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First Early Cretaceous theropod dinosaur from Brazil with comments on Spinosauridae
Abstract
The discovery of the first theropod dinosaur from the Early Cretaceous of Brazil is reported here. The material consists of the anterior portion of a skull and represents a new taxon, Angaturama limai n.g. n.sp. This dinosaur has an extremely laterally compressed rostrum with a well developed premaxillary sagittal crest. Several synapomorphies suggest that Angaturama is related to the unusual theropods attributed to Spinosauridae, known from Cretaceous strata of different localities in Africa, and Baryonyx, known from the Early Cretaceous of England.
... The dinosaur fossil record from Cretaceous Brazil is especially rich, with about 25 valid genera having been described until recently (Arid and Vizotto 1971;Kellner and Campos 1996;Martill et al. 1996;Kellner 1999, 1999;Kellner and Azevedo 1999;Kellner and Campos 2002;Carvalho et al. 2003;Naish et al. 2004;Kellner et al. 2005Kellner et al. , 2006Salgado and Carvalho 2008;Zaher et al. 2011;Bandeira et al. 2016;Carvalho et al. 2017;Delcourt and Iori 2018;Langer et al. 2019;Lindoso et al. 2019;Sayão et al. 2020;Zaher et al. 2020;Brum et al. 2021;Iori et al. 2021;Souza et al. 2021;Silva-Junior et al. 2022). Up to this date, many studies have aimed at reviewing and cataloguing this known dinosaur diversity in Brazil (Kellner 1996;Bittencourt and Langer 2011;Medeiros et al. 2014;Brusatte et al. 2017;Sales, Martinelli et al. 2017), with most of them focusing mainly on the possible palaeobiogeographic implications Brusatte et al. 2017). ...
... Yet, even amidst the rich theropod record from the Romualdo Formation (n = 149; Table 1) there is a peculiarity: the impressively low number of isolated teeth known for this formation (n = 1). In a similar manner to the Quiricó Formation, the majority of the dinosaur dental record from the Romualdo Formation is known by in situ teeth from spinosaurid genera such as Irritator challengeri (Martill et al. 1996;Sues et al. 2002) and the dubious Angaturama limai (?) (Kellner and Campos 1996;see; for further discussion). In contrast, there is only a single record of an isolated dinosaur tooth for this unit, a spinosaurid tooth embedded in a pterosaur cervical vertebra (Buffetaut et al. 2004). ...
... Spinosaurids have the largest tooth record among the dinosaurs from Cretaceous Brazil (n = 339; Figure 3B), those amounting to 85% ( Figure 4D) of all spinosaurid specimens described in the country up to this moment. The occurrences of Spinosauridae in Brazil are limited to Early Cretaceous to early Late Cretaceous units (Kellner and Campos 1996;Sues et al. 2002;Buffetaut et al. 2004 2018; Carvalho & Santucci 2018), with the oldest South American record of this clade being prospected from the Berriasian-Valanginian Feliz Deserto Formation (Sales, Liparini et al. 2017). The Alcântara Formation has the highest incidence of spinosaurid specimens, with more than 280 isolated teeth (Medeiros 2006) and the problematic taxon Oxalaia quilombensis (Kellner et al. 2011;see Sales &Shultz 2017 and for discussion on the synonymy of this taxon with Spinosaurus), which also has in situ teeth in its holotype. ...
Dinosaur fossils are commonly found in Brazilian Mesozoic strata, with teeth being frequently found in Cretaceous outcrops. Many studies have been made with the aim of reviewing the known diversity of Brazilian dinosaurs, mostly focusing solely on their palaeobiogeography. However, few tried to typify these body fossils in order to see which kinds were most prevalent in the fossil record. This study aims to count and identify all occurrences of non-avian dinosaur teeth (isolated or in situ) unearthed from Brazilian Cretaceous strata in order to objectively measure their representativity in the national fossil record. Our literature search showed that almost half of the dinosaur fossil record from the Cretaceous of Brazil is composed by teeth, those being particularly abundant in the Alcântara, Adamantina and Marília formations. Theropod teeth are more abundant in comparison to sauropod teeth, with spinosaurids, deinonychosaurs, and abelisaurids being amply found. Despite representing a large share of the Brazilian fossil record, dinosaur teeth have only recently been studied in depth, with many specimens being reassessed through the current identification methods. Other aspects besides primary taxonomic identification can also be further explored using the latest technologies on dinosaur teeth, allowing a deeper understanding of the palaeobiology of these reptiles.
... Apenas em 1986 um novo táxon, com material diagnóstico o suficiente, foi nomeado: Baryonyx walkeri Charig & Milner, 1986, dessa vez no Reino Unido, Formação Weald Clay (Barremiano) da Bacia Weald (Charig & Milner, 1986, 1997. Posteriormente, mais táxons e elementos indeterminados foram encontrados na África, América do Sul, Europa, Ásia e elementos que talvez pertençam a Spinosauridae na Austrália, Estados Unidos e Argentina (Kellner & Campos, 1996;Sereno et al., 1998;Benton et al., 2000;Buffetaut & Ouaja, 2002;Sues et al., 2002;Yoshikazu et al., 2003;Canudo et al., 2004;Medeiros, 2006;Buffetaut, 2010Buffetaut, , 2012Buffetaut et al., 2008;Bertin, 2010;Hone et al., 2010;Barret et al., 2011;Kellner et al., 2011;Allain et al., 2012;Medeiros et al., 2014;Hendrickx et al., 2016;Candeiro et al., 2017;Hone & Holtz, 2017, 2021. ...
... Este espinossaurídeo também é proveniente da Formação Romualdo, seu holótipo é o espécime USP GP/2T-5 (Kellner & Campos, 1996;Machado & Kellner, 2007). A diagnose revisada por Sales & Schultz (2017) é a seguinte: espinossaurídeo que difere de Baryonyx, Suchomimus e Cristatusaurus pela ausência de dentículos nas carenas; o primeiro dente pré-maxilar é relativamente menor se comparado aos táxons mencionados anteriormente; ele apresenta uma crista pré-maxilar que se estende até quase o fim do crânio que provavelmente começava mais anteriormente do que no Baryonyx, Suchomimus e Cristatusaurus e a presença de uma projeção ântero-dorsal arredondada na sutura dos pré-maxilares. ...
... Apesar da presença de Irritator na mesma formação geológica, o Angaturama pode ser distinguido de todos os espinossaurídeos que possuem pré-maxilares preservados. Caso mais elementos comprovem que Angaturama e Irritator sejam diferentes indivíduos da mesma espécie, o último nome teria prioridade (Kellner & Campos, 1996 Este foi o primeiro Spinosauridae asiático definitivo, o qual pertence a Formação Grès supérieurs na Bacia de Savannakhet, Laos, e seu holótipo MDS BKS 10-01 15 possui as seguintes autapomorfias: uma vela sinusoidal dorsosacral; o penúltimo processo espinhoso é três vezes mais alto que o centro vertebral e possui um processo ântero-distal com forma de dedo; os processos espinhosos sacrais 3 e 4 têm forma de leque; processos transversos da primeira vértebra caudal com perfil sigmóide em vista dorsal; fossas pré-zigoapofisárias centro-diapofisárias profundas na primeira vértebra caudal; lâmina ilíaca longa com razão de comprimento ilíaco-pubiano mais alta do que em quaisquer outros terópodes; costelas posteriores dorsais articuladas com complexo esternal; placa púbica proximal com forame isquiático e diáfise isquiática achatada médio-lateralmente (Allain et al., 2012;Hone & Holtz 2017). ...
Spinosauridae family (Dinosauria: Theropoda): taxonomy, paleobiogeography and paleoecology (a revision). Spinosauridae is a family of Tetanuran theropod dinosaurs that was widely distributed during the Early Cretaceous. Here we revised the state of art of the family’s taxonomy, paleobiogeography and paleoecology. We compiled updated diagnosis for the holotypes of the 20 species attributed to the family since 1841, alongside with the different hypotheses related to the family’s paleobiogeography and paleoecology. We also compiled updated diagnosis for a series of indeterminate elements that are relevant in literature. We conclude that out of these 20 taxa six can be regarded as nomina dubia (Ostafrikasaurus crassiserratus, Suchosaurus girardi, Spinosaurus maroccanus, Siamosaurus suteethorni, Sinopliosaurus fusuiensis, Suchosaurus cultridens) due to the lack of diagnostic material and autapomorphies. Out of these, three were regarded as incertae sedis (Ostafrikasaurus crassiserratus, Suchosaurus girardi, Suchosaurus cultridens) for the same reasons and the possibility of belonging to previously already established taxa inside Spinosauridae and for one of these (Ostafrikasaurus crassiserratus) for possibly being a member of Ceratosauria. As for paleobiogeography, the fossil evidence suggests that the family might have originated in Laurasia (Western Europe), but the existence of a tooth older than the European taxa might indicate that the family might have originated in Gondwana (Brazil). Finally, regarding paleoecology, the most accepted hypothesis is that they were generalist predators of the margins of aquatic environments (i.e. riparian zone), and waders in shallow waters like modern herons and storks, and if necessary also resorting to terrestrial environments. They would be capable of alternating between resources and environments, in addition to sharing their habitats with theropods of the Abelisauridae and Carcharodontosauridae families and even with other spinosaurids, if the environmental conditions favored it. Keywords: Theropoda, Spinosauridae, Spinosaurinae, Baryonychinae, paleobiogeography, paleoecology.
... Spinosaurids probably fed on prey items considerably smaller than their own body size, maybe predominantly, but not exclusively, fish (e.g., Taquet, 1984;Milner, 1986, 1997;Sereno et al., 1998 Hone and Holtz, 2021). However, although the name-giving genus Spinosaurus was described more than 100 years ago (Stromer, 1915), the skull and the postcranial osteology of spinosaurids is still rather poorly known due to the fragmentary nature of most of the recovered material (e.g., Kellner and Campos, 1996;Taquet and Russel, 1998;Milner, 2003 Most spinosaurid specimens described so far have no or only very limited skull remains. The original material of Spinosaurus only included the anterior ends of the mandibles and a fragment of the maxilla (Stromer, 1915), and only few and fragmentary skull remains were referred to the clade up to the mid-1990s (Taquet, 1984;Buffetaut, 1989Buffetaut, , 1992. ...
... Premaxillae and braincase material are also known for the recently described Riparovenator milnerae and Ceratosuchops inferodios (Barker et al., 2021), and a snout and further isolated cranial elements, including a braincase, have been referred to Suchomimus tenerensis (Sereno et al., 1998;Hendrickx et al., 2016;Sereno et al., 2022), but the latter three taxa lack detailed osteological descriptions. Other specimens mainly include partial snouts (Kellner Campos, 1996;Taquet and Russell, 1998 The spinosaurid Irritator from the Araripe Basin of north-eastern Brazil was initially briefly described and assigned to Maniraptora by Martill et al. (1996). The authors examined the specimen with aid of computed tomography (CT), revealing that the upper jaw was artificially elongated, but, due to the technical limitations of CT devices at that time, little anatomical detail could be gathered from the scans. ...
... The authors examined the specimen with aid of computed tomography (CT), revealing that the upper jaw was artificially elongated, but, due to the technical limitations of CT devices at that time, little anatomical detail could be gathered from the scans. In the same year, Kellner (1996) suggested that Irritator represents a spinosaurid, which was later supported in a more detailed description of the specimen by Sues et al. (2002), after the skull had been more completely prepared. The spinosaurids Irritator and Angaturama limai, both from the Romualdo Member of the Santana Formation (as formerly considered, see below) of Brazil, were described within a period of one month (Martill et al., 1996;Kellner and Campos, 1996). ...
Although originally described almost three decades ago, the holotype of Irritator challengeri from the Lower Cretaceous Romualdo Formation of Brazil still represents the most complete spinosaurid skull known to science. Here, we present a detailed description of the skull of Irritator based on digital reconstructions from medical and micro computed tomography (μCT) data. Segmentation reveals the near-complete palatal complex and braincase, an unusual morphology of the retroarticular process, a large, ventrally inclined surangular shelf and the tooth replacement pattern. The digitally reconstructed skull anatomy indicates a robust dentition, a field of binocular vision in front of the skull with an inclined snout orientation, a relatively weak but fast bite, as well as laterally spreading and rotating lower jaw rami during jaw opening. We modified an existing phylogenetic matrix of Tetanurae to account for new observations on the morphology of Irritator and analysed this using parsimony and Bayesian methods. Results support Spinosauridae as members of Megalosauroidea and recover a monophyletic Carnosauria (Megalosauroidea + Allosauroidea). Parsimony analysis recovers Monolophosaurus nested within Megalosauroidea as sister taxon to spinosaurids, but this is not supported by the Bayesian analysis. Bayesian time-calibration and evolutionary rate analysis indicate that spinosaurid evolution happened fast, despite a long ghost lineage of at least 35 million years. High evolutionary rates over a prolonged time can explain the highly derived skull morphology of spinosaurids. This study provides an in-depth look into the evolution of spinosaurid skull anatomy and refines our understanding of these specialized Mesozoic predators.
... Although they have an origin, probably in the Jurassic (Milner 2003;Carrano et al. 2012;Barker et al. 2021) fossil remains are rare to absent during the Jurassic and Lower Cretaceous, and several occurrences of the clade refer to materials of Barremian and Aptian/Albian ages (Bertin 2010;Hone et al. 2010;Novas et al. 2013). Spinosaurids have a widespread distribution; fossils have been recorded in Africa, Europe, Asia, and South America (e.g., Kellner & Campos 1996;Charig & Milner 1997;Sereno et al. 1998;Kellner et al. 2011;Allain et al. 2012;Novas et al. 2013;Barker et al. 2021;Mateus & Estraviz-López 2022) being absent only in North America, Oceania and Antarctica to date. ...
