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Macroinvertebrates of the Capivari marine bed, late Paleozoic glacial Itararé Group, northeast Paraná Basin, Brazil: Paleoenvironmental and paleogeographic implications
Journal Pre-proof Mollusks and brachiopods of the Capivari marine bed, late Paleozoic glacial Itararé Group, northeast Paraná Basin, Brazil: Paleoenvironmental and paleogeographic implications Marcello Guimarães Simões, Jacqueline Peixoto Neves, Arturo César Taboada, Maria Alejandra Pagani, Filipe Giovanini Varejão, Mário Luis Assine PII:
S0895-9811(19)30443-2
DOI:
https://doi.org/10.1016/j.jsames.2019.102433
Reference:
SAMES 102433
To appear in:
Journal of South American Earth Sciences
Received Date: 27 August 2019 Revised Date:
19 November 2019
Accepted Date: 20 November 2019
Please cite this article as: Simões, Marcello.Guimarã., Neves, J.P., Taboada, Arturo.Cé., Pagani, M.A., Varejão, F.G., Assine, Má.Luis., Mollusks and brachiopods of the Capivari marine bed, late Paleozoic glacial Itararé Group, northeast Paraná Basin, Brazil: Paleoenvironmental and paleogeographic implications, Journal of South American Earth Sciences (2019), doi: https://doi.org/10.1016/ j.jsames.2019.102433. This is a PDF file of an article that has undergone enhancements after acceptance, such as the addition of a cover page and metadata, and formatting for readability, but it is not yet the definitive version of record. This version will undergo additional copyediting, typesetting and review before it is published in its final form, but we are providing this version to give early visibility of the article. Please note that, during the production process, errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. © 2019 Published by Elsevier Ltd.
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Mollusks and brachiopods of the Capivari Marine Bed, late Paleozoic glacial
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Itararé Group, northeast Paraná Basin, Brazil: paleoenvironmental and
3
paleogeographic implications
4 5
Marcello Guimarães Simões
6
Instituto de Biociências, Departamento de Zoologia, Universidade Estadual Paulista, Distrito de
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Rubião Junior, Botucatu, São Paulo, Brazil.
8 9
Jacqueline Peixoto Neves*
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Universidade Tecnológica Federal do Paraná, campus Dois Vizinhos, Dois Vizinhos, Paraná,
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Brazil.
12
*Corresponding autor
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Arturo César Taboada
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Centro de Investigación Esquel de Montaña y Estepa Patagónicas, CONICET-UNPSJB, Esquel
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U9200, Chubut, Argentina.
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Maria Alejandra Pagani
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Museo Paleontológico Egidio Feruglio – CONICET, Avenida Fontana nº 140, Trelew
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U9100GYO, Chubut, Argentina.
21 22
Filipe Giovanini Varejão
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Instituto de Geociências e Ciências Exatas, Departamento de Geologia Aplicada, Universidade
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Estadual Paulista, Campus de Rio Claro, Rio Claro, São Paulo, Brazil.
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26 27
Mário Luis Assine
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Instituto de Geociências e Ciências Exatas, Departamento de Geologia Aplicada, Universidade
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Estadual Paulista, Campus de Rio Claro, Rio Claro, São Paulo, Brazil.
30 31
1
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Abstract.—A 2-m-thick silty shale bed within the Taciba Formation, Itararé Group, Paraná
33
Basin, State of São Paulo, southeastern Brazil, records marine sedimentation in a siliciclastic-
34
dominated, low-energy, shelf setting, during a short-lived deglacial event. The bed is located
35
100–150 m below the base of the lower Permian, post-glacial Tatui Formation. The marine
36
assemblage is dominated by rhynchonelliform brachiopods, with subordinate bivalves,
37
gastropods and crinoids, recording the highest phylum-level diversity so far identified within a
38
given fossil-bearing horizon in the uppermost portion of the Itararé Group. Two new species are
39
described, one brachiopod Biconvexiella saopauloensis and one gastropod Peruvispira
40
brasilensis. Additionally, shells of Lyonia rochacamposi, Rhynchopora grossopunctata,
41
Quinquenella rionegrensis, Phestia tepuelensis, Streblopteria aff. S. lagunensis, Limipecten
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capivariensis, Praeundulomya cf. subelongata and Mourlonia (Woolnoughia)? sp.are identified.
43
Crinoid columns were assigned to øPentaridica sp. (a genus based on elements of the columnal).
44
This is the first systematic description of members of the Eurydesma-Lyonia fauna in the
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northeastern part of the Paraná Basin, Brazil. The overwhelming majority of brachiopods belong
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to Biconvexiella saopauloensis, followed by Rhynchopora grossopunctata. The record of Lyonia
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rochacamposi closely resembles that of the uppermost part of the Taciba Formation in southern
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Brazil. Hence, the Capivari marine fauna correlates approximately with that of the upper part of
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the Taciba Formation. Lyonia rochacamposi also indicates correlation with Permian units of the
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Sauce Grande-Colorado (Argentina), Huab (Hardap shale of the Dwyka Group), Aranos area
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(Namibia), southwest Africa, and the Carnavon (Western Australia) basins. These correlations
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support a latest Asselian-earliest Sakmarian age for the fauna.
53
2
54
Highlights
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A marine cool-water benthic fauna developed during Late Paleozoic Ice Age is described;
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Fauna developed during a transgression coupled with a time of climatic amelioration;
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Assemblage flourisched under restrict paleogeograhic conditions.
58 59
Keywords: Permian, Taciba Formation, macroinvertebrate, Paraná Basin, Capivari.
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1. Introduction
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Glaciogenic rocks of the Carboniferous-Permian Itararé Group, Paraná Basin, Brazil, are known
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since Derby (1878) and White (1908). The succession includes one of the most complete
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sedimentary records of the Late Paleozoic Ice Age (LPIA, sensu Shi and Waterhouse, 2010) that
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waxed and waned across the central Gondwana (Rocha-Campos, 1967; França and Potter, 1988;
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Holz et al., 2010). However, due to the large size of the basin (>1,500,000 km2) and the complex
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depositional processes associated with the glacial and post-glacial events, no consensus exists
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about the correlation, age and extent of the Itararé Group (Loczy, 1964; Rocha-Campos, 1967;
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1970; Saad, 1977; Rocha-Campos and Rösler, 1978; França and Potter, 1991; Castro, 1999; Holz
70
et al., 2010).
71
Within the glaciomarine successions of the Itararé Group, as shown by Santos et al. (1996),
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the sea-level rise following ice retreats was often accompanied by widespread marine deposits,
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usually ice-rafted, debris-free, dark-gray, fossil-poor shales. Five regionally continuous
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stratigraphic intervals with marine beds are recognized (Saad, 1977; Rocha-Campos and Rösler,
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1978; Petri and Souza, 1993; Santos et al., 1996; Holz et al., 2010), which are known by their
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local geographic names, as follows: (1) Araçoiaba, (2) Mafra-Ortigueira, (3) Lontras-Guaraúna
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(upper Pennsylvanian), (4) Capivari, and (5) Passinho (Rocha-Campos and Rösler, 1978; Santos
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et al., 1996), whose stratigraphic position within the glacial succession varies according to 3
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distinct authors (Tables 1, 2). These marine beds record extensive transgressive events associated
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with regional episodes of glacier retreats (see Loczy, 1964, p. 37; França and Potter, 1988;
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Castro, 1999; Vesely and Assine, 2006; Mineropar, 2007; Neves et al., 2014a, b; Taboada et al.,
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2016). Hence, some of these deposits (i.e., Lontras and Passinho shales) may have represented
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the maximum flooding surfaces within the depositional sequence (Vesely and Assine, 2006;
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Holz et al., 2008; 2010). Yet, within these marine beds at least eight distinct marine benthic
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assemblages are known (Rocha-Campos and Rösler, 1978; Neves et al., 2014a, b; Taboada et al.,
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2016).
87
As shown in Table 2, at the outcrop belt of the eastern border of the Paraná Basin, fossil-
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rich marine intercalations of the Itararé Group are difficult to correlate regionally (Holz et al.,
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2010). This is the case for the fossil-bearing beds of the Taciba Formation (former Capivari
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Formation, Table 3) exposed in the central-eastern region of the State of São Paulo, southeastern
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Brazil (Figs. 1 and 2). Some authors have correlated these fossil-rich marine beds with the
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Lontras shale (Soares, 1991; Castro, 1999) at the top of the Campo Mourão Formation (França
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and Potter, 1991). On the other hand, Loczy (1964) and Rocha-Campos (1969) correlated the
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Capivari fossil beds with the marine assemblages of the Teixeira Soares marine beds (Passinho
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shale and associated rocks), uppermost part of the Itararé Group, State of Paraná, southern Brazil.
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Alternatively, Taboada et al. (2016) positioned the Capivari fossil-rich beds in between the
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Lontras and Passinho fossil-bearing shales of the southern Brazil (Figs. 1, 2).
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The precise taxonomic composition, and age of the fossil-bearing beds of the Capivari
99
region (see Fig. 2) can shed light on intrabasinal biocorrelation of the Late Paleozoic marine
100
invertebrates faunas of the Paraná Basin with stratigraphic and paleogeographic implications for
101
the depositional history of the glacial succession of the Itararé Group. Therefore, in this
102
contribution we will attempt to better constrain those fossil-bearing beds. To achieve this, we
4
103
reassessed the invertebrate fauna originally described by Mendes (1952), also complementing
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our analysis with new fossil findings.
105 106
1.1. Stratigraphic framework
107
During the Paleozoic, the intraplate Paraná Basin was a large depositional area in the SW
108
Gondwanaland (central/southeastern part of the South America) (Milani, 1997). The basin was
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elongated in the N-S direction and filled with Late Ordovician to Late Cretaceous sedimentary
110
rocks, encompassing six supersequences or unconformity-bounded units (Milani, 1997). The
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2,500-m-thick succession of upper Paleozoic to Mesozoic rocks is referred as the Gondwana I
112
Supersequence (Milani, 1997; Milani et al., 2007) that was generated in a variety of continental,
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transitional and marine sedimentary settings, including upper Carboniferous-Permian glacial to
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Triassic arid depositional environments.
115
Rocks of the glacial succession are included in the Itararé Group (eastern border), which is
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one of the most complete stratigraphic successions of the LPIA in southwestern Gondwana.
117
Resting either directly upon Precambrian basement, lower Paleozoic or on Devonian beds, the
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Itararé Group is the thickest sedimentary package of the Paraná Basin, sedimentation lasting for
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at least 36 million years (França and Potter, 1991). Conglomerates, diamictites, sandstones,
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mudstones, rhythmites, turbidites, and thin coal beds are the main lithotypes of the group
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(Rocha-Campos, 1967; França and Potter, 1991; Holz et al., 2010). The precise age of the rocks
122
of the Itararé Group is one of the main issues of the Gondwana geology in Brazil. The Itararé
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Group is usually referred as upper Mississippian-lower Cisuralian in age (Souza and Marques-
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Toigo, 2005; Guerra-Sommer et al., 2008a, b; Rocha-Campos et al., 2008; Holz et al., 2010;
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Neves et al., 2014b; Taboada et al., 2016). However, U–Pb radiometric ages have dated the
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topmost glacial deposits of the Itararé Group at 307.7 ± 3.1 Ma (Kasimovian−Moscovian,
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Pennsylvanian, late Carboniferous) and the base of the overlying post-glacial deposits of the 5
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coal-bearing Rio Bonito Formation at 298.8 ± 1.9 Ma (Ghzelian−Asselian, Carboniferous–
129
Permian boundary) (Cagliari et al., 2014; 2016). Indeed, a radiometric Carboniferous age for the
130
upper Itararé Group was reinforced by newely U–Pb laser ablation ICP-MS detrital zircon dates
131
of rocks of this unit (Tedesco et al., 2019). However, we are following the chronostratigraphic
132
chart of Holz et al. (2010), which present the most complete information of age for the entire
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Itararé Group (Fig. 1).
134
Since the seminal work by Barbosa and Almeida (1949a), various lithostratigraphic
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subdivisions for the Itararé Group appeared in the literature. In southeastern Brazil, the group
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was subdivided into several formations, which include, in ascending order, the Itu, Capivari,
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Gramadinho, Tietê and Itapetininga formations (Barbosa and Almeida, 1949a). However, the
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original designations of these authors are now in disuse. Schneider et al. (1974) proposed an
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alternate litostratigraphic framework for outcrops in southern Brazil, in which the Itararé
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comprises three main units, including from the base to top, the Campo do Tenente, Mafra, and
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Rio do Sul formations.
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Based on a detailed subsurface analysis, França and Potter (1991) recognized three fining-
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upward glacially-influenced sedimentary cycles and classified them as formations named, from
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base to top, as Lagoa Azul, Campo Mourão and Taciba. These authors also noted that in the
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subsurface the maximum thickness (~1300 m) of the Itararé Group is reached in the State of São
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Paulo, where our study area is located (Fig. 1).
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Trying to correlate the glacial cycles defined by França and Potter (1991) in outcrop area
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of the Itararé Group, Vesely and Assine (2006) recognized five glacially-influenced depositional
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sequences in the State of Paraná. These are also laterally traceable for nearly 400 km in well logs
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across the central portion of the basin. The oldest depositional sequence was correlated with the
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Lagoa Azul Formation of França and Potter (1991), the youngest with the Taciba Formation, and
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the three middle sequences with the whole Campo Mourão Formation. A retrogradational 6
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stratigraphic stacking pattern characterizes all these cycles, which correspond to major ice-retreat
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phases or deglaciation sequences (Vesely and Assine, 2006). Complex facies associations are
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generated during deglaciation, mainly because tillites and outwash facies formed during an
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oscillatory but overall glacier-retreat and are interfingered with contemporaneous glacial-marine
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deposits.
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In the outcrops of the studied area, the Taciba Formation is equivalent to the Capivari
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Formation (mainly fine-grained facies) and the overlying Gramadinho Formation (mainly
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diamictites) of Barbosa and Almeida (1949a, b). Yet, the Taciba Formation is the youngest and
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most widespread unit of the Itararé Group, corresponding to the Rio do Sul Formation of
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Schneider et al. (1974). Finally, as commented above, the Taciba Formation is the sedimentary
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record of the last deglacial sequence (sequence 5 of Vesely and Assine, 2006), representing the
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final ice-retreat in the basin, onlapping the south border of the basin.
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Based on subsurface data from three boreholes (C-IG/93, R-IG/94 and R-IG/95) drilled in
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the Capivari and Rafard counties, Petri et al. (1996) subdivided the upper part of the Itararé
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Group into four main sedimentary packages, as follows (from the base to the top): (a) package D,
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represented by conglomerates and coarse sandstones; (b) package C that includes mudstones,
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siltstones and rhythmites; (c) package B, which is dominated by sandstones, and (d) package A,
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represented by transgressive mudstones. This succession is, in general, similar to that recorded
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by Saad (1977) to the Itararé Group in the south and central portions of the State of São Paulo.
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1.2. Background: the Capivari fossil bed unearthed
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The marine macroinvertebrate assemblage herein described comes from a ~2-m-thick interval of
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silty shales recorded in the middle part of the Taciba Formation, upper part of the Itararé Group
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in the State of São Paulo, formerly refered to the Capivari Formation (Figs. 1 and 2).
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Theassemblage is thought to be the most diverse one of the Itararé Group (Rocha-Campos and 7
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Rösler, 1978; Simões et al., 1998), and is 500 km far away of those invertebrate-rich beds of the
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Taciba Formation (Passinho Shale) from southern Brazil (Neves et al., 2014b; Taboada et al.,
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2016) (Fig. 2). As described and commented below, the fossil-bearing silty shale bed crops out
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in the vicinity of the Capivari county (Fig. 2). Fossils from these localities were first discovered
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in 1947 by F.F.M. Almeida, S. Petri and O. Barbosa (see information in Mendes, 1952), but the
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first publication on these fossil-bearing sites appeared in the literature only two years later
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(Barbosa and Almeida 1949a). It was only in March of 1949, in a short note to the Brazilian
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Academy of Sciences, that Barbosa and Almeida (1949a) briefly discussed the stratigraphic
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position of the Capivari marine fauna within the Itararé Group. In August of that year, a more
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comprehensive and detailed study of the geology and stratigraphy of the area was published. By
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combining both subsurface and surface data, Barbosa and Almeida (1949b) constrained the
189
marine assemblage to the Capivari Formation and noted that the fossil bed is intercalated
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between rhythmites and diamictites. The authors recorded the presence of brachiopods
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(Ambocoelia, Rhynchopora), bivalves (Aviculopecten), and gastropods (that were subsequently
192
lost, see Rocha-Campos, 1966, p. 10), but the fossils were neither described, nor illustrated.
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Despite this, Barbosa and Almeida (1949b) noted that the fossils are not identical to those
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already described from the fossil-rich beds of the Itararé Group (i.e., Passinho shale and coeval
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strata of the Teixeira Soares region, State of Paraná) from the State of Paraná, southern Brazil.
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J.C. Mendes formally described the marine macroinvertebrate fossils of the “Capivari beds”
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in 1952, basing on those specimens previously collected by Barbosa and Almeida (1949a, b),
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plus additional material offered by the geologist S. Mezzalira. He reported the presence of
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rhynchonelliform brachiopods (Crurithyris aff. planoconvexa, Rhynchopora grossopunctata)
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and bivalves (Aviculopecten capivariensis n. sp., Nuculana? sp., and undetermined specimens)
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(Mendes, 1952). Subsequently, Rocha-Campos (1966) recorded the presence of the gastropod
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Peruvispira delicata. 8
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After the contributions above, the marine fauna of the Capivari beds was almost forgotten
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for over twenty years, mainly because the outcrops of the Itararé Group in the region are deeply
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weathered (see also Mendes, 1952) and/or covered by dense commercial plantations (e.g., sugar
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cane). During this period, data about the Capivari marine fauna appeared only in review papers,
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as those by Rocha-Campos and Röser (1978) and Petri and Souza (1993). In both contributions,
208
the stratigraphic position and biocorrelation of the Capivari marine invertebrate fauna with those
209
of the southern parts of Brazil were tentatively constrained and discussed. In this context, a
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probable northward extension of those beds was mentioned by Rocha-Campos and Rösler (1978,
211
p. 5). Indeed, these authors reported the presence of similar fossil-bearing rocks near the city of
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Hortolândia, State of São Paulo, including the following bivalve genera: Phestia, Nuculopsis,
213
and Edmondia, plus undetermined pholadomyid shells.
214
In the mid-1990s, faculty of the Institute of Geosciences, University of São Paulo
215
(IGc/USP) carried out various field trips to the area originally mapped by Barbosa and Almeida
216
(1949b), “rediscovering” the fossil-bearing beds of Capivari (A.C. Rocha-Campos personal
217
communication, 1996). Consequently, new sampling efforts were carried-out, significantly
218
contributing to the find of new specimens. This enlarged substantially the original fossil
219
collection made by Barbosa and Almeida (1949a, b) and Mendes (1952). Unfortunately, at the
220
end of the 1990s, the original exposure of the Capivari outcrop was almost destroyed (Anelli et
221
al., 2000, p. 151) during the road works to enlarge the original path of the old, unpaved route
222
between the cities of Capivari and Salto (see Fig. 2). Despite that, renewed interest on those
223
fossils reappeared during the extensive study of the coeval late Paleozoic marine invertebrate
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faunas of the Itararé Group from southern Brazil (Simões et al., 2012; Neves et al., 2014b;
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Taboada et al., 2016). These studies better constrained the stratigraphic position, age and
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biocorrelation of the marine assemblages of the Taciba Formation, upper part of the Itararé
9
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Group, with other well-dated Gondwanan faunas (Simões et al., 2012; Neves et al., 2014b;
228
Taboada et al., 2016).
229
Based on the literature (e.g. Vesely and Assine, 2006) and personal communications from
230
various paleontologists and stratigraphers (A.C. Rocha-Campos, S. Petri), we relocated what was
231
left of the fossil-bearing beds in the Capivari-Salto road. At the same time, significant efforts
232
were made to gather the original material collected by previous authors, such as O. Barbosa,
233
F.F.M. Almeida, S. Petri, J.C. Mendes, and A.C. Rocha-Campos. Hence, in this contribution we
234
describe the fossil-bearing sedimentary succession of the Itararé Group near Capivari, and revise
235
the marine assemblage based on new material as well as the original fossils studied by previous
236
authors.
237 238
2. Material and methods
239
In total, 150 specimens of macroinvertebrates of the Capivari marine beds were examined, as
240
follows: 115 brachiopods, seven bivalves, three gastropods, and 25 crinoid fragments (Table 4).
241
All analyzed specimens (Tables 5-7) come from the classical outcrop located at ~6 km south of
242
the Capivari, on the old road between the Capivari and Salto towns (Figs. 2, 3).
243
In the laboratory, the fossil material was prepared according to standard paleontological
244
procedures (see Feldmann et al., 1989) using precision tools. Latex casts and plasticine molds
245
were also prepared for most of the specimens. Shells were coated with magnesium oxide
246
sublimate to enhance diagnostic morphological characters for photography (see Neves et al.,
247
2014a, b, and references therein). The ecological guilds (i.e., inferred invertebrate lifestyles)
248
were determined based on Aberhan and Kiessling (2015, p. 2).
249 250
Repositories and institutional abbreviations
10
251
All studied specimens are housed in the Institute de Geosciences, University of São Paulo, São
252
Paulo city, State of São Paulo. The following abbreviations are used in the text for institutional
253
acronyms: GP-1T, this prefix is reserved for type specimens belonging to the Invertebrate
254
Collection deposited in the Geosciences Institute, São Paulo University, Brazil; GP-1E, this
255
prefix is used for the invertebrate specimens that are housed in the non-type collection of that
256
institution; DGM, this prefix indicates that the specimens belong to the fossil collection of the
257
Geology and Mineralogy Division, Nacional Department of Mineral Production, State of Rio de
258
Janeiro, Brazil; IGG, this prefix is used for the material reposited in the Geographical and
259
Geological Institute of the State of São Paulo, now Geological Institute, Secretariat of the
260
Environment, São Paulo State, Brazil; CP.I, this prefix refers to the CENPALEO fossil
261
collection, Contestado University, Mafra, State of Santa Catarina, southern Brazil.
262 263
3. Systematic paleontology
264
The applied classification for Brachiopoda largely follows Waterhouse (2013) for Suborder
265
Linproductidina, Savage et al. (2002) for Order Rhynchonellida, Carter et al. (2006) for
266
Suborder Spiriferidina, Racheboeuf (2000) for Suborder Chonetidina. Taxonomic analysis of
267
Bivalvia follows Waterhouse, (2008, 2010), Bieler et al. (2010), Carter et al. (2011) for
268
suprageneric categories, and Dickins (1957; 1963), Morris et al. (1991), Newell and Boyd (1995)
269
and Neves et al. (2014b) at the generic and species level. Finally, the name of crinoid columnals
270
and pluricolumnals are preceded by the prefix ‘‘ø’’, as suggested by Le Menn (1987, 1988) and
271
Scheffler et al. (2015).
272 273
Class Strophomenata Williams et al., 1996
274
Order Productida Sarytcheva and Sokolskaja, 1959
275
Suborder Linoproductidina Waterhouse, 2013 11
276
Superfamily Proboscidelloidea Muir-Wood and Cooper, 1960
277
Family Auriculispinidae Waterhouse, 1986
278
Subfamily Lyoniinae Waterhouse, 2001
279
Tribe Lyoniini Waterhouse, 2001
280 281
Genus Lyonia Archbold, 1983
282 283
Type species: Linoproductus (Cancrinella) cancriniformis var. lyoni Prendergast 1943, from the
284
Lyons Group (lower Permian), Carnarvon Basin, Western Australia.
285 286
Remarks: A set of finely ribbed, spinose linoproductids such as Cancrinella Fredericks, 1928,
287
Auriculispina Waterhouse,1975, Costatumulus Waterhouse, 1983a, and Magniplicatina
288
Waterhouse, 1983b are externally closely similar to Lyona. Nevertheless, when compared to
289
Lyonia, Cancrinella possesses a more inflated shell, smaller size, crowded spines on the ears and
290
abundant spines on the dorsal valve (see discussion of this last character in Grigorjeva et al.,
291
1977; Waterhouse, 1982a; Shen et al., 2000; Li et al., 2012). Auriculispina is distinguished from
292
Lyonia by the presence of two or three rows of spines on the ears, stronger rugae on both valves,
293
more weakly swollen ventral spine ridges and a lack of dorsal spines (Waterhouse, 1975;
294
Archbold, 1983). Likewise, Costatumulus could be confused to Lyonia but the former has a more
295
inflated concave-convex profile, a lack of dorsal spines and possesses double rows of spines on
296
the ears (Waterhouse, 1983a, 1986). Magniplicatina also has ventral spines in one ou more hinge
297
rows and lacks dorsal spines but exhibits strongly developed concentric wrinkles or rugae as well
298
as coarser ventral spines on elongate spine ridges (Waterhouse, 1983b; Li et al., 2012), unlike
299
Lyonia. Closest genera to Lyonia such as Nambucalinus Waterhouse, 2001 and Bandoproductus
300
Jin and Sun, 1981, were discussed in a previously published paper (Taboada et al., 2016). 12
301 302
Lyonia rochacamposi Taboada et al., 2016
303
Figure 4.29–4.32
304 305
1969 Cancrinella sp.; Rocha-Campos, p. 105, pl. X, figs. 1-5, 8.
306
1969 Linoproductus sp.; Rocha-Campos, p. 106, pl. X, figs. 6-7.
307
2016 Lyonia rochacamposi; Taboada et al., p. 443, figs. A-O.
308 309
Holotype: CP.I 769I, CENPALEO (Universidade do Contestado), from the Taciba Formation,
310
Itararé Group, southern Paraná Basin, Brazil.
311 312
Occurrence: Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
313
Basin, Brazil. Latest Asselian–earliest Sakmarian.
314 315
Description: Lyonia with transverse outline and moderately convex ventral valve. Ventral valve
316
maximum width close to hinge line, up to 30 mm wide, 24 mm long, and width/length ratio 1.25.
317
Ventral valve with large flattened ears (partially broken) and ornamentation of fine radial
318
costellae (8-9 per 5 mm on the venter); weak rugae on venter slightly stronger on ears. Costellae
319
number increasing by splitting into two ribs anteriorly from slightly swollen minute spine bases
320
in roughly quincuncial arrangement. A single row of spines, coarser than those on the venter
321
appears to be on hinge line, although this character is not well preserved.
322 323
Materials: Two fragmentary specimens, one external mold of dorsal valve, GP/1E 4350a; one
324
ventral valve, GP/1E 4350b.
325 13
326
Measurements: See Table 5 for Lyonia rochacamposi shell measurements.
327 328
Remarks: The studied specimens are scarce and limited to fragmentary remains of ventral
329
valves but exhibit comparable size, moderate convex profile, transverse outline, large flattened
330
ears, single row of hinge spines, distinct fine costellae and ventral spines quincunxial
331
arrangement that suggest its assignment to Lyonia rochacamposi Taboada et al., 2016. This
332
species was recently described from the post-glacial section of the Taciba Formation in the
333
Brazilian Santa Catarina State. The Capivari’s L. rochacamposi specimens, as well as those
334
from the southern Taciba Formation, are distinguished from L. lyoni from Lyons Group of
335
Western Australia mainly by their more transverse outline and finer costellation.
