Simon P. Lailvaux

Performance capacity, fighting tactics and the evolution of life-stage male morphs in the green anole lizard (**Anolis carolinensis**)

The evolution of alternative male phenotypes is probably driven by male–male competition for access to reproductive females, but few studies have examined whether whole–organism performance capacities differ between male morphs, and if so whether any such differences affect fighting ability. We show how ontogenetic changes in performance and morphology have given rise to two distinct life–stage male morphs exhibiting different fighting tactics within the green anole lizard (Anolis carolinensis). Field studies show a bimodal distribution of adult males within a single population: larger ‘heavyweight’ males have relatively large heads and high bite forces for their size, whereas smaller ‘lightweight’ males have smaller heads and lower bite forces. In staged fights between size–matched heavyweight males, males with greater biting ability won more frequently, whereas in lightweight fights, males with greater jumping velocity and acceleration won more often. Because growth in reptiles is indeterminate, and the anole males examined are sexually mature, we propose that the heavyweight morph arose through selection against males with small heads and poor bite forces at the lightweight–heavyweight size transition. Our findings imply that one may not be able to predict male fighting success (and hence potential mating success) by examining aspects of male ‘quality’ at only one life stage.

A functional perspective on sexual selection: insights and future prospects

A large number of sexual selection studies have focused on examining the morphological and behavioural factors involved in male combat and female choice, such as whether large males achieve higher reproductive success compared with smaller males. However, until recently, the mechanistic reasons why such cues are linked to male dominance or female choice have been elusive. An emerging body of work shows that physiological and whole-organism performance capacities are important in individual reproductive success. Males with high performance or other physiological capacities (e.g. endurance, biting) often enjoy an advantage over males with poorer performance capacities during male–male contests. In contrast, few studies have examined links between performance and female choice. Here, we highlight recent key literature integrating sexual selection, performance and physiology. We also point to areas where a more rigorous investigation of underlying physiological processes may yield insights into sexual selection. In particular, we note that current progress in several important areas may be hampered by an inadequate physiological understanding of condition. We suggest a conceptual approach that may shed light on the physiological factors underlying condition, and we point out several other potentially important avenues for future research.

The Evolution of Performance‐Based Male Fighting Ability in Caribbean Anolis Lizards

Despite the empirical and theoretical attention paid to the role of sexual signals in resolving agonistic interactions between conspecific males, few studies have applied a comparative perspective, particularly across species that vary in combat intensity. We investigated the relative roles of a male sexual signal (dewlap size) and whole‐organism performance capacity (bite force) on male combat outcomes in nine species of Caribbean Anolis lizards that differ markedly in territoriality, as indicated by sexual size dimorphism. We found that (1) dewlap size was generally an honest signal of bite force in dimorphic but not less dimorphic species; (2) maximum bite force consistently predicted male combat success in dimorphic but not less dimorphic species; (3) in contrast to a priori predictions, dewlap size significantly predicted male combat success in less dimorphic but not dimorphic species; and (4) the incidence of biting but not dewlapping increases as species become more dimorphic. These findings suggest that more dimorphic (and hence more territorial) species escalate to biting during fights more readily compared with less territorial species. The ecological and behavioral qualities of species may therefore modify both the shape and the size of sexually selected traits as well as the nature of the information those traits convey.

Sex‐Based Differences and Similarities in Locomotor Performance, Thermal Preferences, and Escape Behaviour in the Lizard Platysaurus intermedius wilhelmi

Differences between sexes in physiological performance have received little attention in animals. We tested for sex differences in maximum sprint speed and maximal exertion over a range of temperatures in a population of Platysaurus intermedius wilhelmi lizards. We also examined sex‐based differences in selected temperature range, mean field body temperatures (Tb), and thermal activity limits. Finally, we conducted field studies to quantify male and female responses to a potential predator, which may be affected by their respective performance capabilities. Males were faster than females at all temperatures, and body size had no significant effect on sprint speeds. Males and females also selected similar Tbs when placed in a thermal gradient, but in the field, male lizards’ Tbs were different from those of the females. However, predicted sprint speeds for males and females at their field Tbs are similar. No significant differences were found between males and females with regard to maximal exertion. When approached in the field, adult male lizards took refuge significantly earlier than did adult females and also fled over shorter distances, suggesting that females rely on crypsis as an escape strategy.

Defining individual quality over lifetimes and selective contexts

Individual quality has been measured as a variety of different traits and in several different contexts. However, the implications of such measurements in terms of overall fitness are less straightforward than has generally been appreciated. Here we outline some key issues in this regard that have yet to be addressed. Specifically, we consider the importance of both variation in selection on individual and multivariate suites of traits, and of context-specific plasticity in allocation strategies. We argue that an explicit life-history perspective is crucial for understanding variation in quality, as both the strength and direction of selection and an individuals response to it can vary within a breeding season. Hence, ‘quality’ is not a static characteristic that can be measured by taking longitudinal measures of single traits across a population, but rather a dynamic, multivariate suite of traits that is dependent not only on the selective context, but also on the nature and intensity of selection operating at any given time. We highlight these points by considering recent research on selection and plasticity.

