The Walking Whales

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by J G M Hans Thewissen


  Fish,  “Mechanics,  Power  Output,  and  Efficiency  of  the  Swimming  Muskrat

  ( Ondatra zibethicus),”  Journal of Experimental Biology 110 (1984): 183–210.

  5.  F. E.  Fish, “Dolphin  Swimming: A  Review,”  Mammal Review  4  (1991):

  181–95. F. E. Fish, “Power Output and Propulsive Efficiency of Swimming Bot-

  tlenose Dolphins ( Tursiops truncatus),”  Journal of Experimental Biology 185

  (1993): 179–93.

  6.  U. M.  Norberg,  “Flying,  Gliding,  Soaring,”  in  Functional Vertebrate

  Morphology, ed. M. Hildebrand, D. M. Bramble, K. F. Liem, and D. B. Wake

  (Cambridge, MA: Belknap Press, 1985), 129–58.

  7. P. W. Webb and R. W. Blake, “Swimming,” in  Functional Vertebrate Mor-

  phology, ed. M. Hildebrand, D. M. Bramble, K. F. Liem, and D. B. Wake (Cam-

  bridge, MA: Belknap Press, 1985), 110–28.

  8. Humans do create lift with their feet when doing the butterfly stroke.

  9.  F. E.  Fish, “Kinematics  and  Estimated  Thrust  Production  of  Swimming

  Harp and Ringed Seals,”  Journal of Experimental Biology 137 (1988): 157–73.

  10. A. W. English, “Limb Movements and Locomotor Function in the Cali-

  fornia Sea Lion,”  Journal of Zoology, London, 178 (1976): 341–64. F. E. Fish,

  “Influence  of  Hydrodynamic  Design  and  Propulsive  Mode  on  Mammalian

  Swimming Energetics,”  Australian Journal of Zoology 42 (1993): 79–101.

  11. G. C. Hickman, “Swimming Ability in Talpid Moles, with Particular Ref-

  erence  to  the  Semi-Aquatic  Mole  Condylura cristata,”  Mammalia  48  (1984): 505–13.

  12.  F. E.  Fish,  “Transitions  from  Drag-Based  to  Lift-Based  Propulsion  in

  Mammalian Swimming,”  American Zoologist 36 (1996): 628–41.

  13.  T. M.  Williams,  “Locomotion  in  the  North  American  Mink,  a  Semi-

  Aquatic Mammal, I: Swimming Energetics and Body Drag,”  Journal of Experi-

  mental Zoology 103 (1983): 155–68.

  14. F. E. Fish, “Association of Propulsive Mode with Behavior in River Otters

  ( Lutra canadensis),”  Journal of Mammalogy 75 (1994): 989–97.

  15. J. G. M. Thewissen and F. E. Fish, “Locomotor Evolution in the Earliest

  Cetaceans:  Functional  Model,  Modern  Analogues,  and  Paleontological  Evi-

  dence,”  Paleobiology 23 (1997): 482–490.

  16. S. Bajpai and J. G. M. Thewissen, “A New, Diminuitive Whale from Kach-

  chh  (Gujarat,  India)  and  Its  Implications  for  Locomotor  Evolution  of  Ceta-

  ceans,”  Current Science (New Delhi) 79 (2000): 1478–82. J. G. M. Thewissen

  Notes    |    219

  and  S.  Bajpai,  “New  Skeletal  Material  for  Andrewsiphius  and  Kutchicetus,

  Two  Eocene  Cetaceans  from  India,”  Journal of Paleontology  83  (2009):

  635–63.

  17. P. D. Gingerich, “Land-to-Sea Transition in Early Whales: Evolution of

  Eocene Archaeoceti (Cetacea) in Relation to Skeletal Proportions and Locomo-

  tion of Living Semiaquatic Mammals,”  Paleobiology 29 (2003): 429–54.

  18. E. A. Buchholtz, “Implications of Vertebral Morphology for Locomotor

  Evolution in Early Cetacea,” in  The Emergence of Whales: Evolutionary Pat-

  terns in the Origin of Cetacea, ed. J. G. M. Thewissen (New York, NY: Plenum

  Press, 1998), 325–52.

  19. R. Dehm and T. zu Oettingen-Spielberg, “Palaeontologische und geolo-

  gische Untersuchungen im Tertiaer von Pakistan, 2: Die mitteleozaenen Sauege-

  tiere von Ganda Kas bei Basal in Nord-West Pakistan,”  Abhandlungen der Bay-

  erischen Akademie der Wissenschaften, Mathematisch.-Naturwissenschaftliche

  Klasse 91 (1958): 1–53.

