Asiatic Herpetological Research

April 1990

Vol. 3, pp. 1-36

Three Species in the Vipera kaznakowi Complex (Eurosiberian Group)
Their Present Distribution, Possible Genesis, andthe Caucasus:
Phylogeny*

in

NIKOLAI L. ORLOV AND BORIS S. TUNIYEV 2
1^Zoological Institute. USSR Academy of Sciences, Leningrad, USSR
2
Causasian State Biosphere Reserve, Sochi, USSR

Abstract.-Thiee valid species: Vipera ka2nakowi Nikolsky, V. dinniki Nikolsky and V. darevskii"
Vedmedeja, Orlov, and Tuniyev of the "Vipera kaznakowi complex belonging to the Eurosiberian viper
group are recognized. The distributions of the closely related species, V. kaznakowi and V. dinniki aredefined. The habitat of V. kaznakowi is present along the Black Sea coast, from Khopa Village (Turkey)
in the south, to Maikop (USSR) in the north. Vipera kaznakowi is associated with montane areas fromsea level up to 1000 m. The range of V. dinniki covers the northern and southern slopes of the Great(Main) Caucasus Ridge, ranging from the Fisht-Oshten Massive in the west to Shkhara Mountain in theeast. Vipera dinniki tends to be restricted to alpine and subalpine zones at elevations from 1 500 m to3000 m. Vipera darevskii occurs in the southeastern part of the Dzhavachet Mountain Ridge, Armenia,"
near the border with Georgia. The history of studies on the vipers of the "Vipera kaznakowi complex issummarized. The possible genesis of the present distributions of these vipers and their phylogenetic
relationships are discussed. Comparative morphological and ecological characters of the three species arelisted.

Key Words:

Reptilia, Serpentes, Viperidae, Vipera, USSR, Caucasus, systematics, ecology.

Introduction

The study of the following viper speciesof the Caucasus, Vipera kaznakowi
Nikolsky 1909, V. dinniki(Nikolsky1913), and V. darevskii
Vedmederja,
Orlov and Tuniyev 1986 and the history of
their distributions is of great interest forunderstanding the phylogenetic links of theEurosiberian shield-headed vipers from the
Caucasus (subgenus Pelias, Merrem 1808)and the formation of the present snake
fauna in the west Caucasus Isthmus.
Ever since Nikolsky (1909) describedcomplex. All of them occur
primarily in the western part of the
Caucasus Isthmus. The eastern boundaryof the species range needs to be defined.

kaznakowi

The Caucasus vipers (Fig. 1) have beenrecorded from the following localities:

Western Dagestan in the vicinity of
Khasav-Yurt (Krasovsky 1929, 1932).

Along the slope of Makh-khokh
Mountain from the highlands of Ingushetia(Chernov 1929).

On Legli Mountain in the Gukasyansky

the Caucasus Viper, Vipera kaznakowi, theregion, Armenia (Darevsky 1956).

understanding of the taxonomic position ofthis species has constantly varied. Also,
ideas regarding the habitats of those forms
referred to this species have varied (Orlov

In the vicinity of Ushguli at the foothillsof Shkhara Mountain in Svanetia, Georgia

and Tuniyev 1986).

(Muskhelishvili 1959).

We recognize at least

three closely related species within the V.

In

Borzhomi

Canyon,(Bakradze 1969, 1975;

Georgia
Bannikov et al.

1977).
f

This publication combines material previously
published in Russian by Orlov and Tuniyev (1986)
and Vedmederja et al. (1986) with additional
information.

1990 by Asiatic Herpetological ResearchIn

Lagodekhi, Georgia (Zoological

Institute, the

USSR Academy of Sciences,

Nos. 8389 and 13769).

Asiatic Herpetological ResearchVol. 3, p. 2

April 1990

FIG. 1. The distributions of the vipers from the Vipera kaznakowi complex and the Vipera ursini
complex in the Caucasus. 1. V. kaznakowi, 2. V. dinniki, 3. V. darevskii, 4. V. ursini renardi
A -V.
(northern population) and V. ursini eriwanensis (southern population). Type localities:
kaznakowi; B, B' -V. dinniki; C-V. darevskii. Localities: 1 -Makh-Khokh Mountain; 2 -thesettlement of Khasav-Yurt; 3 -Lagodekhi; 4 -Benis-Kheri Canyon; 5 -the settlement of Khopa, Turkey;
6 -Mikhailovsky Pass; 7 -the settlement of Ubinskaya; 8 -the town of Maikop; 9 -the mouth of the
Urushten River; 10 -the Fisht-Oshtenovsky Massive; 11 -Shkhara Mountain. Extreme points of the
distributions are marked.

Caucasus vipers
even neighboring populations.

All three species, Vipera kaznakowi, V.
dinnikiand V. darevskii, differ inoverall range of themorphology and ecology (Vedmederja

Hence, there is an accumulation of uniquecharacteristics in populations.
Very

1984; Orlov and Tuniyev 1986).

isolates

complex situations emerge in sympatricareas of closely related species

History of the study of the Vipera
kaznakowi complex.

Thetaxonomic

problems

complex.

and

This

of"

this

may be connected withnatural hybridization.
Occasionally theidentification of some individuals fromcaused contradictions and has confused
zoologists for nearly a hundred years. Theintergrading populations is difficult. Thisparticularly concerns Vipera dinniki in theareas of its interactions with V. kaznakowi,
and additionally with the vipers of the V.

fragmented highland landscapes within theursiniinterrelationships of the viper forms of theEurosiberian group in the Caucasus hascomplex (Bonaparte).

The high

April 1990

Asiatic Herpetological ResearchVol. 3, p. 3

FIG. 2. Type localities of the vipers of the Vipera kaznakowi complex proposed over the history of thestudy. A -V. kaznakowi Nikolsky, described in 1909, collected from Tsebelda, the vicinity of Sukhumi,
Abkhazia, Georgia. B, B' -V. Berus dinniki Nikolsky, described in 1913, from the upper reaches of the
Malaya Laba River, the Caucasus Reserve, Krasnodarsky Territory and Svanetia, Georgia (respectively).

C -V. tigrina

Tzarewsky, described in 1916, taken from the Northern Caucasus. D-V. berus ornata
Basoglu, described in 1947 from the settlement of Khopa, Artviisky Vilayet, Turkey. E -V. darevskii
Vedmederja, Orlov, and Tuniyev, 1986, from Legli Mountain, the Mokrye Mountains, Gukasyansky
Region, Armenia. F-V. kaznakowi orientalis Vedmederja, 1984 from the eastern part of the Main

Caucasus Ridge.

degree of phenotypic polymorphism invipers from the V. kaznakowi complex
also creates additional problems in theevaluation of their taxonomic position.

colored" vipers in the Northern Caucasus
defined as Vipera berus. Boettger (1893)
regarded the dispersal of two closely related
viper species, V. berus

and V. renardi

from the Caucasus as a phenomenon that

Due to the complexity in identification ofthese vipers and the absence of definite
localities for specimens an analysis of oldliterature gives only an approximate idea
regarding the relationships of the Vipera

kaznakowi complex.

Rossikow (1890) mentioned "multi-

deserves attention.
Nikolsky (1905)
considered V. berus to be associated withTranscaucasia whereas V. renardi inhabits
steppe areas of the Precaucasus andmountains of the Northern Caucasus.
Leister (1908a) in his sketch on thegeographic distribution of V. renardi and
V. berus within the Caucasus wrote that V.

Asiatic Herpetological ResearchVol. 3, p. 4

renardiassociated with steppe areasisApril 1990

vicinity of Grozny is a sympatric area).

which separate the Caucasus from thehabitat of V. berus, and also the mountainsof the Northern Caucasus.
He alsoconfirmed Boettger's and Nikolsky's
opinions that V. berus

also appears inTranscaucasia.
In
Transcaucasia there

theisnorthernanisolated

population. According to Boettger (1899),
the individuals of V. berus which arepreserved at the Caucasus Museum (nowthe Museum of Georgia) were collectedfrom Suani,

Tiflis, Avar, Kodzhori,
Khasav-Yurt, and Kazikoparan, whereas

those from the Senckenburg Museum(Frankfurt-am-Main, West Germany) weretaken from Sukhumi, Georgia.
Leister (1908a,b,c) gives a list of Vipera
specimens in the collection of the

berus

Zoological Museum of the Academy of
Sciences (now Zoological Institute of the
USSR Academy of Sciences) taken from

Caucasus:
1) from Tiflis and
Lagodekhi regions, Georgia and 2) from
Yelenovka near the Gochka Lake (now theSevan Lake). Leister then states that hefound V. renardi on the bank of the
Gochka Lake at the same locality where
Kessler and Brandt collected V. berus (thisspecimen, ZIN 5478, collected by Brandt
we determined to be V. ursini ). On thebasis of the findings of Kessler and Brandt
and his own findings, Leister comes to theconclusion that both species, V. berus andtheV. renardi, live sympatrically. Concerning
the specimens collected by Kessler and

Brandt, Nikolsky (1905) writes that theirrostral touches only one apical scale like in

V. renardi. In all other characters they look

more

like

V.
to

(1908a)intermediate

berus.

subspecies, V. berus dinniki. He defined
the viper's range as the northern andsouthern slopes of the Caucasus Ridge
from the Malaya (Small) Laba River
headwaters up to Elbrus Mountain. Morits(1916) also recorded V. berus in the

Northern Caucasus.

Tsarevsky (1916)

described a new form V. tigrina that was
and V.
closely related to V. renardi

kaznakowi.

Unfortunately he cited thespecies locality solely as "the Northern
Caucasus."

