<?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">vavilov</journal-id><journal-title-group><journal-title xml:lang="ru">Вавиловский журнал генетики и селекции</journal-title><trans-title-group xml:lang="en"><trans-title>Vavilov Journal of Genetics and Breeding</trans-title></trans-title-group></journal-title-group><issn pub-type="epub">2500-3259</issn><publisher><publisher-name>Institute of Cytology and Genetics of Siberian Branch of the RAS</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="doi">10.18699/vjgb-25-115</article-id><article-id custom-type="elpub" pub-id-type="custom">vavilov-4893</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>ЭКОЛОГИЧЕСКАЯ И ПОПУЛЯЦИОННАЯ ГЕНЕТИКА</subject></subj-group><subj-group subj-group-type="section-heading" xml:lang="en"><subject>ECOLOGICAL AND POPULATION GENETICS</subject></subj-group></article-categories><title-group><article-title>Компьютерное моделирование пространственной динамики и первичной генетической дивергенции в системе популяций на кольцевом ареале</article-title><trans-title-group xml:lang="en"><trans-title>Computer modeling of spatial dynamics and primary genetic divergence for a population system in a ring areal</trans-title></trans-title-group></title-group><contrib-group><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-7060-2731</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Кулаков</surname><given-names>М. П.</given-names></name><name name-style="western" xml:lang="en"><surname>Kulakov</surname><given-names>M. P.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Биробиджан</p></bio><bio xml:lang="en"><p>Birobidzhan</p></bio><email xlink:type="simple">k_matvey@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/0000-0002-3090-986X</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Жданова</surname><given-names>О. Л.</given-names></name><name name-style="western" xml:lang="en"><surname>Zhdanova</surname><given-names>O. L.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Владивосток</p></bio><bio xml:lang="en"><p>Vladivostok</p></bio><xref ref-type="aff" rid="aff-2"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0003-1629-2610</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Фрисман</surname><given-names>Е. Я.</given-names></name><name name-style="western" xml:lang="en"><surname>Frisman</surname><given-names>E. Ya.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Биробиджан</p></bio><bio xml:lang="en"><p>Birobidzhan</p></bio><xref ref-type="aff" rid="aff-1"/></contrib></contrib-group><aff-alternatives id="aff-1"><aff xml:lang="ru">Институт комплексного анализа региональных проблем Дальневосточного отделения Российской академии наук<country>Россия</country></aff><aff xml:lang="en">Institute for Complex Analysis of Regional Problems of the Far Eastern Branch of the Russian Academy of Sciences<country>Russian Federation</country></aff></aff-alternatives><aff-alternatives id="aff-2"><aff xml:lang="ru">Институт автоматики и процессов управления Дальневосточного отделения Российской академии наук<country>Россия</country></aff><aff xml:lang="en">Institute of Automation and Control Processes of the Far Eastern Branch of the Russian Academy of Sciences<country>Russian Federation</country></aff></aff-alternatives><pub-date pub-type="collection"><year>2025</year></pub-date><pub-date pub-type="epub"><day>12</day><month>12</month><year>2025</year></pub-date><volume>29</volume><issue>7</issue><fpage>1109</fpage><lpage>1121</lpage><permissions><copyright-statement>Copyright &amp;#x00A9; Кулаков М.П., Жданова О.Л., Фрисман Е.Я., 2025</copyright-statement><copyright-year>2025</copyright-year><copyright-holder xml:lang="ru">Кулаков М.П., Жданова О.Л., Фрисман Е.Я.</copyright-holder><copyright-holder xml:lang="en">Kulakov M.P., Zhdanova O.L., Frisman E.Y.</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://vavilov.elpub.ru/jour/article/view/4893">https://vavilov.elpub.ru/jour/article/view/4893</self-uri><abstract><p>   Одна из ключевых задач современной эволюционной биологии – изучение процессов, приводящих к первичному разделению (дивергенции) популяций на различающиеся генотипами группы особей.