The Behavior, Physiology, and Anatomy of Lactation in the Pinnipedia

  • Olav T. Oftedal
  • Daryl J. Boness
  • Raymond A. Tedman


Lactation represents a major proportion of maternal reproductive effort. In ungulates the energetic costs of lactation comprise 75–80% of the total energetic costs of reproduction (Oftedal, 1985). Although pinnipeds feed at sea, they return to land or ice to give birth and to nurse their young (Fig. 1) (Bonner, 1984). Thus, the bulk of a female’s reproductive expenditure is in an environment in which she cannot feed. The temporal separation of feeding and lactation is achieved by a wide range of behavioral, physiological, and anatomic adaptations.


Elephant Seal Grey Seal Weddell Seal Harp Seal Southern Elephant Seal 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Ackman, R. G., and Burgher, R. D., 1963, Component fatty acids of the milk of the grey (Atlantic) seal, Can. J. Biochem. Physiol. 41:2501–2505.PubMedGoogle Scholar
  2. Alcorn, D. J., 1984, The Hawaiian Monk Seal on Laysan Island: 1982, NOAA Technical Memorandum, National Marine Fisheries Service, Southwest Fisheries Center, No. 42, pp. 1-37.Google Scholar
  3. Amoroso, E. C, and Matthews, L. H., 1952, Reproduction and lactation in the seal, in: Report of the II International Congress of Physiology and Pathology of Animai Reproduction and of Artificial Insemination, Volume 1, Copenhagen, pp. 193-201.Google Scholar
  4. Amoroso, E. C., Goffin, A., Halley, G., Matthews, L. H., and Mathews, D. J., 1951, Lactation in the grey seal, J. Physiol. 113:4–5.Google Scholar
  5. Amoroso, E. C., Bourne, G. H., Harrison, R. L., Matthews, L. H., Rowlands, J. W., and Sloper, J. C., 1965, Reproductive and endocrine organs of foetal newborn and adult seals, J. Zool. Lond. 147:430–486.Google Scholar
  6. 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.Google Scholar
  7. 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.Google Scholar
  8. Anderson, S. S., Burton, R. W., and Summers, C. F., 1975, Behaviour of grey seals (Halichoerus grypus) during a breeding season on North Rona, J. Zool. Lond. 177: 179–195.Google Scholar
  9. Ashworth, U. S., Ramaiah, G. D., and Keyes, M. C, 1966, Species difference in the composition of milk with special reference to the northern für seal, J. Dairy Sci. 49: 1206–1211.Google Scholar
  10. Bartholomew, G. A., 1959, Mother-young relations and the maturation of pup behavior in the Alaska für seal, Anim. Behav. 7:163–171.Google Scholar
  11. Bartholomew, G. A., and Hoel, P. G., 1953, Reproductive behavior of the Alaska für seal, Callorhinus ursinus, J. Mammal. 34:417–437.Google Scholar
  12. Belkin, A. N., Kosygin, G. M., and Panin, K. I., 1969, New materials on island seals (Phoca insularis), in: Morskie Mlekopitayshchie (V. A. Arsen’ev, B. A. Zenkovich, and K. K. Chapskii, eds.), Nauka, Moscow, pp. 157-175 [in Russian].Google Scholar
  13. Belov, L., 1971, Histological organization of the mammary gland in the seals Pagophilus groenlandicus and Cystophora cristata, J. Evol. Biochem. Physiol. 7:585–589.Google Scholar
  14. Berger, J., 1979, Weaning conflict in desert and mountain bighorn sheep (Ovis canadensis): An ecological interpretation, Z. Tierpsychol. 50:188–200.Google Scholar
  15. Bertram, G. C. L., 1940, The biology of the Weddell and crabeater seals: With a study of the comparative behaviour of the Pinnipedia, in: Scientific Reports British Graham Land Expedition, 1934–37, Volume 1, pp. 1-139.Google Scholar
  16. Best, H. A., 1974, A Preliminary Report on the Natural History of Hooker’s Sea Lion at Enderby Island, New Zealand Ministry of Agriculture and Fisheries Report No. 132.Google Scholar
  17. Bester, M. N., 1977, Habitat selection, seasonal population changes, and behaviour of the Amsterdam für seal, Arctocephalus tropicalis, on Gough Island, Ph. D. thesis, University of Pretoria, Pretoria, South Africa.Google Scholar
  18. Bester, M. N., 1981, Seasonal changes in the population composition of the für seal, Arctocephalus tropicalis, at Gough Island, S. Afr. J. Wildl. Res. 11:49–55.Google Scholar
  19. Bigg, M. A., 1969, The Harbor Seal in British Columbia, Bull. Fish. Res. Board Can. 172: 1–33.Google Scholar
  20. Bigg, M. A., 1973, Census of the California sea lions on southern Vancouver Island, British Columbia, J. Mammal. 54:285–287.Google Scholar
  21. Bishop, S. C, 1967, Reproduction, age determination, and behavior of the harbor seal, Phoca vitulina L., in the Gulf of Alaska, M. Sc. Thesis, University of Alaska, Fairbanks.Google Scholar
  22. Bock, H.-D., Wunsche, J. Hoffmann, B., Linke, K., and Fricke, G., 1968, Über die Aminosäuren-Zusammensetzung der Milch der Kegelrobbe, Halichoerus grypus Fab. und des Schweinswals, Phocoena phocoena L., Zool. Gart. Hpz. 35:69–73.Google Scholar
  23. Boness, D. J., 1979, The social system of the grey seal, Halichoerus grypus (Fab.), on Sable Island, Nova Scotia, Ph. D. thesis, Dalhousie University, Halifax, Nova Scotia.Google Scholar
  24. Boness, D. J., and James, H., 1979, Reproductive behaviour of the grey seal (Halichoerus grypus) on Sable Island, Nova Scotia, J. Zool. Lond. 188:477–500.Google Scholar
  25. Boness, D. J., Anderson, S. S., and Cox, C. R., 1982, Functions of female aggression during the pupping and mating season of grey seals, Halichoerus grypus (Fabricius), Can. J. Zool. 60:2270–2278.Google Scholar
  26. Boness, D. J., Dabek, L, Ono, K. A., and Oftedal, O. T., 1985a, Female attendance behavior in California sea lions, in: Abstracts Sixth Biennial Conference on the Biology of Marine Mammals, Vancouver, British Columbia.Google Scholar
