I

Vol. 6, pp. 1-26

Asiatic Herpetological Research

June 1995

Systematics of the Vipers of the Caucasus:

Polymorphism or SiblingSpecies?

GORAN NILSON BORIS S. TUNIYEV 2 NIKOLAI ORLOV^, MATS HOGGREN4 AND CLAES
1,,

ANDREN 1'Department of Zoology, Goteborg University, Sweden
2
Caucasian State Biosphere Reserve, Sochi, Russia^Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia^Department of Genetics, University of Uppsala, Russia
Abstract. -Inter- and intramorphological variation were examined in sympatric and allopatric polymorphic
and monomorphic populations of the Vipera ursinii and Vipera kaznakovi complexes. The alpine Vipera
dinniki populations in upper Great Caucasus show a pronounced, and to a certain extent geographical,

Color patterns include among others 'kaznakovi', 'tigrina', 'berus', 'bronze', and 'ursinii'
litter in certain populations.
Viperatypes. Several of these patterns can be represented within the sametheursiniioftheCaucasianwithisdinnikicomplex in some areas. This lastViperarepresentative
sympatric

polymorphism.

taxon shows a similar degree of polymorphism, which is unique for this complex, and due to morphologicaland molecular distinction, we consider it to be a Caucasian evolutionary species within the ursinii complex

Vipera lotievi sp.n.

Key Words:

V. renardi, V. lotievi
Reptilia, Squamata, Viperidae, Vipera dinniki, V. kaznakovi, V. ursinii,

sp.n., Caucasus, Russia, Georgia, taxonomy, morphology, polymorphism.

Introductionsynonym of V. kaznakovi until Vedmederja
et al(1986) recognizeditas a separate

The taxonomy of the vipers of Caucasus
has for a long time been confusing andspecies inhabiting alpine and subalpine
meadows in the Caucasus, and thus

contradictory. According to the traditionalview a single species, Vipera kaznakovi, isrestricting V. kaznakovi to lower altitudes in

distributed in the moist and warm lowlandsof the western Caucasus as well as in themountain valleys towards the east. In theeast the habitat is drier and along the range
the vipers gradually change toward Vipera
ursinii in appearance. In the east Caucasus
only this last viper was

supposed to occur.

western Caucasus and adjacent moist
lowland habitats along the eastern Black Sea

Thereby the problem of the gradualtransformation from V. kaznakovi in thewest of the Caucasus to V. ursinii in the eastisrestricted to the high Caucasus

coast.

populations, now including V. dinniki and
V. ursinii. Vipera kaznakovi is well definedandThus there seemed to be a somewhat clinalandtransformation from "pure" V. kaznakovi inthe west to "pure" V. ursinii in the east.
Vipers from the intermediate region could be
Within the samedifficult to determine.
locality some specimens look like V.
kaznakovi, other ones are more like V.

geographically separated from all the otherursinii, while still some can be intermediate.

Nikolsky (1913) separated the alpine
populations into the taxon Vipera berus
dinniki, which was based on alpine
specimens of the conventional V. kaznakovi
from high altitudes in the western Caucasus(Malaya Laba River- terra typica restricta
and Svanetia) as well as from other places.
The name dinniki was long considered as a

© 1995 by Asiatic Herpetological Research

restrictedindistribution,

viper species in the region.

The complex history of nomenclatureand taxonomy has been clarified to a greatpart in some recent publications (see Orlov& Tuniyev, 1986; 1990), together withhypotheses about the phylogeny of thisgroup. Concurrently the need of genetic
studies was stressed, and this has led to thepresent work where we in a series of papers
intend to clarify the taxonomy and evolutionof the vipers of this region.

The work is

broad perspectiveplannedincluding phenetic and phylogenetic
analyses, habitat choice, niche-breadth, and
to

have

a

Asiatic Herpetological ResearchVol. 6, p. 2

Different methods takereproduction.
different time and in this first paper themorphology

isreexamined,

based onavailable material in Museum collections andfreshly collected material from a number of
new places, as well as on inheritance ofcolor pattern.

Genetic structures based onphenetic analyses of allozyme data are alsopresented.

The morphological distinction between
Vipera dinniki and V. kaznakovi has beenpresented elsewhere (Orlov and Tuniyev,
1986; 1990) and will not be repeated in thisstudy. In the present paper we are focusing
on patterns of morphological and molecularvariation within and between the different
populations in the Caucasus, and thetaxonomy of these populations.

June 1995

Altogether about 300 preserved or live
specimens of vipers from the Caucasus andadjacent regions have been seen during thestudy.

Joint field trips

were madeindifferent parts of the Caucasus in1990 and
1992, but two of us (Tuniyev and Orlov)
have performed extensive research in theregion prior to that. For morphometricstudies 183 preserved snakes within theursinii and kaznakovi complexes have been
examined more carefully, and for most ofthese specimens 30 different items of data
have been collected. This information was

down

population level, inmorphological descriptions, taxonomical
andconclusions
about
analyseszoogeography and range overlap.
used,

to

Inheritance of color pattern

was studied

based on 23 pregnant females and theiroffspring.

Material and Methods

The work has mainly been a study ofvariation in external morphology, allozymes

and reproduction in order to reveal patternsof sympatry and sibling species. Additional
genetic studies will follow when material
becomes available in suitable samples(presently delayed due to political reasons).
Thus live and preserved museum material
has been examined concordantly withstudies of reproduction in the laboratory.

