A phylogenetic classification of Leptodirini (Coleoptera, Leiodidae, Cholevinae)
Authors: Fresneda Javier , Giachino Pier Mauro , Salgado José María , Faille Arnaud , Bourdeau Charles , Cieslak Alexandra , Ribera† Ignacio 
Published: December 2024

The tribe Leptodirini of the beetle family Leiodidae is one of the most diverse radiations of subterranean animals, with a distribution centered north of the Mediterranean basin from the Iberian Peninsula to Iran. It currently comprises some 1340 species and subspecies in around 250 genera and subgenera. We present a molecular phylogeny of the tribe, and subsequently a revision of their taxonomy taking into account our phylogenetic results. We investigated and established the phylogenetic relationships of 558 species using a combination of fragments of four mitochondrial (two fragments of COI, 16s and cytochrome b) as well as two nuclear (18s and 28s) genes, analysed using maximum likelihood methods. In agreement with the results of the phylogenetic analysis we establish the following synonymies: Leptodirini Lacordaire, 1854 [1849] = Bathysciae Horn, 1880 syn. nov. = Pholeuones Reitter, 1886 syn. nov. = Anthroherpona Jeannel, 1910 syn. nov. = Bathysciotina V.B. Guéorguiev, 1974 syn. nov. = Spelaeobatina V.B. Guéorguiev, 1974 syn. nov. We divided the tribe Leptodirini into three subtribes, two of them monogeneric, Platycholeina and Adelopsellina subtr. nov., and a third one including all of the other Leptodirini, the Leptodirina sensu novo. We further informally divide Leptodirina sensu novo into several monophyletic «series», the most of them strongly supported by the phylogeny, as well as geographically well-defined and morphologically coherent.
20 new genera and one subgenera are established: Ariejasia gen. nov. (type species Speonomus carrerei Fourès, 1954), Casalella gen. nov. (type species Adelops meridionalis Jacquelin du Val, 1854), Catalaniana gen. nov. (type species Bathysciola zariquieyi Bolívar, 1919), Dupreia gen. nov. (type species Euryspeonomus beruetei Dupré, 1991), Galliana gen. nov. (type species Adelops linderi Abeille de Perrin, 1875), Georgiana gen. nov. (type species Bathysciola suramensis Jeannel, 1930), Hypogesia gen. nov. (type species Speocharis minos Jeannel, 1909), Lakedemoniana gen. nov. (type species Lakedemoniana mizerakisi sp. nov.), Luquesia gen. nov. (type species Quaestus pasensis Salgado, Labrada & Luque, 2010), Latellana gen. nov. (type species Bathysciola fortesculpta Jeannel, 1914), Nanobathysciola gen. nov. (type species Bathyscia ovata Kiesenwetter, 1850), Perreauia gen. nov. (type species Perreauia dalmatica sp. nov.), Pyrenaeola gen. nov. (type species Adelops lapidicola Saulcy, 1872), Pyrenaeola (Garunnia) subgen. nov. (type species Bathyscia larcennei Abeille de Perrin, 1883), Schaufussia gen. nov. (type species Speocharis pachecoi Bolívar, 1915), Serbiana gen. nov. (type species Serbiana latitarsis sp. nov.), Uhagoniella gen. nov. (type species Speocharis zariquieyi Jeannel, 1924), Valdesiella gen. nov. (type species Dellabeffaella olmii Casale, 1980), Vallespirana gen. nov. (type species Adelops grenieri Saulcy, 1872), Vanderberghia gen. nov. (type species Adelops speluncarum Delarouzée, 1857) and Zoiana gen. nov. (type species Phaneropella (Hittitia) kociani Švec, 2016).
We describe 29 new species: Aphaobius gorenjskanus sp. nov., Aphaobius luckae sp. nov., Archeoboldoria martinottii sp. nov., Bathyscidius crnagorensis sp. nov., Bathysciola balestrae sp. nov., Bathysciola chestai sp. nov., Bathysciola lanai sp. nov., Bathysciola oliveroi sp. nov., Batinoscelis degiovannii sp. nov., Batinoscelis onnisi sp. nov., Catopsinus rudoyi sp. nov., Lakedemoniana mizerakisi sp. nov., Latellana benellii sp. nov., Latellana degiovannii sp. nov., Latellana magrinii sp. nov., Latellana petruzziellana sp. nov., Latellana petruzzielloi sp. nov., Nafarroa ericduprei sp. nov., Paratroglophyes juberthiei sp. nov., Perreauia dalmatica sp. nov., Proleonhardella epimikes sp. nov., Salfia angelinii sp. nov., Salfia laudis sp. nov., Schaufussia sinuatipenis sp. nov., Serbiana latitarsis sp. nov., Tamaniniella pivai sp. nov., Troglocharinus bellesi sp. nov., Troglocharinus rizzoae sp. nov. and Vanderberghia corbazi sp. nov.
