First reports of Grania (Clitellata: Enchytraeidae) from Africa and South America: Molecular phylogeny and descriptions of nine new species

Abstract

In this article, our knowledge of the geographic distribution of Grania species is expanded by describing seven new species, Grania bekkoucheisp.nov., Grania brasiliensissp.nov., Grania capensissp.nov., Grania chilensissp.nov., Grania crypticasp.nov., Grania hinojosaisp.nov., and Grania simonaesp.nov., from poorly investigated regions of the Southern Hemisphere, plus two new species, Grania carolinensissp.nov. and Grania unithecasp.nov., from off the east coast of the USA. An immature achaetous specimen that we call Grania cf. levis was also included. The newly generated data were combined with a previously published data set in order to update the hypothesis of phylogenetic relationships among Grania species. All new species except G.crypticasp.nov. are supported by both morphological and molecular data. In addition, we find that G.chilensissp.nov. is structured in at least four distinct populations along the Chilean coast. The species described from South Africa form a monophyletic clade where two are morphologically indistinguishable but diverging in both mitochondrial and nuclear ribosomal genes, and thus we describe them as different species, G.bekkoucheisp.nov. and G.crypticasp.nov. Among the North Carolinian species, G.unithecasp.nov. is indicated as a close relative of Grania monospermathecaErséus & Lasserre, 1976, and G.carolinensissp.nov. is indicated as a close relative of G.cf.levis. The updated phylogeny is strongly concordant with geographical species distributions, thus supporting a low level of dispersal within this genus, as has previously been hypothesized.