I

Asiatic Herpetological Research

February 1992~

Historical Biogeography of theVol. 4, pp. 76-98

Phrynocephalus Species of the USSRNATALIA B.ANANJEVA AND BORIS S.TUNIYEV 2
1Academy of Sciences, St. Petersburg, Russia^Caucasian State Biosphere Reserve, Sochi, Russia^Zoological Institute,Some species inhabit
andindeserts.
hard
soil
and
while
othersoccur
present day distributions are
gravelSpeciation
sandy deserts,
a result of geologic events such as mountain building causing the isolation of valleys and basins andchanges in the direction of river courses.
Abstract. -There are 22 species and subspecies of Phrynocephalus in the USSR.

Key words:

Reptilia, Sauria, Agamidae, Phrynocephalus, Armenia, Azerbaijan, Kazakhstan, Kirgizistan,

Russia, Tadjikistan, Turkmenistan, Uzbekistan, USSR, biogeography, distribution, evolution.

possible ways of the development andformation of the distributions of different

Introduction

The reconstruction of the genesis of
Phrynocephalus species can not be basedon the paleontological evidence since datain this field are extremely poor. The only
fossil locality is from the Pliocene ofTurkey (Zerova and Chkhikvadze, 1984).
a comparison of recent
arealographic patterns of species studied
with known ideas about the historical
geography and paleogeography of theregion where the species occur may be used
as one method of research. The data aboutclimate and the genesis of landscapes andvegetation are of great importance.

However,

forms independendy from the disagreementon the opinion about their taxonomic status.
We discuss here the information about theranges of these lizards and although thestatus of some of them may be problematic,
this does not so strongly influence our
speculations about historical biogeography.

Most Phrynocephalus species inhabit
Middle Asia and Kazakhstan territories.
Only a few species penetrate to the easternpart of the northern Caucasus and eastern
Transcaucasia. Phrynocephalus versicolor
kulagini extends to the southern regions ofTuva Autotomous Republic, Russia (Fig.
1).

Accordingandto recent ideas,

subspeciesofPhrynocephalus

live(Bannikov

et

al.,

1977;

22 species

the
in

the

USSR

Borkin andDarevsky, 1987). We use here the last list
and do not try to reflect more recent andoften opposing ideas about Phrynocephalus
taxonomy (Golubev, 1989a, 1989b;
Mezhzherin and Golubev, 1989; Semenov,
1987; Semenov and Dunayev, 1989;
Semenov and Shenbrot, 1982, 1990;
Shenbrot and Semenov, 1987), and about
the status of some populations, subspecies,
and species. This paper does not consider
problematic nomenclature problems thatneed special research. We try here tounderstand the present complicateddistribution of

USSR.

Phrynocephalus

This includes

populations.

theirDiscussion

genus

in thedisjunct

We also try to explain the© 1992 by Asiatic Herpetological Research

As is well known, it is impossible tounderstand the history of the fauna without
knowing the history of the flora andclimatic and geological evolution. The
history of the flora in the Caspian region is
known from the Upper CretaceousPaleogene (Korovin, 1961) when the

Tethys transgression flooded most of
Middle Asia and central Kazakhstan. In
this region a continental regime wasretained only in the Tien Shan area and in
the eastern part of the Kazakh hummock
topography (Gvozdezky and Mikhailov,
1987), (Fig. 1). At that time the Central
Asian land already had an arid regime(Agakhanyanz, 1981). The middle of theGobi was probably real desert surroundedby steppe landscapes (Serebrovsky, 1936).

February 1992

FIG. 1.

Asiatic Herpetological ResearchVol. 4, p. 77

Sea basin and land (hatched) in Middle Asia and Kazakhstan in Upper Eocene - beginning of the

Oligocene.

1- the Lower Cretaceous records of plants in Er-Olian-Duz Depression (Serebrovsky, 1936); 2-

Eocene - Oligocene records of reptiles and amphibians in Zaissan Depression (Bakradze and Chkhikvadze,1988).

A warm tropical and subtropical climate,humid, but from time to time with seasonalaridity and probably with seasonal falls oftemperature, was dominant on theparts of Middle
andMikhailov, 1987).
(Gvozdezky

continental

Asia

It may have promoted the growth ofsuch species as Taxodium distichum,
Populus balsamifera, Juglans acuminata,
Fagus antiposii, Liquidambar europaeum,
Diospiros sp., Gingko biloba, andLiriodendron tulipifera. The analogous
flora was distributed throughout all of theSiberian territory (Serebrovsky 1936).

Luxuriant thermophilous flora of Middle
Asia was accompanied by a highly rich anddiverse fauna of the Late Cretaceous suchas salamanders and frogs from the families
Scapherpetontidae, Batrachosauroidae,
Prosirenidae and Pelobaudae (Bakradze andLizards of the
Chlihikvadze, 1988).
families Parasaniwidae, Teiidae, Anguidae,
Agamidae, Saniwidae, Gekkonidae, andVaranidae
were also present (Nesov,
1981a, 1981b).

