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Continental fossil vertebrates from the mid-Cretaceous (Albian–Cenomanian)Alcântara Formation, Brazil, and their relationship with contemporaneousfaunas from North Africa

Carlos Roberto A. Candeiro a,⇑, Federico Fanti b, François Therrien c, Matthew C. Lamanna d

a Laboratório de Geologia, Curso de Geografia, Universidade Federal de Uberlândia, Av. João Naves de Ávilla, 2160, Sta. Mônica, Uberlândia, Minas Gerais, Brazilb Dipartimento di Scienze della Terra e Geologico-Ambientali, Alma Mater Studiorum, Università di Bologna, Via Zamboni 67, I-40127 Bologna, Italyc Royal Tyrrell Museum of Palaeontology, Box 7500, Drumheller, Alberta T0J 0Y0, Canadad Section of Vertebrate Paleontology, Carnegie Museum of Natural History, 4400 Forbes Ave., Pittsburgh, PA 15213, USA

a r t i c l e i n f o

Article history:Received 13 January 2010Received in revised form 8 February 2011Accepted 9 February 2011Available online 19 February 2011

Keywords:PaleobiogeographyBrazilNorth AfricaCretaceousFaunal exchange

a b s t r a c t

The Albian–Cenomanian Alcântara Formation of northeastern Brazil preserves the most diverse continen-tal vertebrate fauna of this age yet known from northern South America. The Alcântara vertebrate assem-blage, consisting of elasmobranchs, actinopterygians, sarcopterygians, turtles, crocodyliforms, pterosaurs,and non-avian dinosaurs, displays close similarities to contemporaneous faunas from North Africa. Theco-occurrence of as many as eight freshwater or estuarine fish taxa (Onchopristis, Bartschichthys, Lepidotes,Stephanodus, Mawsonia, Arganodus, Ceratodus africanus, and possibly Ceratodus humei) and up to seventerrestrial archosaur taxa (Sigilmassasaurus, Rebbachisauridae, Baryonychinae, Spinosaurinae, Carcharod-ontosauridae, possibly Pholidosauridae, and doubtfully Bahariasaurus) suggests that a land route connect-ing northeastern Brazil and North Africa existed at least until the Albian. Interestingly, most componentsof this mid-Cretaceous northern South American/North African assemblage are not shared with coevalsouthern South American faunas, which are themselves characterized by a number of distinct freshwaterand terrestrial vertebrate taxa (e.g., chelid turtles, megaraptoran and unenlagiine theropods). Theseresults suggest that, although mid-Cretaceous faunal interchange was probably possible between north-ern South America and North Africa, paleogeographic, paleoclimatic, and/or paleoenvironmental barriersmay have hindered continental vertebrate dispersal between northern and southern South America dur-ing this time.

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1. Introduction

Knowledge of the continental vertebrate faunas of the exten-sive, mid-Cretaceous to ?Cenozoic, coastal to continental depositsof the Itapecuru Group in northern Maranhão State, Brazil(Fig. 1), has improved dramatically over the past 30 years (e.g.,Cunha and Ferreira, 1980; Carvalho and Campos, 1988; Carvalho,1994, 1995; Carvalho and Gonçalves, 1994; Medeiros and Schultz,2001a, 2002, 2004; Nobre and Carvalho, 2002; Carvalho et al.,2003; Castro et al., 2004, 2007; Nobre, 2004; Medeiros, 2006; Eliaset al., 2007a,b; Medeiros et al., 2007; Kellner et al., 2009).Vertebrate fossils from the Itapecuru Group include those of fishes,turtles, plesiosaurs, mosasaurs, crocodyliforms, pterosaurs, andnon-avian dinosaurs. Within the Itapecuru Group, the Albian–Cenomanian Alcântara Formation has yielded the greatest diversityand abundance of continental vertebrates. However, few studies

documenting the variety of taxa known from this unit have beenpublished (e.g., Corrêa Martins, 1997; Medeiros and Schultz,2001a, 2002; Medeiros et al., 2007). In this paper, we review andrevise the continental (i.e., freshwater, estuarine, and terrestrial)vertebrate taxa recovered from the Alcântara Formation, and com-pare the fauna of this unit to those of other mid-Cretaceous WestGondwanan (i.e., African and other South American) deposits, tobetter establish the paleobiogeographic relationships of SouthAmerica and Africa during this interval.

2. Geological setting

Located in northeastern Brazil (Fig. 1), the Itapecuru Groupcomprises intercalated marine, paralic, and continental sedimentsthat range in age from the Early Cretaceous (Albian) to possiblythe early Cenozoic (Pedrão et al., 1993; Rossetti, 1997; Rossettiand Truckenbrodt, 1997). Rifting between South America andAfrica that began during the Early Cretaceous led to rapid marinetransgression and, eventually, the deposition of the Itapecuru

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⇑ Corresponding author. Tel.: +55 54 34 3268 6118; fax: +55 51 34 3269 2389.E-mail addresses: candeiro@yahoo.com.br, candeiro@pontal.ufu.br (Carlos Ro-

berto A. Candeiro).

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Group. Although the lowermost portion of the Itapecuru Groupconsists of undifferentiated deposits (informally termed the‘‘Undifferentiated Unit’’ by some authors [e.g., Castro et al.,2007]), two formal stratigraphic units are recognized in the upperportion: the Alcântara and Cujupe formations (Rossetti, 1997;Rossetti and Truckenbrodt, 1997, 1999). The Alcântara Formationis extensively exposed in the São Luís Basin (Fig. 1), where it con-formably overlies undifferentiated strata of the lower ItapecuruGroup (Fig. 2). Considered Albian–Cenomanian in age by mostworkers (e.g., Rossetti and Truckenbrodt, 1997, 1999), this forma-tion reaches a maximum thickness of 35 m and is composed offine- to medium-grained sandstones interbedded with mudstonesand limestones (Rossetti, 1997). Late Cretaceous through possiblyearly Cenozoic deposits of the Cujupe Formation overlie theAlcântara Formation (Rossetti and Truckenbrodt, 1997; Dias Limaand Rossetti, 1999) (Fig. 2).

