Hostname: page-component-78c5997874-mlc7c Total loading time: 0 Render date: 2024-11-13T01:06:55.811Z Has data issue: false hasContentIssue false

Osteological redescription of the skull of Microcleidus homalospondylus (Sauropterygia, Plesiosauria) from the Lower Jurassic of England

Published online by Cambridge University Press:  14 July 2015

David Brown
Affiliation:
School of Dental Sciences, Newcastle University, Framlington Place, Newcastle upon Tyne, NE2 4BW, UK,
Peggy Vincent
Affiliation:
CR2P-UMR 7207 CNRS, Muséum National d'Histoire Naturelle, Univ. Paris 06, 57 rue Cuvier, CP38, F-75005, Paris, France, ; Staatliches Museum für Naturkunde, Rosenstein 1, D-70191 Stuttgart, Germany
Nathalie Bardet
Affiliation:
CR2P-UMR 7207 CNRS, Muséum National d'Histoire Naturelle, Univ. Paris 06, 57 rue Cuvier, CP38, F-75005, Paris, France, ;

Abstract

The plesiosaur specimen NHMUK 36184 from the Lower Jurassic of Whitby, Yorkshire and kept in the Natural History Museum of London, comprises an almost entire skeleton with nearly complete skull. It was described as one of two syntypes of Plesiosaurus homalospondylus by Owen, and selected as the lectotype by Lydekker. Extensive preparation of the skull has revealed it as one of the most complete and best-preserved Jurassic plesiosaurian skulls known, and its description adds much-needed data to our knowledge of the cranial osteology of the Plesiosauria. The three-dimensional preservation permits a relatively reliable reconstruction of its form. Microcleidus homalospondylus displays an interesting combination of cranial characters present in Jurassic plesiosauroids and Cretaceous Elasmosauridea. Its snout presents a very distinctive sculpture; the first pair of premaxillae teeth are extremely reduced; the frontal is partially overlain by the premaxillae, contacts the pineal foramen but does not contact the temporal fenestra; the jugal does not contact the orbit nor the temporal fenestra; the squamosal contacts the postorbital but not the maxilla and presents a bulb; the postfrontal contacts the posterolateral orbit margin; the anterior interpterygoid vacuity is absent; the pterygoids meet posterior to the posterior interpterygoid vacuities and are pierced by a foramen at this level; the quadrate ramus of the pterygoid presents a ventromedial flange; the parasphenoid is crested; the epipterygoid contacts the parietal; the paroccipital process is spatulate distally; the prootic presents an anteroventral process; the mandibular symphysis is keeled and bears four pairs of teeth. Microcleidus appears very similar to Hydrorion and Occitanosaurus, and the three taxa share a great number of plesiomorphic characters with basal plesiosaurians and pliosauroids.

Type
Research Article
Copyright
Copyright © The Paleontological Society 

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Andrews, C. W. 1909. On some new Plesiosauria from the Oxford Clay of Peterborough. Annals and Magazine of Natural History, London, 48:418429.CrossRefGoogle Scholar
Andrews, C. W. 1910. A descriptive catalogue of the marine reptiles of the Oxford Clay, Part 1. British Museum (Natural History), London, 205 p., 10 pls.Google Scholar
Andrews, C. W. 1913. A descriptive catalogue of the Marine Reptiles of the Oxford Clay, Part II. British Museum (Natural History), London, 205p.Google Scholar
Bardet, N. 1992. Stratigraphic evidence for the extinction of the ichthyosaurs. Terra Nova, 4:649656.CrossRefGoogle Scholar
Bardet, N., Godefroit, P., and Sciau, J. 1999. A new elasmosaurid plesiosaur from the Lower Jurassic of Southern France. Palaeontology, 42:927952.CrossRefGoogle Scholar
