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15===== DYNAMICS OF ECOSYSTEMS AND THEIR COMPONENTS ==
UDC 599.735.53:591.5
THE PAST AND PRESENT OF SAIGA IN RUSSIA: IS THERE A FUTURE?
© 2022. T.Yu. Karimova*, **, A.A. Lushchekina*, V.M. Neronov*, Yu.N. Arylov***, N.Yu. Pyurvenova****
*A.N. Sever tsov Institute of Ecology and Evolution of the Russian Academy of Sciences
33, Leninsky Prospekt, Moscow, 119071, Russia. E-mail: rusmabcom@gmail.com **Water Problems Institute of the Russian Academy of Sciences 3, Gubkina Str., Moscow, 119333, Russia. E-mail: katayur@gmail.com
***B.B. Gorodovikov Kalmyk State University 11, Pushkina Str., Elista, 358000, Russia. E-mail: kalmsaiga@mail.ru ****Independent Researcher 54a, Germasheva Per., Elista, 358001, Russia. E-mail: arylova@gmail.com
Received November 01, 2022. Revised November 10, 2022. Accepted December 01, 2022.
The evolutionary strategies of the saiga antelope, such as an early reproductive ability, high female fertility, polygamy, herd instinct and migration, have been helping them to survive since the Late Pleistocene in the changing environmental conditions. In the Holocene they were forced to coexist with humans, and so a new stage in their history began. Aside from the mass hunting, exterminating the saigas, the human impact on their habitat has also increased due to the widespread agriculture, especially in Western Europe, which eventually reduced the species' range. By the early 20th century, only a few patches of their large range remained: the untouched areas of the lower reaches of the Volga River in Europe; Ustyurt, Betpak-Dala, the Ili-Karatal interfluve, China and Mongolia in Asian territory. The conservation measures implemented in the 1920s by the Soviet Union government preserved five saiga populations that currently exist in the world. Four of them (Northwest Pre-Caspian population in Russia; Ural population in Kazakhstan, Russia; Ustyurt population in Kazakhstan, Uzbekistan, Turkmenistan, Russia; Betpakdala population in Kazakhstan, Russia) belong to the nominative subspecies S. t. tatarica, while the fifth one (S. t. mongolica (=S. borealis)) inhabit Western Mongolia. Since the end of the 20th century, the state of the Northwest Pre-Caspian population has been of particular concern. The reason for that is the severe decrease in its number (from 800,000 in 1958 to 5,000 in 2015), and of its main habitat (from 60,000-70,000 km2 to 2,000-3,000 km2). A significant part of this population switched to a sedentary lifestyle in the protected areas of the "Chernyye Zemli" ecological region. A long-term shortage of mature males (<10%) slowed down the population growth and, consequently, decreased its numbers. However, thanks to various protective measures, the Northwest Pre-Caspian population has been gradually growing since 2016, and reached 18,000-19,000 in 2022. The further growth depends on the effectiveness of the actions that are supposed to be carried out as part of the " Strategy for the Conservation of the Saiga in the Russian Federation". In addition to effective protection and full-scale monitoring, the strategy includes the removal of various obstacles that hinder saiga migration and cause habitat fragmentation; improving the quality of the habitats; creating new protected areas on different levels in the most suitable habitats and their integration into a single network via ecological corridors; development and expansion of environmental education activities.
Keywords: saiga, saiga population, animal numbers, range, Northwest Pre-Caspian Region, saiga preservation.
DOI: 10.24412/2542-2006-2022-4-28-49 EDN: QVMVAB
In recent years biodiversity has been decreasing at an unprecedented rate, seriously undermining the ability of the biosphere to support life on planet Earth. According to experts,
about a million species of plants and animals are currently endangered (The Global ..., 2019). As a result, the most important ecosystems that provide many ecological services, including those needed for food production and agriculture development, are being destroyed rapidly, which may lead to food shortages and health deterioration of the humanity (State of ..., 2019). The disappearance of some populations among the most species of large mammals is of particular concern, because it has been happening for the past 100-200 years, while their numbers and the areas of their ranges keep generally declining (Ceballosa et al., 2017). The reasons for that are climate change, environmental pollution, changing systems of land use, as well as the direct danger of recreational hunting, large-scale fishing and poaching (Briggs, 2017). As a result, the loss of one population at first and the entire species of vertebrates later will lead to the destruction of complex ecological networks that include animals, plants and microorganisms.
Today all ungulates that have ever lived in the arid territories of Russia, such as saiga, Mongolian gazelle, onager and Przewalski's horse, are listed in the "Red Data Book of the Russian Federation" (2021), but only saiga and gazelle can still be found in the country. The state of the saiga population has been of great concern since the late 90s of the 20th century, because it is the only European antelope the habitat of which is located in the Northwest Pre-Caspian Region, although in the recent past it was a common and numerous species in the southern regions of the European part of Russia.
Evolution, Taxonomy and Range of Saiga
Despite the long history of saiga and its belonging to the so-called "mammoth" fauna of Eurasia, it is unknown where the genus Saiga originates from. Those fossil remains that were found in sediments are never older than the Middle Pleistocene, which indicates a wide distribution of the species in Eurasia and America, as well as that it does not differ from the modern species both morphologically and ecologically. These facts suggest that the saiga ancestors came from the Caprinae group in the second half of the Miocene or in the early Pliocene (Baryshnikov et al., 1998). During the Quaternary period, the saiga inhabited vast tundra steppes, from the British Isles in the west to Alaska and northwestern Canada in the east, from the New Siberian Islands in the north to the Caucasus in the south. The increasing climate warmth and humidity at the turn of the Pleistocene and Holocene erased tundra steppes, making the saiga adapt to xerophytic steppe and desert landscapes on the southern periphery of its former range. The dynamics of the spatial structure of its range was always determined by the physical and geographical factors, but a new factor was added at the beginning of the Holocene: the necessity to learn and coexist with humans. In addition to mass hunting, the influence of people on the saiga habitat began to increase due to their widespread farming, especially in Western Europe, which then started to shape the species range.
According to the many scientists (Baryshnikov et al., 1998; Kuznetsova et al., 2002), this antelope is the only species of the genus Saiga, which has 5 subspecies: 3 are extinct -Saiga tatarica binagadensis (Transcaucasia), S. t. prisca (Europe and Western Siberia), S. t. borealis (Eastern Siberia and Alaska); 2 are still present - the nominative Saiga tatarica tatarica Linnaeus, 1766 (distributed on the vast plains in the Northwest Pre-Caspian Region, Kazakhstan, Central Asia, Western Dzungaria) and Saiga tatarica mongolica Bannikov, 1946 (small range in Mongolia). Other scientists (Sher, 1967; Baryshnikov, Tikhonov, 1994) suggest that there were two species in the Pleistocene: the European-Kazakh (S. tatarica) and the East Siberian-American (S. borealis). Based on this information and on the slight morphological differences between these two subspecies, the latest edition of "Mammal Species of the World" (2005) gave them the status of species, and the Mongolian saiga was named S. borealis Tschersky, 1876, as reflected in some of the official documents that define the distribution of S. tatarica
and S. borealis in Eurasia.
As a mass species of the open arid territories, the saiga was a valuable commercial animal and played an important role in the life of steppe nomads. In the early 18th century, its range extended from the Black Sea steppes in the west to the arid territories of China and Mongolia in the east (Kirikov, 1966; Dinesman, 1998; Fig. 1). However, in the first half of the 19th century, due to hunting and land development, its range was divided into 3 separate parts: European (Eastern Europe), Middle Asian (Kazakhstan, Uzbekistan, Turkmenistan), Central Asian (Mongolia, China). In the second half of the 19th century, due to people quickly populating the steppes, the saiga almost completely disappeared from Europe, and its Asian range also decreased drastically. As a result, by the beginning of the 20th century, only a few patches of its once vast range remained: the untouched areas of the lower reaches of the Volga River in Europe; Ustyurt, Betpak-Dala, the Ili-Karatal interfluve, China and Mongolia in Asian territory. The total number of saiga in the territory of the former USSR was about 1000 heads (Bannikov et al., 1961), meaning that the species was on the verge of extinction, and so it was decided to increase protection measures. After the saiga hunting was banned in the European part of Russia in 1921, and in Kazakhstan and Central Asia in 1923, as well as after the export of the saiga horns over the border of the USSR was stopped in the early 1930s, the species finally began to recover (Zhirnov, 1982). In 1952 in the Northwest Pre-Caspian Region there were 180,000 animals, while the maximum number in the USSR in the entire history of census was recorded in 1974, when the amount of saiga reached 1,650,000.
Fig. 1. The range of the saiga. Legend: 1 - in the Holocene (Baryshnikov et al., 1998), 2 - Saiga tatarica tatarica (E - Northwest Pre-Caspian population; K1 - Ural population, K2 - Ustyurt population, K3 - Betpakdala population), 3 - meeting points of S. t. tatarica in recent years, 4 - S. t. mongolica (=S. borealis).
These days there are 5 saiga populations on the planet. Four of them are the nominative subspecies S. t. tatarica (the Northwest Pre-Caspian population in Russia; the Ural population in Kazakhstan, Russia; the Ustyurt population in Kazakhstan, Uzbekistan and Turkmenistan, Russia; the Betpakdala population in Kazakhstan, Russia. The fifth population (S. t. mongolica (=S. borealis)) inhabits Western Mongolia (Fig. 1). One more population of S. t. tatarica that used to live in the northwest of China, in the basins of the Dzhungar and Tacheng Rivers, as well as in
the adjacent southwestern areas of Mongolia, was exterminated in 1960-1970s for their horns that were an ingredient of Chinese medicine. However, the local shepherds claimed that they saw small groups of 3-4 saigas until the very 1984 (Gao et al., 2011).
