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The Pursuit of Human Nature in Sociobiology and Evolutionary Sociology Author(s): Alexandra Maryanski Reviewed work(s): Source: Sociological Perspectives, Vol. 37, No. 3 (Autumn, 1994), pp. 375-389 Published by: University of California Press Stable URL: http://www.jstor.org/stable/1389502 . Accessed: 28/07/2012 08:00
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Sociological Perspectives

Vol. 37, No. 3, pp. 375-389 ISSN0731-1214

THE PURSUIT OF HUMAN NATURE IN SOCIOBIOLOGYAND EVOLUTIONARYSOCIOLOGY

ALEXANDRA MARYANSKI University of California, Riverside

ABSTRACT: Outsideof sociology, evolutionary theoryis once again in attention social science. sat Having outthespirited commanding widespread over in are debates sociobiology the1970s,mostsociologists largely unaware thatthefieldhas prospered is nowa respected, and science interdisciplinary witha growing number influential scholars within social the This sciences. of article takesa critical at sociobiology a consideration bothits look with of historical and theoretical whichis exemplified stance, origins its nowmodified In article'Toward Neo-Darwinian a Crippen's by Timothy Sociology." an this that addition, essaysummarizes alternative approach mightbecalled it While alsoincorporates Modern the it "evolutionary sociology." Synthesis, usesestablished methods theory, and withprimate and data sociological along the fossil and archaeological to the records, consider biological legacyof humankind.

Manactson.& is acted by[the]organic inorganic on and like agents thisearth. every of other animal. CharlesDarwin(Notebook 1838-1839) E, A range of opinions now exists on whether or not to incorporatebiology into sociology. One option is to ignore biologicalthinking,much as sociologists did in 1975 after E.O. Wilson introduced "sociobiology"as a new science that was to "reformulate foundationsof the social sciences."Although Wilson questioned the at the time whether the social sciences could be "trulybiologicized"for inclusion into the Modem Synthesis, sociobiologicalmethods and theory are now well entrenchedin the social sciences,especiallyin researchprogramsin anthropology and psychology. In contrast, few sociologists have welcomed what some have called a "new paradigm"(van den Berghe1983;Barash1982).Having sat out the
* Directall to: of correspondence Alexandra Riverside, Maryanski, Department Sociology,Universityof California, CA92521-0419.

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heated discussions over sociobiology in the 1970s, many sociologists are unaware that sociobiology is now a reputable discipline with theories that command widespread attention. Moreover, the discipline itself is now carved up into various subfields which include evolutionary biology, behavioral ecology and evolutionary psychology, with a noticeable cleavage between those committed to the early biological determinism of "hard-core" sociobiology and the more "soft-jointed" sociobiology with its moderate mode and coemphasis on genes and learning. Thus, despite predictions to the contrary, sociobiology has prospered. One objective of this article is to persuade sociologists to consider a more open scientific attitude toward biological arguments, if only to reject them once again. Let me begin with a critical review of sociobiology theory by sketching its historical background, basic logic, key assumptions, successes, and enduring problems. Then, Timothy Crippen's article "Toward a Neo-Darwinian Sociology" (1994, this issue) will be used to illustrate sociobiology's "new look" which, Crippen hopes, will make it more appealing to sociologists and more in touch with observable reality. The second objective of this essay is to present an alternative to sociobiology by outlining a strategy which could be called "evolutionary sociology." This alternative draws from sociobiology in its concern with how biological processes impinge upon humans and human societies and in its use of the Moder Synthesis. However, it differs from sociobiology in its nonreductionist stance, its compatibility with sociological methods and theory, and its unwillingness to build in preconceived notions about human nature. SOCIOBIOLOGY: HISTORY, GENERAL PRINCIPALS, ACHIEVEMENTS, AND SOME ENDURING PROBLEMS Historical Synopsis The basis for sociobiology was forged with the synthesis of two historical traditions-theoretical genetics and selectionist theory. It was into the antiselectionist milieu of the 1920s that R.A. Fisher (1930) triggered the restoration of Darwin's selectionist doctrine and, simultaneously, laid the footings for of Theory NaturalSelection sociobiology. What Fisher essentially did in his TheGenetical was to convincingly refute the then popular "mutation theory" "or saltation theory" (see de Vries 1909-1910) by illustrating that organisms are, in general, intricately adapted to their environment and that, as a consequence, large mutations would be generally harmful (Fisher 1930:41).By restructuring Darwinian ideas on natural selection and casting them in terms of population genetics, Fisher restored natural selection to its rightful place as the central force in evolution. Above all, Fisher emphasized the overriding importance of fitness (defined as the average worth in reproductive terms for each genotype) in his "Fundamental Theorem of Natural Selection," which asserts that: "The rate of increase in fitness of any organism at any time is equal to its genetic variance in fitness at that time" (Fisher 1930:35). In other words, the mean fitness of a population will generally

