Movatterモバイル変換

[0]ホーム
URL:

Skip to main content
Springer Nature Link
Log in

Astragali of Pakicetidae and other early-to-middle Eocene archaeocetes (Mammalia, Cetacea) of Pakistan: locomotion and habitat in the initial stages of whale evolution

  • Research Paper
  • Published:
PalZ Aims and scope Submit manuscript

Abstract

Richard Dehm and colleagues of the Bayerische Staatssammlung für Paläontologie und Geologie in Munich made an important collection of early-to-middle Eocene mammals at Ganda Kas in Pakistan during the winter of 1955/56. The genera and speciesIchthyolestes pinfoldi andGandakasia potens were named from this collection. Both are now recognized as early and primitive archaeocete cetaceans. In addition, Dehm’s group collected 16 complete or partial astragali of archaeocetes that were misidentified as artiodactyls. These bring the total number of archaeocete astragali known from Ganda Kas to 28. They separate clearly into four species distinguished by size: from smallest to largestIchthyolestes pinfoldi Dehm and Oettingen-Spielberg,Pakicetus attocki (West),Gandakasia potens Dehm and Oettingen-Spielberg, andAmbulocetus natans Thewissen et al. Ganda Kas artiodactyls are smaller and rare in comparison.Ichthyolestes andPakicetus are pakicetid archaeocetes,Gandakasia is presently indeterminate to family, andAmbulocetus is an ambulocetid. Tooth size and astragalus size are highly correlated, corroborating reference of astragali to the first three archaeocete taxa based on teeth. Multivariate morphometric comparison (Auto3Dgm) shows that pakicetid astragali overlap almost completely in shape with those of early artiodactyls. Middle Eocene protocetid astragali are divergent from both. Retention of an astragalus indistinguishable from that of artiodactyls shows that pakicetids are closely related to artiodactyls phylogenetically, but does not makeIchthyolestes andPakicetus terrestrial or cursorial. Other skeletal elements and bone microstructure indicate that pakicetids were semiaquatic like later protocetids. Tropical riverine and marginal marine facies of the Kuldana Formation are likely habitats for initial stages of the transition from terrestrial artiodactyls to semiaquatic and fully aquatic archaeocetes.

Kurzfassung

Richard Dehm machte mit Kollegen der Bayerischen Staatsammlung für Paläontologie und Geologie München im Winter 1955/56 bedeutende Aufsammlungen von Wirbeltierfossilien aus dem Unter- bis Mittel-Eozän von Ganda Kas in Pakistan. Die Gattungen und ArtenIchthyolestes pinfordi undGandakasia potens beruhen u. a. auf diesem Material. Beide werden jetzt als frühe und primitive Archaeoceten angesehen. Die Dehm-Gruppe sammelte darüber hinaus 16 vollständige Astragali von Archaeoceten, die seinerzeit den Paarhufern zugeordnet wurden. Somit beläuft sich jetzt die Zahl der Astragali von Archaeoceten von Ganda Kas auf 28. Diese lassen sich aufgrund ihrer Größe den nachfolgenden vier Arten (in aufsteigender Größe) zuordnen:Ichthyolestes pinfoldi Dehm und Oettingen-Spielberg,Pakicetus attocki (West),Gandakasia potens Dehm und Oettingen-Spielberg sowieAmbulocetus natans Thewissen et al. Die Artiodactyla aus Ganda Kas sind im Vergleich dazu kleiner und seltener. Die ArchaeocetenIchthyolestes undPakicetus sind Pakicetidae,Gandakasia kann zurzeit noch keiner Familie zugeordnet werden;Ambulocetus repräsentiert die Ambulocetidae. Da die Größe der Zähne und die der Astragali sehr gut korrelieren, bestätigt sich die Zuordnung der Astragali zu den ersten drei Taxa, die aufgrund der Zähne unterschieden wurden. Ein multivariater morphometrischer Vergleich (Auto3Dgm) zeigt, dass sich die Astragali der Pakicetidae in der Gestalt fast vollständig mit denen der frühen Artiodactyla decken. Die Astragali der mitteleozänen Protocetidae unterscheiden sich aber von beiden. Die Beibehaltung eines Astragalus, der von dem der Artiodactyla nicht unterscheidbar ist, zeigt dass die Pakicetiden mit den Artiodactylen in einem engen phylogenetischen Bezug stehen. Daraus folgt aber nicht, dassIchthyolestes undPakicetus terrestrisch lebten und vorwiegend Läufer waren. Andere Skelettelemente und die Mikrostruktur der Knochen zeigen, dass diese Pakicetiden ebenso semiaquatisch waren, wie die jüngeren Protocetidae. Die Fazies tropischer Flussmündungen und die randlichen marinen Bereiche, die in der Kuldana-Formation belegt sind, bilden den Lebensraum für einen beginnenden Übergang von terrestrischen Artiodactylen über semiaquatische Formen zu den voll aquatischen Archaeoceten.

