ArticlePDF Available

Amphibians and Reptiles in the Subterranean Cavities of the Crimean Mountains

  • T. I. Vyazemsky Karadag Scientific Station - Nature Reserve - Branch of the Institute of Biology of the Southern Seas of the RAS


The current paper summarizes old and recent herptiles records in the karst cavities of the Crimean Mountains, highlighting also the means of entry of poikilothermic vertebrates into the karst. It was found that the underground habitats are on purpose utilized by some amphibians and reptiles, characterized by mostly crepuscular and/or night activity as well as by the pronounced ability to move on the vertical surfaces and, in many cases, by the synanthropization tendency. One of the prerequisites for the animals' entry to the karst might be their habitat's conditions such as arid climate, absence of the forest, harsh winters at the upper distribution limit.
DOI: 10.30906/1026-2296-2019-26-1-29-53
Ilya S. Turbanov,1,2 Oleg V. Kukushkin,3,4* and Robert S. Vargovitsh5
Submitted August 17, 2018
The current paper summarizes old and recent herptiles records in the karst cavities of the Crimean Mountains,
highlighting also the means of entry of poikilothermic vertebrates into the karst. It was found that the underground
habitats are on purpose utilized by some amphibians and reptiles, characterized by mostly crepuscular and/or night
activity as well as by the pronounced ability to move on the vertical surfaces and, in many cases, by the
synanthropization tendency. One of the prerequisites for the animals’ entry to the karst might be their habitat’s
conditions such as arid climate, absence of the forest, harsh winters at the upper distribution limit.
Keywords: Amphibia; Reptilia; karst; subterranean habitats; Crimean Peninsula.
The Crimean Mountains embrace the area of circa
7000 km2organized in three more or less parallel ridges
of different origin and age (Yena et al., 2004). The most
outstanding part of these mountains is the Main (Moun-
tain) Ridge consisting of Upper Jurassic limestone dis-
played aboveground as block low mountains and giant
massifs with plateau-like summits (latter called yailas).
The cuesta ridges of the Premountaun (Piedmont, or
Foothills) are composed from the younger limestones of
Upper Cretaceous, Paleocene and Eocene (Inner Ridge),
or Neogene limestones (Exterior Ridge). Current land-
scapes of the Crimean Mountains are supposed to be
formed in Late Pliocene as well as in Quaternary periods
(Muratov and Nikolaev, 1940; Muratov, 1954), what de-
fines mainly the Pleistocene age of the Crimean karst
(Dublyansky, 1966, 1977). The high intensity of karst
processes in the Crimean Mountains takes place mostly
due to the abundancy and remarkable thickness of the
water-soluble rock deposits (limestones), which are com-
pletely denuded by water-wind erosion at far distances as
well as by the specific climate characterized by heavy
rains in summer period.
The Crimean karst is usually related to the Mediterra-
nean (open) type and is called holokarst, or bare karst
(Maksimovich, 1963). However, it has to be noticed that
in many upland parts and in the mountain hollows a sod-
ded or covered (underground) karst prevails (Amelichev
and Matyushkin, 2011). Today in karst landscapes of the
Crimean Mountains the combination of all surface and
underground karst types including ponors, pits, caves,
grottos, etc. can be observed. Recent Cadaster of karst
caverns of the Crimea has catalogued over 1500 caves
with a total length of about 120 km: 590 caves on the
Ai-Petri Yaila, 422 on the Karabi Yaila, 155 on the
Chatyrdag Yaila massifs, and 101 in the Piedmont area
(Amelichev et al., 2014). Therefore, the Mountain Cri-
mea is a significant karst area, even in the scale of Alpine
Mediterranean folded belt. Currently, karstologists in-
clude the Crimean Mountains into the Crimean-Cauca-
sian karst country as a special province (Vakhrushev,
Relatively old age (mostly Middle or Late Pleisto-
cene), the presence of multiple underground cavities and
a high diversity of the conditions determine altogether
1026-2296/2019/2601-0029 © 2019 Folium Publishing Company
Russian Journal of Herpetology Vol. 26, No. 1, 2019, pp. 29 – 53
1I. D. Papanin Institute for Biology of Inland Waters of Russian Acad-
emy of Sciences, Borok, Yaroslavl oblast’, Russia;
2Cherepovets State University, Cherepovets, Vologda oblast’, Russia.
3T. I. Vyazemski Karadag Scientific Station — Nature Reserve, Cri-
mea; e-mail:
4Zoological Institute of Russian Academy of Sciences, St. Petersburg,
5I. I. Schmalhausen Institute of Zoology of Academy of Science of
Ukraine, Ukraine; e-mail:
* Corresponding author.
the uniqueness of the Crimean cave fauna characterized
by a high level of endemism primarily among the crusta-
ceans (Isopoda), false scorpions (Pseudoscorpiones),
myriapods (Diplopoda), springtails (Collembola), insects
(Coleoptera) and other groups of animals. The Crimean
Peninsula, according to the latest underground fauna sur-
vey of the former USSR (Turbanov et al., 2016a, 2016b,
2016c), is a karst-enriched glacial refugium of the
nemoral biota with 44 troglo-stigobiont invertebrate spe-
cies. At the same time, true troglobionts among poikilo-
thermic vertebrates are not registered in the Crimean
cave fauna, while their several species are known from
the caves of Western Eurasia mountain group (Parin,
1983; Sket, 2017). Such deficiency can be explained by
many reasons, for instance, the long-term insularity of
the Crimea, the flooding of the Crimean karst during ma-
rine transgressions, and the immersion under the sea
level of a significant part of the mountain “core” (Mura-
tov, 1960). Moreover, an unfavorable conditions for the
cave fauna development and survival within the cold and
dry Pleistocene periods, and also, probably, in the mod-
ern epoch — primarily due to the drastic seasonal fluct-
uations of water level in caves and scarce trophic
resources insufficient to the existence of vertebtates’pop-
ulations might input to this deficiency as well (Dublyan-
sky, 1977; Vakhrushev and Amelichev, 2001).
Nevertheless, the Crimean Peninsula has not been
fully explored yet in faunistic relation. As for the subter-
ranean cavities, their systemic zoological studies have
been started only since the last decade. The majority of
the known cave systems have been poorly investigated,
and many karst cavities, undoubtedly, remain undiscov-
ered. For instance, Mamut-Chokrak Cave in Bayidar-
skaya valley (territory of Sevastopol City), the second
largest cave on the Peninsula, was found by the first
author of this paper in 2010. At the moment, the total
length of the traversed passes of this cave comprises
more than 5450 m. Another example illustrating that the
Crimean Karst is still terra incognita is the discovery of a
very large (with a length over 1 km) Tavrida Cave in the
Eocene Nummulite limestones of the Inner Ridge in
Until recently, the Crimean caves have been known
mainly as a shelter for some troglophilous bat species
(Chiroptera). Meanwhile, not only mammals, but also
other vertebrates, including amphibians and reptiles,
have been regularly registered in the Crimean karst, but,
unfortunately, without the relevant description in the sci-
entific literature.
Surprisingly, the first record of the amphibians in the
Crimean Karst refers to an European olm (or “human
fish”), Proteus anguinus Laurenti, 1768 — a member of
tailed amphibians, which, in fact, has not been found in
the Crimea yet. For example, V. Kondaraki reported that
the olm (“Proteus anquineus [sic!]”) dwells “ the un-
derground basins and generally near by the sources of
rivers, where there are gloomy grottos filled with water”
(Kondaraki, 1875, p. 37). Furthermore, A. Brauner
(1904) did not exclude the presence of the European olm
in the Crimea, since in his field guide to reptiles and am-
phibians of the Crimea even the morphological descrip-
tion of this amphibian can be found. At the same work A.
Brauner discusses the members of herpetofauna absent at
the Crimean Peninsula, though occuring in the adjascent
areas: “Finally, probably in caves with large reservoirs,
the olm (Proteus anguinus), which is still not found in the
Crimea, dwells” (Brauner, 1904, p. 57). Later many re-
searchers emphasized on the question of olm’s presence
in the Crimea (Birstein, 1963; Sharygin, 1998; Kotenko,
2010; Pysanets and Kukushkin, 2016). However, the
most detailed survey of this problem is given in mono-
graph “Herpetologica Taurica” (Szczerbak, 1966) with
the references to two findings of olm-like animals resid-
ing in the Crimean caves in the first half of the 20th cen-
tury. It has to be mentioned that M. M. Szczerbak has
been also searching the olm personally in cavities of
Chatyrdag (Suuk-Koba, Bin-Bash-Koba, and Dlinnaya)
and Kizil-Koba caves at the Dolgorukovskaya Yaila on
June 1 – 2, 1958, which were discovered only recently
and, therefore, not visited by tourists (see Voinstvensky,
Unfortunately, it has to be admitted that there is no
any reason for hope to find troglobiont amphibians like
the Proteus in the Crimean karst, since the representa-
tives of the extinct genus Mioproteus Estes et Darevsky,
1977, known from Early Pleistocene of the Eastern Eu-
rope including the territories adjacent to the Crimea were
reophilic forms, not associated with the underground and
generally mountain habitats (Averinov, 2001; Ratnikov,
Whereas the first mention about the record of am-
phibians in the Crimean caves belonged to the legendary
“olm,” the second concerned a species that was consid-
ered as extinct in the Crimea at that times. Only weakly
calcinated subfossil remnants of a common spade-footed
toad, Pelobates fuscus (Laurenti, 1768) (currently re-
ferred to as a Pallas’ spadefoot, Pelobates vespertinus
(Pallas, 1771)) was found in a remote hall of one of the
caves on large upland called Karabi Yaila (Burchak-
Abramovich, 1936). It is reported that “... in October
1929 I found in one of the karst caves at Karabi Yaila
(1 km the north from the foot of the Kara-Tau Mt.), along
the other osteological material, three sacral bones of Pe-
lobates fuscus (Laur.)” and also “... it is more correct to
attribute them to the subfossils category, very close to our
time. It is very likely that spadefoot toad is still living
30 Ilya S. Turbanov et al.
near this cave, in limestone crevices of the yaila”
(Burchak-Abramovich, 1936, p. 131). According to this
author, amphibians might entered the caves hiding from
the daylight, or be brought there by owls as a prey (most
likely — little owl, Athene noctua (Scopoli, 1769), which
is known for its preference of amphibians and reptiles in
a diet).
In addition, the records of rare findings even of an
ordinary members of the Crimean batrachofauna refer
mainly to the second half of the 20th – the beginning of
the 21st centuries. Thus, V. Taraschuk in his monograph
mentioned the record of a marsh frog Pelophylax ridi-
bundus (Pallas, 1771) in the subterranean habitats:
“Marsh frogs have been occasionally registered in the un-
derground water reservoirs — in the caves of the Crimea”
(Taraschuk, 1959, p. 93). The same information was
given by M. Szczerbak (1966) without the description of
the specific localites. Only few records of an Eastern tree
frog Hyla orientalis (Bedriaga, 1890) in the Karabi Yaila
underground cavities were published recently (Pysanets
and Kukushkin, 2016). It is noteworthy that an adult
specimen of this frog was found in “Myshinaya Schel’”
grotto (“Bat’s Fissure”) at the Coastal Crest of the Kara-
dag mountain group on June 25, 1980 (Szczerbak, 1984).
It has to be emphasized that the amphibian travelled to
the distance of 20 m from the entrance to the cave. There-
fore, the question, how has the frog got so deep into an
abrasion grotto on a bare cliff at the sea level remains
Moreover, a green toad, Bufotes viridis (Laurenti,
1768), was discovered near one of the entrances to the
Kizil-Koba Cave on February 26, 2001 (Zagorodniuk,
2004). The author suggested that the amphibian felt into
the trap in search of a suitable place for wintering.
Finally, an amateur, though valuable paper dedicated to
the Karelin’s newt, Triturus karelinii (Strauch, 1870)
findings in caves of the south-western Crimea (Matyush-
kin, 2010) has to be mentioned here.
The information about the reptiles records in the Cri-
mean caves is scarce as well. In observations in summer-
time 1939 some lizards were reported as a blood sourse
for the sandflies Phlebotomus sequens (= Sergentomyia
minuta Rondani, 1843), which reside in caves near Seva-
stopol and at the Southern Coast of the Crimea, however,
without their exact locations: “... specific habitats of
these sandflies are caves... Females of this species in
the Crimea apparently feed mainly on the blood of rep-
tiles — lizards, which are many in those caves, where
sandflies live,” and “... sandflies were caught mainly in
shallow caves, mostly close to the entrance, staying in
numerous cracks and clefts, where lizards usually hide”
(Perfiliev, 1941, p. 275). It is known that the sandflies
usually attack reptiles with a twilight-night rhythm of ac-
tivity, and geckos in particular (Alekseev, 2008). S. minu-
ta, along with S. dentata (Sinton, 1933) feed on Ko-
tschy’s gecko, Mediodactylus kotschyi (Steindachner,
1870) in the Eastern Mediterranean region (Naucke,
2002). Therefore, it was suggested that, regarding the
work of Perfiliev, we can refer only to the Crimean (Da-
nilewski’s) gecko, M. danilewskii (Strauch, 1887), but
not to any other lizard species (Turbanov et al., 2015).
It can be anticipated that the geckos in the southern Cri-
mea caves might be found before the Second World War,
however, this fact was not adequately highlighted in the
scientific literature and/or analyzed by herpetologists.
And, finally, in the publication on the biogeographi-
cal status of the Crimean populations of M. danilewskii
and the leopard snake, Zamenis situla (Linnaeus, 1758),
some data about the use of relic karst cavities by both of
these species in Balaklava and Aya Cape surroundings
were presented (Kukushkin et al., 2017).
Therefore, the current review is the first attempt to
summarize all data regarding the findings of poikilother-
mic vertebrates in underground cavities of the Crimean
Mountains. The main objectives of our study were to ana-
lyse the data on the amphibians and reptiles records in
this area, to evaluate the possibility and scale of animal
entry into the caves, as well as to identify the regularities
of underground habitats use by various representatives of
the Crimean herpetofauna.
All materials were collected at the territory of the
Crimean Peninsula within 2003 – 2018. Hardly access-
ible caves and mountain trials on the Baydarskaya and
Ai-Petrinskaya Yailas were explored mainly by the first
author using special speleological and climbing equip-
ment. The caves near the top of the Aya Cape (Balaklava
District, Sevastopol) were repeatedly monitored by the
second author in 2015 – 2018, technically supported by
P. Oksinenko (Simferopol). The Karadag grottos were
also surveyed during the whole period of observations by
the second author as well.
Laser distometer, an inclinometer and a mountain
compass were used for the topographical studies in
caves. The geographical coordinates and the altitudes of
the terrain above the sea level were determined using the
GPS navigator Garmin 64st. The air temperature in the
cavities was measured by the Exo-Terra digital thermo-
meter with the accuracy of 0.1°C.
The zonation of the Crimean karst is given according
to the B. Vakhrushev’s scheme (Vakhrushev, 2009).
A significant and exclusively important part of our
data on reliable amphibians and reptiles findings in the
Cave Herpetofauna of the Crimea 31
32 Ilya S. Turbanov et al.
TABLE 1. Records of Amphibians in Caves of Mountainous Crimea
Karst area,
affiliation of the
point of record
of relief
The name
of the cavity
(its cadastral
m a.s.l.
