Braincase, neuroanatomy, and neck posture of Amargasaurus cazaui (Sauropoda, Dicraeosauridae) and its implications for understanding head posture in sauropods

LITERATURE CITED

• Alexander, R.McN. 1985. Mechanics of posture and gait of some large dinosaurs. Zoological Journal of the Linnean Society 83:1–25.
   Web of Science ®Google Scholar
• Apesteguía, S. 2007. The sauropod diversity of the La Amarga Formation (Barremian), Neuquén (Argentina). Cretaceous Research 12:533–546.
   Google Scholar
• Bakker, R.T. 1978. Dinosaur feeding behaviour and the origin of flowering plants. Nature 274:661–663.
   Web of Science ®Google Scholar
• Bakker, R.T. 1986. The Dinosaur Heresies: New Theories Unlocking the Mystery of the Dinosaurs and Their Extinction. William Morrow and Company, New York, 481 pp.
   Google Scholar
• Balanoff, A.M., G.S. Bever, and T. Ikejiri. 2010. The braincase of Apatosaurus (Dinosauria: Sauropoda) based on computed tomography of a new specimen with comments on variation and evolution in sauropod neuroanatomy. American Museum Novitates 3677:1–29.
   Web of Science ®Google Scholar
• Bonaparte, J.F. 1996. Dinosaurios de America del Sur. Museo Argentino de Ciencias Naturales ‘Bernardino Rivadavia,’ Buenos Aires, Argentina, 174 pp.
   Google Scholar
• Calvo, J.O. 1994. Jaw mechanics in sauropod dinosaurs. Gaia 10:183–193.
   Google Scholar
• Calvo, J.O., and A.W. Kellner. 2006. Description of a sauropod dinosaur braincase (Titanosauridae) from the Late Cretaceous Rio Colorado Subgroup, Patagonia. Anais da Academia Brasileira de Ciências 78:175–182.
   PubMed Web of Science ®Google Scholar
• Carballido, J.L., and P.M. Sander. 2014. Postcranial axial skeleton of Europasaurus holgeri, (Dinosauria, Sauropoda) from the Upper Jurassic of Germany: implications for sauropod ontogeny and phylogenetic relationships of basal Macronaria. Journal of Systematic Palaeontology 12:335–387.
   Web of Science ®Google Scholar
• Carballido, J.L., D. Pol, I. Cerda, and L. Salgado. 2011a. The osteology of Chubutisaurus insignis del Corro, 1975 (Dinosauria, Sauropoda) from the ‘middle’ Cretaceous of central Patagonia, Argentina. Journal of Vertebrate Paleontology 31:93–110.
   Web of Science ®Google Scholar
• Carballido, J.L., O.W. M. Rauhut, D. Pol, and L. Salgado. 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.
   Web of Science ®Google Scholar
• Chatterjee, S., and Z. Zheng. 2002. Cranial anatomy of Shunosaurus, a basal sauropod dinosaur from the Middle Jurassic of China. Zoological Journal of the Linnean Society 136:145–169.
   Web of Science ®Google Scholar
• Chatterjee, S., and Z. Zheng. 2005. Neuroanatomy and dentition of Camarasaurus lentus; pp. 199–211 in V. Tidwell and K. Carpenter (eds.), Thunder-lizards: The Sauropodomorph Dinosaurs. Indiana University Press, Bloomington and Indianapolis, Indiana.
   Google Scholar
• Christian, A. 2010. Some sauropods raised their necks—evidence for high browsing in Ehuelopus zdanskyi. Biology Letters 6:823–825.
   PubMed Web of Science ®Google Scholar
• Christian, A., and G. Dzemski. 2007. Reconstruction of the cervical skeleton posture of Brachiosaurus brancai Janensch, 1914 by an analysis of the intervertebral stress along the neck and a comparison with the results of different approaches. Fossil Record 10:37–48.
