A revision of the Palaearctic Pimeliini (Coleoptera: Tenebrionidae): a comparative analysis and systematic position of Eastern European and Asian taxa with dorso-lateral eyes

Keywords: Tenebrionidae, Pimeliini, new taxa, synonymy, nomenclature

Abstract

A taxonomic review of tenebrionid platyopoid genera of the subfamily Pimeliinae from Eastern Europe, Central Asia, Afghanistan, Iran and Pakistan is given. This group of taxa was known before 1994 as the tribe Platyopini Motschulsky, 1849, which is now interpreted as a junior synonym of Pimeliini Latreille, 1802. The group is different from other Pimeliini in having dorso-lateral eyes, located above the level of the genae, and it includes the following ultrapsammophilic genera at least from Central and Southern Asia: Apatopsis Semenov, 1891, Habrochiton Semenov-Tjan-Shansky, 1907, Habrobates Semenov, 1903 [= Kawiria Schuster, 1935 syn. nov.], Dietomorpha Reymond, 1938, Przewalskia Semenov, 1893, Mantichorula Reitter, 1889, Platyope Fischer von Waldheim, 1820 [= Homopsis Semenov, 1893 syn. nov.], Earophanta Semenov, 1903. These genera are distributed in almost all large deserts of Palaearctic Asia: Karakum, Kyzylkum, Muyunkum, Taklamakan, Gobi, Registan, Dasht-e-Kawir, Dasht-e-Lut, as well as in other arid and semi-arid sandy landscapes from European Russia to the south of Eastern Siberia. The group of platyopoid genera is polyphyletic. We propose at least two monophyletic branches: the Habrobates genus group (the first four genera mentioned above), which represents the subtribe Habrobatina Nabozhenko & S. Chigray subtrib. nov. and the Platyope genus group (latter four genera) within the nominotypical subtribe. A new species is described from Pakistan (Balochistan): Dietomorpha gonzalesi S. Chigray & Nabozhenko sp. nov. Platyope granulata Fischer von Waldheim, 1820 is recorded for Kazakhstan for the first time. The following synonymy is resurrected: Apatopsis grombczewskii Semenov, 1890 Apatopsis conradti Semenov, 1890, syn. resurr. Two new combinations resulting from the synonymy of genera are given: Habrobates gabrieli Schuster, 1935 comb. nov. (from Kawiria), Platyope grumi Semenov, 1893 comb. nov. (from Homopsis). Lectotypes are designated for the following taxa: Apatopsis grombczewskii (Semenov, 1891), Apatopsis conradti Semenov, 1891, Habrochiton vernus Semenov-Tjan-Shansky, 1907, Habrobates vernalis Semenov, 1903, Kawiria gabrieli Schuster, 1935, Platyope dilatata Reitter, 1887; Mantichorula semenowi Reitter, 1889, Mantichorula grandis Semenov, 1893, Homopsis grumi Semenov, 1893, Platyope serrata Semenov, 1893, Platyope planidorsis Reitter, 1889, Platyope tomentosa Semenov, 1893. Additional information for type specimens studied by the authors is given for Habrochiton primaeveris Semenov-Tjan-Shansky, 1907 (holotype), Habrobates vejisovi Kelejnikova, 1977, Platyope ordossica Semenov-Tjan-Shansky, 1907 (holotype), Earophanta autumnalis Semenov, 1903 (holotype, junior synonym of E. planidorsis Reitter, 1889), Earophanta loudoni Semenov, 1903 (holotype, junior synonym of Earophanta pilosissima Reitter, 1895), Earophanta pubescens Skopin, 1960 (holotype, paratypes), Earophanta beludzhistana Bogatchev, 1957 (holotype).

References

Abdurakhmanov G.M. & Nabozhenko M.V. 2011. Keys to and Catalogue of Darkling Beetles (Coleoptera: Tenebrionidae s. str.) of the Caucasus and South of European Part of Russia. KMK, Moscow. [In Russian.]

Abdurakhmanov G.M. & Nabozhenko M.V. 2016. On faunogenesis of tenebrionid beetles (Coleoptera: Tenebrionidae) of Middle Asia. South of Russia: Ecology, Development 11 (2): 170–177. https://doi.org/10.18470/1992-1098-2016-2-170-177 [In Russian.]

