Asiatic Herpetological Research

December 1993

Vol. 5

pp. 65-73

1Geographic Variation and Diversity in Three Species of Phrynocephalus
the Tengger Desert, Western China
YlTEZHAO WANG AND HUIZHAO WANG 2
1'Chengdu Institute of Biology, Accidentia Sinica, Chengdu, Sichuan, China^
Chengdu Library. Academia Sinica, Chengdu, Sichuan, China
Abstract. -Univariate and multivariate statistical analysis of 5 merisuc characters, 4 metric characters and
8 ratio characters recorded for 9 samples of lizards were used to assess non-geographic variation andA

indicated that there werecluster analysis
geographic variation in the southeast area of the Tengger Desert.
three major groups which represented Phrynocephalus versicolor, P. pnewalskii, and P. frontalis, based on
showed that amongmorphological characters, respectively. The cluster analysis and canonical analysis
and divergence
notwasDispersal
always predicted by geographic proximity.
samples, phenetic similarityof these species of Phrynocephalus with the relationships of paleogeography and paleoclimatology arediscussed. It is evident that the Yellow River fails to cause geographic isolation.

Key words:

Reptilia, Sauria, Agamidae, Phrynocephalus, China, biogeography.

Strauch (1876) described four new
species of Phrynocephalus from the Altan
Desert (including the Tengger Desert) andthe Mu Us Desert (Ordos). They were P.
pnewalskii, P. versicolor, P. affinis, andHe pointed out that P. affinis

P. frontalis.

was very similar to P. pnewalskii. Pope(1935), with out any discussion,considered P. affinis to be synonymouswith P. pnewalskii. Leroy (1939), whothestudied

variation

geographicanddistribution of P. pnewalskii, P. frontalis,and P. vlangalii, failed to recognize P.

frontalis and P. versicolor, which werequite different on morphological characters.
Therefore, his distribution of P. frontalis

Zhao (1979)
versicolor
ofdistribution
thereachedalmostinChinaPhrynocephalus
the shore of the Bohai Sea and in the Altan
Desert (including the Tengger Desert) nodistribution for P. frontalis was mentioned.
included P.

deducedIn.

that

thispaper

wediscuss

(1)

the

geographic variations and distributions ofP. frontalis, and P.
P. pnewalskii,and (2) the dispersal andtractsof the three species in thedivergenceversicolor,Tengger Desert.
Materials and Methods
Study Areas.

inwestern Nei Mongol
Autonomous Region, China. The east edge
reaches the Helan Mountains, and thesouthern edge, the Yellow River. In thesituatedIntroduction—The Tengger Desert© 1993 hy Asiatic Herpetological Researchwest and north, the study area connectswith the Badain Jaran Desert and the Ulan

Behou Desert, respectively. The climate inThethese areas is arid-continental.
istopography
gobi.

moving sand dunes andIn this area the vegetation is scarce.

Potaninia sp.,sp.,
Teraena sp., and Caragana sp. are present.

Ammopiptanthus

Populations of P. pnewalskii, P. frontalis,and P. versicolor were sampled from nine
localities in the southeast area of theTengger Desert from the south and northshore of the Yellow River, throughsoutheast Tengger Desert to Hala Woo
Valley of the Helan Mountains, and in thewest at Xial Hong Shan (Fig. 1 and Table1).

Methods.

—

Five meristic characters,and eight ratio
characters
metricfour
characters were chosen for study.
characters are shown in Table 2.

These—For analysis of
Statistical analysis.
were groupedlizards
variation,
geographic
into ninesamples(Fig.

1and Table1)

within which gene flow was assumed tooccur freely. Sexual variation in metriccharacters

was examinedinsample1(representing P. pnewalskii), and samples
is2 and 3 (representing P. frontalis).

We

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TABLE

66

1 .

Population

Asiatic Herpetological Research

December 1993

Locality data for Phrynocephalus from the Tengger Desert, western China used in this study.

December 1993

Asiatic Herpetological Research

TABLE 2. Characters used in analysis of variationin three species of Phrynocephalus.

TABLE 3.

Sexual variation in three samples ofPhrynocephalus (sample number as in Fig. 1 andTable
A. Meristic Characters

1.

NSPL

2.

NIFL

4.

NSDT
NSNT

5.

NDVT

3.

No. of supralabials
No. of infralabials
No. of subdigital lamellae on longest toe
Number of scales between nostrils
No. of ventral dark bands on tailB. Metric Characters

SVLTL
8. LFL9. LHL
6.

Snout-vent length

7.

Tail LengthForeleg length (including fingers)
Hindleg length (including longest toe)

C. Ratio characters

10.

Vol. 3, p. 67

Levels of significance are indicated.
(*=p<0.05: **=p<0.01)
1.

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68

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December 1993

TABLE 4. Variable coefficients for canonical variates

I

Vol. 3, p. 69

and II and estimated % influence of each vector fornine samples of Phrynocephalus.

