{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,21]],"date-time":"2026-10-21T12:22:12Z","timestamp":1792585332034,"version":"4.5.0"},"reference-count":55,"publisher":"Public Library of Science (PLoS)","issue":"7","license":[{"start":{"date-parts":[[2020,7,30]],"date-time":"2020-07-30T00:00:00Z","timestamp":1596067200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100006769","name":"Russian Science Foundation","doi-asserted-by":"publisher","award":["19-14-00037"],"award-info":[{"award-number":["19-14-00037"]}],"id":[{"id":"10.13039\/501100006769","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.plosone.org"],"crossmark-restriction":false},"short-container-title":["PLoS ONE"],"DOI":"10.1371\/journal.pone.0236749","type":"journal-article","created":{"date-parts":[[2020,7,30]],"date-time":"2020-07-30T20:33:29Z","timestamp":1596141209000},"page":"e0236749","update-policy":"https:\/\/doi.org\/10.1371\/journal.pone.corrections_policy","source":"Crossref","is-referenced-by-count":12,"title":["Independent acoustic variation of the higher- and lower-frequency components of biphonic calls can facilitate call recognition and social affiliation in killer whales"],"prefix":"10.1371","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-1533-4807","authenticated-orcid":true,"given":"Olga A.","family":"Filatova","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2020,7,30]]},"reference":[{"issue":"1871","key":"pone.0236749.ref001","doi-asserted-by":"crossref","first-page":"20172171","DOI":"10.1098\/rspb.2017.2171","article-title":"Imitation of novel conspecific and human speech sounds in the killer whale (Orcinus orca).","volume":"285","author":"JZ Abramson","year":"2018","journal-title":"Proc Royal Soc B Biol Sci"},{"key":"pone.0236749.ref002","doi-asserted-by":"crossref","first-page":"1454","DOI":"10.1139\/z91-206","article-title":"Vocal traditions among resident killer whales (Orcinus orca) in coastal waters of British Columbia.","volume":"69","author":"JKB Ford","year":"1991","journal-title":"Can J Zool"},{"issue":"2","key":"pone.0236749.ref003","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1017\/S0140525X0100396X","article-title":"Culture in whales and dolphins","volume":"24","author":"L Rendell","year":"2001","journal-title":"Behav Brain Sci"},{"issue":"3","key":"pone.0236749.ref004","doi-asserted-by":"crossref","first-page":"150","DOI":"10.1002\/evan.10111","article-title":"Do animals have culture?","volume":"12","author":"KN Laland","year":"2003","journal-title":"Evol Anthropol"},{"issue":"2","key":"pone.0236749.ref005","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1163\/1568539X-00003417","article-title":"Social complexity and cultural transmission of dialects in killer whales.","volume":"154","author":"OA Filatova","year":"2017","journal-title":"Behav"},{"key":"pone.0236749.ref006","doi-asserted-by":"crossref","first-page":"513","DOI":"10.1007\/s00114-010-0657-z","article-title":"The structure of stereotyped calls reflects kinship and social affiliation in resident killer whales (Orcinus orca).","volume":"97","author":"VB Deecke","year":"2010","journal-title":"Naturwissenschaften"},{"key":"pone.0236749.ref007","doi-asserted-by":"crossref","first-page":"449","DOI":"10.1007\/s00359-005-0085-2","article-title":"Diversity in sound pressure levels and estimated active space of resident killer whale vocalizations","volume":"192","author":"PJO Miller","year":"2006","journal-title":"J Comp Physiol A"},{"key":"pone.0236749.ref008","doi-asserted-by":"crossref","first-page":"3932","DOI":"10.1121\/1.2722056","article-title":"Caller sex and orientation influence spectral