<?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">seamed</journal-id><journal-title-group><journal-title xml:lang="ru">Морская медицина</journal-title><trans-title-group xml:lang="en"><trans-title>Marine Medicine</trans-title></trans-title-group></journal-title-group><issn pub-type="ppub">2413-5747</issn><issn pub-type="epub">2587-7828</issn><publisher><publisher-name>Research Institute of Industrial and Maritime Medicine of the Federal Medical and Biological Agency</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="doi">10.22328/2413-5747-2026-12-3-103-111</article-id><article-id custom-type="edn" pub-id-type="custom">NAAJHO</article-id><article-id custom-type="elpub" pub-id-type="custom">seamed-1014</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>Оригинальные статьи</subject></subj-group><subj-group subj-group-type="section-heading" xml:lang="en"><subject>Original articles</subject></subj-group></article-categories><title-group><article-title>Штамм Staphylococcus hominis 23-2 из образца многолетней мерзлоты мыса Сопочная Карга Таймырского Долгано-Ненецкого муниципального района Красноярского края: оригинальная статья</article-title><trans-title-group xml:lang="en"><trans-title>Strain of Staphylococcus hominis 23-2 from a permafrost sample of cape Sopochnaya Karga, Taimyr Dolgan-Nenets municipal district, Krasnoyarsk Territory: original article</trans-title></trans-title-group></title-group><contrib-group><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-5206-6656</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Гончаров</surname><given-names>А. Е.</given-names></name><name name-style="western" xml:lang="en"><surname>Goncharov</surname><given-names>A. E.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Гончаров Артемий Евгеньевич — доктор медицинских наук, заведующий лабораторией микробиологического мониторинга биологических рисков отдела молекулярной микробиологии им. академика А. А. Тотоляна</p><p>197022, Санкт-Петербург, ул. Академика Павлова, д. 12</p></bio><bio xml:lang="en"><p>Artemy E. Goncharov — Cand. of Sci. (Med.), Head of the Laboratory of Microbiological Monitoring of Biological Risks of the Department of Molecular Microbiology named after Academician A. A. Totolian</p><p>197022, Saint Petersburg, Academician Pavlov Str., 12</p></bio><email xlink:type="simple">phage1@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/0000-0003-2483-5144</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Азаров</surname><given-names>Д. В.</given-names></name><name name-style="western" xml:lang="en"><surname>Azarov</surname><given-names>D. V.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Азаров Даниил Валерьевич — кандидат медицинских наук, старший преподаватель кафедры эпидемиологии, паразитологии и дезинфектологии</p><p>191015, Санкт-Петербург, ул. Кирочная, д. 41</p></bio><bio xml:lang="en"><p>Daniil V. Azarov — Cand. of Sci. (Med.), Senior Lecturer of the Department of Epidemiology, Parasitology and Disinfection</p><p>191015, Saint Petersburg, Kirochnaya Str., 41</p></bio><email xlink:type="simple">denazarov.da@yandex.ru</email><xref ref-type="aff" rid="aff-2"/></contrib><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Хорошилов</surname><given-names>В. Ю.</given-names></name><name name-style="western" xml:lang="en"><surname>Khoroshilov</surname><given-names>V. Yu.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Хорошилов Владимир Юрьевич — кандидат медицинских наук, лаборант-исследователь лаборатории микробиологического мониторинга биологических рисков отдела молекулярной микробиологии им. академика А. А. Тотоляна</p><p>197022, Санкт-Петербург, ул. Академика Павлова, д. 12</p></bio><bio xml:lang="en"><p>Vladimir Yu. Khoroshilov — Cand. of Sci. (Med.), Researcher of the Laboratory of Microbiological Monitoring of Biological Risks of the Department of Molecular Microbiology named after Academician A. A. Totolian</p><p>197022, Saint Petersburg, Aсademician Pavlov Str., 12</p></bio><email xlink:type="simple">khoroshiloff@yandex.ru</email><xref ref-type="aff" rid="aff-3"/></contrib><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Бартова</surname><given-names>А. В.