{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,31]],"date-time":"2026-07-31T20:23:32Z","timestamp":1785529412958,"version":"3.56.0"},"reference-count":163,"publisher":"Springer Science and Business Media LLC","issue":"12","license":[{"start":{"date-parts":[[2025,10,28]],"date-time":"2025-10-28T00:00:00Z","timestamp":1761609600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"},{"start":{"date-parts":[[2025,10,28]],"date-time":"2025-10-28T00:00:00Z","timestamp":1761609600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"}],"funder":[{"DOI":"10.13039\/501100000038","name":"Gouvernement du Canada | Natural Sciences and Engineering Research Council of Canada","doi-asserted-by":"publisher","award":["RGPIN-2018-05305"],"award-info":[{"award-number":["RGPIN-2018-05305"]}],"id":[{"id":"10.13039\/501100000038","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000243","name":"W. Garfield Weston Foundation","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100000243","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Nat Ecol Evol"],"DOI":"10.1038\/s41559-025-02872-8","type":"journal-article","created":{"date-parts":[[2025,10,28]],"date-time":"2025-10-28T16:04:00Z","timestamp":1761667440000},"page":"2223-2237","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Mid-Cenozoic rhinocerotid dispersal via the North Atlantic"],"prefix":"10.1038","volume":"9","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-0228-2617","authenticated-orcid":false,"given":"Danielle","family":"Fraser","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0089-3649","authenticated-orcid":false,"given":"Natalia","family":"Rybczynski","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marisa","family":"Gilbert","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mary R.","family":"Dawson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,10,28]]},"reference":[{"key":"2872_CR1","doi-asserted-by":"publisher","first-page":"145","DOI":"10.1007\/s00114-010-0747-y","volume":"98","author":"R Tabuce","year":"2011","unstructured":"Tabuce, R., Clavel, J. & Antunes, M. T. A structural intermediate between triisodontids and mesonychians (Mammalia, Acreodi) from the earliest Eocene of Portugal. Naturwissenschaften 98, 145\u2013155 (2011).","journal-title":"Naturwissenschaften"},{"key":"2872_CR2","doi-asserted-by":"crossref","unstructured":"Rose, K. D. The Beginning of the Age of Mammals (JHU Press, 2006).","DOI":"10.56021\/9780801884726"},{"key":"2872_CR3","first-page":"29","volume":"49","author":"P Missiaen","year":"2011","unstructured":"Missiaen, P. An updated mammalian biochronology and biogeography for the early Paleogene of Asia. Vert. PalAs. 49, 29\u201352 (2011).","journal-title":"Vert. PalAs."},{"key":"2872_CR4","doi-asserted-by":"crossref","unstructured":"Lofgren, D. L., Lillegraven, J. A., Clemens, W. A., Gingerich, P. D. & Williamson, T. A. in Late Cretaceous and Cenozoic Mammals of North America: Biostratigraphy and Biochronology (ed. Woodburne, M.O.) 43\u2013105 (Columbia Univ. Press, 2004).","DOI":"10.7312\/wood13040-005"},{"key":"2872_CR5","unstructured":"McKenna, M. C. & Bell, S. K. Classification of Mammals Above the Species Level (Columbia Univ. Press, 1997)."},{"key":"2872_CR6","unstructured":"Lucas, S. G. in Evolution of Tertiary Mammals of North America (eds Janis, C. M. et al.) 274\u2013283 (Cambridge Univ. Press, 1998)."},{"key":"2872_CR7","doi-asserted-by":"publisher","first-page":"521","DOI":"10.1080\/02724631003617449","volume":"30","author":"CJ L\u00fcthje","year":"2010","unstructured":"L\u00fcthje, C. J., Mil\u00e0n, J. & Hurum, J. H. Paleocene tracks of the mammal pantodont genus Titanoides in coal-bearing strata, Svalbard, Arctic Norway. J. Vertebr. Paleontol. 30, 521\u2013527 (2010).","journal-title":"J. Vertebr. Paleontol."},{"key":"2872_CR8","doi-asserted-by":"publisher","first-page":"613","DOI":"10.1130\/0016-7606(1947)58[613:HMFACR]2.0.CO;2","volume":"58","author":"GG Simpson","year":"1947","unstructured":"Simpson, G. G. Holarctic mammal faunas and continental relationships during the Cenozoic. Geol. Soc. Am. Bull. 58, 613\u2013688 (1947).","journal-title":"Geol. Soc. Am. Bull."},{"key":"2872_CR9","doi-asserted-by":"publisher","first-page":"472","DOI":"10.2307\/2666183","volume":"86","author":"SR Manchester","year":"1999","unstructured":"Manchester, S. R. Biogeographical relationships of North American tertiary floras. Ann. Mo. Bot. Gard. 86, 472\u2013522 (1999).","journal-title":"Ann. Mo. Bot. Gard."},{"key":"2872_CR10","doi-asserted-by":"publisher","first-page":"240","DOI":"10.1086\/297343","volume":"157","author":"SR Manchester","year":"1996","unstructured":"Manchester, S. R. & Shuang-Xing, G. Palaeocarpinus (extinct Betulaceae) from northwestern China: new evidence for Paleocene floristic continuity between Asia, North America, and Europe. Int. J. Plant Sci. 157, 240\u2013246 (1996).","journal-title":"Int. J. Plant Sci."},{"key":"2872_CR11","doi-asserted-by":"crossref","unstructured":"Marincovich, L. et al. in The Arctic Ocean Region (eds Grantz, A. et al.) 403\u2013426 (Geological Society of America, 1990).","DOI":"10.1130\/DNAG-GNA-L.403"},{"key":"2872_CR12","first-page":"175","volume":"12","author":"AY Gladenkov","year":"2004","unstructured":"Gladenkov, A. Y. Onset of connections between the Pacific and Arctic oceans through the Bering Strait in the Neogene. Stratigr. Geol. Correl. 12, 175\u2013187 (2004).","journal-title":"Stratigr. Geol. Correl."},{"key":"2872_CR13","doi-asserted-by":"publisher","first-page":"1550","DOI":"10.1038\/ncomms2516","volume":"4","author":"N Rybczynski","year":"2013","unstructured":"Rybczynski, N. et al. Mid-Pliocene warm-period deposits in the High Arctic yield insight into camel evolution. Nat. Commun. 4, 1550 (2013).","journal-title":"Nat. Commun."},{"key":"2872_CR14","doi-asserted-by":"publisher","first-page":"439","DOI":"10.1139\/e01-078","volume":"39","author":"JH Hutchison","year":"2002","unstructured":"Hutchison, J. H. & Harington, C. A peculiar new fossil shrew (Lipotyphla, Soricidae) from the High Arctic of Canada. Can. J. Earth Sci. 39, 439\u2013443 (2002).","journal-title":"Can. J. Earth Sci."},{"key":"2872_CR15","doi-asserted-by":"publisher","first-page":"17722","DOI":"10.1038\/s41598-017-17657-8","volume":"7","author":"X Wang","year":"2017","unstructured":"Wang, X., Rybczynski, N., Harington, C. R., White, S. C. & Tedford, R. H. A basal ursine bear (Protarctos abstrusus) from the Pliocene High Arctic reveals Eurasian affinities and a diet rich in fermentable sugars. Sci. Rep. 7, 17722 (2017).","journal-title":"Sci. Rep."},{"key":"2872_CR16","doi-asserted-by":"publisher","first-page":"1017","DOI":"10.1111\/1475-4983.00108","volume":"42","author":"J Hulbert","year":"1999","unstructured":"Hulbert, J., Richard, C. & Harington, C. R. An early Pliocene hipparionine horse from the Canadian Arctic. Palaeontology 42, 1017\u20131025 (1999).","journal-title":"Palaeontology"},{"key":"2872_CR17","doi-asserted-by":"publisher","first-page":"1728","DOI":"10.1111\/jbi.12765","volume":"43","author":"S Koblm\u00fcller","year":"2016","unstructured":"Koblm\u00fcller, S. et al. Whole mitochondrial genomes illuminate ancient intercontinental dispersals of grey wolves (Canis lupus). J. Biogeogr. 