Middle Cretaceous dinosaur assemblages from northern Brazil and northern Africa and their implications for northern Gondwanan composition

Accepted Manuscript Middle Cretaceous dinosaur assemblages from northern Brazil and northern Africa and their implications for northern Gondwanan composition Carlos Roberto A. Candeiro PII:

S0895-9811(14)00142-4

DOI:

10.1016/j.jsames.2014.10.005

Reference:

SAMES 1337

To appear in:

Journal of South American Earth Sciences

Received Date: 4 May 2014 Revised Date:

6 October 2014

Accepted Date: 19 October 2014

Please cite this article as: Candeiro, C.R.A., Middle Cretaceous dinosaur assemblages from northern Brazil and northern Africa and their implications for northern Gondwanan composition, Journal of South American Earth Sciences (2014), doi: 10.1016/j.jsames.2014.10.005. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.

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ACCEPTED MANUSCRIPT Middle Cretaceous dinosaur assemblages from northern Brazil and northern Africa and their implications for northern Gondwanan composition

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Carlos Roberto A. Candeiro Laboratório de Geologia, Curso de Geografia, Campus Pontal, Universidade Federal de

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Uberlândia, Rua 20, n. 1.600, Bairro Tupã, Ituiutaba, Minas Gerais State, Brazil

Abstract

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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

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fundamental data about distribution and composition of sauropods and theropods in northern Gondwana during the middle of the Cretaceous Period and confirm these

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assemblages to be among the most important dinosaur faunas in the north Gondwana areas.

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Key-words: Gondwana, dinosaur faunas, Africa, Brazil, Cretaceous.

Introduction

The three major clades of dinosaurs (Ornithischia, Sauropoda and Theropoda)

were widespread and diverse during the middle of the Cretaceous Period in Gondwana. Cretaceous dinosaur assemblages have long been known from southern continents where appropriate rocks are exposed. This northern Gondwana dinosaur fauna has considerable potential for new discoveries (e.g., Brazil: Hauterivian Corda Formation

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in north Tocantins State; Albian Santana Formation in eastern Piauí State; Lybia: Aptian-Albian Chicla Formation; Sudan: Cenomanian Wadi Milk Formation). Gondwanan dinosaur records from these southern continents could foster an increased

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understanding of the global composition of the fauna during the middle Cretaceous Period.

This analysis is based on the dinosaur taxa in northern Brazil and northern

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Africa, which provide a significant amount of information on dinosaur faunas and allow an understanding of the composition of these assemblages from the Aptian to the

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Cenomanian on northern Gondwanan continents.

Gondwana paleobiogeography during the Cretaceous Period has been studied and analyzed by many geoscientists (e.g., Krause et al., 1997; Hay et al., 1999; Novas et al., 2005, Yoder and Nowak, 2006; Upchurch, 2008; Novas, 2009; Candeiro, 2010; Candeiro et al., 2011; Jacobs et al., 2011; Fanti, 2012; Fanti et al., 2012, 2013, 2014),

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and northern Brazil and northern Africa are prominent topics of these studies due to the close similarities between their close geological and dinosaur faunal content. In this

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paper I review the northern Brazilian and African Cretaceous dinosaur fauna

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distribution during the middle of the Cretaceous Period.

Methods

Extensive records were collected from the literature for three major dinosaur groups: ornithischians, sauropods and non-avian theropods. These clades represent the major dinosaur groups using a logical diversity and distribution of comparison. Geological and chronological data were obtained from Itapecuru Group (Rossetti and Truckenbrodt, 1997), Santana (Martill, 1993), Elrhaz and Echkar (Taquet, 1976), Wadi Milk (Werner, 1994), Bahariya (Catuneanu et al., 2006), Chicla (Barale et al., 1997; Barale and Ouaja,

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2002), Douiret and Ain El Guettar formations (Bouaziz et al., 1988; Bondin et al., 2010), and Kem Kem beds (Cavin et al., 2010).

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Cretaceous dinosaurs record from North Brazil and North Africa The dinosaur record (Fig. 1) analyzed here is restricted to continental middle Cretaceous rocks from Brazil, Egypt, Lybia, Morocco, Niger, Sudan and Tunisia. Most

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information on the middle Cretaceous dinosaur faunas of north Brazil and north Africa come from Albian and Cenomanian stages and, above all, from collection made of

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Albian-Early Cenomanian Itapecuru Group and Albian Santana in North Brazil; AptianAlbian Ain El Guettar Formation, Aptian-Albian Elrhaz Formation, Aptian-Albian Chicla Formation, Aptian Douret Formation, Early Cenomanian Bahariya, Cenomanian Wadi Milk Formation and Cenomanian Kem Kem beds in North Africa. On the tectonic divisions, North Brazil and North Africa belong to the same ancient Gondwana plate,

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and this is also supported by Cretaceous dinosaur content.

