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CARBALLIDO, J. L., MARPMANN, J. S., SCHWARZ-WINGS, D., PABST, B. 2012. New information on a juvenile sauropod specimen from the Morrison Formation and the reassessment of its systematic position. Palaeontology, 55, 3, 567–582.
José L. Carballido, Jean S. Marpmann, Daniela Schwarz-Wings and Ben Pabst Morphological changes in the ontogeny of sauropods are poorly known, making difficult to establish the systematic affinities of very young individuals. New information on an almost complete juvenile sauropod (SMA 0009) with an estimated total length of about 2 m is here presented. The specimen was described as a diplodocid owing to the presence of some putative synapomorphies of this group. However, recent further preparation revealed the absence of diplodocid characters and the presence of macronarian derived characters. To test the affinities of this specimen, a phylogenetic analysis was conducted. The strict consensus tree recovers the specimen as a basal titanosauriform, in an unresolved relation with Brachiosaurus and Giraffatitan. Nevertheless, a brachiosaurid assignment is here suggested in base of the widely accepted monophyly of this group (only recovered when SMA 0009 is placed within this group). Although the existence of a new taxon cannot be completely ruled out, the combination of derived and plesiomorphic characters in the specimen suggests its assignment to Brachiosaurus. Sixteen extra steps are needed to place this specimen within Diplodocidae. The high cost to place this specimen within this group is owing to the fact that several diplodocid characters are absent in SMA 0009, such as the absence of divided centroprezygapophyseal lamina in cervical vertebrae, procoelous anterior caudal centra, composed lateral lamina in anterior caudal vertebrae, elongated middle caudal vertebrae, short cervical ribs and caudolateral projection of distal condyle of metatarsal I. Finally, the systematic position reveals few major ontogenetic transformations. These affect the pneumatic structures (e.g. change from simple pleurocoels in the cervical vertebrae to complex pleurocoels and the development of lateral excavations in the dorsal vertebrae) but also include unrecorded transformations of the neural spine (e.g. the development of the spinodiapophyseal lamina, the widening of the neural spines in the dorsal vertebrae) and allometric growth in some limb bones.- ALLAIN, R. and AQUESBI, N. 2008. Anatomy and phylogenetic relationships of Tazoudasaurus naimi (Dinosauria, Sauropoda) from the late Early Jurassic of Morocco. Geodiversitas, 30, 345–424.
- APESTEGUÍA, S. 2005. The evolution of titanosaur metacarpus. 321–345. In TIDWELL, V. and CARPENTER, K. (eds). Thunder-lizards: the Sauropodomorph dinosaurs. Indiana University Press, Bloomington, Indianopolis, 495 pp.
- BARCO, J. L. 2009. Sistemática e implicaciones filogenéticas y paleobiogeográficas del saurópodo Galvesaurus herreroi (Formación Villar del Arzobispo, Galve, España). Universidad de Zaragoza, Zaragoza, 389 pp.
- BARCO, J. L., CANUDO, J. I., CUENCA-BESCÓS, G. and RUIZ-OMEÑACA, J. I. 2005. Un nuevo dinosaurio saurópodo Galvesaurus herreroi. Naturaleza Aragonesa, 15, 4–17.
- BONNAN, M. F. 2004. Morphometric analysis of humerus and femur shape in Morrison sauropods: implications for functional morphology and paleobiology. Paleobiology, 30, 444–470.
- CANUDO, J. I., ROYO-TORRES, R. and CUENCA-BESCÓS, G. 2008. A New Sauropod: Tastavinsaurus sanzi gen. et sp. nov. from the Early Cretaceous (Aptian) of Spain. Journal of Vertebrate Paleontology, 28, 712–731.
- CARBALLIDO, J. L., POL, D., CERDA, I. and SALGADO, L. 2011a. The osteology of Chubutisaurus insignis del Corro, 1975 (Dinosauria, Neosauropoda) from the ‘middle’ Cretaceous of Central Patagonia, Argentina. Journal of Vertebrate Paleontology, 31, 1–19.
- CARBALLIDO, J. L., RAUHUT, O. W., POL, D. and SALGADO, L. 2011b. Osteology and phylogenetic relationships of Tehuelchesaurus benitezii (Dinosauria, Sauropoda) from the Upper Jurassic of Patagonia. Zoological Journal of the Linnean Society, 163, 605–662.
- CARPENTER, K. and McINTOSH, J. 1994. Upper Jurassic sauropod babies from the Morrison Formation. 265–278. In CARPENTER, K., HIRSCH, K. F. and HORN, J. R. (eds). Dinosaur eggs and babies. Cambridge University Press, Cambridge, 372 pp.
- FOSTER, J. R. 2005. New juvenile sauropod material from western Colorado, and the record of juvenile sauropods from the Upper Jurassic Morrison Formation. 141–153. In TIDWELL, V. and CARPENTER, K. (eds). Thunder lizards. Indiana University Press, Indiana, 495 pp.
- GALLINA, P. A. and OTERO, A. 2009. Anterior caudal transverse processes in sauropod dinosaurs: morphological, phylogenetic and functional aspects. Ameghiniana, 46, 165–176.
