Asiatic Herpetological ResearchVol. 5, pp. 74-84

December 1993

Sympatric Amphibians of the Yew-box Grove, Caucasian State BiosphereReserve, Sochi, Russia
BORIS. S. TUNIYEV AND SVETLANA. Yll. BEREGOVAYA
Caucasian State Biosphere Resen>e, Sochi, Russia

Abstract. -The Yew-box Grove of the Caucasian State Biosphere Reserve is home to seven species ofThese species occur in a wide range of aquatic environments. The species composition,
physical characteristics and history of each aquatic site was evaluated. The reproductive biology and food
habits of each species was studied. These amphibians divide their niches on daily activity, seasonalamphibian.

activity, breeding site, microhabitat and food habits.

The highest amphibian diversity and species overlap

occurs in the most stable aquatic environments.

Key Words: Amphibia, Russia, Caucasus, ecology.

Introduction
It is important when studying
theinfluence of environmental factors on life
history characteristics to distinguish thosefactors that are significant and those that areof the "neutral background"part
The study of(Monchadsky, 1958).
environmental influences is accomplishedon sympatric species, usually closelyrelated species (Orr and Maple, 1978;Ananjeva, 1981) at the population level(Pianka, 1973; Schoener, 1974; Schoener,1977; Lyapkov and Severtsev, 1981). Thisis the study of the ecological niche (Pianka,et al., 1979.
In the former Soviet Union one of theareas of highest amphibian diversity isfound in the western Caucuses. The Yewbox Grove of the Caucasian State
Biosphere Reserve is inhabited by eight

amphibian species: Triturus vulgaris lantzi,T. cristatus karelini, T. vittatus ophryticus,

P elodyte scaucasicus,Bufoarborea
verrucosissimus,
Hylaschelkownikowi, Rana ridibunda, and R.
macrocnemis.

FIG. 1. Study sites of sympatric amphibians in theYew-box Grove, Caucasian State BiosphereReserve.

Methods

1-Spring 118; 2- Opolznevaya Ravine:

3- Labirintovaya Ravine; 4- Glubokaya Ravine; 5-Khosta River; 6- Samshit Pond; 7- Pond on
Field studies were conducted from 19801982 in the Yew-box Grove (approximate302 ha)

Caucasian State
Biosphere Reserve and on adjacent land.
Transect routes and study sites wereselected on the basis of local forest
typography (Gulisashvili, et al., 1975).
Observations on these study sites along thearea

inthe© 1993 by Asiatic Herpetological Research

Malaya Khosta River.
transects

were made throughout the year

The intensity of calls wasrecorded in spring (2-3 times per week),summer (1-2 times per month) and fall (2-3times per week). Over 200 individuals
(Fig.

1).

were recorded.

Asiatic Herpetological Research

December 1993

ynyiyiy

Noteixyuixn

xixVol. 5 p. 75:

at the depth of 5 cm

- river Khosta

at the depth of 1C cm- spring 118-at the depth of 15 cm- ravine Labirintovaya

Bt the depth ofc'Ocm- ravine
Glubokaya

Winter soil temperature in the Yew-Box
Grove (1982-1983).
FIG. 4.

FIG. 2. Mean monthly temperatures of perennial
bodies of water in the Yew-box Grove (1 980-1982).

Yew-box Grove

Air(Fig. 2).
thesurfaceaboveandhumidity
temperature
were recorded weekly by a thermograph(M-16AN) and a hydrograph (M-16AN)
placed in a meteorological kiosk (Fig. 3).
Soil temperatures were recorded at 5, 10,
15 and 20 cm depths (Fig. 4).
the2CWater samples were taken periodicallychemical analysis by establishedmethods (Anonymous, 1978). Ambient
meter andlight was measured with a light

10forconverted into percent relative illumination.

Climagrams were made (Formozov, 1934).
91

in

II

iy

yyi

yn

yn

ix

xxi

xn

95

Species composition of each biotope wascalculated using conventional methods(Kashkarov,1927;1952).

ioo
humidity

habitsandwereKaletskaya,
studied using non-lethal methods(Verzhutsky and Zhuravlev, 1977).

Results

FIG. 3. Climatram of the Yew-box Grove (1982).

The location, weather conditions, air andbody temperature, and behavior of eachspecimen was recorded. During 1982,
detailed microclimate records were made inDinesman
Food

Description of Study Sites

Spring 118. This site
contains a small perennial stream running

Study Site

1.

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TABLE 1.

Asiatic Herpetological ResearchSome hydrochemical indices of the study sites.

