Asiatic Herpetological Research

December 1993

Vol. 5, pp. l-To]

iU

Coluber atayevi Sp. Nov. (Ophidia, Colubridae) from the Kopet-Dag
Mountains of Turkmenistan
BORIS S. TUNIYEV AND SAHAT M. SHAMMAKOV 2
1

Caucasian Slate Biosphere Reserve, Sochi, Russia^Institute of Zoology, Turkmenistan Academy of Sciences, Azadi Street 6,744000 Ashgabat,

Turkmenistan

Abstract. -An examination snakes, formerly considered to be Coluber najadum, from the Kopet-Dag
Mountains of Turkmenistan, leads us to believe that this population represents an undescribed species. We
here describe this population as Coluber atayevi.

Key words:

Reptilia, Ophidia, Colubridae, Coluber, Turkmenistan, systematics.

8) form ofmandibular scutes; 9) shape of the head;10) distribution of scutes on throat; 11)
L/L. cd.= body length/tail length; 12) Pr.

number and size of sublabials;

Introduction

During field work in the Kopet-Dag
Mountains, Turkmenistan, we observed
and analyzed many individuals of Coluber
najadum under natural conditions.

Comparison of these individuals with
Caucasian material led to the conclusionthat the Kopet-Dag snakes belong to aThispreviously unrecognized species.
conclusion is supported by significant
morphological divergence from the otherrepresentatives of the najadum-rubriceps

oc.= number of preorbital scales; 13) Post.
oc.= number of postorbital scales;
14)
numberofscales;15)
temporal
Temp.=
A.= form of anal scutes.

For numerical features and indices, wehave calculated the mean (x), mean error
2(m), mean square deviation (S ), using theformulas for small samples (Lakin, 1980)

complex.

When

Methods

We analyzed 10 specimens of Coluber
najadum (Eichwald) from various areas ofdescribing the biotopes, we
determined the plant species according to
Nitikin and Geldykhanov (1988); thegeneral vegetation type follows Korovin(1934), with some corrections.

the Caucasian Isthmus and 10 specimensfrom the Kopet-Dag belonging to the newtaxon. Morphometric data were comparedwith the available information in theliterature about Coluber najadum in its
natural habitat throughout its range(Terentjev and Chernov, 1949; Bannikov etal.,1977) and for separate regions(Ananjeva

andOrlov,

1977;

History of the Study of ''Coluber

najadum'" in the Kopet-Dag

Zamenisdahli Fitzinger was

firstmentioned from the environs of Sulukli
Spring and the Kuchan road by Varentzov(1894). Nikolskij (1905, 1916) observed
this species in the vicinity of Ashkhabad.

Muzskheleshvili, 1970).

On the basis of these records, Coluber

The following features and indices havebeen used: 1) L. = length of body, mm; 2)
L. cd. = length of tail, mm;3) Sq.=
number of scales around body; 4) Ventr.=
number of ventral scutes; 5) S. cd.=
number of subcaudal scutes; 6) Lab.=
number of upper labials; 7) Sublab.=

najadum (Eichwald) was included in the listof the Turkmenistan herpetofauna(Chernov, 1934; Terentjev and Chernov,
All previous
1949; Bogdanov, 1962).
collections from the territory of
Turkmenistan and the neighboring parts ofIran have been analyzed by Ananjeva andOrlov (1977). They added the localities of© 1993 by Asiatic Herpetological ResearchVol. 5, p. 2

TABLE 1.

Asiatic Herpetological Research

December 1993

Morphometry of Coluber atayevi paratypes in the collection of the CaucasianReserve, Sochi, Russia (see text for abbreviations).

No.
421

L.

L.cd. Sq. L./L.cd Ventr. S.cd.

Lab. Preoc.

Postoc.

Temp.

Sublab.

Tuniyev and Shammakow
Asiatic Herpetological Research

Adult Coluber atayevi.

Type locality of Coiluber atayevi, environs of Saivan Village, Saivan-Nokhur Plateau,western Kopet-Dag, Bakharden Region, Turkmenistan.

December 1993

FIG.

1 .