Additional preserved material used inthis study originates from the Natural

Museum in Goteborg (GNM);
Dipl.-Biol. F. J. Obst, Staatliches Museum
History

fur Naturkunde, Dresden(MTKD D); Aram

Institute, Academy ofAgasian, Zoological
Sciences, Eriwan, Armenia. Abbreviations
for museums as used in the text are: CNRCaucasian State Biosphere Reserve,Collection of Boris Tuniyev at Yew-box
Groove, Sochi; GNM- Goteborg Natural

History

Museum, Goteborg; MTKD-

Staatliches

Museumfor

Naturkunde,

ZIEr- Zoological Institute,
Academy of Sciences, Eriwan; ZIGDepartment of Zoology, University of
Goteborg; Goteborg- (authors' collection,
which later will be incorporated in GNM);
ZIN- Zoological Institute, Academy ofDresden;Sciences, St. Petersburg.

Data collected were: total length and taillength; number of preventrals, ventrals,
subcaudals, anterior and mid-body dorsal
scale rows, apical plates, supralabials,
sublabials, circumocular scales, loreals,
second chinshields, mentals, crown scales(=intercanthals+ intersupraoculars), andzig-zag windings in dorsal band.

Furtherrostral index (height/breadth) and head index(breadth/length) were calculated. Divisionof parietals, frontal, and nasalia was noted,
as was the color of dorsal and ventral sides,

and iris (in live specimens). Further, upper
preocular size; and head, labial and lateralbody patterns, as well as distinctiveness ofcanthus rostralis were examined. Details
about these methods are found in Nilson and

Andren (1986).
Morphologylphenetics

Standard errors accompanying mean
were used as relative
measurement of dispersion.
For theanalysis of intra- and interpopulational

character ratios

morphological variation (phenetic analysis)
the samples were divided into subsamples
Thus
depending on questions raised.
besides an analysis of morphologicalvariation also a pattern confirming or
rejecting the present taxonomic pattern could
be achieved. This pattern could also beAsiatic Herpetological Research

June 1995

Vol. 6, p. 3

TABLE 1 Number of specimens used in the genetic analyses and localities (Russia if nothing else is stated).

for the examined taxa.

kaznakovi:
1.

2.

Dagomys, north of Sochi. Six specimens.
Rudorova, inland locality, 900 m alt. Four specimens.

dinniki:

3.

4.

Fisht/Oshten, the westernmost locality of the main Caucasus range. Seven specimens.
Lake Impsi, 1,980 m. alt., at a tributary to the Little Laba River on the northern slopeof the main range. Seven specimens.
Aishkha-II on the southern slope of the main range. Three specimens.
5
6. Lake Kardyvach at upper Mzymta River on the southern slope of the main range
Seventeen specimens.
.

lotievii:

7.

Armkhi, Checheno-Ingushetia. 2,000 m altitude. Seven specimens.

berus:
8.

Uppsala (terra typica), Sweden. Eleven specimens.

eriwanensis:

pooled sample from Asbua and Cildir, Kars, east Turkey; and Sevan, Armenia. Six
specimens.

9.

supported or rejected by the parallel
biochemical studies. Thereby it is possible
to state or reject the occurrence ofthe eight Caucasus populations
studied (see Table 1, for locality data and

among

A

convergent or parallel evolution, i.e. sibling

sample sizes).
potential risk of sampling
error due to syntopic occurrence of two taxa

species.

may

Estimation of the different color patternfrequencies in local populations was basedon observations during the field work.
Small museum samples collected by others
were not included in this analysis due touncertainty of randomness in sampling(unusual morphs might have been collected
and preserved at a higher degree).

Biochemical data

Enzyme electrophoresis.

—

Sixty-eightspecimens representing different taxa of
Caucasus vipers, and Vipera berus frombeavoided by testing observed

genotype distribution within a locality
against Hardy-Weinberg expectations (seeresults).

Fresh or frozen tissues (-75°C)

from liver and skeletal muscle were
homogenized in distilled water. The extracts
were centrifuged for 10 min at 10,000 rpm
and 4°C and the supernatants were thenStandard
stored at -75°C until used.
horizontal starch gel electrophoresis was
carried out, as described by Harris andHopkinson (1976) and Murphy et al.
Gels (11% w/v) were prepared(1990).
from Sigma starch (Sigma Chemical Co.,
St. Louis, Mo). Two buffer systems weretreated as nine independent operationaltaxonomic units (OTUs) in the genetic
analysis, in order to avoid a priori

M tris-0.005 M citric
M lithium hydroxide0.03 M boric acid, pH 8.0 (Ridgway et1970). (B) Gel: 0.002 M citric acid, pH
6.1; Electrode: 0.04 M citric acid, pH

assumptions of taxonomic relationships

adjusted with N-(3-amino-propyl)morpholin

Sweden were examined. The samples wereused: (A) Gel: 0.03

acid; Electrode: 0.06

al.,

Vol. 6, p. 4

TABLE 2.

Asiatic Herpetological Research

June 1995

Enzymes and electrophoretic conditions of the polymorphic loci scored in this study.
Nomenclature and commission numbers following the International Union of Biochemistry, Nomenclature
Committee (1984). Abbreviations for tissue sources are: L=liver and M= skeletal muscle.
EnzymeFIG..

1.

Vol. 6, p. 5

Asiatic Herpetological Research

June 1995

The typical "dinniki" pattern type of Vipera dinniki, with unicolored lateral sides typical for

Fisht/Oshten (ZIG).

