Invertebrate Zoology, 2018, 15(2): 153–213
© INVERTEBRATE ZOOLOGY, 2018
Subterranean biota of the European part of Russia:
A review
Sergei I. Golovatch1, Dmitry M. Palatov2, Ilya S. Turbanov3,4,
Vladimir A. Kniss5, Suren Gazaryan6, Vladimir P. Snit’ko7,
†Vasile Decu8, Christian Juberthie9, George Nazareanu10,11
1
Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, Leninsky pr., 33,
Moscow, 119071 Russia. E-mail: sgolovatch@yandex.ru
2
Department of Hydrobiology, Biological Faculty, Moscow State University, Leninskie Gory 1/12,
Moscow, 119234 Russia; E-mail: triops@yandex.ru
3
Papanin Institute of the Biology of Inland Waters, Russian Academy of Sciences, Borok, Nekouzskiy
District, Yaroslavl Region, 152742 Russia. E-mail: turba13@mail.ru
4
Cherepovets State University, Lunacharskogo pr., 5, Cherepovets, Vologda Region, 162600
Russia.
5
Bashkirian University, 32 rue Frunze, Ufa, 450074 Russia.
6
UNEP/EUROBATS Secretariat, Platz der Vereinten Nationen 1, D-53113 Bonn, Germany. E-mail:
s.gazaryan@gmail.com
7
Il’men State Nature Reserve, Chelyabinsk Region, Miass, 456317 Russia. E-mail:
snitko@ilmeny.ac.ru
8
Dept. of Biospeleology and Karst Edaphobiology,“Emil Racovitza” Institute of Speleology,
Romanian Academy, Calea 13 Septembrie, 13, Bucharest, 050731 Romania.
9
Encyclopédie Biospéologique, Edition. 1 Impasse Saint-Jacques, F-09190 Saint-Lizier, France. Email: christianjuberthie@wanadoo.fr
10
“Grigore Antipa” Nationa Museum of Natural History, Sos. Kiseleff, 1, 011341 Bucharest,
Romania.
11
Karin’s Kids Academy – Nature School, Sos. Bucuresti–Ploiesti, 182A, 013697 Bucharest,
Romania. E-mail: george.nazareanu@karinkids.ro
ABSTRACT. The endogean flora and fauna of European Russia, including those of the
northwestern Caucasus, but excluding those of Crimea, presently comprise at least 389
species or subspecies (some still unidentified) from 229 genera (a few still unidentified),
150 families, 75 orders, 48 classes and 25 phyla, among which 96 species or subspecies
represent presumed stygo- or troglobionts. The taxonomically most diverse phylum is
Arthropoda which encompasses the bulk of the fauna, in particular due to crustaceans,
beetles and collembolans. Whereas the troglofauna of northern Russia is completely devoid
of such strongly cave-adapted animals, in the Urals, Cis-Urals and central Russia there are
a few, but the greatest diversity of hypogean species in the region is observed in the
northwestern Caucasus, in particular its maritime western part, as an “Ice Age” refugium of
nemoral biota rich in karst.
How to cite this article: Golovatch S.I., Palatov D.M., Turbanov I.S., Kniss V.A., Gazaryan
S., Snit’ko V.P., Decu V., Juberthie Ch., Nazareanu G. 2018. Subterranean biota of the
European part of Russia: A review // Invert. Zool. Vol.15. No.2. P.153–213. doi: 10.15298/
invertzool.15.2.01
KEY WORDS: Eastern Europe, Urals, Cis-Urals, northern Caucasus, endogean invertebrates, bats, stygophiles, stygobionts, troglophiles, troglobionts.
154
S.I. Golovatch et al.
Подземная биота европейской части России: Обзор
С.И. Головач1, Д.М. Палатов2, И.С. Турбанов3,4, В.А. Книсс5,
Сурен Газарян6, В.П. Снитько7, †Василе Деку8,
Кристиан Жюберти9, Георге Назаряну10,11
Институт проблем экологии и эволюции РАН, Ленинский проспект, 33, Москва, 119071
Россия.
2
Кафедра гидробиологии, Биологический факультет МГУ, Ленинские Горы, 1/12, Москва,
119234 Россия
3
Институт биологии внутренних вод РАН, пос. Борок, Ярославская обл., 152742 Россия.
4
Череповецкий государственный университет, проспект Луначарского, 5, Вологодская обл.,
162600 Россия.
3
Башкирский государственный университет, ул. Фрунзе, 32, Уфа, 450074 Россия.
6
Секретариат UNEP/EUROBATS, Бонн, Германия.
7
Ильменский государственный заповедник, г. Миасс, Челябинская обл., 456317 Россия.
8
Институт спеологии, Румынская академия наук, Бухарест, Румыния.
9
Издательство Биоспелеологической энциклопедии, Сен-Лизье, Франция.
10
Музей естественной истории, Бухарест, Румыния.
11
Детская академия Карины, Бухарест, Румыния.
1
РЕЗЮМЕ. Подземные флора и фауна европейской части России, включая таковые
Северо-Западного Кавказа, но исключая таковые Крыма, в настоящее время насчитывают по меньшей мере 389 вида и подвида (некоторые пока не определены) из 229
родов (малая часть пока не определена), 150 семейств, 75 отрядов, 48 классов и 25
типов, среди которых 96 видов и подвидов — предположительно стиго- и троглобионты. Таксономически самый разнообразный тип — это Arthropoda, который включает большинство фауны, особенно за счет ракообразных, жуков и коллембол. В то
время, как троглофауна северной части России совершенно лишена сильно адаптированных в жизни в пещерах и подземных водах животных, на Урале, в Предуралье и
центре России таковых немного, а наибольшее разнообразие гипогейных видов в
регионе наблюдается на Северо-Западном Кавказе, прежде всего, в его приморской
западной части, как богатого карстом «ледникового» рефугиума неморальной биоты.
Как цитировать эту статью: Golovatch S.I., Palatov D.M., Turbanov I.S., Kniss V.A.,
Gazaryan S., Snit’ko V.P., Decu V., Juberthie Ch., Nazareanu G. 2018. Subterranean biota
of the European part of Russia: A review // Invert. Zool. Vol.15. No.2. P.153–213. doi:
10.15298/invertzool.15.2.01
КЛЮЧЕВЫЕ СЛОВА: Восточная Европа, Урал, Предуралье, Северный Кавказ,
подземные беспозвоночные, летучие мыши, стигофилы, стигобионты, троглофилы,
троглобионты.
Introduction
The present paper is the result of a compromise. It was initiated by one of us (CJ) to
become a chapter entitled “RUSSIA (Russian
Federation: European part)” in a forthcoming
volume of “ENCYCLOPAEDIA BIOSPEOLOGICA”. The chapter was duly prepared, but
then remained shelved for more than a year
waiting for some other contributions to be com-
pleted for the same volume. However, because
of uncertainty and an apparent delay in publication, let alone an unexpected abortion of the part
pertaining to mammals in our prepared draft, we
have finally decided to publish this material
elsewhere in a reputed journal and before it is
outdated. This decision actually follows that of
Juberthie et al. (2016) who published their chapter on the cave faunas of Siberia and the Russian
Far East as a journal paper. The same concerns
Subterranean biota of European Russia
the chapter on the Mexican hypogean fauna by
Palacios-Vargas et al. (2017). As regards the
present contribution that covers European Russia, a separate chapter will be devoted to the
troglofauna of Crimea.
I. General description of the area
The Russian Federation is the largest country in the world (17,075,000 km2), spanning
10,000 km from the Baltic Sea in the West to the
Pacific (the Sea of Japan, the Okhotsk Sea, the
Bering Sea) in the East. The Ural Mountain
Range represents the traditional border between
European Russia and Siberia (Asian Russia),
lying between Europe and Asia.
The European part consists of large plains;
the southern part of European Russia consists of
the northern and western Caucasus, including a
part of Transcaucasia stretched along the Black
Sea coast until Sochi and the Psou River, both
bordering on Abkhazia. Central Siberia is mostly a large plateau; southern Siberia bordering on
Mongolia and China, as well as the Far East as
far as the Sea of Okhotsk and the Sea of Japan
are mountainous regions.
Climate. Temperate and continental to the
West, with marine influence of the Baltic Sea
and the Sea of Japan; cold and subpolar to the
North, nearly subtropical in the Sochi region
along the Black Sea. The latitude, the distance
to the ocean and the localization of mountains
account for climate continentality; the climate
of Siberia is characterized by mostly very severe and harsh winters.
Vegetation. From north to south, the pattern
of nature zonation is formed by a marked succession of the following biomes: polar deserts,
tundras, taiga (= boreal, mostly coniferous forest), broadleaved forest, forested steppes, steppes
(= grasslands), semi-deserts and deserts.
II. History
II. 1. Karst
In Russia, the first descriptions of karst
forms and caves were made as early as the 18th
century by several academic expeditions, fol-
155
lowing the conquest of the huge territories of the
Urals, Siberia and, later, the Far East. The first
detailed field manual on caves was published by
J. Gmelin as a result of the Great North Expedition carried out between 1733 and 1743.
At the beginning of the 19th century, V.
Severgin published a review of the caves in the
Russian Empire.
In 1887, Yuri Listov published a field manual on cave studies with notes on their geology,
morphology, hydro-geology, speleogenesis, sediments and microclimate.
In 1900, Alexander Kruber published an
important book, “About karst phenomena in
Russia”. The role of Kruber in Russia is somewhat comparable to that of E.A. Martel in Western Europe. Martel visited Russia in 1903.
From the beginning of the 20th century, the
number of regional karst cave studies continued
to grow in all of the main regions of Russia.
The revolution of 1917 interrupted karst and
cave research in the country, but it resumed in
the USSR.
The first Karst Conference in the USSR was
held in 1933 at Kizel, Perm Region. Since 1947,
the Perm State University sponsored all-Union
conferences on karst and played important roles
in the coordination of karst speleological investigations.
Regional thematic conferences on karst and
speleology were organized by the Karst Commission, for instance on Applied Karstology,
Russian Plain karst, Bashkirian karst, Primorskyi Krai karst, Caves, engineering karstology
etc.
According to A. Klimchouk (2003c), modern speleology in the USSR was born in 1958,
when the Academy of Sciences created the
Interdepartmental Commission on Geological
and Geographical Karst Investigations.
General studies on Russian karst were published by Klimchouk (2003b), Maksimovitch
(1962, 1963), Maksimovitch and Kostarev
(1973), Gorbunova et al. (1992) and many others. Special studies were published on gypsum
karsts of the Russian Eastern-European Plain
and the Pre-Ural region by Andrejchuk (1996),
and Andrejchuk and Klimchouk (1996).
156
S.I. Golovatch et al.
Until 1958, only about 500 caves and shafts
were reported in the Soviet Union as a whole.
From 1958 to 1962, cave-exploring groups were
formed, with as many as 3,800 caves discovered
in the USSR (Russia, Ukraine, Caucasus)
(Dublyanskiy, Ilyukhin, 1982; Kiselyov, Klimchouk, 1991).
II. 2. Cave fauna
The first mention concerning Russia’s cave
fauna belonged to P. Pallas, a famous academician and traveler, who, in 1811, reported the
presence of bat colonies in caves of Crimea and
the Urals. Then around the middle of the 19th
century, as industrial development required resources, geological research was conducted in
the eastern regions such as the Urals, the Altais
and Transbaikalia, as well as in the Caucasus
and other regions of the Russian Empire.
Victor von Motschoulsky, a famous entomologist, first visited several Caucasian caves
in 1850, then some caves in Crimea.
In 1889, G. Jakobson and R. Schmidt sampled fauna in the Murandimovskaya and Kapova caves in the southern Urals and recorded the
dipteran Blepharoptera modesta, Culicidae larvae and adults, Collembola and Aranei. From
among the species collected, Skorikow (1899)
described a new species of springtail, Tomocerus baschkiricus.
Understanding the importance of cave studies in resolving biogeographical problems and
those linked to evolutionary theory, G. Kozhevnikov, professor of the Moscow Universty, created in the early 20th century, inside the Zoological Museum, a “dark room” for experimenting
on the impact of darkness upon Crustacea and
fishes (Ognev, 1910; Kapterev, 1910, 1912).
Nevertheless, the true beginning of biospeleological research in Russia is dated 1912,
when A. Shugurov generalized the then available data, relatively few, on the cave fauna of the
Caucasus and Crimea (Shugurov, 1912).
Biospeleological investigations were resumed only in 1926–1927, already in the USSR,
with explorations of some caves in the Urals and
the description by E. Borutzky in 1928 of the
first stygobiont, the amphipod Crangonyx chlebnikovi from the central Urals. In 1929, Borutz-
ky joined the first biospeleological survey of
two karstic caves in the Kutaisi region of Georgia, Transcaucasia, and published the first data
about the cave fauna of Transcaucasia (Borutzky, 1930, 1934).
In 1927 and 1930, A.N. Derzhavin published two papers containing descriptions of
new cave amphipods.
At the end of 1930, K. Verhoeff described a
diplopod, Leucogeorgia longipes, from a cave
near Kutaisi, Georgia (Verhoeff, 1930).
Since 1935, Ya. Birstein undertook 6 trips in
the Caucasus and explored 39 caves: 31 along
the Black Sea coast, 4 in the Imeretian region of
Georgia and 3 in Crimea.
In 1940, Birstein and Lopashov published a
synthesis of the cave fauna of the USSR as
revealed in 1935–1940. That paper also marked
the onset of the series “Biospelogica Sovietica”
(Birstein, Lopashov, 1940). At the same time, I.
Lapshov (1940) reviewed the pseudoscorpion
cave fauna of Transcaucasia.
In 1950, Birstein generalized and updated
all available information on the cave fauna of
western Transcaucasia (Birstein, 1950).
In 1950, E. Borutzky described a stygobiont
copepod, Speocyclops lussianus, from Ciscaucasia (Borutzky, 1950).
From 1950 to 1970, Birstein alone or together with Ljovuschkin, Borutzky, Levanidov or
Lopashov, published mostly on aquatic cave
fauna. This work culminated in a milestone
synthesis of biospeleological research in the
USSR (Birstein, Ljovuschkin, 1967a). During
the same period, Ljovuschkin alone or with coauthors described a number of cave beetles,
crustaceans and arachnids, Borutzky described
numerous new species of cave copepods and
isopods, Ya. Starobogatov and P. Matiokin
worked on molluscs, N. Zalesskaja on lithobiomorphs, I. Malevich on earthworms, etc.
Since the 1970’s, V. Kniss has studied the
fauna of caves and springs, mostly in the Urals
and in Siberia. In 2001, he published a summarizing monograph, “Cave fauna of Russia and
adjacent regions”. From 1984 to 2006, V. Kniss
published mostly on springtail cave fauna (Kniss,
1984a, b, 1985, 1989, 1991, 2001, 2004, 2006).
Subterranean biota of European Russia
It is Birstein who may be credited as the
pioneer of biospeleology in the USSR, with
several of his colleagues or direct students having provided the foundations for our present
knowledge of the endogean faunas of the exUSSR countries.
The cave invertebrate fauna of the former
Soviet Union, which represents the bulk of
endogean biodiversity, has very recently been
revised and summarized (Turbanov et al., 2016a,
b, c). Members of 17 phyla, 38 classes, 90
orders and 278 families of invertebrates which
contain at least 308 species or subspecies of
mostly presumed stygo– or troglobionts and no
fewer than 735 species or subspecies of stygo–
or troglophiles are currently known to populate
the caves and subterranean waters of Russia and
other countries of the former USSR. The main
evolutionary burst in the endogea, including the
MSS (milieu souterrain superficiel, or mesovoid shallow stratum), is due to arthropods,
primarily crustaceans, collembolans and beetles. The major centres of taxonomic diversity
among stygo– and troglobionts thereby remain
the Caucasus (181, or almost 59%) and Crimea
(44, or over 14%), both montane “glacial” refugia of nemoral biota rich in karst, whereas the
contributions of the other major regions, sometimes strongly karstified as well, are considerably smaller, gradually decreasing towards Central Asia (35, or more than 11%), the Far East
(33, or nearly 11%), the Ukrainian Carpathians
together with Podolia (12, or almost 4%), the
Russian Plain (7, or over 2%), the Urals and
Cisuralia (7, also more than 2%) and Siberia (5,
or 1.6%). The problem of inventorying is still
among the most topical in assessing the stygo–
and troglofaunas of the territories in question.
Although occasional findings of bats in caves
were reported by many Soviet zoologists (see
Kuzyakin (1950)) and the importance of these
sites was obvious, Petr Petrovich Strelkov was
the first Russian bat researcher who focused on
underground sites and conducted an extensive
survey of those caves and artificial underground
roosts that were known at that time in European
Russia (Strelkov, 1958), establishing a tradition
for winter bat monitoring. Following his pilot
157
survey, underground sites had become a key
element of all regional studies of bat fauna.
They gained a momentum after the first allUSSR bat research conference that was held in
Leningrad (now Saint Petersburg) in 1974 and
where the commission on bats was organized
under the auspices of the Soviet Theriological
Society. The conference had become a regular
event with a number of attendees from the
former USSR who reported new findings in a
compendium of proceedings (1974, 1980, 1982,
1988, 1990). Altogether, more than 120 papers
and short communications on the distribution of
bats in the territory of the former USSR had
been published by 1990 (Strelkov, 1990); a
great deal of them containing data collected in
caves and underground sites.
After the dissolution of the Soviet Union, the
coordination of bat studies in Russia was taken
over by the Russian Bat Research Group that was
established at Seventh All-Russian Bat Research
Conference (13–16 April 1999). The Group issues its own periodical, Plecotus et al., and
maintains a website at http://zmmu.msu.ru/bats
Apart from papers and proceedings, several
comprehensive regional bat surveys have recently been conducted and subsequently published as dissertations and monographs. With
respect to the cave bat fauna, it is worth mentioning a study on bats of the Western Caucasus
(Gazaryan, 2002) and those for the central and
southern Ural regions (Orlov, 2000; Snit’ko,
2004). The results of the latter two studies
largely constitute the monograph “Bats of the
Urals” (Bol’shakov et al., 2005) that was later
amended by several new publications of
Vladimir Snit’ko, who explored summer and
winter bat assemblages in the caves of Bashkiria
and the Orenburg and Chelyabinsk regions. In
1999, S. Gazaryan and A. Ostapenko published
the first list of 49 caves which are important for
bats in the Russian Caucasus and should be
legally and physically protected; the paper included recommendations to cavers concerning
bat-friendly conduct and the timing of visits to
these caves. In a series of further publications S.
Gazaryan appended this list with many new
important sites which he had discovered during
his field trips in 1999–2012.
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S.I. Golovatch et al.
Since 1975, several groups have become
particularly well studied in the Russian Caucasus, such as cave diplopods (Golovatch, 1975,
1978, 1983, 1984–1985, 1990, 2011; Golovatch, Enghoff, 1990; Golovatch, Chumachenko, 2013; Golovatch et al., 2016; Antić, Makarov,
2016; Antić et al., 2018); cave carabids (Belousov, 1989, 1999, 1998; Belousov, Zamotajlov, 1999, 1997; Belousov, Koval, 2009, 2011)
and a few other higher taxa, mostly with descriptions of numerous new species.
A highly detailed survey of the invertebrate
cave fauna of the former Soviet Union, including Russia, and a complete bibliography was
published recently by Turbanov et al. (2016a, b,
c). The present paper is not only restricted to the
European part of Russia, including the northern
Caucasus, but excluding Crimea (the latter is
to be treated separately), but it incorporates all
previously unpublished data contained in the
PhD Thesis of S.I. Ljovuschkin (1966). It is also
slightly updated using relevant information, both
new and missed old, but in addition it covers
bats as a highly important component of cave
fauna. Besides this, a geological background is
provided here.
No zoogeographical analysis is undertaken
here since such is available and still quite valid
as regards the entire subterranean fauna of the
former USSR (Kniss, 2004). In addition, European Russia, especially the northern Caucasus
with its particularly rich endogean fauna, is an
artificial and heterogeneous region from a biogeographical point of view.
III. Karst and caves
The Precambrian craton occupies most of
the European part of Russia. Karst is mainly
developed in intrastratal settings in limestones,
dolomites, chalk, gypsum and salt of different
ages. Most of the caves are in gypsum.
The karstic caves were developed in rocks
from a wide range of geological age: Archaean,
Proterozoic, Palaeozoic, Triassic, Jurassic, in
limestones, dolomites, mixted carbonate rocks,
carbonate conglomerates, and gypsum.
Pseudokarstic caves are known in sandstones.
Fig. 1. Karsts of European Russia. A — Carbonated
rocks. B — Carbonated and Gypsum rocks. Numbers:
1 — Valdai-Kuloisk: limestone and gypsum karsts;
2 — Volga-Kama; 3 — Timan Plateau; 4 — Urals;
5 — Great and Lesser Caucasus. * Pinega gypsum
caves. (After Pulina (2005), modified; drawing by B.
Juberthie and G. Nazareanu).
Рис. 1. Карсты европейской части России. А —
карбонатные породы. B — карбонатные и гипсовые породы. Номера: 1 — Валдай-Кулой:
известняковые и гипсовые карсты; 2 — ВолгоКамские; 3 — Тиманское плато; 4 — Урал; 5 —
Большой и Малай Кавказ. * Пинежские гипсовые
пещеры. (По Pulina (2005) c изменениями; рисунок B. Juberthie и G. Nazareanu).
A. Chikishev (1966, 1973, 1978, 1984) distinguished 12 speleological areas in the former
USSR. To date, based on morpho-structural
criteria, we distinguish 5 karstic-speleologic
regions in the European part of Russia: ValdaiKuloisk, Volga-Kama, Timan Plateau, Russian
Great Caucasus, Ural (Fig. 1).
Subterranean biota of European Russia
159
More than 360 gypsum caves are known,
1. The Valdai-Kuloisk speleological
their total length is over 100 km. The longest
region. The northern part of the Russian caves occur in gypsum karst and include the
Kulogorskaya-1-2-Troya (14 km), KonstitutPlain
Located in the northwestern part of European Russia, between Moscow and St. Petersburg,
this region is characterized by a very large, but
fragmented limestone and gypsum karst strip,
50–100 km broad, 1000 km from south to north
near the Baltic Sea.
These northern karsts of European Russia
are subjected to hard climate below the arctic
anticyclone, with oceanic traits. The rugged
karst landscape is mostly overgrown with taiga
forest of birch and spruce.
The northwestern Russian area includes some
of the largest gypsum karst regions in Europe.
The gypsum-anhydrate sequence, often intercalated with limestones and dolomites, is usually
40–60 m thick and lies at shallow depths. Near
40% of the caves are active, partly phreatic, air
and water temperature often being near 0 °C.
sionaya (5.8 km) and Olimpiyskaya (5.5 km)
caves (Malkov, Gurkalo, 1999; Klimchouk,
2003b).
They include Pinega gypsum caves in a
Permian gypsum outcrop west of the Urals,
around the small town of Pinega, east of Arkhangelsk, just outside the Arctic zone of permafrost, in the main bed of evaporites (gypsum and
anhydrite). The ground never freezes to depths
of more than a few meters, and the cave streams
continue flowing throughout the winter. Over
50 km of passages have been mapped in the
group of Pinega caves, among them 22 caves are
longer than 1 km each. Three of the most important caves are the Olimpiyskaya, Lomonosovskaya and Muzeynaya (= Muzeyskaya) caves,
in the Iron Gates Sanctuary and the Pinega
(=Pinezhskiy) Nature Reserve (Fig. 2) (Waltham, 1994, 2003; Waltham, Cooper, 1998).
Fig. 2. Pinega Gypsum caves. Maps of three caves located at the Iron Gates (near the Pinezhskiy Nature
Reserve). After surveys by “Arkhangelsk Geologiya” and Waltham (2003); coloured by B. Juberthie.
Рис. 2. Пинежские гипсовые пешеры. Карта трех пещер, расположенных у Железных Ворот (рядом
с Пинежским заповедником). По обследованиям «Архангельской геологии» и Waltham (2003);
раскрасил B. Juberthie).
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S.I. Golovatch et al.
Many caves are truncated in the valley walls.
A large proportion of caves have dendritic passages that carry or carried subterranean streams.
The Simfoniya (3.2 km long) and Golubinskaya (1.6 km long) caves are both maze caves
without stream, located at higher levels in the
gypsum plateau. They are comparable in appearance to those in the gypsum of Ukraine.
Below some large entrances cave glaciers
are created; the oldest ice has been dated to
3000 years ago (Andrejchuk, Klimchouk, 1996;
Malkov et al., 1986, 1988).
2. Volga-Kama (the Volgo-Kamsky
region)
This region lies between the Volga and the
Urals. Karst is developed in Permian limestones, dolomites and sulfates, and is in Carboniferous and Devonian carbonate rocks.
