Night-life of Bryde’s whales: ecological implications of resting in a baleen whale

  • Sahar Izadi
  • Mark Johnson
  • Natacha Aguilar de Soto
  • Rochelle Constantine
Original Article


Many animals require intervals of rest or sleep in which their vigilance level is reduced. For marine fauna, including large baleen whales, resting potentially increases the risk of predation and vessel-strike. However, there is scarce information about how, and how often, whales rest which makes it difficult to assess the severity of this risk. Here we examine resting patterns of Bryde’s whales (Baleaenoptera edeni/brydei), using data collected by sound and movement archival tags (DTAGs) deployed on four whales in the Hauraki Gulf, New Zealand. To identify low activity levels associated with resting, we used RMS jerk and mean flow noise (as proxies for activity and speed, respectively), as well as changes in dive patterns (dive depth and shape), fluking, and respiration rates. The tagged whales showed strong diel differences in behavior with long periods of low activity consistent with resting occurring exclusively during the night. This pattern indicates that either (i) Bryde’s whales rely on senses that are less effective in the dark to locate prey, or (ii) that prey aggregate less densely at night, making foraging less efficient. Thus, Bryde’s whales conserve energy through rest during times when the net benefit of foraging effort is low. However, by reducing their interaction level with their environment, night-time resting also makes Bryde’s whales more vulnerable to vessel strikes, an important source of mortality for cetaceans.

Significant statement

All mammals need to rest periodically and whales are no exception. But while resting land mammals can be observed directly, little is known about when and how whales rest; even though lower vigilance levels during resting could make them more vulnerable to threats such as collisions with boat traffic. We used sound and movement logging tags on resident Bryde’s whales in a busy gulf to study their daily activity patterns. We found that, while whales were active during daytime making energetic lunges to capture tonnes of plankton, they dedicated much of the night to rest. This suggests that whales may rely on vision to find prey or that prey are less densely aggregated at night making foraging less efficient. However, this near-surface resting behavior which may also be shared by the other giant baleen whales increases the risk of ship strikes.


Rest Sleep Whales Diel behavior Accelerometry Biologging 



We thank the Marine Alliance for Science and Technology Scotland, the field assistants from the Marine Mammal Ecology Group, RV Hawere and Dolphin Explorer crews, Phil Brown, Vivian Ward, Tim Higham, Kevin Chang, and tāngata whenua of the Gulf. We also acknowledge the anonymous reviewers for careful and thorough review of our manuscript and for their useful comments.


This work was supported by the Auckland Council, the Department of Conservation, the University of Auckland including a Doctoral Scholarship to SI, and a Marie Curie Post-Doctoral Fellowship to NAS.

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

Ethical approval

The research was conducted in accordance with permits from the University of Auckland Animal Ethics Committee (Permit #AEC03/2008/R636 and 910) and the New Zealand Department of Conservation (Permits #PER02/2009/01) issued to RC.


