Structure of vegetation of steppe hills in the Altai piedmont

Structure of Vegetation of Steppe Hills in the Altai Piedmont

A. Yu. Korolyuka, A. V. Egorova

a, I. E. Smelansky

b, and N. V. Filippova

c

a

Central Siberian Botanic Garden, Siberian Branch of the RAS, ul. Zolotodolinskaya 101, Novosibirsk, 630090 Russiab

Siberian Environmental Center, ul. Pirogova 20/2, Novosibirsk, 630090 Russiac

Tomsk State University, prosp. Lenina 36, Tomsk, 634050 Russia

Abstract—The phytocenotic diversity of the Altai steppe piedmont is described. Plant communities were

ordinated along the moisture gradient. The results were used to analyze the structure of vegetation of steppe hill

massifs. Distribution of steppe vegetation over an ideal hill was simulated, and the main patterns of distribution

were typified.

DOI: 10.1134/S1995425508010062

INTRODUCTION

The Altai piedmont is one of few Russian regions

where steppe ecosystems have remained on vast areas

in all their diversity. We observe there rich floras and

vegetation communities [1, 2]; in steppe hill massifs,

many plant species and types of phytocenoses exist,

which are rare and merit protection [3, 4]. Vegetation of

the piedmont of northern and western Altai is studied

less thoroughly than in the adjacent regions. The avail-

able publications give at best a general idea of dominat-

ing types within geobotanical zones [5, 6]. There are

only few papers describing the vegetation of some

steppe massifs in detail [7–9]. Analysis of new field

data, together with maps and remote sensing data, al-

lows us to describe the contemporary state of vegeta-

tion in the Altai piedmont. Given the considerable

anthropological impact and fragmentation of plant

cover, this is a necessary step to develop measures for

preserving the steppe biota.

AREA UNDER STUDY

This paper is based on data of the 1994 field studies

conducted in Tret’yakovo, Lokot’, Zmeinogorsk, Ku-

r’ya, Krasnoshchekovo, Charysh, Shipunikha, and Ust’-

Kalman districts of Altai krai. The territory under study

is a piedmont plain and hill uplands of north- western

Altai [5] and northern cis-Altai physicogeographical

province: Mid-Charysh and Upper Alei regions [10].

The steppe piedmont is located in the Alei, Charysh

and Anui basins to form a belt up to 50 km wide,

stretching from southwest to northeast for over 200 km.

The foothills fringe the Altai upland from northeast and

are a regular combination of elevated valleys and low

mountains. The ridge relief, frequently partitioned by

erosion to ridge-and-hill and hill relief, is dominant

there, often with vast valleys and gullies. In many

cases, hill massifs border the valleys of large rivers

(Alei, Charysh, and their main tributaries) and marginal

zones of sharp tectonic cliffs (“northern front of Altai”).

Frequently, hill massifs consist of ancient hills buried

under loose alluviums re-exhumed by denudation [11].

The inner parts of watersheds are distinguished by

rather poorly differentiated and gently undulatory re-

lief, which is disrupted only by relict hills. In the south

numerous granite outcrops are present in the relief as

relict rocks often characterized by pillow cleavage.

Within the steppe piedmont, from southeast to

northwest, the climatic zone of dry steppe grades into

the zone of piedmont and low-mountain steppe [12].

These zones correspond to semiarid, dry subhumid

zone, and weakly drought-affected subhumid zones ac-

cording to the UNEP classification of climate aridity

[13]. In general, the climate is continental and is charac-

terized by low winter temperatures (down to –50°C, av-

erage January temperature is –16.4 to –15°C) and high

summer temperatures (up to +40°C, average July tem-

perature is +19 to +20.4°C). Annual rainfall makes up

380 mm in the zone of arid steppe to 550 mm in the

zone of low-mountain steppe, and the moisture coeffi-

cient varies from 0.25 to 0.60. Light frosts in spring and

summer droughts are rather common. The scatter of cli-

matic indicators throughout the piedmont is quite regu-

lar and is controlled by decreasing aridity and tem-

perature contrast of climate from valleys to moun-

tains [12].

