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An Upper Paleozoic bio-chronostratigraphic scheme for the western
margin of Gondwana
Silvia N. Césari
a,
⁎, Carlos O. Limarino
b
, Erik L. Gulbranson
c
a
Museo Argentino de Cs. Naturales, “B. Rivadavia”, Av. Ángel Gallardo 470, Buenos Aires 1405, Argentina
b
Universidad de Buenos Aires, Departamento de Geología, Ciudad Universitaria, Pabellón 2, Buenos Aires 1428, Argentina
c
Department of Geology, University of California-Davis, One Shields Avenue, Davis, CA 95616, USA
abstractarticle info
Article history:
Received 26 July 2010
Accepted 31 January 2011
Available online 12 February 2011
Keywords:
Upper Paleozoic
biostratigraphy
radiometric ages
palynozonation
western Argentina
Gondwana
The Carboniferous and Permian fossiliferous sequences of the central-western Argentina contain abundant
plant remains, palynomorphs and invertebrates. They include a continuous record of large distribution in the
Paganzo, Rio Blanco, Calingasta-Uspallata and San Rafael Basins. The most recent biostratigraphic schemes
recognize a floristic succession represented by the biozones: Archaeosigillaria–Frenguellia (AF Biozone),
Frenguellia eximia–Nothorhacopteris kellaybelenensis–Cordaicarpus cesarii (FNC Biozone), Nothorhacopteris–
Botrychiopsis–Ginkgophyllum (NBG Biozone), Interval Biozone and Gangamopteris Biozone. The associated
palynological record is represented by the biozones: Reticulatisporites magnidictyus–Verrucosisporites
quasigobbetti (MQ Biozone), Raistrickia densa–Convolutispora muriornata (DM Biozone), Pakhapites fusus–
Vittatina subsaccata (FS Biozone), and Lueckisporites–Weylandites (LW Biozone). The precise age of the Upper
Paleozoic western Gondwanan biozones has been under discussion and remains controversial to date in some
regions. The main issue hampering an integrated comparison of the Gondwanan biozones was its imprecise
chronostratigraphic framework. However, new studies in some Argentinian stratigraphic sections bearing
floras and faunas have yielded several radiometric ages. From these
206
Pb/
238
U zircon datings it is possible to
determine the chronostratigraphic range of many fossiliferous assemblages in this sector of Gondwana. In
this way, the AF and MQ Biozones are restricted to the Late Mississippian and they would be not younger
than 335 Ma according to radiometric ages.
206
Pb/
238
U ages suggest that the NBG, DMa and DMb Biozones
characterize the Late Serpukhovian glacial deposits and persisted up to the Late Bashkirian. Beds containing
the Interval and DMc Biozones have yielded
206
Pb/
238
U ages of 312.82± 0.11 Ma and 310.71±0.1 Ma which
would indicate that both zones characterize the Moscovian. The remains of Gangamopteris Biozone found in
the Paganzo Basin overlie basalt levels ranging between 308 ± 6 and 293 ± 6 Ma. Therefore, the incoming
of the first glossopterids was closely associated to the Carboniferous–Permian boundary in this part of
Gondwana. The data presented in this paper are used for establishing comparisons with other Gondwanan
biozones, constrained by absolute ages.
© 2011 Elsevier B.V. All rights reserved.
Contents
1. Introduction .............................................................. 150
2. Carboniferous Argentinian biostratigraphy ................................................ 150
2.1. Mississippian .......................................................... 150
2.2. Pennsylvanian ......................................................... 153
2.3. Carboniferous–Permian boundary ................................................ 153
3. Palaeoclimate and vegetational composition ............................................... 154
4. Comparisons with other Gondwanan assemblages ............................................ 155
5. Conclusions .............................................................. 157
Acknowledgments ............................................................. 158
References ................................................................. 158
Earth-Science Reviews 106 (2011) 149–160
⁎Corresponding author. Tel.: +54 11 49826670.
E-mail addresses: scesari@macn.gov.ar (S.N. Césari), limar@gl.fcen.uba.ar (C.O. Limarino).
0012-8252/$ –see front matter © 2011 Elsevier B.V. All rights reserved.
doi:10.1016/j.earscirev.2011.01.012
Contents lists available at ScienceDirect
Earth-Science Reviews
journal homepage: www.elsevier.com/locate/earscirev
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1. Introduction
The fossiliferous sequences from the western area of Argentina
are recognized by their abundant plant remains, palynomorphs and
invertebrates. They comprise a continuous record of wide distribution
in the Paganzo, Rio Blanco, Calingasta-Uspallata and San Rafael Basins
(Archangelsky et al., 1987, 1996; Azcuy et al., 2007; Césari et al.,
2007). In Gondwana there are few regions with a continuous fossil
record from the Early Carboniferous to the Permian. For example,
India, South Africa and Antarctica contain well-documented Permian
sequences and Permian fossil associations in Australia are better
known than the Carboniferous ones. In addition, the scarcity of
absolute ages related to fossil assemblages is a common feature of
Gondwana. This last issue is very important when trying to refer the
biozones to the international time scale, which is calibrated using
fossils that are absent in the Gondwanic endemic biota.
Accompanying the advancement in palaeontological knowledge,
biostratigraphic proposals have been submitted for western Argentina
during the last twenty years. The latest schemes, formally defined,
recognize a Carboniferous–Permian floristic succession represented by
the biozones: Archaeosigillaria–Frenguellia (AF Biozone), Frenguellia
eximia–Nothorhacopteris kellaybelenensis–Cordaicarpus cesarii (FNC
Biozone), Nothorhacopteris–Botrychiopsis–Ginkgophyllum (NBG Bio-
zone), Interval Biozone and Gangamopteris Biozone. The associated
palynological record is represented by the biozones: Reticulatisporites
magnidictyus/Verrucosisporites quasigobbetti (MQ Biozone), Raistrickia
densa–Convolutispora muriornata (DM Biozone), Phakapites fusus–
Vittatina subsaccatta (FS Biozone) and Lueckisporites–Weylandites (LW
Biozone). The marine faunas are referred to diverse biozones in some
cases attributed to dissimilar ages (Cisterna, 2010; Taboada, 2010 and
cites therein). The precise age of the associations has been debated
because of the absence of species having worldwide biochronologic
value. However, recent studies on key sections of the Precordillera
region (Fig. 1) have yielded absolute ages in various strata bearing
some floras and faunas from the Upper Paleozoic (Gulbranson et al.,
2010). From these radiometric ages it is possible to determine the
chronostratigraphic range of many fossiliferous associations in this
sector of Gondwana.
