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Selected spores and zygospores. 1, Gleicheniidites senonicus, CR.P.CV CH1 15/105; 2, Baculatisporites comaumensis, CR.P.CV CH1d 21/98; 3, Ceratosporites equalis, CR.P.CV CH14 20/90; 4, Gabonisporis vigourouxii, CR.P.CV CH1c 4/94; 5, Gabonisporis vigourouxii, CR.P.CV CH1c 10/94; 6, Cicatricosisporites sp. 1, CR.P.CV CH1d 18/95.5; 7, Cicatricosisporites sp. 2, CR.P.CV CH1d 10/95; 8, Ruffordiaspora sp., CR.P.CV CH2a 15/100; 9, Zlivisporis reticulatus, CR.P.CV CH14-1 9/85; 10, Zlivisporis reticulatus, CR.P.CV CH2a 22/105; 11, Catinipollis geiseltalensis, CR.P.CV CH1e 15/105; 12, Catinipollis geiseltalensis, CR.P.CV CH1b 5/93; 13, Azolla sp.1, microspore massula with septate glochidia, CR.P.CV CH2b 16/109; 14, Azolla sp. 2, microspore massula fragment, CR.P.CV CH2c 22/102; 15, Azolla, megaspore apparatus, CR.P.CV CH14m 11/92; 16, Ariadnaesporites micromedusus, microspore, CR.P.CV CHo 18/96.5. Scale bars= 10 μm.
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This paper reports on the biostratigraphic, paleobiogeographic and paleoecologic aspects of a palynoflora recovered from sedimentary rocks preliminarily assigned to the Lago Colhué Huapi Formation, in the Golfo San Jorge Basin. The present palynological assemblage includes Maastrichtian marker species of the austral Proteacidites/Nothofagidites Pro...
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Several Upper Cretaceous plesiosaur specimens recovered from southernmost Chile are described here. These were collected from upper levels of the Dorotea Formation exposed on three different localities (Sierra Baguales, Cerro Castillo, and Dumestre). The new material includes the first record of Aristonectes (Plesiosauria, Elasmosauridae), previous...
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... These are significant in supporting a (para)autochthonous evolutionary history for all extant genera of this family in South America once the family reached there from Africa (see below). While the work of Vallati et al. (2016), yielding six named Proteaceae pollen in Central Patagonia ~70 Ma, is too recent to provide migratory insights, it does indicate that this family had diversified greatly in the most southern part of South America by this time. On this point, there may be significance in their record of Tricolpites sp in the Albian?-Cenomanian (112-93.6 Ma) (Vallati, 2006) that has the diagnostic features of Peninsulapollis gillii (earlier classified as Tricolpites gillii) which they recorded later at the same site (Vallati et al., 2016). ...
... While the work of Vallati et al. (2016), yielding six named Proteaceae pollen in Central Patagonia ~70 Ma, is too recent to provide migratory insights, it does indicate that this family had diversified greatly in the most southern part of South America by this time. On this point, there may be significance in their record of Tricolpites sp in the Albian?-Cenomanian (112-93.6 Ma) (Vallati, 2006) that has the diagnostic features of Peninsulapollis gillii (earlier classified as Tricolpites gillii) which they recorded later at the same site (Vallati et al., 2016). If our identification is confirmed it means that the origin of Beauprea would need to be changed from Antarctica to S South America and its age increased by at least 5 My to 94 Ma (He, Lamont and Fogliani, 2016). ...
... De acuerdo con el mismo autor, el inicio de la compresión andina en el Cretácico tardío produjo una extensa cuenca de antepaís y controló la subsidencia de carga tectónica de los depósitos continentales del Grupo Chubut (Lesta y Ferello, 1972), ya sin vinculación con el Océano Pacífico (Paredes et al., 2016). El Grupo Chubut representa gran parte de la sedimentación continental desde el Barreamiano (Vallati, 2013) hasta el Maastrichtiano Vallati et al., 2016) de la cuenca del Golfo San Jorge ( Fig. 1.2). Entre las unidades fluviales y lacustres que lo integran, adquiere especial interés paleontológico y bioestratigráfico la Formación Bajo Barreal (Teruggi y Rosetto, 1963) cuya edad, en base a dataciones radiométricas y el registro paleontológico, se asigna al Cenomaniano temprano-Turoniano tardío . ...