This paper is aimed at analyzing typological diver-

sity and structure of vegetation of the steppe piedmont

of Altai. Our tasks were to distinguish the remnant

steppe massifs, to describe phytocenotical diversity of

the steppe type of vegetation, and to analyze the struc-

ture of vegetation of steppe hill massifs determined by

spatial change of moisture character.

MATERIALS AND METHODS

This paper is based on about 800 releves, including

over 400 releves of steppe communities. To store and

analyze materials, we used the IBIS software [14] and

ISSN 1995-4255, Contemporary Problems of Ecology, 2008, Vol. 1, No. 1, pp. 49–58. © Pleiades Publishing, Ltd., 2008.

Original Russian Text © A.Yu. Korolyuk, A.V. Egorova, I.E. Smelansky, N.V. Filippova, 2005, published in Sibirskii Ekologicheskii Zhurnal, 2005,

Vol. 12, No. 6, pp. 999-1011.

49

original processing software. Latin names of the plants

are given according to Cherepanov [15].

To compare species richness of different kinds of

steppe, we used average species saturation on a site

100 m

2

in area and species richness of the cenoflora

core, i.e. species with a frequency over 20%. The spe-

cies richness of a cenoflora depends strongly on the

number of releves while the species richness of its core

is more constant. It has empirically been established

that this indicator is stable when there are over 8–12

releves.

The steppe massifs that escaped from ploughing up

were distinguished by processing land-use maps (scale

1:100 000) and Landsat-7 ETM+ images. According to

the land-use maps, we singled out tillages, pastures,

hayfields, shrubs, forests, and inhabited areas. Pastures

and hayfields were classified into nonimproved and im-

proved; in addition, we marked out tame perennial

grasses in croplands. Pastures, shrubs and hayfields

were considered slightly disturbed grasslands. The next

step was smoothing the borders of grasslands using the

averaging filter of the Spatial Analyst module (Arc-

View3.2a GIS). The map was cleared of all polygons

whose linear dimensions in all directions did not exceed

50 m; also, all narrow strips connecting steppe massifs

were eliminated from the map. As a result, a geo-

information system was assembled, with one of its lay-

ers being a map of distribution of existing steppe

massifs (Fig.1).

To ordinate plant communities, ecological scales of

plants were used [16]. The status of a particular descrip-

tion and richness-salinity of soil were computed using

the method proposed by Canadian ecologists [17, 18]:

Stat

opt i

N

i

N

�

�

�( )

1

,

where Stat is the status of releve, opt(i) is the optimum

of the i-th species along the axis of a factor (moisture or

richness-salinity), and N is the number of species in the

releve. The optimum of the species is the mean ampli-

tude of the greatest species abundance, taken from the

ecological scales [16]. The status of a syntaxon was cal-

culated as the arithmetical mean of the statuses of its

releves. For all types of steppe, we have computed aver-

age moisture (M) and richness-salinity (RS) of soils.

As in most other steppe regions, grasslands in the

Altai piedmont escaped from ploughing up mainly in ar-

eas not suitable for tillage, on hill slopes and between

small hills. In these landscapes, the structure of vegeta-

tion is determined by the distribution of communities

over slopes of various steepness, exposure, and shape.

