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The geography of behaviour: an evolutionary perspective

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Abstract

Traditional approaches to the study of behaviour have typically assumed that behavioural patterns, especially elements of reproductive behaviour, are invariant within species. Recent research on a diversity of behavioural traits in a wide array of taxa provides evidence that genetically based geographic variation in behaviour is common. Comparisons of populations that display geographic variation in behaviour can offer substantial insight into mechanisms of adaptive divergence and constraining or generative roles of gene flow, initial stages of speciation, and the roles of phenotypic plasticity and ontogeny in determining patterns of behavioural evolution.

Section snippets

Divergence in behaviour among populations

Characteristics distinguishing gene pools considered to be subsets of the same species are likely to differ from those distinguishing species because populations tend to have had shorter periods of independent evolution. In particular, novelties (derived behavioural patterns not detected elsewhere within the clade) should appear in population clades less often than in higher-order clades. This is because genetically based behavioural novelties probably appear rarely and, when they do, can often

Adaptive inference

Population comparisons provide some of our best insights into the causes and mechanisms of adaptive differentiation, especially when the geographical distribution of ecotypes is both complex and unlikely to follow a cline in selective regimes2. However, even with an ideal scenario, in which parallel, independent evolution of similar behavioural ecotypes (homoplasies) can be inferred from molecular or geographical data1, methods other than population comparison are necessary to demonstrate the

Plasticity, learning and ontogeny

Many different types of behaviour in many taxa are extremely responsive to environmental conditions. In some instances, measurements of behavioural phenotypes under common conditions appropriately reflect population differences, but in others, measures of adaptive phenotypic plasticity (reaction norms) are more appropriate19, 23. Essentially, the evaluation of behavioural reaction norms is the evaluation of conditional responses to environmental variation, and these can differ among

Speciation

If, as is so often argued, allopatric differentiation is the most common means by which new species arise, speciation is implicitly geographical in nature. Given this observation, the paucity of instances in which population comparisons have been incorporated into research programmes exploring speciation is stunning5, 6, 26. Equally surprising is the tendency to view behavioural phenotypes involved in courtship and mate choice as invariant across populations when these could be the very traits

Conclusions

Behavioural traits, like other phenotypic characteristics, often exhibit geographic variation, and the variation often proves to be the expression of underlying genetic differentiation. Consequently, judicious comparisons across populations can help us understand the patterns and causes of both microevolutionary change and speciation. This approach is most likely to be of value when the taxa under consideration have relatively low dispersal potential or are separated by effective barriers to

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Acknowledgements

I thank R.G. Coss, W.A. Cresko, S.E. Riechert, S.I. Rothstein, P.A. Verrell and M.E. West for comments that improved the quality of this article. R.J. Scott and J.A. Walker drew all or parts of figures. I was partially supported by NSF Presidential Faculty Fellowship DEB-9253718.

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