Se analizan los factores intrínsecos y extrínsecos que controlaron la preservación del saurópodo rebaquisáurido Katepensaurus goicoecheai, procedente de la Formación Bajo Barreal (Grupo Chubut-Cretácico Superior) en el centro-sur de Chubut, cuenca del Golfo San Jorge, Patagonia central, Argentina. Los restos se preservaron en un depósito de desbordamiento no canalizado, de geometría lobulada, de 30 cm de potencia y vinculado a canales fluviales arenosos. Litológicamente está integrado por areniscas gruesas con abundante matriz pelítica y tobácea,que permiten caracterizarlo como un flujo hiperconcentrado. Los elementos recuperados, que corresponden principalmente al esqueleto axial, yacían sobre depósitos pelíticos de una planicie de inundación distal mal drenada y fueron parcialmente cubiertos por el lóbulo de desbordamiento, quedando estructuras expuestas que fueron intensamente erosionadas. Los restos se encontraron incompletos, desarticulados y exhibiendo evidencias de una exposición subaérea prolongada. Fueron poco movilizados por procesos hidráulicos y reorientados de forma normal a la dirección del desbordamiento. La dispersión lateral muestra grupos de huesos producto de entrampamientos hidráulicos causados por los restos de mayor tamaño en la parte central del lóbulo y los más pequeños hacia los laterales acorde a la distribución de energía del depósito. La escasa potencia del desbordamiento no sepultó completamente los restos. Finalmente, un nuevo desbordamiento cubrió completamente los materiales. La acumulación de huesos de Katepensaurus sería una concentración ósea residual y representa una acumulación parautóctona de origen sedimentológico.
... However, most of these specimens were described and/or illustrated with informal names or were assigned to species previously reported in Cretaceous and Cenozoic deposits of Australia and New Zealand. Some of them are apparently conspecific with P. baibianae, for example those reported from Maastrichtian deposits of Chubut Province as P. scaboratus (Vallati et al., 2016(Vallati et al., , 2020. However, SEM studies are required to ensure the correct assignment of most of the Cretaceous and Tertiary proteaceous Patagonian species. ...
Abstract
There are few studies focused on spore and pollen clumps in paleopalynological samples, and these are only reports from the Northern Hemisphere. These aggregates may be of animal or foral origin. The goal of this contribution is to provide the frst detailed study of spore and pollen clumps from the Southern Hemisphere, and to discuss their possible origin, botanical afnities, and pollination modes, based on their morphological characteristics, preservation and comparison with putative liv-
ing representatives. Three fern spore clumps and 18 angiosperm pollen clumps were recognized in Maastrichtian-Danian La Colonia Formation sediments that outcrop at Chubut Province, Patagonia, Argentina. Most clumps are monospecifc and com-
posed of undamaged elements but some of them have two types of spore/pollen with corroded and/or fragmented exines. These fndings represent the most diverse and abundant record of fern and angiosperm clumps from the Late Cretaceous and early
Paleocene so far known from South America and the Southern Hemisphere. These results are indicative of the need for comprehensive large-scale studies on pollination of modern taxa and careful processing of palynological samples to lessen the
already large bias in paleopalynological interpretations. The paucity of information on clumps in the fossil record has impaired our comprehension of dispersion/pollination in deep time.
... Analysis was carried out with the PAST software Package (Hammer et al., 2001). The data matrix was constructed using previously published palynological data from the Late Cretaceous of Argentina (Freile, 1972;Archangelsky, 1973;Romero, 1973;Archangelsky and Romero, 1974;Archangelsky and Zamaloa, 1986;Baldoni, 1992;Papú, 1988aPapú, , 1988bPapú, , 1989Papú, , 1990Papú, , 1993Papú, , 1997Papú, , 2002Papú et al., 1988;Sepúlveda et al., 1989;Baldoni and Askin, 1993;Archangelsky et al., 1994;Zamaloa and Andreis, 1995;Papú and Sepúlveda, 1995;Zamaloa and Andreis, 1995;Archangelsky et al., 1999;Volkheimer et al., 2007;Povilauskas et al., 2008;Scafati et al., 2009;Vallati, 2010;Povilauskas, 2011Povilauskas, , 2012Povilauskas, , 2013Povilauskas, , 2016Povilauskas, , 2017Barreda et al., 2012;Clyde et al., 2014Clyde et al., , 2021Cúneo et al., 2014;Gallego et al., 2014;Guler et al., 2014;Puebla et al., 2015;Vallati et al., 2016Vallati et al., , 2017Vallati et al., , 2020De Benedetti et al., 2018Hermsen et al., 2019;Santamarina et al., 2020;Perez Loinaze et al., 2021;Perez Pincheira and di Pasquo, 2021), Antarctica (Barreda et al., 2019) and Biozones commonly used in Australia (Helby et al., 1987) and New Zealand (Mildenhall, 1977;Raine, 1984).The resulting data matrix is given in Appendix A. ...