Date of
Depth of
the record
from the
surface, m
Distance from
entrance, m
(for the caves with
horizontal stretch)
of individuals
Source of
T. karelinii AY, Yalta btw. Mt. Byuzyuka
and Mt. Balchik-
(= Uparennaya)
(4425/3353 – 3)
940 2nd half
of May 1999 5 2 A. Yushko, O. Peretyatko
BB, Sevastopol Canyon of Chernaya
River, Mt. Khlama Grot Anny Cave
354 31.05.2003 At entrance ca. 10 (ad., juv.) Matyushkin (2010)
AY, Sevastopol locality
Karadagsky Les Kristalnaya (= Maksimovich)
834 2003 – 2004 Up to 64 Several adults
and larvas Kotenko,
Kukushkin (2010)
AY, Sevastopol Ai-Petrinskaya
Yaila Gibel’ Yaponii
1051 15.12.1990 20 5 Matyushkin (2010)
05.07.2004 1
AY, Sevastopol Ai-Petrinskaya Yaila Aie-Kale (4428/3356-10) 1096 11.08.2004 10 1 Matyushkin (2010)
AY, Sevastopol Talakanskaya Hollow Unnamed karst ponor 810 13.10.2007 under stone
at the
bottom of
a karstic
2 juv. Matyushkin (2010)
AY, Sevastopol Ai-Petrinskaya Yaila Raskop Medvezhyi
871 18.01.2008 8 1 Matyushkin (2010)
BB, Sevastopol Bayidarskaya Hollow Mamut-Chokrak
280 08.10.2016 3 – 4 1 S. Kebets, A. Akimov
AY, Sevastopol Mt. Kurt-Kaya Entuziastov
296 20.11.2016 27 3 Ye. Yanovskyi (personal
AY, Sevastopol Ai-Petrinskaya Yaila Gelektitovaya-Vesennyaya
1075 02.04.2017 40 2 A. Akimov, S. Kebets
BB, Sevastopol Canyon of Chernaya
River Chernorechenskaya
77 05.05.2017 30 – 40 1 I. Turbanov
AY, Sevastopol Bolshoyi Babulgan
Hollow Kuban’ (4428/3356-4) 956 15.07.2017 26 1 I. Turbanov, S. Arefiev,
O. Kukushkin
H. orientalis KY, Belogorsk Karabi Yaila Gvozdetsky (4452/3433-23) 985 Late October
1990 180 1 A. Papyi (personal
2011 1 Pysanets, Kukushkin
KY, Belogorsk Karabi Yaila Monastyr-Chokrak
(= Su-Teshigy)
998 10.08.1993,
06.08.2003 90 Several specimens R. Vargovitsh
Early July
2007 110 adult and several
lumps of eggs
in the pool
of condensate
M. Grishkov (personal
BB, Sevastopol Bayidarskaya Hollow Mamut-Chokrak
280 19.08.2010 30 1 I. Turbanov
Cave Herpetofauna of the Crimea 33
Karst area,
affiliation of the
point of record
of relief
The name
of the cavity
(its cadastral
m a.s.l.
Date of
Depth of
the record
from the
surface, m
Distance from
entrance, m
(for the caves with
horizontal stretch)
of individuals
Source of
AY, Yalta Ai-Petrinskaya Yaila Oreshek (4427/3359-3) 1070 23.10.2010 24 1 A. Papyi (personal
BB, Sevastopol Bayidarskaya Hollow,
Mt. Biyuk-Sinor Sakhtykh (4425/3348-1) 639 11.08.2012 7 1 S. Bogdanenko (personal
B. viridis SY, Simferopol Dolgorukovskaya
Yaila Kizil-Koba (= Krasnaya)
585 26.02.2001 4 – 6 1 Zagorodniuk (2004)
AH, Sudak Mt. Shpil’,
Agarmysh massif Pogreb (4500/3500-1) 420 27.11.2005 7 – 8 1 V. Nuzhdenko (personal
BB, Sevastopol Ravine
Kobalar-Dere Karan’-Koba (= Temnaya)
339 18.11.2006 16 1 Matyushkin (2010)
KY, Belogorsk Karabi Yaila,
Ridge Bagalma Kara-Murza
625 22.08.2009 130 1 P. Oksinenko (personal
BB, Sevastopol Ridge Spilia,
Mt. Asketi Asketi-2 (not
included in the Cadastre) ca. 320 Mid-Aug.
2014 5 – 7 1 P. Oksinenko (personal
26.09.2015 5 – 7 1 O. Kukushkin,
P. Oksinenko
BY, Alushta Babugan Yaila,
area of the Mt.
Unnamed cave (not
included in the Cadastre) 1460 Nov. 2014 3 – 4 1 Yu. Balakhtinova,
S. Klushin (personal
AY, Sevastopol locality
Karadagsky Les Bizyukskaya (not
included in the Cadastre) 756 01.04.2017 106 1 R. Potapov (personal
KY, Belogorsk Karabi Yaila,
Egiz-Tinakh Valley Yabushkan-Koba
967 11.08.2017 10 1 O. Kukushkin, I.
KD, Theodosia Ridge
Khoba-Tepe Grotto under the Rock Mayak
(not included in the Cadastre) ca. 200 19.08.2017 6 – 7 1 O. Kukushkin, P.
KY, Alushta Karabi Yaila Tserkov’ (4451/3436-1) ca. 985 02.12.2017 10 1 Yu. Balakhtinova, S.
P. (ridi-
AY, Yalta Ai-Petrinskaya Yaila,
Mt. Morcheka Druzhba
969 01.01.2007 45 1 juv. A. Shendrik (personal
AY, Sevastopol Mt. Karadag Skelskaya
(4427/3352 – 3)
304 09.04.2009 20 1 I. Turbanov
11.04.2010 2 – 3 1
06.03.2015 2 – 3 1
Sept. 2017 15 50 up to 10 juv. K. Moskvina (personal
04.03.2018 15 50 3 juv. I. Turbanov. A.
AY, Sevastopol Mt. Villya-Burun Lyagushach’ya
905 23.02.2010 23 1 S. Kebets (personal
KY, Belogorsk NE spures of Karabi
Yaila, area of the
Mt. Gol’-Kaya
390 30.08.2015 8 1 A. Akimov, Yu.
Balakhtinova (personal
34 Ilya S. Turbanov et al.
TABLE 2. Records of Reptiles in Caves of Mountainous Crimea
Karst area,
affiliation of the
point of record
Mountain massif,
element of relief
The name
of the cavity
(its cadastral
m a.s.l.
Date of
Depth of the
from the
surface, m
Distance from
entrance, m
(for the caves
with horizontal
of indivi-
of information
M. danilewskii KD, Theodosia maritime slope of the
Ridge Karagach Unnamed grotto (not
included in the Cadastre) ca. 20 July 2001 2 1 M. Afanasiev (personal
BB, Sevastopol Cape Aiya,
Mt. Kalafatlar Gekkonovaya
496 08.09.2012 5 – 12 3 Yu. Balakhtinova
28.12.2012 3 – 20 ca. 18 I. Turbanov, Yu.
20.03.2016 3 – 5 1 O. Kukushkin,
P. Oksinenko
09.10.2016 2 – 3 1 O. Kukushkin,
A. Nadolny
BB, Yalta Bayidarskaya Yaila,
Mt. Chelebi Gravitational peel break
on the climbing route
“Kamin v kamine”
610 16.09.2012 1 1 I. Turbanov
BB, Sevastopol Cape Aiya,
Mt. Gurush Arochnaya
(4428/3338-2) and
403 05.12 and
15.12.2012 — from7–10
to 15 – 25 ca. 10 and
up to 20 I. Turbanov, Yu.
385 26.09.2015 3 1 O. Kukushkin, P.
BB, Sevastopol Ridge Spilia,
Mt. Asketi Asketi-1 and Asketi-2
(not included in the
ca. 320 Mid-August
2014 5 – 12 ca. 16 P. Oksinenko, Ye.
Oksinenko (personal
Asketi-1 26.09.2015 5 – 7 1 O. Kukushkin, P.
BB, Sevastopol Karanskie Rocks,
Kaya-Bash Heights Unnamed cave (not
included in the Cadastre) ca. 270 10.06.2016 At entrance 1 I. Turbanov
L. agilis AY, Sevastopol Ai-Petrinskaya Yaila Tri Slepykh
860 14.05.1994 10 10 1 female
and its egg Matyushkin (2010)
C. austriaca AY, Yalta maritime cliff
of the Mt. Morcheka Medovaya
910 19.06.2006 30 1 juv. A. Shendrik (personal
CY, Simferopol Chatyrdag,
Lower plateau Vyalov-Azimutnaya
(4445/3417 – 6)
1020 07.08.2006 30 1 (dead
specimen) E. Bednarskaya
KY, Belogorsk Karabi Yaila Skazka (4452/3433-7) 1002 Early July 2009 45 1 I. Turbanov
KY, Belogorsk Karabi Yaila Monastyr-Chokrak
(= Su-Teshigy)
998 18.07.2010 95 1 À. Akimov (personal
KY, Belogorsk Karabi Yaila Kastere
1011 24 – 27.07.2010 70 1 I. Turbanov
Cave Herpetofauna of the Crimea 35
Karst area,
affiliation of the
point of record
Mountain massif,
element of relief
The name
of the cavity
(its cadastral
m a.s.l.
Date of
Depth of the
from the
surface, m
Distance from
entrance, m
(for the caves
with horizontal
of indivi-
of information
KY, Belogorsk KarabiYaila Kruber (4451/3430 – 8) 998 24 – 27.07.2010 12 1 I. Turbanov
D. caspius KD, Theodosia Ridge
Khoba-Tepe Kostrovy Grotto (not
included in the Cadastre) ca. 230 30.05.2008 4 – 5 1 O. Kukushkin, E.
Gladilina, A. Nadolny
DY, Simferopol Demerdzhi Yaila Maloy Akademii Nauk
(= MAN) (4446/3422-2)
1170 Summer 2008 1 – 2 1 E. Bednarskaya
BB, Sevastopol Cape Aiya,
Mt. Kalafatlar Gekkonovaya 496 20.03.2016 10 1 O. Kukushkin, P.
BB, Sevastopol Bayidarskaya Hollow Urkusta-Chokrak-Koba
385 06.08.2017 30 1 (dead
specimen) I. Turbanov,
S. Arefjev
Z. situla BB, Sevastopol Bayidarskaya Hollow,
Mt. Biyuk-Sinor Sakhtykh (4425/3348-1) 639 11.08.2012 7 1 S. Bogdanenko
BB, Sevastopol Cape Aiya,
Mt. Gurush Unnamed grotto (not
included in the Cadastre) ca. 400 02.12.2012 2 1 Yu. Balakhtinova
BB, Sevastopol Cape Aiya,
Mt. Kalafatlar Gekkonovaya 496 05.12.2012 10 1 juv. I. Turbanov
BB, Sevastopol Ridge Spilia,
Mt. Asketi Asketi-2 300 – 330 26.09.2015 At entrance 1 (moulted
skin) O. Kukushkin, P.
BB, Sevastopol Cape Aiya,
Mt. Gurush Dvukhkupolnaya 385 26.09.2015 At entrance 1 (moulted
skin) O. Kukushkin, P.
BB, Sevastopol Cape Aiya, Mt.
Kalafatlar Gnomov (not included
in the Cadastre) ca. 490 31.03.2016 3 1 O. Kukushkin, E.
BS, Bakhchisarayi Mt. Baba-Dag Mangup-Kale-14 Cave
ca. 550 03.05.2018 10 14 1 I. Turbanov
E. sauromates BB, Sevastopol Bayidarskaya Hollow Mamut-Chokrak
280 13.05.2013 At entrance 1 L. Bogdanenko
V. renardi CY, Simferopol the head of
Orlinoe gorge Monashjya (not included
in the Cadastre) 1025 17.05.2009 At entrance 1 E. Bednarskaya
Ophidia indet. CY, Simferopol Chatyrdag,
Lower plateau Viking (not included in
the Cadastre) ca. 950 05.04.2006 ca. 50 1 E. Bednarskaya
CY, Simferopol Chatyrdag,
Lower plateau Golubinaya-2
ca. 990 28.07.2007 15 1 E. Bednarskaya
CY, Simferopol Chatyrdag,
Lower plateau Tryokhetazhnaya
1009 12.11.2007 10 1 E. Bednarskaya
underground cavities confirmed with the photos was
given by our correspondents: speleologists, mountain-
eers, biologists, and local historians. Findings of 1990s –
2010s were organized in chronological order (Tables 1
and 2).
The free accessible “Caves. Information retrieval
system” Internet resource ( was
used for the specification of the cavities cadastral number
(for each cavity — at the first mention in the tables). The
current taxonomy of the amphibians and reptiles has to
be reffered according to D. Frost and P. Uetz et al. data-
bases (both were accessed on August 17, 2018).
In the tables the following abbreviations were used
for the Crimean Mountains karst areas: BB — Bayidar-
sko-Balaklavsky; AY — Ai-Petrinsky; BY — Babugan-
sky; CY — Chatyrdagsky; SY — Dolgorukovsky; DY —
Demerdzhinsky; KY — Karabyisky; AH — Agarmysh-
sky; KD — Sudaksky (karst areas of the Mountainous
Crimea); BS — Bakhchisaraysky (karst area of the Cri-
mean Piedmont).
In our paper, a four category cave animals ecological
classification proposed by B. Sket was used: (i) troglo-
bionts species strictly associated with hypogean habitats;
(ii) eutroglophilic, originally epigean species able to exist
permanently in subterranean habitats; (iii) subtroglophi-
lic species able to live in subterranean habitats tempo-
rarily or permanently, though occasionally going out to
the surface to support their biological functions such as
feeding or reproduction; (iv) trogloxenic species occa-
sionally got into the caves (Sket, 2008). However, the dif-
ferences between subtroglophiles and trogloxenes seem
to be quite vague at the moment (Trajano and de Car-
valho, 2017), and the representatives of the same species
under the diverse circumstances can be classified into the
third or the fourth category. Moreover, the periodicity of
visiting underground cavities and the reason for their us-
age by the Crimean amphibians and reptiles should be
studied more carefully. Obviously, in such a cases, the
simple categorization of animals found in caves can be
challenging. Therefore, for the characterisation of the
amphibians and reptiles, which get into the underground
cavities purposefully in some phases of their life cycle
and then able to travel back to their typical habitats, the
definition of a facultative cavernicole is used. In turn, an
accidental trogloxenes get into the caves unintentionally
and after a while, without the opportunity to go out to the
surface, die there.
The data on the amphibians and reptiles found at the
Crimean underground cavities within the 1980s – 2010s
are summarized, respectively, in Tables 1 and 2.
Four amphibian species from four families were re-
corded in the cave habitats (Fig. 1A).
Order Caudata
Karelin’s newt — Triturus karelinii (Strauch, 1870)
This Caudata species has the maximum number of
individuals (more than 30 in total) as well as localities
(12 in total) found. Most of the records belong to Ai-Pet-
rinsky karst area (the western part of the Ai-Petrinskaya
Yaila and its spurs). Single individuals or groups, some-
times including up to 10 newts, were also seen in the
Bayidarsko-Balaklavsky karst area within the known
range of the species (more often in the Chernaya River
basin). The altitudes range of the findings varies between
250 and 1100 m a.s.l., which is only slightly lower as
compared to the limiting altitude of the species records in
the Crimea (1200 m a.s.l.) (Szczerbak, 1966). The newts
were registered at depths up to 64 m with the maximum
distance from the horizontal caves entrance of 30 – 40 m.
