Abstract: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.

Abstract: Social learning is fundamental to social life across the animal kingdom, but we still know little about how natural selection has shaped social learning abilities on a proximate level. Sometimes, complex social learning phenomena can be entirely explained by Pavlovian processes that have little to do with the evolution of sociality. This implies that the ability to learn socially could be an exaptation, not an adaptation, to social life but not that social learning abilities have been left untouched by natural selection. I discuss new empirical evidence for associative learning in social information use, explain how natural selection might facilitate the associative learning process and discuss why such studies are changing the way that we think about social learning.

Abstract: Mortality is high in zoo-housed black rhinoceros (Diceros bicornis), and the reproductive rates of captive white rhinoceros (Ceratotherium simum) are unsustainably low. To determine the possible role of stress in the causation of these problems, we analyzed weekly fecal samples collected for 1 year from black (10 males and 16 females) and white (six males and 13 females) rhinoceroses at 16 zoos for corticoid metabolite concentrations. Fecal corticoid profiles were examined in relation to behavior as rated by keepers in a questionnaire, luteal phase ovarian cycles of females (Brown et al., 2001), and socioenvironmental factors. We compared individual fecal corticoid profiles by examining hormone means and variability (i.e., standard deviation (SD) and coefficient of variation (CV)). For the black rhinos, higher mean corticoid concentrations were found at zoos where rhinos were maintained in enclosures that were exposed to the public around a greater portion of the perimeter. Higher variability in corticoid excretion was correlated with higher rates of fighting between breeding partners and higher institutional mortality rates. Black rhino pairs that were kept separated exhibited lower corticoid variability and less fighting activity when they were introduced during female estrous periods compared to pairs that were kept together every day. For white rhinos, significantly lower mean corticoids were found for individuals that rated higher on “friendliness to keeper.” Higher corticoid variability was found in noncycling as compared to cycling white rhino females. Noncycling females exhibited higher rates of stereotypic pacing and lower frequencies of olfactory behaviors. Interindividual differences in mean corticoids in both species appeared to be related to responsiveness to humans, whereas corticoid variability was related to intraspecific social relationships. More importantly, high corticoid variability appeared to be an indicator of chronic or “bad” stress, because of its association with potentially deleterious consequences in each species (i.e., fighting and mortality (black rhino), and reproductive acyclicity (white rhino)). Our results provide evidence that social stressors may cause chronic stress in black and white rhinos, and that this contributes to the captive-population sustainability problems observed in each species. Zoo Biol 0:1–18, 2005. © 2005 Wiley-Liss, Inc.

Abstract: The Przewalski's horse (Equus caballus przewalskii) was extinct in the wild by the mid 1960s. The species has survived because of captive breeding only. The Takhin Tal reintroduction project is run by the International Takhi Group; it is one of two projects reintroducing horses to the wild in Mongolia. In 1997 the first harem group was released. The first foals were successfully raised in the wild in 1999. Currently, 63 Przewalski's horses live in Takhin Tal. Little information exists on causes of mortality before the implementation of a disease-monitoring program in 1998. Since 1999, all dead horses recovered (n = 28) have been examined and samples collected and submitted for further investigation. Equine piroplasmosis, a tick-transmitted disease caused by Babesia caballi or Theileria equi, is endemic in Takhin Tal and was identified as the cause of death of four stallions and one stillborn foal. In December 2000, wolf predation was implicated in the loss of several Przewalski's horses. However, thorough clinical, pathologic, and bacteriologic investigations performed on dead and surviving horses of this group revealed lesions compatible with strangles. The extreme Mongolian winter of 2000-2001 is thought to have most probably weakened the horses, making them more susceptible to opportunistic infection and subsequent wolf predation. Other occasional causes of death since 1999 were trauma, exhaustion, wasting, urolithiasis, pneumonia, abortion, and stillbirth. The pathologic examination of the Przewalski's horses did not result in a definitive diagnosis in each case. Several disease factors were found to be important in the initial phase of the reintroduction, which could potentially jeopardize the establishment of a self-sustaining population.

Abstract: Relative brain size (measured as gross brain size after body size effects are removed) differs systematically between families of rodents, insectivores and lagomorphs. The Sciuridae have the largest relative brain size, the Soricidae and Bathyergidae the smallest. These results are discussed and compared with previous analyses of relative brain sizes among primates and bats. These differences complicate comparisons between relative brain size across phylogenetically diverse species and attempts to relate differences in relative brain size to ecological variables. To overcome these problems, best fit relationships were estimated for each family, and values for each genus were expressed as deviations from the lines of best fit. We refer to these values as Comparative Brain Size (CBS). Differences in CBS are related to differences in habitat type (forest-dwelling genera have larger CBS' than grassland forms), in diet (folivores have smaller CBS' than generalists or insectivores, frugivores and granivores), in zonation (arboreal genera have larger CBS' than terrestrial ones) and in activity timing (nocturnal genera have larger CBS' than dirurnal ones). However, these ecological categories are interrelated and, when the effects of other ecological differences are taken into account using analyses of variance, only the differences associated with diet, and possibly habitat remain.