12
An Evolutionary Theory of Commons Management

Peter J. Richerson, Robert Boyd, and Brian Paciotti

Common property and common-pool resources dilemmas are examples of the broader problem of cooperation, a problem that has long interested evolutionists. In both the Origin and Descent of Man, Darwin worried about how his theory might handle cases such as the social insects in which individuals sacrificed their chances to reproduce by aiding others. Darwin could see that such sacrifices ordinarily would not be favored by natural selection. He argued that honeybees and humans were similar: Among honeybees a sterile worker who sacrificed her own reproduction for the good of the hive would enjoy a vicarious reproductive success through her sibling reproductives. Humans, Darwin (1874:178-179) thought, competed tribe against tribe as well as individually, and the “social and moral faculties” evolved under the influence of group competition:

It must not be forgotten that although a high standard of morality gives but slight or no advantage to each individual man and his children over other men of the tribe, yet that an increase in the number of well-endowed men and an advancement in the standard of morality will certainly give an immense advantage to one tribe over another. A tribe including many members who, from possessing in a high degree the spirit of patriotism, fidelity, obedience, courage, and sympathy, were always ready to aid one another, and to sacrifice themselves for the common good, would be victorious over most other tribes; and this would be natural selection.

More than a century has passed since Darwin wrote, but the debate among evolutionary social scientists and biologists is still framed in similar terms—the conflict between individual and prosocial behavior guided by selection on individuals versus selection on groups. In the meantime, social scientists have devel-



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The Drama of the Commons 12 An Evolutionary Theory of Commons Management Peter J. Richerson, Robert Boyd, and Brian Paciotti Common property and common-pool resources dilemmas are examples of the broader problem of cooperation, a problem that has long interested evolutionists. In both the Origin and Descent of Man, Darwin worried about how his theory might handle cases such as the social insects in which individuals sacrificed their chances to reproduce by aiding others. Darwin could see that such sacrifices ordinarily would not be favored by natural selection. He argued that honeybees and humans were similar: Among honeybees a sterile worker who sacrificed her own reproduction for the good of the hive would enjoy a vicarious reproductive success through her sibling reproductives. Humans, Darwin (1874:178-179) thought, competed tribe against tribe as well as individually, and the “social and moral faculties” evolved under the influence of group competition: It must not be forgotten that although a high standard of morality gives but slight or no advantage to each individual man and his children over other men of the tribe, yet that an increase in the number of well-endowed men and an advancement in the standard of morality will certainly give an immense advantage to one tribe over another. A tribe including many members who, from possessing in a high degree the spirit of patriotism, fidelity, obedience, courage, and sympathy, were always ready to aid one another, and to sacrifice themselves for the common good, would be victorious over most other tribes; and this would be natural selection. More than a century has passed since Darwin wrote, but the debate among evolutionary social scientists and biologists is still framed in similar terms—the conflict between individual and prosocial behavior guided by selection on individuals versus selection on groups. In the meantime, social scientists have devel-

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The Drama of the Commons oped parallel theories of cooperation—rational choice theory takes an individualistic approach while functionalism analyzes the prosocial aspects of institutions. In this chapter we review the evolutionary theory relevant to the question of human cooperation and compare the results to other theoretical perspectives. Then we review some of our own work distilling a compound explanation that we believe gives a plausible account of human cooperation and selfishness. This account leans heavily on group selection on cultural variation but also includes lower level forces driven by both micro-prosocial and purely selfish motives. Next, we review the empirical literature in commons management. Although much work remains to be done on the problem, we conclude that the existing evidence is consistent with our account. Then, we use our hypothesis to derive lessons for applied research in institution building for commons management. On the one hand, the theory of cultural group selection suggests that humans have cooperative sentiments usually assumed to be absent in rational choice theories. On the other hand, the slow rate at which cooperative institutions evolve suggests that considerable friction will afflict our ability to grow up commons management institutions if they do not already exist and to readapt existing institutions to rapid technological and economic change. A better understanding of the way cooperative institutions arise in the long run promises better tools to foster their more rapid evolution when needed and to regulate their performance as necessary. THEORIES OF COOPERATION Our ideas about cooperation are drawn from many sources. Folk sources include diverse religious doctrines, norms and customs, and folk psychology. Anthropologists and historians document an immense diversity of human social organizations and most of these are accompanied by moral justifications, if often contested ones. Johnson and Earle (1987) provide a good introduction to the vast body of data collected by sociocultural anthropologists. The cross-cultural study of commons management is already a well-advanced field drawing on the disciplines of anthropology, political science, and economics (Agrawal, this volume:Chapter 2; Baland and Platteau, 1996; Bardhan and Dayton-Johnson, this volume:Chapter 3; Berkes, this volume:Chapter 9; McCay, this volume:Chapter 11; (Ostrom, 1998). Human Cooperation Is Extensive and Diverse Human cooperation has a number of features begging explanation: Humans are prone to cooperate, even with strangers. Thus many people cooperate in anonymous one-shot prisoners’ dilemma (PD) games (Marwell and Ames, 1981), and often vote altruistically (Sears and Funk, 1990). People begin

