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Cooperation, Evolution Of the increase in species richness with island area is much with latitude, many metacommunities may be sampled, more gradual than in a comparable continental region and the spatial autocorrelation among resources may be (that is, many island species–area relationships have very low for regions at different extremes along the gradi- much smaller exponents than in continental regions). ent. In such cases, the trends in species diversity observed Distribution–abundance relationships and related across space must refl ect differences in the capacity of the patterns are a consequence of asymmetries in ecological environment to support species. attributes among species such that some species are able to fi nd more resources across geographic space than others. SEE ALSO THE FOLLOWING ARTICLES Thus, within metacommunities, the relative abundances Allometry and Growth / Diversity Measures / Metacommunities / of species will also be asymmetric, and such asymmetries Neutral Community Ecology / Species Ranges in relative abundances in metacommunities are preserved even under short-term ecological drift. When a collection FURTHER READING of local communities are sampled using sampling units Blackburn, T. M., and K. J. Gaston, eds. 2003. Macroecology: concepts and of the same size (thus controlling for area effects), the consequences. Proceedings of the 43rd Annual Symposium of the British asymmetries in relative abundances of species within the Ecological Society (17–19 April 2002). Malden, MA: Blackwell Publishing. metacommunity will ensure that some species show up in Brown, J. H. 1995. Macroecology. Chicago: University of Chicago Press. Gaston, K. J., and T. M. Blackburn. 2000. Pattern and process in samples more often than others (Fig. 2). Abundances of macroecology. Malden, MA: Blackwell Science. species at local scales will also refl ect these asymmetries; Hubbell, S. P. 2001. The unifi ed neutral theory of biodiversity and biogeog- hence, species that show up more often in samples will raphy. Princeton, NJ: Princeton University Press. Huston, M. A. 1994. Biological diversity. Cambridge, UK: Cambridge also tend to be represented by more individuals within University Press. those samples. Maurer, B. A. 1999. Untangling ecological complexity: the macroscopic When examining species diversity along environmental perspective. Chicago: University of Chicago Press. Maurer, B. A. 2009. Spatial patterns of species diversity in terrestrial envi- gradients, the sampled geographic region may represent ronments. In S. A. Levin, ed. Princeton guide to ecology. Princeton, NJ: only a subset of sites contained within the metacommu- Princeton University Press. nity of the region. Since there is spatial autocorrelation in Rosenzweig, M. L. 1995. Species diversity in space and time . Cambridge, environmental conditions, sites that have more resources UK: Cambridge University Press. will typically have more species, and these sites will tend to be clustered together along the gradient. In more extensive samples, such as samples of species diversity CONTROL THEORY SEE OPTIMAL CONTROL THEORY COOPERATION, EVOLUTION OF MATTHEW R. ZIMMERMAN, RICHARD MCELREATH, AND PETER J. RICHERSON University of California, Davis Cooperation occurs when individuals act together for FIGURE 2 A schematic diagram showing how asymmetries in distribu- benefi cial results. Most evolutionary theory of coopera- tions of di4 erent species across geographic space result in di4 erences tion addresses the evolution of altruism, the most diffi cult in species diversity among a collection of sampling locations. Each type of cooperation to explain. However, observed coop- unimodal curve represents the geographic distribution of a single species. Note that some species are found across a greater span of eration can also result from environments favoring other geographic space than others. This would be a consequence of the kinds of cooperation, including mutualism and coordina- fact that some species will have attributes that will allow them to use tion. Theoreticians model the evolution of cooperation by a wider range of environmental conditions than others. The dashed lines represent geographic locations where a survey is taken. Surveys accounting for the effects of cooperative behavior at differ- are assumed to be of standard size. ent levels of analysis, including the genes, the individual, COOPERATION, EVOLUTION OF 155 and the group. Different mechanisms have been proposed as ones that encourage cooperative behavior, including limited dispersal, signaling, reciprocity, and biased trans- mission. All of these mechanisms are based on cooperators assortatively interacting with other cooperators. TYPES OF COOPERATION Mutualism Mutualistic behavior is helping behavior for