Sex differences in nutrient-dependent reproductive ageing

Evolutionary theories of aging predict that fitness‐related traits, including reproductive performance, will senesce because the strength of selection declines with age. Sexual selection theory predicts, however, that male reproductive performance (especially sexual advertisement) will increase with age. In both bodies of theory, diet should mediate age‐dependent changes in reproductive performance. In this study, we show that the sexes exhibit dramatic, qualitative differences in age‐dependent reproductive performance trajectories and patterns of reproductive ageing in the cricket Teleogryllus commodus. In females, fecundity peaked early in adulthood and then declined. In contrast, male sexual advertisement increased across the natural lifespan and only declined well beyond the maximum field lifespan. These sex differences were robust to deviations from sex‐specific dietary requirements. Our results demonstrate that sexual selection can be at least as important as sex‐dependent mortality in shaping the signal of reproductive ageing.

The Life History of Whole-Organism Performance

For almost 40 years, studies of whole-organism performance have formed a cornerstone of evolutionary physiology. Although its utility as a heuristic guide is beyond question, and we have learned much about morphological evolution from its application, the ecomorphological paradigm has frequently been applied to performance evolution in ways that range from unsatisfactory to inappropriate. More importantly, the standard ecomorphological paradigm does not account for tradeoffs among performance and other traits, nor between performance traits that are mediated by resource allocation. A revised paradigm that includes such tradeoffs, and the possible ways that performance and fitness-enhancing traits might affect each other, could potentially revivify the study of phenotypic evolution and make important inroads into understanding the relationships between morphology and performance and between performance and Darwinian fitness. We describe such a paradigm, and discuss the various ways that performance and key life-history traits might interact with and affect each other. We emphasize both the proximate mechanisms potentially linking such traits, and the likely ultimate factors driving those linkages, as well as the evolutionary implications for the overall, multivariate phenotype. Finally, we highlight several research directions that will shed light on the evolution and ecology of whole-organism performance and related life-history traits.

Performance is no proxy for genetic quality: trade-offs between locomotion, attractiveness, and life history in crickets

The genetic relationships among traits contributing to overall fitness are an important subject of inquiry because such relationships influence how suites of traits respond to selection. Within the field of sexual selection, these relationships are also of interest for assessing whether any given trait can be used as a proxy for total fitness. A growing number of studies have demonstrated close links between whole-organism performance traits and determinants of individual fitness; however, an understanding of the genetic relationships between performance and important aspects of genetic quality is currently lacking. We present the results of a quantitative genetic study in which we estimate covariation between a locomotor performance trait (maximal jumping ability), calling effort, sexual attractiveness, and life-history traits in male Teleogryllus commodus crickets. We show that the major axis of genetic variation (gmax) is characterized by a contrast between jump performance and life-history traits associated with mating success. Moreover, two additional axes of significant multivariate genetic variation exist, each characterized by strong contrasts among traits. These results argue against the existence of a single axis representing genetic quality, favoring instead the idea that resource allocation strategies shape multiple dimensions of genetic quality through trade-offs among key life-history traits, including performance.

Sexual conflict and the maintenance of multivariate genetic variation.

Mate choice should erode additive genetic variation in sexual displays, yet these traits often harbor substantial genetic variation. Nevertheless, recent developments in quantitative genetics have suggested that multivariate genetic variation in the combinations of traits under selection may still be depleted. Accordingly, the erosion and maintenance of variation may only be detectable by studying whole suites of traits. One potential process favoring the maintenance of genetic variance in multiple trait combinations is the modification of sexual selection via sexually antagonistic interactions between males and females. Here we consider how interlocus sexual conflict can shape the genetic architecture of male sexual traits in the cricket, Teleogryllus commodus. In this species, the ability of each sex to manipulate insemination success significantly alters the selection acting on male courtship call properties. Using a quantitative genetic breeding design we estimated the additive genetic variation in these traits and then predicted the change in variation due to previously documented patterns of sexual selection. Our results indicate that female choice should indeed deplete multivariate genetic variance, but that sexual conflict over insemination success may oppose this loss of variance. We suggest that changes in the direction of selection due to sexually antagonistic interactions will be an important and potentially widespread factor in maintaining multivariate genetic variation.

A Species-Level Phylogeny of Extant Snakes with Description of a New Colubrid Subfamily and Genus.

Background With over 3,500 species encompassing a diverse range of morphologies and ecologies, snakes make up 36% of squamate diversity. Despite several attempts at estimating higher-level snake relationships and numerous assessments of generic- or species-level phylogenies, a large-scale species-level phylogeny solely focusing on snakes has not been completed. Here, we provide the largest-yet estimate of the snake tree of life using maximum likelihood on a supermatrix of 1745 taxa (1652 snake species + 7 outgroup taxa) and 9,523 base pairs from 10 loci (5 nuclear, 5 mitochondrial), including previously unsequenced genera (2) and species (61). Results Increased taxon sampling resulted in a phylogeny with a new higher-level topology and corroborate many lower-level relationships, strengthened by high nodal support values (> 85%) down to the species level (73.69% of nodes). Although the majority of families and subfamilies were strongly supported as monophyletic with > 88% support values, some families and numerous genera were paraphyletic, primarily due to limited taxon and loci sampling leading to a sparse supermatrix and minimal sequence overlap between some closely-related taxa. With all rogue taxa and incertae sedis species eliminated, higher-level relationships and support values remained relatively unchanged, except in five problematic clades. Conclusion Our analyses resulted in new topologies at higher- and lower-levels; resolved several previous topological issues; established novel paraphyletic affiliations; designated a new subfamily, Ahaetuliinae, for the genera Ahaetulla, Chrysopelea, Dendrelaphis, and Dryophiops; and appointed Hemerophis (Coluber) zebrinus to a new genus, Mopanveldophis. Although we provide insight into some distinguished problematic nodes, at the deeper phylogenetic scale, resolution of these nodes may require sampling of more slowly-evolving nuclear genes.