... However, most approaches recognize the monophyly of Baryonychinae, but with a species composition and valid taxa different from previous hypotheses (e.g., Sereno et al. 1998;Allain et al. 2012;Carrano et al. 2012;Malafaia et al. 2020;Barker et al. 2021;Lacerda et al. 2022;Mateus & Estraviz-López 2022). Regardless of this, even with the robustness of the traditional dichotomy of the group being questioned , the distinction of craniodental features, such as alveoli arrangement, tooth morphology, retracted external nares, and snout morphology, are quite evident and useful to distinguish genera from both clades (Kellner & Campos 1996;Charig & Milner 1997;Sereno et al. 1998;Milner Material and Methods ...
... Interestingly, the presence of more than one spinosaurid species from the Spinosaurinae and Baryonychinae, has not yet been ruled out in other Brazilian basins. Although, the co-occurrence of more than one spinosaurine has already been hypothesized considering the possible presence of both taxa of Albian from the Araripe region, Angaturama and Irritator (Kellner & Campos 1996;Sues et al. 2002), as well as the more than one dental morphotype of spinosaurines from the Cenomanian of Alcântara Formation (Medeiros & Schultz 2002;Medeiros 2006). Based on this, the contemporary occurrence of baryonychine and spinosaurine spinosaurids for the Lower Cretaceous of the Feliz Deserto Formation represents a novelty in this assemblage with ecological implications (e.g., competing medium to large predators). ...
We report the first occurrence in South America of spinosaurid specimens belonging to the clade Baryonychinae. The material comprises three tooth crowns recovered from the Lower Cretaceous (Berriasian–Valanginian) of the Feliz Deserto Formation, State of Sergipe, northeast Brazil. The three specimens have the typical conidont condition, displaying a fluted morphology, veined enamel texture and carinae extending to the cervix region, conditions similar to those shared among spinosaurid theropods. Furthermore, a notable feature is the presence of a serrated carinae composed of 6–7 diminutive denticles per millimetre, a key feature shared by baryonychine spinosaurids. Phylogenetic and multivariate analyses performed on dentition-based and crown-based datasets of theropod taxa confirm the relationship of the three tooth crowns described here with the well-supported spinosaurid clade. Our results retrieve the taxa which comprise the “classically” baryonychine clade as individual branches within a polytomy which includes the three tooth crowns and a subclade of spinosaurines (Spinosaurus + Angaturama/Irritator). Additionally, the discriminant and cluster analyses suggest that the materials are more similar to Suchomimus than to Baryonyx morphometrically, but clearly bearing similarities with baryonychines (sharing a similar morphospace). These new findings paleogeographically expand the occurrence of this clade to western Gondwana, and comprise one of the oldest spinosaurid records worldwide, which brings implications on hypotheses for the emergence and evolution of the clade. Furthermore, these new records indicate the presence of more than one spinosaurid taxon in the Lower Cretaceous deltaic paleoenvironment of the Feliz Deserto Formation.
... (1) elongated skull in the rostral region (hypertrophy of maxilla and premaxilla); (2) laterally compressed rostrum (oreinirostral morphology sensu Busbey 1995); (3) external nares posteriorly and laterally positioned; (4) conidont dentition (cone-shaped teeth) and (5) six or seven premaxillary teeth distributed in a 'terminal rosette' Milner 1986, 1997;Kellner and Campos 1996;Sereno et al. 1998;Dal Sasso et al. 2005;Hendrickx et al. 2015aHendrickx et al. , 2015bHone and Holtz 2017;Sales and Schultz 2017;Arden et al. 2019). The post-cranium also shows highly derived features that are observed in some species, such as: (i) hypertrophy of neural spines; (ii) robust forelimbs; (iii) hypertrophied ungual; (iv) flattened pedal ungual; (v) tail specialisations and (vi) high long bone compactness (Stromer 1915;Charig and Milner 1997;Sereno et al. 1998;Smith et al. 2006;Allain et al. 2012;Carrano et al. 2012;Ibrahim et al. 2014Ibrahim et al. , 2020Hendrickx et al. 2015a;Aureliano et al. 2018;Gasca et al. 2018;Maganuco and Dal Sasso 2018). ...
... For the ventral view, besides the main analysis, an additional exploratory analysis (AEA) was performed including three more specimens. Two poorly preserved specimens, NHMUK PV R 16420 and Angaturama limai USP GP/2T-5, were not included in the main analysis because they present a high degree of diagenetic alteration on the bone surface, affecting mainly the ventral border of the premaxillae (see Kellner and Campos 1996; and Sales and Schultz 2017 for comments on A. limai). FSAC-KK-7281 was not included because, due to ontogeny or taphonomy (Lakin and Longrich 2019), it does not have all structures that allow the acquisition of the total set of LMs (described in Table 2). ...
... Two articulated premaxillae from the Elrhaz Formation (Niger), illustrated and initially described as anterior portions of the mandible by Taquet (1984), are the first spinosaurid premaxillae to be published in the literature. The premaxillae were recognised as such by Kellner and Campos (1996) and used as the holotypic specimens of the new species Cristatusaurus lapparenti by Taquet and Russel (1998). The latter referred additional premaxillae and other fragmentary cranial and postcranial material to this taxon. ...
Spinosauridae presents an extensive geographical and temporal distribution, with records from Gondwana and Laurasia, and a temporal range from Barremian (~129 Ma, Lower Cretaceous) to Cenomanian (~95 Ma, Upper Cretaceous). To date, 13 species were described, besides several specimens identified at a broader taxonomic level. One of the most notable cranial features of spinosaurids is their elongated rostrum (hypertrophied premaxilla-maxilla). Leastwise five species possess preserved premaxillae: Angaturama limai, Baryonyx walkeri, Cristatusaurus lapparenti, Oxalaia quilombensis and Suchomimus tenerensis, besides materials tentatively attributed to Spinosaurus aegyptiacus. We studied the premaxillae shape of 10 specimens of the above-mentioned species and other materials through geometric morphometrics, reviewing diagnoses and morphological descriptions. Clear allometric and phylogenetic signals could be identified by ordination methods. We carried out a phylogenetic analysis to test spinosaurid relationships, by inclusion of new landmarks-characters from the premaxillae into a published tetanuran character-taxon matrix. The phylogenetic inference recovered C. lapparenti outside Baryonychinae, which was composed by B. walkeri and S. tenerensis. Spinosaurinae was recovered as (A. limai, (O. quilombensis, (MSNM V4047, Irritator challengeri))). Our results suggest that the premaxillae provide useful phylogenetic information and that the inclusion of landmarks-characters improves our knowledge of this enigmatic clade.
... Canudo et al., 2008), and comparatively little information has been published about South-American rep- resentatives. The South-American records are mostly restricted to the Cenomanian Alc^ antara Formation (S~ ao Luís-Grajaú Basin) (Medeiros & Schultz, 2002;Medeiros, 2006;Kellner et al., 2011) and the Aptian-Albian Romualdo Formation (Araripe Basin) (Kellner & Campos, 1996;Sues et al., 2002) of Northeast Brazil. However, there is also a single tooth attributed to Spinosauridae recently found in Feliz Deserto Formation, Sergipe-Alagoas Basin, also in NE Brazil, in BerriasianeValanginian strata ( , which would represent the oldest Spinosauridae record in South America. ...
... Nevertheless, it also yielded fossil plants and invertebrates (e.g., Coimbra et al., 2002;Lima et al., 2012;Pinheiro et al., 2014), besides crocodyliform, chelonian and pterosaur remains (e.g., Price, 1959;Wellnhofer, 1991;Oliveira & Kellner, 2007;Kellner et al., 2013). Dinosaur fossils are rare, and only theropod material has been recovered so far (Kellner & Campos, 1996;Bittencourt & Langer, 2011). The Romualdo Formation dinosaurs include: two Spinosaurinae, Irritator challengeri Martill et al., 1996 andAngaturama limai Kellner & Campos, 1996;an indeterminate coelurosaur, Santanaraptor placidus Kellner, 1999;a large compsognathid, Mirischia asymmetrica Naish et al., 2004; and a megaraptoran ( Rolando et al., 2017). ...
... Spinosaurinae fossils are unequivocally the most common dinosaur remains found in this geological context to date (Bittencourt & Kellner, 2010). Spinosauridae fossils include two partial skulls Kellner & Campos, 1996;Sues et al., 2002); a sacro-caudal vertebral sequence associated with chevrons (Bittencourt & Kellner, 2004); a rib tentatively assigned to the clade (Machado & Kellner, 2007); a pelvis, parts of anterior and posterior limbs, and further sacral and caudal vertebrae still undescribed Kellner & Campos, 1996;Kellner, 2001;Machado & Kellner, 2005;Machado, 2010). The two Romualdo Formation spinosaurids were erected on the recovered cranial material. ...
Spinosaurinae are known to have a strong relationship with aquatic environments, involving several anatomical adaptations. Nonetheless, this group of theropods remains enigmatic, due to the relative incompleteness of its fossil record. A large partial tibia from the Aptian-Albian Romualdo Formation, Northeast Brazil, is herein described through anatomical comparisons and paleohistological analyzes. It features characteristics previously only observed in Spinosaurus aegyptiacus, which includes a reduced fibular crest and an osteosclerotic condition. The later, a character supported as correlated with semi-aquatic habits in many limbed vertebrates. The results presented here support high bone compactness being already present in Brazilian Spinosaurinae millions of years before the Moroccan Spinosaurus. Furthermore, histological analyses demonstrate the Romualdo Formation specimen was a young subadult still growing fast by the time of its death, and suggests Araripe Basin Spinosaurinae could have grown larger than previously thought. This work contributes to a better paleobiological and ecological understanding of South American spinosaurs, and helps fill a gap in the macroevolutionary comprehension of Spinosaurinae. Ultimately, it also contributes to further advancing the paleoecological characterization of the Romualdo Formation.
... Canudo et al., 2008), and comparatively little information has been published about South-American representatives. The South-American records are mostly restricted to the Cenomanian Alcântara Formation (São Luís-Grajaú Basin) (Medeiros & Schultz, 2002;Medeiros, 2006;Kellner et al., 2011) and the Aptian-Albian Romualdo Formation (Araripe Basin) (Kellner & Campos, 1996;Sues et al., 2002) of Northeast Brazil. However, there is also a single tooth attributed to Spinosauridae recently found in Feliz Deserto Formation, Sergipe-Alagoas Basin, also in NE Brazil, in BerriasianeValanginian strata , which would represent the oldest Spinosauridae record in South America. ...
... Nevertheless, it also yielded fossil plants and invertebrates (e.g., Coimbra et al., 2002;Lima et al., 2012;Pinheiro et al., 2014), besides crocodyliform, chelonian and pterosaur remains (e.g., Price, 1959;Wellnhofer, 1991;Oliveira & Kellner, 2007;Kellner et al., 2013). Dinosaur fossils are rare, and only theropod material has been recovered so far (Kellner & Campos, 1996;Bittencourt & Langer, 2011). The Romualdo Formation dinosaurs include: two Spinosaurinae, Irritator challengeri Martill et al., 1996 andAngaturama limai Kellner &Campos, 1996; an indeterminate coelurosaur, Santanaraptor placidus Kellner, 1999;a large compsognathid, Mirischia asymmetrica Naish et al., 2004; and a megaraptoran (Rolando et al., 2017). ...
... Dinosaur fossils are rare, and only theropod material has been recovered so far (Kellner & Campos, 1996;Bittencourt & Langer, 2011). The Romualdo Formation dinosaurs include: two Spinosaurinae, Irritator challengeri Martill et al., 1996 andAngaturama limai Kellner &Campos, 1996; an indeterminate coelurosaur, Santanaraptor placidus Kellner, 1999;a large compsognathid, Mirischia asymmetrica Naish et al., 2004; and a megaraptoran (Rolando et al., 2017). Spinosaurinae fossils are unequivocally the most common dinosaur remains found in this geological context to date (Bittencourt & Kellner, 2010). ...
Spinosaurinae are known to have a strong relationship with aquatic environments, involving several anatomical adaptations. Nonetheless, this group of theropods remains enigmatic, due to the relative incompleteness of its fossil record. A large partial tibia from the Aptian-Albian Romualdo Formation, Northeast Brazil, is herein described through anatomical comparisons and paleohistological analyzes. It features characteristics previously only observed in Spinosaurus aegyptiacus, which includes a reduced fibular crest and an osteosclerotic condition. The later, a character supported as correlated with semi-aquatic habits in many limbed vertebrates. The results presented here support high bone compactness being already present in Brazilian Spinosaurinae millions of years before the Moroccan Spinosaurus. Furthermore, histological analyses demonstrate the Romualdo Formation specimen was a young subadult still growing fast by the time of its death, and suggests Araripe Basin Spinosaurinae could have grown larger than previously thought. This work contributes to a better paleobiological and ecological understanding of South American spinosaurs, and helps fill a gap in the macroevolutionary comprehension of Spinosaurinae. Ultimately, it also contributes to further advancing the paleoecological characterization of the Romualdo Formation.
... However, since the 1980s, new remains have been reported from various parts of the world, providing insights on the anatomy of these theropods (e.g., [1,2,4,5,7,11,[15][16][17][18]). These records extended the geographic range of Spinosauridae to Europe, Asia, and Gondwanan regions other than Africa, such as South America and perhaps Australia, and its temporal distribution back into the Jurassic [2,16,[19][20][21][22][23]. Some authors have also suggested their possible presence in the Late Jurassic of North America [16,24,25]. ...
... Spinosaurid remains from Brazil have played an important role in discussions regarding this clade. Firstly, they comprised the first unequivocal record out of Africa and Europe [19,26], despite putative reports from Asia by that time [27]. Secondly, the holotype of Irritator challengeri Martill et al., 1996, from the Romualdo Formation (Albian, Araripe Basin), is the spinosaurid specimen with the most complete preserved skull [15]. ...
... It may have come from Buxexé, a locality near Santana do Cariri Municipality, Ceará State [15,49]. On the other hand, the concretion enclosing Angaturama was considered as typically resembling those from the Romualdo Formation (Romualdo Member of the Santana Formation in Kellner and Campos [19]). Both cranial remains are distinct yet complementary to each other. ...