In the middle Volga and Kama basins, four
relatively large zones of limestone and gypsum
karst are recorded. They consist in gypsum
beds, thickness 10–45 m, interstrated to limestone and dolomite of lower Permian age. There
are springs, commonly discharging sulfate-rich
water from deep-seated or semi-confined aquifers (Stupishin, 1965; Andrejchuk, Klimchouk,
1996).
The caves are not numerous, their length
modest (as far as 100 m), and they still remain
poorly-studied in a biospeological aspect (Klimchouk, 2003b).
3. The Timanskiy Mountain Ridge
Within the taiga, or the boreal forest belt,
from the northern Urals to the Arctic Ocean, lies
the Timanskiy Mountain Ridge which contains Devonian and lower Permian gypsum karsts
intercalated in dolomites and limestones (Torsuev, 1975). Boreholes and mines have intercepted numerous cavities yielding sulfate-rich
water.
4. The Urals (Fig. 3)
Karst, gypsum karst, and caves were explored, described and studied by Andrejchuk
(1996), Andrejchuk and Klimchouk (1996),
Fig. 3. Karsts of the Urals: carbonated and sulfated
rocks.
After Salomon and Pulina (2005), modified by G.
Nazareanu.
Рис. 3. Карсты Урала: карбонатные и сульфатные
породы.
По Salomon and Pulina (2005), изменения G. Nazareanu.
Subterranean biota of European Russia
Chikishev (1973), Dublyanskiy and Ilyukhin
(1982), Gorbunova (1965, 1977, 1979), Gorbunova et al. (1992); Kiselyov and Klimchouk
(1991), Maksimovitch (1963), Maksimovitch
and Gorbunova (1958), Maksimovitch and Kostarev (1973) etc.
The Ural Mountain Ridge covers 2000 km
from north to south and represents the traditional boundary between Europe and Asia. It is a
Hercynian folded and eroded system largely
covered by woodlands. Approximately 1,800
caves are recorded in gypsum and anhydrite
rocks of Permian age, as well as in Devonian and
Carboniferous limestones and dolomites.
The largest caves are located in the middle
and southern Urals: the Sumgan-Kutuk (9.9
km), Divya (9.1 km) and Kinderlinskaya (7.9
km) caves, as well as the Kungurskaya Cave
(5.4 km) show in the Perm Region. The Shulgan-Tash Cave, 2.6 km of passages, is famous
for its Palaeolithic paintings. The most important karst springs are located in the western
Urals: the Goluboe Ozero springs and the Krasnyi Klyuch Spring.
In the northern Urals, near the Polar Circle
(Paykhoy and arctic regions), karst was developed under and inside the permafrost (Klimchouk, 2003b).
Gypsum karst in the Urals is widespread,
mainly present along the Urals’ western periphery (Maksimovitch, Gorbunova, 1958; Gorbunova, 1965, 1977, 1979; Maksimovitch, Kostarev,
1973; Lysenin, 1981; Makukhin, Molodkin,
1988; Gorbunova et al., 1992; Andrejchuk,
1996).
These extend from the Arctic islands of
Novaya Zemlya in the North as far as the hot
deserts of the northern Caspian region in the
South.
In the central Urals, several hundred publications concern gypsum karsts. About 200 caves
have been explored in the gypsum: the largest
are the Kungurskaya (= Kungur Ice Cave) (5.6
km) and the Zuyatskaya (1,410 m long) caves.
Karstic springs are numerous.
Studies on cave fauna were mainly performed in caves of the western macro-slope of
the central Urals.
161
5. The Russian Great Caucasus
The Great Caucasus, or Caucasus Major
(Bolshoy Kavkaz), is the highest mountain ridge
in European Russia, where many mountain peaks
are 3,000–5,000 m in elevation (Mt Elbrus,
volcanic peak, 6,642 m a.s.l.), spanning 1,250
km from the Black Sea to the Caspian Sea.
Only the northern macro-slope (Ciscaucasia, or Circassia) and the westernmost part of the
Caucasus, from Novorossiysk as far as Sochi
and the Psou River, neighbouring to Abkhazia,
along the Black Sea coast, is Russian.
On the northern macro-slope, the karsts were
developed in monoclinal and folded structures,
mostly in Jurassic and lower Cretaceous limestones, dolomites, and gypsum. Alpine karst
and deep caves are present in the northern slope
and Russian western region. As of 2003, the
Gorlo Barloga Cave was the deepest (–900 m),
located in the Zagedan Massif in Devonian and
Carboniferous metamorphosed limestones and
schists. Among other caves, there are the
Zagedanskaya (–570 m, 5,300 m long) and
Alekseeva (–465 m, 4,410 long).
In the Dzhentu Massif, the Mayskaya Cave
is –500 deep and 4,410 m long.
In the alpine Fisht Massif at the western
flank of the core zone and the westernmost
summit of the Caucasus Major Range, explored
from 1980 to 1996, deep caves in Upper Jurassic limestones are present: the Krestik-Turist (633 m, 14 km long), Paryashchaya Ptitsa (-595
m, 4,500 m long) and Ol’ga (–520 m, 3,500 m
long) caves; most springs are located at the
range bottom 1,200 m a.s.l. (Reisner, Shelepin
1997; Klimchouk 2003a).
Numerous zones of gypsum karst are present
along the northern macro-slope of the Great
Caucasus. Many caves support subterranean
streams. The longest are the Popova (1,670 m),
Ammonal’naya (1,460 m) and Seteney (980 m)
caves (Kazanbiev, 1975; Klimchouk et al., 1996;
Makukhin, Molodkin, 1988; Ostapenko, 1993,
1994; Sukhovey, 1992).
On the northern macro-slope of the Caucasus Major, the most important biospeleological
areas are located along the Black Sea in the
Krasnodar Province (the Alek, Akhtsu, Voron-
162
S.I. Golovatch et al.
tsovky, Dzykhra, Akhshtyr, Akhun and FishtOshten-Lagonaki massifs of Jurassic dolomites).
As of 2003, the longest and deepest caves are
the Vorontsovkaya (10.6 km) and RucheinayaShkol’naya-Zabludshikh (–601 m) (Gvozdetski, 1965a, 1965b).
Cave fauna is relatively well known.
IV. Subterranean biota
Birstein and Ljovuschkin (1950) divided
Russia into two major biospeleological zones:
1. A northern zone covering most of the
Russian territory: the Russian Plain and the
northern Urals, both characterized by very poor
subterranean faunas, with only a few stygobionts belonging to copepods, Bathynella syncarids and amphipods that correspond to old
widespread groups.
2. A southern zone (Ciscaucasia and western
Transcaucasia), characterized by numerous endemic stygo- and troglobionts.
About 100 exclusively subterranean taxa
have been recorded from European Russia, more
than 50% being aquatic. Most of them inhabit
the Russian Great Caucasus.
Based on actual knowledge, there are three
biospeological zones: central Russia, the Urals
and the Russian Great Caucasus.
IV. 1. Valdai-Kuloisk. The northern
part of the Russian Plain
Mostly microorganisms are surveyed in gypsum caves of the Belomorsko-Kuloiskoe Plateau: the Golubinskaya, Pekhorovskiy Proval
and G-1 caves, all by Semikolennykh (1997).
These gypsum caves are embedded 3–15 m
deep under the surface and are very wet, with
high-level flood waters annually. The mean
temperature is about 2–8 °C in summer and
nearly 0 °C in winter.
The following species have been identified,
none of which represents troglo– or stygobionts:
Domain Bacteria Woese, Kandler et Wheelis,
1990
Division Actinobacteria Goodfellow, 2012
Order Micromonosporales Genilloud, 2015
Family Micromonosporaceae Krassilnikow,
1938 em. Zhi, Li et Stackebrandt, 2009
– Micromonospora sp., from mineral clay
samples;
Order Actinomycetales Buchanan, 1917
Family Streptomycetaceae Waksman et Henrici, 1943
– Streptomyces sp., from mineral clay samples;
Unidentified sulfate-reducing bacteria were
also discovered.
Domain Eukaryota (Chatton, 1925) Whittaker
et Margulis, 1978
Kingdom Fungi Linnaeus, 1753 stat. nov. Necker, 1783 em. Cavalier-Smith, 1998
Division Ascomycota Berkeley, 1857 stat. nov.
Cavalier-Smith, 1987
Class Eurotiomycetes O.E. Eriksson et Winka,
1997
Order Eurotiales G.W.Martin ex Benny et
Kimbrough, (1980)
Family Trichocomaceae E. Fischer, (1897)
– Penicillium italicum (Wehmer, 1894), on
dead insects;
– Penicillium viridicatum Westling, 1911,
with stable colonies on cave clay;
Class Dothideomycetes O.E. Eriksson et Winka, 1997
Order Pleosporales Luttrell ex M.E. Barr,
(1987)
Family: Incertae sedis
– Pyrenochaeta sp.;
Division Zygomycota Barr, 1982
Order Mucorales Fries, 1832
Family Mucoraceae Dumortier, (1822)
– Rhizopus sp., on organic matter (flooded
or penetrated through infiltration);
Kingdom Viridiplantae Cavalier-Smith, 1981
Division Chlorophyta Reichenbach, 1828, em.
Pascher, 1914, em. Lewis et McCourt, 2004
Order Chlorococcales Pascher, 1915
Family Chlorococcaceae Blackman et Tansley, 1902
– Chlorococcus sp., forming green spots
near the entrance, always situated in reflected light.
Kingdom Protozoa R. Owen, 1858
Subkingdom Sarcomastigota Cavalier-Smith,
1983
Subterranean biota of European Russia
Group Excavata (Cavalier-Smith), 2002
Phylum Amoebozoa Lühe, 1913
Class Tubulinea Smirnov, Nassonova, Berney,
Fahrni, Bolivar et Pawlowski, 2005
Order Arcellinida Kent, 1880
Family Centropyxidae Jung, 1942
– Centropyxis aerophila Deflandre, 1929.
Found in a cave at the bank of Pechora
River within the Pechoro-Ilychskiy Nature
Reserve (Troitsk-Pechorsky District, Komi
Republic) (Mazei et al., 2012).
Family Phryganellidae Jung, 1942
– Phryganella hemisphaerica Penard, 1902.
Found in a cave at the bank of Pechora
River within the Pechoro-Ilychskiy Nature
Reserve (Troitsk-Pechorsky District, Komi
Republic) (Mazei et al., 2012).
Kingdom Animalia Linnaeus, 1758
Phylum Nematoda Diesing, 1861
Nematoda indet. Unidentified nematodes
have been recorded in the caves Olimpiyskaya and Muzeynaya, Pinezhskiy Nature
Reserve, Arkhangelsk Region (Chertoprud
et al., 2011).
Phylum Annelida Lamarck, 1809
Class Oligochaeta Grube, 1850
Order Enchytraeida Vejdovský, 1879
Family Enchytraeidae Vejdovský, 1879
Enchytraeidae indet. Unidentified enchytraeids have been recorded in the Olimpiyskaya Cave, Pinezhskiy Nature Reserve,
Arkhangelsk Region (Chertoprud et al.,
2011).
Phylum Arthropoda Siebold, 1848
Subphylum Crustacea Brünnich, 1772
Class Copepoda Milne-Edwards, 1840
Order Cyclopoida Burmeister, 1834
Family Cyclopidae Rafinesque, 1815
– Megacyclops viridis (Jurine, 1820). Recorded in waterbodies inside the Muzeynaya Cave, Pinezhskiy Nature Reserve,
Arkhangelsk Region (Chertoprud et al.,
2011).
Class Malacostraca Latreille, 1817
Order Amphipoda Latreille, 1816
Family Pallaseidae Tachteew, 2000
– Pallasea sp. Recorded in waterbodies
inside the Kitezh Cave, Pinezhskiy Nature
163
Reserve, Arkhangelsk Region (Sidorov et
al., 2011; Chertoprud et al., 2011).
Class Arachnida Cuvier, 1812
Subclass Acari Leach, 1817
The fauna of free-living gamasid mites of
the Pinezhskiy Nature Reserve is unusually
diverse and rich (117 species) (Makarova,
2009), but its caves support only few presumed trogloxenes. Apparently, the caves
are too cold to sustain a characteristic terrestrial fauna.
Superclass Hexapoda Blainville, 1816
Class Collembola Lubbock, 1870
The springtail fauna of the Pinezhskiy Nature Reserve is unusually diverse and rich
(126 species) (Babenko, 2008), but its caves
support only few presumed trogloxenes.
Apparently, the caves are too cold to sustain
a characteristic terrestrial fauna.
Class Insecta Linnaeus, 1758
Order Ephemeroptera Hyatt et Arms, 1891
Family Baetidae Leach, 1815
– Baetis rhodani (Pictet, 1843). Recorded
in waterbodies inside the Olimpiyskaya
Cave, Pinezhskiy Nature Reserve, Arkhangelsk Region (Chertoprud et al., 2011).
– Centroptilum luteolum Müller, 1776. Recorded in waterbodies inside the Olimpiyskaya and Muzeynaya caves, Pinezhskiy Nature Reserve, Arkhangelsk Region
(Chertoprud et al., 2011).
Family Siphlonuridae Ulmer, 1920
– Siphlonurus lacustris Eaton, 1870. Recorded in waterbodies inside the Muzeynaya
Cave, Pinezhskiy Nature Reserve, Arkhangelsk Region (Chertoprud et al., 2011).
Order Plecoptera Burmeister, 1839
Family Nemouridae Newman, 1853
– Nemurella pictetii (Klapálek, 1900). Recorded in waterbodies inside the
Olimpiyskaya Cave, Pinezhskiy Nature
Reserve, Arkhangelsk Region (Chertoprud
et al., 2011).
– Nemoura sahlbergi Morton, 1896. Recorded in waterbodies inside the Olimpiyskaya Cave, Pinezhskiy Nature Reserve,
Arkhangelsk Region (Chertoprud et al.,
2011).
164
S.I. Golovatch et al.
– Nemoura aff. avicularis Morton, 1894.
Recorded in waterbodies inside the Muzeynaya Cave, Pinezhskiy Nature Reserve,
Arkhangelsk Region (Chertoprud et al., 2011).
Family Perlodidae Klapálek, 1912
– Diura nanseni (Kempny, 1900). Recorded in waterbodies inside the Olimpiyskaya
Cave, Pinezhskiy Nature Reserve, Arkhangelsk Region (Chertoprud et al., 2011).
Family Taeniopterygidae Klapálek, 1905
– Taeniopteryx nebulosa (Linnaeus, 1758).
Recorded in waterbodies inside the Olimpiyskaya and Muzeynaya caves, Pinezhskiy Nature Reserve, Arkhangelsk Region
(Chertoprud et al., 2011).
Order Hemiptera Linnaeus, 1758
Family Corixidae Leach, 1815
– Sigara semistriata (Fieber, 1848). Recorded in waterbodies inside the Olimpiyskaya Cave, Pinezhskiy Nature Reserve,
Arkhangelsk Region (Chertoprud et al.,
2011).
Order Diptera Linnaeus, 1758
Suborder Nematocera Schiner, 1862
Family Limoniidae Speiser, 1909
– Eloeophila mundata (Loew, 1871). Recorded in waterbodies inside the Olimpiyskaya Cave, Pinezhskiy Nature Reserve,
Arkhangelsk Region (Chertoprud et al.,
2011).
Table 1. List of *stygobionts, stygophile and troglophile species from the northern part
of the Russian Plain.
Таблица 1. Список *стигобионтов, стигофилов и троглофилов из северной части
Русской равнины.
A — Stygobionts*: stygophiles
BACTERIA
Micromonosporaceae
Micromonospora sp.
Streptomycetaceae
Streptomyces sp.
FUNGI
Trichocomaceae
Penicillium italicum (Wehmer, 1894);
Penicillium viridicatum Westling, 1911
Pleosporales
Pyrenochaeta sp.
Mucoraceae
Rhizopus sp.
VIRIDIPLANTAE
Chlorococcaceae
Chlorococcus sp.
PROTOZOA
Centropyxidae
Centropyxis aerophila Deflandre, 1929
Phryganellidae
Phryganella hemisphaerica Penard, 1902
NEMATODA
Nematoda indet.
OLIGOCHAETA
Enchytraeidae indet.
COPEPODA
Cyclopidae
Megacyclops viridis (Jurine, 1820)
Pallaseidae
Pallasea sp.
INSECTA
EPHEMEROPTERA
Siphlonuridae
Siphlonurus lacustris Eaton, 1870
Baetidae
Centroptilum luteolum Müller, 1776;
Baetis rhodani (Pictet, 1843)
PLECOPTERA
Perlodidae
Diura nanseni (Kempny, 1900)
Taeniopterygidae
Taeniopteryx nebulosa (Linnaeus, 1758)
Nemouridae
Nemurella pictetii (Klapálek, 1900);
Nemoura sahlbergi Morton, 1896;
Nemoura af. avicularis Morton, 1894
HETEROPTERA
Corixidae
Sigara semistriata (Fieber, 1848)
DIPTERA
Pediciidae
Dicranota bimaculata (Schummel, 1829)
Limoniidae
Eloeophila mundata (Loew, 1871)
Simuliidae
Cnetha sp.
Chironomidae
Guttipelopia guttipennis (Wulp, 1874)
CHIROPTERA
Vespertilionidae
Myotis brandtii (Eversmann, 1845)
Eptesicus nilssonii (Keyserling et Blasius, 1839)
Subterranean biota of European Russia
Family Pediciidae Osten-Sacken, 1860
– Dicranota bimaculata (Schummel, 1829).
Recorded in waterbodies inside the Olimpiyskaya Cave, Pinezhskiy Nature Reserve,
Arkhangelsk Region (Chertoprud et al.,
2011).
Family Simuliidae Newman, 1834
–Cnetha sp. Recorded in waterbodies inside the Olimpiyskaya Cave, Pinezhskiy
Nature Reserve, Arkhangelsk Region (Chertoprud et al., 2011).
Family Chironomidae Jacobs, 1900
– Guttipelopia guttipennis (Wulp, 1874).
Recorded in waterbodies inside the Muzeynaya Cave, Pinezhskiy Nature Reserve,
Arkhangelsk Region (Chertoprud et al.,
2011).
Phylum Chordata Haeckel, 1874
Subphylum Vertebrata J.-B. Lamarck, 1801
Class Mammalia Linnaeus, 1758
Order Chiroptera Blumenbach, 1779
Famaly Vespertilionidae Gray, 1821
– Myotis brandtii (Eversmann, 1845). Solitarily hibernating individuals of this species were reported from the Golubinskiy
Proval Cave, Pinezhskiy Nature Reserve,
where they were found twice in 1980 (Rykov, 2014).
– Eptesicus nilssonii (Keyserling et Blasius, 1839). Hibernating Northern bats were
reported from several karst caves of the
Belomorsko-Kuloiskoe Plateau, with the
maximum number of 16 bats observed in
one visit to the Golubinskiy Proval Cave
(Rykov, 2014).
IV. 2. Central Russia
The cave biota of central Russia is littleknown and poorly studied. Below is only a
preliminary evaluation.
This karstic region lies too northerly to be
subjected to either Mediterranean or Balkan, or
Asiatic influence.
Bacteria, Algae and Fungi were surveyed in
the Poneretka Cave, Borovichskiy District,
Novgorod Region, located in limestones of the
Valdai Plateau, temperature 9 °C, by Semikolennykh (1997). The numbers of microorgan-
165
isms are about 107 per sediment gram.
Domain Bacteria Woese, Kandler et Wheelis,
1990
Division Actinobacteria Goodfellow, 2012
Order Actinomycetales Buchanan, 1917
Family Mycobacteriaceae Chester, 1897
– Mycobacterium hyalinum Söhngen, 1913;
– M. licheniforme Krassilnikow, 1949;
– M. luteum Söhngen, 1913;
– M. mucosum Krassilnikow, 1941
Family Propionibacteriaceae Delwiche, 1957
– Propionibacterium sp.
Family Streptomycetaceae Waksman et Henrici, 1943
– Streptomyces sp.;
– Streptoverticillium sp.;
– Streptosporangium sp.
Family Micrococcaceae Pribram, 1929
– Arthrobacter globiformis Conn et Dimmick, 1947
Class Bacilli Ludwig, Schleifer et Whitman,
2010
Order Bacillales Prévot, 1953
Family Bacillaceae Fischer, 1895
– Bacillus mycoides Flügge, 1886
Division Cyanobacteria Stanier, 1973
Order Nostocales Cavalier-Smith, 2002
Family Nostocaceae C.A. Agardh, 1824 ex
Kirchner, 1898
– Nostoc sp.
Division Proteobacteria Stackebrandt, Murray
et Trüper, 1988, Garrity, Bell et Lilburn, 2005
Class Gammaproteobacteria Garrity, Bell et
Lilburn, 2005
Order Pseudomonadales Orla-Jensen, 1921
Family Pseudomonadaceae Winslow, Broadhurst, Buchanan, Krumwiede, Rogers et
Smith, 1917
– Pseudomonas desmolyticum Cray et
Thornton, 1928
Kingdom Fungi Linnaeus, 1753 stat. nov. Necker, 1783 em. Cavalier-Smith, 1998
Division Ascomycota Berkeley, 1857 stat. nov.
Cavalier-Smith, 1987
Class Eurotiomycetes O.E. Eriksson et Winka,
1997
Order Onygenales Ciferri ex Benny et Kimbrough, (1980)
166
S.I. Golovatch et al.
Family Gymnoascaceae Baranetsky, (1872)
– Pseudoarachniotus sp.
Kingdom Viridiplantae Cavalier-Smith, 1981
Division Chlorophyta Reichenbach, 1828, em.
Pascher, 1914, em. Lewis et McCourt, 2004
Class Trebouxiophyceae Friedl, 1995
Chlorellales sp. is the only species collected.
Protists are extremely poorly known in caves
of the central part of European Russia. Only two
papers deal with this group in the region in
question. One is by Semikolennykh (1997),
dedicated to the study of Protista in the Poneretka Cave, Borovichskiy District, Novgorod Region. The other is by Mazei et al. (2012), which
focuses on microfauna in artificial adits of the
Zhiguli Hills, Stavropolskiy District, Samara
Region.
Kingdom Chromista Cavalier-Smith, 1981
Phylum Ciliophora Doflein, 1901
Class Oligohymenophorea de Puytorac et al.,
1974
Order Pleuronematida Fauré-Fremiet in Corliss, 1956
Family Pleuronematidae Kent, 1881
– Balantiophorus elongatus Schewiakoff,
1892. Recorded from the Poneretka Cave,
Borovichskiy District, Novgorod Region
(Semikolennykh, 1997).
Kingdom Protozoa R. Owen, 1858
Subkingdom Sarcomastigota Cavalier-Smith,
1983
Group Excavata (Cavalier-Smith), 2002
Phylum Euglenozoa Cavalier-Smith, 1981
Class Euglenophyceae Schoenichen, 1925
Order Peranemida Bütschli, 1884
Family Bodonaceae Bütschli, 1884
– Bodo globosus Stein, 1878. Recorded
from the Poneretka Cave, Borovichskiy
District, Novgorod Region (Semikolennykh,
1997).
– B. lens (O.F. Müller) Klebs (1892). Recorded from the Poneretka Cave, Borovichskiy District, Novgorod Region (Semikolennykh, 1997).
– Bodo sp. Recorded from the Poneretka
Cave, Borovichskiy District, Novgorod
Region (Semikolennykh, 1997).
Phylum Amoebozoa Lühe, 1913
Class Tubulinea Smirnov, Nassonova, Berney,
Fahrni, Bolivar et Pawlowski, 2005
Order Arcellinida Kent, 1880
Family Arcellidae Ehrenberg, 1843
– Arcella arenaria compressa Chardez,
1974. Recorded from artificial adits of the
Zhiguli Hills, Stavropolskiy District, Samara Region (Mazei et al., 2012).
Family Centropyxidae Jung, 1942
– Centropyxis aerophila Deflandre, 1929.
Recorded from artificial adits of the Zhiguli
Hills, Stavropolskiy District, Samara Region (Mazei et al., 2012).
– C. aerophila sphagnicola Deflandre,
1929. Recorded from artificial adits of the
Zhiguli Hills, Stavropolskiy District, Samara Region (Mazei et al., 2012).
– C. plagiostoma Thomas et Bonnet, 1955.
Recorded from the Poneretka Cave, Borovichskiy District, Novgorod Region (Semikolennykh, 1997).
Family Hyalospheniidae Schultze, 1877
– Hyalosphenia papilio Leidy, 1879. Recorded from artificial adits of the Zhiguli
Hills, Stavropolskiy District, Samara Region (Mazei et al., 2012).
Family Nebelidae Taranek, 1882
– Nebela parvula Cash, 1909. Recorded
from artificial adits of the Zhiguli Hills,
Stavropolskiy District, Samara Region
(Mazei et al., 2012).
– N. tincta Leidy, 1879. Recorded from
artificial adits of the Zhiguli Hills, Stavropolskiy District, Samara Region (Mazei et al.,
2012).
