Caribbean Anolis Research Articles

Adaptive Radiation Without Independent Stages of Trait Evolution in a Group of Caribbean Anoles.

Adaptive radiation involves diversification along multiple trait axes, producing phenotypically diverse, species-rich lineages. Theory generally predicts that multi-trait evolution occurs via a "stages" model, with some traits saturating early in a lineage's history, and others diversifying later. Despite its multidimensional nature, however, we know surprisingly little about how different suites of traits evolve during adaptive radiation. Here, we investigated the rate, pattern, and timing of morphological and physiological evolution in the anole lizard adaptive radiation from the Caribbean island of Hispaniola. Rates and patterns of morphological and physiological diversity are largely unaligned, corresponding to independent selective pressures associated with structural and thermal niches. Cold tolerance evolution reflects parapatric divergence across elevation, rather than niche partitioning within communities. Heat tolerance evolution and the preferred temperature evolve more slowly than cold tolerance, reflecting behavioral buffering, particularly in edge-habitat species (a pattern associated with the Bogert effect). In contrast to the nearby island of Puerto Rico, closely related anoles on Hispaniola do not sympatrically partition thermal niche space. Instead, allopatric and parapatric separation across biogeographic and environmental boundaries serves to keep morphologically similar close relatives apart. The phenotypic diversity of this island's adaptive radiation accumulated largely as a by-product of time, with surprisingly few exceptional pulses of trait evolution. A better understanding of the processes that guide multidimensional trait evolution (and nuance therein) will prove key in determining whether the stages model should be considered a common theme of adaptive radiation.

Comparison of Hydric and Thermal Physiology in an Environmentally Diverse Clade of Caribbean Anoles.

As the world becomes warmer and precipitation patterns less predictable, organisms will experience greater heat and water stress. It is crucial to understand the factors that predict variation in thermal and hydric physiology among species. This study focuses on investigating the relationships between thermal and hydric diversity and their environmental predictors in a clade of Hispaniolan anole lizards, which are part of a broader Caribbean adaptive radiation. This clade, the "cybotoid" anoles, occupies a wide range of thermal habitats (from sea level to several kilometers above it) and hydric habitats (such as xeric scrub, broadleaf forest, and pine forest), setting up the possibility for ecophysiological specialization among species. Among the thermal traits, only cold tolerance is correlated with environmental temperature, and none of our climate variables are correlated with hydric physiology. Nevertheless, we found a negative relationship between heat tolerance (critical thermal maximum) and evaporative water loss at higher temperatures, such that more heat-tolerant lizards are also more desiccation-tolerant at higher temperatures. This finding hints at shared thermal and hydric specialization at higher temperatures, underscoring the importance of considering the interactive effects of temperature and water balance in ecophysiological studies. While ecophysiological differentiation is a core feature of the anole adaptive radiation, our results suggest that close relatives in this lineage do not diverge in hydric physiology and only diverge partially in thermal physiology.

Range Expansion Theory and Recolonization of Secondary Forests by a Caribbean Anole

Range Expansion Theory and Recolonization of Secondary Forests by a Caribbean Anole

Recolonization of secondary forests by a locally extinct Caribbean anole through the lens of range expansion theory.

Disturbance and recovery dynamics are characteristic features of many ecosystems. Disturbance dynamics are widely studied in ecology and conservation biology. Still, we know less about the ecological processes that drive ecosystem recovery. The ecological processes that mediate ecosystem recovery stand at the intersection of many theoretical frameworks. Range expansion theory is one of these complementary frameworks that can provide unique insights into the population-level processes that mediate ecosystem recovery, particularly fauna recolonization. Although the biodiversity patterns that follow the fauna recolonization of recovering forests have been well described in the literature, the ecological processes at the population level that drive these patterns remain conspicuously unknown. In this study, we tested three fundamental predictions of range expansion theory during the recolonization of recovering forests in Puerto Rico by a shade specialist anole, Anolis gundlachi. Range expansion theory predicts that individuals at the early stages of recolonization (i.e., younger forests) would have a high prevalence of dispersive traits, experience less density dependence, and suffer less parasitism. To test these predictions, we conducted a chronosequence study applying space-for-time substitution where we compared phenotypic traits (i.e., body size, body condition, and relative limb size), population density, population growth rates, and Plasmodium parasitism rates among lizard populations living in young (<30 years), mid (~40-70 years), and old-growth forests (>75 years). Lizard populations in younger forests had lower densities, higher population growth rates, and lower rates of Plasmodium parasitism compared with old-growth forests. Still, while we found that individuals had larger body sizes, and longer forelimbs in young forests in one site, this result was not consistent among sites. This suggests a potential trade-off between the traits that provide a dispersal advantage during the initial stages of recolonization and those that are advantageous to establish in novel environmental conditions. Overall, our study emphasizes the suitability of range expansion theory to describe fauna recolonization but also highlights that the ecological processes that drive recolonization are time-dependent, complex, and nuanced.

