Asiatic Herpetological Research

February 1992

Vol. 4, pp. 113-122|

Intrapopulational and Geographic Variation of Eremias przewalskii StrauchinMongolia

VALENTIN A F. ORLOVA

1^Zoological Museum of the Moscow Lomonosov State University, Russia

Abstract. -Body size and proportions, characters of pholidosis, coloration and pattern of Eremias
przewalskii from western, southern and eastern Mongolia were studied. Sexual dimorphism is displayed in
the relative sizes of the tail, head, legs, number of scales around 9- 10th tail ring and ventralia. Somecharacters of pholidosis, as number of scales around the mid-body, number of femoral pores on the rightside of the thigh and others displayed clear geographic variability. Live specimens from western Mongolia

have blue ocelli on the body flanks. The lizards from southern Mongolia significantly differ from thespecimens from other parts of the range by a larger number of scales around the mid-body, number offemoral pores on the right side of the thigh and other features. The intraspecific structure of E. przewalskii
is discussed.

Key words:

Reptilia,

Squamata, Lacertidae, Eremias przewalskii, Mongolia, geographic variation,

morphometries, pholidosis, population variation.

Rarely, E. przewalskii lives on saline soil,
almond-bush. In

Introductionin the dry gullys with

Eremiasprzewalskii Strauchisacommon species of western and southern
Mongolia (Bannikov, 1958; Dely, 1979,
1980; Munkhbayar, 1973, 1976; Obst,
1963; Orlova, 1984, 1989; Orlova and
Semenov, 1986; Szczerbak, 1970, 1974;
Terbish, 1989). Earlier it was thought that
only occured along the boundaries ofsoutheastern Mongolia (Bannikov, 1958;
Szczerbak, 1974). Now it is known to beitmost widely distributed lizard in thisregion (Semenov and Shenbrot, 1986).
Outside of Mongolia this lizard is
distributed in northern China and Russia, in

Bayan-Drag it is found among the stones ofCretaceous red sandstone precipices
As(Borkin, 1986; our observations).
thisto
Eremiasmultiocellata,
compared
species does not extend into highmountains, inhabiting either hollows andfoothills with elevations 760-1800 m above
sea level (Szczerbak, 1974) or in anarrower range of elevation, 1030-1650 m
above sea level (Borkin, 1986).

thethe southern portion of Tuva Autonomous

Republic (Bedriaga, 1909;

Flint,

przewalskii

who have observed E.

in thewild have noted

its

preference for soft soils. This species
inhabits sands overgrown with Nitraria sp.,sand dunes with

Tamarix sp.

Haloxylon

sp.

and

In the Transaltai Gobi, E.

przewalskii rarely occurs in gravel areas
adjacent to sandy habitats (Borkin et al.,In Uws-Nuur hollow and on thebankof the river Khowd-gol it occurs
right
on semi-anchored sands with Caragana.

1983).

1992 by Asiatic Herpetological Research

its range, E.
przewalskii iswithE.
multiocellata(e.g., near
syntopic
the well Buiesengijn-khuduk, 35-40 km
northeast of Ba-Tsagan, in Bayan-Dzag).

1960;

Pope, 1935; Schmidt, 1927; Strauch, 1876;
Szczerbak, 1974).
All researchers

Withinofexternal
variability
E.
featuresofprzewalskii
morphologicalhas been noted repeatedly in the literature.
Such investigations were started by Strauch
in his description of three Eremias species
from China in 1876. Later (Boulenger,
1921; Nikolsky, 1915; Szczerbak, 1969,1974) these forms (E. brachydactyla, E.

