Journal of Biosciences

, Volume 34, Issue 5, pp 673–686 | Cite as

The origin and early evolution of whales: macroevolution documented on the Indian Subcontinent

  • S. BajpaiEmail author
  • J. G. M. Thewissen
  • A. Sahni


The origin of whales (order Cetacea) from a four-footed land animal is one of the best understood examples of macroevolutionary change. This evolutionary transition has been substantially elucidated by fossil finds from the Indian subcontinent in the past decade and a half. Here, we review the first steps of whale evolution, i.e. the transition from a land mammal to obligate marine predators, documented by the Eocene cetacean families of the Indian subcontinent: Pakicetidae, Ambulocetidae, Remingtonocetidae, Protocetidae, and Basilosauridae, as well as their artiodactyl sister group, the Raoellidae. We also discuss the influence that the excellent fossil record has on the study of the evolution of organ systems, in particular the locomotor and hearing systems.


Cetacea Eocene evolution India whales 


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Bajpai S and Gingerich P D 1998 A new Eocene archaeocete (Mammalia, Cetacea) from India and the time of the origin of whales; Proc. Natl. Acad. Sci. USA 95 15464–15468CrossRefGoogle Scholar
  2. Bajpai S and Thewissen J G M 1998 Middle Eocene cetaceans from the Harudi and Subathu Formations of India; in The emergence of whales: evolutionary patterns in the origin of Cetacea (ed.) J G M Thewissen (New York: Plenum Press) First Edition, pp 213–234CrossRefGoogle Scholar
  3. Bajpai S and Thewissen J G M 2000 A new, diminuitive whale from Kachchh (Gujarat, India) and its implications for locomotor evolution of cetaceans; Curr. Sci. 79 1478–1482Google Scholar
  4. Bianucci G and Landini W 2007 Fossil history; in Reproductive biology and phylogeny of Cetacea (ed.) D E Miller (Enfield, NH: Science Publ) First edition, pp 35–94CrossRefGoogle Scholar
  5. Boisserie J R, Lihoreau F and Brunet M 2005 Origins of Hippopotamidae (Mammalia, Cetartiodactyla): towards resolution; Zool. Scr. 34 119–143CrossRefGoogle Scholar
  6. Buchholtz E A 1998 Implications of vertebral morphology for locomotor evolution in early Cetacea; in The emergence of whales: evolutionary patterns in the origin of Cetacea (ed.) J G M Thewissen (New York: Plenum Press) First edition, pp 325–352CrossRefGoogle Scholar
  7. Dehm R and zu Oettingen-Spielberg T 1958 Paläeontologische und geologische Untersuchungen im Tertiär von Pakistan. 2. Die mitteleocäenen Säugetiere von Ganda Kas bei Basal in Nordwest-Pakistan; Bayer. Akad. Wiss., Math.-Naturwiss. Kl. 91 54 3 plGoogle Scholar
  8. Fish F E 1994 Association of propulsive swimming mode with behaviour in river otters (Lutra canadensis); J. Mammal. 75 989–997CrossRefGoogle Scholar
  9. Fish F E 1996 Transitions from drag-based to lift-based propulsion in mammalian swimming; Am. Zool. 36 628–641CrossRefGoogle Scholar
  10. Fish F E 2000 Biomechanics and energetics in aquatic and semiaquatic mammals: platypus to whale; Physiol. Biochem. Zool. 73 683–698CrossRefGoogle Scholar
  11. Fish F E and Baudinette R V 2008 Energetics of swimming by the ferret: consequences of forelimb paddling; Comp. Biochem., Physiol., A., Mol. Integr. Physiol. 150 136–143CrossRefGoogle Scholar
  12. Fordyce E 2008 Cetacean evolution; in Encyclopedia of marine mammals (eds) W Perrin, B Würsig and J G M Thewissen (Elsevier) pp 193–199Google Scholar