</p><p>   Для дивергенции очевидно необходима репродуктивная изоляция, которая делает невозможным контакт особей или существенно затрудняет обмен генетической информацией между популяциями. Настоящее исследование изучает возможность изоляции, в основе которой лежат не очевидные географические барьеры, удаленность популяций или экологическая специализация, а лишь наследственные механизмы, дрейф и поток генов, а также отбор против гетерозигот. Для этого предложена и исследована динамическая модель с дискретным временем, которая описывает динамику частот и численностей в системе миграционно связанных лимитированных популяций. Рассматривается панмиктичная популяция с менделевскими правилами наследования, монолокусным отбором, действием плотностно-зависимых факторов, лимитирующих рост численности. Особи свободно скрещиваются и перемещаются вдоль одномерного кольцевого ареала. Модель верифицирована с использованием данных эксперимента над ящичной системой популяций дрозофил Drosophila melanogaster, проведенного под руководством Ю. П. Алтухова. При достаточно простых предположениях модель описывает некоторые механизмы возникновения и сохранения на однородном ареале существенных генетических различий (первичной генетической дивергенции), сопровождаемых неоднородностью в частотах аллелей и численностях. В этом случае формируется несколько больших групп генетически однородных субпопуляций, развивающихся независимо. В местах их контакта активно идет гибридизация, а полиморфизм сохраняется за счет миграции с сопредельных однородных участков. Обнаружено, что устойчиво поддерживать такое пространственное распределение может только дизруптивный (разрывающий) отбор, направленный против гетерозигот. При движущем отборе дивергенция существует непродолжительное время как часть переходного процесса. За счет пониженной приспособленности гетерозигот (гибридов) и низкой скорости роста на этих участках (зонах гибридизации) существенно затрудняется обмен генами между смежными участками с противоположными гомозиготными генотипами (фенотипами). В результате зоны гибридизации выполняют функцию географического барьера, который фактически останавливает обмен генов между разными группами в случае смежной симпатрии.</p></abstract><trans-abstract xml:lang="en"><p>   One of the main goals of modern evolutionary biology is to understand the mechanisms that lead to the initial differentiation (primary divergence) of populations into groups with genetic traits.</p><p>   This divergence requires reproductive isolation, which prevents or hinders contact and the exchange of genetic material between populations. This study explores the potential for isolation based not on obvious geographical barriers, population distance, or ecological specialization, but rather on hereditary mechanisms, such as gene drift and flow and selection against heterozygous individuals. To this end, we propose and investigate a dynamic discrete-time model that describes the dynamics of frequencies and numbers in a system of limited populations coupled by migrations. We consider a panmictic population with Mendelian inheritance rules, one-locus selection, and density-dependent factors limiting population growth. Individuals freely mate and randomly move around a one-dimensional ring-shaped habitat. The model was verified using data from an experiment on the box population system of Drosophila melanogaster performed by Yu.P. Altukhov et al. With rather simple assumptions, the model explains some mechanisms for the emergence and preservation of significant genetic differences between subpopulations (primary genetic divergence), accompanied by heterogeneity in allele frequencies and abundances within a homogeneous area. In this scenario, several large groups of genetically homogeneous subpopulations form and independently develop. Hybridization occurs at contact sites, and polymorphism is maintained through migration from genetically homogeneous nearby sites. It was found that only disruptive selection, directed against heterozygous individuals, can sustainably maintain such a spatial distribution. Under directional selection, divergence may occur for a short time as part of the transitional evolutionary process towards the best-adapted genotype. Because of the reduced adaptability of heterozygous (hybrid) individuals and low growth rates in these sites (hybrid zones), gene flow between adjacent sites with opposite genotypes (phenotypes) is significantly impeded. As a result, the hybrid zones can become effective geographical barriers that prevent the genetic flow between coupled subpopulations.