Journal of South American Earth Sciences 63 (2015) 417e435

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Journal of South American Earth Sciences

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First record of Ginkgo-related fertile organs (Hamshawvia,

Stachyopitys) and leaves (Baiera, Sphenobaiera) in the Triassic of Brazil,
Santa Maria formation
^nia Lindner Dutra*
Ronaldo Barboni, Ta
~o em Geologia/LaViGæa, MHGEO, Universidade do Vale do Rio dos Sinos, UNISINOS Av. Unisinos 950, 93022-000 Sa
s-Graduaca
Programa de Po ~o Leopoldo,
RS, Brazil

a r t i c l e i n f o a b s t r a c t

Article history: This paper describes new ginkgophytes recently discovered in a new exposure of the original type
Received 24 November 2014  Basin succession. The
section of the Passo das Tropas Member, Santa Maria Formation, linked to Parana
Received in revised form well-preserved fossils come from a very fine red laminated mudstone representing a lacustrine interval,
28 July 2015
within a broader low-sinuosity fluvial succession. The material occurs in a Dicroidium-dominated
Accepted 4 August 2015
Available online 10 August 2015
assemblage and is represented by the leaves Baiera and Sphenobaiera, accompanied by the fertile
structures Hamshawvia and Stachyopytis. Cuticle fragments preserved on leaf impressions of Spheno-
baiera schenkii show xeromorphic features. The geology, paleogeography and paleoclimate of the Passo
Keywords:
Baiera
das Tropas Member (PTM), Santa Maria Formation, is discussed. By comparing these ginkgophytes with
Sphenobaiera those from other Gondwanan assemblages, the Brazilian deposits earlier assigned to AnisianeLadinian
Hamshawvia age can now be extended to the Carnian age.
Stachyopitys © 2015 Elsevier Ltd. All rights reserved.
Taphonomy
Paleogeography
Paleoclimate
South Brazil
Santa Maria formation
MiddleeLate Triassic

1. Introduction the Pennsylvanian to recent times. Bauer et al. (2013a) provided a

Since the end of the Paleozoic and through the Mesozoic, the
Ginkgoales have been an important component in the floras
reaching a great diversity in the end of Triassic and onwards
(Anderson and Anderson, 1989; Zhou and Wu, 2006; Bauer et al.,
2013a). Some taxonomic uncertainties remain due to morpholog-
ical characters shared with the Peltaspermales, Dicranophyllales
and Coniferales (Stewart and Rothwell, 1993; De Franceschi and
Vozenin-Serra, 2000; Taylor et al., 2009). This is the case with the
Permian genera Trichopitys, Polyspermophyllum and Dicranophyllum
(Meyen, 1987; Archangelsky and Cúneo, 1990; DiMichele et al.,
2013). The fossil record of ginkgophytes extends over Laurasia
(Seward, 1919; Meyen, 1987; Kerp, 1996; Zhou, 2009) and Gond-
wana (Archangelsky, 1996; Silva and Iannuzzi, 2000), dating from
* Corresponding author.
E-mail addresses: ronaldobarboni@gmail.com (R. Barboni), tdutra@unisinos.br,
dutratl@gmail.com (T.L. Dutra).
thorough synthesis of the history of ginkgophytes.
The establishment of relations between Ginkgoales in the fossil
record and their younger representatives are complex. More
recently, reproductive structures and leaf anatomy have shown
some features that allow researchers to correlate the Paleozoic
ginkgophytes and those from the Mesozoic, and with some aber-
rant leaves produced by modern Ginkgo (Stewart and Rothwell,
1993; Fischer et al., 2010). Anomalous inflorescences, with multi-
ple ovules, were detected in Trichopitys Saporta from the French
Permian, and in Karkenia Archangelsky from the Lower Cretaceous
of Santa Cruz in Argentina (Archangelsky, 1970). A revaluation of
the ginkgophytes from the Kupferschiefer in Germany revealed that
truly petiolate leaves were present by the end of Permian, sug-
gesting the establishment and diversification of the ginkgophytes
by that time (Bajpai, 1991; Bauer et al., 2013a).
A major innovation is record in the Late Triassic with the first
appearance of dwarf shoots, the differentiation of leaves into
lamina and petiole, and a reduction in the number and increase in

http://dx.doi.org/10.1016/j.jsames.2015.08.001
0895-9811/© 2015 Elsevier Ltd. All rights reserved.
418 R. Barboni, T.L. Dutra / Journal of South American Earth Sciences 63 (2015) 417e435
size of the ovules (Zhou and Wu, 2006). Between the end of the
Triassic and Early Cretaceous the group reached its acme in abun-
dance and a global distribution, showing distinct leaf morphol-
ogies, most of which broadly resemble modern Ginkgo and are
assigned to the fossil taxon Ginkgoites (Zhou and Wu, 2006;
Anderson et al., 2007). Others, with more dissected leaves, were
included in Baiera and Sphenobaiera genera (Florin, 1936; Zhou,
1997; Watson et al., 1999). Fossil seedlings and twigs found in the
Middle Triassic of France show some analogy with those of modern
Ginkgo biloba and attest to the development and evolution of the
group in the early Mesozoic (Bauer et al., 2013b).
Ginkgoales were an important group (second only to cor-
ystosperms) in the Middle Late Triassic Dicroidium floras of Gond-
wana (Du Toit, 1927; Artabe, 1985; Anderson and Anderson, 1989;
Gnaedinger and Herbst, 1999; Troncoso and Herbst, 1999; Artabe
et al., 2007; Moisan et al., 2010; Lutz et al., 2011; and
Supplementary online data). In the Nymboida Flora from Australia,
leaves with intergradational morphologies found in unique as-
semblages, were included in morphospecies complexes. This pre-
vented taxonomic multiplication of names (species) which would
probably reflect only subtle morphological variations, due to ad-
aptations in the local environment (Holmes and Anderson, 2007).
Microsporangiate and ovuliferous structures, although rarer,
was globally found after the end of the Permian (Shirley, 1898;
Walkon, 1917; Frenguelli, 1942; Jain and Delevoryas, 1967;
Krassilov, 1982; Zhou et al., 2002; Holmes and Anderson, 2007;
Naugolnykh, 2007; Yang et al., 2008; Zhou et al., 2012).
In the Upper Triassic Molteno Formation, in South Africa, gink-
goalean foliages were included in Sphenobaiera, Ginkgoites and
Paraginkgo. The ovuliferous structures were assigned to the new
genus Hamshawvia, and to a new order, the Hamshawviales and an
immature form of Hamshawvia, attached to leaves of Sphenobaiera
“schenckii”, was described (Anderson and Anderson, 2003).
Anderson and Anderson (2003) transferred to this new genus,
Umkomasia cacheutensis originally described from Argentina by
Frenguelli (1942). In the Nymboida Flora, Australia, Hamshawvia,
Stachyopitys and Sphenobaiera schenkii was founded associate,
albeit without organic connections (Holmes and Anderson, 2007,
2013).
In Brazil, the previous fossil record of Ginkgoales was scarce and
restrict to the southern Parana  Basin (Fig. 1A). The oldest examples,
from the Lower Permian Itarare  Group and Rio Bonito Formation,
include incomplete leaves of Ginkgoites eximia Seward, Ginkgo-
phytopsis Høeg, Dicranophyllum and Cheirophyllum speculare Ce sari
and Cúneo, the latter exhibiting dubious affinities to the Ginkgoales
(Silva and Iannuzzi, 2000). They were associated with assemblages
composed mostly by pteridosperms (Glossopteris and Gang-
amopteris), accompanied by sphenopsids and primitive conifers
(Guerra-Sommer et al., 1999; Guerra-Sommer and Klepzig, 2000).
In the Triassic, the ginkgophytes were record in levels correlated
to those herein studied (Santa Maria Formation, Passo das Tropas
Member, PTM), represented by ex situ fossil woods (Bardola et al.,
2009), sparse seeds, reproductive structures (Stenorrachis) and
incomplete leaves of Sphenobaiera and Ginkgoites antarctica (Pinto,
1956; Bortoluzzi et al., 1983).
Besides the dominant corystosperms component (70% of esti-
mated relative frequency), the Ginkgoales were the second most
abundant group in the PTM, which includes Neocalamites, conifers,
Taeniopteris and rare Cladophlebis (Gordon and Brown, 1952; Pinto,
1956; Bortoluzzi et al., 1983; Guerra-Sommer and Klepzig, 2000).
The poorly preserved Sphenobaiera leaves was compare with
S. ugotheriensis and S. argentinae, respectively from Australia and
Argentina (Bortoluzzi et al., 1983). In the AnisianeLadinian age
proposed for the Passo das Tropas Member (PTM) was taking into
account the range of Dicroidium species (Mastroberti, 1995; Guerra-
Sommer et al., 1999).
In this study new ginkgoalean fossils are described from an
outcrop near Santa Maria city, central Rio Grande do Sul (Fig. 1B),
not far from the original type-section of the PTM (Bortoluzzi, 1974).
Along with the other Triassic deposits in Brazil, PTM beds are
exclusive to the southern areas of the Paran a Basin.
1.1. Geological background
According to Zerfass et al. (2004), the facies and geometry of the
Brazilian Triassic deposits are related to an extensional phase of an
intraplate rifting associate with the Gondwanides orogeny, that
also originate the back-arc rift basins from Argentina and South
Africa (Central West Gondwana basin complex from Linol et al.,
2014). The Early Triassic initial compressional phase was follow
by successive extensive events spanning the Middle to Late Triassic
transition and which anticipated the initial stages of Gondwanan
break-up. The Sierra de La Ventana and Cape Fold Belt orogenic
belts (Uliana et al., 1989; Kokogian et al., 2001; Zerfass et al., 2004;
Faccini, 2007; Barredo et al., 2011) also controlled those efforts.
In this scenario the Karoo Basin and probably the Santa Cruz
basin in Southern Argentina, were located at 30
S of paleolatitude,
influenced by extratropical warm and wet temperate climates. The
Ischigualasto, Paganzo and the southern Parana  basins, located in
between 30
e15
S, varied from a transitional condition or the in-
fluence of drier subtropics (Spalletti et al., 2003; Holz, 2015). Only
the La Ternera Basin in Chile may have been located at <10
S
(Zerfass et al., 2004).
According to Milani et al. (1998), six stratigraphic mega-
sequences bounded by interregional unconformities characterize
the Parana  Basin. The three upper (MiddleeLate Triassic to Late
Cretaceous) attest a gradual continentalization, expressed by
fluvial, lacustrine and wind generated red bed deposits, covered by
the Jurassic e Cretaceous volcanic rocks (Bortoluzzi, 1974; Andreis
et al., 1980; Milani et al., 1998). In the State of Rio Grande do Sul
these sedimentary sequences are exposed along an eastewest
central belt (Fig. 1BeC).
Faccini and Paim (2001) and Zerfass et al. (2003), applying the
Sequence Stratigraphy principles to the Triassic deposits from
southern Brazil consider that the basal Sanga do Cabral (Early
Triassic) and upper Santa Maria (MiddleeLate Triassic) super-
sequences represent 2 s-order cycles. To the Santa Maria super-
sequence they proposed three sequences (Santa Maria 1, 2 and 3),
with a southward located source area and depositional architec-
tures, comparable to those from other Western Gondwana rift
basins.
In this approach, the ginkgophyte and the Dicroidium flora from
the Passo das Tropas Member (PTM) were assign to the basal part of
Santa Maria 2 Sequence (SM2), were the tetrapod fauna indicates a
Carnian age (Schultz et al., 2000; Langer et al., 2007). In the
sequential framework proposed by Zerfass et al. (2004), the mud
facies of the PTM deposits containing the fossil flora represent
phases of base level changes that led to the massive lacustrine beds
containing the tetrapod fossils. The Carnian age proposed to the
PTM deposits is in agreement with the associated coleopteran
faunas (Pinto and Ornellas, 1974), comparable to that identified in
Los Rastros, Cacheuta, Ischichuca and Santa Joana formations, in
Argentina (Mancuso et al., 2007; Martins-Neto et al., 2011).
The lenticular geometry of the plane-bedded mudstones from
the PTM (Fig. 2A) suggest flood plains/lacustrine deposits formed in
abandoned low areas (or channels) protected from erosion, of a
broad and dominant fluvial system of low sinuosity rivers (Zerfass
et al., 2003; Faccini, 2007).
In the outcrop herein studied, located at geographical co-
ordinates 29
440 37, 8500 S and 53º470 31, 1200 W (Fig. 1C, Fig. 2AeD),
 R. Barboni, T.L. Dutra / Journal of South American Earth Sciences 63 (2015) 417e435 419

