In the Cretaceous of South America

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dos Anjos Candeiro, Carlos Roberto; Martinelli, Agustín Guillermo ABELISAUROIDEA AND CARCHARDONTOSAURIDAE (THEROPODA, DINOSAURIA) IN THE CRETACEOUS OF SOUTH AMERICA. PALEOGEOGRAPHICAL AND GEOCRONOLOGICAL IMPLICATIONS Sociedade & Natureza, vol. 17, núm. 33, diciembre, 2005, pp. 5-19 Universidade Federal de Uberlândia Uberlândia, Minas Gerais, Brasil

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How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative Abelisauroidea and carchardontosauridae (theropoda, dinosauria) in the cretaceous of south america. Paleogeographical and geocronological implications Carlos Roberto dos Anjos Candeiro, Agustín Guillermo Martinelli

ABELISAUROIDEA AND CARCHARDONTOSAURIDAE (THEROPODA, DINOSAURIA) IN THE CRETACEOUS OF SOUTH AMERICA. PALEOGEOGRAPHICAL AND GEOCRONOLOGICAL IMPLICATIONS

Abelisauroidea e Carcharodontosauridae (Theropoda, Dinosauria) na América do Sul durante do Cretáceo. Implicações paleogeograficas e geocronologicas

Carlos Roberto dos Anjos Candeiro Depart. Geologia, Universidade Federal do Rio de Janeiro [email protected], [email protected] Agustín Guillermo Martinelli Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Av. Ángel Gallardo 470, C1405DJR, Buenos Aires, Argentina [email protected]

Artigo recebido em 18/10/2004 e aceito para publicação em 31/08/2005

ABSTRACT: In this contribution an up-to-date list of abelisauroid ceratosaurians and carcharodontosaurid allosaurians recognized in South America is presented. Abelisauroids and carcharodontosaurids in South America show rich species diversity and a wide range of temporal and geographical distribution. At least eight formally described species of Abelisauroidea are recognized in Argen- tina and only one in Brazil; in contrast, only one species of Carcharodontosauridae is known for all South America. The fossil record of abelisauroids and carcharodontosaurids in South America shows a dominance of abelisauroids in the upper late Cretaceous, while the dominance of carcharodontosaurids as large predators was during the Cenomanian-Turonian. Although knowledge of the evolution of Abelisauroidea and Carcharodontasauridae in South America, as well as in the rest of Gondwana is still far for being complete, intensive explorations in recent years have provided greater insight into the composition of theropod faunas in the Cretaceous of Gondwana.

Keywords: Abelisauroidea; Carcharodontosauridae; Cretaceous; South America.

RESUMO: Na presente contribuição é apresentada uma lista atualizada dos ceratossaurios abelissauroideos e dos alossaurios carcarodontosaurídeos sul-americanos. Os abelissauródeios e carcaro- dontossaurídeos possuem uma grande diversidade específica e com uma grande distribuição temporal e geográfica. São reconhecidas oito espécies de Abelisauroidea na Argentina e somente uma no Brasil, em contraste, só é conhecida uma espécie de Carcharodontosauridae na América do Sul. O registro fóssil denota uma importância dos abelissauroideos no Cretáceo mais tardio, contudo o predomínio dos carcarodontossaurídeos como os grandes predadores somente ocorreram durante o Cenomaniano-Turoniano. Apesar da escassez do conhecimento da evolução dos Abelisauroidea e Carcharodontasauridae na América do Sul como o resto do Gondwana, a intensiva exploração nos últimos anos tem produzido um grande incremento de dados e brindado

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5 Abelisauroidea and carchardontosauridae (theropoda, dinosauria) in the cretaceous of south america. Paleogeographical and geocronological implications Carlos Roberto dos Anjos Candeiro, Agustín Guillermo Martinelli

com novas luzes para a melhor compreensão da composição faunística desses terópodes do Cretáceo do Gondwana.

Palavras-chave: Abelisauroidea; Carcharodontosauridae; Cretáceo; América do Sul.

