Evolutionary Social Theory

Evolutionary Social Theory The Current State of Knowledge Joseph Carroll University of Missouri, St. Louis

abstract: The evolution of human sociality is a field in ferment, with writers s truggling to isolate elementary causal forces and organize them systematically. The elements of a usable model for evolved human sociality have become available only within the past few years. Those elements are scattered throughout the books here under review and a small set of articles. None of the books or articles fully exemplifies the whole model. After laying out the model, I use it to evaluate the books, describing how each contributes to it, and measuring each against it. The central idea in a usable model of human sociality is that the identity of the social group is integral to individual identity. In addition to that one central idea, a minimum of seven concepts is necessary to construct a model of sociality that includes the complex forms of organization in post-agricultural societies: (1) dominance, (2) egalitarianism or reverse dominance, (3) leadership, (4) internalized norms, (5) strong reciprocity or third-party enforcement of norms, (6) legal institutions, and (7) legitimacy in the exercise of power. These seven concepts can be reduced to four components: power, values, individuals, and groups. This model of evolved human sociality moves beyond the inconclusive debate between proponents of inclusive fitness and proponents of group selection. It also offers a distinct alternative to the identity politics that currently pervade literary and cultural study.

B ooks Under R eview Boehm, Christopher. Moral Origins: The Evolution of Virtue, Altruism, and Shame. New York: Basic Books, 2012. Print. Bowles, Samuel, and Herbert Gintis. A Cooperative Species: Human Reciprocity and Its Evolution. Princeton, NJ: Princeton UP, 2011. Print. Fukuyama, Francis. The Origins of Political Order: From Prehuman Times to the French Revolution. New York: Farrar, Straus and Giroux, 2011. Print. ———. Political Order and Political Decay: From the Industrial Revolution to the Globalization of Democracy. 1st ed. New York: Farrar, Straus and Giroux, 2014. Print. Haidt, Jonathan. The Righteous Mind: Why Good People Are Divided by Politics and Religion. New York: Pantheon Books, 2012. Print. Style, Vol. 49, No. 4, 2015. Copyright © 2015 The Pennsylvania State University, University Park, PA

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Nowak, Martin A., and Roger Highfield. Supercooperators: Altruism, Evolution, and Why We Need Each Other to Succeed. New York: Free P, 2011. Print. Pinker, Steven. The Better Angels of Our Nature: Why Violence Has Declined. New York: Viking, 2011. Print. Wilson, Edward O. The Social Conquest of Earth. New York: Liveright, 2012. Print.

A U SAB LE M ODE L OF EVO LVED H U MAN SOCIA LITY “Biocultural theory” is an integrative research program designed to investigate the causal interactions between biological adaptations and cultural constructions. The central premise of biocultural theory is that human behavior is produced by interactions between human nature and culture. “Human nature” in this usage designates a species-typical array of evolved, genetically transmitted features of anatomy, physiology, and neurology. “Culture” designates a collective, transmissible body of shared skills, practices, beliefs, values, and imaginative experiences (Baumeister; Carroll, “Truth”; Hill; Richerson and Boyd; Sterelny; Tomasello et al.). From the biocultural perspective, cultural processes are rooted in the b iological necessities of the human life cycle: specifically human forms of birth, growth, survival, mating, parenting, and life in a social group ( Muehlenbein and Flinn). Conversely, from the biocultural perspective, human biological processes are constrained, organized, and developed by culture, which includes technology, culturally specific socioeconomic and political structures, religious and ideological beliefs, and artistic practices such as music, dance, painting, and storytelling. Because culture is social, biocultural theory must include a good basic model of evolved human sociality. The elements of such a model have become available only within the past few years. Those elements are scattered throughout the books here under review and a small set of recent articles. None of the books or articles fully exemplifies the whole model. After laying out the model, I shall use it to evaluate the books, describing how each contributes to it, and measuring each against it. Until very recently, most discussions of evolved human sociality turned endlessly on an inconclusive debate between proponents of “inclusive fitness” theory and proponents of “group selection” (Alexander; Boehm, Moral Origins; Hamilton, “Rule,” “Aptitudes”; Nesse; Pinker, “Allure”; Sober and Wilson; Trivers; D. S. Wilson, “Critique”; Wilson and Wilson;


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E. O. Wilson, Social Conquest). Advocates of “inclusive fitness” reduced social motives to nepotism, direct reciprocation (mutual back-scratching), and indirect reciprocation (giving credit to people with a reputation for reciprocating) (Boehm, “Bullies,” Moral Origins; Nowak). Proponents of group selection felt rightly that these three sources could not adequately account for human prosociality, but they usually acceded to a misleading formula: selection favors selfishness within groups and favors altruism only in conflict between groups. As David Sloan Wilson and Edward O. Wilson put it, “in virtually all cases, traits labeled cooperative and altruistic are selectively disadvantageous within the groups and require between-group selection to evolve” (335). The nonprogressive character of the debate between the proponents of inclusive fitness and the proponents of group selection can be attributed in good part to the false alternatives offered in the formula repeated by Wilson and Wilson. That formula fails to register that individuals can choose to cooperate with cooperators and punish defectors—social cheats and bullies, “free riders” who take social benefits and do not contribute in a compensatory way to the maintenance of the group. The likelihood that cooperators will selectively interact with other cooperators is given different names by d ifferent theorists. Chudek and Henrich call it “phenotypic assortment” (19). Christopher Boehm calls it “social selection” (Moral Origins). Martin Nowak calls it “network reciprocity” and identifies it as one of five main sources of cooperation, the other four being kinship, direct reciprocity, indirect reciprocity, and group selection (Nowak; Nowak and Highfield; Nowak, Tarnita, and Antal). “Altruism,” strictly defined, cannot evolve. Genes that sacrifice themselves for the good of unrelated genes are a biological impossibility. Cooperation, however, can evolve, and has evolved many times over. It is a central principle behind the evolution of all complex life (Nowak and Highfield xviii, 280). It is at work in every “major transition” in evolution: from single replicators to ensembles of genes, from nonnucleated to nucleated singlecell organisms, from single-cell organisms to multi-cellular organisms, and from isolated individual organisms to social organisms (Bowles and Gintis, Cooperative Species; Calcott and Sterelny; Haidt; Maynard Smith and Szathmaáry; Michod; Simpson). In all cases of major transition, the systemic logic of the larger body is encoded in the genome of its particular parts. Individuals cease to be individuals—“selfish genes” or selfish people—and