  27. Boness, D. J., Oftedal, O. T., Ono, K. A., and Dabek, L., 1985b, Observations on yearling California sea lions at breeding beaches on San Nicolas Island, California, in: Abstracts Sixth Biennial Conference on the Biology of Marine Mammals, Vancouver, British Columbia.Google Scholar
  28. Bonner, W. N., 1968, The Fur Seal of South Georgia, British Antarctic Survey Scientific Report No. 56, pp. 1-81.Google Scholar
  29. Bonner, W. N., 1979a, Largha seal, FAO Fish. Ser. No. 5 1979:63–66.Google Scholar
  30. Bonner, W. N., 1979b, Harbour seal, FAO Fish. Ser. No. 5 1979:58–63.Google Scholar
  31. Bonner, W. N., 1984, Lactation strategies in pinnipeds: Problems for a marine mammalian group, Symp. Zool. Soc. Lond. 51:253–272.Google Scholar
  32. Bonner, W. N., and Witthames, S. R., 1974, Dispersal of common seals (Phoca vitulina), tagged in the Wash, East Anglia, J. Zool. Lond. 174:528–531.Google Scholar
  33. Boulva, J., 1979, Mediterranean monk seal, FAO Fish. Ser. No. 5 1979:95–101.Google Scholar
  34. Boulva, J., and McLaren, I. A., 1979, Biology of the Harbor Seal, Phoca vitulina, in Eastern Canada, Bull Fish. Res. Board Can. 200:1–24.Google Scholar
  35. Bowen, W. D., and Sergeant, D. E., 1983, Mark-recapture estimates of harp seal pup (Phoca groenlandica) production in the northwest Atlantic, Can. J. Fish. Aquat. Sci. 40:728–742.Google Scholar
  36. Bowen, W. D., Oftedal, O. T., and Boness, D. J., 1985, Birth to weaning in four days: Remarkable growth in the hooded seal, Cystophora cristata, Can. J. Zool. 63:2841–2846.Google Scholar
  37. Bowen, W. D., Myers, R. A., and Hay, K., 1986a, Abundance of a dispersed, dynamic population: Hooded seals (Cystophora cristata) in the Northwest Atlantic, Can. J. Fish. Aquat. Sci., in press.Google Scholar
  38. Bowen, W. D., Boness, D. J. and Oftedal, O. T., 1986b, Mass transfer from mother to pup and subsequent mass loss by the weaned pup in the hooded seal, Cystophora cristata, Can. J. Zool, in press.Google Scholar
  39. Brambell, F. W. R., 1970, The Transmission of Passive Immunity from Mother to Young, North-Holland, Amsterdam.Google Scholar
  40. Brodie, P., and Beck, B., 1983, Predation by sharks on the grey seal (Halichoerus grypus) in Eastern Canada, Can. J. Fish. Aquat. Sci. 40:267–271.Google Scholar
  41. Bryden, M. M., 1968, Lactation and suckling in relation to early growth of the southern elephant seal, Aust. J. Zool. 16:739–747.Google Scholar
  42. Bryden, M. M., and Stokes, G. B., 1969, Metabolism of fatty acids in the southern elephant seal, Mirounga Jeonina L., Can. J. Biochem. 47:757–760.PubMedGoogle Scholar
  43. Bryden, M. M., and Tedman, R. A., 1974, Studies of lactation in Weddell seals, Antarct. J. U. S. 9:105–106.Google Scholar
  44. Bunt, J. S., 1968, Microalgae of the Antarctic pack ice zone, in: Symposium on Antarctic Oceanography, Scott Polar Research Institute, Cambridge, England, pp. 198-218.Google Scholar
  45. Burns, J. J., 1970, Remarks on the distribution and natural history of pagophilic pinnipeds in the Bering and Chukchi seas, J. Mammal. 51:445–454.Google Scholar
  46. Burns, J. J., 1981a, Bearded seal—Erignathus barbatus, in: Handbook of Marine Mammals (S. H. Ridgway and R. J. Harrison, eds.), Academic Press, New York, Volume 2, pp. 145–170.Google Scholar
  47. Burns, J. J., 1981b, Ribbon seal—Phoca fasciata, in: Handbook of Marine Mammals (S. H. Ridgway and R. J. Harrison, eds.), Academic Press, New York, Volume 2, pp. 89–109.Google Scholar
  48. Burns, J. J., and Frost, K. J., 1979, The Natural History and Ecology of the Bearded Seal, Erignathus barbatus, Final Report, Alaska Fish and Game, Contract #02-5-022-53, pp. 1-77.Google Scholar
  49. Burns, J. J., Ray, G. C., Fay, F. H., and Shaughnessy, P., 1972, Adoption of a strange pup by the ice-inhabiting harbor seal, Phoca vitulina Jargha, J. Mammal. 53:594–598.Google Scholar
  50. Burton, R. W., Summers, C. F., and Anderson, S. S., 1975, Perinatal activities in the grey seal (Haiichoerus grupus), J. Zool. Lond. 177:197–201.Google Scholar
  51. Campagna, C., 1985, The breeding cycle of the southern sea lion, Otaria byronia, Mar. Mammal Sci. 1:210–219.Google Scholar
  52. Carnio, J., 1982, Observations on the mother and young interactions in captive Cape für seals (Arctocephalus pusillus), Aquat. Mamm. 9:50–56.Google Scholar
  53. Carrick, R., Csordas, S. E., Ingham, S. E., and Keith, K., 1962, Studies on the southern elephant seal, Mirounga Jeonina. 3. The annual cycle in relation to age and sex, CSIRO Wildl. Res. 7:119–160.Google Scholar
  54. Christenson, T. E., and LeBoeuf, B. J., 1978, Aggression in the female northern elephant seal, Mirounga angustirostris, Behaviour 64:158–171.PubMedGoogle Scholar
  55. Cody, M. M., 1984, Substances without vitamin status, in: Handbook of Vitamins. Nutritional, Clinical, and Biochemical Aspects (L. J. Machlin, ed.), Marcel Dekker, New York, pp. 571–585.Google Scholar
  56. Condit, R., and LeBoeuf, B. J., 1984, Feeding habits and feeding grounds of the northern elephant seal, J. Mammal. 65:281–291.Google Scholar
  57. Condy, P. R., 1979, Annual cycle of the southern elephant seal, Mirounga Jeonina (Linn.) at Marion Island, S. Afr. J. Zool. 14:95–102.Google Scholar
  58. Condy, P. R., 1980, Postnatal development and growth in southern elephant seals, Mirounga Jeonina, at Marion Island, S. Afr. J. Wildl. Res. 8:159–168.Google Scholar
  59. Cook, H. W., and Baker, B. E., 1969, Seal milk. I. Harp seal (Pagophilis groenlandicus) milk: Composition and pesticide residue content, Can. J. Zool. 47:1129–1132.PubMedGoogle Scholar
  60. Costa, D. P., and Gentry, R. L., 1986, Free-ranging energetics of northern für seals, in: Fur Seals: Maternal Strategies on Land and at Sea (R. L. Gentry and G. L. Kooyman, eds.), Princeton University Press, Princeton, pp. 79–101.Google Scholar