Fig. 1. A, Map of Brazil and main sedimentary basins and terrains; B, State of Rio Grande do Sul with the distribution of the PermianeTriassic sedimentary sequences, outcropping
along East-Western belt; C, geological map of the central Rio Grande do Sul and the areas exposing the deposits of Santa Maria Formation. The star indicates the location of the
studied outcrop at Passo das Tropas Stream, near Santa Maria city.

the plant remains occur in rhythmites (siltstones-mudstones)
overlying by intraformational conglomerates, and covered by cross-
bedded sandstones deposits (Fig. 2AeB). Common facies and
paleontological content point to a correlation of this outcrop with
the type section of PTM, as determined by Bortoluzzi (1974). The
plant remains found by Bortoluzzi (1974) in the mud clasts of the
basal conglomerate enables us to infer that older and similar
lacustrine horizons with plant remains may have existed.
2. Materials and methods
A high-resolution analysis was conduct along the rhythmites in
order to observe the taphonomic constraints of the floral assem-
blage. Quadrats of 20 cm2 were examined along the 2 m high
section, 10 cm apart from each other (an adapted methodology to
that proposed by Pfefferkorn et al., 1975). For each level, all well-
preserved macrofossils were registered and collected.
A total of 638 samples containing plant fossil impressions were
recovered from the 20 levels raised, including complete leaves,
partial fragments, insect wings, carapaces of Spinocaudata, fish-
scales and evidences of planteinsect interaction (Corre ^a, 2014;
Dutra and Barboni, 2014). The first three basal levels are charac-
terized by the exclusive presence of isolate Heidiphyllum leaves,
whereas the two upper ones were sterile. Four intermediary levels
showed plant remains covered by iron oxy-hydroxide coats
(Fig. 2C).
In order to prevent the physical (contraction and expansion) and
chemical disintegration, common in mudstones, the samples were
wrapped in a PVC plastic film during the fieldworks and there
maintained until their complete dehydration (Dutra, 2002; Barboni
et al., 2008). In order to make handling easier, delicate samples as
well as the large ones had their bases inserted into asphalt blankets
(3 mm thick) and paraffin wax (Barboni et al., 2008).
In the evaluation of ginkgophytes representativeness along the
PTM levels, a matrix of presence/absence was produced, using the
PAST software (Hammer et al., 2001). Within the fossil material 75
samples exhibited Ginkgo-related remains and 51 displayed well-
preserved leaf impressions and female/male strobili. The samples
420 R. Barboni, T.L. Dutra / Journal of South American Earth Sciences 63 (2015) 417e435

Fig. 2. A. Geological section from the Passo das Tropas outcrop and its facies, highlighting the lacustrine shales with plant fossils; B, intraformational conglomerates containing mud
clasts; C, detail of the mudstone interval, with dispersed lenses of iron oxy-hydroxide deposits (black lenses); D, trough cross-bedding sandstones from the sand bars; E, detail of a
conchostracan valve among the Sphenobaiera schenkii leaf segments.

were house under the label ULVG at the Paleontological Labora-
tory (LaViGæa) of the Post-graduate Program in Geology e UNI-
SINOS. The leaf material was photographed with a Fuji Finepix
S3300 digital camera under inclined daylight, with polarized light
filters in order to improve the contrast and definition of details.
Poorly preserved specimens or those with fine venations were
draw in a camera lucida, through an Olympus SZH
stereomicroscopic.
Fossil impressions covered by iron oxy-hydroxide films or crusts
allowed the analysis of epidermal anatomical features which were
photographed and analyzed in natura, under a ZEISS EVOMA15
Scanning Electron Microscope. To the establishment of taxonomic
relations all of the fossils (in situ and ex situ samples) were
analyzed.
2.1. Taxonomic aspects of Triassic ginkgophytes
Distinctions among isolated ginkgophyte leaves in the Triassic
record (e.g. Baiera, Sphenobaiera and Ginkgoites) are difficult to
establish (Archangelsky, 1970; Retallack, 1980; Meyen, 1987;
Anderson and Anderson, 1989; Stewart and Rothwell, 1993;
Zhou, 1997; Gnaedinger and Herbst, 1999; Watson et al., 1999;
Bauer et al., 2013a). The major problem is related to the rather
wide spectrum of transitional morphologies or foliar poly-
morphism (Zhou, 1997), and the detection of what is identified as
a “distinct petiole” of Baiera and Ginkgoites forms (Bauer et al.,
2013a). Moreover, in the study of the abundant fossil record of
ginkgophytes from South America, most of the well-described
diagnostic features were published in Spanish (Frenguelli, 1946;
Mene ndez, 1951), making it difficult for researchers to gain a
thorough knowledge of the ginkgophytes diversity in the area. It
was only after 1980 that detailed analysis and illustrations were
produced in Argentina (Baldoni, 1980), in South Africa (Anderson
and Anderson, 1989, 2003), in Australia (Retallack, 1980, 1987;
Holmes and Anderson, 2007). More recently, detailed studies
on the Permian floras of the Northern Hemisphere were made
(Zhou and Wu, 2006; Bauer et al., 2013a). However, distinct
conceptions remain about what is the better way to identify the
genera and species of the group (Zhou, 1997, 2009; Bauer et al.,
2013a).
In order to simplify this approach while also allowing the
analysis of partially preserved materials, a resume of the diagnostic
anatomical and morphological features that distinguish Baiera from
Sphenobaiera is supply in Table 1. This represents an intention of
summarizing many of the aspects proposed by different re-
searchers, with the emended diagnosis proposed by Florin (1936)
to Baiera and can be useful to determine the main morphological
and anatomical features that distinguish Baiera and Sphenobaiera.
The description of reproductive structures is based on the
nomenclature proposed by Anderson and Anderson (2003).
 R. Barboni, T.L. Dutra / Journal of South American Earth Sciences 63 (2015) 417e435 421

Table 1
Morphological and anatomical leaf characters in Baiera and Sphenobaiera highlighting in bold those that distinguishes the two genus. Observations: (þ) similar to Ginkgoites in
morphology and anatomical features; (*) similar to Czekanowskia, but multiveined (Stewart and Rothwell, 1993); (#) To Zhou (1997), however rarer, deeply dissected lamina
can also occur in Sphenobaiera. 1. Seward (1919), Meyen (1987), Gnaedinger and Herbst (1999); Moisan et al. (2010); 2. Braun (1843), Florin (1936), Harris and Millington (1974);
3. To Meyen (1987), Ginkgo and Ginkgoites have disorderly and scattered stomata; 4. Anderson and Anderson (1989); 5. Bauer et al. (2013a).

Baiera (Braun) Florin 1936 (þ) Sphenobaiera (Florin) Harris and Millington 1974 (*)

Lamina Broadly triangular, deeply dissected (#), wedge-shaped down to the base (2), highly dissected, 1e2 bifurcated (1)
many times bifurcate (1,2,5).
Segments Numerous, linear in form (2), narrow Few (2e5) broad segments (arranged into two groups), commonly ovalate in form
(symmetrical arranged), deeply bifurcate
Veins 2e4 parallel veins (1,2), forking many times (5). Variable in number (few to many), in general no forking, with parallel course (1e2)
Apices round to truncate round, obtuse or emarginated
Petiole Distinct, with consistent width (1,2,5) Dilute or inexistent (1), gradatively originating the segments (2, 5), anatomically ~ to the leaf (4)
Epidermis Longitudinally orientated stomata in distinct rows. Longitudinally orientated stomata in indistinct rows, amphistomatic leaf, guard cells containing papilla
(3) Elongate resin bodies and thickness (“Florin rings”). Round or oval resin bodies

2.2. Systematic paleontology
Division: Ginkgophyta Bessey, 1907.
Class Ginkgoopsida Meyen, 1984.
Order Ginkgoales Goro zankin 1904.
Genus Baiera Braun, 1843.
Type species: Baiera muensteriana (Presl in Sternberg, 1833)
Heer.
Baiera africana Baldoni (1980)
(Fig. 3AeE)
Material. ULVG 9770aA/9770bA; 9758 (more complete leaves);
ULVG 8899; 8914; 9260; 9264; 9478; 9490; 9504; 9517; 9518;
9520; 9529; 9545a/9545b; 9548a/9548bA; 9662; 9762; 9769a/
9769b; 11081a/11081b; 11103a/11103b; 11095B; 11112; 11199
(partially preserved materials).
Temporal distribution. Rhaetian, in Gondwana (Table 2).
Occurrence. Middle to upper part of the profile (PTM, Santa
Maria Formation).
Description. Fragmentary impressions of petiolate (petiole
25 mm long and 2 mm wide), symmetrical and triangular leaves,
the most complete one with a maximum of 80 mm long and 60 mm
wide and a sharp acute basal angle (25
). The deeply dissected
lamina bifurcates four times and gives place to many narrow and
elongated segments (more than 16 are preserved) with parallel
margins (2 mm wide) and round apices. Four weak parallel veins
cover each segment.
Remarks. This is the most abundant morphotype among the
Ginkgoales in the PTM profile. The proposed assignment to Baiera
are based on the original diagnosis made by Baldoni (1980), which
indicated the presence of a distinct petiole, numerous linear seg-
ments, and a small acute basal angle for B. africana. It also takes into
account the criteria shown in Table 1 and mainly by the consistent
width of the petiole and number of veins by segment. It is impor-
tant to note, however, that Anderson and Anderson (1989, 2003), in
their description of the Carnian Molteno assemblages, associate
this morphotype to Sphenobaiera and do not use the genus Baiera.
In Argentina, related forms was recorded in Can ~ adon Largo, Los
Rastros and Ischigualasto formations (Middleeearly Late Triassic)
and extends through to the Rhaetian Llantenes and Paso Flores
formations (Spalletti et al., 1999; Gnaedinger and Herbst, 1999;
Stipanicic and Marsicano, 2002; Lutz et al., 2011). B. africana also
occurs in the Upper Triassic of Chile, associated with abundant
Dicroidium, Heidiphyllum and Taeniopteris. The only distinct aspect
in relation to the assemblage discussed herein is the occurrence in
Chile deposits of the more advanced Baiera furcata species (Moisan
et al., 2010), also found in the Jurassic of China (Wu et al., 2006).
The proportional abundance of B. africana and its homogeneous
presence along the profile, considering the restricted wise area
exposure of PTM levels in Brazil, resemble that from the Molteno
Formation and is too distinct from its scarce occurrence in
Argentina. The specimens assigned to Sphenobaiera sp. by Guerra-
Sommer and Klepzig (2000, Fig. 2d) to PTM levels from Brazil
seems to represent the upper segments of the same leaves herein
described.
Baiera sp. 1.
(Fig. 4AeB)
Material. ULVG 9761a/9761b.
Occurrence. Middle part of the profile (PTM, Santa Maria
Formation).
Description. Part and counterpart of a partially preserved fan-
shape lamina with delicate texture (84 mm long). The woody,
long and straight petiole (33 mm long) maintains its consistent
width (2 mm), only expanding apically (3 mm wide). Basal angle of
bifurcation sharply acute (15
). At least nine single veined, slight
and curved terminal segments (1.5 mm wide). Apices not
preserved.
Remarks. The unique leaf preserved shows exclusive characters.
The well-developed petiole, the base expanding into a leaf sheath,
and the great number of slender segments, attest to a relation with
Baiera (Table 1).
Judging by the curved disposition of the lamina, Baiera sp. 1
resembles both the Permian form Tricopithys Krausel
(T. heteromorpha), and Sphenobaiera pontifolia from the Molteno
Floras (Anderson and Anderson, 1989), later included in Baiera
(Lutz et al., 2011). However, both have a lower number of segments
(4e6) due to the restricted number of bifurcations (2e3), in relation
to the herein described leaves. The size and curved segments of the
PTM material is also comparable to S. pontifolia from the Upper
Triassic assemblage of Northern Chile, where the curvature of the
segments were consider a result of the taphonomic processes and/
or post-mortem inclination of the lamina (Troncoso and Herbst,
1999). The incomplete preservation and scarce material herein
detected prevents better comparison.
Baiera sp.1 is in association with Equisetales (Neocalamites), in-
sect wings, seeds, and pinnules of Zuberia feistmantelli Frenguelli.
Genus Sphenobaiera (Florin) Harris and Millington, 1974.
Type species. Sphenobaiera spectabilis (Nathorst) Florin 1936.
Sphenobaiera tenuinervis sp. nov.
(Fig. 4C)
1999. Sphenobaiera sp. C (in Gnaedinger and Herbst, pg. 295,
Figs. 12DeG and 13 HeI).
Diagnosis. Fan to wedge-shaped leaf impression, coriaceous,
broad basal angle and 2e3 times symmetrically dissected. In
distinct petiole gradually enlarging to originate cuneate segments,
slight embossed and coriaceous. Segments with depressed broad
margins and covered by very fine longitudinal veins.
Derivation of name. Refers to the delicate, weak venation of the
leathery segments.
422 R. Barboni, T.L. Dutra / Journal of South American Earth Sciences 63 (2015) 417e435

Fig. 3. AeD, Baiera africana Baldoni (ULVG 9770 aA-bA). A, leaf impression with long and narrow segments. A seed (in lower left) and two recurve cupules (middle right) of
Umkomasia sp. are also preserved; B, counterpart of A; C, detail of the apical portions of the segments; D, detail of the petiole. The arrow indicates probable insect galls; E, specimen
ULVG 9758 showing two superimposed laminae and an indeterminate seed. Scale bars: Figs. AeD ¼ 10 mm; E ¼ 5 mm.

