Delimitation of tribes in the subfamily Leptanillinae (Hymenoptera: Formicidae), with a description of the male of Protanilla lini Terayama, 2009

The subfamily Leptanillinae Emery, 1910 (Hymenoptera, Formicidae) is a clade of cryptic subterranean ants, which is restricted to the tropics and warm temperate regions of the Old World. Due to acquisition bias against the minute and hypogaeic workers, most known leptanilline specimens are male, with four genera described solely from males. The sexes have been associated in only two out of 68 described species, meaning that redundant naming of taxa is likely. Herein the phylogeny of the Leptanillinae, sampled with emphasis on largely undescribed male material, is inferred from ultra-conserved elements (UCEs) using maximum-likelihood inference. This method associates the male of Protanilla lini Terayama, 2009 with corresponding workers collected on Okinawa-jima, Japan, allowing the first published description of male ants belonging to the Anomalomyrmini Taylor, 1990, one of the two established tribes within the Leptanillinae. The first male-based diagnosis of these tribes is provided, al...