Mitt. Mus. Nat.kd. Berl., Geowiss. Reihe 5 (2002) 239-263 10.11.2002 On the systematic relationships of CearadactyZus atrox, an enigmatic Early Cretaceous pterosaur from the Santana Formation of Brazil David M. Unwinl With 6 figures Abstract Cearadactylus atrox, a large pterodactyloid pterosaur represented by an incomplete skull and lower jaw from the Lower Cretaceous Santana Formation of Brazil, is a valid species. Diagnostic characters include a mandibular symphysis with a transversely expanded ‘spatulate’ anterior end that is considerably wider than the rostral spatula, and a third rostral tooth that has a basal diameter more than three times that of the fifth tooth. Additional diagnostic characters, contingent upon assign- ment of Cearadactylus atrox to the Ctenochasmatidae, include: anterior ends of jaws divaricate and containing 7 pairs of rostral teeth and 6 pairs of mandibular teeth; marked dimorphodonty, with an abrupt change in tooth morphology; and a “high cheek”. ‘Ceurudactylus? ligubuei’ Dalla Vecchia, 1993, based on an incomplete skull, also from the Santana Formation, is not related to Cearadactylus utrox, exhibits several ornithocheirid synapomorphies and is referred, tentatively, to Anhun- gueru. Ceurudactylus atrox exhibits various synapomorphies of the Ctenochasmatidae (rostrum anterior to nasoantorbital fe- nestra greater than half total skull length, teeth in anterior part of dentition relatively elongate and pencil-shaped, premaxilla has at least 7 pairs of teeth), the defining synapomorphy of the Gnathosaurinae (rostrum with dorsoventrally compressed laterally expanded spatulate anterior expansion), and shares two synapomorphies with the Chinese gnathosaurine Huanhep- terus quingyangensis (anterior tips of jaws divaricate, teeth restricted to anterior half of mandible). Two elongate cervical vertebrae, also from the Santana Formation and previously assigned to ‘Suntunadactylus brusilensis’, are tentatively referred to Ceuruductylus. Reconstruction of the temporal history of the Ctenochasmatidae suggests that while ctenochasmatines became increasingly specialised for filter feeding, gnathosaurines changed from sieve feeding to piscivory, acquiring several cranial characters that are similar to those of ornithocheirids, a group that also includes large aerial piscivores that used a terminal tooth grab for prey capture. Key words: Pterosaur, Lower Cretaceous, Brazil, Cearaductylus, Ctenochasmatidae. Zusammenfassung Cearadactylus atrox aus der Santana-Formation (Unterkreide, NO-Brasilien) ist eine valide Art. Eine Revision des Taxons, von dem ein unvollstandiger Schadel mit Unterkiefer vorliegt, ergab folgende diagnostische (autapomorphe) Merkmale. Die Symphyse hat ein transversal verbreitertes spatelformiges Vorderende, das deutlich breiter ist als das Schnauzenende. Der dritte rostrale Zahn erreicht einen basalen Durchmesser, der jenen des fiinften Zahns um das Dreifache ubertrifft. Hinzu kommen Merkmale, die C. atrox mit der Ctenochasmatidae gemein hat, darunter die vorn auseinanderklaffenden Kieferran- der, sieben rostrale Zahnpaare, sechs Unterkieferzahnpaare, eine ausgepragte Dimorphodontie sowie eine hohe Wangenre- gion. ‘Cearadactylus ? Zigubuei‘ Dalla Vecchia 1993, ebenfalls mit einem unvollstandigen Schadel belegt, ist nicht naher mit C. atrox venvandt. Im Gegensatz zu letzterem zeigt ‘C. ? Zigubuei’ signifikante Ahnlichkeiten mit den Omithocheiridae. Unter Vorbehalt wird er hier der Gattung Anhanguera zugeordnet. C. atrox hat neben eindeutigen Synapomorphien der Ctenochas- matidae, z. B. erreicht das Rostrum anterior des nasoantorbitalen Fensters mehr als die halbe Schadellange, die vordersten Zahne sind verlangert und stiftformig und die das Pramaxillare tragt mindestens sieben Zahnpaare. Daneben besitzt C. atrox auch noch die entscheidende Synapomorphie der Gnathosaurinae, namlich ein Rostrum mit dorsoventral komprimierter vor- derem Auswuchs. AuBerdem ist C. atrox gekennzeichnet durch zwei Autapomorphien des Gnathosaurinen Huanhepterus quin- gyungensis aus China: divergierende Schnauzenenden und Zahne begrenzt auf vordere Kieferhalfte. SchlieBlich werden zwei lange Halswirbel, die auch aus der Santana Formation stammen und bislang zu Suntunadactylus brusiliensis gerechnet wurden, unter Vorbehalt zu Ceuruductylus gestellt. Die Evolutionsgeschichte der Ctenochasmatidae ist durch eine zunehmende Spezia- lisierung auf filternde Ernahrungsweise gekennzeichnet. Die Gnathosaurinen dagegen stellten sich von der filternden auf eine piscivore Ernahrung um, wobei sie eine Reihe von Schadelmerkmalen erworben haben, die den Ornithocheiriden konvergent ahnlich ist. Schliisselworter: Pterosaurier, Unter-Kreide, Brasilien, Ornithocheiridae, Ctenochasmatidae. Museum fiir Naturkunde der Humboldt-Universitat, Institut fur Palaontologie, Invalidenstr. 