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Epidermal Morphology of the Cordaitalean Leaf Noeggerathiopsis Brasiliensis Nom. Nov. from the Southern Paraná Basin

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Epidermal Morphology of the Cordaitalean Leaf Noeggerathiopsis Brasiliensis Nom. Nov. from the Southern Paraná Basin SILEIR RA A D B E E G D E A O D L E O I G C I A O ARTICLE BJGEO S DOI: 10.1590/2317-4889201920190020 Brazilian Journal of Geology D ESDE 1946 Epidermal morphology of the cordaitalean leaf Noeggerathiopsis brasiliensis nom. nov. from the southern Paraná Basin (Lower Permian, Rio Bonito Formation) and paleoenvironmental considerations Isabela Degani-Schmidt1* , Margot Guerra-Sommer1 Abstract A Brazilian cordaitalean leaf is reassigned and redescribed based on cuticular analysis. The new nameNoeggerathiopsis brasiliensis is erected as a replacement for Rufloria gondwanensis Guerra-Sommer. The leaf adpressions come from a tonstein layer within a Sakmarian coal seam of the Faxinal Coalfield, southern Paraná Basin, Brazil (U–Pb SHRIMP mean age 291 ± 1.3 Ma, Rio Bonito Formation). The leaf micromor- phological pattern, observed under optical transmitted light and field emission gun — scanning electron microscopy, is distinct from the Euramerian Cordaites Unger and very similar to the Angaran Rufloria Meyen. The lower epidermis has conspicuous longitudinal stomatiferous furrows typical of Noeggerathiopsis (Feistmantel) McLoughlin & Drinnan covered by tightly interwoven trichomes, epidermal and probably also hypodermal highly cutinized sheaths at the borders of furrows, and haplocheilic stomatal apparati surrounded by long conical trichomes. The xeromorphic cuticle structures are similar to those ofGlossopteris pubescens (Guerra-Sommer) Degani-Schmidt & Guerra-Sommer, the dominant leaf species preserved in the tonstein layer, indicating convergent ecological adaptations to abiotic stress factors in distinct plant groups in the peat-forming paleoenvironment. KEYWORDS: Gondwana; Cordaitales; Cuticular analysis; Trichomes; Paleozoic; Fossil plants. INTRODUCTION the diagnostic reproductive structures and vegetative organs. Cordaitalean plants, represented by leaves, woods and This also hampers an understanding of their phylogenetic rela- reproductive structures have been recorded from plant assem- tionships with Euroamerian forms. blages in various geological provinces from the Mississippian Since leaves are the most common cordaitalean fossils, var- to the Permian. These plants were important components of ious analyses based on gross morphology, venation, epidermal floristic associations in late Paleozoic mires and are very well and anatomical characters have been undertaken to determine documented in paleotropical Europe and North America shared characters between the morphogenera identified in dif- whence several cordaitaleans have been reconstructed as ferent paleogeographic provinces. whole plants. Although most examples have been reported The characteristics of EuramerianCordaites Unger leaves are from upland and lowland peat mires, others have been linked generally well known based on anatomical details of material to mineral soils and even coastal settings (Tayloret al . 2009, from coal balls and on epidermal patterns identified in compres- and citations therein). sions (Harms & Leisman 1961, Good & Taylor 1970, Rothwell Cordaitaleans were less common in sub-tropical Cathaysia, & Warner 1984) and more recently, on Fourier-transform infra- but some diagnostic features found in whole reconstructed red spectroscopy (FTIR) and elemental analyses (Zodrow plants (e.g., Shanxioxylon) have provided important milestones et al. 2000a, 2000b, and citations therein). The epidermis of in understanding their phylogenetic and evolutionary history Cordaites is characterized by stomata arranged in stomatifer- (Shi-Jun et al. 2003, Hilton & Bateman 2006, Hilton et al. 2009). ous bands between the veins, and sclerenchymatous strands The habit and paleoecology of cordaitaleans in Angaran on the lower surface. Papillae are very rare, and trichomes have and Gondwanan Paleotemperate Realm remain poorly under- not been found (Šimůnek 2007, Šimůnek et al. 2009). This set stood, mainly due to the absence of direct linkages between of leaf characteristics differs from those recognized in many Gondwanan and Angaran leaves. The present study deals with Noeggerathiopsis (Feistmantel) 1Programa de Pós-Graduação em Geociências, Universidade Federal do Rio McLoughlin & Drinnan, a leaf morphogenus of uncertain Grande do Sul – Porto Alegre (RS), Brazil. E-mails: [email protected], [email protected] although presumed cordaitalean affinities owing to the absence *Corresponding author. of associated fertile organs (McLoughlin & Drinnan 1996). Noeggerathiopsis has been recorded from the Mississippian to © 2019 The autors. This is an open access article distributed under late Permian of the Southern Hemisphere, albeit