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The Drama of the Commons contributing substantially to public goods sectors in economic experiments (Falk et al., this volume:Chapter 5; Kopelman et al., this volume:Chapter 4; Ostrom, 1998). The experimental results accord with common experience. Most of us have traveled in foreign cities, even poor foreign cities filled with strange people for whom our possessions and spending money are worth a small fortune, and found risk of robbery and commercial chicanery to be small. Cooperation is contingent on many things. Not everyone cooperates. Aid to distressed victims increases substantially if a potential altruist’s empathy is engaged (Batson, 1991). Being able to discuss a game beforehand and to make promises to cooperate affect success (Dawes et al., 1990). The size of the resource, technology for exclusion and exploitation of the resource, and similar gritty details affect whether cooperation in commons management arises (Ostrom, 1990:202-204). Scientific findings again correspond well to personal experience. Sometimes we cooperate enthusiastically, sometimes reluctantly, and sometimes not at all. People vary considerably in their willingness to cooperate even under the same environmental conditions. Institutions matter. People from different societies behave differently because their habits have been inculcated by long participation in societies with different institutions. In repeated play common property experiments, initial defections induce further defections until the contribution to the public-goods sector approaches zero. However, if players are allowed to exercise strategies they might use in the real world, for example to punish those who defect, participation in the commons stabilizes (Fehr and Tyran, 1996). The strategies for successfully managing commons are generally institutionalized in sets of rules that have legitimacy in the eyes of the participants (Ostrom, 1990:Chapter 2). Families, local communities, employers, nations, and governments all tap our loyalties with rewards and punishments and greatly influence our behavior. Institutions are the product of evolution. The elegant studies by Nisbett’s group show how people’s affective and cognitive styles become intimately entwined with their social institutions (Cohen and Vandello, 2001; Nisbett and Cohen, 1996; Nisbett et al., in press). Because such complex traditions are so deeply ingrained, they are slow both to emerge and to decay. Many commons management institutions have considerable time depths (Ostrom, 1990:Chapter 3). Throughout most of human history, institutional change was so slow as to be nearly imperceptible by individuals. Today, change is rapid enough to be perceptible. Even universities, impeded as they are by conservative faculties deeply suspicious of change, change measurably on the time scale of a generation. Variation in institutions is huge. Already with its very short list of societies and games, the experimental ethnography approach of Henrich et al. (2001) and Nisbett et al. (in press) has uncovered striking differences. The cross-cultural commons work has uncovered much more, suggesting that a rich trove awaits the experimentalists. Agrawal (this volume:Chapter 2) describes the large number of conditions (38 and counting) that have been shown to affect whether local coop-