Family Phryganellidae Jung, 1942
– Phryganella acropodia (Hertwig et Lesser, 1874). Recorded from artificial adits of
the Zhiguli Hills, Stavropolskiy District,
Samara Region (Mazei et al., 2012).
– P. hemisphaerica Penard, 1902. Recorded from artificial adits of the Zhiguli Hills,
Stavropolskiy District, Samara Region
(Mazei et al., 2012).
Phylum Amoebozoa Lühe, 1913 em. CavalierSmith, 1998
Class Archamoebea Cavalier-Smith, 1998
Family Mastigamoebidae Goldschmidt, 1907
– Mastigamoeba sp. Recorded from the
Poneretka Cave, Borovichskiy District,
Novgorod Region (Semikolennykh, 1997).
Subterranean biota of European Russia
Class Tubulinea Smirnov, Nassonova, Berney,
Fahrni, Bolivar et Pawlowski, 2005
Order Tubulinida Smirnov, Nassonova, Berney, Fahrni, Bolivar et Pawlowski, 2005
Family Hartmannellidae Volkonsky, 1931
– Hartmannella sp. Recorded from the Poneretka Cave, Borovichskiy District, Novgorod Region (Semikolennykh, 1997).
Kingdom Rhizaria Cavalier-Smith, 2002
Phylum Cercozoa Cavalier-Smith, 1998
Class Imbricatea Cavalier-Smith, 2003
Order Euglyphida Copeland, 1956
Family Euglyphidae Wallich, 1864
– Assulina seminulum (Ehrenberg, 1848).
Recorded from artificial adits of the Zhiguli
Hills, Stavropolskiy District, Samara Region (Mazei et al., 2012).
– Euglypha rotunda Wailes, 1915. Recorded from artificial adits of the Zhiguli Hills,
Stavropolskiy District, Samara Region
(Mazei et al., 2012).
Family Trinematidae Hoogenraad et de Groot,
1940
– Trinema complanatum Penard, 1890. Recorded from artificial adits of the Zhiguli
Hills, Stavropolskiy District, Samara Region (Mazei et al., 2012).
– T. enchelys (Ehrenberg, 1838). Recorded
from artificial adits of the Zhiguli Hills,
Stavropolskiy District, Samara Region
(Mazei et al., 2012).
Kingdom Animalia Linnaeus, 1758
Phylum Rotifera Cuvier, 1817
Phylum Annelida Lamarck, 1809
Class Oligochaeta Grube, 1850
Order Crassiclitellata Jamieson, 1988
Family Lumbricidae Claus, 1876
– Eisenia foetida (Savigny, 1826), first recorded by Michaelsen in 1926 (Malevich,
1947) from moist clay in the deep part of a
cave on the right bank of Volga River,
downstream the mouth of Kama River. It
was found again by Birstein in several caves
of the Russian Caucasus and in Abkhazia.
The species is trogloxene in Western Europe and abundant in farmyard manure. It is
perhaps troglophilic in Russian and Georgian caves with bat guano.
167
– Eiseniella tetraedra (Savigny, 1826). Recorded from wells in the city of Saratov
(Behning, 1928).
Order Enchytraeida Vejdovský, 1879
Family Enchytraeidae Vejdovský, 1879
– Henlea ventriculosa (d’Udekem, 1854).
Recorded from wells in the city of Saratov
(Behning, 1928).
– Enchytraeus albidus Henle, 1837. Recorded from wells in the city of Saratov
(Behning, 1928).
– Fridericia bulbosa (Rosa, 1887). Recorded from wells in the city of Saratov
(Behning, 1928).
Order Tubificida Brinkhurst, 1982
Family Naididae Ehrenberg, 1828
– Nais communis Piguet, 1906. Repeatedly
recorded from numerous wells in European
Russia, including those in the city of Saratov (Behning, 1928).
– N. pseudobtusa Piguet, 1906. Recorded
from wells in the city of Saratov (Behning,
1928).
Family Tubificidae Vejdovský, 1884
– Potamothrix hammoniensis (Michaelsen,
1901). Recorded from wells in the city of
Saratov (Behning, 1928).
Phylum Nematoda Diesing, 1861
Nematodes have almost never been studied
in caves of the central part of European Russia.
The only relevant paper is that by Behning
(1928) which puts on record 11 species of the
group found in wells in the city of Saratov. This
city is located on the Volga Upland within the
Saratov Region:
Class Enoplea Inglis, 1983
Order Monhysterida Filipjev, 1929
Family Dorylaimidae De Man, 1876
– Eudorylaimus carteri (Bastian, 1865);
– E. carteri rotundatus (Micoletzky, 1922);
– E. monohystera (De Man, 1880);
– E. obtusicaudatus (Bastian, 1865);
– Laimydorus pseudostagnalis (Micoletzky, 1927).
Family Monhysteridae De Man, 1876
– Monhystera paludicola De Man, 1881.
Order Mononchida Jairajpuri, 1969
Family Mononchidae Chitwood et Chitwood,
1937
– Mononchus macrostoma Bastian, 1865;
– M. papillatus Bastian, 1865.
168
S.I. Golovatch et al.
Order Triplonchida Cobb, 1920
Family Tobrilidae Filipjev, 1918
– Tobrilus allophysoides (Micoletzky,
1925).
Class Chromadorea Inglis, 1932
Family Rhabditidae Örley, 1880
– Rhabditis curvicaudata (Schneider, 1866).
Class Secernentea Lorenzen, 1981
Order Tylenchida Thorne, 1949
Family Cephalobidae Filipjev, 1934
– Cephalobus rigidus Schneider, 1866.
Phylum Arthropoda Siebold, 1848
Subphylum Crustacea Brünnich, 1772
Class Copepoda Milne-Edwards, 1840
Order Cyclopoida Burmeister, 1834
– Diacyclops bicuspidatus (Claus, 1857).
Stygoxene, recorded from the Bornukovskaya Cave in the Nizhniy Novgorod
Region (Birstein, Ljovuschkin, 1966).
– Acanthocyclops vernalis (Fischer, 1853).
Stygophile, recorded from the Bornukovskaya Cave in the Nizhniy Novgorod
Region (Birstein, Ljovuschkin, 1966).
Order Harpacticoida G.O. Sars, 1903
Family Canthocamptidae Brady, 1880
– Canthocamptus staphylinus (Jurine,
1820). Stygoxene, recorded from wells near
the city of Saratov (Behning, 1928).
Family Parastenocarididae Chappuis, 1940
– Parastenocaris fonticola Borutzky, 1926,
decribed from a well at Kosino near Moscow City (Borutzky, 1926), currently known
to be a widespread species (Borutzky, 1926).
Class Ostracoda Latreille, 1802
Order Podocopida G.O. Sars, 1866
Family Cyprididae Baird, 1845
– Potamocypris wolfi Brech, 1920. Crenobiont, found in underground streams near
the city of Voronezh below Tertiary sandstone plates (Bronstein, 1947).
Class Malacostraca Latreille, 1817
Superorder Syncarida Packard, 1885
Order Bathynellacea Chappuis, 1915
Family Bathynellidae Grobbon, 1904
– Bathynella sp. Formerly referred to as
Bathynella chappuisi Delachaux, 1919,
from a well 65 km W of Kalach-on-Don,
Volgograd (Schäfer, 1951). The species
identity requires confirmation (Birstein,
Ljovuschkin, 1967b).
Order Amphipoda Latreille, 1816
Family Crangonyctidae Bousfield, 1973
– Synurella ambulans (F. Müller, 1847)
(= S. meschtscherica Borutzky, 1929). Stygophile, widespread across European Russia. Recorded from springs in the Ryazan
(Borutzky, 1927), Moscow (Chertoprud,
2006), Vladimir, Kaluga, Pskov and Bryansk
regions (Sidorov, Palatov, 2012).
– Synurella derzhavini Behning, 1928. Stygobiont, eyes and integument unpigmented. Described from wells near the city of
Saratov (Behning, 1928).
– Synurella donensis (Martynov, 1919).
Stygobiont, described from springs in the
Kiziterinka River valley, now within the
city of Rostov-on-Don (Martynov, 1919).
Class Arachnida Cuvier, 1812
Order Aranei Clerck, 1758
Family Nesticidae Simon, 1894
– Nesticus cellulanus (Clerck, 1758). Troglophile, found in caves at Kamennoe, Nizhniy Novgorod Region (Kapralov, Chernorudskiy, 2009).
Family Tetragnathidae Menge, 1866
– Metellina merianae (Scopoli, 1763). Troglophile, widespread in Europe at cave entrances, found in caves at Kamennoe, Nizhniy Novgorod Region (Kapralov, Chernorudskiy, 2009).
Superclass Hexapoda Blainville, 1816
Class Insecta Linnaeus, 1758
Order Coleoptera Linnaeus, 1758
Family Cryptophagidae Kirby, 1937
– Cryptophagus setulosus Sturm, 1845. Troglophile, recorded from caves in the Novgorod Region (Kapralov, 2015).
– C. schmidti Sturm, 1845. Troglophile,
recorded from caves of the Novgorod Region (Kapralov, 2015).
– Henoticus serratus (Gyllenhal, 1808).
Troglophile, recorded from the Baskunchakskaya Cave, Akhtubinskiy District,
Astrakhan Region (Kapralov, 2015).
Family Leiodidae Fleming, 1821
– Catops morio (Fabricius, 1792). Troglophile, recorded from caves in the Novgorod
Region (Kapralov, 2015).
Subterranean biota of European Russia
– Choleva glauca (Britten, 1918). Subtroglophile, recorded from caves in the Novgorod Region (Kapralov, 2015).
– Ch. lederiana Reitter, 1902. Subtroglophile, a mass species both in natural and
artificial caverns in the Novgorod Region,
also reported from quarries in the Tver and
Ryazan regions (Kapralov, 2015).
– Cholevinus pallidus (Ménétriés, 1832).
Subtroglophile, recorded in a cave near
Lake Baskunchak, Akhtubinskiy District,
Astrakhan Region (Perkovsky, 1991).
Order Lepidoptera Linnaeus, 1758
Family Noctuidae Latreille, 1809
– Scoliopteryx libatrix (Linnaeus, 1758).
Troglophile, recorded in European Russia
from numerous caves in the Novgorod and
Astrakhan regions (Kapralov, Chernorudskiy, 2009; Mukhanov, Kapralov, 2010;
Kapralov, 2015).
Order Diptera Linnaeus, 1758
Suborder Nematocera Schiner, 1862
Family Trichoceridae Rondani, 1841
– Trichocera maculipennis Meigen, 1818.
Troglophile, common in caves of European
Russia in the Novgorod and Astrakhan regions, as well as the Perm Province (Kapralov, Chernorudskiy, 2009; Pan’kov et al.,
2009b; Kapralov, 2015).
– Trichocera regelationis (Linnaeus, 1758).
Troglophile, common in caves of European
Russia in the Novgorod and Astrakhan regions (Kapralov, Chernorudskiy, 2009;
Kapralov, 2015).
Suborder Brachycera Zetterstedt, 1842
Family Phoridae Gray, 1840
Phoridae gen. spp. Likely troglophiles, larvae found in numerous caves of European
Russia, but identified closer to neither genera nor species (Kapralov, 2015).
Phylum Chordata Haeckel, 1874
Subphylum Vertebrata J.-B. Lamarck, 1801
Class Mammalia Linnaeus, 1758
Order Chiroptera Blumenbach, 1779
Family Vespertilionidae Gray, 1821
Strictly speaking, only a few records of bats
have been made in natural undergrounds due to
scarcely available karst caves. Hence, the ma-
169
jority of findings refer to mines, cellars, rockcut cells and cave monasteries. In many cases
naturally presented cavities and crevices were
deepened and transformed into artificial underground chambers; therefore, it is virtually impossible to estimate the past importance of these
roosts to bats. Whereas no bats reproduce in
central Russia, underground chambers, regardless of their origin, represent the only kind of
winter shelter suitable for the hibernation of
seven resident species. Aside from many dozens
of other records from various underground
roosts, all seven of these species were recorded
from the natural caves of the Ichalkovskiy Nature Reserve, Novgorod Region, where they
have been either netted at entrances during
swarming or observed hibernating (Bakka, Bakka, 1999):
– Myotis dasycneme (Boie, 1825);
– Myotis daubentonii (Kuhl, 1817);
– Myotis nattereri (Kuhl, 1817);
– Myotis brandtii (Eversmann, 1845);
– Myotis mystacinus (Kuhl, 1817);
– Plecotus auritus (Linnaeus, 1758);
– Eptesicus nilssonii (Keyserling et Blasius, 1839).
Conclusion. The terrestrial and aquatic cave
biota in central Russia is scarce because the
Quaternary glaciations must have destroyed all
old cave species. Apparently, the only way for
survival was in subterranenan phreatic and hyporheic habitats (Table 2) (Kniss, Smirnov,
1990; Kniss, 2001, 2004).
IV. 3. Ural
In the Urals, caves are relatively well studied compared to northern Russia’s caverns.
Microarthropod cave fauna is diversified. The
subterranean fauna consists of numerous troglophilic species, as well as several troglobionts
and stygobionts. Troglobionts are known only
amongst Collembola.
Studies on microorganisms in the sediment
of the Ledyanaya and Kungurskaya ice caves in
the central Urals show them to range from
44,000 to 500,000 ex. per gram from winter to
spring (Volodin, Pshenichnikov, 1949).
170
S.I. Golovatch et al.
Table 2. List of *stygobionts, stygophile and troglophile species from central Russia.
Таблица 2. Список *стигобионтов, стигофильных и троглофильных видов
из Центральной России.
A — Stygobionts *: stygophiles
BACTERIA
Mycobacteriaceae
Mycobacterium hyalinum Söhngen, 1913
Mycobacterium licheniforme Krassilnikow, 1949
Mycobacterium luteum Söhngen, 1913
Mycobacterium mucosum Krassilnikow, 1941
Propionibacteriaceae
Propionibacterium sp.
Streptomycetaceae
Streptomyces sp.
Streptoverticillium sp.
Streptosporangiaceae
Streptosporangium sp.
Micrococcaceae
Arthrobacter globiformis Conn et Dimmick, 1947
Bacillaceae
Bacillus mycoides Flügge, 1886
Nostocaceae
Nostoc sp.
Pseudomonadaceae
Pseudomonas desmolyticum Cray et Thornton, 1928
FUNGI
Gymnoascaceae
Pseudoarachniotus sp.
VIRIDIPLANTAE
Chlorophyta: Trebouxiophyceae
Chlorellales sp.
CILIOPHORA
Pleuronematidae
Balantiophorus elongatus Schewiakoff, 1892
EUGLENOZOA
Bodonaceae
Bodo globosus Stein, 1878
Bodo lens (O.F. Müller) Klebs, (1892)
Bodo sp.
RHIZARIA
Euglyphidae
Assulina seminulum (Ehrenberg, 1848)
Euglypha rotunda Wailes, 1915
Trinematidae
Trinema complanatum Penard, 1890
Trinema enchelys (Ehrenberg, 1838)
AMOEBOZOA
B — Troglobionts*: troglophiles
Oligochaeta
Lumbricidae
Eisenia foetida Savigny, 1826
Eiseniella tetraedra (Savigny, 1826)
Aranei
Tetragnatidae
Metellina merianae (Scopoli, 1763)
Arcellidae
Arcella arenaria compressa Chardez, 1974
Centropyxidae
Centropyxis aerophila Deflandre, 1929
Centropyxis aerophila sphagnicola Deflandre, 1929
Centropyxis plagiostoma Thomas et Bonnet, 1955
Hyalospheniidae
Hyalosphenia papilio Leidy, 1879
Nebelidae
Nebela parvula Cash, 1909
Nebela tincta Leidy, 1879
Phryganellidae
Phryganella acropodia (Hertwig et Lesser, 1874)
Phryganella hemisphaerica Penard, 1902
Mastigamoebidae
Mastigamoeba sp.
Hartmannellidae
Hartmannella sp.
Oligochaeta
Enchytraeidae
Henlea ventriculosa (d’Udekem, 1854)
Enchytraeus albidus Henle, 1837
Fridericia bulbosa (Rosa, 1887)
Haplotaxida: Naididae
Nais communis Piguet, 1906
Nais pseudobtusa Piguet, 1906
Haplotaxida: Tubificidae
Potamothrix hammoniensis (Michaelsen, 1901)
COPEPODA
Cyclopoida: Cyclopidae
Acanthocyclops vernalis Fischer, 1853
Harpacticoida: Parastenocarididae
*Parastenocaris fonticola Borutzky, 1926
OSTRACODA
Podocopida: Cyprididae
Potamocypris wolfi Brech, 1920
Syncarida
Bathynellacea: Bathynellidae
*Bathynella sp.
Amphipoda
Crangonyctidae
*Synurella derzhavini Behning, 1925
*Synurella donensis (Martynov, 1919)
Synurella ambulans (F. Müller, 1846)
Nesticidae
Nesticus cellulanus (Clerck, 1758)
COLEOPTERA
Cryptophagidae
Cryptophagus setulosus Sturm, 1845
Cryptophagus schmidti Sturm, 1845
Henoticus serratus (Gyllenhal, 1808)
Subterranean biota of European Russia
171
Leiodidae
Catops morio (Fabricius, 1792)
Choleva glauca (Britten, 1918)
Choleva lederiana Reitter, 1902
Cholevinus pallidus (Ménétriés, 1832)
LEPIDOPTERA
Noctuidae
Scoliopteryx libatrix (Linnaeus, 1758)
DIPTERA
Trichoceridae
Trichocera maculipennis Meigen, 1818
Trichocera regelationis (Linnaeus, 1758)
Phoridae
Phoridae gen. spp.
CHIROPTERA
Vespertilionidae
Myotis dasycneme (Boie, 1825)
Myotis daubentonii (Kuhl, 1817)
Myotis nattereri (Kuhl, 1817)
Myotis brandtii (Eversmann, 1845)
Myotis mystacinus (Kuhl, 1817)
Plecotus auritus (Linnaeus, 1758)
Eptesicus nilssonii (Keyserling et Blasius, 1839)
Phylum Annelida Lamarck, 1809
Class Oligochaeta Grube, 1850
Order Crassiclitellata Jamieson, 1988
Family Lumbricidae Claus, 1876
– Dendrobaena octaedra (Savigny, 1826).
Troglophile, recorded from the ShulganTash Cave, Burzianskiy District, Bashkortostan (= Republic of Bashkiria) (Kniss,
1984b, 2001).
– Dendrodrilus rubidus (Savigny, 1826).
Troglophile, recorded from caves of Bashkortostan (Kniss, 2001).
– Eiseniella tetraedra (Savigny, 1826).
Likely troglophile, recorded from the Shulgan-Tash Cave, Burzianskiy District, Bashkortostan (Kniss, 1984b, 2001).
Phylum Arthropoda Siebold, 1848
Subphylum Crustacea Brünnich, 1772
Class Malacostraca Latreille, 1817
Order Amphipoda Latreille, 1816
Family Crangonyctidae Bousfield, 1973
– Crangonyx chlebnikovi chlebnikovi
Borutzky, 1928 (= Crangonyx chlebnikovi
maximovitshi Pankov et Pankova, 2004).
Stygobiont, found in several caves in the
Perm Province of the Urals (Pan’kov, 2008b;
Pan’kov, Pan’kova, 2004; Pan’kov, Starova, 2009a, 2009c; Pan’kov et al., 2005,
2010, 2011; Sidorov et al., 2010, 2012).
Class Arachnida Cuvier, 1812
Order Aranei Clerck, 1758
Family Linyphiidae Blackwall, 1859
– Improphantes improbulus (Simon, 1929).
Troglophile, recorded from the Kurmanaevskaya and Grot Tashastinskiy caves, both
in Bashkortostan (Esyunin, Efimik, 1999).
– Megalepthyphantes pseudocollinus
Saaristo, 1997. Troglophile, recorded from
the Kungurskaya Cave in the Perm Province (Pan’kov, 2008a; Pan’kov et al.,
2009b).
– Neriene montana (Clerck, 1758). Troglophile, recorded from the Kungurskaya Cave
in the Perm Province (Pan’kov, 2008a;
Pan’kov et al., 2009b).
– Neriene radiata (Walckenaer, 1841). Troglophile, recorded from the Chudesnitsa
Cave in the Perm Province (Pan’kov, 2008a;
Pan’kov et al., 2009b).
Family Nesticidae Simon, 1894
– Nesticus cellulanus (Clerck, 1758). Troglophile, recorded from the Shulgan-Tash
(= Kapovaya) Cave in Bashkortostan (Abdullin et al., 2012), as well as in the Obvalnaya and Letuchikh Myshey caves in the
Perm Province (Pan’kov et al., 2009).
Subclass Acari Leach, 1817
Superorder Acariformes Zakhvatkin, 1952
Order Prostigmata Kramer, 1877
Suborder Anystina Van der Hammen, 1972
Family Rhagidiidae Oudemans, 1922
– Rhagidia breviseta Zacharda, 1995. Apparently troglobiont, recorded from the Shulgan-Tash (= Kapovaya) Cave in Bashkortostan (Kapralov, 2015).
– Rhagidia sp. Apparently troglobiont, recorded from several caves in Bashkortostan,
southern Urals, Russia (Kniss, 1984b, 2001).
– Foveacheles sp. Troglobiont, recorded
from the Shulgan-Tash (= Kapovaya) Cave
in Bashkortostan (Abdullin et al., 2012;
Kapralov, 2015).
172
S.I. Golovatch et al.
Suborder Eleutherengona Oudemans, 1909
Family Pygmephoridae Cross, 1965
– Pygmephorus sp. Troglophile, recorded
from several caves in Bashkortostan, southern Urals (Kniss, 1984b, 2001).
Order Sarcoptiformes Reuter, 1909
Suborder Oribatida Van der Hammen, 1968
Family Aleurodamaeidae Paschoal et
Johnston, 1984
– Epidamaeus sp. Troglophile, recorded
from several caves in Bashkortostan, southern Urals (Kniss, 1984b, 2001).
Family Mochlozetidae Grandjean, 1960
– Sphaerobates sp. Troglophile, recorded
from several caves in Bashkortostan, southern Urals (Kniss, 1984b, 2001).
Superclass Hexapoda Blainville, 1816
Class Collembola Lubbock, 1870
As many as 34 troglophile and 4 troglobiont
species of springtails are known from Ural caves
(Table 3).
Order Poduromorpha Börner, 1913, sensu
D’Haese, 2002
Superfamily Neanuroidea Massoud, 1967,
sensu D’Haese, 2002
Family Neanuridae Börner, 1906
– Philotella olgae Kniss et Thibaud, 1995.
Likely troglobiont, described and only
known from caves in Bashkortostan (Kniss,
Thibaud, 1995; Kniss, 2001, 2006).
Superfamily Hypogastruroidea Salmon,
1964, sensu Deharveng, 2004
Family Hypogastruridae Börner, 1906
– Schaefferia baschkirica Kniss, 1985. Troglobiont, described from the Klyka and
Kiekbayevskaya caves, both on the right
bank of Belaya River, as well as from the
Kanskaya-2 Cave on the left bank of Kana
River, all in Bashkortostan (Kniss, 1985,
2001; Babenko et al., 1994).
– Ceratophysella kapoviensis (Babenko,
1994). Troglobiont with faint traces of troglomorphism. Described from the ShulganTash (= Kapovaya) Cave in Bashkortostan
(Babenko et al., 1994; Kniss, 2001).
Order Entomobryomorpha Börner, 1916,
sensu Soto-Adames et al., 2008
Superfamily Isotomoidea Szeptycki, 1979,
sensu Soto-Adames et al., 2008
Family Isotomidae Schäffer, 1896
– Heteroisotoma stebajevae (Rusek, 1991).
Troglophile, recorded from the Planovaya3 Cave in the Burzianskiy District of Bashkortostan, as well as from the entrance to
the Druzhba Cave near Serga River, Sverdlovsk Region (Jie et al., 2011).
– Sericeotoma knissi Potapov, 1991. Troglobiont, described from the Yakshingulovskaya-1 Cave on the right bank of Belaya River in Bashkortostan (Potapov,
1991).
Superfamily Tomoceroidea Szeptycki, 1979
Family Tomoceridae Schäffer, 1896
– Plutomurus baschkiricus (Skorikow,
1899), troglobiont, endemic to many caves
in Bashkortostan (Turbanov et al., 2016b),
populations studied by Kniss (1984a).
– Tomocerus asiaticus Martynova, 1969
and four other troglophile Tomocerus spp.
are recorded from the Urals.
All theses species are recent cave-dwellers
which must have survived in caves during the
glacial Pleistocene period (Kniss, Thibaud,
1999; Kniss, 1984a, 1984b, 1985, 2004).
Class Insecta Linnaeus, 1758
Order Psocoptera Shipley, 1904
Family Psyllipsocidae Lienhard et Smithers,
2002
– Psyllipsocus sp. Troglophile, recorded
from caves of Bashkortostan, southern Urals
(Kniss, 2001).
