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: We report morphological data on brains of four African, Loxodonta africana, and three Asian elephants, Elephas maximus, and compare findings to literature. Brains exhibit a gyral pattern more complex and with more numerous gyri than in primates, humans included, and in carnivores, but less complex than in cetaceans. Cerebral frontal, parietal, temporal, limbic, and insular lobes are well developed, whereas the occipital lobe is relatively small. The insula is not as opercularized as in man. The temporal lobe is disproportionately large and expands laterally. Humans and elephants have three parallel temporal gyri: superior, middle, and inferior. Hippocampal sizes in elephants and humans are comparable, but proportionally smaller in elephant. A possible carotid rete was observed at the base of the brain. Brain size appears to be related to body size, ecology, sociality, and longevity. Elephant adult brain averages 4783 g, the largest among living and extinct terrestrial mammals; elephant neonate brain averages 50% of its adult brain weight (25% in humans). Cerebellar weight averages 18.6% of brain (1.8 times larger than in humans). During evolution, encephalization quotient has increased by 10-fold (0.2 for extinct Moeritherium, approximately 2.0 for extant elephants). We present 20 figures of the elephant brain, 16 of which contain new material. Similarities between human and elephant brains could be due to convergent evolution; both display mosaic characters and are highly derived mammals. Humans and elephants use and make tools and show a range of complex learning skills and behaviors. In elephants, the large amount of cerebral cortex, especially in the temporal lobe, and the well-developed olfactory system, structures associated with complex learning and behavioral functions in humans, may provide the substrate for such complex skills and behavior.

Abstract: The behavior of feral populations of the African wild ass (Equus africanus) were studied in the Northern Panamint Range of Death Valley National Monument for 20 months from 1970 to 1973 [Moehlman, P.D., 1974. Behavior and ecology of feral asses (Equus asinus). PhD dissertation, University of Wisconsin, Madison, 251 pp.; Moehlman, P.D., 1979. Behavior and ecology of feral asses (Equus asinus). Natl. Geogr. Soc. Res. Reports, 1970: 405-411]. Maintenance behavior is described and behavior sequences that were used in social interactions are quantified by sex and age class. Agonistic, sexual, and greeting behavior patterns are described and analyzed in conjunction with the responses they elicited. Mutual grooming mainly occurred between adult males, and between females and their offspring. Five types of vocalizations were distinguished: brays, grunts, growls, snorts, and whuffles. A second population was studied for 1 month on Ossabaw Island, GA (Moehlman, 1979). This population had more permanent social groups and had a higher rate of mutual grooming and foal social play.

Abstract: Feral asses were studied on Volcano Alcedo, Galapagos Islands, Ecuador, during the wet season of 1980. On the volcano rim during March/April, two stable groups were observed to have a `female (harem) defense' polygynous mating system [Emlen, S.T., Oring, S.W., 1977. Ecology, sexual selection, and the evolution of mating systems. Science 197 (4300), pp. 215-223] and social behavior patterns and feeding ecology similar to feral asses living in a habitat where forage and climate are similar, e.g., Ossabaw Island, Georgia [Moehlman, P.D., 1979. Behavior and ecology of feral asses (Equus asinus). Nat. Geogr. Soc. Res. Rep., 1970, pp. 405-411; Moehlman, P.D., 1997. Feral asses (Equus africanus): intraspecific variation in social organization in arid and mesic habitats. J. Appl. Anim. Behav. Sci., this issue; McCort, W.D., 1980. The feral asses (Equus asinus) of Ossabaw Island, Georgia., PhD Dissertation, Pennsylvania State University, University Park, 219 pp.].

Abstract: An ethogram of agonistic and related behaviors among equid bachelor band members was developed. Several key English-language studies on equids were reviewed to derive a preliminary inventory of specific behaviors to be included in the ethogram. A bachelor band of domestic pony stallions pastured together was observed for approximately 50 daylight hours to obtain detailed descriptions of each behavior, enable photographic and video documentation of behaviors, and identify any behaviors to be added to the preliminary inventory. An initial draft of the ethogram was sent to 65 equine researchers for review. Twenty-eight critical reviews were received and their suggestions considered for the final draft. A total of 49 elemental behaviors including five distinct vocalizations was included in the ethogram. Three complex behavioral sequences were also included. Most of the behaviors catalogued from the direct observation of pastured pony stallions were also found in the equid literature. For many, references to these behaviors specifically among males or bachelor band members were not found. The results offer a practical tool for quantitative research and other studies of equid inter-male behavior as well as for teaching of equid behavior, and should facilitate progress toward development of a complete ethogram for the horse and other equids.