Abstract: According to the socioecological framework, transitivity (or linearity) in dominance relationships is related to competition over critical resources. When a population is structured into groups, the intensity of between- versus within-group competition influences the form and function of its social organization. Few studies have compared the type and relative intensity of competition at these two levels. African elephants have well-structured social relations, providing an exemplary system for such a study. We report on dominance hierarchies among free-ranging elephants and evaluate the factors that drive their socioecological structure to lie in a region of the three-dimensional nepotism/despotism/tolerance space rarely observed among social species; namely, where non-nepotistic, transitive dominance hierarchies within groups emerge despite kin-based philopatry and infrequent agonistic interactions over widely distributed resources. We found significant transitivity in dominance hierarchies between groups. Dominance relations among the matriarchs of different social groups were primarily age based, rather than driven by physical or group size, and group matriarch rank influenced the dominance relationships among nonmatriarchal females in the population. Our results suggest that between-group dominance relationships induce tolerance among group members, which in combination with high group relatedness, reduces the benefits of nepotism. We postulate that cognitive abilities and high risk of injury in contests enhance winner and loser effects, facilitating the formation of transitive dominance relationships, despite widely distributed resources over which infrequent competition occurs. The interplay of cognitive abilities, winner and loser effects, resource distribution, and within- and between-group dominance relationships may produce behaviour in other strongly social mammals that differs from that predicted by a superficial application of current socioecological models.

Abstract: Individual humans, and members of diverse other species, show consistent differences in
aggressiveness, shyness, sociability and activity. Such intraspecific differences in
behaviour have been widely assumed to be non-adaptive variation surrounding
(possibly) adaptive population-average behaviour. Nevertheless, in keeping with recent
calls to apply Darwinian reasoning to ever-finer scales of biological variation, we sketch
the fundamentals of an adaptive theory of consistent individual differences in behaviour.
Our thesis is based on the notion that such .personality differences. can be selected for if
fitness payoffs are dependent on both the frequencies with which competing strategies
are played and an individual`s behavioural history. To this end, we review existing models
that illustrate this and propose a game theoretic approach to analyzing personality
differences that is both dynamic and state-dependent. Our motivation is to provide
insights into the evolution and maintenance of an apparently common animal trait:
personality, which has far reaching ecological and evolutionary implications.

Abstract: Predictions of the model of van Schaik (1989) of female-bonding in primates are tested by systematically comparing the ecology, level of within-group contest competition for food (WGC), and patterns of social behaviour found in two contrasting baboon populations. Significant differences were found in food distribution (percentage of the diet from clumped sources), feeding supplant rates and grooming patterns. In accord with the model, the tendencies of females to affiliate and form coalitions with one another, and to be philopatric, were strongest where ecological conditions promoted WGC. Group fission in the population with strong WGC was “horizontal” with respect to female dominance rank, and associated with female-female aggression during a period of elevated feeding competition. In contrast, where WGC was low, females' grooming was focused on adult males rather than other females. Recent evidence suggests that group fission here is initiated by males, tends to result in the formation of one-male groups, and is not related to feeding competition but to male-male competition for mates. An ecological model of baboon social structure is presented which incorporates the effects of female-female competition, male-male competition, and predation pressure. The model potentially accounts for wide variability in group size, group structure and social relationships within the genus Papio. Socio-ecological convergence between common baboons and hamadryas baboons, however, may be limited in some respects by phylogenetic inertia.

Abstract: Socio-ecological explanations for intra- and interspecific variation in the social and spatial organization of animals predominate in the scientific literature. The socio-ecological model, developed first for the Bovidae and Cervidae, is commonly applied more widely to other groups including the Equidae. Intraspecific comparisons are particularly valuable because they allow the role of environment and demography on social and spatial organization to be understood while controlling for phylogeny or morphology which confound interspecific comparisons. Feral horse (Equus caballus Linnaeus 1758) populations with different demography inhabit a range of environments throughout the world. I use 56 reports to obtain 23 measures or characteristics of the behaviour and the social and spatial organization of 19 feral horse populations in which the environment, demography, management, research effort and sample size are also described. Comparison shows that different populations had remarkably similar social and spatial organization and that group sizes and composition, and home range sizes varied as much within as between populations. I assess the few exceptions to uniformity and conclude that they are due to the attributes of the studies themselves, particularly to poor definition of terms and inadequate empiricism, rather than to the environment or demography per se. Interspecific comparisons show that equid species adhere to their different social and spatial organizations despite similarities in their environments and even when species are sympatric. Furthermore, equid male territoriality has been ill-defined in previous studies, observations presented as evidence of territoriality are also found in non-territorial equids, and populations of supposedly territorial species demonstrate female defence polygyny. Thus, territoriality may not be a useful categorization in the Equidae. Moreover, although equid socio-ecologists have relied on the socio-ecological model derived from the extremely diverse Bovidae and Cervidae for explanations of variation in equine society, the homomorphic, but large and polygynous, and monogeneric Equidae do not support previous socio-ecological explanations for relationships between body size, mating system and sexual dimorphism in ungulates. Consequently, in spite of the efforts of numerous authors during the past two decades, functional explanations of apparent differences in feral horse and equid social and spatial organization and behaviour based on assumptions of their current utility in the environmental or demographic context remain unconvincing. Nevertheless, differences in social cohesion between species that are insensitive to intra- and interspecific variation in habitat and predation pressure warrant explanation. Thus, I propose alternative avenues of inquiry including testing for species-specific differences in inter-individual aggression and investigating the role of phylogenetic constraints in equine society. The Equidae are evidence of the relative importance of phylogeny and biological structure, and unimportance of the present-day environment, in animal behaviour and social and spatial organization.

Abstract: The status of the Gobi khulan Equus hemionus subsp. is recorded as “insufficiently known” in the Species Survival Commission's Equid Action Plan. Recent counts confirm that Mongolia holds the most important population of the whole species. Since 1953, the animals have benefited from a protected status, but this is now challenged. A 5-year study in the B part of the Gobi National Park on one subpopulation showed that it has remained stable over the past 15 years with an adequate mean reproductive rate of 15% and a 50% survival rate over the first year. Age/sex related mortality and prey analysis indicate that wolf predation probably has some impact on the population, in particular for 4-6-year-olds of both sexes at the start of reproduction. Desert and mountain steppes are the khulan's year-round preferred habitat, but `oases', play an important role at the beginning of lactation. Anthropogenic factors affect both home range and habitat use through direct intervention or permanent occupation of the scarce water sources. Khulans of this subpopulation, unlike other Asian and African wild asses, form year-round stable, non-territorial families. These families and all-male groups join together into “bands” in winter, and herds of several hundred animals, where reproductive rate is highest, form throughout the year. The existence of such herds may thus be critical for the breeding success of the population. Our study provides the first detailed quantitative data for this subspecies, which will help to monitor changes in the future.