Pilot Whales: Delphinid Matriarchies in Deep Seas

  • Jim BoranEmail author
  • Sara Heimlich
Part of the Ethology and Behavioral Ecology of Marine Mammals book series (EBEMM)


Pilot whales are distributed worldwide with two species, the cold-water long-finned pilot whale, Globicephala melas, and the warm-water short-finned pilot whale, G. macrorhynchus. Long-finned pilot whales have an anti-tropical distribution in separated northern and southern hemisphere groupings, with the North Pacific group extinct, while short-finned pilot whales occur in all tropical oceans. These two species exhibit different life history patterns, with long-finned pilot whales maturing faster and reproducing up to the end of life and dying sooner, while short-finned pilot whales have long maturation (esp. for males) and a post-reproductive phase where females live up to 30 years beyond the birth of their last calf. Photo-identification studies of short-finned pilot whales in Macaronesia (the Canary Islands, Azores, and Madeira) show long-term relationships maintained over hundreds of kilometers and exhibit a wide variety of site fidelity patterns. Mixing between core residents and visiting transients in high productivity areas suggest more fluid interactions than those observed for resident ecotype killer whales or orca (Orcinus orca) that maintain isolated groups from other, co-occurring ecotypes. Four features of orca matrilineal society, (1) matrilineal associations between females, (2) male natal philopatry, (3) extra-group mating or natal exogamy, and (4) post-reproductive lifespan, are compared with short-finned pilot whales, and we conclude that pilot whale societies have numerous matrilineal features, albeit less specialized than those exhibited by orca. Divergent prey preferences most likely account for these differences.


Pilot whale Photo-ID Matrilineal Orca Macaronesia 


  1. Aguilar Soto N, Johnson MP, Madsen PT, Díaz F, Domínguez I, Brito A, Tyack P (2008) Cheetahs of the deep sea: deep foraging sprints in short-finned pilot whales off Tenerife (Canary Islands). J Anim Ecol 77(5):936–947. CrossRefPubMedGoogle Scholar
  2. Alves F (2013) Population structure, habitat use and conservation of short-finned pilot whales (Globicephala macrorhynchus) in the Archipelago of Madeira. PhD Thesis. Universidade da MadeiraGoogle Scholar
  3. Alves F, Quérouil S, Dinis A, Nicolau C, Ribeiro C, Freitas L et al (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 Mar Freshwat Ecosyst 23(5):758–776. CrossRefGoogle Scholar
  4. Alves F, Dinis A, Nicolau C, Ribeiro C, Kaufmann M, Fortuna C, Freitas L (2015) Survival and abundance of short-finned pilot whales in the archipelago of Madeira, NE Atlantic. Mar Mamm Sci 31(1):106–121. CrossRefGoogle Scholar
  5. Alves F, Alessandrini A, Servidio A, Mendonça AS, Hartman KL, Prieto R et al (2019) Complex biogeographical patterns support an ecological connectivity network of a large marine predator in the north-east Atlantic. Divers Distrib 25:269–284.
