Biostratigraphy and geochronology of the late Cenozoic of Córdoba Province (central Argentina)

Journal of South American Earth Sciences 42 (2013) 250e259

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Biostratigraphy and geochronology of the late Cenozoic of Córdoba Province (central Argentina) Laura Edith Cruz* CONICET, Consejo Nacional de Investigaciones Científicas y Técnicas, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Av. Ángel Gallardo 470, Ciudad Autónoma de Buenos Aires C1405DJR, Argentina

a r t i c l e i n f o

a b s t r a c t

Article history: Received 23 December 2011 Accepted 23 June 2012

In the last twenty years, several geological and stratigraphical studies have been undertaken in Córdoba Province, and they have provided useful bases for biostratigraphic work in the late Cenozoic. However, paleontological contributions have been limited to preliminary analyses of mammal assemblages, or specific discoveries. The aim of this work is to contribute to biostratigraphic knowledge of Argentina through the study of late Cenozoic mammals from Córdoba Province. Five localities have been analyzed: San Francisco, Miramar, Río Cuarto, Isla Verde, and Valle de Traslasierra. Through biostratigraphic analysis the first records of several taxa were established, and mammal assemblages with the description and correlation of the sedimentary strata were confirmed. Finally, three Assemblage Zones (Biozonas de Asociación) were proposed: 1) Neosclerocalyptus paskoensiseEquus (Amerhippus) assemblage zone with type area and profile based on the San Francisco locality, referred to the Lujanian (late Pleistoceneeearly Holocene), and comparable to the Equus (Amerhippus) neogeus Biozone of Buenos Aires Province; 2) Neosclerocalyptus ornatuseCatonyx tarijensis assemblage zone with type area and profile based on the San Francisco locality, referred to the Ensenadan (early Pleistocene) and comparable to the Mesotherium cristatum Biozone of Buenos Aires Province, and 3) Nonotherium hennigiePropanochthus bullifer assemblage zone with type area and profile based on the Los Sauces river, Valle de Traslasierra, referred to the MontehermosaneChapadmalalan interval (Pliocene), and comparable to the Trigodon gaudryi, Neocavia depressidens and/or Paraglyptodon chapadmalensis Biozones of Buenos Aires Province. Ó 2012 Elsevier Ltd. All rights reserved.

Keywords: Mammals Late Cenozoic Quaternary Biostratigraphy Córdoba Argentina

1. Introduction

Abbreviations: AGM, Archivo Gráfico y Museo Histórico de la ciudad de San Francisco, Córdoba, Argentina; AMNH, American Museum of Natural History, New York, USA; CB, Colección Botet, Museo de Ciencias naturales, Ayuntamiento de Valencia, Spain; CORD-PZ, Universidad Nacional de Córdoba (UNC), Córdoba, Argentina; DGM-M, Divisião de Geologia e Mineralogia, Coleção do mamíferos fósseis, del Departamento Nacional da Produção Mineral, Río de Janeiro, Brazil; MACN-A, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Colección Ameghino, Ciudad Autónoma de Buenos Aires, Argentina; MACN-PV, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Colección Paleontología Vertebrados, Ciudad Autónoma de Buenos Aires, Argentina; MAMM-PV, Museo Aníbal Montes, Miramar, Córdoba, Argentina; MCNC-PV, Museo Provincial de Ciencias Naturales “Dr. Arturo U. Illia”, Córdoba, Argentina; MLP, Museo de La Plata, La Plata, Buenos Aires, Argentina; MNHN-PAM, Muséum National d’Histoire Naturelle, collection Pampéen, París, France; MUFyCA, Museo Universitario Florentino y Carlos Ameghino, Rosario, Santa Fe, Argentina; UNRC-Pv, Universidad Nacional de Río Cuarto, Colección Paleontología de Vertebrados, Río Cuarto, Córdoba, Argentina; LATyR, Laboratorio de Análisis de Tritio y Radiocarbono, MLP; OSL, optically stimulated luminescence; TL, thermoluminescence. * Tel./fax: þ54 11 4982 6595x193. E-mail address: [email protected]. 0895-9811/$ e see front matter Ó 2012 Elsevier Ltd. All rights reserved. http://dx.doi.org/10.1016/j.jsames.2012.06.015

Mammals from the late Cenozoic of South America (particularly Argentina, Uruguay, Bolivia, Paraguay, Chile, Peru, Venezuela, and Brazil) are well known. However, the chronological scale and continental biostratigraphy of South America of this lapse were established principally on the mammals from Buenos Aires Province, Argentina (Cione and Tonni, 1995a,b, 1999, 2005; Verzi et al., 2004, 2008; Tonni, 2009; Deschamps et al., 2012). Ameghino (1889) created the first stratigraphic scheme for the Pampean Region (Argentina). In recent times, Pascual et al. (1965) established a scheme based on Edades-Mamífero (“South American Land Mammal Ages” e SALMAs) that was later expanded to almost the entire South American continent. Cione and Tonni (1995a,b, 1996, 1999, 2005) examined the theoretical aspects behind the “SALMA” scheme, the units of which they considered to be informal in nature and of limited value. Consequently, Cione and Tonni (1995b, 1999) and Tonni et al. (1999) proposed a return to chronostratigraphic/geochronological classifications with a stratigraphic base. Cione and Tonni (2005) and Tonni (2009) presented

