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As anthropogenic activities are increasing the frequency and severity of droughts, understanding whether and how fast populations can adapt to sudden changes in their hydric environment is critically important. Here, we capitalize on the introduction of the Cuban brown anole lizard (Anolis sagrei) in North America to assess the contemporary evolution of a widespread terrestrial vertebrate to an abrupt climatic niche shift. We characterized hydric balance in 30 populations along a large climatic gradient. We found that while evaporative and cutaneous water loss varied widely, there was no climatic cline, as would be expected under adaptation. Furthermore, the skin of lizards from more arid environments was covered with smaller scales, a condition thought to limit water conservation and thus be maladaptive. In contrast to environmental conditions, genome-averaged ancestry was a significant predictor of water loss. This was reinforced by our genome-wide association analyses, which indicated a significant ancestry-specific effect for water loss at one locus. Thus, our study indicates that the water balance of invasive brown anoles is dictated by an environment-independent introduction and hybridization history and highlights genetic interactions or genetic correlations as factors that might forestall adaptation. Alternative water conservation strategies, including behavioral mitigation, may influence the brown anole invasion success and require future examination.

The G matrix, which quantifies the genetic architecture of traits, is often viewed as an evolutionary constraint. However, G can evolve in response to selection and may also be viewed as a product of adaptive evolution. Convergent evolution of G in similar environments would suggest that G evolves adaptively, but it is difficult to disentangle such effects from phylogeny. Here, we use the adaptive radiation of Anolis lizards to ask whether convergence of G accompanies the repeated evolution of habitat specialists, or ecomorphs, across the Greater Antilles. We measured G in seven species representing three ecomorphs (trunk-crown, trunk- ground, and grass-bush). We found that the overall structure of G does not converge. Instead, the structure of G is well conserved and displays a phylogenetic signal consistent with Brownian motion. However, several elements of G showed signatures of convergence, indicating that some aspects of genetic architecture have been shaped by selection. Most notably, genetic correlations between limb traits and body traits were weaker in long-legged trunk-ground species, suggesting effects of recurrent selection on limb length. Our results demonstrate that common selection pressures may have subtle but consistent effects on the evolution of G, even as its overall structure remains conserved.

In the state of Florida, USA, lizards of the genus Anolis are well represented with at least nine established non-native species and a single native species, A. carolinensis. The most recently introduced species is A. allisoni, a close relative to both the native A. carolinensis and one of the introduced species (A. porcatus). Anolis allisoni is thought to have been present in two locations in Florida since at least 2013 based on photographic evidence. Here, we analyzed mitochondrial DNA (mtDNA) sequences from these three closely related Anolis species to infer the most likely region of origin in the native range and confirm the establishment of the recent invader in Tampa, Florida. We found a single haplotype belonging to A. allisoni, which was closely related to native sequences from east-central Cuba. The most likely geographic origin is a tourist destination in the province of Sancti Spiritus, suggesting the potential for human-mediated introduction of A. allisoni to Florida. Given the evidence of hybridization within the carolinensis subgroup, the presence and establishment of the phylogenetically related and ecomorphologically similar A. allisoni may create novel opportunities for interspecific genetic exchange.

Rapid technological improvements are democratizing access to high quality, chromosome-scale genome assemblies. No longer the domain of only the most highly studied model organisms, now non-traditional and emerging model species can be genome-enabled using a combination of sequencing technologies and assembly software. Consequently, old ideas built on sparse sampling across the tree of life have recently been amended in the face of genomic data drawn from a growing number of high-quality reference genomes. Arguably the most valuable are those long-studied species for which much is already known about their biology; what many term emerging model species. Here, we report a highly complete chromosome-scale genome assembly for the brown anole, Anolis sagrei - a lizard species widely studied across a variety of disciplines and for which a high-quality reference genome was long overdue. This assembly exceeds the vast majority of existing reptile and snake genomes in contiguity (N50 = 253.6 Mb) and annotation completeness. Through the analysis of this genome and population resequence data, we examine the history of repetitive element accumulation, identify the X chromosome, and propose a hypothesis for the evolutionary history of fusions between autosomes and the X that led to the sex chromosomes of A. sagrei.

Rapid technological improvements are democratizing access to high quality, chromosome-scale genome assemblies. No longer the domain of only the most highly studied model organisms, now non-traditional and emerging model species can be genome-enabled using a combination of sequencing technologies and assembly software. Consequently, old ideas built on sparse sampling across the tree of life have recently been amended in the face of genomic data drawn from a growing number of high-quality reference genomes. Arguably the most valuable are those long-studied species for which much is already known about their biology; what many term emerging model species. Here, we report a new, highly complete chromosome-scale genome assembly for the brown anole, Anolis sagrei – a lizard species widely studied across a variety of disciplines and for which a high-quality reference genome was long overdue.

Body size is a key trait linked to many aspects of an organism’s life history, physiology, and behavior. Variation in body size can thus have important fitness consequences across a broad range of ecological contexts. We utilized the variation in multiple abiotic and biotic factors found among urban habitats to test simultaneously their relative effects on body size variation in urban populations of brown anole lizards (Anolis sagrei). These factors represent major hypotheses regarding body size variation in ectotherms and other animal groups: temperature, food availability, conspecific abundance and predator abundance. We also performed a tethered intruder experiment to determine whether the attack behavior of predatory curly-tailed lizards (Leiocephalus carinatus) depends on the body size of their brown anole prey. Lastly, we conducted a common garden experiment to assess whether brown anole body size differences between sites with and without curly-tailed lizard predators are genetically based. Predator abundance was the primary predictor of brown anole body size for both males and females. No other hypotheses were consistently supported. Brown anole body size increased along with increasing predator abundance, and predator abundance was negatively related to body size variation. Additionally, predators approached larger brown anoles less often and at longer latencies compared to smaller ones. Finally, male brown anoles from sites with predators had faster growth rates compared to those from sites without predators under common conditions in the lab. Brown anole body size differences among populations may result from higher survival of larger lizards during predatory confrontations, and this trait is at least partially heritable in males. Therefore, our results suggest that curly-tailed lizard predators may be important agents of evolution by natural selection for brown anole populations in urban habitats. To further investigate the ecological and evolutionary consequences of predator–prey interactions in urban habitats, future research should focus on measuring predator-induced selection in these novel environments.

