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Physiology of behaviour in pinnipeds

  • Chapter
The Behaviour of Pinnipeds

Abstract

Behaviour is an evolutionarily labile species trait that responds to environmental opportunity within the constraints imposed by evolutionarily more conservative traits such as anatomy and physiology. Because of their different degrees of adaptation to a marine environment, pinnipeds provide an interesting group in which to look at these relationships between physiological constraints and environmental opportunities. Although behaviour is shaped by the interaction between environment and physiology, it in tum influences physiology and restricts the environments to which the animals are exposed. This chapter looks at the interplay between anatomy, physiology, environment and behaviour in producing the social and individual behaviours observed in a broad range of pinnipeds. Because pinniped behaviour and environmental parameters are more easily observed than physiological mechanisms are delineated, the contribution of physiology must often be inferred by comparison with animals from other orders.

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References

  • Anderson, S. S. (1978) Day and night activity of grey seal bulls. Mammal Rev. 8: 43–46.

    Article  Google Scholar 

  • Anderson, S. S. and Fedak, M. A. (1985) Grey seal males: Energetic and behavioural links between size and sexual success. Anim. Behav. 33: 829–838.

    Article  Google Scholar 

  • Anderson, S. S. and Harwood, J. (1985) Time budgets and topography: How energy reserves and terrain determine the breeding behaviour of grey seals. Anim. Behav. 33: 1343–1348.

    Article  Google Scholar 

  • Anderson, S. S., Burton, R. W. and Summers, C. F. (1975) Behaviour of grey seals (Halichoerus grypus) during a breeding season at North Rona. J. Zool., London 177: 179–195.

    Article  Google Scholar 

  • Anderson, S. S., Baker, J. R., Prime, J. H. and Baird, A. (1979) Mortality in grey seal pups: Incidence and causes. J. Zool. London 189: 407–417.

    Article  Google Scholar 

  • Arnold, A. P. (1975) The effects of castration and androgen replacement on song, courtship, and aggression in zebra finches (Poephila guttata). J. of Exp. Zool. 191: 309–326.

    Article  CAS  Google Scholar 

  • Arnold, W. and Trillmich, F. (1985) Time budget in Galapagos fur seal pups: The influence of the mother’s presence and absence on pup activity and play. Behaviour 92: 302–321.

    Google Scholar 

  • Ashwell-Erickson, S., Fay, F. H., Elsner, R. and Wartzok, D. (1986) Metabolic and hormonal correlates of molting and regeneration of pelage in Alaskan harbour and spotted seals (Phoca vitulina and Phoca largha). Can. J. of Zool. 64: 1086–1094.

    Article  CAS  Google Scholar 

  • Ashworth, U. S., Ramaiah, G. D. and Keyes, M. C. (1966) Species difference in the composition of milk with special reference to the northern fur seal. J. Dairy Sci. 49: 1206–1211.

    Article  Google Scholar 

  • Bailey, K. M. and Ainley, D. G. (1982) The dynamics of California sea lion predation on Pacific hake. Fish. Res. 1: 163–176.

    Article  Google Scholar 

  • Baker, J. R. (1984) Mortality and morbidity in grey seal pups (Halichoerus grypus): Studies on its causes, effects of environment, the nature and sources of infectious agents and the immunological status of pups. J. Zool., London 203: 23–48.

    Article  Google Scholar 

  • Baker, J. R. and Baker, R. (1988) Effects of environment on grey seal (Halichoerus grypus) pup mortality: Studies on the Isle of May. J. Zool., London 216: 529–537.

    Article  Google Scholar 

  • Baker, J. R. and McCann, T. S. (1989) Pathology and bacteriology of adult male antarctic fur seals, Arctocephalus gazella, dying at Bird Island, South Georgia. Br. Vet J. 145: 263–275.

    Article  PubMed  CAS  Google Scholar 

  • Barber, R. T. and Chavez, F. P. (1983) Biological consequences of El Niño. Science 222: 1203–1210.

    Article  PubMed  CAS  Google Scholar 

  • Bartholomew, G. A. (1953) Behavioural factors affecting social structure in the Alaska fur seal, in Transactions of the Eighteenth North American Wildlife Conference: 48–502.

    Google Scholar 

  • Bartholomew, G. A. (1959) Mother-young relations and the maturation of pup behaviour in the Alaska fur seal. Anim. Behav. 7: 163–171.

    Article  Google Scholar 

  • Bartholomew, G. A. (1966) Interaction of physiology and behaviour under natural conditions, in The Galapagos: 39–45 (R. I. Bowman, ed.), Berkeley, CA: University of California Press.

    Google Scholar 

  • Bartholomew, G. A. (1970) A model for the evolution of pinniped polygyny. Evolution 24: 546–559.

    Article  Google Scholar 

  • Bartholomew, G. A. and Hoel, P. G. (1953) Reproductive behaviour of the Alaska fur seal, Callorhinus ursinus. J. of Mammal. 34: 417–436.

    Article  Google Scholar 

  • Bartholomew, G. A. and Wilke, F. (1956) Body temperature in the northern fur seal, Callorhinus ursinus. J. of Mammal. 37: 327–337.

    Article  Google Scholar 

  • Beddington, J. R., Beverton, R. J. H. and Lavigne, D. M. (eds) (1985) Marine Mammals and Fisheries. London: George Allen & Unwin.

    Google Scholar 

  • Beentjes, M. P. (1989) Haul-out patterns, site fidelity and activity budgets of male Hooker’s sea lions (Phocarctos hookeri) on the New Zealand mainland. Marine Mamm. Sci. 5: 281–297.

    Article  Google Scholar 

  • Beier, J. C. and Wartzok, D. (1979) Mating behaviour of captive spotted seals (Phoca largha). Anim. Behav. 27: 772–781.

    Article  Google Scholar 

  • Bengtson, J. L. and Laws, R. M. (1985) Trends in crabeater seal age at maturity: An insight into antarctic marine interactions, in Antarctic Nutrient Cycles and Food Webs: 669–675 (W. R. Siegfried, P. R. Condy and R. M. Laws, eds), Berlin: Springer-Verlag.

    Google Scholar 

  • Bennett, P. B., Blenkarn, G. D., Roby, J. and Youngblood, D. (1974) Suppression of the high pressure nervous syndrome in human deep dives by He-N2-O2. Undersea Biomed. Res. 1: 221–237.

    PubMed  CAS  Google Scholar 

  • Blix, A. S. and Steen, J. B. (1979) Temperature regulation in newborn polar homeotherms. Physiol. Rev. 59: 285–304.

    PubMed  CAS  Google Scholar 

  • Blix, A. S., Grav, H. J. and Ronald, K. (1975) Brown adipose tissue and the significance of the venous plexuses in pinnipeds. Acta Physiol. Scand. 94:133–135.

    Article  PubMed  CAS  Google Scholar 

  • Blix, A. S., Grav, H. J. and Ronald, K. (1979a) Some aspects of temperature regulation in newborn harp seal pups. Am. J. Physiol. 236: R188–R197.

    PubMed  CAS  Google Scholar 

  • Blix, A. S., Miller, L. K., Keyes, M. C., Grav, H. J. and Eisner, R. (1979b) Newborn northern fur seals (Callorhinus ursinus): Do they suffer from cold?. Am. J. Physiol. 236: R322–R327.

    PubMed  CAS  Google Scholar 

  • Blix, A. S., Fay, F. H., and Ronald, K. (1983) On testicular cooling in phocid seals. Polar Res. 1 n.s.: 231–233.

    Article  Google Scholar 

  • Boness, D. J. (1984) Activity budget of male gray seals, Halichoerus grypus. J. of Mammal. 65: 291–297.

    Article  Google Scholar 

  • Boness, D. J. and James, H. (1979) Reproductive behaviour of the grey seal (Halichoerus grypus) on Sable Island, Nova Scotia. J. Zool., London 188: 477–500.

    Article  Google Scholar 

  • Boness, D. J., Anderson, S. S. and Cox, C. R. (1982) Functions of female aggression during pupping and mating season of grey seals, Halichoerus grypus (Fabricius). Can. J. Zool. 60: 2270–2278.

