Kellner et al., 2009 by Felipe Elias

ISSN 1175-5326 (print edition)

Article

ZOOTAXA ISSN 1175-5334 (online edition)

A new crocodyliform from the Alcântara Formation (Cenomanian), Cajual Island, Brazil ALEXANDER W. A. KELLNER1, ANDRÉ E. P. PINHEIRO2, SERGIO A. K. AZEVEDO3, DEISE D. R. HENRIQUES4, LUCIANA BARBOSA DE CARVALHO5 & GUSTAVO R. OLIVEIRA6 Universidade Federal do Rio de Janeiro, Museu Nacional, Departamento de Geologia e Paleontologia, Setor de Paleovertebrados, Quinta da Boa Vista, Rio de Janeiro – RJ. Brazil. E-mails: 1kellner@mn.ufrj.br; 2paleolones@yahoo.com.br; 3sazevedo@mn.ufrj.br; 4deiseh@acd.ufrj.br; 5lucbc@acd.ufrj.br; 6 gustavoliveira@gmail.com

Abstract A new mesoeucrocodylian (Crocodyliformes) is described from the Laje do Coringa site, earliest Late Cretaceous (early Cenomanian) of the São Luís Basin, northeastern Brazil. Due to the likely hetorodonty indicated by distinct alveoli shapes, Coringasuchus anisodontis gen. et sp. nov. is tentatively referred to the Notosuchia and distinguished from other members of this clade by the presence of obliquely implanted teeth with the main axis directed anterolingually-toposterolabially and the presence of alveoli that are distinctively raised above the level of the dorsal margin of the dentary. The material further confirms the interpretation that the fossil concentration of the Laje do Coringa site is the result of multiple reworking events from previous deposits, but the degree of time-averaging was possibly higher than previously suspected. Key words: Coringasuchus anisodontis, Mesoeucrocodylia, early Cenomanian, Maranhão, Brazil, Alcântara Formation, Laje do Coringa

Introduction The site known as Laje do Coringa, situated on the coast of Cajual Island in Maranhão State, is one of the few bone-beds known from Brazil (Fig. 1). This deposit is regarded of early Cenomanian age and belongs to the Alcântara Formation, Itapecuru Group, of the São Luís Basin (Corrêa-Martins 1997; Medeiros & Schultz 2001), which was formed during the break up of Gondwana when the South American and the African continents drifted apart (Aranha et al. 1990). Explored since 1994, the Laje do Coringa site has yielded hundreds of isolated elements, preserved in a coarse conglomerate intercalated with sandstones. This region of Brazil is strongly influenced by the tidal regime that exposes but at the same time significantly damages the fossils. Subjected to strong erosion processes, part of this deposit had already been destroyed prior to discovery and covered by sand transported from the shore. Elements are found disarticulated and clearly represent end products of transportation (Medeiros et al. 2007). Fossils are invariably badly fragmented and in most cases heavily abraded (Fig. 2). Among the specimens collected are logs of conifers and pteridophytes (Mussa et al. 2000), fish remains including Lepidotes-like scales and isolated elements of the coelacanth Mawsonia Woodward (e.g., Medeiros et al. 2007), pterosaur teeth (Elias et al. 2007), putative mosasaur and plesiosaur teeth (Vilas-Bôas & Carvalho 2001), and a diverse dinosaurian fauna (e.g., Medeiros & Schultz 2001) that differs from that of other deposits in the Brazilian Cretaceous (Kellner & Campos 1999, 2000; Medeiros & Schultz 2002). Although the report of marine reptiles such as plesiosaurs should be regarded with caution, the nature of the

Accepted by R. Butler: 3 Feb. 2009; published: 9 Mar. 2009

fossil content of the Laje do Coringa site suggests that this accumulation resulted from the reworking of previous and distinct deposits. Previously, the crocodyliform record at the Laje do Coringa site has been limited to osteoderms and rare teeth, the latter having been only briefly mentioned or figured without further details (Medeiros & Schultz 2001; Nobre et al. 2002; Medeiros et al. 2007). Here, we report a lower jaw, the first to be recovered from this site, that is assigned to a new mesoeucrocodylian taxon, Coringasuchus anisodontis gen. et sp. nov. The specimen was found as surface float and had been encrusted by recent bivalves which were removed mechanically. Coringasuchus anisodontis is the first new species described at the Laje do Coringa site.

Systematic paleontology Crocodylomorpha Walker, 1970 Crocodyliformes Hay, 1930 (sensu Benton & Clark, 1988) Mesoeucrocodylia Whetstone & Whybrow, 1983 ?Notosuchia Gasparini, 1971 Coringasuchus gen. nov. Type species: Coringasuchus anisodontis gen. sp. nov.

