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7Трансформация экосистем Ecosystem Transformation
ISSN 2619-094X Print ISSN 2619-0931 Online www.ecosysttrans.com
Article
Beetles (Coleoptera) from the burrows of the great gerbil Rhombomys opimus (Lichtenstein, 1823) (Mammalia: Muridae) in the southern Turkmenistan
Alexey S.Sazhnev1* , Nikolay M. Ermakov2 , Evgeniy N. Kondratyev2
1 Papanin Institute for Biology of Inland Waters, Russian Academy of Sciences, Borok 109, Nekouz District, Yaroslavl Region, 152742, Russia
2 Russian Research Anti-Plague Institute "Microbe", ul. Universitetskaya 46, Saratov, 410005 Russia
Received: 27.05.2022 Revised: 04.06.2022 Accepted: 06.06.2022
Abstract. The study is the first attempt to generalize the data on the burrow biocenoses of the great gerbil (Rhombomys opimus (Lichtenstein, 1823)) in the region of persistent plague epizootics in the lowland part of the Karakum Desert (Southern Turkmenistan). The material was collected in May, November, and December of 1987-1988 using trapping cylinders in residential and non-residential colonies of the great gerbil. Coleoptera made up about 40% of the species composition of invertebrates; their share of the total abundance was 19%. The beetle fauna of the studied burrows was represented by fifteen species belonging to seven families (another four taxa from the Staphylinidae family were identified down to the genus level). Atheta flagellicornis G. Benick, 1967 was the most abundant species, common species were Sepedophilus rufulus (Hochhuth, 1849), Aleochara ja-cobsoni Kirshenblat, 1935 (Staphylinidae), and large zoophagous Eremosphodrus dvorshaki Casale et Vereschagina, 1986 (Cara-bidae). The photographs of Atheta flagellicornis (Staphylinidae) and Bradyus pygmaeus (Fischer von Waldheim, 1821) (Tenebri-onidae) are provided for the first time.
Published online: 22.02.2023
DOI: 10.23859/estr-220527 UDC 595.76; 574.472; 599.325.2
Translated by D.M. Martynova
Keywords: nidicolous beetles, Karakum Desert, beetles, mammals, rodents
To cite this article. Sazhnev, A.S. et al., 2023. Beetles (Coleoptera) from the burrows of the great gerbil Rhombomys opimus (Lichtenstein, 1823) (Mammalia: Muridae) in the southern Turkmenistan. Ecosystem Transformation 6 (1), 39-50. https://doi.org/10.23859/estr-220527
Научная статья
Жесткокрылые (Coleoptera) из нор большой песчанки Rhombomys opimus (Lichtenstein, 1823) (Mammalia: Muridae) на юге Туркмении
А.С. Сажнев1* , Н.М. Ермаков2 , Е.Н. Кондратьев2
1 Институт биологии внутренних вод им. И.Д. Папанина РАН, 152742, Россия, Ярославская обл., Некоузский р-н, пос. Борок, д. 109
2 Российский противочумный институт «Микроб» Роспотребнадзора, 410005, Россия, г. Саратов, ул. Университетская, д. 46
Поступила в редакцию: 27.05.2022 Доработана: 04.06.2022 Принята к печати: 06.06.2022 Опубликована онлайн: 22.02.2023
DOI: 10.23859^г-220527 УДК 595.76; 574.472; 599.325.2
Аннотация. Представлена первая попытка обобщить данные по жукам микробиоценозов нор большой песчанки (Rhombomys opimus (Lichtenstein, 1823)) в районе стойких эпизоотий чумы в низменной части Каракумов (Южная Туркмения). Материал был собран в мае, ноябре и декабре 198788 гг. при помощи ловчих цилиндров в жилых и нежилых колониях большой песчанки. Жесткокрылые составили около 40% видового состава беспозвоночных; их доля от общего числа особей насчитывает 19%. Фауна жесткокрылых исследованных нор представлена 15 видами из 7 семейств (еще 4 таксона из семейства Staphylinidae были определены до рода). Самым массовым в сборах был вид Atheta flagellicomis G. Benick, 1967, обычными - Sepedophilus rufulus (Hochhuth, 1849) и Aleochara jacobsoni Kirshenblat, 1935 (Staphylinidae), а также крупный зоофаг Eremosphodrus dvorshaki Casale et Verescha-gina, 1986 (Carabidae). В статье впервые приводятся фотографии Atheta flagellicomis (Staphylinidae) и Bradyus pygmaeus (Fischer von Waldheim, 1821) (Tenebrionidae).
Ключевые слова: нидиколы, пустыня Каракумы, жуки, млекопитающие, грызуны
Для цитирования. Сажнев, А.С. и др., 2023. Жесткокрылые (Coleoptera) из нор большой песчанки Rhombomys opimus (Lichtenstein, 1823) (Mammalia: Muridae) на юге Туркмении. Трансформация экосистем 6 (1), 39-50. https://doi.org/10.23859/estr-220527
Introduction
The natural focality of some especially dangerous diseases (plague, rickettsiosis, and viral infections) is closely related to the nest/burrow biocenoses of small mammals. A burrow is considered as an elementary focus of such infections. The pathogenic agents circulate in different ways and along different bioceno-tic chains, in which many systematic groups of liv-
ing organisms participate. For this reason, the study of the fauna of burrows of common rodent species, considered the carriers of natural focal infections, is of particular interest. Taking into account the regional nature of a number of natural focal infections, arid territories occupy a special place in epizootic studies.
