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Amphibians and Reptiles in the Subterranean Cavities of the Crimean Mountains

February 2019
Russian Journal of Herpetology 26(1):29-53

DOI: 10.30906/1026-2296-2019-26-1-29-53

Authors:

Ilya Turbanov

Russian Academy of Sciences

Oleg V. Kukushkin

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

Robert S. Vargovitsh

National Academy of Sciences of Ukraine

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.

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Content may be subject to copyright.

DOI: 10.30906/1026-2296-2019-26-1-29-53

AMPHIBIANS AND REPTILES IN THE SUBTERRANEAN CAVITIES

OF THE CRIMEAN MOUNTAINS

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.

INTRODUCTION

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,

2009).

Relatively old age (mostly Middle or Late Pleisto-

cene), the presence of multiple underground cavities and

a high diversity of the conditions determine altogether

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;

e-mail: turba13@mail.ru

2Cherepovets State University, Cherepovets, Vologda oblast’, Russia.

3T. I. Vyazemski Karadag Scientific Station — Nature Reserve, Cri-

mea; e-mail: Mtasketi2018@gmail.com

4Zoological Institute of Russian Academy of Sciences, St. Petersburg,

Russia.

5I. I. Schmalhausen Institute of Zoology of Academy of Science of

Ukraine, Ukraine; e-mail: arete@izan.kiev.ua

* 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

2018.

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 “...in 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,

2006).

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,

2010).

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

open.

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.

MATERIAL AND METHODS

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

Species

Karst area,

administrative

affiliation of the

point of record

Mountain

massif,

element

of relief

The name

of the cavity

(its cadastral

number)

Altitude,

m a.s.l.

Date of

observation

Depth of

the record

from the

surface, m

Distance from

entrance, m

(for the caves with

horizontal stretch)

Number

of individuals

Source of

information

T. karelinii AY, Yalta btw. Mt. Byuzyuka

and Mt. Balchik-

Kaya

Razocharovanie

(= Uparennaya)

(4425/3353 – 3)

940 2nd half

of May 1999 5 — 2 A. Yushko, O. Peretyatko

(personal

communication)

BB, Sevastopol Canyon of Chernaya

River, Mt. Khlama Grot Anny Cave

(4431/3345-1)

354 31.05.2003 — At entrance ca. 10 (ad., juv.) Matyushkin (2010)

AY, Sevastopol locality

Karadagsky Les Kristalnaya (= Maksimovich)

(4426/3354-2)

834 2003 – 2004 Up to 64 — Several adults

and larvas Kotenko,

Kukushkin (2010)

AY, Sevastopol Ai-Petrinskaya

Yaila Gibel’ Yaponii

(4427/3355-24)

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

funnel

— 2 juv. Matyushkin (2010)

AY, Sevastopol Ai-Petrinskaya Yaila Raskop Medvezhyi

(4428/3354-6)

871 18.01.2008 8 — 1 Matyushkin (2010)

BB, Sevastopol Bayidarskaya Hollow Mamut-Chokrak

(4425/3345-2)

280 08.10.2016 3 – 4 — 1 S. Kebets, A. Akimov

(personal

communication)

AY, Sevastopol Mt. Kurt-Kaya Entuziastov

(4427/3351-1)

296 20.11.2016 27 — 3 Ye. Yanovskyi (personal

communication)

AY, Sevastopol Ai-Petrinskaya Yaila Gelektitovaya-Vesennyaya

(4428/3356-6)

1075 02.04.2017 40 — 2 A. Akimov, S. Kebets

(personal

communication)

BB, Sevastopol Canyon of Chernaya

River Chernorechenskaya

(4431/3342-2)

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

communication)

December

2011 1 Pysanets, Kukushkin

(2016)

KY, Belogorsk Karabi Yaila Monastyr-Chokrak

(= Su-Teshigy)

(4452/3432-2)

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

communication)

BB, Sevastopol Bayidarskaya Hollow Mamut-Chokrak

(4425/3345-2)

280 19.08.2010 — 30 1 I. Turbanov

Cave Herpetofauna of the Crimea 33

Species

Karst area,

administrative

affiliation of the

point of record

Mountain

massif,

element

of relief

The name

of the cavity

(its cadastral

number)

Altitude,

m a.s.l.

