Morphology, microanatomy, and histology of titanosaur (Dinosauria, Sauropoda) osteoderms from the Upper Cretaceous of Patagonia

LITERATURE CITED

• Avilla, L. S., R. Fernández, and D. F. B. Ramos. 2004. Bite marks on a crocodylomorph from the Upper Cretaceous of Brazil: evidence of social behavior? Journal of Vertebrate Paleontology 24:971–973.
   Web of Science ®Google Scholar
• Azevedo, S. A. K., and A. W. A. Kellner. 1998. A titanosaurid (Dinosauria, Sauropoda) osteoderm from the Upper Cretaceous of Minas Gerais, Brazil. Boletim Do Museu Nacional (Río de Janeiro) 44:1–6.
   Google Scholar
• Barrett, P. M., J. B. Clarke, D. B. Brinkman, S. D. Chapman, and P. C. Ensom. 2002. Morphology, histology and identification of the ‘granicones’ from the Purbeck Limestone Formation (Lower Cretaceous: Berriasian) of Dorset, southern England. Cretaceous Research 23:279–295.
   Web of Science ®Google Scholar
• Bonaparte, J. F., and J. E. Powell. 1980. A continental assemblage of tetrapods from the Upper Cretaceous beds of El Brete, northwestern Argentina (Sauropoda-Coelurosauria-Carnosauria-Aves). Mémoires de la Société Géologique de France 139:19–28.
   Google Scholar
• Buchwitz, M., F. Witzmann, S. Voigt, and V. Golubev. 2012. Osteoderm microstructure indicates the presence of a crocodylian-like trunk bracing system in a group of armoured basal tetrapods. Acta Zoologica 93:260–280.
   Web of Science ®Google Scholar
• Buffrénil, V. de, J. O. Farlow, and A. de Ricqlès. 1986. Growth and function of Stegosaurus plates: evidence from bone histology. Paleobiology 12:459–473.
   Web of Science ®Google Scholar
• Buffrénil, V. de, J.-Y. Sire, and J.-C. Rage. 2010. The histological structure of glyptosaurine osteoderms (Squamata: Anguidae), and the problem of osteoderm development in squamates. Journal of Morphology 267:729–737.
   Google Scholar
• Burns, M. E. 2008. Taxonomic utility of ankylosaur (Dinosauria, Ornithischia) osteoderms: Glyptodontopelta mimus Ford, 2000: a test case. Journal of Vertebrate Paleontology 28:1102–1109.
   Web of Science ®Google Scholar
• Butler R. J., A. Yates, O. W. M. Rauhut, and C. Foth. 2013. A pathological tail in a basal sauropodomorph dinosaur from South Africa: evidence of traumatic amputation? Journal of Vertebrate Paleontology 33:224–228.
   Web of Science ®Google Scholar
• Cerda, I. A., and J. B. Desojo. 2011. Dermal armour histology of aetosaurs (Archosauria: Pseudosuchia), from the Upper Triassic of Argentina and Brazil. Lethaia 44:416–428.
   Web of Science ®Google Scholar
• Cerda, I. A., and J. E. Powell. 2010. Dermal armor histology of Saltasaurus loricatus, an Upper Cretaceous sauropod dinosaur from Northwest Argentina. Acta Palaeontologica Polonica 55:389–398.
   Web of Science ®Google Scholar
• Cerda, I. A., L. Salgado, and J. E. Powell. 2012. Extreme postcranial pneumaticity in sauropod dinosaurs from South America. Paläontologische Zeitschrift 86:441–449.
   Web of Science ®Google Scholar
• Chiappe, L. M., R. A. Coria, L. Dingus, F. Jackson, A. Chinsamy, and M. Fox. 1998. Sauropod dinosaur embryos from the Late Cretaceous of Patagonia. Nature 396:258–261.
   Web of Science ®Google Scholar
• Chinsamy, A., and M. A. Raath. 1992. Preparation of fossil bone for histological examination. Palaeontologia africana 29:39–44.
