The eastern macroslope of the Prepolar Urals is
inhabited by 40 mammalian species. Unique species
diversity associated with a great variety of mountain
landscapes is preserved in this relatively small area
because of a low degree of anthropogenic impact. This
study is an attempt to estimate the consequences of
considerable intensification of human activity planned
in connection with the exploitation of placer deposits.
Previously, such work caused slight damage to terres-
trial ecosystems because it affected the area of only
0.6--1.0 km 2 in each case. Nevertheless, these examples
allow one to predict, to a certain extent, the trend of
subsequent development of the anthropogenous pro-
cesses in natural landscapes of the region.
For a number of years, I studied the state of mam-
malian populations in the regions of placer deposits,
other mining enterprises, and wilderness areas.
Large mammals. Route censuses of animals and
their tracks were taken. Among large mammals, moose
and bears prevail in the region (Flerov, 1933). On aver-
age, one moose track and one bear track crossing the
route were registered each 1.5 and 2 km, respectively.
The tracks following along the route were not found on
rock dumps of gold mines but occurred in river valleys
not disturbed by mining. The tracks across the route
were often near ravines joining the valley. The absence
of differences between the results of censuses taken on
rock dumps and in undisturbed areas is explained by
the fact that the home ranges of these animals are con-
centrated in ravines and on the slopes of mountain
spurs. Bears in summer also occur in mountain tundras;
they cross river valleys during daily migrations from
one slope to another. Moose prefer forested slopes;
hence, their routes across river valleys were mainly in
the middle reaches, i.e., in the zone where gold placers
are commonly located. Consequently, further develop-
ment of gold mining in this region can have serious
ecological consequences. The exploitation of large
river valleys and neighboring tributaries of the same
river system will result in the fragmentation of animal
home ranges. In the summer and winter seasons, each
individual range occupies 2.5-39.0 and 0.8-7.5 km 2,
respectively (Filonov, 1993). Therefore, the loss of sev-
eral parts of these ranges, even as small as i km 2 in
area, will result in the substantial reduction of the total
area inhabited by individual animals, and, conse-
quently, in the decrease of animal abundance.
Further development of mining in the Prepolar
Urals will interfere with seasonal migrations of moose,
preserved population of wild reindeer, and a number of
rare (e.g., wolverine) and valuable commercial species
(sable and marten). The point is that these migrations
are generally directed from the plain to the mountains
and back; hence, as mining is carried out on tributaries
of large rivers flowing down from the main watershed,
areas with depleted deposits in river valleys cut across
the migration routes. As shown in some other regions of
the Urals (Bukhmenov, 1975; Filonov, 1993), the gen-
eral migration flow separates in such cases into discrete
streams, traditional migration routes are displaced to
new, less convenient locations, and the intensity of
migration decreases. As a result, some animals winter
under less favorable conditions, and their mortality
increases.
An important role belongs to the effects of other
industrial activities and anthropogenic factors associ-
ated with mining, such as road construction, land clear-
ing for house building, cutover and burned-out areas
appearing in the forests, uncontrolled hunting, and anx-
iety. Apparently, their adverse consequences will
become even more serious with the expansion of min-
ing industry. The road network will create additional
barriers to animal migration; tree cutting and burning in
large areas will bring about significant changes in food
composition and supply of both herbivorous and pred-
atory mammals, which lead to a short-term increase
and subsequent decrease in animal abundance
(Smirnov, 1987). Anxiety among animals will have the
gravest consequences, making them migrate to the
areas remote from the zone of industrial development
and concentrate there. This primarily concerns herbiv-
orous mammals. In the Prepolar Urals, where the productivity of ecosystems is relatively low, this process
will soon result in the depletion of food resources and
the consequent decrease in animal population size, as it
occurred with wild reindeer. To date, the local popula-
tions of large mammals have not been seriously
affected by uncontrolled hunting. In the Man'ya River
basin, for example, the estimated size of moose popu-
lation is approximately 200 animals, commercial hunt-
ing is virtually absent, and only two or three geologic
field crews usually work in this region; as members of
each crew shoot one or two moose per year, the total
annual loss is only three to six animals, i.e., 1.5-3%.
However, hunting pressure on these populations can
rapidly exceed the allowable limit and result in their
decline.
The aforementioned consequences of industrial
development are equally unfavorable for representa-
tives of the family Mustelidae. Small mustelids, such as
weasels and ermines, have home ranges of several doz-
ens of hectares (Danilov et al., 1979) but never inhabit
the areas of rock dumps; hence, the appearance of each
depleted mining site reduces the populations of these
species by one or several individuals. Damage from
mining is virtually irreversible, as the disturbed biogeo-
cenoses will apparently recover for centuries (the areas
of mines abandoned half a century ago are almost in the
same state). Small mustelids face the risk of losing con-
siderable proportions of their existing populations. This
especially concerns ermines, which prefer floodplain
biotopes. As to large mustelids, their home ranges
cover dozens of square kilometers, and the effects of
mining on these animals will be comparable to those on
bears and ungulates.
A relatively low population density of large mam-
mals is characteristic of the entire Prepolar Urals (Fle-
rov, 1933; Laptev, 1958; Berdyugin, 1997), and its
gradual decrease under the effects of anthropogenic
factors can rapidly bring many species to the point of
extinction. Therefore, instead of developing the mining
industry in this region, it would be expedient to orga-
nize there a specially protected area (e.g., national
park) providing for the conservation of the entire com-
plex of natural conditions, including the species diver-
sity of mammals. This measure will also prevent pollu-
tion of the rivers that provide spawning grounds for
valuable fish species.
Small mammals. The species of this group (rodents
and shrews), owing to their great abundance and diver-
sity, play an important role in the processes occurring
in natural communities. Differing from each other in
biological requirements, they perform different func-
tions in the communities and can promote their devel-
opment in one direction or another, depending on exist-
ing conditions.
The material on small mammals was collected in the
main types of their habitats, both natural (27 types) and
anthropogenically transformed to a greater or lesser
extent (11 types). The results of this investigation are
shown in the table.