Current Condition of Saiga Populations
Every population suffers with its own unique issues, but the general reasons that negatively affect their abundance are the same for the entire saiga range. They are climatic, such as harsh winters, zud (i.e. the lack of food during winters due to ice crust that prevents them from grazing) and droughts during the growing seasons; biotic, such as diseases, parasites and predators; anthropogenic, which can be direct, such as fishing and poaching, or indirect, such as reduction and fragmentation of natural habitats due to the increasing rates of agricultural land development, decreasing fodder capacity of pastures, construction of various obstacles, such as canals, roads and fences across the fields and pastures where the saiga migrations and seasonal migrations take place.
There are 4 subpopulations of the "Mongolian" saiga (S. t. mongolica=S. borealis) in Western Mongolia, near the Mankhan Somon, in Shargiin Gobi, Khuisiin Gobi and in the Dorgon Steppe (Buuveibaatar et al., 2013). According to some scientists (Zhirnov et al., 1998b), the sedentary lifestyle of this subspecies, their low breeding potential and, consequently, low numbers determine their high vulnerability to the negative effects of climatic, biotic (i.e. diseases) and anthropogenic factors. Although saiga hunting has been banned since 1930 (Milner-Gulland et al., 2020), there were fewer than 1,000 "Mongolian" saiga in the remote parts of the Gobi in the 1970-1980s (Lushchekina et al., 1999). However, with the help of such conservation measures as the inclusion of the species in the Red Book of Mongolia in 1987, creation of new protected areas that include about 24% of the saiga range (Clark et al., 2006), increased protection and active educational work with the locals, the situation changed for the better. According to the censuses of 2014, the population size was about 15,000 heads, while the range was almost 4,700 km2 (Chimeddorzh et al., 2016). Unfortunately, after the mass death in 2017 caused by the plague among the small ruminants, and due to severe weather, only 3,800 animals remained by the end of 2018 (Fewer than ..., 2019). The measures that took place in 2019-2020, such as livestock vaccination, extra feeding during the winter season and increased protection rate of the saiga, turned out to be successful: the population expanded up to 10,077 individuals in October 2021 (Mongolian ..., 2021).
The preservation measures aimed at the nominative subspecies (S. t. tatarica) that were taken by the USSR government in 1920-1930, yielded good results as well. By the middle of the 20th century the number of saiga increased to the point that in 1951 hunting was allowed once again in the Northwest Pre-Caspian Region, with some of the Kazakhstan regions following in 1954. Despite the annual catch of 10,000-115,000 saigas in the Pre-Caspian Region and 31,000501,000 in Kazakhstan, by the beginning of the 1980s the total number was about 1.2 million: 400,000 in the Pre-Caspian Region, and 800,000 in Kazakhstan, Uzbekistan and Turkmenistan (Bekenov, Grachev, 1998; Zhirnov, Maksimuk, 1998; Fig. 2). The collapse of the Soviet Union in the 1990s and the following prolonged economic crisis, the opened borders and conditionally free overseas trade, the increased demand for saiga horns in China and countries of Southeast Asia increased the poaching rates throughout the antelopes' range. Over 10 years the total number of 3 populations (Ural, Betpakdala, Ustyurt) decreased by 45 times, from 976,000 in 1993 to 21,100 in 2003 (Grachev et al., 2009); and by 8 times in the Northwest Pre-Caspian Region, from 148,000 to 18,500 (Bliznyuk, 2009). It is important to note that in 2003 the number of saiga in Russia and Kazakhstan, as well as in Uzbekistan and Turkmenistan, were comparable enough, however, after 20 years these figures have changed significantly to 18,500 in the Northwest Pre-Caspian Region
(Saigas ..., 2022) and 1 million 318,000 in Kazakhstan (The Fate of ..., 2022). It should also be taken into account that from 2010 to 2015, almost 230,000 animals of the Ural and Betpakdala populations died to epizootics (Karimova et al., 2021).
Our study showed that in the early 2000s the main biological parameters that characterize the four populations (Northwest Pre-Caspian, Betpakdala, Ustyurt, Ural) were comparable. They include fertility and barrenness of female saiga, sex ratio of their newborns, offspring, sex and age composition of the population that can be determined in July-August. However, the proportion of adult males and the yield per female in July-August in the Pew-Caspian region (11.5±1.9% and 0.81±0.18) were higher than those in Kazakhstan (8.7±3.8% and 0.48±0.40; Karimova et al., 2021). This data allow us to guess that the numbers differ so widely due to the external factors.
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2 3 -4
Fig. 2. Dynamics of the Betpakdala (1), Ustyurt (2), Ural (3) and Northwest Pre-Caspian (4) saiga populations.
The government of the Republic of Kazakhstan, along with the international non-governmental and intergovernmental organizations, invested a lot of resources to developed a network of specially protected natural areas in the country and ensure protection of three local populations. Since 2006, the territory of protected areas in the saiga range has increased by 2.5 times after two new areas were founded, the "Bokeyorda" and "Altyn Dala" nature reserves that include the territory of the "Sarykopinsky" sanctuary; and after the existing ones were expanded, i.e. the "Andasaisky" sanctuary, "Irgiz-Torgay" and "Korgalzhyn" nature reserves and the "Ile-Alatau" national park. The final area eventually amounted to more than 6 million ha (On Approval ..., 2017). In 2014 the first ecological corridor was constructed, connecting the key areas (Bragina, 2015). Furthermore, 30 mobile inspection groups that were common in the USSR were created once again; each consisted of 150 inspectors, provided with modern means of transportation, satellite communication and navigation devices, audio and video surveillance devices (Minoransky, Dankov, 2016). Many projects are currently underway, such as scientific researches and environmental educational work with the locas. Every year, saiga is monitored in Kazakhstan with the help of aerial surveys and ground censuses, and a special program was made to track individual animals that were tagged with satellite-linked collars (Milner-Gulland et al., 2020). Improved protection and tougher
penalties that were implemented in 2018 as part of "The Criminal Code of the Republic of Kazakhstan" (2022) concerning the saiga derivatives and saiga hunting, significantly reduced the pressure caused by poaching and eventually created conditions for a rapid growth of the species.
An individual saiga population of Russia inhabits the "Chernye Zemli" ecological region in the Northwest Pre-Caspian Region, which includes the eastern territories of the Republic of Kalmykia and the south-western parts of the Astrakhan Region. The Astrakhan, Volgograd and Saratov Regions adjacent to Kazakhstan also include part of the Ural population range (Neronov et al., 2013). Recently, due to the growth of the said population and the spring-summer drought in Kazakhstan, the saiga herds consisting of many thousands animals started entering these territories for a short period every year, causing some discontent among local farmers (Saigas ..., 2022). Besides, the Betpakdala population sometimes can be sighted in the Orenburg Region (Levykin et al., 2015).
The Fate of the Northwest Pre-Caspian Population
It was mentioned above that in a few decades of the early 20th century the saiga, previously an endangered species of the USSR, became a commercial one. By 1940 its range on the right bank of the Volga River covered most regions, and by the end of the 1940s there were more than 100,000 animals (Zhirnov, Maksimuk, 1998). In the 1950s-1960s, the area of its main habitat was 60,00070,000 km2, increasing up to 150,000-170,000 during droughts and snowy winters (Fig. 3a),
Fig. 3. The area of distribution of the Northwest Pre-Caspian saiga population a) in the 1950-1960s (Bannikov et al., 1961; Zhirnov, 1982), b) in the 1970-1990s (Zhirnov et al., 1998a). Legend: 1 -saiga's main range, 2 - areas they visited.
while their total number was approximately 800,000 individuals (Zhirnov et al., 1998a; Bliznyuk, 2009). Due to the numbers being so high and the saiga habit to form herds the food resources on the pastures quickly ran out, which forced them to move to new territories, searching for food. During their long-distance migrations that would sometimes cover 420 km, the saigas
could be encountered almost everywhere outside the "Chernye Zemli" ecological region: in Astrakhan, Volgograd and Rostov Regions, Stavropol Krai, the Republics of Kalmykia, Dagestan and Chechnya (Zhirnov et al., 1998a). However, there were areas of their seasonal gatherings. In winter saigas preferred the south, i.e. the eastern slopes of the Yergeni Elevation and the "Chernye Zemli" region; in spring they moved to the north, to calve in the Sarpinsky Lowland; in summer they used to disperse, but the majority grazed to the north and west of the territory where their calving took place (Bannikov et al., 1961).
By the 1980s the large-scale agricultural development of the territory near the Sarpinsky Lakes that started in the 1970s, such as construction of roads and settlements, a network of irrigation canals, and plowing of land (Bliznyuk, 1995), reduced the area of the main saiga habitat to 30,000 km2 (Fig. 3b). The amount of small livestock in Kalmykia that grew up to 3.04 million in 1981, the increased competition for food resources, and sometimes harsh weather conditions forced the numerous herds of saigas, the number of which varied from 240,000 up to 715,000 from the early 1970s to the mid-1980s (Bliznyuk, 2009), migrate over almost 100,000 km2 in search of the better pastures. During that period they were spotted rarely in the Rostov Region, Stavropol Krai and the Republic of Dagestan (Minoransky, Dankov, 2016). Economy, development of agriculture, oil and gas fields, and construction of necessary infrastructures eventually became the reason of the reduction of saiga, so by the end of the 1980s their number was about 150,000-160,000 (Zhirnov, Maksimuk, 1998).