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The Drama of the Commons eration in commons management arises. Plausibly, design complexity, coordination equilibria, and other phenomena generate multiple evolutionary equilibria and much historical contingency in the evolution of particular institutions (Boyd and Richerson, 1992c). We all have at least some experience of how differently different communities, different universities, and different countries solve the same problems. Evolutionary Models Can Explain the Nature of Preferences and Institutions These facts present a challenge to rational actor theories. High levels of cooperation are difficult to reconcile with the usual assumption of self-regarding preferences, and the diversity of institutional solutions is a challenge to any theory based on a universal human nature. The “second generation” bounded rational choice theory championed by Ostrom (1998), and the “situated” rational choice characterized by McCay (this volume:Chapter 11), address these challenges from within the rational choice tradition. These approaches add a psychological basis and institutional constraints to the standard rational choice theory. Although psychological and social structures are invoked to explain individual behavior and its variation, an explanation for psychology and social structure is not part of the theory. Evolutionary theory permits us to address the origin of preferences. A number of economists have noted the neat fit between evolutionary theory and economic theory (Becker, 1976; Hirshleifer, 1977). Evolution, they observed, explains what organisms want, and economics explains how they should go about getting what they want. Without evolution, preferences are exogenous, to be estimated empirically, but not explained. To do a satisfactory job of explaining human social behavior, we need to expand the spare concept of preferences to include the conceptually richer properties of individuals and institutions of bounded and situated rationality. Then, to explain why humans have the unusual forms of social behavior depicted in our list of stylized facts, we need to appeal, we believe, to the special properties of cultural evolution. Evolutionary models have both intellectual and practical payoffs. The intellectual payoff is that evolutionary models link answers to contemporary puzzles to crucial long time-scale processes. The most important economic phenomenon of the past 500 years is the rise of capitalist economies and their tremendous impact on every aspect of human life. Expanding the time scale a bit, the most important phenomena of the past 10 millennia are the evolution of ever more complex social systems and ever more sophisticated technology following the origins of agriculture. A real explanation of both current behavior and its variation must be linked to such long-run processes, where the times to reach evolutionary equilibria are measured in millennia. More practically, the dynamism of

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The Drama of the Commons the contemporary world creates major stresses on the institutions that are used to manage commons. Evolutionary theory often will be useful because it will lead to an understanding of how to accelerate institutional evolution to better track rapid technological and economic change. (For an analogous argument in the context of medical practice, see Nesse and Williams, 1995.) Evolutionary Models Account for the Processes That Shape Heritable Genetic and Cultural Variation Through Time Evolutionary explanations are recursive. Individual behavior results from an interaction of inherited attributes and environmental contingencies. In most species genes are the main inherited attributes, but in humans inherited cultural information is also important. Individuals with different inherited attributes may develop different behaviors in the same environment. Every generation, evolutionary processes—natural selection is the prototype—impose environmental effects on individuals as they live out their lives. Cumulated over the whole population, these effects change the pool of inherited information, so that the inherited attributes of individuals in the next generation differ, usually subtly, from the attributes in the previous generation. Over evolutionary time, a lineage cycles through the recursive pattern of causal processes once per generation, more or less gradually shaping the gene pool and thus the succession of individuals that draw samples of genes from it. Statistics that describe the pool of inherited attributes, such as gene frequencies, are basic state variables of evolutionary analysis. They are what change over time. Note that in a recursive model, we explain individual behavior and population-level processes in the same model. Individual behavior depends, in any given generation, on the gene pool from which inherited attributes are sampled. The pool of inherited attributes depends in turn on what happens to a population of individuals as they express those attributes. Evolutionary biologists have a long list of processes that change the gene frequencies, including natural selection, mutation, and genetic drift. However, no organism experiences natural selection. They either live or die; reproduce or fail to reproduce. If, in a particular environment, some types of individuals do better than others and if this variation has a heritable basis, then we label as “natural selection” the resulting changes in gene frequencies. We use abstract categories like selection to describe such specific events because we wish to build up, concrete case by concrete case, some useful generalizations about evolutionary process. Few would argue that evolutionary biology is the poorer for investing effort in the generalizing project. Although the processes that lead to cultural change are very different from those that lead to genetic change, their logic is the same. For example, the cultural generation time is short in the case of ideas that spread rapidly, but modeling rapidly evolving cultural phenomena like semiconductor technology presents no special problems (Boyd and Richerson, 1985:68-69). Similarly, human choices

