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Home , Armadillosuchus, Baurusuchus, Caipirasuchus, Campinasuchus, Pissarrachampsa, Stratiotosuchus

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Cretaceous Research xxx (xxxx) xxx

55 Contents lists available at ScienceDirect 56 57 58 Research 59 60 journal homepage: www.elsevier.com/locate/CretRes 61 62 63 64 65 1 A new Pissarrachampsinae specimen from the Bauru Basin, , 66 2 67 3 adds data to the understanding of the 68 4 (, ) distribution in the of 69 5 70 6 Q7 South America 71 7 72 8 a, * b, c d Q6 Gustavo Darlim , Ismar de Souza Carvalho , Sandra Aparecida Simionato Tavares , 73 9 Max Cardoso Langer a 74 10 75 11 a Universidade de Sao~ Paulo, Faculdade de Filosofia, Ci^encias e Letras de Ribeirao~ Preto, Laboratorio de Paleontologia, Av. Bandeirantes, 3900, Ribeirao~ Preto, 76 12 SP, Brazil 77 b ^ 13 Q2 Universidade Federal do Rio de Janeiro, Instituto de Geociencias, CCMN, 21.910-200 Av. Athos da Silveira Ramos 273, Rio de Janeiro, RJ, Brazil c Centro de Geoci^encias da Universidade de Coimbra, Portugal 78 14 d Museu de Paleontologia “Prof. Antonio^ Celso de Arruda Campos”, Praça do Centenario, Centro, Monte Alto, SP, Brazil 79 15 80 16 81 17 article info abstract 82 18 83 19 Article history: Baurusuchidae is one of the most diverse notosuchian groups, represented by ten formally described 84 20 Received 7 January 2021 species from the Upper Cretaceous deposits of the Bauru and Neuquen basins, respectively in Brazil and 85 21 Received in revised form Argentina. Among these, recent phylogenetic analyses placed Wargosuchus australis, 86 18 July 2021 22 dinizi, and sera within Pissarrachampsinae, whereas Baurusuchinae is composed by 87 Accepted in revised form 18 July 2021 23 Aphaurosuchus escharafacies, Aplestosuchus sordidus, albertoi, Baurusuchus pachecoi, Baur- Available online xxx 88 24 usuchus salgadoensis, and maxhechti. All baurusuchines, except for Baurusuchus pachecoi 89 and Stratiotosuchus maxhechti, were collected from the same locality of the Adamantina/Vale do Rio do 25 Keywords: 90 Peixe Formation, i.e., Fazenda Buriti, near General Salgado, Sao~ Paulo. Recent works suggested that 26 Notosuchia 91 27 Baurusuchidae pissarrachampsines of that stratigraphic unit come only from outcrops in Minas Gerais, with a possibly  92 Campinasuchus dinizi older age equivalent to those of the Neuquen Basin members of the group. Here, we present a new 28 93 Distribution Pissarrachampsinae specimen from Fazenda Sao~ Jose, a site near Fazenda Buriti, which challenges the 29 Paleobiogeography proposed chronostratigraphic separation between the Sao~ Paulo and Minas Gerais baurusuchids, sug- 94 30 gesting that both groups lived at about the same time. 95 31 © 2021 Elsevier Ltd. All rights reserved. 96 32 97 33 98 34 99 35 100 36 101 37 1. Introduction species were described in the last decade (Nascimento & Zaher, 102 38 2010; Montefeltro et al., 2011; Carvalho et al., 2011; Marinho et al., 103 39 Baurusuchidae is a well-known group of terrestrial and predatory 2013; Godoy et al., 2014; Cotts et al., 2017; Darlim et al., 2021). The 104 40 notosuchians from the Late Cretaceous of Gondwana (Montefeltro group is restricted to the Bauru Basin, except for the Argentinean 105 fi 41 et al., 2020). It was rst described based on Baurusuchus pachecoi species rothi and Wargosuchus australis from the 106 42 Price, 1945, which remained as the single species for the until Bajo de la Carpa Formation, and possibly the Pakistani Pabweshi 107 43 the description of Baurusuchus salgadoensis by Carvalho et al. (2005). pakistanensis, currently comprising eleven species (Woodward and 108 fi 44 Since then, other baurusuchids were identi ed, such as Cyn- Woodward, 1896; Wilson et al., 2001; Martinelli and Pais, 2008; Q3 109 45 odontosuchus rothi (Gasparini 1972)andStratiotosuchus maxhechti Darlim et al., 2021). Baurusuchidae is traditionally diagnosed by a 110 46 (Campos et al., 2001). Yet, much of the morphological and phylo- reduced tooth formula and ziphodont dentition, a laterally com- 111 47 genetic information about Baurusuchidae is quite recent, as most pressed rostrum, and a verticalized quadrate (sensu Price, 1945). It 112 48 corresponds to one of the most diverse notosuchian , with two 113 49 subgroups recognized by Montefeltro et al. (2011),Baurusuchinae 114 50 * Corresponding author. and Pissarrachampsinae (Darlim et al., 2021). The former group in- 115 51 E-mail addresses: [email protected] (G. Darlim), [email protected] cludes Baurusuchus albertoi, Baurusuchus pachecoi, Baurusuchus 116 (I. de Souza Carvalho). 52 117 53 https://doi.org/10.1016/j.cretres.2021.104969 118 54 0195-6671/© 2021 Elsevier Ltd. All rights reserved. 119

Please cite this article as: G. Darlim, I. de Souza Carvalho, S.A.S. Tavares et al., A new Pissarrachampsinae specimen from the Bauru Basin, Brazil, adds data to the understanding of the Baurusuchidae (Mesoeucrocodylia, Notosuchia) distribution in the Late Cretaceous of South America, Cretaceous Research, https://doi.org/10.1016/j.cretres.2021.104969 YCRES104969_proof ■ 29 July 2021 ■ 2/13

