ISSN 20751117, Russian Journal of Biological Invasions, 2015, Vol. 6, No. 4, pp. 223–237. © Pleiades Publishing, Ltd., 2015.

Original Russian Text © S.N. Bazha, P.D. Gunin, E.V. Danzhalova, Yu.I. Drobyshev, T.I. Kazantseva, E. Ariunbold, D. Myagmarsuren, S. Khadbaatar, G. Tserenkhand, 2015,
published in Rossiiskii Zhurnal Biologicheskikh Invasii, 2015, No. 3, pp. 2–21.

Invasive Successions as the Indicator of Desertification

of Dry Steppe by Way of Example of Central Mongolia
S. N. Bazhaa, P. D. Gunina, E. V. Danzhalovaa, Yu. I. Drobysheva, T. I. Kazantsevab,
E. Ariunboldc, D. Myagmarsurend, S. Khadbaatare, and G. Tserenkhandc
aSevertsov
Institute of Ecology and Evolution, Russian Academy of Sciences, pr. Leninskii 33, Moscow, 119071 Russia
email: monexp@mail.ru
bKomarov Botanical Institute, Russian Academy of Sciences, ul. Professora Popova 2, St. Petersburg, 197376 Russia

email: bulgancum@gmail.com
cInstitute of Botany, Academy of Science of Mongolia, pr. Zhukova 77, Ulaanbaatar, 210351 Mongolia

email: gtseren@yahoo.com
d
Institute of Geoecology, Academy of Science of Mongolia, Ulaanbaatar, Mongolia
eMongolian State University of Education, Baga Toiruu14, Ulaanbaatar, 210648 Mongolia

email: hadbaatar@mail.ru
Received June 16, 2014

Abstract—The studies of the steppe ecosystems in Central Mongolia have showed that the simplification of
steppe communities has taken place over recent decades. This occurred by reduction of species diversity and
abundance of indigenous dominants—tussock grasses—as a result of sharp rise in pasture loads and a long
dry period. We have identified two types of introduction of invasive species from different landscapes:
(a) extra and intrazonal and (b) zonal. The first type of succession is characterized by focal distribution of
Ephedra sinica from ecosystems of low mountains to the surrounding mountain plains. The second type of
succession in dry steppes is associated with the expansion of Allium polyrrhizum, whose distribution has
largely a frontal character and is caused by weakening of the competitiveness of indigenous species of grass
communities because of their significant digression. A further factor in ensuring the conditions for invasion
is the aeolian alkalization of the upper horizons of zonal chestnut soils. The ecological and biological features
of these two species, widespread in the desertsteppe and desert landscapes and penetrating into the steppe
ecosystems, make it possible to speak about biological desertification. The wide area of Ephedra sinica and
Allium polyrrhizum indicates a progressive character of the studied types of succession, and as a result of this,
the borders of these areas have reached at present the southern periphery of the Baikal Lake basin. The paper
describes the processes which lead to the reduction of the fodder value of pastures and jeopardize the main
tenance of cattle breeding in Central Mongolia.

Keywords: succession, Ephedra sinica, Allium polyrrhizum, invasive species, dry steppes, pasture digression,
desertification, Central Mongolia
DOI: 10.1134/S2075111715040025

INTRODUCTION differences in the mechanical composition of the soil

According to E.M. Lavrenko (Lavrenko et al.,
1991), steppes as a type of vegetation are formed by
herbaceous communities of the northern temperate
zone with the dominance of tussock species, mainly
large and small tussock grasses from the genera Stipa,
Festuca, Agropyron, Koeleria, Cleistogenes, and Helito
trichon, and less commonly of sedges (Carex) and
onions (Allium) which form the grass stand and the
maximum of phytomass. Forbs are widely represented
in steppes; their species composition and production
decreases from the north to the south. In addition,
welldefined sinusia of subshrubs (Artemisia) and
shrubs (Spiraea, Caragana, etc.) in steppes are due to
forming rocks.
Dry bunchgrass steppes on chestnut soils are the
most widespread subzonal type of steppes in Mongo
lia. They form a wide belt on plains in its central and
eastern parts, occupying almost 35% of the steppe area
or 15% of the area of the country (Ecosystems of Mon
golia, 2005). Such species as Stipa krylovii, Cleistogenes
squarrosa, Agropyron cristatum, and Koeleria cristata
and less commonly Poa botryoides and Stipa grandis
prevail among tussock grasses (Sukhie stepi…, 1984;
Lavrenko et al., 1991). Dry steppes are characterized
by a limited share of onions and forbs (Allium anisopo
dium, A. bidentatum, A. tenuissium, Potentilla acaulis,
Sibbaldianthe adpressa). Xerophytic shrubs Caragana

