Song of my people: dialect differences among sympatric social groups of short-finned pilot whales in Hawai’i

  • Amy M. Van CiseEmail author
  • Sabre D. Mahaffy
  • Robin W. Baird
  • T. Aran Mooney
  • Jay Barlow
Original Article


In many social species, acoustic dialects are used to differentiate among social groups within a local population. These acoustic dialects and their corresponding social groups are often related to distinct foraging behaviors or spatial movement patterns, and it is possible that vocal repertoire variability is one of the proximate mechanisms driving or maintaining genetic and ecological diversity at a subspecies level in social species. Short-finned pilot whales (Globicephala macrorynchus) inhabiting Hawaiian waters have a stable hierarchical social structure, with familial social units associating in larger social clusters within island-associated communities. In this study, we test the hypothesis that sympatric social groups of short-finned pilot whales have acoustically differentiated dialects, which may be used to maintain the social structure. We first examined call composition of social calls collected from photographically identified social clusters of short-finned pilot whales around the Main Hawaiian Islands, using a catalog of manually classified calls, and found that call composition differed among clusters. We then conducted ANOVA and support vector machine (SVM) learning analyses of the acoustic features of social calls. Social clusters were significantly differentiated in their acoustic features, and the SVM classification accuracy was 60%. These results indicate that vocal repertoire reflects social segregation in short-finned pilot whales and may be a driving mechanism of differentiation, potentially contributing to genetic diversity within populations. This suggests divergent acoustic population structure; however, the small sample size in this study decreases the ability to detect acoustic differences among groups. Additional sampling will improve our power to detect acoustic differences among social clusters of Hawaiian pilot whales and improve classification accuracy. The pattern described here highlights the importance of increasing the spatial and temporal resolution of conservation and management plans for this species, in order to conserve subpopulation genetic and social structure.

Significance statement

In some species, vocal repertoires differ among social groups or populations of a species that use the same habitat. These differences, called dialects, are thought to be important to maintaining segregation among groups of animals with overlapping distributions, and in some cases may increase intra-specific ecological or genetic variability. This study is the first to provide evidence that sympatric social clusters of short-finned pilot whales have different vocal repertoires, and that vocal repertoire within groups may change with behavioral context. In terrestrial (e.g., elephants) and marine (e.g., killer whales, sperm whales) species with similarly stable social hierarchies, where acoustic dialects, genetic diversity, and/or ecological variability are linked with social structure, anthropogenic stressors have precipitated rapid declines in abundance with slow or nonexistent recovery. Given the myriad threats faced by short-finned pilot whales in the Hawaiian Islands, including fisheries bycatch, military and commercial anthropogenic noise, and vessel strikes, understanding intra-population social structure and its links with genetic structure and ecological variability is imperative to the proper conservation and management of this species.


Acoustic communication Short-finned pilot whale Dialect Social groups Globicephala macrorhynchus 



We thank Daniel Webster (Cascadia Research Collective) and Max Kaplan (Woods Hole Oceanographic Institution) for their efforts in collecting the acoustic data used in this study. We appreciate the volunteer efforts of Sabrina Pawlek, Madelaine Xochi Perez, and Lisa Valtierra in manually tracing calls included in this study. We also thank Andrew Solow, who provided important insight for statistical analyses used in this study. Finally, we thank Lisa Levin, Bill Hodgkiss, James Fowler, Phil Morin, and Ron Burton for their valuable input in the analysis and interpretation of this study.

Funding information

Funding for data collection was provided by grants from the Pacific Islands Fisheries Science Center and Office of Naval Research, as well as Commander, Pacific Fleet. The SoundTrap was purchased with funding from the SIO/NSF IGERT fellowship program to AVC. DMON data collection was funded by the Office of Naval Research (award numbers: N000141110612 (TAM RWB) and N00014-15-1-2299 (MAR); Program Manager Michael J. Weise) and WHOI Marine Mammal Center and the Sawyer and Penzance Endowed Funds to TAM.

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

Ethical approval

All applicable international, national, and/or institutional guidelines for the care and use of animals were followed. All procedures performed in this study were in accordance with the ethical standards of the institution or practice at which the studies were conducted. These data were collected under NMFS permit #15530 to RWB and WHOI Institutional Animal Care and Use Committee approval (BI15245.00).