Nikolsky (1909, 1911, 1913, 1916), theauthor of Vipera kaznakowi and V. berus
dinnikidescriptions, studied therelationships of the species V. kaznakowi,
V. berus, and V. renardi (= V. ursini
renardi ) and noted the difficulties in
identification of these forms.
Nikolsky(1913) described V. berus dinniki from asingle specimen collected by N. Y. Dinnik
at the upper reaches of the Malaya Laba

certainRiver, and from three specimens found by
Shelkovnikov in Svanetia, Georgia. In hisanpossibleand V. renardi.

next monograph, Nikolsky (1916)
discussed both forms:
Coluber berusThis led Leisterconsiderthatspeciesisancestral form of V. berus

Since Nikolsky (1909) described Vipera
kaznakowi, a number of synonyms havebeen proposed for this species as new
forms have been described and new
combinations proposed. A number offorms, primarily from the Northern
Caucasus and from the eastern range of V.
kaznakowi have confused taxonomists as
to their relationship to V. kaznakowi.
These forms were placed in different
combinations within V. berus and V.
ursini (Nikolsky 1913, 1916; Basoglu
1947; Knoepfler and Sochurek 1955;
Kramer 1961). Nikolsky (1913) assigned
the common Caucasus viper to a newa—Nikolsky (1913) referring to the vipers
from Transcaucasia listed by Kessler,
Brandt and Leister nevertheless concluded
that they are V. renardi. Leister (1908a,b),

having analyzed the collections, concludes
that the territory of the Northern Caucasus
and Transcaucasia are inhabited by V.
berus and V. renardi. He thought that V.
berus and V. renardi live sympatrically in
the Northern Caucasus (for instance, theand Coluber kaznakowi. Theofthe former was defined as therange
Caucasus Mountains on both sides of theGreat Caucasus Ridge. The upper reachesof the Bolshaya (Big) Laba River was
designated as a sympatric zone for V. berus
dinniki and V. renardi. In his opinion, V.
renardi was closely associated with the
Caucasus Black Sea coast and the northernslope of the Caucasus Ridge.
dinnikiApril 1990

Asiatic Herpetological Research

Dinnik (1926) noted that Vipera berus
and V. kaznakowi occurred in the Northern
Caucasus. Krasovsky (1929) indicated thatV. berus inhabited Khasav-Yurt andChernov
Rutulsky Kanton, Dagestan.
in
berus
dinnikirecordedColuber(1929)
the highlands of Ingushetia on the southernThe
slope of Makh-khokh Mountain.
D.
collected
werebyKrasovsky
specimensin 1926 (Fig. 1).
(1933)
Krasovsky
included both V. kaznakowi and V. berus
in the fauna of the Caucasus State Reserve.
Bartenev and Reznikova (1935) concluded
that V. kaznakowi and V. berus dinniki
were distributed in the western Caucasus,
whereas in the alpine, V. ursini renardi
also occurred.
Rostombekov (1939)
recorded V. kaznakowi as part of the fauna
A new form, V. berusof Abkhazia.
ornata, was described from northeast
Anatolia (Basoglu 1947). Later on Mertens(1952a) synonymized it with V
kaznakowi.
Terentyev and Chernov(1949) regarded V. tigrina Tzarevsky andV. berus dinniki
Nikolsky as juniorofV.
kaznakowi
synonyms
Nikolskywhich is, in their opinion, related to V.
.

Vol. 3, p. 5

tigrina and V. berus ornata are mentioned
as junior synonyms of V. kaznakowi.
Vedmederja (1977) recorded V. kaznakowi

In his opinion
Adgaria, Georgia.
V.
kaznakowi is a
(Vedmederja 1984)
polytypic species comprising four
subspecific forms. Tertyshnikov (1977),
in defining ecological and zoogeographic
subdivisions of the herpetofauna of theinNorthern Caucasus, noted that V.
kaznakowi tends to be restricted to westernand southwestern montane regions. The
most recent taxonomic works dealing withthese vipers state that V. berus occurs
neither in the Caucasus nor in the
Precaucasus. Hence, V. kaznakowi is thesole valid name with regard to shieldheaded vipers from the western Caucasus
The
Isthmus and northeast Anatolia.
taxonomic status of the eastern populations
from Dagestan and Armenia was notdiscussed (Terentyev and Chernov 1949;
Mertens 1952a; Mertens and Wermuth
1960; Klemmer 1963; Bannikov et al.
1977; Baran 1977; Harding and Welch1980).

The study of old collections andshows that the majority ofursini renardi.

literature(1956) presented a new
Darevskycombination of names for the viper fromthe Gukasyansky region, Armenia: Vipera

researchers did differentiate the vipers fromthe Vipera ursini complex and the V.

Fyodorov (1956) recorded
V. berus in the forests of the premontane
area of the Stavropolsky Territory and V.

difficulties

kaznakowi

of the V. kaznakowi complex. Referencesthat V. berus occurs in the Northern
Caucasus and Transcaucasia more often
than not concern snakes from the V.
kaznakowi complex rather than those fromursini renardi.

in the subalpine belt. In hisofthesnake fauna of Abkhazia,
surveyGeorgia, Milyanovsky (1957) mentions V.
kaznakowi. Muskhelishvili (1959) records
V. kaznakowi on Mount Shkhara from thevicinity of Ushguli, Svanetia, Georgia.

Kramer (1961) regards V. tigrina

kaznakowi

The

complex.
indefining

the

major

systematic

position and taxonomic status concerned
primarily the vipers from the eastern range

the V. ursini complex.

a junior

synonym of V. kaznakowi, whereas V.
berus dinniki should be synonymized with
V. ursini renardi. Bakradze (1969) found aAn

analysis of literature data andmorphometric characteristics of 141 viper

specimens allowed Vedmederja

et

al.

female V. kaznakowi in the town of BanisKhevi, near Borzhomi, Georgia. On thebasis of this finding, he suggested that the
species habitat covered the entire AdzharoImeretinsky Ridge and some of the(1986) to designate three species within the

Trialetsky Ridge.

2. V. dinniki

In the field guide of Bannikov et al.
(1977) the names Vipera berus dinniki, V.

Tuniyev, 1986

Vipera kaznakowi
1.

V. kaznakowi

3. V.

complex:
Nikolsky, 1909

Nikolsky, 1913

darevskii Vedmederja, Orlov andAsiatic Herpetological ResearchVol. 3, p. 6

Fig. 2 represents type localities of theforms proposed for the Vipera kaznakowi

complex over the history of its study.

Systematic Accounts
Vipera kaznakowi

Nikolsky, 1909

(Fig. 3, 4, 14a, Plate 1)

Chronology of species description
Vipera renardi (Christoph)
1903, 30:37 (part).

-

Silantyev,

Vipera kaznakowi Nikolsky, 1909:174;
Nikolsky, 1910:81, table 1.

kaznakowi

Vipera-Nikolsky,

1913:179-181; colored plate III (=Fig. 3,
this paper).

kaznakowi

Coluber-

Nikolsky,

1916:244-247.
Vipera berus ornata Basoglu, 1947:182190.

Vipera ursini kaznakowi
Sochurek, 1955:185-188.

-

Knoepfler andDiagnosis:

April 1990

A large snake for theEurosiberian group.

Total length reaches

650-700 mm. Dorsally the head is covered
with large scales. In size and shape theydiffer considerably from body scales.
Nostril is cut through either in the middle orslightly closer to the lower edge of thenasal.
Upper-lateral edge of snout is
Rostral normally reaches twopointed.
on upper snout. Upper edge
scalesapicalof the nasalorostral scale is slightly curved
at obtuse angles.

Scales of anterior frontal

have weak longitudinal keels. Head isbroad and normally black dorsally. Head isseparated from body by a sharp nuchal
collar.

Description: The ratio of body length to
tail length is 5.7 to 6.4 in males and 7.5 to
10.9 in females.
Unlike other species

kaznakowi
assigned to the Vipera
redandcolorscomplex,yellowprevail in
V. kaznakowi. Melanistic specimens arecommon. However, unlike complete
melanistic individuals of V. dinniki, thoseof V. kaznakowi preserve yellow or redcolor on either upper or lower labials.
Vipera kaznakowi is either black or darkbrown striped dorsally and laterally. Often
merge so that only red or yellow
remainbetween them. Ventrum is
spots
stripesVipera kaznakowi - Terentyev andChernov, 1949:270-277 (Map 15);
Bannikov et al., 1977:323-324 (map 133;
colored plate 31, 4).

kaznakowi

Viperablack.
dorsally.

Head is very broad, impressed
This fact is emphasized by aslightly pointed upper edge of snout.
Cheeks are greatly swollen. Head is wellkaznakowi -separated from body by a thin nuchal
Comparative data on scalation andsize characters are listed in Tables 1-4.

Vedmederja, 1984:8.

collar.

The English common name is the Caucasus
Viper, or Kaznakow's Viper.

Variability and comparative remarks

No. 4408, an adult female.
Collected by Y. V. Voronov fromTsebelda, the vicinity of Sukhumi,
Abkhasia, the Caucasus. It is stored in the
Caucasian Museum (now the Museum ofLectotype 1:

Georgia, Tbilisi).

+

Kramer (1961) was mistaken regarding

thisspecimen as the holotype, because Nikolsky (1909)
had not singled out a holotype from the fivespecimens from which he described the species.

kaznakowi(Fig. 4, Plate 1) is
moreundoubtedly
closely related to V.
V.

berus and V. ursini. Supposedly, wheninteracting with V. ursini renardi and V.
ursini eriwanensis it forms two species of
V. dinniki
and V.
hybrid genesis:
darevskii. From all listed forms it differs
by: 1) the extraordinarily great width of thetriangular head, 2) the head width of
specimens of this species equals the
distance between tip of snout and posterior
angle of mouth slot, 3) the "cheeks" aregreatly swollen, hence a broad furrowAsiatic Herpetological Research

April 1990

Nikolski.

A. 3. E.

Nova species viperae.

Tom J.

Vol. 3, p. 7

H3B. K»bk. My3. T. V.

III.

Vipera kaznakovi sp

KiiiniiKiir C,

TI-

E.

n

FIG. 3. Type specimen of Vipera kaznakowi Nikolsky, 1913 (reproduced from Plate III, originallyprinted in color, Nikolsky 1913).

