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Novtautesamerican MUSEUM PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y

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Novtautesamerican MUSEUM PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y NovtautesAMERICAN MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3175, 34 pp., 23 figures June 26, 1996 Description of the Braincase of Two Early Cretaceous Pterosaurs (Pterodactyloidea) from Brazil ALEXANDER WILHELM ARMIN KELLNER' ABSTRACT The braincase of Tapejara wellnhoferi and An- sp. revealed the presence ofan interlaminar pneu- hanguera sp., two pterodactyloid taxa from the matic cavity with a complex system of trabeculae Early Cretaceous Santana Formation (Romualdo above the cranial cavity. Therefore, the actual Member) of the Araripe Basin, Brazil, are de- braincase is placed deeper inside the skull than scribed and compared with other archosaurian was previously supposed, raising questions about braincases. The presence of the orbitosphenoid, whether some pterosaur endocasts reported in the previously found in dinosaurs but absent in other literature express the true internal surface of the archosaurs, is reported in pterosaurs for the first braincase or just of the exocranial lamina. The time, suggesting that this bone is an ornithodiran braincase of Anhanguera sp. has some birdlike synapomorphy. Contrary to other archosaurs, there features (reduced olfactory bulbs and lateral place- is an ossification (the pseudomesethmoid) on the ment of the optic lobes) but its size indicates that anteroventral portion of those pterosaur brain- the pterosaur brain was more reptilian than was cases. A horizontal section through Anhanguera previously thought. INTRODUCTION The braincase of pterosaurs is very poorly ten damaged during deposition and diagen- known mainly for two reasons. First, because esis. The skull, in particular, is vulnerable to oftheir fragility, pterosaurian remains are of- crushing and flattening, severely limiting de- 1 Student, Department of Vertebrate Paleontology, American Museum of Natural History and Department of Geological Sciences, Lamont Doherty Geological Observatory, Columbia University; Fellow, Conselho Nacional de Desenvolvimento Cientifico e Tecnol6gico-CNPq, Brasilia. Copyright X American Museum of Natural History 1996 ISSN 0003-0082 / Price $4.20 2 AMERICAN MUSEUM NOVITATES NO. 3175 tailed anatomical studies of the posterior re- Terra-DNPM) in Rio de Janeiro, Brazil. gion. Second, specimens with three-dimen- Both were encased in calcareous nodules and sionally preserved bones are rare and valu- are preserved three dimensionally. able. Thus, detailed preparation ofthe brain- The most complete specimen (MCT 1500- case, which in many cases involves removing R) is referred to the tapejarid Tapejara welln- the posterior region ofthe skull or bones that hoferi. It comprises the skull and mandible are obstructing anatomical features, is often that were broken offand glued together (using discouraged. an epoxy compound) with the mandible and Very few studies of pterosaur braincases the anterior portion ofthe skull inverted. Un- are reported in the literature. Seeley (1870, fortunately, this and other artificial restora- 1871, 1901) was among the first researchers tions, is common practice among fossil col- to describe pterosaur cranial material in some lectors in the Araripe region. The second detail. Most of the specimens he studied, specimen (MCT 1501 -R) is referred to the however, were fragmentary, and therefore Anhangueridae and comprises only the dor- provided only limited information about the soposterior region of the skull. Both speci- braincase. Newton (1888) described the skull mens were prepared in detail, revealing sev- of Parapsicephalus purdoni from the Alum eral features previously unknown for ptero- Shale (Upper Lias), but focused primarily on saurs. the endocast. Edinger (1927, 1941) described specimens from Solnhofen and the Cam- bridge Greensand, but also centered her ob- ABBREVIATIONS servations on endocasts. More recently, bo basioccipital Wellnhofer (1985) and Campos and Kellner bs basisphenoid (1985) described three-dimensionally pre- clr columellar recess served pterosaurian skulls with preserved dep depression braincases from Brazil. Those specimens, eo exoccipital however, were not fully prepared and their f frontal observation were limited to the dermal skull fm foramen magnum roof and the occipital region. Bennett (1991) fo foramen described the braincase ofPteranodon in some gr groove detail, but the specimens are crushed, lim- ifl intermediate frontal lamina iting some observations. ios interorbital septum This paper describes two pterosaur brain- i jugal cases found in Cretaceous rocks of the Ar- la lacrimal aripe Basin, Northeast Brazil, and compares ls laterosphenoid them with braincases from other archosaurs. n