Asiatic Herpetological Research

2001

Vol.9, pp. 113-121

Lizards of the Northern Mongolian Deserts: Densities and Community

Structure

KONSTANTIN A. ROGOVIN

1,

DMITRY V. SEMENOV

1,

AND GEORGY I. SHENBROT 2'A.N.Severtzov Institute of Ecology and Evolution, Russian Academy of Science, Leninsky pr. 33 Moscow
1 17071,

Russia,~Ramon Science Center, Ben-Gurion University of the Negev, P.O. Box 194, Mizpe Ramon,
80600, Israel

Abstract.- Spatial organization and population densities of three-species lizard community was studied in theGobi Desert, Mongolia. To evaluate the effect of habitat variables on the distribution and abundance of eachspecies

we used the stepwise procedure of factor selection with ANOVA on each step. To describe thewe used stepwise discriminantdistribution of species' spatial niches in the space of environmental variables,

function analysis (DFA). The number of species in 1-ha grid areas varied fromto 4. Phrynocephalus versicolor
was the only species distributed over the 91% of grids occupied. There was a positive relationship between
distribution and local species abundance. A set of two to three habitat variables determined the abundance of eachspecies. The result of

DFA signify to the well pronounced segregation, but not even distribution of species spatial

niches in the space of resources.

Key words.- Lizards, Mongolia, community, ecology, density

91*

Fig

1.

Map

106°

101*

96*

of Mongolia and location of sites where data were collected.

111*

116'

Southern Gobi, 2-Western Gobi, 3-

Eastern Gobi, 4-Barun-Churay Basin.
at sites, a low level

Introduction
The reptile communities of Mongolian deserts

tats,

are

characterized by several specific features discussed

elsewhere (Ananjeva and Semenov, 1986; Borkin and

Semenov,

1984; Munkhbajar,

1976;

Semenov andBorkin, 1986; Semenov and Shenbrot, 1988). There
are few species in the fauna with low species richnessof species turnover between habi-

low abundance of most species and high domi-nance of only one species, Phrynocephalus versicolor.

A few common species have rather broad spatial
niches, diverse behavioral and physiological characteristics (diverse

range of thermobiological patterns,

wide active search for food items, etc.).

Asiatic Herpetological ResearchVol. 9, p. 114

The degree of interest in comparative studies ofstructureandfunctionofreptilecommunities

increased dramatically after seminal papers written by
E. Pianka (Pianka, 1973, 1975). Most of the ensuingstudies

were devoted to the species rich and diverse

communities of desert

2001with no specific features of vegetation.

transitiveThus, only the southern (extra-arid) and northern subzones are well pronounced.

The southern desert occurs mostly in the TransGobi and is characterized by a few very dry,Altai

south-

unproductive biotopes inhabited by five lizard species

western North America and southern Africa (Case,(two agamids, two gekkonids, and one Eremias species). Among this group only one species, Phryno-

1983; Fuentes,

lizards

in Australia,

1976; Henle,

1977; Millado et1989; Inger, Colwell,1975; Pianka,1986; Scheibe,al.,1987; Shenbrot et al., 1992; Simbotwe, 1984). At thesame time study of species poor communities in theCentral Asian desert can provide a significant information not only in comparison with other continents,
but also can help us to understand better which factors
rule in reality structure and dynamic of lizard communities of many species.

Up to now there were only two

examples of such studies made in China (Chang et al.,cephalus versicolor is common in
subzone (Semenov and Borkin, 1986).

The main objective of this paper was to study the
specific features of spatial organizationand popula-northern

The northern deserts, which extend to the southwest, west, and south-east ofMongolia and along
both slopes of the Mongolian Altai and Gobi-Altai
mountains, are characterized by pronounced microretypes are thealthough the main landscapesame as in the southern subzone. The

inrock and gravel valleys consists oflief and rich vegetation,

vegetation

1993;Luietal.,1992).

theperennial grass (Stipa), forbs, onions and succulents,
and a variety of annual plant species. Shrub vegetation
is often associated with foothills and sand dunes, or istion densities of three-species lizard community in the

spread along the dry river beds (Lavrenko, 1978). The

Gobi Desert, Mongolia. The study was conducted during a long-term research program on the biodiversity

difference in climate between western and easternof the Mongolian desert biota, and was sponsored bythePermanent Soviet-Mongolian Biological Expedi-

Mongolian desert is not pronounced (Murzaev, 1952); some differences exists inparts of the northern

the composition of the flora (Yunatov, 1950).

tion.

Lizard species

Material and Methods

There are four lizard species inhabiting northern

Mongolian desert

Mongolian deserts: Alsophylax pipiens, Phrynocephalus versicolor, Eremias przewalskii and E. multiocel-

A map (Fig.

1)

illustrates

thelocationof desert

regions of Mongolia. Three desert regions to the south

lata.

