Vegetation of the woodland-steppe transition at the
southeastern edge of the Inner Mongolia Plateau
Hongyan Liu1, 2, *, Haiting Cui1, Richard Pott2, Martin Speier2
1
Department of Geography, Peking University, Beijing, 100871, P. R. China
Institut für Geobotanik der Universität Hannover, Nienburger Strasse 17, D-30167, Hannover,
2
Germany
*Tel. +86 10 62751181; Fax +86 10 62751187; Email lhy@urban.pku.edu.cn
Published on Journal of Vegetation Sciences, 2000, 11(4): 525~532
Abstract:
The vegetation of the woodland-steppe transition in southeastern Inner Mongolia, where the East
Asian monsoon climate reaches its northwestern edge, is described and analysed in this paper. The
communities are classified in a phytocoenological way. 12 major types of woodland, shrubland,
meadow, fen, open woodland and steppe are differentiated and described according to 133 relevés.
DCA results show that precipitation plays a crucial role in the distribution of grassland communities
while woodland and shrubland communities are controlled by both warmth and humidity conditions.
Four vegetation zones can be distinguished. From woodland zone to woodland-grassland zone, the
temperature decreases and the precipitation increases with the rising of the altitude, which leads to the
conditions suitable for the meadow and fen communities. Transiting to the woodland-steppe zone, the
temperature increases while the precipitation decreases with the gradual lowering of altitude, steppe
communities form a matrix and woodlands have patchy distribution. From woodland-steppe zone to
steppe zone, the precipitation rather than temperature decreases obviously, as a result the woodland
communities disappear gradually. In a local scale, geomorphologic conditions determine the
vegetation pattern of the study area. The existence of the sandy land permits different woodland types
occur together in the woodland-steppe transition. The distribution of woodland and steppe
communities in a discrete site is depended on slope conditions. In addition, Human disturbance has
also left overprints upon the composition of plant communities.
1
Keywords: Inner Mongolia China, DCA, phytocoenological approach, vegetation gradient,
woodland-steppe
Nomenclature: Anon. (1989a, 1992, 1994).
Introduction
Under the influence of the East Asian monsoon climate, the precipitation in the eastern part of
China decreases from SE to NW with the weakening of the summer monsoon (SE monsoon).
Consequently, the vegetation shifts from woodland to steppe near the northwestern edge of the
summer monsoon (Wu 1980; Hou 1988; Lavrenko et al. 1993). Walter (1977) called this type of
transition as a "zonoecotone between deciduous broad-leaved forest and steppe". As "ecotone" and
"ecocline" are two confused concepts in ecology (Jeník 1992), we prefer "transitional zone" in this
paper.
Monsoon climate is usually regarded as fluctuating dramatically near its northwestern edge. The
corresponding woodland-steppe transitional zone was inferred to be sensitive to climatic change and
thus could be used as a monitor of regional environmental change (Wang et al. 1991; Liu et al. 1992).
How does climate affect vegetation pattern in this area is a key to reconstruct the vegetation
development under the dynamics of the monsoon climate. Unfortunately, the exact vegetation gradient
and the vegetation-climate relationship in the woodland-steppe transitional zone in China are poorly
understood.
The woodland-steppe transitional zone in southeastern Inner Mongolia is selected as the study
area of this paper. The former research was focused on macro-scale vegetation survey (e.g. Anon.
1985, 1988, 1989b, 1996) according to a few dominant species. In this paper, Braun-Blanquet method,
numerical method (DCA) and landscape transects are combined to describe the vegetation patterns
and to reveal their relations to the environmental conditions in the study area.
2
Study area and method
Geomorphologic features
The geographical coordinates of the study area are 42º-43.75ºN, 115.75º-117.75ºE (Fig. 1). Its
main part is located in the southeastern edge of the Inner Mongolian Plateau with an elevation of
1100-1400m. It displays a great variation in its geomorphologic features. The Xilinguole Lava
Platform is situated in the north. The Otindag (Hunshandak) sandy land, which consists of many E-W
oriented dune belts, dominates the western part of the study area. It extends eastward as far as the
valley of the Xilamulun River. In the east of the Otindag sandy land, the Mandaxile hills covered with
sands stretch in an N-S orientation. The Greater Hinggan Mountains to the northeast and the Jibei
Mountains to the south and southeast border the Inner Mongolian Plateau in the study area.