2.1. Paleogeographic context

Cretaceous faunal similarities between continental Africa andSouth America are widely documented in the dinosaurian fossil

record (e.g., Calvo and Salgado, 1995; Russell, 1996; Sereno et al.,1996, 1998, 1999, 2004; Forster, 1999; Sues et al., 2002; Upchurchet al., 2002; Candeiro et al., 2004, 2006; Holtz et al., 2004; Coriaand Currie, 2006; Wilson, 2006; Sereno and Brusatte, 2008; Cavinet al., 2010; Le Loeuff et al., 2010; Smith et al., 2010). However, itis important to view paleontological discoveries in light of strati-graphic and paleogeographic data when generating and evaluatingpaleobiogeographic hypotheses. According to current paleogeo-graphic models, the South Atlantic began to open northward inthe Early Cretaceous, with emplacement of oceanic crust betweensouthern Africa and southern South America initiating atapproximately 135 Ma (Viramonte et al., 1999; Jokat et al., 2003;Macdonald et al., 2003). Continental extension between WestAfrica and Brazil resulted in the development of the evaporites ofthe South Atlantic salt basin, Barremian-aged oceanic crust, and riftpropagation from east to west across the West African margin(Karner and Gamboa, 2007). The barrier to southern Atlantic mar-ine incursions relates to the magmatic constructions of a proto-Walvis Ridge and the long-lived anomalous topography of thesoutheastern Brazilian highlands (Gradstein et al., 2004; Karnerand Gamboa, 2007). In the Early Cretaceous, isolated rift/

Fig. 1. Location and schematic geological map of the São Luis Basin in northeastern Maranhão State, Brazil.

80 Carlos Roberto A. Candeiro et al. / Journal of African Earth Sciences 60 (2011) 79–92

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strike–slip basins provided intermittent shallow-water connec-tions in relatively small and isolated basins, and during the Ap-tian–Albian a major rift phase led to deeper basins withrestricted deep water circulation (Pletsch et al., 2001; Macdonaldet al., 2003). No later than the early Cenomanian, a mature spread-ing stage allowed a continuous and permanent deep-water con-nection between the southern and northern Atlantic, interruptingpossible terrestrial connections and faunal interchanges betweenSouth America and Africa (Petri, 1987; Reyment and Dingle,1987; Pletsch et al., 2001; Tello Saenz et al., 2003; Guedes et al.,2005; Bodin et al., 2010).

3. Fossil vertebrates from the Alcântara Formation, and theirpaleogeographic distributions

Vertebrate remains reported from the Alcântara Formation per-tain to freshwater, estuarine, and terrestrial taxa, including elas-mobranchs, actinopterygians, coelacanths, lungfishes, turtles,plesiosaurs, mosasaurs, crocodyliforms, pterosaurs, and non-aviandinosaurs (Table 1). Putative records of plesiosaurs and mosasaursfrom this unit (Eugênio, 1994; Carvalho et al., 1997, 2000; VilasBôas and Carvalho, 2001; Marinho et al., 2004; Elias et al., 2005;Marinho and Carvalho, 2005) were recently questioned by Kellneret al. (2009, p. 49); consequently, these aquatic reptiles will not beconsidered here.

3.1. Fishes

Fossils of chondrichthyans and actinopterygians are among themost common fish remains found in the Alcântara Formation.Chondrichthyans are known from fin spines of a Tribodus-like hyb-odontid shark (Marques da Silva and Medeiros, 2003; Medeiroset al., 2007) and material referable to Hylaeobatis sp. (Medeiros,2001), Myliobatidae (Ferreira et al., 1995), and the sclerorhynchidsawfish Onchopristis numidus (Pereira and Medeiros, 2003, 2007).Most recently, Pereira and Medeiros (2008) described the newsclerorhynchid taxon Atlanticopristis equatorialis based on rostral

teeth from Cajual Island in northern Maranhão. Actinopterygiansare represented by material referred to several taxa: the polypteridBartschichthys (Pereira and Medeiros, 2003), an indeterminatepycnodontiform (Sousa et al., 2004), the semionotiform Lepidotes(e.g., Ferreira et al., 1995; Medeiros and Schultz, 2001a; Medeiroset al., 2007), and the teleosts Eotrigonodon and Stephanodus(Pereira and Medeiros, 2003) (Table 1). Although the material inquestion is still undescribed, the report of Bartschichthys is ofparticular interest. If confirmed, this material would representthe oldest record of Polypteriformes outside of continental Africa,and one of the most ancient in the world (see Meunier and Gayet,1996; Gayet et al., 2002).

Sarcopterygian remains from the Alcântara Formation includefossils referred to the giant coelacanthiform Mawsonia sp.(Medeiros and Vilas Bôas, 1999; Medeiros and Schultz, 2001a;Medeiros et al., 2007; Amiot et al., 2010) and multiple dipnoan taxa(Arganodus cf. tiguidiensis [Castro et al., 2004], Ceratodus africanus[Cunha and Ferreira, 1979, 1980; Medeiros and Schultz, 2001a;Toledo and Bertini, 2005; Medeiros et al., 2007], Ceratodus humei[Toledo et al., 2005], and an unidentified, possibly new, ceratodontid[Sousa et al., 2005]) (Table 1).

The coelacanthiform genus Mawsonia has been reported fromEarly and mid-Cretaceous deposits of various sedimentary basinsthroughout Brazil and Africa (e.g., Egypt, Algeria, Morocco, Niger,Democratic Republic of the Congo) (Maisey, 2000; Cavin and Forey,2004; Yabumoto and Uyeno, 2005; Carvalho and Maisey, 2008).Cranial remains and fin rays of Mawsonia from the AlcântaraFormation (Medeiros and Vilas Bôas, 1999; Medeiros and Schultz,2001a; Medeiros et al., 2007; Amiot et al., 2010) are among themost geologically recent, if not the most recent, records of thiscoelacanth yet known from South America.

The lungfish Arganodus tiguidiensis was first recognized (asCeratodus tiguidiensis) from the Late Jurassic and Early Cretaceousof North Africa (Tabaste, 1963). Recently, Castro et al. (2004)inferred the presence of Arganodus (as Asiatoceratodus) cf. tiguidien-sis in the Alcântara Formation, on the basis of two tooth plates(Fig. 3A and B). (Kemp (1998) considered the genus Arganodus tobe a junior synonym of Asiatoceratodus, but this proposed synono-my has not been universally accepted [see Cavin et al., 2007;Agnolin, 2010; Soto and Perea, 2010].) Ceratodus africanus (some-times assigned to the genus Neoceratodus [e.g., Martin, 1982,1984], though probably incorrectly [Kemp, 1997]) occurs widelyin the Cretaceous of North Africa (e.g., Haug, 1905; Peyer, 1925;Arambourg and Joleaud, 1943; Tabaste, 1963; Martin, 1984;Churcher and De Iuliis, 2001; Churcher et al., 2006). Tooth platesreferable to this taxon have also been described from the AlcântaraFormation (Cunha and Ferreira, 1979, 1980; Medeiros and Schultz,2001a; Toledo and Bertini, 2005; Medeiros et al., 2007; Fig. 3C).With the exception of a recent report of A. tiguidiensis and C.africanus from the Late Jurassic–?earliest Cretaceous of Uruguay(Soto and Perea, 2010), the Alcântara specimens constitute the onlyreliable occurrences of these species outside of Africa, demonstrat-ing the similarity of the mid-Cretaceous dipnoan faunas of north-eastern South America and North Africa.

