Why Do Curly Tail Lizards (Genus Leiocephalus) Curl Their Tails? An Assessment of Displays Toward Conspecifics and Predators

Animal display behaviors are used to convey specific messages to other animals, including potential mates, rivals, and predators. However, because these different types of interactions can be mediated by a single behavioral display, or conversely, multiple signals can be used to convey one specific message, interpretation of any particular behavioral display can be difficult. Leiocephalus lizards (i.e., curly tails) provide an excellent opportunity to study the use of display behaviors across multiple contexts. Previous research has demonstrated that the use of tail curling in these lizards is associated with predation risk, but less is known regarding the use of this behavior in social interactions with conspecifics. The goal of this study was to determine the extent to which tail curling display behavior is used to mediate both social and predatory interactions in two species, Leiocephalus barahonensis and L. carinatus. We found that in lizards of both species, tail curling was used in interactions with both conspecifics and potential (human) predators. However, tail curl intensity did not differ between lizards involved in social encounters and solitary lizards, although L. barahonensis lizards performed more headbobs during social than non-social observations. Further, L. carinatus lizards exhibited greater intensity of tail curling upon fleeing from a human predator than during observations in which individuals interacted with conspecifics, and lizards that exhibited tighter tail curls fled from predators for a longer distance. Finally, tail curl intensity was not correlated with headbob displays in either species, suggesting that these two components of display communicate different information. Our results suggest that tail curling displays, while consistently a component of interactions with potential predators, are not a necessary component of social interactions. These data contribute to a more complete understanding of how and why visual signals evolve for use in communication across multiple contexts

The Evolution of Androgen Receptor Expression and Behavior in \u3cem\u3eAnolis\u3c/em\u3e Lizard Forelimb Muscles

The motor systems that produce behavioral movements are among the primary targets for the action of steroid hormones, including androgens. Androgens such as testosterone bind to androgen receptors (AR) to induce physiological changes in the size, strength, and energetic capacity of skeletal muscles, which can directly influence the performance of behaviors in which those muscles are used. Because tissues differentially express AR, resulting in tissue-specific sensitivity to androgens, AR expression may be a major target of selection for the evolution of behavior. Anolis lizards (i.e., anoles) provide a robust system for the study of androgen-regulated traits, including the behavioral traits that facilitate social display and locomotion. In this study, we examined six anole species that demonstrate significant variation in the behavioral use of the forelimbs to measure the proportion of myonuclei in the bicep muscles that express AR. Using phylogenetic comparative analyses, we found that species with a greater proportion of nuclei positive for AR expression in the biceps exhibited greater frequencies of locomotor movements and pushup displays. These results suggest that AR expression in skeletal muscles may influence the evolution of androgen-regulated behaviors in this group

The Incredible Shrinking Dewlap: Signal Size, Skin Elasticity, and Mechanical Design in the Green Anole Lizard (Anolis Carolinensis)

The expression of male secondary sexual traits can be dynamic, changing size, shape, color, or structure over the course of different seasons. However, the factors underlying such changes are poorly understood. In male Anolis carolinensis lizards, a morphological secondary sexual signal called the dewlap changes size seasonally within individuals. Here, we test the hypothesis that seasonal changes in male dewlap size are driven by increased use and extension of the dewlap in spring and summer, when males are breeding, relative to the winter and fall. We captured male green anole lizards prior to the onset of breeding and constrained the dewlap in half of them such that it could not be extended. We then measured dewlap area in the spring, summer, and winter, and dewlap skin and belly skin elasticity in summer and winter. Dewlaps in unconstrained males increase in area from spring to summer and then shrink in the winter, whereas the dewlaps of constrained males consistently shrink from spring to winter. Dewlap skin is significantly more elastic than belly skin, and skin overall is more elastic in the summer relative to winter. These results show that seasonal changes in dewlap size are a function of skin elasticity and display frequency, and suggest that the mechanical properties of signaling structures can have important implications for signal evolution and design

The Evolution of Copulation Frequency and the Mechanisms of Reproduction in Male Anolis Lizards

The evolution of many morphological structures is associated with the behavioral context of their use, particularly for structures involved in copulation. Yet, few studies have considered evolutionary relationships among the integrated suite of structures associated with male reproduction. In this study, we examined nine species of lizards in the genus Anolis to determine whether larger copulatory morphologies and higher potential for copulatory muscle performance evolved in association with higher copulation rates. In 10–12 adult males of each species, we measured the size of the hemipenes and related muscles, the seminiferous tubules in the testes, and the renal sex segments in the kidneys, and we assessed the fiber type composition of the muscles associated with copulation. In a series of phylogenetically-informed analyses, we used field behavioral data to determine whether observed rates of copulation were associated with these morphologies.We found that species with larger hemipenes had larger fibers in the RPM (the retractor penis magnus, a muscle that controls hemipenis movement), and that the evolution of larger hemipenes and RPM fibers is associated with the evolution of higher rates of copulatory behavior. However, the sizes of the seminiferous tubules and renal sex segments, and the muscle fiber composition of the RPM, were not associated with copulation rates. Further, body size was not associated with the size of any of the reproductive structures investigated. The results of this study suggest that peripheral morphologies involved in the transfer of ejaculate may be more evolutionarily labile than internal structures involved in ejaculate production

