Person:

Losos, Jonathan

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.

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.

Background: Comparative studies of amniotes have been hindered by a dearth of reptilian molecular sequences. With the genomic assembly of the green anole, Anolis carolinensis available, non-avian reptilian genes can now be compared to mammalian, avian, and amphibian homologs. Furthermore, with more than 350 extant species in the genus Anolis, anoles are an unparalleled example of tetrapod genetic diversity and divergence. As an important ecological, genetic and now genomic reference, it is imperative to develop a standardized Anolis gene nomenclature alongside associated vocabularies and other useful metrics. Results: Here we report the formation of the Anolis Gene Nomenclature Committee (AGNC) and propose a standardized evolutionary characterization code that will help researchers to define gene orthology and paralogy with tetrapod homologs, provide a system for naming novel genes in Anolis and other reptiles, furnish abbreviations to facilitate comparative studies among the Anolis species and related iguanid squamates, and classify the geographical origins of Anolis subpopulations. Conclusions: This report has been generated in close consultation with members of the Anolis and genomic research communities, and using public database resources including NCBI and Ensembl. Updates will continue to be regularly posted to new research community websites such as lizardbase. We anticipate that this standardized gene nomenclature will facilitate the accessibility of reptilian sequences for comparative studies among tetrapods and will further serve as a template for other communities in their sequencing and annotation initiatives.

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.

Ecological character displacement (ECD), the evolutionary divergence of competing species, has oscillated wildly in scientific opinion. Initially thought to play a central role in community assembly and adaptive radiation, ECD recovered from a 1980s nadir to present-day prominence on the strength of many case studies compiled in several influential reviews. However, we reviewed recent studies and found that only nine of 144 cases are strong examples that have ruled out alternative explanations for an ECD-like pattern. We suggest that the rise in esteem of ECD has outpaced available data and that more complete, rather than simply more, case studies are needed. Recent years have revealed that evolutionary change can be observed as it occurs, opening the door to experimental field studies as a new approach to studying ECD.

Understanding how quickly physiological traits evolve is a topic of great interest, particularly in the context of how organisms can adapt in response to climate warming. Adjustment to novel thermal habitats may occur either through behavioural adjustments, physiological adaptation or both. Here, we test whether rates of evolution differ among physiological traits in the cybotoids, a clade of tropical Anolis lizards distributed in markedly different thermal environments on the Caribbean island of Hispaniola. We find that cold tolerance evolves considerably faster than heat tolerance, a difference that results because behavioural thermoregulation more effectively shields these organisms from selection on upper than lower temperature tolerances. Specifically, because lizards in very different environments behaviourally thermoregulate during the day to similar body temperatures, divergent selection on body temperature and heat tolerance is precluded, whereas night-time temperatures can only be partially buffered by behaviour, thereby exposing organisms to selection on cold tolerance. We discuss how exposure to selection on physiology influences divergence among tropical organisms and its implications for adaptive evolutionary response to climate warming.

Anolis lizards exhibit a remarkable degree of diversity in the shape, colour, pattern and size of their dewlaps. Asymmetry, where one side of the dewlap differs in pattern or colour from the other, has only been reported in one species, Anolis lineatus, and then on only one of the two islands from which it occurs. Given the importance of the dewlap in intra- and interspecific signalling, we expanded on previous work by (1) investigating whether the reported asymmetry actually occurs and, if so, whether it occurs on animals from both Aruba and Curaçao; (2) examining whether populations differ in other aspects of their morphology or ecology; and (3) resolving the evolutionary relationships and the history of the two populations. We confirmed the presence of the asymmetrical dewlap on Curaçao and found that the asymmetry extends to populations on Aruba as well. Animals on Curaçao were smaller overall than populations from Aruba with relatively shorter metatarsals, radii, and tibias but relatively deeper heads, longer jaws, and wider and more numerous toepads on fore and hind feet. Habitat use did not differ significantly between the islands. We found populations on Aruba and Curaçao to be reciprocally monophyletic with an early Pleistocene divergence of populations on the two islands. Neutrality tests indicate that neither population has seen any recent reduction in population size, making it unlikely that the asymmetry is a result of founder effects or is some other consequence of reduced genetic variation. A variety of factors likely account for the remarkable and unique dewlap morphology exhibited by this species, although more detailed field studies are required to test these hypotheses.

Aim:

To test the role of climate, dispersal limitation and biotic interactions in limiting species' distributions within an island adaptive radiation by integrating species traits, phylogeny and estimates of dispersal cost into climate-based species distribution models. Location:

Hispaniola. Methods:

Focusing on 26 species of Anolis lizards, we used multivariate adaptive regression splines to evaluate the contribution of climate, species interactions, phylogenetic history and dispersal limitation to species distributional limits. For each species, we mapped the morphological similarity of congenerics using traits of known ecological import and predicted that species would be less likely to occur in climatically suitable areas if they were inhabited by ecologically similar species. Dispersal limitation was incorporated by generating spatially explicit estimates of dispersal cost, based on inferred habitat suitability. We compared models including morphological similarity, dispersal cost, phylogeny and climate with climate-only models. Results were evaluated against a null model that conserved the spatial structure of species occurrences. Results:

Climate had a dominant role in shaping species distributions. However, for over one-third of species we also found evidence consistent with supplemental effects of species interactions, i.e. ecological niche incumbency. These species were less likely to occur in climatically suitable areas inhabited by a morphologically similar species. Dispersal limitation also supplemented climatic limits in most species. These results were robust to co-variation with phylogeny and to comparison with our null model. Conclusions:

These results suggest that, rather than act as mutually exclusive alternatives, multiple dimensions of the ecological niche, including climatic limits, biotic interactions and dispersal capacity, interact to shape species distributions and that local interactions can influence the broad-scale geography of species in a predictable way.

Investigating the properties of ecological landscapes that influence gene flow among populations can provide key insights into the earliest stages of biological divergence. Both ecological and geographical factors can reduce gene flow, which can lead to population divergence, but we know little of the relative strengths of these phenomena in nature. Here, we use a novel application of structural equation modelling to quantify the contributions of ecological and geographical isolation to spatial genetic divergence in 17 species of Anolis lizards. Our comparative analysis shows that although both processes contributed significantly, geographical isolation explained substantially more genetic divergence than ecological isolation (36.3 vs. 17.9% of variance respectively), suggesting that despite the proposed ubiquity of ecological divergence, non-ecological factors play the dominant role in the evolution of spatial genetic divergence.

The evolutionary processes that produce adaptive radiations are enigmatic. They can only be studied after the fact, once a radiation has occurred and been recognized, rather than while the processes are ongoing. One way to connect pattern to process is to study the processes driving divergence today among populations of species that belong to an adaptive radiation, and compare the results to patterns observed at a deeper, macroevolutionary level. We tested whether evolution is a deterministic process with similar outcomes during different stages of the adaptive radiation of Anolis lizards. Using a clade of terrestrial–scansorial lizards in the genus Anolis, we inferred the adaptive basis of spatial variation among contemporary populations and tested whether axes of phenotypic differentiation among them mirror known axes of diversification at deeper levels of the anole radiation. Nonparallel change associated with genetic divergence explains the vast majority of geographic variation. However, we found phenotypic variation to be adaptive as confirmed by convergence in populations occurring in similar habitats in different mountain ranges. Morphological diversification among populations recurs deterministically along two axes of diversification previously identified in the anole radiation, but the characters involved differ from those involved in adaptation at higher levels of anole phylogeny.