which the producer’s costs are smaller than the producer’s ben- efi ts. Intraspecifi c mutualisms lack clear incentives for individuals to withhold cooperation, and so natural se- lection always favors cooperation over noncooperation. (Community ecologists reserve the term mutualism for interspecies relationships, though here, as in evolution- FIGURE 1 (A) A simple game illustrating mutualistic hunting in a hypothetical wolf population. Wolves get meat only if they hunt ary game theory generally, we use the term primarily cooperatively, thus the benefi ts of cooperation are always positive. for intraspecifi c interactions.) To see how mutualisms (B) The adaptive dynamics of the wolf population when individuals evolve, imagine a population of wolves that can either are randomly paired. Because it is always better to cooperate, natural selection increases the frequency of cooperative hunting away from pair up to hunt deer cooperatively or hunt deer nonco- the unstable noncooperative equilibrium to the stable equilibrium at operatively. (Of course, wolves can cooperatively hunt full cooperation. in larger groups, but the underlying logic of mutualisms holds even if we restrict our analysis to two individuals.) unstable since any introduction of cooperative wolves, If either of the wolves hunts individually, they cannot through genetic mutation or migration from other popu- bring down large game and go hungry. However, if the lations, would allow natural selection to push the popula- wolves hunt together, they can capture a deer and eat tion to the stable equilibrium. three units of meat. In some models of mutualisms, since cooperative in- How would cooperation evolve in a population of dividuals always achieve greater benefi ts than noncoop- wolves made up of cooperative hunters and noncoopera- erative individuals, the evolution of cooperative behavior tive hunters? We will assume that wolves that are more does not present much of a puzzle and does not require successful hunters are more likely to produce offspring. special explanations for why they persist in a population. The potential payoffs to the wolves in this situation are However, the origins of mutualisms can be more diffi - shown in Figure 1A. Both wolves have a choice of co- cult to explain, as initial benefi ts may not be suffi cient to operative hunting (C) and noncooperative hunting (N). exceed individual costs. For example, if in a proto-wolf The numbers in the grid show the payoffs for each com- population hunters are mainly solitary, then it stands to bination of behaviors, with the payoff to the Wolf A fi rst, reason that the skills necessary for successful cooperative followed by the payoff to Wolf B. Notice that a wolf only hunting will not exist. Therefore, early cooperative hunters has a chance at getting meat if it tries to hunt coopera- are not assured the large gains that may arise later, once tively. Thus, wolves that hunt cooperatively will tend cooperation gets a foothold through other mechanisms. In to produce more offspring and the trait for cooperative this way, other types of cooperation, such as coordination hunting will tend to spread in the population. (discussed next ), can evolve over time into mutualisms, The line in Figure 1B shows the adaptive dynamics of once cooperation is stabilized through other means. this wolf population. The arrows represent the direction of natural selection and show that, no matter where the Coordination mixture of hunting strategies starts, selection will tend In coordination contexts, natural selection favors common to push the population toward a stable cooperative equi- behavior, whether it is cooperative or noncooperative. For librium, represented by the solid circle. The open circle example, imagine that our population of wolves lives in at the other end of the line represents a noncooperative an environment with both deer and rabbits. Wolves now equilibrium where there are no cooperative wolves and have two strategies: hunt deer cooperatively, or solitarily they cannot reproduce. However, this equilibrium is hunt rabbits. If a wolf and its partner hunt cooperatively, 156 COOPERATION, EVOLUTION OF they manage to catch a deer with three units of meat the noncooperative rabbit-hunting equilibrium. These apiece as before. If, however, one of the wolves hunts deer equilibria are stable because the introduction of a non- and the other hunts rabbits, the deer hunter fails to bring conforming wolf to the population will have lower pay- down the game, but the rabbit hunter catches two rab- offs than the rest of the population obtains and thus be bits for two units of meat. However, if both wolves hunt outcompeted. Because, in coordination, the equilibrium rabbits individually, they are competing for the easy game preferred
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