Journal of Mammalian Evolution

, Volume 10, Issue 1–2, pp 23–129 | Cite as

Morphological Evidence for the Phylogeny of Cetacea

  • Jonathan H. Geisler
  • Albert E. Sanders
Article

Abstract

A cladistic analysis of 54 extant and extinct cetacean taxa scored for 304 morphological characters supports a monophyletic Odontoceti, Mysticeti, Autoceta, and Cetacea. Forcing a sister-group relationship between Mysticeti and Physeteridae, as suggested by some, but not all, molecular studies, requires an additional 72 steps. In agreement with recent molecular studies, morphological data divide extant mysticetes into two clades: Balaenopteroidea (Eschrichtiidae + Balaenopteridae) and Balaenoidea (Balaenidae + Neobalaenidae). Cetotheriopsinae is removed from Cetotheriidae, elevated to Family Cetotheriopsidae, and placed within the Superfamily Eomysticetoidea. All extant mysticetes and all cetotheriids are placed in a new Parvorder Balaenomorpha, which is diagnosed by many morphological characters, including fusion of the anterior and posterior processes of petrosal to ectotympanic bulla, pronounced median keel on palate, and absence of ventral margin of sigmoid process of bulla. Many of the clades within Odontoceti in the most parsimonious trees of this study are at odds with recent phylogenetic analyses. For example, Platanistidae is not closely related to the extinct odontocete families Squalodontidae and Squalodelphinidae. Instead, it is more closely related to extant river-dwelling odontocetes (i.e., Lipotes, Inia), suggesting a single dispersal of odontocetes into freshwater habitats. We found several characters to support Physeteroidea (Physeteridae + Ziphiidae), a taxon considered paraphyletic by several molecular and some morphological analyses. Lack of agreement on the phylogeny within Odontoceti indicates that additional analyses, which include molecular and anatomical data as well as extant and extinct taxa, are needed.

Cetacea Odontoceti Mysticeti morphology 

Preview

Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.