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------------------------------------- Figure 1 near here

North Brazil

Santana Formation (Arapipe Basin, sensu Martill, 1993) The Albian Santana Formation has produced only theropod records with two

described spinosaurid species and some additional postcranial remains (Kellner 1996; Kellner and Campos 1996; Martill et al., 1996; Sues et al., 2002; Bittencourt and Kellner 2004; Machado and Kellner, 2005). Irritator challengeri Martill, Cruickshank, Frey, Small and Clarke, 1996 was first described based on a partial skull from the

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Santana Formation. Later, I. challengeri was considered a senior synonym of Angaturama limai by Charig and Milner (1997), and Sereno et al. (1998) confirmed it as a spinosaurid species (Sues et al., 2002). Frey and Martill (1995) described a sacrum

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and assigned it to Oviraptorosauria indet., but it has been later suggested that this theropod needs to be revised (Makovicky and Sues 1998; Kellner 1996; Agnolin and Martinelli, 2007). The small-bodied Santanaraptor placidus Kellner, 1999 was

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atributed by Holtz (2004) and Porfiri et al. (2014) to Tyrannosauroidea. Mirischia asymmetrica Naish, Martill and Frey, 2004 was described as the first Brazilian

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Compsognathidae from the Santana Formation, based on incomplete skeletal remains.

Itapecuru Group (Rossetti and Truckenbrodt, 1997)

The Itapecuru Group (Albian age “Undifferentiated Unit” and Early Cenomanian Alcântara; Rossetti and Truckenbrodt, 1997) dinosaur fauna is composed

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of sauropods and theropods and a possible ornithischian record (Avilla et al., 2003; Candeiro et al., 2011). The sauropod dinosaur fauna recorded from the Albian

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“Undifferentiated Unit” includes the diplodocoid Amazonsaurus maranhensis Carvalho, Avilla and Salgado, 2003 and Titanosauria indet. (Castro et al. 2007; Lindoso et al.,

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2013). From the Early Cenomanian Alcântara Formation, large-bodied theropods are represented by the spinosaurid Oxalaia quilombesis Kellner, Azevedo, Machado, Carvalho and Henriques, 2011, Carcharodontosaurus sp., Spinosaurus sp. (Medeiros et al., 2014), and Baryonychinae indet. (Furtado and Candeiro, 2009a, 2009b). An isolated vertebra atributted to Sigilmassasaurus brevicollis Russel, 1996 was briefly described by Medeiros and Schultz (2002), but this Brazilian record needs to be revised and will probably be assigned to Spinosauridae indet. (Bradley Mcfeeters pers. comm.). Small theropod dinosaurs are represented by Dromaeosauridae indet. that have been recently

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confirmed by Medeiros et al. (2014) as cf. Masiakasaurus (Lindoso et al., 2012). The sauropod dinosaur fauna recorded from this unit includes the rebbachisaurid cf. Limaysaurus (Medeiros and Schultz, 2004; Medeiros et al., 2014) and Diplodocoidea

North Africa Egypt Bahariya Formation (Catuneanu et al., 2006)

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indet. (Medeiros and Shultz, 2004).

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The dinosaur fauna of the Egypt comes from Early Cenomanian Bahariya Formation (Catuneanu et al., 2006; Le Loeuff et al., 2012) and includes titanosaurs, ceratosaurs, and tetanuran theropods; titanosaurians are represented by Aegyptosaurus baharijensis Stromer, 1932 and Paralititan stromeri Smith, Lamanna, Lacovara, Dodson, Smith, Poole, Giegengack and Attia, 2001. Large-bodied theropods have also

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been reported from Bahariya Formation: ceratosaurian Bahariasaurus ingens Stromer, 1934, the carcharodontosaurid Carcharodontosaurus saharicus Stromer, 1931, and the

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spinosaurid Spinosaurus aegyptiacus Stromer, 1915.