  6. Amos B, Schltterer C, Tautz D (1993) Social structure of pilot whales revealed by analytical DNA profiling. Science 260(5108):670–672. CrossRefGoogle Scholar
  7. Andersen LW, Siegismund HR (1994) Genetic evidence for migration of males between schools of the long-finned pilot whale Globicephala melas. Mar Ecol Prog Ser 105:1–7CrossRefGoogle Scholar
  8. Augusto JF (2017) Social structure of the pilot whales (Globicephala melas) off Cape Breton, Nova Scotia, Canada. Dalhousie UniversityGoogle Scholar
  9. Augusto JF, Frasier TR, Whitehead H (2017) Social structure of long-finned pilot whales (Globicephala melas) off northern Cape Breton Island, Nova Scotia. Behaviour 154(5):509–540. CrossRefGoogle Scholar
  10. Baird RW (2016) The lives of Hawai“i”s dolphins and whales: natural history and conservation. University of Hawai’i Press, HonoluluGoogle Scholar
  11. Baird RW, Dill LM (1996) Ecological and social determinants of group size in transient killer whales. Behav Ecol 7(4):408–416. CrossRefGoogle Scholar
  12. Baird RW, Whitehead H (2000) Social organization of mammal-eating killer whales: group stability and dispersal patterns. Can J Zool 78:2096–2105CrossRefGoogle Scholar
  13. Baird RW, Borsani JF, Hanson MB, Tyack PL (2002) Diving and night-time behavior of long-finned pilot whales in the Ligurian sea. Mar Ecol Prog Ser 237:301–305. CrossRefGoogle Scholar
  14. Balcomb KC (2018) The Center for Whale Research. Accessed 20 Aug 2018
  15. Balcomb KC, Boran JR, Heimlich SL (1982) Killer whales in greater puget sound. Rep Int Whal Comm 32:681–685Google Scholar
  16. Barham EG (1966) Deep scattering layer migration and composition: observations from a diving saucer. Science 151(3716):1399–1403. CrossRefPubMedGoogle Scholar
  17. Barrett-Lennard LG (2000) Population structure and mating patterns of killer whales (Orcinus orca) as revealed by DNA analysis. PhD Thesis. The University of British ColumbiaGoogle Scholar
  18. Barton ED, Aristegui J, Tett P, Canton M, García-Braun J, Hernández-León S et al (1998) The transition zone of the Canary Current upwelling region. Prog Oceanogr 41(4):455–504. CrossRefGoogle Scholar
  19. Behar DM, Villems R, Soodyall H, Blue-Smith J, Pereira L, Metspalu E et al (2008) The dawn of human matrilineal diversity. Am J Hum Genet 82(5):1130–1140. CrossRefPubMedPubMedCentralGoogle Scholar
  20. Bigg MA, Olesiuk PF, Ellis GM (1990) Social organization and genealogy of resident killer whales (Orcinus orca) in the coastal waters of British Columbia and Washington State. Rep Int Whal Comm 12:383–405Google Scholar
  21. Boran J, Heimlich S (1999) Social learning in cetaceans: hunting, hearing and hierarchies. In: Box H, Gibson K (eds) Mammalian social learning. Cambridge University Press, Cambridge, pp 282–307.
  22. Brent LJN, Franks DW, Foster EA, Balcomb KC, Cant MA, Croft DP (2015) Ecological knowledge, leadership, and the evolution of menopause in killer whales. Curr Biol 25(6):746–750. CrossRefGoogle Scholar
  23. Carrillo M, Ritter F (2010) Increasing numbers of ship strikes in the Canary Islands: proposals for immediate action to reduce risk of vessel-whale collisions. J Cetacean Res Manag 11(2):131–138Google Scholar
  24. Chapais B, Girard M, Primi G (1991) Non-kin alliances, and the stability of matrilineal dominance relations in Japanese macaques. Anim Behav 41(3):481–491. CrossRefGoogle Scholar
  25. Clutton-Brock TH (1984) Reproductive effort and terminal investment in iteroparous animals. Am Nat 123(2):212–229CrossRefGoogle Scholar
  26. Clutton-Brock TH (1989) Female transfer and inbreeding avoidance in social mammals. Nature 337:70–72CrossRefGoogle Scholar