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be proportional to some component of the genetic variance, and the rate of evolution will also be confined to how much variance exists in the breeding contributionsby S.W.Wright(1931)and populationor gene pool. Otherformidable Haldine (1932)followed Fisher's lead by integratingselection theory with J.B.S. Mendeliangenetics. The constructivework of Fisher,Haldine,and Wrightfirmly models of these strappedselection to a genetic foundation,while the mathematical mainarchitectsgave precisionto the processesof Darwinianevolutionby allowing for quantitativemeasurementof the effect of selection on gene frequencies.And, research theirpioneerworkprovidedthe blueprintforthe broad-based historically, foundationthat eventually crystallizedinto the "Modem Synthesis"in the 1940s (fordiscussions, see Mayr1942;Huxley 1942;Dobzhansky1937). Whereas the Modem Synthesis underscoredthe role of selection for physical But selection itselfwas rarelyinvoked for socialcharacters. this was characteristics, to change in 1962over the question of why some animalsshow altruistic behaviors, an altruistbeing someone who acts to help others at some cost to his or her own in fitness.This problemwas firsttackledby W.C.Wynne-Edwards his classicwork to In what is essentially a in Animal Dispersion Relation Social Behavior (1962). Durkheimianmodel, Wynne-Edwardsargued that although selection normally acts on individuals,in groupliving the needs of the collectiveoften takeprecedence over individual needs; thus, social traits that involve mutual cooperation are selected for at the population level, with the "strugglefor existence" shifting to behaviorsevolved to promote the grouplevel. ForWynne-Edwards, then, altruistic with selectionoperatingupon the "goodof the group"or the "goodof the species," variations between groups through differentialextinction, or "group selection" 1962). (Wynne-Edwards The academickickofffor sociobiology came when G. Williams,W.D.Hamilton, and R. Triversmarshaledan attackagainst group selectionisttheory. In Adaptation andNatural Selection (1966),Williamsdismantled the group-selectionistargument by logicallydemonstratingthat naturalselection workingon individualsalone can account for all the phenomena proposed by group selection. Rather than hypothetical emergent properties,Williamsstated that group level adaptations actuallyreflectindividualselection,with most gregariousnessand other collective summationof individualadaptation" (Williams phenomena merely the "statistical Williamswas equally pervasivein casting aside the individualorganism 1966:258). as the actualselected entity, arguing that because the genotype and phenotype are only temporary manifestationsunique to each individual, the selected unit must, therefore,evidence a higherrate of stabilityand a lower rate of endogamous change. For Williams,the gene qualifiedas the only entity to meet these criteria Williams also repeated Fisher's earlier emphasis on fitness, (Williams1966:23). contending that all adaptationsmust relate to reproductionwith every part of an animal organized for some function to meet the ultimate goal of reproductive success (Williams1966). Hamiltonresolved the problem of why animals display altruisticbehaviorsby invoking a pseudo-form of group selection. In his "The Evolution of Altruistic