G. Darlim, I. de Souza Carvalho, S.A.S. Tavares et al. Cretaceous Research xxx (xxxx) xxx

1 salgadoensis, Stratiotosuchus maxhechti, Aplestosuchus sordidus,and of left jugal, the left ectopterygoid, and the anterior portion of both 66 2 Aphaurosuchus escharafacies,andthelatterPissarrachampsa sera, hemimandibles (anterior to the external mandibular fenestra), 67 3 Campinasuchus dinizi,andWargosuchus australis (Campos et al., including dentary and splenial. Furthermore, the dentition of both 68 4 2001; Nascimento and Zaher, 2010; Montefeltro et al., 2011; upper and lower jaws is mostly preserved. Although incomplete, the 69 5 Carvalho et al., 2011; Godoy et al., 2014; Darlim et al., 2021). How- recovered material is in good state of preservation (Fig. 2). 70 6 ever, these subgroups have not been retrieved as monophyletic in The specimen was originally found with the right lateral surface 71 7 some recent phylogenetic analyses (Fiorelli et al., 2016; Barrios et al., of the premaxilla and exposed on the rock-matrix. The 72 8 2018; Leardi et al., 2018; Fernandez-Dumont et al., 2020). material was prepared by SAST at the Paleontology Museum “Prof. 73 9 All Brazilian baurusuchids come from the Vale do Rio do Peixe Antonio Celso de Arruda Campos” in Monte Alto-SP. The rock ma- 74 10 Formation of Fernandes & Coimbra (2000), which is part of the trix was removed using low impact hammers, and the fossil was 75 11 Adamantina Formation described by Soares et al. (1980; see also stabilized using white glue Cascorez in different concentrations. At 76 12 Batezelli, 2015). Several of these were also collected in the same the end of the preparation, the and mandible were 77 13 locality, known as Fazenda Buriti, near Prudencio^ e Morais district, disarticulated. 78 14 General Salgado, Sao~ Paulo, including Aplestosuchus sordidus, Baur- 79 15 usuchus alberoi, Baurusuchus salgadoensis,andGondwanasuchus 3.1. Comparative material 80 16 scabrosus, all of which, except for the latter, are baurusuchines 81 17 (Marinho et al., 2013; Godoy et al., 2014). Pissarrachampsines, on the Aphaurosuchus escharafacies LPRP 0697, Aplestosuchus sordidus 82 18 other hand, are restricted to the Adamantina/Vale do Rio do Peixe LPRP 0229a, arrudai UFRJ DG 303R, MPMA 64- 83  19 Formation of Minas Gerais and Bajo de la Carpa Formation, Neuquen 0001-04; Baurusuchus pachecoi DGM 299-R, Baurusuchus salga- 84 20 Basin, in Argentina. Based on the age of the latter unit, Montefeltro doensis MPMA 62-001-02; UFRJ DJ 288-R, 308-R; Campinasuchus 85 21 et al. (2011) discussed a possible older age (i.e., ) for the dinizi CPPLIP 1235, 1234, 1236, 1237, 1436; Gondwanasuchus scab- 86 22 Pissarrachampsinae-bearing deposits of Minas Gerais, compared to rosus UFRJ DG 408-R; Pissarrachampsa sera LPRP 0019, 0018, 0020; 87 ~ 23 those yielding baurusuchines in Sao Paulo. Stratiotosuchus maxhechti DGM 1477. 88 24 Here, based on a first-hand comparative analysis of most rele- 89 25 vant specimens, we describe a baurusuchid partial skull discovered 3.2. Institutional abbreviations 90 26 in Fazenda Sao~ Jose( Fig. 1), near Fazenda Buriti, which has enough 91 27 morphological evidence to be assigned to Pissarrachampsinae, CCPLIP- Centro de Pesquisas Paleontologicas, L.I. Price, Uberaba; 92 fi 28 speci cally to Campinasuchus dinizi. Campinasuchus dinizi was DGM- Museu de Ciencias^ da Terra do Departamento Nacional de 93 fi 29 described based rst on cranial (Carvalho et al., 2011) and, poste- Produçao~ Mineral, Rio de Janeiro; LPRP- Laboratorio de Paleon- 94 ^ 30 riorly, on postcranial (Cotts et al., 2017). material from the Tres tologia de Ribeirao~ Preto; MPMA- Museu de Paleontologia “Prof. 95 fi 31 Antas Farm, in Minas Gerais. This is the rst record of pissarra- Antonio Celso de Arruda Campos”; UCR- Universidade Estadual 96 ~ 32 champsines in Sao Paulo, challenging previous assumptions that Paulista, Rio Claro; UFRJ-DG- Coleçao~ de Macrofosseis da Uni- 97 33 the taxon was restricted to the of Minas Gerais. versidade Federal do Rio de Janeiro, Departamento de Geologia. 98 34 99 35 2. Geological settings 100 4. Systematic paleontology 36 101 37 The specimen herein described was found at Fazenda Sao~ Jose(20  102 Mesoeucrocodylia Whetstone & Whybrow, 1983 sensu Benton & 38 360 59,800 S50 290 18,300 W), Prudencio^ e Morais district, near General 103 Clark, 1988 39 Salgado, Sao~ Paulo State, in a field work leaded by Prof. Joao~ Tadeu 104 Notosuchia Gasparini 1971 sensu Sereno et al., 2001 40 Arruda. The deposits exposed at Fazenda Sao~ Jose (see Castro et al., 105 Eunotosuchia Ruiz, Bronzati, Ferreira, Martins, Queiroz, Langer & 41 2018) belong to the Vale do Rio do Peixe Formation described by 106 Montefeltro, 2021 42 Fernandes & Coimbra (2000), which corresponds to most of the 107 Xenodontosuchia Ruiz, Bronzati, Ferreira, Martins, Queiroz, Langer 43 Adamantina Formation as described by Soares et al. (1980).Suchbeds 108 & Montefeltro, 2021 44 were radioisotopicaly dated in Fazenda Buriti by Castro et al. (2018), 109 Baurusuchidae Price, 1945 [Darlim, Montefeltro & Langer, 2021] 45 resulting in a late Coniacian to late maximum deposi- 110 Pissarrachampsinae Montefeltro, Larsson & Langer 2011 [Darlim, 46 tional age, although the authors considered as the best 111 Montefeltro & Langer, 2021] 47 age estimate. Previous studies based on biostratigraphic data from 112 Campinasuchus dinizi Carvalho, Teixeira, Ferraz, Ribeiro, Martinelli, 48 vertebrates, ostracods, and Charophyta broadly inferred a to 113 Neto, Sertich, Cunha, Cunha, Ferraz, 2011 49 Maastrichtian age for the Adamantina/Vale do Rio do Peixe Formation 114 50 (Gobbo-Rodrigues et al., 1999; Dias-Brito et al., 2001; Santucci and Locality and Horizon. Adamantina/Vale do Rio do Peixe Formation, 115 51 Bertini, 2001; Marinelli et al., 2011). Bauru Group (Soares et al., 1980; Fernandes & Coimbra, 1996; 116 52 Apart from the already mentioned baurusuchids Baurusuchus Batezelli, 2005); Upper Cretaceous (ca. ConiacianeCampanian) of 117 53 salgadoensis, Baurusuchus albertoi, Gondwanasuchus scabrosus, and the Bauru Basin (Castro et al., 2018). 118 54 Aplestosuchus sordidus (Carvalho et al., 2005; Nascimento & Zaher, Emended diagnosis. Baurusuchid crocodyliform characterized by the 119 55 2010; Marinho et al., 2013; Godoy et al., 2014), Fazenda Buriti has following combination of features ( marked with 120 56 also yielded other crocodyliform material, including eggs and the an asterisk): a short, low, laterally compressed rostrum; cranial 121 57 sphagesaurid stenognathus, as well as a table higher than dorsal border of rostrum; four premaxillary and 122 58 tooth, and snake vertebrae (Zaher et al., 2003; Pol et al., 2014; five maxillary teeth; marked heterodonty, with third maxillary and 123 59 Castro et al., 2018), showing the fossiliferous richness of the out- fourth dentary teeth extremely enlarged relative to other teeth; last 124 60 crops around General Salgado. maxillary tooth placed almost at the level of anterior border of 125 61 palatal (suborbital) fenestra; posteroventrally projecting quadrate; 126 62 3. Material and methods pit on the premaxilla for reception of first mandibular tooth placed 127 63 lateral to first premaxillary tooth (or between first and second 128 64 UFRJ DG 402-R consists of a partially preserved skull containing premaxillary teeth); marked constriction of the skull posterior to 129 65 both premaxillae, maxillae, nasals, and palatines, the anterior portion last maxillary tooth; ten dentary teeth; large anteroposterior 130

