Tardigrade Augean stables—a challenging phylogeny and taxonomy of the family Ramazzottiidae (Eutardigrada: Hypsibioidea)

Abstract

Abstract Tardigrade taxonomy is most often hindered by prevalent outdated species descriptions, lack of integrative redescriptions, scarce genetic information, and fragmentary sampling. Here, we diagnose the problems obscuring phylogenetic inference and the taxonomy of the cosmopolitan family Ramazzottiidae. We carried out the most extensive phylogenetic analysis of this family to date, with a considerable influx of new genetic data from poorly sampled regions of the world. We found two new distinct evolutionary lineages defined by distinct morphological traits (dorsal cuticular sculpturing and/or body shape), characterized by restricted geographic distributions, and we delineated them as new species complexes: the Neotropical Ramazzottius baumanni complex and the Afro-Oriental Ramazzottius szeptyckii complex. Nevertheless, we could not confidently ascertain the taxonomic status of both complexes due to: (i) the current state of the ramazzottiid taxonomy, with outdated and/or imprecise species descriptions with heterogeneous terminology and, in many cases, without accounting for intraspecific variation; (ii) the missing genetic information for key taxa; and (iii) the possible lack of monophyly of Cryoconicus and Ramazzottius as suggested by our results. In addition to diagnosing the problems of ramazzottiid phylogeny and systematics, we also propose possible solutions that could accelerate the progress in our understanding of the evolution of this group.