In order to equally and formally describe vegetation of

the hill massifs, we have developed a graphic model of

an “ideal hill” (Fig. 2). It is oriented according to cardi-

nal points and is, correspondingly, divided into a series

of sectors. Four circular structures are slopes of

different steepness: the outer ring comprises plain sites,

followed by zones of gentle (up to 5°) and medium

steepness (5 to 15°), and steep slopes (over 15°); and

the inner ring is composed of bulging stony tops of hills

and ridges. For each large steppe massif, the releves

were arranged in the hill scheme according to the expo-

sition and steepness of the slope. The status of moisture

was calculated for each releve, and a density model was

compiled to reflect distribution of communities with

different degree of moisture over the hill slopes. Phyto-

cenoses of shallows were eliminated from analysis as

they are moist irrespective of exposure. Their use for

analysis would lead to a significant variance of mois-

ture indicators on the slopes of any exposure. Neverthe-

less, an analysis of concave fragments of the slopes, in

our case mainly occupied by shrubs, is important as the

communities inhabiting them are the integral part of

vegetation of hill massifs and frequently imparts pecu-

liarity to the landscapes. For this reason shrub commu-

nities of shallow gullies are used to characterize and

sometimes entitle types of combinations of hill vegeta-

tion. In our opinion, applying a unified method of

ecocenotical analysis of vegetation will allow a correct

approach to a relative study of vegetation of hill massifs

and then to geobotanical zoning.

RESULTS AND THEIR DISCUSSION

The diversity of vegetation of the Altai piedmont is

composed of steppe, shrub, meadow, forest, and wet-

land types. Grasslands are predominant in the land-

scapes, and of them, steppes of the Kazakhstanian type

prevail. According to the Russian steppe science [19,

20], within the studied area, there are subtypes of

meadow and true steppes, the latter being restricted to

the class of true bunchgrass steppe, divided into two

groups of formations, rich and poor forbs. The frame-

work developed for steppes of Central Kazakhstan [21]

is most appropriate for the steppes of Altai. The frame-

work includes four latitudinal strips of the first level,

corresponding to the classes of formations: meadows,

reach forbs- and forbs-bunchgrass steppes dominated

by Stipa spp., dry bunchgrass steppes dominated by

Stipa and Festuca spp., and desertified steppes. Our

studies have shown that there are no desert steppes in

the Russian Altai piedmont. Evidence of their presence

in the southeastern part of West Siberia [22, 23] is erro-

neous, and the communities described as desert steppes

are arid variants of true steppes or solonetz communi-

ties. Zonal variant of the arid steppe was described by

us only from one locality at the southern border of the

territory. In true steppes, rich forbs-bunchgrass steppes

are dominant and forbs-bunchgrass steppes are domi-

nant too, although to a lesser extent. In some areas of

the piedmont, especially in the eastern, more elevated

part, meadow steppes and steppe-like meadows domi-

nate. The diversity of steppes of the surveyed territory

is represented by 11 types of communities, three of

CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 1 No. 1 2008

50 KOROLYUK et al.

which are zonal vegetation communities of true and

meadow steppes and the rest are petrophytic and

calciphytic communities (Table 1).

Zonal Variants of Steppe

Dry steppe is described only at one locality on the

border of Altai krai and Kazakhstan (Berezovka River,

Lokot’ district). The community is located in the belt

along the border of the river valley and is restricted by

tillage at watershed. Stipa sareptana dominates there,

and Stipa capillata and Festuca valesiaca are codo-

minants (Table 2). Thirteen plant species have been reg-

istered at the plot with the plant cover of 50%.

