Free-Ranging Energetics of Penguins

  • Randall W. Davis
  • Gerald L. Kooyman


Penguins exhibit the greatest degree of adaptation to the aquatic environment of all seabirds. They are powerful swimmers that pursue their prey at depth allowing them to exploit the water column more than any other seabird. Dives 240 m deep have been recorded for King Penguins (Kooyman et al., 1982) and as deep as 265 m for Emperor Penguins (Kooyman et al., 1971). Intermediate sized Gentoo and Chinstrap Penguins dive from 70–100 m (Conroy and Twelves, 1972; Lishman and Croxall, in prep) and even the smallest species, the Fairy Penguin, can dive to 60 m (Montague, 1982). How does swimming as a primary mode of locomotion at sea influence the average daily metabolic rate (ADMR) and foraging energetics of penguins relative to seabirds that fly to and from feeding areas?


Standard Metabolic Rate Tritiated Water Gentoo Penguin Diving Behavior Diving Depth 
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. Aschoff, J., and Pohl, H., 1970, Rhythmic variations in energy metabolism, Fed. Proc., 29: 1541.PubMedGoogle Scholar
  2. Barre, H., 1975, Le jeune du manchot royal (Aptenodytes patagonica J. F. Miller) a l’ile de la Possession, Comptes rendus, Academie des sciences, Paris, Serie D, 280: 2885.Google Scholar
  3. Brown, C. R., In press, Resting metabolic rate and energetic cost of incubation in macaroni penguins Eudyptes chrysolophus) and rockhopper penguins (E. chrysocome), Comp. Biochem. Physiol.Google Scholar
  4. Conroy, J. W. H., and Twelves, E. L., 1972, Diving depths of the gentoo penguin Pygoscelis papua and blue-eyed shag Phalacrocorax atriceps from the South Orkney Islands, Br. Antarc. Surv. Bull., 30:10’.Google Scholar
  5. Copestake, P. G., Croxall, J. P., and Prince, P. A., 1983, Food digestion and energy consumption experiments on a king penguin Aptenodytes patagonicus, Br. Antarct. Surv. Bull., 58: 83.Google Scholar
  6. Croxall, J.P., 1982,. Energy costs of incubation and moult in petrels and penguins, J. Anim. Ecol., 51: 177.CrossRefGoogle Scholar
  7. Croxall, J. P., and Prince, P. A., 1980, The food of gentoo penguins, Pygoscelis papua, and macaroni penguins, Eudyptes chrysolophus, at South Georgia, Ibis, 122: 245.CrossRefGoogle Scholar
  8. Davis, R. W., Kooyman, G.L., and Croxall, J.P., 1983, Water flux and estimated metabolism of free-ranging gentoo and macaroni penguins at South Georgia, Polar Biol., 2: 41.CrossRefGoogle Scholar
  9. Dewasmes, G., Le Maho, Y., Cornet, A., and Groscolas, R., 1980, Resting metabolic rate and cost of locomotion in long-term fasting emperor penguins, J. Appl. Physiol., 49: 888.PubMedGoogle Scholar
  10. Hails, C. J., and Bryant, D. M., 1979, Reproductive energetics of a free-living bird, J. Anim. Ecol., 48: 471.CrossRefGoogle Scholar
  11. Kendeigh, S. C., Dol’nik, V. R., and Gavrilov, V. M., 1977, Avian energetics, In: “Granivorous Birds in Ecosystems,” J. Pinowski and S. C. Kendeigh, ed., Cambridge University Press, Cambridge, New York, pp. 127–204.Google Scholar
  12. King, J. R., 1974, Seasonal allocation of time and energy resources in birds. In: “Avian Energetics,” R. A. Paynter, Jr., ed., Nuttal Ornithological Club, Cambridge, Mass.Google Scholar
  13. Kooyman, G. L., Drabek, C. M., Elsner, R., and Campbell, W. B., 1971, Diving behavior of the emperor penguin, Aptenodytes forsteri, The Auk, 88: 775.Google Scholar
  14. Kooyman, G. L., Davis, R. W., Croxall, J. P., and Costa, D. P., 1982, Diving depths and energy requirements of king penguins, Science, 217: 726.PubMedCrossRefGoogle Scholar
  15. Le Maho, Y., Philippe, D., and Chatonnet, J., 1976, Thermoregulation in fasting emperor penguins under natural conditions, Am. J. Physiol., 231: 913.PubMedGoogle Scholar
  16. Lishman, G. S., and Croxall, J. P., In Prep., Diving depths of the chinstrap penguin Pygoscelis antarctica.Google Scholar
  17. Montague, T. L., 1982, The food and feeding ecology of the little penguin (Eudyptula minor) at Phillip Island, Victoria, Australia, Doctoral Thesis, Monash University.Google Scholar
  18. Mougin, J. L., 1974, Enregistrements continus de temperatures internes chez quelques Spheniscidae. 2. Le manchot royal Aptenodytes patagonica de l’ile de la Possession (archipel Crozet), Com. Nat. Franc. Recherch. Antarct., 33: 29.Google Scholar
  19. Nagy, K. H., 1980, CO production in animals: analysis of potential errors in the doubly labeled water method, Am. J. Physiol., 238: R456.Google Scholar
  20. Pinshow, B., Fedak, M. P,., Battles, D. R., and Schmidt-Nielsen, K., 1976, Energy expenditure for thermoregulation and locomotion in emperor penguins, Am. J. Physiol., 231: 903.PubMedGoogle Scholar
  21. Pinshow, B., Fedak, M. A., and Schmidt-Nielsen, K., 1977,. Terrestrial locomotion in penguins: It costs more to waddle, Science, 195: 592.PubMedCrossRefGoogle Scholar
  22. Richards, P. A., and Tickell, W. L. N., 1968, Comparison between the weather at bird island and King Edward Point, South Georgia, Br. Antarc. Surv. Bull., 15: 63.Google Scholar
  23. Schoeller, D. A., and van Santen, E., 1982, Measurement of energy expenditure in humans by doubly labeled water method, J. Appl. Physiol.:Respirat. Environ. Exercise Physiol., 53: 955.Google Scholar
  24. Shoemaker, V. H., Nagy, K. A., and Costa, W. R., 1976, Energy utilization and temperature regulation by jackrabbits Lepus californicus in the Mojave desert, Physiol. Zool., 49: 364.Google Scholar
  25. Stahel, C. D., and Nicol, S. C., 1982, Temperature regulation in the little penguin, Eudyptula minor in air and water, J. Comp. Physiol., 148: 93.Google Scholar
  26. Stonehouse, B., 1960, The king penguin Aptenodytes patagonica of South Georgia I. Breeding behaviour and development, F.I.D.S. Scientific Reports, 23: 1.Google Scholar
  27. Wilson, R. P., and Bain, C. A. R., In press, An inexpensive depth gauge for penguins, J. Wildl. Manage.Google Scholar

Copyright information

© Plenum Press, New York 1984

Authors and Affiliations

  • Randall W. Davis
    • 1
  • Gerald L. Kooyman
    • 1
  1. 1.Physiological Research LaboratoryScripps Institution of OceanographyLa JollaUSA

Personalised recommendations