Date of

observation

Depth of

the record

from the

surface, m

Distance from

entrance, m

(for the caves with

horizontal stretch)

Number

of individuals

Source of

information

AY, Yalta Ai-Petrinskaya Yaila Oreshek (4427/3359-3) 1070 23.10.2010 24 — 1 A. Papyi (personal

communication)

BB, Sevastopol Bayidarskaya Hollow,

Mt. Biyuk-Sinor Sakhtykh (4425/3348-1) 639 11.08.2012 7 — 1 S. Bogdanenko (personal

communication)

B. viridis SY, Simferopol Dolgorukovskaya

Yaila Kizil-Koba (= Krasnaya)

(4452/3420-7)

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

communication)

BB, Sevastopol Ravine

Kobalar-Dere Karan’-Koba (= Temnaya)

(4430/3348-1)

339 18.11.2006 — 16 1 Matyushkin (2010)

KY, Belogorsk Karabi Yaila,

Ridge Bagalma Kara-Murza

(4454/3433-5)

625 22.08.2009 130 — 1 P. Oksinenko (personal

communication)

BB, Sevastopol Ridge Spilia,

Mt. Asketi Asketi-2 (not

included in the Cadastre) ca. 320 Mid-Aug.

2014 — 5 – 7 1 P. Oksinenko (personal

communication)

26.09.2015 5 – 7 1 O. Kukushkin,

P. Oksinenko

BY, Alushta Babugan Yaila,

area of the Mt.

Zeytin-Kosh

Unnamed cave (not

included in the Cadastre) 1460 Nov. 2014 — 3 – 4 1 Yu. Balakhtinova,

S. Klushin (personal

communication)

AY, Sevastopol locality

Karadagsky Les Bizyukskaya (not

included in the Cadastre) 756 01.04.2017 106 — 1 R. Potapov (personal

communication)

KY, Belogorsk Karabi Yaila,

Egiz-Tinakh Valley Yabushkan-Koba

(4452/3434-1)

967 11.08.2017 — 10 1 O. Kukushkin, I.

Turbanov

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.

Oksinenko

KY, Alushta Karabi Yaila Tserkov’ (4451/3436-1) ca. 985 02.12.2017 — 10 1 Yu. Balakhtinova, S.

Klushin

P. (ridi-

bun-dus)

complex

AY, Yalta Ai-Petrinskaya Yaila,

Mt. Morcheka Druzhba

(4425/3354-10)

969 01.01.2007 45 — 1 juv. A. Shendrik (personal

communication)

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

communication)

04.03.2018 15 50 3 juv. I. Turbanov. A.

Aschepkova

AY, Sevastopol Mt. Villya-Burun Lyagushach’ya

4428/3356-3)

905 23.02.2010 23 — 1 S. Kebets (personal

communication)

KY, Belogorsk NE spures of Karabi

Yaila, area of the

Mt. Gol’-Kaya

Karasu-Bashy

(4458/3434-1)

390 30.08.2015 8 — 1 A. Akimov, Yu.

Balakhtinova (personal

communication)

34 Ilya S. Turbanov et al.

TABLE 2. Records of Reptiles in Caves of Mountainous Crimea

Species

Karst area,

administrative

affiliation of the

point of record

Mountain massif,

element of relief

The name

of the cavity

(its cadastral

number)

Altitude,

m a.s.l.

Date of

observation

Depth of the

record

from the

surface, m

Distance from

entrance, m

(for the caves

with horizontal

stretch)

Number

of indivi-

duals

Source

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

communication)

BB, Sevastopol Cape Aiya,

Mt. Kalafatlar Gekkonovaya

(4427/3338-2)

496 08.09.2012 — 5 – 12 3 Yu. Balakhtinova

(personal

communication)

28.12.2012 3 – 20 ca. 18 I. Turbanov, Yu.

Balakhtinova

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

Dvukhkupolnaya

(4428/3338-1)

403 05.12 and

15.12.2012 — from7–10

to 15 – 25 ca. 10 and

up to 20 I. Turbanov, Yu.

Balakhtinova

385 26.09.2015 3 1 O. Kukushkin, P.

Oksinenko

BB, Sevastopol Ridge Spilia,

Mt. Asketi Asketi-1 and Asketi-2

(not included in the

Cadastre)

ca. 320 Mid-August

2014 — 5 – 12 ca. 16 P. Oksinenko, Ye.

Oksinenko (personal

communication)

Asketi-1 26.09.2015 5 – 7 1 O. Kukushkin, P.

