ISSN 1995-4255, Contemporary Problems of Ecology, 2008, Vol. 1, No. 2, pp. 214–220. © Pleiades Publishing, Ltd., 2008.
Original Russian Text © Yu.S. Ravkin, V.A. Yudkin, S.M. Tsybulin, V.N. Kuranova, O.B. Borisovich, N.A. Bulakhova, S.V. Patrakov, R.R. Shamgunova, 2007,
published in Sibirskii Ekologicheskii Zhurnal, 2007, Vol. 14, No. 4, pp. 557–565.
Spatial-Typological Structure and Mapping of Reptile
Population of West Siberia
Yu. S. Ravkina, V. A. Yudkina, S. M. Tsybulina, V. N. Kuranovab, O. B. Borisovicha,
N. A. Bulakhovab, S. V. Patrakovb, and R. R. Shamgunovac
a
Institute for Systematics and Ecology of Animals, Siberian Branch of the Russian Academy of Sciences,
ul. Frunze 11, Novosibirsk, 630091 Russia
b
Tomsk State University, ul. Lenina 36, Tomsk, 634050 Russia
c
Surgut State University, ul. Energetikov 14, Surgut, 628400 Russia
Abstract—West Siberian reptiles were studied on the basis of the materials collected in forest, forest-steppe,
and steppe zones of the Ob’ region during 12 years. Their abundance in certain biotopes has been determined.
The relationship between the heterogeneity of reptile community and basic structure-forming environmental
factors has been estimated, and a map of population of this faunal class was compiled.
DOI: 10.1134/S1995425508020070
ternate with corn fields (3) and is absent from low
floodplains and other noninundated areas. In subtaiga
forests the sand lizard is typical of low vegetation at
highmoor bogs (9), birch-aspen and pine forests (6), as
well as of inundated plains of large rivers (2). Occasionally it can be found in small forests which alternate with
fields (0.5); was not found in lowland bogs and noninundated meadows. In northern forest-steppe it is common in birch-aspen and pine forests (5), and in small
forests alternating with corn fields (2); it is rare in inundated plains of large rivers (0.4). In southern forest-steppe this lizard is widely distributed in inundated
plains of large rivers (9), in birch-aspen and pine forests
(6 and 3), as well as in afforested lowland bogs (2). In
open lowland bogs, field woodland, and meadows the
lizard is rare (0.3–0.5) and is not met in steppe meadows and meadow steppe. In the steppe zone it is commonly found in fields and open lowland bogs (1),
rarely — in meadows (0.2).
Thus, the sand lizard has the widest distribution
throughout the subzone of subtaiga forests decreasing
in amount northward and southward. High floodplains
and small-leaved noninundated forests are the preferable habitats of this lizard.
The viviparous lizard (common lizard) in West Siberia is distributed from south through southern bushy
tundras [9]. The route estimations of northern taiga
have covered only the Sibirskie Uvaly though these lizards have been repeatedly trapped into cylinders for
mammals in terraces above inundated plains of the Ob’
near Polnovat Village (Berezovo district of Tjumen’
oblast). The common lizard is the most widespread in
full-grown pine riams of degenerative bogs (0.5 individual per hectare). It is almost absent from bird cherry
and willow brushwoods of inundated plains of small
rivers (0.09), subshrub-sphagnum and grass-moss hummock-ridge aapa bogs and pine forests (0.06 and 0.04).
Reptiles are usually counted within limited territories, that is why the data on their population are utterly
insufficient and do not provide a full view of their distribution over regions. In Russia, reptiles of Altai, as
well as of forest-steppe and taiga plain landscapes of
the Upper and Middle Ob’ region, are better studied in
this respect [1–6]. This work aims at studying the distribution of reptiles within natural boundaries and landscapes of three natural zones of the Ob’ region, clearing
up the territorial organization of this class of vertebrates, and mapping their population.
MATERIAL AND METHODS
The inventories of reptiles on one-meter-wide transects were carried out in May–July of 1967, 1976,
1977, 1985, 1991, 1992, 1999–2001, 2003–2005. Most
of the habitats were examined just once. The route of
each study was usually about 5 km long. In all, 359
variants of population were studied. The total route extent reaches approximately 1200 km. All the data were
converted per one hectare and averaged within contours
on the map “Flora of the West Siberian Plain” (scale 1 :
1 500 000, 1976) without regard to route area and extent. The species are named and ordered as in the
taxonomic catalogue of amphibians and reptiles of
Russia [7].