Biological invasions are both a pressing environmental challenge and an opportunity to investigate fundamental ecological processes, such as the role of top predators in regulating biodiversity and food-web structure. In whole-ecosystem manipulations of small Caribbean islands on which brown anole lizards (Anolis sagrei) were the native top predator, we experimentally staged invasions by competitors (green anoles, Anolis smaragdinus) and/or new top predators (curly-tailed lizards, Leiocephalus carinatus). We show that curly-tailed lizards destabilized the coexistence of competing prey species, contrary to the classic idea of keystone predation. Fear-driven avoidance of predators collapsed the spatial and dietary niche structure that otherwise stabilized coexistence, which intensified interspecific competition within predator-free refuges and contributed to the extinction of green-anole populations on two islands. Moreover, whereas adding either green anoles or curly-tailed lizards lengthened food chains on the islands, adding both species reversed this effect—in part because the apex predators were trophic omnivores. Our results underscore the importance of top-down control in ecological communities, but show that its outcomes depend on prey behaviour, spatial structure, and omnivory. Diversity-enhancing effects of top predators cannot be assumed, and non-consumptive effects of predation risk may be a widespread constraint on species coexistence. Whole-ecosystem manipulations of Caribbean islands occupied by brown anoles, involving the addition of competitors (green anoles) and/or top predators (curly-tailed lizards), demonstrate that predator introductions can alter the ecological niches and destabilize the coexistence of competing prey species.

Urbanization significantly alters habitats for arboreal species, increasing the frequency of very smooth substrates by substituting artificial objects, such as metal poles and painted walls, for some trees. Because they experience these novel substrates more often, urban animals may use strategies to overcome challenges from substrate smoothness that animals from natural habitats do not. We assessed locomotor performance and two-dimensional hindlimb kinematics of two species of Anolis lizards (Anolis cristatellus and Anolis sagrei) from both urban and natural habitats in Miami, Florida. We ran lizards on six racetracks, crossing three substrates of increasing smoothness (rough bark, concrete blocks, and smooth, unpainted wood) with two inclinations (37° and vertical). We found that on vertical tracks with smooth substrates, lizards ran slower, took shorter strides and exhibited more contracted limb postures at the end of their stance than when running on the inclined track. Urban lizards, which are likely to be exposed more often to smooth substrates, did not adjust their movement to increase performance relative to lizards from natural habitats. This result, and the similarity of kinematic strategies between the two species, suggests the locomotor responses of lizards to substrate properties are highly conserved, which may be a mitigating factor that dampens or obviates the effects of natural selection on locomotor behaviour.

Extreme climate events such as droughts, cold snaps, and hurricanes can be powerful agents of natural selection, producing acute selective pressures very different from the everyday pressures acting on organisms. However, it remains unknown whether these infrequent but severe disruptions are quickly erased by quotidian selective forces, or whether they have the potential to durably shape biodiversity patterns across regions and clades. Here, we show that hurricanes have enduring evolutionary impacts on the morphology of anoles, a diverse Neotropical lizard clade. We first demonstrate a transgenerational effect of extreme selection on toepad area for two populations struck by hurricanes in 2017. Given this short-term effect of hurricanes, we then asked whether populations and species that more frequently experienced hurricanes have larger toepads. Using 70 y of historical hurricane data, we demonstrate that, indeed, toepad area positively correlates with hurricane activity for both 12 island populations of Anolis sagrei and 188 Anolis species throughout the Neotropics. Extreme climate events are intensifying due to climate change and may represent overlooked drivers of biogeographic and large-scale biodiversity patterns.

Some of the most important insights into the ecological and evolutionary processes of diversification and speciation have come from studies of island adaptive radiations, yet relatively little research has examined how these radiations initiate. We suggest that Anolis sagrei is a candidate for understanding the origins of the Caribbean Anolis adaptive radiation and how a colonizing anole species begins to undergo allopatric diversification, phenotypic divergence and, potentially, speciation. We undertook a genomic and morphological analysis of representative populations across the entire native range of A. sagrei, finding that the species originated in the early Pliocene, with the deepest divergence occurring between western and eastern Cuba. Lineages from these two regions subsequently colonized the northern Caribbean. We find that at the broadest scale, populations colonizing areas with fewer closely related competitors tend to evolve larger body size and more lamellae on their toepads. This trend follows expectations for post‐colonization divergence from progenitors and convergence in allopatry, whereby populations freed from competition with close relatives evolve towards common morphological and ecological optima. Taken together, our results show a complex history of ancient and recent Cuban diaspora with populations on competitor‐poor islands evolving away from their ancestral Cuban populations regardless of their phylogenetic relationships, thus providing insight into the original diversification of colonist anoles at the beginning of the radiation. Our research also supplies an evolutionary framework for the many studies of this increasingly important species in ecological and evolutionary research., We are grateful for funding from the Putnam Fund for Research and Exploration from the Museum of Comparative Zoology and the John Templeton Foundation. We are also grateful for funding from the National Museum of Natural History Research Opportunities Fund (to KdQ), National Museum of Natural History Biological Diversity Program (to KdQ), the National Science Foundation (#9318642 to JBL, KdQ, and A. Larson), the European Commission Marie Sklodowska‐Curie actions (to MLD), and Tenerife 2030 (P. INNOVA 2016–2021; MLD).

