Phylogenetic relationships and revised classification of the true bug infraorder Dipsocoromorpha (Insecta: Hemiptera: Heteroptera)

Abstract

Currently comprising only about 430 species, Dipsocoromorpha or minute litter bugs are one of the small infraorders of Heteroptera. They are classified into five morphologically distinct families--Ceratocombidae, Dipsocoridae, Hypsipterygidae, Schizopteridae and Stemmocryptidae--but relationships among and within these families are poorly understood owing to the lack of phylogenetic studies. A phylogenetic hypothesis based on combined molecular and morphological data is important to both evaluate and revise the higher-level classification and to explore the evolutionary history of morphological features including elytriform (or beetle-like) forewings, that seem to have evolved multiple times in this group, jumping devices, and the highly modified and diverse male genitalia. We here use a dataset combining Sanger-derived (~4500 bp; 108 taxa) and Illumina-generated (~7500 bp; 24 taxa) sequence data with a morphological matrix (159 characters) and taxon sampling that comprises all currently recognized family-, subfamily-and tribal-level taxa and comprehensive genus-level sampling to investigate phylogenetic relationships within litter bugs. Our results support the monophyly of Dipsocoromorpha, Schizopteridae and Dipsocoridae, whereas Ceratocombidae and the schizopterine subfamily "Ogeriinae" are polyphyletic and paraphyletic, respectively. A new classification is proposed that recognizes six families, including Trichotonannidae, stat.n., and two subfamilies each within the two larger families Ceratocombidae and Schizopteridae. Ancestral state reconstructions outline the complex evolutionary history of many morphological characters, including 15 independent origins of elytriform forewings, and at least five shifts in the degree of genitalic asymmetry.

Capítulo 25: Hemiptera Linnaeus, 1758

“…Fêmea com ovipositor bem desenvolvido (Štys 1995b). clado (Knyshov et al 2020). Assim, a família ficou apenas com Schizopterinae e Hypselosomatinae.…”

A synopsis of the numbers of testicular follicles and ovarioles in true bugs (Heteroptera, Hemiptera) – sixty-five years of progress after J. Pendergrast’s review

“…The small infraorder Dipsocoromorpha or minute litter bugs comprises ~ 430 species from 70 genera that are classified into six morphologically distinct families (Knyshov et al 2021). Data on the number of follicles and/or ovarioles are currently available for the families Dipsocoridae, Ceratocombidae, and Schizopteridae.…”

Taxonomic review of Kokeshia Miyamoto, 1960 from China, with description of ten new species (Hemiptera: Heteroptera: Schizopteridae)

“…Their tiny body size and cryptic habits cause them to be one of the least known groups in Heteroptera Latreille, 1810. This infraorder currently comprises 6 families and about 430 species worldwide (Štys 1995;Weirauch et al 2018;Knyshov et al 2020). Schizopteridae Reuter, 1891, the largest family of Dipsocoromorpha, had been divided into 3 subfamilies: Hypselosomatinae Esaki & Miyamoto, 1959, Ogeriinae Emsley, 1969 andSchizopterinae Reuter, 1891 by Emsley (1969), and this classification system was accepted by several subsequent authors Hill 2004;Weirauch & Štys 2014).…”