1 INTRODUCTION groups were unearthed with increasing frequency in recent decades. The abelisauroid ceratosaurians and carcharodontosaurid allosaurians are the best-known large 2 OBJECTIVES AND METHODOLOGY predatory theropod dinosaurs of the Cretaceous of Gondwana. Abelisauroids have been recognized in In this contribution, we present an up-to- several localities of Argentina (BONAPARTE, date list of Abelisauroidea and Carcharodonto- 1991A, 1996; BONAPARTE; NOVAS, 1985; BO- sauridae taxa recognized in the Cretaceous of South NAPARTE; POWELL, 1980; BONAPARTE; NO- America. Furthermore, we evaluate the paleogeo- VAS; CORIA, 1990; CORIA; CHIAPPE; DIN- graphical and chronostratigraphical distribution of GUS, 2002; LAMANNA; MARTINEZ; SMITH, both groups in South America during the Cretaceous 2002; MARTINEZ ET AL. 1986; RAUHUT ET as the result of Gondwanan plate tectonics. AL. 2003), Brazil (BERTINI, 1996; BITTEN- COURT; KELLNER, 2002; CANDEIRO ET. The data on abelisauroid and carcharo- 2004A,B; KELLNER; CAMPOS, 2002;), India dontosaurid species for the present review are (CHATTERJEE, 1978; HUENE; MATLEY, 1933; mostly based in literature sources as well as in the WILSON ET AL. 2003; NOVAS; AGNOLIN; direct observation of the specimens. The term Abe- BANDYOPADYAY, 2004), and Madagascar (LA- lisauroidea is used sensu Bonaparte (1991b), No- VOCAT, 1955; CARRANO; SAMPSON; FORS- vas (1992), Carrano, Sampson, Forster (2002), and TER, 2002; SAMPSON; CARRANO; FORSTER, Wilson et al. (2003), as the clade of ceratosaurians 2001; SAMPSON ET AL. 1998;). Beside Gondwa- that includes Abelisauridae plus Noasauridae. The nan regions, abelisauroids are also documented in term Carcharodontosauridae follows the definition France (BUFFETAUT ET AL. 1988) and Spain of Coria and Currie (2003) as a family of Allo- (ASTIBIA ET AL. 1990). sauroidea that comprises all descendants of the most recent common ancestor of Giganotosaurus and The record of carcharodontosaurids is less Carcharodontosaurus. abundant and mainly restricted to Argentina (CO- RIA; SALGADO, 2000; VICKERS-RICH ET AL. For the stratigraphic units in the Neuquén 1999) and Continental Africa (Morocco; SERENO Basin we follow the arrangement proposed by ET AL. 1996; STROMER, 1915); nevertheless, a Leanza (1999), and Leanza and Hugo (2001). For few isolated tooth remains were also collected in other stratigraphic units we follow: Southern Basins, the upper Cretaceous beds of Brazil (CANDEIRO Page et al. (1999); Salta Basin, Salfity and Marquilla ET AL., 2002, 2004A,B). (1999); San Luis Basin, Medeiros and Schultz (2002), and Bauru Basin, Fernandes and Coimbra (1996). The fossil record of abelisauroids and carcharodontosaurids in South America shows rich species diversity and a wide range of temporal and geographical distribution. Thus, remains attributed to both

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3 PALEOGEOGRAPHY AND UPPER Finally, it is worth mentioning that during the CRETACEOUS TECTONIC EVENTS: late Cretaceous South American areas shared the A BRIEF REVIEW same paleogeographical history; however during the Maastrichtian the southern-most part of South During most of the Mesozoic the continen- America was affected by the Maastrichtian-Danian tal masses were principally arranged in two sepa- marine transgression, which separated the Argen- rated continents: Laurasia in the north and Gondwana tinean and Brazilian basins (Salta and Neuquén Ba- in the south (BONAPARTE, 1986; FORSTER, sins in Argentina and Bauru Basin in Brazil) 1999; HAY ET AL, 1999; PITMAN ET AL, 1993). (PASCUAL; ORTIZ JAUREGUIZAR; PRADO, By the end of the Mesozoic these masses started to 1996; ZAMBRANO, 1987). This disconnection separate into their modern conformation. could have caused same faunal differentiation between the northern and southern parts of South The separation of Gondwana started in the America. late Jurassic as a product of rifting along the present South America and Africa borders. This separation 4 ABELISAUROIDEA AND continued, by the Cretaceous times the South At- CARCHARODONTOSAURIDAE lantic Ocean started to form, and Gondwana split BACKGROUND into the Eastern and Western landmasses (PITMAN ET AL., 1993). Eastern Gondwana comprised of Abelisauroidea is a ceratosaurian clade that Africa, India-Madagascar; whereas Western includes small to large-sized predators that evolved Gondwana included South America, Antarctica, and during the Cretaceous in Gondwanan landmasses. Oceania. By the late Cretaceous India separated The group is composed of two families, Abelisauridae from Madagascar and started the northwest move- and Noasauridae, which are widely distributed in both ments that lead to a collision with Eurasia. Western temporal and geographical rank. Abelisauridae was Gondwanan landmasses were still connected by the created by Bonaparte and Novas (1985) to accom- beginning of the Tertiary (HAY ET AL., 1999; modate Abelisaurus comahuensis. After this first PASCUAL; ORTIZ JAUREGUIZAR;PRADO, discovery, knowledge of the group was increased 1996; SAMPSON ET AL. 1998). considerably by new discoveries, and by the rein- terpretation of fragmentary and isolated remains; Two main paleogeographic features allowed which permitted the recognizion of an important ra- abelisaurids and carcharodontosaurids to attain a diation on Gondwanan landmasses during the Cre- widespread distribution during Cretaceous times. taceous including several adaptative morphotypes These were the Gondwanan Bridge that linked (BONAPARTE, 1991B). India-Madagascar and the South America-Oceania platforms, and the North bridge, connecting north Noasauridae was erected by Bonaparte and Africa and Europe (e.g. HAY ET AL. 1999). In Powell (1980) to include a single species, Noasaurus addition, South America and North America were leali from Lecho Formation (Salta Province). Sub- sporadically connected through Central America at sequently this family was related to abelisaurids the end of the Cretaceous (BONAPARTE, 1986; (BONAPARTE, 1991B; BONAPARTE; NOVAS; BONAPARTE ET AL. 1984). Thus, these transient CORIA, 1990). geographic connections between West Gondwana and other landmasses during the late Cretaceous Beside South America (Table 1), abelisaurid permitted faunal interchanges that directly influenced species were also recognized in India: Indosuchus the faunal composition of the terrestrial ecosystems raptorius Huene and Matley, Indosaurus matleyi (BONAPARTE, 1991B; SAMPSON ET AL. 1998). Huene (BONAPARTE, 1986, 1991B; BONA- PARTE; NOVAS; CORIA, 1990; CHATTERJEE,

