Phylogenetic comparative methods and the geography of speciation

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Abstract

The geography of speciation has long been contentious. In recent years, phylogenetic approaches have been proposed to determine the geographical mode of speciation. If reliable, these methods not only provide a means of settling the debate about the geography of speciation, but also indicate that sympatric speciation is surprisingly common and that peripatric speciation is relatively rare. Similar to any phylogenetic inference, reconstructions of speciation mode are only useful if the underlying assumptions of the method are met. In this case, the key assumption is that the geographical range of both extant and ancestral species at the time of speciation can be inferred from present-day distributions. We discuss whether, and under what circumstances, such assumptions could be met. We conclude that interspecific phylogenies are unable to test alternative hypotheses concerning the geography of speciation rigorously because of the lability of geographical ranges and the lack of correlation between the role of adaptive processes and geographical mode of speciation.

Section snippets

Methods and assumptions

Most interspecific phylogenetic approaches to the geography of speciation work by examining the distribution of sister taxa: for example, if they are sympatric, then speciation is inferred to have been sympatric and if they are allopatric, speciation is inferred to have been allopatric (Fig. 1). For extant taxa, this approach leads to the comparison of sister species; for deeper nodes in a phylogeny, the procedure becomes more complicated because the geographical distribution of ancestral taxa

Shifts in range overlap through time

Several studies have used phylogenies to test geographical modes of speciation by examining changes in geographical range size and overlap of sister taxa through time 8, 9, 10, 21, 24 (Fig. 4). If, for example, allopatric speciation is predominant, then recently diverged sister taxa will tend to be geographically nonoverlapping, and the degree of overlap is likely to increase between deeper phylogenetic clades as a result of geographical range shifts. Conversely, if sympatric speciation is the

The future of phylogenetics in the study of the geography of speciation

Because speciation occurs at the population/species interface, methods are needed that focus at that level 25, 72. One recent approach has been to use phylogenetic approaches to focus solely on recent divergence events, either between populations within a species or within closely related species (reviewed in [73]). These approaches propose that the phylogenetic structure of genetic variation among populations and very recently diverged species will differ depending on geographical mode of

Conclusions

The past 15 years have seen a remarkable revolution in comparative biology: phylogenetic historical perspectives have gone from being completely ignored to being ubiquitous. As with many conceptual breakthroughs, however, the pendulum has swung too far the other way and re-examination of the utility of these methods has begun (e.g. 74, 75, 76). Phylogenetic approaches, similar to any other analytical method, have their own particular assumptions. When those assumptions are met, phylogenetic

Acknowledgements

We thank M. Francis Benard, L. Harmon, J. Kolbe, J. Wakeley, D. Ziolkowski and three anonymous reviewers for comments. Peter Jackson shared his expertise on the geographical range of wild felids. This work was supported by the National Science Foundation (DEB 9982736).

Glossary

Glossary

Allopatric speciation:
speciation resulting from divergent evolution of populations that are geographically isolated from each other.
Parapatric speciation:
speciation resulting from divergent evolution of populations that are geographically adjacent to each other.
Peripatric speciation:
a subset of allopatric speciation in which a peripherally isolated population diverges to become a new species.
Peripheral isolates:
a geographically isolated population on the periphery of a species' range.
Speciation:

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