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become components within a more complex biological entity. The success of each component depends on the success of the more complex entity. Within an organism, interactions among genes are shaped by the functional structure of the organism. Each cell contains the genome of the whole organism. Protein coding genes are switched on and off through networks of regulatory genes that subordinate individual cells to the systemic needs of the organism. In parallel fashion, within social animals, interactions among the individual organisms are shaped by the functional structure of the social group. Herbert Gintis puts this idea into clear focus: “Just as the genome codes for the patterns of interactions of loci in the genome, so it codes for the characteristic patterns of interactions of loci in two or more carriers; i.e., the genome codes for the social structure of the organism it creates” (“Inclusive Fitness” 497; and see Gintis, “Clash” 990). Human societies are less integrated than cells in an organism but more integrated than aggregates such as schools of fish, flocks of birds, or herds of ruminants (Simpson). No integration is perfect. Even in biological organisms, cancerous cells defect from the social body. Human societies are much messier, less coherent, than biological organisms. The needs and impulses of individuals usually work in some tension with the demands of society. Individuals are enmeshed in families and kin groups that contain internal conflicts and that also often work in tension with the demands of the larger society. Individuals and kin groups coalesce into classes, castes, trades, ethnic groups, cities, and geographical regions that have to negotiate conflicting interests (Fukuyama, Origins and Decay). In complex post-agricultural societies, all social power structures constitute temporary equilibria among diverse forces pulling in many different directions simultaneously. Nonetheless, most humans do in fact feel that they are members of some larger social body or set of social bodies. The identity of a social group is deeply embedded within the identity of the individuals that compose it. Individuals are not simply in competitive conflict with other members of their group; the group itself is part of each individual in it. All tribes have myths of origin. Every ethnic group, religious sect, nation, political party, and ideological movement makes itself imaginatively distinct through symbols, historical narratives, monuments, songs, pictorial representations, ceremonies, and aesthetic styles (Brown; Dissanayake; Hill; Hill, Barton, and Hurtado). Individual humans share in the collective imagination of their social groups, and within those groups, they construct


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narratives about their own individual lives (McAdams, “Life Narratives,” “Personal Narratives,” Power, “The Psychology of Life Stories,” Redemptive Self, Stories We Live By; McAdams and Bowman; McAdams and Ochberg; de St. Aubin, McAdams, and Kim). They envision their lives extending into the future, and they look beyond their own individual lives to the continuing life of their descendants and their communities. In contrast to chimpanzees, humans do not live in communities populated solely by the conspecifics with whom they come directly into contact. Their communities include fabled ancestors, generations of the future, and every person—living or dead—who shares beliefs and values that subordinate individuals to some collective body. The central idea in a usable model of human sociality is that the identity of the social group is integral to individual identity. In addition to that one central idea, a minimum of seven concepts is necessary to construct a model of sociality that includes the complex forms of organization in post-agricultural societies: (a) dominance, (b) egalitarianism or reverse dominance, (c) leadership, (d) internalized norms, (e) strong reciprocity or third-party enforcement of norms, (f ) legal institutions, and (g) legitimacy in the exercise of power. Over evolutionary and historical time scales, hominid and human social organization has alternated between dominance and egalitarianism. Dominance is the exercise of controlling force on individual members of a social group. In chimpanzee societies, one adult male, or a small coalition of adult males, dominate all other chimpanzees in a band (Boehm, Hierarchy; de Waal, Chimpanzee Politics, Inner Ape; Wrangham and Peterson). All adult males dominate all adult females; all adults dominate all juveniles. Using violence or the threat of violence, dominant males gain first access to prime foods and also to fertile females. Egalitarianism, a term used interchangeably with “reverse dominance,” is a uniquely human form of social organization and is the form prevalent among hunter-gatherer bands. Reverse dominance means that groups of adult males enforce equality by collectively suppressing dominance behavior in individual males. The group collectively dominates the individual. Individual males who assert dominance by bullying or by taking more than their share of food face sanctions extending from shaming through banishment to execution (Boehm, “Bullies,” Hierarchy, Moral Origins). In the evolutionary history of prehuman and human social organization, alternations of dominance and reverse dominance have passed through four


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main stages: (a) chimpanzee-like dominance centered in individual males or small coalitions of males; (b) hunter-gatherer egalitarianism among adult males; (c) the resurgence of personal dominance as an organizing principle of post-agricultural societies; and (d) the resurgence of egalitarianism in liberal democracies. Boehm argues that simple chimpanzee-like dominance characterized the behavior of the last common ancestors of hominids and chimpanzees; that hunters and gatherers almost universally practice reverse dominance; that the hunting and gathering ecology lasted long enough for egalitarianism to be deeply embedded in evolved human social dispositions; and that in human social groups, dispositions for collectively repressing individual dominance have always been held in strong tension with dispositions for individual dominance. Agriculture makes cumulable and defensible resources readily a vailable and thus alters the egalitarian equilibrium of hunter-gatherer social dynamics. “Material wealth allows aspirants to positions of social dominance to control enough allies and resources to offset the capacity of subordinate individuals to disable and kill them” (Gintis and Van Schaik 34). The resurgence of dominance in post-agricultural societies manifests itself in the prevalence of chiefs in tribal societies and in oligarchies, monarchies, aristocracies, and dictatorships. Reverse dominance resurfaced in Greek city-states and in the Roman republic but became a worldwide phenomenon only during the past two centuries (Fukuyama, Origins and Decay; Roberts; Wrangham and Peterson). Boehm’s arguments have had a major influence on evolutionary social theory and have also influenced evolutionary literary studies (Bowles, “Warriors”; Bowles and Gintis, Cooperative Species and “Strong Reciprocity”; Carroll et al.; Gintis and Van Schaik; Haidt; Richerson and Henrich; van Vugt and Ronay D. S. Wilson, Evolution; Wilson and Wilson). The exercise of power in post-agricultural societies combines the dominance hierarchies of our primate heritage with the willing cooperation that characterizes hunter-gatherer societies. In an essay summarizing evolutionary research on leadership, van Vugt and Ronay define leadership “in terms of the coordination of the actions of two or more individuals to accomplish joint goals” (2). The stipulation “joint goals” distinguishes leadership from raw assertions of dominance designed only to benefit the dominant individual or his kin (13). Leaders achieve legitimacy by eliciting the willing cooperation of individuals and directing them toward common goals in accordance with community norms. Qualities that have been found to


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be universally valued in leaders help them fulfill those functions: “integrity, persistence, humility, competence, decisiveness, and vision” (6). In both hunter-gatherer groups and complex modern societies, cooperative effort depends on the human ability to internalize norms. Analyzing the mechanisms of social learning that contribute to the human ability to internalize norms, Chudek and Henrich single out “prestige and conformity biases” as particularly important (219). Prestige bias is a disposition for imitating the behavior of high-status individuals. Conformity bias is a disposition for imitating the behavior of a majority. Both learning biases contribute to phenotypic assortment, the likelihood that “regularly interacting individuals resemble one another” (219). Social learning enables cooperators to interact selectively with other cooperators while excluding or punishing people who do not cooperate. Extended over evolutionary time scales, cooperative social practices function as a selective force: “culturally transmitted social strategies involving reputation, punishment and signaling substantially increase phenotypic assortment and facilitate the spread of self-reinforcing cooperative norms, creating genetic selection for a prosocial psychology” (220). The ability to internalize norms manifests itself in strong reciprocity: the willingness of a person to incur costs in order to enforce collective values even when that person is not directly affected by the violation of norms (Bowles and Gintis, Cooperative Species and “Strong Reciprocity”; Buckholtz and Marois). Picture a scene in which a man sees another man abusing a child. The observer does not know either the child or the abuser. He nonetheless intervenes, at considerable personal risk, to stop the abuse. Because he has nothing personally to gain from the intervention, he is a “third-party” enforcer of norms. Because people take risks and incur costs to enforce norms, strong reciprocity gives evidence that individuals care about the systemic logic that sustains a society even when they are not directly harmed by infractions against that logic. Strong reciprocity confirms that the functional structure of a social system has been encoded in the motivational repertory of individual people within that system. Institutions for the enforcement of norms distinguish complex, hierarchically organized societies from loosely aggregated band-level societies. In hunter-gatherer bands, norms are enforced by a perpetual stream of gossip serving as a medium for social monitoring (Boehm, Hierarchy and Moral Origins). In larger, more complex societies, face-to-face interaction and the


