Adaptive Radiation, Ecological Opportunity, and Evolutionary Determinism

Adaptive radiation refers to diversification from an ancestral species that produces descendants adapted to use a great variety of distinct ecological niches. In this review, I examine two aspects of adaptive radiation: first, that it results from ecological opportunity and, second, that it is deterministic in terms of its outcome and evolutionary trajectory. Ecological opportunity is usually a prerequisite for adaptive radiation, although in some cases, radiation can occur in the absence of preexisting opportunity. Nonetheless, many clades fail to radiate although seemingly in the presence of ecological opportunity; until methods are developed to identify and quantify ecological opportunity, the concept will have little predictive utility in understanding a priori when a clade might be expected to radiate. Although predicted by theory, replicated adaptive radiations occur only rarely, usually in closely related and poorly dispersing taxa found in the same region on islands or in lakes. Contingencies of a variety of types may usually preclude close similarity in the outcome of evolutionary diversification in other situations. Whether radiations usually unfold in the same general sequence is unclear because of the unreliability of methods requiring phylogenetic reconstruction of ancestral events. The synthesis of ecological, phylogenetic, experimental, and genomic advances promises to make the coming years a golden age for the study of adaptive radiation; natural history data, however, will always be crucial to understanding the forces shaping adaptation and evolutionary diversification.Organismic and Evolutionary Biolog

Seeing the Forest for the Trees: The Limitations of Phylogenies in Comparative Biology

The last 30 years have seen a revolution in comparative biology. Prior to that time, systematics was not at the forefront of the biological sciences, and few scientists considered phylogenetic relationships when investigating evolutionary questions. By contrast, systematic biology is now one of the most vigorous disciplines in biology, and the use of phylogenies is not only requisite in macroevolutionary studies, but has been applied to a wide range of topics and fields that no one could possibly have envisioned 30 years ago. My message is simple: phylogenies are fundamental to comparative biology, but they are not the be all and end all. Phylogenies are powerful tools for understanding the past, but like any tool, they have their limitations. In addition, phylogenies are much more informative about pattern than they are about process. The best way to fully understand the past—both pattern and process—is to integrate phylogenies with other types of historical data as well as with direct studies of evolutionary process.Organismic and Evolutionary Biolog

Area, climate heterogeneity, and the response of climate niches to ecological opportunity in island radiations of Anolis lizards

Aim Rates of climate niche evolution underlie numerous fundamental ecological processes and patterns. However, while climate niche conservatism varies markedly among regions and clades, the source of this variation remains poorly understood. We tested whether ecological opportunity can stimulate radiation within climate niche space at biogeographic scales, predicting that rates of climate niche evolution will scale with geographic area and climate heterogeneity. Location Caribbean Methods We quantified two temperature axes (mean temperature and temperature seasonality of species' localities) of the climate niche for 130 Anolis species on Cuba, Hispaniola, Puerto Rico, Jamaica and the northern and southern Lesser Antilles. Using a species-level phylogeny, we fitted macroevolutionary models that either constrained rates of climate niche evolution or allowed them to vary among regions. Next, we regressed region-specific evolutionary rates against area, species richness and climate heterogeneity. We evaluated whether results were robust to uncertainty in phylogenetic and biogeographic reconstructions and the assumed mode of evolution. Results For both niche axes, an Ornstein-Uhlenbeck model that allowed the net rate of evolution (σ2) to vary among island groups fit the data considerably better than a single-rate Brownian motion model. Nagelkerke pseudo-R2 values of the fit of these OU models to mean temperature and seasonality axes were 0.43 and 0.66, respectively. Evolutionary rates for both axes were higher in larger areas, which also have more species. Only the rate of mean occupied temperature evolution was positively related to climate heterogeneity, and only after accounting for region size. Conclusions Rates of climate niche evolution scale consistently with the area available for radiation, but responses to climate heterogeneity vary among niche axes. For the mean temperature axis, climate heterogeneity generated additional opportunities for radiation, but for seasonality it did not. Overall, the physical setting in which a clade diversifies can influence where it falls on the evolutionary continuum, from climate niche conservatism to radiation

Detective Work in the West Indies: Integrating Historical and Experimental Approaches to Study Island Lizard Evolution

Evolutionary biology is a historical science, like astronomy and geology. Understanding how and why evolution has occurred requires synthesizing multiple lines of inquiry. Historical studies, such as those that estimate phylogenetic trees, can detail the pattern of evolutionary diversification, whereas studies on living species can provide insight into the processes that affect ecological interactions and evolutionary change. The evolutionary radiation of Anolis lizards in the Greater Antilles illustrates the interplay between historical and modern-day approaches and strongly supports the hypothesis that interspecific interactions drive adaptive diversification. Studies of these species also demonstrate the role that manipulative experiments can play in understanding evolutionary phenomena.Organismic and Evolutionary Biolog

Convergence, Adaptation, and Constraint

Convergent evolution of similar phenotypic features in similar environmental contexts has long been taken as evidence of adaptation. Nonetheless, recent conceptual and empirical developments in many fields have led to a proliferation of ideas about the relationship between convergence and adaptation. Despite criticism from some systematically minded biologists, I reaffirm that convergence in taxa occupying similar selective environments often is the result of natural selection. However, convergent evolution of a trait in a particular environment can occur for reasons other than selection on that trait in that environment, and species can respond to similar selective pressures by evolving nonconvergent adaptations. For these reasons, studies of convergence should be coupled with other methods—such as direct measurements of selection or investigations of the functional correlates of trait evolution—to test hypotheses of adaptation. The independent acquisition of similar phenotypes by the same genetic or developmental pathway has been suggested as evidence of constraints on adaptation, a view widely repeated as genomic studies have documented phenotypic convergence resulting from change in the same genes, sometimes even by the same mutation. Contrary to some claims, convergence by changes in the same genes is not necessarily evidence of constraint, but rather suggests hypotheses that can test the relative roles of constraint and selection in directing phenotypic evolution.Organismic and Evolutionary BiologyOther Research Uni