Climate
The contour map of the average temperature in January, the average temperature in July, the
annual average temperature and the annual precipitation are shown in Figure 2. The annual
precipitation has a sharp tendency from SE to NW. It decreases from about 450 mm in the SE to about
320 mm in the NW. The temperature parameters are higher in the valleys and lower on the high peaks
of the Jibei Mountains, but there is no obvious gradient inside the plateau.
Soils
The soil types of the study area are diversified owing to the climatic gradient and the
geomorphologic pattern. From the Jibei Mountains in the SE to the Xilinguole Lava Platform in the
NW, the soil types change from brown soil, grey forest soil, light chernozem to chestnut soil. Sandy
soils are prevalent on the Otindag sandy land. Other azonal soil types, for example subalpine meadow
soil, appear locally (Anon. 1988, 1989b).
3
Study method
71 sites were selected for the field survey. The typical sites of woodlands, open woodlands,
shrublands, steppes, grasslands and fens have been examined by relevé method comprehensively
described by Mueller-Dombois and Ellenberg (1974). 148 plots have been made in total. The
Braun-Blanquet method (Braun-Blanquet, 1964) is used for the classification of communities in this
paper. The constancy and coverage degrees are described following Dierschke (1994).
The software DECORANA (Hill, 1979) is used to execute DCA. DCA for relevés according to
their species composition is used to show the community-climate relationship while a converse one is
used to show the relationship between species distribution and climate conditions (Gauch 1982, Zhang
1994). Ecological groups of species are divided according to the AX1 and AX2 values of each species
on the DCA surface by using the axes classification suggested by Zhang (1994). Contour map of
species distribution is obtained by using Kriging interpolation (Matheron, 1963) of the percentages of
ecological groups in each sample site.
Kira's Warmth Index (see Xu, 1985) and the Humidity Index suggested by Li (1983), which are
demonstrated to be useful to describe the vegetation-climate relationship in East Asia, are also
calculated in this paper. The calculation of these two indexes are described as follows:
WI=(ti-5)
HI= (pi/2-ti)
Where ti is the average monthly temperature when it >5 C, and pi is the monthly precipitation
when the average monthly temperature>5 C.
Result
Vegetation classification
The differential species of main community types are given in table 1, including 133 plots.
Several types with less than 5 plots, such as Larix principis-ruprechtii-woodland, Betula
fruticosa-scrub, Stipa krylovii-steppe, Filifolium sibiricum-steppe and Thymus serphyllum-steppe, are
4
omitted. For details of each community type see Liu (1998).
1. Quercus mongolica-woodland
The Quercus mongolica-woodland is a zonal community type in the northern part of the
temperate forest zone in China. It mainly occurs on rocky shady slopes with brown soils of Jibei
Mountains. On shady slopes covered with sands near Haoluku, it is occasionally distributed. The
elevation of its distribution is usually below 1400 m in the study area. Caused by human disturbances,
the woodland patches are relatively small with a more or less open canopy.
2. Betula platyphylla-woodland
The
Betula platyphylla-woodland occurs at an elevation from 1100 m to 1750 m. It is
usually distributed on shady slopes of sand-covered hills. The top 10-20 cm of its soil mainly consists
of sands. Two subunits can be distinguished in regard to the species composition. One of them is
limited to an elevation of over 1450 m, containing a group of differential species such as Valeriana
officinalis, Polemonium caeruleum, Maianthemum bifolium and Equisetum sylvaticum, which have a
preference for a cool and humid habitat. This subunit can be called ‘Valeriana officinalis-Betula
platyphylla-woodland’. The other is confined to an altitude of less than 1450 m. It contains the
differential species Ostryopsis davidiana, Bupleurum chinense, Galium verum, Viola variegata,
Potentilla tanacetifolia and Rubia cordifolia and can be called ‘Ostryopsis davidiana-Betula
platyphylla woodland’.
3. Betula dahurica-woodland
This type characterizes those regions in the Jibei Mountains with an elevation of 1200-1600 m.
The differential understory species include Prunus padus, Galium aparine var. tenerum, Adenophora
divaricata, Adenophora boreale. Betula dahurica-woodland usually occurs from the upper slope to the
ridge, while Quercus mongolica-woodland commonly occurs on the foot slope (Wu 1980). The
distribution of these two community types in the study area is approximately coincident with the
above description, except that the elevation range for the distribution of Betula dahurica-woodland is
sometimes higher than that of Quercus mongolica-woodland.