43, D-10115 Berlin, Germany. Received April, accepted July 2002 240 Unwin, D. M., Cearadactylus from the Santana Formation Introduction from other deposits. Two distinct approaches are discernable. Some workers (e.g. Price 1971, The Romualdo Member of the Lower Cretaceous Wellnhofer 1977, 1985, BuisonjC 1980, Kellner Santana Formation of Brazil has yielded a re- 1984, Campos & Kellner 1985, Leonardi & Bor- markably well preserved and diverse pterosaur as- gomanero 1985, Kellner 1989) have erected new semblage (e.g. Campos & Kellner 1985, Wellnho- genera and even families for Santana pterosaurs, fer 1985, 1987, 1991a, b, c, Unwin 1988, Kellner subsequently assigning similar forms from other 1990, Martill 1993, Kellner & Tomida 2000). Un- Cretaceous deposits to these taxa (e.g. Kellner & like most fossils of pterosaurs, skeletons are often Tomida 2000). Others (e.g. Fastnacht 2001, Un- preserved uncrushed (e.g. Wellnhofer 1988, win 2001) have argued that some species of San- 1991c), although they are rarely complete (Kell- tana pterosaurs belong within previously estab- ner & Tomida 2000), and some examples of soft lished genera such as Ornithocheirus Seeley, tissue preservation have also been reported 1869 and Coloborhynchus Owen, 1874, both (Campos et al. 1984, Martill & Unwin 1989, Mar- based on material from the late Lower Cretac- till et al. 1990, Kellner 1997). A surprising aspect eous of England (see Unwin 2001). The situation is the apparent diversity of this assemblage: 17 is even more confused at higher taxonomic le- species distributed among 9 genera representing vels with, for example, some authors continuing at least 3 families have been reported so far to use the name Anhangueridae Campos & Kell- (Kellner & Tomida 2000: table 9). This excludes ner, 1985, even though as I have noted else- taxa from the Crato Formation which has been where (Unwin 2001) the content and diagnosis included in the Santana Formation (e.g. Maisey of this family is practically identical to that of 1991), but is now recognised as a distinct unit the Ornithocheiridae Seeley, 1870, an older and (Martill & Wilby 1993), perhaps as much as thus senior synonym. 10 million years older, although with a rather simi- Revision of the taxonomy and relationships of lar pterosaur assemblage (Frey & Martill 1994, Santana pterosaurs is needed if the potential of Martill & Frey 1998,1999, Saygo & Kellner 2000). this important assemblage for understanding the Various workers (e.g. Wellnhofer 1987, 1991c, anatomy, functional morphology, ecology and Unwin 1988, 2001, Kellner & Tomida 2000) have evolutionary history of pterosaurs is to be fully suggested that the apparent species diversity of realised. Here, attention is focused on one of the the Santana Formation assemblage may have earliest and most unusual pterosaurs to be de- been inflated by excessive taxonomic splitting, scribed from the Santana Formation, Cearadacty- leading to a situation that closely resembles early lus, currently represented by two species: Ceara- stages in the development of the taxonomic un- dactylus atrox Leonardi and Borgomanero, 1985 derstanding of other pterosaur assemblages such and ‘Cearadactylus? ligabuei’ Dalla Vecchia, as the Solnhofen Limestone (Wellnhofer 1970, 1993, although the latter is only tentatively as- 1975a) and the Cambridge Greensand (Welln- signed to this genus. This study set out to answer hofer 1978, Unwin 2001). In the case of the San- four questions: (1) Is Cearadactylus atrox a dis- tana Formation, oversplitting can be partially at- tinct, diagnosable and valid taxon? (2) Can the tributed to the non-comparability of some relationship of ‘Cearadactylus? ligabuei’ to C. species because of incomplete preservation atrox be resolved? (3) What are the relationships (Wellnhofer 1987, 1991c, Kellner & Tomida of C. atrox to other pterosaurs? (4) Can any 2000). More problematic, however, has been the other fossil material from the Santana Formation assignment of decisive taxonomic value to ques- be assigned to C. atrox? tionable characters such as size, the degree of Two circumstances allow these questions to be fusion of skeletal elements and, most controver- answered. First, recent systematic revisions of sially, the morphology and position of cranial Santana pterosaurs (e.g. Fastnacht 2001) and re- crests. The latter are widely distributed among lated assemblages such as that from the Cam- Santana pterosaurs and have often been cited as bridge Greensand (Unwin 2001), together with key diagnostic features (e.g. Campos & Kellner work now underway on other undescribed speci- 1985, Kellner & Tomida 2000), even though their mens, are helping to resolve the taxonomy of variation and distribution are