ISSN 1067-4136, Russian Journal of Ecology, 2016, Vol. 47, No. 1, pp. 68–73. © Pleiades Publishing, Ltd., 2016.

Original Russian Text © A.M. Rusanov, M.A. Bulgakova, 2016, published in Ekologiya, 2016, No. 1, pp. 56–61.

Composition and Abundance of Soil Macrofauna

in Pasture Ecosystems of the Steppe Zone of the Urals
A. M. Rusanov and M. A. Bulgakova
Orenburg State University, pr. Pobedy 13, Orenburg, 460018 Russia
e-mail: soilec@esoo.ru
Received February 16, 2015

Abstract—Changes in the abundance and species compositions of the main groups of soil invertebrates have
been analyzed in ecosystems long used as pastures and exposed to different degrees of grazing. The sum of the
results makes it possible to conclude that the zoological method of indication of pasture ecosystems can be
used in ecological monitoring of steppe landscapes.

Keywords: pasture ecosystems, xerophytic vegetation, pasture digression, Carabidae, Tenebrionidae, cherno-
zem, regeneration of landscapes
DOI: 10.1134/S1067413616010136

Steppe ecosystems with chernozem soils and plant
communities dominated by herbaceous xerophytes
(Mirkin and Naumova, 1997) have been actively used
as natural pastures during the past few decades. Long-
term effects of livestock grazing on steppe grasses have
been studied by Gorchakovskii and Ryabinina (1984),
Lavrenko et al. (1991), Mirkin (1984), Mullagulov
et al. (2009), Krivolutskii (1994), and others. Effects
of such land use on the genetic properties of soils have
been analyzed by Vladychenskii et al. (1994), Gur’ev
(1992), Mustafaev (1960), Rusanov (1993, 1995), and
Tesl’ (2009). Their studies deal with changes in the
chemical and physical properties of chernozems and
in the species composition of the natural vegetation,
but they provide little data on the effects of livestock
on the soil fauna.
The founder of the science of the biosphere Ver-
nadsky (1935), discussing the “ubiquity of life” on
Earth, put special emphasis on the soil as one of the
most important “living concentrations.” The soil
occupies the boundary between the lithosphere,
hydrosphere, and atmosphere; it is a product of the
interaction of these environments with each other and
with living organisms. The soil is a combination of ele-
ments in three states: liquid, solid, and gaseous, which
predetermines favorable conditions for the develop-
ment of living matter in its maximum biological diver-
sity. These conditions manifest themselves most
strongly in chernozem soils, which have a relatively
thick humus horizon where the entire spectrum of
nutrients is concentrated and the combination of ther-
mal, hydraulic, and pneumatic soil properties is opti-
mal. Hence, changes in plant formations and genetic
properties of chernozems in pasture biogeocenoses
should apparently be linked to changes in species com-
positions and quantitative parameters of soil organ-
isms.
Considerable progress has been made in the devel-
opment of zoological methods for diagnosis of differ-
ent soil types (Ghilarov, 1965; Krivolutskii, 1994;
Mordkovich, 1977; Striganova, 1980, 2006), but the
effects of overgrazing on the soil macrofauna of pas-
tures situated within the geographic range of a partic-
ular soil type has not been studied sufficiently. More-
over, it is necessary to further evaluate the feasibility of
zoological diagnostic methods for assessing the extent
of changes occurring in pasture ecosystems under the
impact of grazing. The purpose of this study was to
address these issues as applied to the steppe zone of the
Urals, using the example of the Cisural region. No
such studies have yet been performed in this region.
MATERIAL AND METHODS
Studies were performed in 2011 to 2013 in areas
with quasi-natural vegetation conditionally consid-
ered virgin (reference areas) and areas of pastures
degraded to different degrees on ordinary chernozems
of the true steppe located on the flattened Ural–
Sakmara watershed.
Geobotanical descriptions and diagnosis of test
areas with respect to the degree of pasture digression
were made according to Ramenskii (1971). Abo-
veground phytomass was determined by the hay-har-
vest method in 1 × 1-m plots, drying the plants to con-
stant weight in a drying oven. Belowground phytomass
was determined in 20 × 20 × 20-cm soil monoliths the
monoliths were washed with water on a sieve and the