  1. Acevedo-Gutierrez A, Caroll DA, Tershy BR (2002) High feeding costs limit dive time in the largest whales. J Exp Biol 205:1747–1753PubMedGoogle Scholar
  2. Aguilar Soto N, Johnson MP, Madsen PT, Diaz F, Dominguez 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 77:936–947CrossRefPubMedGoogle Scholar
  3. Alves F, Dinis A, Cascão I, Freitas L (2010) Bryde’s whale (Balaenoptera brydei) stable associations and dive profiles: new insights into foraging behavior. Mar Mammal Sci 26:202–212CrossRefGoogle Scholar
  4. Amlaner CJ, Ball NJ (1983) A synthesis of sleep in wild birds. Behaviour 87:85–119CrossRefGoogle Scholar
  5. Andersen V, Nival P (1991) A model of the diel vertical migration of zooplankton based on euphausiids. J Mar Res 49:153–175CrossRefGoogle Scholar
  6. Anderson JR (2000) Sleep-related behavioural adaptations in free-ranging anthropoid primates. Sleep Med Rev 4:355–373CrossRefPubMedGoogle Scholar
  7. Baker AN, Madon B (2007) Bryde’s whales (Balaenoptera cf. brydei Olsen 1913) in the Hauraki Gulf and northeastern New Zealand waters, Science for Conservation, vol 272. Department of Conservation, WellingtonGoogle Scholar
  8. Baker CS, Chilvers BL, Constantine R, DuFresne S, Mattlin RH, van Helden A, Hitchmough R (2010) Conservation status of New Zealand marine mammals (suborders Cetacea and Pinnipedia). New Zeal J Mar Freshw Res 44:1–15CrossRefGoogle Scholar
  9. Bates D, Maechler M, Bolker B, Walker S (2015) Fitting linear mixed-effects models using lme4. J Stat Softw 67:1–48CrossRefGoogle Scholar
  10. Benoit-Bird KJ, Kuletz K, Heppell S, Jones N, Hoover B (2011) Active acoustic examination of the diving behavior of murres foraging on patchy prey. Mar Ecol-Prog Ser 443:217–235CrossRefGoogle Scholar
  11. Best PB (2001) Distribution and population separation of Bryde’s whale Balaenoptera edeni off southern Africa. Mar Ecol-Prog Ser 220:277–289CrossRefGoogle Scholar
  12. Blix S, Folkow L (1995) Daily energy expenditure in free living minke whales. Acta Physiol Scand 153:61–66CrossRefPubMedGoogle Scholar
  13. Booth WE, Søndergaard M (1989) Picophytoplankton in the Hauraki gulf, New Zealand. New Zeal J Mar Freshw Res 23:69–78CrossRefGoogle Scholar
  14. Calambokidis J, Schorr GS, Steiger GH, Francis J, Bakhtiari M, Marshall G, Oleson EM, Gendron D, Robertson K (2007) Insights into the underwater diving, feeding, and calling behavior of blue whales from a suction-cup-attached video-imaging tag (Crittercam). Mar Technol Soc J 41:19–29CrossRefGoogle Scholar
  15. Campbell SS, Tobler I (1984) Animal sleep: a review of sleep duration across phylogeny. Neurosci Biobehav Rev 8:269–300CrossRefPubMedGoogle Scholar
  16. Christiansen F, Lynas NM, Lusseau D, Tscherter U (2015) Structure and dynamics of minke whale surfacing patterns in the Gulf of St. Lawrence, Canada. PLoS One 10:e0126396CrossRefPubMedPubMedCentralGoogle Scholar
  17. Constantine R, Brunton DH, Dennis T (2004) Dolphin-watching tour boats change bottlenose dolphin (Tursiops truncatus) behaviour. Biol Conserv 117:299–307CrossRefGoogle Scholar
  18. Constantine R, Johnson M, Riekkola L, Jervis S, Kozmian-Ledward L, Dennis T, Torres LG, Aguilar de Soto N (2015) Mitigation of vessel-strike mortality of endangered Bryde’s whales in the Hauraki Gulf, New Zealand. Biol Conserv 186:149–157CrossRefGoogle Scholar
  19. Croll DA, Tershy BR, Newton KM (2009) Filter feeding. In: Perrin WF, Würsig B, Thewissen JGM (eds) Encyclopedia of marine mammals, 2nd edn. Thewissen, Academic Press, London, pp 421–425Google Scholar
  20. De Robertis A (2000) Size-dependent visual predation risk and the timing of vertical migration: an optimization model. Limnol Oceanogr 47:925–933CrossRefGoogle Scholar
  21. Dolphin WF (1987) Ventilation and dive patterns of humpback whales, Megaptera novaeangliae, on their Alaskan feeding grounds. Can J Zool 65:83–90CrossRefGoogle Scholar
  22. Fiedler PC, Reilly SB, Hewitt RP, Demer D, Philbrick VA, Smith S, Armstrong W, Croll DA, Tershy BR, Mate BR (1998) Blue whale habitat and prey in the Channel Islands. Deep-Sea Res Pt II 45:1781–1801CrossRefGoogle Scholar