   Google Scholar
• Clark, A.H. 2005. On the vestibular labyrinth of Brachiosaurus brancai. Journal of Vestibular Research 15:65–71.
   PubMed Web of Science ®Google Scholar
• Curry Rogers, K.A. 2009. The postcranial osteology of Rapetosaurus krausei (Sauropoda: Titanosauria) from the Late Cretaceous of Madagascar. Journal of Vertebrate Paleontology 29:1046–1086.
   Web of Science ®Google Scholar
• Curry Rogers, K.A., and C. Forster. 2004. The skull of Rapetosaurus krausei (Sauropoda: Titanosauria) from the Late Cretaceous of Madagascar. Journal of Vertebrate Paleontology 24:121–144.
   Web of Science ®Google Scholar
• Dzemski, G., and A. Christian. 2007. Flexibility along the neck of the ostrich (Struthio camelus) and consequences for the reconstruction of dinosaurs with extreme neck length. Journal of Morphology 268:701–714.
   PubMed Web of Science ®Google Scholar
• Fraas, E. 1908. Ostafrikanische Dinosaurier. Palaeontographica 55:105–144.
   Google Scholar
• Franzosa, J.W. 2004. Evolution of the brain in Theropoda (Dinosauria). Ph.D. dissertation, The University of Texas at Austin, Austin, Texas, 357 pp.
   Google Scholar
• Galton, P.M. 1985. Cranial anatomy of the prosauropod dinosaur Plateosaurus from the Knollenmergel (Middle Keuper, Upper Triassic) of Germany. Geologica et Palaeontologica 19:119–159.
   Google Scholar
• García, R., A. Paulina Carabajal, and L. Salgado. 2008. Un nuevo basicráneo de titanosaurio de la Formación Allen (Campaniano–Maastrichtiano), Provincia de Río Negro, Patagonia, Argentina. Geobios 41:625–633.
   Web of Science ®Google Scholar
• Gee, C.T. 2011. Dietary options for the sauropod dinosaurs from an integrated botanical and paleobotanical perspective; pp. 34–56 in N. Klein, K. Remes, C.T. Gee, and P.M. Sander (eds.), Biology of the Sauropod Dinosaurs. Indiana University Press, Bloomington and Indianapolis, Indiana.
   Google Scholar
• Gilmore, C.W. 1925. A nearly complete articulated skeleton of Camarasaurus, a saurischian dinosaur from the Dinosaur National Monument, Utah. Memoirs of the Carnegie Museum 10:347–384.
   Google Scholar
• Gilmore, C.W. 1936. Osteology of Apatosaurus, with special reference to specimens in the Carnegie Museum. Memoirs of the Carnegie Museum 11:172–298.
   Google Scholar
• Harris, J.D. 2006. Cranial osteology of Suuwassea emiliae (Sauropoda: Diplodocoidea: Flagellicaudata) from the Upper Jurassic Morrison Formation of Montana, U.S.A. Journal of Vertebrate Paleontology 26:88–102.
   Web of Science ®Google Scholar
• Harris, J.D., and P. Dodson. 2004. A new diplodocoid sauropod dinosaur from the Upper Jurassic Morrison Formation of Montana, U.S.A. Acta Palaeontologica Polonica 49:197–210.
   Web of Science ®Google Scholar
• Hatcher, J.B. 1901. Diplodocus (Marsh): its osteology, taxonomy, and probable habits, with a restoration of the skeleton. Memoirs of the Carnegie Museum 1:1–63.
   Google Scholar
• Holland, W.J. 1906. The osteology of Diplodocus Marsh. Memoirs of the Carnegie Museum 2:225–278.
   Google Scholar
• Hopson, J.A. 1979. Paleoneurology; pp. 39–146 in C. Gans (ed.), Biology of the Reptilia, Volume 9: Neurology. Academic Press, London.