Amphibians occured both in comparatively dry cave sec-
tions (Fig. 2a) and puddles of the condensed water
(Fig. 2b). It is well-known fact that the newts are able to
climb on vertical surfaces, and they were often found on
the rock ledges of several meters in hight from the cavity
floor. According to the seasonal distribution of records,
newts visit caves all the year round. Individuals observed
in winter were quite active and were in a good fit (Ma-
tyushkin, 2010). Besides the newts, in the same caves the
numerous invertebrates such as troglophilic spiders, cen-
tipedes, and insects (Trichoptera, Lepidoptera, Diptera)
as well as vertebrates — horseshoe bats (Rhinolophus
sp.) were observed. The adult male of T. karelinii found
in July 2017 at the bottom of the Kuban’ Cave in the Bol-
shoy Babulgan hollow (26 m in depth, air temperature at
midday 8.2°C) was inert and almost irresponsive to vari-
ous stimuli. However, after being transfered above-
ground, it became active within the several minutes and
took an indicative alert position typical for the newts
from T. karelinii group, exhibited in the display of ventral
aposematic coloration. Moreover, T. karelinii large larvae
were recorded in the Skelskaya Cave (Table 1), though
this information needs to be verified.
Order Anura
Family Bufonidae
Green toad — Bufotes viridis (Laurenti, 1768)
Toads in the underground habitats occurs in the Cri-
mean Mountain from Balaklava in the west to Karadag
and Agarmysh massives in the east (Table 1). Generally,
more than 10 individuals were recorded in 10 localities in
total. The altitude range of the findings is much higher
36 Ilya S. Turbanov et al.
Cave Herpetofauna of the Crimea 37
Fig. 1. Records of poikilothermic vertebrates in the karst cavities of the Mountainous Crimea: A, records of amphibians: red circle,Triturus
karelinii: 1, Mamut-Chokrak Cave; 2, Chernorechenskaya Cave; 3, Grot Anny Cave; 4, Razocharovanie Cave; 5, Entuziastov Cave; 6, Kristalnaya
Cave; 7, unnamed karst ponor in Talakanskaya Hollow; 8, Aie-Kale and Raskop Medvezhyi caves; 9, Gibel’Yaponii Cave; 10, Gelektitovaya-Ve-
sennyaya Cave; 11, Kuban’ Cave; green triangle,Pelobates vespertinus: unknown cave, 1 km to the north from the Kara-Tau Mt., Karabi Yaila;
blue square,Bufotes viridis: 1, Asketi-2 Cave; 2, Karan’-Koba Cave; 3, Bizyukskaya Cave; 4, unnamed cave on the Babugan Yaila; 5, Kizil-Koba
Cave; 6, Yabushkan-Koba Cave; 7, Tserkov’ Cave; 8, Pogreb Cave; 9, grotto under the Rock Mayak, Khoba-Tepe Ridge; yellow rhombus,Hyla
orientalis: 1, Mamut-Chokrak Cave; 2, Sakhtykh Cave; 3, Oreshek Cave; 4, Monastyr-Chokrak Cave; 5, Gvozdetsky Cave; 6, seaside grotto on the
Khoba-Tepe Ridge; violet hexahedron,Pelophylax (ridibundus) complex: 1, Skelskaya Cave; 2, Lyagushach’ya Cave; 3, Druzhba Cave; 4, Kara-
su-Bashy Cave; B, records of reptiles: blue circle,Mediodactylus danilewskii: 1, unnamed cave on the Karanskie Rocks; 2, Asketi-1 and Asketi-2
caves; 3, Arochnaya and Dvukhkupolnaya caves; 4, Gekkonovaya Cave; 5, gravitational peel break on the climbing route “Kamin v kamine”; 6, un-
named grotto on the Ridge Karagach; orange star,Lacerta agilis: Tri Slepykh Cave; red square,Coronella austriaca: 1, Medovaya Cave;
2, Druzhba Cave; 3, Vyalov-Azimutnaya Cave; 4, Tryokhetazhnaya Cave; 5, Monastyr-Chokrak, Kastere and Skazka caves; 6, Kruber Cave; green
rhombus,Dolichophis caspius: 1, Gekkonovaya Cave; 2, Urkusta-Chokrak-Koba Cave; 3, Maloyi Akademii Nauk Cave; 4, Kostrovyi Grotto on
the Khoba-Tepe Ridge; yellow hexahedron,Zamenis situla: 1, Asketi-2 Cave; 2, Dvukhkupolnaya Cave and unnamed grotto on the Gurush Mt.;
3, Gekkonovaya and Gnomov caves; 4, Sakhtykh Cave; 5, Mangup-Kale-14 Cave; violete triangle, Ophidia indet. (unidentified snakes): 1, Golubi-
naya-2 Cave; 2, Tryokhetazhnaya Cave; 3, Viking Cave.
than those of other amphibian species, varying from 200
to circa 1500 m a.s.l., while the maximum depth com-
prises up to 130 m, and the distance from the entrance in
the horizontal caves exceeds 16 m. The entrances to the
caves, where the toads were observed, were sometimes
located on hardly accessible cliffs (for example, as the
Asketi Cave near Balaklava) (Pysanets and Kukushkin,
2016). An adult toad was caught on a narrow rock cor-
nice at 560 m a.s.l. during the climbing of “Ukho” (“The
Ear”) alpinistic trail on the Mshatka-Kayasy Mountain
(the western part of the Ai-Petrinskaya Yaila) on August
24, 2012. The rock wall height below the cornice was
almost 250 m. Toads are recorded in caves all the year
round. In the hot season (July – September), their find-
ings in the cavities are usually confined to the humid soil
areas in moisture infiltration zone after an abundant
precipitation or to the sources of the condensed water
(Fig. 2c). Numerous records of B. viridis in the same cav-
ities within the long time intervals indicate that toads sys-
tematically use the caves as summer shelters (Table 1).
It has to be also noticed that the air temperature in a
site of a large B. viridis female finding in the Asketi-2
Cave was 17.8°C, while at the entrance –26.5°C on 10:43
a.m. at the end of September 2015.
The altitude of B. viridis records at the Crimean
upland is of a great interest. Thus, a toad has been ob-
served in nameless cave at an altitude of 1460 m. a.s.l.
(44.620917° N 34.291278° E) near the Babugan Yaila
southeastern spur, what is supposed to be the highest
finding of B. viridis in the Crimea. Moreover, this is the
first record of a green toad at Babugan Yaila in general,
despite it was reported before that this toad may dwell on
its northwestern spur in the cordon Alabach area of the
Crimean Nature Reserve at 1200 m. a.s.l. (S. Kostin, per-
sonal communication). In 2000s the targeted expeditions
were undertaken by T. Kotenko aimed to reveal toads,
since it was anticipated that a common toad, Bufo bufo
(Linnaeus, 1758) could be found in the cool and humid
wooded areas adjacent to Yaila. According to some
sources, this species resided the Crimea in the 19th cen-
38 Ilya S. Turbanov et al.
a b
Fig. 2. Amphibians in the Crimean caves: a,Triturus karelinii in Entuziastov Cave, 20 November 2016, photo by Ye. Yanovsky; b,Triturus kareli-
nii in Gelektitovaya-Vesennyaya Cave, April 2, 2017, photo by A. Akimov; c,Bufotes viridis in Asketi-2 Cave, Mid-August 2014, photo by P. Ok-
sinenko; d,Hyla orientalis in Mamut-Chokrak Cave, 19 August 2010, photo by I. Turbanov.
tury (Kotenko, 2010). However, these attempts were pre-
dictably unsuccessful. At the same time, at Chatyrdag
massif B. viridis was found up to 1030 m a.s.l. (its
spawning ocurrs in the deep gorges below 700 m), at Ka-
rabi Yaila — up to 990 m (toad reproduction was ob-
served on the northern side of the plateau at 680 – 830 m
a.s.l.), and at Ai-Petrinskaya Yaila — up to 1160 m a.s.l.
A large number of larvae close to the final stage of meta-
morphosis were found at this elevation in the mountain
meadow steppe to the west from Mt. Bedene-Kyr in the
muddy road puddles on July 15th, 2017. Approximately
at the same altitudes (1100 – 1150 m a.s.l.) the spawning
reservoirs of B. viridis on the Chuchel’ Pass in the Cri-
mean Nature Reserve were recorded (Szczerbak, 1966).
Family Hylidae
Eastern tree frog —
Hyla orientalis Bedriaga, 1890
Circa 10 individuals (mostly adults, less often juve-
nils) were observed in 5 caves, 3 of which are located in
the southwestern Crimea, including the Ai-Petrinskaya
Yaila, and2—inthecentral part of the Karabi Yaila pla-
teaus. The majority of the tree frogs were observed in the
Karabi Yaila caves. The maximum depth of findings in
karst is up to 180 m, while the horizontal distance from
the cave entrance is about 30 m. The altitude range of the
findings varies from 250 to 1100 m a.s.l. The maximum
altitudes of amphibians records outside cavities are
1000 m a.s.l. at Dolgorukovskaya and Karabi Yailas, up
to 1150 m at the Chuchelsky Pass, up to 1200 m at Ai-
Petrinskaya Yaila, and 1230 m a.s.l. at Demerdzhi Yaila
(Pysanets and Kukushkin, 2016). All H. orientalis find-
ings in caves occurred in the second half of the year —
from July to December. Only once the tree frog spawning
was observed in mid-summer (July 2007) in a complete
darkness in the bottom of one of the Karabi Yaila caves
(Table 1). The pattern (presence and/or absence of the
spots) and coloration of the amphibians found in the
caves varied in many shades of green from salad-light to
olive-green (Fig. 2d). Several clutches of whitish eggs
were on the wet rock surface in condensate streams and
in small puddles on the cavity bottom.
Family Ranidae
Marsh frog – Pelophylax (ridibundus) complex
In 4 localities about 10 individuals in total were re-
corded. A marsh frog is more closely related to the
aquatic habitats as compared to other amphibian species
of the Crimean Peninsula, what might explain its attrac-
tion to the cave springs. Only one large adult was found
on the northern part of the Karabi upland, while most of
the findings are confined to the western part of the
Ai-Petri karst area — primarily to the Skelskaya Cave
(Table 1). The maximum depth of the findings is up to
45 m, while the horizontal distance from the cave en-
trance is up to 50 m. In the Skelskaya Cave, groups of
5 10 juveniles were observed in the puddles of circa
0.25 m in depth (for example, in September 2017). How-
ever, P. ridibundus has not been observed in the cave
lakes of the Crimea, although in other karst areas (for in-
stance, in the Balkans) the large clusters of the Ranidae
family members in the underground reservoirs were ob-
served (Koller, 2017). The altitude range of P. ridibundus
records in the Crimean caves comprised 300 – 1000 m
a.s.l., with the maximum elevation of the species in the
Crimea of 1150 – 1200 m a.s.l. on the Mt. Chuchel’ and
Ai-Petrinskaya Yaila (Pysanets and Kukushkin, 2016).
At least 8 species of reptiles from 4 families (3 spe-
cies of lizards and not less than 5 species of snakes) have
been discovered inside or at a high proximity to the en-
trance of underground cavities (Fig. 1b).
Order Sauria
Family Gekkonidae
Crimean (Danilewski’s) gecko —
Mediodactylus danilewskii (Strauch, 1887)
During the daytime in July 2001, a student M. Afa-
nasyev observed an adult gecko moving in the shade on
the shallow grotto vault located on a very steep talus of
the Karagach ridge in the Karadag Nature Reserve
(Fig. 3a). A few years later, in 2005 2006, a large
M. danilewskii population was found at the western part
of the coastal Karadag ridge (Kukushkin, 2009). It has to
be emphasized that it is the northernmost Danilewski’s
gecko population in the world living in natural land-
scapes, which was revealed for the first time by the sec-
ond author of this paper.
In the evening twilight on September 16, 2012, an
adult gecko was photographed on the southern cliff of
Mt. Chelebi (657 m a.s.l.) on “Kamin v kamine” (“Fire-
place inside the fireplace”) alpinistic trail above the Fo-
ros settlement (Fig. 3b). Since the lizard was found at
610 m a.s.l., it is now recognized as one of the highest lo-
cations of Danilewski’s gecko at the Crimea. Moreover,
the highest point of M. danilewskii habitat in the Crimea
is Mt. Ilyas-Kaya (above the Sarych Cape), which peak
rises 682 m a.s.l., where the gecko was tracked in deep
cracks along the brink of a great southern cliffs (Kukush-
kin, 2004, 2009). It is anticipated that the Crimean gecko
can be found in other gravigenic cracks on the Bayidar-
skaya Yaila southern cliffs and the Bayidaro-Kastropol
Wall of Ai-Petrinskaya Yaila as well, though unlikely —
on the yiala cliffs to the east from the Shaytan-Merdven
Cave Herpetofauna of the Crimea 39
During 2012 – 2017 geckos have been repeatedly ob-
served in caves of Baidarsko-Balaklavsky karst area:
near the northern border of the “Cape Aya” State Land-
scape Reserve, and both to the west and east from Bala-
klava Bay (Table 2). This species was recorded in 6 relic
erosion-karst systems in the cracks of Upper Jurassic
conglomerates and limestones. The cavities are opened
by narrow entrances (in some cases circa 0.5–1mindi
ameter) on the gigantic cliffs of southern and western
slope exposures (Fig. 3c,d). The total length of these al-
most horizontal systems exceeds 58 – 64 m (e.g., Gekko-
novaya (Gecko’s) and Arochnaya (Arch) Caves), while
the thickness of the cave’s vault varies from 0.5–2mon
Mt. Kalafatlar to up to 80 m on Mt. Gurush (Fig. 4a). The
total length of the Mt. Asketi cave system is about 50 m;
the height of the entrances to the caves approximates
7 – 15 m, whereas the total height of the cliffs reaches in
some cases 35 – 40 m (Fig. 4b).
Geckos were concentrated mostly at depths of 3
25 m from the entrance in the dim light of meanders and
halls as well as in the dark distant passages. The width of
cavities in the sites of lizards records varied from 0.3
0.5 to 3 m, while the height was 0.6–2m.Intotal, about
70 M. danilewskii individuals were observed in caves —
close to the Cape Aya summit (2012) and in Balaklava
town vicinity (2014) (Table 2). Groups of the lizards
comprising up to 15 – 20 individuals were found both in
winter (Aya Cape caves, December 2012) and in summer
(Mt. Asketi, mid-August 2014). It is remarkable that
within one cavity the lizards were distributed more or
less evenly, but sometimes 2 – 3 individuals were found
close to each other. In winter time geckos remained quite
active and moved along the cave walls, vaults and clay
40 Ilya S. Turbanov et al.
Fig. 3. Landscapes where Mediodactylus danilewskii was observed in the cavities: a, grotto at the foot of the Ridge Karagach in Karadag
paleovolcanic group, photo by O. Kukushkin; b, southern cliffs of the Upland Bayidarskaya Yaila, Mt. Chelebi, photo by O. Kukushkin; c, maritime
cliffs of the Cape Aiya, view from the sea, photo by O. Kukushkin; d, upper edge of western cliffs of the Mt. Kalafatlar, photo by O. Kukushkin.
bottoms (Fig. 5a–c). During the monitoring of the same
caves in 2015 – 2018 no large groups of geckos were re-
corded, only single individuals.