Literature Cited

  1. Abel, O. (1901). Les dauphins longirostres du Boldérien (Miocène supérieur) des environs d'Anvers. Mem. Mus. Roy, Hist. Natur. Belg. 1: 1–95.Google Scholar
  2. Abel, O. (1902). Les dauphins longirostres du Boldérien (Miocène supérieur) des environs d'Anvers. Mem. Mus. Roy, Hist. Natur. Belg. 2: 1–188.Google Scholar
  3. Abel, O. (1913). Die Vorfahren der Bartenwale. Denkschriften der Akademie der Wissenschaften, Wien, Mathematisch-naturwissenschaftliche Klasse 90: 155–224.Google Scholar
  4. Alexander, L. J., and Beattie, C. W. (1992). The sequence of porcine beta-casein cDNA. Anim. Genet. 23: 369–371.Google Scholar
  5. Allen, G. M. (1921). A new fossil cetacean. Harv. Univ. Bull. Mus. Comp. Zool. 65: 3–15.Google Scholar
  6. Andrews, R. C. (1914). The California gray whale (Rhachianectes glaucas Cope). Mem. Am. Mus. Nat. Hist. 1: 227–287.Google Scholar
  7. Archie, J. W. (1985). Methods for coding variable morphological features for numerical taxonomic analysis. Syst. Zool. 34: 326–345.Google Scholar
  8. Arnason, U., and Gullberg, A. (1994). Relationship of baleen whales established by cytochrome b gene sequence comparison. Nature 367: 726–728.Google Scholar
  9. Baker, A. N. (1985). Pygmy right whale Caperea marginata (Gray, 1846). In: Handbook of Marine Mammals, Vol. 3, S. H. Ridgway and R. Harrison, eds., pp. 345–354, Academic Press, New York.Google Scholar
  10. Balcomb, K. C., III (1989). Baird's beaked whale Berardius bairdii Stejneger, 1883: Arnoux's beaked whale Berardius arnuxii Duvernoy, 1851. In: Handbook of Marine Mammals, Vol. 4, S. H. Ridgway and R. Harrison, eds., pp. 261–288, Academic Press, New York.Google Scholar
  11. Barnes, L. G. (1976). Outline of eastern North Pacific fossil cetacean assemblages. Syst. Zool. 25: 321–344.Google Scholar
  12. Barnes, L. G. (1984). Whales, dolphins and porpoises: Origin and evolution of the Cetacea. In: Mammals. Notes for a Short Course Organized by P. D. Gingerich and C. E. Badgely, T. W. Broadhead, ed., pp. 139–154, Studies in Geology 8 (1–4), University of Tennessee Department of Geological Sciences, Knoxville.Google Scholar
  13. Barnes, L. G. (1985). Fossil pontoporiid dolphins (Mammalia: Cetacea) from the Pacific coast of North America. Nat. Hist. Mus. Los Angel. County Contrib. Sci. 363: 1–34.Google Scholar
  14. Barnes, L. G. (1990). The fossil record and evolutionary relationships of the genus Tursiops. In: The Bottlenosed Dolphin, S. Leatherwood and R. R. Reeves, eds., pp. 3–26, Academic Press, New York.Google Scholar
  15. Barnes, L. G. (2002). An Early Miocene long-snouted marine platanistid dolphin (Mammalia, Cetacea, Odontoceti) from the Korneuberg Basin (Austria) [Das Karpat des Korneuburger Beckens, Vol. 2, edited by Wolfgang Sovis and Brigitta Schmid]. Beitrage zur Paläontologie 27: 407–418.Google Scholar
  16. Barnes, L. G., and Mitchell, E. D. (1978). Cetacea. In: Evolution of African Mammals, V. J. Maglio and H. B. S. Cooke, eds., pp. 582–602. Harvard University Press, Cambridge, MA.Google Scholar
  17. Barnes, L. G., and Sanders, A. E. (1996). The transition from archaeocetes to mysticetes: Late Oligocene toothed mysticetes from near Charleston, South Carolina. In: Sixth North American Paleontological Convention Abstracts of Papers, J. E. Repetski, ed., p. 24, Paleontol. Soc., Washington, D.C., Special Publication No. 8.Google Scholar
  18. Barnes, L. G., Kimura, M., Furusawa, H., and Sawamura, H. (1994). Classification and distribution of Oligocene Aetiocetidae (Mammalia; Cetacea; Mysticeti) from western North America and Japan. Island Arc 3: 392–431.Google Scholar
  19. Barnes, L. G., Goedert, J. L., and Furusawa, H. (2002). The earliest known echolocating toothed whales (Mammalia; Odontoceti): Preliminary observations of fossils from Washington State. Mesa Southwest Mus. Bull. 8: 91–100. [Western Association of Vertebrate Paleontologists with Mesa Southwest Museum and Southwest Paleontological Society, Mesa, Arizona. First Meeting of the new Millenium]Google Scholar
  20. Beneden, P. J. van (1865). Recherches sur les ossements provenant du Crag d'Anvers–Recherches sur les squalodonts. Memoires del' Academie Royal des Sciences, des Lettres et des Beaux-arts de Belgique,Series 2, 35: 1–85.Google Scholar
  21. Best, R. C., and da Silva, V. M. F. (1989). Amazon River Dolphin, Boto Inia geoffrensis (de Blainville, 1817). In: Handbook of Marine Mammals, Vol. 4, S. H. Ridgway and R. Harrison, eds., pp. 1–23, Academic Press, New York.Google Scholar
  22. Bremer, K. (1988). The limits of amino acid sequence data in angiosperm phylogenetic reconstruction. Evolution 42: 785–803.Google Scholar
  23. Bremer, K. (1994). Branch support and tree stability. Cladistics 10: 295–304.Google Scholar