  23. Ford J (2009) Killer whale. In: Perrin WF, Würsig B, Thewissen JGM (eds) Encyclopedia of marine mammals, 2nd edn. Thewissen, Academic Press, London, pp 650–657CrossRefGoogle Scholar
  24. Fox J, Weisberg S (2011) An {R} companion to applied regression, 2nd edn. Sage, Thousand OaksGoogle Scholar
  25. Friedlaender AS, Tyson RB, Stimpert AK, Read AJ, Nowacek DP (2013) Extreme diel variation in the feeding behavior of humpback whales along the western Antarctic Peninsula during autumn. Mar Ecol-Prog Ser 494:281–289CrossRefGoogle Scholar
  26. Gauthier-Clerc M, Tamisier A, Cezilly F (1998) Sleep-vigilance trade-off in green-winged teals (Anas crecca crecca). Can J Zool 76:2214–2218CrossRefGoogle Scholar
  27. Goldbogen JA, Calambokidis J, Shadwick RE, Oleson EM, McDonald MA, Hildebrand JA (2006) Kinematics of foraging dives and lunge-feeding in fin whales. J Exp Biol 209:1231–1244CrossRefPubMedGoogle Scholar
  28. Goldbogen JA, Calambokidis J, Croll DA, Harvey JT, Newton KM, Oleson EM, Schorr G, Shadwick RE (2008) Foraging behavior of humpback whales: kinematic and respiratory patterns suggest a high cost for a lunge. J Exp Biol 211:3712–3719CrossRefPubMedGoogle Scholar
  29. Goldbogen JA, Calambokidis J, Oleson E, Potvin J, Pyenson ND, Schorr G, Shadwick RE (2011) Mechanics, hydrodynamics and energetics of blue whale lunge feeding: efficiency dependence on krill density. J Exp Biol 214:131–146CrossRefPubMedGoogle Scholar
  30. Goldbogen JA, Friedlaender AS, Calambokidis J, McKenna MF, Simon M (2013) Integrative approaches to the study of baleen whale diving behavior, feeding performance, and foraging ecology. Bioscience 63:90–100CrossRefGoogle Scholar
  31. Goley PD (1999) Behavioral aspects of sleep in Pacific white-sided dolphins. Mar Mammal Sci 15:1054–1064CrossRefGoogle Scholar
  32. Griebel U, Peichl L (2003) Colour vision in aquatic mammals—facts and open questions. Aquat Mamm 29:18–30CrossRefGoogle Scholar
  33. Hart NS (2001) Variations in cone photoreceptor abundance and the visual ecology of birds. J Comp Physiol 187:685–697CrossRefGoogle Scholar
  34. Jarman SN, Wiseman N, Baker CS, Gales NJ (2006) Incidence of prey DNA types in Bryde’s whale scats. SC/58/E29. Scientific Committee of the International Whaling Commission, St Kitts and NevisGoogle Scholar
  35. Johnson MP, Tyack PL (2003) A digital acoustic recording tag for measuring the response of wild marine mammals to sound. IEEE J Ocean Eng 28:3–12CrossRefGoogle Scholar
  36. Johnson M, Madsen PT, Zimmer WMX, Aguilar de Soto NA, Tyack PL (2004) Beaked whales echolocate on prey. Proc R Soc Lond B 271:S383–S386CrossRefGoogle Scholar
  37. Johnson M, Aguilar de Soto N, Madsen P (2009) Studying the behaviour and sensory ecology of marine mammals using acoustic recording tags: a review. Mar Ecol-Prog Ser 395:55–73CrossRefGoogle Scholar
  38. Johnson M, Partan J, Hurst T (2013) Low complexity lossless compression of underwater sound recordings. J Acoust Soc Am 133:1387–1398CrossRefPubMedGoogle Scholar
  39. Kato H, Perrin WF (2009) Bryde’s whales (Balaenoptera edeni/brydei). In: Perrin WF, Würsig B, Thewissen JGM (eds) Encyclopedia of marine mammals, 2nd edn. Thewissen, Academic Press, London, pp 158–163CrossRefGoogle Scholar
  40. Ketten DR (2000) Cetacean ears. In: WWL A, Popper AN, Fay RR (eds) Hearing by whales and dolphins. Springer, New York, pp 43–108CrossRefGoogle Scholar
  41. Kronfeld-Schor N, Dayan T (2003) Partitioning of time as an ecological resource. Annu Rev Ecol Evol Sci 34:153–181CrossRefGoogle Scholar
  42. Lampert W (1989) The adaptive significance of diel vertical migration of zooplankton. Funct Ecol 3:21–27CrossRefGoogle Scholar
  43. Lesku JA, Roth TC II, Rattenborg NC, Amlaner CJ, Lima SL (2009) History and future of comparative analyses in sleep research. Neurosci Biobehav Rev 33:1024–1036CrossRefPubMedGoogle Scholar
  44. Lima SL, Rattenborg NC, Lesku J, Amlaner CJ (2005) Sleeping under the risk of predation. Anim Behav 70:723–736CrossRefGoogle Scholar