Forbs-bunchgrass steppes dominated by Stipa capil-

lata and Festuca valesiaca are common in the piedmont

and, together with stipa steppes (Stipa capillata, Stipa

lessingiana), form an arid wing of the steppe vegeta-

tion. They are true steppes but to attribute particular

releves either to forbs-bunchgrass or rich forbs-bunch-

grass variants is often difficult because they are dis-

turbed by overgrazing. Poor floristic composition, dull

colors, and absolute dominance of Festuca valesiaca

and Stipa capillata allow them to be classified as forbs

steppes. Lavrenko [19] defined their status in the same

way after analyzing forbs-bunchgrass steppes of the

woodless meadow subzone of West Siberia described

by Krylov [24]. At the same time, one should account

for possible replacement of rich forbs-bunchgrass

steppe by needle-grass fescue steppe as a result of

long-term overgrazing. The secondary character of

some plots of forbs-bunchgrass communities is ex-

pressed in the participation of relatively mesophytic

plants as well as in the presence of suppressed domi-

nants of rich forbes steppes, Stipa zalesskii and He-

lictotrichon desertorum. The forbs-bunchgrass steppes

dominated by Stipa capillata and Festuca valesiaca are

especially widespread in the western part of the area un-

der study. They occupy nonploughed plain sites, slope

fans, and river terraces and also occur in most arid west-

ern parts of the piedmont along minor gentle depres-

sions surrounded by dry petrophytous steppes. Habitats

are slightly rubbled; only sometimes one can observe

solitary medium-sized and small stones on the soil sur-


STRUCTURE OF VEGETATION OF STEPPE HILLS 51

Fig. 1. Map of distribution of steppe massifs. Study site: 1 – Ust’yanka, 2 – Ploskoe, 3 – Shipunikha, 4 – Ruch’evo, 5 – Trusovo, 6 – Ozerki,

7– Suetka.


52 KOROLYUK et al.

Fig. 2. Models of hills.

face, which were brought there from adjacent slopes.

Most of the described sites of steppes are disturbed by

overgrazing. Fescue-stipa communities are close as the

plant cover makes up 60–80%, with the two-layer verti-

cal structure. Low bunchgrasses are dominant (Festuca

valesiaca and Koeleria cristata) forming the lower sub-

layer, which mainly consists of pasture-resistant worm-

wood (Artemisia austriaca). The upper sublayer is

composed of needle-grass Stipa capillata. During its

mass flowering and fruitage the communities look like

monodominant needle-grass areas. The communities

are poor, the average number of species is 25 per

100 m

2

. The piedmont forbs-bunchgrass steppes are

characterized by persistent low abundance of petro-

phytic plants. As a rule, these are facultative petro-

phytes such as Centaurea sibirica, Artemisia commu-

tata, Allium rubens, and others. The communities with

insignificant number of these species are considered

hemipetrophytic variants of true steppes. Their vast oc-

currence is related to the almost complete ploughing of

plains occupied by steppes without petrophytes. Non-

ploughed zonal steppes were preserved near villages or,

in the form of tiny contours, on field margins. Sparse

plant coverage and high portion of stony habitats

occupied by bunchgrass petrophytic steppes mean

greater participation of petrophytic plants in species

composition of the fescue-stipa steppes.

Rich forbs-bunchgrass steppes are the prevalent

zonal type in the steppe piedmont of Altai. They occupy

gentle plain and slightly convex slopes of different ex-

posure; the soil surface is very little rubbled, with soli-

tary buried big stones observed there. One can suggest

that the ploughed areas were earlier occupied by rich

forbs-tall bunchgrass steppes. Sites of the steppes

which escaped from total ploughing up are mainly rep-

resented by hemipetrophytic variants of Helictotrichon

desertorum—Stipa zalesskii steppes. Nearly all sites of

this kind are more or less intensely used as pastures and,

less often, as hayfields. Overgrazed communities are

close to forbs-bunchgrass fescue-stipa steppes in the

habit and dominants but differ from them in the pres-

ence of mesoxerophylic species. Stipa zalesskii is re-

placed by Stipa capillata here and many forbs species

are absent.

Relatively high moisture of the Altai piedmont,

which is due to submontane moisture zoning, deter-

mines the richness and abundance of species of rich

forbs-bunchgrass steppes. The shrub layer is usually

not developed and the base of communities is formed

by four equally ranked dominants, bunchgrass Stipa

capillata, S. zalesskii, Helicotrichon desertorum, and

Festuca valesiaca. Joint plant cover of these species

may reach 50–60%. Diversity and abundance of peren-

nial forbs (Iris ruthenica, Artemisia latifolia, Pulsatilla

patens, Medicago falcate, Thymus marschallianus,

Fragaria viridis, and many others) determine a colorful

habit of the steppes. The communities are rich in spe-

cies, with 45 species per 100 m

2

, and even 65–70 in

some communities.