  27. Clutton-Brock TH, Lukas D (2012) The evolution of social philopatry and dispersal in female mammals. Mol Ecol 21(3):472–492. CrossRefPubMedGoogle Scholar
  28. Connor RC (2000) Group living in whales and dolphins. In: Mann J, Connor RC, Tyack PL, Whitehead H (eds) Cetacean societies: field studies of dolphins and whales. University of Chicago Press, Chicago, pp 199–218Google Scholar
  29. Davies JL (1963) The antitropical factor in cetacean speciation. Evolution 17:107–116CrossRefGoogle Scholar
  30. de Stephanis R, Verborgh P, Pérez S, Esteban R, Minvielle-Sebastia L, Guinet C (2008) Long-term social structure of long-finned pilot whales (Globicephala melas) in the Strait of Gibraltar. Acta Ethol 11:81–94. CrossRefGoogle Scholar
  31. Desportes G, Mouritsen R (1988) Diet of the pilot whale, Globicephala melas, around the Faroe Islands. ICES Report C.M. 1988/N: 12. CopenhagenGoogle Scholar
  32. Desportes G, Andersen LW, Aspholm PE, Bloch D, Mouritsen R (1992) A note about a male-only pilot whale school observed in Faroe Islands. Fróđskaparrit 40:31–37Google Scholar
  33. Donovan GP, Lockyer CH, Martin AR (1993) Biology of northern hemisphere pilot whales. Rep Int Whal Comm 14:x+479Google Scholar
  34. Doubleday ZA, Prowse TAA, Arkhipkin A, Pierce GJ, Semmens J, Steer M et al (2016) Global proliferation of cephalopods. Curr Biol 26(10):R406–R407. CrossRefPubMedGoogle Scholar
  35. Findlay KP, Best PB, Ross GJB, Cockcroft VG (1992) The distribution of small odontocete cetaceans off the coasts of South Africa and Namibia. S Afr J Mar Sci 12(1):237–270. CrossRefGoogle Scholar
  36. Foote AD (2008) Mortality rate acceleration and post-reproductive lifespan in matrilineal whale species. Biol Lett 4(2):189–191. CrossRefPubMedPubMedCentralGoogle Scholar
  37. Ford JKB, Fisher HD (1983) Group-specific dialects of killer whales (Orcinus orca) in British Columbia. In: Payne RS (ed) Communication and behavior of whales. Westview Press, Boulder CO, pp 129–161Google Scholar
  38. Ford JKB, Ellis GM, Balcomb KC (2000) Killer whales. University of British Columbia Press, VancouverGoogle Scholar
  39. Ford JKB, Ellis GM, Olesiuk PF, Balcomb KC (2010) Linking killer whale survival and prey abundance: food limitation in the oceans’ apex predator? Biol Lett 6(1):139–142. CrossRefGoogle Scholar
  40. Ford MJ, Hanson MB, Hempelmann JA, Ayres KL, Emmons CK, Schorr GS et al (2011) Inferred paternity and male reproductive success in a killer whale (Orcinus orca) population. J Hered 102(5):537–553. CrossRefPubMedGoogle Scholar
  41. Foster EA, Franks DW, Mazzi S, Darden SK, Balcomb KC, Ford JKB, Croft DP (2012) Adaptive prolonged postreproductive life span in killer whales. Science 337(6100):1313. CrossRefPubMedPubMedCentralGoogle Scholar
  42. Fullard KJ, Early G, Heide-Jorgensen MP, Bloch D, Rosing-Asvid A, Amos W (2000) Population structure of long-finned pilot whales in the North Atlantic: a correlation with sea surface temperature? Mol Ecol 9(7):949–958. CrossRefPubMedGoogle Scholar
  43. Furuichi T (2011) Female contributions to the peaceful nature of bonobo society. Evol Anthropol 20:131–142. CrossRefPubMedGoogle Scholar
  44. Gannon DP, Read AJ, Craddock JE, Fristrup KM, Nicolas JR (1997) Feeding ecology of long-finned pilot whales Globicephala melas in the western North Atlantic. Mar Ecol Prog Ser 148(1–3):1–10. CrossRefGoogle Scholar
  45. Gibson-Lonsdale JJ (1990) Pilot whaling in the Faroe Islands–its history and present significance. Mammal Rev 20(1):44–52. CrossRefGoogle Scholar
  46. Gray JE (1846) On the cetaceous animals. In: Richardson EJ, Gray JE (eds) The zoology of the voyage of H.M.S. Erebus and Terror, vol 1. Mammalia, birds. Longmans, Brown, Green and Longmans, London, pp 13–53Google Scholar