Etymology: The generic name is derived from the name of the site (Laje do Coringa) where the specimen was found and souchos, the Greek word for crocodile. Diagnosis: As for the type and only species.

Coringasuchus anisodontis sp. nov. Holotype: MN 7128-V (Figs. 3-5), partial right lower jaw (dentary) housed at the Department of Geology and Paleontology of the Museu Nacional/Universidade Federal do Rio de Janeiro. Etymology: The specific name is derived from the Greek anisos (unequal, irregular) and odontos (tooth), in allusion to the odd variation of alveolar morphology that indicates a strongly heterodont dentition in the new taxon. Locality: Laje do Coringa, on the most eastern beach of Cajual Island, Maranhão State, Northeast Brazil, about 25 km west of São Luís. The specimen was found as surface float (UTM coordinates 2070559882 / 9726080). Horizon and age: Alcântara Formation, Itapecuru Group, São Luís Basin, earliest Late Cretaceous (early Cenomanian). Diagnosis: Mesoeucrocodylian with the following combination of characters that distinguish it from other members of this group: strong size variation of dentary teeth; comparatively small anterior rounded alveoli followed by larger elliptical alveoli; main axis of elliptical alveoli directed anterolingually-toposterolabially; some, but not all, alveoli distinctively raised above the level of the dorsal margin of the dentary. Description. The holotype of Coringasuchus anisodontis consists of an incomplete right mandibular ramus. Based on general morphology the preserved part appears to be from close to the posterior end of the mandibular symphysis (Figs. 3–5). The preserved part of the alveolar margin is 29.6 mm long, the maximum preserved height is approximately 23 mm, and the maximum transverse width (at the anterior end) is

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approximately 10.5 mm. The specimen has a dark brownish color, which is typical of fossils from the Laje do Coringa site. Despite having the margins and teeth broken, there are no signs of deformation that could have altered the shapes of the alveoli. Some parts of the element have been abraded and the external bone surface is slightly damaged, suggesting that this specimen was freed from the matrix and laid exposed as surface float for some time prior to collection, an interpretation supported by the presence of encrusting bivalves (removed from the specimen). Based on the nature of the broken edges, MN 7128-V was likely more complete before being exposed and broken by wave action.

FIGURE 1. Location of Cajual Island and the Laje do Coringa site (modified from Medeiros & Schultz 2001).

The dentary is the sole bone preserved. The lateral surface (Fig. 3A–B) is smooth and lacks strong ornamentation. Some occlusal scars, made by the contact of the upper dentition with the lower jaw, are present. At the posterior preserved end the lateral surface is flat or slightly concave and becomes gradually more convex towards the anterior end. Neurovascular foramina are not visible on the lateral surface, which is possibly a taphonomic artefact. The medial surface (Fig. 3C–D) is gently convex and shows scars (weakly preserved) and fractured areas. The splenial is not preserved and the position of the dentary-splenial suture could not be identified with certainty. Several small neurovascular foramina are found scattered on the medial surface (Fig. 3D). The occlusal view (Figs. 4–5) provides the most important anatomical information. No complete teeth are preserved, but the position of the alveoli demonstrates that the teeth were located close to the medial margin. At the preserved anterior end, the lateral border is quite broad and slightly concave, indicating that some of the teeth were slightly displaced medially. Five alveoli (four of which contain part of the tooth root in place) are preserved and the remnant of a possible sixth alveolus is also present (Fig. 3). The specimen is broken at the first preserved alveolus, exposing a thin layer of the dentine of the root. The second alveolus is sub-circular (diameter of approximately 4.3 mm) and is raised above the level of the dorsal margin of the dentary. The third alveolus (Fig. 5) is also subcircular (diameter of approximately 3 mm) but much smaller than the more anterior ones. In

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alveoli two and three a cross-section through the tooth exposes the enamel, dentine and the pulp cavity. The fourth alveolus (Fig. 5), the only one that lacks part of the root, is larger than the preceding ones, and has an elliptical shape with the long axis directed lingually at approximately 60 degrees to the long axis of the dentary. The fifth alveolus is also elliptical and raised above the level of the dorsal margin of the dentary. It is the largest of all those preserved and possesses the same general inclination of its long axis as alveolus four but at a much lower angle (approximately 10 degrees to the long axis of the dentary). The medial portion of the fifth alveolus is broken exposing part of the enamel and dentine of the tooth. The root of this tooth is quite long relative to the height of the lower jaw, a common feature in crocodylomorphs. A small concavity at the posterior preserved end of the jaw indicates the presence of another alveolus.

FIGURE 2. Partial view of the Laje do Coringa site, showing the isolated and broken nature of the fossil remains.