4. Populus davidiana-woodland
5
In the study area, Populus davidiana-woodland mainly occurs on the shady slopes of the
sand-covered hills in the Inner Mongolia Plateau with an elevation of less than 1450 m. The Populus
davidiana-woodland
occupies
similar
habitat
with
the
Ostryopsis
davidiana-Betula
platyphylla-woodland. Betula platyphylla and Quercus mongolica are commonly present in the upper
part of the tree layer.
Betula platyphylla-woodland and Populus davidiana-woodland (or Populus tremula-woodland)
are generally regarded as one type with the very similar ecological requirements (Hilbig & Knapp
1983; Hilbig 1995). In this study area, the Populus davidiana-woodland occurs below 1400 m and has
a similar habitat with the Ostryopsis davidiana-Betula platyphylla-woodland. When it exceeds 1400 m,
no Populus davidiana-woodland occurs.
5. Picea meyeri-woodland
Picea meyeri reaches the northern edge of its areal in the southeastern edge of the Inner
Mongolia Plateau (Anon. 1989). Only a few patches of Picea meyeri-woodland can be found in some
dunes in the study area. Betula platyphylla grows in the tree layer, as well as Artemisia frigida in the
herb layer. Both species have a high constancy, which leads to the fact that the Picea meyeri-woodland
is not typical in the study area in comparison with its distribution center. Picea meyeri has a
preference for cool and humid climate. Although the climate is dry in the study area, the good water
supply of the sandy land is probably favorable for its growth.
6. Pinus tabulaeformis-woodland
Pinus tabulaeformis grows well in the Jibei Mountains. But in the Inner Mongolia Plateau, the
occurrence of Pinus tabulaeformis is confined to several dunes. The largest patch occupies an area of
a few hundred square meters, the smallest one is built by only a few individuals.
Some researchers have made reference to the fact that 44ºN characterizes the northern border
line of Pinus tabulaeformis woodland in Inner Mongolia (Anon, 1989). Near to this border, the
regeneration of Pinus tabulaeformis is bad, for example in Narsu (43.09ºN, 116.60ºE) and
Ganqinarsi (43.24ºN, 116.70ºE), no seedlings of Pinus tabulaeformis can be found.
7. Ostryopsis davidiana-shrubland
6
This type is a prevalent shrubland type in the study area. It lacks differential species of itself. Two
subunits can be divided, one has the differential species Spiraea aquilegifolia, Veronica incana,
Bupleurum chinense and Leontopodium longifolium while the other one lacks this group of species.
8. Polygonum viviparum-meadow
This community type occurs in mountains with an elevation of over 1450 m. It forms a
finger-staggered
pattern
with
woodlands,
especially
with
Valeriana
officinalis-Betula
platyphylla-woodland. The differential species of this type is Polygonum viviparum.
9. Ranunculus japonica-fen
The Ranunculus japonica-fen usually occurs in the Jibei Mountains with an elevation of over
1400 m. Some Salix shrubs present occasionally. The differential species of this type is Ranunculus
japonicus.
10. Stipa baicalensis-steppe
This type belongs to the meadow-steppe (Zhu 1993). Similar to the Betula platyphylla-woodland,
it can also be divided into two subunits. One is confined to an elevation of more than 1400 m and
contains the two differential species Sanguisorba officinalis and Artemisia laciniata, which prefer a
humid habitat. The other subunit is limited to an elevation of less than 1400 m and does not contain
the above mentioned species.
11. Leymus chinensis-steppe
The Leymus chinensis-steppe occurs vastly in the study area. Artemisia frigida, Cleistogenes
squarrosa, Carex korshinskii and Artemisia capillaris have high coverage and constancy under
different degrees of human disturbance. The dominant species are different in relation to the degree of
human disturbance. Two subunits can be classified. One is differentiated by Carex korshinskii and can
be named as Carex korshinskii-Leymus chinensis-steppe. It is distributed mainly at the foot of the
slopes in the southeastern part of the study region and forms a mosaic together with the Stipa
baicalensis-steppe. The other subunit is differentiated by Carex pediformis and is mostly distributed in
the northwestern part of the study area.
