News from Baltic amber: a new tribe of extinct Paleogene ironclad beetles (Coleoptera: Zopheridae: Zopherinae).

A new extinct monotypic genus of the ironclad beetles, Zopheromimus auriborussiensis gen. et sp. nov., is described and illustrated based on an inclusion in Baltic amber. The new Eocene fossil taxon is placed in Zopheromimini trib. nov., a new tribe within the subfamily Zopherinae, and compared with related extant beetles of the subfamily. The genus Yantaroxenos Nabozhenko, Kirejtshuk et Merkl, 2016 is transferred from the tribe Belopini (Tenebrionidae: Lagriinae) to the newly described tribe of Zopheridae.

Oracula campbelli sp. n.-the first fossil darkling beetle (Coleoptera: Tenebrionidae: Alleculinae) from the late Eocene Rovno amber (Ukraine).

A new species of the currently Oriental alleculine genus Oracula Novák, 2019 is described from the Eocene Rovno amber, Ukraine: O. campbelli sp. n. This is the first report of a representative of the family Tenebrionidae from this Lagerstätte. Extant species of Oracula occur in Nepal, southern China (Yunnan), Thailand, Laos and Indonesia. The newly discovered extinct species is similar to extant O. amica Novák, 2019 from Nepal.

A new tribe, genus and species of the tenebrionid subfamily Lagriinae (Coleoptera: Tenebrionidae) from the Eocene Baltic amber.

A new genus and species are described from the Eocene Baltic amber: Archaeoluprops groehni gen. et sp. nov. This fossil darkling beetle of the subfamily Lagriinae (Tenebrionidae) is externally similar to species of the tribes Lupropini and Goniaderini having the robust pubescent body and estriate elytra, but differs in the absence of intersegmental membranes between abdominal ventrites 3-5 (tentyrioid hinging) and accordingly the primary absence of defensive glands. Combination of plesiomorphic and apomorphic characters and other differences from all extinct and extant lagriine tribes allows to establish the new tribe Archaeolupropini trib. nov. (type genus Archaeoluprops), which is probably a basal group in relation to the non-basal clade Lagriini-Chaerodini-Lupropini-Pycnocerini-Goniaderini-(probably Gonialaenini) + Laenini. All Baltic amber lagriine tribes are absent in extant faunas of middle latitudes of Europa.

Contribution to the knowledge of genus Nalassus Mulsant, 1856 (Coleoptera: Tenebrionidae) from the Eastern Caucasus, Russia.

A new species of darkling beetle of the tribe Helopini is described from the Eastern Caucasus (Russia): Nalassus magomedrasuli Nabozhenko sp. n. It was originally described as the Dagestan population of N. kalashiani Nabozhenko, 2001 based on the similar structure of the male aedeagus. The new species is similar to N. kalashiani in that it differs in the structure of the eyes, male antennae, the shape of the pronotum, the scutellar shield, the male abdomen and habitats. Nalassus magomedrasuli occurs in xerophytic limestone mountain phryganoid steppes in Dagestan, whereas N. kalashiani is distributed in wet alpine meadows in the Chechen Republic and Ingushetia (Russia). The female of N. kalashiani collected in the State Natural reserve Erzi (Ingushetia) is described for the first time.

Review of the subgenus Turcmenicola Bogatchev, 1952, the genus Colposcelis Dejean, 1834 (Coleoptera: Tenebrionidae: Tentyriini).

The subgenus Turcmenicola Bogatchev, 1952 (genus Colposcelis Dejean, 1834) is reviewed. Representatives of the studied taxon occur in the most lifeless rocky, gravelly and clayey areas of the deserts of Middle Asia (Ustyurt plateau, Kyzylkum desert, foothills of southern Kugitang, Betpak-Dala desert). Turcmenicola comprises five species from which two are described here as new: C. ustyurtensis Bekchanov & Nabozhenko sp. n. (Uzbekistan, Karakalpakstan, eastern part of Ustyurt plateau) and C. khudaybergani Bekchanov, Nabozhenko & Bekchanova, sp. n. (Uzbekistan, Karakalpakstan, Amudarya and Beruni districts). The following new synonymy is proposed: Colposcelis jachontovi (Bogatchev, 1952) = Colposcelis kerzhneri Medvedev, 1970, syn. n. A key to species of Turcmenicola is given.