    Article  Google Scholar 

  • Boness, D. J., Bowen, W. D. and Oftedal, O. T. (1988) Evidence of polygyny from spatial patterns of hooded seals (Cystophora cristata). Can. J. Zool. 66: 703–706.

    Article  Google Scholar 

  • Bonner, W. N. (1972) The grey seal and common seal in European waters. Oceanogr. Marine Biol. Annual Rev. 10: 461–507.

    Google Scholar 

  • Bonner, W. N. (1984) Lactation strategies of pinnipeds: problems for a marine mammalian group. Symp. Zool. Soc. London 51: 253–272.

    Google Scholar 

  • Bowen, W. D., Capstick, C. K. and Sergeant, D. E. (1981) Temporal changes in the reproductive potential of female harp seals (Pagophilus groenlandicus). Can. J. Fish. Aquatic Sci. 38: 495–503.

    Article  Google Scholar 

  • Bowen, W. D., Oftedal, O. T. and Boness, D. J. (1985) Birth to weaning in 4 days: Remarkable growth in the hooded seal, Cystophora cristata. Can. J. Zool. 63: 2841–2846.

    Article  Google Scholar 

  • Boyd, I. L. (1984) The relationship between body condition and the timing of implantation in pregnant grey seals (Halichoerus grypus). J. Zool., London 203: 113–123.

    Article  Google Scholar 

  • Boyd, I. L. and McCann, T. S. (1989) Pre-natal investment in reproduction by female antarctic fur seals. Behav. Ecol. Sociobiol. 24: 377–385.

    Article  Google Scholar 

  • Boyd, J. M. and Campbell, R. N. (1971) The grey seal (Halichoerus grypus) at North Rona, 1959 to 1968. J. Zool., London 164: 469–512.

    Article  Google Scholar 

  • Brownell, R. L. and Ralls, K. (1986) Potential for sperm competition in baleen whales, in Behaviour of Whales in Relation to Management: 97–112 (G. P. Donovan, ed.), Reports of the International Whaling Commission (Special Issue 8). International Whaling Commission, Cambridge.

    Google Scholar 

  • Bryden, M. M. (1968a) Control of growth in two populations of elephant seals. Nature 217: 1106–1108.

    Article  Google Scholar 

  • Bryden, M. M. (1968b) Lactation and suckling in relation to early growth of the southern elephant seal, Mirounga leonina (L.). Aust. J. Zool. 16: 739–748.

    Article  Google Scholar 

  • Bryden, M. M. and Stokes, G. B. (1969) Metabolism of fatty acids in the southern elephant seal Mirounga leonina L. Can. J. Biochem. 47: 757–760.

    PubMed  CAS  Google Scholar 

  • Bryden, M. M., Griffiths, D. J., Kennaway, D. J. and Ledingham, J. (1986) The pineal gland is very large and active in newborn antarctic seals. Experientia (Basel) 42: 564–566.

    Article  CAS  Google Scholar 

  • Burns, J. J. (1981) Bearded seal Erignathus barbatus Erxleben, 1777, in Handbook of Marine Mammals: Vol. 2. Seals: 145–170 (S. H. Ridgway and R. J. Harrison, eds), New York: Academic Press.

    Google Scholar 

  • Bums, J. J., Ray, G. C., Fay, F. H. and Shaughnessy, P. D. (1972) Adoption of a strange pup by the ice-inhabiting harbour seal, Phoca vitulina largha. J. of Mammal. 53: 594–598.

    Article  Google Scholar 

  • Campagna, C. and Le Boeuf, B. J. (1988a) Reproductive behaviour of southern sea lions. Behaviour 104: 233–262.

    Article  Google Scholar 

  • Campagna, C. and Le Boeuf, B. J. (1988b) Thermoregulatory behaviour of southern sea lions and its effect on mating strategies. Behaviour 107: 72–90.

    Article  Google Scholar 

  • Campagna, C., Le Boeuf, B. J. and Cappozzo, H. L. (1988a) Group raids: A mating strategy of male southern sea lions. Behaviour 105: 224–249.

    Article  Google Scholar 

  • Campagna, C., Le Boeuf, B. J. and Cappozzo, H. L. (1988b) Pup abduction and infanticide in southern sea lions. Behaviour 107: 44–60.

    Article  Google Scholar 

  • Cane, M. A. (1983) Oceanographie events during El Niño. Science 222: 1189–1195.

    Article  PubMed  CAS  Google Scholar 

  • Carrick, R., Csordas, S. E. and Ingham, S. E. (1962) Studies on the southern elephant seal, Mirounga leonina (L.). IV. Breeding and development. CSIRO Wildlife Res. 7: 161–197.

    Article  Google Scholar 

  • Castellini, M. A. (1988) Visualizing metabolic transitions in aquatic mammals: Does apnea plus swimming equal ‘diving’? Can. J. Zool. 66: 40–44.

    Article  CAS  Google Scholar 

  • Castellini, M. A., Somero, G. N. and Kooyman, G. L. (1981) Glycolytic enzyme activities in tissues of marine and terrestrial mammals. Physiol. Zool. 54: 242–252.

    CAS  Google Scholar 

  • Castellini, M. A., Davis, R. W. and Kooyman, G. L. (1988) Blood chemistry regulation during repetitive diving in Weddell seals. Physiol. Zool. 61: 379–386.

    Google Scholar 

  • Christenson, T. E. and Le Boeuf, B. J. (1977) Aggression in the female northern elephant seal Mirounga angustirostris. Behaviour 64: 158–172.

    Article  Google Scholar 

  • Clutton-Brock, T. H. and Albon, S. D. (1981) Parental investment in male and female offspring in mammals, in Current Problems in Sociobiology: 223–247 (Eds. Kings College Sociobiology Group) Cambridge University Press, Cambridge.

    Google Scholar 

  • Collins, J. C., Pilkington, T. C. and Schmidt-Nielsen, K. (1971) A model of respiratory heat transfer in a small mammal. Biophys. J. 11: 886–914.

    Article  PubMed  CAS  Google Scholar 

  • Costa, D. P. and Gentry, R. L. (1986) Free-ranging energetics of northern fur seals, in Fur Seals: Maternal Strategies on Land and at Sea: 79–101 (R. L. Gentry and G. L. Kooyman, eds), Princeton, NJ: Princeton University Press.

    Google Scholar 

  • Costa, D. P. and Kooyman, G. L. (1984) Contribution of specific dynamic action to heat balance and thermoregulation in the sea otter Enhydra lutris. Physiol. Zool. 57: 199–203.

    Google Scholar 

  • Costa, D. P. and Trillmich, F. (1988) Mass changes and metabolism during the perinatal fast: A comparison between antarctic (Arctocephalus gazella) and Galapagos fur seals (Arctocephalus galapagoensis). Physiol. Zool. 61: 160–169.

    Google Scholar 

  • Costa, D. P., Le Boeuf, B. J., Huntley, A. C. and Ortiz, C. L. (1986) The energetics of lactation in the northern elephant seal Mirounga angustirostris. J. Zool., London (A) 209: 21–33.

    Article  Google Scholar 

  • Costa, D. P., Trillmich, F. and Croxall, J. P. (1988) Intraspecific allometry of neonatal size in the antarctic fur seal (Arctocephalus gazella). Behav. Ecol. Sociobiol. 22: 361–364.

    Google Scholar 

  • Costa, D. P., Croxall, J. P. and Duck, C. D. (1989) Foraging energetics of antarctic fur seals in relation to changes in prey availability. Ecology 70: 596–606.

    Article  Google Scholar 

  • Cox, C. R. and Le Boeuf, B. J. (1977) Female incitation of male competition: A mechanism in sexual selection. Am. Nat. 111: 317–335.

    Article  Google Scholar 

  • Craig, A. B. and Pasche, A. (1980) Respiratory physiology of freely diving harbour seals (Phoca vitulina). Physiol. Zool. 53: 419–432.