12. Ulmus pumila-open woodland
7
Ulmus pumila-open woodlands usually grow at the foot of the dunes within the Otindag sandy
land. Only one species, Ulmus pumila, appears in the tree layer. The coverage of the tree layer is
usually less than 30%. Spiraea aquilegifolia has a relatively high constancy and coverage in the shrub
layer. Hilbig (1995) called it in Mongolia as Spiraeo aquilegifoliae-Ulmetum pumilae (elm bush
forest). But in the study area, Spiraea aquilegifolia can not be regarded as a differential species
because of its higher constancy in other types. The steppe elements, for example Artemisia frigida,
Cleistogenes squarrosa, Carex korshinskii and Potentilla accaulis, have high degrees of constancy in
the herb layer. In contrast, elements of summer-green broadleaved forest are relatively short.
Therefore this community type has a character closer to steppe than to woodland.
Vegetation-climate relationship
For woodland, open woodland and shrubland communities, the DCA result is expressed in Figure
3. AX1 represents the warmth gradient and a better warmth condition is expressed by a higher AX1
value. The warmth condition of woodland and shrubland communities can be summarized as follow:
Ulmus pumila-open woodland>Pinus tabulaeformis-woodland and Picea meyeri-woodland>Populus
davidiana-woodland and Quercus mongolica-woodland>Betula platyphylla-woodland and Betula
duhurica-woodland. AX2 represents a humidity gradient. The humidity condition of woodland and
shrubland communities can be summarized as follow: Pinus tabulaeformis-woodland>Betula
platyphylla-woodland and Betula dahurica-woodland>Populus davidiana-woodland and Quercus
mongolica-woodland>Picea meyeri-woodland>Ulmus pumila-open woodland.
In general, this result coincides with the community-climate relationship in North China (cf.
Anon. 1985, 1989, 1996, Zhou 1997). But the DCA result for Picea meyeri is abnormal. Picea meyeri
has a preference for cool and humid climate and Picea meyeri-woodland has the similar warmth and
humidity requirement with Betula platyphylla-woodland in North China. In the study area, Picea
meyeri distributes on some dunes as extra-zonal community, which depends on the good water supply
of the sandy land although precipitation is relative short.
The DCA result of steppe and grassland communities is expressed in Figure 4. AX1 represents a
8
humidity gradient. Stipa bacalensis-steppe and Filifolium sibiricum-steppe have higher humidity
requirement than other steppe communities, which meets the normal situation in North and Northeast
China (Danert et al. 1961; Anon. 1985, 1996; Zhu 1993). AX2 represents a warmth gradient. No
obvious difference is found between various community types, which shows that temperature is not a
decisive factor for the distribution of steppe and grassland communities in the study area.
The vegetation gradient from woodland to steppe
Four zones can be distinguished according to the vegetation landscape. They are woodland zone,
woodland-grassland zone, woodland-steppe zone and steppe zone (Fig. 1). The landscapes of these
zones are distinctly differentiated and this division has been demonstrated by the remotely sensed
images (Guo et al., 1999).
Figure 5 shows the Warmth Index, Humidity Index, altitude and vegetation along a transect from
SE to NW in the study area. From woodland zone to woodland-steppe zone, the temperature decreases
and the precipitation increases with the rising of the altitude, which leads to the conditions suitable for
the grassland and fen communities, the vegetation changes from woodland to a combination of
woodland and grassland. Transiting to the woodland-steppe zone, the temperature increases while the
precipitation decreases with the gradual lowering of altitude, steppe communities form a matrix. From
woodland-steppe zone to steppe zone, the precipitation rather than warmth condition decreases
obviously, as a result the woodland communities disappear gradually.
The above result shows that vegetation has an obvious gradient from SE to NW in the study area,
which coincides with the precipitation gradient. Species distribution also reflects this gradient.
Ecological groups of species are distinguished according to the DCA values of AX1 and AX2 that
reflect their requirement of habitat conditions. The DCA result for species shows that AX1 represents
the warmth conditions while AX2 represents the humidity conditions. Mean value of all the species in
AX1 axis (AX1) and that of AX2 axis (AX2) are calculated respectively. Four groups are then divided
and compared with species distribution in northern China.