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a new chronological scheme for Buenos Aires Province, which has been used until now in Argentina and extended to South America. The application of this scheme to Córdoba Province (central Argentina) (Fig. 1) is now underway. The substantial number of geological and stratigraphical studies conducted in the last twenty years (e.g. Cantú, 1992; Carignano, 1999; Iriondo, 1999; Kröhling, 1999a,b; Kröhling and Iriondo, 1999; Zárate, 2003; Kemp et al., 2004, 2006) provides a useful basis for biostratigraphic studies of the Quaternary. However, until the present, paleontological contributions have been minor, principally consisting of reports of isolated discoveries (Tauber, 1990, 1997, 1999; Tauber and Di Ronco, 2000, 2003) and preliminary groupings of mammals (Tauber, 2000; Tauber and Álvarez, 2003; Tauber et al., 1997a,b, 2008; Zurita et al., 2007). Although brief paleontological sections have been reported frequently in geological and stratigraphic researches (e.g. Cantú, 1992; Bonalumi et al., 1999; Sanabria et al., 1996), these usually refer to only a handful of specimens. Recently, the first studies of mammal assemblages have begun (Cruz, 2007; Cruz et al., in press), starting with biostratigraphic analysis. These studies, combining analysis of mammalian assemblages and geology, are a useful tool for different kinds of research. Because of the interest and utility of

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mammals for the bio-chronostratigraphy of Argentina and South America, particular attention is paid in these works to determine the stratigraphy as precisely as possible. The aim of the present paper is to provide a biostratigraphical and chronological framework for the Córdoba province based on the study of the fossil mammals (which have precise stratigraphic and geographic provenances) and geological units, combined with analysis of the results of previous studies carried out by other researchers. In this way, this study will contribute to the knowledge of the biostratigraphy of one province of Argentina, through the study of Plioceneeearly Holocene mammals, their geological setting, and their ages. 2. Material and methods 2.1. Collections More than seventy fossil mammals with precise stratigraphic and geographic provenances were selected for study. The specimens are housed in the collections of the MUFyCA, MACN, MLP, and also in local museum collections such as MAMM, AGM, and UNRC.

Fig. 1. Location map. 1. San Francisco; 2. Miramar; 3. Valle de Traslasierra; 4. Río Cuarto; 5. Isla Verde.

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The materials of AGM and MAMM correspond to the regional museums of San Francisco and Miramar. Some of the materials studied from AGM were published by Zurita et al. (2007) and reanalyzed by Cruz et al. (in press). The fossil material of Río Cuarto is housed in the UNRC-PV and was studied by Cruz (2007). The fossil material of Isla Verde is housed in the CORD-PZ and MCNP. Finally, materials collected in the Valle de Traslasierra are housed in the MUFyCA in Rosario; the collection studied here is part of the collection of Castellanos, published by him in several papers (e.g. Castellanos, 1936, 1942, 1944). The systematic of the different groups follows Gaudin and Wible (2006) for Dasypodidae, Fernicola (2008) and Porpino et al. (2010) for Glyptodontia (Xenarthra, Cingulata), Gaudin (2004) for Folivora (Xenarthra), Nasif et al. (2000) for Toxodontidae (Notoungulata), Alberdi and Prado (2004) for Perissodactyla, Soibelzon (2004) for Ursidae (Carnivora), Berman (1994) for Felidae (Carnivora), and McKenna and Bell (1997) for Rodentia. Table 1 shows the complete systematic list of the mammals studied in all localities. The fossils used in this work (Table 1) were compared to holotypes and referred material of almost all species of genera represented for the taxonomic revisions. For this purpose the author visited different collections of fossil and extant mammals (e.g. Argentina, Brazil, France, Spain, USA). The discussion about this topic was realized by the author in your PhD thesis (Cruz, 2011). 2.2. Geology and geochronology Five localities of Córdoba Province were studied: San Francisco, Miramar, Valle de Traslasierra, Río Cuarto, and Isla Verde (Fig. 1). Fieldwork included a general geological survey and field recognition and description of several units of the late Cenozoic. Also, old localities in the Valle de Traslasierra were prospected in order to recognize the stratigraphic levels of the material from old collections. Grain size and mineralogical analyses were carried out on exposed sedimentary units to obtain a preliminary characterization of geological units. These analyses were performed at the Laboratorio de Sedimentos de Facultad de Ciencias Exactas y Naturales, Universidad Nacional de La Pampa. Geochronologic data were extracted from the literature (e.g. Cantú et al., 2004, 2006; Cruz et al., 2010; Kemp et al., 2004, 2006; Kröhling, 1999a,b; Kröhling and Iriondo, 1999). 2.3. Biostratigraphic analysis For biostratigraphic analysis, taxa were listed in the profile of each locality with their stratigraphic provenance. In this way, records of several taxa were established and the assemblages of mammals were documented with the description and correlation of the sedimentary bearing strata. Taxa exclusive to the Plioceneeearly Holocene of Córdoba Province are here reported, biostratigraphic units are proposed and compared with those of the biochronological scheme developed for the Pampean Region. The definition of biostratigraphic units, type areas, and type sections follows the Código Argentino de Estratigrafía (Comité Argentino de Estratigrafía, 1992). The units here proposed are Assemblage Zones (NACSN, 2005; Biozonas de Asociación sensu CAE, 1992). 3. Results and discussion 3.1. Geology and geochronology In San Francisco, almost all material comes from the Cava de Carobolante (31240 30.2400 S and 62 080 15.4700 O, see Cruz et al., in