Signals of differential introgression in the genome of natural hybrids of Caribbean anoles.

Hybridization facilitates recombination between divergent genetic lineages and can be shaped by both neutral and selective processes. Upon hybridization, loci with no net fitness effects introgress randomly from parental species into the genomes of hybrid individuals. Conversely, alleles from one parental species at some loci may provide a selective advantage to hybrids, resulting in patterns of introgression that do not conform to random expectations. We investigated genomic patterns of differential introgression in natural hybrids of two species of Caribbean anoles, Anolis pulchellus and A. krugi in Puerto Rico. Hybrids exhibit A. pulchellus phenotypes but possess A. krugi mitochondrial DNA, originated from multiple, independent hybridization events, and appear to have replaced pure A. pulchellus across a large area in western Puerto Rico. Combining genome-wide SNP datasets with bioinformatic methods to identify signals of differential introgression in hybrids, we demonstrate that the genomes of hybrids are dominated by pulchellus-derived alleles and show only 10%-20% A. krugi ancestry. The majority of A. krugi loci in hybrids exhibit a signal of non-random differential introgression and include loci linked to genes involved in development and immune function. Three of these genes (delta like canonical notch ligand 1, jagged1 and notch receptor 1) affect cell differentiation and growth and interact with mitochondrial function. Our results suggest that differential non-random introgression for a subset of loci may be driven by selection favouring the inheritance of compatible mitochondrial and nuclear-encoded genes in hybrids.

DIFFERENCES IN THE HINDLIMB ANATOMY IN THE TWO SPECIES OF THE LATE TRIASSIC DREPANOSAUROMORPH DIAPSID &lt;em&gt;MEGALANCOSAURUS&lt;/em&gt; INDICATE HABITAT PARTITIONING WITHIN THE ARBOREAL ENVIRONMENT

Discovery of two different morphologies of the hindlimb in specimens ascribed to the Triassic drepanosauromorph diapsid Megalancosaurus led to the erection of two distinct species within this genus: M. preonensis and M. endennae. In the present study a re-examination of the hindlimb architecture of both Megalancosaurus species indicates that M. preonensis and M. endennae exploited different microhabitats within the arboreal environment as for many extant species belonging to the same genus, like caribbean Anoles. M. endennae, with a stronger lower leg, a larger grasping foot, an opposable clawless hallux, and deeper and more recurved claws on pedal digits 2-5 was better adapted for locomotion on narrow and intricate supports like narrow twigs on terminal branches. M.preonensis with more slender hindlimbs, absence of an opposable hallux and longer, but thinner and less recurved claws on all pedal digits may have been better adapted to live on larger supports and could also have used the long pedal claws as hooks in a sloth-like fashion. In both Megalancosaurus species the tail acted both as a prop to assume a stable tripodal gait and as a clinging device that ensured grip during bridging as in other arboreal drepanosauromorphs which skeleton is adequately known. The overall hindlimb adaptation of Megalancosaurus species are more similar to that of some small arboreal mammals rather than that of chameleons.

Identifying individual and spatial drivers of heterogeneous transmission and virulence of malaria in Caribbean anoles

AbstractHeterogeneous distributions are a fundamental principle of ecology, manifesting as natural variability within ecological levels of organization from individuals to ecosystems. In disease ecology, variability in biotic and abiotic factors can result in heterogeneous patterns of transmission and virulence—broadly defined here as the negative consequences of infection. Still, our classic theoretical understanding of disease dynamics comes from models that assume homogeneous transmission and virulence. Here, we test this assumption by assessing the contribution of various sources of individual and spatial heterogeneity to patterns of transmission and sublethal measurements of virulence in two lizard–malaria systems: a three‐parasite assemblage (Plasmodium floridense,Plasmodium leukocytica, andPlasmodium azurophilum) infecting the lizardAnolis gundlachiin the rainforest of Puerto Rico and a single‐parasite system (P. floridense–Anolis sagrei) in Florida. Using a Bayesian model selection framework, we evaluated whether individual host differences (i.e., body size and sex) or spatial variability (i.e., habitat type and local‐scale host spatial structure) drive heterogeneity in the probability of infection or its associated health costs (i.e., body condition, blood chemistry). We found that the probability of infection increases with increasing lizard body size in both systems. However, in Florida, we found the relationship to be subdued in deforested habitats compared to the adjacent urban hydric forests. Furthermore, infection was spatially clustered within sampling sites, with “hot” and “cold” spots across the landscape. Nevertheless, we found no clear evidence of costs of infection on lizard health in any of the measures assessed and hence no grounds for inference regarding heterogeneous virulence. Ultimately, the consistency of our results across systems suggests prominent roles of individual and spatial heterogeneities as driving factors of transmission of vector‐borne diseases.