Benson, R. B., Butler, R. J., Lindgren, J., and Smith, A. S. 2010. Mesozoic marine tetrapod diversity: Mass extinctions and temporal heterogeneity in geological megabiases affecting vertebrates. Proceedings of the Royal Society B, 277:829834.CrossRefGoogle ScholarPubMed
Benson, R. B. J., Ketchum, H. F., Noè, L. F., and Gomez-Perez, M. 2011a. New information on Hauffiosaurus (Reptilia, Plesiosauria) based on a new species from the Alum Shale Member (lower Toarcian: Lower Jurassic) of Yorkshire, U.K. Palaeontology, 54:547571.CrossRefGoogle Scholar
Benson, R. B. J., Bates, K. T., Johnson, M. R., and Withers, P. J. 2011b. Cranial anatomy of Thalassiodracon hawkinsii (Reptilia, Plesiosauria) from the Early Jurassic of Somerset, United Kingdom. Journal of Vertebrate Paleontology, 31:562574.CrossRefGoogle Scholar
Benson, R. B. J., Evans, M., and Druckenmiller, P. S. 2012. High diversity, low disparity and small body size in plesiosaurs (Reptilia, Sauropterygia) from the Triassic–Jurassic boundary. PLoS ONE, 7:e31838.CrossRefGoogle ScholarPubMed
Benton, M. J. and Taylor, M. A. 1983. Marine reptiles from the Upper Lias (lower Toarcian, Lower Jurassic) of the Yorkshire Coast. Proceedings of the Yorkshire Geological Society, 44:399429.CrossRefGoogle Scholar
Blainville, H. D. de. 1835. Description de quelques espèces de reptiles de la Californie, précédée de l'analyse d'un système général d'Erpétologie et d'Amphibiologie. Nouvelles Annales du Muséum (national) d'Histoire Naturelle de Paris, 4:233296.Google Scholar
Brown, D. S. 1981. The English Upper Jurassic Plesiosauroidea (Reptilia) and a review of the phylogeny and classification of the Plesiosauria. Bulletin of the British Museum (Natural History), Geology Series, 35:253347.Google Scholar
Brown, D. S. 1993. A taxonomic reappraisal of the families Elasmosauridae and Cryptoclididae (Reptilia: Plesiosauria). Revue de Paléobiologie, special volume, 7:916.Google Scholar
Cabrera, A. 1941. Un plesiosaurio nuevo del Cretáceo del Chubut. Revista del Museo de la Plata, 2:113130.Google Scholar
Carpenter, K. 1997. Comparative cranial anatomy of two North American Cretaceous plesiosaurs, p. 191216. InCallaway, J. M. and Nicholls, E. L.(eds), Ancient Marine Reptiles. Academic Press, San Diego, 501 p.CrossRefGoogle Scholar
Carpenter, K. 1999. Revision of North American elasmosaurs from the Cretaceous of the Western Interior. Paludicola, 2:148173.Google Scholar
Cheng, Y-N., Sato, T., Wu, X-C., and Li, C. 2006. First complete pistosauroid from the Triassic of China. Journal of Vertebrate Paleontology, 26:501504.CrossRefGoogle Scholar
Conybeare, W. D. 1824. On the discovery of an almost perfect skeleton of the Plesiosaurus. Transactions of the Geological Society of London, Second Series, 1:381389.CrossRefGoogle Scholar
Cruickshank, A. R. I. 1994a. Cranial anatomy of the Lower Jurassic pliosaur Rhomaleosaurus megacephalus (Stutchbury) (Reptilia: Plesiosauria). Philosophical Transactions of the Royal Society of London, Series B, 343:247260.CrossRefGoogle Scholar
Cruickshank, A. R. I. 1994b. A Juvenile plesiosaur (Plesiosauria: Reptilia) from the Lower Lias (Hettangian: Lower Jurassic) of Lyme Regis, England: A pliosauroid-plesiosauroid intermediate? Zoological Journal of the Linnean Society, 112:151178.CrossRefGoogle Scholar
Cruickshank, A. R. I. and Fordyce, R. E. 2002. A new marine reptile (Sauropterygia) from New Zealand: Further evidence for a late Cretaceous Austral radiation of cryptoclidid plesiosaurs. Palaeontology, 45:557575.CrossRefGoogle Scholar
Dames, W. 1895. Die Plesiosaurier der süddeutschen Liasformation. Abhandlungen der Königliche Preussische Akademie der Wissenschaften zu Berlin, 1895:183.Google Scholar