336
Cancrinelloides monticulus Waterhouse, 1982b, from the Asselian Pucket Group of south
337
Thailand was considered moderately close to Lyonia lyoni, but the Australian species is smaller,
338
with more emphasized concentric ornament, finer radial ornament, and slightly shorter spine
339
bases (Waterhouse, 1982b). Likewise, Lyonia rochacamposi is distinguished from the Thailand
340
species by its more transverse outline, finer costellation and presence of dorsal spines anteriorly.
341
Cancrinelloides monticulus was assigned latter to Bandoproductus (although with subgeneric
342
hierarchy) by Waterhouse (1982b) and shortly after to the same genus by Archbold (1983).
343 344
Order Rhynchonellida Kuhn, 1949
345
Superfamily Rhynchoporoidea Muir-Wood, 1955
346
Family Rhynchoporidae Muir-Wood, 1955
347
Subfamily Rhynchophoriinae Muir-Wood, 1955
348 349
Genus Rhynchopora King, 1865
350 14
351
Type species: Terebratula geinitziana de Verneuil, 1845, from the upper Permian (Kazanian) of
352
Arkhangelsk Region, Russia.
353 354
Rhynchopora grossopunctata Mendes, 1952
355
Figure 4.1–4.15
356 357
1952 Rhynchopora grossopunctata; Mendes, p. 12, figs. 4-7.
358
1966 Rhynchopora grossopunctata; Mendes, Mezzalira, pl. 2, figs. 3-4 (copy Mendes, 1952,
359
figs. 4-5).
360
1967, Rhynchopora grossopunctata; Mendes, Rocha-Campos, pl. 31, figs. 1-3 (copy Mendes,
361
1952 figs. 4-5, 7).
362
1989 Rhynchopora grossopunctata; Mendes, Mezzalira, pl. 2, figs. 3-4 (copy Mendes, 1952, figs.
363
4-5).
364 365
Holotype: Mendes (1952) did not choose a holotype but illustrated the syntype series of
366
Rhynchopora grossopunctata with three specimens labeled DGM 4189-4190 and IGG 518-I,
367
respectively. The IGG repository (Instituto Geográfico e Geológico do Estado de São Paulo) is
368
no longer available, while the specimens under the DGM acronym (Divisão de Geologia e
369
Mineralogia, Departamento Nacional da Produção Mineral, Estado de Rio de Janeiro) were
370
transferred to the fossil collection of the Instituto de Geociências da Universidade de São Paulo,
371
although they are currently lost. An external mold of conjugated valves, labeled GP/1E-4342,
372
housed in the Instituto de Geociências da Universidade de São Paulo, is here selected as neotype
373
of R. grossopunctata Mendes.
374
15
375
Diagnosis: Large, subpentagonal in outline with maximum width mid-length. Five costae on
376
sulcus and fold, eight to nine on lateral flanks. Interspaces anteriorly develop coarse projections
377
extending as short tongues. Five punctae per mm2. Dental plates diverge 30º, up to one fourth of
378
valve length; dorsal interior with a thin median septum one third of valve length.
379 380
Occurrence: Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
381
Basin, Brazil. Latest Asselian–earliest Sakmarian.
382 383
Description: Large-sized Rhynchopora, subpentagonal in outline with maximum width up to 18
384
mm at mid-length and length up 16 mm, with an average W/L ratio of 1.15. Ventral valve gently
385
convex with maximum convexity at umbo and dorsal valve more inflated with slightly raised
386
fold and maximum convexity at two thirds of valve length. Umbonal angle of 95º-100º; cardinal
387
extremities rounded. Sulcus very shallow, barely noticeable anteriorly from first third of valve
388
length and widening anteriorly (~40º) up to one fourth of maximum width. Five costae on sulcus
389
and fold, and 8-9 on lateral flanks, with finer costae developed postero-laterally. Costae wider at
390
commissure. Interspaces anteriorly develop long coarse projections extending as short tongues.
391
Five punctae per mm2. Dental plates diverge 30º, up to one fourth of valve length; dorsal interior
392
with a distinct thin median septum extending one third of valve length. Others features unknown.
393 394
Materials: One external mold of conjugated valves is the neotype, GP/1E-4342. Twenty
395
topotypic specimens. Two ventral valve external molds, GP/1E-443a, 443b; two ventral valve
396
internal molds GP/1E-2507, 2545; three dorsal valve external molds, GP/1E-451, 1915, 4405;
397
three dorsal valve internal molds, GP/1E-1912, 2492, 4325. Other fragmentary specimens,
398
GP/1E-445b, 446-447, 460, 1913, 2502, 2524, 2536, 4339.
399 16
400
Measurements: See Table 5 for Rhynchopora grossopunctata shell measurements.
401 402
Remarks: Although not preserved in the material here described, a spondylium is clearly present
403
in the syntype series of R. grossopunctata (Mendes, 1952). Species of Rhynchopora reported
404
from South America include Kozlowski (1914) R. nikitini (not Tschernyschew, see Cooper and
405
Grant, 1976) from the Copacabana Formation (Early Permian) of Bolivia (later synonymysed
406
under R. aff. illinoisensis [Worthen] by Chronic in Newell et al., 1953), and R. amotapensis
407
Chronic (in Newell et al., 1953), the latter two from the Tarma Group (Late Carboniferous) of
408
Peru. The last species shares a slightly transverse subpentagonal outline with R. grossopunctata,
409
as well as shallow sulcus and flatten fold with a similar number of costae, but the Brazilian
410
species is larger and exhibits more costae on the lateral flanks. R. aff. illinoisensis is smaller and
411
possesses more costae on sulcus, fold and lateral flanks when compared to R. grossopunctata.
412
Another South American species is Rhynchopora sp. from the Río del Peñón Formation
413
(Moscovian) of central-western Argentina (Cisterna and Simanauskas, 2000). It differs from R.
414
grossopunctata by having a smaller size, rounded subtriangular outline, deeper sulcus, fewer
415
costae on flanks and longer dorsal median septum. Rhynchopora australassica Archbold, 1995
416
(see also Archbold and Hogeboom, 2000), from the Lyons Group (Early Permian) of Western
417
Australia, has a larger size, subtriangular outline, fewer costae on flanks, deeper sulcus and a
418
more raised fold when compared to R. grossopunctata. Rhynchopora culta Waterhouse, 1982b,
419
from south Thailand and western Peninsular Malaysia (Shi and Waterhouse, 1991) is comparable
420
to R. grossopunctata, although the former has a subtrigonal shell shape, reduced number of
421
costae on flanks and almost double the number of punctae per millimeter.
422 423
Order Spiriferida Waagen, 1883
424
Superfamily Ambocoeloioidea George, 1931 17
425
Family Ambocoeliidae George, 1931
426
Subfamily Ambocoeliinae George, 1931
427
Genus Biconvexiella Waterhouse, 1983a
428 429
Type species: Attenuatella convexa Armstrong, 1968 from the Tiverton Formation, (Early
430
Permian), Bowen Basin, Queensland, Australia.
431 432
Remarks: Biconvenxiella shares with Attenuatella Stehli, 1954 and Crurithyris George, 1931,
433
comparable shell shape and a similar external spinose microornamentation. Biconvexiella
434
possesses a subcircular shell outline, slightly incurved umbo, slightly concave to gently convex
435
dorsal valve and two pairs of adductor scars, the posterior pair scars being smaller than anterior
436
pair (Shi and Waterhouse, 1996; He et al., 2007). Attenuatella is distinguished from
437
Biconvenxiella by the presence in the former of an attenuated outline, a stronger convex ventral
438
valve and strongly incurved umbo, as well as a different arrangement and proportions of muscle
439
scars (Stehli, 1954; Waterhouse, 1964; Landis and Waterhouse, 1966; Shi and Waterhouse, 1996:
440
He et al., 2007, 2012). Crurithyris is characterized by a subrounded to transversely ovate shell
441
outline, a moderately to strongly incurved ventral umbo, a pair of centrally depressed ventral
442
adductor scars bisected by a thin ridge (may be absent), a pair of laterally depressed diductor
443
scars, which are separated from their adjacent adductor scars by ridges, and a pair of dorsal
444
adductor scars bisected by a median ridge (George, 1931; Stehli, 1954; He et al., 2007, 2012).
445 446
Biconvexiella saopauloensis n. sp.
447
Figure 4.16–4.25
448 449
1952 Crurithyris aff. planoconvexa (Shumard); Mendes, p. 10-11, pl. I, figs. 1-3. 18
450 451 452 453
1966 Crurithyris aff. planoconvexa (Shumard); Mezzalira, pl. 2, figs. 1-2 (copy Mendes, 1952 pl. I, figs. 1-2). 1967 Crurithyris aff. planoconvexa (Shumard); Rocha-Campos, pl. XXXI, figs. 4-5 (copy Mendes, 1952, pl. I, figs. 2-3).
454
1969 Attenuatela paulistana; Rocha-Campos, p. 108-112, text-fig. 12, pl. XI, figs. 1-14.
455
1989 Crurithyris aff. planoconvexa (Shumard); Mezzalira, pl. II, figs. 1-2 (copy Mendes,
456 457
1952 pl. I, figs. 1-2). 2016 Biconvexiella spp.; Taboada et al., pl. 4, fig. J.
458 459
Holotype: GP/1E 4339; Paratypes: GP/1E 450, 453, 2495, 2513, 2520, 2534, 4322, 4327, 4328,
460
4356c, all from the Taciba Formation, Paraná Basin, State of São Paulo, southeastern Brazil.
461 462
Diagnosis: Average-sized Biconvexiella of biconvex profile with slightly convex dorsal valve
463
and strongly convex ventral valve, and subcircular outline. Ventral valve subequidimensional;
464
dorsal valve variably transverse; hinge line shorter than maximum width. Ventral median sinus
465
shallow and narrow. Dorsal valve convex with a narrow slightly raised fold.
466 467
Occurrence: Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
468
Basin, Brazil. Latest Asselian–earliest Sakmarian.
469 470
Description: Average-sized Biconvexiella of biconvex profile and subcircular outline. Ventral
471
valve subequidimensional; dorsal valve variably transverse; hinge line shorter than maximum
472
width; which is located at valve midlength. Maximum width up to 14 mm, maximum length up
473
to 11.5 mm with a dorsal W/L ratio varying between 1.23-1.43 and W/L average of 1.33. Ventral
474
valve strongly convex with its stronger convexity in the umbonal region. Ventral median sinus 19
475
shallow and narrow. Dorsal valve slightly convex with a narrow fold not always developed.
476
Ornamentation of fine concentric growth lines sometimes lamellose and subconcentric rows of
477
minute subcircular spine bases (9-10/mm at valve mid lenght). Interior of ventral valve with long
478
(3/4 ventral valve length), narrow median adductor ridge bounded by two narrow elongate
479
depressed adductor scars and slightly rised semielliptical diductor scars. Interior of dorsal valve
480
(holotype) with small subelliptical (0.3 mm width, 1 mm length) depressed posterior adductor
481
scars separated by a short thin median ridge. Anterior adductor scars barely impressed, elongate
482
(2 mm length) tongue-shaped (1 mm of maximum width anteriorly) with inner side curved.
483
Crural plates short, divergent (30º-50º); dental sockets deep; cardinal process tuberculate.
484 485
Etymology: From the State of São Paulo, southeastern Brazil.
486 487
Materials: Eighty-six specimens. Three external molds of conjugated valves: GP/1E 2495, 2513,
488
4328; two exterior of dorsal valves: GP/1E 2534, 4327; five dorsal valve external molds: GP/1E
489
448, 449, 2413, 4323a, 4354d1; fifteen ventral valve external molds; GP/1E 2503, 2504, 2522,
490
2526, 2544, 2552, 4318a, 4322, 4323b, 4324, 4326, 4332, 4341, 4354d2; eleven dorsal valve
491
internal molds: GP/1E 450, 453, 2493, 2498, 2536-2538, 2543, 4339, 4351; twelve ventral valve
492
internal molds: GP/1E 1910, 2489, 2520, 2525, 2527, 2528, 2551, 4354b, 4355b, 4356a, 4356b,
493
4338; other fragmentary material: GP/1E 1906, 1907, 1911, 1914, 2496, 2497, 2499, 2500, 2505,
494
2506, 2508, 2510, 2514-2517, 2521, 2524, 2529, 2530, 2532, 2533, 2535, 2539, 2541, 2546-
495
2550, 2553, 2555, 4320, 4330, 4331, 4334, 4340, 4357.
496 497
Measurements: See Table 5 for Biconvexiella saopauloensis n. sp. shell measurements.
498
20
499
Remarks: Ambocoelids from the Taciba Formation were first mentioned by Barbosa and
500
Almeida (1949a, b), later briefly described as Crurithyris aff. planoconvexa (Shumard) by
501
Mendes (1952) but interpreted as belonging to a new species (named paulistana) of
502
Attenuatella Stehli, 1954, by Rocha-Campos (1969). Attenuatella paulistana was never
503
published, failing to satisfy article 13 of the ICZN, therefore being a nomen nudum. The
504
Capivari’s ambocoelids exhibit shell shape, slightly incurved umbo, strong ventral median
505
ridge supporting muscle scars, dorsal thin median ridge separating posterior adductors scars and
506
larger anterior adductor scars, that collectively support assingnment to Biconvexiella, as was
507
recently indicated (Taboada et al., 2016).
508
Biconvexiella saopauloensis n. sp. is comparable to Attenuatella convexa Armstrong,
509
1968, type species of Biconvexiella Waterhouse, 1983a, which occurs in the Tiverton
510
Formation (Sakmarian), Bowen Basin, Queensland, and also present in the Farley Formation,
511
Sydney Basin, New South Wales, Australia (Armstrong and Telford, 1970). It shares with B.
512
saopauloensis n. sp. a biconvex profile, ventral sinus and dorsal fold, as well as comparable
513
internal muscle scars. Nevertheless, B. saopauloensis n. sp. has a more transverse outline, less
514
pronounced ventral valve sinus and shallower dorsal valve fold lacking lateral bounding ridges.
515
B. saopauloensis n. sp. is close to B. roxoi (Oliveira, 1936) (see also Taboada et al., 2016) from
516
the upper part of the Taciba Formation, Paraná Basin, southern Brazil, although the Capivari’
517
species is more transverse with a convex dorsal valve, a stronger and longer ventral median
518
septum and more divergent crural plates. Ogilviecoelia inflata Shi and Waterhouse, 1996, from
519
the Early Permian Jungle Creek Formation, northern Yukon Territory, Canada, resembles B.
520
saopaulensis n. sp. in shell outline and dorsal valve profile, although the Canadian material
521
possesses a micro-ornamentation of fine elongate grooves instead spinose ornament, and a large,
522
depressed and well differentiated ventral muscle field and undifferentiated dorsal adductors,
523
unlike the Brazilian species. 21
524 525
Order Chonetida Nalivkin, 1979
526
Suborder Chonetidina Muir-Wood, 1955
527
Superfamily Chonetoidea Bronn, 1862
528
Family Rugosochonetidae Muir-Wood, 1962
529
Subfamily Quinquenellinae Archbold, 1981
530 531
Genus Quinquenella Waterhouse, 1975
532 533
Type species: Quinquenella glabra Waterhouse, 1975 from the Nambdo Member of the Senja
534
Formation (late Permian), east Jumbla city, north-west Nepal.
535 536
Remarks: Both Tornquistia Paeckelmann, 1930 and Quinquenella are smooth externally and
537
have similar dorsal interiors. Nevertheless, the former is smaller, has a hump-shaped strong
538
convave-convex profile and stout accessory septa in the dorsal valve, unlike the low profile and
539
weaker accessory septa of Quinquenella.
540 541
Quinquenella rionegrensis (Oliveira, 1930)
542
Figure 4.26–4.28
543 544
1930 Chonetes rionegrensis; Oliveira, p. 19, unnumbered figure.
545
2016 Quinquenella rionegrensis (Oliveira, 1930); Taboada et al., 2016, p. 449-450, pl. III,
546
figs. D-L.
547
22
548
Holotype: The holotype of Quinquenella rionegrensis (Oliveira, 1930) is lost (see Taboada et
549
al., 2016, p. 449, 450). Lectotype GP/1E 4339b is from the Taciba Formation, Itararé Group,
550
Paraná Basin, Teixeira Soares region, southern Brazil.
551 552
Occurrence: Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
553
Basin, Brazil. Latest Asselian–earliest Sakmarian.
554 555
Materials: Seven specimens. Two interior molds of dorsal valves, GP/1E-4354c, GP/1E 4358;
556
one exterior mold of dorsal valve, GP/1E 4355a. Other fragmentary material, GP/1E 449, 456,
557
1909, 2523.
558 559
Measurements: See Table 5 for Quinquenella rionegrensis shell measurements.
560 561
Remarks: Available dorsal valves exhibit moderate concavity, subelliptical transverse outline
562
and maximum width at hinge line. Maximum width up 9 mm; maximum length up 5.5 mm with
563
an W/L ratio varying between 1.55-1.76 and W/L average of 1.65. Interior with lines of pustules
564
radially aligned, short lateral septa, thin long accessory septa, ill-defined median septum fused
565
posteriorly surrounding a small, circular shallow alveolus anterior to a small bifid cardinal
566
process. Despite scarcity of material, internal characters and dimensions of dorsal valve fit finely
567
with those observed in Quinquenella rionegrensis (Oliveira, 1930) (see also Taboada et al., 2016)
568
from the upper part of the Taciba Formation, Paraná Basin, southern Brazil. Finally, the
569
Brazilian species was compared to Quinquenella species from Western Australia (see Archbold
570
1981) by Taboada et al. (2016).
571 572
Class Bivalvia Linnaeus, 1758 23
573
Subclass Protobranchia Pelseneer, 1889
574
Order Nuculanida Carter, Campbell and Campbell, 2000
575
Superfamily Nuculanoidea Adams and Adams, 1858 (Gray, 1854)
576
Family Polidevciidae Kumpera, Prantl and Ruzicka, 1960
577 578
Genus Phestia Chernyshev, 1951
579 580
Type species: Leda inflatiformis Chernyshev, 1939 from the Carboniferous of
581
Russia, by original designation.
582 583
Remarks: González (2010) proposed the genus Waterhouseus to include the species Phestia
584
tepuelensis, but the generic diagnosis is not precise. Besides, the differences of Phestia and
585
other closely related genera, such as Polidevcia are not clear in the discussion. Thus, we decide
586
to keep the original name for the Patagonian species (Argentina). A revision of these genera
587
may be necessary but exceeds the scope of this contribution and is an objective of a future
588
research.
589 590
Phestia tepuelensis Gonzalez, 1969
591
Figure 5.1–5.4
592 593
Holotype: MLP 11021 from the Sierra de Languiñeo, Tepuel-Genoa Basin, Chubut, Argentina
594
(Gonzalez, 1969; Pagani, 2004).
595 596
Occurrence: Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
597
Basin, Brazil. Latest Asselian–earliest Sakmarian. 24
598 599
Description: Shell very inequilateral, tapering posteriorly with a developed rostrum. Umbones
600
slightly opisthogyrous, small, very depressed, positioned at anterior fourth of shell length. The
601
greatest height of the shell occurs at umbones. Anterior margin rounded forming an obtuse angle
602
with a convex ventral margin; posterior-dorsal margin slightly concave. Surface ornamentation
603
of thick commarginal costae, which are crossed by radial costae; with a pronounced
604
discontinuity in the coarseness of the commarginal ribs from the anterior lobe to the lateral flank.
605
Escutcheon, hinge features and muscle scars not observed.
606 607
Materials: One internal mold of right valve (GP/1E 455) and one external mold of left valve
608
(GP/1E 452).
609 610
Measurements: See Table 6 for Phestia tepuelensis shell measurements.
611 612
Remarks: Phestia tepuelensis was firstly described for the Tepuel-Genoa Basin, Argentina
613
(González, 1969), and revised by Pagani (2004). This species shows strong concentric costae
614
crossed by fine radial ribs, a feature that distinguishes it from seven Argentinean species. Three
615
of them come from Northwestern Argentina: Phestia sp. from San Juan Province (Taboada,
616
1989); Phestia sp. from Río Blanco Basin (González, 1994), both Carboniferous, and Phestia aff.
617
P. bellistriata (Stevens) from Permian deposits of the Tupe Formation (Sterren, 2004). The other
618
four species come from the Tepuel-Genoa Basin, in Patagonia, described by Pagani (2004): P.
619
regularis, P. sabattiniae, Phestia sp. II and Phestia sp. III. The Capivari specimens differ from
620
Phestia aff. P. sabattiniae from the Itararé Group (Taciba Formation, see Neves et al., 2014b),
621
by having an opisthogyrous umbones and a concave posterior-dorsal margin, while in the second
622
species the umbones are orthogyrous and the posterior-dorsal margin is straight. Yet, P. 25
623
tepuelensis presents radial ribs crossing the commarginal costae, a feature absent in the Brazilian
624
specimens. Phestia tepuelensis also differs from two Australian species Phestia darwini
625
(=Nuculana darwini) (Dickins, 1957) and Phestia lyonensis Dickins, 1956. The Brazilian species
626
have narrower umbones, which are very pronounced in P. darwini and the surface of shell is
627
ornamented by commarginal ribs, while in P. lyonensis they can form a shallow V-shaped
628
ornamentation (see Dickins, 1963, p. 37).
629 630
Order Pectinida Gray, 1854
631
Superfamily Chaenocardioidea Miller, 1889
632
Family Streblochondriidae Newell, 1938
633
Subfamily Saturnellinae Astafieva, 1994
634
Genus Streblopteria McCoy, 1851
635 636
Type species: Meleagrina laevigata McCoy, 1844 from Carboniferous of Ireland, by posterior
637
designation of Meek and Worthen (1866, p. 333).
638 639
Remarks: González (2006) described the species Streblopteria montgomeryi for the Mojón de
640
Hierro Formation in Patagonia, Argentina; at the same time Pagani (2006) described the species
641
Streblopteria lagunensis from the same localities. Both species are synonyms and Gonzalez
642
(2006) and Pagani (2006) have the same date of publication. In this way, the rule of
643
“Determination by the First Reviser” (ICZN Article 24.2) is applied, and we use Streblopteria
644
lagunensis in the sense of Pagani (2006). Later, Waterhouse (2010 p. 105) designated the
645
Argentinian specimens of Streblopteria montgomeryi Gonzalez (2006) under a new genus
646
Montorbicula, based on the absence of a resilifer and by the presence of a platyvincular hinge.
647
However, the generic diagnosis is imprecise. Besides, the differences with Streblopteria and 26
648
other related new genera proposed by Waterhouse (2010) are unclear in the discussion. Thus we
649
decided to keep the original generic assignation for the Patagonian species.
650 651
Streblopteria aff. S. lagunensis Pagani, 2006
652
Figure 5.5–5.7
653 654
1978 Streblopteria sp. nom. nud.; in Rocha-Campos and Rösler (1978, p. 5).
655
1988 Streblopteria sp. nom. nud.; in Simões et al. (1998, p. 445).
656 657
Occurrence: Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
658
Basin, Brazil. Latest Asselian–earliest Sakmarian.
659 660
Description: Internal mold of right valve, suborbicular, slightly longer than higher, inflated,
661
acline. Umbones slightly project above the cardinal margin. Anterior auricle separated from the
662
shell by a moderately deep byssal notch. Posterior auricle small, subtriangular, with its posterior
663
margin forming an obtuse angle with the postero-dorsal margin. Cardinal margin straight;
664
anterior margin seems to be more expanded than the posterior one; ventral, posterior and,
665
possibly, anterior (fragmented) margins rounded. External surface of shell ornamented by faint
666
and fine growth lines, at least in the ventral margin. Inner cardinal features and muscle scars not
667
observed.
668 669
Material: One internal mold of right valve, GP/1E 457.
670 671
Measurements: See Table 6 for Streblopteria aff. S. lagunensis shell measurements.
672 27
673
Remarks: Streblopteria aff. S. lagunensis of the Taciba Formation closely resembles
674
Streblopteria lagunensis (=Streblopteria montgomeryi) recorded in lower Permian (Asselian)
675
beds of Patagonia, Argentina (Pagani, 2006, p. 468, fig. 3 M-S; González, 2006, p. 140, fig. 8).
676
The last author suggested that the Argentinean species have smooth shells and inconspicuous
677
commarginal ornamentation, which can also be verified in the Capivari specimen. However, the
678
Brazilian specimen seems to be ornamented by very closely spaced fine growth lines (Figure
679
5.7). Thus, we kept the Capivari species as Streblopteria aff. S. lagunensis.
680 681
Superfamily Heteropectinoidea Beurlen, 1954
682
Family Limipectinidae Newell and Boyd, 1990
683
Subfamily Limipectininae Newell and Boyd, 1990
684 685
Genus Limipecten Girty, 1904
686 687
Type species: Limipecten texanus Girty, 1904 from the Carboniferous of Mexico, by
688
subsequent designation of Newell (1938, p. 68).
689 690
Remarks: Mendes (1952) assigned the pectinid of the Taciba Formation, Capivari County, to
691
Aviculopecten capivariensis. He pointed out that these specimens had just one order of costae
692
(Mendes, 1952, p. 13). However, Rocha-Campos (1969) referred them to Limipecten
693
capivariensis, since they show surface ornament formed by two or three order costae (Rocha-
694
Campos, 1969, p. 48). He referred this bivalve to Limipecten capivariensis nom. nud. We agree
695
with this interpretation and here this species is formally described for the first time.
696 697
Limipecten capivariensis (Mendes), 1952 28
698
Figure 5.8–5.10
699 700
1952 Aviculopecten capivariensis; Mendes (p. 12, plate I, figs. 8 e 9).
701
1967 Aviculopecten capivariensis; Rocha-Campos (plate XXIX, fig. 14).
702
1969 Limipecten capivariensis nom. nud.; in Rocha-Campos (p. 47, plate II, F).
703
1988 Limipecten capivariensis nom. nud.; in Simões et al. (1988, p. 445).
704 705
Holotype: The holotype (DGM 4121) of Limipecten capivariensis (Mendes), 1952 is lost.
706
Lectotype GP/1E 2494 from the Taciba Formation, Itararé Group, Paraná Basin, Capivari
707
county, State of São Paulo, southeastern Brazil.
708 709
Occurrence: Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
710
Basin, Brazil. Latest Asselian–earliest Sakmarian.
711 712
Diagnosis: Convex left valve. Valve ornamented by, at least, ten coarse first order radial ribs,
713
intercalated by second order costae; interspaces among ribs wider than in the right valve. Surface
714
of shell crossed by concentric lamellae, forming scalloped lines. Right valve moderately convex.
715
Surface of shell ornamented by twenty-two first order radial ribs, intercalated by second and
716
third order costae; ribs finer and numerous than in the left valve; costae are crossed by concentric
717
regular lamellae.
718 719
Description: Shell orbicular, inequivalved, with length slightly greater than width. Convex left
720
valve; umbo and auricles not preserved (fragmented). Surface of valve ornamented by, at least,
721
ten coarse first order radial ribs; second order ribs increase by intercalation; interspaces among
722
ribs are wider in the left valve. Surface of shell crossed by concentric lamellae, forming 29
723
scalloped lines. Right valve moderately convex, mainly in the ventral margin. Umbo poorly
724
preserved. Anterior ear sharply defined, separated from the body shell by a well-marked byssal
725
sinus; seven first order ribs crossed by concentric regularly spaced lamellae. The posterior ear is
726
poorly preserved. Surface of shell is ornamented by twenty-two first order radial ribs,
727
intercalated by second and third order costae; ribs are much finer and numerous than in the left
728
valve; surface of shell is crossed by concentric regular lamellae, which are also finer and fainter
729
than in the left valve. Internal features of hinge plate not observed.