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The Drama of the Commons include ones that modify inherited attributes directly rather than indirectly by natural selection. These “Lamarckian” effects are added easily to models, and the models remain evolutionary so long as rationality remains bounded. The degenerate case, of course, needs no recursion because everything happens in the first generation (instantly in a typical rational choice model). Evolutionary models are a natural extension of the concept of bounded rational choice. They help explain how the innate and cultural constraints on choice and on rationality arise (Boyd and Richerson, 1993). Evolution is Multilevel Evolutionary theory is always multilevel; at a minimum it keeps track of properties of individuals, like their genotypes, and of the population, such as the frequency of a particular gene. Other levels may also be important. Phenotypes are derived from many genes interacting with each other and the environment. Populations may be structured, perhaps divided into social groups with limited exchanges of members. Thus, evolutionary theories are systemic, integrating every part of biology. In principle, everything that goes into causing change through time plays its proper part in the theory. This in-principle completeness led Mayr (1982) to speak of “proximate” and “ultimate” causes in biology. Proximate causes are those that physiologists and biochemists generally treat by asking how an organism functions. These are the causes produced by individuals with attributes interacting with environments and producing effects on them. Do humans use innate cooperative propensities to solve commons problems or do they have only self-interested innate motives? Or are the causes more complex than either proposal? Ultimate causes are evolutionary. The ultimate cause of an organism’s behavior is the history of evolution that shaped the gene pool from which our samples of innate attributes are drawn. Evolutionary analyses answer why questions. Why do human communities typically solve at least some of the commons dilemmas and other cooperation problems on a scale unknown in other apes and monkeys? Human-reared chimpanzees are capable of many human behaviors, but they nevertheless retain many chimp behaviors and cannot act as full members of a human community (Temerlin, 1975). Thus we know that humans have different innate influences on their behavior than chimpanzees, and these must have arisen in the course of the two species’ divergence from our common ancestor. In Darwinian evolutionary theories, the ultimate sources of cooperative behavior are classically categorized into three evolutionary processes operating at different levels of organization. Individual-level selection. Individuals and the variants they carry are obviously a locus of selection. Selection at this level favors selfish individuals who

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The Drama of the Commons are evolved to maximize their own survival and reproductive success. Pairs of self-interested actors can cooperate when they interact repeatedly (Axelrod and Hamilton, 1981; Trivers, 1971). Alexander (1987) argued that such reciprocal cooperation also can explain complex human social systems, but most formal modeling studies make this proposal doubtful (Boyd and Richerson, 1988, 1989; Leimar and Hammerstein, 2001; Nowak and Sigmund, 1998). Kin selection. Hamilton’s (1964) papers showing that kin should cooperate to the extent that they share genes identical by common descent offer one of the theoretical foundations of sociobiology. Kin selection can lead to cooperative social systems of a remarkable scale, as illustrated the colonies of termites, ants, and some bees and wasps. However, most animal societies are small because individuals have few close relatives. It is the fecundity of insects, and in one case rodents, that permits a single queen to produce huge numbers of sterile workers and hence large, complex societies composed of close relatives (Campbell, 1983). Group selection. Selection can act on any pattern of heritable variation that exists (Price, 1970). Darwin’s model of the evolution of cooperation by intertribal competition is perfectly plausible, as far as it goes. The problem is that genetic variation between groups other than kin groups is hard to maintain unless the migration between groups is very small or unless some very powerful force generates between-group variation (Aoki, 1982; Boorman and Levitt, 1980; Eshel, 1972; Levin and Kilmer, 1974; Rogers, 1990; Slatkin and Wade, 1978; Wilson, 1983). In the case of altruistic traits, selection will tend to favor selfish individuals in all groups, tending to aid migration in reducing variation between groups. The success of kin selection in accounting for the most conspicuous and highly organized animal societies (except humans) has convinced most, but by no means all, evolutionary biologists that group selection is of modest importance in nature (see Sober and Wilson, 1998, for a group selectionist’s eye view of the controversy). We could make this picture much more complex by adding higher and lower levels and cross-cutting forms of structure. Many examples from human societies will occur to the reader, such as gender. Indeed, Rice (1996) has demonstrated elegantly that selection on genes expressed in the different sexes sets up a profound conflict of interest between these genes. If female Drosophila are prevented from evolving defenses, male genes will evolve that seriously degrade female fitness. The genome is full of such conflicts, usually muted by the fact that an individual’s genes are forced by the evolved biology of complex organisms to all have an equal shot at being represented in one’s offspring. Our own bodies are a group-selected community of genes organized by elaborate “institutions” to ensure fairness in genetic transmission, such as the lottery of meiosis that gives each chromosome of a pair a fair chance at entering the functional gamete (Maynard Smith and Szathmáry, 1995).