Abdurakhmanov G.M., Nabozhenko M.V., Abdurakhmanov A.G., Ivanushenko Yu.Yu. & Daudova M.G. 2016a. Geographic relations of darkling beetles (Coleoptera: Tenebrionidae) of the Palaearctic Tethys desert-steppe region with the historical review. South of Russia: Ecology, Development 11 (3): 35–89. https://doi.org/10.18470/1992-1098-2016-3-35-89 [In Russian.]

Abdurakhmanov G.M., Shokhin I.V., Teymurov A.A., Abdurakhmanov A.G., Gadzhiev A.A., Daudova M.G., Magomedova M.Z. & Ivanushenko Yu.Yu. 2016b. The use of the elements of morphoecological adaptations of organisms to the environment under paleogeographic reconstructions of biotas of Tetiysky desert-steppe region (building schemes of historical formation of flora and fauna). South of Russia: Ecology, Development 11 (2): 9–31. https://doi.org/10.18470/1992-1098-2016-2-9-31 [In Russian.]

Abushama F.T. 1990. Epigean (ground) insects. In: Sahara (translation from English): 155–170. Progress, Moscow.

Bogatchev A.V. 1949. New species of Tenebrionidae of Azerbaijan and other areas of the Palaearctic. Doklady Akademii Nauk Azerbaydzhanskoy SSR 5 (1): 38–41. [In Russian.]

Bogatchev A.V. 1960a. A new genus of darkling beetles from Tajikistan Allotadzhikistania gen. nov. (Tenebrionidae, Pimeliini). Doklady Akademii Nauk Tadzhikskoy SSR 3 (1): 43–46. [In Russian.]

Bogatchev A.V. 1960b. New unusual genus of darkling beetles, found in Tajikistan – Tadzhikistania gen. nov. (Coleoptera, Tenebrionidae). Doklady Akademii Nauk Tadzhikskoy SSR 3 (3): 35–37. [In Russian.]

Bogachev A.V. & Nepesova M.G. 1980. Tenebrionid beetles of the genus Pachyscelis Sol. (Coleoptera, Tenebrionidae) from Turkmenia. Entomologicheskoe Obozrenie 59: 607–612. [In Russian.]

Bouchard P., Lawrence J.F., Davies A.E. & Newton A.F. 2005. Synoptic classification of the world Tenebrionidae (Insecta: Coleoptera) with a review of family-group names. Annales zoologici 55 (4): 499–530.

Bouchard P., Bousquet Y., Davies A.E., Alonso-Zarazaga M.A., Lawrence J.F., Lyal C.H.C., Newton A.F., Reid C.A.M., Schmitt M., Ślipiński S.A. & Smith A.B.T. 2011. Family-group names in Coleoptera (Insecta). ZooKeys 88: 1–972. https://doi.org/10.3897/zookeys.88.807

Chigray I. & Ivanov A. 2020. A Review of the genus Blaps (Coleoptera: Tenebrionidae) of Central and South Kazakhstan with description of two new species. Acta Zoologica Academiae Scientiarum Hungaricae 66 (2): 111–162. https://doi.org/10.17109/azh.66.2.111.2020

Chigray I., Nabozhenko M., Abdurakhmanov G. & Keskin B. 2019. A systematic review of the genus Dila Fischer von Waldheim, 1844 (= Caenoblaps König, 1906, syn.n.) (Coleoptera: Tenebrionidae) from the Caucasus, Turkey and boundary territories of Iran. Insect Systematics & Evolution 51 (4): 1–30. https://doi.org/10.1163/1876312x-00001006

Chigray I., Nabozhenko M., Merkl O. & Kovalev A. 2018. A review of the genus Prosodes Eschscholtz, 1829 (Coleoptera: Tenebrionidae) of Iran. Zootaxa 4379 (4): 451–483. https://doi.org/10.11646/zootaxa.4379.4.1

Chigray S.N. 2018. A revision of the genus Ascelosodis L. Redtenbacher, 1868 (Coleoptera: Tenebrionidae: Pimeliinae: Edrotini) of Pamir. Caucasian Entomological Bulletin 14 (2): 165–169. https://doi.org/10.23885/181433262018142-165169

Chigray S.N. 2019. Revision of the Palaearctic Pimeliini (Coleoptera, Tenebrionidae): the genus Podhomala Solier, 1836. Entomological Review 99 (7): 924–948. https://doi.org/10.1134/s0013873819070066