    Google Scholar 

  • Crawley, M. C. and Wilson, G. J. (1976) The natural history and behaviour of the New Zealand fur seal (Arctocephalus forsteri). Tuatara 22: 1–29.

    Google Scholar 

  • Crompton, A. W., Taylor, C. R. and Jagger, J. A. (1978) Evolution of homeothermy in mammals. Nature 272: 333–336.

    Article  PubMed  CAS  Google Scholar 

  • Croxall, J. P. and Gentry, R. L. (eds) (1987) Status, Biology, and Ecology of Fur Seals. NOAA Technical Report NMFS 51. US Department of Commerce, National Marine Fisheries Service, Seattle, WA.

    Google Scholar 

  • Croxall, J. P. and Hiby, L. (1983) Fecundity, survival and site fidelity in Weddell seals, Leptonychotes weddelli. J. Appl. Ecol. 20: 19–32.

    Article  Google Scholar 

  • David, J. H. M. and Rand, R. W. (1986) Attendance behaviour of South African fur seals, in Fur Seals: Maternal Strategies on Land and at Sea: 126–141 (R. L. Gentry and G. L. Kooyman, eds), Princeton, NJ: Princeton University Press.

    Google Scholar 

  • Depocas, F. and Hart, J. S. (1971) Sea water drinking and water flux in starved and fed harbour seals, Phoca vitulina. Can. J. Physiol, and Pharmacol. 49: 53–62.

    Article  CAS  Google Scholar 

  • Deutsch, C. J., Haley, M. P. and Le Boeuf, B. J. (in press) Reproductive effort of male northern elephant seals: Estimates from mass loss. Can. J. Zool.

    Google Scholar 

  • Doidge, D. W. and Croxall, J. P. (1985) Diet and energy budget of the antarctic fur seal, Arctocephalus gazella, at South Georgia, in Antarctic Nutrient Cycles and Food Webs: 543–550 (W. R. Siegfried, P. R. Condy and R. M. Laws, eds), Berlin: Springer-Verlag.

    Google Scholar 

  • Doidge, D. W. and Croxall, J. P. (1989) Factors affecting weaning weight in antarctic fur seals Arctocephalus gazella at South Georgia. Polar Biol. 9: 155–160.

    Article  Google Scholar 

  • Doidge, D. W., Croxall, J. P. and Baker, J. R. (1984) Density-dependent pup mortality in the antarctic fur seal Arctocephalus gazella at South Georgia. J. Zool., London 202: 449–460.

    Article  Google Scholar 

  • Doidge, D. W., McCann, T. S. and Croxall, J. P. (1986) Attendance behaviour of antarctic fur seals, in Fur Seals: Maternal Strategies on Land and at Sea: 102–114 (R. L. Gentry and G. L. Kooyman, eds), Princeton, NJ: Princeton University Press.

    Google Scholar 

  • Dykes, R. W. (1974a) Factors related to the dive reflex in harbour seals: Respiration, immersion bradycardia, and lability of the heart rate. Can. J. Physiol. Pharmacol. 52: 248–258.

    Article  PubMed  CAS  Google Scholar 

  • Dykes, R. W. (1974b) Factors related to the dive reflex in harbour seals: Sensory contributions from the trigeminal region. Can. J. Physiol. Pharmacol. 52: 259–265.

    Article  PubMed  CAS  Google Scholar 

  • Elden, C. A., Keyes, M. C. and Marshall, C. E. (1971) Pineal body of the northern fur seal (Callorhinus ursinus): A model for studying the probable function of the mammalian pineal body. Am. J. Vet. Res. 32: 639–647.

    PubMed  CAS  Google Scholar 

  • Elsner, R. and Gooden, B. (1983) Diving and Asphyxia. Cambridge: Cambridge University Press.

    Book  Google Scholar 

  • Eisner, R., Kooyman, G. L. and Drabek, C. M. (1970) Diving duration in pregnant Weddell seals, in Antarctic Ecology: 477–482 (M. W. Holdgate, ed.), New York: Academic Press.

    Google Scholar 

  • Elsner, R., Hammond, D. D., Denison, D. and Wybum, R. (1977) Temperature regulation in the newborn Weddell seal, in Adaptations within Antarctic Ecosystems: 531–540 (G. A. Llano, ed.), Washington, DC: Smithsonian Institution.

    Google Scholar 

  • Elsner, R., Wartzok, D., Sonafrank, N. B. and Kelly, B. P. (1989) Behavioural and physiological reactions of arctic seals during under-ice pilotage. Can. J. Zool. 67: 2506–2513.

    Article  Google Scholar 

  • Engelhardt, F. R. and Ferguson, J. M. (1980) Adaptive hormone changes in harp seals, Phoca groenlandica, and gray seals, Halichoerus grypus, during the postnatal period. Gen. Comp. Endocrinol. 40: 434–445.

    Article  PubMed  CAS  Google Scholar 

  • Fagen, R. (1981) Animal Play Behaviour. New York: Oxford University Press.

    Google Scholar 

  • Farentinos, R. C. (1971) Some observations on the play behaviour of the Steller sea lion (Eumetopias jubata). Z. Tierpsychol. 28: 428–438.

    Article  PubMed  CAS  Google Scholar 

  • Fay, F. H. and Burns, J. J. (1988) Maximal feeding depth of walruses. Arctic 41: 239–240.

    Google Scholar 

  • Fedak, M. A. and Anderson, S. S. (1982) The energetics of lactation: Accurate measurements from a large wild mammal, the grey seal (Halichoerus grypus). J. Zool., London 198: 473–479.

    Google Scholar 

  • Feldkamp, S. D., DeLong, R. L. and Antonelis, G. A. (1989) Diving patterns of California sea lions, Zalophus californianus. Can. J. Zool. 67: 872–883.

    Article  Google Scholar 

  • Feltz, E. T. and Fay, F. H. (1966) Thermal requirements in vitro of epidermal cells from seals. Cryobiology 3: 261–264.

    Article  PubMed  CAS  Google Scholar 

  • Fisher, R. A. (1930) The Genetical Theory of Natural Selection. Oxford: Oxford University Press.

    Google Scholar 

  • Folkow, L. P. and Blix, A. S. (1987) Nasal heat and water exchange in gray seals. Am. J. Physiol. 253: R883–R889.

    PubMed  CAS  Google Scholar 

  • Frisch, R. E. (1980) Pubertal adipose tissue: Is it necessary for normal sexual maturation? Evidence from the rat and human female. Fed. Proc. 39: 2395–2400.

    PubMed  CAS  Google Scholar 

  • Frost, K. J. and Lowry, L. F. (1981) Ringed, Baikal and Caspian seals, in Handbook of Marine Mammals: Vol. 2. Seals: 29–53 (S. H. Ridgway and R. J. Harrison, eds), London: Academic Press.

    Google Scholar 

  • Gallivan, G. J. and Ronald, K. (1979) Temperature regulation in freely diving harp seals (Phoca groenlandica). Can. J. Zool. 57: 2256–2263.

    Article  PubMed  CAS  Google Scholar 

  • Gentry, R. L. (1970) Social behaviour of the Steller sea lion. PhD thesis, University of California, Santa Cruz, 113 pp.

    Google Scholar 

  • Gentry, R. L. (1973) Thermoregulatory behaviour of eared seals. Behaviour 46: 73–93.

    Article  PubMed  CAS  Google Scholar 

  • Gentry, R. L. (1974) The development of social behaviour through play in the Steller sea lion. Am. Zool. 14: 391–403.

    Google Scholar 

  • Gentry, R. L. and Holt, J. R. (1986) Attendance behaviour of northern fur seals, in Fur Seals: Maternal Strategies on Land and at Sea: 41–60 (R. L. Gentry and G. L. Kooyman, eds), Princeton, NJ: Princeton University Press.

    Google Scholar 

  • Gentry, R. L., Kooyman, G. L. and Goebel, M. E. (1986a) Feeding and diving behaviour of northern fur seals, in Fur Seals: Maternal Strategies on Land and at Sea: 61–78 (R. L. Gentry and G. L. Kooyman, eds), Princeton, NJ: Princeton University Press.