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1978; CHATTERJEE; RUDRA, 1996; NOVAS; and Giganotosaurus carolinii Coria and Salgado AGNOLIN; BANDYOPADYAY, 2004), Lameta- from Argentina (CORIA; SALGADO, 1995; SE- saurus indicus Matley (WILSON ET AL. 2003), RENO ET AL. 1996) were included. Most recent and Rajasaurus narmadensis Wilson, Sereno, research supports the hypothesis that the Carcha- Srivastava, Bhatt, Khosla, and Sahni (WILSON ET rodontosauridae are allosaurians; nevertheless, it is AL. 2003); Madagascar: Majungatholus atopus noteworthy that Novas (1997) found several derived Lavocat (SAMPSON ET AL. 1998); Niger: Rugo- features shared between carcharodontosaurids and pus primus and Spinostropeus gautieri Sereno, abelisauroids suggesting a close phylogenetic affi- Wilson, Conrad (SERENO. WILSON; CONRAD, nity with the latter. 2004); Sereno, Duthiel, Larochene, Larsson, Lyon, Magwene, Sidor, Varicchio and Wilson: Deltadro- 5 ABELISAUROIDEA AND meus agilis (SERENO ET AL. 1996); and France: CARCHARODONTOSAURIDAE Tarascosaurus salluvicus Le Loeuff and Buffetaut SPECIES (LE LOEUFF; BUFFETAUT, 1990). Noasaurids discovered in India include Laevisuchus indicus The following is a brief, annotated list of Huene (CARRANO; SAMPSON; FORSTER, abelisauroid and carcharodontosaurid taxa from the 2002; NOVAS; AGNOLIN; BANDYOPADYAY, Late Cretaceous of South America (Tables 1 and 2004); and from Madagascar Masiakasaurus 2). The Argentinean and Brazilian species and knopfleri Sampson, Carrano and Forster (CARRA- records are in chronological order (from early to late NO; SAMPSON; FORSTER, 2002; SAMPSON; forms). CARRANO; FORSTER, 2001). 5.1 Abelisauroidea Other non-South American abelisauroids of uncertain affinities are Jubbulpuria tenuis Huene 5.1.1 Argentina and Matley, Ornithomimoides mobilis Huene and Matley, O.? barasimlensis Huene and Matley, Ligabueino andensi Bonaparte, 1996. Ligabueino Coeluroides largus Huene and Matley, and andensi comes from the La Amarga Formation Dryptosauroides grandis Huene and Matley, from (Hauterivian-Barremian), La Amarga locality, south- India nomen nudum (NOVAS;AGNOLIN; ern Neuquén Province (BONAPARTE, 1996). BANDYOPADYAY, 2004); Elaphrosaurus bam- Ligabueino is a very small theropod that is based bergi Janensch from Tanzania (CARRANO; on; a cervical and two dorsal vertebral neural arches, SAMPSON; FORSTER, 2002; WILSON ET AL. right ilium, two incomplete pubis, a left femur, and 2003); and Genusaurus sisteronis Accarie, Beau- two pedal phalanges. This species is considered to doin, Dejaxm Friès, Michard and Taquet from France be a basal abelisauroid (BONAPARTE, 1996) with (ACCARIE ET AL. 1995). possible noasaurid affinities (CARRANO; SAMP- SON; FORSTER, 2002). Carcharodontosauridae is a family of allosauroid theropods, which is composed of the larg- Ilokelesia aguadagrandensis Coria and Salgado, est theropod species (CORIA; SALGADO, 2000; 2000. Ilokelesia aguadagrandensis is from the Table 2). The first recognized member of this fam- Huincul Formation (Cenomanian-Turonian; Río ily was Carcharodontosaurus saharicus Stromer Limay Subgroup) in the Aguada Grande locality, from Continental Africa (Algeria, Niger, Tunisia, eastern Neuquén Province. Ilokelesia is based on Sudan and Morroco; SERENO ET AL. 1996; STRO- some skull bones, two cervical, one dorsal, and five MER, 1915; 1931); then later followed Acrocantho- articulated vertebrae, fragmentary ribs, some haemal saurus atokensis Stoval and Langston from United arches, and some pedal phalanges (CORIA; SAL- States (HOLTZ, 2000; SERENO ET AL., 1996), GADO, 2000). This species is considered to be a