Asiatic Herpetological ResearchVol. 5, p. 3

Holotype of Coluber atayevi sp. nov. (Collection of the Caucasian Reserve no. 420).

ofSnoutholotype:tail 172 mm; headlength 19.6 mm, head width 8.4 mm, head

Description

vent length 500 mm,

height 6.2 mm. Head smoothly rounded,narrow, covered with large regular scutes;
8 upper labials, 5th upper labial touches

lower postorbital and large lower temporalwith its extended upper posterior side; 9lower labials, 6th largest, last two pairs
almost covered by upper labials; a single
preorbitalon either side, 2 small scutes

Colorationinpreservative:
Dorsum gray, venter grayish; 5 large dark
ocelli bordered with light circles on sides ofneck; lateral row of small black dots ending
abruptly on anterior third of body. Eyewith white lines extending
anteriorly and posteriorly; lower white
stripe extends over upper labials; a narrow
black streak running posteriorly and downfrom eye, situated on 5th upper labial andoutlinedslightly touching 6th.

below; posterior pair of upper temporals
slightly larger than anterior one; seen fromabove, rostral extends slightly between
internasals.
Narrow genial scutes incontact along mental groove, no space

CountsDescription of paratypes:
and measurements of the paratypes in thecollection of the Caucasian Reserve, Sochi,
Russia are given in Table 1.

between posterior genials.
Diagnosis:
Nineteen scale rows at midbody; 206
abdominal scutes; 97 pairs of subcaudals;
anal divided. Scales bordering abdominal
scutes of same size as other lateral scales;
body scales smooth, rhombic; ridge on
lateral aspect of abdominal scutes indistinct,
almost absent.

Comparatively

small

snake (Fig. 2), smaller than C. n. najadum,
C. n. dahli, and C. r. rubriceps indimensions. It is comparable in size with
the European subspecies, C. rubriceps
In contrast to C. najadum, whose
constitutes 1/3 of its total length, C.
atayevi has a comparatively short tail,thracius.
tailVol. 5, p. 4

Asiatic Herpetological Research

December 1993'-•iSfe,-"FIG. 2. Representatives of the najadum-rubriceps" -complex. Left- Largest specimen of Coluber atayevi
sp. nov. (Collection of the Caucasian Reserve no. 421).
Right- Medium-sized specimen of Coluber
Reserve no. 94).
CollectionoftheCaucasianenvirons,
Akhun,(SochinajadumMalyapproximately 1/4 of total length,characteristic of C. rubriceps.

also

Habitus and elements of coloration of C.
atayevi are intermediate between C
najadum and C. rubriceps; its narrow,
sharp, and flat head with the rostrum
beveled downward is closer to the C.
rubriceps head shape than to that of C.
najadum, with its comparatively wide,
rounded, and high head, where the upperand lower surfaces of head are parallel.
.

Color pattern of C. atayevi resembles thatof C. najadum, but brown colors prevail
instead of olive-green ones.
Lateralabdominal ridges are practically absent in
C. atayevi, in contrast to the abovementioned species, and consequently the
body is round in cross-section and notrectangular as in C. najadum and C.

rubriceps.

Genial scutes of C. atayevi contact one
another along the mental groove, rarely
having a few isolated granules between theposterior pair, whereas between the widely
separated posterior genials of C. najadum
there are always 2-4 rows of well-developed scales (Fig. 3). Posterior upperlabials of C.
atayevi are weakly

December 1993

Asiatic Herpetological Research

900-1400 meters elevation.

Vol. 5, p. 5

The most

typical biotopes of the species are ecotonesof mesophilous derivatives of deciduous
forest and meadow-steppe coenosis

the edges of small ravines having

along

depositsof limestone and argillaceous slates.
Indicators of the deciduous forest are
isolated old trees of Oriental plane (Platanus
orientalis) and English walnut (Juglans
regia). Forest plots of "prashiblyak" aretypified by Aceretum fruticans and Acer
turcomanicum, a subdominant role playedby Crataegus turcomanica, Lonicera
floribunda, Prunus cerasifera, Cotoneasternummularioides, Cotoneaster ovatus, andRubus anatolicus. In the herb layer, there
are such species as Alliaria petiolata,Lamium album, Geranium pussilum, Arumjuquemontii, and Allium paradoxum.

FIG.

Rocky-shrubby vegetation is usual forthe ecotone of forest ravines on the rocky3.

Distribution

of

head

scalesonrepresentatives of the Coluber najadum-Coluber
rubriceps-complex: a, b, c- a young specimen of C.
najadum; d, e, f- a young specimen of C. atayevi
sp. nov.

distinguished from the throat scales,
whereas all upper labials of C. najadum arestrongly pronounced.

Geographic distribution: The rangeof Coluber atayevi includes the western andcentral Kopet-Dag, from thesurroundingsof the Kara-Kala settlement in the west to
the Sulukli Spring in the east.

