{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T16:19:31Z","timestamp":1785773971779,"version":"3.56.0"},"reference-count":102,"publisher":"The Royal Society","issue":"2075","license":[{"start":{"date-parts":[[2026,7,15]],"date-time":"2026-07-15T00:00:00Z","timestamp":1784073600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,7,15]],"date-time":"2026-07-15T00:00:00Z","timestamp":1784073600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,7,15]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Pinnipeds (true seals, eared seals and walruses) are the only mammals that can hear effectively in both air and water. How and when they achieved the ability to negotiate such contrasting auditory media remains unknown. Here, we apply 3D shape and phylogenetic comparative analyses to a large dataset of extant and extinct caniform carnivorans (119 species, 217 specimens) to study the emergence of amphibious hearing in pinnipeds despite significant evolutionary constraints. We find support for the cavernous tissue as a functional and evolutionary mechanism for amphibious hearing. This tissue fills with blood during diving to equalize air pressure in the ear, enabling a shift from in-air to underwater hearing by matching the ear\u2019s acoustic impedance to that of the surrounding water. We found that early diverging freshwater pinnipeds had impaired hearing underwater. The first ancestral marine pinnipeds could hear amphibiously but with limited hearing ranges, probably reducing attenuation and harm from loud underwater sounds. Subsequently, otariids (eared seals) and phocids (true seals) independently acquired middle ear adaptations that expanded their underwater hearing range. This iterative evolution probably facilitated the exploration of novel auditory adaptive zones by crown pinnipeds, resulting in rare acoustic abilities like ultrasonic singing, vocal learning and rhythm.<\/jats:p>","DOI":"10.1098\/rspb.2026.0178","type":"journal-article","created":{"date-parts":[[2026,7,15]],"date-time":"2026-07-15T00:19:50Z","timestamp":1784074790000},"source":"Crossref","is-referenced-by-count":1,"title":["The origin and evolution of amphibious hearing in pinnipeds"],"prefix":"10.1098","volume":"293","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-7256-2393","authenticated-orcid":false,"given":"James P.","family":"Rule","sequence":"first","affiliation":[{"name":"Natural History Museum, 1","place":["London,, UK"],"department":["Science Group,"]},{"name":"Monash University, 2","place":["Clayton,, Australia"],"department":["School of Biological Sciences,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9492-8859","authenticated-orcid":false,"given":"Travis","family":"Park","sequence":"additional","affiliation":[{"name":"Natural History Museum, 1","place":["London,, UK"],"department":["Science Group,"]},{"name":"Monash University, 2","place":["Clayton,, Australia"],"department":["School of Biological Sciences,"]},{"name":"Museums Victoria, 3","place":["Melbourne,, Australia"],"department":["Museums Victoria Research Institute,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Moganavalli","family":"Kattan","sequence":"additional","affiliation":[{"name":"Natural History Museum, 1","place":["London,, UK"],"department":["Science Group,"]},{"name":"Imperial College London, 4","place":["Ascot,, UK"],"department":["Department of Life Sciences,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6697-7151","authenticated-orcid":false,"given":"Camille","family":"Groh\u00e9","sequence":"additional","affiliation":[{"name":"UMR 7262, CNRS & Universit\u00e9 de Poitiers, 5","place":["Poitiers,, France"],"department":["Laboratoire PALEVOPRIM,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3970-5270","authenticated-orcid":false,"given":"Roxana","family":"Taszus","sequence":"additional","affiliation":[{"name":"University of Liverpool, 6","place":["Liverpool,, UK"],"department":["Department of Musculoskeletal Biology,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0008-9297-7455","authenticated-orcid":false,"given":"Stephanie M.","family":"Palmer","sequence":"additional","affiliation":[{"name":"Lake Erie College of Osteopathic Medicine, 7","place":["Greensburg,, PA,, USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6848-1208","authenticated-orcid":false,"given":"David P.","family":"Hocking","sequence":"additional","affiliation":[{"name":"Tasmanian Museum and Art Gallery, 8","place":["Hobart,, Tasmania,, Australia"],"department":["Department of Zoology,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6214-9850","authenticated-orcid":false,"given":"Justin W.","family":"Adams","sequence":"additional","affiliation":[{"name":"Museums Victoria, 3","place":["Melbourne,, Australia"],"department":["Museums Victoria Research Institute,"]},{"name":"Monash University, 9","place":["Clayton,, Australia"],"department":["Anatomy and Developmental Biology,"]},{"name":"University