... Scotese, 1991) showing provenance of extra-Patagonian palynological assemblages compared in this work, and main palynological provinces recognized at the end of the Cretaceous (modified from Vajda and Bercovici, 2014). NB: Neuquen Basin, CAB: Cañadon Asfalto Basin, GSJB: Golfo San Jorge Basin, MB: Malvinas Basin, 1: Bajo de la Carpa Formation (Vallati, 2010), 2: Loncoche Formation (Papú, 1990(Papú, , 1997(Papú, , 2002Papú et al., 1988;Sepúlveda et al., 1989;Puebla et al., 2015), 3: Jagüel Formation (Perez Pincheira and di Pasquo, 2021), 4: Allen Formation (Vallati, 2010), 5: Los Alamitos Formation (Papú and Sepúlveda, 1995), 6: Angostura Colorada Formation (Papú et al., 1988;Sepúlveda et al., 1989), 7: La Colonia Formation (Archangelsky et al., 1999;Cúneo et al., 2014;Guler et al., 2014;Gallego et al., 2014;De Benedetti et al., 2018Clyde et al., 2021), 8: Paso del Sapo Formation (Papú, 1988a, 1988b), 9: Lefipán Formation (Baldoni, 1992Baldoni and Askin, 1993;Barreda et al., 2012), 10: Bororó and Salamanca formations (Archangelsky, 1973;Archangelsky and Romero, 1974;Archangelsky and Zamaloa, 1986;Volkheimer et al., 2007;Scafati et al., 2009;Zamaloa and Andreis, 1995;Clyde et al., 2014;Hermsen et al., 2019), 11: Colhué Huapi Formation (Vallati et al., 2016(Vallati et al., , 2017(Vallati et al., , 2020, 12: Cañadón Seco Formation (Archangelsky et al., 1994;Perez Loinaze et al., 2021), 13: La Anita and La Irene formations (Povilauskas et al., 2008;Santamarina et al., 2020), 14: Cerro Cazador, Monte Chico and Cerro Dorotea formations (Freile, 1972;Povilauskas, 2011Povilauskas, , 2012Povilauskas, , 2013Povilauskas, , 2016Povilauskas, , 2017 This species was originally described by Stough (1968) from two samples obtained from continental deposits outcropping between the Lago Argentino and Lago Viedma (Santa Cruz province, Argentina), which the author assigned to the mid-Cenomanian Mata Amarilla Formation . However, recent studies of these continental deposits have clarified the stratigraphy of this area and it is now evident that the samples analyzed by Stough (1968) were obtained from the Campanian-Maastrichtian Cerro Fortaleza Formation (e.g., Sickmann et al., 2018;Ghiglione et al., 2021). ...
... Nevertheless, this species reaches the Maastrichtian in Australia and Antarctica (Dettmann, 1989(Dettmann, , 1994. Gabonisporis vigorouxii, originally described by Boltenhagen, 1967 from Upper Cretaceous strata of Gabon, was reported also from Coniacian-Maastrichtian units of Brazil (Regali et al., 1974;Ashraf and Stinnesbeck, 1988) and Santonian-Danian of Argentina (Vallati, 2010;Volkheimer et al., 2007;Vallati et al., 2016) (Fig. 4). Gabonisporis cristatus presents a wide record from upper Campanian to Eocene strata in the Northern Hemisphere . ...
... ;Arai and Dias-Brito, 2018; among others). Also sparsely recorded from the Maastrichtian-Danian of southern Argentina(Volkheimer et al., 2007;Vallati et al., 2016, ...
