Skip to main content Accessibility help
×
Home
Hostname: page-component-564cf476b6-z65vl Total loading time: 0.281 Render date: 2021-06-20T04:56:59.975Z Has data issue: true Feature Flags: { "shouldUseShareProductTool": true, "shouldUseHypothesis": true, "isUnsiloEnabled": true, "metricsAbstractViews": false, "figures": true, "newCiteModal": false, "newCitedByModal": true, "newEcommerce": true }

First remingtonocetid archaeocete (Mammalia, Cetacea) from the middle Eocene of Egypt with implications for biogeography and locomotion in early cetacean evolution

Published online by Cambridge University Press:  21 March 2016

Ryan M. Bebej
Affiliation:
Department of Biology, Calvin College, 1726 Knollcrest Circle SE, Grand Rapids, Michigan 49546-4403, USA 〈bebej@calvin.edu〉
Iyad S. Zalmout
Affiliation:
Mammals Research Chair, Department of Zoology, College of Science, King Saud University, PO Box 2455, Riyadh 11451, Saudi Arabia 〈izalmout@ksu.edu.sa〉
Ahmed A. Abed El-Aziz
Affiliation:
Egyptian Environmental Affairs Agency, Wadi Al-Hitan World Heritage Site, Fayum, Egypt 〈ahmedreactors@gmail.com〉, 〈wrpashark@yahoo.com〉
Mohammed Sameh M. Antar
Affiliation:
Egyptian Environmental Affairs Agency, Wadi Al-Hitan World Heritage Site, Fayum, Egypt 〈ahmedreactors@gmail.com〉, 〈wrpashark@yahoo.com〉
Philip D. Gingerich
Affiliation:
Department of Earth and Environmental Sciences, Museum of Paleontology, University of Michigan, 1109 Geddes Ave, Ann Arbor, Michigan 48109-1079, USA 〈gingeric@umich.edu〉

Abstract

Remingtonocetidae are Eocene archaeocetes that represent a unique experiment in cetacean evolution. They possess long narrow skulls, long necks, fused sacra, and robust hind limbs. Previously described remingtonocetids are known from middle Eocene Lutetian strata in Pakistan and India. Here we describe a new remingtonocetid, Rayanistes afer, n. gen. n. sp., recovered from a middle to late Lutetian interval of the Midawara Formation in Egypt. The holotype preserves a sacrum with four vertebral centra; several lumbar and caudal vertebrae; an innominate with a complete ilium, ischium, and acetabulum; and a nearly complete femur. The ilium and ischium of Rayanistes are bladelike, rising sharply from the body of the innominate anterior and posterior to the acetabulum, and the acetabular notch is narrow. These features are diagnostic of Remingtonocetidae, but their development also shows that Rayanistes had a specialized mode of locomotion. The expanded ischium is larger than that of any other archaeocete, supporting musculature for powerful retraction of the hind limbs during swimming. Posteriorly angled neural spines on lumbar vertebrae and other features indicate increased passive flexibility of the lumbus. Rayanistes probably used its enhanced lumbar flexibility to increase the length of the power stroke during pelvic paddling. Recovery of a remingtonocetid in Egypt broadens the distribution of Remingtonocetidae and shows that protocetids were not the only semiaquatic archaeocetes capable of dispersal across the southern Tethys Sea.

Type
Articles
Copyright
Copyright © 2016, The Paleontological Society 

Access options

Get access to the full version of this content by using one of the access options below.