of Johannesburg, 10","place":["Johannesburg,, South Africa"],"department":["Palaeo-Research Institute,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4078-4693","authenticated-orcid":false,"given":"Alistair R.","family":"Evans","sequence":"additional","affiliation":[{"name":"Monash University, 2","place":["Clayton,, Australia"],"department":["School of Biological Sciences,"]},{"name":"Museums Victoria, 3","place":["Melbourne,, Australia"],"department":["Museums Victoria Research Institute,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4416-5224","authenticated-orcid":false,"given":"Ian G.","family":"Brennan","sequence":"additional","affiliation":[{"name":"Natural History Museum, 1","place":["London,, UK"],"department":["Science Group,"]},{"name":"Australian National University, 11","place":["Acton,, Australia"],"department":["Division of Ecology & Evolution,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5605-9069","authenticated-orcid":false,"given":"Tahlia I.","family":"Pollock","sequence":"additional","affiliation":[{"name":"Monash University, 2","place":["Clayton,, Australia"],"department":["School of Biological Sciences,"]},{"name":"University of Bristol, 12","place":["Bristol,, UK"],"department":["Palaeobiology Research Group,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0713-5226","authenticated-orcid":false,"given":"Daniela","family":"Sanfelice","sequence":"additional","affiliation":[{"name":"Instituto Federal de Educa\u00e7\u00e3o Ci\u00eancia e Tecnologia do Rio Grande do Sul, 13","place":["Porto Alegre,, Brazil"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1029-4001","authenticated-orcid":false,"given":"Felix G.","family":"Marx","sequence":"additional","affiliation":[{"name":"Museum of New Zealand Te Papa Tongarewa, 14","place":["Wellington,, New Zealand"],"department":["Natural History,"]},{"name":"University of Otago, 15","place":["Dunedin,, New Zealand"],"department":["Department of Geology,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5329-4063","authenticated-orcid":false,"given":"Naoki","family":"Kohno","sequence":"additional","affiliation":[{"name":"National Museum of Nature and Science, 16","place":["Tsukuba,, Japan"],"department":["Department of Paleontology and Anthropology,"]},{"name":"University of Tsukuba, 17","place":["Tsukuba,, Japan"],"department":["Graduate School of Life and Environmental Sciences,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2064-4631","authenticated-orcid":false,"given":"Martin","family":"Sabol","sequence":"additional","affiliation":[{"name":"Comenius University in Bratislava, 18","place":["Bratislava,, Slovakia"],"department":["Department of Geology & Palaeontology,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2434-2542","authenticated-orcid":false,"given":"Alexander","family":"Stoessel","sequence":"additional","affiliation":[{"name":"Friedrich Schiller University Jena, 19","place":["Jena,, Germany"],"department":["Institute of Zoology and Evolutionary Research,"]},{"name":"Max Planck Institute for Evolutionary Anthropology, 20","place":["Leipzig,, Germany"],"department":["Department of Archaeogenetics,"]},{"name":"Max Planck Institute for Evolutionary Anthropology, 21","place":["Leipzig,, Germany"],"department":["Department of Human Origins,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4705-3591","authenticated-orcid":false,"given":"John J.","family":"Flynn","sequence":"additional","affiliation":[{"name":"American Museum of Natural History, 22","place":["New York,, NY,, USA"],"department":["Division of Paleontology,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4919-8655","authenticated-orcid":false,"given":"Natalie","family":"Cooper","sequence":"additional","affiliation":[{"name":"Natural History Museum, 1","place":["London,, UK"],"department":["Science Group,"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"175","published-online":{"date-parts":[[2026,7,15]]},"reference":[{"key":"2026071420194562200_R1","first-page":"267","article-title":"The origins of the cochlea and impedance matching hearing in synapsids","volume":"61","author":"Laa\u00df","year":"2015","journal-title":"Acta Palaeontol. 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Biol."},{"key":"2026071420194562200_R8","doi-asserted-by":"crossref","first-page":"1550","DOI":"10.1121\/1.2221532","article-title":"Underwater auditory localization by a swimming harbor seal (Phoca vitulina)","volume":"120","author":"Bodson","year":"2006","journal-title":"J. Acoust. Soc. Am."},{"key":"2026071420194562200_R9","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1121\/1.5114809","article-title":"Underwater sound localization abilities of harbor seals (Phoca vitulina) for high-frequency noise band stimuli in the median plane","volume":"146","author":"Byl","year":"2019","journal-title":"J. Acoust. Soc. Am."