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 behaviors: What are the proximate causes, ontogeny, survival value, and phylogeny of a behavior? If an organism has a trait that provides a selective advantage (i.e., has adaptive significance) in its environment, then natural selection favors it. Adaptive significance refers to the expression of a trait that affects fitness, measured by an individual's reproductive success. Adaptive traits are those that produce more copies of the individual's genes in future generations. Maladaptive traits are those that leave fewer. For example, if a bird that can call more loudly attracts more mates, then a loud call is an adaptive trait for that species because a louder bird mates more frequently than less loud birds—thus sending more loud-calling genes into future generations. Conversely, loud calling birds may attract the attention of predators more often, decreasing their presence in the gene pool. Individuals are always in competition with others for limited resources, including food, territories, and mates. Conflict occurs between predators and prey, between rivals for mates, between siblings, mates, and even between parents and 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.

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Life history theory
Life history theory is an analytical framework designed to study the diversity of life history strategies used by different organisms throughout the world, as well as the causes and results of the variation in their life cycles. It is a theory of biological evolution that seeks to explain aspects of organisms' anatomy and behavior by reference to the way that their life histories—including their reproductive development and behaviors, post-reproductive behaviors, and lifespan (length of time alive)—have been shaped by natural selection.
Foraging
Foraging is searching for wild food resources. It affects an animal's fitness because it plays an important role in an animal's ability to survive and reproduce. Foraging theory is a branch of behavioral ecology that studies the foraging behavior of animals in response to the environment where the animal lives. Behavioral ecologists use economic models and categories to understand foraging; many of these models are a type of optimal model. Thus foraging theory is discussed in terms of optimizing a payoff from a foraging decision.
Sociobiology
Sociobiology is a field of biology that aims to examine and explain social behavior in terms of evolution. It draws from disciplines including psychology, ethology, anthropology, evolution, zoology, archaeology, and population genetics. Within the study of human societies, sociobiology is closely allied to evolutionary anthropology, human behavioral ecology, evolutionary psychology, and sociology. Sociobiology investigates social behaviors such as mating patterns, territorial fights, pack hunting, and the hive society of social insects.
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