RESULTS AND DISCUSSION
Distribution of Species
The sand lizard (Lacerta agilis) does not occur in
northern taiga. In middle taiga it is found along the railway to Surgut [8]; in southern taiga it is abundant (24
individuals per hectare) in inundated forests of high
ecological level, i.e. forests rarely flooded during high
water. It is a common species in small forests which al214
SPATIAL-TYPOLOGICAL STRUCTURE AND MAPPING
In dark coniferous, mixed coniferous-broad-leaved and
small-leaved forests, as well as in logged areas the
lizard was not found.
In middle taiga the lizard is common in low vegetation riams of raised bogs and birch-aspen forests (5 and
3), rare in dark coniferous-small-leaved and pine forests (0.8 and 0.2), not found in inundated plains of large
rivers. In southern taiga the common lizard is found in
all the habitats. The widest distribution falls on dark coniferous-small-leaved forests and small forests which
alternate with fields (14–15), as well as on small-leaved
forests (11). This lizard is common in pine forests, in
forests and meadows of inundated plains of large rivers
(3–4), in dark coniferous forests and open lowland bogs
(2). It is rare in vast raised bogs (0.8).
In subtaiga forests the widest distribution of the
common lizard falls on low vegetation riams of raised
bogs and pine forests (13–14). It is common in afforested and open lowland bogs (7 and 4), as well as in
birch-aspen forests and small forests which alternate
with corn fields (2). In inundated plains of large rivers
the lizard is rare (0.4).
In northern forest-steppe the common lizard is numerous in birch-aspen forests (13), common in pine and
birch-pine forests (2), and rare in small forests which alternate with fields (0.3). It was not found in inundated
plains of large rivers.
In southern forest-steppe the widest distribution of
the lizard falls on birch-aspen forests and afforested
lowland bogs (4). Moreover the species is common in
pine forests (2) and in noninundated meadows alternating with shrubs (1), and rare in open lowland bogs (0.4).
In steppe zone the common lizard was not found.
Thus, the common lizard is widespread in southern
taiga decreasing in amount northward and southward
from the subzone. In northern and middle taiga most of
the species inhabit suppressed pine forests and bogs
(riams), in southern — dark coniferous-small-leaved
and small-leaved forests, in subtaiga forests — low
vegetation pine riams and pine forests, in foreststeppe — small-leaved forests.
The grass snake (Natrix natrix) in southern taiga
was found only in inundated plain of the Ob’ (1 individual per hectare) and in open lowland bogs (0.01). In
subtaiga it occurs in noninundated meadows (0.001)
and, not inventoried, has been noticed in pine forests
and settlements. In northern forest-steppe the species is
rare in pineries, birch-aspen forests, and small woods
which alternate with fields (0.4–0.7). In southern forest-steppe it is common for pineries (2), rare in inundated plains of large rivers and birch-aspen forests (0.7
and 0.4), and very rare in small forests which alternate
with fields (0.07). In lowland bogs, meadows, meadow
steppes and vast open fields. In the steppe zone it was
not registered during the estimation work.
Thus, the the grass snake is distributed from southern taiga to southern forest-steppe; pinery is a prefera-
215
ble habitat. Most of the population falls on southern forest-steppe.
The Pallas’s coluber (Elaphe dione), as inferred
from the literature, occurs in forest-steppe and steppe in
the south of West Siberia, though we found no supporting evidence.
The mamushi (Agkistrodon halys) is registered in
the forest-steppe zone on rocks of steep banks of the
Berd’ middle reaches. The rocky south-facing slopes
with mountain-steppe flora is the optimal habitat,
where mamushi local population density comprises no
less than 10–15 individuals per hectare [10].
The adder’s (Vipera berus) range in West Siberia extends from south up to northern taiga [9].
In northern taiga the widest distribution of the adder
was detected in full-grown pine riams of degenerating
bogs (0.5 individual per hectare). It is very rare in
subshrub-sphagnous and grass-moss hummock-ridge
aapa-bogs and bird-cherry and willow brushwoods of
inundated plains of small rivers (0.06 and 0.04), as well
as in pine forests (0.02). The species is almost absent
(0.003) from light coniferous-fir-cedar riparian forests,
and was not found in coniferous-broad-leaved and
small-leaved forests and in logged areas.