This is a preview of subscription content,log in via an institution to check access.

Access this article

Log in via an institution

Subscribe and save

Springer+ Basic
¥17,985 /Month
  • Get 10 units per month
  • Download Article/Chapter or eBook
  • 1 Unit = 1 Article or 1 Chapter
  • Cancel anytime
Subscribe now

Buy Now

Price includes VAT (Japan)

Instant access to the full article PDF.

Fig. 1
Fig. 2
Fig. 3
Fig. 4
Fig. 5
Fig. 6
Fig. 7
Fig. 8
Fig. 9
Fig. 10
Fig. 11
Fig. 12
Fig. 13
Fig. 14
Fig. 15
Fig. 16

Similar content being viewed by others

Explore related subjects

Discover the latest articles and news from researchers in related subjects, suggested using machine learning.

References

  • Bajpai, S., V.V. Kapur, D.P. Das, B.N. Tiwari, N. Saravanan, and R. Sharma. 2005. Early Eocene land mammals from Vastan Lignite Mine, district Surat (Gujarat), western India.Journal of the Palaeontological Society of India 50: 101–113.

    Google Scholar 

  • Boisserie, J.-R., F. Lihoreau, and M. Brunet. 2005. The position of Hippopotamidae within Cetartiodactyla.Proceedings of the National Academy of Sciences USA 102: 1537–1541.

    Article  Google Scholar 

  • Boyer, D.M., J. Puente, J.T. Gladman, C. Glynn, S. Mukherjee, G.S. Yapuncich, and I. Daubechies. 2015. A new fully automated approach for aligning and comparing shapes.Anatomical Record 298: 249–276.

    Article  Google Scholar 

  • Cifelli, R.L. 1981. Patterns of evolution among the Artiodactyla and Perissodactyla (Mammalia).Evolution 35: 433–440.

    Google Scholar 

  • Cooper, L.N., E.R. Seiffert, M. Clementz, S.I. Madar, S. Bajpai, S.T. Hussain, and J.G.M. Thewissen. 2014. Anthracobunids from the middle Eocene of India and Pakistan are stem perissodactyls.PLoS One 9: e109232.

    Article  Google Scholar 

  • Cooper, L.N., J.G.M. Thewissen, S. Bajpai, and B.N. Tiwari. 2011. Postcranial morphology and locomotion of the Eocene raoellidIndohyus (Artiodactyla: Mammalia).Historical Biology 24: 279–310.

    Google Scholar 

  • Cooper, L.N., J.G.M. Thewissen, and S.T. Hussain. 2009. New middle Eocene archaeocetes (Cetacea: Mammalia) from the Kuldana Formation of northern Pakistan.Journal of Vertebrate Paleontology 29: 1289–1299.

    Article  Google Scholar 

  • Cotter, G.P. de. 1933. The geology of the part of the Attock District west of longitude 72°45′E.Memoirs of the Geological Survey of India 55 (2): 63–161.

    Google Scholar 

  • Dehm, R., T. zu Oettingen-Spielberg, and H. Vidal. 1958. Paläontologische und geologische Untersuchungen im Tertiär von Pakistan. 1. Die Münchner Forschungsreise nach Pakistan 1955–1956.Abhandlungen der Bayerischen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse, München, Neue Folge 90: 1–12.

    Google Scholar 

  • Dehm, R., and T. zu Oettingen-Spielberg. 1958. Paläontologische und geologische Untersuchungen im Tertiär von Pakistan. 2. Die mitteleocänen Säugetiere von Ganda Kas bei Basal in Nordwest-Pakistan.Abhandlungen der Bayerischen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse, München, Neue Folge 91: 1–54.