  24. Brodie, P. F. (1989). The white whale Delphinapterus leucas (Pallas, 1776). In: Handbook of Marine Mammals, Vol. 4, S. H. Ridgway and R. Harrison, eds., pp. 119–144, Academic Press, New York.Google Scholar
  25. Brownell, R. L., Jr. (1989). Franciscana Pontoporia blainvillei (Gervais and d'Orbigny, 1844). In: Handbook of Marine Mammals, Vol. 4, S. H. Ridgway and R. Harrison, eds., pp. 45–67, Academic Press, New York.Google Scholar
  26. Caldwell, D. K., and Caldwell, M. C. (1989). Pygmy sperm whale Kogia breviceps (de Blainville, 1838): Dwarf sperm whale Kogia simus Owen, 1866. In: Handbook of Marine Mammals, Vol. 4, S. H. Ridgway andR. Harrison, eds., pp. 235–260, Academic Press, New York.Google Scholar
  27. Cappellini, J. (1904). Avanzi di Squalodonte, nella arenaria di Grumi dei Frati presso Schio. Memorie della Regia Accademia delle Scienze all'Instituto di Bologna, 10: 413–419.Google Scholar
  28. Cassens, I., Vicario, S., Waddell, V. G., Balchowsky, H., Van Belle, D., Ding, W., Fan, C., Lal Mohan, R. S., Simões-Lopes, P. C., Bastida, R., Meyer, A., Stanhope, M. J., and Milinkovitch, M. C. (2000). Independent adaptation to riverine habitats allowed survival of ancient cetacean lineages. Proc. Natl. Acad. Sci. USA 97: 11343–11347.Google Scholar
  29. Cerchio, S., and Tucker, P. (1998). Influence of alignment on the mtDNA phylogeny of Cetacea: Questionable support for a Mysticeti/Physeteroidea clade. Syst. Biol. 47: 336–344.Google Scholar
  30. Cranford, T. W., Amundin, M., and Norris, K. S. (1996). Functional morphology and homology in the odontocete nasal complex: Implications for sound generation. J. Morphol. 228: 223–285.Google Scholar
  31. Cummings, W. C. (1985). Right whales Eubalaena glacialis (Müller, 1776) and Eubalaena australis (Desmoulins, 1822). In: Handbook of Marine Mammals, Vol. 3, S. H. Ridgway and R. Harrison, eds., pp. 275–304, Academic Press, New York.Google Scholar
  32. Doran, A. H. G. (1876). Morphology of the mammalian ossicula auditûs. Linn. Soc. Lond. Trans. 2nd Ser. Zool. 1: 371–497.Google Scholar
  33. Dubrovo, I. A., and Sanders, A. E. (2000). A new species of Patriocetus (Mammalia, Cetacea) from the Late Oligocene of Kazakhstan. J. Vertebr. Paleontol. 20: 577–590.Google Scholar
  34. Emlong, D. (1966). A new archaic cetacean from the Oligocene of Northwest Oregon. Bull. Mus. Nat. Hist. Univ. Oregon 3: 1–51.Google Scholar
  35. Evans, H. E. (1993). Miller's Anatomy of the Dog, 3rd edn., W. B. Saunders, Philadelphia.Google Scholar
  36. Farris, J. S. (1983). The logical basis of phylogenetic analysis. In: Advances in Cladistics, Vol. 2, N. I. Platnick and V. A. Funk, eds., pp. 7–36, Columbia University, New York.Google Scholar
  37. Flower, W. H. (1867). Description of the skeleton of Inia geoffrensis and of the skull of Pontoporia blainvillii with remarks on the systematic position of these animals in the Order Cetacea. Trans. Zool. Soc. Lond. 6: 87–116.Google Scholar
  38. Flower, W. H. (1869). On the osteology of the cachalot or sperm-whale (Physeter macrocephalus). Trans. Zool. Soc. London 6: 309–372.Google Scholar
  39. Flower, W. H. (1872). On the recent ziphioid whales, with a description of the skeleton of Berardius arnouxi. Trans. Zool. Soc. Lond. 8: 203–234.Google Scholar
  40. Flynn, T. T. (1948). Description of Prosqualodon davidi Flynn, a fossil cetacean from Tasmania. Trans. Zool. Soc. 26: 153–197.Google Scholar
  41. Fordyce, R. E. (1977). The development of the circum-antarctic current and the evolution of the Mysticeti (Mammalia: Cetacea). Palaeogeog. Palaeoclimat. Palaeoecol. 21: 265–271.Google Scholar
  42. Fordyce, R. E. (1981). Systematics of the odontocete whale Agorophius pygmaeus and the family Agorophiidae (Mammalia, Cetacea). J. Paleontol. 55: 1028–1045.Google Scholar
  43. Fordyce, R. E. (1983). Rhabdosteid dolphins (Mammalia: Cetacea) from the middle Miocene, Lake Frome area, South Australia. Alcheringa 7: 27–40.Google Scholar
  44. Fordyce, R. E. (1994). Waipatia maerewhenua, new genus and new species (Waipatiidae, new family), an archaic late Oligocene dolphin (Cetacea: Odontoceti: Platanistoidea) from New Zealand. In: Contributions in Marine Mammal Paleontology Honoring Frank Whitmore Jr., A. Berta and T. A. Deméré, eds., pp. 147–176, Proc. San Diego Soc. Nat. Hist. 29.Google Scholar
  45. Fordyce, R. E. (2002). Simocetus rayi (Odontoceti: Simocetidae, New Family): A bizarre new archaic Oligocene dolphin from the Eastern Pacific. In: Cenozoic Mammals of Land and Sea, Tributes to the Career of Clayton E. Ray, R. E. Emry, ed., pp. 185–222, Smithson. Contrib. Paleobiology 93.Google Scholar
  46. Fordyce, R. E., and Barnes, L. G. (1994). The evolutionary history of whales and dolphins. Annu. Rev. Earth Planet. Sci. 1994, 22: 419–455.Google Scholar