In the Prepolar Urals, this animal group is repre-
sented by ten species, two of which occur only in their
natural habitats and are absent from anthropogenically
transformed biotopes. A decrease in species diversity
indicates degradation of animal communities. On the
other hand, water voles appeared in the same areas. The
occurrence of this species in the mountains provides
evidence that human activities resulted in the expansion
of river segments with a slow current and loose soils on
the banks. This also follows from the fact that the abun-
dance and proportion of root voles--rodents character-
ized by a similar mode of life increased in the com-
munities of anthropogenically transformed habitats
(Berdyugin, 1985). The expansion of such landscape
elements in the mountains can have grave conse-
quences, as strong spring and rain floods, which are
common in this zone, inevitably wash the soil away and
denude the banks.
In general, the structure of rodent communities in
natural habitats is characterized by the following fea-
tures. The total number of rodents is relatively low: this
is evidence that the productivity of both rodent commu-
nities and biocenoses of the Prepolar Ural Mountains as
a whole is also low. The northern red-backed vole is the
dominant species accounting for almost half the total
number of rodents in the communities. The large-
toothed red-backed vole, a specific mountain species in
the Urals, is subdominant. Three species--bank, field,
and Middendorff voles--are common and fairly
numerous components of the communities. Other spe-
cies are relatively rare. In anthropogenically trans-
formed habitats, the species ratio changes substantially:
the proportions of the dominant species and large-
toothed red-backed voles decrease, whereas those of
bank voles (more typical for the southern taiga) and
Middendorff voles (inhabitants of open tundra areas)
increase. Changes in the proportion of root voles in
rodent communities were considered above.
All distinctive features of rodent communities in the
anthropogenically transformed habitats indicate that
these areas are losing the landscape pattern characteris-
tic of the Prepolar Urals, i.e., coniferous forests are
replaced by deciduous ones, treeless areas increase in
size, and the structure of the herbaceous layer changes.
In addition, the system of slowly flowing streams is
formed (see above). Thus, landscape formations char-
acteristic of plains and atypical for highlands appear in
the mountain area. They are extremely insecure under
conditions of the mountain relief and cannot provide
for its stabilization. These events in their natural course
can lead to a general landscape crisis (Makunina,
1974).
The survey of areas exposed to less destructive
anthropogenic influences several decades ago showed
that changes in plant and rodent communities are gen-
erally interconnected. When the cessation of mining is
followed by the development of birch, mixed conifer-
ous-birch, and, at later stages, birch--coniferous herba-
ceous forest communities, bank voles become domi-
nant in the rodent community, and field voles increase
in number. Rodents can be abundant in such forest bio-
cenoses. Because of certain ecological peculiarities
(primarily those of feeding), these species interfere
with the reverse transformation of biocenoses into their
initial state, i.e., the state of equilibrium under given
environmental conditions. When anthropogenic effects
result in predominant development of herbaceous asso-
ciations, rodent communities are characterized by
increasing proportions of field voles and, in mountain
tundras, Middendorff's, bank, or gray-sided-backed
voles. The ecological effect of this phenomenon is the
same as in the previous case. Disturbed sites are often
occupied by plant communities with a weakly devel-
oped herbaceous-moss layer and low productivity
(even by local criteria). They are usually inhabited by
northern red-backed voles, but the density of these ani-
mals is extremely low. Finally, the variant closely
resembling the situation on fresh rock dumps was
observed in the areas where the plant cover and soils
were largely (but incompletely) destroyed in the course
of mining. The recovery of the biota in this case is com-
plicated and proceeds extremely slowly. For example, a
site of the sedge-moss tundra exposed to such an
impact more than 50 years ago is still uninhabited by
rodents.
To date, the biotic complex has been completely
destroyed in relatively small areas. However, further
development of gold mining and the resulting expan-
sion of such areas in the Prepolar Urals may have grave
consequences, up to the point of ecological disaster.
On the Problem of Steppe Ecosystem Conservation in the Central Caucasus
Key words: the Caucasus, steppe ecosystems, Arik Ridge, nature reserve.
The steppe biomes of Russia are endangered, and remaining steppe ecosystems have degraded to differ-ent extents under the impact of human activities. The main destructive factor is extensive agriculture, which cannot be profitable without expansion to new areas. The Caucasus as a whole and particularly the Northern Caucasus, one of the most densely populated regions of Russia with an economic system based primarily on agriculture, are not an exception.
Natural steppe ecosystems in the Northern Cauca-sus occupied western and central Ciscaucasia within Krasnodar and Stavropol regions, the Adygei and Kab-ardino-Balkar republics, and North Ossetia, giving way to semidesert ecosystems in plain and foothill areas lying farther east. Although the natural range of steppe cenoses in the region is limited and exposed to signifi-cant anthropogenic impact, there is not a single nature reserve established in order to conserve the steppe eco-systems of the Northern Caucasus. Piedmont forests are the main protected biomes in the Caucasian, Teberda, and North Ossetia nature reserves; subalpine and alpine meadows, in the Kabardino-Balkar High Mountain Reserve. In the Dagestan Reserve, which consists of two separate areas, attention is focused on the conser-vation of desert landscapes and coastal ecosystems of the Caspian Sea region. The necessity of establishing a forest–steppe nature reserve was substantiated by spe-cialists of Stavropol State University in 2000 (Godzevich et al., 2000).
Although the flora and fauna of the Northern Cauca-sus are unique among mountain regions of Russia, the total size of specially protected natural areas (SPNAs) in the region is the smallest, and the pattern of their dis-tribution (in clusters) does not comply with the require-ment for representativeness of the protected biota (Tishkov and Belonovskaya, 2004). Among measures to conserve steppe ecosystems in the Kabardino-Balkar Republic, of primary importance is the establishment of a nature reserve as the most effective type of SPNAs with regard to protection of ecosystems and their com-ponents. Basic criteria for identifying areas of nature-conser-vation significance are the systems of parameters char-acterizing the state of botanical and zoological objects.