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8 Abelisauroidea and carchardontosauridae (theropoda, dinosauria) in the cretaceous of south america. Paleogeographical and geocronological implications Carlos Roberto dos Anjos Candeiro, Agustín Guillermo Martinelli basal abelisaurid (CORIA; SALGADO, 2000). saurus was unearthed from the same mudstone and siltstone strata that has produced several remains Xenotarsosaurus bonapartei Martínez, Giménes, of fossilized nests, eggs, and embryos of still unde- Rodríguez, and Bochatey, 1986. Xenotarsosaurus termined but possible titanosaur neosauropods bonapartei comes from the Bajo Barreal Forma- (CHIAPPE; CORIA, 2000). Aucasaurus and Car- tion (middle Cenomanian-late Turonian) in the notaurus (see below) were nested together in the Sarmiento Department, in the southern part of the Tribe Carnotaurini as the most derived members of Chubut Province. Xenotarsosaurus is a medium- the family Abelisauridae (CORIA; CHIAPPE; DIN- sized abelisaurid based on two incomplete cervico- GUS, 2002). dorsal vertebrae, a right femur, tibia and fibula, and a fused astragalus-calcaneous (MARTÍNEZ ET Carnotaurus sastrei Bonaparte, 1985. Carnotau- AL. 1986). rus sastrei from La Colonia Formation (Maastri- chtian) of Chubut Province is, without a doubt, the Velocisaurus unicus Bonaparte, 1991a. Veloci- best-known abelisaurid. The skeleton is almost com- saurus unicus comes from the Bajo de la Carpa plete (with exception of the distal portion of the tibiae, Formation (Santonian; Río Colorado Subgroup) of both fibulae, the hind feet, and the medial and distal Neuquén city, Neuquén Province. This species is a end of the tail), and exceptionally well preserved small theropod, which is only known from the right including the skin impressions (BONAPARTE; tibia and almost complete right foot. Velocisaurus NOVAS; CORIA, 1990). After the original taxo- was tentatively interpreted as a Ceratosauria (BO- nomic designation of Carnotaurus (BONAPARTE, NAPARTE, 1991A); in addition some authors have 1985), a complete description of the whole skeleton found a few features in common with noasaurids was performed by Bonaparte, Novas, Coria (1990); (CARRANO; SAMPSON; FORSTER, 2002). Ne- revealing a lightly-built theropod about 7 meters long, vertheless, new materials should confirm the taxo- with the skull high and short bearing two prominent nomic status of this taxon. frontal horns (Fig. 3).

Abelisaurus comahuensis Bonaparte and Novas, Noasaurus leali Bonaparte and Powell, 1980. 1985. Abelisaurus comahuensis comes from the Noasaurus leali comes from the Lecho Formation Anacleto Formation (Santonian-Early Campanian; (Maastrichtian), southern Salta Province. Noasaurus Río Colorado Subgroup; HEREDIA; SALGADO, is a small theropod based on; a complete left max- 1999) in the Río Negro Province. Abelisaurus is a illa, a right quadrate, a cervical neural arch, a dorsal medium to large (6 to 7 m long) theropod that is only vertebral body, cervical ribs, the right metatarsal II, known from a partial skull of about 0.85 meters (Figs. a pedal phalanx, and an ungual phalanx of the foot. 1 and 2). Since the discovery of this species, the di- Because of derived features found in the maxilla versity and phylogenetic relationships of abelisauroids and in the cervical neural arch, Bonaparte and Powell have begun to be more understood, it has also pro- (1980) erected the family Noasauridae, which pre- vided new insights on the biogeographic distribution viously was regarded as part of the gondwanan ra- of this gondwanic lineage of theropods (BONA- diation of abelisauroids (BONAPARTE, 1991B; PARTE, 1986, 1991B). NOVAS, 1997). In addition, closely related forms to Noasaurus were reported from the Maastrichtian Aucasaurus garridoi Coria, Chiappe, and Dingus, of Madagascar (CARRANO; SAMPSON; FORS- 2002. Aucasaurus garridoi was recently described TER, 2002; SAMPSON; CARRANO; FORSTER, from an almost complete skeleton from the Anacleto 2001). Formation (early Campanian; Río Colorado Sub- group) from Auca Mahuevo, eastern Neuquén Prov- Other abelisauroid remains. In the Cerro Barcino ince (CORIA; CHIAPPE; DINGUS, 2002). Auca- Formation (Hauterivian-Barremian) at Cerro Chivo,