Group I (AX1> AX1, AX2>AX2): This group of species requires a warm and humid climate and
9
is distributed mostly in the deciduous broad-leaved woodlands in northern China (Anon. 1985, 1996;
Zhou 1997)
Group II (AX1>AX1, AX2<AX2): This group of species needs a cold and dry climate and is
distributed mostly in the steppe communities.
Group III (AX1<AX1, AX2>AX2): This group of species needs a cold and humid climate.
Group IV (AX1<AX1, AX2<AX2): This group of species needs a warm and dry climate.
Figure 6 shows the contour map of the above four groups obtained by Kriging interpolation.
Group II decreases markedly from NW to SE while Group I has a converse tendency, which is
corresponding to the precipitation gradient. Figure 6e is the contour map of the ratio between group II
and group I. A marked decrease from NW to SE can be found, which shows that woodland species
decreases and steppe species increases gradually when climate becomes drier. This gradient of species
distribution is the basis for the above vegetation gradient.
Discussion
Climate and vegetation gradient
Walter (1977, 1990) has discussed the characteristics of the woodland-steppe transitional zone in
detail. Unlike Savanna, the zonoecotone between temperate deciduous woodland and steppe is a
landscape mosaic of woodlands and steppes in a large scale. With the drying of the climate, the
woodlands change from matrix to smaller and smaller patches, and finally disappear. The insufficient
water supply lead to the disappearance of woodland. A similar conclusion can be drawn in the study
area, precipitation rather than temperature plays a crucial role in the vegetation gradient from
woodland zone to steppe zone. The disappearance of woodland in the steppe zone is due to the low
precipitation.
The climate variance also leads to community differentiation. For example, The Leymus
chinensis-steppe can be distinguished into two subunits. One occurs in the woodland-grassland zone
and woodland-steppe zone, and is differentiated by Carex korshinskii. The other one occurs in the
steppe zone and is differentiated by Carex pediformis.
10
Geomorphologic conditions and vegetation patterns
Climate variances caused by geomorphologic factors also play an important role in the vegetation
gradient of the study area. The occurrence of the woodland-grassland zone in the study area is due to a
cold and humid climate caused by a marked rising of the altitude. The differentiation of the
woodland-grassland zone and the woodland-steppe zone leads to the differentiation of plant
communities. For example, two subunits of the Stipa baicalensis-steppe occur in the
woodland-grassland zone and the woodland-steppe zone respectively. The distribution of two subunits
of Betula platyphylla-woodland, Valeriana officinalis-Betula platyphylla-woodland and Ostryopsis
davidiana-Betula platyphylla-woodland, is similar. The species distributions of group III and group IV
have also relations with geomorphic conditions. High values of Group III can be found in the high
mountains and the Xilinguole Lava Platform while that of group IV can be found in the Otindag sandy
land.
Lavrenko et al. (1993) has referred to the diversity of plant communities owing to the monsoon
climate in North China. In our study area, the high diversity is not caused directly by the monsoon
climate. The monsoon climate has led to higher diversity of plant communities in the temperate
woodland region and the geomorphic variance in the woodland-steppe transitional zone allows various
woodland community types to join together and form a complicated landscape mosaic with steppe
communities. For example, the good water supply of sandy land in the study area allows Picea
meyeri-woodland and Pinus tabulaeformis-woodland to distribute. The occurrence of Ulmus
pumila-open woodland is also limited to sandy land.
Walter (1974) has concluded that soil condition is the key factor to the distribution of woodland
and steppe in a discrete site of the woodland-steppe transitional zone in flat areas. He suggested that
woodlands occur at well-drained coarse soil while steppes occur at bad-drained fine soils. In our study
area, the distribution of woodland and steppe is determined by gradient and exposure of slopes in a
discrete site. Woodlands occur usually at shady slopes over 20 degrees while steppes usually occur at
sunny slopes and shady slopes less than 20 degrees. We guess that water supply and light condition
11
rather than soil texture are critical factors for the distribution of woodland in the study area.
Effects of Human disturbances on the vegetation
Although the study area has only a history of human cultivation for about one century, the rapid
growth of human population has lead to inappropriate land use and left overprints upon the
composition of plant communities. For example, Ostryopsis davidiana-shrubland is a degradation of
the broadleaved woodlands in the study area and thus has no differential species. The Valeriana
officinalis-Betula
platyphylla-woodland
is
a
degradation
phase
of
the
Larix
principis-ruprechtii-woodland, its differential species Valeriana officinalis, Polemonium caeruleum,
Maianthemum bifolium and Equisetum sylvaticum, are also commonly distributed under the Larix
principis-ruprechtii-woodland (Anon. 1985). Steppe communities are usually dominated by such
species as Artemisia frigida, Potentilla acaulis, Cleistogenes squarrosa under overgrazing.