Ceratodus humei was originally described on the basis of toothplates from the Late Cretaceous (Campanian) of Quft, Egypt (Priem,1914). Although additional African Cretaceous specimens have beenreferred to C. humei, according to Churcher and De Iuliis (2001), onlythe tooth plates from Upper Cretaceous (probably Campanian) sed-iments of the Dakhleh Oasis, Egypt that these authors described def-initely pertain to this species. Toledo et al. (2005) recently referredtooth plates and fragmentary mandibular elements from the Alcânt-ara Formation to C. humei (as Protopterus humei). If this referral is cor-rect, these Brazilian specimens would constitute the first occurrenceof C. humei outside of Africa, and possibly the oldest record of thespecies wolrdwide. Finally, Sousa et al. (2005) briefly reported a

Fig. 2. Age and stratigraphy of the Itapecuru Group and its subunits (modified fromRossetti and Truckenbrodt (1997)).

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Table 1Vertebrate fauna of the Alcântara Formation (excluding ichnofossils). Principal references for each record are provided in parentheses.

ChondrichthyesElasmobranchii

Hybodontidae indet. (aff. Tribodus sp. [Marques da Silva and Medeiros, 2003; Medeiros et al., 2007])Ptychodontidae

Hylaeobatis sp. (Medeiros, 2001)Batoidea

Myliobatidae indet. (Ferreira et al., 1995)Sclerorhynchidae

Atlanticopristis equatorialis (Pereira and Medeiros, 2008)Onchopristis numidus (Pereira and Medeiros, 2003, 2007)

OsteichthyesActinopterygii

PolypteridaeBartschichthys sp. (Pereira and Medeiros, 2003)

NeopterygiiPycnodontiformes indet. (Sousa et al., 2004)Semionotiformes

Lepidotes sp. (Ferreira et al., 1995; Medeiros and Schultz, 2001a; Medeiros et al., 2007)Teleostei

Eotrigonodon sp. (Pereira and Medeiros, 2003)Stephanodus sp. (Pereira and Medeiros, 2003)

SarcopterygiiCoelacanthiformes

Mawsonia sp. (Medeiros and Vilas Bôas, 1999; Medeiros and Schultz, 2001a; Medeiros et al., 2007; Amiot et al., 2010)Dipnoi

Arganodus cf. tiguidiensis (Castro et al., 2004)Ceratodus africanus (Cunha and Ferreira, 1979, 1980; Medeiros and Schultz, 2001a; Toledo and Bertini, 2005; Medeiros et al., 2007)Ceratodus humei (Toledo et al., 2005)Ceratodontidae sp. nov.? (Sousa et al., 2005)

TestudinesPleurodira indet. (Moraes-Santos et al., 2001a, 2001b; Oliveira and Romano, 2007)

?Sauropterygia?Plesiosauria indet. (Carvalho et al., 1997, 2000; Vilas Bôas and Carvalho, 2001; Marinho et al., 2004; Elias et al., 2005; Marinho and Carvalho, 2005)

?Lepidosauria?Mosasauridae indet. (Eugênio, 1994; Vilas Bôas and Carvalho, 2001; Marinho and Carvalho, 2005)

CrocodylomorphaCrocodyliformes

Crocodyliformes indet. (Medeiros and Schultz, 2001a; Medeiros et al., 2007)Mesoeucrocodylia

?NotosuchiaCoringasuchus anisodontis (Kellner et al., 2009)

?Pholidosauridae indet. (Nobre et al., 2002; Elias et al., 2005; Kellner et al., 2009)

PterosauriaOrnithocheiroidea

Ornithocheiroidea indet. (Elias et al., 2007a; Lindoso et al., 2008)Anhangueridae indet. (Elias et al., 2007a; Lindoso et al., 2008)

DinosauriaDinosauria incertae sedis

Sigilmassasaurus brevicollis (Medeiros and Schultz, 2001a, 2002; Medeiros et al., 2007)Saurischia

SauropodaSauropoda indet. (Price, 1947; Medeiros et al., 2007; = cf. Astrodon [Medeiros and Schultz, 2001a, 2002])Diplodocoidea

Rebbachisauridae indet. (= Rebbachisaurus sp., = Rebbachisaurus? sp.,=aff. Limaysaurus [‘‘Rayososaurus’’, ‘‘Rebbachisaurus’’] tessonei, = aff.‘‘Rebbachisaurustamesnensis’’ [Medeiros and Schultz, 2001a,b, 2003, 2004; Medeiros and Avilla, 2005; Medeiros et al., 2007])

TitanosauriformesTitanosauriformes indet. (= ‘‘Andesauridae’’ indet. [Medeiros and Schultz, 2001a, 2002; Medeiros et al., 2007])Titanosauria indet. (= Saltasaurinae indet., = ‘‘Titanosauridae’’ indet., = cf. Aegyptosaurus, = cf. ‘‘Titanosauridae’’ indet., = aff. Malawisaurus dixeyi [Medeiros and

Schultz, 2001a, 2002; Medeiros, 2002; Medeiros and Avilla, 2005; Freire et al., 2007; Medeiros et al., 2007])Theropoda

Theropoda indet. (Price, 1960; = aff. ‘‘Elaphrosaurus iguidiensis’’ [Medeiros et al., 2007])Theropoda incertae sedis

?cf. Bahariasaurus ingens (Medeiros and Schultz, 2001a, 2002; Medeiros et al., 2007)Spinosauridae

Spinosauridae indet. (Medeiros, 2006; Medeiros et al., 2007)Spinosauridae sp. nov? (Machado et al., 2009)Baryonychinae indet. (Furtado and Candeiro, 2009a, 2009b)Spinosaurinaecf. Spinosaurus sp. (Medeiros and Vilas Bôas, 1999; Medeiros and Schultz, 2001a, 2002; Medeiros, 2006; Medeiros et al., 2007)

Carcharodontosauridae indet. (= Carcharodontosaurus sp. [Vilas Bôas et al., 1999; Medeiros and Schultz, 2001a, 2002; Medeiros et al., 2007])Maniraptora indet. (= Velociraptorinae indet. [Vilas Bôas, 1999; Elias et al., 2007b])

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possible new taxon of ceratodontid lungfish based on tooth platesfrom the Alcântara Formation that these authors likened to un-named material from the Paleocene of Bolivia.

