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Pterosauria, Pterodactyloidea, Anhangueridae); CRANIUM 22,1 (2005) Brasileodactylus (Pterosauria, Pterodactyloidea, Anhangueridae); an update André+J. Veldmeijer Marco Signore and Hanneke Meijer Summary Brazil and The majority of the teethed pterosaurs from the Araripe Basin in (Santana Crato Formations) have premaxillary sagittal crests and dentary sagittal crests. A few skulls, however, lack such crests and ever since these sexual fossils have been published, discussion continueswhether these are independent taxa, ontogenetic variants or dimorphics of other taxa. The present work presents an update, discussing the dentitionand other morphological features, previously unnoticed, in order to evaluate the systematic position of the taxon. Samenvatting Brazilië Crato Het merendeel van de getande pterosauriërs die afkomstig zijn uit het Araripe Basin in (Santana en kam kam. Er echter die Formaties) hebben een premaxilla sagittale en een dentary sagittale zijn een paar taxa geen kammen hebben deze de schedel vanafhetallereerstemoment dat deze fossielen bekend is op plekken op en waren, of deze of dat deze dieren of sexueel dimorfische varianten er gespeculeerd een geldig taxon zijn ontogenetische zijn kort van andere taxa. Deze bijdrage presenteert een update waarin het gebit gepresenteerd en besproken wordt, alsmede enkele morfologische kenmerken dieeerder over het hoofdwerden gezien, met als doel de systematische evalueren. positie van het taxonBrasileodactylus te Introduction measurements are most of the time not included (exceptions are Fastnacht (2003), Lee (1994) and first this Ever since the pterosaur was discovered, Veldmeijer (2003b)). Therefore, measurement of of vertebrates has been the extraordinary group teethand alveoli are includedin the present work subject of much debate and discussion. This is and discussed. A few problems, however, occur skeleton partly dueto the fact that the pterosaur is which complicate the taxonomic issue. and there extremely fragile as a consequence, are only relatively few fossils and even fewer well preserved complete specimens which have been Materials and methods adequately prepared. Nowadays, there are a few sites for of which the major pterosaurs, Araripe The used for following specimens are compa- basin in Brazil is the most important one. rison: The aim of the work is to present present an of - Skull Anhanguera blittersdorffi, MN-4805-V of the toothedpterodactyloid taxon Brasi- update - Skull and mandible of referred specimen of leodactylus on the basis of the skull, since long the Anhanguera blittersdorffi, n. 40 Pz-DBAV-UERJ most distinctive element in the pterosaur - Skull Anhanguera santanae, AMNH 22555 skeleton. Consequently, only comparable taxa - Mandible 22573 Anhanguera sp, AMNH from Brazil will be discussed of which cranial - Mandible SAO 200602 Anhanguera sp, Arthurdactylusmaterial is known and therefore - Anhanguerid skull in the Iwaki Museum, Araripedactylus& Martill, 1994, conandoylei Frey IMNH 1053 castilhoi Price, 1971, Araripedactylus dehmi Brasileodactylus- Skull and mandible cf. arari- Wellnhofer, 1977, Santanadactylus brasilensis De pensis, MN-4797-V 1980, Santanadactylus pricei Wellnhofer, Buisonje, - BrasileodactylusAnterior part mandible araripen- 1985 and 1985) Santanadactylus spixi (Wellnhofer, sis, MN-4804-V are left out. - Anterior mandible Brasileodactyluspart MN-6517-V Dentition is sometimes used for systematic ? araripensis, - Skull and mandible Unwin used the Brasileodactylus sp., AMNH purposes (especially (2001) 24444 relative size, but the material he used is extremely - Partial skull BSP19911 27 fragmentated and no reliable picture of the entire Brasileodactylus sp, - Anterior skull dentition can be obtained), although the part Coloborhynchus clavirostris, BMNH 1822 45 Brasileodactylus (Pterosauria, Pterodactyloidea, Anhangueridae); an update = Staatliches Museum fur - MandibleColoborhynchus robustus, BSP1987147 Karlsruhe; SMNS - Skull and mandible Coloborhynchus piscator, Naturkunde, Stuttgart. NSM-PV19892 All measurementsof alveoli and diastemae have skull and mandible - Anterior parts Coloborhyn- been taken laterally except for the anterior and chus robustus, SMNK 2302 PAL anterolateral alveoli. These are measured - Skull Coloborhynchus sp, MN-4735-V lateral-lateral and anterolateral-ventrolateral SAO 16494 - Skull Coloborhynchus sp, respectively. The patterns are visualised in a - Skull and mandible ? Coloborhynchus sp., graph (graph 1 & 2). MN-6687-V ? MN-6503-V - Skull Coloborhynchus sp, - Mandible ? Coloborhynchus sp., MN-6687-V and The known - Skull mandible Coloborhynchus spielbergi, specimens RGM 401880 Type specimen ofBrasileodactylus andholotype of - Skull and mandible Criorhynchus mesembrinus, Brasileodactylus araripensis (Fig. 1). BSP1987146 