A taxonomic review of the genus Euboeus Boieldieu, 1865 (Coleoptera: Tenebrionidae: Helopini) of the Caucasus, Iran and Turkmenistan.

A taxonomic review of the genus Euboeus from the Caucasus, Iran and Turkmenistan is given. All revised species belong to the subgenus Pelorinus Vauloger de Beaupr, 1900 and can be divided into two groups: the tenebricosus species-group and the huedepohli species-group. The first one consists of mainly forest low mountain species, while the second one includes high mountain, alpine or steppe taxa. Only one species, E. horasanicus (Medvedev, 1976), has mixed characters and unknown habitats. In total, 11 species of the genus are known from Iran, one species is distributed in Iran and Turkmenistan. Euboeus prometheus is hereby excluded from the fauna of Azerbaijan, the species occurs on Iranian part of Talysh Mts. Euboeus quadricollis (Mntris, 1832) is recorded from Iran for the first time. The following new species are described: Euboeus grimmi sp. nov. (Iran, Western Elburs), E. kasatkini sp. nov. (Iran, Central Elburs), E. krivokhatskyi sp. nov. (Iran, Western Elburs and Zagros), E. merkli sp. nov. (Iran, Central Elburs), E. arzanovi sp. nov. (Iran, Central Elburs), E. zubovi sp. nov. (Iran, Zagros, Kordestan Prov.), E. kalashiani sp. nov. (Armenia, Azerbaijan: Nakhchivan, North-Western Iran). The following new synonimies are established: Helops corrugosus Seidlitz, 1895 = Probaticus parthorum G. Medvedev, 1976, syn. nov.; Helops vicinus Allard, 1876 = Probaticus (Pelorinus) medvedevi Abdurakhmanov et Nabozhenko, 2011, syn. nov. The lectotype for Helops corrugosus is designated to fix the taxonomic status of that species. In addition, illustrated identification keys for males and females of Caucasian, Iranian and Turkmen Euboeus are provided.

Contribution to the knowledge of the genus Dendarus Dejean, 1821 (Coleoptera: Tenebrionidae: Blaptinae: Dendarini) from Iran, Turkmenistan and some adjacent territories.

The taxonomic review of the genus Dendarus Dejean, 1821 from Iran and Turkmenistan is given. In total, five species are distributed in Iran and four species in South Turkmenistan; three species occur in both countries. The new species D. matthewsi sp. n. is described from Iran and compared with D. simplex Seidlitz, 1893. One taxon is resurrected from a synonymy with D. transcaspicus: D. vagabundus sp. resurr. As a result, one Afghan subspecies of D. transcaspicus belongs to another species: D. vagabundus afghanicus Kaszab, 1973. The following new synonymy is proposed: Dendarus vagabundus vagabundus Reitter, 1904 = D. transcaspicus medvedevi Kaszab, 1973, syn. n. Lectotypes of Dendarus armeniacus Baudi di Selve, 1876 and D. leonhardi Schuster, 1940 are designated. The following erroneous interpretations of Iranian and Turkmenian species are indicated: D. vagabundus = D. armeniacus sensu G. Medvedev = D. transcaspicus sensu Kaszab; D. armeniacus = D. simplex sensu G. Medvedev. New distribution data are given: D. armeniacusN Iran (excluded from faunistic lists of Armenia, Afghanistan, Turkmenistan and Turkey); D. crenulatus (Mntris, 1832)Turkey, Transcaucasia, the North Caucasus (Russia: Dagestan, Chechen Republic), N Iran, Turkmenistan (new record for the country); D. leonhardiNE Iran, S Turkmenistan; D. matthewsi Nabozhenko sp. n.W Iran (Zagros); D. transcaspicus Brancsik, 1899S Turkmenistan (Kopetdag); D. vagabundus vagabundus Reitter, 1904N Iran, S Turkmenistan (Kopetdag). Two species must be excluded from Iranian faunistic list: D. simplexTurkey, Syria, Iraq; D. extensus (Faldermann, 1837)Georgia, Armenia, W Azerbaijan, Turkey (new record for the country). Well-illustrated keys to males and females of Iranian and Turkmenian species of the genus are given.