The study of the inhabitants of burrows and nests in various post-Soviet regions has been carried out
by employees of anti-plague stations and parasitologists since the beginning of the XX century. The most detailed studies of Coleoptera have been performed by Ya.D. Kirshenblat (1935, 1936a, b, 1937a, b), who considered the fundamental issues of the emergence of nidicolous beetles and the general provisions of biocenology in addition to faunistics and taxonomy. Subsequent works on the structure of burrow biocenoses were devoted to the regions adjacent to Turkmenistan, on the example of gerbil burrows (Kashcheev and Iskakov, 1981; Kashcheev, 1982b; Klimova et al., 1970; Klimova and Nelzina, 1971; Nelzina et al., 1978, etc.).
Ya.P. Vlasov (1937) with colleagues (Vlasov and Ioff, 1937; Vlasov and Kirichenko, 1937, etc.) were among the pioneer researchers of rodent nests (mainly different gerbil species) in Turkmenistan, followed by a number of authors (Bakhaeva, 1962; Dudnikova, 1960; Zagniborodova, 1960) studying certain groups of arthropods in the biocenoses of rodent burrows in Turkmenistan. Most publications were devoted to parasite fauna and did not consider other groups of invertebrates, including Coleoptera. The most complete information about the insects of burrowing consortia of gerbils in Turkmenistan (eastern Karakum) was presented by V.A. Krivokhatsky (1982, 1983, 1985) together with V.A. Kashcheev (Krivokhatsky and Kashcheev, 1986).
The study aims to characterize the species composition of beetles (Insecta: Coleoptera) in burrow biocenoses of the great gerbil Rhombomys opimus (Lichtenstein, 1823), which is the most common rodent species of the lowland part of the Central Karakum (Turkmenistan). This report is the first attempt to summarize the data on biocenoses of the great gerbil burrows in this part of its range, i.e., in the area of persistent plague epizootics. Given the present difficulties in studying the fauna and flora of Turkmenistan for foreign scientists, the collected material is largely unique.
Materials and methods
The material was collected by N.M. Ermakov in May, November, and December of 1987-1988 in the natural foci of the plague in the lowland part of the Karakum in the colonies of the great gerbil:
Turkmenistan, Akhal Velayat, Gokdepe Etrap, lowland part of the central Karakum Desert, environs of the city of Gokdepe (before 1993, Geok-Tepe), ridge-cellular sands.
The method of trapping cylinders was used to collect the material (Fasulati, 1971). Sampling was carried out both in residential and non-residential colonies of the great gerbil. Glass jars (0.4-0.5-L volume) were used as the trapping cylinders. A 1% formadehyde solution was used as a fixative; in cold (frosty) weather, glycerol was added. Cylinders
were placed in the burrows at a 25-30-cm distance from the entrance. The upper cut of the cylinder was placed on a par with the soil surface in the burrow or 2-3 mm lower. Two cylinders (internal and external) were installed into each burrow. The total number of cylinders ranged from 5 to 10, depending on the size of the colony. A strip of metal foil (or glass) was inserted into the gap between the cylinders across the course of the burrow, rising 3-5 cm above the surface of the burrow and closely adhering to its walls (Fig. 1). It largely served as a barrier to animals moving along the burrow, which made it possible to estimate the direction of movement of invertebrates in the burrow (from the depth of the burrow or from the surface).
After removal of the cylinders, invertebrates were picked up manually or concentrated onto a fine sieve, then fixed in a 70% alcohol solution. During the field work, 77 samples containing beetles were obtained; in total, about 230 imago beetle specimens were collected. The material was stored in alcohol and mounted dry. Most of material is now stored in the collection of invertebrates of the Papanin Institute for Biology of Inland Waters, Russian Academy of Sciences (Borok, Russia). Some specimens were granted to the collection of the Zoological Institute of the Russian Academy of Sciences (St. Petersburg, Russia). Species nomenclature and general distribution data are taken according to the latest editions of the Catalogue of Coleoptera of the Palearctic (Alonso-Zarazaga et al., 2022; Catalogue..., 2007, 2015, 2017, 2020).
The illustrations were made by the first author using a Leica M165C stereo microscope on a Leica MC170 HD (12 MP) digital camera. Photos were processed and stacked in Sketchbook and Helicon Focus 7.7.4.
The Coleoptera were defined by A.S. Sazhnev with the help of colleagues studying particular taxonomic groups: K.V. Makarov (Carabidae), S. Anla§ and A.V. Kovalev (Staphylinidae), V.O. Kozminykh (Histeridae), L.V. Egorov (Tenebrionidae), and I.A. Zabaluev (Curculionidae).
Results and discussion
Great gerbil is a rodent species common for the lowland part of the central Karakum Desert, it is distributed everywhere in the region, so wider than other gerbils. As a rule, the great gerbils settle in colonies, their burrows are both temporary and permanent, simple and complex. The number of exits/entrances in burrows ranges from 7 to 900. The depth of the nesting chamber in the colonies may reach 2.7 m below the ground surface (Kamnev et al., 1968).