Chigray S.N. & Abakumov E.V. 2019. Taxonomic Revision of the Genus Calyptopsis Solier, 1835 (Coleoptera, Tenebrionidae: Pimeliinae: Tentyriini) from Iran. Entomological Review 99 (2): 209–244. https://doi.org/10.1134/s001387381902009x

Chigray S., Nabozhenko M., Keskin B. & Abdurakhmanov G. 2018. Taxonomic review of the genus Calyptopsis of Turkey and adjacent areas (Coleoptera: Tenebrionidae: Tentyriini). Acta entomologica Musei Nationalis Pragae 58 (2): 331–346. https://doi.org/10.2478/aemnp-2018-0028

Cloudsley-Thompson J.L. & Chadwick M.J. 1964. Life in Deserts. Dufour, Philadelphia.

Doyen J.T. 1972. Familial and subfamilial classification of the Tenebrionoidea (Coleoptera) and a revised generic classification of the Coniontini (Tentyriidae). Quaestiones entomologicae 8: 357–376.

Doyen J.T. 1994. Cladistic relationships among Pimeliine Tenebrionidae (Coleoptera). Journal of the New York Entomological Society [1993] 101 (4): 443–514.

Edney E.B. 1971. The body temperature of tenebrionid beetles in the Namib Desert of Southern Africa. The Journal of Experimental Biology 55: 253–272.

Fischer von Waldheim G. 1820. Entomographie de la Russie (Entomographia Imperii Rossici). Auctoritate Societatis Caesareae Mosquensis naturae scrutatorum collecta et in lucem edita. Vol. 1. A. Semen, Moscow.

Fischer von Waldheim G. 1822. Entomographie de la Russie (Entomographia Imperii Rossici). Auctoritate Societatis Caesareae Mosquensis naturae scrutatorum collecta et in lucem edita. Vol. 2. A. Semen, Moscow.

Fischer von Waldheim G. 1837. Notice sur les mélasomes. Bulletin de la Société impériale des Naturalistes de Moscou 10 (4): 3–18, pl. ii.

Gebien H. 1937. Katalog der Tenebrioniden (Col. Heteromera) Teil I. Pubblicazioni del Museo Entomologico ‘Pietro Rossi’ 2: 505–883.

Gebien H. 1940. Die von Dr. Trinkler in Tibet gesammelten Tenebrioniden (Col. Tenebr.). Mitteilungen der Münchener Entomologischen Gesellschaft 30: 1–3.

Grimm R. 2015. Tenebrionidae (Insecta: Coleoptera) from Iran. Vernate 34: 299–318.

Herbst J.F.W. 1799. Natursystem aller bekannten in- und ausländischen Insekten, als eine Fortsetzung der von Buffonschen Naturgeschichte. Der Käfer achter Theil. Geh. Commerzien-Raths Pauli, Berlin. https://doi.org/10.5962/bhl.title.156919

ICZN. 1999. International Code of Zoological Nomenclature. 4th Edition. Adopted by the International Union of Biological Sciences. The International Trust for Zoological Nomenclature. London, UK. Available from http://iczn.org [accessed 9 Mar. 2022].

IUSS Working Group WRB. 2015. World Reference Base for Soil Resources 2014, update 2015. International soil classification system for naming soils and creating legends for soil maps. World Soil Resources Reports No. 106. FAO, Rome, Italy.

Iwan D., Löbl I., Bouchard P., Bousquet Y., Kamiński M.J., Merkl O., Ando K. & Schawaller W. 2020. Family Tenebrionidae Latreille, 1802. In: Iwan D. & Löbl I. (eds) Catalogue of Palaearctic Coleoptera. Vol. 5: Tenebrionoidea: 104–476. Brill, Leiden. https://doi.org/10.1163/9789004434998

Jacquelin du Val C. 1861. Manuel entomologique. Genera des coléoptères d’Europe comprenant leur classification en familles naturelles, la description de tous les genres, des tableaux synoptiques destinés à faciliter l’étude, le Catalogue de toutes les espèces de nombreux dessins au trait de charactères et plus de treize cents types représentant un ou plusieurs insectes de chaque genre dessinés et peints d’après nature avec le plus grand soin par M. Jules Migneaux. Vol. 3. Deyrolle, Paris.

Kaplin V.G. 2019. Distribution, life forms and ecological peculiarities of darkling beetles (Coleoptera, Tenebrionidae) of the Eastern Kara Kum. Entomological Review 99 (7): 987–1004. https://doi.org/10.1134/s0013873819070091

Kaszab Z. 1957. Zehn neue Tenebrioniden aus Asien (Coleoptera). Annales Historico-Naturales Musei Nationalis Hungarici. Series nova 8: 289–299.