press for more details). In this site there are four stratigraphic levels, and the fossils came from levels 2 and 3, level 3 is assigned to the Tezanos Pinto Formation (Fig. 2A). In Miramar (30 550 2600 S and 62 4101200 O) the fossils were found in one single level (level 1, assigned to the Tezanos Pinto Formation) (Fig. 2B). The Tezanos Pinto Formation is typical loess, composed of silt, with clay in variable and subordinate amounts, and with rare fine to very fine sand (Kröhling and Iriondo, 1999). The top of this formation is marked by a Hypsithermal palaeosoil, partially eroded, and typically represented by a Bt horizon (Kröhling, 1999a,b; Kröhling and Iriondo, 1999). The Tezanos Pinto Formation was dated by Kröhling (1999a,b) and Kröhling and Iriondo (1999) by a series of TL age estimates in Santa Fe and Córdoba provinces, and more recently by Kemp et al. (2004) by OSL at the type locality of Tortugas (Santa Fe Province). TL dates at the base of the Tezanos Pinto Formation gave ages of 35.89  1.03 ky BP, 31.69  1.62 ky BP, and 32.00 ky BP. OSL dates obtained by Kemp et al. (2004) suggest that the base of this unit may be considerably older, ca. 149.4  9.5 ky BP. At the top of this formation, TL dates gave ages of 8.15  0.40 ky BP below the paleosol and 9.39  0.63 ky BP (Kröhling, 1999a,b; Kröhling and Iriondo, 1999). The fossil materials of Isla Verde (331401900 S and 62 250 2200 O) were found in stratigraphic level 2 (Fig. 2C). This level is composed by eolian facies of sandy silt sediments. It contains nodular concretions of carbonate in the lower and middle sections and rhizoconcretions in the upper section. Taking into account these characteristics, this level is probably best assigned to the La Invernada Formation described by Cantú (1992), and Cantú et al. (2004, 2006). In Río Cuarto Deparment fossil materials were collected from Elena (32 320 5600 S and 64 260 4200 O), Sampacho (33 2301000 S and 64 430 0200 O), Santa Catalina stream (33160 5600 S and 64 2103600 O) and Las Lajas stream (331501100 S and 64 250 5000 O). The profile here presented correspond to the Las Lajas stream, the fossil materials were collected from stratigraphic levels 2 and 3, assigned to the Chocancharava and La Invernada formations respectively (Fig. 3A). Chocancharava Formation (Cantú, 1992) lies disconformably on the Estancia El Cerrito paleosoil. This formation, a pedocomplex named La Colacha (Cantú, 1992), was developed in alluvial plain facies of the abandoned channel. These facies are composed of clayey silts in which three paleosoils are intercalated (Cantú et al., 2004). La Invernada Formation (Cantú, 1992) is a very fine sandy silt with lenses of coarse sand; these lenses are intercalated by other lenses of very fine sandy silt. The top of this formation is marked by Las Tapias geosol, developed in a loess mantle (La Invernada Formation) and stratigraphically equivalent to the hypsithermal paleosoil of Kröhling (1999a). There are no numerical dates for Chocancharava Formation, but absolute datings were reported for lower (Pampiano Formation) and upper (La Invernada Formation) stratigraphic levels. These units were dated by TL and ISRL using sediments from the Las Lajas (Cantú et al., 2004) and Santa Catalina (Cantú et al., 2006) streambeds. The Pampiano Formation was dated between 227.9  30.2 ky BP and 126.3  13.5 ky BP, and La Invernada Formation was dated between 85.7  11.6 ky BP and 12.8  1.5 ky BP. Recently, Cruz et al. (2010) reported a radiocarbon dating obtained from a ground sloth Scelidotherium leptocephalum (Xenarthra, Folivora), found in sediments of the La Invernada Formation. This specimen gave an age estimate of 7.55  0.60 ky 14C BP. The temporal assignations proposed by Cantú (1992, 1998) and Cantú et al. (2004, 2006) are late Pleistocene (interstadial) for the Chocancharava Formation and late Pleistocene (roughly LGM) to early Holocene for the La Invernada Formation. In Valle de Traslasierra (Fig. 3B) the fossil mammals came from Los Chiflones stream (3141045.000 S and 65 01044.300 O), Heredia Cliff (31480 25.700 S and 65 01030.600 O), and Paso del Río Arriba and

