Hydromantes

Abstract: A recently presented model of tongue projection dynamics is used to generate a series of predictions concerning morphologies to be expected under selection for increased distance of projection, increased speed of projection, and increased directional versatility. A general understanding of biomechanical events and the model are used as points of departure for making specific predictions concerning details of structure in skeletal, muscular and connective tissue components of the tongue and associated structures. Comparative methods are used to examine these predictions in the genera of plethodontid salamanders. These salamanders are known to project their tongues to different degrees, and this knowledge is used to test the hypotheses concerning morphological specialization. Three distinct groups of plethodontid salamanders have evolved specializations for long distance projection, and these genera differ from one another in important ways in respect to specific character complexes. For example, the tropical genera and Hydromantes use CBII as the major force transmission element in the skeleton, while Eurycea and its allies use CBI in this role. Hydromantes differs from both in having a uniquely proportioned and structured hyobranchial skeleton and associated musculature. Less extreme specializations for tongue projection are found in different combinations in three other groups. Finally, two distinct groups of generalized species having only limited tongue projection capabilities are recognized, each having a unique complex of inter-related features. Each of these eight groups is recognized and characterized as a functional mode, and hypotheses concerning the biomechanical meaning of the character complexes of each are formulated.

Abstract: Deep phylogenetic relationships of the largest salamander family Plethodontidae have been difficult to resolve, probably reflecting a rapid diversification early in their evolutionary history. Here, data from 50 independent nuclear markers (total 48,582 bp) are used to reconstruct the phylogeny and divergence times for plethodontid salamanders, using both concatenation and coalescence-based species tree analyses. Our results robustly resolve the position of the enigmatic eastern North American four-toed salamander (Hemidactylium) as the sister taxon of Batrachoseps + Tribe Bolitoglossini, thus settling a long-standing question. Furthermore, we statistically reject sister taxon status of Karsenia and Hydromantes, the only plethodontids to occur outside the Americas, leading us to new biogeographic hypotheses. Contrary to previous long-standing arguments that plethodontid salamanders are an old lineage originating in the Cretaceous (more than 90 Ma), our analyses lead to the hypothesis that these salamanders are much younger, arising close to the K-T boundary (~66 Ma). These time estimates are highly stable using alternative calibration schemes and dating methods. Our data simulation highlights the potential risk of making strong arguments about phylogenetic timing based on inferences from a handful of nuclear genes, a common practice. Based on the newly obtained timetree and ancestral area reconstruction results, we argue that (i) the classic "Out of Appalachia" hypothesis of plethodontid origins is problematic; (ii) the common ancestor of extant plethodontids may have originated in northwestern North America in the early Paleocene; (iii) origins of Eurasian plethodontids likely result from two separate dispersal events from western North America via Beringia in the late Eocene (~42 Ma) and the early Miocene (~23 Ma), respectively.