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Blaps subalpina M��n��tri��s, 1832 stat. resurr. (Figs 10, 23C, D) This species was described from fortress Groznaya (now Groznyi city, Chechen Republic, Russia) or its vast surroundings; type locality: ���Caucase, pr��s de Grosna��a���; the lectotype (ZIN) was designated and studied by Abdurakhmanov & Nabozhenko (2011). Since Allard (1880, 1882), many authors have interpreted this name as a subspecies of Blaps lethifera based on its smooth elytra. Abdurakhmanov & Nabozhenko (2011) interpreted this taxon as a subspecies of Blaps scabriuscula M��n��tri��s, 1832 and proposed the synonymy Blaps subalpina = Blaps montana Motschulsky, 1839. In fact, B. subalpina is a separate species (not subspecies), which differs from B. verrucosa (= B. montana) by the shape of the narrow pronotum, narrowed to apical margin, smooth elytra, little wider mucro in males and the absence of the mucro in females. In addition, both species differ in their bionomics. Blaps verrucosa is cavernicolous species, which inhabits caves, niches and deep cracks in the rocks. High mountain populations (usually more or strongly rugose) of B. verrucosa prefer wet meadow mountain habitats with rocks, while lower mountain Transcaucasian populations (for example, in Gobustan desert) occur in rocks or clusters of huge boulders. Blaps subalpina inhabits xerophytic mountain steppes and slopes only on the North Caucasus (Dagestan, Chechnya) and can often be found in phrygana-like steppes at the foot of Astragalus spp. (milkvetches; Fabaceae), Stipa spp. (feather grasses; Poaceae) and in rodents burrows. These two species are allopatric in natural non-transformed landscapes. However, both species tend to a synanthropic lifestyle and are sometimes sympatric in ruins of abandoned mountain villages in Dagestan and Chechnya. Blaps verrucosa is a species demonstrating great morphological variation. High mountain populations are strongly rugose and slender (Fig. 23B), while almost smooth (but with coarse elytral punctation) specimens have been found in subdesert Gobustan populations around Baku, Azerbaijan (Fig. 23A). Given that the species status of Blaps subalpina is also corroborated by molecular data (Fig. 22) (at least for specimens from Dagestan), the species rank for this taxon is resurrected., Published as part of Nabozhenko, Maxim V., Chigray, Ivan A., Ntatsopoulos, Konstantinos & Papadopoulou, Anna, 2022, A key to Russian and Eastern European species of Blaps Fabricius, 1775 (Coleoptera: Tenebrionidae: Blaptinae) with the description of a new species from the North Caucasus supported by morphological and molecular data, pp. 267-291 in Zootaxa 5116 (2) on page 286, DOI: 10.11646/zootaxa.5116.2.5, http://zenodo.org/record/6367323, {"references":["Menetries, E. (1832) Catalogue raissonne des objets de Zoologie recueillis dans un voyage au Caucase et jusqu'aux frontieres actuelles de la Perse. St. Petersbourg, L'Imprimerie de l'Academie Imperiale des Sciences, 272 pp., 5 pls. https: // doi. org / 10.5962 / bhl. title. 51784","Abdurakhmanov, G. M. & Nabozhenko, M. V. (2011) Keys and catalogue to darkling beetles (Coleoptera: Tenebrionidae s. str.) of the Caucasus and south of European part of Russia. KMK Scientific Press Ltd, Moscow, 361 pp. [in Russian]","Motschulsky, V. de (1839) Insectes du Caucase et des Provinces Transcaucausiennes. Bulletin de la Societe Imperiale des Naturalistes de Moscou, 12, 44 - 93, pls. I, II, V, VI."]}

Blaps lethifera Marsham, 1802 (Fig. 23E, F) = Blaps pterotapha M��n��tri��s, 1832, syn. n. The species Blaps pterotapha M��n��tri��s, 1832 was described from Baku and long time was interpreted as a subspecies of Blaps lethifera (see full bibliography in Abdurakhamnov & Nabozhenko (2011)). Transcaucasian populations named ��� B. lethifera pterotapha ��� differ from the European populations only by weakly emarginated lateral margins of the pronotum at the base. This character is very variable, and specimens with emarginated or weakly emarginated or non-emarginated lateral pronotum margins can be found even in one Transcaucasian micropopulation. This unreliable character���s entire spectrum of variability can be observed in specimens from the east Caspian coast to western Turkey. The extensive variability of this species has been beautifully illustrated