   Google Scholar
• Hummel, J., and M. Clauss. 2011. Sauropod feeding and digestive physiology; pp. 11–33 in N. Klein, K. Remes, C.T. Gee, and P.M. Sander (eds.), Biology of the Sauropod Dinosaurs. Indiana University Press, Bloomington and Indianapolis, Indiana.
   Google Scholar
• Janensch, W. 1914. Übersicht über die Wirbeltierfauna der Tendaguruschichten, nebst einer kurzen Charakterisierung der neu aufgeführten Arten von Sauropoden. Archiv für Biontologie 3:81–110.
   Google Scholar
• Janensch, W. 1929. Die Wirbelsäule der Gattung Dicraeosaurus. Palaeontographica Supplement 7:39–133.
   Google Scholar
• Janensch, W. 1935–1936. Die Schädel der Sauropoden Brachiosaurus, Barosaurus und Dicraeosaurus aus den Tendaguru-Schichten Deutsch-Ostafrikas. Palaeontographica Supplement 7:147–298.
   Google Scholar
• Janensch, W. 1950. Die Wirbelsäule von Brachiosaurus brancai. Palaeontographica Supplement 7:27–93.
   Google Scholar
• Knoll, F., and D. Schwarz-Wings. 2009. Paleoneuroanatomy of Brachiosaurus. Annales de Paléontologie 95:165–175.
   Web of Science ®Google Scholar
• Knoll F., P.M. Galton, and R. López-Antoñanzas. 2006. Paleoneurological evidence against a proboscis in the sauropod dinosaur Diplodocus. Geobios 39:215–221.
   Web of Science ®Google Scholar
• Knoll F., R.C. Ridgely, F. Ortega, J.L. Sanz, and L.M. Witmer. 2013. Neurocranial Osteology and Neuroanatomy of a Late Cretaceous Titanosaurian Sauropod from Spain (Ampelosaurus sp.). PLoS ONE 8(1): e54991. doi:10.1371/journal.pone.0054991
   PubMed Web of Science ®Google Scholar
• Knoll, F., L.M. Witmer, F. Ortega, R.C. Ridgely, and D. Schwarz-Wings. 2012. The braincase of the basal sauropod dinosaur Spinophorosaurus and 3D reconstructions of the cranial endocast and inner ear. PLoS ONE 7:e30060. doi: 10.1371/journal.pone.0030060.
   PubMed Web of Science ®Google Scholar
• Marsh, O.C. 1877. Notice of new dinosaurian reptiles from the Jurassic formations. American Journal of Science, Series 3 14:514–516.
   Google Scholar
• Marsh, O.C. 1880. Principal characters of American Jurassic dinosaurs. Part 3. American Journal of Science, Series 3 19:253–259.
   Google Scholar
• Marsh, O.C. 1884. Principal characters of American Jurassic dinosaurs. Part VII. On the Diplodocidae, a new family of the Sauropoda. American Journal of Science, Series 3 27:161–168.
   Google Scholar
• Martin, J. 1987. Mobility and feeding of Cetiosaurus: why the long neck? Occasional Paper of the Tyrrell Museum of Paleontology 3:150–155.
   Google Scholar
• Martinelli, A.G., and A.M. Forasiepi. 2004. Late Cretaceous Vertebrates from Bajo de Santa Rosa (Allen Formation), Rio Negro province, Argentina, with the description of a new sauropod dinosaur (Titanosauridae). Revista Museo Argentino Ciencias Naturales 6:257–305.
   Google Scholar
• Osborn, H.F. 1912. Crania of Tyrannosaurus and Allosaurus. Memoirs of the American Museum of Natural History 1:1–30.
   Google Scholar
• Osborn, H.F., and C.C. Mook. 1921. Camarasaurus, Amphicoelias, and other sauropods of Cope. Memoirs of the American Museum of Natural History 3:247–287.