From data about the Cretaceous flora insouthern Middle Asia, we hypothesize thepresence of a more dry and hot climate

Vol. 4, p. 78

Asiatic Herpetological Research(Korovin, 1961).

The Lower Tertiary findings in the ErOilan-Duz Depression in Badkhyz contains
Dryrandra schrenkii, Celastrophyllum
turcmenicus and other typical xerophilous
plants which also indicates adaptation ofthese plants to the survival during the hot
and dry periods during the vegetation

period (Serebrovsky, 1936), (Fig.

1).

From the Eocene fossils of the family
Agamidae were found in some localities in
Kazakhstan including the Zaissan Basin(Bakradze and Chkhicvadze, 1988).
In the Oligocene the sea retreated and theformation of a continental landscape began.
However, the development of relief took

February 1992

Paleogene-Neogene time in arid regions ofCentral Asia and in the southern part of
Middle Asia (Fig. 2). Until middle or late
Pliocene the herpetofauna of Central Asia
and Turan represented, more or less, asingle unit (Chkhikvadze et al., 1983). The
independent formation of Central Asian andMiddle Asian centers of different faunabegan readily after their separation by
mountain structures of Alpic orogenesis(Ananjeva, 1986; Chernov, 1959; Geptner,
1938; Peters, 1984).
In discussing the center of origin of thegenus Phrynocephalus, Moody's (1980)
opinion should be noted. He suggested
that themost primitive Phrynocephalus

P. vlangalii inhabiting northspecies
eastern Tibet and Qinghai. The validity ofisbecause heplace in different ways. In the Kara-Kum
and Kyzil-Kum deserts and the Ustyurt

this conclusion is problematic

Plateau, anticlinal and synclinal structures
were formed in the Neogene. In spite ofhis phylogenetic and biogeographic studyof agamids.

studied only two species of this genus in

their platformal type of structure, they are

sufficiently sharp with an angle ofdeclination of more than 10°. The Turgai

Plateau and western Betpak-Dala wereslightly touched by the most recent
orogenetic movements on these small
territories. This not only resulted in theformation of different geomorphologicalstructures which will be discussed below,
but also in the difference of the amplitude
raising and sinking of wholeterritorial divisions. These differences in
the amplitude of movements have resultedof thein the isolation and formation of the specificrelief in each of the plains in Middle Asia

(Voskresensky, 1968).
In themiddle Oligocene there was a

sharp change in the composition of theherpetofauna of the Zaisan Depression.
The early Oligocene giant salamanders(Zaissanurus), giant snakes (Boidae),Glyptosaurinae, etc. were replaced byamphibians of the families Pelobabidae,
Ranidae, and Bufonidae and by boids ofgenus Bransateryx (Bakradze andChkhikvadze, 1988).
theOf special interest for understanding theorigin of Phrynocephalus is the finding of anew

Phrynocephalus
species,langwalaensis (Sharma, 1970), from theRadjastan Desert, in western India.

Whiteman(1978)

suggestedthat

Phrynocephalus probably originated in the
early

On

figure12) illustrating the hypothesized

Miocene in Central Asia.
Whiteman's map (Whiteman, 1978:

his

dispersal of Phrynocephalus, he showed
this point in southern Middle Asia. The

reason of such term confusion is connectedwith the absence of separating, in English,
the terms Central Asia and Middle Asia
traditionally used in German and Russian
geographical and zoological literature.

Middle Asia is the part of Asian territoryof the USSR from the Caspian Sea in thewest to the Chinese border in the east, andalso from the Aral-Irtysh drainage divide in
the north to the border of Iran and
Afghanistan in the south.
Central Asia is defined as the desert and

The formation of two centers ofspeciation of the genus Phrynocephalus
probably began on the boundary of thesemidesert plains, tableland and highplateaus which are limited to the east by thesouthern part of Great Khingan andFebruary 1992

Asiatic Herpetological ResearchVol. 4, p. 79

Sea basin and land (hatched) during the Lower - middle Miocene. Hypothesized centers ofPhrynocephalus speciation: A- Middle Asian center; B- western edge of the Central Asian center. Lower
middle Miocene records (Bakradze and Chkhikvadze, 1988): 1- northern Aral Sea region; 2- Turgai; 3Zaissan Depression. The arrow shows the hypothesized direction of movement of ancestral forms of the

FIG. 2.

Phrynocephalus guttatus complex.

Taikhanshan ridge and to the south by thelongitudinal tectonic basin of the upper
Indus River and Brahmaputra (Tsangpa).
In the west and in the north the border ofCentral Asia corresponds to the mountainridges of eastern Kazakhstan, Altai,
western and eastern Sayan, approximately
coinciding with the state border between the
USSR on the one hand, and China andMongolia on the other hand.

