Late Carboniferous postglacial brachiopod faunas in ...

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Late Carboniferous postglacial brachiopod faunas in the Southwestern Gondwana margin Gabriela A. Cisterna a,∗ , Andrea F. Sterren b a

Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET)-Universidad Nacional de La Rioja (UNLAR), Av. Dr. Luis M. de la Fuente s/n, La Rioja 5300, Argentina b CICTERRA, CONICET, Universidad Nacional de Córdoba, Av. Vélez Sarsfield 1611, Ciudad Universitaria, X5016GCA Córdoba, Argentina Received 15 January 2016; received in revised form 8 June 2016; accepted 14 July 2016

Abstract The glaciomarine sediments related to the Late Paleozoic Ice Age (LPIA) have an excellent stratigraphic record in Argentina, particularly those associated to the Late Carboniferous glacial episode identified along the southwestern margin of South America: Bolivia (Tarija Basin), west central Argentina (Calingasta-Uspallata Basin) and Patagonia (Tepuel-Genoa basins). The aim of this contribution is mainly a biostratigraphy update of the carboniferous brachiopod faunas that occur in the earliest postglacial interval (late Serpukhovian–Bashkirian) in the west central Argentina (i.e., Levipustula and Aseptella-Tuberculatella/Rhipidomella-Micraphelia faunas) and its regional correlation with those equivalents in the nearby basins. Components of these faunas are recognized from the Bolivia to Argentine Patagonia and their compositional variations appear to be controlled principally by a paleolatitudinal factor. The affinities showed by the postglacial faunas of the Calingasta-Uspallata Basin and the faunal assemblages that integrate the Lanipustula and Tuberculatella biozones in Patagonia differ from the significant contrast proposed by other authors, based on the paleogeographical position of Patagonia in the Late Paleozoic. Paleoecological studies focused on the paleoenvironmental controls related with the glacial dynamic are suggested to understand the complex relationship between these postglacial faunas. © 2016 Elsevier B.V. and Nanjing Institute of Geology and Palaeontology, CAS. All rights reserved. Keywords: Brachiopods; Carboniferous; Postglacial; South America

1. Introduction Postglacial marine faunas related to the Late Paleozoic Ice Age (LPIA) have an excellent record in the southwestern Gondwana margin, particularly those associated with the Late Carboniferous glacial episode identified in South America (López Gamundí, 1997). The deglaciation processes connected to this event produced an important sea level rise and consequently an extended postglacial transgression in the areas located along the Paleo-Pacific basins, from Bolivia to Southern Argentina (Fig. 1). A rich marine invertebrate fauna appears usually associated with the Late Carboniferous postglacial succession;

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Corresponding author. E-mail addresses: [email protected] (G.A. Cisterna), [email protected] (A.F. Sterren).

brachiopods, as one of the main components of this fauna, can be considered the group of more biostratigraphical value. The aim of this contribution is mainly a biostratigraphy update of the Late Carboniferous brachiopod faunas that occur in the earliest postglacial interval (late Serpukhovian–Bashkirian) of glaciomarine sequences from Argentine Precordillera (Fig. 2A) as well as the regional correlation with those equivalents in the nearby basins of Bolivia (Tarija Basin, Fig. 2B) and Patagonia (Tepuel-Genoa Basin, Fig. 2C). Postglacial brachiopod faunas from Precordillera have been widely studied (Keidel and Harrington, 1938; Amos et al., 1963; Amos, 1979; Taboada, 1989; Taboada and Cisterna, 1996; Cisterna, 1999; Simanauskas and Cisterna, 2001; Cisterna and Sterren, 2008, 2010; Taboada and Shi, 2011; Cisterna et al., 2013), and they have an excellent stratigraphic control, with a clearly defined position within the glacial-postglacial sequences that allow to consider them as a key postglacial brachiopod assemblage to inter-basinal biostratigraphic correlation.

http://dx.doi.org/10.1016/j.palwor.2016.07.005 1871-174X/© 2016 Elsevier B.V. and Nanjing Institute of Geology and Palaeontology, CAS. All rights reserved.

Please cite this article in press as: Cisterna, G.A., Sterren, A.F., Late Carboniferous postglacial brachiopod faunas in the Southwestern Gondwana margin. Palaeoworld (2016), http://dx.doi.org/10.1016/j.palwor.2016.07.005

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Bolivia), characterized by the glacial sediments of the Macharetí and Mandiyutí groups; the Calingasta-Uspallata Basin, in the west central Argentina, where the typical glacial-postglacial sequences can be recognized in several stratigraphic units that will be listed below; and the Tepuel Genoa Basin in Patagonia, represented by glaciomarine deposits of the Pampa de Tepuel and Las Salinas formations. The Late Carboniferous postglacial deposits in each of these basins are characterized respectively by distinctive marine invertebrate faunas. Brachiopods are one of the most important components of the postglacial assemblages studied and their distribution is analyzed herein. 2.1. Calingasta-Upallata Basin Fig. 1. Paleogeographic reconstruction of Gondwana Supercontinent with glacial evidence in the studied basins. Glacial episodes after López Gamundí (1997).

2. The postglacial faunas in the South American basins Important successions of glacial diamictites overlain by transgressive postglacial marine shales were deposited during the Late Carboniferous interval in different South American basins: the Tarija Basin (northernmost part of Argentina and

The Carboniferous glaciomarine deposits of the CalingastaUspallata Basin outcrop disconnected and irregularly along western of the Tontal Hill, west of Argentina (Fig. 2A). The complete sedimentary succession that characterizes this basin does not exceed the 1500 m of thickness in different sections studied, and the postglacial marine interval, in particular, reaches only a few hundred meters (about 100–200 m). The characteristic glacial-postglacial transition from basal deposits of proximal glacial-marine environment with sediment gravity flows to postglacial marine fossiliferous mudstones

Fig. 2. Geographic location of Carboniferous postglacial faunas in South America. (A) Calingasta-Uspallata Basin (LC, La Capilla Formation; CT, Ciénaga Larga del Tontal Foramtion; HV, Hoyada Verde Formation; EP, El Paso Formation; Le, Leoncito Formation; Ya, Yalguaraz Formation; AJ, Agua del Jagüel Formation), central-west of Argentina. (B) Tarija Basin. (C) Tepuel Genoa Basin (Patagonia) (LS, Las Salinas Formation; PT, Pampa de Tepuel Formation).




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Fig. 3. Hoyada Verde and El Paso formations, Calingasta-Uspallata Basin, with graphics showing the percentage of the generic abundance of brachiopods vs. bivalves in each fauna studied. AT/RM, Aseptella-Tuberculatella/Rhipidomella-Micraphelia fauna.

