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The expensive tissue hypothesis predicts a trade-off between investments in the brain and other energetically costly organs due to the costs associated with their growth and maintenance within the finite energy resources available. However, few studies address the strength of relationships between brain size and investments in precopulatory (ornaments and armaments) and postcopulatory (testes and ejaculates) sexual traits. Here, in a broad comparative study, we tested the prediction that the relationship between brain size and investment in sexual traits differs among taxa relative to the importance of sperm competition within them. We found that brain size was negatively correlated with sexual size dimorphism (SSD) in anurans and primates, and it tended to decrease with SSD in ungulates and cetaceans. However, brain size did not covary significantly with armaments (e.g., canine length, horn, antler, and muscle mass). Brain size was not correlated with postcopulatory sexual traits (testes and ejaculates). The intensity of covariance between brain size and precopulatory sexual traits decreased with increasing relative testis size.

Competition over mates is a powerful force shaping trait evolution. For instance, better cognitive abilities may be beneficial in male-male competition and thus be selected for by intrasexual selection. Alternatively, investment in physical attributes favoring male performance in competition for mates may lower the resources available for brain development, and more intense male mate competition would coincide with smaller brains. To date, only indirect evidence for such relationships exists, and most studies are heavily biased toward primates and other homoeothermic vertebrates. We tested the association between male brain size (relative to body size) and male-male competition across N = 30 species of Chinese anurans. Three indicators of the intensity of male mate competition—operational sex ratio (OSR), spawningsite density, and male forelimb muscle mass—were positively associated with relative brain size, whereas the absolute spawning group size was not. The relationship with the OSR and male forelimb muscle mass was stronger for the male than for the female brains. Taken together, our findings suggest that the increased cognitive abilities of larger brains are beneficial in male-male competition. This study adds taxonomic breadth to the mounting evidence for a prominent role of sexual selection in vertebrate brain evolution.

Sperm competition is often considered the primary selective force underlying the rapid and diversifying evolution of ejaculate traits. Yet, several recent studies have drawn attention to other forms of selection with the potential of exceeding the effects of sperm competition. Since ejaculates are complex, multivariate traits, it seems plausible that different ejaculate components vary in their responses to different selective pressures. Such information, however, is generally lacking as individual ejaculate traits tend to be studied in isolation. Here, we studied the macroevolutionary patterns of ejaculate volume, sperm number, sperm length and the proportion of viable normal sperm in response to varying levels of sperm competition, body size and the duration of female sperm storage in pheasants and allies (Phasianidae). Ejaculate volume, sperm number and sperm viability were all relatively higher in polygamous than in monogamous mating systems. However, whereas ejaculate volume additionally covaried with body size, sperm number instead increased with the female sperm-storage duration, in conjunction with a decrease in sperm length. Overall, our results revealed important details on how different forms of selection can jointly shape ejaculates as complex, composite traits.

The evolution of sperm quality and quantity is shaped by various selective processes, with sperm competition generally considered the primary selective agent. Particularly in external fertilizers, however, sperm limitation through gamete dispersal can also influence gamete investments, but empirical data examining this effect are limited. Here, we studied the relative importance of sperm competition and the spawning conditions in explaining the macroevolutionary patterns of sperm size and number within two taxa with external fertilization but differences in their reproductive biology. In frogs, sperm swim slowly but for up to hours as they penetrate the gelatinous egg coating, whereas fish sperm typically swim fast, are very short-lived (seconds to minutes), and often face a relatively higher risk of being moved away from the ova by currents. Our phylogenetic models and path analyses revealed different trajectories of ejaculate evolution in these two taxa. Sperm size and number responded primarily to variation in sperm competition in the anurans, but more strongly to egg number and water turbulence in the fishes. Whereas the results across anurans align with the general expectation that sexual selection is the main driver of ejaculate evolution, our findings across the fishes suggest that sperm limitation has been underappreciated.

Sexual selection theory predicts a trade-off between premating (ornaments and armaments) and postmating (testes and ejaculates) sexual traits, assuming that growing and maintaining these traits is costly and that total reproductive investments are limited. The number of males in competition, the reproductive gains from investing in premating sexual traits, and the level of sperm competition are all predicted to influence how males allocate their finite resources to these traits. Yet, empirical examination of these predictions is currently scarce. Here, we studied relative expenditure on pre- and postmating sexual traits among frog species varying in their population density, operational sex ratio, and the number of competing males for each clutch of eggs. We found that the intensifying struggle to monopolize fertilizations as more and more males clasp the same female to fertilize her eggs shifts male reproductive investment toward sperm production and away from male weaponry. This shift, which is mediated by population density and the associated level of male–male competition, likely also explains the trade-off between pre- and postmating sexual traits in our much broader sample of anuran species. Our results highlight the power of such a multilevel approach in resolving the evolution of traits and allocation trade-offs.