  61. Costa, D. P., Thorson, P., Feldkamp, S., Gentry, R. L., DeLong, R., Antonelis, G., and. Croxall, J. P., 1985, At sea foraging energetics of three species of pinnipeds, Fed. Proc. 44:1000.Google Scholar
  62. Costa, D. P., LeBoeuf, B. J., Huntley, A. C., and Ortiz, C. L., 1986, The energetics of lactation in the northern elephant seal, J. Zool. Lond. 209:21–33.Google Scholar
  63. Coulson, J. C., and Hickling, G., 1964, The breeding biology of the grey seal, Halichoerus grypus (Fab.) on the Farne Islands, Northumberland, J. Anim. Ecol. 33:483–512.Google Scholar
  64. Cowie, A. T., and Tindal, J. S., 1971, The Physiology of Lactation, Edward Arnold, London.Google Scholar
  65. Cross, B. A., 1977, Comparative physiology of milk removal, Symp. Zool. Soc. Lond. 41: 193–210.Google Scholar
  66. Croxall, J. P., Everson, I., Kooyman, G. L., Ricketts, C., and Davis, R. W., 1985, Fur seal diving behavior in relation to vertical distribution of krill, J. Anim. Ecol. 54:1–8.Google Scholar
  67. Curry-Lindahl, K., 1975, Ecology and conservation of the grey seal, Halichoerus grypus, common seal, Phoca vitulina, and ringed seal, Phoca hispida in the Baltic Sea, Rapp. P. V. Reun. Cons. Int. Explor. Mer. 169:527–532.Google Scholar
  68. David, J. H. M., and Rand, R. W., 1986, Attendance behavior of female South African für seals, in: Fur Seals: Maternal Strategies on Land and at Sea, (R. L. Gentry and G. L. Kooyman, eds.), Princeton University Press, Princeton, pp. 126–141.Google Scholar
  69. Davies, J. L., 1949, Observations of the grey seal (Halichoerus grypus) at Ramsey Island, Pembrokeshire, Proc. Zool. Soc. Lond. 119:673–692.Google Scholar
  70. Demaster, D. P., 1979, Weddell seal, FAO Fish. Ser. No. 5 1979:130–134.Google Scholar
  71. Doidge, D. W., McCann, T. S., and Croxall, J. P., 1986, Attendance behavior of Antarctic für seals, in: Fur Seals: Maternal Strategies on Land and at Sea (R. L. Gentry and G. L. Kooyman, eds.), Princeton University Press, Princeton, pp. 102–114.Google Scholar
  72. Dosako, S., Taneya, S., Kimura, T., Ohmori, T., Daikoku, H., Suzuki, N., Sawa, J., Kano, K., and Katayama, S., 1983, Milk of northern für seal: Composition, especially carbohydrate and protein, J. Dairy Sci. 66:2076–2083.PubMedGoogle Scholar
  73. Drewett, R. F., 1983, Sucking, milk synthesis, and milk ejection in the Norway rat, in: Parental Behaviour of Rodents (R. W. Elwood, ed.), Wiley, London, pp. 181–203.Google Scholar
  74. Dylewski, D. P., and Keenan, T. W., 1983, Compound exocytosis of casein micelles in mammary epithelial cells, Eur. J. Cell. Biol. 31:114–124.PubMedGoogle Scholar
  75. Edie, A. G., 1977, Distribution and movements of steller sea lion cows (Eumetopias jubata) on a pupping colony, M. Sc. Thesis, University of British Columbia, Vancouver, British Columbia.Google Scholar
  76. FAO, 1982, Mammals in the Seas, FAO Fisheries Ser. No. 5, Volume 4.Google Scholar
  77. Fay, F. H., 1982, Ecology and Biology of the Pacific Walrus, Odohenus rosmarus divergens Illiger, U. S. Fish and Wildlife Service, North American Fauna, No. 74, pp. 1-279.Google Scholar
  78. Fedak, M. A., and Anderson, S. S., 1982, The energetics of lactation: Accurate measurements from a large wild mammal, J. Zool. Lond. 198:473–479.Google Scholar
  79. Fiscus, F. H., and Baines, G. A., 1966, Food and feeding behavior of Steller and California sea lions, J. Mammal. 47:195–200.Google Scholar
  80. Fogden, S. C. L., 1971, Mother—young behaviour at grey seal breeding beaches, J. Zool. Lond. 164:61–92.Google Scholar
  81. Freed, L. M., York, C. M., Hamosh, M., Sturman, J. T., Oftedal, O. T., and Hamosh, P., 1986, Bile salt stimulated lipase: The enzyme is present in nonprimate milk, in: Human Lactation, Volume 2: Maternal-Environmental Factors (M. Hamosh and A. S. Goldman, eds.), Plenum Press, New York, pp. 595–601.Google Scholar
  82. Frost, K. J., and Lowery, L. F., 1980, Feeding of ribbon seals (Phoca fasciata) in the Bering Sea in spring, Can. J. Zool. 58:1601–1607.Google Scholar
  83. Gentry, R. L., 1970, Social behavior of Steller sea lions, Ph. D. thesis, University of California, Santa Cruz.Google Scholar
  84. Gentry, R. L., and Holt, J. R., 1986, Attendance behavior of northern für seals, in: Fur Seals: Maternal Strategies on Land and at Sea (R. L. Gentry and G. L. Kooyman, eds.), Princeton University Press, Princeton, pp. 41–60.Google Scholar
  85. 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., 1986, Synthesis and conclusions, in: Fur Seals: Maternai Strategies on Land and at Sea (R. L. Gentry and G. L. Kooyman, eds.), Princeton University Press, Princeton, pp. 220–278.Google Scholar
  86. Glass, R. L., and Jenness, R., 1971, Comparative biochemical studies of milks—VI. Constituent fatty acids of milk fats of additional species, Comp. Biochem. Physiol. 38B: 353–359.Google Scholar
  87. Glass, R. L., Troolin, H. A., and Jenness, R., 1967, Comparative biochemical studies of milks—IV. Constituent fatty acids of milk fats, Comp. Biochem. Physiol. 22:415–425.PubMedGoogle Scholar
  88. Golt’sev, V. N., 1971, Feeding of the harbor seal, Ekologiya 2:62–70.Google Scholar
  89. Golt’sev, V. N., and Fedoseev, G. A., 1970, Dynamics of the age composition of rookeries and the replacement capacity of harbor seal populations, TINRO 71:309-317 [Fisheries Research Board of Canada Translation Series No. 2401 (1974)].Google Scholar
  90. Hall, W. G., 1983, Suckling isn’t feeding, or is it? A search for developmental continuities, Adv. Stud. Behav. 13:219–254.Google Scholar