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Table 1 Systematic list and number of materials of the taxa studied, up to the levels in which the material could be determined. Order Cingulata Illiger, 1811 Family Dasypodidae Gray, 1821 Eutatus seguiniGervais, 1867 Zaedyus pichiy (Desmarest, 1804) Osgood, 1919 Tolypeutes matacusDesmarest, 1804

MUFyCA 367 e fragment of the dorsal carapace; CORD-PZ 1037 e osteoderm CORD-PZ 1072 e mandible, left humerus and osteoderms from the scapular carapace CORD-PZ 1039 e osteoderms

Suborder Glyptodontia Ameghino, 1889 Phlyctaenopyga sp.

MUFyCA 1017 e fragment of the dorsal carapace

Superfamily Glyptodontoidea Gray, 1869 Tribe Lomaphorini Hoffstetter, 1958 Lomaphorus sp.

Family Panochthidae Castellanos, 1927 Tribe Neosclerocalyptini Paula Couto, 1957 Neosclerocalyptus sp.

Neosclerocalyptus ornatusOwen, 1845 Neosclerocalyptus paskoensis Zurita, 2002 Tribe Panochthini Castellanos, 1927 Panochthus sp. Propanochthus bullifer Burmeister, 1874 Family Glyptodontidae Gray, 1869 Tribe Plohophorini Castellanos, 1932 Plohophorus sp. Tribe Glyptodontini Gray, 1869 Glyptodon sp.

AGM 007 e mandible, right humerus, fragment of the dorsal carapace, and caudal tube; AGM 023 e proximal fragment of the caudal tube; MAMM-PV 016 e fragment of the dorsal carapace and caudal tube; CORD-PZ s/n e osteoderms.

AGM 003 e humerus, ulna, femur, fragment of the dorsal carapace and caudal tube; AGM 004 e postero region of the skull, fragment of the dorsal carapace and fragment of the caudal tube; AGM 022 e dorsal carapace and caudal tube; MAMM-PV 021 e distal fragment of caudal tube; MAMM-PV 022 e postero region of the skull, dorsal carapace and caudal tube; UNRC-PV 005 e left mandible, left humerus and fragment of the dorsal carapace AGM 006 e skull and fragment of the dorsal carapace AGM 002 e skull, mandible, lateral region of the dorsal carapace and caudal tube; MCNC-PV 278 e skull, fragment of the carapace and post-skeleton UNRC-PV 003 e osteoderms of the dorsal carapace; MAMM-PV 024 e osteoderms of the dorsal carapace and fragment of the caudal tube; MUFyCA 383 e caudal tube MACN-Pv 1761 (Holotipo) e fragment of the postero-dorsal carapace and caudal tube; MUFyCA 370 e osteoderms of the dorsal carapace

MUFyCA 396 e fragment of the dorsal carapace MAMM-PV 025 e osteoderms of the dorsal carapace; MUFyCA 354 e fragment of the dorsal carapace; MUFyCA 359 e fragment of the dorsal carapace; UNRC-PV 006 e osteoderms of the dorsal carapace

Order Tardigrada Latham & Davies, 1795 Family Mylodontidae Ameghino, 1889 Scelidotheriinae indet. Catonyx tarijensis (Gervais y Ameghino), 1880 Scelidotherium sp. Scelidotherium leptocephalum Owen, 1840 Mylodon darwini Owen, 1839

AGM 025 e femur, tibia, fibula, and astragalus; CORD-PZ 1047 e astragalus AGM 026 e left mandible, both radius, cervical vertebrae, upper teeth, and both calcaneum MAMM-PV 017 e fragment of the right mandible and aislated teeth UNRC-PV 001 e skull, mandible, both tibia, right astragalus, both navicular, and left calcaneum UNRC-PV 002 e fragment of the skull

Family Megatheriidae Owen, 1842 Megatherium sp. Megatherium americanum Cuvier 1796

MUFyCA 392 e right femur, tibia, proximal fragment of the fibula, astragalus, and calcaneum UNRC PV 004 e fragment of the left mandible