Eremias sp., Varanus,

Ophisaurus,

Eryx, and Protestudo were found in the late
Miocene deposites in eastern Kazakhstan(Bakradze and Chkhikvadze, 1988). The
ancestor of Phrynocephalus maculatus may
have already existed during the Neogene in
Middle Asia in the condition of southernsavannas and xerophytous vegetation of thesouthern and southeastern Transcaspian
region. The ancestral form of P. raddei(Fig. 3) may have already been distributed
along all the southern part of the Thetys
geosyncline from the Caspian Sea to the
Pamir. We can hypothesize this because
fossil remains of giant land tortoises andmonitors are known from the Pliocene in
Tadjikistan.

Somelizards,

TrapelusVol. 4, p. 80

FIG. 3.

Asiatic Herpetological Research

Sea basin and land during the Upper Miocene (Sarmat Sea is hatched).

February 1992

1-

Hypothesized

distribution of Phrynocephalus maculatus ancestor; 2- Hypothesized distribution of Phrynocephalus raddei

ancestor.

The arrow shows the direction of continuing dispersal of the forms of the Phrynocephalus

guttatus complex.

sanguinolentus, Eremias sp., and Varanus
Pliocene incf. griseus are known from the
Turkmenia (Ananjeva and Gorelov, 1981;

Bakradze and Chkhikvadze, 1988)
the
In
the
Pliocene,genus
Phrynocephalus could have divided into
species complexes or into the genera
Phrynocepnalus and Megalochilus(Ananjeva, 1986) on the territory of thesouthern Kara-Kum Desert. Federovitch(1946) assumes that one should look forthe origin of typical recent sandy desert
vegetation associations in the Neogene in
the Kara-Kum (Fig. 4).

The Miocene may be considered as thetime when the ancestral form of the
Phrynocehalus guttatus complex (Figs. 5
and 6) from the Central Asian center
penetrated as far as the eastern boundariesof the Tethys (recent regions of Pamir- Alaiand Gissar-Darvaz mountains), (Fig. 2).
This territory, now occupied by mountains
and intermountain depressions, resembled
low mountain relief now present northwestof Samarkand and Djizak (Voskresensky,

The subsequent dispersion of thisgroup to the west was along the northern1968).

shore of the Thetys (later the Sarmat Sea,
Fig. 3). Further spreading to the north wasFebruary 1992

Asiatic Herpetological ResearchVol. 4, p. 81

FIG. 4.

The middle Pliocene changes of Phrynocephalus. 1- Lower Pliocene distribution of
Phrynocephalus reticulatus; V- middle Pliocene populations of Phrynocephalus reticularis on emerged
land; 2- Lower Pliocene distribution of Phrynocephalus maculatus; 2'- middle Pliocene populations
separated by alpic orogenesis of the Kopeth-Dag. The arrows show the direction of dispersal of the species
A- hypothesized place for divergence of Phrynocephalus andBof
Megalochilus;
hypothesized place
origin for Phrynocephalus rossikowi.
of the Phrynocephalus guttatus complex.

prevented by phytogeographical conditions
since the plains of central Kazakhstan werecovered by deciduous forests of Populussp., Salix sp., Alnus sp., Zelkowa sp.,

Ulmus sp., Acer sp., i.e. the vegetation
was intermediate between the Turgai andrecent types. The xerophilous formations
were only beginning to develope in thisterritory (Gvozdezky and Mikhailov,
1987).

During the first half of the Neogene, a
was predominate on thelake regime

elevated plains of Middle Asia such as the
Turgai tableland and western Betpak-Dala.
The southwestern branch of the P. guttatuscomplex dispersion could have penetrated
this area.
The process of uplifting took
place across the entire plains of Middle Asia
towards the end of Sarmatian time and tothe beginning of the Pliocene. The sea
basins disappeared and erosional divisionof the region took place (Voskresensky,1968). During this period, Phrynocephalus
reticulatus could have dispersed widelyover the entire plain area from the CaspianVol. 4, p. 82

Asiatic Herpetological Research

February 1992

FIG. 5. Phrynocephalus guttatus from the west side of the Caspian Sea in Chechen-Ingush, Russia.

Sea in the west to the Fergan Depression inThis species apparently did notreach beyond the limits of the dry subtropic
climatic belt. The same is observed at theIn such context, thepresent (Fig. 4).
opinion of Golubev (1989b) on the unity ofthe origin of P. moltschanovi and P.
reticulatus from the forms penetrating here
from the north in the middle Pleistocene
seems to us doubtful.
the east.