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Behavior," Hamilton (1963) introduced "kin selection" as a concept that accounted for cooperation and self-sacrifice through a quantity he labeled "inclusive fitness." This concept rests on the premise that blood relatives share many identical genes inherited from one or more ancestors. Hamilton alleged that selection will positively favor the evolution of "altruistic" social behaviors when individuals aid close relatives; for in this way their own fitness can also be maximized by helping those who share some identical genes. Finally, Trivers, in his "The Evolution of Reciprocal Altruism" (1971), attacked the problem of why unrelated individuals still engaged in altruistic behaviors. In a number of sophisticated modeling procedures, Trivers showed that altruistic behaviors are likely to evolve in any social species when the following biological parameters exist: (1) a long lifetime to maximize the chances of altruistic encounters; (2) a low dispersal rate to increase the chances that individuals will interact repeatedly with the same set of conspecifics; and (3) a group-living population to increase the degree of mutual dependence among individuals. Under the above conditions, altruistic behaviors are likely to occur because of the potential reciprocity in future interactions. Thus, unlike kin selection where fitness is optimized through the helping of relatives with cognate genes, in reciprocal altruism nonrelatives can also help to maximize fitness, but only when individuals are reasonably sure that circumstances will permit a return on their altruism (Trivers 1971:37-39; see Rosenberg 1992 for an excellent appraisal of biological altruism). Finally, a number of influential scholars worked to codify the approach into a new scientific discipline. Among the most influential were J.Maynard-Smith (1974), who merged game theory with genetic evolutionary theory to describe the process of fitness; Richard Dawkins (1976), who wrote a popularized version of William's "genic selection" with his portrayal of human bodies as robot vehicles blindly programmed to serve selfish genes; R.D. Alexander (1971), who underscored the importance of viewing humans as organisms whose behaviors evolved to maximize The reproductive fitness; and E.O. Wilson, whose magum opus Sociobiology: New Synthesis (1975) formally labeled the field, drew together its central concepts, and established it as a new research program with a voluminous amount of empirical data (for historical discussions, see Barlow 1991; Maryanski and Turner 1991). Sociobiology's Theory and Applications Sociobiology has greatly enlarged its scope within the last fifteen years in traditional biology and zoology, as well as in the social sciences, in part because of its novelty, theoretical sophistication, elegant mathematical models, innovative concepts, and dynamic evolutionary thrust. Those directly under the sociobiology umbrella can normally be identified by their commitment to a strategy metaphor, their strict application of Darwinian selection, their focus on ultimate causation, and their application of such pivotal concepts as kin selection, inclusive fitness, and reciprocal altruism. Essentially, sociobiology is a theory-driven science that is rooted in the evolutionary principle that the ultimate goal of all organisms is

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to survive and reproduce.However, in place of the classic selection model which conjures up an image of relatively passive individuals being acted upon by the environment,sociobiologistsconcentrateon the interactionsamong conspecifics and portray organisms as selfish and calculating actors who actively seek to maximize their reproductive success by any means imaginable (although no consciousintent or moralundertonesareever implied).In these efforts,actorsmust adopt strategieswith differentrelative outcomes under particularcircumstances. Sociobiology has also revamped the selectionist doctrine by deemphasizing the individualas the unit of selection and highlighting"selfish" genes, which Williams had earlierhighlighted as the units actuallyaffectedby the selection process. But, Hamiltonunquestionablyforced this shift with his "inclusivefitness"concept as the additive quantity that could maximizeindividualfitness by helping relatives who share identicalgenes. In applying this strategy, sociobiologistsrely upon the following logic: In the past, natural selection favored particularkinds of behavioraladaptations which increased the fitnesses of individuals in a given environment. Nonadaptive behaviors were selected out long ago, leaving in place morphologicaland social traits that now serve the functionalgoal of maximizingfitness. Researchersthen seek to discover how particular social behaviors are adaptive for a fitness payoff. Generally, sociobiologists focus on particulargroups of behaviors where the influence of kin selection and reciprocalaltruismcan be examined and, then, be associated(throughproximatefactors)with reproductivesuccess. Forexample,in behavioralecology (a subfieldof sociobiology),animalresearcherscollect data on specificbehavioralpatterns,with the comparativemethod used to assess whether species differences are related to ecological differences. Hypotheses are then formulatedabout particularbehavioralstrategies;and experimentsare designed to assess the "costs" and "benefits"of different behavioral strategies. Finally, decisions and to make optimalitymodels are then constructedto analyze "fitness" strategy(fordiscussions, quantitativepredictionsabout the benefitsof a particular see Krebsand Davies 1987;Gray1985).Some outcomes which are framedin terms of "costs"and "benefits" (1974,1978)concept actuallyrely upon Maynard-Smith's of an "evolutionary stable strategy"(ESS), which uses game theory forfixed fitness payoffs. Here, the optimal strategy will hinge upon "net benefit" since it must include the tactics of a number of conspecificswho play out differentstrategies within a given social field. In human sociobiology, earlier models of absolute determinism are usually models which involve an interface supplanted with relative "dual-inheritance" between genes and culture. Most sociobiologists now assume that human behaviors are highly flexible in their expression and that humans have a much greaterlatitudeforlearningand choice than other animals.However,in theirview, human action is still the result of past adaptations and, hence, humans are of predisposed to assess the "costs"and "benefits" social interactions.Thus, even cooperativeand altruisticbehaviorsareviewed as adaptivestrategiesto maximize payoffs for "selfishgenes" that compete in the "strugglefor survival."