Order Notoungulata Rot, 1903 Suborder Typotheria Zittel, 1892 Family Hegetotheriidae Ameghino, 1894 Paedotherium bonaerense Ameghino, 1887

Family Toxodontidae Gervais, 1847 Nonotherium hennigi Castellanos, 1942 ? Nonotherium hennigi Castellanos, 1942 Toxodon sp. Toxodon platensis Owen, 1837 Order Litopterna Ameghino, 1889 Family Macraucheniidae Gill, 1872 Macrauchenia sp. Macrauchenia patachonicaOwen, 1839 Order Artiodactyla Owen, 1848 Family Camelidae Gray, 1821 Hemiauchenia paradoxa Gervais y Ameghino, 1880

MUFyCA 372 e fragment of the right mandible with p4 to m1; MUFyCA 373 e fragment of the right mandible with p4 to m3 and metacarpian; MUFyCA 374 e fragment of the maxillary with right P2 to M3 and left P2 to M2; MUFyCA 375 e fragment of the right mandible with p4 to m2; MUFyCA 764 e fragment of the left mandible with p2 to m3 MUFyCA 1 (Holotipo) e skull MUFyCA 380 (Cotipo) e fragment of the right mandible with m3 AGM 011 e fragment of the upper incisor MUFyCA 39 e mandible

MUFyCA 393 e distal fragment of the femur; MUFyCA 379 e distal fragment of the right humerus; CORD-PZ 1071 e vertebrae, ribs, and fragment of the pelvis AGM 029 e fragment of the both mandible with left m2 and m3, and right m1 to m3

AGM 028 e right radius plus ulna, and metacarpal

(continued on next page)

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Table 1 (continued )

Order Perissodactyla Owen, 1848 Family Equidae Gray, 1821 Hippidion sp. Equus (Amerhippus) sp

MUFyCA 360 e right lower m2 to m3 AGM 010 e right p3 or p4; MAMM-PV 001 e left m1; MAMM-PV 002 e right M?; MAMM-PV 003 e left M3; MAMM-PV 004 e left pm2; MAMM-PV 005 e right p3; MAMM-PV 006 e right p3; MAMM-PV 007 e right p2; MAMM-PV 008 e left p2; CORD-PZ 1054 e left mandible with p2 to m3

Order Rodentia Family Chinchillidae Bennett, 1833 cf. Lagostomus

AGM 030 e left mandible with p4 to m3; CORD-PZ 1050 e fragment of the left mandible with p4 to m3

Family Octodontidae Waterhouse, 1839 Ctenomys sp.

CORD-PZ 1048 e skull and mandible

Superfamily Cavioidea Kraglievich, 1930:60 Family Caviidae Waterhouse, 1839 Galea sp.

CORD-PZ 1078 e fragment of the right mandible with m1 and m2; CORD-PZ 1081 e fragment of the right mandible with m1 and m2; CORD-PZ 1082 e fragment of the left maxillary with M1 and M2

Order Carnivora Bowdich, 1821 Family Felidae Gray, 1821 Smilodon populator Lund, 1842 Panthera onca (Linnaeus, 1758)

AGM 005 e skeleton almost complete AGM 012 e right tibia, fibula, astragalus, calcaneum, navicular, and cuboid

Family Ursidae Arctotherium sp.

AGM 027 e right femur

Pedernera Cliff (31470 3000 S and 65 000 5800 O), from stratigraphic level 1 and stratigraphic level 2 plus 3. Stratigraphic level 1 is assigned to the Brochero Formation. This formation is normally graded and composed of two facies. One is a matrix-supported conglomerate, the grain size of which is granular and the matrix is sandy silt. The upper facies is a silty to fine sandy paleosol with discontinuous laminate and nodulate tosca with carbonates. The time of deposition of these sediments is controversial, as it has previously been based on mammal fossils without any numerical age. First, Castellanos (1942, 1944) referred this formation to the interval post Chapadmalalanepre Ensenadan, as suggested by the presence of Propanochthus, Nopachthus coagmentatus,

Paedotherium, and Nonotherium. Later, Bonalumi et al. (1999) and Tauber and Álvarez (2003) referred the Brochero Formation to the Montehermosan (late Mioceneeearly Pliocene), based on the presence of N. coagmentatus and the preliminary synonymy of Nonotherium with Xotodon. Different exposures of the Brochero Formation are covered by younger sediments assigned to the Las Rabonas, Mina Clavero, Toro Muerto, and Charbonier formations. The deposition of these formations occurred in different intervals of the Pleistocene (Bonalumi et al., 1999; Carignano, 1999). But, particularly in this section, the Las Rabonas Formation lies disconformably over the Brochero Formation. Las Rabonas Formation is composed of a sandy matrix-supported pebble conglomerate

Fig. 2. Biostratigraphy of the studied localities. A. San Francisco; B. Miramar; C. Isla Verde.