   Google Scholar
• Chinsamy-Turan, A. 2005. The Microstructure of Dinosaur Bone: Deciphering Biology with Fine-Scale Techniques. Johns Hopkins University Press, Baltimore and London, 195 pp.
   Google Scholar
• Csiki, Z. 1999. New evidence of armoured titanosaurids in the Late Cretaceous–Magyarosaurus dacus from the Hateg Basin (Romania). Oryctos 2:93–99.
   Google Scholar
• Curry-Rogers, K., M. D. D’Emic, M. Vickaryous, and A. Cagan. 2011. Sauropod dinosaur osteoderms from the Late Cretaceous of Madagascar. Nature Communications 2:564. doi: 10.1038/ncomms1578.
   PubMed Web of Science ®Google Scholar
• D’Emic, M. D., J. A. Wilson, and S. Chatterjee. 2009. The titanosaur (Dinosauria: Sauropoda) osteoderm record: review and first definitive specimen from India. Journal of Vertebrate Paleontology 29:165–177.
   Web of Science ®Google Scholar
• Depérét, C. 1896. Note sur le dinosauriens sauropodes et théropodes du Crétacé Supérieur de Madagascar. Bulletin de la Société Géologique de France 24:176–194.
   Google Scholar
• Díez Díaz, V., X. Pereda Suberbiola, and J. L. Sanz. 2013. Appendicular skeleton and dermal armour of the Late Cretaceous titanosaur Lirainosaurus astibiae (Dinosauria: Sauropoda) from Spain. Palaeontologia Electronica 16:2; 19A; 18p. Available at www.palaeo-electronica.org/content/2013/502-titanosaur-skeleton.
   Google Scholar
• Dingus, L., J. Clarke, G. R. Scott, C. C. Swisher III, L. M. Chiappe, and R. A. Coria. 2000. Stratigraphy and magnetostratigraphic/faunal constraints for the age of sauropod embryo-bearing rocks in the Neuquén Group (Late Cretaceous, Neuquén Province, Argentina). American Museum Novitates 3290:1–11.
   Google Scholar
• Dodson, P., D. W. Krause, C. A. Forster, S. D. Sampson, and F. Ravoavy. 1998. Titanosaurid (Sauropoda) osteoderms from the Late Cretaceous of Madagascar. Journal of Vertebrate Paleontology 18:563–568.
   Web of Science ®Google Scholar
• Erickson, G. M., and C. M. Brochu. 1999. How the ‘terror crocodile’ grew so big. Nature 398:205–206.
   Web of Science ®Google Scholar
• Erickson, G. M., A. de Ricqlès, V. de Buffrénil, R. E. Molnar, and M. K. Bayless. 2003. Vermiform bones and the evolution of gigantism in Megalania—how a reptilian fox became a lion. Journal of Vertebrate Paleontology 23:966–970.
   Web of Science ®Google Scholar
• Francillon-Vieillont, H., V. de Buffrénil, J. Castanet, J. Geraudie, F. J. Meurier, J. Y. Sire, L. Zylberberg, and A. de Ricqlès. 1990. Microstructure and mineralization of vertebrate skeletal tissues; pp. 471–530 in J. G. Carter (ed.), Skeletal Biomineralization: Patterns, Processes and Evolutionary Trends, Volume 1. Van Nostrand Reinhold, New York.
   Google Scholar
• García, R. A., and L. Salgado. 2013. The titanosaur sauropods from the late Campanian–early Maastrichtian Allen Formation of Salitral Moreno, Río Negro, Argentina. Acta Palaeontologica Polonica 58:269–284.
   Web of Science ®Google Scholar
• Gomani, E. M. 2005. Sauropod dinosaurs from the Early Cretaceous of Malawi, Africa. Palaeontologia Electronica 8: Issue 1; 27A:37p. Available at http://palaeo-electronica.org/paleo/2005_1/gomani27/issue1_05.htm.
   Web of Science ®Google Scholar
• González Riga, B. J. 2003. A new titanosaur (Dinosauria, Sauropoda) from the Upper Cretaceous of Mendoza Province, Argentina. Ameghiniana 40:155–172.
