Molecular phylogeographic analyses and species delimitations reveal that Leopoldamys edwardsi (Rodentia: Muridae) is a species complex

Abstract

Leopoldamys edwardsi is a species with wide distribution ranges in southern China but is not discussed in studies on geographic variation and species differentiation. We used 2 mitochondrial (Cytb, CO1) and 3 nuclear (GHR, IRBP and RAG1) genes to clarify species phylogeography and geographical differentiation. Maximum likelihood (ML) and Bayesian phylogenetic inference (BI) trees consistently indicated that L. edwardsi is a species complex containing 3 main lineages with high Kimura-2-parameter (K2P) divergences (i.e. lineages L_N , L_S and L_HN ) found in the northern and southern China and Hainan Island, respectively. The 3 species delimitation methods, automated barcoding gap discovery, Bayesian poisson tree process analysis and Bayesian phylogenetics and phylogeography, consistently supported the existence of cryptic species. Divergence times among the main lineages were inferred to be during the Pleistocene, with L_HN /L_S split at 1.33 Ma and L_N /(L_HN +L_S ) at 2.61 Ma; the diversifications of L. edwardsi complex might be caused by the rapid uplifts of Tibetan Plateau, paleoclimate change and complex topography. The divergence between L_HN and L_S was probably related to the separation of Hainan Island from the mainland via the formation of the Qiongzhou Strait. Lineages L_N and (L_S +L_HN ) likely diverged due to the Wuyi-Nanling mountain range forming a dispersal barrier. Our results suggested that L. edwardsi complex contains at least 3 distinct species: L_HN represents L. hainanensis, endemic to Hainan Island and previously considered as a subspecies L. e. hainanensis; L_S represents a cryptic species distributed throughout the southern Chinese continent; and L_N represents the nominotypical species L. edwardsi.

Keywords: Leopoldamys edwardsi complex; Leopoldamys hainanensis; ancient hybridization; cryptic species; species delimitation.