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Fig. 3. Biostratigraphy of the studied localities. A. Río Cuarto; B. Valle de Traslasierra. Some references in Fig. 2.

with rounded clasts and intercalated medium to fine sands. This facies also presents lenses of pebble- to boulder-supported conglomerate. Overlying these sediments is the Mina Clavero Formation. This formation is a fluvial facies composed of intercalations of fluvial sand and silt with typical loess. 3.2. Biostratigraphic analysis At San Francisco locality two mammal assemblages were identified (Fig. 2A). One from stratigraphic level 3, assigned to the Tezanos Pinto Formation, and is composed by Neosclerocalyptus paskoensis (Fig. 4A), Equus (Amerhippus) sp. (Fig. 4B), Toxodon sp., Arctotherium sp., and a possible Lagostomus. The other mammal assemblage comes from stratigraphic level 2 is composed of Lomaphorus sp., Neosclerocalyptus ornatus (Fig. 4C), Catonyx

tarijensis (Fig. 4D), Scelidotheriinae indet., Macrauchenia patachonica, Hemiauchenia paradoxa, Smilodon populator, and Panthera onca. At Miramar locality only one mammal assemblage was recognized (Fig. 2B) from stratigraphic level 1, assigned to the Tezanos Pinto Formation. It is composed of Lomaphorus sp., Neosclerocalyptus sp., Panochthus sp., Glyptodon sp., Scelidotherium sp., Toxodon sp., and Equus (Amerhippus) sp. At the Isla Verde locality one mammal assemblage was identified (Fig. 2C), from stratigraphic level 2 and including Eutatus seguini, Zaedyus pichiy, Tolypeutes matacus, Neosclerocalyptus sp., N. paskoensis, Lomaphorus sp., Scelidotheriinae indet., Macrauchenia sp., Equus (Amerhippus) sp., Ctenomys sp., cf. Lagostomus and Galea sp. At Río Cuarto department two mammal assemblages were identified (Fig. 3A). One of them, from stratigraphic level 2 and

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Fig. 4. Species taken into account for the definition of biozones A. Neosclerocalyptus paskoensis, AGM 002, lateral view of the skull, scale 5 cm; B. Equus (Amerhippus) sp., AGM 010, right lower premolar, scale 3 cm; C. Neosclerocalyptus ornatus, AGM 006, lateral view of the skull, scale 5 cm; D. Catonyx tarijensis, AGM 026, left mandible, scale 5 cm; E. Propanochthus bullifer, MUFyCA 370, osteoderms of the dorsal carapace, scale 5 cm; F. Nonotherium hennigi, MUFyCA 1, occlusal view of the skull, scale 10 cm.

assigned to Chocancharava Formation, contains only a two taxa, Panochthus sp. and Glyptodon sp. The other comes from stratigraphic level 3; it is assigned to the La Invernada Formation and is composed of Neosclerocalyptus sp., S. leptocephalum, and Megatherium americanum. Finally, at the Valle de Traslasierra locality two mammal assemblages were recognized (Fig. 3B). The first one from stratigraphic level 1, is assigned to the Brochero Formation and composed of Phlyctaenopyga sp., Propanochthus bullifer (Fig. 4E), Plohophorus sp., Paedotherium bonaerense, and Nonotherium hennigi (Fig. 4F); and the second from stratigraphic levels 2 and 3, featuring E. seguini, Panochthus sp., Glyptodon sp., Megatherium sp., Macrauchenia sp., Toxodon platensis, and Hippidion sp. The correlation among localities (Fig. 5) was established by evident biostratigraphic, stratigraphic relationships and numerical age. From a biostratigraphic point of view, stratigraphic levels 3 at San Francisco, 1 at Miramar, and 2 at Isla Verde are strongly correlated, in view of: 1) the presence of Equus (Amerhippus) sp. in all levels; 2) the occurrence of N. paskoensis and cf. Lagostomus sp. in San Francisco and Isla Verde; and 3) the presence of Toxodon sp. in Miramar and San Francisco. These biostratigraphic correlations are supported too by stratigraphic relationships, because stratigraphic level 1 of Miramar and stratigraphic level 3 of San Francisco are assigned to the same formation (Tezanos Pinto Formation). According to numerical ages, stratigraphic level 1 of Miramar is correlated with stratigraphic level 3 of Río Cuarto. Thus, this