Order Coleoptera Linnaeus, 1758
Family Carabidae Latreille, 1802
– Bembidion obscurellum turanicum Csiki,
1928. Troglophile, recorded from caves of
Bashkortostan, southern Urals (Kniss,
2001).
– Bembidion quadrimaculatum (Linnaeus,
1761). Mistakenly referred to as a troglophile, recorded from the Zapovednaya Cave,
Beloretskiy District, Bashkortostan (Kniss,
2001), but actually, just like the previous
subspecies, a trogloxene.
Family Cryptophagidae Kirby, 1937
– Micrambe nigricollis Reitter, 1876. Troglophile, recorded from the Shulgan-Tash
Cave, Burzianskiy District, Bashkortostan
(Kapralov, 2015).
Subterranean biota of European Russia
173
Table 3. List of *stygobionts, stygophiles and *troglobionts, troglophiles from the Urals.
Таблица 3. Список *стигобионтов, стигофилoв и *троглобиотов, троглофилов с Урала.
A — Stygobionts *: stygophiles
AMPHIPODA
Crangonyctidae
*Crangonyx chlebnikovi chlebnikovi Borutzky, 1928
B — Troglobionts *: troglophiles
Oligochaeta
Lumbricidae
Eiseniella tetraedra (Savigny, 1826)
Dendrobaena octaedra (Savigny, 1826)
Dendrodrilus rubidus (Savigny, 1826)
ARANEI
Linyphiidae
Improphantes improbulus (Simon, 1929)
Megalepthyphantes pseudocollinus Saaristo, 1997
Neriene montana (Clerck, 1758)
Nesticidae
Nesticus cellulanus (Clerck, 1758)
ACARI
Prostigmata: Eupodina: Rhagidiidae
*Rhagidia breviseta Zacharda, 1995
Rhagidia sp.
*Foveacheles sp.
Sarcoptiformes: Oribatida: Damaeidae
Epidamaeus sp.
Sarcoptiformes: Oribatida: Mochlozetidae
Sphaerobates sp.
ACARI PARASITI
Parasitidae
Parasitus oudemansi Berlese, 1903
Parasitus remberti (Oudemans, 1912)
Rhodacaridae
Cyrtolaelaps mucronatus Canestrini, 1881
CHILOPODA
Monotarsobius curtipes C.L. Koch, 1847
COLLEMBOLA
Dicyrtomidae
Dicyrtoma fusca (Lubbock, 1873)
Hypogastruridae
*Ceratophysella armata (Nicolet, 1842)
Ceratophysella bengtssoni (Agren, 1904)
Ceratophysella denticulata (Bagnall, 1941)
Ceratophysella kapoviensis (Babenko, 1994)
Ceratophysella sigillata (Uzel, 1890)
Ceratophysella succinea (Gisin, 1949)
Hypogastrura sahlbergi (Reuter, 1895)
Hypogastrura sensilis (Folsom, 1919)
Hypogastrura subboldorii Delamare et Jacquemin,
1962
Hypogastrura.vernalis (Carl, 1901)
*Schaefferia baschkirica Kniss, 1985
Typhlogastrura cf. alabamensis Thibaud, 1975
Entomobryidae
Entomobrya puncteola Uzel, 1891
Lepidocyrtus curvicollis (Bourlet, 1839)
Lepidocyrtus lanuginosus Gmelin, 1788
Lepidocyrtus violaceus (Fourcroy, 1785)
Isotomidae
Desoria olivacea (Tullberg, 1871)
Folsomia diplophthalma (Axelson, 1902)
Folsomia multiseta Stach, 1947
Folsomia quadrioculata Tullberg, 1871
Folsomia spinosa Kseneman, 1936
Heteroisotoma stebajevae Rusek, 1999
Isotoma albella Packard, 1873
Isotoma knissi Potapov, 1991
Isotoma viridis Bourlet, 1839
Proisotoma notabilis (Schaeffer, 1896)
Neanuridae
*Philotella olgae Kniss et Thibaud, 1996
Onychiuridae
Hymenaphorura sibirica (Tullberg, 1876)
Onychiuroides granulosus (Stach, 1930)
Onychiurus furcifer (Börner, 1901)
Onychiurus schoetti Lie-Pettersen, 1896
Protaphorura armata (Tullberg, 1869)
Tomoceridae
*Plutomurus baschkiricus (Skorikow, 1899)
Tomocerina minuta (Tullberg, 1876)
Tomocerus asiaticus Martynova, 1969
Tomocerus minor (Lubbock, 1862)
Tomocerus cf. punctatus Yosii, 1967
Tomocerus vulgaris (Tullberg, 1871)
Arrhopalitidae
Arrhopalites principalis Stach, 1945
PSOCOPTERA
Psyllipsocidae
Psyllipsocus sp.
COLEOPTERA
Carabidae
Bembidion obscurellum turanicum Csiki, 1928
Bembidion quadrimaculatum (Linnaeus, 1761)
Cryptophagidae
Micrambe nigricollis Reitter, 1876
Leiodidae
Choleva glauca (Britten, 1918)
Choleva lederiana Reitter, 1902
Staphylinidae
Atheta sp.
Falagria thoracica Curtis, 1833
Geostiba circellaris Gravenhorst, 1802
174
S.I. Golovatch et al.
LEPIDOPTERA
Noctuidae
Scoliopteryx libatrix (Linnaeus, 1758)
DIPTERA
Mycetophilidae
Speolepta leptogaster (Winnertz, 1863)
Sciaridae
Sciaridae gen. spp.
Trichoceridae
Trichocera maculipennis Meigen, 1818
Phoridae
Phoridae gen. spp.
Family Leiodidae Fleming, 1821
– Choleva glauca (Britten, 1918). Subtroglophile, recorded from the Shulgan-Tash
Cave, Burzianskiy District, Bashkortostan
(Kapralov, 2015).
– Choleva lederiana Reitter, 1902. Subtroglophile, recorded from the Kungurskaya
Ice Cave (Koz’minykh, 2012).
Family Staphylinidae Lameere, 1900
– Atheta sp. Troglophile, recorded from
caves of the southern Urals: Ishcheevskaya2, Ishimbayskiy District, and Shulgan-Tash,
Burzianskiy District, both Bashkortostan
(Kniss, 2001).
– Falagria thoracica Curtis, 1833. Troglophile, recorded from the Bortovoy Otpor
Cave and the Zilimskiy grotto, Gafuriyskiy
District, Bashkortostan (Kniss, 2001).
– Geostiba circellaris Gravenhorst, 1802.
Troglophile, recorded from caves of Bashkortostan, southern Urals (Kniss, 2001).
Order Lepidoptera Linnaeus, 1758
Family Noctuidae Latreille, 1809
– Scoliopteryx libatrix (Linnaeus, 1758).
Troglophile, recorded from numerous caves
in the Perm Province and Bashkortostan
(Pan’kov et al., 2009b; Kapralov, 2015).
Order Diptera Linnaeus, 1758
Suborder Nematocera Schiner, 1862
Family Mycetophilidae Newman, 1835
– Speolepta leptogaster (Winnertz, 1863).
Troglophile, recorded from the ShulganTash Cave, Burzianskiy District, Bashkortostan; the characteristic hunting nets also
reported from the Dudkinskaya adit, Ufa
City, and the Butylka Cave, both Bashkortostan as well (Kapralov, 2015).
Scathophagidae
Scathophagidae gen. spp.
CHIROPTERA
Vespertilionidae
Myotis dasycneme (Boie, 1825)
Myotis daubentonii (Kuhl, 1817)
Myotis nattereri (Kuhl, 1817)
Myotis brandtii (Eversmann, 1845)
Myotis mystacinus (Kuhl, 1817)
Myotis davidii Peters, 1869
Plecotus auritus (Linnaeus, 1758)
Eptesicus nilssonii (Keyserling et Blasius, 1839)
Family Sciaridae Billberg, 1820
– Sciaridae gen. spp. Likely troglophiles,
larvae encountered in several caves of European Russia, including the southern Urals.
A reliable record concerns the Starosaitovskaya-1 Cave, Ishimbayskiy District,
Bashkortostan (Kniss, 2001; Kapralov,
2015).
Family Trichoceridae Rondani, 1841
– Trichocera maculipennis Meigen, 1818.
Troglophile, common in caves of the Perm
Province (Pan’kov et al., 2009b; Kapralov,
2015).
Suborder Brachycera Zetterstedt, 1842
Family Phoridae Gray, 1840
– Phoridae gen. spp. Likely troglophiles,
larvae encountered in several caves in the
southern Urals, but remain unidentified as
to genus and species (Kapralov, 2015).
Family Scathophagidae Robineau-Desvoidy,
1830
Specimens unidentified to genus and species have been encountered in the Babinogorskaya Cave, Kungurskiy District,
Perm Province (Pan’kov et al., 2009b).
Phylum Chordata Haeckel, 1874
Subphylum Vertebrata J.-B. Lamarck, 1801
Class Mammalia Linnaeus, 1758
Order Chiroptera Blumenbach, 1779
Family Vespertilionidae Gray, 1821
According to recent data, eight bat species
of the 14 occurring in the Urals are resident in
the region, while others perform seasonal longdistance migrations and depart in winters. No
maternity roosts have been found underground
so far, as all records in caves were related to
swarming or hibernating bats belonging to the
Subterranean biota of European Russia
following eight species (see Bol’shakov et al.
(2005), Snit’ko (2007, 2009, 2011), and Snit’ko
and Snit’ko (2016, 2017a, 2017b) for details):
– Myotis dasycneme (Boie, 1825) (Fig.
4A). This species, common in the region,
was found in more than 50 cave hibernacula
located from north to south along the Urals,
with the largest known aggregations in the
Smolinskaya and Arakaevskaya caves in
the Sverdlovsk Region (up to 2,000 and
over 200, respectively). In the southern
Urals, no substantial winter colonies have
been discovered, although this bat is relatively abundant at cave entrances during
swarming time.
– Myotis daubentonii (Kuhl, 1817) (Fig.
4D). Daubenton’s bat is as fairly common
in the Urals as elsewhere within its distribution range in Russia; it was recorded in over
50 caves throughout the region. Unlike the
Pond Bat, it forms no large winter aggregations. Therefore, it is usually undercounted
during surveys at hibernation sites. The
Smolinskaya and Arakaevskaya caves in
the Sverdlovsk Region alongside the Nadezhda and Sukhokamenskaya caves in the
Chelyabinsk Region, and the Kueshta Cave
in Bashkiria comprise the largest winter
colonies which range from 50 to 110 individuals.
– Myotis nattereri (Kuhl, 1817). The species was considered rare in the region because only few records from the middle and
southern Urals were reported in the 20th
century. However, Natterer’s bats turned
out to be very common in caves of the
southern Urals in late summer and autumn,
where these bats were caught in large numbers at cave entrances as they swarmed,
with the largest number of 1,000 animals
counted in the Kyzyl-Yar Cave in Bashkiria.
– Myotis brandtii (Eversmann, 1845) (Fig.
4E). Brandt’s bat was found in >20 and 29
underground roosts in the middle and southern Urals, respectively. The Divya Cave in
the Perm Province represents the northernmost hibernation roost in the Urals with the
175
largest winter colony (up to 1,000 individuals); only a few Brandt’s bats were found
hibernating in other caves. Despite its relative rarity in winter, a number of Brandt’s
bats were netted at swarming sites in the
southern Urals, where M. brandtii was the
second most abundant species following E.
nilssonii in Bashkiria and following P. auritus in the Chelyabinsk Region.
– Myotis mystacinus (Kuhl, 1817) (Fig.
4F). A few reliable records from the northern Urals were made in the Divya Cave. In
the southern Urals, the species is abundant
in swarming and hibernation undergrounds
in Bashkiria, where about 50 bats were
netted at the entrances to the caves Kueshta,
Laklinskaya, Oktyabrskaya, Nadezhda and
Sukhokamenskaya.
– Myotis davidii Peters, 1869. It is unclear
yet, which of the previous records of bats
from the M. mystacinus morpho-group
should be assigned to this species (see Benda et al. (2016) for a taxonomic review).
Two individuals identified as undoubtedly
M. davidii were found in the Podarok Cave,
Orenburg Region in September 2016.
– Plecotus auritus (Linnaeus, 1758) (Fig.
4B). The Brown long-eared bat is widely
distributed, albeit not abundant in hibernation sites in the Urals. It was found in more
than 60 caves throughout the region, northwards up to the Divya Cave. It was also very
common in samples netted during swarming in the southern part of the region (Chelyabinsk Region and Bashkiria), where it
constituted 15 and 24% of all captured bats,
respectively.
– Eptesicus nilssonii (Keyserling et Blasius, 1839) (Fig. 4C). The Northern Bat is
the most widely distributed and abundant
species in the Urals. It has been found in
more than 70 underground sites across the
region, with the largest winter aggregation
in the Arakaevskaya Cave, Perm Province.
E. nilssonii comprised a large proportion of
swarming bats at the entrances to caves in
the southern Urals, being the most abundant
species in the Chelyabinsk Region.
176
S.I. Golovatch et al.
Fig. 4. Some species of bats during hibernation in caves of the Urals. A — hibernating colonies of the Pond
bat, Myotis dasycneme (Boie, 1825) in the Smolinskaya Cave, Sverdlovsk Region. B — a hibernating Brown
long-eared bat, Plecotus auritus (Linnaeus, 1758) in the Nadezhda Cave, Chelyabinsk Region. C — a
hibernating Northern bat, Eptesicus nilssonii (Keyserling et Blasius, 1839) in the Nukatovskaya Cave,
Bashkiria. D — a hibernating Daubenton’s bat, Myotis daubentonii (Kuhl, 1819) in the Sukhaya Atya Cave,
Chelyabinsk Region. E — a hibernating Brandtґs bat, Myotis brandtii (Eversmann, 1845) in the Kyzyl-Yar
Cave, Bashkiria. F — a hibernating Whiskered bat, Myotis mystacinus (Kuhl, 1817) in the Sukhokamenskaya
Cave, Chelyabinsk Region. All pictures by Ksenia Snit’ko.
Subterranean biota of European Russia
Conclusion. The subterranean fauna of the
Urals, both terrestrial and aquatic, is mainly
troglophile and stygophile, with a reduced number of endemic and relict cave species (Table 3).
The impact of glacial Pleistocene phases may
account for the scarcity of troglobionts and the
relatively high number of recent troglophiles.
IV. 4. The Russian Great Caucasus
Charitonov (= Kharitonov) (1947), Malevich
(1947), Borutzky (1934, 1948a, b, 1950a, b,
1961, 1965, 1967a, b), Birstein (1948, 1950,
1952, 1954, 1967), Starobogatov (1962), Ljovuschkin and Starobogatov (1963), Rudjakov
(1963), Zalesskaja (1963, 1973a, b, 1978), Ljovuschkin and Matiokin (1965), Ljovuschkin
(1962, 1963, 1965, 1966, 1970, 1972a, b, 1973),
Birstein and Ljovuschkin (1967a), Danielopol
(1969), Golovatch (1975, 1978, 1983, 19841985, 1990, 2011), Read (1992), Belousov
(1999), Kniss and Thibaud (1999), Kantor and
Sysoev (2005), Belousov and Koval (2009,
2011), Chemeris (2009), Karaman (2012), Antić and Makarov (2016) and several other authors have provided much of the information on
invertebrates of the region in question. A full
updated checklist and an as complete bibliography as possible concerning the entire territory of
the former Soviet Union are available in Turbanov et al. (2016a, b, c).
The Russian Caucasus consists of Ciscaucasia (= Circassia) and the western part of Transcaucasia as far south as Psou River, bordering
on Abkhazia. The subterranean biota can be
estimated as being relatively well known in
Russia, also being the richest in endemic and
relict species with more than 80 troglo- and
stygobionts revealed (Table 4).
Kingdom Chromista Cavalier-Smith, 1981
Phylum Ochrophyta Cavalier-Smith, 1995
177
Class Chrysophyceae Pascher, 1914
Order Ochromonadales Pascher, 1910
Family Paraphysomonadaceae Preisig et Hibberd, 1983
– Paraphysomonas bandaiensis (Hibberd,
1979). Likely stygophile, recorded from
the Vorontsovskaya Cave, Khostinsky City
District, Greater Sochi, Krasnodar Province (Myl’nikov et al., 2006).
Family Chromulinaceae Engler, 1897
– Spumella sp. Likely stygophile, recorded
from the Vorontsovskaya Cave, Khostinsky City District, Greater Sochi, Krasnodar
Province (Myl’nikov et al., 2006).
Class Raphidophyceae Chadefaud et Silva, 1980
Order Thaumatomonadida Shirkina, 1987
Family Thaumatomastigidae Patterson et
Zolffel, 1991
– Thaumatomonas seravini Mylnikov and
Karpov, 1993. Likely stygophile, recorded
from the Vorontsovskaya Cave, Khostinsky City District, Greater Sochi, Krasnodar
Province (Myl’nikov et al., 2006).
Superphylum Alveolata Cavalier-Smith, 1991
Phylum Ciliophora Doflein, 1901
Class Phyllopharyngea de Puytorac et al., 1974
Subclass Suctoria Claparède et Lachmann,
1858
– Suctoria gen. sp. Epibiont forms found on
hypogean amphipods in the Beloskal’skaya
and Pionerskaya (= Ushchel’naya) caves,
Khostinsky City District, Greater Sochi,
Krasnodar Province (Birstein, 1950).
Kingdom Rhizaria Cavalier-Smith, 2002
Phylum Cercozoa Cavalier-Smith, 1998
Class Sarcomonadea Cavalier-Smith, 1993
Order Glissomonadida Howe, Bass, Vickerman, Chao et Cavalier-Smith, 2009
Family Allapsidae Howe, Bass, Vickerman,
Chao et Cavalier-Smith, 2009
Рис. 4. Некоторые виды летучих мышей во время зимовки в пещерах Урала. A — зимующие колонии
ночницы прудовой (Myotis dasycneme (Boie, 1825)) в Смолинской пещере (Свердловская обл.). B —
зимующий ушан (Plecotus auritus (Linnaeus, 1758)) в пещере Надежда (Челябинская обл.). С —
зимующий северный кожанок (Eptesicus nilssonii (Keyserling et Blasius, 1839)) в пещере Нукатовская
(Башкирия). D — зимующая водяная ночница (Myotis daubentonii (Kuhl, 1819)) в пешере Сухая Атья
(Челябинская обл.). E — зимующая ночница Брандта (Myotis brandtii (Eversmann, 1845)) в пещере
Кызыл-Яр (Башкирия). F — зимующая усатая ночница (Myotis mystacinus (Kuhl, 1817)) в пещере
Сухокаменская (Челябинская обл.). Все фотографии Ксении Снитько.
178
S.I. Golovatch et al.
Table 4. List of *stygobionts, possible stygobionts, stygophiles, possible troglobionts
and troglophiles from the Caucasus.
Таблица 4. Список *стигобионтов, вероятных стигобионтов, стигофилoв, возможных
троглобиотов и троглофилов с Кавказа.
A — Stygobionts *: stygophiles
CHROMISTA
Paraphysomonadaceae
Paraphysomonas bandaiensis (Hibberd, 1979)
Chromulinaceae
Spumella sp.
RHIZARIA
Thaumatomastigidae
Thaumatomonas seravini Mylnikov et Karpov, 1993
Thaumatomonas seravini Mylnikov et Karpov, 1993
Allapsidae
Allantion tachyploon Sandon, 1924
Heteromitidae
Heteromita globosa (Stein, 1878)
Heteromita sp.
ALVEOLATA
Suctoria gen. sp.
MYZOZOA
Protaspidaceae
Protaspis simplex Vors, 1992
Colponemidae
Colponema edaphicum Mylnikov et Tikhonenkov,
2007
SARCOMASTIGOTA
Bodonaceae
Bodo designis Skuja, 1948
Bodo saliens Larsen et Paterson, 1990
Bodo saltans Ehrenberg, 1838
Cercomonas granulifera (Hollande, 1942)
Cercomonas laciniaegerens (Krassilstschick, 1886)
Cercomonas sp.
Dimastigella mimosa Frolov, Mylnikov et Malysheva, 1997
Rhynchomonas nasuta (Stokes) Klebs, 1892
AMOEBOZOA
Centropyxidae
Centropyxis constricta (Ehrenberg, 1841)
Centropyxis ecornis (Ehrenberg, 1841)
Centropyxis orbicularis Deflandre, 1929
Centropyxis plagiostoma Bonnet et Thomas, 1955
Centropyxis platystoma (Penard, 1890)
Centropyxis spinosa Cash, 1905
Difflugiidae
Difflugia avellana Penard, 1890
Difflugia avellana gigas Gauthier-Lievre et Thomas, 1958
Difflugia penardi Hopkinson, 1909
Difflugia oblonga Ehrenberg, 1838
Pontigulasia incisa Rhumbler, 1896
Heleoperidae
Heleopera sphagni Leidy, 1874
MYCETOZOA
Physaridae
Physarum flagellatum (Alexeieff, 1923)
PLATYHELMINTHES
Tricladida gen. sp.
Dugesiidae
Dugesia taurocaucasica (Livanov, 1951)
Dendrocoelidae
Dendrocoelidae gen. sp.
*Dendrocoelum sp. 1
*Dendrocoelum sp. 2
MOLLUSCA: GASTROPODA
Hydrobiidae
Belgrandiella caucasica Starobogatov, 1962
*“Geyeria” valvataeformis Starobogatov, 1962
*“Lartetia” sp.
*Paladilhiopsis orientalis Starobogatov, 1962
*Paladilhiopsis pulcherrima Starobogatov, 1962
*Paladilhiopsis subovata Starobogatov, 1962
MOLLUSCA: BIVALVIA
Euglesidae
Euglesa personata (Malm, 1855)
*Euglesa cavatica (Shadin, 1952)
HIRUDINEA
Erpobdellidae
Erpobdella octoculata (Linnaeus, 1758)
COPEPODA
Cyclopoida
Cyclopidae
*Acanthocyclops venustus stammeri (Kiefer, 1930)
Diacyclops bisetosus (Rehberg, 1880)
Paracyclops fimbriatus imminutus Kiefer, 1929
*Speocyclops cinctus Monchenko, 1982
*Speocyclops lussianus Borutzky, 1950
*Speocyclops psezuapsensis Borutzky, 1965
Harpacticoida
Ameiridae
*Nitocrella hirta caucasica Borutzky, 1967
*Megastygonitocrella ljovuschkini (Borutzky, 1967)
Canthocamptidae
Attheyella crassa (G.O. Sars, 1863)
*Bryocamptus aquaeductus Borutzky, 1940
Bryocamptus hostensis Borutzky, 1972
*Bryocamptus innominatus Borutzky, 1940
Bryocamptus pygmaeus (G.O. Sars, 1863)
Bryocamptus tarnogradskyi Borutzky, 1934
Bryocamptus zschokkei caucasicus Borutzky, 1930
Canthocamptus staphylinus (Jurine 1820)
Elaphoidella bidens coronata (G.O. Sars, 1904)
*Pilocamptus georgevitchi (Chappuis, 1923)
*Elaphoidella czerkessica Borutzky, 1972
*Moraria hostensis Borutzky, 1972
Subterranean biota of European Russia
*Moraria operculata Borutzky, 1948
*Moraria poppei (Mrazek, 1893)
*Moraria varica (Graeter, 1910)
Parastenodarididae
*Parastenocaris tenuis Borutzky, 1948
*Parastenocaris sp.
Ostracoda
Candonidae
*Trapezicandona ljovuschkini (Rudjakov, 1963)
Cyprididae
Cypria reptans Bronstein, 1947
Limnocytheridae
*Kovalevskiella rudjakovi (Danielopol, 1969)
Syncarida
Bathynellidae
*Antrobathynella stammeri ciscaucasica (Birstein
et Ljovuschkin, 1964)
*Bathynella natans natans Vejdovsky, 1882
Isopoda
Asellota: Asellidae
*Proasellus linearis (Birstein, 1967)
*Proasellus ljovuschkini (Birstein, 1967)
*Proasellus similis (Birstein, 1967)
179
Asellota: Microparasellidae
*Microcharon tantalus Birstein et Ljovuschkin, 1964
AMPHIPODA
Crangonyctidae
*Lyurella shepsiensis Sidorov, 2015
Niphargidae
*Niphargus abchasicus Martynov, 1932
*Niphargus caelestis G. Karaman, 1982
*Niphargus cubanicus Birstein, 1954
*Niphargus krasnodarus G. Karaman, 2012
*Niphargus latimanus Birstein, 1952
*Niphargus cf. magnus Birstein, 1940
*Niphargus potamophilus Birstein, 1954
*Niphargus pseudolatimanus Birstein, 1952
*Niphargus smirnovi Birstein, 1952
*Niphargus submersus (Derzhavin, 1945)
Typhlogammaridae
*Zenkevitchia sp.