The eastern branch of the P. guttatuscomplex, i. e. Phrynocephalus versicolor
was widespread north and northeast of the
Tien Shan (Mountains), (Fig. 4). Isolationof Phrynocephalus rossikowi (Fig. 4)
could have taken place on the dense river
sediments of the Amu Darya (River) which
flowed into the Caspian Sea at that time.
The Pliocene raising of Asia Minor and the
Iranian Plateau had apparently already led
to disjunction of the area inhabited by theancestor of Phrynocephalus helioscopus
and also by some species of the genus
Trapelus with similar ecologicalrequirements. The diverged populations of
P. helioscopus, P. helioscopus persicus (P.
persicus, Meszszerin and Golubev, 1989;
Nikolsky, 1915), could probably haveseparated in the Pliocene. In the middle of
the Pliocene a sinking process occured in
the sand deserts of Middle Asia to theslightly elevated Zaunguz Plateau.
However, on the Turgai Plateau in western
Betpak-Dala and on the Ustyrt Plateau theraising of the Kysil-Kum and Mangyshlak
was no longer restored by the regime of

The relief continued todevelope slowly by an erosion andaccumulation.

denudation process (Voskresensky, 1968).
It was in the middle of the Pliocene that the
disjunction of the continuous range of
Phrynocephalus reticulatus took place.
That led to isolation of three relict

Asiatic Herpetological Research

February 1992

Vol. 4, p. 83

**$,.

*
FIG. 6. Habitat of Phrynocephalus guttatus on the west side of the Caspian Sea in Chechen-Ingush,
Russia.

populations on the plateau islands notrelease themselves from the captivity of thecoveredsand during movement of sand dunes. It
should be noted that ridges formed from thestabilized aeolian landforms in the
Pleistocene when mountain structures,
which mainly determine the direction of the
air streams, were formed.
It is assumed
that the direction of sand movement

South Ustyurt
by sea:Krasnovodsk, Kiysil-Kum and Fergana

Depression (Fig. 4).

The formation of the Phrynocephlalus(orcomplexgenus

mystaceus

Megalochilus), (Figs. 7 and 8), and of theparallel sand inhabiting P. interscapulariscomplex, continued in the extreme southern
portion of Middle Asia under conditions ofsandy desert formation. The adaptive
radiation of Phrynocephalus in sympatry,according to Peters (1984), could havebeen accompanied by increasing differencesin the body size. This seems to have been
important in the evolution of P. mystaceus.
On quick moving sand dunes with steep
slopes, the largest specimens could
survive.
They were able to dig
uncrumbling deep holes protecting themfromsummertemperatures.

heat and low winter
They were also able toremained the same at least to the Upper

The
Neogene (Voskresensky, 1968).
withformedmesoandplainsnewly
microrelief created before the Quarternary

drying and cooling by wind activity were
alluvial plains with all the typical features(Voskresensky, 1968).
P.

mystaceus evolved under theof blowing sand.
The

conditions

decreasing of body size of P. mystaceus
in comparison with P.
mystaceus galli may be indirect evidence infavour of this hypothesis. There may becorrelations between this change of body

mystaceusVol. 4, p. 84

Asiatic Herpetological Research

February 1992

FIG. 7. Phrynocephalus mystaceus from Repetek (38° 34' N 63° 11' E), Turkmenistan.

and the inhabiting of P. mystaceus
mystaceus in the comparatively stabilized
Terek-Kuma rivers sands on the west sideof the Caspian Sea in Russia. Smaller sizes
sizeThis formof a unique
under
the
conditions
developed
isolated sand dune with a special wind

Dune, Dagestan, Russia.

regime (Khonyakina, 1962).

of specimens of P. mystaceus are typical of
the populations

from the Sari-Kum Sand

The

restricteddistribution

of

P

.

Asiatic Herpetological Research

February 1992

Vol. 4, p. 85

FIG. 8. Habiat of Phrynocephalus mystaceus (large sand dunes), Phrynocephalus interscapularis (sanddune edges), Phrynocephalus raddei (hard packed soil to the left) from the Kara Kum Desert 80 km north of
Ashkabad (37° 57' N 58° 23' E), Turkmenistan.

mystaceus andpoor foodinterscapularis may be explained by someavailability of the sand dunes could have
driven the small P. interscapularis back toecological peculiarities. This lizard is verysmall. It is not capable of digging deep into
the sand, and it also has a greater tolerance

the dune valleys.

commonrelatively

This species, sharing a

origin with P
mystaceus, could not spread beyond thelimit of the subtropical climatic belt during
Its present
the next geological epoch.
distribution almost completely lies in the
climatic zone of the continental southern
Turanian region with a small penetration to
the extreme southern part of the continental
north Turanian region (after Alisov, 1969).
These species occur were the temperatureduring January -3°C in the north, up to 2°C
in the south and annual precipitation from100 to 200 mm. According to Kashkarov

southern.

and Korovin (1936) P. interscapularis
Mediterranian deserts with awinter-spring period of precipitation andinhabits

vegetation of the ephemeral type (Table 1).
The relatively restricted range of P.

to high temperatures.
The temperature
preference of P. interscapularis may
fluctuate only 3°C, whereas in P.

mystaceus it may fluctuate 4°C and in
Eremias grammica up to 5°C (Cherlin and
Muzicnenko, 1983).