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Sociobiology'sSuccesses Sociobiology theory is used to interpret a variety of social behaviors among animalsand humans. For example,evolutionarymodels have been developed to explain the existence of dominance hierarchiesin female rhesus monkeys as a mechanismfor femalesto optimize their inclusive fitness (Chapais1981).Also, the rangingpatternsof male and femaleprimateshave been shown to follow different food resources,with findings reproductivestrategiesthroughthe choiceof different in line with expected sociobiologypredictions(Wrangham 1979).The significance of kinship bonds in social living animals has been particularlywell-established. For example,in bird species the "inclusivefitness"concept has been successfully upheld with documented field studies showing that in some bird colonies, individualsforegotheir own directindividualfitness and turn to helping close kin rear their offspring by alloparenting (becoming helpers) (Woolfenden and Fitzpatrick 1984; Krebs and Davies 1987). Also, in agonistic conflicts among primates,close relatives often aid each other much more than distant relatives (Kurland1977;see also Gray1985). In human sociobiology,researchersfocus on a range of social issues where key sociobiological concepts can be applied.Forexample,to accountforthe widespread and altruismamong humans, kin selection theory has been widely cooperation applied to such traditional sociological topics as murder, racism, infanticide, despotism, male dominance,gender roles, sexual jealously, mate selection, rape, child abuse,and parentalcare,with predictionsthat relateto fitness maximization. Formal game theory employing the notion of inclusive fitness is often used to establish the conditions under which particularforms of altruismcan evolve, or conversely,the cause for constraintson kinshipinvestment.Forexample,parental investment models are used in conjunction with cross-cultural data to test hypotheses that include such variables as the relationship between parental investment and paternal certainty (which is predicted to effect residence and descent rules),the degree of genetic relatednessand child abuse (with predictions that biologicalchildrenareless likelyto be abused),the degree of genetic closeness and the transmissionof property (with the rule of inheritanceconsidered a kind of parentalinvestment),the relationshipbetween the likelihoodof infanticideand chronologicalage (with the predictionthat the younger the child, the more likely s/he is to be a victimof infanticide), the probability maleor femaleinfanticide and of (with predictionsthat female infanticidewill likely prevail with scarce resources since males are often favored for inheritanceand have a greateroverall fertility potential)(see, for example,Voland 1984;Dickemann1985;Daly and Wilson 1981, Archer1991). 1984,1988;van den Berghe1990; On Fitness Maximizationand Other EnduringProblems One of sociobiology's great virtues is its clear methodologicalprotocol which addresses broad and basic questions in terms of one set of coherent concepts. As a result,investigatorsfollowa systematicresearchprocedurewhich, in turn,allows