    Google Scholar 

  • Gentry, R. L., Costa, D. P., Croxall, J. P., David, J. H. M., Davis, R. W., Kooyman, G. L., Majluf, P., McCann, T. S. and Trillmich, F. (1986b) Synthesis and Conclusions, in Fur Seals: Maternal Strategies on Land and at Sea: 220–264 (R. L. Gentry and G. L. Kooyman, eds), Princeton, NJ: Princeton University Press.

    Google Scholar 

  • Godsell, J. (1988) Herd formation and haul-out behaviour in harbour seals (Phoca vitulina). J. Zool., London 215: 83–98.

    Article  Google Scholar 

  • Grav, H. J. and Blix, A. S. (1976) Brown adipose tissue: A factor in the survival of harp seal pups. Can. J. Physiol. Pharmacol. 54: 409–412.

    Article  PubMed  CAS  Google Scholar 

  • Grav, H. J. and Blix, A. S. (1979) A source of non-shivering thermogenesis in fur seal skeletal muscle. Science 204: 87–89.

    Article  PubMed  CAS  Google Scholar 

  • Green, K. and Burton, H. R. (1988) Annual and diurnal variations in the underwater vocalizations of Weddell seals. Polar Biol. 8: 161–164.

    Article  Google Scholar 

  • Guppy, M., Hill, R. D., Schneider, R. C., Qvist, J., Liggins, G. C., Zapol, W. M. and Hochachka, P. W. (1986) Microcomputer-assisted metabolic studies of voluntary diving of Weddell seals. Am. J. Physiol. 250: R175–R187.

    PubMed  CAS  Google Scholar 

  • Harcourt, A. H., Harvey, P. H., Larson, S. G. and Short, R. V. (1981) Testis weight, body weight and breeding system in primates. Nature 293: 55–57.

    Article  PubMed  CAS  Google Scholar 

  • Harrison, R. J. and Kooyman, G. L. (1968) General physiology of the pinnipedia, in The Behaviour and Physiology of Pinnipeds: 211–296 (R. J. Harrison, R. C. Hubbard, R. S. Peterson, C. E. Rice and R. J. Schusterman, eds), New York: Appleton-Century-Crofts.

    Google Scholar 

  • Heinrich, B. (1977) Why have some animals evolved to regulate a high body temperature?. Am. Nat. 111: 623–640.

    Article  Google Scholar 

  • Hill, R. D., Schneider, R. C., Liggins, G. C. et al. (1987) Heart rate and body temperature during free diving of Weddell seals. Am. J. Physiol. 253: R344–R351.

    PubMed  CAS  Google Scholar 

  • Hochachka, P. W. and Guppy, M. (1987) Metabolic Arrest and the Control of Biological Time. Cambridge, MA: Harvard University Press.

    Google Scholar 

  • Hochachka, P. W. and Somero, G. N. (1973) Strategies of Biochemical Adaptation. Philadelphia: Saunders.

    Google Scholar 

  • Hogg, J. T. (1984) Mating in bighorn sheep: Multiple creative male strategies. Science 225: 526–529.

    Article  PubMed  CAS  Google Scholar 

  • Huber, H. R. (1987) Natality and weaning success in relation to age of first reproduction in northern elephant seals. Can J. Zool. 65: 1311–1316.

    Article  Google Scholar 

  • Huntley, A. C., Costa, D. P. and Rubin, R. D. (1984) The contribution of nasal countercurrent heat exchange to water balance in the northern elephant seal, Mirounga angustirostris. J. Exp. Biol. 113: 447–454.

    PubMed  CAS  Google Scholar 

  • Hyvärinen, H. (1989) Diving in darkness: whiskers as sense organs of the ringed seal (Phoca hispida saimensis). J. Zool., London 218: 663–678.

    Article  Google Scholar 

  • Innes, S., Lavigne, D. M., Earle, W. M. and Kovacs, K. M. (1987) Feeding rates of seals and whales. J. Anim. Ecol. 56: 115–130.

    Article  Google Scholar 

  • Irving, L. (1938) Changes in the blood flow through the brain and muscles during the arrest of breathing. Am J. Physiol. 122: 207–214.

    Google Scholar 

  • Irving, L. (1939) Respiration in diving mammals. Physiol. Rev. 19: 112–134.

    Google Scholar 

  • Irving, L. (1972) Arctic Life of Birds and Mammals Including Man. New York: Springer-Verlag.

    Book  Google Scholar 

  • Irving, L., Scholander, P. F. and Grinnell, S. W. (1941) Significance of the heart rate to the diving ability of seals. J. Cell. Comp. Physiol. 18: 283–297.

    Article  CAS  Google Scholar 

  • Irving, L., Scholander, P. F. and Grinnell, S. W. (1942) The regulation of the arterial blood pressure in the seal during diving. Am. J. Physiol. 135: 557–566.

    Google Scholar 

  • James, H. and Dykes, R. W. (1978) Some experiments on navigation in the harbour seal, Phoca vitulina, in Animal Migration, Navigation and Homing: 395–404 (K. Schmidt-Koenig and W. T. Keeton, eds), New York: Springer-Verlag.

    Google Scholar 

  • Jangaard, P. M. and Ke, P. J. (1968) Principal fatty acids of depot fat and milk lipids from harp seal (Pagophilus groenlandicus) and hooded seal (Cystophora cristata). J. Fish. Res. Board Can. 25: 2419–2426.

    Article  CAS  Google Scholar 

  • John, T. M., Ronald, K. and George, J. C. (1987) Blood levels of thyroid hormones and certain metabolites in relation to moult in the harp seal (Phoca groenlandica). Comp. Biochem. Physiol. 88A: 655–657.

    Article  CAS  Google Scholar 

  • Johnstone, R. M. and Davis, L. S. (1987) Activity in a non-breeding colony of New Zealand fur seals, Arctocephalus forsteri (Note). NZ J. Marine and Freshwater Res. 21: 153–155.

    Article  Google Scholar 

  • Kajimura, H. (1985) Opportunistic feeding by the northern fur seal, (Callorhinus ursinus), in Marine Mammals and Fisheries: 300–318 (J. R. Beddington, R. J. H. Beverton and D. M. Lavigne, eds), London: George Allen & Unwin.

    Google Scholar 

  • Kenagy, G. J. and Trombulak, S. C. (1986) Size and function of mammalian testes in relation to body size. J. Mammal. 67: 1–22.

    Article  Google Scholar 

  • Kennedy, G. C. and Mitra, J. (1963) Body weight and food intake as initiation factors for puberty in the rat. J. Physiol. (London) 166: 408–418.

    CAS  Google Scholar 

  • Keyes, M. C., Kim, K. C., Dailey, M. D. and Vendros, N. A. (1981) Fur seal investigations 1980. Northwest and Alaska Fisheries Center. Processed Reports of the US Department of Commerce, Seattle, Washington 81-2: 33–34.

    Google Scholar 

  • Kleiber, M. (1975) The Fire of Life, 2nd edn, revised. New York: Wiley.

    Google Scholar 

  • Kooyman, G. L. (1966) Maximum diving capacities of the Weddell seal, Leptonychotes weddelli. Science 151: 1553–1554.

    Article  PubMed  CAS  Google Scholar 

  • Kooyman, G. L. (1968) An analysis of some behavioural and physiological characteristics related to diving in the Weddell seal, in Antarctic Research Series: Vol. 11. Biology of the Antarctic Seas III: 227–261. (W. L. Schmitt and G. A. Llano, eds), Washington, DC: American Geophysical Union.

    Google Scholar 

  • Kooyman, G. L. (1972) Deep diving behaviour and effects of pressure in reptiles, birds, and mammals. Symp. Soc. Exp. Biol. 26: 295–311.

    PubMed  CAS  Google Scholar 

  • Kooyman, G. L. (1975) A comparison between day and night diving in the Weddell seal. J. Mammal. 56: 563–574.

    Article  Google Scholar 

  • Kooyman, G. L. (1981) Weddell Seal: Consummate Diver. Cambridge: Cambridge University Press.