2. Carboniferous Argentinian biostratigraphy
The biostratigraphy of Upper Paleozoic deposits of central western
Argentina has long been based on palynofloras, macrofloras and
marine faunas (e.g. Archangelsky et al., 1987, 1996; Azcuy et al., 2007;
Césari et al., 2007). Palynomorphs are most valuable for biostratig-
raphy in the Gondwana region, as shown in many publications, for
example, by Césari and Gutiérrez (2001), Eyles et al. (2002, 2006),
Holz et al. (2010) and Stephenson (2008); therefore biostratigraphic
discussion in this paper is mainly focused on palynological data. In
this way, the main palynological species belonging to the different
associations recognized in western Argentina are illustrated in Fig. 2.
During nearly forty years the only radiometric ages in the re-
gion were from Thompson and Mitchell (1972),obtainedinbasalts
from La Torre and Paganzo localities (Paganzo Basin). A K–Ar age
range of 293± 6–308 ± 6 Ma was obtained from six samples over-
lying the Carboniferous NBG flora, and underlying the first record
of Gangamopteris leaves (Thompson and Mitchell, 1972; Césari,
2007). These ages were traditionally used for establishing the
Carboniferous–Permian boundary in Argentina.
In order to clarify the stratigraphic location of the fossiliferous
associations and the radiometric data, the most important strati-
graphic sections are shown in Fig. 3, as well as an integrated column
where Mississippian and Pennsylvanian deposits are plotted. Missis-
sippian sequences are represented in western Argentina by the
Angualasto Group (Limarino and Césari, 1993), the Malimán and the
Cortaderas Formations in ascending stratigraphic order are included
as reference units. The Pennsylvanian is classically represented by the
Guandacol, Tupe and Patquía Formations. The Guandacol Formation
possesses glacial and postglacial marine transgressive deposits
(Limarino et al., 2006) that represent the most important glacial
event along the western margin of Gondwana and results in a key
horizon for correlation with equivalent units in the region. Another
transgressive event is recognized in the middle–upper part of the
Tupe Formation characterized by rich faunal associations (Limarino
et al., 2006). The Patquia Formation comprises mainly red beds de-
posits bearing scarce fossils.
The time scale used in this paper follows the global Carboniferous
chronostratigraphic time scale recently calibrated with over 25 high
precision ID-TIMS U–Pb zircon ages from the Donets Basin, Ukraine
(Davydov et al., 2010). It must be emphasized that the time scale is
being continually modified resulting in changes that make difficult to
relate the ages offered in the literature to an updated scale. Therefore,
when it is possible, we try to use the available absolute ages related to
Gondwanic biozones for establishing comparisons.
2.1. Mississippian
Although the presence of Mississippian marine fauna and macro-
floristic remains in the Angualasto Group (Rio Blanco Basin) has
been known since the mid-twentieth Century, the first proposals of
zonation were presented by Sessarego and Césari (1988) and González
(1993). Palynological assemblages from the Angualasto Group were
mentioned by Sessarego and Césari (1988), Césari and Limarino (1992,
1995) and later described and improved by Amenábar et al. (2006,
2007) and Perez Loinaze (2007, 2008). Most recent biostratigraphic
schemes include the Late Tournaisian–Early Visean Protocanites
scalabrini–Azurduya chavelensis (PA) Zone based on invertebrates
(Sabattini et al., 2001), and the floristic Archaeosigillaria–Frenguellia
Biozone (Sessarego and Césari, 1988; Arrondo et al., 1991) from the
Malimán Formation and equivalent units. Although, Carrizo and
Azcuy (2006a) proposed the Gilboaphyton argentinus–Malimaniun
furquei and Frenguellia–Paulophyton Biozones for replacing the AF
Biozone, they have not been defined according to guidelines con-
ventionally established, and some of its diagnostic species have not yet
been described (see Azcuy et al., 2007). Palynological assemblages
Fig. 1. Location map showing the main Upper Paleozoic basins from western Argentina.
Darker areas represent palaeotopographic highs (Limarino et al., 2006). Asterisks show
the locations of stratigraphic sections with radiometric data.
150 S.N. Césari et al. / Earth-Science Reviews 106 (2011) 149–160
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from these levels may be partially referred to the Cordylosporites/
Verrucosisporites Biozone early defined by Césari and Gutiérrez (2001).
The Late Visean MQ palynozone (Perez Loinaze, 2007) was defined for
the overlying Cortaderas Formation, which contains glacial deposits in
its upper section. The faunistic record related to this glacial event
would correspond to the Rugosochonetes–Bulahdelia (RB) marine
fauna (Taboada, 1989, 2010) recorded in the El Paso Formation (San
Juan province).
A
206
Pb/
238
U 335.99 ±0.06 Ma age (2σanalytical uncertainty)
from an andesite in the uppermost Punta del Agua Fm. was obtained
Fig. 2. 1, Verrucosisporites congestus Playford, BAPal 5756, X250; 2, Verrucosisporites cortaderensis Pérez Loinaze, BAPal 5756; X500; 3, Reticulatisporites magnidictyus Playford and
Helby, BA Pal 5760–1, X750: 4, Dibolisporites malimanensis Pérez Loinaze BAPal 5754, X750; 5, Verrucosisporites quasigobbettii Jones and Truswell; BA Pal 5756, X 500;
6, Reticulatisporites asperidictyus Playford and Helby, BAPal 6134, X 500; 7, Foveosporites hortonensis (Playford) Azcuy, BAPal 6134, X500; 8, Raistrickia rotunda Azcuy, BAPal 6134,
X 500; 9, Cristatisporites menendezii (Menéndez and Azcuy) Playford, BAPal 6134, X 500; 10, Plicatipollenites densus Srivastava, BAPal 6134, X300; 11, Ahrensisporites cristatus Playford
and Powis, BAPal 6134, X 300; 12, Convolutispora muriornata Menéndez, BAPal 6134, X 500; 13, Granulatisporites austroamericanus (=Microbaculispora tentula); 14, Spelaeotriletes
ybertii (Marques Toigo) Playford and Powis, BAPal 6134, X300; 15, Protohaploxypinus sp., BAFCPl 229, X500; 16, Apiculatisporis variornatus di Pasquo, Azcuy y Souza, BA Pal 5846,
X500; 17, Lundbladispora braziliensis (Pant y Srivastava) Marques Toigo y Pons, BAPal 6133, X 500; 18, Raistrichia cephalata ; 19, Horriditriletes uruguaiensis ; 20, Protohaploxypinus sp.