characteristics of \u201ctwo-voice\u201d stereotyped calls produced by free-ranging killer whales.","volume":"121","author":"PJO Miller","year":"2007","journal-title":"J Acoust Soc Amer"},{"key":"pone.0236749.ref009","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1007\/s10211-009-0056-7","article-title":"Usage of monophonic and biphonic calls by free-ranging resident killer whales (Orcinus orca) in Kamchatka, Russian Far East.","volume":"12","author":"OA Filatova","year":"2009","journal-title":"Acta Ethol"},{"key":"pone.0236749.ref010","doi-asserted-by":"crossref","first-page":"599","DOI":"10.1111\/j.1439-0310.2008.01496.x","article-title":"Temporal and contextual patterns of killer whale (Orcinus orca) call type production.","volume":"114","author":"AD Foote","year":"2008","journal-title":"Ethol"},{"key":"pone.0236749.ref011","doi-asserted-by":"crossref","first-page":"3710","DOI":"10.1121\/1.2799907","article-title":"Intra-and intergroup vocal behavior in resident killer whales, Orcinus orca","volume":"122","author":"BM Wei\u00df","year":"2007","journal-title":"J Acoust Soc Am"},{"key":"pone.0236749.ref012","doi-asserted-by":"crossref","first-page":"1 71","DOI":"10.1080\/09524622.1998.9753394","article-title":"Subharmonics, biphonation and deterministic chaos in mammal vocalization.","volume":"9","author":"I Wilden","year":"1998","journal-title":"Bioacoustics."},{"key":"pone.0236749.ref013","doi-asserted-by":"crossref","first-page":"1435","DOI":"10.1121\/1.1289208","article-title":"Nonlinear phenomena and their anatomical basis in the natural howling of a female dog-wolf breed","volume":"108","author":"T Riede","year":"2000","journal-title":"J Acoust Soc Am"},{"key":"pone.0236749.ref014","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1080\/09524622.2002.9753490","article-title":"Biphonation as a prominent feature of dhole Cuon alpinus sound.","volume":"13","author":"IA Volodin","year":"2002","journal-title":"Bioacoustics"},{"key":"pone.0236749.ref015","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1046\/j.1439-0310.2001.00630.x","article-title":"Acoustic features of female chacma baboon barks.","volume":"107","author":"J Fischer","year":"2001","journal-title":"Ethol"},{"key":"pone.0236749.ref016","doi-asserted-by":"crossref","first-page":"2114","DOI":"10.1121\/1.1528930","article-title":"Laryngeal biomechanics and vocal communication in the squirrel monkey (Saimiri boliviensis).","volume":"113","author":"CH Brown","year":"2003","journal-title":"J Acoust Soc Am"},{"key":"pone.0236749.ref017","doi-asserted-by":"crossref","first-page":"277","DOI":"10.1002\/ajp.20078","article-title":"Nonlinear acoustics in pant hoots of common chimpanzees (Pan troglodytes): frequency jumps, subharmonics, biphonation, and deterministic chaos.","volume":"64","author":"T Riede","year":"2004","journal-title":"Am J Primatol"},{"issue":"3","key":"pone.0236749.ref018","doi-asserted-by":"crossref","first-page":"1365","DOI":"10.1121\/1.2756263","article-title":"Nonlinear phenomena in the vocalizations of North Atlantic right whales (Eubalaena glacialis) and killer whales (Orcinus orca).","volume":"122","author":"RB Tyson","year":"2007","journal-title":"J Acoust Soc Am"},{"issue":"1","key":"pone.0236749.ref019","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1111\/mms.12452","article-title":"Two\u2010component calls in short\u2010finned pilot whales (Globicephala macrorhynchus).","volume":"34","author":"N Quick","year":"2018","journal-title":"Mar Mam Sci"},{"key":"pone.0236749.ref020","doi-asserted-by":"crossref","first-page":"407","DOI":"10.1006\/anbe.2001.1912","article-title":"Calls out of chaos: the