In the 1990s the unregulated poaching that intensified severely after the collapse of the USSR, as well as the following collapse of the budgetary structures, including those necessary for nature protection, led to a sharp drop in the saiga population in the region: in 2006 there were 15,00020,000 animals (Melnikov, Sidorov, 2009), and only 4,000-5,000 remained by 2015 (Karimova, Lushchekina, 2018). According to some experts (Kuhl et al., 2009), this is the fastest, the most catastrophic decline in the number of mammals ever recorded in our world. Trying to explain the drop from the late 1990s to the early 2000s, various hypotheses were discussed. Some authors linked it with long-term cyclic fluctuations and the fact that the saiga population potential was declining (Sidorov, Bukreeva, 1999). Others believed it was caused by the changes in vegetation cover throughout the saiga habitats (Abaturov, 2007; Abaturov et al., 2008). Another considered reason was the increased wolf population. Over time, however, only the version of A.A. Danilkin (2005) was confirmed; he explained the decline in numbers was caused by the unregulated, insatiable manner that the saiga resources were used with and the lack of proper animal protection.
Aside from the poaching for meat during that period, the saiga males were hunted for their horns that were of high value in Chinese medicine, which consequently disrupted the sex and age structure of the entire population. The high fertility of the Northwest Pre-Caspian population, while abundance of the animals was so low (Karimova et al., 2020), could not be fulfilled due to the lack of mature males, because a normal reproduction cycle requires at least 15-20% of male animals, but by August 2002 their number had dropped to 1.1%, declining further to the critical level of 0.6% by the rutting season (Milner-Gulland et al., 2003). Despite the measures taken, such unbalanced proportion persisted in subsequent years as well. Thus, in August 2014, the proportion of mature males in the population was only 5.7%, and by December 2014 it even decreased to 0.72% (Chronicle of 2014). There were critical drops before, indeed, but they never lasted for so long. For example, in the winter of 1958-1959 after the excessive commercial hunting the proportion of adult males decreased to 1-2% by the rutting season (Bannikov et al., 1961). However, due to the high pliability of the saiga species and their adaptive abilities developed during evolution, as well as a decrease in the number of hunted animals, the population recovered in just a few years.
The improvement in the age and sex structure observed in recent years for this population ECOSYSTEMS: ECOLOGY AND DYNAMICS, 2022, Vol. 6, No. 4
(Fig. 4) allows us to hope for its prolonged depression to be near the end (Bogun, 2019). It is also confirmed by the data of the expert assessment of the species abundance, because in July 2022 there were already 18,000-19,000 saigas (Saigas ..., 2022).
Due to the decline in the population, the main habitat area of the saiga also decreased (Fig. 5). In recent years, despite the rapid growth in the number of small livestock from 312.8 thousand in 1998 to 1328.3 thousand in 2019 (Livestock ..., 2019) in the east of the Republic of Kalmykia (Chernozemelsky, Yustinsky and Yashkulsky Regions), the not so numerous saiga herds no longer migrate in search of new pastures. Instead they prefer to stay in protected areas, such as the steppe area of the "Chernye Zemli" state nature reserve in the Republic of Kalmykia and the "Stepnoy" sanctuary in the Astrakhan region, or near them, in an area of about 3,000 km2 (Karimova, Lushchekina, 2018), which is only 10.2% of the optimal habitats offered by this region (Paltsyn, 2016). A study of the vegetation of the northwestern part of the Pre-Caspian Lowland that was carried out in 2021-2022 by the World Wildlife Fund (WWF), as part of the "Saiga Habitat Assessment of the Northwest Pre-Caspian Region" project, showed that plant communities in most of the territory are in good condition (according to I.N. Safronova's report), and their diversity and species composition are suitable enough to expand the modern saiga range, if its abundance continues to grow.
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Fig. 4. Dynamics of sexually mature saiga males in the Northwest Pre-Caspian population.
Preservation Measures Taken to Protect the Saiga Population in the Northwest Pre-Caspian Region
In recent decades the problem of the saiga species conservation became an important task of global significance. In 1995 the species was included into Appendix II of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). At the 18th CITES Conference of the Parties in 2019, Geneva an annotation was accepted to "establish a zero export quota for specimens that are removed from nature for commercial purposes", which is equal to the species being temporarily listed in Appendix I of CITES. In 2002 the International Union for Conservation of Nature (IUCN/IUCN) added the saiga to its Red List. It is also included in Appendix II of the Convention on the Conservation of Migratory Species of Wild Animals
(Bonn Convention), so that the countries where the listed species live must focus on making agreements that are important for the preservation of those species and prioritize the species the safety of which remains in question. The Russian Federation is not currently a Party to the Bonn Convention, but it joined the Memorandum of Understanding Concerning Conservation, Restoration and Sustainable Use of the Saiga Antelope in 2009. The Memorandum works though the Medium-Term International Work Programs and includes every country of the saiga range, as well as some public organizations.
Fig. 5. Distribution of saiga in the Northwest Pre-Caspian region in 2000-2022. Legend: 1 -the main range, 2 - the sites of sightings.
In 2012 the Ministry of Natural Resources and Ecology of the Russian Federation and the Ministry of Agriculture of the Republic of Kazakhstan signed a bilateral interdepartmental
"Agreement on the Protection, Reproduction and Use of the Ural Saiga Population". In 2020 the Ministry of Ecology, Geology and Natural Resources of the Republic of Kazakhstan and the Ministry of Natural Resources and Ecology of the Russian Federation signed the "Agreement on the Protection, Reproduction and Use of Transboundary Saiga (Saiga tatarica tatarica) Populations".
Over the past 30 years the Russian Federation also took a number of measures to preserve the saiga and its habitats. In 1998 the saiga hunting was banned; in 2001 the President of Kalmykia signed a decree "On Emergency Measures for the Protection and Conservation of the Kalmyk Saiga Population"; 2010 was named the "Year of the Saiga Antelope" in Kalmykia; in 2013 the species was included in the list of "especially valuable wild animals and water biological resources belonging to the species of the Red Data Book of the Russian Federation and (or) protected by international treaties of the Russian Federation", the illegal extraction, keep, acquisition, storage, transportation, shipment and sale of which is criminally punished in accordance with Article 258.1 of the Criminal Code of the Russian Federation (2017). The species is also listed in the Red Data Books of the Republic of Kalmykia (2013), Rostov Region (2014), Astrakhan Region (2014) and the Russian Federation (2021).
In 1990 in order to ensure the protection of saiga habitats and monitor the species successfully, the state nature reserve "Chernye Zemli" was founded in Kalmykia, the steppe area of which is 94,000 ha, with a protection zone of 57,000 ha. In 2010 this reserve accepted the management of three other federal sanctuaries: "Mekletinsky" - 102,500 ha, "Sarpinsky" - 195,900 ha, "Kharbinsky" - 163,000 ha. For more than 20 years the "Stepnoy" regional sanctuary has been effectively operating in the Astrakhan Region, with its small but professional team protecting saiga over an area of about 104,000 ha. In addition, the research works are carried out in the sanctuary. Since 2004, the local inspectors have been monitoring the saiga species, while many local and foreign researchers have chosen the territory as a model range for their researches, where the biology, ethology and ecology of the saiga are studied along with the biodiversity of the region itself. The biotechnical measures are also being taken to prevent fires in the territory of the "Stepnoy" sanctuary. Plus, to provide saigas and other animals with water during the droughts, several artesian wells were cleared to create watering places for them (Fig. 6), with the support of the "Russian Caucasus" Branch of the WWF Russia. The employees of the sanctuary work a lot with the local people to raise their environmental awareness and responsibility by welcoming them to participate in various ecological activities. This work, including exhibitions, competitions and field trips, has an especially fruitful result among children from various educational institutions of the Astrakhan Region and neighboring territories.
Many attempts to keep and breed saigas in various zoos around the world have failed (Karimova et al., 2018). However, in order to preserve its gene pool three breeding centres were created in Russia. In 2000, the "Yashkul" captive breeding station was built on the basis of the Centre for Wild Animals of the Republic of Kalmykia (abolished in 2014); in 2003, the "Saiga" captive breeding centre was created on the basis of the "Astrakhan" State Hunting Farm in the Astrakhan Region, as well as the Centre of Rare Animals of European Steppes of the "Wild Nature of the Steppe" Association in the Rostov Region (Karimova et al., 2017). After annexation of Crimea in 2014, another semi-free centre for saigas was founded in the Tarkhankut National Nature Park, where 10 animals were brought to in 2013 from the "Askania Nova" bio sphere reserve (Ukraine). However, there are no saigas there at the moment. The listed centres study the biological and ethological characteristics of the species, carry out various veterinary activities, including artificial insemination, and raise animals to release them into the wild.