Signal detection shapes ornament allometry in functionally convergent Caribbean Anolis and Southeast Asian Draco lizards.

Visual ornaments have long been assumed to evolve hyper-allometry as an outcome of sexual selection. Yet growing evidence suggests many sexually selected morphologies can exhibit other scaling patterns with body size, including hypo-allometry. The large conspicuous throat fan, or dewlap, of arboreal Caribbean Anolis lizards was one ornament previously thought to conform to the classical expectation of hyper-allometry. We re-evaluated this classic example alongside a second arboreal group of lizards that has also independently evolved a functionally equivalent dewlap, the Southeast Asian Draco lizards. Across multiple closely related species in both genera, the Anolis and Draco dewlaps were either isometric or had hypo-allometric scaling patterns. In the case of the Anolis dewlap, variation in dewlap allometry was predicted by the distance of conspecifics and the light environment in which the dewlap was typically viewed. Signal efficacy, therefore, appears to have driven the evolution of hypo-allometry in what was originally thought to be a sexually selected ornament with hyper-allometry. Our findings suggest that other elaborate morphological structures used in social communication might similarly exhibit isometric or hypo-allometric scaling patterns because of environmental constraints on signal detection.

Infection with a novel pentastome (Raillietiella sp.) in a juvenile endangered Galápagos tortoise (Chelonoidis niger)

Pentastomes are crustacean endoparasites in the lungs of herpetofauna and rarely mammals and birds. To date, the only pentastomes reported in chelonians, all aquatic turtles, are Diesingia megastomum from Brazil and Pelonia africana from South Africa. In March 2021, 185 juvenile tortoises (Chelonoidis niger) were confiscated after an attempted illegal exportation from the Galápagos. It is believed the tortoises were removed from nests on Santa Cruz Island. The young tortoises were individually wrapped in plastic and at seizure ten were dead and another 25 tortoises subsequently died. One of the tortoises, estimated to be 3 months old, had 11 pentastomes in the lungs. The pentastomes were identified as a Raillietiella sp. based on morphology. The specimens had a claviform body that tapers into a bifid tail, a 165.3 μm × 92.1 μm buccal cadre, and sharp tipped anterior and posterior hooks with the posterior hooks being larger than the anterior. The males have smooth copulatory spicules with a rounded, smooth base. All females were gravid. Molecular analysis confirmed that the parasites were a Raillietiella sp. Based on the COI gene, it was most similar (82.7% identical) to an undescribed Raillietiella species from a Caribbean anole (Anolis cristatellus) found in Florida followed by R. hebitihamata (81%), R. indica (80.7%), and R. orientalis (78.8–80.7%). Based on the 18S rRNA gene sequence (1799 bp), it was most similar (99.3% identical) to two undescribed Raillietiella species followed by R. aegypti from a berber skink (Eumeces schneideri) from Saudia Arabia. Phylogenetically, with both molecular targets, the Raillietiella sp. from the Galápagos tortoise grouped with other Raillietiella spp. and was basal within the group. Currently, the origin of this parasite (native to Galápagos or introduced) and the life cycle are unknown. Because some pentastome species, especially when in aberrant hosts, can be pathogenic, additional studies of parasites in native and introduced reptile and amphibian species in the Galápagos are needed to better understand the risk this parasite poses to Galápagos tortoises.

No evidence of predicted phenotypic changes after hurricane disturbance in a shade-specialist Caribbean anole.