adaptive significance of nonlinear phenomena in mammalian vocal production","volume":"63","author":"WT Fitch","year":"2002","journal-title":"Anim Behav"},{"key":"pone.0236749.ref021","doi-asserted-by":"crossref","first-page":"1081","DOI":"10.1098\/rspb.2000.1112","article-title":"Penguins use the twovoice system to recognize each other","volume":"267","author":"T Aubin","year":"2000","journal-title":"Proc R Soc Lond B Biol Sci"},{"issue":"8","key":"pone.0236749.ref022","doi-asserted-by":"crossref","first-page":"815","DOI":"10.1111\/j.1439-0310.2006.01231.x","article-title":"Biphonation may function to enhance individual recognition in the dhole, Cuon alpinus.","volume":"112","author":"EV Volodina","year":"2006","journal-title":"Ethol"},{"issue":"9","key":"pone.0236749.ref023","doi-asserted-by":"crossref","first-page":"e106317","DOI":"10.1371\/journal.pone.0106317","article-title":"Identification and characteristics of signature whistles in wild bottlenose dolphins (Tursiops truncatus) from Namibia.","volume":"9","author":"HJ Kriesell","year":"2014","journal-title":"PLoS One."},{"issue":"3","key":"pone.0236749.ref024","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1080\/09524622.2015.1041158","article-title":"Biphonic calls as signature whistles in a free-ranging bottlenose dolphin.","volume":"24","author":"E Papale","year":"2015","journal-title":"Bioacoustics"},{"key":"pone.0236749.ref025","doi-asserted-by":"crossref","first-page":"629","DOI":"10.1006\/anbe.2000.1454","article-title":"Dialect change in resident killer whales: implications for vocal learning and cultural transmission","volume":"60","author":"VB Deecke","year":"2000","journal-title":"Anim Behav"},{"key":"pone.0236749.ref026","doi-asserted-by":"crossref","first-page":"1103","DOI":"10.1006\/anbe.2002.3012","article-title":"Cultural transmission within maternal lineages: vocal clans in resident killer whales in southern Alaska","volume":"63","author":"H Yurk","year":"2002","journal-title":"Anim Behav"},{"key":"pone.0236749.ref027","doi-asserted-by":"crossref","first-page":"1037","DOI":"10.1139\/z95-124","article-title":"Pod-specific call repertoires and compound calls of killer whales, Orcinus orca Linnaeus, 1758, in the waters of northern Norway","volume":"73","author":"H Strager","year":"1995","journal-title":"Can J Zool"},{"issue":"2","key":"pone.0236749.ref028","doi-asserted-by":"crossref","first-page":"191228","DOI":"10.1098\/rsos.191228","article-title":"Cold call: the acoustic repertoire of Ross Sea killer whales (Orcinus orca, Type C) in McMurdo Sound, Antarctica.","volume":"7","author":"R Wellard","year":"2020","journal-title":"Royal Soc Open Sci."},{"key":"pone.0236749.ref029","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1121\/1.4922704","article-title":"Killer whale call frequency is similar across the oceans, but varies across sympatric ecotypes","volume":"138","author":"OA Filatova","year":"2015","journal-title":"J Acoust Soc Amer"},{"key":"pone.0236749.ref030","doi-asserted-by":"crossref","first-page":"908","DOI":"10.1101\/gr.102954.109","article-title":"Complete mitchondrial genome phylogeographic analysis of killer whales (Orcinus orca) indicates multiple species.","volume":"20","author":"PA Morin","year":"2010","journal-title":"Genome Res"},{"issue":"1","key":"pone.0236749.ref031","first-page":"3","article-title":"Photo-identification of individual killer whales.","volume":"17","author":"MA Bigg","year":"1983","journal-title":"Whalewatcher"},{"key":"pone.0236749.ref032","volume-title":"The Killer Whales of Eastern Kamchatka","author":"AM Burdin","year":"2006"},{"key":"pone.0236749.ref033","doi-asserted-by":"crossref","first-page":"198","DOI":"10.1016\/j.mambio.2009.03.006","article-title":"The social organization of resident-type killer whales (Orcinus orca) in Avacha Gulf, Northwest Pacific, as revealed through association patterns and acoustic similarity.","volume":"75","author":"TV Ivkovich","year":"2010","journal-title":"Mamm Biol."