  1. Alves F, Quérouil S, Dinis A, Nicolau C, Ribeiro C, Freitas L, Kaufmann M, Fortuna C (2013) Population structure of short-finned pilot whales in the oceanic archipelago of Madeira based on photo-identification and genetic analyses: implications for conservation. Aquat Conserv 23:758–776. CrossRefGoogle Scholar
  2. Amézquita A, Lima AP, Jehle R, Castellanos L, Ramos Ó, Crawford AJ, Gasser H, Hödl W (2009) Calls, colours, shape, and genes: a multi-trait approach to the study of geographic variation in the Amazonian frog Allobates femoralis. Biol J Linn Soc 98:826–838. CrossRefGoogle Scholar
  3. Ansmann IC, Parra GJ, Lanyon JM, Seddon JM (2012) Fine-scale genetic population structure in a mobile marine mammal: inshore bottlenose dolphins in Moreton Bay, Australia. Mol Ecol 21:4472–4485. CrossRefPubMedGoogle Scholar
  4. Baird RW (2016) The lives of Hawai’i’s dolphins and whales: natural history and conservation. University of Hawai’i Press, Honolulu, Hawai’iGoogle Scholar
  5. Baird RW, Webster DL, Aschettino JM, Schorr G, McSweeney DJ (2013) Odontocete cetaceans around the Main Hawaiian Islands: habitat use and relative abundance from small-boat sighting surveys. Aquat Mamm 39:253–269. CrossRefGoogle Scholar
  6. Baird RW, Abrams PA, Dill LM (1992) Possible indirect interactions between transient and resident killer whales: implications for the evolution of foraging specializations in the genus Orcinus. Oecologia 89:125–132. CrossRefPubMedGoogle Scholar
  7. Baker MC, Cunningham MA (1985) The biology of bird-song dialects. Behav Brain Sci 8:85. CrossRefGoogle Scholar
  8. Balcazar NE, Tripovich JS, Klinck H, Nieukirk SL, Mellinger DK, Dziak RP, Rogers TL (2015) Calls reveal population structure of blue whales across the southeast Indian Ocean and southwest Pacific Ocean. J Mammal 96:1184–1193. CrossRefPubMedPubMedCentralGoogle Scholar
  9. Beecher MD, Campbell SE (2005) The role of unshared songs in singing interactions between neighbouring song sparrows. Anim Behav 70:1297–1304Google Scholar
  10. Bennett KP, Campbell C (2000) Support vector machines: hype or hallelujah? SIGKDD Explor 2:1–13CrossRefGoogle Scholar
  11. Cantor M, Shoemaker LG, Cabral RB, Flores CO, Varga M, Whitehead H (2015) Multilevel animal societies can emerge from cultural transmission. Nat Commun 6:8091. CrossRefPubMedPubMedCentralGoogle Scholar
  12. Conner DA (1982) Dialects versus geographic variation in mammalian vocalizations. Anim Behav 30:297–298CrossRefGoogle Scholar
  13. Crance JL, Bowles AE, Garver A (2014) Evidence for vocal learning in juvenile male killer whales, Orcinus orca, from an adventitious cross-socializing experiment. J Exp Biol 217:1229–1237. CrossRefPubMedGoogle Scholar
  14. Crockford C, Herbinger I, Vigilant L, Boesch C (2004) Wild chimpanzees produce group-specific calls: a case for vocal learning? Ethology 110:221–243. CrossRefGoogle Scholar
  15. de la Torre S, Snowdon CT (2009) Dialects in pygmy marmosets? Population variation in call structure. Am J Primatol 71:333–342. CrossRefPubMedGoogle Scholar
  16. Deecke VB, Barrett-Lennard LG, Spong P, Ford JKB (2010) The structure of stereotyped calls reflects kinship and social affiliation in resident killer whales (Orcinus orca). Naturwissenschaften 97:513–518. CrossRefPubMedGoogle Scholar
  17. Deecke VB, Ford JKB, Slater PJB (2005) The vocal behaviour of mammal-eating killer whales: communicating with costly calls. Anim Behav 69:395–405. CrossRefGoogle Scholar
  18. Deecke VB, Ford JKB, Spong P (2000) Dialect change in resident killer whales: implications for vocal learning and cultural transmission. Anim Behav 60:629–638. CrossRefPubMedGoogle Scholar
  19. Esser KH, Schubert J (1998) Vocal dialects in the lesser spear-nosed bat Phyllostomus discolor. Naturwissenschaften 85:347–349. CrossRefGoogle Scholar