3.2. Turtles

The only testudine remains definitively known from the Alcânt-ara Formation consist of indeterminate pleurodiran material(Moraes-Santos et al., 2001a,b; Oliveira and Romano, 2007) ofnegligible paleobiogeographic significance. Multiple publications(e.g., Kischlat and Carvalho, 2000; Batista, 2005; Batista andCarvalho, 2007) have described fossils of the pelomedusoid pleuro-dire Araripemys barretoi from Early Cretaceous sediments in north-eastern Brazil that are assigned to the ‘‘Itapecuru Formation’’.Similarly, Batista and Carvalho (2006a,b) reported a possiblepodocnemidid from the ‘‘Itapecuru Formation’’. Unfortunately,however, because the former ‘‘Itapecuru Formation’’ is now knownas the Itapecuru Group, and its Lower Cretaceous deposits havebeen divided between the Albian ‘‘Undifferentiated Unit’’ and theAlbian–Cenomanian Alcântara Formation (e.g., Rossetti andTruckenbrodt, 1997, 1999; Rossetti, 2001; Castro et al., 2007), wehave not been able to conclusively determine whether or not theseturtle fossils were recovered from the latter unit. Consequently,these specimens will not be considered here.

3.3. Crocodyliforms

Remains of crocodyliforms are known from the Alcântara For-mation; however, most reports (e.g., Medeiros and Schultz,2001a; Nobre et al., 2002; Medeiros et al., 2007) of these archo-saurs from the unit consist only of isolated teeth and/or osteo-derms. Nobre et al. (2002), Elias et al. (2005), and Kellneret al. (2009) suspected that at least some of these teeth mightbelong to Pholidosauridae, a Middle Jurassic to mid-Cretaceousneosuchian clade that is known from most continents andincludes the gigantic, well-known genus Sarcosuchus (de Broinand Taquet, 1966; Buffetaut and Taquet, 1977; Sereno et al.,2001). Recently, Kellner et al. (2009) described the new mesoeu-crocodylian Coringasuchus anisodontis based on a dentaryfragment from the Alcântara Formation of the Laje do Coringalocality on Cajual Island. These authors tentatively assignedCoringasuchus to Notosuchia, a morphologically diverse mesoeu-crocodylian clade that is abundant and widespread in continen-tal Cretaceous strata, mostly in the Gondwanan landmasses.

Well-preserved fossils of the notosuchian Candidodon itapecuru-ense occur in some abundance in Lower Cretaceous (Albian)Itapecuru Group deposits near the municipality of Itapecuru-Mirim in northern Maranhão (Carvalho and Campos, 1988;Carvalho, 1994; Nobre and Carvalho, 2002; Nobre, 2004).

Fig. 3. Representative continental vertebrate fossils from the Albian–Cenomanian Alcântara Formation of northeastern Brazil. A and B, tooth plates of the dipnoan Arganodus cf.tiguidiensis; C, tooth plate of the dipnoan Ceratodus africanus; D, skull element Mawsonia; E, Lepidotes scale; F, Atlanticopristis tooth; G and H, incomplete caudal vertebrae of anindeterminate rebbachisaurid sauropod; I, Coringasuchus dentary; J, Pterosauria tooth; K, Spinosaurinae toofh; L, Carcharodontosauridae tooth; M, Maniraptora tooth. (A–Cmodified from Medeiros (2001), D and E modified from Medeiros et al. (2007), F modified from Pereira and Medeiros (2008), G and H modified from Medeiros and Schultz (2004), Imodified from Kellner et al. (2009), J modified from Elias et al. (2007a), K modified from Medeiros (2006), L modified from Elias (2006), M modified from Elias et al. (2007b).)

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Candidodon is a peculiar, small-bodied mesoeucrocodylian withlarge orbits and a short snout (Nobre and Carvalho, 2002). As inseveral other notosuchians, it possesses a strikingly heterodontdentition with a surprisingly mammal-like aspect (Carvalho andCampos, 1988; Carvalho, 1994). Regrettably, however, thestratigraphic position(s) of the fossil-bearing sediments inthe Itapecuru-Mirim region within the Itapecuru Group have notyet been established with certainty (Rossetti, pers. comm., 2010);consequently, it is presently unclear whether known material ofCandidodon pertains to the Alcântara Formation (as stated by Eliaset al. (2005)) or the ‘‘Undifferentiated Unit’’ (as stated by Elias(2006, p. 53)). We therefore cannot confidently regard this crocody-liform as a component of the Alcântara Formation fauna, and as such,we will not consider the taxon further here. It is worth noting,however, that most recent phylogenetic analyses that have includedCandidodon (e.g., Turner and Buckley, 2008; Novas et al., 2009;O’Connor et al., 2010) have recovered this notosuchian as the sistertaxon of, or at least very closely related to, Malawisuchusmwakasyungutiensis (Gomani, 1997) from the Early Cretaceous ofMalawi (Africa).

3.4. Pterosaurs

Brazilian pterosaur fossils are known primarily from mid-Cretaceous sediments in the Araripe Basin. Recently, however, Eliaset al. (2007a) and Lindoso et al. (2008) reported the first pterosaurremains from the Alcântara Formation. This record consists of iso-lated teeth from Laje do Coringa that Elias et al. (2007a) referred toOrnithocheiroidea (as Ornithocheiroidea indet. and Anhangueridaeindet.).

3.5. Dinosaurs

Fossils of a variety of non-avian dinosaurs have been collectedfrom the Alcântara Formation (Table 1). More specifically, the dino-saur fauna of this unit presently includes sauropods, theropods,and Sigilmassasaurus brevicollis, an otherwise North African formof controversial taxonomic validity and systematic position.Originally described as a bizarre theropod of uncertain affinities(Russell, 1996), several recent works (Sereno et al., 1996, 1998;Brusatte and Sereno, 2007) have argued that Sigilmassasaurus isactually a junior synonym of the carcharodontosaurid allosauroidCarcharodontosaurus. Other publications (Novas et al., 2005a,b;Canale et al., 2008; Cau and Maganuco, 2009; Novas, 2009) havemaintained the distinction of the two genera; in addition, severalof these latter works (Novas et al., 2005a; Canale et al., 2008;Novas, 2009) have even suggested that at least some materialassigned to Sigilmassasaurus might pertain to the ornithischiandinosaur clade Ornithopoda.