DECAPODA
Caridea: Atyidae
*Troglocaris (Xyphocaridella) jusbaschjani Birstein,
1948
Brachyura: Potamidae
Potamon ibericum tauricum (Czerniavsky 1884)
B — Troglobionts *: troglophiles
OLIGOCHETA
Lumbricidae
Allolobophora cavatica Michaelsen, 1910
Dendrobaena veneta crassa (Malevics, 1947)
Eisenia fetida (Savigny, 1826)
Naididae
Homochaeta sp.
Peloscolex sp.
NEMATODA
Monhysteridae
Monchystera paludicola De Man, 1880
Dorylaimidae
Dorylaimus stagnalis Dujardin, 1845
Dorylaimus callosus Skwarra, 1921
Eudorylaimus laticollis (De Man, 1907)
Eudorylaimus condamni (Vanha, 1893)
Eudorylaimus iners (Bastian, 1865)
Actinolaimidae
Trachactinolaimus radulatus Andrássy, 1963
Mononchidae
Mononchus truncatus Bastian, 1865
Prionchulus muscorum (Dujardin, 1845)
Tripylidae
Tripyla papillata Bütschli, 1873
Tripyla filicaudata De Man, 1880
Prismatolaimidae
Prismatolaimus dolichurus De Man, 1880
Criconematidae
Criconema fimbriatum Cobb in Taylor, 1936
Criconema sp.
Criconemoides sp.
Mollusca
Orculidae
Lauria sp.
Trigonochlamydidae
*Troglolestes sokolovi Ljovuschkin et Matiokin,
1965
Orculidae
*Euxinolauria vitrea (Schileyko, 1988)
Clausiliidae
Scrobifera taurica (L. Pfeiffer, 1848)
Serrulina sp.
Zonitidae
*Conulopolita cavatica (Riedel, 1966)
Oxychilus (Longiphallus) sp.
ISOPODA TERRESTRIA
Oniscidea
Ligiidae
Ligidium zaitzevi Borutzky, 1950
*Ligidium cavaticum Borutzky, 1950
Buddelundiellidae
*Pseudobuddelundiella hostensis Borutzky, 1967
*Pseudobuddelundiella ljovuschkini Borutzky, 1967
*Trichoniscus pygmaeus tuapsensis Borutzky, 1972
Cylisticidae
Cylisticus birsteini Borutzky, 1961
Trichoniscidae
*Caucasocyphonethes cavaticus Borutzky, 1948
*Psachonethes czerkessicus Borutzky, 1969
ARANEI
Agelenidae
Tegenaria abchasica Charitonov, 1941
180
S.I. Golovatch et al.
Tegenaria pontica Charitonov, 1947
Tegenaria sp.
Hahniidae
Iberina ljovuschkini Pichka, 1965
Tetragnathidae
Meta bourneti Simon, 1922.
Metellina merianae (Scopoli, 1763)
Linyphiidae
Troglohyphantes birsteini Charitonov, 1943
Nesticidae
Carpathonesticus birsteini (Charitonov, 1947)
Carpathonesticus ljovuschkini (Pichka, 1965)
Carpathonesticus zaitzevi (Charitonov, 1939)
Carpathonesticus sp.
Aituaria pontica (Spassky, 1932)
Pholcidae
Hoplopholcus longipes (Spassky, 1934)
PSEUDOSCORPIONES
Neobisiidae
*Neobisium (Neobisium) speleophilum Krumpal, 1986
*Neobisium sp.
OPILIONES
Nemastomatidae
Histricostoma caucasicum (Redikorzev 1936)
*Nemaspela abchasica (Ljovuschkin et Starobogatov, 1963)
*Nemaspela sokolovi (Ljovuschkin et Starobogatov, 1963)
*Nemaspela kovalii Chemeris, 2009
Phalangiidae
Nelima pontica Charitonov, 1941
ACARI
Phthiracaridae
Phthiracarus globosus (C.L. Koch, 1841)
Ixodidae
Ixodes vespertilionis C.L. Koch, 1844
CHILOPODA
Lithobiidae
Harpolithobius perplexus Zalesskaja, 1973
Lithobius liber Lignau, 1903
Lithobius reconditus Zalesskaja, 1973
Lithobius stuxbergii Seliwanoff, 1880
DIPLOPODA
Glomeridellidae
Typhloglomeris caucasica Golovatch, 1975
Doderiidae
Trachysphaera costata (Waga, 1857)
Trachysphaera minuta Golovatch, 1976
Anthroleucosomatidae
*Caucaseuma elephantum Antić et Makarov, 2016
*Caucaseuma fanagoriyskaya Antić et Makarov,
2016
*Caucaseuma lohmanderi Strasser, 1970
*Caucaseuma minellii Antić et Makarov, 2016
*Heterocaucaseuma feminaepectorum Antić et
Makarov, 2016
Metamastigophorophyllon giljarovi (Lang, 1959)
Blaniulidae
*Nopoiulus ammonites Enghoff, 1984
Nopoiulus kochii (Gervais, 1847)
Julidae
Archileucogeorgia sp.
Cylindroiulus placidus (Lignau, 1903)
Cylindroiulus pterophylacum Read, 1992
Cylindroiulus schestoperovi Lohmander, 1932
Paradoxosomatidae
Strongylosoma kordylamythrum Attems, 1898
Polydesmidae
Brachydesmus furcatus Lohmander, 1936
Trichopolydesmidae
*Caucasodesmus inexpectatus Golovatch, 1985
Collembola
Hypogastruridae
*Typhlogastrura preobrazhenskyi Babenko, 1987
Isotomidae
Desoria fennica (Reuter, 1895)
Tomoceridae
*Plutomurus jelesnovodskii Kniss et Thibaud, 1999
*Plutomurus kelasuricus (Martynova, 1969)
*Plutomurus sorosii Kniss et Thibaud, 1999
ORTHOPTERA
Rhaphidophoridae
*Dolichopoda euxina Semenov, 1901
TRICHOPTERA
Polycentropodidae
Plectrocnemia sp. sensu Lepneva, 1940
COLEOPTERA
Carabidae
*Caucasaphaenops molchanovi Belousov, 1999
*Caucasorites kovali amplicolis Belousov, 1999
*Caucasorites kovali Belousov, 1999
*Caucasorites shchurovi Belousov et Zamotajlov,
1997
*Caucasorites victori Belousov, 1999
Cimmerites circassicus Reitter, 1888
*Cimmerites kryzhanovskii Belousov 1999
*Cimmerites maximovitchi Belousov et Koval, 2011
Cimmerites zamotajlovi Belousov, 1998
*Duvalius miroshnikovi Belousov et Zamotajlov,
1995
Duvalius gusevi Belousov, 1989
*Jeannelius birsteini Ljovuschkin, 1965
Jeannelius zhicharevi Lutshnik, 1915
Laemostenus koenigi (Reitter, 1887)
Laemostenus tschitscherini Semenov, 1909
Meganophthalmus irinae Belousov et Zamotajlov,
1999
*Meganophthalmus kravetzi Komarov, 1993
*Nannotrechus ciscaucasiens (Ljovuschkin, 1972)
Nannotrechus fishtensis Belousov, 1989
Nannotrechus kovali Belousov, 1989
Porocimmerites imitator Belousov, 1998
Pterostichus lacunosus (Chaudoir 1844)
Trechus heniochicus Ljovuschkin, 1970
Subterranean biota of European Russia
181
Cryptophagidae
Cryptophagus pilosus Gyllenhal, 1827
Curculionidae
*Otiorhynchus vargovitchi Davidian, 2007
Elmidae
Limnius colchicus Delève, 1963
Leiodidae
Catops subfuscus Kellner, 1846
Choleva sp.
Pselaphidae
Bryaxis balneator Besuchet et Kurbatov, 2007
Bryaxis kovali Besuchet et Kurbatov, 2007
*Seracamaurops komarovi Hlaváč, Kodada et Koval, 1999
Ptiliidae
Ptenidium intermedium Wankowicz, 1896
Scirtidae
Odeles sp.
Staphylinidae
Bisnius parcus (Sharp, 1874)
Heinzia caucasica Gusarov et Koval, 2002
DIPTERA
Limoniidae
Limonia nubeculosa Meigen, 1804
Sciaridae
Neosciara sp.
Heleomyzidae
Heteromyza atricornis Meigen, 1830
CHIROPTERA
Rhinolophidae
Rhinolophus euryale Blasius, 1853
Rhinolophus ferrumequinum (Schreber, 1774)
Rhinolophus hipposideros (Borkhausen, 1797)
Rhinolophus mehelyi Matschie, 1901
Vespertilionidae
Barbastella barbastellus (Schreber, 1774)
Barbastella caspica Satunin, 1908
Eptesicus serotinus (Schreber, 1774)
Hypsugo savii (Bonaparte, 1837)
Myotis alcathoe von Helversen et Heller, 2001
Myotis davidii Peters, 1869
Myotis bechsteinii (Kuhl, 1817)
Myotis blythii (Tomes, 1857)
Myotis brandtii (Eversmann, 1845)
Myotis dasycneme (Boie, 1825)
Myotis daubentonii (Kuhl, 1817)
Myotis emarginatus (Geoffroy, 1806)
Myotis mystacinus (Kuhl, 1817)
Myotis nattereri (Kuhl, 1817)
Nyctalus noctula (Schreber, 1774)
Pipistrellus pipistrellus (Schreber, 1774)
Plecotus auritus (Linnaeus, 1758)
Plecotus macrobullaris Kuzyakin, 1965
Miniopteridae
Miniopterus schreibersii (Kuhl, 1817)
Molossidae
Tadarida teniotis (Rafinesque, 1814)
– Allantion tachyploon Sandon, 1924. Likely stygophile, recorded from the Vorontsovskaya Cave, Khostinsky City District,
Greater Sochi, Krasnodar Province (Myl’nikov et al., 2006).
Order Cercomonadida Poche, 1913
Family Heteromitidae Kent, 1880
– Heteromita globosa (Stein, 1878). Likely
stygophile, recorded from the Vorontsovskaya Cave, Khostinsky City District,
Greater Sochi, Krasnodar Province (Myl’nikov et al., 2006).
– Heteromita sp. Likely stygophile, recorded from the Vorontsovskaya Cave, Khostinsky City District, Greater Sochi, Krasnodar
Province (Myl’nikov et al., 2006).
Phylum Myzozoa Cavalier-Smith et Chao, 2004
Class Dinophyceae Fritsch, 1927
Order Desmomastigales Bourrelly, 1970
Family Protaspidaceae Skuja, 1939
– Protaspis simplex Vors, 1992. Likely
stygophile, recorded from the Vorontsovskaya Cave, Khostinsky City District,
Greater Sochi, Krasnodar Province (Myl’nikov et al., 2006).
Class Colponemea Cavalier-Smith, 1993
Order Colponemida Cavalier-Smith, 1993
Family Colponemidae Cavalier-Smith et
Chao, 2004
– Colponema edaphicum Mylnikov et
Tikhonenkov, 2007. Likely stygophile, described from the Vorontsovskaya Cave,
Khostinsky City District, Greater Sochi,
Krasnodar Province (Myl’nikov et al., 2006;
Myl’nikov, Tikhonenkov, 2007).
Kingdom Protozoa R. Owen, 1858
Subkingdom Sarcomastigota Cavalier-Smith,
1983
Group Excavata (Cavalier-Smith), 2002
Phylum Euglenozoa Cavalier-Smith, 1981
Class Kinetoplastea Honigberg, 1963
Order Eubodonida Vickerman, in Moreira et
al., 2004
Family Bodonaceae Bütschli, 1884
– Bodo designis Skuja, 1948. Likely stygophile, recorded from the Vorontsovskaya
182
S.I. Golovatch et al.
Cave, Khostinsky City District, Greater
Sochi, Krasnodar Province (Myl’nikov et
al., 2006).
– Bodo saliens Larsen et Paterson, 1990.
Likely stygophile, recorded from the
Vorontsovskaya Cave, Khostinsky City
District, Greater Sochi, Krasnodar Province (Myl’nikov et al., 2006).
– Bodo saltans Ehrenberg, 1838. Likely
stygophile, recorded from the Vorontsovskaya Cave, Khostinsky City District,
Greater Sochi, Krasnodar Province (Myl’nikov et al., 2006).
– Cercomonas granulifera (Hollande,
1942). Likely stygophile, recorded from
the Vorontsovskaya Cave, Khostinsky City
District, Greater Sochi, Krasnodar Province (Myl’nikov et al., 2006).
– Cercomonas laciniaegerens (Krassilstschick, 1886). Likely stygophile, recorded from the Vorontsovskaya Cave,
Khostinsky City District, Greater Sochi,
Krasnodar Province (Myl’nikov et al., 2006).
– Cercomonas sp. Likely stygophile, recorded from the Vorontsovskaya Cave,
Khostinsky City District, Greater Sochi,
Krasnodar Province (Myl’nikov et al.,
2006).
– Dimastigella mimosa Frolov, Myl’nikov
et Malysheva, 1997. Likely stygophile, recorded from the Vorontsovskaya Cave,
Khostinsky City District, Greater Sochi,
Krasnodar Province (Myl’nikov et al., 2006).
– Rhynchomonas nasuta (Stokes) Klebs,
1892. Likely stygophile, recorded from the
Vorontsovskaya Cave, Khostinsky City District, Greater Sochi, Krasnodar Province
(Myl’nikov et al., 2006).
Phylum Amoebozoa Lühe, 1913
Class Tubulinea Smirnov, Nassonova, Berney,
Fahrni, Bolivar et Pawlowski, 2005
Order Arcellinida Kent, 1880
Family Centropyxidae Jung, 1942
– Centropyxis constricta (Ehrenberg, 1841).
Stygophile, recorded from the Labirintovaya
Cave, Khostinsky City District, Greater
Sochi, Krasnodar Province (Chibisova,
1967).
– Centropyxis ecornis (Ehrenberg, 1841).
Likely stygophile, recorded from sulfuric
springs in the karst canyon of Agura River,
Khostinsky City District, Greater Sochi,
Krasnodar Province (Chibisova, 1967).
– Centropyxis orbicularis Deflandre, 1929.
Stygophile, recorded from the Labirintovaya
Cave, Khostinsky City District, Greater
Sochi, Krasnodar Province (Chibisova,
1967).
– Centropyxis plagiostoma Bonnet et Thomas, 1955. Stygophile, recorded from the
Labirintovaya Cave, Khostinsky City District, Greater Sochi, Krasnodar Province
(Chibisova, 1967; Ljovuschkin 1972b).
– Centropyxis platystoma (Penard, 1890).
Stygophile, recorded from the Labirintovaya
Cave, Khostinsky City District, Greater
Sochi, Krasnodar Province (Chibisova,
1967).
– Centropyxis spinosa Cash, 1905. Stygophile, recorded from the Labirintovaya
Cave, Khostinsky City District, Greater
Sochi, Krasnodar Province (Chibisova,
1967).
Family Difflugiidae Wallich, 1864
– Difflugia avellana Penard, 1890. Stygophile, recorded from the Labirintovaya
Cave, Khostinsky City District, Greater
Sochi, Krasnodar Province (Chibisova,
1967).
– Difflugia avellana gigas Gauthier-Lievre
et Thomas, 1958. Stygophile, recorded from
the Pionerskaya (= Ushchel’naya) Cave,
Khostinsky City District, Greater Sochi,
Krasnodar Province (Chibisova, 1967).
– Difflugia penardi Hopkinson, 1909. Stygophile, recorded from the Labirintovaya
Cave, Khostinsky City District, Greater
Sochi, Krasnodar Province (Chibisova,
1967).
– Difflugia oblonga Ehrenberg, 1838. Stygophile, recorded from the Labirintovaya
Cave, Khostinsky City District, Greater
Sochi, Krasnodar Province (Chibisova,
1967).
– Pontigulasia incisa Rhumbler, 1896. Stygophile, recorded from the Labirintovaya
Subterranean biota of European Russia
Cave, Khostinsky City District, Greater
Sochi, Krasnodar Province (Chibisova,
1967).
Family Heleoperidae Jung, 1942
– Heleopera sphagni Leidy, 1874 (= Heleopera picta Leidy, 1879). Likely stygophile, recorded from the Labirintovaya Cave,
Khostinsky City District, Greater Sochi,
Krasnodar Province (Chibisova, 1967).
Subphylum Conosa Cavalier-Smith, 1998
Infraphylum Mycetozoa de Bary, 1859
Class Myxogastria Macbride, 1899
Order Physarida Macbride, 1922
Family Physaridae Rostafinski, 1873
– Physarum flagellatum (Alexeieff, 1923)
(= Hyperamoeba flagellata Alexeieff,
1923). Likely stygophile, recorded from
the Labirintovaya Cave, Khosta District,
Greater Sochi, Krasnodar Province (Myl’nikov et al., 2006).
Kingdom Animalia Linnaeus, 1758
Phylum Platyhelminthes Claus, 1887
Class Turbellaria Ehrenberg, 1831
Order Tricladida Lang, 1884
Unidentified unpigmented planarians have
long been recorded from the Bolshaya
Vorontsovskaya (= Vorontsovskaya) Cave,
Khostinsky City District, Greater Sochi,
Krasnodar Province (Birstein, 1950). Recorded from a tectonic cave on the Psebe
River, Tuapse District, as well as the Kirovskaya (Lazarevsky City District, Greater
Sochi) and Labirintovaya caves (Khostinsky City District, Greater Sochi), all in
Krasnodar Province (Ljovuschkin, 1966).
Family Dugesiidae Ball, 1974
– Dugesia taurocaucasica (Livanov, 1951).
Stygophile, recorded from a cave on Mt
Armovka, Tryu-Yatyrgvarta Massif, Mostovskoy District, as well as from the Fanagoriyskaya Cave near Goryachiy Klyuch, both
Krasnodar Province (Shumeev, 2008).
Family Dendrocoelidae Hallez, 1892
– Dendrocoelidae gen. sp. Likely stygobiont, closer unidentified, recorded from a
spring inside the Anglo-Russkaya Cave on
Mt Pshekha-Su, Maikopsky District, Adygea Republic (Shumeev, 2008).
183
– Dendrocoelum sp. 1. Stygophile, an undescribed species from the running waters
of the Tryu-Yatyrgvarta Massif, including a
spring inside a cave on Mt Armovka, Mostovskoy District, Krasnodar Province
(Shumeev, 2008).
– Dendrocoelum sp. 2. Stygobiont, an undescribed species from the running waters
inside the Fanagoriyskaya Cave near Goryachiy Klyuch, Krasnodar Province
(Shumeev, 2008).
Phylum Nematoda Diesing, 1861
Uncertain species of nematodes have been
recorded from the Pionerskaya (= Ushchel’naya) Cave, Khostinsky City District, Greater Sochi, Krasnodar Province (Ljovuschkin, 1966).
Class Enoplea Inglis, 1983
Order Monhysterida Filipjev, 1929
Family Monhysteridae De Man, 1876
– Monchystera paludicola De Man, 1880.
Troglophile, recorded from the Labirintovaya (= Novaya) Cave, Greater Sochi,
Krasnodar Province (Ljovuschkin, 1966).
Family Actinolaimidae Thorne, 1939
– Trachactinolaimus radulatus Andrássy,
1963. Troglophile, recorded from the Tonnel Kuzmenko Cave, Khostinsky City District, Greater Sochi, Krasnodar Province
(Ljovuschkin, 1966).
Family Dorylaimidae De Man, 1876
– Dorylaimus stagnalis Dujardin, 1845.
Troglophile, recorded from the Bolshaya
Kazachebrodskaya Cave, Adlersky City
District, Greater Sochi, Krasnodar Province (Ljovuschkin, 1966).
– Dorylaimus callosus Skwarra, 1921. Troglophile, recorded from the karst waters on
the Agura River, Greater Sochi, Krasnodar
Province (Ljovuschkin, 1966).
– Eudorylaimus laticollis (De Man, 1907).
Troglophile, recorded from the Labirintovaya (= Novaya) Cave, Greater Sochi,
Krasnodar Province (Ljovuschkin, 1966).
– Eudorylaimus condamni (Vanha, 1893).
Troglophile, recorded from the Dolgaya
Cave, Khostinsky City District, Greater
Sochi, Krasnodar Province (Ljovuschkin,
1966).
184
S.I. Golovatch et al.
– Eudorylaimus iners (Bastian, 1865). Troglophile, recorded from the Vorontsovskaya Cave, Khostinsky City District, Greater
Sochi, Krasnodar Province (Ljovuschkin,
1966).
Order Mononchida Jairajpuri, 1969
Family Mononchidae Chitwood et Chitwood,
1937
– Mononchus truncatus Bastian, 1865. Troglophile, recorded from the Zapovednaya
(= Podparapetnaya) Cave, Khostinsky City
District, Greater Sochi, Krasnodar Province (Ljovuschkin, 1966).
– Prionchulus muscorum (Dujardin, 1845).
Troglophile, recorded from the Dolgaya
Cave, Khostinsky City District, Greater Sochi, Krasnodar Province (Ljovuschkin,
1966).
Order Triplonchida Cobb, 1920
Family Tripylidae (De Man, 1876)
– Tripyla papillata Bütschli, 1873. Troglophile, recorded from the karst waters on the
Agura River, Greater Sochi, Krasnodar
Province (Ljovuschkin, 1966).
– Tripyla filicaudata De Man, 1880. Troglophile, recorded from the Krasnoaleksandrovskaya, Labirintovaya (= Novaya) and
Bolshaya Kazachebrodskaya caves, Greater Sochi, Krasnodar Province (Ljovuschkin, 1966).
Family Prismatolaimidae Micoletzky, 1922
– Prismatolaimus dolichurus De Man, 1880.
Troglophile, recorded from the Labirintovaya (= Novaya) Cave, Greater Sochi,
Krasnodar Province (Ljovuschkin, 1966).
Class Secernentea Lorenzen, 1981
Order Tylenchida Thorne, 1949
Family Criconematidae Taylor, 1936
– Criconema fimbriatum Cobb in Taylor,
1936. Troglophile, recorded from the
Krasnoaleksandrovskaya and Pionerskaya
(= Ushchel’naya) caves, Greater Sochi,
Krasnodar Province (Ljovuschkin, 1966).
– Criconema sp. Troglophile, recorded from
the Fanagoriyskaya Cave near Goryachiy
Klyuch, Krasnodar Province (Ljovuschkin,
1966).
– Criconemoides sp. Troglophile, recorded
from the Labirintovaya (= Novaya) Cave,
Greater Sochi, Krasnodar Province (Ljovuschkin, 1966).
Phylum Annelida Lamarck, 1809
Class Hirudinea Lamarck, 1818
Order Arhynchobdellida Blanchard, 1894
Family Erpobdellidae Blanchard, 1894
– Erpobdella octoculata (Linnaeus, 1758),
trogloxene, recorded by Birstein and Ljovuschkin (1967a) from the Fanagoriyskaya
Cave near Goryachiy Klyuch, Krasnodar
Province; also reported from several Balkan caves (Montenegro, Republic of Macedonia, Greek Macedonia).
Class Oligochaeta Grube, 1850
Order Crassiclitellata Jamieson, 1988
Family Lumbricidae Claus, 1876
– Allolobophora cavatica Michaelsen,
1910. Troglophile, described from a cave at
the Khodz’ River, Adygea Republic or Krasnodar Province. Also recorded from epigean
habitats in Transcaucasia (Lenkoran, Azerbaijan) (Michaelsen, 1910).
– Dendrobaena veneta crassa (Malevics,
1947). Troglophile, described as a separate
form from the Vorontsovskaya Cave,
Khostinsky City District, Greater Sochi,
Krasnodar Province (Malevich, 1947).
– Eisenia foetida (Savigny, 1826). Troglophile, recorded from the Podzemnaya Khosta (in the publication of Malevich, it was
referred to as Khostinskaya Mokraya) and
Bolshaya Kazachebrodskaya (= VerkhneMzymtinskaya, Mzymtinskaya) caves, both
in Greater Sochi, Krasnodar Province (Malevich, 1947).
Order Enchytraeida Vejdovský, 1879
Family Enchytraeidae Vejdovský, 1879
Uncertain enchytraeid species have been
recorded from the Fanagoriyskaya, Zapovednaya (= Podparapetnaya), Labirintovaya (= Novaya) and Vorontsovskaya
caves, all Krasnodar Province (Ljovuschkin, 1966).
Order Tubificida Brinkhurst, 1982
Family Naididae Ehrenberg, 1828
– Homochaeta sp. Troglophile, recorded
from the Zapovednaya (= Podparapetnaya)
Cave, Khostinsky City District, Greater
Subterranean biota of European Russia
Sochi, Krasnodar Province (Ljovuschkin,
1966).
Peloscolex sp. Troglophile, recorded from
the Labirintovaya (= Novaya) Cave, Greater Sochi, Krasnodar Province (Ljovuschkin, 1966).