The mode of preference of temperaturesin P. mystaceus and Eremias grammica is

39°C which may be comparable with thevery high level of tolerance known for
Dipsosaurus dorsalis. However, for P.
interscapularis this index is still higher
This may be considered an(41.3°C).
outstanding example of adaptation of asmall lizard to extremely arid conditions.
In the north and northwestern deserts, P.

Asiatic Herpetological ResearchVol. 4, p. 86

February 1992

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Asiatic Herpetological Research

February 1992

mystaceus occurs on steep slopes of sanddunes were as the sympatric P. guttatus
occurs in the valleys between sand dunes.
Large mountain ranges were formed inplace of residial plains and low
mountain relief with a Paleozoic folded
thestructure. The peneplain was subjected tofolded deformations (Gvozdesky and
Mikhailov, 1987). In this context it isdifficult to share Golubev's (1989b)
opinion that the conditions of the early
Pleistocene were favorable for the
penetration of Phrynocephalus from Central
Asia to eastern Kazakhstan. However,Vol. 4, p. 89

have occupied all of Middle Asia, from the
Caspian Sea in the west to Balkhash Lakein the east, and from the plains near theslopes of Kopeth-Dagh and Hindu Kush in
the south up to the Naryn Sands in thenorth.

Its

primary Aralo-Caspianand distribution isallsupported byzoogeographers(Anderson, 1968; Chernov, 1954;
Rustamov and Sczcerbak, 1985;
Vereshagin, 1966). The penetration to thedeserts north of the Caucasus Mountains(Turanian)originprobably took place around the northern
Caspian Sea (Chernov, 1954; Darevsky,
1957).

Golubev (1989b) noted that owing to active
alpic orogenetic processes in the Junggar
Alatau and the Tian Shan, the contact of thereptiles between the Balkhash-Alakol and

the Junggar Depression became graduallymore difficult or was interrupted. This
period wasprobably characterized by

disjunction of continuous ranges and by theisolation in the intermountain depressionsof the central branch of the P. guttatuscomplex. Further history of the speciationof each population seems to have resulted
in the formation, during the Pleistocene, of
more clearly isolated taxonomical forms in
different depressions:Hi- (P. alpherakii
Peters, 1984; Golubev,
1989b, or P. versicolor paraskivi according

accordingtoto Semenov, 1987); Alakol Depression- (P.

versicolor ssp.), and Zaissan Depression(P. salenskyi according to Peters, 1984;

Golubev,

1989b,orP.

melanurus

according to Semenov, 1987). Definitive
formation of eastern populations of the P.
reticulatus complex, which subsequently
led to the isolation of P. strauchi, may havetaken place at the same time. Its speciation
occured because of the isolation of the
Fergana Valley. Retaining of isolated

populations in intermountain depressions
was possible due to the absence of repeatedleveling of the relief in Neotectonic time.
This is supported by geological evidence
such as the composition and thickness of
sediments in the Hi and Fergana
depressions (Voskresensky, 1968).

Most of the distribution of P. mystaceus
seems to have been formed during the endof the Neogene when these lizards could

In the Pleistocene, the Central Asianelements of the flora dispersed from theeast to the north of Middle Asia and central
Kazakhstan. The other important center
which influenced the development ofMiddle Asian vegetation was the eastern
Mediterranian center (Gvosdezky and
Mikhailov, 1987). All the plains of thisextensive region are classified by vegetation

type into two kinds of deserts:
Mediterranian (after Kashkarov andKorovin, 1936), and subtropical deserts of
the northern zone (after Gvosdezky and
Mikhailov, 1987), or accordingly, thedeserts of the northern zone (after
Gvosdezky and Mikhailov, 1987). The
present boundary between the two zones
approximately corresponds to the boundary
distinguished by Alisov (1969) for the
climatic regions of continental northern

Turanianandcontinental

southern

Turanian.
In the Pleistocene, speciation of the P.
guttatus complex occured in the northern
deserts of the Caspian and Aral Regions.

During the Upper Pleistocene, river beds of
the Amu-Darya and Syr-Darya rivers turnedto the Aral Sea (Voskresensky, 1968).
This resulted in a change from sandmassive on the southern coast of the Aral
Sea to loess and clay plains of river origin.
Under these conditions of hard soils, theformation of the isolated southwestern
population of the P. guttatus group,
considered now as a separate species, P.
moltschanowi (Semenov and Shenbrot,1982) took place (Fig. 9). The taxonomic

Asiatic Herpetological ResearchVol. 4, p. 90

February 1992

FIG. 9. Upper Pliocene changes of Phrynocephalus. A- middle Pliocene distribution of the species of the
Phrynocephalus guttatus complex. The disjunction of the continuous distribution of this complex occuredin the Upper Pliocene by: B- Junggar Alatau (Altai mountains); C- Saur and Tarbagatai mountain ridges.
1- Phrynocephalus melanurus; 2- Phrynocephalus versicolor ssp.; 3- Phrynocephalus versicolor paraskivi;

