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Density and distribution of the greater mole rat (Spalax microphthalmus) on the northern border of the area in a region of European Russia

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There are few subterranean mammals in Russia. The aim of the paper is to study the distribution and abundance of greater mole rat in the local population. The characteristic of the local population of the greater mole rat (Spalax microphthalmus) on the northern border of the range with reference to geographical coordinates is given. Our research has been conducted in Republic of Mordovia (European Russia). The field studies were carried out in 2007-2019. The population density in Mordovia is from 2 to 6 individuals/ ha. The number of greater mole rat is estimated at 190 individuals. There are 27 to 200 mounds per one hectare of registered area. The distance of the greater mole rat habitat in Mordovia is from 40 to 372 km from other populations of neighboring regions. It is revealed that animals in the region are confined to pastures and dacha areas. The population density of subterranean rodent populations, such as mole rats, is in equilibrium with the capacity of the environment in which they live. The results on the population density and the number of greater mole rats on the northern border of the species range indicate the oppression of the local population.
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Ecological Questions 32 (2021) 2: 71-80 http://dx.doi.org/10.12775/EQ.2021.015
Density and distribution of the greater mole rat
(Spalax microphthalmus) on the northern border of the area
in a region of European Russia
Alexey V. Andreychev1*, Alexandr S. Lapshin2
1Department of Zoology, National Research Mordovia State University, Bolshevistskaya Street 68,
430005, Saransk, Russia
2Biological Museum, National Research Mordovia State University, Ul’yanova Street 26 A,
430019, Saransk, Russia
*corresponding author e-mail: teriomordovia@bk.ru
Received: 20 April 2020 / Accepted: 27 October 2020
Abstract. There are few subterranean mammals in Russia. The aim of the paper is to study the distribution and abundance of greater
mole rat in the local population. The characteristic of the local population of the greater mole rat (Spalax microphthalmus) on the
northern border of the range with reference to geographical coordinates is given. Our research has been conducted in Republic of
Mordovia (European Russia). The field studies were carried out in 2007-2019. The population density in Mordovia is from 2 to 6 in-
dividuals/ha. The number of greater mole rat is estimated at 190 individuals. There are 27 to 200 mounds per one hectare of registered
area. The distance of the greater mole rat habitat in Mordovia is from 40 to 372 km from other populations of neighboring regions.
It is revealed that animals in the region are confined to pastures and dacha areas. The population density of subterranean rodent popu-
lations, such as mole rats, is in equilibrium with the capacity of the environment in which they live. The results on the population
density and the number of greater mole rats on the northern border of the species range indicate the oppression of the local population.
Key words: subterranean mammals, mound, relative abundance, rodent, Republic of Mordovia.
casus (Topachevskii, 1969; Flint et al., 1970; Puzachenko,
1993; Savchenko & Ronkin, 2006). The Northern bound-
ary runs from Kiev to the southern part of the Chernigov
oblast, the Eastern part of the Kursk and Orel oblasts, in the
southern outskirts of Tula, Ryazan, Tambov, Penza oblasts,
Republic of Mordovia, Nizhny Novgorod oblast, the Chu-
vash Republic, Ulyanovsk oblast. The extreme Northern
point of detection does not rise above the 56th parallel
of the northern latitude (flood plain of the Bolshoi Civil,
Chuvash Republic). The extreme North-Eastern point of
detection is located in the Samara region (Samara oblast).
The southern border of the area runs along the coast of
the sea of Azov to the East in the South-Western part of
the Krasnodar territory and rises in the Stavropol territo-
1. Introduction
Herbivores, especially subterranean mammals, can influ-
ence per plants, soil structure and chemistry by moving
soil (Williams & Cameron, 1984; Andersen, 1987; Inouye
et al., 1987; Benedix, 1993). There are few subterranean
mammals in Russia. However, their role is significant in
biotopes. They are almost unknown in many regions.
The habitat of the greater mole rat Spalax microph-
thalmus Guldenstaedt 1770 (Rodentia, Spalacidae) it cov-
ers areas with fertile soil and abundant vegetation. It lives
in the steppe and forest-steppe of South-Eastern Europe
between the Dnieper and the Volga, in the Western Cau-
72 Alexey V. Andreychev, Alexandr S. Lapshin
ry to the North-East. The border then goes to the Volga
North of Volgograd. South-Eastern borders have not been
established yet (Puzachenko, 2011).