  91. Hamilton, J. E., 1934, The southern sea lion, Otaria hyronia, Discovery Rep. 8:269–318.Google Scholar
  92. Hanwell, A., and Peaker, M., 1977, Physiological effects of lactation on the mother, Symp. Zool. Soc. Lond. 41:297–312.Google Scholar
  93. Harrison, R. J., 1960, Reproduction and reproductive organs in common seals (Phoca vitulina) in the Wash, East Anglia, Mammalia 24:372–385.Google Scholar
  94. Harrison, R. J., 1969, Reproduction and reproductive organs, in Biology of Marine Mammals (H. T. Anderson, ed.), Academic Press, New York, pp. 253–348.Google Scholar
  95. Harrison, S. H., and Ridgway, R. J. (eds.), 1981, Handbook of Marine Mammals, Volumes 1 and 2, Academic Press, London.Google Scholar
  96. Hewer, H. R., 1960, Behaviour of the grey seal (Halichoerus grypus) in the breeding season, Mammalia 24:401–420.Google Scholar
  97. Hewer, H. R., 1974, British Seals, Collins, London.Google Scholar
  98. Higgins, L. V., 1984, Material behavior and attendance patterns of the Steller sea lion in California, M. Sc. thesis, University of California, Santa Cruz.Google Scholar
  99. Hill, K. J., Noakes, D. E., and Lowe, R. E., 1970, Gastric digestive physiology of the calf and piglet, in: Physiology of Digestion and Metabolism in the Ruminant (A. T. Phillipson, ed.), Oriel Press, Newcastle, pp. 166–179.Google Scholar
  100. Hollmann, K. H., 1974, Cytology and fine structure of the mammary gland, in: Lactation: A Comprehensive Treatise, Volume 1 (B. L. Larson and V. R. Smith, eds.), Academic Press, London, pp. 3–95.Google Scholar
  101. Holt, C., 1983, Swelling of Golgi vesicles in mammary secretory cells and its relation to the yield and quantitative composition of milk, J. Theor. Biol. 101:247–261.PubMedGoogle Scholar
  102. Hook, O., and Johnels, A. G., 1972, The breeding and distribution of the grey seal (Halichoerus grypus Fab.) in the Baltic Sea, with observations on other seals of the area, Proc. R. Soc. Ser. B 182:37–58.Google Scholar
  103. Ivashin, M. V., Popov, L. A., and Tsapko, A. S., 1972, Morskie Mlekopitayushchie, Pishchevaia Promyshlennost, Moscow [Fisheries Research Board of Canada, Translation, Series No. 2783 (1973)].Google Scholar
  104. Jangaard, P. M., and Ke, P. J., 1968, Principal fatty acids of depot fat and milk lipids from harp seal (Pagophilis groenlandius) and hooded seal (Cystophora cristata), J. Fish. Res. Board Can. 25:2419–2426.Google Scholar
  105. Jenness, R., 1979, Comparative aspects of milk proteins, J. Dairy Res. 46:197–210.PubMedGoogle Scholar
  106. Jenness, R., and Sloan, R. E., 1970, The composition of milks of various species: A review, Dairy Sci. Abstr. 32:599–612.Google Scholar
  107. Jenness, R., Erickson, A. W., and Craighead, J. J., 1972, Some comparative aspects of milk from four species of bears, J. Mammal. 53:34–47.PubMedGoogle Scholar
  108. Johnson, J. D., 1975, Comparative aspects of lactose digestion and synthesis in mammals — The California sea lion, Mod. Probl. Paediatr. 15:194–200.Google Scholar
  109. Johnson, J. D., Christiansen, R. O., and Kretchmer, N., 1972, Lactose synthetase in mammary gland of the California sea lion, Biochem. Biophys. Res. Commun. 47:393–397.PubMedGoogle Scholar
  110. Kaufman, G. W., Siniff, D. B., and Reichle, R., 1975, Colony behavior of Weddell seals, Leptonychotes weddelli, at Hutton Cliffs, Antarctica, Rapp. P. V. Reun. Cons. Int. Explor. Mer. 169:228–246.Google Scholar
  111. Keenan, T. W., Franke, W. W., Mather, I. H., and Moore, D. J., 1978, Endomembrane composition and function in milk formation, in: Lactation: A Comprehensive Treatise, Volume 4 (B. L. Larson, ed.), Academic Press, London, pp. 405–436.Google Scholar
  112. Kenyon, K. W., and Rice, D. W., 1959, Life history of the Hawaiian monk seal, Pac. Sci. 13:215–252.Google Scholar
  113. Kerley, G. I. H., 1983, Comparison of seasonal haul-out patterns of für seals, Arctocephalus tropicalis and A. gazella, on subantarctic Marion Island, S. Afr. J. Wildl. Res. 13:71–77.Google Scholar
  114. Kerry, K. R., and Messer, M., 1968, Intestinal glycosidases of three species of seals, Comp. Biochem. Physiol. 25:437–446.PubMedGoogle Scholar
  115. King, J. E., 1983, Seals of the World, 2nd ed., Oxford University Press, Oxford.Google Scholar
  116. Knox, G. A., 1970, Antarctic marine ecosystems, in: Antarctic Ecology, Volume 1 (N. W. Holdgate, ed.), Academic Press, London, pp. 69–97.Google Scholar
  117. Kooyman, G. L., and Drabek, C. M., 1968, Observations on the milk, blood, and urine constituents of the Weddell seal, Physiol. Zool. 41:187–194.Google Scholar
  118. Kormondy, E. J., 1969, Concepts of Ecology, Prentice-Hall, Englewood Cliffs, New Jersey.Google Scholar
  119. Kovacs, K., 1986, Maternal investment and early behavioural development in harp seals (Phoca groenlandica) and grey seals (Halichoerus grypus), Ph. D. thesis, Guelph University, Guelph, Ontario.Google Scholar
  120. Kovacs, K., and Lavigne, D. M., 1986, Maternal investment and neonatal growth in phocid seals, J. Anim. Ecol. 55:(in press).Google Scholar
  121. Lander, R. H., 1979, Alaskan or northern für seal, EAO Fish. Ser. No. 5 1979:19–24.Google Scholar
  122. Lander, R. H., and Kajimura, H., 1982, Status of northern für seals, FAO Fish. Ser. No. 5 1979:319–347.Google Scholar
  123. Lascelles, A. K., and Lee, C. S., 1978, Involution of the mammary gland, in: Lactation: A Comprehensive Treatise, Volume 4 (B. L. Larson, ed.), Academic Press, London, pp. 115–177.Google Scholar
  124. Lauer, B. H., and Baker, B. E., 1977, Amino acid composition of casein isolated from the milk of different species, Can. J. Zool. 55:231–236.PubMedGoogle Scholar
  125. Lavigne, D. M., 1976, Counting harp seals with ultraviolet photography, Polar Record 18:269–272.Google Scholar
