Abstract: 1. Dominance/subordinance is a relationship between two individuals in which one defers to the other in contest situations. Each such relationship represents an adaptive compromise for each individual in which the benefits and costs of giving in or not giving in are compared. Familiar associates in groups or neighbours on nearby territories may develop relatively stable dominant-subordinate relationships based on individual recognition. Although the aggressive aspects of dominance are usually emphasized, the less conspicuous actions of the subordinate individual are actually more important in maintaining a stable relationship. 2. In evolutionary terms, dominance essentially equals priority of access to resources in short supply. Usually the subordinate, who would probably lose in combat anyway, is better off to bide its time until better able to compete at another time or another place. Both individuals save time, energy, and the risk of injury by recognizing and abiding by an established dominant-subordinate relationship. 3. Dominance can be either absolute or predictably reversible in different locations or at different times. Of the various forms of dominance behaviour, rank hierarchies and territoriality represent the two extremes of absolute and relative dominance, respectively. A dominance hierarchy is the sum total of the adaptive compromises made between individuals in an aggregation or organized group. Many animals seem to be capable of both absolute and relative dominance, and within species-specific limits the balance may shift toward one or the other. High density, or a decrease in available resources, favours a shift from relative to absolute dominance. Some species may exhibit both simultaneously. Social mammals may have intra-group hierarchies and reciprocal territoriality between groups, while the males of lek species may exhibit 'polarized territoriality' by defending small individual territories, with the most dominant males holding the central territories where most of the mating takes place. 4. Territoriality is a form of space-related dominance. Most biologists agree that its most important function is to provide the territory holder with an assured supply of critical resources. Territoriality is selected for only when the individual's genetic fitness is increased because its increased access to resources outweighs the time, energy, and injury costs of territorial behaviour. 5. Territoriality was first defined narrowly as an area from which conspecifics are excluded by overt defence or advertisement. The definition has been variously expanded to include all more or less exclusive areas without regard to possible defence, and finally to include all areas in which the owner is dominant. I define territory as a fixed portion of an individual's or group's range in which it has priority of access to one or more critical resources over others who have priority elsewhere or at another time. This priority of access must be achieved through social interaction. 6. My definition excludes dominance over individual space and moving resources, and includes areas of exclusive use maintained by mutual avoidance. It differs from most other definitions in its explicit recognition of time as a territorial parameter and its rejection of exclusivity and overt defence as necessary components of territorial behaviour. There is an indivisible continuum of degrees of trespass onto territories, and functionally it is priority of access to resources that is important rather than exclusive occupancy. 7. There is a similarly indivisible continuum in the intensity of behaviour needed to achieve priority of access to resources. Deciding whether or not an exclusive area is defended leads to the pointless exercise of trying to decide which cues indicating the owner's presence are conspicuous enough to merit being called defence. Concentrating on overt defence emphasizes the aggressive aspects of territorial behaviour rather than the equally or more important submissive aspects such as passive avoidance.

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: Darwin argued that between-species differences in intelligence were differences of degree, not of kind. The contemporary ecological approach to animal cognition argues that animals have evolved species-specific and problem-specific processes to solve problems associated with their particular ecological niches: thus different species use different processes, and within a species, different processes are used to tackle problems involving different inputs. This approach contrasts both with Darwin's view and with the general process view, according to which the same central processes of learning and memory are used across an extensive range of problems involving very different inputs. We review evidence relevant to the claim that the learning and memory performance of non-human animals varies according to the nature of the stimuli involved. We first discuss the resource distribution hypothesis, olfactory learning-set formation, and the 'biological constraints' literature, but find no convincing support from these topics for the ecological account of cognition. We then discuss the claim that the performance of birds in spatial tasks of learning and memory is superior in species that depend heavily upon stored food compared to species that either show less dependence upon stored food or do not store food. If it could be shown that storing species enjoy a superiority specifically in spatial (and not non-spatial) tasks, this would argue that spatial tasks are indeed solved using different processes from those used in non-spatial tasks. Our review of this literature does not find a consistent superiority of storing over non-storing birds in spatial tasks, and, in particular, no evidence of enhanced superiority of storing species when the task demands are increased, by, for example, increasing the number of items to be recalled or the duration of the retention period. We discuss also the observation that the hippocampus of storing birds is larger than that of non-storing birds, and find evidence contrary to the view that hippocampal enlargement is associated with enhanced spatial memory; we are, however, unable to suggest a convincing alternative explanation for hippocampal enlargement. The failure to find solid support for the ecological view supports the view that there are no qualitative differences in cognition between animal species in the processes of learning and memory. We also argue that our review supports our contention that speculation about the phylogenetic development and function of behavioural processes does not provide a solid basis for gaining insight into the nature of those processes. We end by confessing to a belief in one major qualitative difference in cognition in animals: we believe that humans alone are capable of acquiring language, and that it is this capacity that divides our intelligence so sharply from non-human intelligence.

Abstract: ABSTRACT Reciprocity is probably the most debated of the evolutionary explanations for cooperation. Part of the confusion surrounding this debate stems from a failure to note that two different processes can result in reciprocity: partner control and partner choice. We suggest that the common observation that group-living animals direct their cooperative behaviours preferentially to those individuals from which they receive most cooperation is to be interpreted as the result of the sum of the two separate processes of partner control and partner choice. We review evidence that partner choice is the prevalent process in primates and propose explanations for this pattern. We make predictions that highlight the need for studies that separate the effects of partner control and partner choice in a broader variety of group-living taxa.