Oksinenko

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

(4428/3354-2)

860 14.05.1994 10 10 1 female

and its egg Matyushkin (2010)

C. austriaca AY, Yalta maritime cliff

of the Mt. Morcheka Medovaya

(4425/3354-9)

910 19.06.2006 — 30 1 juv. A. Shendrik (personal

communication)

CY, Simferopol Chatyrdag,

Lower plateau Vyalov-Azimutnaya

Togerik-Alan-Khosar)

(4445/3417 – 6)

1020 07.08.2006 30 — 1 (dead

specimen) E. Bednarskaya

(personal

communication)

KY, Belogorsk Karabi Yaila Skazka (4452/3433-7) 1002 Early July 2009 45 — 1 I. Turbanov

KY, Belogorsk Karabi Yaila Monastyr-Chokrak

(= Su-Teshigy)

(4452/3432-2)

998 18.07.2010 95 — 1 À. Akimov (personal

communication)

KY, Belogorsk Karabi Yaila Kastere

(4452/3432-7)

1011 24 – 27.07.2010 70 — 1 I. Turbanov

Cave Herpetofauna of the Crimea 35

Species

Karst area,

administrative

affiliation of the

point of record

Mountain massif,

element of relief

The name

of the cavity

(its cadastral

number)

Altitude,

m a.s.l.

Date of

observation

Depth of the

record

from the

surface, m

Distance from

entrance, m

(for the caves

with horizontal

stretch)

Number

of indivi-

duals

Source

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

(personal

communication)

BB, Sevastopol Cape Aiya,

Mt. Kalafatlar Gekkonovaya 496 20.03.2016 — 10 1 O. Kukushkin, P.

Oksinenko

BB, Sevastopol Bayidarskaya Hollow Urkusta-Chokrak-Koba

(4431/3349-2)

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

(personal

communication)

BB, Sevastopol Cape Aiya,

Mt. Gurush Unnamed grotto (not

included in the Cadastre) ca. 400 02.12.2012 — 2 1 Yu. Balakhtinova

(personal

communication)

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.

Oksinenko

BB, Sevastopol Cape Aiya,

Mt. Gurush Dvukhkupolnaya 385 26.09.2015 At entrance 1 (moulted

skin) O. Kukushkin, P.

Oksinenko

BB, Sevastopol Cape Aiya, Mt.

Kalafatlar Gnomov (not included

in the Cadastre) ca. 490 31.03.2016 — 3 1 O. Kukushkin, E.

Gladilina

BS, Bakhchisarayi Mt. Baba-Dag Mangup-Kale-14 Cave

(4435/3348-2)

ca. 550 03.05.2018 10 14 1 I. Turbanov

E. sauromates BB, Sevastopol Bayidarskaya Hollow Mamut-Chokrak

(4425/3345-2)

280 13.05.2013 — At entrance 1 L. Bogdanenko

(personal

communication)

V. renardi CY, Simferopol the head of

Orlinoe gorge Monashjya (not included

in the Cadastre) 1025 17.05.2009 — At entrance 1 E. Bednarskaya

(personal

communication)

Ophidia indet. CY, Simferopol Chatyrdag,

Lower plateau Viking (not included in

the Cadastre) ca. 950 05.04.2006 ca. 50 — 1 E. Bednarskaya

(personal

communication)

CY, Simferopol Chatyrdag,

Lower plateau Golubinaya-2

(4447/3417-3)

ca. 990 28.07.2007 15 — 1 E. Bednarskaya

(personal

communication)

CY, Simferopol Chatyrdag,

Lower plateau Tryokhetazhnaya

(4447/3417-7)

1009 12.11.2007 10 — 1 E. Bednarskaya

(personal

communication)

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 (https://speleoatlas.ru) 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.

RESULTS

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.

Amphibia

Four amphibian species from four families were re-

corded in the cave habitats (Fig. 1A).

Order Caudata

Salamandridae

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).

REPTILIA

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

Pass.

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).

DISCUSSION

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.,

2017).

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

revealed.

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

slopes.

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.

CONCLUSIONS

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).

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Cave Herpetofauna of the Crimea 53

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Ilya Turbanov · Oleg V. Kukushkin · Robert S. Vargovitsh

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addendum_Table 2_true version.doc

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Ilya Turbanov · Oleg V. Kukushkin · Robert S. Vargovitsh

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Fig. 4 b_true version.tif

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Ilya Turbanov · Oleg V. Kukushkin · Robert S. Vargovitsh

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Ilya Turbanov · Oleg V. Kukushkin · Robert S. Vargovitsh

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TABLE 2. Records of reptiles in caves of Mountainous Crimea

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Ilya Turbanov · Oleg V. Kukushkin · Robert S. Vargovitsh

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Table 2_true version.doc

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Ilya Turbanov · Oleg V. Kukushkin · Robert S. Vargovitsh

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Amphibians and reptiles in the caves of the Greater Caucasus

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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.

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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.

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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.

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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.

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Katarina Koller Šarić

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Apr 2004
ZOOL ZH

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%).

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