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The evolution of hybrid male sterility, or Haldane\u2019s rule, typically characterizes later stages of species formation, when reproductive isolation is strongest. Yet, understanding how quickly reproductive barriers evolve and their consequences for maintaining genetic boundaries between emerging species remains a challenging task because it requires studying taxa that hybridize in nature. Here, we address these questions using the meadow grasshopper<jats:italic>Pseudochorthippus parallelus<\/jats:italic>, where populations that show multiple reproductive barriers, including hybrid male sterility, hybridize in two natural hybrid zones. Using mitochondrial data, we infer that such populations diverged some 100,000\u00a0years ago, at the beginning of the last glacial cycle in Europe. Nuclear data show that contractions at multiple glacial refugia, and post\u2010glacial expansions have facilitated genetic differentiation between lineages that today interact in hybrid zones. We find extensive introgression throughout the sampled species range, irrespective of the current strength of reproductive isolation. Populations exhibiting hybrid male sterility in two hybrid zones show repeatable patterns of genomic differentiation, consistent with shared genomic constraints affecting ancestral divergence or with the role of those regions in reproductive isolation. Together, our results suggest that reproductive barriers that characterize late stages of species formation can evolve relatively quickly, particularly when associated with strong demographic changes. Moreover, we show that such barriers persist in the face of extensive gene flow, allowing future studies to identify associated genomic regions.<\/jats:p>","DOI":"10.1111\/mec.16406","type":"journal-article","created":{"date-parts":[[2022,2,22]],"date-time":"2022-02-22T07:59:13Z","timestamp":1645516753000},"page":"2384-2399","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":23,"title":["Extensive introgression at late stages of species formation: Insights from grasshopper hybrid zones"],"prefix":"10.1111","volume":"31","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-1441-6931","authenticated-orcid":false,"given":"Linda","family":"Hagberg","sequence":"first","affiliation":[{"name":"Division of Evolutionary Biology, Faculty of Biology II Ludwig\u2010Maximilians\u2010Universit\u00e4t M\u00fcnchen Planegg\u2010Martinsried Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Enrique","family":"Celem\u00edn","sequence":"additional","affiliation":[{"name":"Division of Evolutionary Biology, Faculty of Biology II Ludwig\u2010Maximilians\u2010Universit\u00e4t M\u00fcnchen Planegg\u2010Martinsried Germany"},{"name":"Unit of Evolutionary Biology\/Systematic Zoology, Institute of Biochemistry and Biology Universit\u00e4t Potsdam Potsdam Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Iker","family":"Irisarri","sequence":"additional","affiliation":[{"name":"Institute for Microbiology and Genetics, Department of Applied Bioinformatics University of Goettingen G\u00f6ttingen Germany"},{"name":"Campus Institute Data Science (CIDAS) G\u00f6ttingen Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8010-4157","authenticated-orcid":false,"given":"Oliver","family":"Hawlitschek","sequence":"additional","affiliation":[{"name":"Leibniz Institute for the Analysis of Biodiversity Change Zoological Museum Hamburg Germany"},{"name":"Zoologische Staatssammlung (SNSB\u2010ZSM) Munich Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jos\u00e9 L.","family":"Bella","sequence":"additional","affiliation":[{"name":"Departamento de Biolog\u00eda (Gen\u00e9tica), Facultad de Ciencias Universidad Aut\u00f3noma de Madrid Madrid Spain"},{"name":"Centro de Investigaci\u00f3n en Biodiversidad y Cambio Global (CIBC\u2010UAM) Universidad Aut\u00f3noma de Madrid Madrid Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tam\u00ed","family":"Mott","sequence":"additional","affiliation":[{"name":"Instituto de Ci\u00eancias Biol\u00f3gicas e da Sa\u00fade Universidade Federal de Alagoas Macei\u00f3 Brazil"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8076-4822","authenticated-orcid":false,"given":"Ricardo J.","family":"Pereira","sequence":"additional","affiliation":[{"name":"Division of Evolutionary Biology, Faculty of Biology II Ludwig\u2010Maximilians\u2010Universit\u00e4t M\u00fcnchen Planegg\u2010Martinsried Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2022,3,5]]},"reference":[{"key":"e_1_2_9_2_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.es.16.110185.000553"},{"key":"e_1_2_9_3_1","doi-asserted-by":"publisher","DOI":"10.1038\/341497a0"},{"key":"e_1_2_9_4_1","volume-title":"Heredity and variation in modern lights","author":"Bateson W.","year":"1909"},{"key":"e_1_2_9_5_1","doi-asserted-by":"publisher","DOI":"10.1038\/hdy.1992.49"},{"key":"e_1_2_9_6_1","doi-asserted-by":"publisher","DOI":"10.1017\/S001667230002886X"},{"key":"e_1_2_9_7_1","doi-asserted-by":"publisher","DOI":"10.1111\/mec.13276"},{"key":"e_1_2_9_8_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2012.08.023"},{"key":"e_1_2_9_9_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467\u2010017\u201001761\u20104"},{"key":"e_1_2_9_10_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.91.14.6491"},{"key":"e_1_2_9_11_1","doi-asserted-by":"publisher","DOI":"10.1016\/0169\u20105347(87)90193\u20105"},{"key":"e_1_2_9_12_1","first-page":"367","volume-title":"Endless forms: Species and speciation","author":"Butlin R. 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