Extreme climatic events (ECEs) such as hurricanes have been hypothesized to be a major driving force of natural selection. Recent studies argue that, following strong hurricane disturbance, Anolis lizards in the Caribbean undergo selection for traits such as longer forelimbs or smaller body sizes that improve their clinging ability to their substrates increasing their chances of surviving hurricane wind gusts. Some authors challenge the generalization of this hypothesis arguing that other mechanisms may explain these phenotypic changes or that they may not necessarily be generalizable across systems. To address this issue, we compared body size and relative forelimb length of Anolis gundlachi, a trunk-ground anole living in closed-canopy forests in Puerto Rico, before, four months after, and 15 months after Hurricanes Irma and Maria in 2017. Overall, our results show no clear evidence of a temporal decrease in body size or increase forelimb length (relative to body size) challenging the generalizability of the clinging ability hypothesis. Understanding how animals adapt to ECE is an emerging field. Still, we are quickly learning that this process is complex and nuanced.

Parameters of the adhesive setae and setal fields of the Jamaican radiation of anoles (Dactyloidae: Anolis): potential for ecomorphology at the microscopic scale

Abstract The subdigital adhesive pads of Caribbean Anolis lizards are considered to be a key innovation that permits occupation of novel ecological niches. Although previous work has demonstrated that subdigital pad morphology and performance vary with habitat use, such investigations have only considered the macroscale aspects of these structures (e.g. pad area). The morphological agents of attachment, however, are arrays of hair-like fibres (setae) that terminate in an expanded tip (spatula) and have not been examined in a similar manner. Here we examine the setal morphology and setal field configuration of ecologically distinct species of the monophyletic Jamaican Anolis radiation from a functional and ecological perspective. We find that anoles occupying the highest perches possess greater setal densities and smaller spatulae than those exploiting lower perches. This finding is consistent with the concept of contact splitting, whereby subdivision of an adhesive area into smaller and more densely packed fibres results in an increase in adhesive performance. Micromorphological evidence also suggests that the biomechanics of adhesive locomotion may vary between Anolis ecomorphs. Our findings indicate that, in a similar fashion to macroscale features of the subdigital pad, its microstructure may vary in relation to performance and habitat use in Caribbean Anolis.

Can Sensory Drive Explain the Evolution of Visual Signal Diversity in Terrestrial Species? A Test with Anolis Lizards.

AbstractAnimal signal colors evolve to efficiently stimulate conspecific visual systems. The sensory drive hypothesis proposes that species differences in habitat light conditions favor the evolution of color diversity. The strongest support comes from aquatic systems, while terrestrial systems offer fewer convincing examples. Anolis lizards occupy diverse habitats and signal with a colorful dewlap. Dewlap visibility depends on perceived chromatic contrast with the background. Visual system modeling has shown that red dewlaps are most visible in most habitat types. However, a majority of species possess white or yellow dewlaps. In a recent behavioral study, we showed that low light conditions can sometimes make yellow and white colors more visible by altering chromatic contrast perception with the background. Using 17 Caribbean Anolis species, we showed that cut-on wavelength, a measure of dewlap color in a white to red continuum, correlates with habitat light intensity. Pairwise comparisons revealed that red dewlaps are most visible in bright habitats, whereas yellow and white are more visible in darker habitats. We conclude that sensory drive has contributed to the evolution of dewlap color differences through the interactive effects of total habitat light intensity and chromatic contrast perception and may provide a mechanism for speciation among anoles.

Animal trait variation at the within-individual level: erythrocyte size variation and malaria infection in a tropical lizard.

High levels of within-individual variation (WIV) in reiterative components in plants such as leaves, flowers, and fruits have been shown to increase individual fitness by multiple mechanisms including mediating interactions with natural enemies. This relationship between WIV and fitness has been studied almost exclusively in plant systems. While animals do not exhibit conspicuous reiterative components, they have traits that can vary at the individual level such as erythrocyte size. It is currently unknown if WIV in animals can influence individual fitness by mediating the outcome of interactions with natural enemies as it has been shown in plants. To address this issue, we tested for a relationship between WIV in erythrocyte size, hemoparasite infection status, and body condition (a proxy for fitness) in a Caribbean anole lizard. We quantified the coefficient of variation of adult erythrocytes size in $n = 95$ infected and $n = 107$ non-infected lizards. We found higher degrees of erythrocyte size variation in infected lizards than in non-infected individuals. However, we found no significant relationship between infection status or erythrocyte size variation, and lizard body condition. These results suggest that higher WIV in erythrocyte size in infected lizards is not necessarily adaptive but likely a consequence of the host response to infection. Many hemoparasites destroy their host cells as part of their life cycle. To compensate, the host lizard may respond by increasing production of erythrocytes resulting in higher WIV. Our results emphasize the need to better understand the role of within-animal variation as a neglected driver or consequence of ecological and evolutionary interactions.