   Google Scholar
• Ostrom, J.H., and J.S. McIntosh. 1966. Marsh's Dinosaurs. Yale University Press, New Haven, Connecticut, 388 pp.
   Google Scholar
• Ouyang, H., and Y. Ye. 2002. The First Mamenchisaurian Skeleton with Complete Skull: Mamenchisaurus youngi. Sichuan Science and Technology Press, Chengdu, China, 88 pp. [Chinese, with English summary]
   Google Scholar
• Paulina Carabajal, A. 2012a. First insights into the dicraeosaurid (Sauropoda: Diplodocoidea) inner ear: the endocranial morphology of Amargasaurus cazaui studied using CT scans. Journal of Vertebrate Paleontology, Program and Abstracts 2012:155A.
   Google Scholar
• Paulina Carabajal, A. 2012b. Neuroanatomy of titanosaurid dinosaurs from the Upper Cretaceous of Patagonia, with comments on endocranial variability within Sauropoda. The Anatomical Record 295:2141–2156.
   Google Scholar
• Paulina Carabajal A., and L. Salgado. 2007. El basicráneo de un titanosaurio (Dinosauria, Sauropoda) del Cretácico Superior del norte de Patagonia: descripción y aportes al conocimiento del oído interno de los dinosaurios. Ameghiniana 44:109–120.
   Google Scholar
• Paulina Carabajal, A., R.A. Coria, and L.M. Chiappe. 2008. An incomplete Late Cretaceous braincase (Sauropoda: Titanosauria): new insights about the dinosaurian inner ear and endocranium. Cretaceous Research 29:643–648.
   Web of Science ®Google Scholar
• Rauhut, O.W. M., K. Remes, R. Fechner, G. Cladera, and P. Puerta. 2005. Discovery of a short-necked sauropod dinosaur from the Late Jurassic period of Patagonia. Nature 435:670–672.
   PubMed Web of Science ®Google Scholar
• Remes, K. 2006. Revision of the Tendaguru sauropod dinosaur Tornieria africana (Fraas) and its relevance for sauropod paleobiogeography. Journal of Vertebrate Paleontology 26:651–669.
   Web of Science ®Google Scholar
• Remes, K., F. Ortega, I. Fierro, U. Joger, R. Kosma, J.M. Marín Ferrer, O.A. Ide, and A. Maga. 2009. A new basal sauropod dinosaur from the Middle Jurassic of Niger and the early evolution of Sauropoda. PLoS ONE 4:e6924. doi: 10.1371/journal.pone.0006924.
   PubMed Web of Science ®Google Scholar
• Salgado, L. 1996. The macroevolution of the Diplodocimorpha (Dinosauria; Sauropoda): a developmental model. Ameghiniana 36:203–216.
   Google Scholar
• Salgado, L., and J.F. Bonaparte. 1991. Un nuevo saurópodo Dicraeosauridae, Amargasaurus cazaui gen. et sp. nov., de la Provincia del Neuquén, Argentina. Ameghiniana 28:333–346.
   Google Scholar
• Salgado, L., and J.O. Calvo. 1992. Cranial osteology of Amargasaurus cazaui Salgado and Bonaparte (Sauropoda, Dicraeosauridae) from the Neocomian of Patagonia. Ameghiniana 29:337–346.
   Google Scholar
• Salgado, L., R.A. Coria, and L.M. Chiappe. 2005. Osteology of the sauropod embryos from the Upper Cretaceous of Patagonia. Acta Palaeontologica Polonica 50:79–92.
   Web of Science ®Google Scholar
• Sander, P.M., and M. Clauss. 2008. Perspective: sauropod gigantism. Science 322:200–201.
   PubMed Web of Science ®Google Scholar
• Sander, P.M., O. Mateus., T. Laven, and N. Knötschke. 2006. Bone histology indicates insular dwarfism in a new Late Jurassic sauropod dinosaur. Nature 441:739–741.