Holotype. ULVG 10880, LaViGæa Collection, UNISINOS.
Other materials. ULVG 9477a/9477b.
Occurrence. Upper part of the profile (PTM, Santa Maria
Formation).
Description. Incomplete (without apices) and triangular leaves,
70e80 mm long and 40e50 mm wide in the preserved form, with a
gently widened petiole (15 mm long, 2 mm wide in the holotype)
that apically originates a gradually and symmetrical tapered lam-
ina, two to three times deeply dissected. The partially preserved
apices suggest that an extra forking could be present. The surfaces
of its four broad proximal segments (1 cm wide) shows a gentle
relief, prominent in the middle portion, and grooved at the margins
(0.5e1 mm wide). Bifurcated basal veins enter into segments and
become fine striations.
Remarks. The nearly indistinct petiole and the symmetrical
wedge deeply dissected lamina carrying broad segments, relate this
form to Sphenobaiera (Table 1). The general size, and the segments
with broad depressed margins, covered by few and weak visible
veins, give consistence to the proposition of the new species.
Moreover it leads to the inclusion of Sphenobaiera sp. C (from the
Upper Triassic Can ~ adon Largo Formation, in South Argentina),
described by Gnaedinger and Herbst (1999), into the Sphenobaiera
tenuinervis described herein. The Brazilian material has the
advantage of exhibiting details of the venation, absent in the
Argentine samples. The exclusive morphological features pointed
above, up until now unknown to other Ginkgo-related fossils from
Gondwana, gives consistency to the new species, despite the scarce
known materials. However, except by a greater number of seg-
ments and low developed marginal grooves S. tenuinervis have
similar morphological features to that of Sphenobaiera sulcata sp.
nov. herein described.
S. tenuinervis occurs in the upper levels of the profile, where
Table 2
Age and places of occurrence of ginkgophytes in the Triassic of Gondwana. Observations: þ, data from this work. Orange colors and its variants (clear columns in black and
white version) indicate seasonal drier areas under the influence of monsoonal climates, green ones (gray in black and white version) those more wet, with plant fossils
occurring in coal measure deposits.
424 R. Barboni, T.L. Dutra / Journal of South American Earth Sciences 63 (2015) 417e435

Fig. 4. AeB, Baiera sp. 1 (ULVG 9761a), characterized by linear segments and distinct petiole; B, detail of the petiole showing the thickened nature and basal expansion; C,
Sphenobaiera tenuinervis sp. nov. (Holotype, ULVG 10880); D, Sphenobaiera sulcata sp. nov. (Holotype, ULVG11386) leaf and the characteristic longitudinal grooves; EeF, Sphenobaiera
sulcata sp. nov. (ULVG 9763), partial lamina and a detail of the leaf blade, highlighting the leathery aspect; GeI, Sphenobaiera insecta Anderson and Anderson (ULVG9502). In G,
incomplete leaf impression; H, basal part of the leaf (arrow indicates the two basal veins); I, detail of the segment. The arrows indicate the presence of galls. Scale bars:
Figs. AeC ¼ 10 mm; DeG ¼ 5 mm; HeI ¼ 2.5 mm.

associates with Baiera africana, Sphenobaiera sulcata nov. sp., Neocalamites sp., undetermined seeds and carapaces of
Dicroidium obtusifolium, D. odontopteroides, Xylopteris elongata, Spinicaudata.
Johnstonia stelzneriana, Pteruchus, Taeniopteris, Rissikia media, Sphenobaiera sulcata sp. nov.
 R. Barboni, T.L. Dutra / Journal of South American Earth Sciences 63 (2015) 417e435
(Fig. 4DeF)
Synonym
1989 Sphenobaiera sectina Anderson and Anderson, plate 63,
Figs. 4 and 19.
1999 Sphenobaiera sectina Gnaedinger and Herbst, Fig. 10EeF.
2007 Sphenobaiera sectina Holmes and Anderson, Fig. 24.
Diagnosis. Wedge-shaped, apparently coriaceous and sym-
metrical leaves, with a unique shallow-dissected lamina. Each
segment presents two marginal grooves and a central high relief
area. Though presenting a weak preserved venation pattern, at least
five longitudinal veins are visible in each lamina.
Derivation of name. Refers to the three longitudinal grooves
over the segments.
Holotype. ULVG 11386, LaViGæa Collection, UNISINOS.
Material. ULVG 9403; 9537; 9763; 11019.
Occurrence. Upper part of the profile (PTM, Santa Maria For-
mation), Paran a Basin.
Description. Impression of a cuneate and thickened leaf;
46e48 mm long and 11e17 mm wide in the preserved portion,
apparently once bifurcated in a very acute angle (15e20
). The two
wedge-shape segments (33 mm in size) have a wavy surface and
become broader apically (3.5e4 mm to 5e6 mm wide). Marginal
grooves, 1 mm wide. Fine veins covered the high relief intermediary
areas.
Remarks. This morphotype is rare in the assemblage, however
likewise seen in previous form, exhibits an exclusive morphology
when compared with previously known records of ginkgophytes.
However, in terms of the general morphology, the type of incision
and the depressed margin are reminiscent of some forms illus-
trated as S. sectina by Anderson and Anderson (1989, Pl. 63, Figs. 4
and 19) from the Molteno, South Africa, and by Holmes and
Anderson (2007, Fig. 24) from the Nymboidea Coal Measures, in
East Australia. The presence of longitudinal grooves over the seg-
ments approximates this morphotype to some forms of S. sectina
(Kurtz) Frenguelli, described to Can ~ adon Largo Formation by
Gnaedinger and Herbst (1999, Figs. 10EeF), that include part of the
S. argentinae material described by Azcuy y Baldoni (1990, Lam.1,
Figs. 10 and 13) in the synonym of S. sectina. The bigger and cori-
aceous leaves, with numerous, bifurcate and thick veins of
S. argentinae (Frenguelli, 1946; Artabe, 1985; Anderson and
Anderson, 1989; Morel et al., 2011) are also very distinct from the
herein described material.
Except by the low number of segments and bifurcations, the
form here described shares a general morphology (presence of
alternate high and low relief areas over the lamina) with the pre-
viously herein described S. tenuinervis.
Sphenobaiera insecta J.M. Anderson and H.M. Anderson 1989.
(Fig. 4GeI)
Material. ULVG 8946; 9502, from the LaViGæa Collection,
UNISINOS.
Temporal distribution. Late Triassic, except in Argentina where
they also occurs in the Middle Triassic levels (Table 2).
Occurrence. Exclusive from the middle levels of the profile
(PTM, Santa Maria Formation).
Description. Partially preserved (without apices), wedge-
shaped and elongated leaf (45 mm long, 15 mm wide at basal
part), deeply dissected and one time bifurcated. Petiole not distinct
(Fig. 4H). Sharp acute basal angle (19
), the lowest found among the
leaf morphotypes here studied. The lamina is divide by a deep
incision into two broad linear segments (4e5 mm wide) with two
basal veins, twice divided upwards and leading to at least five
parallel and well-visible veins in each segment.
Remarks. The diluting “petiole”, deeply dissected lamina and
bifurcate veins attest the relation to the genus Sphenobaiera
(Table 1). Judging by the linear-oblanceolate character of the leaf,
425
divided into two broad segments, and its well-defined basal sector
with two veins, approximates to S. insecta described to the Molteno
Flora by Anderson and Anderson (1989).
Additionally to the South Africa record, Sphenobaiera insecta was
also record in the Los Rastros Formation, in Argentina (Lutz et al.,
2011 and Table 2). According to Anderson and Anderson (1989)
the elongated laminas if preserved in fragmentary state (only
segments) could assume a similar morphology to that of Heidi-
phyllum and only distinguished in the presence of cuticles. How-
ever, the illustrated material of Sphenobaiera from Molteno
(Anderson and Anderson, 1989) shows a characteristic vein di-
chotomy distributed over the leaves and the lack of interveinal
striae, distinct from Heidiphyllum that only dichotomizes in the
basal part of the leaf (Retallack, 1981).
Evidences of planteinsect interactions are scattered over the
laminas of S. insecta from the PTM (Fig. 4I) and were also previously
described to the Molteno Formation (Scott et al., 2004). Here, their
oval shape and marked margins indicate oviposition activity.
However, the oval shape, with long axis aligned to the venation,
and the hardened out tissue, could also be found in galling activity
(Labandeira et al., 2007).
The S. insecta leaves herein described, still rarer in the assem-
blage, represents its first record in Brazil.
Sphenobaiera schenkii (Feistmantel) Florin 1936.
(Figs. 5AeG)
Material. ULVG 8900; 9279; 9505a/9505b; 9525A; 9757; 9772;
11095A; 11101; 11115a/11115b; 11151; 11198a/11198b, from the
LaViGæa Collection, UNISINOS.
Temporal distribution. Middle (Chile) to Upper Triassic
(Table 2).
Occurrence. Restrict to the middle part of the profile (PTM,
Santa Maria Formation).
Description. Big and wedge-shaped asymmetric leaves, the
most complete being 70 mm long and 45 mm wide. The leaf blade
(10 mm long and 2e2.5 mm wide) basally dilutes in width to form a
“semi” petiole. Four times deeply dissected lamina, originates 8e9
straight or, more rarely curved segments, each one with two to five
parallel weak veins. Basal acute angle varies from 25
to 30
. Seg-
ments varying in width, from 1.5 mm near the first dissection, to
3 mm apically. Longitudinal rows of red points over the laminas
represent the stomatal complexes (Figs. 5DeE).
Epidermal features (ULVG 11198a, Figs. 5FeG). The extraction
of the iron oxy-hydroxide coats in some sectors of the lamina
allowed the observation of molds and casts of some sectors of the
epidermis. The SEM analysis showed a thickened cuticle with costal
rectangular cells (110e191 mm long, 24 mm wide), longitudinally
aligned, with oblique contacts and straight smooth walls (Fig. 5F).
Anderson and Anderson (1989, 2003, pg. 222) illustrated similar
rectangular epidermal cells of S. “schenckii” from Molteno beds and
showed that they are rarer and restricted to specific sectors on the
adaxial side, and more common on the abaxial side. The dispersed
sunken stomata complexes (40 mm long, 25 mm wide) shows belong
to the brachyparaeactinocytic type, with five subsidiary cells in
radial arrangement, and occurs aligned along the intercostal areas,
(Fig. 4G). The subsidiary cells have prominent spherical papillae
projecting upwards (10e20 mm in diameter) and overarching the
oval pit mouth and the guard cells. Additional dispersed papillae
also appear in the surrounding epidermal cells (10e6 mm in
diameter), also longitudinally arranged. No striations linking the
stomatal complexes were observed.
Remarks. The morphological diversity of S. schenkii leaf mor-
photypes in the Triassic enables us to understand what motivated
Holmes and Anderson (2007) to classify them as members of the
Sphenobaiera schenkii complex. Associated with the transitional or
intergrading characters, it is difficult to determine if the apparently
426 R. Barboni, T.L. Dutra / Journal of South American Earth Sciences 63 (2015) 417e435

Fig. 5. AeB, Sphenobaiera schenkii (ULVG 9525), general aspect of the lamina; C, S. schenkii (ULVG 11198) showing the segments partially covered by iron coats. At center left a
fragment of a Dicroidium pinnae and planteinsect interaction; DeE, detail of C. The red points in E coincide with the longitudinally aligned rows of stomata; F-G, SEM images from
the epidermis, F. shows elongate epidermical cells preserved as a mold in the interior of the iron coats, G, a sunken actinocytic stomatal complex of C, showing the central pore
surrounded by six spherical papillae. Scale bars: AeC ¼ 10 mm; DeE ¼ 2, 5 mm; F ¼ 20 mm; G ¼ 10 mm.

