The xerophilic genera Xerobiotus and Pseudohexapodibius (Macrobiotidae; Tardigrada): biodiversity, biogeography and phylogeny

The genera Xerobiotus and Pseudohexapodibius are xerophilic and characterized by reduced appendages and claws as adaptations to move between small interstices. To increase the knowledge of their biodiversity, biogeography and phylogeny, several specimens from European countries and Australia were analysed using an integrative approach, i.e. morphological, karyological and molecular studies (18S, 28S, cox1, cytb and ITS2). The phylogenetic position of Xerobiotus with respect to the three evolutionary lineages of Macrobiotus remains unchanged with respect to what was previously identified in the phylogeny of Macrobiotidae. The following new Xerobiotus species are described: Xerobiotus litus sp. nov., Xerobiotus arenosum sp. nov. and Xerobiotus reductus sp. nov. It is proposed that Macrobiotus naginae should be transferred to Xerobiotus (Xerobiotus naginae comb. nov.). Pseudohexapodibius degenerans clusters within Xerobiotus and shares morphological and genetic characters with this genus. Some discrepancies between genetic and phenotypic data are found among the analysed species. Even between the results obtained with analysed genes disagreements are found, with cox1 and cytb generally agreeing with phenotypic results more than ITS2. Genotypic and phenotypic data are useful tools for species identification, but they have to be evaluated critically to obtain reliable results.