</given-names></name><name name-style="western" xml:lang="en"><surname>Bartova</surname><given-names>A. V.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Бартова Арина Викторовна — ведущий геолог (начальник отряда), отдел региональной геологии и полезных ископаемых восточных районов России</p><p>199106, Санкт-Петербург, Средний проспект, д. 74</p></bio><bio xml:lang="en"><p>Arina V. Bartova — Leading Geologist, Department of Regional Geology and Mineral Resources of the Eastern Regions of Russia</p><p>199106, Saint Petersburg, Sredny prospekt, 74</p></bio><email xlink:type="simple">arina_bartova@mail.ru</email><xref ref-type="aff" rid="aff-4"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-6890-8096</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Асланов</surname><given-names>Б. И.</given-names></name><name name-style="western" xml:lang="en"><surname>Aslanov</surname><given-names>B. I.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Асланов Батырбек Исмелович — доктор медицинских наук, профессор, директор института профилактической медицины, заведующий кафедрой эпидемиологии, паразитологии и дезинфектологии</p><p>191015, Санкт-Петербург, ул. Кирочная, д. 41</p></bio><bio xml:lang="en"><p>Batyrbek I. Aslanov — Cand. of Sci. (Med.), Professor, Director of the Institute of Preventive Medicine, Head of the Department of Epidemiology, Parasitology and Disinfection</p><p>191015, Saint Petersburg, Kirochnaya Str., 41</p></bio><email xlink:type="simple">batyra@gmail.com</email><xref ref-type="aff" rid="aff-2"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-9900-5332</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Косякова</surname><given-names>К. Г.</given-names></name><name name-style="western" xml:lang="en"><surname>Kosyakova</surname><given-names>K. G.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Косякова Карина Георгиевна — кандидат медицинских наук, доцент кафедры медицинской микробиологии</p><p>191015, Санкт-Петербург, ул. Кирочная, д. 41</p></bio><bio xml:lang="en"><p>Karina G. Kosyakova — Cand. of Sci. (Med.), Associate Professor of the Department of Medical Microbiology</p><p>191015, Saint Petersburg, Kirochnaya Str., 41</p></bio><email xlink:type="simple">karina.kosyakova@szgmu.ru</email><xref ref-type="aff" rid="aff-2"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-2152-9881</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Бердичевский</surname><given-names>Г. М.</given-names></name><name name-style="western" xml:lang="en"><surname>Berdichevsky</surname><given-names>G. M.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Бердичевский Григорий Михайлович — кандидат биологических наук, заведующий лабораторией биомедицинской и фармацевтической масс-спектрометрии отдела молекулярной микробиологии им. академика А. А. Тотоляна</p><p>197022, Санкт-Петербург, ул. Академика Павлова, д. 12</p></bio><bio xml:lang="en"><p>Grigory M. Berdichevsky – Cand. of Sci. (Med.), Head of the Laboratory of Biomedical and Pharmaceutical Mass Spectrometry of the Department of Molecular Microbiology named after Academician A.A. Totolyan</p><p>197022, Saint Petersburg, Akademika Pavlova Street, 12</p></bio><email xlink:type="simple">grishaberd@gmail.com</email><xref ref-type="aff" rid="aff-3"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0009-0003-5188-8900</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Эргашев</surname><given-names>О. Н.</given-names></name><name name-style="western" xml:lang="en"><surname>Ergashev</surname><given-names>O. N.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Эргашев Олег Николаевич — доктор медицинских наук, профессор, исполняющий обязанности директора</p><p>197022, Санкт-Петербург, ул. Академика Павлова, д. 12</p></bio><bio xml:lang="en"><p>Oleg N. Ergashev – Dr. of Sci. (Med.), Professor, Acting Director</p><p>197022, Saint Petersburg, Academician Pavlov Str., 12</p></bio><email xlink:type="simple">iem@iemspb.ru</email><xref ref-type="aff" rid="aff-3"/></contrib></contrib-group><aff-alternatives id="aff-1"><aff xml:lang="ru"><institution>Институт экспериментальной медицины;&#13;