</p></trans-abstract><kwd-group xml:lang="ru"><kwd>метапопуляция</kwd><kwd>миграция</kwd><kwd>пространственно-временная динамика</kwd><kwd>математическое моделирование</kwd><kwd>генетическая дивергенция</kwd><kwd>поток генов</kwd><kwd>гибридные зоны</kwd><kwd>изоляция</kwd></kwd-group><kwd-group xml:lang="en"><kwd>metapopulation</kwd><kwd>migration</kwd><kwd>spatiotemporal dynamics</kwd><kwd>mathematical modeling</kwd><kwd>genetic divergence</kwd><kwd>gene flow</kwd><kwd>hybrid zones</kwd><kwd>isolation</kwd></kwd-group><funding-group xml:lang="en"><funding-statement>This work was carried out within the framework of the state task of the Institute for Complex Analysis of Regional Problems of the Far Eastern Branch of the Russian Academy of Sciences</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">Aguillon S.M., Rohwer V.G. Revisiting a classic hybrid zone: movement of the northern flicker hybrid zone in contemporary times. Evolution. 2022;76(5):1082­1090. doi: 10.1111/evo.14474</mixed-citation><mixed-citation xml:lang="en">Aguillon S.M., Rohwer V.G. Revisiting a classic hybrid zone: movement of the northern flicker hybrid zone in contemporary times. Evolution. 2022;76(5):1082­1090. doi: 10.1111/evo.14474</mixed-citation></citation-alternatives></ref><ref id="cit2"><label>2</label><citation-alternatives><mixed-citation xml:lang="ru">Allee W.C. The Social Life of Animals. Beacon Press, 1958</mixed-citation><mixed-citation xml:lang="en">Allee W.C. The Social Life of Animals. Beacon Press, 1958</mixed-citation></citation-alternatives></ref><ref id="cit3"><label>3</label><citation-alternatives><mixed-citation xml:lang="ru">Altukhov Yu.P. Genetic Processes in Populations. Moscow: Akadem kniga Publ., 2003 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Altukhov Yu.P. Genetic Processes in Populations. Moscow: Akadem kniga Publ., 2003 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit4"><label>4</label><citation-alternatives><mixed-citation xml:lang="ru">Altukhov Yu.P., Bernashevskaya A.G. Experimental modeling of genetic processes in a population system of Drosophila melanogaster corresponding to a circular stepping­stone model: 2. Stability of al­lelic composition and periodic relationship of allele frequency with distance. Soviet Genetics. 1981;17(6):1052­1059 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Altukhov Yu.P., Bernashevskaya A.G. Experimental modeling of genetic processes in a population system of Drosophila melanogaster corresponding to a circular stepping­stone model: 2. Stability of al­lelic composition and periodic relationship of allele frequency with distance. Soviet Genetics. 1981;17(6):1052­1059 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit5"><label>5</label><citation-alternatives><mixed-citation xml:lang="ru">Altukhov Yu.P., Bernashevskaya A.G., Milishnikov A.N. Experimen­tal modeling of genetic processes in the population system of Drosophila melanogaster corresponding to the ring step model. Soviet Genetics. 1979;15(4):646­655 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Altukhov Yu.P., Bernashevskaya A.G., Milishnikov A.N. Experimen­tal modeling of genetic processes in the population system of Drosophila melanogaster corresponding to the ring step model. Soviet Genetics. 1979;15(4):646­655 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit6"><label>6</label><citation-alternatives><mixed-citation xml:lang="ru">Bazykin A.D. Reduced fitness of heterozygotes in a system of adjacent populations. Soviet Genetics. 1972;8(11):155­161 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Bazykin A.D. Reduced fitness of heterozygotes in a system of adjacent populations. Soviet Genetics. 1972;8(11):155­161 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit7"><label>7</label><citation-alternatives><mixed-citation xml:lang="ru">Blair W.F. Mating call and stage of speciation in the Microhyla olivacea–M. carolinensis complex. Evolution. 1955a;9(4):469­480. doi: 10.1111/j.1558­5646.1955.tb01556</mixed-citation><mixed-citation xml:lang="en">Blair W.F. Mating call and stage of speciation in the Microhyla olivacea–M. carolinensis complex. Evolution. 