In middle taiga the adder is found in low vegetation
riams of raised bogs (0.6) and small-leaved forests
(0.3). The species was documented neither in coniferous and coniferous-broad-leaved forests nor in floodplains of large rivers. In southern taiga the abundance
of adder is typical of willow-birch forests along small
rivers and raised bogs (3–4). For afforested lowland
bogs and fir-cedar damp taiga the figures are
1.5–2 times lower (2). Rarely the adder is found in
small-leaved forests and coppices alternating with
fields, in coniferous-broad-leaved semi-swamped forests (0.3–0.5), open lowland bogs and inundated willow shrubs combined with meadows (0.2). It does not
occur in upland forests (dark coniferous, dark coniferous-small-leaved and basic), as well as in inundated
water meadows which are frequently flooded for long
periods. In subtaiga forests it is common in afforested
lowland and raised bogs, as well as in small forests alternating with fields (1–2). In birch-pine, pine and
small-leaved forests the species is rare (0.7 and 0.1). In
meadows and in open lowland bogs it does not occur.
In northern forest-steppe the adder is common in
birch-aspen forests (1) and almost absent from fields
which alternate with small forests (0.02). In pine forests
and floodplains of large rivers of this subzone the species was not found. In southern forest-steppe it is rare in
small-leaved forests, floodplains, afforested lowland
bogs (0.5), and noninundated dump meadows (0.1). It is
almost absent from fields which alternate with shrubs
(0.01) and not registered in open lowland bogs, in
steppe meadows, and meadow steppes. In the steppe
zone the adder does not occur, either.
Thus, southern taiga hosts the greatest amount of the
adder, which reduces northward and southward. In the
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RAVKIN et al.
forest zone the species generally prefers suppressed
uliginous pine forests, and small-leaved forests in forest-steppe.
A total of six species of reptiles occur in the
near-Ob’ West Siberian Plain: two lizard species (common lizard and sand lizard) and four species of snakes
(adder, grass snake, mamushi, and Pallas’s coluber).
The species are listed in the order of decreasing abundance. The common lizard and adder are most frequent
in southern taiga, the sand lizard — in subtaiga forests,
the grass snake and mamushi — in forest-steppe zone,
the Pallas’s coluber — in steppe zone. The common lizard and the adder both prefer uliginous pine forests
(riams). The lizard is also frequent in dry pineries and
less frequent in birch-aspen and dark coniferous-small-leaved forests. The largest distribution of the
grass snake falls on pine forests, of the sand lizard — on
raised, inundated plains of rivers which are rarely
flooded and noninundated birch-aspen forests. The
mamushi occurs only on rocky streamside southern
slopes, and the Pallas’s coluber evidently prefers steppe
areas.
Reptile Population
Using the frequency-abundance index for West Siberia extrapolated from the index for the Ob’ region, we
can classify the communities under study. The cluster
analysis has indicated seven types of population, one of
which (subextremal territories) is subdivided into two
subtypes, and another (suboptimal territories) — into
six.
The detailed classification reads as follows:
Types of population:
1 — extremal territories (subzones of arctic and
northern subarctic tundras, bogs of southern subarctic
and hummocky northern taiga – no reptiles found);
2 — subextremal territories.
Subtypes of population:
2.1 — subzones of southern subarctic tundras (except bogs), open boreal woodland and northern taiga
(except inundated plains and hummocky bogs; percentage of population: common lizard — 93, adder — 7;
density of population — 1 individual per square
kilometer);
2.2 — northern- and middle-taiga inundated plains
(common lizard — 1).
Types of population:
3 — pessimal middle taiga territories (pineries:
common lizard — 24);
4 — suboptimal territories (common and sand lizards — 51 and 33, adder and grass snake — 9 and 6;
546).
Subtypes of population:
4.1 — middle taiga (except pineries, inundated
plains and bogs; common lizard — 86, adder — 14;
403);
4.2 — non-pine forests and afforested lowland bogs
from southern taiga up to forest-steppe and raised bogs
of middle and southern taiga (common and sand lizards — 71 and 19, adder and grass snake — 9 and 0.8;
939).
Classes of population:
4.2.1 — middle and southern taiga (common and
sand lizards — 81 and 11, grass snake — 9; 948);
4.2.2 — subtaiga forests and forest-steppe (common
lizard — 57, sand lizard — 31, adder and grass snake —
11 and 2; 928).