730 731
Materials: Two external molds, one fragment of left valve, with well preserved surface
732
ornamentation (GP/1T-116), and one almost complete right valve (GP/1E 2494).
733 734
Measurements: See Table 6 for Limipecten capivariensis shell measurements.
735 736
Remarks: Limipecten capivariensis differs from the type species L. texanus, from Texas, USA,
737
by showing much wider interspaces between ribs in the left valve. Additionally, the right valve
738
of the Brazilian shell is much taller than the North American one. The Capivari shells also differ
739
from Limipecten sp. of the Ciénega Larga de Tontal Formation, San Juan, Argentina (Lech and
740
Milana, 2006), by the absence of protuberances in the meeting of ribs and growth lines, which is
741
a remarkable feature of these specimens. Limipecten capivariensis also differs from another
742
Argentinean species: Limipecten? sp. described by González (1972) in the Las Salinas Formation,
743
Chubut, later revised by Pagani (2006) as Limipecten herrerai. In the Brazilian shells, the radial
744
ribs are well spaced than in L. herrerai. Yet, the commarginal lamellae are more well spaced
745
than in the Argentinian specimens.
746 747
Infrasubcohort Cardiidia Férussac, 1822 30
748
Superfamily Grammysioidea Miller, 1877
749
Family Sanguinolitidae Miller, 1877
750
Subfamily Sanguinolitinae Miller, 1877
751
Genus Praeundulomya Dickins, 1957
752 753
Type-species. Praeundulomya concentrica Dickins, 1957 from the Carnarvon Basin, lower
754
Permian, Western Australia.
755 756
Praeundulomya cf. subelongata Dickins, 1963
757
Figure 5.11–5.12
758 759
Occurrence. Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
760
Basin, Brazil. Latest Asselian–earliest Sakmarian.
761 762
Description. Internal mold strongly inequilateral, subequivalve, very elongate posteriorly.
763
Umbones subterminal; beaks slightly prosogyrous. Very narrow sinus extending from the
764
umbones to ventral margin; well-developed postumbonal ridge extends to posteroventral
765
extremity. Anterodorsal margin very short; anterior margin rounded; ventral margin slightly
766
convex; posterior extremity rounded and defined by the respiratory margin. Lunule and
767
escutcheon were not observed. Surface ornament of very coarse, regularly spaced
768
commarginal rugae. Hinge edentulous. Muscle scars not observed.
769 770
Material. One internal mold of conjugated valves (GP/1E-5242).
771 772
Measurements: See Table 6 for Praeundulomya cf. subelongata shell measurements. 31
773 774
Remarks: Praeundulomya subelongata was first described by Dickins (1963), in the Fossil Cliff
775
Formation, Western Australia. Recently, this species was reported as Praeundulomya cf.
776
subelongata, in the Taciba Formation, Itararé Group (Neves et al., 2014b). The Capivari
777
specimen closely resembles that of the Taciba Formation. For this reason, we assign the Capivari
778
material to Praeundulomya cf. subelongata. This species differs from Praeundulomya moreli
779
Gonzalez, 2006 from Early Permian beds of Tepuel-Genoa Basin, Argentina. The Brazilian shell
780
is very elongated posteriorly, with the postero-ventral angle defined by a well-marked
781
postumbonal carina, while the shell of P. moreli shells is more subquadrate in outline, with a
782
truncate dorsal-posterior margin, and the postero-ventral angle forming a gentle carina.
783 784
Class Gastropoda Cuvier, 1797
785
Order Vertigastropoda Salvini-Plawen, 1980
786
Superfamily Eotomarioidea Wenz, 1938
787
Family Eotomarridae Wenz, 1938
788
Subfamily Neilsoniidae Knight, 1956
789 790
Genus Peruvispira Chronic, 1949
791 792
Type species: Peruvispira delicata Chronic, 1949 from the Copacabana Group (lower Permian),
793
Peru.
794 795
Remarks: Peruvispira is a genus that close resembles Ptycomphalina Fischer 1885 (see Dickins,
796
1963, p. 126), and Pleurocintosa Fletcher (1958). Indeed, according to Dickins (1961),
797
Peruvispira is similar to Ptycomphalina but ‘characterized by the possession of a distinct 32
798
revolving concave area about as wide as and below the slit-band’. On the other hand, the
799
relationship of Peruvispira and other Late Paleozoic gastropods, such as Pleurocintosa, was
800
recently discussed by Taboada et al. (2015). We agree with them that Pleurocintosa cannot be
801
regarded as a valid genus, since it was erected based on inconspicuous (e.g., weak spiral filae)
802
and highly variable characters (e.g., presence of a third and peribasal carina bounding the
803
alveozone) found in species of Peruvispira (see Taboada et al., 2015, p. 219 for details).
804 805
Peruvispira brasilensis n. sp.
806
Figure 6.1–6.2
807 808
1966 Peruvispira delicata nom. nud.; in Rocha-Campos, p. 10, fig. 3 a, b.
809
1970 Peruvispira delicata nom. nud.; Rocha-Campos, p. 607, fig. 3.
810 811
Holotype: GP/1E-4319 from the Taciba Formation, State of São Paulo, southeastern Brazil.
812 813
Diagnosis: Pleurotomarid shell with 4-5 very concave whorls; deep and strongly concave
814
selenizone, ornamented by 8-9 lunulae per mm, bounded by two prominent carinae. Shell
815
ornamentation of thin growth lines (7-8 per mm).
816 817
Occurrence: Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
818
Basin, Brazil. Latest Asselian–earliest Sakmarian.
819 820
Description: Small and turbiniform pleurotomarid shell, moderately thick; apex and upper whorl
821
slightly convex; shell with 4-5 very concave whorls; selenizone deep and strongly concave,
822
moderately wide, ornamented by 9-10 concave lunulae per mm, bounded by two prominent 33
823
carinae; shell ornamented by 7-8 thin growth lines per mm. Lower face continuously rounded
824
and convex, not flattened; sutures well impressed; inner lip poorly preserved; outer lip well
825
developed, convex and rounded, becoming almost perpendicular to the last whorl to form a sinus.
826
Shell body thicker near aperture.
827 828
Etymology: From Brazil.
829 830
Materials: Three external molds, GP/1E-4319 and 4348A, B.
831 832
Measurements: See Table 6 for Peruvispira brasiliensis n. sp. shell measurements.
833 834
Remarks: Peruvispira brasilensis n. sp. differs from the type species P. delicata Chronic, 1949,
835
from the Carboniferous of Peru, by the presence of very concave whorls, limited by a deep and
836
strongly concave selenizone. The Brazilian species closely resemble Peruvispira cf. umariensis
837
(Dickins, 1963, p. 126), from Western Australia, however they differ in the density of the growth
838
lines. Peruvispira brasilensis n. sp. is ornamented by 7-8 growth lines per mm, while P. cf.
839
umariensis is nearly 4 per mm. Additionally, the Brazilian species shows higher and more
840
concave whorls. Peruvispira brasilensis n. sp. can be distinguished from P. sueroi from the Late
841
Paleozoic beds of Chubut, Argentina (Sabatini and Norait, 1969) by having a very concave
842
whorl area, in which the spires are gradually higher toward the anterior region. In P. sueroi, the
843
whorl area is slightly convex and the basal whorl is much higher than the other ones. Peruvispira
844
australis is another comparable Argentinean species from San Juan, Mendoza and Chubut
845
(Sabatini and Noirat, 1969, p 104). P. australis differs from P. brasilensis n. sp. by the same
846
characters of P. sueroi, for being three times wider and having more widely spaced growth lines
847
(2-3 per mm) as well. 34
848 849
Class Crinoidea Miller, 1821
850
Subclass and order uncertain
851
Group Pentameri Moore, Jeffords and Miller, 1968
852
Family øPentacauliscidae Moore, Jeffords and Miller, 1968
853 854
Genus øPentaridica Moore, Jeffords and Miller, 1968
855 856
Type species: øPentaridica rothi Moore, Jeffords and Miller, 1968.
857 858
øPentaridica sp.
859
Figure 6.4-6.6
860 861
Occurrence. Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
862
Basin, Brazil. Latest Asselian–earliest Sakmarian.
863 864
Materials: Four individual discs (articular facet) GP/1E-4321; 4349 B and C; 4350C, and one
865
pluricolumn, GP/1E 4349A.
866 867
Measurements: See Table 7 for crinoid discs measurements.
868 869
Remarks: The overwhelming majority of Paleozoic crinoids are preserved as disarticulated
870
skeletal elements, mainly stem ossicles. In most cases linking them with other anatomical crinoid
871
parts (i.e., crowns) is virtually impossible. Therefore, in many cases, as is the case of the genus
872
øPentaridica, the taxonomy is based only on stem anatomy. Thus, stem-based genera have a 35
873
minimal chance to express true phylogenetic relationships (Głuchowski, 2002). Despite the
874
difficult determining the biological consistency of crinoid species on stem morphology alone (for
875
detailed discussion see Moore et al., 1968; Donovan 2001; Głuchowski, 2002) stem elements are,
876
in many cases, unique and allow specific identification.
877
The studied specimens are represented by 22 individual discs (articular facet) and three
878
pluricolumn remains (stems), which are, in the most part, poorly preserved. However, the
879
specimens GP/1E 4319 and 4349 B, C shows articular facets with well-preserved peripheral
880
crenularia, which can be referred to øPentaridica sp. (Dr. Julio Hlebszevitsch, personal
881
communication, April 2016).
882 883
4. Results
884
A total of 150 specimens were studied, belonging to Brachiopoda (77%), Mollusca (7%) and
885
Echinodermata, Crinoidea (16%) (Table 4). They were assigned to four brachiopod species
886
(Lyonia rochacamposi, Rhynchopora grossopunctata, Biconvexiella saopauloensis n. sp,
887
Quinquenella rionegrensis), four bivalve species (Phestia tepuelensis, Streblopteria aff. S.
888
lagunensis, Limipecten capivariensis, Praeundulomya cf. subelongata), and two gastropod
889
species [Peruvispira brasilensis n. sp, and Mourlonia (Woolnoughia)? sp.] (Tables 5, 6). Crinoid
890
columns were assigned to øPentaridica (Table 7). The overwhelming majority of brachiopod
891
shells belong to Biconvexiella saopauloensis (n= 86, 74,8%), and is followed by Rhynchopora
892
grossopunctata (n=20, 1,7%). Table 4 shows the total number of the remaining brachiopod
893
specimens. On the other hand, the bivalves are less common with just a few specimens of each
894
taxon (Table 4). The gastropods are also rare and represented by three shells only, two assigned
895
to Peruvispira brasilensis, and one to Mourlonia (Woolnoughia)? sp., this last one just illustrated
896
here (Fig. 6). Therefore, the assemblage is dominated by shells of Biconvexiella saopauloensis,
897
and is here referred as the Bs assemblage. 36
898
As noted above, the taphonomic information available for the bioclastic remains in the Bs
899
assemblage is limited, but some observations are noteworthy. First, the majority of both
900
brachiopod and bivalve shells are disarticulated. The most preserved specimens (articulated
901
shells) are illustrated in the figures 4 and 5. However, the shells are complete and the
902
ornamentation (when visible) is well preserved (i.e., signs of abrasion, bioerosion, and
903
encrustation are missing), and various size classes are represented. Closed articulated bivalve
904
mollusk shells are also observed. Indeed, Praeundulomya cf. subelongata is preserved with
905
closed articulated shells, and with the long axis inclined ~40° to bedding. Due to compaction, the
906
specimen is slightly compressed through the anterior-posterior axis of the shell. Thus, the angle
907
between the long axis and bedding has probably been reduced.
908
Finally, the crinoid remains are represented mainly by disarticulated skeletal ossicles
909
(mostly columnals), which are disperse in the matrix. Crinoid remains varies in size from 1.5 to
910
3.5 mm, and abrasion of crenelations and edges are not visible (i.e., columnals are pristine, see
911
Figure 6). Signs of encrustation, bioerosion and corrosion are also missing.
912 913
5. Discussion
914
5.1. Depositional environment
915
In the study area, the investigated marine fossil bed is recorded in the Taciba Formation
916
(formerly referred as Capivari Formation) and placed 100-150 m from the contact with the
917
overlying, post-glacial Early Permian Tatui Formation. The vertical profile reveals the interplay
918
of distinct sedimentary processes in a subaqueous environment and the presence of an
919
invertebrate marine fauna in a glacio-marine context. A clear change in the marine setting from a
920
low- to a high-energy environment is recorded, probably due to a glacier advance event in the
921
depositional setting (Fig. 3). Initially, sedimentation took place in a distal, low-energy
922
environment in an epeiric sea with free-floating icebergs. In this context, massive mudstones 37
923
overlying dark shales suggests increased sediment input (Fig. 3). The presence of bottom
924
currents, which resulted in current-rippled medium- to fine-grained sandstones are indicative of
925
bedform migration, coupled with sediment transportation by suspension (Fig. 3). Trough cross-
926
bedded strata interbedded with laminated siltstone occurs in the middle portion of the studied
927
section and records increased high-energy processes. This suggests migrating 3D bedforms
928
formed by subaqueous meltwater flows and/or turbiditic currents. Due to the rapid sedimentation
929
of these beds, flame structures were formed by the overloading of medium-grained massive
930
sandstones on fine-grained facies. Association with crudely parallel-laminated, plant-bearing
931
siltstones contrasts with the dysoxic environment during deposition of dark gray shales recorded
932
in the lowermost portion of the studied section. Increased sediment supply and input of
933
phytoclasts are suggestive of terrestrial contribution probably derived from contemporaneous
934
coastal progradational systems, or directly supplied by subaqueous outwash currents associated
935
with oscillating glacier margins in the context of a wet-based grounded ice shelf. To the top,
936
fossil-bearing silty shales record the environmental change to a low-energy shelfal depositional
937
setting (Fig. 3). The intervening of an episode of glacial retreat is corroborated by the absence of
938
iceberg-related dropstones and meltwater bottom currents. The relatively calm offshore
939
environment that was colonized by the marine invertebrates was eventually disrupted by high-
940
energy sedimentation events that enhanced fossil preservation. After that, a new glacial advance
941
event induced the sedimentation of pebbly mudstones that may have inhibited the bottom
942
colonization by shelly benthos. In the examined sedimentary succession, the percentage of
943
pebbles increases upward and the transition to pebbly silty sandstone is indicative of renewed
944
glacial sedimentation in the depositional setting.
945 946
5.2. Taphonomy
38
947
As commented above the taphonomic information available for the bioclastic remains in the Bs
948
assemblage is limited and mainly based on museum specimens that were not properly collected
949
for taphonomic studies. Anyway, some observations are worth to mention. Most of brachiopods
950
and bivalves are represented by disarticulated shells. However, the shells are complete and the
951
ornamentation (when visible) is well preserved, including various size classes. These features
952
indicate that after death most of the shells had, probably, limited exposure at the sediment/water
953
interface, suffering only minor lateral transport. These are probably parautochthonous (sensu
954
Kidwell et al., 1986) elements in this assemblage. By contrast, specimens of Praeundulomya cf.
955
subelongata were preserved in situ. Similar preservational conditions are also noted in
956
specimens of Praeundulomya and other associated Grammysioidea bivalves in the uppermost
957
part of the Taciba Formation, State of Paraná, southern Brazil (see Neves et al., 2014b). There,
958
the shells are preserved in fine to very fine sandstones/siltstones with hummocky cross-
959
stratification and intercalated mudstones forming stacked storm deposits in distal shoreface
960
settings (Neves et al., 2014b). As in the case of Praeundulomya from the Capivari beds, those
961
shells are closed articulated, compacted, and with the anterior margin pointing downward. The
962
long axis of the shell inclined 40°-50° to bedding. As convincingly demonstrated by Anelli et al.
963
(1998), such specimens were preserved in life position, and a similar condition is inferred for
964
Praeundulomya from the Bs assemblage. Hence, this species is the unique autochthonous
965
element so far identified in this assemblage.
966
The crinoid remains are all usually small (1.5 to 3.5 mm) disarticulated ossicles or rarely
967
articulated stem segments, without any signs of abrasion, encrustation, bioerosion and/or
968
corrosion. These taphonomic signatures indicates that after death and disarticulation, which can
969
occur in a few weeks, the columnals were not intensely transported and sorted (see Llewellyn
970
and Messing, 1993). Thus, the crinoid remains are probably parautochthonous. Finally, the
971
presence of comminuted plant remains (Fig. 3) is also worth mentioning, suggesting the 39
972
contribution of allochthonous materials from coastal settings during events of enhanced riverine
973
freshwater input.
974 975
5.3. Paleoautoecology
976
The Bs assemblage is numerically dominated by ambocoelids, a group of small, thin-shelled
977
brachiopods with an open delthyrium and a pedicle during life (see He et al., 2012). However,
978
the fine-grained, soft, siliciclastic-dominated facies where Biconvexiella saopauloensis is
979
recorded may have offered limited site opportunities for pedicle attachment. Indeed, the shells
980
are not found in clusters, and signs of encrustation are unknown in the studied specimens. Hence,
981
this tiny and spinose brachiopod species may have lived on soft substrates with the ventral valve
982
partially embedded within the substrate, a condition also shown by other ambocoelids (e.g.,
983
Ambocoelia gregaria Hall, 1860; in Zambito and Schemm-Gregory, 2013). The presence of
984
spine bases suggest that the shell was covered by tinny spines in life.
985
Given the dominance of Biconvexiella saopauloensis in the assemblage, this species may
986
haveacted as a pioneer species in muddy, dysoxic bottoms during deposition of host sediments.
987
Indeed, as for other ambocoelids (e.g., Attenuatella mengi He et al., 2012), this shell form is
988
usually associated with low oxygen and/or limited trophic resources in the sedimentary settings
989
inhabited by those brachiopods (see He et al., 2012 and references therein). Their small size is a
990
good indicator that the conditions were stressful, probably coupled with oscillations in the
991
oxygen content, salinity and food-supply. The linoproductids, such as Lyonia rochacamposi,
992
with spinose convex ventral valves, are interpreted as a quasi-infaunal, suspension feeders that
993
lived partially buried in soft substrates (see Cisterna et al., 2017). The other two brachiopods in
994
the Bs assemblage (Rhynchopora grossopunctata, Quinquenella rionegrensis) were epifaunal
995
suspension feeders.
40
996
The bivalves Streblopteria aff. S. lagunensis and Limipecten capivariensis were both
997
stationary epifaunal suspension feeders (Stanley, 1970, 1972), whereas Praeundulomya cf.
998
subelongata with a posteriorly elongated shell (Stanley, 1970) was a facultatively mobile
999
infaunal suspension feeder. As other nuculanoids, Phestia tepuelensis was a facultatively mobile
1000
infaunal deposit feeder/suspension feeder (Stanley, 1970). The other mollusks in the assemblage,
1001
the small gastropod Peruvispira and Mourlonia (Woolnoughia)? sp. were both facultatively
1002
mobile epifaunal suspension feeders or grazers (Aberhan and Kiessling, 2015). Yet, Peruvispira
1003
is known for its broad environmental tolerances in Early Permian marine assemblages from
1004
Australia (Clapham and James, 2008). Finally, the crinoid was a stationary epifaunal suspension
1005
feeder.
1006 1007
5.4. Paleocommunity and paleoenvironment
1008
With only 11 genera, the Biconvexiella saopauloensis assemblage is diverse when compared
1009
with other coeval benthic marine assemblages associated with LPIA sedimentary successions in
1010
South America (i.e., Argentina) and Australia (see data in Cisterna et al., 2019 and references
1011
therein). However, among the known macroinvertebrate marine faunas of the upper part of the
1012
Taciba Formation, the Bs assemblage is the most diverse one, including members of three major
1013
phyla (Brachiopoda, Mollusca and Echinodermata). The assemblage encompasses benthic
1014
invertebrates occupying distinct substrate tierings (Table 8). The epifaunal tiering encompassed
1015
(a) the low-level stationary epifaunal suspension feeders, including all the brachiopods, and
1016
epifaunal bivalves. Numerically, this was the dominant ecological group in the assemblage, (b)
1017
the stationary upper-level epifaunal suspension feeders (i.e., crinoids), and (c) vagile epifaunal
1018
suspension feeders or grazers (Aberhan and Kiessling (2015), including all the small gastropods.
1019
The shallow infaunal tiering was occupied by (d) facultatively mobile infaunal deposit
1020
feeders/suspension feeders, such as the protobranchiate bivalves (i.e., Phestia), and (e) the 41
1021
facultatively mobile shallow infaunal suspension feeders (i.e., Praeundulomya). Both infaunal
1022
groups colonized the fine-grained, siliclastic-dominated substrates of the Capivari marine beds
1023
just a few centimeters below the sediment water/interface. Surprisingly, planktic and nektic
1024
organisms are missing in the assemblage. The absence of these groups may be a result of a
1025
combination of processes related to taphonomic and sampling biases and/or stressful
1026
paleoenvironmental conditions. The fossil-bearing mudstones of the Taciba Formation are dark
1027
grey/green (or even yellowish when weathered), and finely laminated (horizontal bedding)
1028
indicating the settling of muds in quiet shelf, with restricted oxygen and food supply. Indeed, as
1029
commented by Harper and Moran (1997, p. 235) brachiopods, the numerically dominant group
1030
in the assemblage, were minimalist animals, “operating as a ciliary suspension feeder, if
1031
necessary, at very low oxygen levels.” The general small body size of the Capivari brachiopod
1032
fauna is in accordance with these observations.
1033
The presence of members of the rhynchonelliform brachiopods, sanguinolitid and pectinid
1034
bivalves, pleurotomariid gastropods and crinoids clearly indicates that the Bs assemblage is fully
1035
marine. However, the restricted vertical distribution of these marine groups in the uppermost part
1036
of the Itararé Group in the São Paulo State suggests that the background conditions were
1037
generally unfavourable to the profuse development of the benthic fauna or their preservation. As
1038
noted above, the invertebrates lived and/or were preserved in a distal glaciomarine environment
1039
during more proximal marine deglaciation (Figs. 7, 8, see Vesely and Assine, 2006, p. 164). The
1040
clay-rich, fossil-bearing marine horizon containing members of the Bs assemblage show no
1041
evidence of glacial influence (ice-rafted dropstones), indicating that this may correspond to a
1042
time of ice retreat during the sequence time span (Fig. 7, 8). This interval may correspond to a
1043
time of climatic amelioration, also supported by the presence of plant remains recorded in
1044
laminated siltstones, located nearly 2 meters below the marine fossil-bearing interval (Figs. 3, 7,
1045
8). This may indicate the presence of plant covered, deglaciated continental terrains (Fig. 8). 42
1046
Finally, Cisterna et al. (2017, and references therein) suggested that a complex array of
1047
abiotic factors encompassing variation in salinity and oxygen contents, water turbidity and depth,
1048
and food-supply are among the main controlling factors in the development of marine benthic
1049
faunas during deglacial events of LPIA. Notably, a comparison between the gross taxonomic
1050
composition and ecological structure of the Capivari marine fauna with those coevals from the
1051
upper part of the Taciba Formation, southern Brazil (Simões et al., 2012; Neves et al., 2014b;
1052
Taboada et al., 2016), strongly support the observations made by Cisterna et al. (2017). The
1053
benthic marine assemblages recorded in the so-called Rio da Areia sandstone and Baitaca
1054
siltstone, in the upper part of the Taciba Formation (Neves et al., 2014b), which are dominated
1055
by epifaunal and infaunal suspension feeding bivalves lived in quite distinct paleoenvironmental
1056
conditions compared to those of the Bs assemblage. Taphonomic and sedimentologic
1057
information (Neves et al., 2014b) conclusively indicates that the benthic faunas from the
1058
southern part of the basin lived in coastal to shelf areas commonly affected by moderate to
1059
strong bottom currents and episodically affected by high-energy flows and currents generated by
1060
storms, which is not the case of the Bs assemblage.
1061 1062
5.5. Biostratigraphic implications
1063
Our data support the correlation of the Bs assemblage with those from the uppermost part of the
1064
Taciba Formation, from southern Brazil, as previously pointed out by Taboada et al. (2016, p.
1065
437). Particularly noteworthy is the record of Lyonia rochacamposi that is common in the Taciba
1066
Formation from the Bela Vista do Sul assemblage (Taboada et al., 2016). Additionally, close
1067
species of Quinquenella, Phestia, Limipecten, and Praeundulomya are also recorded in the
1068
Teixeira Soares assemblages (Neves et al., 2014b; Taboada et al., 2016). Hence, the Bs
1069
assemblage correlates with the upper portion of the Itararé Group from southern Brazil,
1070
suggesting an early Permian (latest Asselian-earliest Sakmarian) age. These data indicate that the 43
1071
marine fossil-bearing beds of Capivari and Teixeira Soares regions are stratigraphic records of
1072
northward transgressive events during the end of LPIA in the Paraná Basin. Faunal similarities
1073
suggest that the latest Asselian-earliest Sakmarian Paraná marine assemblages correlate with
1074
those of the Sauce Grande-Colorado (Argentina), Huab (Hardap shale of the Dwyka Group),
1075
Aranos area (Namibia), southwest Africa, and the Carnarvon (Western Australia) basins, and
1076
characterize the Eurydesma-Lyonia chronozone, reinforcing the paleogeography scenario of a
1077
W-E trans-Gondwanan marine seaway linking those basins during the Early Permian, as detailed
1078
discussed in Taboada et al. (2016).
1079 1080
6. Significance and concluding remarks
1081
In this contribution, we report a cool-water fauna from the LPIA, which was found in a fossil-
1082
rich bed of the upper part of the Itararé Group from the Capivari county, on the eastern border of
1083
the Paraná Basin. The fauna was described and its stratigraphic position within the glacial Itararé
1084
Group better constrained. The measured sedimentary succession is placed between 100-150 m
1085
below the contact with the overlying, post-glacial Early Permian Tatuí Formation.
1086
Members of the Biconvexiella saopauloensis assemblage flourished in a rather confined
1087
paleogeographic setting in the northeastern border of the Paraná Basin, under stressful conditions
1088
mainly linked to changes in oxygen content and food-supply. Although evidences of freshwater
1089
discharge from coastal areas are recorded in the studied sedimentary succession, the Bs
1090
assemblage is fully marine, suggesting that salinity conditions were normal, at least during the
1091
time of benthic faunal development.