(López Gamundí and Martínez, 2000) have been recognized in several stratigraphic units (Hoyada Verde, El Paso, La Capilla, Leoncito, Yalguaraz formations). One of these typical glacial-postglacial sequences has been described from the Barreal Hill (San Juan province), in the Hoyada Verde Formation (late Serpukhovian–Bashkirian) (Fig. 3), in which the postglacial deposits contain the very well-known Levipustula fauna (Cisterna and Sterren, 2008, 2010). In an equivalent section located to south of the Barreal Hill (Fig. 2A), El Paso Formation (Fig. 3), a different fauna composed of the Aseptella-Tuberculatella and Rhipidomella-Micraphelia assemblages was identified (Cisterna et al., 2013). These two faunas with significant taxonomic, taphonomic, and paleoecological differences appear associated with the earliest postglacial transgression in the Carboniferous diamictite-bearing sequences of the Calingasta-Uspallata Basin in Precordillera (Fig. 2A): the Levipustula fauna, widely studied because of its biostratigraphic, paleocologic, and paleobiogeographic implications (Cisterna, 1999; Cisterna and Sterren, 2008, 2010) and a less known faunal assemblages from the El Paso Formation that integrate the Aseptella-Tuberculatella/Rhipidomella-Micraphelia fauna (Cisterna et al., 2013).

2.1.1. Levipustula fauna The Levipustula fauna is composed mainly of bivalves, brachiopods, and bryozoans with less abundant gastropods, crinoids, and conulariids. Besides being characterized by these three components, this fauna shows a particular generic richness of bivalves vs. brachiopods, the dominant groups in the Levipustula fauna (Fig. 3). Brachiopods that characterize the Levipustula fauna are dominated by spiriferids (Costuloplica leoncitensis (Harrington), Kitakamithyris booralensis Campbell, Kitakamithyris immensa Campbell, Kitakamithyris ssp., Torynifer tigrensis Taboada and Cisterna, Spiriferellina octoplicata (Sowerby), Septosyringothryis keideli (Harrington)), accompanied by the ubiquitous productid Levipustula levis Maxwell and the terebratulid Beecheria sp. (Fig. 4). The detailed geographic and stratigraphic distribution of the Levipustula fauna in the different sections of the Calingasta-Uspallata Basin (i.e., Hoyada Verde, Leoncito, La Capilla and Yalguaraz formations) has been provided by Cisterna and Sterren (2010). The Levipustula fauna has been particularly studied from the postglacial shales of the Hoyada Verde Formation in the core of Barreal Hill, San Juan province, where it occurs either as shell beds of 1–5 cm thick or as nests in a stratigraphic interval




Fig. 4. Brachiopods of the Levipustula fauna. (A–C) Levipustula levis Maxwell; (A) ventral valve, IPI 3238, scale bar = 1 cm; (B) internal mould of dorsal valve, IPI 4501, scale bar = 1 cm; (C) dorsal valve partially decorticated, IPI 3442, scale bar = 1 cm. (D–H) Costuloplica leoncitensis (Harrington); (D) dorsal valve partially decorticated, IPI 3223, scale bar = 0.95 cm; (E) ventral valve, IPI 3221, scale bar = 3 cm; (F) ventral valve, IPI 3214, scale bar = 1.69 cm; (G) internal mould of ventral valve, CEGH-UNC 22904, scale bar = 2.37 cm; (H) ventral valve, IPI 3208, scale bar = 0.95 cm. (I, J) Kitakamithyris sp.; (I) internal mould of ventral valve incomplete, IPI 4502, scale bar = 1.5 cm; (J) internal mould of dorsal valve, IPI 4503, scale bar = 2.5 cm. (K, L) Kitakamithyris sp. A; (K) internal mould of dorsal valve, CEGHUNC 22905, scale bar = 2.35 cm; (L) internal mould of ventral valve, CEGH-UNC 22907, scale bar = 2.35 cm. (M, N) Kitakamithyris sp. B; (M) internal mould of




that represents the transition from maximum flooding interval (MFI), at the top of a transgressive systems tract (TST), to the highstand systems tract (HST) (López Gamundí and Martínez, 2000). In terms of paleoecology the Levipustula fauna is dominated by epifaunal bivalves, brachiopod spiriferids, and bryozoans, all suspension feeders that would have developed in some areas of the basin with open connection with the oceanic masses (Cisterna et al., 2013). Studies of the paleocommunities that integrate the Levipustula fauna in the Hoyada Verde Formation indicate a stable marine environment, such as an open shelf with moderate bottom currents and availability of nutrients spread in the water column. The variations recognized in these communities along the fossiliferous interval would have been controlled by substrate types and food supply fluctuations during the postglacial transgression (Cisterna, 1999; Cisterna and Sterren, 2010). These parameters would have played the most important role in the distribution of post-glacial faunas. The age of the Levipustula fauna and the relationship with its occurrence in Australia have been widely discussed in previous studies (Cisterna and Sterren, 2010; Taboada, 2010 and references therein). The recent radiometric data from Precordillera have allowed to constrain the age of this fauna to the interval late Serpukhovian–early Bashkirian (Césari et al., 2011). This age has been assigned by the correlation of the Hoyada Verde Formation that contains the typical Levipustula fauna, with the glacial deposits of the Guandacol Formation dated by U-Pb method (Gulbranson et al., 2010). 2.1.2. Aseptella-Tuberculatella/Rhipidomella-Micraphelia fauna The faunal assemblages that integrate the AseptellaTuberculatella/Rhipidomella-Micraphelia fauna consist mainly of brachiopods, bivalves, and gastropods, accompanied by conulariids, nautiloids, and corals; brachiopods and bivalves are dominant in term of generic richness (Fig. 3). Two brachiopod assemblages have been recognized in this fauna: the lower, the Aseptella-Tuberculatella assemblage, dominated by productids (Aseptella sp. aff. A. patriciae Simanauskas, Tuberculatella peregrina Reed, Overtoniinae indet., and Linoproductoidea indet.), accompanied by chonetids (Micraphelia indianae Simanauskas, Micraphelia ? sp.) and orthids (Rhipidomella sp., a new species of Rhipidomella pending to be formally described), and less abundant “inarticulates” brachiopods (lingulids indet. and Orbiculoidea sp.), athyrids indet. and terebratulids (Beecheria patagonica Amos); and the upper, the

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Rhipidomella-Micraphelia assemblage, where the chonetids (Micraphelia indianae) and orthids (Rhipidomella sp.) are dominant, is also accompanied by less abundant “inarticulates” brachiopods (Orbiculoidea sp.) and terebratulids (Beecheria patagonica Amos) (Fig. 5). The fossil assemblages that integrate this fauna appear very scattered in thick mudstone packages with calcareous concretions in the upper part of the glaciomarine succession of the El Paso Formation. The Aseptella-Tuberculatella assemblage, dominated by quasi infaunal brachiopods and infaunal detritus-feeding bivalves that suggest a soft-bottom with scarce material in suspension, occurs in the initial transgressive interval characterized by sediment starvation and relatively minor current circulation (López Gamundí and Martínez, 2003). Brachiopods and bivalves of the upper Rhipidomella-Micraphelia assemblage, which appears at the onset of the highstand systems tract (HST), show important paleoecological changes in relation with the lower assemblage Aseptella-Tuberculatella (i.e., decrease in taxonomic abundance, disappearance of the dominant guild and replacement of infaunal deposit feeders by epifaunal suspension feeders). This faunal turnover recognized along the two faunal assemblages could be explained as fluctuations in the depositional environment that range from starvation to shallowing conditions, reflecting higher energy and more active circulation of nutrients. The AseptellaTuberculatella/Rhipidomella-Micraphelia fauna would have developed in a relatively more isolated or restricted area (Cisterna et al., 2013), such as a “paleofjord”, similar to those described in other parts of the basin (Kneller et al., 2004). Some typical brachiopods of Aseptella-Tuberculatella/ Rhipidomella-Micraphelia fauna have also been identified from the other localities in the Calingasta-Uspallata Basin, i.e., the Ciénaga Larga del Tontal Formation in San Juan province, to the north and the Agua del Jagüel Formation in Mendoza province, to the south (Fig. 2A). The age of the Aseptella-Tuberculatella/RhipidomellaMicraphelia fauna has been recently defined by palynological data (Vergel et al., 2008, 2015) as late Serpukhovian–early Bashkirian, and it is discussed herein in the paragraph corresponding to biostratigraphic considerations. 2.2. Tarija Basin The Tarija Basin in the Sub Andean Bolivian area is developed mainly in Bolivia and only the southernmost part is located in Argentine territory (particularly in the Salta province and less