Kaszab Z. 1967. Die Tenebrioniden der Westmongolei (Coleoptera). Acta Zoologica Academiae Scientiarum Hungaricae 13 (3–4): 279–351.

Kaszab Z. 1982. Insects of Saudi Arabia. Coleoptera: fam. Tenebrionidae (part 2). Fauna of Saudi Arabia 4: 124–243.

Koch C. 1964. Zur Synonymie von Dietomorpha pardalis Reymond (Col., Tenebrionidae). Entomologische Arbeiten aus dem Museum G. Frey 15: 741–742.

Koch C. 1969. Storthocnemis kulzeri, ein bisher unbekanntes Bindeglied zwischen den Tribus Pimeliini/Platyopini und Leucolaephini (Col., Tenebrionidae). Entomologische Arbeiten aus dem Museum G. Frey 20: 484–493.

Kryzhanovsky O.L. 1965. Composition and Origin of Terrestrial Fauna of Middle Asia (Based Mainly on the Material of Beetles). Nauka, Moscow-Leningrad. [In Russian.]

Kühnelt W. 1957. Ergebnisse der Österreichischen Iran-Expedition 1949/50. Die Tenebrioniden Irans. Sitzungsberichte der Österreichischen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse 166 (2): 65–102.

Kwieton E. 1978a. Sur la division de l’ancien genre Pachyscelis Sol. (Col., Tenebrionidae). Bulletin de la Société Entomologique de Mulhouse 1978: 28–34.

Kwieton E. 1978b. Essai synoptique sur le genre Thriptera Solier (Col. Tenebrionidae). Annotationes Zoologicae et Botanicae 123: 1–15.

Kwieton E. 1980. Révision du genre Pachyscelis Solier et notes sur des relations inter-génériques chez les Pimeliini (Col., Tenebrionidae. Annotationes Zoologicae et Botanicae 137: 1–16.

Kwieton E. 1981. Exposé sommaire des faits concernant la phylogenèse et la paléogéographie de la sous-famille des Pimeliinae de l’Ancient Monde. Bulletin de la Société entomologique de Mulhouse 25: 1–7.

Kwieton E. 1982. Notes sur les Tenebrionidae Pimeliinae du Pakistan. Bulletin de la Société entomologique de Mulhouse 26: 21–26.

Lacordaire J.T. 1859. Histoire naturelle des insectes. Genera des coléoptères ou exposé méthodique et critique de tous les genres proposés jusqu’ici dans ce ordre d’Insectes. Vol. 5. Librairie Encyclopédique de Roret, Paris.

Leo P. 2016. Osservazioni su Spectrocnera anguliceps Kwieton, 1981, descrizione della femmina e prima segnalazione per l’Oman (Coleoptera, Tenebrionidae, Pimeliinae, Pimeliini). Revista gaditana de Entomología 7 (1): 469–478.

Lillig M. & Pavlíček T. 2002. Astorthocnemis becvarorum, a new genus and new species from Middle East (Coleoptera: Tenebrionidae: Pimeliinae: Platyopini). Mitteilungen des Internationalen Entomologischen Vereins 27: 97–104.

Löbl I., Merkl O., Ando K., Bouchard P., Lillig M., Masumoto K. & Schawaller W. 2008. Family Tenebrionidae Latreille, 1802. In: Löbl I. & Smetana A. (eds) Catalogue of Palaearctic Coleoptera. Vol. 5: Tenebrionoidea: 105–352. Apollo books, Stenstrup.

Matthews E.G. 1993. Classification, relationships and distribution of the genera of Heleini (Coleoptera: Tenebrionidae). Invertebrate Taxonomy 7 (5): 1025–1095. https://doi.org/10.1071/it9931025

Matthews E.G. & Bouchard P. 2008. Tenebrionid Beetles of Australia: Descriptions of Tribes, Keys to Genera, Catalogue of Species. Australian Biological Resources Study, Canberra.

Matthews E.G., Lawrence J.F., Bouchard P., Steiner W.E., Ślipiński Jr. & Ślipiński S.A. 2010. Tenebrionidae Latreille, 1802. In: Handbook of zoology. Arthropoda: Insecta. Part 38. Coleoptera, beetles. Vol. 2. Morphology and Systematics (Elateroidea, Bostrichiformia, Cucujiformia partim): 574–659. Walter de Gruyter, Berlin.

