Evolution of Thermoregulatory Function in Cetacean Reproductive Systems

  • D. Ann Pabst
  • Sentiel A. Rommel
  • William A. McLellan
Part of the Advances in Vertebrate Paleobiology book series (AIVP, volume 1)


Modern cetaceans possess a suite of morphological adaptations that permit their existence in the marine environment (e.g., Howell, 1930; Slijper, 1936, 1979). Their streamlined body shape, hypertrophied axial musculoskeletal system, thick blubber layer, and de novo dorsal fin and flukes are morphological features that reduce the energetic costs of both swimming (e.g., Fish and Hui, 1991; Williams et al., 1992; Fish, 1993a,b; Pabst, 1996) and whole body thermoregulation (e.g., Worthy and Edwards, 1990; Koopman et al., 1996).


Bottlenose Dolphin Dorsal Aorta Harbor Porpoise Superficial Vein Sigmoid Flexure 
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. Alexander, G., Hales, J. R. S., Stevens, D., and Donnelly, J. B. 1987. Effects of acute and prolonged exposure to heat on regional blood flows in pregnant sheep. J. Dev. Physiol. 9:1–15.PubMedGoogle Scholar
  2. Anthony, R. 1920. L’exorchidie du Mesoplodon et la remontee des testicules as cours de la phylogenese des cetaces. C. R. Acad. Sci. 170:529–531.Google Scholar
  3. Arkowitz, R. A., and Rommel, S. A. 1985. Force and bending moment of the caudal muscles in the short finned pilot whale. Mar. Mamm. Sci. 1(3):203–209.CrossRefGoogle Scholar
  4. Arnason, U., and Gullberg, A. 1996. Cytochrome b nucleotide sequences and the identification of five primary lineages of extant cetaceans. Mol. Biol. Evol. 13(2):407–417.PubMedCrossRefGoogle Scholar
  5. Bailey, F. R., and Miller, A. M. 1909. Textbook of Embryology. William Wood & Co., York, PA.Google Scholar
  6. Barnett, C. H., Harrison, R. J., and Tomlinson, J. D. W. 1958. Variations in the venous systems of mammals. Biol. Rev. 33:442–487.CrossRefGoogle Scholar
  7. Bedford, J. M. 1977. Evolution of the scrotum: the epididymis as the prime mover, in: J. H. Calaby and C. H. Tyndale-Biscoe (eds.), Reproduction and Evolution, pp. 171–182. Australian Academy of Science, Canberra City.Google Scholar
  8. Bell, A. W. 1987. Consequences of severe heat stress for fetal development, in: J. R. S. Hales and D. A. B. Richards (eds.), Heat Stress: Physical Exertion and Environment, pp. 313–333. Elsevier, Amsterdam.Google Scholar
  9. Benirschke, K., and Cornell, L. H. 1987. The placenta of the killer whale, Orcinus orca. Mar. Mamma. Sci. 3(1):82–86.CrossRefGoogle Scholar
  10. Boice, R. C., Swift, M. L., and Roberts, J. C., Jr. 1964. Cross-sectional anatomy of the dolphin. Nor. Hvalfangst Tid. 7:178–193.Google Scholar
  11. Brodie, P. F., and Paasche, A. 1985. Thermoregulation and energetics of fin and sei whales based on postmortem, stratified temperature measurements. Can. J. Zool. 63:2267–2269.CrossRefGoogle Scholar
  12. Carrick, F. N., and Setchell, B. P. 1977. The evolution of the scrotum, in: J. H. Calaby and C. H. Tyndale-Biscoe (eds.), Reproduction and Evolution, pp. 165–170. Australian Academy of Science, Canberra City.Google Scholar
  13. Cartmill, M., Hylander, W. L., and Shafland, J. 1987. Human Structure. Harvard University Press, Cambridge, MA.Google Scholar
  14. Cephalo, R. C., and Hellegers, A. E. 1978. The effect of maternal hyperthermia on maternal and fetal cardiovascular and respiratory function. Am. J. Obstet. Gynecol. 131:687–694.Google Scholar
  15. Cowles, R. B. 1958. The evolutionary significance of the scrotum. Evolution XII:417–418.CrossRefGoogle Scholar
  16. Cowles, R. B. 1965. Hyperthermia, aspermia, mutation rates and evolution. Q. Rev. Biol. 40:341–367.CrossRefGoogle Scholar
