An increase in ecological opportunities, either through changes in the environment or acquisition of new traits, is frequently associated with an increase in species and morphological diversification. However, it is possible that certain ecological settings might prevent lineages from diversifying. Arboreality evolved multiple times in vipers, making them ideal organisms for exploring how potentially new ecological opportunities affect their morphology and speciation regimes. Arboreal snakes are frequently suggested to have a very specialized morphology, and being too large, too small, too heavy, or having short tails might be challenging for them. Using trait-evolution models, we show that arboreal vipers are evolving towards intermediate body sizes, with longer tails and more slender bodies than terrestrial vipers. Arboreality strongly constrains body size and circumference evolution in vipers, while terrestrial lineages are evolving towards a broader range of morphological variants. Trait-dependent diversification models, however, suggest similar speciation rates between microhabitats. Thus, we show that arboreality might constrain morphological evolution but not necessarily affect the rates at which lineages generate new species.
Vipers are among the most misunderstood and persecuted animals. They occupy most terrestrial ecosystems around the world, often at high population densities. However, certain aspects of their biology (e.g., low fecundity and slow growth) have resulted in vipers being disproportionately threatened by extinction. Despite increased extinction risk, relatively little is known about viper biology, severely limiting the development and implementation of conservation initiatives. Here, we review the conservation status of vipers globally, map species richness, and develop three indices (one reactive; one proactive; one combined index emphasising irreplaceable species facing severe threats) to identify species for which conservation action should be prioritised. Moreover, we map species richness weighted by each index to identify regions for conservation prioritisation. We ranked prioritisation scores for all species for which data were available. In doing so we identify species for which valuable data are missing and that should be prioritised for research. We additionally show that 17 species, currently listed as Not Assessed or Data Deficient by the IUCN, score sufficiently high on our Threat Index to be considered as Threatened in the future. We identify multiple regions for which viper conservation action should be prioritised. These areas broadly correlate with species richness patterns, suggesting that species richness may be an effective proxy for conservation planning. Finally, we discuss the major gaps in knowledge, as well as the major threats facing vipers.
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