   Web of Science ®Google Scholar
• Haines, R. W., and A. Mohuiddin. 1968. Metaplastic bone. Journal of Anatomy 103:527–538.
   PubMed Web of Science ®Google Scholar
• Hayashi, S., K. Carpenter, and D. Suzuki. 2009. Different growth pattern between the skeleton and osteoderms of Stegosaurus (Ornithischia: Thyreophora). Journal of Vertebrate Paleontology 29:123–131.
   Web of Science ®Google Scholar
• Hayashi, S., K. Carpenter, M. Watabe, and L. A. D. McWhinney. 2012. Ontogenetic histology of Stegosaurus plates and spikes. Palaeontology 55:145–161.
   Web of Science ®Google Scholar
• Hayashi, S., K. Carpenter, T. M. Scheyer, M. Watabe, and D. Suzuki. 2010. Function and evolution of ankylosaur dermal armor. Acta Palaeontologica Polonica 55:213–228.
   Web of Science ®Google Scholar
• Heredia, S. E., and L. Salgado, 1999. Posición estratigráfica de los sustratos supracretácicos portadores de dinosaurios en Lago Pellegrini, Patagonia septentrional, Argentina. Ameghiniana 36:229–234.
   Google Scholar
• Hill, R. V. 2006. Comparative anatomy and histology of xenarthran osteoderms. Journal of Morphology 267:1441–1460.
   PubMed Web of Science ®Google Scholar
• Hill, R. V. 2010. Osteoderms of Simosuchus clarki (Crocodyliformes: Notosuchia) from the Late Cretaceous of Madagascar. Society of Vertebrate Paleontology Memoir 10:154–176.
   Web of Science ®Google Scholar
• Hill, R. V., and S. G. Lucas. 2006. New data on the anatomy and relationships of the Paleocene crocodylian Akanthosuchus langstoni. Acta Palaeontologica Polonica 51:455–464.
   Web of Science ®Google Scholar
• Huene, F. von. 1929. Los saurisquios y ornitisquios del Cretáceo Argentino. Anales del Museo de La Plata (Series 2) 3:1–196.
   Google Scholar
• Hutton, J. M. 1986. Age determination of living Nile crocodiles from the cortical stratification of bone. Copeia 1986:332–341.
   Web of Science ®Google Scholar
• Katsura, Y. 2004. Paleopathology of Toyotamaphimeia machikanensis (Diapsida, Crocodylia) from the Middle Pleistocene of Central Japan. Historical Biology 16:2–4.
   Google Scholar
• Kellner, A. W. A., D. A. Campos, Z. A. K. Azevedo, M. N. F. Trota, D. D. R Enriques, M. M. T. Craik, and H. Paula Silva. 2006. A new titanosaur sauropod from the Bauru Group, Late Cretaceous of Brazil. Boletim do Museu Nacional 74:1–31.
   Google Scholar
• Klein, N., T. Scheyer, and T. Tütken. 2009. Skeletochronology and isotopic analysis of a captive individual of Alligator mississippiensis Daudin, 1802. Fossil Record 12:121–131.
   Web of Science ®Google Scholar
• Le Loeuff, J., E. Buffetaut, L. Cavin, M. Martin, V. Martin, and H. Tong, 1994. An armoured titanosaurid sauropod from the Late Cretaceous of Southern France and the occurrence of osteoderms in the Titanosauridae. Gaia 10:155–159.
   Google Scholar
• Levrat-Calviac, V., and L. Zylberberg. 1986. The structure of the osteoderms in the gekko: Tarentola mauritanica. The American Journal of Anatomy 176:437–466.
   PubMedGoogle Scholar
• Lindoso, R. M., T. S., Marinho, R. M. Santucci, M. A. Medeiros, and I. S. Carvalho. 2013. A titanosaur (Dinosauria: Sauropoda) osteoderm from the Alcântara Formation (Cenomanian), São Luís Basin, Northeastern Brazil. Cretaceous Research 45:43–48.