A great contribution was made to the study of the great-
er mole rat in Russian theriology by S. L. Ovchinnikova
and A. Yu. Puzachenko. More attention to the ecology and
biology of the species was given in Kursk, Voronezh, and
Belgorod oblasts. For other regions, there is mainly frag-
mentary information, which, as a rule, is limited to a de-
scription of the distribution and status. The mole-rat has
a rare species status in Tambov, Ryazan, Samara, and Uly-
anovsk oblasts, as well as the republics of Mordovia and
Chuvashia. In the context of the foregoing, the research in
other regions, in particular, where relict local populations
are preserved, is of significance. It is no coincidence that
A.Yu. Puzachenko proposes to consider isolated popula-
tions as vulnerable or potentially vulnerable. The fact is
that, at the end of the Pleistocene, mole-rats inhabited arid
steppes, and with the onset of forests in the Holocene, the
northern line of its range, as well as many other steppe spe-
cies, shifted markedly southward and only a few popula-
tions remained in the former distribution boundaries. In the
Volga region, mole-rat isolates were preserved in the refu-
gia of steppe vegetation in Mordovia, Chuvashia, Nizhny
Novgorod, and Samara oblasts.
Many researchers rightly point out that direct obser-
vations of the biology and ecology of mole-rat are diffi-
cult. This is due to the large length of individual burrows
(up to hundreds of meters), seasonal changes in the con-
figuration and direction of burrows on individual sites
(Ovchinnikova, 1971; Rado & Terkel, 1989; Puzachenko,
1993; Vlasov & Puzachenko, 1993).
The aim of the paper is to study the distribution and
abundance of greater mole rat in the local population at
the border of the area. These tasks are relevant, since in
the literature these aspects are not adequately addressed.
In addition, in local animal populations under suboptimal
conditions, differences in population density can be ob-
served compared with populations of the main range of
the species.
2. Study area
The studies were carried out in Ruzaevsky district of the
Republic of Mordovia and Saransk urban district (geo-
graphical coordinates 54º05’–54º09’N and 44º59’–45º06’
E) (Fig. 1) in 2007–2019 and covered the entire habitat of
mole rats in the region. The climate of the region is mod-
erately continental with pronounced seasons throughout
the year. The influx of direct solar radiation varies from
5.0 in December to 58.6 kJ/cm2 in June. Total radiation
throughout the year is 363.8 kJ/cm2; the radiation balance
is 92.1 kJ/cm2. The average annual air temperature var-
ies from 3.5 to 4.0 °C. The average temperature of the
coldest month (January) is in the range of –11.5...–12.3
°C. Temperature drops down to –47 °C occur. The average
temperature of the warmest month, i.e. July, is in the range
of +18.9...+19.8 °C. Extreme temperatures in the summer
reach 37 °C. The average annual precipitation in the ter-
ritory is 480 mm. Over the course of observation lasting
many years, periods of more and less humidification were
noted, ranging between the minimum and maximum values
of 120-180 mm. Distribution of precipitation across the
territory is not very diverse. The average long-term value
of evaporation is calculated to be in the range of 390-460
mm (Yamashkin, 1998).
3. Material and Methods
The degree of population of the territory on individual
areas was determined by calculating the amount of land
emissions on 1-hectare sites (Dukel’skaya, 1932; Guly-
aevskaya, 1954; Pavlov et al., 1963; Puzachenko & Vlasov,
1993). We carried out selective catches from burrows to
determine the number of animals in burrow systems with
subsequent release (Fig. 1). Catching was carried out in
burrows until the noises of the passage of animals were
recorded on dictaphones in the burrow. Studies were car-
ried out according to a new methodology that we devel-
oped for studying of subterranean mammals (Andreychev,
2018; 2019; Andreychev et al., 2020). They also noted the
absence of small animals after catching them in burrows
where the ground was not covered with drafts.
Initially, the boundaries of individual habitats of mole
rats were revealed, then the directions of the foraging pas-
sages were determined. The activity of animals may be
surveyed with the use of voice recorders in different parts
of colonies. The location of recorders was preliminarily
determined on the basis of a map with consideration of the
boundaries of animal colonies. A GPS-navigator was used
for quick detection of recorders in the field.
When conducting scientific research, we used АIМР
1.75 (2007) and AUDACITY2.1.1 (2015) programs (An-
dreychev, 2018). During the research period, 200 sound
recordings were obtained and processed, with a total length
of about 16000 hours. We carried out 496 measurements of
distances between mounds mole rats.
