Thursday, September 19, 2019

My amazon.com comments on Eisenberg JF (1981)...Mammal "radiations" (Clara B. Jones)

My amazon.com comments on Eisenberg JF (1981)...Mammal "radiations"...

The mammalogist, John F. Eisenberg's, oeuvre remains one of the most highly respected and important in his field. Because most of his research investigated Neotropical mammals, rather than mammals of the Old World, his is not a household name. Because of my own specializations, in this brief review, I limit my comments to Eisenberg's treatments of mammal Behavior and Social Organization [group-living, interindividual interactions, cooperation, and altruism]. With E.O. Wilson's 1975 important chapter on patterns of sociality in Class: Mammalia ["The secret to the evolution of sociality in mammals is milk."], Eisenberg's "radiations" is the first attempt to identify PATTERNS of Behavior & Social Biology across all group-living mammals for which there was data by 1981. No subsequent treatment has attempted successfully to summarize the social biology of mammals comparable to Wilson's 1971 treatment of Social Insects [necessary reading for Behavioral Ecologists & Social Biologists] or Holldobler & Wilson's 1990 treatment of the Ants. Though Tim Clutton-Brock recently published a book [2016]  titled Mammal Societies [sic], it stands as a highly selective literature review, rather than a synthesis. The limited and highly selective chapter by Smith et al. in Rubenstein & Abbott's 2017 Comparative Social Evolution omits Primates and posits bats as the Order needing research as a model for the evolution of sociality in mammals [a view, possibly, derived from Wilson, 1975's discussion of bats]. To the contrary, bats are a highly derived group not suited for a model of general patterns. Rodents are the taxon of choice, including large-bodied and small-bodied groups, sexually-segregated ["solitary"] to highly social species [including the social mole-rats], and taxa with generalized as well as specialized phenotypes--living across virtually all ecotones, and with a commensal relation to humans--that will be most helpful in the early phases of identifying common features across mammals, as well, possibly, across vertebrates. Additionally, critical to any scientific treatment, and as Eisenberg points out in "radiations," the marsupials are the only group of mammals that can serve as a "control group" for tests of hypotheses and apparent patterns [see comments on this idea in Wilson '75]. My 2014 book, The evolution of mammalian sociality in an ecological perspective, is synthetic, emphasizes Ecology, but is brief. The Class is sorely in need of a synthesis across taxa for which data are available; unfortunately, behavior & sociality of rodents are not well known [but, see, Wolff & Sherman's 2007 volume, Rodent Societies]. Wilson, 1971, 1975, are critical for the standardization of terminology, as well as, factors critical to the study of Social Biology and patterns of group life [e.g., the evolution of Communication, "polyethism"], and Robert Trivers', Social Evolution, as well as, James Costs's, The other social insects, should be consulted. Critically, as a few mammalogists have pointed out [e.g., Bob Selander], initial attempts to synthesize patterns across group-living mammals should begin with analyses that study large and small mammals separately. Finally, we want to assess the observation that the structure of mammal [vertebrate?] groups results from the tendency of females to select rich patches of food and that of males to select the largest relative aggregation[s] of females. Other patterns are identifiable in existing literature. Since the time of John Hurrell Crook [Behavior Monograph X], we know that patterns of group organization depend upon the distribution and abundance [dispersion]  of limiting resources [e.g,, food, mates, sleeping sites]. Also, related to this overview, females are, cet. par., "energy maximizers;" males, "time-minimizers."

General Mammalian Patterns, esp., Patterns of Group-living Species [compiled by Clara B. Jones]

General Mammalian Patterns [?--in no particular order] with an emphasis on Group-living taxa, especially, Social Biology & Behavioral Ecology--also see Twitter @cbjones1943... Conceptual Framework... 

FIRST PRINCIPLES OF BEHAVIORAL ECOLOGY:: E[nergy]: Acquisition->Consumption->Allocation====> Worker &/or Reproductive &/or Dependent...(Males, T[ime] Minimizers; Females, E[nergy] Maximizers)


Mammals: Anisogamy-->Sexually-Antagonistic Selection-->Sexual Conflict-->Differential T-E investment, including, Life History (males, time-minimizers; females, energy-maximizers)-->Sexual Dimorphism, inc, Body Size...constrained by dispersion of limiting resources [especially, food for females, mates for males]--that may or may not not favor group-living...[apply, for example, to evolution of male Alternative Reproductive Behaviors (ARBs) & female "counterstrategies"]...


For analysis, and where data permit, we will separate comments for large [>100 kg] & small mammals--as suggested in literature, e.g., https://pdfs.semanticscholar.org/3281/ad106c0d6a9fdc831b7725dc5d3460222136.pdf

Terminology & Context: Terminology in the field of Social Biology, here, related to Social Evolution, is not standardized ... for example, WD Hamilton & Stu A. West, define "social" in different ways, the former limiting use of "social" to Cooperation & Altrusim, the two [of four] categories of behavior [Selfish, Cooperation, Altruism, Spite] in which Recipient gains "fitness" benefits ... West, on the other hand, defines "social" as any interindividual interaction where either or both interactants benefit reproductively ... recall that "interdependence" is a necessary precursor to Cooperation, while Cooperation is a necessary precursor to the evolution of Complex Sociality ... here, I am thinking of Cooperation as per Hamilton--an interindividual interaction in which both interactants benefit in "fitness"  ... related to the Major Transitions Approach, Complex Sociality [division-of-labor (cooperation between specialists); task, role, &/or morphological specialization] is, particularly, problematic, it seems to me ... if we view specialization along a continuum, say, from feeding specialization to DoL to reproductive DoL with "totipotent" workers or "helpers" to reproductive DoL with more or less "sterile castes" [social insect workers], do we accept the typical usage whereby Advanced Eusocial taxa are the most "complex" societies [the "pinnacle" of sociality as per Wilson (1971)]? ... though many Social Scientists would claim that interindividual interactions in humans represent the "pinnacle" of Complex Sociality, expert consensus among Social Biologists (e.g., B Crespi 2014) have concluded that humans cannot be classified, Eusocial, because this species does not display "reproductive division-of-labor" ... it is worth pointing out that the wide variety of human reproductive groups have not been systematically described, classified, or diagnosed, including, analysis of populations in which more than one architecture is found, such as all modern nation-states [e.g., monogamy, polygamy, promiscuity, etc.] ... whether & how & why these various conformations interact over Time & Space remains to be determined ... it is assumed that 1st Principles [T, E, acquisition, consumption, allocation] relative to abiotic [e.g., climate] & biotic [e.g., food dispersion; breeding sites] environmental regimes, as well as, constraints of limiting resources, as well as, species-typical female traits, always apply ...