Druckenmiller, P. S. 2002. Osteology of a new plesiosaur from the lower Cretaceous (Albian) Thermopolis Shale of Montana. Journal of Vertebrate Paleontology, 22:2942.CrossRefGoogle Scholar
Druckenmiller, P. S. and Knutsen, E. M. 2012. Phylogenetic relationships of Upper Jurassic (middle Volgian) plesiosaurians (Reptilia: Sauropterygia) from the Agardhfjellet Formation of central Spitsbergen, Norway. Norwegian Journal of Geology, 92:277284.Google Scholar
Druckenmiller, P. S. and Russell, A. P. 2008. A phylogeny of Plesiosauria (Sauropterygia) and its bearing on the systematic status of Leptocleidus Andrews, 1922. Zootaxa, 1863:1120.CrossRefGoogle Scholar
Evans, M. 1999. A new reconstruction of the skull of the Callovian elasmosaurid plesiosaur Muraenosaurus leedsii Seely. Mercian Geologist, 14:191196.Google Scholar
Fritsch, K. von. 1894. Beitrag zur Kenntnis der Saurier des Halle'schen unteren Muschelkalkes. Abhandlungen der Naturforschenden Gesellschaft zu Halle, 20:273302.Google Scholar
Gasparini, Z. 1997. A new pliosaur from the Bajocian of the Neuquen Basin, Argentina. Paleontology, 40:135147.Google Scholar
Gasparini, Z., Salgado, L., and Casadıo, S. 2003a. Maastrichtian plesiosaurs from northern Patagonia. Cretaceous Research, 24:157170CrossRefGoogle Scholar
Gasparini, Z., Bardet, N., Martin, J. E., and Fernandez, M. 2003b. The elasmosaurid plesiosaur Aristonectes Cabrera from the latest Cretaceous of South America and Antarctica. Journal of Vertebrate Paleontology, 23:104115.CrossRefGoogle Scholar
Gray, J. E. 1825. A synopsis of the genera of reptiles and Amphibia, with a description of some new species. Annals of Philosophy, 26:193217.Google Scholar
Groβmann, F. 2007. Preliminary description and phylogenetic position of a new plesiosaur (Reptilia: Sauropterygia) from the Toarcian of Holzmaden, Germany. Journal of Paleontology, 78:973988.Google Scholar
Huene, F. R. von. 1923. Ein neuer Plesiosaurier aus dem oberen Lias Württembergs. Jahreshefte vaterländ. Naturkunde Württemberg, 79:123.Google Scholar
International Commission on Zoological Nomenclature. 1999. International Code of Zoological Nomenclature, fourth edition. International Trust for Zoological Nomenclature, London, 306p.Google Scholar
Kear, B. P. 2005. A new elasmosaurid plesiosaur from the Lower Cretaceous of Queensland, Australia. Journal of Vertebrate Paleontology, 25:792805.CrossRefGoogle Scholar
Kear, B. P. 2007. Taxonomic clarification of the Australian elasmosaurid genus Eromangasaurus, with reference to other austral elasmosaur taxa. Journal of Vertebrate Paleontology, 27:241246.CrossRefGoogle Scholar
Kear, B. P., Schroeder, N. I. and Lee, M. S. Y. 2006. An archaic crested plesiosaur in opal from the Lower Cretaceous high-latitude deposits of Australia. Biology Letters, 2:615619.CrossRefGoogle ScholarPubMed
Ketchum, H. F. and Benson, R. B. 2010. Global interrelationships of Plesiosauria (Reptilia, Sauropterygia) and the pivotal role of taxon sampling in determining the outcome of phylogenetic analyses. Biological Reviews, 85:361392.CrossRefGoogle ScholarPubMed
Ketchum, H. F. and Benson, R. B. 2011. A new pliosaurid (Sauropterygia, Plesiosauria) from the Oxford Clay Formation (Middle Jurassic, Callovian) of England: Evidence for a gracile longirostrine grade of Early-Middle Jurassic pliosaurids. Special Papers in Palaeontology, 86:109129.Google Scholar
Lydekker, R. 1889. Catalogue of the fossil Reptilia and Amphibia in the British Museum (Natural History). Part. II. Containing the orders Ichthyopterygia and Sauropterygia. British Museum (Natural History), London, 307p.Google Scholar