On microevolutionary timescales, adaptive evolution depends upon both natural selection and the underlying genetic architecture of traits under selection, which may constrain evolutionary outcomes. Whether such genetic constraints shape phenotypic diversity over macroevolutionary timescales is more controversial, however. One key prediction is that genetic constraints should bias the early stages of species divergence along “genetic lines of least resistance” defined by the genetic (co)variance matrix, G. This bias is expected to erode over time as species means and G matrices diverge, allowing phenotypes to evolve away from the major axis of variation. We tested for evidence of this signal in West Indian Anolis lizards, an iconic example of adaptive radiation. We found that the major axis of morphological evolution was well aligned with a major axis of genetic variance shared by all species despite separation times of 20–40 million years, suggesting that divergence occurred along a conserved genetic line of least resistance. Further, this signal persisted even as G itself evolved, apparently because the largest evolutionary changes in G were themselves aligned with the line of genetic least resistance. Our results demonstrate that the signature of genetic constraint may persist over much longer timescales than previously appreciated, even in the presence of evolving genetic architecture. This pattern may have arisen either because pervasive constraints have biased the course of adaptive evolution or because the G matrix itself has been shaped by selection to conform to the adaptive landscape. National Science Foundation NSF:DEB 0519658 NSF:0650078 NSF:DEB NSF:0519777 NSF:0722475 Virginia Tech University of Virginia

Persistence of animals in urban habitats, a stark environmental contrast to natural habitats, can be explained through evaluating the mechanisms behind organism-habitat interactions. One of the most notable effects of urbanization is the change in structural habitat; vegetation is removed and modified, favoring large trees and adding artificial structures in cities, which may alter how organismal preferences for aspects of the habitat are realized. We evaluated the mechanisms by which structural habitat changes associated with urbanization alter the available vegetation and substrates on which two species of Anolis lizards perch in urban and natural forest sites in Miami, FL. We also experimentally assessed habitat preference in the lab to establish the mechanism behind habitat selection. We found that vegetation was broader in urban areas compared to natural habitats, and artificial structures in urban areas were more than twice the diameter of available natural perches. Lizards expressed their preference for broad perches by selecting broader vegetation and artificial structures compared to their availability in both habitats. With the increased availability of broad substrates in urban areas, perch diameters selected by lizards resulted in an expansion of this aspect of the structural habitat niche for both species. The two species differed, however, in other responses to altered urban habitats. Anolis cristatellus tended to avoid artificial substrates, whereas A. sagrei used both natural and artificial structures in proportion to their availabilities. This study provides a mechanistic explanation for how urbanization alters structural habitats, leading to niche expansion for organisms living in cities.

Predation favors the unadventurous Selection is likely to shape behavior by acting on behavioral differences between individuals. Testing this idea has been challenging. Lapiedra et al. took advantage of a chain of small islands in the Caribbean colonized by anole lizards. A series of repeated behavioral selection experiments were set up in which brown anole populations were established with and without predators. On predator-free islands, animals that were more exploratory were favored, whereas when predators were present, less adventurous animals survived better. Selection for behavior occurred simultaneously with morphological selection but was predominant when predators were present. Science , this issue p. 1017

Only a handful of multi-generational experiments in natural systems of eco-evolutionary dynamics currently exist, despite Fussmann et al.'s call for more such studies nearly a decade ago. To perform such a study, in 2008 we introduced the lizard Leiocephalus carinatus, a predator (and possible food competitor) of the lizard Anolis sagrei, to seven islands having A. sagrei, with seven unmanipulated islands having A. sagrei as controls. Almost immediately, L. carinatus, which is larger and more terrestrial than A. sagrei, caused a major habitat shift in A. sagrei away from the ground and toward higher and thinner perches; focal behavioral surveys showed that on islands where its predator was introduced, A. sagrei had less conspicuous visual displays. The expected pattern for density of A. sagrei is that it would decrease markedly at first via predation from the larger lizard, but then it would increase as the habitat shift selected for individuals better able to live in higher vegetation. Data through 2015 ...

Human-mediated dispersal has reshaped distribution patterns and biogeographic relationships for many taxa. Long-distance and over-water dispersal were historically rare events for most species, but now human activities can move organisms quickly over long distances to new places. A potential consequence of human-mediated dispersal is the eventual reintroduction of individuals from an invasive population back into their native range; a dimension of biological invasion termed “cryptic back-introduction.” We investigated whether this phenomenon was occurring in the Cayman Islands where brown anole lizards (Anolis sagrei) with red dewlaps (i.e., throat fans), either native to Little Cayman or invasive on Grand Cayman, have been found on Cayman Brac where the native A. sagrei have yellow dewlaps. Our analysis of microsatellite data shows strong population-genetic structure among the three Cayman Islands, but also evidence for non-equilibrium. We found some instances of intermediate multilocus genotypes (possibly 3–9% of individuals), particularly between Grand Cayman and Cayman Brac. Furthermore, analysis of dewlap reflectance data classified six males sampled on Cayman Brac as having red dewlaps similar to lizards from Grand Cayman and Little Cayman. Lastly, one individual from Cayman Brac had an intermediate microsatellite genotype, a red dewlap, and a mtDNA haplotype from Grand Cayman. This mismatch among genetic and phenotypic data strongly suggests that invasive A. sagrei from Grand Cayman are interbreeding with native A. sagrei on Cayman Brac. To our knowledge, this is the first evidence of cryptic back-introduction. Although we demonstrate this phenomenon is occurring in the Cayman Islands, assessing its frequency there and prevalence in other systems may prove difficult due to the need for genetic data in most instances. Cryptic back-introductions may eventually provide some insight into how lineages are changed by the invasion process and may be an underappreciated way in which invasive species impact native biodiversity.