Among the fossils that Russell (1996) allocated to Sigilmassa-saurus are caudal vertebrae from Egypt and Morocco that possessunusual characters, including a transversely compressed centrumwith a subrectangular contour in anteroposterior view, and anelevated, posterodorsally inclined neural spine that expands intransverse dimension towards its apex. Two caudals from theAlcântara Formation of the Laje do Coringa site exhibit the samesuite of features, and have accordingly been referred to thisproblematic genus (Medeiros and Schultz, 2001a, 2002; Medeiroset al., 2007). Provided that, as seems likely, they do not pertainto Carcharodontosauridae (which is already represented in theAlcântara Formation fauna by teeth [see below]), the occurrenceof these vertebrae at Laje do Coringa constitutes yet anothersimilarity between the mid-Cretaceous vertebrate faunas ofnorthern Brazil and North Africa (Novas et al., 2005a; Novas,2009).

3.5.1. OrnithischiansUnless Sigilmassasaurus, or at least the caudal vertebral mor-

photype referred to this taxon, pertains to Ornithopoda (as sug-gested by Novas et al. (2005a), Canale et al. (2008), and Novas(2009)), no ornithischian dinosaur fossils are definitively knownfrom the Alcântara Formation. Avilla et al. (2003) reported a prob-able ornithopod caudal from ‘‘Aptian–Albian strata from the Ita-pecuru Formation’’. However, as stated above, the former‘‘Itapecuru Formation’’ is now known as the Itapecuru Group, andits Lower Cretaceous strata have been partitioned between thelower-middle Albian ‘‘Undifferentiated Unit’’ and the upperAlbian–Cenomanian Alcântara Formation (e.g., Rossetti andTruckenbrodt, 1997, 1999).

3.5.2. SauropodsReasonably abundant but disarticulated and often fragmentary

remains of sauropod dinosaurs have been recovered from theAlcântara Formation (Table 1). Price (1947) provided the first re-port of sauropod fossils from sediments that are now assigned tothis unit. Subsequent works (e.g., Medeiros and Schultz, 2001a,2002, 2004; Medeiros and Avilla, 2005; Freire et al., 2007;Medeiros et al., 2007) have considerably improved our understand-ing of the Alcântara sauropod fauna. Medeiros and Schultz (2001a,2002) referred a single middle or posterior caudal vertebral cen-trum from Laje do Coringa to the titanosauriform Astrodon sp.However, given the fragmentary nature of the specimen, and thefact that the current diagnosis of Astrodon (Carpenter and Tidwell,2005) does not include characters pertaining to the middle or pos-terior caudals, we concur with Medeiros et al. (2007:409) inregarding this Brazilian vertebra as Sauropoda indet.

The presence of rebbachisaurid diplodocoids in the AlcântaraFormation is indicated by disarticulated vertebral remains(Medeiros and Schultz, 2001a,b, 2003, 2004; Medeiros and Avilla,2005; Medeiros et al., 2007; Fig. 3D–F) and possibly by teeth (Fre-ire et al., 2007). Medeiros and Schultz (2004) referred many ofthese fossils to the genus Rayososaurus based on their resemblanceto corresponding skeletal elements of the Patagonian rebbachisau-rid ‘‘Rayososaurus’’ (now known as Limaysaurus) tessonei. More re-cently, however, Medeiros and Avilla (2005) and Medeiros et al.(2007) regarded this Alcântara Formation material as referableonly to Rebbachisauridae indet., an assessment that we followhere. Because rebbachisaurids occurred in at least Europe, SouthAmerica, and North Africa during the Early and mid-Cretaceous(e.g., Mannion, 2009; Apesteguía et al., 2010), the paleobiogeo-graphic utility of the Alcântara material is limited.

Represented by a fragmentary postcranial skeleton from EarlyCretaceous sediments exposed at the Mata locality in northernMaranhão, Amazonsaurus maranhensis is currently the most com-plete sauropod known from the Itapecuru Group and the only dipl-odocoid genus to be named from Brazil (Carvalho et al., 2003). Arecent, comprehensive phylogenetic analysis of Diplodocoidea(Whitlock, in press) suggests that Amazonsaurus may be a basalmember of this clade. Unfortunately, however, as is the case forthe crocodyliform Candidodon, the stratigraphic position of theonly known specimen of Amazonsaurus within the Itapecuru Grouphas not been resolved (Rossetti, pers. comm., 2010); as such, thissauropod may pertain to either the ‘‘Undifferentiated Unit’’ or theAlcântara Formation. As a result, we cannot regard this taxon as amember of the Alcântara fauna and as such do not consider it here.

Titanosauriform sauropods are represented in the AlcântaraFormation fauna by teeth and vertebrae (Medeiros and Schultz,2001a, 2002; Medeiros and Avilla, 2005; Freire et al., 2007;Medeiros et al., 2007). Medeiros and Schultz (2001a, 2002) referredtwo amphiplatyan or shallowly amphicoelous caudal centra fromLaje do Coringa (positioned in the middle-posterior and posteriorregions of the tail, respectively) to ‘‘Andesauridae’’; Medeiros

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et al. (2007:408) implicitly supported this referral. ‘‘Andesauridae’’was erected by Bonaparte and Coria (1993) to encompass thetitanosaurian genera Andesaurus, Argentinosaurus, and Epachtho-saurus. Nevertheless, as pointed out by Wilson and Upchurch(2003, p. 154), ‘‘. . .Andesauridae is based on primitive charactersthat by definition specify a paraphyletic group. Until taxa are foundsharing synapomorphies with Andesaurus, ‘Andesauridae’ will re-main an informal name.’’ Given that Bonaparte and Coria’s(1993) three ‘‘andesaurid’’ genera are now widely (e.g., Curry Rog-ers, 2005; Wilson, 2006; Filippi et al., in press) regarded as basalmembers of Titanosauria (i.e., non-lithostrotian titanosaurians orbasal lithostrotians), the Alcântara caudals may be those of basaltitanosaurians as well. However, the purportedly ‘‘andesaurid’’characters cited by Medeiros and Schultz (2001a, 2002) and Medei-ros et al. (2007) (amphiplatyan-amphicoelous articular surfaces, ananteriorly-placed neural arch, and other general resemblances tocomparably-positioned vertebrae of Andesaurus) also occur in mid-dle and posterior caudals of non-titanosaurian titanosauriforms.Accordingly, we regard these vertebrae from Laje do Coringa asTitanosauriformes indet.

Despite the uncertainty regarding the affinities of these ‘‘ande-saurid’’ vertebrae within Titanosauriformes, definitive titanosauri-an fossils are known from the Alcântara Formation. Described byMedeiros and Schultz (2001a, 2002), Freire et al. (2007), andMedeiros et al. (2007), these remains consist of dissociated teethand vertebrae. Perhaps the most anatomically informative of thesespecimens is an anterior caudal vertebra, preserving the completecentrum and the base of the neural arch, that was illustrated byMedeiros and Schultz (2001a, 2002) and Medeiros et al. (2007).The centrum of this specimen is strongly procoelous and its neuralarch is anteriorly positioned and anterodorsally inclined, charac-ters that confirm its placement within Titanosauria. Unusuallyamong titanosaurian anterior caudals, the centrum is transverselycompressed (such that it is substantially taller than wide) and sub-rectangular in anteroposterior view.