Rich forbs-bunchgrass meadow steppes are ob-

served in southern, more elevated and rugged, territo-

ries grading into forest-steppe and meadow-steppe

hills. Meadow steppes occur in different parts of slopes

of the eastern, northern and western exposition. In

moist forest-steppe areas where shadowed slopes are

occupied by larch or deciduous forests, meadow



Table 1. Types of steppes of the Altai piedmont

Class of formations

according to [21]

Communities

zonal petrophytic calciphytic

Dry bunchgrass steppes Dry steppe with Stipa

sareptana, S. capillata, and

Festuca valesiaca

Petrophytic steppes with

Stipa capillata, Artemisia

frigida, and A.

sublessingiana

Forbs-bunchgrass steppes

with Stipa lessingiana and S.

capillata

Forbs-bunchgrass steppes Forbs-bunchgrass steppes

with Stipa capillata and

Festuca valesiaca

Petrophytic bunchgrass

steppes with Festuca

valesiaca, Stipa capillata,

and Koeleria cristata

Forbs-bunchgrass steppes

with Stipa lessingiana

Rich forbs-bunchgrass

steppes

Rich forbs-bunchgrass

steppes with S. zalesskii, S.

capillata, and Helictotrichon

desertorum

Petrophytic bunchgrass

steppes with Festuca

valesiaca, Helictotrichon

desertorum, and Carex

humilis

Calciphytic

forbs-bunchgrass steppes

with Helictotrichon

desertorum. Carex humilis,

and Stipa zalesskii

Meadow steppes Rich forbs meadow steppesPetrophytic

shrub-bunchgrass steppes

with Carex humilis,

Helictotrichon desertorum,

and Spiraea trilobata

Observed in low mountains;

probably absent from

piedmont


54 KOROLYUK et al.

steppes occur on gentle plain or convex southern

slopes. The shrub layer in meadow-steppe communities

is poorly developed. The herbage is dense, with the

plant cover of 80–95%, and its base is composed of

constant dominants: Helictotrichon desertorum in the

upper layer and Carex humilis in the lower layer, with

the total cover averaging about 30%. Three other spe-

cies are codominants and less often dominants; these

are Stipa zalesskii, Iris ruthenica, and Filipendula vul-

garis. A picturesque character of herbage is determined

by abundance of many species including Artemisia

sericea, Poa transbaicalica, Dracocephalum ruyschi-

ana, Fragaria viridis, Carex pediformis, Helictotri-

chon pubescens, Festuca valesiaca, and Pulsatilla

patens. The species saturation averages 64 species per

100 m

2

.

Petrophytic Variants of Steppes

Petrophytic steppes hold the lead in area among

piedmont natural rangelands. The reason is the com-

plete ploughing-up of zonal habitats and vast develop-

ment of hill relief, where stony slopes are common.

Petrophytic steppes are located on the slopes of differ-

ent steepness, as a rule, convex ones, and at the tops of

hills and ridges. The communities may be diagnosed by

plants which are usual for stony arid habitats of Altai:

Allium rubens, Youngia altaica, Patrinia intermedia,

Veronica pinnata, Seseli buchtormense, Orostachys

spinosa, Gypsophila patrinii, Artemisia commutata,

Artemisia frigida, Centaurea sibirica, Sedum

hybridum, Coluria geoides, etc. Stony steppes are di-

vided into four variants and each of them corresponds

to a certain zonal type.

Petrophytic Stipa-Artemisia steppes are considered

variants of the dry steppe subtype. These communities

are observed only on the right bank of the Alei river up

to Gilevo Village and south of the town of Gornyak.

The communities occupy southern arid and stony

slopes and ridges of hills and uplands. We have com-

piled only two releves, which is not sufficient to distin-

guish in full measure the composition and structure of

cenoflora. Rare plants growing there at the northwest-

ern border of the habitat are: Galitzkya spathulata,

Silene incurvifolia and Artemisia sublessingiana.

Petrophytic short bunchgrass steppes are stony vari-

ants of forbs-bunchgrass steppes dominated by Stipa

capillata and Festuca valesiaca. They are frequent in

piedmont hill massifs of the steppe zone in Altai krai.

They are most usual in the western part of the region.