Fossil sites from Brazil have yielded specimens of spinosaurid theropods, among which the most informative include the cranial remains of Irritator, Angaturama, and Oxalaia. In this work some of their craniodental features are reinterpreted, providing new data for taxonomic and evolutionary issues concerning this particular clade of dinosaurs. The mesial-most tooth of the left maxilla of the holotype of Irritator is regarded as representing the third tooth position, which is also preserved in the holotype of Angaturama. Thus, both specimens cannot belong to the same individual, contrary to previous assumptions, although they could have been the same taxon. In addition, the position of the external nares of Irritator is more comparable to those of Baryonyx and Suchomimus instead of other spinosaurine spinosaurids. In fact, with regards to some craniodental features, Brazilian taxa represent intermediate conditions between Baryonychinae and Spinosaurinae. Such a scenario is corroborated by our cladistic results, which also leave open the possibility of the former subfamily being non-monophyletic. Furthermore, the differences between spinosaurids regarding the position and size of the external nares might be related to distinct feeding habits and degrees of reliance on olfaction. Other issues concerning the evolution and taxonomy of Spinosauridae require descriptions of additional material for their clarification.
... Other traits, like the degree of labiolingual compression and the distribution of flutes, can vary within the tooth row (Dal Sasso et al., 2005;Hendrickx and Mateus, 2014). Finally, although exhibiting flutes only on the lingual side, the sixth right premaxillary crown of the spinosaurid Angaturama limai is similar to LPUFS 5737 both in size and the general aspect of the flutes (Kellner, 1996;Kellner and Campos, 1996). ...
... Other traits, like the degree of labiolingual compression and the distribution of flutes, can vary within the tooth row (Dal Sasso et al., 2005;Hendrickx and Mateus, 2014). Finally, although exhibiting flutes only on the lingual side, the sixth right premaxillary crown of the spinosaurid Angaturama limai is similar to LPUFS 5737 both in size and the general aspect of the flutes (Kellner, 1996;Kellner and Campos, 1996). ...
... Given the BerriasianeValanginian age of the Feliz Deserto Formation, LPUFS 5737 represents a spinosaurid theropod in the pre-Aptian Early Cretaceous of northeastern Brazil, being the oldest occurrence of a South American spinosaur. Known and undisputed spinosaurid records from South America include three described taxa, Irritator challengeri, Angaturama limai, and Oxalaia quilombensis, along with post-cranial remains and isolated teeth, from the Albian and the AlbianeCenomanian of Araripe and São Luís-Grajaú basins, respectively ( Fig. 3; Kellner and Campos, 1996;Sues et al., 2002;Medeiros, 2006;Kellner, 2007, 2008;Kellner et al., 2011;Medeiros et al., 2014;Sales et al., 2016). ...
A new fossil site, called ‘Canafístula 01’, has yielded the first archosaur remains from the Berriasian–Valanginian Feliz Deserto Formation, Sergipe-Alagoas Basin, northeastern Brazil. Most of them comprise crocodylomorph teeth and osteoderms. However, the most remarkable specimen is a partial tooth assigned to Spinosauridae, based on the unique combination of the following features: (1) unfluted root almost as wide as the crown base, with a large pulp cavity; (2) straight and more regularly spaced flutes of the crown, formed by both the enamel and the dentine; and (3) unserrated carina on a mesiodistal plane coinciding with the main plane of curvature of the crown. This is the oldest occurrence of a spinosaurid from South America. In addition, given the unserrated distal carina, this tooth is referred to the subfamily Spinosaurinae, representing also the oldest spinosaurine record worldwide. It indicates that the cladogenetic event between Baryonychinae and Spinosaurinae must have occurred even before the Barremian, and that the latter was already present in South America in pre-Aptian ages. Thus, the occurrence of a spinosaurid in the Feliz Deserto Formation points to a latent potential for new relevant findings in northeastern Brazil and the necessity for greater collection efforts in this region.
... In Baryonyx (Charig and Milner, 1997) there are 6 teeth on the left and 7 on the right premaxilla. In MSNM V4047, alveolus 1 is much smaller than its counterpart in the Baryonychinae, whereas it is comparable to that of S. maroccanus and to that of Angaturama (Kellner and Campos, 1996); alveoli 2 and 3 are the largest; alveoli 4 and 5 are coupled and separated from the other premaxillary teeth by two short diastemata. A third larger, asymmetrical diastema (up to 76 mm on the right side) is present between alveolus 6 and the maxillary teeth. ...
... The preserved portions of the tooth crowns (left 3 and right 3) closely resemble the teeth previously assigned to Spinosaurus (Stromer, 1915;Bouaziz et al., 1988), both being nearly straight, elongate, conical, and sub-circular in transverse section. In lateral view, the dentigerous margin of the premaxilla resembles S. maroccanus (Taquet and Russell, 1998) in being strongly downturned towards the tip, such that the front of the rostrum is not elevated above the line of the maxillary tooth row, as in Baryonyx (Charig and Milner, 1997), Suchomimus (Sereno et al., 1998), and Angaturama (Kellner and Campos, 1996). In the diastema region, the rostrum has a sub-oval cross section with minimum circumference of 303 mm; the dentigerous margin is smoothly concave and, at scale, seems to fit on the convex dorsal margin of the rostral portion of the dentary figured by Stromer (1915:taf I, fig. ...
... According to several authors (Charig and Milner, 1997;Sereno et al., 1998;Buffetaut and Ouaja, 2002;Sues et al., 2002), Angaturama limai is a junior synonym of Irritator challengeri, as the two holotypes in all likelihood pertain to the same taxon (Fig. 4c, d). For this reason, information about the rostrum in Irritator is here taken from Angaturama (Kellner and Campos, 1996). ...
New specimens of the unusual theropod Spinosaurus cf. S. aegyptiacus from the Late Cretaceous (early Cenomanian) of Morocco reveal new information about the structure of the snout and the very large adult body size attained by the species. The external naris is retracted farther caudally on the snout than in other spinosaurids and is bordered exclusively by the maxilla and nasal. The fused nasals preserve a longitudinal, fluted crest. The size of the snout suggests that Spinosaurus may well have exceeded the maximum adult body size of other large Cretaceous theropods such as Tyrannosaurus and Giganotosaurus. The new material also supports the monophyly of the Spinosaurinae and the separation of Spinosaurus and Irritator.
... The main dinosaur-bearing localities come from Lower Cretaceous deposits, especially in the Northeast region of the country (Figure 1; see also Supplementary Data). The dinosaur components are mainly represented by rebbachisaurids ) and titanosaurian sauropods (Zaher et al. 2011;Ghilardi et al. 2016;Carvalho et al. 2017), associated with abelisauroid (Zaher et al. 2020;De Souza et al. 2021), spinosaurid (Kellner and Campos 1996;Sues et al. 2002, França et al. 2022Lacerda et al. 2023), carcharodontosaurid (Medeiros et al. 2014Pereira et al. 2020), megaraptoran (Aranciaga-Rolando et al. 2018), and coelurosaurian theropods (Kellner 1999;Naish et al. 2004;Sayão et al. 2020), including avialans (Carvalho et al. 2015. ...
... Prolific spinosaurine remains come from the Aptian record of the Araripe Basin (e.g. Kellner and Campos 1996;Martill et al. 1996;Sues et al. 2002;Bittencourt and Kellner 2004;Machado and Kellner 2007;Aureliano et al. 2018), from the Albian-Cenomanian São Luis-Grajaú Basin (Medeiros and Schultz 2002;Medeiros 2006;Kellner et al. 2011;França et al. 2022), and recently by a Spinosauridae tooth from the Albian-Cenomanian of the Potiguar Basin (Pereira et al. 2020). A dubious tooth was from the Cenomanian-Turonian of Argentina (Canudo et al. 2004), which potentially comprises the southernmost record of a spinosaurid in South America. ...
Supposed dinosaur remains were collected between 1859 and 1906 in the Lower Cretaceous Recôncavo Basin (Northeast Brazil). Since these materials remained undescribed, and most were considered lost. Recently, some of these historical specimens were rediscovered in the Natural History Museum of London, providing an opportunity to revisit them after 160 years. The specimens come from five different sites, corresponding to the Massacará (Berriasian-Barremian) and Ilhas (Valanginian-Barremian) groups. Identified bones comprise mainly isolated vertebral centra from ornithopods, sauropods, and theropods. Appendicular remains include a theropod pedal phalanx, humerus, and distal half of a left femur with elasmarian affinities. Despite their fragmentary nature, these specimens represent the earliest dinosaur bones discovered in South America, enhancing our understanding of the Cretaceous dinosaur faunas in Northeast Brazil. The dinosaur assemblage in the Recôncavo Basin resembles coeval units in Northeast Brazil, such as the Rio do Peixe Basin, where ornithopods coexist with sauropods and theropods. This study confirms the presence of ornithischian dinosaurs in Brazil based on osteological evidence, expanding their biogeographic and temporal range before the continental rifting between South America and Africa. Additionally, these findings reinforce the fossiliferous potential of Cretaceous deposits in Bahia State, which have been underexplored since their initial discoveries.
... The fossil record of Spinosauridae is, in most cases, fragmentary, with only a few well-known taxa. Comparison with the spinosaurid fossil record is limited by the absence of material overlapping between Riojavenatrix and the European Ceratosuchops and Riparovenator, the African Cristatusaurus and Sigilmassasaurus, and the South American Angaturama Kellner andCampos, 1996, Irritator, andOxalaia Kellner et al., 2011. Nevertheless, the fossil remains of Riojavenatrix lacustris overlap with enough material of other African, Asian, and European spinosaurids (Charig and Milner 1997, Sereno et al. 1998, Allain et al. 2012, Ibrahim et al. 2014, Sánchez-Hernández and Benton 2014, Malafaia et al. 2018, 2020a, Mateus and Estraviz-López 2022 to allow us to make comparisons and evaluate the possible synonymy between them, especially between the herein described taxon and the other Iberian spinosaurids. ...
... The presence of different spinosaurid taxa in a restricted area and time interval has also been reported, for instance, in south-eastern England, with Ceratosuchops and Riparovenator from the Barremian Wessex Formation, and the large-sized indeterminate spinosaurid from the overlying upper Barremian Vectis Formation of the Wessex sub-basin, and Baryonyx from the Barremian Upper Weald Clay Formation of the Weald subbasin (Charig and Milner 1986, Barker et al. 2021. The same has been suggested in other Gondwanan localities, such as the Araripe Basin, which has yielded holotype specimens of the spinosaurines Angaturama and Irritator (Kellner and Campos 1996, Martill et al. 1996, Sales and Schultz 2017 but for the possible synonymy, see e.g. Charig and Milner 1997, Sereno et al. 1998, Buffetaut and Oujada 2002, Dal Sasso et al. 2005, and also in the Kem Kem beds (Evers et al. 2015), the Baharyia Oasis (Stromer 1934), and the Elrhaz Formation (Taquet and Russell 1998). ...
A new member of Spinosauridae from the Enciso Group (uppermost Barremian–lower Aptian) from Igea (La Rioja, Spain) is here erected on the
basis of axial, pelvic girdle, and hindlimb elements that exhibit a unique combination of characters. Riojavenatrix lacustris gen. et sp. nov. is one of
the latest Iberian and European spinosaurid taxa. It retains a triangular pubic boot, like the megalosaurids, and a medial condyle of the femur that
shows a transitional stage between the anteroposteriorly oriented long axis of non-spinosaurid theropods and the posteromedially oriented long
axis of Spinosauridae. The spinosaurid record of Iberia ranges from the late Hauterivian–early Barremian to the latest Barremian–early Aptian
so far, and both the oldest and the most recent evidence comes from the Cameros Basin, where spinosaurid remains are especially abundant in
the Barremian deposits. A review of the spinosaurid record has allowed us to dismiss the presence of the genus Baryonyx from Iberia; hence, only
Camarillasaurus, Iberospinus, Protathlitis, Riojavenatrix gen. nov., and Vallibonavenatrix are considered to be present in the Early Cretaceous of
Iberia. According to this study, Riojavenatrix is one of the youngest baryonychines in the fossil record.
... Additionally, one specimen (LPUFS 5864) has two marks of tooth replacement (Fig. 9D2), a pattern already observed in spinosaurids (Isasmendi et al., 2022;Lacerda et al., 2022). In general terms, all described teeth share the morphology of flutes and enamel texture similar to spinosaurids, such as Angaturama (Kellner and Campos, 1996), Spinosaurus (Hasegawa et al., 2010;Hendrickx et al., 2015) and the indeterminate spinosaurine from the same locality and previously described (Sales et al., 2017). In contrast to baryonychine spinosaurids such as Baryonyx (Charig and Milner, 1997) and Iberospinus (Mateus and Estraviz-L opez, 2022), the teeth presented here lack the denticles in the mesial and distal carinae, which is a discernible feature shared among spinosaurines (Sereno et al., 1998;Hendrickx et al., 2015;Alonso and Canudo, 2016). ...
... Similarly, the fossil record of spinosaurids in South America comes from several localities in northeastern Brazil (e.g., Kellner and Campos, 1996;Medeiros et al., 2014;Sales et al., 2017). Based on our observations, the materials described here can be attributed to Spinosaurinae, which is the only spinosaurid clade recorded in Brazil to date. ...
The Feliz Deserto Formation (Berriasian–Valanginian, Sergipe-Alagoas Basin, NE Brazil) preserved some of the earliest South American fossil records of the rifting stages which resulted in the Gondwana supercontinent break-up during the Lower Cretaceous. Recently, the first spinosaurid theropod record of this formation was described, based on a tooth recovered from Canafístula 01 fossil locality in Sergipe State. We add herein twenty-seven isolated specimens to the fossil record of the Lower Cretaceous Feliz Deserto Formation. The new material includes seven isolated spinosaurid theropod teeth of the spinosaurine clade, as well as an indeterminate theropod preungual pedal phalanx. In addition, we describe an isolated crocodyliform osteoderm, as well as eighteen isolated teeth, some of which were taxonomically identified in three distinct morphotypes of neosuchian crocodyliforms. These findings expand the Gondwanan fossil record of both spinosaurine theropods and neosuchian crocodyliforms. Despite the fragmented nature of the specimens, these new fossils allowed the characterization of their general taphonomic features with low fluvial transport of bioclast prior to the burial. The depositional paleoenvironment of the Canafístula 01 locality is compatible with the deltaic system unit, that characterizes part of the Feliz Deserto Formation during the Lower Cretaceous. These fossil findings exemplify the co-occurrence of spinosaurids and more than one taxon of crocodyliforms in the deltaic-lacustrine paleoenvironment represented by the Feliz Deserto Formation. These new occurrences reinforce the fossiliferous potential of the Canafístula 01 locality, especially related to the paleovertebrates from the Lower Cretaceous of Brazil.