Maisch, M. 1998. Notes on the cranial osteology of Muraenosaurus Seeley, 1874 (Sauropterygia, Jurassic), with special reference to the neurocranium and its implications for sauropterygian phylogeny. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 207:207253.CrossRefGoogle Scholar
Meyer, H. von. 1842. Simosaurus, die Stumpfschnauze, ein Saurier as dem Muschelkalke von Luneville. Neues Jahrbuch für Mineralogie, Geognosie, Geologie und Petrefakten-Kunde, 1842:184197.Google Scholar
Noè, L. F. 2001. A taxonomic and functional study of the Callovian (Middle Jurassic) Pliosauroidea (Reptilia, Sauropterygia). Unpublished Ph.D. dissertation, University of Derby, UK, vol. 1, 354. p.; vol. 2, 182 p.Google Scholar
O'Keefe, R. F. 2001. A cladistic analysis and taxonomic revision of the Plesiosauria (Reptilia: Sauropterygia). Acta Zoologica Fennica, 213:163.Google Scholar
O'Keefe, R. F. 2004. Preliminary description and phylogenetic position of a new plesiosaur (Reptilia: Sauropterygia) from the Toarcian of Holzmaden, Germany. Journal of Paleontology, 78:973988.2.0.CO;2>CrossRefGoogle Scholar
O'Keefe, R. F. 2006. Morphologic and ontogenetic patterns in elasmosaur neck length, with comments on the taxonomic utility of neck length variables. Paludicola, 5:207229.Google Scholar
Otschev, V. G. 1976. A New Plesiosaur from the Upper Cretaceous of the Penza Region. Paleontologicheskii Zhurnal, 2:135138.Google Scholar
Otschev, V. G. 1977. On the Replacement of the Preoccupied Name Georgia penzensis. Paleontologicheskii Zhurnal, 4:118.Google Scholar
Owen, R. 1838. A Description of a Specimen of Plesiosaurus macrocephalus (Conybeare). Transactions of the Geological Society of London Series 2, 5:515535.CrossRefGoogle Scholar
Owen, R. 1860. On the orders of fossil and recent Reptilia, and their distribution in time. Reports of the British Association for the Advancement of Science, 29:153166.Google Scholar
Owen, R. 1865. Monograph on the fossil Reptilia of the Liassic formations, Part.1: Sauropterygia. The Palaeontographical Society, London, 40p.Google Scholar
Rieppel, O. 1989. A new pachypleurosaur (Reptilia: Sauropterygia) from the Middle Triassic of Monte San Giogio, Switzerland. Philosophical Transactions of the Royal Society of London, 323:173.Google Scholar
Rieppel, O. 1994. The braincase of Simosaurus and Nothosaurus: Monophyly of the Nothosauridae (Reptilia: Sauropterygia). Journal of Vertebrate Paleontology, 14:923.CrossRefGoogle Scholar
Rieppel, O., Sander, P. M., and Storrs, G. W. 2002. The skull of the pistosaur Augustasaurus from the Middle Triassic of northwestern Nevada. Journal of Vertebrate Paleontology, 22:577592.CrossRefGoogle Scholar
Sachs, S. 2005. Tuarangisaurus australis sp. nov. (Plesiosauria: Elasmosauridae) from the Lower Cretaceous of northeastern Queensland, with additional notes on the phylogeny of the Elasmosauridae. Memoirs of the Queensland Museum, 50:425440.Google Scholar
Sander, P. M. 1989. The pachypleurosaurids (Reptilia: Nothosauria) from the middle Triassic of Monte San Giorgio (Switzerland) with the description of a new species. Philosophical Transactions of the Royal Society of London, 325:61670.Google Scholar
Sander, P. M., Rieppel, O., and Bucher, H. 1997. A new pistosaurid (Reptilia, Sauropterygia) from the Middle Triassic of Nevada and its implications for the origin of the plesiosaurs. Journal of Vertebrate Paleontology, 17:526533.CrossRefGoogle Scholar
Sato, T. 2002. Description of plesiosaurs (Reptilia: Sauropterygia) from the Bearpaw Formation (Campanian–Maastrichtian) and a phylogenetic analysis of the Elasmosauridae. Unpublished Ph.D. dissertation, University of Calgary, Alberta, Canada, 391p.Google Scholar