Temperature strongly affects performance in ectotherms. As ocean warming continues, performance of marine species will be impacted. Many studies have focused on how warming will impact physiology, life history, and behavior, but few studies have investigated how ecological and behavioral traits of organisms will affect their response to changing thermal environments. Here, we assessed the thermal tolerances and thermal sensitivity of swimming performance of two sympatric mysid shrimp species of the Northwest Atlantic. Neomysis americana and Heteromysis formosa overlap in habitat and many aspects of their ecological niche, but only N. americana exhibits vertical migration. In temperate coastal ecosystems, temperature stratification of the water column exposes vertical migrators to a wider range of temperatures on a daily basis. We found that N. americana had a significantly lower critical thermal minimum (CTmin) and critical thermal maximum (CTmax). However, both mysid species had a buffer of at least 4 °C between their CTmax and the 100-year projection for mean summer water temperatures of 28 °C. Swimming performance of the vertically migrating species was more sensitive to temperature variation, and this species exhibited faster burst swimming speeds. The generalist performance curve of H. formosa and specialist curve of N. americana are consistent with predictions based on the exposure of each species to temperature variation such that higher within-generation variability promotes specialization. However, these species violate the assumption of the specialist-generalist tradeoff in that the area under their performance curves is not constant. Our results highlight the importance of incorporating species-specific responses to temperature based on the ecology and behavior of organisms into climate change prediction models.

Urban landscapes are a mixture of built structures, human-altered vegetation, and remnant semi-natural areas. The spatial arrangement of abiotic and biotic conditions resulting from urbanization doubtless influences the establishment and spread of non-native species in a city. We investigated the effects of habitat structure, thermal microclimates, and species coexistence on the spread of a non-native lizard (Anolis cristatellus) in the Miami metropolitan area of South Florida (USA). We used transect surveys to estimate lizard occurrence and abundance on trees and to measure vegetation characteristics, and we assessed forest cover and impervious surface using GIS. We sampled lizard body temperatures, habitat use, and relative abundance at multiple sites. At least one of five Anolis species occupied 79 % of the 1035 trees surveyed in primarily residential areas, and non-native A. cristatellus occupied 25 % of trees. Presence and abundance of A. cristatellus were strongly associated with forest patches, dense vegetation, and high canopy cover, which produced cooler microclimates suitable for this species. Presence of A. cristatellus was negatively associated with the ecologically similar non-native A. sagrei, resulting in reduced abundance and a shift in perch use of A. cristatellus. The limited spread of A. cristatellus in Miami over 35 years is due to the patchy, low-density distribution of wooded habitat, which limits dispersal by diffusion. The presence of congeners may also limit spread. Open habitats—some parks, yards and roadsides—contain few if any A. cristatellus, and colonization of isolated forest habitat appears to depend on human-mediated dispersal.

In allopatric species, reproductive isolation evolves through the accumulation of genetic incompatibilities. The degree of divergence required for complete reproductive isolation is highly variable across taxa, which makes the outcome of secondary contact between allopatric species unpredictable. Since before the Pliocene, two species of Anolis lizards, Anolis carolinensis and Anolis porcatus, have been allopatric, yet this period of independent evolution has not led to substantial species‐specific morphological differentiation, and therefore, they might not be reproductively isolated. In this study, we determined the genetic consequences of localized, secondary contact between the native green anole, A. carolinensis, and the introduced Cuban green anole, A. porcatus, in South Miami. Using 18 microsatellite markers, we found that the South Miami population formed a genetic cluster distinct from both parental species. Mitochondrial DNA revealed maternal A. porcatus ancestry for 35% of the individuals sampled from this population, indicating a high degree of cytonuclear discordance. Thus, hybridization with A. porcatus, not just population structure within A. carolinensis, may be responsible for the genetic distinctiveness of this population. Using tree‐based maximum‐likelihood analysis, we found support for a more recent, secondary introduction of A. porcatus to Florida. Evidence that ~33% of the nuclear DNA resulted from a secondary introduction supports the hybrid origin of the green anole population in South Miami. We used multiple lines of evidence and multiple genetic markers to reconstruct otherwise cryptic patterns of species introduction and hybridization. Genetic evidence for a lack of reproductive isolation, as well as morphological similarities between the two species, supports revising the taxonomy of A. carolinensis to include A. porcatus from western Cuba. Future studies should target the current geographic extent of introgression originating from the past injection of genetic material from Cuban green anoles and determine the consequences for the evolutionary trajectory of green anole populations in southern Florida.

Biological invasions are both a pressing environmental challenge and an opportunity to investigate fundamental ecological processes, such as the role of top predators in regulating biodiversity and food-web structure. In whole-ecosystem manipulations of small Caribbean islands on which brown anole lizards (Anolis sagrei) were the native top predator, we experimentally staged invasions by competitors (green anoles, Anolis smaragdinus) and/or new top predators (curly-tailed lizards, Leiocephalus carinatus). We show that curly-tailed lizards destabilized the coexistence of competing prey species, contrary to the classic idea of keystone predation. Fear-driven avoidance of predators collapsed the spatial and dietary niche structure that otherwise stabilized coexistence, which intensified interspecific competition within predator-free refuges and contributed to the extinction of green-anole populations on two islands. Moreover, whereas adding either green anoles or curly-tailed lizards lengthened food chains on the islands, adding both species reversed this effect-in part because the apex predators were trophic omnivores. Our results underscore the importance of top-down control in ecological communities, but show that its outcomes depend on prey behaviour, spatial structure, and omnivory. Diversity-enhancing effects of top predators cannot be assumed, and non-consumptive effects of predation risk may be a widespread constraint on species coexistence.

The urban heat island effect, where urban areas exhibit higher temperatures than less-developed suburban and natural habitats, occurs in cities across the globe and is well understood from a physical perspective and at broad spatial scales. However, very little is known about how thermal variation caused by urbanization influences the ability of organisms to live in cities. Ectotherms are sensitive to environmental changes that affect thermal conditions, and therefore, increased urban temperatures may pose significant challenges to thermoregulation and alter temperature-dependent activity. To evaluate whether these changes to the thermal environment affect the persistence and dispersal of ectothermic species in urban areas, we studied two species of Anolis lizards (Anolis cristatellus and Anolis sagrei) introduced to Miami-Dade County, FL, USA, where they occur in both urban and natural habitats. We calculated canopy openness and measured operative temperature (Te ), which estimates the distribution of body temperatures in a non-thermoregulating population, in four urban and four natural sites. We also captured lizards throughout the day and recorded their internal body temperature (Tb ). We found that urban areas had more open canopies and higher Te compared to natural habitats. Laboratory trials showed that A. cristatellus preferred lower temperatures than A. sagrei. Urban sites currently occupied by each species appear to lower thermoregulatory costs for both species, but only A. sagreihad field Tb that were more often within their preferred temperature range in urban habitats compared to natural areas. Furthermore, based on available Te within each species' preferred temperature range, urban sites with only A. sagrei appear less suitable for A. cristatellus, whereas natural sites with only A. cristatellus are less suitable for A. sagrei. These results highlight how the thermal properties of urban areas contribute to patterns of persistence and dispersal, particularly relevant for studying species invasions worldwide.