   Web of Science ®Google Scholar
• Main, R. P., A. de Ricqlès, J. R. Horner, and K. Padian. 2005. The evolution and function of thyreophoran dinosaur scutes: implications for plate function in stegosaurs. Paleobiology 31:291–314.
   Web of Science ®Google Scholar
• Malkani, M. S. 2003. Pakistani Titanosauria; are armored dinosaurs? Geological Bulletin of University of Peshawar 36:85–91.
   Google Scholar
• Marinho, T. S., and C. R. A. Candeiro. 2005. Titanosaur (Dinosauria: Sauropoda) osteoderms from the Maastrichtian of Uberaba, Minas Gerais State, Brazil. Gondwana Research 8:473–477.
   Web of Science ®Google Scholar
• Marinho, T. S., and F. V. Iori. 2011. A large titanosaur (Dinosauria, Sauropoda) osteoderm with possible bite marks from Ibirá, São Paulo State, Brazil; pp. 369–379 in I. D. Souza Carvalho, N. K. Srivastava, O. Strohschoen Jr., and C. C. Lana (eds.), Paleontologia: Cenários de Vida, Volume 4. Editorial Interciência, Rio de Janeiro.
   Google Scholar
• Martinelli, A. G., and A. M. Forasiepi. 2004. Late Cretaceous vertebrates from Bajo de Santa Rosa (Allen Formation), Río Negro province, Argentina, with the description of a new sauropod dinosaur (Titanosauridae). Revista del Museo de Ciencias Naturales 6:257–305.
   Google Scholar
• Molnar, R. E. 2011. New morphological information about Cretaceous sauropod dinosaurs from the Eromanga Basin, Queensland, Australia. Alcheringa 35:329–339.
   Web of Science ®Google Scholar
• O’Leary, M. A., E. M. Roberts, J. J. Head, F. Sissoko, and M. L. Bouaré. 2004. Titanosaurian (Dinosauria: Sauropoda) remains from the ‘Continental Intercalaire’ of Mali. Journal of Vertebrate Paleontology 24:923–930.
   Web of Science ®Google Scholar
• Parker, W. G., M. R. Stocker, and R. R Irmis. 2008. A new desmatosuchine aetosaur (Archosauria: Suchia) from the Upper Triassic Tecovas Formation (Dockum Group) of Texas. Journal of Vertebrate Paleontology 28:692–701.
   Web of Science ®Google Scholar
• Powell, J. E. 1980. Sobre la presencia de armadura dérmica en algunos dinosaurios titanosáuridos. Acta Geológica Lilloana 15:41–47.
   Google Scholar
• Powell, J. E. 2003. Revision of South American titanosaurid dinosaurs: palaeobiological, palaeobiogeographical and phylogenetic aspects. Records of the Queen Victoria Museum 111:1–173.
   Google Scholar
• Ramos, V. A. 1981. Descripción geológica de la Hoja 33 c. Los Chihuidos Norte, Provincia del Neuquén. Servicio Geológico Nacional, Boletín 182:1–103.
   Google Scholar
• Reid, R. E. H. 1996. Bone histology of the Cleveland-Lloyd dinosaurs and of dinosaurs in general. Part I: Introduction to bone tissues. Brigham Young University Geology Studies 41:25–72.
   Google Scholar
• Ricqlès, A. de, F. J. Meunier, J. Castanet, and H. Francillon-Vieillot. 1991. Comparative microstructure of bone; pp. 1–78 in B. B. K. Hall (ed.), Bone, Volume 3: Bone Matrix and Bone Specific Products. CRC Press, Boca Raton, Florida.
   Google Scholar
• Ricqlès, A. de, X. Pereda Suberbiola, Z. Gasparini, and E. Olivero. 2001. Histology of the dermal ossifications in an ankylosaurian dinosaur from the Late Cretaceous of Antartica. Asociación Paleontológica Argentina, Publicación Especial 7:171–174.