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direct collective enforcement of norms are not possible. Societies have to develop institutions that enforce social norms. Buckholtz and Marois argue that the codification of “norms into laws, and the attendant establishment of state-administered systems of criminal justice that are charged with norm compliance” are “one of the most important developments in human culture” (657). Distinguishing states from tribes, Francis Fukuyama argues that states possess a centralized authority backed by “a monopoly of the legitimate means of coercion, in the form of an army and/or police” (Origins 80). Internalizing a norm signifies that a person regards socially enforced standards of behavior as just or legitimate. As Fukuyama puts it, “Legitimacy means that the people who make up the society recognize the fundamental justice of the system as a whole and are willing to abide by its rules” (Origins 42). Jonathan Haidt also distinguishes legitimate authority from pure coercion: “Human authority” is “not just raw power backed by the threat of force. Human authorities take on responsibility for maintaining order and justice” (143). Leaders and members of legal institutions achieve legitimacy by exercising power in accordance with the shared values of the social group (Fukuyama, Origins and Decay; Gintis and Van Schaik; Haidt; van Vugt and Ronay). All societies have a structure of power, and in all societies, that power is used for the enforcement of norms. Even egalitarian hunter-gatherer bands exercise power in collectively suppressing dominance among individual males. If people are to internalize norms, they must also internalize power structures. The seven concepts in this model of evolved human sociality can be reduced to four components: power, values, individuals, and groups (Carroll, “Evolved Human Sociality”). Dominance and reverse dominance signify different ways in which social power is exercised, either by dominant individuals or by groups who suppress dominance behavior in individuals. Leaders are individuals who organize and direct the power latent in groups. Internalized norms are shared values that influence individuals. In strong reciprocity, individuals use power to enforce shared values. Legal institutions are contrivances for delegating strong reciprocity to individuals and groups such as the police and the judiciary. Leaders and members of legal institutions achieve legitimacy by exercising power in accordance with the shared values of the social group. Identifying four specific components as common denominators in all human social interaction provides a framework for a rational evolutionary comparison


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of societies extending from hunter-gatherer bands through modern postindustrial economies. CONTRIB UTIONS TO CU RRENT KNOWL EDGE FROM TH E WORKS U NDER REVIEW The evolution of human sociality is a field in ferment, with writers struggling to isolate elementary causal forces and organize them systematically. The books here under review are far removed from the settled status of textbook knowledge. In the history of science, ideas that will eventually become parts of settled science are at first often confusedly intermingled with other ideas that will not stand up to the test of time. Potentially permanent contributions to knowledge jostle along beside failed efforts to extend valid ideas to adjacent fields, mid-level analytic categories that are mistaken for conceptual primitives, incompatible basic concepts used by the same writer, and unwitting projections of ideological bias that limit the explanatory potential of good ideas. And yet it is from such ferments that new paradigms eventually are formed (Dolnick; Hallam; Mayr; Young). Assimilating Group Identity to Individual Identity: Nowak, Boehm, and Bowles and Gintis

Many theorists are struggling toward an explanation for the evolution of human prosociality, but a good deal of confusion persists in identifying the nature of the human social group. Some theorists who acknowledge prosocial dispositions in human nature attribute them to group selection or to some form of “multilevel selection,” selection that operates on both individuals and groups. In group selection, individuals are sometimes envisioned as semiautonomous entities in a collective loosely identified as a group. That kind of thinking fails to register that for humans the identity of the group is encoded in the individual. Except for sociopaths, humans are animated both by selfish motives and by motives aimed at sustaining the integrity of the social group. On the surface, multilevel selection seems to register that tension, but proponents of multilevel selection often wrongly suppose that all selection within the group selects for selfishness and that only selection between groups can promote prosociality. They thus overlook the idea of “network reciprocity” formulated by Nowak and the more specifically human idea of “social selection” formulated by Boehm.


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Nowak’s concept “network reciprocity” offers an alternative to kin selection, reciprocity, and also group selection (Nowak; Nowak and Highfield). But Nowak’s mathematical formulas remain abstract, fairly far removed from the psychology of human behavior, from the basic organizing structures of human life history, and from the socioeconomic and political institutions in which modern human behavior is enmeshed. Boehm has a clear understanding of the ecological conditions that shape social relations for hunter-gatherers, and he bears down effectively on the social psychology of internalizing norms. He does not, though, fully envision the way group identity is encoded in individual identity. His conceptions of cultural imagination include only moral norms for cooperation and sharing. Religion, ideology, and the arts play no part in his exposition. Moreover, his scope is limited to hunter-gatherers. He develops no theory of legal or governmental institutions like those necessitated by complex, post-agricultural economies. Similar limitations appear in the work of Bowles and Gintis. Boehm surveys the history of thinking about the evolution of human sociality, beginning with Darwin. In a famous passage, Darwin had argued that tribes containing men willing to sacrifice themselves for the good of the tribe would prevail over other tribes, but Darwin was puzzled about how those self-sacrificing genes could arise in a group to begin with, since self-sacrificing men would be at a selective disadvantage within the group (Darwin, Descent 1: 162–65). Boehm’s theory is designed to solve that puzzle. In truth, Boehm’s solution was pointed to by Darwin as his own speculative solution to the puzzle; Darwin suggests that selection for reputation would entice men to engage in heroic behavior. Boehm argues that over evolutionary time scales the coercive enforcement of group norms among huntergatherers acted as a selective force, giving an advantage to people who most readily internalized group norms, subordinating individual impulses of greed or dominance to cooperative interaction. That argument identifies a plausible evolutionary mechanism for the internalization of norms, but Boehm does not give much attention to the inter-group conflict that was the chief context of the problem Darwin had formulated. Boehm is considering only how cooperation and sharing could arise in a group considered apart from other groups. In his previous book on dominance and egalitarianism, Hierarchy in the Forest, Boehm had appealed to group selection, but in Moral Origins he discounts the significance of group selection and distinguishes it clearly from “social selection.” Bowles and Gintis put much more emphasis


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than Boehm on conflict between groups. They argue that warfare among ancient humans was sufficiently chronic and deadly to make group selection a major active force in human evolution (also see Bowles, “Did Warfare,” “Group Competition,” “Warriors”). Several theorists have registered that the theory of major transitions in evolution is the right conceptual context for the problem of incorporating the identity of the group into the genome of the individual. Nowak, Bowles/ Gintis, and Haidt all reference major transitions. Once a major transition is complete, the subordinate elements that have coalesced have ceased to be individuals and have become subordinate parts of a new individual (Calcott and Sterelny; Maynard Smith and Szathmaáry). Much of the confusion surrounding the nature of human sociality probably derives from the intermediate character of human sociality. Humans are both individuals and members of social groups. Human social groups are real entities, not just aggregates of discrete individuals, but the individuals are not as fully integrated as the cells that make up the individual organisms. Rather than focusing on this crucial conceptual problem, both Boehm and Haidt offer weighted assessments of the selfish and group-oriented parts of human nature. Boehm describes humans as intensely egoistic and nepotistic and moderately altruistic—that is, moderately given to “extra-familial g enerosity” (Moral Origins 10). With mock precision, Haidt affirms that humans are 90 percent selfish and 10 percent groupish (220). Ideas on the internalization of norms are now widely disseminated. Versions of Boehm’s theory appear in the books by Haidt and by Bowles and Gintis. Chudek and Henrich and Fukuyama do not reference Boehm but formulate similar ideas. The mechanisms for social selection proposed by Darwin, Boehm, and others will, I think, become a permanent part of the paradigm that will eventually form in this field. However, internalizing norms for cooperation and sharing is not quite the same thing as internalizing the identity of the social group as a whole. Norms specify patterns of behavior, and following such patterns can be mechanically triggered by stimuli in the same way that an environmental cue triggers instinctive behavior in an animal. To move from mechanically following rules to internalizing the identity of the group, individual people have to have an imaginative sense of their own identity and the identity of the group. They have to have an idea, a mental image, not just a mechanical reflex, but that idea must also be so deeply embedded in their motivational systems that it can, at