ventral valve, CEGH-UNC 22908, scale bar = 2.59 cm; (N) internal mould of dorsal valve, CEGH-UNC 22906, scale bar = 1.7 cm. (O–R) Septosyringothyris keideli (Harrington); (O) internal mould of dorsal valve, CEGH-UNC 22943, scale bar = 2.35 cm; (P) ventral valve, CEGH-UNC 22939, scale bar = 1.9 cm; (Q) specimen articulated, view of the ventral area, CEGH-UNC 22940, scale bar = 2 cm; (R) internal mould of ventral valve, CEGH-UNC 22941, scale bar = 1.85 cm. (S–W) Spiriferellina octoplicata (Sowerby); (S) internal mould of dorsal valve, CEGH-UNC 22931, scale bar = 1.27 cm; (T) external mould of ventral valve, CEGH-UNC 22932, scale bar = 0.95 cm; (U) internal mould of ventral valve, CEGH-UNC 22933, scale bar 0.85 cm; (V) ventral valve incomplete, IPI 3232, scale bar = 0.48 cm; (W) ventral valve, CEGH-UNC 22930, scale bar = 0.79 cm. (X, Y) Beecheria sp.; (X) internal mould of dorsal valve, IPI 4505, scale bar = 0.37 cm; (Y) internal mould of ventral valve, IPI 4504, scale bar = 0.43 cm. (A–E, I, J, V, X, Y) Material from the upper part of Hoyada Verde Formation (Cisterna and Sterren, 2010); (F–H, K–N) material from La Capilla Formation; (O–U, W) material from Leoncito Formation (Cisterna and Sterren, 2008). Repository: The fossils figured in this paper are registered by the prefix IPI (Instituto de Paleontología, sección Invertebrados) housed in the collections of the Miguel Lillo Foundation (San Miguel de Tucumán), and with repository number CEGH-UNC (Centro de Investigaciones Paleobiológicas) housed in the CICTERRA – Córdoba University, Argentina.




Fig. 5. Brachiopodos of Aseptella-Tuberculatella/Rhipidomella-Micraphelia fauna from the El Paso Formation. (A–C, E) Micraphelia indianae Simanauskas and Cisterna; (A, E) external mould of dorsal and ventral valve, CEGH-UNC 26714a-b, scale bar = 1.75 mm; (B) ventral valve partially decorticated, CEGH-UNC 26715, scale bar = 1.75 mm; (C) internal mould of ventral valve, CEGH-UNC 26716, scale bar = 3.25 mm. (D, F) Micraphelia ? sp.; (D) internal mould of ventral valve, CEGH-UNC 26721a, scale bar = 4.5 mm; (F) internal mould of dorsal valve, CEGH-UNC 26719, scale bar = 4.5 mm. (G–L) Tuberculatella peregrina (Reed); (G, J) internal and external mould of ventral valve, CEGH-UNC 26723a-b, scale bar = 4.5 mm; (H) internal mould of ventral valve, posterior view, CEGH-UNC 26725, scale bar = 4.5 mm; (I) internal mould dorsal valve, CEGH-UNC 26726, scale bar = 3.5 mm; (K) internal mould of ventral valve, CEGH-UNC 26724, scale bar = 4.5 mm; (L) dorsal external mould, CEGH-UNC 26727, scale bar = 4.5 mm. (M, N) Overtoniinae indet., internal and external mould of ventral valve, CEGH-UNC 26728, scale bar = 4.5 mm. (O–Q, T) Rhipidomella sp.; (O, Q) external and internal mould of ventral valve, CEGH-UNC 26706a-b, scale bar = 1.5 mm; (P, T) internal and




represented in the Jujuy province, Fig. 2B). The important sedimentary succession that characterizes this basin has been widely studied from the sequence stratigraphic viewpoint but the marine invertebrate faunas restricted to the north part of the basin in Bolivia are not well known. Glacial-postglacial Carboniferous sediments can be recognized in two Super-sequences, which coincide roughly with the Macharetí and Mandiyutí groups, and they are composed of sandstones and diamictites with silt and shale levels of 2000 m thick. The Tarija Formation is the most representative unit of the Macharetí Group and is characterized by an important diamictitic succession. The overlying postglacial succession of the Taiguatí Formation has been defined only in Bolivia. From this unit Trujillo Ikeda (1989) reports at least six fossiliferous localities with marine invertebrates. The fossil assemblage is dominated by bivalves, accompanied by less abundant brachiopods and gastropods, and scarce orthoceratids, crinoids, ichnites, and plant remains. The key brachiopod Levipustula levis Maxwell has been identified only in the Parapetí River (Charagua Hill locality). This species was illustrated by Rocha-Campos et al. (1977) from the upper part of the Taiguatí Formation and the interval containing this fauna was assigned to the Serpukhovian–Bashkirian interval. In the recent review of the brachiopods Levipustulini from Argentina, Taboada and Shi (2011) indicate that the material identified in Tarija Basin seems to be comparable in shell size to Levipustula specimens from western Argentina rather than to the Australian type material of Levipustula levis. 2.3. Tepuel Genoa Basin An important glacial-postglacial succession containing abundant invertebrate marine faunas, with a significant thicknesses that generally reach 2900 m, characterizes the Tepuel Genoa Basin located in the Argentine Patagonia. The Pampa de Tepuel and Las Salinas formations, in Chubut province, represent the stratigraphic record of the Carboniferous glacial event in this part of Gondwana. These units are characterized by diamictites and shales with dropstones and/or faceted clast, with sandstones to conglomerates intercalated (González, 1972; Page et al., 1984; Archangelsky et al., 1986; among other references). The fossiliferous horizons occur mainly in shale and siltstones levels throughout the whole diamictite interval. The most representative invertebrate groups in the Late Palaeozoic marine deposits of Patagonia are brachiopods, bivalves, gastropods, cephalopods, hyolithids, pelmatozoans, ostracods, and cnidarians. Two faunas related to the Carboniferous glacial episode were identified by Simanauskas and Sabattini (1997) to define the Lanipustula biozone (late Serpukhovian–Gzhelian, Simanauskas and Sabattini, 1997) and

