Glotov.indd UDC UDC 595.763(477:292.451/.454) THE GENUS ATHETA (COLEOPTERA, STAPHYLINIDAE, ALEOCHARINAE) IN THE UKRAINIAN CARPATHIANS S. Glotov1,5*, K. Hushtan1,2, H. Hushtan1, N. Koval3,4, V. Diedus3 1State Museum of Natural History NAS of Ukraine, Teatralna st.,18, Lviv, 79008 Ukraine 2Ecological College of Lviv National Agrarian University, Zamarstynivska st., 167, Lviv, 79068 Ukraine 3Uzhgorod National University, A. Voloshyn st., 32, Uzhgorod, 88000 Ukraine 4Uzhansky National Nature Park, Nezalezhnosti st., 7, village Velykyi Bereznyi, Ukraine 5 InstitutionNational Antarctic Scientifi c Center, Ministry of Education and Science of Ukraine, 16 Taras Shevchenko blvd., Kyiv, 01601 Ukraine 5*Corresponding author E-mail: sergijglotov@gmail.com S. Glotov (https://orcid.org/0000-0003-3986-0844) K. Hushtan (https://orcid.org/0000-0002-4005-9029) H. Hushtan (https://orcid.org/0000-0001-6999-6043) V. Diedus (https://orcid.org/0000-0002-0957-5156) Th e Genus Atheta (Coleoptera, Staphylinidae, Aleocharinae) in the Ukrainian Carpathians. Glotov, S., Hushtan, K., Hushtan, H., Koval, N., Diedus, V. — Th e Carpathian species of the genus Atheta Th omson, 1858 are reviewed. Th e list contains 65 species, of which 8 species: Atheta kochi Roubal, 1937, A. interme- dia (Th omson, 1852), A. setigera (Sharp, 1869), A. foveicollis (Kraatz, 1856), A. luctuosa (Mulsant & Rey, 1853), A. cribrata (Kraatz, 1856), A. mortuorum Th omson, 1867, A. picipes (Th omson, 1856), are recorded for Ukraine for the fi rst time. Species composition, data on bionomics and distribution genus Atheta in the studied region are discussed. K e y w o r d s : Coleoptera, Staphylinidae, Aleocharinae, Atheta, new records, Ukraine, Carpathians. Introduction Th e rove beetle genus Atheta Th omson, 1858, includes 1900 described species in the world fauna (Orlov, 2019). Of them, 1500 species are known from the Palaearctic Region (Schülke & Smetana, 2015); including 78 species in the fauna of Ukraine (Glotov, 2021). Th e taxonomic and faunistic knowledge of Atheta is extremely fragmentary both at the global and Palaearctic scale. Th e fauna of Northern and Central Europe is better studied, with comprehensive papers including keys to species known to occur in these territories (Münster, 1925; Brundin, 1954 a, b, c, d; Strand & Vik 1964; Palm, 1972; Lohse, 1974). Th e Eastern and Southern Europe is less studied. Th ere were no special publications on the fauna of this genus in Ukraine. Th e present paper is an attempt to fi ll the gap in the local knowledge of Atheta, an ecologically notable genus of the rove beetles. Th e aim of this paper is to overview species composition of Atheta in the Ukrainian Carpathians. As a result of extensive fi eld investigation of rove beetles (Coleoptera, Staphylinidae) in the Ukrainian Carpathians in the last ten years more than 150 species were added to the list. Th is paper provides an overview of the genus Atheta species in the Ukrainian Carpathians. Zoodiversity, 56(1):91–110, 2022 DOI 10.15407/zoo2022.02.091 92 S. Glotov, K. Hushtan, H. Hushtan, N. Koval, V. Diedus An analysis of all available collection and literature information on the fi ndings of species of the genus Atheta has shown that 65 species were known in the fauna of the Ukrainian Carpathians. Th ey belong to 19 sub- genera (Anopleta  — 2 species, Atheta  — 22 species, Badura  — 3  species, Bessobia  — 2 species, Chaetida  — 1 species, Coprothassa  — 1 species, Datomicra  — 4 species, Dimetrota  — 11 species, Microdota  — 7 species, Mycetota — 1 species, Oreostiba — 1 species, Oxypodera — 1 species, Pachyatheta — 2 species, Rhagocneme — 1 species, Tetropla — 2 species, Th inobaena — 1 species, Traumoecia — 1 species, Trochanterella — 1 species, Xenota  — 1 species). Among the 65 species of the genus Atheta, presence of 45  species is confi rmed by the collection material and 20 species were known for the region only from literature data, among them 8 species (Atheta kochi (Roubal, 1937), A. intermedia (Th omson, 1852), A. setigera (Sharp, 1869), A. foveicollis (Kraatz, 1856), A. luctuosa (Mulsant & Rey, 1853), A. cribrata (Kraatz, 1856), A. mortuorum (Th omson, 1867), A. picipes (Th omson, 1856) are reliably recorded for Ukraine for the fi rst time in this paper. Th e fi rst fundamental contribution on the list of the genus Atheta was made by A. Łomnicki (1868, 1884, 1890, 1891, 1913), who at that time actively studied the beetle fauna of Western Ukraine and provided a list of 42 species for the Ukrainian Carpathians, including: Atheta corvina (Th omson, 1856), A. aquatica (Th om- son, 1852), A. brunneipennis (Th omson, 1852), A. castanoptera (Mannerheim, 1830), A. crassicornis (Fabricius, 1792), A. divisa (Märkel, 1844), A. hypnorum (Kiesenwetter, 1850), A. fungicola (Th omson, 1852), A. gramini- cola (Gravenhorst, 1806), A. euryptera (Stephens, 1832), A. laevicauda (J. Sahlberg, 1876), A. oblita (Erichson, 1839), A. pilicornis (Th omson, 1852), A. ravilla (Erichson, 1839), A. vaga (Heer, 1839), A. xanthopus (Th omson, 1856), A. cauta (Erichson, 1837), A. occulta (Erichson, 1837), A. monticola (Th omson, 1852), A. longicornis (Gravenhorst, 1802), A. melanaria (Mannerheim, 1830), A. celata (Erichson, 1837), A. sordidula (Erichson, 1837), A. zosterae (Th omson, 1856), A. atramentaria (Gyllenhal, 1810), A. cadaverina (Brisout de Barnev- ille, 1860), A. europaea (Likovský, 1984), A. intermedia (Th omson, 1852), A. laevana (Mulsant & Rey, 1852), A. marcida (Erichson, 1837), A. nigripes (Th omson, 1856), A. putrida (Kraatz, 1856), A. subtilis (Scriba, 1866), A. amicula (Stephens, 1832), A. atomaria (Kraatz, 1856), A. foveicollis (Kraatz, 1856), A. laticollis (Stephens, 1832), A. tibialis (Heer, 1839), A. cribrata (Kraatz, 1856), A. subsinuata (Erichson, 1839), A. nigritula (Graven- horst, 1802), A. vestita (Gravenhorst, 1806), A. picipes (Th omson, 1856), A. cribripennis (J. Sahlberg, 1890). Information about the fi ndings of the genus Atheta in the Ukrainian Carpathians in the territories of modern Lviv and Ivano-Frankivsk Regions are contained in the works of L. Miller (1868), who studied beetles in Kolomyia, Kosiv, Verkhovyna, Bystrets River and in Gadzhyna District and recorded fi ve species: A. cas- tanoptera Mannerheim, 1830, A. fungicola (Th omson, 1852), A. euryptera (Stephens, 1832), A. tibialis (Heer, 1839), A. nigritula (Gravenhorst, 1802). In addition, M. Nowicki (1873), recorded fi ve species of Atheta in the territories covering the north-east- ern macroslope of the Ukrainian Carpathians: Atheta castanoptera (Mannerheim, 1830), A. fungicola (Th om- son, 1852), A. euryptera (Stephens, 1832), A. tibialis (Heer, 1839), A. nigritula (Gravenhorst, 1802) and 3 species are cited by J. Weise (1876): Atheta castanoptera (Mannerheim, 1830), A. longicornis (Gravenhorst, 1802), A. tibialis (Heer, 1839). M. Rybiński (1903) mentioned three species: Atheta laevicauda (J. Sahlberg, 1876), A. atramentaria (Gyllenhal, 1810), A. nigripes (Th omson, 1856), and E. Lokay (1912) eight species: Atheta al- locera (Eppelsheim, 1893), A. autumnalis (Erichson, 1839), A. diversa diversa (Sharp, 1869) A. hypnorum (Kie- senwetter, 1850) A. laevicauda (J. Sahlberg, 1876), A. macrocera (Th omson, 1856), A. laevana (Mulsant & Rey, 1852), A. cribripennis (J. Sahlberg, 1890) and four species are cited in the works of J. Fleischer and co-authors (Fleischer et al., 1922, 1924, 1925) including: Atheta aquatica (Th omson, 1852), A. contristata (Kraatz, 1856), A. cadaverina (Brisout de Barneville, 1860), A. subtilis (Scriba, 1866). C. Hormuzaki (1888, 1891) listed six species Atheta from the south-western macroslope of the Ukrai- nian Carpathians and Transcarpathian Region: Atheta occulta (Erichson, 1837), A. melanaria (Mannerheim, 1830), A. celata (Erichson, 1837), A. sordidula (Erichson, 1837), A. amicula (Stephens, 1832), A. laticollis (Ste- phens, 1832). D. Kuthy (1896) recorded additional six species from this territory: Atheta boletophila (Th omson, 1856), A. oblita (Erichson, 1839), A. melanaria (Mannerheim, 1830), A. fi morum (Brisout de Barneville, 1860), A. subsinuata (Erichson, 1839), A. liturata (Stephens, 1832). Th e most signifi cant work for this area of the modern territory of Slovakia and Transcarpathia (Subcarpathian Rus’) was made by J. Roubal (1930), in which he mentioned 17 species belonging to the genus Atheta: A. aquatica (Th omson, 1852), A. hypnorum (Kiesen- wetter, 1850), A. hypnorum (Kiesenwetter, 1850), A. incognita (Sharp, 1869), A. pilicornis (Th omson, 1852), A. vaga (Heer, 1839), A. spatula (Fauvel, 1875), A. sordidula (Erichson, 1837), A. zosterae (Th omson, 1856), A. cadaverina (Brisout de Barneville, 1860), A. europaea (Likovský, 1984), A. laevana (Mulsant & Rey, 1852), A. putrida (Kraatz, 1856), A. subrugosa (Märkel & Kiesenwetter, 1848), A. aegra (Heer, 1841), A. excelsa (Ber- nhauer, 1911), A. tibialis (Heer, 1839). O. Marcu (1936) listed 5 species: Atheta divisa (Märkel, 1844), A. oblita (Erichson, 1839), A. ravilla (Erichson, 1839), A. monticola (Th omson, 1852), A. subtilis (Scriba, 1866). More recently, O. Mateleshko (2008) recorded three additional species: Atheta crassicornis (Fabricius, 1792), A. oblita (Erichson, 1839), A. vaga (Heer, 1839). Material and methods Th e material consists of 311 examined specimens deposited in the following collections examined through the kindness of: State Museum of Natural History of the National Academy of Sciences of Ukraine, Lviv (SMNH); I. I. Schmalhausen Institute of Zoology National Academy of Sciences of Ukraine, Kyiv (SIZK); 93 Th e Genus Atheta (Coleoptera, Staphylinidae, Aleocharinae) in the Ukrainian Carpathians Zoological Museum of Taras Shevchenko National University, Kyiv (ZMTSNU); collection of V.  O.  Chu- mak, Uzhgorod (KCHM); collection of N. P. Koval, Velykyi Bereznyi (KKOV); collection of Yu. V. Kanarsky deposited in SMNH. Vast comparative material from Western Europe was studied through the kindness of A. Solodovnikov Natural History Museum of Denmark (ZMUC), Copenhagen. Current taxonomic status, nomenclature of the species and distribution follow the “Catalogue of Palaearc- tic Coleoptera” (Schülke & Smetana, 2015) taking into account taxonomic changes based on the molecular phylogenetic analysis (Elven et al., 2010, 2012). Abbreviations of the collecting localities with their coordinates are as follows: Ivano-Frankivsk Region (IFR): An- heliv tract — Rozhniativskyi District, Osmoloda village (48.672552 N, 24.039903 E); Kosiv — Kosiv town (48.319828 N, 25.097768 E); mountain Dantsyzh  — Nadvirnianskyi District, Vorokhta village (48.135544 N, 24.530363 E); mountain range Pozhyzhevska — Nadvirnianskyi District, Vorokhta village (48.144413 N, 24.523402 E); mountain Rebrovach  — Nadvirnianskyi District, Vorokhta village (48.298046 N, 24.605371 E); Pistyn  — Kosivskyi District, Pistyn village (48.356373 N, 25.019105 E); Tatariv — Nadvirnianskyi District, Tatariv village (48.343931 N, 24.572832 E); Vorokhta — Nadvirnianskyi District, Vorokhta village (48.284073 N, 24.559030 E). Lviv Region (LWI): Bibrka — Peremyshlianskyi District, Bibrka town (49.639129 N, 24.293517 E); Dit- kivtsi  — Brodivskyi District, Ditkivtsi village (50.082033 N, 25.187976 E); Ivano-Frankove  — Yavorivskyi District, Ivano-Frankove village (49.921699 N, 23.722952 E); Pasiky-Zubrytski  — Pustomytivskyi District, Pasiky-Zubrytski village (49.7728439 N, 24.0712486 E); Roztochia Biosphere Reserve — Yavorivskyi District, Ivano-Frankove village (49.947253 N, 23.655512 E); Zubra — Pustomytivskyi District, Zubra village (49.771909 N, 24.051012 E); Lviv — Lviv City (49.839337 N, 24.030204 E); Holosko — Lviv City (49.870974 N, 24.018913 E); Kryvchytsi  — Lviv City (49.849492 N, 24.084754 E); Pohulanka, Lviv City (49.8245 N, 24.0650 E); Ri- asne — Lviv City (49.869202 N, 23.953784 E); Sykhiv — Lviv City (49.790710 N, 24.064629 E); Zamarstyniv — Lviv City (49.8634595 N, 24.0193791 E). Transcarpathian Region (ZAK): mountain range Chornohora (48.160040 N, 24.500105 E), Mala Uhol- ka — Tiachivskyi District, Mala Uholka village (48.2637 N, 23.6232 E); mountain Yavirnyk — Volovetskyi Dis- trict, Skotarske village, south-western slope, the outskirts of the mountain, beech derivative forest, 60 years old, litter (48.759725 N, 23.26430 E); Stinka mountain range — Velykobereznianskyi District, Kniahynia village, 961 m, mountain meadow, (48.999267 N, 22.506984 Е), mountain range Yavirnyk  — Velykobereznianskyi District, Mochar village, 1010 m, mountain meadow (48.911741 N, 22.556342 Е), Polianskyi — Rakhiv District, Dilove village, Maramaros Mts, Mt Polianskyi, 850 m a. s. l., fi r-beech forest 160–180 years old (47.936723 N, 24.132009 E); Kuzii — Rakhiv District, Luh village, Maramaros Mts, Kuzii site, 380 a. s. l., derivate beech forest mixed with fi r and spruce 60–80 years old (47.934847 N, 24.125106 E). Chernivtsi Region (СER): Chernivtsi City (48.317560 N, 25.917437 E). Results Genus Atheta Th omson, 1858 D i a g n o s i s . Body ovoid or subparallel, rather loosely articulated, length 2.5–4.8 mm in most species; color: dark brown to black, integument usually with meshed microscu- lpture; pronotum with pubescence on disc forming distinct pattern, usually directed lat- eroposteriad from midline, but with specifi c patterns in diff erent subgenera; hypomera completely visible in lateral view; mesocoxae narrowly separated; process of mesoven- trite slender; tarsal formula 4–5–5; athetine bridge located in dorso-basal part of median lobe of aedeagus; spermatheca with club-shaped capsule and long stem that is narrowly looped posteriorly in most species (there are many spermathecal forms presented in Atheta). Th is genus is treated here as Atheta sensu lato, and embraces a wide range of spe- cies with diverse morphology forming the Atheta complex. It includes several more or less clearly defi ned subgenera that are not suffi ciently delimited to justify elevating them to generic status, and such an act would bring only more confusion. Th e subgenera of Atheta are listed below in alphabetical order because the relationships between them are not clear. For species identifi cation, dissection and examination of genital structures is necessary (Klimaszeski et al., 2018). Subgenus Anopleta Mulsant & Rey, 1874 Atheta corvina (Th omson, 1856) M a t e r i a l . 1 ex. IFR.: Pistyn [Pistyń], date not specifi ed, 1 ex. (SMNH). 94 S. Glotov, K. Hushtan, H. Hushtan, N. Koval, V. Diedus D i s t r i b u t i o n . Europe, Siberia (Schülke & Smetana, 2015); Ukrainian Carpathians: Ivano-Frankivsk Region, between Pistyn (Glotov & Hushtan, 2020). “Poland”, without indication of region (Łomnicki, 1913) B i o n o m i c s . Th e species occurs in mixed and deciduous forests, in the mountains, where they live in forest litter, decaying organic residues, and in moss (Nikitskij et al., 1996; Semenov, 2007). Adults occur in V–VII (Ganglbauer, 1895). Atheta kochi Roubal, 1937 M a t e r i a l . 