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Lybia

Chicla Formation (Barale et al., 1997; Barale and Ouaja, 2002) The Aptian-Albian Chicla Formation (Barale et al., 1997; Barale and Ouaja,

2002) dinosaur record is poorly known. However, Smith and Dalla Vechia (2006) described abelisaurid teeth that resemble Indosuchus, and confirmed the fossiliferous character of this unit.

Morroco

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The Cenomanian Kem Kem beds (Cavin et al., 2010) have a remarkable dinosaur record, especially with respect to theropods. Among sauropods, the taxa present include

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the diplodocoid Rebbachisaurus garasbae Lavocat, 1954, Somphosponyli indet. (Mannion and Barrett, 2013; Lamanna and Hasegawa, 2014), Rebbachsauridae indet. (Mannion and Barrett, 2013), and Lithostrotia indet. (Cavin et al., 2010). From these

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beds, the giant theropods are represented by the Deltadromeus agilis Sereno et al. 1996 (ceratosaur sensu Carrano and Sampson, 2002, 2008; Wilson et al., 2003; Sereno et al., abelisaurid

cf.

Majungasaurus

(Russell,

1996),

carcharodontosaurids

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2004),

Carcharodontosaurus saharicus Stromer, 1931 (Sereno et al., 1996; Brusatte and Sereno, 2007) and Sauroniops pachytholus Cau, Dalla Vecchia and Fabbri, 2013, spinosaurid Spinosaurus cf. aegyptiacus described by Dal Sasso et al. (2005), spinosaurid Sigilmassasaurus brevicollis Russell, 1996 (Evers et al., 2012) and

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indeterminate teeth of Abelisauridae, Carcharodontosauridae (Richter et al., 2013) and Dromaeosauridae indet. (Amiot et al., 2004; Richter et al., 2013). Spinosaurus

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maroccanus was described by Russel (1996), but Sereno et al. (1998) and Dal Sasso et al. (2005) considered it a nomen dubium. Recently, Evans et al. (2014) described a

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small-bodied Averostra indet. and Noasauridae indet. from Morrocan Kem Kem beds, and McFeeters (2013) reported Noasauridae indet. and na Saurischia incertae sedis. Several ornithischian, sauropod and theropod tracks have been found in this fossiliferous unit (Ibrahim et al., 2014).

Niger Elrhaz Formation (Taquet, 1976)

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The Nigerian Aptian-Albian Elrhaz Formation (Taquet, 1976) is an representative geological unit from the middle Creteaceous of northern Gondwana, with ornithischians and saurischians in the dinosaur fauna. The herbivorous dinosaurs

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include the dryosaurid Valdosaurus nigeriensis Galton and Taquet, 1982 (according Galton, 2009), ornithopod Lurdusaurus arenatus Taquet and Russell, 1999 (McDonald et al., 20012), Ouranosaurus nigeriensis Taquet, 1976, and the rebbachisaurid

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Nigersaurus taqueti Sereno, Beck, Dutheil, Larsson, Lyon, Moussa, Sadleir, Sidor, Varricchio, Wilson and Wilson, 1999. The most common theropods from the Elrhaz

Brusatte,

2008,

Rugops

primus

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Formation are large-bodied theropods: the abelisaurid Kryptops palaios Sereno and Sereno,

Wilson

and

Conrad,

2004,

the

carcharodontosaurid Eocarcharia dinops Sereno and Brussate, 2008, and the spinosaurid Suchomimus tenerensis Sereno, Beck, Dutheil, Gado, Larsson, Lyon, Marcot, Rauhut, Sadleir, Sidro, Varricchio, Wilson and Wilson, 1998. However, S.

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tenerensis was redefined as Baryonyx tenerensis by Hutt and Newbery (2004). Cristatusaurus lapparenti Taquet and Russell, 1998 is known from fragmentary

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dubium.

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material, but Sereno et al. (1998) and Sues et al. (2002) consider this species a nomen

Echkar Formation (Taquet, 1976) The Echkar Formation, from the Tegama Series unit, is a geological formation

of Cenomanian age (Taquet, 1976) that have records of sauropods and theropods. The sauropod record is represented by Rebbachisauridae indet. and Titanosauriformes indet. (Sereno et al., 2004). The large abelisaurid Rugops primus Sereno, Wilson and Conrad, 2004, and the tetantuans Carcharodontosaurus iguidensis Brusatte and Sereno 2007,

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Sigilmassasaurus cf. brevicollis (McFeeters et al. 2013), and Spinosaurus sp. (Sereno et al., 2004) were also reported in the Echkar Formation.