1092
The fossil-bearing marine beds of Capivari county may represent short-lived transgressive
1093
events during periods of ice retreats. These beds are overlain by pebbly mudstones and silty
1094
sandstones recording high energy processes under glacial conditions. Since Bs assemblage is
1095
placed in between subaqueous glacial facies associations, they may correspond to high frequency 44
1096
interglacials. Consequently, the fossil-bearing marine beds generated in this stratigraphic context
1097
are thin and difficult to find within the thick glacial package. This is particularly true when
1098
proper subsurface data are missing and/or outcrops are deeply weathered and covered by dense
1099
vegetation, which is the case for the studied fossil-bearing beds. Faunal composition indicates
1100
that the transgressive fossil-bearing beds of the Capivari region correlates with those of the upper
1101
part of the Taciba Formation from the southern part of the Paraná Basin (States of Paraná and
1102
Santa Catarina), suggesting an Early Permian (latest Asselian-earliest Sakmarian) age. Indeed, in
1103
the upper part of the Taciba Formation, the spore-pollen assemblages are referable to the
1104
Vittatina costabilis Biozone of the Paraná Basin indicative of an Early Permian age (Souza and
1105
Callegari, 2004; Souza and Toigo, 2005; Souza, 2006). These marine faunas also indicate the
1106
increase of glacial-marine depositional conditions in the uppermost part of the Itararé Group,
1107
which is tied to the last cycle of ice advance and retreat during the LPIA.
1108 1109
Acknowledgments
1110
The authors are very indebted to the Instituto de Geociências, Universidade de São Paulo, São
1111
Paulo, for access and support in the study of the invertebrate collection of the Capivari fossils.
1112
We also thank Ivone C. Gonzales, IGc/USP, for assistance with this collection. Julio
1113
Hlebszevitsch helped us with crinoid identification and Suzana A. Matos, IBB/UNESP, assisted
1114
us in the field. Hence, both are here acknowledged. Many thanks also to Paul A. Johnston, and
1115
two anonymous reviewers for their suggestions, corrections and comments that improved the
1116
final version of the manuscript. Editor-in-Chief, Francisco J. Vega is also acknowledged for the
1117
useful suggestions and corrections. Finally, we also thank the paleoillustrator Julio L.C. Bezerra
1118
for the “Capivari sea” paleoscene. Financial support was offered by FAPESP and CNPq, Brazil,
1119
and CONICET, Argentina. JPN was a fellow of FAPESP (grant 2013/25317-7). This project is a
45
1120
contribution to the following FAPESP projects: 2013/25317-7 and 2014/09149-0. CNPq grants:
1121
304925/2017-9 to MLA and 301023/94-3 to MGS.
1122 1123
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Cincinnati, 253 p. Miller, S.A., 1889. North American Geology and Paleontology: Cincinnati, Ohio, Western Methodist Book Concern Press, 793 p. Mineropar., 2007. Mapeamento geológico da folha de Ponta Grossa. Governo do Estado do Paraná, scale 1:100.000. Moore, R.C.; Jeffords, R.M., and Miller, T.H., 1968. Morphological features of crinoid columns. The University of Kansas Contributions, 8, 1–30. Morris, N.J., Dickins, J.M., and Astafieva-Urbaitis, K., 1991. Upper Paleozoic
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Newell, N.D., 1938. Late Paleozoic pelecypods: Pectinacea. University of Kansas State Geological Survey. Kansas, 10, 1–123. Newell, N.D., and Boyd, D.W., 1990. Nacre in a Carboniferous pectinoid mollusc and a new subfamily Limipectininae: American Museum Novitates, 2970, 1–7. Newell, N.D., and Boyd, D.W., 1995. Pectinoid bivalves of the Permian–Triassic crisis: American Museum of Natural History Bulletin, 227, 1–95. Newell, N.D., Chronic, J., and Roberts, T.G., 1953. Upper Paleozoic of Peru: Geological Society of America, Memoir (Boulder), 58, 1–276. Oliveira, E.P., 1930. Fósseis marinhos na Série Itararé no Estado de Santa Catarina: Anais da Academia Brasileira de Ciências, 2, 17–21. Oliveira, E.P., 1936. Um novo braquiópode da Serie Itararé: Notas Preliminares e Estudos, Divisão de Geologia e Mineralogia, 5, 8–10.
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1447
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1499 1500 1501 1502 1503 1504
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1507 1508 1509
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1511
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1512
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1519 1520
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1521
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1522
Waterhouse, J.B., 2013. The Evolution and Classification of Productida (Brachiopoda):
1523 1524 1525 1526 1527 1528
Earthwise, 10, 1–535. Wenz, W. 1938. Gastropoda, Teil I. In: Schindewolf, O.H. (Ed.), Handbuch der Paläozoologie, Verlag Gebrüder Bornträger, 1639 p. White, I.C., 1908. Relatório sobre as Coal Measures e rochas associadas ao sul do Brasil: Comissão das Minas de Carvão de Pedra do Brasil, Rio de Janeiro, 300 p. Williams, A., Carlson, S.J., Brunton, C.H.C., Holmer, L.E., and Popov, L.E., 1996. A supra-
1529
ordinal classification of the Brachiopoda: Philosophical transaction of the Royal Society of
1530
London (series B), 351, 1171–1193.
1531
Zambito, J., and Schemm-Gregory, M., 2013. Revised taxonomy and autecology for the
1532
brachiopod genus Ambocoelia in the Middle and Late Devonian northern Appalachian Basin
1533
(USA): Journal of Paleontology, 87, 277–288.
1534 62
1535
Captions
1536 1537
Figure 1. Stratigraphic chart of Itararé Group showing the main marine mudstone intercalations
1538
and their lateral correlations. Capivari assemblage (= Biconvexiella saopauloensis assemblage
1539
here described). Araçoiaba, Ortigueira and Mafra assemblages are recorded in thin marine
1540
intercalations in a thick sandstone package. Explanation: LPTS, Late Paleozoic Third Order
1541
Sequence; SB, Sequence Boundary; ass., Assemblage. Modified from Holz et al. (2010) and
1542
Taboada et al. (2016).
1543 1544
Figure 2. A: Distribution of the upper Paleozoic strata (Itararé Group) in the Paraná Basin,
1545
southeastern Brazil and location of the studied section (Capivari marine beds). The location of
1546
the Passinho and Lontras marine beds, southern Brazil, is also indicated. B: Geological map with
1547
the distribution of the upper Paleozoic rocks in the studied area (black star). Source of data: IPT
1548
(1981).
1549 1550
Figure 3. Columnar section of the upper portion of the Itararé Group, Taciba Formation, in the
1551
Capivari region, State of São Paulo, Brazil. The fossil-bearing interval is highlighted.
1552
Abbreviations: C=clay; S = silt; FS = fine sand; MS= medium sand; CS= coarse sand. A: Pebbly
1553
mudstone in the upper portion of the section. B: Fossil-bearing silty shale. C: Sample of
1554
laminated siltstone with rich concentrations of fragmented plant material. Scale bars 1 cm.
1555 1556
Figure 4.
1557
Brachiopods of the Taciba Formation, Capivari county. 1-12: Rhynchopora grossopunctata
1558
Mendes, 1952. 1: conjugated valves, ventral valve view, GP/1T 117a; 2: dorsal valve, GP/1E-
1559
451; 3-5: ventral valves; 3: GP/1E 117c; 4: GP/1E-2507; 5: GP/1E-2545; 6, 11: conjugated 63
1560
valves, GP/1E-4342; 6: dorsal valve view; 11: posterior view; 7: cast of external mold of dorsal
1561
valve, GP/1E-4405; 8: ventral valve, GP/1E-443a; 9, 10, 14, 15: dorsal valves, GP/1E-4325; 9:
1562
dorsal valve view; 10: anterior view; 14: lateral view; 15: posterior view; 12: external mold of
1563
ventral valve, GP/1T 117b; 13: detail of the ventral valve of GP/1E 443b, showing punctae
1564
(5/mm2); 16-25: Biconvexiella saopauloensis sp. nov., 16-19: dorsal valves; 16: cast of GP/1E-
1565
4339 (holotype); 17: natural mold; 18: GP/1E-453; 19: GP/1E 450; 20: external mold of ventral
1566
valve, GP/1E 4322; 21, 22: GP/1E 2513; 21: conjugated valves; 22: close-up of the
1567
subconcentric rows of spine bases; 23: cast of external mold of dorsal valve, GP/1E-4327; 24-25:
1568
ventral valve, 24: GP/1E 2520; 25: GP/1E-4338. 26-28: Quinquenella rionegrensis (Oliveira), 26:
1569
dorsal valve, GP/1E 4355c; 27-28: dorsal valve interior, 27: GP/1E-4354c; 28: GP/1E 4358; 29-
1570
32: Lyonia rochacamposi Taboada et al., 2016; 29: ventral valve in oblique view, GP/1E-4350b;
1571
30: CP.I 769I, specimen from Bela Vista do Sul locality (Southern Brazil), showing a single row
1572
of spines (white arrow); 31: cast of a fragmentary ventral valve, external mold, GP/1E 4350b; 32:
1573
fragmentary ventral valve, external mold, GP/1E 4350a. Scale bar = 10 mm, except in 19; 20;
1574
26-28 = 5 mm. Specimens 2, 6, 11-13; 20-22, 26 are external molds. 3-5; 8-10, 14-19, 24, 25 are
1575
internal molds.
1576 1577
Figure 5.
1578
Bivalves of the Taciba Formation, Capivari county. 1-4: Phestia tepuelensis, 1-3: GP/1E-452; 1:
1579
external mold of right valve; 2: latex cast of 1; 3: detail of 1, showing the radial ribs crossing the
1580
commarginal costae; 4: internal mold of right valve, GP/1E-455. 5-7: Streblopteria aff.
1581
lagunensis, GP/1E-457, internal mold of right valve; 5: dorsal view; 6: antero-posterior view. 7:
1582
right valve view; 8-10: Limipecten capivariensis (Mendes), 8: latex cast of left valve fragment,
1583
GP/1T-116; 9: cast of external mold of right valve, GP/1E 2494; 10: internal mold of
64
1584
undetermined valve, GP/1E 2511; 11-12: Praeundulomya cf. subelongata Dickins, GP/1E-4252;
1585
11: right valve view; 12: ventral margin view. Scale bar = 10 mm, except in 9 and 10 = 5 mm.
1586 1587 1588
Figure 6. Gastropod and crinoid remains of the Taciba Formation, Capivari county. 1-3:
1589
Gastropods, 1-2: Peruvispira brasilensis n. sp. 1: latex cast of external mold ,GP/1E 4348
1590
(holotype); 2: cast of external mold, GP/1E-4319; 3: Mourlonia (Woolnoughia)? sp., GP/1E-454;
1591
4-6: øPentaridica sp. (personal communication Dr. Julio Hlebszevitsch, April 2016); 4: Articular
1592
facet showing the peripheral crenularium [L2]GP/1E-4321; 5: Detail of a pluricolumn, GP/1E-
1593
4349A; 6: two articular facets with well-preserved peripheral crenularium, GP/1E-4349. Scale
1594
bar = 5 mm.
1595 1596
Figure 7. Sedimentary facies relationships of the Taciba Formation, Capivari County, State of
1597
São Paulo, Brazil. Note the dropstone-bearing shales in the distal areas, and the glacier melting
1598
associated with the cross-stratified sandstones intercalated with plant-rich mudstones. Members
1599
of the Biconvexiella saopauloensis assemblage are recorded in the upper, fossil-bearing silty
1600
shales.
1601 1602
Figure 8. Reconstruction of the hypothesized siliciclastic-dominated, low-energy, shelfal
1603
bottoms of the “Capivari sea”, during a short-lived deglacial event. Note the dominance of
1604
rhynchonelliform brachiopods, with subordinated bivalves, gastropods and crinoids. Explanation:
1605
(1) øPentaridica sp.; (2) Peruvispira brasilensis n. sp.; (3) Quinquenella rionegrensis; (4)
1606
Lyonia rochacamposi; (5) Rhynchopora grossopunctata; (6) Biconvexiella saopauloensis n. sp.;
65
1607
(7) Limipecten capivariensis; (8) Streblopteria aff. S. lagunensis; (9) Phestia tepuelensis and (10)
1608
Praeundulomya cf. subelongata. Illustration by J.L.C. Bezerra.
1609 1610
Table 1. Age of the Biconvexiella saopauloensis assemblage within the glacial succession of the
1611
Itararé Group, Paraná Basin, according to various authors.
1612 1613
Table 2- Correlation of the Itararé Group macroinvertebrate faunas, based on literature data.
1614 1615
Table 3. Faunal composition, suggested age and biocorrelation of the Biconvexiella
1616
saopauloensis assemblage, Taciba Formation, Itararé Group, State of São Paulo, Brazil.
1617 1618
Table 4. Taxonomic composition of the Biconvexiella saopauloensis marine assemblage, Taciba
1619
Formation, Itararé Group, Paraná Basin, southeastern Brazil. Explanation: *species illustrated in
1620
this contribution, but not formally described.
1621 1622
Table 5. Measurements of the brachiopod shells, Biconvexiella saopauloensis assemblage,
1623
Taciba Formation, Itararé Group, Paraná Basin, southeastern Brazil. Explanation: V= ventral
1624
valve; D= dorsal valve; F= fragment; C= conjugated valves.
1625 1626
Table 6. Measurements of the mollusk shells, Biconvexiella saopauloensis assemblage, Taciba
1627
Formation, Itararé Group, Paraná Basin, southeastern Brazil. Explanation: R= right valve; L=
1628
left valve; C= conjoined (articulated) valves; U= undetermined valve.
1629
66
1630
Table 7. Measurements of the crinoid remains, Biconvexiella saopauloensis assemblage, Taciba
1631
Formation, Itararé Group, Paraná Basin, southeastern Brazil. Explanation: *individual discs;
1632
**stem (pluricolumn).
1633 1634
Table 8. Guild diversity and abundance of the marine invertebrates, Biconvexiella saopauloensis
1635
assemblage, Taciba Formation, Itararé Group, Paraná Basin, southeastern Brazil. Guild
1636
classification based on Aberhan and Kiessling (2015).
67
Authors Itararé faunas
Rocha-Campos and Rösler (1978)
Petri and Souza (1993)
Holz et al. (2010)
Simões et al. (2012)
Neves et al. (2014b)
Taboada et al. (2016)
This study
São Paulo
Paraná
Santa Catarina
Artinskian-Kungurian
Kungurian
---
---
---
---
Itaporanga
---
Kungurian
---
---
---
---
Araçoiaba
Late Carboniferous (Stephanian)-Early Permian
Late Carboniferous (WestphalianStephanian)
---
---
---
---
---
Guaraúna
Kungurian
Artinskian
---
---
---
---
---
Ortigueira
Artinskian
Artinskian
---
---
---
---
---
Teixeira Soares
Kungurian
Artinskian
---
Late AsselianSakmarian
Asselian
Latest AsselianEarly Sakmarian
Latest AsselianEarly Sakmarian*
Artinskian-Kungurian
Artinskian
Gzelian-Asselian (C-P boundary)
---
---
---
---
Mafra
Artinskian
Artinskian
---
---
---
---
---
Bela Vista do Sul (Butiá)
Kungurian
Kungurian
---
Late AsselianSakmarian
Late AsselianEarly Sakmarian
Late AsselianEarly Sakmarian
Latest AsselianEarly Sakmarian*
Capivari
Lontras
Latest AsselianEarly Sakmarian
Faunas São Paulo Studies
Capivari
Hortolândia
Mezzalira (1956) Loczy (1964) Rocha-Campos (1969)
X
Paraná Guaraúna
Santa Catarina Ortigueira
Bela Vista do Sul
Itaporanga
Teixeira Soares
X
X
X
X
X
Mafra
X
Saad (1977)
X
X
Rocha-Campos & Rösler (1978)
X
X X
Canuto (1985)
X X
Simões et al. (2012)
Lontras
Rio Grande do Sul
X
X X
Budó
Capivari Marine Invertebrate Fauna Authors
Fossils
Age
Correlation
Barbosa and Almeida (1949a, b) Ambocoelia planoconvexa; Rhynchopora; Aviculopecten sp. gastropods
-------------
Late Paleozoic deposits from Bolivia and Amazon Basin, Brazil
Mendes (1952)
Crurithyris aff. planoconvexa; Rhynchopora grossopunctata Aviculopecten capivariensis; Nuculana ? sp.
Late Carboniferous /Permian
Teixeira Saores, Bela Vista do Sul and Bagé marine beds (=Taciba Formation), Paraná Basin, and Barreal beds, Argentina
Barbosa and Gomes (1958)
RochaCampos (1966)
Peruvispira delicata -------------
Early Pennsylvanian
-------------
Early Permian
Copacabana Formation, Peru
RochaCampos (1967)
as in RochaCampos (1966)
-------------
-------------
RochaCampos (1969) Attenuatella paulistana sp. nov. (nom. nud.) Limipecten capivariensis (nom. nud.);
Older than Teixeira Soares marine fauna (middle portion of the Itararé Group)
Teixeira Soares marine bed (=Taciba Formation), Paraná Basin, Brazil
Saad (1977)
Attenuatella sp.; Rhynchopora grossopunctata Limipecten capivariensis (nom. nud.); Phestia sp.; Streblopteria sp.; Peruvispira delicata; crinoids
Artinskian
Hortolândia shale fauna, Paraná Basin, Brazil
Rocha-Campos and Rösler (1978) Attenuatella sp. nov.; Rhynchopora grossopunctata Phestia sp.; Streblopteria sp.; Peruvispira delicata crinoids
Early Permian (middle portion of the Itararé Group)
Hortolândia shale fauna, Paraná Basin, Brazil
Petri and Souza (1993)
-------------
Artinskian
Teixeira Soares marine bed (=Taciba Formation), Paraná Basin, Brazil
This study
Phestia tepuelensis Streblopteria aff. lagunensis Limipecten capivariensis Praeundulomya cf. subelongata Peruvispira brasiliensis n. sp. Mourlonia (Woolnoughia)? sp. Lyonia rochacamposi Rhynchopora grossopunctata Biconvexiella saopauloensis n. sp. Quinquenella rionegrensis Pentaridica sp.
Latest Asselian-Earliest Sakmarian
Uppermost part of the Taciba Formation (Teixeira Soares and Butiá assemblages)
Bivalvia
Gastropoda
Brachiopoda
Crinoidea
Family
Species
Studied specimens 2
Polidevciidae
Phestia tepuelensis
Streblochondriidae
Streblopteria aff. lagunensis
1
Limipectinidae
Limipecten capivariensis
3
Sanguinolitidae
Praeundulomya cf. subelongata
1
Eotomariidae
Peruvispira brasilensis n. sp.
2
Gosseletinidae
Mourlonia (Woolnoughia)? sp.*
1
Auriculispinidae
Lyonia rochacamposi
2
Rhynchoporidae
Rhynchopora grossopunctata
20
Ambocoeliidae
Biconvexiella saopauloensis n. sp.
86
Rugosochonetidae
Quinquenella rionegrensis
7
øPentacauliscidae
Pentaridica sp.
25
Measurements (mm) Width
Length
Thickness
GP/1E-4342 (C)
19
16
4
GP/1E-443a (V)
14
15
-
GP/1E-443b (V)
~17
~15
-
GP/1E-2507 (V)
~14
~12
-
GP/1E-2545 (V)
~16
~18
~3
GP/1E-451 (D)
18
16
-
GP/1E-1912 (D)
17
~16
-
GP/1E-4325 (D)
14
~13
9
GP/1E-4405 (D)
13
14
-
GP/1E-2527 (V)
~7
~6
-
GP/1E-2504 (V)
~10
9
4
GP/1E-4338 (V)
~6
9
~2
GP/1E-2520 (V)
9
~9
5
GP/1E-4354b (V)
8
9
4
Biconvexiella saopauloensis
GP/1E-4327 (D)
10
8
-
sp. nov
GP/1E-2543 (D)
10
9
-
GP/1E-2538 (D)
~11
~8
-
GP/1E-453 (D)
11
9
-
GP/1E-4339 (D)
11
9
-
GP/1E-2513 (D)
~11
8
-
GP/1E-4358 (D)
5
9
-
GP/1E-459 (V)
10
9
-
GP/1E-4354c (D)
11
8
-
GP/1E-4350a (V)
~13
~22
-
GP/1E-4350b (F)
~35
~26
-
GP/1E-4351a (F)
~31
~15
-
Species
Rhynchopora grossopunctata
Quinquinella rionegrensis
Lyonia rochacamposi
Specimen (valve)
Measurements (mm)
Bivalvia
Gastropoda
Species
Specimen
Length
Height
Width
Phestia tepuelensis
GP/1E-452 (R)
21
11
---
GP/1E-455 (R)
~23
11
---
Streblopteria aff. lagunensis
GP/1E-457 (R)
~21
20
~4
Limipecten capivariensis
GP/1T-116 (L)
~8
~90
---
GP/1E-2511 (U)
~6
~9
---
GP/1E-2494 (R)
9
6
---
Praeundulomya cf. subelongata
GP/1E-5242 (C)
69
27
~13
Peruvispira brasilensis n. sp.
GP/1E-4319
5,3
7,8
---
GP/1E-4348
3,8
~4,5
---
Crinoidea
Specimen
øPentaridica sp.
GP/1E-4321
Major axis (mm) 3,5*
GP/1E-4349 A
2,5 **
GP/1E-4349 B
2,0*
GP/1E-4349 C
1,5*
GP/1E-4350C
~1,2*
Taxon Lyonia Rhynchophora Biconvexiella Quinquenella Streblopteria Limipecten Phestia Praeundulomya
Mourlonia (Woolnoughia) Peruvispira øPentaridica Total
Stationary lowlevel epifaunal suspension feeder 2 20 86 7 1 3 ----------119
Facultatively mobile, infaunal, deposit feeder ------------1 --------1
Facultatively mobile infaunal suspension feeder --------------2 ------2
Epifaunal grazer ----------------1 2 3
Stationary upperlevel epifaunal suspension feeder --------------------25 25
Asselian
Pennsylvanian
303.4 +/- 0.9
Kasimovian 307.2 +/- 1.0
Moscovian 311.7 +/- 1.1
Bashkirian 318.1 +/- 1.3
Itararé Group
298.9 +/- 0.1
Gzheian
Late Carboniferous
Taciba Fm.
EurydesmaLyonia fauna
295.0 +/- 0.1
Tubarão Supergroup
Permian
Cisuralian
Sakmarian
MAH
subzone*
São Paulo Capivarí ass.
Chronostratigraphic Chart Paraná Teixeira Soares ass.
South-Southeast
Rio Grande do Sul
Santa Catarina Butiá BA ass.
Passinho Shale
Rio do Sul Formation
Budó facies
Taciba Guaraúna ass. Araçoiaba ass.
Ortigueira ass.
Sequences
Lontras ass. Mafra ass.
SB-3 Suspiro facies
Lontras shale (Carb./Permian transition)
LPTS -2 Mudstones
Campo Mourão Fm.
Campo Mourão
Sandstones Lapa/ Vila Velha incised valley fill
Lagoa Azul Fm.
Lagoa Azul
“Folhelho Roncador” transgression
Conglomerates Hiatus
SB-2
Incised valley Lithostratigraphic LPTS -1 contact 900 km SB-1 aprox. horiz. scale
Gondwana 1 Supersequence (Milani et al., 2007)
North-Northwest Geochronology Lithostratigraphy Biostratigraphy
A
B
47°30'
47°20'
N
Cuiabá Brasilia
GO
MT
5 km
700
Monte Mor 18º
MG 70
0
900
MS
23°00'
Capivari
1100 1300
1-AB-1-SP
SP
1100
ld
úc
pi
São Paulo
na enti
ad
r
ve
Arg
Ro
Ri 23°10'
ad
lto
e
N
Rio do Sul Lontras Shale
500
Atlantic Ocean 24ºS
Sa
Curitiba
t Tie
Teixeira Soares Passinho Shale
700
Ro
ri-
ar
va
Paraguay
Aç
Ca
PR
900
O
Capivari
Capivari Beds
Salto
SC
300
200 km
Brazil
RS
0
10
0
10
Paraná Basin 54º
Porto Feliz
Porto Alegre
50ºW
Basin border Isopachs (m) Itararé Group outcrop belt Exploration well
Intrusive rocks
Tatuí Fm.
Corumbataí Fm.
Itararé Group
Irati Fm.
Itu Suit
Cities Outcrop Roads
Secondary Road Rivers
25 m
A
A 20
B
B 15
C
C 10
Dark shale with dropstones
5
Shale
Flame structure Plant remains
Silt Shale
Brachiopods
Massive mudstone
Bivalves
Pebbly mudstone
Gastropods
Siltstone
Crinoids
Pebbly silty sandstone Massive sandstone Current-rippled sandstone
0
C S FS MS CS
Trough cross-stratified sandstone
1
2
3
4
5
6
7
8
9
10
11
12
13
14
16
17
18
20
19
26
25
23
21
15
22
24
30 27
28
29
31
32
1
2
4
5
3
6
t ea tr re e Ic Silty shale Cross-stratified sandstone Massive mudstone Shale with dropstones Vegetation
1
2
4
3
5
7
6
8
11
10
9
12
Simões et al., JSAES, CRediT author’s statements
Title: Mollusks and brachiopods of the Capivari Marine Bed, late Paleozoic glacial Itararé Group, northeast Paraná Basin, Brazil: paleoenvironmental and paleogeographic implications Simões Marcello: Supervision; conceptualization; writing (with input from all authors); field collecting (geological and paleontological); funding acquisition; Neves Jaqueline: General systematic analysis; data curation; writing (with input from all authors); figure design and visualization; Taboada Arturo: Conceptualization; general systematic analysis; data curation, writing (with input from all authors); Pagani Maria: General systematic analysis; data curation; writing (with input from all authors); Varejão Filipe: Field collecting (geological and paleontological) data; geological interpretations; figure design and visualization; writing (review and editing); Assine Mário: Field collecting (geological and paleontological) data; geological interpretations; writing (review and editing).
S0895-9811(19)30443-2
DOI:
https://doi.org/10.1016/j.jsames.2019.102433
Reference:
SAMES 102433
To appear in:
Journal of South American Earth Sciences
Received Date: 27 August 2019 Revised Date:
19 November 2019
Accepted Date: 20 November 2019
Please cite this article as: Simões, Marcello.Guimarã., Neves, J.P., Taboada, Arturo.Cé., Pagani, M.A., Varejão, F.G., Assine, Má.Luis., Mollusks and brachiopods of the Capivari marine bed, late Paleozoic glacial Itararé Group, northeast Paraná Basin, Brazil: Paleoenvironmental and paleogeographic implications, Journal of South American Earth Sciences (2019), doi: https://doi.org/10.1016/ j.jsames.2019.102433. This is a PDF file of an article that has undergone enhancements after acceptance, such as the addition of a cover page and metadata, and formatting for readability, but it is not yet the definitive version of record. This version will undergo additional copyediting, typesetting and review before it is published in its final form, but we are providing this version to give early visibility of the article. Please note that, during the production process, errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. © 2019 Published by Elsevier Ltd.
1
Mollusks and brachiopods of the Capivari Marine Bed, late Paleozoic glacial
2
Itararé Group, northeast Paraná Basin, Brazil: paleoenvironmental and
3
paleogeographic implications
4 5
Marcello Guimarães Simões
6
Instituto de Biociências, Departamento de Zoologia, Universidade Estadual Paulista, Distrito de
7
Rubião Junior, Botucatu, São Paulo, Brazil.
8 9
Jacqueline Peixoto Neves*
10
Universidade Tecnológica Federal do Paraná, campus Dois Vizinhos, Dois Vizinhos, Paraná,
11
Brazil.
12
*Corresponding autor
13 14
Arturo César Taboada
15
Centro de Investigación Esquel de Montaña y Estepa Patagónicas, CONICET-UNPSJB, Esquel
16
U9200, Chubut, Argentina.
