angel insects; diversity; morphology; evolution; Polyneoptera

Zoraptera is a small and predominantly tropical insect order with an unresolved higher classification due to the extremely uniform external body morphology. We, therefore, conducted a multigene molecular phylogeny of extant Zoraptera and critically re-evaluated their morphological characters in order to propose a natural infraordinal classification. We recovered a highly-resolved phylogeny with two main clades representing major evolutionary lineages in Zoraptera, for which we propose family ranks. The two families exhibit striking differences in male genitalia and reproductive strategies. Each family contains two subclades (subfamilies) supported by several morphological synapomorphies including the relative lengths of the basal antennomeres, the number and position of metatibial spurs, and the structure of male genitalia. The newly proposed higher classification of Zoraptera includes the family Zorotypidae stat. revid. with Zorotypinae Silvestri, 1913 (Zorotypus stat. revid., Usazoros Kukalova-Peck and Peck, 1993 stat. restit.) and Spermozorinae subfam. nov. (Spermozoros gen. nov.), and Spriralizoridae fam. nov. with Spiralizorinae subfam. nov. (Spiralizoros gen. nov., Scapulizoros gen. nov., Cordezoros gen. nov., Centrozoros Kukalova-Peck and Peck, 1993, stat. restit., Brazilozoros Kukalova-Peck and Peck, 1993, stat. restit.), and Latinozorinae subfam. nov. (Latinozoros Kukalova-Peck and Peck, 1993, stat. restit.). An identification key and morphological diagnoses for all supraspecific taxa are provided.