  13. Fordyce E and Muizon Cd 2001 Evolutionary history of cetaceans: a review; in Secondary adaptation of tetrapods to life in water (eds) J M Mazin and Vd Buffrénil (Munich: Verlag Friedrich Pfeil) First edition, pp 169–233Google Scholar
  14. Gatesy J and O’Leary M A 2001 Deciphering whale origins with molecules and fossils; Tr. Ecol. Evol. 16 562–570CrossRefGoogle Scholar
  15. Geisler C D 1998 From sound to synapse: physiology of the mammalian ear (New York: Oxford University Press)Google Scholar
  16. Geisler J H and Theodor J M 2009 Hippopotamus and whale phylogeny; Nature (London) 458, E5, doi:10.1038/Nature07776CrossRefGoogle Scholar
  17. Geisler J H and Uhen M D 2003 Morphological support for a close relationship between hippos and whales; J. Vertebr. Paleontol. 23 991–996CrossRefGoogle Scholar
  18. Geisler J H and Uhen M D 2005 Phylogenetic relationships of extinct Cetartiodactyls: results of simultaneous analyses of molecular, morphological, and stratigraphic data; J. Mamm. Evol. 12 145–160CrossRefGoogle Scholar
  19. Geisler J H, Theodor J M, Uhen M D and Foss S E 2007 Phylogenetic relationships of cetaceans to terrestrial artiodactyls; in The evolution of artiodactyls (eds) D R Prothero and S E Foss Baltimore, MD: Johns Hopkins Univ. Press) First edition, pp 19–31Google Scholar
  20. Gingerich P D 2003 Land-to-sea transition of early whales: evolution of Eocene Archaeoceti (Cetacea) in relation to skeletal proportions and locomotion of living semiaquatic mammals; Paleobiology 29 429–454CrossRefGoogle Scholar
  21. Gingerich P D, Smith B H, Simons E L 1990 Hind limbs of Basilosaurus isis: evidence of feet in whales; Science 229 154–157CrossRefGoogle Scholar
  22. Gingerich P D, Raza S M, Arif M, Anwar M and Zhou X 1994 New whale from the Eocene of Pakistan and the origin of cetacean swimming; Nature (London) 368 844–847CrossRefGoogle Scholar
  23. Gingerich P D, Haq M, Zalmout I S, Khan I H and Malkani M S 2001 Origin of whales from early artiodactyls: hands and feet of Eocene Protocetidae from Pakistan; Science 293 2239–2242CrossRefGoogle Scholar
  24. Gingerich P D, ul-Haq M, Koenigswald W V, Sanders W J, Smith B H, Zalmout S 2009 New protocetid whale from the Middle Eocene of Pakistan: Birth on land, precocial development, and sexual dimorphism; PLOS One 4 e4366. doi:10.1371/journal.pone.0004366CrossRefGoogle Scholar
  25. Grey N M, Kainec K, Madar S, Tomko L, and Wolfe S 2007 Sink or swim? Bone density as a mechanism for buoyancy control in early cetaceans; Anat. Rec. Adv. Integr. Anat. Evol. Biol. 290 638–653CrossRefGoogle Scholar
  26. Hemilä S, Nummela Z and Reuter T 1999 A model of the odontocete middle ear; Hear Res. 133 82–97CrossRefGoogle Scholar
  27. Hulbert R C Jr, Petkewich R M, Bishop G A, Burky D and Aleshire D P 1998 A new middle Eocene protocetid whale (Mammalia: Cetacea: Archaeoceti) and associated biota from Georgia; J. Paleontol. 72 905–925CrossRefGoogle Scholar
  28. Hulbert R C Jr 1998 Postcranial osteology of the North American middle Eocene protocetid Georgiacetus; in The emergence of whales: evolutionary patterns in the origin of Cetacea (ed.) J G M Thewissen (New York: Plenum Press) First edition, pp 235–267CrossRefGoogle Scholar
  29. Kellogg R 1936 A review of the Archaeoceti (Carnegie Institution of Washington, DC) 366 pp, 37 plGoogle Scholar