FIG. 2. The "dinniki" pattern type of Vipera dinniki, with a tendency towards the "tigrina" morph. From
Kardyvach (ZIG).

River.

Results

Also situated on the southern slopeof the main range.

—

at 1980 m. alt., situated at
Lake Impsi
the Tsahvoa River, a tributary of the Little(Malaya) Laba River. The locality is mainlyon the slopes of the Damhorts Range at thenorthern part of the Reserve, and partly on
Akaragvarta Mountain situated on thenorthern slope of the main range.

Theof the analyses ofandenzyme electrophoresis
morphometries
are presented separately.
Intra- and interpopulational variation inmorphology

A great number of different color
morphsIn addition much information on Vipera

kaznakovi was gathered at the lowland
Black Sea coast localities of Dagomys, northof Sochi (Russia) and Hopa, Artvin
Province (Turkey).

resultsare expressed in the Caucasian

Although several stages ofvipers.
overlapping and intermediate forms could beseen, we define the following major patterntypes:

Nilson et al.

Plate

Asiatic Herpetological Researcha.

The "ursinii" pattern type of Vipera dinniki, from Kardyvach (ZIG).

b.

The "tigrina" morph of Vipera dinniki, from Impsi with partly divided transverse bars

(ZIG).

1

Nilson et al.

Plate 1

Asiatic Herpetological Researchc.

The "bronze" morph of Vipera dinniki, from Impsi (ZIG).

d.

The "bronze" morph of Vipera lotievi from Itum Kali, Checheno-Ingushetia.

Vol. 6, p. 6

Asiatic Herpetological Research

June 1995

FIG. 3. The "kaznakovi" morph of Vipera dinniki, from Fisht/Oshten (Mt.Oshten - Armenian pass).

A. "dinniki": a more or less continuousand pronounced lateralblotches. Sometimes nebulous in pattern.
Sometimes rather Vipera berus like (Figs.

D.

"tigrina":

dorsal pattern fragmented

zig-zag band,

into broad or

1,2).

Also spotted pattern could be seen in some
populations. This pattern type is most close
to "tigrina", but differ by being divided
along the vertebral line thus resulting in two
rows of dark spots along the dorsal side ofB.

"kaznakovi": pronounced blackand dorsal longitudinal bands, andyellow or orange ground color. The dorsal
band is waving or expressed as a straightband resulting in a contrasting more or lessnarrow transverse bands.

lateralthe body (Fig. 6, Plate lb).

bilineated pattern. Besides these we could
define a "bilineated" morph which is similarE.
"bronze":a uniform greyish tobrownish or blackish ground color coveringallparts of body except the head.
Sometimes with a darker narrow or broadto the "kaznakovi" type of pattern but muchvertebral stripe (Figs. 7, 8, Plate lc).

lighter. In the analysis below it is includedin the "kaznakovi" type (Fig. 3, compare

Fig. 4).

F.
"melanism": black, with a highproduction of melanin covering all othercolor patterns.
C.

"ursinii":

a black-edged continuous

dark brown zig-zag band on a paler groundcolor and lighter sides of body (Fig. 5, Platela).

The different morphs were represented
in different frequencies at the different
localities examined, and some morphs

seemed to be restricted to one or a few

Asiatic Herpetological Research

June 1995

FIG. 4.

Vol. 6, p. 7

A typical Vlpera kaznakovi from the inland locality cordon Babuk-Aul at the foothills of Mt. Fisht(ZIG).

localities (Table 3). Also variation in othercolor characteristics was obvious whencomparing populations. The number ofwindings in the dorsal zig-zag band variedmarkedly with low number in the western

isolated Fisht population and high in the
more eastern populations (Fig. 9). This was

especially pronounced when comparing
Fisht with Kardyvach, characterized by ahigh frequency of "tigrina" morphs (Table
3). Almost no overlap was detected as in
Kardyvach the vipers have 68 or more
windings while in the Fisht population thecorresponding figures are 69 or less (Table
In general the west Caucasian samples(except Fisht) have higher number of dorsal
windings or transverse bars than central andeast Caucasian dinniki vipers. Also V.
4).

FIG.

The

pattern type of Viper adinniki, with more unicolored lateral sides from Mt.
5.

Fisht (ZIG).

"ursinii"

kaznakovi has fewer windings.

Vol. 6, p. 8

Asiatic Herpetological Research

June 1995

FIG. 7. The "bronze" morph of Vipera dinniki, from Mt. Loyub (ZIG).

TABLE 3.

Distribution of color morphs in samples of V. dinniki and V. kaznakovi (Dagomys) in absolute
numbers and in percentage (in some cases) in 1991. Sample size in parenthesis.

June 1995

Vol. 6, p. 9

Asiatic Herpetological Research

FIG. 8.

The "bronze" morph of Vipera dinniki,from Mt. Loyub (ZIG).

Dorsal bars

FIG. 6. The narrow-banded "ligrina" morph of
Vipera dinniki, from Kardyvach (ZIG).

Of the juveniles taken together half(50%) were "tigrina". When considering
Kardyvach material alone (10 broods

thewith 29 juveniles) 52% were "tigrina" while
24% were "bronze". Of 47 adults observedin the field in July 1991 at this locality 24%
The number of adult
were "bronze".
observed(33%) was lower than the"tigrina"

frequency of juvenile "tigrina" produced
while the number of adult "ursinii" wasrather high (27%). This slight reduction ofthe "tigrina" pattern between juveniles andadults can have an ontogenetic explanation

FIG. 9.