   Google Scholar
• Ruibal, R., and V. Shoemaker. 1984. Osteoderms in anurans. Journal of Herpetology 18:313–328.
   Web of Science ®Google Scholar
• Salgado, L. 1996. Pellegrinisaurus powelli nov. gen. et sp. (Sauropoda, Titanosauridae) from the Upper Cretaceous of Lago Pellegrini, Northwestern Patagonia, Argentina. Ameghiniana 33:355–365.
   Google Scholar
• Salgado, L. 2000. Evolución y paleobiología de los saurópodos titanosáuridos. Ph.D. dissertation, Universidad Nacional de La Plata, La Plata, Argentina, 300 pp.
   Google Scholar
• Salgado, L. 2003. Considerations on the bony plates assigned to titanosaurs (Dinosauria, Sauropoda). Ameghiniana 40:441–456.
   Web of Science ®Google Scholar
• Salgado, L., and C. Azpilicueta. 2000. Un nuevo saltasaurino (Sauropoda, Titanosauridae) de la Provincia de Río Negro (Formación Allen, Cretácico Superior), Patagonia, Argentina. Ameghiniana 37:259–264.
   Web of Science ®Google Scholar
• Salgado, L., and J. F. Bonaparte. 2007. Sauropodomorpha; pp. 188–228 in Z. Gasparini, L. Salgado, and R. A. Coria (eds.), Patagonian Mesozoic Reptiles. Indiana University Press, Bloomington, Indiana.
   Google Scholar
• Salgado, L., and R. A. Coria, 1993. El género Aeolosaurus (Sauropoda, Titanosauridae) en la Formación Allen (Campaniano–Maastrichtiano) de la provincia de Río Negro. Ameghiniana 30:119–128.
   Google Scholar
• Salgado, L., S. Apesteguía, and S. E. Heredia. 2005. A new specimen of Neuquensaurus australis, a Late Cretaceous saltasaurine titanosaur from North Patagonia. Journal of Vertebrate Paleontology 25:623–634.
   Web of Science ®Google Scholar
• Sander, P. M. 2000. Long bone histology of the Tendaguru sauropods: implications for growth and biology. Paleobiology 26:466–488.
   Web of Science ®Google Scholar
• Sander, P. M., N. Klein, E. Buffetaut, G. Cuny, V. Suteethorn, and J. Le Loeuff. 2004. Adaptive radiation in sauropod dinosaurs: bone histology indicates rapid evolution of giant body size through acceleration. Organisms, Diversity & Evolution 4:165–173.
   Web of Science ®Google Scholar
• Sanz, J. L., and A. D. Buscalioni. 1987. New evidence of armored dinosaurs in the Upper Cretaceous of Spain, pp. 199–204 in P. M. Currie and E. H. Koster (eds.), 4th Symposium of Mesozoic Terrestrial Ecosystems, Drumheller, Alberta, 10–14 August 1987. Short Papers. Royal Tyrrell Museum, Paleontology, Drumheller, Alberta.
   Google Scholar
• Scheyer, T. M., and M. R. Sánchez-Villagra. 2007. Carapace bone histology in the giant pleurodiran turtle Stupendemys geographicus: phylogeny and function. Acta Palaentologica Polonica 52:137–154.
   Web of Science ®Google Scholar
• Scheyer, T. M., and P. M. Sander. 2004. Histology of ankylosaur osteoderms: implications for systematics and function. Journal of Vertebrate Paleontology 20:874–893.
   Web of Science ®Google Scholar
• Scheyer, T. M., and P. M. Sander. 2007. Shell bone histology indicates terrestrial palaeoecology of basal turtles. Proceedings of the Royal Society B 274:1885–1893.
   Google Scholar
• Scheyer, T. M., and P. M. Sander. 2009. Bone microstructures and mode of skeletogenesis in osteoderms of three pareiasaur taxa from the Permian of South Africa. Journal of Evolutionary Biology 22:1153–1162.