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9 Abelisauroidea and carchardontosauridae (theropoda, dinosauria) in the cretaceous of south america. Paleogeographical and geocronological implications Carlos Roberto dos Anjos Candeiro, Agustín Guillermo Martinelli northern Chubut Province, two isolated caudal ver- fibula. Pycnonemosaurus is a medium to large-sized tebrae of a medium sized abelisaurid were reported theropod of approximately 6 to 7 meters in length, (RAUHUT ET AL. 2003). Furthermore, an isolated regarded as an abelisaurid because it share several but nearly complete left maxilla with some teeth was derived features with other well known member of discovered in the Bajo Barreal Formation (middle the group, such as Ilokelesia, Carnotaurus, and Cenomanian-late Turonian) in the Sierra de San Aucasaurus (KELLNER; CAMPOS, 2002). Bernardo, Chubut Province, and subsequently was assigned to the subfamily Carnotaurinae (LA- Abelisaurid remains. The first abelisaurid from MANNA; MARTINEZ; SMITH, 2002). Brazil was discovered by Bertini (1996). This mate- rial consists of a partial premaxilla with a tooth. The 5.1.2 Brazil specimen was discovered in Adamantina Formation sediments (Bauru Group, Turonian-Santonian) in São Pycnonemosaurus nevesi Kellner and Paulo (Santo Anastácio area) and Minas Gerais Campos, 2002. Pycnonemosaurus nevesi comes (Prata area) States. Furthermore, Candeiro (2002) from the Adamantina Formation (Turonian- and Candeiro et al. (2002, 2004a,b) reported Santonian) of the Cambebe area, Mato Grosso State, abelisaurid teeth from the states of Minas Gerais and represents the first Brazilian abelisaurid formally (Peirópolis area) and São Paulo (Alfredo Marondes, described (KELLNER; CAMPOS, 2002). This spe- Flórida Paulista, Santo Anastácio) in the Adamantina cies is based on five teeth (BITTENCOURT; and Marília Formations (Late Maastrichtian) of the KELLNER, 2002), parts of seven caudal vertebrae, Bauru Group. These remains are very fragmentary, fragments of ribs, and an incomplete right tibia and making an accurate identification difficult.

Figure 1 Skulls of Abelisauridae. A, Abelisaurus comahuensis Bonaparte and Novas (modified from BONAPARTE AND NOVAS, 1985); B, Carnataurus sastrei Bonaparte (modified from BONAPARTE, NOVAS, CORIA, 1990). Scale bar represents 10 cm.

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Table 1 Abelisauroidea records from South America.

Abelisauroidea Country Stratigraphy Reference Selected Ligabueino andensi Argentina La Amarga, Barremian Bonaparte (1996) Llokelesia aguadagrandensis Argentina Huincul Fm., Cenomanian-Turonian Coria and Salgado (2000) Xenotarsosaurus bonapartei Argentina Bajo Barreal Fm., middle Martinez et al. (1986) Cenomanian-Late Turonian Velocisaurus unicus Argentina Bajo de La Carpa, Turonian Bonaparte (1991a) Pycnonemosaurus nevéis Brazil Adamantina Fm., Turonian-Santonian Campos and Kellner (2002) Abelisaurus comahuensis Argentina Anacleto Fm., Santonian-Early Campanian Bonaparte (1991b) Aucasaurus garridoi Argentina Anacleto Fm., Santonian-Early Campanian Coria, Chiape and Dingus (2002) Carnotaurus sastrei Argentina La Colonia Fm., Maastrichtian Bonaparte, Novas and Coria. (1990) Noasaurus leali Argentina Lecho Fm., Maastrichtian Carrano, Sampson, Forster (2002) Carnotaurinae Argentina Bajo Barreal Fm., middle Lamanna, Martinez and Smith Cenomanian-Late Turonian (2002) Abelisauridae Argentinal Cerro Barcino Fm., Hauterivian-Barremian Rauhut et al. (2003) Abelisauridae Brazil Adamantina Fm., Turonian-Santonian; Candeiro et al. (2004a,b) Marília Fm., Late Maastrichtian