4- the place of speciation of the southwestern branch of the Phrynocephalus guttatus
Phrynocephalus moltschanovi.

statusof thisspecies

was discussed

recendy by Golubev (1989b).
It should be noted that the flow ofMiddle Asian rivers to the Caspian Seachanged to the Aral Sea, as a result of thedownwarp of the region (Voskresensky,1968). This was of crucial importance forthe reconstruction of the ranges of mostsclerobiont Phrynocephalus species.
Probably before the Amu-Darya River(Uzboi), changed its course P. raddei wasdistributed on the clay and loess ares fromthe Caspian Sea to Kukhistan. Its range

complex anddecreased considerably from the north andfrom the south owing to orogenesis (Fig.
9). In the south this species remained on
the incline plain of the Kopeth Dagh and in
the loess regions in the estuaries of the

Murgab and Tedjen rivers.
After the Amu Darya changed its flow to
the Aral Sea, which coincided in time withthemaximal development of the sanddeserts, the area inhabited by P. raddei wasa number of isolated
include the piedmont
These
populations.
andthe Kopeth Dagh,in
Kukhistan
plainsdividedintoFebruary 1992

Asiatic Herpetological ResearchVol. 4, p. 91

FIG. 10. Pleistocene changes of Phrynocephalus. 1 to 1 '- the river course of the Amu Darya River (Kelif
Uzboi and Uzboi) flowing in to the Caspian Sea before the Lower Pliocene. The distribution of
Phrynocephalus raddei before the change of the Amu Darya River course to the Aral Sea is hatched. The
relictual populations (Upper Pliocene to present) after the change of the

Amu Darya River course to theAral Sea are depicted by cross hatching. A- present distribution of Phrynocephalus guttatus kushakewichi;
B- the transformed distribution of Phrynocephalus rossikowi during the Upper Pleistocene from prePleistocene center of speciation (B').

remaining loess and other valley originatedforms of the mouth of the Murgab andTedjen rivers, and the dry bed of the
Uzboi. P. raddei boettgeri was formed in
the eastern isolated part and it is possible

correlated with the change of course of the

Amu Darya River. This area decreased in
the south and reached the southern coast of
the Aral Sea in the north (Fig. 10). Owing
to the constant change of the configurationthat the western populations also present
combinations of different forms (now thenominative subspecies P. raddei raddei,of the Amu Darya estuary also observednow (Voskresensky, 1968). The northernpart of the range of P. rossikowi was also

(Fig. 10).

changing repeatedly, resulting in theisolation and long existence of this isolated

The genesis of the area inhabited by P.
rossikowi (Figs. 11 and 12) is also

western population.
recentlyasIt

a distinct

was describedsubspecies, P.

Asiatic Herpetological ResearchVol. 4, p. 92

FIG.
(39°

1 1 .

15'

Phrynocephalus rossikowi (size x 2) from along the Amu Darya River, 30 km
1 1' E), Turkmenistan.

WNW of Deynau

N 63°rossikowi shammakovi. It is also possiblethat the extreme northeastern population has
been isolated from the main distribution fora long time and represents a distinct
taxonomical form.
After the formation of sand ridges andthe deeping of the dune valleys slowed, theof washing away the subtle
material from the ridges to the valleys withthe formation of the "takyr" landscapes

processbegan

February 1992

in the Pleistocene(Voskresensky,

Under such new conditions P.
helioscopus became widerspread in the1968).

plains of Middle Asia.

The Pleistocene glaciation in Europe
resulted in the sharp displacement ofvegetation zones in the southern Russian
plains and vegetation belts in the Caucasus
Mountains. As a result, the distribution of

P. mystaceus

in the deserts northof theCaucasus Mountains was separated into a
number of isolated parts.
Its range
increased in the piedmont regions north of
the Caucasus Mountains to the westward,

probably in the postglacial xerothermic timeof the Holocene. At this time P. mystaceus
reached the present border of Dagestan andStavropolsky Territory along the Terek-

Kuma rivers sands.
During the Pleistocene, P. mystaceus
and P. interscapularis dispersed into
mountainous Kukhistan along the sands
formed from the alluvial of the Amu Darya(River). Phrynocephalus sogdianus
evolved as a result of the disjunction of theKukhistan enclave during the Upper
Quaternary from the continuous range of P.
interscapularis. This species was describedby Chernov (1959) as a subspecies, P.
interscapularis sogdianus. This form wasgiven the status of a distinct species after

Asiatic Herpetological Research

February 1992

Vol. 4, p. 93

FIG. 12. Habitat of Phrynocephalus rossikowi (size x 2) from along the Amu Darya River, 30 km

WNW

of Deynau (39° 15' N 63° 1 1' E), Turkmenistan.

