Unveiling the Evolutionary History of a Puzzling Antlion Genus Gatzara Navás (Neuroptera: Myrmeleontidae: Dendroleontinae) Based on Systematic Revision, Molecular Phylogenetics, and Biogeographic Inference

The antlion genus Gatzara Navás, 1915 is one of the major lineages of the subfamily Dendroleontinae Banks, 1899 (Neuroptera: Myrmeleontidae) from Asia, but has a complex systematic background. Here we present a comprehensive systematic revision and mitochondrial phylogenomic analysis to clarify the identity of this genus, and to unravel its evolutionary history. Combining morphological and molecular evidence, we found that the species currently placed in Gatzara belong to two clades, and most of the Gatzara species are considered to be affiliated to the genus Nepsalus Navás, 1912. The dated phylogeny with ancestral area reconstruction indicates that the common ancestor of Gatzara and Nepsalus might have been widely distributed in East Asia and these two genera may have diverged during the late Miocene. The speciation of most Nepsalus species that are allopatric in distribution might have been driven by a series of vicariance events related to the rise of the Himalayas and the formation of the major islands of East Asia during the late Miocene and Pliocene. A new species, namely Nepsalus chikuni sp. n., is described from Tibet. New taxonomic changes include the six new combinations: Nepsalus caelestis (Krivokhatsky, 1997) comb. n., Nepsalus decorillus (Yang, 1997) comb. n., Nepsalus decorosus (Yang, 1988) comb. n., Nepsalus insolitus (Walker, 1860) comb. n., Nepsalus jezoensis (Okamoto, 1910) comb. n., and Nepsalus petrophilus (Miller & Stange in Miller et al., 1999) comb. n.