Distribution of number of bars and/or

windings in the dorsal zig-zag band in the different
Vipera dinniki populations, running from west
towards east in high Caucasus (1: Fisht, 2: Loyub,
3:
Kardyvach, 4: Impsi, 5: "central Caucasus"(=Elbrus and surroundings), 6: "east Caucasus"(=mountains above Lagodechi, Georgia), and V.
kaznakovi populations (7: Sochi-Adler, 8: Hopa,

as pattern often fades with age.

Artwin).

In other populations examined the
combinations of morphs were different,

At Impsi the typical "dinniki" pattern waswith other patterns dominating. The numberof windings is much lower in Fisht
compared to the Kardyvach sample (with"tigrina" predominating). "Melanism" wasonly observed at Fisht, but it is known also
from Aishkha-II and the Bezymyanka River.
"Bronze" was only observed at Kardyvach

andImpsi.

When comparing

field

observations and all available specimens incollections, this morph was not documented

from any other locality along the entire area.

dominating (56%).
Analysis ofscalation characters in the
different dinniki populations

Geographic variationin colormorph

also reflected in scalation
frequencies
characters (Table 4). Going from the west
is

Caucasus towards

east, certain changes
could be observed. The central and east
Caucasus samples of dinniki have higher
mean number of preventral plates, lower

Vol. 6, p. 10

TABLE 4.

Asiatic Herpetological Research

June 1995

Variation in scalation and color pattern characters between different isolated populations of Vipera

The localities include a series of isolates at close distance in the Caucasus State Biosphere Reserve (1:Mt. Loyub; 3: Lake Kardyvach; 4: Lake Impsi), and 5: central Caucasian population
and
6: east Caucasian population (the mountain region of Lagodekhi) sample. Given as
Elbrus(Mt
region)dinnikiMt Fisht/Oshten; 2:

Mean value, S.E. and range (except for preventrals and apicals). Number of specimens in parentheses.

June 1995

Asiatic Herpetological ResearchVol. 6, p.

Crownscales 25
-

-

o
o
o
o
o
o
o
o
o

o
o
o
o
o
o

o
o

o
o
o
o
o
o
o
o
o
o

FIG. 10.

o
o
o
o
o

o
o

•

o
o
o
o
o

o

o
o
o

o
o
o

o
o
o

o
o
o

-

o
o

o
o
o

-

o
o

Distribution of ventral numbers in the

different Vipera dinniki populations, running fromwest towards east in high Caucasus (1:Fisht, 2:

Loyub, 3: Kardyvach, 4: Impsi, 5: "central
Caucasus" (=Elbrus and surroundings), 6: "eastCaucasus" (=mountains above Lagodechi, Georgia),
and V. kaznakovi populations (7: Sochi-Adler, 8:

Hopa, Artwin).

head index

iQO

rostral index

FIG. 12. Head index (breadth/length) and rostralindex (height/ breadth) in sympatric sibling speciesof east Caucasus. White circles= V. dinniki ("eastdinniki"); black squares= V. lotievi.

mean number of apical scales, and lower
number of loreal scales. Also the number ofventrals showed a slight decrease towards
the east, a pattern also observed between
northwestern and southern populations ofVipera kaznakovi (Fig. 10).
Analysis of scalation characters in the
different dinniki morphs
In subalpine and alpine mountain belts of thewest Caucasus (eastward to the basin of the

Big Laba River) the "ursinii" morph belongs
to the same species as the rest of the

1 1

Vol. 6, p. 12

Asiatic Herpetological Research

June 1995

TABLE 6. Variation in scalation and color pattern characters between the northern and southern Vipera
kaznakovi populations. The samples are from northeastern Black Sea regions in Russia (Sochi-Adler); andfrom northeastern Turkish Anatolia (Hopa, Artvin province). Given as Mean value, S.E. and range (except
for preventrals and apicals). Number of specimens in parentheses.
Sochi-Adler (17)Ventrals

1.4L+0.19
133.8+0.6, 130-138

Rostral index

1.1±0.02, 1.0-1.27

Preventrals

Apicals

Circumoculars*
Loreals*

Crown scales
Zig-zag windings***

Hopa (13)1.54±0.18
130.4±0.9, 124-136
1.1±0.06, 0.87-1.5

1.50±0.13

1.64±0.14

20.0±0.48, 16-23
11.06±0.76, 7-16

19.31±0.43, 15-21
8.69±0.60, 5-12

14.9410.92, 10-23
56.33±0.67, 55-57

17.33+1.08, 11-22

50.75+1.55, 48-55

Counted as sum of both sides.
completely bilineate and melanistic specimens are excluded.

** Four

TABLE 7.

Allele frequencies of polymorphic loci (see Table 2 for locus abbreviations).

abbreviations, sample sizes and localities, see Table 1.

Locus

For taxon

Asiatic Herpetological Research

June 1995

Vol. 6, p. 13

TABLE 8. Genetic distances (above diagonal) and genetic identities (below diagonal) of eight OTUs of
Caucasus vipers and Vipera berus.

1- kaznakovi,

Dagomys; 2- kaznakovi, Rudorova; 3- dinniki, Fisht; 4-dinniki, Impsi; 5- dinniki, Aishka; 6- dinniki, Kardyvach; 7- lotievii; 8- berus; 9- eriwanensis.