As for the caves microclimate, it can be generally
characterized as ventillated by a warm scheme with
the heat accumulation in cavities during the cold season.
The temperature varies in these cavities, since in the sec-
ond half of December 2012 the temperature outside was
–2 to –4°C, while inside the caves it was 10 – 14°C at a
distance of 10 – 20 m from the entrance, where most of
the lizards were concentrated). In turn, on September
26th, 2015 at 11.00 a.m. the temperature outside was
27°C, though at the entrance hall of the Asketi-1 Cave it
was only 18°C. Furthermore, the same day at 3.00 p.m.,
at the entrance of the Arch Cave it was 25.8°C, but at the
site of gecko tracking at the beginning of the narrow hol-
low upright path starting immediately after the entrance
to the hall it comprised 23.6°C. At the same time, in
March 31st, 2016 at 1.00 p.m. at the entrance to the
Gekkonovaya Cave the temperature was 13.5°C, while at
10 – 12 m from the entrance –7.6°C.
Geckos were not the only inhabitants of these caves,
since they were accompanied by the numerous inverte-
brates such as spiders (Meta sp.), centipedes (Pachyiulus
varius,Scutigera coleoptrata; Lithobiidae), Coleoptera
(Tenebrionidae), Lepidoptera (Noctuidae), and Diptera.
Besides that, under the cave vaults small groups or sole
horseshoe bats [Rhinolophus hipposideros (Borkhausen,
1797) and Rh. ferrumequinum (Schreber, 1774)] were re-
peatedly observed.
Therefore, the height range of gecko’s findings in
karst cavities varied from almost 300 to 500 m a.s.l.
Within the last two decades the predominant point of
view that the Crimean gecko is an exclusively synanthro-
pic species confined to the lower sea-side zone (Szczer-
bak, 1966) has been subjected to critical correction.
Throughout the coast of the Crimean Mountains from
Mt. Kaya-Bash and the southern border of the Balaklava
Valley in the west to the Karadag mountain group in the
east, numerous natural M. danilewskii populations were
revealed. The upper limit of this species distribution in
the Crimea “shifted” from previously established 150
200 m to 600 – 700 m a.s.l. depending on the orographic
and climatic characteristics of the region (Kukushkin,
2004, 2009; author’s unpublished data).
Family Lacertidae
Lindholm’s rock lizard —
Darevskia lindholmi (Szczerbak, 1962)
There were a plethora of findings of this species at
the illuminated parts of karst caves and grottos uniformly
throughout the territory of the Crimean Mountains and
Cave Herpetofauna of the Crimea 41
Fig. 4. Sketchs of some underground systems where large aggregations of Mediodactylus danilewskii were observed: a, Gekkonovaya Cave,
Mt. Kalafatlar, Cape Aiya, made by I. Turbanov and A. Kurichenko; b, Asketi cave system, Ridge Spilia, environs of Balaklava, made by P. Oksi-
nenko and A. Kurichenko.
Piedmont at a small distance (up to 2 m) from the en-
trance, but in rare cases — up to 7 – 10 m (for example,
in artificial cavities of Mangup-Kale “cave town”). Gen-
erally, single individuals getting into the grotto were re-
corded during the daytime with high temperatures. Occa-
sionally, small groups of Lindholm’s rock lizards were
observed at the entrance to shallow, but fairly wide (0.5 –
1 m) vertical crevices in the stony mountain-meadow
steppe (for instance, at the top of Mt. Takja-Tepe on the
Karabi Yaila, 1250 m a.s.l., or in the limestone pavement
landscapes of the western part of Ai-Petrinskaya Yaila).
Unlike the crepuscular geckos, petrophilic lacertids are
active during the day hours, use the cavity entrances only
as a temporary shelter and for the thermoregulation dur-
ing the warm time of the day. Therefore, they were not
registered in caves during the wintering. The data on this
lizard species is not included in Table 2.
Taurida’s sand lizard —
Lacerta agilis tauridica Suchow, 1926
On some stony plots of yaila characterized by poor
vegetation, sand lizards live in karst funnels with the
abundant grasses. Large olive-green specimen (appar-
ently female) was observed on May 14, 1994 in one of
the caves on Ai-Petrinskaya Yaila (Table 2). It seems to
42 Ilya S. Turbanov et al.
Fig. 5. Reptiles in the Crimean caves: a,b,Mediodactylus danilewskii in the relic karst caves of Cape Aiya environs, December 2012, photos
by Yu. Balakhtinova; c,Mediodactylus danilewskii in Asketi cave system, Balaklava surroundings, Mid-August 2014, photo by P. Oksinenko;
d,Zamenis situla, Sakhtykh Cave, Mt. Biyuk-Sinor, 11 August 2012, photo by S. Bogdanenko.
be interesting that a laid egg close to the lizard was found
(Matyushkin, 2010).
Order Serpentes
Family Colubridae
Smooth snake —
Coronella austriaca Laurenti, 1768
To our knowledge, at least 8 findings of the smooth
snake were registered in karst fitting the known range of
this species at the Crimea, while one half of our records
attributed to the central part of Karabi Yaila. Majority of
specimens were recorded in summer, what indicates the
occasional entry of snakes into the cavities within the
peak of the seasonal activity. The maximum depth of the
findings is up to 95 m, while the horizontal distance from
the entrance into the caves is circa 30 m. It is noteworthy
that C. austriaca is the most common snake species on
the Crimean yailas up to the Shaitan-Merdven Pass in the
west and can be seen not only near the plateau edge, but
also in its central parts. On Karabi plateau this snake is
quite ordinary: up to 3 records during the day in June
2011 at the altitude range between 900 1050 m a.s.l.
On Ai-Petrinskaya Yaila and the Karatau Upland (Karabi
Yaila) the smooth snake is observed up to 1200 m a.s.l.,
while at Chatyrdag and Dolgorukovskaya Yaila — up to
1000 m (Szczerbak, 1966; own observations).
Caspian whipe snake —
Dolichophis caspius (Gmelin, 1789)
There are data on 4 findings of this species in cavities
throughout the Crimean Mountains with the altitude
range from about 200 m to almost 1200 m a.s.l. The high-
est locality on the western slope of Demerdzhi Yaila is of
unique importancy, since usually D. caspius in the Cri-
Cave Herpetofauna of the Crimea 43
Fig. 6. Habitats of the Crimean anurans in terrains where they penetrate into the caves: a,Bufotes viridis habitats on the Coastal Crest of Karadag
mountain group, Ridge Khoba-Tepe, photo by O. Kukushkin; b,Hyla orientalis and B. viridis habitats on the Karabi Yaila Plateau, photo by
A. Nadolny; c,Triturus karelinii habitats in Juniperus sparse forest at NW spures of Ai-Petrinskaya Yaila, photo by O. Kukushkin; d, in canyon of
the River Chernaya, photo by I. Turbanov.
mea does not occur in the mountains higher than 1000 m
a.s.l. However, due to the climatic characteristics of this
area, extraordinary high findings of thermophilic reptiles
are described from here: for example, the Balkan wall liz-
ard, Podarcis tauricus (Pallas, 1814) was registered on
Mt. South Demerdzhi at up to 1200 m a.s.l.
Caspian whipe snake uses caves for wintering (one
case) and visits them while hunting for food (2 cases).
A small adult snake was found on hibernation in a rock
crack in the Gekkonovaya Cave at the end of March 2016
(Table 2). At midday of May 30, 2008 the snake of about
1 m long was observed on a narrow ledge under one of
the Karadag’s grotto vaults at the altitudes of more than
5 m. The snake has been hunting for two Rh. hipposide-
ros individuals under the wedge-shaped cave arch. The
disturbed snake disappeared in a deep crack on the sur-
face with a negative inclination angle. Next time, a large
whipe snake has been wading through the bush branches
to the swift (Apus sp.) nest on the rock at the cave en-
trance. In the eastern part of the Inner Ridge of the Sa-
bah-Kogey Valley an active whipe snake was recorded
inside a deep (over 1.5 m) niche under the rock canopy at
a wall foot hunting for lacertids on October 7, 2017 (data
not shown). Recently, the decayed adult snake, most
likely to be the prey of a fox (Vulpes vulpes krymea-mon-
tana Brauner, 1914) was found in one of the caves on
the northern side of the Bayidarskaya Valley (Table 2;
Fig. 7a).
Blotched rat snake —
Elaphe sauromates (Pallas, 1814)
The only record of this species is known from the en-
trance to the Mamut-Chokrak Cave in the Bayidarskaya
Valley in mid-May 2013.
Leopard snake — Zamenis situla (Linnaeus, 1758)
Fresh moulted skin found in underground cavities as
well as findings of snakes themselves allowed to register
7 records within the known range of the species at the
Crimea. The distance from the entrance to the cave was
up to 14 m, the altitudes varied from 300 m to almost
650 m a.s.l., and the highest finding of Z. situla in the
subterranean cavities was detected in the northern
macroslope inside the ponor of the Sakhtyh Cave on the
northern flank of Mt. Biyuk-Sinor on August 2012
(Fig. 5d). The fact of Z. situla occurence at relatively
high altitudes under the harsh mountain climate is worth
of the special attention. For example, in summertime of
1998 and 1999 the leopard snakes were found on
Ai-Petrinskaya Yaila in the retaining wall of the old road
near the Bayidarsky Pass and in the deep cracks on the
top of Mt. Ilyas-Kaya at circa 630 m a.s.l. In the
mid-June 2011 Z. situla was observed on the north-west-
ern slope of Mt. Kilse-Burun at about 700 m a.s.l. More-
over, a young Z. situla female was found on
Ai-Petrinskaya Yaila in a stony steppe near the top of Mt.
Merdven-Kaya (688 m a.s.l.) on May 23, 2012.
Leopard snakes have been often recorded in the karst
area close to the top of Cape Aya (Table 2), what is con-
sistent with the previously published data on the
abundancy of Z. situla in this region (Kukushkin and
Tsvelykh, 2004). In one case, a hibernating young snake
was found in a shallow rock crack at a distance of 10 m
from the entrance, while in other two cases (in early De-
cember 2012 and at the end of March 2016), active
snakes were observed close to the entrance to the cave.
On September 26, 2015, quite fresh and still wet moulted
skins of Z. situla were found on the vertical rock surfaces
at a height of 1.2 – 1.5 m from the ground at the entrance
44 Ilya S. Turbanov et al.
Fig. 7. Remains of snakes in the Crimean caves: a, decomposed corpse of Dolichophis caspius in Urkusta-Chokrak-Koba Cave (Diptera puparia
are clearly visible on the right side of the photo), photo by I. Turbanov; b, skeleton of a colubrid snake (most likely D. caspius) in recently discov-
ered Tavrida Cave (border of Simferopol and Belogorsk District, surroundings of the settlement Zuya), photo by D. Tikhomirov.
of 2 of 4 caves explored at a site between Mt. Asketi to
Mt. Gurush.
The data of locals from the mid-1990s on aggrega-
tions of the hibernating leopard snakes in artificial cavi-
ties of the “cave towns” Eski-Kermen and Mangup in the
western part of the Inner Ridge still has to be checked and
verified. It is noteworthy, however, that these localities in
the South-Western Piedmont are included into the species
range. A young snake was caught on May 3, 2018 in the
Mangup-Kale-14 Cave with the entrance turned to
south-east, what favors the sufficient warming in the
morning hours. The air temperature at the location of the
snake’s finding on the far wall of the cavity was 28.8°C at
11.30 a.m.
Leopard snake found in the Sakhtyh Cave was full,
with a clearly visible thickening on its belly. Probably,
the snake might hunt small bats in the same manner as
D. caspius (see above), since this cave is a shelter for a
small Rh. hipposideros colony and these bats are in the
diet of European colubrids belonging to several genera:
Zamenis longissimus (Laurenti, 1768), Hemorrhois hip-
pocrepis (Linnaeus, 1758), and E. sauromates (Kauch
and Bala, 2005; Garrido-García et al., 2013; Ostrov-
skikh et al., 2015).
Family Viperidae
Puzanov’s (Crimean) steppe viper —
Vipera (Pelias)renardi puzanovi Kukushkin, 2009
The steppe viper was observed only once in stones
near the Monashya (Monk) Cave entrance on the Lower
Chatyrdag plateau at 1025 m a.s.l. in mid-May 2009.
This location almost reaches the upper limit of this spe-
cies range in the Crimea.
Ophidia indet.
All three unknown snake records (due to the limited
observation time and lack of photo proofs) were from
the Lower Chatyrdag plateau at 950 – 1000 m a.s.l.
(E. Bednarskaya, personal communication). With high
probability it might be the most common Chatyrdag’s
snake species, namely C. austriaca and/or V. renardi.
However, according to the responsdent’s descriptions,
it can be assumed that at least in one case the snake be-
longed to the genus Natrix Laurenti, 1768 and might be
identified as a common grass snake, N. natrix (Linnaeus,
1758), or the dice snake, N. tessellata (Laurenti, 1768).
It has to be noticed in this regard that N. tessellata
occuring in a very small number in the pre-yaila forest-
steppe of the northern slope of Chatyrdag massif, has
been registered at altitudes not higher than 750 m a.s.l.
This mainly ichthyophagous snake climbs Chatyrdag
slopes from the Ayan water reservoir, but keeps the dis-
tance from it not more that 2 km. On other Crimean up-
lands N. tessellata has not been recorded yet. Unlike
N. tessellata,N. natrix was observed on Yaila on October
6, 2003 in one of the streams of Besh-Tekne Hollow in
western part of Ai-Petrinskaya Yaila at about 1000 m
a.s.l.6At the beginning of the 20th century, the grass
snake was also registered on the Lower Plateau of Cha-
tyrdag (Puzanov, 1931).
From five known Crimean amphibian species only
P. vespertinus was not detected in the subterranean cavi-
ties, since till the end of the 20th century this species has
been considered to be extremely rare or even extinct at
the Crimean Peninsula. According to the numerous find-
ings of this very hidden burrowing species at the eastern
part of the Crimean Mountains in 2003 2016, the
chances to record it at Karabi Plateau are relatively high
(Pysanets and Kukushkin, 2016). In karst areas character-
ized with drought climatic conditions and heavy or stony
soils this anurans find a shelter in caves and can reach up
to 100 m from the entrance to the subterranean cavities
(Ostrovskikh, 2012). Almost complete absense of water
reservoirs at Karabi Yaila is not necessarily a limiting
factor for P. vespertinus, because our long-time observa-
tions in the arid zones of the South-Eastern Crimea sug-
gest that the reproduction of this amphibian can occur in
small stagnant reservoirs (in fact, puddles) with muddy
water, which do not exist annually. However, P. vesperti-
nus in the Crimea has not been found at altitudes above
250 m a.s.l. yet, and the question of its occurence on the
upland remains open.
The analysis and the detailed study of the amphibian
records in the Crimean caves revealed that they are not
accidental trogloxenes, but cave visitors or facultative
cavernicoles. Undoubtedly, amphibians in some cases
might enter the underground cavities accidentally, first of
all, just fall down into the vertical shafts during their sur-
face activity. Such tunnels and holes in karst caves are
often ending up in a dense grassy or shrubby vegetation
and trap the terrestrial animals, which have only two
ways — to die or to escape. Another way of amphibians’
entry into the underground cavities is through the sys-
tems of caverns and cracks in the epikarst zone.