    Google Scholar 

  • Ducrocq, S., and F. Lihoreau. 2006. The occurrence of bothriodontines (Artiodactyla, Mammalia) in the Paleogene of Asia with special reference toElomeryx: paleobiogeographical implications.Journal of Asian Earth Sciences 27: 885–891.

    Article  Google Scholar 

  • Ehlert, A., J. Ferguson, and K. Gerlach. 2011. Magnetic resonance imaging and cross-sectional anatomy of the normal bovine tarsus.Anatomia, Histologia, Embryologia 40: 234–240.

    Article  Google Scholar 

  • Fish, F.E. 2016. Secondary evolution of aquatic propulsion in higher vertebrates: Validation and prospect.Integrative and Comparative Biology 56: 1285–1297.

    Article  Google Scholar 

  • Geais, G. 1934. LeBrachyodus borbonicus des Argiles de St-Henri (près Marseille).Travaux du Laboratoire de Géologie du Faculté des Sciences, Lyon 25: 1–55.

    Google Scholar 

  • Gingerich, P.D. 1977. A small collection of fossil vertebrates from the middle Eocene Kuldana and Kohat formations of Punjab (Pakistan).Contributions from the Museum of Paleontology, University of Michigan 24: 190–203.

    Google Scholar 

  • Gingerich, P.D. 1989. New earliest Wasatchian mammalian fauna from the Eocene of northwestern Wyoming: Composition and diversity in a rarely sampled high-floodplain assemblage.University of Michigan Papers on Paleontology 28: 1–97.

    Google Scholar 

  • Gingerich, P.D. 2003a. 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: 429–454.

    Article  Google Scholar 

  • Gingerich, P.D. 2003b. Stratigraphic and micropaleontologic constraints on the middle Eocene age of the mammal-bearing Kuldana Formation of Pakistan.Journal of Vertebrate Paleontology 23: 643–651.

    Article  Google Scholar 

  • Gingerich, P.D. 2005. Cetacea. InPlacental Mammals: Origin, Timing, and Relationships of the Major Extant Clades, ed. K.D. Rose, and J.D. Archibald, 234–252. Baltimore: Johns Hopkins University Press.

    Google Scholar 

  • Gingerich, P.D. 2012. Evolution of whales from land to sea.Proceedings of the American Philosophical Society 156: 309–323.

    Google Scholar 

  • Gingerich, P.D., M. Arif, I.H. Khan, M. ul Haq, J.I. Bloch, W.C. Clyde, and G.F. Gunnell. 2001a. Gandhera Quarry, a unique mammalian faunal assemblage from the early Eocene of Baluchistan (Pakistan). InEocene Biodiversity: Unusual Occurrences and Rarely Sampled Habitats, ed. G.F. Gunnell, 251–262. New York: Kluver Academic/Plenum Publishers.

    Chapter  Google Scholar 

  • Gingerich, P.D., M. ul Haq, W. von Koenigswald, W.J. Sanders, B.H. Smith, and I.S. Zalmout. 2009. New protocetid whale from the middle Eocene of Pakistan: birth on land, precocial development, and sexual dimorphism.PLoS One 4 (e4366): 1–20.

    Google Scholar 

  • Gingerich, P.D., M. Haq, I.S. Zalmout, I.H. Khan, and M.S. Malkani. 2001b. Origin of whales from early artiodactyls: hands and feet of Eocene Protocetidae from Pakistan.Science 293: 2239–2242.

    Article  Google Scholar 

  • Gingerich, P.D., S.M. Raza, M. Arif, M. Anwar, and X. Zhou. 1994. New whale from the Eocene of Pakistan and the origin of cetacean swimming.Nature 368: 844–847.

    Article  Google Scholar 

  • Gingerich, P.D., and D.E. Russell. 1981.Pakicetus inachus, a new archaeocete (Mammalia, Cetacea) from the early–middle Eocene Kuldana Formation of Kohat (Pakistan).Contributions from the Museum of Paleontology, University of Michigan 25: 235–246.

    Google Scholar 

  • Gingerich, P.D., and D.E. Russell. 1990. Dentition of early Eocene Pakicetus (Mammalia, Cetacea).Contributions from the Museum of Paleontology, University of Michigan 28: 1–20.

    Google Scholar 

  • Gingerich, P.D., D.E. Russell, D. Sigogneau-Russell, J.-L. Hartenberger, S.M.I. Shah, M. Hassan, K.D. Rose, and R.H. Ardrey. 1979. Reconnaissance survey and vertebrate paleontology of some Paleocene and Eocene formations in Pakistan.Contributions from the Museum of Paleontology, University of Michigan 25: 105–116.