Summary As animals move through their environments, they encounter a variety of substrates, which have important effects on their locomotor performance. Habitat modification can alter the types of substrates available for locomotion. In particular, many types of artificial substrates have been added to urban areas, but effects of these novel surfaces on animal locomotion are little known. In this study, we assessed locomotor performance of two Anolis lizard species (A. cristatellus and A. stratulus) on substrates that varied in inclination and surface roughness. Rough substrates represented the tree trunks and branches typically used in natural forest habitats, whereas smooth, vertical substrates captured the qualities of artificial surfaces, such as posts and walls, available in human-modified habitats. We then observed habitat use to test the habitat constraint hypothesis – that lizards should more frequently occupy portions of the habitat in which they perform better. Increased inclination and decreased surface roughness caused lizards to run slower. Both A. cristatellus and A. stratulus ran slowest on the smooth, vertical surface, and A. cristatellus often slipped and fell on this surface. In contrast to predictions, both species frequently used smooth, vertical substrates in the wild. Anolis cristatellus occupied artificial substrates 73% of the time in human-modified habitats despite performing worse than A. stratulus on the smooth, vertical track. We therefore rejected the habitat constraint hypothesis for anoles in these human-modified habitats. Despite overall poor performance on the smooth, vertical track, A. cristatellus had a significant morphology–performance relationship that supports the prediction that selection should favour smaller lizards with relatively longer limbs in human-modified habitats. The smaller bodied A. stratulus performed better than A. cristatellus on smooth, vertical substrates and therefore may not be exposed to the same selective pressures. We contend that habitat modification by humans may alter morphology-performance–habitat use relationships found in natural habitats. This may lead to changes in selective pressures for some species, which may influence their ability to occupy human-modified habitats such as cities.

Hurricanes are catastrophically destructive. Beyond their toll on human life and livelihoods, hurricanes have tremendous and often long-lasting effects on ecological systems1,2. Despite many examples of mass mortality events following hurricanes3–5, hurricane-induced natural selection has not previously been demonstrated. Immediately after we finished a survey of Anolis scriptus—a common, small-bodied lizard found throughout the Turks and Caicos archipelago—our study populations were battered by Hurricanes Irma and Maria. Shortly thereafter, we revisited the populations to determine whether morphological traits related to clinging capacity had shifted in the intervening six weeks and found that populations of surviving lizards differed in body size, relative limb length and toepad size from those present before the storm. Our serendipitous study, which to our knowledge is the first to use an immediately before and after comparison6 to investigate selection caused by hurricanes, demonstrates that hurricanes can induce phenotypic change in a population and strongly implicates natural selection as the cause. In the decades ahead, as extreme climate events are predicted to become more intense and prevalent7,8, our understanding of evolutionary dynamics needs to incorporate the effects of these potentially severe selective episodes9–11. Two populations of Anolis lizards that survived Hurricanes Irma and Maria had larger toepads, longer forelimbs and shorter hindlimbs relative to the pre-hurricane populations, which suggests hurricane-induced natural selection.

On microevolutionary timescales, adaptive evolution depends upon both natural selection and the underlying genetic architecture of traits under selection, which may constrain evolutionary outcomes. Whether such genetic constraints shape phenotypic diversity over macroevolutionary timescales is more controversial, however. One key prediction is that genetic constraints should bias the early stages of species divergence along “genetic lines of least resistance” defined by the genetic (co)variance matrix, G. This bias is expected to erode over time as species means and G matrices diverge, allowing phenotypes to evolve away from the major axis of variation. We tested for evidence of this signal in West Indian Anolis lizards, an iconic example of adaptive radiation. We found that the major axis of morphological evolution was well aligned with a major axis of genetic variance shared by all species despite separation times of 20–40 million years, suggesting that divergence occurred along a conserved genetic line of least resistance. Further, this signal persisted even as G itself evolved, apparently because the largest evolutionary changes in G were themselves aligned with the line of genetic least resistance. Our results demonstrate that the signature of genetic constraint may persist over much longer timescales than previously appreciated, even in the presence of evolving genetic architecture. This pattern may have arisen either because pervasive constraints have biased the course of adaptive evolution or because the G matrix itself has been shaped by selection to conform to the adaptive landscape.

Since the invention of electric lighting, artificial light at night (ALAN) has become a defining, and evolutionary novel, feature of human-altered environments especially in cities. ALAN imposes negative impacts on many organisms, including disrupting endocrine function, metabolism, and reproduction. However, we do not know how generalized these impacts are across taxa that exploit urban environments. We exposed brown anole lizards, an abundant and invasive urban exploiter, to relevant levels of ALAN in the laboratory and assessed effects on growth and reproduction at the start of the breeding season. Male and female anoles exposed to ALAN increased growth and did not suffer increased levels of corticosterone. ALAN exposure induced earlier egg-laying, likely by mimicking a longer photoperiod, and increased reproductive output without reducing offspring quality. These increases in growth and reproduction should increase fitness. Anoles, and potentially other taxa, may be resistant to some negative effects of ALAN and able to take advantage of the novel niche space ALAN creates. ALAN and both its negative and positive impacts may play a crucial role in determining which species invade and exploit urban environments.

