title: The braincase of the basal sauropod dinosaur Spinophorosaurus and 3D reconstructions of the cranial endocast and inner ear.
abstract: BACKGROUND
  Sauropod dinosaurs were the largest animals ever to walk on land, and, as a result, the evolution of their remarkable adaptations has been of great interest. The braincase is of particular interest because it houses the brain and inner ear. However, only a few studies of these structures in sauropods are available to date. Because of the phylogenetic position of Spinophorosaurus nigerensis as a basal eusauropod, the braincase has the potential to provide key evidence on the evolutionary transition relative to other dinosaurs.
  
  
  RESULTS
  The only known braincase of Spinophorosaurus ('Argiles de l'Irhazer', Irhazer Group; Agadez region, Niger) differs significantly from those of the Jurassic sauropods examined, except potentially for Atlasaurus imelakei (Tilougguit Formation, Morocco). The basisphenoids of Spinophorosaurus and Atlasaurus bear basipterygoid processes that are comparable in being directed strongly caudally. The Spinophorosaurus specimen was CT scanned, and 3D renderings of the cranial endocast and inner-ear system were generated. The endocast resembles that of most other sauropods in having well-marked pontine and cerebral flexures, a large and oblong pituitary fossa, and in having the brain structure obscured by the former existence of relatively thick meninges and dural venous sinuses. The labyrinth is characterized by long and proportionally slender semicircular canals. This condition recalls, in particular, that of the basal non-sauropod sauropodomorph Massospondylus and the basal titanosauriform Giraffatitan.
  
  
  CONCLUSIONS
  Spinophorosaurus has a moderately derived paleoneuroanatomical pattern. In contrast to what might be expected early within a lineage leading to plant-eating graviportal quadrupeds, Spinophorosaurus and other (but not all) sauropodomorphs show no reduction of the vestibular apparatus of the inner ear. This character-state is possibly a primitive retention in Spinophorosaurus, but due the scarcity of data it remains unclear whether it is also the case in the various later sauropods in which it is present or whether it has developed homoplastically in these taxa. Any interpretations remain tentative pending the more comprehensive quantitative analysis underway, but the size and morphology of the labyrinth of sauropodomorphs may be related to neck length and mobility, among other factors.
authors.affiliation: Departamento de Paleobiología, Museo Nacional de Ciencias Naturales-CSIC, Madrid, Spain. knoll@mncn.csic.es
authors.forename: Fabien
authors.initials: F
authors.lastname: Knoll
authors.forename: Lawrence M
authors.initials: LM
authors.lastname: Witmer
authors.forename: Francisco
authors.initials: F
authors.lastname: Ortega
authors.forename: Ryan C
authors.initials: RC
authors.lastname: Ridgely
authors.forename: Daniela
authors.initials: D
authors.lastname: Schwarz-Wings
mesh_terms: D000818:Animals; D005075:Biological Evolution; D025061:Dinosaurs / Q000033:anatomy & histology* / Q000145:classification / Q000235:genetics; D007758:Ear, Inner / Q000033:anatomy & histology*; D005580:Fossils; D021621:Imaging, Three-Dimensional / Q000379:methods*; D008953:Models, Anatomic; D010163:Paleontology / Q000379:methods; D010802:Phylogeny; D012886:Skull / Q000033:anatomy & histology*; D019291:Skull Base / Q000033:anatomy & histology; D013997:Time Factors; D014057:Tomography, X-Ray Computed