We compiled the fossil record of the heterosporous water ferns (Salviniales) including macro- and microfossils from South America and the Antarctic Peninsula. Both extant families, Marsileaceae and Salviniaceae, are well represented and several fossil spore genera that cannot be placed within the extant families are included as Incertae sedis. Marsileaceae is first recorded in the Middle to Late Jurassic. Incertae sedis genera are recorded for first time in the Early Cretaceous while Salviniaceae in the Late Cretaceous. Two diversity spikes are recognized: one spanned the Aptian–Albian (Early Cretaceous) and is linked to an increase in species diversity within Marsileaceae; and the other occurred during the Campanian–Maastrichtian (Late Cretaceous) and it is associated with the diversification at generic level of the entire Salviniales. Two decreases in diversity are recognized: one during the Cenomanian–Santonian and affected Marsileaceae at specific level but not its generic diversity, and the second is related to the Cretaceous–Paleogene mass extinction and affected all Salviniales at generic and specific levels. From the Paleocene onwards there is a steady decline in the fossil record of the group, with most remains belonging to the extant genera. This study suggests that Southern Hemisphere aquatic ferns underwent a sharp radiation during the Late Cretaceous and a deep decline in the Paleocene that parallels that occurred in the Northern Hemisphere. Probably, the temporal co-occurrence of geological and climate events and the availability of suitable lineages promoted the evolutionary changes of the water ferns in the studied area, including the high diversification and distribution in the Cretaceous and the subsequent post-Maastrichtian decline.
... Zippi (1998) related the presence of zygospores with repeated cycles of arid conditions as well as moderate to higher warm temperatures that triggered their germination (Zavattieri and Pr amparo, 2006;Mautino, 2007). The marine phytoplankton is characteristic of shallow waters (Wrenn and Hart, 1988;van Geel and Grenfell, 1996;Guy-Ohlson, 1996). The low frequency of the latter forms suggests deposition in a shallow brackish to marginal marine environment with low energy and terrestrial input from lowlands and freshwater bodies. ...
... Species of Nothofagidites and most of the gymnospermic pollen grains of Podocarpidites, Microcachryidites antarcticus and Phyllocladidites mawsonii, which are especially typical of the N/P Province, are not documented at Cerro Azul. Therefore, characteristic species of Proxapertites, Longapertites and Spinozonocolpites of the Paleotropical Palmae Province, which are documented in our Danian A2-3 and MaastrichtianePaleocene palynofloras of northern Patagonia (Volkheimer et al., 2007;Scafati et al., 2009;Vallati, 2010;Povilauskas, 2013;Vallati et al., 2016), confirm that they belong to the Mixed Paleofloral Province (Fig. 12). This is in agreement with Romero (1986), Vajda-Santivanez (1999), Vajda and Bercovici (2012) and Vallati (2013). ...
The palynology of the Jagüel Formation at Cerro Azul locality Río Negro province; in the Neuquén Basin, was studied. The distribution of palynomorphs (spores, pollen grains, chlorophytes, and miscellaneous forms) across the outcrops allowed the definition of three palynological associations: A1 (upper Maastrichtian), A2 and A3 (Danian). The palynological results, together with micro- and invertebrate fossils found in this locality contributed to the paleoenvironmental reconstruction of this unit. A1 is deposited in mixed (coastal) to shallow marine environments with low energy associated to freshwater bodies. Whereas it is interpreted that A2 and A3 are deposited in coastal lagoons near shallow and internal platform marine environments. The largest number of palm pollen grains of Proxapertites and other angiosperms Retitrescolpites baculatus and Striatopollis for the Danian of Argentina has been registered here. These associations shared species with Maastrichtian and Danian palynofloras of the Colorado and western Neuquén basins. Species with affinities from northern latitudes of South America and Africa, and a few other species distributed more widely (cosmopolitan) and from southern regions (Argentina, Chile, Antarctica, New Zealand) predominate in the Danian. Therefore, we can confirm that the palynofloras of the Jagüel Formation are part of the Mixed Floristic Realm.
... Therefore, the materials described herein are the first hadrosaurid fossils from the Golfo San Jorge Basin to be associated with precise geographic and stratigraphic information, which in turn casts light on the biostratigraphic and paleobiogeographic range and distribution of South American hadrosaurids during the latest Cretaceous. Furthermore, UNPSJB-PV 1050 and UNPSJB-PV 1061 were associated with palynomorphs and fossil leaves (Vallati et al., 2016 as well as freshwater stromatolites . Therefore, the paleoflora Vallati et al., 2017) and paleoenvironmental setting of the remains are well known, which in turn supports inferences regarding the association of hadrosaurids with coastal, marine-influenced paleoenvironments and how these environmental preferences shaped the distribution of these dinosaurs. ...