Medeiros (2002) referred a fragmentary vertebra from Laje doCoringa to the derived titanosaurian subclade Saltasaurinae; how-ever, Medeiros and Avilla (2005) reassigned this material toSauropoda indet.. Other previous studies (e.g., Medeiros andSchultz, 2001a, 2002; Freire et al., 2007; Medeiros et al., 2007) havenoted the resemblances of certain Alcântara Formation titanosau-rian fossils to particular genera (e.g., Aegyptosaurus, Malawisaurus).Nevertheless, due to the fragmentary nature of all describedAlcântara titanosaurian material, and the currently insufficientstate of knowledge regarding character distributions within theclade, we prefer to regard this material as Titanosauria indet. Sincetitanosaurians enjoyed an effectively global distribution during theLate Cretaceous (see, for example, Wilson (2006) or Molnar andWiffen (2007)), these fossils do not allow us to draw any precisepaleobiogeographic conclusions.

Additional sauropod fragments from the Itapecuru Group wererecently described by Castro et al. (2007). Recovered from expo-sures near the municipality of Coroatá in Maranhão, these fossilspertain to the lower-middle Albian ‘‘Undifferentiated Unit’’, notthe Alcântara Formation, and consequently are not taken into ac-count here.

3.5.3. TheropodsLike other kinds of dinosaurs, non-avian theropods are repre-

sented in the Alcântara Formation only by isolated teeth andfragmentary postcranial remains (Table 1). Price (1960) wasthe first to note the occurrence of theropod material in depositsthat modern geologists regard as belonging to the Alcântara For-mation. Decades later, multiple publications (e.g., Vilas Bôaset al., 1999; Medeiros and Schultz, 2001a, 2002; Medeiros,2006; Elias et al., 2007b; Medeiros et al., 2007; Machado et al.,

2009) have reported theropod fossils from this unit, revealing acarnivorous dinosaur assemblage that includes representativesof several major lineages. Some Alcântara theropod remains can-not presently be assigned to particular clades, however. Forexample, Medeiros et al. (2007) described a series of teeth fromthe Laje do Coringa bonebed that these authors likened to thoseof ‘‘Elaphrosaurus iguidiensis’’ from the Cretaceous of North Africa(Lapparent, 1960). Because ‘‘E. iguidiensis’’ is widely considered anomen dubium (e.g., Makovicky et al., 2004; Carrano and Samp-son, 2008), and because material assigned to this taxon is cur-rently regarded as that of one or more indeterminatetheropods (e.g., Carrano and Sampson, 2008; Smith et al.,2010), Medeiros et al. (2007) prudently regarded the Alcântarateeth as Theropoda indet. as well.

Medeiros and Schultz (2001a, 2002) and Medeiros et al. (2007)have compared an isolated ?caudal vertebral centrum from Laje doCoringa to a centrum from In Abangarit, Niger that Lapparent(1960) referred to the enigmatic North African theropod Baharia-saurus ingens. Indeed, in the captions to their illustrations of theBrazilian vertebra, Medeiros and Schultz (2002, Fig. 2E) andMedeiros et al. (2007: Figs. 3.3–3.4) referred to this specimen as‘‘cf. Baharijasaurus (sic)’’ and ‘‘cf Baharijasaurus (sic) ingens’’,respectively. However, in our view, this putative record ofBahariasaurus from the Alcântara Formation is highly doubtful.Although the Laje do Coringa centrum does resemble the vertebrafrom Niger (Lapparent, 1960, pl. V, Fig. 4), the possibility that eitherfossil actually pertains to Bahariasaurus is open to question. Theholotype and all specimens originally referred to B. ingens byStromer (1934) were destroyed in World War II (Rauhut, 1995;Nothdurft et al., 2002; Smith et al., 2006); consequently, compari-sons of more recently collected material to these remains arenecessarily limited to Stromer’s (1934) text and illustrations. Theonly confidently referred caudal vertebrae of Bahariasaurus illus-trated by Stromer (1934, pl. II, Figs. 11, 16, 25, 26)) do not appearclosely similar to either the Alcântara centrum or that from InAbangarit. For example, as also noted by Medeiros and Schultz(2001a, p. 213), the lateral pneumatic fossae (‘‘pleurocoels’’) inthe centrum of the Laje do Coringa vertebra are proportionallymuch larger than those of B. ingens (compare Stromer’s (1934) pl.II, Fig. 25 to Medeiros and Schultz’s (2002) Fig. 2e). We thereforeregard the possible record of Bahariasaurus in the Alcântara Forma-tion as extremely tentative. Indeed, we cannot even be certain thatthis Brazilian vertebra pertains to Theropoda. Interestingly, thespecimen shares several characters with dorsal and sacral centraof juvenile sauropods, including modest size, proportionally enor-mous lateral pneumatic fossae, and unfused neurocentral sutures(compare Medeiros and Schultz’s (2002) Fig. 2e to, for example,Carpenter and McIntosh’s (1994) Fig. 17.2H, Rauhut’s (1999)Fig. 11, Carpenter and Tidwell’s (2005) Figs. 3.5E–I, and/or Schwarzet al.’s (2007) Figs. 5D and E).

In contrast to Bahariasaurus, spinosaurids are unquestionablypresent in the fauna of the Alcântara Formation, with the teethof these aberrant theropods being particularly abundant in the unit(Medeiros and Vilas Bôas, 1999; Medeiros and Schultz, 2001a,2002; Medeiros, 2006; Medeiros et al., 2007; Furtado and Candeiro,2009a, 2009b). Furtado and Candeiro (2009b) recently assigned sixof these teeth to the spinosaurid subclade Baryonychinae. This re-port is of paleobiogeographic significance in that, although spino-saurids as a whole are well-documented from South America,Europe, Asia, and Africa (e.g., Milner et al., 2007; Buffetaut et al.,2008; Buffetaut, 2009; Furtado and Candeiro, 2009a; Hone et al.,2010), baryonychine fossils were previously known only from thelatter three continents. The Alcântara teeth therefore constitutethe first baryonychine occurrence in South America, and possiblythe geologically youngest record of this clade in the Gondwananlandmasses.