Among these, only the three last mentioned

species are abundant and relatively widespread.

of the Altai Mountains are partly separated from one
another by chains of low mountains and hills. These

Data collection

Gobi (South), Alashan Gobi
(East) and Sungarian Gobi or Barun-Churay Basin

We collected data during three field trips to the Mon-

three are Trans-Altai(West) (Yunatov, 1950). Besides these deserts there

golian northern deserts in June-August 1985, 1986
and 1988. Forty five 1-ha grids were established in theare desert areas between the Altai and Hangai moun-

Northern desert subzone (see map, Fig

tains, usually called

Western cold deserts, and somearid lands in the Great

Lakes and Ubsu-Nur Basins

Three arid subzones of the Mongolian desert are

Grids were—

sampled equally. Two factors determined the numberaridof grids at a desert region: diversity of habitat types

mm of rainfall per year), real desert (50—

and abundance of lizards. Each grid was divided into
25 smaller sample plots, 20 x 20 m. The centers of(Sokolovdesert (<50).

the clay basins and sand dunes. Each habitat type was(northwest).

defined1

distributed so that they covered the whole range of
habitats from the middle slopes of the mountains toandGunin,

100 mm per year) and steppe —1986):

extra

like desert (100—150

mm per year). However, the climatic border that
restricts the distributionof plants (Kazantseva, 1986;

Volkova et al., 1986) and animals (Podtyazhkin and
Orlov, 1986; Semenov and Borkin, 1986) exists only

sample plots were marked with 50 cm aluminum
stakes. Lizards were sampled by repeated, regularsearch of established grids during two to four consecutive days during periods of theirmaximal diurnal

encountered lizards were captured

between the southern part of Trans- Altai Gobi (< 50activity.

mm per year) and the northern waste belt of desert
precipitation (100-200

by hand. Most of the surveys accounting were conducted before the appearance of hatchlings. In the rest

mm per year). The narrow real desert subzone appearsof the cases hatchlings were not counted. Each cap-

lands withmore predictable

NearlyallAsiatic Herpetological Research

2001

Table 1. Density (no. ha</>)of lizard species.

s.

a

O
w
c

(A

"(5

TO

c
3
O

0)

'o
a>

N

Vol. 9, p. 115

2001

Asiatic Herpetological ResearchVol. 9, p. 116

cies but with the pronounced habitat preference. The

Results

second was E. multeocellata, and the third was E.

Densities anddistribution

through habitat

types
The results of lizard density and diversity estimations
on the 1-ha grids indicate the low local species diversity in

Mongolian deserts (Table

1).

The number of

species in our grid areas varied from zero to three.
Among 45 grids there were two grids with no lizards,

przewalskii. The last species had the most restricted
habitat preference, namely sandy-loess hills in saline

depressions with shrub vegetation of Nitraria sp.

The abundance of P. versicolor was positively correlated with its broad distribution. The density of thisspecies varied from1

to106 individuals per 1-ha

21 grids with only one species. 18 grids with two spe-

(Table 1). The second most abundant E. przewalskii(44 individuals per lha maximum), and the third wasand four grids with three species. Phrynoceph-

E. multiocelata (11 individuals per 1-ha maximum).

alus versicolor was the only species distributed over

All three species coexisted at rather high densities inmost number of grids (91% of grids occupied).
Eremias przewalskii was found on 33%, E. multioce-

Nitraria sp. shrubs. There were no pronounced nega-

4% of grids. Regarding

tive correlation between the densities of two dominant

cies

the

lata on 24% and A. pipiens on

distribution through the main habitat types (TableP.

1),versicolor was also the most widely distributed spe-saline depressions with sandy-loess hills covered withspecies on grids situated within this habitat typeversicolor-E. przewalskii: R"=0.04, n=20, ns).

Table 2. Designation and description for the 23 habitat variables included in the analysis.
Unit

Mnemonic

Variable

RCK

Content of rocks in the soil

GRW

Content of gravel in the soil

CLY

Content of clay in the soil

SCS

Sand cover area

%%
%%
%%
%%

SCH

Sand cover height

cm

WDS
WDD

Dry river bed area%%

Dry river bed depth

cm

NRB

Number of rodent burrows

no/sq.m.

FRB

Abundance of annual forbs

no/sq.m.

AGR

Abundance of annual grasses

no/sq.m.

ANN

Overall abundance of annual grasses and forbs

no/sq.m.

ALL

Perennial Allium covet

PGR

Perennial grass cover

MIC

Cover of microphyllous shrubs

HAL

Cover of halophytuos shrubssueCover of small succulent shrubs

HLX

Cover of Haloxylon

NIT

Cover of Nitraria

/o /o

SHC

Overall shrub cover

o/ o/
/o /o

SV1

Perennial plant crown volume at the level 0-25 cm

SV2

Perennial plant crown volume at the level 25-50 cm

SV3

Perennial plant crown volume at the level 0.5-1

SV4

Perennial plant crown volume at the level 1-2

%%
%%
%%
%%
%%
%%

m

m

%%
%%
0/ O/
/o /o

o/o.

Vo%

(P.

Vol. 9, p. 117

Asiatic Herpetological Research

2001

NIT

AGR
1

1

0.5

0.5
-0.5

-0.5

-1

-1

SHV2

SHV2
1

0.5

-0.5
-1

CLY
1

GRV

0.5

1

-0.5

0.5

-1

-0.5
Fig. 3. Habitat use profiles of-1

E przewalskii. For expla-

nation see Fig. 2.