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the youngest Tuberculatella biozone (Sakmarian, Simanauskas and Sabattini, 1997). 2.3.1. Lanipustula biozone (Simanauskas and Sabattini, 1997) Based on the database available from the Patagonian Basin (Pagani and Taboada, 2010), the Lanipustula fauna is dominated particularly by bivalves and bryozoans, accompanied by gastropods, echinoderms, ostracods, and cnidarians with scarce cephalopods and trilobites. Brachiopods appear to be less diversified and they are represented mainly by the key species Lanipustula patagoniensis Simanauskas and Lanipustula kletsi Taboada and Shi (Taboada and Shi, 2011), although the species Krotovia ? sp., Kitakamithyris cf. K. immensa (Campbell), Spiriferellina octoplicata ? (Sowerby), Spiriferellina ? sp., Alispirifer ? sp., Costuloplica sp., Spiriferiidae spp., and Beecheria ? sp. have been cited from the Pampa de Tepuel Formation (Pagani and Taboada, 2010). Brachiopods that characterize the Lanipustula biozone were recognized in the laterally equivalent Pampa de Tepuel and Las Salinas formations, which are exposed in at least two areas (Fig. 2C). Fossiliferous levels containing brachiopods of the Lanipustula biozone in these glaciomarine sequences occur in the diamictite-dominated interval, as well as in the upper postglacial deposits. The diagnostic species Lanipustula patagoniensis Simanauskas has been described from the middleupper part of the Pampa de Tepuel Formation (Amos, 1961; Simanauskas, 1996a; Taboada and Shi, 2011), and Lanipustula patagoniensis Simanauskas along with Lanipustula kletsi Taboada and Shi, in the lower to upper part of the Las Salinas Formation (Taboada and Shi, 2011). In both cases the species described occur in shale and siltstones beds located both below and above diamictite horizons. In the Las Salinas Formation that outcrops in the northern part of Languiñeo Hill (Fig. 2C), nine fossiliferous members based mainly on the bivalve faunas were described by González (1972). More recently, Pagani and Taboada (2010) resampled some of these members in an attempt to correlate this section with the type section of Pampa de Tepuel Formation at Tepuel Hill; these authors emphasized the paleontological importance of three members: LS 3 with bivalves, LS 5 characterized by a bryozoan fauna, and LS 9 that houses the most abundant and diverse fauna also containing the Lanipustula kletsi (Taboada and Shi, 2011). Besides, Alispirifer tranversus Maxwell and Neospirifer ? sp. have also been described from this last member (Cisterna, 1997). The presence of Levipustula levis Maxwell in Patagonia was mentioned by Amos (1979) in several localities of the Tepuel Genoa Basin, and by Archangelsky et al. (1986) in the Member LS 5 of the Las Salinas Formation. Simanauskas (1996a) redefined this material as Lanipustula patagoniensis and proposed

external mould of dorsal valve, CEGH-UNC 26712a-b, scale bar = 1.25 mm. (R, S) Beecheria patagonica Amos; (R) internal mould of articulated specimen, dorsal view, CEGH-UNC 26736, scale bar = 3.5 mm; (S) internal mould of articulated specimen, dorsal and ventral views, CEGH-UNC 26737, scale bar = 3.5 mm. (U–W) Aseptella sp. aff. A. patriciae Simanauskas; (U, V) external and internal mould of dorsal valve, CEGH-UNC 26731a-b, scale bar = 2.75 mm; (W) external mould of ventral valve, CEGH-UNC 26733, scale bar = 2.75 mm. Material comes from the lower fossiliferous interval of the El Paso Formation; (B, C, P, T, S), from the upper fossiliferous interval of this unit.




the Member LS 5 of the Las Salinas Formation as the parastratotype of the Lanipustula biozone, whereas the holostratotype was defined in the Sierra de Tepuel (level with “Fenestelids and Productids”, Simanauskas and Sabattini, 1997), in the Pampa de Tepuel Formation. Simanauskas (1996a) firstly discussed the presence of the genus Lanipustula in Patagonia. He reassigned the species Levipustula levis described by Amos from the Pampa de Tepuel Formation (Amos, 1961) to Lanipustula patagoniensis Simanauskas. The genus Lanipustula Klets was defined in the Carboniferous of Transbaikal region, Russia (Klets, 1983), to include species close to Levipustula but distinguishable from it by the position of the cardinal ridges and the shape and outline of anterior adductor scars: in Lanipustula the cardinal ridges do not diverge from the cardinal margin and the anterior adductor scars exhibit a teardrop shape. However, other authors have considered these characters as minor ‘intraspecific’ differences and proposed that the significant differences between Lanipustula and Levipustula are in the abundance of dorsal spines, and also in the development of concentric ornamentation observed in Lanipustula (Brunton et al., 2000; Taboada, 2006; Taboada and Shi, 2011). The age initially proposed by this fauna was late Serpukhovian–Gzhelian (Simanauskas and Sabattini, 1997), and then it was limited to the late Serpukhovian–Moscovian by Taboada (2008) and Taboada and Shi (2011). 2.3.2. Tuberculatella biozone (Simanauskas and Sabattini, 1997) This biozona, mainly composed of bivalves, brachiopods, gastropods, cephalopods and corals, was defined by Simanauskas and Sabattini (1997), with the holostratotype located in the Tepuel Hill, in the uppermost part of the Pampa de Tepuel Formation, and the parastratotypes in the Tecka and Languiñeo hills. The diagnostic brachiopods initially suggested to define this biozone were Tuberculatella laevicaudata (Amos), Aseptella patriciae Simanauskas, and Beecheria patagonica Amos (Simanauskas and Sabattini, 1997; Taboada et al., 2005). The age suggested to Tuberculatella Biozone by Simanauskas and Sabattini (1997) would be Sakmarian but it was widely discussed in further studies. Taboada (2008) indicates that the brachiopod assemblage that characterizes the Tuberculatella biozone in the type locality of Tepuel Hill, uppermost part of the Pampa de Tepuel Formation, is composed by the species Tuberculatella ? laevicaudata (Amos), Amosia sueroi Simanauskas, Verchojania archboldi Taboada, and Verchojania inacayali Taboada. Species of the genus Verchojania occur closely to the base and top of the biozone and below the Sakmarian Cimmeriella biozone, which appears in the overlying Mojón de Hierro Formation (Taboada, 2008). The genus Verchojania Abramov in Patagonia includes material previously described as Levipustula levis Amos (Amos, 1961). In this sense, Verchojania Abramov is a genus defined in the Upper Carboniferous (Bashkirian–Moscovian) of northern Asia (Abramov, 1970), which has been compared with Levipustula because its ornamentation pattern (Taboada, 2008). However, Verchojania would