The characteristic features of the microclimate (lack of direct influence of sunlight, moderate thermal
Fig. 1. Scheme of placement of trapping cylinders in rodent burrows. 1 plates.
conical cylinder, 2 - level of the fixative liquid, 3 - separating
conditions, and high relative humidity) make the burrow a special kind of microbiotope (Beklemishev, 1959; Vlasov, 1937). It serves as a habitat both for the host (consortium founder) and for many invertebrates and microorganisms (consorts). In the lowland part of the central Karakum Desert, beetles make up 40% of the species composition of invertebrates in the burrows of the great gerbil, their share in the total abundance of the main taxonomic groups is relatively permanent (19%).
Some beetles belonging to the Staphylinidae family (genera Falagria, Haploglossa, Oxypoda and Heterothops) were not identified down to the species level due to poor preservation and lack of serial material. An annotated list of the species found in the samples is given below.
Annotated list of Coleoptera species from the burrows of the great gerbil in the southern Turkmenistan
Carabidae
Eremosphodrus (Rugisphodrus) dvorshaki Casale et Vereschagina, 1986
Fairly common in our samples (31 ind. in
6 samples). Asian species, distributed in Kazakhstan, Turkmenistan, Uzbekistan, and Tajikistan. Zoophagous. Like many representatives of the Sphodrini tribe, it is a facultative nidicolous beetle.
Staphylinidae
Dropephylla caucasica (Kolenati, 1846) Found singly, but quite constantly (7 ind. in
7 samples). The species was recorded in the central European part of Russia (probably, mistakenly);
noted for the Caucasus, Azerbaijan, Afghanistan, Iran, the Asian part of Turkey, Turkmenistan, and Kyrgyzstan. Probably, a saprophage. The bionomics of the species of the genus Dropephylla is poorly studied, indications for nests of mammals are known (Jaszay and Hlavac, 2006).
Sepedophilus rufulus (Hochhuth, 1849) Common in samples (38 ind. at 30 samples). The species is distributed in the south of the European part of Russia, in Azerbaijan, Armenia, Georgia, Afghanistan, Iran, Iraq, Asian Turkey, Turkmenistan, Uzbekistan, in the south of Western Siberia, in Pakistan, and India (Kashmir). Probably, a saprophage. Common in gerbil burrows (Krivokhatsky and Kashcheev, 1986), where it appears as a facultative nidicolous beetle.
Tachyporus (Palporus) nitidulus (Fabricius, 1781) In the burrows of gerbils, it is represented by a single find (2 ind. in 1 sample). The species has a cosmopolitan distribution; it has been introduced to the New World, Africa, and Australian zoogeographic region. Ubiquist; probably, the finds in the burrows are accidental.
Aleochara (Xenochara) jacobsoni Kirshenblat, 1935
Common in samples (13 ind. in 16 samples). Described from the Mirzachol Steppe (clay-saline desert in Central Asia, located in south Kazakhstan, Turkmenistan, and Uzbekistan) from the burrows of the long-clawed ground squirrel Spermophilop-sis leptodactylus (Lichtenstein, 1823) and the great gerbil (Kirschenblat, 1935). In the Palearctic Catalogue (Catalogue..., 2015), the species is listed for Kazakhstan and Turkmenistan, it is also indicated as the most common species in the burrows of the great gerbil in these regions (Krivokhatsky and Kashcheev, 1986). Recorded in burrows of little ground squirrel (Spermophilus pygmaeus (Pallas, 1778)) (Kirschenblat, 1937a). Obligate facultative nidicolous beetle.
Atheta (Traumoecia) flagellicornis G. Benick, 1967 (Fig. 2)
The most common species in samples (131 ind. in 36 samples). The species is known from the Asian part of Turkey, Syria, and Turkmenistan (Assing, 2006; Catalogue..., 2015), from where it has been described from the vicinity of Ashgabat (Benick, 1967). In Turkey and Syria, the species is found in the litter, on Tamarix sp., on the river bank (Assing, 2006); in Turkmenistan, it is an obligate nidicolous beetle. Perhaps, the taxon includes a complex of species.
Sunius nidicola (Kashcheev, 1982) Relatively rare in samples (3 ind. in 3 samples). The species was described from the nesting chamber of the great gerbil from Kazakhstan (type locality "280 km east of Nukus (Uzbekistan)" (Kashcheev, 1982a) was specified in the revision of the genus (Assing, 2010), where the original holotype label
"Karakalpakia, Chaban-Kazgan" is given (now Kazakhstan, city of Shobankazgan on the border with Uzbekistan)), also known from Turkmenistan. Obligate nidicolous beetle.
Heterothops cf. praevius Erichson, 1839
Relatively common in samples (6 ind. in 6 samples). The species is widely distributed in the Palearctic, known in Central Asia from Uzbekistan and Southeast Kazakhstan (Catalogue..., 2015; Kashcheev, 2002). Like other representatives of the genus, it may be nidicolous beetle optionally.
Philonthus (Philonthus) cf. variabilis Eppelsheim, 1892
Only 1 ind. (female) has been found, so undoubtful identification is problematic. The species is known only from Turkmenistan and Uzbekistan. Possibly may be nidicolous beetle optionally.