Vol. 3, p. 8

TABLE 1.

Asiatic Herpetological ResearchComparative scale characters of the three viper species of the "Vipera kaznakowi

April 1990"

complex.

Asiatic Herpetological Research

April 1990

Vol. 3, p. 9

TABLE 4. Comparative morphology of vipers in the "Vipera kaznakowi " complex.
V. dinniki, 68 specimens

Vipera kaznakowi, 64 specimens^475 mm
600 mm

• SVL 8 8

SVL V 986/ CI. x 5.6-6.4
SVL 9 9/ CL as 7.5-10.9• SVL• Head is either impressed or flat

• SVL 8 8412

SVL 9 9y<V. darevskii, 9 specimens

mm^^486 mm• SVL 6 6slightly

SVL 88 / CL a 6.0-6.5
SVL 9 9 / CL a8.4-9.3• Headeitherisflat• Head is either flat or slightly

or slightly

protuberant dorsally• Edge of snout is roundedroundedmm
mm

SVL 9 9/ CL as 7.9-13 5protuberant dorsally
is

258
421

• SVL 8 8/ CL a 5.9-7.4

dorsally• Edge of snout?

SVL 9 9?*• Edges of snout are slightly pointed
Anterior edge

laterally.

isalittlerounded• Rostral is broad.

Usually touches<

two apical scales (in 91% specimens)

Rostral(inisnarrow reaching either one

48.6%) or two (in 51.4%) apicals

rarely one apical (in 9%)

itsof the frontal and the rostralisparietals or equalsi• The width of the frontal
1.48-1.71 of

The distance between anterior edge ofthe frontal and the rostral

equal

is

equal to0.77-1.35 of the frontal edge

edge

either smaller thanisThe width of the frontal is equal to1.13-1.83 of its length• The distance between anterior edge

• Frontalilength

to 0.75-1.05 of the frontal

one (in three specimens) or two (in six
specimens) apical scales• The width of the frontal is equal to1.21-1.72 of• Rostral is more narrow and touches

(

them

Frontal

is

its

is

equal tolength• The distance between anterior edge ofand the rostral is equal to

the frontal

0.67-1.07 of the frontal edge

either smaller than parietals• Frontal is larger than parietals

or equals them

• Large lower oculars are separatediLarge lower oculars are separated fromfrom frontal either by one row (infrontal either

87.0%) or rarely by two rows of

two rows (in 41.5%)

by one row (in 58.5%) or• Large lower oculars are separated fromfrontal only by a single

scales (inrow of smallnine specimens)allsmall scales (in 12.4%)• The upper pre -ocular does not touchthe nasal in 96.1%.

In(

3.9% it does

The upper pre-ocular does not touchthe nasal in 86.6%.

In other casesitreaches the nasal• Nostril

either cut through in the

Nostriliscut through in the center or

middle of nasal or is slightly shiftednasal.

It

may be shifted downwards

downwards

exceedingly rarelyisi• Nasal does not touch rostral<• Body scales with expressed keels;<

Nasal does not touch rostralBody

the nasal in

two specimens.

specimens

does

it

In

seven

• Nostril is cut through in the lower partof the nasal• Nasal does not touch rostral76%

scales have expressed keels.

some scale rows that reach ventrals

of scales that reach ventrals have no

have no keels

keels,

18.4% have slightly expressed

keels,

5.6% have well-expressed keels

supporting ribs are given in Tables 1-4.

• The upper pre-ocular does not touch• Body scales have expressed keels.

Some scale rows that reach ventrals
have no keels

dorsal stripe merges with lateral stripes sothat either red or yellow spots arranged inColoration. Vipera

kaznakowi

aretwo rows along the dorsum remain.

anddorsal stripe can be either zig-zag shaped or

extraordinarily bright among shield-headed
Red hues prevail in the color
vipers.
Newborns
are similarly bright inpattern.
color, primarily red brown, unlike the grayin the shape of an even broad line.
Ventrum is black. Head pattern of adultincredibly

diverseincolor

young of V. dinniki. The typical intensive
red color appears in V. kaznakowi after
they have shed twice. Complete melanists
are frequent in populations. Often the blackThe

specimens normally blends with the dorsal
In immature specimens, the head
stripe.
pattern may be separated from the dorsal
stripe by a light interval thatafter maturation.

disappears

Vol. 3, p. 10

Asiatic Herpetological Research

April 1990

FIG. 4. Vipera kaznakowi (ZIN 1 1529) from the isolated forest of Babuk-Aul.

PLATE 1 Vipera kaznakowi (ZIN 1529) from the isolated forest of Babuk-Aul.
PLATE 2. One pattern variation of Vipera dinniki from Krasnaya Polyana (ZIN 12153), a male.
PLATE 3. Vipera ursini eriwanensis from the valley of Kassach, the foothills of the Ara-Fler Mountains.
.

1Orlov and Tuniyev

Plate

Asiatic Herpetological Research1% Muy^ \JUVipera kaznakowi

Orlov and Tuniyev

Plate 2

Asiatic Herpetological Research

Vipera dinniki

\ au^

if\\

Orlov and Tuniyev

Plate 3

Asiatic Herpetological Research

Vipera ursini eriwanensis

1 2Uu#W

Asiatic Herpetological Research

April 1990

Maximum bodylength is longer in females (up to 600 mm)

Sexualdimorphism.

than in males (does not exceed 457 mm).

Males have longer tails (Table 2).
Accordingly, the number of ventrals isgreater in females, whereas the number ofsubcaudals is greater in males. Males are
more slender. Sexual dimorphism incoloration is feebly marked. Melanistic
individuals are more frequent amongfemales.

Age variability. Vipera kaznakowi

areborn with the typical adult color andHowever, in newborns thesepattern.
characters are less pronounced than in
adults.
Newborn snakes may be eitherorreddish.
According to our
pinkishobservations on birth and development ofsnakes from the Sochi-Khosta populations,the coloration becomes stronger with each
Maximal color
subsequent shedding.
isachieved
theseason after theintensitybyfirst hibernation.

Melanistic specimens are born with thetypical species pattern, but their colorationis darker. The coloration becomes darker
during subsequent sheddings and elementsof the pattern merge.

New born litters are homogeneously
On maturing, the coloration

colored.

becomes

diverse.
Phenotypic
polymorphism in mature vipers of a litter isEither partial or nearly complete
great.
melanism is observed in all vipers from thispopulation. This is typical for a number ofother animals from humid subtropic areas
adjacent to the Black Sea coast (Bartenev
and Reznikova 1935). Specimens showing

Vol. 3, p. 11

northern slope of the Main Caucasus
Ridge. Here Vipera kaznakowi occurs
along the foothills from the settlement of
Ubinskaya in the west to the town of
Maikop, USSR in the north and the mouthof the Urushten River in the east (Fig. 1
The
from Vedmederja et. al. 1986).
toanelevation
species generally occurs upof 800 m. Along the river valleys of theBlack Sea coast, it may occur up to anor even higher.
elevation of 1000m

Vipera kaznakowi is a forest dwelling
It occurs on montane wooded
slopes, in the bottoms of humid canyons,and in meadows adjacent to forests. It is
recorded in Quercetum azaleorum andspecies.

Quercetum coggygriosecornosum oakgroves, in mixed subtropic forests withevergreensubforestsofQuercusiberica, Alnus
barbata, Fagus orientalis, Taxus baccata,
Laurocerasus officinalis, Buxux colchicus,

hartwissiana,

QuercusCastanetum colchicum, Fagetum nudum,
Salictum fontenale, and Alnetum
In addition, thisstrut hiopteridor sum.
in
found
is
polydominant forests inspeciesriver terraces and large overgrown
outcroppings. At the upper limits of its
elevational range, the species reaches
coniferous forests. It is recorded within theecotone of Fageto-Abieta

athyridosa-

but the viper neverinto
coniferous forests.
penetrates deep
kaznakowiisalso present within
Viperatransformed areas such as meadows formed
after forests are cut, fruit orchards, kitchen

maxtoherbosa,gardens, vineyards, and dilapidated parks(Red Data Book of the USSR 1978, 1984).
As a rule, vipers occur within sites where
the density of lizards is high.

maximalmelanism always preserve
elements of orange and red on the throat
scales and chin shields, the rostral, upperlabials and subcaudals.
MelanisticofV.
dinnikican havespecimens
completely black coloration by maturity.

Geographic range and ecology.

The

species ranges along the BlackSea coastfrom the town of Khopa, Turkey andSuramsky Pass in the east, then throughout
Kolchida (Colchis) up to Mikhailovsky
Pass in the west. It is then found up to the

Typical biotypes are as follows:smallmeadows and other illuminated spots inforests with an

exposure providing highsolar radiation in conditions of humid
subtropical climate of the Black Sea coastof the Caucasus. The sites are located in
the vicinity of standing water, where rocks
exit that are suitable for hibernation. The

climate of the Caucasus coast is greatly
dependent on topographic conditions of thearea that forms a narrow line between the
Caucasus Ridge and the Black Sea. It isAsiatic Herpetological ResearchVol. 3, p. 12

situated from northwest to southeast along

The wall of the Ridge starts from
Near
Novorossiisk it reaches an
Anapa.
elevation of up to 600 m. Near Tuapse, thewall exceeds 1000 m above sea level. At
the latitude of Sochi, it is 3000 m high.
The Great Caucasus Ridge presents a
the coast.

barrier for cold northeastern winds.