sometimes the terms of the Creative Commons license. placed under Cordaites. Leaves are linear to spathulate, elongate 1 Braz. J. Geol. (2019), 49(2): e20190020 and with sub-parallel venation (Feistmantel 1879, McLoughlin purposes (Guerra-Sommer 1988, 1989, 1992, Cazzulo-Klepzig 1992). Maithy (1964) emended the generic diagnosis adding et al. 2007, 2009, Guerra-Sommer et al. 2008, Simas et al. epidermal patterns, which were not included in the original 2013, Degani-Schmidt et al. 2011, Degani-Schmidt & Guerra- description based on impressions. Sommer 2016a, 2016b, Jasper et al. 2011, Boardman et al. 2012, The diagnostic characteristics ofNoeggerathiopsis distin- Degani-Schmidt et al. 2015). guishing it from Cordaites were, according to Pant & Verma This study aims to correct, by means of cuticular anal- (1964), simple veins of uniform thickness throughout the ysis, the taxonomic identity of the cordaitalean leaf species lamina, ill-defined files of stomata along the stomatiferous from the tonstein layer of the Faxinal Coalfield previously bands and absence of interstitial fibers. Anatomical analy- described by Guerra-Sommer (1989) as Rufloria gondwanensis sis was decisive in understanding the leaf organization, con- Guerra-Sommer, renaming it and contributing to the under- firming the lack of fiber bundles in the leaf structure, leading standing of leaf micromorphology and diversity of cordaital- McLoughlin & Drinnan (1996) to propose an emended diag- ean plants from Gondwana. nosis for the morphogenus so far characterized by discernible stomatiferous furrows in the lower epidermis, protected by densely arranged trichomes. GEOLOGICAL AND The probable presence of northern cordaitalean elements PALEONTOLOGICAL CONTEXT in the Gondwanan floras was supported by the identification The Paraná Basin is a large (1,500,000 km2) intracratonic sag of Cordaites bifolius Césari & Hünicken (2013), which shares basin covering part of southern Brazil, Paraguay, Uruguay and features with Cordaites (e.g., heterophylly, needle-like leaves, Argentina (Fig. 1). Basin subsidence, in addition to Paleozoic leaf scars similar to Cordaicladus Grand’Eury, fertile organs sea-level changes, created six second-order sequences depos- similar to Cordaitanthus Feistmantel bearing Cordaicarpus-type ited from the Ordovician to the late Cretaceous, separated by of seeds). The occurrence of GondwananCordaites was pre- regional unconformities (Milani et al. 2007). The coal-bearing viously reported by Archangelsky & Leguizamón (1980), Rio Bonito Formation, which is the subject of this study, occurs Archangelsky & Cúneo (1991), Chandra & Srivastava (1991) at the base of the second order Carboniferous — Lower Triassic and Srivastava (1996) based only on leaf gross morphology sequence (Gondwana I Supersequence). Extensive peat swamps and venation patterns. The absence of anatomic and epider- developed during the deposition of Rio Bonito Formation in mal features in the Gondwanan type material of both Cordaites Paraná Basin at a paleolatitude of approximately 50°S, included and Noeggerathiopsis preserved as impressions precludes in the cool temperate biome (Rees 2002). Most of the peat-form- close comparisons. ing areas have been identified as part of a back-barrier lagoonal The epidermal patterns of GondwananNoeggerathiopsis paleoenvironment (Holz 2003). compressions are very similar to those of the Angaran Rufloria The study area comprises the Faxinal Coalfield, a former Meyen. Morphological similarities were attributed to homoplasy opencast coal mine closed in 2009, in the southeastern out- by Meyen (1970, 1979, 1984), though a floristic interchange crop belt of Rio Bonito Formation in southern Brazil, Rio via disjunct landmasses in the Tethyan — East Asian region Grande do Sul State. This coalfield overlies the Precambrian during the Carboniferous — was suggested by McLoughlin basement rock (Fig. 1). & Drinnan (1996). The present study focuses on a fossiliferous argillite 7–12 cm In the Brazilian Permian, cordaitalean leaves are known from thick, interbedded in the upper coal seam S. Petrographic stud- interglacial, periglacial and post-glacial deposits from the late ies revealed that this layer represents a tonstein (Simas et al. Carboniferous (Pennsylvanian) to the early Permian (Sakmarian 2013). U–Pb SHRIMP dating of zircons from the tonstein — Artinskian) in the Paraná Basin. Impressions of isolated leaf by Simas et al. (2012) yielded a mean age of 291 ± 1.3 Ma fragments were identified either as Noeggerathiopsis (e.g., Millan (Sakmarian), corresponding to the Protohaploxypinus goraiensis 1972, 1987, Millan et al. 1982, Zampirolli 2001, Bernardes- Subzone within the palynostratigraphic framework defined by de-Oliveira et al. 2016) or Cordaites (Millan & Dolianiti 1981, Souza & Marques-Toigo (2005) and to the phytostratigraphic Millan et al., 1982, Guerra-Sommer & Cazzulo-Klepzig
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