BAPal 6133, X 300; 21, Colpisaccites granulosus Archangelsky y Gamerro, BAPal 6131, X250; 22, Hamiapollenites fusiformis Marques Toigo emend. Archangelsky y Gamerro, BAPal
6131, X300; 23, Kraeuselisporites sanluisensis Menéndez, BAPal 6131, X500; 24, Verrucosisporites patelliformis (Menéndez) Gutiérrez y Césari, BAPal 6131, X500; 25, Vittatina
subsaccata Samoilovich emend. Jansonius, BAPal 6131, X 500; 26, Lueckisporites stenotaeniatus Menéndez, 5419, X500; 27, Vittatina fasciolata (Balme y Hennelly) Bharadwaj, 5419,
X500; 28, Weylandites magmus Bose y Kar, 5415, X500; 29, Staurosaccites cordubensis Archangelsky and Gamerro, BAPal 5416, X 250; 30, Lueckisporites latisaccus Archangelsky y
Gamerro, BAPm 5424, X250.
151S.N. Césari et al. / Earth-Science Reviews 106 (2011) 149–160
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by Gulbranson et al. (2010). These authors correlated the upper part
of the Punta del Agua Formation with the diamictites reported from
the Cortaderas Fm. and therefore suggested a mid-late Visean mini-
mum age for the MQ palynozone and the glacial episode. This Late
Visean glacial event was characterized and distinguished by Perez
Loinaze et al. (2010a) from the Serpukhovian glacial episode
registered in the Guandacol Formation. The most significant palyno-
logical difference between both deposits is the appearance of
monosaccate pollen in diamictites of the Guandacol Formation (Césari
and Gutiérrez, 2001; Limarino et al., 2002; Perez Loinaze et al., 2010a).
Recently, Balseiro et al. (2009) described a small but significant
floristic association that referred to a new biozone: Frenguellia eximia–
Nothorhacopteris kellaybelenensis–Cordaicarpus cesarii (FNC). The as-
semblage was found in rocks underlying the Guandacol Formation at
Loma de Los Piojos, near Huaco area (Fig. 3). This is the first record of
an intermediate flora between the AF and NBG assemblages and
Fig. 3. Summary time-space framework for the Carboniferous and Permian of the western Argentina basins. Generalized graphic sedimentological log (left) and detailed logs
of representative formations in different localities (right). The location of the palaeofloristic assemblages and radiometric age control points (green lines) are shown.
152 S.N. Césari et al. / Earth-Science Reviews 106 (2011) 149–160
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would be characterized by the coexistence of lycophytes like
Frenguellia, the pteridosperm Nothorhacopteris and probably cordai-
tales represented by Cordaicarpus among other taxa. The age of this
flora can be constrained as younger than 335 Ma and older than
323 Ma according to Gulbranson et al. (2010). This early Serpukho-
vian age agrees with the assumption of Balseiro et al. (2009) about the
stratigraphic position of the biozone.
2.2. Pennsylvanian
The Guandacol Formation and equivalent units such as the Jejenes
Formation and lower section of the Río del Peñón Formation contain
some imprints of floristic remains of the Nothorhacopteris–Botry-
chiopsis–Ginkgophyllum (NBG) Biozone (Vazquez Nistico and Césari,
1987; Césari and Limarino, 1988; Césari et al., 1988a, 1988b; Gutiérrez
and Pazos, 1994; Gutiérrez et al., 1995). Some of these and other
contemporary stratigraphic units (e.g. Hoyada Verde Formation) are
also characterized by the presence of abundant marine invertebrates
belonging to the Levipustula fauna (Taboada, 1997, 2010). This fauna
is distinguished by the presence of brachiopods, bivalves, bryozoan,
gastropods, together with cnidaria, crinoids and ostracods (see ref-
erences in Césari et al., 2007 and Azcuy et al., 2007). Fossil woods have
also been described associated to these marine deposits (Pujana and
Césari, 2007).
The palynology of the Guandacol Fm. was initially analyzed by
Césari and Vazquez Nístico (1988) and has been improved with the
contributions of Ottone and Azcuy (1989), Ottone (1991) and Césari
and Limarino (2002).Césari and Gutiérrez (2001) referred the paly-
nological assemblages from Guandacol and equivalents formations,
to the Subzone A of the DM Biozone (Namurian in age).
206
Pb/
238
U
319.57 ±0.09 Ma and 318.79 ±0.10 Ma ages (Gulbranson et al., 2010)
from postglacial transgressive facies, immediately above the basal
glacial deposits (Fig. 3), confirm the biostratigraphic age of Subzone
A, and provide a minimum age constraint on this subzone (Late
Serpukhovian–Bashkirian).
Although the well known NBG flora is present in the Guandacol
Formation, it has been mainly characterized in the coal beds of the
lower section of the Tupe Formation and equivalents units (Fig. 3). This
floristic association and the related palynological assemblages were
analyzed in its stratotype by Césari (1984, 1985, 1986a, 1987).
Imprints of Cordaitales, sphenopsids, pteridosperms and lycophytes
are the most common components of the flora (Archangelsky et al.,
1987; Vega and Archangelsky, 2001; Césari and Perez Loinaze, 2005;
Césari et al., 2007), with ancillary wood remains (Brea and Césari,
1995; Césari et al., 2005; Pujana, 2005). The fourteen palynological
associations from different localities, originally referred to the
Subzone B of the DM Biozone by Césari and Gutiérrez (2001), have
been increased with new findings in the last years (i.e. Di Pasquo
et al., 2010). Subzone B is characterized by the presence of
diverse monosaccate pollen and scarce taeniate pollen (usually
Protohaploxypinus) that suggests a Late Carboniferous (Westphalian
in Césari, 1986b) age for the unit. A
206
Pb/
238
U age of 318.79 ±0.10 Ma
obtained at the upper levels of the Guandacol Formation (see Fig. 3;
Gulbranson et al., 2010) constrains the maximum age of Subzone B,
while its upper limit is approximately marked by an age of 315.46 ±
0.07 Ma from tonsteins included in coal beds of floodplain deposits
belonging to the Tupe Formation (Fig. 3;Gulbranson et al., 2010).
Therefore, Subzone B may be constrained to the Late Bashkirian.