17 18
Maria Alejandra Pagani
19
Museo Paleontológico Egidio Feruglio – CONICET, Avenida Fontana nº 140, Trelew
20
U9100GYO, Chubut, Argentina.
21 22
Filipe Giovanini Varejão
23
Instituto de Geociências e Ciências Exatas, Departamento de Geologia Aplicada, Universidade
24
Estadual Paulista, Campus de Rio Claro, Rio Claro, São Paulo, Brazil.
25
26 27
Mário Luis Assine
28
Instituto de Geociências e Ciências Exatas, Departamento de Geologia Aplicada, Universidade
29
Estadual Paulista, Campus de Rio Claro, Rio Claro, São Paulo, Brazil.
30 31
1
32
Abstract.—A 2-m-thick silty shale bed within the Taciba Formation, Itararé Group, Paraná
33
Basin, State of São Paulo, southeastern Brazil, records marine sedimentation in a siliciclastic-
34
dominated, low-energy, shelf setting, during a short-lived deglacial event. The bed is located
35
100–150 m below the base of the lower Permian, post-glacial Tatui Formation. The marine
36
assemblage is dominated by rhynchonelliform brachiopods, with subordinate bivalves,
37
gastropods and crinoids, recording the highest phylum-level diversity so far identified within a
38
given fossil-bearing horizon in the uppermost portion of the Itararé Group. Two new species are
39
described, one brachiopod Biconvexiella saopauloensis and one gastropod Peruvispira
40
brasilensis. Additionally, shells of Lyonia rochacamposi, Rhynchopora grossopunctata,
41
Quinquenella rionegrensis, Phestia tepuelensis, Streblopteria aff. S. lagunensis, Limipecten
42
capivariensis, Praeundulomya cf. subelongata and Mourlonia (Woolnoughia)? sp.are identified.
43
Crinoid columns were assigned to øPentaridica sp. (a genus based on elements of the columnal).
44
This is the first systematic description of members of the Eurydesma-Lyonia fauna in the
45
northeastern part of the Paraná Basin, Brazil. The overwhelming majority of brachiopods belong
46
to Biconvexiella saopauloensis, followed by Rhynchopora grossopunctata. The record of Lyonia
47
rochacamposi closely resembles that of the uppermost part of the Taciba Formation in southern
48
Brazil. Hence, the Capivari marine fauna correlates approximately with that of the upper part of
49
the Taciba Formation. Lyonia rochacamposi also indicates correlation with Permian units of the
50
Sauce Grande-Colorado (Argentina), Huab (Hardap shale of the Dwyka Group), Aranos area
51
(Namibia), southwest Africa, and the Carnavon (Western Australia) basins. These correlations
52
support a latest Asselian-earliest Sakmarian age for the fauna.
53
2
54
Highlights
55
A marine cool-water benthic fauna developed during Late Paleozoic Ice Age is described;
56
Fauna developed during a transgression coupled with a time of climatic amelioration;
57
Assemblage flourisched under restrict paleogeograhic conditions.
58 59
Keywords: Permian, Taciba Formation, macroinvertebrate, Paraná Basin, Capivari.
60 61
1. Introduction
62
Glaciogenic rocks of the Carboniferous-Permian Itararé Group, Paraná Basin, Brazil, are known
63
since Derby (1878) and White (1908). The succession includes one of the most complete
64
sedimentary records of the Late Paleozoic Ice Age (LPIA, sensu Shi and Waterhouse, 2010) that
65
waxed and waned across the central Gondwana (Rocha-Campos, 1967; França and Potter, 1988;
66
Holz et al., 2010). However, due to the large size of the basin (>1,500,000 km2) and the complex
67
depositional processes associated with the glacial and post-glacial events, no consensus exists
68
about the correlation, age and extent of the Itararé Group (Loczy, 1964; Rocha-Campos, 1967;
69
1970; Saad, 1977; Rocha-Campos and Rösler, 1978; França and Potter, 1991; Castro, 1999; Holz
70
et al., 2010).
71
Within the glaciomarine successions of the Itararé Group, as shown by Santos et al. (1996),
72
the sea-level rise following ice retreats was often accompanied by widespread marine deposits,
73
usually ice-rafted, debris-free, dark-gray, fossil-poor shales. Five regionally continuous
74
stratigraphic intervals with marine beds are recognized (Saad, 1977; Rocha-Campos and Rösler,
75
1978; Petri and Souza, 1993; Santos et al., 1996; Holz et al., 2010), which are known by their
76
local geographic names, as follows: (1) Araçoiaba, (2) Mafra-Ortigueira, (3) Lontras-Guaraúna
77
(upper Pennsylvanian), (4) Capivari, and (5) Passinho (Rocha-Campos and Rösler, 1978; Santos
78
et al., 1996), whose stratigraphic position within the glacial succession varies according to 3
79
distinct authors (Tables 1, 2). These marine beds record extensive transgressive events associated
80
with regional episodes of glacier retreats (see Loczy, 1964, p. 37; França and Potter, 1988;
81
Castro, 1999; Vesely and Assine, 2006; Mineropar, 2007; Neves et al., 2014a, b; Taboada et al.,
82
2016). Hence, some of these deposits (i.e., Lontras and Passinho shales) may have represented
83
the maximum flooding surfaces within the depositional sequence (Vesely and Assine, 2006;
84
Holz et al., 2008; 2010). Yet, within these marine beds at least eight distinct marine benthic
85
assemblages are known (Rocha-Campos and Rösler, 1978; Neves et al., 2014a, b; Taboada et al.,
86
2016).
87
As shown in Table 2, at the outcrop belt of the eastern border of the Paraná Basin, fossil-
88
rich marine intercalations of the Itararé Group are difficult to correlate regionally (Holz et al.,
89
2010). This is the case for the fossil-bearing beds of the Taciba Formation (former Capivari
90
Formation, Table 3) exposed in the central-eastern region of the State of São Paulo, southeastern
91
Brazil (Figs. 1 and 2). Some authors have correlated these fossil-rich marine beds with the
92
Lontras shale (Soares, 1991; Castro, 1999) at the top of the Campo Mourão Formation (França
93
and Potter, 1991). On the other hand, Loczy (1964) and Rocha-Campos (1969) correlated the
94
Capivari fossil beds with the marine assemblages of the Teixeira Soares marine beds (Passinho
95
shale and associated rocks), uppermost part of the Itararé Group, State of Paraná, southern Brazil.
96
Alternatively, Taboada et al. (2016) positioned the Capivari fossil-rich beds in between the
97
Lontras and Passinho fossil-bearing shales of the southern Brazil (Figs. 1, 2).
98
The precise taxonomic composition, and age of the fossil-bearing beds of the Capivari
99
region (see Fig. 2) can shed light on intrabasinal biocorrelation of the Late Paleozoic marine
100
invertebrates faunas of the Paraná Basin with stratigraphic and paleogeographic implications for
101
the depositional history of the glacial succession of the Itararé Group. Therefore, in this
102
contribution we will attempt to better constrain those fossil-bearing beds. To achieve this, we
4
103
reassessed the invertebrate fauna originally described by Mendes (1952), also complementing
104
our analysis with new fossil findings.
105 106
1.1. Stratigraphic framework
107
During the Paleozoic, the intraplate Paraná Basin was a large depositional area in the SW
108
Gondwanaland (central/southeastern part of the South America) (Milani, 1997). The basin was
109
elongated in the N-S direction and filled with Late Ordovician to Late Cretaceous sedimentary
110
rocks, encompassing six supersequences or unconformity-bounded units (Milani, 1997). The
111
2,500-m-thick succession of upper Paleozoic to Mesozoic rocks is referred as the Gondwana I
112
Supersequence (Milani, 1997; Milani et al., 2007) that was generated in a variety of continental,
113
transitional and marine sedimentary settings, including upper Carboniferous-Permian glacial to
114
Triassic arid depositional environments.
115
Rocks of the glacial succession are included in the Itararé Group (eastern border), which is
116
one of the most complete stratigraphic successions of the LPIA in southwestern Gondwana.
117
Resting either directly upon Precambrian basement, lower Paleozoic or on Devonian beds, the
118
Itararé Group is the thickest sedimentary package of the Paraná Basin, sedimentation lasting for
119
at least 36 million years (França and Potter, 1991). Conglomerates, diamictites, sandstones,
120
mudstones, rhythmites, turbidites, and thin coal beds are the main lithotypes of the group
121
(Rocha-Campos, 1967; França and Potter, 1991; Holz et al., 2010). The precise age of the rocks
122
of the Itararé Group is one of the main issues of the Gondwana geology in Brazil. The Itararé
123
Group is usually referred as upper Mississippian-lower Cisuralian in age (Souza and Marques-
124
Toigo, 2005; Guerra-Sommer et al., 2008a, b; Rocha-Campos et al., 2008; Holz et al., 2010;
125
Neves et al., 2014b; Taboada et al., 2016). However, U–Pb radiometric ages have dated the
126
topmost glacial deposits of the Itararé Group at 307.7 ± 3.1 Ma (Kasimovian−Moscovian,
127
Pennsylvanian, late Carboniferous) and the base of the overlying post-glacial deposits of the 5
128
coal-bearing Rio Bonito Formation at 298.8 ± 1.9 Ma (Ghzelian−Asselian, Carboniferous–
129
Permian boundary) (Cagliari et al., 2014; 2016). Indeed, a radiometric Carboniferous age for the
130
upper Itararé Group was reinforced by newely U–Pb laser ablation ICP-MS detrital zircon dates
131
of rocks of this unit (Tedesco et al., 2019). However, we are following the chronostratigraphic
132
chart of Holz et al. (2010), which present the most complete information of age for the entire
133
Itararé Group (Fig. 1).
134
Since the seminal work by Barbosa and Almeida (1949a), various lithostratigraphic
135
subdivisions for the Itararé Group appeared in the literature. In southeastern Brazil, the group
136
was subdivided into several formations, which include, in ascending order, the Itu, Capivari,
137
Gramadinho, Tietê and Itapetininga formations (Barbosa and Almeida, 1949a). However, the
138
original designations of these authors are now in disuse. Schneider et al. (1974) proposed an
139
alternate litostratigraphic framework for outcrops in southern Brazil, in which the Itararé
140
comprises three main units, including from the base to top, the Campo do Tenente, Mafra, and
141
Rio do Sul formations.
142
Based on a detailed subsurface analysis, França and Potter (1991) recognized three fining-
143
upward glacially-influenced sedimentary cycles and classified them as formations named, from
144
base to top, as Lagoa Azul, Campo Mourão and Taciba. These authors also noted that in the
145
subsurface the maximum thickness (~1300 m) of the Itararé Group is reached in the State of São
146
Paulo, where our study area is located (Fig. 1).
147
Trying to correlate the glacial cycles defined by França and Potter (1991) in outcrop area
148
of the Itararé Group, Vesely and Assine (2006) recognized five glacially-influenced depositional
149
sequences in the State of Paraná. These are also laterally traceable for nearly 400 km in well logs
150
across the central portion of the basin. The oldest depositional sequence was correlated with the
151
Lagoa Azul Formation of França and Potter (1991), the youngest with the Taciba Formation, and
152
the three middle sequences with the whole Campo Mourão Formation. A retrogradational 6
153
stratigraphic stacking pattern characterizes all these cycles, which correspond to major ice-retreat
154
phases or deglaciation sequences (Vesely and Assine, 2006). Complex facies associations are
155
generated during deglaciation, mainly because tillites and outwash facies formed during an
156
oscillatory but overall glacier-retreat and are interfingered with contemporaneous glacial-marine
157
deposits.
158
In the outcrops of the studied area, the Taciba Formation is equivalent to the Capivari
159
Formation (mainly fine-grained facies) and the overlying Gramadinho Formation (mainly
160
diamictites) of Barbosa and Almeida (1949a, b). Yet, the Taciba Formation is the youngest and
161
most widespread unit of the Itararé Group, corresponding to the Rio do Sul Formation of
162
Schneider et al. (1974). Finally, as commented above, the Taciba Formation is the sedimentary
163
record of the last deglacial sequence (sequence 5 of Vesely and Assine, 2006), representing the
164
final ice-retreat in the basin, onlapping the south border of the basin.
165
Based on subsurface data from three boreholes (C-IG/93, R-IG/94 and R-IG/95) drilled in
166
the Capivari and Rafard counties, Petri et al. (1996) subdivided the upper part of the Itararé
167
Group into four main sedimentary packages, as follows (from the base to the top): (a) package D,
168
represented by conglomerates and coarse sandstones; (b) package C that includes mudstones,
169
siltstones and rhythmites; (c) package B, which is dominated by sandstones, and (d) package A,
170
represented by transgressive mudstones. This succession is, in general, similar to that recorded
171
by Saad (1977) to the Itararé Group in the south and central portions of the State of São Paulo.
172 173
1.2. Background: the Capivari fossil bed unearthed
174
The marine macroinvertebrate assemblage herein described comes from a ~2-m-thick interval of
175
silty shales recorded in the middle part of the Taciba Formation, upper part of the Itararé Group
176
in the State of São Paulo, formerly refered to the Capivari Formation (Figs. 1 and 2).
177
Theassemblage is thought to be the most diverse one of the Itararé Group (Rocha-Campos and 7
178
Rösler, 1978; Simões et al., 1998), and is 500 km far away of those invertebrate-rich beds of the
179
Taciba Formation (Passinho Shale) from southern Brazil (Neves et al., 2014b; Taboada et al.,
180
2016) (Fig. 2). As described and commented below, the fossil-bearing silty shale bed crops out
181
in the vicinity of the Capivari county (Fig. 2). Fossils from these localities were first discovered
182
in 1947 by F.F.M. Almeida, S. Petri and O. Barbosa (see information in Mendes, 1952), but the
183
first publication on these fossil-bearing sites appeared in the literature only two years later
184
(Barbosa and Almeida 1949a). It was only in March of 1949, in a short note to the Brazilian
185
Academy of Sciences, that Barbosa and Almeida (1949a) briefly discussed the stratigraphic
186
position of the Capivari marine fauna within the Itararé Group. In August of that year, a more
187
comprehensive and detailed study of the geology and stratigraphy of the area was published. By
188
combining both subsurface and surface data, Barbosa and Almeida (1949b) constrained the
189
marine assemblage to the Capivari Formation and noted that the fossil bed is intercalated
190
between rhythmites and diamictites. The authors recorded the presence of brachiopods
191
(Ambocoelia, Rhynchopora), bivalves (Aviculopecten), and gastropods (that were subsequently
192
lost, see Rocha-Campos, 1966, p. 10), but the fossils were neither described, nor illustrated.
193
Despite this, Barbosa and Almeida (1949b) noted that the fossils are not identical to those
194
already described from the fossil-rich beds of the Itararé Group (i.e., Passinho shale and coeval
195
strata of the Teixeira Soares region, State of Paraná) from the State of Paraná, southern Brazil.
196
J.C. Mendes formally described the marine macroinvertebrate fossils of the “Capivari beds”
197
in 1952, basing on those specimens previously collected by Barbosa and Almeida (1949a, b),
198
plus additional material offered by the geologist S. Mezzalira. He reported the presence of
199
rhynchonelliform brachiopods (Crurithyris aff. planoconvexa, Rhynchopora grossopunctata)
200
and bivalves (Aviculopecten capivariensis n. sp., Nuculana? sp., and undetermined specimens)
201
(Mendes, 1952). Subsequently, Rocha-Campos (1966) recorded the presence of the gastropod
202
Peruvispira delicata. 8
203
After the contributions above, the marine fauna of the Capivari beds was almost forgotten
204
for over twenty years, mainly because the outcrops of the Itararé Group in the region are deeply
205
weathered (see also Mendes, 1952) and/or covered by dense commercial plantations (e.g., sugar
206
cane). During this period, data about the Capivari marine fauna appeared only in review papers,
207
as those by Rocha-Campos and Röser (1978) and Petri and Souza (1993). In both contributions,
208
the stratigraphic position and biocorrelation of the Capivari marine invertebrate fauna with those
209
of the southern parts of Brazil were tentatively constrained and discussed. In this context, a
210
probable northward extension of those beds was mentioned by Rocha-Campos and Rösler (1978,
211
p. 5). Indeed, these authors reported the presence of similar fossil-bearing rocks near the city of
212
Hortolândia, State of São Paulo, including the following bivalve genera: Phestia, Nuculopsis,
213
and Edmondia, plus undetermined pholadomyid shells.
214
In the mid-1990s, faculty of the Institute of Geosciences, University of São Paulo
215
(IGc/USP) carried out various field trips to the area originally mapped by Barbosa and Almeida
216
(1949b), “rediscovering” the fossil-bearing beds of Capivari (A.C. Rocha-Campos personal
217
communication, 1996). Consequently, new sampling efforts were carried-out, significantly
218
contributing to the find of new specimens. This enlarged substantially the original fossil
219
collection made by Barbosa and Almeida (1949a, b) and Mendes (1952). Unfortunately, at the
220
end of the 1990s, the original exposure of the Capivari outcrop was almost destroyed (Anelli et
221
al., 2000, p. 151) during the road works to enlarge the original path of the old, unpaved route
222
between the cities of Capivari and Salto (see Fig. 2). Despite that, renewed interest on those
223
fossils reappeared during the extensive study of the coeval late Paleozoic marine invertebrate
224
faunas of the Itararé Group from southern Brazil (Simões et al., 2012; Neves et al., 2014b;
225
Taboada et al., 2016). These studies better constrained the stratigraphic position, age and
226
biocorrelation of the marine assemblages of the Taciba Formation, upper part of the Itararé
9
227
Group, with other well-dated Gondwanan faunas (Simões et al., 2012; Neves et al., 2014b;
228
Taboada et al., 2016).
229
Based on the literature (e.g. Vesely and Assine, 2006) and personal communications from
230
various paleontologists and stratigraphers (A.C. Rocha-Campos, S. Petri), we relocated what was
231
left of the fossil-bearing beds in the Capivari-Salto road. At the same time, significant efforts
232
were made to gather the original material collected by previous authors, such as O. Barbosa,
233
F.F.M. Almeida, S. Petri, J.C. Mendes, and A.C. Rocha-Campos. Hence, in this contribution we
234
describe the fossil-bearing sedimentary succession of the Itararé Group near Capivari, and revise
235
the marine assemblage based on new material as well as the original fossils studied by previous
236
authors.
237 238
2. Material and methods
239
In total, 150 specimens of macroinvertebrates of the Capivari marine beds were examined, as
240
follows: 115 brachiopods, seven bivalves, three gastropods, and 25 crinoid fragments (Table 4).
241
All analyzed specimens (Tables 5-7) come from the classical outcrop located at ~6 km south of
242
the Capivari, on the old road between the Capivari and Salto towns (Figs. 2, 3).
243
In the laboratory, the fossil material was prepared according to standard paleontological
244
procedures (see Feldmann et al., 1989) using precision tools. Latex casts and plasticine molds
245
were also prepared for most of the specimens. Shells were coated with magnesium oxide
246
sublimate to enhance diagnostic morphological characters for photography (see Neves et al.,
247
2014a, b, and references therein). The ecological guilds (i.e., inferred invertebrate lifestyles)
248
were determined based on Aberhan and Kiessling (2015, p. 2).
249 250
Repositories and institutional abbreviations
10
251
All studied specimens are housed in the Institute de Geosciences, University of São Paulo, São
252
Paulo city, State of São Paulo. The following abbreviations are used in the text for institutional
253
acronyms: GP-1T, this prefix is reserved for type specimens belonging to the Invertebrate
254
Collection deposited in the Geosciences Institute, São Paulo University, Brazil; GP-1E, this
255
prefix is used for the invertebrate specimens that are housed in the non-type collection of that
256
institution; DGM, this prefix indicates that the specimens belong to the fossil collection of the
257
Geology and Mineralogy Division, Nacional Department of Mineral Production, State of Rio de
258
Janeiro, Brazil; IGG, this prefix is used for the material reposited in the Geographical and
259
Geological Institute of the State of São Paulo, now Geological Institute, Secretariat of the
260
Environment, São Paulo State, Brazil; CP.I, this prefix refers to the CENPALEO fossil
261
collection, Contestado University, Mafra, State of Santa Catarina, southern Brazil.
262 263
3. Systematic paleontology
264
The applied classification for Brachiopoda largely follows Waterhouse (2013) for Suborder
265
Linproductidina, Savage et al. (2002) for Order Rhynchonellida, Carter et al. (2006) for
266
Suborder Spiriferidina, Racheboeuf (2000) for Suborder Chonetidina. Taxonomic analysis of
267
Bivalvia follows Waterhouse, (2008, 2010), Bieler et al. (2010), Carter et al. (2011) for
268
suprageneric categories, and Dickins (1957; 1963), Morris et al. (1991), Newell and Boyd (1995)
269
and Neves et al. (2014b) at the generic and species level. Finally, the name of crinoid columnals
270
and pluricolumnals are preceded by the prefix ‘‘ø’’, as suggested by Le Menn (1987, 1988) and
271
Scheffler et al. (2015).
272 273
Class Strophomenata Williams et al., 1996
274
Order Productida Sarytcheva and Sokolskaja, 1959
275
Suborder Linoproductidina Waterhouse, 2013 11
276
Superfamily Proboscidelloidea Muir-Wood and Cooper, 1960
277
Family Auriculispinidae Waterhouse, 1986
278
Subfamily Lyoniinae Waterhouse, 2001
279
Tribe Lyoniini Waterhouse, 2001
280 281
Genus Lyonia Archbold, 1983
282 283
Type species: Linoproductus (Cancrinella) cancriniformis var. lyoni Prendergast 1943, from the
284
Lyons Group (lower Permian), Carnarvon Basin, Western Australia.
285 286
Remarks: A set of finely ribbed, spinose linoproductids such as Cancrinella Fredericks, 1928,
287
Auriculispina Waterhouse,1975, Costatumulus Waterhouse, 1983a, and Magniplicatina
288
Waterhouse, 1983b are externally closely similar to Lyona. Nevertheless, when compared to
289
Lyonia, Cancrinella possesses a more inflated shell, smaller size, crowded spines on the ears and
290
abundant spines on the dorsal valve (see discussion of this last character in Grigorjeva et al.,
291
1977; Waterhouse, 1982a; Shen et al., 2000; Li et al., 2012). Auriculispina is distinguished from
292
Lyonia by the presence of two or three rows of spines on the ears, stronger rugae on both valves,
293
more weakly swollen ventral spine ridges and a lack of dorsal spines (Waterhouse, 1975;
294
Archbold, 1983). Likewise, Costatumulus could be confused to Lyonia but the former has a more
295
inflated concave-convex profile, a lack of dorsal spines and possesses double rows of spines on
296
the ears (Waterhouse, 1983a, 1986). Magniplicatina also has ventral spines in one ou more hinge
297
rows and lacks dorsal spines but exhibits strongly developed concentric wrinkles or rugae as well
298
as coarser ventral spines on elongate spine ridges (Waterhouse, 1983b; Li et al., 2012), unlike
299
Lyonia. Closest genera to Lyonia such as Nambucalinus Waterhouse, 2001 and Bandoproductus
300
Jin and Sun, 1981, were discussed in a previously published paper (Taboada et al., 2016). 12
301 302
Lyonia rochacamposi Taboada et al., 2016
303
Figure 4.29–4.32
304 305
1969 Cancrinella sp.; Rocha-Campos, p. 105, pl. X, figs. 1-5, 8.
306
1969 Linoproductus sp.; Rocha-Campos, p. 106, pl. X, figs. 6-7.
307
2016 Lyonia rochacamposi; Taboada et al., p. 443, figs. A-O.
308 309
Holotype: CP.I 769I, CENPALEO (Universidade do Contestado), from the Taciba Formation,
310
Itararé Group, southern Paraná Basin, Brazil.
311 312
Occurrence: Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
313
Basin, Brazil. Latest Asselian–earliest Sakmarian.
314 315
Description: Lyonia with transverse outline and moderately convex ventral valve. Ventral valve
316
maximum width close to hinge line, up to 30 mm wide, 24 mm long, and width/length ratio 1.25.
317
Ventral valve with large flattened ears (partially broken) and ornamentation of fine radial
318
costellae (8-9 per 5 mm on the venter); weak rugae on venter slightly stronger on ears. Costellae
319
number increasing by splitting into two ribs anteriorly from slightly swollen minute spine bases
320
in roughly quincuncial arrangement. A single row of spines, coarser than those on the venter
321
appears to be on hinge line, although this character is not well preserved.
322 323
Materials: Two fragmentary specimens, one external mold of dorsal valve, GP/1E 4350a; one
324
ventral valve, GP/1E 4350b.
325 13
326
Measurements: See Table 5 for Lyonia rochacamposi shell measurements.
327 328
Remarks: The studied specimens are scarce and limited to fragmentary remains of ventral
329
valves but exhibit comparable size, moderate convex profile, transverse outline, large flattened
330
ears, single row of hinge spines, distinct fine costellae and ventral spines quincunxial
331
arrangement that suggest its assignment to Lyonia rochacamposi Taboada et al., 2016. This
332
species was recently described from the post-glacial section of the Taciba Formation in the
333
Brazilian Santa Catarina State. The Capivari’s L. rochacamposi specimens, as well as those
334
from the southern Taciba Formation, are distinguished from L. lyoni from Lyons Group of
335
Western Australia mainly by their more transverse outline and finer costellation.
336
Cancrinelloides monticulus Waterhouse, 1982b, from the Asselian Pucket Group of south
337
Thailand was considered moderately close to Lyonia lyoni, but the Australian species is smaller,
338
with more emphasized concentric ornament, finer radial ornament, and slightly shorter spine
339
bases (Waterhouse, 1982b). Likewise, Lyonia rochacamposi is distinguished from the Thailand
340
species by its more transverse outline, finer costellation and presence of dorsal spines anteriorly.
341
Cancrinelloides monticulus was assigned latter to Bandoproductus (although with subgeneric
342
hierarchy) by Waterhouse (1982b) and shortly after to the same genus by Archbold (1983).
343 344
Order Rhynchonellida Kuhn, 1949
345
Superfamily Rhynchoporoidea Muir-Wood, 1955
346
Family Rhynchoporidae Muir-Wood, 1955
347
Subfamily Rhynchophoriinae Muir-Wood, 1955
348 349
Genus Rhynchopora King, 1865
350 14
351
Type species: Terebratula geinitziana de Verneuil, 1845, from the upper Permian (Kazanian) of
352
Arkhangelsk Region, Russia.
353 354
Rhynchopora grossopunctata Mendes, 1952
355
Figure 4.1–4.15
356 357
1952 Rhynchopora grossopunctata; Mendes, p. 12, figs. 4-7.
358
1966 Rhynchopora grossopunctata; Mendes, Mezzalira, pl. 2, figs. 3-4 (copy Mendes, 1952,
359
figs. 4-5).
360
1967, Rhynchopora grossopunctata; Mendes, Rocha-Campos, pl. 31, figs. 1-3 (copy Mendes,
361
1952 figs. 4-5, 7).
362
1989 Rhynchopora grossopunctata; Mendes, Mezzalira, pl. 2, figs. 3-4 (copy Mendes, 1952, figs.
363
4-5).