Habitats differ in stoniness as some areas can be cov-

ered by detrital rocks and bedrock outcrops for

70–80%. Depending on this factor, plant cover of forbs

varies from 20 to 80%. The structure of the communi-

ties is determined by dominance of bunchgrass and

petrophytic forbs. The main dominant with the plant

cover of 10–30% is Festuca valesiaca, forming a sub-

layer 8–15 cm high. Stipa capillata, Koeleria cristata,

and Artemisia austriaca are codominants. Sometimes

wormwoods Artemisia frigida and Artemisia commu-

tata are abundant in the communities, as well as repre-

sentatives of petrophile forbs Gypsophila patrinii,

Sedum hybridum, and Orostachys spinosa. The shrub

layer is not developed in most comminities but the vari-

ants with abundant Spiraea hypericifolia or Juniperus

sabina are also common. The communities are not rich,

their species saturation on the average is 30 species per

100 m

2

which is somewhat greater than in zonal steppes

whose petrophytic variant they are. Of rare communi-

ties, one should note eastern-stipa (Stipa orientalis)

steppe described by us at the sole locality on the right

bank of the Alei River in the Lokot’ district [4].

Petrophytic bunchgrass steppes (serial variants of

bunchgrass steppes) are one of dominating elements of

vegetation in Altai krai. In addition to grass, turf sedges

Carex humilis and Carex pediformis may dominate the

communities. The described stony steppes occupy the

same habitats as their drier analogs but areas with their

prevalence are located nearer to low mountains (to the

south and southeast), absolute altitude marks are a bit

greater and moisture is also greater. The communities

are dense and their average plant cover is 60%. The

structure and habit of each community is determined by

the ratio of short and tall bunchgrass. Short bunchgrass

(Festuca valesiaca, Carex humilis, and Koeleria crista-

ta) can cover up to 40–45% of the soil. The cover of tall

bunchgrass (Helictotrichon desertorum and Stipa za-

lesskii) reaches 35–40% in some communities. The

community usually consists of two sublayers 8–15 cm

and 30–40 cm high. The ratio of dominant biomorphs

may also vary and thus leads to the diversity of variants

easy to distinguish by the eye. The analyzed 76 releves

of stony steppes were divided into three groups:

27 releves were given to short bunchgrass steppes (the

cover of short bunchgrass is 3 and more times over the

cover of tall bunchgrass), 43 releves to tall and short

bunchgrass steppes (approximate equality), and six to

tall bunchgrass steppes (tall bunchgrass cover is more

than 3 times over that of short bunchgrass). Thus,

among stony steppes of this type, communities with

well-expressed short and tall bunchgrass layers are

dominating, and short bunchgrass variants are also

common. Of characteristic features, worthy of note is

the abundance of forbs, usually only a little inferior to

bunchgrass and having an average cover of 20–25%.

This determines the rich forbs habit of the communities,

especially in early summer, in the season of mass flow-

ering. Species richness makes up 42 species per 100 m

2

and is close to that of zonal rich forbs and tall bunch-

grass steppes, with these communities being its petro-

phytic variant.

Petrophytic shrubs-bunchgrass steppes dominate

over southern stony slopes under the conditions of for-

est-steppe hills of the Altai periphery. They are more

mesophytic and mountainous than other stony steppes

and the area of their mass occurrence is related to the



forest-steppe low mountains of the Altai-Sayan moun-

tain. Small contours of the communities are also fre-

quent in the north of the piedmont but their habitat is

not continuous there, as they are relatively rare. Shrub-

bunchgrass steppes occupy mainly steep (10–15°)

slopes exposed to light. The structure of communities is

determined by a vast development of shrub and grass

layers. The former is dominated by Spiraea trilobata,

with the cover of 10–30%. The herbage is dense and ho-

mogeneous. The base of the lower layer is Carex hu-

milis. The upper layer is Helictotrichon desertorum.

Together with sedge, it covers 30 to 50% of the ground.

There are no constant codominants here but many spe-

cies may occur in abundance, such as Thymus serpyl-

lum sensu lato, Festuca valesiaca, Sedum hybridum,

Artemisia gmelinii. Iris ruthenica, Pulsatilla patens,

Carex pediformis, Stipa capillata, Patrinia sibirica,

Coluria geoides, Stipa pennata, and others. Average

species saturation is 51 species per 100 m

2

.