IUCN Conservation Status Does Not Predict Glucocortoid Concentrations in Reptiles and Birds

Circulating glucocorticoids (GCs) are the most commonly used biomarkers of stress in wildlife. However, their utility as a tool for identifying and/or managing at-risk species has varied. Here, we took a very broad approach to conservation physiology, asking whether International Union for the Conservation of Nature (IUCN) listing status (concern versus no obvious concern) and/or location within a geographic range (edge versus non-edge) predicted baseline and post-restraint concentrations of corticosterone (CORT) among many species of birds and reptiles. Even though such an approach can be viewed as coarse, we asked in this analysis whether CORT concentrations might be useful to implicate species at risk. Indeed, our effort, relying on HormoneBase, a repository of data on wildlife steroids, complements several other large-scale efforts in this issue to describe and understand GC variation. Using a phylogenetically informed Bayesian approach, we found little evidence that either IUCN status or edge/non-edge location in a geographic distribution were related to GC levels. However, we did confirm patterns described in previous studies, namely that breeding condition and evolutionary relatedness among species predicted some GC variation. Given the broad scope of our work, we are reluctant to conclude that IUCN status and location within a range are unrelated to GC regulation. We encourage future more targeted efforts on GCs in at-risk populations to reveal how factors leading to IUCN listing or the environmental conditions at range edges impact individual performance and fitness, particularly in the mammals, amphibians, and fish species we could not study here because data are currently unavailable

Metabolic Scaling of Stress Hormones in Vertebrates

Glucocorticoids (GCs) are stress hormones that can strongly influence physiology, behavior, and an organism’s ability to cope with environmental change. Despite their importance, and the wealth of studies that have sought to understand how and why GC concentrations vary within species, we do not have a clear understanding of how circulating GC levels vary within and across the major vertebrate clades. New research has proposed that much interspecific variation in GC concentrations can be explained by variation in metabolism and body mass. Specifically, GC concentrations should vary proportionally with mass-specific metabolic rates and, given known scaling relationships between body mass and metabolic rate, GC concentrations should scale to the -1/4 power of body mass and to the power of 1 with mass-specific metabolic rate. Here, we use HormoneBase, the newly compiled database that includes plasma GC concentrations from free-living and unmanipulated vertebrates, to evaluate this hypothesis. Specifically, we explored the relationships between body mass or mass-specific metabolic rate and either baseline or stress-induced GC (cortisol or corticosterone) concentrations in tetrapods. Our phylogenetically-informed models suggest that, whereas the relationship between GC concentrations and body mass across tetrapods and among mammals is close to -1/4 power, this relationship does not exist in amphibians, reptiles, and birds. Moreover, with the exception of a positive association between stress-induced GC concentrations and mass-specific metabolic rate in birds, we found little evidence that GC concentrations are linked to metabolic rate, although the number of species sampled was quite limited for amphibians and somewhat so for reptiles and mammals. Nevertheless, these results stand in contrast to the generally accepted association between the two and suggest that our observed positive association between body mass and GC concentrations may not be due to the well-established link between mass and metabolism. Large-scale comparative approaches can come with drawbacks, such as pooling and pairing observations from separate sources. However, these broad analyses provide an important counterbalance to the majority of studies examining variation in GC concentrations at the population or species level, and can be a powerful approach to testing both long-standing and new questions in biology

HormoneBase, a Population-Level Database of Steroid Hormone Levels Across Vertebrates

Hormones are central regulators of organismal function and flexibility that mediate a diversity of phenotypic traits from early development through senescence. Yet despite these important roles, basic questions about how and why hormone systems vary within and across species remain unanswered. Here we describe HormoneBase, a database of circulating steroid hormone levels and their variation across vertebrates. This database aims to provide all available data on the mean, variation, and range of plasma glucocorticoids (both baseline and stress-induced) and androgens in free-living and un-manipulated adult vertebrates. HormoneBase (www.HormoneBase.org) currently includes \u3e6,580 entries from 476 species, reported in 648 publications from 1967 to 2015, and unpublished datasets. Entries are associated with data on the species and population, sex, year and month of study, geographic coordinates, life history stage, method and latency of hormone sampling, and analysis technique. This novel resource could be used for analyses of the function and evolution of hormone systems, and the relationships between hormonal variation and a variety of processes including phenotypic variation, fitness, and species distributions