  20. Filatova OA, Deecke VB, Ford JKB, Matkin CO, Barrett-Lennard LG, Guzeev MA, Burdin AM, Hoyt E (2012) Call diversity in the North Pacific killer whale populations: implications for dialect evolution and population history. Anim Behav 83:595–603. CrossRefGoogle Scholar
  21. Filatova OA, Fedutin ID, Nagaylik MM, Burdin AM, Hoyt E (2009) Usage of monophonic and biphonic calls by free-ranging resident killer whales (Orcinus orca) in Kamchatka, Russian Far East. Acta Ethol 12:37–44. CrossRefGoogle Scholar
  22. Filatova OA, Guzeev MA, Fedutin ID, Burdin AM, Hoyt E (2013) Dependence of killer whale (Orcinus orca) acoustic signals on the type of activity and social context. Biol Bull 40:790–796. CrossRefGoogle Scholar
  23. Fischer J, Hammerschmoidt K, Todt D (1998) Local variation in Barbary macaque shrill barks. Anim Behav 56:623–629CrossRefGoogle Scholar
  24. Foote AD (2012) Investigating ecological speciation in non-model organisms: a case study on killer whale ecotypes. Evol Ecol Res 14:447–465Google Scholar
  25. Foote AD, Newton J, Piertney SB, Willerslev E, Gilbert MTP (2009) Ecological, morphological and genetic divergence of sympatric North Atlantic killer whale populations. Mol Ecol 18:5207–5217. CrossRefPubMedGoogle Scholar
  26. Foote AD, Morin PA (2016) Genome-wide SNP data suggest complex ancestry of sympatric North Pacific killer whale ecotypes. Heredity 117:316–325Google Scholar
  27. Foote AD, Nystuen JA (2008) Variation in call pitch among killer whale ecotypes. J Acoust Soc Am 123:1747–1752. CrossRefPubMedGoogle Scholar
  28. Foote AD, Osborne RW, Hoelzel AR (2004) Environment: whale-call response to masking boat noise. Nature 428:910. CrossRefPubMedGoogle Scholar
  29. Foote AD, Vijay N, Avila-Arcos M et al (2016) Genome-culture coevolution promotes rapid divergence in the killer whale. Nat Commun 7:11693. CrossRefPubMedPubMedCentralGoogle Scholar
  30. Ford JKB (1991) Vocal traditions among resident killer whales (Orcinus orca) in coastal waters of British Columbia. Can J Zool 69:1454–1483CrossRefGoogle Scholar
  31. Ford JKB, Fisher HD (1982) Killer whale (Orcinus orca) dialects as an indicator of stocks in British Columbia. Rep IWC 32:671–679Google Scholar
  32. Freeberg TM, Dunbar RIM, Ord TJ (2012) Social complexity as a proximate and ultimate factor in communicative complexity. Philos Trans R Soc B 367:1785–1801. CrossRefGoogle Scholar
  33. Garland EC, Goldizen AW, Lilley MS, Rekdahl ML, Garrigue C, Constantine R, Hauser ND, Poole MM, Robbins J, Noad MJ (2015) Population structure of humpback whales in the western and central South Pacific Ocean as determined by vocal exchange among populations. Conserv Biol 29:1198–1207. CrossRefPubMedGoogle Scholar
  34. Gero S, Whitehead H, Rendell L (2016) Individual, unit and vocal clan level identity cues in sperm whale codas. R Soc Open Sci 3:150372. CrossRefPubMedPubMedCentralGoogle Scholar
  35. Gillespie D, Caillat M, Gordon J, White P (2013) Automatic detection and classification of odontocete whistles. J Acoust Soc Am 134:2427–2437. CrossRefPubMedGoogle Scholar
  36. Gillespie D, Gordon J, McHugh R, Mclaren D, Mellinger D, Redmond P, Thode A, Trinder P, Deng XY (2009) PAMGUARD: Semiautomated, open source software for real-time acoustic detection and localisation of cetaceans. Proc Ins Ac 30:2547Google Scholar
  37. Green S (1975) Dialects in Japanese monkeys: vocal learning and cultural transmission of locale-specific vocal behavior? Z Tierpsychol 38:304–314. CrossRefPubMedGoogle Scholar
  38. Ivkovich T, Filatova OA, Burdin AM, Sato H, Hoyt E (2010) The social organization of resident-type killer whales (Orcinus orca) in Avacha Gulf, Northwest Pacific, as revealed through association patterns and acoustic similarity. Mamm Biol 75:198–210CrossRefGoogle Scholar