1... most Mammals are "sexually segregated" [sometimes termed, "solitary" in the literature]; however, aggregations [temporary clumping of animals] may occur around clumped, limiting resources, especially, food [as, white-tailed deer may do in Spring when food is abundant & clumped]; other aggregations may form when animals migrate from one feeding &/or breeding ground to another; the term "herds" often refers to an aggregation ... the term, "group," is reserved for a reproductive unit ... Hamilton's Rule [rb - c>0 ----> rb > c] is widely accepted as a general formulation of Social Evolution ... according to Hamilton (1964), "social" behavior is limited to Cooperation & Altruism--the only conformations of interindividual interactions [Selfish, Cooperation, Altruism, Spite] whereby Recipient gains in "fitness" ... this definition of "social" should be derivable from Hamilton's equations ...



2... Several authors have claimed that large mammals have, cet. par., generalized phenotypes (e.g., Eisenberg 1981), presumably, related to evolution in heterogeneous regimes ["environmental grain" theory proposed by Richard Levins 1968. Wilson '75 posits that [1] phenotypic flexibility/plasticity is a general feature of social evolution of large mammals [n.b., the evolution of "learning" should be discussed here, including, the evolution of "social" learning requisite to Culture]; however, generalized phenotypes may represent [2] "canalization." Whether [1] or [2] is an effect of temporal variability & "grain" will depend upon patterns of variability [e.g., rainfall, hurricanes] relative to Generation Time [Is the environment trackable?] . For all of the topics in #2, you might want to take a look at my 2012 Springer Brief. Furthermore, to the extent that mammals exhibit generalized phenotypes, this trait would be expected to inhibit evolution of specialized morphologies, such as, morphologically distinct castes. See sections on generalized phenotypes in Eisenberg (1981).

3... Mammalian phenotypes are usually generalized [see Eisenberg op. cit.] ... the evolution of Complex Sociality requires Specialization [division-of-labor; task, role, &/or morphological specialization]; Interdependence [among interacting group members] is a necessary precursor to the evolution of Cooperation ... since Cooperation is a necessary precursor to the evolution of Complex Sociality, specialization must evolve at some point during the Cooperation "grade" ... Hamilton (op. cit.) classifies "cooperation" as an interaction in which both Actor & Recipient benefit reproductively ... as a significant aside, in taxonomic studies, "specialization" is used to diagnose "primitive"/"ancestral" from "derived"/recent characters, where "specialization" is a criterion for "derived" classification ...

4... depending upon environmental regime, an advantage of a generalized phenotype is promotion of phenotypic flexibility [reversible response] &/or phenotypic plasticity [irreversible response]--responses that are usually condition-, context-, situation-dependent ... in extreme &/or highly time-varying environments, responses may be "decided" statistically or probabilistically, even under Hamilton's Rule [op. cit.; often termed "kin selection"] ... it is worth pointing out that a] Hamilton's Rule [rb - c >0 ----> rb>c] does not predict that it is always beneficial to assist the reproduction of kin] the evolution of sociality does not necessarily require individual recognition, e.g., "greenbeards" may operate or "decisions" may be based on probabilities or likelihoods ...

5... the observation that most mammalian phenotypes are generalized is usually attributed to evolution in time-varying environments ... because most mammalian phenotypes are generalized & because Complex Sociality requires the evolution of specialization, it is expected that Complex Sociality, &, possibly, Cooperation, would be uncommon in the Class & in other taxa with generalized phenotypes ... except for cooperatively-breeding mammals & the [eu]social mole rats, [reproductive] division-of-labor [cooperation between specialists] has not been described to date in mammals; task, role, & morphological specialization are described so far only for the social [eusocial] mole rats ... several other species of mammals are potential candidates for specialization [see David Macdonald's edited reviews of Mammal Orders ... in humans, whose phenotypes are generalized in the mammalian mode, the evolution of learning mechanisms permits [learned] specializations such as division-of-labor and task, role specialization ... learning may have evolved in mammals with generalized phenotypes, in part, to minimize the costs of solitary living and maximize the benefits of group-living ... the question of whether morphological specialization has evolved in humans has not, to my knowledge, been systematically studied in humans; however, Race [ecotype] may be a candidate for morphological specialization in Homo sapiens ...

6... since most mammals' phenotypes are generalized, the evolution of Complex Sociality [&, possibly, Cooperation] is expected to be severely constrained in the Class ... on the other hand, some mammal species display specialized phenotypes, & many species display both generalized and specialized traits ... selection may favor some characters [e.g., Learning; phenotypic flexibility (reversible) & phenotypic plasticity (irreversible)] because they minimize the costs &/or maximize the benefits of generalization &/or specialization ... thinking in terms of differential costs & benefits [to "fitness"], research has led to the conclusion that specialists are good competitors but vulnerable to extinction while generalists are poor competitors but comparatively less vulnerable to extinction ...



7... Selander & Kaufman (1973) and others [see paper linked in #1 above] argue that large mammals have higher tolerance for genetic monomorphism. 

8... The basis for Social Evolution in mammals is "milk", according to EO Wilson [1975, Sociobiology, Belknap: HUP] n.b. a number of authors claim that the basis for Social Evolution in mammals is the primitively [non-cooperative, non-altruistic] communal female group leading to polygyny [c.f. Wittenberger 1980]. The basis of Wilson's point, however, is that Mammal females are obligate caretakers of young, predisposing them to certain life-history trajectories, including, probably, receiving help, all limited by energetics. Recall the litany that females are "energy-maximizers," males, "time-minimizers." Some female mammals, extend maternal care [&, often, lifespan], a trait that will retard the evolution of "complex" sociality [specialization, eusociality] because extended maternal care prevents the evolution of reduced maternal care and the evolution of the de-coupling of Survival and Fecundity. Importantly, in humans, social insects, & "primitively" eusocial mole-rats [& "cooperatively-breeding" mammals?], an apparently rare life-history strategy is found [among females] whereby extended lifespan [a "slow" life-history trait] is combined with a high reproductive rate [a "fast" life-history trait]. In humans, however, extended lifespan is, apparently, combined with increased investment in maternal care; while, in social insects, social mole-rats, & "cooperatively-breeding" mammals [?],  extended lifespan in associated with reduced maternal care--a necessary, though, not sufficient, condition for the evolution of higher "grades" of sociality [specialization, eusociality, genotypic altruism]. Further, reduced maternal care is a necessary trait associated with decoupling of Survival & Fecundity.