7 ex. ZAK: Mala Uholka, 15.08.2017, 5 ex., same locality, 05.09.2017, 2 ex., leg. V. Chumak (KCHM). D i s t r i b u t i o n . Central and South Europe (Schülke & Smetana, 2015); Ukraine (fi rst record). B i o n o m i c s . Biology has not been studied. Th e entire material is collected with the help of window traps. Subgenus Atheta Thomson, 1858 Atheta allocera Eppelsheim, 1893 M a t e r i a l . Literature data only. D i s t r i b u t i o n . Europe, East Siberia, Mongolia, Japan (Schülke & Smetana, 2015); in the Ukrainian Carpathians known from Ivano-Frankivsk Region, mountain Dantsyzh (Lokay, 1912). B i o n o m i c s . Th e species occurs in lowlands and foothills along rivers and reservoirs, in forest litter and in remains of plants. Atheta aquatica (Th omson, 1852) M a t e r i a l . 27 ex. ZAK: Mala Uholka, 15.07.2017, 14 ex., same locality, 15.08.2017, 11 ex., same locality, 15.09.2017, 2 ex., leg. V. Chumak (KCHM). D i s t r i b u t i o n . Europe, Asia Minor, Siberia, Central Asia (Schülke & Smetana, 2015); in the Ukrainian Carpathians known from Transcarpathian Region, mountain Hoverla (Fleischer et al., 1924), Kuzii tract (Roubal, 1930), “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs in lowlands and foothills along rivers and reservoirs, in swampy areas and in places of accumulation of moisture, in sediments and in remains of plants (Roubal, 1930; Glotov, 2021). Atheta autumnalis (Erichson, 1839) M a t e r i a l . 1 ex. CER: Chernivtsi [Bucowina. Czernowitz], data not specifi ed, 1 ex. (ZMTSNU). D i s t r i b u t i o n . Europe (Schülke & Smetana, 2015); in the Ukrainian Carpathians known from Ivano-Frankivsk Region, between Vorokhta and Tatarov (Lokay, 1912). B i o n o m i c s . Th e remains of plants species occurs in lowlands and foothills, in forest litter and in remains of plants. Atheta boletophila Th omson, 1856 M a t e r i a l . Literature data only. D i s t r i b u t i o n . Europe, Far East (Schülke & Smetana, 2015); in the Ukrainian Car- pathians known from Transcarpathian Region, Rakhiv (Kuthy, 1896). B i o n o m i c s . Th e species occurs in lowlands and in foothills where they are wide- spread in Inonotus obliquus (Pers.) Pil., sometimes found on other fungi (Bogdanov, 1985; Semenov, 2007). 95 Th e Genus Atheta (Coleoptera, Staphylinidae, Aleocharinae) in the Ukrainian Carpathians Fig. 1. Habitus: a — Atheta coriaria (Kraatz, 1856); b — A. divisa (Märkel, 1844); c — A. graminicola (Grave- nhorst, 1806); d  — A. oblita (Erichson, 1839); e  — A. ravilla (Erichson, 1839); f  — A. macrocera (Th omson, 1856); g — A. occulta (Erichson, 1837); h — A. longicornis (Gravenhorst, 1802); i — A. melanaria (Manner- heim, 1830); j — A. zosterae (Th omson, 1856); k — A. atramentaria (Gyllenhal, 1810); l — A. marcida (Erich- son, 1837); m — A. subtilis (Scriba, 1866); n — A. amicula (Stephens, 1832); o — A. laticollis Stephens, 1832; p — A. liturata (Stephens, 1832); q — A. nigritula (Gravenhorst, 1802). 96 S. Glotov, K. Hushtan, H. Hushtan, N. Koval, V. Diedus Atheta brunneipennis (Th omson, 1852) M a t e r i a l . Literature data only. D i s t r i b u t i o n . Europe, Eastern Siberia, Far East (Schülke & Smetana, 2015); “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs in lowlands and foothills in forest litter and in remains of plants (Glotov, 2021). Atheta castanoptera Mannerheim, 1830 M a t e r i a l . 13 ex. IFR: Vorokhta [Worochta. Cp. or. ], 24.6. [19]11, 1 ex., leg. Dr. Lokay, coll. Grolle, same locality but, 1 ex., same locality but, leg. Stoeekel, 4 ex. LWI: Bibrka [Bobr], 9.7. [year not specifi ed], 1 ex.; Ditkivtsi [D], 26.8. [year not specifi ed], 1 ex.; Lviv [ok. Lwowa], date not specifi ed, 5 ex., coll. Grolle. (SMNH); Roztochia Biosphere Reserve, 24.08.1988, 4 ex., leg. A. Petrenko (SIZK). D i s t r i b u t i o n . Palaearctic (Schülke & Smetana, 2015); in the Ukrainian Carpathians known from Ivano-Frankivsk Region, Bystrets (Miller, 1868), Verkhovyna (Łomnicki, 1868) and Transcarpathian Region, mountain range Chornohora (Nowicki, 1873; Weise, 1876). Without specifying the territory is given in the catalogs of “Galicia and Poland” (Łomnicki, 1884, 1913). B i o n o m i c s . Th e species occurs in lowlands and foothills where they live along rivers and in remains of plants (Glotov, 2021). Atheta contristata (Kraatz, 1856) M a t e r i a l . Literature data only. D i s t r i b u t i o n . West and South Europe (Schülke & Smetana, 2015); in the Ukrainian Carpathians known from Transcarpathian Region, mountain Blyznytsia (Fleischer et al., 1924). B i o n o m i c s . Th e species occurs in lowlands and foothills where they live in forest litter and in remains of plants (Fleischer et al., 1924). Atheta coriaria (Kraatz, 1856) (fi g. 1, a) M a t e r i a l . 1 ex. LWI: Lviv, Riasne [Rzęsna], 14.4.1918, 1 ex., det. Dr. Lokay. (SMNH). D i s t r i b u t i o n . Cosmopolitan (Schülke & Smetana, 2015); fi rst record for the Ukrainian Carpathians. B i o n o m i c s . Th e species occurs in lowlands and in foothills where they live in forest litter (Semenov, 2007). Atheta crassicornis (Fabricius, 1792) M a t e r i a l . 54 ex. CER: Chernivtsi [Bucowina. Czernowits], data not specifi ed, 3 ex. (ZMTSNU). IFR: Vorokhta [Worochta], date not specifi ed, 6 ex.; same locality but, date not specifi ed, 1 ex., coll. Grolle. LWI: Ivano-Frankove [Janów], 10.4. [year not specifi ed], 1 ex., leg. A. Stoeckel, same locality but, date not specifi ed, 2 ex.; Roztochia Biosphere Reserve, 24.08.1988, 1 ex., leg. A. Petrenko (SIZK). Lviv [Lwów], 18.3. [1]925, 1 ex.; same locality but, 28.3. [1]925, 1 ex.; same locality but, 24.4. [1]925, 1 ex.; same locality but, date not specifi ed, 3 ex.; same locality but, 29.3. [year not specifi ed], 2 ex., coll. Grolle; same locality but, [ok. Lwowa], date not specifi ed, 1 ex., coll. Grolle; same locality but, Holosko [H], 13.9. [year not specifi ed], 1 ex.; same locality but, Pohulanka [Pohulanka], 14.4. 1917, 1 ex.; same locality but, 5.5. [19]16, 1 ex., leg. Grolle, coll. Grolle; locality not specifi ed, 25.7. [year not specifi ed], 1 ex. (SMNH). ZAK: Mala Uholka, 16.10.1984, 4 ex., leg. V. Chumak; same locality but, 13.08.1990, 3 ex., leg. V. Chumak (SIZK); mountain Yavirnyk, 21.05. — 02.07.2009, 1 ex., leg. Yu. Kanarskyi; Polianskyi, 14.05–19.06.2009, 1 ex., leg. Yu. Kanarskyi; Kuzii, 15.04.  — 15.05.2009, 1 ex., leg. Yu. Kanarskyi; same locality, 15.05–20.06.2009, 1 ex., leg. Yu. Kanarskyi (SMNH); Stinka mountain range, eco- tone, 21.06.2018, 7 ex.; same locality but, ecotone, 23.08.2018, 3 ex., same locality but, forest, 18.05.2019, 2 ex., mountain range Yavirnyk, mountain meadow, 27.06.2018, 9 ex., same locality but, ecotone, 29.08.2018, 2 ex. leg. N. Koval (KKOV). 97 Th e Genus Atheta (Coleoptera, Staphylinidae, Aleocharinae) in the Ukrainian Carpathians D i s t r i b u t i o n . Palaearctic (excluding Far East) (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from the Ukrainian Carpathian Mountains (Mateleshko, 2008), Transcarpathian Region, Kuzii locality, Mountain Polianskyi (Glotov et al., 2020) and Lviv Region, Pohulanka, Klepariv, Holosko (Łomnicki, 1890; 1891). “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs in lowlands, in foothills as well as high in the mountains in deciduous and mixed forests, in meadows and in artifi cial plantations, in forest litter, along rivers and reservoirs, under wet leaves and in plant remains, as well as in many agaric and pore fungi (Łomnicki, 1891; Glotov, 2021). Atheta diversa diversa (Sharp, 1869) M a t e r i a l . 