Thisbarrier separates the warm humid coastfrom the comparatively cold continental
Prekuban area (Korostelyov 1933). In the

east

the

Adzharo-Imeretinsky

Ridgeseparates the humid subtropical Kolchida(Colchis) from the arid regions of theEastern Precaucasia. The climatic zone of
the Black Sea coast of the Caucasus
appears to be extremely favorable forVipera kaznakowi. The northwestern most
(Mikhailovsky
Pass and Stanitsa Ubinskaya) coincide with
Pontic and northwestern floristic regions.
localities for this species

climate, and vegetation thenorthwestern region is a continuation of thesouthern coast of the Crimea.
In relief,is marked by 1)
inthe foothills, 2)
subtropical vegetation

The Pontic region

temperature stability, and 3) highhumidity (Korostelyov 1933). The limited
dispersal of the species along the northernslope of the Big Caucasus is in the regionof the "Kolchidian (Colchis) Gates." Dueto the lowering of the western portion of
the Main Caucasus Ridge, interchange offlora and fauna of Kolchida (Colchis) and
the Prekuban area occurred in the past andpresently occurs. Warm humid air from theBlack Sea breaks through in this part of the
Ridge. On the northern slope of the Big
Caucasus, it creates a small refugium on a
Kolchidian (Colchis) type associated with
great

Castanea satyra, Buxux colchicus, Ostrya
carpinifolia, Corylus colurna, and others(Galushko 1974; Kharadze 1974;

Kholyavko

etherpetofaunaal.

1978).

of

Kolkhida

Of theB ufoverucossimus, Triturus vittatus ophryticus,
Pelodytes caucasicus, and Lacerta derjugini
occur along with V. kaznakowi. Along theentire area of of the Black Sea coast of the
Caucasus, the viper is rare. In a number of
In somedisappeared.
specimens of V.
kaznakowi might be encountered.

spots

it

has

situations, only single

April 1990

The highest density of Vipera kaznakowi
have been observed are in rocky

thatoutcroppings within the forest belt in themountains of the Caucasus Reserve. In
beech forests along a road in the valley ofthe Achipse River, we have recorded three
specimens per kilometer of road. Single

specimens have been recorded in thevicinity of Khosta, Babuk-Aul, Guzeripl,
and Kisha Kordons. The density of theviper is also low in the southern habitats in
Adzharia and Lazistan, Georgia (Basoglu
1947; Vedmederja 1977; Basoglu andBaran 1977).
Anthropogenic factors are responsiblefor declining numbers and populations ofVipera kaznakowi. There are pressures due
to recreational use of health resorts along

the Black Sea coast,ploughing of sites
near the foothills, and to a smaller extent,
hay-mowing (Red Data Book of the USSR
1978, 1984). Within some resort areas V.
kaznakowi is completely extinct.
feeds on various
Different populations show
specific feeding patterns with regard to preyavailable. According to data recorded in
captivity, individual preference is observed.
When stimulated to regurgitate, thefollowing food species were recorded in theDiet. Vipera kaznakowi

animals.

sylvaticus, a forest
mouse, A. agrarius, a field mouse,
Microtus majori, a juvenile specimen, M.
field:

Apodemusgud, Sorex raddei, Lacerta saxicola, L.
derjugini, L. praticola and L. agilis. In thecollection of the Zoological Museum of the

Moscow State University a viper specimen
from Bebysyry Lake is preserved. In its
stomach a juvenile grass snake (Natrix
natrix ) was found. Immature individuals
feed on juvenile lizards of the above
mentioned species, and to a lesser extent,
on Orthoptera.
In captivity adult vipers readily take any

small rodents, fledgling sparrows, lizards,

and pieces of chicken.
Young vipers
usually start on small lizards and crickets
and Achaeta(Grillus bimaculatus
domestica ). After some months they
After theusually take newborn mice.
in 5-7diesbiteanormally
prey
viper'sAsiatic Herpetological Research

April 1990

The snake never persecutes its

minutes.

finds the prey some time later
using its olfactory organs. Swallowing
ranges from one minute to 3.5 hours
depending on prey size and the state of asnake. Complete digestion in the wild
takes up to five days. In captivity digestion
It

prey.

30 to 40 hours at stabletemperature. Optimal day temperature is26-30°C, and night temperature 18°C

may

take

during activity period.

Shedding. Mature vipers normally shed 2
to 3 times during the activity period.
General shedding is observed in June.
New born vipers shed in the first hours
after birth. Before entering hibernation,
they shed again.
Reproduction. Mating is recorded from lateMarch to April. Birth occurs in lateAugust. Females give birth to 3 to 5
young. The observed time of birth lasts forabout two hours with intervals of 20 to 40

minutes (Zinyakova and Trofimov 1977).
In captivity, the majority of females givebirth at night, between 2400 and 0600 hrs.
Some individuals are born in transparent
capsules, which the neonates leave in thefirst minutes after birth. Females reproduce
annually. Gravid females continue to takefood right up to birth.

Development of young. Neonate Vipera
kaznakowi have a mean body length ofVol. 3, p. 13

Like other vipers, Vipera
kaznakowi is conservative in territory use.
The same individuals can be encountered in
Territoriality.

the same places during different seasons.

Vipera kaznakowi utilizes considerablylarger individual ranges than V. dinniki.

Seasonal and daily activity. Along theBlack Sea coast of the Caucausus, Vipera
kaznakowi emerge after hibernation in
March at an altitude of 600 to 800 m. On
the northern slope of the Big Caucasus thevipers appear in the second half of April toearly May when the mean day temperature
is 13°to 16°C. In the foothills the vipers

enter hibernation at the beginning ofNovember at an altitude of up to 600 m. In
the upper elevational limits of its range V.
kaznakowi hibernate in late September toearly October. New born vipers are more
active than those of other age groups.

Two sharply marked peaks of daily
activity can be observed in Vipera
kaznakowi. In the morning the period ofdaily activity ranges from 0730 to 1130
hrs, and in the evening from 1630 to 1830
hrs.

At those times the soil temperaturedoes not exceed 30-32°C

inthe sites

inhabited by V. kaznakowi.

Sympatric species. Along with Vipera
kaznakowi the following species of reptiles
occur: Lacerta derjugini, L. saxicola, L.
L.

After birth the activity

Anguis fragilis,
Colubernajadum, C.
Pseudopus apodus,
jugularis, Elaphe longissima, Natrix natrix,
N. tessellata, and Coronella austriaca.
Narrow sympatric areas appear with
Testudo graeca, Lacerta trilineata, L.

2.5 months, whereas
newborns of V. dinniki never take food

Elaphe hohenackeri, Vipera ursini, and V.

144.75 mm, tail length of 14.0 mm. Meanbody weight is 4.1 g (n=8). After first

shedding on the second day after birth,
newborns start actively feeding on insects
or small lizards.

period

is

1.5

to

and almost immediately enter hibernation,
during which snakes increase in lengthfrom 10 to 20 mm. Vipera kaznakowi lose
0.3 g of initial weight during the firstmonth after birth. In the second monththey restore their initial weight and thenincreaseitenteringhibernation.

approximately

1One

g beforeyear oldmm

specimens have a body length of 200
and a tail length of 24 mm. Vipers reachpraticola,

agilis,caucasica, L. rudis, L. parvula, L. mixta,
dinniki.

Vipera dinniki

Nikolsky, 1913

(Fig. 5, 6, 7, 12, 14ab,16, Plate 2)

Species description in chronology

Radde,
Vipera berus - Boettger in
1899:286; Nikolsky, 1905:304 (adCaucasus).

sexual maturity by the third year at a SVL

of 350-400 mm.

Pelias chersea-

Menetries, 1 832:73 (part).

Asiatic Herpetological ResearchVol. 3, p. 14

April 1990

Vipera xanthina -Dinnik, 1902:34.
Silantyer, 1903, 30:37
Vipera renardi(part).

Vipera berus dinniki Nikolsky, 1913:176179.

Coluber berus-

dinnikiNikolsky,

1916:240-244.
Vipera tigrina

Tzarevsky, 1916:32-37.

Vipera ursini renardi-

Kramer, 1961:715.

Vipera ursini kaznakowi
Sochurek, 1955:185-188.

-

Knoepfler and

FIG. 5. Lectotype of Vipera dinniki (TheMuseum of Natural History of Kharkov State
University 26044).

exceed 486 mm. The ratio of body length
to tail length is 5.9 to 7.4 in males; 7.8 to

Terentyev andVipera kaznakowi
1949:270-271
Chernov,(map 5);Bannikov et al., 1977:323-324 (map 133,

13.5 in females. General coloration is not
kaznakowi.
as bright as in Vipera
However, specimens with bright yellow

colored plate 31,4).

and orange elements can be observed.
(Fig. 6,.Plate 2)
Normally V. dinniki
have light brown, grey, silver greyish or
green greyish color which never occurs in
V. kaznakowi. Some specimens have adark even dorsal stripe along the center ofVipera kaznakowi dinniki-

Vedmederja,

1984:8.

Vipera kaznakowi orientalis-

Vedmederja,

1984:9, nomen nudum.

the body.

The English common name is Dinnik's
Viper or the Caucusus Subalpine Viper.

Lectotype:

No. 26044, an adult female

collected by N. Y. Dinnik (Fig. 5) fromthe upper reaches of the Malaya (Small)Laba River, Northern Caucasus andSvanetia, Georgia (Fig. 2: B, B'). The
specimen is preserved at the Museum of
Natural History, Kharkov State University,
Ukraine.

Diagnosis: Total length reaches 500 to
550 mm. Dorsally the head is covered with
cut through in thelarge scales. Nostril
center of the nasal. Upperlateral snout edge
Rostralis rounded and slightly pointed.
istouches either one or two apical scales on
upper head. Three to four scale rows withno keels are between the rostral and frontal.
Head is not broad. Nuchal collar is not
expressed.

Description: In males body length is not
greater than 412 mm; in females it does not

The latter substitutes for the zig-

zag shaped stripe typical for the majority ofplate-headed vipers. Ventrum is either dark
and light spotted or light grey and dark
Neonates of V. dinniki
speckled.
have the red color of the
don't
occasionallykaznakowi. They canforV.
body typicalbe bom grey brown. The head is relatively
narrower than that of V. kaznakowi. The
nuchal collar is not expressed, unlike in V.
kaznakowi. Upper edge of snout is either
rounded or slightly pointed. Head can beeither protuberant or flat dorsally, but neverimpressed like that of V. kaznakowi. Body
is thinner andmore delicate. Comparative

data on pholidosis and size characters arelisted in Tables 1-4.