  47. Fordyce, R. E., and Muizon, C. de (2001). Evolutionary history of cetaceans: A review. In: Secondary Adaptations of Tetrapods to Life in Water, J.-M. Mazin and V. de Buffrénil, eds., pp. 169–233, Verlag Dr. Friedrich Pfeil, München, Germany.Google Scholar
  48. Fordyce, R. E., Quilty, P. G., and Daniels, J. (2002). Australodelphis mirus, a bizarre new toothless ziphiid-like fossil dolphin (Cetacea: Delphinidae) from the Pliocene of Vestfold Hills, East Antarctica. Antarctic Sci. 14: 37–54.Google Scholar
  49. Fraser, F. C., and Purves, P. E. (1960). Hearing in cetaceans–Evolution of the accessory air sacs and the structure and function of the outer and middle ear in recent cetaceans. Bull. Br. Mus. (Nat. Hist.) Zool. 7: 1–140.Google Scholar
  50. Gambell, R. (1985). Fin whale Balaenoptera physalus (Linnaeus, 1758). In: Handbook of Marine Mammals,Vol. 3, S. H. Ridgway and R. Harrison, eds., pp. 171–192, Academic Press, New York.Google Scholar
  51. Gatesy, J. (1997). More DNA support for a Cetacea/Hippopotamidae clade: The blood-clotting protein gamma-fibrinogen. Mol. Biol. Evol. 14: 537–543.Google Scholar
  52. Gatesy, J. (1998). Molecular evidence for the phylogenetic affinities of Cetacea. In: The Emergence of Whales,J. G. M. Thewissen, ed., pp. 63–111, Plenum, New York.Google Scholar
  53. Gatesy, J., Hayashi, C., Cronin, A., and Arctander, P. (1996). Evidence from milk casein genes that cetaceans are close relatives of hippopotamid artiodactyls. Mol. Biol. Evol. 13: 954–963.Google Scholar
  54. Gatesy, J., Milinkovitch, M., Waddell, V., and Stanhope, M. (1999). Stability of cladistic relationships between Cetacea and higher-level artiodactyl taxa. Syst. Biol. 48: 6–20.Google Scholar
  55. Geisler, J. H. (2001a). New morphological evidence for the phylogeny of Artiodactyla, Cetacea, and Mesonychidae. Amer. Mus. Novit. 3344: 1–53.Google Scholar
  56. Geisler, J. H. (2001b). Morphological and Molecular Evidence for the Phylogeny of Cetacea and Artiodactyla: Explaining Incongruence Between Types of Data, Unpublished Ph.D. Dissertation, Columbia University, New York.Google Scholar
  57. Geisler, J. H., and Luo, Z. (1996). The petrosal and inner ear of Herpetocetus sp. (Mammalia: Cetacea) and their implications for the phylogeny and hearing of archaic mysticetes. J. Paleontol. 70: 1045–1066.Google Scholar
  58. Geisler, J. H., and Luo, Z. (1998). Relationships of Cetacea to terrestrial ungulates and the evolution of cranial vasculature in Cete. In: The Emergence of Whales, J. G. M. Thewissen, ed., pp. 163–212, Plenum, New York.Google Scholar
  59. Geisler, J. H., Sanders, A. E., and Luo, Z. (1996). A new protocetid cetacean from the Eocene of South Carolina, U.S.A.: Phylogenetic and biogeographic implications. Paleontol. Soc. Spec. Publ. 8: 139.Google Scholar
  60. Gingerich, P. D., and Russell, D. E. (1981). Pakicetus inachus, a new archaeocete (Mammalia, Cetacea) from the early-middle Eocene Kuldana Formation of Kohat (Pakistan). Contrib. Mus. Paleontol. Univ. Mich. 25: 235–246.Google Scholar
  61. 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 368: 844–847.Google Scholar
  62. Gingerich, P. D., Arif, M., and Clyde, W. C. (1995). New archaeocetes (Mammalia, Cetacea) from the middle Eocene Domanda Formation of the Sulaiman Range; Punjab (Pakistan). Contrib. Mus. Paleontol. Univ. Mich. 29: 291–230.Google Scholar
  63. Gingerich, P. D., Haq, M. u., Zalmout, I. S., Khan, I. H., and Malakani, M. S. (2001). Origin of whales from early artiodactyls: Hands and feet of Eocene Protocetidae from Pakistan. Science 293: 2239–2242.Google Scholar
  64. Goloboff, P. (1993). NONA Version 1.9. Computer program and documentation http://www.cladistics.com/about_nono.htm.Google Scholar
  65. Gottfried, M. D., Bohaska, D. J., and Whitmore, F. C., Jr. (1994). Miocene cetaceans of the Chesapeake Group. In: Contributions in Marine Mammal Paleontology Honoring Frank Whitmore Jr., A. Berta and T. A. Deméré, eds., pp. 229–238, Proc. San Diego Soc. Nat. Hist. 29.Google Scholar
  66. Gray, J. E. (1868). Synopsis of the Species of Whales and Dolphins in the Collection of the British Museum, Bernard Quaritch, London.Google Scholar
  67. Graybeal, A. (1998). Is it better to add taxa or characters to a difficult phylogenetic problem. Syst. Biol. 47: 9–17.Google Scholar
  68. Gregory, W. K., and Kellogg, R. (1927). A fossil porpoise from California. Am. Mus. Novitates 269: 1–7.Google Scholar
  69. Hanke, H. von (1914). Ein Beitrag zur Kenntnis der Anatomie des äusseren und mittlere Ohres der Bartenwale. Jena. Z. Naturwiss. 3: 487–524.Google Scholar
  70. Hendy, M. D., and Penny, D. (1989). A framework for the quantitative study of evolutionary trees. Syst. Zool. 38: 297–309.Google Scholar
  71. Heyning, J. E. (1989a). Comparative facial anatomy of beaked whales (Ziphiidae) and a systematic revision among the families of extant Odontoceti. Nat. Hist. Mus. Los Angle. Cty. Contrib. Sci. 405: 1–64.Google Scholar