Histeridae
Erebidus vlasovi (Reichardt, 1941)
Rare in samples (3 ind. in 2 samples).A comparison with a sibling species E. reichardti Tishechkin & Lackner, 2012, revealed clear differences (reported by V.O. Kozminykh). Endemic to the arid Turan Province, known from single finds from Kazakhstan, Turkmenistan and Uzbekistan. Nidicolous species, associated with sands and loess soils (Kryzhanovsky and Reichard, 1976). Recorded in burrows of gerbils, long-clawed ground squirrel, etc. (Krivokhatsky and Kashcheev, 1986; Kryzhanovsky and Reichard, 1976).
Pholioxenus phoenix (Reichardt, 1929)
Rare in our samples (2 ind. in 2 samples), common in the burrows of the great gerbil (Krivokhatsky and Kashcheev, 1986). Turanian species, noted for Central Asia (Kazakhstan, Turkmenistan, and Uzbekistan). Nidicolous beetles, associated with gerbil burrows on loess and clayey soils, less often on fixed sands (Kryzhanovsky and Reichard, 1976).
Coccinellidae
Coccinella (Coccinella) septempunctata L., 1758
Widely distributed in the Palaearctic. Zoophagous, polyphagous; it mainly consumes aphids that live on both woody and herbaceous vegetation. It is not associated with mammalian burrows; one specimen found is a random case.
Cryptophagidae
Cryptophagus quadrimaculatus Reitter, 1877
Known in samples from 1 sample (4 ind.). Widely distributed in the Palearctic; known from Kazakhstan, Kyrgyzstan, Turkmenistan, Tajikistan, and Uzbekistan in Central Asia. Probably, may be nidicolous beetle optionally.
Tenebrionidae
Bradyus pygmaeus (Fischer von Waldheim, 1821) (Fig. 3)
Relatively rare in samples (3 ind. in 2 samples). Central Asian species: Afghanistan, Kazakhstan, Turkmenistan, and Uzbekistan. Widespread in the
Fig. 2. Atheta (Traumoecia) flagellicornis G. Benick, 1967: A - spermatheca (lateral view), B - tergite VIII and sternite of female, C -aedeagus (dorsal and lateral view), D - general view (male).
Karakum and Kyzylkum deserts (Medvedev and Nepesova, 1985), may be nidicolous beetle optionally.
Curculionidae
Sitona macularius (Marsham, 1802)
In training camps random (1 ind.). The West Palearctic species is associated in development with various types of legumes. First reported for Turkmenistan (Alonso-Zarazaga et al., 2022).
Conclusions
In Turkmenistan, beetles make up about 40% of the species composition of invertebrates and 19% of the total abundance in the burrows of the great gerbil. In the lowland part of the Central Karakum, the beetle fauna of the studied burrows is represented by 15 species from 7 families (another 4 taxa from the Staphylinidae family are identified down to the genus level). Obligate and facultative nidicolous beetles are found, as well as randomly sampled species that are
not the representatives of the burrow fauna. The beetles of the Staphylinidae family (12 taxa) are the most represented in the samples, both quantitatively and qualitatively. The most abundant is Atheta flagellicor-nis, followed by two other species of rove beetles, Sepedophilus rufulus and Aleochara jacobsoni. The large zoophagous Eremosphodrus dvorshaki (Cara-bidae) is also quite common in the samples.
Funding
The work of A.S. Sazhnev was a part of the State Task of the Ministry of Science and Higher Education of the Russian Federation, no. 121051100109-1.
Acknowledgments
The authors are grateful to K.V. Makarov (Moscow State Pedagogical University, Moscow, Russia), S. Anla§ (Ege Universitesi, izmir, Turkey), A.V. Kova-lev (Zoological Institute, Russian Academy of Scienc-
es, St. Petersburg, Russia), V.O. Kozminykh (Perm State Humanitarian Pedagogical University, Perm, Russia), L.V. Egorov (Prisursky State Nature Reserve, Cheboksary, Russia), and I.A. Zabaluev (Severtsov Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, Moscow, Russia).
ORCID
A.S. Sazhnev 0000-0002-0907-5194 N.M. Ermakov 0000-0001-9105-7448 E.N. Kondratyev 0000-0002-7508-4355
References
Alonso-Zarazaga, M.A., Barrios, H., Borovec, R., Bouchard, P., Caldara, R. et al., 2022. Cooperative catalogue of Palaearctic Coleoptera Curculionoidea, Monografías Electrónicas S.E.A. 8, 1-556.
Assing, V., 2006. New species and records of Staphylinidae from Turkey IV, with six new synonymies (Coleoptera: Staphylinidae). Koleopterologische Rundschau 76, 223-276.
Assing, V., 2010. A revision of Palaearctic Sunius. XII. New species, new synonymies, and additional records (Coleoptera: Staphylinidae: Paederinae). Linzer biologische Beiträge 42, 1045-1061.
Bakhaeva, A.V., 1962. Fauna kleshchei Turkmenii [Fauna of ticks of Turkmenistan]. Izvestiia AN TSSR. Seriia biologicheskie nauki [Proceedings of the Turkmen SSR Academy of Sciences. Biological Sciences Series] 5, 349-356. (In Russian).
