<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD JATS (Z39.96) Journal Publishing DTD v1.3 20210610//EN" "JATS-journalpublishing1-3.dtd">
<article article-type="research-article" dtd-version="1.3" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xml:lang="ru"><front><journal-meta><journal-id journal-id-type="publisher-id">sevin</journal-id><journal-title-group><journal-title xml:lang="ru">ПОВОЛЖСКИЙ ЭКОЛОГИЧЕСКИЙ ЖУРНАЛ</journal-title><trans-title-group xml:lang="en"><trans-title>Povolzhskiy Journal of Ecology</trans-title></trans-title-group></journal-title-group><issn pub-type="ppub">1684-7318</issn><issn pub-type="epub">2541-8963</issn><publisher><publisher-name>Федеральное государственное бюджетное учреждение науки Институт проблем экологии и эволюции им. А. Н. Северцова РАН; Товарищество научных изданий КМК.</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="doi">10.35885/1684-7318-2025-1-31-47</article-id><article-id custom-type="elpub" pub-id-type="custom">sevin-627</article-id><article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject></subj-group><subj-group subj-group-type="section-heading" xml:lang="ru"><subject>Статьи</subject></subj-group></article-categories><title-group><article-title>Трансформация растительности как лимитирующий фактор существования популяций грызунов открытых пространств (на примере Spermophilus suslicus)</article-title><trans-title-group xml:lang="en"><trans-title>Vegetation transformation as a limiting factor in the existence of rodent populations in open area (using the example of Spermophilus suslicus)</trans-title></trans-title-group></title-group><contrib-group><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0001-9508-5008</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Леонова</surname><given-names>Н. А.</given-names></name><name name-style="western" xml:lang="en"><surname>Leonova</surname><given-names>N. A.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Леонова Наталья Алексеевна, кафедра зоологии и экологии факультета физико-математических и естественных наук </p><p>440026, г. Пенза, ул. Красная, д. 40</p></bio><bio xml:lang="en"><p>Natalia A. Leonova, department of Zoology and Ecology of the Faculty of Physical, Mathematical and Natural Sciences</p><p>40 Krasnaya St., Penza 440026</p></bio><email xlink:type="simple">na_leonova@mail.ru</email><xref ref-type="aff" rid="aff-1"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0001-9480-7523</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Картавов</surname><given-names>Н. А.</given-names></name><name name-style="western" xml:lang="en"><surname>Kartavov</surname><given-names>N. A.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Картавов Никита Александрович</p><p>440026, г. Пенза, ул. Красная, д. 40</p></bio><bio xml:lang="en"><p>Nikita A. Kartavov</p><p>40 Krasnaya St., Penza 440026</p></bio><email xlink:type="simple">nikitakartavov@yandex.ru</email><xref ref-type="aff" rid="aff-1"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0009-0007-8287-9221</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Чернышова</surname><given-names>О. В.</given-names></name><name name-style="western" xml:lang="en"><surname>Chernyshova</surname><given-names>O. V.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Чернышова Ольга Валерьевна</p><p>440026, г. Пенза, ул. Красная, д. 40</p></bio><bio xml:lang="en"><p>Olga V. Chernyshova</p><p>40 Krasnaya St., Penza 440026</p></bio><email xlink:type="simple">oliarabbit@yandex.ru</email><xref ref-type="aff" rid="aff-1"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0009-0004-4106-7188</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Волкова</surname><given-names>Е. К.</given-names></name><name name-style="western" xml:lang="en"><surname>Volkova</surname><given-names>E. K.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Волкова Елена Константиновна</p><p>440026, г. Пенза, ул. Красная, д. 40</p></bio><bio xml:lang="en"><p>Elena K. Volkova</p><p>40 Krasnaya St., Penza 440026</p></bio><email xlink:type="simple">elena.dad30@gmail.com</email><xref ref-type="aff" rid="aff-1"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0001-9634-1157</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Титов</surname><given-names>С. В.</given-names></name><name name-style="western" xml:lang="en"><surname>Titov</surname><given-names>S. V.