Северо-Западный государственный медицинский университет им. И. И. Мечникова</institution><country>Россия</country></aff><aff xml:lang="en"><institution>Institute of Experimental Medicine;&#13;
North-Western State Medical University named after I.I. Mechnikov</institution><country>Russian Federation</country></aff></aff-alternatives><aff-alternatives id="aff-2"><aff xml:lang="ru"><institution>Северо-Западный государственный медицинский университет им. И. И. Мечникова</institution><country>Россия</country></aff><aff xml:lang="en"><institution>North-Western State Medical University named after I. I. Mechnikov</institution><country>Russian Federation</country></aff></aff-alternatives><aff-alternatives id="aff-3"><aff xml:lang="ru"><institution>Институт экспериментальной медицины</institution><country>Россия</country></aff><aff xml:lang="en"><institution>Institute of Experimental Medicine</institution><country>Russian Federation</country></aff></aff-alternatives><aff-alternatives id="aff-4"><aff xml:lang="ru"><institution>Всероссийский научно-исследовательский геологический институт им. А. П. Карпинского</institution><country>Россия</country></aff><aff xml:lang="en"><institution>All-Russian Research Geological Institute named after A. P. Karpinsky</institution><country>Russian Federation</country></aff></aff-alternatives><pub-date pub-type="collection"><year>2026</year></pub-date><pub-date pub-type="epub"><day>10</day><month>10</month><year>2026</year></pub-date><volume>12</volume><issue>3</issue><fpage>103</fpage><lpage>111</lpage><permissions><copyright-statement>Copyright &amp;#x00A9; Гончаров А.Е., Азаров Д.В., Хорошилов В.Ю., Бартова А.В., Асланов Б.И., Косякова К.Г., Бердичевский Г.М., Эргашев О.Н., 2026</copyright-statement><copyright-year>2026</copyright-year><copyright-holder xml:lang="ru">Гончаров А.Е., Азаров Д.В., Хорошилов В.Ю., Бартова А.В., Асланов Б.И., Косякова К.Г., Бердичевский Г.М., Эргашев О.Н.</copyright-holder><copyright-holder xml:lang="en">Goncharov A.E., Azarov D.V., Khoroshilov V.Y., Bartova A.V., Aslanov B.I., Kosyakova K.G., Berdichevsky G.M., Ergashev O.N.</copyright-holder><license xml:lang="ru" license-type="creative-commons-attribution" xlink:href="https://creativecommons.org/licenses/by/4.0/" xlink:type="simple"><license-p>Данная работа распространяется под лицензией Creative Commons Attribution 4.0.</license-p></license><license xml:lang="en" 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://seamed.elpub.ru/jour/article/view/1014">https://seamed.elpub.ru/jour/article/view/1014</self-uri><abstract><p>ВВЕДЕНИЕ. Многолетняя мерзлота способна сохранять жизнеспособность древних микроорганизмов, в том числе патогенов животных и человека, на протяжении по меньшей мере тысячелетий. Изучение древних микробиомов, включая детерминанты устойчивости к антибиотикам, позволяет оценить потенциальные риски, связанные с распространением генетического материала древних микроорганизмов в современных экосистемах.ЦЕЛЬ. Определить генетические особенности и патогенный потенциал штамма Staphylococcus hominis 23-2, выделенного из отложений древних (неоплейстоценовых) многолетнемерзлых пород на мысе Сопочная Карга (северо-запад полуострова Таймыр, Красноярский край), а также оценить разнообразие детерминант антибиотикорезистентности мерзлотного микробиома, компонентом которого является данный штамм.МАТЕРИАЛЫ И МЕТОДЫ. Образцы мерзлых пород отбирали с соблюдением протоколов предотвращения контаминации и использовали для выделения ДНК и микробиологического посева на питательные среды общего назначения. Геном выделенного штамма Staphylococcus hominis секвенировали на приборе DNBSEQ-G50 (MGI, Китай) с длиной прочтений 2 × 100 пар нуклеотидов (далее — п. н.). Метагеномное секвенирование проводили на секвенаторе DNBSEQ-G400 с длиной парноконцевых прочтений 150 п. н. Биоинформационная обработка результатов секвенирования штамма Staphylococcus hominis включала филогеномный анализ последовательности генома, идентификацию генов лекарственной устойчивости и патогенности. Результаты метагеномного секвенирования использовали для идентификации генов устойчивости к антибиотикам.