Vector II (8.67%)

Vector I (77.58%)% influence

Variable

Character

NSPL

NSDT
NSNT
NDVT
SVLTTL
SVL/LFL

SVL/LHL
LNE/LN
LFL/LHL

HL/HW
TL/LFLTL/LHL% influence

coefficientscoefficients

NIFL

Variable

0.332
0.355
0.871
0.035
-0.093-0.0080.026-0.004
0.032

8.040
9.780
58.862
0.097
0.007
0.000

-0.001

0.000
0.000
0.000
0.000

0.715
0.715
-0.450-0.245-0.036-0.004-0.0430.005-0.010
0.001
0.009

0.001

0.000
0.001

0.003
0.007
0.023

-0.003

0.031

4.430
4.430
1.763
0.011
0.001
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000

TABLE 5. Characters for three species of population-groups of Phrynocephalus.
CharacterP. przewalskiiP. frontalis

Range

P. versicolor(Mean)

Range (Mean)

45.8-57.2(51.5)

50.7-71.4(61.6)

46.1-55.6(49.5)

58.4-82.1 (72.4)

53.9-69.4 (60.8)

23.3-29.5 (26.1)

76.1-101.5(86.3)
27.4-35.9 (31.4)

37.1-48.7(43.1)

43.7-59.6(51.1)

36.5-43.4 (39.1)

NSNT

2-4(3.2)

2-4 (2.9)

2-4 (2.0)

NSPL

11-17 (14.9

15-20(17.6)

13-17(15.0)

NIFL

12-17(15.1)

14-20(17.1)

12-16(14.0)

NDVT
NSDT

2-5 (2.8)

1-4(2.2)

0-3 (2.0)

24-30 (26.4)

27-34 (29.9)

21-26(22.0)

Range (Mean)

SVLTLLFL
LHL

Color on ventral surface of tail tip

Color on armpits
Color on back of body

22.7-26.9 (24.4)

darkdark

dark

grey-whitegrey-whitegrey-white

2-5 pairs of black spots

irregular spots

2-4 transverse black-longitudinal line

reddish bands

10

No. of teeth on maxillary1111

No. of teeth on dentary

111211

Cartilage

Cartilage

Posterior 1/2 is cartilage

Covered by splenial

Covered by splenial

Native not covered byIshium
Meckel's cartilage

splenial(sample 9) was separated
IIand vector I. The thirdvector
alonggroup, represented by samples 2, 3, 4, 5,6, and 8 in clustering, was plotted along thein clustering

centroid in Fig. 2.

to vector I and vector II are NSDT, NSPL,and NIFL, respectively.

Only males were used in univariate andmultivariate analysis for geographicvariation.

Characters having the greatest influence

Vol.

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-'.1-on.2
Xi

20-ea"mQ.

=00

15-

1070

Asiatic Herpetological Research

December 1993

Asiatic Herpetological Research

December 1993

80°

60°

140°

120°

100°50 ov

r- iS<mSi:Vol. 3, p. 71

MONGOLIAy"Tyyj-:-\/'

40°I.

I30 c"V,""*/ '*\1P.

,,'V-versicolor

P. przewalskii

20 cP. frontalis
1200 km_1FIG. 6. Geographic distribution of P. versicolor, P. przewalskii, and P. frontalis.

(Table 5; Wang, 1987), and showed thecharacters replacement. For example, thepro-half part of the ischiopubis in P.
versicolor is ossified and the post-half partstill is cartilage.
The ischiopubis in P.
przewalskii and P. frontalis, conversely, is
all cartilage.
Additionally, the Meckle's
notiscoveredby the splenial in P.
cartilage
versicolor but it is covered by the splenialinP.

przewalskii andElectrophoresis

alsoP.

frontalis.

showedthatP.

versicolor is different from P. przewalskii

and P. frontalis, the latter two species being
similar (R.
Macey pers. comm.).
Therefore we suggest that P. versicolor
was probably derived earlier from theancestral stock than P. przewalskii and P.
frontalis.

The distribution of P. versicolor is from
central Asia through western China to thecentral parts of Mongolia (Boblov, 1986)

and Nei Mongol (Zhao, 1978; 1975).
Phrynocephalus przewalskii occurs fromZhang Ye, Gansu Province through theTengger Desert to the west edge of theHelan Mountains.
Phrynocephalus
frontalis is found from Zhang Ye (Yao,1983) through the Tengger desert to the Mu
Us Desert (Ordos) (Schmidt, 1927) andforward to the southin the east part ofGansu Province. There is also an isolated
population at Xun Yang in Shaanxi
Province (Song,1987) (Fig. 6).
Phrynocephalus is distributed to the east toabout 1 12°E but does not reach the shore ofthe Bohai Sea as Zhao (1979) stated.
In the Early Tertiary, there was an aridsubtropical continental climate belt fromcentral Asia through the southern edge ofthe depression areas in the Tian Shan
Mountains and the north edge of the Tarim
Depression, the north part of the QinghaiXizang "initial plateau", the Qaidam
Depression, depression areas in the QilianQinling Mountains, and the south edge ofdepression areas in the Helan-Liupan

Vol.

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Asiatic Herpetological Research72

Mountains to the central part of China (Li,In some areas, desert had been1984).
forming and the ancestral stock of theAgamidae might have followed this belt toinvade the areas of the south side of thedepression areas in the Helan-Liupan
Mountains from central Asia throughwestern China. As evidence, some fossilsof Agamidae were discovered from theEocene of western and central China (Li,1984). We suggest that P. versicolor might
have derived from the ancestral stockduring that time. After the Late Tertiary
and Early Quaternary, with the uplift of theQinghai-Xizang Plateau, this aridcontinental climate belt shifted forward tonorthern regions, including the Tengger
Desert (Li, 1984). The ancestral stock ofPhrynocephalus and P. versicolor might
have followed the shift of this belt again todisperse into northern regions, including
the Tengger Desert and P. przewalskii andP. frontalis derived from ancestral stock in
the Tengger Desert during that time. ThenP. frontalis dispersed further east to theMu

Us Desert (Ordos) and south part of theQinling Mountains.

Accordingto

thedistribution of P. frontalis, it is obviousthat it had a wider range in the past.

December 1993Literature Cited

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BOBLOV, V. V.

1986. [Zoogeographical analysisof herpetofauna in Mongolia]. Symposium ofHerpetological Research. USSR Academy ofSciences, Moscow. Pp. 85-119.

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between males and females was the resultof sexual selection.

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H.

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