... A newly discovered outcrop of the Lago Colhué Huapi Formation at the headwaters of the Río Chico, informally named Cerro del Hadro, has yielded, to date, an association of fossils including stromatolites , palynomorphs (Vallati et al., 2016, wood fragments, indeterminate eggshells, recently recovered non-avian theropod teeth (BNA pers. obs.), and the hadrosaurid remains that are the focus of this study. ...
... 1G (Casal et al., 2020: Table 1; Vallati et al., 2020)]. Vallati et al. (2016) studied the palynoflora of these levels and reported the presence of Maastrichtian species that became extinct at the Cretaceous/Paleogene boundary (Vajda and Bercovici, 2014). Among them, Quadraplanus brossus and Tubulifloridites lilliei characterize the austral Nothofagidites/Proteacidites Province (Herngreen, 1980), whereas Buttinia andreevi and Gabonisporis vigourouxii typify the tropical to subtropical Palmae Province (Herngreen, 1980). ...
We describe and incorporate fragmentary new cranial and postcranial materials of hadrosaurid ornithopods into the non-avian dinosaur assemblage of the Upper Cretaceous (Coniacian–Maastrichtian) Lago Colhué Huapi Formation of central Patagonia, south-central Chubut Province, Argentina. The fossils come from the upper part of the formation, probably from a stratigraphic interval close to the Cretaceous/Paleogene boundary. The materials belong to at least two ontogenetically distinct individuals that are assigned to Hadrosauridae due to their possession of anatomical features that characterize this derived ornithopod group. Sedimentological inferences reveal that the paleoenvironment of these hadrosaurids was characterized by high-sinuosity, meandering-type fluvial channels, whereas palynological data suggest the existence of low-energy freshwater bodies in the floodplains of these rivers and a warm and humid paleoclimate. The deposition of the upper part of the Lago Colhué Huapi Formation was probably influenced by the Patagonian Atlantic marine transgression, which may have played an important role in the distribution of this sedimentary unit. The new Lago Colhué Huapi Formation hadrosaurid specimens constitute the most stratigraphically recent records of this clade from Argentina. The materials contribute to the interpretation of Upper Cretaceous hadrosaurid paleobiogeography in Patagonian basins; moreover, sedimentological and palynological data suggest that paleoenvironmental conditions may have exerted an important influence on South American hadrosaur distribution, supporting hypotheses of ornithopod faunal turnover during the Late Cretaceous of central Patagonia. Finally, the new remains add to the generally depauperate record of Late Cretaceous ornithopods in the Southern Hemisphere.
... Thereby, the Laguna Palacios Formation and the Lago Colhué Huapi Formation (restricted to the western domain) represent the last fluvial infill of the Chubut Group to the Maastrichtian (e.g. Genise et al., 2007;Casal et al., 2015;Vallati et al., 2016). They are unconformably covered by Paleogene successions stratigraphically arranged as follows: Danianlower Paleocene marine Salamanca Formation restricted to the east due to the existence of positive reliefs at deposition time (e.g. ...
First low-temperature thermochronology data across the central Patagonia and thermal modeling provide information on the thermal history of the intraplate San Bernardo fold and thrust belt (44-46 • S). Apatite (U-Th-Sm)/He (AHe), fission tracks (AFT) ages and inverse thermal modeling indicate that the sedimentary rocks presently at the surface of the southwestern San Bernardo FTB have experienced a broadly similar thermal history: i) cooling at the deposition time during the Cretaceous, ii) a significant Eocene-early Miocene thermal event which totally reset the AHe data and partially the AFT data, and iii) a cooling since the early Miocene. Although former thermochronology studies in surrounding areas do not evidence any Neogene thermal event, our regionally consistent ages and thermal modeling most likely indicate a large-scale thermal event through the Eocene to the early Miocene. As sedimentary burial was not significant in this area through the Cenozoic, we propose that this heating episode might have been caused by coeval widespread and long-term intraplate volcanic processes associated with a modest burial and the increase of the geothermal gradient in the entire area, which has been strong enough to significantly affect the low-temperature thermochronometers regionally. A subsequent slow cooling phase starting from early Miocene may have been related with the termination of this intraplate volcanic phase and subsequent decrease of the thermal gradient to average values, and with a modest exhumation of ~1 km at most related to a mild deformation episode of the broken foreland and dynamic topography processes during the middle-late Miocene, as well as the weathering of the Oligo-Miocene basalts.