High

przewalskii and E.

kessleri)E.
withprzewalskii.
synonymized

wereTreatment of collections made by the
Herpetological Department of the Joint
Soviet-Mongolian Complex BiologicalAsiatic Herpetological Research

Vol.4, p. 114

i

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February 1992

Asiatic Herpetological Research

February 1992

characters of pholidosis were taken into
consideration: 1 - number of scales aroundthe midbody (Sq.); 2 - number of scales
along mid-line of throat (G.); 3 number offemoral pores on the right side of the thigh- distance between the internal(P. fm.); 4
sides of the rows of femoral pores; 5 number of transversal rows of pectoral andventral scales (ventrale); 6 - number ofscales around the 9- 10th tail ring (Sq. c.
number of subdigital lamellae oncd.); 7the 4th toe of right hindleg; 8 - number
number of
supralabial scales (labialia); 9

scales (infralabialia); 10 number of dorsal scales between parietals
and level of anus; 11 - number offrontonasal scales.
infralabial

characters mentioned
above, pattern and body coloration wererecorded, including the presence or absence
In additionto

For the treatment of material standard
statistical methods were used (Lakin,1980)

with the calculation X, m x and t-criterion
for revealing sexual dimorphism andResults

dimorphism.

— Eremias

przewalskii males and females in Mongolia
differ from each other by there snout-vent
length in all samples, except those from

Aymag (5) and East-Gobi

Aymag (6). Males are slightly larger thanfemales. The maximum difference in linear
size between the sexes was in South-Gobi

Aymag sample

(5)

from

southern

Mongolia, but statistically significant
differences were not revealed, as in allother samples (Tables 1 and 2). The tail inthe males is longer than in the females

samples from Gobi-Altaiand
Gobi-Altai Aymag (3)]; as
Aymag (2)[except

theis length of the hind legs

also

sex

differences, althoughdisplayed
Szczerbak (1970, 1974) considered suchsexual dimorphism to be absent in E.

przewalskii.

In

all

the samples

weinvestigated the number of scales aroundthe 9- 10th tail ring is larger in males than infemales (Fig.

2).

In the samples

fromGobi-Altai Aymag (3), South-Gobi Aymag

and East-Gobi Aymag (6) suchtendency is displayed in the number oftransversal rows of the pectoral and ventral
In specimens from westernscales.
Mongolia (samples 1-4) the number ofscales around the middle of the body(5)

differed between thesexes, but thesedifferences are statistically insignificant.

Coloration and pattern.

—We did not

find clear differences in coloration andpattern between males and females. The

western part of its range, E. przewalskii
has blue spots on the flanks which is lessbright in old females.

Thus,

geographic differences.

South-Gobi

Some characters of pholidosis

ventral surface of the body, legs and tail isalways entirely white in both sexes. In theof blue spots on the body sides.

SexualVol.4, p. 115[except the sample

from East-Gobi Aymag (6)]. As to thehead proportions, there is a stabledifference in relative head length (P<0.01,
At the same time, otherP<0.001).
proportions, mainly Hp/Lp, slightly
differed from each other in both sexes.

sexualdimorphism

in

E

.

przewalskii is displayed in relatively longtail, head, both pairs of legs, number ofscales around the 9- 10th tail ring andventrals in males as compared to females.

Geographic Variability—

Body size and proportions. The largest(maximum length 84.5 mm) lizards inhabit
the southern and southeastern parts of thecountry, and the smallest lizards in thenorthern and northwestern regions.
Moreover, animals of minimum size havebeen found near lake Alag-Nuur, GobiAltai Aymag (3). Slender specimens withrelatively long tail and hind legs are seen in
the samples from Gobi-Altai Aymag (3)and South-Gobi Aymag (5). Lizards fromthe southeast of Mongolia [East-Gobi

Aymag (6) sample] are similar to thosefrom the south [South-Gobi Aymag (5)] in
the relative tail length, but they have a morerobust habitus and shorter hind legs. Thus,
within the range of E. przewalskii, its bodysize, relative tail and hind leg length

Vol. 4,

Asiatic Herpetological Research

February 1992

Vol. 4, p. 117

40r

35

B
30-

25

D

FIG. 2. Sexual dimorphism and geographic variation in the number of scales around 9- 10th tail ring (Sq.
c. cd.)

The

external

Between these colored areas there are 4

morphological features includes, along withothers, the number of scales along the spine(from parietals to posterior border of hindAymag (5) sample this feature shows alongitudinal rows of white ocelli or larger
eroded light spots. The legs (especiallyhindlegs) are covered by a pattern of lightspots surrounded by brown. The upper tailsurface (about one third of the tail length)

similar situation as inmany others, i.e.

has a similar pattern, caudally it is divided

number is markedly higher than in

into single dark spots, and the tip of the tail

leg).

their

characteristicsof

For the lizards from the South-Gobi

lizards from the western part of the range.

is light.