5.2 CARCHARODONTOSAURIDAE nian-Maastrichtian) of the Salitral Ojo de Agua locality, Río Negro Province. This species consists of 5.2.1 Argentina only a distal portion of a right femur and a complete right tibia. Quilmesaurus was originally regarded Giganotosaurus carolinii Coria and Salgado, 1995. as a non-ceratosaur theropod possibly related to Giganotosaurus carolinii comes from the Candele- Giganotosaurus (CORIA, 2001). Later, Quilme- ros Formation (Cenomanian, Río Limay Subgroup) saurus was considered an abelisauroid (Kellner and of the El Chocón locality, Neuquén Province. This Campos, 2002), but recently Juárez Valieri, Fiorelli species is considered to be the largest theropod in and Cruz (2004) regarded it as nomen dubium due the world with a skull length of about 1.7 meters to the lack of derived features needed to construct (CALVO; CORIA, 2000; CORIA; SALGADO, a robust taxon. 1995) (Figs. 3 and 4). Giganotosaurus is based on approximately 70% of the skeleton, including sev- Carcharodontosaurid remains. Isolated teeth and eral skull bones, teeth, and most of the postcranial skeletal remains are recognized from the Cerro elements (CORIA; CURRIE, 2003; CORIA; SAL- Barcino (Hauterivian-Barremian; Chubut Province; GADO, 1995;). In addition, isolated teeth and a frag- NOVAS; DE VALAIS, 2001; VICKERS-RICH ET ment of a left dentary from the same stratigraphic AL. 1999;), Mata Amarilla (Turonian; Chubut Prov- horizons of the holotype of Giganotosaurus were ince; NOVAS ET AL. 1999), Anacleto (Santonian; described and assigned to the genus (CALVO, 1999; Neuquén Province; ALCOBER ET AL. 1998), Los DE VALAIS; APESTEGUÍA, 2001). Alamitos (Late Campanian; Río Negro Province; APESTEGUÍA ET AL., in press.), and Allen (Cam- Quilmesaurus curriei Coria, 2001. Quilmesaurus panian-Maastrichtian; Río Negro Province; MARTI- curriei comes from the Allen Formation (Campa- NELLI; FORASIEPI, in press) Formations.

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11 Abelisauroidea and carchardontosauridae (theropoda, dinosauria) in the cretaceous of south america. Paleogeographical and geocronological implications Carlos Roberto dos Anjos Candeiro, Agustín Guillermo Martinelli

Figure 2 Life reconstruction of Abelisaurus comahuensis Bonaparte and Novas (left) and Giganotosaurus carolinii Coria and Salgado (right) performed by the paleoartist Jorge Blanco. Both species are in scale.

5.2.2 Brazil dontosaurid teeth have been found in the Ceno- manian-Turonian beds of Maranhão State in recent Carcharodontosaurid remains. Recently carcha- years. Medeiros and Shultz (2002) attributed these rodontosaurid remains were reported in the Adaman- teeth coming from the São Luis Basin (VILAS- tina (Turonian-Santonian, Bauru Group; CANDEI- BOAS ET AL., 1999) as being of Carcharodo- RO ET AL. 2002, 2004A), and Marília Formations tosaurus saharicus; nevertheless, we prefer to con- (Maastrichtian, Bauru Group; CANDEIRO, 2002; sider as carcharodontosaurid indet. due to the lack CANDEIRO ET AL., 2004B). Many carcharo- of diagnostic features.

Figure 3 Reconstruction of the skeleton of the carcharodontosaurid Giganotosaurus carolinii Coria and Salgado performed by the paleoartist Jorge Blanco. Estimated length is of about 14 meters.

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12 Abelisauroidea and carchardontosauridae (theropoda, dinosauria) in the cretaceous of south america. Paleogeographical and geocronological implications Carlos Roberto dos Anjos Candeiro, Agustín Guillermo Martinelli

Table 2 Carcharodontosauridae records from South America

Carcharodontosauridae Country Stratigraphy ReferenceSelected Giganotosaurus carolinii Argentina Candeleros Fm., Cenomanian Coria and Currie (2003) Quilmesaurus curriei Argentina Allen Fm., Campanian-Maastrichtian Coria (2001) Carcharodontosauridae Argentina Cerro Barcino Fm., Hauterivian-Barremian; Martinelli and Forasiepi (2004) Mata Amarilla Fm., Turonian; Anacleto Fm., Santonian-Early Campanian; Allen Fm., Campanian-Maastrichtian; Los Alamitos Fm, Campanian Carcharodontosauridae Brazil São Luis Basin, Albian-Cenomanian; Medeiros and Schultz (2002), Adamantina Fm., Turonian-Santonian; Candeiro et al. (2004a,b) Marília Fm., Late Maastrichtian

Figure 4 Solid square, Abelisauroidea. Brazil: 1, Pycnemosaurus; Abelisauridae, 2, Peirópolis; 3, Prata; 4, Santo Anastácio; 6, Flórida Paulista; Argentina; 7, Noasaurus; 8, Abelisaurus; 9, Aucasaurus; 10, Ilokelesia; 11, Ligabueino; 12, Velocisaurus; 13, Xenotarsosaurus; 14, Carnotaurinae; 15, Abelisauridae. Solid circle, Carcharodontosauridae. Brazil: A, Carcharodontosauridae; B, Peirópolis; C, Prata; D, Alfredo Marcondes; Argentina: E, Giganotosaurus; F, Quilmesaurus; Carcharodontosauridae, G, Plaza Huincul area; H, Los Alamitos area; I, Bajo Santa Rosa; J, Cerro Barcino; K, Mata Amarilla, Chubut Province.