4. Results
It is revealed that the mole rat lives only in the central
part of the region within the Saransk urban district and
Ruzaevsky district. Natural and anthropogenic factors in
Mordovia have long been limiting the further territorial
settlement of this species. The settlement of mole rats to
[73]
4
Figure 1. Geographical position of the Republic of Mordovia in Russia and distance of the local
population of Spalax microphthalmus from other populations of neighboring regions.
Note: Remoteness of population with the distance indicated (km)
Points of closest populations
Figure 1. Geographical position of the Republic of Mordovia in Russia and distance of the local population of Spalax microphthalmus
from other populations of neighboring regions.
Note:– Remoteness of population with the distance indicated (km)
Points of closest populations
74 Alexey V. Andreychev, Alexandr S. Lapshin
the west and south was limited by the Levzha River and
its left tributaries, to the north, by the forest of Saransk,
and to the east, by a road and the Insar River. A distinction
was made using the OziExplorer program for 13 mole rat
habitats (Table 1). The territory of detection of mole rat is
currently more than 100 km2. The area was previously 26.8
km2 in 2010. However, the actual inhabited area is 568.3
ha. In 2010, it was 297.7 ha (11.1% of the total area). In
the study area, the species does not populate fields sown
with grain crops, tall grasses, and forest belts. It prefers
meadows, pastures, summer cottages, and hayfields. It also
settles near ponds and marshy areas.
However, the population of the mole rat is experienc-
ing significant anthropogenic pressure. The mole rat was
recorded on pastures, summer cottages, hayfields, and ar-
able lands (Fig. 2). The animals prefer pastures and sum-
mer cottages, where from 27 to 200 mounds occur on 1
ha (Table 2).
Table 1. Number and area of the greater mole rat’s habitat.
Number habitat Locality Habitat area, ha Number, individuals
1 Levzhenskii 32.4 25
2 Dobrovol’nyi 16.8 4
3Klyucharevo (South-East) 20.6 8
4Klyucharevo (North) 183.8 15
5Klyucharevo (West) 3.4 6
6 Popovka 12.5 60
7 Rybnyi 28.2 11
8Klyucharevskie Vyselki 110.6 24
9 Levzha 77.3 12
10 Yalga 51.1 18
11 Kulikovka 10.7 3
12 Monastyrskoe 15.8 2
13 Novaya Nechayevka 5.1 2
7
51%
22%
16%
11%
1
2
3
4
Figure 2. Occurrence of greater mole rat in various anthropogenic habitats in Mordovia.
Note: 1 pastures; 2 dacha areas, 3 haymaking, 4 arable land
Table 2. Characteristics of trial registration areas (1 ha), habitats of S. microphthalmus.
Site number
Locality
Length of
burrows, m
Burrow
direction
(N,S,W,E)
Density of
mounds per 1 ha
I
Klyucharevskie
Vyselki
160
N-S
200
II
Klucharevo
134
N-S
56
III
Popovka
125
W-E
105
IV
Levzhenskii
110
W-E
53
V
Levzha
71
N-S
25
VI
Yalga
281
N-S
50
VII
Kulikovka
95
N-S
30
Figure 2. Occurrence of greater mole rat in various anthropogenic habitats in Mordovia.
Note: 1 – pastures; 2 – dacha areas, 3 – haymaking, 4 – arable land
75
Density and distribution of the greater mole rat (Spalax microphthalmus)
The number of the greater mole rat in the total area of
habitat in Mordovia was 190 individuals. Moreover, the
distribution of land plots is uneven (Table 1). The popula-
tion density varied from 2 individuals / ha (the villages of
Monastyrskoe, the of Kulikovka, the of Novaya Nechayev-
ka) to 6 individuals / ha (the villages of Klyucharevskie
Vyselki, of the Klyucharevo, of the Popovka).
The primacy of animals changes to different anthropo-
genic habitats. Mole rats prefer to settle in the district of
summer cottages near the village of Klucharevo. Mole rat
inhabits the villages Rybnyi and Levzhenskii in the pas-
tures around ponds. It should be noted that pastures in the
Kursk region are considered a suboptimal biotope for the
blind (Puzachenko & Vlasov, 2012). This once again high-
lights the differences in the animal’s biotopes in different
regions.
