IDecember 1993

Asiatic Herpetological ResearchVol. 5, pp. 5 1-58

The Variegated Toad Agama in Djungar Gate (Eastern Kazakstan) withNotes on Certain Systematic Problems of Phrynocephalus versicolor Str.
(Reptilia:Agamidae)

M. L. GOLUBEV
Instituteof Zoology. Academy of Sciences, Kiev, Ukraine

Abstract.-The distribution of Phrynocephalus versicolor in Djungar (=Junggar) Gate (Eastern Kazakstan)
was investigated. The characteristics of these lizards are: absence of red axillary spots and the presence ofP. versicolor
red-orange subcaudal coloration. The taxonomic status of this population and of subspecies ofis discussed.

A^y Words:

The variegated toad agama is presumed to be a "composed" species.

Reptilia, Sauria,distribution.
systematics,

75
sa-oo'Agamidae, Phrynocephalus versicolor, Kazakstan, China, Djungar Gate,1Vol. 5, p. 52

Asiatic Herpetological Research

December 1993

FIG. 2. Habitat of Phrynocephalus versicolor in Djungar Gale near Druzhba Railway Station:
stony and gravel semidesert covered with boyalych (Sal sola orbuscula).

Methods

We studied the distribution of thevariegated toad agama in June 1991 during
investigations of the Djungar Gate territory(Fig. 1).

crushed-gravels and fine gravels on a loess basewith the finer sorted materials deposited
farther from the mountains slopes.
Southeast of Lake Zhalanashkol the valley
floor is a broad alkaline plain (20-25 km.)with subsurface water.

Results and Discussion

The dominate plant, Salsola orbuscula,(common Russian name = boyalych, also

The Djungar Gate is a relatively narrowpass between the Balkhash-Alakol

known in the United States as Russian

Depression and Chinese Djungaria. This
pass is oriented from the northwest to thesoutheast and rising in elevation towards
the southeast. The pass enters China near

alluvial plains of the valley.

the Druzhba ("Friendship") railway station

dominate among theandthegrasses
nearlyonly plant on flats

along the Lankol Valley. The Djungar Gate
Valley is formed by the broad alluvial plain
and gentle foothills of Maily Ridge and theDjavlau Mountains to the northeast and themore abrupt upthrust of the Djungar Alatau
to the southwest. The surface of the valley
alluvial plains varied with the degree ofslope and ranged from boulders, rubble.

thistle) isfound on the lower slopes andIt is

morewidely distributed on the northeastern
slopes but it is sometimes replaced bysaxaul(Haloxylon(Artemisia sp.)

sp).

Wormwood

iswithout shrubs.

From the Lankol Valley the toad agamas
are distributed along both the northeasternand southwestern slopes above the Djungar
Gate. Along the southwestern alluvial plain
the toad agama is distributed 15-20 km

Asiatic Herpetological Research

December 1993

from the Druzhba station to the northwest.
The lizard occurs along the foothills of theMaily Ridge and the Djavlau Mountains for
55-60 km where the northern limit of its
distribution coincides with the border of thebeVol. 5, p. 53

Currently P. versicolor is considered toa polymorphic species and it isinteresting to determine the subspecificposition of the form inhabiting the Djungar
Gate.

boyalych dominate, gently sloping alluvial
plain composed of rock rubble and gravels.
Further north, on the steeper slopes of thealluvial plain composed of larger rockrubble and dominated by wormwood, thesunwatcher (P. helioscopus) is found. It ispossible that P. versicolor occurs muchfurther to the northeast into the Alakol

Depression along thefoothills,

howeverfield work in this area is difficult because of

the presence of military installations along

A small isolated area withthe border.
conditions which would make good habitat
for this species is found along the railway

tracks between the Zhalanashkol Stationand the 19th Station near the mouth of theIrgaty River.

Toad agamas are found under single
bushes in small groups composed of 1-2males and 2-5 females of different ages. In
addition, groups of up to 10 subadults wereobserved. The density of the lizards isvariable, with higher densities in gravelly
areas with boyalych (Fig. 2) as well as inareas of colonies of the great gerbil(Rhombomys opimus ) that have excavatedthrough the darker colored gravels and rockrubble and where the lighter loes makes up
the predominant coloration of the surface.
It is interesting to note that the lizards
inhabiting the gravel plains in the Djungar
Gate have retained the sand burrowing
behavior,
involving rapid lateral

movements ofbody, observedpopulations inhabiting sandy areas.
theinPregnant P. versicolor as well as femalesof other Phrynocephalus species assume
the "copulation avoidance" posture whenpursued by males (Polynova, 1982; 1989;
Rogovin, 1991; and our observations of P.
strauchi in the Fergan Valley). To assume
the "copulation avoidance" posture thefemale turns onto her back as the maleapproaches and maintains this position
while he is nearby.