- GALTON, P. M. 2000. The prosauropod dinosaur Plateosaurus engelhardti Meyer, 1837 (Saurischia: Sauropodomorpha). I. The syntypes of P. engelhardti Meyer, 1837 (Upper Triassic, Germany), with notes on other European prosauropods with ‘distally straight’ femora. Geologie und Palontologie Abhandlungen, 216, 233–275.
- GILMORE, C. W. 1936. Osteology of Apatosaurus, with special reference to specimens in the Carnegie Museum. Memoirs of the Carnegie Museum, XI, 172–298.
- GOLOBOFF, P. A., FARRIS, S. and NIXON, K. 2008a. TNT, a free program for phylogenetic analysis. Cladistics, 24, 774–786.
- GOLOBOFF, P. A., FARRIS, S. and NIXON, K. 2008b. TNT: Tree analysis using New Technology, vers. 1.1 (Will Henning Society Edition). Program and documentation. Available at: http://www.zmuc.dk/public/phylogeny/tnt.
- HARRIS, J. D. 2006. The significance of Suuwassea emilieae (Dinosauria: Sauropoda) for Flagellicaudatan intrarelationships and evolution. Journal of Systematic Palaeontology, 4, 185–198.
- HATCHER, J. B. 1901. Diplodocus (Marsh): its osteology, taxonomy, and probable habits, with a restoration of the skeleton. Memoirs of the Carnegie Museum, 1, 347–355.
- HATCHER, J. B. 1903. Osteology of Haplocanthosaurus with description of a new species, and remarks on the probable habits of the Sauropoda, and the age and origin of the Atlantosaurus beds. Additional Remarks on Diplodocus. Memoirs of the Carnegie Museum, 2, 1–75.
- IKEJIRI, T. 2004. Anatomy of Camarasaurus lentus (Dinosauria: Sauropoda) from the Morrison Formation (Late Jurassic), Thermopolis, central Wyoming, with determination and interpretation of ontogenetic, sexual dimorphic, and individual variation in the genus. Unpublished MS Thesis, University of Fort Hays State, Hays, 336 pp.
- IKEJIRI, T., TIDWELL, V. and TREXLER, D. L. 2005. New adult specimen of Camarasaurus lentus highlight ontogenetic variation within the species. 154–179. In TIDWELL, V. and CARPENTER, K. (eds). Thunder lizards. Indiana University Press, Indiana, 495 pp.
- JANENSCH, W. 1950. Die Wirbelsaüle von Brachiosaurus brancai. Paleontographica, 3 (Suppl 7), 27–93.
- KILBOURNE, B. M. and MAKOVICKY, P. J. 2010. Limb bone allometry during postnatal ontogeny in non-avian dinosaurs. Journal of Anatomy, 217, 135–152.
- KLEIN, N. and SANDER, M. 2008. Ontogenetic stages in the long bone histology of sauropod dinosaurs. Paleobiology, 34, 247–263.
- KSEPKA, D. T. and NORELL, M. A. 2010. The Illusory Evidence for Asian Brachiosauridae: new Material of Erketu ellisoni and a Phylogenetic Reappraisal of Basal Titanosauriformes. American Museum Novitates, 3700, 1–27.
- LEHMAN, T. M. and COULSON, A. B. 2002. A juvenile specimen of the sauropod dinosaur Alamosaurus sanjuanensis from the Upper Cretaceous of Big Bend National Park, Texas. Journal of Paleontology, 76, 156–172.
- LOVELACE, D. M., HARTMAN, S. A. and WILLIAM, R. 2007. Morphology of a specimen of Supersaurus (Dinosauria, Sauropoda) from the Morrison Formation of Wyoming, and a re-evaluation of diplodocid phylogeny. Arquivos do Museu Nacional, Rio de Janeiro, 65, 527–544.
- MANNION, P. D. and UPCHURCH, P. 2010a. A quantitative analysis of environmental associations in sauropod dinosaurs. Paleobiology, 32, 253–282.
- MANNION, P. D. and UPCHURCH, P. 2010b. Completeness metrics and the quality of the sauropodomorph fossil record through geological and historical time. Paleobiology, 32, 283–302.
- MARTIN, V., SUTEETHORN, V. and BUFFETAUT, E. 1999. Description of the type and referred material of Phuwiangosaurus sirindhornae Martin, Buffetaut and Suteethorn, 1994, a sauropod from the Lower Cretaceous of Thailand. Oryctos, 2, 39–91.
- McINTOSH, J. 2005. The Genus Barosaurus (Marsh). 38–77. In CARPENTER, K. and TIDWELL, T. (eds). Thunder lizards: the Sauropodomorph dinosaurus. Indiana University Press, Indiana, 503 pp.
- McINTOSH, J., MILLER, W. E., STADTMAN, K. L. and GILLETTE, D. D. 1996. The osteology of Camarasaurus lewisi (Jensen, 1988). Brigham Young University, Geology Studies, 41, 73–115.
- OSBORN, H. F. 1899. A skeleton of Diplodocus. Memoirs of the American Museum of Natural History, 1, 191–214.
