Ecomorphology and bone microstructure of Proterochampsia from the Chanares Formation

• Autorzy: Arcucci A. , Previtera E. , Mancuso A.C.
• Języki publikacji: EN

Abstrakty

EN

Proterochampsians are a South American endemic group of non-archosaurian archosauriforms with morphological characteristics recollecting Recent crocodilians, and therefore have been proposed as aquatic species. However, this has not been based on careful examination of anatomical and histological features. We provide a review of the morphological and histological evidence present in the skeleton of proterochampsids and discuss its implications for inferring the lifestyles of these organisms. Anatomical features such as a secondary palate, marginal dentition, palatine teeth, morphology of the tail, limb modification, and dermal armor are reviewed, and details of histological structures are described based on bone thin sections. Histological examination reveals a predominance of fibrolamellar bone tissue, suggesting rapid periosteal osteogenesis and therefore overall fast bone growth. The existence of discontinuities (LAGs) demonstrates that these animals responded to changes in their environment. Ecomorphological features do not provide definitive evidence for the lifestyles of proterochampsids, but allow us to propose a terrestrial/amphibious condition. The same is true of the histological features, particularly compactness of the bone.

Słowa kluczowe

EN

Argentina; Triassic; Upper Triassic; Carnian; Archosauriformes; Proterochampsia; bone microstructure; fossil; fossil vertebrate; anatomical feature; morphological feature; ecomorphology; paleozoology; paleobiology

• Wydawca: -
• Czasopismo: Acta Palaeontologica Polonica
• Rocznik: 2019
• Tom: 64
• Numer: 1
• Opis fizyczny: p.157-170,fig.,ref.

Twórcy

autor

Arcucci A.

• Area de Zoologia, Universidad Nacional de San Luis, IMIBIO CONICET, Av. Ejercito de los Andes 950, 5700 San Luis, Argentina

autor

Previtera E.

• Instituto Argentino de Nivologia, Glaciologia y Ciencias Ambientales (IANIGLA), CCT-Mendoza, CONICET. Av. Adrian Ruiz Leal s/n, Parque General San Martin CC 330 (CP 5500) Mendoza, Argentina

autor

Mancuso A.C.

• Instituto Argentino de Nivologia, Glaciologia y Ciencias Ambientales (IANIGLA), CCT-Mendoza, CONICET. Av. Adrian Ruiz Leal s/n, Parque General San Martin CC 330 (CP 5500) Mendoza, Argentina