References

Alexander, R.M., Dimery, N.J., and Ker, R.F., 1985, Elastic structures in the back and their role in galloping in some mammals: Journal of Zoology, v. 207, p. 467482.CrossRefGoogle Scholar
Bajpai, S., and Gingerich, P.D., 1998, A new Eocene archaeocete (Mammalia, Cetacea) from India and the time of origin of whales: Proceedings of the National Academy of Sciences USA, v. 95, p. 1546415468.CrossRefGoogle ScholarPubMed
Bajpai, S., and Thewissen, J.G.M., 1998, Middle Eocene cetaceans from the Harudi and Subathu formations of India, in Thewissen, J.G.M., ed., The Emergence of Whales: Evolutionary Patterns in the Origin of Cetacea, New York, Plenum Press, p. 213233.CrossRefGoogle Scholar
Bajpai, S., and Thewissen, J.G.M., 2000, A new diminutive Eocene whale from Kachchh (Gujarat, India) and its implications for locomotor evolution of cetaceans: Current Science, v. 79, p. 14781482.Google Scholar
Bajpai, S., Thewissen, J.G.M., and Conley, R.W., 2011, Cranial anatomy of middle Eocene Remingtonocetus (Cetacea, Mammalia) from Kutch, India: Journal of Paleontology, v. 85, p. 703718.CrossRefGoogle Scholar
Beadnell, H.J.L., 1905, The topography and geology of the Fayum Province of Egypt, Cairo, Survey Department of Egypt, 101 p.Google Scholar
Bebej, R.M., Haq, M., Zalmout, I.S., and Gingerich, P.D., 2012, Morphology and function of the vertebral column in Remingtonocetus domandaensis (Mammalia, Cetacea) from the middle Eocene Domanda Formation of Pakistan: Journal of Mammalian Evolution, v. 19, p. 77104.CrossRefGoogle Scholar
Brisson, M.-J., 1762, Le Regnum Animale in Classes IX Distributum, sive Synopsis Methodica: Leiden, Lugduni Batavorum, Theodorum Haak, 296 p.Google Scholar
Carlson, H., 1978, Morphology and contraction properties of cat lumbar back muscles: Acta Physiologica Scandinavica, v. 103, p. 180197.CrossRefGoogle ScholarPubMed
Deban, S.M., Schilling, N., and Carrier, D.R., 2012, Activity of extrinsic limb muscles in dogs at walk, trot and gallop: Journal of Experimental Biology, v. 215, p. 287300.CrossRefGoogle ScholarPubMed
Dumas, G.A., Beaudoin, L., and Drouin, G., 1987, In situ mechanical behavior of posterior spinal ligaments in the lumbar region, an in vitro study: Journal of Biomechanics, v. 20, p. 301310.CrossRefGoogle ScholarPubMed
English, A.W., 1980, The functions of the lumbar spine during stepping in the cat: Journal of Morphology, v. 165, p. 5566.CrossRefGoogle Scholar
Evans, H.E., 1993, Miller’s Anatomy of the Dog, Philadelphia, Saunders, 1113 p.Google Scholar
Fisher, R.E., Scott, K.M., and Adrain, B., 2010, Hind limb myology of the common hippopotamus, Hippopotamus amphibius (Artiodactyla: Hippopotamidae): Zoological Journal of the Linnean Society, v. 158, p. 661682.CrossRefGoogle Scholar
Fleagle, J.G., and Meldrum, D.J., 1988, Locomotor behavior and skeletal morphology of two sympatric pitheciine monkeys, Pithecia pithecia and Chiropotes satanas: American Journal of Primatology, v. 16, p. 227249.CrossRefGoogle Scholar
Fraas, E., 1904, Neue Zeuglodonten aus dem unteren Mitteleocän vom Mokattam bei Cairo: Geologische und Paläontologische Abhandlungen, Neue Folge, Jena, v. 6, p. 197220.Google Scholar
Gál, J.M., 1993, Mammalian spinal biomechanics: II. Intervertebral lesion experiments and mechanisms of bending resistance: Journal of Experimental Biology, v. 174, p. 281297.Google ScholarPubMed
Geisler, J.H., Sanders, A.E., and Luo, Z., 2005, A new protocetid whale (Cetacea: Archaeoceti) from the late middle Eocene of South Carolina: American Museum Novitates, v. 3480, p. 165.CrossRefGoogle Scholar
Getty, R., 1975, Ruminant myology, in Getty, R., ed., Sisson and Grossman’s the Anatomy of Domestic Animals, 5th ed.., Philadelphia, W.B. Saunders Co, p. 791860.Google Scholar
Gingerich, P.D., 1992, Marine mammals (Cetacea and Sirenia) from the Eocene of Gebel Mokattam and Fayum, Egypt: stratigraphy, age, and paleoenvironments: University of Michigan Papers on Paleontology, v. 30, p. 184.Google Scholar
Gingerich, P.D., 1998, Paleobiological perspectives on Mesonychia, Archaeoceti, and the origin of whales, in Thewissen, J.G.M., ed., Emergence of Whales: Evolutionary Patterns in the Origin of Cetacea, New York, Plenum Press, p. 423449.CrossRefGoogle Scholar
Gingerich, P.D., 2008, Early evolution of whales: a century of research in Egypt, in Fleagle J.G., and Gilbert, C.C., eds., Elwyn Simons: A Search for Origins, New York, Springer, p. 107124.CrossRefGoogle Scholar
Gingerich, P.D., 2010, Cetacea, in Werdelin, L., and Sanders, W.J., eds., Cenozoic Mammals of Africa, Berkeley, University of California Press, p. 873899.CrossRefGoogle Scholar