The vegetation coverage has also decreased under strong human disturbances. As a result, sandy
land activated as well as expanded its area, especially at the eastern boundary of the Otindag sandy
land. For the prediction of future vegetation development in the study area, factors of both climate
changes and human disturbances must be considered.
Acknowledgments
This paper is part of the results of the research project ‘ Shifting of landscape borders
in the semi-humid to semi-arid transitional zone in North China’ which is financially
supported by the National Natural Science Foundation of China (NSFC No. 49571069). It
is a part of a bilateral scientific work about the reconstruction of vegetation in
mountainous and alpine area in the Eurasia continent supported by NSFC and the German
National Science Foundation (DFG). We thank Huang Yongmei, Cao Yanli, Liu Liping, Li
Dihua, Feng Xiahong and Wang Yulin for taking part in the field survey. Guo Qinghua
and Zhang Zepu have helped us with data management. We express our thanks for their
help.
12
References
Anon. 1985. Vegetation of Inner Mongolia, Science Press, Beijing. (in Chinese)
Anon. 1988. Serial maps of resources of Inner Mongolia, Science Press, Beijing. (in Chinese).
Anon. 1989a. Flora of Hebei (Vol. I, Vol. II). Hebei Science and Technology Press, Shijiazhuang. (in
Chinese)
Anon. 1989b. Forest of Inner Mongolia. China Forestry Publishing House, Beijing. (in Chinese)
Anon. 1992. Flora of Hebei (Vol. III). Hebei Science and Technology Press, Shijiazhuang. (in
Chinese)
Anon. 1994. Flora of Inner Mongolia (Vol. II-V). Inner Mongolia Remin Press, Huhhot. (in Chinese)
Anon. 1996. Vegetation of Hebei Province. Science Press, Beijing. (in Chinese)
Braun-Blanquet, J. 1964. Pflanzensoziologie. 3. Aufl. Springer Verlag, Wien, New York.
Denart, S., Geier, S. & Hanelt, P. 1961. Vegetationskundliche Studien in Nordostchina (Mandschurei)
und der Inneren Mongolei. Feddes Repertorium, 139: 5-144.
Dierschke, H. 1994. Pflanzensoziologie. Verlag Eugen Ulmer, Stuttgart.
Gauch, H. G. 1982. Multivariates analysis in community ecology. Cambridge University Press,
London, New York.
Guo, Q., Yu, H., Zhang, Z. & Cao, Y. 1999. Study of the woodland-steppe transitional zone in North
China with remotely sensed data. Journal of Peking University (Natural Science), 35: 550-556.
Hilbig, W. & Knapp, H.D. 1983. Vegetationsmosaik und Florenelemente an der Wald-Steppe-Grenze
im Chentej-Gebirge (Mongolei). Flora, 174:1-89.
Hilbig, W. 1995. The vegetation of Mongolia. SPB Academic Publishing, Armsterdam.
Hill, M. O. 1979. DECORANA——A Fortran program for Detrended Correspondence Analysis.
Cornell University, Ithaca, New York.
Hou, H.-Y. 1988. Physical geography of China: Phytogeography (II). Science Press, Beijing. (in
Chinese)
Jeník, J. 1992. Ecotone and ecocline: Two questionable concepts in ecology, Ekologia (CSFR), 11:
243-250.
13
Lavrenko, E. M. & Karamysheva, Z.V. 1993. Steppes of the former Soviet Union and Mongolia. In:
Coupland R. T. (ed.) Natural Grassland－Eastern Hemisphere and Résumé, pp 3-59. Elsevier,
Amsterdam.
Li, W. 1983. A quantitative analysis of the relationship between vegetation distribution and
warmth-humidity factors in Hengduan Mountains, Yunnan Province, China, In: Anon (ed.)
Proceedings of an integrated survey in Hengduan Mountains, pp 185-204. Yunnan Renmin Press,
Kunmin. (in Chinese)
Liu, H. 1998. Past and present woodland-steppe ecotone in the southeastern edge of the Inner
Mongolia Plateau. Unpublished Dissertation of the University of Hannover.