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least on occasion, over-rule the most powerful instincts, such as the instinct for survival. A soldier who throws himself on a grenade has internalized a sense of the group that transcends his own individual identity. Less dramatically, any effort that sacrifices immediate personal gain for the benefit of a group could give evidence that the individual has internalized the identity of the group within his or her own personal identity. Among the theorists here under review, Wilson, Haidt, and Fukuyama are most alert to the way humans use ideological narratives to evoke their sense of group identity. The Grand Old Manner: Wilson’s Social Conquest

Edward O. Wilson has done bold and innovative work in many fields over a long career. His discovery that pheromones serve as media of communications has created whole research programs for zoological specialists. His work on island biogeography set a new standard for scientific study in evolutionary ecology, and he has for many decades been one of the world’s most prominent and effective advocates for biodiversity and ecological preservation. In company with Charles Lumsden, he pioneered the field of gene-culture coevolution, a line of thought central to unifying the humanities and the sciences. He is one of the world’s foremost authorities in his own particular area of scientific specialization, the ants, and he has been one of the most successful authors of biological works aimed at generally educated readers. Consilience: The Unity of Knowledge, published in 1998, has inspired a generation of scholars and scientists who believe, along with Wilson, that nature forms a unitary causal order and that all knowledge can and should form a seamless web leading from physics and chemistry through biology to the social sciences and the humanities. Reviewing Consilience shortly after it was published, I described Wilson as a “cultural hero” (“Wilson’s Consilience” 396). Because of all his previous work, nothing could take that status away from Wilson, but the book currently under review, The Social Conquest of Earth, will not, I think, contribute to his reputation. I judge the book as, on the whole, a failure. It seems designed as both a revolutionary manifesto and a triumphal magnum opus—both a declaration of the new Wilsonian orthodoxy in theoretical biology and a magisterial summary of all his main themes about evolved human sociality. It is aimed at both a generally educated audience and at the audience of theoretical biologists preoccupied with the debate over inclusive fitness and group selection. The theoretical discussion deteriorates into confused and contradictory assertion.


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The popular-science presentation on evolved human sociality says little that is striking for novelty or depth of insight, and it lacks the evocative power of great essayistic prose. Wilson has never been a great stylist. His virtues have been virtues of substance, and the chief virtue of his style has been simple, functional clarity. In this book, Wilson has assumed the manner of the Grand Style without the substance to support it. The manner is merely self-important—combative, self-aggrandizing, and complacent. Wilson’s summary of human evolution might have some value for readers relatively unfamiliar with the subject. He usefully takes a zoological bird’seye view of humans, delineating the ecological conditions and morphological and behavioral characteristics that made human evolution possible (45–50). Humans are a land-dwelling species with a large body, grasping hands, spatulate fingers, and an upright gait. They have a diet that includes a large proportion of meat, and they occupy settled campsites centered on the controlled use of fire. They live in highly organized social groups that display an efficient division of labor. Despite his emphasis on meat-eating, Wilson offers a purely social account of the development of human intelligence (227), thus overlooking one of the most important lines of research in the evolutionary social sciences: “human life history theory,” which explains how the specifically human adaptations for provisioning, reproduction, and social organization have co-evolved in systemic interdependence with one another (Kaplan, Gurven, and Winking; Muehlenbein and Flinn). In attempting to delineate the higher reaches of imaginative life in art, literature, and philosophy, Wilson seldom rises above platitude. It is wholesome to hear, again, that the mind includes powers of imaginative synthesis and that the arts have adaptive utility. It is less valuable to hear, again, that literature consists essentially of metaphor and feeling (ch. 26). Common observation informs us that people internalize the identity of social groups, but providing a theoretical rationale for the internalization of group identity has proved difficult. Aggressively affirming a strict opposition between inclusive fitness theory and group selection, Wilson reiterates the idea that selection within groups can only favor selfishness and that cooperative behavior can arise only through group selection. He thus fails to register the significance of Nowak’s “network reciprocity” and Boehm’s “social selection.” While arguing for group selection, Wilson also makes a strained and equivocal effort to identify humans, along with the social insects and a few other species, as “eusocial.” Eusocial species are “superorganisms.” They have


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morphologically distinct reproductive castes. In the theory of eusociality Wilson developed to explain the organization of insect colonies, the whole hive or colony is essentially a single reproductive organism centered in the activity of the queen. An alternative way of saying that is that the nonreproductive workers are all robotic extensions of the queen (E. O. Wilson, “Giant Leap” and Social Conquest; Wilson and Hölldobler). The description of humans as “eusocial” can make sense only if “eusocial” is redefined to mean “very social” or “ultra-social.” Humans do not have morphologically distinct reproductive castes. Wilson acknowledges this, but once it is acknowledged, the only reasonable thing to do is to stop trying to use the term “eusocial” to describe human sociality. No other writer here reviewed uses that term for this purpose. The confusions in Wilson’s Social Conquest have a backstory in a 2007 article Wilson coauthored with David Sloan Wilson and in a 2010 article Wilson coauthored with Nowak and with Corina Tarnita (Nowak, Tarnita, and Wilson; Wilson and Wilson). In the 2007 paper, Wilson and Wilson had criticized inclusive fitness theory and had proposed multilevel selection, including group selection, as an alternative. In their 2010 paper, Nowak, Tarnita, and Wilson had also criticized inclusive fitness theory but on different grounds and with a different mathematical alternative. Nowak, Tarnita, and Wilson had argued that inclusive fitness theory requires strictly additive effects and pairwise interactions; it eliminates synergistic effects and thus models relations that seldom appertain in the actual world—an argument repeated in SuperCooperators. As an alternative to inclusive fitness, they had offered a game-theoretic framework that left room for favoring kin and also for group selection as specific sources of cooperation. In Social Conquest, Wilson makes rather a hash of all this. He confuses Nowak’s alternative to inclusive fitness with the kind of multilevel selection theory he and David Sloan Wilson had advocated in 2007. Evidently associating “inclusive fitness” (the general idea) with kin selection (the more particular application of the general idea), Wilson sometimes says that kin selection exists (76, 109), sometimes that it does not (53, 106), and sometimes that it might or might not exist but that if it does exist it does not count for much (174, 181). Advocates of multilevel selection often argue that inclusive fitness and multilevel selection are mathematically interchangeable ways of identifying “relatedness” and that relatedness can be phenotypic and behavioral, not exclusively genetic (Hamilton, “Aptitudes”; Marshall; Okasha; West, Griffin, and Gardner; D. S. Wilson, “Social Semantics”; Wilson and Wilson). Wilson