Gingerich, P.D., Antar, M.S., and Zalmout, I.S., 2013, Faunas of whales and sea cows (Cetacea and Sirenia) from middle and upper Eocene strata in western Fayum Province, Egypt (abstract): Journal of Vertebrate Paleontology, Program and Abstracts, v. 2013, p. 134135.Google Scholar
Gingerich, P.D., Arif, M., Bhatti, M.A., and Clyde, W.C., 1998, Middle Eocene stratigraphy and marine mammals (Cetacea and Sirenia) of the Sulaiman Range, Pakistan, Bulletin of Carnegie Museum of Natural History, v. 34, p. 239259.Google Scholar
Gingerich, P.D., Arif, M., and Clyde, W.C., 1995a, New archaeocetes (Mammalia, Cetacea) from the middle Eocene Domanda Formation of the Sulaiman Range, Punjab (Pakistan): Contributions from the Museum of Paleontology, University of Michigan, v. 29, p. 291330.Google Scholar
Gingerich, P.D., Arif, M., Bhatti, M.A., Raza, H.A., and Raza, S.M., 1995b, Protosiren and Babiacetus (Mammalia, Sirenia and Cetacea) from the middle Eocene Drazinda Formation, Sulaiman Range, Punjab (Pakistan): Contributions from the Museum of Paleontology, University of Michigan, v. 29, p. 331357.Google Scholar
Gingerich, P.D., Haq, M., Khan, I.H., and Zalmout, I.S., 2001a, Eocene stratigraphy and archaeocete whales (Mammalia, Cetacea) of Drug Lahar in the eastern Sulaiman Range, Balochistan (Pakistan): Contributions from the Museum of Paleontology, University of Michigan, v. 30, p. 269319.Google Scholar
Gingerich, P.D., Haq, M., Koenigswald, W.v., Sanders, W.J., Smith, B.H., and Zalmout, I.S., 2009, New protocetid whale from the middle Eocene of Pakistan: birth on land, precocial development, and sexual dimorphism: PLoS ONE, v. 4, p. e4366.CrossRefGoogle Scholar
Gingerich, P.D., Haq, M., Zalmout, I.S., Khan, I.H., and Malkani, M.S., 2001b, Origin of whales from early artiodactyls: hands and feet of Eocene Protocetidae from Pakistan: Science, v. 293, p. 22392242.CrossRefGoogle ScholarPubMed
Gingerich, P.D., Raza, S.M., Arif, M., Anwar, M., and Zhou, X., 1993, Partial skeletons of Indocetus ramani (Mammalia, Cetacea) from the lower middle Eocene Domanda Shale in the Sulaiman Range of Pakistan: Contributions from the Museum of Paleontology, University of Michigan, v. 28, p. 393416.Google Scholar
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, v. 368, p. 844847.CrossRefGoogle Scholar
Gingerich, P.D., Wells, N.A., Russell, D.E., and Shah, S.M.I., 1983, Origin of whales in epicontinental remnant seas: new evidence from the early Eocene of Pakistan: Science, v. 220, p. 403406.CrossRefGoogle ScholarPubMed
Hulbert, R.C., Petkewich, R.M., Bishop, G.A., Bukry, D., and Aleshire, D.P., 1998, A new middle Eocene protocetid whale (Mammalia: Cetacea: Archaeoceti) and associated biota from Georgia: Journal of Paleontology, v. 72, p. 907927.CrossRefGoogle Scholar
Kappelman, J., 1988, Morphology and locomotor adaptations of the bovid femur in relation to habitat: Journal of Morphology, v. 198, p. 119130.CrossRefGoogle ScholarPubMed
Kumar, K., and Sahni, A., 1986, Remingtonocetus harudiensis, new combination, a middle Eocene archaeocete (Mammalia, Cetacea) from western Kutch, India: Journal of Vertebrate Paleontology, v. 6, p. 326349.Google Scholar
Madar, S.I., 1998, Structural adaptations of early archaeocete long bones, in Thewissen, J.G.M., ed., The Emergence of Whales: Evolutionary Patterns in the Origin of Cetacea, New York, Plenum Press, p. 353378.CrossRefGoogle Scholar
Madar, S.I., 2007, The postcranial skeleton of early Eocene pakicetid cetaceans: Journal of Paleontology, v. 81, p. 176200.CrossRefGoogle Scholar
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: Journal of Vertebrate Paleontology, v. 22, p. 405422.CrossRefGoogle Scholar
McLeod, S.A., and Barnes, L.G., 2008, A new genus and species of Eocene protocetid archaeocete whale (Mammalia, Cetacea) from the Atlantic Coastal Plain, Science Series: Natural History Museum of Los Angeles County, v. 41, p. 7398.Google Scholar
Pabst, D.A., 1993, Intramuscular morphology and tendon geometry of the epaxial swimming muscles of dolphins: Journal of Zoology, London, v. 230, p. 159176.CrossRefGoogle Scholar
Pabst, D.A., 2000, To bend a dolphin: convergence of force transmission designs in cetaceans and scombrid fishes: American Zoologist, v. 40, p. 146155.Google Scholar
Passalacqua, N.V., 2009, Forensic age-at-death estimation from the human sacrum: Journal of Forensic Sciences, v. 54, p. 255262.CrossRefGoogle ScholarPubMed