Sokolowsky(1975)

discovered

considerable karyotypical differences
betweenP.
sogdianus and P
interscapularis. With the alternations of the.

interglacial

The Khvalynskaya transgression of theepoches, the pluvial and xero-thermic
climatic
periods were connected.
However, during the whole Quaternary

Caspian Sea defined the western part of therange of P. raddei. The flooding of theSary Ishik Otrau sands near the southerncoast of Balkhash Lake resulted in thealmost total disappearence of P. guttatus
and P. mystaceus in this region. They areretained probably only near the foot of
isolated island mountains that have risen

Quarternary

glacialanddisappeared in the middle Quaternary
period from the Muyun Kum Desert. After
which only P. mystaceus could inhabit it.

period, the climate was sharply continental.
The desert or desert steppe (in pluvial

epoches) regime was retained on the plainsof Middle Asia (Gvozdezky and Mikhailov,
1987).

the

Quaternary, the lastaccumulation changing of the relief on theplains in Middle Asia took place. This may

Duringexplain the present configuration of the
The last
ranges of desert animals.
considerable accumulation (Khvalynskaya)
included the Caspian and low land KaraKum Desert, Muyn Kum and Sary Ishik

Otrau.

Toad headed agamids completely

recendy among the sands in the eastern partIshik
of Sary
Otrau (Fig.
10).
Subsequently, dispersal from these refugia
and isolated areas could have led to theformation of P. guttatus kuschakevitschi inthe Balkhash sands.

Thus, the history of the formation of
Phrynocephalus distributions, which is thesclerobionts (hard soils) depressions andAsiatic Herpetological ResearchVol. 4, p. 94

the blooms of psammobionts, is correlatedwith step wise development of the sanddeserts of the Middle Asian plains. It was
influenced by geological processes. All
this wide belt is correlated with the zone ofFebruary 1992

by special valves; 4) special "scaled"eyelids close the eyes; 5) tympanum absent
or hidden under the skin; 6) granularmost recent downwarp along the peripheralsmooth scales; 7) comparatively high speedof locomotion, sometimes bipedal; 8)
increased finger surface, "sand ski"; 9) thepart of the mountain massives of Middle
Asia with the accumulation in them This

capacity to bury into the sand; 10) shortrecurveable tail.

was followed by transformation of riverand estuary sediments under arid

However a more detailed study of thesecharactersconditions.
In some Phrynocephalus groups, species
secondarly inhabiting sands are known,(Semenov, 1987). For example this patternobserved in the P. guttatus complex,
apart from typical sclerobiont forms.
Chernov (1948, 1959) noted that P.
isrefutesthesimplified
determination of their correlation with ahabitat in the sand. It is doubtful that thedorsoventral depression of the body may be
an indicator of inhabiting sand biotopes.
This character is in the basis of theidentification key in the Agamidae family(Boulenger,

1885),

separating

more

guttatus inhabits different types of sandy
biotopes from P. mystaceus and P

specialized tree agamids from all terrestrial,
rock and desert forms. It is well known

interscpularis.

such form of the body of mountainagamids of the genus Stellio is not anindicator of their origin in sand deserts.

.

In

that

connection

with

the

problems

discussed, it is necessary to mention the
problem of Phrynocephalus origin. As was
already stated above, paleontological data
are available for Phrynocephalus only fromthe Pliocene of Turkey. This is not enough

data
about
fossilsignificant
The
data
of
Phrynocephalus.
present
distribution and life history allow us to
speculate about the primary character of
habitats typical of these lizards. Golubev(1989b) wrote, correctly in our opinion,
about the development of the most primitive
Phrynocephalus in gravel and sand-stone

Most herpetologists(Gobi) deserts.
(Chernov, 1948; Semenov, 1987; Whitman,1978) suggest that the primary character issand biotypes.
Thus, Chernov (1948)
assumed that Phrynocephalus originallyadapted to sand habitats and then moved tohard soils. The same opinion is shared by

Whiteman (1978) and Semenov (1987).

It

interesting that all these herpetologists
use as the most serious argument, theisSuch characters as closed upper lipcovering the mouth, nostrils closed by
special valves, and special "horny scaled"eyelids closing the eyes undoubtedly may
be considered as defensive structures.

However they can be developed in different
kinds of deserts (not only sand deserts)
with a typical windy regime.

Considering the ideas about the origin
oiPhrynocephalus and the so called"primary substratum" one may assume thatthe terms "desert" in general and sanddesert are sometimes confused.
Thus,
Chernov (1948, p. 132) was absolutely
right that Phrynocephalus "originated anddeveloped in the conditions of rather sparseThis, however, does notvegetation."
thesepermitcomplexes of landscapes andsand desert to be considered as equal. The
latter is only one type of desert and it is the

youngest from a geological aspect.

number of morphological adaptations
shared by all species of Phrynocephalus
and are typical of many other lizards

adapted to

life

in

deserts.