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and Wilson had said that inclusive fitness theory and group selection theory “are largely intertranslatable” and that they both “correctly predict what evolves in the total population” (336). In Social Conquest, the elder Wilson appeals to his work with Nowak and Tarnita to discredit inclusive fitness theory, reverts to a simple opposition between inclusive fitness theory and group selection, champions group selection, and declares that the difficulties with inclusive fitness theory “render incorrect the oft-repeated claim that group selection is the same as kin selection expressed through inclusive fitness” (171). Some general readers will no doubt be vaguely impressed by Wilson’s authoritative manner and a grand scope, especially if they feel that the main general substantive claims are ideologically unobjectionable. Many readers who are politically liberal and who are also sympathetic to biology will presumably assent readily enough to claims that people are divided between selfish and altruistic impulses, that humans have an ethical and aesthetic obligation to respect biological diversity, and that there are no significant variations among races (a claim differing from the claims made in Wilson’s 1978 book On Human Nature). For readers not too troubled by the opacities and contradictions in Wilson’s theoretical formulations, having yet another general review of human evolution might offer some satisfaction. For readers geared toward substantive developments in the evolutionary social sciences and evolutionary humanities, to which Wilson has contributed so much, this book is likely to be disappointing. Fukuyama on Political Development

Fukuyama’s two-volume set aims at giving a comprehensive evolutionary and historical overview of sociopolitical organization from prehuman times to the present. In his first volume, The Origins of Political Order, Fukuyama sketches out social organization in hunter-gatherer bands and in tribes, then describes in detail the history of political organization in China, India, and other societies from the beginning of state organization up to the period just before the French Revolution. The second volume, Political Order and Political Decay, continues the story from the French Revolution to the present. Fukuyama attempts to ground his historical account in an evolutionary view of human nature. Evidently believing that old-fashioned inclusive fitness theory is settled science, he confidently announces that reciprocation and favoring kin are the biological sources of all social behavior (Origins 30,


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439, Decay 8–9). He seems unaware that inclusive-fitness theory has ever been questioned, but he appears to see no conflict between this theory and ideas about the internalization of group norms, legitimacy in the exercise of power, and narratives of group identity—all of which play a major role in his books. He makes no reference to group selection or to major transitions theory. He nowhere explicitly formulates the idea that individual identity incorporates the logic of the larger social body. Since that idea is implicit in the internalization of group norms, legitimate authority, and narratives of group identity, his analytic descriptions of particular societies are out of sync with his basic theoretical formulations. A similar failure in deep theoretical coherence mars Fukuyama’s effort to synthesize evolutionary social science with specifically political theory. He draws a sharp line between band-level societies, based on kinship and reciprocity, and state-level societies: “With the advent of the state, we exit out of kinship and into the realm of political development proper” (Origins 81). Fukuyama argues that nepotism and reciprocity are the default form of social organization to which people regress when political structures decay. In his understanding, then, specifically political structures are not grounded in the evolved social dispositions that regulate hunter-gatherer behavior. They arise only out of demographic, ecological, and economic conditions. Fukuyama thus partially reinstates the dualistic idea that culture operates independently of evolved and adapted structures in the human mind. He agrees with Aristotle “that human beings are political by nature” (Decay 540), but he also suggests that “higher-level institutions are in some sense quite unnatural” (Origins 44). Though he does not formulate an adequate theory about the biological and psychological basis of states, Fukuyama usefully distinguishes states from tribes and also identifies key features that enable states to emerge out of tribes: states differ from tribes in having a centralized source of authority backed by the legitimate means of coercion; the state is territorial rather than kin based; states are more stratified than tribes; and states are legitimated by more elaborate forms of religious belief presided over by priests (Origins 80–81). Main factors generating the emergence of states from tribes are population density, geographical obstacles that prevent people from simply running away from rulers, geographical space sufficient for organizational scale, and the threat of extinction from other populations.


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With respect to societies in the period before the rise of modern democracy, Fukuyama evaluates political development by counter-posing the power of the state against other social institutions such as organized classes, religious institutions, municipal and regional powers, and guilds. China, in Fukuyama’s telling, developed a precocious state before society had achieved sufficient internal structure to balance against it. India, in contrast, had a weak state and a highly structured class system intertwined with a complex religious system. More fortunate, politically, than China and India, England produced the golden mean: a balance between a strong state and a highly structured social order with widely distributed centers of power. With respect to societies in the period since the French Revolution, Fukuyama uses three criteria to evaluate the degree of political development in a society: (a) the strength of the state; (b) the rule of law, that is, the existence of laws to which even rulers must bow; and (c) degrees of democratic participation. To describe variations in the political development of modern bourgeois democracies, he isolates three causal factors that can appear in different chronological sequences and in different combinations: (a) economic development; (b) social mobilization or the formation of new socioeconomic groups that are conscious of their identity as groups with distinct interests; and (c) ideas/ideology, like those manifested in the American Declaration of Independence or the Communist Manifesto (Decay 40–51). Modern liberal bourgeois democracy is for Fukuyama the normative, teleological condition to which all development tends. Even though democratic participation has developed fully only in the past two hundred years and has never been universal, it fulfills, Fukuyama believes, a basic need in human nature (Decay 540–41). As with his intuitive feel for narratives of group identity, Fukuyama’s intuitive feel for the psychological satisfactions of democratic participation exceeds his capacity for causal explanation. Darwin describes The Origin of Species as “one long argument” (Origin 459), meaning that his vast array of facts from multiple domains are all brought to bear as evidence for a chain of tightly linked causal processes summarized in the phrase “descent with modification by means of natural selection.” Fukuyama’s books are not in this sense an argument. They are analytic surveys deploying an array of conceptual categories that have good descriptive utility but no systematic relationship to a core set of primary causal terms.


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Haidt’s Moral Foundations

Haidt explains that as a graduate student he accepted “the common liberal belief that hierarchy = power = exploitation = evil” (143). By studying with Alan Fiske and through Fiske gaining access to Richard Shweder’s categories for social psychology, Haidt expanded his vision to include conservative criteria for social cohesion. Shweder had identified three clusters of moral themes: autonomy (the individual self ), community (the self in its relations to others), and divinity (the self in its relation to divinity) (in Haidt, 99). On the basis of his own research, Haidt further subdivides this set of themes, originally producing five “foundations” of moral psychology, and then later adding a sixth (125–54, 169–73). Each “foundation” is joined by a slash with an opposite that signifies the violation of the positive criterion. Haidt argues that the first two foundations, Care/harm and Fairness/cheating, are the basis for liberal or left-wing political values. Care is the concern for the welfare of individuals, the prohibition against harming them. Fairness was first envisioned by Haidt as reciprocity and also equality but was later modified, under criticism from conservatives, to include equity or proportionality of reward to merit (169). The inverse of Fairness, cheating, is a criterion stigmatizing free riders. The other three foundations of the original five are designed to encompass Shweder’s ethics of “community” and “divinity.” Loyalty/betrayal and Authority/subversion are values oriented to the integrity of the social body; Sanctity/degradation is Haidt’s version of religious feeling. The sixth foundation, Liberty/oppression, was added during the rethink of fairness. Liberty/oppression is based on Boehm’s concepts of dominance and reverse dominance; it is essentially individual resistance against dominance. For each foundation, Haidt identifies an evolved, instinctive origin. Care originates in care of children; Fairness originates in reciprocity and also in resistance to dominance (egalitarianism); Loyalty derives from primate dispositions for forming coalitions; Authority derives from dispositions for negotiating dominance hierarchies; and Sanctity is ascribed to disgust reflexes against disease-bearing substances such as feces and rotting flesh. These six criteria for moral judgment serve well as descriptive categories for a range of moral feelings, but they have a certain ad hoc, provisional feel to them. They derive originally not from an evolutionary conception of the basic structures of the human life cycle but only from analytically organized impressions of responses to surveys. Adding a sixth foundation that overlaps with the Authority and Fairness foundation—all three involve dealing