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scholarsfromdiverse fields to convergeupon a common set of issues. Sociobiology has also brought some fresh ways of looking at sociality, forcing scientists to become more sensitive to how selection pressures might operate to produce adaptationsthat enhance reproductivefitness. Nevertheless,criticismcontinues to be leveled at sociobiological arguments(see, for example, Lernerand von Eye 1992;Kitcher1985;Saunders 1988;Travis and As Yeager1991). long as this theory was appliedto such organismsas socialinsects, it was successful. But when researchersused sociobiology theory to explain the complexityof socialbehaviorsin highermammals,problemsbegan to emerge.One problemis whether or not all social behavioris adaptive. Generally,for a trait to be consideredan adaptation,it mustbe shown to have been historicallycast-that is, "if and only if it is the product of selection for the trait in question" (Burian 1992:8,his emphasis; see also West-Eberhard1992). This point has been most emphatically argued by Stephen Jay Gould and Elizabeth Vrba (1982), who maintain that sociobiology is dominated by an "overly adaptationist neoDarwiniantheory"which ignoresthe crucialdistinctionbetween historicalgenesis and currentutility. Charactersutilized today may indeed have been selected for in the past, but a trait'sformerfunction might now be usurped;or alternatively, a once-adaptivetraitmight now be converted for other functionsthan its original utility;or, perhaps a charactermight be importantnow, but it may have arisen originallyby being entailed to other structuresand then co-opted for currentuse (Gould 1991).In human sociobiology, this issue is especially pertinent because humans evolved in small bands of less than 100 individualsand lived as nomadic food-collectors.Even if we grant the assumption that all current behaviors are adaptive, the shift from food foraging to a sedentary lifestyle without a correspondingshift in human biology is likely to have skewed the directhistorical relationbetween behaviorand adaptation. A second majorimpediment is the validity of sociobiology's core assumption to that organisms are alwaysseeking enhance the number of genes they leave behind whether directly through their own offspring, indirectly through the offspring of their relatives, or through a reciprocal exchange of services with nonrelatives.Yet, in long-lived animals with complex social behaviors, how can fitness be measured?How can it be determinedif a behaviorinvolves "reciprocal altruism"?How can we begin to assess the actual "costs"and "benefits"of any behavior in terms of later reproductivesuccess? Sociobiologiststry to surmount these obstacles with probability models and game theory to predict expected Darwinianfitness throughthe playing of particular strategies,makingevery effort to test their hypotheses with comparative data and (when applicable) crossspecies data. By fitting the data to the theory, findings are often in the predicted direction,but most are limited to making simple associations (see Archer 1991). Criticschargethat sociobiology narrativesare easy to constructand provide only pseudo-explanations that simply confirm unsubstantiated assumptions. Also, analogous to classical sociological functionalism,sociobiology explanations can slip into tautologies: Why does a social behavior exist? It exists because it

