Abstract: Feral pony mares, Equus caballus, at Assateague Island, Maryland, formed linear dominance hierarchies within bands. Generally, older mares dominated younger mares, and larger mares dominated smaller mares. Large mares initiated aggression more often than small mares when age was controlled for but, surprisingly, older mares initiated aggression less often than younger mares when size was controlled for. Thus, mares peak in aggressiveness fairly soon after achieving full size and then, while maintaining or improving their rank in the domainance hierarchy, progressively reduce their involvement in aggression as they grow older, Involvement in aggression per mare increased as number of mares in the group increased; this effect was independent of nearest-mare distances. Frequency of involvement in aggression did not differ between mares that had changed bands within the year and mares whose band association had continued for a year or more. Aggression was directed more frequently than expected at subordinate mares who were nursing, and also occurred more frequently than expected at water holes. The proportion of aggressive encounters during grazing closely matched the total proportion of time spent grazing. Subordinate mares with foals received aggression more often than subordinate mares without foals. The high frequency of aggression associated with foals and nursing suggests that interference with reproduction of subordiantes is an important mode of competition between mares. Such interference may be common in animals that feed on dispersed resources and live in small, cohesive groups.

Abstract: In a wide range of animal species, males coerce females to mate with them, either by physically forcing them to mate, by harassing them until they mate or by punishing persistent refusal to mate. The first section of this paper argues that the possibility of forced copulation can generate arms races between males and females that may have substantial costs to both sexes. In the second section, it is suggested that sexual harassment commonly represents a `war of attrition' between the sexes; existing game theory models that may apply to sexual conflict over mating decisions are reviewed. The third section develops a simple prospective model for the evolution of intimidation by punishment in situations where males can raise the probability that females will accept their advances in future by punishing them for refusal to mate. Where the benefits of sexual coercion to males are high, all three male strategies may develop to a point where they have substantial costs to females. In the final section, evidence that female behaviour is adapted to minimizing these costs is reviewed.

Abstract: In many cooperatively breeding species, group members form a dominance hierarchy or queue to inherit the position of breeder. Models aimed at understanding individual variation in helping behavior, however, rarely take into account the effect of dominance rank on expected future reproductive success and thus the potential direct fitness costs of helping. Here we develop a kin-selection model of helping behavior in multimember groups in which only the highest ranking individual breeds. Each group member can invest in the dominant's offspring at a cost to its own survivorship. The model predicts that lower ranked subordinates, who have a smaller probability of inheriting the group, should work harder than higher ranked subordinates. This prediction holds regardless of whether the intrinsic mortality rate of subordinates increases or decreases with rank. The prediction does not necessarily hold, however, where the costs of helping are higher for lower ranked individuals: a situation that may be common in vertebrates. The model makes two further testable predictions: that the helping effort of an individual of given rank should be lower in larger groups, and the reproductive success of dominants should be greater where group members are more closely related. Empirical evidence for these predictions is discussed. We argue that the effects of rank on stable helping effort may explain why attempts to correlate individual helping effort with relatedness in cooperatively breeding species have met with limited success.

Abstract: During 1992 and 1993, 14 reintroduced Przewalski Horses or Takhi (Equus ferus przewalskii) were studied in the Hustain Nuruu Mountain Steppe reserve in Mongolia. Most of the individuals did not know each other before reintroduction. These Takhi were the first of five groups due to be released in the reserve after an acclimatisation period of at least 1 year. During acclimatisation the Takhi, lived visually and acoustically separately, in fenced enclosures of approximately 45 ha each. The observations, mostly scan-sampling, were carried out in each season. The observation bouts were divided over six periods and over two harem herds. Two of the periods were in the same consecutive seasons, so comparison over the years was possible. Social integration within the Takhi herds was very high from the beginning, as described by the spatial relation and synchronisation data. Between 50 and 89% of the observation time, the behaviour of all herd members was synchronised. The amount of time spent grazing by the Takhi (30-68% of the daylight period) was similar to that of feral horses and Takhi in captivity and semi-reserves. The Takhi tended to rest in the morning and have a bimodal period of grazing at dawn and in the afternoon. The Takhi displayed clear habitat preferences for certain activities. They had a strong preference to rest at the highest point in their enclosure. They fed preferably on two or three different vegetation types (with five types available in each enclosure). The amount of time spent grazing during the non-growing seasons (49 +/- 15%) indicates that the feeding value and availability of food were sufficient. Health changes were detected adequately using condition scoring sheets. No supplementary food or water was supplied during the harsh winters. Moreover, low mortality rates and high reproductive success show that the mountain steppe is a habitat which is potentially suitable for establishing a healthy Takhi population. Takhi is the first species to return to its native habitat after living only in zoos for so many generations.

Abstract: A short-term study was made of 2 groups of Przewalski horses, a bachelor group of 4 juvenile stallions in Ooij Polder and a harem group of 1 stallion and 4 mares. All social interactions were recorded and the nearest and farthest neighbour was noted. Correspondence analysis was used to determine what parameters determined the relationships among the horses. There was a linear hierarchy among the bachelor stallions. The dominant stallion of the group was also the oldest. The hierarchy was not linear in the harem group, and the 3-year-old stallion was subordinate to the 5-year-old mares. He was also most likely to be farthest from other horses. The mares of the same age, who had also arrived in the park at the same time, tended to be one another's nearest neighbours. The frequency of aggression is higher among Przewalski horses than among domestic horses of similar ages. Correspondence analysis revealed that head-threats and other forms of aggression accounted for more of the variance in the data than any other behaviour, but submission, play and social interactions also contributed.