Sato, T. 2003. Terminonatator ponteixensis, a new elasmosaur (Reptilia: Sauropterygia) from the Upper Cretaceous of Saskatchewan. Journal of Vertebrate Paleontology, 23:89103.CrossRefGoogle Scholar
Sato, T., Hasegawa, Y., and Manabe, M. 2006. A new elasmosaurid plesiosaur from the Upper Cretaceous of Fukushima, Japan. Palaeontology, 49:467484.CrossRefGoogle Scholar
Sato, T., Cheng, Y.-N., Wu, X.-C., and Li, C. 2010: Osteology of Yunguisaurus Cheng et al., 2006 (Reptilia; Sauropterygia), a Triassic pistosauroid from China. Paleontological research, 14:179195.CrossRefGoogle Scholar
Sato, T., Wu, X-C., Tirabasso, A., and Bloskie, P. 2011. Braincase of a polycotylid plesiosaur (Reptilia: Sauropterygia) from the Upper Cretaceous of Manitoba, Canada. Journal of Vertebrate Paleontology, 31:313329.CrossRefGoogle Scholar
Sauvage, M. H. E. 1873. Notes sur les reptiles fossiles. Bulletin de la Societe Geologique de France, Paris, 3:365380.Google Scholar
Sciau, J., Crochet, J. Y., and Mattei, J. 1990. Le premier squelette de plésiosaure de France sur le Causse du Larzac (Toarcien, Jurassique inférieur). Géobios, 23:111116.CrossRefGoogle Scholar
Seeley, H. G. 1865. On Plesiosaurus macropterus, a new species from the Lias of Whitby. Annals of the Magazine of Natural History, Series 3, 15:4953.CrossRefGoogle Scholar
Seeley, H. G. 1874. On Muraenosaurus leedsii, a plesiosaurian from the Oxford Clay, Part 1. Quarterly Journal of the Geological Society of London, 30:197208.CrossRefGoogle Scholar
Seeley, H. G. 1882. On Neusticosaurus pusillus (Fraas), an amphibious reptile having affinities with terrestrial Nothosauria and with marine Plesiosauria. Quarterly Journal of the Geological Society of London, 38:350366.CrossRefGoogle Scholar
Seeley, H. G. 1892. The nature of the shoulder girdle and clavicular arch in Sauropterygia. Proceedings of the Royal Society of London, 51:119151.Google Scholar
Sennikov, A. G. and Arkhangelsky, M. S. 2010. On a typical Jurassic sauropterygian from the Upper Triassic of Wilczek Land (Franz Josef Land, Arctic Russia). Paleontological Journal, 44:567572.CrossRefGoogle Scholar
Smith, A. S. 2003. Cladistic analysis of the Plesiosauria (Reptilia: Sauropterygia). MSc. thesis in palaeobiology, University of Bristol, 91p.Google Scholar
Smith, A. S. and Dyke, G. J. 2008. The skull of the giant predatory pliosaur Rhomaleosaurus cramptoni: Implications for plesiosaur phylogenetics. Naturwissenschaften, 95:975980.CrossRefGoogle ScholarPubMed
Smith, A. S. and Vincent, P. 2010. A new genus of pliosaur (Reptilia: Sauropterygia) from the Lower Jurassic of Holzmaden, Germany. Palaeontology, 53:10491063.CrossRefGoogle Scholar
Smith, A. S., Araújo, R., and Mateus, O. 2012. Lusonectes sauvagei, a new plesiosauroid from the Toarcian (Lower Jurassic) of Alhadas, Portugal. Acta Palaeontologica Polonica, 57:257266CrossRefGoogle Scholar
Storrs, G. W. 1991. Anatomy and Relationships of Corosaurus alcovensis (Diapsida: Sauropterygia) and the Triassic Alcova Limestone of Wyoming. Peabody Museum of Natural History Yale University, Bulletin, 44:1151.Google Scholar
Storrs, G. W. 1997. Morphological and taxonomic clarification of the genus Plesiosaurus, p. 145190. InCallaway, J. M. and Nicholls, E. L., (eds), Ancient Marine Reptiles. Academic Press, San Diego, 501 p.CrossRefGoogle Scholar
Storrs, G. W. and Taylor, M. A. 1996. Cranial anatomy of a new plesiosaur genus from the lowermost Lias (Rhaetian/Hettangian) of Street, Somerset, England. Journal of Vertebrate Paleontology, 16:403420.CrossRefGoogle Scholar