43, 1728\u20131738 (2016).","journal-title":"J. Biogeogr."},{"key":"2872_CR18","doi-asserted-by":"publisher","first-page":"6407","DOI":"10.1111\/mec.16267","volume":"31","author":"AT Salis","year":"2022","unstructured":"Salis, A. T. et al. Lions and brown bears colonized North America in multiple synchronous waves of dispersal across the Bering Land Bridge. Mol. Ecol. 31, 6407\u20136421 (2022).","journal-title":"Mol. Ecol."},{"key":"2872_CR19","doi-asserted-by":"publisher","first-page":"748","DOI":"10.1126\/science.adr2355","volume":"388","author":"Y Running Horse Collin","year":"2025","unstructured":"Running Horse Collin, Y. et al. Sustainability insights from Late Pleistocene climate change and horse migration patterns. Science 388, 748\u2013755 (2025).","journal-title":"Science"},{"key":"2872_CR20","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-018-07828-0","volume":"9","author":"S Kender","year":"2018","unstructured":"Kender, S. et al. Closure of the Bering Strait caused mid-Pleistocene transition cooling. Nat. Commun. 9, 5386 (2018).","journal-title":"Nat. Commun."},{"key":"2872_CR21","doi-asserted-by":"publisher","first-page":"1287","DOI":"10.1002\/2015PA002840","volume":"30","author":"KP Knudson","year":"2015","unstructured":"Knudson, K. P. & Ravelo, A. C. North Pacific Intermediate Water circulation enhanced by the closure of the Bering Strait. Paleoceanography 30, 1287\u20131304 (2015).","journal-title":"Paleoceanography"},{"key":"2872_CR22","doi-asserted-by":"publisher","first-page":"1036","DOI":"10.1111\/jbi.12310","volume":"41","author":"L Brikiatis","year":"2014","unstructured":"Brikiatis, L. The De Geer, Thulean and Beringia routes: key concepts for understanding early Cenozoic biogeography. J. Biogeogr. 41, 1036\u20131054 (2014).","journal-title":"J. Biogeogr."},{"key":"2872_CR23","doi-asserted-by":"publisher","first-page":"459","DOI":"10.2307\/2992083","volume":"70","author":"MC McKenna","year":"1983","unstructured":"McKenna, M. C. Holarctic landmass rearrangement, cosmic events, and Cenozoic terrestrial organisms. Ann. Mo. Bot. Gard. 70, 459\u2013489 (1983).","journal-title":"Ann. Mo. Bot. Gard."},{"key":"2872_CR24","doi-asserted-by":"crossref","unstructured":"McKenna, M. C. in Structure and Development of the Greenland-Scotland Ridge: New Methods and Concepts Vol. 8 (eds Bott, M. H. P. et al.) 351\u2013399 (Springer, 1983).","DOI":"10.1007\/978-1-4613-3485-9_19"},{"key":"2872_CR25","first-page":"243","volume":"66","author":"BH Tiffney","year":"1985","unstructured":"Tiffney, B. H. The Eocene North Atlantic land bridge: its importance in Tertiary and modern phytogeography of the northern hemisphere. J. Arnold Arbor. 66, 243\u2013273 (1985).","journal-title":"J. Arnold Arbor."},{"key":"2872_CR26","first-page":"269","volume":"144","author":"FS Szalay","year":"1971","unstructured":"Szalay, F. S. & McKenna, M. C. Beginning of the age of mammals in Asia: the late Paleocene Gashato fauna, Mongolia. Bull. Am. Mus. Nat. Hist. 144, 269\u2013318 (1971).","journal-title":"Bull. Am. Mus. Nat. Hist."},{"key":"2872_CR27","doi-asserted-by":"crossref","unstructured":"Tucholke, B. E. & McCoy, F. W. in The Western North Atlantic Region (eds Vogt, P. R. & Tucholke, B. E.) 589\u2013602 (Geological Society of America, 1986).","DOI":"10.1130\/DNAG-GNA-M.589"},{"key":"2872_CR28","doi-asserted-by":"publisher","first-page":"621","DOI":"10.1029\/98TC01890","volume":"17","author":"AE Blythe","year":"1998","unstructured":"Blythe, A. E. & Kleinspehn, K. L. Tectonically versus climatically driven Cenozoic exhumation of the Eurasian plate margin, Svalbard: fission track analyses. Tectonics 17, 621\u2013639 (1998).","journal-title":"Tectonics"},{"key":"2872_CR29","doi-asserted-by":"publisher","first-page":"110","DOI":"10.14430\/arctic2822","volume":"28","author":"MR Dawson","year":"1975","unstructured":"Dawson, M. R., West, R. M., Ramaekers, P. & Hutchison, J. H. New evidence on the palaeobiology of the Eureka Sound Formation, Arctic Canada. Arctic 28, 110\u2013116 (1975).","journal-title":"Arctic"},{"key":"2872_CR30","doi-asserted-by":"publisher","first-page":"153","DOI":"10.1016\/0031-0182(90)90127-S","volume":"80","author":"RD M\u00fcller","year":"1990","unstructured":"M\u00fcller, R. D. & Spielhagen, R. F. Evolution of the Central Tertiary Basin of Spitsbergen: towards a synthesis of sediment and plate tectonic history. Palaeogeogr. Palaeoclimatol. Palaeoecol. 80, 153\u2013172 (1990).","journal-title":"Palaeogeogr. Palaeoclimatol. Palaeoecol."},{"key":"2872_CR31","first-page":"115","volume":"592","author":"C Lee","year":"2008","unstructured":"Lee, C., Lehnert, O. & Nowlan, G. Sedimentology, stratigraphy, and clast biostratigraphy of Cretaceous and Tertiary strata, northeastern Ellesmere Island, Nunavut. Geol. Surv. Can. Bull. 592, 115\u2013167 (2008).","journal-title":"Geol. Surv. Can. Bull."},{"key":"2872_CR32","doi-asserted-by":"publisher","first-page":"335","DOI":"10.2307\/2395200","volume":"62","author":"MC McKenna","year":"1975","unstructured":"McKenna, M. C. Fossil mammals and early Eocene North Atlantic land continuity. Ann. Mo. Bot. Gard. 62, 335\u2013353 (1975).","journal-title":"Ann. Mo. Bot. Gard."},{"key":"2872_CR33","doi-asserted-by":"publisher","first-page":"381","DOI":"10.1007\/BF01820951","volume":"60","author":"F Strauch","year":"1970","unstructured":"Strauch, F. Die Thule-Landbr\u00fccke als Wanderweg und Faunenscheide zwischen Atlantik und Skandik im Terti\u00e4r. Geol. Rundsch. 60, 381\u2013417 (1970).","journal-title":"Geol. Rundsch."},{"key":"2872_CR34","doi-asserted-by":"publisher","first-page":"629","DOI":"10.2307\/2398982","volume":"70","author":"DI Axelrod","year":"1983","unstructured":"Axelrod, D. I. Biogeography of oaks in the Arcto-Tertiary province. Ann. Mo. Bot. Gard. 70, 629\u2013657 (1983).","journal-title":"Ann. Mo. Bot. Gard."},{"key":"2872_CR35","unstructured":"Saunders, A. D., Fitton, J. G., Kerr, A. C., Norry, M. J. & Kent, R. W. in Large Igneous Provinces: Continental, Oceanic, and Planetary Flood Volcanism (eds Coffin, M. F. & Mahoney, J. J.) 45\u201393 (American Geophysical Union, 1997)."},{"key":"2872_CR36","doi-asserted-by":"publisher","first-page":"255","DOI":"10.1130\/G32461.1","volume":"40","author":"MW Hohbein","year":"2012","unstructured":"Hohbein, M. W., Sexton, P. F. & Cartwright, J. A. Onset of North Atlantic Deep Water production coincident with inception of the Cenozoic global cooling trend. Geology 40, 255\u2013258 (2012).","journal-title":"Geology"},{"key":"2872_CR37","doi-asserted-by":"publisher","first-page":"627","DOI":"10.1144\/0016-764902-064","volume":"159","author":"JR Smallwood","year":"2002","unstructured":"Smallwood, J. R. & Gill, C. E. The rise and fall of the Faroe\u2013Shetland Basin: evidence from seismic mapping of the Balder Formation. J. Geol. Soc. 159, 627\u2013630 (2002).","journal-title":"J. Geol. Soc."