OTU

1Asiatic Herpetological ResearchVol. 6, p. 14

II and Impsi) belong to one polymorphic
taxonomic unit (V. dinniki).

Discussion

The

original

question,whetherpolymorphism or sibling species are theJune 1995

The warm and wet adapted Vipera
kaznakovi is distributed in this area today,
and it has been considered that this species
has had an occurrence in this region for along time (Orlov and Tuniyev, 1990;
Tuniyev, 1990) due to the climatic stability.

A scenario for the arise of various

prevailing phenomenon in the Caucasian
populations of vipers, must be answeredwith yes in both cases. In the western
Caucasus a large number of morphs can bepolymorphic V. dinniki populations could
have taken place in two steps and been like

separate six different ones
recognized.
in that region while the number is restricted
to three in the eastern and central parts of themountain range (Fig. 14). As seen in the
reproductive studies all six morphs

If the Pliocene Colchis region decreased in

this:

We

1.

First, the evolution of Vipera dinniki.

range during a Pleistocene glacial period
there are two alternatives for the species in("melanism", "bilineat-kaznakovi", typicalthat part of the old range that have turned"dinniki" (including "nebulosa"), "tigrina","bronze", and "ursinii") in the mountainhabitats in the western Caucasus belong to acold:

This is also verified bysingle species.
cladistic analyses of biochemical andforced down to the remaining warm zone.
In both these two cases the net result will bea reduced range for the single species

morphological data (Nilson et al., 1994).
The different morphs can be seen sympatricand syntopic in suitable rocky, vegetationrich, and moist habitats. There seems to bea certain inheritance pattern, but in principle
most morphs can occur in the same brood(Table 5).

the viper could disappear or it could

adapt. If it disappears it can be done in two
ways: the snake becomes extinct or it is(kaznakovi) in the remaining warm and wet
Colchis refuge. In the second case, if theviper is adapted to the new climate, "the coldzone", it would be a new, physiologicallydifferent race.

Certain areas seem to have acertain range of frequency of the various
morphs, which might be unique for thatmust be kept in mind that
only a random number of populations havebeen investigated, and it is likely that
additional morphs will be described fromparticular area.

Itother isolated localities.

The isolation of thedifferent mountains in the Caucasus is muchcomparable with e.g. the Andes of Ecuador,or the Galapagos islands, and a high degreeof isolation has obviously taken place
between the different mountain peaks.

How could this particular polymorphic
pattern in the subalpine Vipera dinniki
populations have evolved? The present
warm and wet subtropical Colchis area ofwestern Transcaucasia at the Black Sea coast
has, as indicated by botanical evidence,
served as a refuge for animals and plants

the whole of Pliocene and
The total
Pleistocene (Tuniyev, 1990).
region has varied much in size during the
Pleistocene glacial and interglacial periods
but remained a permanent relict key area.

In an interglacial period when the climate

becomes warmer again the warm and wet
Colchis zone expands and "the cold zone" isforced upwards to higher altitudes in themountains. Now, if there is a cold-adapted
physiological race, again two alternatives are
open: first, it could adapt, or second, it
could disappear. If it is adapted back to the

new warm environment, it can either goback into and unite with the old species(kaznakovi) which under the new climaticconditions can expand its range, or it could
form a sympatric but physiological distinct
In the case of sympatry there is agood possibility that it would disappear due

taxon.

to competition (if not ecological distinct). In

both these last situations there is a greatthat the original species(kaznakovi) would return and again cover its

probability

original range.

duringIf the new physiological race disappears

from the region it could again be done in
two ways: it simply becomes extinct due tothe new "severe" climatic conditions, or itJune 1995

Vol.6, p. 15

Asiatic Herpetological Researchmigrates upwards following the pushed up"cold zone". In the first case again only the

The present geographical distribution
and morphological pattern of Vipera dinniki,

original species (kaznakovi) will remain in
its new expanded (=original) warm and wet
range. In the second case there will be aas well as the fact that in all subalpine
mountain regions where V. dinniki isnumber of cold-adapted populationsathigher altitudes (the present different
dinniki, "east-dinniki", darevskii
populations). This would mean a number ofisolated populations as the mountains of the
Caucasus are rather steep andsubalpine/alpine habitats are in many cases

Further the
climatologically isolated.
isvery complex reflecting a highgeologyof
local
endemism among plants anddegreeanimals. One can postulate that the different
populations, or groups of populations, musthave become adapted to local conditions.
2. This may have been the prerequisite
for an evolution of a polymorphic Vipera
dinniki.
During Pleistocene there wereseveral glacial and interglacial periods, andthe scenario postulated above would havebeen repeated several times, and during eachand its cold
withforceddownwards
wereadapted viper

interglacial the "cold zone"

the result of a secondary contact with

Unique morphs

neighboring populations.
could by this be spreadtoadjacent

populations etc.

Today we can see a polymorphic colorpattern in the cold adapted Vipera dinniki
that to a high degree is unique for one or asmall

number of populations

in

close

connection, e.g. the "bronze" and "tigrina"
morphs, but not seen in all populations.
Vipera dinniki could be a polymorphic
species, constituent of a number oflocated today, there also are fragmented
more or less subtropical Colchian refugia at
lower altitudes, inhabited by V. kaznakovi
This supports the(Tuniyev, 1990).
evolutionary pattern postulated above.

Further east in the mountains, thehabitats get drier with moist areas restricted
to stream surroundings and lake shores. In
central Caucasus (Mt. Elbrus) the number ofmorphs decreases

to

two

("ursinii"

and"dinniki") or three in the eastern Caucasus(where again a form of "bronze" morph

appears) (Fig. 14).