   PubMed Web of Science ®Google Scholar
• Sander, P.M., A. Chirstian, M. Clauss, R. Fechner, C.G. Gee, E. Grieber, H. Guna, J. Hummel, H. Mallison, S.F. Perry, H. Preuschoft, O.W. M. Rauhut, K. Remes, T. Tütken, O. Wings, and U. Witzel. 2011. Biology of the sauropod dinosaurs: the evolution of gigantism. Biological Reviews 86:117–155.
   PubMed Web of Science ®Google Scholar
• Schwarz, D., and G. Fritsch. 2006. Pneumatic structures in the cervical vertebrae of the Late Jurassic Tendaguru sauropods Brachiosaurus brancai and Dicraeosaurus. Eclogae Geologicae Helvetiae 99:65–78.
   Google Scholar
• Schwarz, D., F. Eberhard, and C.A. Meyer. 2007. Novel reconstruction of the orientation of the pectoral girdle in sauropods. The Anatomical Record 290:32–47.
   Google Scholar
• Schweitzer, M.H., and C.L. Marshall. 2001. A molecular model for the evolution of endothermy in the theropod-bird lineage. Journal of Experimental Zoology 291:317–338.
   PubMedGoogle Scholar
• Sereno, P.C., J.A. Wilson, L.M. Witmer, J.A. Whitlock, A. Maga, O. Ide, and T.A. Rowe. 2007. Structural extremes in a Cretaceous dinosaur. PloS ONE 2:e21230. doi: 10.1371/journal.pone.0001230.
   Google Scholar
• Sereno, P.C., A.L. Beck, D.B. Dutheil, H.C. E. Larsson, G.H. Lyon, B. Moussa, R.W. Sadleir, C.A. Sidor, D.J. Varricchio, G.P. Wilson, and J.A. Wilson. 1999. Cretaceous sauropods from the Sahara and the uneven rate of skeletal evolution among dinosaurs. Science 286:1342–1347.
   PubMed Web of Science ®Google Scholar
• Snively, E., and A. Russell. 2007. Functional variation of neck muscles and their relation to feeding style in Tyrannosauridae and other large theropod dinosaurs. The Anatomical Record 290:934–957.
   Google Scholar
• Stevens, K.A., and J.M. Parrish. 1999. Neck posture and feeding habits of two Jurassic sauropod dinosaurs. Science 284:798–800.
   PubMed Web of Science ®Google Scholar
• Stevens, K.A., and J.M. Parrish. 2005. Neck posture, dentition, and feeding strategies in Jurassic sauropod dinosaurs; pp. 212–232 in V. Tidwell and K. Carpenter (eds.), Thunder-lizards: The Sauropodomorph Dinosaurs. Indiana University Press, Bloomington and Indianapolis, Indiana.
   Google Scholar
• Taylor, M.P., M.J. Wedel, and D. Naish. 2009. Head and neck posture in sauropod dinosaurs inferred from extant animals. Acta Palaeontologica Polonica 54:213–220.
   Web of Science ®Google Scholar
• Tschopp, E., and O. Mateus. 2013. The skull and neck of a new flagellicaudatan sauropod from the Morrison Formation and its implication for the evolution and ontogeny of diplodocid dinosaurs. Journal of Systematic Palaeontology 11:853–888.
   Web of Science ®Google Scholar
• Tsuihiji, T. 2005. Homologies of the transversospinalis muscles in the anterior presacral region of sauria (crown Diapsida). Journal of Morphology 263:151–178.
   PubMed Web of Science ®Google Scholar
• Tütken, T. 2011. The diet of sauropod dinosaurs–Implications from carbon isotope analysis of teeth, bones, and plants; pp. 57–79 in N. Klein, K. Remes, C.T. Gee, and P.M. Sander (eds.), Biology of the Sauropod Dinosaurs: Understanding the Life of Giants. Indiana University Press, Bloomington and Indianapolis, Indiana.