1955a;9(4):469­480. doi: 10.1111/j.1558­5646.1955.tb01556</mixed-citation></citation-alternatives></ref><ref id="cit8"><label>8</label><citation-alternatives><mixed-citation xml:lang="ru">Blair W.F. Size difference as a possible isolation mechanism in Micro­hyla. Am Nat. 1955b;89(848):297­301. doi: 10.1086/281894</mixed-citation><mixed-citation xml:lang="en">Blair W.F. Size difference as a possible isolation mechanism in Micro­hyla. Am Nat. 1955b;89(848):297­301. doi: 10.1086/281894</mixed-citation></citation-alternatives></ref><ref id="cit9"><label>9</label><citation-alternatives><mixed-citation xml:lang="ru">Blinov V.N., Zheleznova T.K. Black Corvus corone and grey C. cornix crows: controversial issues about status (races, semispecies or species?), origin (allo­ or sympatric?) and the phenomenon of stable hybrid zones. Russkiy Ornitologicheskiy Zhurnal = Russian Ornithological Journal. 2020;29(1958):3596­3601 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Blinov V.N., Zheleznova T.K. Black Corvus corone and grey C. cornix crows: controversial issues about status (races, semispecies or species?), origin (allo­ or sympatric?) and the phenomenon of stable hybrid zones. Russkiy Ornitologicheskiy Zhurnal = Russian Ornithological Journal. 2020;29(1958):3596­3601 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit10"><label>10</label><citation-alternatives><mixed-citation xml:lang="ru">Dey S., Joshi A. Stability via asynchrony in Drosophila metapopulations with low migration rates. Science. 2006;312(5772):434­436. doi: 10.1126/science.1125317</mixed-citation><mixed-citation xml:lang="en">Dey S., Joshi A. Stability via asynchrony in Drosophila metapopulations with low migration rates. Science. 2006;312(5772):434­436. doi: 10.1126/science.1125317</mixed-citation></citation-alternatives></ref><ref id="cit11"><label>11</label><citation-alternatives><mixed-citation xml:lang="ru">Filchak K., Roethele J., Feder J. Natural selection and sympatric divergence in the apple maggot Rhagoletis pomonella. Nature. 2000; 407(6805):739­742. doi: 10.1038/35037578</mixed-citation><mixed-citation xml:lang="en">Filchak K., Roethele J., Feder J. Natural selection and sympatric divergence in the apple maggot Rhagoletis pomonella. Nature. 2000; 407(6805):739­742. doi: 10.1038/35037578</mixed-citation></citation-alternatives></ref><ref id="cit12"><label>12</label><citation-alternatives><mixed-citation xml:lang="ru">Frisman E.Y. Primary Genetic Divergence (Theoretical analysis and modeling). Vladivostok, 1986 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Frisman E.Y. Primary Genetic Divergence (Theoretical analysis and modeling). Vladivostok, 1986 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit13"><label>13</label><citation-alternatives><mixed-citation xml:lang="ru">Haring E., Däubl B., Pinsker W., Kryukov A., Gamauf A. Genetic divergences and intraspecific variation in corvids of the genus Corvus (Aves: Passeriformes: Corvidae) – a first survey based on museum specimens. J Zool Syst Evol Res. 2012;50(3):230­246. doi: 10.1111/j.1439­0469.2012.00664.x</mixed-citation><mixed-citation xml:lang="en">Haring E., Däubl B., Pinsker W., Kryukov A., Gamauf A. Genetic divergences and intraspecific variation in corvids of the genus Corvus (Aves: Passeriformes: Corvidae) – a first survey based on museum specimens. J Zool Syst Evol Res. 2012;50(3):230­246. doi: 10.1111/j.1439­0469.2012.00664.x</mixed-citation></citation-alternatives></ref><ref id="cit14"><label>14</label><citation-alternatives><mixed-citation xml:lang="ru">Kapitonova L.V., Formozov N.A., Fedorov V.V., Kerimov A.B., Seliva­nova D.S. Peculiarities of behavior and ecology of the Great tit Parus major Linneus, 1758 and Japanese tit P. minor Temmink et Schlegel, 1848 as possible factors of maintaining the stability of speciesspecific phenotypes in the area of sympatry and local hybridi zation in the Amur Region. Dal’nevostochnyy Ornitologicheskiy Zhurnal = Far Eastern Journal of Ornithology. 2012;3:37­46 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Kapitonova L.V., Formozov N.A., Fedorov V.V., Kerimov A.B., Seliva­nova D.S. Peculiarities of behavior and ecology of the Great tit Parus major Linneus, 1758 and Japanese tit P. minor Temmink et Schlegel, 1848 as possible factors of maintaining the stability of speciesspecific phenotypes in the area of sympatry and local hybridi zation in the Amur Region. Dal’nevostochnyy Ornitologicheskiy Zhurnal = Far Eastern Journal of Ornithology. 