РЕЗУЛЬТАТЫ. Выделен штамм Staphylococcus hominis 23-2. В структуре его генома идентифицированы последовательности гена устойчивости к макролидным антибиотикам (msr(A)) плазмидной локализации и гена устойчивости к фузидиевой кислоте (fusB). Филогеномный анализ свидетельствует в пользу близости генома изучаемого штамма к штаммам современных генетических линий. В структуре метагенома идентифицировано 78 уникальных генов устойчивости к антибиотикам, включая гены эффлюкс-насосов семейств MFS и RND, гены β-лактамаз и гены ферментов модификации аминогликозидов. В составе резистома присутствуют последовательности генов резистентности к антибиотикам «последней линии»: гликопептидам (vanR), карбапенемам (металло-β-лактамазы, SHN-1, OXA-60) и полимиксину (pmr).ОБСУЖДЕНИЕ. Результаты исследования свидетельствуют о широкой представленности генов лекарственной устойчивости в образце арктической мерзлоты на мысе Сопочная Карга. При этом наличие в структуре генома штамма Staphylococcus hominis 23-2 генов лекарственной устойчивости, наряду с плазмидными и профаговыми последовательностями, позволяет предположить возможность мобилизации данным штаммом компонентов древнего резистома.ЗАКЛЮЧЕНИЕ. Исследование продемонстрировало принципиальную возможность сохранения в отложениях неоплейстоценовой мерзлоты мыса Сопочная Карга штаммов стафилококков, несущих гены лекарственной устойчивости и обладающих патогенным потенциалом. Полученные результаты обосновывают целесообразность мониторинга динамики распространения генов лекарственной устойчивости в качестве элемента региональных программ оценки биологических рисков, связанных с изменением климата в прибрежных экосистемах Арктики.</p></abstract><trans-abstract xml:lang="en"><p>INTRODUCTION. Permafrost can preserve the viability of ancient microorganisms, including animal and human pathogens, for at least millennia. Studying ancient microbiomes, including antibiotic resistance determinants, allows us to assess the potential risks associated with the spread of ancient microbial genetic material into modern ecosystems.OBJECTIVE. To determine the genetic characteristics and pathogenic potential of the Staphylococcus hominis strain 23-2 isolated from ancient (Neopleistocene) permafrost deposits at Cape Sopochnaya Karga (northwest of the Taimyr Peninsula, Krasnoyarsk Krai), and to evaluate the diversity of antibiotic resistance determinants in the permafrost microbiome of which this strain is a component.MATERIALS AND METHODS. Permafrost samples were collected following contamination prevention protocols and used for DNA extraction and microbiological culture on general-purpose nutrient media. The genome of the isolated Staphylococcus hominis strain was sequenced on a DNBSEQ-G50 system (MGI, China) with a read length of 2 × 100 base pairs (bp). Metagenomic sequencing was performed on a DNBSEQ-G400 sequencer with a paired-end read length of 150 bp. Bioinformatic processing of the Staphylococcus hominis strain sequencing results included phylogenomic analysis of the genome sequence and identification of drug resistance and pathogenicity genes. The metagenomic sequencing results were used to identify antibiotic resistance genes.RESULTS. A strain of Staphylococcus hominis 23-2 was isolated. Its genome was identified to contain sequences of the plasmid-localized macrolide antibiotic resistance gene (msr(A)) and the fusidic acid resistance gene (fusB). Phylogenomic analysis indicates that the genome of the studied strain is similar to strains of modern genetic lineages. Seventy-eight unique antibiotic resistance genes were identified in the metagenome structure, including genes encoding efflux pumps of the MFS and RND families, β-lactamase genes, and genes encoding aminoglycoside modification enzymes. The resistome contains sequences of genes encoding resistance to last-line antibiotics: glycopeptides (vanR), carbapenems (metallo-β-lactamases, SHN-1, OXA-60), and polymyxin (pmr).DISCUSSION. The study results demonstrate a high abundance of drug resistance genes in the Arctic permafrost sample from Cape Sopochnaya Karga. Furthermore, the presence of drug resistance genes in the genome structure of Staphylococcus hominis strain 23-2, along with plasmid and prophage sequences, suggests the possibility that this strain may mobilize components of an ancient resistome.CONCLUSION. The study demonstrated the fundamental possibility of preserving staphylococcal strains carrying drug resistance genes and having pathogenic potential in the Neopleistocene permafrost sediments of Cape Sopochnaya Karga. The results obtained justify the feasibility of monitoring the dynamics of the spread of drug resistance genes as an element of regional programs for assessing biological risks associated with climate change in coastal ecosystems of the Arctic.