Abstract: For the purpose of distinguishing between mutualism and reciprocity in nature, recent work on the evolution of cooperation has both oversimplifed and undersimplified the distinction between these two categories of cooperation. This article addresses the resulting issues of model testability, clarifies the role of time and argues that the category of `pseudo-reciprocity' is an unnecessary complication.

Abstract: An experimental analysis of the movements of predator-approaching fish is presented. The experiments evaluated two competing hypotheses. (1) Predator-approaching fish play the game-theoretical strategy Tit for Tat. Alternatively, (2) the movements of predator-approaching fish superficially resemble Tit for Tat, because fish independently orient to a predator and simultaneously attempt to stay close together. Experimental subjects were mosquito fish,Gambusia affinisapproaching a green sunfish,Lepomis cyanellusTwo experiments were performed. Experiment 1 replicated results of Milinski (1987) and Dugatkin (1991), showing thatGambusiacome closer to a visible predator when a mirror is oriented parallel to their direction of travel. Experiment 2 attempted to separate the effects of common orientation and social cohesion in accounting for the frequency of Tit-for-Tat-like motions in pairs of predator-approachingGambusia. Results of experiment 2 suggest that a simple additive combination of the effects of (1) social cohesion in the absence of a visible predator and (2) orientation to a visible predator in the absence of a visible companion can account for the observed frequency of Tit-for-Tat-like motions for pairs of predator-approachingGambusia. It is concluded that predator approach in shoaling fishes is probably a simple by-product mutualism, rather than cooperation maintained by reciprocity in a Prisoner's Dilemma.

Abstract: Lactating female baboons, Papio cynocephalusoften maintain close associations with particular males. There are at least three proposed benefits of 'friendships' to females: (1) male protection against potentially infanticidal males; (2) male protection against harassment by dominant females; (3) male attachment to an infant that develops into future care of juveniles. These hypotheses were examined in a population of chacma baboons, P. c. ursinusin which male infanticide accounted for at least 38% of infant mortality. Almost all mothers of young infants formed strong bonds with one or two males with whom they had copulated during the cycle in which they conceived their infants. Females were primarily responsible for maintaining friendships during lactation, but they terminated these relationships if their infants died. In playbacks of females' screams, male friends responded more strongly than control males. They also responded more strongly to the screams of female friends than to the screams of control females. Following an infant's death, however, male friends responded less strongly than control males to the same females' screams. Finally, male friends responded more strongly than control males to playback sequences in which female screams were combined with the threat vocalizations of a potentially infanticidal alpha male, but not when female screams were combined with the threat calls of a non-infanticidal male or the alpha female. Both observations and experiments suggest that the benefits of friendships to females derive from the protection of their infants against infanticide.1997The Association for the Study of Animal Behaviour

Abstract: Risk-sensitive foraging theory is based on the premise that unpredictable runs of good or bad luck can cause a variable food source to differ in fitness value from a fixed food source yielding the same average rate of gain but no unpredictability. Thus, risk-sensitive predictions are dependent on the food intake from variable sources being not only variable but also unpredictable or `risky' in outcome. This study tested whether unpredictability is a component of the value that foraging starlings,Sturnus vulgarisattribute to food sources that are variable in the delay to obtain food. Two groups of birds chose between a fixed and a variable delay option; the variable option was unpredictable in the risky group and predictable in the risk-free group in the overall rate of intake it yielded. In both groups the fixed option was adjusted by titration to quantify the magnitude of preference for predictable and unpredictable variance. On negative energy budgets both groups were significantly risk-prone, with the risky group being significantly more risk-prone than the risk-free group. Switching the birds to positive budgets by doubling the size of each food reward had no significant effect on preference, and similar trends to those found with negative budgets were observed. These results are not readily explained by risk-sensitive foraging theory, but may be explained by the algorithm used by the birds to attribute value to average expected rewards.