2012;3:37­46 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit15"><label>15</label><citation-alternatives><mixed-citation xml:lang="ru">Keymer J.E., Galajda P., Muldoon C., Park S., Austin R.H. Bacterial metapopulations in nanofabricated landscapes. Proc Natl Acad Sci USA. 2006;103(46):17290­17295. doi: 10.1073/pnas.0607971103</mixed-citation><mixed-citation xml:lang="en">Keymer J.E., Galajda P., Muldoon C., Park S., Austin R.H. Bacterial metapopulations in nanofabricated landscapes. Proc Natl Acad Sci USA. 2006;103(46):17290­17295. doi: 10.1073/pnas.0607971103</mixed-citation></citation-alternatives></ref><ref id="cit16"><label>16</label><citation-alternatives><mixed-citation xml:lang="ru">Kryukov A.P. Phylogeography and hybridization of corvid birds in the Palearctic Region. Vavilov J Genet Breed. 2019;23(2):232­238. doi: 10.18699/VJ19.487</mixed-citation><mixed-citation xml:lang="en">Kryukov A.P. Phylogeography and hybridization of corvid birds in the Palearctic Region. Vavilov J Genet Breed. 2019;23(2):232­238. doi: 10.18699/VJ19.487</mixed-citation></citation-alternatives></ref><ref id="cit17"><label>17</label><citation-alternatives><mixed-citation xml:lang="ru">Kulakov M., Frisman E.Ya. Primary genetic divergence in a system of limited population coupled by migration in a ring habitat. Маthematical Biology and Bioinformatics. 2025;20(1):1­30. doi: 10.17537/2025.20.1 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Kulakov M., Frisman E.Ya. Primary genetic divergence in a system of limited population coupled by migration in a ring habitat. Маthematical Biology and Bioinformatics. 2025;20(1):1­30. doi: 10.17537/2025.20.1 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit18"><label>18</label><citation-alternatives><mixed-citation xml:lang="ru">Láruson Á.J., Reed F.A. Stability of underdominant genetic polymorphisms in population networks. J Theor Biol. 2016;390:156­163. doi: 10.1016/j.jtbi.2015.11.023</mixed-citation><mixed-citation xml:lang="en">Láruson Á.J., Reed F.A. Stability of underdominant genetic polymorphisms in population networks. J Theor Biol. 2016;390:156­163. doi: 10.1016/j.jtbi.2015.11.023</mixed-citation></citation-alternatives></ref><ref id="cit19"><label>19</label><citation-alternatives><mixed-citation xml:lang="ru">Littlejohn M.J. Premating isolation in the Hyla ewingi complex (Anura: Hylidae). Evolution. 1965;19(2):234­243. doi: 10.2307/2406376</mixed-citation><mixed-citation xml:lang="en">Littlejohn M.J. Premating isolation in the Hyla ewingi complex (Anura: Hylidae). Evolution. 1965;19(2):234­243. doi: 10.2307/2406376</mixed-citation></citation-alternatives></ref><ref id="cit20"><label>20</label><citation-alternatives><mixed-citation xml:lang="ru">Matsumoto M., Nishimura T. Mersenne twister: a 623­dimensionally equidistributed uniform pseudorandom number generator. ACM Trans Model Comput Simul. 1998;8(1):3­30. doi: 10.1145/272991.272995</mixed-citation><mixed-citation xml:lang="en">Matsumoto M., Nishimura T. Mersenne twister: a 623­dimensionally equidistributed uniform pseudorandom number generator. ACM Trans Model Comput Simul. 1998;8(1):3­30. doi: 10.1145/272991.272995</mixed-citation></citation-alternatives></ref><ref id="cit21"><label>21</label><citation-alternatives><mixed-citation xml:lang="ru">Murphy M.A., Dezzani R., Pilliod D.S., Storfer A. Landscape genet­ics of high mountain frogmetapopulations. Mol Ecol. 2010;19(17): 3634­3649. doi: 10.1111/j.1365­294X.2010.04723.x</mixed-citation><mixed-citation xml:lang="en">Murphy M.A., Dezzani R., Pilliod D.S., Storfer A. Landscape genet­ics of high mountain frogmetapopulations. Mol Ecol. 2010;19(17): 3634­3649. doi: 10.1111/j.1365­294X.2010.04723.x</mixed-citation></citation-alternatives></ref><ref id="cit22"><label>22</label><citation-alternatives><mixed-citation xml:lang="ru">Orsini L., Corander J., Alasentie A., Hanski I. Genetic spatial structure in a butterfly metapopulation correlates better with past than present demographic structure. Mol Ecol. 2008;17(11):2629­2642. doi: 10.1111/j.1365­294X.2008.03782.x</mixed-citation><mixed-citation xml:lang="en">Orsini L., Corander J., Alasentie A., Hanski I. Genetic spatial structure in a butterfly metapopulation correlates better with past than present demographic structure. Mol Ecol. 2008;17(11):2629­2642. doi: 10.1111/j.1365­294X.2008.03782.x</mixed-citation></citation-alternatives></ref><ref id="cit23"><label>23</label><citation-alternatives><mixed-citation xml:lang="ru">Poelstra J.W., Vijay N., Bossu C.M., Lantz H., Ryll B., Müller I., Baglione V., Unneberg P., Wikelski M., Grabherr M.G., Wolf J.B.W. The genomic landscape underlying phenotypic integrity in the face of gene flow in crows. Science. 