},{"key":"2872_CR38","first-page":"TC1002","volume":"27","author":"M Shaw Champion","year":"2008","unstructured":"Shaw Champion, M., White, N., Jones, S. & Lovell, J. Quantifying transient mantle convective uplift: an example from the Faroe-Shetland basin. Tectonics 27, TC1002 (2008).","journal-title":"Tectonics"},{"key":"2872_CR39","doi-asserted-by":"publisher","first-page":"562","DOI":"10.1038\/ngeo1191","volume":"4","author":"RA Hartley","year":"2011","unstructured":"Hartley, R. A., Roberts, G. G., White, N. & Richardson, C. Transient convective uplift of an ancient buried landscape. Nat. Geosci. 4, 562\u2013565 (2011).","journal-title":"Nat. Geosci."},{"key":"2872_CR40","first-page":"697","volume":"170","author":"KC Beard","year":"1999","unstructured":"Beard, K. C. & Dawson, M. R. Intercontinental dispersal of holarctic land mammals near the Paleocene\/Eocene boundary; paleogeographic, paleoclimatic and biostratigraphic implications. Bull. Soc. Geol. Fr. 170, 697\u2013706 (1999).","journal-title":"Bull. Soc. Geol. Fr."},{"key":"2872_CR41","doi-asserted-by":"crossref","unstructured":"Gingerich, P. D. in Causes and Consequences of Globally Warm Climates in the Early Paleogene (eds Wing, S. L. et al.) 463\u2013478 (Geological Society of America, 2003).","DOI":"10.1130\/SPE369"},{"key":"2872_CR42","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1206\/0003-0082(2005)474[0001:AACOFL]2.0.CO;2","volume":"3474","author":"GJ Bowen","year":"2005","unstructured":"Bowen, G. J., Koch, P. L., Meng, J., Ye, J. & Ting, S. Age and correlation of fossiliferous late Paleocene\u2013early Eocene strata of the Erlian Basin, Inner Mongolia, China. Am. Mus. Novit. 3474, 1\u201326 (2005).","journal-title":"Am. Mus. Novit."},{"key":"2872_CR43","doi-asserted-by":"publisher","first-page":"11223","DOI":"10.1073\/pnas.0511296103","volume":"103","author":"T Smith","year":"2006","unstructured":"Smith, T., Rose, K. D. & Gingerich, P. D. Rapid Asia\u2013Europe\u2013North America geographic dispersal of earliest Eocene primate Teilhardina during the Paleocene\u2013Eocene thermal maximum. Proc. Natl Acad. Sci. USA 103, 11223\u201311227 (2006).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2872_CR44","doi-asserted-by":"publisher","first-page":"1107","DOI":"10.1139\/e01-010","volume":"38","author":"MR Dawson","year":"2001","unstructured":"Dawson, M. R. Early Eocene rodents (Mammalia) from the Eureka Sound Group of Ellesmere Island, Canada. Can. J. Earth Sci. 38, 1107\u20131116 (2001).","journal-title":"Can. J. Earth Sci."},{"key":"2872_CR45","first-page":"179","volume":"3","author":"MR Dawson","year":"1993","unstructured":"Dawson, M. R., McKenna, M. C., Beard, K. C. & Hutchison, J. H. An early Eocene plagiomenid mammal from Ellesmere and Axel Heiberg Islands, Arctic Canada. Kaupia 3, 179\u2013192 (1993).","journal-title":"Kaupia"},{"key":"2872_CR46","doi-asserted-by":"publisher","first-page":"126","DOI":"10.1016\/j.gr.2020.05.011","volume":"86","author":"EO Straume","year":"2020","unstructured":"Straume, E. O., Gaina, C., Medvedev, S. & Nisancioglu, K. H. Global Cenozoic paleobathymetry with a focus on the northern hemisphere oceanic gateways. Gondwana Res. 86, 126\u2013143 (2020).","journal-title":"Gondwana Res."},{"key":"2872_CR47","doi-asserted-by":"publisher","first-page":"231","DOI":"10.1038\/s43247-023-00899-y","volume":"4","author":"AP Lasabuda","year":"2023","unstructured":"Lasabuda, A. P. et al. Paleobathymetric reconstructions of the SW Barents Seaway and their implications for Atlantic\u2013Arctic ocean circulation. Commun. Earth Environ. 4, 231 (2023).","journal-title":"Commun. Earth Environ."},{"key":"2872_CR48","doi-asserted-by":"publisher","first-page":"Q02015","DOI":"10.1029\/2007GC001824","volume":"9","author":"J Knies","year":"2008","unstructured":"Knies, J. & Gaina, C. Middle Miocene ice sheet expansion in the Arctic: views from the Barents Sea. Geochem. Geophys. Geosyst. 9, Q02015 (2008).","journal-title":"Geochem. Geophys. Geosyst."},{"key":"2872_CR49","doi-asserted-by":"publisher","first-page":"Q06010","DOI":"10.1029\/2005GC001085","volume":"7","author":"H Poore","year":"2006","unstructured":"Poore, H., Samworth, R., White, N., Jones, S. & McCave, I. Neogene overflow of northern component water at the Greenland-Scotland Ridge. Geochem. Geophys. Geosyst. 7, Q06010 (2006).","journal-title":"Geochem. Geophys. Geosyst."},{"key":"2872_CR50","doi-asserted-by":"publisher","first-page":"393","DOI":"10.1038\/255393a0","volume":"255","author":"P Robertson","year":"1975","unstructured":"Robertson, P. & Mason, G. D. Shatter cones from Haughton Dome, Devon Island, Canada. Nature 255, 393\u2013394 (1975).","journal-title":"Nature"},{"key":"2872_CR51","doi-asserted-by":"crossref","unstructured":"Dinerstein, E. The Return of the Unicorns: The Natural History and Conservation of the Greater One-Horned Rhinoceros (Columbia Univ. Press, 2003).","DOI":"10.7312\/dine08450"},{"key":"2872_CR52","doi-asserted-by":"publisher","first-page":"355","DOI":"10.1017\/S0030605315000472","volume":"50","author":"RG Havm\u00f8ller","year":"2016","unstructured":"Havm\u00f8ller, R. G. et al. Will current conservation responses save the critically endangered Sumatran rhinoceros Dicerorhinus sumatrensis?. Oryx 50, 355\u2013359 (2016).","journal-title":"Oryx"},{"key":"2872_CR53","unstructured":"Khan, M. K. B. M. & Strien, N. J. V. Asian Rhinos: Status Survey and Conservation Action Plan Vol. 32 (IUCN, 1997)."},{"key":"2872_CR54","doi-asserted-by":"publisher","first-page":"91","DOI":"10.69649\/pachyderm.v52i.311","volume":"52","author":"K Rookmaaker","year":"2012","unstructured":"Rookmaaker, K. & Antoine, P.-O. New maps representing the historical and recent distribution of the African species of rhinoceros: Diceros bicornis, Ceratotherium simum and Ceratotherium cottoni. Pachyderm 52, 91\u201396 (2012).","journal-title":"Pachyderm"},{"key":"2872_CR55","doi-asserted-by":"crossref","unstructured":"Antoine, P.-O., Becker, D., Pandolfi, L. & Geraads, D. in Rhinos of the World: Ecology, Conservation and Management (eds Melletti, M. et al.) 31\u201348 (Springer, 2025).","DOI":"10.1007\/978-3-031-67169-2_2"},{"key":"2872_CR56","doi-asserted-by":"publisher","first-page":"396","DOI":"10.1016\/j.jtbi.2010.09.010","volume":"267","author":"T Stadler","year":"2010","unstructured":"Stadler, T. Sampling-through-time in birth\u2013death trees. J. Theor. Biol. 267, 396\u2013404 (2010).","journal-title":"J. Theor. Biol."},{"key":"2872_CR57","doi-asserted-by":"publisher","first-page":"251","DOI":"10.1146\/annurev-ecolsys-102220-030855","volume":"53","author":"AM Wright","year":"2022","unstructured":"Wright, A. M., Bapst, D. W., Barido-Sottani, J. & Warnock, R. C. Integrating fossil observations into phylogenetics using the fossilized birth\u2013death model. Annu. Rev. Ecol. Evol. Syst. 53, 251\u2013273 (2022).","journal-title":"Annu. Rev. Ecol. Evol. Syst."