Liu, Q. & Li, H. 1992. Holocene environmental change of the agriculture-pasture transitional zone in
northern China (Daihai-Huangqihai region), In Zhou T. and Zhang L. (eds.) Holocene
environmental change and its prediction of the agriculture-pasture transitional zone in northern
China, pp.16-50. Geological Publishing House, Beijing.
Matheron, G. 1963. Principles of geostatistics. Economic geography, 58: 1246-1266.
Mueller-Dombois, D. & Ellenberg, H. 1974. Aims and methods of vegetation ecology. John Wiley &
Sons, New York.
Walter, H. 1974. Die Vegetation Osteuropas, Nord- und Zentralasiens. Gustav Fischer Verlag,
Stuttgart.
Walter, H. 1977. Vegetation of the earth and ecological systems of the geobiosphere. 2nd ed. Springer
Verlag, New York
Walter, H. 1990. Vegetation und Klimazonen. 6. Aufl. Verlag Eugen Ulmer, Stuttgart.
Wang, S. & Feng, M. 1991. Environmental change and its relation with the weakening of the summer
monsoon in Daihai, Inner Mongolia. Science in China (Ser. B), 21: 759-768.
Wu, C. Y. (ed.). 1980. Vegetation of China. Science Press, Beijing. (in Chinese)
Xu, W. 1985. Kira's Warmth Index and its application in the vegetation of China. Chinese Journal of
Ecology, .5: 53-59. (in Chinese with English abstract)
Zhou, Y. 1997. Geography of the vegetation in Northeast China. Science Press, Beijing. (in Chinese)
14
Zhang, J. 1994. The classification method based on DCA axes and its applications. Chinese Journal of
Ecology, 13: 73-75. (in Chinese with English abstract)
Zhu T.-C. 1993. Grassland of China. In: Coupland R.T. (ed.) Natural Grassland－Eastern Hemisphere
and Résumé, pp. 61-82. Elsevier, Amsterdam.
15
Table 1. Synoptic table of major communities in the study area. The first number in a cell
indicates the constancy while the second one indicates the mean coverage of each species
in the plots of the correspondent community type.
Woodland and shrubland communities:
1. Quercus mongolica-woodland; 2: Betula dahurica-woodland;
3: Betula platyphylla-woodland;
4: Populus davidiana-woodland;
5: Pinus tabulaeformis-woodland; 6: Picea meyeri-woodland;
7: Ostryopsis davidiana-shrubland
8-9: Grassland and fen community
8: Polygonum viviparum-meadow; 9 Ranunculus japonicus-fen
10-12: Steppe and open woodland community
10: Stipa baicalensis-steppe;
11: Leymus chinensis-steppe;
12: Ulmus pumila-open woodland
Community type
1
2
3
4
5
6
7
Number of Relevés
5
5
17 11 5
8
6
Average Coverage of Tree Layer (%) 50 55 55 55 40 60 0
Average Coverage of Shrub Layer (%) 30 20 20 25 20 5
55
Average coverage of Herb Layer (%)
35 30 45 30 25 15 25
Carex lanceolata
V,2 V,1 V,1 V,1 IV,1 III,1 IV,1
Vicia unijuga
IV,+ V,+ V,+ V,+ II,+ IV,1 I,+
Deyeuxia arundinacea
III,+ II,+ I,+ III,+ III,+ III,+ I,+
Polygonatum odoratum
III,+ II,+ I,+ III,+ .
II,+ I,+
Rosa davurica
II,+ III,+ IV,2 V,1 I,1 IV,1 V,1
Spiraea pubescens
V,2 I,+ II,1 III,1 III,1 II,1 II,1
Ostryopsis davidiana
III,1 II,1 II,1 IV,2 IV,2 II,1 V,3
Sedum aizoon
V,+ II,+ III,+ II,+ V,+ III,+ IV,+
Polygonum viviparum
.
I,+ II,+ .
.
.
.
Picris japonica
I,+ II,+ II,+ II,+ .
.
I,+
Ranunculus japonicus
.
.
r,+ .
.
.
.
Carex appendiculata
.
.
.
.
.
.
.
Carex korshinskii
.
.
.
.
I,+ .
1,1
Cleistogenes squarrosa
.
.
.
.
.
.