364 365
Holotype: Mendes (1952) did not choose a holotype but illustrated the syntype series of
366
Rhynchopora grossopunctata with three specimens labeled DGM 4189-4190 and IGG 518-I,
367
respectively. The IGG repository (Instituto Geográfico e Geológico do Estado de São Paulo) is
368
no longer available, while the specimens under the DGM acronym (Divisão de Geologia e
369
Mineralogia, Departamento Nacional da Produção Mineral, Estado de Rio de Janeiro) were
370
transferred to the fossil collection of the Instituto de Geociências da Universidade de São Paulo,
371
although they are currently lost. An external mold of conjugated valves, labeled GP/1E-4342,
372
housed in the Instituto de Geociências da Universidade de São Paulo, is here selected as neotype
373
of R. grossopunctata Mendes.
374
15
375
Diagnosis: Large, subpentagonal in outline with maximum width mid-length. Five costae on
376
sulcus and fold, eight to nine on lateral flanks. Interspaces anteriorly develop coarse projections
377
extending as short tongues. Five punctae per mm2. Dental plates diverge 30º, up to one fourth of
378
valve length; dorsal interior with a thin median septum one third of valve length.
379 380
Occurrence: Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
381
Basin, Brazil. Latest Asselian–earliest Sakmarian.
382 383
Description: Large-sized Rhynchopora, subpentagonal in outline with maximum width up to 18
384
mm at mid-length and length up 16 mm, with an average W/L ratio of 1.15. Ventral valve gently
385
convex with maximum convexity at umbo and dorsal valve more inflated with slightly raised
386
fold and maximum convexity at two thirds of valve length. Umbonal angle of 95º-100º; cardinal
387
extremities rounded. Sulcus very shallow, barely noticeable anteriorly from first third of valve
388
length and widening anteriorly (~40º) up to one fourth of maximum width. Five costae on sulcus
389
and fold, and 8-9 on lateral flanks, with finer costae developed postero-laterally. Costae wider at
390
commissure. Interspaces anteriorly develop long coarse projections extending as short tongues.
391
Five punctae per mm2. Dental plates diverge 30º, up to one fourth of valve length; dorsal interior
392
with a distinct thin median septum extending one third of valve length. Others features unknown.
393 394
Materials: One external mold of conjugated valves is the neotype, GP/1E-4342. Twenty
395
topotypic specimens. Two ventral valve external molds, GP/1E-443a, 443b; two ventral valve
396
internal molds GP/1E-2507, 2545; three dorsal valve external molds, GP/1E-451, 1915, 4405;
397
three dorsal valve internal molds, GP/1E-1912, 2492, 4325. Other fragmentary specimens,
398
GP/1E-445b, 446-447, 460, 1913, 2502, 2524, 2536, 4339.
399 16
400
Measurements: See Table 5 for Rhynchopora grossopunctata shell measurements.
401 402
Remarks: Although not preserved in the material here described, a spondylium is clearly present
403
in the syntype series of R. grossopunctata (Mendes, 1952). Species of Rhynchopora reported
404
from South America include Kozlowski (1914) R. nikitini (not Tschernyschew, see Cooper and
405
Grant, 1976) from the Copacabana Formation (Early Permian) of Bolivia (later synonymysed
406
under R. aff. illinoisensis [Worthen] by Chronic in Newell et al., 1953), and R. amotapensis
407
Chronic (in Newell et al., 1953), the latter two from the Tarma Group (Late Carboniferous) of
408
Peru. The last species shares a slightly transverse subpentagonal outline with R. grossopunctata,
409
as well as shallow sulcus and flatten fold with a similar number of costae, but the Brazilian
410
species is larger and exhibits more costae on the lateral flanks. R. aff. illinoisensis is smaller and
411
possesses more costae on sulcus, fold and lateral flanks when compared to R. grossopunctata.
412
Another South American species is Rhynchopora sp. from the Río del Peñón Formation
413
(Moscovian) of central-western Argentina (Cisterna and Simanauskas, 2000). It differs from R.
414
grossopunctata by having a smaller size, rounded subtriangular outline, deeper sulcus, fewer
415
costae on flanks and longer dorsal median septum. Rhynchopora australassica Archbold, 1995
416
(see also Archbold and Hogeboom, 2000), from the Lyons Group (Early Permian) of Western
417
Australia, has a larger size, subtriangular outline, fewer costae on flanks, deeper sulcus and a
418
more raised fold when compared to R. grossopunctata. Rhynchopora culta Waterhouse, 1982b,
419
from south Thailand and western Peninsular Malaysia (Shi and Waterhouse, 1991) is comparable
420
to R. grossopunctata, although the former has a subtrigonal shell shape, reduced number of
421
costae on flanks and almost double the number of punctae per millimeter.
422 423
Order Spiriferida Waagen, 1883
424
Superfamily Ambocoeloioidea George, 1931 17
425
Family Ambocoeliidae George, 1931
426
Subfamily Ambocoeliinae George, 1931
427
Genus Biconvexiella Waterhouse, 1983a
428 429
Type species: Attenuatella convexa Armstrong, 1968 from the Tiverton Formation, (Early
430
Permian), Bowen Basin, Queensland, Australia.
431 432
Remarks: Biconvenxiella shares with Attenuatella Stehli, 1954 and Crurithyris George, 1931,
433
comparable shell shape and a similar external spinose microornamentation. Biconvexiella
434
possesses a subcircular shell outline, slightly incurved umbo, slightly concave to gently convex
435
dorsal valve and two pairs of adductor scars, the posterior pair scars being smaller than anterior
436
pair (Shi and Waterhouse, 1996; He et al., 2007). Attenuatella is distinguished from
437
Biconvenxiella by the presence in the former of an attenuated outline, a stronger convex ventral
438
valve and strongly incurved umbo, as well as a different arrangement and proportions of muscle
439
scars (Stehli, 1954; Waterhouse, 1964; Landis and Waterhouse, 1966; Shi and Waterhouse, 1996:
440
He et al., 2007, 2012). Crurithyris is characterized by a subrounded to transversely ovate shell
441
outline, a moderately to strongly incurved ventral umbo, a pair of centrally depressed ventral
442
adductor scars bisected by a thin ridge (may be absent), a pair of laterally depressed diductor
443
scars, which are separated from their adjacent adductor scars by ridges, and a pair of dorsal
444
adductor scars bisected by a median ridge (George, 1931; Stehli, 1954; He et al., 2007, 2012).
445 446
Biconvexiella saopauloensis n. sp.
447
Figure 4.16–4.25
448 449
1952 Crurithyris aff. planoconvexa (Shumard); Mendes, p. 10-11, pl. I, figs. 1-3. 18
450 451 452 453
1966 Crurithyris aff. planoconvexa (Shumard); Mezzalira, pl. 2, figs. 1-2 (copy Mendes, 1952 pl. I, figs. 1-2). 1967 Crurithyris aff. planoconvexa (Shumard); Rocha-Campos, pl. XXXI, figs. 4-5 (copy Mendes, 1952, pl. I, figs. 2-3).
454
1969 Attenuatela paulistana; Rocha-Campos, p. 108-112, text-fig. 12, pl. XI, figs. 1-14.
455
1989 Crurithyris aff. planoconvexa (Shumard); Mezzalira, pl. II, figs. 1-2 (copy Mendes,
456 457
1952 pl. I, figs. 1-2). 2016 Biconvexiella spp.; Taboada et al., pl. 4, fig. J.
458 459
Holotype: GP/1E 4339; Paratypes: GP/1E 450, 453, 2495, 2513, 2520, 2534, 4322, 4327, 4328,
460
4356c, all from the Taciba Formation, Paraná Basin, State of São Paulo, southeastern Brazil.
461 462
Diagnosis: Average-sized Biconvexiella of biconvex profile with slightly convex dorsal valve
463
and strongly convex ventral valve, and subcircular outline. Ventral valve subequidimensional;
464
dorsal valve variably transverse; hinge line shorter than maximum width. Ventral median sinus
465
shallow and narrow. Dorsal valve convex with a narrow slightly raised fold.
466 467
Occurrence: Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
468
Basin, Brazil. Latest Asselian–earliest Sakmarian.
469 470
Description: Average-sized Biconvexiella of biconvex profile and subcircular outline. Ventral
471
valve subequidimensional; dorsal valve variably transverse; hinge line shorter than maximum
472
width; which is located at valve midlength. Maximum width up to 14 mm, maximum length up
473
to 11.5 mm with a dorsal W/L ratio varying between 1.23-1.43 and W/L average of 1.33. Ventral
474
valve strongly convex with its stronger convexity in the umbonal region. Ventral median sinus 19
475
shallow and narrow. Dorsal valve slightly convex with a narrow fold not always developed.
476
Ornamentation of fine concentric growth lines sometimes lamellose and subconcentric rows of
477
minute subcircular spine bases (9-10/mm at valve mid lenght). Interior of ventral valve with long
478
(3/4 ventral valve length), narrow median adductor ridge bounded by two narrow elongate
479
depressed adductor scars and slightly rised semielliptical diductor scars. Interior of dorsal valve
480
(holotype) with small subelliptical (0.3 mm width, 1 mm length) depressed posterior adductor
481
scars separated by a short thin median ridge. Anterior adductor scars barely impressed, elongate
482
(2 mm length) tongue-shaped (1 mm of maximum width anteriorly) with inner side curved.
483
Crural plates short, divergent (30º-50º); dental sockets deep; cardinal process tuberculate.
484 485
Etymology: From the State of São Paulo, southeastern Brazil.
486 487
Materials: Eighty-six specimens. Three external molds of conjugated valves: GP/1E 2495, 2513,
488
4328; two exterior of dorsal valves: GP/1E 2534, 4327; five dorsal valve external molds: GP/1E
489
448, 449, 2413, 4323a, 4354d1; fifteen ventral valve external molds; GP/1E 2503, 2504, 2522,
490
2526, 2544, 2552, 4318a, 4322, 4323b, 4324, 4326, 4332, 4341, 4354d2; eleven dorsal valve
491
internal molds: GP/1E 450, 453, 2493, 2498, 2536-2538, 2543, 4339, 4351; twelve ventral valve
492
internal molds: GP/1E 1910, 2489, 2520, 2525, 2527, 2528, 2551, 4354b, 4355b, 4356a, 4356b,
493
4338; other fragmentary material: GP/1E 1906, 1907, 1911, 1914, 2496, 2497, 2499, 2500, 2505,
494
2506, 2508, 2510, 2514-2517, 2521, 2524, 2529, 2530, 2532, 2533, 2535, 2539, 2541, 2546-
495
2550, 2553, 2555, 4320, 4330, 4331, 4334, 4340, 4357.
496 497
Measurements: See Table 5 for Biconvexiella saopauloensis n. sp. shell measurements.
498
20
499
Remarks: Ambocoelids from the Taciba Formation were first mentioned by Barbosa and
500
Almeida (1949a, b), later briefly described as Crurithyris aff. planoconvexa (Shumard) by
501
Mendes (1952) but interpreted as belonging to a new species (named paulistana) of
502
Attenuatella Stehli, 1954, by Rocha-Campos (1969). Attenuatella paulistana was never
503
published, failing to satisfy article 13 of the ICZN, therefore being a nomen nudum. The
504
Capivari’s ambocoelids exhibit shell shape, slightly incurved umbo, strong ventral median
505
ridge supporting muscle scars, dorsal thin median ridge separating posterior adductors scars and
506
larger anterior adductor scars, that collectively support assingnment to Biconvexiella, as was
507
recently indicated (Taboada et al., 2016).
508
Biconvexiella saopauloensis n. sp. is comparable to Attenuatella convexa Armstrong,
509
1968, type species of Biconvexiella Waterhouse, 1983a, which occurs in the Tiverton
510
Formation (Sakmarian), Bowen Basin, Queensland, and also present in the Farley Formation,
511
Sydney Basin, New South Wales, Australia (Armstrong and Telford, 1970). It shares with B.
512
saopauloensis n. sp. a biconvex profile, ventral sinus and dorsal fold, as well as comparable
513
internal muscle scars. Nevertheless, B. saopauloensis n. sp. has a more transverse outline, less
514
pronounced ventral valve sinus and shallower dorsal valve fold lacking lateral bounding ridges.
515
B. saopauloensis n. sp. is close to B. roxoi (Oliveira, 1936) (see also Taboada et al., 2016) from
516
the upper part of the Taciba Formation, Paraná Basin, southern Brazil, although the Capivari’
517
species is more transverse with a convex dorsal valve, a stronger and longer ventral median
518
septum and more divergent crural plates. Ogilviecoelia inflata Shi and Waterhouse, 1996, from
519
the Early Permian Jungle Creek Formation, northern Yukon Territory, Canada, resembles B.
520
saopaulensis n. sp. in shell outline and dorsal valve profile, although the Canadian material
521
possesses a micro-ornamentation of fine elongate grooves instead spinose ornament, and a large,
522
depressed and well differentiated ventral muscle field and undifferentiated dorsal adductors,
523
unlike the Brazilian species. 21
524 525
Order Chonetida Nalivkin, 1979
526
Suborder Chonetidina Muir-Wood, 1955
527
Superfamily Chonetoidea Bronn, 1862
528
Family Rugosochonetidae Muir-Wood, 1962
529
Subfamily Quinquenellinae Archbold, 1981
530 531
Genus Quinquenella Waterhouse, 1975
532 533
Type species: Quinquenella glabra Waterhouse, 1975 from the Nambdo Member of the Senja
534
Formation (late Permian), east Jumbla city, north-west Nepal.
535 536
Remarks: Both Tornquistia Paeckelmann, 1930 and Quinquenella are smooth externally and
537
have similar dorsal interiors. Nevertheless, the former is smaller, has a hump-shaped strong
538
convave-convex profile and stout accessory septa in the dorsal valve, unlike the low profile and
539
weaker accessory septa of Quinquenella.
540 541
Quinquenella rionegrensis (Oliveira, 1930)
542
Figure 4.26–4.28
543 544
1930 Chonetes rionegrensis; Oliveira, p. 19, unnumbered figure.
545
2016 Quinquenella rionegrensis (Oliveira, 1930); Taboada et al., 2016, p. 449-450, pl. III,
546
figs. D-L.
547
22
548
Holotype: The holotype of Quinquenella rionegrensis (Oliveira, 1930) is lost (see Taboada et
549
al., 2016, p. 449, 450). Lectotype GP/1E 4339b is from the Taciba Formation, Itararé Group,
550
Paraná Basin, Teixeira Soares region, southern Brazil.
551 552
Occurrence: Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
553
Basin, Brazil. Latest Asselian–earliest Sakmarian.
554 555
Materials: Seven specimens. Two interior molds of dorsal valves, GP/1E-4354c, GP/1E 4358;
556
one exterior mold of dorsal valve, GP/1E 4355a. Other fragmentary material, GP/1E 449, 456,
557
1909, 2523.
558 559
Measurements: See Table 5 for Quinquenella rionegrensis shell measurements.
560 561
Remarks: Available dorsal valves exhibit moderate concavity, subelliptical transverse outline
562
and maximum width at hinge line. Maximum width up 9 mm; maximum length up 5.5 mm with
563
an W/L ratio varying between 1.55-1.76 and W/L average of 1.65. Interior with lines of pustules
564
radially aligned, short lateral septa, thin long accessory septa, ill-defined median septum fused
565
posteriorly surrounding a small, circular shallow alveolus anterior to a small bifid cardinal
566
process. Despite scarcity of material, internal characters and dimensions of dorsal valve fit finely
567
with those observed in Quinquenella rionegrensis (Oliveira, 1930) (see also Taboada et al., 2016)
568
from the upper part of the Taciba Formation, Paraná Basin, southern Brazil. Finally, the
569
Brazilian species was compared to Quinquenella species from Western Australia (see Archbold
570
1981) by Taboada et al. (2016).
571 572
Class Bivalvia Linnaeus, 1758 23
573
Subclass Protobranchia Pelseneer, 1889
574
Order Nuculanida Carter, Campbell and Campbell, 2000
575
Superfamily Nuculanoidea Adams and Adams, 1858 (Gray, 1854)
576
Family Polidevciidae Kumpera, Prantl and Ruzicka, 1960
577 578
Genus Phestia Chernyshev, 1951
579 580
Type species: Leda inflatiformis Chernyshev, 1939 from the Carboniferous of
581
Russia, by original designation.
582 583
Remarks: González (2010) proposed the genus Waterhouseus to include the species Phestia
584
tepuelensis, but the generic diagnosis is not precise. Besides, the differences of Phestia and
585
other closely related genera, such as Polidevcia are not clear in the discussion. Thus, we decide
586
to keep the original name for the Patagonian species (Argentina). A revision of these genera
587
may be necessary but exceeds the scope of this contribution and is an objective of a future
588
research.
589 590
Phestia tepuelensis Gonzalez, 1969
591
Figure 5.1–5.4
592 593
Holotype: MLP 11021 from the Sierra de Languiñeo, Tepuel-Genoa Basin, Chubut, Argentina
594
(Gonzalez, 1969; Pagani, 2004).
595 596
Occurrence: Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
597
Basin, Brazil. Latest Asselian–earliest Sakmarian. 24
598 599
Description: Shell very inequilateral, tapering posteriorly with a developed rostrum. Umbones
600
slightly opisthogyrous, small, very depressed, positioned at anterior fourth of shell length. The
601
greatest height of the shell occurs at umbones. Anterior margin rounded forming an obtuse angle
602
with a convex ventral margin; posterior-dorsal margin slightly concave. Surface ornamentation
603
of thick commarginal costae, which are crossed by radial costae; with a pronounced
604
discontinuity in the coarseness of the commarginal ribs from the anterior lobe to the lateral flank.
605
Escutcheon, hinge features and muscle scars not observed.
606 607
Materials: One internal mold of right valve (GP/1E 455) and one external mold of left valve
608
(GP/1E 452).
609 610
Measurements: See Table 6 for Phestia tepuelensis shell measurements.
611 612
Remarks: Phestia tepuelensis was firstly described for the Tepuel-Genoa Basin, Argentina
613
(González, 1969), and revised by Pagani (2004). This species shows strong concentric costae
614
crossed by fine radial ribs, a feature that distinguishes it from seven Argentinean species. Three
615
of them come from Northwestern Argentina: Phestia sp. from San Juan Province (Taboada,
616
1989); Phestia sp. from Río Blanco Basin (González, 1994), both Carboniferous, and Phestia aff.
617
P. bellistriata (Stevens) from Permian deposits of the Tupe Formation (Sterren, 2004). The other
618
four species come from the Tepuel-Genoa Basin, in Patagonia, described by Pagani (2004): P.
619
regularis, P. sabattiniae, Phestia sp. II and Phestia sp. III. The Capivari specimens differ from
620
Phestia aff. P. sabattiniae from the Itararé Group (Taciba Formation, see Neves et al., 2014b),
621
by having an opisthogyrous umbones and a concave posterior-dorsal margin, while in the second
622
species the umbones are orthogyrous and the posterior-dorsal margin is straight. Yet, P. 25
623
tepuelensis presents radial ribs crossing the commarginal costae, a feature absent in the Brazilian
624
specimens. Phestia tepuelensis also differs from two Australian species Phestia darwini
625
(=Nuculana darwini) (Dickins, 1957) and Phestia lyonensis Dickins, 1956. The Brazilian species
626
have narrower umbones, which are very pronounced in P. darwini and the surface of shell is
627
ornamented by commarginal ribs, while in P. lyonensis they can form a shallow V-shaped
628
ornamentation (see Dickins, 1963, p. 37).
629 630
Order Pectinida Gray, 1854
631
Superfamily Chaenocardioidea Miller, 1889
632
Family Streblochondriidae Newell, 1938
633
Subfamily Saturnellinae Astafieva, 1994
634
Genus Streblopteria McCoy, 1851
635 636
Type species: Meleagrina laevigata McCoy, 1844 from Carboniferous of Ireland, by posterior
637
designation of Meek and Worthen (1866, p. 333).
638 639
Remarks: González (2006) described the species Streblopteria montgomeryi for the Mojón de
640
Hierro Formation in Patagonia, Argentina; at the same time Pagani (2006) described the species
641
Streblopteria lagunensis from the same localities. Both species are synonyms and Gonzalez
642
(2006) and Pagani (2006) have the same date of publication. In this way, the rule of
643
“Determination by the First Reviser” (ICZN Article 24.2) is applied, and we use Streblopteria
644
lagunensis in the sense of Pagani (2006). Later, Waterhouse (2010 p. 105) designated the
645
Argentinian specimens of Streblopteria montgomeryi Gonzalez (2006) under a new genus
646
Montorbicula, based on the absence of a resilifer and by the presence of a platyvincular hinge.
647
However, the generic diagnosis is imprecise. Besides, the differences with Streblopteria and 26
648
other related new genera proposed by Waterhouse (2010) are unclear in the discussion. Thus we
649
decided to keep the original generic assignation for the Patagonian species.
650 651
Streblopteria aff. S. lagunensis Pagani, 2006
652
Figure 5.5–5.7
653 654
1978 Streblopteria sp. nom. nud.; in Rocha-Campos and Rösler (1978, p. 5).
655
1988 Streblopteria sp. nom. nud.; in Simões et al. (1998, p. 445).
656 657
Occurrence: Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
658
Basin, Brazil. Latest Asselian–earliest Sakmarian.
659 660
Description: Internal mold of right valve, suborbicular, slightly longer than higher, inflated,
661
acline. Umbones slightly project above the cardinal margin. Anterior auricle separated from the
662
shell by a moderately deep byssal notch. Posterior auricle small, subtriangular, with its posterior
663
margin forming an obtuse angle with the postero-dorsal margin. Cardinal margin straight;
664
anterior margin seems to be more expanded than the posterior one; ventral, posterior and,
665
possibly, anterior (fragmented) margins rounded. External surface of shell ornamented by faint
666
and fine growth lines, at least in the ventral margin. Inner cardinal features and muscle scars not
667
observed.
668 669
Material: One internal mold of right valve, GP/1E 457.
670 671
Measurements: See Table 6 for Streblopteria aff. S. lagunensis shell measurements.
672 27
673
Remarks: Streblopteria aff. S. lagunensis of the Taciba Formation closely resembles
674
Streblopteria lagunensis (=Streblopteria montgomeryi) recorded in lower Permian (Asselian)
675
beds of Patagonia, Argentina (Pagani, 2006, p. 468, fig. 3 M-S; González, 2006, p. 140, fig. 8).
676
The last author suggested that the Argentinean species have smooth shells and inconspicuous
677
commarginal ornamentation, which can also be verified in the Capivari specimen. However, the
678
Brazilian specimen seems to be ornamented by very closely spaced fine growth lines (Figure
679
5.7). Thus, we kept the Capivari species as Streblopteria aff. S. lagunensis.
680 681
Superfamily Heteropectinoidea Beurlen, 1954
682
Family Limipectinidae Newell and Boyd, 1990
683
Subfamily Limipectininae Newell and Boyd, 1990
684 685
Genus Limipecten Girty, 1904
686 687
Type species: Limipecten texanus Girty, 1904 from the Carboniferous of Mexico, by
688
subsequent designation of Newell (1938, p. 68).
689 690
Remarks: Mendes (1952) assigned the pectinid of the Taciba Formation, Capivari County, to
691
Aviculopecten capivariensis. He pointed out that these specimens had just one order of costae
692
(Mendes, 1952, p. 13). However, Rocha-Campos (1969) referred them to Limipecten
693
capivariensis, since they show surface ornament formed by two or three order costae (Rocha-
694
Campos, 1969, p. 48). He referred this bivalve to Limipecten capivariensis nom. nud. We agree
695
with this interpretation and here this species is formally described for the first time.
696 697
Limipecten capivariensis (Mendes), 1952 28
698
Figure 5.8–5.10
699 700
1952 Aviculopecten capivariensis; Mendes (p. 12, plate I, figs. 8 e 9).
701
1967 Aviculopecten capivariensis; Rocha-Campos (plate XXIX, fig. 14).
702
1969 Limipecten capivariensis nom. nud.; in Rocha-Campos (p. 47, plate II, F).
703
1988 Limipecten capivariensis nom. nud.; in Simões et al. (1988, p. 445).
704 705
Holotype: The holotype (DGM 4121) of Limipecten capivariensis (Mendes), 1952 is lost.
706
Lectotype GP/1E 2494 from the Taciba Formation, Itararé Group, Paraná Basin, Capivari
707
county, State of São Paulo, southeastern Brazil.
708 709
Occurrence: Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
710
Basin, Brazil. Latest Asselian–earliest Sakmarian.
711 712
Diagnosis: Convex left valve. Valve ornamented by, at least, ten coarse first order radial ribs,
713
intercalated by second order costae; interspaces among ribs wider than in the right valve. Surface
714
of shell crossed by concentric lamellae, forming scalloped lines. Right valve moderately convex.
715
Surface of shell ornamented by twenty-two first order radial ribs, intercalated by second and
716
third order costae; ribs finer and numerous than in the left valve; costae are crossed by concentric
717
regular lamellae.
718 719
Description: Shell orbicular, inequivalved, with length slightly greater than width. Convex left
720
valve; umbo and auricles not preserved (fragmented). Surface of valve ornamented by, at least,
721
ten coarse first order radial ribs; second order ribs increase by intercalation; interspaces among
722
ribs are wider in the left valve. Surface of shell crossed by concentric lamellae, forming 29
723
scalloped lines. Right valve moderately convex, mainly in the ventral margin. Umbo poorly
724
preserved. Anterior ear sharply defined, separated from the body shell by a well-marked byssal
725
sinus; seven first order ribs crossed by concentric regularly spaced lamellae. The posterior ear is
726
poorly preserved. Surface of shell is ornamented by twenty-two first order radial ribs,
727
intercalated by second and third order costae; ribs are much finer and numerous than in the left
728
valve; surface of shell is crossed by concentric regular lamellae, which are also finer and fainter
729
than in the left valve. Internal features of hinge plate not observed.
730 731
Materials: Two external molds, one fragment of left valve, with well preserved surface
732
ornamentation (GP/1T-116), and one almost complete right valve (GP/1E 2494).
733 734
Measurements: See Table 6 for Limipecten capivariensis shell measurements.
735 736
Remarks: Limipecten capivariensis differs from the type species L. texanus, from Texas, USA,
737
by showing much wider interspaces between ribs in the left valve. Additionally, the right valve
738
of the Brazilian shell is much taller than the North American one. The Capivari shells also differ
739
from Limipecten sp. of the Ciénega Larga de Tontal Formation, San Juan, Argentina (Lech and
740
Milana, 2006), by the absence of protuberances in the meeting of ribs and growth lines, which is
741
a remarkable feature of these specimens. Limipecten capivariensis also differs from another
742
Argentinean species: Limipecten? sp. described by González (1972) in the Las Salinas Formation,
743
Chubut, later revised by Pagani (2006) as Limipecten herrerai. In the Brazilian shells, the radial
744
ribs are well spaced than in L. herrerai. Yet, the commarginal lamellae are more well spaced
745
than in the Argentinian specimens.
746 747
Infrasubcohort Cardiidia Férussac, 1822 30
748
Superfamily Grammysioidea Miller, 1877
749
Family Sanguinolitidae Miller, 1877
750
Subfamily Sanguinolitinae Miller, 1877
751
Genus Praeundulomya Dickins, 1957
752 753
Type-species. Praeundulomya concentrica Dickins, 1957 from the Carnarvon Basin, lower
754
Permian, Western Australia.
755 756
Praeundulomya cf. subelongata Dickins, 1963
757
Figure 5.11–5.12
758 759
Occurrence. Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
760
Basin, Brazil. Latest Asselian–earliest Sakmarian.