The upper transgression of the Tupe Formation contains in-
vertebrates referred to the Tivertonia jachalensis–Streptorhynchus
inaequiornatus (TS) Biozone (Sabattini et al., 1991) originally referred
to the Late Carboniferous. Recently, some authors extended the age of
the fossiliferous levels up to the Early Permian (Cisterna et al., 2002a,b;
Sterren, 2004; Cisterna et al., 2005; Coturel and Gutiérrez, 2005;
Cisterna et al., 2006; Vergel, 2008). Furthermore, Coturel and Gutiérrez
(2005) and Cisterna (2010) proposed the upper section of the Tupe
Formation, containing TS fauna, as the stratotype of the Carbonifer-
ous–Permian boundary. Recently, Desjardins et al. (2009) analyzed in
detail the transgressive deposits correlating the sequences outcrop-
ping in the nearby localities of Mina La Ciénaga and Mina La Delfina.
The fossiliferous assemblages from the latter locality were also
correlated by Cisterna et al. (2006) with those from La Herradura
creek where the TS biozone was defined by Sabattini et al. (1991).
Moreover, Vergel (2008) described in La Herradura creek a palyno-
logical association characterized by 5% of taeniate pollen assigned to
the FS Biozone, obtained from a sample several meters above the strata
bearing the TS fauna. Recent palynological study on the same strata
containing the marine invertebrates allows to refer the assemblages to
the Subzones B and C (Perez Loinaze et al., 2010c).
Radiometric dates (Gulbranson et al., 2010) confirm in part the
Westphalian (=Moscovian)–Stephanian age originally proposed by
Sabattini et al. (1991) for the TS Biozone. A sample from shallow-
marine deposits at Mina La Ciénaga locality (facies association VII by
Desjardins et al., 2009) yields a
206
Pb/
238
U age of 312.82 ±0.11 Ma
(Fig. 3), whereas overlying fluvial deposits of the Patquia Formation
yield a
206
Pb/
238
U age of 310.71 ±0.1 Ma (Fig. 3;Gulbranson et al.,
2010). Both ages discard an Early Permian age for the top of the Tupe
Formation. Another radiometric date that constrains the age range
of the TS Biozone is available from the middle part of the Agua de
Jagüel Formation (Mendoza Province) where Lech (2002) reported a
radiometric age of 307.2 ±5.2 Ma from volcanic rocks related to
marine fauna referred by this author to the TS Biozone (Fig. 3).
The floristic assemblages from this Moscovian stratigraphic
interval, referred by Archangelsky and Cúneo (1991) to the Interval
Biozone, show several compositional changes by the usual presence
of ferns (Pecopteris,Asterotheca) and conifers (Paranocladus and
Kraeuselcladus), some of them anatomically preserved (Crisafulli,
2002). Carrizo and Azcuy (2006b) proposed the Kraeuselcladus–
Asterotheca Biozone for including these floras, but still need to define
this biozone according to guidelines conventionally established.
Palynofloras from this stratigraphic interval were referred to the
Subzone C (DM Biozone) by Césari and Gutiérrez (2001). These as-
semblages did not show significant changes in its composition except
for the recognition of brackish and marine palynomorphs and the
increase of taeniate pollen (Césari and Gutiérrez, 2001; Perez Loinaze
and Césari, 2004; Gutiérrez and Limarino, 2006).
2.3. Carboniferous–Permian boundary
The Argentinian palynological FS Biozone defined by Césari and
Gutiérrez, 2001) was originally referred to the Early Permian in
western Argentina, although Césari (2007,figs. 1, 2) remarked
the uncertainty about the age of its lower boundary which could be
in the latest Carboniferous. It is characterized by the abundance of
bisaccate taeniate (Protohaploxypinus,Vittatina,Hamiapollenites, and
Striatopodocarpites), plicate (Pakhapites and Marsupipollenites) pollen
and some specimens referred to Converrucosisporites confluens. This
biozone was recognized in the isolated outcrops of Bajo de Véliz and
Tasa Cuna localities (Césari et al., 1999; Gutiérrez and Césari, 2000;
Balarino and Gutiérrez, 2006), Los Sauces Formation and the upper
sections of the Santa Máxima, Río del Peñón and El Imperial For-
mations (see Ottone, 1989; García, 1995, 1996; Gutiérrez and
Limarino, 2006; Di Pasquo et al., 2010).
Recently, a
206
Pb/
238
U age of 310.63 ±0.1 Ma was reported by
Gulbranson et al. (2010) from fluvio-deltaic deposits in the middle
part of the Rio del Peñón Formation (Fig. 3), approximately at the
same level where the boundary between the DM and FS Biozones was
suggested (Gutiérrez and Limarino, 2006). Therefore, the FS Biozone
would have begun in the Late Carboniferous if the few diagnostic
specimens (see appendix, Gutiérrez and Limarino, 2006) reported in
the Rio del Peñón Formation are considered valid for the recognition
of the biozone in that place.
153S.N. Césari et al. / Earth-Science Reviews 106 (2011) 149–160
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Besides the abundance of taeniate pollen the former reference to
the Carboniferous–Permian boundary of the FS Biozone was inferred
because some of the assemblages also contain glossopterids leaves. In
addition, the basaltic horizon interbedded in the lower part of the
La Colina Formation were dated between 293 ±6 Ma and 308 ±6 by
K/Ar method (Thompson and Mitchell, 1972) suggesting a maximum
age of 298–301 Ma for the glossopterid remains of this unit (Limarino
and Césari, 1984). Recently Gulbranson et al. (2010) dated zircons
extracted from a rhyolitic tuff yielding an age of 296.09± 0.085 Ma.
The tuff was interbedded with eolian deposits located some tens of
meters above the basaltic rocks (Fig. 3).
Marine environments were developed during the Early Permian
and are characterized by the Costatumulus amosi (ex-Cancrinella
farleyensis) fauna. This biozone was defined and characterized by
Taboada (1998, 2006), who suggested an Asselian–Tastubian age for
the biozone. Recent palynological studies (Perez Loinaze et al., 2010b)
of the same fossiliferous strata confirm the Permian age of the
fauna by the presence of assemblages with abundant taeniate
pollen, including specimens of Lueckisporites,Striatopodocarpites and
Hamiapollenites among others.
The palynological LW Biozone succeeds the FS Biozone and is
characterized by the cleardominance of taeniate pollen grains(48–53%)
including Lueckisporites,Weylandites,Vittatina and Marsupipollenites.
The base of the biozone was defined by the first appearance of
Lueckisporites spp. One of the reference sections of the biozone is the
Yacimiento Los Reyunos Formation, a volcaniclastic unit related to the
Permian Andean volcanism (references in Césari, 2007).
3. Palaeoclimate and vegetational composition
One of the most attractive aspects of the southwestern basins of
Gondwana is the complex palaeoclimatic history that they exhibit
(Lopez Gamundi et al., 1992; Isbell et al., 2003; Limarino et al., 2006).