</p></trans-abstract><kwd-group xml:lang="ru"><kwd>морская медицина</kwd><kwd>многолетняя мерзлота</kwd><kwd>палеомикробиология</kwd><kwd>резистом</kwd><kwd>метагеномика</kwd><kwd>Staphylococcus hominis</kwd><kwd>горизонтальный перенос генов</kwd></kwd-group><kwd-group xml:lang="en"><kwd>marine medicine</kwd><kwd>maritime medicine</kwd><kwd>permafrost</kwd><kwd>paleomicrobiology</kwd><kwd>resistome</kwd><kwd>metagenomics</kwd><kwd>Staphylococcus hominis</kwd><kwd>horizontal gene transfer</kwd></kwd-group><funding-group><funding-statement xml:lang="ru">Исследование выполнено в рамках государственного задания Института экспериментальной медицины № 102204110 1032-1-1.6.2 (FGWG-2025-0010).</funding-statement><funding-statement xml:lang="en">The study was carried out as part of the state assignment of the Institute of Experimental Medicine № 102204110 1032-1-1.6.2 (FGWG-2025-0010).</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">Гончаров А. Е., Колоджиева В. В. Медицинская палеомикробиология: проблемы и перспективы // Антибиотики и химиотерапия. 2021. T. 66, № 5–6. С. 72–77 doi: 10.24411/0235-2990-2021-66-5-6-72-77.</mixed-citation><mixed-citation xml:lang="en">Goncharov A. E., Kolodzhieva V. V. Medical paleomicrobiology: problems and prospects. Antibiotics and chemotherapy, 2021, Vol. 66, No. 5–6, pp. 72–77 (In Russ.)  doi: 10.24411/0235-2990-2021-66-5-6-72-77.</mixed-citation></citation-alternatives></ref><ref id="cit2"><label>2</label><citation-alternatives><mixed-citation xml:lang="ru">Saleem M. M., Elahi N., Athar R., Gul A., Adil M., Ellahi A., Kashif H., Hojeij M. The Petri dish under the ice: permafrost pathogens and their impact on global healthcare and antibiotic resistance. Ann Med Surg (Lond), 2024, Vol. 86, No. 12, pp.7193–7201. doi: 10.1097/MS9.0000000000002650.</mixed-citation><mixed-citation xml:lang="en">Saleem M. M., Elahi N., Athar R., Gul A., Adil M., Ellahi A., Kashif H., Hojeij M. The Petri dish under the ice: permafrost pathogens and their impact on global healthcare and antibiotic resistance. Ann Med Surg (Lond), 2024, Vol. 86, No. 12, pp.7193–7201. doi: 10.1097/MS9.0000000000002650.</mixed-citation></citation-alternatives></ref><ref id="cit3"><label>3</label><citation-alternatives><mixed-citation xml:lang="ru">Grigoriev S. E., Fisher D. C., Obadă T., Shirley E. A., Rountrey A. N., Savvinov G. N., Garmaeva D. K., Novgorodov G. P., Cheprasov M. Y., Vasilev S. E., Goncharov A.E., Masharskiy A., Egorova V. E. , Petrova P. P., Egorova E. E., Akhremenko Ya. A., van der Plicht J., Galanin A. A., Fedorov S. E., Ivanov E. V., Tikhonov A. N. A woolly mammoth (Mammuthus primigenius) carcass from Maly Lyakhovsky Island (New Siberian Islands, Russian Federation). Quaternary Int, 2017, Vol. 445, pp. 89–103.</mixed-citation><mixed-citation xml:lang="en">Grigoriev S. E., Fisher D. C., Obadă T., Shirley E. A., Rountrey A. N., Savvinov G. N., Garmaeva D. K., Novgorodov G. P., Cheprasov M. Y., Vasilev S. E., Goncharov A.E., Masharskiy A., Egorova V. E. , Petrova P. P., Egorova E. E., Akhremenko Ya. A., van der Plicht J., Galanin A. A., Fedorov S. E., Ivanov E. V., Tikhonov A. N. A woolly mammoth (Mammuthus primigenius) carcass from Maly Lyakhovsky Island (New Siberian Islands, Russian Federation). Quaternary Int, 2017, Vol. 445, pp. 89–103.</mixed-citation></citation-alternatives></ref><ref id="cit4"><label>4</label><citation-alternatives><mixed-citation xml:lang="ru">Goncharov A., Grigoriev S., Karaseva A., Kolodzhieva V., Azarov D., Akhremenko Y., Tarasova L., Tikhonov A., Masharskiy A., Zueva L., Suvorov A. Draft Genome Sequence of Enterococcus faecium Strain 58m, Isolated from Intestinal Tract Content of a Woolly Mammoth, Mammuthus primigenius. Genome Announc, 2016, Vol. 4, No. 1, pp. e01706–01715. doi:10.1128/genomeA.01706-15.