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with dominance hierarchies, and equality is divided between Fairness and Liberty—suggests how far this system is from cutting moral psychology at its joints. Haidt says that there are at least six foundations but might be even more—he suggests honesty, ownership, self-control, and waste as possible candidates. Despite the open-ended suggestion of further possible candidates, he says he is confident that these six are the main ones (181, 347 n. 25, 350 n. 53). That confidence seems gratuitous. The proliferation of candidates for moral foundations has a parallel with the tendency in early evolutionary psychology to postulate open-ended lists of “modules” supposedly geared toward specific adaptive challenges—and indeed, Haidt himself explicitly links his concept of foundations with the concept of modularity. The weak theoretical structure of open-ended lists in early evolutionary psychology was effectively countered by human life history theory—the theory of a systemic logic in the reproductive cycle, with its specific phases, family roles, and social roles. A system of moral psychology better grounded than Haidt’s would rationalize its categories within human life history theory. A grounding in human life history would also overcome a blind spot in Haidt’s purely social version of human psychology. In its basic scheme of human psychology, human life history theory incorporates the need for acquiring food. That need involves developing skills, thus playing an evolutionary role in the extension of human childhood, and it entails, in ancestral populations, sex-specific divisions of labor (Kaplan, Gurven, and Winking). The systemic integration of foraging, hunting, bearing and tending children, and forming cooperative groups, some all-male, some all-female, and some male and female, have had lasting consequences in human anatomy, physiology, and psychology (Boehm, Moral Origins; Flinn, Geary, and Ward; Foley and Gamble; Geary and Flinn; Gintis and Van Schaik; Kaplan, Gurven, and Lancaster; Kaplan, Hooper, and Gurven; Muehlenbein and Flinn; Wrangham). When Haidt declares that “moral psychology” is “the key to understanding politics, religion, and our spectacular rise to planetary dominance” (315), he betrays a blind spot peculiar to people who have never been cold, hungry, or in physical danger. Like most academic psychologists, Haidt has presumably spent most of his life in modern bourgeois environments beautifully engineered for safety and comfort. For people whose own immediate experience gives them no sense of challenges from the physical environment, the only challenges that seem real can be those associated with


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“forging beneficial relationships within hierarchies” (144). Our actual human nature is broader, deeper, and tougher than that. Haidt’s own political ideology, self-professedly, is liberal. In other words, it is an ideology oriented to the fulfillment of individual needs so long as they do no harm to others. Though he acknowledges “Authority” as a principle, in describing it, he does not describe reverence toward a power structure that embodies the moral ethos of a community. Instead, he speaks of opportunistically negotiating hierarchies so as to gain “the protection of superiors and the allegiance of subordinates” (144). In developing the idea of group identity, he does not develop the idea that group identity is integral with individual identity. Instead, he appeals to the work of Émile Durkheim, a founder of sociology and a chief architect of the Standard Social Science Model (Degler; Fox; Pinker, Blank Slate; Tooby and Cosmides). As Haidt himself acknowledges, Durkheim is a proponent of the idea that social institutions and social group phenomena operate independently of evolved psychological dispositions lodged in individual minds. “As a sociologist, Durkheim focused on social facts—things that exist outside of any individual mind” (270). Haidt’s examples of such social facts include “families, laws” and “shared networks of meaning.” That kind of thinking contains an obvious fallacy. Families are constituted by more than one person and thus exist not only in one individual mind but in the minds of all the individuals in the family. There is a kind of sophistical sleight of hand at work in removing families, laws, and religions from experiences originating in individual minds. That sort of sophistry is endemic to Durkheim’s thinking. The split in Haidt’s sources—evolutionary psychology for the origins of moral foundations, Durkheim for corporate social identity—reappears in Haidt’s version of multilevel selection. He declares that “most of human nature was shaped by natural selection operating at the level of the individual. Most, but not all. We have a few group-related adaptations too” (219). This is a split-level version of human identity. Most of the time, Haidt suggests, we are egoistic isolates, unconcerned with the social body. But 10 percent of the time, when social cues trigger a switch, we become “hivish” creatures, like bees. That image of human nature—“We are 90 percent chimp and 10 percent bee” (220)—suggests a basic failure to overcome an original belief in the primacy of individuals as a unit of selection. In reality, we are never an asocial species. We are always a species in which social identity is integral to individual identity. Haidt has not developed an integrated view


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of human nature. He has simply shoved together two anomalous pieces: the egoistic isolate that makes up, supposedly, 90 percent of human motives and the superorganism that makes up the other 10 percent. Given this odd incoherence in his basic conception of human nature, Haidt is to be credited with achieving as much as he does achieve. He makes reference to Boehm, to major transitions theory, and to Dan P. McAdams’s theories about ideological narratives. He thus brings together major elements in a usable model of human nature. His own synthesis of these elements is fairly superficial, and his chief constructive contribution, the elaboration of Shweder’s categories, is both derivative and insufficiently grounded in human life history theory. Still, he provides the rudiments of a moral social psychology associated with evolutionary thinking, and in that respect he at least provisionally occupies a crucial position in which he has little competition. The closest available competition is the kind of five-factor personality psychology that forms the core of Dan P. McAdams’s research. Pinker’s Liberal Individualism

Fukuyama affirms old-fashioned inclusive fitness theory but then deploys a set of categories presupposing the reality of corporate identity. Pinker affirms old-fashioned inclusive fitness theory and actually means it (“False Allure”). He believes that self-interest, nepotism, and reciprocation exhaust the sources of human social behavior. In concord with that individualistic vision of human nature, Pinker identifies Thomas Hobbes as a chief inspiration for the political philosophy at work in The Better Angels of Our Nature: Why Violence Has Declined. For Hobbes, humans are inherently aggressive, selfish, and asocial. They are brought together into cooperative social activity only through the repressive, coercive power of the state, the Leviathan. In The Origins of Political Order, Fukuyama gives an astute critique of Hobbes’s false vision of human nature and pairs it with a critique of the inverse but equally false vision of human nature in Rousseau. Pinker simply takes Hobbes at face value. Most evolutionary psychologists would not make that mistake, but Pinker can at least be credited with a certain consistency. The Hobbesian vision is compatible with the cynical vision of human nature in Richard Dawkins’s The Selfish Gene and in Richard Alexander’s The Biology of Moral Systems (D. S. Wilson, “Critique”). In The Meaning of Human Existence, published after Social Conquest, Edward O. Wilson rightly criticizes that vision for taking the individual organism as the main unit