Remarks and variability.
greatly resembles
Vipera dinniki
morphologically V. kaznakowi, V. ursini
renardi and V. berus. In size V. dinniki issmaller than V. kaznakowi and larger thanV. ursini renardi(Table 2). The headnormally is slightly protuberant, seldom flatand never as broad as that of V.

April 1990

FIG. 6.

Asiatic Herpetological ResearchVol. 3, p. 15

A Female Vipera dinniki from the upper part of the Laba River (ZDN 17281).

kaznakowi. Hence, the nuchal collar ishardly marked. In body proportions it
primarily resembles V. berus. Normallythe parietals are shorter than the frontal.
Three to four lower labials reach the lower

jaw scale.

yellow or orange elements occur. Often V.
dinniki have greyish, silver greyish or
green greyish color. This is never recorded
in V. kaznakowi. For all shield-headed
vipers a zig-zag shaped dorsal stripe is acommon pattern element. In V. dinniki theeven broad dark line
which runs along the dorsum. This isoccasionally seen in V. kaznakowi, where
stripe is often an

Color. Normally Vipera dinniki is not askaznakowi.
brightly colored as V.
However specimens with either bright

it is usually represented

by a row of oblique

Vol. 3, p. 16

FIG. 7.

Asiatic Herpetological Research

April 1990

A male Vipera dinniki from the vicinity of Krasnaya Polyana (Red Meadow), [ZIN 12153].

diametrical spots. The dorsal stripe isseparated from the dark sides by lighter
lateral stripes. The ventrum is either darkwith light spots or light grey. The numberof melanistic specimens in populations

ranges from 20 to 25%.

Completemelanistic individuals of V. dinniki do nothave a single light spot in color, unlike
melanists of V. kaznakowi.

Sexual dimorphism.

Maximum bodylength is greater in females (up to 486 mm),and smaller in males (up to 412 mm).
Males have longer tails, characteristicallyApril 1990

FIG. 8.

Asiatic Herpetological Research

A Vipera dinniki from Lagodekhi, an eastern population (ZIN 13769).

thicker at the tail base (Table 2). The
number of ventral scales is greater infemales. The number of subcaudals isgreater in males. Males are more delicately
built than females. Sexual dimorphism incolor is hardly expressed. Coloration in

males

Vol. 3, p. 17

isgenerally brighter and more

contrasting.

Neonate vipers arevariability.
like
adult
individuals.
However,patterned
the general color is normally grey, unlike
bright red neonates of Vipera kaznakowi.
Only after the third shedding, faintAgeAsiatic Herpetological ResearchVol. 3, p. 18

April 1990

stronger with eachsubsequent shedding. Maximum color
strength is reached by maturity. Melanistic
specimens are born with a specific color.
They gradually darken with each shedding.
By the third year, they acquire a blackoccurs widely in moraines
moss, lichens andovergrown
The
Thymus.
biotype of V. Kaznakowi isin the vicinity of water.
locatedalwaysHibernation sites are also located in theimmediate vicinity of summer biotypes. All
types of rock outcrops are inhabited.
Vipera dinniki can be found in limestone,

velvet color.

slate, and crystalline outcrops.

Geographical range and ecology. The
species ranges from the Fisht-Oshtenovsky
Mountain Range in the west up to Mount
Shkhara in the east. The eastern limits ofthis species distribution has not been
worked out and additional eastern localities

Climatic conditions within the range ofVipera dinniki are much more severe thanthose of V. kaznakowi.
However, theinseveredistribution
viper'shighlands
coincides with the mildest climatic places inthis severe area. Vipera dinniki commonlyoccurs on slopes with a southern or
southeastern exposure. For instance, on

coloration typical for the species (yellow,
reddish, greenish) emerges in V. dinniki.

Colorbecomesare probable.

The southern distributionalruns along the dark coniferous
highland ecological belt on the southernslope of the Main Caucasus and South

limit

Frontal Ridges.

The northern limit goesfrom Mount Shkhara to the west along thecrest of the Main Caucasus Ridge up to theBolshaya (Big) Laba River head where it
passes on to a northern macroslope. In thenorth it occurs on the Peredovoy (Frontal)
Ridge and reaches the Fisht-Oshtenovsky
Mountain Massive (Orlov and Tuniyev
Research during 1987-1989
1986).

showed that Vipera dinniki occurs further
to the east than previously known (Fig. 9).

The problem of subspecific status of theeastern populations and their interaction
with the vipers of the V. ursini complex
will be addressed in a future paper.

The elevational distribution is generally
between 1500 and 3000 m. Occasionallythe viper may descend slightly lower.
Vipera dinniki is a subalpine montane-

meadow species. It tends to be restricted tothe upper forest belt, subalpine and alpine

meadows, rocky outcroppings and montane
moraines.

can be observed inVipera dinniki
vegetation associations of Betuletum
calamagrostidosum, Pinetum mystillosum

subalpinum, Fageto Betuleto-Sorbetum
altherbosa-subalpinum, also in Aceretum

trantaltherbosum

subalpinum.

It isassociated with rock outcrops interspacedwith shrubs of Rhodoretum caucasicus
subalpinus, subalpine highland herbs androck debris.

It

with

Mount Aibga, 23 km away from the sea, at
an altitude in excess of 1800 m, the mean
temperature in January is -0.1°C(Korostelyov 1933; Shkadova 1979).
Mean temperature in July is only 13.7°C on
Mount Achisko at an altitude of 1750 m (37km away from the sea). However, great
temperature drops never occur in this area,even in winter. As regards to soil freezing,

minimum temperature in January is -7°to 8°C (Selyaninov 1933). Due to solar
radiation on slopes with southern andsoutheastern exposures, the vipers manage
to maintain high body temperatures during
activity periods.

Vipera kaznakowi reaches the edge, butdoes not penetrate deep into the dark
coniferous belt. Vipera dinniki, at its lower
limits, reaches the edge but never penetrates
the dark coniferous belt. Thus, the dark
coniferous belt is a barrier between them.
Occasionally, in spots where this belt iseither absent or is fragmented, for instancealong river valleys, both species occur
sympatrically, forming a narrow line withintergrading characters. The valley of the
Mzymta River may serve as an example ofwhere the two species come in
The limited distribution of thespecies on the northern slope of the Main
Caucasus Ridge might be connected witha sitecontact.

the increase of the mountain's aridity andsevere climate towards the east.

Xerophytization is observed from west toeast in the subsequent change of beechAsiatic Herpetological Research

April 1990

Vol. 3, p. 19|*| V 'ipera darevskii
J Vipera dinnik

1 Vipera kaznakowi| Viperaursini eriwanensis

J Vipera ursini renardi

FIG. 9. Distribution map of shield-headed vipers in the Caucasus.
forests

and Abies

and further(Adamyants

forests to pine forests,

east,
steppe areas
Kharadze
1971,
1974,
Lavrenko 1980; Agakhanyants 1981). Thischange is noted in amphibians and reptiles.
Along with V. dinniki, Lacerta derjugini
and Pelodytes caucasicus fall out andmore xeric species such as Bufo viridis,
Lacerta agilis, and Vipera ursini renardi
dominate.
toPopulation density is different in various
It is maximal on rockparts of the habitat.

outcrops and moraines in the subalpine beltof the Caucasus Reserve.
In July, in
subalpine meadows of the Gertsen Ridge

and along the valley of the Bezymyanka(Without name) River, 1700 to 1900 m, we
have recorded as many as 5 to 7 vipers per
1 km of a route.
At the same time, on anm

we have recorded up to 6
This
is at an elevation of 1800
specimens.
on the Aishkha Ridge in a subalpine
meadow of Aceretum trantaltherbosum
area of 500

2

,

m

subalpinum. In late June to early Julyalong the Moloshnaya (Milk) andSumasshedshya (Crazy) rivers of the

Mzymta River basin, up to 4 specimens per

100 to 500 m of a route were recorded. In
August, on a bank of the high altitude
Kardavych Lake, we recorded up to 8
specimens per 300 m of a route. On the
Aspidny Ridge, 2000 m above sea level,
we recorded 5 to 6 specimens per 300 to

400 m of a route.

According to Bozhansky (1979), thedensity in the subalpine belt is 2 to 6
specimens per hectare.
Occasionally,seasonal accumulations of up to 30-40

Asiatic Herpetological ResearchVol. 3, p. 20

individuals per hectare are possible. In
other regions the population densities aremuch lower. Degradation of localities isdue to intensive cattle grazing in subalpine
meadows. Overall numbers of both V.
kaznakowi and V. dinniki are estimated atsome dozens of thousands (Bannikov andMakeyev 1976). Apparently the figure may
primarily concern V. dinniki as the numberof V. kaznakowi

is rather small.

Diet. Adult Vipera dinniki eat fewer types
Forof food items than V. kaznakowi.

instance, in the highlands there are viper
populations that prey either on lizards or on
small mammals as no other food items are

April 1990

conditions, female Vipera dinniki have a

reproductive cycle of many years.

Development of the young. In late July toearly August viper embryos reach 70 mm inlength (Orlova 1973). Mean body length ofmm, tail length is
14.8 mm, mean weight is 3.1 g (N=28).
In the highlands almost right after birth thevipers enter hibernation. Unlike newlyborn Vipera kaznakowi, they do not feed
until the next season. By the third year thevipers become mature.
the neonates is 131.0

Gravid females tend to moveon small areas ranging from 1-4 to 51 m 2
Territoriality.