This fact offers a good opportunity to evaluate if climatic shifts were
intense enough to affect Late Paleozoic floras, and to what degree.
Synthesis of the climatic evolution of Carboniferous and Permian
Argentinian floras have been presented by Lopez Gamundi et al.
(1992) and Limarino et al. (1997) and considerably improved by
López-Gamundi and Martínez (2000), Perez Loinaze et al. (2010a),
Gulbranson et al. (2010), Spalletti et al. (2010) among others.
Presently the palaeoclimatic evolution of this part of Gondwana can
be synthesized following seven major stages: 1. Preglacial climate
(Late Tournaisian–Early Visean), 2. Early glacial event (Late Visean), 3.
Interglacial period (Early Serpukhovian), 4. Late glacial event (Late
Serpukhovian–Early Bashkirian), 5. Humid postglacial climates (Late
Bashkirian), 6. Warming semiarid conditions (Moscovian–Asselian)
and Permian aridization (Middle Permian?).
The preglacial interval, equivalent to palaeoclimatic stage 1 of
Lopez Gamundi et al. (1992), records a large span of time charac-
terized by humid and temperate conditions (Lopez Gamundi et al.,
1992; Limarino et al., 1997). During this time, the flora was dominated
by small and primitive lycophytes belonging to the AF Biozone and
palynological assemblages included in CV and MQ Biozones that
are characterized by the lack of pollen grains (Fig. 4). A dramatic
palaeoclimatic change occurred in the Late Visean when the oldest
deposits of proven glacial origin in Argentina are recognized in
Precordillera (Perez Loinaze et al., 2010a). Despite the fact that
macrofloristic remains have not been found neither in the diamictites
nor in periglacial shales, palynological records of the MQ Biozone
indicate that Cordaitales has not yet appeared taking into account the
lack of monosaccate pollen grains related to this group. Stratigraphic
levels above the diamictites are composed of sandstones and carbo-
naceous mudstones representing the interglacial period. This interval,
likely characterized by temperate and very humid climate, documents
a gradual replacement of the AF flora by the incorporation of new taxa
(e.g. Nothorhacopteris), giving rise to the FNC Biozone (Fig. 4).
In the Late Serpukhovian–Early Bashkirian a new glacial phase
affected all the western basins of Argentina forming glacial diamic-
tites, well-known in the Paganzo, Río Blanco, Calingasta–Uspallata
and San Rafael Basins. At the end of the glaciation, postglacial
conditions resulted in temperate and very humid climate (Fig. 4)
leading to the widespread formation of coal beds in paralic and
fluvial environments. Under this new climate, deep modifications in
the composition of both macro and microfloras have been recorded in
Argentina. In this way palynological associations included in
the NBG Biozone and Subzones A and B of the DM Biozone contain
the first records of pollen grains indicating the progressive coloniza-
tion of upland areas by Cordaites. The nearly synchronous appearance
of monosaccate pollen grains and Cordaites in Gondwana (Césari
and Gutiérrez, 2001) suggests a close relationship between them.
Moreover, small lycophytes were rapidly replaced by bigger ones
including the genera Brasilodendron (Césari, 1982) and Bumbudendron
(Archangelsky et al., 1981; Gutierrez et al., 1986). The Gondwanan
pteridosperms (Nothorhacopteris,Botrychiopsis and Fedekurtzia) as-
sociated with cosmopolitan ones like Eusphenopteris (Césari et al.,
1988a) were becoming more dominant towards the Late Bashkirian.
Fig. 4. Palaeofloristic succession in western Argentinian Basins and palaeoclimatic
events. Radiometric data (♦), see references in Fig. 6.
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As from the Moscovian, a progressive warming and drying in
environmental conditions led to the disappearance of coal beds, the
dominance of calcisols in alluvial plains and extensive formation of
red beds sequences towards the Asselian. This climate change was
also associated to the onset of volcanism in the Andean region that
very probably stressed deeply the vegetal communities in the area. In
this sense the recent finding of nurse logs palaeoecological strategy
(Césari et al., 2010) in a Late Carboniferous forest corroborates the
development of adverse environments. This new scenario was firstly
characterized by subtle changes in the vegetation (Interval Biozone
and the palynological Subzone C) and later by the occurrence of the
first glossopterids. The Interval Biozone is characterized by the first
macrofloristic records of conifers and ferns remains. In the same way,
the palynological Subzone C shows a more usual content of taeniate
pollen related to conifers. The increase in bisaccate taeniate pollen is
continuous through the FS and LW Biozones accompanying the
progressive Permian aridization (Fig. 4). Thus, palynological associa-
tions recovered from shallow lacustrine and playa lake deposits of the
La Veteada Formation (Zavattieri et al., 2008) would be representing
the uppermost terms of the LW Biozone.
In short, in our interpretation the Late Paleozoic climates played a
main role in the floristic changes. The two glacial events are likely to
have been responsible of the extinction of the small lycophytes of the
AF flora which was replaced after the cessation of glacial conditions by
arborescent lycophytes, abundant pteridosperms and cordaitales
belonging to the NBG flora. Likewise the apparition of conifers and
glossopterids seems to be related to a progressive climatic warming in
this part of Gondwana.
4. Comparisons with other Gondwanan assemblages
Correlations of palynological biozones of Gondwana have been
hampered by several reasons including plant provinciality, taxonomy,
different standards of documentation of palynological data among
others (see Stephenson, 2008). Additional difficulties are the potential
contamination of subsurface palynological samples (Wood et al.,
1996) and the incompleteness of stratigraphic Late Palaeozoic record
across the Gondwana region supporting the biozones. For example,
palynological knowledge of Australia and Brazil mostly is based on
wells, whereas in central-western Argentina sampling was made on
outcrops of continuous and well referenced stratigraphic sequences.
Also, palynostratigraphy of central-western Argentina shows some
discrepancies with the stratigraphic ranges of some key species used
in other regions. The first appearances of species like Microbaculispora
tentula (=Granulatisporites austroamericanus) and Horriditriletes spp.
are considered Asselian in Oman, Saudi Arabia and Australia
(Stephenson, 2008). However, in the more complete Argentinian
record these species are registered since the Late Serpukhovian–
Bashkirian. In the same way Crucisaccites monoletus appears together
with Ahrensisporites cristatus in Argentina while in Brazil both species
are markers of successive zones. Spelaeotriletes ybertii in Australia is
considered coeval with the first record of monosaccate pollen
although in Argentina the incoming of this species is younger, related
with the first taeniate pollen.