- OSBORN, H. F. and MOOK, C. C. 1921. Camarasaurus, Amphicoellias and other sauropods of Cope. Memoirs of the American Museum of Natural History, 3, 247–387.
- RIGGS, E. S. 1903. Brachiosaurus altithorax, the largest known dinosaur. American Journal of Science, 4, 299–306.
- RIGGS, E. S. 1904. Structure and relationships of opisthocoelian dinosaurs. Part II, the Brachiosauridae. Field Columbian Museum, Geological Series, 2, 229–247.
- ROMER, A. S. 1956. Osteology of the Reptiles. University of Chicago Press, Chicago, 772 pp.
- ROSE, P. 2007. A new Titanosauriform sauropod (Dinosauria: Saurischia) from the Early Cretaceous of Central Texas and its phylogenetic relationships. Paleontologia Electronica, 10, 1–65.
- SALGADO, L. and POWELL, J. E. 2010. Reassessment of the vertebral laminae in some South American titanosaurian sauropods. Journal of vertebrate paleontology, 30, 1760–1772.
- SALGADO, L., CORIA, R. A. and CALVO, J. O. 1997. Evolution of titanosaurid sauropods. I: Phylogenetic analysis based on the postcranial evidence. Ameghiniana, 34, 3–32.
- SANDER, M. P., OCTÁVIO, M., THOMAS, L. and NILS, K. 2006. Bone histology indicates insular dwarfism in a new Late Jurassic sauropod dinosaur. Nature, 441, 739–741.
- SCHWARZ, D., IKEJIRI, T., BREITHAUPT, B. H., SANDER, P. M. and KLEIN, N. 2007. A nearly complete skeleton of an early juvenile diplodocid (Dinosauria: Sauropoda) from the Lower Morrison Formation (Late Jurassic) of north central Wyoming and its implications for early ontogeny and pneumaticity in sauropods. Historical Biology, 19, 225–253.
- SERENO, P. C., BECK, A. L., DUTHEIL, D. B., LARSSON, H. C. E., LYON, G. H., MOUSSA, B., SADLEIR, R. W., SIDOR, C. A., VARRICCHIO, D. J., WILSON, G. P. and WILSON, J. A. 1999. Cretaceous Sauropods from the Sahara and the Uneven Rate of Skeletal Evolution Among Dinosaurs. Science, 286, 1342–1347.
- TAYLOR, M. P. 2009. A re-evaluation of Brachiosaurus altithorax Riggs 1903 (Dinosauria, Sauropoda) and its generic separation from Giraffatitan brancai (Janensch 1914). Journal of Vertebrate Paleontology, 29, 787–806.
- TIDWELL, V. and WILHITE, R. D. 2005. Ontogenetic variation and isometric growth in the forelimb of the Early Cretaceous sauropod Venenosaurus. 187–196. In TIDWELL, V. and CARPENTER, K. (eds). Thunder-lizards: the Sauropodomorph dinosaurs. Indiana University Press, Bloomington, Indianopolis, 495 pp.
- TIDWELL, V., CARPENTER, K. and BROOKS, W. 1999. New sauropod from the Lower Cretaceous of Utah, USA. Oryctos, 2, 21–37.
- UPCHURCH, P. 1995. The Evolutionary History of Sauropod Dinosaurs. Philosophical Transactions: Biological Sciences, 349, 365–390.
- UPCHURCH, P. 1998. The phylogenetic relationships of sauropod dinosaurs. Zoological Journal of the Linnean Society, 124, 43–103.
- UPCHURCH, P. and MARTIN, J. 2002. The Rutland Cetiosaurus: the anatomy and relationships of a Middle Jurassic British sauropod dinosaur. Palaeontology, 45, 1049–1074.
- UPCHURCH, P., BARRET, P. M. and DODSON, P. 2004. The Sauropods. 259–354. In WEISHAMPEL, D. B., DODSON, P. and OSMÓLSKA, H. (eds). The dinosauria. University of California, Berkley, 880 pp.
- WILSON, J. A. 1999. A nomenclature for vertebral laminae in sauropods and other saurischian dinosaurs. Journal of Vertebrate Paleontology, 19, 639–653.
- WILSON, J. A. 2002. Sauropod dinosaur phylogeny: critique and cladistic analysis. Zoological Journal of the Linnean Society, 136, 215–275.
- WILSON, J. A. and SERENO, P. C. 1998. Early evolution and higher-level phylogeny of sauropod dinosaurs. Memoir (Society of Vertebrate Paleontology), 5, 1–68.
- WILSON, J. A. and UPCHURCH, P. 2003. A revision of Titanosaurus Lydekker (Dinosauria - Sauropoda), the first dinosaur genus with a “Gondwanan” distribution. Journal of Systematic Palaeontology, 1, 125–160.
- WILSON, J. A. and UPCHURCH, P. 2009. Redescription and reassessment of the phylogenetic affinities of Euhelopus zdanskyi (Dinosauria: Sauropoda) from the Early Cretaceous of China. Journal of Systematic Palaeontology, 7, 199–239.