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.

A new species is described from the Central part of the North Caucasus (Russia): Blaps caucasica M. Nabozhenko et I. Chigray sp. n. This taxon was interpreted earlier as B. scabriuscula subalpina Mntris, 1832, but both our morphological and molecular genetic analyses showed that it is in fact a separate new species. The following new synonymies are proposed: Blaps verrucosa Adams, 1817 = Blaps scabriuscula Mntris, 1832 syn. n., = Blaps montana Motschulsky, 1839 syn. n.; Blaps lethifera Marsham, 1802 = Blaps pterotapha Mntris, 1832 syn. n. The rank of one species is reinstated: Blaps subalpina Mntris, 1832 stat. resurr. The name Blaps sinuatocollis Solier, 1848 was unambiguously proposed for an infrasubspecific entity and must be excluded from zoological nomenclature. A phylogenetic hypothesis was reconstructed using mitochondrial Cytochrome c oxidase subunit I (COI) sequences for some closely related taxa from the North Caucasus and Ciscaucasia: B. caucasica sp. n., B. lethifera, B. subalpina and B. verrucosa. The resulting tree supports the assignment of two specimens from the Lower Don area (Rostov Region of Russia), earlier interpreted as B. scabriuscula subalpina, to B. lethifera. All known Blaps from Russia and Eastern Europe (countries of the former USSR) are illustrated, and keys on males and females are provided. The status of an invasive species Blaps aff. oblonga Kraatz, 1883 from South Siberia is discussed.

The first described darkling beetle of the tribe Metaclisini (Coleoptera: Tenebrionidae) from Eocene Baltic amber.

The description of an extinct species of Metaclisa Jacquelin du Val, 1861 (Tenebrionidae) is presented. This genus and the tribe Metaclisini are recorded as fossils for the first time, from Eocene Baltic amber. The new species Metaclisa ottoi sp. nov. belongs to the subgenus Trichometaclisa subgen. nov. and differs from all other Metaclisini in possessing short, fine recumbent setation on the pronotum and elytra; in addition, the prosternal process in Metaclisa ottoi sp. nov. is roundly bent down and weakly projected behind the procoxae, which differs from extant species.

Two new species and new records of lichen-feeding darkling beetles (Coleoptera: Tenebrionidae: Helopini) from Turkey with notes on bionomics and trophic relations.

Turkonalassus mavi M. Nabozhenko B. Keskin, sp. n. and Odocnemis rufocruralis M. Nabozhenko B. Keskin, sp. n., two distinctive darkling beetle species of the tribe Helopini, are described from the Southeastern Anatolia region of Turkey, based on both their external morphology and genital structures. Turkonalassus mavi sp. n. differs from all congeners by the bluish dorsal surface of the body, the pronotum widest before middle, and wide merged baculi of the median lobe of the aedeagus. Odocnemis rufocruralis sp. n. belongs to the praelonga species-group. Odocnemis rufocruralis sp. n. is similar to O. torosica Nabozhenko Keskin, 2016, O. terminassianae (Nabozhenko, 2011) and O. kakunini Nabozhenko Keskin, 2016 but differs from all three taxa by the reddish-brown body with red legs, male apical maxillary palpomere wider than in female, elevate and narrowly separate male protrochanters, and the structure of the very long and narrow apical piece of the aedeagus. New data on distribution, bionomics, and trophic relations of several species of Helopini from Turkey are given. Host lichens are determined for nine species. The majority of studied adult beetles feed on lichens from the families Physciaceae and Parmeliaceae. Some examined taxa feed on Cladoniaceae. Feeding on crustose lichens is registered for the first time for Coleoptera, specifically Odocnemis rufocruralis sp. n. was observed to feed on Pertusaria sp. (on Prunus) in Mardin Province.