In the former case, these are parameters such as rarity of plant communities, their floristic and phytosociolog-ical significance, reduction of their ranges, and the risk of their extinction (Zhuravleva, 1999). In 2006, special-ists of the Institute of the Ecology of Mountain Areas surveyed steppe ecosystems made in Maiskii, Prokhlad-nenskii, and Terskii raions of the Kabardino-Balkar Republic. The results of this survey show that relatively small areas of virgin land on the Arik Ridge fully sat-isfy the above criteria. The fact that these areas still retain their natural steppe vegetation has also been noted by other authors (Kos, 1959; Kerefov and Fiap-shev, 1977; Shkhagapsoev and Volkovich, 2002). Physiographic characteristics.The Arik Ridge, located in the northwest of the republic (43°20′–43°50′N, 44°–45°E), is actually a system of spurs of the Terek Ridge with a subdued topography and elevations of no more than 450 m a.s.l. The watershed and slopes are composed of Pliocene sand–clay conglomerate rock masses. Prevailing soils are micellar, calcareous, ordi-nary chernozems with low or medium humus contents (Kerefov and Fiapshev, 1968). According to physio-graphic zoning of the Northern Caucasus (Chupakhin, 1974), this area is in the Kabarda sloping-plain region of the Mineralnye Vody–Terek district of the Stavropol–Terek province (Central Ciscaucasia). Its climate is moderately continental, with annual average temperature and precipitation of 11.0°C and 522.6 mm (according to meteorological data from the city of Terek between 1987 and 2002) (Ashabokov et al., 2005). The bulk of precipitation falls in summer but is largely lost by evaporation and surface runoff. The area has no natural sources of water and receives it from the irrigation system that includes the Malo-Kabardinskii, Akbashskii, and Tambovskii canals.
Phytocenotic and floristic diversity.According to florogenetic zoning of the Central Caucasus (Galushko, 1976), the Arik Ridge is in the Terek–Sunzha region of the Ciscaucasian district of the Kuban–Terek piedmont steppe province. According to the results of the 2006 survey, its natural vegetation is composed mainly by herb–grass, grass–legume–herb, grass–wormwood, herb–licorice, and, to a lesser extent, beard grass, feather grass, and shrub–herb phytocenoses. Among grasses, common are steppe species such asKoeleria cristata(L.) Pers., Phleum phleoides(L.) Karst., Poa angustifoliaL., Festuca valesiacaGaudin, Helictotri-chon pubescens(Huds.) Pilg., Bromopsis riparia (Rehm.) Holub., and Bothryochloa ischaemum(L.) Keng. The last species is dominant in places, forming local beard grass communities. Feather grasses (Stipa lessingianaTrin. et Rupr. and S. pennataL.) are rare, communities with their participation are small and have limited distribution. Stipa pennataL. is included in the Red Data Books of the Russian Federation and Stavropol krai (Krasnaya kniga…, 1998, 2002). How-ever, this species is not listed in the Red Data Book of the Kabardino-Balkar Republic (Krasnaya kniga…, 2000), probably because its distribution in the republic has not been studied sufficiently. According to Kos (1959), the vegetation of the Arik Ridge in the mid-20th century included one more feather grass species, Stipa pulcherrima(included in the Red Data Books of the Russian Federation, Kabardino-Balkar Republic, and Stavropol krai), but we have not found it on the slopes surveyed.
Ephemeral grasses such as Anisantha tectorum(L.) Nevski, A. sterilis(L.) Nevski, Bromus japonicus Thunb., and Poa bulbosaL. abundantly develop in spring. In trampled areas,Hordeum leporinumLink. is dominant.
Herbage is rich in species. The dominant group includes Salvia verticillataL.,S. tesquicolaKlok. et Pobed., Filipendula vulgaris Moench, Agrimonia eupatoriaL., Galium verticillatum Danth., Centaurea dealbataWilld., and Scabiosa ochroleucaL. Some communities also contain large proportions of Polygala anatolicaBoiss. et Heldr., Fragaria viridis (Duch.) Weston, Ajuga orientalisL., and Poterium polygamum Waldst. et Kit. Areas used as grazing grounds are often overgrown with Thymus marschallianusWilld. and the legume Onobrychis bobroviiGrosshm., which is hardly eaten by livestock because of abundant pubescence. An area of natural steppe vegetation with species rarely occurring in the republic and other red-list plants was discovered on the southwestern slope of the Arik Ridge 5 km east of the village having the same name (230 m a.s.l.). The floristic diversity of two communi-ties described there in a 10 ×10-m plot on May 25, 2006, reached 35 species in the herb–grass community and 39 species in the shrub–herb community. In both cases, the herbaceous layer had 85% coverage. The group of rare species consisted ofPaeonia tenuifoliaL., Asparagus verticillatusL. (another species of the genus,A. officinalisL., is more common), Dictamnus caucasicus(Fisch. et C.A. Mey) Grossh., Clematis lathyrifoliaBess. et Reichenb., Adonis flammeaJacq., Amygdalus nanaL., and some other plants.
The fern-leaved peony P. tenuifolia(Paeoniaceae) occurs in Prokhladnenskii raion of the republic, in typ-ical steppe habitats on the slopes of Dzhenali, Terek, and Arik ridges (Kos, 1959; Shkhagapsoev and Slonov, 1987; Krasnaya kniga…, 2000; our collections of 2006 from the Arik Ridge). The species is included in the Red Data Book of North Ossetia (Krasnaya kniga…, 1981), where it occurs along the Sunzha Ridge, and in the Red Data Books of the Russian Federation and Kab-ardino-Balkar Republic (Krasnaya kniga…, 1988, 2000). On the Arik Ridge, the average population den-sity of P. tenuifoliain the communities mentioned above is 7 ind./m 2 . In the same communities, single individuals of Bieberstein’s peony P. biebersteiniana Rupr. can be found. This species visually differs from P. tenuifoliain having broader leaf segments, and the dates of blooming and fruiting in these two species are also different. According to our observations, most P. tenuifoliaplants on May 16, 2006, already entered the stage of fruiting, while P. biebersteinianaplants were still blooming. It should be noted, however, that the taxonomic status of the latter species is ambiguous, since some botanists regard it as a subspecies of P. tenuifolia. In any case, it is listed as a true species in the Red Data Book of Stavropol Krai (Krasnaya kniga…, 2002).
