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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.

Multivariate adaptation to climatic shifts may be limited by trait integration that causes genetic variation to be low in the direction of selection. However, strong episodes of selection induced by extreme climatic pressures may facilitate future population-wide responses if selection reduces trait integration and increases adaptive potential (i.e., evolvability). We explain this counter-intuitive framework for extreme climatic events in which directional selection leads to increased evolvability and exemplify its use in a case study. We tested this hypothesis in two populations of the lizard Arias scriptus that experienced hurricane-induced selection on limb traits. We surveyed populations immediately before and after the hurricane as well as the offspring of post-hurricane survivors, allowing us to estimate both selection and response to selection on key functional traits: forelimb length, hindlimb length, and toepad area. The direct selection was parallel in both islands and strong in several limb traits. Even though overall limb integration did not change after the hurricane, both populations showed a non-significant tendency toward increased evolvability after the hurricane despite the direction of selection not being aligned with the axis of most variance (i.e., body size). The population with comparably lower between-limb integration showed a less constrained response to selection. Hurricane-induced selection, not aligned with the pattern of high trait correlations, likely conflicts with selection occurring during normal ecological conditions that favours functional coordination between limb traits, and would likely need to be very strong and more persistent to elicit a greater change in trait integration and evolvability. Future tests of this hypothesis should use G-matrices in a variety of wild organisms experiencing selection due to extreme climatic events.

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.

With urbanization expanding into natural areas, it is increasingly important to understand how species subject to human-induced habitat alteration respond to novel opportunities and stressors. A pervasive consequence of urbanization is artificial light at night (ALAN), which previous studies have found introduces both costs and benefits for vertebrates. This understanding, however, primarily reflects findings from laboratory-controlled experiments or comparisons of wild populations in areas with long-standing differences in ALAN regimes. Here, we investigated the short-term costs and benefits for Anolis lizards during the period of initial exposure to ALAN using realistic light levels for urban areas (mean ± SD = 87.9 ± 36.7 lx at a distance of 3 m). As compared to controls, we hypothesized that adding ALAN would result in behavioral and physiological changes over the short term for brown anoles and their arthropod prey. In contrast to predictions, ALAN did not increase arthropod abundance or extend anole activity into the night. Structural habitat and sleep site use changed little in response to ALAN, which exposed about one-third of sleeping anoles in ALAN plots to light at night due to our manipulation. However, this direct light exposure resulted in lizards being more easily roused from sleep compared to lizards sleeping in the dark in control plots or in shadows in ALAN plots. The apparent inability of some anoles to adjust their sleep sites to avoid ALAN exposure may have contributed to their increased responsiveness at night and decreased locomotor endurance in the day. Our study suggests brown anoles can experience higher short-term costs than benefits during initial exposure to ALAN.

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.

Human activities have reshaped patterns of island biogeography for many groups of amphibians and reptiles. In Trinidad, an island in the West Indies with only one native Anolis lizard species, four additional anole species have been introduced since the 1800s. The most recent introduction, Anolis wattsi, native to Antigua, has become established despite the presence of multiple species of resident anoles. We used genetic data (mitochondrial DNA) to infer the geographic origin and genetic structure of introduced A. wattsi on Trinidad. We then gathered published data to compare eco-morphological traits of all anole species currently established on the island. We found three mtDNA haplotypes in Trinidad that clustered with two different clades from the northern part of the native range in Antigua, rejecting the hypothesis that the lizards originated in the pineapple-growing region of the south. However, a lack of fine-scale population structure precluded precise identification of the origin in Antigua of haplotypes introduced to Trinidad. Compared to the other anole species on Trinidad, A. wattsi is smaller, perches lower, and has a higher field body temperature. Thus, the successful establishment and spread of A. wattsi should not be surprising given the success of two previous introductions of anole species that were more similar ecomorphologically to the native species and each other, and the fact that Trinidad has relatively few anole species for its size.

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.