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1 66 2 67 3 68 4 69 5 70 6 71 7 72 8 73 9 74 10 75 11 76 12 77 13 78 14 79 15 80 16 81 17 82 18 83 19 84 20 85 21 86 22 87 23 88 24 89 25 90 26 91 27 92 28 93 29 94 30 95 31 96 32 97 33 98 34 99 35 100 36 101 37 102 38 103 39 104 40 105 41 106 42 107 43 108 44 109 45 110 46 111 47 112 48 113 49 114 50 115 51 116 52 117 53 118 54 119 55 120 56 121 57 122 Fig. 1. Distribution of baurusuchids. A, Map of Brazil focusing on the Adamantina/Vale do Rio do Peixe Fm. in Sao~ Paulo and Minas Gerais states (modified from Fernandes, 2004); B, 58 123 Schematic drawing showing the locality of the main outcrops near General Salgado-SP. Silhouettes indicate the baurusuchids subgroups: Baurusuchinae of Fazenda Buriti in black 59 (Baurusuchus albertoi, Baurusuchus salgadoensis, and Aplestosuchus sordidus þ the non baurusuchinae Gondwanasuchus scabrosus); Pissarrachampsine in pink occurring in Minas 124 60 Gerais (Pissarrachampsa sera, and Campinasuchus dinizi) and in Sao~ Paulo (Campinasuchus dinizi). Star indicates de locality of UFRJ DG 402-R. (For interpretation of the references to 125 61 colour in this figure legend, the reader is referred to the Web version of this article.) 126 62 127 63 128 64 depression on each palatine between palatal fenestrae; ventrally suture; slender posterior process of jugal; dorsal nasal-maxillary 129 fl 65 at ectopterygoid surface; anteriorly semi-circular maxilla-jugal suture strongly laterally concave; nasal-frontal contact reduced; 130

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1 66 2 67 3 68 4 69 5 70 6 71 7 72 8 73 9 74 10 75 11 76 12 77 13 78 14 79 15 80 16 81 17 82 18 83 19 84 20 85 21 86 22 87 23 88 24 89 25 90 26 91 27 92 28 93 29 94 30 95 31 96 32 97 33 98 34 99 35 100 36 101 37 102 38 103 39 104 40 105 41 106 42 107 43 108 44 109 45 110 46 111 47 112 48 113 49 114 50 115 51 116 52 117 53 118 54 119 55 120 56 121 57 Fig. 2. Campinasuchus dinizi specimen UFRJ DG 402-R. A right lateral view of cranium and mandible articulated; B ventral view of the cranium; C dorsal view of cranium; D close up 122 58 of premaxilla-maxilla lateral suture within the notch for the reception of D4; E, left lateral view of cranium and mandible articulated. d, dentary; ect, ectopterygoid; f, foramen; if, 123 foramen incisivum; j, jugal; l, lacrimal; m1, maxillary tooth 1; mx, maxilla; n, nasal; nd, nasal depression; op, occlusal pit; pal, palatine; pmx, premaxilla. Scale bar 10 cm (2 cm in D). 59 124 60 125 61 ventral depression on splenial at mandibular symphysis; anteriorly 5. Description 126 62 convex dentary-splenial suture at mandibular symphysis in ventral 127 63 view; conspicuously flattened ventral surface of palatine*. Premaxilla. Both premaxillae are well preserved and articu- 128 64 lated. The bone is subtriangular in lateral view. Anteriorly, its dorsal 129 65 130