Shrub communities consist mainly of Amygdalus nanaL. accompanied by Frangula alnusMill. and Rhamnus pallasiiFisch. et C.A. Mey. Communities described above contain no endemic species. Similar communities probably grow also on the Terek and Sunzha ridges but are absent in other regions of the republic. On this basis, such communi-ties may be classified as rare. The fauna of the region is poorly studied, and reli-able information on its present-day species diversity is almost absent. According to available data, large mam-mals are represented by the red fox and jackal. The ornithofauna includes the steppe eagle and small birds of prey; little bustard; among gallinaceous birds, quail; among passerines, the bee-eater Merops apiaster, which nests on ravine slopes.
Thus, the Arik Ridge is exposed to considerable anthropogenic impact: the major part of land is plowed, and most of the remaining part is under uncontrolled grazing load. In addition, population decline in some plant species (e.g., P. teniofoliaand feather grasses) is also explained by their commercial harvesting for dec-orative purposes. These factors are responsible for deg-radation of primary phytocenoses, expansion of weeds. and destruction of habitats favored by different species of the local fauna.
Thus, to conserve the steppe cenoses of the Central Caucasus within the Kabardino-Balkar republic, it is necessary (1) to establish a steppe nature reserve up to 10 000 ha in area on the Arik Ridge and (2) to include vascular plant species such as Stipa pennata, S. lessin-giana, Asparagus verticillatus, and Amygdalus nanain the Red Data Book of the republic.
The establishment of such a reserve will provide a basis for measures to restore populations of the little and great bustards, unique species of typical steppe ornithofauna that had been widespread in steppe eco-systems, including those of the Northern Caucasus.
In the mid-20th century, the little bustards used to nest on the Arik Ridge (A.N. Kudaktin, personal communi-cation) and the great bustard was recorded during the periods of flight and local winter migrations in the Cen-tral Caucasus (Beme, 1958).
The steppe biomes of Russia are endangered, and remaining steppe ecosystems have degraded to differ-ent extents under the impact of human activities. The main destructive factor is extensive agriculture, which cannot be profitable without expansion to new areas. The Caucasus as a whole and particularly the Northern Caucasus, one of the most densely populated regions of Russia with an economic system based primarily on agriculture, are not an exception.
Natural steppe ecosystems in the Northern Cauca-sus occupied western and central Ciscaucasia within Krasnodar and Stavropol regions, the Adygei and Kab-ardino-Balkar republics, and North Ossetia, giving way to semidesert ecosystems in plain and foothill areas lying farther east. Although the natural range of steppe cenoses in the region is limited and exposed to signifi-cant anthropogenic impact, there is not a single nature reserve established in order to conserve the steppe eco-systems of the Northern Caucasus. Piedmont forests are the main protected biomes in the Caucasian, Teberda, and North Ossetia nature reserves; subalpine and alpine meadows, in the Kabardino-Balkar High Mountain Reserve. In the Dagestan Reserve, which consists of two separate areas, attention is focused on the conser-vation of desert landscapes and coastal ecosystems of the Caspian Sea region. The necessity of establishing a forest–steppe nature reserve was substantiated by spe-cialists of Stavropol State University in 2000 (Godzevich et al., 2000).
Although the flora and fauna of the Northern Cauca-sus are unique among mountain regions of Russia, the total size of specially protected natural areas (SPNAs) in the region is the smallest, and the pattern of their dis-tribution (in clusters) does not comply with the require-ment for representativeness of the protected biota (Tishkov and Belonovskaya, 2004). Among measures to conserve steppe ecosystems in the Kabardino-Balkar Republic, of primary importance is the establishment of a nature reserve as the most effective type of SPNAs with regard to protection of ecosystems and their com-ponents. Basic criteria for identifying areas of nature-conser-vation significance are the systems of parameters char-acterizing the state of botanical and zoological objects.
In the former case, these are parameters such as rarity of plant communities, their floristic and phytosociolog-ical significance, reduction of their ranges, and the risk of their extinction (Zhuravleva, 1999). In 2006, special-ists of the Institute of the Ecology of Mountain Areas surveyed steppe ecosystems made in Maiskii, Prokhlad-nenskii, and Terskii raions of the Kabardino-Balkar Republic. The results of this survey show that relatively small areas of virgin land on the Arik Ridge fully sat-isfy the above criteria. The fact that these areas still retain their natural steppe vegetation has also been noted by other authors (Kos, 1959; Kerefov and Fiap-shev, 1977; Shkhagapsoev and Volkovich, 2002). Physiographic characteristics.The Arik Ridge, located in the northwest of the republic (43°20′–43°50′N, 44°–45°E), is actually a system of spurs of the Terek Ridge with a subdued topography and elevations of no more than 450 m a.s.l. The watershed and slopes are composed of Pliocene sand–clay conglomerate rock masses. Prevailing soils are micellar, calcareous, ordi-nary chernozems with low or medium humus contents (Kerefov and Fiapshev, 1968). According to physio-graphic zoning of the Northern Caucasus (Chupakhin, 1974), this area is in the Kabarda sloping-plain region of the Mineralnye Vody–Terek district of the Stavropol–Terek province (Central Ciscaucasia). Its climate is moderately continental, with annual average temperature and precipitation of 11.0°C and 522.6 mm (according to meteorological data from the city of Terek between 1987 and 2002) (Ashabokov et al., 2005). The bulk of precipitation falls in summer but is largely lost by evaporation and surface runoff. The area has no natural sources of water and receives it from the irrigation system that includes the Malo-Kabardinskii, Akbashskii, and Tambovskii canals.