223
224 BAZHA et al.
(C. microphylla, C. stenophylla, and C. pygmaea) often
play an edificatory role in dry steppes. Their distribu
tion is associated with a high content of pebble in the
soil substrate or sandification of the surface soil hori
zons (Yunatov, 1950). In addition, dry steppes are
characterized by admixture of subshrubs (Artemisia
frigida, A. adamsii, and Kochia prostrata). The pres
ence of summer–autumn annual plants of the genera
Chenopodium, Artemisia, Dontostemon, and Chamaer
hodos, which grow heavily in wet years, is typical of dry
steppes (Sukhie stepi…, 1984).
Signs of pasture digression were recorded as early as
the middle of the past century. On the basis of the
results of the studies conducted in the 1950s–1960s,
Yunatov (1950), Miroshnichenko (1964, 2004), and
Chognii (1988) reported subshrub Artemisia frigida as
a species which grows actively in pastures and is resis
tant to cattle grazing. The authors also noted that such
species as Leymus chinensis, Carex duriuscula, Ther
mopsis lanceolata, Artemisia changaica, Potentilla acau
lis, and Schizonepeta multifida positively responded to
grazing. Davazhamts (1954), who conducted studies
in steppes in the Uburkhangai aimag, indicated an
increasing role of Carex duriuscula and Artemisia
frigida, as well as Kochia prostrata and Allium anispo
dium, under pasture intensification. The studies which
were conducted at the BayanUndzhul somon station
in the 1970s also demonstrated that Caragana–true
grasses–Artemisia frigida pastures are a variant in a
digressive series of Caragana–true grasses pastures
which are subject to extensive grazing. A distinct fea
ture of a digressive variant of forb–true grasses pas
tures is the abundance of Carex duriuscula and Cleisto
genes squarrosa, which makes it possible to conclude
the formation of Cleistogenes–sedge–Artemisia frigida
communities in pastures subjected to extensive grazing
(Sukhiye stepi…, 1984).
Over the past 20 years, increasing pasture load
caused by the transition of Mongolia to a market
economy and increase in the number of cattlebreed
ing farms and a two to threefold increase in livestock
(Gunin et al., 2009; Ariunbold, 2014) have intensified
digression of vegetation communities and degradation
of the ecosystem. Thus, studies which were conducted
along the TransMongolian railway showed that areas
with strongly and very strongly disturbed vegetation
cover began to prevail among vegetation communities
of steppes (Miklyaeva et al., 2004; Miklyaeva and
Fakkhire, 2004). When comparing the areas which
have been more than 50 years in the trackside zone of
the railway with their landscapeecological analogs
under grazing, a positive response of shrubs of the
genus Caragana and subshrubs of the genus Artemisia
(A. adamsii, A. frigida) to pasturing was found; among
other species, rhizomatous plants Carex duriuscula,
Leymus chinensis, and Potentilla acaulis play an active
role in pastures.
Over the past decade (2004–2014), the transfor
mation of the vegetation cover in ecosystems of dry
RUSSIAN JOURNAL OF BIOLOGICAL INVASIONS
steppes occurred because of the continuous increase in
the anthropogenic load and repeated droughts. We
assume that the transformation is such state of the veg
etation community when the role of native edificators
becomes insignificant or they completely disappear
from the phytocenosis (Bazha et al., 2008, 2012). An
important feature of this process in the subzone of dry
steppes is the invasion into vegetation communities
and expansion of habitats of such desertsteppe spe
cies as Peganum nigellastrum, Stipa inebrians, Ephedra
sinica, and Allium polyrrhizum (Gunin et al., 2013).
According to the classification of invasive plants by the
degree of their aggressiveness and features of distribu
tion, the species under consideration can be consid
ered as transformers (Richardson et al., 2000), which
are characterized by an active penetration into natural
and seminatural communities leading to changes in
the ecosystems and disturbance of existing phyto
cenotic relations. These species begin to play the role
of edificators and dominants, forming overgrowths
and (or) preventing restoration of native flora (Akatov
and Akatova, 2010; Notov et al., 2011). The studies of
communities in dry steppes of Central Mongolia have
demonstrated that at present the communities with
the dominance of Ephedra sinica and Allium polyrrhi
zum have been formed in vast areas, which necessitates
more detailed consideration of ecological and biolog
ical features of these two species and the causes of their
spreading.
MATERIALS AND METHODS
The state of dry steppes was studied in the southern
part of the Central aimag and the northern part of the
Middle Gobi aimag. The relief in the surveyed territo
ries is represented by gently undulating and lowhill
plains and lowmountain massifs. The climate is
extremely continental with severe winters (the average
temperature in January is –22°C), warm summers
(the average temperature in July is +20°C), and low
amount of precipitation (165–280 mm yearly), most
of which falls in the second half of July and August,
and frequently repeated droughts observed for 2–
3 years in 10 (Beresneva, 1984). According to the
botanical and geographical zoning proposed by
Lavrenenko (1970), the steppes under study belong to
the Middle Khalkha subprovince of the Mongolian
province in Eurasian steppes. In the latest zoning pro
posed by Kamelin (2010), the surveyed region belongs
to the Khalkha okrug of the DarigangEastern Mon
golia transition territory.
Field studies were conducted during the height of
herbage (midJuly–midAugust) from 2008 to 2013.
In each plot, detailed geobotanical descriptions of the
communities in the area of 100 m2 were made. Phyto
cenotic parameters (the number, projective cover, and
aboveground phytomass of herbaceous plants) were
calculated in plots of 1 m2 in three to five replicates.
Biomorphometric parameters of invasive species were
Vol. 6 No. 4 2015