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of selection: “With inclusive fitness the unit of selection had passed subtly from the gene to the individual” (69). For all its theoretical limitations, Haidt’s taxonomy of modern left–right political ideology has a good deal of descriptive utility. It is for instance adequate for describing Pinker’s liberal individualism. Though affiliated with a cynically narrow understanding of human nature, Pinker’s own value system is wholly oriented to the Harm/care foundation. He loathes violence. He has a kindly disposition. War and cruelty are abhorrent to him. Like the typical liberal ideologues characterized by Haidt, Pinker is blind to the moral character of identification with a social group. He usually cannot even register the reality of group identity, but he can on occasion, locally, make use of it to explain the cruelty of groups who engage in religious or ethnic persecution (522–25). That is, groups, insofar as they exist at all, are agencies of evil. Pinker has one main scholarly purpose in Better Angels: to give evidence that violence has declined over historical time. For this purpose, he has amassed a huge collection of facts, and much of his evidence—so far as domestic and civil life go—is convincing. One might obviously object that violence has moved up the chain of social organization, shifting from individuals and small groups to nation states, ethnic enclaves, and ideological blocs. To counter that claim, Pinker must deprecate the significance of the two World Wars, the Holocaust, Stalin’s purges and the famine Stalin intentionally created in Ukraine, the Chinese Cultural Revolution, and the Cambodian killing fields. Through ingenious special pleading, Pinker contrives not to explain but to explain away all such events: “We will never really know how to explain the most calamitous events of the 20th century. The ideologies prepared the ground and attracted the men, the absence of democracy gave them the opportunity, but tens of millions of deaths ultimately depended on the decisions of just three individuals [Stalin, Hitler, and Mao]” (343). The solution is clear. We can never be sure of getting rid of all the bad apples—like Hitler, Stalin, and Mao—but we can resolutely turn away from all evil ideologies, that is, all ideologies that involve any concession to group identity. Accordingly, Pinker unequivocally affirms “the Enlightenment ideal that the ultimate value in the political realm is the individual human being” (258). Pinker shares with the Enlightenment not only an emphasis on the rights of the individual but also a specific vision of history. Histories written by Enlightenment thinkers such as Voltaire and Gibbon are unilinear and


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progressive. People used to be superstitious, stupid, and cruel, but they have gradually become rational, intelligent, and kind. There is something thin and shallow in this way of thinking. It can identify improved conditions of material life and improved standards of civil behavior, but it cannot grasp the inner logic that makes sense of previous cultural periods. Because it is blind to the elements of human nature that produced the brutalities of previous historical periods, it has no real explanatory grasp on the forces still at work in our own contemporary circumstances. Pinker is immensely clever and extraordinarily well informed. Nonetheless, of all the books here under review, Better Angels offers the least toward a usable model of evolved human sociality. Pinker’s understanding of theoretical biology is dogmatic, narrow, and shallow, and his handling of facts is subordinated to an ideology: a structure of values supported by false ideas about human nature and human social interaction. CONCLU SION Evolutionary studies in the humanities necessarily depend on biocultural theory—analytic accounts of the interaction among innate human dispositions and specific environmental conditions, including the built and imagined environments produced by crafts, engineering, technology, religions, ideologies, philosophy, the sciences, and the arts. Good biocultural theory, in turn, depends on adequate models of human nature. Since humans are a profoundly social species, adequate models of human nature depend on valid theories about evolved human sociality. The books and articles reviewed here represent the best of current thinking on evolved human sociality. Nothing has been decisively settled in the debates between proponents of inclusive fitness theory and proponents of group selection. No single theorist contains all the elements necessary for a usable basic model of human nature. But we now have enough elements, scattered across the field, to see beyond the cynical and narrow vision of human nature promulgated by the proponents of selfish gene theory. We know that humans internalize norms and have innate predispositions for engaging in cooperative behavior with others. The one most important idea in a usable model of human nature—the idea that individual identity contains the identity of the social group—has been suggested and implied much more often than it has been clearly formulated. Because it is true and important, and because it is clearly


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implied by the most advanced evolutionary social thinkers, it is virtually certain to become a leading idea in the further development of evolutionary social theory. For literary studies, the intersection of individual identity and social group identity can be applied to analyzing the interplay between narratives of the individual self and narratives of cultural group identity. Every author, every reader, and every character depicted in a poem, play, or story has a point of view, a personality, an individual identity. For each person or character, that identity is both idiosyncratic, individual, unique, and also stereotypical, recognizably compounded of the basic features of human nature interacting with the ideas, feelings, and values that are available in the culture in which a person or character lives. Each author who depicts a character is a unique individual participating in a shared cultural medium. In contriving imagined worlds, authors affirm their own complex identity themes—attitudes and beliefs, tastes, ideas, and values—and they seek to engage the responses of readers who also have world views. Creating and responding to literary depictions is thus an ultra-social form of activity. Literature provides a vast field of play for uniquely developed human capacities for “theory of mind,” that is, entering imaginatively into the experience of other people, real and fictional. To be adequate to its subject, literary theory must include a good model of evolved human sociality. Biocultural literary analysis offers a distinct alternative to the “identity politics” that currently pervade literary and cultural studies. For some three decades or more, a majority of literary scholars have adopted a Foucauldian perspective on social power relations. From the Foucauldian perspective, society is a system subserving impulses of domination—domination of one class by another class, or one gender, race, or ethnic group by another such group (Armstrong; Butler; Miller). Freudian forces of repression within the subconscious supposedly keep the processes of power hidden (Jameson). The function of criticism, for the Foucauldian, is to expose the subterranean machinations of social dominance. As Foucault puts it, “power is always exercised at the expense of the people” (211). Consequently, “the intellectual’s role” (207) is to engage in “a struggle against power, a struggle aimed at revealing and undermining power where it is most invisible and insidious” (208). These Foucauldian formulas give a stripped down and distorted image of social identity (Carroll, “Correcting”). Social relations include dominance, subjugation, and resistance, but they also include


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internalized norms, willing cooperation, and identification with a social body. Most literary authors intuitively recognize this more complete set of principles at work in social relations. Literary scholars need not be only end users of biocultural theory produced by evolutionary biologists and social scientists. Given the unsettled condition of evolutionary social theory at the present time, literary scholars who want to use that theory have no choice but to engage with it critically. But even in future times, when evolutionary social theory will have become more of a settled paradigm, literary scholars will still have a vital role to play. Literary scholars have training and aptitudes that enable them to give expert accounts of imaginative structures. Making judicious use of the information available in evolutionary social science should greatly enhance the explanatory capacity of their own research and also contribute to the collective effort, across the sciences, to understand human culture.

joseph carroll is Curators’ Professor in the English department at the University of Missouri—St. Louis. In addition to monographs on Matthew Arnold and Wallace Stevens, he is the author of Evolution and Literary Theory (1995), Literary Darwinism: Evolution, Human Nature, and Literature (2004), and Reading Human Nature: Literary Darwinism in Theory and Practice (2011); he is the lead author of Graphing Jane Austen: The Evolutionary Basis of Literary Meaning (2012). He has produced an edition of Darwin’s Origin of Species (2003) and is a co-editor of Evolution, Literature, and Film: A Reader (2010) and Darwin’s Bridge: Uniting the Sciences and the Humanities (forthcoming). WORKS CITED Alexander, Richard D. The Biology of Moral Systems. Foundations of Human Behavior. Hawthorne, NY: de Gruyter, 1987. Print. Armstrong, Nancy. Desire and Domestic Fiction: A Political History of the Novel. New York: Oxford UP, 1987. Print. Baumeister, Roy F. The Cultural Animal: Human Nature, Meaning, and Social Life. Oxford: Oxford UP, 2005. Print. Boehm, Christopher. “Bullies: Redefining the Human Free-Rider Problem.” Darwin’s Bridge: Uniting the Humanities and Sciences. Eds. Joseph Carroll, Daniel P. McAdams, and Edward O. Wilson. New York: Oxford UP, forthcoming. Print. ———. Hierarchy in the Forest: The Evolution of Egalitarian Behavior. Cambridge, MA: Harvard UP, 1999. Print.


