Abstract: Predator odours and habitat structure are thought to influence the behaviour of small mammalian prey, which use them as cues to reduce risks of predation. We tested this general hypothesis for house mice, Mus domesticus, by manipulating fox odour density via addition of fox scats and habitat via patchy mowing of vegetation, for populations in 15 x 15-m field enclosures. Using giving-up densities (GUDs), the density of food remaining when an animal quits harvesting a patch, we measured foraging behaviours in response to these treatments. Mice consistently avoided open areas, leaving GUDs two to four times greater in these areas than in densely vegetated patches. However, mouse GUDs did not change in response to the addition of fox scats, even immediately after fresh scats were added. There was no interaction between fox odour and habitat use. We then tested whether habituation to fox odours had occurred, by comparing the individual responses to scats of eight mice born into enclosures with fox scats to those of eight mice born into scat-free enclosures and five wild mice. In smaller enclosures, GUDs of trays with scats did not differ from GUDs of trays without scats for any treatment. We conclude that exposure to high levels of fox odours did not alter the foraging behaviour of mice, but that mice did reduce foraging in areas where habitat was removed, perceiving predation risk to be greater in these areas than controls. We suggest further that studies using the `scat-at-trap' technique, which have shown avoidance of predator odours by mice and other small mammals, may overestimate the general avoidance of predator odours by free-living prey, which must forage with a constant background of predator odours.

Abstract: Flehmen behaviour in Welsh pony (Equus caballus) mares and foals living on pasture was observed during 807 h of focal sampling. A series of flehmens performed at one site was defined as a flehmen incident. Colts exhibited flehmen incidents and performed flehmen more frequently during an incident than did fillies or mares. Filies exhibited flehmen incidents more frequently than did mares, but did not flehmen more frequently during an incident. Colts exhibited a peak frequency of performing flehmen and of flehmen incidents during weeks 1-4 with a subsequent linear decrease in frequency up to weeks 17-20. Usually, flehmen occurred without the subject having had direct contact of the nostrils, lips, or tongue with a possible stimulant. Twenty-six per cent of the flehmen incidents occurred during or after urination by another pony. Seven per cent of the incidents occurred during or after urination by the pony showing flehmen.

Abstract: Breeding groups with multiple stallions occur sympatrically with single-stallion breeding groups in feral horse, Equus caballus, populations. Mutualism and reciprocal altruism between stallions have been proposed to explain the origin and functioning of multistallion bands. However, empirical support for these hypotheses is contradictory and incomplete. Furthermore, there are no explicit tests of the predictions that each hypothesis makes about stallion behaviour and social structure. We compared nine multistallion and 18 single-stallion bands in the Kaimanawa Ranges, New Zealand. Compared with agonistic behaviours, affiliative behaviours were relatively unimportant in the relationships between stallions within bands. The number of stallions in the band did not have a positive influence on mare group size, stability, home range quality or reproductive success in bands. Furthermore, there was a positive relationship between aggression ('intolerance') by the dominant towards subordinate stallions and the subordinates' effort in mare group defence ('helping') but a negative relationship between helping effort by subordinates and their proximity to, and mating with, the bands' mares. Therefore, the predictions of the mutualism and reciprocal altruism hypotheses were not supported. Indeed, for some of the predictions we found the opposite outcomes to be true. Multistallion bands had significantly poorer reproductive success, and dominant stallions were less tolerant of subordinates that helped most and reduced their access to mares. Nevertheless, in all other respects Kaimanawa stallions in multistallion bands behaved like those described elsewhere. Thus, we reject cooperative hypotheses for multimale breeding groups in horses and discuss the mate parasitism and consort hypotheses as better alternatives. Copyright 2000 The Association for the Study of Animal Behaviour.

Abstract: Females often aggregate at particular sites for feeding or shelter, thus giving adult males the opportunity to defend harems and increase male reproductive success. Rival males compete for control of harems via ritualized displays or direct combat using weaponry. Contests for harems or the resources required by females can be settled based on asymmetries in fighting ability or resource ownership. Males that accurately assess a rival's fighting ability prior to engaging in potentially costly combat should be favoured by selection. Game theory and optimality models provide three models to explain how individuals decide to persist in or flee from a fight. These models are the energetic war of attrition, the sequential assessment model and the cumulative assessment model. Using staged contests in the laboratory, I tested predictions of these models using the Wellington tree weta, Hemideina crassidens, a sexually dimorphic insect native to New Zealand. Male H. crassidens use their enlarged mandibles as weapons in fights for access to adult females that reside in cavities in trees. My results supported a prediction common to each assessment model: contest duration was negatively correlated with the asymmetry in opponent's weapon size. The sequential assessment model of contest settlement was partially supported but the strongest support was for the cumulative assessment model. Predictions of the latter model were supported because: (1) fights are apparently settled based on own-size assessment; (2) fights occur in a single phase and escalate; and (3) contests involve physical combat and injury. I suggest that, in nocturnal species, cumulative assessment will generally be most applicable.

Abstract: A study of a herd of Camargue horsesEquus caballus, showed that while the majority of high-ranking stallions held single-male harems, some sons of low-ranking mares, being low ranking themselves, formed alliances that could last a lifetime. The two stallions were each other's closest associate and preferential grooming partner. Alliances were based on coalitions in which either both partners confronted an intruder synchronously or the dominant of the pair tended the female(s) while the subordinate simultaneously displayed towards the rival. Alliance partners were of similar age but were not more closely related to each other than to other stallions in the herd. Long-term paternity data revealed that subordinates sired close to a quarter of the foals born into the alliance group, and significantly more foals than low-ranking stallions in the herd adopting a `sneak'-mating strategy. The dominant appeared to benefit from the presence of his subordinate partner. Fights occurred all year round, and the subordinate stallion of each alliance pair fought outside competitors more than twice as often as the dominant. Forming short-term alliances before defending mares on their own may enhance long-term reproductive success for both partners. Other benefits to both partners include higher survivorship of their foals and increased access to proven reproductive mares. These results suggest that the relationship between alliance partners is based on mutualism, but several conditions for reciprocity seem to be fulfilled: the benefit to the dominant (assistance in fights), and the benefit to the subordinate (access to reproduction), are both costly to the other partner and delayed in time.