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13 Abelisauroidea and carchardontosauridae (theropoda, dinosauria) in the cretaceous of south america. Paleogeographical and geocronological implications Carlos Roberto dos Anjos Candeiro, Agustín Guillermo Martinelli

6 DISCUSSION AND CONCLUSIONS FORASIEPI, in press), Turonian-Santonian Adamantina Formation (Brazil; CANDEIRO ET The paleogeographical distribution of Abeli- AL. 2002), and in the Maastrichtian Marília Forma- sauroidea and Carcharodontosauridae remains is the tion (Brazil; CANDEIRO, 2002; CANDEIRO ET product of the long, isolated evolution of both groups AL., 2003, 2004B). Therefore, if the taxonomic iden- on Gondwanan landmasses (BONAPARTE, 1986). tification of these records is correct and the features used for recognizing carcharodontosaurid af- The early occurrence of Abelisauridae finities are unequivocal (e.g. wrinkles in the crown comes from the Hauterivian-Barremian Cerro tooth), the group persisted into the latest Upper Cre- Barcino Formation (Chubut Province; RAUHUT taceous. Moreover, it is worthy to mention that pre- ET AL. 2003); suggesting that the abelisauroids split sumable Campanian-Maastrichtian carcharodon- into the two recognized families (Abelisauridae and tosaurids are small to medium-sized, contrasting with Noasauridae) at least in the early Lower Cretaceous. older taxa characterized by their large size. From these times, Ligabueino andensi is also known from the La Amarga Formation (Neuquén Province; When the carchorodontosaurid record is BONAPARTE, 1996), and is regarded as a basal more abundant (in the Cenomanian-Turonian), me- abelisauroid (BONAPARTE, 1996; CARRANO; dium to large size abelisaurids are also known (CO- SAMPSON; FORSTER, 2002); however further RIA AND SALGADO, 2000; MARTÍNEZ ET AL., studies and materials should clarify its affinities within 1986). The most derived abelisaurids appear well this group. Both Neuquén and Chubut discoveries documented in the Late Cretaceous with extremely represent the oldest Cretaceous records of theropods specialized forms such as Carnotaurus sastrei in South America (BONAPARTE, 1996; RAUHUT (BONAPARTE; NOVAS; CORIA, 1990). Carno- ET AL. 2003). taurins are known in the Santonian-Campanian (CO- RIA; CHIAPPE; DINGUS, 2002) and in the In the early Lower Cretaceous, carcharo- Maastrichtian (BONAPARTE; NOVAS; CORIA, dontosaurid remains are also found at Cerro Barcino 1990; PASCUAL ET AL. 2000); nevertheless, (Chubut Province, NOVAS; DE VALAIS, 2001; Lamanna, Martinez and Smith (2002) described a VICKERS-RICH ET AL., 1999;) indicating that both carnotaurin maxilla from the Bajo Barreal Forma- groups of large theropods (abelisauroids and tion (late Cenomanian-early Turonian) and suggested carcharodontosaurids) appeared together in the fossil that the origin of this subfamily took place before record as two distinctive gondwanic clades. During the middle Cretaceous. We believe that more mate- the Cenomanian-Turonian, the occurence of carcha- rial is required to confirm this suggestion. rodontosaurids is high in South America (CORIA; SALGADO, 1995; DE VALAIS; APESTEGUÍA, Noasauridae is still poorly known in South 2001; NOVAS ET AL. 1999) as well as in Conti- America but its importance as a derived, endemic nental Africa (e.g. Sereno et al., 1996), and com- group of theropods in Gondwana is remarkable prises the largest theropod forms. Some authors (BONAPARTE, 1991B). Moreover, a new taxon postulated that carcharodontosaurids became extinct from the Maastrichian of Madagascar provides a in the Turonian (APESTEGUÍA, 2002); neverthe- wide knowledge of this bizarre family (CARRANO; less, some isolated teeth and postcranial remains SAMPSON; FORSTER, 2002; SAMPSON; suggest the presence of a member of this family in CARRANO; FORSTER, 2001). the Santonian Anacleto Formation (Neuquén Prov- ince; ALCOBER ET AL., 1998), Late Campanian In summary, the fossil record of abeli- Los Alamitos (Río Negro Province; Apesteguía et sauroids and carcharodontosaurids in South America al. in press), Campanian-Maastrichtian Allen For- shows a dominance of abelisauroids in the upper mation (Río Negro Province; MARTINELLI; Late Cretaceous, while the dominance of large,

