The phylogeny of Ptiliidae (Coleoptera: Staphylinoidea) – the smallest beetles and their evolutionary transformationsстатья
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Аннотация:The smallest beetles and the smallest non-parasitic insects belong to the staphylinoid family Ptiliidae. Their adult body length
can be as small as 0.325 mm and is generally smaller than 1 mm. Here we address the phylogenetic relationships within the family using
formal analyses of adult morphological characters and molecular data, and also a combination of both for the frst time. Strongly supported
clades are Ptiliidae + Hydraenidae, Ptiliidae, Ptiliidae excl. Nossidium, Motschulskium and Sindosium, Nanosellini, and a clade comprising
Acrotrichis, Smicrus, Nephanes and Baeocrara. A group comprising Actidium, Oligella and Micridium + Ptilium is also likely monophyletic. Limulodes + Cephaloplectus, strongly supported as a clade (Cephaloplectus included only in morphological analysis), are deeply
nested within Ptiliidae in the morphology-only non-weighted and in the molecular analyses, but placed as sister to the remaining Ptiliidae
(excl. Nossidium, Motschulskium and Sindosium) after successive reweighting. We propose that Ptiliidae should be taxonomically divided
into two subfamilies: the most recently established Nossidiinae and Ptiliinae, the latter currently composed of seven tribes: Acrotrichini,
Cephaloplectini, Discheramocephalini, Nanosellini, Ptenidiini, Ptiliini and Ptinellini, although the monophyly and taxonomic status is still
uncertain in some cases. Important morphological innovations evolved in the stem group of Hydraenidae and Ptiliidae, including partly
internalized mouthparts suitable for saprophagy and sporophagy, a labral-mandibular locking device, a specifc elytral locking mechanism
with elongated alacristae, wings with fringes of hairs, and a very unusual spermathecal pump. A complex feature of Ptiliidae linked with
miniaturization is the transformation of the alae into “feather wings”, with a highly effcient fight mechanism unique in beetles. Nanosellini include the smallest ptiliid species and display features correlated with extremely small body size, such as simplifcation of the
endoskeleton of the head and thorax, far-reaching reduction of the wing venation, and a reduced number of antennomeres.