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 pbylogenetically-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 larg- er 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.

Trait allometries generate super-honesty in Anolis dewlaps and may underlie sexual dimorphism

Whether or not sexually selected traits consistently exhibit positive allometry(i.e.are disproportionately large in larger individuals)is an ongoing debate.Multiple models and exceptions to this rule suggest that the underlying drivers of sexual trait allometry are nuanced.Here,we compare allometries of sexual and non-sexual traits of a species(Anolis aquaticus)within a well-studied lizard genus to test the competing hypotheses that sexual traits are,or are not,defined by positive allometry.We further consider the relationships of trait functions,which are relatively well understood in the genus Anolis,and allometry to identify potential drivers of allometric patterns.In particular,we explore how trait allometries interact to influence total organism function and generate sexual dimorphism.We quantified size(of targeted traits)and color of a sexual signal(the dewlap)in Anolis aquaticus in the field.The dewlap conveyed information relevant to intra-sexual combat and exhibited positive allometry.Overall,our results suggest that using single-trait allometries as indicators of past selection provides only an incomplete understanding of trait evolution.Although the function of positive allometry in some individual sexual signals(e.g.those conveying“super-honest”information)may be straightforward,we illustrate how scaling relationships interact synergistically to influence the function of phenotypes and propose avenues for future research.

Visual ＂playback＂ of colorful signals in the field supports sensory drive for signal detectability

Colorful visual signals are important systems for investigating the effects of signaling environ ments and receiver physiology on signal evolution as predicted by the sensory drive hypothesis. Support for the sensory drive hypothesis on color signal evolution is mostly based on document ing correlations between the properties of signals and habitat conditions under which the signals are given （i.e., a correlational approach） and less commonly on the use of mathematical models that integrate representations of visual environments, signal properties, and sensory systems （i.e., a functional approach）. Here, we used an experimental approach in the field to evaluate signal efficacy of colorful lizard throat fans called dewlaps that show geographic variation in the lizard Anolis cristatellus. We used a remote controlled apparatus to display ＂fake dewlaps＂ to wild lizards to test for adaptive divergence in dewlap brightness （i.e., perceived intensity） among populations in situ. We found evidence of local adaptation in dewlap brightness consistent with the sensory drive hypothesis. Specifically, dewlaps that had the brightness characteristics of local lizards were more likely to be detected than those with the brightness characteristics of nonlocal lizards. Our findings indicate that simplified mathematical representations of visual environments may allow robust estimates of relative detectability or conspicuousness in natural habitats. We have shown the feasibility of evaluating color signal efficacy experimentally under natural conditions and dem onstrate the potential advantages of presenting isolated components of signals to an intended re ceiver to measure their contribution to signal function.

Dewlap size in male water anoles associates with consistent inter-individual variation in boldness

Male sexually selected signals can indicate competitive ability by honestly signaling fitnessrelevant traits such as condition or performance. However, behavior can also influence contest outcomes;in particular, boldness often predicts dominance rank and mating success. Here, we sought to determi ne whether male ornament size is associated with consiste nt in dividual differences in boldness in water anoles Anol is aquaticus. We measured the relative size of the dewlap, a flap of skin under the chin that is a sexually selected ornament in Anolis lizards, and tested for associations with responses to a novel and potentially risky environment: time to emerge from a refuge into an arena and number of head sea ns post-emergence. We found that in dividuals con siste ntly differed in both time to emerge and head sea nning (i.e., in dividual resp on ses were repeatable), and that dewlap size was negatively related to nu mber of head sea ns. This suggests that orname nt size could indicate male boldness if seanning represents antipredator vigilance. We found that males that had larger relative dewlaps were also in better body condition, but boldness (i.e., head scann ing) was not related to condition. Lastly, we found con siste nt d iff ere nces in behavior betwee n trials, showing that anoles were becoming habituated or sensitized to the testing arena. Overall, our study shows that in additi on to indicating con dition and perform a nee, dewlap size could also honestly indicate male boldness in Anolis lizards.