9... According to Wilson (1975, p 379), the four "pinnacles" of social evolution are: the colonial invertebrates [e.g., corals, bryozoans], the social insects, the non-human mammals, & humans]. n.b. reflective of a "major transitions" approach to the Tree of Life

10... Traits of the non-human mammals: aggressiveness & discord carried further in vertebrate, including, mammal, societies than in social insects...selfishness rules inter-individual interactions...no sterile castes [though reproduction may be suppressed chemically or by "choice"]...acts of altruism infrequent and usually directed toward kin, especially, offspring...cooperation usually rudimentary; though inequity & hierarchy virtually ubiquitous [wherever there is division of labor there will be inequity]...by human standards, some acts "brutal" [e.g., treatment of infirm & dying]...mostly after Wilson '75, p 380

11... Human social traits: humans remain "essentially vertebrate" in their social structure, but more "complex," not by reduction of selfishness but by evolution of intelligence and language [relate to phenotypic plasticity; learning, including, social learning (cultural mechanism); social parasitism; persuasion; manipulation; exploitation, etc]...elaboration of kin relations [cost-benefit]...human societies "approach" levels of cooperation in insect societies & far exceed social insect societies in communication...mostly after Wilson '75, p 380

12... Most "interesting" mammal orders: rodents, artiodactyls, primates, marsupials--"control group" for Class after Wilson '75 & Eisenberg '81 [social biology of largest mammal orders--bats, rodents--virtually unknown as of 1975]...mostly after Wilson '75, p 468 x

13... Traits labile [e.g., mode of intrasexual cooperation, degree of cohesion, openness of societies (c.f. "open" vs. "closed" groups)]. Many forms of interaction change seasonally, & patterns differ at species level. [after Wilson '75]

14... Phylogenetically, more primitive living marsupials and insectivores tend to be solitary. Species that forage nocturnally or underground are, also, mainly solitary. Eisenberg [1981] & Wilson [1975] posit that marsupials are the only "control group" we have for Class Mammalia as a whole.

15... As a rule, the most complex social systems within each Order occur in the physically largest numbers...true of marsupials, rodents, ungulates, carnivores, and primates [Why? maybe because diurnal and because of large brains (intelligence?)? Wilson '75 Also, refer to J.H. Crook's work with weaver birds and other work in Behavioral Ecology: need to evaluate dispersion (distribution and abundance in time and space) of limiting resources relative to behavior & social biology].

16... Species adapted to life in open environments more likely to be social. Rodents known to form coteries of mixed sexes are all inhabitants of grasslands. Ungulates--great herds predominantly found in grasslands and on savannas, though herds mostly loosely structured--but see, horses, mountain sheep, elephants, and a few others [zebras?] Wilson '75

17... The structure of mammal [vertebrate?] groups is thought to result from the tendency of females to select rich patches of food and that of males to select large aggregations of females. Because patch richness and the consequent number [and quality] of females [female group size] is expected to vary, the relative reproductive success [RRS] of females may, also, vary over time and space. See the following paper, for example:

http://eprints.uberibz.org/1173/1/jones_2008.pdf

Also, see:

Jones, CB (March, 1996) Relative reproductive success [RRS] in the mantled howler monkey: implications for conservation. Neotropical Primates 4(1), pp 21-23.

18... Chapters on Mammals in Rubenstein & Abbott (2017), may not be very useful for treatments of general patterns of Social Biology in Mammals; however, see stimulating final, summary chapter in this ambitious edited volume.

19. IMO, the chapters in Wilson (1975) on non-human primates and on humans are not very useful for identifying patterns across the Primates, in Homo sapiens, across Mammals [or, across vertebrates].

20... Tim Clutton-Brock's text, Mammal Societies, not very useful for identifying general patterns of Social Biology in Mammals. This book is not a Synthesis for the Mammals in the way that Wilson's [1971], Insect Societies, is a Synthesis for the Social Insects--even though TC-B titles his book, Mammal Societies. This book is a highly selective literature review. See, however, the useful table of contents & #generateideas. re: sub-eusocial insects--their traits & trends--see James Costa's book, The other insect societies which includes many ideas for research.

21... Neotropical social mammals require thorough treatment, review, and integration into the broader literature on mammals which is dominated by research in the Paleotropics. In South America, see, especially, caviomorph rodents [infraorder, Caviomorpha], uniting all So. American hystricognaths. n.b. all or most of these spp. are small-bodied, & they are related to Old World [hystricognath] social mole rats.

22... General Patterns...Gene Evolution...Convergence...Gene E. Robinson Lab [see Jenny Tung's preliminary, though, cutting-edge, work on mammals @jtung5]:

https://www.pnas.org/content/108/18/7472

23... Temporal division of labor [age (-graded) polyethism: Wilson 1971] is ubiquitous in social insects; how common is it in Mammals & other vertebrates? See [especially, tables]: Jones CB (June, 1996) Temporal division of labor in a primate: age-dependent foraging behavior. Neotropical Primates 4(2): 50-53.

also, Wilson EO (1971) The Insect Societies. Belknap [HUP], Cambridge, MA.

24. The following book by Bernie Crespi discusses correlates of sociality [p 66, pp 73-74]...note, especially, significance of "high value" of limiting resources [abode, mates, food, etc.]...again, shows need to study limiting resource dispersion [distribution & abundance in time & space]...

Crespi BJ, Morris DC, Mound LA (2004) Evolution of ecological and behavioural diversity: Australian Acacia thrips as model organisms. Australian Biological Resources Study, Canberra.

https://www.environment.gov.au/science/abrs/publications/thrips

25... How much of the variation in mammal behavior & social biology is explained by body size?

26. Nursing [mammary glands] predisposes mammal females to caretaking, dependency, subordinate [e.g., "helper"] status, & favoritism of kin &/or other females. But how are these adaptations affected by evolution in suboptimal ["poor"] conditions or conditions of intense competition [for limiting resources], or, say, drought or collapse of prey population?

27. See Part 4 of Eisenberg [1981] for discussion [also, tables] of mammalian behavior, including mating systems & social biology--see, especially, Chapters 31-33.. Eisenberg [1981] classifies mammal "mating systems" into 10 categories, reduced to 3 basic types: polygyny [1 male, >1 females]; polyandry [multiple-male : multiple female], in which females mate with >1 male [not necessarily "promiscuous", i.e., females may demonstrate selectivity or preferences; also, there may be "favoritism" in which an adult female & an adult male are likely to mate exclusively or commonly with each other]; monogamy [uncommon] 

See Eisenberg's outdated but still interesting and classic paper, Eisenberg JF (1966) The social organizations of mammals, Handbuch der Zoologie, Band 8. Lieferung 39: 10(7): 1-92.

28... Feldhamer GA, Drickamer LC, Vessey SH, Merritt JF, Krajewski C (2007) Mammalogy. Johns Hopkins UP, MD: "typical mammalian pattern of preferring unrelated distant kin as mates;" Emlen's "ecological constraints" model + Trivers' reciprocity [+ "pay to stay"?] explain "helping" in mammals?--especially, patterns in social mole rats [do naked mole rats have castes since there is a specialized "disperser form" in this species?]...see Jarman [1974] for classification of mammalian social organization x feeding selectivity: least selective most gregarious, large, polygamous, & sexually dimorphic--Jarmon PJ [1974] The social organization of antelope in relation to their ecology. Behaviour 48: 215-267...need to integrate Jarmon's findings with those of Crook [1964, Behaviour Monograph X] on weaver birds, i.e. relationship between food dispersion [distribution & abundance] and social structure/organization...