Behavioral adjustments may allow organisms to cope with the drastic environmental change caused by urbanization. We compared the escape behavior of Anolis lizards during simulated predator approaches on trees in both urban and forest habitats and on artificial structures (i.e., cement walls and metal posts) in urban areas. We found that urban lizards were less wary on trees, likely due at least in part to habituation to humans. In contrast, lizards on cement walls showed greater wariness, which may reflect a behavioral modification when escaping because of poor locomotor performance on this substrate. In both habitats, lizards modulated escape responses in ways consistent with their performance abilities on the various substrates. Escape by jumping decreased when lizards used wider and more isolated perches, which are characteristic of urban sites. Urban lizards squirreled (i.e., moved to the opposite side of a structure) more on wider trees and metal posts, and sprinted more but never jumped from cement walls. These behavioral responses for predator escape are consistent with the structural habitat changes caused by urbanization. Urban habitats had larger diameter trees, sparser ground vegetation, and many artificial structures. We found that urban lizards showed behavioral modulations because of exposure to humans as well as the locomotor constraints of using some artificial structures, such as cement walls. The ability of urban lizards to alter their escape responses and adjust to novel features of the habitat may be important traits of species able to colonize and persist in urban areas.

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.

Determining whether organisms have preferences for certain aspects of the habitat is important for understanding the process of habitat selection. Anolis lizards (anoles) have evolved to occupy distinct parts of the arboreal environment. Depending on the population or species, anoles perch on different aspects of the structural habitat varying in diameter, height, and type of substrate. We used laboratory and mesocosm trials to assess whether 2 species of Anolis lizards have preferences for the diameter and type of substrate they use for perching. Results reveal A. cristatellus and A. sagrei both prefer larger diameter perches. Lizards from natural forests and urban populations showed no difference in perch-diameter preference. In contrast, A. sagrei from an urban site overwhelmingly preferred to perch on wood substrates over polyvinyl chloride and metal ones. The larger diameter and wood substrates preferred by anoles in our trials should confer increased locomotor performance for these species based on previous studies. Rapid environmental change caused by humans can decouple the relationship between habitat preferences and fitness, so it is essential to test for changing preferences in organisms that experience urbanization and other forms of global change.

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 ...

Aim We examine the influence of fluctuating sea levels in a land-bridge archipelago on the apportioning of intraspecific genetic diversity and divergence in the widespread Puerto Rican crested anole (Anolis cristatellus). We compare three alternative scenarios for genetic diversification in an archipelagic species that contrast the relative influences of periodic isolation versus island connectedness driven by fluctuating sea levels. Our approach combines information from geography and population genetics to assess the influence of island size, island isolation, island historical geography, and population genetic processes such as drift on the contemporary distribution of genetic variation within and among islands. Location The Puerto Rico Bank in the Caribbean focusing primarily on the Spanish, British and U.S. Virgin Islands. Methods We used nuclear and mitochondrial DNA sequences and microsatellite genotypes sampled from A. cristatellus populations to investigate: (1) the broad-scale pattern of phylogeographical divergence across Puerto Rico Bank islands and (2) diversification within the Virgin Islands archipelago. For the first component, we used sequence data to reconstruct the relationships among 542 samples from across the species range. For the second component, we examined the relative influences of island size, isolation, and population genetic processes on the distribution of genetic diversity across the Virgin Islands. Results In the Virgin Islands, A. cristatellus is represented by a monophyletic clade except on the island of Vieques, where two divergent clades coexist. We found evidence for non-equilibrium dynamics in the Virgin Islands, suggesting spatial population expansion during intraglacial periods of low sea level. Main conclusions We found limited evidence that periods of island isolation affected patterns of genetic diversity and differentiation. Instead, we found that the patterns of genetic diversity and divergence in A. cristatellus in the Virgin Islands archipelago are likely shaped by long-term persistence in the region and periods of population spatial expansion.

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.

Foraging decisions reflect a trade-off between the benefits of acquiring food and the costs of movement. Changes in the biotic and abiotic environment associated with urbanization can alter this trade-off and modify foraging decisions. We experimentally manipulated foraging opportunities for two Anolis lizard species – the brown anole (A. sagrei) in Florida and the crested anole (A. cristatellus) in Puerto Rico – to assess whether foraging behavior differs between habitats varying in their degree of urbanization. In both urban and natural forest habitats, we measured the latency of perched anoles to feed from an experimental feeding tray. We manipulated perch availability and predator presence, while also taking into account population (e.g., conspecific density) and individual-level factors (e.g., body temperature) to evaluate whether and how these contribute to between-habitat differences in foraging behavior. In both species, urban anoles had longer latencies to feed and lower overall response rates compared to lizards from forests. Urban anoles were also larger (i.e., snout-vent length and mass) in both species and urban A. sagrei were in better body condition than the natural forest population. We postulate that the observed patterns in foraging behavior are driven by differences in perceived predation risk, foraging motivation, or neophobia. Although we are unable to identify the mechanism(s) driving these differences, the substantial differences in urban versus forest anole foraging behavior emphasizes the importance of understanding how urbanization influences animal populations and their persistence in anthropogenically-modified environments.