nasal The material studied here comes from the oc occipital condyle Romualdo Member, upper stratigraphic unit op opisthotic of the Santana Formation (Beurlen, 1971). Os orbitosphenoid The Santana Formation is known worldwide p parietal for the quantity and exquisite preservation pcf postcranial fenestra of fossil remains (see Maisey, 1991, for a re- pfo pneumatic foramen view). The Romualdo Member essentially Po postorbital comprises shales of Albian age (Lima, 1978; pr prootic Pons et al., 1990), which contain calcareous psm pseudomesethmoid nodules. Fossils are found in the nodules and ptf posttemporal fenestra in the shales, but only the former are excep- q quadrate tionally well preserved. res resin The two pterosaur specimens described sac sagittal crest here belong to the Desiree Collection, which scp sclerotic plate is housed permanently in the Paleontology so supraoccipital Section of the Departamento Nacional da sq squamosal Produgcio Mineral (Museu de Ciencias da utf upper temporal fenestra 1 996 KELLNER: BRAINCASE, CRETACEOUS PTEROSAURS 3 (1 or r following abbreviations indicates left pared, allowing a complete view of the spec- or right side; roman numerals indicate fo- imen. Subsequently, a horizontal section was ramina for cranial nerves.) cut through the top of the skull (embedded in resin), allowing preparation of the pneu- MATERLALS AND METHODS matic cavity (or interlaminar pneumatic cav- The pterosaur braincases studied here were ity) present between the external (exocranial) prepared utilizing chemical techniques. Al- and internal (endocranial) laminae of the though the employment of acids in fossils is skull. a very old method (see Kellner, 1995a, for a Some material was removed mechanically review), the refinement of this technique for from the interlaminar pneumatic cavity of vertebrate fossils was done only in the 1950s the anhanguerid braincase and etched in a (Rixon, 1949; Toombs and Rixon, 1959). 3% formic acid solution. After drying, sam- The procedures adopted here were devel- ples were mounted on a conductive material oped particularly for three-dimensional tet- (carbon-based tape). Some were coated with rapods preserved in limestone (Kellner, 1991, a gold-palladium layer, while others were 1995a; Rutzky et al., 1994). This technique coated with a carbon layer. Both were ex- is carried out by immersing the specimen in amined by scanning electron microscope varying concentrations of formic acid solu- (SEM model: Zeiss DSM 950). The carbon- tions, which are saturated with calcium phos- coated samples were submitted to energy dis- phate to prevent bone deterioration. The persive spectroscopy for elemental charac- specimen is subsequently neutralized in plain terization in order to establish the compo- water and dried. The exposed parts of the sition of several parts of the specimen (e.g., bone are protected with different solutions of bone, matrix). a methacrylate resin (Paraloid B-72; Acriloid Comparisons were made with Recent spec- B-67) and the specimen is further immersed imens ofcrocodilians and birds to help iden- in the acid solution. This procedure is re- tify the cranial foramina. Previous interpre- peated until the desired preparation stage is tations of other fossil archosaur braincases reached. were also used, particularly the descriptions Due to the fragility of both specimens, a available forAllosaurus (Madsen, 1976), Sin- weak 3% formic acid solution was used. In raptor (Currie and Zhao, 1993a), Troodon some exceptionally delicate areas, a thin coat (Currie, 1985; Currie and Zhao, 1 993b), Syn- ofcyanoacrylate adhesive (super glue) diluted tarsus (Raath, 1985), Sphenosuchus (Walker, in acetone was applied to provide greater sta- 1990), Rhamphorhynchus (Wellnhofer, 1975), bility during the preparation process. Anhanguera (Wellnhofer, 1985; Campos and Several bones were covered with a reddish Kellner, 1985), and Pteranodon (Bennett, layer rich in iron, which was formed second- 1991). The anatomical terminology follows arily during diagenesis. This layer was not Baumel and Witmer (1993), using the English dissolved by acid, but had to be removed equivalents of the Latin terms. mechanically. The tapejarid specimen (MCT 1500-R) was SYSTEMATIC PALEONTOLOGY submitted to CT-Scan during an intermedi- ate preparation stage. The results, however, To date, 17 different pterosaur species have were limited, apparently due to the small been named from the Romualdo Member (for density contrast between the bone and the a review see Campos and Kellner, 1985; Kell- calcareous matrix. ner, 1990; Wellnhofer, 1991 b; Dalla Vecchia, The anhanguerid braincase was divided 1993; Kellner and Campos, 1994). This rel- during the fossilization processes on a hori- atively high number of named taxa seems to zontal plane passing through the posttem- be inflated, especially
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