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.
BackgroundWith 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).ResultsIncreased 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.ConclusionOur 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.
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.
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.
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 (T(b)), 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 T(b)'s when placed in a thermal gradient, but in the field, male lizards' T(b)'s were different from those of the females. However, predicted sprint speeds for males and females at their field T(b)'s 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.
SummaryEvolutionary theories of aging predict that fitnessrelated 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.
Summary1. In many animals, the size of secondary sexual ornaments is known to be related to the probability of victory in fights between males, and hence to fighting ability. However, few studies have attempted to link fighting ability to any physical performance measures. 2. Here we show that horn size in the dung beetle Euoniticellus intermedius accurately predicts two types of whole-organism performance, independent of body size, that are likely to play an important role in male contests: the force required to pull a beetle out of a tunnel, and the distance the beetle was able to run before exhaustion (maximum exertion). 3. Body length is also a statistically significant predictor of pulling force, but not of exertion, suggesting that horn size is a more reliable predictor of performance than body size alone, a result that is consistent with a previous finding that horn size becomes more important in determining victory in male-male contests as body size increases. 4. This study is the first to establish direct links between whole-organism performance abilities, male armaments and fighting ability in the same species. Our findings suggest that physiological performance capacities are important factors underlying the evolution of signal expression in E. intermedius , and should be considered in future studies of the evolution of animal signalling.
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