Interspecific Gene Flow and Mitochondrial Genome Capture during the Radiation of Jamaican Anolis Lizards (Squamata; Iguanidae).

Gene flow and reticulation are increasingly recognized as important processes in the diversification of many taxonomic groups. With the increasing ease of collecting genomic data and the development of multispecies coalescent network approaches, such reticulations can be accounted for when inferring phylogeny and diversification. Caribbean Anolis lizards are a classic example of an adaptive radiation in which species have independently radiated on the islands of the Greater Antilles into the same ecomorph classes. Within the Jamaican radiation at least one species, Anolis opalinus, has been documented to be polyphyletic in its mitochondrial DNA, which could be the result of an ancient reticulation event or incomplete lineage sorting (ILS). Here, we generate mtDNA and genotyping-by-sequencing (GBS) data and implement gene tree, species tree, and multispecies coalescent network methods to infer the diversification of this group. Our mtDNA gene tree recovers the same relationships previously inferred for this group, which is strikingly different from the species tree inferred from our GBS data. Posterior predictive simulations suggest that our genomic data violate commonly adopted assumptions of the multispecies coalescent model (MSCM), so we use network approaches to infer phylogenetic relationships. The inferred network topology contains a reticulation event but does not explain the mtDNA polyphyly observed in this group; however, coalescent simulations suggest that the observed mtDNA topology is likely the result of past introgression. How common a signature of gene flow and reticulation is across the radiation of Anolis is unknown; however, the reticulation events that we demonstrate here may have allowed for adaptive evolution, as has been suggested in other, more recent, adaptive radiations. [Adaptive radiation; hybridization; introgression; multispecies network coalescent; posterior predictive simulation.].

Phylogeographic and phenotypic outcomes of brown anole colonization across the Caribbean provide insight into the beginning stages of an adaptive radiation.

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 speciesbegins 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 suppliesan evolutionary framework for the many studies of this increasingly important species in ecological and evolutionary research.

Bridging the Process-Pattern Divide to Understand the Origins and Early Stages of Adaptive Radiation: A Review of Approaches With Insights From Studies of Anolis Lizards.

Understanding the origins and early stages of diversification is one of the most elusive tasks in adaptive radiation research. Classical approaches, which aim to infer past processes from present-day patterns of biological diversity, are fraught with difficulties and assumptions. An alternative approach has been to study young clades of relatively few species, which may represent the putative early stages of adaptive radiation. However, it is difficult to predict whether those groups will ever reach the ecological and morphological disparity observed in the sorts of clades usually referred to as adaptive radiations, thereby making their utility in informing the early stages of such radiations uncertain. Caribbean Anolis lizards are a textbook example of an adaptive radiation; anoles have diversified independently on each of the 4 islands in the Greater Antilles, producing replicated radiations of phenotypically diverse species. However, the underlying processes that drove these radiations occurred 30-65 million years ago and so are unobservable, rendering major questions about how these radiations came to be difficult to tackle. What did the ancestral species of the anole radiation look like? How did new species arise? What processes drove adaptive diversification? Here, we review what we have learned about the cryptic early stages of adaptive radiation from studies of Anolis lizards, and how these studies have attempted to bridge the process-pattern divide of adaptive radiation research. Despite decades of research, however, fundamental questions linking eco-evolutionary processes to macroevolutionary patterns in anoles remain difficult to answer.

Morphological divergence and reduced ectoparasite prevalence in an introduced population of a Caribbean anole

AbstractAs biological invasions become increasingly common understanding what factors drive a species’ ability to rapidly colonize new habitats will be important for future conservation management. Theoretical models predict that spatial sorting of an invasive population will select for enhanced dispersal‐related traits and lower prevalence of parasitic infections. Supporting these, there is a growing body of evidence that demonstrates both features are common in invasive populations, although, these observations stem from mainland systems, over larger spatial scales. Here, we investigated the morphology and ectoparasite prevalence of Maynard's anole (Anolis maynardi) in its native range Little Cayman, and across its introduced range Cayman Brac where it invaded 31 years ago. We tested for morphological divergence, ectoparasite prevalence between the native population and the core, intermediate and range edge populations within the introduced range. In addition, we compared the overall body condition of lizards from each sex between islands using the scaled‐mass index. Our results suggest that spatial sorting could have selected for less parasitized individuals on the range edge of the introduced population, across an area of just 39 km2. However, there was no difference in hind‐limb length, the character that is expected to enhance dispersal ability, across the range of the introduced population. Instead, the greatest variation in morphology across the introduced range was found in fore‐limb length. Finally, both males and females from the introduced population were of significantly higher body condition than the native population, yet ectoparasite intensity had no effect on the body condition of infested hosts. These results highlight the rapid change in forelimb length and overall morphology has occurred rapidly since the species’ first introduction and a clear reduction in parasite prevalence on the range edge of the introduced population. Together these attributes have likely contributed to the successful colonization of Cayman Brac by A. maynardi.