Medvedev G.S. 1959. Types of mouthparts in darkling beetles (Coleoptera, Tenebrionidae) of Turmenistan. Zoologicheskiy zhurnal 38 (8): 1214–1229. [In Russian.]

Medvedev G.S. 1962. A new subfamily of darkling beetles (Coleoptera, Tenebrionidae) from Turkmenia. Zoologicheskii Zhurnal 41 (8): 1184–1189. [In Russian.]

Medvedev G.S. 1965a. 71. Fam. Tenebrionidae – darkling beetles. In: Key to the Insects of the European Part of the USSR. Vol. 2. Coleoptera and Strepsiptera: 365–381. Nauka, Leningrad. [In Russian.]

Medvedev G.S. 1965b. The types of adaptations of legs structure of desert darkling beetles (Coleoptera, Tenebrionidae). Entomologicheskoe obozrenie 44 (4): 803–826. [In Russian.]

Medvedev G.S. 1970. Eco-morphological types of darkling beetles (Coleoptera, Tenebrionidae) of Middle Asia. In: The Fauna and Ecology of Insects of Middle Asia: 26–44. Irfon, Dushanbe. [In Russian.]

Medvedev G.S. 1990. Key to the darkling beetles of Mongolia. Proceedings of the Zoological Institute 220. Zoological Institute of the USSR, Leningrad. [In Russian.]

Medvedev G.S. 1992. 97. Fam. Tenebrionidae – Darkling beetles. In: Key to the Insects of the Far East of the USSR. Vol. 3. Coleoptera, or beetles. Part 2: 621–659. Dal’nauka, Vladivostok. [In Russian.]

Medvedev G.S. 1993. To the knowledge of darkling beetles of the genus Lasiostola Sol. (Coleoptera, Tenebrionidae). Entomologicheskoe Obozrenie 73 (1): 106–118. [In Russian.]

Medvedev G.S. 2001. Evolution and System of Darkling Beetles of the Tribe Blaptini (Coleoptera, Tenebrionidae). Lectures in commemoration of N.A. Cholodkovskogo. Vol. 53. Russian Entomological Society Publ., St. Petersburg. [In Russian.]

Medvedev G.S. 2005. On connections of the sand desert faunas of Tenebrionidae (Coleoptera) of Middle Asia, Iran, and Afghanistan. In: Konstantinov A.S., Tishechkin A.K. & Penev L. (eds) Contribution to Systematics and Biology of Beetles. Papers Celebrating the 80th Birthday of Igor Konstantinovich Lopatin: 299–314. Pensoft, Moscow.

Medvedev G.S. & Nepesova M.G. 1985. Key to Darkling Beetles of Turkmenistan. Ylym, Ashgabat. [In Russian.]

Motschulsky V. de. 1849. Coléoptères reçus d’un voyage de M. Handschuh dans le midi de l’Espagne, énumerés et suivis de notes. Bulletin de la Société impériale des Naturalistes de Moscou 22 (3–4): 52–163.

Nabozhenko M.V. & Ando K. 2018. Subtribal, generic and subgeneric composition of darkling beetles of the tribe Helopini (Coleoptera: Tenebrionidae) in the Eastern Palaearctic. Acta zoologica Academiae Scientiarum Hungaricae 64 (4): 277–327. https://doi.org/10.17109/azh.64.4.277.2018

Nabozhenko M.V. & Grimm R. 2019. New species and new records of darkling beetles of the tribe Helopini (Coleoptera: Tenebrionidae) from the Western Palaearctic. Caucasian Entomological Bulletin 15 (1): 107–116. https://doi.org/10.23885/181433262019151-107116

Nabozhenko M. & Purchart L. 2019. Socotraphanes – a new endemic genus of darkling beetles (Coleoptera: Tenebrionidae) from Socotra Island. Annales zoologici 69 (1): 149–157. https://doi.org/10.3161/00034541anz2019.69.1.010

Nabozhenko M.V. & Sadeghi S. 2017. Foranotum perforatum gen. et sp. nov. – a new troglobitic darkling beetle (Coleoptera: Tenebrionidae: Kuhitangiinae: Foranotini trib. nov.) from a cave in Southern Zagros, Iran. Zootaxa 4338 (1): 163–172. https://doi.org/10.11646/zootaxa.4338.1.9

Pierre F. 1958. Ecologie et peuplement entomologique des sables vifs du Sahara Nord-Occidental. Publications du Centre de Recherches sahariennes. Série biologie, Paris.

