—a type of within-group “helping” behavior, are graded by age, with the oldest female foragers ["helpers"] engaged in most foraging bouts, younger adult females accounting for less “social foraging” [“helping”]. Adult female life-history parameters were calculated from a population census and shown to correspond to patterns of temporal environmental cycles, in particular, the 6-month [seasonal] pattern of rainfall associated with limiting
food availability. To my knowledge, this is the first demonstration of "temporal division-of-labor" in primates, and the methods are novel. Eusocial mole-rats have been shown to exhibit both “temporal” and “reproductive” division-of-labor [Nigel Bennett, personal communication, 2021]. The present results indicate feedback loops among cyclical rainfall patterns; ephemeral food availability; and, adult females operating in the context of “contest competition” for limiting nutrients from which female life-history traits and the rare “age-reversed” dominance system have emerged. The present study, also, has implications for the evolutionary causes and consequences of cumulative acquisition of information by adult females over time who share a group range.

References

Jones CB (1978) Aspects of reproduction in the mantled howler monkey (Alouatta palliata Gray). Unpublished Ph.D. dissertation, Cornell University, Ithaca, NY.

Jones CB (1980) The functions of status in the mantled howler monkey (Alouatta palliata Gray): intraspecific competition for group membership in a folivorous Neotropical primate. Primates 21: 389-405.

18. Mammal social behavior and group-living ["social organization" from the perspective of Social Parasitism ... A mechanistic approach to studying mammalian populations [2021] ... available in hard copy at lulu dot com ...


Skill Sets: Reading & conceptualizing 3-D visualization graphs/maps; Writing for publication [empirical research papers, synthetic theoretical/review articles, book chapters, technical monographs [2 Springer Briefs],  notes, newspaper [Salisbury NC] & newsletter articles, book reviews]; Field Research [Methods, Design, & Analysis, especially Invasive Field Experiments]; Animal Immobilization & Translocation [field]; Animal predator-Plant prey Manipulations [field]; Technical Networking; Solution-oriented Problem-solving [including Brainstorming & Negotiation]; Conflict-resolution; Teaching [Undergraduate, Graduate]; Coaching [Solution-focused life tactics and strategies]; Administration [inc. Program Director, Department Head, Division Head]; Reviews of technical papers, chapters, proposals; Private-, NGO-, regional-, community-, & government-entity collaborations related to conservation biology; Development of academic courses [e.g., Genetic Aspects of Behavior]

Footnotes

*Jones CB [2005] Social parasitism in mammals with particular reference to neotropical primates. Mastozoologia Neotropical 12: 19-35

**Animals [Group 5 and Group 12] aged and marked by Norman J. Scott [USFW, retired] and his assistants [including CB Jones]; 2 raw data, 3-ring binders [Group 5: Riparian Habitat; Group 12: Deciduous Habitat] archived via Dr. Todd Vision @ National Evolutionary Synthesis Center [NESCent], Duke University, Durham, NC

***Telemetry equipment provided and 1 adult female ["TC"] fitted with transmitter collar by Scott

****Concept first suggested by Jack W. Bradbury [Cornell University, communication early 1970s], who, also, highlighted the importance of using multiple field assistants systematically recording observations concurrently, an essential procedure for predator-prey experiment

*****with Robin Brockett & Rob Horwich

****** http://pin.primate.wisc.edu/news/cons/COMBELEN.html


******Using mapped survey data [Robert H. Horwich's raw data] of the two species' distributions in Belize [the only non-human primate taxa in Belize]




Photo by Liz Williams www.makemesomeart.com