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1 projection forms an anteroposteriorly and lateromedially thin is also heavily ornamented at the ventral and anteroventral borders 66 2 internarial bar, the dorsal tip of which is slightly curved posteriorly, of the orbit and is marked dorsally by a conspicuous infraorbital 67 3 forming an anteriorly convex surface. Dorsally, that tip contacts the ridge (sensu Montefeltro et al., 2011). Ventral to the ridge, elongated 68 4 pointed anteroventral tip of the nasal. Posterior to that, there is a and dorsoventrally directed grooves are seen along the bone. Also, 69 5 relatively shallow perinarial fossa that is displaced laterally and the ventral margin of the infraorbital region is conspicuously in- 70 6 extended along the entire premaxillary margin of the nasal cavity. clined anteroventrally towards the suture with the maxilla. The 71 7 Bordering the cavity, the fossa gradually becomes less marked antorbital portion of the jugal contacts the maxilla and the lacrimal, 72 8 anteriorly, towards the base of the internarial bar. In lateral view, but the sutures are not discernible due to the high ornamentation 73 9 the premaxillary body has a concave posterior margin that forms of that surface. The dorsal contact with the lacrimal is inferred by 74 10 the anterior border of the premaxilla-maxilla notch for the recep- the dorsal end of the infraorbital ridge (Fig. 3D). Ventrally, the jugal 75 11 tion of the dentary caniniform (D4). Within this notch, the articulates with the dorsal projection of the ectopterygoid. 76 12 premaxilla-maxilla is marked by an ovate foramen (Fig. 2D). From Lacrimal. The lacrimal is only preserved in the left side, but lacks 77 13 the anterior margin of the notch, there is an elongated depression its posteriormost portion. Laterally, the lacrimal surface is less 78 14 extending anteroventrally at the lateral surface of the premaxilla. ornamented than that of the jugal and maxilla. Anteriorly, the 79 15 The premaxilla is well ornamented, the right bone bears one fo- lacrimal contacts the maxilla by an extended dorsoventrally linear 80 16 ramen right posterior to the perinarial fossa, whereas two foramina suture. From its ventral contact with the jugal, the lacrimal extends 81 17 in the same portion are seen on the left side. dorsomedially, projecting laterally at its dorsal margin to form a 82 18 In ventral view, the premaxilla has a subtriangular shape. The platform for the anterior palpebral. Such contact between the 83 19 medial margin of the alveoli is marked by thin and elongated grooves lacrimal and the anterior palpebral precludes a dorsal exposure of 84 20 and few foramina scattered along the surface. Also, two ante- the former bone. Furthermore, the dorsolateral projection provides 85 21 roposteriorly elongated occlusal pits for the insertion of the first to the lacrimal a laterally concave shape in posterior view. In that 86 22 dentary teeth are present between the first and second premaxillary same view, a perforation is seen slightly below the lateral inflexion of 87 23 alveoli. The posteromedial corners of the ventral surface of the the bone, which may represent the lacrimal duct. 88 24 premaxillae forms the anterior margin of the foramen incisivum. Anterior palpebral. The anterior palpebral is poorly preserved 89 25 Maxilla. The maxilla is a heavily ornamented bone contacting the and only its anteriormost portion can be observed at the left side. In 90 26 nasal dorsally, the premaxilla anteriorly, the lacrimal posterodorsally, lateral view, the bone is ornamented and linear being slightly 91 27 and the jugal posteroventrally. The lateral alveolar margin of the anterior to posteriorly inclined, and it can be seen in articulation 92 28 bone is pierced by nine small foramina. Anteriorly, the maxilla has a with the dorsal surface of the lacrimal. In dorsal view, a clear suture 93 29 convex lateral projection that forms the posterior portion of the is present possibly contacting anteriorly a posterolateral small 94 30 premaxilla-maxilla notch for the insertion of the caniniform dentary portion of the nasal (Fig. 3), and medially the lateral surface of the 95 31 tooth (D4). In lateral view, the ventral margin of the maxilla is convex highly damaged prefrontal. 96 32 where the bone bears teeth and becomes straight posterior to the Palatine. The palatine pair forms a subtriangular element in 97 33 last maxillary tooth (M5). This straight portion forms the posterior ventral view (Fig. 2B). It contacts the maxilla anteriorly by a slightly 98 34 ramus of maxilla, which is lateromedially constricted. As mentioned anteriorly convex suture, which converges posteriorly near the 99 35 above, the anterior suture with the premaxilla is marked by an ovate sagittal line. The slightly concave lateral margin of the palatines 100 36 foramen, at the premaxilla-maxilla notch (Fig. 2D). Forming the forms the medial margin of the suborbital fenestrae. The anterior 101 37 posterior border of such foramen, the ventral portion of the anterior portion of the palatine is tubular, with a generally lateromedially 102 38 margin of the maxilla is anteriorly projected. As in the premaxilla, convex ventral surface, but bearing a parasagittally elongated sul- 103 39 medial to the alveoli, the ventral surface of the maxilla is orna- cus. More posteriorly, the bone gradually expands laterally, forming 104 40 mented and bears scattered foramina. In that area, between the a flat ventral surface. It is possible to observe some foramina scat- 105 41 fourth and fifth maxillary teeth, there is a marked occlusal pit for the tered on both the sulci and the flattened area. The posterolateral 106 42 reception of the ninth dentary tooth (D9). ramus of the palatine contacts the ectopterygoid. 107 43 The maxilla articulates ventromedially with its antimere via a In posterior view, even that the dorsal area is damaged, it is 108 44 linear suture. Parasagitally to this suture, at its posteriormost possible to observe at the anterior tubular portion the dorsal con- 109 45 portion that contacts palatines, the maxilla bears two inconspic- tact with the ventral area of the vomer, forming a continuous sur- 110 46 uous elongated depressions. Posteriorly, the maxilla articulates face laterally. Medially it is possible to observe that the contact of 111 47 with the palatine via a slightly anteriorly convex suture and forms both bones forms the nasopharyngeal duct (sensu Witmer, 1995) 112 48 the anteromedial border of the suborbital fenestra, the antero- which is filled with rock matrix. 113 49 lateral margin of which is formed by the posterior ramus of the Ectopterygoid. The ectopterygoid is only preserved on the left 114 50 maxilla. That ramus contacts the dorsal projection of the ectop- side and presents two major portions: the ventral or ectopterygoid 115 51 terygoid medially and the anteriormost portion of jugal laterally. wings and the dorsolateral projection. The ventral portion of the 116 52 Nasal. The nasals are partially fused to one another, but the ectopterygoid contacts the palatine medially and the pterygoid 117 53 suture is still recognizable anteriorly (Fig. 3C). It is an ante- posteriorly (Fig. 2B). It is ventrally flattened and the ventralmost tip 118 54 roposteriorly elongated bone, with an anteriorly project and of the ectopterygoid wing is posteriorly oriented. The dorsolateral 119 55 ventrally curved portion of the internarial bar that contacts an projection contacts the posterior ramus of the maxilla anteriorly 120 56 equivalent process of the premaxilla. In dorsal view, this projection and the medial portion of the jugal laterally. Its medial margin 121 57 is conspicuous and gives this part of the skull a pointed shape. The forms the posterior and posterolateral borders of the suborbital 122 58 dorsal surface of the nasal is flat and ornamented. More posteriorly fenestra. 123 59 the bone bears a rugose concavity, bordered by the anterior 124 60 palpebral and supposedly by the anterior margin of the prefrontals, 5.1. Mandible 125 61 however this portion is damaged making it difficult to access such 126 62 information. Laterally, the nasal contacts the dorsal surface of the Dentary. The dentaries are articulated to one another, but only 127 63 premaxillae and maxillae via linear suture. the portion anterior to the external mandibular fenestra is pre- 128 64 Jugal. The jugal is only partially preserved on the left side, served. It is a highly ornamented bone, including the symphyseal 129 65 including part of the infraorbital and antorbital portions. The bone area and the diverging posterior portion. The conjoined mandibular 130

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1 pmx 66 2 67 3 68 4 69 5 A B pm3 70 6 pm4 71 7 n 72 8 if 73 9 74 10 75 11 76 12 mx 77 13 78 14 79 15 80 16 m5 op 81 17 pmx 82 18 83 19 m1 84 20 m3 85 21 86 22 d 87 23 j 88 24 89 25 ect 90 26 n 91 27 C 92 28 pmx pal 93 29 94 30 95 31 ect 96 32 97 33 98 34 ap 99 35 mx 100 36 D 101 37 l 102 38 103 39 nd n 104 40 j 105 41 ap spmx 106 42 107 43 108 44 j 109 45 110 46 111 47 112 48 113 49 pmx 114 50 115 51 116 52 pal d 117 53 118 54 119 55 120 56 ect 121 57 122 58 Fig. 3. Schematic drawings of Campinasuchus dinizi UFRJ DG 402-R. A right lateral view of the cranium and mandible articulated; B ventral view of cranium; C dorsal view of 123 cranium; D left lateral view of cranium and mandible articulated. ap, anterior palpebral; d, dentary; ect, ectopterygoid; if, foramen incisivum; j, jugal; l, lacrimal; mx, maxilla; m1-3, 59 maxillary teeth; n, nasal; nd, nasal depression; op, occlusal pit; pal, palatine; pmx, premaxilla; pm 3e4, premaxillary teeth. Scale bar: 10 cm. 124 60 125 61 126 symphyseal area is ovate in dorsal/ventral views, with its most starts diverging posterior to that. In lateral view, the symphyseal 62 127 mediolaterally wide portion at the level of D4 (Fig. 4A). The lower area is dorsally projected, as typical of baurusuchids, whereas and 63 128 jaw is constricted posterior to that, until the seventh dentary tooth. the dorsal margin of the mandibular rami gets conspicuously in- 64 129 This is the most mediolaterally narrow part of the mandible, which clined dorsally as it approaches the external mandibular fenestra. 65 130