As Stephenson (2008) correctly pointed out, palynostratigraphic
schemes can vary from basin to basin within a region due to
provincialism. To this we add the need for a comprehensive
knowledge of the basin stratigraphy for establishing biozones.
Biozones are intimately related to ecological patterns, and local
environmental and palaeogeographic conditions can conditionate, at
least partially, the stratigraphic ranges of some species. Therefore, we
have compared the Gondwanan fossiliferous assemblages on the basis
of existing radiometric ages as tiepoints across regions. In addition to
the U–Pb calibrated Late Paleozoic palynological record of Argentina,
we incorporated the radiometrically calibrated biostratigraphy of
Brazil (Paraná Basin), Australia and South Africa (Fig. 5).
Different palynostratigraphic subdivisions for the Upper Paleozoic
stratigraphy of Australia were outlined by Kemp et al. (1977), Powis
(1984), Jones and Truswell (1992). In eastern Australia (Fig. 6), the
Early Carboniferous is characterized by the Anapiculatisporites
largus Assemblage (early to late Visean), which is succeeded by the
Grandispora maculosa (Gm) Assemblage. This last biozone is identified
in the Italia Road Formation and at the top of the Mount Johnstone
Formation (Playford and Helby, 1968; Jones and Truswell, 1992) just
below the Paterson Volcanic dated as 329.3±3.8 Ma and 328.9±
4.2 Ma. The PatersonVolcanic also constrained the base of theextended
Australian Nothorhacopteris Flora associated to the Spelaeotriletes
ybertii Assemblage in Western Australia, where the first appearance
of monosaccate pollen is registered. According to Perez Loinaze
(2007) the Gm Assemblage shares some biostratigraphic key spe-
cies (Reticulatisporites magnidictyus,Rugospora australiensis and
Verrucosisporites quasigobbettii) with the Argentinian MQ Biozone.
Iannuzzi and Pfefferkorn (2002) proposed the name Paraca Floral
Realm for relating the Australian palynological biozone and some
macrofloras from South America, India and Africa (Niger) developed
during the late Visean–Early Serpukhovian. Recently, Balseiro et al.
(2009) reported in Argentina the finding of the FNC Biozone, as a
coeval of the Paraca flora (Fig. 3). The Eastern Australian Joe Joe Group
is a key sequence because its lower section was proposed by Jones and
Fielding (2004) as the only unequivocal record of the Namurian–
Westphalian glaciation outside South America. Palynological studies
on this stratigraphic unit allowed to Jones and Truswell (1992) the
characterization of the Spelaeotriletes queenslandensis (=Grandispora
queenslandensis sensu Playford et al., 2001) Superzone (equivalent to
the Spelaeotriletes ybertii Assemblage) subdivided into three Oppel-
zones (A–C). The basal Oppel-zone A was correlated by Fielding et al.
(2008) with the glacial interval C2 and contains monosaccate pollen,
without taeniate specimens, like the Subzone A of the Argentinian DM
Biozone. Levipustula levis fauna characterizes coeval marine deposits
that are dated in approximately 321–323 Ma (Roberts et al., 1995)in
Australia. This age agrees with the inferred age of the Argentinian
glacial event of the Guandacol Formation and coeval glacial-marine
deposits (i.e. Hoyada Verde Formation) bearing rich invertebrate
assemblages belonging to the Levipustula Biozone (Fig. 6).
Perez Loinaze et al. (2010a) also proposed a probable correlation
of the lowermost glacial levels identified in the eastern area of the
Brazilian Itararé Group (Rocha-Campos et al., 2008) with the tillite
Fig. 5. Palaeogeographic reconstruction of Gondwana showing the location of the
stratigraphic sections with radiometric data. 1. Paganzo and Río Blanco Basins, 2. Paraná
Basin, 3. Namibia, 4. western Australian Basins, 5. eastern Australian Basins.
155S.N. Césari et al. / Earth-Science Reviews 106 (2011) 149–160
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of the Guandacol Formation. These glacigenic rocks, included in the
subsurface Lagoa Azul Formation, have yielded palynological assem-
blages with monosaccate pollen referred to the Ahrensisporites
cristatus Zone (Souza, 2006). This AcZ is correlated with the Argen-
tinian Subzone A by sharing many species (Fig. 6). Radiometric ages
reported by Rocha-Campos et al. (2006, 2008) suggest a Serpukhovian
age for the onset of the glaciation in the Paraná Basin while Holz
et al. (see appendix A, 2010) proposed that the glacigenic sedimen-
tation began in the Bashkirian. As in Argentina, Brazilian macrofloras
from this interval are characterized by the species Nothorhacopteris
argentinica and Botrychiopsis weissiana, without glossopterids remains
(Holz et al., 2010).
Australian palynological Oppel-zone B could be also coeval with
Argentinian Subzone A considering the absence of bisaccate taeniate
pollen, whereas the upper part of the Spelaeotriletes Assemblage
distinguished by the Oppel-zone C or Diatomozonotriletes birkheadensis
(sensu Jones and Truswell, 1992) and characterized by the incoming
of Protohaploxypinus, results comparable to the Argentinian Subzone B.
The Australian Potonieisporites Assemblage (=Oppel-zoneD of Jones
and Truswell, 1992) is recognized in the lower and middle sections of
the Seaham Formation which upper section, dated as 312.2± 3.2 and
310.6± 4.0 Ma (Moscovian), is characterized by an increase of taeniate
pollen in palynological assemblages. Retallack (1999) described plant
fossils collected from this interval as a flora lacking diversity and
Fig. 6. Correlation of the main biozones for Argentina, Brazil, South Africa and Australia according to the available radiometric data (♦). Tentative or uncertain boundaries are shown
by dotted lines. References are in text unless otherwise noted. PG: Phyllotheca–Gangamopteris,GB: Glossopteris–Brasilodendron, PG: Polysoneloxylon/Glossopteris LPT: Lycopodiopsis
derbyi–Psaronius–Tietea, Pp: Pseudoreticulatispora pseudoreticulata, Sf: Striatopodocarpites fusus, Mb: Marginirigus barringtonensis. 1. Punta del Agua Formation 335.99 ±0.06 Ma
(Gulbranson et al., 2010); 2. Lower section of the Río del Peñon Formation and Guandacol Formation, 319.57 ±0.09 and 318.79 0.10 Ma (Gulbranson et al., 2010); 3. Tupe Formation
315.46 ±0.07 Ma (Gulbranson et al., 2010); 4. Tupe Formation 312.82 ±0.11 Ma; 5. Patquía Formation, 310.71±0.11, 309.89± 0.08; middle Rio del Peñón Formation 310.63± 0.07
(Gulbranson et al., 2010); 6. Agua del Jagüel Formation 307.2 ±5.2 Ma (Lech, 2002). 7. La Colina Formation, 295 ±6 Ma (Thompson and Mitchell, 1972); 8. La Colina Formation
296.09±0.08 (Gulbranson et al., 2010); 9. Itararé Group 323.5 ±15 Ma (Rocha-Campos et al., 2006, 2008); 10. Middle Rio Bonito Formation, 299 ±2.6 to 296 ±1.4 Ma (Guerra-
Sommer et al., 2005); 298.5 ±2.6 Ma (Rocha-Campos et al., 2006); 11. Irati Formation 279.9 ±4.8 Ma and 278 ±2.2 Ma (Rocha-Campos et al., 2006; Santos et al., 2006); 12.