Pierre F. 1964. Storthocnemis nouveaux de la zone sahélienne et du Sahara. Remarques concernant les Leucolaephini trib. nov. (Col. Tenebrionidae). Bulletin de l’Institut français d’Afrique noire (Série A) 26: 865–874.

Pierre F. 1968. Contribution à la faune de l’Iran 12. Étude écologique et biogéographique des ténébrionides Pimeliinae s. nov. des régions désertiques et semi-désertiques de l’Iran. Annales de la Société entomologique de France (N.S.) 4: 997–1036.

Reitter E. 1887. Insecta in itinere Cl. N. Przewalskii in Asia Centrali novissime lecta. IX. Tenebrionidae. Horae Societatis Entomologicae Rossicae 21: 355–389.

Reitter E. 1889. Insecta, a cl. G. N. Potanin in China et in Mongolia novissime lecta. XIII. Tenebrionidae. Horae Societatis Entomologicae Rossicae 23: 678–710.

Reitter E. 1893. Bestimmungs-Tabelle der unechten Pimeliden aus der palaearctischen Fauna. Verhandlungen des Naturforschenden Vereines in Brünn 31 (1892): 201–250.

Reitter E. 1900. Neue Coleopteren aus Europa, den angrenzenden Ländern, russisch Asien und der Mongolei. Deutsche Entomologische Zeitschrift 1899: 273–281.

Ren G.-D. & Ba Y.-B. 2009. Revision of the genus Ocnera Fischer von Waldheim, 1822 (Coleoptera: Tenebrionidae: Pimeliini) from China. Caucasian Entomological Bulletin 5 (2): 221–225. https://doi.org/10.23885/1814-3326-2009-5-2-221-225

Ren G.-D. & Dong S. 2001. The beetles of the tribe Platyopini in China and description of two new species (Coleoptera: Tenebrionidae). Journal of Hebei University (Natural Science Edition) 21 (3): 296–302.

Ren G.-D. & Yu Y. 1999. The Darkling Beetles from Deserts and Semideserts of China (Coleoptera: Tenebrionidae). Hebei University Publishing House, Baoding. [In Chinese.]

Ren G.-D., Wang X.-P. & Ma F. 1993. Two new species and one new record of Platyope F.-W. from China. (Coleoptera: Tenebrionidae). Journal of the Ningxia Agricultural College 14 (Supplement): 44–49. [In Chinese.]

Schuster A. 1935. Die Tenebrioniden-Ausbeute Dr. Gabriels in Persien. Koleopterologische Rundschau 21: 20–29.

Schuster A. 1938. Neue Tenebrioniden (Col.) von Persisch-Belutschistan. Koleopterologische Rundschau 24: 77–90.

Schuster A. 1940. Die Tenebrioniden (Col.) des Museums Hoang ho-Pei ho in Tientsin. Koleopterologische Rundschau 26 (1/2): 15–24.

Schuster A. & Reymond A. 1937. Quatre nouveaux ténébrionides provenant de la Mission Citroën-Centre-Asie (Col.). Bulletin de la Société entomologique de France 42: 234–238.

Semenov A. P. 1889. Diagnoses Coleopterorum novorum ex Asia centrali et orientali. Horae Societatis Entomologicae Rossicae 24 [1889–1890]: 193–226.

Semenov A.P. 1891. Diagnoses coleopterorum novorum ex Asia centrali et orientali. Horae Societatis Entomologicae Rossicae 25: 262–382.

Semenov A.P. 1893. Symbolae ad cognitionem pimeliidarum. I–III. Horae Societatis Entomologicae Rossicae 27 [1892–1893]: 249–264.

Semenov A.P. 1903a. Coleoptera novae faunae turanicae. Revue russe ďEntomologie 3: 8–13.

Semenov A.P. 1903b. Analecta coleopterologica. iii. Revue russe d’Entomologie 3: 169–173.

Semenov-Tjan-Shansky A.P. 1907. Synopsis generum tribus Platyopinorum (Coleoptera, Tenebrionidae Pimeliini). Horae Societatis Entomologicae Rossicae 38 [1907–1908]: 175–184.