The remoteness of the local mole rat population from
other populations of neighboring regions differs. From the
populations of the southern regions, namely, Penza and
Ulyanovsk oblasts, which are part of the continuous range
of the species, the habitat of mole rats from Mordovia is
separated by a distance of about 40 and 125 km, respec-
tively. In Penza oblast, the closest biotopes where mole-
rats live are Issinskii and Luninskii districts (Il’in et al.,
2006). In Ulyanovsk oblast, the nearest habitat with respect
to the population from Mordovia is located in the Bazarno-
syzganskii district in the vicinity of the Yasachnyi Syzgan
village (Abrakhina et al., 1993). As for identical isolated
populations from the Chuvashia and Ryazan and Nizhny
Novgorod oblasts, the distances to them from the extreme
habitats of the mole rat in Mordovia are 120, 372, and 131
km, respectively. In Chuvashia, only four local populations
were noted, the nearest of which is in the Alatyr district
(Dimitriev et al., 2010). In Nizhny Novgorod oblast, there
are only two isolates in the Mezhp’yan’e (Sergachskii and
Krasnooktyabr’skii districts) and on the left bank of the
southern branch of the P’yana (the border of the Gaginskii
and Shatkovskii districts) (Bakka, 2014). In Ryazan oblast,
the habitat is known only on the slopes of the Panika River
valley to the southeast of the Chernavskaya oak grove in
Miloslavskii district (Didorchuk & Kotyukov, 2012).
5. Discussion
We compared the current range boundaries of the mole
rat with data on its distribution in Mordovia in the past
(Astradamov et al., 1976; Andreychev et al., 2010; An-
dreychev, 2020). It should be noted that it significantly ex-
panded the areas of their habitat (Fig. 3). In recent years the
mole rat has managed to expand their habitat to the East to
the village Kulikovka, West to the village Klyucharevskie
Vyselki, South to the village Levzha. No settlement was
registered in the North. In Mordovia, the distance between
Table 2. Characteristics of trial registration areas (1 ha), habitats of S. microphthalmus.
Site number Geographical
coordinates Locality Length of
burrows, m
Burrow
direction
(N,S,W,E)
Density of mounds
per 1 ha
I54.1507 N,
44.9473 E
Klyucharevskie
Vyselki
160 N-S 200
II 54.1340 N,
45.0160 E
Klucharevo 134 N-S 56
III 54.1200 N,
45.0289 E
Popovka 125 W-E 105
IV 54.0998 N,
45.0964 E
Levzhenskii 110 W-E 53
V54.0928 N,
45.0192 E
Levzha 71 N-S 25
VI 54.1329 N,
45.1316 E
Yalga 281 N-S 50
VII 54.1170 N,
45.1770 E
Kulikovka 95 N-S 30
VIII 54.0779 N,
45.1644 E
Monastyrskoe 181 N-S 60
76 Alexey V. Andreychev, Alexandr S. Lapshin
the edge detection points of mole rats from north to the
south is currently 15 km, and from the west to the east, 10
km. In Mordovia, the distance between the edge detection
points of mole rats from the north to the south was 10 km
and from the west to the east, 8.8 km (Andreychev et al.,
2010).
According to the results obtained in Mordovia and in
the territories where the population density of S. microph-
thalmus is higher (Kursk oblast) (Puzachenko & Vlasov,
1993), animals from our region have low (10-50 mounds
per 1 ha) and medium (100-200 mounds per 1 ha) intensity
of burrowing activity. The smallest occurrence of mounds
10
Figure 3. Geographic distribution of greater mole rat in the Republic of Mordovia.
Note: A solid black line marks the former border of the greater mole rat habitat
(Astradamov et al., 1976), and a dotted black line marks the current border of the
greater mole rat habitat (our data, 2019). Black squares indicate the experimental
areas for the registration of the greater mole rat (1 ha each). IVII-site location
numbers (see symbols in Table 1)
Figure 3. Geographic distribution of greater mole rat in the Republic of Mordovia. Note: A solid black line marks the former border
of the greater mole rat habitat (Astradamov et al., 1976), and a dotted black line marks the current border of the greater
mole rat habitat (our data, 2019). Black squares indicate the experimental areas for the registration of the greater mole rat
(1 ha each). I–VII-site location numbers (see symbols in Table 1)
77
Density and distribution of the greater mole rat (Spalax microphthalmus)
mole rats was noted on agricultural arable lands and veg-
etable gardens; the first type of plots is rejected by the
mounds itself, and from vegetable gardens it is actively
expelled by humans. Under optimal habitat conditions, the
mole rats can form up to 380 mounds per 1 ha, for exam-
ple, in the steppe of the Lugansk oblast (Gulyaevskaya,
1954).