  47. Greenwood PJ (1980) Mating systems, philopatry and dispersal in birds and mammals. Anim Behav 28(1960):1140–1162. CrossRefGoogle Scholar
  48. Hamilton WD (1964) The genetical evolution of social behavior. J Theor Biol 7:17–52CrossRefGoogle Scholar
  49. Hartny-Mills L (2015) Site fidelity, social structure and spatial distribution of short-finned pilot whales, Globicephala macrorhynchus, off the South West Coast of Tenerife. PhD Thesis, University of PortsmouthGoogle Scholar
  50. Heide-Jørgensen MP, Bloch D, Stefansson E, Mikkelsen B, Ofstad LH, Dietz R (2002) Diving behaviour of long-finned pilot whales Globicephala melas around the Faroe Islands. Wildl Biol 8(4):307–313CrossRefGoogle Scholar
  51. Heimlich-Boran JR (1986a) Fishery correlations with the occurrence of killer whales in greater Puget Sound. In: Kirkevold B, Lockard J (eds) Behavioral biology of killer whales. A. R. Liss, New York, pp 113–131Google Scholar
  52. Heimlich-Boran SL (1986b) Cohesive relationship among Puget Sound killer whales. In: Kirkevold B, Lockard JS (eds) Behavioral biology of killer whales. A. R. Liss, New York, pp 251–284Google Scholar
  53. Heimlich-Boran JR (1988a) Behavioral ecology of killer whales (Orcinus orca), in the Pacific Northwest. Can J Zool 66(3):565–578. CrossRefGoogle Scholar
  54. Heimlich-Boran SL (1988b) Association patterns and social dynamics of killer whales (Orcinus orca) in Greater Puget Sound. MA Thesis. San Jose State UniversityGoogle Scholar
  55. Heimlich-Boran JR (1993) Social organisation of the short-finned pilot whale, Globicephala macrorhynchus, with special reference to the comparative social ecology of delphinids. PhD Thesis. University of Cambridge.
  56. Johnson DDP, Kays R, Blackwell PG, MacDonald DW (2002) Does the resource dispersion hypothesis explain group living? Trends Ecol Evol 17(12):563–570CrossRefGoogle Scholar
  57. Kasuya T (1975) Past occurrence of Globicephala melaena in the western North Pacific. Sci Rep Whales Res Inst 27(27):95–110Google Scholar
  58. Kasuya T, Marsh H (1984) Life history and reproductive biology of the short-finned Pilot whale, Globicephala macrorhynchus, off the Pacific coast of Japan. Rep Int Whal Comm 6:259–310. Google Scholar
  59. Kasuya T, Sergeant DE, Tanaka K (1988) Re-examination of life history parameters of long-finned pilot whales in the Newfoundland waters Canada. Sci Rep Whales Res Inst Tokyo 39:103–120Google Scholar
  60. LeDuc RG, Perrin WF, Dizon AE (1999) Phylogenetic relationships among the delphinid cetaceans based on full Cytochrome B sequences. Mar Mamm Sci 15(3):619–648. CrossRefGoogle Scholar
  61. Lee PC, Fishlock V, Webber CE, Moss CJ (2016) The reproductive advantages of a long life: longevity and senescence in wild female African elephants. Behav Ecol Sociobiol 70:337–345. CrossRefPubMedPubMedCentralGoogle Scholar
  62. Lesson RP (1828) Cétacés. Complément des oeuvres de Buffon ou Histoire Naturelle des animaux rares découverts par les naturalistes et les voyageurs depuis la mort de Buffon, vol 1. Badouin Freres, ParisGoogle Scholar
  63. Mahaffy SD (2012) Site fidelity, associations and long-term bonds of short-finned pilot whales off the Island of Hawai’i. MSc Thesis, Portland State UniversityGoogle Scholar
  64. 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 Mamm Sci 31(4):1427–1451. CrossRefGoogle Scholar
  65. Marsh H, Kasuya T (1984) Changes in the ovaries of the short-Finned pilot whale, Globicephala macrorhynchus, with age and reproductive activity. Rep Int Whal Comm 6:311–335Google Scholar
  66. Marsh H, Kasuya T (1986) Evidence for reproductive senescence in female cetaceans. Rep Int Whal Comm 8:57–74Google Scholar