This is ansuchvegetation types as "prashiblyak," "broad-upland species, associated withleavedforest"(Kamelin,

1970)

and

"phrygana," and in the western part of theuplands, where these plant associations
occur at lower elevations, individuals of C.
atayevi are found at elevations of 400-1600

meters (Shcherbak et al., 1991). At theeastern end of its range (Dushak
Mountain), the snake has been found at
2000 meters elevation (Shcherbak et al.,1986).

Biotopes:

Accordingtoourobservations, Coluber atayevi is found on
the highest parts of the Saivano-Nokhur
Plateau and on the crests of mountains atand scree slopes, with isolated trees of
Celtis caucasica and many different shrubs
and semi-shrubs:
Colutea gracillis,
Ephedra equisitina, Thelycrania meyeri,
Rhamnus coriacea, Hymenocrater
bituminosus, and Artemisia turcomanica.
Representatives of Coluber atayevi arefound in Cousinia smirnovii associations
and Astragalus piletoclados groups(phrygana type) in the immediate vicinity of
the forest and shrubby communities withsteppe wedges, mainly in overgrazed
places. The region is comparatively wellwatered, because almost every ravine has aspring or stream.

The vegetation of the eastern border ofthe distribution of C. atayevi is describedby Korovin (1934): "Dushak mountain is
an isolated massif, formed by light
limestone. Its steep slopes serve as a home
for typical mountainous xerophytes. Here

we find juniper, both isolated trees andgroups of them. A number of semishrubs
and xerophytic herbs form flora of thesemountains." Later, the author mentions thedomination of shrubs of Astragalus
piletoclados, as well as groups of different
species of Acantholimon and here very
common gray cushion-like groups ofOnobrychis comma. Korovin concludesAsiatic Herpetological ResearchVol. 5, p. 6that the cushion-like xerophytes of theKopet-Dag are better developed on the topsofmountains(about2000

elevation).
zone, so phrygana belongs to high-altitude
vegetation.

To our regret, absence of data about thebiotopes of species from the other places
gives us no opportunity to characterize the
Cenozoic ties of the species throughout the

Population density: Coluber atayevi
is the most numerous snake species on the
Saivano-Nokhur Plateau. Six specimens
were found during a three-hour excursion
in the vicinity of Saivan village in May,
1990; 12 specimens were noticed duringThenumber of snakes (as many as 5
specimens per 300 meters) was among thethe same period of time in May, 1991.

largestshrubs of rocky-scree plots of ravines.
Isolated specimens of snakes were met instoneless places. The fact that this species
is rather common for the western Kopet-

Dag is proved by the data of Ataev et al.
All other authors mention onlyisolated findings, reckoning

speciesinsunnyweather.
Diet:

Lizards in the habitat of Coluber

atayevi are Ablepharus pannonicus, Stellio

caucasicus, and Pseudopus apodus.
Cyrtopodion caspius and Eremias strauchi
are common, though not so numerous,
while Mabuya aurata and Eumecestaeniolatus are rare. Considering the small
size of the head and body of Coluber
atayevi, the bulk of its diet must be formed

whole area.

rarer

where they are usually met

metersThis is higher than the steppe(1991).

December 1993

of

the

among theKopet-Dag.
it

Apparently, the sporadic distribution of thespecies and the considerable altitudes atwhich its habitat occurs, are the reasons
why it is rarely met. It is not excluded thatits population density in the eastern part ofits range is significantly lower than in its
western part.

Seasonal and daily activity:
Presumably, the species' activity begins inthemiddle of April, considering
temperature conditions of this mountainzone (Babaev, et al., 1982) and preferable
temperatures of daily activity, noted inMay, 1990-1991. Coluber atayevi is adiurnal species with two-peak activity in
May; the morning peak (9:00-11:00) is
strongly pronounced and the evening one isfeebly marked. The snakes are active insunny, windless weather. We have neverseen them when it rains, there are strong
winds, or heavy overcast. In bad weather
the snakes are absent not only on thesurface, but from under the plates of slate,caucasicus, young
only of Stellio
specimens of Eremias strauchi, andpossibly Cyrtopodion caspius. Evidently,
C. atayevi is a saurophage exclusively,
since the common and abundant Microtus
socialis and Saxetania cultricolis (amicromammal and an orthopteran insect,respectively) are too large to be eaten by
this snake.