},{"key":"2872_CR58","doi-asserted-by":"publisher","first-page":"3836","DOI":"10.1002\/grl.50745","volume":"40","author":"KE Young","year":"2013","unstructured":"Young, K. E. et al. Impact thermochronology and the age of Haughton impact structure, Canada. Geophys. Res. Lett. 40, 3836\u20133840 (2013).","journal-title":"Geophys. Res. Lett."},{"key":"2872_CR59","doi-asserted-by":"publisher","first-page":"233","DOI":"10.1111\/j.1945-5100.1988.tb01285.x","volume":"23","author":"EK Jessberger","year":"1988","unstructured":"Jessberger, E. K. Ar-40-Ar-39 dating of the Haughton impact structure. Meteoritics 23, 233\u2013234 (1988).","journal-title":"Meteoritics"},{"key":"2872_CR60","doi-asserted-by":"publisher","first-page":"1603","DOI":"10.1126\/science.237.4822.1603","volume":"237","author":"G Omar","year":"1987","unstructured":"Omar, G. et al. Fission-track dating of Haughton astrobleme and included biota, Devon Island, Canada. Science 237, 1603\u20131605 (1987).","journal-title":"Science"},{"key":"2872_CR61","doi-asserted-by":"publisher","first-page":"1005126","DOI":"10.3389\/fevo.2023.1005126","volume":"11","author":"X-K Lu","year":"2023","unstructured":"Lu, X.-K., Deng, T. & Pandolfi, L. Reconstructing the phylogeny of the hornless rhinoceros Aceratheriinae. Front. Ecol. Evol. 11, 1005126 (2023).","journal-title":"Front. Ecol. Evol."},{"key":"2872_CR62","doi-asserted-by":"publisher","first-page":"803","DOI":"10.1111\/j.2041-210X.2012.00223.x","volume":"3","author":"DW Bapst","year":"2012","unstructured":"Bapst, D. W. paleotree: an R package for paleontological and phylogenetic analyses of evolution. Methods Ecol. Evol. 3, 803\u2013807 (2012).","journal-title":"Methods Ecol. Evol."},{"key":"2872_CR63","doi-asserted-by":"publisher","first-page":"242","DOI":"10.21425\/F55419694","volume":"5","author":"NJ Matzke","year":"2013","unstructured":"Matzke, N. J. Probabilistic historical biogeography: new models for founder-event speciation, imperfect detection, and fossils allow improved accuracy and model-testing. Front. Biogeogr. 5, 242\u2013248 (2013).","journal-title":"Front. Biogeogr."},{"key":"2872_CR64","doi-asserted-by":"publisher","first-page":"951","DOI":"10.1093\/sysbio\/syu056","volume":"63","author":"NJ Matzke","year":"2014","unstructured":"Matzke, N. J. Model selection in historical biogeography reveals that founder-event speciation is a crucial process in island clades. Syst. Biol. 63, 951\u2013970 (2014).","journal-title":"Syst. Biol."},{"key":"2872_CR65","unstructured":"Matzke, N. J. BioGeoBEARS: stochastic mapping under biogeographical models. R version v. 1.1.3 (2016)."},{"key":"2872_CR66","doi-asserted-by":"publisher","first-page":"4","DOI":"10.1080\/10635150701883881","volume":"57","author":"RH Ree","year":"2008","unstructured":"Ree, R. H. & Smith, S. A. Maximum likelihood inference of geographic range evolution by dispersal, local extinction, and cladogenesis. Syst. Biol. 57, 4\u201314 (2008).","journal-title":"Syst. Biol."},{"key":"2872_CR67","doi-asserted-by":"crossref","unstructured":"Owen, R. On the Archetype and Homologies of the Vertebrate Skeleton (John van Voorst, 1848).","DOI":"10.5962\/bhl.title.118611"},{"key":"2872_CR68","unstructured":"Owen, R. Odontography (Hippolyte Bailliere, 1845)."},{"key":"2872_CR69","first-page":"296","volume":"15","author":"JE Gray","year":"1821","unstructured":"Gray, J. E. On the natural arrangement of vertebrate mammals. Lond. Clin. Med. Repository 15, 296\u2013310 (1821).","journal-title":"Lond. Clin. Med. Repository"},{"key":"2872_CR70","unstructured":"Abel, O. Kritische Untersuchungen \u00fcber die pal\u00e4ogenen Rhinocerotiden Europas (H\u00f6lder, 1910)."},{"key":"2872_CR71","first-page":"1","volume":"170","author":"U Uhlig","year":"1999","unstructured":"Uhlig, U. Die Rhinocerotoidea (Mammalia) aus der unteroligoz\u00e4nen Spaltenf\u00fcllung M\u00f6hren 13 bei Treuchtlingen in Bayern. Abh. Bayer. Akad. Wiss. Math.-Naturwiss. Kl. 170, 1\u2013254 (1999).","journal-title":"Abh. Bayer. Akad. Wiss. Math.-Naturwiss. Kl."},{"key":"2872_CR72","first-page":"121","volume":"53","author":"M B\u00f6hme","year":"2013","unstructured":"B\u00f6hme, M. et al. Na Duong (northern Vietnam)-an exceptional window into Eocene ecosystems from Southeast Asia. Zitteliana A 53, 121\u2013167 (2013).","journal-title":"Zitteliana A"},{"key":"2872_CR73","doi-asserted-by":"publisher","first-page":"947","DOI":"10.1080\/14772019.2012.699007","volume":"11","author":"D Becker","year":"2013","unstructured":"Becker, D., Antoine, P.-O. & Maridet, O. A new genus of Rhinocerotidae (Mammalia, Perissodactyla) from the Oligocene of Europe. J. Syst. Paleontol. 11, 947\u2013972 (2013).","journal-title":"J. Syst. Paleontol."},{"key":"2872_CR74","doi-asserted-by":"publisher","first-page":"200633","DOI":"10.1098\/rsos.200633","volume":"7","author":"J Tissier","year":"2020","unstructured":"Tissier, J., Antoine, P.-O. & Becker, D. New material of Epiaceratherium and a new species of Mesaceratherium clear up the phylogeny of early Rhinocerotidae (Perissodactyla). R. Soc. Open Sci. 7, 200633 (2020).","journal-title":"R. Soc. Open Sci."},{"key":"2872_CR75","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1111\/j.1096-3642.2009.00597.x","volume":"160","author":"P-O Antoine","year":"2010","unstructured":"Antoine, P.-O. et al. A revision of Aceratherium blanfordi Lydekker, 1884 (Mammalia: Rhinocerotidae) from the Early Miocene of Pakistan: postcranials as a key. Zool. J. Linn. Soc. 160, 139\u2013194 (2010).","journal-title":"Zool. J. Linn. Soc."},{"key":"2872_CR76","doi-asserted-by":"publisher","first-page":"416","DOI":"10.1093\/zoolinnean\/zlab009","volume":"194","author":"P-O Antoine","year":"2022","unstructured":"Antoine, P.-O. et al. A new rhinoceros clade from the Pleistocene of Asia sheds light on mammal dispersals to the Philippines. Zool. J. Linn. Soc. 194, 416\u2013430 (2022).","journal-title":"Zool. J. Linn. Soc."},{"key":"2872_CR77","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1206\/3943.1","volume":"2019","author":"J Eberle","year":"2019","unstructured":"Eberle, J. et al. The first Tertiary fossils of mammals, turtles, and fish from Canada\u2019s Yukon. Am. Mus. Novit. 2019, 1\u201328 (2019).","journal-title":"Am. Mus. Novit."},{"key":"2872_CR78","doi-asserted-by":"publisher","first-page":"324","DOI":"10.14430\/arctic1628","volume":"43","author":"C Whitlock","year":"1990","unstructured":"Whitlock, C. & Dawson, M. R. Pollen and vertebrates of the Early Neogene Haughton Formation, Devon Island, Arctic Canada. Arctic 43, 324\u2013330 (1990).","journal-title":"Arctic"},{"key":"2872_CR79","doi-asserted-by":"publisher","first-page":"221","DOI":"10.1111\/j.1945-5100.1988.tb01284.x","volume":"23","author":"LJ Hickey","year":"1988","unstructured":"Hickey, L. J., Johnson, K. R. & Dawson, M. R. The stratigraphy, sedimentology, and fossils of the Haughton Formation: a post-impact crater-fill, Devon Island, NWT, Canada. Meteoritics 23, 221\u2013231 (1988).","journal-title":"Meteoritics"},{"key":"2872_CR80","doi-asserted-by":"publisher","first-page":"719","DOI":"10.1038\/s41586-025-09231-4","volume":"643","author":"RS Paterson","year":"2025","unstructured":"Paterson, R. S. et al. Phylogenetically informative proteins from an Early Miocene rhinocerotid. Nature 643, 719\u2013724 (2025).","journal-title":"Nature"},{"key":"2872_CR81","doi-asserted-by":"crossref","unstructured":"Antoine, P.