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The Drama of the Commons Culture Evolves In theorizing about human evolution, we must include processes affecting culture in our list of evolutionary processes alongside those that affect genes. Culture is a system of inheritance. We acquire behavior by imitating other individuals much as we get our genes from our parents. A fancy capacity for high-fidelity imitation is one of the most important derived characters distinguishing us from our primate relatives (Tomasello, 1999). We are also an unusually docile animal (Simon, 1990) and unusually sensitive to expressions of approval and disapproval by parents and others (Baum, 1994:218-219). Thus parents, teachers, and peers can rapidly, easily, and accurately shape our behavior compared to training other animals using more expensive material rewards and punishments. Finally, once children acquire language, parents and others can communicate new ideas quite economically. Our own contribution to the study of human behavior is a series of mathematical models in the Darwinian style of what we take to be the fundamental processes of cultural evolution (e.g., Boyd and Richerson, 1985). The application of Darwinian methods to the study of cultural evolution was advocated forcefully by Campbell (1965, 1975). Cavalli-Sforza and Feldman (1973) constructed the first mathematical models to analyze cultural recursions (see also Durham, 1991). The list of processes that shape cultural change includes: Biases. Humans do not passively imitate whatever they observe. Rather, cultural transmission is biased by decision rules that individuals apply to the variants they observe or try out. The rules behind such selective imitation may be innate or the result of earlier imitation or a mixture of both. Many types of rules might be used to bias imitation. Individuals may try out a behavior and let reinforcement guide acceptance or rejection. Or they may use various rules of thumb to reduce the need for costly trials and punishing errors. The use of a conformist rule of the form “when in Rome do as the Romans do” is an example that is important in our hypothesis about the origins of cooperative tendencies in human behavior. Nonrandom variation. Genetic innovations (mutations, recombinations) are random with respect to what is adaptive. Human individual innovation is guided by many of the same rules that are applied to biasing ready-made cultural alternatives. Bias and learning rules have the effect of increasing the rate of evolution relative to what can be accomplished by random mutation, recombination, and natural selection. We believe that culture originated in the human lineage as an adaptation to the Plio-Pleistocene ice-age climate deterioration, which included much rapid, high-amplitude variation of just the sort that would favor adaptation by biased innovation and imitation (Richerson and Boyd, 2000). Natural selection. Because selection operates on any form of heritable variation and imitation and teaching are forms of inheritance, selection will influence cultural as well as genetic evolution. However, selection on culture is liable

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The Drama of the Commons to favor behaviors different from those favored by selection on genes. Because we often imitate peers, culture is liable to selection at the subindividual level, potentially favoring pathogenic cultural variants—selfish memes (Blackmore, 1999). On the other hand, rules like conformist imitation have the opposite effect. By tending to suppress cultural variation within groups such rules protect variation between them, potentially exposing our cultural variation to much stronger group selection effects than our genetic variation (Henrich and Boyd, 1998; Soltis et al., 1995). Human patterns of cooperation may owe much to cultural group selection. Evolutionary Models Are Consistent with a Wide Variety of Theories Evolutionary theory prescribes a method, not an answer, and a wide range of particular hypotheses can be cast in an evolutionary framework. If population-level processes are important, we can set up a system for keeping track of heritable variation, and the processes that change it through time. Darwinism as a method is not at all committed to any particular picture of how evolution works or what it produces. The view that many social scientists have of Darwinism is influenced too heavily by the work of human sociobiologists. Many things can be said in defense of this enterprise (Borgerhoff-Mulder et al., 1997) and much useful work goes on under its major research programs, human behavioral ecology (Cronk et al., 2000) and evolutionary psychology (Barkow et al., 1992). However, these research programs have two major weaknesses: neglect of culture and a taboo against group selection. Sociobiologists typically assume that culture is a strictly proximate phenomenon, akin to individual learning (e.g., Alexander, 1979), or constrained so strongly by genes as to be virtually proximate (Wilson, 1998). As Alexander (1979:80) puts it, “Cultural novelties do not replicate or spread themselves, even indirectly. They are replicated as a consequence of the behavior of vehicles of gene replication.” Commons institutions are deeply rooted in cultural traditions. Theoretical models show that the processes of cultural evolution can behave differently in critical respects from those only including genes. If such effects are important in the real world, neglecting them is a bad bet to get the approximately correct answers we hope to win using evolutionary theory. Most evolutionary biologists believe that group beneficial behavior is always a side effect of individual payoffs. We have already noted the problems with maintaining variation between groups in theory and the seeming success of alternative explanations. Persuaded by the biologist’s arguments, most social science scholars from the Darwinian tradition have followed the argument forcefully articulated by Williams (1966) and have anathematized group selection.1 However, cultural variation is more plausibly susceptible to group selection than is genetic variation. For example, if people use a somewhat conformist bias in acquiring important social behaviors, the variation between groups needed for group selec-