Skopin N.G. 1959. Beetle larvae of the tribe Platyopini (Coleoptera, Tenebrionidae). Zoologicheskii zhurnal 38: 69–75. [In Russian.]

Skopin N.G. 1960. Neue Tenebrioniden aus Zentralasien. I. Annales Historico-Naturales Musei Nationalis Hungarici 52: 295–311.

Skopin N.G. 1961. Materials on the fauna and ecology of darkling beetles (Coleoptera, Tenebrionidae) of South-east Kazakhstan. Trudy Nauchno-Isledovatelskogo Instituta Zashchtity Rasteniy Kazakhstanskoy Akademii Selskokhozyastvennykh Nauk 6: 172–208. [In Russian.]

Skopin N.G. 1962. Larvae of the subfamily Pimeliinae (Coleoptera, Tenebrionidae). Trudy Nauchno-Issledovatelskogo Instituta Zashchity Rastenii Kazakhstanskoy Akademii Selskokhozyastvennykh Nauk 7: 191–298. [In Russian.]

Skopin N.G. 1964. Neue Tenebrioniden aus Zentralasien. iii, nebst einigen systematischen sowie synonymischen Bemerkungen. Annales Historico-Naturales Musei Nationalis 56: 389–412.

Skopin N.G. 1968a. Eine neue Tenebrioniden-Gattung und -Art aus Zentralasien (Coleoptera-Tenebrionidae-Pimeliini). Entomologische Arbeiten aus dem Museum G. Frey 19: 300–304.

Skopin N.G. 1968b. Darkling-beetles (Coleoptera, Tenebrionidae) of Southern Kazakhstan and their economic importance. Trudy Kazakhckogo Nauchno-Issledovatelskogo Instituta Zashchity Rastenii 10: 73–114. [In Russian.]

Skopin N.G. 1971. Monographische Übersicht der Arten der Gattungen Diesia Fisch.-W. und Platyesia gen. nov. Entomologische Abhandlungen des Staatlichen Museum für Tierkunde in Dresden 38: 325–349.

Skopin N.G. 1972. Revision der Tenebrioniden-Gattung Stalagmoptera Solsky, 1875 (Coleoptera). Entomologische Arbeiten aus dem Museum G. Frey 23: 298–313.

Skopin N.G. 1973. Revision der Tenebrioniden-Gattungsgruppe Trigonoscelis-Sternoplax (Coleoptera). Entomologische Arbeiten aus dem Museum G. Frey 24: 104–185.

Skopin N.G. 1974. Revision der Gattung Pterocoma Dejean; Solier, 1836 (Coleoptera, Tenebrionidae). Entomologische Abhandlungen des Staatlichen Museum für Tierkunde in Dresden 40: 127–164.

Soldati L. 2009. Darkling Beetles of Qatar (Insecta: Coleoptera: Tenebrionidae). Natura optima dux Foundation, Warszawa.

Solier A.J.J. 1836. Essai sur les collaptèrides (suite). 4e Tribu. Pimélites. Annales de la Société Entomologique de France 5 [1835–1836]: 5–200.

Solsky S.M. 1876. Matériaux pour l’entomographie des provinces asiatiques de la Russie. Horae Societatis Entomologicae Rossicae 11 [1875–1876]: 253–272, 273–299.

Strand E. 1917. Übersicht der in Gistel’s ‘Achthundert und zwanzig neue oder unbeschriebene wirbellose Thiere’ (1857) behandelten Insekten. Archiv für Naturgeschichte (A) 82 (5) [1916]: 75–101.

Turner J.S. & Lombard A.T. 1990. Body color and body temperature in white and black Namib Desert beetles. Journal of Arid Environments 19 (3): 303–315. https://doi.org/10.1016/s0140-1963(18)30795-x

Wu W., Fan Z. & Huang R. 2005. Three new species of Platyope (Coleoptera: Tenebrionidae) from Xinjiang, China. Entomotaxonomia 27 (4): 283–288.

Published
2022-04-05
How to Cite
Chigray, S. N., Nabozhenko, M. V., Chigray, I. A., & Abakumov, E. V. (2022). A revision of the Palaearctic Pimeliini (Coleoptera: Tenebrionidae): a comparative analysis and systematic position of Eastern European and Asian taxa with dorso-lateral eyes. European Journal of Taxonomy, 809(1), 1–71. https://doi.org/10.5852/ejt.2022.809.1719
Section
Monograph