Beklemishev, V.N., 1959. Populiatsii i mikropopuliatsii parazitov i nidicolov [Populations and micropopulations of parasites and nidicolous beetles]. Zoologicheskii zhurnal [Zoological Journal] 38 (8), 1128-1137. (In Russian).
Benick, G., 1967. Eine neue Atheta (Traumoecia) aus Turkmenien (Coleoptera, Staphylinidae). Entomologische Blätter für Biologie und Systematik der Käfer 63, 30-31.
Catalogue of Palaearctic Coleoptera., 2007. Elateroidea - Derodontoidea - Bostrichoidea -Lymexyloidea - Cleroidea - Cucujoidea. Vol. 4. Löbl, I. and Smetana, A. (eds.). Apollo Books, Stenstrup, Denmark, 935 p.
Catalogue of Palaearctic Coleoptera, 2015. Hydrophiloidea - Staphylinoidea. Revised and updated edition. Vol. 2/1. Löbl, I. and Löbl, D (eds.). Brill, Leiden - Boston, 1702 p.
Catalogue of Palaearctic Coleoptera, 2017. Archostemata - Myxophaga - Adephaga. Revised and updated edition. Vol. 1. Lobl, I. and Lobl, D. (eds.). Brill, Leiden - Boston, 1443 p.
Catalogue of Palaearctic Coleoptera, 2020. Tenebrionoidea. Revised and updated second edition. Vol. 5. Lobl, I. and Smetana, A. (eds.). Apollo Books, Stenstrup, Denmark, 935 p.
Dudnikova, A.F., 1960. Fauna i dinamika chislennosti blokh gryzunov Zapadnykh Karakumov. Soobshchenie 2. Blokhi poludennoi i krasnokhvostoi peschanok, tonkopalykh suslikov i tarbaganchikov [Fauna and population dynamics of rodent fleas in the Western Karakum desert. Report 2. Fleas of midday and libyan gerbils, long-clawed ground squirrels and dwarf fat-tailed jerboas]. In: Fenyuk, B.K. (ed.), Voprosy prirodnoi ochagovosti i epizootii chumy v Turkmenii [Issues of Natural Foci and Epizootics of Plague in Turkmenistan]. Poligraphkombinat, Ashgabat, USSR, 335-341. (In Russian).
Fasulati, K.K., 1971. Polevoe izuchenie nazemnykh bespozvonochnykh [Field study of terrestrial invertebrates]. Vysshaia Shkola, Moscow, USSR, 424 p. (In Russian).
Jaszay, T., Hlavac, P., 2006. A revision of the Palaearctic species of the genus Dropephylla (Coleoptera: Staphylinidae: Omaliinae). Entomological Problems 36 (1), 31-62.
Kamnev, P.I., Zhernovov, I.V., Trofimova, R.K., Skvortsov, G.N., Bogdanova, A.I., Panova, O.M. 1968. Kharakter poselenii gryzunov i ikh ektoparazitov v Kopet-Dage v svyazi s problemoi enzootii chumy [Characteristics of the settlements of rodents and their ectoparasites in Kopet-Dag in connection with the problem of plague enzootics]. In: Fenyuk, B.K. (ed.), Gryzuny i ikh ektoparazity [Rodents and their ectoparasites]. Saratov University Publishing House, Saratov, USSR, 140149. (In Russian).
Kashcheev V.A., 1982a. Novye vidy zhukov-stafilinov (Coleoptera, Staphylinidae) iz pustyni Kyzylkum [New species of rove beetles (Coleoptera, Staphylinidae) from the Kyzylkum desert]. Entomologicheskoe obozrenie [Entomological Review] 61 (3), 537-541. (In Russian).
Kashcheev, V.A., 1982b. Struktura mikrobiotsenoza nory bol'shoi peschanki severnykh i tsentral'nykh Kyzylkumakh [The structure of the microbiocenosis
of the burrow of the great gerbil in the northern and central Kyzylkum Desert]. Izvestiia AN Kazakhskoi SSR. Seriia biologicheskie nauki [Proceedings of the Kazakh SSR Academy of Sciences. Biological Sciences Series] 3, 31-38. (In Russian).
Kashcheev, V.A., 2002. Naselenie stafilinid (Col., Staph.) v agrotsenozakh iugo-vostoka Kazakhstana [The population of Staphilinid beetles (Col., Staph.) in the agrocenoses of the south-east of Kazakhstan]. Tethys Entomological Research 7, 179-192. (In Russian).
Kashcheev, V.A., Iskakov, B.V., 1981. Stafilinidy (Coleoptera, Staphylinidae) iz kolonii nor bol'shoi peschanki (Rhombomis opimus Licht.) v pustyne Kyzylkum [Staphylinids (Coleoptera, Staphylinidae) from colonies of burrows of the great gerbil (Rhombomis opimus Licht.) in the Kyzylkum Desert]. Izvestiia AN Kazakhskoi SSR. Seriia biologicheskie nauki [Proceedings of the Kazakh SSR Academy of Sciences. Biological Sciences Series] 5, 35-40. (In Russian).
Kirshenblat, Ya.D., 1935. K voprosu o proiskhozhdenii nekotorykh vidov nidicolov (obitatelei gnezd) [On the question of the origin of some types of nidicolous beetles (nest dwellers)]. Doklady AN SSSR [Proceedings of the USSR Academy of Sciences] 2 (3-4), 332-337. (In Russian).