Wadi Milk Formation (Werner, 1994)

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Sudan

Most dinosaur specimens collected from the Cenomanian Wadi Milk Formation

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(Werner, 1994) in Sudan are fragmentary materials which were described as cf. Ouranosaurus, Dicraeosauridae indet., Lithostrotia indet., Bahariasaurus sp.,

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Carcharodontosaurus sp., and Dromaeosauridae indet. (Weishampel et al., 1990, 2004; Rauhut and Werner, 1995; Rauhut, 1999).

Tunisia

Douiret and Ain El Guettar formations (Bouaziz et al., 1988; Bodin et al., 2010)

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The fossiliferous potential of the Tunisia Early Cretaceous units is highly significant. The dinosaur record comes from Aptian Douiret and Aptian-Albian Ain El

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Guettar formations of the Tataouine Basin. Dinosaur records from the Early Aptian Douiret Formation (Bouaziz et al., 1988) are represented by Ornithischia indet. (Benton

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et al., 2000) and Carcharodontosaurus sp. (Srarfi et al., 2004). Two members from the Ain El Guettar Formation can be distinguished (Bodin et al., 2010), the lower one, the Aptian Chenini Member, and the upper one, the Early Albian Oum ed Diab (Fanti et al., 2012). The Chenini Member dinosaur records are represented by Carcharodontosaurus sp. (Srarfi et al., 2004), Spinosaurus sp., Abelisauridae indet., Rebbachisauridae indet., and Iguanodontidae indet. described by Fanti et al. (2012). The Early Albian Oum Ed Diab Member dinosaur fauna consists of the Ornithopoda indet., Rebbachisauidae indet., the rebbachisaurid Tataouinea hannibalis Fanti, Cau, Hassine and Contessi,

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2012, and the large-bodied theropods Carcharodontosaurus saharicus (Sereno and Brusatte, 2008), Spinosaurus sp. (Russell, 1998; Buffetaut and Ouaja, 2002), and Abelisauridae indet. (Fanti et al., 2012). Fanti et al. (2014) recently described a diverse

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theropod assemblage from Ain El Guettar beds represented by specimens of Carcharodontosaurus sp., Spinosaurus sp., Baryonichiane indet., Abelisauridae indet.,

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and Theropoda indet.

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Discussion

The dinosaur diversity presented here occurs in twelve lithostratigraphic units distributed over two large continents (Africa and South America) (see Fig. 1 and Box. 1).

The middle Cretaceous North Gondwanan assemblage is commonly found in

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northern African and Brazilian strata. The age of these dinosaur-bearing units ranges from Aptian to Cenomanian. This assemblage is characterized by the co-occurrence of

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Pangean (diplodocoid, compsognathid and tyrannosauroid) and Gondwanan (abelisaurid and carcharodontosaurid) dinosaur taxa. This implies that elements of clades dispersed

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from Laurasia during the Early Cretaceous (e.g., the rebbachisaurid Amazonzaurus maranhensis – according Gallina et al., 2014 the compsognathid Myrischia asymmetrica). Nevertheless, taxa such as Rebbachisauridae and Carcharodontosaurinae do not have records outside Gondwana, which suggests a low Gondwanan endemism during the middle Cretaceous. These latter endemic clades include rebbachisaurid (Rebbachisaurus garasbae, cf. Limaysaurus) sauropods; basal titanosauriforms (Aegyptosaurus baharijensis); and basal abelisaurids (Rugops primus). Most of the North Gondwanan dinosaur fauna has been used to suggest certain similarities with the

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middle Cretaceous Patagonian fauna (Coria and Salgado, 2005; Ibiricu et al., 2012). However, although records of titanosaurians were also found in the Itapecuru Group and Bahariya Fomation, this taxon is not a good indicator to distinguish the assemblages

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from these deposits from Patagonia and northern Gondwana assemblages. The presence in northern Brazilian and northern African geological units of latest basal Titanosauria (A. baharijensis and P. stromeri; Stromer, 1932; Smith et al.,

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2001), carcharodontosaurids (Carcharodontosaurus saharicus, Eoacarcharia dinops, Carcharodontosaurus sp.; Stromer, 1931; Sereno et al., 1998; Rauhut, 1999; Srarfi et

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al., 2004; Sereno and Brussate, 2008) and spinosaurids (Irritator challengeri, Baryonyx tenerensis, Spinosaurus aegyptiacus, Spinosaurus cf. aegyptiacus, Oxalaia quilombesis and Spinosaurus sp.; Stromer, 1915; Martill et al., 1996; Sereno et al., 1998; Buffetaut and Oueja, 2002; Dal Sasso et al. 2005; Medeiros, 2006; Kellner et al., 2011; Medeiros et al., 2014) greatly indicates that some of these taxa were part of the Laurasian fauna.