  20. S. Bajpai and P. D. Gingerich, “A New Archaeocete (Mammalia, Cetacea)

  from  India  and  the Time  of  Origin  of Whales,”  Proceedings of the National

  Academy of Sciences 95 (1998): 15464–68.

  21. J. G. M. Thewissen, E. M. Williams, and S. T. Hussain, “Eocene Mammal

  Faunas from Northern Indo-Pakistan,”  Journal of Vertebrate Paleontology 21

  (2001): 347–66.

  22.  K. K.  Smith, “The  Evolution  of  the  Mammalian  Pharynx,”  Zoological

  Journal of the Linnean Society 104 (1992): 313–49.

  23. J. S. Reidenberg and J. T. Laitman, “Anatomy of the Hyoid Apparatus in

  Odontoceti (Toothed Whales): Specializations of their Skeleton and Muscula-

  ture Compared with Those of Terrestrial Mammals,”  Anatomical Record 240

  (1994): 598–624.

  24. The hyoid of humans is a single bone, located in the midline of the neck,

  but embryologically, it consists of three bones. In most mammals, there are even

  more: a dog has nine, for instance.

  25. E. J. Slijper,  Whales (New York, NY: Basic Books, 1962).

  26. S. Nummela, S. T. Hussain, and J. G. M. Thewissen, “Cranial Anatomy of

  Pakicetidae  (Cetacea,  Mammalia) , ”  Journal of Vertebrate Paleontology  26

  (2006): 746–59.

  27. B. Møhl, W. W. L. Au, J. Pawloski, and P. E. Nachtigall, 1999, “Dolphin

  Hearing: Relative Sensitivity as a Function of Point of Application of a Contact

  Sound Source in the Jaw and Head Region,”  Journal of the Acoustical Society of

  America 105 (1999): 3421–24.

  28. S. Nummela, J. G. M. Thewissen,  S.  Bajpai, T. Hussain,  and K. Kumar,

  “Sound Transmission in Archaic and Modern Whales: Anatomical Adaptations

  for  Underwater  Hearing,”  Anatomical Record  290  (2007):716–33.  S.  Num-

  mela, J. G. M. Thewissen, S. Bajpai, S. T. Hussain, and K. K. Kumar, “Eocene Evo-

  lution of Whale Hearing,”  Nature 430 (2004): 776–78.

  29. S. I. Madar, J. G. M. Thewissen, and S. T. Hussain, “Additional Holotype

  Remains of  Ambulocetus natans (Cetacea, Ambulocetidae), and Their Implica-

  tions for Locomotion in Early Whales,”  Journal of Vertebrate Paleontology 22

  (2002): 405–22.

  220    |    Notes

  30. Y. Narita and S. Kuratani, “Evolution of the Vertebral Formulae in Mam-

  mals: A Perspective on Developmental Constraints,”  Journal of Experimental

  Zoology Part B: Molecular and Developmental Evolution 15 (2005): 91–106. J.

  Müller, T. M.  Scheyer,  J. J.  Head,  P.M.  Barrett,  I. Werneburg,  P. G.  Ericson,  D.

  Polly,  and  M. R.  Sánchez-Villagra,
 “Homeotic  Effects,  Somitogenesis  and  the

  Evolution of Vertebral Numbers in Recent and Fossil Amniotes,”  Proceedings of

  the National Academy of Sciences 107 (2010): 2118–23.

  31. M. M. Moran, S. Bajpai, J. C. George, R. Suydam, S. Usip, and J. G. M.

  Thewissen, “Intervertebral and Epiphyseal Fusion in the Postnatal Ontogeny of

  Cetaceans and Terrestrial Mammals,”  Journal of Mammalian Evolution (2014),

  doi:10.1007/s10914–014–9256–7.

  32. J. G. M. Thewissen, S. I. Madar, and S. T. Hussain, “Ambulocetus natans,

  an Eocene Cetacean (Mammalia) from Pakistan,”  Courier Forschungs.-Institut

  Senckenberg  190  (1996):  1–86.  L. J.  Roe,  J. G. M.  Thewissen,  J.  Quade,  J. R.