Pickford, M., 2008, Differences in ligamenta flava among some mammals: Iowa Orthopaedic Journal, v. 15, p. 141146.Google Scholar
Ponseti, I.V., 1995, The myth of the hippo-like anthracothere: the eternal problem of homology and convergence: Revista Española de Paleontología, v. 23, p. 3190.Google Scholar
Robertson, R.A., and Shadle, A.R., 1954, Osteologic criteria of age in beavers: Journal of Mammalogy, v. 35, p. 197203.CrossRefGoogle Scholar
Sahni, A., and Mishra, V.P., 1972, A new species of Protocetus (Cetacea) from the middle Eocene of Kutch, western India: Palaeontology, v. 15, p. 490495.Google Scholar
Sahni, A., and Mishra, V.P., 1975, Lower Tertiary vertebrates from western India: Palaeontological Society of India, Monographs, v. 3, p. 148.Google Scholar
Sánchez-Villagra, M.R., 2002, Comparative patterns of postcranial ontogeny in therian mammals: an analysis of relative timing of ossification events: Journal of Experimental Zoology (Molecular and Developmental Evolution), v. 294, p. 264273.Google ScholarPubMed
Schilling, N., and Carrier, D.R., 2010, Function of the epaxial muscles in walking, trotting and galloping dogs: implications for the evolution of epaxial muscle function in tetrapods: Journal of Experimental Biology, v. 213, p. 14901502.CrossRefGoogle ScholarPubMed
Schilling, N., Fischbein, T., Yang, E.P., and Carrier, D.R., 2009, Function of the extrinsic hindlimb muscles in trotting dogs: Journal of Experimental Biology, v. 212, p. 10361052.CrossRefGoogle ScholarPubMed
Slijper, E.J., 1946, Comparative biologic-anatomical investigations on the vertebral column and spinal musculature of mammals: Verhandlingen der Koninklijke Nederlandse Akademie van Wetenschappen, Afdeling Natuurkunde, Tweede Sectie, v. 42, p. 1128.Google Scholar
Strougo, A., 2008, The Mokattamian stage: 125 years later: Middle East Research Center, Ain Shams University, Cairo, Earth Science Series, v. 22, p. 47108.Google Scholar
Strougo, A., Abul-Nasr, R.A., and Haggag, M.A., 1982, Contribution to the age of the middle Mokattam beds of Egypt: Neues Jahrbuch für Geologie und Paläontologie, Monatshefte, Stuttgart, v. 1982, p. 240243.Google Scholar
Thewissen, J.G.M., and Bajpai, S., 2009, New skeletal material of Andrewsiphius and Kutchicetus, two Eocene cetaceans from India: Journal of Paleontology, v. 83, p. 635663.CrossRefGoogle Scholar
Thewissen, J.G.M., and Hussain, S.T., 2000, Attockicetus praecursor, a new remingtonocetid cetacean from marine Eocene sediments of Pakistan: Journal of Mammalian Evolution, v. 7, p. 133146.CrossRefGoogle Scholar
Trivedy, A.N., and Satsangi, P.P., 1984, A new archaeocete (whale) from the Eocene of India: Abstracts of 27th International Geological Congress, Moscow, v. 1, p. 322323.Google Scholar
Uhen, M.D., 2010, The origin(s) of whales: Annual Review of Earth and Planetary Sciences, v. 38, p. 189219.CrossRefGoogle Scholar
Uhen, M.D., 2013, A review of North American Basilosauridae: Alabama Museum of Natural History Bulletin, v. 31, p. 145.Google Scholar
Uhen, M.D., Pyenson, N.D., Devries, T.J., Urbina, M., and Renne, P.R., 2011, New middle Eocene whales from the Pisco Basin of Peru: Journal of Paleontology, v. 85, p. 955969.CrossRefGoogle Scholar
Underwood, C.J., Ward, D.J., King, C., Antar, M.S., Zalmout, I.S., and Gingerich, P.D., 2011, Shark and ray faunas in the late Eocene of the Fayum area, Egypt: Proceedings of the Geologists’ Association, London, v. 122, p. 4766.Google Scholar
West, R.M., 1980, Middle Eocene large mammal assemblage with Tethyan affinities, Ganda Kas region, Pakistan: Journal of Paleontology, v. 54, p. 508533.Google Scholar
White, J.L., 1993, Indicators of locomotor habits in xenarthrans: evidence for locomotor heterogeneity among fossil sloths: Journal of Vertebrate Paleontology, v. 13, p. 230242.CrossRefGoogle Scholar
Williams, E.M., 1998, Synopsis of the earliest cetaceans: Pakicetidae, Ambulocetidae, Remingtonocetidae, and Protocetidae, in Thewissen, J.G.M., ed., The Emergence of Whales: Evolutionary Patterns in the Origin of Cetacea, New York, Plenum Press, p. 128.Google Scholar
Zalmout, I.S., and Gingerich, P.D., 2012, Late Eocene sea cows (Mammalia, Sirenia) from Wadi Al Hitan in the Western Desert of Fayum, Egypt: University of Michigan Papers on Paleontology, v. 37, p. 1158.Google Scholar
Zhou, X., Sanders, W.J., and Gingerich, P.D., 1992, Functional and behavioral implications of vertebral structure in Pachyaena ossifraga (Mammalia, Mesonychia): Contributions from the Museum of Paleontology, University of Michigan, v. 28, p. 289319.Google Scholar
12
Cited by