  126. Lavigne, D. M., Stewart, R. E. A., and Fletcher, F., 1982, Changes in composition and energy content of harp seal milk during lactation, Physiol. Zool. 55:1–9.Google Scholar
  127. Laws, R. M., 1956, The Elephant Seal (Mirounga leonina Linn). II General, Social, and Reproductive Behavior, Falkland Islands Dependency Survey, Scientific Report No. 13, pp. 1-88.Google Scholar
  128. Laws, R. M., 1958, Growth rates and ages of crabeater seals, Lobodon carcinophagus, Ptoc. Zool. Soc. Lond. 130:275–288.Google Scholar
  129. Laws, R. M., 1959, Accelerated growth in seals, with special reference to the Phocidae, Nor. Hvalfangst-Tid. 48:425–452.Google Scholar
  130. LeBoeuf, B. J., 1974, Male-male competition and reproductive success in elephant seals, Am. Zool. 14:163–176.Google Scholar
  131. LeBoeuf, B. J., and Ortiz, C. L., 1977, Composition of elephant seal milk, J. Mammal. 58: 683–685.Google Scholar
  132. LeBoeuf, B. J., and Panken, K. J., 1977, Elephant seals breeding on the mainland in California, Proc. Calif. Acad. Sci. Ser. 4 41:267–280.Google Scholar
  133. LeBoeuf, B. J., Whiting, R. J., and Gantt, R. F., 1972, Perinatal behavior of northern elephant seal females and their young, Behaviour 43:121–156.Google Scholar
  134. Lincoln, D. W., and Paisley, A. C., 1982, Neuroendocrine control of milk ejection, J. Reprod. Fertil. 65:571–586.PubMedGoogle Scholar
  135. Lindsey, A. A., 1937, The Weddell seal in the Bay of Whales, Antarctica, J. Mammal. 18:127–144.Google Scholar
  136. Ling, J. K., and Walker, G. E., 1976, Seal studies in South Australia: Progress report for the year 1975, S. Aust. Nat. 50:59–72.Google Scholar
  137. Linzell, J. L., Peaker, M., and Taylor, J. C., 1975, The effects of prolactin and oxytocin on milk secretion and on the permeability of the mammary epithelium in the rabbit, J. Physiol. 253:547–563.PubMedGoogle Scholar
  138. Loughrey, A. G., 1959, Preliminary investigation of the Atlantic walrus, Odobenus rosmarus rosmarus, Can. Wildl. Serv. Bull. Ser. No. 14 1959:1–23.Google Scholar
  139. Macy, S. K., 1982, Mother-pup interactions in the northern für seal, Ph. D. thesis, University of Washington, Seattle.Google Scholar
  140. Mansfield, A. W., 1958, The biology of the Atlantic walrus, Odobenus rosmarus rosmarus (Linnaeus), in the Eastern Canadian Arctic, Fish. Res. Board Can. Manuscr. Rep. Ser. No. 653 1958:1–146.Google Scholar
  141. Mansfield, A. W., 1967, Seals of Arctic and Eastern Canada, 2nd ed., Bull. Fish. Res. Board Can. 137:1–30.Google Scholar
  142. Mansfield, A. W., and Beck, B., 1977, The grey seal in Eastern Canada, Can. Fish. Mar. Serv. Tech. Rep. 704:1–81.Google Scholar
  143. Marlow, B. J., 1975, The comparative behaviour of the Australian sea lions, Neophoca cinerea and Phocartos hookeri, Mammalia 39:159–230.Google Scholar
  144. Mate, B. R., and Gentry, R. L., 1979, Northern (Steller) sea lion, FAO Fish. Ser. No. 5 1979:1–5.Google Scholar
  145. Mather, I. H., and Keenan, T. W., 1983, Function of endomembranes and the cell surface in the secretion of organic milk constituents, in: Biochemistry of Lactation (T. B. Mepham, ed.), Elsevier, New York, pp. 231–283.Google Scholar
  146. Maule Walker, F. M., 1984, Exocrine and endocrine secretions of the mammary gland: Local control in ruminants, Symp. Zool. Soc. Lond. 51:171–191.Google Scholar
  147. McCann, T. S., 1982, Aggressive and maternal activities of female southern elephant seals (Mirounga leonina), Anim. Behav. 30:268–276.Google Scholar
  148. McLaren, I. A., 1958, The Biology of the ringed seal (Phoca groenlandica Schreber) in the Eastern Canadian Arctic, Bull. Fish. Res. Board Can. 118:1–97.Google Scholar
  149. McLaren, I. A., 1960, On the origin of the Caspian and Baikal seals and the paleoclimatological implications, Am. J. Sci. 258:35–50.Google Scholar
  150. McNab, A. G., and Crawley, M. C, 1975, Mother and pup behaviour of the New Zealand für seal, Arctocephalus forsten (Lesson), Mauri Ora 3:77–88.Google Scholar
  151. Meara, M. L., 1952, The component fatty acids of the milk fat of a grey Atlantic seal, Biochem. J. 51:190–193.PubMedGoogle Scholar
  152. Miller, E. H., 1971, Social and thermoregulatory behaviour of the New Zealand für seal (Arctocephaius forsten), M. Sc. thesis, University of Cantebury, Christchurch.Google Scholar
  153. Miller, E. H., 1975, Annual cycle of für seals, Arctocephalus forsten (Lesson), on the Open Bay Islands, New Zealand, Pac. Sci. 29:139–152.Google Scholar
  154. Miller, E. H., and Boness, D. J., 1983, Summer behaviour of the Atlantic walrus (Odobenus rosmarus rosmarus) on Coats Island, NWT, Canada, Z. Säugetierkd. 48:298–313.Google Scholar
  155. Murie, J., 1874, Researches upon the anatomy of the Pinnipedia, Part II, Descriptive anatomy of the sea-lion (Otaria jubata), Trans. Zool. Soc. Lond. 7:527–596.Google Scholar
  156. Naito, Y., and Nishiwaki, M., 1972, The growth of two species of the harbour seal in the adjacent waters of Hokkaido, Whaie Res. Inst. Sci. Rep. 24:127–144.Google Scholar
  157. Oftedal, O. T., 1984a, Milk composition, milk yield, and energy output at peak lactation: A comparative review, Symp. Zool. Soc. Lond. 51:33–85.Google Scholar
  158. Oftedal, O. T., 1984b, Body size and reproductive strategy as correlates of milk energy output in lactating mammals, Acta Zool. Fenn. 171:183–186.Google Scholar
  159. Oftedal, O. T., 1985, Pregnancy and lactation, in: The Bioenergetics of Wild Herbivores (R. J. Hudson and R. G. White, eds.), CRC Press, Boca Raton, Florida, pp. 215–238.Google Scholar