4.3 — forest-steppe pineries (adder — 72, common
and sand lizards — 25 and 4; 540);
4.4 — inundated plains from southern taiga up to
forest-steppe (sand and common lizards — 70 and 20,
grass snake and adder — 8 and 2; 893);
4.5 — lowland open subtaiga bogs (common and
sand lizards — 82 and 10, adder and grass snake — 8
and 0.3; 353);
4.6 — lowland open forest-steppe bogs (common
and sand lizards — 84 and 9, adder — 7; 151).
Types of population:
5 — optimal territories (southern taiga and subtaiga
pineries; common and sand lizards — 67 and 33, grass
snake — 0.4; 4410);
6 — pessimal forest-steppe territories (meadows,
steppes, fields; common and sand lizards — 86 and 14;
21);
7 — pessimal steppe territories (meadows, steppes,
fields — sand lizard — 109).
The diagram constructed on the basis of this classification (Fig. 1) illustrates the change in communities in
line with latitudinally decreasing warmth provision, the
influence of prairiefication and ploughing up, salinity
and swamping, as well as with the composition of forest-forming tree species and floods. As the warmth provision increases from north to south the population
becomes denser up to subtaiga forests, but then in open
habitats of forest-steppe and steppe zone it decreases
because of prairiefication, ploughing up and the intrusion of steppe. High waters in inundated plains of
rivers, particularly in the north through middle taiga
lead to a decline in population density. Southward, under general soil dehydration, despite or owing to floods,
the habitats of inundated territories are more suitable
for reptiles than those of noninundated areas. The maximum total abundance of reptiles is recorded in southern-taiga and subtaiga pineries (4410 individuals per
square kilometer). In pine forests of middle taiga the
number decreases nearly 200 times, on the rest territory
of this subzone — 11 times. In non-pine forests and in
afforested lowland and transitory bogs of southern taiga
up to forest-steppe, and the raised bogs of middle and
southern taiga the reptile is 4 times less frequent than
within the territories optimal for them.
With increasing prairiefication, ploughing up and
intrusion of steppe the density of reptile population de-
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 1 No. 2 2008
Fig. 1. Spatial-typological structure of reptile population of West Siberia. Symbolic notations: triangle, apex up — population of open habitats poor in productivity; triangle, apex
down — the same, rich in productivity; circle — mosaic; square — afforested. Inside each symbol there are numbers of taxa of corresponding classification, index — intragroup likeness. Continuous lines between symbols indicate likeness above threshold, dash lines — for maximum value below threshold. Beside the lines is given intergroup likeness, beside the
symbols — occurring species, density of population and the number of species registered. The arrows beside the list of basic structure-forming factors of the environment show the
growth direction of their influence and population trends.
SPATIAL-TYPOLOGICAL STRUCTURE AND MAPPING
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 1 No. 2 2008
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RAVKIN et al.
creases by a factor of 30—44 and even 200 in comparison with the optimal biotopes. The minimum value,
disregarding territories with no reptiles found, reaches
1 individual per square kilometer to 4410 individuals
per square kilometer in optimal conditions.
Based on this classification, a map of reptile population has been compiled (Fig. 2). In general, it clearly
shows that zonal differentiation of population does not
exactly coincide with the zonal differentiation of West
Siberia. For instance, the population of arctic tundras is
of one type, while the communities of subarctic tundra,
forest-tundra and northern taiga, of another. The fourth
type and partly the populations of the third and fifth
types occupy forest zone from middle taiga to subtaiga
forests and forest in forest-steppe. Two other belts of
population fall on forest-steppe and steppe zones. The
general background shows the penetration of the first
type (population of extremal territories) through hummocky bogs up to northern taiga inclusively and into inundated plains within the tundra and forest-tundra
zones. The second type (population of subextremal territories) penetrates through the inundated plains southward up to middle taiga inclusive.
Specific populations of pineries are well recognized
on the map within the forest zone (from middle taiga):
those of middle taiga, of southern subzones of forest
zone – and, of separate open lowland bogs (southern subzones of the forest zone as opposed to foreststeppe).