Phytocenotic and floristic diversity.According to florogenetic zoning of the Central Caucasus (Galushko, 1976), the Arik Ridge is in the Terek–Sunzha region of the Ciscaucasian district of the Kuban–Terek piedmont steppe province. According to the results of the 2006 survey, its natural vegetation is composed mainly by herb–grass, grass–legume–herb, grass–wormwood, herb–licorice, and, to a lesser extent, beard grass, feather grass, and shrub–herb phytocenoses. Among grasses, common are steppe species such asKoeleria cristata(L.) Pers., Phleum phleoides(L.) Karst., Poa angustifoliaL., Festuca valesiacaGaudin, Helictotri-chon pubescens(Huds.) Pilg., Bromopsis riparia (Rehm.) Holub., and Bothryochloa ischaemum(L.) Keng. The last species is dominant in places, forming local beard grass communities. Feather grasses (Stipa lessingianaTrin. et Rupr. and S. pennataL.) are rare, communities with their participation are small and have limited distribution. Stipa pennataL. is included in the Red Data Books of the Russian Federation and Stavropol krai (Krasnaya kniga…, 1998, 2002). How-ever, this species is not listed in the Red Data Book of the Kabardino-Balkar Republic (Krasnaya kniga…, 2000), probably because its distribution in the republic has not been studied sufficiently. According to Kos (1959), the vegetation of the Arik Ridge in the mid-20th century included one more feather grass species, Stipa pulcherrima(included in the Red Data Books of the Russian Federation, Kabardino-Balkar Republic, and Stavropol krai), but we have not found it on the slopes surveyed.
Ephemeral grasses such as Anisantha tectorum(L.) Nevski, A. sterilis(L.) Nevski, Bromus japonicus Thunb., and Poa bulbosaL. abundantly develop in spring. In trampled areas,Hordeum leporinumLink. is dominant.
Herbage is rich in species. The dominant group includes Salvia verticillataL.,S. tesquicolaKlok. et Pobed., Filipendula vulgaris Moench, Agrimonia eupatoriaL., Galium verticillatum Danth., Centaurea dealbataWilld., and Scabiosa ochroleucaL. Some communities also contain large proportions of Polygala anatolicaBoiss. et Heldr., Fragaria viridis (Duch.) Weston, Ajuga orientalisL., and Poterium polygamum Waldst. et Kit. Areas used as grazing grounds are often overgrown with Thymus marschallianusWilld. and the legume Onobrychis bobroviiGrosshm., which is hardly eaten by livestock because of abundant pubescence. An area of natural steppe vegetation with species rarely occurring in the republic and other red-list plants was discovered on the southwestern slope of the Arik Ridge 5 km east of the village having the same name (230 m a.s.l.). The floristic diversity of two communi-ties described there in a 10 ×10-m plot on May 25, 2006, reached 35 species in the herb–grass community and 39 species in the shrub–herb community. In both cases, the herbaceous layer had 85% coverage. The group of rare species consisted ofPaeonia tenuifoliaL., Asparagus verticillatusL. (another species of the genus,A. officinalisL., is more common), Dictamnus caucasicus(Fisch. et C.A. Mey) Grossh., Clematis lathyrifoliaBess. et Reichenb., Adonis flammeaJacq., Amygdalus nanaL., and some other plants.
The fern-leaved peony P. tenuifolia(Paeoniaceae) occurs in Prokhladnenskii raion of the republic, in typ-ical steppe habitats on the slopes of Dzhenali, Terek, and Arik ridges (Kos, 1959; Shkhagapsoev and Slonov, 1987; Krasnaya kniga…, 2000; our collections of 2006 from the Arik Ridge). The species is included in the Red Data Book of North Ossetia (Krasnaya kniga…, 1981), where it occurs along the Sunzha Ridge, and in the Red Data Books of the Russian Federation and Kab-ardino-Balkar Republic (Krasnaya kniga…, 1988, 2000). On the Arik Ridge, the average population den-sity of P. tenuifoliain the communities mentioned above is 7 ind./m 2 . In the same communities, single individuals of Bieberstein’s peony P. biebersteiniana Rupr. can be found. This species visually differs from P. tenuifoliain having broader leaf segments, and the dates of blooming and fruiting in these two species are also different. According to our observations, most P. tenuifoliaplants on May 16, 2006, already entered the stage of fruiting, while P. biebersteinianaplants were still blooming. It should be noted, however, that the taxonomic status of the latter species is ambiguous, since some botanists regard it as a subspecies of P. tenuifolia. In any case, it is listed as a true species in the Red Data Book of Stavropol Krai (Krasnaya kniga…, 2002).
Shrub communities consist mainly of Amygdalus nanaL. accompanied by Frangula alnusMill. and Rhamnus pallasiiFisch. et C.A. Mey. Communities described above contain no endemic species. Similar communities probably grow also on the Terek and Sunzha ridges but are absent in other regions of the republic. On this basis, such communi-ties may be classified as rare. The fauna of the region is poorly studied, and reli-able information on its present-day species diversity is almost absent. According to available data, large mam-mals are represented by the red fox and jackal. The ornithofauna includes the steppe eagle and small birds of prey; little bustard; among gallinaceous birds, quail; among passerines, the bee-eater Merops apiaster, which nests on ravine slopes.
Thus, the Arik Ridge is exposed to considerable anthropogenic impact: the major part of land is plowed, and most of the remaining part is under uncontrolled grazing load. In addition, population decline in some plant species (e.g., P. teniofoliaand feather grasses) is also explained by their commercial harvesting for dec-orative purposes. These factors are responsible for deg-radation of primary phytocenoses, expansion of weeds. and destruction of habitats favored by different species of the local fauna.
Thus, to conserve the steppe cenoses of the Central Caucasus within the Kabardino-Balkar republic, it is necessary (1) to establish a steppe nature reserve up to 10 000 ha in area on the Arik Ridge and (2) to include vascular plant species such as Stipa pennata, S. lessin-giana, Asparagus verticillatus, and Amygdalus nanain the Red Data Book of the republic.
The establishment of such a reserve will provide a basis for measures to restore populations of the little and great bustards, unique species of typical steppe ornithofauna that had been widespread in steppe eco-systems, including those of the Northern Caucasus.