1 ex. IFR: Vorokhta [Worochta. Cp. or. ], 24.6. [19]11, 1 ex., leg. Dr. Lokay, coll. Grolle. (SMNH). D i s t r i b u t i o n . Europe (Schülke & Smetana, 2015); in the Ukrainian Carpathians known from Ivano-Frankivsk Region, between Vorokhta and Tatarov 9.08. (Lokay, 1912). B i o n o m i c s . Th e species occurs in lowlands and in foothills of coniferous and de- ciduous forests, where they live in forest litter, mosses and fungi, in particular it is noted on Ganoderma lipsiense (Batsch) G. F. Atk. (= applanatum (Pers.) Pat.) (Semenov, 2007). Atheta divisa (Märkel, 1844) (fi g. 1, b) M a t e r i a l . 7 ex. LWI: Ivano-Frankove [Janów], 22.7. [1]917, 2 ex.; Lviv [ok. Lwowa], 22.5. [year not specifi ed], 5 ex. (SMNH). D i s t r i b u t i o n . Europe (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Chernivtsi Region, Chernivtsi, Convent of the Nativity of the Virgin “Horecea”, Zhuchka (Marcu, 1936). “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs in wet places in remains of plant and animal origin, on the fl owing sap of trees. At dusk, they fl y to light. Atheta hypnorum (Kiesenwetter, 1850) M a t e r i a l . Literature data only. D i s t r i b u t i o n . Europe, Asia Minor, West Siberia, Mongolia (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Ivano-Frankivsk Region, mountain Dantsyzh (Lokay, 1912) and Transcarpathian Region (Roubal, 1930). “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs in lowlands and highlands where they live in the forest litter, on the banks of rivers and reservoirs, in mosses and under the bark of trees as well as in rotten wood (Roubal, 1930). Atheta fungicola Th omson, 1852 M a t e r i a l . Literature data only. D i s t r i b u t i o n . Europe, Asia Minor (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Ivano-Frankivsk Region, Bystrets (Miller, 1868), Verkhovyna (Łomnicki, 1868) and Transcarpathian Region, mountain range Chornohora (Nowicki, 1873). Without specifying the territory is given in the catalog of Galicia (Łomnicki, 1884). B i o n o m i c s . Th e species occurs in lowlands and in foothills of coniferous and deciduous forests, where they live in the forest litter, in fungi, in particular it is noted in fungi of the genus Russula (Semenov, 2007). 98 S. Glotov, K. Hushtan, H. Hushtan, N. Koval, V. Diedus Atheta graminicola (Gravenhorst, 1806) (fi g. 1, c) M a t e r i a l . 10 ex. LWI: Ivano-Frankove [Janów], date not specifi ed, 4 ex.; Lviv [L], [132], date not speci- fi ed, 1 ex.; same locality but, 6.9. [year not specifi ed], 1 ex.; same locality but, Sykhiv [Sichów], date not specifi ed, 1 ex.; Zubra [Zubra], 7.5. [year not specifi ed], 1 ex. (SMNH). ZAK: Mala Uholka, 13.08.1990, 2 ex., leg. V. Chu- mak (SIZK). D i s t r i b u t i o n . Palaearctic (excluding North Africa) (Schülke & Smetana, 2015). Without specifying the territory is given in the catalogs of Galicia and Poland (Łomnicki, 1884, 1913); fi rst record for Ukrainian Carpathians. B i o n o m i c s . Th e species occurs in lowlands and foothills in deciduous and mixed forests, in swamps, along rivers and reservoirs, where they live in cover and coastal sediments, in mosses, in sedge and cereal mounds, in dry grass; in warm evenings, they oft en gather in mowing around vegetation. At dusk, they fl y to light (Semenov, 2007; Glotov, 2021). Atheta euryptera (Stephens, 1832) M a t e r i a l . 1 ex. LWI: Lviv [Lw], 5.5. [year not specifi ed], 1 ex. (SMNH). D i s t r i b u t i o n . Palaearctic (excluding North Africa) (Schülke & Smetana, 2015); in the Ukrainian Carpathians known from Ivano-Frankivsk Region, Bystrets (Miller, 1868), Verkhovyna (Łomnicki, 1868), Transcarpathian Region, mountain range Chornohora (Nowicki, 1873) and Lviv Region, Lviv, Vulka (Łomnicki, 1890). Without specifying the territory is given in the catalogs of Galicia and Poland (Łomnicki, 1884, 1913). B i o n o m i c s . Th e species occurs in lowlands and foothills in deciduous and mixed forests, where they live in forest litter, on fresh and fermented sap of birches, oaks, elms, willows and other deciduous trees, sometimes occur in rotten fungi and rotten plant remains (Łomnicki, 1890; Semenov, 2007). Atheta incognita (Sharp, 1869) M a t e r i a l . Literature data only. D i s t r i b u t i o n . Europe, Asia Minor (Schülke & Smetana, 2015); in the Ukrainian Carpathians known from Transcarpathian Region, Yesenia 3 km SW, Diana tract (Roubal, 1930). B i o n o m i c s . Th e species occurs in foothills and highlands in deciduous and mixed forests, where they live in forest litter, under bark and rotten wood and in fungi (Roubal, 1930). Atheta laevicauda J. Sahlberg, 1876 M a t e r i a l . Literature data only. D i s t r i b u t i o n . Europe (Schülke & Smetana, 2015); in the Ukrainian Carpathians known from Transcarpathian Region, mountain range Chornohora, mountain Hoverla (Rybiński, 1903) and Ivano-Frankivsk Region, mountain Dantsyzh (Lokay, 1912). “Po- land”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs in foothills and highlands in deciduous and mixed forests, where they live in fungi (Roubal, 1930). Atheta oblita (Erichson, 1839) (fi g. 1, d) M a t e r i a l . 5 ex. Ukraine: CER: Chernivtsi [Bucowina. Czernowitz], data not specifi ed, 4 ex. (ZMTSNU); LWI: Lviv, Holosko [H], 21.10. [year not specifi ed], 1 ex. (SMNH). 99 Th e Genus Atheta (Coleoptera, Staphylinidae, Aleocharinae) in the Ukrainian Carpathians D i s t r i b u t i o n . Europe, Caucasus, Asia Minor, Iran, Syria, Mongolia (Schülke & Smetana, 2015); in the Ukrainian Carpathians known from the Ukrainian Carpathian Mountains (Mateleshko, 2008), Chernivtsi Region, Chernivtsi, mountain Tsetsyno (Marcu, 1936), Lviv Region, Holosko (Łomnicki, 1890, 1891) and Transcarpathian Region, Uzhhorod (Kuthy, 1896). “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs in foothills and highlands in deciduous and mixed forests, where they live in forest litter, in rotten wood and in remains of plants, as well as in the nests of small mammals, oft en found in fungi: Laetiporus sulphureus (Bull.) Murr., Inonotus dryadeus (Pers.) Murr. та Polyporus squamosus Huds. (Roubal, 1930; Semenov, 2007; Glotov, 2021). Atheta pilicornis (Th omson, 1852) M a t e r i a l . Literature data only. D i s t r i b u t i o n . Europe, East Siberia (Schülke & Smetana, 2015); in the Ukrainian Carpathians known from Lviv Region, Lviv, Pohulanka (Łomnicki, 1890, 1891) and Trans- carpathian Region, Kozmeschik tract (Roubal, 1930). “Poland”, without indication of re- gion (Łomnicki, 1913). B i o n o m i c s . Th e species occurs in foothills and mountain forests, where they live in fungi (Roubal, 1930). Atheta ravilla (Erichson, 1839) (fi g. 1, e) M a t e r i a l . 12 ex. CER: Chernivtsi [Bucowina. Czernowitz], data not specifi ed, 4 ex. (ZMTSNU). IFR: Vorokhta [Worochta], date not specifi ed, 2 ex.; LWI: Lviv [Lwów], 7.04.1925, 1 ex.; same locality but, [L], 27.3. [year not specifi ed], 1 ex.; Zubra [Zubra], date not specifi ed, 1 ex.; Pasiky-Zubritski [Pasieki], 6.05. 1917, 1 ex.; same locality but, 12.05. [1]917, 2 ex. (SMNH). D i s t r i b u t i o n . Europe, Caucasus, North Africa (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Lviv Region, Lviv (Łomnicki, 1890, 1891), Lviv and Chernivtsi Region, Chernivtsi, mountain Tsetsyno (Marcu, 1936). “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs in foothills and mountain forests, where they live mainly in the burrows and nests of mammals, in remains of plants, sometimes found on the fl owing sap of trees and in fungi: Phallus impudicus Pers., Laetiporus sulphureus (Bull.) Murr. (Łomnicki, 1890; Semenov, 2007). Atheta vaga (Heer, 1839) M a t e r i a l . 5 ex. ZAK: Stinka mountain range, 18.05.2019, 1 екз.; same locality but, 30.06.2019, 1 ex., leg.  N.  