in the work of ��ustek (1982). Another illustrative example is two very unusual specimens from the Don River delta area. These specimens were collected after deep excavation by an excavator in sandy alluvial soils near the banks of Don River in the burrows of Spalax microphthalmus G��ldenst��dt, 1770. The first two authors interpreted them as an isolated, distant micropopulation of Blaps scabriuscula (Chigray et al. 2015). However, COI analysis showed that this Don micropopulation is genetically highly similar with specimens assigned to Blaps lethifera (Fig. 22), despite significant morphological differences. Considering the information mentioned above, we propose the following new synonymy: Blaps lethifera Marsham, 1802 = Blaps pterotapha M��n��tri��s, 1832 syn. n. In addition, we examined the specimen, which was listed as Blaps pterosticha Fischer von Waldheim, 1844 from Orenburg Region of Russia (Shapovalov et al. 2011; Kozminykh 2015). The specimen is a typical B. lethifera, so B. pterosticha must be excluded from the Russian faunistic list. Blaps sinuatocollis Solier, 1848 (recorded as nominotypical subspecies from Southern Russia (Nabozhenko & Chigray 2020)) was described as the variation B of Blaps reflexicollis (now a junior synonym of B. lethifera). Types of both taxa are lost (personal communication with Antoine Mantilleri (MNHN)). In our opinion, this name is infrasubspecific and must be excluded from zoological nomenclature because Solier���s description unambiguously reveals that the name was proposed for an infrasubspecific entity (article 45.6.4 of ICZN). Solier indicated that var. B (sinuatocollis) is just one male deviating from the typical form, and if its characters were stable, then it could be a species (Solier 1848, pp. 315, 316)., Published as part of Nabozhenko, Maxim V., Chigray, Ivan A., Ntatsopoulos, Konstantinos & Papadopoulou, Anna, 2022, A key to Russian and Eastern European species of Blaps Fabricius, 1775 (Coleoptera: Tenebrionidae: Blaptinae) with the description of a new species from the North Caucasus supported by morphological and molecular data, pp. 267-291 in Zootaxa 5116 (2) on pages 286-288, DOI: 10.11646/zootaxa.5116.2.5, http://zenodo.org/record/6367323, {"references":["Menetries, E. (1832) Catalogue raissonne des objets de Zoologie recueillis dans un voyage au Caucase et jusqu'aux frontieres actuelles de la Perse. St. Petersbourg, L'Imprimerie de l'Academie Imperiale des Sciences, 272 pp., 5 pls. https: // doi. org / 10.5962 / bhl. title. 51784","Sustek, Z. (1982) Contribution to the synonymy of Blaps lethifera Marsham, 1802 (Coleoptera Tenebrionidae). Acta entomologica bohemoslovaca, 79, 143 - 153.","Chigray, I. A., Abdurakhmanov, G. M., Nabozhenko, M. V. & Shmatko, V. Yu. (2015) Morphological diversity and distribution of Blaps scabriuscula Menetries, 1832 (Coleoptera: Tenebrionidae). Yug Rossii: Ekologiya, Razvitiye, 10 (4), 59 - 68. [in Russian] https: // doi. org / 10.18470 / 1992 - 1098 - 2015 - 4 - 59 - 68","Shapovalov, A. M., Grigoriev, V. E., Nemkov, V. A., Rusakov, A. V. & Kazakov, E. P. (2011) Interesting finds of beetles (Insecta, Coleoptera) in the Orenburg Region. Trudy Orenburgskogo otdeleniya Russkogo entomologicheskogo obshchestva, 1, 39 - 48. [in Russian]","Kozminykh, V. O. (2015) Faunistic data on tenebrionid beetles (Coleoptera, Tenebrionidae) of the Orenburg Region. Vestnik of Orenburg State Pedagogical University. Electronic Scientific Journal, 1 (13), 16 - 42. [in Russian]","Solier, A. J. J. (1848) Essai sur les collapterides. 14 e Tribu. Blapsites. In: Baudi di Selve: Studi Entomologici. Pubblicati per cura di Flaminio Baudi e di Eugenio Truqui. Tomo 1. Torino: Stamperia Degli Artisti Tipografi, pp. 149 - 370, pls. iv - xv.","Nabozhenko, M. V. & Chigray, I. A. (2020) Tribe Blaptini Leach, 1815. In: Iwan, D. & Lobl, I. (Eds.), Catalogue of Palaearctic Coleoptera. Volume 5. Tenebrionoidea. Leiden, Brill, pp. 268 - 296. https: // doi. org / 10.1163 / 9789004434998 _ 004"]}