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The Drama of the Commons tion to operate is protected from the variance-reducing force of migration between groups (Boyd and Richerson, 1985:Chapter 7). We believe considerable evidence supports the hypothesis that cultural group selection has played an important role in human social evolution (Richerson and Boyd, 2001). Evolutionary Models Are Widely Used in the Social Sciences Although evolutionary tools are not yet commonplace in the study of human behavior, the general approach we advocate has a long history (Campbell, 1965, 1975) and several vigorous currently active branches. We mentioned evolutionary psychology and human behavioral ecology already. Others include evolutionary economics (Alchian, 1950; Day and Chen, 1993; Gintis, 2000; Hodgson, 1993; Witt, 1992), evolutionary sociology (Dietz and Burns, 1992; Luhmann, 1982; Maryanski and Turner, 1992; McLaughlin, 1988), evolutionary organization science (Baum and McKelvey, 1999; Hannan and Freeman, 1989), evolutionary epistemology (Callebaut and Pinxten, 1987; Derksen, 1998; Hull, 1988), evolutionary behavior analysis (Baum, 1994), and applied mathematics (Vose, 1999). The concepts of the meme (Blackmore, 1999), of complex adaptive systems (Holland, 1995), and of universal Darwinism (Dennett, 1996) have attracted much attention. Some of the most interesting evidence for the importance of evolutionary theory in the study of culture comes from the not infrequent reinvention of basic Darwinism when scholars in the social sciences find themselves in need of it. Empirical research traditions with strongly Darwinian overtones include historical linguistics (Mallory, 1989), sociolinguistics (Labov, 1973), studies of the diffusion of innovations (Rogers, 1995), human social learning theory (Bandura, 1986), experimental cultural evolution (Insko et al., 1983), and religious demography (Roof and McKinney, 1987). Weingart and colleagues (1997) attempt a comprehensive survey of the issues involved in integrating the historically abiological and non-Darwinian theories of the social sciences with Darwinian theory from biology. EVOLUTION OF COOPERATIVE INSTITUTIONS Here we summarize a theory of institutional evolution that we have developed elsewhere in more detail (Richerson and Boyd, 1998, 1999, 2001). The theory is rooted in a mathematical analysis of the processes of cultural evolution and is, we argue in these papers, consistent with much empirical data. We make limited claims for our particular hypotheses, although we think that the thrust of the empirical data as summarized by the stylized facts already noted is much harder on current alternatives. We make a much stronger claim that a dual gene-culture theory of some kind will be necessary to account for the evolution of human cooperative institutions. Understanding the evolution of contemporary human cooperation requires attention to two different time scales. First, a long period of evolution in the

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The Drama of the Commons Pleistocene shaped the innate “social instincts” that underpin modern human behavior. During this period, much genetic change occurred as a result of humans living in groups with social institutions heavily influenced by culture, including group-selected culture (Richerson and Boyd, 2000). On this time scale genes and culture coevolve, and cultural evolution is plausibly a leading rather than lagging partner in this process. Then, only about 10,000 years ago, the origins of agricultural subsistence systems laid the basis for revolutionary changes in the scale of social systems. The evidence suggests that genetic changes in the social instincts over the past 10,000 years are insignificant. Rather, the evolution of complex societies has involved the relatively slow cultural accumulation of institutional “work-arounds.” These take advantage of a psychology evolved to cooperate with distantly related and unrelated individuals belonging to the same symbolically marked tribe while coping more or less successfully with the fact that these social systems are larger, more anonymous, and more hierarchical than the tribal scale ones of the late Pleistocene (Richerson and Boyd, 1998, 1999). Tribal Social Instincts Hypothesis Our hypothesis is premised on the idea that group selection plays a more important role in shaping culturally transmitted variation than it does in shaping genetic variation. As a result, humans have lived in social environments characterized by high levels of cooperation for as long as culture has played an important role in human development. To judge from the other living apes, our remote ancestors had only rudimentary culture (Tomasello, 1999) and lacked cooperation on a scale larger than groups of close kin (Boehm, 1999). The difficulty of constructing theoretical models of group selection on genes favoring cooperation matches neatly with the empirical evidence that cooperation in most social animals is limited to kin groups. In contrast, rapid cultural adaptation can lead to ample variation among groups whenever multiple stable social equilibria exist, due to conformist social learning, symbolically marked boundaries, or moralistic enforcement of norms (Boyd and Richerson, 1992a). Such models of group selection are relatively powerful because they only require the social, not physical, extinction of groups. Formal theoretical models suggest that conformism is an adaptive heuristic for biasing imitation under a wide variety of conditions (Boyd and Richerson, 1985:Chapter 7; Henrich and Boyd, 1998; Simon, 1990). Similarly, symbolic group marking arises for adaptive reasons in cultural evolution models in which either ecological differences or different solutions to games of coordination make the imitation of behaviors common in neighboring groups maladaptive in one’s own group (Boyd and Richerson, 1987; McElreath et al., no date). Models of moralistic punishment (Boyd and Richerson, 1992c) lead to multiple stable social equilibria and to reductions in noncooperative strategies if punishment is prosocial. A consequence, we believe, is that a growing reliance on cultural evolution led to larger, more cooperative societies among humans over the past 250,000 years or so.