},{"key":"pone.0236749.ref034","doi-asserted-by":"crossref","first-page":"261","DOI":"10.1080\/09524622.2007.9753581","article-title":"The structure of the discrete call repertoire of killer whales Orcinus orca from Southeast Kamchatka","volume":"16","author":"OA Filatova","year":"2007","journal-title":"Bioacoustics."},{"key":"pone.0236749.ref035","first-page":"1169","article-title":"A catalogue of discrete calls of resident killer whales (Orcinus orca) from the Avacha Gulf of Kamchatka Peninsula.","volume":"83","author":"OA Filatova","year":"2004","journal-title":"Zoologicheskii Journal"},{"key":"pone.0236749.ref036","doi-asserted-by":"crossref","first-page":"595","DOI":"10.1016\/j.anbehav.2011.12.013","article-title":"Call diversity in the North Pacific killer whale populations: implications for dialect evolution and population history.","volume":"83","author":"OA Filatova","year":"2012","journal-title":"Anim Behav"},{"issue":"1","key":"pone.0236749.ref037","doi-asserted-by":"crossref","first-page":"645","DOI":"10.1121\/1.2139067","article-title":"Automated categorization of bioacoustic signals: avoiding perceptual pitfalls","volume":"119","author":"VB Deecke","year":"2006","journal-title":"J Acoust Soc Amer"},{"key":"pone.0236749.ref038","doi-asserted-by":"crossref","first-page":"1134","DOI":"10.1121\/1.427121","article-title":"Killer whale (Orcinus orca) hearing: Auditory brainstem response and behavioral audiograms.","volume":"106","author":"MD Szymanski","year":"1999","journal-title":"J Acoust Soc Am"},{"issue":"4","key":"pone.0236749.ref039","doi-asserted-by":"crossref","first-page":"2387","DOI":"10.1121\/1.4979116","article-title":"Killer whale (Orcinus orca) behavioral audiograms.","volume":"141","author":"BK Branstetter","year":"2017","journal-title":"J Acoust Soc Amer"},{"issue":"3","key":"pone.0236749.ref040","doi-asserted-by":"crossref","first-page":"262","DOI":"10.1007\/s00265-002-0508-9","article-title":"Mixed-directionality of killer whale stereotyped calls: A direction of movement cue?","volume":"52","author":"PJO Miller","year":"2002","journal-title":"Behav Ecol Sociobiol."},{"key":"pone.0236749.ref041","doi-asserted-by":"crossref","first-page":"1613","DOI":"10.1121\/1.3676694","article-title":"Directional properties of bottlenose dolphin (Tursiops truncatus) clicks, burst-pulse, and whistle sounds.","volume":"131","author":"BK Branstetter","year":"2012","journal-title":"J Acoust Soc Amer"},{"issue":"5","key":"pone.0236749.ref042","doi-asserted-by":"crossref","first-page":"3755","DOI":"10.1121\/1.4967369","article-title":"Physical constraints of cultural evolution of dialects in killer whales","volume":"140","author":"OA Filatova","year":"2016","journal-title":"J Acoust Soc Amer"},{"key":"pone.0236749.ref043","volume-title":"Underwater acoustics: Analysis, design and performance of sonar","author":"RP Hodges","year":"2011"},{"issue":"2","key":"pone.0236749.ref044","doi-asserted-by":"crossref","first-page":"394","DOI":"10.1111\/j.1748-7692.2002.tb01045.x","article-title":"Underwater noise of whale\u2010watching boats and potential effects on killer whales (Orcinus orca), based on an acoustic impact model.","volume":"18","author":"C Erbe","year":"2002","journal-title":"Mar Mam Sci."