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Титов Сергей Витальевич</p><p>440026, г. Пенза, ул. Красная, д. 40</p></bio><bio xml:lang="en"><p>Sergey V. Titov</p><p>40 Krasnaya St., Penza 440026</p></bio><email xlink:type="simple">svtitov@yandex.ru</email><xref ref-type="aff" rid="aff-1"/></contrib></contrib-group><aff-alternatives id="aff-1"><aff xml:lang="ru">Пензенский государственный университет<country>Россия</country></aff><aff xml:lang="en">Penza State University<country>Russian Federation</country></aff></aff-alternatives><pub-date pub-type="collection"><year>2025</year></pub-date><pub-date pub-type="epub"><day>09</day><month>04</month><year>2025</year></pub-date><volume>0</volume><issue>1</issue><fpage>31</fpage><lpage>47</lpage><permissions><copyright-statement>Copyright &amp;#x00A9; Леонова Н.А., Картавов Н.А., Чернышова О.В., Волкова Е.К., Титов С.В., 2025</copyright-statement><copyright-year>2025</copyright-year><copyright-holder xml:lang="ru">Леонова Н.А., Картавов Н.А., Чернышова О.В., Волкова Е.К., Титов С.В.</copyright-holder><copyright-holder xml:lang="en">Leonova N.A., Kartavov N.A., Chernyshova O.V., Volkova E.K., Titov S.V.</copyright-holder><license license-type="creative-commons-attribution" xlink:href="https://creativecommons.org/licenses/by/4.0/" xlink:type="simple"><license-p>This work is licensed under a Creative Commons Attribution 4.0 License.</license-p></license></permissions><self-uri xlink:href="https://sevin.elpub.ru/jour/article/view/627">https://sevin.elpub.ru/jour/article/view/627</self-uri><abstract><p>Приводится сравнительный анализ состава и структуры растительных сообществ в двух местах былого обитания крапчатого суслика (Spermophilus suslicus) в Николаевском районе Ульяновской области. Анализ современного состояния популяций S. suslicus в Поволжье показал, что на протяжении последних пятидесяти лет наблюдается критическое снижение численности вида, а в ряде регионов – его полное исчезновение. Рядом исследователей отмечается, что определенное влияние на эти процессы оказывают динамические изменения растительности. Поэтому целью нашей работы являлось сравнительное изучение растительности биотопов на момент высокой численности зверьков на фоне интенсивного выпаса и при отсутствии поселений суслика более 15 – 20 лет при отсутствии выпаса. Наши исследования показали, что происходит увеличение видового богатства и структурного разнообразия сообществ за счет внедрения в их состав видов боровой, неморальной и бореальной экологоценотических групп. Фитоценозы остепненных лугов с высоким участием Festuca valesiaca, Poa pratensis, Helictotrichon pubescens, Elytrigia repens после прекращения выпаса сменяются ценозами луговых степей с доминированием Bromopsis riparia, Poa angustifolia, Calamagrostis epigeios. Происходит увеличение общего проективного покрытия травостоя (с 40 до 60 – 70%). Кормовые виды, являющиеся фоновыми на момент высокой численности зверьков, резко сокращают свое участие в травостое. Характерны изменения экологических условий биотопов: современные сообщества формируются в более засушливых, менее богатых азотом и «кислых» местообитаниях. Сохранение и возможное восстановление поселений крапчатого суслика в Поволжье возможно при соблюдении научно обоснованного антропогенного воздействия, реализуемого в форме сенокошения и выпаса.</p></abstract><trans-abstract xml:lang="en"><p>A comparative analysis of the composition and structure of plant communities in two places of the former habitat of Spermophilus suslicus in the Nikolaevsky district of the Ulyanovsk region is provided. Our analysis of the current state of S. suslicus populations in the Volga region has shown that over the past fifty years there has been a critical decline in the numbers of the species, and even its complete disappearance has been noted in some regions. A number of researchers note that dynamic changes in vegetation have certain influence on these processes. Therefore, the goal of our work was a comparative study of the vegetation of biotopes at the time of high numbers of animals against the background of intensive grazing and in the absence of gopher settlements for more than 15–20 years in the absence of grazing. Our studies have shown an increase in the species richness and structural diversity of communities due to the introduction of species from the upland, nemoral and boreal ecological-coenotic groups into their composition. Phytocenoses of steppe meadows with a high participation of Festuca valesiaca, Poa pratensis, Helictotrichon pubescens, and Elytrigia repens after the cessation of grazing are replaced by cenoses of meadow steppes with the dominance of Bromopsis riparia, Poa angustifolia, and Calamagrostis epigeios. There is an increase in the total projective cover of the grass stand (from 40 up to 60– 70%). Forage species that are background ones when the number of animals is high sharply reduce their participation in the grass stand. Changes in the ecological conditions of biotopes are characteristic, namely: modern communities are formed in drier, less nitrogen-rich and “acidic” habitats. The preservation and possible restoration of spotted ground squirrel settlements in the Volga region is possible subject to scientifically based anthropogenic impact, implemented in the form of haymaking and grazing.