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The Drama of the Commons reforms. One of the main reasons the German General Staff worked so well was that the prestigious and rather scholarly staff officers routinely served in line roles and earned the respect of line officers. In a few disciplines—engineering, economics—the flow of personnel from academic to practical line and staff roles is perhaps routine enough to resemble an informal general staff. In most disciplines academic and practitioner roles are mutually exclusive, practically speaking. The various agricultural extension services and other applied science organizations could be prospected for models. A practical scheme to “grow” innovative commons management institutions is perhaps only an inspirational innovation or two away from practicality. The two senior authors, who have had considerable, interesting, and rewarding experience as staffers in applied science and policy contexts, must admit that they found no way in the end to combine such work with an academic career. CONCLUSIONS In this chapter, we have tried to tie together the literature on the evolution of cooperation with the literature on commons management institutions. We believe an interesting parallel exists between the sophisticated bounded rationality models necessary to account for the behavior of people toward commons and dual inheritance or gene-culture coevolutionary theory. People behave in experiments and in the field as if they have strong—perhaps innate—dispositions to cooperate, although dispositions vary considerably from person to person, society to society, and time to time. The variation is best explained by the existence of complex cultural traditions of social behavior, the collective results of which we call social institutions. Our ability to organize cooperation on a scale considerably larger than predicted by theory based on unconstrained selfish rationality, or by most evolutionary mechanisms, is one of the most striking features of our species. Another striking feature is our extraordinary facility for imitation and teaching. Our main hypothesis is that the co-occurrence of culture and cooperation in our species is not a coincidence. Group selection on cultural variation provides a plausible mechanism by which large-scale cooperation might arise. Cultural group selection is a slow process, at least in some models we have studied, so supplementary processes are likely to be more important in the shorter run evolution of cooperative institutions. The cooperative dispositions, cultural or innate, favored originally by cultural group selection or some similar process will inevitably act as biases of cultural innovation and transmission. All else equal, people will tend to favor innovations that seem fair, that are efficient producers of public goods, and that contribute to their ingroup’s position relative to competing outgroups. As team sports show, people play games of cooperation for fun. We can even organize institutions to promote desirable institutional evolution, ranging from research universities and political parties to village assemblies. Of course, people are