  160. Oftedal, O. T., Boness, D. J., and Iverson, S. J., 1983, The effect of sampling method and stage of lactation on the composition of California sea lion milk, in: Abstracts Fifth Biennial Conference on the Biology of Marine Mammals, Boston, Massachusetts, p. 73.Google Scholar
  161. Oftedal, O. T., Iverson, S. J., and Boness, D. J., 1985, Energy intake in relation to growth rate of the California sea lion, in: Abstracts Sixth Biennial Conference on the Biology of Marine Mammals, Vancouver, British Columbia.Google Scholar
  162. Ono, K. A., Boness, D. J., Oftedal, O. T., and Iverson, S. J., 1985, Effects of the 1983 El Niño on the mother-pup relationship in the California sea lion, Abstracts Sixth Biennial Conference on the Biology of Marine Mammals, Vancouver, British Columbia.Google Scholar
  163. Ortiz, C. L., Costa, D. P., and LeBoeuf, B. J., 1978, Water and energy flux in elephant seals pups fasting under natural conditions, Physiol. Zool. 238:166–178.Google Scholar
  164. Ortiz, C. L., LeBoeuf, B. J., and Costa, D. P., 1984, Milk intake of elephant seal pups: An index of parental investment, Am. Nat. 124:416–422.Google Scholar
  165. Pastukhov, V. N., 1975a, Birth time and duration of pupping period of Baikal seals, Morsk. Mlekopitayushchiye 4:41–43.Google Scholar
  166. Pastukhov, V. N., 1975b, Number and distribution of the postparturent females of the Baikal seal, Morsk. Mlekopitayushchiye 4:39–41.Google Scholar
  167. Payne, M. R., 1979, Growth in the Antarctic für seal, Arctocephaius gazella, J. Zool. Lond. 187:1–20.Google Scholar
  168. Payne, P. R., and Wheeler, E. F., 1968, Comparative nutrition in pregnancy and lactation, Proc. Nutr. Soc. 27:129–138.PubMedGoogle Scholar
  169. Peaker, M., 1977, The aqueous phase of milk: Ion and water transport, Symp. Zool. Soc. Lond. 41:113–134.Google Scholar
  170. Peaker, M., and Goode, J. A., 1978, The milk of the für seal Arctocephaius tropicalis gazella, in particular the composition of the aqueous phase, J. Zool. Lond. 185: 469–476.Google Scholar
  171. Peterson, R. S., 1965, Behavior of the northern für seal, D. Sc. thesis, Johns Hopkins University, Baltimore, Maryland.Google Scholar
  172. Peterson, R. S., 1968, Social behavior in pinnipeds with particular reference to the northern für seal, in: Behavior and Physiology of Pinnipeds, (R. J. Harrison, R. C. Hubbard, R. C. Peterson, D. E. Rice, and R. J. Schusterman, eds.), Appleton-Century-Crofts, New York, pp. 1-53.Google Scholar
  173. Peterson, R. S., and Bartholomew, G. A., 1967, The Natural History and Behavior of the California Sea Lion, American Society of Mammalogists.Google Scholar
  174. Pierotti, R., and Pierotti, D., 1980, Effects of cold on the evolution of pinniped breeding systems, Evolution 34:494–507.Google Scholar
  175. Pierson, M. O., 1978, Breeding behavior of the Guadalupe für seal, ArctocephaJus townsendi, Ph. D. thesis, University of California, Santa Cruz.Google Scholar
  176. Pilson, M. E. Q., and Kelly, A. L., 1962, Composition of the milk from Zalophus californianus, the California sea lion, Science 135:104–105.PubMedGoogle Scholar
  177. Pitcher, K. W., and Calkins, D. G., 1981, Reproductive biology of Steller sea lions in the Gulf of Alaska, J. Mammal. 62:599–605.Google Scholar
  178. Ponce de Leon, A., 1984, Lactancia y Composicion Cuantitativa de la Leche del Lobo Fino Sudamericano, Arctocephalus australis (Zimmerman, 1783), Industria Lobera Pesquera Estado, Montevideo, Uruguay, Anales 1984:43–58.Google Scholar
  179. Popov, L. A., 1979, Caspian seal, FAO Fish. Ser. No. 5 1979:63–66.Google Scholar
  180. Popov, L. A., and Krylov, V. I., 1978, On the period of reproduction in the Weddell seal (Leptonychotes weddelli) on the King Georges Island (Waterloo, Antarctica), Zool. Zh. 57:1250–1255.Google Scholar
  181. Poulter, T. C., Pinney, T. C., Jennings, R., and Hubbard, R. C., 1965, The rearing of Steller sea lions, in: Proceedings Second Annual Conference Biological Sonar and Diving Mammals (C. E. Rice, ed.), Stanford Research Institute, Fremont, Calif., pp. 49–62.Google Scholar
  182. Pumphrey, R. S. H., 1977, A comparative study of plasma cells in the mammary gland in pregnancy and lactation, Symp. Zool. Soc. Lond. 41:261–276.Google Scholar
  183. Rand, R. W., 1955, Reproduction in the Cape für seal, Arctocephalus pusillus (Schreber), Proc. Zool. Soc. Lond. 124:717–740.Google Scholar
  184. Rand, R. W., 1959, The Cape für seal (ArctocephaJus pusiJJus). 2. Distribution, abundance and feeding habits off the southwestern coast of the Cape Province, S. Afr. Dept. Comm. Ind. Div. Fish. Invt. Rep. 34:1–65.Google Scholar
  185. Ray, C. E., 1976, Geography of phocid evolution, Syst. Zool. 25:391–406.Google Scholar
  186. Reinhardt, V., and Reinhardt, A., 1981, Natural sucking performance and age of weaning in Zebu cattle (Bos indicus), J. Agric. Sci. Camh. 96:309–312.Google Scholar
  187. Reiter, J., Stinson, N. L., and LeBoeuf, B. J., 1978, Northern elephant seal development: The transition from weaning to nutritional independence, Behav. Ecol. Sociohiol. 3:337–367.Google Scholar
  188. 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. Lond. 202:583–590.Google Scholar
  189. Renouf, D., and Diemand, D., 1984, Behavioural interactions between harbour seal mothers and pups during weaning (Pinnipedia: Phocidae), Mammalia 48:53–58.Google Scholar
  190. Renouf, D., Gaborko, L., Galway, G., and Finlayson, R., 1981, The effects of disturbance on the daily movements of harbour seals and grey seals between the sea and their hauling grounds at Miquelon, Appl. Anim. Ethol. 7:373–379.Google Scholar
  191. Repenning, C. A., 1976, Adaptive evolution of sea lions and walruses, Syst. Zool. 25: 375–391.Google Scholar