When field collected vipers were
stimulated to regurgitate, the following prey

Their individual places to a great extent (upwere observed: Apodemus sylvaticus,
Microtus majori, Sicista caucasica,
fledglings of ground nesting birds such as
Anthus spinoletta, and Lacerta caucasica.
Immature specimens feed on Orthoptera
and small lizards. In captivity V. dinniki

vipers actively utilize only

available.

takes similar food items as V. kaznakowi.
It is often observed that a bitten mouse or afield-vole makes a few desparate leaps,
falls into a deep crack between rocks anddies there. The viper skillfully catches its
prey between its teeth, drags it out onto alevel spot, then drops it, examines it fromall sides and having found the prey's head,swallows it. This behavior is typical for

vipers from the
V.
such
as
raddei, and V.
Caucasus,other rock dwelling

ammodytes.
Shedding.

Overall, shedding is observed

June and in late August to early
September. Neonate vipers shed during theinfirst hours after birth.

to98%) overlap.

Within their sites the
2 or 3 places

where they may be encountered at different
times and under various weather
conditions. During the morning the vipers
can be found in places with a western
exposure. Normally, this place is on a rock
The vipers
surface shaded by a bush.
utilize direct sun for a short time, leaving apart of the body under the sun and a part
under a half shaded area. From 1 100-1200
the vipers retreat to burrows and normallyIn gloomy
appear after 1500 hours.
weather they lie flat on a rock during theentire period of activity. This increases thesurface of the body contiguity with the
On rare hot days, after midday thevipers never reappear on the surface.
Males and non-reproducing females emerge
much more seldom, normally after they
have taken food.
Bozhansky (1984)rock.

observed some specimens using the same
territories for three seasons.

Some days later they
Insolation is of great importance in theenter hibernation.

Gravid females take foodtwomonths before giving birth,andrarelyviper's life.

Reproduction. Copulation occurs in late
April to May (Bozhansky 1984). Birth ofneonates on the northern slope of the Main
Caucasus Ridge occurs in August. On thesouthernitoccursslope
throughout
September. Bozhansky (1983) found twogroups of specimens during the summer.
One group containing males and females
which do not reproduce that year, and theother group containing gravid females.
Bozhansky suggests that in montane

Territory utilization isentirely dependent on optimal insolation

they stop eating.

Absence of aggressivenesspatterns.
allows all adult females in a territorial group
to utilize the warmest microhabitats(Bozhansky 1983, 1984). These data werecollected by Bozhansky during threesummers at an elevation of 2000 m along

the border between the forest and subalpine
belts on Aishkha Ridge, the right ridge ofAsiatic Herpetological Research

April 1990

the Mzymta River valley, the western
Caucasus.
It is amazing that Vipera
kaznakowi from the low altitudes of thecoast have considerably larger individualsites. The females do not have a multi-year
cycle and never stop taking food whilegravid. The adaptations in the highland V.
dinniki are very important in relatively low

temperatures, rainy periods, and when it
suddenly becomes cold.

Vol. 3, p. 21

shifted from 1700 to 2000 hrs. In gloomy
weather snakes are active throughout

daylight at temperatures higher than 10°C.

At 8°C vipers are not seen on the surface.
Gravid females are seen on the surface evenin drizzling rain. Even if the temperature is
10°C, body temperature in vipers is 30°C,
and cloacal temperature is 26-28°C due toaccumulation of warmth from solar
radiation.

This is an important thermal

adaptation of a number of highland reptiles.

Seasonal

dynamics in populations.
monitoring in highland
populations from the Caucasus Reserve
Biologicaltestify to their climax state (the collectionsof the Zoological Museum of the Moscow

State

University

Institute, theSympatric species. Throughout nearly theentire range Vipera dinniki is sympatricwith Lacerta caucasica, L. saxicola, Anguis
fragilis, and Coronella austrica.

and the Zoological

USSR Academy of Scienceswere adopted as a zero counting point).
Seasonal dynamics within climax

At thepopulations is monotypical.
beginning of spring, males emerge first.
Then the percentage of female occurrences
gradually increases. By early-mid June thesex ratio is 1:1. In mid-late summer males
are seen on the surface less frequently thanfemales.
During the period of prehibernation at the end of summer, males

become more common again. From lateAugust to mid September, 80% of thesnakes observed are gravid females andnew born individuals. The later can beobserved on the surface even when adults
have entered hibernation. During thisperiod, the location of new bom individuals
does not necessarily correlate with rockoutcrops which are used as winter shelters.
The newborn snakes migrate much morethan adult snakes.

Vipera darevskii Vedmederja, Orlov,
andTuniyev, 1986
(Fig. 10, 11, 12, 13)

Chronology of species description
Vipera kaznakowi dinniki-

Darevsky,

1956:128.

Vipera kaznakowi darevskii-

Vedmederja,

1984:8, nomen nudum.

The English common name is Darevsky's
Viper.

Holotype: ZIN 19934, an adult female
from Legli Mountain, the Mokrye (Wet)
Mountains, Gukasynsky region, Armenia.
The specimen was collected in June, 1980
The specimen isby I. S. Darevsky.
preserved at the Zoological Institute, the
USSR Academy of Sciences, Leningrad(Fig. 10).

Seasonal and daily activity. In the spring
vipers emerge from mid April to May whenmean day temperature on

the surface

reaches

11°C.
Seasonal activity isonweather
conditions. In thedependentoftheCaucasus
Reserve the firsthighlands
snow usually falls in the second half ofSeptember and snow is present until May.
Snow cover is 7-8 m thick. Hence, Vipera
dinniki inhabiting the subalpine belt enter
hibernation in the second half ofAt elevations of 1800 to
September.
2400 m morning activity is hardly
expressed,

whereas evening activity is

Paratypes:

ZIN 16546 a and b.

The

specimens were collected May 28, 1954;
ZIN 17545, the specimen was collected
August 6, 1955; ZIN 19935, the specimen
was collected June, 1980 by I. S. Darevsky(Fig. 11).

Holotype description: Body length is
421 mm, head included. Tail length is 46

mm. A female. Head is slightly impressed

Lateral snout edges are slightly
Anterior
pointed.
edge of snout is slightly
rounded. Rostral is narrow. Frontal isdorsally.

Vol. 3, p. 22

Asiatic Herpetological Research

April 1990

The latter is separated from the rostral by abroad scale.
Upper labials and lower
labials are both 9 to the right and 8 to theleft.
There are 5 rows of throat scales.
Around the center of the body there are 21
scale rows with strongly expressed keels,
except for two scale rows on both sides
adjacent to the ventrals, which are smooth.
The number of ventrals is 138. There are

25 pairs of subcaudals.

The color is yellowish grey.

A zig-zagshaped brown stripe runs along thedorsum. At the center of the body its width

A row of hardly
nearly 8 mm.
conspicuous spots is present laterally. The
isspots merge into a light brown stripe.
Dorsally, the head has light yellowish spots
along the edges of the frontal, parietals andlower oculars with yellowish temporals.

Ventrum is blackish marked by light
contours of ventrals (Fig. 10).
Paratypes: Morphological characters of 8paratypes are listed in the tables 1-4.
General color resembles that of theholotype, except for 2 specimens. In thelatter the dorsal stripe is interrupted in theanterior part of the body (Fig. 11).

Diagnosis: This viper is not large. SVL
reaches 460 to 489 mm. Head is slightly
impressed dorsally, covered by big scales.
Lateral edges of snout are slightly pointed.
Anterior snout edge is rounded. Nostril iscut through in lower part of the nasal.

Head is narrow, hardly separated frombody.

Remarks and variability.
FIG. 10.
19934).

Holotype of Vipera darevskii (ZINa. dorsal view, b. ventral view, c. head.

narrow. Its length is equivalent to 1.66
times its width. Parietals are slightly longer
than the frontal. The frontal is separated

from supraoculars, which protrude overlateral edge of the snout, by a row of three
Prefrontal is triangular, three times
shorter than the frontal, and is separated
from the rostral by 2 scale rows. Nostril iscut through in the lower part of the nasal.
scales.

Morphologically,

Viperadarevskii

occupies an intermediate position between
V. kaznakowi and V. ursini eriwanensis.
To be more precise, morphologically V.

darevskii occupies a middle position
between the two species of the "Vipera"
kaznakowi
complex (V. kaznakowi and
V. dinniki ), on the one hand, and the"

steppe vipers from the "V. ursini
complex, on the other hand. This viper isconsiderably smaller in body size than V.
kaznakowi. The head is narrower and thenuchal collar is less expressed. It differs

Asiatic Herpetological Research

April 1990

Vol. 3, p. 23

mrf - •**<*£*>«^T1*.

from V. ursini eriwanensis by greater headheight and much less pointed upper anterior

Coloration

Pholidosis data are listed in

T i>v» Y»£*^aa., b.

dorsal view, c, d. ventral view.

Yellow-greyish hues are prevalent.

Thepattern is more homogeneous. The necktransition is hardly expressed, like in V.
It
differs from V.
ursinieriwanensis by 1) a relatively high head, 2)yellowish general coloration, 3) clear
contrasted pattern, and 4) specialdinniki.
isyellowish or yellowish

grey, which never occurs in Vipera ursini.
Along the dorsum a contrasting zig-zag

Theshaped brown stripe is present.
ventrum is dark grey, speckled black andwhite.

.

.

FIG. 11. The paratypes of Vipera darevskii (ZIN 19935).

snout edge.
Tables 1-3.

Jr.- •>".:..'.-Of the small number of known

specimens, no melanistic individuals havebeen found. The coloration of V. darevskii
is evidently more stable than that of thepolymorphic and motley colored V.
kaznakowi and V. dinniki (Fig. 13).

pholidosis.

Sexual dimorphism. Maximum body size
is greater in females than males (Table 2).
Males have longer tails. Number of ventral
scales is greater infemales (Fig. 14).
Number of subcaudal scales is greater inmales. Sexual dimorphism in color is notrecorded.

Asiatic Herpetological ResearchVol. 3, p. 24

<fc*

'

April 1990k

FIG. 12. Vipera darevskii

Age variability.

in the field.

