Integrative systematic revision of a Mediterranean earthworm family: Hormogastridae (Annelida, Oligochaeta)

Abstract

The problem of reconciling earthworm taxonomy and phylogeny has shown advances with the application of molecular techniques, yet they have proven insufficient. Integrative systematics could solve this by combining multiple sources of evolutionary information. Relatively low diversity, restricted range and low nomenclatural conflict make Hormogastridae Michaelsen, 1900 a desirable target for an integrative systematics approach. The main systematic conflicts within this family are the polyphyly of the species Hormogaster pretiosa Michaelsen, 1899, the widespread presence of cryptic lineages, the lack of resolution of supraspecific relationships and the paraphyly of the genus Hormogaster Rosa, 1877 (found to be composed of four well-supported genus-level clades by molecular phylogenetic inference). This work integrates all the existing information by performing phylogenetic inference based on morphological, molecular and total evidence datasets, comparing their performance with the topology obtained by phylogenomic analyses. It also includes a comparative study of representatives of the main clades based on microcomputed tomography (µCT) reconstructions. The addition of morphological characters improved the resolution of the Hormogastridae tree; ancestral state reconstruction displayed the evolution of character states and provided morphological diagnoses for the genera within a new system, which incorporates information about ecological niches and biogeography.