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promotes fitness. How do we know that it promotes fitness? It promotes fitness because, otherwise, it would not exist. In human sociobiology, "fitness maximization" generates heckling even within sociobiology, with some critics charging that "there are absolutely no grounds for proposing such a general principle" (Harpending, Rogers, and Draper 1987:132). Although research on some animal societies has shown a positive association between such proximate factors as higher status and greater fertility, human males usually do not follow expected predictions (for detailed discussions, see Hill 1984; Gray 1985). More seriously, the adaptive value of male "cultural success" when measured in terms of power, prestige, and wealth in industrial societies actually manifests a negative relationship: higher-status males have reduced fertility rates over lower-status males. In regarding this serious problem, two recent articles with very different solutions take up the challenge of why genic fitness is not maximized (or at least not fully operative) in human societies. Daniel Perusse (1993) in his "Cultural and Reproductive Success in Industrial Societies" concludes that although an inverse relationship undoubtedly exists between career success and reproductive success, his research did find a strong positive association between a male's social rank and his frequency of sexual intercourse. In short, Perusse maintains that industrialized males are still striving to optimalize fertility, thereby behaving no differently than in the past. But in modern societies, a male's "potential fertility" is blocked by two unnatural fecundity disrupters: a monogamous mating pattern and artificial birth control which, he says, has derailed the adaptive system. In comparison, Timothy Crippen (1994, this volume) says it is "patently false" to assume that all social behavior is adaptive. For Crippen, fitness is still the ultimate to evolutionary goal, but organisms should be seen as only tending behave as though they were optimizing fitness. Equally important, in humans the motivation to optimize is now often subverted by proximate factors (whether genetic, environmental, or sociocultural) which once enhanced fitness but are now likely to result in maladaptive and even genuinely altruistic behaviors. In light of these obstacles, Crippen calls for an "amended maximization principle" along the lines suggested by Lopreato (1984,1989), which would highlight the probabilistic nature of adaptation and shift attention away from a focus on adaptive behavior to a focus on the proximate, constraining factors that subvert the tendency to maximize fitness. Crippen avers that humans are equipped with "deep-seated behavioral predispositions" derived from humans' long tenure as food-foragers, which on average and in specific circumstances is still invoked to enhance fitness. But, with changing environmental circumstances, this motivation is now often conditioned, or supplanted, by proximate conditions such as needs for "creature comforts," which undermine genetic self-interest. This realization calls for a modified optimizing principle which would preserve the "covering law" of fitness and adaptation but, at the same time, promote and guide the investigation of the proximate conditions-material and emotional rewards, maximization of leisure,

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and so forth-that override and constrain the human motivation to maximize inclusive fitness. Thus, Crippen's selectionist approach is fundamentally different from traditional sociobiology which is mostly concerned with ultimate causation. Instead, while Crippen concurs that behavior is inherently and functionally designed to optimize fitness, he also seeks to investigate the proximate factors (whether environmental, sociocultural or genetic) that, under varying conditions, are likely to promote, constrain, or override the optimizing process. By advocating this dual emphasis, Crippen seeks to bridge the gap between sociobiology and sociological theory through the use of a common theoretical framework for both ultimate and proximate causation. Crippen is aware, we can only hope, that his approach is fraught with the same problems, revolving around issues of adaptation and causation, evident in traditional sociobiology, while his focus on the nature of proximate factors is likely to augment these methodological difficulties. Still, if Crippen's strategy does lead to some convincing results, it would no doubt stimulate lively debates in sociology. EVOLUTIONARY SOCIOLOGY Even the harshest of opponents credit sociobiology with highlighting what has too long been ignored in the social sciences: humans are animals who evolved just like every other organism, with selection favoring particular traits over others. Yet, despite this emphasis, sociobiology has won few converts in sociology, in part because of the field's biological reductionism but also because of its cynical and decidedly unpleasant portrayal of humans as manipulative, fitness maximizers. In short, the discipline "has a distinct penchant for bad news and the 'ugly' side of human behavior" (Arens 1989:403, his emphasis), which recently led one sociobiology critic to question: "can they all really mean what they seem to be saying?" (Settle 1993:62). Notwithstanding this inclination, sociobiologists have provided a much-needed corrective for the extreme social determinism of the past, although, if truth be told, sociologists have always entertained notions of "human nature." They just viewed these as inconsequential for understanding human behavior. But with the nature/ nurture problem now actively being explored within the social sciences, it would benefit general sociology to at least begin to consider how biology might influence human behavior and organization. But the question remains: how to proceed. First, it is important to emphasize that despite Wilson's skepticism, quite a number of sociologists have already incorporated the Modem Synthesis into sociology but withoutthe sociobiological thesis (see, for example, Lenski, Lenski, and Nolan 1991; Grove 1987; Kemper 1990; Freese 1988, 1994 [this volume]; Baldwin and Baldwin 1981;Machalek 1992;Catton 1992; Tenhouten 1991; Sanderson 1992; see also Masters 1991). Second, in considering additional ways to foster an evolutionary sociology, we might start by taking into account the evolutionary history of primates, which includes over