Discussion Based on the available material of Coringasuchus anisodontis, this species is not a primitive member of the Crocodylomorpha or Crocodyliformes, which are mainly found in Triassic and Jurassic deposits (e.g., Steel 1973; Clark 1994). Non-mesoeucrocodylian crocodyliforms such as Zosuchus and Edentosuchus are also found in Cretaceous deposits (Young 1973; Pol & Norell 2004; Pol et al. 2004), but although some possess heterodonty, they lack the remarkable size variation present in Coringasuchus anisodontis and lack elliptical alveoli. The clade Mesoeucrocodylia contains a relatively diverse array of crocodyliforms, including extinct longirostrine forms such dyrosaurids and pholidosaurids, some short-faced taxa traditionally referred or closely related to the Notosuchia, and the Eusuchia (e.g., Clark 1994; Pol et al. 2004). Longirostrine forms tend to have isometric teeth, or at least lack the strong variation in size and shape of closely located alveoli (e.g., Steel 1973; Buffetaut & Taquet 1977; Jouve 2007; Barbosa et al. 2008) which is present in Coringasuchus anisodontis. For the same reasons, Coringasuchus anisodontis can be excluded from the Eusuchia (e.g., Brochu 2001, 2003; Broin 2002) and closely related taxa (e.g., Schwarz 2002; Salisbury et al. 2006). Some eusuchians show expanded alveolar rims (e.g., crocodylids) but not the variation presented in Coringasuchus, where the second and fifth preserved alveoli are distinctively raised above the level of the dorsal margin of the dentary (Fig. 3).

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FIGURE 3. Photographs and drawings of the lower jaw of Coringasuchus anisodontis gen. et. sp. nov. (MN 7128-V) in lateral (A, B) and medial (C, D) views. Scale bars = 10 mm. Anatomical abbreviations: a1-a5, alveoli; a6 ?, alveolus ?; d, dentary; fa, fractured area. Neurovascular foramina are indicated in the circles.

According to Harris et al. (2000), heterodonty, and possibly a deviation from strict carnivory, occurred multiple times within Crocodylomorpha (e.g., Phyllodontosuchus lufengensis; Edentosuchus tienshanensis). However, the group that shows extreme variation in teeth morphology is Notosuchia, a Cretaceous group (mainly from Gondwana) of small–medium mesoeucrocodylians erected by Gasparini (1971). There is ongoing debate regarding the taxonomic content of this clade (e.g., Sereno et al. 2001; Pol 2003; Andrade & Bertini 2008a; Fiorelli & Calvo 2008), but authors agree that a number of comparatively odd, small terrestrial taxa such as Notosuchus, Sphagesaurus, Comahuesuchus, Malawisuchus and Candidodon form a monophyletic group for which the name Notosuchia is available. Some authors additionally consider the large bodied Baurusuchidae referable to this clade, which differ from all other taxa (and also Coringasuchus anisodontis) by the possession of a theropod-like dentition (Price 1945; Riff & Kellner 2001). The variation in shape and size of the alveoli suggest that Coringasuchus anisodontis had teeth of different sizes and shapes, features observed in many taxa with heterodontous dentition. A comparable example is the notosuchian Sphagesaurus, where the obliquely placed teeth are positioned in elliptical alveoli and the more rounded teeth are positioned in circular to subcircular alveoli (e.g., Pol 2003). As a result we tentatively refer the new species to Notosuchia. Several notosuchids exhibit unusual adaptations, primarily involving the dentition, which are related to feeding behavior (Price 1950; Bonaparte 1991, 1996; Pol 2003, Vasconcellos & Carvalho 2005; Andrade & Bertini 2008a, c). The elliptical alveoli (indicating the presence of obliquely oriented teeth) present in Coringasuchus anisodontis differ from notosuchians such as

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Araripesuchus, Mariliasuchus, Candidodon, Comahuesuchus, Anatosuchus, and Malawisuchus, where the long axes of the teeth tend to be parallel to the alveolar border (Price 1959; Carvalho & Campos 1988; Carvalho & Bertini 1999; Sereno et al. 2003; Zaher et al. 2006).

FIGURE 4. Photographs and drawings of the lower jaw of Coringasuchus anisodontis gen. et. sp. nov. (MN 7128-V) in occlusal view (A, B). Scale bars = 10 mm. Anatomical abbreviations: a1-a5, alveoli; a6?, alveolus ?; e, enamel; lab, labial dentary process; os, occlusal scar.