consistent width of the basal part conforms to a distinct petiole. All
these characters make S. schenkii one of the most difficult forms in
the proposition of generic assignation. More detailed cuticle studies
on the genus would be valuable.
Previous epidermal features of ginkgophytes have been
described in the Permian (e.g. Glossophyllum florinii) and Triassic
(Sphenobaiera) periods, including those of S. schenkii. The
anatomical features presented attest the sunken stomata, the five
subsidiary cells radially arranged and the rounded papilla projected
over the stomatal aperture, or Florin rings (Krassilov, 1982;
Anderson and Anderson, 1989; Kerp, 1990; Wang et al., 2005;
Pott et al., 2007; Fischer et al., 2010; Pott and McLoughlin, 2011).
However, analysis of Baiera epidermis is almost unknown, with the
only exception being a study performed by Baldoni (1980) and
Gnaedinger and Herbst (1999, Table 1) to the Triassic, and those
from the Jurassic of the Northern Hemisphere (Wu et al., 2006). In
each case, the epidermal features found are the same as presented
by Anderson and Anderson (1989, 2003) for S. schenckii. However,
in the PTM material the number of stomata seems to be lower.
Sphenobaiera schenkii is a common component of the Triassic
Dicroidium floras (Table 2 and Supplementary online data), and
recorded in all landmasses of the Gondwana (Baldoni, 1980;
Anderson and Anderson, 1989; Stipanicic et al., 1996; Troncoso
and Herbst, 1999; Leppe Cartes, 2005; Holmes and Anderson,
 R. Barboni, T.L. Dutra / Journal of South American Earth Sciences 63 (2015) 417e435
2007; Troncoso and Herbst, 2007; Lutz et al., 2011; Escapa et al.,
2011). The abundant and overlapped leaf remains was assigned to
“autumnal banks” when detected in the Basin Creek Formation,
Australia (Holmes and Anderson, 2007; McLoughlin, 2010).
Once again, it is the first record of the species in Brazil. The leaf
imprints of S. schenkii in the PTM occur in association with vege-
tative and reproductive structures of Corystospermales (Dicroidium,
Zuberia, Xylopteris and Pteruchus), with other ginkgophytes, and
with molds of clam shrimps valves (Fig. 2E).
Morphotypes with intergrading characters and conflicting
generic affinities
The two following taxa are a good example (see introduction)
about the intergrading or transitional characters (or “problematic
contiguity”) that characterize Sphenobaiera and Baiera representa-
tives, and that make their distinction difficult in some cases (Bauer
et al., 2013a and Table 1).
Sphenobaiera steinmannii (Solms-Laubach) Anderson and
Anderson, 1989.
(Figs. 6AeB)
Basionym. Baiera (?) steinmanni Solms-Laubach 1899.
Material. ULVG 9766a/9766b; 9492, LaViGæa Collection,
UNISINOS.
Temporal distribution. Late Triassic (Carnian-Norian).
Occurrence. Middle-upper part of the profile (PTM, Santa Maria
Formation).
Description. Part and counterpart of a fragmentary impression
of lamina showing 12 (sample 9492) to 16 (sample ULVG 9766)
narrow and elongated segments (90 mm long and 1e2 mm wide on
average), with five parallel and longitudinal veins. Few irregularly
distributed dissections are preserved (Fig. 6A). Bases and apices are
absent.
Remarks. The presence of over four veins for each segment
relates this morphotype to Sphenobaiera, supported by the criteria
set out in Table 1. The previous record of similar morphotypes was
restricted to those identified by Solms-Laubach (1899) from La
Ternera Formation, in Chile. Originally assigned to Baiera, this
species was transfer to Sphenobaiera by Anderson and Anderson
Fig. 6. Ginkgophyte leaves with transitional characters. AeB, Sphenobaiera steinmannii (ULVG9766a), general view of the numerous and slender segments, irregularly bifurcate
(arrow). In B (ULVG9766b), a detail of the counterpart highlighting the segments and the five longitudinal veins; C, Sphenobaiera stormbergensis (ULVG 9771a), view of the
symmetrical dissected lamina with flattened, linear segments with round scars of planteinsect interactions. Scale bars: AeB ¼ 10 mm; C ¼ 5 mm.
427
(1989). The same status was used in forms from the Norian-
Raethian beds of Quebrada del Salitre in Northern Chile (Troncoso
et al., 2002; Troncoso and Herbst, 2012).
Since it lacks basal features, it is difficult to imply a generic af-
finity. However, this is a kind of preservation common for the
species, and does not impede our ability to detect some mixing of
features. The broad lamina, with many deep dissections and elon-
gate segments with linear margins is consider typical of Baiera,
while the presence of more than four veins by segment (except in
the apical ones) is a feature of Sphenobaiera.
Sphenobaiera steinmannii calls attention to its large size, only
comparable to that of Sphenobaiera stormbergensis, which makes it
one of the largely ginkgophyte leaf in the Triassic deposits of the
Gondwana. The great number of segments is a character also share
with B. africana.
In the Passo das Tropas Member it occurrence, still rare, co-
incides with the more diversified assemblages from the middle-
upper levels.
Sphenobaiera stormbergensis (Seward) Frenguelli 1948.
Type species: Sphenobaiera spectabilis (Nathorst) Florin 1936.
(Fig. 6C)
Basionym. Baiera stormbergensis Seward, 1903 (page 64, plate 8,
Fig. 3).
Synonym. See Artabe (1985).
Material. ULVG 9771/9771b.
Temporal distribution. LadinianeCarnian, Rhaetian?
Occurrence. Restricted to the basal part of the profile (PTM,
Santa Maria Formation).
Description. Cuneate asymmetric leaf (7 cm long, 5 cm wide)
preserved in a curved general form and composed by five flattened,
broad and linear segments (5 mm wide), originating from 3 to 4
irregularly distributed incisions. Basal acute angle is of 50
. In the
segments 5e6 veins are preserved, the central one more promi-
nent, with weak interveinal striae. The preserved apex shows a
straight and irregular dentate form. The covering of margins by iron
oxy-hydroxides suggests a coriaceous texture.
Remarks. Seward (1903), working on South African material,
428 R. Barboni, T.L. Dutra / Journal of South American Earth Sciences 63 (2015) 417e435
assigned this species to Baiera genus. Frenguelli (1948) afterwards
transferred it to Sphenobaiera drawing the attention to the large
size (12 cm long, 10 cm wide), deeply dissected lamina distally
divided in broad linear segments, and the presence of up to ten
parallel veins. He also pointed out that it could represent a bigger
form of B. “schenki”, as described by Feistmantel (1889). In size, the
form described by Seward (1903) can be only compared to that
assigned to S. stormbergensis by Gnaedinger and Herbst (1999) from
levels of the Can ~ adon Largo Formation, in Santa Cruz, Argentina,
which is still large (15 cm long). The Argentine morphotypes also
display some visible interveinal striae and the segments have
similar width to that described herein, the apical ones with 8 veins.
However, all those materials are twice the size of the PTM
morphotype and those from the Molteno Formation (Anderson and
Anderson, 1989), both up to 7 cm long. By the absence of a distinct
petiole (considered semi-petiolate), the Karoo Basin forms were
also assigned to Sphenobaiera (Anderson and Anderson, 1989).
Other shared features are the wedge-shaped and curved form of
the leaves, the number of veins in the segments and the broad and
flattened form.
Sphenobaiera stormbergensis, with a variable number of veins in
the segments, occurs in other basins of Argentina, Chile (Troncoso
and Herbst, 1999; Artabe et al., 2007), and at Nymboida, in
Australia (Holmes and Anderson, 2007). The latter have well-
preserved basal parts that show a short and robust semi-petiole,
to which many segments are associated.
By the criteria expressed in Table 1, those features fall into
Baiera, a procedure also adopted by Lutz et al. (2011) to the Los
Rastros Formation morphotypes. It seems clear that the taxonomic
insertion of some forms related to the two genera is still complex
and surrounded by many dubious aspects.
In the PTM beds S. stormbergensis is associated with diversified
Corystospermales, leaves of other ginkgophytes, with Hamshawvia,
and with carapaces of Spinocaudata conchostracans. Round single
foliar scars, with prominent reaction rims over the segments, seem
to represent oviposition of insects (Fig. 6C).
Reproductive structures associated to Ginkgophyte leaves
The following female and male fructifications were also identi-
fied in the South African Molteno floras where they occur in organic
connection with leaves related respectively to Sphenobaiera
schenkii and S. sectina. By their exclusive character in relation to
other ginkgoopsid, Anderson and Anderson (2003) include all those
forms (vegetative and reproductive ones) in the order
Hamshawviales.
Order Hamshawviales Anderson and Anderson, 2003.
Genus Hamshawvia Anderson and Anderson.
Type species. Hamshawvia baccata Anderson and Anderson,
2003.
Hamshawvia balausta sp. nov.
(Fig. 7AeE)
Diagnosis. Female strobili, basal axis or stalk apically bifurcated,
with a single distal pair of elliptic and pedunculated megasporo-
phylls of wavy and coriaceous surface, sinuous margins and
longitudinally folded. Ovules/seeds disposed into five oval cavities
(seminal fields) on each side of the central axis and separated by
fine walls, slight marked in the interior side and well visible in the
outer surface.
Derivation of name. Refers to its similarity with the modern
pomegranate balausta fruit, with a leathery pericarp and seeds
disposed in cavities (locules).
Holotype. ULVG 11176, LaViGæa Collection, UNISINOS.
Material. ULVG 11175A; 11177; 11183 aC.
Occurrence. Through all the Passo das Tropas stream profile
(levels 5, 6, 11 and 13) (PTM, Santa Maria Formation).
Description. Female structure with an incomplete stalk, apically
divided into two peduncles (1 mm width, angle of 30
), each one
giving place to a broadly ovate strobili, bilaterally symmetrical
(12e14 mm long and 8e9 mm wide). Distinct samples exposed the
inner (ventral, Fig. 7AeB) and external (dorsal) part (Fig. 7CeD) of
the megasporophyll, the first with the proposed five ovate seminal
fields (6e7 mm long and 2e2.5 mm wide) in each side of the
central vein (Fig. 7E), well-visible in the outer surface. Small and
round ovules/seeds (0.3 mm in diameter), dispersed and rarer in
the internal view (arrows in Figs. 7AeB), seem to connect to walls
limiting the seminal fields. In the dorsal surface (Fig. 7C) it is
possible to see a typical hemi-lamina of Hamshawvia, preserved in a
folded position and showing to be supported by two curved or
pending peduncles, probably due to the heavier (mature ones?) and
bigger seeds contained within (0.5 mm in diameter). The crenu-
lated margins of those structures are also well-visible (Fig. 7D). A
reconstruction of the ventral surface of the megasporophyll is
propose in Fig. 7E.
Remarks: This new species of female strobili differs from the
previously known Hamshawvia species from the Molteno Forma-
tion (Anderson and Anderson, 2003), since it contains a high
number of seminal fields and ovule/seeds, and wavy surface and
margins. However, as it happens with forms found in South Africa,
each sample seems to represent distinct stages of maturation (e.g.
H. baccata). The form herein chosen to represent the holotype
(Fig. 7CeD) probably refers to a mature head due to the apparently
more coriaceous character and the slightly bigger and expressive
number of seeds. In some other aspects, H. balausta sp. nov. co-
incides with the morphological and reproductive characters of
H. baccata, such as the general size and form of the megasporophyll.
H. baccata is distinct however, by showing a lower number of ovule/
seeds in radial disposition around the central axis.
The new species also differs from Hamshawvia longipedunculata
mostly because of its ovate form and the coriaceous nature of its
ovuliferous lamina (see below), which are analogous distinctions to
those that differentiate H. baccata from H. longipedunculata
(Anderson and Anderson, 2003). In order to confirm that those
kinds of heteromorphies were not influenced by taphonomic pro-
cesses (or stages of maturation) it is important to observe the
relation among the megasporophyll and its peduncle, a character
not influenced by the diversified sections shown in each mold or
cast.
H. balausta sp. nov. still shares features with Hamshawvia lini-
semina, those being the size and the five seminal fields in each
valve. Although, in this species the seminal fields are linear-
rectangular in shape and contain a unique ovule. In Hamshawvia
octosemina, the eight big seeds/ovules are even more distinct.
H.amshawviadistichos Holmes and Anderson from the Basin
Creek Formation, in Australia, has also a similar megasporophyll in
terms of form and size, but as in H. octosemina it contains eight
ovules/seeds disposed in two rows on each side of the mid-rib
(Holmes and Anderson, 2007).
In Argentina Hamshawvia is known by a unique form preserved
in the Cacheuta Formation (Cuyana Basin). Originally assigned to
Umkomasia by Frenguelli (1942), it was later transferred to Ham-
shawvia (Hamshawvia cacheutensis) by Anderson and Anderson
(2003). H. cacheutensis has three elliptical round seeds, inserted
into radial ridges and aligned on each side of the mid vein. An
analysis of Frenguelli's (1942) material confirmed the generic af-
finity proposed by Anderson and Anderson (2003).
Hamshawvia longipedunculata Anderson and Anderson, 2003.
(Fig. 7F)
Material. ULVG 9759, from the LaViGæa Collection, UNISINOS.
Temporal distribution. Ladinian (Australia)eCarnian (South
Africa).
Occurrence. In the lower e middle boundary (level 5) of the
 R. Barboni, T.L. Dutra / Journal of South American Earth Sciences 63 (2015) 417e435 429