Signaling individuals must effectively capture and hold the attention of intended conspecific receivers while limiting eavesdropping by potential predators. A possible mechanism for achieving this balance is for individuals to modulate the physical properties of their signals or to alter the proportion of time spent signaling, depending upon local levels of predation pressure. We test the hypothesis that prey can alter their visual signaling behavior to decrease conspicuousness and potentially limit predation risk via modulation of signal properties or display rate. To do so, we conducted a manipulative experiment in nature to evaluate the possible effect of predation pressure on the physical properties of movement-based signals and on the proportion of time spent signaling by using a well-understood predator-prey system in the Bahamas, the semiarboreal lizard Anolis sagrei, and one of its main predators, the curly-tailed lizard Leiocephalus carinatus. We find that on islands onto which the predator was introduced, male anoles reduce the maximum amplitude of head-bob displays but not the proportion of time spent signaling, in comparison with control islands lacking the predator. This reduction of amplitude also decreases signal active space, which might alter the reproductive success of signaling individuals. We suggest that future studies of predator-prey interactions consider the risk effects generated by changes in signals or signaling behavior to fully determine the influence of predation pressure on the dynamics of prey populations.

In polygynous lizards, male–male competition is an important driver of morphologic and behavioral traits associated with intraspecific dominance. The extent to which females engage in aggressive behavior and thus contribute to competition-driven morphologic variation is not well studied. We used injury frequencies of brown anoles (Anolis sagrei) in 16 island populations to test the hypothesis that injury-inducing aggressive encounters increase with population density in both male and female lizards. We further asked whether intraspecific competition is a potential driver of phenotypic traits related to dominance by using population density as proxy for intraspecific competition. We found that the proportion of individuals with injuries was greater in populations with higher densities, suggesting that agonistic competitive interactions increase with population density. Size-adjusted head length of male and female lizards increased with population density, suggesting that larger heads might be advantageous when intraspecific competition is strong. We detected differences in morphology and injury frequency among islands for both males and females, which suggests that agonistic competitive interactions among females may be stronger than previously appreciated. Further research is needed to determine whether aggressive encounters involving females are restricted to intrasexual competition or whether they also involve males, and how morphologic traits of females are related to competitive dominance and reproductive success.

The northeastern United States has experienced dramatic alteration to its landscape since the time of European settlement. This alteration has had major impacts on the distribution and abundance of wildlife populations, but the legacy of this landscape change remains largely unexplored for most species of freshwater turtles. We used microsatellite markers to characterize and compare the population genetic structure and diversity between an abundant generalist, the eastern painted turtle (Chrysemys p. picta), and the rare, more specialized, spotted turtle (Clemmys guttata) in Rhode Island, USA. We predicted that because spotted turtles have disproportionately experienced the detrimental effects of habitat loss and fragmentation associated with landscape change, that these effects would manifest in the form of higher inbreeding, less diversity, and greater population genetic structure compared to eastern painted turtles. As expected, eastern painted turtles exhibited little population genetic structure, showed no evidence of inbreeding, and little differentiation among sampling sites. For spotted turtles, however, results were consistent with certain predictions and inconsistent with others. We found evidence of modest inbreeding, as well as tentative evidence of recent population declines. However, genetic diversity and differentiation among sites were comparable between species. As our results do not suggest any major signals of genetic degradation in spotted turtles, the southern region of Rhode Island may serve as a regional conservation reserve network, where the maintenance of population viability and connectivity should be prioritized.

The transformation of natural habitats into urban landscapes dramatically alters thermal environments, which in turn, can impact local biota. Ectothermic organisms that are oviparous are particularly sensitive to these altered environments because their embryos cannot behaviorally thermoregulate and the surrounding environment determines the temperature experienced during development. We studied the effects of urban and forested thermal environments on embryo development and hatchling phenotypes in two non-native lizards (Anolis sagrei and A. cristatellus) in metropolitan Miami, Florida. To determine if embryos from urban and forested sites are adapted to their respective thermal environments, we incubated eggs from each site using temperatures that simulate likely nest conditions in both urban and forested environments. For both species, urban thermal environments accelerated embryonic development, but had no impact on egg survival or any of the phenotypic traits that were measured (e.g., body size, running performance, and locomotor behavior). Our results provide no evidence that embryos from urban and forested sites are adapted to their respective thermal environments. Instead, the lack of any major effects suggest that embryos of both species are physiologically robust with respect to novel environments, which could have facilitated their success in establishing in non-native ranges and in human-modified landscapes.

Novel selective pressures derived from human activities challenge the persistence of animal populations worldwide. Behavior is expected to be a major factor driving animals' responses to global change because it largely determines how animals interact with the environment. However, the role of individual variation in behavior to facilitate the persistence of animals in changing environments remains poorly understood. Here, we adopted an animal personality approach to investigate whether different behavioral traits allow animals to deal with two major components of global change: urbanization and biological invasions. By studying six populations of Anolis sagrei lizards, we found for the first time that anoles vary consistently in their behavior across different times and contexts. Importantly, these animal personalities were consistent in the wild and in captivity. We investigated whether behavioral traits are pulled in different directions by different components of global change. On the one hand, we found that lizards from urban areas differ from nearby forest lizards in that they were more tolerant of humans, less aggressive, bolder after a simulated predator attack, and they spent more time exploring new environments. Several of these risk-taking behaviors constituted a behavioral syndrome that significantly differed between urban and forest populations. On the other hand, the behavior of urban A. sagrei coexisting with the invasive predatory lizard Leiocephalus carinatus was associated with dramatic changes in their foraging niche. Overall, we provide evidence that differences in animal personalities facilitate the persistence of animals under novel selective regimes by producing adaptive behaviors relevant to their ecology such as predator avoidance. Our results suggest that natural selection can favor certain behaviors over others when animals are confronted with different ecological challenges posed by global change. Therefore, we underscore the need to incorporate behavioral ecology into the study of how animals adaptively respond to human-induced environmental changes.