-O., Becker, D., Pandolfi, L. & Geraads, D. in Rhinos of the World: Ecology, Conservation and Management Fascinating Life Sciences (eds. Melletti, M. et al.) 31\u201348 (Springer, 2025).","DOI":"10.1007\/978-3-031-67169-2_2"},{"key":"2872_CR82","unstructured":"Prothero, D. R. The Evolution of North American Rhinoceroses (Cambridge Univ. Press, 2005)."},{"key":"2872_CR83","doi-asserted-by":"publisher","first-page":"zlad183","DOI":"10.1093\/zoolinnean\/zlad183","volume":"202","author":"D Sun","year":"2024","unstructured":"Sun, D., Deng, T. & Wang, S. The first record of the genus Prosantorhinus (Perissodactyla: Rhinocerotidae) of East Asia. Zool. J. Linn. Soc. 202, zlad183 (2024).","journal-title":"Zool. J. Linn. Soc."},{"key":"2872_CR84","first-page":"154","volume":"60","author":"DE Savage","year":"1971","unstructured":"Savage, D. E. The Sparnacian-Wasatchian mammalian fauna, Early Eocene, of Europe and North America. Abh. Hess. Landesamt. Bodenf. 60, 154\u2013158 (1971).","journal-title":"Abh. Hess. Landesamt. Bodenf."},{"key":"2872_CR85","unstructured":"Dawson, M. R. in The Miocene Land Mammals of Europe (eds R\u00f6ssner, G. & Heissig, K.) 473\u2013483 (Verlag Dr Friedrich Pfeil, 1999)."},{"key":"2872_CR86","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1016\/j.revpalbo.2004.11.002","volume":"134","author":"F Gr\u00edmsson","year":"2005","unstructured":"Gr\u00edmsson, F. & Denk, T. Fagus from the Miocene of Iceland: systematics and biogeographical considerations. Rev. Palaeobot. Palynol. 134, 27\u201354 (2005).","journal-title":"Rev. Palaeobot. Palynol."},{"key":"2872_CR87","doi-asserted-by":"crossref","unstructured":"Stepien, C. A. & Haponski, A. E. in Biology and Culture of Percid Fishes: Principles and Practices (eds Kestemont, P. et al.) 3\u201360 (Springer, 2015).","DOI":"10.1007\/978-94-017-7227-3_1"},{"key":"2872_CR88","doi-asserted-by":"publisher","first-page":"156","DOI":"10.1111\/bij.12114","volume":"110","author":"AE Haponski","year":"2013","unstructured":"Haponski, A. E. & Stepien, C. A. Phylogenetic and biogeographical relationships of the Sander pikeperches (Percidae: Perciformes): patterns across North America and Eurasia. Biol. J. Linn. Soc. 110, 156\u2013179 (2013).","journal-title":"Biol. J. Linn. Soc."},{"key":"2872_CR89","unstructured":"Stepien, C., Behrmann-Godel, J, Bernatchez, L. in Biology of Perch (eds Moyer, G & Couture, P) Ch. 2 (CRC Press, 2015)."},{"key":"2872_CR90","doi-asserted-by":"publisher","first-page":"98","DOI":"10.1016\/j.tecto.2015.12.002","volume":"691","author":"W Jokat","year":"2016","unstructured":"Jokat, W., Lehmann, P., Damaske, D. & Nelson, J. B. Magnetic signature of North-East Greenland, the Morris Jesup Rise, the Yermak Plateau, the central Fram Strait: constraints for the rift\/drift history between Greenland and Svalbard since the Eocene. Tectonophysics 691, 98\u2013109 (2016).","journal-title":"Tectonophysics"},{"key":"2872_CR91","doi-asserted-by":"publisher","first-page":"25","DOI":"10.1007\/s11001-013-9165-9","volume":"34","author":"B-M Ehlers","year":"2013","unstructured":"Ehlers, B.-M. & Jokat, W. Paleo-bathymetry of the northern North Atlantic and consequences for the opening of the Fram Strait. Mar. Geophys. Res. 34, 25\u201343 (2013).","journal-title":"Mar. Geophys. Res."},{"key":"2872_CR92","doi-asserted-by":"publisher","first-page":"986","DOI":"10.1038\/nature05924","volume":"447","author":"M Jakobsson","year":"2007","unstructured":"Jakobsson, M. et al. The early Miocene onset of a ventilated circulation regime in the Arctic Ocean. Nature 447, 986\u2013990 (2007).","journal-title":"Nature"},{"key":"2872_CR93","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1016\/j.tecto.2008.01.002","volume":"450","author":"\u00d8 Engen","year":"2008","unstructured":"Engen, \u00d8, Faleide, J. I. & Dyreng, T. K. Opening of the Fram Strait gateway: a review of plate tectonic constraints. Tectonophysics 450, 51\u201369 (2008).","journal-title":"Tectonophysics"},{"key":"2872_CR94","doi-asserted-by":"crossref","unstructured":"Kristoffersen, Y. in Geological History of the Polar Oceans: Arctic Versus Antarctic (eds Bleil, U. & Thiede, J.) 63\u201376 (Springer, 1990).","DOI":"10.1007\/978-94-009-2029-3_4"},{"key":"2872_CR95","first-page":"159","volume":"151","author":"A Myhre","year":"1995","unstructured":"Myhre, A. et al. Site 909. Proc. Ocean Drill. Program Initial Rep. 151, 159\u2013220 (1995).","journal-title":"Proc. Ocean Drill. Program Initial Rep."},{"key":"2872_CR96","doi-asserted-by":"publisher","first-page":"3815","DOI":"10.1073\/pnas.0710180105","volume":"105","author":"KC Beard","year":"2008","unstructured":"Beard, K. C. The oldest North American primate and mammalian biogeography during the Paleocene\u2013Eocene Thermal Maximum. Proc. Natl Acad. Sci. USA 105, 3815\u20133818 (2008).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2872_CR97","doi-asserted-by":"publisher","first-page":"276","DOI":"10.3732\/ajb.0900195","volume":"97","author":"T Denk","year":"2010","unstructured":"Denk, T., Gr\u00edmsson, F. & Zetter, R. Episodic migration of oaks to Iceland: evidence for a North Atlantic \u2018land bridge\u2019 in the latest Miocene. Am. J. Bot. 97, 276\u2013287 (2010).","journal-title":"Am. J. Bot."},{"key":"2872_CR98","doi-asserted-by":"publisher","first-page":"271","DOI":"10.1144\/GSL.SP.1995.090.01.17","volume":"90","author":"C Wold","year":"1995","unstructured":"Wold, C. Palaeobathymetric reconstruction on a gridded database: the northern North Atlantic and southern Greenland-Iceland-Norwegian Sea. Geol. Soc. Spec. Publ. 90, 271\u2013302 (1995).","journal-title":"Geol. Soc. Spec. Publ."},{"key":"2872_CR99","doi-asserted-by":"publisher","first-page":"55","DOI":"10.1144\/GSL.SP.1998.131.01.04","volume":"131","author":"AT Ramsay","year":"1998","unstructured":"Ramsay, A. T., Smart, C. W. & Zachos, J. C. A model of early to middle Miocene deep ocean circulation for the Atlantic and Indian oceans. Geol. Soc. Spec. Publ. 131, 55\u201370 (1998).","journal-title":"Geol. Soc. Spec. Publ."},{"key":"2872_CR100","doi-asserted-by":"crossref","unstructured":"Denk, T. et al. in Late Cainozoic Floras of Iceland: 15 Million Years of Vegetation and Climate History in the Northern North Atlantic Topics in Geobiology (eds Denk, F. G. T. et al.) 647\u2013668 (Springer, 2011).","DOI":"10.1007\/978-94-007-0372-8_12"},{"key":"2872_CR101","unstructured":"Poore, R. H. Neogene Epeirogeny and the Iceland Plume. PhD thesis, Univ. of Cambridge, (2008)."