Introduced species as moving targets: changes in body sizes of introduced lizards following experimental introductions and historical invasions

Introduced species usually fail to establish, but when they succeed, may undergo character release and rapid evolutionary divergence in novel environments. We collected brown anoles (Anolis sagrei: Lacertilia: Iguanidae) from a single Florida population and released them onto two ecologically different dredge-spoil islands in central Florida (forested and non-forested) and measured differences in population growth, individual growth, body size, and condition over four years. The population on the non-forested island expanded twice as fast as the forested island population and reached a density of ca. 12,000 lizards ha^–1 and a biomass of ca. 43.3 kg ha^–1, among the largest values recorded for non-aggregated terrestrial vertebrates. First-year progeny grew larger than their surviving parents on both islands, indicating character release occurred in early stages of both invasions. However, in subsequent years, lizards became larger on the forested island, but smaller on the non-forested island. Body condition declined over time on both islands, but the effect was most dramatic on the non-forested island. Lizards on the forested island had the lowest survival rates and highest tail autotomy frequencies. These results were attributed to differences in abiotic and biotic conditions on the two islands. Brown anoles are generally larger on islands where they have been introduced than on their native Caribbean islands, and are much larger on mainlands than on islands, indicating character release occurred at larger geographic scales as well. Habitat influences the morphology of introduced species possessing the ability to rapidly adapt to local conditions, presenting invasive species managers with moving targets.     

Physiological evolution during adaptive radiation: A test of the island effect in Anolis lizards

Phenotypic evolution is often exceptionally rapid on islands, resulting in numerous, ecologically diverse species. Although adaptive radiation proceeds along various phenotypic axes, the island effect of faster evolution has been mostly tested with regard to morphology. Here, we leveraged the physiological diversity and species richness of Anolis lizards to examine the evolutionary dynamics of three key traits: heat tolerance, body temperature, and cold tolerance. Contrary to expectation, we discovered slower heat tolerance evolution on islands. Additionally, island species evolve toward higher optimal body temperatures than mainland species. Higher optima and slower evolution in upper physiological limits are consistent with the Bogert effect, or evolutionary inertia due to thermoregulation. Correspondingly, body temperature is higher and more stable on islands than on the American mainland, despite similarity in thermal environments. Greater thermoregulation on islands may occur due to ecological release from competitors and predators compared to mainland environments. By reducing the costs of thermoregulation, ecological opportunity on islands may actually stymie, rather than hasten, physiological evolution. Our results emphasize that physiological diversity is an important axis of ecological differentiation in the adaptive radiation of anoles, and that behavior can impart distinct macroevolutionary footprints on physiological diversity on islands and continents.     

Selection at behavioural,developmental and metabolic genes is associated with the northward expansion of a successful tropical colonizer

What makes a species able to colonize novel environments? This question is key to understand the dynamics of adaptive radiations and ecological niche shifts, but the mechanisms that underlie expansion into novel habitats remain poorly understood at a genomic scale. Lizards from the genus Anolis are typically tropical, and the green anole (Anolis carolinensis) constitutes an exception since it expanded into temperate North America from subtropical Florida. Thus, we used the green anole as a model to investigate signatures of selection associated with colonization of a new environment, namely temperate North America. To this end, we analysed 29 whole‐genome sequences, covering the entire native range of the species. We used a combination of recent methods to quantify both positive and balancing selection in northern populations, including F_ST outlier methods, machine learning and ancestral recombination graphs. We naively scanned for genes of interest and assessed the overlap between multiple tests. Strikingly, we identified many genes involved in behaviour, suggesting that the recent successful colonization of northern environments may have been linked to behavioural shifts as well as physiological adaptation. Using a candidate genes strategy, we determined that genes involved in response to cold or behaviour displayed more frequently signals of selection, while controlling for local recombination rate, gene clustering and gene length. In addition, we found signatures of balancing selection at immune genes in all investigated genetic groups, but also at genes involved in neuronal and anatomical development.     

The visual ecology of Puerto Rican anoline lizards: habitat light and spectral sensitivity

The visual ecology of six closely related species of Puerto Rican anoline lizards was investigated and they were found to occupy four distinct habitat types in terms of light conditions: “full shade”, “partial shade”, “no shade”, and “forest canopy.”The habitats differed substantially in total radiance and irradiance as well as in the shape of the irradiance spectrum. The shape of the radiance spectrum was similar in all of the habitats. We used electroretinogram (ERG) flicker photometry to measure spectral sensitivity and found the curves for all six species to be similar. The spectral sensitivity peaked in the range 550–560 nm, which matched the peak in spectral radiance for all of the habitats. The shape of the spectral-sensitivity curve was similar to those of a number of other terrestrial vertebrates. We suggest that the convergence of the shape of the photopic ERG-determined spectral-sensitivity curve in many terrestrial vertebrates may, in part, be due to the fact that the background radiance of many terrestrial habitats is dominated by the reflectance spectrum of green vegetation which peaks at 550 nm. Accepted: 14 May 1997     