Sample site
No. of samples

December 1993

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Asiatic Herpetological ResearchVol. 5 p. 77

Distribution of sympatric amphibians at Spring 118 in the Yew Box Grove. 1- Triturus vittatus
ophryticus; 2- Pelodytes caucasicus; 3- Rana ridibunda; 4- Hyla arborea schelkownikowi; 5- Rana
macrocnemis; 6- Bufo verrucosissimus.

FIG. 5.

Spring temperature ranges from 11-16° C.

The water chemistry of the spring water
was hydrocarbonic-calcic with moderatemineralization, and moderately hard (2.42 3.6 mg-eq/1). Ammonium concentration islow (maximum = 0.04 mg/1) and in winter
drops

to 0.

Nitrates are highest during

varies fromto 0.07 mg/1, but during
floods it can reach 2.32 mg/1. This stream
issubject

toConcentrationoccasionalofnitratesflooding.
reaches amaximum during floods of 1.44 mg/1.
Dissolved

oxygenis10-15mg/l

andcarbonic acid is 10mg/l.

summer low water period (up to 0.37mg/1).

Phosphorus concentration is verylow (>0.01 mg/1). Bufo verrucosissimus,
macrocnemis, P. caucasicus, R.

R.

ridibunda and T.

v.

ophryticus are foundliving at this study site and the last three
species breed here.

Study Site 5. Khosta River. This site isa small mountain stream 21 km long thatdrains a watershed of approximately 96

km 2 The stream flows through a canyon.

formedCretaceous limestone at anaverage of 5 m-Vsec. Water flow is derived
from runoff and springs arising in karst
rock formations. Vegetation is a broadleafed, subtropical Colchis type forest.
Relative illumination is as high as 100%.
The water chemistry of this stream ishydrocarbonic-calcic with moderatemineralization and basic pH (7.7-8.5).
Ammonium concentration is not high andin

The Nizhe-Khostinsky Spring, flowing
out of a karst formation, contributes 1-1.5m-Vsec of flow at 11-13° C to the Khosta
In low water periods the KhostaRiver.
River above the spring nearly dries up andits temperature ranges from 0.6 (the riverfreezes) 26° C. During low waters on theKhosta River the Nizhe-Khostinsky Springprovides a relatively stable flow andtemperature regime. This spring serves asa barrier for dispersal of amphibians at thisstudy site. Rana ridibunda and Hylaarborea schelkownikowi live and reproduce
above the spring. Pelodytes caucasicus andBufo verrucoisissimus live and breed

below the spring.

isStudy Site 6. Samshit Pond. This sitea small pond with flowing water in astand of hornbeam {Carpinus betulus) inthebroad-leafed forest.
RelativeVol. 5 p. 78

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Ranamacrocnemis

I—Uo.

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December 1993

TABLE 4. Comparison of development periods of Triturus vittatus ophryticus and Pelodytes caucasicus.
MONTHS:
SPECIES:December 1993

Asiatic Herpetological Research

TABLE 5. Daily activity of amphibians of the Yew-box Grove.

Vol. 5 p. 81

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TABLE 6. Faeces compostion of sympatric amphibian species of the Yew-box Grove
SPECIES:

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TABLE 7. Size limits of feeding objects of sypatric amphibians in the Yew-box Grove
SIZE OF FEEDING OBJECTS ( in mm)

SPECIES:Vol. 5 p. 83

Asiatic Herpetological ResearchVol. 5 p. 84

terrestrial invertebrates (all other species).

Rana ridibunda feeds on both aquatic andterrestrial invertebrates (Table 6, 7).

December 1993

FORMOZOV, A. N.

[Special type of1934.

for ecological studies].

climagrams

ScientificTransactions of Moscow University 2:271-274.
(In Russian).

In a given area species diversity
dependent upon niche separation (Pianka,The seven amphibian species
1981).
studied here are characterized by biotopic

GULISASHVILI, V. Z., L. B. MAHATADZE, AND
1975.
L. I. PRILIPKO.
[Vegetation of theNauka
Caucasus].
Publishing House, Moscow.

(including breeding site choice, hibernationsites and summer activity ranges), seasonal

KASHKAROV, D. N.

isactivity period, daily activity period andfood habits isolation. In the Yew-box
Grove the highest level of species diversity
is achieved in the most stable aquaticenvironments. Water temperature during

the breeding season, level of illumination

and water chemistry appeartobe an

227 pp.

(In Russian).

[MethodsofTashkent. 23 pp. (In Russian).

LYAPKOV, S. M. AND A. S. SEVERTSEV.

1981.

[Mechanism of co-existence of two species ofFar-Eastern Anura]. Zoological Journal 3:398408. (In Russian).

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Acknowledgments

We wish to express our gratitude to N.
B. Ananjeva for valuable consultations andto I. V. Marchukaitis for preparing theillustrations.

ORR, L. P. AND W.
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PIANKA, E. R.
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