correlation allows addition to the faunistic group of a genus, Scelidotherium, present in both localities, as well as a species, M. americanum. The relationship between stratigraphic levels 2 of Río Cuarto, San Francisco, and the upper stratigraphic levels of the Brochero Formation in Valle de Traslasierra is uncertain. Principally because information is insufficient to be able to know which are the upper stratigraphic levels of the Brochero Formation (Valle de Traslasierra) in which Castellanos found the material. But, due to the superposition of strata, the faunal assemblages of stratigraphic level 2 at San Francisco and Río Cuarto are older than the faunal assemblages of level stratigraphic 3 at the same localities. From a regional perspective, and considering the biostratigraphic distribution of the mammals and the correlations previously described, three Assemblage Zones (Biozonas de Asociación) are proposed (Fig. 5). The Assemblage Zones are, from youngest to oldest: 1) Neosclerocalyptus paskoensiseEquus (Amerhippus) assemblage zone. This biozone is defined by the co-occurrence of N. paskoensis, Equus (Amerhippus) sp., S. leptocephalum, M. americanum, cf. Lagostomus, Toxodon sp., Arctotherium sp., E. seguini, Z. pichiy, and T. matacus. Type area: Cava de Carobolante, San Francisco locality. Type profile: Represented in Fig. 2. The stratotype is a composite of stratigraphic level 1 from Miramar, 2 from Isla

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Fig. 5. Bio-chronostratigraphic scheme of the Córdoba province.

Verde, 3 from San Francisco, and 3 from Río Cuarto. The first two are assigned to the Tezanos Pinto Formation, and the last to the La Invernada Formation. Age: This Biozone is correlated with the Lujanian (late Pleistoceneeearly Holocene) of the Pampean region. Remarks: This biozone has two biostratigraphic indicators (N. paskoensis and Equus [Amerhippus]), both being guide fossils of the Lujanian in Argentina (Cione and Tonni, 2005; Soibelzon et al., 2010; Tonni, 2009; Zurita et al., 2005, 2009). On the other hand, this biozone has a numerically supported temporal age of the Tezanos Pinto and La Invernada formations (Cantú et al., 2004, 2006; Cruz et al., 2010; Kemp et al., 2004; Kröhling, 1999a,b; Kröhling and Iriondo, 1999). 2) Neosclerocalyptus ornatuseC. tarijensis assemblage zone. This biozone is based on the co-occurrence of N. ornatus, C. tarijensis, M. patachonica, H. paradoxa, S. populator, and P. onca. Type area: “Cava de Carobolante”, San Francisco locality. Type profile: Represented in Fig. 2. The stratotype is defined as stratigraphic level 2 at San Francisco. Age: This Biozone is correlated with the Ensenadan (earlyemiddle Pleistocene) of the Pampean region. Remarks: this biozone has two biostratigraphic indicators (N. ornatus and C. tarijensis), both guide fossils of the Ensenadan in Argentina (Cione and Tonni, 2005; McDonald, 1987; McDonald and Perea, 2002; Pujos, 2000; Soibelzon et al., 2010; Tonni, 2009; Zurita et al., 2005, 2009). Additionally, this biozone has a stratigraphic correlation with the one previously described by superposition of strata. 3) Nonotherium hennigieP. bullifer assemblage zone. This biozone is defined by the co-occurrence of N. hennigi, P. bullifer, Phlyctaenopyga sp., Plohophorus sp., and P. bonaerense.

Type area: Los Sauces River, Valle de Traslasierra locality. Type profile: Represented in Fig. 3. The stratotype is defined as stratigraphic level 1 from Valle de Traslasierra, assigned to the Brochero Formation. Age: This Biozone is correlated with the Montehermosane Chapadmalalan (early Plioceneelate Pliocene) of the Pampean region. Remarks: This biozone has two biostratigraphic indicators (Phlyctaenopyga sp. and Plohophorus sp.). Phlyctaenopyga is recorded in Catamarca, from levels 15 to 32 with numerical ages between 6.68 My BP (level 8) and 3.54 My BP (level 30) (Marshall and Patterson, 1981) and is correlated with the MontehermosaneChapadmalalan interval. Plohophorus is a guide fossil present in the MontehermosaneChapa dmalalan of Argentina (Cione and Tonni, 2005; Tonni, 2009). Full establishment of this regional model is hindered by several factors. First, further studies are needed to elucidate the age and guide fossils of the N. hennigieP. bullifer and N. ornatuseC. tarijensis assemblage zones which are only registered at two different sites in Córdoba. Two, it is important to discover other sites with vertebrates of the PlioceneePleistocene in order to corroborate this model and to allow the study of the important faunal turnover that took place during this interval. 4. Conclusions Three “assemblage zones” (“biozonas de asociación”) are proposed in this paper: 1) N. paskoensiseEquus (Amerhippus) assemblage zone with type area and profile based on the San Francisco locality, referred to the Lujanian (late Pleistoceneeearly Holocene), and comparable to the Equus (Amerhippus) neogeus