For comparison, the population density of S. microph-
thalmus was 2.6-4.9 individuals / ha in the Volga forest-
steppe reserve in the Penza oblast in 1998-2004 (Il’yin
et al., 2006). There, the blind man lives in a single undi-
vided population. The population density for the Voronezh
oblast varied from 3 to 20 individuals/ha (Ovchinnikova,
1971). The population density in the virgin steppe of the
Lugansk oblast was from 1 to 11 individuals / ha (on aver-
age 3.7-3.8) (Gulyaevskaya, 1954).
The results of studying the burrowing activity of the
mole rat are poorly displayed in many regions. It is advis-
able to provide the available data for some regions (Ta-
ble 3). The data obtained for the territory of Mordovia
are consistent with the results of other researchers (Pu-
zachenko & Vlasov, 1993; Didorchuk & Kotyukov, 2012).
However, the average depth of the upper edge of the stern
passage is less in Mordovia (13.2±1.49 cm) than in the
Kursk and Belgorod oblasts (22.1±1.17 cm). The obtained
average value of distances between adjacent mounds for
Mordovia. It is 3.5 m. The average length of the burrow
system is 70.7 m (n=30) in Mordovia.
It was found that animals have different indicators
of feeding passages in different types of anthropogenic
changed habitats. They are slightly different from the same
indicators in natural areas (meadows). It is noted that the
most common land emissions occur in pastures and dacha
plots. Sparseness is shown for haymaking and arable land.
The lowest average depth of the upper edge of the stern
passage is marked for suburban areas. The greatest weight
of the thrown out earth from the passages is registered for
haymaking. However, the regularity in the direction of the
feeding course is shown from the North-East to the South-
West for almost all types of anthropogenic habitats (more
than 70%) of the mole rat in Mordovia.
Cattle graze annually in the number of more than
150 individuals in many areas of the mole rat is habitat.
In particular, this is typical for the village of Levzhenskii.
We observed numerous failures of feeding passages in pas-
tures. They were formed as a result of trampling the soil
by cattle. S. microphthalmus in places of failures of feed
passages conduct renewable activity. Gradually, new earth
emissions appear. Researchers have found that the appear-
ance of new mounds is a reliable marker of employment
of the optimal territory (Zhukov et al., 2011).
Many researchers note that the mole rat can often be
found in forest belts (Topachevskii, 1969; Ovchinnikova,
1971). The mole rat avoids these habitats in Mordovia.
The nearest mounds were found at a distance of 40 meters.
It was on a pasture near the village of Levzhenskii and 60
meters away on a pasture near the village of Klyucharevo.
It is very interesting to compare the features of burrow-
ing activity in different types of mole rat. For example, the
Riesen-Blindmaus Spalax giganteus in the Terek-Kumskoe
interfluve in the Caucasus. The number of mounds per
1 ha varied from 32 to 456, (on average 181) (Pavlov et al.,
1963). The population density of the S. giganteus reached
0.5-2 individuals / 1 ha, in the area of the Talovka River
(Dagestan) in August 1991 (Puzachenko, 1999). At the
same time, the average distance between the mounds was
about 5 m.
According to A. Yu. Puzachenko, the same burrow sys-
tem is inhabited by a sexually Mature female and male
greater mole rat. Besides them, fingerlings can be caught.
In this regard, the researcher introduced a classification
of burrow systems by the size and composition of their
inhabitants. He identified three groups of settlements of
the greater mole rat. The first group consists of complex
burrow systems extending for hundreds of meters, where
adult males, females and youngsters live together. The sec-
Table 3. Some characteristics of burrowing activity of S. microphthalmus in different regions
Region
Characteristic
Source
Average distance
between neighboring
mounds, m
Average depth
of burrow from the
soil surface, cm
Average mounds soil
weight, kg
Kursk and Belgorod
oblasts
1.9±0.07
(n=491)
22.1±1.17
(n=285)
10.6±1.63
(n=25)
Puzachenko, Vlasov,
1993
Ryazan oblast 1.4±0.2
(n=14)
Didorchuk, Kotyukov,
2012
Mordovia 3.5±0.12
(n=496)
13.2±1.49
(n=280)
10.3±0.44
(n=280)
Own data, 2019
78 Alexey V. Andreychev, Alexandr S. Lapshin
ond group includes systems that are less extensive (tens
or hundreds of meters) and contain only one individual.