  45. Lyamin OI, Mukhametov LM, Siegel JM, Manger PR, Shpak OV (2001) Resting behavior in a rehabilitating gray whale calf. Aquat Mamm 27:256–266Google Scholar
  46. Lyamin OI, Mukhametov LM, Siegel JM, Nazarenko EA, Pollyakova LG, Shpak OV (2002) Unihemispheric slow wave sleep and the state of eyes in a white whale. Behav Brain Res 129:125–129CrossRefPubMedGoogle Scholar
  47. Lyamin OI, Manger PR, Ridgway SH, Mukhametov LM, Siegel JM (2008) Cetacean sleep: an unusual form of mammalian sleep. Neurosci Biobehav Rev 32:1451–1484CrossRefPubMedGoogle Scholar
  48. Martin Lopez ML (2016) Fine-scale kinematics and energetics: a detailed insight into the biomechanical strategies of cetacean locomotion. PhD thesis, The University of St Andrews, Scotland, UKGoogle Scholar
  49. Miller PJO, Aoki K, Rendell LE, Amano M (2008) Stereotypical resting behaviour of the sperm whale. Curr Biol 18:21–23CrossRefGoogle Scholar
  50. Mobley JRJ, Helweg DA (1990) Visual ecology and cognition in cetaceans. In: Thomas JA, Kastelein RA (eds) Sensory abilities of cetaceans, laboratory and field evidence. Springer, New York, pp 1689–1999Google Scholar
  51. Mukhametov LM (1987) Unihemispheric slow-wave sleep in the Amazonian dolphin, Inia geoffrensis. Neurosci Lett 79:128–132CrossRefPubMedGoogle Scholar
  52. Mukhametov LM, Supin AY, Polyakova IG (1977) Interhemispheric asymmetry of the electroencephalographic sleep patterns in dolphins. Brain Res 134:581–584CrossRefPubMedGoogle Scholar
  53. Murase H, Tamura T, Kiwada H, Fujise Y, Watanabe H, Ohizumi H, Yonezaki S, Okamura H, Kawahara S (2007) Prey selection of common minke (Balaenoptera acutorostrata) and Bryde’s (Balaenoptera edeni) whales in the western North Pacific in 2000 and 2001. Fish Oceanogr 16:186–201CrossRefGoogle Scholar
  54. Norris KS, Dohl TP (1980) Behavior of the Hawaiian spinner dolphin, Stenella longirostris. Fish Bull 77:821–849Google Scholar
  55. Nowacek DP, Johnson MP, Tyack PL (2004) North Atlantic right whales (Eubalaena glacialis) ignore ships but respond to alerting stimuli. Proc R Soc Lond B 271:227–231CrossRefGoogle Scholar
  56. Nummela S (2009) Hearing. In: Perrin WF, Würsig B, Thewissen JGM (eds) Encyclopedia of marine mammals, 2nd edn. Thewissen, Academic Press, London, pp 553–562CrossRefGoogle Scholar
  57. Ohman MD, Frost BW, Cohen EB (1983) Reverse diel vertical migration: an escape from invertebrate predators. Science 220:1404–1407CrossRefPubMedGoogle Scholar
  58. Oleson EM, Calambokidis J, Burgess WC, McDonalds MA, LeDuc CA, Hildebrand JA (2007) Behavioral context of call production by eastern North Pacific blue whales. Mar Ecol-Prog Ser 330:269–284CrossRefGoogle Scholar
  59. Panigada S, Pesante G, Zanardelli M, Oehen S (2003) Day and night-time behaviour of fin whales in the Western Ligurian Sea. In: Proceedings of the oceans 2003, San Diego, CA, pp 466–471Google Scholar
  60. Parks SE, Searby A, Célérier A, Johnson MP, Nowacek DP, Tyack PL (2011) Sound production behavior of individual North Atlantic right whales: implications for passive acoustic monitoring. Endanger Species Res 15:63–76CrossRefGoogle Scholar
  61. Paul LJ (1968) Some seasonal water temperature patterns in the Hauraki Gulf, New Zealand. New Zeal J Mar Freshw Res 2:535–258CrossRefGoogle Scholar
  62. Peichl L, Behrmann G, Kröger RHH (2001) For whales and seals the ocean is not blue: a visual pigment loss in marine mammals. Eur J Neurosci 13:1520–1528CrossRefPubMedGoogle Scholar
  63. Penry G, Cockcroft V, Hammond P (2011) Seasonal fluctuations in occurrence of inshore Bryde’s whales in Plettenberg Bay, South Africa, with notes on feeding and multispecies associations. Afr J Mar Sci 33:403–414CrossRefGoogle Scholar
  64. Pivorunas A (1979) The feeding mechanisms of baleen whales. Am Sci 67:432–440Google Scholar
  65. R Core Team (2018) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL
  66. Rattenborg NC, Amlaner CJ, Lima SL (2000) Behavioral, neurophysiological and evolutionary perspectives on unihemispheric sleep. Neurosci Biobehav Rev 24:817–842CrossRefPubMedGoogle Scholar