Some amphibians due to their ecological preferences
are pre-adapted to the life in the karst cavities or at least
their limited use. A green toad such as a very eurytopic
Cave Herpetofauna of the Crimea 45
6In the annotated list of amphibians and reptiles of the Crimean re-
serves (Kotenko and Kukushkin, 2010), the maximum elevation of
N. natrix finding at 850 m a.s.l. in Yalta Mountain Forest Reserve is
incorrect, since, in fact, it is much higher — circa 1030 m a.s.l.
species expresses itself as a typical petrophil at the Cri-
mean Mountains and during the out-of-breeding season
prefers rock clumps, grottos, or ruins as a safe shelter
with favorable microclimate (Fig. 6a). Moreover, toads
are excellent climbers and have been recorded even on
rock ledges while passing the climbing trails. According
to our observations, B. viridis purposefully uses caves
and grottos both during the summer drought and hiberna-
tion (Table 1). Throughout the Crimean Mountains, win-
tering toads have been repeatedly observed in the cavities
of walls and in cracks of rocky cliffs opened on vertical
surfaces at a height of 1.5–2mfrom the ground, some-
times sharing their shelters with lizards — M. danilewskii
and D. lindholmi.
Probably, it is not accidental to find single individu-
als or small groups of H. orientalis in arid karst areas in
the caves primarily on Karabi Yaila plateau (Fig. 6b).
Basing on the frequency of occurrence in some caves, the
frogs can use underground habitats for wintering and/or
the prolongation of their active period intentionally, espe-
cially on the upland with severe climate. It was estab-
lished that a growling grass frog, Litoria raniformis Ke-
ferstein, 1867 (Hylidae) demonstrates trogloxenic behav-
ior differentially: it appears in the caves in one area in
some cases, though is totally absent in others (Fordham,
1985), what can take place in the Crimea as well in a sim-
ilar way. Inspite of the finding of H. orientalis eggs in
completely dark cave, its successful reproduction even in
small cavities is still doubtful; however, it can not be
completely excluded, since the reproductive biology of
this species is very flexible. Thus, in areas with a hot arid
climate (such as Karadag in the South-Eastern Crimea)
during an extremely hot summers, when all natural reser-
voirs dry up for a long time, the spawning and develop-
ment of the Eastern tree frog larvae still occurs in small
artificial reservoirs such as darkened concrete cisterns
and shallow muddy wells. Nevertheless the success of
the reproduction in such cases is extremely low, a limited
number of juveniles successfully finish metamorphosis
(Pysanets and Kukushkin, 2016). The use of the reser-
voirs of this type for the reproduction can be understood
as a step towards the underground reproduction, for ex-
ample, in the entrance part of the caves.
One more amphibian, T. karelinii, also visits various
artificial reservoirs with vertical walls (wells, collectors,
etc.) periodically. Definitely, in karst areas with the lack
of surface waters a stable caves microclimate attracts am-
phibians not only for the time of the breeding season. For
instance, newts in karst were mostly found on dry south-
western slopes of the western part of Ai-Petrinskaya
Yaila covered with sparce juniper forests (Fig. 6c) (Ma-
tyushkin, 2010). However, this tendency is not so strict,
and the newts during the hot season can live in suitable
biotopes in thick leaf litter of hornbeam or beech forests
near the entrance to the caves on the Yaila plateau.
Today the biology of T. karelinii in the Crimea re-
mains poorly studied. It is known that this amphibian is
adapted to quite an arid conditions, characterized by a
comparatively short water phase of life cycle (sometimes
less than 2 months) with a reproduction period extended
from late-February to late-August (Pysanets and Kukush-
kin, 2016). The existence of spawning reservoirs of T. ka -
relinii in the Crimea is unstable: from time to time they
dry up and remains critically shallow or empty for a long
time, sometimes for several years (Kukushkin and Ku-
schan, 2015). In especially arid and hot areas of the
southern macroslope within the territory of Sevastopol
(Laspi amphitheatre), the aggregations of adult newts
were observed near small (several meters in diameter)
shallow puddles covered with branch and leaf litter on
the bottom under the forest canopy near the springs (May
7, 2018). Such situation is supposed to be typical for
newts, because the fish in these reservoirs is completely
absent or does not have opportunity to form large popula-
tions. Some of the newts might migrate to the shallow
karst to survive the dry seasons.
There are some prerequisites for the reproduction of
T. karelinii in underground habitats. For example, in
1999 – 2004 in the Ulu-Uzen River Valley near Alushta a
small “ruinous” newts population has been reproducing
in darkness of the flooded basement of the abandoned
bulding, where females attached eggs to the floating de-
bris and even to the concrete bottom of the reservoir
(Pysanets and Kukushkin, 2016). Of course, this case of
newt’s reproduction in the flooded ruins does not confirm
the possibility of its reproduction in the underground cav-
ities, though the specifics of the conditions in the caves
(primarily, the temperature) is not an insurmountable
limitation for it, especially since the conditions in the Cri-
mean karst are quite variable. For example, it became
evident recently that a fire salamander, Salamandra sala-
mandra (Linnaeus, 1758) uses the caves systematically
not only for wintering, but also for the reproduction7
(Manenti et al., 2009, 2011; Ianc et al., 2012). S. sala-
mandra life cycle is toughly connected with the caves in
46 Ilya S. Turbanov et al.
7Within the territory of the former USSR S. salamandra has been fre-
quently observed in underground cavities (caves, grottos, wells, tun-
nels, etc.) of the Ukrainian Carpathians (Pokynchereda, 1989; R. Var-
govitsh, unpublished data). These amphibians form the winter aggre-
gations of up to 20 – 35 individuals (for example, in a semi-watered
20-m long abandonen adit with a narrow entrance near Mukachevo
Town) retaining only basic activity. Probably, they can also feed on
invertebrates, hibernating in caves. In a number of localities (for in-
stance, near Hlyboke Village, Uzhgorod district) in the adits with the
permanent water flow salamanders larvae at different developmental
stages have been regularly observed, what migh indirectly prove the
reproduction of salamanders in a complete darkness.
karst-enriched areas, which allows to categorize this spe-
cies as a subtroglophile or facultative troglophile (Balo-
gová et al., 2015, 2017).
The reports about the findings of the mysterious
“white newts” in the underground waters of the south-
western Crimea (Sharygin, 1998) is undoubtedly related
to T. karelinii larvae with light coloration of integuments
even under normal insolation, which are able to bleach
(lose the pigmentation quickly) under light-deficiency
conditions. However, the means of their entry into the
karst are still unclear. The larvae might enter the under-
ground habitats by water streams during floods or
through the system of cracks in the epikarst zone, which
may, apparently, become easier by the regular use of the
karst wells by newts for the reproduction (Kotenko and
Kukushkin, 2010; Matyshkin, 2010). It has to be noticed
that the development of larvae is delayed in cold-water
oligotrophic reservoirs. T. karelinii hibernated larvae
have been repeatedly found in wells, spring reservoirs,
and drinking water collectors (Pysanets and Kukushkin,
2016) with the majority of findings at the western spurs
of Ai-Petrinskaya Yaila and the Bayidarskaya Hollow.
At the same time, the reproduction of single newts in
shallow karst in case of drying up of water reservoirs
suitable for spawning remains possible. The small size of
cave reservoirs seems not to be an insurmountable obsta-
cle. For instance, in underground habitats an onset of fire
salamander larvae in small puddles at a distance of more
than 20 m from the entrance to the cave was observed,
what can be considered anomalous, as far as the animals
living on the surface spawn in brooks (Balogová et al.,
The eggs (portions of 2 – 10 eggs in a clutch) were
laid in wet moss in the terrarium by the Karelin’s newt fe-
males within 1 – 15 days after being captured. One group
of females was obtained under a beech forest canopy at a
distance of 10 – 50 m from the shallowed lake in the area
of the Angarsky Pass in the second half of April 2016.
In turn, the second group was collected at the bottom of
the dried lake with the poor vegetation in the forest-
steppe near the Mt. Mangup. Some of the eggs developed
normally in the water film on the moss. At 16 – 20°C the
embryonic development before the appearance of larvae
comprised two weeks.
Assuming that T. karelinii lay fertilized eggs outside
the water reservoirs may also occur in nature, the embry-
onic development takes place in wet litter near the water
sources, under the stones deeply anchored in the ground
or in cavities of the epikarst zone. In case of the abundant
precipitation, the water level in reservoirs suitable for
spawning rises rapidly, flooding their banks, the hatching
of larvae and further development continue under normal
conditions — in the water.
In Mediterranean some cases of atypical localization
of salamandrid species clutches were described. For in-
stance, females of Sardinian brook salamander, Euproc-
tus platycephalus (Gravenhorst, 1829) sometimes lay
eggs near or above the water level under the moist condi-
tions, although normally eggs of this species are laid into
the streams (Sotgiu et al., 2017).
It is necessary to distinguish several records of T. ka -
relinii in the peak of breeding season in caves at a small
distance from the affluent Chernaya River which is un-
avaliable for the newts’ life because of the strong current
and predatory fishes abundance (Fig. 6d). Presumably,
T. karelinii might use the karst cavities (or, in any case,
karst and erosional relief forms) for their reproduction.
Breeding of newts might occur as well in whirlpool de-
pressions (evorsion hollows) in ephemeral stream beds or
at the entrance to the caves with the underground
streams, what has to be studied in details.
It can be generally concluded that T. karelinii has
closer ecological bonds with the underground habitats
than other amphibians living in the Crimea, and that dur-
ing different phases of its life cycle it uses underground
cavities in karst landscapes systematically. At the same
time, in spite of a certain similarity of manifestations, this
bond is undoubtedly much less close than that of the
eutroglophilic European lungless cave salamanders (Ple-
thodontidae). These salamanders reveal well-developed
troglophily, despite being found in both cavities and sur-
face habitats, such as rock outcrops, dry-stone walls, etc.
(Vignoli et al., 2008; Lunghi et al., 2015).
Almost in all cavities, where amphibians were found,
numerous invertebrates were registered as well. Observa-
tions of T. karelinii in the terrestrial phase of its life cycle
characterize it as a nimble and voracious predator with a
broad feeding spectrum, which actively searches for a
prey just like lizards. However, the direct observations of
the amphibian’s diet in caves are lacking, hence, it is
doubtful, whether it continues to feed at low tempe-
ratures deep in underground. At the same time, tailless
amphibians (e.g., marsh frogs and green toads) hunt near
the entrances to caves and grottos. Thus, in stomachs of
three P. ridibundus juveniles collected in Skelskaya Cave
in early spring 2018 (Table 1) no food remnants were
In turn, three Crimean reptile species have never
been recorded in karst. One of them is the European pond
turtle, Emys orbicularis (Linnaeus, 1758) toughly con-
nected to water reservoirs (usually lower than 500 m
a.s.l.) and absent in the open Yaila karst zone — a possi-
ble explanation, why the probability of its entry into the
underground cavities is not so high. Nevertheless, there is
an evidence for the presence of E. orbicularis in an un-
derground lake in Croatia, where it could have entered
Cave Herpetofauna of the Crimea 47
through an underground tunnel about a hundred meter
long (Koller, 2017).
Two thermophilic species of lizards common in the
Crimea, both in the plain area and in the mountains — the
Balkan wall lizard and the glass-lizard, or zheltopuzik,
Pseudopus apodus (Pallas, 1775) have not been found in
the caves as well. Both species in the Crimea prefer gen-
tle slopes with weakly developed karst, usually at alti-
tudes lower than 700 m a.s.l. Despite that fact, some
steppe populations of P. tauricus are distinguished by
their preference of bare rock cliffs and low precipices,
what is always supposed to be the lizards’ specific traits,
since in danger they seek shelter on the rocks and navi-
gate vertically almost as nimbly as D. lindholmi.P. tauri-
cus‘s entries were noted in shallow karst grottos with
wide entry holes, but usually not further than one meter
from the entrance. As for P. apodus, this large anguid liz-
ard was observed only in artificial analogs of caves (road
collectors, various tanks with low overboard, etc.), where
they fall into accidentally. We also observed P. apodus in
rock shelters and at the entrance to the shallow grottos on
the slopes of the gulches and valleys at the Western Foot-
hills. However, such findings are of no interest for the
key issue of the present article.
There is also a lack of reliable evidence for the
snakes from the genus Natrix as well as E. sauromates
and V. renardi entering caves, although some of the spe-
cies mentioned above were registered near the entrances
into the underground cavities (Table 2). Only a few repre-
sentatives of Natrix genus, as it was noticed before, were
recorded on the plateau and on the Crimean Upland’s
In future the dice snakes are likely to be found in
caves along the sides of the river valleys. However, the
vast Crimean foothills are still very poorly studied re-
garding the use of the caves by reptiles, and evidences for
it have to be accumulated. As for the steppe viper, it is a
quite common, though not numerous snake species of the
forest-steppe pre-Yaila slopes at altitudes of about 850 –
900 m a.s.l. Within the last decade the data on its habitats
on the steppe plateaus of the central and eastern uplands
was obtained. In particular, this snake was found at
Mt. Chatyrdag not far from the Suuk-Koba Cave (found
by I. Turbanov, April 13, 2009, 1070 m a.s.l.) and in the
Egiz-Tinakh Hollow in the southeastern part of Karabi
plateau (found by E. Bednarskaya, May 23th, 2009,
1000 m a.s.l.). However, V. renardi abundance on the
mountain plateaus is quite low.
A blotched rat snake is the only species of the Cri-
mean ophidiofauna that never appears in the middle
mountain belt. It has not been registered higher than
600 m a.s.l. both on the northern and southern macro-
slopes. In this regard, it is noteworthy that in the Middle
Valday deposits of the Emine-Bair-Khosar Cave at
Chatyrdag massif (the entrance to the cave is located at
980 m a.s.l.), not only the remains of the representatives
of the regional contemporary ophidiofauna were re-
corded (e.g., C. austriaca and V. cf. renardi), but also
those of the species, which currently do not inhabit the
uplands — Z. situla (Vremir and Ridush, 2005) and E. cf.
sauromates (Ratnikov, 2015).
At the same time, several species of reptiles have
been repeatedly registered in the karst cavities of the Cri-
mea. Among them there are two most thermophilic and
brightly petrophilic species characterized by the predom-
inantly crepuscular and night activities (Crimean gecko
and Leopard snake), which are apparently more related to
cave habitats than others.
The numerous facts of the use of underground shel-
ters by gekkotans (it refers to the infraorder Gekkota)
have to be mentioned here as well. A large number of
cave-dwelling species of a bent-toed geckos of the genus
Cyrtodactylus Gray, 1827 related to the karst regions of
Southeast Asia has been described. Usually, these geckos
are found in dry cave areas and should be described as a
facultative cavernicoles (Bauer et al., 2002) or troglo-
xenes using the caves as a shelter and for the breeding,
but feeding aboveground (Ellis and Pauwels, 2012).
Moreover, various representatives of Phyllodactylidae
and Gekkonidae (including species of the genus Medio-
dactylus Szczerbak et Golubev, 1977) are not rare in
caves of Near East (Iraq Kurdistan, Zagros in Iran) (Afra-
siab and Mohamad, 2009; Esmaeili-Rineh et al., 2016).