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Bowles, Samuel. “Did Warfare among Ancestral Hunter-Gatherers Affect the Evolution of Human Social Behaviors?” Science 324.5932 (2009): 1293–98. Print. ———. “Group Competition, Reproductive Leveling, and the Evolution of Human Altruism.” Science 314.5805 (2006): 1569. Print. ———. “Warriors, Levelers, and the Role of Conflict in Human Social Evolution.” Science 336.6083 (2012): 876–79. Print. Bowles, Samuel, and Herbert Gintis. “The Evolution of Strong Reciprocity: Cooperation in Heterogeneous Populations.” Theoretical Population Biology 65.1 (2004): 17–28. Print. Brown, Donald E. Human Universals. Philadelphia: Temple UP, 1991. Print. Buckholtz, Joshua W., and Rene Marois. “The Roots of Modern Justice: Cognitive and Neural Foundations of Social Norms and Their Enforcement.” Nature Neuroscience 15.5 (2012): 655–61. Print. Butler, Judith. Gender Trouble: Feminism and the Subversion of Identity. New York: Routledge, 1990. Print. Calcott, Brett, and Kim Sterelny. The Major Transitions in Evolution Revisited. Cambridge, MA: MIT P, 2011. Print. Carroll, Joseph. “Correcting for the Corrections: A Darwinian Critique of a Foucauldian Novel.” Style 47.1 (2013): 87–118. Print. ———. “Evolved Human Sociality and Literature.” Evolutionary Analysis in the Social Sciences. Eds. Richard Malachek, Jonathan Turner, and Alexandra Maryanski. Boulder, CO: Paradigm, 2015. 572–608. Print. ———. “The Truth About Fiction: Biological Reality and Imaginary Lives.” Style 46.2 (2012): 129–60. Print. ———. “Wilson’s Consilience and Literary Study.” Philosophy and Literature 23.2 (1999): 393–413. Print. Carroll, Joseph, Jonathan Gottschall, John A. Johnson, and Daniel Kruger. Graphing Jane Austen: The Evolutionary Basis of Literary Meaning. New York: Palgrave Macmillan, 2012. Print. Chudek, Maciej, and Joseph Henrich. “Culture-Gene Coevolution, Norm-Psychology and the Emergence of Human Prosociality.” Trends in Cognitive Sciences 15.5 (2011): 218–26. Print. Darwin, Charles. The Descent of Man, and Selection in Relation to Sex. Eds. John Tyler Bonner and Robert M. May. Princeton, NJ: Princeton UP, 1981. Print. ———. On the Origin of Species by Means of Natural Selection. London: J. Murray, 1859. Print. Dawkins, Richard. The Selfish Gene. New York: Oxford UP, 1976. Print. Degler, Carl N. In Search of Human Nature: The Decline and Revival of Darwinism in American Social Thought. New York: Oxford UP, 1991. Print. de St. Aubin, Ed, Dan P. McAdams, and Tae-Chang Kim. The Generative Society: Caring for Future Generations. 1st ed. Washington, DC: American Psychological Association, 2003. Print. de Waal, Frans. Chimpanzee Politics: Power and Sex Among Apes. 25th Anniversary ed. Baltimore, MD: Johns Hopkins UP, 2007. Print.


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538 Style ———. Our Inner Ape: A Leading Primatologist Explains Why We Are Who We Are. New York: Riverhead Books, 2005. Print. Dissanayake, Ellen. Art and Intimacy: How the Arts Began. Seattle: U of Washington P, 2000. Print. Dolnick, Edward. The Clockwork Universe: Isaac Newton, the Royal Society, and the Birth of the Modern World. New York: Harper, 2011. Print. Flinn, Mark V., David C. Geary, and Carol V. Ward. “Ecological Dominance, Social Competition, and Coalitionary Arms Races: Why Humans Evolved Extraordinary Intelligence.” Evolution and Human Behavior 26.1 (2005): 10–46. Print. Foley, Robert, and Clive Gamble. “The Ecology of Social Transitions in Human Evolution.” Philosophical Transactions of the Royal Society B: Biological Sciences 364.1533 (2009): 3267–79. Print. Foucault, Michel. Language, Counter-Memory, Practice: Selected Essays and Interviews. Trans. Donald F. Bouchard and Sherry Simon. Ed. Donald F. Bouchard. Ithaca, NY: Cornell UP, 1977. Print. Fox, Robin. The Search for Society: Quest for a Biosocial Science and Morality. New Brunswick, NJ: Rutgers UP, 1989. Print. Geary, David C., and Mark V. Flinn. “Evolution of Human Parental Behavior and the Human Family.” Parenting: Science and Practice 1.1/2 (2001): 5–61. Print. Gintis, Herbert. “Clash of the Titans.” BioScience 62.11 (2012): 987–91. Print. ———. “Inclusive Fitness and the Sociobiology of the Genome.” Biology and Philosophy 29 (2014): 477–515. Print. Gintis, Herbert, and Carel P. Van Schaik. “Zoon Politicon: The Evolutionary Roots of Human Sociopolitical Systems.” Cultural Evolution. Eds. Peter J. Richerson and Morten H. Christiansen. Cambridge, MA: MIT Press, 2012. 25–44. Print. Hallam, Anthony. Great Geological Controversies. Oxford: Oxford UP, 1983. Print. Hamilton, W. D. “Hamilton’s Rule. The Genetical Evolution of Social Behavior, I and II.” Narrow Roads of Gene Land: The Collected Papers of W. D. Hamilton. Vol. 1. Oxford: W. H. Freeman, 1996. 11–82. Print. ———. “Innate Social Aptitudes of Man: An Approach from Evolutionary Genetics.” Biosocial Anthropology. Ed. Robin Fox. New York: Wiley, 1975. 135–55. Print. Hill, Kim. “Evolutionary Biology, Cognitive Adaptations, and Human Culture.” The Evolution of Mind: Fundamental Questions and Controversies. Eds. Steven W. Gangestad and Jeffry A. Simpson. New York: Guilford, 2007. 348–56. Print. Hill, Kim, Michael Barton, and A. Magdalena Hurtado. “The Emergence of Human Uniqueness: Characters Underlying Behavioral Modernity.” Evolutionary Anthropology: Issues, News, and Reviews 18.5 (2009): 187–200. Print. Jameson, Fredric. The Political Unconscious: Narrative as a Socially Symbolic Act. Ithaca, NY: Cornell UP, 1981. Print. Kaplan, Hillard S., Michael Gurven, and Jane B. Lancaster. “Brain Evolution and the Human Adaptive Complex: An Ecological and Social Theory.” The Evolution of Mind: Fundamental Questions and Controversies. Eds. Steven W. Gangestad and Jeffry A. Simpson. New York: Guilford, 2007. 269–79. Print.


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