  39. Janik VM, Slater PJB (1997) Vocal learning in mammals. Adv Study Behav 26:59–100CrossRefGoogle Scholar
  40. Jensen FH, Perez JM, Johnson M, Soto NA, Madsen PT (2011) Calling under pressure: short-finned pilot whales make social calls during deep foraging dives. Proc R Soc Lond B 278:3017–3025. CrossRefGoogle Scholar
  41. Kaplan MB, Mooney TA, Sayigh LS, Baird RW (2014) Repeated call types in Hawaiian melon-headed whales (Peponocephala electra). J Acoust Soc Am 136:1394–1401. CrossRefPubMedGoogle Scholar
  42. Kasuya T, Miyashita T, Kasamatsu F (1988) Segregation of two forms of short-finned pilot whales off the Pacific coast of Japan. Sci Rep Whal Res Inst 39:77–90Google Scholar
  43. Kershenbaum A, Ilany A, Blaustein L, Geffen E (2012) Syntactic structure and geographical dialects in the songs of male rock hyraxes. Proc R Soc Lond B 279:2974–2981. CrossRefGoogle Scholar
  44. Kessler SE, Radespiel U, Hasiniaina AIF, Leliveld LMC, Nash LT, Zimmermann E (2014) Modeling the origins of mammalian sociality: moderate evidence for matrilineal signatures in mouse lemur vocalizations. Front Zool 11:14. CrossRefPubMedPubMedCentralGoogle Scholar
  45. Kindt R, Coe R (2005) Tree diversity analysis. A manual and software for common statistical methods for ecological and biodiversity studies. World Agroforestry Centre (ICRAF), NairobiGoogle Scholar
  46. Krebs JR, Kroodsma DE (1980) Repertoires and geographical variation in bird song. Adv Study Behav 11:143–177. CrossRefGoogle Scholar
  47. Lachlan RF, Slater PJB (1999) The maintenance of vocal learning by gene-culture interaction: the cultural trap hypothesis. Proc R Soc Lond B 266:701–706. CrossRefGoogle Scholar
  48. Lemasson A, Gautier JP, Hausberger M (2003) Vocal similarities and social bonds Campbell’s monkey (Cercopithecus campbelli). CR Biol 326:1185–1193. CrossRefGoogle Scholar
  49. Lovell SF, Ross LM (2005) Individual recognition of neighbors by song in a suboscine bird, the alder flycatcher Empidonax alnorum. Behav Ecol Sociobiol 57:623–630. CrossRefGoogle Scholar
  50. MacDougall-Shackleton EA, MacDougall-Shackleton SA (2001) Cultural and genetic evolution in mountain white-crowned sparrows: song dialects are associated with population structure. Evolution 55:2568–2575. CrossRefPubMedGoogle Scholar
  51. Madsen PT, Jensen FH, Carder D, Ridgway S (2012) Dolphin whistles: a functional misnomer revealed by heliox breathing. Biol Lett 8:211–213. CrossRefPubMedGoogle Scholar
  52. Mahaffy SD, Baird RW, McSweeney DJ, Webster DL, Schorr GS (2015) High site fidelity, strong associations, and long-term bonds: short-finned pilot whales off the island of Hawai’i. Mar Mammal Sci 31:1427–1451. CrossRefGoogle Scholar
  53. Martien KK, Chivers SJ, Baird RW, Archer FI, Gorgone AM, Hancock-Hanser BL, Mattila D, McSweeney DJ, Oleson EM, Palmer C, Pease VL, Robertson KM, Schorr GS, Schultz MB, Webster DL, Taylor BL (2014) Nuclear and mitochondrial patterns of population structure in North Pacific false killer whales (Pseudorca crassidens). J Hered 105:611–626. CrossRefPubMedGoogle Scholar
  54. McComb K, Semple S (2005) Coevolution of vocal communication and sociality in primates. Biol Lett 1:381–388. CrossRefPubMedPubMedCentralGoogle Scholar
  55. McCullagh P, Nelder JA (1989) Generalized linear models. Chapman and Hall, LondonCrossRefGoogle Scholar
  56. McDonald MA, Mesnick SL, Hildebrand JA (2006) Biogeographic characterisation of blue whale song worldwide: using song to identify populations. J Cetacean Res Manag 8:55–65Google Scholar
  57. Meyer D, Dimitriadou E, Hornik K, Weingessel A, Leisch F (2018) E1071: misc functions of the Department of Statistics. Probability Theory Group (formerly: E1071) TU, Wien Google Scholar