  O’Neil, S. Bajpai, A. Sahni, and S. T. Hussain, “Isotopic Approaches to Under-

  standing  the  Terrestrial  to  Marine  Transition  of  the  Earliest  Cetaceans,”  in

  The Emergence of Whales: Evolutionary Patterns in the Origin of Cetacea,

  ed.  J. G. M.  Thewissen  (New  York,  NY:  Plenum  Press,  1998),  399–421.  S. I.

  Madar, J. G. M. Thewissen, and S. T. Hussain, “Additional Holotype Remains of

  Ambulocetus natans  (Cetacea,  Ambulocetidae),  and  their  Implications  for

  Locomotion in Early Whales,”  Journal of Vertebrate Paleontology  22 (2002):

  405–22.

  33. D. Gish, “When Is a Whale a Whale?”  Acts & Facts 23 (1994, No. 4).

  http://www.icr.org/article/when-whale-whale/.

  34.  K.  Miller,  Finding Darwin’s God: A Scientist’s Search for Common

  Ground between God and Evolution (New York, NY: HarperCollins, 1999).

  35. L. Van Valen, “Deltatheridia: A New Order of Mammals,”  Bulletin of the

  American Museum of Natural History 132 (1966): 1–126.

  36. M. Goodman, J. Czelusniak, and J. E. Beeber, “Phylogeny of the Primates

  and  Other  Eutherian  Orders:  A  Cladistics  Analysis  Using  Amino  Acids  and

  Nucleotide Sequence Data,”  Cladistics 1 (1985): 171–85.

  chapter 5. when the mountains grew

  1. The term  Himalayas is used in two different senses. It refers loosely to all

  the mountains on the northern side of India, Pakistan, and Bangladesh. More

  specifically, it refers to one particular mountain range in that area, with a geo-

  logical history that is very different from the others.

  2.  University  of  California  Museum  of  Paleontology,  “Alfred  Wegener

  (1880–1930),” http://www.ucmp.berkeley.edu/history/wegener.html.

  3. G. E. Pilgrim, “Middle Eocene Mammals from Northwest India,”  Proceed-

  ings of the Zoological Society 110 (1940): 124–52.

  4.  R.  Dehm  and  T.  zu  Oettingen-Spielberg, “Paläontologische  und  geolo-

  gische Untersuchungen im Tertiär von Pakistan, 2: Die mitteleozänen Säugetiere

  von  Ganda  Kas  bei  Basal  in  Northwest  Pakistan,”  Abhandlungen der Bayer-

  ischen Akademie der Wissenschaften, Mathematisch.-Naturwissenschaftliche

  Klasse 91 (1958): 1–54.

  Notes    |    221

  5. R. M. West, “Middle  Eocene  Large Mammal Assemblage with Tethyan

  Affinities,  Ganda  Kas  Region,  Pakistan,”  Journal of Paleontology  54  (1980):

  508–33.

  6.  J. G. M.  Thewissen,  S. I.  Madar,  and  S. T.  Hussain,  1996, “Ambulocetus

  natans,  an Eocene Cetacean (Mammalia) from Pakistan,”  Courier Forschungs-

  Institut Senckenberg  190 (1996):  1–86. Some  years later we  were able  to go

  back  and to  excavate  the remainder  of the  holotype of  Ambulocetus natans.

  Those fossils are described in S. I. Madar, J. G. M. Thewissen, and S. T. Hussain,

  “Additional Holotype Remains of  Ambulocetus natans (Cetacea, Ambuloceti-

  dae), and Their Implications for Locomotion in Early Whales,”  Journal of Ver-

  tebrate Paleontology 22 (2002): 405–22.

  7. A. Sahni, “Enamel Ultrastructure of Fossil Mammalia: Eocene Archaeoceti

  from Kutch,”  Journal of the Palaeontological Society of India 25 (1981): 33–37.

  8. M. C. Maas and J. G. M. Thewissen, “Enamel Microstructure of  Pakicetus

  (Mammalia: Archaeoceti),”  Journal of Paleontology 69 (1995): 1154–63.

  chapter 6. passage to india

  1. Panjab is a state in India; Punjab is a province of Pakistan. When the Brit-

  ish ruled India, these were one; when the country broke into two, the province

  was divided, too.

  2. A. B. Wynne, “Memoir on the Geology of Kutch,”  Memoirs of the Geo-

  logical Survey of India 9 (1872).

  3. A. Sahni and V. P. Mishra, “A New Species of  Protocetus from the Middle

  Eocene of Kutch, Western India,”  Palaeontology 15 (1972): 490–95.