</p></trans-abstract><kwd-group xml:lang="ru"><kwd>Spermophilus suslicus</kwd><kwd>растительность</kwd><kwd>видовое богатство</kwd><kwd>структура сообществ</kwd><kwd>трансформация растительности</kwd><kwd>Поволжье</kwd></kwd-group><kwd-group xml:lang="en"><kwd>Spermophilus suslicus</kwd><kwd>vegetation</kwd><kwd>species richness</kwd><kwd>community structure</kwd><kwd>vegetation transformation</kwd><kwd>Volga region</kwd></kwd-group><funding-group xml:lang="ru"><funding-statement>Работа выполнена при финансовой поддержке Российского научного фонда (проект № 22-24-20099, https://rscf.ru/project/24-24-20099).</funding-statement></funding-group><funding-group xml:lang="en"><funding-statement>This work was financially supported by the Russian Science Foundation (Project number 22-24-20099, https://rscf.ru/project/24-24-20099).</funding-statement></funding-group></article-meta></front><back><ref-list><title>References</title><ref id="cit1"><label>1</label><citation-alternatives><mixed-citation xml:lang="ru">Abaturov B. D. Microdepressions microrelief of Caspian lowland and mechanisms of its formation. Arid Ecosystems, 2010, vol. 16, no. 5, pp. 31–45 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Abaturov B. D. Microdepressions microrelief of Caspian lowland and mechanisms of its formation. Arid Ecosystems, 2010, vol. 16, no. 5, pp. 31–45 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit2"><label>2</label><citation-alternatives><mixed-citation xml:lang="ru">Abrakhina I. B. Influence of land plowing on the number and distribution of ground squirrels in the Ulyanovsk region. In: Vliyanie antropogennoi transformatsii landshafta na naselenie nazemnykh pozvonochnykh zhivotnykh: tezisy Vsesoyuznogo soveshchaniya [Influence of Anthropogenic Transformation of the Landscape on the Population of Terrestrial Vertebrates: Abstracts of the All-Union Meeting]. Moscow, VNII okhrany prirody i zapovednogo dela Gosagroproma SSSR, 1987, pt. 1, pp. 308–310 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Abrakhina I. B. Influence of land plowing on the number and distribution of ground squirrels in the Ulyanovsk region. In: Vliyanie antropogennoi transformatsii landshafta na naselenie nazemnykh pozvonochnykh zhivotnykh: tezisy Vsesoyuznogo soveshchaniya [Influence of Anthropogenic Transformation of the Landscape on the Population of Terrestrial Vertebrates: Abstracts of the All-Union Meeting]. Moscow, VNII okhrany prirody i zapovednogo dela Gosagroproma SSSR, 1987, pt. 1, pp. 308–310 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit3"><label>3</label><citation-alternatives><mixed-citation xml:lang="ru">Artemyev Yu. T. Ecological and Morphological Sketch of Squirrels of the Volga-Kama Region. Thesis Diss. Cand. Sci. (Biol.). Kazan, 1966. 24 p. (in Russian).</mixed-citation><mixed-citation xml:lang="en">Artemyev Yu. T. Ecological and Morphological Sketch of Squirrels of the Volga-Kama Region. Thesis Diss. Cand. Sci. (Biol.). Kazan, 1966. 24 p. (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit4"><label>4</label><citation-alternatives><mixed-citation xml:lang="ru">Assessment and Conservation of Forest Cover Biodiversity in the Reserves of European Russia. Moscow, Nauchnyi mir, 2000. 196 p. (in Russian).</mixed-citation><mixed-citation xml:lang="en">Assessment and Conservation of Forest Cover Biodiversity in the Reserves of European Russia. Moscow, Nauchnyi mir, 2000. 196 p. (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit5"><label>5</label><citation-alternatives><mixed-citation xml:lang="ru">Barabash-Nikiforov I. I. Zveri yugo-vostochnoi chasti Chernozemnogo Tsentra [Beasts of the South-Eastern Part of the Black Earth Center]. Voronezh, Voronezhskoe knizhnoe izdatel'stvo, 1957, pp. 219–230 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Barabash-Nikiforov I. I. Zveri yugo-vostochnoi chasti Chernozemnogo Tsentra [Beasts of the South-Eastern Part of the Black Earth Center]. Voronezh, Voronezhskoe knizhnoe izdatel'stvo, 1957, pp. 219–230 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit6"><label>6</label><citation-alternatives><mixed-citation xml:lang="ru">Bogdanov M. N. Birds and mammals of the chernozem zone in the Volga region and valleys of both Middle Volga and Lower Volga (biogeographic materials). Proceedings of the Society of Naturalists in the Imperial Kazan University, 1871, vol. 1, pp. 4–226 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Bogdanov M. N. Birds and mammals of the chernozem zone in the Volga region and valleys of both Middle Volga and Lower