Fig. 7. Ginkgophyte reproductive structures. AeE, female strobilus of Hamshawvia balausta sp. nov. (ULVG 11175A): A, general ventral or internal view of the apical pair of
megasporophyll. Points of iron deposits organized in transversal rows (arrow) seems suggests the ovule/seed disposition (arrow); B, detail of A, showing the probable ovules or
immature seeds disposed along a central wall (or mid-rib) of the seminal fields (arrow); C, holotype of H. balausta sp. nov. (ULVG 11176), partially folded megasporophylls, showing
seminal fields (SF) and probable seeds in distinct maturation stages linked to vascular traces (black arrow); D, detail of C to highlight the crenulated margins (black arrow) and well-
developed seeds (white arrow); E, reconstruction of H. balausta sp. nov. megasporophyll with the disposition of seminal fields and ovules/seeds; F, Hamshawvia longipedunculata
(ULVG 9759), and the elongate and apically bifurcated axis supporting one of the rounded and fleshy megasporophylls; G, fertile heads (microsporophyll) of Stachyopitys cf.
S. lacrisporangia (ULVG11010) with numerous clavate microsporangia, alternately inserted. Scale bars: AeC, E, G ¼ 5 mm; D and F ¼ 2.5 mm.

profile (PTM, Santa Maria Formation).
Description. A three dimensional impression of a partially
preserved fleshy megasporophyll (1 cm in diameter), showing one
of the two round ovuliferous lamina, inserted over an erected and
short peduncle (17 mm long, 0.7 mm wide), originated from the
apical bifurcation (angle of 20
) of the straight and erected axis
(1.4 mm wide). Veins or striations radially diverge from the
peduncle and central vein with the distal end coinciding with a
few round points (immature seeds?), of 0.7 to 0.25 mm in
diameter.
Remarks. H. longipedunculata differs from other species of
Hamshawvia due to its characteristic round megasporophyll, with a
smooth surface of an apparent fleshy nature and erect disposed
peduncle that diverts into low angles from the axis. However,
comparisons between the herein studied material and the African
form in respect to the ovules/seeds disposition are impossible due
430 R. Barboni, T.L. Dutra / Journal of South American Earth Sciences 63 (2015) 417e435
to its 3D preservation that only allowed us to see the external
surface or a section of the megasporophyll structure.
H. longipedunculata specimens from the Molteno Flora
(Anderson and Anderson, 2003) also occurs in the Nymboida Coal
Measures in Australia, but like here, without ovules/seeds pre-
served (Holmes and Anderson, 2007).
Despite the incomplete preservation of the PTM form, the shape
and arrangement of the probable ovules/seeds and the elongate
axis, better approximates to those present in the South African
H. longipedunculata. They only differ in the slightly larger sizes of
the African forms, which could be related to stages of maturation.
Anderson and Anderson (2003) found leaves of Sphenobaiera
schenkii and the immature strobili of H. longipedunculata attached
to the same short shoot in the Molteno Flora.
Genus Stachyopitys Schenck 1867
Type species: Stachyopitys preslii Schenk, 1867.
Stachyopitys cf. S. lacrisporangia Anderson and Anderson,
2003.
(Fig. 7G)
Material. ULVG 11010, from the LaViGæa Collection, UNISINOS.
Temporal distribution. Ladinian (Australia) e Carnian (South
Africa).
Occurrence. Unknown (ex situ sample).
Description. A lateral branch with delicate peduncles (2 mm in
diameter) showing at least four radially symmetrical microsporo-
phyll or fertile heads (8 mm long, 7.5 mm wide), each one bearing c.
14 elliptical microsporangia (2.3 mm long, 0.7 mm wide). The
delicate axis that supports this structure, seen only as a vestige in
the preserved form, seems to be a flexible axis or stalk.
Remarks. The fragmentary nature of this male strobilus only
allows us to relate them to S. lacrisporangia, characterized and
illustrated by Anderson and Anderson (2003, pl. 76) to Molteno
and thereafter identified in the MiddleeLate Triassic Nymboidea
flora of Australia (Holmes and Anderson, 2007). This affinity is
expressed in the general disposition of the fertile heads and in the
form and number of the microsporangia. However, the herein
described one is twice as big as that from Australia, and does not
contain the shallow longitudinal striations of the African repre-
sentatives. For those reasons, the proposed relation among the
forms is uncertain. In terms of the general morphology and radial
arrangement of the microsporophyll, it also has some similarities
to S. annularioides from Ipswich levels (Shirley, 1898). However,
the PTM impressions are smaller and present a greater number of
microsporophylls, densely inserted in a single point of the
apparently flexible axis.
The other Stachyopitys species proposed by Anderson and
Anderson (2003) have distinct characters and with more elon-
gated microsporangia (e.g. S. maziranus), or differ by the low
number and elongated shape of the microsporangia, as in
S. matatilongus, S. matatiramus and S. gypsianthus. Additionally,
S. rotundisporangia, as suggested in its epithet, has rounded
microsporangia. In South Africa, Anderson and Anderson (2003)
found S. lacrisporangia and S. sectina leaves attached to a bulbous
base.
Stachyopitys is not exclusive from Gondwana, having been found
in many Northern Hemisphere floras in both the Triassic and the
Lower Jurassic (Wcislo-Luraniec, 1992; Wang et al., 2007; Pott et al.,
2007; Van Konijnenburg-Van Cittert, 2010), where it is associated
with distinct kinds of foliage and ovulate organs (Zhou, 2009).
Firstly regarded as a conifer related structure, they were later linked
to an immature male reproductive structure of Baiera münsteriana
(Weber, 1968). However, the European and Chinese types are
distinct from the Gondwana representatives and from the speci-
mens described here.
Once again, the described form represents a first record to Brazil.
3. Discussion
3.1. Fossil assemblage and age constrains
The analysis made along the mudstones profile of the new
outcrop from Passo das Tropas Member, basal Santa Maria Forma-
tion, attests to a Ginkgophyte diversity previously unknown in the
Brazilian Triassic, and whose abundance (16%) is only lower than
that of the pteridosperms (70%). Similar role were detected in other
Gondwana floras, especially in those from Argentina and South
Africa (Anderson and Anderson, 1989, pag.12). Additionally, they
occur in a dispersed manner along all of the profile, yet with some
species showing high concentrations in specific levels (e.g.,
B. africana and S. schenkii leaves), suggesting deciduous habits (or
the autumnal banks of Holmes and Anderson, 2007). The other
components, like the isolated leaves of Taeniopteris (5%), and co-
nifers (3%), is low represented and allows infer a growth in more
distal areas from the waterbody.
Among the ginkgophytes, Baiera is rarer (B. africana and Baiera
sp. 1.) and Sphenobaiera represented by two new species
(S. tenuinervis and S. sulcate), by S. insect and S. schenkii, and by
some transitional forms like S. stormbergensis and S. steinmannii.
The cuticle characters confirm the relation of S. schenkii with
Sphenobaiera genus and with those described by Anderson and
Anderson (1989).
For otherwise, the Ginkgophyte affinity is confirmed by the
presence of two species of Hamshawvia and one of Stachyopitys,
additionally attesting the next relation with South African deposits
and, in a lesser degree, with those from Argentina and Australia
(Table 2, Supplementary online data). The occurrence of Spheno-
baiera cf. S. steinmannii, exclusive from the Upper Triassic assem-
blages of Chile, is an additional support with the South American
deposits.
In terms of age, the presence of Baiera africana, Sphenobaiera
steinmannii and Hamshawvia longipedunculata, which until now are
restricted to the Carnian (Table 2), allows us to extend the previous
AnisianeLadinian age assigned to the basal part Santa Maria For-
mation to include Carnian deposits. This inference have support
also in the Carnian age proposed to the associate coleopterans
fauna (Pinto, 1956; Martins-Neto et al., 2011), which correlates with
those from Argentina (Lara et al., 2012; Corre ^a, 2014).
In South America, similar fossil plant assemblages were pre-
served in Northwest Argentina, at Cuyana and Ischigualasto-Villa
Unio  n rift basins, dated to late Middle Triassiceearly Late Triassic
(Stipanicic and Marsicano, 2002). The Agua de la Zorra Formation,
in Mendoza (Ottone et al., 2005) and the lacustrine deposits of the
Los Rastros Formation (Mancuso and Marsicano, 2008; Lutz et al.,
2011), share some comparable plant fossils and the paleoenviron-
ment, with the only exception been the strong influence of volca-
nism in Agua de la Zorra Formation. Taking into account the
difficult distinction between Baiera and Sphenobaiera in the fossil
assemblages (sometimes being arbitrary), the use of morphospe-
cies instead of the generic affiliation seems to give more adequate
results when the focus is on the establishment of age and paleo-
geographic routes of the ginkgophytes along the Triassic.
Distinct ages have been proposed to the Los Rastros Formation,
in accord with analyzed kind of data (stratigraphy, palinology and
macrofossils). Based on the ~227 Ma radiometric age obtained in
the lower part of the uppermost Ischigualasto Formation (Barredo
et al., 2011; basal Norian, to Gradstein et al., 2012), a Carnian age
seems to be the most parsimonious for Los Rastros, and was sug-
gested by the palinological content (Zavattieri, 2002). Meanwhile,
in South Africa the Carnian age is well-established (Anderson and
Anderson, 1983, 2003; Bordy et al., 2005). So, comparable ages
that includes lattermost Middle Triassic to the beginning of the
 R. Barboni, T.L. Dutra / Journal of South American Earth Sciences 63 (2015) 417e435
Carnian, can be suggested for the basal part of Santa Maria For-
mation due to the presence of ginkgophytes.
The new upper limit to the Carnian and the expansion in age to
Norian, proposed by Gradstein et al. (2012), associated with the age
data recently obtained to South Africa (Bordy et al., 2005;
Catuneanu et al., 2005; Abdala et al., 2012), and in Argentina
(Barredo et al., 2011), have a strong influence on the age of those
floras.
Finally, for biostratigraphical purposes it is important to take
into account that there are some preferential tendencies in the
proportional presence of Baiera and Sphenobaiera in the Triassic
assemblages (Table 2). The data obtained showed that Baiera be-
comes proportionally more common than Sphenobaiera through
the end of Triassic of the Gondwana (the only exception been the
western Argentina and Chile deposits). Along with Ginkgo and
Ginkgoites, Baiera is part of similar communities to those from
Laurasia in the same time (Taylor et al., 2009).
3.2. Occupation of the Gondwanaland by the Ginkgophyta in the
Triassic
The ginkgophyte distribution compiled for the Triassic of
Gondwana (Table 2; Supplementary online data) shows that
ancient forms of ginkgophytes were recorded in the Early Triassic of
Northern Argentina (Saportea) and India (Sidhiphyllites).
In the Middle Triassic, more typical Mesozoic physiognomies
appear and diversify, firstly in the Anisian-Ladinian Basin Creek
Formation, Nymboidea, in Australia (Holmes and Anderson, 2007),
and after then, in Potrerillos and Cacheuta formations, Argentina
(Spalletti et al., 1999). In the latter area, following a decrease in
abundance, the ginkgophytes is another time recorded in the upper
levels of Cortaderita Formation (San Juan Province), the source beds
for the material of Frenguelli (1942, 1948) and corresponding to the
late Middle Triassiceearly Late Triassic OL Biozone of Spalletti et al.
(1999). According to Spalletti et al. (2003), the upper Cortadiretian
Stage coincides with the transition between evergreen to decidu-
ous subtropical vegetations, and a maximum diversification of the
Dicroidium floras. The Molteno assemblages (Anderson and
Anderson, 1989, 2003) attest that these events only took place in
the Carnian at South Africa, however it also could been influenced
by the extensive sampling made in the coal-rich facies of that unit.
To Zhou and Wu (2006), the Carnian was characterized by a
major change in leaf morphologies of the Ginkgo-related forms in
the Southern Hemisphere, mainly expressed by lamina and petiole
differentiation and by the reduction in number and increase in size
of the seeds. In the fossil woods, anatomical changes were also
detected, with mixed types giving place to the abietinoid ones
(Zhou, 2009; Gnaedinger, 2012).
Less diversified, the ginkgophytes are also characteristic from
the Ipswich Coal Measures, in Australia (Walkon, 1917; Jones and
De Jersey, 1947; Hill et al., 1965; Pattemore and Rigby, 2005;
Webb and Mitchell, 2006) and in those from East Antarctica
(Rigby, 1985; Escapa et al., 2011). The Carnian Blackstone Formation
from Australia includes the youngest flora containing ginkgophytes,
with a community that mainly includes Ginkgo and Ginkgoites
(McLoughlin, 2013).
In Argentina, the first “Jurassic-like” form appears in the Triassic
levels of the northern basins, accompanying the decrease of the
Dicroidium floras (Supplementary online data). In the south, a mix
of advanced (Baiera furcata and Sphenobaiera taeniata) and more