29... Ricklefs RE, Miller GL [2000] Ecology, 4th Edition. W.H. Freeman & Co., NY: Part 4, Population Ecology, Section 4.2, The dispersion of individuals reflects habitat heterogeneity and social interactions--"The dispersion of individuals within a population describes their spacing with respect to one another."--clumped [aggregation (maybe poor choice of word)]--clumped [limiting] resources may be most common determinant of these patterns, also, tendency of offspring to remain in natal group --see #13 above; spaced [even spacing= hyperdispersion]; random ["In the absence of social antagonism, which results in a spaced dispersion pattern or mutual attraction, which leads to clumping, individuals may distribute themselves at random" Can use Poisson methods to evaluate dispersion patterns...also, see Index of Dispersion [Clark & Evans, 1954]...also, can evaluate "nearest-neighbor distance"--"interindividual distance" [IID] in primatology literature...again, see Crook, 1964, Behavior Monograph X...

30... When discussing patterns of grouping or "social" organization [in populations], I have the strong impression that Mammalogists tend to attribute the patterns to phylogeny, philopatry [patterns of dispersal x sex], and/or predation...a Crookian [Behavioral Ecology] framework seems not to influenced Mammalogy very much where a "Crookian framework" would entail the mapping of animals in a population on to the dispersion of limiting resources [distribution and abundance of limiting resources in time and space--ceteris paribus & on average relative to differential engergetics x sex--males, at least, adult males, expected to be "time-minimizers," females, at least, adult females, "energy-maximizers"]...

31... The (2007) edited volume on Rodents by Wolff & Sherman [link] should be read in association with John F. Eisenberg's 1981 book on mammalian "radiations" [op. cit.]...Eisenberg provides broad context for many topics [e.g., parental behavior, scent marking, infanticide]...

https://www.amazon.com/s?k=wolff+rodents&i=stripbooks&ref=nb_sb_noss

32... General patterns in Social Mammals [Social Vertebrates? Social Animals?]: EO Wilson [Insect Societies, Belknap (Harvard), 1971] discussed "age polyethism" ["temporal division-of-labor": TDL; "primitive" (totipotent) eusociality] in social insects as a feature of eusociality, particularly, "primitive" eusociality [taxa displaying "totipotency," the capacity to switch from a reproductive to a non-reproductive state & vice versa, as opposed to "advanced" eusociality, taxa characterized by sterile castes]...this article* provides the first empirical evidence of temporal division-of-labor in a mammal [1996], the mantled howler monkey, Alouatta palliata Gray, studied by the present author in Canas, Guanacaste, Costa Rica. Temporal division-of-labor ["age polyethism," "primitive" (totipotent) eusociality] may occur widely among social mammals, social vertebrates, and, possibly, all social animals, where Cooperation and/or Altruism [including, reciprocity] have evolved in group-structured populations...i would speculate that TDL ["age polyethism;" "primitive" (totipotent) eusociality] is more energetically efficient than more opportunistic, and, perhaps, more flexible or plastic, modes of [social] behavioral expression & might be expected to evolve/occur in contexts favoring conservation of energy [e.g., in heterogeneous regimes]...since females are expected to be "energy maximizers" [metabolically, reproductively], TDL may be more likely to be observed in females compared to males ["time-minimizers"]...TDL is expected to be developmentally programmed [within some (statistically normal) species-typical, population-typical, range (x sex)], & conscious & aware processes may not be required for its generation...in ////?, Jones reported age-dependent expressions of conflict-resolution among adult males of the same Alouatta species in the same location & group; these results require intense investigation, lab & field experiments, & modeling, including, for humans...it is important to investigate those conditions under which different energy-conservation strategies evolve x sex...since males are expected to be more robust under energy constraints, it seems likely that thresholds will be identified differentially x sex for the same endogenous and exogenous, including, social, stimuli/factors/conditions.

EO Wilson's 2019 book, Genesis, advances the idea that many [social] Mammals, including, humans [ see**], may be "eusocial." Where "temporal division-of-labor" ["age polyethism"] is demonstrated, taxa can be classified, "primitively eusocial" and/or Totipotent Eusocial [TE] this classification would apply, also, to any other Social Vertebrates or, indeed, to any other Social Animals, where "age polyethism" [TDL; "primitive" eusocial; TE] is identified. If "tradeoffs" [e.g., energetic, reproductive, survival] are most likely to be observed in "poor" conditions [e.g., heterogeneous regimes where "fitness" is compromised; recurrent drought, unpredictable food or water supply], "age polyethism" may evolve to minimize energetic costs in time and space. While females are expected to be most sensitive to energetic effects, males, also, may benefit, under some conditions, from age-dependent responses. Furthermore, there  may be energetic [reproductive, survival] benefits in coordinating many maturational [age-dependent] and developmental [age-dependent] milestones or markers with one another as genetic and physiological energy-savings tactics and strategies.

33... Wolff & Sherman (2007, link) includes some interesting & useful chapters, though, terminology is not standardized [& is often confusing (e.g., "single-breeding," "multiple-breeding"--particularly, as they are discussed in relation to philopatry...see, for example, Chapter 21 by Lacey & Sherman, who needed to make Emlen's work on Ecological Constraints fundamental to their discussion of "philopatry")], and "species" is often employed where "population" is, rather, the correct term. Each chapter in this book has a Summary at the end that can be read before digging into the whole chapter. Some chapters in Sections, "Introduction," "Social Behavior," & "Comparative Sociality" are of particular interest to students of Behavioral Ecology & Social Biology. There are many allusions to the importance of environmental [exogenous] factors, including, dispersion of resources; however, there is no explicit integration of [John Hurrel] Crookian socioecology [Behavior Monograph X] nor is there integration of Emlen's Ecological Constraints model which is fundamental when combined with Crookian socioecology. For context & references (especially, Emlen, Crespi), see blogpost, this blog, of EO Wilson's, Genesis.

Wolff & Sherman [2007], amazon.com:

https://www.amazon.com/Rodent-Societies-Ecological-Evolutionary-Perspective/dp/0226905365/ref=sr_1_1?keywords=jerry+wolff+rodents&qid=1580071530&s=books&sr=1-1

EO Wilson Genesis review [blogpost, this blog]:***

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

34... Fundamental questions in Behavioral Ecology pertain to Population Structure relative to dispersion [distribution & abundance] of limiting resources in T(ime) & S(pace), also to Population Structure relative to Basic Laws of Ecology--Acquisition, Consumption, Allocation. Ultimately, we should be able to reduce "fundamental questions" to Thermal Biology [all Biology reduces to Heat], & all material phenomena reduceable to Physics----->expressible in mathematical terms.