  67. Martin AR, Rothery P (1993) Reproductive parameters of female long-finned pilot whales (Globicephala melas) around the Faroe Islands. Rep Int Whal Comm 14:263–304Google Scholar
  68. Mate BR, Lagerquist BA, Winsor M, Geraci JR, Prescott JH (2005) Movements and dive habits of a satellite-monitored long-finned pilot whale (Globicephala melas) in the Northwest Atlantic. Mar Mamm Sci 21:136–144. CrossRefGoogle Scholar
  69. McAuliffe K, Whitehead H (2005) Eusociality, menopause and information in matrilineal whales. Trends Ecol Evol 20(12):650. CrossRefPubMedGoogle Scholar
  70. McComb K, Moss C, Durant SM, Baker L, Sayialel S (2001) Matriarchs as repositories of social knowledge in African elephants. Science 292(5516):491–494. CrossRefPubMedGoogle Scholar
  71. Mendonça AS (2012) Estudo da distribuição e abundância da Baleia pi-loto-tropical (Globicephala macrorhynchus) no Arquipélago dos Açores. PhD Thesis, University of Algarve, PortugalGoogle Scholar
  72. Miralles L, Lens S, Rodríguez-Folgar A, Carrillo M, Martín V, Mikkelsen B, Garcia-Vazquez E (2013) Interspecific introgression in cetaceans: DNA markers reveal post-F1 status of a pilot whale. PLoS One 8(8):e69511. CrossRefPubMedPubMedCentralGoogle Scholar
  73. Miralles L, Oremus M, Silva MA, Planes S, Garcia-vazquez E (2016) Interspecific hybridization in pilot whales and asymmetric genetic introgression in northern Globicephala melas under the scenario of global warming. PLoS One 11(8):1–15. CrossRefGoogle Scholar
  74. Möller LM, Wiszniewski J, Parra G, Beheregaray L (2012) Sociogenetic structure, kin associations and bonding in delphinids. Mol Ecol 21(3):745–764. CrossRefGoogle Scholar
  75. Newman MEJ, Girvan M (2014) Finding and evaluating community structure in networks. Phys Rev E 69(2):26113. CrossRefGoogle Scholar
  76. Nores C, Peréz C (1988) Overlapping range between Globicephala macrorhynchus and Globicephala melaena in the northeastern Atlantic. Mammalia 52(1):51–55CrossRefGoogle Scholar
  77. Nyakaana S, Arctander P (1999) Population genetic structure of the African elephant in Uganda based on variation at mitochondrial and nuclear loci: evidence for male-biased gene flow. Mol Ecol 8(7):1105–1115. CrossRefPubMedGoogle Scholar
  78. Olson PA (2018) Pilot whales: Globicephala melas and G. macrorhynchus. In: Würsig B, Thewissen JGM, Kovacs KM (eds) Encyclopedia of marine mammals, 3rd edn. Academic, New York, pp 701–705. CrossRefGoogle Scholar
  79. Oremus M (2008) Genetic and demographic investigation of population structure and social system in four delphinid species. PhD Thesis, University of AuklandGoogle Scholar
  80. Oremus M, Gales R, Dalebout ML, Funahashi N, Endo T, Kage T et al (2009) Worldwide mitochondrial DNA diversity and phylogeography of pilot whales (Globicephala spp.). Biol J Linn Soc 98(4):729–744. CrossRefGoogle Scholar
  81. Ottensmeyer CA, Whitehead H (2003) Behavioural evidence for social units in long-finned pilot whales. Can J Zool 81(8):1327–1338. CrossRefGoogle Scholar
  82. Packer C, Tatar M, Collins A (1998) Reproductive cessation in mammals. Nature 392(1989):807–811CrossRefGoogle Scholar
  83. Pavelka MSM, Brent LJN, Croft DP, Fedigan LM (2018) Post-fertile lifespan in female primates and cetaceans. In: Kalbitzer U, Jack KM (eds) Primate life histories, sex roles, and adaptability. Springer, Basel, pp 37–55. CrossRefGoogle Scholar
  84. 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(1):20–31. CrossRefPubMedGoogle Scholar
  85. Prieto R, Fernandes M (2007) Revision of the occurrence of the long-finned pilot whale Globicephala melas (Traill, 1809), in the Azores. Arquipélago 24(1861):65–69.