  30. Kumar K and A Sahni 1986 Remingtonocetus harudiensis, new combination, a middle Eocene archaeocete (Mammalia, Cetacea) from Kutch, western India; J. Vertebr. Paleontol. 6 326–349CrossRefGoogle Scholar
  31. Lombard R E, Hetherington T E 1993 Structural basis of hearing and sound transmission; in The skull (eds) J H Hanken and B K Hall (London: University of Chicago Press) pp 241–302Google Scholar
  32. Lucas F A 1900 The pelvic girdle of Zeuglodon, Basilosaurus cetoides (Owen), with notes on other portions of the skeleton; Proc. US Natl. Mus. 23 327–331CrossRefGoogle Scholar
  33. Madar S I 1998 Structural adaptations of early archaeocette long bones; in The emergence of whales: evolutionary patterns in the origin of Cetacea (ed.) J G M Thewissen (New York: Plenum Press) First edition, pp 353–378CrossRefGoogle Scholar
  34. Madar S I 2007 The postcranial skeleton of pakicetid cetaceans; J. Paleontol. 81 176–200CrossRefGoogle Scholar
  35. Madar S I, Thewissen J G M and Hussain S T 2002 Additional holotype remains of Ambulocetus natans (Cetacea, Ambulocetidae), and their implications for locomotion in early whales; J. Vertebr. Paleontol. 22 405–422CrossRefGoogle Scholar
  36. Nikaido M, Rooney A P and Okada N 1999 Phylogenetic relationships among cetartiodactyls based on insertions of short and long interspersed elements: hippopotamuses are the closest extant relatives of whales; Proc. Natl. Acad. Sci. USA 96 10261–10266CrossRefGoogle Scholar
  37. Norris K S 1968 The evolution of acoustic mechanisms in odontocete cetaceans; in Evolution and environment (ed.) E T Drake (New Haven: Yale University Press) pp 297–324Google Scholar
  38. Nummela S, Thewissen J G M, Bajpai S, Hussain S T and Kumar K 2004 Eocene evolution of whale hearing; Nature (London) 430 776–778CrossRefGoogle Scholar
  39. Nummela S, Hussain S T and Thewissen J G M 2006 Cranial anatomy of Pakicetidae (Cetacea, Mammalia); J. Vertebr. Paleontol. 26 746–759CrossRefGoogle Scholar
  40. Nummela S, Thewissen J G M, Bajpai S, Hussain T and Kumar K 2007 Sound transmission in archaic and modern whales: anatomical adaptations for underwater hearing; Anat. Rec. 290 716–733CrossRefGoogle Scholar
  41. Price S A, Bininda-Edmonds O R P and Gitttleman J L 2005 A complete phylogeny of the whales, dolphins, and even-toed hoofed mammals (Cetartiodactyla); Biol. Rev. 80 445–473CrossRefGoogle Scholar
  42. Ranga Rao A 1971 New mammals from Murree (Kalakot zone) of the foot-hills near Kalakot, J & K State; J. Geol. Soc. India 12 125–134Google Scholar
  43. Sahni A and Mishra V P 1972 A new species of Protocetus (Cetacea) from the middle Eocene of Kutch, Western India; Palaeontology 15 490–495Google Scholar
  44. Sahni A and Mishra V P 1975 Lower Tertiary vertebrates from western India; Monogr. Palaeontol. Soc. India 3 48, 6 plGoogle Scholar
  45. Schaeffer B 1947 Notes on the origin and function of the artiodactyls tarsus; Am. Mus. Novit. 1356 1–24Google Scholar
  46. Shedlock A M, Milinkovitch M C and Okada N 2000 SINE evolution, missing data and the origin of whales; Syst. Biol. 49 808–817CrossRefGoogle Scholar
  47. Struthers J 1893 On the rudimentary hind-limb of a great fin-whale (Balaenoptera musculus) in comparison with those of the humpback and the Greenland right-whale; J. Anat. Phys. 7 291–335Google Scholar
  48. Thewissen J G M and Fish F E 1997 Locomotor evolution in the earliest cetaceans: functional model, modern analogues, and paleontological evidence; Paleobiology 23 482–490CrossRefGoogle Scholar
