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.

中文翻译：

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真虫亚目双翅目（昆虫纲：半翅目：异翅目）的系统发育关系和修订分类

目前仅包括约 430 种，Dipsocoromorpha 或微小的垃圾虫是异翅目的小型亚目之一。它们被分为五个形态上不同的科 - Ceratocombidae，Dipsocoridae，Hypsipterygidae，Schizopteridae和Stemmocryptidae - 但由于缺乏系统发育研究，这些科之间和内部的关系知之甚少。基于组合分子和形态学数据的系统发育假说对于评估和修改更高级别的分类以及探索包括鞘翅形（或甲虫状）前翅在内的形态特征的进化历史非常重要，这些特征似乎在此进化了多次群体，跳跃装置，以及高度修饰和多样化的男性生殖器。我们在这里使用一个结合了 Sanger 衍生的数据集（~4500 bp；108 个分类单元）和 Illumina 生成的（~7500 bp；24 个分类单元）序列数据，具有形态矩阵（159 个字符）和分类单元采样，包括所有当前公认的科、亚科和部落级分类单元和综合属级抽样调查垃圾虫内的系统发育关系。我们的结果支持 Dipsocoromorpha、Schizopteridae 和 Dipsocoridae 的单系，而 Ceratocombidae 和裂翅目亚科“Ogeriinae”分别是多系和并系的。提出了一种新的分类，可识别六个科，包括滴虫科、亚科和部落级分类群和全面的属级抽样，以研究垃圾虫内的系统发育关系。我们的结果支持 Dipsocoromorpha、Schizopteridae 和 Dipsocoridae 的单系，而 Ceratocombidae 和裂翅目亚科“Ogeriinae”分别是多系和并系的。提出了一种新的分类，可识别六个科，包括滴虫科、亚科和部落级分类群和全面的属级抽样，以研究垃圾虫内的系统发育关系。我们的结果支持 Dipsocoromorpha、Schizopteridae 和 Dipsocoridae 的单系，而 Ceratocombidae 和裂翅目亚科“Ogeriinae”分别是多系和并系的。提出了一种新的分类，可识别六个科，包括滴虫科、统计信息 ，以及两个较大的角翅目和裂翅目科中的两个亚科。祖先状态重建概述了许多形态特征的复杂进化历史，包括 15 个独立的鞘翅状前翅起源，以及至少五个生殖器不对称程度的变化。