Bibliografia

• Arcucci, A.B. 1990. Un nuevo Proterochampsidae (Reptilia–Archosauriformes) de la fauna local de Los Chañares (Triásico Medio), La Rioja, Argentina. Ameghiniana 27: 365–378.
• Arcucci, A.B. 2011. Sistemática y filogenia de los Proterochampsidae (Amniota, Diápsida, Archosauriformes) del Triásico de América del Sur, y sus implicancias en el origen de Archosauria. 302 pp. Ph.D. Thesis, Universidad Nacional de San Juan, San Juan.
• Barberena, M.C. 1982. Uma nova espécie de Proterochampsa, P. nodosa sp. nov. do Triássico do Brasil. Anais da Academia Brasileira de Ciências 54: 127–141.
• Betz, O. 2006. Ecomorphology: Integration of form, function, and ecology in the analysis of morphological structures. Mitteilungen der Deutschen Gesellschaft für Allgemeine und Angewandte Entomologie 15: 409–416.
• Bininda-Emonds, O.R.P., Gittleman, J.L., and Kelly, C.K. 2001. Flippers versus feet: comparative trends in aquatic and non-aquatic carnivores. Journal of Animal Ecology 70: 386–400.
• Bock, W.J. 1994. Concepts and methods in ecomorphology. Journal of Biosciences 19: 403–413.
• Bonaparte, J.F. 1971. Cerritosaurus binsfeldi Price, tipo de una nueva familia de tecodontes (Pseudosuchia–Proterochampsia). Anais da Academia Brasileira de Ciencias 43: 417–422.
• Bonaparte, J.F. 1978. El Mesozoico de América del Sur y sus Tetrápodos. Opera Lilloana 26: 1–596.
• Bonaparte, J.F. 1997. El Triásico de San Juan-La Rioja Argentina y sus Dinosaurios.190 pp. Museo Argentino de Ciencias Naturales, Buenos Aires.
• Botha-Brink, J. and Smith, R.M.H. 2011. Osteohistology of the Triassic archosauromorphs Prolacerta, Proterosuchus, Euparkeria and Erythrosuchus from the Karoo Basin of South Africa. Journal of Vertebrate Paleontology 31: 1238–1254.
• Buffrénnil, V. de 1980. Mise en evidence de l’incidence des conditions de milieu sur la croissance de Crocodylus siamensis (Schneider, 1801) et valeur des marques de croissance squelettiques pour l’evaluation de l’age individual. Archives de Zoologie Experimentale Generale 121: 63–76.
• Buffrénil, V. de and Mazin, J.M. 1990. Bone histology of the ichthyosaurs: comparative data and functional interpretation. Paleobiology 16: 435– 447.
• Buffrénil, V. de, Mazin J.M., and Ricqlès, A.J. de 1987. Caractères structuraux et mode de croissance du fémur d’Omphalosaurus nisseri, ichthyosaurien du Trias moyen de Spitsberg. Annales De Paleontologie 73: 195–216.
• Buffrénil, V. de, Ricqlès, A.J. de, Ray, C.E., and Domning, D.P. 1990a. Bone histology of the ribs of the archaeocetes (Mammalia: Cetacea). Journal of Vertebrate Paleontology 10: 455–466.
• Buffrénil, V. de, Ricqlès, A.J. de, Sigogneau-Russell, D., and Buffetaut, E. 1990b. L’histologie osseuse des champsosauridés: données descriptives et interprétation fonctionnelle. Annales De Paleontologie 76: 255–275.
• Buffrénil, V. de, Canoville, A., D’Anastasio, R., Domning, and Daryl P., 2010. Evolution of sirenian pachyosteosclerosis, a model-case for the study of bone structure in aquatic tetrapods. Journal of Mammalian Evolution 17: 101–120.
• Canoville, A. and Laurin, M. 2009. Microanatomical diversity of the humerus and lifestyle in lissamphibians. Acta Zoológica 90: 110–122.
• Canoville, A. and Laurin, M. 2010. Evolution of humeral microanatomy and lifestyle in amniotes, and some comments on paleobiological inferences. Biological Journal of the Linnean Society 100: 384–406.
• Carroll, R.L. 1985. Evolutionary constraints in aquatic diapsid reptiles. Palaeontology 33: 45–155.
• Casinos, A. 1996. Bipedalism and quadrupedalism in Megatherium: an attempt at biomechanical reconstruction. Lethaia 29: 87–96.
• Castanet, J., Francillon-Vieillot, H., Meunier, F.J., and Ricqlès, A.J. de 1993. Bone and individual aging. In: B.K. Hall (ed.), Bone Growth, 245–283. CRC Press, London.
• Chinsamy, A. 1997. Assessing the biology of fossil vertebrates through bone histology. Palaeontologia Africana 33: 29–35.