The third group is formed by the norms of fingerlings that
have separated along the periphery of the family’s habitat.
The length of such burrows is insignificant (tens of me-
ters) (Puzachenko, 1993). According to this classification,
the first two groups of settlements are found in Mordo-
via. They are inhabited by several individuals (settlements
near the village of Levzhenskii, the village of Klyucharevo,
Popovka, Klyucharevskie Vyselki) and only one individual
(Dobrovol’nyi, Novaya Nechayevka, Monastyrskoe, Ku-
likovka).
The population density of the Palestine mole rat Spal-
ax ehrenbergi in Israel ranged from 91 to 177 individuals
per 1 km2 (1 to 1.8 individuals / ha) (Nevo et al., 1982).
According to other data, the population density of Spalax
golani varied in Israel from 2 to 10 individuals / ha (Lovy
et al., 2015). However, the population density of different
researchers is within the range of variation for this spe-
cies from 0.1 to 20 individuals/ha (Savic & Nevo, 1990).
The population density of other subterranean mammals is
4.6-5.2 individuals / ha (Sumbera et al., 2003, 2008). These
figures in terms of population density do not differ signifi-
cantly from S. microphthalmus.
The area and length of the burrows of the greater mole
rat are similar to other subterranean mammals. The esti-
mated density of mound clusters was 26 ± 21 clusters per
km2. The average and estimated cluster sizes were four
mounds per cluster. Estimated density of 94 ± 79 mounds
per km2 (Quinn et al., 2010).
The density Pocket gophers (Geomys breviceps) is 1.35
gophers per acre in Texas (Davis et al., 1938). In Texas,
densities Geomys bursarius ranged from 1.3, 16.0, and
18.7 individuals / ha in pastureland, riparian sandbar, and
hayfield, respectively (Broussard, 1996). Average density
estimates per hectare were 20.4 (range 4.0-60.4) in Ar-
kansas (Connior et al., 2010), 7.4-9.9 in Indiana (Mohr
& Mohr, 1936), 12.9 in Wisconsin (Zinnel, 1992), and 5.4-
22.0 in Minnesota (Adams, 1966). Smallwood and Morri-
son (1999) determined that generally much of the variation
can be explained by the size of the study area (Connior,
2011). Average home range sizes for adult female and adult
male of the Ozark pocket gopher (G. bursarius ozarkensis)
in Arkansas over 2 y of 287.1 m2 and 291.8 m2 (Connior
& Risch, 2010).
The habitat of the zokor is larger than that of the mole
rat. The home range of a male zokor exceeds 1500 m2 and
is larger than a female range of less than 500 m2 (Zhou
& Dou, 1990). The distribution and population density of
plateau zokors are limited by elevation, vegetation, precipi-
tation, and anthropogenic disturbance (Zhang et al., 1999).
It was estimated that they occupied a range of approxi-
mately 3.8×106 ha at an average density of 15 individuals
/ ha, but ranging from five to more than 70 per hectare in
Qinghai province in the 1980s (Wang & Fan, 1987; Zhang
et al., 1999; Zhang et al., 2003; Zhang, 2007). This popula-
tion density of the zokor is several times higher than that
of the greater mole rat.
6. Conclusions
Thus, it can be concluded that the population density of
subterranean rodent populations, such as mole rats, is in
equilibrium with the capacity of the environment in which
they live. The population density of the greater mole rat in
Mordovia varied from 2 to 6 individuals / ha. The number
in the total area of habitat was 190 individuals. The results
on the population density and the number of greater mole
rats on the northern border of the species range indicate the
oppression of the local population. In most subterranean
mammals species a population’s growth rate is a decreas-
ing function of density. This explains the relative stability
of animal populations, which never continue to increase
at rates their fertility would allow, and rarely decrease to
extinction (Tanner, 1966).
Acknowledgement
We are grateful to G. F. Salmov and R. M. Kulahmetov for
support in carrying out of field studies. We are grateful to
two anonymous reviewers for their constructive comments
and feedback on an earlier version of this paper.
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