... Spinosaurids were particularly diverse during the Cretaceous (Bertin 2010) and share synapomorphies such as a pre-maxillary rosette (Charig and Milner 1997;Hone and Holtz 2017), external nostrils positioned posteriorly at the row of pre-maxillary teeth, well-elongated jaws, taller dorsal neural spines (twice as high as the centre) (Sereno et al. 1998;Hone and Holtz 2017), conical teeth with subcircular tooth crowns (Hone et al. 2010). Their fossils have been found in South America (Brazil), Africa (particularly North Africa), Europe (Portugal, Spain, England), Asia (China, Thailand, Laos), and Oceania (Australia) (e.g., Kellner and Campos 1996;Benton et al. 2000;Sues et al. 2002;Medeiros 2006;Buffetaut et al. 2008; Barrett et al. 2011;Bittencourt and Langer 2011;Kellner et al. 2011;Hendrickx et al. 2016;, with no occurrences in North America. In Brazil, spinosaurid remains include three species from the northeast: Angaturama limai (Kellner and Campos 1996) and Irritator challengeri (Martill et al. 1996; redescribed by Sues et al. 2002), both from the Romualdo Formation (Albian), as well as Oxalaia quilombensis (Kellner et al. 2011) from the Alcântara Formation (Cenomanian; Góes and Rossetti 2001). ...
... Their fossils have been found in South America (Brazil), Africa (particularly North Africa), Europe (Portugal, Spain, England), Asia (China, Thailand, Laos), and Oceania (Australia) (e.g., Kellner and Campos 1996;Benton et al. 2000;Sues et al. 2002;Medeiros 2006;Buffetaut et al. 2008; Barrett et al. 2011;Bittencourt and Langer 2011;Kellner et al. 2011;Hendrickx et al. 2016;, with no occurrences in North America. In Brazil, spinosaurid remains include three species from the northeast: Angaturama limai (Kellner and Campos 1996) and Irritator challengeri (Martill et al. 1996; redescribed by Sues et al. 2002), both from the Romualdo Formation (Albian), as well as Oxalaia quilombensis (Kellner et al. 2011) from the Alcântara Formation (Cenomanian; Góes and Rossetti 2001). performed a reinterpretation of spinosaurids' craniodental features and concluded that Angaturama and Irritator would fall between Baryonychinae and Spinosaurinae regarding their morphological similarities, with Oxalaia closer to other African Spinosaurinae. ...
Despite being poorly known from limited skeletal remains as teeth and a few postcranial material, spino-saurid remains have been reported from two different Lower Cretaceous localities in Brazil, with three officially proposed taxa up to now: Angaturama, Irritator and Oxalaia. Here, we report the first record of a spinosaurid pedal ungual from the Itapecuru Formation, Parnaíba Basin, Lower Cretaceous rocks of Maranhão State. The specimen retains a quite flat ventral surface that is proportionally almost two times broader than its proximal depth, which is a feature also found in Spinosaurus. Moreover, size inferences are made for this immature specimen.
... Stromer 1915) and South America (e.g. Kellner and Campos 1996;Sues et al. 2002;Bittencourt and Kellner 2004;Machado and Kellner 2005). ...
... Russell 1996;Dal Sasso et al. 2005;Ibrahim et al. 2014Ibrahim et al. , 2020aMaganuco and Dal Sasso 2018), and the Albian of both Algeria (e.g. Taquet and Russell 1998) and Tunisia (Buffetaut and Ouaja 2002); Irritator challengeri (including the specimen USP GP/2T-5 first described as Angaturama limai by Kellner and Campos 1996) from the Albian of Brazil (Sues et al. 2002;Sales and Schultz 2017); Oxalaia quilombensis from the Cenomanian of Brazil (Kellner et al. 2011); and Sigilmassasaurus brevicollis from the Cenomanian of Morocco (Russell 1996;McFeeters et al. 2013;Evers et al. 2015;Lakin andLongrich 2019see Ibrahim et al. (2020b) and Smyth et al. (2020) for a different interpretation regarding the validity of S. brevicollis). Besides, fragmentary specimens attributed to spinosaurids have been described in Valanginian (possibly extending to the Berriasian) levels of the Hastings beds of England (Fowler 2007), the upper Hauterivian-lower Aptian of Spain (e.g. ...
The fossil record of spinosaurid theropods from the Lower Cretaceous of the Iberian Peninsula is relatively abundant, but not very informative. This record, as generally occurs in other geographic areas, is mainly represented by isolated teeth, probably due to the distinctive features of the dentition of these theropods. However, cranial and postcranial elements attributed to spinosaurids have been described and reported from different Iberian fossil sites. The currently known record of these theropods spans from the upper Hauterivian to the lower Aptian of the Cameros and Maestrazgo basins in Eastern Spain whereas it is early Barremian in age in the Lusitanian Basin of Western Portugal. Spinosaurid remains from the Iberian fossil record were traditionally attributed to Baryonyx, firstly described in the upper Barremian–lower Aptian of United Kingdom. However, the recent description of a specimen collected in the locality of Santa Águeda (Vallibona, Castellón), from sediments of the Morella Formation, implies the presence of a new taxon with a combination of derived features more closely related with some Gondwanan spinosaurines than with Baryonyx. Therefore, the currently known Iberian spinosaurid record would be made up of at least three taxa: Baryonyx (or a closely related baryonychine), the spinosaurine Vallibonavenatrix cani, and possibly Camarillasaurus cirugedae, which was recently re-interpreted as belonging to a megalosauroid tentatively related with spinosaurids. This faunal diversity is also compatible with the assortment of spinosaurid tooth morphotypes (distinguished mostly by the presence or absence of serrated mesial carina) that has been described in the Lower Cretaceous of the Iberian Peninsula.
... Most Brazilian theropod nominal species and the best-known specimens come from the Lower Cretaceous Santana Formation in the northeast of the country (i.e., Irritator challengeri, Angaturama limai, Santanaraptor placidus, Mirischia asymmetrica; Kellner and Campos, 1996;Martill et al., 1996;Kellner, 1999;Sues et al., 2002;Naish et al., 2004;Carvalho et al., 2015). However, in recent years a more diverse Late Cretaceous theropod assemblage has been described in other geological units of the country, but still known from mostly fragmentary specimens, including abelisaurids (e.g., Pycnomemosaurus nevesi; Kellner and Campos, 2002;Delcourt, 2017;Delcourt and Grillo, 2017), spinosaurids (e.g., Oxalaia quilombensis; Kellner et al., 2011), probable carcharodontosaurids (Vilas Bôas et al., 1999;Candeiro et al., 2004Candeiro et al., , 2006, megaraptorans (M endez et al., 2012;Martinelli et al., 2013;Sales et al., 2017), and possibly noasaurids (Lindoso et al., 2012) and maniraptorans (Elias et al., 2004;Bittencourt and Langer, 2011). ...
... Despite being rare, all non-avian dinosaur specimens from the Araripe Basin have been discovered within the carbonate concretions of the Romualdo Member. Besides SMNS 58023, they include the holotype of the spinosaurids Irritator challengeri and Angaturama limai, the possible tyrannosauroid Santanaraptor placidus, and the possible compsognathid Mirischia asymmetrica, together with other postcranial remains of indeterminate theropods (Maisey, 1991;Kellner, , 1999Kellner and Campos, 1996;Martill et al., 1996Martill et al., , 2000Naish et al., 2004;Kellner, 2007, 2008). This non-avian dinosaur assemblage is unusual because consists only of theropods so far (Naish et al., 2004). ...
A specimen composed of a partial sacrum articulated to two anterior caudal vertebrae and an ilium (SMNS 58023) from the Lower Cretaceous (Albian) Santana Formation of Brazil was originally described as an oviraptorosaur, but it is here re-interpreted as the oldest megaraptoran of South America. The phylogenetic relationships of SMNS 58023 were tested quantitatively for the first time including it in the two most comprehensive phylogenetic data sets focused on non-maniraptoran theropods –including megaraptorans. The Brazilian specimen was consistently found as a megaraptoran in both analyses because of the presence of sacral centra longer than tall, absence of a median transverse constriction of sacral centra, and the morphology and position of sacral pleurocoels. SMNS 58023 sheds light on a region of the body that is poorly known in megaraptorans and pulls back the temporal range of the clade in South America. This re-interpretation reinforces the absence of oviraptorosaurs in Gondwana.
... REVISTA BRASILEIRA DE PALEONTOLOGIA, 18(3), 2015 Other groups with similar confi guration of teeth are the spinosaurids. For example, Irritator challengeri Martill, Cruickshank, Frey, Small, Clarke, 1996 and Angaturama limai Kellner & Campos, 1996 from the Early Cretaceous (Aptian-Albian) Romualdo Formation (see stratigraphic discussion in Kellner et al., 2013); shows slightly curved conical teeth with a rounded transverse section and longitudinal ridges ( Figure 3). However, Irritator and Angaturama bear fi ne and short enamel ''wrinkles'' (Kellner & Campos, 1996;Sues et al., 2002), as is the case of other spinosaurids (e.g. ...
... For example, Irritator challengeri Martill, Cruickshank, Frey, Small, Clarke, 1996 and Angaturama limai Kellner & Campos, 1996 from the Early Cretaceous (Aptian-Albian) Romualdo Formation (see stratigraphic discussion in Kellner et al., 2013); shows slightly curved conical teeth with a rounded transverse section and longitudinal ridges ( Figure 3). However, Irritator and Angaturama bear fi ne and short enamel ''wrinkles'' (Kellner & Campos, 1996;Sues et al., 2002), as is the case of other spinosaurids (e.g. Sereno et al.,1998;Kellner et al., 2011) that are not present in the Thoracosaurus bahiensis. ...
In the Recôncavo Basin, Bahia State, Samuel Allport and latter Charles F. Hartt collected several fossils, being some of them attributed to reptile teeth. Based on the morphotypes of these teeth Othniel C. Marsh nominated two species, the first with “delicately wrinkled surface” as Crocodylus hartti (currently Sarcosuchus hartti), and the second based on teeth with “strong continuous striae, and coarse riblets” as Thoracosaurus bahiensis, this latter will be the focus of this work. Marsh did not designate any holotype and, thus, the five teeth collected and described by Allport became the type series and subsequently the syntypes of the species. However, these are currently lost. Comparing the diagnoses of T. bahiensis with the type species of the genus and with others crocodyliformes as well as spinosaurids, we noted that the supposed autapomorphy of the species is highly variable. Therefore, we propose that T. bahiensis should be considered as nomen dubium.
... Los dientes de la Parreta presentan una sección mucho más redondeada que la presente en terópodos con dientes de similar tamaño como pueden ser alosaurios o abelisaurios. La forma cónica es un carácter derivado en Archosauria que aparece, convergentemente, en cocodrilos y en los terópodos Spinosauridae (Charig y Milner, 1997;Sereno et al., 1998), pudiendo ser una sinapomorfía de estos últimos (Kellner y Campos, 1996). ...
... Estrías longitudinales con patrones más irregulares que los encontrados en el material de la Parreta, también se han citado en dientes de terópodos enigmáticos como Paronychodon del Cretácico Superior de Norteamérica (Currie et al., 1990) y formas afines del Cretácico Inferior de la Península Ibérica (Zinke y Rauhut, 1994). Por otro lado, las formas con ornamentación consistente en crestas longitudinales del Cretácico Inferior son asignadas a Spinosauridae (Kellner y Campos, 1996;Martill y Hutt, 1996;, que está compuesto por dos clados hermanos: los Baryonychinae (los espinosáuridos más cercanos a Baryonyx que a Spinosaurus), y los Spinosaurinae (los espinosáuridos más cercanos a Spinosaurus que a Baryonyx). Los miembros de estos clados presentan dos tipos distintos de dientes. ...
In this work we study twenty teeth of spinosaurid theropod dinosaurs from the Early Aptian (Early Cretaceous) of Morella (Castellón, Spain). The teeth exhibit a mosaic of derived theropod characters such as ornamentation in longitudinal crests together with very small, uniformly sized denticles, which resemble baryonychine spinosaurids such as Baryonyx. However, they also display other derived characters not found in European Lower Cretaceous spinosaurids. Furthermore, the teeth share certain characters with Gondwanan taxa such as Suchomimus and Irritator.
... followed by other authors (CHARIG & MILNER, 1997;SERENO et al., 1998;SUES et al., 2002;TAQUET & RUSSEL, 1998). Angaturama limai, also a spinosaurid, is based on the anterior end of a skull (KELLNER & CAMPOS, 1996), and the much smaller tyrannoraptoran Santanaraptor placidus is based on an incomplete pelvis, caudal vertebrae, and hind limbs (KELLNER, 1999). Other theropod remains from the Romualdo Member include an undetermined sacrum with fragments of the right ilium (FREY & MARTILL, 1995), a partial skeleton of a small coelurosaurian theropod (MARTILL et al., 2000), and a large pelvis with segments of the vertebral column and limbs, figured (CAMPOS & KELLNER, 1991;KELLNER, 2001) but not described yet. ...