Volga (biogeographic materials). Proceedings of the Society of Naturalists in the Imperial Kazan University, 1871, vol. 1, pp. 4–226 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit7"><label>7</label><citation-alternatives><mixed-citation xml:lang="ru">Dufrene M., Legendre P. Species assemblages and indicator species: The need for a flexible asymmetrical approach. Ecological Monographs, 1997, vol. 67, no. 3, pp. 345–366. https://doi.org/10.2307/2963459</mixed-citation><mixed-citation xml:lang="en">Dufrene M., Legendre P. Species assemblages and indicator species: The need for a flexible asymmetrical approach. Ecological Monographs, 1997, vol. 67, no. 3, pp. 345–366. https://doi.org/10.2307/2963459</mixed-citation></citation-alternatives></ref><ref id="cit8"><label>8</label><citation-alternatives><mixed-citation xml:lang="ru">Fedorovich F. F. Animals and birds of the Penza province. Trudy Penzenskogo obshchestva lyubiteley prirody, 1915, iss. 2, pp. 41–76 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Fedorovich F. F. Animals and birds of the Penza province. Trudy Penzenskogo obshchestva lyubiteley prirody, 1915, iss. 2, pp. 41–76 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit9"><label>9</label><citation-alternatives><mixed-citation xml:lang="ru">Hill M. O., Gauch H. G. Detrended correspondence analysis: An improved ordination technique. Vegetatio, 1980, vol. 42, iss. 1–3, pp. 47–58. https://doi.org/10.1007/BF00048870</mixed-citation><mixed-citation xml:lang="en">Hill M. O., Gauch H. G. Detrended correspondence analysis: An improved ordination technique. Vegetatio, 1980, vol. 42, iss. 1–3, pp. 47–58. https://doi.org/10.1007/BF00048870</mixed-citation></citation-alternatives></ref><ref id="cit10"><label>10</label><citation-alternatives><mixed-citation xml:lang="ru">Krylova T. V., Deistfeldt L. A. Patterns of stationary distribution of settlements of the small mountain gopher. Ecologiya, 1987, no. 1, pp. 59–66 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Krylova T. V., Deistfeldt L. A. Patterns of stationary distribution of settlements of the small mountain gopher. Ecologiya, 1987, no. 1, pp. 59–66 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit11"><label>11</label><citation-alternatives><mixed-citation xml:lang="ru">Lobkov V. A. Spotted Sousliks of the Northeastern Black Sea Region: Biology and Population Functioning. Odessa, AstroPrint, 1999, pp. 113–120 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Lobkov V. A. Spotted Sousliks of the Northeastern Black Sea Region: Biology and Population Functioning. Odessa, AstroPrint, 1999, pp. 113–120 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit12"><label>12</label><citation-alternatives><mixed-citation xml:lang="ru">Lobkov V. A. Ecological reasons for changes in the number and distribution of the speckled ground squirrel Spermophilus suslicus (Guldenstaedt, 1770). Bulletin of Moscow Society of Naturalists. Biological Series, 2006, vol. 111, iss. 5, pp. 59–64 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Lobkov V. A. Ecological reasons for changes in the number and distribution of the speckled ground squirrel Spermophilus suslicus (Guldenstaedt, 1770). Bulletin of Moscow Society of Naturalists. Biological Series, 2006, vol. 111, iss. 5, pp. 59–64 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit13"><label>13</label><citation-alternatives><mixed-citation xml:lang="ru">Magerramov S. V., Martsokha K. S., Yakovlev S. A., Mandzhieva V. S., Bondarev V. A., LidzhiGaryaeva G. V., Matrosov A. N., Popov N. V. Modern climate warming impact on the population dynamics of the small ground squirrel (Spermophilus pygmaeus Pallas, 1778) (Rodentia, Mammalia) in the Ilmenno-Prideltovy region of the Caspian lowland. Povolzhskiy Journal of Ecology, 2022, no. 1, pp. 17–33 (in Russian). https://doi.org/10.35885/1684-7318-2022-1-17-33</mixed-citation><mixed-citation xml:lang="en">Magerramov S. V., Martsokha K. S., Yakovlev S. A., Mandzhieva V. S., Bondarev V. A., LidzhiGaryaeva G. V., Matrosov A. N., Popov N. V. Modern climate warming impact on the population dynamics of the small ground squirrel (Spermophilus pygmaeus Pallas, 1778) (Rodentia, Mammalia) in the Ilmenno-Prideltovy region of the Caspian lowland. Povolzhskiy Journal of Ecology, 2022, no. 1, pp. 17–33 (in Russian). https://doi.org/10.35885/1684-7318-2022-1-17-33</mixed-citation></citation-alternatives></ref><ref id="cit14"><label>14</label><citation-alternatives><mixed-citation xml:lang="ru">Magerramov Sh. V., Matrosov A. N., Bocharnikova T. A., Grazhdanov A. K., Martsokha K. S., Kuznetsov A. A., Sludsky A. A., Popov N. V. Decrease in the population of the small ground squirrel – Spermophilus pygmaeus Pallas, 1779 (Rodentia, Sciuromorpha, Sciuridae) in