conservative forms (B. africana and S. robusta) occur in Paso Flores,
Llantenes and Chihuido formations (Late Triassic DLM Zone from
Spalletti et al., 1999).
431
3.3. Taphonomy, paleoenvironment and paleoecology
The taphonomic analysis performed in the nearly 2- meter
section of the red mudstones from Passo das Tropas (PTM) outcrop
attests to a unique perturbation in its homogeneity. This process is
showing in the recurrent presence of iron rich beds or lenses that
sometimes covered the frond impressions with iron coats that
maintains the three dimensional aspect of its thicker or lignified
structures. Similar kind of authigenic preservation was previouslly
detected in works made with the younger mudstone deposits of
Caturrita Formation (Late TriassiceEarly Jurassic, Fig. 1), also con-
taining floral remains (Barboni and Dutra, 2013). In that region, iron
coats covered conifer twigs and reveals a composition by goethite
minerals, also preserving molds of the cuticule features in its
interior (Wilberger et al., 2008). Lele (1962) detected such kind of
preservation over Dicroidium fronds from the South Rewa Basin in
India.
They seem supports the presence of perennial lakes, with pe-
riodical variations of the base level, and periods of deep-water
conditions that favored suboxide or anoxic environments. Under
these constraints, anaerobic bacteria could act in the water-
sediment limit (Frankel and Bazylinski, 2003) and promote the
microbial Fe (II) oxidation processes (Melton et al., 2014). In the
PTM the great amount of organic matter preserved, also indicates a
rapid cover and cementation that enabled the sustainability of
aerobic bacteria during a short period.
According to Crowe et al. (2007), the oxygen depletion and low
pH environment (that oxidizes the soluble Fe III and Mn IV), could
open space to anaerobic bacteria action that synthetize the fine iron
coatings (Fig. 3B, D and 4C), and to associated fine beds or lenses of
pure iron oxide-hydroxides (Fig. 2C). Apparently, the previous
phase of suboxide conditions furthered the genesis of the iron
covers and prevented or delayed the cuticle decay. Dunn et al.
(1997), in case of resulting from variations of the water level,
those iron deposits could be used to infer seasonal conditions of the
lake.
In the PTM deposition these taphonomic processes suggest that
in the broader fluvial context, some places sustain more permanent
water bodies, seasonally stratified and with no currents influence.
Bauer et al. (2013b) inferred similar conditions to the Middle
Triassic deposits of the Voltzia Sandstone, in France, where silte-
clay lenses with well-preserved leaves and seedlings represents
pond deposits, scattered between the fine-grained sandstone of the
fluvial sinuous channels.
The presence of Spinicaudata carapaces in the PTM profile
(Fig. 2E) supports oxygenated marginal areas on those perennial
lakes, with the likelihood of having deeper waters (anoxic/suboxic)
which protected the carapaces from predators (Gray, 1988; Rohn
et al., 2005). Gore (1986) gather evidences that apparently throw
doubt on the previous conception that Spinicaudata would indicate
temporary lakes, based mainly on the associate occurrence of fish
scales/bones and clam shrimps in Mesozoic lakes.
In paleofloristic terms the presence of Equisetales along the
profile suggests that wet soils exist around the lake, with the cor-
ystosperms and ginkgophytes been part of woodlands growing in
near elevated grounds or over the sand bars or river banks. The
rarity of conifer seems to indicate that they were part of a more
distal forest developed in drainer areas. On the other hand, the
abundance of Taeniopteris leaves supports a deciduous habit and/or
the “autumnal banks” commented herein. In this context, the
paleovegetation points to wooded open areas subjected to seasonal
dry atmospheric conditions, which prevented the growth of dense
forests.
When compared to previously studied basins from Gondwana,
such king of paleocommunity is similar to those found in the north
432 R. Barboni, T.L. Dutra / Journal of South American Earth Sciences 63 (2015) 417e435
of Argentina, in Chile and in Australia, differing in humidity from
those from the Coal Measures, in Molteno and Waterberg Basin
from Africa, with extensive coal deposits. However, in composition
the PTM flora is not so distinct of the megaplant associations of
Sphenobaiera and Dicroidium alluded by Anderson and Anderson
(1984). In the Late Triassic, the assemblages of southern
Argentina, similar to those from South Africa, shows that also water
restricted conditions exists still at higher latitudes (e.g. Passo Flores
deposits).
Taking those biological and environmental inferences into ac-
count, the narrow and in general fewer segments of the ginkgo-
phytes in the PTM, support the presence of monsoonal (seasonal)
conditions, as proposed by Parrish (1993) and Preto et al. (2010) to
the interior areas of Pangea, located in the boundary of tropi-
calesubtropical climate belts (lower than 30
of paleolatitude).
Similar conditions are indicated to the early Late Triassic palino-
logical provinces of Ipswich-Onslow, Easternmost Gondwana,
which age limits was recently revised (Dolby and Balme, 1976;
Cesari and Colombi, 2013). This proposal has support in the pres-
ence of heavily cutinized epidermis covered by numerous papillae,
and deeply sunken stomata in the cuticle (Pott et al., 2007).
More recently, such sclerophyllic leaf characters were shown to
be related, not only to a hydric deficiency, but to distinct environ-
mental and biological aspects, all being probably present in the
Triassic interior areas, e.g. poor soils, open areas with high lighting
incidence and leaf resistance to insect attack (Salleo and Nardini,
2000). According to Salleo et al. (1997), it may derive from
anatomical structures formerly adapted to humid environments,
and that assures the survival of the group in its later migration to
arid zones.
However, the presence of a diversified Dicroidium Flora in the
PTM indicates a climate amelioration that anticipates the more
continuous wet intervals that culminate in the deposition of the
thick sequence of mudstones (with a diversified tetrapod fauna)
that characterizes the uppermost Alemoa Member. Apparently the
“Carnian humid pulse” (Simms and Ruffel, 1989; Preto et al., 2010),
is testify along the Santa Maria Formation. To Holz (2015) it could
explain the explosion of life and diversity occurred in many parts of
the Gondwana, even during a drastic warm greenhouse interval.
4. Conclusions
The ginkgophytes from the basal Santa Maria Formation fill a
gap in the group distribution in Gondwana, enlarged and confirmed
by the presence of Hamshawvia and Stachyopitys reproductive
structures, accompanied by Sphenobaiera and Baiera leaves. The
exclusive Middle Triassic age previously proposed to the Passo das
Tropas Member (PTM) is now extend to include deposits and floras
of also a Carnian age. In this is consider the correlation between the
studied flora and those preserved in Molteno beds, South Africa,
and on the Los Rastros, Ischigualasto and Can ~ adon Largo formations
from Argentina. This broader interval of age opens the opportunity
of a coeval occurrence of floras and tetrapod faunas from the
Triassic of South Brazil.
In the paleoassemblage Sphenobaiera-related forms are the
most diversified, whereas Baiera africana is the most abundant,
displaying analogous condition to that from other assemblages in
westernmost Gondwana records. However, the distinct opinions
about the taxonomic relations of some forms, and intergraded
features between Baiera and Sphenobaiera, hamper more accurate
inferences about the distribution of the Ginkgophyta in the Triassic
of Gondwana.
The lacustrine diversified taphocenosis, inserted mostly in a low
sinuosity fluvial red-bed succession, confirms that warm and sea-
sonal climates existed in the interior areas of Gondwana in the
Middle to Late Triassic boundary, especially at latitudes lower than
30
S. The energy assumed from the fluvial context where the
fossiliferous mudstones are included indicates intervals of strong
rainfall and scarce vegetation cover in distal areas, which fed the
river systems. Taking both aspects into account (paleolatitude and
environment), the Southern Brazilian Dicroidium floras with gink-
gophytes show a stronger correlation to those from northwestern
Argentina and South Africa, than to the slightly older and devel-
oped at higher latitudes from Australia, Antarctica and South
America.
Acknowledgements
We are indebted to Silvia Gnaedinger (CECOAL/CONICET, Cor-
rientes, Argentina), for the ever-friendly time spent discussing the
Ginkgophyta affinities, and by the access to the paleobotanical
collection from CECOAL. To Heidi M. Anderson (ESI Palaeobotany,
University of Witwatersrand, Johannesburg, South Africa), for the
kind support with the bibliography and valuable assistance in
taxonomic and nomenclatural issues, and to Ubiratan F. Faccini
(UNISINOS), for the support in the facies analysis and stratigraphy.
To the anonymous reviewers, our deep thanks to the improvement
of the text and its results. Financial support was provided by the
Development of Brazilian Paleontology Program from CNPq (Na-
tional Council to Scientific and Technological Development, Procs.
401780/2010-4 and 401854/2010-8) and by FAPERGS (Research
Foundation from the State of Rio Grande do Sul, Proc. 1010122),
which made the field trips possible and improved the paleonto-
logical collection infrastructure. We are also grateful to the CAPES
Foundation, Brazil, for the Doctoral grant awarded to RB.
The authors also want to pay homage to Drs. John M. Anderson
and Heidi Anderson for their splendid and detailed work on the
Molteno floras.
Appendix A. Supplementary data
Supplementary data related to this article can be found at http://
dx.doi.org/10.1016/j.jsames.2015.08.001.
References
Abdala, F., Marsicano, C.A., Smith, R.M.H., Swart, R., 2012. Strengthening western
gondwanan correlations: a brazilian dicynodont (Synapsida, Anomodontia) in
the middle triassic of Namibia. Gondwana Res. 23, 151e162. http://dx.doi.org/
10.1016/j.gr.2012.07.011.
Anderson, J.M., Anderson, H.M., 1983. Palaeoflora of southern africa: molteno for-
mation. In: Part 1. Introduction/Part 2. Dicroidium, 1. A.A. Balkema, Rotterdam,
The Netherlands, p. 226.
Anderson, J.M., Anderson, H.M., 1984. The Fossil Content of the Upper Triassic
Molteno Formation, South Africa, 25, pp. 39e59. Palaeontologia Africana.
Anderson, J.M., Anderson, H.M., 1989. Palaeoflora of southern africa: molteno for-
mation (Triassic). In: Gymnosperms (Excluding Dicroidium), 2. Balkema, Rot-
terdam, The Netherlands, p. 567.
Anderson, J.M., Anderson, H.M., 2003. Heyday of the Gymnosperms: Systematics
and Biodiversity of the Late Triassic Molteno Fructifications. Strelitzia 15. Na-
tional Botanical Institute, Pretoria, p. 398.
Anderson, J.M., Anderson, H.M., Cleal, C.J., 2007. Brief History of the Gymnosperms:
Classification, Biodiversity, Phytogeography and Ecology. Strelitzia. National
Botanical Institute, Pretoria, p. 280.
Andreis, R.R., Bossi, G.E., Montardo, D.K., 1980. O Grupo Rosario do sul (Tri
assico) no
Rio Grande do sul. In: Congresso Brasileiro de Geologia, 31, Balnea rio Camboriú,
Sociedade Brasileira de Geologia, Anais, vol. 2, pp. 659e673.
Archangelsky, S., 1970. Fundamentos de Paleobot anica. Universidad de La Plata,
FCNM, Se rie Te
cnica y Didactica, 10, p. 340, 213 pl.
Archangelsky, S., 1996. Aspects of Gondwana paleobotany: gymnosperms of the
Paleozoic-Mesozoic transition. Rev. Palaeobot. Palynol. 90, 287e302.
Archangelsky, S., Cúneo, R., 1990. Polyspermophyllum, a new Permian gymnosperm
from Argentina, with considerations about the Dicranophyllales. Rev. Palaeobot.
Palynol. 63, 117e135.
Artabe, A.E., 1985. Estúdio sistema tico de la taphoflora triasica de Los Menucos,
Província Rio Negro, Argentina. Part 2. Cycadophyta, Ginkgophyta y Con-
iferophyta. Ameghiniana 22, 159e180.
 R. Barboni, T.L. Dutra / Journal of South American Earth Sciences 63 (2015) 417e435
Artabe, A.E., Morel, E.M., Spalletti, L.A., Ganuza, D.G., Zavattieri, A.M., 2007. La
paleoflora tria sica de Potrerillos, provincia de Mendoza, Argentina. Ame-
ghiniana 44, 79e301.
Azcuy, C.A., Baldoni, A.M., 1990. La flora tri asica del Grupo El Tranquilo, província de
Santa Cruz (Patagonia). Pt 3. Ginkgoales. In: 5th Congreso Argentino de Pale-
ontología y Bioestratigrafía, Tucum an, Argentina. Actas, vol. 7, pp. 109e115.
Bajpai, U., 1991. On ginkgoites leaves from the early permian of Rajmahal Hills,
Bihar, India. Ameghiniana 28, 145e148.
Baldoni, A., 1980. Baiera africana, una nueva especie de Ginkgoal del Tri asico de
Sud africa. Ameghiniana 17, 156e162.
Barboni, R., Dutra, T.L., 2013. New “flower” and leaves of Bennettitales from
Southern Brazil and their implication in the age of the Lower Mesozoic deposits.
Ameghiniana 50, 14e32.
Barboni, R., Silva, J.M., Lisboa, V.H.M., 2008. Te cnica de estabilizaça ~o de fo sseis em
pelitos laminados: aplicaç~ ao em níveis com plantas do Tri assico Superior da
Bacia do Parana , RS. Brasil. Gæa J. Geoscience 4, 38e41.
Bardola, T.P., Schmidt, I.D., Sommer, M.G., Schultz, C.L., 2009. Ginkgophyta wood in
the petrified forest of the Upper Triassic from Rio Grande do Sul, Brazil. Rev.
Bras. Paleontol. 12, 139e148.