Newly born vipers are

uniquely colored.

invasion

of

bare

rock

Characteristic

components

outcrops.
of thesemeadows

asfollows:

Festucaare

Geographic range and ecology. The
species range shows relict characters. It

woronowii, F. ovina, Bromopsis variegata,B. villosula, and Carex tristis. Lower

covers the southeast part of the submeridian
Dzhavakhetsky Ridge within the territory of
Armenia (here this ridge has the namealong the slopes at elevations from 2300 to2400
the formations of transition

Mokrye (Wet) Mountains) and evidently
adjacent regions of Georgia (Fig. 1: 9).
Vertical distribution lies within a narrowinterval at elevations from 2600 (rarely
It is

m

on Mount Legli, Georgia.
a montane meadow subalpine species

2500) to 3000

slopes with great
inhabiting
amounts of big black outcrops of volcanic
rocks at angles of 35°-45° (Darevsky pers.
comm.). At recorded elevations, according
to Tumadzhanov(1980), subalpine
meadows (so called Transcaucasian oats
outcrops) are widely represented. It is aleading formation on steep slopes with all
exposures which form the first stages of
detritusmmontane-meadow steppe are replaced bymontane Festuca valesidea and Stipa(feather grass). From elevations of 2200 m
down to 1000 m Festuca valesiaca steppes
Here in theoccur (Lavrenko 1980).
montane-steppe highland ecological belt,
Vipera ursini eriwanensis occurs in contact
with V. darevskii in the transition zone of
the meadow steppes.

The numbers of Vipera darevskii are notgreat.

To date it is known only from thetype locality cited as "Mount Legli", where
it occurs within a narrow band in subalpine

meadows. The species biology is nearly
unknown.

Asiatic Herpetological Research

April 1990

Vol. 3, p. 25

FIG. 13. Polymorphism in Vipera dinniki from the population that occurs in the valley of the upper partof the Mzymta River.

Discussion
Phytogeny and the history of present viper
distributions in the Vipera kaznakowi
complex in the Caucasus Isthmus.

The head of the vipers assigned

toregular head plates, on the basis of skull
kinesis and analysis of the head muscles.
Without going into extensive detail on thetaxonomic position of these vipers withlarge regular head scales from the EuroSiberian group, we shall simply refer tothem as shield-headed vipers. These vipers

ursini,Vipera berus,
kaznakowi complex (Fig. 15) is covered

show great morphological resemblance.
They are share a common color pattern,with rectangular scales (the rostral, nasals,behavior, and reproductive biology.

V.

and the V.

nasal-rostrals, upper oculars, parietals, andthe frontal), unlike in V. lebetina, V.

xanthina, V. raddei, and V. persica. In thelatter species, small scales on the upper

head are typical.

Marx and Rabb (1965) consider thesecharacters important in the assessment ofphylogenetic relationships of the vipers.
On the basis of body vertebrae close
relationships and isolation of Vipera berus,
V. ursini and V. kaznakowi
within thehave been proposed
genus Vipera
andZerova 1983). They
(Chkhikvadze
restore the prior generic name of Pelias tothe vipers.
Reuss (1935) noted theisolation of the vipers that have large

We consider Vipera berus
kaznakowi

to be close relatives.

and V
These
.

vipers probably diverged from a mesophilicform in the Miocene. Northern populations

tend to be restricted to humid biotypes up to
tundra areas, whereas for southern
populations, humid subtropical forests are
In Kramer's (1961) bookinhabited.

concerning the taxonomy of V. ursini andV. kaznakowi, in the chapter on phylogeny
and history of the vipers distribution, heregards V. berus and V. ursini as a united
circle of races, a rassenkreis. Such facts
that V. ursini renardi was subspecifically
included in V. berus also emphasizes close
relationships of shield-headed vipers.

Vol. 3, p. 26

FIG. 14.

Asiatic Herpetological Research

April 1990

A female Vipera darevskii from the vicinity of Gukasyansky Region, Armenia (ZIN 19934).

Basoglu (1947) recognized the following
varieties within V. berus:

berus, renardi,

ornata. His view point was criticized andthe V. ursini renardi
was restored(Terentyev and Chernov 1949; Mertens
1952 a, b). Kramer (1961) noted greatmorphological resemblance in V. berus
and V. kaznakowi. He considered them

April 1990

Asiatic Herpetological Research

FIG. 15. The shield-headed vipers from the Caucasus,
renardi, d. V. dinnikia.

Vipera kaznakowi, b. V. dinniki,

Vol. 3, p. 27

c. V.

ursini(Lagodekhi, Georgia. ZIN 8389).

phylogenetically close species and felt thatmontane populations of the vipers played a
big role in the formation and isolation of theforms. In his opinion, when it became cold
they were forced out of the mountains onto
the plains by glaciers. When the climate
became warm again, they retreated into themountains. In the mountains it was easy tochoose temperature and humidity optima,
whereas on the plains this was notpossible. Kramer felt that in some placeson the plains where refugia might bepresent, the climate remained more or lessstable.
For the vipers in the V. ursini

complex, Kramer suggests first separationin the Miocene and Pliocene and later

independent development of the eastern andwestern forms. The complex formation of
the climate and relief in the Caucasus
coincides with regular regressions andtransgressions of the sea. Breaks in links
between the faunas of the Caucasus and theEuropean platform, the Balkans and Turkey
stipulate emergence and isolation of originalviper forms, beginning with the Miocene
and Pliocene. At this time the formation ofthe

Caucasus as a montane country
Schwarz
1938).

occurred (Bogachev(1936) suggests that the viper fauna of theMediterranean islands are Miocene relicts.
Present ranges of shield-headed vipers in

Vol. 3, p. 28

Asiatic Herpetological Research

April 1990

the Caucasus have precise altitudinal andthe plains area along the northern slope inecological limits to their distributions.
They are supported by natural historic

the Big Caucasus.

Overlap of ranges occurs inThecomparatively small areas.
"kaznakowi "like vipers evidently invaded
the Caucasus during the Miocene from thesouth, when the Caucasus island wasconnected with Middle Asian land. The
southern invasion of the Caucasus in upper
Miocene by different mammal species wasobserved by Vereschagin (1958); that oflizards from the Podarchis-Archaeolacerta
groups was noted by Darevsky (1967).
Fossils of Vipera
have been known in
since
the
Miocene(Tatarinov
Europe1964). Of the European Miocene fossils,
Provipera boettgeri Kinkelin 1892, wasdescribed. Of those from lower Pliocene,
V. gedulyi Bolkay 1913, was described by
Marx and Rabb (1965). This was at about
the time the Caucasus Mountains were
events.

A warm1938).
climatewhich
stimulatedrich
subtropicalofdevelopment
mesophilic vegetation, andcontributed to the formation and wide
distribution of warmth and mesophilic
formed (Bogachev

species like V. kaznakowi in the Caucasus,
including the Adzharo-Imeretinsky Ridge

where even until the Pliocene, bay trees,
rubber plants, araucarias and palms were
preserved (Vereschagin 1958).

mammalfossils:

Different

Middle Asian and

Caucasus hamsters, Mesocricetus,
and Talpa
Prometheomys, Sorex,(Vereschagin 1958) and also those of
insects: Orthoptera, Hemiptera, Blattoidea,
Coleoptera (Rodendorf 1939) testify to thepresence of good foodstuff for the shieldheaded vipers in the Miocene.

The end of the Tertiary period wasmarked by a decrease in tectonics and withthe emergence of a broad link between the
Caucasus and the Balkans via the Crimea
Steppes arose innorthern areas adjacent to the Black Sea
coast (Pidoplichko 1954; Scherbak 1966).
During this period an arid adapted"urisini "-like viper associated with steppeareas penetrated into western Precaucasia(Vereschagin 1958).

from theeast.
It might have alreadyfromVipera ursini renardi andseparatedhave become widely distributed throughout

The middle-upper Pliocene, when theridges of both the Big and Small Caucasus
were subjected to considerable glaciation,
should be considered the beginning of theinitial break in the range of Vipera

kaznakowi(Markov et al. 1965).

The

Kolchida (Colchis) had become a basic
nucleus in V. kaznakowi dispersal. There,
even in the epochs when it became severely
cold in the Pleistocene, a relatively warmloving vegetation of a Caucasus type waspreserved (Vereschagin 1958). Along with
Kolchida (Colchis) much smaller refugia
were sporadically preserved along theBlack Sea coast up to the town ofLazorevskoye. The same is true for thenorthern slope of the Main Caucasus Ridge

between the Pshekha and Malaya rivers.
This is supported by the present
distribution of the Tertiary vegetation of theKolchida (Colchis) type in the western
Caucasus (Adamyants 1971; Kharadze
1974; Pechorin and Lozovoy 1980;

Kholyavko et al. 1978; Tuniyev 1990, thisvolume).
It isin the narrow humid canyons with arelatively regular thermal regime that Vipera

kaznakowi is preserved.

Also, isolatedbepopulations mightpreserved in midlands
where the refugia with Kolchida (Colchis)vegetation are known in regions of the
Fisht-Oshterovsky Massive, the Lago-Naki
Plateau, and even in the Central Caucasus(Kharadze 1974; Kholyavko et al. 1978).
Small refugia might also be preserved along
the southern slope of the eastern Big
Caucasus.
Similar refugia
forArchaeolacerta were recorded by Darevsky(1967). Most highland populations of V.

kaznakowi

undoubtedly went extinct

during the glacial period. Those that were
preserved in refugia have beenDataaccumulating unique characters.
obtained by Takhtadzhan (1946) andMaruashvili (1956) reveal that in glacial
epochs, mean annual temperature neverdropped lower than 1.5-2.0°C, and theamount of precipitation was never less than
1500-2000 mm. These data support the
proposed preservation of relictual V.

Asiatic Herpetological Research

April 1990

kaznakowi populations in the mountains.

