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, along with a
dichotomous key to both all described male-based species within the
Leptanillinae and male morphospecies sequenced in this study. With
genome-scale data enabling the association of separately collected sexes
and phylogenomic inference contextualizing morphological observations, the
parallel taxonomy that afflicts this enigmatic group of ants can begin to
be resolved.