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: 1. The niche separation of three species of alcelaphine antelope (wildebeest, topi and hartebeest) with similar body size was compared by measuring bite weight, bite rate, intake rate and selectivity of tame animals in plots containing grass at different growth stages. 2. On growing swards, hartebeest had a smaller bite weight and lower intake rate, and were also less selective of green leaf, than either topi or wildebeest. On senescent swards, hartebeest were more selective of leaf than the other two species. 3. Wildebeest had a faster bite rate than either topi or hartebeest on swards with low biomass and high protein content of green leaf (green flush). Bite weight and intake rate of wildebeest and topi were similar despite the difference in breadth of their incisor rows. 4. Topi were significantly more selective of green leaf than the other two species and were the only species to maintain a rapid bite rate on swards with high green leaf biomass. 5. The feeding experiments did not reveal significant cross-overs between species in the rate of food intake on different grass types, but each species was most proficient either in leaf selection or bite rate when feeding on grass swards in a particular growth stage. We suggest that growth stage is a primary determinant of niche separation. 6. In Serengeti, grazing ungulates which migrate are specialists of the earlier growth stages of grass which tend to be transient, while those that are residential specialize on late growth stages which are more enduring. The mobility of species, and the spatial and temporal dynamics of pastures containing different growth stages of grass, contribute to niche separation.

Abstract: During contests animals typically exchange information about fighting ability. Among feral horses these signals involve olfactory or acoustical elements and each type can effectively terminate contests before physical contact becomes necessary. Dung transplant experiments show that for stallions, irrespective of rank, olfactory signals such as dung sniffing encode information about familiarity suggesting that such signals can be used as signatures. As such they can provide indirect information about fighting ability as long as opponents associate identity with past performance. Play-back experiments, however, show that vocalizations, such as squeals, directly provide information about status regardless of stallion familiarity. Sonographs reveal that squeals of dominants are longer than those of subordinates and that only those of dominants have at their onset high-frequency components.

Abstract: 1. The role of lion Panthera leo predation in the dynamics of blue wildebeest Connochaetes taurinus and zebra Equus burchelli populations was investigated through simulation models. The data used in the models were from intensive observations over 4 years in the south-east of the Kruger National Park. 2. Population estimates of wildebeest and zebra were made from aerial surveys, sex and age ratios from ground counts. Lion numbers were determined from observations of marked and radio-collared animals. Predation was studied by following lions for continuous periods of up to 336 h. 3. Two models were constructed. Model 1 ascertained the number of killing lions (adult females) that could be supported by each prey population while remaining stable. A single model was constructed for the sedentary wildebeest population. A summer and winter model was constructed for the semi-migratory zebra population. The sensitivity of the parameters in the model was tested by changing their value by 10%. In model 2, the kill age structure for each species was changed to determine the number of killing lions the altered prey selection parameters could support. 4. There was no difference in the vulnerability of either species to predation. Zebra foals (<1 year) were killed more frequently than expected. No selection for sex or by season could be found for either species. 5. Model 1 predicted that the wildebeest population stabilizes with 7.7 killing lions, close to the number in the study area. The winter zebra population stabilizes with 6.8 killing lions and the summer zebra population with 19.4. Manipulation of kill rate followed by adult fecundity rate had the greatest effect on population size of both species. In model 2, wildebeest predation was made selective towards calves and zebra predation was made non-selective for sex and age. With these parameters the wildebeest population stabilizes with 10.7 killing lions and the zebra population with 5.4 in winter and 15.1 in summer. 6. The models suggest that lion predation affected wildebeest more severely than zebra during the study. This was through the way in which lions selected their prey, and because of the sedentary behaviour of the wildebeest, as opposed to the semi-migratory behaviour of the zebra.

Abstract: Summary The introduction of game-theoretical thinking into evolutionary biology has laid the groundwork for a heuristic view of animal behaviour in which individuals employ “strategies” – rules that instruct them how to behave in a given circumstance to maximize relative fitness. Axelrod and Hamilton (1981) found that a strategy called Tit-For-Tat (TFT) is one robust cooperative solution to the iterated Prisoner's Dilemma game. There exists, however, little empirical evidence that animals employ TFT. Predator inspection in fish provides one ecological context in which to examine the use of the TFT strategy.