• Chinsamy, A. and Raath, M.A. 1992. Preparation of fossil bone for histological examination. Palaeontologia Africana 29: 39–44.
• Chinsamy-Turan, A. 2005. The Microstructure of Dinosaur Bone. 216 pp. The Johns Hopkins University Press, Baltimore.
• Chinsamy-Turan, A. 2011. Forerunners of Mammals: Radiation, Histol ogy, Biology. 352 pp. Indiana University Press, Bloomington.
• Cerda, I.A. and Chinsamy, A. 2012. Biological implications of the bone microstructure of the Late Cretaceous ornithopod dinosaur Gasparinisaura cincosaltensis. Journal of Vertebrate Paleontology 32: 355–368.
• Cerda, I.A., Desojo, J.B., Trotteyn, M.J., and Scheyer, T.M. 2015. Osteoderm histology of Proterochampsia and Doswelliidae (Reptilia: Archosauriformes) and their evolutionary and paleobiological implications. Journal of Morphology 276: 385–402.
• Currey, J.D. 2002. Bones: Structure and Mechanics. 456 pp. Princeton University Press, Princeton.
• Curry, K.A. 1999. Ontogenetic histology of Apatosaurus (Dinosauria: Sauropoda) new insights on growth rates and longevity. Journal of Vertebrate Paleontology 19: 654–665.
• Dilkes, D. 1998. The Early Triassic rhynchosaur Mesosuchus browni and the interrelationships of basal archosauromorph reptiles. Philosophical Transactions of the Royal Society of London B 353: 501–541.
• Dilkes, D. and Arcucci, A. 2012. Proterochampsa barrionuevoi (Archosauriformes: Proterochampsia) from the Late Triassic (Carnian) of Argentina and a phylogenetic analysis of Proterochampsia. Palaeontology 5: 853–885.
• Dilkes, D. and Sues, H. 2009. Redescription and phylogenetic relationships of Doswellia kaltenbachi (Diapsida: Archosauriformes) from the Upper Triassic of Virginia. Journal of Vertebrate Paleontology 29: 58–79.
• Ewer, R.F. 1965. The anatomy of the thecodont reptile Euparkeria capensis Broom. Philosophical Transactions of the Royal Society of London B 248: 379–435.
• Ezcurra, M.D., Desojo, J.B., and Rauhut, O.W.M. 2015. Redescription and phylogenetic relationships of the proterochampsid Rhadinosuchus gracilis (Diapsida: Archosauriformes) from the early Late Triassic of southern Brazil. Ameghiniana 52: 391–417.
• Ezcurra, M.D., Lecuona, A., and Martinelli, A.G. 2010. A new basal diapsid archosauriform from the Early Triassic of Argentina. Journal of Vertebrate Paleontology 39: 1–18.
• Fish, F.E. and Stein, B.R. 1991. Functional correlates of differences in bone density among terrestrial and aquatic genera in the family Mustelidae (Mammalia). Zoomorphology 110: 339–345.
• Francillon-Vieillot, H., Buffrenil, V. de, Castanet, J., Geraudie, J., Meunier, F.J., Sire, J.Y., Zylberberg, L., and Ricqles, A. de 1990. Microstructure and mineralization of vertebrate skeletal tissues. In: J.G. Carter (ed.), Skeletal Biomineralization: Patterns, Processes and Evolutionary Trends, 471–530. Van Nostrand Reinhold 1, New York.
• Flynn, J.J., Nesbitt, S.J., Parrish, J.M., Ranivoharimanana, L., and Wyss, A.R. 2010. A new species of Azendohsaurus (Diapsida: Archosauromorpha) from the Triassic Isalo Group of Southwestern Madagascar: cranium and mandible. Palaeontology 53: 669–688.
• Garcia Marsà, J.A., Agnolín, F.L., and Novas, F. 2017. Bone microstructure of Lewisuchus admixtus Romer, 1972 (Archosauria, Dinosauri-formes). Historical Biology [published online, https://doi.org/10.1080 /08912963.2017.1347646]
• Gauthier, J.A. 1994. The diversification of the amniotes. In: D.R. Prothero and R.M. Schoch (eds.), Major Features of Vertebrate Evolution, 129–159. The Paleontological Society, Knoxville.
• Germain, D. and Laurin, M. 2005. Microanatomy of the radius and lifestyle in amniotes (Vertebrata Tetrapoda). Zoologica Scripta 34: 335–350.
• Horner, J.R., Ricqlès, A.J. de, and Padian, K. 1999. Variation in dinosaur skeletochronology indicators: Implications for age assessment and physiology. Paleobiology 25: 295–304.