Sues, H. D. 1987. Postcranial skeleton of Pistosaurus and interrelationships of the Sauropterygia (Diapsida). Zoological Journal of the Linnean Society, 90:109131.CrossRefGoogle Scholar
Taylor, M. A. 1992. Functional anatomy of the head of the large aquatic predator Rhomaleosaurus zetlandicus (Plesiosauria; Reptilia) from the Toarcian (Lower Jurassic) of Yorkshire, England. Philosophical Transactions of the Royal Society of London, Series B, 335:247280.Google Scholar
Taylor, M. A. and Cruickshank, A. R. I. 1993. Cranial anatomy and functional morphology of Pliosaurus brachyspondylus (Reptilia: Plesiosauria). Philosophical Transactions of the Royal Society of London B, Biological Sciences, 341:399418.Google Scholar
Vincent, P. 2011. A re-examination of Hauffiosaurus zanoni, a Pliosauroidea from the Lower Jurassic of Germany. Journal of Vertebrate Paleontology, 31:340351.CrossRefGoogle Scholar
Vincent, P. 2012. Re-description of a basal plesiosaur (Reptilia, Sauropterygia) from the Lower Jurassic of England. Journal of Paleontology, 86:11871196.CrossRefGoogle Scholar
Vincent, P. and Benson, R. B. 2012. Anningasaura, a basal plesiosaurian (Reptilia, Plesiosauria) from the Lower Jurassic of Lyme Regis, United Kingdom. Journal of Vertebrate Paleontology, 32:10491063.CrossRefGoogle Scholar
Vincent, P. and Smith, A. S. 2009. A redescription of Plesiosaurus propinquus Tate and Blake, 1876 (Reptilia, Plesiosauria), from the Lower Jurassic (Toarcian) of Yorkshire, England. Proceedings of the Yorkshire Geological Society, 57:133142.CrossRefGoogle Scholar
Vincent, P., Bardet, N., Pereda Suberbiola, X., Bouya, B., Amaghzaz, M., and Meslouh, M. S. 2011. Zarafasaura oceanis, a new elasmosaurid (Reptilia: Sauropterygia) from the Maastrichtian Phosphates of Morocco and the palaeobiogeography of latest Cretaceous plesiosaurs. Gondwana Research, 19:10621073.CrossRefGoogle Scholar
Watson, D. M. S. 1909. A preliminary note on two new genera of Upper Liassic plesiosaurs. Memoirs and Proceedings of the Manchester Literary and Philosophical Society, 54:128.Google Scholar
Watson, D. M. S. 1911. The Upper Liassic Reptilia, Part III. Microcleidus macropterus (SEELEY) and the limbs of Microcleidus homalospondylus (OWEN). Memoirs of the Manchester Literary and Philosophical Society, 55:19.Google Scholar
Welles, S. P. 1943. Elasmosaurid plesiosaurs with a description of new material from California. Memoirs of the University of California, 13:125254.Google Scholar
Welles, S. P. 1949. A new elasmosaur from the Eagle Ford Shale of Texas. Fordren Science Series, 1:128.Google Scholar
Welles, S. P. 1962. A new species of elasmosaur from the Aptian of Colombia and a review of the Cretaceous plesiosaurs. University of California Publications in Geological Sciences, 46:196.Google Scholar
Wegner, T. 1914. Brancasaurus brancai n.g. n.sp., ein Elasmosauride aus dem Wealden Westfalens, p. 235305. InSchoendorf, F.et al., Branca-Festschrift. Verlag von Gebrüder Borntraeger, Leipzig.Google Scholar
Wiffen, J. and Moisley, W. L. 1986. Late Cretaceous reptiles (Families Elasmosauridae and Pliosauridae) from the Mangahouanga Stream, North Island, New Zealand. New Zealand Journal of Geology and Geophysics, 29:205252.CrossRefGoogle Scholar
Williston, S. W. 1890. A new plesiosaur from the Niobrara Cretaceous of Kansas. Transaction of the Kansas Academy of Science, 12:174178.Google Scholar
Williston, S. W. 1903. North American plesiosaurs, Part.1. Field Colombian Museum Publication (Geology), 73:177.Google Scholar
Williston, S. W. 1907. The skull of Brachauchenius, with special observations on the relationships of the plesiosaurs. United States National Museum Proceedings, 32:477489.CrossRefGoogle Scholar