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The Cretaceous Crocodyliform Caipirasuchus: Behavioral Feeding Mechanisms

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The Cretaceous Crocodyliform Caipirasuchus: Behavioral Feeding Mechanisms Cretaceous Research 84 (2018) 181e187 Contents lists available at ScienceDirect Cretaceous Research journal homepage: www.elsevier.com/locate/CretRes Short communication The Cretaceous crocodyliform Caipirasuchus: behavioral feeding mechanisms * Fabiano Vidoi Iori a, b, , Ismar de Souza Carvalho c a Museu de Paleontologia “Prof. Antonio Celso de Arruda Campos”, Centro de Artes, Praça do Centenario, 15910-000, Monte Alto, SP, Brazil b Museu de Paleontologia “Pedro Candolo”, Estaçao~ Cultura, Praça Farmac^eutico Bruno Garisto, 15890-000, Uchoa, SP, Brazil c Universidade Federal do Rio de Janeiro, Instituto de Geoci^encias, Departamento de Geologia, Cidade Universitaria, Ilha do Fundao,~ Av. Athos da Silveira Ramos, 274, 21941-916, Rio de Janeiro, RJ, Brazil article info abstract Article history: Caipirasuchus is a sphagesaurid crocodyliform from the Cretaceous of the Bauru Basin. Three species, Received 6 June 2017 with terrestrial habits, leaned bodies and measuring less than 1.5 m in length, have been described up to Received in revised form now: Caipirasuchus montealtensis, C. paulistanus and C. stenognathus. Similar to other advanced noto- 12 November 2017 suchians, they present a complex mastication mechanism that includes propalinal movements, side Accepted in revised form 24 November 2017 alternation during chewing and tooth-to-tooth occlusion. This study analyzes the anatomic structures Available online 2 December 2017 and dental wear related to such mastication mechanisms in Caipirasuchus, in corresponding to feeding strategies. Keywords: © Caipirasuchus 2017 Elsevier Ltd. All rights reserved. Sphagesauridae Mastication processes 1. Introduction in the southeastern portion of the South American Plate (Fernandes and Coimbra, 1996) between the Coniacian and Maastrichtian Notosuchia was one of the most peculiar groups during the (Fernandes and Coimbra, 2000). Dias-Brito et al. (2001) suggested Cretaceous of Gondwana. These crocodyliforms were essentially two sedimentary episodes: one in the Turonian-Santonian interval terrestrial and had very specialized dentition, indicating that had (represented by the Adamantina, Araçatuba and Sao~ Jose do Rio occupied various ecological niches (Carvalho and Bertini, 1999). Preto formations) and the other in the Maastrichtian (the Marília Among these crocodyliforms is Sphagesauridae, mainly character- Formation). Menegazzo et al. (2016) suggested that the history of ized by presenting a set of teeth adapted to herbivory/omnivory the basin spans from the Cenomanian to the early Paleocene. and a complex mastication mechanism (Pol, 2003; Marinho and Caipirasuchus was a genus widely distributed in the Bauru Basin, Carvalho, 2009; Novas et al., 2009; Iori and Carvalho, 2011; Iori and the three know species, Caipirasuchus montealtensis, et al., 2011; Kellner et al., 2011a). The diagnoses of this clade C. paulistanus and C. stenognathus, have been recovered from were initially based on two isolated posterior teeth, that differ sandstones of the Adamantina Formation (Upper Cretaceous), strongly from the dentition of other reptiles (Price, 1950; Kuhn, deposited in different environmental contexts. In the deposits of 1968). the Monte Alto region, the crocodyliform specimens are commonly In the Bauru Basin, Sao~ Paulo state, Brazil, five genera attributed found in paleosols, whereas the Catanduva specimens come from to sphagesaurids were discovered: Sphagesaurus, Ada- flooded plains (Iori et al., 2013). mantinasuchus, Armadillosuchus, Caipirasuchus and Caryonosuchus We have analyzed here the morphological features and tooth (Pol, 2003; Nobre and Carvalho, 2006; Andrade and Bertini, 2008; wear observed in Caipirasuchus. The new observation has contrib- Marinho and Carvalho, 2009; Iori and Carvalho, 2011; Kellner uted to enhance our knowledge on the mastication mechanism in et al., 2011a; Iori et al., 2013; Pol et al., 2014). This basin (Fig. 1)is Caipirasuchus and enabled a greater understanding of its a broad cratonic depression developed during the Late Cretaceous paleoautoecology. * Corresponding author. E-mail addresses: [email protected] (F.V. Iori), [email protected] (I.S. Carvalho). https://doi.org/10.1016/j.cretres.2017.11.023 0195-6671/© 2017 Elsevier Ltd. All rights reserved. 182 F.V. Iori, I.S. Carvalho / Cretaceous Research 84 (2018) 181e187 Fig. 1. Caipirasuchus crocodyliforms analyzed from the Bauru Basin and their geographical distribution. 