have weaker concentric ornamentation especially on the dorsal valve, longer and abundant halteroid spines on flanks and ears and thin prostrate spines on the venter in relation with the stronger and suberect spines with more expanded spine bases over the venter that characterize the typical material of Levipustula, as it was indicated by Taboada (2008, p. 309). This author has also suggested that the genus Levipustula is non-sulcate, has shorter trail and lesser geniculate form, and possesses a different outline of ventral and dorsal muscle scars, in comparison with Verchojania. Based on the bryozoans associated with Verchojania inacayali Taboada, the base of the Tuberculatella biozone has been placed in the early Late Pennsylvanian (Gzhelian), and the top 700 m above in the uppermost levels of the Pampa de Tepuel Formation. This biozone would be no younger than Asselian since the Cimmeriella biozone of early Cisuralian age (Sakmarian) (Taboada, 2001; Simanauskas and Archbold, 2002; Taboada et al., 2005) occurs immediately above the Tuberculatella biozone (Taboada, 2008). 3. Biostratigraphic, paleoecological and paleobiogeographic considerations The studies of the postglacial marine invertebrates associated with the diamictite bearing sequences of the late Serpukhovian–Bashkirian interval in the Calingasta-Uspallata basin allowed to define two assemblages: the Levipustula fauna, widely known by previous studies (Cisterna and Sterren, 2008, 2010 and references therein), and the AseptellaTuberculatella/Rhipidomella-Micraphelia fauna, which has recently been proposed as a postglacial fauna. The significant compositional differences identified between these chronologically equivalent faunas have been explained by paleoecological controls (Cisterna et al., 2013): Levipustula fauna can be considered as a postglacial open sea fauna, whereas AseptellaTuberculatella/Rhipidomella-Micraphelia fauna would be a restricted marine assemblage developed in a relatively more isolated part of the basin. The age of AseptellaTuberculatella/Rhipidomella-Micraphelia fauna was widely discussed since its initial definition by Simanauskas and Cisterna (2001). These authors located the assemblages that integrate this fauna in the latest Carboniferous–earliest Permian interval, according the age of the Tuberculatella biozone in Patagonia, defined on the brachiopods Aseptella patriciae Simanauskas (Simanauskas, 1996b) and Tuberculalla laevicaudata (Amos) (Simanauskas, 1996c; Simanauskas and Sabattini, 1997). However, the palynological assemblages associated with this fauna in the El Paso Formation can be referred to the Subzone A of the Raistrickia densa-Convolutispora muriornata (DM) biozone (Césari et al., 2011), which indicates a late Serpukhovian–Bashkirian age (Vergel et al., 2008, 2015). This evidence as well as the sequence stratigraphic studies of the deposits containing the Levipustula fauna and the AseptellaTuberculatella/Rhipidomella-Micraphelia fauna, suggest that they are chronologically equivalent. Aseptella-Tuberculatella/Rhipidomella-Micraphelia fauna has also important intrabasinal implication, which has been




9

Fig. 6. Biostratigraphic scheme of the postglacial Carboniferous faunas studied (modified from Taboada, 2008; Pauls, 2014; Vergel et al., 2008, 2015). AT/RM, Aseptella-Tuberculatella/Rhipidomella-Micraphelia fauna. Black triangles indicate diamictitic levels.

recently discussed by Cisterna et al. (2013). Brachiopods of the assemblages that integrate this fauna are identified from the different glacial-postglacial transition sequences of the Calingasta-Uspallata Basin in Precordillera (Fig. 2A). In the lower part of the Ciénaga Larga del Tontal Formation, Lech et al. (1998) have described brachiopods of Aseptella-Tuberculatella assemblage (Aseptella ? sp. and Productella sp. (= Aseptella sp. aff. A. patriciae and Tuberculatella peregrina, Simanauskas and Cisterna, 2001)). To the south of the basin, near Uspallata locality (Fig. 2A), species of the Rhipidomella-Micraphelia assemblage (Rhipidomella sp. and Micraphelia indianae) have been identified associated with the early postglacial interval of the Agua del Jagüel Formation (Martínez et al., 2001; Simanauskas and Cisterna, 2001; Cisterna et al., 2013). Components of the postglacial faunas recognized in the west central of Argentina have also been identified in other South American basins (Cisterna and Sterren, 2015) (Fig. 6). In Bolivia, the invertebrate marine assemblage associated with the postglacial interval of the Taiguatí Formation (Macharetí Group) is not well known but the occurrence of the diagnostic species Levipustula levis Maxwell has been recently confirmed (Taboada and Shi, 2011). Levipustula fauna is the only postglacial fauna recognized to date in this basin, and it would represent the most northern record of Levipustula in South America. Some components that integrate the postglacial faunas of the Precordillera can be recognized also in the faunal assemblages that characterize the Lanipustula and Tuberculatella biozones in Patagonia. The Levipustula fauna could be considered chronologically equivalent to the Lanipustula biozone in the Patagonian Tepuel Genoa Basin. The age initially proposed for this biozone was late Serpukhovian–Gzhelian (Simanauskas and Sabattini, 1997), but later it was restricted to late Serpukhovian–Moscovian interval (Taboada, 2008; Taboada and Shi, 2011; Pauls, 2014). The occurrence of trilobites with the species Australosutura gardneri (Mitchell) in the Languiñeo and Tepuel hills has been taken as evidence to suggest a Carboniferous age (Amos et al., 1960). It is important to note that this biozone was renamed as Lanipustula

because the generic reassignment of the species Levipustula levis Maxwell (Amos, 1961) to Lanipustula patagoniensis Simanauskas (Simanauskas, 1996a). As it was mentioned above (see Section 2.3.1), Lanipustula Klets has been defined from the Carboniferous of Transbaikal region (Klets, 1983) and, even though Taboada and Shi (2011) suggest a possible occurrence in Europe, the genus has been described only in Patagonia besides Russia. On the other hand, as it was indicated by Taboada and Shi (2011), the validity of the genus has been questioned by some authors (Roberts et al., 1993, 1995). Although the mollusks (particularly bivalves) appear to be the dominant and more diversified component in the Lanipustula and Tuberculatella biozones of Patagonia (databases have been taken from Simanauskas and Sabattini (1997) and Taboada and Pagani (2010)), other invertebrate groups have been identified also in these biozones. As in the Levipustula fauna described from the Calingasta-Uspallata Basin, the bryozoans are an important component of the Lanipustula biozone. The brachiopod composition of Lanipustula would be also dominated by spiriferids (Kitakamithyris, Spiriferellina, Alispirifer, Costuloplica, and Spiriferiidae spp.), accompanied by productids (Lanipustula) and terebratulids (Beecheria), based always on the database of Pagani and Taboada (2010). Hence, in terms of paleoecology, the morphotypes identified in the Levipustula fauna of the Precordillera (Cisterna, 1999; Cisterna and Sterren, 2008, 2010) can be recognized in the Lanipustula biozone: the epifaunal liberosessile spiriferids and the productid Lanipustula adapted to a quasi infaunal mode of life. Brachiopod assemblages that characterize the Tuberculatella biozone in Patagonia appear to be dominated by productids (Tuberculatella, Amosia, Verchojania), as in the postglacial Aseptella-Tuberculatella/Rhipidomella-Micraphelia fauna from Precordillera. Also, some common species (Aseptella patriciae Simanauskas and Beecheria patagonica) have been previously described from the Tepuel-Genoa Basin (Amos, 1958; Simanauskas, 1996c). The occurrence of the genus Aseptella in both basins would have important paleobiogeographic implications. The oldest records of this genus are in the uppermost Visean–lower Serpukhovian horizons of the