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190 species, classified into 25%prosimians, 70%monkeys, and 5%apes and humans. In the middle 1930s, detailed and long-term field studies on primates were initiated. These studies have now provided great insights into primate behaviors and organization. In particular, they allow scholars to appreciate how labile primate behaviors and patterns of social arrangements are and how much environmental/ ecological conditions and phylogenetic heritage (i.e., retention of characters from taxa of origin) interact in complex ways to shape patterns of primate organization. In short, the nonhuman primates are much like those who study them. Also, just as general patterns and forms of social organization and behaviors can be depicted for humans, there are discernible social structural and behavioral forms among nonhuman primates. Thus, by placing humans within this more inclusive primate framework, we can then entertain questions about "human nature" that move well beyond cross-cultural research. Such an approach has a number of advantages: First, it is empirically driven, keeping us in touch with reality by using the archaeological and fossil records as this can best be documented. Second, this method would incorporate the accumulated data on human and nonhuman primates, which could then guide inferences about humans' most basic behavioral, interactional, and organizational propensities. Third, this approach would allow us to employ established sociological research methods and theory and avoid excessive reductionism by stressing that, at times, selection processes operate at other than the genic level. Fourth, this approach would embrace the Moder Synthesis, which is not merely neo-Darwinian but a broad-based evolutionary paradigm that subsumes both micro and macro levels of evolution. Undoubtedly, this strategy lacks the elegance and precision of sociobiology models, but what it lacks in refinement it makes up in being congruent with humans' long-term evolutionary legacy. In recent years, this approach has been used to address such issues as hominoid sociality (Maryanski and Turner 1992; Maryanski 1992). Essentially, humans and apes are descendants of a large adaptive radiation of hominoids that flourished during the Miocene epoch (about 20 million years ago), although contemporary hominoids [i.e., the chimpanzee (Pan), gorilla (Gorilla),Orangutan (Pongo), gibbon (Hylobates), and human (Homo)] make up only a handful of primate species. Using a cladistic/network analysis for the organizational tendencies of the Last Common Ancestor (LCA) to humans and present-day hominoids (Maryanski 1992; Maryanski and Turner 1992),a striking conclusion can be drawn: like contemporary apes, the Last Common Ancestor of apes and humans evidenced a fluid organizational structure, consisting of a low level of sociality and a lack of intergenerational group continuity over time. The reasons for this structure are a combination of several forces: (1) female (and also male) dispersal from the natal unit at puberty, which is the opposite trend from monkeys, where male-biased dispersal leaves females to form intergenerational matrilines; (2) shifting mating patterns that make it impossible to know paternity (with the gibbon being the exception); and (3) relatively low social bonding among most adults. These features are associated with the dramatic

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decline in species of apes in the middle Miocene epoch, during which species of Old World monkeys suddenly proliferated and, according to the fossil record, moved into the former ape niches. One explanation for this replacement is that Cercopithecoid monkeys developed a competitive, dietary edge over the hominoids. In turn, this forced apes to evolve a novel skeletal structure with forelimb dominant locomotion in order to feed in borderline ecological zones, such as the terminal branches of fruit trees where quadrupedal monkey could not range (see Andrews 1981; Temerin and Cant 1983). Whatever the explanation, the fossil record confirms that ape niches during the Miocene were usurped by monkeys and that, during this replacement phase, apes were undergoing anatomical modification for forelimb dominant locomotion and other novel skeletal features that characterize both apes and humans today (Conroy, 1990;Andrews, 1981). But, in addition to skeletal modifications, movement into a new niche must have entailed organizational changes. Since terminal branch feeding or any marginal habitat is, by definition, one with thinly distributed resources, selection pressures would surely have worked against large, intergenerational groupings or stable cliques through time. Instead, if the social network reconstruction of the LCA population is correct (see Maryanski 1992; Maryanski and Turner 1992), pressures in the new adaptive zone selected for: (1) a dispersal of females from mother at puberty (a rare phenomenon among mammals in general and primates in particular), thereby preventing the continuance of the mother-daughter relationship known to be the primary building block for prosimians and monkey networks and for group continuity over time; and (2) behavioral tendencies for relatively low sociality, high individualism, and high autonomy, that created, at the hominoid organizational level loose and fluid social networks (which a comparative network analysis of the underlying structural relations of contemporary apes clearly makes evident). But at some point in time, descendant ape populations branched off from the LCA population and moved into forest zones, where selection seemingly favored the building of network ties, a trend evidenced by the novel organizational structures of the extant apes, whose network ties reflect efforts to work around the phylogenetic stumbling block of female dispersal at puberty. Moreover, the overall lack of density in ape organization seemingly stems from the hominoid general propensity for loose, fluid, and weak ties-a striking contrast to the tightknit and high density female cliques that characterize most monkey species. And what of hominids (i.e., humans or those early ancestors near to, or on, the line to humans)? While space forbids a discussion here (see Maryanski and Turner 1992; Maryanski 1993), some conclusions can be summarized. First, to the extent that we are willing to consider inferences from our closest ape relatives-where, for example in chimpanzees, our DNA differs from theirs by 1.1% [which "is far smaller than that between species within a genus of mice, frogs or flies" (King and Wilson 1978; see also Goodman et al. 1990; Sibley et al. 1990)1-hominoid social structure has an inherent tendency to be built around a predominance of weak ties over strong ties, which facilitates fluid, individual movement around a shared