2. Materials the quadrate; pm, premaxillary tooth; pt, pterygoid; q, quadrate; r, root; rap, retroarticular process; rlf, retroarticular lateral flange; 2.1. Specimens of Caipirasuchus rdg, ridges; sa, surangular; sp, splenial; sph, sphagesauriform tooth. Cranial elements related to the mastication mechanisms of four specimens of Caipirasuchus (Fig. 1), all coming from the sandstones 3. Morphological features and mastication of the Adamantina Formation (Bauru Group, Bauru Basin), were here studied (see Fig. 1). Specimen MPMA 67-0001/00 is the ho- The description of a nearly full skull and the anterior part of the lotype of Caipirasuchus paulistanus and was found in Monte Alto. mandible of a specimen of Sphagesaurus huenei by Pol (2003) The specimens of C. montealtensis are MPMA 68-0003/12 (an adult) allowed the inference of relevant aspects of mastication in the and MPMA 68-0004/12 (a youth) were from Catanduva. The ho- species, such as tooth-by-tooth occlusion, propalinal movements lotype of C. stenognathus (MZSP-PV 139) is from General Salgado. and side alternation in mandible movements, as well as mecha- nisms that so far had not been noticed in crocodyliforms. The 2.2. Institutional abbreviations movements involved in mastication, inferred in Sphagesaurus huenei, have been found in other members of the family, such as MPMA, Museu de Paleontologia “Prof. Antonio Celso de Arruda Armadillosuchus, Caryonosuchus and Caipirasuchus (Marinho and Campos”, Monte Alto, SP, Brazil; MZSP-PV, Museu de Zoologia, Carvalho, 2009; Iori and Carvalho, 2011; Kellner et al., 2011a; Pol Universidade de Sao~ Paulo, Sao~ Paulo, Brazil. et al., 2014). Iori and Carvalho (2011) simulated the bite of Caipirasuchus 2.3. Anatomical abbreviations paulistanus in plasticine and found two areas of action: an anterior area of apprehension, and another for effective food processing in amf, anterior surface of retroarticular medial flange; an, the posterior area. angular; art, articular; aw, apical wear; c, crown; d, dentary tooth; In a detailed study on the mastication mechanisms of hetero- € dent, dentary; gc, gleinoid crest; lcq, lateral condyle of the quad- dontiform crocodylomorphs, Osi (2013) corroborates previous rate; lw, lingual wear; m, maxillary tooth; mcq, medial condyle of studies on the mandibular movements of sphagesaurids and F.V. Iori, I.S. Carvalho / Cretaceous Research 84 (2018) 181e187 183 observes that, in this clade, propalinal movement is greater than in C. paulistanus,butinC. montealtensis (MPMA 68-0003/12) and any other crocodyliforms. C. stenognathus is almost intact (Figs. 2 and 3). Both lack the The osteological analysis of the quatrate-articular joint, the glenoid fossa typical of crocodyliforms (with a lateral-medial dentition arrangement, diastemata and the relationships between orientation and a crest separating it in two parts). The glenoid lower and upper dentition of Caipirasuchus allowed the mastication crest, an antero-posteriorly elongate intumescence, is bordered process reconstruction and the inference of the possible eating on its sides by a shallow fossa and medially by a small bone habits of Caipirasuchus. expansion. The glenoid crest fits between the quadrate condyles and is about twice as long as the quadrate region that is in contact with it (Figs. 2 and 3). 3.1. Quadrate-articular joint When in occlusion on the anteriormost position of the mandible € (Fig. 3A), the quadrate articulates with the posterior half of the Osi (2013) points out the unusual medial hemicondyle of the articular intumescence and the posterior margin of the quadrate quadrate bone observed in Caipirasuchus paulistanus (MPMA 67- condyles aligns to the beginning of the retroarticular process 0001/00), which is very conspicuous and has an edge of about 1 cm (Fig. 3A2 and A3). In this position, all mandibular teeth are exter- ventral to the lateral hemicondyle. However, he stressed that no nally bordered by the skull teeth. When the mandible is articulated sphagesaurid had so far presented a fully preserved quadrate- in its furthest posterior position (Fig. 3B), the apices of the five last articular contact. A specimen of Caipirasuchus montealtensis mandibular teeth are almost aligned to the anterolateral margin of (MPMA 68-0003/12) and the holotype of Caipirasuchus sten- the suborbital fenestra. ognathus preserve that region, and as in the C. paulistanus, the ventral surface extends further ventrally on the medial than on the 3.2. Dentition arrangement lateral hemicondyle (Fig. 2)(Iori et al., 2013; Pol et al., 2014). The articulars in the specimes of Caipirasuchus present similar The mandibular teeth in Caipirasuchus paulistanus and C. mon- aspects: they are wider than taller; they have a semi-circular tealtensis are arranged into two straight lines. The 1st through 6th medial contour; and the retroarticular processes are very small teeth are arranged in one line, parallel to the sagittal plane, another (Iori and Carvalho, 2011; Iori et al., 2013; Pol et al., 2014). The line begins at the 6th tooth and diverges laterally by about 40 from largest mediolateral
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