- Behavioral ecology
Behavioral ecology is the study of the ecological and evolutionary basis for
animal behavior, and the roles of behavior in enabling an animal to adapt to its environment (both intrinsic and extrinsic). Behavioral ecology emerged from ethologyafter Niko Tinbergen(a seminal figure in the study of animal behavior), outlined the four causes of behavior.
The two causes that contribute to ultimate causation are phylogenetic constraints that contribute to the development of behavior and the second is adaptive significance.
Phylogeneticconstraints are generally factors that might stop certain lineages developing certain behavioral or morphological traits. Hence, it is no coincidence that generally birds are able to flyand mammals cannot. The evolutionary history of these lineages have made it profitable for birds to fly and for mammalian feet to remain planted on the ground. Adaptive significance is akin to asking what a trait is good for in an evolutionary context. Therefore, the adaptive significance of flight in birds might have enabled avian ancestors to escape from predators. However, it is not sufficient to apply these explanations where they seem convenient. These have been labeled Just So Storiesby some biologists after Rudyard Kipling's tales for children about how animals came to be the way they are. To be rigorous, one must suppose a hypothesis and then test it scientifically. Hence, for avian flight, one can suppose that when birds are not at risk of being eaten, they might lose the ability to fly.
Proximate causation is also divided into two factors which are ontogenetic and mechanistic. Ontogenetic factors are the entire sum of experience throughout the lifetime of an individual from embryo to death. Hence, factors included are learning the genetic factors giving rise to behavior in individuals. Mechanistic factors, as the name implies, are the processes of the body that give rise to behavior such as the effects of
hormoneson behavior and neuronal basis of behavior.
Behavioral ecology, along with other areas of
evolutionary biology, has incorporated a number of techniques which have been borrowed from optimization theory. Optimization is a concept that stipulates strategies that offer the highest return to an animal given all the different factors and constraints facing the animal. One of the simplest ways to arrive at an optimal solution is to do a cost/benefit analysis. By considering the advantages of a behavior and the costs of a behavior, it can be seen that if the costs outweigh the benefits then a behavior will not evolve and vice versa. This is also where the concept of the trade-off becomes important. This is because it rarely pays an animal to invest maximally in any one behavior. For example, the amount of time an ectothermicanimal such as a lizard spends foraging is constrained by its body temperature. The digestive efficiency of the lizard also increases with increases in body temperature. Lizards increase their body temperature by basking in the sun. However, the time spent basking decreases the amount of time available for foraging. Basking also increases the risk of being discovered by a predator. Therefore, the optimal basking time is the outcome of the time necessary to sufficiently warm itself to carry out its activities such as foraging. This example shows how foraging is constrained by the need to bask (intrinsic constraint) and predation pressure (extrinsic constraint).
Differential reproductive success
Ultimately, all behavior is subject to
natural selectionas with any other trait of an animal. Therefore animals that employ optimal behavioral strategies specific to their environment will generally leave greater numbers of offspring than their suboptimal conspecifics. Animals that leave a greater number of offspring than others of their own species are said to have greater fitness than their suboptimal cousins. However, over time environments change meaning that what might be good behavior today might not be the best behavior in 10,000 years time or even 10 years time. Recent glaciation and future global warmingmean that one thing will be certain. The behavior of animals has and will continue to change in response to the environment. Behavioral ecology is one of the best ways to study these changes. As the great geneticist Theodosius Dobzhanskyfamously wrote, "nothing in biology makes sense except in the light of evolution."
Evolutionarily stable strategies
Another driving force in the evolution of animal behavior is the concept of an
evolutionarily stable strategy(or ESS), a term derived from economic game theorywhich became prominent after John Maynard Smith's 1982 book, Evolution and the Theory of Games. However, the concept can be traced back (as with most evolutionary ideas) to W.D. Hamilton, R.A. Fisher and Charles Darwin. In short, evolutionary game theoryasserts that only strategies that, when common in the population, cannot be "invaded" by any alternative rare (mutant) strategy will be ESSs, and thus maintained in the population. Therefore, animal behavior can be said to be governed not only by what is optimal, but also by what other strategies are found in the population. Furthermore, the relative frequencies of each strategy can influence the fitness of the other strategies in the population (frequency dependence). It is important to consider that evolution is not only driven by the physical environment, but also the interactions between other individuals.
Agent based model
Dear enemy recognition
Gene-centered view of evolution
Human behavioral ecology
Life history theory
Marginal value theorem
Phylogenetic comparative methods
r/K selection theory
* J.R. Krebs and Nicholas Davies, "An Introduction to Behavioural Ecology" [ISBN 0-632-03546-3]
* J.R. Krebs and Nicholas Davies, "Behavioural Ecology: an evolutionary approach" [ISBN 0-86542-731-3] (These two are respectively first/second college year level, and third/fourth college year level)
* Maynard Smith, J. 1982. Evolution and the Theory of Games.
Readings on "Human" Behavioral Ecology
* Borgerhoff Mulder, M. (2003). Human Behavioural Ecology. "Nature Encyclopedia of Life Sciences". [http://www.anthro.ucdavis.edu/faculty/monique/MBMWeb/MBMData/Nature-Enc.pdf Full text]
* Hames, R. (2001). Human Behavioral Ecology In
N. J. Smelserand Paul B. Baltes (Eds.) "International Encyclopedia of the Social & Behavioral Sciences". Pergamon, Oxford. 6946-695. [http://www.unl.edu/rhames/ms/behavioral-ecology-hames.pdf#search='human%20behavioral%20ecology' Full text]
* Winterhalder, Bruce & Smith, Eric Alden (2000). Analysing Adaptive Strategies: Human Behavioral Ecology at Twenty-Five. "Evolutionary Anthropology: Issues, News, and Reviews", Volume 9, Issue 2. [http://www.hbes.com/HBES/evolanth.pdf Full text]
* [http://blog.behavioralecology.net/ Behavioral Ecology Blog]
* [http://www-personal.umich.edu/~nesse/fourquestions.pdf The Four Areas of Biology]
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