31. Towards general principles [GUTs] via studies of Biological Assembly...for group-living---->sociality, see literature in Physics on "queuing"...

... reintegratingbiology.org/wp-content/upl

...as "assembly" patterns & rules are investigated and identified, it is important to address the Physics literature on "queuing" [see my 2014 Springer Brief on mammalian social evolution, Chapter 4]...

...as "assembly" patterns & rules are investigated and identified, it is important not to ignore Thermal Biology pertaining to Social Biology...

35... Needs lots of research: Resource Partitioning in Group-living vertebrates--search for patterns search for rules [within & between trophic levels, including, differential focus on "frugivores," "carnivores," "omnivores," etc.]

36. Mammalian Patterns [here, Rodents]: Chapter 3 in #Rodent Societies [Jerry Wolff & Paul Sherman, Eds, U of Chicago Press, 2007] concludes that most rodents are polygynous or promiscuous [the author prefers the latter]...the litany for #Mammals is that most mammals are polygynous...

37... Mammalian Patterns, Ch. 4 on female reproductive strategies in Wolff & Sherman, op. cit. does not identify general patterns but exposes several problems inherent to the literature on mammalian sexual selection in this book & in many other treatments [e.g., Mammal Societies by TCB, op. cit.]...for example: 1...the reason treatments of male reproductive strategies dominate the literature is that females are a LIMITING resource for males; males are not a limiting resource for females [------>selection]; 2...many treatments, such as this chapter, do not integrate optimality language and logic into their analyses; 3...many/most treatments are not derived from Crookian model of Behavioral Ecology/Social Biology [dispersion [distribution & abundance] of limiting resources [food, mates, sleeping sites, etc.] in Time & Space; 4...many treatments do not integrate Hamilton's Rule & logic into their analyses; 5...many treatments, especially, in Primatology/Anthropology, do not consider the various hypotheses advanced in an attempt to explain Multiple-Mating by Females [in many reports, only Hrdy's idea is advanced and/or the hypothesis that females mate multiply in order to diversify the genetic composition of litters]; 6...Trivers' 1972 chapter in Campbell's volume seems not to have been digested in many treatments of female reproductive strategies [e.g., mammal females, after parturition and after some threshold investment in "current" offspring, are understood to begin investment in future offspring (e.g., repair, mate choice, fertilization, gestation, etc.)]; this generalization may need to be and has been modified for older/old females; 7...females are, cet. par., "energy-maximizers"; males, "time-minimizers"...

38... this is a classic chapter by Jerry Wolff on alternative reproductive strategies in non-primate mammals...

https://www.cambridge.org/core/books/alternative-reproductive-tactics/alternative-reproductive-tactics-in-nonprimate-male-mammals/88AB2B7352642314032233C426A4A231

39.... following Chapter 7, by John L. Koprowski, in Wolff & Sherman...: "Conflict, both intersexual and intrasexual, has been remarkably influential in the evolution of mating systems of tree squirrels. Conflict among males appears to have been particularly important in promoting evasive behavior in females, led to reduced copulatory duration, promoted alternative reproductive tactics, and influenced postcopulatory tactics for minimizing multiple paternity. Conflict among the sexes likely acts to diminish reproductive success of any dominant individual male, as females mate with multiple males and often avoid active pursuit males. The reproductive skew among males, however, is substantial, and active pursuit males clearly are most successful. Intersexual conflict appears to increase the number of males that are able to mate due directly to the evasive behavior of females, a behavior that maintains the satellite tactic among males. The benefits to females of manipulating the types of competition remain unclear. Future research must quantify the costs and benefits of male and female tactics under different environmental and social conditions...." [95]

40... Gene E. Robinson and other researchers have pointed out that, in social insects, "coordination and control" is decentralized, while, in [most?] human societies, "coordination and control" is centralized...is C & C in [most?] Mammal societies [i.e., in most group-living Mammals], centralized? ... my initial impression based upon my knowledge of all mammalian Orders is that centralized group structure is not common ... dominance hierarchies, for example, would be characterized as a type of "Interdependence," a necessary precursor to social behavior [Cooperation] ... thus, perhaps, the evolution of centralized group architecture is evolutionarily "derived" ...

41... In Wolff & Sherman [op. cit.], Chapter Eight by Dobson & Oli, [using PCA] body mass & phylogeny explain most of the variation in life-history traits [age at maturity; age at last reproduction; juvenile & adult survival; fecundity]...these traits predict a species' position on the "fast-slow [life-history] continuum...should this dataset, also, be analyzed by body size separately--"small" Mammals vs "large" Mammals [unless i am mistaken, a "small" Mammal is considered one < 100 lbs]...

42... in the final analysis, is the most important reason to study Primates the fact that such a large proportion of species is group-living ["social"]--structured around philopatric females, often kin?

43... In Wolff & Sherman [op. cit.], Chapter 13 by Scott Nunes, "dispersal and philopatry" is discussed: "The dispersal process can be categorized as either natal or post-breeding.... In mammals, natal dispersal is more common than breeding dispersal, occurring in nearly all species. Natal dispersal has a strong sex-bias in mammals, with males typically emigrating in new home areas at higher rates or over greater distances than females." [150] "A variety of hypotheses have [sic] been proposes to explain the prevalence of female philopatry in rodents and other mammals.... The evolution of complex social systems among rodents and mammals in general has been suggested to be predicated upon philopatry and cooperation among kin living near each other [especially, philopatric females, often kin]." [162]

44... though authors may mention environmental or ecological factors, in general, few, if any, of the chapters in Wolff & Sherman integrate a Crookian model into their analyses [dispersion (distribution and abundance) of limiting resources in Time & Space]--but see Section, Comparative Socioecology, Chapters 29-37...these excellent and important chapters address Families, Genera, & Species that could reasonably become model systems for the investigation of general mammalian patterns, Behavioral Ecology, & Social Biology [Rattus & Mus are already model systems in research Biology and Medicine]...taxa covered are: ground squirrels; marmots; semifossorial desert rodents; Rattus & Mus; Capybaras and Maras; Octodontid & Ctenomyid rodents; rock-dwelling rodents; African mole-rats; black-tailed gunnison's & Utah prairie dogs...