  58. Mitani JC, Hasegawa T, Gros-Louis J, Marler P, Byrne R (1992) Dialects in wild chimpanzees? Am J Primatol 27:233–243. CrossRefGoogle Scholar
  59. Mundinger PC (1980) Animal cultures and a general theory of cultural evolution. Ethol Sociobiol 1:183–223. CrossRefGoogle Scholar
  60. Murray SO, Mercado E, Roitblat HL (1998) Characterizing the graded structure of false killer whale (Pseudorca crassidens) vocalizations. J Acoust Soc Am 104:1679–1688 CrossRefGoogle Scholar
  61. Nottebohm F (1969) The song of the chingolo, Zonotrichia capensis, in Argentina: description and evaluation of a system of dialects. Condor 71:299–315CrossRefGoogle Scholar
  62. Oremus M, Gales R, Dalebout ML et al (2009) Worldwide mitochondrial DNA diversity and phylogeography of pilot whales (Globicephala spp.). Biol J Linn Soc 98:729–744. CrossRefGoogle Scholar
  63. Oswald JN, Oswald M (2013) ROCCA (Real-time Odontocete Call Classification Algorithm) user’s manual. Prepared for Naval Facilities Engineering Command Atlantic, Norfolk, Virginia under HDR Environmental, Operations and Construction, Inc. Contract No. CON005-4394-009, Subproject 1647. Bio-Waves Inc., Encinitas, CAGoogle Scholar
  64. Papale E, Azzolin M, Gannier A, Lammers MO, Martin VM, Oswald J, Perez-Gil M, Giacoma C (2013) Geographic variability in the acoustic parameters of striped dolphin’s (Stenella coeruleoalba) whistles. J Acoust Soc Am 133:1126–1134CrossRefGoogle Scholar
  65. Parreira BR, Chikhi L (2015) On some genetic consequences of social structure, mating systems, dispersal, and sampling. P Natl Acad Sci USA 112:E3318–E3326. CrossRefGoogle Scholar
  66. Perla BS, Slobodchikoff CN (2002) Habitat structure and alarm call dialects in Gunnison’s prairie dog (Cynomys gunnisoni). Behav Ecol 13:844–850. CrossRefGoogle Scholar
  67. Pilot M, Dahlheim ME, Hoelzel AR (2010) Social cohesion among kin, gene flow without dispersal and the evolution of population genetic structure in the killer whale (Orcinus orca). J Evol Biol 23:20–31. CrossRefPubMedGoogle Scholar
  68. Podos J (2010) Acoustic discrimination of sympatric morphs in Darwin’s finches: a behavioural mechanism for assortative mating? Philos Trans R Soc B 365:1031–1039. CrossRefGoogle Scholar
  69. Prohl H, Koshy RA, Mueller U, Rand AS, Ryan MJ (2006) Geographic variation of genetic and behavioral traits in northern and southern Tungara frogs. Evolution 60:1669–1679. CrossRefPubMedGoogle Scholar
  70. R Core Team (2016) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria Google Scholar
  71. Rendell L, Mesnick SL, Dalebout ML, Burtenshaw J, Whitehead H (2012) Can genetic differences explain vocal dialect variation in sperm whales, Physeter macrocephalus? Behav Genet 42:332–343. CrossRefPubMedGoogle Scholar
  72. Rendell LE, Whitehead H (2003) Vocal clans in sperm whales (Physeter macrocephalus). Proc R Soc Lond B 270:225–231. CrossRefGoogle Scholar
  73. Riesch R, Barrett-Lennard LG, Ellis GM, Ford JKB, Deecke VB (2012) Cultural traditions and the evolution of reproductive isolation: ecological speciation in killer whales? Biol J Linn Soc 106:1–17. CrossRefGoogle Scholar
  74. Riesch R, Ford JKB, Thomsen F (2006) Stability and group specificity of stereotyped whistles in resident killer whales, Orcinus orca, off British Columbia. Anim Behav 71:79–91. CrossRefGoogle Scholar
  75. Sayigh L, Quick N, Hastie G, Tyack P (2013) Repeated call types in short-finned pilot whales, Globicephala macrorhynchus. Mar Mammal Sci 29:312–324. CrossRefGoogle Scholar
  76. Slabbekoorn H, Smith TB (2002) Bird song, ecology and speciation. Philos Trans R Soc B 357:493–503. CrossRefGoogle Scholar
  77. Soha JA, Nelson DA, Parker PG (2004) Genetic analysis of song dialect populations in Puget Sound white-crowned sparrows. Behav Ecol 15:636–646. CrossRefGoogle Scholar