the Volga– Ural steppe natural focus of plague. Povolzhskiy Journal of Ecology, 2025, no. 1, pp. 48–63 (in Russian). https://doi.org/10.35885/1684-7318-2025-1-48-63</mixed-citation><mixed-citation xml:lang="en">Magerramov Sh. V., Matrosov A. N., Bocharnikova T. A., Grazhdanov A. K., Martsokha K. S., Kuznetsov A. A., Sludsky A. A., Popov N. V. Decrease in the population of the small ground squirrel – Spermophilus pygmaeus Pallas, 1779 (Rodentia, Sciuromorpha, Sciuridae) in the Volga– Ural steppe natural focus of plague. Povolzhskiy Journal of Ecology, 2025, no. 1, pp. 48–63 (in Russian). https://doi.org/10.35885/1684-7318-2025-1-48-63</mixed-citation></citation-alternatives></ref><ref id="cit15"><label>15</label><citation-alternatives><mixed-citation xml:lang="ru">McCune B., Grace J. B. Analysis of Ecological Communities. Gleneden Beach, Oregon, MjM Software Design, 2002. 300 p.</mixed-citation><mixed-citation xml:lang="en">McCune B., Grace J. B. Analysis of Ecological Communities. Gleneden Beach, Oregon, MjM Software Design, 2002. 300 p.</mixed-citation></citation-alternatives></ref><ref id="cit16"><label>16</label><citation-alternatives><mixed-citation xml:lang="ru">McCune B., Mefford M. J. Multivariate Analysis of Ecological Data. Gleneden Beach, Oregon, MjM Software Design, 1999. 237 p.</mixed-citation><mixed-citation xml:lang="en">McCune B., Mefford M. J. Multivariate Analysis of Ecological Data. Gleneden Beach, Oregon, MjM Software Design, 1999. 237 p.</mixed-citation></citation-alternatives></ref><ref id="cit17"><label>17</label><citation-alternatives><mixed-citation xml:lang="ru">Nedosekin V. Yu., Ushakov M. V. On the state of the speckled ground squirrel in the Lipetsk region. In: Ground Squirrels of Eurasia (Genera Spermophilus, Spermophilopsis): Origin, Taxonomy, Ecology, Behavior, and Species Diversity Conservation: Proceedings of the Russian Scientific Conference. Moscow, KMK, 2005, pp. 65–66 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Nedosekin V. Yu., Ushakov M. V. On the state of the speckled ground squirrel in the Lipetsk region. In: Ground Squirrels of Eurasia (Genera Spermophilus, Spermophilopsis): Origin, Taxonomy, Ecology, Behavior, and Species Diversity Conservation: Proceedings of the Russian Scientific Conference. Moscow, KMK, 2005, pp. 65–66 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit18"><label>18</label><citation-alternatives><mixed-citation xml:lang="ru">Nitsenko A. A. On the problem of investigation of ecological structure of the vegetational cover. Botanicheskii zhurnal, 1969, vol. 54, no. 7, pp. 1002–1014 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Nitsenko A. A. On the problem of investigation of ecological structure of the vegetational cover. Botanicheskii zhurnal, 1969, vol. 54, no. 7, pp. 1002–1014 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit19"><label>19</label><citation-alternatives><mixed-citation xml:lang="ru">Okulova N. M. A burrow of the little souslik as a consortia. Parazitologiya, 2003, vol. 37, iss. 5, pp. 361–380 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Okulova N. M. A burrow of the little souslik as a consortia. Parazitologiya, 2003, vol. 37, iss. 5, pp. 361–380 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit20"><label>20</label><citation-alternatives><mixed-citation xml:lang="ru">Oparin M. L., Oparina O. S. Bird and mammal complex transformation of steppe ecosystems under ploughing up (with Saratov steppes as examples). Povolzhskiy Journal of Ecology, 2010, no. 4, pp. 361–373 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Oparin M. L., Oparina O. S. Bird and mammal complex transformation of steppe ecosystems under ploughing up (with Saratov steppes as examples). Povolzhskiy Journal of Ecology, 2010, no. 4, pp. 361–373 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit21"><label>21</label><citation-alternatives><mixed-citation xml:lang="ru">Persson S. Ecological indicator values as an aid in the interpretation of ordination diagrams. Journal of Ecology, 1981, vol. 69, no. 1, pp. 71–84. https://doi.org/10.2307/225981</mixed-citation><mixed-citation xml:lang="en">Persson S. Ecological indicator values as an aid in the interpretation of ordination diagrams. Journal of Ecology, 1981, vol. 69, no. 1, pp. 71–84. https://doi.org/10.2307/225981</mixed-citation></citation-alternatives></ref><ref id="cit22"><label>22</label><citation-alternatives><mixed-citation xml:lang="ru">Pivanova S. V., Shubina Yu. E. Factors affecting the abundance of speckled ground squirrel (Spermophilus suslicus Güldenstaedt, 1770) in natural and anthropogenic landscapes. In: Pest Management and Problems of Biodiversity Conservation: Materials of the Russian ScientificPractical Conference with International Participation. Moscow, A. N. Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences Publ., 2009, pp. 48–49 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Pivanova S. V., Shubina Yu. E. Factors affecting the abundance of speckled ground squirrel (Spermophilus suslicus Güldenstaedt, 1770) in natural and anthropogenic landscapes. In: Pest Management and Problems of Biodiversity Conservation: Materials of the Russian ScientificPractical Conference with International Participation. Moscow, A. N. Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences Publ., 2009, pp. 48–49 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit23"><label>23</label><citation-alternatives><mixed-citation xml:lang="ru">Popov V. A. Mlekopitaiushchie Volzhsko-Kamskogo kraya (nasekomoyadnye, rukokrylye, gryzuny) [Mammals of the Volga-Kama Region (Insectivora, Chiroptera, Rodentia)]. Kazan, Kazan Branch of the USSR Academy of Sciences Publ., 1960. 468 p. (in Russian).</mixed-citation><mixed-citation xml:lang="en">Popov V. A. Mlekopitaiushchie Volzhsko-Kamskogo kraya (nasekomoyadnye, rukokrylye, gryzuny) [Mammals of the Volga-Kama Region (Insectivora, Chiroptera, Rodentia)]. Kazan, Kazan Branch of the USSR Academy of Sciences Publ., 1960. 468 p. (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit24"><label>24</label><citation-alternatives><mixed-citation xml:lang="ru">Rodimtsev A. S. New settlements of spotted souslik (Spermophilus suslicus Güld., 1770) in Tambov region. Tambov University Reports. Series: Natural and Technical Sciences, 2012, vol. 17, iss. 2, pp. 784–785 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Rodimtsev A. S. New settlements of spotted souslik (Spermophilus suslicus Güld., 1770) in Tambov region. Tambov University Reports. Series: Natural and Technical Sciences, 2012, vol. 17, iss. 2, pp. 784–785 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit25"><label>25</label><citation-alternatives><mixed-citation xml:lang="ru">Sapelnikov S. F., Sapelnikova I. I. Postnatal development of speckled ground squirrel (Spermophilus suslicus Güld, 1770) (Sciuridae, Mammalia) in Captivity. Povolzhskiy Journal of Ecology, 2019, no. 1, pp. 47–60. https://doi.org/10.35885/1684-7318-2019-1-47-60</mixed-citation><mixed-citation xml:lang="en">Sapelnikov S. F., Sapelnikova I. I. Postnatal development of speckled ground squirrel (Spermophilus suslicus Güld, 1770) (Sciuridae, Mammalia) in Captivity. Povolzhskiy Journal of Ecology, 2019, no. 1, pp. 47–60. https://doi.org/10.35885/1684-7318-2019-1-47-60</mixed-citation></citation-alternatives></ref><ref id="cit26"><label>26</label><citation-alternatives><mixed-citation xml:lang="ru">Sapelnikov S. F., Sapelnikova I. I. Retrospective analysis of condition of populations of Spermophilus suslicus Güldenstädt, 1770 in Central Black Earth region and in adjacent territories and possible ways of preserving the species. Field Biologist Journal, 2021, vol. 3, no. 2, pp. 167– 212 (in Russian). https://doi.org/10.52575/2658-3453-2021-3-2-167-212</mixed-citation><mixed-citation xml:lang="en">Sapelnikov S. F., Sapelnikova I. I. Retrospective analysis of condition of populations of Spermophilus suslicus Güldenstädt, 1770 in Central Black Earth region and in adjacent territories and possible ways of preserving the species. Field Biologist Journal, 2021, vol. 3, no. 2, pp. 167– 212 (in Russian). https://doi.org/10.52575/2658-3453-2021-3-2-167-212</mixed-citation></citation-alternatives></ref><ref id="cit27"><label>27</label><citation-alternatives><mixed-citation xml:lang="ru">Shilova S. A. Abundance control and conservation of sousliks in Russia (G. spermophilus). Arid Ecosystems, 2011, vol. 17, no. 4, pp. 267–272. https://doi.org/10.1134/S2079096111040147</mixed-citation><mixed-citation xml:lang="en">Shilova S. A. Abundance control and conservation of sousliks in Russia (G. spermophilus). Arid Ecosystems, 2011, vol. 17, no. 4, pp. 267–272. https://doi.org/10.1134/S2079096111040147</mixed-citation></citation-alternatives></ref><ref id="cit28"><label>28</label><citation-alternatives><mixed-citation xml:lang="ru">Shilova S. A., Neronov V. V., Shekarova O. N., Savinetskaya L. E. Dynamics of colonies of the speckled ground squirrel (Spermophilus suslicus Guld., 1770) on the northern boundary of the habitat. Biology Bulletin, 2010, vol. 37, no. 5, pp. 532–536. https://doi.org/10.1134/S1062359010050134</mixed-citation><mixed-citation xml:lang="en">Shilova S. A., Neronov V. V., Shekarova O. N., Savinetskaya L. E. Dynamics of colonies of the speckled ground squirrel (Spermophilus suslicus Guld., 1770) on the northern boundary of the habitat. Biology Bulletin, 2010, vol. 37, no. 5, pp. 