Send article to Kindle

To send this article to your Kindle, first ensure no-reply@cambridge.org is added to your Approved Personal Document E-mail List under your Personal Document Settings on the Manage Your Content and Devices page of your Amazon account. Then enter the ‘name’ part of your Kindle email address below. Find out more about sending to your Kindle. Find out more about sending to your Kindle.

Note you can select to send to either the @free.kindle.com or @kindle.com variations. ‘@free.kindle.com’ emails are free but can only be sent to your device when it is connected to wi-fi. ‘@kindle.com’ emails can be delivered even when you are not connected to wi-fi, but note that service fees apply.

Find out more about the Kindle Personal Document Service.

First remingtonocetid archaeocete (Mammalia, Cetacea) from the middle Eocene of Egypt with implications for biogeography and locomotion in early cetacean evolution
Available formats
×

Send article to Dropbox

To send this article to your Dropbox account, please select one or more formats and confirm that you agree to abide by our usage policies. If this is the first time you use this feature, you will be asked to authorise Cambridge Core to connect with your <service> account. Find out more about sending content to Dropbox.

First remingtonocetid archaeocete (Mammalia, Cetacea) from the middle Eocene of Egypt with implications for biogeography and locomotion in early cetacean evolution
Available formats
×

Send article to Google Drive

To send this article to your Google Drive account, please select one or more formats and confirm that you agree to abide by our usage policies. If this is the first time you use this feature, you will be asked to authorise Cambridge Core to connect with your <service> account. Find out more about sending content to Google Drive.

First remingtonocetid archaeocete (Mammalia, Cetacea) from the middle Eocene of Egypt with implications for biogeography and locomotion in early cetacean evolution
Available formats
×
×

Reply to: Submit a response

Please enter your response.

Your details

Please enter a valid email address.

Conflicting interests

Do you have any conflicting interests? *