Abstract: Theoretical models have demonstrated the possibility of stable cost-free signalling of need between relatives. The stability of these cost-free equilibria depends on the indirect fitness cost of cheating and deceiving a donor into giving away resources. We show that this stability is highly sensitive to the distribution of need among signallers and receivers. In particular, cost-free signalling is likely to prove stable only if there is very large variation in need (such that the least-needy individuals stand to gain much less than the most-needy individuals from additional resources). We discuss whether these conditions are likely to be found in altricial avian breeding systems--the most intensively studied instance of signalling of need between relatives. We suggest that cost-free signalling is more likely to prove stable and will provide parents with more information during the earlier phases of chick growth, when parents can more easily meet the demands of a brood (and chicks are more likely to reach satiation). Later, informative yet cost-free signalling is unlikely to persist.

Abstract: Animals frequently use signals that travel further than the spacing between individuals. For every intended recipient of a given signal there are likely to be many other individuals that receive information. Eavesdropping on signalling interactions between other individuals provides a relatively cost-free method of assessing future opponents or mates. Male great tits (Parus major) extract relative information from such interactions between individuals unknown to them. Here, we show that male great tits can take information gathering a stage further and obtain more information about a previously unencountered intruder, by the hitherto unknown capability of combining information gathered by eavesdropping with that derived from their own direct interaction with an individual. Prior experience with an intruder (A) was achieved by subjecting a focal male to different levels of intrusion simulated using interactive playback. This intruder (A) then took part in a simulated interaction with an unknown male (B) outside the territorial boundary of the focal males. In response to subsequent intrusion by the second male (B), focal males showed low song output in response to males that had lost to a male that the subject was able to beat. Males of known high quality, or those about which information was ambiguous, elicited a high level of song output by focal males. We discuss the implications of this finding for the evolution of communication and social behaviour.

Abstract: In several lek-breeding populations of birds and mammals, females arriving on leks tend to join males that already have females in their territories. This might occur either because females have an evolved preference for mating with males that are attractive to other females, or because they join groups of other females to obtain greater safety from predation or dangerous harassment by males. We have previously used controlled experiments to show that oestrous fallow deer females join males with established harems because they are attracted to female groups rather than to the males themselves. Here we demonstrate that the preference for males with females over males without females is specific to oestrous females and weak or absent in anoestrous ones, and that it is not associated with a preference for mating with males that have previously been seen to mate with other females. Furthermore, oestrous females given the choice between males that do not already have females with them show no significant preference for antlered over deantlered males or for older males over younger ones. We conclude that female attraction to other females on the lek is likely to be an adaptation to avoiding harassment in mixed-sex herds. In this situation, a male's ability to maintain the cohesion of his harem may be the principal cause of variation in mating success between males.

Abstract: An animal's within-group spatial position has several important fitness consequences. Risk of predation, time spent engaging in antipredatory behavior and feeding competition can all vary with respect to spatial position. Previous research has found evidence that feeding rates are higher at the group edge in many species, but these studies have not represented the entire breadth of dietary diversity and ecological situations faced by many animals. In particular the presence of concentrated, defendable food patches can lead to increased feeding rates by dominants in the center of the group that are able to monopolize or defend these areas. To fully understand the tradeoffs of within-group spatial position in relation to a variety of factors, it is important to be able to predict where individuals should preferably position themselves in relation to feeding rates and food competition. A qualitative model is presented here to predict how food depletion time, abundance of food patches within a group, and the presence of prior knowledge of feeding sites affect the payoffs of different within-group spatial positions for dominant and subordinate animals. In general, when feeding on small abundant food items, individuals at the front edge of the group should have higher foraging success. When feeding on slowly depleted, rare food items, dominants will often have the highest feeding rates in the center of the group. Between these two extreme points of a continuum, an individual's optimal spatial position is predicted to be influenced by an additional combination of factors, such as group size, group spread, satiation rates, and the presence of producer-scrounger tactics.

Abstract: Conflict management is one of the primary requirements for social complexity. Of the many forms of conflict management, one of the rarest and most interesting is third-party policing, or intervening impartially to control conflict. Third-party policing should be hard to evolve because policers personally pay a cost for intervening, while the benefits are diffused over the whole group. In this study we investigate the incidence and costs of policing in a primate society. We report quantitative evidence of non-kin policing in the nonhuman primate, the pigtailed macaque. We find that policing is effective at reducing the intensity of or terminating conflict when performed by the most powerful individuals. We define a measure, social power consensus, that predicts effective low-cost interventions by powerful individuals and ineffective, relatively costly interventions by low-power individuals. Finally, we develop a simple probabilistic model to explore whether the degree to which policing can effectively reduce the societal cost of conflict is dependent on variance in the distribution of power. Our data and simple model suggest that third-party policing effectiveness and cost are dependent on power structure and might emerge only in societies with high variance in power.