  192. Repenning, C. A., Ray, C. E., and Grigorescu, D., 1979, Pinniped biogeography, in: Historical Biogeography, Plate Tectonics and the Changing Environment (J. Grey and A. J. Boucot, eds.), Oregon State University Press, Corvallis, pp. 357–369.Google Scholar
  193. Riedman, M., and Ortiz, C. L., 1979, Changes in milk composition during lactation in the northern elephant seal, Physiol. Zool. 52:240–249.Google Scholar
  194. Roux, J. P., and Hess, A. D., 1984, The seasonal haul-out of the für seal, Arctocephalus tropicalis (Gray, 1872) on Amsterdam Island, Mammalia 48:377–389.Google Scholar
  195. Salter, R., 1978, Normal Behavior and Disturbance of Walruses (Odobenus rosmarus L.) during Terrestrial Haul-out, Eastern Bathurst Island, N.W.T., Report to Polar Gas Environmental Program, LGL Ltd, Toronto, Ontario.Google Scholar
  196. Sandegren, F. E., 1970, Breeding and maternal behavior of the Steller sea lion, M. Sc. thesis, University of Alaska, Fairbanks.Google Scholar
  197. Scheffer, V. B., 1962, Pelage and Surface Topography of the Northern Fur Seal, U. S. Fish and Wildlife Service, North American Fauna, No. 64, pp. 1-206.Google Scholar
  198. Scheffer, V. B., 1970, The Year of the Seal, Scribner’s, New York.Google Scholar
  199. Schmidt, D. V., Walker, L. E., and Ebner, K. E., 1971, Lactose synthetase activity in northern für seal milk, Biochim. Biophys. Acta 252:439–442.PubMedGoogle Scholar
  200. Schroeder, C. R., and Wegeforth, H. M., 1935, The occurrence of gastric ulcers in sea mammals of the California coast, their etiology and pathology, J. Am. Vet. Med. Assoc. 87:333–342.Google Scholar
  201. Scolaro, J. A., 1976, Censo de Elefantes Marinos (Mirounga Jeonina) en el Territorio Continental Argentino, Cent. Nac. Patagonica, Argentine Inf. Tec. 1:1–12.Google Scholar
  202. Shaughnessy, P. D., 1974, An electrophoretic study of blood and milk proteins of the southern elephant seal, Mirounga leonina, J. Mammal. 55:796–808.PubMedGoogle Scholar
  203. Shepeleva, V. K., 1971, Adaptations of seals to life in the Arctic, in: Morphology and Ecology of Marine Mammals (K. K. Chapskii and V. E. Sokolov, eds.), Wiley, New York, pp. 1–58.Google Scholar
  204. Shergin, A. L., Shergina, V. P., and Beloborofov, A. G., 1969, Composition of hooded seals’ milk, Dairy Sci. Abstr. 31:1086.Google Scholar
  205. Sinha, A. A., and Erickson, A. W., 1974, Ultrastructure of the placenta of Antarctic seals during the first third of pregnancy, Am. J. Anat. 141:263–280.PubMedGoogle Scholar
  206. Siniff, D. B., Stirling, I., Bengston, J. L., and Reichle, R. A., 1979, Social and reproductive behavior of crabeater seals during the austral spring, Can. J. Zool. 57:2243–2255.Google Scholar
  207. Sivertsen, E., 1941, On the biology of the harp seal, Phoca groenlandica, Erxl. Hvalraadets Skr. 26:1–164.Google Scholar
  208. Smith, E. A., and Burton, R. W., 1970, Weddell seals of Signey Island, in: Antarctic Ecology, Volume 1 (N. W. Holdgate, ed.), Academic Press, London, pp. 415–428.Google Scholar
  209. Smith, T. G., 1973, Population dynamics of the ringed seal, Phoca hispida, in the Canadian Eastern Arctic, Bull. Fish. Res. Board Can. 181:1–55.Google Scholar
  210. Smith, T. G., and Hammil, M. O., 1981, Ecology of the ringed seal, Phoca hispida, in its fast ice breeding habitat, Can. J. Zool. 59:966–981.Google Scholar
  211. Smith, T. G., and Stirling, I., 1978, Variation in the density of ringed seal (Phoca hispida) birth lairs in the Amundsen Gulf, NWT, Can. J. Zool. 56:1066–1070.Google Scholar
  212. Stewart, B. S., and Yochem, P., 1984, Seasonal abundance of pinnipeds at San Nicolas Island, California, 1980–1982, Bull. S. Calif. Acad. Sci. 83:121–132.Google Scholar
  213. Stewart, R. E. A., 1983, Behavioral and energetic aspects of reproductive effort in female harp seals, Ph. D. thesis, Guelph University, Guelph, Ontario.Google Scholar
  214. Stewart, R. E. A., and Lavigne, D. M., 1984, Energy transfer and female condition in nursing harp seals Phoca groenlandica, Holarct. Ecol. 7:182–194.Google Scholar
  215. Stewart, R. E. A., and Murie, D. J., 1986, Food habits of lactating harp seals (Phoca groenlandica) in the Gulf of St. Lawrence in March, J. Mammal. 67:186–188.Google Scholar
  216. Stewart, R. E. A., Lightfoot, N., and Innes, S., 1981, Parturition in harp seals, J. Mammal. 62:845–850.Google Scholar
  217. Stewart, R. E. A., Webb, B. E., Lavigne, D. M., and Fletcher, F., 1983, Determining lactose content of harp seal milk, Can. J. Zool. 61:1094–1100.Google Scholar
  218. Stirling, I., 1970, Observations on the behaviour of the New Zealand für seal (Arctocephalus forsten), J. Mammal. 51:766–778.Google Scholar
  219. Stirling, I., 1971, Studies on the behaviour of the South Australian für seal, Arctocephalus forsten (Lesson). 2. Adult females and pups, Aust. J. Zool. 19:267–273.Google Scholar
  220. Stirling, I., 1975, Adoptive suckling in pinnipeds, J. Aust. Mammal. Soc. 1:389–391.Google Scholar
  221. Stirling, I., 1983, The evolution of mating systems in pinnipeds, in: Advances in the Study of Mammalian Behavior (J. F. Eisenberg and D. G. Kleiman, eds.), American Society of Mammalogists, pp. 489-527.Google Scholar
  222. Stirling, I., and McEwan, E. H., 1975, Caloric value of whole ringed seals (Phoca hispida) in relation to polar bear (Ursus maritimus) ecology and hunting, Can. J. Zool. 8: 1021–1027.Google Scholar
  223. Stirling, I., and Warneke, R. M., 1971, Implications of a comparison of the airborne vocalizations and some aspects of the behaviour of two Australian für seal species on the evolution and present taxonomy of the genus, Aust. J. Zool. 19:227–241.Google Scholar