761 762
Description. Internal mold strongly inequilateral, subequivalve, very elongate posteriorly.
763
Umbones subterminal; beaks slightly prosogyrous. Very narrow sinus extending from the
764
umbones to ventral margin; well-developed postumbonal ridge extends to posteroventral
765
extremity. Anterodorsal margin very short; anterior margin rounded; ventral margin slightly
766
convex; posterior extremity rounded and defined by the respiratory margin. Lunule and
767
escutcheon were not observed. Surface ornament of very coarse, regularly spaced
768
commarginal rugae. Hinge edentulous. Muscle scars not observed.
769 770
Material. One internal mold of conjugated valves (GP/1E-5242).
771 772
Measurements: See Table 6 for Praeundulomya cf. subelongata shell measurements. 31
773 774
Remarks: Praeundulomya subelongata was first described by Dickins (1963), in the Fossil Cliff
775
Formation, Western Australia. Recently, this species was reported as Praeundulomya cf.
776
subelongata, in the Taciba Formation, Itararé Group (Neves et al., 2014b). The Capivari
777
specimen closely resembles that of the Taciba Formation. For this reason, we assign the Capivari
778
material to Praeundulomya cf. subelongata. This species differs from Praeundulomya moreli
779
Gonzalez, 2006 from Early Permian beds of Tepuel-Genoa Basin, Argentina. The Brazilian shell
780
is very elongated posteriorly, with the postero-ventral angle defined by a well-marked
781
postumbonal carina, while the shell of P. moreli shells is more subquadrate in outline, with a
782
truncate dorsal-posterior margin, and the postero-ventral angle forming a gentle carina.
783 784
Class Gastropoda Cuvier, 1797
785
Order Vertigastropoda Salvini-Plawen, 1980
786
Superfamily Eotomarioidea Wenz, 1938
787
Family Eotomarridae Wenz, 1938
788
Subfamily Neilsoniidae Knight, 1956
789 790
Genus Peruvispira Chronic, 1949
791 792
Type species: Peruvispira delicata Chronic, 1949 from the Copacabana Group (lower Permian),
793
Peru.
794 795
Remarks: Peruvispira is a genus that close resembles Ptycomphalina Fischer 1885 (see Dickins,
796
1963, p. 126), and Pleurocintosa Fletcher (1958). Indeed, according to Dickins (1961),
797
Peruvispira is similar to Ptycomphalina but ‘characterized by the possession of a distinct 32
798
revolving concave area about as wide as and below the slit-band’. On the other hand, the
799
relationship of Peruvispira and other Late Paleozoic gastropods, such as Pleurocintosa, was
800
recently discussed by Taboada et al. (2015). We agree with them that Pleurocintosa cannot be
801
regarded as a valid genus, since it was erected based on inconspicuous (e.g., weak spiral filae)
802
and highly variable characters (e.g., presence of a third and peribasal carina bounding the
803
alveozone) found in species of Peruvispira (see Taboada et al., 2015, p. 219 for details).
804 805
Peruvispira brasilensis n. sp.
806
Figure 6.1–6.2
807 808
1966 Peruvispira delicata nom. nud.; in Rocha-Campos, p. 10, fig. 3 a, b.
809
1970 Peruvispira delicata nom. nud.; Rocha-Campos, p. 607, fig. 3.
810 811
Holotype: GP/1E-4319 from the Taciba Formation, State of São Paulo, southeastern Brazil.
812 813
Diagnosis: Pleurotomarid shell with 4-5 very concave whorls; deep and strongly concave
814
selenizone, ornamented by 8-9 lunulae per mm, bounded by two prominent carinae. Shell
815
ornamentation of thin growth lines (7-8 per mm).
816 817
Occurrence: Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
818
Basin, Brazil. Latest Asselian–earliest Sakmarian.
819 820
Description: Small and turbiniform pleurotomarid shell, moderately thick; apex and upper whorl
821
slightly convex; shell with 4-5 very concave whorls; selenizone deep and strongly concave,
822
moderately wide, ornamented by 9-10 concave lunulae per mm, bounded by two prominent 33
823
carinae; shell ornamented by 7-8 thin growth lines per mm. Lower face continuously rounded
824
and convex, not flattened; sutures well impressed; inner lip poorly preserved; outer lip well
825
developed, convex and rounded, becoming almost perpendicular to the last whorl to form a sinus.
826
Shell body thicker near aperture.
827 828
Etymology: From Brazil.
829 830
Materials: Three external molds, GP/1E-4319 and 4348A, B.
831 832
Measurements: See Table 6 for Peruvispira brasiliensis n. sp. shell measurements.
833 834
Remarks: Peruvispira brasilensis n. sp. differs from the type species P. delicata Chronic, 1949,
835
from the Carboniferous of Peru, by the presence of very concave whorls, limited by a deep and
836
strongly concave selenizone. The Brazilian species closely resemble Peruvispira cf. umariensis
837
(Dickins, 1963, p. 126), from Western Australia, however they differ in the density of the growth
838
lines. Peruvispira brasilensis n. sp. is ornamented by 7-8 growth lines per mm, while P. cf.
839
umariensis is nearly 4 per mm. Additionally, the Brazilian species shows higher and more
840
concave whorls. Peruvispira brasilensis n. sp. can be distinguished from P. sueroi from the Late
841
Paleozoic beds of Chubut, Argentina (Sabatini and Norait, 1969) by having a very concave
842
whorl area, in which the spires are gradually higher toward the anterior region. In P. sueroi, the
843
whorl area is slightly convex and the basal whorl is much higher than the other ones. Peruvispira
844
australis is another comparable Argentinean species from San Juan, Mendoza and Chubut
845
(Sabatini and Noirat, 1969, p 104). P. australis differs from P. brasilensis n. sp. by the same
846
characters of P. sueroi, for being three times wider and having more widely spaced growth lines
847
(2-3 per mm) as well. 34
848 849
Class Crinoidea Miller, 1821
850
Subclass and order uncertain
851
Group Pentameri Moore, Jeffords and Miller, 1968
852
Family øPentacauliscidae Moore, Jeffords and Miller, 1968
853 854
Genus øPentaridica Moore, Jeffords and Miller, 1968
855 856
Type species: øPentaridica rothi Moore, Jeffords and Miller, 1968.
857 858
øPentaridica sp.
859
Figure 6.4-6.6
860 861
Occurrence. Capivari County, State of São Paulo. Taciba Formation, Itararé Group, Paraná
862
Basin, Brazil. Latest Asselian–earliest Sakmarian.
863 864
Materials: Four individual discs (articular facet) GP/1E-4321; 4349 B and C; 4350C, and one
865
pluricolumn, GP/1E 4349A.
866 867
Measurements: See Table 7 for crinoid discs measurements.
868 869
Remarks: The overwhelming majority of Paleozoic crinoids are preserved as disarticulated
870
skeletal elements, mainly stem ossicles. In most cases linking them with other anatomical crinoid
871
parts (i.e., crowns) is virtually impossible. Therefore, in many cases, as is the case of the genus
872
øPentaridica, the taxonomy is based only on stem anatomy. Thus, stem-based genera have a 35
873
minimal chance to express true phylogenetic relationships (Głuchowski, 2002). Despite the
874
difficult determining the biological consistency of crinoid species on stem morphology alone (for
875
detailed discussion see Moore et al., 1968; Donovan 2001; Głuchowski, 2002) stem elements are,
876
in many cases, unique and allow specific identification.
877
The studied specimens are represented by 22 individual discs (articular facet) and three
878
pluricolumn remains (stems), which are, in the most part, poorly preserved. However, the
879
specimens GP/1E 4319 and 4349 B, C shows articular facets with well-preserved peripheral
880
crenularia, which can be referred to øPentaridica sp. (Dr. Julio Hlebszevitsch, personal
881
communication, April 2016).
882 883
4. Results
884
A total of 150 specimens were studied, belonging to Brachiopoda (77%), Mollusca (7%) and
885
Echinodermata, Crinoidea (16%) (Table 4). They were assigned to four brachiopod species
886
(Lyonia rochacamposi, Rhynchopora grossopunctata, Biconvexiella saopauloensis n. sp,
887
Quinquenella rionegrensis), four bivalve species (Phestia tepuelensis, Streblopteria aff. S.
888
lagunensis, Limipecten capivariensis, Praeundulomya cf. subelongata), and two gastropod
889
species [Peruvispira brasilensis n. sp, and Mourlonia (Woolnoughia)? sp.] (Tables 5, 6). Crinoid
890
columns were assigned to øPentaridica (Table 7). The overwhelming majority of brachiopod
891
shells belong to Biconvexiella saopauloensis (n= 86, 74,8%), and is followed by Rhynchopora
892
grossopunctata (n=20, 1,7%). Table 4 shows the total number of the remaining brachiopod
893
specimens. On the other hand, the bivalves are less common with just a few specimens of each
894
taxon (Table 4). The gastropods are also rare and represented by three shells only, two assigned
895
to Peruvispira brasilensis, and one to Mourlonia (Woolnoughia)? sp., this last one just illustrated
896
here (Fig. 6). Therefore, the assemblage is dominated by shells of Biconvexiella saopauloensis,
897
and is here referred as the Bs assemblage. 36
898
As noted above, the taphonomic information available for the bioclastic remains in the Bs
899
assemblage is limited, but some observations are noteworthy. First, the majority of both
900
brachiopod and bivalve shells are disarticulated. The most preserved specimens (articulated
901
shells) are illustrated in the figures 4 and 5. However, the shells are complete and the
902
ornamentation (when visible) is well preserved (i.e., signs of abrasion, bioerosion, and
903
encrustation are missing), and various size classes are represented. Closed articulated bivalve
904
mollusk shells are also observed. Indeed, Praeundulomya cf. subelongata is preserved with
905
closed articulated shells, and with the long axis inclined ~40° to bedding. Due to compaction, the
906
specimen is slightly compressed through the anterior-posterior axis of the shell. Thus, the angle
907
between the long axis and bedding has probably been reduced.
908
Finally, the crinoid remains are represented mainly by disarticulated skeletal ossicles
909
(mostly columnals), which are disperse in the matrix. Crinoid remains varies in size from 1.5 to
910
3.5 mm, and abrasion of crenelations and edges are not visible (i.e., columnals are pristine, see
911
Figure 6). Signs of encrustation, bioerosion and corrosion are also missing.
912 913
5. Discussion
914
5.1. Depositional environment
915
In the study area, the investigated marine fossil bed is recorded in the Taciba Formation
916
(formerly referred as Capivari Formation) and placed 100-150 m from the contact with the
917
overlying, post-glacial Early Permian Tatui Formation. The vertical profile reveals the interplay
918
of distinct sedimentary processes in a subaqueous environment and the presence of an
919
invertebrate marine fauna in a glacio-marine context. A clear change in the marine setting from a
920
low- to a high-energy environment is recorded, probably due to a glacier advance event in the
921
depositional setting (Fig. 3). Initially, sedimentation took place in a distal, low-energy
922
environment in an epeiric sea with free-floating icebergs. In this context, massive mudstones 37
923
overlying dark shales suggests increased sediment input (Fig. 3). The presence of bottom
924
currents, which resulted in current-rippled medium- to fine-grained sandstones are indicative of
925
bedform migration, coupled with sediment transportation by suspension (Fig. 3). Trough cross-
926
bedded strata interbedded with laminated siltstone occurs in the middle portion of the studied
927
section and records increased high-energy processes. This suggests migrating 3D bedforms
928
formed by subaqueous meltwater flows and/or turbiditic currents. Due to the rapid sedimentation
929
of these beds, flame structures were formed by the overloading of medium-grained massive
930
sandstones on fine-grained facies. Association with crudely parallel-laminated, plant-bearing
931
siltstones contrasts with the dysoxic environment during deposition of dark gray shales recorded
932
in the lowermost portion of the studied section. Increased sediment supply and input of
933
phytoclasts are suggestive of terrestrial contribution probably derived from contemporaneous
934
coastal progradational systems, or directly supplied by subaqueous outwash currents associated
935
with oscillating glacier margins in the context of a wet-based grounded ice shelf. To the top,
936
fossil-bearing silty shales record the environmental change to a low-energy shelfal depositional
937
setting (Fig. 3). The intervening of an episode of glacial retreat is corroborated by the absence of
938
iceberg-related dropstones and meltwater bottom currents. The relatively calm offshore
939
environment that was colonized by the marine invertebrates was eventually disrupted by high-
940
energy sedimentation events that enhanced fossil preservation. After that, a new glacial advance
941
event induced the sedimentation of pebbly mudstones that may have inhibited the bottom
942
colonization by shelly benthos. In the examined sedimentary succession, the percentage of
943
pebbles increases upward and the transition to pebbly silty sandstone is indicative of renewed
944
glacial sedimentation in the depositional setting.
945 946
5.2. Taphonomy
38
947
As commented above the taphonomic information available for the bioclastic remains in the Bs
948
assemblage is limited and mainly based on museum specimens that were not properly collected
949
for taphonomic studies. Anyway, some observations are worth to mention. Most of brachiopods
950
and bivalves are represented by disarticulated shells. However, the shells are complete and the
951
ornamentation (when visible) is well preserved, including various size classes. These features
952
indicate that after death most of the shells had, probably, limited exposure at the sediment/water
953
interface, suffering only minor lateral transport. These are probably parautochthonous (sensu
954
Kidwell et al., 1986) elements in this assemblage. By contrast, specimens of Praeundulomya cf.
955
subelongata were preserved in situ. Similar preservational conditions are also noted in
956
specimens of Praeundulomya and other associated Grammysioidea bivalves in the uppermost
957
part of the Taciba Formation, State of Paraná, southern Brazil (see Neves et al., 2014b). There,
958
the shells are preserved in fine to very fine sandstones/siltstones with hummocky cross-
959
stratification and intercalated mudstones forming stacked storm deposits in distal shoreface
960
settings (Neves et al., 2014b). As in the case of Praeundulomya from the Capivari beds, those
961
shells are closed articulated, compacted, and with the anterior margin pointing downward. The
962
long axis of the shell inclined 40°-50° to bedding. As convincingly demonstrated by Anelli et al.
963
(1998), such specimens were preserved in life position, and a similar condition is inferred for
964
Praeundulomya from the Bs assemblage. Hence, this species is the unique autochthonous
965
element so far identified in this assemblage.
966
The crinoid remains are all usually small (1.5 to 3.5 mm) disarticulated ossicles or rarely
967
articulated stem segments, without any signs of abrasion, encrustation, bioerosion and/or
968
corrosion. These taphonomic signatures indicates that after death and disarticulation, which can
969
occur in a few weeks, the columnals were not intensely transported and sorted (see Llewellyn
970
and Messing, 1993). Thus, the crinoid remains are probably parautochthonous. Finally, the
971
presence of comminuted plant remains (Fig. 3) is also worth mentioning, suggesting the 39
972
contribution of allochthonous materials from coastal settings during events of enhanced riverine
973
freshwater input.
974 975
5.3. Paleoautoecology
976
The Bs assemblage is numerically dominated by ambocoelids, a group of small, thin-shelled
977
brachiopods with an open delthyrium and a pedicle during life (see He et al., 2012). However,
978
the fine-grained, soft, siliciclastic-dominated facies where Biconvexiella saopauloensis is
979
recorded may have offered limited site opportunities for pedicle attachment. Indeed, the shells
980
are not found in clusters, and signs of encrustation are unknown in the studied specimens. Hence,
981
this tiny and spinose brachiopod species may have lived on soft substrates with the ventral valve
982
partially embedded within the substrate, a condition also shown by other ambocoelids (e.g.,
983
Ambocoelia gregaria Hall, 1860; in Zambito and Schemm-Gregory, 2013). The presence of
984
spine bases suggest that the shell was covered by tinny spines in life.
985
Given the dominance of Biconvexiella saopauloensis in the assemblage, this species may
986
haveacted as a pioneer species in muddy, dysoxic bottoms during deposition of host sediments.
987
Indeed, as for other ambocoelids (e.g., Attenuatella mengi He et al., 2012), this shell form is
988
usually associated with low oxygen and/or limited trophic resources in the sedimentary settings
989
inhabited by those brachiopods (see He et al., 2012 and references therein). Their small size is a
990
good indicator that the conditions were stressful, probably coupled with oscillations in the
991
oxygen content, salinity and food-supply. The linoproductids, such as Lyonia rochacamposi,
992
with spinose convex ventral valves, are interpreted as a quasi-infaunal, suspension feeders that
993
lived partially buried in soft substrates (see Cisterna et al., 2017). The other two brachiopods in
994
the Bs assemblage (Rhynchopora grossopunctata, Quinquenella rionegrensis) were epifaunal
995
suspension feeders.
40
996
The bivalves Streblopteria aff. S. lagunensis and Limipecten capivariensis were both
997
stationary epifaunal suspension feeders (Stanley, 1970, 1972), whereas Praeundulomya cf.
998
subelongata with a posteriorly elongated shell (Stanley, 1970) was a facultatively mobile
999
infaunal suspension feeder. As other nuculanoids, Phestia tepuelensis was a facultatively mobile
1000
infaunal deposit feeder/suspension feeder (Stanley, 1970). The other mollusks in the assemblage,
1001
the small gastropod Peruvispira and Mourlonia (Woolnoughia)? sp. were both facultatively
1002
mobile epifaunal suspension feeders or grazers (Aberhan and Kiessling, 2015). Yet, Peruvispira
1003
is known for its broad environmental tolerances in Early Permian marine assemblages from
1004
Australia (Clapham and James, 2008). Finally, the crinoid was a stationary epifaunal suspension
1005
feeder.
1006 1007
5.4. Paleocommunity and paleoenvironment
1008
With only 11 genera, the Biconvexiella saopauloensis assemblage is diverse when compared
1009
with other coeval benthic marine assemblages associated with LPIA sedimentary successions in
1010
South America (i.e., Argentina) and Australia (see data in Cisterna et al., 2019 and references
1011
therein). However, among the known macroinvertebrate marine faunas of the upper part of the
1012
Taciba Formation, the Bs assemblage is the most diverse one, including members of three major
1013
phyla (Brachiopoda, Mollusca and Echinodermata). The assemblage encompasses benthic
1014
invertebrates occupying distinct substrate tierings (Table 8). The epifaunal tiering encompassed
1015
(a) the low-level stationary epifaunal suspension feeders, including all the brachiopods, and
1016
epifaunal bivalves. Numerically, this was the dominant ecological group in the assemblage, (b)
1017
the stationary upper-level epifaunal suspension feeders (i.e., crinoids), and (c) vagile epifaunal
1018
suspension feeders or grazers (Aberhan and Kiessling (2015), including all the small gastropods.
1019
The shallow infaunal tiering was occupied by (d) facultatively mobile infaunal deposit
1020
feeders/suspension feeders, such as the protobranchiate bivalves (i.e., Phestia), and (e) the 41
1021
facultatively mobile shallow infaunal suspension feeders (i.e., Praeundulomya). Both infaunal
1022
groups colonized the fine-grained, siliclastic-dominated substrates of the Capivari marine beds
1023
just a few centimeters below the sediment water/interface. Surprisingly, planktic and nektic
1024
organisms are missing in the assemblage. The absence of these groups may be a result of a
1025
combination of processes related to taphonomic and sampling biases and/or stressful
1026
paleoenvironmental conditions. The fossil-bearing mudstones of the Taciba Formation are dark
1027
grey/green (or even yellowish when weathered), and finely laminated (horizontal bedding)
1028
indicating the settling of muds in quiet shelf, with restricted oxygen and food supply. Indeed, as
1029
commented by Harper and Moran (1997, p. 235) brachiopods, the numerically dominant group
1030
in the assemblage, were minimalist animals, “operating as a ciliary suspension feeder, if
1031
necessary, at very low oxygen levels.” The general small body size of the Capivari brachiopod
1032
fauna is in accordance with these observations.
1033
The presence of members of the rhynchonelliform brachiopods, sanguinolitid and pectinid
1034
bivalves, pleurotomariid gastropods and crinoids clearly indicates that the Bs assemblage is fully
1035
marine. However, the restricted vertical distribution of these marine groups in the uppermost part
1036
of the Itararé Group in the São Paulo State suggests that the background conditions were
1037
generally unfavourable to the profuse development of the benthic fauna or their preservation. As
1038
noted above, the invertebrates lived and/or were preserved in a distal glaciomarine environment
1039
during more proximal marine deglaciation (Figs. 7, 8, see Vesely and Assine, 2006, p. 164). The
1040
clay-rich, fossil-bearing marine horizon containing members of the Bs assemblage show no
1041
evidence of glacial influence (ice-rafted dropstones), indicating that this may correspond to a
1042
time of ice retreat during the sequence time span (Fig. 7, 8). This interval may correspond to a
1043
time of climatic amelioration, also supported by the presence of plant remains recorded in
1044
laminated siltstones, located nearly 2 meters below the marine fossil-bearing interval (Figs. 3, 7,
1045
8). This may indicate the presence of plant covered, deglaciated continental terrains (Fig. 8). 42
1046
Finally, Cisterna et al. (2017, and references therein) suggested that a complex array of
1047
abiotic factors encompassing variation in salinity and oxygen contents, water turbidity and depth,
1048
and food-supply are among the main controlling factors in the development of marine benthic
1049
faunas during deglacial events of LPIA. Notably, a comparison between the gross taxonomic
1050
composition and ecological structure of the Capivari marine fauna with those coevals from the
1051
upper part of the Taciba Formation, southern Brazil (Simões et al., 2012; Neves et al., 2014b;
1052
Taboada et al., 2016), strongly support the observations made by Cisterna et al. (2017). The
1053
benthic marine assemblages recorded in the so-called Rio da Areia sandstone and Baitaca
1054
siltstone, in the upper part of the Taciba Formation (Neves et al., 2014b), which are dominated
1055
by epifaunal and infaunal suspension feeding bivalves lived in quite distinct paleoenvironmental
1056
conditions compared to those of the Bs assemblage. Taphonomic and sedimentologic
1057
information (Neves et al., 2014b) conclusively indicates that the benthic faunas from the
1058
southern part of the basin lived in coastal to shelf areas commonly affected by moderate to
1059
strong bottom currents and episodically affected by high-energy flows and currents generated by
1060
storms, which is not the case of the Bs assemblage.
1061 1062
5.5. Biostratigraphic implications
1063
Our data support the correlation of the Bs assemblage with those from the uppermost part of the
1064
Taciba Formation, from southern Brazil, as previously pointed out by Taboada et al. (2016, p.
1065
437). Particularly noteworthy is the record of Lyonia rochacamposi that is common in the Taciba
1066
Formation from the Bela Vista do Sul assemblage (Taboada et al., 2016). Additionally, close
1067
species of Quinquenella, Phestia, Limipecten, and Praeundulomya are also recorded in the
1068
Teixeira Soares assemblages (Neves et al., 2014b; Taboada et al., 2016). Hence, the Bs
1069
assemblage correlates with the upper portion of the Itararé Group from southern Brazil,
1070
suggesting an early Permian (latest Asselian-earliest Sakmarian) age. These data indicate that the 43
1071
marine fossil-bearing beds of Capivari and Teixeira Soares regions are stratigraphic records of
1072
northward transgressive events during the end of LPIA in the Paraná Basin. Faunal similarities
1073
suggest that the latest Asselian-earliest Sakmarian Paraná marine assemblages correlate with
1074
those of the Sauce Grande-Colorado (Argentina), Huab (Hardap shale of the Dwyka Group),
1075
Aranos area (Namibia), southwest Africa, and the Carnarvon (Western Australia) basins, and
1076
characterize the Eurydesma-Lyonia chronozone, reinforcing the paleogeography scenario of a
1077
W-E trans-Gondwanan marine seaway linking those basins during the Early Permian, as detailed
1078
discussed in Taboada et al. (2016).
1079 1080
6. Significance and concluding remarks
1081
In this contribution, we report a cool-water fauna from the LPIA, which was found in a fossil-
1082
rich bed of the upper part of the Itararé Group from the Capivari county, on the eastern border of
1083
the Paraná Basin. The fauna was described and its stratigraphic position within the glacial Itararé
1084
Group better constrained. The measured sedimentary succession is placed between 100-150 m
1085
below the contact with the overlying, post-glacial Early Permian Tatuí Formation.
1086
Members of the Biconvexiella saopauloensis assemblage flourished in a rather confined
1087
paleogeographic setting in the northeastern border of the Paraná Basin, under stressful conditions
1088
mainly linked to changes in oxygen content and food-supply. Although evidences of freshwater
1089
discharge from coastal areas are recorded in the studied sedimentary succession, the Bs
1090
assemblage is fully marine, suggesting that salinity conditions were normal, at least during the
1091
time of benthic faunal development.
1092
The fossil-bearing marine beds of Capivari county may represent short-lived transgressive
1093
events during periods of ice retreats. These beds are overlain by pebbly mudstones and silty
1094
sandstones recording high energy processes under glacial conditions. Since Bs assemblage is
1095
placed in between subaqueous glacial facies associations, they may correspond to high frequency 44
1096
interglacials. Consequently, the fossil-bearing marine beds generated in this stratigraphic context
1097
are thin and difficult to find within the thick glacial package. This is particularly true when
1098
proper subsurface data are missing and/or outcrops are deeply weathered and covered by dense
1099
vegetation, which is the case for the studied fossil-bearing beds. Faunal composition indicates
1100
that the transgressive fossil-bearing beds of the Capivari region correlates with those of the upper
1101
part of the Taciba Formation from the southern part of the Paraná Basin (States of Paraná and
1102
Santa Catarina), suggesting an Early Permian (latest Asselian-earliest Sakmarian) age. Indeed, in
1103
the upper part of the Taciba Formation, the spore-pollen assemblages are referable to the
1104
Vittatina costabilis Biozone of the Paraná Basin indicative of an Early Permian age (Souza and
1105
Callegari, 2004; Souza and Toigo, 2005; Souza, 2006). These marine faunas also indicate the
1106
increase of glacial-marine depositional conditions in the uppermost part of the Itararé Group,
1107
which is tied to the last cycle of ice advance and retreat during the LPIA.
1108 1109
Acknowledgments
1110
The authors are very indebted to the Instituto de Geociências, Universidade de São Paulo, São
1111
Paulo, for access and support in the study of the invertebrate collection of the Capivari fossils.
1112
We also thank Ivone C. Gonzales, IGc/USP, for assistance with this collection. Julio
1113
Hlebszevitsch helped us with crinoid identification and Suzana A. Matos, IBB/UNESP, assisted
1114
us in the field. Hence, both are here acknowledged. Many thanks also to Paul A. Johnston, and
1115
two anonymous reviewers for their suggestions, corrections and comments that improved the
1116
final version of the manuscript. Editor-in-Chief, Francisco J. Vega is also acknowledged for the
1117
useful suggestions and corrections. Finally, we also thank the paleoillustrator Julio L.C. Bezerra
1118
for the “Capivari sea” paleoscene. Financial support was offered by FAPESP and CNPq, Brazil,
1119
and CONICET, Argentina. JPN was a fellow of FAPESP (grant 2013/25317-7). This project is a
45
1120
contribution to the following FAPESP projects: 2013/25317-7 and 2014/09149-0. CNPq grants:
1121
304925/2017-9 to MLA and 301023/94-3 to MGS.
1122 1123
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1124
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Boletim IG-USP, Série Científica, 12, 71–82.