    Book  Google Scholar 

  • Kooyman, G. L. (1989) Diving physiology of marine mammals, in Comparative Pulmonary Physiology: Current Concepts: 721–734 (S. C. Wood, ed.), New York: Marcel Dekker.

    Google Scholar 

  • Kooyman, G. L. and Campbell, W. B. (1972) Heart rates in freely diving Weddell seals, Leptonychotes weddelli. Comp. Biochem. Physiol. 43A: 31–36.

    Article  Google Scholar 

  • Kooyman, G. L. and Trillmich, F. (1986a) Diving behaviour of Galapagos fur seals, in Fur Seals: Maternal Strategies on Land and at Sea: 186–195 (R. L. Gentry and G. L. Kooyman, eds), Princeton, NJ: Princeton University Press.

    Google Scholar 

  • Kooyman, G. L. and Trillmich, F. (1986b) Diving behaviour of Galapagos sea lions, in Fur Seals: Maternal Strategies on Land and at Sea: 209–219 (R. L. Gentry and G. L. Kooyman, eds), Princeton, NJ: Princeton University Press.

    Google Scholar 

  • Kooyman, G. L., Hammond, D. D. and Schroeder, J. P. (1970) Bronchograms and tracheograms of seals under pressure. Science 169: 82–84.

    Article  PubMed  CAS  Google Scholar 

  • Kooyman, G. L., Wahrenbrock, E. A., Castellini, M. A., Davis, R. W. and Sinnett, E. E. (1980) Aerobic and anaerobic metabolism during voluntary diving in Weddell seals: evidence of preferred pathways from blood chemistry and behaviour. J. Comp. Physiol. 138: 335–346.

    CAS  Google Scholar 

  • Kooyman, G. L., Castellini, M. A. and Davis, R. W. (1981) Physiology of diving in marine mammals. Annu. Rev. Physiol. 43: 343–356.

    Article  PubMed  CAS  Google Scholar 

  • Kooyman, G. L., Castellini, M. A., Davis, R. W. and Maue, R. A. (1983) Aerobic diving limits of immature Weddell seals. J. Comp. Physiol. B 151: 171–174.

    Article  Google Scholar 

  • Kooyman, G. L., Davis, R. W. and Croxall, J. P. (1986) Diving behaviour of antarctic fur seals, in Fur Seals: Maternal Strategies on Land and at Sea: 115–125 (R. L. Gentry and G. L. Kooyman, eds), Princeton, NJ: Princeton University Press.

    Google Scholar 

  • Kovacs, K. M. (1987) Maternal behaviour and early behavioural ontogeny of grey seals (Halichoerus grypus) on the Isle of May, UK. J. Zool., London 213: 697–715.

    Article  Google Scholar 

  • Kovacs, K. M. and Lavigne, D. M. (1986) Maternal investment and neonatal growth in phocid seals. J. Anim. Ecol. 55: 1035–1051.

    Article  Google Scholar 

  • Krieber, M. and Barrette, C. (1984) Aggregation behaviour of harbour seals at Forillon National Park, Canada. J. Anim. Ecol. 53: 913–928.

    Article  Google Scholar 

  • Lavigne, D. M. and Øritsland, N. A. (1974) Ultraviolet photography: A new application for remote sensing of mammals. Can. J. Zool. 52: 939–941.

    Article  PubMed  CAS  Google Scholar 

  • Lavigne, D. M., Barchard, W., Innes, S. and Øritsland, N. A. (1982a) Pinniped Bioenergetics. FAO Fish Ser. No. 5 4: 191–235.

    Google Scholar 

  • Lavigne, D. M., Stewart, R. E. A. and Fletcher, F. (1982b) Changes in composition and energy content of harp seal milk during lactation. Physiol. Zool. 55: 1–9.

    Google Scholar 

  • Lavigne, D. M., Innes, S., Worthy, G. A. J., Kovacs, K. M., Schmitz, O. J. and Hickie, J. P. (1986) Metabolic rates of seals and whales. Can. J. Zool. 64: 279–284.

    Article  Google Scholar 

  • Laws, R. M. (1956a) The elephant seal (Mirounga leonina, Linn.) I. General, social and reproductive behaviour. FIDS Scientific Report No. 13, 88 pp.

    Google Scholar 

  • Laws, R. M. (1956b) Growth and sexual maturity in aquatic mammals. Nature 178: 193–194.

    Article  Google Scholar 

  • Laws, R. M. (1959) Accelerated growth in seals, with special reference to the Phocidae. Nor Hvalfangst-T 9: 425–452.

    Google Scholar 

  • Laws, R. M. (1977) The significance of vertebrates in the antarctic marine ecosystem, in Adaptations Within Antarctic Ecosystems: Proceedings of the Third SCAR Symposium on Antarctic Biology: 411–438 (G. A. Llano, ed.), Washington, DC: Smithsonian Institution.

    Google Scholar 

  • Lawson, J. and Renouf, D. (1985) Parturition in the Atlantic harbour seal. J. Mammal. 66: 395–398.

    Google Scholar 

  • Le Boeuf, B. J. (1972) Sexual behaviour in the northern elephant seal, Mirounga angustirostris. Behaviour 41: 1–26.

    Article  Google Scholar 

  • Le Boeuf, B. J. (1974) Male-male competition and reproductive success in elephant seals. Am. Zool. 14: 163–176.

    Google Scholar 

  • Le Boeuf, B. J. and Mesnick, S. (in press) Sexual behaviour of male northern elephant seals I: Lethal injuries to adult females. Behaviour.

    Google Scholar 

  • Le Boeuf, B. J. and Briggs, K. T. (1977) The cost of living in a seal harem. Mammalia 41: 167–195.

    Google Scholar 

  • Le Boeuf, B. J. and Reiter, J. (1988) Lifetime reproductive success in northern elephant seals, in Reproductive Success: 344–362 (T. Clutton-Brock, ed.), Chicago: University of Chicago Press.

    Google Scholar 

  • Le Boeuf, B. J., Whiting, R. J. and Gantt, R. F. (1972) Perinatal behaviour of northern elephant seal females and their young. Behaviour 43: 121–156.

    Article  PubMed  Google Scholar 

  • Le Boeuf, B. J., Riedman, M. and Keyes, R. S. (1982) White shark predation on pinnipeds in California coastal waters. Fish. Bull. 80: 891–895.

    Google Scholar 

  • Le Boeuf, B. J., Costa, D. P., Huntley, A. C. and Feldkamp, S. D. (1988) Continuous, deep diving in female northern elephant seals, Mirounga angustirostris. Can. J. Zool. 66: 446–458.

    Article  Google Scholar 

  • Le Boeuf, B. J., Condit, R. and Reiter, J. (1989a) Parental investment and secondary sex ratio in northern elephant seals. Behav. Ecol. Sociobiol. 25: 109–117

    Article  Google Scholar 

  • Le Boeuf, B. J., Naito, Y., Huntley, A. C. and Asaga, T. (1989b) Prolonged, continuous, deep diving by northern elephant seals. Can. J. Zool. 67: 2514–2519.

    Article  Google Scholar 

  • Lever, M. J., Miller, K. W., Paton, W. D. M. and Smith, E. B. (1971) Pressure reversal of anaesthesia. Nature 231: 368–371.

    Article  PubMed  CAS  Google Scholar 

  • Limberger, D., Trillmich, F., Biebach, H. and Stevenson, R. D. (1986) Temperature regulation and microhabitat choice by free-ranging Galapagos fur seal pups (Arctocephalus galapagoensis). Oecologia (Berlin) 69: 53–59.

    Article  Google Scholar 

  • Ling, J. K. (1974) The integument of marine mammals, in Functional Anatomy of Marine Mammals: 1–44 (R. J. Harrison, ed.), New York: Academic Press.

    Google Scholar 

  • Loughlin, T. R., Bengtson, J. L., and Merrick, R. L. (1987) Characteristics of feeding trips of female northern fur seals. Can. J. Zool. 65: 2079–2084.

    Article  Google Scholar 

  • Marlow, B. G. (1975) The comparative behaviour of the Australasian sea lions Neophoca cinerea and Phocarctos hookeri (Pinnipedia: Otariidae). Mammalia 39: 159–230.