  17. De Smet, W. M. A. 1977. The position of the testes in cetaceans, in: R. J. Harrison (ed.), Functional Anatomy of Marine Mammals, Volume 3, pp. 361–386. Academic Press, London.Google Scholar
  18. Eisner, R., Pirie, J., Kenney, D. D., and Schemmer, S. 1974. Functional circulatory anatomy of cetacean appendages, in: R. J. Harrison (ed.), Functional Anatomy of Marine Mammals, Volume 2, pp. 143–159. Academic Press, London.Google Scholar
  19. Evans, H. E., and Christensen, G. C. 1993. The urogenital system, in: H. E. Evans (ed.), Miller’s Anatomy of the Dog, 3rd ed., pp. 494–558. Saunders, Philadelphia.Google Scholar
  20. Fish, F. E. 1993a. Power output and propulsive efficiency of swimming bottlenose dolphins (Tursiops truncatus). J. Exp. Biol. 185:179–193.Google Scholar
  21. Fish, F. E. 1993b. Influence of hydrodynamic design and propulsive mode on mammalian swimming energetics. Aust. J. Zool. 42:79–101.CrossRefGoogle Scholar
  22. Fish, F. E., and Hui, C. A. 1991. Dolphin swimming—a review. Mammal. Rev. 21:181–195.CrossRefGoogle Scholar
  23. Gatesy, J., Hayashi, C., Cronin, M. A., and Arctander, P. 1996. Evidence from milk casein genes that cetaceans are close relatives of hippopotamid artiodactyls. Mol. Biol. Evol. 13(7):954–963.PubMedCrossRefGoogle Scholar
  24. Gilbert, R. D., and Power, G. G. 1986. Fetal and uteroplacental heat production in sheep. J. Appl. Physiol. 61:2018–2022.PubMedGoogle Scholar
  25. Gilbert, R. D., Schroder, HL, Kawamura, T., Dale, P. S., and Power, G. G. 1985. Heat transfer pathways between fetal lamb and ewe. J. Appl Physiol. 59:634–638.PubMedGoogle Scholar
  26. Graur, D., and Higgins, D. G. 1994. Molecular evidence for the inclusion of cetaceans within the order Artiodactyla. Mol. Biol. Evol. 11(3):357–364.PubMedGoogle Scholar
  27. Harrison, R. J. 1948. The comparative anatomy of the blood-supply of the mammalian testis. Proc. Zool Soc. London 11:325–334 (plates I–V).Google Scholar
  28. Harrison, R. J. 1969. Reproduction and reproductive organs, in: H. T. Anderson (ed.), The Biology of Marine Mammals, pp. 253–348. Academic Press, New York.Google Scholar
  29. Harrison, R. J., and Tomlinson, J. D. W. 1956. Observations on the venous system in certain pinnipedia and cetacea. Proc. Zool Soc. London 126:205–233.CrossRefGoogle Scholar
  30. Hart, F. M., and Farber, J. J. 1985. Fetal and maternal temperatures in rabbits. J. Appl Physiol. 20:737–741.Google Scholar
  31. Howell, A. B. 1930. Aquatic Mammals: Their Adaptations to Life in the Water. Thomas, Springfield, IL.Google Scholar
  32. Hudlicka, O., and Tyler, K. R. 1986. Angiogenesis. Academic Press, Orlando, FL.Google Scholar
  33. Hyman, L. H. 1942. Comparative Vertebrate Anatomy. University of Chicago Press, Chicago.Google Scholar
  34. Kanwisher, J., and Sundes, G. 1965. Physiology of a small cetacean. Hvalradets Skr. 48:45–53.Google Scholar
  35. Koopman, H. N., Iverson, S. J., and Gaskin, D. E. 1996. Stratification and age-related differences in blubber fatty acids of the male harbour porpoise (Phocoena phocoena). J. Comp. Physiol. B 165:628–639.PubMedCrossRefGoogle Scholar
  36. Lotgering, F. K., Gilbert, R. D., and Longo, L. D. 1985. Maternal and fetal responses to exercise during pregnancy. Physiol. Rev. 65(1):1–29.PubMedGoogle Scholar
  37. Mackay, R. S. 1964. Deep body temperature of an untethered dolphin recorded by radio transmitter. Science 144:864–866.PubMedCrossRefGoogle Scholar
  38. Meek, A. 1918. The reproductive organs of cetacea. J. Anat. 52:186–210.PubMedGoogle Scholar
  39. Moore, C. R. 1926. The biology of the mammalian testis and scrotum. Q. Rev. Biol. 1:4–50.CrossRefGoogle Scholar