    Google Scholar 

  • Gingerich, P.D., N.A. Wells, D.E. Russell, and S.M.I. Shah. 1983. Origin of whales in epicontinental remnant seas: New evidence from the early Eocene of Pakistan.Science 220: 403–406.

    Article  Google Scholar 

  • Haq, B. ul, J.A. Hardenbol, and P.R. Vail. 1987. Chronology of fluctuating sea levels since the Triassic.Science 235: 1156–1167.

    Article  Google Scholar 

  • Heissig, K. 1993. The astragalus in anoplotheres and oreodonts, phylogenetical and paleogeographical implications.Kaupia, Darmstädter Beiträge zur Naturgeschichte, Darmstadt 3: 173–178.

    Google Scholar 

  • Hellmund, M. 1991. Revision der europäischen Species der GattungElomeryx Marsh 1894 (Anthracotheriidae, Artiodactyla, Mammalia)—odontologische Untersuchungen.Palaeontographica, Abteilung A 220: 1–101.

    Google Scholar 

  • Hussain, S.T., H. de Bruijn, and J.J.M. Leinders. 1978. Middle Eocene rodents from the Kala Chitta Range (Punjab, Pakistan).Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen, Series B 81: 74–112.

    Google Scholar 

  • Hutchinson, G.E. 1959. Homage to Santa Rosalia or why are there so many kinds of animals?American Naturalist 93: 145–159.

    Article  Google Scholar 

  • Janis, C.M. 1989. A climatic explanation for patterns of evolutionary diversity in ungulate mammals.Palaeontology 32: 463–481.

    Google Scholar 

  • Jolly, A., and A. Sahni. 1990. Raoellid astragali from the Kalakot zone (Subathu Group): Evidence for assignment to the Artiodactyla.Current Science 59: 102–105.

    Google Scholar 

  • Khan, I.H., and W.C. Clyde. 2013. Lower Paleogene tectonostratigraphy of Balochistan: Evidence for time-transgressive late Paleocene–early Eocene uplift.Geosciences 3: 466–501.

    Article  Google Scholar 

  • Kripp, D. von. 1935. Bau, Funktion und Ableitung des Bewegungsmechanismus der Hinterextremitäten bei Paarhufern und Unpaarhufern unter besonderer Berücksichtigung der Fusswurzel.Morphologisches Jahrbuch 76: 259–278.

    Google Scholar 

  • Kumar, K., and A. Jolly. 1986. Earliest artiodactyl (Diacodexis, Dichobunidae: Mammalia) from the Eocene of Kalakot, north-western Himalaya, India.Indian Society of Geosciences Bulletin, Lucknow 2: 20–30.

    Google Scholar 

  • Kumar, K., K.D. Rose, R.S. Rana, L. Singh, T. Smith, and A. Sahni. 2010. Early Eocene artiodactyls (Mammalia) from western India.Journal of Vertebrate Paleontology 30: 1245–1274.

    Article  Google Scholar 

  • Kumar, K., and A. Sahni. 1985. Eocene mammals from the upper Subathu group, Kashmir Himalaya, India.Journal of Vertebrate Paleontology 5: 153–168.

    Article  Google Scholar 

  • Latif, M.A. 1970. Explanatory notes on the geology of southeastern Hazara to accompany the revised geological map.Jahrbuch der Geologischen Bundesanstalt Wien, Sonderband 15: 5–20.

    Google Scholar 

  • Lydekker, R. 1880. A sketch of the history of the fossil vertebrata of India.Journal of the Asiatic Society of Bengal. Part II. Physical Science 69: 8–40.

    Google Scholar 

  • Madar, S.I. 2007. The postcranial skeleton of early Eocene pakicetid cetaceans.Journal of Paleontology 81: 176–200.

    Article  Google Scholar 

  • Madar, S.I., J.G.M. Thewissen, and S.T. Hussain. 2002. Additional holotype remains ofAmbulocetus natans (Cetacea, Ambulocetidae), and their implications for locomotion in early whales.Journal of Vertebrate Paleontology 22: 405–422.

    Article  Google Scholar 

  • Martinez, J.-N., and J. Sudre. 1995. The astragalus of Paleogene artiodactyls: Comparative morphology, variability and prediction of body mass.Lethaia 28: 197–209.