Koval; Yavirnyk mountain range, 27.05.2019, 1 ex.; same locality but, 19.06.2019, 2 ex., leg.  N.  Koval (KKOV). D i s t r i b u t i o n . Europe, Caucasus, North Africa, Siberia, Mongolia (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Carpathian Mountains (Mateleshko, 2008) and Transcarpathian Region, mountain range Chornohora; mountain Sheshul and Uzhhorod (Roubal, 1930). “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs from lowlands to highlands in deciduous and mixed forests, in meadows, along rivers and reservoirs in coastal sediments, in remains of plant and animal origin, in mammals’ burrows and in birds’ nests, on fermented birch sap, in rotten fungi (Roubal, 1930; Semenov, 2007). 100 S. Glotov, K. Hushtan, H. Hushtan, N. Koval, V. Diedus Atheta xanthopus (Th omson, 1856) M a t e r i a l . 5 ex. LWI: Ivano-Frankove [Janów], date not specifi ed, 1 ex.; same locality but, Sykhiv [Si- chów], date not specifi ed, 2 ex.; Zubra [Zubra], 1 ex., same locality but, 24.4. [year not specifi ed], 1 ex. (SMNH). D i s t r i b u t i o n . Europe, North Africa (Schülke & Smetana, 2015). “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs in lowlands and foothills in deciduous and mixed forests, where they live in forest litter and in remains of plants (Semenov, 2007). Subgenus Badura Mulsant & Rey, 1873 Atheta cauta (Erichson, 1837) M a t e r i a l . Literature data only. D i s t r i b u t i o n . Europe, Asia Minor, Siberia, Far East (Schülke & Smetana, 2015); “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs in lowlands and foothills in deciduous and mixed forests, where they live in forest litter, in remains of plants as well as in manure (Semenov, 2007). Atheta macrocera (Th omson, 1856) (fi g. 1, f) M a t e r i a l . Literature data only. D i s t r i b u t i o n . Europe, Caucasus, Kazakhstan (Schülke & Smetana, 2015); іn the Ukrainian Carpathians known from Ivano-Frankivsk Region, between Vorokhta and Tatarov (Lokay, 1912). B i o n o m i c s . Th e species occurs in lowlands and foothills in deciduous and mixed forests, where they live in manure, plant and animal remains, on tree sap, rotten fungi, es- pecially in Laetiporus sulphureus (Bull.) Murr. (Semenov, 2007; Glotov, 2021). Atheta spatula (Fauvel, 1875) M a t e r i a l . Literature data only. D i s t r i b u t i o n . Central and South Europe (Schülke & Smetana, 2015); in the Ukrai- nian Carpathians known from Transcarpathian Region, Lazeshchyna 8 km SSE, Kozmes- chik tract (Roubal, 1930). B i o n o m i c s . Th e species occurs mainly in foothills and high in the mountains, in deciduous and mixed forests, in plains where they live in intact forests in forest litter, but more oft en it occurs in fungi (Roubal, 1930). Subgenus Bessobia Thomson, 1858 Atheta occulta (Erichson, 1837) (fi g. 1, g) M a t e r i a l . 30 ex. CER: Chernivtsi [Bucowina. Czernowitz], data not specifi ed, 4 ex. (ZMTSNU). LWI: Lviv [Lwiw], date not specifi ed, 4 ex.; same locality but, 23.4[19]16, 1 ex.; same locality but, 28.4[1]925, 1 ex.; same locality but, 2.4. [year not specifi ed], 1 ex.; same locality but, [Lm], 8.4. [year not specifi ed], 1 ex.; same locality but, [Lm], 6.4. [year not specifi ed], 1 ex.; same locality but, Sykhiv [Sichów], date not specifi ed, 1 ex. (SMNH). ZAK: Stinka mountain range, forest, 23.08.2018, 2 ex.; same locality but, ecoton, 18.05.2019, 1 ex., same locality but, ecotone, 04.06.2019, 2 ex., same locality but, mountain meadow, 19.07.2019, 2 ex., mountain range Yavirnyk, ecotone, 05.06.2018, 1 ex., same locality but, ecotone, 27.06.2018, 2 ex., same locality but, moun- tain meadow, 12.05.2019, 2 ex., same locality but, mountain meadow, 08.07.2019, 4 ex., leg. Koval N. (KKOV). D i s t r i b u t i o n . Europe, Caucasus, Asia Minor, North Africa, Siberia, Far East (Schül- ke & Smetana, 2015); іn the Ukrainian Carpathians known from Chernivtsi Region, Cher- nivtsi, in spring under leaves (Hormuzaki, 1891) and Lviv Region, Lviv, Lychakiv District 101 Th e Genus Atheta (Coleoptera, Staphylinidae, Aleocharinae) in the Ukrainian Carpathians (Łyczakowskie przedmieście) (Łomnicki, 1890). Without specifying the territory is given in the catalogs of Galicia and Poland (Łomnicki, 1884, 1913). B i o n o m i c s . Th e species occurs in lowlands and foothills in deciduous and mixed forests on forest glades, meadows, in forest litter and on animal cadavers (Semenov, 2007). Atheta monticola (Th omson, 1852) M a t e r i a l . 1 ex. IFR: Vorokhta [Worochta], data not specifi ed, 1 ex. (SMNH). D i s t r i b u t i o n . Europe (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Chernivtsi Region, Chernivtsi, mountain Tsetsyno (Marcu, 1936). “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs in lowlands and foothills in deciduous and mixed forests, where they live in remains of animal origin, on animal cadavers (Semenov, 2007). Subgenus Chaetida Mulsant & Rey, 1873 Atheta longicornis (Gravenhorst, 1802) (fi g. 1, h) M a t e r i a l . 6 ex. LWI: Lviv [Lwów], 22.9. [1]917, 1 ex.; same locality but, 1.5. [1]917, 1 ex., same locality but, 23.4. [year not specifi ed], 1 ex.; same locality but, Holosko [H], 23.7. [year not specifi ed], 1 ex. (SMNH). ZAK: Mala Uholka, 05.05.2017, 2 ex., leg. V. Chumak (KCHM). D i s t r i b u t i o n . Palaearctic, Oriental Regions, North America (Schülke & Smeta- na, 2015). In the Ukrainian Carpathians known from Lviv Region, Lviv, Znesinnia, in the second half of July (Łomnicki, 1890) and Transcarpathian Region, mountain range Chornohora (Weise, 1876). Without specifying the territory is given in the catalogs of Gali- cia and Poland (Łomnicki, 1884, 1913). B i o n o m i c s . Th e species occurs mainly in foothills and high in the mountains, in deciduous and mixed forests where they live in forest litter, in manure, in remains of plant and animal origin, in rotten fungi, as well as on the fermented sap of trees (Roubal, 1930; Semenov, 2007). Subgenus Coprothassa Thomson, 1859 Atheta melanaria (Mannerheim, 1830) (fi g. 1, i) M a t e r i a l . 4 ex. LWI: Lviv [Lwów], 22.9. [1]917, 1 ex.; same locality but, date not specifi ed, 2 ex. (SMNH). ZAK: Mala Uholka, 13.08.1990, 1 ex., leg. V. Chumak (SIZK). D i s t r i b u t i o n . Palaearctic, Oriental Regions (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Chernivtsi Region, Chernivtsi (Hormuzaki, 1888), Lviv Region, the vicinity of Lviv (Łomnicki, 1886, 1890) and Transcarpathian Region, Bere- hovo (Kuthy, 1896). Without specifying the territory is given in the catalogs of Galicia and Poland (Łomnicki, 1884, 1913). B i o n o m i c s . Th e species occurs in lowlands and foothills in the manure of cattle and wild mammals (Roubal, 1930; Semenov, 2007). Subgenus Datomicra Mulsant & Rey, 1874 Atheta canescens (Sharp, 1869) M a t e r i a l . 2 ex. LWI: Ivano-Frankove [Janów], 9. [1]899, 1 ex.; same locality but, date not specifi ed, 1 ex. (SMNH). D i s t r i b u t i o n . Europe, Siberia, Mongolia (Schülke & Smetana, 2015). B i o n o m i c s . Th e species occurs in lowlands and in foothills, in deciduous and mixed forests, in meadows, along rivers and reservoirs, where they live in forest litter, in sediments, in manure, in remains of plant and animal origin, on animal cadavers (Semenov, 2007). 102 S. Glotov, K. Hushtan, H. Hushtan, N. Koval, V. Diedus Atheta celata (Erichson, 1837) M a t e r i a l . 