   PubMed Web of Science ®Google Scholar
• Scheyer, T. M., B. Brüllmann, and M. R. Sánchez-Villagra. 2008. The ontogeny of the shell in side-necked turtles, with emphasis on the homologies of costal and neural bones. Journal of Morphology 269:1008–1021.
   PubMed Web of Science ®Google Scholar
• Scheyer, T. M., M. P. Sander, W. G. Joyce, W. Böhme, and U. Witzel. 2007. A plywood structure in the shell of fossil and living softshelled turtles (Trionychidae) and its evolutionary implications. Organisms, Diversity & Evolution 7:136–144.
   Web of Science ®Google Scholar
• Sire, J.-Y., P. C. J. Donoghue, and M. K. Vickaryous. 2009. Origin and evolution of the integumentary skeleton in non-tetrapod vertebrates. Journal of Anatomy 214:409–440.
   PubMed Web of Science ®Google Scholar
• Soto, M., D. Perea, and A. Cambiaso. 2012. First sauropod (Dinosauria: Saurischia) remains from the Guichón Formation, Late Cretaceous of Uruguay. Journal of South American Earth Sciences 33:68–79.
   Web of Science ®Google Scholar
• Taborda, J. R., I. A. Cerda, and J. B. Desojo. 2013. Growth curve of Aetosauroides scagliai Casamiquela 1960 (Pseudosuchia: Aetosauria) inferred from osteoderm histology; pp. 413–423 in S. J. Nesbitt, J. B. Desojo, and R. B. Irmis (eds.), Anatomy, Phylogeny and Palaeobiology of Early Archosaurs and Their Kin. Special Publications of the Geological Society of London 379.
   Google Scholar
• Tucker, A. D. 1997. Validation of skeletochronology to determine age of freshwater crocodiles (Crocodylus johnstoni). Marine and Freshwater Research 48:343–351.
   Web of Science ®Google Scholar
• Uliana, M. A., and D. A. Dellape. 1981. Estratigrafía y evolución paleoambiental de la sucesión Maastrichtiano-Eoterciaria del Engolfamiento Neuquino (Patagonia Septentrional); pp. 673–711 in 8’ Congreso Geológico Argentino, San Luis, Argentina, September 20-26. Asociación Geológica Argentina.
   Google Scholar
• Upchurch, P., P. M. Barrett, and P. Dodson. 2004. Sauropoda; pp. 295–322 in D. B. Weishampel, P. Dodson, and H. Osmólska (eds.), The Dinosauria, second edition. University of California Press, Berkeley, California.
   Google Scholar
• Vickaryous, M. K., and B. K. Hall. 2008. Development of the dermal skeleton in Alligator mississippiensis (Archosauria, Crocodylia) with comments on the homology of osteoderms. Journal of Morphology 269:398–422.
   PubMed Web of Science ®Google Scholar
• Vickaryous, M. K., and J.-Y. Sire. 2009. The integumentary skeleton of tetrapods: origin, evolution, and development. Journal of Anatomy 214:441–464.
   PubMed Web of Science ®Google Scholar
• Wilson, J. A., and P. Upchurch. 2003. A revision of Titanosaurus Lydekker (Dinosauria-Sauropoda), the first dinosaurian genus with a ‘Gondwanan’ distribution. Journal of Systematic Palaentology 1:125–160.
   Web of Science ®Google Scholar
• Witzmann, F., and R. Soler-Gijón. 2008. The bone histology of osteoderms in temnospondyl amphibians and in the chroniosuchian Bystrowiella. Acta Zoologica 89:1–19.
   Google Scholar
• Wolff, E. D. S., D. W. Fowler, and J. W. Bonde. 2007. A possible case of necrotizing dermatitis in the crocodylian Diplocynodon, from the Oligocene of the Isle of Wight, United Kingdom. Historical Biology 19:203–207.
   Google Scholar
• Zylberberg, L., and J. Castanet. 1985. New data on the structure and growth of the osteoderms in the reptile Anguis fragilis L. (Anguidae, Squamata). Journal of Morphology 186:327–342.
   PubMed Web of Science ®Google Scholar