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.

Green Iguanas (Iguana iguana) are invasive in Puerto Rico due to a variety of negative economic effects, yet we know very little about their ecological impacts. Because they are herbivorous, defecate intact seeds, move through the forest, and have long gut-passage times, Green Iguanas may affect seed germination and seed dispersal. In summer 2013, a total of 258 Green Iguana scat samples were collected at the Humacao Natural Reserve in southeastern Puerto Rico. Seeds extracted from scat and collected from fruit were planted under common garden conditions using experimental treatments designed to tease apart the effects of feces, fruit, and ingestion on seed germination. Green Iguanas decreased the time for seeds to germinate in Ficus spp. by removing fruit pulp, but had no effect on germination of native Annona glabra seeds. For non-native P. pterocarpus and Pterocarpus spp., Green Iguanas produced conflicting results, decreasing the percentage of seeds germinating, but at the same time, reducing the time for seeds to germinate. Green Iguanas likely disperse most seeds beyond the canopies of parental tree at our site. Government and economic resources are being used to eradicate Green Iguana populations in Puerto Rico, but the lack of consistent effects of Green Iguanas on seed germination for the plant species consumed at our site complicates generalizing about their ecological effects and developing management plans that minimize negative effects for native plant communities. We recommend additional studies that target both species of particular concern, such as threatened native or invasive species, as well as studies of sensitive habitats in Puerto Rico.

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.

Most previous studies of clinging ability in Anolis lizards have focused on how toepads adhere to smooth, flat surfaces; yet lizards in nature use a wide variety of substrates ranging from smooth to rough and narrow to broad. I used an ecologically relevant measure of performance to determine the effects of hind-limb length and substrate diameter on clinging ability for two species of Anolis lizards from the British Virgin Islands. Clinging force was greater on the 12-mm smooth, wooden dowel compared to the 33-mm one for both species. Male Anolis cristatellus had considerably larger hind limbs than female A. cristatellus and both sexes of Anolis stratulus, and consequently performed substantially better on both dowel diameters. Results suggest two important components of clinging ability on cylindrical substrates: first, the ability of lizards to reach around and interlock their limbs with the substrate increases with longer limbs and narrower substrates; and second, the limb strength to maintain a grip on the dowel when limbs wrap fully around and the ability to apply compressive force when limbs wrap only partially around the dowel. These abilities are likely related to muscle size (e.g., cross-sectional area) and may be correlated with limb length. Further studies to understand how multiple morphological traits and different substrates affect clinging performance will contribute to a better understanding of this morphology-performance-habitat use relationship.

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.

Invasive species often encounter novel climatic conditions when they spread outside of their native ranges. Invading populations can respond to novel conditions by acclimation or adaptation of physiological capacities, which may facilitate their spread. We investigated differences in physiological traits among three populations of an invasive lizard, the brown anole (Anolis sagrei), along the latitudinal extent of its invasion in the southeastern United States. We predicted latitudinal clines for most traits based on models of adaptation to climate. Consistent with the latitudinal cline in temperature and moisture, the mean critical thermal minimum and the mean rate of water loss were lowest for lizards in the northern population. Furthermore, these traits acclimated to either temperature or humidity in a direction consistent with adaptive phenotypic plasticity. By contrast, metabolic rates varied among populations but did not conform to our prediction based on a latitudinal cline in temperature. Critical thermal maxima, endurances, and sprint speeds were similar among populations. Despite the idea that tropical lizards have limited capacity for acclimation, we found variation among invasive populations of brown anoles, which could have partially resulted from acclimation. This physiological variation within the invasive range raises questions about the roles of plasticity and adaptation in the success of the invasion.