    Article  Google Scholar 

  • Middlemiss, C.S. 1896. The geology of Hazara and Black Mountains.Geological Survey of India, Memoirs 26: 1–302.

    Google Scholar 

  • Missiaen, P., and P.D. Gingerich. 2014. New basal Perissodactyla (Mammalia) from the lower Eocene Ghazij Formation of Pakistan.Contributions from the Museum of Paleontology, University of Michigan 32: 139–160.

    Google Scholar 

  • Montgelard, C., F.M. Catzeflis, and E. Douzery. 1997. Phylogenetic relationships of artiodactyls and cetaceans as deduced from the comparison of cytochromeb and 12S rRNA mitochondrial sequences.Molecular Biology and Evolution 14: 550–559.

    Article  Google Scholar 

  • Nummela, S., S.T. Hussain, and J.G.M. Thewissen. 2006. Cranial anatomy of Pakicetidae (Cetacea, Mammalia).Journal of Vertebrate Paleontology 26: 746–759.

    Article  Google Scholar 

  • Pilgrim, G.E. 1940. Middle Eocene mammals from northwest India.Proceedings of the Zoological Society of London, Series B 110: 127–152.

    Google Scholar 

  • Pinfold, E.S. 1918. Notes on the structure and stratigraphy of the north-west Punjab.Records of the Geological Survey of India 49: 137–160.

    Google Scholar 

  • Qureshi, M.K.A., and M. Ahmad. 2001.Geological Map of the Kala Chitta Range, Northern Punjab, Pakistan. Quetta: Geological Survey of Pakistan.

    Google Scholar 

  • Ranga Rao, A. 1971. New mammals from Murree (Kalakot Zone) of the Himalyan foot hills near Kalakot, Jammu and Kashmir State, India.Journal of the Geological Society of India 12: 125–134.

    Google Scholar 

  • Ranga Rao, A. 1972. Further studies on the vertebrate fauna of Kalakot, India: new mammalian genera and species from the Kalakot zone of Himalayan foot hills near Kalakot, Jammu and Kashmir state, India.Directorate of Geology, Oil and Natural Gas Commission, Dehra Dun, Special Papers 1: 1–21.

    Google Scholar 

  • Rose, K.D. 1982. Skeleton ofDiacodexis, oldest known artiodactyl.Science 216: 621–623.

    Article  Google Scholar 

  • Rose, K.D. 1985. Comparative osteology of North American dichobunid artiodactyls.Journal of Paleontology 59: 1203–1226.

    Google Scholar 

  • Rose, K.D., A.E. Chew, R.H. Dunn, M.J. Kraus, H.C. Fricke, and S.P. Zack. 2012. Earliest Eocene mammalian fauna from the Paleocene–Eocene Thermal Maximum at Sand Creek Divide, southern Bighorn Basin, Wyoming.University of Michigan Papers on Paleontology 36: 1–122.

    Google Scholar 

  • Rose, K.D., L.T. Holbrook, R.S. Rana, K. Kumar, K.E. Jones, H.E. Ahrens, P. Missiaen, A. Sahni, and T. Smith. 2014. Early Eocene fossils suggest that the mammalian order Perissodactyla originated in India.Nature Communications 5 (5570): 1–9.

    Google Scholar 

  • Sahni, A., and S.K. Khare. 1972. Three new Eocene mammals from Rajauri District, Jammu and Kashmir.Journal of the Palaeontological Society of India 16: 41–53.

    Google Scholar 

  • Sahni, A., and S.K. Khare. 1973. Additional Eocene mammals from the Subathu Formation of Jammu and Kashmir.Journal of the Palaeontological Society of India 17: 31–49.

    Google Scholar 

  • Schaeffer, B. 1947. Notes on the origin and function of the artiodactyl tarsus.American Museum Novitates 1356: 1–24.

    Google Scholar 

  • Scott, W.B. 1894. The structure and relationships ofAncodus.Academy of Natural Sciences, Philadelphia, Journal 9: 461–497.

    Google Scholar 

  • Scott, W.B. 1940. The mammalian fauna of the White River Oligocene. Part IV. Artiodactyla.Transactions of the American Philosophical Society 28: 363–746.

    Article  Google Scholar 

  • Simpson, G.G. 1953.The Major Features of Evolution. New York: Columbia University Press.