</mixed-citation><mixed-citation xml:lang="en">Goncharov A., Grigoriev S., Karaseva A., Kolodzhieva V., Azarov D., Akhremenko Y., Tarasova L., Tikhonov A., Masharskiy A., Zueva L., Suvorov A. Draft Genome Sequence of Enterococcus faecium Strain 58m, Isolated from Intestinal Tract Content of a Woolly Mammoth, Mammuthus primigenius. Genome Announc, 2016, Vol. 4, No. 1, pp. e01706–01715. doi:10.1128/genomeA.01706-15.</mixed-citation></citation-alternatives></ref><ref id="cit5"><label>5</label><citation-alternatives><mixed-citation xml:lang="ru">Timofeev V., Bahtejeva I., Mironova R., Titareva G., Lev I., Christiany D., Borzilov A., Bogun A., Vergnaud G. Insights from Bacillus anthracis strains isolated from permafrost in the tundra zone of Russia. PLoS One, 2019, Vol. 14, No. 5, pp. e0209140. doi:10.1371/journal.pone.0209140.</mixed-citation><mixed-citation xml:lang="en">Timofeev V., Bahtejeva I., Mironova R., Titareva G., Lev I., Christiany D., Borzilov A., Bogun A., Vergnaud G. Insights from Bacillus anthracis strains isolated from permafrost in the tundra zone of Russia. PLoS One, 2019, Vol. 14, No. 5, pp. e0209140. doi:10.1371/journal.pone.0209140.</mixed-citation></citation-alternatives></ref><ref id="cit6"><label>6</label><citation-alternatives><mixed-citation xml:lang="ru">Liskova E. A., Egorova I. Y., Selyaninov Y. O., Razheva I. V., Gladkova N. A., Toropova N. N., Zakharova O. I., Burova O. A., Surkova G. V., Malkhazova S. M., Korennoy F. I., Iashin I. V., Blokhin A. A. Reindeer Anthrax in the Russian Arctic, 2016: Climatic Determinants of the Outbreak and Vaccination Effectiveness. Front Vet Sci, 2021, Vol. 24, No. 8, pp. 668420. doi: 10.3389/fvets.2021.668420.</mixed-citation><mixed-citation xml:lang="en">Liskova E. A., Egorova I. Y., Selyaninov Y. O., Razheva I. V., Gladkova N. A., Toropova N. N., Zakharova O. I., Burova O. A., Surkova G. V., Malkhazova S. M., Korennoy F. I., Iashin I. V., Blokhin A. A. Reindeer Anthrax in the Russian Arctic, 2016: Climatic Determinants of the Outbreak and Vaccination Effectiveness. Front Vet Sci, 2021, Vol. 24, No. 8, pp. 668420. doi: 10.3389/fvets.2021.668420.</mixed-citation></citation-alternatives></ref><ref id="cit7"><label>7</label><citation-alternatives><mixed-citation xml:lang="ru">Rigou S., Christo-Foroux E., Santini S., Goncharov A., Strauss J., Grosse G., Fedorov A. N., Labadie K., Abergel C., Claverie J. M. Metagenomic survey of the microbiome of ancient Siberian permafrost and modern Kamchatkan cryosols. Microlife, 2022, Vol. 7, No. 3, pp. 003. doi: 10.1093/femsml/uqac003.</mixed-citation><mixed-citation xml:lang="en">Rigou S., Christo-Foroux E., Santini S., Goncharov A., Strauss J., Grosse G., Fedorov A. N., Labadie K., Abergel C., Claverie J. M. Metagenomic survey of the microbiome of ancient Siberian permafrost and modern Kamchatkan cryosols. Microlife, 2022, Vol. 7, No. 3, pp. 003. doi: 10.1093/femsml/uqac003.</mixed-citation></citation-alternatives></ref><ref id="cit8"><label>8</label><citation-alternatives><mixed-citation xml:lang="ru">Гусев Е. А., Молодьков А. Н., Деревянко Л. Г. Cопкаргинский мамонт, время и условия его обитания (cевер Западной Сибири) // Успехи современного естествознания. 2015. № 1–3. С. 432–435</mixed-citation><mixed-citation xml:lang="en">Gusev E. A., Molodkov A. N., Derevyanko L. G. The Sopkarginsky Mammoth, Its Time and Habitat (Northern West Siberia). Advances in Modern Natural Sciences, 2015., No. 1-3, pp. 432-435 (In Russ.)</mixed-citation></citation-alternatives></ref><ref id="cit9"><label>9</label><citation-alternatives><mixed-citation xml:lang="ru">Pitulko V. V., Tikhonov A. N., Pavlova E. Y., Nikolskiy P. A., Kuper K. E., Polozov R. N. Paleoanthropology. Early human presence in the Arctic: Evidence from 45,000-year-old mammoth remains. Science, 2016, Vol. 351, No. 6270, pp. 260–263. doi: 10.1126/science.aad0554.</mixed-citation><mixed-citation xml:lang="en">Pitulko V. V., Tikhonov A. N., Pavlova E. Y., Nikolskiy P. A., Kuper K. E., Polozov R. N. Paleoanthropology. Early human presence in the Arctic: Evidence from 45,000-year-old mammoth remains. Science, 2016, Vol. 351, No. 6270, pp. 260–263. doi: 10.1126/science.aad0554.