In the mid-20th century, the little bustards used to nest on the Arik Ridge (A.N. Kudaktin, personal communi-cation) and the great bustard was recorded during the periods of flight and local winter migrations in the Cen-tral Caucasus (Beme, 1958).
Carbon Concentrations and Caloric Value of Organic Matter in Northern Forest Ecosystems
Key words: north, taiga, forest ecosystems, carbon, caloric value.
The data on the carbon content in different plant organs and their caloric value are necessary for evaluat-ing the bioproduction process in phytocenoses and for studying the carbon cycle and energy and mass exchange in forest biogeocenoses. According to published data, the carbon content in individual biomass fractions amounts to 50–57% of their dry weight in conifers and to 42–48% in deciduous woody plants (Risser, 1985; Kobak, 1988; Vogt, 1991). Most of researchers estimating carbon stock in forest communities assume that it accounts for 50% of the absolutely dry weight of trunk, roots, and branches and for 45–50% of the weight of green plant parts (Makarevskii, 1991; Bidsey, 1990; Uglerod v ekosiste-makh…, 1994; Tsel’niker and Malkina, 1994; Kobak, 1989; Isaev et al., 1993, 1996). The caloric value of plant components in the ecosystems of forest zone is better studied (Ovington, 1961; Golley, 1961; Kur-batskii, 1962; Molchanov, 1971; Kononenko, 1975; Dadykin and Kononenko, 1975; Dem’yanov, 1974, 1981; Ivask, 1983, 1985; Vookova, 1985).
Utkin (1986) thoroughly analyzed the available data on the caloric values of plants and animals. He found that the heat of plant combustion as a physical parame-ter is characterized by a relatively high variability, being dependent on plant species, growing conditions, morphological structure, age, period of sampling, and other factors. However, many aspects of plant differen-tiation with respect to their caloric value remain unclear. Special studies are necessary for elucidating the relationships between the heat of plant combustion and multiple environmental factors, the intensity of physiological processes, and the biochemical composi-tion of organic matter synthesized and accumulated by plants. Moreover, publications provide virtually no data on the carbon content and caloric value of organic mat-ter in forest ecosystems of the European Northeast. The purpose of this work was to determine the car-bon content and caloric value of different phytomass fractions. The following plants were studied: trees Pinus sylvestrisL., Picea obovataLedeb., Betula pen-dulaRoth., Populus tremulaL., and Larix sibirica Ledeb., Fl. Alt.; dwarf shrubs Vaccinium vitis-idaeaL., V. myrtillusL., and V. uliginosumL.; mosses Pleuro-zium schreberi, Hylocomium splendens, Polytrichum commune, and Sphagnumsp.; mixed herbaceous sam-ples including Trientalis europeaL., Maianthemum bifoliumL., Equisetum silvaticumL., and Agrostis tenuisSibth.; and lichens Cladinasp. The main compo-nents of plant fall and litter were also analyzed. The study was carried out in pine and spruce phyto-cenoses of the middle taiga subzone in the Komi Republic (62¡N, 50¡20′E). Plant samples for analysis were collected in the end of the growing period (August to September) simultaneously with estimating the phy-tomass of the plant communities. The carbon concen-trations in phytomass fractions were determined in an ANA-1500 automatic nitrogen and carbon analyzer (Carboro Erba, Italy); the caloric value, by the combus-tion method according to Kochan (1982). Measure-ments were made in ten biological and three to eight analytical replications. The experimental data were processed statistically by conventional methods. As follows from Tables 1 and 2, the range of carbon concentrations in different phytomass fractions of trees was 44.6–50.3% dry weight; in plants of the herb– dwarf shrub layer, 41.9–53.4%; in mosses and lichens, 42.3–45.4%; in forest litter, 45.8–48.2%; and in the components of plant fall, 44.6–53.1%. The carbon con-centrations in trees varied insignificantly: the coeffi-cient of variation (CV) was 2.4% for individual species and 1.5–4.3% for phytomass components within a spe-cies. The range of carbon concentrations in plants of ground vegetation was slightly higher, but the variation of this parameter by species did not exceed 10% in these plants and 2.8% in mosses and lichens. The car-bon concentrations in individual fractions of plant fall and different types of litter differed by 5 and 8.6%, respectively.
The analysis of data on the caloric value of tree plants shows that this parameter of individual phyto-mass fractions varied from 16.81 to 21.77 kJ/g in spruce, from 16.40 to 22.91 kJ/g in pine, from 17.91 to 21.56 kJ/g in larch, and from 16.66 to 20.95 kJ/g in birch (Table 3). The coefficients of variation were 10.3, 8.8, 7.0, and 5.2%, respectively. Higher energy values were typical of trunk wood and large roots. The caloric value of plants in the lower layers of coniferous communities varied from 17.44 to 19.76 kJ/g; of forest litter, from 17.37 to 18.46 kJ/g; and of plant fall, from 16.58 to 19.89 kJ/g. The variation in this parameter both in plants of ground vegetation and in the litter was insignificant: the coefficients of variation were 3.0 and 2.4%, respectively These data can be used for making up the energy and carbon balance and for studying energy flows in forest ecosystems of the taiga zone..
The data on the carbon content in different plant organs and their caloric value are necessary for evaluat-ing the bioproduction process in phytocenoses and for studying the carbon cycle and energy and mass exchange in forest biogeocenoses. According to published data, the carbon content in individual biomass fractions amounts to 50–57% of their dry weight in conifers and to 42–48% in deciduous woody plants (Risser, 1985; Kobak, 1988; Vogt, 1991). Most of researchers estimating carbon stock in forest communities assume that it accounts for 50% of the absolutely dry weight of trunk, roots, and branches and for 45–50% of the weight of green plant parts (Makarevskii, 1991; Bidsey, 1990; Uglerod v ekosiste-makh…, 1994; Tsel’niker and Malkina, 1994; Kobak, 1989; Isaev et al., 1993, 1996). The caloric value of plant components in the ecosystems of forest zone is better studied (Ovington, 1961; Golley, 1961; Kur-batskii, 1962; Molchanov, 1971; Kononenko, 1975; Dadykin and Kononenko, 1975; Dem’yanov, 1974, 1981; Ivask, 1983, 1985; Vookova, 1985).