... The chevrons of MN 4743-V also lack those processes, what suggest that it possible represents a spinosaurid theropod. It should be noted that at the same deposit where MN 4743-V has yielded two spinosaurid taxa, both based solely on cranial material: Irritator challengeri (MARTILL et al., 1996) and Angaturama limai (KELLNER & CAMPOS, 1996 , 1995). This assignment was criticized by MAKOVICKY & SUES (1998), who argue that this character is present in several theropod taxa, and appears to have taken place several times in the Theropoda evolution (HOLTZ, 2000). ...
Besides being rare, most theropod remains from fossil deposits of Brazil are incomplete. Up to date the Romualdo Member (Aptian/Albian) of the Santana Formation yielded six theropod specimens. To those we add the description of a sequence of three posterior sacral and six anterior caudal vertebrae with three chevrons (MN 4743-V). Differences between MN 4743-V and members of the major theropod clades such as Ceratosauria, Allosauroidea and Coelorusauria do not allow its assignment to one of those groups. Instead, MN 4743-V is referred to the Spinosauroidea (Spinosauridae plus Torvosauridae), based on the presence of three robust laminae below the transverse process of the anterior caudals, which delimit three fossae. The lack of paired processes on the chevrons suggests that within Spinosauroidea MN 4743-V is a member of the Spinosauridae. This is congruent with previous findings of spinosaurids in the Romualdo Member. MN 4743-V differs from the remaining specimens that present sacrocaudal elements indicating the co-existence of at least five theropod species in this deposit.
... Dinosaur material is also rare (Figures 5a, b) and very few specimens were reported (Campos and Kellner, 1991; Kellner, 1996b). Two taxa were formally described: Irritator (Martill et al., 1996) and Angaturama (Kellner and Campos, 1996). Both were large animals representing Spinosauridae (Kellner, 1994; 1996b), a rare theropod clade previously recorded in the continental Cretaceous strata of Africa (see Kellner and Campos, 1996, for a review of spinosaurids). ...
... Two taxa were formally described: Irritator (Martill et al., 1996) and Angaturama (Kellner and Campos, 1996). Both were large animals representing Spinosauridae (Kellner, 1994; 1996b), a rare theropod clade previously recorded in the continental Cretaceous strata of Africa (see Kellner and Campos, 1996, for a review of spinosaurids). Small theropods are also found (Kellner, 1996a ) but were not formally described. ...
A review of the vertebrate fossil diversity in Brazil is presented. The best known faunas are the fish and reptiles from the Santana Formation (both, Crato and Romualdo Members). Also comparatively well known are the mammalian faunas from Pleistocene deposits, which is the result of extensive research done in the last decades. Poorly known are the Paleozoic vertebrates, which is possible due to the limited outcrops in the country. Paradoxically, the Late Cretaceous vertebrate faunas (fishes, reptiles, and mammals) from the Bauru Group are also poorly known, despite the large extension of these sedimentary rocks in the country. Following the achievements in the last decades, continuous systematic fieldwork has to be done, with the employment of various collecting techniques (e.g., screen washing), in order to improve the knowledge about the Brazilian paleovertebrate faunas.
... Moreover, the occurrence of these two tracksites confirms previous evidence for a geographic distribution of large theropod dinosaurs on the western margin of Gondwana ). In the absence of bones in the fossil record, these tracksites are the sole indication for the presence in this region of spinosaurids and carcharodonto− saurids, the potential trackmakers known in South America during the Cretaceous (e.g., Coria and Salgado 1995;Kellner and Campos 1996;Currie 2002, 2006;Candeiro et al. 2004;Novas et al. 2005;Medeiros 2006;Salgado et al. 2009). ...
... Large carnivorous dinosaurs in South America with sizes comparable to those of the Peruvian and Chilean track− makers, as estimated from the footprint length (hip height up to 3 m), pertain to spinosaurids and carcharodontosaurids, for which bone remains are known from the Aptian to the Santonian of Argentina and Brazil (e.g., Coria and Salgado 1995;Kellner and Campos 1996;Currie 2002, 2006;Candeiro et al. 2004;Novas et al. 2005;Medeiros 2006;Salgado et al. 2009). We should mention that known abelisauroids from the Early Cretaceous have feet smaller than 0.3 m and therefore could not have produced the large footprints recorded in the Querulpa Chico, Chacarilla and Calama tracksites (e.g., . ...
In modern terrestrial ecosystems, the population size of large predators is low, and a similar pattern has usually been assumed for dinosaurs. However, fossil finds of monospecific, large theropod accumulations suggest that population dynamics were more complex. Here, we report two Early Cretaceous tracksites dominated by large theropod footprints, in Querulpa Chico (Peru) and Chacarilla (Chile). The two sites correspond to distinct depositional environments-tidal basin/delta (Querulpa Chico) and meandering river (Chacarilla)-with both subject to extensive arid or semiarid palaeoclimatic conditions. Although most trackways show no preferred orientation, a clear relationship between two trackmakers is observed in one instance. This observation, coupled with the high abundance of trackways belonging to distinct large theropods, and the exclusion of tracks of other animals, suggests some degree of grouping behaviour. The presence of freshwater sources in a dry climate and perhaps social behaviour such as pair bonding may have promoted interactions between large carnivores. Further, the occurrence of these two tracksites confirms that large theropod dinosaurs, possibly spinosaurids and/or carcharodontosaurids, existed on the western margin of Gondwana as early as the earliest Cretaceous.
... Several new theropod dinosaurs from Brazil have been described in the last few decades and are helping to fill gaps in the evolution of ceratosaurs (Langer et al. 2019, Zaher et al. 2020, de Souza et al. 2021) and spinosaurs (Kellner and Campos 1996, Sues et al. 2002, Sales and Schultz 2017, Schade et al. 2023, for example. Other records and species, however, have been described based on very fragmentary materials (Delcourt and Iori 2020, Brum et al. 2021, Iori et al. 2021), but they are still important because they help to reveal part of the hidden diversity of dinosaurs from central South America. ...
In central South America, theropod remains are relatively scarce in comparison to the southern part of the continent, with shed teeth being the primary fossils found in this region. We examined 179 isolated teeth from the Bauru Basin, Brazil, using linear discriminant analysis (LDA; N = 178) and phylogenetic analysis (N = 174). The LDA used eight measurements, and the phylogenetic analysis used seven morphotypes. Although the results of the LDA suggest the presence of various South American theropod clades, such as Carcharodontosauridae, Noasauridae, and Spinosauridae, the phylogenetic analysis using a constrained dentition-based matrix classified the morphotypes as Abelisauridae (morphotypes I–IV, VI, and VII) and Therizinosauria (morphotype V). Given the considerable number of homoplastic characters and missing data, the phylogenetic analyses could not precisely determine the taxonomy of morphotypes V, VI, and VII in the unconstrained dataset. Morphological comparisons, nevertheless, strongly suggest that all morphotypes should be classified as abelisaurid theropods. We propose that the palaeogeographical distribution of Abelisauridae in South America was influenced by climatic conditions. These apex carnivores are likely to have adapted well to increased climate changes that led to semi-arid conditions. Our research sheds light on the evolutionary and ecological aspects of theropod dinosaurs in this region, contributing to a better understanding of the ancient ecosystems of central South America.
... The group named the animal Irritator challengeri, after the painstaking restoration work, which apparently caused them much irritation. The second group, composed of Brazilian palaeontologists, unaware of Irritator, were working on a similar fossil and named it Angaturama limai (Kellner & Campos, 1996). These turned out to be the same animal, with the British-German group having 'priority' on the name by only a few days, as outlined by the International Code of Zoological Nomenclature. ...
The fascination with ancient flora and fauna is a centuries-old phenomenon and is one of the main drivers of fossil crimes. Fossils have been sought by the rich and famous since the advent of colonialism, propelling development in the field of palaeontology but also in how fossils make their way through various, sometimes illegal, corridors to end up in museums and collections across the world. As demand for fossils increased in the nineteenth century, so did commercial avenues for these objects that were being sold not only to private individuals but also to scientific institutions. The most complete and visually impressive fossils in terms of size or uniqueness determine the market value of these fossils with some selling for millions of United States dollars. This has also led to the tampering or forging of fossils by many dealers in an attempt to inflate prices. While fossil crimes have been prevalent for centuries, the legal frameworks within which these crimes can be controlled or stopped are regularly challenged or even circumvented not only due to the lack of enforcements of these regulations but also due to how paleontological objects are classified in the first place as well as other legal loopholes.
... Os dinossauros não avianos da Formação Romualdo são representados por cinco espécies de Theropoda, incluindo os espinossaurídeos Irritator challengeri e Angaturama limai e os coelurossaurídeos Santanaraptor placidus, Mirischia assymetrica e Aratasaurus museunacionali, sendo apenas este último reconhecido nas concreções carbonáticas dos folhelhos negros na porção inferior da unidade, enquanto todos os demais foram identificados em rochas da porção superior. Aratasaurus museunacionali constitui o primeiro coelurossauriano basal reconhecido na Bacia do Araripe, sendo sugestivo de que este grupo era amplamente distribuído durante o Cretáceo Inicial (Kellner & Campos, 1996;Martill et al., 1996;Kellner, 1999;Aureliano et al., 2018;Sayão et al., 2020). Diferentes autores consideraram a possibilidade de os restos de Angaturama limai representarem a terminação anterior do rosto de Irritator challengeri (Sereno et al., 1998;Sues et al., 2002). ...
O rifteamento do Gondwana resultou em mudanças importantes na
composição química e distribuição de nutrientes dos oceanos, além de ter contribuído para modificações na circulação de massas de água, o que acarretou na abertura de novos espaços ecológicos. Com a reativação de sistemas de falhas do embasamento durante a ruptura, tem-se a formação de lagos de riftes perenes e temporários, por vezes alimentados por incursões fluviais. Além disso, os pulsos de incursões marinhas durante os eventos de transgressão no final do Cretáceo Inicial também geraram importantes modificações na biota da Bacia do Araripe. A gênese desses novos ecoespaços resultou em grandes transformações bióticas, criando novas possibilidades para o surgimento, a diversificação e a extinção de espécies em um curto espaço temporal. As rochas da Bacia do Araripe registram este momento em que ficam evidentes as relações intrínsecas entre as alterações dos espaços ecológicos e as consequentes biotas registradas em ambientes continentais e marinhos. A integração entre os dados sedimentológicos, estratigráficos e paleontológicos
das rochas da Bacia do Araripe permite categorizar três biotas, Missão Velha, Crato e Romualdo, produtos das dinâmicas tectônicas globais associadas à ruptura do Gondwana e suas decorrentes variações climáticas e oceanográficas. Essas biotas apresentam registros paleobiológicos singulares, com animais e vegetais abundantes, diversificados e muitas vezes de caráter endêmico. Muitos dos fósseis identificados são espécies com preservação
excepcional, que possibilitam o entendimento de aspectos-chave para a evolução biológica e a relação com as espécies viventes.
... Both species were published in the same month, only a few days apart. Irritator was described exclusively by non-Brazilian researchers, based on the posterior portion of a skull, which was acquired from fossil traders [30]; and Angaturama was described by Brazilian researchers, based on the anterior portion of a skull, acquired through a donation from a private collection to the Universidade de São Paulo [195]. The situation caused some authors to speculate that the fossils belonged to the same individual [196,197], which was, much later, discarded [198]. ...
Scientific practices stemming from colonialism, whereby middle- and low-income countries supply data for high-income countries and the contributions of local expertise are devalued, are still prevalent today in the field of palaeontology. In response to these unjust practices, countries such as Mexico and Brazil adopted protective laws and regulations during the twentieth century to preserve their palaeontological heritage. However, scientific colonialism is still reflected in many publications describing fossil specimens recovered from these countries. Here, we present examples of ‘palaeontological colonialism’ from publications on Jurassic–Cretaceous fossils from NE Mexico and NE Brazil spanning the last three decades. Common issues that we identified in these publications are the absence of both fieldwork and export permit declarations and the lack of local experts among authorships. In Mexico, access to many fossil specimens is restricted on account of these specimens being housed in private collections, whereas a high number of studies on Brazilian fossils are based on specimens illegally reposited in foreign collections, particularly in Germany and Japan. Finally, we outline and discuss the wider academic and social impacts of these research practices, and propose exhaustive recommendations to scientists, journals, museums, research institutions and government and funding agencies in order to overcome these practices.
... Whole dentition. Unserrated teeth are present in the whole dentition of: the noasaurid Limusaurus [Wang et al.'s (2017a) data matrix]; spinosaurines such as Irritator (Sues et al., 2002;Figure 11.3), Angaturama (Kellner and Campos, 1996), and Spinosaurus (Stromer, 1915); the neocoelurosaur Fukuivenator (Azuma et al., 2016); basal ornithomimosaurs such as Nqwebasaurus (Choiniere et al., 2012;Figure 11.6), Pelecanimimus (Pérez-Moreno et al., 1994), and Shenzhousaurus (Ji et al., 2003), and all alvarezsaurids such as Shuvuuia Figure 11.7) and Mononykus (Perle et al., 1993). Unserrated crowns are also present in the whole dentition of the basal oviraptorosaur Caudipteryx (IVPP 12430; NGMC 97 4 A; Figure 11.8), and all halszkaraptorine and unenlagiine dromaeosaurids such as Halszkaraptor , Mahakala (Turner et al., 2011), Buitreraptor (Gianechini et al., 2011b;Figure 11.9) and Austroraptor ). ...
Isolated theropod teeth are some of the most common fossils in the dinosaur fossil record and are continually reported in the literature. Recently developed quantitative methods have improved our ability to test the affinities of isolated teeth in a repeatable framework. But in most studies, teeth are diagnosed on qualitative characters. This can be problematic because the distribution of theropod dental characters is still poorly documented, and often restricted to one lineage. To help in the identification of isolated theropod teeth, and to more rigorously evaluate their taxonomic and phylogenetic potential, we evaluated dental features in two ways. We first analyzed the distribution of 34 qualitative dental characters in a broad sample of taxa. Functional properties for each dental feature were included to assess how functional similarity generates homoplasy. We then compiled a quantitative data matrix of 145 dental characters for 97 saurischian taxa. The latter was used to assess the degree of homoplasy of qualitative dental characters, address longstanding questions on the taxonomic and biostratigraphic value of theropod teeth, and explore the major evolutionary trends in the theropod dentition.