</mixed-citation></citation-alternatives></ref><ref id="cit10"><label>10</label><citation-alternatives><mixed-citation xml:lang="ru">Zhang L., Thomas J. C., Miragaia M., Bouchami O., Chaves F., d’Azevedo P. A., Aanensen D. M., de Lencastre H., Gray B. M., Robinson D. A. Multilocus sequence typing and further genetic characterization of the enigmatic pathogen, Staphylococcus hominis. PLoS One, 2013, Vol. 8, No. 6, pp. e66496. doi: 10.1371/journal.pone.0066496.</mixed-citation><mixed-citation xml:lang="en">Zhang L., Thomas J. C., Miragaia M., Bouchami O., Chaves F., d’Azevedo P. A., Aanensen D. M., de Lencastre H., Gray B. M., Robinson D. A. Multilocus sequence typing and further genetic characterization of the enigmatic pathogen, Staphylococcus hominis. PLoS One, 2013, Vol. 8, No. 6, pp. e66496. doi: 10.1371/journal.pone.0066496.</mixed-citation></citation-alternatives></ref><ref id="cit11"><label>11</label><citation-alternatives><mixed-citation xml:lang="ru">Kashuba E., Dmitriev A. A., Kamal S. M., Melefors O., Griva G., Römling U., Ernberg I., Kashuba V., Brouchkov A. Ancient permafrost staphylococci carry antibiotic resistance genes. Microb Ecol Health Dis, 2017, Vol. 28, No. 1, pp. 1345574. doi: 10.1080/16512235.2017.1345574.</mixed-citation><mixed-citation xml:lang="en">Kashuba E., Dmitriev A. A., Kamal S. M., Melefors O., Griva G., Römling U., Ernberg I., Kashuba V., Brouchkov A. Ancient permafrost staphylococci carry antibiotic resistance genes. Microb Ecol Health Dis, 2017, Vol. 28, No. 1, pp. 1345574. doi: 10.1080/16512235.2017.1345574.</mixed-citation></citation-alternatives></ref><ref id="cit12"><label>12</label><citation-alternatives><mixed-citation xml:lang="ru">Sannino C., Qi W., Rüthi J., Stierli B., Frey B. Distinct taxonomic and functional profiles of high Arctic and alpine permafrost-affected soil microbiomes. Environ Microbiome, 2023, Vol. 18, No. 1, pp. 54. doi: 10.1186/s40793-023-00509-6.</mixed-citation><mixed-citation xml:lang="en">Sannino C., Qi W., Rüthi J., Stierli B., Frey B. Distinct taxonomic and functional profiles of high Arctic and alpine permafrost-affected soil microbiomes. Environ Microbiome, 2023, Vol. 18, No. 1, pp. 54. doi: 10.1186/s40793-023-00509-6.</mixed-citation></citation-alternatives></ref><ref id="cit13"><label>13</label><citation-alternatives><mixed-citation xml:lang="ru">Marimuthu K. Isolation and characterization of Staphylococcus hominis JX961712 from oil contaminated soil. Journal of pharmacy research, 2013, Vol. 7, No. 3, pp. 252–256.</mixed-citation><mixed-citation xml:lang="en">Marimuthu K. Isolation and characterization of Staphylococcus hominis JX961712 from oil contaminated soil. Journal of pharmacy research, 2013, Vol. 7, No. 3, pp. 252–256.</mixed-citation></citation-alternatives></ref><ref id="cit14"><label>14</label><citation-alternatives><mixed-citation xml:lang="ru">Aftab Uddin M., Akter S., Ferdous M., Haidar B., Amin A., Shofiul Islam Molla A. H. M., Khan H., Islam M. R. A plant endophyte Staphylococcus hominis strain MBL_AB63 produces a novel lantibiotic, homicorcin and a position one variant. Sci Rep, 2021, Vol. 11, No. 1, pp. 11211. doi: 10.1038/s41598-021-90613-9.</mixed-citation><mixed-citation xml:lang="en">Aftab Uddin M., Akter S., Ferdous M., Haidar B., Amin A., Shofiul Islam Molla A. H. M., Khan H., Islam M. R. A plant endophyte Staphylococcus hominis strain MBL_AB63 produces a novel lantibiotic, homicorcin and a position one variant. Sci Rep, 2021, Vol. 11, No. 1, pp. 11211. doi: 10.1038/s41598-021-90613-9.</mixed-citation></citation-alternatives></ref><ref id="cit15"><label>15</label><citation-alternatives><mixed-citation xml:lang="ru">Rakitin A. L., Ermakova A. Y., Beletsky A. V., Petrova M., Mardanov A. V., Ravin N. V. Genome Analysis of Acinetobacter lwoffii Strains Isolated from Permafrost Soils Aged from 15 Thousand to 1.8 Million Years Revealed Their Close Relationships with Present-Day Environmental and Clinical Isolates. Biology (Basel), 2021, Vol. 10, No. 9, pp. 871. doi: 10.3390/biology10090871.