For a long time it was assumed that ineastern Kazakhstan this toad agama wasfound in three isolated populations: theZaissan Depression, Alakol Depression andthe Hi River Depression (Paraskiv, 1956;Bannikov et al., 1977). Peters (1984)
considered the Zaissan Depression and HiRiver Depression lizards to be two seperate
Bedr. and P.
species: P. salenskyi
Bedr.
Threealpheraki
years later a newP.
versicolor
subspecies,
paraskiwi(Semonov et al., 1987), was describedfrom the Hi River Depression. Theseauthors speculated that the two Chinese
Djungar Depression subspecies, P. v.
hispida Bedr. and P. v. doriai Bedr. wereHowever, becuase of aconspecifics.
shortage of material, they did not determine
the taxonomic status of the Alakol
Depression variegated toad agama. Soon
after the most significant attempt to analyze

the intraspecific variaton of P. versicolor

was undertaken (Semenov and Shenbrot,1989).

Semenov and Shenbrot (1989) examined
675 specimens: 580 from Mongolia andTuva, 65 from the Hi River Depression, 19
from Chinese Kuldja (now Yining,Xinjiang, China), 11 from the Alakol
Depression, but no specimens from theremainder of the range of this species in
China. The authors, using discriminant
analysis techniques, felt their material wasadequate to discuss all known subspeciesof the variegated toad agama.

Semenov and Shenbrot (1989) indicatedv. paraskiwi was detached from themain group. Also, P. v. doriai fromthat P.

from western
wereresurrected.
The Alakol
Mongolia
Kuldja and P. v. kulagini

agama wasvariegated toadsingularly

These authors
were unable to distinguish the lizards from
Mongolian Djungaria from "typical" P. v.
hispida, however they did not indicate
which P. v. hispida they considered
attributed to P.

v.

doriai.

Vol.

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54

December 1993

Asiatic Herpetological ResearchVol. 5, p. 55

FIG. 3. Dorsal view of Phrynocephalus versicolor from Djungar Gate (Djungar Railway Station).

10 years and these spots might havedisappeared during this time. Also there
are no detailed data on the distribution ofthis character in lizards from eastern
Djungaria. It is known only that such spots
are present in the Mongolian part of therange of the toad agama (Semenov andShenbrot, 1989). However, the nearly

isolated

Mongolian Djungaria (Barun-Khuray Depression) differsfrom Chinese

Djungaria in several geogrphic parameterssuch as altitude.

The type specimens of P. v. hispida aredated 1879 (ZIN 6637 and 6638) and 1880
Nikolay M. Przewalsky's
Third Central Asian Expedition (FirstRidge and the plains of south eastern
Djungaria. On May 18 (May 30 by themodern calendar) he reached Barkul anddid not return to Djungaria during thisIn
expedition (Przewalsky, 1883).
thetoadagamas
Przewalsky's journal{Phrynocephalus sp.) are casuallymentioned for the middle and lower reachesof the river. Thus, it is possible to draw
two conclusions: (1) the P. v. hispida typespecimens may have been collected indifferent parts of Djungaria and thus
include different forms, (2) the date ofcollection for the ZIN 6639 sample isincorrect.

(ZIN 6639).

Tibetan Expedition) took place at this time

(Dubrovin, 1980). Przewalsky left the cityof Zaissan on March 21 (April 4 by themodern calendar) and reached Ulungur
Lake (Ulungur Hu) on March 31 (April 12by the modern calendar). He followed theUrungu River (Ulungur He) andits

tributary the Bulugun to the Barun-Khuray

Depression and crossed it from north tosouth. He then crossed the Baytik Shan

Thereisan overlap between bothof toad agamas from thesubspecies
Djungar Gate region in body proportions(females 0.6-1.06; males 0.64-0.75),
number of scales across (19-29; notcounting the supraocular scales) and along(8-14) the top of the head. The scales on
the thigh are smooth and dorsal and ventral
coloration are highly variable (Figs. 3, 4,
In the material we collected, a slight5).
shift in these character's values towardAsiatic Herpetological ResearchVol. 5, p. 56

December 1993

FIG. 4. Ventral view of Phrynocephalus versicolor from Djungar Gale (Djungar Railway Station).

hispida can be noted. It is important to notethat the subcaudal surface in living lizardsof both sexes is a red-orange color, thatwith age looses its lustre and disappears(Fig. 4). We also discovered remnants of
the red-orange coloration in one of 13

specimens from "Alakol Depression"(ZoologicalMuseum of Moscow State
Recently weUniversity, MSU R7779).

examined specimens from China with thesame subcaudal coloration in the California

Academyof Sciences collected in thecentral (northeast of the city of Karamay)

and southeastern (east from the city ofUrumqi) Djungaria. In all specimens
examined there are dark transverse bars on
the ventral tail surface and this agrees withBedriaga (1909) but not with Semenov

indicate the existence of a new subspecies?