Mediterranen house gecko, Hemidactylus turcicus (Lin-
naeus, 1758) has been recorded in caves of Sardinia Is-
land forming colonies of more than 20 individuals in the
moist cave passages in early spring (Rivera et al., 2011).
In the same cavities, a European leaf-toed gecko, Eulep-
tes europaea (Gené, 1838) (Sphaerodactylidae), and
plethodontid salamanders are occurred. Geckos were
found no deeper than 15 m from the entrance — in the
place coinsiding with the concentration of invertebrates.
In turn, an information about the common M. ko-
tschyi findings in the caves of southern Europe and the
Near East is practically absent today. The only record of
M. danilewskii is known from the 1960s at the entrance
of the Orlova Chuka Cave in the Cherny Lom River val-
ley in northern Bulgaria (Koynova et al., 2017). Later
there were no novel findings in this area, and, most likely,
it was a single case of delivery from the Black Sea coast
or from the nearby Danube harbor of Ruse (B. Petrov,
personal communication). Thus, the observation of M.
danilewskii aggregations in the Crimean caves at a quite
high altitudes above the sea level enriches the knowledge
on its biology.
48 Ilya S. Turbanov et al.
In the Crimea, geckos’ habitats reach their maximum
heights on particularly steep and narrow parts of the
coast, which at the same time are good-insulated (Ku-
kushkin, 2004). The hibernation of this species usually
takes place in rock cracks and wall crevices in close prox-
imity to the surface, where geckos form aggregations of
several individuals (up to 12) in one place. Shallow win-
tering shelters sometimes are freezed for short periods,
which usually does not lead to the death of lizards (Ku-
kushkin, 2005). Moreover, some populations of M. dani-
lewskii retain the basic level of activity even during the
wintering period. Geckos, leaving their shelters with the
beginning of darkness at an air and substrate tempera-
tures of 7.2 9.8°C or lower have been regularly ob-
served on the surface of walls or rocks throughout the
cold season. Nevertheless, despite some resistance to low
temperatures, according to the biotope nature and habitat
type, M. danilewskii is the most thermophilic species of
the Crimean herpetofauna. Although during the extreme-
ly cold winters with the prolonged frosty periods geckos
die on hibernation. Sometimes the overall frost devasta-
tion of local populations in the northernmost areas of the
range has been observed (Kukushkin, 2007). Therefore,
the usage of the karst cavities with the inert microclimate
for wintering should definitely increase the probability of
geckos survival during the cold season — especially near
the upper limit of species distribution. The deep gravige-
nic separation cracks also contribute to geckos upward
movement on the Yaila cliffs and isolated rocks. Lizards
find favorable conditions for wintering in caves and deep
crevices with a stable positive temperatures and the abun-
dance of invertebrates for the occasional feeding. Under
the experimental conditions, the hunting of the Crimean
geckos for the spiders has been observed even in January
in the dark at temperatures above 10.5°C (Kukushkin,
2005). This temperature threshold level corresponds to
the winter microclimate of the caves, where geckos ag-
gregations were found. However, their aggregations in
karst cavities with the sources of condensate have been
also observed during the hottest period of the year, while
in winter they were not found here (Table 2). Altogether,
it can be concluded that the part of the lizard’s population
uses underground shelters on purpose to optimize their
living conditions not only during the cold season, but also
in periods of extreme drought.
Among the Eurasian ophidiofauna representatives,
troglophily is a quite rare phenomenon. In relation to the
cave habitats, few Colubridae species have been men-
tioned, and the most frequently a striped racer, or a
cave-dwelling ratsnake, Orthriophis taeniurus (Cope,
1861), and a Moellendorff’s trinket snake, O. moellen-
dorfi (Boettger, 1886) (McClur et al., 1967; Zhou et al.,
2012). Comparatively regular Z. situla findings in the
Crimean karst suggest that in some localities it is both a
cave visitor and facultative cavernicole, which systemati-
cally uses karst cavities for wintering, molting, and prob-
ably feeding. This snake, characterized mainly by the
twilight-night activity during the hot season with the
preferance of the rocky habitats (Kukushkin and Tsve-
lykh, 2004) is pre-adapted for living in caves to some
extent as the Crimean gecko.
A Caspian whipe snake as the diurnal species with a
well-developed climbing ability, supllements well the list
of the potentially nocturnal species of reptiles mentioned
above. It has been established that D. caspius visits caves
in search of prey or can use them for wintering. In shal-
low caves these snakes are able to feed by bats and ro-
dents, whose burrows can be often met in cave alluvium,
and also, probably, geckos (hunting D. caspius on M. da-
nilewskii has been observed repeatedly — on walls,
rocks, trunks of fallen trees, etc.).
Other snakes species of the Crimean fauna in most
cases find themselves in the underground cavities acci-
dentally and, if they do not escape, die there. First of all,
this is true for C. austriaca, the most common mainly
saurofag species in the open Yaila karst areas, which may
fall into the cracks and karst holes while hunting lizards.
Another conclusion concerns the origin of fossil re-
mains of herpetofauna in the underground cavities of the
Crimea. Usually the findings of the reptile fossils of late
Pleistocene and Holocene age in caves and grottos are as-
sociated with the activity of predators — diurnal birds of
prey or owls (Burchak-Abramovich, 1936; Gromov,
1961; Vremir and Ridush, 2005). Based on our data, the
possibility of this is not excluded, however, the extent of
reptile’s entry to the caves during their active phase of
life cycle should not be underestimated and it can be
quite significant. The aditional evidence for this is the
brand new record of the full skeleton of colubrid snake
found near the entrance to the newly discovered Tavrida
Cave in July 2018 (Fig. 7b).
Therefore, these ecological characteristics may be
the background for the systemic use of cave habitats by
amphibians or reptiles: petrophylity or, at least, a well-
developed climbing skills; mainly twilight-night type of
activity related to the ability to remain active at a rela-
tively low temperatures; a sort of time budget, when the
animal spends most of the time in shelters with a rela-
tively low amplitude of diurnal fluctuations of tempera-
ture and humidity.
It is evident that the current data on the usage of un-
derground habitats in the Crimean Mountains by amphib-
Cave Herpetofauna of the Crimea 49
ians and reptiles is still insufficient and mostly frag-
mented in many respects. However, even at this stage it
seems possible to draw several conclusions.
The majority of the Crimean herpetofauna represen-
tatives are trapped in underground cavities occasionally
and can be categorized as accidental trogloxenes. More-
over, several amphibian and reptile species of the Cri-
mean Peninsula enter the karst cavities on purpose and
systematically use them, what allows to categorize them
as facultative cavernicoles. Among the amphibians, the
closest ecological relation to the karst is revealed by the
T. karelinii, though its nature remains unclear, as well as
of B. viridis. Apparently, amphibians enter underground
habitats for the survival during the dry period, recovery
of water balance within the hot season, wintering and
feeding in the entrance parts of the caves.
Among the reptiles, M. danilewskii and Z. situla are
considered to be the species, associated with the under-
ground habitats to high extent. The use of cavities by rep-
tiles (mainly relic caves of karst origin with the warm-
type ventilation) might be related with the optimization
of living conditions in winter and summer as well as with
molting and feeding.
The intensity of underground cavities usage might be
determined by the conditions of the specific locality.
Thus, the mesophilous amphibian species enter karst
more actively in arid zones with a harsh climate and poor
vegetation (T. karelinii and H. orientalis), while the most
thermophilic reptile species (M. danilewskii and Z. situ-
la) — near the upper limit of their range.
Some parallels between the synatropization degree of
the species and the preference of the natural underground
cavities should be also noticed. Among the amphibians
this relation is clearly visible on B. viridis, while on rep-
tiles — on M. danilewskii and Z. situla.
For accidental trogloxenes, there is a relationship be-
tween the species entry into the underground cavities and
its presence and population density in open karst zone.
For instance, the most frequently recorded snake species
in the Crimean Yaila cavities is C. austriaca, which is, at
the same time, the most abundant species there.
In mountainous areas, speleoresearch plays an im-
portant role for the establishment of the upper limit of
distribution and detalization of batracho- and herpeto-
fauna species range. In this regards, the cases of B. viridis
and M. danilewskii are quite illustrative, since the highest
finding of a green toad in the Crimea was made in a karst
cave at Babugan Yaila, while a single finding of Danilew-
ski’s gecko in a grotto on the Karadag coast triggered
more thorough study of this area resulted in the identifi-
cation of large population of this lizard of exclusive sci-
entific importance.
At the moment, the hottest issue of the Crimean spe-
leozoology is the study of the amphibians’ (mainly
newts) episodic reproduction in the underground habitats
and trophic bonds of both amphibians and reptiles visit-
ing caves regularly. In this regards, the snakes feeding on
bats is the issue of a particular interest.
Acknowledgments. The authors are sincerely grateful to
friends and colleagues for the information on amphibians and
reptiles findings in karst cavities, assistance in field research
and professional advices: Aleksey Akimov (Association of the
Speleologists of Sevastopol), Sergey Arefiev (Sevastopol),
Yulia Balakhtinova (Speleological club “Zelenye Kamneedy,”
Sevastopol – Yalta), Elena Bednarskaya (Denisova) (Simfero-
pol), Svetlana Bogdanenko and Larisa Bogdanenko (Sevasto-
pol), Elena Gladilina and Aleksey Ivanov (State historical-ar-
chaeological Museum-Reserve “Tauric Chersonese,” Sevasto-
pol), Maksim Grishkov (Sevastopol), Maksim Kashirsky (“Ki-
zil-Koba” company, Simferopol), Sergey Kebets (Speleological
club “Zelenye Kamneedy,” Sevastopol), Sergey Klushin (Spe-
leological club “Zelenye Kamneedy,” Yalta), Sergey Kostin
(Yalta), Gennadyi Krepakov (Sevastopol tourist club), Alek-
sandra Kurichenko (Association of the Speleologists of Sevas-
topol), Sergey Leonov (Simferopol), Boris Matyushkin(Seva-
stopol), Ksenia Moskvina (Sevastopol), Pavel Moroz (Kyiv),
Anton Nadolny (Sevastopol), Valentin Nuzhdenko (Simfero-
pol), Pavel Oksinenko and Yevgeny Oksinenko (Simferopol),
Aleksey Papyi (Yalta speleosection), Oleg Peretyatko (Large
Sevastopol speleosection), Boyan Petrov (National Museum of
Natural History, Bulgarian Academy of Sciences, Sofia), Vasil
Pokinchereda (Carpathian Biosphere Reserve of National
Academy of Sciense of Ukraine, Rakhiv), Roman Potapov
(Sevastopol tourist club), Grigory Prokopov (Simferopol),
Andrey Shendrik (Sevastopol tourist club), Ruslan Svetlov
(Speleological club “Karst”, Simferopol), Elena Sviridenko
(Alushta – Kyiv), Dmitryi Tikhomirov (Association of speleo-
logists of Sevastopol), Sergey Tokarev (Simferopol), Aleksandr
Trofimov (Sevastopol), Anton Yushko (St. Petersburg), Yevge-
ny Yanovsky (Association of speleologists of Sevastopol). Spe-
cial thanks to Yuliya Krasylenko (Olomouc, Czech Republic),
Natalia Aschepkova (Morbegno, Italy), and Anastasia Aschep-
kova (Moscow) for the translation and proof-read of the English
version of the manuscript.
This research was covered by the state assignment
of FASO of Russian Federation (topics No. AAAA-A17-
117030310017-8, AAA-A16-116022510087-5, AAAA-A18-
118012690106-7, AAAA-A18-118012690105, and AAAA-
A19-119012490044-3) and partially supported by RFBR (pro-
ject No. 17-54-40017 Abkh_a).
Afrasiab S. R. and Mohamad S. I. (2009), “A study on cave-
dwelling geckos in Iraq, with the description of a new spe-
cies from Saffine mountain (Reptilia; Gekkonidae),” Zool.
Middle East,47,49–56.
50 Ilya S. Turbanov et al.
Alekseev E. V. (2008), Bloodsucking and Venomous Arthro-
pods of the Crimea, DIAYPI, Simferopol [in Russian].
Amelichev G. N., Klimchouk A. B., Tokarev S. V., and Me-
metova E. I. (2014), “The Cadaster of karst caverns of the
Crimea: past, present and future,” Geopolit. Ékogeodin.
Reg.,10(1), 345 – 351 [in Ukrainian].
Averinov A. O. (2001), “New records of proteid salamanders
(Amphibia, Caudata) from the Pliocene of Ukraine and
lower pleistocene of Moldova,” Vestn. Zool.,35(1), 43 – 46.
Balogová M., Apfelová M., Flajc T., Jablonski D., Kaut-
man J., Krišovsky P., Krištín A., Papáè V., Puchala P.,
Urban P., and Uhrin M. (2015), “Distribution of the fire
salamander (Salamandra salamandra) in Slovakia,” Folia
Faun. Slovaca,20(1), 67 – 93.
Balogová M., Jeliæ D., Kyselová M., and Uhrin M. (2017),
“Subterranean systems provide a suitable overwinterings
habitat for Salamandra salamandra,” Int. J. Speleol.,46(3),
321 – 329.
Bauer A. M., Pauwels O. S. G., and Chanhome L. (2002),
“A new species of cave-dwelling Cyrtodactylus (Squamata:
Gekkonidae) from Thailand,” Nat. Hist. J. Chulalongkorn
Univ.,2(2), 19 – 29.
Birstein Ya. A. (1963), “Some results of the study of subterra-
nean fauna of the Crimea,” in: Collection of scientific pa-
pers of the complex karst expedition of Academy of Sciences
of UkrSSR. Vol. 1. Investigation of the karst of the Crimea,
Izd. AN UkrSSR, Kiev, pp. 123 – 134 [in Russian].
Brauner A. (1904), “A brief guide of reptiles and amphibians
of the Crimea and steppe belt of European Russia,” in: Bul-
letin of the Crimean Alpine club. Vols.3–9, A. F. Sokolov-
sky’s Tipo-Lithography, Odessa,1–64[inRussian].
Burchak-Abramovich M. O. (1936), “Remains of Pelobates
fuscus (Laur.) from the karst cave at Karabi-Yaila (the Cri-
mea),” in: Collection of scientific paper of Zoological Mu-
seum of Ukrainian Academy of Sciences. Vol. 16 (Pr. Inst.
Zool. Biol.,11), 129 – 133 [in Ukrainian].
Caves. Information retrieval system. (ac-
cessed on August 17, 2018).
Dublyansky V. N. (1966), “Age of the deep karst cavities
of the Mountain Crimea,” Peshchery,6(7), 70 – 81 [in
Dublyansky V. N. (1977), Karst Caves and Mines of the
Mountain Crimea (Genesis, Sediments, Hydrogeological
Significance), Nauka, Leningrad. otd., Leningrad [in
Ellis M. and Pauwels O. S. (2012), “The bent-toed geckos
(Cyrtodactylus) of the caves and karst of Thailand,” Cave
Karst Sci.,39(1), 16 – 22.
Esmaeili-Rineh S., Akmali V., Fathipour F., Heidari N., and
Rastegar-Pouyani N. (2016), “New distribution records of
cave-dwelling gekkonid lizards (Sauria, Gekkonidae and
Phyllodactylidae) in the Zagros Mountains of Iran,”
Subterr. Biol.,18,39–47.