Other features, such as the number ofsupra- and infralabials, are less variable.

olive-grey in immature specimens without
spots, or with a few spots in the parietal
and supraocular scales. There is a dark and—Coloration and pattern.
The coloration
varies from sandy or grey to dark-brownand black. The pattern may be made up
either of relatively thin lines, or of rather
wide, interwoven, waved stripes and spots.
The dorsal pattern of specimens from theGreat Lakes Hollow (Fig. 5) is formed bysandy or light-coffee wavy lines or spots.

The upper surface of the head islight striped pattern on the side of the parital

scaleswhich

sometimes reaches thesupraorbital. In individual specimens theentire head surface is covered by darkwhich are more or less apparent.
Temporalia are covered by a pattern of dark
spots,

spots or stripes alternating with light spots
or ocelli. The ventral surface of the bodyVol.4, p. 118

Asiatic Herpetological Research

TABLE 3. Geographic variation in characters of pholidosis.

CharacterFebruary 1992

Asiatic Herpetological Research

February 1992

Vol. 4, p. 119

lizards have the most bright and contrasting
dorsal pattern (Figs. 5, 6 and 7). It is black
and white, with the pileus pattern as bright.
The ventral surface is white, but a slightly

yellowish tint may be present. Regarding
the blue spots on the body flanks, one
could not make a definite conclusion.
Fixed specimens may have blue spots, butthese, as a rule, are positioned on the parts
not characteristic for their usual occurrence.

This

could

be

a

consequence

preservation in alcohol.

ofIntrapopulationofpattern polymorphism is characteristic
Lake
the
2
from
Beger-Nuursamplewhere

the following

environs (Fig. 6),
specimens occur: 1- with black wavy spots(combined with white ones) on the bodyflanks, with a brown tint in the middorsum; 2- with an unclear dorsal pattern
and three rows of white spots (surroundedby black) on the body flanks; under theserows is a row of blue ocelli on each side; 3with a pattern of thinner transversallyelongated, black, wavy stripes, connectedwith each other or isolated, and with single
white ocelli. The same pattern covers also
the anterior one third of the tail, and is thendivided into single small spots of darkcolor. The head in most cases is light-grayor beige, with a pattern on the parietal and

FIG. 5.

The dorsal surface pattern in Eremias

przewalskii from the northern coast of Char-UsNuur Lake, Somon Urdgol (=Chandman) sample 1.

temporal parts (mainly in specimens with a

These blue spots arevisible in almost all thefine pattern - type 3).

more or lessspecimens.

Discussion
Analysis of Eremias przewalskii
intrapopulation size and proportions

The dorsal surface pattern in Eremias
Lake Begerprzewalskii: from Gobi-Altai Aymag,

FIG. 6.

Nuur, sample 2.

variability in Mongolia reveal clear sexual
They are expressed todifferences.

different degrees in populations studied

southern andthewestern,southeastern parts of Mongolia. These
differences do not concern, as a rule, theabsolute sizes of the body (L) but
tailrelatively, length of the head, limbs andare larger in males than in females. We
have also shown dimorphism of somefromcharacters of pholidosis in our material,
including samples of adult specimens fromeach locality. Such sexual dimorphism had

not been

shown by

Szczerbak (1974)

during his researches of Tuva (Russia),

FIG. 7.

The dorsal surface pattern in Eremias

przewalskii: from South-Gobi Aymag, sample

5.

Asiatic Herpetological ResearchVol. 4, p. 120

northwestern Mongolia, China and single
specimens of Eremias from southern
Mongolia. The number of scales aroundthe 9- 10th rings of the tail differs in all thesamples and the number of ventrals differs
in the samples from Gobi- Altai Aymag (3)and East-Gobi Aymag (6). The colorationand the pattern of adult specimens have nosignificant sexual differences.

The wide range of pholidosis variability
is clearly expressed to various degrees in all

the samples from Mongolia.