Calciphytic Variants of Steppes

Dry forbs-stipa steppes are most xerophytic variants

of steppes in the Altai piedmont. They are classified as

a subtype of dry steppes and are their calciphytic vari-

ant lying on carbonate soils. Within the studied area,

they are rare communities, not covering vast areas, and

their occurrence is limited to the north by the right bank

of the Alei river (upstream to the village of Gilevo). The

communities are floristically poor but their composi-

tion includes many species rare in Russia which, along

with rareness and vulnerability of stipa steppes, deter-

mines their high conservation value [4]. They are in

peril to a greater degree than the other steppes in plains

and steppe piedmont of Altai. This is related to the fact

that sites occupied by this type of phytocenoses are

small in size, most of them are subject to overgrazing

and are often used for country roads, garbage dumps,

and quarries. These steppes are unique in the Altai terri-

tory because they have an arid steppe florocenotical

complex of the Kazakhstanian type at its northeastern

margin. The habit and structure of the communities is

determined by the dominance of forbs (S. lessingiana,

S. capillata, and sometimes S. korshinskyi) over the up-

per sublayer of herbage, with the plant cover of up to

40–60%. The lower sublayer is significantly less devel-

oped (average plant cover is about 20%), and is com-

posed of fescue (Festuca valesiaca) and wormwoods

(Artemisia schrenkiana and A. austriaca). Dry steppe is

one of the poorest steppe types in the south of West

Siberia. Average species saturation is 12 species per

100 m

2

.

North of the Gilevo reservoir, we have described

floristically richer variants of forbs-stipa steppes con-

sidered calciphytic variants of forbs-stipa steppes. Hab-

itats occupied by rich Stipa lessingiana steppes are the

same as above. Probably their floristical richness is to a

certain extent related to climatic change as a result of

the influence of the Gilevo reservoir.

Calciphytic forbs-bunchgrass steppes are described

from the limestone massif in the basin of the Loktevka

River where they form large tracts on convex stony tops

and slopes of hills. The communities of this type are

rare, and analysis of their occurrence requires special

studies of the limestone massifs. In the upper layer of

herbage, Helictotrichon desertorum and Stipa zalesskii

are dominants while the lower layer is dominated by

Carex humilis and representatives of petrophile forbs

Eritrichium altaicum, Dracocephalum discolor, Thy-

mus serpullum s. l., and Orostachys spinosa. Abun-

dance of Stipa korshinskyi, a calciphylous plant, is the

characteristic feature of the communities. Herbage is of

medium density, with a plant cover of 40–60%. Hori-

zontal and vertical structure of the community is deter-

mined by the character of distribution of tall and short

bunchgrass as well as low petrophile forbs on open

stony sites. Species saturation varies from 24 to 31 spe-

cies per 100 m

2

.

Spatial Structure of Vegetation

The main factor influencing spatial distribution of

plant communities of the steppe piedmont is relief-de-

pendent moisture. To locate different steppe subtypes

along the moisture gradient, we have analyzed the sta-

tus of each releve we made in the south of West Siberia,

Altai and Eastern Kazakhstan as well as numerous pub-

lished releves from Northern Kazakhstan [25, 26].

Analysis of ordination results shows that the described

steppes have the medium, meadow, and arid steppe

types of moisture [16]. This subdivision does not corre-

spond to the ecological status of steppe communities in

the south of West Siberia and Northern Kazakhstan, and

for this reason we studied grades of moisture in detail.

Average moisture statuses of different phytocenoses of

dry steppes lie in the segment of 41–45 grades, forbs-

bunchgrass steppes take up positions from 45 to 49, rich

forbs-bunchland steppes — from 49 to 53, and rich

forbs meadow steppes — from 53 to 57.