532–536. https://doi.org/10.1134/S1062359010050134</mixed-citation></citation-alternatives></ref><ref id="cit29"><label>29</label><citation-alternatives><mixed-citation xml:lang="ru">Smirnov V. E., Khanina L. G., Bobrovsky M. V. Validation of the ecological-coenotical groups of vascular plant species for European Russian forests on the basis of ecological indicator values, vegetation releves and statistical analysis. Bulletin of Moscow Society of Naturalists. Biological Series, 2006, vol. 111, iss. 2, pp. 36–47 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Smirnov V. E., Khanina L. G., Bobrovsky M. V. Validation of the ecological-coenotical groups of vascular plant species for European Russian forests on the basis of ecological indicator values, vegetation releves and statistical analysis. Bulletin of Moscow Society of Naturalists. Biological Series, 2006, vol. 111, iss. 2, pp. 36–47 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit30"><label>30</label><citation-alternatives><mixed-citation xml:lang="ru">Smirnova O. V., ed. Forests of Eastern Europe: Holocene History and Modern Times: in 2 books. Moscow, Nauka, 2004, 1 books. 476 p. (in Russian).</mixed-citation><mixed-citation xml:lang="en">Smirnova O. V., ed. Forests of Eastern Europe: Holocene History and Modern Times: in 2 books. Moscow, Nauka, 2004, 1 books. 476 p. (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit31"><label>31</label><citation-alternatives><mixed-citation xml:lang="ru">Stoyko T. G., Denisov V. P., Gudkov A. S. Speckled ground squirrel as an indicator of anthropogenic impact on steppe communities of the forest-steppe zone. In: Materialy V soveshchaniya po gryzunam [Proceedings of the V Meeting on Rodents]. Moscow, Nauka, 1980, pp. 447–449 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Stoyko T. G., Denisov V. P., Gudkov A. S. Speckled ground squirrel as an indicator of anthropogenic impact on steppe communities of the forest-steppe zone. In: Materialy V soveshchaniya po gryzunam [Proceedings of the V Meeting on Rodents]. Moscow, Nauka, 1980, pp. 447–449 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit32"><label>32</label><citation-alternatives><mixed-citation xml:lang="ru">Tikhvinsky V. I. Materialy po izucheniyu suslika v Tatrespublike [Materials on the Study of the souslik in the Tatar Republic]. Kazan, Oblastnaya poligrafshkola FZU im. Lunacharskogo, 1931. 11 p. (in Russian).</mixed-citation><mixed-citation xml:lang="en">Tikhvinsky V. I. Materialy po izucheniyu suslika v Tatrespublike [Materials on the Study of the souslik in the Tatar Republic]. Kazan, Oblastnaya poligrafshkola FZU im. Lunacharskogo, 1931. 11 p. (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit33"><label>33</label><citation-alternatives><mixed-citation xml:lang="ru">Titov S. V. Current distribution and changes in number of the spotted souslik, Spermophilus suslicus, in the eastern part of its range. Zoologicheskiy zhurnal, 2001, vol. 80, no. 2, pp. 230–235 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Titov S. V. Current distribution and changes in number of the spotted souslik, Spermophilus suslicus, in the eastern part of its range. Zoologicheskiy zhurnal, 2001, vol. 80, no. 2, pp. 230–235 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit34"><label>34</label><citation-alternatives><mixed-citation xml:lang="ru">Titov S. V. Habitat preference in Spermophilus major and S. suslicus in recently developed sympatric zone. Zoologicheskiy zhurnal, 2000, vol. 79, no. 1, pp. 64–72 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Titov S. V. Habitat preference in Spermophilus major and S. suslicus in recently developed sympatric zone. Zoologicheskiy zhurnal, 2000, vol. 79, no. 1, pp. 64–72 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit35"><label>35</label><citation-alternatives><mixed-citation xml:lang="ru">Titov S. V. Relationships Between the Speckled Ground Squirrel and the Great Ground Squirrel in the Recently Emerged Sympatry Zone. Thesis Diss. Cand. Sci. (Biol.). Moscow, 1999. 24 p. (in Russian).</mixed-citation><mixed-citation xml:lang="en">Titov S. V. Relationships Between the Speckled Ground Squirrel and the Great Ground Squirrel in the Recently Emerged Sympatry Zone. Thesis Diss. Cand. Sci. (Biol.). Moscow, 1999. 