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1 When the skull is articulated, it is possible to notice that the the anterior border of the internal mandibular fenestra and the 66 2 inclination of the ventral margin of the jugal fits that of the medial wall of the Meckelian canal. The splenial is only orna- 67 3 mandible (Fig. 2E). This area also bears a longitudinal sulcus, mented in the dorsal portion of its medial surface, where a bulged 68 4 positioned dorsal to the fenestra. In dorsal view, the medial portion suture with its antimere is present in the symphyseal area. This 69 5 of the symphyseal area is also ornamented and the suture with the suture ends posteriorly in a peg-like protuberance (Fig. 4A). In 70 6 antimere is slightly bulged. The ventral surface of the dentary is as ventral view, lateral to the symphysis, each splenial bears an 71 7 ornamented as the lateral, with the medial suture with the splenial anteroposteriorly directed shallow sulcus (Figs. 4D, 5D). The 72 8 and angular clearly seen. ventral extension if this sulcus, at the posterior face of the sym- 73 9 Splenial. Both splenials are completely preserved. The splenial physeal area, is deeper than the inter-splenial suture. A large, 74 10 comprises in a thin bone, located at the anteromedial portion of anteroposteriorly elongated intramandibularis oralis foramen 75 11 the mandible, contacting the dentary anterior and laterally, and perforates the medial surface of the splenial, slightly posterior to 76 12 the angular posteriorly. The posterior margin of the splenial forms the peg-like protuberance. 77 13 78 14 79 15 80 16 81 17 82 18 83 19 84 20 85 21 86 22 87 23 88 24 89 25 90 26 91 27 92 28 93 29 94 30 95 31 96 32 97 33 98 34 99 35 100 36 101 37 102 38 103 39 104 40 105 41 106 42 107 43 108 44 109 45 110 46 111 47 112 48 113 49 114 50 115 51 116 52 117 53 118 54 119 55 120 56 121 57 122 58 123 59 124 60 125 61 126 62 127 63 128 64 Fig. 4. Mandible of Campinasuchus dinizi UFRJ DG 402-R in A, dorsal; B, left lateral; C, right lateral; and D, ventral views. a, angular; d, dentary; d1-d9, dentary teeth; emf, external 129 65 mandibular fenestra; msy, mandibular symphysis; pms, peg-like projection of mandibular symphysis; spl, splenial; svd, splenial ventral depression. Scale bar: 10 cm. 130

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1 Angular. Only the anteriormost portion of both angulars are and positioned right below the base of nasal septum. The remaining 66 2 preserved. This part of the bone contacts the splenial medially and premaxillary tooth crowns are broken, but it is possible to notice 67 3 the dentary laterally (Fig. 5D). It has a tubular shape, forming the that the teeth gradually increase in size until they reach the hy- 68 4 floor of the Meckelian canal, which extends anteriorly between pertrophied pm3. On the other hand, pm4, located at the anterior 69 5 the dentary and the splenials. margin of the premaxilla-maxilla notch, is the smallest premaxil- 70 6 lary tooth (Fig. 2A). 71 7 5.2. Dentition The maxilla bears five teeth (m1-5), all also with crowns pos- 72 8 teriorly curved on their apicalmost portion and with serrated distal 73 9 The premaxilla possesses four teeth (pm1-4), the crowns of and mesial carinae. M1 is the smallest of those teeth, positioned on 74 10 which are slightly curved posteriorly on their apicalmost portion, the posterior border of the premaxilla-maxilla notch (Figs. 2A, 3A). 75 11 with serrated distal and mesial carinae. Both pm1 are preserved In occlusion, its tip aligns to the posterior portion of the alveolus of 76 12 77 13 78 14 d1 79 15 A B 80 16 81 17 d4 82 18 d4 83 19 84 20 d9 85 21 msy 86 22 87 23 d1 88 24 emf 89 25 90 26 91 27 92 28 93 a 29 pms 94 30 d 95 31 96 32 spl 97 33 D 98 34 d 99 35 100 36 d 101 37 102 38 103 39 104 40 105 41 106 42 107 43 108 44 109 45 d4 110 46 svd 111 C 47 d9 112 48 113 49 114 50 spl 115 51 116 52 117 53 118 54 119 55 120 56 121 57 d 122 58 123 59 a 124 60 125 61 126 62 127 63 128 Fig. 5. Schematic drawings of the mandible of Campinasuchus dinizi UFRJ DG 402-R in A, dorsal; B, left lateral; C, right lateral; and D, ventral views. a, angular; d, dentary; d1-d9, 64 dentary teeth; emf, external mandibular fenestra; msy, mandibular symphysis; pms, peg-like protuberance of mandibular symphysis; spl, splenial; svd, splenial ventral depression. 129 65 Scale bar: 10 cm. 130

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1 66 2 67 3 68 4 69 5 70 6 71 7 72 8 73 9 74 10 75 11 76 12 77 13 78 14 79 15 80 16 81 17 82 18 83 19 84 20 85 21 86 22 87 23 88 24 89 25 90 26 91 27 92 28 93 29 94 30 95 31 96 32 97 33 98 34 99 35 100 36 101 37 102 38 103 39 104 40 105 41 106 42 107 43 108 44 109 45 110 46 111 47 112 48 113 49 114 50 Fig. 6. Comparison between the choanal region of Baurusuchidae subgroups. A, Pissarrachampsinae (Pissarrachampsa sera, LPRP 0019); B, Baurusuchinae (Aphaurosuchus 115 escharafacies, LPRP 0697). C, schematic drawing LPRP 0019 and D, LPRP 0697 respectively. cs, choanal septum; ec, ectopterygoid; pcf, parachoanal fenestra; pcfo, parachoanal fossae; 51 pl, palatine; pt, pterygoid. Scale bar: 5 cm. 116 52 117 53 118 54 the fourth dentary tooth (d4). The maxillary dentition also in- dorsally until the dorsal surface of the nasal (Fig. 2E). D9 is located 119 55 creases in size posteriorly, with m2 almost three times longer than in a dorsally raised area of the dentary and inserts in a poster- 120 56 m1, and m3 representing a conspicuously hypertrophied tooth, omedial occlusal pit of the maxilla. The remaining teeth are smaller 121 57 extending in occlusion until the ventral border of the dentary. M4 is and of almost the same size (Figs. 4A, 5A). 122 58 in almost of the same size as m2, as is m5 in relation to m1. 123 59 The dentary bears ten teeth (d1-d10), also with the ziphodont 6. Discussion 124 60 configuration, with three of them being the largest of the series. D1 125 61 is procumbent (sensu Godoy, 2011) forming an angle of ca. 45 to In the original definition of Pissarrachampsinae, Montefeltro 126 62 the general tooth line (Fig. 4B). In occlusion, it is inserted on an et al. (2011) listed key features for the identification of the group 127 63 anteroposterioly elongated depression in the premaxilla, between and its differentiation from Baurusuchinae (e.g., posterior portion 128 64 pm1 and pm2. D4 is the hypertrophied caniniform tooth that in of the nasal bearing a rugose broad depression, approximation of 129 65 occlusion inserts in the premaxilla-maxilla notch, extending the prefrontals along their medial edges anteriorly, and presence of 130