Carlos Roberto A. Candeiro et al. / Journal of African Earth Sciences 60 (2011) 79–92 85

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Most other spinosaurid teeth from the Alcântara Formation arereferable to the subclade Spinosaurinae. Some of these possiblypertain to the genus Spinosaurus itself (e.g., Medeiros and Schultz,2001a, 2002), while others may indicate the presence of a secondspinosaurine taxon in the fauna (Medeiros, 2006). Machado et al.(2009) recently reported what is probably the most anatomicallyand systematically informative spinosaurid fossil yet discoveredfrom the Alcântara Formation. Consisting of paired premaxillae,the specimen exhibits probable synapomorphies of Spinosaurinae(e.g., unserrated teeth) and represents a very large animal, seem-ingly comparable in size to Spinosaurus (see Dal Sasso et al.,2005). With a possible exception from the Barremian of Spain(Sánchez-Hernández et al., 2007), spinosaurines are known only

from mid-Cretaceous sediments of Brazil and North Africa, thusconstituting another notable faunal commonality between thetwo areas.

Carcharodontosaurid theropods are also known from theAlcântara Formation, represented by their distinctive teeth.Although most previous authors (Vilas Bôas et al., 1999; Medeirosand Schultz, 2001a, 2002; Medeiros et al., 2007) have referredthese teeth to Carcharodontosaurus, Candeiro and Martinelli(2005, p. 12) reinterpreted this material as pertaining to indetermi-nate carcharodontosaurids due to its lack of genus-level diagnosticfeatures. Since carcharodontosaurids were widespread during themid-Cretaceous (occurring in North America, Asia, South America,Africa, and possibly Europe [e.g., Brusatte et al., 2010; Ortega et al.,

Fig. 4. Geographic distribution of selected continental vertebrate taxa in northwestern Gondwana during the mid-Cretaceous. (Not all northwestern Gondwanan mid-Cretaceous localities are shown.) (Paleogeographical map modified from Maisey (2000).)

86 Carlos Roberto A. Candeiro et al. / Journal of African Earth Sciences 60 (2011) 79–92

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Tabl

e2

Dis

trib

utio

nof

cont

inen

talv

erte

brat

eta

xash

ared

betw

een

the

Alc

ânta

raFo

rmat

ion

and

sele

cted

Nor

thA

fric

anan

dSo

uth

Am

eric

anm

id-C

reta

ceou

sst

rati

grap

hic

unit

s.D

ata

sour

ces

are

asfo

llow

s:Ba

hari

yaFo

rmat

ion

(Str

omer

,193

6;Sm

ith

etal

.,20

01);

Chen

iniF

orm

atio

n(B

ento

net

al.,

2000

;A

nder

son

etal

.,20

07);

Kem

Kem

beds

(Dut

heil,

1999

;Ca

vin

etal

.,20

10);

Elrh

azFo

rmat

ion

(Taq

uet,

1976

;Se

reno

and

Brus

atte

,200

8);

Wad

iMilk

Form

atio

n(W

erne

r,19

94;

Wer

ner

and

Gay

et,1

997;

Rauh

ut,1

999)

;A

lcân

tara

Form

atio

n(t

his

pape

r,an

dre

fere

nces

here

in);

Mis

são

Vel

haan

dM

arfi

mfo

rmat

ions

(Mai

sey,

2000

);A

çuFo

rmat

ion

(Ass

ine,

1992

);Sa

ntan

aFo

rmat

ion

(Mai

sey,

2000

;M

arti

ll,20

07);

Loha

nCu

raFo

rmat

ion

(Mar

tine

lliet

al.,

2007

).

Nor

thA

fric

aSo

uth

Am

eric

a

Egyp

tTu

nis

iaM

oroc

coN

iger

Suda

nB

razi

lA

rgen

tin

a

Bah

ariy

aFm

.(C

enom

ania

n)

Ch

enin

iFm

.(A

ptia

n?–

Alb

ian

)

Kem

Kem

beds

(Cen

oman

ian

)

Elrh

azFm

.(A

ptia

n–

Alb

ian

)

Wad

iM

ilk

Fm.

(Cen

oman

ian

)

Alc

ânta

raFm

.(A

lbia

n–

Cen

oman

ian

)

Mis

são

Vel

ha

Fm.

(Apt

ian

)

Mar

fim

Fm.

(Hau

teri

vian

–B

arre

mia

n)

Açu

Fm.

(Alb

ian

–C

enom

ania

n)

San

tan

aFm

.(A

lbia

n)

Loh

anC

ura

Fm.(

Apt

ian

–A

lbia

n)

Can

dele

ros

Fm.

(Cen

oman

ian

)

Hyb

odon

tida

e�

��

��

��

��

Myl

ioba

tida

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�O

ncho

pris

tis

��

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rtsc

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thys

��

�Py

cnod

onti

form

es�

��

��

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dote

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odus

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rica

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urus

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tan

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ria

��

��

��

�Ba

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rus

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?B

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inos

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Man

irap

tora

��

��

Carlos Roberto A. Candeiro et al. / Journal of African Earth Sciences 60 (2011) 79–92 87

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2010]), the occurrence of these large-bodied predatory dinosaursin the Albian–Cenomanian of northeastern Brazil is not surprising.Finally, Vilas Bôas (1999) and Elias et al. (2007b) described teethpurportedly belonging to velociraptorine dromaeosaurids fromthe Alcântara Formation. Nevertheless, Gianechini et al. (in press)recently reinterpreted this material as Maniraptora indet., asystematic interpretation that we accept here.

4. Paleobiogeographic implications of the Alcântara Formationvertebrate fauna

As noted by many previous authors (e.g., Medeiros and Schultz,2001a, 2002; Medeiros et al., 2007), numerous similarities existbetween the continental vertebrate assemblages of the AlcântaraFormation of Brazil and multiple mid-Cretaceous units of NorthAfrica, supporting the hypothesis that faunal interchange occurredbetween western Gondwanan landmasses during the Early (andpossibly the early Late) Cretaceous. This interpretation is also sup-ported by the presence of mid-Cretaceous abelisauroid, carchar-odontosaurid, rebbachisaurid, and titanosaurian dinosaurs in bothNorth Africa (e.g., Stromer, 1931; Lavocat, 1954; Russell, 1996;Sereno et al., 1996, 1999, 2004; Smith et al., 2001; Sereno andBrusatte, 2008; Contessi, 2009) and Argentina (e.g., Calvo andSalgado, 1995; Coria and Salgado, 1995; Bonaparte, 1996; Lamannaet al., 2002; Martínez et al., 2004; Salgado et al., 2004; Novas et al.,2005b; Coria and Currie, 2006). Among the many African mid-Cretaceous units that preserve faunas similar to that of theAlcântara Formation are the Bahariya Formation of Egypt, theChenini Formation of Tunisia, the Elrhaz Formation of Niger, theWadi Milk Formation of Sudan, and the Kem Kem beds of Morocco(Fig. 4; Table 2). With the possible exceptions of the Chenini andElrhaz formations, which have been dated to the Aptian?–Albianand the Aptian–Albian, respectively, all of these units are approxi-mately equivalent in age to the Alcântara Formation (Pedrãoet al., 1993; Werner, 1994; Rossetti, 1997; Rossetti and Truckenb-rodt, 1997; Smith et al., 2001; Anderson et al., 2007; Sereno andBrusatte, 2008; Cavin et al., 2010). As is also the case for thisBrazilian formation, each of these African units preserves remainsof many of the following low-level (i.e., ‘‘family-level’’ or less inclu-sive) continental vertebrate taxa: Hybodontidae, Myliobatidae,Onchopristis, Bartschichthys, Lepidotes, Stephanodus, Mawsonia,Arganodus, Ceratodus africanus, Pholidosauridae, Sigilmassasaurus,Rebbachisauridae, Bahariasaurus, Baryonychinae, Spinosaurinae,Carcharodontosauridae, and possibly Ceratodus humei (though, asnoted above, the alleged pre-Campanian African records of this dip-noan species were questioned by Churcher and De Iuliis (2001)).