    Article  Google Scholar 

  • Maynard Smith, J. (1980) A new theory of sexual investment. Behav. Ecol. Sociobiol. 7: 247–251.

    Article  Google Scholar 

  • Mayr, E. (1974) Behaviour programs and evolutionary strategies. Am. Sci. 62: 650–659.

    PubMed  CAS  Google Scholar 

  • McCann, T. S. (1980) Territoriality and breeding behaviour of adult male antarctic fur seal, Arctocephalus gazella. J. Zool., London 192: 295–310.

    Article  Google Scholar 

  • McCann, T. S. (1981) Aggression and sexual activity of male southern elephant seals, Mirounga leonina. J. Zool., London 195: 295–310.

    Article  Google Scholar 

  • McCann, T. S. (1982) Aggressive and maternal activities of female southern elephant seals (Mirounga leonina). Anim. Behav. 30: 268–276.

    Article  Google Scholar 

  • McCann, T. S. (1983) Activity budgets of southern elephant seals Mirounga leonina during the breeding season. Z. Tierpsychol. 61: 111–126.

    Article  Google Scholar 

  • McLaren, I. A. (1958) The biology of the ringed seal, Phoca hispida, in the eastern Canadian arctic. Bull. Fish. Res. Board Can., No. 118.

    Google Scholar 

  • McLaren, I. A. (1967) Seals and group selection. Ecology 48: 104–110.

    Article  Google Scholar 

  • Miller, E. H. (1975a) Annual cycle of fur seals, Arctocephalus forsteri (Lesson), on the Open Bay Islands, New Zealand. Pacific Sci. 29: 139–152.

    Google Scholar 

  • Miller, E. H. (1975b) Social and evolutionary implications of territoriality in adult male New Zealand fur seals, Arctocephalus forsteri (Lesson, 1828), during the breeding season, in Rapports et Procès-Verbaux des Réunions: Vol 169. Biology of the Seal: 170–187 (K. Ronald and A. W. Mansfield, eds), Conseil International pour L’Exploration de la Mer, Charlottenlund Slot, Denmark.

    Google Scholar 

  • Miller, E. H. and Boness, D. J. (1979) Remarks on display functions of the snout of the grey seal, Halichoerus grypus (Fab.), with comparative notes. Can. J. Zool. 57: 140–148.

    Article  Google Scholar 

  • Miller, L. K. (1978) Energetics of the northern fur seal in relation to climate and food resources of the Bering Sea. Final Report to the US Marine Mammal Commission. NTIS, US Department of Commerce, Springfield, VA, 27 pp.

    Google Scholar 

  • Montagna, W. (1967) Comparative anatomy and physiology of the skin. Arch. Dermatol. 96: 357–363.

    Article  PubMed  CAS  Google Scholar 

  • Mortola, J. P. and Lanthier, C. (1989) Normoxic and hypoxic breathing pattern in newborn grey seals. Can. J. Zool. 67: 483–487.

    Article  Google Scholar 

  • Nottebohm, F. (1970) Ontogeny of bird song. Science 167: 950–956.

    Article  PubMed  CAS  Google Scholar 

  • Odell, D. K. (1974) Behavioural thermoregulation in the California sea lion. Behav. Biol. 10: 231–237.

    Article  PubMed  CAS  Google Scholar 

  • Oftedal, O. T., Boness, D. J. and Tedman, R. A. (1987) The behaviour, physiology, and anatomy of lactation in the pinnipedia. Curr. Mammal. 1: 175–245.

    Google Scholar 

  • Oftedal, O. T., Boness, D. J. and Bowen, W. D. (1988) The composition of hooded seal (Cystophora cristata) milk: An adaptation for postnatal fattening. Can. J. Zool. 66: 318–322.

    Article  CAS  Google Scholar 

  • Oliver, G. W. (1978) Navigation in mazes by a grey seal, Halichoerus grypus (Fabricius). Behaviour 67: 97–114.

    Article  Google Scholar 

  • Ono, K. A., Boness, D. J. and Oftedal, O. T. (1987) The effect of a natural environmental disturbance on maternal investment and pup behaviour in the California sea lion. Behav. Ecol. Sociobiol 21: 109–118.

    Article  Google Scholar 

  • Øritsland, N. A. (1968) Variations in the body surface temperature of the harp seal. Acta Physiol. Scand. 73: 35A–36A.

    Article  Google Scholar 

  • Øritsland, N. A. (1970) Energetic significance of absorption of solar radiation in polar homeotherms, in Antarctic Ecology: 464–470 (M. W. Holdgate, ed.), New York: Academic Press.

    Google Scholar 

  • Øritsland, N. A. and Ronald, K. (1978) Aspects of temperature regulation in harp seal pups evaluated by in vivo experiments and computer simulations. Acta Physiol Scand. 103: 263–269.

    Article  PubMed  Google Scholar 

  • Ortiz, C. L., Costa, D. and Le Boeuf, B. J. (1978) Water and energy flux in elephant seal pups fasting under natural conditions. Physiol Zool 51: 166–178.

    CAS  Google Scholar 

  • Parker, E. D. (1984) Sperm competition and the evolution of animal mating strategies, in Sperm Competition and the Evolution of Animal Mating Systems: 1–60 (R. L. Smith, ed.), Orlando, FL: Academic Press.

    Google Scholar 

  • Payne, M. R. (1977) Growth of a fur seal population. Philosophical Trans. R. Soc. London (B) 279: 67–79.

    Article  Google Scholar 

  • Payne, M. R. (1979) Growth in the antarctic fur seal Arctocephalus gazella. J. Zool, London 187: 1–20.

    Article  Google Scholar 

  • Pedersen, J. I. and Grav, H. J. (1972) Physiologically-induced loose coupling of brown-adipose-tissue mitochondria correlated to endogenous fatty acids and adenosine phosphates. Eur. J. Biochem. 25: 75–83.

    Article  PubMed  CAS  Google Scholar 

  • Perez, M. A. and Bigg, M. A. (1986) Diet of northern fur seals, Callorhinus ursinus, off western North America. Fish. Bull. 84: 957–971.

    Google Scholar 

  • Perez, M. A. and Mooney, E. E. (1986) Increased food and energy consumption of lactating northern fur seals Callorhinus ursinus. Fish. Bull. 84: 371–381.

    Google Scholar 

  • Perry, E. A. and Renouf, D. (1988) Further studies of the role of harbour seal (Phoca vitulina) pup vocalizations in preventing separation of mother-pup pairs. Can. J. Zool. 66: 934–938.

    Article  Google Scholar 

  • Peterson, R. S. (1968) Social behaviour in pinnipeds with particular reference to the northern fur seal, in The Behaviour and Physiology of Pinnipeds: 3–53 (R. J. Harrison, R. C. Hubbard, R. S. Peterson, C. E. Rice and R. J. Schusterman, eds), New York: Appleton-Century-Crofts.

    Google Scholar 

  • Peterson, R. S. and Bartholomew, G. A. (1967) The Natural History and Behaviour of the California Sea Lion. Special Publication Number 1. Shippensburg, PA: American Society of Mammalogists.

    Google Scholar 

  • Pianka, E. R. and Parker, W. S. (1975) Age-specific reproductive tactics. Am. Nat. 109: 453–464.

    Article  Google Scholar 

  • Pilson, M. E. Q. and Kelly, A. L. (1962) Composition of the milk from Zalophus californianus, the California sea lion. Science 135: 104–105.

    Article  PubMed  CAS  Google Scholar 

  • Plötz, J. (1986) Summer diet of Weddell seals (Leptonychotes weddelli) in the eastern and southern Weddell Sea, Antarctica. Polar Biol. 6: 97–102.

    Article  Google Scholar 

  • Ralls, K. (1977) Sexual dimorphism in mammals: avian models and unanswered questions. Am. Nat. 111: 917–938.

    Article  Google Scholar 

  • Rand, R. W. (1955) Reproduction in the female Cape fur seal, Arctocephalus pusillus (Schreber). Proc. Zool. Soc. London 124: 717–740.

