Behavioral ecology, also spelled behavioural ecology, is the study of the evolutionary basis for animal behavior due to ecological pressures. Behavioral ecology emerged from ethology after Niko Tinbergen outlined four questions to address when studying animal behavior which are the proximate causes, ontogeny, survival value, and phylogeny of behavior.
If an organism has a trait which provides them with a selective advantage (i.e. has an adaptive significance) in its environment, then natural selection can potentially favor it. Adaptive significance therefore refers to the beneficial qualities (such as in terms of increased survival and reproduction), any given modified trait conveys. For example, genetic differences between individuals may lead to behavioral differences, some of which in turn may drive differences in reproductive success, and ultimately over generations, the increased dominance of individuals with those favoured traits, i.e. evolution.
Individuals are always in competition with others for limited resources, including food, territories, and mates. Conflict will occur between predators and prey, between rivals for mates, between siblings, mates, and even between parents and their offspring.
The value of a social behavior depends in part on the social behavior of an animal's neighbors. For example, the more likely a rival male is to back down from a threat, the more value a male gets out of making the threat. The more likely, however, that a rival will attack if threatened, the less useful it is to threaten other males. When a population exhibits a number of interacting social behaviors such as this, it can evolve a stable pattern of behaviors known as an evolutionarily stable strategy (or ESS). This term, derived from economic game theory, became prominent after John Maynard Smith (1982) recognized the possible application of the concept of a Nash equilibrium to model the evolution of behavioral strategies.