Several other mid-Cretaceous Brazilian units preserve a conti-nental vertebrate fauna similar to that of the Alcântara Formation.For example, the fauna of the Albian Santana Formation includesHybodontidae, Lepidotes, Mawsonia, and the spinosaurine spino-saurid Irritator challengeri (Maisey, 2000; Sues et al., 2002; Martill,2007), while the Aptian Missão Velha Formation has yielded thesethree fish taxa plus Ceratodus (Brito et al., 1994; Maisey, 2000).Additionally, the Albian–Cenomanian Açu Formation has producedLepidotes remains, while the Hauterivian–Barremian Marfim For-mation yields this semionotiform genus plus Hybodontidae andMawsonia (Fig. 4, Table 2). Thus, the presence of closely relatednonmarine vertebrate taxa in middle Cretaceous deposits of NorthAfrica and Brazil (Sereno et al., 1996, 1998, 1999, 2004; Forster,1999; Candeiro et al., 2004, 2006; Holtz et al., 2004; Wilson,2006; Sereno and Brusatte, 2008; this study), as well as paleogeo-graphic reconstructions, supports the hypothesis that a land routeconnecting the western Gondwanan landmasses may have per-sisted until the early Late Cretaceous, possibly in the equatorial re-

gion (Petri, 1987; Reyment and Dingle, 1987; Pitman et al., 1993;Pletsch et al., 2001; Tello Saenz et al., 2003; Guedes et al., 2005).

Interestingly, certain continental vertebrate taxa that occur inboth the Alcântara Formation and approximately coeval NorthAfrican strata, such as Bartschichthys, Mawsonia, Arganodus,Ceratodus africanus, Sigilmassasaurus, and Spinosauridae, have notbeen discovered in contemporaneous deposits in Argentina. (Weconcur with Buffetaut (2009) in regarding a recent report of aspinosaurid tooth from the Turonian of Patagonia [Salgado et al.,2009] with caution, as this tooth exhibits morphologies that areunknown in definitive spinosaurid taxa.) Similarly, mid-Cretaceousterrestrial tetrapod faunas from Argentina include several taxa thathave not been found in Brazil or North Africa, such as chelid turtles(e.g., Lapparent de Broin and de la Fuente, 2001), dicraeosauridsauropods (Salgado and Bonaparte, 1991), and megaraptoran(Novas, 1998; Benson et al., 2009) and unenlagiine (e.g., Makovickyet al., 2005; Gianechini et al., in press) theropods. These differencesin faunal composition suggest that the mid-Cretaceous continentalvertebrate assemblages of northern South America may have beenmore similar to coeval North African faunas than to those fromsouthern South America.

Although further paleontological discoveries may falsify thisinterpretation, the distinction observed between the northernSouth American/North African and southern South American fau-nal assemblages supports the idea that paleogeographic, paleocli-matic, and/or paleoenvironmental barriers existed betweennorthern South America/North Africa and southern South Americaduring the middle stages of the Cretaceous. Geological features ofSouth America have previously been hypothesized to have playeda role in acting as physical barriers between the northern andsouthern portions of the continent (Arid, 1977; Garcia et al.,2005; Candeiro et al., 2006; Candeiro, 2010). One of the most plau-sible candidates is the Paranaíba High (=Alto Paranaíba Arch), anelevated ridge between central and southern South America thatcould have acted as a structural barrier preventing or hinderingfaunal exchange during the mid-Cretaceous. The emplacement ofthis barrier could have resulted in the divergence of northernand southern South American faunas and in the formation of dis-tinct paleobiogeographic provinces in both areas of the continent.

5. Conclusions

The continental vertebrate fauna of the Albian–CenomanianAlcântara Formation of northeastern Brazil shows strongsimilarities with penecontemporaneous assemblages from NorthAfrica. The co-occurrence of up to eight fish taxa (Onchopristis,Bartschichthys, Lepidotes, Stephanodus, Arganodus, Ceratodusafricanus, Mawsonia, and possibly Ceratodus humei) and as manyas seven low-level terrestrial archosaur clades (Sigilmassasaurus,Rebbachisauridae, Baryonychinae, Spinosaurinae, Carcharodonto-sauridae, possibly Pholidosauridae, and doubtfully Bahariasaurus)in the Alcântara Formation and coeval African deposits indicatesthat either a significant vicariant event took place when NorthAfrica and northern South America separated, or that a terrestrialcorridor at least intermittently connected these areas until the lateEarly or early Late Cretaceous. Interestingly, many of the faunalcommonalities between Brazil and North Africa are not sharedwith southern South America, which is instead characterized bylargely distinct continental vertebrate assemblages. These resultssuggest that, although mid-Cretaceous faunal interchange wasprobably possible between northern South America and North Afri-ca, paleogeographic, paleoclimatic, and/or paleoenvironmentalbarriers may have impeded faunal dispersal between northernand southern South America until the later stages of the LateCretaceous.

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Acknowledgments

We thank D.M. Martill (University of Portsmouth, UK), F.Lapparent de Broin (Muséum National d’Histoire Naturelle,France), D.F. Rossetti (Museu Paraense Emílio Goeldi, Brazil), andA.G. Martinelli (Universidade de Uberaba, Brasil) for providingvaluable comments in their reviews of earlier versions of this man-uscript. C.R.A. Candeiro was supported by a Jurassic Foundationgrant and Conselho Nacional de Desenvolvimento Científico eTecnológico (CNPq)/ Bolsista de Produtividade em Pesquisa.

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