A comprehensive molecular phylogeny of Geometridae (Lepidoptera) with a focus on enigmatic small subfamilies

Our study aims to investigate the relationships of the major lineages within the moth family Geometridae, with a focus on the poorly studied Oenochrominae-Desmobathrinae complex, and to translate some of the results into a coherent subfamilial and tribal level classification for the family. We analyzed a molecular dataset of 1,206 Geometroidea terminal taxa from all biogeographical regions comprising up to 11 molecular markers that includes one mitochondrial (COI) and 10 protein-coding nuclear gene regions (wingless, ArgK, MDH, RpS5, GAPDH, IDH, Ca-ATPase, Nex9, EF-1alpha, CAD). The molecular data set was analyzed using maximum likelihood as implemented in IQ-TREE and RAxML. We found high support for the subfamilies Larentiinae, Geometrinae and Ennominae in their traditional scopes. Sterrhinae becomes monophyletic only if <i>Ergavia</i> Walker, <i>Ametris</i> Hübner and <i>Macrotes</i> Westwood, which are currently placed in Oenochrominae, are formally transferred to Sterrhinae. Desmobathrinae and Oenochrominae are found to be polyphyletic. The concepts of Oenochrominae and Desmobathrinae required major revision and, after appropriate rearrangements, these groups also form monophyletic subfamily-level entities. Oenochrominae <i>s.str</i>. as originally conceived by Guenée is phylogenetically distant from <i>Epidesmia</i> and its close relatives. The latter is hereby described as the subfamily Epidesmiinae Murillo-Ramos, Sihvonen & Brehm, <b>subfam. nov.</b> Epidesmiinae are a lineage of "slender-bodied Oenochrominae" that include the genera <i>Ecphyas</i> Turner, <i>Systatica</i> Turner, <i>Adeixis</i> Warren, <i>Dichromodes</i> Guenée, <i>Phrixocomes</i> Turner, <i>Abraxaphantes</i> Warren, <i>Epidesmia</i> Duncan & Westwood and <i>Phrataria</i> Walker. Archiearinae are monophyletic when <i>Dirce</i> and <i>Acalyphes</i> are formally transferred to Ennominae. We also found that many tribes were para- or polyphyletic and therefore propose tens of taxonomic changes at the tribe and subfamily levels. Archaeobalbini <b>stat. rev.</b> Viidalepp (Geometrinae) is raised from synonymy with Pseudoterpnini Warren to tribal rank. Chlorodontoperini Murillo-Ramos, Sihvonen & Brehm, <b>trib. nov.</b> and Drepanogynini Murillo-Ramos, Sihvonen & Brehm, <b>trib. nov.</b> are described as new tribes in Geometrinae and Ennominae, respectively.