Phylogeny of Itaquasconinae in the light of the evolution of the flexible pharyngeal tube in Tardigrada

摘要

The family Hypsibiidae comprises four subfamilies, among which Itaquasconinae are the most abundant in genera. Although being morphologically diverse and cosmopolitan, itaquasconins are scarcely reported and their taxonomy is entangled. Here, we present a phylogeny based on three DNA markers and morphological clues obtained from a detailed scanning electron microscopy analysis of bucco-pharyngeal apparatuses of 25 species representing seven genera. Our study revealed that (a) the polyphyletic Adropion consists of at least three evolutionary lineages, with Guidettion gen. nov. being the sister group of all remaining itaquasconins; (b) stylet supports were independently lost in Astatumen and Insulobius gen. nov.; (c) pharyngeal tube annulation can be divided into two general types, simple and complex, and is stable at the genus level, but being homoplasious (similarly to the apophyses for the insertion of stylet muscles and the shape of the claw bases), it is not useful in inferring phyletic relationships within the subfamily; and (d) Astatumen and Platicrista contain numerous similar species, challenging the current taxonomic classification; moreover, (e) three new taxa are proposed to accommodate the newly recognised lineages: Guidettion gen. nov. (the former Adropion prorsirostre group), Insulobius gen. nov. (for a new Malayan species with a mixture of Astatumen and Itaquascon traits), and Raribius gen. nov. (former Itaquascon spp. with an elongated and narrowed pharyngeal tube terminating with a spherical pharynx). The simple type of annulation is synapomorphic to Hypsibiidae, with single reversals to the uniform buccal tube in Parascon (Itaquasconinae) and in Hypsibiinae. The pharyngeal tube evolved convergently several times, and is of different character in classes Heterotardigrada and Eutardigrada.