Frogs and toads (Anura) exhibit some of the most diverse parental strategies in vertebrates. Identifying the evolutionary origins of parenting is fundamental to understanding the relationships between sexual selection, social evolution and parental care systems of contemporary Anura. Moreover, parenting has been hypothesized to allow the invasion of terrestrial habitats by the ancestors of terrestrial vertebrates. Using comprehensive phylogenetic analyses of frogs and toads based on data from over 1000 species that represent 46 out of 55 Anura families, we test whether parental care is associated with terrestrial reproduction and several life-history traits. Here, we show that both the duration of care and offspring protection by males and females have coevolved with terrestrial reproduction. Sexual size dimorphism is also related to care, because the large male size relative to female size is associated with increased paternal care. Furthermore, increased egg size and reduced clutch volume are associated with increased care in bivariate but not in multivariate analyses, suggesting that the relationships between care, egg size and clutch volume are mediated by terrestrial reproduction. Taken together, our results suggest that parenting by males and females has coevolved, and complex parenting traits have evolved several times independently in Anura in response to breeding in terrestrial environments.
Vipera ursinii graeca is a restricted-range, endemic snake of the Pindos mountain range in the southwestern Balkans. The subspecies was previously reported from eight localities in Greece and one locality in southern Albania. We used species distribution modelling based on climate data from known localities in Greece to estimate the potential distribution of the subspecies. The model predicted suitable areas for eleven mountains in southern Albania, which we visited in ten field expeditions in four years. Based on 78 live individuals and 33 shed skins, we validated the presence of the snake on eight of the eleven mountains. Six populations (Dhëmbel, Llofiz, Griba, Shendelli, Tomorr and Trebeshinë Mountains) are reported here for the first time. Morphological characters undoubtedly supported that all individuals found at these new localities belong to V. u. graeca. Genetic analysis of mitochondrial DNA sequences also confirmed the identity of the snakes as V. u. graeca and a low number of identified haplotypes suggested low genetic variability among populations despite significant spatial isolation. All localities were subalpine-alpine calcareous meadows above 1600 m. These high montane habitats are separated by deep valleys and are threatened by overgrazing, soil erosion, and a potential increase in the elevation of the tree line due to climate change. Our surveys increased the number of known populations by 60% and the known geographical range of the subspecies by approximately 30%. Our study serves as a baseline for further ecological research and for conservation measures for one of the least known European viperid snakes.
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