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 COMPOSITION AND ABUNDANCE OF SOIL MACROFAUNA
Table 1. Average values of main parameters characterizing chemical and physical properties of chernozems
Parameter
virgin land
Humus content in layer 0–20 cm, % 4.8
Humus supply in layer 0–20 cm, t/ha 108.1
Degree of organic substance humification, % 41.2
Chumic acids : Cfulvic acids 2.22
Soil density, g/cm3 1.07
Rate of water permeability, mm/h 126
Coefficient of structuredness 1.62
extracted parts of plants were dried and weighed. All
samples were taken in three replicates. The abo-
veground, belowground, and total phytomass values
were recalculated and expressed as metric centners per
hectare (cwt/ha).
Soil density and moisture were determined by vol-
umetric and gravimetric methods, respectively
(Vadyunina and Korchagina, 1986); total humus, by
Tyurin’s method (Agrokhimicheskie…, 1975); its frac-
tion-group composition, by Tyurin’s method modi-
fied by Ponomareva and Plotnikova (see Kaurichev,
1980).
Zoological samples were taken by the method of
Ghilarov (1965) from 0.25-m2 plots (50 × 50 cm)
evenly distributed over each test area, in 15 replicates.
Each sample was taken to the depth at which inverte-
brates ceased to occur. A total of 360 samples were
analyzed, and over 10 000 soil invertebrates were
extracted. Beetles were collected by lines of pitfall
traps filled with a fixative (formaldehyde solution)
(Ghilarov, 1965; Mordkovich, 1977). The total
amount of work was 2400 trap–days.
RESULTS AND DISCUSSION
The degradation of pastures in the study region
resulted from long-term irrational use, especially after
the development of virgin and fallow lands in 1954–
1962, when cultivated areas markedly increased at the
expense of natural pastures and the numbers of graz-
ing livestock increased as well; as a result, the pressure
on pasture ecosystems became much greater than their
ecological capacity allowed.
The quasi-natural vegetation of the reference area
was represented by a herb–feather grass–fescue com-
munity (Stipa lessingiana Trin. еt Rupr. + Festuca vale-
siaca Gaudin) with 85–90% coverage. Although this
area was conditionally considered virgin, sporadic
occurrence of Stipa tirsa Stev. and Artemisia austriaca
Jacq. indicated that it had been exposed to an insignif-
icant grazing impact. The weakly degraded pasture
area was occupied by a wormwood–fescue–feather
grass community (Festuca valesiaca Gaudin + Arte-
RUSSIAN JOURNAL OF ECOLOGY Vol. 47 No. 1
69
Degree of vegetation digression
slight moderate strong
4.6 4.5 4.1
108.6 110.0 106.6
42.5 41.4 36.1
2.31 2.06 1.79
1.08 1.26 1.34
96 77 65
1.22 1.12 1.06
misia austriaca Jacq. + Stipa lessingiana Trin. еt Rupr.)
with 70–75% coverage. The moderately degraded area
was occupied by a June grass–bluegrass–fescue com-
munity (Koeleria gracilis Pers. + Poa bulbosa L. + Fes-
tuca valesiaca Gaudin.) with 40–45% coverage. The
area under a fescue–June grass–wormwood commu-
nity (Artemisia austriaca Jacq. + Koeleria gracilis Pers. +
Festuca valesiaca Gaudin.) with 15–20% coverage was
considered classified as a strongly degraded pasture.
The stocks of the aboveground and belowground (0–
20 cm) phytomass in the series from conditional virgin
land to strongly degraded pasture decreased by a factor
of 4.2, to 2.3 metric cwt/ha.
Long-tem excessive anthropogenic pressure on
pasture ecosystems results not only in alteration of flo-
ristic and geobotanical parameters of natural vegeta-
tion but also in deterioration of quantitative and qual-
itative parameters of organic matter and physical
properties of chernozem soils. Thus, the content of
humus in the studied soils decreased from 4.8% in the
reference area to 4.1% in the strongly degraded pas-
ture, with the density of the upper soil layer increasing
by 25.2% (Table 1).
The impact of long-term, uncontrolled livestock
grazing on pasture ecosystems manifested itself not
only in digression of vegetation and impairment of
genetic soil properties, but also in the species compo-
sition and abundance of soil invertebrates. The exper-
iment made it possible to determine the abundance
and recognize the dominant groups of soil inverte-
brates living in pastures of different degrees of digres-
sion.
A total of 49 species of beetles (Coleoptera) from
seven families were recorded over the study period in
the virgin and pasture areas (Table 2). Most of these
species (12) belonged to Carabidae (ground beetles);
the second most abundant family (eight species) was
Tenebrionidae (darkling beetles); and the third most
abundant family (four species) was Silphidae (carrion
beetles). In addition to beetles, earthworms (Lumbri-
cidae), centipedes (Chilopoda), true bugs (Hemip-
tera), and arachnids (Arachnida) were collected.
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70 RUSANOV, BULGAKOVA