• Horner, J.R., Ricqlès, A.J. de, and Padian, K. 2000. Long bone histology of the hadrosaurid dinosaur Maiasaura peeblesorum: growth dynamics and physiology based on an ontogenetic series of skeletal elements. Journal of Vertebrate Paleontology 20: 115–129.
• Houssaye, A., Lindgren J., Pellegrini R, Lee A.H., Germain, D., and Polcyn, M.J. 2013. Microanatomical and histological features in the long bones of mosasaurine mosasaurs (Reptilia, Squamata)—implications for aquatic adaptation and growth rates. PLoS ONE 8: e76741.
• Houssaye, A., Scheyer, T.M., Kolb, C., Fischer, V., and Sander, P.M. 2014. A new look at ichthyosaur long bone microanatomy and histology: Implications for their adaptation to an aquatic life. PLoS ONE 9: e95637.
• Hsiou, A., Adbala, F., and Arcucci, A.B. 2002. Novo registro de Proterocampsídeo (Reptilia, Archosauriformes) do Triássico Médio–Superior do Brasil. Revista Brasileira de Palentologia 3: 48–55.
• Huene, F. von 1938. Die fossilen Reptilien des südamerikanischen Gondwanalandes. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie 3: 142–151.
• Kardong, K.V. 1999. Vertebrados: Anatomía Comparada, Función, Evolución. 732 pp. McGraw-Hill Interamericana, Madrid.
• Keaveny, T.M., Morgan, E.F., and Yeh, O.C. 2003. Bone mechanics. In: M. Kurtz (ed.), Standard Handbook of Biomedical Engineering and Design, 8.1–8.23. McGraw-Hill Press, New York.
• Klein, N. 2012. Postcranial morphology and growth of the Winterswijk pachypleurosaur Anarosaurus heterodontus (Sauropterygia) from the Lower Muschelkalk of Winterswijk, The Netherlands. Paläontologische Zeitschrift 86: 389–408.
• Klein, N., Houssaye, A., Neenan, J.M., and Scheyer, T.M. 2015. Long bone histology and microanatomy of Placodontia (Diapsida: Sauropterygia). Contributions to Zoology 84: 59–84.
• Klein, N. and Sander, M. 2008. Ontogenetic stages in the long bone histology of sauropod dinosaurs. Paleobiology 34: 247–263.
• Kischlat, E.E. 2000. Tecodôncios: a aurora dos arcossáurios no Triássico. In: M. Holz and L.F. De Ross (eds.), Paleontologia do Rio Grande do Sul, 273–316. CIGO/UFRGS, Porto Alegre.
• Kriloff, A., Germain, D., Canoville, A., Vincent, P., Sache, M., and Laurin, M. 2008. Evolution of bone microanatomy of the tetrapod tibia and its use in palaeobiological inference. Journal of Evolutionary Biology 21: 807–826.
• Langer, M.C., Ramezani, J., and Da Rosa, A.A.S. 2018. U-Pb age constraints on dinosaur rise from south Brazil. Gondwana Research 57: 133–140.
• Laurin, M., Canoville, A., and Germain, D. 2011. Bone microanatomy and lifestyle: a descriptive approach. Comptes Rendu Palevol 10: 381–402.
• Laurin, M., Germain, D., Steyer, J.S., and Girondot, M. 2006. Données microanatomiques sur la conquête de l’environnement terrestre par les vertébrés. Comptes Rendu Palevol 5: 603–618.
• Laurin, M., Girondot, M., and Loth, M.M. 2004. The evolution of long bone microanatomy and lifestyle in lissamphibians. Paleobiology 30: 589–613.
• Laurin, M., Meunier, F.J., Germain D., and Lemoine, M. 2007. A microanatomical and histological study of the paired fin skeleton of the Devonian sarcopterygian Eusthenopteron foordi. Journal of Paleontology 81: 143–153.
• Le, P., Mouton, F.N., Geertsema, H., and Visagie, L. 2000. Foraging mode of a group-living lizard, Cordylus cataphractus (Cordylidae). African Zoology 35: 1–7.
• Lee, M.S.Y. 1997. Reptile relationships turn turtle. Nature 389: 245–246.
• Mancuso, A.C. 2005. Revisión y aportes a la estratigrafía de la sección inferior del Grupo Agua de la Peña (Triásico Medio, Argentina). XVI Congreso Geológico Argentino, La Plata, Argentina, Actas 3: 415–422.
• Mancuso, A.C., Gaetano, L.C., Leardi, J.M., Abdala, F., and Arcucci, A.B. 2014. The Chañares Formation: a window to the palaeobiology of a Middle Triassic vertebrate fauna. Lethaia 47: 244–265.
• Mancuso, A.C., Previtera, E., Benavente, C.A., and Hernández del Pino, S. 2017. Evidence of bacterial decay and early diagenesis in a partially-articulated tetrapod from the Triassic Chañares Formation. Palaios 32: 367–381.
