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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.

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Aim The study of cosmopolitan earthworms could be even more interesting than that of endemic species in revealing evolutionary processes. Previous research on the cosmopolitan worm Eiseniella tetraedra has indicated some phylogeographic structure among populations, but the factors responsible remain unresolved. We hypothesized that environmental factors and dispersal have shaped the distribution of the species' lineages. Location Spain and Portugal; Iberian Peninsula. Taxon Eiseniella tetraedra (Lumbricidae, Oligochaeta, Annelida). Methods We collected 739 specimens of Eiseniella tetraedra from 65 localities around the Iberian Peninsula between 2012 and 2016. We performed phylogenetic analysis (Bayesian Inference and maximum likelihood) using two mitochondrial (COI and 16S) and one nuclear marker (28S). Furthermore, we studied their genetic diversity and historic demography based on the COI gene. Correlations between genetic diversity and 22 environmental factors were tested. Results Eiseniella tetraedra showed high diversity in the Iberian Peninsula, with eight different lineages nested in two clades. We found lineages mostly restricted to the northern region, while others were distributed throughout the Iberian Peninsula. Habitat stability, that is, constant availability or lack of water, also correlated with genetic diversity. Thus, although no clear phylogeographic pattern was found, environmental factors (such as precipitation, temperature, and soil pH) and habitat stability influenced the distribution of genetic variability. Main Conclusions Eiseniella tetraedra is an earthworm with great genetic variability. We show that the ranges of species with high relative dispersal ability and ambiguous phylogeographic patterns may be better explained by influence of environmental conditions rather than specific geographic features. Adaptation to unstable conditions has been shown to confer more success on one of the two major genetic clades recovered, pointing to ecological plasticity as a key for evolutionary success.

The earthworm family Hormogastridae is a relatively diverse group in the Western Mediterranean basin. Since 1887, around thirty species have been described and assigned to four genera. However, from 2010 on, molecular, ecological and morphological studies have questioned the validity of those genera. Meanwhile, new species were discovered and assigned to them, pending a formal systematic revision; such a revision has been performed recently by integrating all the existing sources of information. The resulting classification consists of nine genera, including four newly erected ones. This revised systematic background is used in the current work as a base for the description of six new hormogastrid species: Diazcosinia sacrarium Marchán, Fernández, Díaz Cosín & Novo, sp. nov., Boucheona martae Marchán, Fernández & Díaz Cosín, sp. nov., Boucheona rosae Marchán, Díaz Cosín & Novo, sp. nov., Norana emiliae Marchán, Fernández, Díaz Cosín & Novo, sp. nov., Norana xylocerasi Marchán, Fernández, Díaz Cosín & Novo, sp. nov., and Norana beatrizae Marchán, Fernández, Díaz Cosín & Novo, sp. nov. Norana is a new replacement name for the preoccupied Nora Marchán, Fernández, Díaz Cosín & Novo, 2018. Likewise, Xanina is proposed to replace the preoccupied Xana Díaz Cosín, Briones & Trigo, 1989. We provide an overview of the currently known diversity of the different genera, and we further propose common names in several languages for some of the species of Hormogastridae.

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