In smaller phylogenetic datasets for Theropoda, dental characters exhibit higher levels of homoplasy than non-dental characters, yet they still provide useful grouping information and optimize as local synapomorphies of smaller clades. In broader phylogenetic datasets, the degree of homoplasy displayed by dental and non-dental characters is not significantly different. Dental features on crown ornamentations, enamel texture, and tooth microstructure have significantly less homoplasy than other dental features and can be used to identify many theropod taxa to ‘family’ or ’sub-family’ level, and some taxa to genus or species. These features should, therefore, be a priority for investigations seeking to classify isolated teeth.
Our observations improve the taxonomic utility of theropod teeth and in some cases can help make isolated teeth useful as biostratigraphic markers. This proposed list of dental features in theropods should, therefore, facilitate future studies on the systematic paleontology of isolated teeth.
... Spinosaurids, a group of tetanuran theropod dinosaurs most distinctly characterized by their laterally compressed snouts and subconical crocodile-like teeth, have undergone a substantial surge in study and interest in recent years (Bertin, 2010). Skeletal remains attributed to this group have been described from the Cenomanian of Egypt (Stromer, 1915), the Aptian of Niger (Sereno et al., 1998), the Barremian of England (Charig and Milner, 1997), and the Albian and Cenomanian of South America (Kellner and Campos, 1996;Sues et al., 2002;Bittencourt and Kellner, 2004;Machado and Kellner, 2005). Aside from these records, spinosaurid material generally tends to be fragmentary, most frequently consisting of isolated teeth and bone fragments (Holtz et al., 2004;Bertin, 2010;Hone and Holtz, 2017). ...
A new medium-sized theropod dinosaur from the Arcillas de Morella Formation (upper Barremian) in Vallibona (Castellón, Spain) is described based on a partial skeleton. Vallibonavenatrix cani, gen. et sp. nov., is diagnosed by five autapomorphies: moderately high dorsal neural spines relative to the centrum height, the presence of deep pleurocoelous fossae and pneumatic foramina in the sacral vertebrae, a strongly pneumatic ilium with large internal cameras, the presence of a broad and flat platform on the ventromedial surface of the postacetabular blade of the ilium, and the strong ventral torsion of the ischium diaphysis. Phylogenetic analysis recovered Vallibonavenatrix cani as a spinosaurid megalosauroid and revealed that the new Iberian form is more closely related to Gondwanan spinosaurines, such as Spinosaurus, Irritator or Angaturama and the Asian taxon Ichthyovenator than it is to its synchronic contemporary European taxon Baryonyx walkeri. The Gondwanan or Asian affinities of Vallibonavenatrix indicate a complex palaeobiogeographic pattern and may be interpreted simultaneously as evidence for contact between Europe and North Africa and between Europe and Asia before the Aptian. This specimen is currently the most complete evidence of a spinosaurid theropod known in the fossil record of the Iberian Peninsula.
... The surface lacks ornamentation, unlike other theropods (i.e spinosaurids). The crowns of the maxillary teeth are mediolaterally compressed, with an elliptical cross-section, and slightly curved, as in most theropods except derived spinosaurids (Kellner and Campos, 1996;Serrano-Martínez et al., 2015). Maxillary and premaxillary teeth have crenulated carinae as most theropods. ...
Concavenator corcovatus is a carcharodontosaurid dinosaur represented by an almost complete and articulated skeleton from the ‘Las Hoyas’ fossil site (Lower Cretaceous, Spain). The skull of Concavenator is almost complete, missing only the anteriormost and posteriormost regions. Here, a review of the cranial anatomy of Concavenator has been carried out to test its phylogenetic relationships. This description provides a detailed revision of the cranial anatomy of carcharodontosaurids, which is relevant to their general phylogenetic relationships. The results show that the skull of Concavenator has several carcharodontosaurid synapomorphies such as: (1) a lacrimal-postorbital contact, (2) a well-developed postorbital boss, (3) anteroventrally oriented postorbital ventral process, (4) an intraorbital process in the postorbital, (5) rostrally projected roughness in the dorsal surface of the postorbital, (6) a notch in the ascending ramus of the maxilla, (7) a sulcus on the anterior margin of the lacrimal ventral ramus, (8) curved dorsal surface of the lacrimal, and (9) fused frontal-parietal contact. Finally, Concavenator has three cranial autapomorphies: (1) connection between the different recesses on the lateral surface of the nasal, (2) elongated and acuminate posterior narial fossa, and (3) a rounded morphology of the ventral surface of the postorbital boss. In addition, anatomical comparison has enabled to create a complete 3D reconstruction of the skull of Concavenator.
... Os peixes destacam-se por ser o grupo de macrofósseis que ocorrem em maior abundância, além de ter uma grande paleobiodiversidade, incluindo Chondrichthyes e Osteichthyes (Actinopterygii e Sarcopterygii), os táxons mais comuns são Vinctifer comptoni (Agassiz, 1841), Rhacolepis buccalis (Agassiz, 1841) e Tharrhias araripis (Jordan & Branner, 1908). Dentre os tetrápodes, há cinco espécies de quelônios, Araripemys barretoi (Price, 1973), Santanachelys gaffneyi (Hirayama, 1998), Brasilemys josai (Broin, 2000), Cearachelys placidoi (Gaffney et al., 2001), e Euraxemys essweini (Gaffney et al., 2006), os arcossauros compreendem os crocodilianos Araripesuchus gomesii (Price, 1959) e Caririsuchus camposi (Kellner, 1987), os dinossauros Angaturama limai (Kellner & Campos, 1996), Irritator challengeri (Martill et al., 1996), Santanaraptor placidus (Kellner, 1999) e Mirischia asymmetrica (Naish et al., 2004) Tharrhias, Notelops, Araripelepdotes, Brannerion, Beurlenichthys, celacantídeos, quelônios, sendo um plastrão e ossos articulados de Araripemys barretoi e uma carapaça completa de uma outra espécie, em processo de identificação, Pterosauria, incluindo fragmentos de ossos de membro anterior e, parte anterior de um rostro (ou mandíbula). ...
Here are new and old inventory of fossiliferous sites in the Romualdo Formation, Albian of the Araripe Basin, in the municipality of Exu, Pernambuco, Northeast of Brazil, with the survey of its fossiliferous diversity, with emphasis on paleovertebrates. Seven localities were studied, and 16 taxa of paleovertebrates were identified, including fishes: Vinctifer
comptoni, Rhacolepis buccalis, Calamopleurus cylindricus, Cladocyclus gardneri, Neoproscinetes penalvai, Paraelops cearenses, Tharrhias araripis, Notelops brama, Araripelepdotes temnurus, Brannerion sp. e Beurlenichthys ouricuriensis; new occurrences of pterosaurs Anhangueridae and; chelonians (Araripemys barretoi). The Romualdo Formation has been studied since 1800 and it is internationally recognized as Fossillagerstätte. However, the Pernambuco region of the Araripe Basin lacks systematic studies when compared to the region of Ceará, which indicates the need to register the
occurrence of fossiliferous sites and diversity, contributing to the knowledge of vertebrate paleontology in the southwest portion of the Araripe Basin and expanding the paleontological collections of the state of Pernambuco.
... Baryonychine teeth have serrated carinae with minute denticles, more labiolingually compressed teeth than spinosaurines and a more pronounced distal curvature of the crown (Charig and Milner 1997;Canudo et al. 2008a;Alonso and Canudo 2016). Also, spinosaurine teeth have unserrated carinae, as exemplified by Irritator, Angaturama and Spinosaurus (Stromer, 1915;Kellner and Campos 1996;Sues et al. 2002). In general, all these differences are plesiomorphies in baryonychine teeth, or apomorphies in spinosaurine teeth. ...
Introduction: Barranco del Hocino-1 is a new fossil site located near Estercuel, Teruel province, Spain. The fossil site is located geologically within the Oliete sub-basin, in the Blesa Formation (Barremian in age). Barranco del Hocino-1 shows a diverse assemblage of tetrapod vertebrates similar to other sites in the Blesa Formation. Materials and methods: Six isolated teeth belonging to Theropoda have been found. A study of their qualitative and quantitative characters, along with statistical (DFA) and cladistic analyses, enable us to identify four different dental morphotypes. Results: These morphotypes belong to separate tetanuran theropod taxa. One is related to Spinosauridae. The other morphotypes show affinities with non-spinosaurid tetanurans, probably related to Carcharodontosauria. Conclusions: The results are congruent with the known theropod record of the Iberian Peninsula and western Europe. This work is a new contribution to what is known of the palaeobiodiversity and distribution of large-bodied theropods from the Barremian of the Iberian Peninsula. © 2018, Springer International Publishing AG, part of Springer Nature.
... Spinosaurids -the group of long-snouted and possibly semiaquatic theropods -are one of the most distinctive dinosaur groups. Fossil remains of these predators are known from many localities from the Lower Cretaceous of Africa (Algeria, Egypt, Niger, Sudan, Tanzania, Tunisia), Europe (England, Portugal, Spain), South America (Brazil), Asia (China, Laos, Thailand), and Australia (e.g., Kellner & Campos, 1996;Sereno et al., 1998;Benton et al., 2000;Buffetaut & Ouaja, 2002;Sues et al., 2002;Medeiros, 2006;Buffetaut, 2008Buffetaut, , 2012Buffetaut et al., 2008;Hone et al., 2010;Barrett et al., 2011;Kellner et al., 2011;Allain et al., 2012;Medeiros et al., 2014;Hendrickx et al., 2016;Sales et al., 2017). Knowledge about spinosaurids has increased significantly in the last few decades as new fossil remains, including various new species, have been found in several new Gondwanan and Laurasian localities (Buffetaut & Ingavat, 1986;Sereno et al., 1998;Benson et al., 2009;Buffetaut, 2012;Allain et al., 2012;Hendrickx et al., 2016). ...
We review the fossil records of spinosaurid dinosaurs in order to discuss this group’s evolution and distribution
in Europe and North Africa during the Early Cretaceous. Along with their eastern Laurasian distribution during the
Cretaceous, these theropods have been found in coastal deposits of Europe and North Africa dated from the Barremian
to the Cenomanian. The main occurrences of spinosaurid remains are in the deposits of northern Gondwana and western
Laurasia, which suggests that these regions were very important in spinosaurid evolution prior to the Cenomanian. Later,
spinosaurids were seemingly replaced in northern Gondwana by other top predator groups, including the abelisauroids.
... Baryonychine teeth have an ornamentation based on ridges, a cross-section that is not as subcircular as in spinosaurine teeth, and a more pronounced distal curvature of the crown Alonso and Canudo 2016). Spinosaurine teeth also tend to have unserrated carinae, as in Irritator, Angaturama and Spinosaurus (Stromer 1915;Kellner and Campos 1996;Sues et al. 2002;Hendrickx 2015). In addition, the sporadic variation in size of denticles has been proposed as a feature of Baryonychinae (Mateus et al. 2011). ...
Introduction
The relationship between dinosaur carcasses and isolated theropod teeth is well known in the fossil record. The usual explanation is that theropod dinosaurs fed on the herbivore carcass. Other evidence is provided by theropod tooth marks on the surface of herbivore skeletal remains. In this work we study isolated theropod teeth and the tooth marks in evidence on the bones of the sauropod from El Oterillo II (Salas de los Infantes, Spain, Lower Cretaceous). Theropod and crocodylomorph teeth have been found in relationship with the sauropod carcass. The fossils lie on channel lag deposits composed of sandstones with quartzite gravel.
Materials and methods
Thirty theropod teeth have been studied using qualitative features, statistical and cladistics analysis in addition to the tooth marks present on the vertebrae.
Results
The morphology of the theropod teeth has revealed greater palaeobiodiversity in these faunas than previously known, including baryonychine spinosaurids, basal tetanurans, dromaeosaurids and a singular coelurosaurian. The presence of tooth marks and isolated theropod teeth in close relationship with the sauropod carcass could also provide new evidence of the scavenging of theropod dinosaurs on the sauropods of this age and location.
Conclusions
Six morphotypes of theropod teeth have been distinguished; the combination of basal and derived tetanurans is congruent with the known record from the Early Cretaceous of the Iberian Peninsula. The most probable explanation for the association of isolated theropod teeth and sauropod remains is the scavenging of the carcass by theropod dinosaurs.
... Baryonychine teeth have an ornamentation based on ridges, a cross-section that is not as subcircular as in spinosaurine teeth, and a more pronounced distal curvature of the crown Alonso and Canudo 2016). Spinosaurine teeth also tend to have unserrated carinae, as in Irritator, Angaturama and Spinosaurus (Stromer 1915;Kellner and Campos 1996;Sues et al. 2002;Hendrickx 2015). In addition, the sporadic variation in size of denticles has been proposed as a feature of Baryonychinae (Mateus et al. 2011). ...
... [Kellner etCampos, 1996]Carcharodontosauridae ...
Le plateau des Hamadas, situé au sud du Haut Atlas marocain, contient de nombreux
fossiles de vertébrés qui constituent une succession de faunes provenant d’environnements divers
sur une période de près de 10 millions d’années à la base du Crétacé supérieur. La base de la
série crétacée est constituée des formations Ifezouane et Aoufous, correspondant à des plaines
alluviales et des deltas pour la première, et à des milieux paraliques et de sebkhas pour la seconde.
Les fossiles de l’assemblage composite des « Kem Kem beds » se trouvent le long d’une falaise de
250 km située le long de la frontière algérienne et au nord-est de la ville d’Erfoud. Les origines
géographique et stratigraphique de nombreuses découvertes scientifiquement importantes
sont imprécises car les fossiles sont souvent découverts par les personnes de la région qui les
exploitent dans un but commercial. La majorité des spécimens semblent provenir du sommet
de la formation Ifezouane. Cet assemblage, comprenant des requins, des poissons osseux, des
amphibiens, des squamates, des tortues, des crocodiles, des dinosaures sauropodes et théropodes
ainsi que des ptérosaures, fait partie d’un ensemble de faunes continentales rencontrées dans
une grande partie de l’Afrique du Nord dans la série nommée « Continental intercalaire ». Les
différents assemblages du « Continental intercalaire » sont d’âges variables, s’échelonnant de la
base du Crétacé inférieur (peut-être même du Jurassique) jusqu’à la base du Crétacé supérieur.