5 ex. IFR: Vorokhta [Worochta], date not specifi ed, 1 ex.; same locality but, 20.6. [19]11, 1 ex., leg. Dr. Lokay; LWI: Lviv [ok. Lwowa], 22.5. [year not specifi ed], 1 ex.; same locality but, Pohulanka, 4.5. [year not specifi ed], 2 ex. (SMNH). D i s t r i b u t i o n . Holarctic (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Chernivtsi Region, Chernivtsi (Hormuzaki, 1888). Without specifying the territory is given in the catalogs of Galicia and Poland (Łomnicki, 1884, 1913). B i o n o m i c s . Th e species occurs in lowlands and foothills, in deciduous and mixed forests, in meadows, along rivers and reservoirs, where they live in forest litter, rotten wood and hay, in rotten fungi, in coastal sediments, sometimes on animal cadavers and in manure (Semenov, 2007; Glotov, 2021). Atheta sordidula (Erichson, 1837) M a t e r i a l . 5 ex. LWI: Ivano-Frankove [Janów], date not specifi ed, 2 ex.; same locality but, 4.9. [1]917, 1 ex.; same locality but, 9.9. [1]917, 1 ex.; Lviv [Lw], 16.7. [year not specifi ed], 1 ex. (SMNH). D i s t r i b u t i o n . Europe, Caucasus, Asia Minor, Kazakhstan, Siberia, Far East, Oriental Regions (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Chernivtsi Region, Chernivtsi (Hormuzaki, 1888, 1889) and Transcarpathian Region, Uzhhorod (Roubal, 1930). Without specifying the territory is given in the catalogs of Galicia and Poland (Łomnicki, 1884, 1913). B i o n o m i c s . Th e species occurs both in lowlands and high in the mountains, in deciduous and mixed forests, meadows and mountain meadow, along rivers and reservoirs, where they live in forest litter, in rotten wood, in rotten hay and grass, on the fl owing sap of trees, in mature and in rotten fungi, in coastal deposits, sometimes on animal cadavers and in manure, and burrows and nests of mammals (Roubal, 1930; Semenov, 2007). Atheta zosterae (Th omson, 1856) (fi g. 1, j) M a t e r i a l . 7 ex. IFR: Pistyn [Pištyn], date not specifi ed, 1 ex.; LWI: Ivano-Frankove [Janów], date not specifi ed, 1 ex.; Lviv [ok. Lwowa], 22.5. [year not specifi ed], 3 ex.; same locality, [Lwіw], date not specifi ed, 2 ex. (SMNH). D i s t r i b u t i o n . Europe, North Africa, Asia Minor, Kazakhstan (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Transcarpathian Region, Uzhhorod (Roubal, 1930). Without specifying the territory is given in the catalogs of Galicia and Poland (Łomnicki, 1884, 1913). B i o n o m i c s . Th e species occurs in forests and on open steppe and meadow areas, at foothills and high in the mountains, where they live in forest litter, in decaying plant residues, in fungi, in holes of rodents, in nests of birds, in ant hills, and on fl owing sap of trees. Adults occur in V–IX (Bogdanov 1985; Semenov, 2007; Glotov, 2021). Subgenus Dimetrota Mulsant & Rey, 1873 Atheta atramentaria (Gyllenhal, 1810) (fi g. 1, k) M a t e r i a l . 2 ex. IFR: Vorokhta [Worochta], date not specifi ed, 2 ex. (SMNH). D i s t r i b u t i o n . Holarctic, Afrotropical and Oriental Regions (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Transcarpathian Region, mountain Hoverla (Rybiński, 1903). “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs both in lowlands and high in the mountains, almost exclusively in open, unforested areas, in meadows, forest glades and meadows, or on border of forests and meadows in the manure of cattle and wild animals (Semenov, 2007). 103 Th e Genus Atheta (Coleoptera, Staphylinidae, Aleocharinae) in the Ukrainian Carpathians Atheta cadaverina (Brisout de Barneville, 1860) M a t e r i a l . 3 ex. IFR: Vorokhta [Worochta], date not specifi ed, 1 ex.; LWI: Lviv [Lwów], 9.4. [19]24, 1 ex.; same locality but, [L], 22.3. [year not specifi ed], 1 ex. (SMNH). D i s t r i b u t i o n . Europe (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Transcarpathian Region, mountain Blyznytsia (Fleischer et al., 1924), Uzhhorod (Roubal, 1930). “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . A rare species, which occurs both in lowlands and high in the mountains in remains of plant and animal origin, on animal cadavers, in cattle manure and in underground nests of mammals (Roubal, 1930; Semenov, 2007). Atheta europaea Likovský, 1984 M a t e r i a l . 5 ex. LWI: Ivano-Frankove [Janów], date not specifi ed, 2 ex.; Lviv [Lwów], date not specifi ed, 1 ex.; same locality but, [ok. Lwowa], 11.4. [1]917, 1 ex.; Zubra [Zubra], date not specifi ed, 1 ex. (SMNH). D i s t r i b u t i o n . West and South Europe (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Transcarpathian Region, mountain Sheshul (Roubal, 1930). “Po- land”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs both in lowland and high in the mountains in decidu- ous and mixed forests, in meadows and mountain meadows, in fl oodplains, on forest glades in forest litter, in remains of plants, in coastal sediments, in manure of cattle and wild ani- mals as well as in fungi (Roubal, 1930; Semenov, 2007). Atheta intermedia (Th omson, 1852) M a t e r i a l . 4 ex. IFR: Vorokhta [Worochta], date not specifi ed, 1 ex.; same locality but, 5. [1]926, 1 ex.; same locality but, 10. [1]926, 1 ex.; LWI: Lviv [Lwów], 28.3. [1]925, 1 ex. (SMNH). D i s t r i b u t i o n . Europe, East Siberia, Far East (Schülke & Smetana, 2015). Without indication for the territory is given in the catalog of Poland (Łomnicki, 1913); Ukraine (fi rst record). B i o n o m i c s . Th e species occurs in lowlands and in foothills, in deciduous and mixed forests, in meadows, on the banks of rivers and reservoirs, where they live in forest litter, in sediments, in manure, in remains of plant and animal origin, on animal cadavers (Semenov, 2007). Atheta laevana (Mulsant & Rey, 1852) M a t e r i a l . 7 ex. IFR: mountain Rebrovach [Rebrowac Cp. or.], date not specifi ed, 1 ex., leg. Dr. Lokay. LWI: Ivano-Frankove [Janów], date not specifi ed, 5 ex.; Lviv, Pohulanka [Pohulanka], 6.4. [1]917, 1 ex. (SMNH). D i s t r i b u t i o n . Europe, North Africa, Asia Minor, Siberia (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Ivano-Frankivsk Region, mountain Rebrovach (Lokay, 1912) and Transcarpathian Region, mountain range Chornohora, mountain Sheshul (Roubal, 1930). Without specifying the territory is given in the catalogs of Galicia and Poland (Łomnicki, 1884, 1913). B i o n o m i c s . Th e species occurs mainly in mountains in deciduous and mixed forests, in meadows and mountain meadows, in forest litter, in coastal sediments, in hay and rotten grass as well as in manure (Roubal, 1930). Atheta marcida (Erichson, 1837) (fi g. 1, l) M a t e r i a l . Literature data only. D i s t r i b u t i o n . Europe, North Africa (Schülke & Smetana, 2015); “Poland”, without indication of region (Łomnicki, 1913). 104 S. Glotov, K. Hushtan, H. Hushtan, N. Koval, V. Diedus B i o n o m i c s . Th e species occurs mainly in lowlands and in foothills in deciduous and mixed forests in forest litter (Glotov, 2020). Atheta nigripes (Th omson, 1856) M a t e r i a l . 1 ex. IFR: Kosiv [Kossiw], date not specifi ed, 1 ex. (SMNH). D i s t r i b u t i o n . Europe, Mongolia, North Korea (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Transcarpathian Region, mountain Hoverla (Rybiński, 1903). “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Mixed and deciduous forests. Th e species occurs mainly in lowlands and in foothills in deciduous and mixed forests in forest litter (Glotov, 2020). Atheta putrida (Kraatz, 1856) M a t e r i a l . 