A factor that indicates successful saiga maintenance and breeding in the breeding centres is its population dynamics. The best results were achieved in the "Yashkul" captive breeding station that took in only 56 individuals and then increased the saiga number to 220 individuals over 15 years (2000-2014; Fig. 7). In the "Saiga" captive breeding centre in the Astrakhan Region, despite it
being built in an unsuitable habitat and therefore being exposed to frequent outbreaks of pasteurellosis and other diseases, the livestock of 30-40 individuals was maintained for quite a long time thanks to well-established zootechnical and veterinary measures, and some males were released into the wild (Karimova et al., 2017). Unfortunately, over the past 5 years the situation has changed drastically, and now only 8 animals (6 males and 2 females) remain in the breeding centre (according to S.A. Kalashnikov's report). The Centre of Rare Animals of European Steppes in Rostov Region currently considers itself the most successful captive breeding centre in Russia (Minoransky, Dankov, 2016). However, in 2010-2013 the number of their saiga decreased gradually due to no new animals being introduced to the Centre population from the wild; they declined from 70 to 43 individuals. Only in 2015-2016 the numbers grew after 32 animals were transferred to a much larger enclosure near the Manych Village; currently there are about 80 individuals that live in its territory (Minoransky et al., 2021).
Fig. 6. A wolf in the background and a black-winged stilt (Red Data Book of the Russian Federation) near an artesian well (taken by a camera trap, from the archive of the "Stepnoy" sanctuary, July 2022).
Breeding saigas in the breeding centres is quite promising, especially now that the methods for animals' keeping, breeding, transporting and capturing are well explored. The success of the "Yashkul" breeding station showed that the number of animals can be increased tenfold in a fairly short period of time. However, successful breeding of saigas requires several conditions, such as a naturally suitable area, a large and safe enclosure, constant veterinary monitoring, passionate specialists and state support.
Fig. 7. Saigas in the "Yashkul" Captive Breeding Station (photo by T.Yu. Karimova, June 2012).
Since 2019 the saiga is listed among the 11 priority animal species of the "Ecology" National Project, the purpose of which is to preserve and restore rare and endangered species as part of the federal project "Conservation of Biological Diversity and Development of Ecological Tourism". The said project released several documents that define the main areas of activity for the study, conservation, protection and restoration of the saiga population:
- a roadmap for the conservation and restoration of the saiga until 2024;
- an conservation plan for the saiga in the Russian Federation until 2025, created in accordance with the Memorandum of Understanding Concerning Conservation, Restoration and Sustainable Use of the Saiga Antelope;
- a conservation strategy for the saiga in the Russian Federation until 2030 and a plan for its implementation.
The "Strategy for the Conservation of the Saiga in the Russian Federation" approved in 2021 (Order ..., 2021) suggests some measures aimed at preserving the species in the country. According to it, the number of viable saiga population of the Northwest Pre-Caspian Region should be at least 20,000 individuals by 2030, while its range area should increase to 20,000 km2. In order to achieve these goals, it is necessary to solve many problems: to improve the legal and methodological base of saiga conservation; develop a network of protected areas; support the conservation of the saiga and its most important habitats; conduct a complex of researches and regular monitoring; optimize nature management throughout the saiga habitats; preserve the population of saigas in the semi-free enclosures; improve educational work with the local people and development of ecological tourism.
The data collected over the recent decades on the size of the Northwest Pre-Caspian saiga population are preliminary. One of its collection methods is a census conducted via ground vehicles, but it is unable to cover the entire range. Before that, the low-flying aircrafts or helicopters were used, but it is not possible anymore due to the noise, which can frighten and cause death of the
saigas. Therefore, the method of satellite data is the most promising one (Rozhnov et al., 2014), but the cost of high resolution images is very high. In May 2022, the Ministry of Natural Resources of Russia adopted the methodology developed by the WWF Russia for counting saiga in the Northwest Pre-Caspian Region using unmanned aircraft, which can make it possible to count the population size, as well as to determine its sex and age structure (Census ..., 2022). The advantage of this technique is the low disturbance factor, while the use of computer programs will eliminate significant errors during calculations. We would like to hope that in the near future we will find out the actual number of saigas in the Northwest Pre-Caspian Region.
Conclusions
The evolutionary strategies of the saiga antelope, such as an early reproductive ability, high female fertility, polygamy, herding and migration, have been helping them to survive since the Late Pleistocene in the changing environment. Usually, when their abundance is threatened by the natural factors like harsh winters, droughts, diseases and predators, they restore it rather quickly. Unfortunately, the saiga was unable to get used to the human's disruptive activities, such as agricultural land development, hunting and poaching.
The preservation of the Northwest Pre-Caspian saiga population and the restoration of its numbers currently depend on the different measures that should be taken in the future. In addition to effective protection and full-scale monitoring, this includes removal of various obstacles that hinder saiga migration and cause habitat fragmentation, improving the quality of the said habitats, creating new protected areas in the most suitable habitats and their integration into a single network via ecological corridors, development and expansion of environmental education activities. We hope and expect that the "Saiga Conservation Strategy in the Russian Federation" that was approved by the Ministry of Natural Resources of Russia in 2021 (Order ..., 2021) will contribute greatly to this.
Acknowledgements. The authors express their sincere gratitude to the staff of the "Stepnoy" sanctuary and its director V.G. Kalmykov for all these years of fruitful cooperation. The authors also thank A.N. Gilev for the photographs that were taken in the "Stepnoy" sanctuary.
Funding. The work was carried out for the A.N. Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences as part of the research work "Fundamental Problems of Wildlife Protection and Rational Use of Bioresources", State Assignment No. 1021062812203-8, as well as for the Water Problems Institute of the Russian Academy of Sciences as part of the research work for 2022-2024 "Study of Geoecological Processes in Hydrological Systems of Land, Formation of the Quality of Surface and Ground Waters, Problems of Water Resources Management and Water Use under Conditions of Climate Change and Anthropogenic Impact", No. FMWZ-2022-0002, State Registration No. AAAA-A18-118022090104-8.
REFERENCES REFERENCES
1. Abaturov BD. Saiga population in Russia and 1. Абатуров Б.Д. 2007. Популяция сайгака в problems of its conservation [Populyatsiya России и проблемы ее сохранения // saygaka v Rossii i problemy yeye sokhraneniya] Вестник Российской академии наук. Т. Bulletin of the Russian Academy of Sciences 77. № 9. С. 785-793.
[Vestnik Rossiyskoy akademii nauk], 2007;77 2. Абатуров Б.Д., Ларионов К.О., Джапова
(9):785-793. Р.Р., Колесников М.П. 2008. Качество
2. Abaturov BD, Larionov KO, Dzhapova RR, кормов и обеспеченность сайгаков (Saiga Kolesnikov MP. Forage quality and food supply tatarica) пищей в условиях for saigas (Saiga tatarica) under conditions of восстановительной смены regenerative vegetation change in the Chernye растительности на Черных Землях
Zemli of Kalmykia [Kachestvo kormov i obespechennost' saygakov (Saiga tatarica) pishchey v usloviyakh vosstanovitel'noy smeny rastitel'nosti na Chernykh Zemlyakh Kalmykii] Zoological Journal. 2008;87 (12):1524-1530.
3. Bannikov AG, Zhirnov LV, Lebedeva LS, Fandeev AA. Biology of the saiga [Biologiya saygaka]. Moscow: Izd-vo sel'skokhozyaystvennoy literatury, zhurnalov i plakatov, 1961:336.
4. Baryshnikov GF, Dmitrieva EL, Krakhmalnaya TV, Sher AV. Origin, evolution and systematics of the saiga [Proiskhozhdeniye, evolyutsiya i sistematika saygaka] Saiga: phylogeny, systematics, ecology, protection and use [Saygak: filogeniya, sistematika, ekologiya, okhrana i ispol'zovaniye]. Moscow: Tipografiya Rossel'khozakademii, 1998:9-20.
5. Bekenov AB, Grachev YuA. The number of saigas in the 20th century. Kazakhstan [Chislennost' saygakov v KHKH veke. Kazakhstan] Saiga: phylogeny, taxonomy, ecology, protection and use [Saygak: filogeniya, sistematika, ekologiya, okhrana i ispol'zovaniye]. Moscow: Tipografiya Rossel'khozakademii, 1998:225-227.
6. Bliznyuk AI. The role of economic development of the territory in changing the number of the Kalmyk saiga population [Rol' khozyaystvennogo osvoyeniya territorii v izmenenii chislennosti kalmytskoy populyatsii saygaka] Biota and natural environment of Kalmykia [Biota i prirodnaya sreda Kalmykii]. Moscow-Elista, 1995:222-244.
7. Bliznyuk AI. Saiga of the Kalmyk population [Saygak kalmytskoy populyatsii]. Elista: ZAO "NPP 'Dzhangar'", 2009:554.
8. Bogun SA. The state of the saiga population in the Chernye Zemli Nature Reserve: problems and prospects for its conservation [Sostoyaniye populyatsii saygaka v zapovednike «Chernyye zemli»: problemy i perspektivy yeye sokhraneniya] Scientific Works of the Khvalynsky National Park [Nauchnyye trudy Natsional'nogo parka «Khvalynskiy»] Collection of scientific articles of the VI International Scientific and Practical Conference "Specially Protected Natural Territories: Past, Present, Future", Khvalynsk, October 17-18, 2019 [Sbornik nauchnykh statey VI Mezhdunarodnoy nauchno-prakticheskoy konferentsii "Osobo okhranyayemyye prirodnyye territorii: proshloye, nastoyashcheye, budushcheye"]. Saratov: Amirit,
Калмыкии // Зоологический журнал. Т. 87. № 12. С. 1524-1530.
3. Банников А.Г., Жирное Л.В., Лебедева Л.С., Фандеев А.А. 1961. Биология сайгака. М.: Изд-во сельскохозяйственной литературы, журналов и плакатов. 336 с.