Phylum Mollusca Linnaeus, 1758
Class Gastropoda Cuvier, 1797
Order Littorinimorpha Golikov et Starobogatov, 1975
Family Hydrobiidae Stimpson, 1865
– Belgrandiella caucasica Starobogatov,
1962 (= B. nemethi Schütt in Schütt et
Șeșen, 1993). Stygo- and crenobiont, described from a spring inside the Krasnoaleksandrovskaya (= Peshchera Ved’m) Cave,
Lazarevsky City District, Greater Sochi,
Krasnodar Province (Starobogatov, 1962).
Inhabiting cave (e.g. the Dolgaya Cave)
and creek waters along the Black Sea coast
between Tuapse and Psou (Starobogatov,
1962; Schütt, Șeșen, 1993; Palatov, Vinarski, 2015; Vinarski, Kantor, 2016).
– “Geyeria” valvataeformis Starobogatov,
1962. Stygobiont, known only from the
type locality, a spring inside the Krasnoaleksandrovskaya (= Peshchera Ved’m) Cave,
Lazarevsky City District, Greater Sochi,
Krasnodar Province (Starobogatov, 1962;
Vinarski, Kantor, 2016).
– “Lartetia” sp. Recorded from the Bolshaya
Kazachebrodskaya (= Verkhne-Mzymtinskaya) Cave, Adlersky City District, Greater Sochi, Krasnodar Province (Birstein,
1950), but actually it seems to concern one
of the Transcaucasian species of Paladilhiopsis (Turbanov et al., 2016a).
– Paladilhiopsis orientalis Starobogatov,
1962. Stygobiont, known only from the
type locality, a spring inside the Krasnoaleksandrovskaya (= Peshchera Ved’m) Cave,
Lazarevsky City District, Greater Sochi,
Krasnodar Province (Starobogatov, 1962;
Vinarski, Kantor, 2016).
– Paladilhiopsis pulcherrima Starobogatov, 1962. Stygo- and crenobiont, described
from a spring inside the Krasnoaleksandrovskaya (= Peshchera Ved’m) Cave, Laz-
185
arevsky City District, Greater Sochi, Krasnodar Province (Starobogatov, 1962). Inhabiting also helo- and rheocrens along the
Black Sea coast between Tuapse and Psou
(Starobogatov, 1962; Palatov, Vinarski,
2015; Vinarski, Kantor, 2016).
– Paladilhiopsis subovata Starobogatov,
1962. Stygobiont, known only from the
type locality, a spring inside the Krasnoaleksandrovskaya (= Peshchera Ved’m) Cave,
Lazarevsky City District, Greater Sochi,
Krasnodar Province (Starobogatov, 1962;
Vinarski, Kantor, 2016).
Order Geophila Ferrusac, 1821
Family Orculidae Steenberg, 1925
– Lauria sp. Likely troglobiont, recorded
from the Shirokopokosskaya and Malaya
Kazachebrodskaya caves, Greater Sochi,
Krasnodar Province (Birstein, 1950; Ljovuschkin, 1966).
Family Trigonochlamydidae Hesse, 1882
– Troglolestes sokolovi Ljovuschkin and
Matiokin, 1965. Troglobiont, known only
from the type locality, the Vorontsovskaya
Cave, Khostinsky City District, Greater
Sochi, Krasnodar Province (Ljovuschkin,
Matiokin, 1965).
Family Orculidae Steenberg, 1925
– Euxinolauria vitrea (Schileyko, 1988).
Likely troglobiont, described from the
Bolshaya Kazachebrodskaya (= VerkhneMzyimtinskaya) Cave near Mzymta River
(Schileyko 1988), also recorded from the
nearby Malaya Kazachebrodskaya (=Nizhne-Mzymtinskaya, Akhshtyrskaya) Cave,
Adlersky City District, Greater Sochi, Krasnodar Province (Egorov, Greke, 2005).
Family Clausiliidae Gray, 1855
– Scrobifera taurica (L. Pfeiffer, 1848).
Troglophile, recorded from the Shirokopokosskaya Cave, Khostinsky City District,
Greater Sochi, Krasnodar Province, previously referred to as Clausilia foveicollis
Parr. (Birstein, 1950).
– Serrulina sp. Troglophile, recorded from
the Beloskalskaya Cave, as well as a cave
on the Agura River, Greater Sochi, Krasnodar Province (Ljovuschkin, 1966).
186
S.I. Golovatch et al.
Family Zonitidae Morch, 1864
– Conulopolita cavatica (Riedel, 1966).
Troglobiont, described from the Shirokopokosskaya Cave (Khostinsky City District), also known from the nearby Bolshaya
Kazachebrodskaya and Malaya Kazachebrodskaya caves (Adlersky City District),
as well as the Chortova Nora Cave (Khostinsky City District), all in Greater Sochi,
Krasnodar Province (Birstein, 1950; Ljovuschkin, 1966; Riedel, 1966; Koval,
2004a).
– Oxychilus (Longiphallus) sp. Likely troglophile, recorded from the Shakal’ya, Chortova Nora, Bolshaya Kazachebrodskaya,
Malaya Kazachebrodskaya, Beloskalskaya,
Shirokopokosskaya (= Bozhyey Materi) and
Vorontsovskaya caves, all in Greater Sochi, Krasnodar Province (Birstein, 1950;
Ljovuschkin, 1966; Koval, 2004a).
Class Bivalvia Linnaeus, 1758
Order Veneroida Adams et Adams, 1856
Family Euglesidae Pirogov et Starobogatov,
1974
– Euglesa (s. str.) personata (Malm, 1855).
Likely stygophile, widespread in Europe, the
Caucasus and Asia Minor (Kantor et al.,
2010; Vinarski, Kantor, 2016), recorded from
wells and springs near Adler, Greater Sochi,
Krasnodar Province (Starobogatov, 1962).
– Euglesa (s. str.) cavatica (Shadin, 1952).
Stygobiont, described from a spring inside
the Pionerskaya (= Ushchel’naya) Cave in
the upper reaches of Vostochnaya (=
Bolshaya) Khosta River, Khostinsky City
District, Greater Sochi, Krasnodar Province. Reported also from caves in Abkhazia
(Starobogatov, 1962; Chertoprud et al.,
2016; Vinarski, Kantor, 2016, Turbanov et
al., 2016a), but with some minor differences from the typical form.
Phylum Arthropoda Siebold, 1848
Subphylum Crustacea Brünnich, 1772
Class Copepoda Milne-Edwards, 1840
Order Cyclopoida Burmeister, 1834
Family Cyclopidae Rafinesque, 1815
– Acanthocyclops venustus stammeri (Kiefer, 1930). Stygobiont, interstitial-dweller.
Recorded from the imterstitial of Psezuapse River, Lazarevsky City District, Greater Sochi, and in the interstitial of Djubga
River near Djubga, Tuapse District, both in
the Krasnodar Province (Monchenko,
1984).
– Diacyclops bisetosus (Rehberg, 1880).
Stygophile, recorded from the Bolshaya
Kazachebrodskaya Cave, Adlersky City
District, Greater Sochi, Krasnodar Province (Ljovuschkin, 1966).
– Paracyclops fimbriatus imminutus Kiefer, 1929. Stygophile, recorded from a karst
spring on the Vostochnaya (= Bolshaya)
Khosta River and from the Vorontsovskaya
Cave, Khostinsky City District, Greater
Sochi, Krasnodar Province (Borutzky,
1950a; Ljovuschkin, 1966).
– Speocyclops cinctus Monchenko, 1983.
Interstitial of Alibek River, KarachaevoCherkessia (Monchenko, 1983).
– Speocyclops demetiensis (Scourfield,
1932). Stygobiont, interstitial-dweller. Recorded from the imterstitial of Psezuapse,
Khosta, Mzymta and Psou rivers in the
Krasnodar Province (Monchenko, 1986).
– Speocyclops lussianus Borutzky, 1950.
Stygobiont, described from the Vorontsovskaya and Pionerskaya (= Ushchel’naya)
caves, Khostinsky City District, Greater
Sochi, Krasnodar Province (Borutzky,
1950a; Ljovuschkin, 1966).
– Speocyclops psezuapsensis Borutzky,
1965. Sygobiont, described from the
Kirovskaya Cave, Psezuapse River, Lazarevsky City District, Greater Sochi, Krasnodar Province (Borutzky, 1965).
Order Harpacticoida G.O. Sars, 1903
Family Ameiridae Boeck, 1865
– Nitocrella hirta caucasica Borutzky,
1967. Stygobiont, described from the Zapovednaya (= Podparapetnaya) Cave, Khostinsky City District, Greater Sochi, Krasnodar
Province (Borutzky, 1967a, 1972b).
– Megastygonitocrella ljovuschkini
(Borutzky, 1967). Stygobiont, described or
reported from the Dolgaya and Pionerskaya
(= Ushchel’naya) caves near Sochi, and
Subterranean biota of European Russia
from the Krasnoaleksandrovskaya Cave
near Lazarevskoe, all in the Krasnodar Province (Borutzky, 1967a, 1972b).
Family Canthocamptidae Brady, 1880
– Attheyella crassa (G.O. Sars, 1863). Stygophile, recorded from the Fanagoriyskaya
Cave near Goryachiy Klyuch, from the Krasnoaleksandrovskaya Cave (Lazarevsky City
District) and from the Pionerskaya (=
Ushchel’naya), Labirintovaya and Dolgaya
caves, Khostinsky City District, Greater
Sochi, all in the Krasnodar Province (Borutzky, 1948a, 1972b).
– Bryocamptus aquaeductus Borutzky,
1934. Stygobiont, described from a water
pipeline in the town of Teberda, Karachaevo-Chekessia (Borutzky, 1934).
– Bryocamptus innominatus Borutzky,
1940. Stygobiont, described from the Pionerskaya (= Ushchel’naya) Cave, Khostinsky City District, Greater Sochi, Krasnodar
Province (Borutzky, Mikhailova-Neikova,
1970; Borutzky, 1972b).
– Bryocamptus pygmaeus (G.O. Sars, 1863).
Stygophile, recorded from the Dolgaya and
Labirintovaya caves, Khostinsky City District, Greater Sochi, Krasnodar Province
(Borutzky, 1972b).
– Bryocamptus tarnogradskyi Borutzky,
1934. Stygophile, recorded from the
Fanagoriyskaya Cave near Goryachiy
Klyuch, from the Krasnoaleksandrovskaya
Cave near Lazarevskoe, and from the Labirintovaya, Podparapetnaya (= Zapovednaya) and Tonnel Kuzmenko caves near
Khosta, Greater Sochi, all in the Krasnodar Province (Borutzky, 1948a, 1972b;
Ljovuschkin, 1966).
– Bryocamptus zschokkei caucasicus
Borutzky, 1930. Stygophile, recorded from
the Bolshaya Kazachebrodskaya (= Verkhne-Mzymtinskaya), Pionerskaya (= Ushchel’naya), Labirintovaya and Dolgaya caves
near Sochi, from the Krasnoaleksandrovskaya Cave near Lazarevskoe, and from
the Fanagoriyskaya Cave near Goryachiy
Klyuch, all in the Krasnodar Province
(Borutzky, 1948a, 1972b).
187
– Bryocamptus hostensis Borutzky, 1972.
Likely stygobiont, described from the Labirintovaya Cave, Khostinsky City District,
Greater Sochi, Krasnodar Province (Borutzky, 1972b).
– Bryocamptus sp. The first copepodid instar recorded from the Vorontsovskaya Cave,
Khostinsky City District, Greater Sochi,
Krasnodar Province (Borutzky, 1952).
– Canthocamptus staphylinus (Jurine,
1820). Stygophile, recorded from the karst
waters on the Agura River, Greater Sochi,
Krasnodar Province (Ljovuschkin, 1966).
– Pilocamptus georgevitchi (Chappuis,
1924). Stygobiont, recorded from the Krasnoaleksandrovskaya Cave (Lazarevsky City
District), from the Dolgaya, Labirintovaya
and Nikity caves, Greater Sochi, Krasnodar
Province (Borutzky, Mikhailova-Neikova,
1970; Borutzky 1972b).
– Elaphoidella bidens coronata (G.O. Sars,
1904). Stygophile, discovered in the interstitial (Chappuis pit) of Mzymta River at
Adler, Greater Sochi, Krasnodar Province
(Borutzky, 1972b).
– Elaphoidella czerkessica Borutzky, 1972.
Stygobiont, described from the Pionerskaya
(= Ushchel’naya) Cave, Khostinsky City
District, Greater Sochi, Krasnodar Province (Borutzky, 1972b).
– Moraria operculata Borutzky, 1948. Stygobiont, described from the Bolshaya Kazachebrodskaya (= Verkhne-Mzymtinskaya) and Dolgaya caves, as well as from
the interstitial (Chappuis pit) at a bank of the
Mzymta River, all in Greater Sochi, Krasnodar Province (Borutzky, 1948a, 1972b).
– Moraria hostensis Borutzky, 1972. Stygobiont, described from the Zapovednaya
(= Podparapetnaya) Cave, Khostinsky City
District, Greater Sochi, Krasnodar Province (Borutzky, 1972b).
– Moraria poppei (Mrazek, 1893). Stygobiont, recorded from the Dolgaya Cave,
Khostinsky City District, Greater Sochi,
Krasnodar Province (Borutzky, 1972b).
– Moraria varica (Graeter, 1910). Stygobiont, recorded from the Fanagoriyskaya
188
S.I. Golovatch et al.
Cave near Goryachiy Klyuch and from the
Labitintovaya Cave, Khostinsky City District, Greater Sochi, both in the Krasnodar
Province (Borutzky, 1972b).
– Moraria sp. Nauplia recorded from the
Dolgaya Cave, Khostinsky City District,
Greater Sochi, Krasnodar Province (Borutzky, 1972b).
Family Parastenocarididae Chappuis, 1940
– Parastenocaris tenuis Borutzky, 1948.
Stygobiont, described from the Fanagoriyskaya Cave near Goryachiy Klyuch, Krasnodar Province (Borutzky, 1948a, 1972b).
– Parastenocaris sp. Stygobiont, recorded
from the interstitial waters on the banks of
Mzymta River (Ljovuschkin, 1966).
Class Ostracoda Latreille, 1802
Order Podocopida G.O. Sars, 1866
Family Candonidae Kaufmann, 1900
– Trapezicandona ljovuschkini (Rudjakov,
1963). Stygobiont, described and still known
only from the Krasnoaleksandrovskaya (=
Peshchera Ved’m) Cave, Lazarevsky City
District, Greater Sochi, Krasnodar Province (Rudjakov, 1963).
– Candona sp. Recorded without an exact
species identification from the Krasnoaleksandrovskaya (= Peshchera Ved’m) Cave
(Lazarevsky City District) and the Labirintovaya Cave (Khostinsky City District), both
in Greater Sochi, Krasnodar Province (Rudjakov, 1963).
Family Cyprididae Baird, 1845
– Cypria reptans Bronstein, 1928. Crenobiont, found in the Dolgaya Cave, Khostinsky City District, Greater Sochi, Krasnodar
Province (Rudjakov, 1963).
Family Limnocytheridae Klie, 1938
– Kovalevskiella (Cordocythere) rudjakovi
(Danielopol, 1969). Stygobiont, described
and still known only from the Krasnoaleksandrovskaya (= Peshchera Ved’m) Cave,
Lazarevsky City District, Greater Sochi,
Krasnodar Province (Danielopol, 1969).
Earlier referrred to as Metacypris sp. (Rudjakov, 1963).
Class Malacostraca Latreille, 1817
Superorder Syncarida Packard, 1885
Order Bathynellacea Chappuis, 1915
Family Bathynellidae Grobbon, 1904
– Antrobathynella stammeri ciscaucasica
(Birstein et Ljovuschkin, 1964). Stygobiont, interstitial-dweller, described from
a spring in the Fanagoriyskaya Cave near
Goryachiy Klyuch, Krasnodar Province
(Birstein, Ljovuschkin, 1964, 1967b; Serban, 1993).
– Bathynella natans natans Vejdovsky,
1882. Stygobiont, interstitial-dweller, discovered in a waterbody inside the Pionerskaya (= Ushchel’naya) Cave, upper reaches of Vostochnaya (= Bolshaya) Khosta
River, Greater Sochi, Krasnodar Province
(Birstein, Ljovuschkin, 1967b).
Order Isopoda Latreille, 1817
Suborder Asellota Latreille, 1803
Family Asellidae Latreille 1803, sensu
Rafinesque-Schmaltz, 1815
– Proasellus linearis (Birstein, 1967). Stygobiont, described from a spring at Evstafieva Shchel’ near Gelendjik, Krasnodar
Province (Birstein, 1967). There are also
records of P. cf. linearis from near Tuapse,
in springs and oozing waters in the valleys
of Agoy and Nebug rivers, as well as in
streams of Kadosh Park where the animals
are washed out from subterranean waters
during strong rain flashes (Sokolova, Palatov, 2015).
– Proasellus ljovuschkini (Birstein, 1967).
Stygobiont, described from a karst vaucluse in the lower reaches of Khosta River,
Greater Sochi, Krasnodar Province (Birstein, 1967).
– Proasellus similis (Birstein, 1967). Stygobiont, described from a waterbody inside
the Nikity Cave in the Psakho River basin,
a tributary of Kudepsta River, Greater Sochi, Krasnodar Province (Birstein, 1967).
– Proasellus sp. (Fig. 5A). Stygobiont, possibly a new species, recorded from the Avgust Cave, Adlersky City District, Greater
Sochi, Krasnodar Province (Turbanov et
al., 2016a).
Family Microparasellidae Karaman, 1933
– Microcharon tantalus Birstein et Ljovuschkin, 1965. Stygobiont, described from
Subterranean biota of European Russia
189
Fig. 5. Some stygo- or troglobiont crustaceans in nature (all caves from Greater Sochi, Krasnodar Province).
A — Proasellus sp., a stygobiont water louse from the Avgust Cave; B — Ligidium cavaticum Borutzky,
1950, a troglobiont wood louse from the Giganov Cave; C — Niphargus sp., a stygobiont amphipod from
the Gigantov Cave; D — Troglocaris (Xyphocaridinella) jusbaschjani Birstein, 1948, a stygobiont shrimp
from sulphurous springs on the Agura River. All pictures by Ilya Turbanov.
Рис. 5. Некоторые стиго- и троглобионтные ракообразные в природе (все пещеры из Большого Сочи,
Краснодарский край). A — Proasellus sp., стигобионтный водяной ослик из пещеры Август; B —
Ligidium cavaticum Borutzky, 1950, троглобионтная мокрица из пещеры Гигантов; C — Niphargus sp.,
стигобионтный бокоплав из пещеры Гигантов; D — Troglocaris (Xyphocaridinella) jusbaschjani Birstein,
1948, стигобионтная креветка из серных ручьев у реки Агура. Все фотографии Ильи Турбанова.
the interstitial of Mzymta River, Greater
Sochi, Krasnodar Province (Birstein, Ljovuschkin, 1965).
Suborder Oniscidea Latreille, 1802
Family Ligiidae Brandt, 1883
– Ligidium zaitzevi Borutzky, 1950. Troglophile, described from a series of syntypes stemming from the Vorontsovskaya
Cave, Khostinsky City District, Greater
Sochi, Krasnodar Province, as well as the
Mikhailovskaya Cave and the cave near
Venetian Bridge, both latter near Sukhum,
Abkhazia (Borutzky, 1950b, 1972a).
– Ligidium hypnorum (Cuvier, 1792). Troglophile, recorded for the Shirokopokosskaya Cave, Greater Sochi, Krasnodar
Province (Borutzky, 1950b, 1972a).
– Ligidium cavaticum Borutzky, 1950 (Fig.
5B). Troglobiont, described from a series
of syntypes coming from the Pervomayskaya, Podzemnaya Khosta, Shirokopokosskaya (= Bozhyey Materi), Vorontsovskaya,
Beloskalskaya, Navalishinskaya, Partizanskaya, Pionerskaya (= Ushchel’naya), Zapovednaya (= Podparapetnaya), Malaya Kazachebrodskaya, Bolshaya Kazachebrod-
190
S.I. Golovatch et al.
skaya, Kamenskaya and Giganov (new data)
caves, all in Greater Sochi, Krasnodar Province (Borutzky, 1950b, 1972a; Ljovuschkin, 1966).
Family Buddelundiellidae Verhoeff, 1930
– Pseudobuddelundiella hostensis Borutzky, 1967. Troglobiont, described from the
Labirintovaya and Pionerskaya (= Ushchel’naya) caves, Greater Sochi, Krasnodar Province (Borutzky, 1967b, 1972a).
– Pseudobuddelundiella ljovuschkini
Borutzky, 1967. Troglobiont, described
from the Kirovskaya Cave, Lazarevsky City
District, Psezuapse River, Krasnodar Province (Borutzky, 1967b, 1972a).
Family Trichoniscidae Sars, 1899
– Caucasocyphonethes cavaticus Borutzky, 1948. Troglobiont, represented by 4
subspecies from caves on the Mzymta, Khosta and Kudepsta rivers, Greater Sochi, Krasnodar Province: C. c. cavaticus Borutzky,
1948 from the Malaya Kazachebrodskaya
Cave; C. c. chostensis Borutzky, 1948 from
the Podzemnaya Khosta and Beloskalskaya
caves; C. c. msymticus Borutzky, 1948 from
the Bolshaya Kazachebrodskaya Cave; and
C. c. adlerensis Borutzky, 1948 from the
Shirokopokosskaya and Nikity caves (Borutzky, 1948b, 1972a).
– Psachonethes czerkessicus Borutzky,
1969. A troglomorphic species described
from a cave on the Psakho River, a tributary
of Kudepsta River, Greater Sochi, Krasnodar Province (Borutzky, 1969c, 1972a).
– Trichoniscus pygmaeus tuapsensis Borutzky, 1972. Troglobiont, described from
a cave on the Psebe River, Tuapse District,
Krasnodar Province (Borutzky, 1972a).
– Trichoniscus sp. A record is available of
a troglomorphic woodlouse from the Shirokopokosskaya Cave, Khostinsky City
District, Greater Sochi, Krasnodar Province (Borutzky, 1972a).
Family Cylisticidae Verhoeff, 1949
– Cylisticus birsteini Borutzky, 1961. Troglophile, described from a cave on the Psakho River, a tributary of Kudepsta River,
recorded from the Navalishinskaya (= Mu-
zeynaya) and Podzemnaya Khosta (=
Khostinskaya Mokraya) caves, Greater
Sochi, Krasnodar Province (Borutzky, 1961,
1972a; Ljovuschkin, 1966).
Order Amphipoda Latreille, 1816
Family Crangonyctidae Bousfield, 1973
– Lyurella shepsiensis Sidorov, 2015. Stygo- and crenobiont, described from springs
in the Shepsi River basin (Sidorov 2015),
also recorded in springs of the Ashe River
basin near Kalezh and in the Krasnoaleksandrovskaya Cave (Lazarevsky City District), all in the Krasnodar Province (Sokolova, Palatov, 2015).
Family Niphargidae Bousfield, 1977
– Niphargus abchasicus Martynov, 1932.
Stygophile, described from springs in the
interfluve of Khosta and Kudepsta rivers,
Greater Sochi, Krasnodar Province (Ìartynov, 1932), also recorded there later (Ljovuschkin 1963), as well as from springs
within and near Sochi (Derzhavin, 1945).
– Niphargus caelestis G.S. Karaman, 1982
(= N. stygius longidactylus Birstein, 1952).
Stygobiont, described from the Pionerskaya
(= Ushchel’naya) Cave, Greater Sochi, Krasnodar Province (Birstein, 1952). According to the Code of Zoological Nomenclature the original name was changed since
“longidactylus” had been preoccupied by
N. kochianus longidactylus Ruffo, 1937,
from springs at Verona, Italy (Karaman,
1982).
– Niphargus cubanicus Birstein, 1954.
Likely stygophile, described from a wellwarmed fish pond at Goryachiy Klyuch,
Krasnodar Province (Birstein, 1954).
– Niphargus krasnodarus Karaman, 2012.
Stygobiont, described from the Fanagoriyskaya Cave near Goryachiy Klyuch, Krasnodar Province (Karaman, 2012).
– Niphargus latimanus Birstein, 1952 (= N.
stygius latimanus Birstein, 1952). Stygobiont, described from a stream in the Vorontsovskaya Cave, Greater Sochi, Krasnodar
Province (Birstein, 1952). Morphologically similar forms recorded also from springs
near Tuapse (Sokolova, Palatov, 2015).
Subterranean biota of European Russia
– Niphargus cf. magnus Birstein, 1940.
Stygobiont, a possibly new species morphologically similar to N. magnus, found in
springs near Tuapse, Krasnodar Province
(Sokolova, Palatov, 2015).
– Niphargus potamophilus Birstein, 1954.
Likely stygophile, described from experimental fish ponds in the mouth of Don River
near the city of Rostov-on-Don, Rostov
Region, as well as in Akhtyrskie fish ponds
in the mouth of Kuban River, Abinsk District, Krasnodar Province (Birstein, 1954).