  86. Pusey A, Wolf M (1996) Inbreeding avoidance in animals. Trends Ecol Evol 11(5):201–206. CrossRefPubMedGoogle Scholar
  87. Ralls K, Brugger K, Glick A (1980) Deleterious effects of inbreeding in a herd of captive Dorcas gazelle Gazella dorcas. Int Zoo Yearb 20(1):137–146. CrossRefGoogle Scholar
  88. Richard KR, Dillon MC, Whitehead H, Wright JM (1996) Patterns of kinship in groups of free-living sperm whales (Physeter macrocephalus) revealed by multiple genetic analyses. Proc Natl Acad Sci 93:8792–8795CrossRefGoogle Scholar
  89. Sabatier E, Pante E, Dussud C, Van Canneyt O, Simon-Bouhet B, Viricel AA (2015) Genetic monitoring of pilot whales, Globicephala spp. (Cetacea: Delphinidae), stranded on French coasts. Mammalia 79(1):111–114. CrossRefGoogle Scholar
  90. Santos MB, Monteiro SS, Vingada JV, Ferreira M, Lopez A, Martínez Cedeira JA et al (2014) Patterns and trends in the diet of long-finned pilot whales (Globicephala melas) in the northeast Atlantic. Mar Mamm Sci 30(1):1–19. CrossRefGoogle Scholar
  91. Sergeant DE (1962) The biology of the pilot or pothead whale Globicephala melaena (Traill) in Newfoundland waters. Bull Fish Res Bd Can 132:1–84Google Scholar
  92. Servidio A (2014) Distribution, social structure and habitat use of short-finned pilot whale, Globicephala macrorhynchus, in the Canary Islands. PhD Thesis, University of St AndrewsGoogle Scholar
  93. Servidio A, Alves F, Dinis A, Freitas L, Martín V (2007) First record of movement of short-finned pilot whales between two Atlantic oceanic Archipelagos. In: The 17th biennial conference on the biology of marine mammals. The Society for Marine Mammalogy, Cape TownGoogle Scholar
  94. Siemann LA (1994) Mitochondrial DNA sequence variation in North Atlantic long-finned pilot whales, Globicephala melas. PhD Thesis, Massachusetts Institute of TechnologyGoogle Scholar
  95. Sillero-Zubiri C, Gottelli D, MacDonald DW (1996) Male philopatry, extra-pack copulations and inbreeding avoidance in Ethiopian wolves (Canis simensis). Behav Ecol Sociobiol 38(5):331–340. CrossRefGoogle Scholar
  96. Silva MA, Prieto R, Magalhães S, Cabecinhas R, Cruz A, Gonçalves JM, Santos RS (2003) Occurrence and distribution of cetaceans in the waters around the Azores (Portugal), Summer and Autumn 1999–2000. Aquat Mamm 29(1):77–83. CrossRefGoogle Scholar
  97. Silva MA, Prieto R, Cascão I, Seabra MI, Machete M, Baumgartner MF, Santos RS (2014) Spatial and temporal distribution of cetaceans in the mid-Atlantic waters around the Azores. Mar Biol Res 10(2):123–137. CrossRefGoogle Scholar
  98. Sousa A, Alves F, Dinis A, Bentz J, Cruz MJ, Nunes JP (2018) How vulnerable are cetaceans to climate change? Developing and testing a new index. Ecol Indic 98:9–18. CrossRefGoogle Scholar
  99. Traill TS (1809) Description of a new species of whale, Delphinus melas. In a letter from Thomas Stewart Traill, M. D. to Mr. Nicholson. J Nat Philos Chem Arts 1809:81–83Google Scholar
  100. van Bree PJH, Best BB, Ross GJB (1978) Occurrence of the two species of pilot whales (genus Globicephala) on the coast of South Africa. Mammalia 42(1872):323–328Google Scholar
  101. van Cise AM, Martien KK, Mahaffy SD, Baird RW, Webster DL, Fowler JH et al (2017) Familial social structure and socially-driven genetic differentiation in Hawaiian short-finned pilot whales. Mol Ecol 26:6730–6741. CrossRefPubMedGoogle Scholar