Fordham R. A. (1985), “Live Litoria raniformis (Anura: Hyl-
idae) in New Zealand caves: a ‘smuggler’s’ dispersal
route?” N. Zeeland J. Zool.,12(4), 561 – 563.
Frost D. R. (2018), Amphibian Species of the World: an Online
Reference. Ver. 6.0,
amphibia/index.html (accessed on August 17, 2018).
Garrido-García J. A., Schreur G., and Pleguezuelos J. M.
(2013), “Occasional bat predation by the horseshoe whip
snake (Reptilia, Colubride),” Galemys,25,59–61.
Gromov I. M. (1961), “Fossil Upper Quaternary rodents of the
premontane Crimea,” Nauch. Tr. Komis. Izuch. Chetvert.
Period,17, 1 – 190 [in Russian].
Ianc R., Cicort-Lucaciu A.-ª.,Ilieº D., and Kovács É.-H.
(2012), “Note on the presence of Salamandra salamandra
(Amphibia) in caves from Padurea Craiului Mountains, Ro-
mania,” N.-W. J. Zool.,8(1), 202 – 204.
Kaòuch P. and Bala P. (2005), “Bat as a prey of Elaphe
longissima (Laurenti, 1768),” Herpetozoa,18(1/2), 92 –
Koller K. (2017), “Undeground occurences of three species of
amphibians and reptiles with special emphasis on Rana la-
tastei (Amphibia: Anura),” N.-W. J. Zool.,13(1), 176 – 179.
Kondaraki V. Ch. (1875), Universal Description of the Cri-
mea. Part 7. Herptiles (Reptilia s. amphibia), V. Velling
printing house, St. Pertersburg, pp. 35 – 37 [in Russian].
Kotenko T. I. (2010), “Amphibians and reptiles of the Cri-
mea,” in: Scientific Notes of the ‘Cape Martyan’ Nature Re-
serve. Vol. 1. Nikitsky Botan. Garden — National Scientific
Center, Yalta, pp. 171 – 224 [in Russian].
Kotenko T. I. and Kukushkin O. V. (2010), “Annotated lists
of amphibians and reptiles of the Crimean Nature Re-
serves,” in: Scientific Notes of the ‘Cape Martyan’ Nature
Reserve. Vol. 1. Nikitsky Botan. Garden — National Scien-
tific Center, Yalta, pp. 225 – 261 [in Russian].
Koynova T., Tzankov N., Popgeorgiev G., Naumov B., and
Natchev N. (2017), “A new distribution record of the
Kotschy’s Gecko (Mediodactylus kotschyi) from inland
north-eastern Bulgaria,” Herpetol. Notes,10,1–2.
Kukushkin O. V. (2004), “Distribution, habitat allocation and
abundance of the Kotschy’s (Crimean) gecko, Cyrtopodion
kotschyi danilewskii (Strauch, 1887) (Reptilia: Squamata:
Gekkonidae) in the Southern Crimea,” in: Karadag. His-
tory, Geology, Botany, Zoology. Vol. 1, SONAT, Simferopol,
pp. 367 – 396 [in Russian].
Kukushkin O. V. (2005), “A duration of winter diapause and
peculiarities of biology of the Crimean Kotschy’s gecko
(Mediodactylus kotschyi danilewskii) during a hibernation
period. Current issues of ecology and zoology,” in: Proc. of
the International scientific commemorative conference ded-
icated to the 140th anniversary of I. I. Mechnikov Odessa
National University and the 120th anniversary of the birth
of prof. I. I. Puzanov, 22 – 25 April 2005, Odessa, Feniks,
Odessa, pp. 148 – 151 [in Russian].
Kukushkin O. V. (2009), “About inhabitation of Kotschy’s
Naked-toed Gecko, Mediodactylus kotschyi danilewskii
(Reptila: Sauria: Gekkonidae), in the middle forest belt of
southern macroslope of the Crimean Mountains,” Tr. Ukr.
Gerpetol. Obshch.,2, 27 – 36 [in Russian].
Kukushkin O. V. (2007), “Data on cold tolerance during hi-
bernation in the Crimean Kotschyi’s [sic!] Gecko,” in:
Programme & Abstr. of the First Mediterranean Herpetol.
Cave Herpetofauna of the Crimea 51
Congress (CMH1), 16 20 April 2007, Marrakech,
pp. 88 – 89.
Kukushkin O. V. and Kuschan N. B. (2015), “Materials to
the study of Karelin’s newt (Amphibia, Caudata, Salaman-
dridae) in the Crimea,” in: Proc. of the 2nd All-Russian
Sci.-Pract. Conf. Sustainable development of specially
protected natural territories”. Vol. 2, 2 – 4 December 2015,
Donizdat, Sochi, pp. 141 – 151 [in Russian].
Kukushkin O. V., Petrov B. P., Nazarov R. A., and Mel-
nikov D. A. (2016), “The problem of a biogeographical sta-
tus of two reptiles with narrow distribution in the Crimean
Mountains and the importance of Cape Aiya karst caves for
its solution,” in: Proc. of 2nd All-Russian conference of
young scientists “Biospeleological Studies in Russia and
adjacent countries,” 1 2 December 2016, Moscow, Fili-
gran’, Yaroslavl, pp. 56 – 69 [in Russian].
Kukushkin O. V. and Tsvelykh A. N. (2004), “Distribution
and ecological-morphological peculiarities of the leopard
snake, Elaphe situla (Serpentes, Colubridae), in the Cri-
mea,” Zool. Zh.,83(4), 439 – 448 [in Russian].
Lunghi E., Manenti R., and Ficetola G. F. (2015), “Seasonal
variation in microhabitat of salamaders: environmental
variation or shift of habitat selection?” PeerJ. https://
Manenti R., Ficetola G. F., Bianchi B., and De Bernardi F.
(2009), “Habitat features and distribution of Salamandra
salamandra in underground springs,” Acta Herpetol.,4,
143 – 151.
Manenti R., Ficetola G. F., Marieni A., and De Bernardi F.
(2011), “Caves as breeding sites for Salamandra salaman-
dra: habitat selection, larval development and conservation
issues,” N.-W. J. Zool.,7, 304 – 309.
Matyushkin B. (2010), “From the order Tailed Amphibians,”
Svet: Ukr. Speleol. Assots. J.,1(36), 48 – 52 [in Russian].
Maksimovich G. A. (1963), Principles of Karstology. Vol. 1.
Issues of the Karst Morphology, Speleology and Hydro-
geology of Karst, Perm. Knizh. Izd., Perm’ [in Russian].
McClur H. E., Lim B.-L., and Winn S. E. (1967), “Fauna of
the Dark Cave, Batu Caves, Kuala-Lumpur, Malaysia,” Pa-
cific Insects,9(3), 399 – 428.
Muratov M. V. and Nikolaev N. I. (1940), “Quaternary his-
tory and development of relief of the Mountain Crimea,”
Uch. Zap. MGU,48, 65 – 73 [in Russian].
Muratov M. V. (1954), “About Miocene and Pliocene history
of development of the Crimean peninsula,” Byull. Mosk.
Obshch. Estestvoisp. Otd. Geol.,29(1),4–20[inRussian].
Muratov M. V. (1960), A Brief Essay of Geological Structure
of the Crimean Peninsula, Gos. Nauch.-Tekh. Izd. Liter. po
Geol. i Okhrane Mineral. Resursov, Moscow [in Russian].
Naucke T. J. (2002), “Leishmaniose, eine Tropenkrankheit
und deren Vektoren (Diptera, Psychodidae, Phlebotominae)
in Mitteleuropa,” Denisia 6. Zugleich Kataloge des OÖ
Landesmuseums,184, 163 – 178.
Ostrovskikh S. V. (2012), “Amphibians (Amphibia) and rep-
tiles (Reptilia),” in: I. N. Safronov, P. I. Bukharitsyn, and
A. V. Barmin (eds.), Contemporary State and Long-Term
Changes of the Environment at Bogdinsko-Baskunchakskyi
Reserve Territory, Tsaritsyn, Volgograd, pp. 174 – 182 [in
Ostrovskikh S. V., Pestov M. V., and Gnetneva A. N. (2015),
“Reptiles (Reptilia) of the ‘Utrish’ Reserve and adjacent
areas,” in: G. N. Ogureeva and K. B. Gongalsky (eds.),
Protection of Biota in the State Nature Reserve “Utrish.”
Sci. Works. Vol. 3, Poligraf-YUG, Maykop, pp. 272 – 296
[in Russian].
Parin N. V. (1983), “Noemachelius (Troglocobitis)starostini
sp. n. — new cave fish from subterranean waters of Kugi-
tangtau (Turkmenia),” Zool. Zh.,62(1), 83 – 89 [in
Perfiliev P. P. (1941), “Data on the sandfly fauna of the
USSR,” Nauch. Zap. VMA im. S. M. Kirova RKKA,25,
272 – 283 [in Russian].
Pokinchereda V. F. (1989), “Records of amphibians in caves
and galleries of the Carpathians,” in: Abstrs. of the 7th Her-
petol. Conf. “The problems of herpetology,” 26 – 29 Sep-
tember 1989, Naukova Dumka, Kiev, p. 198 [in Russian].
Puzanov I. I. (1931), “Preliminary results of the study of ver-
tebrates fauna of the Crimean Reserve,” in: Collection of
the papers on the study of the fauna of Crimean State Re-
serve, Gosmedizdat, Moscow – Leningrad, pp.5–38 [in
Pysanets E. and Kukushkin O. (2016), Amphibians of the
Crimea, NAS of Ukraine, National Museum of Natural His-
tory, Kyiv [in Ukrainian; in English].
Ratnikov V. Yu. (2010), “A Review of Tailed Amphibian Re-
mains from Late Cenozoic sediments of the East European
Plain,” Russ. J. Herpetol.,17(1), 59 – 66.
Ratnikov V. Yu. (2015), “Squamate reptiles from Upper Pleis-
tocene deposites of the cave Emine-Bair-Khosar in the Cri-
mea,” in: Proc. of 1st All-Russian conference of young sci-
entists “Biospeleology of the Caucasus and other regions of
Russia,” 3 – 4 December 2015, Moscow, Kostromskoy pe-
chat. dom, Kostroma, pp. 64 – 65 [in Russian].
Rivera X., Carranza S., Amat F., Romano A., and Sotgiu G.
(2011), “On the presence of Hemidactylus turcicus (Lin-
naeus, 1758) in caves and mines of the island of Sardinia,
Italy,” Buttl. Soc. Cat. Herpetol.,19,61–65.
Sket B. (2008), “Can we agree on an ecological classification
of subterranean animals?” J. Nat. Hist.,42(21 – 22), 1549 –
Sket B. (2017), “Discoveries the black proteus Proteus angiu-
nus parkelj (Amphibia: Caudata),” Natura Sloveniae,19(1),
27 – 28.
Sharygin S. A. (1998), “To the study of amphibians of South-
ern Coast of the Crimea,” in: Discovery of diversity of ani-
mal kingdom: Scientific papers of Zoological museum of I.
I. Mechnikov, Odessa State University, Vol. 3, pp. 51 – 55
[in Russian].
Sotgiu G., Angelini C., Bovero S., and Tessa G. (2017),
“Unusual egg deposition sites of Euproctus platycephalus
(Gravenhorst, 1829),” Herpetozoa,30(1/2), 66 – 68.
Szczerbak N. N. (1966), Amphibians and reptiles of the Cri-
mea. Herpetologia Taurica, Naukova Dumka, Kiev [in
Szczerbak N. N. (1984), “Amphibians and reptiles of Kara-
dag,” in: Study of the fauna and abundance of terrestrial
vertebrates of Karadag (final report). Annals of Karadag
52 Ilya S. Turbanov et al.
State Reserve of Academy of the Sciences of UkrSSR,1(5),
4 – 32 [in Russian].
Taraschuk V. I. (1959), “Amphibians and reptiles,” in: Fauna
of Ukraine. Vol. 7. Izd. AN UkrSSR, Kyiv [in Ukrainian].
Trajano E. and de Carvalho M. R. (2017), “Towards a bio-
logically meaningful classification of subterranean organ-
isms: a critical analysis of the Schiner-Racovitza system
from a historical perspective, difficulties of its application
and implications for conservation,” Subterr. Biol.,22,
1 – 26.
Turbanov I. S., Oksinenko P. V., and Kukushkin O. V.
(2015), “Records of reptiles in karst cavities of Mountain
Crimea,” in: Proc. of 1st All-Russian conference of young
scientists “Biospeleology of the Caucasus and other re-
gions of Russia,” 3 – 4 December 2015, Moscow, Kostrom-
skoy pech. dom, Kostroma, 90 – 94 [in Russian].
Turbanov I. S., Palatov D. M., and Golovatch S. I. (2016a),
“The state of the art of biospeleology in Russia and other
countries of the former Soviet Union: A review of cave
(endogean) invertebrate fauna. 1. Introduction — Crusta-
cea,” Entomol. Rev.,96(7), 926 – 963.
Turbanov I. S., Palatov D. M., and Golovatch S. I. (2016b),
“The state of the art of biospeleology in Russia and other
countries of the former Soviet Union: A review of cave
(endogean) invertebrate fauna. 2. Arachnida — Acknowl-
edgements,” Entomol. Rev.,96(9), 1297 – 1333.
Turbanov I. S., Palatov D. M., and Golovatch S. I. (2016c),
“The state of art in biospeleology in Russia and other coun-
tries of the former Soviet Union: A review of the cave
(endogean) invertebrate fauna. 3. References,” Entomol.
Rev.,96(9), 1334 – 1358.
Uetz P., Freed P., and Hošek J. (2018), The Reptile Database, (accessed on August 17,
Vakhrushev B. A. and Amelichev G. N. (2001), “On the pos-
sibility of glaciation of the Crimean mountains,” Fiz.
Geogr. Geomorfol.,40, 134 – 153 [in Russian].
Vakhrushev B. A. (2009), “Regionalization of karst of the Cri-
mean Peninsula,” Speleol. Karstol.,3, 39 – 46 [in Russian].
Vignoli L., Caldera F., and Bologna M. A. (2008), “Spatial
niche of the Italian cave salamander, Speleomantes italicus
(Dunn, 1923) (Plethodontidae, Amphibia), in subterranean
system of Central Italy,” Ital. J. Zool.,75(1), 59 – 65.
Voinstvensky M. A. (2006), “Diaries of Crimean expeditions
in 1957 and 1958,” in: M. M. Beskaravayniy (ed.): Avifauna
of Ukraine.Vol.3,pp.2–40[inRussian].
Vremir M. M. and Ridush B. (2005), “The Emine-Bair-
Khosar ‘Mega Trap’(Ukraine),” Mitteilungen der Kommiss.
fur Quartärforsch. Österreich. Akad. der Wissensch.,14,
235 – 239.
Yena V. G., Yena Al. V., and Yena An. V. (2004), Protected
Landscapes of Taurida [in Russian], Biznes-Inform,
Zagorodniuk I. (2004), Cave Fauna of Ukraine. Ser. Proc. of
the Theriological School. No. 6, Kyiv [in Ukrainian].