In GIS the steppe sites were ranged in area and seven

large massifs were selected (see Fig. 1). All of them

were characterized by more or less densely differenti-

ated hill-and-ridge or hill relief. Each massif has been

represented by a sufficient number of releves (35 and

more) related to slopes of various steepness and expo-

sure. For each massif, a model of the “ideal hill” was

developed (see Fig. 2). Comparative ecologo-phyto-

cenotical analysis of the sites suggested their classifica-

tion into three types.

1. True steppe type of hill massif is observed only in

the latitudinal section of the Alei River (from Lokot’ to

Gilevo Village) and in the Zolotukha River basin. This

type corresponds to the belt of forbs-stipa steppes.

These are most arid hill massifs of Altai krai. It is char-

acterized by the dominance of petrophytic short-bunch-


56 KOROLYUK et al.

grass steppes and vast occurrence of forbs-fescue-stipa

communities. Petrophytic forbs-stipa-Artemisia, dry

stipa and rich forbs-bunchgrass steppes are rare. Shrub

communities are common in very concave sites of the

slopes and broad gullies. We conducted a detailed study

of the hill massif near the village of Ust’yanka of Lokot’

district in the territory of the reserve “Loktevskii” now

in construction [8, 9].

2. True rich forbs steppe type of hill massif is a

moister variant characterized by dominance of rich

forbs-tall bunchgrass steppes and vast occurrence of

fescue-stipa cenoses. On northern slopes, fragments of

meadow steppes occur. In eroded relief and broad gul-

lies, there are some shrub communities. This type is

represented by hill massifs near the village of Trusovo

of the Kur’ya district and village of Ozerki of Shipu-

nikha district.

3. Rich forbs steppe type of hill massif is typical of

the Altai piedmont and is marked by codominance

of rich forbs-tall bunchgrass and petrophytic forbs

steppes. We have distinguished a separate shrub steppe

variant in it, characterized by development of poly-

dominant mesophile grass-shrub communities which

occupy concave forms of the relief: slopes, narrow and

broad gullies, and ravines. The shrub steppe variant is

more usual for stony low mountains in the territory of

Tret’yakovo and Zmeinogorsk districts. In Fig. 2, mod-

els of hills are based on releves from several sites in the

vicinity of villages of Shipunikha and Ploskoe (Tret’ya-

kovo district). Northeastward, the contribution of shrub

communities to the vegetation of hill massifs dimini-

shes, and rich forbs steppe variant becomes dominant

there (Suetka Village of Krasnoshchekovo district).

Within this type, we have described a calciphytic vari-

ant (Ruch’evo Village in Kur’ya district) in limestone

outcrops. It is characterized by vast development of

calciphytic stony steppes but at the same time, zonal

types are represented by rich forbs-tall bunchgrass

steppes without calciphytes.

The studies have shown the high degree of safety of

steppes in the Altai piedmont. Steppe vegetation was

preserved in vast areas in a series of massifs whose size

reaches 15 000 hectares and in one case even exceeds

22 000 hectares. Phytocenotic diversity of the steppes

is represented by 11 types of communities, from dry to

meadow steppes. Along the axis of moisture the steppes

of the Altai piedmont are characterized by a broad am-

plitude of 40–58 grades of moisture. Regarding the

structure of vegetation, we have marked out three types

and two variants of hill massifs being territorial units of

vegetation of mesocombination rank. The developed

system of territorial units can serve a basis for geo-

botanical zoning of the Altai piedmont. The proposed

method of ecologocenotical analysis requires unified

standards of vegetation description. One can recom-

mend that scheme of the hill be used in field studies on

the key site (hill massif, mountain ridge, or others) to

estimate the degree of its being filled up.

The works on study and preservation of steppes of

the Altai piedmont are conducting with the financial

support of the Russian Foundation for Basic Research

(grant 05-04-48212-a), of the Integration Project of the

Siberian Branch of the Russian Academy of Sciences

“The Altai Ecoregion” (grant 124), and with the aid

of the Weeden Foundation and Hewlett Foundation

provided via non-commercial organization Pacific En-

vironments (PERC, San Francisco, USA).

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58 KOROLYUK et al.

Documents

Structure of vegetation of steppe hills in the Altai piedmont