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1 66 2 67 3 68 4 69 5 70 6 71 7 72 8 73 9 74 10 75 11 76 12 77 13 78 14 79 15 80 16 81 17 82 18 83 19 84 20 85 21 86 22 87 23 88 24 89 25 90 26 91 27 92 28 93 29 94 30 95 31 96 32 97 33 98 34 99 35 100 36 101 37 102 38 103 39 104 40 105 41 106 42 107 43 108 44 109 45 110 46 111 47 112 48 113 49 114 50 115 51 116 52 117 53 118 54 119 55 120 56 121 57 122 58 123 59 124 60 125 61 Fig. 7. Comparison of the palatine morphology of pissarrachampsines in ventral view. A, palatine of Campinasuchus dinizi (UFRJ DG 402-R) and B, schematic drawing. C, palatine of 126 fl 62 Pissarrachampsa sera (LPRP 0019) and D, schematic drawing. Arrows indicate the extension of the tubular margin. bss, bulged sagittal suture; fs, attened surface; ps, parasagittal 127 sulcus; ts, tubular margin. Scale bar: 5 cm. 63 128 64 129 65 130

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1 a midline longitudinal depression on anterior portion of the fron- (2011), the specimen presents some differences in relation to the 66 2 tal). More recently, following Nascimento (2014), Darlim et al. holotype of Stratiotosuchus maxhechti, such as a foramen between 67 3 (2021) provided a revised definition of the choanal pattern of the the palpebrals, a small posttemporal fenestra and a non-ridged 68 4 group, some of the main features of which include in a pair of choanal septum, morphological features that can be related to 69 5 conspicuous longitudinal row of foramina, two parasagittal sulci on taphonomic distortions and/or ontogenetic variation (Montefeltro 70 6 the ventral surface of the palatine, ectopterygoid and pterygoid et al., 2011, supplementary material). Although the specimen was 71 7 wings posteriorly directed, ectopterygoid mediolaterally flattened, not analyzed first-hand in the present work, we agree that URC 73 72 8 pterygoid forming parachoanal fossae, and choanal septum later- may not represent a Stratiotosuchus maxhechti specimen, and also 73 9 omedially thin (Fig. 6). Although only partially preserved, several of highlight several features in common between URC 73 and pis- 74 10 those diagnostic traits are seen in UFRJ DG 402-R, including a sarrachampsines (sensu Darlim et al., 2021): palatine presenting a 75 11 rugose depression on the posterior portion of the nasals, conspic- flattened ventral surface with two conspicuous parasagittal sulci, 76 12 uous ventral palatine sulci, ectopterygoid mediolaterally flattened posteriorly oriented and mediolaterally flattened ectopterygoid, 77 13 and posteriorly directed ectopterygoid, supporting its identification conspicuously mediolaterally thin choanal septum, and post- 78 14 as a pissarrachampsine. choanal portion of the pterygoid anteroposteriorly short and 79 15 Among pissarrachampsines, the fragmentary nature of Wargo- dorsally recessed at the contact with the parabasisphenoid. 80 16 suchus australis and scarce overlapping of parts between UFRJ DG 81 17 402-R and the Argentinean taxon prevented their comparison. In 6.1. Paleogeographic distribution of Bauru Basin baurusuchids 82 18 relation to Pissarrachampsa sera, UFRJ DG 402-R does not have its 83 19 ridged jugal-ectopterygoid suture exposed in lateral view, which is Based on the Santonian age of the Argentinean pissarra- 84 20 instead restricted the medial portion of the jugal. Also, the lateral champsine Wargosuchus australis, Montefeltro et al. (2011) sug- 85 21 surface of the latter bone is not pierced by small foramina as in gested that the Pissarrachampsinae-bearing deposits of the 86 22 Pissarrachampsa sera. Additionally, the number of maxillary teeth Adamantina/Vale do Rio do Peixe Formation in Minas Gerais could 87 23 could also distinguish those taxa, given that Pissarrachampsa sera be older than those in Sao~ Paulo. Indeed, all Bauru Basin baur- 88 24 was described as having four maxillary teeth, the second of which usuchines come from Sao~ Paulo, whereas pissarrachampsines were 89 25 being the largest. However, given that the anteriormost portion of until now restricted to Minas Gerais. Yet, UFRJ DG 402-R shows that 90 26 the maxilla (i.e., forming the posterior margin of the premaxilla- the latter also occurs in Bauru Basin rocks of Sao~ Paulo, in a 91 27 maxilla notch) is damaged in both the holotype (LPRP/USP 0019) site (Fazenda Sao~ Jose) nearby the most diverse site for Baur- 92 28 and paratype (LPRP/USP 0018) of Pissarrachampsa sera, we do not usuchinae (Fazenda Buriti) and within the same geological context, 93 29 consider this as reliable evidence to differentiate UFRJ DG 402-R revealing the sympatric occurrence of both major Baurusuchidae 94 30 from that taxon. groups (Fig. 1). 95 31 Our comparative analysis also revealed that previously pro- The joint occurrence of pissarrachampsines and baurusuchines in 96 32 posed autapomorphies of Campinasuchus dinizi (i.e., conspicuously Sao~ Paulo falsifies the hypothesis that those groups were chrono- 97 33 large third maxillary and fourth dentary teeth and an ante- logically separated, weakening the assumption that the deposits of 98 34 roposteriorly elongated depression on the palatine) are actually the Adamantina/Vale do Rio do Peixe Formation in Minas Gerais are 99 35 shared with Pissarrachampsa sera (LPRP 0049, see Godoy et al., older than those in Sao~ Paulo. Radioisotopic ages for the 100 36 2018, Fig, 1) and not diagnostic of the former species. However, Pissarrachampsinae-bearing deposits in Minas Gerais and Argentina, 101 37 we propose that a short anterior tubular area and a large flattened together with the discovery of better preserved Argentinean baur- 102 38 ventral surface of the palatine is unique to Campinasuchus dinizi usuchid material will be important to further test these proposals 103 39 (Fig. 7A,B), whereas a more tubular-shaped palatine, with a and to better understand the past distribution of baurusuchids. For 104 40 rounded ventrolateral surface that is more anteroposteriorly now, the tentative Campanian dating of the Adamantina/Vale do Rio 105 41 expanded, is typical of Pissarrachampsa sera (holotype LPRP/USP do Peixe Formation by Castro et al. (2018), suggests that it is slightly 106 42 0019; Fig. 7C,D). Yet, it is important to highlight that even having a younger than the Santonian Bajo de La Carpa Formation. This could 107 43 longer tubular portion, the palatine of Pissarrachampsa sera still suggest a more southern origin for Baurusuchidae and their later 108 44 differs from those of Baurusuchinae (sensu Darlim et al., 2021)by north-western dispersion. Such pattern agrees with the inferred 109 45 having marked parasagittal sulci and not being medially conver- tectonic dynamics of South America during the Cretaceous, where 110 46 gent (e.g., not forming a lateromedially constricted palatine; Fig. 6). drainage connections between southern Argentina and central South 111 47 Based on the above observations, we also suggest the reas- America, from Aptian to the Late Cretaceous, probably allowed the 112 48 signment of some previously published specimens of both pissar- interchange of the crocodyliform fauna (Carvalho et al., 2010). Yet, it 113 49 rachampsines species. LPRP/USP 0018 assigned to Pissarrachampsa is important to highlight that paleobiogeographic reconstructions, 114 50 sera by Montefeltro et al. (2011, Figure 11) has a conspicuously including the identification of ancestral areas and dispersal events, 115 51 flattened ventral surface of the palatine similar to that of Campi- could be biased by several factors, such as the uncertain age of the 116 52 nasuchus dinizi, whereas CPPLIP 1236, one of the paratypes of Adamantina/Vale do Rio do Peixe Formation, the high endemicity of 117 53 Campinasuchus dinizi (Carvalho et al., 2011, Figure 5), preserves only Baurusuchinae, restricted to Sao~ Paulo state, and the high diversity of 118 54 part of the palatine, but clearly with more conspicuously rounded/ baurusuchids in the Adamantina/Vale do Rio do Peixe Formation 119 55 bulged lateral margins, medially restricting the parasagittal sulci, as (Celis et al., 2020) in comparison with the more fragmentary baur- 120 56 seen in the holotype of Pissarrachampsa. sera (Fig. 7C). Hence, LPRP/ usuchid record in the Neuquen Basin, Argentina (Leardi et al., 2018). 121 57 USP 0018 and CPPLIP 1236 are here tentatively reassigned respec- 122 58 tively to Campinasuchus dinizi and Pissarrachampsa sera, but it is 7. Conclusions 123 59 clear that more comprehensive studies are needed to understand 124 60 the morphological variation of the baurusuchids of the Ada- The new specimen described here (UFRJ DG 402-R) is recog- 125 61 mantina/Vale do Rio do Peixe Formation in Minas Gerais. nized as a Pissarrachampsinae, specifically as Campinasuchus dinizi, 126 62 Additionally, the specimen URC 73, consisting of a fragmentary due to the presence of a ventrally flattened and posteriorly directed 127 63 cranium and mandible, referred as Stratiotosuchus maxhechti by ectopterygoid, palatines ventrally flattened with two parasagittal 128 64 Pinheiro et al. (2008) also has morphological similarities with sulci, short anterolateral tubular margins and a large flattened 129 65 pissarrachampsines. As previously noted by Montefeltro et al. ventral surface of the palatine, and a depressed posterior portion of 130