Table 2. Estimated abundance of soil macrofauna in test areas

Degree of pasture digression
Main groups of soil invertebrates
virgin land slight moderate strong
Elateridae S – – S
Dermestidae SD R S D
Carabidae D D D D
Order Coleoptera

Curculionidae S – – –
Tenebrionidae D D D D
Class Insecta

Family

Scarabaeidae S S – SD
Silphidae S – – R
Order Hemiptera S – – –
Class Chilopoda R S S –
Order Acari S – – –
Class Arachnida
Order Araneae S S R R
Class Oligochaeta Order Lumbricidae R R S –
S, sporadic, abundance less than 1%; R, rare, abundance 1–3%; SD, subdominant, abundance 3–10%; D, dominant, abundance over 10%.

Table 3. Species composition and distribution of Carabidae in test areas

Degree of pasture digression
Species
virgin land slight moderate strong
Amara aenea Deg. + + + +
Calathus erratus C.Sahlb. +
C. halensis Schall. + +
C. melanocephalus L. + + +
Cymindis angularis Gyll. + + + +
Harpalus rufipes Deg. + + +
H. smaragdinus Duft. + + + +
H. calceatus Duft. + + + +
Ophonus stictus Steph. + + + +
Poecilus lepidus Leske +
P. sericeus Fisch. + + +
P. versicolor Sturm +
(+) Species presence in the area.

The recorded ground beetles belong to six genera
(Table 3). In contrast to the reference area, which was
colonized by typical mesophilic ground beetles, pas-
tures were colonized by species typical of xerophytic
vegetation (Gaevskaya, 2011; Rusanov and
Gaevskaya, 2011).
The abundance of zoophagous species, above all
litter and litter–soil stratobionts, was lower in pasture
ecosystems than in conditional virgin areas. This is
obviously explained by very poor development of the
litter in the moderately and strongly overgrazed areas.
The total number of zoophagous ground beetles
decreased because many detritophagous species, on
which they feed, disappeared from the biocenosis.
If we examine in detail the relative abundance of
different life forms in the ground beetle assemblages,
the dynamics of their abundance clearly manifests
itself. Thus, the proportion of mixophytophagous geo-
hortobionts increases from 33% in the reference area
to 46% in the strongly degraded pasture, and the pro-
portion of mixophagous stratohortobionts also
increases from 20 to 27%. Because of their ability to
feed on mixed food, these ground beetles have greater
chance to survive than zoophagous species. The abun-
dance of the latter consistently decreases as the bio-