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The Drama of the Commons hardly perfect paragons of cooperation. Our mixture of altruistic and selfish propensities varies across cultures but neither element is ever suppressed entirely. Gene-culture coevolution theory has a natural account of our conditional and incomplete altruism. At root, reproductive competition between the cooperators in human societies means that selection on genes still acts strongly to favor behavior enhancing inclusive fitness. Group selection on culture can only partially mitigate selfish and nepotistic impulses, not eliminate them. Aside from providing an ultimate explanation for the patterns of cooperation we observe in humans, we hope the application of evolutionary theory to the understanding of commons institutions will lead to means to improve commons management. If our particular evolutionary theory is correct, we have good news and bad news for the practitioner. The good news is that we have much better raw material to work with improving commons management than the selfish rationality theorists think we have. The bad news is that institutions to capitalize on our prosocial instincts and traditions evolve relatively slowly and uncertainly. Regress is possible as well as progress. Cooperation within groups is all too often devoted to unhelpful if not destructive conflicts with other groups, as in the conflict between rivalrous national goals and the regulation of the global commons. The new theory of the commons already understands all these things. Evolutionary theory offers a program for investigating just how institutions do evolve. We have outlined a little of the complexity possible when several different evolutionary processes can be at work, some stronger and some weaker, and all depending, at least to some extent, on the case at hand. The products of evolution are not only complex but also diverse. Exploring the tempo and mode of cultural evolution is a long-term project. After all, biologists are still at work on organic evolution a century and a half after Darwin, and they’re still having plenty of fun. Of course, they have so many species to work on and we are only one, albeit a more than ordinarily diverse and complex one. In some ways cultural evolution is easier to study than organic evolution. Cultures change faster than gene pools. Historians and anthropologists have compiled vast amounts of qualitative information about our evolution and diversity and some innovative scholars have produced quantitative data. We believe that all the empirical methods needed to study cultural evolution have been used effectively in some specialized application or another, even if they are not yet in every social scientist’s toolkit. We believe there is nothing to lose—and everything to gain—by developing and verifying a rigorous evolutionary theory of human behavior. NOTE 1   Several prominent modern Darwinians—Hamilton (1975), E.O. Wilson (1975:561-562), Alexander (1987:169), and Eibl-Eibesfeldt (1982)—have given serious consideration to group selection as a force in the special case of human ultrasociality. They are impressed, as we are, by the organization of human populations into units that engage in sustained, lethal combat with other groups, not to mention other forms of cooperation. The trouble with a straightforward group selection hypoth-

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The Drama of the Commons     esis is our mating system. We do not build up concentrations of intrademic relatedness like social insects, and few demic boundaries are without considerable intermarriage. Moreover, the details of human combat are more lethal to the hypothesis of genetic group selection than to the human participants. For some of the most violent groups among simple societies, wife capture is one of the main motives for raids on neighbors, a process that hardly could be better designed to erase genetic variation between groups. REFERENCES Acheson, J.M. 1988 The Lobster Gangs of Maine. Hanover, NH: University Press of New England. Alchian, A.A. 1950 Uncertainty, evolution and economic theory. Journal of Political Economy 58:211-222. Alexander, R.D. 1979 Darwinism and Human Affairs. Seattle: University of Washington Press. 1987 The Biology of Moral Systems. Hawthorne, NY: Aldine de Gruyter. Anderson, B.R. 1991 Imagined Communities: Reflections on the Origin and Spread of Nationalism. Rev. and extended ed. London: Verso. Aoki, K. 1982 A condition for group selection to prevail over counteracting individual selection. Evolution 36:832-842. Arrow, K.J. 1963 Social Choice and Individual Values. 2d ed. New Haven, CT: Yale University Press. Axelrod, R., and W.D. Hamilton 1981 The evolution of cooperation. Science 211:1390-1396. Baland, J.M., and J.P. Platteau 1996 Halting Degradation of Natural Resources: Is There a Role for Rural Communities? Oxford, Eng.: Oxford University Press. Bandura, A. 1986 Social Foundations of Thought and Action: A Social Cognitive Theory. Englewood Cliffs, NJ: Prentice-Hall. Banfield, E.C. 1958 The Moral Basis of a Backward Society. Glencoe, IL: Free Press. Barkow, J.H., L. Cosmides, and J. Tooby 1992 The Adapted Mind: Evolutionary Psychology and the Generation of Culture. New York: Oxford University Press. Batson, C.D. 1991 The Altruism Question: Toward a Social Psychological Answer. Hillsdale, NJ: Lawrence Erlbaum Associates. Baum, J.A.C., and B. McKelvey, eds. 1999 Variations in Organization Science: In Honor of Donald T. Campbell. Thousand Oaks, CA: Sage Publications. Baum, W.B. 1994 Understanding Behaviorism: Science, Behavior, and Culture. New York: HarperCollins. Becker, G.S. 1976 Altruism, egoism, and genetic fitness: Economics and sociobiology. Journal of Economic Literature 14:817-826. Bettinger, R.L. 1991 Hunter-Gatherers: Archaeological and Evolutionary Theory. New York: Plenum Press.

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