    Article  Google Scholar 

  • Rand, R. W. (1956) The Cape fur-seal, Arctocephalus pusillus (Schreber) 1. Its general characteristics and moult. Investigational Report 21. Dept. of Commerce and Industries, Division of Sea Fisheries, South Africa. 52 pp.

    Google Scholar 

  • Rand, R. W.(1967) The Cape fur-seal (Arctocephalus pusillus). 3. General behaviour on land and at sea. Investigational Report 60. Dept. of Commerce and Industries, Division of Sea Fisheries, South Africa. 39 pp.

    Google Scholar 

  • Rasmusson, E. M. and Wallace, J. M. (1983) Meteorological aspects of the El Niño/southern oscillation. Science 222: 1195–1202.

    Article  PubMed  CAS  Google Scholar 

  • Ray, G. C. and Watkins, W. A. (1975) Social function of underwater sounds in the walrus Odobenus rosmarus, in Rapports et Procès-Verbaux des Réunions: Vol. 169. Biology of the Seal: 524–526 (K. Ronald and A. W. Mansfield, eds), Conseil International pour L’Exploration de la Mer, Charlottenlund Slot, Denmark.

    Google Scholar 

  • Reiter, J., Stinson, N. L. and Le Boeuf, B. J. (1978) Northern elephant seal development: The transition from weaning to nutritional independence. Behav. Ecol. Sociobiol. 3: 337–367.

    Article  Google Scholar 

  • Reiter, J., Panken, K. J. and Le Boeuf, B. J. (1981) Female competition and reproductive success in northern elephant seals. Anim. Behav. 29: 670–687.

    Article  Google Scholar 

  • Reiter, R. J. (1973) Comparative physiology: Pineal gland. Annu. Rev. Physiol. 35: 305–328.

    Article  PubMed  CAS  Google Scholar 

  • Renouf, D. (1984) The vocalization of the harbour seal pup (Phoca vitulina) and its role in the maintenance of contact with the mother. J. Zool., London 202: 583–590.

    Article  Google Scholar 

  • Renouf, D. (1985) A demonstration of the ability of the harbour seal Phoca vitulina (L.) to discriminate among pup vocalizations. J. Exp. Marine Biol. Ecol. 87: 41–46.

    Article  Google Scholar 

  • Renouf, D. (1989) Sensory function in the harbor seal. Sci. Am. 260: 90–95.

    Article  PubMed  CAS  Google Scholar 

  • Renouf, D. and Gaborko, L. (1982) Speed sensing in a harbour seal. J. Marine Biol. Assoc. UK 62: 227–228.

    Article  Google Scholar 

  • Renouf, D. and Lawson, J. W. (1986) Harbour seal vigilance: watching for predators or mates. Biol. Behav. 11: 44–49.

    Google Scholar 

  • Renouf, D. and Lawson, J. W. (1987) Quantitative aspects of harbour seal [Phoca vitulina) play. J. Zool, London 212: 267–273.

    Article  Google Scholar 

  • Ribic, C. A. (1988) Maternal aggression in northern elephant seals: The effect of the pup. Can. J. Zool. 66: 1693–1698.

    Article  Google Scholar 

  • Ridgway, S. H. and Robison, C. C. (1985) Homing by released captive California sea lions, Zalophus californianus, following release on distant islands. Can. J. Zool. 63: 2162–2164.

    Article  Google Scholar 

  • Riedman, M. and Ortiz, C. L. (1979) Changes in milk composition during lactation in the northern elephant seal. Physiol. Zool. 52: 240–249.

    CAS  Google Scholar 

  • Riviere, J. E., Engelhardt, F. R. and Solomon, J. (1977) The relationship of thyroxine and cortisol to the moult of the harbour seal Phoca vitulina. Gen. Comp. Endocrinol. 31: 398–401.

    Article  PubMed  CAS  Google Scholar 

  • Ronald, K. and Dougan, J. L. (1982) The ice lover: Biology of the harp seal (Phoca groenlandica). Science 215: 928–933.

    Article  PubMed  CAS  Google Scholar 

  • Ryg, M., Smith, T. G. and Øritsland, N. A. (1988) Thermal significance of the topographical distribution of blubber in ringed seals (Phoca hispida). Can. J. Fish. Aquatic Sci. 45: 985–992.

    Article  Google Scholar 

  • Sadlier, R. M. F. S. (1969) The Ecology of Reproduction in Wild and Domestic Mammals. London: Methuen.

    Book  Google Scholar 

  • Salter, R. E. (1979) Site utilization, activity budgets, and disturbance responses of Atlantic walmses during terrestrial haul-out. Can. J. Zool. 57: 1169–1180.

    Article  Google Scholar 

  • Sandegren, F. E. (1970) Breeding and Maternal Behaviour of the Steller Sea Lion (Eumetopias jubatus) in Alaska. MS thesis, University of Alaska, Fairbanks, 138 pp.

    Google Scholar 

  • Sandegren, F. E. (1976) Courtship display, agonistic behaviour and social dynamics in the Steller sea lion (Eumetopias jubatus). Behaviour 57: 159–172.

    Article  Google Scholar 

  • Schmidt-Nielsen, K. and Schmidt-Nielsen, B. (1952) Water metabolism of desert mammals. Physiol. Rev. 32: 135–166.

    PubMed  CAS  Google Scholar 

  • Scholander, P. F. (1940) Experimental investigations on the respiratory functions in diving mammals and birds. Hvalrdets Skrifter 22: 1–131.

    Google Scholar 

  • Scholander, P. F., Irving, L. and Grinnell, S. W. (1942a) Aerobic and anaerobic changes in the seal muscles during diving. J. Biol. Chem. 142: 432–440.

    Google Scholar 

  • Scholander, P. F., Irving, L. and Grinnell, S. W. (1942b) On the temperature and metabolism of the seal during diving. J. Cell. Comp. Physiol. 19: 67–78.

    Article  CAS  Google Scholar 

  • Sharman, G. B. (1963) Delayed implantation in marsupials, in Delayed Implantation: 3–14 (A. C. Enders ed.) Chicago: University Chicago Press.

    Google Scholar 

  • Shipley, C. and Strecker, G. (1986) Day and night patterns of vocal activity of northern elephant seal bulls. J. Mammal. 67: 775–778.

    Article  Google Scholar 

  • Shipley, C, Hines, M. and Buchwald, J. S. (1986) Vocalizations of northern elephant seal bulls: development of adult call characteristics during puberty. J. Mammal 67: 526–536.

    Article  Google Scholar 

  • Siniff, D. B., Stirling, I., Bengston, J. L. and Reichle, R. A. (1979) Social and reproductive behaviour of crabeater seals (Lobodon carcinophagus) during the austral spring. Can. J. Zool. 57: 2243–2255.

    Article  Google Scholar 

  • Smith, M. S. R. (1965) Seasonal movements of the Weddell seal in McMurdo Sound, Antarctica. J. Wildlife Management 29: 464–470.

    Article  Google Scholar 

  • Smith, R. E. and Horwitz, B. A. (1969) Brown fat and thermogenesis. Physiol. Rev. 49: 330–425.

    PubMed  CAS  Google Scholar 

  • Stewart, R. E. A. (1986) Energetics of age-specific reproductive effort in female harp seals, Phoca groenlandica. J. Zool, London, Series A 208: 503–517.

    Google Scholar 

  • Stewart, R. E. A. (1987) Behavioural reproductive effort of nursing harp seals, Phoca groenlandica. J. Mammal. 68: 348–358.

    Article  Google Scholar 

  • Stewart, R. E. A. and Lavigne, D. M. (1980) Neonatal growth of Northwest Atlantic harp seals, Pagophilus groenlandicus. J. Mammal. 61: 670–680.

    Article  PubMed  CAS  Google Scholar 

  • Stewart, R. E. A. and Lavigne, D. M. (1984) Energy transfer and female condition in nursing harp seals Phoca groenlandica. Holarctic Ecol. 7: 182–194.