1463 1464 1465 1466 1467 1468 1469
Soares, P.C., 1991. Tectônica sinssedimentar cíclica na Bacia do Paraná-controles. Full Professor thesis, Universidade Federal do Paraná. [in Portuguese]. Souza, P.A., 2006. Late Carboniferous palynostratigrafphy of the Itararé Subgroup, northeastern Paraná Basin, Brazil: Review of Palaeobotany and Palylonogy, 138, 9–29. Souza, P.A., and Callegari, L.M., 2004. An Early Permian palynoflora from the Itararé Subgroup, Paraná Basin, Brazil: Revista Española de Micropaleontología, 36, 439–450. Souza, P.A., and Marques-Toigo, M., 2005. Progress on the palynostratigraphy of the Permian
1470
strata in Rio Grande do Sul State, Paraná Basin, Brazil: Anais da Academia Brasileira de
1471
Ciências, 77, 353–365.
1472 1473 1474 1475 1476 1477 1478 1479 1480 1481
Stanley, S.M., 1970. Relation of shell form to life habits of the Bivalvia (Mollusca). Geological Society of America Memoir, 125, 1–296. Stanley, S.M., 1972. Functional morphology and evolution of byssally attached bivalve mollusks. Journal of Paleontology, 46, 165–212. Stehli, F.G., 1954. Lower Leonardian Brachiopoda of the Sierra Diablo: Bulletin of the American Museum of Natural History, 105, 257–358. Sterren, A.F., 2004. Bivalvos pérmicos de la Formación Tupe em la quebrada La Herradura: Ameghiniana, 41, 57–74. Taboada, A.C., 1989. La fauna de la Formación El Paso, Carbónico inferior de la precordillera sanjuanina: Acta Geológica Lilloana, 17, 113–129.
1482
Taboada, A.C., Mory, A., Shi, G.R., and Pinilla, K., 2015. The Early Permian brachiopod fauna
1483
from Pasminco BW 9 borehole (Calytrix Formation) on the Barbwire Terrace, Canning
1484
Basin, Western Australia: Alcheringa, 39, 207–223. 60
1485
Taboada, A.C., Neves, J.P., Weinschütz, L.C., Pagani, M.A., and Simões, M.G., 2016.
1486
Eurydesma–Lyonia fauna (Early Permian) from the Itararé group, Paraná Basin (Brazil): a
1487
paleobiogeographic W–E trans-Gondwanan marine connection: Palaeogeography,
1488
Palaeoclimatology, Palaeoecology, 449, 431–454.
1489
Tedesco, J., Cagliari, J. Chemale Junior, F., Girelli, T., and Lana, C., 2019. Provenance and
1490
paleogeography of the Southern Paraná Basin: Geochemistry and U–Pb zircon
1491
geochronology of the Carboniferous-Permian transition: Sedimentary Geology, on-line,
1492
doi.org/10.1016/j.sedgeo.2019.105539
1493
Verneuil, E. de, 1845. Paléontologie, mollusques, brachiopodes. In: Murchison, R. I., Verneuil,
1494
E., and Keyserling, A. (Eds.), Géologie de la Russie d’Europe et des Montagnes de l’Oural 2,
1495
Part 3: London, Paris, p. 17–395.
1496
Vesely, F.F., and Assine, M.L., 2006. Deglaciation sequences in the Permo-Carboniferous Itararé
1497
Group, Paraná Basin, southern Brazil. Journal of South American Earth Sciences, 22, 1–156.
1498
Waagen, W., 1883. Salt Range Fossils. Productus-Limestone Fossils: Geological Survey of India,
1499 1500 1501 1502 1503 1504
Memoirs, Palaeontologia Indica, 2, 39–546. Waterhouse, J.B., 1964. Permian brachiopods of New Zealand: New Zealand Geological Survey, Paleontological Bulletin, 35, 1–285. Waterhouse, J.B., 1975. New Permian and Triassic brachiopod taxa: Department of Geology of the University of Queensland papers (new series), 7, 1–23. Waterhouse, J.B., 1982a., New Zealand Permian Brachiopod Systematics, Zonation, and
1505
Paleoecology: Geological Survey of New Zealand, Palaeontological Bulletin, 48, 1–158.
1506
Waterhouse, J.B., 1982b. An early Permian cool-water fauna from pebbly mudstones in south
1507 1508 1509
Thailand: Geological Magazine, 119, 337–432. Waterhouse, J.B., 1983a. New Permian invertebrate genera from the East Australian segment of Gondwana: Indian Geologists’ Association, Bulletin, 16, 153–158. 61
1510
Waterhouse, J.B., 1983b. Permian Brachiopods from Pija Member, Senja Formation, in Manang
1511
District of Nepal, with new brachiopod genera and species from other regions: Indian
1512
Geologists' Association, Bulletin, 16, 111-151.
1513
Waterhouse, J. B., 1986. Late Palaeozoic Scyphozoa and Brachiopoda (Inarticulata,
1514
Strophomenida, Productida and Rhynchonellida) from the southeast Bowen Basin, Australia:
1515
Palaeontographica, Abt. A, 193, 1–76.
1516
Waterhouse, J. B., 2001. Late Paleozoic Brachiopoda and Mollusca from Wairaki Downs, New
1517
Zealand.With notes on Scyphozoa and Triassic ammonoids and new classifications of
1518
Linoproductoidea (Brachiopoda) and Pectinida (Bivalvia): Earthwise, 3, 1–195.
1519 1520
Waterhouse, J.B., 2008. Aspects of the evolutionary records for fossils of the bivalve subclass Pteriomorphia Beurlen: Earthwise, 8, 1–220.
1521
Waterhouse, J.B., 2010. New Late Paleozoic Brachipod and Molluscs: Earthwise, 9, 1–134.
1522
Waterhouse, J.B., 2013. The Evolution and Classification of Productida (Brachiopoda):
1523 1524 1525 1526 1527 1528
Earthwise, 10, 1–535. Wenz, W. 1938. Gastropoda, Teil I. In: Schindewolf, O.H. (Ed.), Handbuch der Paläozoologie, Verlag Gebrüder Bornträger, 1639 p. White, I.C., 1908. Relatório sobre as Coal Measures e rochas associadas ao sul do Brasil: Comissão das Minas de Carvão de Pedra do Brasil, Rio de Janeiro, 300 p. Williams, A., Carlson, S.J., Brunton, C.H.C., Holmer, L.E., and Popov, L.E., 1996. A supra-
1529
ordinal classification of the Brachiopoda: Philosophical transaction of the Royal Society of
1530
London (series B), 351, 1171–1193.
1531
Zambito, J., and Schemm-Gregory, M., 2013. Revised taxonomy and autecology for the
1532
brachiopod genus Ambocoelia in the Middle and Late Devonian northern Appalachian Basin
1533
(USA): Journal of Paleontology, 87, 277–288.
1534 62
1535
Captions
1536 1537
Figure 1. Stratigraphic chart of Itararé Group showing the main marine mudstone intercalations
1538
and their lateral correlations. Capivari assemblage (= Biconvexiella saopauloensis assemblage
1539
here described). Araçoiaba, Ortigueira and Mafra assemblages are recorded in thin marine
1540
intercalations in a thick sandstone package. Explanation: LPTS, Late Paleozoic Third Order
1541
Sequence; SB, Sequence Boundary; ass., Assemblage. Modified from Holz et al. (2010) and
1542
Taboada et al. (2016).
1543 1544
Figure 2. A: Distribution of the upper Paleozoic strata (Itararé Group) in the Paraná Basin,
1545
southeastern Brazil and location of the studied section (Capivari marine beds). The location of
1546
the Passinho and Lontras marine beds, southern Brazil, is also indicated. B: Geological map with
1547
the distribution of the upper Paleozoic rocks in the studied area (black star). Source of data: IPT
1548
(1981).
1549 1550
Figure 3. Columnar section of the upper portion of the Itararé Group, Taciba Formation, in the
1551
Capivari region, State of São Paulo, Brazil. The fossil-bearing interval is highlighted.
1552
Abbreviations: C=clay; S = silt; FS = fine sand; MS= medium sand; CS= coarse sand. A: Pebbly
1553
mudstone in the upper portion of the section. B: Fossil-bearing silty shale. C: Sample of
1554
laminated siltstone with rich concentrations of fragmented plant material. Scale bars 1 cm.
1555 1556
Figure 4.
1557
Brachiopods of the Taciba Formation, Capivari county. 1-12: Rhynchopora grossopunctata
1558
Mendes, 1952. 1: conjugated valves, ventral valve view, GP/1T 117a; 2: dorsal valve, GP/1E-
1559
451; 3-5: ventral valves; 3: GP/1E 117c; 4: GP/1E-2507; 5: GP/1E-2545; 6, 11: conjugated 63
1560
valves, GP/1E-4342; 6: dorsal valve view; 11: posterior view; 7: cast of external mold of dorsal
1561
valve, GP/1E-4405; 8: ventral valve, GP/1E-443a; 9, 10, 14, 15: dorsal valves, GP/1E-4325; 9:
1562
dorsal valve view; 10: anterior view; 14: lateral view; 15: posterior view; 12: external mold of
1563
ventral valve, GP/1T 117b; 13: detail of the ventral valve of GP/1E 443b, showing punctae
1564
(5/mm2); 16-25: Biconvexiella saopauloensis sp. nov., 16-19: dorsal valves; 16: cast of GP/1E-
1565
4339 (holotype); 17: natural mold; 18: GP/1E-453; 19: GP/1E 450; 20: external mold of ventral
1566
valve, GP/1E 4322; 21, 22: GP/1E 2513; 21: conjugated valves; 22: close-up of the
1567
subconcentric rows of spine bases; 23: cast of external mold of dorsal valve, GP/1E-4327; 24-25:
1568
ventral valve, 24: GP/1E 2520; 25: GP/1E-4338. 26-28: Quinquenella rionegrensis (Oliveira), 26:
1569
dorsal valve, GP/1E 4355c; 27-28: dorsal valve interior, 27: GP/1E-4354c; 28: GP/1E 4358; 29-
1570
32: Lyonia rochacamposi Taboada et al., 2016; 29: ventral valve in oblique view, GP/1E-4350b;
1571
30: CP.I 769I, specimen from Bela Vista do Sul locality (Southern Brazil), showing a single row
1572
of spines (white arrow); 31: cast of a fragmentary ventral valve, external mold, GP/1E 4350b; 32:
1573
fragmentary ventral valve, external mold, GP/1E 4350a. Scale bar = 10 mm, except in 19; 20;
1574
26-28 = 5 mm. Specimens 2, 6, 11-13; 20-22, 26 are external molds. 3-5; 8-10, 14-19, 24, 25 are
1575
internal molds.
1576 1577
Figure 5.
1578
Bivalves of the Taciba Formation, Capivari county. 1-4: Phestia tepuelensis, 1-3: GP/1E-452; 1:
1579
external mold of right valve; 2: latex cast of 1; 3: detail of 1, showing the radial ribs crossing the
1580
commarginal costae; 4: internal mold of right valve, GP/1E-455. 5-7: Streblopteria aff.
1581
lagunensis, GP/1E-457, internal mold of right valve; 5: dorsal view; 6: antero-posterior view. 7:
1582
right valve view; 8-10: Limipecten capivariensis (Mendes), 8: latex cast of left valve fragment,
1583
GP/1T-116; 9: cast of external mold of right valve, GP/1E 2494; 10: internal mold of
64
1584
undetermined valve, GP/1E 2511; 11-12: Praeundulomya cf. subelongata Dickins, GP/1E-4252;
1585
11: right valve view; 12: ventral margin view. Scale bar = 10 mm, except in 9 and 10 = 5 mm.
1586 1587 1588
Figure 6. Gastropod and crinoid remains of the Taciba Formation, Capivari county. 1-3:
1589
Gastropods, 1-2: Peruvispira brasilensis n. sp. 1: latex cast of external mold ,GP/1E 4348
1590
(holotype); 2: cast of external mold, GP/1E-4319; 3: Mourlonia (Woolnoughia)? sp., GP/1E-454;
1591
4-6: øPentaridica sp. (personal communication Dr. Julio Hlebszevitsch, April 2016); 4: Articular
1592
facet showing the peripheral crenularium [L2]GP/1E-4321; 5: Detail of a pluricolumn, GP/1E-
1593
4349A; 6: two articular facets with well-preserved peripheral crenularium, GP/1E-4349. Scale
1594
bar = 5 mm.
1595 1596
Figure 7. Sedimentary facies relationships of the Taciba Formation, Capivari County, State of
1597
São Paulo, Brazil. Note the dropstone-bearing shales in the distal areas, and the glacier melting
1598
associated with the cross-stratified sandstones intercalated with plant-rich mudstones. Members
1599
of the Biconvexiella saopauloensis assemblage are recorded in the upper, fossil-bearing silty
1600
shales.
1601 1602
Figure 8. Reconstruction of the hypothesized siliciclastic-dominated, low-energy, shelfal
1603
bottoms of the “Capivari sea”, during a short-lived deglacial event. Note the dominance of
1604
rhynchonelliform brachiopods, with subordinated bivalves, gastropods and crinoids. Explanation:
1605
(1) øPentaridica sp.; (2) Peruvispira brasilensis n. sp.; (3) Quinquenella rionegrensis; (4)
1606
Lyonia rochacamposi; (5) Rhynchopora grossopunctata; (6) Biconvexiella saopauloensis n. sp.;
65
1607
(7) Limipecten capivariensis; (8) Streblopteria aff. S. lagunensis; (9) Phestia tepuelensis and (10)
1608
Praeundulomya cf. subelongata. Illustration by J.L.C. Bezerra.
1609 1610
Table 1. Age of the Biconvexiella saopauloensis assemblage within the glacial succession of the
1611
Itararé Group, Paraná Basin, according to various authors.
1612 1613
Table 2- Correlation of the Itararé Group macroinvertebrate faunas, based on literature data.
1614 1615
Table 3. Faunal composition, suggested age and biocorrelation of the Biconvexiella
1616
saopauloensis assemblage, Taciba Formation, Itararé Group, State of São Paulo, Brazil.
1617 1618
Table 4. Taxonomic composition of the Biconvexiella saopauloensis marine assemblage, Taciba
1619
Formation, Itararé Group, Paraná Basin, southeastern Brazil. Explanation: *species illustrated in
1620
this contribution, but not formally described.
1621 1622
Table 5. Measurements of the brachiopod shells, Biconvexiella saopauloensis assemblage,
1623
Taciba Formation, Itararé Group, Paraná Basin, southeastern Brazil. Explanation: V= ventral
1624
valve; D= dorsal valve; F= fragment; C= conjugated valves.
1625 1626
Table 6. Measurements of the mollusk shells, Biconvexiella saopauloensis assemblage, Taciba
1627
Formation, Itararé Group, Paraná Basin, southeastern Brazil. Explanation: R= right valve; L=
1628
left valve; C= conjoined (articulated) valves; U= undetermined valve.
1629
66
1630
Table 7. Measurements of the crinoid remains, Biconvexiella saopauloensis assemblage, Taciba
1631
Formation, Itararé Group, Paraná Basin, southeastern Brazil. Explanation: *individual discs;
1632
**stem (pluricolumn).
1633 1634
Table 8. Guild diversity and abundance of the marine invertebrates, Biconvexiella saopauloensis
1635
assemblage, Taciba Formation, Itararé Group, Paraná Basin, southeastern Brazil. Guild
1636
classification based on Aberhan and Kiessling (2015).
67
Authors Itararé faunas
Rocha-Campos and Rösler (1978)
Petri and Souza (1993)
Holz et al. (2010)
Simões et al. (2012)
Neves et al. (2014b)
Taboada et al. (2016)
This study
São Paulo
Paraná
Santa Catarina
Artinskian-Kungurian
Kungurian
---
---
---
---
Itaporanga
---
Kungurian
---
---
---
---
Araçoiaba
Late Carboniferous (Stephanian)-Early Permian
Late Carboniferous (WestphalianStephanian)
---
---
---
---
---
Guaraúna
Kungurian
Artinskian
---
---
---
---
---
Ortigueira
Artinskian
Artinskian
---
---
---
---
---
Teixeira Soares
Kungurian
Artinskian
---
Late AsselianSakmarian
Asselian
Latest AsselianEarly Sakmarian
Latest AsselianEarly Sakmarian*
Artinskian-Kungurian
Artinskian
Gzelian-Asselian (C-P boundary)
---
---
---
---
Mafra
Artinskian
Artinskian
---
---
---
---
---
Bela Vista do Sul (Butiá)
Kungurian
Kungurian
---
Late AsselianSakmarian
Late AsselianEarly Sakmarian
Late AsselianEarly Sakmarian
Latest AsselianEarly Sakmarian*
Capivari
Lontras
Latest AsselianEarly Sakmarian
Faunas São Paulo Studies
Capivari
Hortolândia
Mezzalira (1956) Loczy (1964) Rocha-Campos (1969)
X
Paraná Guaraúna
Santa Catarina Ortigueira
Bela Vista do Sul
Itaporanga
Teixeira Soares
X
X
X
X
X
Mafra
X
Saad (1977)
X
X
Rocha-Campos & Rösler (1978)
X
X X
Canuto (1985)
X X
Simões et al. (2012)
Lontras
Rio Grande do Sul
X
X X
Budó
Capivari Marine Invertebrate Fauna Authors
Fossils
Age
Correlation
Barbosa and Almeida (1949a, b) Ambocoelia planoconvexa; Rhynchopora; Aviculopecten sp. gastropods
-------------
Late Paleozoic deposits from Bolivia and Amazon Basin, Brazil
Mendes (1952)
Crurithyris aff. planoconvexa; Rhynchopora grossopunctata Aviculopecten capivariensis; Nuculana ? sp.
Late Carboniferous /Permian
Teixeira Saores, Bela Vista do Sul and Bagé marine beds (=Taciba Formation), Paraná Basin, and Barreal beds, Argentina
Barbosa and Gomes (1958)
RochaCampos (1966)
Peruvispira delicata -------------
Early Pennsylvanian
-------------
Early Permian
Copacabana Formation, Peru
RochaCampos (1967)
as in RochaCampos (1966)
-------------
-------------
RochaCampos (1969) Attenuatella paulistana sp. nov. (nom. nud.) Limipecten capivariensis (nom. nud.);
Older than Teixeira Soares marine fauna (middle portion of the Itararé Group)
Teixeira Soares marine bed (=Taciba Formation), Paraná Basin, Brazil
Saad (1977)
Attenuatella sp.; Rhynchopora grossopunctata Limipecten capivariensis (nom. nud.); Phestia sp.; Streblopteria sp.; Peruvispira delicata; crinoids
Artinskian
Hortolândia shale fauna, Paraná Basin, Brazil
Rocha-Campos and Rösler (1978) Attenuatella sp. nov.; Rhynchopora grossopunctata Phestia sp.; Streblopteria sp.; Peruvispira delicata crinoids
Early Permian (middle portion of the Itararé Group)
Hortolândia shale fauna, Paraná Basin, Brazil
Petri and Souza (1993)
-------------
Artinskian
Teixeira Soares marine bed (=Taciba Formation), Paraná Basin, Brazil
This study
Phestia tepuelensis Streblopteria aff. lagunensis Limipecten capivariensis Praeundulomya cf. subelongata Peruvispira brasiliensis n. sp. Mourlonia (Woolnoughia)? sp. Lyonia rochacamposi Rhynchopora grossopunctata Biconvexiella saopauloensis n. sp. Quinquenella rionegrensis Pentaridica sp.
Latest Asselian-Earliest Sakmarian
Uppermost part of the Taciba Formation (Teixeira Soares and Butiá assemblages)
Bivalvia
Gastropoda
Brachiopoda
Crinoidea
Family
Species
Studied specimens 2
Polidevciidae
Phestia tepuelensis
Streblochondriidae
Streblopteria aff. lagunensis
1
Limipectinidae
Limipecten capivariensis
3
Sanguinolitidae
Praeundulomya cf. subelongata
1
Eotomariidae
Peruvispira brasilensis n. sp.
2
Gosseletinidae
Mourlonia (Woolnoughia)? sp.*
1
Auriculispinidae
Lyonia rochacamposi
2
Rhynchoporidae
Rhynchopora grossopunctata
20
Ambocoeliidae
Biconvexiella saopauloensis n. sp.
86
Rugosochonetidae
Quinquenella rionegrensis
7
øPentacauliscidae
Pentaridica sp.
25
Measurements (mm) Width
Length
Thickness
GP/1E-4342 (C)
19
16
4
GP/1E-443a (V)
14
15
-
GP/1E-443b (V)
~17
~15
-
GP/1E-2507 (V)
~14
~12
-
GP/1E-2545 (V)
~16
~18
~3
GP/1E-451 (D)
18
16
-
GP/1E-1912 (D)
17
~16
-
GP/1E-4325 (D)
14
~13
9
GP/1E-4405 (D)
13
14
-
GP/1E-2527 (V)
~7
~6
-
GP/1E-2504 (V)
~10
9
4
GP/1E-4338 (V)
~6
9
~2
GP/1E-2520 (V)
9
~9
5
GP/1E-4354b (V)
8
9
4
Biconvexiella saopauloensis
GP/1E-4327 (D)
10
8
-
sp. nov
GP/1E-2543 (D)
10
9
-
GP/1E-2538 (D)
~11
~8
-
GP/1E-453 (D)
11
9
-
GP/1E-4339 (D)
11
9
-
GP/1E-2513 (D)
~11
8
-
GP/1E-4358 (D)
5
9
-
GP/1E-459 (V)
10
9
-
GP/1E-4354c (D)
11
8
-
GP/1E-4350a (V)
~13
~22
-
GP/1E-4350b (F)
~35
~26
-
GP/1E-4351a (F)
~31
~15
-
Species
Rhynchopora grossopunctata
Quinquinella rionegrensis
Lyonia rochacamposi
Specimen (valve)
Measurements (mm)
Bivalvia
Gastropoda
Species
Specimen
Length
Height
Width
Phestia tepuelensis
GP/1E-452 (R)
21
11
---
GP/1E-455 (R)
~23
11
---
Streblopteria aff. lagunensis
GP/1E-457 (R)
~21
20
~4
Limipecten capivariensis
GP/1T-116 (L)
~8
~90
---
GP/1E-2511 (U)
~6
~9
---
GP/1E-2494 (R)
9
6
---
Praeundulomya cf. subelongata
GP/1E-5242 (C)
69
27
~13
Peruvispira brasilensis n. sp.
GP/1E-4319
5,3
7,8
---
GP/1E-4348
3,8
~4,5
---
Crinoidea
Specimen
øPentaridica sp.
GP/1E-4321
Major axis (mm) 3,5*
GP/1E-4349 A
2,5 **
GP/1E-4349 B
2,0*
GP/1E-4349 C
1,5*
GP/1E-4350C
~1,2*
Taxon Lyonia Rhynchophora Biconvexiella Quinquenella Streblopteria Limipecten Phestia Praeundulomya
Mourlonia (Woolnoughia) Peruvispira øPentaridica Total
Stationary lowlevel epifaunal suspension feeder 2 20 86 7 1 3 ----------119
Facultatively mobile, infaunal, deposit feeder ------------1 --------1
Facultatively mobile infaunal suspension feeder --------------2 ------2
Epifaunal grazer ----------------1 2 3
Stationary upperlevel epifaunal suspension feeder --------------------25 25
Asselian
Pennsylvanian
303.4 +/- 0.9
Kasimovian 307.2 +/- 1.0
Moscovian 311.7 +/- 1.1
Bashkirian 318.1 +/- 1.3
Itararé Group
298.9 +/- 0.1
Gzheian
Late Carboniferous
Taciba Fm.
EurydesmaLyonia fauna
295.0 +/- 0.1
Tubarão Supergroup
Permian
Cisuralian
Sakmarian
MAH
subzone*
São Paulo Capivarí ass.
Chronostratigraphic Chart Paraná Teixeira Soares ass.
South-Southeast
Rio Grande do Sul
Santa Catarina Butiá BA ass.
Passinho Shale
Rio do Sul Formation
Budó facies
Taciba Guaraúna ass. Araçoiaba ass.
Ortigueira ass.
Sequences
Lontras ass. Mafra ass.
SB-3 Suspiro facies
Lontras shale (Carb./Permian transition)
LPTS -2 Mudstones
Campo Mourão Fm.
Campo Mourão
Sandstones Lapa/ Vila Velha incised valley fill
Lagoa Azul Fm.
Lagoa Azul
“Folhelho Roncador” transgression
Conglomerates Hiatus
SB-2
Incised valley Lithostratigraphic LPTS -1 contact 900 km SB-1 aprox. horiz. scale
Gondwana 1 Supersequence (Milani et al., 2007)
North-Northwest Geochronology Lithostratigraphy Biostratigraphy
A
B
47°30'
47°20'
N
Cuiabá Brasilia
GO
MT
5 km
700
Monte Mor 18º
MG 70
0
900
MS
23°00'
Capivari
1100 1300
1-AB-1-SP
SP
1100
ld
úc
pi
São Paulo
na enti
ad
r
ve
Arg
Ro
Ri 23°10'
ad
lto
e
N
Rio do Sul Lontras Shale
500
Atlantic Ocean 24ºS
Sa
Curitiba
t Tie
Teixeira Soares Passinho Shale
700
Ro
ri-
ar
va
Paraguay
Aç
Ca
PR
900
O
Capivari
Capivari Beds
Salto
SC
300
200 km
Brazil
RS
0
10
0
10
Paraná Basin 54º
Porto Feliz
Porto Alegre
50ºW
Basin border Isopachs (m) Itararé Group outcrop belt Exploration well
Intrusive rocks
Tatuí Fm.
Corumbataí Fm.
Itararé Group
Irati Fm.
Itu Suit
Cities Outcrop Roads
Secondary Road Rivers
25 m
A
A 20
B
B 15
C
C 10
Dark shale with dropstones
5
Shale
Flame structure Plant remains
Silt Shale
Brachiopods
Massive mudstone
Bivalves
Pebbly mudstone
Gastropods
Siltstone
Crinoids
Pebbly silty sandstone Massive sandstone Current-rippled sandstone
0
C S FS MS CS
Trough cross-stratified sandstone
1
2
3
4
5
6
7
8
9
10
11
12
13
14
16
17
18
20
19
26
25
23
21
15
22
24
30 27
28
29
31
32
1
2
4
5
3
6
t ea tr re e Ic Silty shale Cross-stratified sandstone Massive mudstone Shale with dropstones Vegetation
1
2
4
3
5
7
6
8
11
10
9
12
Simões et al., JSAES, CRediT author’s statements
Title: Mollusks and brachiopods of the Capivari Marine Bed, late Paleozoic glacial Itararé Group, northeast Paraná Basin, Brazil: paleoenvironmental and paleogeographic implications Simões Marcello: Supervision; conceptualization; writing (with input from all authors); field collecting (geological and paleontological); funding acquisition; Neves Jaqueline: General systematic analysis; data curation; writing (with input from all authors); figure design and visualization; Taboada Arturo: Conceptualization; general systematic analysis; data curation, writing (with input from all authors); Pagani Maria: General systematic analysis; data curation; writing (with input from all authors); Varejão Filipe: Field collecting (geological and paleontological) data; geological interpretations; figure design and visualization; writing (review and editing); Assine Mário: Field collecting (geological and paleontological) data; geological interpretations; writing (review and editing).