  49. Thewissen J G M and Hussain S T 1993 Origin of underwater hearing in whales; Nature (London) 361 444–445CrossRefGoogle Scholar
  50. Thewissen J G M and Williams E M 2002 The early evolution of Cetacea (whales, dolphins, and porpoises); Ann. Rev. Ecol. Syst. 33 73–90CrossRefGoogle Scholar
  51. Thewissen J G M and Nummela S 2007 Toward an integrative approach; in Sensory evolution on the threshold (eds) J G M Thewissen and S Nummela (Berkeley: University of California Press) pp 333–340Google Scholar
  52. Thewissen J G M, Cooper L N, George J C and Bajpai S 2009 From Land to Water: the origin of whales, dolphins, and porpoises; Evol. Edu. Outreach 2 272–288CrossRefGoogle Scholar
  53. Thewissen J G M, Hussain S T and Arif M 1994 Fossil evidence for the origin of aquatic locomotion in archaeocete whales; Science 263 210–212CrossRefGoogle Scholar
  54. Thewissen J G M, Madar S I and Hussain S T 1996 Ambulocetus natans, an Eocene cetacean (Mammalia) from Pakistan; Courier Forschungs-Institut Senckenberg series 191, pp 1–86Google Scholar
  55. Thewissen J G M, Williams E M, Roe L J, and Hussain S T 2001 Skeletons of terrestrial cetaceans and the relationship of whales to artiodactyls; Nature (London) 413 277–281CrossRefGoogle Scholar
  56. Thewissen J G M, Cohn M J, Stevens L S, Bajpai S, Heyning J, Horton W E Jr 2006 Developmental basis for hind limb loss in dolphins and the origin of the cetacean bodyplan; Proc. Natl. Acad. Sci. USA 103 8414–8418CrossRefGoogle Scholar
  57. Thewissen J G M, Cooper L N, Clementz M T, Bajpai S and Tiwari B N 2007 Whales originated from aquatic artiodactyls in the Eocene epoch of India; Nature (London) 450 1190–1195CrossRefGoogle Scholar
  58. Thewissen J G M and Bajpai S 2009 New skeletal material for Andrewsiphius and Kutchicetus, two Eocene cetaceans from India; J. Paleontol. 2 272–288Google Scholar
  59. Uhen M D 1998 Middle to late Eocene basilosaurines; in The emergence of whales: evolutionary patterns in the origin of Cetacea (ed.) J G M Thewissen (New York: Plenum Press) First edition, pp 29–61CrossRefGoogle Scholar
  60. Uhen M D 2004 Form, function, and anatomy of Dorudon atrox (Mammalia, Cetacea): an archaeocete from the middle to late Eocene of Egypt; Univ. Michigan. Pap. Pal. 34 1–222Google Scholar
  61. Uhen M D 2008 The oldest cetaceans from the southern hemisphere: new archaeocetes from the Pisco Basin of Southern Peru; J. Vertebr. Paleontol. Progr. Abstr., Suppl. 28 154AGoogle Scholar
  62. Van Valen L 1966 Deltatheridia, a new order of mammals; Bull. Am. Mus. Nat. Hist. 132 pp.126Google Scholar
  63. West R M 1980 Middle Eocene large mammal assemblage with Tethyan affinities, Ganda Kas region, Pakistan; J. Paleontol. 54 508–533Google Scholar
  64. Williams E M 1998 Synopsis of the earliest cetaceans Pakicetidae, Ambulocetidae, Remingtonocetidae, and Protocetidae; in The emergence of whales: evolutionary patterns in the origin of Cetacea (ed.) J G M Thewissen (New York: Plenum Press) First edition, pp 1–28Google Scholar

Copyright information

© Indian Academy of Sciences 2009

Authors and Affiliations

  1. 1.Department of Earth SciencesIndian Institute of TechnologyRoorkeeIndia
  2. 2.Department of Anatomy and NeurobiologyNortheastern Ohio Universities College of MedicineRootstownUSA
  3. 3.Geology DepartmentPanjab UniversityChandigarhIndia

Personalised recommendations