See Emlen's "ecological constraints" model:

http://max2.ese.u-psud.fr/epc/conservation/PDFs/HIPE/Emlen1982.pdf

See Greenwood's classic 1980 paper on dispersal in birds & mammals:

https://psycnet.apa.org/record/1981-29692-001

45... Wolff & Sherman Chapter 14: Gene dynamics and social behavior [F. Stephen Dobson] uses "F" statistics [Population Genetics] to investigate population structure, finding that, in many mammals, population structure is a function of dispersion of female kin groups...n.b. "social breeding groups may slow the loss of genetic diversity from populations," a process adding to the list of benefits to group-living and, possibly, sociality [cooperation, altruism]...indeed, cooperation & altruism may evolve to insure or enhance these benefits [but what are the tradeoffs, the costs?]...though Anthropologists generally maintain that human sociality evolves from an ancestral state of monogamy, the evidence from Mammals summarized in this blogpost strongly suggests that the ancestral state is the female group, especially, the female kin group, responding to clumped, limiting resources [in heterogeneous regimes?]--as most treatments by mammalogists hold...polygyny [and/or promiscuity?] appear to be the most common mammalian sociosexual system, &, perhaps, as suggested by many researchers, males, in part, monopolize female groups to defend females & their limiting resources [saving energy for female "energy-maximizers"]...as an aside, what role dies Sexual Selection play in all of this...

46... is eusociality much more common than we generally think? Bernie Crespi, WD Hamilton, Nancy Moran quoted in this NY Times article linked below...note importance of protected, reliable food source[s] and of protective, reliable housing [& protection from predators--see Bernie Crespi article linked below]...how about humans?; i have suggested that humans are "facultatively eusocial" in my brief communication, "Are humans cooperative breeders" in Archives of Sexual Behavior discussed on this blog and linked below...

https://www.nytimes.com/1993/02/16/science/social-castes-found-to-be-not-so-rare-in-nature.html

"Are humans cooperative breeders: a call for research" (2011), CB Jones:

https://link.springer.com/article/10.1007/s10508-011-9741-5?shared-article-renderer

...thre conditions for the evolution of eusociality, Bernie Crespi [Are these general?]:

https://www.semanticscholar.org/paper/Three-conditions-for-the-evolution-of-eusociality%3A-Crespi/feb94b7e861ac0e14b8d575f39e33c0a28338419

47... when we speak of "generality" we are often speaking of Convergence--whatever our level[s] of analysis from gene/allele to whole organism...here is link to a now classic paper from Gene E. Robinson's lab; note generalizeability, convergent evolution, & "toolkit" concept...see Table 3.1 in my 2014 Springer brief--phylogenetic & convergent social features of "prehistoric" mammals...

https://www.pnas.org/content/108/18/7472


----------------------------------
*
http://vertebratesocialbehavior.blogspot.com/2020/01/age-polyethism-temporal-division-of.html

**
https://link.springer.com/article/10.1007/s10508-011-9741-5?shared-article-renderer

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




Tuesday, June 25, 2019

Research questions based on Ratnieks et al. 2006 (Clara B. Jones)

Research questions for group-living vertebrates based on Ratnieks et al. 2006*, **

Introduction

Inclusive fitness theory explanations for both cooperation and conflict

What do we mean by conflict resolution?
1. "Potential conflict is any difference in the reproductive optima of individuals or groups within a society."
2. Individuals have three options...to be workers, reproductives, or dependents (e.g., "social parasites") or some combination of these serially or concurrently. Categories can be analyzed by class or rank, by sex, by age, by race or ethnicity, etc.
3. For vertebrates, including, humans, how do we estimate "fitness optima?"

Kinship, coercion, and constraint
"...low relatedness makes wasteful conflicts more likely because the cost of conflict falls upon more distant kin."

Sex allocation
...differential investment in males and/or females & differential costs/benefits...

Conflict-resolution
1. "Coercion and constraint may help to resolve conflict if one party has complete power over sex allocation." ...etc., e.g., over access to limiting resources...
2. "Mixed power seems likely in many but not all situations." ...e.g., "division of task"..."division of labor"...n.b. DoL more efficient than DoT; also note, male & female interests may not coincide since males are expected to be time-minimizers, females, energy-maximizers...
3. Under some conditions, is it more beneficial for an individual to assume the costs of conflict rather than to resolve conflict? ...i.e., When are costs of conflict [over reproduction &/or limiting resources (convertible to offspring] tolerated or preferred relative to benefits of conflict-resolution?
4. Importance of information, especially, competition for information [see EO Wilson 1971 on "communication" in societies]...Humans: language, who shares information with whom, etc...
5. Investigate "policing"*** as a form of conflict-resolution [mitigation of selfishness], mitigation of conflict [e.g., force, coercion, constraint, persuasion, etc. (see French & Raven's "bases of power")]...
6. What factors lower incentives for individual selfishness [& subsequent "tragedy of the commons"]
7. High "r" lowers the proportion of reproductives in a population...
See "reproductive skew" literature, e.g., R Hager & CB Jones 2009 CUP...

Conflicts among totipotent individuals
1. "Totipotency greatly increases the potential for conflict...."
2. If optimal family and/or group size can be estimated, excess individuals are likely to increase potential for conflict...

Discussion & Conclusions
1. Class conflicts relatively easy to resolve [because dominants "police" subordinates?]...
2. Conflict easily resolved when one party "powerless"...
3. Should the incorporation of steriles or other non-reproductives in groups reduce conflict?...relative to what [e.g., "r"]...
4. How do humans escape control in groups? [see French & Raven's "bases of power"]
5. What role does genetics play in conflict-resolution [e.g., genetic conflict]...

*Ratnieks FIW, Foster KR, Wenseleers T (2006) Conflict resolution in insect societies. Annu. Rev. Entomol. 51: 581-608.****
**Sentences in quotation marks are quotations from paper; other statements, comments, or questions are my interpolations relative to group-living vertebrates, including, humans.
***New paper: "'Enforcement' is central to evolution of cooperation"...i.e., &, also, to evolution of repression of selfishness... https://www.nature.com/articles/s41559-019-0907-1

****Topics/questions based on titles of literature cited:
--evolution of male and female traits
--causes & consequences of informational constraint
--conflict over class/SES determination
--status-allocation to offspring
--Does it "pay' kin to favor relatives who are "losers" or inferior reproductives and/or competitors?
--n.b. Charnov EL (1978) Evolution of eusocial behavior: offspring choice or parental parasitism? JTB 75: 451-465.
--n.b. Crespi BJ, Ragsdale JE (2000) A skew model for the evolution of sociality via manipulation: why it is better to be feared than loved. Proc R. Soc. London B Biol. Sci. 267: 821-828.
--facultative policing
--conflict over paternity [n.b. causes & consequences of multiple-mating by females]
--n.b. Keller L (1997) Indiscriminate altruism: unduly nice parents and siblings. TREE 12: 99-103.
--genes regulating complex social behavior
--ancestral states of complex sociality
--sex-ratio determination in groups & populations [e.g., conflict over sex-ratio determination]
--causes & consequences of sexual deception [e.g., in some mammal species, males are indistinguishable from adult females for some time before expressing secondary sexual characteristics (see, e.g., mantled howler monkeys, Alouatta palliata)]
--selfish tactics & strategies that promote sociality [e.g., "social cohesion"]
--implications: "heirs & spares"
--mechanisms of conflict-management & conflict-resolution in eusocial taxa
--n.b. Ratnieks FLW, Wenseleers T (2005) Policing in insect societies. Science 307: 54-56.
--alternative reproductive strategies and conflict-resolution/conflict-management
--n.b. Sachs JL, Mueller UG, Wilcox TP, Bull JJ (2004) The evolution of cooperation. Q. Rev. Biol. 79: 135-160.
--policing x age, sex, class/SES, race, etc.
--Is multiple-mating by females rare in vertebrates?
--causes & effects of totipotency; ubiquitous in vertebrates?
--genetic basis of sterility
--mechanisms of kin discrimination
--What traits differentiate members of classes [within & between]?
--relevance of behavioral flexibility & phenotypic plasticity to conflict-management & conflict-resolution [also, flexibility of group structures]