II,+
Artemisia capillaris
I,+ .
.
.
.
.
I,+
Agrimonia pilosa
III,+ III,+ III,+ II,+ .
.
I,+
Artemisia lavandulaefolia
II,+ III,+ III,+ II,+ I,+ .
I,+
Thalictrum baicalense
III,+ II,+ II,+ II,+ .
.
I,+
Quercus mongolica
V,3 .
I,+ II,1 .
I,+ .
Dictamnus albus
III,1 I,+ I,+ I,+ .
.
.
Betula platyphylla
.
IV,2 V,4 II,1 I,1 III,1 .
Saussurea ussuriensis
I,+ V,+ IV,+ I,+ I,+ II,+ .
Betula dahurica
I,1 V,4 r,+ .
.
.
.
Galium aparine var. tenerum
I,+ V,+ II,+ II,+ .
I,+ I,+
Potentilla fragarioides
I,+ IV,+ I,+ I,+ .
.
.
16
8
8
0
2
75
II,+
V,+
I,+
II,+
IV,1
.
.
.
V,1
IV,+
.
.
II,+
.
I,+
.
.
.
.
.
.
II+
.
III,+
II+
9
5
0
<1
65
.
.
.
.
.
.
.
.
I,1
.
V,1
IV,2
.
.
.
I,+
.
.
.
.
.
.
.
.
.
10
16
0
<1
45
+,+
I,+
+,+
.
I,+
.
+,+
.
11
39
0
<1
35
.
.
.
.
r,+
.
.
r,+
12
8
30
25
35
II,+
.
.
.
II,+
I,1
.
.
+,+
.
.
III,1
II,+
+,+
I,+
.
.
.
.
.
+,+
.
.
.
.
.
.
III,+
IV+
III+
.
r,+
.
.
.
.
.
.
.
.
.
.
.
IV,+
IV,+
IV,+
.
.
.
.
.
.
.
.
.
.
Community type
Adenophora borealis
Prunus padus
Adenophora divaricata
Populus davidiana
Pinus tabulaeformis
Picea meyeri
Thalictrum squarrosum
Cotoneaster melanocarpus
Stipa baicalensis
Filifolium sibiricum
Leymus chinensis
Ulmus pumila
Dontostemon eglandulosus
Androsace septentrionalis
1
I,+
.
I,+
I,1
.
.
.
.
I,+
.
.
.
I,+
I,+
2
IV,+
IV,1
III,+
II,1
.
.
II,+
I,1
.
.
.
.
.
.
3
II,+
I,+
I,+
I,1
.
.
II,+
II,+
.
.
.
.
.
I,+
17
4
I,+
II,1
I,+
V,4
.
.
II,+
I,+
.
.
.
.
.
+,+
5
.
.
.
.
V,3
.
II,+
.
I,+
.
I,+
.
I,+
.
6
.
I,+
.
.
.
V,4
IV,+
IV,+
.
.
.
.
.
.
7
.
II,+
.
.
.
.
II,+
.
I,+
.
.
.
I,+
II,+
8
I,+
.
I,+
.
.
.
.
.
II,+
.
I,+
.
.
.
9
.
.
.
.
.
.
.
.
.
.
II,+
.
.
.
10
.
.
.
.
.
.
II+
.
V,2
VI,1
III,+
.
I,+
.
11
.
.
.
.
.
.
II,+
.
II+
r,+
V,1
.
II,+
I,+
12
.
.
.
.
.
.
II,+
.
.
.
II,+
V,2
IV,+
IV,+
Figure captions:
Fig. 1 Location of the study area and its geomorphologic features.
The black quadrate in the inside map indicates the location of the study area. The number 1,2,3,4 represents
the woodland zone, the woodland-grassland zone, the woodland steppe zone and the steppe zone
respectively.
Fig. 2 Contour map of selected climatic parameters in the study area.
D and J represent the sites of Duolun and Jingpeng respectively
Fig. 3 DCA result of woodlands, shrublands and open woodland
Fig. 4 DCA result of steppes and meadow
Fig. 5 Differentiation of climate, altitude and vegetation from SE to NW in the study area. In this figure, WI
and HI represent the Warmth Index and Humidity Index respectively, for the calculation of which please
refers to the text.
Fig. 6 Contour map of percentages of different ecological groups of species and the ratio of group II to
group I
18