Kirshenblat, Ya.D., 1936a. Zhuki-staphilinidy iz nor gryzunov na Yugo-Vostoke RSFSR [Rove beetles from rodent burrows in the South-East of the RSFSR]. Vestnik mikrobiologii, epidemiologii i parazitologii[Bulletin of Microbiology, Epidemiology, and Parasitology] 15 (2), 249-253. (In Russian).
Kirshenblat, Ya.D., 1936b. Fauna gnezd mlekopitaiushchikh i ptits [Fauna of nests of mammals and birds]. Voprosy ekologii i biotsenologii [Problems of Ecology and Biocenology] 3, 45-56. (In Russian).
Kirshenblat, Ya.D., 1937a. Zhuki-stafiliny v gnezdakh Citellus pygmaeus Pall [Rove beetles in the nests of Citellus pygmaeus Pall.]. Vestnik mikrobiologii, epidemiologii i parazitologii [Bulletin of Microbiology, Epidemiology, and Parasitology] 16 (1-2), 171-185. (In Russian).
Kirshenblat, Ya.D., 1937b. Opredelitelnye tablitsy zhukov stafilinid zhivushchikh v gnezdakh mlekopitaiushchikh i ptits [Key to Staphilinid beetles living in the nests of mammals and birds]. Vestnik mikrobiologii epidemiologii i parazitologii [Bulletin
of Microbiology, Epidemiology, and Parasitology] 16 (1-2), 227-242. (In Russian).
Klimova, Z.I., Borutzky, E.V., Evseeva, V.E., Medvedev, S.I., Nelzina, E.N., Serzhanov, O.S., Kharitonov, D.E., 1970. Struktura nornykh biotsenozov bol'shoi peschanki v severnykh Kyzylkumakh. Otchet I. Taksonomicheskie i ekologicheskie gruppy artropod uchastvuyuschikh v formirovanii mikrobiotsenosov [The structure of burrow microbiocoenoses of the great gerbil in the northern Kizilkum. Report I. Taxonomic and ecological groups of arthropods participating in the formation of microbiocoenoses]. Parazitologiya 5, 437-442. (In Russian).
Klimova, Z.I., Nelzina, E.N., 1971. Morfo-funktsionalnaia organizatsiia norovykh mikrobiotsenozov bolshoi peschanki v Severnykh Kyzylkumakh. Soobshchenie 2. Sezonnye aspekty v strukture norovykh mikrobiotsenozov [Morpho-functional organization of burrow biocenoses of the great gerbil in the Northern Kyzylkum Desert. Report 2. Seasonal aspects in the structure of burrow biocenoses]. Meditsinskaia parazitologiia i parazitarnye bolezni [Medical Parasitology and Parasitic Diseases] 5, 566-571. (In Russian).
Krivokhatskii, V.A., 1982. Chlenistonogie zhivotnye iz khodov nor peschanok v Vostochnykh Karakumakh [Arthropods from the burrows of gerbils in the Eastern Karakum Desert]. Ekologiya [Russian Journal of Ecology] 2, 60-64. (In Russian).
Krivokhatskii, V.A., 1983. K izuchennosti norovykh konsortsii bolshoi peschanki [On the study of burrow consortia of the great gerbil]. Problemy osvoeniia pustyn [Problems of Desert Development] 1, 5968. (In Russian).
Krivokhatskii, V.A., 1985. Nasekomye norovykh konsortsii peschanykh pustyn Srednei Azii [Insects of burrow consortia of sandy deserts of Central Asia.]. Biological sciences PhD thesis abstract. Leningrad, USSR, 1-20. (In Russian).
Krivokhatskii, V.A., Kashcheev, V.A., 1986. Stafilinidy (Coleoptera, Staphylinidae) iz nor gryzunov i drugikh mestoobitanii Repetekskogo zapovednika [Staphilinid beetles (Coleoptera, Staphylinidae) from rodent burrows and other habitats of the Repetek Biosphere State Reserve]. Izvestiia AN TSSR. Seriia biologicheskie nauki [Proceeedings of the Turkmen SSR Academy of Sciences. Biological Sciences Series] 3, 26-31. (In Russian).
Kryzhanovsky, O.L., Reichardt, A.N., 1976. Zhuki nadsemeystva Histeroidea (semeystva Sphaeritidae, Histeridae, Synteliidae) (Fauna SSSR. Zhestkokrylyye. T. 5, vyp. 4) [Beetles of the superfamily Histeroidea (families Sphaeritidae, Histeridae, Synteliidae) (Fauna of the USSR. Coleoptera. Vol. 5, Is. 4)]. Nauka, Moscow-Leningrad, USSR, 435 p. (In Russian).
Medvedev, G.S., Nepesova, M.G., 1985. Opredelitel zhukov-chernotelok Turkmenistana [Key to darkling beetles of Turkmenistan]. Ylym, Ashgabat, USSR, 180 p. (In Russian).
Nelzina, E.N., Klimova, Z.I., Protopopjan, M.G., 1978. Striktura i stepen' nornykh biotsenozov Rhombomys opimus Lichtenstein v razlichnykh chastyakh areala [Structure and degree of burrow microbiocoenoses Rhombomys opimus Lichtenstein in different parts of its area of distribution]. In: Gilyarov, M.S. (ed.), Resul'taty i perspektivy parazitotsenologii v SSSR [Results and Prospects of Parasite Coenology in the USSR]. Nauka, Moscow, USSR, 87-97. (In Russian).