Alba Formation (Winkler Prins and Martínez Chacón, 1998) in the Cantabrian Mountains (Spain), where the Aseptella was previously defined (Martínez Chacón and Winkler Prins, 1977). Argentinean species are particularly large in relation with the Spanish and a migration along the Northwest Gondwana margin, from the Iberian Peninsula through the Tethys Ocean, up to Gondwana realm, has been proposed to explain their occurrence in Calingasta-Uspallata and Tepuel-Genoa basins (Cisterna and Simanauskas, 1999). The genus could have evolved in species adapted to the glacial conditions existing to the Late Carboniferous in the southwestern margin of Gondwana. On the other hand, the presence of corals in the faunal assemblages of Aseptella-Tuberculatella/RhipidomellaMicraphelia fauna in the Calingasta-Uspallata Basin, as well as in the Tuberculatella biozone in Patagonia, should be noted mainly considering that corals are unusual in the Carboniferous glacial deposits of Argentina. Even though the Tuberculatella biozone exhibits some faunal affinity with the Aseptella-Tuberculatella/Rhipidomella-Micraphelia fauna from Precordillera, in Patagonia Tuberculatella biozone has a youngest record that reaches an Asselian age (Taboada, 2008; Pauls, 2014). Taboada (2010) and Taboada and Shi (2011) recognized in Patagonia a glacial episode extended from the latest Carboniferous to earliest Asselian, which would be absent in western Argentina where the youngest fauna identified (the Early Permian Costatumulus Fauna) is associated with an important climatic amelioration recognized in different central western Argentinean basins (Cisterna, 2010; Sterren and Cisterna, 2010). This amelioration of the climatic conditions was also registered at a global level in Gondwana and it would have started at the end of the Kasimovian–Gzhelian (López Gamundí, 1997; Limarino et al., 2014). This condition, as well as the volcanic activity and the action of relatively warmer marine currents, would be important local factors that controlled the benthic fauna distribution (Sterren and Cisterna, 2010). However, for the same interval of time Patagonia would be affected by a glacial episode according to Taboada (2010) and Taboada and Shi (2011). Recent radiometric data (U-Pb analysis of detrital zircons) from the Mojón de Hierro Formation located above of the horizons that contain the Tuberculatella biozone in Patagonia indicate a maximum age Late Carboniferous, constraining the timing of final deposition of glaciomarine sediments in the Tepuel-Genoa Basin (Griffis et al., 2014). According to the new data, the age of the Tubercualatella biozone from Patagonia would be the oldest in relation with that suggested by Taboada (2008), and its paleobiogeographic interpretation should be reconsidered. Taboada (2010) and Taboada and Shi (2011) suggested an important faunal contrast between western Argentina and Patagonia since middle Carboniferous times, and explained this difference in the context of the allochthonous Patagonia hypothesis (Ramos, 1984, 2008). These authors propose that Patagonia would have been located closer to the palaeopole than western Argentina according to a possible local block rotation of Patagonia (Rapalini, 2005), which would allow a displacement between its northern boundary and Gondwana up to about 1000 km in

the middle to late Carboniferous (Ramos, 2008; Rapalini et al., 2010). Even considering the allochthonous Patagonia hypothesis, the faunal assemblages herein analyzed from western Argentina and Tepuel Genoa basin do not exhibit a significant differentiation in relation with the brachiopods, in particular (Cisterna and Sterren, 2015), as well as in the relative abundance of the other invertebrate groups that integrate these faunas. The distribution of the postglacial faunas along the South America shows variations that would be controlled mainly by a paleolatitudinal factor: from the north (Tarija Basin) to south (Tepuel Genoa Basin), the postglacial faunas appear to have a clear differentiation. However, the paleoenvironmental control directly related with glacial retreat dynamics and the configuration of the coast, has also been considered to explain the important taxonomic, taphonomic, and paleoecological differences between the two contemporary postglacial faunas (Levipustula and AseptellaTuberculatella/Rhipidomella-Micraphelia) in the Barreal Hill, Calingasta-Uspallata Basin (Cisterna et al., 2013). This type of study could also help to understand the relationship between the faunal assemblages that integrate the Lanipustula and Tuberculatella biozones in Patagonia. 4. Conclusions The two postglacial faunas associated with the diamictite bearing sequences of the late Serpukhovian–Bashkirian interval in the west central Argentina (Levipustula fauna and the chronologically equivalent Aseptella-Tuberculatella/RhipidomellaMicraphelia fauna) are also recognized in other South American basins affected by the Carboniferous glacial episode in the western margin of Gondwana: the Tarija Basin in Bolivia and the Tepuel Genoa Basin in Patagonia. Along the South America, from the Tarija Basin in the north, to Tepuel Genoa Basin in the south, the postglacial faunas appear to have a clear differentiation that could be controlled mainly by a paleolatitudinal factor. The significant differences in faunal composition between Calingasta-Uspallata and Tepuel Genoa basins proposed by Taboada (2010) and Taboada and Shi (2011), because of the paleogeographical position of Patagonia considered to be a terrane accreted to Gondwana in the Late Paleozoic (Ramos, 2008), could not be confirmed in this paper. The two postglacial faunas described from the Calingasta-Uspallata Basin would have an equivalent in Patagonia. The Lanipustula Fauna dominated by bivalves and bryozoans with diversified spiriferid brachiopods can be compared with the Levipustula fauna from Precordillera. In terms of paleoecology, the morphotypes characterized by epifaunal liberosessile spiriferids and productids adapted to a quasi infaunal mode of life are recognized in both faunas. Brachiopod assemblages that characterize the Tuberculatella biozone in Patagonia appear to be dominated by productids as in the postglacial Aseptella-Tuberculatella/RhipidomellaMicraphelia fauna and these faunas would also have some common elements (Tuberculatella, Aseptella, Beecheria). The Tuberculatella biozone was assigned to the interval of early late Pennsylvanian (Gzhelian)–Asselian by the associated bryozoan