The first fossilized comb-clawed beetle of the genus Asiomira Dubrovina, 1973 (Coleoptera: Tenebrionidae: Alleculinae) from Baltic Amber and notes on the distribution of extant species of the genus.

A new species of comb-clawed beetles of the genus Asiomira Dubrovina, 1973 (A. dubrovinae sp. n.) is described from the Eocene Baltic amber. This newly descovered fossil species displays typical generic characters and is the most similar to the extant Asiomira ophtalmica (Seidlitz, 1896). Both species share a similar shape of the pronotum and the flattened posterior angles of the pronotal disc. Asiomira dubrovinae sp. n. can be distinguished from A. ophtalmica by the smaller body size (4.8 mm in contrast to 68.12 mm), more serrate antennomeres, and finer and sparser pronotal punctation. Extant species of the genus are distributed in the arid landscapes of Central Asia with the highest diversity occurring in Tajikistan. Therefore the discovery of a new fossil species from Eocene Baltic amber suggests that Asiomira could have a wider range, and the modern distribution of this group is the result of a later secondary expansion of the ancestral representatives that survived in mid-mountain areas with arboreal and shrub vegetation. Relevant corrections to the distribution of extant species, Asiomira ophtalmica (Seidlitz, 1896) and Asiomira firjusana (Dubrovina, 1973), are given.

The oldest fossil darkling beetle of the genus Neomida Latreille, 1829 (Coleoptera: Tenebrionidae) from Eocene Baltic amber examined with X-ray microtomography.

Based on a single well-preserved specimen from Eocene Baltic amber, a new tenebrionid beetle Neomida groehni Nabozhenko et Bukejs sp. nov. (Diaperinae: Diaperini) is described and illustrated using phase-contrast X-ray microtomography. This oldest representative of the genus differs from all known extant and extinct congeners by the dorsally very weakly convex pronotum with undulate lateral margins (while other Neomida have a pronotum that is strongly convex along its transverse axis, with evenly rounded lateral margins); distinct, right posterior angles of pronotum; and elevated sutural area of the elytra.

To the knowledge of the Near East species of the genus Odocnemis Allard, 1876 (Coleoptera: Tenebrionidae: Helopini).

A brief review of the eight species of the genus Odocnemis Allard, 1876 from the Near East (Cyprus, Syria, Lebanon, Israel, Jordan, Egypt) is given. All species (excluding probably O. intrusicollis from Cyprus) belong to the praelonga species-group. Odocnemis evae sp. n. is described from Lebanon. The new species is similar to O. praelonga Baudi di Selve, 1876 by the presence of a mucron in females and to O. moabitica Sahlberg, 1908 by the presence of teeth and granules on inner side of metatibiae (at least in large specimens). It differs from both species by the shiny body, the presence a tooth on the outer margin of male pro- and mesotrochanters and by the structure of the aedeagus (very narrow apical piece with ball-like apex). Redescriptions, full bibliographic information and key to the species are given. Odocnemis punctata Allard, 1876 is recorded for Israel for the first time.

Contribution to the knowledge of Mediterranean lichen-feeding darkling beetles of the genus Stenohelops Reitter, 1922 (Coleoptera: Tenebrionidae).

