Abstract: The Marginal Value Theorem (MVT) describes the behaviour that maximizes the ratio of expected gain over expected foraging time in a patchy environment. When travel time is variable, the MVT rationale and its predictions are sensitive only to the mean travel time and not to the spread or skew of the distribution. Two mechanistic arguments contradict these predictions. First, tests of the MVT have previously shown that there is a disproportionate influence of the last travel time, and second, psychological models of information processing suggest that memory for time intervals is strongly dependent on the scatter of the distribution experienced. These mechanistic concepts, combined with Jensen's inequality, suggest that patch exploitation should decrease as the scatter of the travel distribution increases. In a Skinner box experiment with pigeons, Columba livia, the problem was examined by simulating three environments with identical patches and the same mean travel time, but different travel time variability. Patch exploitation decreased with increasing variance in travel time. The results are used to argue in favour of the inclusion of realistic psychological properties as constraints in functional models of behaviour. Although both the MVT and the mechanistic models account for some features of the results, none of them can explain all the findings.

Abstract: In animals, behavioural properties such as aggressive propensity are often consistent over a life span, and they may form part of a behavioural syndrome. We studied how aggressive propensity influences dominance, contest behaviour and growth in the cooperatively breeding cichlid fish Neolamprologus pulcher. We tested whether intrinsic aggressive propensity (1) influences dominance in paired contests, (2) causes different aggression levels in contests with partners matched for aggressive propensity compared to unmatched partners, and how it (3) affects growth rate in groups that were either matched or unmatched for aggressive propensity. Intrinsic aggressive propensity was first scored with a mirror test and classified as high, medium or low. Thereafter we tested fish with either high or low aggressive propensity with partners matched for size and either matched or unmatched for aggressive type in a paired contest for a shelter. We scored dominance, aggression and submission. As predicted, (1) dominance was more clearly established in unmatched than in matched contests and (2) individuals with high aggressive propensity launched more attacks overall than fish with low intrinsic aggressiveness, suggesting a higher propensity to escalate independently of winning or losing the paired contest. However, contrary to expectation, (3) individuals with low aggressiveness grew faster than aggressive ones in unmatched groups, whereas the opposite occurred in matched groups. This suggests that individuals with low aggressive propensity may benefit from conflict evasion, which might allow them to gain dominance in the future owing to larger body size.

Abstract: Newborn rabbits, Oryctolagus cuniculus, discriminate between different categories of adult conspecifics on the basis of their abdominal odour cues. Whether these cues can support the development of filial preferences has not been adequately tested. Using a two-choice paradigm, we assessed the ability of 3-8-day-old pups to orient selectively to the mother versus an unfamiliar female, either spontaneously or after odour conditioning. In experiment 1, nonconditioned pups roamed indifferently over the mother and an unfamiliar female. In experiment 2, pups conditioned to a neutral odorant while nursing or with the mammary pheromone became attracted by the odorant. In experiment 3, pups that had learned the odorant while nursing oriented for longer to any female carrying it, but the unscented mother and a scented unfamiliar female were equally attractive. Finally, in experiment 4, pups that had learned the odorant paired with the mammary pheromone showed a preference for their scented mother, but not systematically for a scented unfamiliar female; furthermore, they were equally attracted by the unscented mother and a scented unfamiliar female. In sum, pups did not spontaneously evince an olfactory preference for the mother when opposed to an unfamiliar female, although they seemed able to detect individual maternal odours. In fact, they appeared to react to both species-specific cues and individual cues that they had learned, and their responses depended on their degree of familiarity with the cues and on the context. The mammary pheromone by itself might act as both a releasing and a reinforcing signal in these early socially oriented behaviours.

Abstract: The function of olfactory signalling in social species is less well understood than in asocial species. Consequently, we examined olfactory communication in the ringtailed lemur, a socially complex primate that retains a functional vomeronasal organ, has well-developed scent glands and shows a suite of scent-marking behaviour. To assess the information content of different types of scent gland secretions, we decoupled olfactory cues from the visual and behavioural modalities with which scent marking is normally associated. We presented male and female subjects (signal receivers) with a series of choice tests between odours derived from conspecific donors (signal senders) varying by sex, age, social status and reproductive condition. We additionally examined the influence of the receivers' reproductive state and familiarity with the signaller. The reproductive condition, social status and familiarity of senders and receivers affected signal transmission; specifically, male receivers attended most to the odours of conspecifics in breeding condition and to the odours of familiar, dominant animals. By contrast, females varied their responses according to both their own reproductive state and that of the sender. Based on male and female patterns of countermarking, we suggest that scent marking serves a function in intergroup spacing and intrasexual competition for both sexes, as might be expected in a female-dominant species. By contrast, minimal female interest in male odours counters a female mate choice function for scent marking in this species. Nevertheless, scent marks are critical to male-male competition and, therefore, may be subject to sexual selection.

Abstract: Feral horse, Equus caballus, breeding groups, called bands, usually include one but sometimes up to five stallions. We found that mares were loyal to single-stallion (SS) or multistallion (MS) bands or were social dispersers (maverick mares, Mv). The spacing and social behaviour of mares and stallions in single- and multistallion bands was measured. Indices of mare well-being were also measured including activity budgets (feeding: MS>SS=Mv; resting: MS<SS=Mv), band and mare travel (MS>SS), maternal effort in maintaining contact with foals (MS=Mv>SS), parasite levels in faeces (MS>Mv>SS), body condition (MS=Mv<SS), fecundity (Mv<MS<SS) and offspring mortality (Mv<MS<SS). We present evidence suggesting that the poorer well-being of maverick mares and multistallion band mares results from greater harassment by stallions. Stallion and mare behaviour and poor reproductive success in multistallion bands were not consistent with explanations for the existence of such bands based on cooperation or alternative mating strategies. We suggest an alternative explanation. Stable relationships between mares and a single stallion may enhance reproductive success by reducing aggression between individuals. Therefore, we propose that there is strong selection pressure for stable, long-term stallion-mare relationships, called consort relationships. We propose the consort hypothesis, that multistallion bands are an artefact of selection for stable relationships that occasionally result in more than one such relationship forming, because mares solicit more than one stallion and stallion dominance changes during band formation. Copyright 1999 The Association for the Study of Animal Behaviour.