4. Барышников Г.Ф., Дмитриева Е.Л., Крахмальная Т.В., Шер А.В. 1998. Происхождение, эволюция и систематика сайгака // Сайгак: филогения, систематика, экология, охрана и использование. М.: Типография Россельхозакадемии. С. 9-20.
5. Бекенов А.Б., Грачев Ю.А. 1998. Численность сайгаков в ХХ веке. Казахстан // Сайгак: филогения, систематика, экология, охрана и использование. М.: Типография Россельхозакадемии. С. 225-227.
6. Близнюк А.И. 1995. Роль хозяйственного освоения территории в изменении численности калмыцкой популяции сайгака // Биота и природная среда Калмыкии. Москва-Элиста. С. 222-244.
7. Близнюк А.И. 2009. Сайгак калмыцкой популяции. Элиста: ЗАО «НПП «Джангар». 554 с.
8. Богун С.А. 2019. Состояние популяции сайгака в заповеднике «Черные земли»: проблемы и перспективы ее сохранения //
Научные труды Национального парка «Хвалынский». Сборник научных статей VI Международной научно-практической конференции «Особо охраняемые природные территории: прошлое, настоящее, будущее». Хвалынск, 1718 октября 2019 г. Саратов: Амирит. С. 714.
9. Брагина Т.М. 2015. Сайгачьи переходы в Казахстане взяты под охрану // Saiga News. № 19. С. 3-4.
10. Грачев Ю.А., Мелдебеков А.М., Бекенов А.Б. 2009. Численность, структура и воспроизводство популяций сайгака в Казахстане // Степной бюллетень. № 27. С. 47-50.
11. Данилкин А.А. 2005. Млекопитающие России и сопредельных территорий. Полорогие. М.: Товарищество научных изданий КМК. 550 с.
12. Динесман Л.Г. 1998. История ареала сайгака в послеледниковое время //
2019:7-14.
9. Bragina TM. Saiga crossings in Kazakhstan are taken under protection [Saygach'i perekhody v Kazakhstane vzyaty pod okhranu] Saiga News. 2015;19:3-4.
10. Grachev YuA, Meldebekov AM, Bekenov AB. Number, structure and reproduction of saiga populations in Kazakhstan [Chislennost', struktura i vosproizvodstvo populyatsiy saygaka v Kazakhstane] Steppe Bulletin [Stepnoy byulleten']. 2009;27:47-50.
11. Danilkin AA. Mammals of Russia and neighboring territories [Mlekopitayushchiye Rossii i sopredel'nykh territoriy] Bovids [Polorogiye]. Moscow: Tovarishchestvo nauchnykh izdaniy KMK, 2005:550.
12. Dinesman LG. History of the saiga range in the post-glacial period [Istoriya areala saygaka v poslelednikovoye vremya] Saiga: phylogeny, taxonomy, ecology, protection and use [Saygak: filogeniya, sistematika, ekologiya, okhrana i ispol'zovaniye]. Moscow: Tipografiya Rossel'khozakademii, 1998:54-57.
13. Zhirnov LV. Brought back to life (ecology, conservation and use of saigas) [Vozvrashchennyye k zhizni (ekologiya, okhrana i ispol'zovaniye saygakov)]. Moscow: Lesnaya promyshlennost', 1982:224.
14. Zhirnov LV, Bekenov AB, Grachev YuA, Dulamtseren S, Lushchekina AA. The area and its change in the 20th century [Areal i yego izmeneniye v XX v. ] Saiga: phylogeny, taxonomy, ecology, protection and us [Saygak: filogeniya, sistematika, ekologiya, okhrana i ispol'zovaniye]. Moscow: Tipografiya Rossel'khozakademii, 1998a:60-66.
15. Zhirnov LV, Dulamtseren S, Lushchekina AA. Population status in China and Mongolia
[Sostoyaniye chislennosti populyatsiy v Kitaye i Mongolii] Saiga: phylogeny, taxonomy, ecology, protection and use [Saygak: filogeniya, sistematika, ekologiya, okhrana i ispol'zovaniye]. Moscow: Tipografiya Rossel'khozakademii, 1998b:227-229.
16. Zhirnov LV, Maksimuk AV. The number of saigas in the 20th century. Northwest Pre-Caspian [Chislennost' saygakov v XX veke. Severo-Zapadnyy Prikaspiy] Saiga: phylogeny, taxonomy, ecology, protection and use [Saygak: filogeniya, sistematika, ekologiya, okhrana i ispol'zovaniye]. Moscow: Tipografiya Rossel'khozakademii, 1998:219-224.
17. Karimova TYu, Lushchekina AA. Features of the
Сайгак: филогения, систематика, экология, охрана и использование. М.: Типография Россельхозакадемии. С. 54-57.
13. Жирное Л.В. 1982. Возвращенные к жизни (экология, охрана и использование сайгаков). М.: Лесная промышленность. 224 с.
14. Жирное Л.В., Бекеное А.Б., Грачев Ю.А., Дуламцэрэн С., Лущекина А.А. 1998а. Ареал и его изменение в ХХ в. // Сайгак: филогения, систематика, экология, охрана и использование. М.: Типография Россельхозакадемии. С. 60-66.
15. Жирное Л.В., Дуламцэрэн С., Лущекина А.А. 19986. Состояние численности популяций в Китае и Монголии // Сайгак: филогения, систематика, экология, охрана и использование. М.: Типография Россельхозакадемии. С. 227-229.
16. Жирное Л.В., Максимук А.В. 1998. Численность сайгаков в ХХ веке. СевероЗападный Прикаспий // Сайгак: филогения, систематика, экология, охрана и использование. М.: Типография Россельхозакадемии. С. 219-224.
17. Каримоеа Т.Ю., Лущекина А.А. 2018. Особенности пространственного размещения и этологической структуры популяции сайгака на территории заказника «Степной» (Астраханская область) // Экосистемы: экология и динамика. Т. 2. № 1. С. 73-91. [Электронный ресурс http://www.ecosystemsdynamic.ru
(дата обращения 22.09.2022)].
18. Каримоеа Т.Ю., Лущекина А.А., Рожное В.В. 2017. Сайгаки в неволе: от содержания и разведения до выпуска в природу. Москва: Т-во научных изданий КМК. 122 с. [Karimova T.Yu., Lushchekina A.A., Rozhnov V.V. 2018. Saiga management at zoos and breeding centres: making effective use of the lessons learnt for the restoration of wild saiga populations. M.: KMK Scientific Press Ltd. 119 p.].
19. Кирикое С.В. 1966. Промысловые животные, природная среда и человек. М.: Наука. 348 с.
20. Красная книга Астраханской области. 2014. Астрахань: Издательский дом «Астраханский университет». 413 с.
spatial distribution and ethological structure of the saiga population on the territory of the Stepnoy sanctuary (Astrakhan region) [Osobennosti prostranstvennogo razmeshcheniya i etologicheskoy struktury populyatsii saygaka na territorii zakaznika «Stepnoy» (Astrakhanskaya oblast')] Ecosystems: Ecology and Dynamics. 2018;2 (1):73-91, Available at
http://www.ecosystemsdynamic.ru (Date of Access 22/09/2022).
18. Karimova TYu, Lushchekina AA, Rozhnov VV. Saiga management at zoos and breeding centres: making effective use of the lessons learned for the restoration of wild saiga populations. Moscow: T-vo nauchnykh izdaniy KMK, 2018;119.
19. Kirikov SV. Game animals, natural environment and man [Promyslovyye zhivotnyye, prirodnaya sreda i chelovek]. Moscow: Nauka, 1966:348.
20. Red Data Book of the Astrakhan region [Krasnaya kniga Astrakhanskoy oblasti]. Astrakhan: Izdatel'skiy dom "Astrakhanskiy universitet", 2014:413.
21. Red Book of the Republic of Kalmykia [Krasnaya kniga Respubliki Kalmykiya] Animals [Zhivotnyye]. Elista: ZAO "NPP 'Dzhangar'", 2013;1:200.
22. Red Data Book of the Russian Federation
23.
24.
AA. Phylogenetic analysis of the 12S and 16S rRNA mitochondrial gene sequences of representatives of the Bovidae family: new data [Filogeneticheskiy analiz posledovatel'nostey mitokhondrial'nykh genov 12S i 16S rRNK predstaviteley semeystva Bovidae: novyye dannyye]. Genetics. 2002;38 (3):942-950.
25. Levykin SV, Kazachkov GV, Yakovlev IG, Grudinin DA. Saiga in the Orenburg region: history, legends, prospects for return [Saygak v Orenburzh'ye: istoriya, legendy, perspektivy vozvrashcheniya] Proc. of the Samara Scientific Center of the Russian Academy of Sciences [Izvestiya Samarskogo nauchnogo tsentra RAN]. 2015;17 (4):174-178.
26. Chronicle of Nature of the Federal State Budgetary Institution of the State Natural Biosphere Reserve
21. Красная книга Республики Калмыкия. Т. 1: Животные. 2013. Элиста: ЗАО «НПП «Джангар». 200 с.
22. Красная книга Российской Федерации. Животные. 2021. М.: ФГБУ «ВНИИ Экология». 1128 с.
23. Красная книга Ростовской области. Т. 1: Животные. 2014. Ростов-на-Дону: Минприроды Ростовской области. 280 с.
24. КузнецоваМ.В., ХолодоваМ.В., Лущекина А.А. 2002. Филогенетический анализ последовательностей митохондриальных генов 12S и 16S рРНК представителей семейства Bovidae: новые данные // Генетика. Т. 38. № 3. С. 942-950.