L’assemblage composite des « Kem Kem beds » peut être rapporté au Cénomanien inférieur sur
la base de similitudes taxonomiques avec la faune bien datée de Bahariya en Égypte. Plusieurs
traits écologiques caractérisent cet ensemble : 1) une abondance de taxons, en particulier chez les
poissons et les dinosaures, représentés par des individus de grande taille ; 2) la présence quasi exclusive de dinosaures saurischiens et 3) la surreprésentation des restes de carnivores par rapport
aux herbivores.
Au-dessus de la formation Ifezouane, la formation Aoufous est surmontée de la formation
Akrabou, constituée de sédiments marins déposés lors de la grande transgression du Cénomano-
Turonien. La faune d’Agoult, située près de cette limite cénomano-turonienne sur le lieu même où
Lavocat découvrit des restes de dinosaures dans les années 1950 (Gara Sbâa), contient des poissons
qui vivaient dans un environnement marin restreint, calme et peu profond. Plus au nord, dans la
région de Goulmima, un assemblage de poissons, essentiellement des téléostéens, et de reptiles
marins (mosasauroïdes, polycotylidés, elasmosauridés, protostegidés) est connu dans les niveaux
du Turonien inférieur.
D’un point de vue palaéobiogéographique, l’assemblage composite des « Kem Kem beds »
est proche d’autres faunes d’Afrique du nord et du nord-est du Brésil, indiquant l’existence
d’écosystèmes relativement homogènes s’étendant sur de gigantesques régions. La faune d’Agoult
montre aussi de fortes affinités avec les faunes de la Téthys centrale et du Nord du Mexique, alors
que l’assemblage de Goulmima est proche des faunes de la Téthys occidentale et de la Western
Interior seaway et, dans une moindre mesure, de l’Atlantique Sud.
... Sob a Chapada do Araripe pode ser encontrada a sequência completa das unidades geológicas que compõem a bacia. É delimitada ao Norte pelo lineamento da Paraíba, a noroeste pela falha de Farias Brito, ao Sul pela Falha de Sítio das Moreiras e a leste pela falha de Conceição (CASTRO; CASTELO-BRANCO, 1999 RICHTER, 1998;MAISEY, 1994;WILBY;BRIGGS, 1997;MARTILL;WASHINGTON-EVANS, 2008), dinossauros terópodes (KELLNER;CAMPOS, 1996;2000;BITTENCOURT;KELLNER, 2004;MACHADO;, pterossauros (PRICE, 1971;BARRETT et al., 2008;KELLNER et al., 2013;VILA NOVA et al., 2014), crocodilomorfos (PRICE, 1959KELLNER, 1987;HECHT, 1991) ...
Uma nova localidade fossilifera, Sitio Baixa Grande, e descrita para a Formacao Romualdo, Cretaceo Inferior do Nordeste do Brasil. Localizada proxima da area urbana da cidade de Potengi, na porcao oeste da Bacia do Araripe, esse sitio foi descoberto em 2010 e apresenta diversidade e grande quantidade de fosseis. As concrecoes, de coloracao bege a avermelhada, com formas regular e irregular, sao facilmente encontradas sobre o solo. Apresentam forma ovoide, com textura pouco laminada. Os macrofosseis estao localizados na porcao mediana da concrecao, podendo apresentar tambem tridimensionalidade, uma feicao comum aos fosseis da Formacao Romualdo. Dentre os fosseis coletados estao tetrapodes, representados por ossos isolados de pterossauros e restos de tartarugas nao-Araripemidedae. Os peixes encontrados incluem: Vinctifer comptoni, Cladocyclus gardineri, Tharrhias araripis, Calamopleurus cilindricus e Neoproscinetes penalvai . Os fosseis aqui apresentados compreendem os unicos com localidade de coleta entre os encontrados na regiao oeste da Bacia do Araripe, uma area onde os afloramentos fossiliferos do Grupo Santana foram considerados como inexistentes. Dessa forma contribuindo para o conhecimento da distribuicao de niveis fossiliferos dessa importante feicao tectonica.
... The carbonate concretions of the Romualdo Member of the Santana Formation, provided a record of at least four non-avian theropods, with the possibility of a fifth (Machado & Kellner, 2007), interpreted as a rib of an unknown theropod. Only theropods were found in this unit so the dinosaur fauna of the Araripe Basin consists of two spinosaurids, Irritator challengeri (Martill et al., 1996) and its possible synonym, Angaturama limai (Kellner & Campos, 1996); and two coelurosaurs, Santanaraptor placidus (Kellner, 1999) and Mirischia asymmetrica (Naish, Martill & Frey, 2004). The latter two likely had some feathery integument (Ji & Ji, 1996; Chen, Dong & Zhen, 1998; Ji et al., 2007). ...
Here we describe three fossil feathers from the Early Cretaceous Santana Formation of the Araripe Basin, Brazil. Feathers are the most complex multiform vertebrate integuments; they perform different functions, occurring in both avian and non-avian dinosaurs. Despite their rarity, fossil feathers have been found across the world. Most of the Brazilian feather fossil record comes from the Santana Formation. This formation is composed of two members: Crato (lake) and Romualdo (lagoon); both of which are predominantly reduced deposits, precluding bottom dwelling organisms, resulting in exceptional preservation of the fossils. Despite arid and hot conditions during the Cretaceous, life teemed in the adjacency of this paleolake. Feathered non-avian dinosaurs have not yet been described from the Crato Member, even though there are suggestions of their presence in nearby basins. Our description of the three feathers from the Crato laminated limestone reveals that, despite the small sample size, they can be referred to coelurosaurian theropods. Moreover, based on comparisons with extant feather morphotypes they can be identified as one contour feather and two downy feathers. Despite their rareness and low taxonomic potential, fossilized feathers can offer insights about the paleobiology of its owners and the paleoecology of the Araripe Basin.
... Despite the considerable number of Lower Cretaceous continental sedimentary deposits in Brazil, body fossil remains of Early Cretaceous dinosaurs have only been formally documented in the AptianeAlbian Araripe and São Luís-Grajaú basins (Kellner and Campos, 1996;Martill et al., 1996;Kellner, 1999;Medeiros and Schultz, 2002;Sues et al., 2002;Carvalho et al., 2003;Naish et al., 2004;Medeiros, 2006;Castro et al., 2007;Bittencourt and Langer, 2011;Kellner et al, 2011;Lindoso et al., 2012;Faria et al., 2015). Curiously though, dinosaur footprints from the Lower Cretaceous of Brazil are pervasive (e.g. ...
Here we describe a new record of a sauropod dinosaur from the Lower Cretaceous (Hauterivian-Barremian) Rio Piranhas Formation, Sousa Basin, NE Brazil. Dinosaur fossil bones from this deposit were
unknown until now. Thus, the discovery of a sauropod fibula from this locality is highly significant. Our
discovery represents an indeterminate titanosaur and the earliest stratigraphic occurrence of this group
in central Gondwana. When compared to chronocorrelate titanosaur trackmakers of this geological unit,
this fossil specimen appears substantially smaller. Histological analysis of the fibula suggests that this is a
relatively young individual (approximately 40-50% adult body size) that had passed its most rapid phase
of early juvenile growth, but had not yet attained somatic maturity. Thus, the fibula recovered is from a
young individual rather than from a small-bodied adult titanosaur.
... ''Outward'' and ''inward'' facing lateral crown surfaces have been described as either being external and internal or labial and lingual in theropods (Dong, 1997a); as lateral and internal in sauropods (Dong, 1997b); as labial and lingual in artiodactyls (Lucas and Emry, 1999), cervids (Azanza and Montoya, 1995), insectivores (X. Wang and Zhai, 1995), marsupials (Cifelli and de Muizon, 1998), armadillos (Vizcaíno and Bargo, 1998), feliforms (Albright, 1996), theropods (Hutt et al., 1996;Kellner and Campos, 1996;Charig and Milner, 1997), ornithischians (Dong, 1997c), sauropods (Upchurch, 1999), and osteichthyians (Kemp, 1997). These same surfaces have been described as buccal and lingual in marsupials (Muirhead and Filan, 1995;Kappelman et al., 1996;Wroe, 1996;Sánchez-Villagra and Kay, 1997), theropods (Kirkland et al., 1993), mosasaurs (Lingham-Soliar, 1998, hadrosaurids (Head, 1998), and ursids (Stiner et al., 1998); as labial and medial in ceratopsian dinosaurs (Dong and Azuma, 1997); and as medial and lateral in carnivores (Biknevicius et al., 1996), alligatoroids (Williamson, 1996), ornithischians (Hunt and Lucas, 1994;Galton, 1995Galton, , 1996Russell and Zhao, 1996;Xu, 1997), sauropods (Sereno et al., 1994), and theropods (Sereno et al., 1996). ...
There is little consistency in the notation and orientation terminology used in discussions of non-mammalian fossil vertebrate dentitions. The standardization of this terminology, as done in the medical and dental sciences, would facilitate all future research on fossil teeth. For mammals, we recommend following convention, where incisors, canines, premolars, and molars are abbreviated as In, Cn, Pn, and Mn (n = tooth number) in upper jaws and as in, cn, pn, and mn in lower jaws. Right, left, and deciduous teeth are indicated by R, L, and D (e.g., DP4, Rp2). For non-mammals, which can have dentigerous premaxillae, maxillae, and dentaries, as well as additional tooth-bearing bones (e.g., vomers, palatines, pterygoids, ectopterygoids, sphenoids, splenials, and even parasphenoids), we encourage identifying teeth using the bone abbreviation (e.g., pmn, mxn, dn, vn, paln). A number and slash (/) combination can be used to distinguish between multiple tooth rows (e.g., Pal1/n, Pal2/n), and specimen-specific maps can be created for very complicated dentitions. We suggest the use of the terms mesial and distal to designate tooth surfaces and directions facing toward and away from the mandibular symphysis. Labial is offered for those surfaces and directions facing the lips or cheeks and lingual for those facing the tongue. We offer the terms basal for the direction toward crown bases, apical for the direction toward crown tips, occlusal for views of the occlusal surfaces, and basal and root apical for views of crown bases and roots, respectively.
... Hone et al. [48] also commented that a taphonomic bias in favor of spinosaurid remains in the fossil record in comparison to other theropod taxa should be expected, as the former might have inhabited preferentially aquatic habitats. However, the rareness of spinosaurid specimens would suggest the contrary, and this could correlate with the rareness of these theropods in the fossil record, although the most abundant dinosaur remains in the Romualdo Formation are those identified as spinosaurids [59][60][61][62][63][64]. Besides, they also suggest that spinosaurids should have been present also in North America (see also [65]) and their absence in the North American fossil record should be considered as evidence further corroborating the hypothesis of their scarcity in paleoecosystems. ...
The view of spinosaurs as dinosaurs of semi-aquatic habits and strongly associated with marginal and coastal habitats are deeply rooted in both scientific and popular knowledge, but it was never statistically tested. Inspired by a previous analysis of other dinosaur clades and major paleoenvironmental categories, here we present our own statistical evaluation of the association between coastal and terrestrial paleoenvironments and spinosaurids, along with other two theropod taxa: abelisaurids and carcharodontosaurids. We also included a taphonomic perspective and classified the occurrences in categories related to potential biases in order to better address our interpretations. Our main results can be summarized as follows: 1) the taxon with the largest amount of statistical evidence showing it positively associated to coastal paleoenvironments is Spinosauridae; 2) abelisaurids and carcharodontosaurids had more statistical evidence showing them positively associated with terrestrial paleoenvironments; 3) it is likely that spinosaurids also occupied spatially inland areas in a way somehow comparable at least to carcharodontosaurids; 4) abelisaurids may have been more common than the other two taxa in inland habitats.
... Spinosaurid teeth seem to be already present in the Kimmeridgian-Tithonian of Tanzania (Buffetaut 2011; but for a different opinion see Rauhut 2011), yet the earliest definitive spinosaurid is currently Baryonyx walkeri ( Figure 9C) from the Barremian of England and Portugal (Charig & Milner 1986Mateus et al. 2011). Spinosauridae are also known from the Aptian and/or Albian of Niger (Suchomimus tenerensis; Sereno et al. 1998;n.b., Suchomimus tenerensis most likely represents the same animal as the non-diagnostic Cristatusaurus lapparenti Taquet & Russell 1998 from the same deposits; Carrano et al. 2012), Brazil (Angaturama limai, Irritator challengeri; Kellner & Campos 1996;Sues et al. 2002;n.b., these two taxa known from non-overlapping cranial material recovered from the same deposits may in fact represent the same taxon/individual; Sereno et al. 1998;Sues et al. 2002;Dal Sasso et al. 2005) and South-eastern Asia (Ichthyovenator laosensis; Allain et al. 2012). The most derived spinosaurid, Spinosaurus aegyptiacus, comes from the Albian-Cenomanian of North Africa (e.g., Stromer 1915;Taquet & Russell 1998;Buffetaut & Ouaja 2002;Dal Sasso et al. 2005;Ibrahim et al. 2014). ...
