Variation in ecological selection pressures has been implicated to explain variation in brain size and architecture in fishes, birds and mammals, but little is known in this respect about amphibians. Likewise, the relative importance of constraint vs. mosaic hypotheses of brain evolution in explaining variation in brain size and architecture remains contentious. Using phylogenetic comparative methods, we studied interspecific variation in brain size and size of different brain parts among 43 Chinese anuran frogs and explored how much of this variation was explainable by variation in ecological factors (viz. habitat type, diet and predation risk). We also evaluated which of the two above-mentioned hypotheses best explains the observed patterns. Although variation in brain size explained on average 80.5% of the variation in size of different brain parts (supporting the constraint hypothesis), none of the three ecological factors were found to explain variation in overall brain size. However, habitat and diet type explained a significant amount of variation in telencephalon size, as well in three composite measures of brain architecture. Likewise, predation risk explained a significant amount of variation in bulbus olfactorius and optic tecta size. Our results show that evolution of anuran brain accommodates features compatible with both constraint (viz. strong allometry among brain parts) and mosaic (viz. independent size changes in response to ecological factors in certain brain parts) models of brain size evolution.

Brain size differs substantially among species, and several hypotheses have been proposed to explain the evolution of brain size. Because the brain is among the most energetically expensive organs in the vertebrate body, trade-offs have been hypothesized to exert constraints on brain size evolution. Prominently, the expensive tissue hypothesis (ETH) proposes that reducing the size of another expensive organ, such as the gut, should compensate for the cost of a large brain. But energetic constraints may also drive covariation between the brain and other costly traits-such as body maintenance, locomotion, or reproduction-as formulated in the energy trade-off hypothesis. To date, these hypotheses have mainly been tested in homeothermic animals and within the ectothermic animals, primarily in fishes. Here, we undertake a comparative test of the interplay between energetic limitations and brain size evolution within amphibians. After controlling for phylogenetic relationships and body size, we find a negative correlation between brain mass and the length of the digestive tract within 30 species of anurans. We further find that the evolution of large brain size is accompanied by an increase in female reproductive investment into egg size. Our results suggest that the evolution of brain size follows general patterns across vertebrate clades.

The Expensive Brain Framework (EBF) and the Cognitive Buffer Hypothesis are commonly used to explain inter-and intraspecific variation in brain size and brain architecture. Using Andrew's toad (Bufo andrewsi) as a model species, we investigated whether brain attributes in amphibians are shaped by the interplay between age and the length of the activity season as determined by altitude and latitude. We found significant differences in absolute and relative brain volume across 12 populations. Supporting the EBF, we observed a positive correlation between absolute and relative brain volume with season length. In particular, populations experiencing long seasons were characterized by having relatively large olfactory nerve and optic tecta. Relative, but not absolute, brain volume was also positively correlated with individual age. When accounting for the effects of body size and brain volume, however, the size of different brain structures (olfactory nerve, olfactory bulbs, telencephalon, optic tecta and cerebellum) was independent from age. Taken together, our findings are largely in agreement with the EBF to determine brain size variation in Andrew's toad.

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

BackgroundEnvironmental variation associated with season length is likely to promote differentiation in life-history traits, but has been little studied in natural populations of ectotherms. We investigated patterns of variation in egg size, clutch size, age at sexual maturity, maximum age, mean age, growth rate and adult body size in relation to growth season length among 17 populations of Andrew’s toad (Bufo andrewsi) at different latitudes and altitudes in the Hengduan Mountains, western China.ResultsWe found that egg size, age at sexual maturity, and mean age increased with decreasing length of the growth season, whereas clutch size showed a converse cline. Body size did not increase with decreasing length of the growth season, but was tightly linked to lifetime activity (i.e. the estimated number of active days during lifetime). Males and females differed in their patterns of geographic variation in growth rates, which may be the result of forces shaping the trade-off between growth and reproduction in different environments.ConclusionsOur findings suggest that growth season plays an important role in shaping variation in life-history traits in B. andrewsi across geographical gradients.Electronic supplementary materialThe online version of this article (doi:10.1186/s12983-016-0138-0) contains supplementary material, which is available to authorized users.

Natural selection is a major force in the evolution of vertebrate brain size, but the role of sexual selection in brain size evolution remains enigmatic. At least two opposing schools of thought predict a relationship between sexual selection and brain size. Sexual selection should facilitate the evolution of larger brains because better cognitive abilities may aid the competition for mates. However, it may also restrict brain size evolution due to energetic trade‐offs between brain tissue and sexually selected traits. Here, we examined the patterns of selection on brain size and brain anatomy in male anurans (frogs and toads), a group where the strength of sexual selection differs markedly among species, using a phylogenetically controlled generalized least‐squared (PGLS) regression analyses. The analysis revealed that in 43 Chinese anuran species, neither mating system, nor type of courtship, or testes mass was significantly associated with relative brain size. While none of those factors related to the relative size of olfactory nerves, optic tecta, telencephalon, and cerebellum, the olfactory bulbs were relatively larger in monogamous species and those using calls during courtship. Our findings support the mosaic model of brain evolution and suggest that while the investigated aspects of sexual selection do not seem to play a prominent role in the evolution of brain size of anurans, they do impact their brain anatomy.

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Theory predicts that the degree of testes asymmetry should be positively correlated with male body condition in species with directional testis asymmetry. We tested this prediction in Rhacophorus omeimontis, a species in which females mate with more than one male. Our results showed that the treefrogs did not exhibit the absence of directional asymmetry in testis size, but rather the occurrence of fluctuating asymmetry. Moreover, we also tested differences in body size, body mass, testis mass, testis asymmetry, and sperm size among initially paired, jointly paired, and unpaired males. We found that body size and mass, testis mass, testis asymmetry and sperm length did not differ among the three male types. Testis mass showed a positive relationship with soma mass, but the correlations between the extent of fluctuating testis asymmetry and sperm length, and between testis mass and sperm length were not significant. Our data suggest that testes size and sperm length do not play an important role in determining male mating success in the presence of sperm competition.