  72. Heyning, J. E. (1989b). Cuvier's beaked whale Ziphius cavirostris G. Cuvier. In: Handbook of Marine Mammals, Vol. 4, S. H. Ridgway and R. Harrison, eds., pp. 289–308, Academic Press, New York.Google Scholar
  73. Heyning, J. E. (1997). Sperm whale phylogeny revisted: Analysis of the morphological evidence. Mar. Mammal Sci. 13: 596–613.Google Scholar
  74. Heyning, J. E., and Mead, J. G. (1990). Evolution of the nasal anatomy of cetaceans. In: Sensory Abilities of Cetaceans, J. Thomas and R. Kastelein, eds., pp. 67–79, Plenum Press, New York.Google Scholar
  75. Hillis, D. M. (1998). Taxonomic sampling, phylogenetic accuracy, and investigator bias. Syst. Biol. 47: 3–8.Google Scholar
  76. Hulbert, R. C., Jr. (1998). Postcranial osteology of the North American Middle Eocene protocetid Georgiacetus. In: The Emergence of Whales, J. G. M. Thewissen, ed., pp. 235–267, Plenum, New York.Google Scholar
  77. 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: 907–927.Google Scholar
  78. Irwin, D. M., and árnason, ú. (1994). Cytochrome b gene of marine mammals: Phylogeny and evolution. J. Mammal. Evol. 2: 37–55.Google Scholar
  79. Kasuya, T. (1973). Systematic consideration of recent toothed whales based on the morphology of tympano-periotic bone. Sci. Rep. Whales Res. Inst. 25: 1–103.Google Scholar
  80. Kellogg, A. R. (1923a). Description of an apparently new toothed cetacean from South Carolina. Smithson. Misc. Coll. 76: 1–7.Google Scholar
  81. Kellogg, A. R. (1923b). Description of two squalodonts recently discovered in the Calvert Cliffs, Maryland; and notes on the shark-toothed cetaceans. Proc. U.S. Natl. Mus. 62: 1–69.Google Scholar
  82. Kellogg, A. R. (1924). A fossil porpoise from the Calvert Formation of Maryland. Proc. U.S. Natl. Mus. 63: 1–39.Google Scholar
  83. Kellogg, A. R. (1925). On the occurrence of remains of fossil porpoises of the genus Eurhinodelphis in North America. Proc. U.S. Natl. Mus. 66: 1–40.Google Scholar
  84. Kellogg, A. R. (1926). Supplementary observations on the skull of the fossil porpoise Zarhachis flagellator Cope. Proc. U.S. Natl. Mus. 67: 1–18.Google Scholar
  85. Kellogg, A. R. (1927). Kentriodon pernix, a Miocene porpoise from Maryland. Proc.U.S. Natl Mus. 69(19): 1–55.Google Scholar
  86. Kellogg, A. R. (1928). The history of whales–Their adaptation to life in the water. Q. Rev. Biol. 3: 29–76, 174–208.Google Scholar
  87. Kellogg, A. R. (1932). A Miocene long-beaked porpoise from California. Smithson. Misc. Coll. 87: 1–11.Google Scholar
  88. Kellogg, A. R. (1936). A review of the Archaeoceti. Carnegie Inst. Washington Publ. 482: 1–366.Google Scholar
  89. Kellogg, A. R. (1965). Fossil marine mammals from the Miocene Calvert Formation of Maryland and Virginia. U.S. Natl. Mus. Bull. 247: 1–63.Google Scholar
  90. Kellogg, A. R. (1968). Fossil marine mammals from the Miocene Calvert Formation of Maryland and Virginia. U.S. Natl. Mus. Bull. 247: 103–201.Google Scholar
  91. Kim, J. (1998). Large-scale phylogenies and measuring the performance of phylogenetic estimators. Syst. Biol. 47: 43–60.Google Scholar
  92. Kimura, T., and Ozawa, T. (2002). A new cetothere (Cetacea: Mysticeti) from the Early Miocene of Japan. J. Vertebr. Paleontol. 22: 684–702.Google Scholar
  93. Kleinenberg, S. E. (1958). K voprosu o proiskhozhdenii kitoobraznykh. Doklady Akademia Nauk S. S. S. R. 122: 950–952.Google Scholar
  94. Kleinenberg, S. E. (1959). On the origin of Cetacea. Proc. XV Int. Cong. Zool. 122: 445–447.Google Scholar
  95. Kluge, A. G. (1997). Testability and the refutation and corroboration of cladistic hypotheses. Cladistics 13: 81–96.Google Scholar
  96. Lancaster, W. C. (1990). The middle ear of the Archaeoceti. J. Vertebr. Paleontol. 10: 117–127.Google Scholar
  97. Leidy, J. (1869). The extinct mammalian fauna of Dakota and Nebraska. J. Acad. Nat. Sci. Phila. 7: 1–472.Google Scholar
  98. Lillie, D. G. L. (1910). Observations on the anatomy and general biology of some members of the larger Cetacea. Proc. Zool. Soc. (Lond.) 2: 769–791.Google Scholar
  99. Lipscomb, D. L. (1992). Parsimony, homology, and the analysis of multistate characters. Cladistics 8: 45–65.Google Scholar
  100. Luo, Z. (1998). Homology and transformation of cetacean ectotympanic structures. In: The Emergence of Whales, J. G. M. Thewissen, ed., pp. 269–301, Plenum, New York.Google Scholar
  101. Luo, Z., Crompton, A. W., and Lucas, S. G. (1995). Evolutionary origins of the mammalian promontorium and cochlea. J. Vertebr. Paleontol. 15: 113–121.Google Scholar
  102. Luo, Z., and Eastman, E. R. (1995). Petrosal and inner ear of a squalodontoid whale: Implications for evolution of hearing in odontocetes. J. Vertebr. Paleontol. 15: 431–442.Google Scholar