24 p. (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit36"><label>36</label><citation-alternatives><mixed-citation xml:lang="ru">Titov S. V., Kartavov N. A., Leonova N. A., Smirnov D. G., Chernyshova O. V., Kuzmin A. A. Current status of populations of speckled ground squirrels (Spermophilus suslicus Güld.) in Volga region: History and scale of extinction. Russian Journal of Ecosystem Ecology, 2024, vol. 9, no. 3, pp. 1–13 (in Russian). https://doi.org/10.21685/2500-0578-2024-3-1</mixed-citation><mixed-citation xml:lang="en">Titov S. V., Kartavov N. A., Leonova N. A., Smirnov D. G., Chernyshova O. V., Kuzmin A. A. Current status of populations of speckled ground squirrels (Spermophilus suslicus Güld.) in Volga region: History and scale of extinction. Russian Journal of Ecosystem Ecology, 2024, vol. 9, no. 3, pp. 1–13 (in Russian). https://doi.org/10.21685/2500-0578-2024-3-1</mixed-citation></citation-alternatives></ref><ref id="cit37"><label>37</label><citation-alternatives><mixed-citation xml:lang="ru">Titov S. V., Kuzmin A. A., Naumov R. V., Ermakov O. A., Zaks S. S., Chernyshova O. V. Dynamics of Habitats and the Current State of Settlements of Terrestrial Squirrels in the Right Bank Regions of the Volga Region. Penza, Penza State University Publ., 2015. 124 p. (in Russian).</mixed-citation><mixed-citation xml:lang="en">Titov S. V., Kuzmin A. A., Naumov R. V., Ermakov O. A., Zaks S. S., Chernyshova O. V. Dynamics of Habitats and the Current State of Settlements of Terrestrial Squirrels in the Right Bank Regions of the Volga Region. Penza, Penza State University Publ., 2015. 124 p. (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit38"><label>38</label><citation-alternatives><mixed-citation xml:lang="ru">Tkachenko V. S. Placement of mountain gopher burrows inside isolated habitats. In: Ekologiya i okhrana gornykh vidov mlekopitayushchikh: materialy III Vsesoyuznoj shkoly [Ecology and Protection of Mountain Species of Mammals: Materials of the III All-Union School]. Moscow, Severtsov Institute of Evolutionary Morphology and Ecology of Animals of the USSR Academy of Sciences Publ., 1987, pp. 183–185 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Tkachenko V. S. Placement of mountain gopher burrows inside isolated habitats. In: Ekologiya i okhrana gornykh vidov mlekopitayushchikh: materialy III Vsesoyuznoj shkoly [Ecology and Protection of Mountain Species of Mammals: Materials of the III All-Union School]. Moscow, Severtsov Institute of Evolutionary Morphology and Ecology of Animals of the USSR Academy of Sciences Publ., 1987, pp. 183–185 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit39"><label>39</label><citation-alternatives><mixed-citation xml:lang="ru">Tsyganov D. N. Fitoindikatsiya ekologicheskikh rezhimov v podzone khvoyno-shirokolistvennykh lesov [Phyto-indication of Ecological Regimes in the Subzone of Coniferous-deciduous Forests.]. Moscow, Nauka, 1983. 196 p. (in Russian).</mixed-citation><mixed-citation xml:lang="en">Tsyganov D. N. Fitoindikatsiya ekologicheskikh rezhimov v podzone khvoyno-shirokolistvennykh lesov [Phyto-indication of Ecological Regimes in the Subzone of Coniferous-deciduous Forests.]. Moscow, Nauka, 1983. 196 p. (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit40"><label>40</label><citation-alternatives><mixed-citation xml:lang="ru">Zhigarev I. V., Rumyantsev V. Yu. Modern situation in spotted ground squirrel (Spermophilus suslicus) populations within the forest steppe of European Russiа. Bulletin of Moscow Society of Naturalists. Biological Series, 2024, vol. 129, iss. 6, pp. 3–13 (in Russian). https://doi.org/10.55959/MSU0027-1403-BB-2024-129-6-3-13</mixed-citation><mixed-citation xml:lang="en">Zhigarev I. V., Rumyantsev V. Yu. Modern situation in spotted ground squirrel (Spermophilus suslicus) populations within the forest steppe of European Russiа. Bulletin of Moscow Society of Naturalists. Biological Series, 2024, vol. 129, iss. 6, pp. 3–13 (in Russian). https://doi.org/10.55959/MSU0027-1403-BB-2024-129-6-3-13</mixed-citation></citation-alternatives></ref><ref id="cit41"><label>41</label><citation-alternatives><mixed-citation xml:lang="ru">Zozulin G. M. Historical formations of vegetation of a part of the USSR. Botanicheskii zhurnal, 1973, vol. 58, no. 8, pp. 1081–1092 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Zozulin G. M. Historical formations of vegetation of a part of the USSR. Botanicheskii zhurnal, 1973, vol. 58, no. 8, pp. 1081–1092 (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit42"><label>42</label><citation-alternatives><mixed-citation xml:lang="ru">Zozulin G. M. Relationships between forest and herbaceous vegetation in the Central Black Earth State Reserve. Proceedings of the Central Chernozem State Reserve, 1955, iss. 3, pp. 102– 234 (in Russian).</mixed-citation><mixed-citation xml:lang="en">Zozulin G. M. Relationships between forest and herbaceous vegetation in the Central Black Earth State Reserve. Proceedings of the Central Chernozem State Reserve, 1955, iss. 3, pp. 102– 234 (in Russian).</mixed-citation></citation-alternatives></ref></ref-list><fn-group><fn fn-type="conflict"><p>The authors declare that there are no conflicts of interest present.</p></fn></fn-group></back></article>