},{"key":"2872_CR102","doi-asserted-by":"crossref","unstructured":"Eldholm, O., Myhre, A. M., Thiede, J. in Cenozoic Plants and Climates of the Arctic Vol. 127 NATO ASI Series (eds Fisher, H. C. & Boulter, M. C.) 35\u201355 (Springer, 1994).","DOI":"10.1007\/978-3-642-79378-3_4"},{"key":"2872_CR103","doi-asserted-by":"crossref","unstructured":"Thiede, J. & Eldholm, O. in Structure and Development of the Greenland-Scotland Ridge: New Methods and Concepts (eds Saxow, S. et al.) 445\u2013456 (Plenum, 1983).","DOI":"10.1007\/978-1-4613-3485-9_23"},{"key":"2872_CR104","doi-asserted-by":"crossref","unstructured":"Trettin, H. in Geology of the Innuitian Orogen and Arctic Platform of Canada and Greenland Vol. 3 Geology of Canada Series (ed. Trettin, H. P.) 59\u201366 (Geological Survey of Canada, 1991).","DOI":"10.4095\/133959"},{"key":"2872_CR105","doi-asserted-by":"publisher","first-page":"B04402","DOI":"10.1029\/2006JB004308","volume":"112","author":"T Funck","year":"2007","unstructured":"Funck, T., Jackson, H. R., Louden, K. E. & Klingelh\u00f6fer, F. Seismic study of the transform-rifted margin in Davis Strait between Baffin Island (Canada) and Greenland: what happens when a plume meets a transform. J. Geophys. Res. 112, B04402 (2007).","journal-title":"J. Geophys. Res."},{"key":"2872_CR106","doi-asserted-by":"publisher","first-page":"B10401","DOI":"10.1029\/2011JB008942","volume":"117","author":"GN Oakey","year":"2012","unstructured":"Oakey, G. N. & Chalmers, J. A. A new model for the Paleogene motion of Greenland relative to North America: plate reconstructions of the Davis Strait and Nares Strait regions between Canada and Greenland. J. Geophys. Res. 117, B10401 (2012).","journal-title":"J. Geophys. Res."},{"key":"2872_CR107","first-page":"1","volume":"74","author":"NJ Van Strien","year":"1974","unstructured":"Van Strien, N. J. Dicerorhinus sumatrensis (Fischer) the Sumatran or two-horned Asiatic Rhinoceros. a study of literature. Mededelingen Landbouwhogeschool 74, 1\u201382 (1974).","journal-title":"Mededelingen Landbouwhogeschool"},{"key":"2872_CR108","doi-asserted-by":"publisher","first-page":"77","DOI":"10.7557\/2.25.4.1773","volume":"25","author":"FL Miller","year":"2005","unstructured":"Miller, F. L., Barry, S. J. & Calvert, W. A. Sea-ice crossings by caribou in the south-central Canadian Arctic Archipelago and their ecological importance. Rangifer 25, 77\u201388 (2005).","journal-title":"Rangifer"},{"key":"2872_CR109","doi-asserted-by":"publisher","first-page":"805","DOI":"10.1007\/s13280-019-01205-x","volume":"49","author":"C Cuyler","year":"2020","unstructured":"Cuyler, C. et al. Muskox status, recent variation, and uncertain future. Ambio 49, 805\u2013819 (2020).","journal-title":"Ambio"},{"key":"2872_CR110","doi-asserted-by":"publisher","first-page":"686","DOI":"10.1126\/science.1059412","volume":"292","author":"J Zachos","year":"2001","unstructured":"Zachos, J., Pagani, M., Sloan, L., Thomas, E. & Billups, K. Trends, rhythms, and aberrations in global climate 65 Ma to present. Science 292, 686\u2013693 (2001).","journal-title":"Science"},{"key":"2872_CR111","doi-asserted-by":"publisher","first-page":"279","DOI":"10.1038\/nature06588","volume":"451","author":"JC Zachos","year":"2008","unstructured":"Zachos, J. C., Dickens, G. R. & Zeebe, R. E. An early Cenozoic perspective on greenhouse warming and carbon-cycle dynamics. Nature 451, 279\u2013283 (2008).","journal-title":"Nature"},{"key":"2872_CR112","doi-asserted-by":"publisher","first-page":"1851","DOI":"10.1029\/2018PA003433","volume":"34","author":"R Stein","year":"2019","unstructured":"Stein, R. The late Mesozoic-Cenozoic Arctic Ocean climate and sea ice history: a challenge for past and future scientific ocean drilling. Paleoceanogr. Paleoclimatol. 34, 1851\u20131894 (2019).","journal-title":"Paleoceanogr. Paleoclimatol."},{"key":"2872_CR113","doi-asserted-by":"publisher","first-page":"1757","DOI":"10.1016\/j.quascirev.2010.02.010","volume":"29","author":"L Polyak","year":"2010","unstructured":"Polyak, L. et al. History of sea ice in the Arctic. Quat. Sci. Rev. 29, 1757\u20131778 (2010).","journal-title":"Quat. Sci. Rev."},{"key":"2872_CR114","doi-asserted-by":"publisher","first-page":"241","DOI":"10.1029\/96PA00813","volume":"11","author":"KL Brice","year":"1996","unstructured":"Brice, K. L., Arthur, M. A. & Marincovich, L. Jr Late Paleocene Arctic Ocean shallow-marine temperatures from mollusc stable isotopes. Paleoceanography 11, 241\u2013249 (1996).","journal-title":"Paleoceanography"},{"key":"2872_CR115","doi-asserted-by":"publisher","first-page":"25302","DOI":"10.1073\/pnas.2003914117","volume":"117","author":"CL O\u2019Brien","year":"2020","unstructured":"O\u2019Brien, C. L. et al. The enigma of Oligocene climate and global surface temperature evolution. Proc. Natl Acad. Sci. USA 117, 25302\u201325309 (2020).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2872_CR116","doi-asserted-by":"publisher","first-page":"112","DOI":"10.1016\/j.epsl.2007.09.045","volume":"265","author":"AK Tripati","year":"2008","unstructured":"Tripati, A. K. et al. Evidence for glaciation in the northern hemisphere back to 44 Ma from ice-rafted debris in the Greenland Sea. Earth Planet. Sci. Lett. 265, 112\u2013122 (2008).","journal-title":"Earth Planet. Sci. Lett."},{"key":"2872_CR117","doi-asserted-by":"publisher","first-page":"185","DOI":"10.1016\/S0277-3791(97)00076-0","volume":"17","author":"J Thiede","year":"1998","unstructured":"Thiede, J. et al. Late Cenozoic history of the Polar North Atlantic: results from ocean drilling. Quat. Sci. Rev. 17, 185\u2013208 (1998).","journal-title":"Quat. Sci. Rev."},{"key":"2872_CR118","doi-asserted-by":"publisher","DOI":"10.1016\/j.gloplacha.2022.103876","volume":"215","author":"J Gruetzner","year":"2022","unstructured":"Gruetzner, J., Matthiessen, J., Geissler, W. H., Gebhardt, A. C. & Schreck, M. A revised core-seismic integration in the Molloy Basin (ODP Site 909): implications for the history of ice rafting and ocean circulation in the Atlantic-Arctic gateway. Glob. Planet. Change 215, 103876 (2022).","journal-title":"Glob. Planet. Change"},{"key":"2872_CR119","doi-asserted-by":"publisher","first-page":"PA1S03","DOI":"10.1029\/2007PA001476","volume":"23","author":"J Backman","year":"2008","unstructured":"Backman, J. et al. Age model and core-seismic integration for the Cenozoic Arctic Coring Expedition sediments from the Lomonosov Ridge. Paleoceanography 23, PA1S03 (2008).","journal-title":"Paleoceanography"},{"key":"2872_CR120","doi-asserted-by":"publisher","first-page":"601","DOI":"10.1038\/nature04800","volume":"441","author":"K Moran","year":"2006","unstructured":"Moran, K. et al. The Cenozoic palaeoenvironment of the Arctic Ocean. Nature 441, 601\u2013605 (2006).","journal-title":"Nature"},{"key":"2872_CR121","doi-asserted-by":"publisher","first-page":"210","DOI":"10.1038\/ngeo2068","volume":"7","author":"DA Darby","year":"2014","unstructured":"Darby, D. A. Ephemeral formation of perennial sea ice in the Arctic Ocean during the middle Eocene. Nat. Geosci. 7, 210\u2013213 (2014).","journal-title":"Nat. Geosci."