  103. Luo, Z., and Gingerich, P. D. (1999). Terrestrial Mesonychia to aquatic Cetacea: Transformation of the basicranium and evolution of hearing in whales. Univ. Michigan Papers Paleontol. 31: 1–98.Google Scholar
  104. Luo, Z., and Marsh, K. (1996). Petrosal (periotic) and inner ear of a Pliocene kogiine whale (Kogiinae, Odontoceti): Implications on relationships and hearing evolution of toothed whales. J. Vertebr. Paleontol. 16: 328–348.Google Scholar
  105. MacPhee, R. D. E. (1981). Auditory regions of primates and eutherian insectivores: Morphology, ontogeny, and character analysis. Contrib. Primatol. 18: 1–282.Google Scholar
  106. MacPhee, R. D. E. (1994). Morphology, adaptations, and relationships of Plesiorycteropus, and a diagnosis of a new order of eutherian mammals. Bull. Amer. Mus. Nat. Hist. 220: 1–214.Google Scholar
  107. Maddison, W. P., and Maddison, D. R. (1992). MacClade Program (3.01), Sinauer and Associates, Sutherland, MA.Google Scholar
  108. McKenna, M. C., and Bell, S. K. (1997). Classification of Mammals Above the Species Level, Columbia University Press, New York.Google Scholar
  109. McLeod, S. A., Whitmore, F. C., Jr., and Barnes, L. G. (1993). Evolutionary relationships and classification. In The Bowhead Whale, J. J. Burns, J. J. Montague, and C. J. Cowles, eds., pp. 45–70, Society for Marine Mammalogy Special Publication No. 2.Google Scholar
  110. Mead, J. G. (1975). Anatomy of the external nasal passage and facial complex in the Delphinidae (Mammalia: Cetacea). Smithson. Contrib. Zool. 207: 1–72.Google Scholar
  111. Mead, J. G. (1989a). Shepherd's beaked whale Tasmacetus shepherdi Oliver, 1937. In: Handbook of Marine Mammals, Vol. 4, S. H. Ridgway and R. Harrison, eds., pp. 309–320, Academic Press, New York.Google Scholar
  112. Mead, J. G. (1989b). Beaked whales of the genus Mesoplodon. In: Handbook of Marine Mammals, Vol. 4, S. H. Ridgway and R. Harrison, eds., pp. 349–430, Academic Press, New York.Google Scholar
  113. Messenger, S. L., and McGuire, J. A. (1998). Morphology, molecules, and the phylogenetics of cetaceans. Syst. Biol. 47: 90–124.Google Scholar
  114. Mickevich, M. F., and Johnson, M. S. (1976). Congruence between morphological and allozyme data in evolutionary inference and character evolution. Syst. Zool. 25: 260–270.Google Scholar
  115. Milinkovitch, M. C., Orti, G., and Meyer, A. (1993). Revised phylogeny of whales suggested by mitochondrial ribosomal DNA sequences. Science 12: 346–348.Google Scholar
  116. Milinkovitch, M. C., Meyer, A., and Powell, J. R. (1994). Phylogeny of all major groups of cetaceans based on DNA sequences from three mitochondrial genes. Mol. Biol. Evol. 11: 939–948.Google Scholar
  117. Milinkovitch, M. C., Orti, G., and Meyer, A. (1995). Novel phylogeny of whales revisited but not revised. Mol. Biol. Evol. 12: 518–520.Google Scholar
  118. Miller, G. S., Jr. (1918). A new river-dolphin from China. Smithson. Misc. Coll. 68: 1–12.Google Scholar
  119. Miller, G. S., Jr. (1923). The telescoping of the cetacean skull. Smithson. Misc. Coll. 75: 1–55.Google Scholar
  120. Mitchell, E. (1989). A new cetacean from the late Eocene La Mesta Formation, Seymour Island, Antarctic Peninsula. Can. J. Fish. Aquat. Sci. 46: 2219–2235.Google Scholar
  121. Montgelard, C., F. M. Catzeflis, and Douzery, E. (1997). Phylogenetic relationships of artiodactyls and cetaceans as deduced from the comparison of cytochrome b and 12S RNA mitochondrial sequences. Mol. Biol. Evol. 14: 550–559.Google Scholar
  122. Morgan, G. S. (1994). Miocene and Pliocene marine mammal faunas from the Bone Valley Formation of central Florida. In: Contributions in Marine Mammal Paleontology Honoring Frank C. Whitmore, Jr., A. Berta and T. A. Deméré, eds., pp. 239–268, Proc. San Diego Soc. Nat. Hist. 29.Google Scholar
  123. Moore, J. C. (1968). Relationships among the living genera of beaked whales. Fieldiana Zool. 53: 209–298.Google Scholar
  124. Muizon, C. de (1984). Les vertébrés fossiles de la Formation Pisco (Pérou). deuxiéme partie: les Odontocétes (Cetacea, Mammalia) du Pliocéne inférieur de Sud-Sacaco. Trav. Inst. Fr. Et. Andines 27: 1–188.Google Scholar
  125. Muizon, C. de (1987). The affinities of Notocetus vanbenedeni, an early Miocene platanistoid (Cetacea, Mammalia) from Patagonia, southern Argentina. Am. Mus. Novit. 2904: 1–27.Google Scholar
  126. Muizon, C. de (1988a). Les relations phylogénétiques des Delphinida (Cetacea, Mammalia). Ann. Paléontol. 74: 159–227.Google Scholar
  127. Muizon, C. de (1988b). Les vertébrés fossiles de la Formation Pisco (Pérou) III: les odontocétes (Cetacea, Mammalia) du Miocéne. Rech. sur Civ., mem. 78: 1–244.Google Scholar