Utkin (1986) thoroughly analyzed the available data on the caloric values of plants and animals. He found that the heat of plant combustion as a physical parame-ter is characterized by a relatively high variability, being dependent on plant species, growing conditions, morphological structure, age, period of sampling, and other factors. However, many aspects of plant differen-tiation with respect to their caloric value remain unclear. Special studies are necessary for elucidating the relationships between the heat of plant combustion and multiple environmental factors, the intensity of physiological processes, and the biochemical composi-tion of organic matter synthesized and accumulated by plants. Moreover, publications provide virtually no data on the carbon content and caloric value of organic mat-ter in forest ecosystems of the European Northeast. The purpose of this work was to determine the car-bon content and caloric value of different phytomass fractions. The following plants were studied: trees Pinus sylvestrisL., Picea obovataLedeb., Betula pen-dulaRoth., Populus tremulaL., and Larix sibirica Ledeb., Fl. Alt.; dwarf shrubs Vaccinium vitis-idaeaL., V. myrtillusL., and V. uliginosumL.; mosses Pleuro-zium schreberi, Hylocomium splendens, Polytrichum commune, and Sphagnumsp.; mixed herbaceous sam-ples including Trientalis europeaL., Maianthemum bifoliumL., Equisetum silvaticumL., and Agrostis tenuisSibth.; and lichens Cladinasp. The main compo-nents of plant fall and litter were also analyzed. The study was carried out in pine and spruce phyto-cenoses of the middle taiga subzone in the Komi Republic (62¡N, 50¡20′E). Plant samples for analysis were collected in the end of the growing period (August to September) simultaneously with estimating the phy-tomass of the plant communities. The carbon concen-trations in phytomass fractions were determined in an ANA-1500 automatic nitrogen and carbon analyzer (Carboro Erba, Italy); the caloric value, by the combus-tion method according to Kochan (1982). Measure-ments were made in ten biological and three to eight analytical replications. The experimental data were processed statistically by conventional methods. As follows from Tables 1 and 2, the range of carbon concentrations in different phytomass fractions of trees was 44.6–50.3% dry weight; in plants of the herb– dwarf shrub layer, 41.9–53.4%; in mosses and lichens, 42.3–45.4%; in forest litter, 45.8–48.2%; and in the components of plant fall, 44.6–53.1%. The carbon con-centrations in trees varied insignificantly: the coeffi-cient of variation (CV) was 2.4% for individual species and 1.5–4.3% for phytomass components within a spe-cies. The range of carbon concentrations in plants of ground vegetation was slightly higher, but the variation of this parameter by species did not exceed 10% in these plants and 2.8% in mosses and lichens. The car-bon concentrations in individual fractions of plant fall and different types of litter differed by 5 and 8.6%, respectively.
The analysis of data on the caloric value of tree plants shows that this parameter of individual phyto-mass fractions varied from 16.81 to 21.77 kJ/g in spruce, from 16.40 to 22.91 kJ/g in pine, from 17.91 to 21.56 kJ/g in larch, and from 16.66 to 20.95 kJ/g in birch (Table 3). The coefficients of variation were 10.3, 8.8, 7.0, and 5.2%, respectively. Higher energy values were typical of trunk wood and large roots. The caloric value of plants in the lower layers of coniferous communities varied from 17.44 to 19.76 kJ/g; of forest litter, from 17.37 to 18.46 kJ/g; and of plant fall, from 16.58 to 19.89 kJ/g. The variation in this parameter both in plants of ground vegetation and in the litter was insignificant: the coefficients of variation were 3.0 and 2.4%, respectively These data can be used for making up the energy and carbon balance and for studying energy flows in forest ecosystems of the taiga zone..
Geographic Trends in the Accumulation of Heavy Metals in Mosses and Forest Litters in Karelia
Heavy metals (HMs) are considered to be among
priority technogenic pollutants. To solve ecological
problems related to the environmental effects of HMs
in the Russian North, it is necessary to make a detailed
inventory of their contents in natural objects in different
areas with regard to the diversity of climatic and soil-geochemical conditions and the degree of industrial
development in these areas. It is known that mosses are
informative indicators of aerotechnogenic environmen-tal pollution. Forest litters are important as the struc-tures retaining and accumulating various pollutants.
The contents of HMs in the soil depends on the distance
from local pollution sources and, to a large extent, on
the pattern of pollutant transfer in the upper layers of
the atmosphere. An important role belongs to region-specific natural factors, i.e., local climate, relief, vege-tation, and soils.
The Republic of Karelia is located on the Baltic shield, which forms the northwestern part of the Russian platform. The vast area of the republic (117300 km 2 )
extends from the north to the south for 672 km; hence, the climate, geological structure, hydrographic net-work, soils, and vegetation in different parts of the republic are heterogeneous.
The climate in Karelia is relatively mild, with a long mild winter and a short cool summer; considerable cloudiness, high humidity, and changeable weather are characteristic of all seasons. The prevailing form of atmospheric circulation over the territory of Karelia is the western transfer of air masses. The formation of precipitation is also accounted for by moisture evapo-rated from the White Sea and numerous lakes and bogs, which cover one-third of the Karelian territory. Vegeta-tion has a considerable effect on the migration of sub-stances. In Karelia, coniferous forests are the dominant type of vegetation.
The spectrum of possible sources of technogenic HM pollution in Karelia is wide. There are 10284 sources of industrial emissions into the atmosphere, and most of them are concentrated in the cities of Petrozavodsk, Segezha, Kostomuksha, and Kondo-poga. The total amount of emissions from large indus-trial enterprises of these cities reaches 128600 tons per year. A complex combination of technogenic factors and natural geochemical conditions in Karelia deter-mines the pattern of HM distribution over its territory. In this work, we studied green mosses (Pleurozium schreberi, Hylocomium splendens) and forest litters. The former indicate the state of the atmosphere over a relatively short period of time (approximately three years), and the chemical composition of the latter reflects the impact of long-term industrial pollution (over more than ten years). The chemical analysis of mosses and litters can provide information about the sources, ranges, and extents of environmental pollu-tion, as well as reveal major pollutants. Our studies were performed by internationally accepted methods (Atmospheric Heavy Metal…, 1996).