Whiteman(1978), refering to Stebbins (1944), names
the following morphological characters: 1)
dorsoventral depressed body form; 2)
protruding labial border; 3) nostrils closedThe

character, tympanum absent or
hidden under the skin, is of special
importance. Analysis of the distribution ofthis character and trends to the reduction of
the middle ear among all the agamids,
shows that it has arisen independently insome evolutionary lines. The loss of theFebruary 1992

Asiatic Herpetological Research

tympanum and tympanum cavity is typicalof the Australian genus Tympanocryptis.
Such reduction may arise even in primitive
forms like Ceratophora, Cophotis andLyriocephalus. These convergent trends
are noted in Otocryptis, Phoxophrys,
Phrynocephalus, and Ptyctolaemus(Moody, 1980). The enumeration of thesegenera shows that side by side with thedesert lizards (but not psammophilous)
there are even forest species.
Thus
is
a
terrestrial
lizardwhich
Otocryptis
prefers to inhabit the vicinity of rivers
shaded by vegetation in India and Sri
Lanka. With this consideration, furtherofthe
new form,
Phrynocephalus laungwalaensis from theRadjastan Desert of India may be

examination

important.

The granular smooth scales also may beobserved not only in psammophilous
agamids but also in Leiolepis, Uromastyx,
and Physignathus (Moody, 1980).
Besides, an examination of the correlationof Phrynocephalus
morphologicalcharacters and the type of substrate
shows
that
1974)
(Galayeva,
have
psammophilous species
granularsmooth scales. Where as lizards inhabitingVol. 4, p. 95

explanations of their function (Chernov,
1948; Fausek, 1906, 1959; Luke, 1986;

Werner, 1987). These structures are really
typical of many sand lizards, but simple
character environment correlation may be
misleading (Luke, 1986; Smith, 1935).
Toe fringes have arisen independently atleast 26 times in seven families of lizards(Luke, 1986) and can be used forlocomotion on shifting sand, across water,
and for digging in some kinds of soil suchas sand and loess (Chernov, 1948; 1959;
Luke, 1986).

The original capability of burying into
the sand with horizontal movements of thewhole bodyvery well expressed in
psammophilous P. mystaceus and P.
inter scapular is. There exists an opinion
that such behavior may evolve only onlarge areas of moving sand (Fausek, 1906).
But one should not excluded thedevelopment of such interesting defensive
behavior on the loose sand from theelements of cryptic and or sit and wait
behavior with similar patterns observed in
P. helioscopus on hard soil. This speciesispresses the depressed widening body to theground with several horizontal movements
before standing still.

hard soil (rock debris desert or arid desert
with clay soil) usually have somewhat
enlarged, imbricate scales and small

seems

protuberances. These data show that there

considerations.

are gradations of morphological charactersworld of the sand deserts and that of thedeserts with hard soils are two formations

among Phrynocephalus from sclerobionts
to psammobionts, and not indisputable
psammophilous morphological adaptations
whole genus.
The possible

In general, the idea of Geptner (1933)

tobefruitful

such

for

He thought that the animal

different in many aspects with their own, toa considerable extent independent, ways ofTheof thedevelopment.

functional importance of small granular
scales in the capillary collection andtransport of the water in many desert lizards

adaptations in the two kinds of landscapesshould be noted (Schwenk and Greene,

To summarize the review of the recent1987).

The comparatively high speed oflocomotion, sometimes bipedal although
the limbs may be weak, are typical of many
agamids inhabiting open areas (Sukhanov,
1968; Cogger,1975) and can not be

restricted to psammophilous species only.

The increased toe fringes are widely
discussed but there are no good

purposeoftheis considerably different.

chorology of Phrynocephalus in the USSR
fauna, it should be noted that thedifferences in the outlines of the
distributions in general correspond to two

The species ofCentral Asian origin have the northern mostmain centers of origin.

distribution, inhabiting totally a moderate
climatic zone and the species of Milddle

Asian origin mainly did not go beyond thelimits of the subtropical climatic zone. The
relatively young species (Phrynocephalus

Vol. 4, p.

%

Asiatic Herpetological Research

February 1992

helioscopus, and P. mystaceus) have thewidest distribution. A large portion of their

Proceedings of the Zoololical Institute, USSR
Academy of Science, Leningrad 157:4-13. (Indistributions were formed in the Pleistocene

Russian).

immediate connection with thedevelopment of the sand deserts andinaccompanied takyrs.

We can distinguish several types of the

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helioscopus persicus, and P. strauchi).

The Pliocene-Pleistocene ranges:a)wide (P. helioscopus heliscopus, and P.
mystaceus); b) subtropical (Pinter scapularis, and P. sogdianus).
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Pleistocene transformed area (P.
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