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Proterotheriidae and Macraucheniidae (Litopterna, Mammalia) from the Pleistocene of Rio Grande Do Sul State, Brazil

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Proterotheriidae and Macraucheniidae (Litopterna, Mammalia) from the Pleistocene of Rio Grande Do Sul State, Brazil Rev. bras. paleontol. 12(3):231-246, Setembro/Dezembro 2009 © 2009 by the Sociedade Brasileira de Paleontologia doi:10.4072/rbp.2009.3.06 PROTEROTHERIIDAE AND MACRAUCHENIIDAE (LITOPTERNA, MAMMALIA) FROM THE PLEISTOCENE OF RIO GRANDE DO SUL STATE, BRAZIL CAROLINA SALDANHA SCHERER, VANESSA GREGIS PITANA & ANA MARIA RIBEIRO Seção de Paleontologia, Museu de Ciências Naturais, Fundação Zoobotânica do Rio Grande do Sul, Av. Salvador França, 1427, 90690-000, Porto Alegre, RS, Brasil [email protected], [email protected], [email protected] ABSTRACT – New materials of the order Litopterna from the late Pleistocene (Lujanian) of Rio Grande do Sul State are presented here. The specimens are an astragalus and calcaneum, assigned to cf. Neolicaphrium recens, and fragments of mandible, humeri, tibia and fibula, astragali, besides vertebrae and metatarsal, all of them attributed to Macrauchenia patachonica. The material described here comes from the municipalities of Santa Vitória do Palmar (Hermenegildo Beach, Chuí Creek), Uruguaiana (Touro Passo Creek), Alegrete (Sanga da Cruz Creek) and Pantano Grande (Sanga Borba Creek). The record of these taxa in the Lujanian of Rio Grande do Sul State shows a greater similarity with the fauna in the same latitudinal belt of Argentina and Uruguay than that of the intertropical region. Key words: Litopterna, Proterotheriidae, Macraucheniidae, Lujanian, Rio Grande do Sul State. RESUMO – São aqui apresentados novos materiais da Ordem Litopterna para o Pleistoceno final (Lujanense) do Estado do Rio Grande do Sul. Os espécimes constam de astrágalo e calcâneo, atribuídos a cf. Neolicaphrium recens; e fragmentos de mandíbula, úmeros, tíbia e fíbula, astrágalos, além de vértebras e metatarsal, todos atribuídos a Macrauchenia patachonica. O material aqui descrito provém dos municípios de Santa Vitória do Palmar (Balneário Hermenegildo e Arroio Chuí), Uruguaiana (Arroio Touro Passo), Alegrete (Sanga da Cruz) e Pantano Grande (Sanga Borba). O registro destes táxons para o Lujanense do RS evidencia, mais uma vez, a maior similaridade da fauna deste estado com a da mesma faixa latitudinal da Argentina e do Uruguai, do que com a da região intertropical. Palavras-chave: Litopterna, Proterotheriidae, Macraucheniidae, Lujanense, Rio Grande do Sul. INTRODUCTION Bergqvist et al., 1998; Bond, 1999; Flynn et al., 2002; Kramarz & Bond, 2005). Since the Mio-Pliocene, their diversity was The Litopterna Ameghino, 1889 are native ungulates of reduced and they were considered extinct in the late Pliocene South America that had a wide distribution on this continent (Bond, 1999). However, Neolicaphrium recens Frenguelli, since the late Paleocene until the late Pleistocene. The order 1921 was described for the Lujanian (late Pleistocene, Cione Litopterna (sensu McKenna & Bell, 1997) includes the families & Tonni, 2005, including Bonaerian, sensu Bond et al., 2001) Protolipternidae Cifelli, 1983 (late Paleocene),PROVAS Notonychopidae of Argentina (Cordoba Province), and further, finds of this Soria, 1989 (late Paleocene), Adianthidae Ameghino, 1891 species were recorded for the Lujanian (including Bonaerian) (early Eocene - middle Miocene), Proterotheriidae Ameghino, of Uruguay (Salto Department) and Argentina (Cordoba, 1887 (late Paleocene - late Pleistocene) and Macraucheniidae Corrientes and Santa Fe Provinces) and also for the Gervais, 1855 (late Paleocene - late Pleistocene), the last two Ensenadan (early to middle Pleistocene, Cione & Tonni, 2005) being the main goal of this work. (Córdoba Province, Argentina) (Alvarez, 1974; Perea et al., The Proterotheriidae include two subfamilies and 18 1998; Tauber, 2000; Bond et al., 2001; Ubilla et al., 2007; Vezzosi genera (sensu McKenna & Bell, 1997), of small and middle et al., 2009). Pitana et al. (2005) reported the presence of an size forms. They are characterized mainly by brachydont to undetermined Proterotheriidae material for the Lujanian of mesodont dentition and a marked trend toward monodactily, southern Brazil (Figure 1). with digit III more developed, digits I and V lost, and digits II The family Macraucheniidae includes three subfamilies and IV reduced (Bond, 1999; Bond et al., 2001). They are and 16 genera of middle size to very large forms. The recorded for the Tertiary in Argentina, Brazil, Bolivia, Chile, representatives of this family are more conservative in dental Colombia and Uruguay (Marshall et al., 1983; Frailey, 1986; and postcranial aspects than proterotherids, and are Bond et al., 1995; Bergqvist, 1996; Cifelli & Guerrero, 1997; characterized mainly by the retraction of the nasals and the 231 