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Oviraptorosaurs have not been found in northern Africa, but there are records in Brazil. However, Maniraptora are present in Aptian-Cenomanian rocks from Niger,

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Sudan, and Morocco (Rauhut and Werner, 1995; Amiot et al., 2004; Richter et al., 2013). The main middle Cretaceous North Gondwana theropod records are giant

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abelisaurids, carcharodontosaurids, and spinosaurids. Northern Gondwanan dinosaur distribution and diversity have basic similarities

among middle Cretaceous dinosaur-bearing deposits from northern Brazil and northern Africa (Amiot et al., 2004; Cavin et al., 2010; Candeiro et al., 2011; Fanti et al., 2012; Medeiros et al., 2014). The presense of the large theropod carcharodontosaurds Carcharodontosaurus

saharicus

and

Carcharodontosaurus

sp.,

spinosaurid

Spinosaurus aegyptiacus and Spinosaurus sp., and distinctive caudal vertebrae possibly belonging to the tetanuran Sigilmassasaurus brevicollis in the diferent areas of northern

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Gondwana reinforces this faunal similarity (Box 1). The most significant aspect of this assemblage of middle Cretaceous large-bodied theropods is the endemism of the large dinosaur fauna during this time. The more primitive, endemic taxa of the Early

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Cretaceous did not persist into the early Late Cretaceous (e.g., Dicraeosauridae, Carcharodontosauridae, and Spinosauridae). Similarly, there is limited evidence that carcharodontosaurids and spinosaurids, the dominant large theropods of the Aptian-

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Cenomanian of North Gondwana (Fig. 2), developed large body sizes and were the top dinosaur predators during the earlier Aptian (Ain El Guettar Formation). The large-

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bodied theropods of the middle Cretaceous recorded in southern South America (e.g., Neuquén and San Jorge basins, Lamanna et al., 2002, Leanza et al., 2004, Novas et al., 2013; Bauru Group, Candeiro et al., 2006, Bittencourt and Langer, 2011) belonged to the same theropod dinosaur clades that dominated as predators during this late Early and Late Cretaceous. This is not surprising in North Gondwana because it was physically

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linked to other southern continents during the Early Cretaceous (Canudo et al., 2009; Fanti, 2012). Gondwanan sauropods (Rebbachisaurus, Limaysaurus) and abelisaurids

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(cf. Masikiasaurus, Indosuchus sp., cf. Majungasaurus; Russell, 1996; Smith and Dalla Vecchia, 2006; Lindoso et al., 2012) represent members of probable edemic taxa during

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the mid-Late Cretaceous that have been recorded in the Late Cretaceous (Maastrichtian Masiakasaurus and Majungasaurus in Madagascar, and Indosuchus in India).

---------------------------------------------Box 1 near here

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The middle Cretaceous dinosaur fauna from northern Gondwana represent important records to studies of paleogeography and paleodiversity of the Gondwanan

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fauna. Although northern Brazil and northern Africa dinosaurs from middle Cretaceous show close similarities, these faunas remain poorly understood.

The affinities of the middle Cretaceous taxa that occur in nine dinosaur-bearing

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geological units from northern Africa and northern Brazil suggest that a basically homogenous faunal assemblage was present on northern Gondwana during the Aptian

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to the Cenomanian. These dinosaur deposits do not present a complete faunal isolation of Laurasia and Gondwana (e.g., such as Europe and southern America). Therefore, it is possible that by the early Cretaceous there was little paleobiogeograhic barriers for dinosaurs to substantially speciate into a wide variety of taxa.

According to the current information on the distribution and diversity of the

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dinosaur record, the faunal composition of the northern Gondwana fauna seems to be correlated with the evolution of the diplodocoid, abelisaurid, carcharodontosaurid, and

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spinosaurid dinosaurs. However, there are still unknown aspects of the middle Cretaceous dinosaur fauna, and there is a seemingly low effort to conduct prospecting

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and collection of more fossil materials. Thus, in order to advance in the knowledge of dinosaur faunas from North Gondwana it is crucial to conduct more studies with integrated paleontological work on this field.