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1 the nasal dorsal surface. This identification resulted from a short Celis, A., Narvaez, I., Arcucci, A., Ortega, F., 2020. Lagerstatte€ effect drives noto- 66 e 2 review of the baurusuchid palatine morphology, allowing the dif- suchian palaeodiversity (, Notosuchia). Historical Biology 1 10. 67 Cotts, L., Pinheiro, A.E.P., Marinho, T.S., Carvalho, I.S., Di Dario, F., 2017. Postcranial 3 ferentiation of Campinasuchus dinizi from Pissarrachampsa sera and skeleton of Campinasuchus dinizi (Crocodyliformes, Baurusuchidae) from the 68 4 a tentative reassignment of previously published specimens of both Upper Cretaceous of Brazil, with comments on the ontogeny and ecomor- 69 e 5 species. UFRJ DG 402-R represents the first record of Pissarra- phology of the species. Cretaceous Research 70, 163 188. 70 Darlim, G., Montefeltro, F.C., Langer, M.C., 2021. 3D skull modelling and description 6 champsinae in the Adamantina/Vale do Rio do Peixe Formation of of a new baurusuchid (Crocodyliformes, Mesoeucrocodylia) from the Late 71 7 Sao~ Paulo. This demonstrates that, during the Late Cretaceous, Cretaceous (Bauru Basin) of Brazil. Journal of Anatomy 1e41. 72  8 pissarrachampsines lived together with baurusuchines in the re- Dias-Brito, D., Musacchio, E.A., de Castro, J.C., Maranhao, M.S.A.S., Suarez, J.M., 73 Rodrigues, R., 2001. Grupo Bauru: uma unidade continental do Cretaceo no 9 gion, without any noticeable palaeobiogeographic barrier. Brasil - concepçoes~ baseadas em dados micropaleontologicos, isotopicos e 74 10 estratigraficos. Revue de Paleobiologie 20, 245e304. 75  11 CRediT authorship contribution statement Fernandes, L.A., 2004. Mapa litoestratigrafico da parte oriental da Bacia Bauru (PR, 76 12 SP, MG), escala 1: 1.000. 000. Boletim Paranaense de Geociencias 55. 77 Fernandes, L.A., Coimbra, A.M., 1996. Bacia Bauru (Cretaceo superior, Brasil). Anais 13 Gustavo Darlim: Writing e review & editing. Ismar de Souza da Academia Brasileira de Ciencias^ 68, 195e205. 78 14 Carvalho: Writing e review & editing. Sandra Aparecida Sim- Fernandes, L.A., Coimbra, A.M., 2000. Revisao~ estratigrafica da parte oriental da 79  e 15 ionato Tavares: e Max Cardoso Langer: Bacia Bauru (Neocretaceo). Revista Brasileira de Geociencias 30, 717 728. 80 Writing review & editing. Fernandez-Dumont, M.L., Bona, P., Pol, D., Apesteguía, S., 2020. New anatomical 16 Writing e review & editing. information on buitreraensis with implications for the system- 81 17 atics of Uruguausuchidae (Crocodyliforms, Notosuchia). Cretaceous Research 82 18 Declaration of competing interest 113, 104494. 83 Fiorelli, L.E., Leardi, J.M., Martín, E., Pol, D., Basilici, G., Grellet-Tinner, G., 2016. A new 19 Late Cretaceous crocodyliform from the western margin of Gondwana (La Rioja 84 20 The authors declare that they have no known competing Province, Argentina). Cretaceous Research 60, 194e209. 85   21 financial interests or personal relationships that could have Gasparini, Z.B., 1971. Los Notosuchia del Cretacico de America del Sur como um 86 nuevo Infraorden de los Mesosuchia (). Ameghiniana 8, e83ee103. Q5 22 appeared to influence the work reported in this paper. Gasparini, Z., 1972. Los (Crocodilia) del territorio argentino. Consid- 87 23 eraciones sobre su "status" taxonomico. Ameghiniana 9, 23e34. 88 ^  24 Acknowledgments Gobbo-Rodrigues, S.R., Petri, S., Bertini, R.J., 1999. Ocorrencias de ostracodes na 89 Formaçao~ Araçatuba do Grupo Bauru, Cretaceo Superior da Bacia do Paranae 25 possibilidades de correlaçao~ com depositos isocronos argentinos - parte i: 90 26 We want to thank Joao~ Tadeu Arruda and Antonio Celso de Familia Hylocyprididae. Acta Geologica Leopoldensia 23, 3e13. 91 27 fi Godoy, P.L., 2011. Evoluçaeo dos dentes procumbentes em Mesoeucrocodylia,com a 92 Arruda Campos (in memorian) for the efforts in the eld work. ̧ e 28 descricao da mandíbula de um possível Baurusuchidae juvenil (Croc- 93 Gustavo Darlim thanks Fellipe Muniz, Julian Silva, and Thiago Fac- odyliformes, Mesoeucrocodylia). Universidade de Sao~ Paulo, Monografia. 