25. Левыкин С.В., Казачков Г.В., Яковлев И.Г., Грудинин Д.А. 2015. Сайгак в Оренбуржье: история, легенды, перспективы возвращения // Известия Самарского научного центра РАН. Т. 17. № 4. С. 174-178.
26. Летопись природы ФГБУ государственного природного биосферного заповедника «Черные земли». Кн. XVIII. 2014. [Электронный ресурс http://zapovednik-chernyezemli. ru/wp -
content/uploads/2015/11/Летопись_природ ы_ 2014_год.pdf (дата обращения 22.09.2022)].
27. Мельников В.В., Сидоров С.В. 2009. Сайгак в России: современное состояние, сохранение и восстановление // Степной бюллетень. № 27. С. 42-46.
28. Милнер-Гулланд Э.Дж., Хагес П., Быкова E., Буувейбаатар Б., Чимеддорж Б., Каримова Т., Лущекина А., Салемгареев А., Мейбом С., Цутер Ш. 2020. Устойчивое использование антилопы сайги: обзор и перспективы. Предварительный отчет для Конвенции ООН по сохранению мигрирующих видов диких животных и Федерального агентства по охране природы Германии. 129 с. [Электронный ресурс https://www.cms. int/sites/default/files/docume nt/unep-
cms_saiga_mos4_doc.7_rev.1_sustainable-use-saiga-antelopes_ru.pdf (дата обращения 22.09.2022)]. [Milner-GullandE.J., Hughes P., Bykova E., Buuveibaatar B.,
Chimeddorj B., Karimova T. Yu.,
Lushchekina A.A., Salemgareyev A.,
[Krasnaya kniga Rossiyskoy Federatsii] Animals
[Zhivotnyye]. Moscow: FGBU
"VNII Ekologiya", 2021: 1128.
Red Data Book of the Rostov Region [Krasnaya
kniga Rostovskoy oblasti] Animals [Zhivotnyye].
Rostov-on-Don: Minprirody Rostovskoy oblasti,
2014;1:280.
Kuznetsova MV, Kholodova MV, Lushchekina
"Chernye Zemli", Book XVIII [Letopis' prirody FGBU gosudarstvennogo prirodnogo biosfernogo zapovednika "Chernyye zemli"]. 2014, Available at http://zapovednik-chernyezemli.ru/wp-content/uploads/2015/11/neTOnHCb_npHpogbi_ 2014_rog.pdf (Date of Access 22/09/2022).
27. Melnikov VV, Sidorov SV. Saiga in Russia: Current State, Conservation and Restoration [Saygak v Rossii: sovremennoye sostoyaniye, sokhraneniye i vosstanovleniye] Steppe Bulletin. 2009;27:42-46.
28. Milner-Gulland EJ, Hughes P, Bykova E, Buuveibaatar B, Chimeddorj B, Karimova TYu, Lushchekina AA, Salemgareyev A, von Meibom S, Zuther S . The Sustainable Use of Saiga Antelopes: Perspectives and Prospects. The Bundesamt fur Naturschutz and the Convention on the Conservation of Migratory Species of Wild Animals, 2021:117.
29. Minoransky VA, Dankov VI. Saiga antelope (Saiga tatarica L.) is an endangered species in Russia [Saygak (Saiga tatarica L.) -ischezayushchiy v Rossii vid] South of Russia: ecology, development (Series "Animal Ecology") [Yug Rossii: ekologiya, razvitiye (seriya "Ekologiya zhivotnykh")]. 2016;11 (1):88-103.
30. Minoransky VA, Uzdenov AM, Dankov VI, Malinovskaya YuV. Problems of conservation of the saiga (Saiga tatarica L.) in Russia and breeding it in artificial conditions [Problemy sokhraneniya saygaka (Saiga tatarica L.) v Rossii i razvedeniya yego v iskusstvennykh usloviyakh] Biological diversity of Asian steppes [Biologicheskoye raznoobraziye aziatskikh stepey] Proc. of the IV International Scientific Conference, April 14, 2022, Kostanay, Kazakhstan [Materialy IV mezhdunarodnoy nauchnoy konferentsii]. Kostanay: KRU im. A. Baytursynova, 2022:32-36.
31. On approval of the list of specially protected natural areas of republican significance [Ob utverzhdenii perechnya osobo okhranyayemykh prirodnykh territoriy respublikanskogo znacheniya] Decree of the Government of the Republic of Kazakhstan No. 593, issued on 26/09/2017 [Postanovleniye Pravitel'stva Respubliki Kazakhstan]. 2022, Available at https://adilet.zan.kz/rus/docs/P1700000593 (Date of Access 22/09/2022).
32. Paltsyn MYu. Assessment of the potential suitability of habitats for steppe ungulates in the Republic of Kalmykia and the Trans-Baikal Territory [Otsenka potentsial'noy prigodnosti mestoobitaniy stepnykh kopytnykh v Respublike
von Meibom S., Zuther S. 2021. The Sustainable Use of Saiga Antelopes: Perspectives and Prospects. The Bundesamt fur Naturschutz and the Convention on the Conservation of Migratory Species of Wild Animals. 117 p.].
29. Миноранский В.А., Даньков В.И. 2016. Сайгак (Saiga tatarica L.) - исчезающий в России вид // Юг России: экология, развитие (серия «Экология животных»). Т. 11. № 1. С. 88-103.
30. Миноранский В.А., Узденов А.М., Даньков В.И., Малиновская Ю.В. 2022. Проблемы сохранения сайгака (Saiga tatarica L.) в России и разведения его в искусственных условиях // Биологическое разнообразие азиатских степей. Материалы IV международной научной конференции, 14 апреля 2022 г., Костанай, Казахстан. Костанай: КРУ им. А. Байтурсынова. С. 3236.
31. Об утверждении перечня особо охраняемых природных территорий республиканского значения. Постановление Правительства Республики Казахстан от 26 сентября 2017 года. № 593. 2022. [Электронный ресурс
https://adilet.zan.kz/rus/docs/P1700000593 (дата обращения 22.09.2022)].
32. Пальцын М.Ю. 2016. Оценка потенциальной пригодности местообитаний степных копытных в Республике Калмыкия и Забайкальском крае. Отчет по проекту ПРООН/ГЭФ 00072294 «Совершенствование системы и механизмов управления ООПТ в степном биоме России» [Электронный ресурс http://savesteppe.org/project/docs/report_final _habitat-model_ungulates2016.pdf (дата обращения 22.09.2022)].
33. Поголовье скота на 1 декабря 2019 г. 2019 [Электронный ресурс https://astrastat.gks.m/storage/mediabank/П оголовье_скота_на_1_декабря_2019.pdf (дата обращения 22.09.2022)].
34. Распоряжение Минприроды России от 11.08.2021 N 30-Р «Об Утверждении Стратегии Сохранения сайгака в Российской Федерации». 2021. [Электронный ресурс https://rulaws.ru/acts/Rasporyazhenie-Minprirody-Rossii-ot-11.08.2021 -N-30-r/
Kalmykiya i Zabaykal'skom kraye] Report on the UNDP/GEF project No. 00072294 "Improving the system and management mechanisms for protected areas in the steppe biome of Russia" [Otchet po proyektu PROON/GEF 00072294 "Sovershenstvovaniye sistemy i mekhanizmov upravleniya OOPT v stepnom biome Rossii"]. 2016, Available at
http://savesteppe.org/project/docs/report_final_ha bitat-model_ungulates2016.pdf (Date of Access 22/09/2022).
33. Livestock as of 01/12/2019 [Pogolov'ye skota na 1 dekabrya 2019 g. ]. 2019, Available at https://astrastat.gks.ru/storage/mediabank/noron OBbe_CKOTa_Ha_1_geKa6pa_2019.pdf (Date of Access 22/09/2022).
34. Order of the Ministry of Natural Resources of Russia No. 30-R, issued on 08/11/2021 [Rasporyazheniye Minprirody Rossii] "On Approval of the Saiga Conservation Strategy in the Russian Federation" ["Ob Utverzhdenii Strategii Sokhraneniya saygaka v Rossiyskoy Federatsii"]. 2021, Available at https://rulaws.ru/acts/Rasporyazhenie-Minprirody-Rossii-ot-11.08.2021 -N-30-r/ (Date of Access 22/09/2022).
35. Rozhnov VV, Yachmennikova AA, Dobrynin DV. On the possibility of identifying the saiga (Saiga tatarica) on high-resolution satellite images [O vozmozhnosti vyyavleniya saygaka (Saiga tatarica) na sputnikovykh snimkakh vysokogo razresheniya] Reports of the Academy of Sciences [Doklady Akademii nauk]. 2014;459 (6):769-773.
36. Saigas destroy the crops of Saratov farmers [Saygaki unichtozhayut posevy saratovskikh fermerov], 2022, Available at https://gtrk-saratov.ru/sajgaki-unichtozhayut-posevy-saratovskih-fermerov/ (Date of Access 22/09/2022).
37. Counting rhyme about saiga [Saygach'ya schitalochka]. 2022, Available at https://zapovednik-chemyezemli.ru/6e3-py6pHKH/caHranba-CHHTanoHKa/ (Date of Access 22/09/2022).
38. Sidorov SV, Bukreeva OM. Population cycles of the saiga in the Northwest Pre-Caspian
[Populyatsionnyye tsikly saygaka Severo-Zapadnogo Prikaspiya] VI Congress of the Theriological Society [VI S'yezd
teriologicheskogo obshchestva]. Moscow: Izdatel'stvo Rossel'khozakademii, 1999:232.