Vlasov, Ya.P., 1937. Nora, kak svoeobraznyi biotop v okrestnostiakh Ashkhabada [Burrow as a kind of biotope near Ashgabat]. Problemy parazitologii i fauny Turkmenii [Problems of parasitology and fauna of Turkmenistan] 9, 223-240. (In Russian).
Vlasov, Ya.P., Ioff, I.G., 1937. Blokhi iz nor v okrestnostiakh Ashkhabada [Fleas from burrows near Ashgabat]. Problemy parazitologii i fauny Turkmenii [Problems of Parasitology and Fauna of Turkmenistan] 9, 277-282. (In Russian).
Vlasov, Ya.P., Kirichenko, A.N., 1937. Hemiptera-Heteroptera iz nor v okrestnostiakh Ashkhabada [Hemiptera-Heteroptera from burrows near Ashgabat]. Problemy parazitologii i fauny Turkmenii [Problems of Parasitology and Fauna of Turkmenistan] 9, 263-268. (In Russian).
Zagniborodova, E.N., 1960. K faune i ekologii blokh Zapadnoi Turkmenii [On the fauna and ecology of fleas in Western Turkmenistan]. In: Fenyuk, B.K. (ed.), Voprosy prirodnoi ochagovosti i epizootii chumy v Turkmenii [Issues of Natural Foci and Epizootics of Plague in Turkmenistan]. Poligraphkombinat, Ashgabat, USSR, 320-333. (In Russian).
Финансирование
Работа А.С. Сажнева проведена в рамках выполнения государственного задания Министерства науки и высшего образования РФ № 121051100109-1.
Благодарности
За помощь в определении авторы признательны К.В. Макарову (МПГУ, Москва), S. Anla§ (Ege Üniversitesi, izmir), А.В. Ковалеву (ЗИН РАН, Санкт-Петербург), В.О. Козьминых (ПГГПУ, Пермь), Л.В. Егорову (Государственный природный заповедник «Присурский», Чебоксары) и И.А. Забалуеву (ИПЭЭ РАН, Москва).
Список литературы
Бахаева, А.В., 1962. Фауна клещей Туркмении. Известия АН ТССР. Серия биологические науки 5, 349-356.
Беклемишев, В.Н., 1959. Популяции и микропопуляции паразитов и нидиколов. Зоологический журнал 38 (8), 1128-1137.
Власов, Я.П., 1937. Нора, как своеобразный биотоп в окрестностях Ашхабада. Проблемы паразитологии и фауны Туркмении 9, 223-240.
Власов, Я.П., Иофф, И.Г., 1937. Блохи из нор в окрестностях Ашхабада. Проблемы паразитологии и фауны Туркмении 9, 277-282.
Власов, Я.П., Кириченко, А.Н., 1937. Hemiptera-Heteroptera из нор в окрестностях Ашхабада. Проблемы паразитологии и фауны Туркмении 9, 263-268.
Дудникова, А.Ф., 1960. Фауна и динамика численности блох грызунов Западных Каракумов. Сообщение 2. Блохи полуденной и краснохвостой песчанок, тонкопалых сусликов и тарбаганчиков. В: Фенюк, Б.К. (ред.), Вопросы природной очаговости и эпизоотий чумы в Туркмении. Полиграфкомбинат, Ашхабад, СССР, 335-341.
Загнибородова, Е.Н., 1960. К фауне и экологии блох Западной Туркмении. В: Фенюк, Б.К. (ред.), Вопросы природной очаговости и эпизоотий чумы в Туркмении. Полиграфкомбинат, Ашхабад, СССР, 320-333.
Камнев, П.И., Жерновов, И.В., Трофимова, Р.К., Скворцов, Г.Н., Богданова, А.И., Панова, О.М., 1968. Характер поселений грызунов и их эктопаразитов в Копет-Даге в связи с проблемой энзоотии чумы. В: Фенюк, Б.К. (ред.), Грызуны и их эктопаразиты. Издательство Саратовского университета, Саратов, СССР, 140-149.
Кащеев В.А., 1982a. Новые виды жуков-стафилинов (Coleoptera, Staphylinidae) из пустыни Кызылкум. Энтомологическое обозрение 61 (3), 537-541.
Кащеев, В.А., 1982b. Структура микробиоценоза норы большой песчанки северных и центральных Кызылкумах. Известия АН Казахской ССР. Серия биологические науки 3, 31-38.
Кащеев, В.А., 2002. Население стафилинид (Col., Staph.) в агроценозах юго-востока Казахстана. Tethys Entomological Research 7, 179-192.
Кащеев, В.А., Искаков, Б.В., 1981. Стафилиниды (Coleoptera, Staphylinidae) из колоний нор большой песчанки (Rhombomis opimus Licht.) в пустыне Кызылкум. Известия АН Казахской ССР. Серия биологические науки 5, 35-40.
Киршенблат, Я.Д., 1935. К вопросу о происхождении некоторых видов нидиколов (обитателей гнезд). Доклады АН СССР 2 (3-4), 332-337.