Body size-dependent mating pattern is thought to a consequence of the sexual selection on phenotypic characters. In the view, we investigated the mating patterns in Bufo andrewsi from three populations at different latitude in three Nature Reserves (i.e. Wanglang, Yele and Fengtongzhai) in China from early- March to late-May 2013 and 2014. The results showed that average body size did not differ significantly between amplectant and non-amplectant males within each population. We found a significantly positive relationship in body size between the amplectant males and amplectant females in Wanglang population, but not in Qiqiao and Yele populations. The strength of size-dependent mating was not correlated with the malebiased operational sex ratios (OSRs), suggesting that intensity of male-male competition at nights do not result in variation in size-dependent mating patterns in B. andrewsi among populations.

Sexual differences in morphological traits are widespread among animals. Theory predicts that dimorphism in secondary sexual characters evolves as a consequence of sexual selection. We investigated the intersexual difference in mass of forelimb muscles of the Andrew's toad, Bufo andrewsi, an elongated species inhabiting montane regions in western China. Our results showed that average body size of females was significantly larger than males. However, when the influence of body size was removed, the forelimb muscle mass of males significantly exceeded that of females, and total mass of forelimb muscles of amplectant males was significantly larger than that of non-amplectant males. These results suggest that the robustness of the forelimbs can allow amplectant males to retain a firm grip on the female in amplexus, which may aid resistance to inference by conspecific males. Our findings are consistent with the prediction that sexual selection favors large forelimb muscle mass in males.

In animals, body size is a prominent trait that generally positively affects ecological and reproductive success. Through field observations and experiments, we investigated the effects of two mechanisms of sexual selection on large male mating advantage in the Andrew's toad, Bufo andrewsi, a species widely distributed in western China. We observed a large male mating advantage in the field. Field data were corroborated by experiments in which large males mated more frequently than smaller males. However, in female preference tests, in which females could choose freely between two males differing in body size, choosing females showed no size preference while many females did not exert mate choice at all. We suggest that the large male mating advantage observed in the field and laboratory is caused by competition among males rather than by female choice.

In amphibians, theory predicts that male mate choice with respect to female body size can be expected to occur when female fecundity is related to body size and when the time and energy invested into one mating are relatively large. Based on experimental observations, we tested whether male mate choice occurs in a population of the Andrew’s toad (Bufo andrewsi), a species in which both assumptions are likely to be met. When a male B. andrewsi was placed with a gravid female and a non-gravid similar-sized female, the male did not discriminate between them. When two gravid females with distinct size differences were provided to a male, the male preferred the larger one. In an experiment in which two different-sized gravid females were put in two separate transparent cylinders to exclude potential chemical cues, males spent more time in proximity to the larger gravid females and jumped more frequently towards the larger gravid females than the smaller ones. These findings suggest that male B. andrewsi recognizes female body size, exhibits mate choice, and prefers to mate with larger females that provide greater reproductive potential.

Background: The degree of postcopulatory sexual selection, comprising variable degrees of sperm competition and cryptic female choice, is an important evolutionary\ud force to influence sperm form and function. Here we investigated the effects of\ud mating system and spawning location on the evolution of sperm morphology in 67\ud species of Chinese anurans. We also examined how relative testes mass as an\ud indicator of the level of sperm competition affected variation in sperm morphology\ud across a subset of 29 species.\ud Results: We found a significant association of mating system and spawning location\ud with sperm morphology. However, when removing the effects of body mass or\ud absolute testes mass for species for which such data were available, this effect became\ud non-significant. Consistent with predictions from sperm competition theory, we found\ud a positive correlation between sperm morphology and relative testes mass after taking\ud phylogeny into account.\ud Conclusions: Our findings suggest that sexual selection in Chinese anurans favors\ud longer sperm when the level of sperm competition is high. Pre-copulatory male-male\ud competition and spawning location, on the other hand, do not affect the evolution of\ud sperm morphology after taking body mass and absolute testes mass into account.

Background Sexual size dimorphism (SSD) is widespread and variable among animals. Sexual selection, fecundity selection and ecological divergence between males and females are the major evolutionary forces of SSD. However, the influences of mating system and habitat types on SSD have received little attention. Here, using phylogenetic comparative methods, we at first examine the hypotheses to that mating system (intensity of sexual selection) and habitat types affect significantly variation in SSD in anurans (39 species and 18 genera). Results Our data set encompass 39 species with female-biased SSD. We provide evidence that the effects of mating system and habitat types on SSD were non-significant across species, also when the analyses were phylogenetically corrected. Conclusions Contrast to the hypotheses, our findings suggest that mating system and habitat types do not play an important role in shaping macro-evolutionary patterns of SSD in anurans. Mating system and habitat types cannot explain the variation in SSD when correcting for phylogenetic effects.