We establish the generic synonymies: Espanoliella V.B. Guéorguiev, 1976 = Samanolla Salgado, 2000 syn. nov. (type species Speocharis oxypterus Bolívar, 1917); Ovobathysciola Jeannel, 1924 = Patriziella Jeannel, 1956 syn. nov. (type species Patriziella sardoa Jeannel, 1956) = Sardostygia Casale, 2019 syn. nov. (type species Sardostygia cogonii Casale, 2019); Quaestus Schaufuss, 1861 = Oresigenus Jeannel, 1948 syn. nov. (type species Oresigenus jaspei Jeannel, 1948) = Speogeus Salgado, 1985 syn. nov. (type species Speocharis avicularis Salgado, 1985) = Asturianella Salgado & Fresneda, 2004 syn. nov. (type species Quaestus incognitus Salgado & Fresneda, 2004); Speonomus Jeannel, 1908 = Metaspeonomus Coiffait, 1959 syn. nov. (type species Speonomus monticola Coiffait, 1959); Bathysciella Jeannel, 1906 = Phacomorphus Jeannel, 1908 syn. nov. (type species Bathyscia Mascarauxi Sainte Claire Deville, 1905) = Phacomorphoides Dupré, 1996 syn. nov. (type species Bathyscia alexinae Jeannel, 1906); Magdelainella Jeannel, 1924 = Kosaniniella S. Ćurčić, Brajković & B. Ćurčić, 2004 syn. nov. (type species Kosaniniella javorensis S. Ćurčić, Brajković & B. Ćurčić in S. Ćurčić et al., 2004); Proleonhardella Jeannel, 1910 = Bozidaria S. Ćurčić & Pavićević in S. Ćurčić et al., 2021 syn. nov. (type species Bozidaria serboocidentalis S. Ćurčić & Pavićević in S. Ćurčić et al., 2021), = Serboleonhardella S. Ćurčić & Schönmann in S. Ćurčić et al., 2008 syn. nov. (type species Proleonhardella remyi Jeannel, 1934); Rozajella S. Ćurčić, Brajković & B. Ćurčić, 2007 in S. Ćurčić et al., 2007 = Nonveilleriella Perreau & Pavićević, 2008 syn. nov. (type species Nonveilleriella ognjenovici Perreau & Pavićević, 2008).
Six subgenera are elevated to the rank of genus: Amphogeus Salgado, 2000 stat. nov. (type species Bathyscia cantabrica Uhagón, 1881); Hittitia Casale, Giachino, Vailati & Rampini, 1991 stat. nov. (type species Bathyscia (Bathyscia) turcica Reitter, 1885); Ochsiella Jeannel, 1947 stat. nov. (type species Bathyscia Bucheti Abeille de Perrin, 1905); Salfia Tamanini, 1955 stat. nov. (type species Bathysciola (Salfia) ruffoi Tamanini, 1955); Uludagites Casale, Giachino, Vailati & Rampini, 1991 stat. nov. (type species Phaneropella (Uludagites) minuta Casale, Giachino, Vailati & Rampini, 1991) and Cernella Jeannel, 1930 stat. nov. (type species Bathyscia (Sophrochaeta) reitteri Frivaldsky, 1884).
Bathyscia schiodtei Kiesenwetter, 1850 = Hoffmannia biokovensis Knirsch, 1928 syn. nov. Zariquieyella Jeannel, 1929 is substitute name for Hoffmannia Knirsch, 1928 nom. praeocc. The name Zariquieyella applies to the genus for the species of the Bathysciola schiodtei group.
A lectotype is designated for Adelops lapidicola Saulcy, 1872. 164 new combinations for species and subspecies are proposed. Close relationships between Coreobathyscia and Proleptodirina with Leptodirini are refuted, with the former genus appearing more closely related to Cholevini and the latter to Ptomaphagini.