  128. Muizon, C. de (1991). A new Ziphiidae (Cetacea) from the early Miocene of Washington State (USA) and phylogenetic analysis of the major groups of odontocetes. Bull. Mus. Natn. Hist. Nat., Section C, 4ème série 12: 279–326.Google Scholar
  129. Muizon, C. de (1994). Are squalodonts related to the platanistoids? In: Contributions in Marine Mammal Paleontology Honoring Frank C. Whitmore, Jr., A. Berta and T. A. Deméré, eds., pp. 35–93, Proc. San Diego Soc. Nat. Hist. 29.Google Scholar
  130. 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. U.S.A. 96: 10261–10266.Google Scholar
  131. Nikaido, M., Matsuno, F., Hamilton, H., Brownell, R. L., Jr., Cao, Y., Ding, W., Zuoyan, Z., Shedlock, A. M., Fordyce, R. E., Hasegawa, M., and Okada, N. (2001). Retroposon analysis of major cetacean lineages: The monophyly of toothed whales and paraphyly of river dolphins. Proc. Natl. Acad. Sci. U.S.A. 98: 7384–7389.Google Scholar
  132. Nixon, K. C. (1999). The parsimony ratchet, a new method for rapid parsimony analysis. Cladistics 15: 407–414.Google Scholar
  133. O'Leary, M. A., and Geisler, J. H. 1999. The position of Cetacea within Mammalia: Phylogenetic analysis of morphological data from extinct and extant taxa. Syst. Biol. 48: 455–490.Google Scholar
  134. Omura, H., Nishiwaki, M., Ichihara, T., and Kasuya, T. (1962). Osteological note of a sperm whale. Sci. Rep. Whales Res. Inst. 16: 35–45.Google Scholar
  135. Omura, H., Ohsumi, S., Nemoto, T., Nasu, K., and Kasuya, T. (1969). Black right whales in the North Pacific. Sci. Rep. Whales Res. Inst. 21: 1–78.Google Scholar
  136. Peixun, C. (1989). Lipotes vexillifer Miller, 1918. In: Handbook of Marine Mammals, Vol. 4, S. H. Ridgway and R. Harrison, eds., pp. 25–43, Academic Press, New York.Google Scholar
  137. Pilleri, G., and Gihr, M. (1976). The function and osteology of the manus of Platanista gangetica and Platanista indi. Invest. Cetacea 7: 109–118.Google Scholar
  138. Popper, K. L. (1968). The Logic of Scientific Discovery. Harper and Row Publishers, New York.Google Scholar
  139. Reeves, R. R., and Brownell, R. L., Jr. (1989). Platanista gangetica (Roxburgh, 1801) and Platanista minor Owev, 1853. In: Handbook of Marine Mammals, Vol. 4, S. H. Ridgway and R. Harrison, eds., pp. 69–99, Academic Press, New York.Google Scholar
  140. Rice, D. W., and Wolman, A. W. (1990). The stomach of Kogia breviceps. J. Mammal. 71: 242–246.Google Scholar
  141. Ridewood, W. G. (1922). Observations on the skull in foetal specimens of whales of the genera Megaptera and Balaenoptera. Philos. Trans. R. Soc. Lond. Ser. B 211: 209–272.Google Scholar
  142. Rommel, S. (1990). Osteology of the bottlenose dolphin. In: The Bottlenose Dolphin, S. Leatherwood and R. R. Reeves, eds., pp. 29–49, Academic Press, San Diego, CA.Google Scholar
  143. Rothausen, K. (1968). Die systematische Stellung de europäischen Squalodontidae (Odontoceti, Mamm.). Pal. Zool. 48: 83–104.Google Scholar
  144. Sanders, A. E. (1980). Excavation of Oligocene marine fossil beds near Charleston, South Carolina. Nat. Geograph. Soc. Res. Rep. 12: 601–621.Google Scholar
  145. Sanders, A. E. (1996). The systematic position of the primitive odontocete Xenorophus sloanii (Mammalia, Cetacea) and two new taxa from the late Oligocene of South Carolina, U.S.A. Paleontol. Soc. Spec. Publ. 8: 338.Google Scholar
  146. Sanders, A. E., and Barnes, L. G. (2002a). Paleontology of the Late Oligocene Ashley and Chandler Bridge Formations of South Carolina, 3: Eomysticetidae, a new family of Oligocene mysticetes (Mammalia: Cetacea). In: Cenozoic Mammals of Land and Sea, Tributes to the Career of Clayton E. Ray, R. E. Emry, ed., pp. 313–356, Smithson. Contrib. Paleobiology 93.Google Scholar
  147. Sanders, A. E., and Barnes, L. G. (2002b). Paleontology of the Late Oligocene Ashley and Chandler Bridge Formations of South Carolina, 2: Micromysticetus rothauseni, a primitive cetotheriid mysticete (Mammalia: Cetacea). In: Cenozoic Mammals of Land and Sea, Tributes to the Career of Clayton E. Ray, R. E. Emry, ed., pp. 271–294, Smithson. Contrib. Paleobiology 93.Google Scholar
  148. Schulte, H. von W. (1917). The skull of Kogia breviceps Blainv. Bull. Amer. Mus. Nat. Hist. 37: 361–404.Google Scholar
  149. Schulte, H. von W., and Smith, M. de F. (1918). The external characters, skeletal muscles, and peripheral nerves of Kogia breviceps (Blainville). Bull. Amer. Mus. Nat. Hist. 38: 7–72.Google Scholar
  150. Shimamura, M., Yasue, H., Ohshima, K., Abe, H., Kato, H. Kishiro, T. Goto, M., Munechika, I., and Okada, N. (1997). Molecular evidence from retroposons that whales form a clade within even-toed ungulates. Nature 388: 666–670.Google Scholar
  151. Simpson, G. G. (1945). The principles of classification and a classification of mammals. Bull. Am. Mus. Nat. Hist. 85: 1–339.Google Scholar