Acknowledgements The author would like to especially thank Michael D´Emic (State University of Stony Book, New Yor, USA) for his friendship and contributions on this article. I also acknowledge two anonymous for all suggested improvements to this paper, and to

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Bradley McFeeters (Carleton University/Canada) and Camila Bernardes (Universidade Federal do Estado do Rio de Janeiro) by comments and grammar improved. This contribution was partially supported by the Conselho Nacional de Ciência e Tecnologia

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Candeiro by Produtividade Pesquisa CNPq fellow.

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Figure Captions

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Evolution, and Systematics 37, 405-431.

Figure 1. Middle Cretaceous dinosaur-bearing units of northern Gondwana (modified

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from Scotese, 2000).

Figure 2. The deep-time paleodistribution of major dinosaur families during middle Cretaceous from northern Gondwana.

Box 1. Dinosaur fauna from northern Gondwana areas recorded in middle Cretaceous dinosaur-bearing units.

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Box 1. Brazil

Egypt

Libya

Morocco

Niger

Sudan

Albian age

Early Cenomanian

Cenomanian Bahariya

Aptian-Albian

Cenomanian Kem Kem

Aptian-Albian

Cenomanian

Cenomanian Wadi

Fm.

“Undifferentia

Alcântara Fm.

Fm.

Chicla Fm.

Beds

Elrhaz Fm.

Echkar Fm.

Milk Fm.

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Albian Santana

ted Unit” Ornithischia Ornithischia indet.

Ouranosaurus

Ouranosaurus sp.

Valdosaurus

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nigeriensis

Aptian Douiret Fm.

Albian Chenini Mb.

Early Albian Oum ed Diab Mb. (Ain El

(Ain El Guettar Fm.) Guettar Fm.)

Ornithischia indet.

Ornithopoda indet.

Ornithopoda indet.

Rebbachisauridae

Tataouinea hannibalis

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nigeriensis

Tunisia

Lurdusaurus arenatus

Sauropoda Rebbachisaurus

baharijensis

garasbae

Diplodocoidea indet. Titanosauria indet.

Paralititan stromeri Titanosauria indet.

Nigersaurus

Rebbachisauridae

taqueti

indet.

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maranhensis

Aegyptosaurus

Titanosauriformes

indet.

indet.

Lithostrotia indet.

Somphosponyli indet. Theropoda

Dicraeosauridae indet.

indet. Lithostrotia indet.

Rebbachisauridae

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cf. Limaysaurus

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Amazonsaurus

Rebbachisauridae indet.

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Irritator

Sigilmassasaurus

challengeri

brevicollis

Bahariasaurus ingens

Indosuchus

Deltadromeus agilis

Kryptops

Sigilmassasaurus

palaios

Carcharodontosaurus

Rugops primus

Bahariasaurus sp.

Sigilmassasaurus

Carcharodontosaurus

cf. brevicollis

sp.

Carcharodontosaurus

Carcharodontosaurus

Carcharodontosaurus

sp.

sp.

sp.

Abelisauridae indet.

Spinosaurus sp.

Spinosaurus sp.

Abelisauridae indet.

Abelisauridae indet.

Carcharodontosaurus sp.

placidus

Spinosaurus sp

Carcharodontosaurus Eoacarcharia

Spinosaurus

Mirischia asymmetrica

Rugops primus

saharicus

saharicus Oxalaia quilombesis

dinops

aegyptiacus Sauroniops

Spinosauridae

cf. Masiakasaurus

pachytholus

indet.

Baryonyx tenerensis

Baryonychinae indet. aegyptiacus

cf. Majungasaurus

Carcharodontosauridae indet

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Abelisauridae indet.

Noasauridae indet.

Dromaeosauridae indet.

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Dromaeosauridae indet.

Averostra indet.

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indet.

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Spinosaurus cf.

Oviraptorosauria

Saurischia incertae sedis

Dromaeosauridae indet.

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Santanaraptor

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brevicollis

Baryonychinae indet.

Theropoda indet.

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Middle Cretaceous dinosaur composition from northern Gondwana were examined. Mainly theropod species are found regularly co-occurring. A comparison with coeval dinosaur reported from Brazil, Egypt, Libya Morocco, Niger, Sudan and Tunisia, support a connection land active during the Middle Cretaceous.