},{"issue":"1","key":"pone.0236749.ref045","doi-asserted-by":"crossref","first-page":"EL27","DOI":"10.1121\/1.3040028","article-title":"Speaking up: killer whales (Orcinus orca) increase their call amplitude in response to vessel noise.","volume":"125","author":"MM Holt","year":"2009","journal-title":"J Acoust Soc Amer"},{"key":"pone.0236749.ref046","doi-asserted-by":"crossref","first-page":"e1657","DOI":"10.7717\/peerj.1657","article-title":"Ship noise extends to frequencies used for echolocation by endangered killer whales.","volume":"4","author":"S Veirs","year":"2016","journal-title":"PeerJ"},{"issue":"1","key":"pone.0236749.ref047","doi-asserted-by":"crossref","first-page":"92","DOI":"10.1121\/1.3664100","article-title":"Underwater radiated noise from modern commercial ships.","volume":"131","author":"MF McKenna","year":"2012","journal-title":"J Acoust Soc Amer."},{"issue":"5","key":"pone.0236749.ref048","doi-asserted-by":"crossref","first-page":"2875","DOI":"10.1121\/1.2890745","article-title":"Mathematics of pulsed vocalizations with application to killer whale biphonation","volume":"123","author":"JC Brown","year":"2008","journal-title":"J Acoust Soc Amer."},{"key":"pone.0236749.ref049","doi-asserted-by":"crossref","first-page":"509","DOI":"10.1098\/rsbl.2006.0525","article-title":"Killer whales are capable of vocal learning.","volume":"2","author":"AD Foote","year":"2006","journal-title":"Biol Lett."},{"issue":"15","key":"pone.0236749.ref050","doi-asserted-by":"crossref","first-page":"2001","DOI":"10.1163\/1568539X-00003317","article-title":"Cultural evolution of killer whale calls: background, mechanisms and consequences.","volume":"152","author":"OA Filatova","year":"2015","journal-title":"Behav"},{"key":"pone.0236749.ref051","doi-asserted-by":"crossref","first-page":"34","DOI":"10.1016\/j.beproc.2013.06.008","article-title":"Is killer whale dialect evolution random?","volume":"99","author":"OA Filatova","year":"2013","journal-title":"Behav Proc."},{"key":"pone.0236749.ref052","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1016\/0162-3095(80)90008-4","article-title":"Animal cultures and a general theory of cultural evolution","volume":"1","author":"PC Mundinger","year":"1980","journal-title":"Ethol Sociobiol"},{"key":"pone.0236749.ref053","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1016\/0140-1750(85)90042-9","article-title":"The relation between biological and cultural evolution","volume":"8","author":"CJ Lumsden","year":"1985","journal-title":"J Soc Biol Struct"},{"key":"pone.0236749.ref054","doi-asserted-by":"crossref","first-page":"481","DOI":"10.1098\/rsbl.2006.0517","article-title":"The influence of social affiliation on individual vocal signatures of northern resident killer whales (Orcinus orca).","volume":"2","author":"AE Nousek","year":"2006","journal-title":"Biol Lett."},{"key":"pone.0236749.ref055","volume-title":"Vocal culture and social stability in resident killer whales (Orcinus orca).","author":"H Yurk","year":"2005"}],"container-title":["PLOS ONE"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0236749","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2020,7,30]],"date-time":"2020-07-30T20:33:58Z","timestamp":1596141238000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0236749"}},"subtitle":[],"editor":[{"given":"William David","family":"Halliday","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2020,7,30]]},"references-count":55,"journal-issue":{"issue":"7","published-online":{"date-parts":[[2020,7,30]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pone.0236749","relation":{},"ISSN":["1932-6203"],"issn-type":[{"value":"1932-6203","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,7,30]]}}}