  152. Slijper, E. J. (1936). Die Cetaceen Vergleichend-Anatomische und Systematisch. Capita Zoologica 7: 1–590.Google Scholar
  153. Sorenson, M. D. (1996). TreeRot. University of Michigan, Ann Arbor.Google Scholar
  154. Swofford, D. L. (1993). PAUP: Phylogenetic Analysis Using Parsimony (3.1.1), Privately distributed by Illinois Natural History Survey, Champaign, IL.Google Scholar
  155. Thewissen, J. G. M., and Hussain, S. T. (1998). Systematic review of Pakicetidae, Early and Middle Eocene Cetacea (Mammalia) from Pakistan and India. Bull. Carnegie Mus. Nat. Hist. 34: 220–238.Google Scholar
  156. 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 413: 277–281.Google Scholar
  157. True, F. W. (1904). The whalebone whales of the North Atlantic compared with those occuring in European waters with some observations on the species of the North Pacific. Smithson. Contrib. Knowledge 33: 297–332.Google Scholar
  158. True, F. W. (1910a). Description of a skull and some vertebrae of the fossil cetacean Diochoticus vanbenedeni from Santa Cruz, Patagonia. Bull. Amer. Mus. Nat. Hist. 28: 19–32.Google Scholar
  159. True, F. W. (1910b). An account of the beaked whales of the family Ziphiidae in the collection of the United States National Museum, with remarks on some specimens in other American museums. Bull. U.S. Natl. Mus. 73: 1–89.Google Scholar
  160. Uhen, M. D. (1998). Middle to late Eocene Basilosaurines and Dorudontines. In: The Emergence of Whales, J. G. M. Thewissen, ed., pp. 29–61, Plenum Press, New York.Google Scholar
  161. Uhen, M. D. (1999). New species of protocetid archaeocete whale, Eocetus wardii (Mammalia: Cetacea) from the middle Eocene of North Carolina. J. Paleontol. 73: 512–528.Google Scholar
  162. Uhen, M. D. (2000). Replacement of deciduous first premolars and dental eruption in archaeocete whales. J. Mammal. 81: 123–133.Google Scholar
  163. Uhen, M. D., and Gingerich, P. D. (2001). New genus of dorudontine archaeocete (Cetacea) from the middle-to-late Eocene of South Carolina. Marine Mammal Sci. 17: 1–34.Google Scholar
  164. van Kampen, P. N. (1905). Die Tympanalgegend des Säugetierschädel. Morphol. Jahrb. 34: 321–722.Google Scholar
  165. Van Valen, L. (1968). Monophyly or diphyly in the origin of whales. Evolution 22: 37–41.Google Scholar
  166. Waddell, V. G., Milinkovitch, M. C., M. Bérubé, M., and Stanhope, M. J. (2000). Molecular phylogenetic examination of the Delphinoidea trichotomy: Congruent evidence from three nuclear loci inicates that porpoises (Phocoenidae) share a more recent common ancestry with white whales (Monodontidae) than they do with true dolphins (Delphinidae). Mol. Phylogenet. Evol. 15: 314–318.Google Scholar
  167. Whitmore, F. C. (1994). Neogene climatic change and the emergence of the modern whale fauna of the north atlantic ocean. In: Contributions in Marine Mammal Paleontology Honoring Frank Whitmore Jr., A. Berta and T. A. Deméré, eds., pp. 223–227, Proc. San Diego Soc. Nat. Hist. 29.Google Scholar
  168. Whitmore, F. C., and Sanders, A. E. (1977). Review of the Oligocene Cetacea. Syst. Zool. 25: 304–320.Google Scholar
  169. Wible, J. R. (1984). The Ontogeny and Phylogeny of the Mammalian Cranial Pattern, Ph.D. Dissertation, Duke University, Durham.Google Scholar
  170. Wiens, J. J. (2001). Character analysis in morphological phylogenetics: Problems and solutions. Syst. Biol. 50: 689–699.Google Scholar
  171. Wilkinson, M. (1992). Ordered versus unordered characters. Cladistics 8: 375–385.Google Scholar
  172. Wilson, L. E. (1935). Miocene marine mammals from the Bakersfield region, California. Peabody Mus. Natur. Hist. Yale Bull. 4: 1–143.Google Scholar
  173. Wolman, A. A. (1985). Gray whale Eschrichtius robustus (Lilljeborg, 1861). In: Handbook of Marine Mammals, Vol. 3, S. H. Ridgway and R. Harrison, eds., pp. 67–90, Academic Press, New York.Google Scholar
  174. Yablokov, A. V. (1964). Covergence or parallelism in the evolution of cetaceans. Int. Geol. Rev. 7: 1461–1468.Google Scholar
  175. Zeigler, C.V., Chan, G. L., Barnes, L.G. (1997). A new late Miocene balaenopterid whale (Cetacea: Mysticeti), Parabalaenoptera baulinensis; (new genus and species) from the Santa Cruz Mudstone, Point Reyes Peninsula, California. Proc. Calif. Acad. Sci. 50: 115–138.Google Scholar
  176. Zhou, K. (1982). Classification and phylogeny of the Superfamily Platanistoidea, with notes on evidence of the monophyly of the Cetacea. Sci. Rep. Whales Res. Inst. 34: 93–108.Google Scholar

Copyright information

© Plenum Publishing Corporation 2003

Authors and Affiliations

  • Jonathan H. Geisler
    • 1
  • Albert E. Sanders
    • 2
  1. 1.Department of Geology and Geography, Georgia Southern MuseumGeorgia Southern UniversityStatesboroUSA
  2. 2.Department of Natural SciencesThe Charleston MuseumCharlestonUSA

Personalised recommendations