... ;Arai and Dias-Brito, 2018; among others). Also sparsely recorded from the Maastrichtian-Danian of southern Argentina(Volkheimer et al., 2007;Vallati et al., 2016, ...
We describe a water fern spore assemblage from the Maastrichtian La Colonia Formation, Chubut Province, Patagonia, Argentina. The assemblage includes three species of Azolla (A. andreisii sp. nov., A. coloniensis, and A. sp. 1), two species of Azollopsis (A. intermedia and A. tomentosa), Crybelosporites pannuceus, Gabonisporis cristata, Ghoshispora sp., Grapnelispora loncochensis, two species of Molaspora (M. lobata and M. reticulata), and Paleoazolla patagonica. A. tomentosa, A. intermedia, G. cristata, and M. reticulata are recorded for the first time in the Southern Hemisphere. Ghoshispora sp. constitutes the southernmost record of the genus and its first mention for Argentina. In addition, the genus Azollopsis is emended to include megaspore apparatuses with more than one megaspore. This water fern spore assemblage is one of the most diverse so far known for South America and substantiates the significant radiation that aquatic ferns underwent worldwide at the end-Cretaceous. Furthermore, it also implies that the presence of widespread suitable paleoenvironments allowed the establishment and rapid evolution of these plant communities in Patagonia.
... The overlying Castillo (Albian) and Bajo Barreal (Cenomanian) Formations mainly register volcaniclastic fluvial systems with abundant tephra influx from the west (Bridge et al., 2000;Umazano et al., 2008Umazano et al., , 2009Umazano et al., , 2012Suárez et al., 2014;Paredes et al., 2015Paredes et al., , 2016Paredes et al., , 2018Genise et al., 2020). Finally, the upper part of the Chubut Group includes a Cenomanian loess-paleosol volcaniclastic succession in the western margin of the basin, which is denominated Laguna Palacios Formation (Sciutto, 1981;Bellosi and Sciutto, 2002;Suárez et al., 2014), and the Coniacian-Maastrichtian fluvial sediments of the Lago Colhue Huapi Formation (Casal et al., 2015;Vallati et al., 2016;. ...
... In Chubut Province, Jurassic records are concentrated in the Cañadón Asfalto Basin (Fig. 6), with abundant cones and Brachyphyllumtype leaves, besides Classopollis pollen grains Olivera et al., 2015;Escapa and Leslie, 2017). Cretaceous records of this province are based mostly on Classopollis pollen (Vallati, 1993(Vallati, , 1996b(Vallati, , 2013Volkheimer et al., 2009;Vallati et al., 2016Vallati et al., , 2019, often associated with drilling studies for hydrocarbons extraction and exploration belonging to Golfo San Jorge Basin (Archangelsky and Seiler, 1980;Archangelsky et al., 1981;Seiler and Moroni, 1984;Jalfin et al., 2002;Barreda et al., 2003Barreda et al., , 2012. Cenozoic studies of Chubut Province have also recorded Classopollis near Bororó, Hansen and Abigarrado Hills Table 4 for references. ...
We studied fossil woods from Matasiete and Castillo Formations (Aptian–Albian) for the first time. The woods, preserved in fluvial channel and floodplain deposits, have anatomy consistent with the fossil-genus Brachyoxylon Hollick and Jeffrey. Specimens from Castillo Formation have poorly distinct growth ring boundaries, exclusively uniseriate with a mixed arrangement intertracheary radial pitting, araucarioid cross-field pitting with 3–8 half-bordered pits with oblique aperture, and uniseriate rays. Specimens from Matasiete Formation are poorly preserved and only one could be assigned to Brachyoxylon with reservations. Brachyoxylon is a fossil-genus with a worldwide distribution and related to the fossil family Cheirolepidiaceae, a group of conifers that developed in various environments. However, our study suggests that this family would have prospered in locally semi-arid conditions.