Now, in the region of the central andeastern Caucasus two different sympatricAs shown in thespecies are involved.
results section above, at several localities thetwo morphs "dinniki" and "ursinii" actually
represents two sympatric species from the
kaznakovi line and the ursinii line
respectively. It is obvious that these two
species are sympatric (but not necessarily
syntopic) in a large number of places inthese eastern and central parts of the main

range of the Caucasus. We have in ourmaterial such records of sympatry from Mt.
Elbrus in the central Caucasus (Figs. 1518); mountains north of Lagodechi in theeastern Caucasus; at Itum- Kali, ChechenoIngushetia (Fig. 19); and various records
from Dagestan, besides several isolated
records of both species from the entire
eastern and central Caucasus range.

the relative genetic differentiation between

Vipers of the kaznakovi group are
known from subalpine meadows, andsnakes of the ursinii group have been foundin the semiarid hollows between the main
range and the Skalisty (Rocky) range. At
several places with connection of subalpine
meadows and semiarid hollows the two
species have a sympatric occurrence (andsyntopic along the ecotones of both types of

examined taxa and degree of genetic

landscapes).

populations that during periods are isolated
from each other, but irregularly have
In some cases thesecondary contact.
isolation could also have been morepermanent resulting in a number of sister
species along the range, a possible

phylogenetic pattern we currently are trying
to solve with genetic studies. But overall,polymorphism were low, indicating a rather
recent divergence.

The ursinii line has probably
never been widely represented in theperpetually humid western Caucasus, as thistaxon is adapted to dry environments, but asstated earlier, in the extreme eastern part ofVol. 6, p. 16

Asiatic Herpetological Research

June 1995

FIG. 15. Male of the east Vipera dinniki ("eastdinniki") at the sympatric locality, Mt Elbrus,
central Caucasus. This specimen was found together

with the Vipera lotievi on figures 17 and 18 (ZIG).

FIG.

16. Ventral side of the male of the east
Vipera dinniki ("east-dinniki") at the sympatriclocality, Mt Elbrus, central Caucasus from figure 15(ZIG).

FIG.

17.

Female of the Vipera

lotievi at thesympatric locality, Mt Elbrus, central Caucasus.
This specimen was sympatric with the Vipera
dinniki ("east-dinniki") on photo 15 and 16 (ZIG).

FIG. 18.

Ventral side of the female of the Vipera

lotievi at the sympatric locality, Mt Elbrus, central

the west Caucasus there aresome isolatedoftheursinii
populations
complex (from the
Abishiz-Akhuba Range to Mt. Elburs).

Caucasus from photo 17 (ZIG).

"nebulosa-dinniki" pattern of eastern dinniki

The eastern form of dinniki is notpolymorphic in the same way as the western
populations. Rather the main color patternis the typical "nebulosa-dinniki" kind ofpattern. In sympatric areas the ursinii taxon
is more or less typical "mountain ursinii" incolor pattern

and the "seoanei-ursinii" pattern of
Caucasian ursinii taxon shows great
similarities, and can in some specimens bedifficult to separate. This is certainly thereason for much of the confusion in earlier
studies of these vipers.

although with a certain

Checheno-Ingushetia, a certain fraction of
the snakes are also "bronze" colored (Plate

However, at a closer examination, thetwo taxa are possible to identify (Table 9).
In ursinii the belly is lighter and the snout
more concave with raised canthus. The
preocular is large and in contact with theThis pattern type has not beenId).
observed in the sympatric populations. The

nasal, and the apical is always single. Also
the crownscales are less fragmented. Theresimilarity to the type of bilineate pattern seenin the southwest European V. seoanei
(Fig.

20).

In some populations of this taxon inJune 1995

Asiatic Herpetological ResearchVol. 6, p. 17

TABLE 9. Main morphological characteristics separating the sympatric Vipera lotievi and "east-dinniki".
lotievi

White belly
Preocular in contact with nasal

Snout concave

Mean no. of ventrals
Always a single apical
Mean no. of crown scales
Parietal ocellated spot
Iris gold-edged in life

Asiatic Herpetological ResearchVol. 6, p. 18

FIG. 20.

Female of Vipera

lotievi sp.n.

June 1995

from the type locality, the surroundings of Armkhi Village,

Checheno-Ingushetia, Nazranovskiy District.

always seems to be an ocellated spot present
is high.

ursinii constitute a distinct evolutionary
lineage (Herrmann et al, 1987; 1992). The
genetic comparisons of the west Caucasianisdinniki and the ursiniion the parietal plate, and the ventral number
In the "east dinniki" taxon the belly
blackish and the snout more flat, thepreocularisalways separated from theChecheno-Ingushetia

taxon

point in

the

from
same

nasal, and there is a higher fragmentation of
the crown scales. The iris always seems todirection (this study; Nilson et al., 1994)
except that the closer relation between

be gold-edged in live specimens (as is thecase for the entire V. kaznakovi complex),

kaznakovi and aspis was not supported.
Thus this morphological similarity between
the two lineages in Caucasus might be a caseand this

specially distinct in youngerThe
ventral number is lower.
specimens.
is

Althoughinmany ways

similar inpholidosis the kaznakovi lineage and theursinii lineage are genetically well separatedof convergent adaptation towards a similar
habitat, although Muellerian mimicry might
be involved.