Order Pterosauria Kaup, 1834 - Mandible cf. Criorhynchus mesembrinus, SMNS Suborder 1901 56994 Pterodactyloidea Plieninger, & Kellner, 1985 Undescribed skeleton of crestless Family Anhangueridae Campos - a pterosaur in GenusBrasileodactylus Kellner, 1984 the Kitakyushu museum and arari- Institutional abbreviations: Type species specimen: Brasileodactylus pensis, anterior part of mandible, MN 4804-V, BMNH = British Museum of Natural History, Museu Nacional, Rio de Janeiro, Brazil. London; BSP = Bayerische Staatsammlung fur Brasileodactylus as diagnosed by Palaontologie und historische Geologie, Munich; Diagnosis: Kellner (1984:580): "Pterosaurier mitUnterkiefer IMNH = Iwaki Museumof Coal Mining & Fossils, aus einer am Ende = gebildet langlichen abgerun- Iwaki; MN Museu Nacional, Rio de Janeiro; detenSymphyse, leicht nachoben gebogen, trian- NSM = National Science Museum, Tokyo; gularem Querschnitt, Schmalerung ab dem Pz-DBAV-UERJ = Geological Museum Univer- Teil, wobei ene an = proximalen Verbreiterung sity of Rio de Janeiro; RGM Nationaal Natuur- dem distalen Bereich ab der dritten Alveole exis- historisch Museum (Naturalis), former tiert, die eine flache Oberflache bildet. Vorhan- Rijksmuseum voor Geologie en Mineralogie, denseineiner medialenFurche an derDorsalseite Leiden; SAO = Sammlung Oberli, St. Gallen; der sehr ab dem SMNK = Staatliches Museum fur Naturkunde, Symphyse, ausgepragt Beginn Fig 1 Type specimen ofBrasileodactylus and holotype of Brasileodactylus araripensis, MN 4804-V. Courtesy of Museu Nacional, Rio de Janeiro, Brasil. Photograph by E. Endenburg/A.J. Veldmeijer MN 4804-V. Met dank Natio- Type specimen van Brasileodactylus en holotype vanBrasileodactylus araripensis, aan naal Museum, Rio de Janeiro, Brazilië. Foto door E. Endenburg / A.J. Veldmeijer 46 CRANIUM 22,1 (2005) Fig 2 The Crato specimen, MN 4797-V. Courtesy of Museu Nacional, Rio de Janeiro, Brasil. Photograph by E. En- denburg / A.J. Veldmeijer Het Crato stuk, MN 4797-V. Met dank aan het NationaalMuseum, Rio de Janeiro, Brazilië. Foto door E. Enden- burg / A.J. Veldmeijer in des Unterkiefers (distaler Teil), die sich proxi- ting on the rostral tip and widening caudally." maler Richtung verbreitet. Alveolen mit elipti- have to be regarded as apomorphies of Brasi- scher und rundlicher Form, Zahnabstande leodactylus.” They regard the degree of expansion ibidem Kellner vergrossern sich in proximaler Richtung. as apomorph ( : 103). (1984) regards Bezahnung bis an die Unterkieferspitze, Zahne Brasileodactylus as Ornithocheirid. Therostral end schmal und spitz, nach vorne stehend." (For starts to expand between the third and fourth English translation, see Kellner & Tomida (2000: alveoli, which is between the fourth and fifth 102). alveoli in Anhanguera and Coloborhynchus. However, the expansion in SMNS 55414 starts Emended diagnosis: Combination of first pair of betweenthe fourth and fifth alveolus as well. The the the alveoli positioned at anterior aspect; expansion inBrasileodactylus is small but distinct, second pair of alveoli positioned anterolateral similar to the in very situation Anhanguera, and is and the third pair of alveoli lateral. The dentary not consistent with the robust expansion in Colob- has small sagittal groove exten- orhynchus. Thefirst threepairs of alveoli are at the ding sub-grooves. anterior, anterolateral and lateral aspect respecti- Discussion of diagnosis: Kellner & Tomida (2000: vely (these are positioned anterodorsally and 103) evaluated Brasileodactylus and came to the laterodorsally in Anhanguera and Coloborhynchus; conclusion that "4) rostral end expand from the both second and third pairs are orientated antero- 3rd alveoli, forming a flat surface. 5) medial dorsally. The remaining alveoli are placed the dorsal of the relative for groove on part symphysis, star- strongly laterodorsally, to, instance, 47 Brasileodactylus (Pterosauria, Pterodactyloidea, Anhangueridae); an update Fig 3 The New York specimen, AMNH 24444. Courtesy of AMNH, New York. Photograph by E. Endenburg/A.J Veldmeijer Het stuk uit New York, AMNH 24444. Met dankaan AMNH, New York. Foto door E. Endenburg/A.J Veldmeijer comparable alveoli in Coloborhynchus as prelimi- The third alveolus is the largest of the alveoli in in nary results of study progress suggests. found the distal expansion; the following alveoli decrease in size continuously but the size In general, thealveoli are small. The first tooth is increases again steadily from the sixth alveolus the smallest, after which the tooth size increases. 48 CRANIUM 22,1 (2005) onwards. This results in the ninth alveolus being The Crato specimen (MN 4797-V; Fig. 2) as large as the third one (remarkable is Kellner's Various fossils have been assigned to Brasileodac- statement on the teeth, as they are not preserved
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