Abstract: Close associations between adult males and lactating females and their dependent infants are not commonly described in non-monogamous mammals. However, such associations [sometimes called friendships (Smuts 1985)] are regularly observed in several primate species in which females mate with multiple males during the fertile period. The absence of mating exclusivity among friends suggests that males should invest little in infant care, raising questions about the adaptive significance of friendship bonds. Using data from genetic paternity analyses, patterns of behavior, and long-term demographic and reproductive records, we evaluated the extent to which friendships in four multi-male, multi-female yellow baboon (Papio cynocephalus) groups in Amboseli, Kenya represent joint parental care of offspring or male mating effort. We found evidence that mothers and infants benefited directly from friendships; friendships provided mother–infant dyads protection from harassment from other adult and immature females. In addition, nearly half of all male friends were the genetic fathers of offspring and had been observed mating with mothers during the days of most likely conception for those offspring. In contrast, nearly all friends who were not fathers were also not observed to consort with the mother during the days of most likely conception, suggesting that friendships between mothers and non-fathers did not result from paternity confusion. Finally, we found no evidence that prior friendship increased a male’s chances of mating with a female in future reproductive cycles. Our results suggest that, for many male–female pairs at Amboseli, friendships represented a form of biparental care of offspring. Males in the remaining friendship dyads may be trading protection of infants in exchange for some resources or services not yet identified. Our study is the first to find evidence that female primates gain social benefits from their early associations with adult males.

Abstract: Network analysis has a long history in the mathematical and social sciences and the aim of this introduction is to provide a brief overview of the potential that it holds for the study of animal behaviour. One of the most attractive features of the network paradigm is that it provides a single conceptual framework with which we can study the social organisation of animals at all levels (individual, dyad, group, population) and for all types of interaction (aggressive, cooperative, sexual etc.). Graphical tools allow a visual inspection of networks which often helps inspire ideas for testable hypotheses. Network analysis itself provides a multitude of novel statistical tools that can be used to characterise social patterns in animal populations. Among the important insights that networks have facilitated is that indirect social connections matter. Interactions between individuals generate a social environment at the population level which in turn selects for behavioural strategies at the individual level. A social network is often a perfect means by which to represent heterogeneous relationships in a population. Probing the biological drivers for these heterogeneities, often as a function of time, forms the basis of many of the current uses of network analysis in the behavioural sciences. This special issue on social networks brings together a diverse group of practitioners whose study systems range from social insects over reptiles to birds, cetaceans, ungulates and primates in order to illustrate the wide-ranging applications of network analysis.

Abstract: The formation of collaborating pairs by individuals belonging to two different classes occurs in the contexts of reproduction and intea-specific cooperation as well as of inter-specific mutualism. There is potential for partner choice and for competition for access to preferred partners in all three contexts. These selective forces have long been recognised as important in sexual selection, but their impact is not yet appreciated in cooperative and mutualistic systems. The formation of partnerships between members of different classes has much in common with the conclusion of trade agreements in human markets with two classes of traders, like producers and consumers, or employers and employees. Similar game-theoretical models can be used to predict the behaviour of rational traders in human markets and the evolutionarily stable strategies used in biological markets. We present a formal model in which the influence of the market mechanism on selection is made explicit. We restrict ourselves to biological markets in which: (1) Individuals do not compete over access to partners in an agonistic manner, but rather by outcompeting each other in those aspects that are preferred by the choosing party. (2) The commodity the partner has to offer cannot be obtained by the use of force, but requires the consent of the partner. These two restrictions ensure a dominant role for partner choice in the formation of partnerships. In a biological market model the decision to cooperate is based on the comparison between the offers of several potential partners, rather than on the behaviour of a single potential partner, as is implicitly assumed in currently accepted models of cooperation. In our example the members of one class A offer a commodity of fixed value in exchange for a commodity of variable value supplied by the other class, B. We show that when the B-class outnumbers the A-class sufficiently and the cost for the A-class to sample the offers of the B-class are low, the choosiness of the A-class will lead to selection for the supply of high value commodities by the B-class (Fig. 3a). Under the same market conditions, but with a high sampling cost this may still be the evolutionariy stable outcome, but another pair of strategies proves to be stable too: relaxed choosiness of class A coupled with low value commodities supplied by class B (Fig. 3b). We give a number of examples of mating, cooperative and mutualistic markets that resemble the low sampling cost situation depicted in Fig. 3a.

Abstract: There is an established and very influential view that primate societies have identifiable, persistent social organizations. It assumes that association patterns reflect long-term strategic interests that are not qualitatively perturbed by short-term environmental variability. We used data from two baboon troops in markedly different habitats over three consecutive seasons to test this assumption. Our results demonstrate pronounced cyclicity in the extent to which females maintained differentiated relationships. When food was plentiful, the companionships identified by social network analysis in the food-scarce season disappeared and were replaced by casual acquaintanceships more representative of mere gregariousness. Data from the fourth, food-scarce, season at one site indicated that few companions were re-united. It is likely that this reflected stochastic variation in individual circumstances. These results suggest that attention could profitably be paid to the effects of short-term local contingencies on social dynamics, and has implications for current theories of primate cognitive evolution.

Abstract: Place learning abilities represent adaptations that contribute also to foraging efficiency under given spatio-temporal conditions. We investigated if this ability in turn constrains decision making in two sympatric vole species: while the herbivorous common vole (Microtus arvalis) feeds on spatio-temporally predictable food resources (e.g. roots, tubers and shoots of plant tubers), the omnivorous bank vole (Myodes glareolus) additionally subsists on temporally unpredictable food resources (e.g. insects and seeds). Here, we compare the spatial reference memory and working memory of the two species. In an automated operant home cage with eight water places, female voles either had to learn the fixed position of non-depletable places (reference memory task) or learn and avoid previously visited water places depleted in a single visit (win-shift task). In the reference memory task, Microtus females required significantly more choices to find all water places, initially performed slightly worse than Myodes females, and displayed slightly lower asymptotic performance. Both species were highly similar in new learning of the same task. In the more complex win-shift task, asymptotic performance was significantly lower in Microtus (72% correct) than in Myodes (79%). Our results suggest that both vole species resemble each other in their efficiency to exploit habitats with low spatio-temporal complexity but may differ in their efficiency at exploiting habitats with temporally changing spatial food distributions. The results imply that spatial ability adjusted to specific food distributions may impair flexible use of habitats that differ in their food distribution and therefore, decrease a species’ chances of survival in highly dynamic environments.