Obliquely inclined dental implantation is known primarily in Sphagesaurus, where the long axes of the alveoli are directed anterolabially-to-posterolingually (Pol 2003; Andrade & Bertini 2008a). This is also the case for Adamantinasuchus and to a lesser extend in Notosuchus and Mariliasuchus (Nobre & Carvalho 2006; Andrade & Bertini 2008 b, c). Coringasuchus anisodontis differs from all those taxa by having the long axes of the alveoli inclined to the opposite direction (e.g., anterolingually-to-posterolabially). In addition to this character Coringasuchus anisodontis differs from all other notosuchians by having some alveoli distinctively raised above the level of the dorsal margin of the dentary. With the exception of the specimen described here (MN 7128-V), the only records of crocodylomorphs from the Laje do Coringa site are some undescribed osteoderms and isolated teeth. Two of the teeth have been figured and both are comparatively large (tooth crown more than 50 mm in apicobasal height) with rather smooth enamel (Medeiros & Schultz 2001; Medeiros et al. 2007). These teeth differ markedly in size from those of Coringasuchus anisodontis and other comparatively small notosuchian taxa known from Brazilian Cretaceous deposits (e.g. Price 1959; Carvalho 1994; Carvalho & Bertini 1999; Nobre & Carvalho 2006). While isolated reptilian teeth are often quite difficult to assign to any specific taxon (e.g., Kellner & Mader 1997), the two isolated teeth from the Laje do Coringa site lack the serrated carinae commonly present in

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baurusuchids (Price 1945; Riff & Kellner 2001) and peirosaurids (Gasparini 1982; Carvalho et al. 2004, 2007), and the strong striations present in dyrosaurids (e.g., Barbosa et al. 2008). Based on their morphology, those teeth do not appear to belong to “trematochampsids” recorded in Cretaceous deposits of the Araripe Basin, northeastern Brazil (Kellner 1987), and are overall more similar to pholidosaurids found in both South America and northern Africa (Broin & Taquet 1966; Buffetaut & Taquet 1977; Sereno et al. 2001).

FIGURE 5. Photograph of the lower jaw of Coringasuchus anisodontis gen. et. sp. nov. (MN 7128-V) showing occlusal view of the third and fourth preserved alveoli. Scale bar = 5 mm.

Final considerations The fossil specimens recovered from the Laje do Coringa site are quite diverse and represent taxa not normally found associated (e.g., plant material, marine fishes, small terrestrial vertebrates, large dinosaur bones). Although taxa with distinct preservation potential that originally constituted a single ecosystem can occasionally be preserved in the same fossil deposit (e.g., Calvo et al. 2007) this is a rare situation in the geological record. Based on the taxic diversity and the isolated nature of the material, Medeiros & Schultz (2001) interpreted the site as a result of multiple reworking events from previous deposits. The fragile nature of the lower jaw of Coringasuchus anisodontis agrees with this interpretation and it is probably reworked from a different deposit to that of most of the large isolated dinosaur remains and tree logs recovered thus far. Medeiros & Schultz (2001) also concluded that the extent of time averaging at the Laje do Coringa site was not significant. However, this contradicts previous studies where intense reworking, common in proximal marine systems such as the one observed here, is normally linked with a high degree of time averaging (e.g., Behrensmeyer 1982). Experimental studies have indicated that reptilian teeth can undergo extensive transport but exhibit little or no wear (Martin 1999), as is the case for some teeth found at this site, suggesting that detailed taphonomic studies are still needed to provide a better understanding of the origin of this fossil deposit. Since the discovery of the Laje do Coringa site, extensive field work has been continuously carried out in this region, demonstrating that small reptilian bones are exceedingly rare. To our knowledge, the holotype of

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Coringasuchus anisodontis is the first small reptilian lower jaw to be recovered from this site, which is consistent with the notion that remains of crocodylomorphs are rare (Medeiros et al. 2007). Despite its fragmentary nature, MN 7128-V shows distinct features allowing the recognition of a new crocodylomorph taxon, shedding new light on morphological variation within notosuchids. Coringasuchus anisodontis is the first species to be named from the Laje do Coringa site. It is possible that the introduction of screen washing techniques, although difficult in this region due to the effects of the tides (Corrêa-Martins 1997), might potentially recover additional specimens that would complement the anatomical information for this taxon.

Acknowledgments We would like to thank Jorge Calvo (Universidad del Comahue, Neuquén), Diogenes de Almeida Campos (Museu de Ciências da Terra/DNPM, Rio de Janeiro), Mark Norell (American Museum of Natural History, New York), and Ismar S. Carvalho, Felipe Vasconcellos and Thiago Marinho (Insituto de Geociêcias/UFRJ, Rio de Janeiro) for access to specimens. We also would like to thank Christopher Brochu (University of Iowa), Marco Brandalise (University of Bristol), and Richard Butler (The Natural History Museum, London) for many suggestions that significantly improved the manuscript. The Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq, grants numbers 486313/2006-9 and 501267/2008-5 to AWAK and grant number140812/2007-5 to GRO) and Fundação Carlos Chagas Filho de Amparo a Pesquisa do Estado do Rio de Janeiro (FAPERJ, grant number E-26/152.885/2006 to AWAK) partially funded this project. The drawings used in this paper were made by AEPP.

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