2014;344(6190):1410­1414. doi: 10.1126/science.1253226</mixed-citation><mixed-citation xml:lang="en">Poelstra J.W., Vijay N., Bossu C.M., Lantz H., Ryll B., Müller I., Baglione V., Unneberg P., Wikelski M., Grabherr M.G., Wolf J.B.W. The genomic landscape underlying phenotypic integrity in the face of gene flow in crows. Science. 2014;344(6190):1410­1414. doi: 10.1126/science.1253226</mixed-citation></citation-alternatives></ref><ref id="cit24"><label>24</label><citation-alternatives><mixed-citation xml:lang="ru">Smith M.J., Osborne W., Hunter D. Geographic variation in the advertisement call structure of Litoria verreauxii (Anura: Hylidae). Copeia. 2003;4:750­758. doi: 10.1643/HA02­133.1</mixed-citation><mixed-citation xml:lang="en">Smith M.J., Osborne W., Hunter D. Geographic variation in the advertisement call structure of Litoria verreauxii (Anura: Hylidae). Copeia. 2003;4:750­758. doi: 10.1643/HA02­133.1</mixed-citation></citation-alternatives></ref><ref id="cit25"><label>25</label><citation-alternatives><mixed-citation xml:lang="ru">Sundqvist L., Keenan K., Zackrisson M., Prodöhl P., Kleinhans D. Directional genetic differentiation and relative migration. Ecol Evol. 2016;6(11):3461­3475. doi: 10.1002/ece3.2096</mixed-citation><mixed-citation xml:lang="en">Sundqvist L., Keenan K., Zackrisson M., Prodöhl P., Kleinhans D. Directional genetic differentiation and relative migration. Ecol Evol. 2016;6(11):3461­3475. doi: 10.1002/ece3.2096</mixed-citation></citation-alternatives></ref><ref id="cit26"><label>26</label><citation-alternatives><mixed-citation xml:lang="ru">Tait C., Kharva H., Schubert M., Kritsch D., Sombke A., Rybak J., Feder J.L., Olsson S.B. A reversal in sensory processing accom­panies ongoing ecological divergence and speciation in Rhagoletis pomonella. Proc Biol Sci. 2021;288(1947):20210192. doi: 10.1098/rspb.2021.0192</mixed-citation><mixed-citation xml:lang="en">Tait C., Kharva H., Schubert M., Kritsch D., Sombke A., Rybak J., Feder J.L., Olsson S.B. A reversal in sensory processing accom­panies ongoing ecological divergence and speciation in Rhagoletis pomonella. Proc Biol Sci. 2021;288(1947):20210192. doi: 10.1098/rspb.2021.0192</mixed-citation></citation-alternatives></ref><ref id="cit27"><label>27</label><citation-alternatives><mixed-citation xml:lang="ru">Yeaman S., Otto S.P. Establishment and maintenance of adaptive genetic divergence under migration, selection, and drift. Evolution. 2011;65(7):2123­2129. doi: 10.1111/j.1558­5646.2011.01277.x</mixed-citation><mixed-citation xml:lang="en">Yeaman S., Otto S.P. Establishment and maintenance of adaptive genetic divergence under migration, selection, and drift. Evolution. 2011;65(7):2123­2129. doi: 10.1111/j.1558­5646.2011.01277.x</mixed-citation></citation-alternatives></ref><ref id="cit28"><label>28</label><citation-alternatives><mixed-citation xml:lang="ru">Yee W.L., Goughnour R.B. Mating frequencies and production of hy­brids by Rhagoletis pomonella and Rhagoletis zephyria (Diptera: Tephritidae) in the laboratory. Can Entomol. 2011;143(1):82­90. doi: 10.4039/n10­047</mixed-citation><mixed-citation xml:lang="en">Yee W.L., Goughnour R.B. Mating frequencies and production of hy­brids by Rhagoletis pomonella and Rhagoletis zephyria (Diptera: Tephritidae) in the laboratory. Can Entomol. 2011;143(1):82­90. doi: 10.4039/n10­047</mixed-citation></citation-alternatives></ref><ref id="cit29"><label>29</label><citation-alternatives><mixed-citation xml:lang="ru">Zhdanova O.L., Frisman E.Y. On the genetic divergence of migration-coupled populations: modern modeling based on the experimental results of Yu.P. Altukhov et al. Russ J Genet. 2023;59:614­622. doi: 10.1134/S1022795423060133</mixed-citation><mixed-citation xml:lang="en">Zhdanova O.L., Frisman E.Y. On the genetic divergence of migration-coupled populations: modern modeling based on the experimental results of Yu.P. Altukhov et al. Russ J Genet. 2023;59:614­622. doi: 10.1134/S1022795423060133</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>