Киршенблат, Я.Д., 1936a. Жуки стафилиниды из нор грызунов на юго-востоке РСФСР. Вестник микробиологии эпидемиологии и паразитологии 15 (2), 249-253.
Киршенблат, Я.Д., 1936b. Фауна гнезд млекопитающих и птиц. Вопросы экологии и биоценологии 3, 45-56.
Киршенблат, Я.Д., 1937a. Жуки-стафилины в гнездах Citellus pygmaeus Pall. Вестник микробиологии, эпидемиологии и
паразитологии 16 (1-2), 171-185.
Киршенблат, Я.Д., 1937b. Определительные таблицы жуков стафилинид живущих в гнездах млекопитающих и птиц. Вестник микробиологии эпидемиологии и паразитологии 16 (1-2), 227242.
Климова, 3.И., Боруцкий, В.Е., Евсеева, Е.В., Медведев, С.И., Нельзина Е.Н.,
Сержанов, О.С., Харитонов, Д.Е., 1970. Структура норовых микробиоценозов большой песчанки в Северных Кызылкумах. Отчет I. Таксономические и экологические группировки нидиколов, участвующих в формировании микробиоценозов. Паразитология 5, 437-442.
Климова, З.И., Нельзина, Е.Н., 1971. Морфо-функциональная организация норовых микробиоценозов большой песчанки в Северных Кызылкумах. Сообщение 2. Сезонные аспекты в структуре норовых микробиоценозов. Медицинская паразитология и паразитарные болезни 5, 566-571.
Кривохатский, В.А., 1982. Членистоногие животные из ходов нор песчанок в Восточных Каракумах. Экология 2, 60-64.
Кривохатский, В.А., 1983. К изученности норовых консорций большой песчанки. Проблемы освоения пустынь 1, 59-68.
Кривохатский, В.А., 1985. Насекомые норовых консорций песчаных пустынь Средней Азии. Автореферат диссертации на соискание ученой степени кандидата биологических наук. Ленинград, СССР, 1-20.
Кривохатский, В.А., Кащеев, В.А., 1986. Стафилиниды (Coleoptera, Staphylinidae) из нор грызунов и других местообитаний Репетекского заповедника. Известия АН ТССР. Серия биологические науки 3, 26-31.
Крыжановский О.Л., Рейхардт А.Н., 1976. Жуки надсемейства Histeroidea (семейства Sphaeritidae, Histeridae, Synteliidae) (Фауна СССР. Жесткокрылые. Т. V, вып. 4). Наука, Москва - Ленинград, СССР, 435 с.
Медведев, Г.С., Непесова, М.Г., 1985. Определитель жуков-чернотелок
Туркменистана. Ылым, Ашхабад, СССР, 180 с.
Нельзина, Е.Н., Климова, З.И., Протопопян, М.Г., 1978. Структура и степень организованности норовых микробиоценозов большой песчанки в разных частях ее ареала. В: Гиляров, М.С. (ред.), Итоги и перспективы исследований по паразитологии в СССР. Наука, Москва, СССР, 87-97.
Фасулати, К.К., 1971. Полевое изучение наземных беспозвоночных. Высшая школа, Москва, СССР, 424 с.
Alonso-Zarazaga, M.A., Barrios, H., Borovec, R., Bouchard, P., Caldara, R. et al., 2022. Cooperative catalogue of Palaearctic Coleoptera Curculionoidea, Monografías Electrónicas S.E.A. 8, 1-556.
Assing, V., 2006. New species and records of Staphylinidae from Turkey IV, with six new synonymies (Coleoptera: Staphylinidae). Koleopterologische Rundschau 76, 223-276.
Assing, V., 2010. A revision of Palaearctic Sunius. XII. New species, new synonymies, and additional records (Coleoptera: Staphylinidae: Paederinae). Linzer biologische Beiträge 42, 1045-1061.
Benick, G., 1967. Eine neue Atheta (Traumoecia) aus Turkmenien (Coleoptera, Staphylinidae). Entomologische Blätter für Biologie und Systematik der Käfer 63, 30-31.
Catalogue of Palaearctic Coleoptera., 2007. Elateroidea - Derodontoidea - Bostrichoidea -
Lymexyloidea - Cleroidea - Cucujoidea. Vol. 4. Lobl, I. and Smetana, A. (eds.). Apollo Books, Stenstrup, Denmark, 935 p.
Catalogue of Palaearctic Coleoptera, 2015. Hydrophiloidea - Staphylinoidea. Revised and updated edition. Vol. 2/1. Lobl, I. and Lobl, D (eds.). Brill, Leiden - Boston, 1702 p.
Catalogue of Palaearctic Coleoptera, 2017. Archostemata - Myxophaga - Adephaga. Revised and updated edition. Vol. 1. Lobl, I. and Lobl, D. (eds.). Brill, Leiden - Boston, 1443 p.
Catalogue of Palaearctic Coleoptera, 2020. Tenebrionoidea. Revised and updated second edition. Vol. 5. Lobl, I. and Smetana, A. (eds.). Apollo Books, Stenstrup, Denmark, 935 p.
Jaszay, T., Hlavac, P., 2006. A revision of the Palaearctic species of the genus Dropephylla (Coleoptera: Staphylinidae: Omaliinae). Entomological Problems 36 (1), 31-62.