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ranging area. A richness of weak ties over strong ties, as theoretically elaborated by Blau (1974), Friedkin (1980), and, in particular, Granovetter (1973), is also seen to provide hominoids with a degree of integration at the macro-population level in contrast to monkey populations where a richness of strong ties over weak ties is seen to provide integration at the micro-group level of organization (Maryanski 1987). Third, a predominance of weak ties among hominoids also suggests that, among humans, there might not be a biological need for "great intimacy" despite what many social philosophers have argued. If there is such a need for primary relations (aside from mother and offspring), it must have been selected for when early proto-hominids moved from the forest to open ranging savanna areas and confronted greatly increased predation pressures, where more intimate bonds might promote survival and reproductive success. Or, alternatively, it might have been selected for at the sociocultural level rather than at the genetic level. This conclusion might help us to understand the constant tension and vacillation in humans over constraint versus freedom, individualism versus collectivism, freedom of choice versus the collective good, and so on. At the biological level, humans may be far more individualistic than the "collectivist" biases of sociology would acknowledge. Also, the constant tension between the individual and the collective could be the result of humans' hominoid legacy of loose weak ties and a relatively low level of sociality. Finally, some of the conclusions outlined here on hominoid sociality might have broader implications for uncovering the origins of some core human social institutions, especially the economy with its economic division of labor between males and females; kinship with its traditional emphasis on patrilocal residence, marriage, and female-biased dispersal after puberty; and religion with its integrative Durkheimian rituals. While these considerations are for now highly speculative, they can provide some rich insights into the dynamics of human organization, if only by questioning implicit presumptions about "human nature" and by allowing for the possibility of discovery. Most significantly, they provide an alternative to "hard core," reductionist sociobiology. REFERENCES Alexander,Richard.1971."TheSearchfor an Evolutionary Philosophyof Man."Proceedings 8: Australia] 99-120. [Melbourne, of theRoyal Society Victoria of Andrews.P. 1981."SpeciesDiversityand Diet in Monkeysand Apes Duringthe Miocene." edited by C.B.Stringer.London:Taylorand of Pp. 26-61 in Aspects HumanEvolution, Francis. Evolution 401-407. 18: Journal Human Arens,W. 1989."Essaysin HumanSociobiology." of BasicConceptsand Limitations." Journal Social Archer, John.1991."HumanSociobiology: of Issues 11-26. 47: New Elsevier. Baldwin,John,and JaniceBaldwin.1981.Beyond Sociobiology. York: Barlow, George. 1991. "Nature-Nurtureand the Debates Surrounding Ethology and American 31: Sociobiology." Zoologist 286-296. D.P.1982.Sociobiology Behavior. York: and New Elsevier. Barash, of American Review 615-635. 39: Sociological Blau,P. 1974."Parameters SocialStructure."

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