Vol. 3, p. 29

Mzymta River head, all the initial types ofcan be found (Fig. 15).
InofV.
kaznakowi
premontane populations
we have not found specimens which havehybrid characters.
Generally allof
V. kaznakowi
premontane populations
show great homogeneity. In the eastern
vipersDarevsky (1967)

regards this argument

as proof that due to the development ofmontane glaciers, a fundamental
reconstruction of all animals and plants inthese areas occured from premontane andmontane refugia. Lizards for example,
might have been preserved in the
Gagrinsky and Bzybsky ridges facing thesea, along with other regions. The end of
the Pleistocene was marked by thedevelopment of steppe landscapes on the
Zakubauskaya sloping plain along withalready formed landscapes and acharacteristic biological community of thepresent type along the Black Sea coast of
the Caucasus (Vereschagin 1958). Thissituation contributed to a much wider
dispersal of Vipera ursini renardi andfurther preservation of V. kaznakowi.

During the xerothermal epoch in the
Holocene, intensive glacial melting and the
upwards elevational shift of vegetation belts
along the mountain slopes occurred
throughout the area. Within a considerable
area of the Caucasus, steppe landscapes
became prevalent. This allowed the steppeanimals, including the steppe viper, Vipera
ursini renardi, to ascend into the mountains(Vereschagin 1958). At that time an initial
contact of V. ursini renardi with montane

kaznakowi might
have occurred in the midmountains of theWestern Caucasus.
This region is
inhabitedV.
dinniki which
presently
byshows mixed morphologies of both V.
kaznakowi and V. ursini renardi. The

relict populations of V.

approximate ways of possible interaction
and characters are shown in Fig. 16.

Darevsky (1967) writes about the formbuilding role of hybridization in the
Caucasus. He observes the genesis of newforms in the lizard group Archaeolacerta.
Hybridogenity in Vipera dinniki isproblematic,

whereasintrogressing
hybridization in the zones of contact

between V. kaznakowi and V. dinniki

ispresently not doubted, age interbreedings

apparently

occurringin

midmontane

populations of V. dinniki. For example, in
one of these populations associated with therange of V. dinniki the species is presently
disjunct geographically from the closely
related V. ursini renardi,
but rarewith an intermediatemorphology between these species areindividuals

present.

Mayr (1968), Borkin and Darevsky(1980) and Solbrig and Solbrig (1982)
report on various types of hybridization.
For instance, it is recorded that hybrids
possessing vital capacity occur between
sympatric species. Some of these hybrids
are capable of recurrent crosses with one orboth parental forms. The fate of both
species between which hybridization occurs
is partially dependent on 1) the intensity ofhybridization occurring between them andthe level of genetic and ecological2)sterilityof hybrids (Solbrig and Solbrig1982).
In most cases the hybrids themselves do
not have greater evolutionary significance.
Rather it is the products of recurrent
interbreeding between the hybrids and theparental forms, i.e. the process of

introgressive hybridization, that may form
new taxa; i.e. the blending of genomes ofancestral taxons, or "borrowing" of parts

from

other

genomesbymeans ofintrogression of genes as a variant ofhybridogenic species formation (Mayr
1968; Borkin and Darevsky 1980).

After the xerothermal epoch, the climate
This factagain became more humid.
contributed to the restoration of the former
limits of the forest belt (Vereschagin 1958).

Throughout the subalpine belt along thesouthern slope of the Main Caucasus Ridge
within the area from the Central Caucasus
up to the Fisht-Oshtenovsky Massive in thewest, subalpine meadows and montane
curved forests are widely developed(Galushko 1974; Dolukhanov 1974;
Kharadze 1974; Kholyavko et al. 1978).

Vol. 3, p. 30

Asiatic Herpetological Research

V. ursini eriwanensis

V. darevskii

I
FIG. 16. Model of the proposed genesis of the vipers from the Vipera kaznakowi complex.

April 1990

Asiatic Herpetological Research

April 1990

In the western part, this vegetation ispresent along the northern slope. To theeast, it gradually changes to steppe(Lavrenko 1980). Definite limits of thesubalpine belt in the area influenced by thewarm Black Sea mark the present
distributional limit of Vipera dinniki.

The final establishment of the present
climate stabilized the range of Vipera
kaznakowi and allowed the joining of
scattered populations from the refugia
within the given region. The favorable
conditions along the Black Sea coast madeitpossible for V.

kaznakowi

to disperse

along warm river valleys up
midmountains and for V. dinniki

totodisperse along the northern slope of the Big

species, V. kaznakowi

Vol. 3, p. 31

and V. darevskii.

In middle Pliocene, Vipera kaznakowi
apparently invaded the Dzhavakhetsky

Ridge (the Mokrye [Wet] Mountains). At
that time the hot, arid climate of the region

was replaced by a more humid, coolclimate.
Later on, however, beginningwith the middle Pleistocene, processes ofaridization and glaciation caused the final
degradation of forests (Agakhanyants
1981). By that time the diverged vipers
were restricted to the highest places in themountains where maximum humidity could
be found. At the same time the newly
formed steppe areas were actively invaded
by V. ursini eriwanensis (Reuss 1935)
from the Anatolyskoye Plateau. Later onviper became widely distributed
throughout the entire montane-steppe belt in

Caucasus to the east up to the Belshaya
Laba River head. Where V. kaznakowi

theand V. dinniki

distributions meet, andthose of V. dinniki and V. ursini renardi,

the Small Caucasus and the Armyanskoyezones of secondary hybridization emerged.
In these zones the following is observed:
intergradation of characters, a high degreeof polymorphism, and the "plucking" ofhybridization, including further formbuilding, evidently existed. The genesis of
V. darevskii
can be explained by thehybridization of V. ursini eriwanensis andspecimens from the "initial" type. In these
populations the specimens are hard to
define.

that split from its major
V. kaznakowi
range in the vicinity of the Mokrye
Mountains (Plate 3).

Phenotypical diversity is great withregard to the Mzymta River valley
Non-identified individuals(Fig. 15).
having characters of both Vipera ursini
renardi and V. kaznakowi are known froma number of localities in the complex

Apparently, introgression of genes from
Vipera ursini eriwanensis which essentiallysurrounded a small Pleistocene populationof V. kaznakowi was very substantial.
Morphological propinquity of V. darevskii
to V. ursini eriwanensis and V. kaznakowi

eastern range of these vipers.

We think that

history of V. darevskii
connected with the invasion
the

is

also

of

V

.

kaznakowi from the south during the
Miocene and its further wide dispersal
throughout the Caucasus.
Viperakaznakowi apparently used to inhabit the
majority of the Maly (Small) Caucasus
area, including the Adzharo-Imeretinsky,

Meskhetsky, and Dzhavakhetsky ridges.
The fact that this range did exist in the pastis demonstrated by a relict

population of V.

kaznakowi that occurs in Baniskhevsky
Canyon, Georgia and by the proposed
distribution of the vipers along the
Trialetsky Ridge (Bakradze 1969). The
latter might be a link between the areas
presently inhabited by the closely related

Plateau.

At that site one more source of

support its possible hybrid genesis (Borkin

and Darevsky 1980).

It

goes without

saying that further ecological, ethological,
cytological and biochemical research ofthese vipers is needed to prove it.
Similarly, Agakhanyants (1981) explains
the floristic richness of the region by acomplex interaction of invaders from thesouth and north in the highlands of the
Small Caucasus. Further aridization of theclimate caused the mesophilic form of V.
darevskii to ascend into the mountains.
Ecological disconnection in the present
high-altitude belts caused complete
separation of V. darevskii and V. ursini
eriwanensis.

The presence of "kaznakowi

"

-like

Asiatic Herpetological ResearchVol. 3, p. 32

vipers in Lagodekhi and other regions of
the Big Caucasus may be accounted forsolely by changes in climate in refugia that

populations and further
divergence in isolated montane canyons. A
number of isolated populations might bymodified due to hybridization with Vipera
ursini renardi in a manner similar to theabove listed scheme.

preserved

During the period of xerophytization in
the Holocene, breaks within the range ofVipera kaznakowi occurred. At that time
isolated populations existed over a long
period.

A number of populations inhydrophilous epochs to follow, would havebeen capable of connecting many times.
This is supported by the restoration of theformer forest belt after Holocene
xerophytization (Galushko 1974; Kharadze1974). Pleistocene glaciation obviously
also influenced the emergence of isolated
populations and changes in the
distributional limits (Markov et al. 1965).

assistance

of

many

(In Russian).

ANONYMOUS.

[Red Data Book of the
Lesnaya Promyshlennost Publishing,

USSR].
Moscow.

1978.

(In Russian).

ANONYMOUS.

1984. [Red Data Book of the
USSR]. Lesnaya Promyshlennost Publishing,
Moscow. (In Russian).

BAKRADZE, M. A.

1969. [New data on the rangeof the Caucasus viper {Vipera kaznakowi
Academic Press,
Nikolsky) in Georgia].
Tbilisi, Georgia 56(2):467-468. (In Russian).

BAKRADZE, M. A.

1975. [Distribution of Vipera
laurenti in south Georgia]. Bulletin of the State

Museum of Georgia, Tbilisi 28a:41 1-412.

(In

Russian).

BANNIKOV, A. G. AND V. M. MAKEYEV.

1976.

[Conservation of venomous snakes of the

USSR]. Priroda Publishing, Moscow 5:71-83.
(In Russian).

BANNIKOV, A. G.,

Acknowledgments
The

Mysl Publishing, Moscow.

of the USSR].

270 pp.

April 1990

I.

S.

DAREVESKY, V. G.

ISCHENKO, A. K. RUSTAMOV, AND N. N.
SCHERBAK. 1977. [Field guide of the USSR
peoplecontributed to the completion of this study.

We would like to extend our special
recognition to II ya S. Darevsky and Natalia
B. Ananjeva from the Leningrad Zoological
Institute, USSR Academy of Sciences who
aided us in the collection of data, analysisof the material, and preparation of thisare grateful to Valentina F.
manuscript
theOrlova from
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