faunas, which allows to propose a glacial episode extended from the latest Carboniferous to earliest Asselian (Taboada, 2010). The occurrence in Patagonia of some components of the late Serpukhovian–Bashkirian Aseptella-Tuberculatella and Rhipidomella-Micraphelia assemblages as well as the new radiometric data from the Mojón de Hierro Formation located above of the horizons containing the Tuberculatella biozone in the Tepuel Hill (Griffis et al., 2014) questions the age of this fauna. Finally, paleoecological studies focused on the paleoenvironmental controls related with the glacial dynamic are suggested to understand the relationship between the faunal assemblages that integrate the Lanipustula and Tuberculatella biozones in Patagonia. Acknowledgements This study was supported by CONICET (Consejo Nacional de Investigaciones Científicas y Técnicas): PIP 0091 (GAC) and PIP 112-201101-00571 (AFS). The authors are especially grateful to the reviewers (Dr. María Luisa Martínez Chacón and the anonymous one), and to Ms. Si-Wei Chen (Editorial Office, Palaeoworld) for their valuable comments and suggestions. References Abramov, B.S., 1970. Biostratigrafiia kamennougol’nykh otlozhenii SetteDabana (Iuzhnoe Verkhoian’e). “Nauka”, Moskva, 178 pp. (in Russian). Amos, A.J., 1958. Some Lower Carboniferous brachiopods from the Volcan Formation, San Juan, Argentina. Journal of Paleontology 32, 838–845. Amos, A.J., 1961. Algunos Chonetacea y Productacea del Carbonífero inferior y superior del Sistema de Tepuel, Provincia de Chubut. Revista de la Asociación Geológica Argentina 15, 81–107. Amos, A.J., 1979. Guía Paleontológica Argentina. Parte I: Paleozoico, faunas carboníferas y pérmicas. Consejo Nacional de Investigaciones Científicas y Técnicas, Buenos Aires, 154 pp. Amos, A.J., Campbell, K.S.W., Goldring, R., 1960. Australosutura gen. nov, (Trilobita) form the Carboniferous of Australia and Argentina. Palaeontology 3, 227–236. Amos, A.J., Rolleri, E.O., Csaky, A., 1963. La fauna del Carbonífero medio de la Formación La Capilla y sus relaciones geológicas. Ameghiniana 3, 123–132. Archangelsky, S., Azcuy, C., González, C.R., Sabattini, N., 1986. Correlación general de las biozonas. In: Archangelsky, S. (Ed.), El Sistema Carbonífero en la República Argentina. Academia Nacional de Ciencias de Córdoba, Córdoba, pp. 281–292. Brunton, C.H.C., Lazarev, S.S., Grant, R.E., Jin, Y.G., 2000. Productidina. In: Kaesler, R.L. (Ed.), Treatise on Invertebrate Paleontology, Part H, Brachiopoda 3 (revised). Geological Society of America and University of Kansas Press, Boulder and Lawrence, pp. 424–565. Césari, S.N., Limarino, C.O., Gulbranson, E.L., 2011. An Upper Paleozoic biochronostratigraphic scheme for the western margin of Gondwana. Earth Science Reviews 106, 149–160. Cisterna, G.A., 1997. Spiriferida (Brachiopoda) en la Formación Las Salinas, Carbonífero superior, provincia de Chubut, Argentina. Ameghiniana 34, 155–161. Cisterna, G.A., 1999. Paleoecología de niveles pelíticos de la sección superior de la Formación Hoyada Verde, Carbonífero Superior, Precordillera de San Juan, Argentina. Ameghiniana 36, 259–267. Cisterna, G.A., 2010. Earliest Permian brachiopod faunas of central western Argentina: implications for defining the Carboniferous–Permian boundary. Palaeogeography, Palaeoclimatology, Palaecology 298, 91–100.

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Cisterna, G.A., Simanauskas, T., 1999. Aseptella (Brachiopoda) en el Paleozoico tardío de Argentina. Revista Española de Paleontología 14, 117–122. Cisterna, G.A., Sterren, A.F., 2008. Late Carboniferous Levipustula fauna in the Leoncito Formation, San Juan province, Argentine Precordillera: Biostratigraphical and palaeoclimatological implications. Proceedings of the Royal Society of Victoria 118, 137–147. Cisterna, G.A., Sterren, A.F., 2010. Late Carboniferous “Levipustula Fauna” in central-western Argentina and its relationships with the Carboniferous glacial event in the southwestern Gondwanan margin. In: López Gamundí, O.R., Buatois, L.A. (Eds.), Late Paleozoic Glacial Events and Postglacial Transgressions in Gondwana. Geological Society of America Special Paper 468, pp. 133–147. Cisterna, G.A., Sterren, A.F., 2015. Carboniferous postglacial brachiopod faunas in the southwestern Gondwana margin. Permophiles, The Brachiopod World, Abstract for IBC7, pp. 21–23. Cisterna, G.A., Sterren, A.F., Vergel, M.M., López Gamundí, O.R., 2013. Reinterpretación bioestratigráfica de las faunas postglaciales carboníferas en la Cuenca Calingasta-Uspallata, Argentina. 6th Simposio Argentino del Paleozoico Superior, Abstract, Buenos Aires, p. 9. González, C.R., 1972. La Formación Las Salinas del Paleozoico superior de Patagonia (República Argentina). Parte I: Estratigrafía, facies y ambientes de sedimentación. Revista de la Asociación Geológica Argentina 27, 95–115. Griffis, N.P., Motañez, I.P., Isbell, J., Gulbranson, E.L., Wimpenny, J., Yin, Q.Z., Cúneo, N.R., Pagani, M.A., Taboada, A.C., 2014. U-Pb detrital zircon geochronologic constraints on depositional age and sediment source terrains of the Late Paleozoic Tepuel-Genoa Basin. American Geophysical Union, Fall Meeting 2014, Abstract, EP21D-3566. Gulbranson, E.L., Montañez, I.P., Schmitz, M.D., Limarino, C.O., Isbell, J.L., Marenssi, S.A., Crowley, J.L., 2010. High-precision U-Pb calibration of Carboniferous glaciation and climate history, Paganzo Group, NW Argentina. Geological Society of America Bulletin 122, 1480–1498. Keidel, J., Harrington, H.J., 1938. On the discovery of Lower Carboniferous tillites in the Precordillera of San Juan. Geological Magazine 75, 103–129. Klets, A.G., 1983. Novyi kamennougol’nyi rod produktid. Paleontologicheskii Zhurnal 1, 74–78 (in Russian). Kneller, B., Milana, J.P., Buckee, C., Al Ja’aidi, O., 2004. A depositional record of deglaciation in a paleofjord (Late Carboniferous [Pennsylvanian] of San Juan Province, Argentina): The role of catastrophic sedimentation. Geological Society of America Bulletin 116, 348–367. Lech, R.R., Milana, J.P., Banchig, A.L., 1998. Braquiópodos del Carbonífero de la Sierra del Tontal, Provincia de San Juan, Argentina. Ameghiniana 35, 405–413. Limarino, C.O., Césari, S.N., Spalletti, L.A., Taboada, A.C., Isbell, J.L., Geuna, S., Gulbranson, E.L., 2014. A paleoclimatic review of southern South America during the late Paleozoic: A record from icehouse to extreme greenhouse conditions. Gondwana Research 25, 1396–1421. López Gamundí, O.R., 1997. Glacial–postglacial transition in the late Paleozoic basins of Southern South America. In: Martini, I.P. (Ed.), Late Glacial and Postglacial Environmental Changes: Quaternary Carboniferous–Permian, and Proterozoic. Oxford University Press, Oxford, pp. 147–168. López Gamundí, O.R., Martínez, M., 2000. Evidence of glacial abrasion in the Calingasta-Uspallata and western Paganzo basins, mid-Carboniferous of western Argentina. Palaeogeography, Palaeoclimatology, Palaeoecology 159, 145–165. López Gamundí, O.R., Martínez, M., 2003. Esquema estratigráfico-secuencial para las unidades neopaleozoicas de la cuenca Calingasta-Uspallata en el flanco occidental de la Precordillera. Revista de la Asociación Geológica Argentina 58, 367–382. Martínez Chacón, M.L., Winkler Prins, C.F., 1977. A Namurian brachiopod fauna from Meré (Province of Oviedo, Spain). Scripta Geologica 39, 1–67. Martínez, M., Cisterna, G.A., Sterren, A.F., 2001. La Formación Agua del Jaguel (Cuenca Calingasta-Uspallata): reconsideraciones estratigráficas y bioestratigráficas. 2nd Simposio Argentino del Paleozoico Superior, Abstract, Trelew, p. 18. Pagani, M.A., Taboada, A.C., 2010. The marine Upper Palaeozoic in Patagonia (Tepuel-Genoa Basin, Chubut province, Argentina): 85 years of work and future prospects. Palaeogeography, Palaeoclimatology, Palaeoecology 298, 130–151.