There are also red tailed toad agamas innorthern Djungaria and the Zaissan
Depression variously described as P.
guttatus, P. salenskyi and P. versicolor.
In the Zaissan

Depression this lizard ismostly sand-dwelling and similar in habits
to P. guttatus and P. frontalis (Golubev,1989), while in northern Djungaria and insome places in the Zaisson Depression they
are found in more stabilized soils. Fromthe Alakol Depression and the Djungar
Gate, where this species is found, there areover 300 km of continuous habitat without
noticeable barriers into central Djungaria
where P. versicolor is found. This may
represent a cline with a gradual transition

from one form to the other.

(1986).

From the above data, it follows thatvariegated toad agamas inhabiting theAlakol Depression and Djungar Gate tosouthern Djungaria (>500 km) differ fromall other forms of P.
versicolor in theabsence of axillary spots and brightly
colored subcaudal surface.
Does thisIn September 1991 repeated copulations
between a male P. salenskyi (ZaissanDepression) and a female P. versicolor(Djungar Gate) were observed in the Kiev

Zoo terrarium.

If precopulation barriers
apparently can be broken interrarium conditions. Both Przewalsky(1883) and Potanin (1948), when travelingexist, theyAsiatic Herpetological Research

December 1993

Vol. 5, p. 57

along the lower reaches of the Urungu
River and the southern shore of Ulungur
Lake (type locality of P. salenskyi ), notedWe

the variable coloration of toad agamas.
discovered fragments of a light longitudinalcaudal stripe (a characteristic of P. g.
salenskyi) in some specimens of P.
versicolor from Djungar Gate (Fig. 5).
Red axillary spots are present not only in
the Alashan variegated toad agama (speciestype locality) but in lizards inhabiting thearea south of Beishan Ridge in the Gashun
Goby. However, here P. versicolor areconnected by coloration and patterntransitions with P. axillaris Blanf.

Thus, the question of the taxonomic
status of the variegated toad agamas from

Djungaria and Alakol Depression should
again be considered open as does thequestion of the position of P. v. doriai.
The Kuldja and western Djungarian
populations are seperated by the Tianshan
Mountains. There are reasons to believethat the Kuldja P. v. doriai is actually an
ecological race of the Hi P.
while the Ebinurian P.

g.
v.

alpherakii
doriai isassignable to the acutirostris group (which
also might be no more than one of the color
variants of the axillaris- guttatus complex.

summary, it appears that only twosubspecies of the variegated toad agama canInbe recognized. P. v. kulagini inhabits
southern Tuva and northwestern Mongolia
and forms a narrow zone of intergradation
with the nominative subspecies P. v.
versicolor. However, there are doubts thattheaxillaryredspots

constituteacharacteristic which allows one to delineate
populations specifically on the level of
It issubspecies.
ofconsists
thatP.
versicolor
possibleisolated, genetically differentiated color
morphs associated with stabilized soils.
Taxonomic seperation of these variants
should occur only after a detailed study ofthe entire group.
Bedriaga (1909)
recognized five subspecies and consideredP. versicolor to be a species composed ofgeographaic race,i.e.

highly variable populations.
Bedriaga expressed concern that his
taxonomic arragement of these subspecies
did not represent a natural assemblage.

manyA specimen of Phrynocephalus versicolor
from Djungar Gate (Djungar Railway Stauon)with

FIG. 5.

light longitudinal caudal stripe.

Further he stated "[only when we are morefamiliar with the fauna of Central Asia, will
it be clear whether I have exaggerateddistinguishing characters]". Leroy (1940)
proposed that the species P. versicolor beabolished. Leroy's point of view may beclosest to the truth.

Acknowledgments
The author is most grateful for the loan

Vol.

5, p.

Asiatic Herpetological Research

58of Phrynocephalus collections from M. E.
A. Dunajev and Dr. V. F. Orlova of theZoological Museum of the Moscow State
University, Russia; Mrs. L. Jogansen andProf. I. S. Darevsky of the Zoological
Institute of the Russian Academy ofSciences, St. Petersberg; and Mr. J. V.
Vindum and Dr. A. E. Leviton of theCalifornia Academy of Sciences, San

Also, I
Francisco, California, U.S.A.
would like to thank Mr. V. B. Zaykovsky,

Mr. G. Makhnin, and Mr. G. I.
Zveryansky of the Railway Antiplague
Service, Alma-Ata, Kazakhstan for thefriendly help during the 1991 field season.

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