Blaps verrucosa Adams, 1817 (Figs 6, 23A, B) = Blaps scabriuscula M��n��tri��s, 1832 syn. n. = Blaps montana Motschulsky, 1839 syn. n. Blaps verrucosa was described from the surroundings of Tiflis (now Tbilisi, Georgia). Adams���s description is detailed and completely corresponds to Blaps scabriuscula M��n��tri��s, 1832 described from Baku (Azerbaijan) (M��n��tri��s 1832) and B. montana Motschulsky, 1839 described from Kurush (Southern Dagestan, Russia) (Motschulsky 1839). Types of all species are lost, but the first author examined specimens from Baku, Kurush and Georgia (Abdurakhmanov & Nabozhenko 2011) and confirmed that all three taxa are morphologically conspecific and differ only in the level of elytral rugosity: from weak, poorly visible wrinkles in some specimens from Kurush and Baku to strongly and coarsely rugose specimens from mountains of Inner Dagestan, Chechnya and southern slopes of the Greater Caucasus in Georgia. Adams mentioned that this very rare species has rugose flattened elytra with mucro, subquadrate pronotum, and long legs. Only one species with such characters occurs in Georgia, and it was listed for the eastern part of the country (Nakalakebi, Kazbegi) by Dzhambazishvili (2000). As a result, the following synonymy is proposed: Blaps verrucosa Adams, 1817 = Blaps scabriuscula M��n��tri��s, 1832 syn. n., = Blaps montana Motschulsky, 1839 syn. n. Notes. A.V. Bogatshev confused the status and subspecies of B. verrucosa and mixed them with other species. The following interpretations were published by Bogatshev: 1. Bogatshev (1934): Blaps montana (= Blaps scabriuscula, as a junior synonym). In addition, he mentioned two of its subspecies B. montana rugosa and B. montana chevsuretica. Both names are nomina nuda, because descriptions are absent. The Greater Caucasus. 2. Bogatshev (1938): Blaps montana. Russia: Dagestan (Kurush, Chadakolob); Azerbaijan: Sary-Bash; Georgia: Kheledula, Latal, Lailchala, Mestia. 3. Bogatshev (1958): Blaps scabriuscula scabriuscula: Azerbaijan: Baku and surrounds; Blaps scabriuscula montana: Greater Caucasus; B. scabriuscula puella Allard, 1881: Armenia, Azerbaijan (Karabakh, Nakhchivan), Eastern Anatolia. Blaps puella, which was interpreted as a subspecies of B. scabriuscula (Bogatshev 1958), belongs to the mortisaga species-group with acute, strongly elongate reservoirs of the spermatheca (Chigray & Nabozhenko 2016, Figs 7A���C) only slightly differing from B. mortisaga., Published as part of Nabozhenko, Maxim V., Chigray, Ivan A., Ntatsopoulos, Konstantinos & Papadopoulou, Anna, 2022, A key to Russian and Eastern European species of Blaps Fabricius, 1775 (Coleoptera: Tenebrionidae: Blaptinae) with the description of a new species from the North Caucasus supported by morphological and molecular data, pp. 267-291 in Zootaxa 5116 (2) on pages 285-286, DOI: 10.11646/zootaxa.5116.2.5, http://zenodo.org/record/6367323, {"references":["Menetries, E. (1832) Catalogue raissonne des objets de Zoologie recueillis dans un voyage au Caucase et jusqu'aux frontieres actuelles de la Perse. St. Petersbourg, L'Imprimerie de l'Academie Imperiale des Sciences, 272 pp., 5 pls. https: // doi. org / 10.5962 / bhl. title. 51784","Motschulsky, V. de (1839) Insectes du Caucase et des Provinces Transcaucausiennes. Bulletin de la Societe Imperiale des Naturalistes de Moscou, 12, 44 - 93, pls. I, II, V, VI.","Abdurakhmanov, G. M. & Nabozhenko, M. V. (2011) Keys and catalogue to darkling beetles (Coleoptera: Tenebrionidae s. str.) of the Caucasus and south of European part of Russia. KMK Scientific Press Ltd, Moscow, 361 pp. [in Russian]","Dzhambazishvili, M. Ya. (2000) Catalogue to darkling beetles (Coleoptera, Tenebrionidae) of Georgia. Trudy instituta zoologii Akademii nauk Gruzii, 20, 185 - 193. [in Russian]","Bogatshev, A. V. (1934) Materials to the knowledge of the fauna of beetles of Apsheron Penisula. Trudy Azerbaijanskogo otdelenia Zakavkazskogo filiala of the AN SSR. Sektor zoologii, 7, 14 - 71.","Bogatshev, A. V. (1938) Review of species of the fam. Tenebrionidae of the Caucasus and neighboring countries. Bulleten' Gosudarstvennogo Muzeya Gruzii, 9, 118 - 127 [in Russian].","Bogatshev, A. V. (1958) Darkling beetles. In: Kryzhanovsky, O. L. & Ter-Minasyan, M. E. Beetles - Coleoptera. In: Animal world of the USSR. Vol. 5. Mountain regions of the European part of the USSR. Academy of Sciences of the USSR, Moscow - Leningrad, pp. 384 - 421.","Chigray, I. A. & Nabozhenko, M. V. (2016) To the knowledge of the genus Blaps Fabricius, 1775 (Coleoptera: Tenebrionidae) from Iran and Transcaucasia. Annales Zoologici, 66 (2), 267 - 275. https: // doi. org / 10.3161 / 00034541 ANZ 2016.66.2.007"]}