Species of the genus Helopelius Reitter, 1922 (type species: Stenomax aeneipennis Allard, 1876) were known from Rhodes and North Africa. We found that two species described from Rhodes belong to the genus Stenohelops Reitter, 1922, and Helopelius can be interpreted as a subgenus within latter, guided by the article 23.2 of ICZN. The following synonyms are proposed: Helopelius, stat. n. (from genus to subgenus) = Stenomaleis Español, 1957, syn. n.; Stenohelops Reitter, 1922 = Gunarellus Reitter, 1922, syn. n.; Stenohelops (Helopelius) aeneipennis (Allard, 1876) = Helopelius disgregus Reitter, 1922, syn. n., = Gunarus gayirbegi Nabozhenko Keskin, 2009, syn. n. As a result, the following species are transferred from the former genus Helopelius to the genus Stenohelops: Stenohelops (Helopelius) nodifer (Kraatz, 1880), comb. n.; Stenohelops (Helopelius) otini (Antoine, 1949), comb. n.; Stenohelops (Helopelius) subsinuatus (Antoine, 1951), comb. n.; Stenohelops (Helopelius) verrucosus (Vauloger de Beaupré, 1900), comb. n.; Stenohelops (Helopelius) zaianus (Antoine, 1949), comb. n. Thus, the subgenus Helopelius contains 11 species from three isolated geographical exclaves: Western Mediterranean, Eastern Mediterranean and China. Lectotypes of Stenomax aeneipennis Allard, 1876, Helops gratus J. Frivaldszky, 1894 and Cylindrinotus (Helopelius) disgregus Reitter, 1922 are designated. Data on fossil species of the genus and the allied extinct taxa, as well as on bionomics of extant species of the subgenus Helopelius are presented. The check-list for extant and extinct species of Stenohelops is given.

Gonialaenini, a new tribe of Lagriinae (Coleoptera: Tenebrionidae) from Eocene Baltic Amber.

Based on well preserved specimen, Gonialaena groehni gen. et sp. nov. (Coleoptera: Tenebrionidae) is described from Eocene Baltic amber and assigned to Gonialaenini trib. nov. within the subfamily Lagriinae due to the combination of the following characters: labrum elongate; antennal sensilla simple; procoxal cavities closed at least externally; narrow membranes present between abdominal ventrites 3-5; mesocoxal cavities partly closed by mesepimeron, trochantine present; ovipositor of primitive lagrioid type-digitate fourth coxite lobe with gonostyles apically. Gonialaena gen. nov. combines characters of various Lagriinae tribes: humeral angles rounded, formed by basally widened epipleura as in Goniaderini, Pycnocerini and Lupropini; narrow acute intercoxal process of abdominal ventrite 1 as in Belopini, Cossyphini and Lagriini; absence (or strong reduction) of metathoracic wings as in most Adeliini and some Lupropini; large, reniform eyes as in Goniaderini, Pycnocerini, Lagriini and many Adeliini; present intersegmental membranes between abdominal ventrites 3-5 as in all lagriine tribes except for Belopini and Cossyphini; subcylindrical terminal maxillary palpomere as in Belopini and Cossyphini. Thus, the position of Gonialaenini trib. nov. within Lagriinae is not fully understood, but combination of certain features (humeral angles rounded but distinct and vertical border present at elytral base, inconspicuous elytral striae, large reniform eyes, narrow intersegmental membranes) points on morphological similarity with Goniaderini, especially some Anaedus Blanchard, 1845 with indistinct elytral striae and related genera.

A new subgenus and species of the genus Hedyphanes Fischer von Waldheim, 1820 (Coleoptera: Tenebrionidae: Helopini) from Israel and Egypt.

A new species, Hedyphanes chikatunovi sp. n., is described from southern Israel (Negev, Gerofit) and Egypt (Northern Sinai). These localities represent the southwestern limit of the distribution of the genus. The new species is placed in a new monotypic subgenus, Microhedyphanes subgen. n.