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: Abstract Hormones play a central role in the physiology and behaviour of animals. The recent development of noninvasive techniques has increased information on physical and social states of individuals through hormone measurements. The relationships among hormones, life history traits and behaviours are, however, still poorly known. For the first time, we evaluated natural winter glucocorticoid and testosterone levels in young ungulates in relation to winter progression, diet quality and social rank. Overwinter, levels of glucocorticoid and testosterone decreased, possibly due to the decline of fawns" body mass. The relationships between hormone levels and diet quality were surprising: Fawns fed the control diet presented higher glucocorticoid and lower testosterone levels then fawns fed the poor diet, suggesting that control fawns faced a higher nutritional stress than those on the poor diet. Similarly to other studies on social mammals, we found no relationship between faecal glucocorticoid levels and social rank, suggesting that social stress was similar for dominant and subordinate fawns during winter. Testosterone levels were not correlated to social rank as found previously in groups of individuals forming stable social hierarchies and maintaining stable dominance relationships. The simultaneous suppression of glucocorticoid and testosterone levels suggests for the first time that young ungulates present a hormonal strategy to prevent fast depletion of limited proteins and fat resources during winter.

Abstract: Considerable interspeci®c variation in female social relationships occurs in gregarious primates, particularly with regard to agonism and cooperation between females and to the quality of female relationships with males. This variation exists alongside variation in female philopatry and dispersal. Socioecological theories have tried to explain variation in female-female social relationships from an evolutionary perspective focused on ecological factors, notably predation and food distribution. According to the current ``ecological model'', predation risk forces females of most diurnal primate species to live in groups; the strength of the contest component of competition for resources within and between groups then largely determines social relationships between females. Social elationships among gregarious females are here characterized as DispersalEgalitarian, Resident-Nepotistic, Resident-Nepotistic-Tolerant, or Resident-Egalitarian. This ecological model has successfully explained i€erences in the occurrence of formal submission signals, decided dominance relation ships, coalitions and female philopatry. Group size and female rank generally a€ect female reproduction success as the model predicts, and studies of closely related species in di€erent ecological circumstances underscore the importance of the model. Some cases, however, can only be explained when we extend the model to incorporate the e€ects of infanticide risk and habitat saturation. We review evidence in support of the ecological model and test the power of alternative models that invoke between-group competition, forced female philopatry, demographic female recruitment, male interventions into female aggression, and male harassment.
Not one of these models can replace the ecological model, which already encompasses the between-group competition. Currently the best model, which explains
several phenomena that the ecological model does not, is a ``socioecological model'' based on the combined importance of ecological factors, habitat saturation and infanticide avoidance. We note some points of similarity and divergence with other mammalian taxa; these remain to be explored in detail.

Abstract: Predictions of the model of van Schaik (1989) of female-bonding in primates are tested by systematically comparing the ecology, level of within-group contest competition for food (WGC), and patterns of social behaviour found in two contrasting baboon populations. Significant differences were found in food distribution (percentage of the diet from clumped sources), feeding supplant rates and grooming patterns. In accord with the model, the tendencies of females to affiliate and form coalitions with one another, and to be philopatric, were strongest where ecological conditions promoted WGC. Group fission in the population with strong WGC was “horizontal” with respect to female dominance rank, and associated with female-female aggression during a period of elevated feeding competition. In contrast, where WGC was low, females' grooming was focused on adult males rather than other females. Recent evidence suggests that group fission here is initiated by males, tends to result in the formation of one-male groups, and is not related to feeding competition but to male-male competition for mates. An ecological model of baboon social structure is presented which incorporates the effects of female-female competition, male-male competition, and predation pressure. The model potentially accounts for wide variability in group size, group structure and social relationships within the genus Papio. Socio-ecological convergence between common baboons and hamadryas baboons, however, may be limited in some respects by phylogenetic inertia.

Abstract: In many mammalian species, female success in raising offspring improves as they age. The residual reproductive value hypothesis predicts that each individual offspring will be more valuable to the mother as she ages because there is less conflict between the current and potential future offspring. Therefore, as mothers age, their investment into individual offspring should increase. Empirical evidence for an influence of declining residual reproductive value on maternal investment is unconvincing. Older mothers may not invest more, but may be more successful due to greater experience, allowing them to target their investment more appropriately (targeted reproductive effort hypothesis). Most studies do not preclude either hypothesis. Mare age significantly influenced maternal investment in feral horses living on the North Island of New Zealand. Older mares, that were more successful at raising foals, were more protective for the first 20 days of life, but less diligent thereafter. Total maternal input by older mothers did not seem to be any greater, but was better targeted at the most critical period for foal survival and a similar pattern was observed in mares that had lost a foal in the previous year. In addition, older mothers were more likely to foal in consecutive years, supporting the hypothesis that they are investing less than younger mares in individual offspring. Therefore, older mothers seem to become more successful by targeting their investment better due to experience, not by investing more in their offspring.