Dely (1979,

February 1992

and pattern are also variable, but among
specimens studied we have not found"rough-spotted" ones. In China, where thedorsal pattern of the lizards is known to bevariable, nobody has mentioned the blue
spots on the body flanks, which arecharacteristic of E.

przewalskii in thewestern part of Mongolia. Strauch (1876)
gave detailed descriptions of single
specimens of the new species, but not in
one case did he note blue spots. Possibly,
southern Mongolian specimens also do nothave these spots. In this connection the(ZIN 5145:

1980) speculated that an explanation of thespecimen of E.

high variability may be ecological isolationand interbreeding, connected with relatively

collection of Zoological Institute, Russian

low population density and low fecundityof the females. It was interesting to find 2
frontonasals in 25% of investigated lizards

from southern Mongolia (Orlova, 1989)
together with other characters (larger size,
contrasting coloration, larger number ofscales around the midbody, number of
dorsal scales between parietals and level ofanus, etc.). It is curious not only as a trait
reflecting the isolation of these populations
that the presence of 2 frontonasals isnormal for E. argus and occurs in E
multiocellata as an exception. It may.

suggest a close relationship of thesenoted by previous authors(Bedriaga, 1912; Szczerbak, 1974).
Szczerbak has not mentioned specimens ofE. przewalskii with 2 frontonasals in
Mongolia and did not report this in thedescription of the species and nominative
species,

subspecies, while Strauch (1876) indicatedin the description of his new species.

it

Strauch's descriptions of 3 species from
China were based on the variability ofcoloration and pattern of E. przewalskii.
Specimens with a "rough-spotted" patternconsisting of black or dark-brown stripesor spots drawn in transverse direction were
described as E. przewalskii.. Specimens
with a "netted" pattern consisting of fine,
fused, wavy, interwoven lines were
described as E. brachydactyla. The third

of

was

designated by(1970) as "transitional",
intermediate between the first and thesecond characteristics of E. kessleri.
Within Mongolia, these lizards' coloration
type

pattern

Szczerbak

kesslieri

Academy of Sciences, St. Petersburg) fromthe lower Tarim, collected by N. M.
Przewalskii, is interesting.
Bedriaga(1912:577) wrote about this specimen:"Das Originalstuck der E. kessleri stammt
aus Gansu; ein anderes soil von Przewalski
am unteren Tarim erbeutet worden sein (N5145). Diese weit westlich vorgeruckte
Fundstelle und die Thatsache, dass nur ein
einziges, an den Seiten sonderbarerweiseblau geaugtes Individuum von dort
mitgebracht worden ist, hat in mir Anfangs
einige Zweifel hinsichtlich der Herkunftdesselben erweckt; dock uberzeugte ich
mich nachtraglich, das es im Jahre 1878

dem akademischen Museum ubergeben
worden ist, und dass der General vorher,
namlich in den Jahren 1876 und 1877, ausseiner Reise nach dem Lob-nor in
Wirklichkeit am Unterlauf des TarimFlusses gewesen ist."

no samples from the enormous
Mongolian territory were such drasticIn

differences recorded, as in
specimens from China, where lizards with
2 pattern types coexist (collections of the
Zoological Institute, Academy of Science,pattern

St.

Petersburg; T.

J.

Papenfuss and J. R.

Macey, pers. comm.).
At present, E. przewalskii is considereda species with two subspecies: nominative
E. p. przewalskii (Strauch) (southern
Mongolia and northern China) and E. p.
tuvensis Szczerbak (Tuva, Russia, andwestern Mongolia) (Szczerbak, 1970,

Dely (1979, 1980), analyzing thevariation of external morphological1974).

February 1992

Asiatic Herpetological Researchcharacters of lizards from Mongolia, noted
the population from the southern Gobi as

most clearly distinct. While the author did
not have the possibility of studying thematerials from Tuva, Russia he suggested
the existance of two subspecies of E.
przewalskii. At the same time, each of the
populations investigated by him wasreferred to the nominative subspecies.

Determination of the structure of thespecies as a whole will depend upon future
research on E. przewalskii from China(with detailed analysis of intrapopulationaland geographic variability, including
biochemical analysis).

Acknowledgments
would like to express my gratitude toDrs. Kh. Terbish, S. L. Kuzmin, E. A.
Dunaev and M. Prutkina for their help inI

collecting material and in the preparation ofthis paper.

Vol. 4, p. 121

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