Page, R., Limarino, C.O., López Gamundí, O.R., Page, S., 1984. Estratigrafía del Grupo Tepuel en su perfil tipo y en la región de El Molle, provincia de Chubut. 9th Congreso Geológico Argentino, Acta 1, Bariloche, pp. 619–632. Pauls, K.N., 2014. Sedimentology and paleoecology of fossil-bearing, highlatitude marine and glacially influenced deposits in the Tepuel Basin, Patagonia, Argentina. PhD Dissertation, University of Milwaukee, Milwaukee, 420 pp. Ramos, V.A., 1984. Patagonia: Un continente paleozoico a la deriva? 9th Congreso Geológico Argentino, Acta 2, Bariloche, pp. 311–325. Ramos, V.A., 2008. Patagonia: a paleozoic continent adrift? Journal of South American Earth Sciences 26, 235–251. Rapalini, A.E., 2005. The accretionary history of southern South America from the latest Proterozoic to the Late Paleozoic: some paleomagnetic constraints. Terrane Processes at the Margins of Gondwana. Special Publications of the Geological Society 246, 305–328. Rapalini, E., López De Luchi, M., Martínez Dopico, C., Linc Klinger, F., Giménez, M., Martínez, P., 2010. Did Patagonia collide with Gondwana in the Late Paleozoic? Some insights from a multidisciplinary study of magmatic units of the North Patagonian Massif. Geologica Acta 8, 349–371. Roberts, J., Jones, P.J., Jenkins, T.B.H., 1993. Revised correlations for Carboniferous marine invertebrate zones of eastern Australia. Alcheringa 17, 353–376. Roberts, J., Claoué-Long, J.C., Jones, P.J., Foster, C.B., 1995. SHRIMP zircon age control of Gondwanan sequences in Late Carboniferous and early Permian Australia. In: Dunay, R.E., Hailwood, E.A. (Eds.), NonBiostratigraphical Methods of Dating and Correlation. Special Publications of the Geological Society 89, pp. 145–174. Rocha-Campos, A., Carvalho, R., Amos, J., 1977. A Carboniferous (Gondwana) fauna from subandean Bolivia. Revista Brasileira de Geociencias 7, 287–303. Simanauskas, T., 1996a. Una nueva especie de Lanipustula (Productoidea, Brachiopoda) del Paleozoico superior de Patagonia, Argentina. Ameghiniana 33, 301–305. Simanauskas, T., 1996b. Braquiópodos (Productoidea) del Pérmico de la Cuenca Tepuel-Genoa, Chubut, Argentina. Ameghiniana 33, 377–383. Simanauskas, T., 1996c. Un supuesto chonetoidea del Pérmico de Patagonia, reasignado a un género de Productoidea (Brachiopoda). Ameghiniana 33, 349–351. Simanauskas, T., Archbold, N.W., 2002. Early Permian Jakutoproductus (Productida: Brachiopoda) from Patagonia, Argentina. Alcheringa 26, 465–474. Simanauskas, T., Cisterna, G.A., 2001. Braquiópodos articulados de la Formación El Paso, Paleozoico superior, Precordillera Argentina. Revista Española de Paleontología 16, 209–222. Simanauskas, T., Sabattini, N., 1997. Bioestratigrafía del Paleozoico superior marino de la Cuenca Tepuel–Genoa, provincia de Chubut, Argentina. Ameghiniana 34, 49–60.

Sterren, A.F., Cisterna, G.A., 2010. Bivalves and brachiopods in the Carboniferous–Early Permian of Argentine Precordillera: diversification and faunistic turnover. Geologica Acta 8, 501–517. Taboada, A.C., 1989. La fauna de la Formación El Paso, Carbónico inferior de la precordillera sanjuanina. Acta Geológica Lilloana 17, 113–129. Taboada, A.C., 2001. Bioestratigrafía del Neopaleozoico del Valle de Tres Lagunas, sierra de Tepuel, Provincia de Chubut. Acta Geológica Lilloana 18, 291–304. Taboada, A.C., 2006. Levipustula Maxwell, 1951 y Lanipustula Klets, 1983 (Brachiopoda, Levipustulini) en Argentina: revisión preliminar. 9th Congreso Argentino de Paleontología y Bioestratigrafía, Abstract, Córdoba, p. 193. Taboada, A.C., 2008. First record of the Late Paleozoic brachiopod Verchojania in Patagonia, Argentina. Proceedings of the Royal Society of Victoria 120, 305–319. Taboada, A.C., 2010. Mississippian–Early Permian brachiopods from western Argentina: tools for middle- to high-latitude correlation, paleobiogeographic and paleoclimatic reconstruction. Palaeogeography, Palaeoclimatology, Palaeoecology 298, 152–173. Taboada, A.C., Cisterna, G.A., 1996. Elythinae (Bachiopoda) del Paleozoico Superior de Argentina. Ameghiniana 33, 83–94. Taboada, A.C., Pagani, M.A., 2010. The coupled occurrence of CimmeriellaJakutoproductus (Brachiopoda: Productidina) in Patagonia: implications for Early Permian high to middle paleolatitudinal correlations and paleoclimatic reconstruction. Geologica Acta 8, 519–540. Taboada, A.C., Shi, G.R., 2011. Taxonomic review and evolutionary trends of Levipustulini and Absenticostini (Brachiopoda) from Argentina: Palaeobiogeographic and palaeoclimatic implications. Memoirs of the Association of Australasian Palaeontologists 41, 87–114. Taboada, A.C., Archbold, N.W., González, C.R., Sabattini, N., 2005. The Late Carboniferous Early Permian Tepuel fauna of Patagonia: updated brachiopods records. In: Pankhurst, R.J., Veiga, G.D. (Eds.), Geological and Biological Heritage of Gondwana, Gondwana 12. Academia Nacional de Ciencias, Córdoba, p. 349. Trujillo Ikeda, H., 1989. Nuevo hallazgo de fósiles de la Formación Taiguati en la Serranía Caipipendi, Santa Cruz, Bolivia. Revista Técnica de Yacimientos Petrolíferos Fiscales Bolivianos 10, 7–11. Vergel, M.M., Cisterna, G.A., Sterren, A.F., 2008. Primera evidencia palinológica en la Formación El Paso, Paleozoico Superior, Precordillera Argentina. Acta Geológica Lilloana 21 (Suplemento), 75–77. Vergel, M.M., Cisterna, G.A., Sterren, A.F., 2015. New palynological records from the glaciomarine deposits of the El Paso Formation (late Serpukhovian–Bashkirian) in Argentine Precordillera: biostratigraphical implications. Ameghiniana 52, 334–349. Winkler Prins, C.F., Martínez Chacón, M.L., 1998. A new species of the productid brachiopod Aseptella from the Lower Carboniferous of the Cantabrian Mountains (Spain). Revista Española de Paleontología 13, 243–249.