Thursday, April 11, 2019

Review of Tim Clutton-Brock's, Mammal Societies (by Clara B. Jones, 2016)

Mammal Societies
Tim Clutton-Brock
2016
Wiley-Blackwell (Oxford, UK)
744 pp
ISBN 97811119095323


“The key to the sociobiology of mammals is milk.” E.O. Wilson (1975)

Reviewed by Clara B. Jones (2016; revised 2020)*

Knowledge about group-living mammals may contribute to an understanding of vertebrate social evolution and the evolution of gregariousness in animals with generalized phenotypes [specialization being a signature of high "grades" of social evolution (specialization, reproductive division-of-labor]. Compared to social insects and birds, the social biology of mammals is poorly known with the exception of ungulates, carnivores, and primates (3 of ~25 Orders). Among many similar papers in the mammalian literature on Social Paleontology, in 2011, Ladevèze et al. reported fossil evidence appearing to document mammalian gregariousness and its associated ecology from the basal Tertiary of Bolivia. These findings suggested that extinct, marsupial-like Pucadelphys andinus were group-living, probably exhibiting frequent interactions, strong sexual dimorphism, and male-male competition, as well as, polygyny. Based on the spatial and ecological settings of their specimens, as well as, the climate, in addition to, physical and situational associations and patterning of adult, sub-adult, and juvenile remains in their sample, these authors speculated that the species may have been "social" [gregarious]. In 2012, employing phylogenetic analyses, Briga et al. showed that relatedness and allomaternal¹ care are positively correlated in Class Mammalia. These papers indicate that, though the population dispersion of most extant mammals is sexually segregated (“solitary”) and though fossil remains cannot definitively preserve Behavior, group-living may have a long history in this Class (also see Jones 2014, Table 3.1, pp 19-25).

Tim Clutton-Brock (henceforth, “TC-B”) is a highly-regarded empiricist at the University of Cambridge (UK), recognized, particularly, for his field studies on primates, red deer, and meerkats. He is a prolific scientist with a knack for asking good questions and choosing animal models that have yielded flagship research. The author will be familiar to most animal behaviorists and behavioral ecologists as a specialist of cooperative breeding and evolutionary aspects of reproduction (e.g., female mating strategies, sexual selection). In the book under review, TC-B notes that his undergraduate training was in Anthropology and that he completed his doctorate under Robert Hinde, an animal behaviorist that Psychology typically claims as one of its own. I have been familiar with TC-B's work since the 1970s, and my personal favorites among his copious publications are his 1995 paper with Geoff Parker and the 2003 volume edited with R.M. Sibley & J. Hone. I am pleased to have the opportunity to review Mammal Societies. I have interacted with TC-B on several occasions, once face-to-face, and, more than once, via e-mail. He has always been generous and courteous to me. 

Previous books by JH Crook, “Griff” Ewer, J Eisenberg, EO Wilson, R Estes, D MacDonald, CB Jones, and others, have treated mammalian social biology to one degree or another. Mammal Societies, however, is the first attempt to provide a comprehensive literature review of the topic. The publisher's description of the volume states that it is intended for “behavioral ecologists, ecologists, and anthropologists,” and TC-B self-identifies as a “behavioral ecologist.” The book is, to all purposes, a literature review emphasizing publications on Old World taxa [a tradition attributable, in particular, to Primatology]. While the Table of Contents presents a detailed outline of topics of interest to social biologists, the book is not organized using ecological [e.g., JH Crook] or evolutionary models [e.g., the "major transitions"]. To provide context, professors using Mammal Societies as a course textbook or reference work are strongly advised to acquaint their students early on with Wilson's (1975) treatment (pp 456-574) presenting an explicitly articulated conceptual framework for mammalian social biology, including, trends, conventional terminology, general and comparative features in the Class, an extensive glossary, as well as, case studies and summary tables, figures, and diagrams. John Eisenberg's [1981], "mammalian radiations," is, also, an invaluable source of information on mammalian patterns, including, behavior, as well as, mating & group architectures.

Chapter 1, “Social evolution,” omits definitions of terms the first time they appear in the book (e.g., “aggregation,” “social”, “cooperation”), leading to obfuscation throughout, particularly, since there is no discussion of how to measure social traits (cooperation, altruism) and to discuss their pertinence to reproductive success. In this chapter, the author might have defined “Mammal” and should tell the reader why mammalian social biology is of import. The reader will want to understand possible trajectories to cooperation and altruism from aggregations to group-formation to group-maintenance and how the (spatial and temporal) distribution of limiting resources favor or disfavor the evolution of mammalian sociality. Chapter 1 is, in great part, a selective account of the history of Animal Behavior combined with some mention of theoretical issues (e.g., Darwinism, competition, reciprocity, game theory). However, for rigorous discussions of verbal and quantitative theory in Behavioral Ecology, as well as, overviews of Methods and G x E interactions, readers are referred to Davies et al. (2012) and Westneat & Fox (2010).

Chapters 2-9 address topics related to features of female behavior, particularly, as they pertain to mating, maternal tendencies, and gregariousness. Focusing on females, their strategies, and their energetic requirements as the primary driver of group-living and patterns of male behavior and dispersion is fundamental to an understanding of mammal societies because fertilizable females are a limiting resource for males and, subsequently, an ultimate determinant of male “fitness." Though these and other important concepts are implicit in some of TC-B's discussions, explicit use of many principles inherent to Behavioral Ecology are unclear or lacking (e.g., integration of Hamilton's rule [rb > c] throughout chapters, acknowledgment of the many competing hypotheses in Ecology pertaining to dispersal or multiple-mating by females, use of optimality formulations). As an example from Chapter 5 (“Maternal care”), TC-B's treatment asserts, accurately, that mammalian females invest heavily in current offspring, but theory holds that, after parturition, female resources, above some critical minimum, are channeled into future reproduction and lifetime reproductive success--“fitness.”