  224. Stull, J. W., Brown, W. H., and Kooyman, G. L., 1967, Lipids of the Weddell seal, Leptonychotes weddelli, J. Mammal. 48:642–645.Google Scholar
  225. Tedman, R. A., 1977, The gross anatomy of the mammary glands of the Weddell seal, Leptonychotes weddelli, J. Anat. 124:524.Google Scholar
  226. Tedman, R. A., 1980, Lactation in the Weddell seal, Leptonychotes weddelli, Ph. D. thesis, University of Queensland, Australia.Google Scholar
  227. Tedman, R. A., 1983, Ultrastructural morphology of the mammary gland with observations on the size distribution of fat droplets in milk of the Weddell seal, Leptonychotes weddelli (Pinnipedia), J. Zool. Lond. 200:131–141.Google Scholar
  228. Tedman, R. A., 1985, The Weddell seal, Leptonychotes weddelli, at McMurdo Sound, Antarctica: Milk production in relation to pup growth, in: Studies of Sea Mammals in South Latitudes (J. K. Ling and M. M. Bryden, eds.), South Australian Museum, Adelaide, pp. 41-52.Google Scholar
  229. Tedman, R. A., and Bryden, M. M., 1979, Cow-pup behaviour of the Weddell seal, Leptonychotes weddelli (Pinnipedia), in McMurdo Sound, Antarctica, Aust. Wildl. Res. 6:19–37.Google Scholar
  230. Tedman, R. A., and Bryden, M. M., 1981a, The mammary gland of the Weddell seal, Leptonychotes weddelli (Pinnipedia), 1. Gross and microscopic anatomy, Anat. Rec. 199:519–529.PubMedGoogle Scholar
  231. Tedman, R. A., and Bryden, M. M., 1981b, Resin injection of milk duct and arterial systems of the mammary glands of Weddell and southern elephant seals, Bull. Aust. Mammal. Soc. 7:61–62.Google Scholar
  232. Tedman, R. A., and Green, B., 1986, Water and sodium fluxes and lactational energetics in suckling pups of Weddell seals (Leptonychotes weddellii), J. Zool. Lond., in press.Google Scholar
  233. Thomas, J. A., Pastukhov, V., Eisner, R., and Petrov, E., 1982, Phoca siberica, Mammal. Species 188:1–6.Google Scholar
  234. Tikhomirov, E. A., 1975, Biology of the ice forms of seals in the Pacific section of the Antarctic, Rapp. P. V. Reun. Cons. Int. Explor. Mer 169:409–413.Google Scholar
  235. Tindal, J. S., 1978, Neuroendocrine control of lactation, in: Lactation A Comprehensive Treatise, Volume 4 (B. L. Larson, ed.), Academic Press, New York, pp. 67–114.Google Scholar
  236. Trillmich, F., 1979, Sea lions and für seals, Not. Galapagos 29:8–14.Google Scholar
  237. Trillmich, F., 1981, Mutual mother—pup recognition in Galapagos für seals and sea lions: Cues used and functional significance, Behaviour 78:21–42.Google Scholar
  238. Trillmich, F., 1986a, Attendance behavior of Galapagos für seals, in: Fur Seals: Maternal Strategies on Land and at Sea (R. L. Gentry and G. L. Kooyman, eds.), Princeton University Press, Princeton, pp. 168–185.Google Scholar
  239. Trillmich, F., 1986b, Attendance behavior of Galapagos sea lions, in: Fur Seals: Maternal Strategies on Land and at Sea (R. L. Gentry and G. L. Kooyman, eds.), Princeton University Press, Princeton, pp. 196–207.Google Scholar
  240. Trillmich, F., and Lechner, E., 1986, Milk of the Galapagos für seal and sea lion, with a comparison of the milk of eared seals (Otariidae), J. Zool. Lond. 209:271–277.Google Scholar
  241. Trillmich, F., and Majluf, P., 1981, First observations on colony structure, behaviour, and vocal repertoire of the South American für seal (Arctocephalus australis Zimmermann, 1783) in Peru, Z. Säugetierkd. 46:310–322.Google Scholar
  242. Trillmich, F., Majluf, P., Sanchez-Grinan, M., and Kooyman, G. L., 1986, Attendance and diving behavior of the South American für seal during “El Nino” in 1983, in: Fur Seais: Maternal Strategies on Land and at Sea (R. L. Gentry and G. L. Kooyman, eds.), Princeton University Press, Princeton, pp. 153–167.Google Scholar
  243. Van Utrecht, W. L., 1968, Notes on some aspects of the mammary glands in the fin whale, Balaenoptera physalus (L), with regard to the criterion’ lactating’, Nor. Hvalfangst-Tid. 57:1–13.Google Scholar
  244. Van Horn, D. R., and Baker, B. E., 1971, Seal milk. II. Harp seal (Pagophilis groenlandicus) milk: Effects of stage of lactation on the composition of the milk, Can. J. Zool. 49: 1085–1088.PubMedGoogle Scholar
  245. Vaz Ferriera, R., 1956, Terrestrial Ethology of ArctocephaJus austraJis in the Uruguayan Islands, Works Concerning Seals and Sea Lions, Oceanographics and Fisheries Service, No. 2, pp. 1-22.Google Scholar
  246. Venables, U. M., and Venables, L. S. V., 1955, Observations on a breeding colony of the seal, Phoca vitulina in Shetland, Proc. Zool. Soc. Lond. 125:521–532.Google Scholar
  247. Watson, D. L., 1980, Immunological functions of the mammary gland and its secretion — Comparative review, Aust. J. Biol. Sci. 33:403–422.PubMedGoogle Scholar
  248. Whitworth, M. R., 1984, Maternal care and behavioural development in pikas, Ochotona princeps, Anim. Behav. 32:743–752.Google Scholar
  249. Wooding, F. B. P., 1977, Comparative mammary fine structure, Symp. Zool. Soc. Lond. 41:1–41.Google Scholar
  250. Worthy, G. A. J., and Lavigne, D. M., 1983a, Energetics of fasting and subsequent growth in harp seal pups, Phoca groenlandica, Can. J. Zool. 61:447–456.Google Scholar
  251. Worthy, G. A. J., and Lavigne, D. M., 1983b, Changes in energy stores during postnatal development of the harp seal, Phoca groenlandica, J. Mammal. 64:89–96.Google Scholar

Copyright information

© Springer Science+Business Media New York 1987

Authors and Affiliations

  • Olav T. Oftedal
    • 1
  • Daryl J. Boness
    • 1
  • Raymond A. Tedman
    • 2
  1. 1.Department of Zoological ResearchNational Zoological Park, Smithsonian InstitutionUSA
  2. 2.Department of Anatomy and HistologyUniversity of AdelaideAdelaideAustralia

Personalised recommendations