Samples of green mosses and forest litters were taken from test plots of the bioindication network cov-ering the entire Karelian territory. The contents of iron, manganese, chromium, copper, nickel, zinc, cobalt, lead, and cadmium in the samples were determined by atomic absorption spectrometry.
We also estimated the effects of climatic parameters (wind rose, precipitation rate) on the distribution of aerotechnogenic pollutants containing HMs over the territory of the republic. The data on each of eight wind directions recorded by the Karelian hydrometeorologi-cal observatory (N, S, W, E, NE, NW, SE, SW) was assessed quantitatively with respect to wind stability, i.e., the frequency of its occurrence as a percentage of the total number of observations (without calm winds). Taking into account wind directions in winter and sum-mer and different weather patterns in the cold or warm periods of the year, the parameters of stability were averaged. Thus, we distinguished cold winters with lit-tle snow from warm, snowy winters and cold, rainy summers from warm, dry summers.
To estimate the correctness of grouping (homogene-ity within each group and heterogeneity of different groups), stepwise discriminant analysis was used. Its results confirmed that all five groups were identified correctly: they proved to be internally homogeneous and did not overlap with one another. The main dis-criminators (major pollutants) in forming regional groups with respect to the pollution of mosses are nickel, cobalt, chromium, and cadmium. According to their significance for group formation, they can be arranged in the following series: Co > Cr > Ni > Cd. In the case of forest litters, the main discriminators arranged in the same order are as follows: Fe > Mn > Pb > Zn.
The results of pairwise comparisons of the regional groups in the three-factor spaces with respect to HM contents in mosses and forest litters (Table 3) demon-strated that differences were significant only for groups I and II, especially concerning the contents of cad-mium. In the second group (Segezhskii and Med-vezh’egorskii raions), differences between HM accu-mulation in mosses and forest litters were significant for the majority of elements (especially for copper) and nonsignificant for zinc and iron.
Thus, we revealed the existence of geographic trends in the distribution of pollutants over the Karelian territory and their accumulation in mosses and forest litters.
The Republic of Karelia is located on the Baltic shield, which forms the northwestern part of the Russian platform. The vast area of the republic (117300 km 2 )
extends from the north to the south for 672 km; hence, the climate, geological structure, hydrographic net-work, soils, and vegetation in different parts of the republic are heterogeneous.
The climate in Karelia is relatively mild, with a long mild winter and a short cool summer; considerable cloudiness, high humidity, and changeable weather are characteristic of all seasons. The prevailing form of atmospheric circulation over the territory of Karelia is the western transfer of air masses. The formation of precipitation is also accounted for by moisture evapo-rated from the White Sea and numerous lakes and bogs, which cover one-third of the Karelian territory. Vegeta-tion has a considerable effect on the migration of sub-stances. In Karelia, coniferous forests are the dominant type of vegetation.
The spectrum of possible sources of technogenic HM pollution in Karelia is wide. There are 10284 sources of industrial emissions into the atmosphere, and most of them are concentrated in the cities of Petrozavodsk, Segezha, Kostomuksha, and Kondo-poga. The total amount of emissions from large indus-trial enterprises of these cities reaches 128600 tons per year. A complex combination of technogenic factors and natural geochemical conditions in Karelia deter-mines the pattern of HM distribution over its territory. In this work, we studied green mosses (Pleurozium schreberi, Hylocomium splendens) and forest litters. The former indicate the state of the atmosphere over a relatively short period of time (approximately three years), and the chemical composition of the latter reflects the impact of long-term industrial pollution (over more than ten years). The chemical analysis of mosses and litters can provide information about the sources, ranges, and extents of environmental pollu-tion, as well as reveal major pollutants. Our studies were performed by internationally accepted methods (Atmospheric Heavy Metal…, 1996).
Samples of green mosses and forest litters were taken from test plots of the bioindication network cov-ering the entire Karelian territory. The contents of iron, manganese, chromium, copper, nickel, zinc, cobalt, lead, and cadmium in the samples were determined by atomic absorption spectrometry.
We also estimated the effects of climatic parameters (wind rose, precipitation rate) on the distribution of aerotechnogenic pollutants containing HMs over the territory of the republic. The data on each of eight wind directions recorded by the Karelian hydrometeorologi-cal observatory (N, S, W, E, NE, NW, SE, SW) was assessed quantitatively with respect to wind stability, i.e., the frequency of its occurrence as a percentage of the total number of observations (without calm winds). Taking into account wind directions in winter and sum-mer and different weather patterns in the cold or warm periods of the year, the parameters of stability were averaged. Thus, we distinguished cold winters with lit-tle snow from warm, snowy winters and cold, rainy summers from warm, dry summers.
To estimate the correctness of grouping (homogene-ity within each group and heterogeneity of different groups), stepwise discriminant analysis was used. Its results confirmed that all five groups were identified correctly: they proved to be internally homogeneous and did not overlap with one another. The main dis-criminators (major pollutants) in forming regional groups with respect to the pollution of mosses are nickel, cobalt, chromium, and cadmium. According to their significance for group formation, they can be arranged in the following series: Co > Cr > Ni > Cd. In the case of forest litters, the main discriminators arranged in the same order are as follows: Fe > Mn > Pb > Zn.
The results of pairwise comparisons of the regional groups in the three-factor spaces with respect to HM contents in mosses and forest litters (Table 3) demon-strated that differences were significant only for groups I and II, especially concerning the contents of cad-mium. In the second group (Segezhskii and Med-vezh’egorskii raions), differences between HM accu-mulation in mosses and forest litters were significant for the majority of elements (especially for copper) and nonsignificant for zinc and iron.
Thus, we revealed the existence of geographic trends in the distribution of pollutants over the Karelian territory and their accumulation in mosses and forest litters.
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