Ganigobis Shale Member, 302 ±3 Ma and 299.2 ±3.2 Ma (Bangert et al., 1999); 13. Top of DS III, 297 ±1.8 Ma (Bangert et al., 1999); 14. Paterson Volcanics 329.3 ± 3.8 Ma, 328.9 ±
4.2 Ma (Roberts et al., 1995); 15. 321–323 Ma (Roberts et al., 1995); 16. Upper section Seaham Formation 312.2±3.2 and 310.6 ±4.0 Ma (Roberts et al., 1995).
156 S.N. Césari et al. / Earth-Science Reviews 106 (2011) 149–160
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characterized by Botrychiopsis plantiana (Carruthers) Archangelsky and
Gamerro and Dichophyllites peruvianus (Gothan) Morris. Coeval
palynofloras in Argentina are represented by the Subzone C and the
Interval macrofloristic biozone. Palynological composition of the
Brazilian Crucisaccites monoletus Zone (CmZ) suggests a correlation
with these assemblages from Australia and Argentina (Fig. 6). Césari
(2007) also proposed a correlation among the palynological Subzone B,
or Quadrisporites Assemblage Zone of Falcon (1975), and their
equivalents: Biozone 1 of Anderson (1977) and Biozone A of MacRae
(1988) in South Africa with the Argentinian Subzone C.
According to Césari (2007) palynological assemblages from the
Carboniferous–Permian boundary of western southern Africa, Brazil
and Argentina show close similarities. This boundary is approximately
represented in southern Africa by the Ganigobis Shale Member,
dated to 302± 3 Ma and 299.2 ± 3.2 Ma (Bangert et al., 1999) and
the Brazilian coal seams of the Rio Bonito Formation (Fig. 6) dated to
299± 2.6 to 296 ±1.4 Ma (Guerra-Sommer et al., 2005).
The Ganigobis Shale Member is 130 m thick at the Ganigobis
locality (Namibia) and it is characterized by continental subglacial
tillite succeeded by through cross-bedded fluvioglacial outwash
conglomerates and glacimarine deposits (Stollhofen et al., 2000).
Glacimarine conditions were established in the upper part repre-
sented by laminated dropstone-free silty mud rocks, which contains
marine fauna. Fossils woods were identified in two stratigraphic levels
in the lower and upper part (Bangert and Bamford, 2001). Recently,
Stephenson (2009) reported from the Ganigobis Shale Member of
Namibia, dated to 302.0 Ma (Gzhelian), a diverse assemblage analo-
gous to the Australian Converrucosisporites confluens Oppel Zone (Cc).
Foster (in Foster and Waterhouse, 1988) interpreted this later biozone
immediately overlying the upper part of the Spelaeotriletes Biozone.
Although Cc Oppel Zone has been considered Asselian to Tastubian in
age (Archbold, 2001), Stephenson (2009) proposed that the lower
limit could extend earlier and be as low as the latest Carboniferous.
The Brazilian Early Permian Vittattina costabilis (VcZ) Interval Zone
(Souza and Marques-Toigo (2003) is characterized by the abundance
and diversity of taeniate pollen grains and is registered in the upper
Itararé Subgroup and Rio Bonito Formation (Marques-Toigo, 1991).
The VcZ was subdivided by Souza and Marques-Toigo (2003) into two
subzones, Protohaploxypinus goraiensis and Hamiapollenites karroensis.
The subzone P. goraiensis is recognized from the uppermost section of
the Itararé Subgroup to the middle section of the Rio Bonito Formation
and contain specimens of Converrucosisporites confluens (Souza and
Callegari, 2004). Guerra-Sommer et al. (2005) obtained a time of
deposition based on ID-TIMS U–Pb zircon ages ranging from 299 ±2.6
to 296± 1.4 Ma for tonsteins located in the middle section of the Rio
Bonito Formation. A similar age (298.5 ±2.6 Ma) was obtained by
Rocha-Campos et al. (2006, 2008) for the same stratigraphic section.
Therefore, the first records of C. confluens in South America seem
approximate in age to the reported in South Africa.
It is noteworthy the coincidence (within analytical uncertainty of
the data reported from Guerra-Sommer et al., 2005) with the
206
Pb/
238
U age of 296.09 ±0.08 Ma obtained from a tuff within eolian
stratigraphy overlying the La Torre basalts in Argentina (Gulbranson
et al., 2010) with those from the Rio Bonito Formation. This reinforces
a close correlation proposed by Césari (2007) between palynological
assemblages from the Rio Bonito Formation (Vittatina costabilis
Biozone) and the Argentinian FS Biozone.
The associated macrofloral remains of this stratigraphic interval
in southern Africa, Brazil and Argentina are characterized by
glossopterids leaves. Archangelsky and Cúneo (1984) defined the
Gangamopteris Biozone to include the floristic assemblages recovered
from this interval in Argentina. Gangamopteris leaves also characterize
the deglaciation facies in the Karoo successions (Catuneanu et al.,
2005) and the Rio Bonito coals from Brazil (Iannuzzi and Souza, 2005).
The top of the depositional sequence III in southern Africa (Visser,
1997), closely related to the Gondwana-wide Eurydesma transgres-
sion, gave a SHRIMP
206
Pb/
238
U age of 297 ± 1.8 Ma (Bangert et al.,
1999). In Argentina, Eurydesma-bearing shales are recognized in the
Bonete Formation (Buenos Aires Province) containing Glossopteris
remains also, but unfortunately nothing is known about their paly-
nological content (Harrington, 1955; Rocha-Campos and de Carvalho,
1975). Some characteristic marine invertebrate species related to
this Australian Fauna (=Lyonia lyoni fauna), were recovered from the
upper part of the Itararé Group (Rocha-Campos and Rosler, 1978),
where the V. costabilis Zone has been identified.