Syntopical species of snakes:
Subdominants of Coluber atayevi are C.
nummifer, C. ravergieri and Vipera
lebetina; Typhlops vermicularis is common;

Agkistrodon halys caucasicus and Natrix
tesselata are rare. At the borders of theEirenismeda,
Psammophis lineolatum, Naja oxiana, andEryx miliaris are met. These species are
more characteristic for smaller hypsometricspecies'biotopesmarks.

Discussion
Coluber atayevi is most closely related to
C. najadum and C. rubriceps, possessing
features of both species; in habitus (headshape, in particular) it resembles C.

rubriceps, but it is similar to C. najadum inshould note that thesecolor pattern.

Wefeatures are characteristic for juvenile
specimens as well as adults; in otherwords, we cannot distinguish ancestralfeatures that would allow us to unite C.
atayevi with either of the species mentioned(Table 2, Fig. 3).

Judging

from

distributionof thethe contemporary
three species, weof
the complex is thethatthecenter
proposeEastern Mediterranean region, so-called dry
land of Asia Minor or Balkan-CaucasianDecember 1993

Asiatic Herpetological ResearchVol. 5, p. 7

Vol. 5, p. 8

Asiatic Herpetological Research

The ancestral form could havebeen widespread in the Upper Miocene-

dry land.

Pliocene over subtropical semiarid regionsof Asia Minor united with the Central Asian
Massif, Caucasian Island and the Balkan
Mountains in the Miocene. Mixed savannahylile landscape, combining alternation ofopen spaces with subtropical forests werecharacteristic for the northern parts of theFossilIranian Plateau in that period.
mammals and birds testify to this fact
In Vereshchagin's(Vereshchagin, 1959).
opinion, similarity of the Upper Miocene
fauna unites Central Asia, the Caucasus,
the Crimea, and the Balkan Mountains.

Analysis of fossil tortoises testifies to thepresence of a bridge between the BalkanCaucasian dry-land and an IranianPakistanian Peninsula (Chkhikvadze,1991).

December 1993

Caspian seas, could have favored thesecondary overlapping of the diverged C.
najadum and C. rubriceps. This sympatry,which can be seen today in some areas,
served to reinforce the species divergence.
In the east, representatives of this complex
could survive only in relict populations on
the northern (Caspian) slope of the Alborz
Range and, in isolated form, in thederivatives of forest coenosis in the western

Kopet-Dag that were never subjected toglaciation (Gvozdetzkij and Mikhailov,
1987). In the Iranian part of the TurkmenKhorasan mountains there are islands ofgyrkana forests to the east of Astrabad,reaching the longitude of GombedezhKabus. Some islands of oak forests reach
Bodgenurd in Khorasan (Menitskij, 1984).
Anderson (1968) wrote about possible
preservation of lizard refuges since the
Pleistocene in the Kopet-Dag mountains.

Alpineorthogenesis that has led to theformation of mountain relief over the wholearea of study, from the Balkan Mountains
to the Kopet-Dag, evidently promoted theIncreasing climatic aridization in the

Holocene can be the cause of disappearanceof C. atayevi in the foothills and severe

breakup of the original area of the ancestralform of the najadum-rubriceps complex. A

restriction of its area in the middle-altitude

number of populations along the sea coasts

centralof that time (Mediterranean, Black, andCaspian Seas) could already have been

breakup of the area into several local
refuges. The "primitive" morphologicalfeatures were preserved in the absence ofisolated in the Pleistocene.

The event has

and high-altitude parts of the western and

Kopet-Dag and, possibly, to thebeen connected with aridization of thecontinental interior and the formation ofcontacts with closely related forms. In any

newvariability inconditions.

These

ecologicalconditions have given impetus to thedevelopment of the highland xerophytic
vegetation of Iran and the formation of thefauna of arid mountains. It should be notedthat the vegetation of the central andwestern Kopet-Dag has more features incommon with the vegetation of southern
and western territories than with the othermountain massifs of Central Asia. Korovin(1934) emphasizes that the Kopet-Dag
are the connecting link with
Mediterranean macchias. The provenanceof highland xerophytes (phrygana), as well
as forest vegetation, Korovin stated to be
Persia and Armenia.
forests

Pleistocene cataclysms, connected withthe glaciation in the Caucasus, Asia Minor,

and the Iranian Plateau, as well as thechanges in the basins of the Black andcase,further

only

study

of

species

specific localities of the
can throw light on thisKopet-Dagquestion.

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