29 cini, for helping during the visits to paleontological collections, and Godoy, P.L., Montefeltro, F.C., Norell, M.A., Langer, M.C., 2014. An additional Baur- 94 fi 30 for Thiago Marinho (CCPLIP) and Rafael Costa (DGM) for allowing usuchid from the cretaceous of Brazil with evidence of interspeci c predation 95 31 among crocodyliformes. PloS One 9, e97138. 96 the study of baurusuchid fossils for comparison. Finally, we thank Godoy, P.L., Ferreira, G.S., Montefeltro, F.C., Vila-Nova, B.C., Butler, R.J., Langer, M.C., 32 the reviewers Ane de Celis and Felipe C. Montefeltro for the time 2018. Evidence for heterochrony in the cranial evolution of fossil crocodyli- 97 33 invested and for the constructive comments that definitely forms. Palaeontology 61, 543e558. 98 34 ~ Leardi, J.M., Pol, D., Gasparini, Z., 2018. New Patagonian baurusuchids (Croc- 99 improved the manuscript. The research was supported by Fundaçao odylomorpha; Notosuchia) from the Bajo de la Carpa Formation (Upper 35 de Amparo a Pesquisa do Estado de Sao~ Paulo (FAPESP), Fundaçao~ Cretaceous; Neuquen, Argentina): New evidences of the early sebecosuchian 100 36 Carlos Chagas Filho de Amparo a Pesquisa do Estado do Rio de diversification in Gondwana. Comptes Rendus Palevol 17 (8), 504e521. 101 Marinho, T.S., Iori, F.V., de Souza Carvalho, I., Vasconcellos, F.M., 2013. Gondwana- 37 Janeiro (FAPERJ) and by Conselho Nacional de Desenvolvimento 102  suchus scabrosus gen. et sp. nov., a new terrestrial predatory crocodyliform 38 Q4 Científico e Tecnologico (CNPq). Gustavo Darlim is funded by (Mesoeucrocodylia: Baurusuchidae) from the Late Cretaceous Bauru Basin of 103 39 FAPESP [2019/06311-4]. Brazil. Cretaceous Research 44, 104e111. 104 40 Martinelli, A.G., Pais, D.F., 2008. A new baurusuchid crocodyliform (Archosauria) 105 from the Late Cretaceous of Patagonia (Argentina). Comptes Rendus Palevol 7, 41 References 371e381. https://doi.org/10.1016/j.crpv.2008.05.002. 106 42 Martinelli, A.G., Riff, D., Lopes, R.P., 2011. Discussion about the occurrence of the 107 43 Barrios, F., Bona, P., Carabajal, A.P., Gasparini, Z., 2018. Re-description of the cranio genus Aelosaurus Powell 1987 (Dinosauria, ) in the Upper Creta- 108 e 44 mandibular anatomy of terrestris (Crocodyliformes, Meso- ceous of Brazil. Gaea 7, 34 40. 109 eucrocodylia) from the Upper Cretaceous of Patagonia. Cretaceous Research 83, Montefeltro, F.C., Larsson, H.C.E., Langer, M.C., 2011. A new baurusuchid (Croc- 45 3e39. odyliformes, Mesoeucrocodylia) from the late cretaceous of Brazil and the 110 46 Batezelli, A., 2015. Continental systems tracts of the Brazilian Cretaceous Bauru phylogeny of Baurusuchidae. PloS One 6, e21916. https://doi.org/10.1371/ 111 47 Basin and their relationship with the tectonic and climatic evolution of South journal.pone.0021916. 112 America. Basin Research 29, 1e25. Montefeltro, F.C., Lautenschlager, S., Godoy, P.L., Ferreira, G.S., Butler, R.J., 2020. 48 Batezelli, A., Gomes, N.S., Perinotto, J.A.D.J., 2005. Petrografia e evoluçao~ diagenetica A unique predator in a unique ecosystem: modelling the apex predator from 113 49 dos arenitos da porçao~ norte e nordeste da Bacia Bauru (Cretaceo Superior). the Late Cretaceous crocodyliform-dominated fauna in Brazil. 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PloS One 9, e93105. 125 Carvalho, I.S., Teixeira, V.P., Ferraz, M.L., Ribeiro, L.C.B., Martinelli, A.G., Neto, F.M., Price, L.I., 1945. A new from the Cretaceous of Brazil. Notas Preliminares e ~  61 2011. Campinasuchus dinizi gen. et sp. nov., a new Late Cretaceous baurusuchid Estudos. Divisao de Geologia e Mineralogia. Ministerio da Agricultura. Rio de 126 62 (Crocodyliformes) from the Bauru Basin, Brazil. Zootaxa 287, 19e42. Janeiro, Brasil 25, 1e8. 127 63 Castro, M.C., Goin, F.J., Ortiz-Jaureguizar, E., Vieytes, E.C., Tsukui, K., Ramezani, J., Ruiz, J.V., Bronzati, M., Ferreira, G.S., Martins, K.C., Queiroz, M.V., Langer, M.C., 128 Batezelli, A., Marsola, J.C.A., Langer, M.C., 2018. A Late Cretaceous mammal from Montefeltro, F.C., 2021. A new species of Caipirasuchus (Notosuchia, Sphage- 64 Brazil and the first radioisotopic age for the Bauru Group. Royal Society of Open sauridae) from the Late Cretaceous of Brazil and the evolutionary history of 129 65 Science 5, 180482. 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