Souza and Marques-Toigo (2003) distinguished the Brazilian
Lueckisporites virkkiae Zone by the dominance of taeniate pollen
and the abundance of Lueckisporites virkkiae Potonié and Klaus,
Marsupipollenites triradiatus Balme and Hennelly, Protohaploxypinus
perfectus (Naumova) Samoilovich, Lunatisporites variesectus among
other species. This biozone is represented in the Palermo, Irati, Serra
Alta and Teresina Formations and referred to the late Cisularian to early
Guadalupian by Souza and Marques-Toigo (2005). A SHRIMP age of
278.4± 2.2 Ma was reported for the biozone in the Iratí Formation
(Santos et al., 2006). An U–Pb zircon dating of 270 ± 1 Ma seems to
constrain the age of the underlying Whitehill Formation that bears the
amphibious reptile Mesosaurus just like the Irati Formation in Brazil. The
Argentinian Lueckisporites–Weylandites Biozone related to radiometric
ages of 282± 13 Ma and 266 ±5 Ma is considered coeval of the Brazilian
assemblages (Césari, 2007). Similar ages have been calculated for the
Zone 3 of Anderson (1977) in South Africa. Beri et al. (2004) proposed
the Striatoabieites anaverrucosus/Staurosaccites cordubensis(AC) Biozone
in the Early Permian of Uruguay (Mangrullo Formation), characterized
by the presence of Lueckisporites and dated radimetrically as no older
than Artinskian (Rocha-Campos et al., 2006).
5. Conclusions
Although some of the Argentinian biozones may seem broad, they
are mostly based on sampling of many well known continuous
stratigraphic sections. Moreover, it should be noted that only floras
with regional distribution were used for defining them. This
biozonation is constrained by recent radiometric data in the following
biostratigraphic intervals:
Late Tournaisian–Early Visean: represented by Archaeosigillaria–
Frenguellia,Cordylosporites/Verrucosisporites and Protocanites
scalabrini–Azurduya chavelensis Biozones. This biota was charac-
terized by essentially shrubby plants (mainly lycophytes and
pteridosperms) together with brachiopods and ammonites in
marine environments.
Late Visean: distinguished by the palynological Reticulatisporites
magnidictyus–Verrucosisporites quasigobbetti and invertebrate
Rugosochonetes–Bulahdelia Biozones closely related to a glacial
event.
Early Serpukhovian: characterized by the Frenguellia eximia–
Nothorhacopteris kellaybelenensis–Cordaicarpus cesarii Biozone, a
typical interval zone, with plant species in common with the under
and overlying floras.
Late Serpukhovian–Early Bashkirian: distinguished by plant and
palynological associations included in the Nothorhacopteris–
Botrychiopsis–Ginkgophyllum Biozone and Subzone A of the
Raistrickia densa–Convolutispora muriornata (DM) Biozone to-
gether with marine invertebrates of the Levipustula Biozone. This
biota is closely related with the most important glacial event in
western Argentina.
Late Bashkirian: characterized by palynological Subzone B of
the DM Biozone and plants referred to the Nothorhacopteris–
Botrychiopsis–Ginkgophyllum Biozone.
Moscovian: distinguished by transgressive marine deposits contain-
ing the Tivertonia jachalensis–Streptorhynchus inaequiornatus fauna
and the vegetation represented by the Interval Biozone and Subzone C.
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Ghzelian?–Asselian: the palynological Pakhapites fusus–Vittatina
subsaccata Biozone characterizes the latest Carboniferous–Early
Permian accompanying the incomingof the first Glossopteris remains.
Sakmarian?–Artinskian: the palynological Lueckisporites–Weylandites
Biozone is distinguished by the dominance of taeniate pollen grains
in stratigraphic sections also characterized by the Costatumulus amosii
fauna.
The integration of the biostratigraphy of the Upper Paleozoic in
southern Gondwana with radiometric ages supports some previous
inferences and highlights several important common bioevents. Visean
fossiliferous assemblages are better known in western Argentina and
Australia where plant remains are typically represented by lycophytes
and primitive pteridosperms. Palynological assemblages from both
areas are characterized by the absence of pollen and share some
spore species. During the Late Visean a glacial event is identified in
western Argentina. The Serpukhovian beds of the same region contain
plants included in the FNC flora, coeval in part with the Australian
Grandispora maculosa Biozone. This FNC flora precedes the Argenti-
nian late Serpukhovian–Early Bashkirian glacial episode which
coincides with the C2 glacial event documented for eastern Australia
(Fielding et al., 2008) and also recognized in Brazil. These glacial and
postglacial deposits are characterized by plants assigned to the NBG
Biozone in Argentina and equivalent floristic assemblages containing
Nothorhacopteris argentinica in Brazil and Australia. Associated paly-
nological records are distinguished in the three areas by the presence of
abundant monosaccate pollen. During Late Bashkirian–Moscovian an
increase of taeniate pollen is recognized in palynological assemblages
from Argentina, Brazil and Australia. First records of Protohaploxypinus
pollen are registered in the Argentinian Subzone B, in the upper part of
the Brazilian Ahrensisporites cristatus Biozone and the uppermost part
of the Spelaeotriletes Assemblage in Australia. The early Moscovian in
western Argentina coincides with a marine transgression characterized
by the TS Biozone and palynological Subzone C. Coeval fossiliferous
associations in Australia are represented by the Potonieisporites
Assemblage and the Crucisaccites monoletus Biozone in Brazil.
Latest Carboniferous interval is represented in South Africa by
glacial and postglacial deposits of the Dwyka Group. Palynological
assemblages dated in approximately 302 Ma are characterized by the
presence of Converrucosisporites confluens,Protohaploxypinus limpidus,
Alisporites indarraensis,Vittatina sp. (Stephenson, 2009), among
other species also identified in Argentina, Brazil and Australia too
(FC, Vc and Cc Biozones). Floristic remains in all areas belong to the
Gangamopteris flora. These biostratigraphic units are well represented
in Early Permian strata. A noticeable change in the palynological
assemblages occurs with the introduction of Lueckisporites and 50%
or more of taeniate pollen in the LW and Lueckisporites virkkiae
Biozones in South America and the Stage 3 in South Africa (Césari,
2007).
Acknowledgments
Dr. Maurice Streel, an anonymous reviewer and Editor Paul
Wignall are thanked for their constructive reviews of the submitted
manuscript. Financial support for this study was provided by ANPCyT
(Agencia Nacional de Promoción Científica y Tecnológica) PICT 20752
and 1499 grants.
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