},{"key":"2872_CR122","doi-asserted-by":"publisher","first-page":"PA1S05","DOI":"10.1029\/2007PA001483","volume":"23","author":"K St. John","year":"2008","unstructured":"St. John, K. Cenozoic ice-rafting history of the central Arctic Ocean: terrigenous sands on the Lomonosov Ridge. Paleoceanography 23, PA1S05 (2008).","journal-title":"Paleoceanography"},{"key":"2872_CR123","doi-asserted-by":"publisher","first-page":"376","DOI":"10.1038\/nature08163","volume":"460","author":"CE Stickley","year":"2009","unstructured":"Stickley, C. E. et al. Evidence for middle Eocene Arctic sea ice from diatoms and ice-rafted debris. Nature 460, 376\u2013379 (2009).","journal-title":"Nature"},{"key":"2872_CR124","doi-asserted-by":"publisher","first-page":"1383","DOI":"10.1126\/science.aba6853","volume":"369","author":"T Westerhold","year":"2020","unstructured":"Westerhold, T. et al. An astronomically dated record of Earth\u2019s climate and its predictability over the last 66 million years. Science 369, 1383\u20131387 (2020).","journal-title":"Science"},{"key":"2872_CR125","first-page":"3","volume":"124","author":"JJ Eberle","year":"2012","unstructured":"Eberle, J. J. & Greenwood, D. R. Life at the top of the greenhouse Eocene world\u2014a review of the Eocene flora and vertebrate fauna from Canada\u2019s High Arctic. Bulletin 124, 3\u201323 (2012).","journal-title":"Bulletin"},{"key":"2872_CR126","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.earscirev.2012.02.005","volume":"112","author":"MJ Pound","year":"2012","unstructured":"Pound, M. J., Haywood, A. M., Salzmann, U. & Riding, J. B. Global vegetation dynamics and latitudinal temperature gradients during the mid to late Miocene (15.97\u20135.33 Ma). Earth-Sci. Rev. 112, 1\u201322 (2012).","journal-title":"Earth-Sci. Rev."},{"key":"2872_CR127","doi-asserted-by":"publisher","first-page":"47","DOI":"10.7557\/2.12.2.1016","volume":"12","author":"DR Klein","year":"1992","unstructured":"Klein, D. R. Comparative ecological and behavioral adaptations of Ovibos moschatus and Rangifer tarandus. Rangifer 12, 47\u201355 (1992).","journal-title":"Rangifer"},{"key":"2872_CR128","doi-asserted-by":"publisher","first-page":"17258","DOI":"10.3402\/polar.v31i0.17258","volume":"31","author":"B Hansen","year":"2012","unstructured":"Hansen, B. & Aanes, R. Kelp and seaweed feeding by High-Arctic wild reindeer under extreme winter conditions. Polar Res. 31, 17258 (2012).","journal-title":"Polar Res."},{"key":"2872_CR129","doi-asserted-by":"publisher","first-page":"261","DOI":"10.1146\/annurev.es.11.110180.001401","volume":"11","author":"JP Bryant","year":"1980","unstructured":"Bryant, J. P. & Kuropat, P. J. Selection of winter forage by subarctic browsing vertebrates: the role of plant chemistry. Annu. Rev. Ecol. Syst. 11, 261\u2013285 (1980).","journal-title":"Annu. Rev. Ecol. Syst."},{"key":"2872_CR130","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.palaeo.2014.03.006","volume":"401","author":"T Wappler","year":"2014","unstructured":"Wappler, T. et al. Before the \u2018Big Chill\u2019: a preliminary overview of arthropods from the middle Miocene of Iceland (Insecta, Crustacea). Palaeogeogr. Palaeoclimatol. Palaeoecol. 401, 1\u201312 (2014).","journal-title":"Palaeogeogr. Palaeoclimatol. Palaeoecol."},{"key":"2872_CR131","first-page":"1","volume":"111","author":"MY Hassan","year":"1953","unstructured":"Hassan, M. Y. Tertiary faunas from Kap Brewster, East Greenland. Medd. Groenl. 111, 1\u201342 (1953).","journal-title":"Medd. Groenl."},{"key":"2872_CR132","doi-asserted-by":"publisher","first-page":"303","DOI":"10.33799\/jokull2008.58.303","volume":"58","author":"F Gr\u00edmsson","year":"2008","unstructured":"Gr\u00edmsson, F. & S\u00edmonarson, L. A. Upper Tertiary non-marine environments and climatic changes in Iceland. J\u00f6kull 58, 303\u2013314 (2008).","journal-title":"J\u00f6kull"},{"key":"2872_CR133","first-page":"189","volume":"59","author":"LA S\u00edmonarson","year":"1990","unstructured":"S\u00edmonarson, L. A. Fyrstu landspend\u00fdraleifarnar \u00far \u00edslenskum tert\u00ederl\u00f6gum. Natturufraedingurinn 59, 189\u2013195 (1990).","journal-title":"Natturufraedingurinn"},{"key":"2872_CR134","doi-asserted-by":"crossref","unstructured":"Wang, X., Flynn, L. J. & Fortelius, M. Fossil Mammals of Asia: Neogene Biostratigraphy and Chronology (Columbia Univ. Press, 2013).","DOI":"10.7312\/wang15012"},{"key":"2872_CR135","doi-asserted-by":"publisher","first-page":"12","DOI":"10.1017\/S0094837300006254","volume":"5","author":"AK Behrensmeyer","year":"1979","unstructured":"Behrensmeyer, A. K., Western, D. & Boaz, D. E. D. New perspectives in vertebrate paleoecology from a recent bone assemblage. Paleobiology 5, 12\u201321 (1979).","journal-title":"Paleobiology"},{"key":"2872_CR136","doi-asserted-by":"publisher","first-page":"103","DOI":"10.1666\/0094-8373(2000)26[103:TAP]2.0.CO;2","volume":"26","author":"AK Behrensmeyer","year":"2000","unstructured":"Behrensmeyer, A. K., Kidwell, S. M. & Gastaldo, R. A. Taphonomy and paleobiology. Paleobiology 26, 103\u2013147 (2000).","journal-title":"Paleobiology"},{"key":"2872_CR137","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1016\/S1631-0683(02)00005-2","volume":"1","author":"P-O Antoine","year":"2002","unstructured":"Antoine, P.-O., Alf\u00e9rez, F. & I\u00f1igo, C. A new elasmotheriine (Mammalia, Rhinocerotidae) from the Early Miocene of Spain. C. R. Palevol 1, 19\u201326 (2002).","journal-title":"C. R. Palevol"},{"key":"2872_CR138","doi-asserted-by":"publisher","first-page":"183","DOI":"10.58782\/flmnh.kpcf8483","volume":"56","author":"RA Short","year":"2019","unstructured":"Short, R. A., Wallace, S. C. & Emmert, L. G. A new species of Teleoceras (Mammalia, Rhinocerotidae) from the late Hemphillian of Tennessee. Bull. Florida Mus. Nat. Hist. 56, 183\u2013260 (2019).","journal-title":"Bull. Florida Mus. Nat. Hist."},{"key":"2872_CR139","doi-asserted-by":"publisher","first-page":"726","DOI":"10.1093\/sysbio\/syw021","volume":"65","author":"S H\u00f6hna","year":"2016","unstructured":"H\u00f6hna, S. et al. RevBayes: Bayesian phylogenetic inference using graphical models and an interactive model-specification language. Syst. Biol. 65, 726\u2013736 (2016).","journal-title":"Syst. Biol."},{"key":"2872_CR140","doi-asserted-by":"publisher","first-page":"753","DOI":"10.1093\/sysbio\/syu039","volume":"63","author":"S H\u00f6hna","year":"2014","unstructured":"H\u00f6hna, S. et al. Probabilistic graphical model representation in phylogenetics. Syst. Biol. 63, 753\u2013771 (2014).","journal-title":"Syst. Biol."},{"key":"2872_CR141","unstructured":"Barido-Sottani, J. et al. in Phylogenetics in the Genomic Era (eds Scornavacca, C. et al.) 5.2:2\u20135.2:22 (No commercial publisher, 2020)."},{"key":"2872_CR142","doi-asserted-by":"publisher","first-page":"E2957","DOI":"10.1073\/pnas.1319091111","volume":"111","author":"TA Heath","year":"2014","unstructured":"Heath, T. A., Huelsenbeck, J. P. & Stadler, T. The fossilized birth\u2013death process for coherent calibration of divergence-time estimates. Proc. Natl Acad. Sci. USA 111, E2957\u2013E2966 (2014).