Speciation, phylogeography and evolution of life history and morphology in plethodontid salamanders of the Eurycea multiplicata complex

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Abstract
Understanding the complex interactions among environment, genotype and ontogeny in determining organismal phenotypes is cental to many biological disciplines. The Eurycea multiplicata complex, endemic to the Interior Highlands (Ozark Plateau and Ouachita Mountains) of eastern North America, comprises a diverse radiation of paedomorphic surface-dwelling ( E. tynerensis ), metamorphic surface-dwelling ( E. multiplicata multiplicata and E. m. griseogaster ) and metamorphic subterranean ( Typhlotriton spelaeus ) hemidactyli- ine plethodontid salamanders. Portions of two mitochondrial genes, cytochrome- b and NADH dehydrogenase-4, totalling 1818 base pairs (bp) were sequenced for 70 ingroup individuals plus numerous outgroup taxa, to examine the biogeography and relationships among these morphologically disparate species. Results show the E. multiplicata complex to be monophyletic, with its two most divergent clades corresponding to geography, not morphology or life history. Transforming surface-dwelling populations from the Ouachi- tas ( E. m . multiplicata ) are sister to the Ozark taxa, including paedomorphic surface- dwelling ( E. tynerensis ), subterranean ( T. spelaeus ) and transforming surface-dwelling salamanders assigned to the 'subspecies' E. m. griseogaster . Among Ozark taxa T. spelaeus (deeply nested within Eurycea ) is sister to a clade that includes E. m. griseogaster and E. tynerensis . Current taxonomy suggests that paedomorphic populations ( E. tynerensis ) from the western Ozarks are distinct from nearby transforming populations ( E. m. griseogaster ). However, paedomorphic and transforming salamanders do not form reciprocally mono- phyletic groups and many populations share almost identical haplotypes. Ancestral state reconstruction of life history traits shows that paedomorphosis arose independently from three to nine times. Most populations are either completely paedomorphic or completely transforming. This suggests that local habitat parameters strongly influence life history mode in this complex, either facultatively or by selection for particular genotypes.