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14 Abelisauroidea and carchardontosauridae (theropoda, dinosauria) in the cretaceous of south america. Paleogeographical and geocronological implications Carlos Roberto dos Anjos Candeiro, Agustín Guillermo Martinelli carcharodontosaurid predators was during the REFERENCES Cenomanian-Turonian. At least eight formally described species of Abelisauroidea are recognized in ACCARIE, H., B.; BEUDOIN, J.; DEJAX, G.; Argentina and only one in Brazil. In contrast, only FRIÈS, J.-G.; MICHARD, A.; TAQUET, A. one species of Carcharodontosauridae is known for Découverte dun dinosaure théropode nouvean all South America. (Genusaurus sisteronis n. g., n. sp.) dans lAlbien marin de Sisteron (Alpes de Haute-Provence, Although the knowledge of the evolution of France) et extension au Crétacé inférieur de la lignée Abelisauroidea and Carcharodontasauridae in South cératosaurienne. Computes Rendus dAcadémie America, as well as in the rest of Gondwana, is still des Sciences. Paris. v. 320. p. 327-334, set. 1995. far for being complete, intensive explorations in recent years have provided greater insight into the ALCOBER, O. et al. A Late Cretaceous carcha- composition of theropod faunas in the Cretaceous rodontosaurid (Theropoda: Allosauroidea) from Ar- of Gondwana. gentina. Journal of Vertebrate Paleontology. Northbrook, v. 18, suppl. 3, p. 23A, 1998. ACKNOWLEDGEMENTS APESTEGUÍA, S. Successional structure in conti- This paper is based on two chapters of the nental tetrapod faunas from Argentina along the R. Candeiro Doctor Thesis from the Departamento Cretaceous. In: SIMPÓSIO SOBRE O CRETÁ- de Geologia, Universidade Federal do Rio Janeiro CEO DO BRASIL E II SIMPÓSIO SOBRE EL under L.P. Bergqvist (Rio de Janeiro, Brazil), F.E. CRETÁCICO DE AMÉRICA DEL SUR, 6, 2002, Novas (Buenos Aires, Argentina), and P.J. Currie São Pedro. Boletim... São Pedro: UNESP/Rio (Drumheller, Canada) advise during Doctor Study. Claro, 2002. p. 135-141. We are grateful to Argentinean and Brazilian re- searchers J. Bonaparte(Buenos Aires), C. Muñoz APESTEGUÍA, S.; CANDEIRO, R.; AGNOLIN, (Cipolleti), R. Coria (Plaza Huincul), S. Valais F. L. Late Cretaceous dwarf carcharodontosaurid (Trelew), R. Martinez (Comodoro Rivadavia), L.P. in Patagônia. Ameghiniana. Buenos Aires, Supple- Bergqvist (Rio de Janeiro), and L.C. Ribeiro ment, in press. (Peirópolis, Uberaba) for having provided access to, and extended loans of, specimens in their care. The ASTIBIA, H.; BUFFETAUT, E.; BUSCALIONI, authors wish to thank A. M. Forasiepi (University A. D.; CAPPETTA, H. The fossil vertebrates from of Louisville, KY, USA), L. Witmer (Ohio Univer- Laño (Bosque Country, Spain); new evidence on the sity, Athens, OH, USA), P. OConnor (Ohio Uni- composition and affinities of the Late Cretaceous versity, Athens, OH, USA), Sarah Earland (Univer- continental faunas of Europe. Terra Nova. Paris, sity of Portmouth, UK) for helpful reviews of previ- v. 2, p. 460-466, mar. 1990. ous versions of this article. We also F. Novas (Museo Argentino de Ciencias Naturales, Buenos Aires) for BERTINI, R. J. Evidências de Abelisauridae reading the penultime draft (prior submission) and (Carnosauria: Saurischia) do Neocretáceo da Bacia for providing many helpful comments and discus- do Paraná. In: SIMPÓSIO CRETÁCEO DO sions on Gondwana faunal distributions, including his BRASIL, 6, 1996. Rio Claro. Boletim... São Paulo, specie sharing of unpublished interpretations. We 1996. p. 267-271. also thank to J. Blanco (Buenos Aires) for drawing the figures 3 and 4. Finally, we are grateful to the BITTENCOURT, J. S., KELLNER, A. W. A. Capes Fellowship and the Jurassic Foundation (to Abelisauria (Theropoda, Dinosauria) teeth from Roberto Candeiro) for funding and support. Brazil. Boletim do Museu Nacional Série Geologia. Rio de Janeiro, v. 62, p. 1-8, mar. 2002.

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