Revised Cornechiniscus (Heterotardigrada) and new phylogenetic analyses negate echiniscid subfamilies and tribes

Echiniscidae are undoubtedly the most thoroughly studied lineage of the class Heterotardigrada. Recently, new subfamilies and tribes grouping echiniscid genera based on traditionally recognized morphological clues have been proposed. Here, by integrative analyses of morphology and DNA sequences of numerous populations of a rare genus Cornechiniscus, we show that this hypothesized classification is artificial. Specifically, we demonstrate that Echiniscinae are diphyletic, as Bryodelphax forms a distinct phyletic lineage within Echiniscidae, and Pseudechiniscinae are polyphyletic, with Mopsechiniscus being indirectly related to Pseudechiniscus, which is closer to the Echiniscus-like genera than to other genera with pseudosegmental plates. Consequently, the subfamilies and tribes are considered as unsupported from the phylogenetic and morphological point of view. The genus Cornechiniscus is revised, and the phenotypic diagnoses of several species are updated thanks to new rich material from Africa, Asia and Europe. Cornechiniscus imperfectus sp. nov. is described from mountains of Kyrgyzstan, being the second appendaged species within the genus and the third known to exhibit dioecy. A taxonomic key to the genus is provided. Systematic positions of Acanthechiniscus and Multipseudechiniscus are also discussed. Acanthechiniscus goedeni is confirmed to be a member of the genus Acanthechiniscus.