},{"key":"2026071420194562200_R10","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1016\/j.heares.2015.10.002","article-title":"High-frequency hearing in seals and sea lions","volume":"331","author":"Cunningham","year":"2016","journal-title":"Hear. Res."},{"key":"2026071420194562200_R11","doi-asserted-by":"crossref","first-page":"1","DOI":"10.3389\/fevo.2021.644497","article-title":"Strange seal sounds: claps, slaps, and multimodal pinniped rhythms","volume":"9","author":"Verga","year":"2021","journal-title":"Front. Ecol. Evol"},{"key":"2026071420194562200_R12","doi-asserted-by":"crossref","first-page":"351","DOI":"10.1007\/978-3-031-77022-7_5","volume-title":"Marine mammal acoustics in a noisy ocean","author":"Parsons","year":"2025"},{"key":"2026071420194562200_R13","doi-asserted-by":"crossref","first-page":"2263","DOI":"10.1121\/1.2775424","article-title":"Underwater localization of pure tones by harbor seals (Phoca vitulina)","volume":"122","author":"Bodson","year":"2007","journal-title":"J. Acoust. Soc. Am."},{"key":"2026071420194562200_R14","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1093\/zoolinnean\/zlad014","article-title":"The shape of water: adaptations of cochlea morphology in seals and otters","volume":"199","author":"Taszus","year":"2023","journal-title":"Zool. J. Linn. Soc."},{"key":"2026071420194562200_R15","doi-asserted-by":"crossref","first-page":"63","DOI":"10.15407\/zoo2021.01.063","article-title":"Can you hear me now? A comparative survey of pinniped auditory apparatus morphology","volume":"55","author":"Koper","year":"2021","journal-title":"ZD."},{"key":"2026071420194562200_R16","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1007\/978-3-031-77022-7_8","volume-title":"Marine mammal acoustics in a noisy ocean","author":"Houser","year":"2025"},{"key":"2026071420194562200_R17","first-page":"307","volume-title":"Functional anatomy of marine mammals","author":"Repenning","year":"1972"},{"key":"2026071420194562200_R18","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1007\/s00359-013-0813-y","article-title":"Comparative assessment of amphibious hearing in pinnipeds","volume":"199","author":"Reichmuth","year":"2013","journal-title":"J. Comp. Physiol. A Neuroethol. Sens. Neural. Behav. Physiol."},{"key":"2026071420194562200_R19","doi-asserted-by":"crossref","first-page":"3463","DOI":"10.1121\/1.2372712","article-title":"High-frequency hearing in phocid and otariid pinnipeds: an interpretation based on inertial and cochlear constraints (L)","volume":"120","author":"Hemil\u00e4","year":"2006","journal-title":"J. Acoust. Soc. Am."},{"key":"2026071420194562200_R20","first-page":"117","volume-title":"Biology of marine mammals","author":"Wartzok","year":"1999"},{"key":"2026071420194562200_R21","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1007\/978-0-387-78365-9_7","volume-title":"Principles of marine bioacoustics","author":"Au","year":"2008"},{"key":"2026071420194562200_R22","doi-asserted-by":"crossref","first-page":"689","DOI":"10.1080\/00016480152583629","article-title":"How can the hooded seal dive to a depth of 1000 m without rupturing its tympanic membrane? A morphological and functional study","volume":"121","author":"Stenfors","year":"2001","journal-title":"Acta Otolaryngol."},{"key":"2026071420194562200_R23","doi-asserted-by":"crossref","DOI":"10.1126\/science.157.3784.99.a","article-title":"Seal ears","volume":"157","author":"Mohl","year":"1967","journal-title":"Science"},{"key":"2026071420194562200_R24","doi-asserted-by":"crossref","first-page":"768","DOI":"10.1126\/science.153.3737.768","article-title":"Pressure regulation in the middle ear cavity of sea lions: a possible mechanism","volume":"153","author":"Odend\u2019hal","year":"1966","journal-title":"Science"},{"key":"2026071420194562200_R25","first-page":"437","article-title":"Uber ein corpus cavernosum tympanicum beim seehnnd","volume":"33","author":"Tandler","year":"1899","journal-title":"Monatschr. 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Paleontol."},{"key":"2026071420194562200_R37","doi-asserted-by":"crossref","first-page":"457","DOI":"10.3389\/fevo.2019.00457","article-title":"A total evidence phylogenetic analysis of pinniped phylogeny and the possibility of parallel evolution within a monophyletic framework","volume":"7","author":"Paterson","year":"2020","journal-title":"Front. Ecol. Evol."},{"key":"2026071420194562200_R38","first-page":"1212","article-title":"Charting the course of pinniped evolution: insights from molecular phylogeny and fossil record integration","volume":"78","author":"Park","year":"2024","journal-title":"Evolution (N Y)"},{"key":"2026071420194562200_R39","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1146\/annurev-earth-082517-010009","article-title":"The origin and evolutionary biology of pinnipeds: seals, sea lions, and walruses","volume":"46","author":"Berta","year":"2018","journal-title":"Annu. Rev. Earth Planet. 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