Host defense triggers rapid adaptive radiation in experimentally evolving parasites.

Adaptive radiation occurs when the members of a single lineage evolve different adaptive forms in response to selection imposed by competitors or predators. Iconic examples include Darwin's finches, Caribbean anoles, and Hawaiian silverswords, all of which live on islands. Although adaptive radiation is thought to be an important generator of biodiversity, most studies concern groups that have already diversified. Here, we take the opposite approach. We experimentally triggered diversification in the descendants of a single population of host‐specific parasites confined to different host “islands.” We show rapid adaptive divergence of experimentally evolving feather lice in response to preening, which is a bird's main defense against ectoparasites. We demonstrate that host defense exerts strong phenotypic selection for crypsis in lice transferred to different colored rock pigeons (Columba livia). During four years of experimental evolution (∼60 generations), the lice evolved heritable differences in color. Strikingly, the observed color differences spanned the range of phenotypes found among congeneric lice adapted to other species of birds. To our knowledge, this is the first real‐time demonstration that microevolution is fast enough to simulate millions of years of macroevolutionary change. Our results further indicate that host‐mediated selection triggers rapid divergence in the adaptive radiation of parasites, which are among the most diverse organisms on Earth.

Climatic Niche Dynamics and Its Role in the Insular Endemism of Anolis Lizards

Insular systems are usually characterized by have a high species diversity, endemism, and evolutionary uniqueness. Although ecological and evolutionary factors shaping insular diversity and endemism are relatively well established, there is a little understanding about climatic niche dynamics for many insular adaptive radiations. Here, we evaluate the tempo and mode of climatic niche evolution in an iconic insular radiation of lizards. By using an extensive dataset of phylogenetic and coarse-grain climatic niches, we evaluated phylogenetic niche divergence and niche conservatism across temporal and spatial scales in the Caribbean Anolis lizard radiation. We found several instances of niche shifts during the anole radiation across islands. Many of these niche shifts converged to similar climatic regimes between different islands. Furthermore, we find evidence that single-island endemic species are more limited by low suitability of climatic conditions outside its native islands than oceanic barriers due to the high climatic heterogeneity observed at least between Greater Antillean islands. These results suggest that within-lineage climatic niche conservatism has been prevalent in short time scales and likely played a role driving the exceptional insular endemism observed today.

Parallel Behavioral Divergence with Macrohabitat inAnolis(Squamata: Dactyloidae) Lizards from the Dominican Republic

The ecomorph concept of the adaptive radiation of Caribbean anoles is characterized by a suite of behavioral, ecological, and morphological traits that are tightly linked to microhabitat use in lizards. However, most studies on the adaptive radiation of anoles have been conducted in a single macrohabitat type—lowland tropical forests. Because behavior can help organisms cope with different environmental conditions, we can predict that there will be key shifts in behavior within ecomorphs when examined across different macrohabitats, although this idea remains empirically underexplored. Here we utilized the replicated evolution of montane endemics from a primarily lowland species in a clade of trunk–ground Anolis lizards to test the hypothesis that shifts in basking behavior, wariness, and display behavior accompany divergence into montane habitats. The montane specialists A. armouri and A. shrevei each independently evolved from the primarily lowland dwelling A. cybotes in two widely separated mountain chains on the island of Hispaniola. We found evidence for a convergent behavioral response to the high-altitude macrohabitat: A. armouri and A. shrevei spend more time basking, utilize more open environments, and are warier than lowland A. cybotes. We also found divergence in display behavior in A. shrevei. We detected no evidence of divergence in locomotor behavior with elevation among active lizards. Together, our results suggest that the ecomorph concept would be enriched by extending observations of behavior (and other aspects of the phenotype) into different macrohabitats. Future work should focus on whether the observed behavioral shifts are clinal, reflecting local adaptation within A. cybotes, or fixed differences between the lowland generalist and montane species. Adaptation to the macrohabitat has previously been underappreciated as a source of behavioral diversity in Anolis lizards; this study is the first step toward documenting intraecomorph behavioral variation across divergent habitats.