    Google Scholar 

  • Sinclair, W.J. 1914. A revision of the bunodont Artiodactyla of the middle and lower Eocene of North America.Bulletin of the American Museum of Natural History 33: 267–295.

    Google Scholar 

  • Thewissen, J.G.M., L.N. Cooper, M.T. Clementz, S. Bajpai, and B.N. Tiwari. 2007. Whales originated from aquatic artiodactyls in the Eocene epoch of India.Nature 450: 1190–1195.

    Article  Google Scholar 

  • Thewissen, J.G.M., P.D. Gingerich, and D.E. Russell. 1987. Artiodactyla and Perissodactyla (Mammalia) from the early–middle Eocene Kuldana Formation of Kohat (Pakistan).Contributions from the Museum of Paleontology, University of Michigan 27: 247–274.

    Google Scholar 

  • Thewissen, J.G.M., and S.T. Hussain. 1990. Postcranial osteology of the most primitive artiodactylDiacodexis pakistanensis (Dichobunidae).Anatomia Histologia Embryologia, Zentralblatt für Veterinärmedizin, Reihe C 19: 37–48.

    Article  Google Scholar 

  • Thewissen, J.G.M., and S.T. Hussain. 1993. Origin of underwater hearing in whales.Nature 361: 444–445.

    Article  Google Scholar 

  • Thewissen, J.G.M., and S.T. Hussain. 1998. Systematic review of the Pakicetidae, early and middle Eocene Cetacea from Pakistan and India.Bulletin of Carnegie Museum of Natural History 34: 220–238.

    Google Scholar 

  • Thewissen, J.G.M., and S.T. Hussain. 2000.Attockicetus praecursor, a new remingtonocetid cetacean from marine Eocene sediments of Pakistan.Journal of Mammalian Evolution 7: 133–146.

    Article  Google Scholar 

  • Thewissen, J.G.M., S.T. Hussain, and M. Arif. 1994. Fossil evidence for the origin of aquatic locomotion in archaeocete whales.Science 263: 210–212.

    Article  Google Scholar 

  • Thewissen, J.G.M., S.I. Madar, and S.T. Hussain. 1996.Ambulocetus natans, an Eocene cetacean (Mammalia) from Pakistan.Courier Forschungsinstitut Senckenberg, Frankfurt am Main 191: 1–86.

    Google Scholar 

  • Thewissen, J.G.M., S.I. Madar, and S.T. Hussain. 1998. Whale ankles and evolutionary relationships.Nature 395: 452.

    Article  Google Scholar 

  • Thewissen, J.G.M., D.E. Russell, P.D. Gingerich, and S.T. Hussain. 1983. A new dichobunid artiodactyl (Mammalia) from the Eocene of northwest Pakistan: dentition and classification.Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen, Series B 86: 153–180.

    Google Scholar 

  • Thewissen, J.G.M., E.M. Williams, and S.T. Hussain. 2001a. Eocene mammal faunas from northern Indo-Pakistan.Journal of Vertebrate Paleontology 21: 347–366.

    Article  Google Scholar 

  • Thewissen, J.G.M., E.M. Williams, L.J. Roe, and S.T. Hussain. 2001b. Skeletons of terrestrial cetaceans and the relationship of whales to artiodactyls.Nature 413: 277–281.

    Article  Google Scholar 

  • Van Valen, L.M. 1965. Paroxyclaenidae, an extinct family of Eurasian mammals.Journal of Mammalogy 46: 388–397.

    Article  Google Scholar 

  • Van Valen, L.M. 1966. Deltatheridia, a new order of mammals.Bulletin of the American Museum of Natural History 132: 1–126.

    Google Scholar 

  • Van Valen, L.M. 1968. Monophyly or diphyly in the origin of whales.Evolution 22: 37–41.

    Article  Google Scholar 

  • Wells, N.A. 1983. Transient streams in sand-poor redbeds: Early–middle Eocene Kuldana Formation in northern Pakistan. InModern and Ancient Fluvial Systems, eds. J.D. Collinson, and J. Lewin, 393–403. Oxford: Blackwell.

    Chapter  Google Scholar 

  • Wells, N.A. 1984.Marine and Continental Sedimentation in the Early Cenozoic Kohat Basin and Adjacent Northwestern Indo-Pakistan. Ph.D. Dissertation, Ann Arbor: University of Michigan.