RUSSIAN JOURNAL OF ECOLOGY Vol. 47 No. 1 2016

 COMPOSITION AND ABUNDANCE OF SOIL MACROFAUNA
Table 4. Estimated abundance of Tenebrionidae in test areas
Species
virgin land
Blaps halophila F.-W. SD
Blaps lethifera F.-W. SD
Crypticus quisquilius L. SD
Gonocephalum granulatum pusillum F. D D
Opatrum sabulosum L. SD
Pedinus femoralis L. SD
Platyscelis hypolithа Pall. SD
Tentyria nomas Pall. D D
S, sporadic, abundance less than 1%; R, rare, abundance 1–3%; SD, subdominant, abundance 3–10%; D, dominant, abundance over 10%.
geocenoses become more xerophytic (from 26% in the
reference area to 13% in the strongly degraded pasture).
Thus, the highest species diversity of carabids was
recorded in the conditional virgin area, and the lowest
diversity, in the strongly degraded pastures. It is
believed that the combination of dominant species at
each studied locality is the most informative parame-
ter for estimating the degree of changes in species
composition. The area occupied by quasi-natural veg-
etation and weakly degraded areas were dominated by
ground beetle species such as the steppe litter–soil
zoophagous stratobiont Poecilus sericeus Fisch.,
meadow litter zoophagous stratobionts Calathus mela-
nocephalus L. and C. erratus C.Sahlb., and meadow–
field litter–soil zoopagous stratobiont Poecilus lepidus
Leske. The strongly degraded pasture was dominated
by the geohortobiont mixiphytophagous Harpalus
smaragdinus Duft. and H. calceatus Duft., mixophy-
tophagos stratohortobiont Ophonus stictus Steph., and
litter–crevice zoophagous stratobiont Cymindis angu-
laris Gyll. It can be concluded that the decreasing pro-
portion of litter species of the genus Poecilus could be
a sign of strongly depleted herbaceous layer in a pas-
ture area. The simultaneous shift of dominance toward
the geohortobiont group (Harpalus Lath. and Amara
Bon.) is indicative of significant changes in the prop-
erties of the upper soil layer, especially its density. Con-
siderable compaction and drying of this layer is also indi-
cated by an increase in the proportion of steppe species of
ground beetles (Calathus halensis Schall., Cymindis
angularis Gyll., and Ophonus stictus Steph.). The
growing abundance of these species is evidence for
aridization of the ecosystem.
Soil detritophages include not only earthworms,
but also darkling beetles (Coleoptera: Tenebrionidae),
which grind, eat, and decompose plant litter, thereby
contributing to its mineralization (Table 4). This
group in the conditional virgin land area was strongly
dominated by Tentyria nomas Pall., Gonocephalum
granulatum pusillum F., and Blaps halophila F.-W.
These species were subdominant in all areas, and their
RUSSIAN JOURNAL OF ECOLOGY Vol. 47 No. 1
71
Degree of pasture digression of vegetation
slight moderate strong
SD SD SD
SD SD SD
SD SD SD
D SD
SD SD SD
SD SD SD
SD SD SD
D D
numbers were the highest in conditional virgin land
(47 ind./100 trap–days) and the lowest in the strongly
degraded pasture (17 ind./100 trap–days), with their
relative abundance being 8.32 and 9.83%, respectively.
The increase in the latter parameter is explained by a
reduced proportion of other darkling beetle species in
the ecosystem. Blaps lethifera F.-W. was subdominant
in all areas except the moderately degraded pasture,
where it was dominant (10.04% of the total abun-
dance) due to the absence of Platyscelis hypolithа Pall.
and Blaps lethifera F.-W.
At the same time, the abundance of mesophilic
species, including Crypticus quisquilius L., consider-
ably decreased as a result of overgrazing (from 54 ind./
100 trap–days in the virgin land to 3 ind./100 trap–days
in the strongly degraded pasture).
Gonocephalum granulatum pusillum F. dominated in
the virgin land and weakly degraded pasture (12.61%),
but its rank dropped to subdominant in the moderately
degraded and strongly degraded pastures. Platyscelis
hypolithos Pall. was subdominant in the virgin land
(55 ind./100 trap–days, 9.73%) but proved to be
absent even in the moderately degraded pasture.
Tentyria nomas Pall. was dominant among darkling
beetles in all areas (in the virgin land, 197/100 trap–
days, 34.87%). Its abundance decreased under the effect
of pasture digression but remained high even in strongly