Zhou Z., Lau M., and Nguyen T. Q. (2012), “Orthriophis
moellendorfi. The IUCN Red List of Threatened Species
2012,” DOI: 10.2305/IUCN.UK.2012-1.RLTS.
Cave Herpetofauna of the Crimea 53
... The present study is the first review of the herpetofauna of the Greater Caucasian caves. As far as the adjacent territories are concerned, a detailed survey of the cave herpetofauna has been published only for the Crimea, where four species of amphibians and eight species of reptiles were recorded (Turbanov et al., 2015(Turbanov et al., , 2019Kukushkin et al., 2017). In Western Ukraine (Carpathians and Podolia), five species of amphibians are shown to be common in caves (Krochko, 1973;Zagorodniuk & Petrushenko, 2003;Vargovitsh & Monich, 2004;Zagorodniuk & Vargovitsh, 2004). ...
... In 1990s-2000s we observed the accumulation of wintering adults of Salamandra salamandra Linnaeus, 1758 (up to 35 specimens) in the underground cavities of the Ukrainian Carpathians, located near the town of Mukachevo (Turbanov et al., 2019). Also their larvae were regularly observed, for several years, in a semi-submerged adit in vicinity of the village Hlyboke near Uzhgorod, which indicates the apparent propensity of this species to troglophily. ...
... Similar observations were carried out in the Crimea. In this region, Mediodactylus kotschyi danilewskii (Strauch, 1887) and Zamenis situla (Linnaeus, 1758) have been observed in small warm caves while wintering, as well as seeking for shelter at other periods of the year, and sometimes looking for food (Turbanov et al., 2015(Turbanov et al., , 2019. ...
Full-text available
The present study is the first review of the recent herpetofauna of the Greater Caucasian caves. Also, it is worth noting some phossylous material on amphibians and reptiles (mainly Pleistocene remains) from the caves in this area. The goal of the study is to fill knowledge gaps and to present long-term data on the cave herpetofauna of the Greater Caucasus. In 1992–2019, amphibians and reptiles were found in 61 Greater Caucasian caves located in an altitude range from 55 m a.s.l. to 1300 m a.s.l. in Russia, Abkhazia, and Georgia. A total of 272 specimens representing ten species have been found. The most frequently observed species were Rana macrocnemis (39.0% of all recorded specimens), Pelodytes caucasicus (17.3%), Bufo verrucosissimus (11.0%), Darevskia brauneri (8.8%), Hyla orientalis (8.1%). The total proportion of the remaining five species (Ommatotriton ophryticus, Hyla savignyi, Pelophylax ridibundus, Anguis colchica, and Darevskia derjugini) was 15.8%. As a rule, representatives of the herpetofauna in the Greater Caucasian caves are quite rare and only single or few individuals are usually found. Exceptions were Fyodorovskaya Cave (Black Sea Region near the city of Sochi), Akshasha Cave, Abshdza Cave and Marshania Verkhnyaya Cave (last three in Abkhazia) where 141 specimens of amphibians and reptiles were observed, i.e. more than half (51.8%) of the totally recorded specimens. They belong to eight species (80.0% of all species found). The caves of the Greater Caucasus foothills, where the vast majority of amphibians and reptiles were observed, are usually characterised by a rich faunal diversity of invertebrate animals – their potential food. Amphibians were recorded from both twilight and deep parts of the caves, whereas lizards were found mainly close to the entrances, where the light penetrates and a relatively high air temperature is observed in the warm period of the year. Probably, some of the Greater Caucasian amphibians and reptiles enter the caves purposefully for wintering, and/or for feeding. The other animals occur accidentally in the caves (especially on the bottom of vertical pits), becoming «prisoners of circumstances». Judging by the length of time spent in the caves and, as a consequence, the change in body pigmentation (lighting of the main colouration tone) and good external condition of many individuals, these amphibians can be considered subtroglophiles. Rana macrocnemis, capable of laying eggs in caves, shows a tendency for eutroglophily.
... The "Baydarskyi", "Cape Aya" and "Laspi" state regional wildlife sanctuaries play the most signi ficant role in preserving the herpetofauna of the region, covering the upper part of the Chernaya River basin (Main Range) and the extreme southwestern part of the Southern Coast of Crimea, as well as the Mekenzievskoe Forestry in the foothills. Kukushkin, O.V. et al., 2019. Ecosystem Transformation 2 (4), 4-62. ...
... The exact location for each animal was recorded, indicating the geographical coordinates, in the WGS-84 system and applied to an electronic map of 1:200 000 scale, and the outlines of everyday routes were displayed. It was the first time that such a significant amount of targeted Kukushkin, O.V. et al., 2019. Ecosystem Transformation 2 (4), 4-62. ...
... 66-PP dated February 8, 2018 "On the Establishment of the "Laspi" state nature landscape sanctuary of regional importance". The boundaries of protected areas were Kukushkin, O.V. et al., 2019. Ecosystem Transformation 2 (4), 4-62. ...
... The number of geckos in the upper part of Cape Aya in the section from Mount Kalafatlar to Mount Kush-Kaya is quite high even at altitudes of 500-660 m above sea level and, in any case, not lower than the values of population density we recorded in the altitude range 200-250 m above sea level (for example, in the rocks under the cliffs of the western part of the Ai-Petri Yayla or in the Choban-Tash Rocks over Cape Sarych) (Kukushkin, 2004a(Kukushkin, , 2009aTurbanov et al., 2019). ...
... Individuals of Pe. ridibundus s. str., identified using molecular markers, were found only in the upper reaches of a number of the most isolated tributaries of the Chernaya River (mainly in canyons, which for most of the year were cut off from the intermontane hollows by long sections of dry channels), some small forest lakes located under the forest canopy, as well as in the Skelskaya Cave Turbanov et al., 2019). The altitude of the localities is 295-815 m above sea level, an average of 467 ± 52.5 m. ...
... Features of amphibian and reptile spatial distribution in Sevastopol city territory. The maximum heights of the species records in the whole Crimea are indicated in parentheses (by:Turbanov et al., 2019). ...
... и, во всяком случае, не ниже значений, отмеченных нами в диапазоне высот 200-250 м н.у.м. (например, в скалах под обрывами западной части Ай-Петринской яйлы или в урочище Чобан-Таш над мысом Сарыч) (Кукушкин, 2004aTurbanov et al., 2019). ...
... Особи Pe. ridibundus s. str., выявляемые с использованием молекулярных маркеров, обнаружены только в верхнем течении ряда наиболее изолированных притоков р. Черная (главным образом в каньонах, которые на протяжении большей части года отрезаны от межгорных котловин протяженными участками сухих русел), в некоторых лесных озерцах, расположенных под пологом леса, а также в Скельской пещере (Кукушкин и др., 2018;Turbanov et al., 2019). Высота локалитетов составляет 295-815 м н.у.м., в среднем -467 ± 52.5 м. ...
... The use of subterranean cavities by amphibians in the Black Sea Region was studied by some researches. Turbanov et al. (2019) reported amphibians and reptiles in the subterranean cavities of the Crimean Mountains. In addition, Tuniyev (2017) provided data on the occurrence of amphibians in the Samshitovaya Cave in Colchida historical area. ...
Full-text available
An inventory belonging to knowledge of the amphibian fauna is provided from the caves in Zonguldak province, western Black Sea Region of Turkey. No systematic field survey of amphibians in Turkish cave environments has yet been performed. The number of known caves in the western Black Sea region is approximately 280 and the province of Zonguldak ranks second with 41 caves in this region. We conducted our field surveys in 31 caves of Zonguldak during the period in 2017 – 2019 and our results revealed occurrences of four amphibian species. Dedicated surveys using appropriate methods will increase the number of amphibian species in the caves.
... Also, a relatively rich sandfly fauna was found in the coastal district of Bar, Montenegro with the presence of Ph. papatasi, Ph. perfiliewi, Ph. tobbi, Ph. neglectus and S. minuta, which area has a typical Mediterranean coastal karstic landscape (Ivović et al. 2003). In Crimea, sandfly species feed on lizards that reside in caves on the southern coast of the peninsula (Turbanov et al. 2019). These observations confirm that several Mediterranean sandfly species can colonise the relatively dry environment. ...
Full-text available
The Oligocene and Miocene tectonic and biogeographical alterations of the peri-Mediterranean area could strongly impact the speciation processes and migrations of Mediterranean sandfly species. To understand the possible former role of this palaeobiogeographic factor on ancestral sandfly species, the potential suitability values of five Larroussius , two Paraphlebotomus and one Phlebotomus species were modelled from the Rupelian to the Tortonian stage in the Central Paratethys. The bioclimatic extrema of this sandfly species and the Coexistence Approach-based palaeoclimatic reconstructions made the basis of hypothesis testing. In the case of the Tortonian stage, a georeferenced climatic model was created. The models indicate that the suitability values could notably vary by species and periods. The monsoon-influenced humid subtropical climate of the Oligocene epoch could be less suitable for the ancestors of Mediterranean sandfly taxa than the later, drier humid subtropical climates-characterised Miocene stages. The Central Paratethys area could be less suitable for ancestors of the extant Paraphlebotomus , and Phlebotomus in the Miocene epoch compared to Larroussius species. It may indicate that the Central Paratethys formed a barrier against the east-to-west migration of the ancestors of Phlebotomus and Paraphlebotomus species . In contrast, Larroussius species could prefer the Miocene climate of the Central Paratethys. In the Tortonian stage, mainly the coastal areas of the sea could be colonised by sandflies. These results indicate that the coasts of the Central Paratethys should be not only considered as a potential former migration route but could be a part of the speciation area of Larroussius sandflies.
... In general, information on the repose and summer roost of frogs in European caves is scarce. The only recent detailed overview of the herpetological fauna covers karst cavities of the Crimean Mountains (Turbanov et al. 2019). Sightings of R. graeca were reported only from entrances to three caves in Bulgaria (Western Rhodopes Mts.: entrances to Uhlovitsa and Gargina Dupka caves near Mostovo village, and the entrance to the mine gallery at Dzurkovo village) (Petrov et al. 2006). ...
Full-text available
The subterranean fauna of both natural and artificial caves in Albania remains poorly studied. Cave colonisation can be clarified by investigating the same process in artificial underground sites. The existence of such a process can be judged by the composition of specific vertebrate and invertebrate species in every site. The biospeleological study carried out in an artificial cave in Përmet in June 2019 provided the following results: two bat species, i.e. Greater horseshoe bat Rhinolophus ferrumequinum (Schreber, 1774) and Mouse-eared bat Myotis myotis species group, probably M. blythii (Tomes, 1857) were observed, a specimen of the Balkan stream frog Rana graeca Boulenger, 1891, representing the first amphibian record for the Albanian cave, was discovered deep inside the cave, nine invertebrate species were found (7 spiders, 1 harvestman and 1 moth), with three of the spider species, i.e. Ceratinella brevis (Wider, 1834), Hogna radiata (Latreille, 1817) and Trachyzelotes barbatus (L. Koch, 1866), being the first records for Balkan caves. All recorded species, except Myotis blythii/myotis (both were previously known), are new to the study area. The total number of species recorded in this site, including the data available in literature, has increased to 29. This high number of species shows that artificial caves, including military installations, represent biodiversity hotspots, comparable to natural underground sites and demand much more attention and further investigations.
Full-text available
Subterranean organisms always attracted the attention of humans using caves with various purposes, due to the strange appearance of several among them and life in an environment considered extreme. According to a classification based on the evolutionary and ecological relationships of these organisms with subterranean habitats, first proposed by Schiner in 1854 and emended by Racovitza in 1907, three categories have been recognized: Troglobites, troglophles and trogloxenes. The Schiner-Racovitza system has been discussed, criticized, emended, the categories have been redefined, subdivided, original meanings have changed, but it is used until now. Herein we analyze in a conceptual framework the main ecological classifications of subterranean organisms, from Schiner to Trajano, in 2012, so far the last author to introduce a relevant conceptual change on the categories definitions, incorporating the source-sink population model. Conceptual inconsistencies are pointed, especially with regards to the generally ill-defined trogloxene category, and the correspondence between categories according to the original sense and in alternative classifications is discussed. Practical criteria for distinction between these categories and difficulties for their application are presented. The importance of rightly classifying subterranean populations according to the Schiner-Racovitza system for conservation of these fragile and mostly threatened habitats is discussed.
Full-text available
At least 308 species or subspecies of stygo- or troglobionts and at least 735 species or subspecies of mostly presumed stygo- or troglophiles representing 17 phyla, 38 classes, 90 orders, and 278 families of invertebrates are currently known to populate the caves and subterranean waters of Russia and other countries of the former Soviet Union. The main evolutionary burst in the endogean habitats including the MSS (milieu souterrain superficiel, or mesovoid shallow stratum) is observed in arthropods, primarily crustaceans, collembolans, and beetles. The major centers of taxonomic diversity among stygo- and troglobionts within the study region are the Caucasus (181 species, or almost 59%) and Crimea (44 species, or over 14%), which are montane karstified “glacial” refugia of the nemoral biota. The contribution of the other major regions including karstified areas is considerably smaller and gradually decreases from Central Asia (35 species; over 11%), the Far East (33; nearly 11%), the Ukrainian Carpathians with Podolia (12; almost 4%), the Russian Plain (7; over 2%), the Urals and Ural region (7; over 2%) to Siberia (5 species; 1.6%). Inventorying remains a topical problem in assessing the stygo- and troglofaunas of the territories in question. The most complete bibliography possible is included.
Full-text available
The distribution of cave-dwelling lizards of the families Gekkonidae and Phylodactylidae (Sauria) was investigated in the Zagros Mountains of Iran. Obtained information includes new distribution records of species from 15 caves. The caves are Bendireh, Taigeh, Ban, Zarrinabad, Ghadah, Kulkani, and Darham-reh in Ilam Province, Pelazh, Gavbar and Asmari in Khuzestan Province, Tadovan, Manian and Sangesh-kan in Fars Province and Dalaki and Khesht in Bushehr Province. In this study, five species belonging to the Gekkonidae and Phylodactylidae families were recorded including: Asaccus elisae, Asaccus nasrullahi, Hemidactylus persicus, Cyrtopodion scabrum, and Cyrtopodion gastrophole.
The distribution, numbers, seasonal and diurnal activities, relation to temperature, biology and morphological features of the leopard snake, Elaphe situla, in the Crimea were investigated. The snake is distributed on the southern slopes and in western and eastern parts of the northern macroslope of the Crimean mountains and on the Geraklea peninsula up to an altitude of 680 m above sea level. It occurs permanently in populated areas, including cities (Sevastopol). The distribution of the snake is sporadic, but in habitats with optimal conditions, its population reaches 2.3-8.9 ind./ha. The number of males in the population 1.7 times exceeds that of females. The snakes are active from March to November at the temperatures of 16-29°C and 16-35°C in air and substrate, respectively. They occur in winter during thaw periods. The maximal activity of these snakes is observed in breeding periods. The way of life is day and night. The night activity is maximal in August. Pairing takes place in April-May; clutch, in June-July; molt, from March to November. The diet is represented by small mammals, birds, and lizards. Sexual dimorphism manifests itself in sizes, the number of ventral and supraucaudal shields, and the proportion between sizes of the skew field and tail. A relative size of the snake's head increases with age. The share of the Situla form is maximal (54%) in the central part of the snake's range. In peripheral areas of the range, the Leopardina forms predominate (84-85%).