Abstract: The iterated prisoner's dilemma game, or IPD, has now established itself as the orthodox paradigm for theoretical investigations of the evolution of cooperation; but its scope is restricted to reciprocity, which is only one of three categories of cooperation among unrelated individuals. Even within that category, a cooperative encounter has in general three phases, and the IPD has nothing to say about two of them. To distinguish among mechanisms of cooperation in nature, future theoretical work on the evolution of cooperation must distance itself from economics and develop games as a refinement of ethology's comparative approach.

Abstract: One specific prediction emerging from trait-group models of natural selection is that when individuals possess traits that benefit other group members, natural selection will favor “cheating” (i.e. not possessing the group-beneficial trait) within groups. Cheating is selected within groups because it allows individuals to avoid bearing the relative costs typically associated with group-beneficial traits, but to still reap the benefits associated with the acts of other group members. Selection between groups favors traits that benefit other group members. The relative strength of within- and between-group selection then determines the equilibrium frequency of those who produce group-beneficial traits and those that do not. Here we demonstrate that individual-level selection, that is selection within groups can also produce an intermediate frequency of such group-beneficial traits by frequency-dependent selection. The models we develop are general in nature, but were inspired by the evolution of antibiotic resistance in bacteria. The theory developed here is distinct from prior work that relies on reciprocity or kinship per'se to achieve cooperation and altruism among group members.

Abstract: To evaluate the role of individual recognition in the evolution of cooperation, we formulated and analyzed a genetic algorithm model (EvCo) for playing the Iterated Prisoner's Dilemma (IPD) game. Strategies compete against each other during each generation, and successful strategies contribute more of their attributes to the next generation. Each strategy is encoded on a `chromosome' that plays the IPD, responding to the sequences of most recent responses by the interacting individuals (chromosomes). The analysis reported in this paper considered different memory capabilities (one to five previous interactions), pairing continuities (pairs of individuals remain together for about one, two, five, or 1000 consecutive interactions), and types of individual recognition (recognition capability was maximal, nil, or allowed to evolve between these limits). Analysis of the results focused on the frequency of mutual cooperation in pairwise interactions (a good indicator of overall success in the IPD) and on the extent to which previous responses by the focal individual and its partner were associated with the partner's identity (individual recognition). Results indicated that a fixed, substantial amount of individual recognition could maintain high levels of mutual cooperation even at low pairing continuities, and a significant but limited capability for individual recognition evolved under selection. Recognition generally increased mutual cooperation more when the recent responses of individuals other than the current partner were ignored. Titrating recognition memory under selection using a fitness cost suggested that memory of the partner's previous responses was more valuable than memory of the focal's previous responses. The dynamics produced to date by EvCo are a step toward understanding the evolution of social networks, for which additional benefits associated with group interactions must be incorporated.

Abstract: Ever since Fisher (1958) formalized models of sexual selection, female mate choice has been assumed to be a genetically determined trait. Females, however, may also use social cues to select mates. One such cue might be the mate choice of conspecifics. Here we report the first direct evidence that a female's preference for a particular male can in fact be reversed by social cues. In our experiments using the Trinidadian guppy (Poecilia reticulata), this reversal was mediated by mate-copying opportunities, such that a female (the 'focal' female) is given the opportunity to choose between two males, followed by a period in which she observes a second female (the 'model' female) displaying a preference for the male she herself did not prefer initially. When allowed to choose between the same males a second time, compared with control tests, a significant proportion of focal females reversed their mate choice and copied the preference of the model female. These results provide strong evidence for the role of non-genetic factors in sexual selection and underlie the need for new models of sexual selection that explicitly incorporate both genetic and cultural aspects of mate choice.

Abstract: Coalition formation has been documented in a diverse array of taxa, yet there has been little formal analysis of polyadic interactions such as coalitions. Here, we develop an optimality model which examines the role of winner and loser effects in shaping coalition formation. We demonstrate that the predicted patterns of alliances are strongly dependent on the way in which winner and loser effects change with contestant strength. When winner and loser effects decrease with the resource-holding power (RHP) of the combatants, coalitions will be favoured between the strongest members of a group, but not between the weakest. If, in contrast, winner and loser effects increase with RHP, exactly the opposite predictions emerge. All other things being equal, intervention is more likely to prove worthwhile when the beneficiary of the aid is weaker (and its opponent is stronger), because the beneficiary is then less likely to win without help. Consequently, intervention is more probable when the impact of victory on the subsequent performance of a combatant increases with that individual's strength because this selects for intervention in favour of weaker combatants. The published literature on hierarchy formation does not reveal how winner and loser effects actually change with contestant strength and we therefore hope that our model will spur others to collect such data; in this light we suggest an experiment which will help to elucidate the nature of winner and loser effects and their impact on coalition formation in animals.