  102. van Waerebeek K, Leaper R, Baker AN, Papastavrou V, Thiele D, Findlay K et al (2010) Odontocetes of the Southern Ocean Sanctuary. J Cetacean Res Manag 11(3):315–346Google Scholar
  103. Verborgh P, De Stephanis R, Pérez S, Jaget Y, Barbraud C, Guinet C (2009) Survival rate, abundance, and residency of long-finned pilot whales in the Strait of Gibraltar. Mar Mamm Sci 25(3):523–536. CrossRefGoogle Scholar
  104. Verborgh P, Gauffier P, Esteban R, Giménez J, Cañadas A, Salazar-Sierra JM, de Stephanis R (2016) Conservation status of long-finned pilot whales, Globicephala melas, in the Mediterranean Sea. In: Notarbartolo Di Sciara G, Podestà M, Curry BE (eds) Advances in marine biology, vol 75. Academic, Oxford, pp 173–203. CrossRefGoogle Scholar
  105. Wells RS, Fougeres EM, Cooper AG, Stevens RO, Brodsky M, Lingenfelser R et al (2013) Movements and dive patterns of short-finned pilot whales (Globicephala macrorhynchus) released from a mass stranding in the Florida Keys. Aquat Mamm 39(1):61–72. CrossRefGoogle Scholar
  106. Werner TB, Northridge S, McClellan K, Young N (2018) Mitigating bycatch and depredation of marine mammals in longline fisheries. ICES J Mar Sci 72(5):1576–1586CrossRefGoogle Scholar
  107. West Eberhard MJ (1975) The evolution of social behavior by kin selection. Q Rev Biol 50(1):1–33Google Scholar
  108. Whitehead H (1990) Rules for roving males. J Theor Biol 145:355–368CrossRefGoogle Scholar
  109. Whitehead H (1998) Cultural selection and genetic diversity in matrilineal whales. Science 282(5394):1708–1711CrossRefGoogle Scholar
  110. Whitehead H (2007) Selection of models of lagged identification rates and lagged association rates using AIC and QAIC. Commun Stat Simul Comput 36(6):1233–1246. CrossRefGoogle Scholar
  111. Whitehead H (2008) Analyzing animal societies: quantitative methods for vertebrate social analysis. University of Chicago Press, ChicagoCrossRefGoogle Scholar
  112. Whitehead H, Rendell L (2014) The cultural lives of whales and dolphins. University of Chicago Press, ChicagoCrossRefGoogle Scholar
  113. Whitehead H, Antunes R, Gero S, Wong SNP, Engelhaupt D, Rendell L (2012) Multilevel societies of female sperm whales (Physeter macrocephalus) in the Atlantic and Pacific: why are they so different? Int J Primatol 33(5):1142–1164. CrossRefGoogle Scholar
  114. Williams GC (1957) Pleiotropy, natural selection, and the evolution of senescence. Evolution 11(4):398–411. CrossRefGoogle Scholar
  115. Wittemyer G, Okello JBA, Rasmussen HB, Arctander P, Nyakaana S, Douglas-Hamilton I, Siegismund HR (2009) Where sociality and relatedness diverge: the genetic basis for hierarchical social organization in African elephants. Proc R Soc B Biol Sci 276(1672):3513–3521. CrossRefGoogle Scholar
  116. Wright BM, Stredulinsky EH, Ellis GM, Ford JKB (2016) Kin-directed food sharing promotes lifetime natal philopatry of both sexes in a population of fish-eating killer whales, Orcinus orca. Anim Behav 115:81–95. CrossRefGoogle Scholar

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© Springer Nature Switzerland AG 2019

Authors and Affiliations

  1. 1.University of ManchesterManchesterUK
  2. 2.Hatfield Marine Science CenterOregon State UniversityNewportUSA

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