39. State of the world's biodiversity for food and
(дата обращения 22.09.2022)].
35. Рожнов В.В., Ячменникова А.А., Добрынин Д.В. 2014. О возможности выявления сайгака (Saiga tatarica) на спутниковых снимках высокого разрешения // Доклады Академии наук. Т. 459. № 6. С. 769-773.
36. Сайгаки уничтожают посевы саратовских фермеров. 2022. [Электронный ресурс https://gtrk-saratov.ru/sajgaki-unichtozhayut-posevy-saratovskih-fermerov/ (дата обращения 22.09.2022)].
37. Сайгачья считалочка [Электронный ресурс https://zapovednik-chernyezemli.ru/без-рубрики/сайгачья-считалочка/ (дата обращения 22.09.2022)].
38. Сидоров С.В., Букреева О.М. 1999. Популяционные циклы сайгака СевероЗападного Прикаспия // VI Съезд териологического общества. М.: Изд-во Россельхозакадемии. С. 232.
39. Состояние биоразнообразия в мире для производства продовольствия и ведения сельского хозяйства. Краткий обзор. 2019. Комиссия по генетическим ресурсам для производства продовольствия и ведения сельского хозяйства ФАО. 16 с. [Электронный ресурс
https://www.fao.org/3/CA3229RU/ca3229ru .pdf (дата обращения 22.09.2022)].
40. Судьба сайгаков в Казахстане: что говорят экологи, ученые и о чем заявляют фермеры. 2022. [Электронный ресурс https://www.inform.kz/ru/sud-ba-saygakov-v-kazahstane-chto-govoryat-ekologi-uchenye-i-o-chem-zayavlyayut-fermery_a3952746 (дата обращения 22.09.2022)].
41. Уголовный кодекс Российской Федерации. 2017. [Электронный ресурс http://stykrf.ru (дата обращения 22.09.2022)].
42. Уголовный кодекс Республики Казахстан от 3 июля 2014 года № 226-V (с изменениями и дополнениями по состоянию на 12.09.2022 г.). 2022. [Электронный ресурс https://online.zakon.kz/document/?doc_id=3 1575252&pos=363;-48#pos=363;-48 (дата обращения 22.09.2022)].
43. Учет сайгаков Северо-Западного Прикаспия будут вести с помощью
agriculture [Sostoyaniye bioraznoobraziya v mire dlya proizvodstva prodovol'stviya i vedeniya sel'skogo khozyaystva] Short Review [Kratkiy obzor]. FAO Commission on Genetic Resources for Food and Agriculture, 2019:16, Available at https://www.fao.org/3ZCA3229RU/ca3229ru 44. (Date of Access 22/09/2022).
40. The fate of saigas in Kazakhstan: what ecologists, scientists say and what farmers say [Sud'ba saygakov v Kazakhstane: chto govoryat ekologi, uchenyye i o chem zayavlyayut fermery]. 2022, Available at https://www.inform.kz/ru/sud-ba- 45. saygakov-v-kazahstane-chto-govoryat-ekologi-uchenye-i-o-chem-zayavlyayut-fermery_a3952746 (Date of Access 22/09/2022).
41. The Criminal Code of the Russian Federation 46. [Ugolovnyy kodeks Rossiyskoy Federatsii]. 2017, Available at http://stykrf.ru (Date of Access 22/09/2022).
42. The Criminal Code of the Republic of 47. Kazakhstan No. 226-V as of 03/06/2014 (with changes and additions as of 09/12/2022) [Ugolovnyy kodeks Respubliki Kazakhstan (s 48. izmeneniyami i dopolneniyami)]. 2022, Available at
https://online.zakon.kz/document/?doc_id=31575 252&pos=363;-48#pos=363;-48 (Date of Access 22/09/2022).
43. Saigas census in the Northwest Pre-Caspian 49. region will be carried out using drones [Uchet saygakov Severo-Zapadnogo Prikaspiya budut vesti s pomoshch'yu bespilotnikov]. 2022, Available at https://tass.ru/obschestvo/14551515?utm_source 50. =google. com&utm_medium=organic&utm_camp aign=google. com&utm_referrer=google. com
(Date of Access 22/09/2022).
44. Chimeddorj B, Sergelen E, Buweibatar B. The impact of anthropogenic factors on the distribution and movement of saigas in Western Mongolia [Vliyaniye antropogennykh faktorov na rasprostraneniye i peremeshcheniye saygakov v Zapadnoy Mongolii] Saiga News. 2016;21:35-37. 51.
45. Sher AV. Fossil saiga in the north of Eastern Siberia and Alaska [Iskopayemaya sayga na severe Vostochnoy Sibiri i Alyaski] Bulletin of the commission for the study of the Quaternary period [Byulleten' komisssii po izucheniyu chetvertichnogo perioda]. 1967;33:97-112.
46. Baryshnikov GF, Tikhonov AN. Notes on skulls 52. of Pleistocene Saiga of Northern Eurasia. Historical Biology. 1994;4:209-234.
47. Briggs JC. Emergence of a sixth mass extinction?
беспилотников. 2022. [Электронный ресурс https://tass.ru/obschestvo/145515157utm_sour ce=google.com&utm_medium=organic&utm_ campaign=google.com&utm_referrer=google. com (дата обращения 22.09.2022)]. Чимеддорж Б., Сергелен Э.,
Бувейбатар Б. 2016. Влияние
антропогенных факторов на
распространение и перемещение сайгаков в Западной Монголии // Saiga News. № 21. С. 35-37. Шер А.В. 1967. Ископаемая сайга на севере Восточной Сибири и Аляски // Бюллетень комисссии по изучению четвертичного периода. № 33. С. 97-112. Baryshnikov G.F., Tikhonov A.N. 1994. Notes on skulls of Pleistocene Saiga of Northern Eurasia // Historical Biology. Vol. 4. P. 209-234.
Briggs J.C. 2017. Emergence of a sixth mass extinction? // Biological Journal of the Linnean Society. Vol. 122. Is. 2. P. 243-248. Buuveibaatar B., Fuller T.K., Fine A.E., Chimeddorj B., Young J.K., Berger J. 2013. Changes in grouping patterns of saiga in relation to intrinsic and environmental factors in Mongolia // Journal of Zoology. Vol. 29. Р. 51-58.
ClarkE.L., Javzansuren M., Dulamtseren J., Baillie E.M., Batsaikhan N., Samiya R., Stubbe M. 2006. Mongolian Red List of Mammals. London: Zoological Society of London. P. 6466.
Ceballosa G., Ehrlichb P.R., Dirzob R. 2017. Biological annihilation via the ongoing sixth mass extinction signaled by vertebrate population losses and declines // Proceedings of the National Academy of Sciences. No. 114 (30). P. Е6089-Е6096. [Электронный ресурс
https://www.pnas.org/doi/full/10.1073/pnas.17 04949114 (дата обращения 22.09.2022)]. Fewer than 3800 Mongolian Saiga antelopes remain in Mongolia: WWF. 2019. [Электронный ресурс
https://www.wwf.mg/7344814/Fewer-than-3800-Mongolian-Saiga-antelopes-remain-in-Mongolia-WWF (дата обращения
22.09.2022)].
Gao X., Xu W., Yang W., Blank D., Qiao J.F., Xu K.F. 2011. Status and distribution of ungulates in Xinjiang, China // Journal of Arid Land. Vol. 3. No. 1. P. 49-
Biological Journal of the Linnean Society. 2017;122 (2):243-248.
48. Buuveibaatar B, Fuller TK, Fine AE, Chimeddorj B, Young JK, Berger J. Changes in grouping patterns of saiga in relation to intrinsic and environmental factors in Mongolia. Journal of Zoology. 2013;29:51-58.
49. Clark EL, Javzansuren M, Dulamtseren J, Baillie EM, Batsaikhan N, Samiya R, Stubbe M. Mongolian Red List of Mammals. London: Zoological Society of London, 2006:64-66.
50. Ceballosa G, Ehrlichb PR, Dirzob R. Biological annihilation via the ongoing sixth mass extinction signaled by vertebrate population losses and declines. Proc. of the National Academy of Sciences. 2017;114 (30):E6089-E6096, Available at
https://www.pnas.org/doi/full/10.1073/pnas.1704 949114 (Date of Access 22/09/2022).
51. Fewer than 3800 Mongolian Saiga antelopes remain in Mongolia: WWF. 2019, Available at https://www.wwf.mg/?344814/Fewer-than-3800-Mongolian-Saiga-antelopes-remain-in-Mongolia-WWF (Date of Access 22/09/2022).
52. Gao X, Xu W, Yang W, Blank D, Qiao JF, Xu KF. Status and distribution of ungulates in Xinjiang, China. Journal of Arid Land. 2011;3 (1);49-60.
53. Karimova TYu, Lushchekina AA, Neronov VM. Saiga Populations of Russia and Kazakhstan: Current Status and Retrospective Analysis of Some Biological Parameters. Arid Ecosystems. 2021;11:164-172.
54. Karimova TYu, Lushchekina AA, Neronov VM, Pyurvenova NYu, Arylov YuN. Biological features of the Northwest Pre-Caspian Saiga population at different sizes. Arid Ecosystems. 2020;10 (4):298-304.
55. Kuhl A, Balinova N, Bykova E, Arylov Yu, Esipov A, Lushchekina A, Milner-Gulland EJ. The role of saiga poaching in rural communities: Linkages between attitudes, socio-economic circum-stances and behavior. Biological Conservation. 2009;143:1442-1449.