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2872_CR143","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1016\/j.jtbi.2018.03.005","volume":"447","author":"T Stadler","year":"2018","unstructured":"Stadler, T., Gavryushkina, A., Warnock, R. C., Drummond, A. J. & Heath, T. A. The fossilized birth-death model for the analysis of stratigraphic range data under different speciation modes. J. Theor. Biol. 447, 41\u201355 (2018).","journal-title":"J. Theor. Biol."},{"key":"2872_CR144","doi-asserted-by":"publisher","first-page":"e88","DOI":"10.1371\/journal.pbio.0040088","volume":"4","author":"AJ Drummond","year":"2006","unstructured":"Drummond, A. J., Ho, S. Y. W., Phillips, M. J. & Rambaut, A. Relaxed phylogenetics and dating with confidence. PLoS Biol. 4, e88 (2006).","journal-title":"PLoS Biol."},{"key":"2872_CR145","first-page":"5","volume":"188","author":"P-O Antoine","year":"2002","unstructured":"Antoine, P.-O. Phylog\u00e9nie et \u00e9volution des Elasmotheriina. Mem. Mus. Natl Hist. Nat. 188, 5\u2013350 (2002).","journal-title":"Mem. Mus. Natl Hist. Nat."},{"key":"2872_CR146","doi-asserted-by":"publisher","first-page":"e14156","DOI":"10.1371\/journal.pone.0014156","volume":"5","author":"SS Tobe","year":"2010","unstructured":"Tobe, S. S., Kitchener, A. C. & Linacre, A. M. Reconstructing mammalian phylogenies: a detailed comparison of the cytochrome b and cytochrome oxidase subunit I mitochondrial genes. PLoS ONE 5, e14156 (2010).","journal-title":"PLoS ONE"},{"key":"2872_CR147","doi-asserted-by":"publisher","first-page":"144","DOI":"10.1111\/cla.12524","volume":"39","author":"PA Goloboff","year":"2023","unstructured":"Goloboff, P. A. & Morales, M. E. TNT version 1.6, with a graphical interface for MacOS and Linux, including new routines in parallel. Cladistics 39, 144\u2013153 (2023).","journal-title":"Cladistics"},{"key":"2872_CR148","doi-asserted-by":"publisher","first-page":"295","DOI":"10.1111\/j.1096-0031.1994.tb00179.x","volume":"10","author":"K Bremer","year":"1994","unstructured":"Bremer, K. Branch support and tree stability. Cladistics 10, 295\u2013304 (1994).","journal-title":"Cladistics"},{"key":"2872_CR149","doi-asserted-by":"publisher","first-page":"199","DOI":"10.18814\/epiiugs\/2013\/v36i3\/002","volume":"36","author":"KM Cohen","year":"2013","unstructured":"Cohen, K. M., Finney, S. C., Gibbard, P. L. & Fan, J.-X. The ICS international chronostratigraphic chart. Episodes 36, 199\u2013204 (2013).","journal-title":"Episodes"},{"key":"2872_CR150","doi-asserted-by":"publisher","first-page":"724","DOI":"10.1111\/2041-210X.12081","volume":"4","author":"DW Bapst","year":"2013","unstructured":"Bapst, D. W. A stochastic rate-calibrated method for time-scaling phylogenies of fossil taxa. Methods Ecol. Evol. 4, 724\u2013733 (2013).","journal-title":"Methods Ecol. Evol."},{"key":"2872_CR151","unstructured":"Geraads, D. et al. A database of Old World Neogene and Quaternary rhino-bearing localities. Rhino Resource Center http:\/\/www.rhinoresourcecenter.com\/about\/fossil-rhino-database.php (2021)."},{"key":"2872_CR152","unstructured":"Nash, J. C., Varadhan, R. & Grothendieck, G. optimx: expanded replacement and extension of the \u2018optim\u2019 function. R version v. 2023-10.21 (2023)."},{"key":"2872_CR153","unstructured":"Lalibert\u00e9, E., Legendre, P. & Shipley, B. FD: measuring functional diversity (FD) from multiple traits, and other tools for functional ecology. R version v. 1.0-12 (2014)."},{"key":"2872_CR154","unstructured":"Tierney, L., Rossini, A. J., Li, N. & Sevcikova, H. snow: simple network of workstations. R version v. 0.4-3 (2018)."},{"key":"2872_CR155","unstructured":"Support for Parallel Computation in R v. 4.0.3 (R Development Core Team, 2020); https:\/\/stat.ethz.ch\/R-manual\/R-devel\/library\/parallel\/doc\/parallel.pdf"},{"key":"2872_CR156","unstructured":"Gubian, S., Xiang, Y., Suomela, B. Hoeng, J. & PMP SA GenSA: R functions for generalized simulated annealing. R version v. 1.1.7 (2018)."},{"key":"2872_CR157","unstructured":"Matzke, N. J., Sidje, R. B. & Schmidt, D. rexpokit: R wrappers for EXPOKIT; other matrix functions. R version v. 0.26.6.7 (2020)."},{"key":"2872_CR158","unstructured":"Matzke, N. J. cladoRcpp: C++ implementations of phylogenetic cladogenesis calculations. R version v. 0.15.1 (2018)."},{"key":"2872_CR159","doi-asserted-by":"publisher","first-page":"2299","DOI":"10.1111\/j.0014-3820.2005.tb00940.x","volume":"59","author":"RH Ree","year":"2005","unstructured":"Ree, R. H., Moore, B. R., Webb, C. O. & Donoghue, M. J. A likelihood framework for inferring the evolution of geographic range on phylogenetic trees. Evolution 59, 2299\u20132311 (2005).","journal-title":"Evolution"},{"key":"2872_CR160","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1093\/sysbio\/46.1.195","volume":"46","author":"F Ronquist","year":"1997","unstructured":"Ronquist, F. Dispersal-vicariance analysis: a new approach to the quantification of historical biogeography. Syst. Biol. 46, 195\u2013203 (1997).","journal-title":"Syst. Biol."},{"key":"2872_CR161","doi-asserted-by":"publisher","first-page":"789","DOI":"10.1093\/sysbio\/syt040","volume":"62","author":"MJ Landis","year":"2013","unstructured":"Landis, M. J., Matzke, N. J., Moore, B. R. & Huelsenbeck, J. P. Bayesian analysis of biogeography when the number of areas is large. Syst. Biol. 62, 789\u2013804 (2013).","journal-title":"Syst. Biol."},{"key":"2872_CR162","doi-asserted-by":"publisher","first-page":"887","DOI":"10.1111\/jbi.12898","volume":"44","author":"J Dupin","year":"2017","unstructured":"Dupin, J. et al. Bayesian estimation of the global biogeographical history of the Solanaceae. J. Biogeogr. 44, 887\u2013899 (2017).","journal-title":"J. Biogeogr."},{"key":"2872_CR163","doi-asserted-by":"publisher","unstructured":"Fraser, D. danielleleefraser\/rhinocerotids: v.1.0 (rhinocerotids.v.1.0) Zenodo https:\/\/doi.org\/10.5281\/zenodo.17049570 (2025).","DOI":"10.5281\/zenodo.17049570"}],"container-title":["Nature Ecology & Evolution"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41559-025-02872-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41559-025-02872-8","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41559-025-02872-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,12,5]],"date-time":"2025-12-05T14:02:47Z","timestamp":1764943367000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41559-025-02872-8"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,10,28]]},"references-count":163,"journal-issue":{"issue":"12","published-online":{"date-parts":[[2025,12]]}},"alternative-id":["2872"],"URL":"https:\/\/doi.org\/10.1038\/s41559-025-02872-8","relation":{},"ISSN":["2397-334X"],"issn-type":[{"value":"2397-334X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,10,28]]},"assertion":[{"value":"4 June 2024","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"15 September 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"28 October 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}]}}