– Niphargus pseudolatimanus Birstein,
1952 (= N. stygius pseudolatimanus Birstein, 1952). Stygobiont, described from a
stream inside the Labirintovaya (= Novaya)
Cave, Greater Sochi, Krasnodar Province
(Birstein, 1952).
– Niphargus smirnovi Birstein, 1952. Stygophile, described from the Bolshaya Kazachebrodskaya (= Verkhne-Mzymtinskaya) Cave in the valley of Mzymta River,
Greater Sochi, Krasnodar Province (Birstein, 1952). Subsequently recorded in
springs of Khosta, Kudepsta and Psou rivers, as well as in the Vorontsovskaya, Labirintovaya, Dolgaya, Pionerskaya (=
Ushchel’naya) and Podzemnaya Khosta
caves in the same region. The species is
thereby fairly variable, each river basin and
cave system supporting its own unique
morphotype. Perhaps we face a group of
closely related species (Ljovuschkin, 1963).
– Niphargus submersus (Derzhavin, 1945)
(= Martynovia submersa Derzhavin, 1945).
Stygophile, described from Sochinka River
under a bridge in the town of Greater Sochi,
Krasnodar Province (Derzhavin, 1945).
Subsequently recorded in springs in the
Vostochnaya (= Bolshaya) Khosta River
basin (Ljovuschkin, 1963).
– Niphargus sp. (Fig. 5C). Stygobiont, possibly a new species, recorded from the Gigantov Cave in the Alek Massif, Greater
Sochi, Krasnodar Province (new data).
Family Typhlogammaridae Bousfield, 1977
– Zenkevitchia sp. Stygobiont, a new, yet
undescribed species recorded in the Avgust
191
Cave, Adlersky City District, Greater Sochi, Krasnodar Province (Turbanov, Marin,
2015, 2017).
Order Decapoda Latreille, 1802
Infraorder Caridea Dana, 1852
Family Atyidae De Haan, 1849
– Troglocaris (Xyphocaridinella) jusbaschjani Birstein, 1948 (Fig. 5D). Described
and still known only from the type locality,
i.e. hydrogen sulfide springs on the Agura
River, Greater Sochi, Krasnodar Province
(Birstein, 1948; Marin, Sokolova, 2014).
Infraorder Brachyura Linnaeus, 1758
Family Potamidae Ortmann, 1896
– Potamon ibericum tauricum (Czerniavsky, 1884). Possibly troglophile, recorded
from the Chortova Nora Cave, Agura River,
Greater Sochi, Krasnodar Province (Koval,
2004a).
Class Arachnida Cuvier, 1812
Order Aranei Clerck, 1758
There seem to be no troglobitic spiders not
only in the Russian Caucasus, but across the entire
former Soviet Union (Turbanov et al., 2016b).
Family Agelenidae C.L. Koch, 1837
– Tegenaria abchasica Charitonov, 1941.
Troglophile, recorded from the Beloskal’skaya Cave, Khostinsky City District,
Greater Sochi, Krasnodar Province (Turbanov, Nadolny, 2017).
– Tegenaria pontica Charitonov, 1947. Troglophile, described from the Fanagoriyskaya
Cave near Goryachiy Klyuch, Krasnodar
Province (Charitonov, 1947).
– Tegenaria sp. Troglophile, a record is
available of a closer unidentified species
from the Chortova Nora Cave, Khostinsky
City District, Greater Sochi (Koval, 2004a).
Family Hahniidae Bertkau, 1878
– Iberina ljovuschkini Pichka, 1965. Troglophile, described from the Shakal’ya
Cave, Lazarevsky City District, Greater
Sochi, Krasnodar Province (Pichka, 1965).
Family Linyphiidae Blackwall, 1859
– Troglohyphantes birsteini Charitonov,
1947. Troglophile, described from the
Bolshaya Kazachebrodskaya and Malaya
Kazachebrodskaya caves, Greater Sochi,
192
S.I. Golovatch et al.
Krasnodar Province (Charitonov, 1947).
Further records concern the Chortova Nora,
Labirintovaya, Vorontsovskaya and Ametist caves, as well as a cave on Kamenka
River, all in the same region (Pichka, 1965;
Koval, 2004a; Turbanov, Nadolny, 2017).
Family Nesticidae Simon, 1894
– Aituaria pontica (Spassky, 1932) (= Nesticus ponticus Spassky, 1932). Troglophile,
described from a wine cellar at Khosta,
Greater Sochi, Krasnodar Province (Spassky
1932). Later recorded from the Bolshaya
Kazachebrodskaya, Chortova Nora and
Ametist caves in the same region, as well as
in a cave in the middle flow of Psakho River
and a karst pit on Agura River, both also in
the Krasnodar Province (Charitonov, 1947;
Pichka, 1965; Koval, 2004a; Turbanov,
Nadolny, 2017).
– Carpathonesticus birsteini (Charitonov,
1947). Troglophile, described from the
Podzemnaya Khosta (in the publication of
Charitonov it is indicated as Khostinskaya
Mokraya) Cave, Greater Sochi, Krasnodar
Province (Charitonov, 1947). Subsequently found in the Akhunskaya, Beloskal’skaya,
Vorontsovskaya, Labirintovaya (= Novaya),
Kamenskaya and Dolgaya caves, as well as
in a cave on the Agura River in the same
region (Pichka, 1965; Ljovuschkin, 1966;
Koval, 2004a, b; Turbanov, Nadolny, 2017).
– Carpathonesticus ljovuschkini (Pichka,
1965). Troglophile, described from the
Shakal’ya Cave, Lazarevsky City District,
Greater Sochi, Krasnodar Province (Pichka, 1965).
– Carpathonesticus zaitzevi (Charitonov,
1939). Troglophile, recorded from the Baribana Cave in the Alek Massif, Greater
Sochi, Krasnodar Province (Turbanov, Nadolny, 2017).
– Carpathonesticus sp. Troglophile, recorded from the Beloskal’skaya Cave, Greater
Sochi, Krasnodar Province (Turbanov,
Nadolny, 2017).
Family Pholcidae C.L. Koch, 1851
– Hoplopholcus longipes (Spassky, 1934).
Troglophile, described from a wine cellar at
Khosta, Greater Sochi, Krasnodar Province
(Spassky, 1934). Subsequently recorded in
the Bolshaya Kazachebrodskaya (= Verkhne-Mzymtinskaya), Shirokopokosskaya,
Beloskal’skaya and BS-2 caves, all in the
same region (Charitonov 1941; Turbanov,
Nadolny, 2017).
Family Tetragnathidae Menge, 1866
– Meta bourneti Simon, 1922. Troglophile,
recorded from the Chortova Nora Cave,
Khostinsky City District, Greater Sochi
(Koval, 2004a), widespread in Europe.
– Metellina merianae (Scopoli, 1763). Troglophile, recorded from the Fanagoriyskaya and Shirokopokosskaya caves in the
Krasnodar Province (Charitonov, 1947),
widespread in Europe.
Order Pseudoscorpiones De Geer, 1778
Family Neobisiidae Chamberlin, 1930
– Neobisium (Neobisium) speleophilum
Krumpal, 1986. Troglobiont, first recorded
without an exact species identification from
the Dolgaya and Pionerskaya caves in the
Vorontsovka Massif near Adler, Greater
Sochi, Krasnodar Province (Birstein, Ljovuschkin, 1960), later described (Krumpal,
1986).
– Neobisium sp. (Fig. 6A). Several records
are available from caves in the Krasnodar
Province: Shkol’naya and Gigantov (new
data) in the Alek Massif, Pechal’naya in the
Dzykhra Massif, and Dolgaya, Pionerskaya
(= Ushchel’naya), Labirintovaya (Khostinsky City District), Greater Sochi (Ljovus-
пещеры Гигантов; B — Nemaspela sokolovi (Ljovuschkin et Starobogatov, 1963), троглобионтный
сенокосец из пещеры Гигантов; C — Heterocaucaseuma feminaepectorum Antić et Makarov, 2016,
троглобионтная двупарноногая многоножка из пещеры Гигантов; D — Dolichopoda euxina Semenov,
1901, троглофильный кузнечик из пещеры Ахунская; E — Caucasaphaenops molchanovi Belousov,
1999, троглобионтный жук-жужелица из пещеры Гигантов; F — Seracamaurops komarovi Hlaváč,
Kodada et Koval, 1999, троглобионтный жук-ощупник из пещеры Гигантов. Все фотографии Ильи
Турбанова.
Subterranean biota of European Russia
193
Fig. 6. Some troglobiont or troglophile arthropods in nature (all caves from Greater Sochi, Krasnodar
Province). A — Neobisium sp., a troglobiont false scorpion from the Gigantov Cave; B — Nemaspela
sokolovi (Ljovuschkin et Starobogatov, 1963), a troglobiont harvestman from the Gigantov Cave; C —
Heterocaucaseuma feminaepectorum Antić et Makarov, 2016, a troglobiont millipede from the Gigantov
Cave; D — Dolichopoda euxina Semenov, 1901, a troglophile grasshopper from the Akhunskaya Cave; E —
Caucasaphaenops molchanovi Belousov, 1999, a troglobiont carabid beetle from the Gigantov Cave; F —
Seracamaurops komarovi Hlaváč, Kodada et Koval, 1999, a troglobiont pselaphid beetle from the Gigantov
Cave. All pictures by Ilya Turbanov.
Рис. 6. Некоторые троглобионтные и троглофильные членистоногие в природе (все пещеры из
Большого Сочи, Краснодарский край). A — Neobisium sp., троглобионтный ложноскорпион из
194
S.I. Golovatch et al.
chkin, 1966; Dashdamirov, Schawaller,
1992; Turbanov et al., 2016b).
Order Opiliones Sundevall, 1833
Family Nemastomatidae Simon, 1872
– Histricostoma caucasicum (Redikorzev,
1936). Likely troglophile, Caucasian endemic, recorded from the Fanagoriyskaya
Cave near Goryachiy Klyuch, Krasnodar
Province (Martens, 2006).
– Nemaspela abchasica (Ljovuschkin et
Starobogatov, 1963). Troglobiont, recorded from the Beloskal’skaya, Kolokol’naya
and Akhunskaya caves, Greater Sochi, Krasnodar Province (Martens, 2006; Chemeris,
2009).
– Nemaspela kovali Chemeris, 2009. Troglobiont, described from the Fontanka,
Omega-12, Omega-15 and Otte-Shik caves
in Kabardino-Balkaria (Chemeris, 2009).
– Nemaspela sokolovi (Ljovuschkin et Starobogatov, 1963) (Fig. 6B). Troglobiont,
known from a few caves in the Greater
Sochi region (Dolgaya, Labirintovaya,
Vorontsovskaya and Gigantov (new data)),
anophthalmic with elongated appendages
(Ljovuschkin, Starobogatov, 1963; Martens, 2006; Chemeris, 2009).
– Nemaspela sp. Troglobiont, without an
exact species identification recorded from
the Akhunskaya Cave, Greater Sochi, Krasnodar Province (Koval, 2004b).
Family Phalangiidae Latreille, 1802
– Nelima pontica Charitonov, 1941. Likely
troglophile, recorded from the Shirokopokosskaya, Bolshaya Kazachebrodskaya (=
Verkhne-Mzymtinskaya) and Malaya Kazachebrodskaya (= Nizhnyaya Kazachebrodskaya, Nizhne-Mzymtinskaya) caves,
Greater Sochi, Krasnodar Province (Charitonov, 1947; Ljovuschkin, Starobogatov,
1963; Ljovuschkin, 1966).
Subclass Acari Leach, 1817
Superorder Acariformes Zakhvatkin, 1952
Order Sarcoptiformes Reuter, 1909
Suborder Oribatida Van der Hammen, 1968
Family Phthiracaridae Perty, 1841
– Phthiracarus globosus (C.L. Koch, 1841).
Troglophile, recorded from the Voronts-
ovskaya Cave, Khostinsky City District,
Greater Sochi, Krasnodar Province (Ljovuschkin, 1966).
Superorder Parasitiformes Reuter, 1909
Order Mesostigmata Canestrini, 1891
Suborder Ixodoidea Leach, 1815
Family Ixodidae C.L. Koch, 1844
– Ixodes vespertilionis C.L. Koch, 1844. A
parasite of bats, recorded from the Pionerskaya (= Ushchel’naya) and Dolgaya caves,
Greater Sochi, Krasnodar Province (Ljovuschkin, 1966).
Subphylum Tracheata Haeckel, 1866
Class Chilopoda Latreille, 1817
Order Lithobiomorpha Pocock, 1895
Family Lithobiidae Newport, 1844
– Harpolithobius perplexus Zalesskaja,
1973. Troglophile, recorded from the Chortova Nora Cave, Greater Sochi, Krasnodar
Province (Koval, 2004a).
– Lithobius liber Lignau, 1903. Troglophile, recorded from the Chortova Nora
Cave, Greater Sochi, Krasnodar Province,
but perhaps belongs to a different species
(Volkova, 2015).
– Lithobius reconditus Zalesskaja, 1973.
Troglophile, recorded from the Pionerskaya
(= Ushchel’naya) Cave, Greater Sochi, Krasnodar Province (Zalesskaja, 1973a, b, 1978).
– Lithobius stuxbergii Sseliwanoff, 1881.
Troglophile, recorded from the Vorontsovskaya, Shirokopokosskaya and Chortova
Nora caves, all in Greater Sochi, Krasnodar
Province (Zalesskaja, 1963, 1973a, b, 1978;
Koval, 2004a).
Class Diplopoda de Blainville in Gervais, 1844
Order Glomerida Leach, 1814
Family Glomeridellidae Cook, 1896
– Typhloglomeris caucasica Golovatch,
1975. Troglophile, recorded from several
caves near Sochi, Krasnodar Province (Partizanskaya, Dolgaya, Vorontsovskaya etc.)
(Golovatch, 1975), but then found epigeically at Khosta (Golovatch, Chumachenko,
2013).
Family Glomeridae Leach, 1815
– Trachysphaera costata (Waga, 1857).
Troglophile, widespread across Central,
Subterranean biota of European Russia
Eastern and Southern Europe, as well as
Anatolia to northwestern Iran, recorded from
a cave in Crimea, from the Fanagoriyskaya
Cave near Goryachiy Klyuch, and from a
karst funnel at Shedok, both Krasnodar
Province (Golovatch, 1990).
– Trachysphaera minuta Golovatch, 1976.
Troglophile, known from two caves near
Sochi — Partizanskaya, Malaya Kazachebrodskaya (= Nizhne-Kazachebrodskaya),
as well as epigeically across much of the
Caucasus (Russia, Georgia and Azerbaijan)
(Golovatch, 1990).
– Hyleoglomeris sp. Troglophile or trogloxene, from a tectonic cave in the upper
basin of Psebe River, Tuapse District, Krasnodar Province (Golovatch, 1984–1985).
Order Chordeumatida Pocock, 1894
Family Anthroleucosomatidae Verhoeff,
1899
– Caucaseuma elephantum Antić et
Makarov, 2016. Troglobiont, known only
from the Yuzhnyi Slon Cave, Mt Dzentu, ca
1800 m a.s.l., Karachaevo-Cherkesskaya
Republic, northern Caucasus; body yellowish white, 11–12 ocelli in 3–4 rows (Antić,
Makarov, 2016).
– Caucaseuma fanagoriyskaya Antić et
Makarov, 2016. Troglobiont, known only
from the Fanagoriyskaya Cave near Goryachiy Klyuch, Krasnodar Province; body
greyish yellow with a brown head, 21–24
ocelli in 6 rows (Antić, Makarov, 2016).
– Caucaseuma lohmanderi Strasser, 1970.
Likely troglobiont, frequent in caves in the
western Caucasus (near Sochi, Krasnodar
Province): Baribana, Labirintovaya, Navalishinskaya (= Muzeynaya), Vorontsovskaya, and since the description, Ametist
Cave (Western Akhtsu Mts), Akhunskaya,
Partizanskaya, Dolgaya, Shirokopokosskaya (= Bozhyey Materi), Tisovaya (perhaps this is the Zapovednaya Cave) caves
(Strasser, 1970; Antić, Makarov, 2016).
– Caucaseuma minellii Antić et Makarov,
2016. Likely troglobiont, known only from
the Sukhaya and Bolshaya Azishskaya
caves, both near Sochi, Krasnodar Prov-
195
ince; body yellowish white, 22–24 ocelli in
6 rows (Antić, Makarov, 2016).
– Heterocaucaseuma feminaepectorum Antić et Makarov, 2016 (Fig. 6C). Likely troglobiont, from the Krasnoyarskaya and
Pechalnaya caves, both in the Dzykhra karst
massif, and the Osennyaya, Medvezhya and
Gigantov caves in the Alek Massif, Greater
Sochi, Krasnodar Province; body yellowish white, 10–18 ocelli in 3–4 rows (Antić,
Makarov, 2016; Antić et al., 2018).
– Metamastigophorophyllon giljarovi
(Lang, 1959). Troglophile, widespread
along the Black Sea coast from Tuapse to
Abkhazia, recorded also from the Shirokopokosskaya (= Bozhyey Materi, = Our Lady)
Cave, Greater Sochi, Krasnodar Province
(Antić, Makarov, 2016).
Order Julida Brandt, 1833
Family Blanulidae C.L. Koch, 1847
– Nopoiulus (s. str.) ammonites Enghoff,
1984. Likely troglobiont, described from
the Ammonitovaya Cave near Shedok, Psebai District, Krasnodar Province (Enghoff
1984), later recorded from the nearby Dedova Cave (Golovatch, Enghoff, 1990).
– Nopoiulus (s. str.) kochii (Gervais, 1847).
Troglophile, abundant populations observed in the Bolshaya Kazachebrodskaya
and Shirokopokosskaya caves near Sochi,
Krasnodar Province (Golovatch, 1981,
1984/85; Golovatch, Enghoff, 1990).
Family Julidae Leach, 1814
– Archileucogeorgia sp. Troglophile?, recorded from the Fanagoriyskaya Cave near
Goryachiy Klyuch, Krasnodar Province
(Ljovuschkin, 1966).
– Cylindroiulus placidus (Lignau, 1903).
Troglophile, widespread across the western
Caucasus, including a record from the Meshok Cave, Maikopsky District, Adygea
Republic (Lohmander, 1936).
– Cylindroiulus pterophylacum Read, 1992.
Troglophile, widespread across the western
Caucasus, including a record from the
Beloskal’skaya Cave, Khostinsky City District, Greater Sochi, Krasnodar Province
(Read, 1992).
196
S.I. Golovatch et al.
– Cylindroiulus schestoperovi Lohmander,
1932. Troglophile, widespread across the
western Caucasus, including records from
the Beloskal’skaya Cave, Khostinsky City
District, Greater Sochi, Krasnodar Province, and a cave in western Georgia (Read,
1992).
Order Polydesmida Peach, 1815
Family Polydesmidae Peach, 1815
– Brachydesmus furcatus Lohmander, 1936.
Troglophile, widespread across the northwestern Caucasus, including records from
the Fanagoriyskaya Cave near Goryachiy
Klyuch, the Baribana and Akhunskaya caves
(Golovatch et al., 2016), as well as the
Medvezhya Cave (new data) near Sochi.
Family Paradoxosomatidae Daday, 1889
– Strongylosoma kordylamythrum Attems,
1898. Troglophile, widespread across the
Caucasus, also recorded from the Psebe
Cave near Tuapse, as well as the Shirokopokosskaya (= Bozhyey Materi) and Labirintovaya caves, Khostinsky City District,
Greater Sochi, Krasnodar Province, and a
cave in Abkhazia (Evsyukov et al., 2016).
Family Trichopolydesmidae Verhoeff, 1910
– Caucasodesmus inexpectatus Golovatch,
1985. Troglobiont, unpigmented, antennae
and legs elongated, from the Nyvdjinlagat
(= Tagardonskaya) Cave in North Ossetia
(Golovatch, 1984–1985).
Superclass Hexapoda Blainville, 1816
Class Collembola Lubbock, 1870
Order Poduromorpha Börner, 1913, sensu
D’Haese, 2002
Superfamily Hypogastruroidea Salmon,
1964, sensu Deharveng, 2004
Family Hypogastruridae Börner, 1906
– Typhlogastrura preobrazhenskyi Babenko, 1987. Troglobiont, described from the
Shuby-Nykhasskaya Cave in North Ossetia
(Babenko, 1987).
Order Entomobryomorpha Börner, 1916,
sensu Soto-Adames et al., 2008
Superfamily Tomoceroidea Szeptycki, 1979
Family Tomoceridae Schäffer, 1896
– Plutomurus jeleznovodskii Kniss et Thibaud, 1999. Troglobiont with traits of tro-
glomorphism, described from a cave near
Zheleznovodsk, Stavropol’ Province (Kniss,
Thibaud, 1999; Kniss, 2006).
– Plutomurus kelasuricus Martynova, 1969.
Troglophile devoid of distinct troglomorphic features, recorded from the Baribana
Cave, Alek Massif, Greater Sochi, Krasnodar Province (Kniss, Thibaud, 1999; Kniss,
2006).
– Plutomurus sorosii Kniss et Thibaud,
1999. Troglobiont with traits of troglomorphism, described from a cave in the Teberda Nature Reserve, Karachaevo-Cherkessia (Kniss, Thibaud, 1999; Kniss, 2006).
Class Insecta Linnaeus, 1758
Order Plecoptera Burmeister, 1839
Family Leuctridae Klapálek, 1905
– Leuctra furcatella Martynov, 1928. Trogloxene. Larvae and adults recorded from
the grotto part of the Krasnoaleksandrovskaya Cave, Lazarevsky City District,
Greater Sochi, Krasnodar Province (Palatov, Sokolova, 2015).
Family Nemouridae Newman, 1853
– Nemoura elegantula Martynov, 1928.
Trogloxene. Larvae and adults recorded
from the grotto part of the Krasnoaleksandrovskaya Cave, Lazarevsky City District,
Greater Sochi, Krasnodar Province (Palatov, Sokolova, 2015).
Order Orthoptera Latreille, 1793
Family Rhaphidophoridae Walker, 1869
– Dolichopoda euxina Semenov, 1901 (Fig.
6D). Troglophile. Described from caves in
the Caucasus (Semenov, 1901), being very
common there. Known from caves in the
Adygea Republic (Shapovalov et al., 2015),
the Krasnodar Province, Abkhazia, western
Georgia (Satunin, 1912; Birstein, Lopashov, 1940; Zaitsev, 1948; Birstein, 1950;
Barjadze et al., 2015) and South Ossetia
(Bey-Bienko, 1965, 1969), altogether over
40 records.
Order Trichoptera Kirby, 1813
Suborder Annulipalpia Martynov, 1924
Family Polycentropodidae Ulmer, 1903
– Plectrocnemia sp. sensu Lepneva, 1940.
Possible troglophile. In the Krasnodar Prov-
Subterranean biota of European Russia
ince, recorded from the Dolgaya Cave,
Greater Sochi. Larvae live under stones and
in weak currents (Lepneva, 1940; Palatov,
Sokolova, 2015).
Order Coleoptera Linnaeus, 1758
Family Carabidae Latreille, 1802
– Caucasaphaenops molchanovi Belousov, 1999 (Fig. 6E) Troglobiont from the
Sokolova and Gigantov (new data) caves,
Greater Sochi, Krasnodar Province (Belousov, 1999).
– Caucasorites kovali Belousov, 1999.
Troglobiont from the Akhunskaya Cave,
Greater Sochi, Krasnodar Province (Belousov, 1999).
– Caucasorites kovali amplicolis Belousov,
1999. Troglobiont from the Psakho Cave,
Greater Sochi, Krasnodar Province (Belousov, 1999).
– Caucasorites shchurovi Belousov et Zamotajlov, 1997. Troglobiont from the Entomologicheskaya Cave, Lazarevsky City
District, Greater Sochi, Krasnodar Province (Belousov, Zamotajlov, 1997).
– Caucasorites victori Belousov, 1999.
Troglobiont from the Akhunskaya Cave,
Khostinsky City District, Greater Sochi,
Krasnodar Province (Belousov, 1999).
– Cimmerites circassicus Reitter, 1888. A
troglomorphic species not yet encountered
in caves. Described from mountain regions
of the Krasnodar Province, between Tuapse and Sochi. Characterized by reduced
eyes and an unpigmented tegument (Reitter, 1888–1889; Winkler, 1926), likely
adapted to the MSS.
– Cimmerites kryzhanovskii Belousov,
1999. Possible troglobiont, described from
the Malaya Kazachebroskaya Cave, Adlersky City District, Greater Sochi, Krasnodar
Province (Belousov, 1999).
– Cimmerites maximovitchi Belousov and
Koval, 2011. Possible troglobiont, described
from the Akhunskaya and Labirintovaya
caves, Khostinsky City District, Greater
Sochi, Krasnodar Province; occurring together with Caucasorites victori on the
197
western slope of Mt Akhun, Sochi region
(Belousov, Koval, 2011).