degraded pasture, reaching 106 ind./100 trap–days
(61.27%).
Strong soil compaction, the removal of vegetation,
and other consequences of overgrazing in most cases
lead to a decrease in the abundance of soil-living
invertebrates. Thus, in the series from conditional vir-
gin land to strongly degraded pasture, the abundance
of the meadow–steppe mesophilic loessobiont Crypti-
cus quisquilius L. decreased by 94.4%, and the abun-
dance of the forest–steppe mesophile Pedinus femora-
lis L. decreased by 95.5%. At the same time, the abun-
dance of thermoxerophilous species of the herbaceous
layer decreased by only 59.5%, and the abundance of
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72 RUSANOV, BULGAKOVA
the steppe xerophile Opatrum sabulosum L. decreased
by 75.5%.
Therefore, the irrational use of pasture ecosystem
leads, above all, to the loss of mesophilous compo-
nents of the beetle fauna (Crypticus quisquilius L., Ped-
inus femoralis L., and others), which give place to dry
steppe species such as Blaps halophila F.-W., Gono-
cephalum granulatum pusillum F., and Platyscelis hypo-
litha Pall.
Another important consequence of excessive graz-
ing is the compaction of the upper soil layer, which
hinders the movement of earthworms, while their food
supply decreases because of reduced amount of plant
litter. These phenomena account for the fact that the
abundance of earthworms during all studied seasons
(spring, summer, and autumn) was low: only one or
two individuals were recorded in some samples, while
in others the number of earthworms reached 13. Stan-
dard deviation varied between the samples from 1.7 to
9.1. These results are evidence for a patchy distribution
of earthworms over pasture areas. Since the dung of
livestock was regularly found in sites where their abun-
dance was the highest, the uneven distribution of
earthworms could be explained by the presence of this
easily accessible food and the microclimate it created.
Therefore, since earthworms are detritophagous,
livestock grazing plays a positive role by improving
their food supply, but overgrazing and its conse-
quences have a suppressing effect on them.
CONCLUSIONS
Transformation of genetic properties of cherno-
zems and change of plant formations caused consider-
able changes in the species composition of soil organ-
isms. Simultaneous impairment of several habitat fac-
tors provoked a rapid decrease in the abundance of
moisture-loving and least flexible species. Their place
in the pasture biocenosis was occupied by highly toler-
ant mixophytophagous species for which the estab-
lished environmental conditions are not limiting.
These changes in the composition of soil macrofauna
develop as the degree of pasture digression increases.
Therefore, the zoological method for indication of
pasture ecosystems in the steppe zone is quite suitable
for identifying and specifying the levels of their eco-
logical state. In addition, the results of this study allow
us to recommend that parameters of the species com-
position and abundance of soil invertebrates be used as
an indicator for monitoring lands of all categories.
It should be specially noted that the past few
decades have been a period of large-scale natural
regeneration of soils and vegetation in steppe land-
scapes conditioned by decreasing intensity of agricul-
tural land use (Rusanov, 2012; Babaeva and Osipova,
2014; etc.). Thus, the upper soil layer in pastures has
loosened after a period of relative dormancy. Plant
communities with signs of strong digression, which in
RUSSIAN JOURNAL OF ECOLOGY
the Cisural region were previously observed in an area
of more than 400 000 ha, can now be found only in the
vicinities of cities and villages. The area occupied by
moderately grazed plant communities has decreased,
while that under quasi-natural communities has
increased significantly.
The results of this study give ground for considering
the simultaneous restoration of the species composi-
tion and abundance of soil invertebrates, an important
component of the “soil living concentration” dis-
cussed by V.I. Vernadsky in the early 20th century.
REFERENCES
Agrokhimicheskie metody issledovaniya pochv (Agrochemical
Methods of Soil Studies) Sokolov, A.V, Ed., Moscow:
Nauka, 1975.
Babaeva, M.A. and Osipova, S.V., Soil as a factor of vegeta-
tion recovery in natural pastures of the Terek–Kuma Low-
land under conditions of intensive anthropogenic pressure,
Tr. Inst. Geol. Dagestan. Nauch. Tsentra Ross. Akad. Nauk,
2014, no. 63, pp. 35–38.
Gaevskaya, M.A., Changes in the properties of forest–
steppe and steppe chernozem soils under different anthro-
pogenic loads as a factor affecting the composition and
abundance of soil macrofauna, Vestn. Orenburg. Gos. Univ.,
2011, no. 12 (131), pp. 49–51.
Ghilarov, M.S., Zoologicheskii metod diagnostiki pochv
(Zoological Method of Soil Diagnosis), Moscow: Nauka,
1965.
Gorchakovskii, P.L. and Ryabinina, Z.N., Steppes of
southern Orenburg oblast (the Ural–Ilek interluve), in Ras-
titel’nye soobshchestva Urala i ikh antropogennaya degradat-
siya (Plant Communities of the Urals and Their Anthropo-
genic Degradation), Sverdlovsk, 1984, pp. 3–64.
Gur’ev, M.A., Physical soil degradation in pasture ecosys-
tems, in Fizika pochv i problemy ekologii: Sb. st. (Soil Physics
and Ecological Problems: Collected Papers), Pushchino:
ONTI PNTs RAN, 1992, pp. 23–24.
Kaurichev, I.S., Praktikum po pochvovedeniyu (A Practical
Course in Soil Science), Moscow: Kolos, 1980.
Krivolutskii, D.A., Pochvennaya fauna v ekologicheskom
kontrole (Soil Fauna in Ecological Control), Moscow:
Nauka, 1994.
Lavrenko, E.M., Karamysheva, Z.V., and Nikulina, R.I.,
Stepi Evrazii (The Steppes of Eurasia), Leningrad: Nauka,
1991.
Mirkin, B.M., Anthropogenic dynamics of vegetation, Itogi
Nauki Tekh., Ser.: Bot., Moscow: VINITI, 1984, no. 5,
pp. 139–232.
Mirkin, B.M. and Naumova, L.G., Populyarnyi ekologich-
eskii slovar' (Popular Ecological Dictionary), Ufa: Kitap,
1997.
Mordkovich, V.G., Zoologicheskaya diagnostika pochv leso-
stepnoi i stepnoi zon Sibiri (Zoological Soil Diagnosis in the
Forest–Steppe and Steppe Zones of Siberia), Novosibirsk:
Nauka, 1977.
Mullagulov, R.T., Suyundukov, Ya.T., and Yamalov, S.M.,
Effect of livestock grazing on the plant and soil components
of steppes in the Bashkir Transural region, Vestn. Orenburg.
Gos. Univ., 2009, no. 6, pp. 589–591.
Vol. 47 No. 1 2016
 COMPOSITION AND ABUNDANCE OF SOIL MACROFAUNA
Mustafaev, K.M., Adverse effect of livestock grazing on the
soil, Dokl. Akad. Nauk SSSR, 1960, no. 2, pp. 175–177.
Ramenskii, L.G., Problemy i metody izucheniya rastitel’nogo
pokrova (Problems and Methods of Studies on the Plant
Cover), Leningrad: Nauka, 1971.
Rusanov, A.M., The humus state of southern chernozems
under natural pastures, Pochvovedenie, 1993, no. 11,
pp. 25–30.
Rusanov, A.M., Influence of erosion on the humus state of
chernozems in the Ural region, Russ, J. Ecol., 1995, no. 2,
pp. 132–135.
Rusanov, A.M., Natural recovery of agrolandscapes in the
steppe and forest–steppe zones of Orenburg oblast, Stepnoi
Byull., 2012, no. 36, pp. 8–12.
Rusanov, A.M. and Gaevskaya, M.A., Changes in the soil–
plant–soil macrofauna community under anthropogenic
impact, Vestn. Orenburg. Gos. Univ., 2011, no. 12 (131),
pp. 129–132.
Striganova, B.R., Pitanie pochvennykh saprofagov (Nutri-
tion of Soil Saprophages), Moscow: Nauka, 1980.
RUSSIAN JOURNAL OF ECOLOGY Vol. 47 No. 1
73
Striganova, B.R., Systemic analysis of biocenotic relation-
ships in soil communities, in Chteniya pamyati M.S. Gil-
yarova (M.S. Ghilarov Memorial Lectures), Moscow: KMK,
2006, pp. 16–38.
Teslya, A.V., Ecological conditions of natural vegetation
recovery in overgrazed pastures in the forest–steppe and
steppe zones of the Cisural region, Extended Abstract of
Cand. Sci. (Biol.) Dissertation, Orenburg, 2009.
Vadyunina, A.F. and Korchagina, Z.A., Metody issledo-
vaniya fizicheskikh svoistv pochv (Methods for Analyzing
Physical Properties of Soils), Moscow: Agropromizdat, 1986.
Vernadsky, V.I., Problemy biogeokhimii. Znachenie bio-
geokhimii dlya poznaniya biosfery (Problems in Biogeo-
chemistry: The Significance of Biogeochemistry for the
Cognition of the Biosphere), Moscow: Akad. Nauk SSSR,
1935.
Vladychenskii, A.O., Ul’yanova, T.Yu., Balandin, S.A., and
Kozlov, I.N., Impact of grazing on the soils of juniper for-
ests in the southwestern Tien Shan, Pochvovedenie, 1994,
no. 7, pp. 45–52.
Translated by P. Petrov
2016