  67. Rechtschaffen A (1998) Current perspectives on the function of sleep. Perspect Biol Med 41:359–390CrossRefPubMedGoogle Scholar
  68. Roth TC II, Lesku JA, Amlaner CJ, Lima S (2006) A phylogenetic analysis of the correlates of sleep in birds. J Sleep Res 15:395–402CrossRefPubMedGoogle Scholar
  69. Rutz C, Hays GC (2009) New frontiers in biologging science. Biol Lett 5:289–292CrossRefPubMedPubMedCentralGoogle Scholar
  70. Sekiguchi Y, Arai K, Kohshima S (2006) Sleep behaviour: sleep in continuously active dolphins. Nature 441:E9–E10CrossRefPubMedGoogle Scholar
  71. Serafetinides EA, Shurley JT, Brooks RE (1972) Electroencephalogram of the pilot whale, Globicephala scammoni, in wakefulness and sleep: lateralization aspects. Int J Psychobiol 2:129–135Google Scholar
  72. Shpak OV, Liamin OI, Manger PR, Siegel JM, Mukhametov LM (2009) Rest and activity states in the Commerson’s dolphin (Cephalorhynchus commersonii). J Evol Biochem Phys 45:97–104CrossRefGoogle Scholar
  73. Siegel JM (2005) Clues to the functions of mammalian sleep. Nature 437:1264–1271CrossRefPubMedGoogle Scholar
  74. Siegel JM (2008) Do all animals sleep? Trends Neurosci 31:208–213CrossRefPubMedGoogle Scholar
  75. Siegel JM (2011) Sleep in animals: a state of adaptive inactivity. In: Kryger MH, Roth T, Dement WC (eds) Principles and practice of sleep medicine, 5th edn. WB Saunders, Philadelphia, pp 126–138CrossRefGoogle Scholar
  76. da Silva J, Terhune JM (1998) Harbour seal grouping as an anti-predator strategy. Anim Behav 36:309–316Google Scholar
  77. Simon M, Johnson M, Tyack P, Madsen PT (2009) Behaviour and kinematics of continuous ram filtration in bowhead whales (Balaena mysticetus). Proc R Soc Lond B 276:3819–3828CrossRefGoogle Scholar
  78. Simon M, Johnson M, Madsen P (2012) Keeping momentum with a mouthful of water: behavior and kinematics of humpback whale lunge feeding. J Exp Biol 215:3786–3798CrossRefPubMedGoogle Scholar
  79. Szczucka J (2009) Acoustic studies of diving birds in the Arctic. In: 3rd International Conference on Underwater Acoustic Measurements: Technologies and Results, Nafplion, Greece, pp 1181–1188Google Scholar
  80. Tershy BR (1992) Body size, diet, habitat use, and social behavior of Balaenoptera whales in the Gulf of California. Am Soc Mammal 73:477–486CrossRefGoogle Scholar
  81. Tyson RB, Friedlaender AS, Ware C, Stimpert AK, Nowacek DP (2012) Synchronous mother and calf foraging behaviour in humpback whales Megaptera novaeangliae: insights from multi-sensor suction cup tags. Mar Ecol-Prog Ser 457:209–220CrossRefGoogle Scholar
  82. Watkins WA, Daher MA, DiMarzio NA (1999) Sperm whale surface activity from tracking by radio and satellite tags. Mar Mammal Sci 15:1158–1180CrossRefGoogle Scholar
  83. Wiseman N (2008) Genetic identity and ecology of Bryde’s whales in the Hauraki Gulf, New Zealand. PhD thesis, The University of Auckland, New ZealandGoogle Scholar
  84. Würsig B, Würsig M (1980) Behavior and ecology of the dusky dolphin, Lagenorhynchus obscurus, in the South Atlantic. Fish Bull 77:871–890Google Scholar
  85. Ydesen KS, Wisniewska DM, Hansen JD, Beedholm K, Johnson M, Madsen PT (2014) What a jerk: prey engulfment revealed by high-rate, super-cranial accelerometry on a harbour seal (Phoca vitulina). J Exp Biol 217:2239–2243CrossRefPubMedGoogle Scholar
  86. Zerbini AN, Secchi ER, Siciliano S, Simoes-Lopes PC (1997) A review of the occurrence and distribution of whales of the genus Balaenoptera along the Brazilian Coast. SC/48/SH4. Scientific Committee of the International Whaling Commission, Monte Carlo, MonacoGoogle Scholar

Copyright information

© Springer-Verlag GmbH Germany, part of Springer Nature 2018

Authors and Affiliations

  1. 1.Institute of Marine ScienceUniversity of AucklandAucklandNew Zealand
  2. 2.Sea Mammal Research UnitUniversity of St AndrewsFifeUK
  3. 3.Department of BioscienceAarhus UniversityAarhusDenmark
  4. 4.BIOECOMACLa Laguna UniversityTenerifeSpain
  5. 5.School of Biological SciencesUniversity of AucklandAucklandNew Zealand

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