The canonical model of sex-chromosome evolution predicts that, as recombination is suppressed along sex chromosomes, gametologs will progressively differentiate, eventually becoming heteromorphic. However, there are numerous examples of homomorphic sex chromosomes across the tree of life. This homomorphy has been suggested to result from frequent sex-chromosome turnovers, yet we know little about which forces drive them. Here, we describe an extremely fast rate of turnover among 28 species of Ranidae. Transitions are not random, but converge on several chromosomes, potentially due to genes they harbour. Transitions also preserve the ancestral pattern of male heterogamety, in line with the ‘hot-potato’ model of sex-chromosome transitions, suggesting a key role for mutation-load accumulation in non-recombining genomic regions. The importance of mutation-load selection in frogs might result from the extreme heterochiasmy they exhibit, making frog sex chromosomes differentiate immediately from emergence and across their entire length.
Summary: Anurans exhibit high levels of growth‐mediated phenotypic plasticity in age and size at metamorphosis. Although temperature and food quality exert a strong influence on larval growth, little is known about the interacting effects of these factors on age and size at metamorphosis. Plasticity in growth rates, maximum larval mass, mass loss, larval period and size at metamorphosis was examined in Iberian Painted Frogs (Discoglossus galganoi Capula, Nascetti, Lanza, Bullini & Crespo 1985) under different combinations of temperature and diet quality. Temperature and diet had strong effects on the maximum size reached by tadpoles throughout the premetamorphic stages. Larval body mass varied inversely with temperature. The effect of diet depended on temperature; larvae fed on a ‘carnivorous’ diet (rich in protein and lipids) achieved a larger size than larvae offered an ‘herbivorous’ diet (rich in carbohydrates) at 17 °C but not at 12 or 22 °C. Larval period was insensitive to diet composition, and varied only with temperature. Primarily the interacting effects of food quality and temperature affected size at metamorphosis. Size at metamorphosis varied inversely with temperature under the plant‐ and the animal‐based diets. However, the carnivorous diet resulted in bigger metamorphs at 17 and 22 °C, but did not influence final mass at 12 °C. Maximum size over the larval period explained most of the variation in mass loss after the premetamorphic growing phase.
The genetic architecture of speciation, i.e., how intrinsic genomic incompatibilities promote reproductive isolation (RI) between diverging lineages, is one of the best-kept secrets of evolution. To directly assess whether incompatibilities arise in a limited set of large-effect speciation genes, or in a multitude of loci, we examined the geographic and genomic landscapes of introgression across the hybrid zones of 41 pairs of frog and toad lineages in the Western Palearctic region. As the divergence between lineages increases, phylogeographic transitions progressively become narrower, and larger parts of the genome resist introgression. This suggests that anuran speciation proceeds through a gradual accumulation of multiple barrier loci scattered across the genome, which ultimately deplete hybrid fitness by intrinsic postzygotic isolation, with behavioral isolation being achieved only at later stages. Moreover, these loci were disproportionately sex linked in one group (Hyla) but not in others (Rana and Bufotes), implying that large X-effects are not necessarily a rule of speciation with undifferentiated sex chromosomes. The highly polygenic nature of RI and the lack of hemizygous X/Z chromosomes could explain why the speciation clock ticks slower in amphibians compared to other vertebrates. The clock-like dynamics of speciation combined with the analytical focus on hybrid zones offer perspectives for more standardized practices of species delimitation. Significance Reproductive isolation is instrumental to the formation of new species (speciation), but it remains largely enigmatic how many incompatibilities are required to prevent hybridization and where they lie across the genome. By studying patterns of admixture in amphibian hybrid zones, we found that reproductive isolation is initiated by numerous small-effect incompatibilities scattered across the genome rather than concentrated in a few important genes. Unlike mammals and birds, in which Y/W degeneracy is a major cause of hybrid dysfunctions, the undifferentiated sex chromosomes of amphibians do not always host more genetic incompatibilities than other chromosomes. These combined results might explain why amphibian speciation is relatively slow, and its clock-like dynamics offer practical perspectives to categorize evolutionary lineages into species or subspecies.