– Cimmerites zamotajlovi Belousov, 1998.
Troglophile, described from forest litter on
Mt Verblyudka near the village of Kalinovoe Ozero, Greater Sochi, Krasnodar Province. Recorded also from other epigean habitats near Khosta, as well as from the Vorontsovskaya Cave, all within Greater Sochi, Krasnodar Province (Belousov, 1998).
– Duvalius gusevi Belousov, 1989. A troglomorphic species not yet recorded from
caves. Described from the bank of a mountain spring near the sources of Kezadon
River, Irafskiy District, North Ossetia, where
it occurs under large boulders (Belousov,
1989). Characterized by a light coloration
and strongly reduced eyes, likely adapted to
the MSS.
– Duvalius miroshnikovi Belousov et Zamotajlov, 1995. Troglobiont from the Baribana Cave, Greater Sochi, Krasnodar Province (Belousov, Zamotajlov, 1995).
– Jeannelius birsteini Ljovuschkin, 1965.
Troglobiont from caves in Greater Sochi,
Krasnodar Province (Vorontsovskaya,
Akhunskaya, Beloskal’skaya) (Ljovuschkin, 1965).
– Jeannelius zhicharevi Lutshnik, 1915. A
troglomorphic species not yet recorded from
caves. Described from the Mt Achishkho
karst massif near Sochi (Luchnik, 1915;
Kryzhanovskij et al., 1995); a species related to J. gloriosus and J. magnificus, both
from Abkhazia.
– Meganophthalmus irinae Belousov et
Zamotajlov, 1999. Troglophile, described
from the Chernogorye karst massif near
Maikop, Adygea Republic, occurring both
in epigean habitats under stones and in
caves (Belousov, Zamotajlov, 1999).
– Meganophthalmus kravetzi Komarov,
1993. Troglobiont, described from the Fontanka Cave in the upper reaches of Belaya
Rechka River, 15 km SW of Nalchik, Cherkesskiy District, Kabardino-Balkaria. Found
25 m off the cave entrance, eyes absent
(Komarov, 1993; Belousov, Koval, 2009).
198
S.I. Golovatch et al.
– Nannotrechus ciscaucasiens (Ljovuschkin, 1972) (= Birsteiniotrechus ciscaucasiens Ljovuschkin, 1972). Likely troglobiont, found in the Ammonitovaya Cave
near Psebai, Kuban River basin, Mostovskoy District, Krasnodar Province (Ljovuschkin, 1972a). This genus belongs to the
Neotrechus lineage and is closely related to
Troglocimmerites.
– Nannotrechus fishtensis Belousov, 1989.
A troglomorphic species not yet recorded
from caves. Described from near Mt Fisht,
at the sources of Armyanka River, Mostovskoy District, Krasnodar Province. Characterized by reduced eyes and an unpigmented tegument (Belousov, 1989), likely
adapted to the MSS.
– Nannotrechus kovali Belousov, 1989. A
troglomorphic species not yet encountered
in caves. Described from the montane areas near Solokh-Aul, Greater Sochi, Krasnodar Province (Belousov, 1989). Characterized by reduced eyes, likely adapted
to the MSS.
– Porocimmerites imitator Belousov, 1998.
A troglomorphic species not yet encountered in caves. Characterized by reduced
eyes and a reddish coloration, occurring in
the low flow region of Mzymta River near
Sochi (Belousov, 1998), likely adapted to
the MSS.
– Trechus heniochicus Ljovuschkin, 1970.
Troglophile, described from the Vorontsovskaya Cave, Khostinsky City District,
Greater Sochi, Krasnodar Province. Found
on rotting wood (Ljovuschkin, 1970).
– Laemostenus koenigi (Reitter, 1887). Troglophile, recorded from the Fanagoriyskaya
and Stalaktitovaya caves, both near Goryachiy Klyuch, Krasnodar Province (Vereshchagina, 1985; Vereshchagina, Makarov,
1986).
– Laemostenus tschitscherini Semenov,
1909. Troglophile, described from the Dakhovskaya Cave, Maikopsky District, Adygea Republic (Semenov-Tian-Shanskiy,
1909), also known from many caves near
Maikop (Belaya, Sukhaya, Budkova etc.)
(Vereshchagina, 1985; Vereshchagina, Makarov, 1986).
– Pterostichus lacunosus (Chaudoir, 1844).
Likely troglophile, recorded from the
Vorontsovskaya Cave, Khostinsky City
District, Greater Sochi, Krasnodar Province (Ljovuschkin, 1962).
Family Cryptophagidae Kirby, 1937
– Cryptophagus pilosus Gyllenhal, 1827.
Troglophile, recorded from the Chortova
Nora Cave, Khostinsky City District, Greater Sochi, Krasnodar Province (Koval,
2004a).
Family Curculionidae Latreille, 1802
– Otiorhynchus vargovitchi Davidian,
2007. Troglobiont, described from the Beloskal’skaya Cave, Khostinsky City District,
Greater Sochi, Krasnodar Province (Davidian, 2007).
Family Elmidae Curtis, 1830
– Limnius colchicus Delève, 1963. Likely
troglophile, both larvae and adults encountered in the waters of the Dolgaya Cave near
Khosta, Greater Sochi, Krasnodar Province. Penetrating inside the cave up to 1 km,
but retaining the typical coloration (Palatov, Sokolova, 2015).
Family Leiodidae Fleming, 1821
– Catops subfuscus Kellner, 1846. Troglophile, recorded in a cave at Svetlaya Balka,
Mostovskoy District, Adygea Republic, at
Eki-Suara, Malokarachaevskiy District,
Karachaevo-Cherkessia, and in a cave at
Ispravnaya, Zelenchuk District, Karachaevo-Cherkessia (Perkovsky, 1991).
– Choleva sp. Subtroglophile, recorded from
the Labirintovaya Cave, Khostinsky City
District, Greater Sochi, Krasnodar Province (Ljovuschkin, 1962).
Family Pselaphidae Latreille, 1802
– Bryaxis balneator Besuchet et Kurbatov,
2007. Troglophile, recorded from the
Akhunskaya Cave, Khostinsky City District, Greater Sochi, Krasnodar Province
(Besuchet, Kurbatov, 2007), but also inhabiting epigean biotopes.
– Bryaxis kovali Besuchet et Kurbatov,
2007. Troglophile, recorded from the La-
Subterranean biota of European Russia
birintovaya Cave, Khostinsky City District,
Greater Sochi, Krasnodar Province; showing no visible adaptations to subterranean
life (Besuchet, Kurbatov, 2007).
– Seracamaurops komarovi Hlaváč, Kodada et Koval, 1999 (Fig. 6F). Troglobiont,
described from the Baribana and Gigantov
(new data) caves in the Alek Massif, Greater Sochi, Krasnodar Province (Hlaváč et
al., 1999).
Family Ptiliidae Heer, 1843
– Ptenidium intermedium Wankowicz,
1896. Troglophile associated in its development with bat guano. Recorded from the
Chortova Nora Cave, Khostinsky City District, Greater Sochi, Krasnodar Province
(Koval, 2004a).
Family Scirtidae Fleming, 1821
– Odeles sp. Troglophile, larvae found under stones in a stream in places with a
weakened current in the Dolgaya Cave,
Khostinsky City District, Greater Sochi,
Krasnodar Province (Palatov, Sokolova,
2015).
Family Staphylinidae Lameere, 1900
– Bisnius parcus (Sharp, 1874). Troglophile, recorded from the Chortova Nora
Cave, Khostinsky City District, Greater
Sochi, Krasnodar Province (Koval, 2004a).
– Heinzia caucasica Gusarov et Koval,
2002. Troglophile, described the Baribana
Cave in the Alek Massif, Greater Sochi,
Krasnodar Province, recorded also from
the Sokolova (= Atsinskaya), Dolgaya and
Vorontsovskaya caves in the same area;
showing no visible adaptations to subterranean life, encountered in the Caucasus in
epigean habitats as well (Gusarov, Koval,
2002).
Order Diptera Linnaeus, 1758
Suborder Nematocera Schiner, 1862
Family Limoniidae Speiser, 1909
– Limonia nubeculosa Meigen, 1804. Troglophile, repeatedly recorded from Transcaucasian caves: Bolshaya Kazachebrodskaya, Malaya Kazachebrodskaya, Akhunskaya, Shirokopokosskaya, Vorontsovskaya
etc., all within Greater Sochi, Krasnodar
199
Province, etc. (Birstein, Lopashov, 1940;
Zaitsev, 1948; Birstein, 1950; Koval,
2004b).
Family Sciaridae Billberg, 1820
– Neosciara sp. Troglophile, recorded from
the Bolshaya Kazachebrodskaya Cave,
Adlersky City District, Greater Sochi, Krasnodar Province (Ljovuschkin, 1966).
Suborder Brachycera Zetterstedt, 1842
Family Heleomyzidae Bezzi, 1911
– Heteromyza atricornis Meigen, 1830.
Troglophile, recorded from the Bolshaya
Kazachebrodskaya (= Verkhne-Mzymtinskaya), Akhunskaya and Chortova Nora
caves, all within Greater Sochi, Krasnodar
Province (Birstein, 1950; Koval, 2004a, b).
Larvae dwelling in bat guano (Zaitsev,
1948).
Phylum Chordata Haeckel, 1874
Subphylum Vertebrata J.-B. Lamarck, 1801
Class Mammalia Linnaeus, 1758
Order Chiroptera Blumenbach, 1779
Caves play a vital role in the life-cycle of
eight (out of 31 reported from the Russian
Caucasus) bat species, which use them as hibernation, breeding and swarming sites. Another
16 species occur in caves and artificial undergrounds in some periods of the life cycle, but are
not strictly dependent on them.
Family Rhinolophidae Gray, 1825
– Rhinolophus euryale Blasius, 1853. Seven distributional records of R. euryale were
reported from the vicinity of Sochi, Krasnodar Province, Russia, including one nursery roost in a small cave near the Nizhnyaya
Shilovka village (Gazaryan, 2016).
– Rhinolophus ferrumequinum (Schreber,
1774) (Fig. 7B). This typical cave-dwelling
species is abundant in the western Caucasus, whereas it is rather rare in the central
part of the Caucasus due to the deficit of
suitable underground chambers. Nursery
roosts and hibernating groups have been
found in the Caucasus only in underground
chambers of different origin, mainly in karst
caves. At least 65 among nearly 100 summer and winter records were made in the
latter roost type.
200
S.I. Golovatch et al.
Fig. 7. Some bats from the Russian Great Caucasus in nature. A — a Ringed European barbastelle,
Barbastella barbastellus (Schreber, 1774), hibernating on the wall of a mine near Derbentskaya, Seversky
District, Krasnodar Province; B — a winter aggregation of Greater Horseshoe bats, Rhinolophus
ferrumequinum (Schreber, 1774), in a gypsum cave near Psebai, Mostovskoy District, Krasnodar Province;
C — a colony of Bent-winged bats, Miniopterus schreibersii (Kuhl, 1817), in a limestone cave near Guamka,
Apsheronsky District, Krasnodar Province; D — a maternity roost of Lesser mouse-eared bats, Myotis blythii
(Tomes, 1857), in a cave near Karabudakhkent, Dagestan. All pictures by Suren Gazaryan.
Рис. 7. Некоторые летучие мыши с Большого Кавказа России в природе. A — широкоушка
европейская (Barbastella barbastellus (Schreber, 1774)), зимующая на стене шахты близ станицы
Дербентская (Северский район Краснодарского края); B — зимнее скопление большого подковоноса
(Rhinolophus ferrumequinum (Schreber, 1774)) в гипсовой пещере близ поселка Псебай (Мостовской
район Краснодарского края); C — колония обыкновенного длиннокрыла (Miniopterus schreibersii
(Kuhl, 1817)) в известковой пещере близ поселка Гуамка (Апшеронский район Краснодарского края);
D — материнский насест остроухой ночницы (Myotis blythii (Tomes, 1857)) в пещере близ села
Карабудахкент (Дагестан). Все фотографии Сурена Газаряна.
– Rhinolophus hipposideros (Borkhausen,
1797). The Lesser horseshoe bat is widely
distributed across the region from the Utrish Peninsula in the West to the vicinity of
Derbent in the Southeast. It hibernates in
caves and other underground spaces, whilst
maternity colonies overwhelmingly occupy
above-ground roosts. In total, this bat has
been reported from nearly 120 geographi-
cal locations in the Russian Caucasus,
nearly half of which are caves.
– Rhinolophus mehelyi Matschie, 1901.
The only maternity and hibernation roost in
Russia has been known since 1972 in the
Karabudakhkentskaya Cave in Dagestan,
with the number of bats gradually depleting
from 1000 in 1972 to 30–40 in 2008.
Subterranean biota of European Russia
Family Vespertilionidae Gray, 1821
– Barbastella barbastellus (Schreber, 1774)
(Fig. 7A). Barbastelles utilize caves and
other underground shelters for hibernation
and swarming. Of the ca 60 location records
of the species, as many as 35 belong to
caves. The hibernation colony in the Kanyon
Cave, Krasnodar Province is the largest
ever reported for the species, accumulating
up to 8 thousand individuals in harsher
winters. Another large winter aggregation
of over 2,000 barbastelles was discovered
in the Mayskaya Cave in KarachaevoCherkessia (Gazaryan, 1999, 2007a; Gazaryan, Panyutina, 2013).
– Barbastella caspica (Satunin, 1908).
Amirkhanov (1980) reportedly found 6 individuals in two small caves near the Amushi
and Tagada villages in Dagestan. The species was also recorded from caves in other
parts of its distribution range.
– Eptesicus serotinus (Schreber, 1774). This
species only occasionally hibernates in
caves; several animals were found in the
Kanyon Cave in winter, few serotines were
also netted at the entrance to Babaylovskaya
Cave in the Caucasian State Nature Reserve
during swarming in August. In summer,
resident males often occupy crevices and
niches at the entrance parts of large caves
and grottos.
– Hypsugo savii (Bonaparte, 1837). The
species uses spacious caves and grottos as
swarming sites. It was netted at the entrances to the neighbouring two caves in the
Mzymta River valley near the city of Sochi:
Bolshaya Kazachebrodskaya (Smirnov,
2001) and Dzykhrinskaya (= Lianovaya).
– Myotis alcathoe von Helversen et Heller,
2001. Several individuals were netted entering or flying out from the Fanagoriyskaya
and Krasnoaleksandrovskaya (= Takhira)
caves, Krasnodar Province during swarming, one male was found hibernating in the
Rozhnova Cave in Karachaevo-Cherkessia.
– Myotis bechsteinii (Kuhl, 1817). Single
hibernating individuals were found in the
karst caves Fanagoriyskaya and Kanyon.
201
Swarming Bechstein’s bats were netted at
the entrances to the caves Malaya and
Bolshaya Kazachebrodskaya (Tsytsulina,
1998; Smirnov, 2001), and Dzykhrinskaya
(= Lianovaya), Akhunskaya and Fanagoriyskaya, all within the Krasnodar Province.
– Myotis blythii (Tomes, 1857) (Fig. 7D).
With ca 110 record locations, the Lesser
Mouse-eared Bat occurs literally everywhere in montane ecosystems of the region.
As everywhere within their range, these
bats occupy both underground and aboveground roosts in summer, although hibernate only in caves and other underground
sites. Large maternity colonies comprising
hundreds of females are sustainably present
in the caves Karabudakhkenskaya (Dagestan) and Samorodnaya (KarachaevoCherkessia); a mass winter aggregation
amounting to over 2,000 bats is known from
the Shubi-Nykhass Cave in North Ossetia
(Gazaryan, 2007b).
– Myotis brandtii (Eversmann, 1845).
Brandt’s bat is common in woodland areas
across the region and the absence of any
records from Dagestan is apparently related
to insufficient research efforts. Among 26
record locations in the Russian Caucasus,
seven belong to the following caves: Fanagoriyskaya, Kanyon, Babaylovskaya and Tru52 in the Krasnodar Region, Mayskaya and
Pogrebok in Karachaevo-Cherkessia, and
Anglo-Russkaya in the Adygea Republic.
During the swarming time, M. brandtii was
abundant among bats netted at the entrances to these caves, even though only single
solitarily hibernating Brandt’s bats were
found there in the winter.
– Myotis dasycneme (Boie, 1825). One
young female was found on 8 November
2003 in the Gunkina-4 Cave in Karachaevo-Cherkessia (Gazaryan, 2004), but the
occurrence has since been revealed nowhere else in the region. Consequently, we
tend to associate this single record with a
vagrant bat.
– Myotis daubentonii (Kuhl, 1817). Until
the end of the 20th century, the species had
202
S.I. Golovatch et al.
been considered as extremely rare or even a
recent colonist in the Caucasus and Transcaucasia. However, after extensive fieldwork the former putative status was reassessed to common and abundant across the
region (Gazaryan et al., 2008). Although it
was recorded at 82 locations in the Russian
Caucasus, only several hibernation sites
containing modest (up to 10 animals) numbers of Daubenton’s bats have been found
so far. Daubenton’s bats do not aggregate in
large clusters in the Caucasus and only sole
animals could be observed in the caves
Fanagoriyskaya, Ared, Kanyon and Chernorechenskaya, Krasnodar Region, as well
as in several mines.
– Myotis davidii Peters, 1869. One genetically identified specimen of this species
was collected in the Razvalka mine near
Pyatigorsk, Stavropol Province. See Benda
et al. (2016) for details.
– Myotis emarginatus (Geoffroy, 1806).
Similar to the Greater Horseshoe Bat, the
distribution of this species in the Caucasus
is limited by the availability of suitable
underground maternity sites. The species
has not been found in the Eastern Caucasus
and reported only twice from the central
part of the region, including one record in a
small cave near the Babugent settlement
(Kabardino-Balkaria). Other 21 record locations lay in karstic areas of the Western
Caucasus within the Kranodar Province and
Adygea Republic. Maternity colonies were
reported from the Khadzhokhskaya (now
abandoned by bats), Dedova Yama, Teshebskaya, Svetlaya, Mordvinovskaya and Ared
caves; hibernating individuals were recorded in the Ambitsukova, Fanagoriyskaya,
Kabaniy Proval, Ared and Dedova Yama
caves.
– Myotis mystacinus (Kuhl, 1817). The
distribution of M. mystacinus in the region
and its dependence on caves is unclear due
to difficulties in a morphological identification of this species in its contemporary
sense. Genetically identified specimens were
collected in the Fanagoriyskaya Cave (Benda et al., 2016).
– Myotis nattereri (Kuhl, 1817). Recorded
from 27 sites in the Russian Caucasus, this
bat has rarely been found in caves and undergrounds. Few individuals were captured as
they swarmed in the Akhunskaya, Pervomayskaya, Babailovskaya, Fanagoriyskaya and
Kanyon caves, Krasnodar Province. In the
latter two caves, single Natterer’s bats were
also found during winter. Nine pregnant females were netted on 4 June 2007 as they
swarmed at the entrance to Kamennye Sarai
Cave, Stavropol Province.
– Nyctalus noctula (Schreber, 1774). Roosting behaviour unusual for this species was
first reported in 1980 for the Proval Cave,
Stavropol Province (Gazaryan, Kazakov,
2002), in 2008 this colony remained in the
cave. Only adult males used this cave in
summer, whereas females were found there
only between September and May.
– Pipistrellus pipistrellus (Schreber, 1774).
A group of 30–40 hibernating pipistrelles
was found in a deep crevice inside the
Kamennye Sarai Cave, Stavropol Province
on 20 December 2009. This is the only
documented evidence of this species hibernating in Russian caves.
– Plecotus auritus (Linnaeus, 1758). Twenty-six of ca 60 records of this species in the
region belong to underground sites, including 20 karst caves, where it was found both
during the hibernation and swarming seasons. The largest number of the brown longeared bats netted at one roost did not exceed
15 animals, and only solitary individuals
could usually be found in winter shelters.
– Plecotus macrobullaris Kuzyakin, 1965.
The sole record in an underground site was
made in a mine at Razvalka Mountain,
Stavropol Province on 21 December 2009.
However, we anticipate future findings of
this species in natural underground roosts.
Family Miniopteridae Dobson, 1875
– Miniopterus schreibersii (Kuhl, 1817)
(Fig. 7C). Among the Russian representatives of Chiroptera, Schreiber’s Bent-wing
bat shows the highest rate of dependence on
caves and is very selective in terms of their
Subterranean biota of European Russia
size, accessibility and microclimate. Hence,
it inhabits only the Western Caucasus with
its vast karst areas, and is not distributed
eastward to the Central and Eastern Caucasus; 34 record locations in the Adygea Republic, Krasnodar Province and Karachaevo-Cherkessia are available. All known
records pertain to natural caves, with two
exceptions when four individuals were observed in abandoned mines and one animal
was found roosting under a bridge. The
largest colonies (up to 5,000 bats) occupy
spacious horizontal caves with large entrances, which make them exposed to disturbance by visitors and susceptible to vandalism and intentional killing. For these
reasons, bats forsook some of the previously reported cave roosts and many colonies
have depleted in numbers since the first
extended survey (Gazaryan, Ostapenko,
1999). To avoid possible further decline
and disturbance, we abstain from listing the
names of these roosting sites.
Family Molossidae Gervais, 1856
– Tadarida teniotis (Rafinesque, 1814).
This species does not inhabit caves in the
strict sense of this term, but colonies could
be found in ceiling crevices of rocky overhangs, where these bats may also swarm.
Under such circumstances the free-tailed
bats were found in a large grotto at the
valley of the Cherek River in KabardinoBalkaria (Gazaryan, Tembotova, 2007).
203
tails and mites, as well as a number of troglophiles amongst earthworms, spiders, beetles
and flies (see above). In 1973, the Bolshaya
Azishskaya and Fanagoriyskaya caves, both in
the Krasnodar Province, were allotted the status
of nature monuments. A good number of karst
cavities are situated inside nature reserve stock
objects of higher rank. Thus, numerous caves
are located within the Sochi National Park or
Caucasian Biosphere State Nature Reserve. Yet
many caves like Shulgan-Tash, Bolshaya Azishskaya, Vorontsovskaya, Bolshaya Kazachebrodskaya etc., despite their protection statuses, are
subjected to economic/excursion activities, this
clearly rendering a negative effect on the biotas
they host.
A number of cave animal species in European Russia are listed in regional red data books.
Thus, the Red Data Book of the Moscow Region
contains the amphipod crustacean Synurella
ambulans (F. Müller, 1847). The Red Data
Book of the Krasnodar Province lists several
cave mollusks, crustaceans and many insects,
while that of the Adygea Republic a few insects
as well, all largely highly local endemic troglobionts (e.g. Turbanov et al. (2016a, b, c) and
references therein). A number of bat species are
likewise listed in several regional red data books.
The main emphasis is thereby put on the conservation of caves as geological bodies, but not as
shelters to the biotic components they support.
VI. Conclusion
V. Conservation of caves and hyAs a result, the subterranean biota of European Russia, including that of the northern Caupogean species
Some of the karst cavities in European Russia are protected as objects of nature reserve
stock, with various statuses allotted. Yet no
dedicated animal life conservation is being performed in such caves. For example, the Shulgan-Tash (= Kapovaya) Cave in Bashkortostan
(= Bashkiria), southern Cisuralia, is part of the
Shulgan-Tash Federal State Nature Reserve as
an object of the national historical and cultural
legacy of the palaeolithic epoch. However, this
very cave also hosts a few troglobiont spring-
casus, but excluding that of Crimea, presently
comprises at least 389 species or subspecies
(some still unidentified) from 229 genera (a few
still unidentified), 150 families, 75 orders, 48
classes and 25 phyla, among which 96 species or
subspecies represent presumed stygo– or troglobionts. The taxonomically most diverse phylum is Arthropoda which encompasses the bulk
of the fauna, in particular due to crustaceans,
beetles and collembolans. Whereas the troglofauna of northern Russia is completely devoid
of such strongly cave-adapted animals, in the
204
S.I. Golovatch et al.
Urals, Cis-Urals and central Russia there are a
few, but the richest hypogean list in the region
concerns the northern Caucasus, especially its
western part where karst is particularly abundant.
Acknowledgements
The authors sincerely thank Yevgeny Zakharov (Sochi Branch of the Russian Geographical Society) for his very valuable comments on
our work, including the polytoponymy of the
caves of Greater Sochi, Krasnodar Province.
Special thanks go to an anonymous referee who
reviewed the submission and provided constructive criticism.
Field studies by Ilya Turbanov and Dmitry
Palatov were performed in the framework of the
state assignment of the Federal Agency for
Scientific Organizations (FASO Russia) (topics
Nos АААА-А18-118012690106-7, ААААА18-118012690105), supported in part also by
the Russian Foundation for Basic Research
(project No. 17-54-40017 Абх_a).
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Responsible editor K.G. Mikhailov