A circadian clock measures photoperiodic time in the male lizard Anolis carolinensis

Summary Photoperiod plays an important role in controlling the annual reproductive cycle of the male lizard Anolis carolinensis. The nature of photoperiodic time measurement in Anolis was investigated by exposing anoles to 3 different kinds of lighting paradigms (resonance, T cycles, and night breaks) to determine if photoperiodic time measurement involves the circadian system. Both the reproductive response and the patterns of entrainment of the activity rhythm were assessed. The results show that the circadian system is involved in photoperiodic time measurement in this species and that a discrete photoinducible phase resides in the latter half of the animals' subjective night. Significantly, the ability of the circadian system to execute photoperiodic time measurement is crucially dependent on the length of the photoperiod. Resonance, T cycle and night break cycles utilizing a photoperiod 10–11 h in duration reveal circadian involvement whereas these same cycles utilizing 6 or 8 h photoperiods do not.Abbreviation CRPP circadian rhythm of photoperiodic sensitivity     

Shared and unique features of diversification in Greater Antillean Anolis ecomorphs

Examples of convergent evolution suggest that natural selection can often produce predictable evolutionary outcomes. However, unique histories among species can lead to divergent evolution regardless of their shared selective pressures-and some contend that such historical contingencies produce the dominant features of evolution. A classic example of convergent evolution is the set of Anolis lizard ecomorphs of the Greater Antilles. On each of four islands, anole species partition the structural habitat into at least four categories, exhibiting similar morphologies within each category. We assessed the relative importance of shared selection due to habitat similarity, unique island histories, and unique effects of similar habitats on different islands in the generation of morphological variation in anole ecomorphs. We found that shared features of diversification across habitats were of greatest importance, but island effects on morphology (reflecting either island effects per se or phylogenetic relationships) and unique aspects of habitat diversification on different islands were also important. There were three distinct cases of island-specific habitat diversification, and only one was confounded by phylogenetic relatedness. The other two unique aspects were not related to shared ancestry but might reflect as-yet-unmeasured environmental differences between islands in habitat characteristics. Quantifying the relative importance of shared and unique responses to similar selective regimes provides a more complete understanding of phenotypic diversification, even in this much-studied system.     

Radioreceptor Assay for α;-Msh Using Mouse B16 Melanoma Cells

Abstract

A radioreceptor assay for α;-MSH is described which is based on cultured mouse B16 melanoma cells and bioactive monoiodinated [Nle⁴]-α;-MSH tracer. The assay was used (1) to study the binding characteristics of α;-MSH to B16 cells, (2) to determine the relative binding activity of MSH peptides, and (3) to measure MSH in tissue extracts. The association of α;-MSH to B16 cells reached a stable plateau after 3 h at 15°C. At 25° or 37°C, the binding was transient and at 0-1°C, the association was very slow. The hormone-receptor complex was relatively stable between 0° and 15°C whereas a 50% dissociation was reached after 90 min at 25°C and after 35 min at 37°C. The mean K_D for α;-MSH of four saturation experiments was 1.3 nM and the number of receptors 9570 per cell. 1,10-Phenanthroline had a stabilizing effect in the binding assay when used at a 0.3 mM concentration. From the MSH peptides tested in the binding assay, some showed similar potencies in three bioassays (tyrosinase, melanin and Anolis skin), whereas others displayed considerably     

A reconsideration of Sphaerodactylus dommeli Böhme, 1984 (Squamata: Gekkota: Sphaerodactylidae), a Miocene lizard in amber

The Miocene gecko in amber, Sphaerodactylus dommeli, is one of the best‐preserved fossil gekkotan species; nonetheless, its identity has been questioned and it has been insinuated to be an iguanian referable to the mega‐diverse genus Anolis. In this paper, we provide digital X‐rays and new osteological evidence that include 11 characters that reaffirms its placement within the infraorder Gekkota and eight characters that specifically place this fossil within the species‐rich genus Sphaerodactylus, in which it was originally described. This contribution seeks to eliminate remaining scepticism about the generic allocation of this species, and to provide a reliable calibration point for ongoing research in squamate phylogenetics, especially for biogeographical studies and molecular dating inference. This contribution also reviews many diagnostic characters for the two genera in question, which is critical for the correct identification of amber‐embedded specimens representing the rich Hispaniolan lizard paleocommunity.