Anolis carolinensis is an emerging model species and the sole member of its genus native to the United States. Considerable morphological and physiological variation has been described in the species, and the recent sequencing of its genome makes it an attractive system for studies of genome variation. To inform future studies of molecular and phenotypic variation within A. carolinensis, a rigorous account of intraspecific population structure and relatedness is needed. Here, we present the most extensive phylogeographic study of this species to date. Phylogenetic analyses of mitochondrial DNA sequence data support the previous hypothesis of a western Cuban origin of the species. We found five well-supported, geographically distinct mitochondrial haplotype clades throughout the southeastern United States. Most Florida populations fall into one of three divergent clades, whereas the vast majority of populations outside Florida belong to a single, shallowly diverged clade. Genetic boundaries do not correspond to major rivers, but may reflect effects of Pleistocene glaciation events and the Appalachian Mountains on migration and expansion of the species. Phylogeographic signal should be examined using nuclear loci to complement these findings.

Random and Directed Natural selection drives populations to adapt to new environments; the raw material or “founder effects” provided by the first colonizing individuals can thus have a formative influence on the population's future. Kolbe et al. (p. 1086 , published online 2 February; see the cover) tested the relative contributions of selection and founder effects in Bahamian lizards. Founders were taken from an island covered in forest: These lizards had long hindlimbs for sprinting across the broad expanses of tree trunks. Long-limbed lizards were introduced to seven smaller islands covered in scrub that, before hurricane Frances in 2004 swept them away, had been populated by lizards with short hindlimbs better suited for navigating a twiggy habitat. After several generations, all the new lizard populations had adapted to their new habitats by evolving shorter hindlimbs but they also retained other morphological and genetic signatures from their founding ancestors. Thus, evolution occurs by a combination of arbitrary events, as well as those shaped by selection.

Plastic responses of embryos to developmental environments can shape phenotypes in ways that impact fitness. The mechanisms by which developmental conditions affect offspring phenotypes vary substantially among taxa and are poorly understood in most systems. In this study, we evaluate the effects of thermal and hydric conditions on patterns of egg water uptake, embryonic development and yolk metabolism in embryos of the lizard Anolis sagrei to gain insights into how these factors shape morphological variation in hatchlings. Our 3 × 2 experimental design (3 thermal and 2 hydric conditions) revealed that developmental temperature has strong effects on rates of development and yolk metabolism, but the impacts of moisture were minimal. Increased water uptake by eggs under relatively wet conditions resulted in larger hatchlings with less internalized residual yolk than hatchlings from dry-incubated eggs. However, the relatively small phenotypic differences among treatments may have small fitness consequences. These results demonstrate that embryos of A. sagrei can tolerate a broad range of environmental conditions without substantial impacts on critical morphological traits. Such embryonic tolerances may facilitate colonization and establishment in novel environments. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105, 25–41.

The adaptive landscape and the G-matrix are keys concepts for understanding how quantitative characters evolve during adaptive radiation. In particular, whether the adaptive landscape can drive convergence of phenotypic integration (i.e., the pattern of phenotypic variation and covariation summarized in the P-matrix) is not well studied. We estimated and compared P for 19 morphological traits in eight species of Caribbean Anolis lizards, finding that similarity in P among species was not correlated with phylogenetic distance. However, greater similarity in P among ecologically similar Anolis species (i.e., the trunk-ground ecomorph) suggests the role of convergent natural selection. Despite this convergence and relatively deep phylogenetic divergence, a large portion of eigenstructure of P is retained among our eight focal species. We also analyzed P as an approximation of G to test for correspondence with the pattern of phenotypic divergence in 21 Caribbean Anolis species. These patterns of covariation were coincident, suggesting that either genetic constraint has influenced the pattern of among-species divergence or, alternatively, that the adaptive landscape has influenced both G and the pattern of phenotypic divergence among species. We provide evidence for convergent evolution of phenotypic integration for one class of Anolis ecomorph, revealing yet another important dimension of evolutionary convergence in this group.

Anolis lizards are the focus of most Caribbean lizard community ecology studies with few studies including other common species that might influence community structure. To study niche relationships and interspecific interactions in Antiguan lizard communities, we used five offshore islands with varying combinations of three diurnal lizards: Ameiva griswoldi, Anolis leachii, and Anolis wattsi. We collected data on perch height, substrate, thermal microhabitat, body size, head length, daily activity, and abundance to characterize the ecological niche of each species. Ameiva griswoldi was more similar to A. leachii in size and daily activity, but more similar to A. wattsi in perch height, and A. leachii and A. wattsi were more similar in thermal microhabitat. This pattern of niche differentiation was consistent with niche complementarity, where species are similar on some niche axes but differ on others. Using the same niche characteristics as in species comparisons, we tested for a niche shift amo...

Molecular genetic analyses show that introduced populations undergoing biological invasions often bring together individuals from genetically disparate native-range source populations, which can elevate genotypic variation if these individuals interbreed. Differential admixture among multiple native-range sources explains mitochondrial haplotypic diversity within and differentiation among invasive populations of the lizardAnolis sagrei. Our examination of microsatellite variation supports the hypothesis that lizards from disparate native-range sources, identified using mtDNA haplotypes, form genetically admixed introduced populations. Furthermore, within-population genotypic diversity increases with the number of sources and among-population genotypic differentiation reflects disparity in their native-range sources. If adaptive genetic variation is similarly restructured, then the ability of invasive species to adapt to new conditions may be enhanced.