  • Wells, N.A., and P.D. Gingerich. 1983. Review of Eocene Anthracobunidae (Mammalia, Proboscidea) with a new genus and species,Jozaria palustris, from the Kuldana Formation of Kohat (Pakistan).Contributions from the Museum of Paleontology, University of Michigan 26: 117–139.

    Google Scholar 

  • West, R.M. 1980. Middle Eocene large mammal assemblage with Tethyan affinities, Ganda Kas region, Pakistan.Journal of Paleontology 54: 508–533.

    Google Scholar 

  • West, R.M., and J.R. Lukacs. 1979. Geology and vertebrate-fossil localities, Tertiary continental rocks, Kala Chitta Hills, Attock District, Pakistan.Contributions in Biology and Geology, Milwaukee Public Museum 26: 1–20.

    Google Scholar 

  • Wynne, A.B. 1874. Notes on the geology of the neighbourhood of Mari Hills station in the Panjab.Records of the Geological Survey of India 7: 64–74.

    Google Scholar 

  • Wynne, A.B. 1877. Note on the Tertiary zone and underlying rocks in the north-west Panjab.Records of the Geological Survey of India 10: 107–132.

    Google Scholar 

Download references

Acknowledgements

We thank Richard Dehm (1907–1996), Therese zu Oettingen-Spielberg (1909–1991), and Helmut Vidal (1919–2002) of the Bayerische Staatssammlung für Paläontologie und Geologie München for the foresight to investigate the Pakistan Eocene and for their effort in collecting the astragali described here. Gertrud Rößner generously allowed us to borrow these for comparative study. Jeremy Hooker granted access to relevant Natural History Museum London specimens, and J. G. M. Thewissen provided casts of Howard University-Geological Survey of Pakistan specimens. We thank Michelle Lynch of the MicroCT Core at the University of Michigan School of Dentistry for computed tomography. R. Ewan Fordyce and an anonymous referee provided manuscript reviews that improved this study substantially. Support for initial field work in Pakistan was provided by the Geological Survey of Pakistan and by the Smithsonian Foreign Currency Program. Follow-up investigation, again in collaboration with the Geological Survey of Pakistan, was funded by grants from the National Geographic Society (CRE 5072-93) and the U. S. National Science Foundation (EAR-9714923). Study of the Dehm collection of astragali was initiated during tenure of an Alexander von Humboldt Foundation research award at the Steinmann-Institut (Paläontologie), Rheinische Friedrich-Wilhelms-Universität Bonn (2006).

Author information

Authors and Affiliations

  1. Museum of Paleontology, University of Michigan, 1109 Geddes Avenue, Ann Arbor, MI, 48109-1079, USA

    Philip D. Gingerich

  2. Bayerische Staatssammlung für Paläontologie und Geologie, Richard-Wagner-Straße 10, 80333, Munich, Germany

    Kurt Heissig

  3. Department of Biology, Calvin College, 1726 Knollcrest Circle SE, Grand Rapids, MI, 49546-4403, USA

    Ryan M. Bebej

  4. Steinmann-Institut für Geologie, Mineralogie und Paläontologie, Rheinische Friedrich-Wilhelms-Universität Bonn, Nussallee 8, 53115, Bonn, Germany

    Wighart von Koenigswald

Authors
  1. Philip D. Gingerich
  2. Kurt Heissig
  3. Ryan M. Bebej
  4. Wighart von Koenigswald

Corresponding author

Correspondence toPhilip D. Gingerich.

Additional information

Handling editor: Thomas Mörs.

Rights and permissions

About this article

Check for updates. Verify currency and authenticity via CrossMark

Cite this article

Gingerich, P.D., Heissig, K., Bebej, R.M.et al. Astragali of Pakicetidae and other early-to-middle Eocene archaeocetes (Mammalia, Cetacea) of Pakistan: locomotion and habitat in the initial stages of whale evolution.PalZ91, 601–627 (2017). https://doi.org/10.1007/s12542-017-0362-8

Download citation

Keywords

Schlüsselwörter

Access this article

Subscribe and save

Springer+ Basic
¥17,985 /Month
  • Get 10 units per month
  • Download Article/Chapter or eBook
  • 1 Unit = 1 Article or 1 Chapter
  • Cancel anytime
Subscribe now

Buy Now

Price includes VAT (Japan)

Instant access to the full article PDF.

Advertisement

[8]ページ先頭
©2009-2025 Movatter.jp