• Marsicano, C.A., Irmis, R.B., Mancuso, A.C., Mundil, R., and Chemale, F. 2016. The precise temporal calibration of dinosaur origins. Proceedings of the National Academy of Sciences of the United States of America 113: 509–513.
• Martínez, R.N., Sereno, P.C., Alcober, O.A., Colombi, C.E., Renne, P.R., Montañez, I.P., and Currie, B.S. 2011. A basal dinosaur from the dawn of the dinosaur era in southwestern Pangaea. Science 331: 206–210.
• Massare, J.A. 1988. Swimming capabilities of Mesozoic marine reptiles. Paleobiology 14: 187–205.
• Nesbitt, S. 2011. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 352: 1–292.
• Nesbitt, S., Stocker, M.R., Small, B., and Downs, A. 2009. The osteology and relationships of Vancleavea campi (Reptilia: Archosauriformes). Zoological Journal of the Linnean Society 157: 814–864.
• Nesbitt, S.J., Desojo, J.B., and Irmis, R.B. 2013. Anatomy, phylogeny and palaeobiology of early archosaurs and their kin. Geological Society, London, Special Publications 379: 1–7.
• Nopcsa, F.B. 1923. Vorläufige Notiz über die Pachyostose und Osteosklerose einiger mariner Wirbeltiere. Anatomischer Anzeiger 56: 353–359.
• Nopcsa, F.B. and Heidsieck, E. 1934. Uber eine pachyostotische Rippe aus der Kreide Rugens. Acta Zoologica 15: 431–455.
• Padian, K. and Lamm, E.T. 2013. Bone Histology of Fossil Tetrapods: Advancing Methods, Analysis, and Interpretation. 285pp. University of California Press, Berkeley, California.
• Ponce, D.A., Cerda, I.A., Desojo, J.B., and Nesbitt, S.J. 2017. The osteoderm microstructure in doswelliids and proterochampsids and its implications for palaeobiology of stem archosaurs. Acta Palaeontologica Polonica 62: 819–831.
• Price, L.I. 1946. Sobre um novo pseudosuquio do Triássico Superior do Rio Grande do Sul. Boletim Divisão de Geologáa e Mineralogía 120: 1–38.
• Raugust, T., Lacerda, M., and Schultz, C.L. 2013. The first occurrence of Chanaresuchus bonapartei Romer 1971 (Archosauriformes, Proterochampsia) of the Middle Triassic of Brazil from the Santacruzodon Assemblage Zone, Santa Maria Formation (Paraná Basin). Geological Society London, Special Publications 379: 303–318.
• Reid, R.E.H. 1990. Zonal growth rings in dinosaurs. Modern Geology 15: 19–48.
• Reid, R.E.H. 1996. Bone histology of the Cleveland-Lloyd dinosaurs and of dinosaurs in general, Part I: Introduction: introduction to bone tissues. BrighamYoung University Research Studies. Geology 41: 25–72.
• Reig, O.A. 1958. A cerca de la posición sistemática de la Familia Rauisuchidae y del género Saurosuchus (Reptilia–Thecodontia). Publicaciones del Museo Municipal de Ciencias Naturales y Tradicionales de Mar del Plata 1: 73–114.
• Reig, O.A. 1959. Primeros datos descriptivos sobre nuevos reptiles arcosaurios del Triásico de Ischigualasto (San Juan, Argentina). Revista de la Asociación Geológica Argentina 13: 257–270.
• Ricqlès, A.J. de 1972. Recherches paléohistologiques sur les os longs des Tétrapodes. Part III, Titanosuchiens, Dinocéphales et Dicynodontes. Annales de Paléon tologie Vertébrés 58: 17–60.
• Ricqlès, A.J. de 1974a. Recherches paléohistologiques sur les os longs des tétrapodes IV. Eothériodontes et pélycosaures. Annales de Paléon tologie 60: 1–39.
• Ricqlès, A.J. de 1974b. Recherches paléohistologiques sur les os longs des tétrapodes V. Cotylosaures et mésosaures. Annales de Paléont ologie 60: 171–216.
• Ricqlès, A.J. de 1976. On bone histology of fossil and living reptiles, with comments on its functional and evolu tionary significance. In: B. Bellairs Cox (ed.), Morphology and Biology of Reptiles, 126–150. Linnean Society of London Academic Press, London.
• Ricqlès, A.J. de 1983. Cyclical growth in the long limb bones of a sauropod dinosaur. Acta Palaeontologica Polonica 28: 225–232.
• Ricqlès, A.J. de, and Buffrenil, V. de 2001. Bone histology, heterochronies and the return of tetrapods to life in water: where are we? In: J.M. Mazin and V. de Buffrenil (eds.), Secondary Adaptation of Tetrapods to Life in Water, 289–310. Verlag Dr. Friedrich Pfeil, München.