Chapters 10-16 pertain to males, especially, mating strategies, relations with females, and paternal care. Characteristic of Mammal Societies as a whole, these chapters are literature reviews of mostly descriptive publications from the Animal Behavior literature. Life history evolution is addressed in this chapter without mentioning the importance of tradeoffs, the distinction between semelparity and iteroparity (“fast” and “slow” life history trajectories, respectively), the importance of life-tables and the role of mortality as a driver of life-history evolution (Stearns 2000). Chapter 17 reviews “Cooperative breeding,” one of TC-B's specializations, and Chapter 18 presents a discussion of “Sex differences." Throughout the book, the author impresses the reader with the centrality of sex, sexual competition, and mating—topics of import in TC-B's career, though one is surprised that more attention is not given to Sexual Selection, per se. Chapters 19 and 20 address hominoids and hominids, including, modern humans, topics often missing or skimmed in other Animal Behavior texts.

TC-B presents at least one controversial formulation in Mammal Societies by asserting, with no supporting evidence or logical arguments, that no mammals are “eusocial”²—that the highest grade of sociality in mammals is “cooperative breeding.” This view is orthogonal to standard practice in Mammalian Social Biology whereby the social mole rats are typically classified as “primitively” eusocial. Technically, according to common usage, “cooperative breeders” might, as well, be classified “primitively” eusocial because of the presence of reproductive division of labor [cooperation between specialists] in the form of totipotent “helpers” (see Jones 2014, p 48-52). Mammal Societies exemplifies the need for practitioners of Natural History, Animal Behavior, and Behavioral Ecology to revisit topics such as standardization of terminology, advancement of the Hamiltonian Project, the roles of quantitative theory and modeling (in particular, agent-based modeling), field experiments, as well as, hypothesis-testing [including, the role of null-hypotheses] based on 1st principles. The text will appeal to professors wanting a Natural History, mostly, non-quantitative, review allowing supplementary reading to be incorporated into a syllabus. Future syntheses of Mammalian Social Biology will rely on mainstream schemas from Ecology & Evolution, in particular, employing a Major Transitions Approach (cf. West et al. 2015), in addition to, Population Ecology, of which Behavioral Ecology is a sub-field.

¹Care of offspring by conspecifics other than the mother
²”The evolution of eusociality, here defined as the emergence of societies with reproductive division of labour and cooperative brood care, has occurred under specific ecological, genetic, and life history conditions. Although sophisticated levels of cooperation have evolved in the largest and more complex societies, conflicts among individuals are still common because, in contrast to cells of an organism, they are not genetically identical,”--i.e., not "clones" (Keller & Chapuisat, 2010)

References

Bradbury JW (1981) The evolution of leks. In Natural selection and social behavior. (RD Alexander, DW Tinkle, eds). Chiron Press, New York, pp 138-169.

Briga M, Pen I, Wright J (2012) Care for kin: within-group relatedness and allomaternal care are positively correlated and conserved throughout the mammalian phylogeny. Biology Letters: p.rsbl20120159

Clutton-Brock TH (2021) Social evolution in mammals. Science 373(6561): doi:10.1126/science.abc9699. 

Clutton-Brock TH, Parker GA (1995) Punishment in animal societies. Nature 373: 209-216.

Davies NB, Krebs JR, West SA (2012) Introduction to behavioral ecology. Wiley-Blackwell, 4th edition. Oxford, UK.

Eisenberg JF (1981) Mammalian radiations. U Chicago Press.

Jones CB (2014) Evolution of mammalian sociality in an ecological perspective. Springer, New York.

Keller L, Chapuisat M (2010) Eusociality and cooperation. In Encyclopedia of life sciences. Macmillan, published online: DOI: 10.1002/9780470015902.a0003670.pub

Ladevèze S, de Muizon C, Beck RMD, Germain D, Cespedes-Paz R (2011) Earliest evidence of mammalian social behaviour in the basal Tertiary of Bolivia. Nature 474: 83-86.

Sibley RM, Hone J, Clutton-Brock TH (eds) (2003) Wildlife population growth rates. The Royal Society: Cambridge University Press, Cambridge University Press, UK.

Stearns SC (2000) Life history evolution: successes, limitations, and prospects. Naturwissenschaften 87: 476-486.

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

Westneat D, Fox C (eds) (2010) Evolutionary behavioral ecology. Oxford University Press, Oxford University Press, UK.

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

Wilson EO (1975) Sociobiology: the new synthesis. Belknap (Harvard), Cambridge, MA.


*Originally published in International Society for Behavioral Ecology Newsletter, 2016.


Addendum: I want to apologize to Tim Clutton-Brock for this review which expresses, as much as anything else, my disappointment in his text--of which I had great expectations. My expectations were so high--I expected a synthetic masterpiece in the context of evolution and the Major Transitions Approach--something on the order of EO Wilson's, The Insect Societies (1971), but, for Mammals. Even a book like JF Eisenberg's, mammalian "radiations" (1981), but about mammalian social evolution--extending Eisenberg's relevant chapters--would have been a major advance for the field. Instead, Clutton-Brock's book, whose title is a misnomer, is little more than a review of many highly selected publications without a conceptual framework [e.g., The Hamiltonian Project (the General Law of Social Evolution); Major Transitions, including, Sociality (Cooperation, after Hamilton 11964), as well as, Complex Sociality: SPECIALIZATION: division-of-labor (Cooperation between specialists); reproductive DoL, task, role, &/or morphological specialization] without any treatment of general patterns, without standardization of terminology, with only a nod to, systematic, quantitative theory--including, Hamilton's rule, and all in the context of a colonial historical-academic framework [e.g., every photograph of people of African descent is of primitive groups save one, and that one of an African guide, apparently, on a hunting safari with white male tourists. Unless I am mistaken, only a very few women are highlighted. Surely, TC-B might have provided a nod to changing worldviews by highlighting one or more of his African & female colleagues who must exist since TC-B has been conducting research in Africa for decades; if TC-B has not had African collaborators, volumes are silently spoken [admittedly, I know nothing about his personal or academic or research affiliations or his views on race, ethnicity, class or gender; and, I "get" it--the "greats" in Animal Behavior are, mostly, white men--even, until recently, in Anthropology, & I defer to the historical record ... nonetheless, it would have been generous to include, say, "Griff" Ewer, as deserving of recognition, for example--if only for the sake of appearances, but, then, my own biases are showing; in the final event, it would not detract from C-B's legacy for him to have been more generous in his historical acknowledgements].In short, as one who has admired TC-B's exhaustively rendered empirical work for decades, my expectations and anticipations, as well as, intellectual curiosity, were totally deflated by the long  literature review treated herein. A synthesis of mammalian Ecology & Evolutionary Biology, in other words, a text on Mammal Societies, awaits future treatment. John F. Eisenberg's 1981 book, "radiations," is the closest thing we have so far to a synthesis of mammalian social biology. 5/9/2020; slight edit, 12/30/2023 ...

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