7 ex. CER: Chernivtsi [Bucovina. Czernowitz], data not specifi ed, 2 ex. (ZMTSNU). IFR: Vorokhta [Worochta], date not specifi ed, 1 ex.; LWI: Lviv [Lwiw], date not specifi ed, 1 ex.; same locality but, Pohulanka [Pohulanka], 6.4. [1]917, 1 ex.; same locality but, Sykhiv [Sichów], 1 ex., Pasiky-Zubrycki [Pasieki], 6.5. [1]917, 1 ex. (SMNH). D i s t r i b u t i o n . Central and South Europe, Asia Minor (Schülke, Smetana, 2015). In the Ukrainian Carpathians known from Lviv Region, Lviv, Pohulanka, in July (Łomnicki, 1890; 1891) and Transcarpathian Region, mountain Sheshul (Roubal, 1930). “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs mainly in lowlands and in foothills in deciduous and mixed forests in forest litter and in rotten wood, in coastal sediments, sometimes they occur with ants Lasius fuliginosus Latr. (Roubal, 1930). Atheta setigera (Sharp, 1869) M a t e r i a l . 3 ex. IFR: mountain Rebrovach [Rebrowac], date not specifi ed, 1 ex., leg. A. Stöeckel; moun- tain range Pozhyzhevska [Pożyżewska Cp. or.], 11.9. [19]08, 2 ex. (SMNH). D i s t r i b u t i o n . Europe (Schülke & Smetana, 2015); Ukraine (fi rst record). B i o n o m i c s . Th e species occurs mainly in foothills and high in the mountains in deciduous and mixed forests, in mountain meadows in the forest litter and in remains of plants. Atheta subrugosa (Märkel & Kiesenwetter, 1848) M a t e r i a l . Literature data only. D i s t r i b u t i o n . Central Europe, Mongolia (Schülke, Smetana, 2015); іn the Ukrainian Carpathians known from Transcarpathian Region, mountain Hoverla; Dubove 8 km E, mountain Apetska (Roubal, 1930). B i o n o m i c s . Th e species occurs mainly in mountains in deciduous and mixed forests, in mountain meadows in cattle manure, in forest litter, in rotten wood and in remains of plants (Roubal, 1930). Atheta subtilis (Scriba, 1866) (fi g. 1, m) M a t e r i a l . 2 ex. LWI: Lviv [Lwiw], date not specifi ed, 1 ex.; same locality but, Pohulanka [Pohulanka], 6.4. 1917, 1 ex. (SMNH). ZAK: mountain Yavirnyk, 03.05–21.05.2009, 1 ex., leg. Yu. Kanarskyi (SMNH). D i s t r i b u t i o n . Europe, Asia Minor, Mongolia (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Chernivtsi Region, Chernivtsi, mountain Tsetsyno, 105 Th e Genus Atheta (Coleoptera, Staphylinidae, Aleocharinae) in the Ukrainian Carpathians Bila (Marcu, 1936) and Transcarpathian Region, mountain Pоp Ivan (Fleischer et al., 1925). “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs mainly in lowlands and foothills in deciduous and mixed forests in forest litter and in rotten wood, in coastal sediments, in rotten fungi, on fermented birch sap, in straw and on animal cadavers (Semenov, 2007). Subgenus Microdota Mulsant & Rey, 1873 Atheta aegra (Heer, 1841) M a t e r i a l . Literature data only. D i s t r i b u t i o n . Europe, Asia Minor, Syria (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Transcarpathian Region, Kuzii tract (Roubal, 1930). B i o n o m i c s . Th e species occurs mainly in lowlands and foothills, in deciduous, mostly oak and beech forests, in forest litter and in remains of plants (Roubal, 1930). Atheta amicula (Stephens, 1832) (fi g. 1, n) M a t e r i a l . 15 ex. LWI: Ivano-Frankove [Janów], date not specifi ed, 1 ex.; Lviv, Sykhiv [Sichów], date not specifi ed, 1 ex.; Zubra [Zubra], date not specifi ed, 1 ex. (SMNH). ZAK: Mala Uholka, 20.06.2017, 12 ex., leg. V. Chumak (KCHM). D i s t r i b u t i o n . Palaearctic, Nearctic and Neotropical Regions (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Chernivtsi Region, Chernivtsi, Convent of the Nativity of the Virgin Horecea, under leaves in the forest (Hormuzaki, 1888). Without specifying the territory is given in the catalogs of Galicia and Poland (Łomnicki, 1884, 1913). B i o n o m i c s . Th e species occurs mainly in foothills and high in the mountains, in forest litter and in remains of plant and animal origin (Glotov, 2021). Atheta atomaria (Kraatz, 1856) M a t e r i a l . Literature data only. D i s t r i b u t i o n . Europe (Schülke & Smetana, 2015); “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs mainly in foothills and high in the mountains, in forest litter and in remains of plant and animal origin. Atheta foveicollis (Kraatz, 1856) M a t e r i a l . 1 ex. IFR: mountain Dantsyzh [Pod Dancerzem Cp. Or. ], 17.6. [19]11, 1 ex., coll. Grolle. (SMNH). D i s t r i b u t i o n . Central Europe, Asia Minor (Schülke & Smetana, 2015); without in- dication for the territory is given in the catalog of Poland (Łomnicki, 1913); Ukraine (fi rst record). B i o n o m i c s . Th e species occurs mainly in foothills and high in the mountains, in for- est litter and in remains of plants (Łomnicki, 1913). Atheta excelsa Bernhauer, 1911 M a t e r i a l . Literature data only. D i s t r i b u t i o n . Europe, East Siberia (Schülke & Smetana, 2015); іn the Ukrainian Carpathians known from Transcarpathian Region, mountain range Chornohora, mountain Hoverla (Roubal, 1930). 106 S. Glotov, K. Hushtan, H. Hushtan, N. Koval, V. Diedus B i o n o m i c s . Th e species occurs mainly high in the mountains in forests and mountain meadows, in cover and remains of plant and animal origin, on animal cadavers and in the manure of wild and domestic animals (Roubal, 1930; Semenov, 2007). Atheta luctuosa (Mulsant & Rey, 1853) M a t e r i a l . 3 ex. LWI: Ivano-Frankove [Janów], date not specifi ed, 1 ex.; Lviv [Lwów], 28.3. [1]925, 2 ex. (SMNH). D i s t r i b u t i o n . West and South Europe, North Africa, Asia Minor (Schülke & Smetana, 2015); Ukraine (fi rst record). B i o n o m i c s . Th e species occurs mainly in lowlands and in foothills, on the banks of rivers and reservoirs, in sediments and forest litter and in remains of plant and animal origin. Atheta palleola (Erichson, 1837) M a t e r i a l . Literature data only. D i s t r i b u t i o n . Europe, East Siberia, North Korea (Schülke & Smetana, 2015). Without specifying the territory is given in the catalogs of Galicia and Poland (Łomnicki, 1884, 1913). B i o n o m i c s . Biology has not been studied. Th e entire material is collected without specifying collection methods. Subgenus Mycetota Ádám, 1987 Atheta laticollis Stephens, 1832 (fi g. 1, o) M a t e r i a l . 10 ex. IFR: Tatariv [Tatarów Cp. or. ], 8.9. [19]08, 1 ex. LWI: Ivano-Frankove [Janów], 9.9. [1]925, 2 ex.; Lviv, Pohulanka [Pohulanka], 6.4. 1917, 2 ex.; Zubra [Zubra], data not specifi ed, 3 ex.; Pasiky- Zubrytski [Pasieki], 6.5. [1]917, 2 ex. (SMNH). D i s t r i b u t i o n . Palaearctic (excluding Far East) (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Chernivtsi Region, Chernivtsi (Hormuzaki, 1888) and Lviv Region, Bohdanivka (Łomnicki, 1890). Without specifying the territory is given in the catalogs of Galicia and Poland (Łomnicki, 1884, 1913). B i o n o m i c s . Biology has not been studied. Th e entire material is collected without specifying collection methods. Subgenus Oreostiba Ganglbauer, 1895 Atheta tibialis Heer, 1839 M a t e r i a l . 5 ex. IFR: mountain Dantsyzh [Pod Dancerzem Cp. Or. ], 17.6. [19]11, 1 ex., coll. Grolle. LWI: Lviv [L] [92], date not specifi ed, 1 ex., same locality but, Kryvchytsi [Kr], 3.7. [year not specifi ed], 1 ex.; same locality but, Zamarstyniv [Zm], 20.7. [year not specifi ed], 1 ex. ZAK: mountain range Chornohora [Czar- nohora], 3.7. [year not specifi ed], 1 ex. (SMNH). D i s t r i b u t i o n . Europe, North Africa, Asia Minor (Schülke & Smetana, 2015); in the Ukrainian known from the Carpathian Mountains. High alpine in damp places, close to the edges of the snow fi elds (Miller, 1868), Ivano-Frankivsk Region, Bystrets 6 km W, Gadzhyna locality (Łomnicki, 1868; 1886) and Transcarpathian Region, mountain Sheshul (Roubal, 1930), mountain range Chornohora (Nowicki, 1873; Weise, 1876), mountain range Chornohora, mountain Rebra-Hoverla (Łomnicki, 1868), mountains Rebra, Szpyci (Łomnicki, 1886). Without specifying the territory is given in the catalogs of Galicia and Poland (Łomnicki, 1884, 1913). 