Abstract: Data were collected on 972 aggressions and 233 acts of mutual grooming in a herd of 9 Przewalski horses in the Hellabrunn Tierpark in Munich, West Germany. The herd was composed of 1 adult stallion, 5 adult mares and 3 foals. A distinct linear dominance hierarchy was present in the herd, with the stallion being the top-ranking animal. Age was significantly correlated with rank. Almost 40% of all aggressions consisted of herding actions by the stallion. Threats to bite (20% of all aggressions) and threats to kick (11.4%) were next in frequency of occurrence. Most mutual grooms (71%) involved grooming the front part of the body. Although mutual grooming may be used to appease higher-ranked animals, most grooming bouts were between related horses. Foals initiate 47.6% of all allogrooming. Mutual grooming may reduce weaning conflict between a mare and her foal or may result in female coalitions that defend against predators or aggression by the herd stallion.

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: New Caledonian crows, Corvus moneduloides, are the most advanced avian tool makers and tool users. We previously reported that captive-bred isolated New Caledonian crows spontaneously use twig tools and cut tools out of Pandanus spp. tree leaves, an activity possibly under cultural influence in the wild. However, what aspects of these behaviours are inherited and how they interact with individual and social experience remained unknown. To examine the interaction between inherited traits, individual learning and social transmission, we observed the ontogeny of twig tool use in hand-reared juveniles. Successful food retrieval was preceded by stereotyped object manipulation action patterns that resembled components of the mature behaviour, demonstrating that tool-oriented behaviours in this species are an evolved specialization. However, there was also an effect of social learning: juveniles that had received demonstrations of twig tool use by their human foster parent showed higher levels of handling and insertion of twigs than did their naive counterparts; a choice experiment showed that they preferred to handle objects that they had seen being manipulated by their human foster parent. Our observations are consistent with the hypothesis that individual learning, cultural transmission and creative problem solving all contribute to the acquisition of the tool-oriented behaviours in the wild, but inherited species-typical action patterns have a greater role than has been recognized.

Abstract: The present investigation was undertaken to study the proximate reasons why and the behavioral patterns of young male Misaki feral horses when they left their natal band or mothers. We observed a total of ten young males twice a month from January 1988 to December 1995. Almost all young males left their natal band or mothers at between 1 and 4 years of age. We found that, during the separation process, all the young males from first parity dams returned several times after the initial separation, indicating a strong attachment between primiparous mares and their male offspring. The other five separated only once without rejoining. Our observations showed five variable behavior patterns of young males at separation time, depending on the consort relation between their mothers and harem stallion and the reason for separation at that time. Eight young males separated in the non-breeding season at average 2.1 years and the other two separated in the breeding season at average 3 years and the average difference was not significant. These results revealed that 80% of the young males separated voluntarily when the natural resources become poor whereas 20% separated when their siblings were born.

Abstract: Przewalski horses (Equus ferus przewalskii), also known as takhi, were first re-introduced to the wild in Hustai National Park, Mongolia, in 1994. Since then the number of free harems increased to a maximum of seven; there are currently six (October 2000). The size of the home range of each of the harems changed among years and among seasons. The horses tended to settle in a home range close to where they were released although they explored the surrounding area. The use of the habitat within each home range changed through the day, with the horses grazing in the valleys during the morning and evening, and moving to higher places to stand rest and use as a refuge from heat and flies during the middle of the day. Range establishment and area, as well as habitat use are discussed.