  40. Morishima, H. O., Glaser, B., Niemann, W. H., and James, L. S. 1975. Increased uterine activity and fetal deterioration during maternal hyperthermia. Am. J. Obstet. Gynecol 121(4):531–538.PubMedGoogle Scholar
  41. Nickel, R., Schummer, A., Seiferle, E., Wilkens, H., Wille, K.-H., and Frewein, J. 1986. The Locomotor System of the Domestic Mammals, Volume 1. Springer-Verlag, Berlin.Google Scholar
  42. Ommanney, F. D. 1932. The urogenital system of the fin whale (Balaenoptera physalus) with appendix: the dimensions and growth of the kidneys of blue and fin whales. Discovery Rep. 5:363–465.Google Scholar
  43. Pabst, D. A. 1996. Morphology of the subdermal connective tissue sheath of dolphins: a new fibre-wound, thin-walled, pressurized cylinder model for swimming vertebrates. J. Zool (London) 238:35–52.CrossRefGoogle Scholar
  44. Pabst, D. A., Rommel, S. A., McLellan, W. A., Williams, T. M., and Rowles, T. K. 1995. Thermoregulation of the intra-abdominal testes of the bottlenose dolphin (Tursiops truncatus) during exercise. J. Exp. Biol. 198:221–226.PubMedGoogle Scholar
  45. Power, G. G., Schroder, H., and Gilbert, R. D. 1984. Measurement of fetal heat production using differential calorimetry. J. Appl Physiol. 57(3):17–22.Google Scholar
  46. Prothero, D. R. 1993. Ungulate phytogeny: molecular vs. morphological evidence, in: F. S. Szalay, M. J. Novacek, and M. C. McKenna (eds.). Mammal Phylogeny, pp. 173–181. Springer-Verlag, Berlin.CrossRefGoogle Scholar
  47. Pycraft, W. P. 1932. On the genital organs of a female common dolphin (Delphinus delphis). Proc. Zool Soc. London 1932:807–811 (3 plates).Google Scholar
  48. Ridgway, S. H. 1965. Medical care of marine mammals. J. Am. Vet. Med. Assoc. 147:1077–1085.PubMedGoogle Scholar
  49. Ridgway, S. H. 1968. The bottlenosed dolphin in biomedical research. Methods Anim. Exp. 3:416–417.Google Scholar
  50. Ridgway, S. H. 1972. Homeostasis in the aquatic environment, in: S. H. Ridgway (ed.), Mammals of the Sea: Biology and Medicine, pp. 590–747. Thomas, Springfield, IL.Google Scholar
  51. Rommel, S. A. 1990. Osteology of the bottlenose dolphin, in: S. Leatherwood and R. R. Reeves (eds.), The Bottlenose Dolphin, pp. 29–49. Academic Press, San Diego, CA.Google Scholar
  52. Rommel, S. A., Pabst, D. A., McLellan, W. A., Mead, J. G., and Potter, C. W. 1992. Anatomical evidence for a countercurrent heat exchanger associated with dolphin testes. Anat. Rec. 232(1):150–156.PubMedCrossRefGoogle Scholar
  53. Rommel, S. A., Pabst, D. A., and McLellan, W. A. 1993. Functional morphology of the vascular plexuses associated with the cetacean uterus. Anat. Rec. 237(4):538–546.PubMedCrossRefGoogle Scholar
  54. Rommel, S. A., Pabst, D. A., McLellan, W. A., Williams, T. M., and Friedl, W. A. 1994. Temperature regulation of the testes of the bottlenose dolphin (Tursiops truncatus): evidence from colonic temperatures. J. Comp. Physioi. B 164:130–134.CrossRefGoogle Scholar
  55. Schmidt-Nielsen, K. 1990. Animal Physiology: Adaptation and Environment, 4th ed. Cambridge University Press, London.Google Scholar
  56. Scholander, P. F., and Schevill, W. E. 1955. Counter-current vascular heat exchange in the fins of whales. J. Appl. Physiol. 8:279–282.PubMedGoogle Scholar
  57. Schroeder, J. P. 1990. Breeding bottlenose dolphins in captivity, in: S. Leatherwood and R. R. Reeves (eds.), The Bottlenose Dolphin, pp. 425–446. Academic Press, San Diego, CA.Google Scholar
  58. Schummer, A., Nickel, R., and Sack, W. O. 1979. The Viscera of the Domestic Mammals. Verlag Paul Parey, Berlin.Google Scholar
  59. Schummer, A., Wilkens, H., Vollmerhaus, B., and Habermehl, K.-H. 1981. The Circulatory System, the Skin, and the Cutaneous Organs of the Domestic Mammals. Verlag Paul Parey, Berlin.Google Scholar