232 REVISTA BRASILEIRA DE PALEONTOLOGIA, 12(3), 2009 backward shift of the narines and by complete dental formula Paraguay, northern and southern Chile, Peru and Bolivia (from (Bond, 1999). They are recorded for the Tertiary of Argenti- the Ñuapua and Tarija faunas, where deposits are assigned na, Brazil, Bolivia, Colombia and Chile (Marshall et al., 1983; to Ensenadan or Lujanian, according to different authors) Perea et al., 1994; Bond et al., 1995; Cifelli & Guerrero, 1997; (Souza-Cunha, 1959; Marshall et al., 1984; Marshall & Bond, 1999; Flynn et al., 2002; Croft et al., 2004, 2007). For Sempere, 1991; Oliveira, 1992; Buchmann, 1994; Ubilla et al., the Pleistocene of the Buenos Aires Province (Argentina), 1994; MacFadden & Shockey, 1997; Bond, 1999; Casamiquela, Ameghino (1888) described Macrauchenia ensenadensis, 1999; Ubilla, 2004; López & Labarca, 2005; Scherer et al., later transferred to the genus Macraucheniopsis by Paula- 2006; Coltorti et al., 2007). Macrauchenia patachonica was Couto (1945). Currently, this species is considered as valid recorded for the northeastern Brazilian states of Pernambuco, for the Ensenadan of the Pampean region of Argentina (Bond, Bahia, Ceará, Paraíba and Minas Gerais (e.g., Winge, 1906; 1999). Macrauchenia patachonica Owen, 1838 is recorded Paula-Couto, 1947, 1954, 1970, 1980; Vidal, 1946, 1955). since the Ensenadan, by only one find for the Pampean region Xenorhinotherium bahiense Cartelle & Lessa, 1988 was of Argentina (Bond, 1999), and more abundantly for the described based on material from states of Bahia Lujanian (including the Bonaerian) of Argentina, in the (northeastern Brazil) and Minas Gerais (southeastern Brazil). Pampean (Buenos Aires, Santa Fe and San Luis Provinces), These authors suggested that all the Macraucheniidae mate- Mesopotamian (Corrientes and Entre Rios Provinces), rial previously recorded from the northeastern Brazil should Patagonian (Santa Cruz Province) and northern regions (For- belong to this taxon. Later records of macraucheniids from mosa and Jujuy Provinces) (Ameghino, 1889; Bond, 1999; states of Bahia, Pernambuco, Ceará, Paraíba and Rio Grande Ferrero et al., 2007; Ferrero, 2009). This species was also do Norte were also assigned to X. bahiense (Oliveira et al., recorded for the Lujanian of southern Brazil, Uruguay, 1989; Bergqvist, 1993; Bergqvist et al., 1997). Recently, Guérin & Faure (2004) suggested that this taxon could be a synonym of Macrauchenia patachonica, including new material from Piauí State. Nevertheless, some authors (e.g., Socorro, 2006) do not follow this proposition, considering X. bahiense as a valid taxon. Xenorhinotherium bahiense was also recently recorded for the Muaco, Taima Taima and Cuenca del Lago de Valencia faunas from Venezuela (Socorro, 2006). Recently, Dias-Neto et al. (2008) referred to the presence of the Macraucheniidae in the Pleistocene of Alagoas State, but these authors did not assign the material to Macrauchenia or Xenorhinotherium due to the absence of diagnostic elements. For Rio Grande do Sul State, Macrauchenia patachonica has been recorded for the Coastal Plain of Rio Grande do Sul, by Souza-Cunha (1959), for Sanga da Cruz Creek (Alegrete) by Oliveira (1992), Lessa & Oliveira (1996), Milder (2000) and Scherer & Da-Rosa (2003), and for the Pantano Grande Municipality by Scherer et al. (2003). Scherer et al. (2006) revised the material of the Macraucheniidae from this state and referred the material from the municipalities of PROVASSanta Vitória do Palmar (Hermenegildo Beach), Uruguaiana (Touro Passo Creek) and Alegrete (Sanga da Cruz Creek) to Macrauchenia patachonica (Figure 1). This work provides a description and comparison of fossil remains of Macraucheniidae and Protherotheriidae from Pleistocene of Rio Grande do Sul State, and includes a discussion about the taxonomic relevance of its osteological and dental features. GEOLOGICAL SETTING Figure 1. Geographic distribution of the Macraucheniidae and Proterotheriidae during the Pleistocene in South America, according The material of Litopterna from Rio Grande do Sul State to Paula-Couto (1945), Alvarez (1974), Marshall et al. (1984), comes from Pleistocene fluvial and lagoonal deposits, from Cartelle & Lessa (1988), Oliveira et al., (1989), Bergqvist (1993), the municipalities of Uruguaiana (Touro Passo Creek), Ale- Bergqvist et al. (1997), MacFadden & Shockey (1997), Perea et al. grete (Sanga da Cruz Creek), Pantano Grande (Sanga Borba (1998), Bond (1999), Casamiquela (1999), Tauber (2000), Bond et al. (2001), Guérin & Faure (2004), Ubilla (2004), López & Labarca Creek) and Santa Vitória do Palmar (Hermenegildo Beach and (2005), Scherer et al. (2006), Coltorti et al. (2007), Ubilla et al. Chuí Creek). (2007), Socorro (2006), and Vezzosi et al. (2009). Touro Passo Creek is a tributary on the left bank of the SCHERER ET AL. – PROTEROTHERIIDAE AND MACRAUCHENIIDAE FROM
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