   Google Scholar
• Upchurch, P., and P.M. Barrett. 2000. The evolution of sauropod feeding mechanisms; pp. 79–122 in H.-D. Sues (ed.), The Evolution of Herbivory in Terrestrial Vertebrates, Perspectives from the Fossil Record. Cambridge University Press, Cambridge, U.K.
   Google Scholar
• Upchurch, P., and P.M. Barrett. 2005. A phylogenetic and taxic perspectives on sauropod diversity; pp. 104–124 in K.A. Curry Rogers and J.A. Wilson (eds.), The Sauropods: Evolution and Paleobiology. University of California Press, Berkeley, California.
   Google Scholar
• Upchurch, P., P.M. Barrett, and P. Dodson. 2004. Sauropoda; pp. 259–322 in D.B. Weishampel, P. Dodson, and H. Osmólska (eds.), The Dinosauria, second edition. University of California Press, Berkeley, California.
   Google Scholar
• White, T.E. 1958. The braincase of Camarasaurus lentus. Journal of Paleontology 32:477–494.
   Web of Science ®Google Scholar
• Whitlock, J.A. 2011a. A phylogenetic analysis of Diplodocoidea (Saurischia: Sauropoda). Zoological Journal of the Linnean Society 161:827–915.
   Web of Science ®Google Scholar
• Whitlock, J.A. 2011b. Inferences of diplodocid (Sauropoda: Dinosauria) feeding behavior from snout shape and microwear analyses. PLoS ONE 6:e18304. doi: 10.1371/journal.pone.0018304.
   PubMed Web of Science ®Google Scholar
• Wilson, J.A. 2002. Sauropod dinosaur phylogeny: critique and cladistic analysis. Zoological Journal of the Linnean Society 136:217–276.
   Web of Science ®Google Scholar
• Wilson, J.A. 2005. Redescription of the Mongolian sauropod Nemegtosaurus mongoliensis Nowinski (Dinosauria: Saurischia) and comments on Late Cretaceous Sauropod diversity. Journal of Systematic Palaeontology 3:238–318.
   Web of Science ®Google Scholar
• Wilson, J.A. 2009. Reassessment of the sauropod dinosaur Jainosaurus (= ‘Antarctosaurus’) septentrionalis from the Upper Cretaceous of India. Contributions from the Museum of Paleontology, University of Michigan 32:17–40.
   Google Scholar
• Wilson, J.A., and P.C. Sereno. 1998. Early evolution and higher level phylogeny of sauropod dinosaurs. Society of Vertebrate Paleontology Memoir 5:1–68.
   Google Scholar
• Wilson, J.A., M.D. D’Emic, T. Ikejiri, E.M. Moacdieh, and J.A. Whitlock. 2011. A nomenclature for vertebral fossae in sauropods and other saurischian dinosaurs. PLoS ONE 6:e17114. doi: 10.1371/journal.pone.0017114.
   PubMed Web of Science ®Google Scholar
• Witmer, L.M., and R. Ridgely. 2009. New insights into the brain, braincase, and ear region of tyrannosaurs (Dinosauria, Theropoda), with implications for sensory organization and behavior. The Anatomical Record 292:1266–1296.
   Google Scholar
• Witmer, L.M., R.C. Ridgely, D.L. Dufeau, and C. Semones. 2008. Using CT to peer into the past: 3D visualization of the brain and ear regions of birds, crocodiles, and nonavian dinosaurs; pp. 67–87 in H. Endo and R. Frey (eds.), Anatomical Imaging: Towards a New Morphology. Springer-Verlag, Tokyo.
   Google Scholar
• Zheng, Z. 1996. Cranial anatomy of Shunosaurus and Camarasaurus (Dinosauria: Sauropoda) and the phylogeny of the Sauropoda. Ph.D. dissertation, Texas Tech University, Lubbock, Texas, 208 pp.
   Google Scholar