    Google Scholar 

  • Stirling, I. (1969) Ecology of the Weddell seal in McMurdo Sound, Antarctica. Ecology 50: 573–586.

    Article  Google Scholar 

  • Stirling, I. (1971) Studies on the behaviour of the South Australian fur seal, Arctocephalus forsteri (Lesson). I. Annual cycle, postures and calls, and adult males during the breeding season. Aust. J. Zool. 19: 243–266.

    Article  Google Scholar 

  • Stirling, I. (1973) Vocalization in the ringed seal [Phoca hispida). J. Fish. Res. Board Can. 30: 1592–1594.

    Article  Google Scholar 

  • Stirling, I. (1983) The evolution of mating systems in pinnipeds, in Recent Advances in the Study of Mammalian Behaviour: 489–527 (J. F. Eisenberg and D. G. Kleiman, eds), Shippensburg, PA: American Society of Mammalogists.

    Google Scholar 

  • Stirling, I., Calvert, W. and Cleator, H. (1983) Underwater vocalizations as a tool for studying the distribution and relative abundance of wintering pinnipeds in the high arctic. Arctic 36: 262–274.

    Google Scholar 

  • Tarasoff, F. J. and Toews, D. P. (1972) The osmotic and ionic regulatory capacities of the kidney of the harbour seal, Phoca vitulina. J. Comp. Physiol. 81: 121–132.

    Article  CAS  Google Scholar 

  • Tedman, R. A. and Bryden, M. M. (1979) Cow-pup behaviour of the Weddell seal, Leptonychotes weddelli (Pinnipedia), in McMurdo Sound, Antarctica. Aust. Wildlife Res. 6: 19–37.

    Article  Google Scholar 

  • Terhune, J. M. (1985) Scanning behaviour of harbour seals on haul-out sites. J. Mammal. 66: 392–395.

    Article  Google Scholar 

  • Terhune, J. M. and Ronald, K. (1976) Examining harp seal behavioural patterns via their underwater vocalizations. Appl. Anim. Ethol. 2: 261–264

    Article  Google Scholar 

  • Terhune, J. M., Terhune, M. E. and Ronald, K. (1979) Location and recognition of pups by adult female harp seals. Appl. Anim. Ethol. 5: 375–380.

    Article  Google Scholar 

  • Thomas, J. A. and DeMaster, D. P. (1983) Diel haul-out patterns of Weddell seal (Leptonychotes weddelli) females and their pups. Can. J. Zool. 61: 2084–2086.

    Article  Google Scholar 

  • Thompson, P. (1988) Timing of mating in the common seal (Phoca vitulina). Mammal Rev. 18: 105–112.

    Article  Google Scholar 

  • Thompson, P. M., Fedak, M. A., McConnell, B. J. and Nicholas, K. S. (1989) Seasonal and sex-related variation in the activity patterns of common seals (Phoca vitulina). J. Appl. Ecol. 26: 521–535.

    Article  Google Scholar 

  • Trillmich, F. (1984) Natural history of the Galapagos fur seal (Arctocephalus galapagoensis, Heller), in Key Environments: Galapagos: 215–223 (R. Perry, ed.), Oxford: Pergamon Press. Trillmich, F. (1986a) Attendance behaviour of Galapagos fur seals, in Fur Seals: Maternal Strategies on Land and at Sea: 168–185 (R. L. Gentry and G. L. Kooyman, ed.), Princeton, NJ: Princeton University Press.

    Google Scholar 

  • Trillmich, F. (1986b) Attendance behaviour of Galapagos sea lions, in Fur Seals: Maternal Strategies on Land and at Sea: 196–208 (R. L. Gentry and G. L. Kooyman, eds), Princeton, NJ: Princeton University Press.

    Google Scholar 

  • Trillmich, F. (1986c) Maternal investment and sex-allocation in the Galapagos fur seal, Arctocephalus galapagoensis. Behav. Ecol. Sociobiol. 19: 157–164.

    Google Scholar 

  • Trillmich, F. and Limberger, D. (1985) Drastic effects of El Niño on Galapagos pinnipeds. Oecologia (Berlin) 67: 19–22.

    Article  Google Scholar 

  • Trillmich, F. and Trillmich, K. G. K. (1984) The mating systems of pinnipeds and marine iguanas: convergent evolution of polygyny. Biol. J. Linnean Soc. 21: 209–216.

    Article  Google Scholar 

  • Trillmich, F., Kooyman, G. L., Majluf, P. and Sanchez-Grinan, M. (1986) Attendance and diving behaviour of South American fur seals during El Niño in 1983, in Fur Seals: Maternal Strategies on Land and at Sea: 153–167 (R. L. Gentry and G. L. Kooyman, eds), Princeton, NJ: Princeton University Press.

    Google Scholar 

  • Trivers, R. L. and Willard, D. E. (1973) Natural selection of parental ability to vary sex ratios of offspring. Science 179: 90–92.

    Article  PubMed  CAS  Google Scholar 

  • Vaz-Ferreira, R. and Achaval, F. (1979) Relati ón y reconocimiento maternofilial en Otaria flavescens (Shaw) ‘Lobo de un pelo’, y reacciones de los machos subadultos ante los cachorros. Acta Zool. Lilloana 35: 295–302.

    Google Scholar 

  • Wartzok, D. and Ray, G. C. (1980) Hauling-out Behaviour of the Pacific Walrus. Final report MM5ACO28. Marine Mammal Commission, Washington, DC v + 46 pp.

    Google Scholar 

  • Wartzok, D., Elsner, R., Kelly, B., Mimken, G. and Quakenbush, L. (1987) Visual, acoustic and vibrissal contributions to under-ice navigation by ringed seals, Phoca hispida. Proc. the Seventh Biennial Conference on the Biology of Marine Mammals, Miami. Abstract.

    Google Scholar 

  • Wartzok, D., Elsner, R., Davis, R. and Mimken, G. (1990) Under-ice movements of Weddell seals. Antarctic J. 24: 205–206.

    Google Scholar 

  • White, F. N. and Odell, D. K. (1971) Thermoregulatory behaviour of the northern elephant seal, Mirounga angustirostris. J. Mammal. 52: 758–774.

    Article  Google Scholar 

  • Whittow, G. C. (1978) Thermoregulatory behaviour of the Hawaiian monk seal (Monachus schauinslandi). Pacific Sci. 32: 47–60.

    Google Scholar 

  • Whittow, G. C, Ohata, C. A. and Matsuura, D. T. (1971) Behavioural control of body temperature in the unrestrained California sea lion. Comm. Behav. Biol. 6: 87–91.

    CAS  Google Scholar 

  • Whittow, G. C, Matsuura, D. T. and Lin, Y. C. (1972) Temperature regulation in the California sea lion (Zalophus californianus). Physiol. Zool. 45: 68–77.

    Google Scholar 

  • Worthy, G. A. J. and Lavigne, D. M. (1983) Energetics of fasting and subsequent growth in weaned harp seal pups, Phoca groenlandica. Can. J. Zool. 61: 447–456.

    Article  Google Scholar 

  • Worthy, G. A. J. and Lavigne, D. M. (1987) Mass loss, metabolic rate, and energy utilization by harp and gray seal pups during the postweaning fast. Physiol. Zool. 60: 352–364.

    Google Scholar 

  • Wyrtki, K. (1982) The southern oscillation, ocean-atmosphere interaction and El Niño. Marine Technol. Soc. J. 16: 3–10.

    Google Scholar 

  • York, A. E. (1979) Analysis of pregnancy rates in female fur seals in the combined United States-Canada pelagic collections, 1950–1974, in Preliminary analysis of pelagic fur seal data collected by the United States and Canada during 1958–1974: 50–122 (H. Kajimura, R. H. Landers, M. A. Perez, A. E. York and M. A. Biggs, eds), Seattle, WA: National Marine Fisheries Service.

    Google Scholar 

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Wartzok, D. (1991). Physiology of behaviour in pinnipeds. In: Renouf, D. (eds) The Behaviour of Pinnipeds. Springer, Dordrecht. https://doi.org/10.1007/978-94-011-3100-1_5

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