A number of nominal taxa related to

Immunologicalthese populations are recognized from thiscomparisons of blood serum albumins
indicate that Vipera kaznakovi and related

geographical region (Russian Republic,
Georgia, Azarbaidjan and Armenia): renardi,

andparaphyletic.

taxa belongs to the berus-aspis branch whileJune 1995

Vol. 6, p. 19

Asiatic Herpetological ResearchDistribution of the vipers of the Caucasus and adjacent areas discussed in this paper. Light
horizontal
stippled= Vipera renardi; dark stippled= Vipera kaznakovi; cross-hatching= Vipera (u.) eriwanensis;

FIG. 21.

hatching= Vipera lotievi; vertical hatching= Vipera dinniki. Due to environmental reasons the distribution of
Also

all taxa are only fragmented within the depictured ranges, a situation especially pronounced in renardi.

of the ursinii
occurring in the region and related to the vipers discussed are the north Iranian populations
inthesoutheastern
intomountains,that
Vipera darevskii (ofTalyshAzarbaidjan
penetrates
complex ('ebneri')the kaznakovi complex) which has its known distribution restricted to northwestern Armenia (Mt. Legli), andV.
pontica from the Artwin province in Turkey. Other species of vipers not discussed here occur
sympatrically in the region.

kaznakovi, darevskii, eriwanensis, dinniki(Fig. 21).

What names are then available for thesetwo different sympatric central and east
Caucasian taxa?
Vipera dinniki wasfrom
described
Malaya Laba andoriginally
Svanetia, localities situated on the westernside of the upper parts of Little Laba Riverand the high-mountain basin of the Inguri
River, respectively (Orlov and Tuniyev,
1986). The type locality has been restricted
to Malaya Laba (by selection of 'The
Museum of Natural History of Kharkov
State University specimen no. 26044' as
lectotype;

Vedmederja et al., 1986).

Thesituated in the western
Caucasus and the polymorphic western

type localityisVol. 6, p. 20

Asiatic Herpetological Research

June 1995

TABLE 10. Variation, given as Mean ±S.E. and range (for apicals in % of specimens with two plates) ofselected morphological characters in Vipera eriwanensis (N=44), Vipera lotievi (N=14, if not otherwise stated),andVipera renardi (N=42).

Asiatic Herpetological Research

June 1995

TABLE 1 1.

Vol. 6, p. 21

Frequency of certain characteristics in the populations (in percentage of investigated specimens)

eriwanensis

Vol. 6, p. 22

Asiatic Herpetological Research

June 1995

FIG. 22. The female holotype of Vipera lotievi (ZIN 20309), Armkhi, Checheno-Ingushetia, below Mt.
Stolovaya, 2000 m altitude.

1986-05-28, leg. B.Tuniyev;

ZIN 20307,

Itum-Kali, Checheno-Ingushetia, 1987-08,

ZIN 20312, Armkhi,
Lotiev;Checheno-Ingushetia, 1987-09, leg. Lotiev;
ZIN 20313, Armkhi and Mt. Stolovaya,
Checheno-Ingushetia, 1986-07-20, leg.
Gizatulin; ZIG 297, Mt. Elbrus, 1986, leg
Filippov, coll. Tuniyev; ZIN 18203,
Teberda, State Reserve, Mt. Bolshaya
Hatipara, 1969, leg.
Zalslavsky; ZIN
18226, Kabardino-Balkaria, vicinity ofvillage Terskol, 1970-08-19, leg. Kireev;leg.

ZIN 11996, Caucasus, Gunib, Dagestan,
1909-05-29, leg. Berg; ZIN 20303,
Lagodechi, 1988-07, leg. Bakradze.

Diagnosis and definition: A speciesof the Vipera ursinii complex characterized
by polymorphism in color-pattern, including"bilineate pattern" of the same kind as in V.
seoanei, and "bronze" unimorphs. External

morphology evolved as typical for mountaintaxa of the ursinii complex but not similar toany of the other in color pattern.

Vol. 6, p. 23

Asiatic Herpetological Research

June 1995

From the sympatric "east-dinniki" itdiffers in several scalation characters and incolor of the belly (Table 9, Fig. 11). In
lotievi the belly is generally white, preocularin contact with nasal, snout concave, 138 or
more ventrals, always a single apical, lesson right and left side respectively, 1 canthaland 1 supranasal scale on each canthus
rostralis, but the two supranasals are partly
united with the apical; 3 intercanthals and 6

fragmentized crown scales (7-16), parietal

intersupraoculars. Height/depth of rostral
3.4/2.7(=1.26), it is bordered by 2
supralabials, 2 internasals and the broad

ocellated spot present, iris not gold-edged in
In "east-dinniki" the belly is black,life.
preocular separated from nasal, snout not"apical"; eye surrounded by 8 circumoculars
on each side, 5 loreals on each side, upper
preocular in contact with nasal on bothconcave, 136 or less ventrals, apical single
or divided, more fragmentized crown scales

sides, nasal partly divided at upper edge, 8(10-21), no parietal ocellated spot, iris gold-edged in life.