</mixed-citation><mixed-citation xml:lang="en">Rakitin A. L., Ermakova A. Y., Beletsky A. V., Petrova M., Mardanov A. V., Ravin N. V. Genome Analysis of Acinetobacter lwoffii Strains Isolated from Permafrost Soils Aged from 15 Thousand to 1.8 Million Years Revealed Their Close Relationships with Present-Day Environmental and Clinical Isolates. Biology (Basel), 2021, Vol. 10, No. 9, pp. 871. doi: 10.3390/biology10090871.</mixed-citation></citation-alternatives></ref><ref id="cit16"><label>16</label><citation-alternatives><mixed-citation xml:lang="ru">Goncharov A., Grigorjev S., Karaseva A., Kolodzhieva V., Azarov D., Akhremenko Y., Tarasova L., Tikhonov A., Masharskiy A., Zueva L., Suvorov A. Draft Genome Sequence of Enterococcus faecium Strain 58m, Isolated from Intestinal Tract Content of a Woolly Mammoth, Mammuthus primigenius. Genome Announc, 2016, Vol. 4, No. 1, pp. e01706-15. doi:10.1128/genomeA.01706-15.</mixed-citation><mixed-citation xml:lang="en">Goncharov A., Grigorjev S., Karaseva A., Kolodzhieva V., Azarov D., Akhremenko Y., Tarasova L., Tikhonov A., Masharskiy A., Zueva L., Suvorov A. Draft Genome Sequence of Enterococcus faecium Strain 58m, Isolated from Intestinal Tract Content of a Woolly Mammoth, Mammuthus primigenius. Genome Announc, 2016, Vol. 4, No. 1, pp. e01706-15. doi:10.1128/genomeA.01706-15.</mixed-citation></citation-alternatives></ref><ref id="cit17"><label>17</label><citation-alternatives><mixed-citation xml:lang="ru">Goncharov A., Grigorjev S., Karaseva A., Kolodzhieva V., Azarov D., Akhremenko Y., Tarasova L., Tikhonov A., Masharskiy A., Zueva L., Suvorov A. Reply to “Isolating Viable Ancient Bacteria: What You Put In Is What You Get Out”. Genome Announc, 2016, Vol. 4, No. 4, pp. e00734-16. doi: 10.1128/genomeA.00734-16.</mixed-citation><mixed-citation xml:lang="en">Goncharov A., Grigorjev S., Karaseva A., Kolodzhieva V., Azarov D., Akhremenko Y., Tarasova L., Tikhonov A., Masharskiy A., Zueva L., Suvorov A. Reply to “Isolating Viable Ancient Bacteria: What You Put In Is What You Get Out”. Genome Announc, 2016, Vol. 4, No. 4, pp. e00734-16. doi: 10.1128/genomeA.00734-16.</mixed-citation></citation-alternatives></ref><ref id="cit18"><label>18</label><citation-alternatives><mixed-citation xml:lang="ru">Rigou S., Christo-Foroux E., Santini S., Goncharov A., Strauss J., Grosse G., Fedorov A. N., Labadie K., Abergel C., Claverie J. M. Metagenomic survey of the microbiome of ancient Siberian permafrost and modern Kamchatkan cryosols. Microlife, 2022, Vol. 7, No. 3, pp. 003. doi: 10.1093/femsml/uqac003.</mixed-citation><mixed-citation xml:lang="en">Rigou S., Christo-Foroux E., Santini S., Goncharov A., Strauss J., Grosse G., Fedorov A. N., Labadie K., Abergel C., Claverie J. M. Metagenomic survey of the microbiome of ancient Siberian permafrost and modern Kamchatkan cryosols. Microlife, 2022, Vol. 7, No. 3, pp. 003. doi: 10.1093/femsml/uqac003.</mixed-citation></citation-alternatives></ref><ref id="cit19"><label>19</label><citation-alternatives><mixed-citation xml:lang="ru">Муратов И. Н., Ибраева А. Ш., Тимергазина Л. Л., Полищук Ю. М. Дистанционное исследование термокарстовых озер арктической тундры Таймыра. Вестник Югорского государственного университета. 2021. № 17(1). C. 62–71doi: 10.17816/byusu20210162-71.</mixed-citation><mixed-citation xml:lang="en">Muratov I. N., Ibraeva A. Sh., Timergazina L. L., Polischuk Yu. M. Remote Study of Thermokarst Lakes in the Arctic Tundra of Taimyr. Bulletin of Yugra State University, 2021, Vol. 17, No. 1, pp. 62–71 (In Russ.) doi: 10.17816/byusu20210162-71.</mixed-citation></citation-alternatives></ref><ref id="cit20"><label>20</label><citation-alternatives><mixed-citation xml:lang="ru">Haan T. J., Drown D. M. Unearthing Antibiotic Resistance Associated with Disturbance-Induced Permafrost Thaw in Interior Alaska. Microorganisms, 2021, Vol. 9, No. 1, pp. 116. doi: 10.3390/microorganisms9010116.</mixed-citation><mixed-citation xml:lang="en">Haan T. J., Drown D. M. Unearthing Antibiotic Resistance Associated with Disturbance-Induced Permafrost Thaw in Interior Alaska. Microorganisms, 2021, Vol. 9, No. 1, pp. 116. doi: 10.3390/microorganisms9010116.</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>