  4.  A.  Sahni  and  V. P.  Mishra,  “Lower  Tertiary  Vertebrates  from  Western

  India,”  Monographs of the Palaeontological Society of India 3 (1975).

  5. R. Kellogg,  A Review of the Archaeoceti (Washington, DC: Carnegie Insti-

  tute of Washington, 1936).

  6.  S.  Bajpai  and  J. G. M.  Thewissen, “Middle  Eocene  Cetaceans  from  the

  Harudi and Subathu Formations of India,” in  The Emergence of Whales: Evo-

  lutionary Patterns in the Origin of Cetacea, ed. J. G. M. Thewissen (New York,

  NY: Plenum Press, 1998), 213–34.

  chapter 7. a trip to the beach

  1. S. K. Biswas, “Tertiary Stratigraphy of Kutch,”  Memoirs of the Geological

  Society of India 10 (1992): 1–29.

  2. S. K. Mukhopadhyay and S. Shome, “Depositional Environment and Basin

  Development  during  Early  Paleaeogene  Lignite  Deposition,  Western  Kutch,

  Gujarat,”  Journal of the Geological Society of India 47 (1996): 579–92.

  chapter 8. the otter whale

  1. S. Bajpai and J. G. M. Thewissen, “A New, Diminuitive Whale from Kach-

  chh (Gujarat, India) and Its Implications for Locomotor Evolution of Cetaceans,”

  Current Science (New Delhi) 79 (2000): 1478–82.

  222    |    Notes

  2.  A.  Sahni  and  V. P.  Mishra,  “Lower  Tertiary  Vertebrates  from  Western

  India,”  Monographs of the Palaeontological Society of India 3 (1975).

  3. K. Kumar and A. Sahni, “Remingtonocetus harudiensis: New Combina-

  tion, a Middle Eocene Archaeocete (Mammalia, Cetacea) from Western Kutch,

  India,”  Journal of Vertebrate Paleontology 6 (1986): 326–49.

  4. P. D. Gingerich, M. Arif
, and W. C. Clyde, “New Archaeocetes (Mammalia,

  Cetacea) from the Middle Eocene Domanda Formation of the Sulaiman Range,

  Punjab, Pakistan,”  Contributions of the Museum of Paleontology, University of

  Michigan 29 (1995): 291–330.

  5.  J. G. M. Thewissen  and  S.  Bajpai, “Dental  Morphology  of  the  Reming-

  tonocetidae (Cetacea, Mammalia),”  Journal of Paleontology 75 (2001): 463–65.

  6.  J. G. M. Thewissen  and  S.  T.  Hussain, “Attockicetus praecursor,   a  New

  Remingtonocetid Cetacean from Marine Eocene Sediments of Pakistan,”  Jour-

  nal of Mammalian Evolution 7 (2000): 133–46.

  7. V.  Ravikant  and  S.  Bajpai, “Strontium  Isotope  Evidence  for  the Age  of

  Eocene  Fossil  Whales  of  Kutch,  Western  India,”  Geological Magazine  147

  (2012): 473–77.

  8. P. D. Gingerich, M. Ul-Haq, W. V. Koenigswald, W. J. Sanders, B. H. Smith,

  and I. S. Zalmout, “New Protocetid Whale from the Middle Eocene of Pakistan:

  Birth on Land, Precicial Development, and Sexual Dimorphism,”  PLoS One 4

  (2009): E4366.

  9. L. N. Cooper, T. L. Hieronymus, C. J. Vinyard, S. Bajpai, and J. G. M. Thewis-

  sen, “Feeding Strategy in Remingtonocetinae (Cetacea, Mammalia) by Constrained

  Ordination,” in  Experimental Approaches to Understanding Fossil Organisms, ed.

  D. I. Hembree, B. F. Platt, and J. J. Smith (Dordrecht, Plenum, 2014), 89–107.

  10. If the teeth had fallen out during life, the space in the jaw (the alveolus)

  that the tooth was anchored in would be filled by new bone.

  11. J. G. M. Thewissen and S. Bajpai, “New Skeletal Material of  Andrews-

  iphius and  Kutchicetus,  Two Eocene Cetaceans from India,”  Journal of Paleon-

  tology 83 (2009): 635–63.

  12. R. Elsner, “Living in Water: Solutions to Physiological Problems,” in  Biol-

  ogy of Marine Mammals, ed. J. E. Reynolds III and S. A. Rommel (Washington,

 

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