Invasive species are classically thought to suffer from reduced within-population genetic variation compared to their native-range sources due to founder effects and population bottlenecks during introduc- tion. Reduction in genetic variation in introduced species may limit population growth, increase the risk of extinction, and constrain adaptation, hindering the successful establishment and spread of an alien species. Results of recent empirical studies, however, show higher than expected genetic variation, rapid evolution, and multiple native-range sources in introduced populations, which challenge the classical scenario of invasive- species genetics. With mitochondrial DNA (mtDNA) sequence data, we examined the molecular genetics of 10 replicate introductions of 8 species of Anolis lizards. Eighty percent of introductions to Florida and the Domini- can Republic were from multiple native-range source populations. MtDNA haplotypes restricted to different geographically distinct populations in the native range of a species commonly occurred as intrapopulation polymorphisms in introduced populations. Two-thirds of introduced populations had two or more sources, and admixture elevated genetic variation in half of the introduced populations above levels typical of native-range populations. The mean pairwise sequence divergence among haplotypes sampled within introduced popula- tions was nearly twice that within native-range populations (2.6% vs. 1.4%). The dynamics of introductions from multiple sources and admixture explained the observed genetic contrasts between native and introduced Anolis populations better than the classical scenario for most introduced populations. Elevated genetic vari- ation through admixture occurred regardless of the mode or circumstances of an introduction. Little insight into the number of sources or amount of genetic variation in introduced populations was gained by knowing the number of physical introductions, the size of a species' non-native range, or whether it was a deliberate or accidental introduction. We hypothesize that elevated genetic variation through admixture of multiple sources is more common in biological invasions than previously thought. We propose that introductions follow a sequential, two-step process involving a reduction in genetic variation due to founder effects and population bottlenecks followed by an increase in genetic variation if admixture of individuals from multiple native-range sources occurs.

Caribbean Anolis lizards are a classic case of adaptive radiation, repeated four times across islands of the Greater Antilles. On each island, very similar patterns of evolutionary divergence have occurred, resulting in the evolution of the same set of ecological specialists—termed ecomorphs—on each island. However, this is only part of the story of the Caribbean anole radiations. Indeed, much of the species diversity of Caribbean Anolis occurs within clades of ecomorphs, which contain as many as 14 ecologically-similar species on a single island. We ask to what extent the classic model of ecological interactions as the driving force in adaptive radiation can account for this aspect of anole evolutionary diversity. Our answer is that it can in part, but not entirely. More generally, the most complete understanding of evolutionary diversification and radiation is achieved by studying multiple hierarchical evolutionary levels from clades to populations.

The positive relationship between hind-limb length and perch diameter is well established in Anolis lizards, both among populations of some species and among species. Interspecific comparisons indicate that longer legs confer an advantage for increased running speed on broad substrates, whereas shorter limbs provide greater maneuverability on narrow surfaces. In this light, phenotypic plasticity for hind-limb length previously detected in Anolis sagrei may be adaptive because hatchlings exposed to only broad substrates developed relatively longer hind limbs for their body size compared to hatchlings exposed to only narrow substrates. We tested the generality of hind-limb length plasticity in Anolis by conducting a hatchling growth experiment on Anolis carolinensis, a distant relative of A. sagrei and a different type of habitat specialist. Similar to A. sagrei, A. carolinensis grown in cages with different sized perches showed hind-limb length plasticity, but the magnitude of difference between treatments and sexes was less for A. carolinensis than for A. sagrei. This finding suggests either hind-limb plasticity is widespread within the genus Anolis or that it has evolved independently at least twice.

Aim We investigate biogeographic relationships within the lizard genus Anolis Daudin, 1802 to test the hypothesis that the mainland (Central and South American) Norops-clade species descended from a West Indian Anolis ancestor. Previous hypotheses have suggested that close island relatives of mainland Norops species (the Cuban Anolis sagrei and Jamaican A. grahami series) represent over-water dispersal from a mainland ancestor. These previous hypotheses predict that the A. sagrei and A. grahami series should be phylogenetically nested within a Norops clade whose ancestral geography traces to the mainland. If Norops is West Indian in origin, then West Indian species should span the deepest phylogenetic divergences within the Norops clade. Location Central and South America and West Indian islands. Methods The phylogenetic relationships of Anolis lizards are reconstructed from aligned DNA sequences using both parsimony and Bayesian approaches. Hypotheses are tested in two ways: (1) by reconstructing the ancestral geographic location for the Norops clade using Pagel & Lutzoni's (2002) Bayesian approach, and (2) by testing alternative topological arrangements via Wilcoxon Signed-Ranks tests (Templeton, 1983) and Shimodaira–Hasegawa tests (Shimodaira & Hasegawa, 1999). Results Our evidence supports an origin of mainland Norops anoles from a West Indian ancestor. A West Indian ancestor to the Norops clade is statistically supported, and alternatives to the biogeographic pattern [Cuban (Jamaican, Mainland)] are statistically rejected by Shimodaira–Hasegawa tests, although not by Wilcoxon Signed-Ranks tests. Main conclusions Our data support the hypothesis of a West Indian origin for mainland Norops. This result contradicts previous hypotheses and suggests that island forms may be an important source for mainland biodiversity.

1999 at an - 1.5 ha study site. In three of five years and overall, nests closer to the water edge had a higher probability of predation; and in one year nests closer to the wooded edge had a higher probability of predation. Although more turtles nested closer to the water edge as the nesting season progressed in some years and overall, this behavior does not explain the observed patterns of nest predation. We present a novel application of the cubic spline analysis to address the dynamics of predation across continuous distances from an edge and identify threshold values where the predation rate levels off. Threshold values of 25-40 m were detected in 1995, 1998, and with all five years combined. However, even though a significant edge effect was detected in 1997, a threshold was not clear. While an edge effect on predation was not detected in each year, this study provides evidence for a strong effect of distance from the water edge on nest predation over significant ecological time. Focusing on turtle nest predation and smaller spatial scales addresses previous taxonomic and spatial bias in edge effects research, and provides further support for the ecological importance of edge effects.

The effect of water dynamics is well known for some aspects of turtle biology, particularly for egg incubation. The role water plays during migration from the nest to the aquatic environment, however, is not well understood. To evaluate water loss and movement patterns of hatchling turtles during migration, we conducted an experimental release of 463 hatchling snapping turtles, Chelydra serpentina, under ecologically relevant field conditions. We recaptured 62.9% of hatchlings released, and 90% of these were caught during the first four days following release. Hatchlings migrating for less time lost more water than those migrating for a longer time; changes in water content ranged from +1.2% to −4.7%. Mean air temperature during the recapture interval was positively correlated with water loss. Furthermore, hatchlings lost significantly more water when recaptured during periods with no rainfall compared to intervals with rainfall (14.5% and 8.3% water loss, respectively). Additionally, more hatchl...