The island–mainland species turnover relationship

Many oceanic islands are notable for their high endemism, suggesting that islands may promote unique assembly processes. However, mainland assemblages sometimes harbour comparable levels of endemism, suggesting that island biotas may not be as unique as is often assumed. Here, we test the uniqueness of island biotic assembly by comparing the rate of species turnover among islands and the mainland, after accounting for distance decay and environmental gradients. We modelled species turnover as a function of geographical and environmental distance for mainland (M–M) communities of Anolis lizards and Terrarana frogs, two clades that have diversified extensively on Caribbean islands and the mainland Neotropics. We compared mainland–island (M–I) and island–island (I–I) species turnover with predictions of the M–M model. If island assembly is not unique, then the M–M model should successfully predict M–I and I–I turnover, given geographical and environmental distance. We found that M–I turnover and, to a lesser extent, I–I turnover were significantly higher than predicted for both clades. Thus, in the first quantitative comparison of mainland–island species turnover, we confirm the long-held but untested assumption that island assemblages accumulate biodiversity differently than their mainland counterparts

What Drives Variation in Habitat Use by Anolis Lizards: Habitat Availability or Selectivity?

Geographic variation in habitat availability may drive geographic variation in a species' habitat use; alternatively, species adapted to particular habitat characteristics may use a habitat regardless of its availability within an environment. In this study, we investigated habitat use of two sympatric species of Anolis lizards that are morphologically specialized to use different microhabitats. We examined variation in microhabitat use and availability among four distinct forest types. In each forest type, we quantified available microhabitats (i.e., perch diameter, angle of inclination, and visibility), as well as microhabitats actually used by each species. We found that species consistently differed in microhabitat use, corresponding to each species' morphological specializations. However, microhabitat use of both species varied among sites. This variation in Anolis gundlachi Peters, 1876 reflected differences in microhabitat availability, while the variation in Anolis krugi Peters, 1876 resulted from differential microhabitat selectivity. These results indicate that both habitat availability and habitat preferences must be examined in multiple localities for a species to understand the causes of variation in its habitat use.Organismic and Evolutionary Biolog

Ecology and co-existence of two endemic day gecko (Phelsuma) species in Seychelles native palm forest

In island ecosystems, reptiles play diverse ecological roles as a result of niche broadening, which increases potential niche overlap between species. Ecological niche partitioning is a means of reducing direct competition between coexisting species and differences in habitat use among island gecko species have been suggested as a by-product of specialization to feeding on certain resources. Here, we examine modes and drivers of niche partitioning of two endemic species of Phelsuma gecko (Phelsuma sundbergi and Phelsuma astriata) in relict native palm forest in the Seychelles to further understanding of congeneric reptile co-existence in native habitats. Phelsuma abundance, microhabitat use and habitat composition were quantified in different macrohabitat types. P. sundbergi showed a clear preference for habitat dominated by the coco de mer palm, Lodoicea maldivica and a strong association with male individuals of this dioecious species. P. astriata density increased significantly with arboreal biodiversity but did not display a relationship with a specific tree type. High levels of resource segregation were determined along the microhabitat axis, based on differential tree preference. Our results suggest that P. sundbergi and P. astriata may have evolved to co-exist in this habitat type through partitioning of microhabitat as members of a divergent specialist/generalist assemblage determined by consumption of L. maldivica pollen by P. sundbergi. Our findings concur with the hypothesis that differences in habitat use among island reptiles are a by-product of trophic specialization and support the conservation of native habitat for maintenance of reptile diversity

Evolutionary biology for the 21st century

New theoretical and conceptual frameworks are required for evolutionary biology to capitalize on the wealth of data now becoming available from the study of genomes, phenotypes, and organisms - including humans - in their natural environments.Molecular and Cellular BiologyOrganismic and Evolutionary Biolog

Do the relationships between hindlimb anatomy and sprint speed variation differ between sexes in Anolis lizards?

The ability of an animal to run fast has important consequences on its survival capacity and overall fitness. Previous studies have documented how variation in the morphology of the limbs is related to variation in locomotor performance. Although these studies have suggested direct relations between sprint speed and hindlimb morphology, few quantitative data exist. Consequently, it remains unclear whether selection acts in limb segment lengths, overall muscle mass or muscle architecture (e.g. muscle fiber length and cross-sectional area). Here, we investigate whether muscle architecture (mass, fiber length and physiological cross-sectional area), hindlimb segment dimensions, or both, explain variation in sprint speed across 14 species of Anolis lizards. Moreover, we test whether similar relationships exist between morphology and performance for both sexes, which may not be the case given the known differences in locomotor behavior and habitat use. Our results show that the main driver of sprint speed is the variation in femur length for both males and females. Our results further show sexual dimorphism in the traits studied and, moreover, show differences in the traits that predict maximal sprint speed in males and females. For example, snout vent length and overall muscle mass are also good predictors of sprint speed in males, whereas no relationships between muscle mass and sprint speed was observed in females. Only a few significant relationships were found between muscle architecture (fiber length, cross-sectional area) and sprint speed in male anoles, suggesting that overall muscles size, rather than muscle architecture, appears to be under selection