• Ricqlès, A.J. de, Padian, K., and Horner, J.R. 2003. On the bone histology of some Triassic pseudosuchian archosaurs and related taxa. Annales de Paléontologie 89: 67–101.
• Ricqlès, A.J. de, Padian, K., Knoll, F., and Horner, J.R. 2008. On the origin of high growth rates in archosaurs and their ancient relatives: Complementary histological studies on Triassic archosauriforms and the problem of a “phylogenetic signal” in bone histology. Annales de Paléontologie 94: 57–76.
• Rimblot-Baly, F., Ricqlès, A.J. de, and Zylberberg, L. 1995. Analyse paléohistologique d’une série de croissance partielle chez Lapparentosaurus madagascariensis (Jurassique moyen): essai sur la dynamique de croissance d’un Dinosaure Sauropode. Annales de Paléontologie 1: 49–86.
• Romer, A.S. 1966. Vertebrate Paleontology. Third Edition, 1–707. University of Chicago Press, Chicago.
• Romer, A.S. 1971. The Chañares (Argentina) Triassic reptile fauna. XI: two new long-snouted thecodonts, Chanaresuchus and Gualosuchus. Breviora 379: 1–22.
• Scheyer, T.M. and Sander, P.M. 2007. Shell bone histology indicates terrestrial palaeoecology of basal turtles. Proceedings of the Royal Society B 274: 1885–1893.
• Sereno, P.C. and Arcucci, A.B. 1990. The monophyly of the crurotarsal archosaurs and the origin of birds and crocodile ankle joints. Neues Jahrbuch fur Geologie und Paläontologie, Abhandlungen 180: 21–52.
• Sill, W.D. 1967. Proterochampsa barrionuevoi and the early evolution of the Crocodilia. Bulletin of the Museum of Comparative Zoology 135: 415–446.
• Smith-Gill, J.S. 1983. Developmental plasticity: developmental conversion versus developmental modulation. American Zoologist 23: 47–55.
• Sookias, R.B. and Butler, R.J. 2013. Euparkeriidae. Geological Society London, Special Publications 379: 35–48.
• Starck, J.M. and Chinsamy, A. 2002. Bone microstructure and developmental plasticity in birds and other dinosaurs. Journal of Morphology 254: 232–246.
• Stein, B.R. 1989. Bone density and adaptation in semiaquatic mammals. Journal of Mammalogy 70: 467–476.
• Steyer, J.S., Laurin, M., Castanet, J., and Ricqlès, A.J. de 2004. First histological and skeletochronological data on temnospondyl growth; palaeo ecological and palaeoclimatological implications. Palaeogeography, Palaeoclimatology, Palaeoecology 206: 193–201.
• Taylor, L.H. 1994. Stone, bone or blubber? Buoyancy control strategies in aquatic tetrapods. In: L. Maddock, Q. Bone, and J.M.V. Rayner (eds.), Mechanics and Physiology of Animal Swimming, 151–161. Cambridge University Press, Cambridge.
• Trotteyn, M.J. 2011. Material postcraneano de Proterochampsa barrionuevoi Reig, 1959 (Diapsida: Archosauriformes) del Triásico Superior del centro-oeste de Argentina. Ameghiniana 48: 424–446.
• Trotteyn, M.J., Arcucci, A.B., and Raugust, T. 2013. Proterochampsia: an endemic archosauriform clade from South America. Geological Society London, Special Publications 379: 59–90.
• Trotteyn, M.J. and Ezcurra, M.D. 2014. Osteology of Pseudochampsa ischigualastensis gen. et comb. nov. (Archosauriformes: Proterochampsidae) from the early Late Triassic Ischigualasto Formation of northwestern Argentina. PloS ONE 9: e111388.
• Van der Klaauw, C.J. 1948. Ecological studies and reviews. IV. Ecological morphology. Bibliotheory 4: 27–111.
• Wall, W. P. 1983. The correlation between high limb-bone density and aqua tic habits in recent mammals. Journal of Paleontology 57: 197– 207.
• Weems, R.E. 1980. An unusual newly discovered archosaur from the Upper Triassic of Virginia, U.S.A. Transactions of the American Philosophical Society 70: 1–53.
• Wiffen, J., Buffrénil, V. de, Ricqlès, A.J. de, and Mazin, J.M. 1995. Ontogenetic evolution of bone structure in Late Cretaceous Plesiosauria from New Zealand. Geobios 28: 625–640.

Identyfikatory

DOI

10.4202/app.00536.2018