As the environment changes, will species be able to adapt? By conducting experiments in natural environments, biologists can study how evolutionary processes such as natural selection operate through time. We predicted that the introduction of a terrestrial predator would first select for longer-legged lizards, which are faster, but as the lizards shifted onto high twigs to avoid the predator, selection would reverse toward favoring the shorter-legged individuals better able to locomote there. Our experimental studies on 12 islets confirmed these predictions within a single generation, thus demonstrating the rapidity with which evolutionary forces can change during times of environmental flux.
Evolutionary ecologists have long debated the extent to which communities in similar environments but different geographic regions exhibit convergence. On the one hand, if species' adaptations and community structure are determined by environmental features, convergence would be expected. However, if historical contingencies have long-lasting effects convergence would be unlikely. Most studies to date have emphasized the differences between communities in similar environments and little quantitative evidence for convergence exists. The application of comparative phylogenetic methods to ecological studies provides an opportunity to further investigate hypotheses of convergence. We compared the evolutionary patterns of structural ecology and morphology of 42 species of iguanian lizards from deserts of Australia and North America. Using a comparative approach, we found that evolutionary convergence of ecology and morphology occurs both in overall, community-wide patterns and in terms of pairs of highly similar intercontinental pairs of species. This result indicates that in these desert lizards, deterministic adaptive evolution shapes community patterns and overrides the historical contingencies unique to particular lineages.
Recent advances in ecological niche modeling (ENM) algorithms, in conjunction with increasing availability of geographic information system (GIS) data, allow species' niches to be predicted over broad geographic areas using environmental characteristics associated with point localities for a given species. Consequently, the examination of how niches evolve is now possible using a regionally inclusive multivariate approach to characterize the environmental requirements of a species. Initial work that uses this approach has suggested that niche evolution is characterized by conservatism: the more closely related species are, the more similar are their niches. We applied a phylogenetic approach to examine niche evolution during the radiation of Cuban trunk-ground anoles (Anolis sagrei group), which has produced 15 species in Cuba. We modeled the niche of 11 species within this group using the WhyWhere ENM algorithm and examined the evolution of the niche using a phylogeny based on approximately 1500 base pairs of mitochondrial DNA. No general relationship exists between phylogenetic similarity and niche similarity. Examination of species pairs indicates some examples in which closely related species display niche conservatism and some in which they exhibit highly divergent niches. In addition, some distantly related species exhibit significant niche similarity. Comparisons also revealed a specialist-generalist sister species pair in which the niche of one species is nested within, and much narrower than, the niche of another closely related species.
Examples of convergent evolution suggest that natural selection can often produce predictable evolutionary outcomes. However, unique histories among species can lead to divergent evolution regardless of their shared selective pressures-and some contend that such historical contingencies produce the dominant features of evolution. A classic example of convergent evolution is the set of Anolis lizard ecomorphs of the Greater Antilles. On each of four islands, anole species partition the structural habitat into at least four categories, exhibiting similar morphologies within each category. We assessed the relative importance of shared selection due to habitat similarity, unique island histories, and unique effects of similar habitats on different islands in the generation of morphological variation in anole ecomorphs. We found that shared features of diversification across habitats were of greatest importance, but island effects on morphology (reflecting either island effects per se or phylogenetic relationships) and unique aspects of habitat diversification on different islands were also important. There were three distinct cases of island-specific habitat diversification, and only one was confounded by phylogenetic relatedness. The other two unique aspects were not related to shared ancestry but might reflect as-yet-unmeasured environmental differences between islands in habitat characteristics. Quantifying the relative importance of shared and unique responses to similar selective regimes provides a more complete understanding of phenotypic diversification, even in this much-studied system.
Overwater dispersal and subsequent allopatric speciation contribute importantly to the species diversity of West Indian Anolis lizards and many other island radiations. Here we use molecular phylogenetic analyses to assess the contribution of overwater dispersal to diversification of the Anolis carolinensis subgroup, a clade comprising nine canopy-dwelling species distributed across the northern Caribbean. Although this clade includes some of the most successful dispersers and colonists in the anole radiation, the taxonomic status and origin of many endemic populations have been ambiguous. New mitochondrial and nuclear DNA sequences from four species occurring on small islands or island banks (Anolis brunneus, Anolis longiceps, Anolis maynardi, Anolis smaragdinus) and one species from the continental United States (A. carolinensis) are presented and analysed with homologous sequences sampled from related species on Cuba (Anolis allisoni and Anolis porcatus). Our analyses confirm that all five non-Cuban species included in our study represent distinct, independently evolving lineages that warrant continued species recognition. Moreover, our results support Ernest Williams's hypothesis that all of these species originated by overseas colonization from Cuban source populations. However, contrary to Williams's hypothesis of Pleistocene dispersal, most colonization events leading to speciation apparently occurred earlier, in the late Miocene-Pliocene. These patterns suggest that overwater dispersal among geologically distinct islands and island banks is relatively infrequent in anoles and has contributed to allopatric speciation. Finally, our results suggest that large Greater Antillean islands serve as centres of origin for regional species diversity.
Convergent evolution has played an important role in the development of the ecological niche concept. We investigated patterns of convergent and divergent evolution of Caribbean Anolis lizards. These lizards diversified independently on each of the islands of the Greater Antilles, producing the same set of habitat specialists on each island. Using a phylogenetic comparative framework, we examined patterns of morphological convergence in five functionally distinct sets of morphological characters: body size, body shape, head shape, lamella number, and sexual size dimorphism. We find evidence for convergence among members of the habitat specialist types for each of these five datasets. Furthermore, the patterns of convergence differ among at least four of the five datasets; habitat specialists that are similar for one set of characters are often greatly different for another. This suggests that the habitat specialist niches into which these anoles have evolved are multidimensional, involving several distinct and independent aspects of morphology.
Population phenomena, which provide much of the underlying basis for the theoretical structure of island biogeography, have received little direct study. We determined a key population trait-survival-in the Bahamian lizard Anolis sagrei on islands with an experimentally introduced predatory lizard and on neighboring unmanipulated islands. On unmanipulated islands, survival declined with several variables, most notably vegetation height: The island with the shortest vegetation had nearly the highest survival recorded for any lizard. On islands with the introduced predator, which forages mostly on the ground, A. sagrei shifted to taller vegetation; unlike on unmanipulated islands, its survival was very low on islands with the shortest vegetation but was higher on the others. Thus, species introduction radically changed a resident species' relation of survival to a key island-biogeographical variable.
Comparative studies have increased greatly in number in recent years due to advances in statistical and phylogenetic methodologies. For these studies, a trade-off often exists between the number of species that can be included in any given study and the number of individuals examined per species. Here, we describe a simple simulation study examining the effect of intraspecific sample size on statistical error in comparative studies. We find that ignoring measurement error has no effect on type I error of nonphylogenetic analyses, but can lead to increased type I error under some circumstances when using independent contrasts. We suggest using ANOVA to evaluate the relative amounts of within- and between-species variation when considering a phylogenetic comparative study. If within-species variance is particularly large and intraspecific sample sizes small, then either larger sample sizes or comparative methods that account for measurement error are necessary.