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The genus-level taxonomy of the ant subfamily Leptanillinae (Hymenoptera: Formicidae) is here revised, with the aim of delimiting genus-level taxa that are reciprocally monophyletic and readily diagnosable based upon all adult forms. This new classification reflects molecular phylogenetics and is informed by joint consideration of both male and worker morphology. Three valid genera are recognized in the Leptanillinae:Opamyrma,Leptanilla(=Scyphodonsyn. nov.,Phaulomyrma,Leptomesites,Noonillasyn. nov.,Yavnellasyn. nov.), andProtanilla(=Anomalomyrmasyn. nov.,Furcotanilla).LeptanillaandProtanillaare further divided into informal, monophyletic species groups. Synoptic diagnoses are provided for all genera and informal supraspecific groupings. In addition, worker-based keys to all described species within the Leptanillinae for which the worker caste is known are provided; and male-based keys to all species for which males are known, plus undescribed male morphospecies for which molecular data are published. The following species are described as new:Protanilla wallaceisp. nov.,Leptanilla acherontiasp. nov.,Leptanilla belantansp. nov.,Leptanilla bethyloidessp. nov., andLeptanilla najaphallasp. nov. 

Abstract The classification of ants (Hymenoptera: Formicidae) has progressed in waves since the first 17 species were described by Linnaeus in the 1758 edition of Systema Naturae. Since then, over 18,000 species-rank names have accumulated for the global myrmecofauna, of which ~14,260 living and ~810 fossil species are valid. Here, we provide a synopsis of ant biodiversity and review the history and classification of the family, while highlighting the massive growth of the field in the new millennium. We observe that major transformation has occurred for ant classification due to advances in DNA sequencing technologies, model-based hypothesis testing, and imaging technologies. We therefore provide a revised and illustrated list of diagnostic character states for the higher clades of Formicidae, recognizing that vastly more work is to be done. To facilitate discussion and the systematic accumulation of evolutionary knowledge for the early evolution of the ants, we suggest an informal nomenclatural system for the higher clades of ants, based on names currently in use and a set of names that have been democratically selected by the authors. To guide future work on ant systematics, we summarize currently available databases and present perspectives on regions in need of biodiversity exploration, challenges facing the field, and the future of ant taxonomy. 

The ant subfamily Leptanillinae (Hymenoptera: Formicidae) consists of minute soil-dwelling species, with several genera within this clade being based solely upon males, including Yavnella Kugler. The dissociation of males and workers has resulted in taxonomic confusion for the Leptanillinae. We here describe the worker caste of Yavnella, facilitated by maximum-likelihood and Bayesian inference from 473 partitioned ultra-conserved element loci, this dataset including 49 other leptanilline species, both described and undescribed. Yavnella laventa sp. nov. is described from seven worker specimens collected in south-western Iran from the Milieu Souterrain Superficiel, a subterranean microhabitat consisting of air-filled cavities among rock and soil fragments, which is subject to similar environmental conditions as caves. This species has bizarrely elongated appendages, which suggests that it is confined to cavities, in contrast with the soil-dwelling behaviour observed in other leptanilline ants. Based on its gracile phenotype relative to other Leptanillinae, Y. laventa shows remarkable adaptations for subterranean life, making it one of a very few examples of this syndrome among the ants. Moreover, the discovery of the worker caste of Yavnella expands our morphological knowledge of the leptanilline ants. We provide worker- and male-based diagnoses of Yavnella, along with a key to the genera of the Leptanillinae for which workers are known. The worker caste of Yavnella as known from this species is immediately recognisable, but the possibility must be noted that described workers of Leptanilla may in fact belong to Yavnella. Further molecular sampling is required to test this hypothesis. ZooBank: urn:lsid:zoobank.org:pub:A54A5766-F35A-4074-9353-1C70FE3955D3 

Although molecular data have proven indispensable in confidently resolving the phylogeny of many clades across the tree of life, these data may be inaccessible for certain taxa. The resolution of taxonomy in the ant subfamily Leptanillinae is made problematic by the absence of DNA sequence data for leptanilline taxa that are known only from male specimens, including the monotypic genus Phaulomyrma Wheeler & Wheeler. Focusing upon the considerable diversity of undescribed male leptanilline morphospecies, the phylogeny of 35 putative morphospecies sampled from across the Leptanillinae, plus an outgroup, is inferred from 11 nuclear loci and 41 discrete male morphological characters using a Bayesian total-evidence framework, with Phaulomyrma represented by morphological data only. Based upon the results of this analysis Phaulomyrma is synonymised with Leptanilla Emery, and male-based diagnoses for Leptanilla that are grounded in phylogeny are provided, under both broad and narrow circumscriptions of that genus. This demonstrates the potential utility of a total-evidence approach in inferring the phylogeny of rare extant taxa for which molecular data are unavailable and begins a long-overdue systematic revision of the Leptanillinae that is focused on male material.