  78. Stoddard PK (1996) Vocal recognition of neighbors by territorial passerines. In: Ecology and Evolution of Acoustic Communication in Birds. Comstock Pub, New York, pp 356–374Google Scholar
  79. Van Cise AM, Martien KK, Mahaffy SD, Baird RW, Webster DL, Fowler JH, Oleson EM, Morin PA (2017b) Familial social structure and socially-driven genetic differentiation in Hawaiian short-finned pilot whales. Mol Ecol 26:6730–6741. CrossRefPubMedGoogle Scholar
  80. Van Cise AM, Morin PA, Baird RW, Lang AR, Robertson KM, Chivers SJ, Brownell RL, Martien KK (2016) Redrawing the map: mtDNA provides new insight into the distribution and diversity of short-finned pilot whales in the Pacific Ocean. Mar Mammal Sci 32:1177–1199. CrossRefGoogle Scholar
  81. Van Cise AM, Roch MA, Baird RW, Mooney TA, Barlow J (2017a) Acoustic differentiation of Shiho- and Naisa-type short-finned pilot whales in the Pacific Ocean. J Acoust Soc Am 141:737–748. CrossRefPubMedGoogle Scholar
  82. Wade PR, Reeves RR, Mesnick SL (2012) Social and behavioural factors in cetacean responses to overexploitation: are Odontocetes less ‘resilient’ than mysticetes? J Mar Biol 15:e567276. CrossRefGoogle Scholar
  83. Watkins WA (1968) The harmonic interval: fact or artifact in spectral analysis of pulse trains. Proc Sec Symp Mar Bio-Acoust 2:15–43. CrossRefGoogle Scholar
  84. Weiß BM, Symonds H, Spong P, Ladich F (2007) Intra- and intergroup vocal behavior in resident killer whales, Orcinus orca. J Acoust Soc Am 122:3710–3716. CrossRefPubMedGoogle Scholar
  85. Whitehead H (1998) Cultural selection and genetic diversity in matrilineal whales. Science 282:1708–1711. CrossRefPubMedGoogle Scholar
  86. Whitehead H (2007) Learning, climate and the evolution of cultural capacity. J Theor Biol 245:341–350. CrossRefPubMedGoogle Scholar
  87. Whitehead H (2017) Gene-culture coevolution in whales and dolphins. P Natl Acad Sci USA 114:7814–7821. CrossRefGoogle Scholar
  88. Whitehead H, Ford JKB (2018) Consequences of culturally-driven ecological specialization: killer whales and beyond. J Theor Biol 456:279–294. CrossRefPubMedGoogle Scholar
  89. Winn HE, Thompson TJ, Cummings WC (1981) Song of the humpback whale—population comparisons. Behav Ecol Sociobiol 8:41–46CrossRefGoogle Scholar
  90. Wright TF, Wilkinson GS (2001) Population genetic structure and vocal dialects in an amazon parrot. Proc R Soc Lond B 268:609–616. CrossRefGoogle Scholar
  91. Yoshino H, Armstrong KN, Izawa M, Yokoyama J, Kawata M (2008) Genetic and acoustic population structuring in the Okinawa least horseshoe bat: are intercolony acoustic differences maintained by vertical maternal transmission? Mol Ecol 17:4978–4991. CrossRefPubMedGoogle Scholar
  92. Yurk H, Barrett-Lennard LG, Ford JKB, Matkin CO (2002) Cultural transmission within maternal lineages: vocal clans in resident killer whales in southern Alaska. Anim Behav 63:1103–1119. CrossRefGoogle Scholar
  93. Zwamborn EMJ, Whitehead H (2017) Repeated call sequences and behavioural context in long-finned pilot whales off Cape Breton, Nova Scotia, Canada. Bioacoustics 26:169–183. CrossRefGoogle Scholar

Copyright information

© This is a U.S. Government work and not under copyright protection in the US; foreign copyright protection may apply 2018

Authors and Affiliations

  • Amy M. Van Cise
    • 1
    • 2
    Email author
  • Sabre D. Mahaffy
    • 3
  • Robin W. Baird
    • 3
  • T. Aran Mooney
    • 2
  • Jay Barlow
    • 1
    • 4
  1. 1.Scripps Institution of Oceanography, UCSDLa JollaUSA
  2. 2.Woods Hole Oceanographic InstitutionWoods HoleUSA
  3. 3.Cascadia Research CollectiveOlympiaUSA
  4. 4.NOAA Southwest Fisheries Science CenterLa JollaUSA

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