1

Fig 2. Habitat use profiles of P. versicolor. For each

FRB

habitat variable rank, mean use by species is plotted
as a difference between capture frequency on plots ofthis rank and capture frequency of all plots. The size ofhistograms therefore denote the selectivity (positive or
negative) of habitat use. For mnemonics see Table 2

Spatial niches

There was a statistically significant influence of habitat

variables on the distribution and abundance of all

MIC

three studied lizard species (Table 3). For each ofthese species we extracted a set of two to three habitat
variables, determining 6.6-28.7% of observed variance in abundance. Densities of two species (P. versicolor,E. przewalskii)

were moderately affected byhabitat variables, whereas density of E. multiocellata

was weakly affected.
There was a statistically significant influence ofhabitat variables on the distribution and abundance ofFig. 4. Habitat use profiles ofE multiocellata. For

explanation see Fig.2.

three studied lizard species (Table 3). For each of

Patterns of habitat usage based on selected vari-these species we extracted a set of two to three habitat

all

vari-

able sets for each species are presented on Figures
2-4.
Phrynocephalus versicolor clearly avoided

ance in abundance. Densities of two species (P. versi-

microsites with low gravel content, very low and very

were moderately affected byhabitat variables, whereas density of E. multiocellata

high annual grass abundance, moderate and highand preshrub crown volume in the level 0.25-0.5

was weakly affected.

ferred microsites with moderate gravel content, mod-variables, determiningcolor,

E. przewalskii)

6.6-28.7% of observedmerate annual

grass abundance and very low shrubVol. 9, p. 118

Asiatic Herpetological Research

2001

Table 3. Summary of ANOVA analyses of influence of habitat variables on individual species' abundance. Values
are proportions of total dispersion determined by given variable. Total proportion of variance determinedby a setof variables may be greater than sum of influences of individual variables as a result ofSpeciesVarhigh-order interactions.

Asiatic Herpetological Research

2001

Vol. 9, p. 119

Fig. 5. Seventy-five percent confidence ellipses for the species observations on two discriminant axes (DF1andDF2). Em- Eremias multiocellata, Ep- Eremias przewalskii, Pv Phrynocephalus vesicolor.

fa

Q

DF1
dance with food abundance and a species-specific tacticof antipredator behavior.

It

preferred microsites

with moderate to high Nitraria cover and moderate tom(Fig. 3). In
high shrub crown volume at 0.25-0.5
summer E. przewalskii feed predominantly on berries

and young green twigs of Nitraria, and also find protection under the dense cover of thecrowns of low

spiny shrubs. In contrast to E. przewalskii, E. multiocellata avoided microsites with very high annual plant

abundance and shrub cover, but also preferred microsites

with moderate and rather high values of these

here means not the size of the species range area, but

number of sites where each species was found.
Phrynocephalus versicolor was encountered on 41

thegrid areas andhad the highest population density(mean: 37.3, median: 26 ind/ha, maximum: 106 ind/ha). With the edge-effect correction (Semenov,1991)

maximum density was 70.02 ind/ha (175.5 g/ha biomass). Eremias przewalskii was found on

1

5 grid areas

and was the second abundant species (mean: 9.5,
median: 4 ind/ha, maximum: 44 ind/ha). Eremias multiocellata

was found on

1

1

grid areasand its maxi-

variables (Fig. 4). This difference can be interpreted

mum density was 12 ind/ha (mean: 3.6, median: 2 ind/in accordance with thermobiological and size charac-ha). Alsophilax pipiens

teristics of two Eremias species. The larger species, E.

density 3 ind/ha.

przewalskii is not so quick as the smaller E. multiocellata. The first one digs well in soft soil, and ther-

moregulates climbing on or escaping under the shrub
periodically, being active throughout the day. Small E.
multiocellata that occupy habitats with relatively low

cover

(with

small

was met on two grids with

Brown (1995) explains this rather common relationship by the "Hutchinsonian niche
also for one species,

Brown

model"(see1984), suggesting that"the species that is slightly more tolerant of some abiotic conditions or biotic interaction or is slightly bet-

sparsely-distributed

ter able to use some resource should not only be able

shrubs) must cross open sites in search for food items

to occur in more places but also to attain higher abun-and escape predator's attacks by quickly rushing into

dance in some of those places." In the approach we

small shrubs or burrows. This species reduces heat byuse here niche breadth reflects microhabitat require-vegetation

escaping into burrow during the day-time.

Another result of the above comparison is the pos-ments of each species, namely the range of microconditions where each lizard species occurred. Diversity

itive relationship between distribution and local abun-of these microhabitats

dance among the species considered. Distribution

with the diversity of macrohabitats as well as with theisnot necessarily correlated

Vol. 9, p. 120

Asiatic Herpetological Research

Table 4. Summary of discriminant analysis of the habitats of lizard species. DF1

and DF2 are the first twocomponents (all are significant, P<0.001). Mnemonics
for habitat variables are from Table 1

.

2001

2001

Asiatic Herpetological Research

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