Barredo, S., Chemale, F., Marsicano, C., Avila, J.N., Ottone, E.G., Ramos, V.A., 2011.
Tectono-sequence stratigraphy and UePb zircon ages of the Rinco  n Blanco
depocenter, northern Cuyo rift, Argentina. Gondwana Res. 21, 624e636. http://
dx.doi.org/10.1016/j.gr.2011.05.016.
Bauer, K., Kustatscher, E., Krings, M., 2013a. The ginkgophytes from the German
Kupferschiefer (Permian), with considerations on the taxonomic history and
use of Baiera and Sphenobaiera. Bull. Geosci. 88, 539e556.
Bauer, K., Grauvogel-Stamm, L., Kustatscher, E., Krings, M., 2013b. Fossil ginkgo-
phyte seedlings from the Triassic of France resemble modern Ginkgo biloba.
BMC Evol. Biol. 13 (177), 8.
Bordy, E.M., Hancox, P.J., Rubidge, B.S., 2005. The contact of the Molteno and Elliot
formations through the main Karoo Basin, South Africa: a second-order
sequence boundary. South Afr. J. Geol. 108, 351e364.
Bortoluzzi, C.A., 1974. Contribuiça ~o a
 Geologia da Regia ~o de Santa Maria, Rio Grande
do Sul, Brasil. Pesquisas 4, 7e86.
Bortoluzzi, C.A., Guerra-Sommer, M., Cazzulo-Klepzig, M., 1983. Tafoflora tri assica
da Formaça ~o Santa Maria, RS, Brasil. 1. Equisetales, Ginkgoales, Coniferales e
Pteridophylla. In: Congresso Brasileiro de Paleontologia, vol. 8, pp. 539e549. Rio
de Janeiro, Anais.
Braun, F.R., 1843. Beitr€ age zur Urgeschichte der Pflanzen. I. Die Fundorte von fos-
silen Pflanzen in der Umgegend von Bayreuth und Geschichte ihres Auffindens.
In: Zu, Münster G.G. (Ed.), Beitr€ age zur Petrefacten-Kunde. Buchner’sche
Buchhandlung, Bayreuth, p. 46.
Catuneanu, O., Wopfner, H., Eriksson, P.G., Cairncross, B., Rubidge, B.S.,
Smith, R.M.H., Hancox, P.J., 2005. The Karoo basins of south-central Africa. J. Afr.
Earth Sci. 43, 211e253. http://dx.doi.org/10.1016/j.jafrearsci.2005.07.007.
sari, S.N., Colombi, C.E., 2013. A new Late Triasssic phytogeographical scenario in
Ce
westernmost Gondwana. Nat. Commun. 4 http://dx.doi.org/10.1038/
ncomms2917 article number1889.
Corre^a, G., da, R., 2014. Fo  sseis de insetos em uma associaça ~o paleoflorística do
Tri
assico do Brasil: resposta tafono ^ mica e paleoambiental (Fossil insects in a
paleofloristic assemblage from the Triassic of Brazil; taphomic and paleo-
environmental aspects). Vale do Rio dos Sinos University, S~ ao Leopoldo, p. 40.
Undergraduate thesis, Biology. http://www.unisinos.br.
Crowe, S.A., Roberts, J.A., Weisener, C.G., Fowle, D.A., 2007. Alteration of iron-rich
lacustrine sediments by dissimilatory iron-reducing bacteria. Geobiology 5,
63e73. http://dx.doi.org/10.1111/j.1472-4669.2006.00086.x.
De Franceschi, D., Vozenin-Serra, C., 2000. Ginko biloba L. origin. Phylogenetical
approach. Comptes Rendus de l'Academie des Sciences Series III. Sci. la Vie 323,
583e592.
DiMichele, D.A., Chaney, D.S., Lucas, S.G., Kerp, H., Voigt, S., 2013. Flora of the lower
permian Abo formation redbeds, western Equatorial pangea, New Mexico. In:
Lucas, S.G., et al. (Eds.), Carboniferous-permian Transition in Central New
Mexico. New Mexico Museum of Natural History and Science, Bulletin, vol. 59,
pp. 265e288.
Dolby, J.H., Balme, B.E., 1976. Triassic palynology of the Carnarvon Basin, western
Australia. Rev. Palaeobot. Palynol. 22, 105e168.
Dunn, K.A., McLean, R.J.C., Upchurch Jr., G.R., Folk, R.L., 1997. Enhancement of leaf
fossilization potential by bacterial biofilms. Geology 25, 1119e1122.
Du Toit, A.L., 1927. The fossil flora of the Upper Karroo beds. Part. 2. Ann. South Afr.
Mus. 22, 289e420.
Dutra, T.L., 2002. Te cnicas e procedimentos para o trabalho com fo  sseis e formas
modernas comparativas (Technics and procedures to work with fossils and
extant comparative forms). In: PROIN/CAPES. UNISINOS, S~ ao Leopoldo, p. 55.
http://www.unisinos.br. mhgeo@unisinos.br].
Dutra, T.L., Barboni, R., 2014. Rediscovering the Brazilian Dicroidium flora: a “fossil-
lagerst€atte” in red beds?. In: International Congress of Palaeontology, 4, Men-
doza, Argentina, Abstracts, p. 701. http://www.ipc4mendoza2014.org.ar/
abstracts/ (accessed in 15.03.15).
Escapa, I.H., Taylor, E.L., Cúneo, R., Bomfleur, B., Bergene, J., Serbet, R., Taylor, T.N.,
2011. Triassic floras of Antarctica: plant diversity and distribution in high
paleolatitude communities. Palaios 26, 522e544. http://dx.doi.org/10.2110/
palo.2010.p10-122r.
Faccini, U.F., 2007. Tectonic and climatic induced changes in depositional styles of
the Mesozoic sedimentary record of southern Parana  Basin, Brazil. In:
Iannuzzi, R., Boardman, D. (Eds.), Problems in Western Gondwana Geology, I
433
Workshop: “South America e Africa correlations: du Toit revisited”, Gramado.
Extended abstracts, RS, Brazil, pp. 42e45.
Faccini, U.F., Paim, P.S.G., 2001. Estratigrafia de seque ^ncias em depo sitos con-
tinentais. In: Ribeiro, H.J.P.S. (Ed.), Estratigrafia de seque ^ncias e fundamentos e
aplicaço~es. Ed. Unisinos, S~ ao Leopoldo, pp. 341e389.
Feistmantel, O., 1889. Die Karoo-Formation und die duesselbe unterlagerden
Schichten. Pt 1. In: Überzschtliche Darstellung der geologisch-
palaeontologischen Verh€ altnisse Süd-Afrikas. Abhandlungen der Bo €hmischen
Gesellschaft der Wissenschaften, Math.-Natur. Cl., ser. 7, vol. 3 no. 6, 80 pp., 4
pls.
Fischer, T.C., Meller, B., Kustatscher, E., Butzmann, R., 2010. Permian ginkgophyte
fossils from the Dolomites resemble extant O-ha-tsuki aberrant leaf-like fruc-
tifications of Ginkgo biloba L. BMC Evol. Biol. 10, 3e17.
Florin, R., 1936. Die fossilen Ginkgophyten von Franz-Joseph-Land nebst Ero € rter-
ungen über vermeintliche Cordaitales mesozoischen Alters. I. Spezieller Teil.
Palaeontogr. B 81, 71e173.
Frankel, D.A., Bazylinski, D.A., 2003. Biologically induced Mineralization by bacteria.
Rev. Mineral. Geochem. 54, 95e114.
Frenguelli, J., 1942. Contribucio n al conocimiento de la flora del Gondwana Superior
en la Argentina VII, Umkomasia cacheutensis n. sp. Notas del Museo de La Plata.
Paleontología 48, 315e321.
Frenguelli, J., 1946. Contribuciones al conocimiento de la flora del Gondwana su-
perior en la Argentina. XXXIII. Ginkgoales de los estratos de Potrerillos en la
precordillera de Mendoza. Notas Museo de La Plata. Paleontología 11, 101e127.
Frenguelli, J., 1948. Estratigrafía y edad del llamado Retico en la Argentina, Gaea. An.
la Soc. Argent. Estratigr. Geogr. 8, 159e309.
Gnaedinger, S., 2012. Ginkgoalean woods from the Jurassic of Argentina: taxonomic
considerations and palaeogeographical distribution. Geobios 45, 187e198.
Gnaedinger, S., Herbst, R., 1999. “La Flora Tria sica del Grupo El Tranquilo, Provincia
de Santa Cruz (Patagonia). Parte VI. Ginkgoales”. Ameghiniana 36, 281e296.
Gordon Jr., M., Brown, R., 1952. Plantas tria ssicas do Rio Grande do Sul. Notas
Preliminares e Estudos, Divis~ ao de Geologia e Mineralogia. Dep. Nac. Produça ~o
Miner. 54, 1e7.
Gore, P.J.W., 1986. Triassic notostracans in the Newark Supergroup, Culpeper Basin,
northern Virginia. J. Paleontol. 60, 1086e1096.
Gradstein, M., Ogg, J.G., Schmitz, M.D., Ogg, G.M., 2012. The Geological Time Scale
2012. Elsevier, UK, p. 1140.
Gray, J., 1988. Evolution of the freshwater ecosystem: the fossil record. In: Gray, J.
(Ed.), Paleolimnology: Aspects of Freshwater Paleoecology and Biogeography.
Elsevier, Amsterdam, p. 214.
Guerra-Sommer, M., Klepzig, M., 2000. The triassic taphoplora from paran a Basin,
southern Brazil: an overview. Rev. Bras. Geocie ^ncias 30, 477e481.
Guerra-Sommer, M., Cazzulo-Klepzig, M., Ianuzzi, R., 1999. The Triassic taphoflora of
the Parana  Basin, southern Brazil: a biostratigraphical approach. J. Afr. Earth Sci.
29, 243e255.
Hammer, O., Harper, D.A.T., Ryan, P.D., 2001. Past - Palaeontological Statistics, ver.
1.75, 86 pp. [versio n original 2001]. Palaeontol. Electron. 4, 1e9.
Harris, T.M., Millington, W., 1974. The Yorkshire Jurassic Flora, IV. Trustees of the
British Museum (Natural History), London, p. 150.
Herbst, R., Troncoso, A., 2012. La flora Tria sica de la Quebrada Don ~ a Ines Chica,
Region de Atacama Chile. Gaea J. Geosci. 8, 55e66.
Hill, D., Playford, G., Woods, J.T., 1965. Triassic fossils of Queensland. Qld. Palae-
ontogr. Soc. 32.
Holmes, W.B.K., Anderson, H.M., 2007. The middle triassic megafossil flora of the
Basin creek formation, nymboida coal measures, new South Wales, Australia.
Part 6. Ginkgophyta Ginkgophya. Proc. Linn. Soc. N. S. W. 128, 155e200.
Holmes, W.B.K., Anderson, H.M., 2013. A synthesis of the rich Gondwana Triassic
megafossil flora from Nymboida, Australia. In: Tanner, L.H., Spielmann, J.A.,
Lucas, S.G. (Eds.), The Triassic System. New Mexico Museum of Natural History
and Science, Bulletin, vol. 61, pp. 296e305.
Holz, M., 2015. Mesozoic paleogeography and paleoclimates. A discussion of the
diverse greenhouse and hothouse conditions of an alien world. J. S. Am. Earth
Sci. 61, 91e107. http://dx.doi.org/10.1016/j.jsames.2015.01.001.
Jain, R.K., Delevoryas, T., 1967. A Middle Triassic flora from the Cacheuta Formation,
Minas de Petro leo, Argentina. Palaeontology 10, 564e589.
Jones, O.A., De Jersey, N.J., 1947. The Flora of the Ipswich Coal Measures:
Morphology and Floral Succession. Papers of the Department of Geology, vol. 3.
University of Queensland, p. 88.
Kerp, H., 1990. The study of fossil gymnosperms by means of cuticular analysis.