  60. Shelton, M. 1964. Relation of environmental temperature during gestation on birth weight and mortality in lambs. J. Anim. Sci. 23:360–364.Google Scholar
  61. Simpson, J. G., and Gardner, M. B. 1972. Comparative microscopic anatomy of selected marine mammals, in: S. H. Ridgway (ed.), Mammals of the Sea, pp. 298–418. Thomas, Springfield, IL.Google Scholar
  62. Slijper, E. J. 1936. Die Cetaceen: Vergleichend-Anatomisch und Systematisch. Asher & Co., Amsterdam, 1972 reprint.Google Scholar
  63. Slijper, E. J. 1966. Functional morphology of the reproductive system in Cetacea, in: K. S. Noms (ed.), Whales, Dolphins, and Porpoises, pp. 277–319. University of California Press, Berkeley.Google Scholar
  64. Slijper, E. J. 1979. Whales. Cornell University Press, Ithaca, NY.Google Scholar
  65. Smith, M. R., Shivji, M. S., Waddell, V. G., and Stanhope, M. J. 1996. Phylogenetic evidence from the IRBP gene for the paraphyly of toothed whales, and mixed support for Cetacea as a suborder of Artiodactyla. Mol. Biol. Evol. 13(7):918–922.PubMedCrossRefGoogle Scholar
  66. Thewissen, J. G. M. 1994. Phylogenetic aspects of cetacean origins: a morphological perspective. J. Mamm. Evol. 2(3):157–184.CrossRefGoogle Scholar
  67. Van Demark, N. L., and Free, M. J. 1970. Temperature regulation and the testis, in: A. D. Johnson, W. R. Gomes, and N. L. Van Demark (eds.), The Testis, Volume III, pp. 233–312. Academic Press, New York.Google Scholar
  68. Van der Schoot, P. 1995. Studies on the fetal development of the gubernaculum in Cetacea. Anat. Rec. 243:449–460.PubMedCrossRefGoogle Scholar
  69. Van Valen, L. 1966. Monophyly or diphyly in the origin of whales. Evolution 22:37–41.CrossRefGoogle Scholar
  70. Walmsley, R. 1938. Some observations on the vascular system of a female fetal finback. Contrib. Embryol. 27:109–178.Google Scholar
  71. Whittow, G. C., Hampton, I. F. G., Matsura, D. T., Ohata, C. A., Smith, R. M., and Allen, J. F. 1974. Body temperature of three species of whales. J. Mammal. 55:653–656.PubMedCrossRefGoogle Scholar
  72. Wiley, E. O. 1981. Phylogenetics: The Theory and Practice of Phylogenetic Systematics. Wiley, New York.Google Scholar
  73. Williams, T. M., Friedl, W. A., Fong, M. L., Yamada, R. M., Sedivy, P., and Haun, J. E. 1992. Travel at low energetic cost by swimming and wave-riding bottlenose dolphins. Nature 355:821–823.PubMedCrossRefGoogle Scholar
  74. Wislocki, G. B. 1933. On the placentation of the harbor porpoise (Phocoena phocoena Linnaeus). Biol. Bull. 65:80–98.CrossRefGoogle Scholar
  75. Wislocki, G. B., and Enders, R. K. 1941. The placentation of the bottle-nosed porpoise (Tursiops truncatus). Am. J.Anat. 68(1):97–125.CrossRefGoogle Scholar
  76. Woollard, H. H. 1922. The development of the principal arterial stems in the forelimb of the pig. Contributions to Embryology, Carnegie Institute of Washington 70:139–154 + plates 1 and 2.Google Scholar
  77. Worthy, G. A. J., and Edwards, E. F. 1990. Morphometric and biochemical factors affecting heat loss in a small temperate cetacean (Phocoena phocoena) and a small tropical cetacean (Stenella attenuata). Physiol. Zool. 63(2):432–442.Google Scholar

Copyright information

© Springer Science+Business Media New York 1998

Authors and Affiliations

  • D. Ann Pabst
    • 1
  • Sentiel A. Rommel
    • 2
  • William A. McLellan
    • 1
  1. 1.Biological Sciences and Center for Marine Science ResearchUniversity of North Carolina at WilmingtonWilmingtonUSA
  2. 2.Florida Marine Research Institute, Florida Department of Environmental ProtectionMarine Mammal Pathobiology LaboratorySt. PetersburgUSA

Personalised recommendations