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Among the few aquatic mammalian relicts, the Saimaa ringed seal (\n                    <jats:italic toggle=\"yes\">Pusa hispida saimensis<\/jats:italic>\n                    ) has been landlocked in Lake Saimaa, Finland, for the last 10,000 y. Saimaa ringed seals show genetic, behavioral, and morphological differences from the other ringed seal subspecies, but the extent these differences stem from the end of the last glacial period remains unclear. Here, we demonstrate with comprehensive sampling and state-of-the-art genomic methods that the Saimaa ringed seals are much older than the lake they inhabit, having formed a separate evolutionary branch for at least 60,000 y. This deep evolutionary origin of the Saimaa ringed seals is further underscored by anatomical differences, including our ecomorphological analyses revealing adaptively distinct features in their dentition and tongue. Overall, glacial relicts may harbor a richer selection of evolutionary history than might be expected from their recent isolation history alone.\n                  <\/jats:p>","DOI":"10.1073\/pnas.2503368122","type":"journal-article","created":{"date-parts":[[2025,6,10]],"date-time":"2025-06-10T11:08:22Z","timestamp":1749553702000},"update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":12,"title":["Deep origins, distinct adaptations, and species-level status indicated for a glacial relict seal"],"prefix":"10.1073","volume":"122","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-5389-6611","authenticated-orcid":false,"given":"Ari","family":"L\u00f6ytynoja","sequence":"first","affiliation":[{"name":"Organismal and Evolutionary Biology Research Programme, Faculty of Biological and Environmental Sciences, University of Helsinki","place":["Helsinki, Finland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1185-3610","authenticated-orcid":false,"given":"Jaakko","family":"Pohjoism\u00e4ki","sequence":"additional","affiliation":[{"name":"Department of Environmental and Biological Sciences, University of Eastern Finland","place":["Joensuu, Finland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mia","family":"Valtonen","sequence":"additional","affiliation":[{"name":"Natural Resources Institute Finland (Luke)","place":["Helsinki, Finland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4040-6588","authenticated-orcid":false,"given":"Juha","family":"Laakkonen","sequence":"additional","affiliation":[{"name":"Division of Veterinary Anatomy and Developmental Biology, Department of Veterinary Biosciences, Faculty of Veterinary Medicine, University of Helsinki","place":["Helsinki, Finland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7443-400X","authenticated-orcid":false,"given":"Wataru","family":"Morita","sequence":"additional","affiliation":[{"name":"Division of Anthropology, Department of Paleontology and Anthropology, National Museum of Nature and Science","place":["Tsukuba, Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mervi","family":"Kunnasranta","sequence":"additional","affiliation":[{"name":"Department of Environmental and Biological Sciences, University of Eastern Finland","place":["Joensuu, Finland"]},{"name":"Natural Resources Institute Finland (Luke)","place":["Joensuu, Finland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0056-5741","authenticated-orcid":false,"given":"Risto","family":"V\u00e4in\u00f6l\u00e4","sequence":"additional","affiliation":[{"name":"Finnish Museum of Natural History, University of Helsinki","place":["Helsinki, Finland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Morten Tange","family":"Olsen","sequence":"additional","affiliation":[{"name":"Section of Molecular Ecology and Evolution, Globe Institute, University of Copenhagen","place":["Copenhagen, Denmark"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3947-4778","authenticated-orcid":false,"given":"Petri","family":"Auvinen","sequence":"additional","affiliation":[{"name":"Institute of Biotechnology, Helsinki Institute of Life Science, University of Helsinki","place":["Helsinki, Finland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6575-8486","authenticated-orcid":false,"given":"Jukka","family":"Jernvall","sequence":"additional","affiliation":[{"name":"Institute of Biotechnology, Helsinki Institute of Life Science, University of Helsinki","place":["Helsinki, Finland"]},{"name":"Department of Geosciences and Geography, University of Helsinki","place":["Helsinki, Finland"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"341","published-online":{"date-parts":[[2025,6,10]]},"reference":[{"key":"e_1_3_4_1_2","first-page":"1","article-title":"The biology of the ringed seal (Phoca hispida Schreber) in the eastern Canadian Arctic","volume":"118","author":"McLaren I. 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