From the allopatric renardi it differs
besides color pattern in morphology byhaving light supralabials (sutures heavily
colored in black in renardi), a higher rostralindex, smaller size, white belly (dark inrenardi), and a different niche by being

alpine {renardi is a lowland steppe
No future reproductive
inhabitant).
cohesion can be postulated.

from the likewisein the Armenian
eriwanensis
allopatricthesemidesert
lowland of thehighlands byKura River Valley, that separates the Big
Caucasus from the Small Caucasus. No
connection can be postulated in anItisseparated

mm

supralabials, with forth below eye, and 9
sublabials on each side, anterior supralabials

not much enlarged compared to posteriorones, 6 second chinshields bordering theanterior ones and 4 scales in the gular row.
Dorsal side of snout concave resulting in apronounced and raised canthus rostralis.

Two preventrals and 141 ventrals, 24+1
subcaudals,

21

dorsal

scale

rows

at

midbody and on neck one head-length
behind the head, 17 dorsal scalerows one
head-length anterior to anal. Reduction
from 21 to 19 dorsal scale rows at level of
Dorsal patternventral number 89.
consisting of a weakly winding zig-zag bandwith 48 windings, lateral body pattern dark
weakly contrasting towards the lighter
dorsal groundcolor.

Head pattern consistsof 2 dark oblique bands which do not unite,
and a posterior band from eye to corner of

evolutionary time frame. Besides color
pattern there is a differentiation inmorphology by eriwanensis having a higher
number of crown scales and a somewhat
lower ventral count, and preocular separated
from nasal to a higher degree (Tables 10 andmouth and somewhat further back along thelateral sides of neck, no dark pattern on chin

11).

ventral side light, throat light.

or in labial region although a very weak
dotted pattern at the supralabial sutures canbe imagined, ground color light brown withdorsal pattern dark brown and black edged,

Ocellated

spot on frontalia.

Descriptionofholotype(Fig.

An adult female, total length 422 mm,
tail 41 mm, latter equal to 10.8 % of total
22):

Length of head, from posteriorborder last supralabial to tip of snout 16.8mm, from posterior border of parietals to tip
of snout 12.2 mm, breadth of head atbroadest part of head 9.5 mm, at level of theeyes 8.0 mm, size of eye horizontally 2.5
and vertically 2.0 mm, distance between
eye and lip 2.6 mm. Head covered withrather large scales or plates. Two large
length.

mm

supraoculars and 1 large frontal plate on topof head, parietals large, frontal separated
from supraoculars by 3 and 2 smaller scales

See Tables 5 and 6.
Variation:
Besides the variation in scalation a
pronounced variation in color and pattern isexpressed. Most striking, and unique forthe entire ursinii complex, is the bronzemorph, which is found in 25 % of theinvestigated specimens (N=40) (Plate Id).
Distribution:

Vipera lotievi isthe semiarid 'hollows'
between the northern slope of the main
Caucasian range and Skalisty range from theupper part of the Kyafar River (range
Abishir-Akhuba) eastward to the interior of

distributedinVol. 6, p. 24

Asiatic Herpetological Research

Daghestan (see map). Altitudinal span inthis region goes from 1200 m up to 1600 m(1800m). Further it is recorded from Mt.
Elbrus in the central Caucasus, andmountains north of Lagodechi in the eastern
Caucasus, besides several isolated records
from the eastern and central Caucasus range.

Habitats:
Typical habitats areoreoxerophytes landscapes with semiarid
light-forests (like Shibliak), phrygana (with'tragakant' astragalus) which are very
similar to east-Mediterranean types ofvegetation. On the upper elevation of the
distribution V. lotievi reaches the subalpine

mountain belt.

"Refuge History": The developmentof the xerophilous vegetation has taken placesince Pliocene in the eastern part of the
Caucasian Isthmus. Four main refuges are

known: two humid (Colchis and TalyshHyrkanian) and two arid-xerophilous(Armenian and Dagestanian). The north
Caucasian refuge of oreoxerophits,
including shibliak and phryganas are
situated along the shale-depression between
the main Caucasian Range and the Skalistiy(Rocky) Range. There are several semiarid
hollows from central Dagestan (Gunibskoe
Plateau) and westward to the beginning of
the Kuban River (at the Mt. Elbrus region).
The vegetation is composed of Juniperus
oblonga, Paliurus spina-christi, Cerasus
incana, Colutea orientalis, Berberis vulgaris,

Astragalus denudatus, Celtis glabrata,
Ephedra procera and others. This vegetation
superficially is very close to the vegetationof the Armenian highland and the mountainsof the Near East, but the regions share

June 1995

Kuznetsov, 1890), while Galushko (1974)
has the opinion that the semiarid hollows ofChecheno-Ingushetia are younger than thehollows of Kardino-Balkaria and Osetia in
the west and Dagestan in the east, andperhaps not older than Holocene. But theremains of xerophilous flora on the crests
between the semiarid hollows are thewitness of the existence of a unitedenormous xerophilous (Mediterranean)
refuge, running from Dagestan to the regionof Mt. Elbrus. Later, in Pleistocene, thisrefuge disintegrated to several micro-refuges
which have persisted to different extent until
present. However, it must be pointed outthat although the xerophilous vegetation(including mountain-steppe) had a wide
distribution along the shale-depression, it
also had the possibility to disperse up to themountains along the river valleys.

Bothways could be used by representatives of the"ursinii-sptcics group" of vipers. Besides
the "ursinii-group" also thermophilous

species like Lacerta strigata, Coluber
najadum, and Elaphe hohenackeri arepresent as isolates in these hollows. One

should also pay attention to the occurrenceof relicts of the xerothermal epoch in thewestern Caucasus near the mountains
Jatyrgvarta and Magisko (Altukhov, 1966),
but the xerophilous vegetation did never

have any wide development in that area.

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