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biozone from Buenos Aires Province; 2) N. ornatuseC. tarijensis assemblage zone with type area and profile based on the San Francisco locality, referred to the Ensenadan- (earlyemiddle Pleistocene) and comparable to the Mesotherium cristatum biozone of Buenos Aires Province, and 3) N. hennigieP. bullifer assemblage zone with type area and profile based on sediments of the Los Sauces River, Valle de Traslasierra, referred to the MontehermosaneChapadmalalan interval (earlyelate Pliocene), and comparable to the Trigodon gaudryi, Neocavia depressidens and/ or Paraglyptodon chapadmalensis Biozones of Buenos Aires Province. The last three biozones are under discussion (see Deschamps et al., 2012). Finally, among new records for Córdoba Province are: Phlyctaenopyga sp. and P. bonaerense from the Brochero Formation (MontehermosaneChapadmalalan), Arroyo Los Chiflones profile, Villa Cura Brochero. Acknowledgements I thank A. Kramarz (MACN), M. Reguero and L. Pomi (MLP), S. Cornero and F. Solomita (MUFyCA), M.T. Milani, A. Bienedell and A. Orellano (AGM), L. Dorazat and H. Giraudo (MAMM), D. Álvarez and C. Luna (MCNC), A. Tauber (Museo de Paleontología, UNC), M. Belinchón (CB, Museo Ciencias Naturales, Valencia) and C. Argot (MNHN) for the access to museum collections. I also thank M.S. Bargo, J.C. Fernicola, and A.G. Kramarz for their help during my research. This work was supported by CONICET, ANPCyT-PICT 38171 (E.P. Tonni), CIC (M.S. Bargo), UNLP N647 and CONICET-PIP 1054 (S.F. Vizcaíno). References Alberdi, M.T., Prado, J.L., 2004. Caballos fósiles de América del Sur, Una historia de tres millones de años. In: INCUAPA serie monográfica, first ed., vol. 3. Universidad Nacional del Centro de la Provincia de Buenos Aires, Buenos Aires, 269 pp. Ameghino, F., 1889. Contribución al conocimiento de los mamíferos fósiles de la República Argentina. Actas de la Academia Nacional de Ciencias de Córdoba 6, 1e1027. Berman, W.D., 1994. Los carnívoros continentales (Mammalia, Carnivora) del Cenozoico en la provincia de Buenos Aires. Ph.D. thesis, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, Unpublished. Bonalumi, A., Martino, R., Baldo, E., Zarco, J., Sfragulla, J., Carignano, C., Kraemer, P., Escayola, M., Tauber, A., Cabanillas, A., Juri, E., Torres, B., 1999. Hoja Geológica 3166-IV, Villa Dolores. Provincias de Córdoba, La Rioja y San Luis, vol. 250. Instituto de Geología y Recursos Minerales, Servicio Geológico Minero Argentino, Buenos Aires, 123 pp. Cantú, M.P., 1992. Provincia de Córdoba. In: Iriondo, M. (Ed.), El Holoceno en la Argentina, vol. 1. CADINQUA, pp. 1e16. Cantú, M.P., 1998. Estudio geocientífico para la evaluación ambiental y la ordenación territorial de una cuenca pedemontana. Caso: Cuenca del Arroyo La Colacha, Dpto. Río Cuarto, provincia de Córdoba. Ph.D. thesis, Universidad Nacional de Río Cuarto. Unpublished. Cantú, M.P., Schiavo, H.F., Musso, T.B., Becker, A.R., 2004. Paleosuelos del Pleistoceno superioreHoloceno del sur de la provincia de Córdoba, Argentina. In: XIX Congreso Internacional de la Ciencias del Suelo, CD 12pp. Cantú, M.P., Schiavo, H.F., Becker, A.R., Zhou, L., Grumelli, M.T., 2006. Pleistoceno superior tardíoeHoloceno de la cuenca media del arroyo Santa Catalina, provincia de Córdoba, Argentina. In: III Congreso Argentino de Cuaternario y Geomorfología, Actas, pp. 777e786. Carignano, C.A., 1999. Late Pleistocene to recent climate change in Córdoba Province, Argentina: geomorphological evidence. Quaternary International 57/58, 117e134. Castellanos, A., 1936. Los sedimentos del Pampeano inferior y del Araucano en el Valle de Los Reartes. In: serie técnico-científica, vol. 6. Publicaciones de la facultad de ciencias matemáticas, físico-químicas y naturales aplicadas a la industria de la Universidad Nacional del Litoral, pp. 1e110. Castellanos, A., 1942. Los sedimentos prepampeanos del Valle de Nono (Sierra de Córdoba), vol. 4. Publicaciones del Instituto de Fisiografía y Geología de la Universidad Nacional del Litoral, Argentina, pp. 1e63. Castellanos, A., 1944. Paleontología estratigráfica de los sedimentos neógenos de la provincia de Córdoba, vol. 23. Publicaciones del Instituto de Fisiografía y Geología de la Universidad Nacional del Litoral, pp. 5e47. Cione, A.L., Tonni, E.P., 1995a. Chronostratigraphy and “Land mammal-ages”: the Uquian problem. Journal of Paleontology 69, 135e159.

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