56. Lushchekina AA, Dulamtseren S, Amgalan L, Neronov VM. The status and prospects for conservation of the Mongolian saiga Saiga tatarica mongolica. Oryx. 1999;33:21-30.
57. Mammal Species of the World. A Taxonomic and Geographic Reference / eds. D.E. Wilson, D.M. Reeder. Baltimore: Johns Hopkins University Press, 2005:2142.
58. Milner-Gulland EJ, Bukreeva OM, Coulson T,
60.
53. Karimova T.Yu., Lushchekina A.A., Neronov V.M. 2021. Saiga Populations of Russia and Kazakhstan: Current Status and Retrospective Analysis of Some Biological Parameters // Arid Ecosystems. Vol. 11. P. 164-172 [Каримова Т.Ю., Лущекина А.А., Неронов В.М. 2021. Современное состояние и ретроспективный анализ популяций сайгака России и Казахстана // Аридные экосистемы. Т. 27. № 2 (87). С. 57-67.].
54. Karimova T.Yu., Lushchekina A.A., Neronov V.M., Pyurvenova N.Yu., Arylov Yu.N. 2020. Biological features of the Northwest Pre-Caspian Saiga population at different sizes // Arid Ecosystems. Vol. 10. No. 4. P. 298-304 [Каримова Т.Ю., Лущекина А.А., Неронов В.М., Пюрвенова Н.Ю., Арылов Ю.Н. 2020. Биологические особенности популяции сайгака Северо-Западного Прикаспия в периоды разной численности // Аридные экосистемы. T. 26. № 4 (85). C. 51-58.].
55. Kuhl A., Balinova N., Bykova E., Arylov Yu., Esipov A., Lushchekina A., Milner-Gulland E.J. 2009. The role of saiga poaching in rural communities: Linkages between attitudes, socio-economic circum-stances and behavior // Biological Conservation. Vol. 143. P. 1442-1449.
56. Lushchekina A.A., Dulamtseren S., Amgalan L., Neronov V.M. 1999. The status and prospects for conservation of the Mongolian saiga Saiga tatarica mongolica // Oryx. Vol. 33. P. 21-30.
57. Mammal Species of the World. A Taxonomic and Geographic Reference. 2005 / Eds. D.E. Wilson, D.M. Reeder. Baltimore: Johns Hopkins University Press. 2142 p.
58. Milner-Gulland E.J., Bukreeva O.M., Coulson T., Lushchekina A.A., Kholodova M.V., Bekenov A.B., Grachev I.A. 2003. Reproductive collapse in saiga antelope harems // Nature. Vol. 422. P. 135.
59. Mongolian saiga population hits 10,077. 2021 [Электронный ресурс https://m.akipress.com/news:664939:Mongo lian_saiga_population_hits_10,077 (дата обращения 22.09.2022)].
60. Neronov V.M., ArylovaN.Yu., Dubinin M. Yu., Karimova T. Yu., Lushchekina A.A. 2013. Current state and
Lushchekina AA, Kholodova MV, Bekenov AB, Grachev IA. Reproductive collapse in saiga antelope harems. Nature. 2003;422:135.
59. Mongolian saiga population hits 10,077. 2021, Available at
https://m.akipress.com/news:664939:Mongolian_ saiga_population_hits_10,077 (Date of Access 22/09/2022).
60. Neronov VM, Arylova NYu, Dubinin MYu, Karimova TYu, Lushchekina AA. Current state and prospects of preserving saiga antelope in Northwest Pre-Caspian region. Arid Ecosystems. 2013;3 (2):57-64.
61. The global assessment report on biodiversity and ecosystem services. Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (IPBES). 2019:60, Available at https://ipbes.net/sites/default/files/inline/files/ipbes_ global_assessment_report_summary_for_policyma kers.pdf (Date of Access 22/09/2022).
prospects of preserving saiga antelope in Northwest Pre-Caspian region // Arid Ecosystems. Vol. 3. P. 57-64 [Неронов В.М., Арылова Н.Ю., Дубинин М.Ю., Каримова Т.Ю., Лущекина А.А. 2013. Современное состояние и перспективы сохранения сайгака в Северо-Западном Прикаспии // Аридные экосистемы. Т. 19. № 2 (55). С. 5-14.].
61. The global assessment report on biodiversity and ecosystem services. 2019. Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (IPBES). 60 p. [Электронный ресурс https ://ipbes.net/sites/default/files/inline/file s/ipbes_global_assessment_report_summary _for_policymakers.pdf (дата обращения 22.09.2022)].
УДК 599.735.53:591.5
ПРОШЛОЕ И НАСТОЯЩЕЕ САЙГАКА В РОССИИ - ЕСТЬ ЛИ БУДУЩЕЕ?
© 2022 г. Т.Ю. Каримова*' **, А.А. Лущекина*, В.М. Неронов*, Ю.Н. Арылов***, Н.Ю. Пюрвенова****
* Институт проблем экологии и эволюции им. А.Н. Северцова РАН Россия, 119071, г. Москва, Ленинский просп., д. 33. E-mail: rusmabcom@gmail.com
**Институт водных проблем РАН Россия, 119333, г. Москва, ул. Губкина, д. 3. E-mail: katayur@gmail.com ***Калмыцкий государственный университет им. Б.Б. Городовикова Россия, 358000, г. Элиста, ул. Пушкина, д. 11. E-mail: kalmsaiga@mail.ru ****Независимый исследователь Россия, 358001, г. Элиста, пер. Гермашева, д. 54а. E-mail: arylova@gmail.com
Поступила в редакцию 01.11.2022. После доработки 10.11.2022. Принята к публикации 01.12.2022.
Выработанные в ходе эволюции жизненные стратегии сайгака (раннее вступление в размножение и высокая плодовитость самок, полигамия, стадность, миграции и др.) позволили виду сохраниться со времен мамонтовой фауны в изменяющихся условиях среды. В голоцене начался новый этап в истории сайгака - сосуществование с человеком. Помимо истребления этих животных в результате массовой охоты, стало расти влияние человека на среду обитания животных за счет широкого распространения земледелия, особенно в Западной Европе, что отразилось на сокращении ареала вида в исторический период. К началу ХХ века от сплошного и обширного ареала сайгака остались лишь отдельные пятна, и в Европе он встречался только в нетронутых человеком глухих уголках нижнего течения реки Волги, а в Азии - на Устюрте, в Бетпак-Дале, в междуречье Или-Каратал, Китае и Монголии. Благодаря предпринятым в 1920-е гг. советским правительством охранным мерам, в настоящее время в мире существует пять популяций сайгака, из которых четыре популяции - Северо-Западного Прикаспия (Россия), Уральская (Казахстан, Россия), Устюртская (Казахстан, Узбекистан, Туркменистан),
Бетпакдалинская (Казахстан, Россия) - относятся к номинативному подвиду £ Шапса, а пятая популяция - 5. (. тощоИса (=5. ЪвгеаИъ) - обитает в Западной Монголии. С конца прошлого века состояние популяции сайгака Северо-Западного Прикаспия вызывает особую озабоченность. Наряду со снижением численности сайгака (с 800 тыс. в 1958 г. до 5 тыс. в 2015 г.) сократилась и площадь зоны его основного обитания (с 60-70 тыс. км2 до 2-3 тыс. км2, соответственно). Значительная часть популяции перешла к оседлому образу жизни на охраняемых территориях экорегиона «Черные земли». Нехватка половозрелых самцов в популяции (менее 10 %) на протяжении продолжительного времени привела к сокращению ее прироста и, следовательно, к снижению численности. Благодаря предпринятым на разных уровнях мерам с 2016 г. популяция сайгака в Северо-Западном Прикаспии начала постепенно увеличиваться (до 18-19 тыс. в 2022 г.). Дальнейший рост ее численности в настоящее время зависит от эффективности действий, которые предполагается осуществить в рамках реализации «Стратегии сохранения сайгака в Российской Федерации» - кроме организации эффективной охраны и полномасштабного мониторинга, это и недопущение фрагментации мест обитания сайгака за счет снятия разного рода препятствий для кочевок, улучшение качества местообитаний, создание новых охраняемых территорий разного уровня в наиболее подходящих местообитаниях и объединение их путем создания экологических коридоров в единую сеть, расширение и активизация эколого-просветительской деятельности. Ключевые слова: сайгак, популяции сайгака, численность, ареал, Северо-Западный Прикаспий, сохранение сайгака.
Благодарности. В заключении авторы выражают искреннюю благодарность сотрудникам заказника «Степной» и его директору В.Г. Калмыкову за многолетнее плодотворное сотрудничество, а также А.Н. Гилеву за предоставленные фотографии, сделанные в заказнике «Степной».
Финансирование. Работа выполнена по теме НИР Института проблем экологии и эволюции им. А.Н. Северцова РАН, Госзадание № 1021062812203-8 «Фундаментальные проблемы охраны живой природы и рационального использования биоресурсов», а также по теме НИР фундаментальных исследований ИВП РАН за 2022-2024 гг. «Исследования геоэкологических процессов в гидрологических системах суши, формирования качества поверхностных и подземных вод, проблем управления водными ресурсами и водопользованием в условиях изменений климата и антропогенных воздействий» (№ FMWZ -2022-0002), № государственной регистрации АААА-А18-118022090104-8. DOI: 10.24412/2542-2006-2022-4-28-49 EDN: QVMVAB
