On the Definitions and Functions of Dominance and Territoriality

Summary 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.