Theropods form a taxonomically and morphologically diverse group of dinosaurs that include extant birds. Inferred relationships between theropod clades are complex and have changed dramatically over the past thirty years with the emergence of cladistic techniques. Here, we present a brief historical perspective of theropod discoveries and classification, as well as an overview on the current systematics of non-avian theropods. The first scientifically recorded theropod remains dating back to the 17th and 18th centuries come from the Middle Jurassic of Oxfordshire and most likely belong to the megalosaurid Megalosaurus. The latter was the first theropod genus to be named in 1824, and subsequent theropod material found before 1850 can all be referred to megalosauroids. In the fifty years from 1856 to 1906, theropod remains were reported from all continents but Antarctica. The clade Theropoda was erected by Othniel Charles Marsh in 1881, and in its current usage corresponds to an intricate ladder-like organization of textquoteleftfamilytextquoteright to textquoteleftsuperfamilytextquoteright level clades. The earliest definitive theropods come from the Carnian of Argentina, and coelophysoids form the first significant theropod radiation from the Late Triassic to their extinction in the Early Jurassic. Most subsequent theropod clades such as ceratosaurs, allosauroids, tyrannosauroids, ornithomimosaurs, therizinosaurs, oviraptorosaurs, dromaeosaurids, and troodontids persisted until the end of the Cretaceous, though the megalosauroid clade did not extend into the Maastrichtian. Current debates are focused on the monophyly of deinonychosaurs, the position of dilophosaurids within coelophysoids, and megaraptorans among neovenatorids. Some recent analyses have suggested a placement of dilophosaurids outside Coelophysoidea, Megaraptora within Tyrannosauroidea, and a paraphyletic Deinonychosauria with troodontids placed more closely to avialans than dromaeosaurids.
... The Spinosauridae constitute a family of theropod dinosaurs ranging in age from the Late Jurassic to the early Late Cretaceous in Africa (e.g. Stromer, 1915;Taquet and Russell, 1998;Buffetaut and Ouaja, 2002;Buffetaut, 2008), South America (Kellner and Campos, 1996;Martill et al., 1996;Kellner et al., 2011), Europe (Charig and Milner, 1997;Buffetaut, 2007;Canudo et al., 2008), Australia (Barrett et al., 2011) and Asia (Buffetaut and Ingavat, 1986;Buffetaut et al., 2008). In North Africa, remains of spinosaurids are relatively common and diversified in a sedimentary series referred to as the 'Continental intercalaire'. ...
The informally called ‘Continental intercalaire’ is a series of continental and brackish deposits that outcrops in several regions of North Africa. The age of the series is not well-constrained, but its upper part, visible in the ‘Kem Kem beds’ in Morocco and in Bahariya in Egypt, is regarded as early Cenomanian in age. Spinosaurid remains are an important component of this series, but records of this dinosaur are surprisingly rare in Algerian localities of the ‘Continental intercalaire’. Here, we describe a vertebrate assemblage from two localities, Kénadsa and Menaguir, situated in the Guir basin, Western Algeria. The assemblage comprises hybodont sharks, sarcopterygian fishes, ray-finned fishes, turtles, crocodiles and dinosaurs. Among the latter, only teeth of theropods have been recovered and 94% belong to Spinosaurus. The assemblage is taxonomically very similar to the Moroccan and Egyptian assemblages mentioned above. This study: 1) suggests a likely early Cenomanian age for the Guir basin deposits containing the assemblage; 2) provides a new evidence of the homogeneity of the early Cenomanian vertebrate fauna throughout North Africa; and 3) confirms the overabundance of theropod dinosaurs, especially spinosaurs, in the assemblage showing a possible shortcut in the vertebrate food chain. The northern most locality, Menaguir, shows sedimentological and ichnological evidence of marine influences indicating that the palaeoenvironment shows spatial heterogeneities.
... However, we feel specimens without direct provenance data may be useful when they present distinct morphology and produce scientific clarity. For example, Irritator challengeri is represented by a skull of a spinosaurid theropod (Martill et al. 1996) missing the rostral end of the skull that had been privately collected prior to purchase by the Staatliches Museum fü r Naturkunde Stuttgart; however, a specimen recovered directly from the Romualdo Formation had been named Angaturama limai (Kellner and Campos 1996) and preserves only the rostral end of a snout, corresponds directly to the missing section in Irritator (Charig and Milner 1997). Indeed, these two specimens are so similar they have been implied to belong to the same individual (Charig and Milner 1997), which certainly implies that Irritator was likely derived from the Romualdo Formation as we know Angaturama to. ...
Numerous taxa make up the Early Cretaceous fauna of Brazil, including Ornithocheiroidea, Tapejaridae,
Thalassodromidae, Chaoyangopteridae and a purported member of Azhdarchidae. Dsungaripteridae has only been tentatively assumed to be present in the form of ‘Santanadactylus’ spixi. New study of NMSG SAO 251093 (a specimen referred to Thalassodromeus sethi) suggests it is a previously unknown species of dsungaripterid, Banguela oberlii, tax. nov., differing from Thalassodromeus and other pterosaurs from the Early Cretaceous of Brazil by a unique combination of characters, including an upturned jaw tip, a short dorsal mandibular symphyseal shelf (dmss), and an autapomorphic thin crest placed halfway along the fused mandibular symphysis without a keel along the ventral margin of the jaw. B. oberlii, tax. nov., is referred to Dsungaripteridae based on a dmss no longer than the ventral shelf, U-shaped caudal margin of the ventral shelf and lateral margins of the mandibular symphysis concave in dorsal view. B. oberlii, tax. nov., is the youngest known dsungaripterid, and expands known morphological diversity in the clade as well as the Early Cretaceous pterosaur fauna of South America.
Notosuchia are terrestrial crocodyliforms that have lived during the Cretaceous. The morphological variation in this group implies different adaptative and physiological factors that can be registered in the bone microstructure. Here, were analyzed long bones and rib fragments belonging to Notosuchia aiming to describe the bone histology, verify if any growth patterns, and, possibly, ecological adaptations. The methodology followed standards of paleohistological protocols. The fibolamelar complex is present in the femora, tíbia and ulna of Stratiotosuchus maxhecthi and Baurusuchus sp. In Pissarrachampsa sera the bone matrix is mostly parallel-fibred bone showing a gradual decrease in depositional rates. The ribs showed lamellar-zonal bone, simples vascular arrangement, and extensive remodeling. Armadillosuchus arrudai shows parallel-fibred bone matrix and simple vascular canals and Mariliasuchus amarali is a juvenile with the formation of a fibrolamellar complex in the limb bones and secondary compact cancellous bone in the hand bones. No external fundamental system was found in any of the samples. Notosuchia shows a variable and cyclical growth very similar to other Crocodyliformes, but the histological arrangement suggests higher depositional rates. The rhythm can be associated with the terrestrial ecology and/or body size which can be the case for Baurusuchidae. Given these results, Baurusuchidae seems to grow like Dinosauria and Sphagesauridae similar to Crocodylia. The absence of the external fundamental system is not directly related to indeterminate growth, which can be associated with the K-P transition. The difference in growth physiology compared to the survivors of Pseudosuchia non-Crocodylomorpha, Neosuchia and Eusuchia suggest that the slow and cyclical growth could have been determinant of the maintenance of these groups after the K-P event.
We describe two partial postcranial skeletons belonging to the enigmatic theropod dinosaur clade Megaraptoridae from the Upper Cretaceous (lower Cenomanian-upper Turonian) Bajo Barreal Formation of southern Chubut Province, central Patagonia, Argentina. The specimens are assigned to Megaraptoridae due to their possession of multiple anatomical features that are considered synapomorphies of that predatory dinosaur group, such as a greatly enlarged, laterally compressed ungual of manual digit I that possesses asymmetrical lateral and medial vascular grooves. Overlapping elements of the two skeletons are nearly identical in morphology, suggesting that they probably represent the same taxon, a large-bodied theropod that was previously unknown from the early Late Cretaceous of southern South America. The Bajo Barreal specimens constitute the most ancient unquestionable records of Megaraptoridae from that continent, and exhibit particularly strong osteological resemblances to penecontemporaneous megaraptorids from the Winton Formation of Australia. Phylogenetic analysis recovers the unnamed Bajo Barreal taxon as the earliest-diverging South American megaraptorid and the oldest-known representative of this clade that likely attained a body length of at least seven meters and a mass of at least one metric ton. Overall, the balance of the evidence suggests that megaraptorids originated in eastern Gondwana (Australia) during the Early Cretaceous, then subsequently dispersed to western Gondwana (South America) during the mid-Cretaceous, where they attained substantially larger body sizes, ultimately coming to occupy the apex predator niches in their respective habitats.
A new locality near Tarda on the northern margin of the Tafilalt, south eastern Morocco exposes extensive sequences of the Ifezouane and Aoufous formations of the fluvial Kem Kem Group (Cretaceous, ?Albian-Cenomanian) on the south western flank of Ikfh n’Oufza escarpment of the Hamada du Meski. The stratigraphic sequence here differs significantly from better known exposures of the Kem Kem Group in the southern Tafilalt, and includes a heterolithic sequence of alternating grey mudstones and fine sandstones and a thin (∼1.5 m) marine limestone. The locality is noteworthy for three vertebrate-bearing horizons within the upper part of the Ifezouane Formation. The upper two (Sites 1 and 2 in ascending order) are dominated by dental remains of the sawfish Onchopristis and the aquatic theropod dinosaur Spinosaurus. Significantly, the remains of terrestrial dinosaurs constitute less than 1 % of the total dental assemblage at Site 1 and 5.6% at Site 2. At Site 2 teeth of Spinosaurus outnumber the rostral “teeth” of Onchopristis. The remarkably high abundance of spinosaur teeth compared to remains of terrestrial dinosaurs, and even some aquatic animals strongly supports Spinosaurus being an aquatic animal spending much of its life in water where its teeth were shed and preserved.
The Romualdo Formation (Araripe Basin) is worldwide known for the large number of well-preserved fossils but the dinosaur record is rather scarce. Here we describe a new coelurosaur, which is the first tetrapod recovered from the basal layers of this stratigraphic unit that consist of dark shales. Aratasaurus museunacionali gen. et sp. nov. is known by an incomplete but articulated right hind limb with the distal portion of the femur, proximal half of tibia and incomplete pes. The new species differs from other coelurosaurs by a medial fossa in the tibia and digits II, III and IV being symmetric. The phylogenetic analysis recovered Aratasaurus museunacionali closely related to Zuolong salleei, forming a basal coelurosaur lineage. The paleohistology indicate that the specimen is a juvenile, with an estimated body length around 3.12 m. The new taxon represents the first occurrence of basal coelurosaurians in the Araripe Basin and suggests a widespread distribution of this group during the Lower Cretaceous.
The spinosaurids represent an enigmatic and highly unusual form of large tetanuran theropods that were first identified in 1915. A recent flurry of discoveries and taxonomic revisions of this important and interesting clade has added greatly to our knowledge. Spinosaur body fossils are however generally rare and most species are known from only limited skeletal remains. Their unusual anatomical adaptations to the skull, limbs and axial column all differ from other large theropods and point to an unusual ecological niche and a lifestyle intimately linked to water.
The study of bone microstructure preserved in fossils provides substantial information about physiology, growth gradients and strategies, and some ecological considerations. Paleohistology is a useful tool for understanding the biological mechanisms of extinct animals. Presented here is the microstructure characterization of two Anhangueria pterosaurs. Thin sections of the first wing phalanx and metacarpal IV of MN 7060-V have been confectioned, as have sections of the radius, ulna and first wing phalanx of MPSC R2090. The histological analysis of bones of MN 7060-V revealed fibrolamellar tissue, few osteocytes and vascular canals. Bones of MPSC R2090 showed a mixed plexiform-fibrolamellar tissue and histovariability, showing that bones from the same individual grew in different patterns. The vascularization was high in the phalanx, intermediate in the ulna and absent in the radius. The absence of canals in the radius may be related to biomechanical issues, due to torsion resistance during flight. The histology and the absence of fused bones suggests that the specimens are not adults. Two distinct moments of growth were established. MN 7060-V is a subadult, with presence of bone porosity and MPSC R2090 is a young animal as determined by the high number of canals and plexiform-fibrolamellar tissues, which indicates fast growth. In this work, we concluded that in the Anhangueria clade, the growth of bones is not compatible with the ontogenetic stage. Young animals may present large proportions, whereas there were older individuals of smaller sizes in the same clade.
Sigilmassasaurus brevicollis is an enigmatic theropod dinosaur from the early Late Cretaceous (Cenomanian) of Morocco, originally based on a few isolated cervical vertebrae. Ever since its original description, both its taxonomic validity and systematic affinities were contentious. Originally considered to represent its own family, Sigilmassasauridae, the genus has variously been suggested to represent a carcharodontosaurid, an ornithischian, and, more recently, a spinosaurid. Here we describe new remains referrable to this taxon and re-evaluate its taxonomic status and systematic affinities. Based on the new remains, a re-evaluation of the original materials, and comparisons with other spinosaurids, the holotype of Sigilmassasaurus brevicollis is identified as an anterior dorsal, rather than a cervical vertebra, and differences between elements referred to this taxon can be explained by different positions of the elements in question within the vertebral column. Many characters used previously to diagnose the genus and species are found to be more widespread among basal tetanurans, and specifically spinosaurids. However, the taxon shows several autapomorphies that support its validity, including the presence of a strongly rugose, ventrally offset triangular platform that is confluent with a ventral keel anteriorly in the mid-cervical vertebral centra and a strongly reduced lateral neural arch lamination, with no or an incomplete distinction between anterior and posterior centrodiapophyseal laminae in the posterior cervical and anterior dorsal vertebrae. We argue furthermore that Spinosaurus maroccanus, also described on the basis of isolated cervical vertebrae from the same stratigraphic unit and in the same paper as Sigilmassasaurus brevicollis, is a subjective synonym of the latter. Both a detailed comparison of this taxon with other theropods and a formal phylogenetic analysis support spinosaurid affintities for Sigilmassasaurus. However, we reject the recently
Dinosaurs are one of the most dominant groups in Cretaceous reptilian faunas. A summary of their record in northern Brazil and northern Africa during the middle of the Cretaceous Period (Aptian-Cenomanian) is presented here. Dinosaurs are represented by 32 species (three ornithischians, six sauropods and 23 theropods) from Brazil, Egypt, Lybia, Morocco, Niger, Sudan and Tunisia. These dinosaur assemblages provide fundamental data about distribution and composition of sauropods and theropods in northern Gondwana during the middle of the Cretaceous Period and confirm these assemblages to be among the most important dinosaur faunas in the north Gondwana areas.