One can compare the described map with population
maps of amphibians and small mammals [11, 12]. As to
both amphibians and reptiles, the latitudinal zonality of
population does coincide neither with natural zonality
nor with the zonality typical of reptile population. The
territories which are extremal for amphibians cover the
whole tundra zone, not only the arctic tundras as in case
of reptiles. Forest-tundra and northern taiga are shared
among types of communities of pessimal and subpessimal territories. The communities of subpessimal areas
are spread through steppe zone along the raised bogs
and dehydrated southern biotopes (fields, meadows,
steppes). Zonal and subzonal strips for amphibians are
distinguished with the ratio of optimal, suboptimal,
pessimal, and subpessimal conditions taken into account. For reptiles the strips are characterized by more
homogeneous population with spots of optimal conditions. Floodplains are more favorable for amphibians
than flood-free territories, while for reptiles – quite the
contrary. The series of optimality on graphs and maps
are more distinct for amphibians than for reptiles, while
the degree of optimality of biotopes plays an important
role for the common and sand lizards, as well as for the
adder. The heterogeneity of reptile population might be
reflected in a concise form by listing the habitats occupied by separate communities, whereas for amphibians
this list is great and diverse, though various limiting
factors have quite the same effect [13].
Small mammals show a larger degree of coincidence
with natural zonality than poikilotherm vertebrates.
Thus, the deviations are marked in the tundra zone
(population of Arctic zone differs from the Subarctic);
forest-tundra population occupies the subzones of open
boreal woodlands and most of northern taiga. The population of middle and southern taiga and northern forest-steppe are significantly similar, while southern
forest-steppe is inhabited by various types of communities. The habitats of small mammals are considerably
less heterogeneous than of poikilotherm vertebrates.
This, however, is consequence of preliminary data levelling (the map is based on contribution of species to the
Shannon index of diversity). The levelling was necessitated by significant annual variations in abundance of
small mammals in comparison to amphibians. Reptiles
show even larger degree of locality of occurrence
within complex landscape tracts than amphibians and
small mammals. The locality of distribution and oscillations variations linked to smaller amount of material
are compensated by preliminary group averaging of
map units on the basis of forest-forming tree composition. On the map “Flora of the West Siberian Plain”
(1976) these groups also include phytocenoses derived
from climax forests of homogeneous compound of forest-forming species. Thus, some differences of the
maps arise from methodological differences in preliminary preparation of data before cluster analysis. This,
however, compensates the drawbacks of the obtained
data which, in turn, allows comparison of the maps.
Population Structure
The assessment of force and commonness of
renations between environmental heterogeneity and
reptile population by linear quality approximation of
matrices of resemblance showed that zonality bears
maximum informativity (41% of variance for quotient
of similarity). Eight times less is the influence of forest-forming species composition (5), and additionally
5 times less is the influence of prairiefication and
solinity of soil (1 each) and humidification (0.8). All
the listed factors cover 43% of variance, 22 and 23% of
variance refer to the informativity of classification and
structure (together 23%). All together the factors and
conditions take into account 45% of variance, which
approximately corresponds to multiple correlation
coefficient equal to 0.67.
As a general rule, the relations between amphibian
population heterogeneity and the same environmental
factors are considerably weaker than of those of reptiles
(in respect to zonality — nearly three times weaker, to
composition of forest-forming species and humidification — the same; general integrated assessment is
half less). In regard to birds the assessments are more
close: zonality is equally informative, the total influence is one and a half less, the informativity of forest-forming species composition and humidification is
less considerably. In regard to small mammals the rela-
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 1 No. 2 2008
Fig. 2. Reptile population of the West Siberian Plain (second half of summer, original scale 1 : 10 000 000). See classification for population descriptions.
SPATIAL-TYPOLOGICAL STRUCTURE AND MAPPING
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RAVKIN et al.
tionship estimates are lower than to reptiles: zonality —
1.3 times, forest-forming species composition — 19
times and humidification — 4 times. In general the explanation is nearly of equal informativity (47 and 45%
of variance).
The integral assessment showed that birds have the
highest value of force and commonness of relations between environmental heterogeneity and population
(69%), which approximately one and a half times less
in regard to small mammals and even less — in regard
to amphibians (almost three times). These differences
in the degree of approximation arise from ecological
heterogeneity (diversity) of population and locality of
distribution of poikilotherms in comparison to homoiothermal animals.
The studies are supported by the international projects of SB RAS # 5.12 and 5.14.
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