107 Th e Genus Atheta (Coleoptera, Staphylinidae, Aleocharinae) in the Ukrainian Carpathians B i o n o m i c s . Th e species occurs mainly in foothills and high in the mountains in for- ests and mountain meadows, in cover and plant remains, in mosses, on animal cadavers and in the manure of wild and domestic animals as well as in mammal burrows: Marmota bobak (Rybinski, 1902; Roubal, 1930). Subgenus Oxypodera Bernhauer, 1915 Atheta fi morum (Brisout de Barneville, 1860) M a t e r i a l . Literature data only. D i s t r i b u t i o n . Europe, Asia Minor (Schülke & Smetana, 2015); in the Ukrainian Carpathians known from Transcarpathian Region, Uzhhorod (Kuthy, 1896). B i o n o m i c s . Th e species occurs mainly in lowlands and in foothills, in forest litter and plant remains, in mosses, fungi and manure (Roubal, 1930). Subgenus Pachyatheta Munster, 1925 Atheta cribrata (Kraatz, 1856) M a t e r i a l . 1 ex. LWI: Lviv [Lw], 20.4. [year not specifi ed], 1 ex. (SMNH). D i s t r i b u t i o n . Europe (Schülke & Smetana, 2015); without indication for the territory is given in the catalog of Poland (Łomnicki, 1913); Ukraine (fi rst record). B i o n o m i c s . Th e species occurs mainly in lowlands and foothills, in forest litter and in mosses (Semenov, 2007). Atheta mortuorum Th omson, 1867 M a t e r i a l . 2 ex. IFR: Vorokhta [Worochta], date not specifi ed, 2 ex. (SMNH). D i s t r i b u t i o n . Europe, North Africa, Siberia, North Korea (Schülke & Smetana, 2015); Ukraine (fi rst record). B i o n o m i c s . Th e species occurs mainly in lowlands and foothills, in forest litter. Subgenus Rhagocneme Munster, 1923 Atheta subsinuata Erichson, 1839 M a t e r i a l . 1 ex. IFR: Vorokhta [Worochta], date not specifi ed, 1 ex. (SMNH). D i s t r i b u t i o n . Europe, North America (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Transcarpathian Region, Berehovo (Kuthy, 1896). “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs mainly in lowlands and foothills, in forest litter, in rotten wood and in mosses. Subgenus Tetropla Mulsant & Rey, 1873 Atheta liturata (Stephens, 1832) (fi g. 1, p) M a t e r i a l . 3 ex. CER: Chernivtsi [Bucowina. Czernowits], data not specifi ed, 3 ex. (ZMTSNU). D i s t r i b u t i o n . Europe (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Transcarpathian Region, mountain Hoverla (Kuthy, 1896) B i o n o m i c s . Th e species occurs mainly in foothills and high in the mountains in forests and mountain meadows, mainly in young mature and rotten fungi, mainly in Laetiporus sulphureus (Bull.) Murr., as well as in forest litter (Roubal, 1930; Glotov, 2020). 108 S. Glotov, K. Hushtan, H. Hushtan, N. Koval, V. Diedus Atheta nigritula (Gravenhorst, 1802) (fi g. 1, q) M a t e r i a l . 10 ex. CER: Chernivtsi [Bucowina. Czernowits], data not specifi ed, 4 ex. (ZMTSNU). IFR: Vorokhta [Worochta], date not specifi ed, 1 ex.; LWI: Lviv, Riasne [Rzęsna], 11.4. [1]925, 1 ex.; same local- ity but, Sykhiv [Schihow], date not specifi ed, 1 ex.; Zubra [Zubra], date not specifi ed, 1 ex.; Pasiky-Zubrytski [Pasieki], 12.5. [1]917, 1 ex.; locality not specifi ed, 11.4. [year not specifi ed], 1 ex. (SMNH). D i s t r i b u t i o n . Europe, North Africa, Asia Minor, East Siberia, North America (Schülke & Smetana, 2015). In the Ukrainian Carpathians known from Ivano-Frankivsk Region, Bystrets (Miller, 1868) and Transcarpathian Region, mountain range Chornohora (Nowicki, 1873). “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs in lowlands and in foothills, mainly in agaric mushrooms, as well as in forest litter (Semenov, 2007). Subgenus Thinobaena Thomson, 1859 Atheta vestita Gravenhorst, 1806 M a t e r i a l . 3 ex. IFR: Kosiv [Kossiw], date not specifi ed, 1 ex. LWI: Zubra [Zubra], 8.4. [year not speci- fi ed], 1 ex., same locality but, 6.4. [year not specifi ed], 1 ex. (SMNH). D i s t r i b u t i o n . Europe, North America (Schülke & Smetana, 2015). “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs in forests and on open steppe and meadow areas, at foothills and high in the mountains, on alkalized plots, along the banks of rivers and reservoirs, on nearshore area, where they live in decaying organic residues. Adults occur in V–VIII (Klimaszeski et al., 2007, 2011). Subgenus Traumoecia Mulsant & Rey, 1873 Atheta picipes (Th omson, 1856) M a t e r i a l . 1 ex. LWI: Lviv [Lwów], date not specifi ed, 1 ex. (SMNH). D i s t r i b u t i o n . Europe, East Siberia (Schülke & Smetana, 2015). Without indication for the territory is given in the catalog of Poland (Łomnicki, 1913); Ukraine (fi rst record). B i o n o m i c s . Th e species occurs in lowlands and in foothills in forest litter, in rotten wood and under the bark of trees aff ected by fungi, are found mainly in fungi: Fomes fomentarius (L.) Fr., Piptoporus betulinus (Bull.) Karst., Polyporus squamosus Huds.: Fr., Trametes hirsuta (Wulf.) Pil., Pleurotus ostreatus (Jack.) Kumm. (Nikitskij et al., 1996; Semenov, 2007). Subgenus Trochanterella Brundin, 1954 Atheta cribripennis J. Sahlberg, 1890 M a t e r i a l . Literature data only. D i s t r i b u t i o n . Europe, West Siberia (Schülke & Smetana, 2015); in the Ukrainian known from the Carpathian Mountains (Łomnicki, 1911) and Ivano-Frankivsk Region, mountain range Chornohora, mountain Dantsyzh (Lokay, 1912). B i o n o m i c s . Th e species occurs in lowlands and in foothills in forest litter and remains of plants. Subgenus Xenota Mulsant & Rey, 1873 Atheta lativentris (Sahlberg, 1876) M a t e r i a l . 11 ex. IFR: Anheliv tract [Ay], 18.9. [year not specifi ed], 1 ex., LWI: Lviv [Lwów], 5.5. [1] 922, 1 ex., leg. A. Stöeckel; same locality but, date not specifi ed, 3 ex.; same locality but, date not specifi ed, 6 ex., coll. Grolle. (SMNH). D i s t r i b u t i o n . Europe, Altai (Schülke & Smetana, 2015). In the Ukrainian Carpathians 109 Th e Genus Atheta (Coleoptera, Staphylinidae, Aleocharinae) in the Ukrainian Carpathians known from Transcarpathian Region, coniferous forests (Roubal, 1930). “Poland”, without indication of region (Łomnicki, 1913). B i o n o m i c s . Th e species occurs in lowlands and in foothills in forest litter, in fungi, and in ant hills (Roubal, 1930). We would like to take this opportunity and thank V. A. Korneyev, M. Yu. Rusin, J. Frisch, H. Schillham- mer, M. M. Bilyashivskiy, V. B. Rizun, A. Yu. Solodovnikov, V. O. Chumak for the opportunity to work with their respective institutional collections, and their valuable remarks on this manuscript. Th e visit (S. Glotov) to the Entomology Department of ZMUC was carried out through funding from Dr. Bøje Benzons Støttefond and was awarded by A. Yu. Solodovnikov, who contributed to this visit, and support from the entire ZMUC- entomology is appreciated. Th e work was performed within the framework of the scientifi c topic: “Estimation of the biotic diversity of model groups of Arthropoda of the Ukrainian Carpathians with the use of modern information technology”. References Bogdanov, Yu. A. 1985. Fauna and ecology of rove beetles Staphilinids of the Transcarpathia Region. Synopsis of thesis of Ph. D (Biology), Kiev, 1–23 [In Russian]. Brundin, L. 1954 a. Die paläarktischen Arten der Atheta-Untergattung Dimetrota Muls. et Rey (Col., Staphyli- nidae). Eine systematische Studie. Arkiv for Zoologi, 5 (2), 369–434. Brundin, L. 1954 b. In: Hansen, V.: Biller XVII. Rovbiller 3. Del. Danmarks Fauna 59. Kobenhavn, G. E. C., Gads Forlag, 1–499. Brundin, L. 1954 c. 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