Palaios 5, 548e569.
Kerp, H., 1996. Post-Variscan late Palaeozoic Northern Hemisphere gymnosperms:
the onset to the Mesozoic. Rev. Palaeobot. Palynol. 90, 263e285.
Kokogia n, D.A., Spalletti, L.A., Morel, E.M., Artabe, A.E., Martínez, R.N., Alcober, O.A.,
Milana, J.P., Zavattieri, A.M., 2001. Estratigrafía del Tria sico argentino. In:
Artabe, A.E., Morel, E.M., Zamuner, A.B. (Eds.), El Sistema Tria sico en la
Argentina, Fundacio n Museo de La Plata “Francisco Pascasio Moreno”, La Plata,
pp. 23e54.
Krassilov, V.A., 1982. Early cretaceous flora of Mongolia. Palaeontogr. B 181, 43.
Labandeira, C.C., Wilf, P., Johnson, K.R., Marsh, F., 2007. Guide to Insect (And Other)
Damage Types on Compressed Plant Fossils. Version 3.0. Smithsonian Institu-
tion, Washington, p. 25.
Langer, M., Ribeiro, A.M., Schultz, C.L., Ferigolo, J., 2007. The continental tetrapod-
bearing Triassic of South Brazil. In: Lucas, S.G., Spielmann, J.A. (Eds.), The
Global Triassic. New Mexico Museum of Natural History and Science Bulletin,
vol. 41, pp. 201e218.
434 R. Barboni, T.L. Dutra / Journal of South American Earth Sciences 63 (2015) 417e435
Lara, M.B., Gallego, O.F., Tassi, L.V., 2012. Mesozoic coleopteran faunas from
Argentina: geological context, diversity, taphonomic observations, and com-
parison with other fossil insect records. Psyche a J. Entomology 1e14, 242563.
http://dx.doi.org/10.1155/2012/242563.
Lele, K.M., 1962. Studies in the Indian middle gondwana Flora: I. On dicroidium from
south rewa gondwana Basin. Palaeobotanists 10, 48e68.
Leppe Cartes, M., 2005. Paleobot sico del valle del rio Biobio, regio
anica del Tria  n del
Biobio, Chile. Concepcio n University, Chile, p. 149. PhD thesis.
Linol, B., Kasanzu, C., De Wit, M., 2014. Sediment sources and basin analysis of the
Central West Gondwana basin complex. In: Gondwana 15, North Meets the
South, p. 99. Abstract Books. http://www.gondwana15.org/wp-content/
uploads/Scientific-Programme_FINAL.pdf.
Lutz, A., Gnaedinger, S., Mancuso, A., Crisafulli, A., 2011. Paleoflora de la Formacio n
Los Rastros (Tria sico medio), Provincia de San Juan, Argentina. Consideraciones
taxono micas y tafono  micas. Ameghiniana 48, 568e588.
Mancuso, A.C., Marsicano, C.A., 2008. Paleoenvironments and taphonomy of a
triassic lacustrine system (Los rastros formation, Central-Western Argentina).
Palaios 23, 535e547. http://dx.doi.org/10.2110/palo.2007.p07-064r.
Mancuso, A.C., Gallego, O.F., Martins-Neto, R.G., 2007. The triassic insect fauna from
the los rastros formation (Bermejo Basin), La Rioja province (Argentina): its
context, taphonomy and paleobiology. Ameghiniana 44, 337e348.
Martins-Neto, R.G., Gallego, O.F., Tassi, L.V., 2011. The Triassic coleopteran fauna of
southern South America: morphometric variation of elytra, paleobiogeography,
and a phylogenetic approach. Gaea J. Geosci. 7, 1e18.
Mastroberti, A.A., 1995. Contribuiça ~o ao estudo do ge ^nero dicroidium (Gothan, 1912)
Towrown, 1957, na Formaça ~o Santa Maria (Rio Grande do Sul, Brasil), Bacia do
Parana . Acta Geol. Leopoldensia 44, 83e86.
McLoughlin, S., 2010. Ginkgo in Australia. Claystone Textbook. Aust. Age Dinosaurs
Mag. 8, 42e47.
McLoughlin, S., 2013. The fossil flora of Dinmore. Claystone Textbook,. Aust. Age
Dinosaurs Mag. 10, 40e49.
Melton, E.D., Schmidt, C., Behrens, S., Schink, B., Kappler, A., 2014. Metabolic flexi-
bility and substrate preference by the Fe(II)-Oxidizing Purple Non-Sulphur
bacterium Rhodopseudomonas palustris Strain TIE-1. Geomicrobiol. J. 31,
835e843.
Mene ndez, C., 1951. La flora Mesozoica de la Formacio n Llantenes (Província de
Mendoza). In: Revista Instituto Nacional de Investigaciones en Ciencias Natu-
rales (Bota nica), vol. 2(3). Museu Argentino de Ciencias Naturales Bernardino
Rivadavia, pp. 147e261.
Meyen, S., 1987. Fundamentals of Paleobotany. Chapman & Hall, London/New York,
p. 432.
Milani, E.J., Faccini, U.F., Scherer, C.M., Araújo, L.M., Cupertino, J.A., 1998. Sequences
and stratigraphic hierarchy of the Paran a Basin (Ordovincian to Cretaceous),
Southern Brazil. Boletim do Instituto de Geocie ^ncias, IGUSP. Serie Científica 29,
125e173.
Moisan, P., Abad, E., Bomfleur, B., Kerp, H., 2010. A late triassic flora from Gomero
(Santa Juana formation), Chile. Neues Jahrb. für Geol. Pala €ontologie
eAbhandlungen 258, 89e106.
Morel, E.M., Artabe, A.E., Ganuza, D.G., Zún ~ iga, A., 2011. La paleoflora triasica del
cerro Cacheuta, provincia de Mendoza, Argentina. Bryopsida, Lycopsida, Sphe-
nopsida, Filicopsida y Gymnospermopsida (Corystospermales y Peltas-
permales). Ameghiniana 47, 3e23.
Naugolnykh, S.V., 2007. Foliar seed-bearing organs of Paleozoic Ginkgophytes and
the early evolution of the Ginkgoales. Paleontological J. 41, 815e859.
Ottone, E.G., Mancuso, A.C., Resano, M., 2005. Miosporas y algas chlorococcales de la
Formacio n Los Rastros, Triasico medio a superior del centro-oeste de Argentina.
Ameghiniana 42, 347e362.
Parrish, J.T., 1993. Climate of the supercontinent Pangea. J. Geol. 101, 215e233.
Pattemore, G.A., Rigby, J.F., 2005. Fructifications and foliage from the Mesozoic of
southeast Queensland. Memoirs Qld. Mus. 50, 329e345.
Pinto, I.D., 1956. Artro podos da Formaç~ ao Santa Maria (Tria ssico Superior) do Rio
Grande do Sul, com notícias sobre alguns restos vegetais. Bol. Soc. Bras. Geol. 5,
75e87.
Pinto, I.D., Ornellas, L., 1974. A new insect Triassoblatta cargnini Pinto et Ornellas, sp.
nov., a Triassic Blattoid from Santa Maria Formation, South Brazil. An. Acad.
Bras. Cie ^ncias 46, 515e521.
Pfefferkorn, H.W., Mustafa, H., Hass, H., 1975. Quantitative charakterisierung ober-
karboner Abdruckfloren. Neues Jahrb. für Geol. Pal€ aontologie Abh. 150,
253e269.
Pott, C., McLoughlin, S., 2011. The rhaetian flora of Ro € gla, northern Scania, Sweden.
Palaeontology 54, 1025e1051.
Pott, C., Krings, M., Kerp, H., 2007. A surface microrelief on the leaves of Glosso-
phyllum florinii (?Ginkgoales) from the Upper Triassic of Lunz, Austria. Botanical
J. Linn. Soc. 153, 87e95, 19 figs.
Preto, N., Kustascher, E., Wignall, P.B., 2010. Triassic climates e state of the art and
perspectives. Palaeogeogr. Palaeoclimatol. Palaeoecol. 290, 1e10. http://
dx.doi.org/10.1016/j.palaeo.2010.03.015.
Retallack, G.J., 1980. Middle triassic megafossil plants and trace fossils from Tank
Gully, Canterbury, New Zealand. J. R. Soc. New Zel. 1, 31e63.
Retallack, G.J., 1981. Middle triassic megafossil plants from long Gully, near Ote-
matata, north Otago, New Zealand. J. R. Soc. N. Z. 11, 167e200.
Retallack, G.J., 1987. Triassic vegetation and geography of the New Zealand portion
of the Gondwana supercontinent. In: Elliot, D.H., Collinson, J.W., McKenzie, G.D.,
Haban, S.M. (Eds.), Gondwana Six, Stratigraphy and Paleontology, American
Geophysical Union, Geophysical Monograph, 41, pp. 29e39.
Rigby, J.F., 1985. Some triassic (Middle gondwana) floras from south Victoria Land,
Antarctica. In: COOPER, R. (Ed.), Hornibrook Symposium, New Zealand
Geological Survey Record, vol. 9, pp. 78e79. Extended Abstracts.
Rohn, R., Yanbin, Shen, Dias-Brito, D., 2005. A new ConiacianeSantonian con-
chostracan genus from the Bauru Group, south-east Brazil: taxonomy, palae-
obiogeography and palaeoecology. Cretac. Res. 26, 581e592.
Salleo, S., Nardini, A., 2000. Sclerophylly: evolutionary advantage or mere epiphe-
nomenon? Plant Biosyst. 134, 247e249. http://dx.doi.org/10.1080/
11263500012331350435.
Salleo, S., Nardini, A., Lo Gullo, M.A., 1997. Is sclerophylly of Mediterranean ever-
greens an adaptation to drought? New Phytol. 135, 603e612. http://dx.doi.org/
10.1046/j.1469-8137.1997.00696.x.
Schenk, A., 1867. Die fossile Flora der Grenzschichten des Keupers und Lias in
Franken. C.W. Kreidel’s Verlag, Wiesbaden, p. 232.
Schultz, C.L., Scherer, C.M.S., Barberena, M.C., 2000. Biostratigraphy of the southern
brazilian upper triassic. Rev. Bras. Geocie ^ncias 30, 495e498.
Scott, A.C., Anderson, J.M., Anderson, H.M., 2004. Evidence of planteinsect in-
teractions in the Upper Triassic Molteno Formation of South Africa. J. Geol. Soc.
161, 401e410. http://dx.doi.org/10.1144/0016-764903-118.
Seward, A.C., 1903. Fossil Flora of the cape Colony. Ann. South Afr. Mus. 4, 122.
Seward, A.C., 1919. Fossil Plants. A Text-book for Students of Botany and Geology. IV.
Ginkgoales, Coniferales, Gnetales. Cambridge University Press, Cambridge,
p. 543.
Shirley, J., 1898. Additions to the fossil flora of Queensland, mainly from the Ipswich
Formation, Trias-Jura System. Bull. Geol. Surv. Qld. 7, 1e5.
Silva, L.P., Iannuzzi, R., 2000. Cheirophyllum speculare Ce sari & Cúneo e ?Dicrano-
phyllum sp., novas formas afins as Ginkgophyta na seqüe ^ncia Gondva ^nica
Neopaleozo ica do Sul do Brasil. Rev. Univ. Guarulhos 5, 40e43 especial.
Simms, M.J., Ruffel, A.H., 1989. Synchroneity of climatic change and extinctions in
the late Triassic. Geology 17, 265e268.
Solms-Laubach, H., 1899. Beschreibung der Pflanzenreste von La Ternera. In: Solms-
Laubach, Hrsg. Von H., Steinmann, G. (Eds.), Das Auftreten und die Flora der
rha€tischen Kohlenschichten von La Ternera (Chile), Neues Jahrbuch für Miner-
alogie,Geologie und Paleontologie, vol. 12, pp. 581e592.
Spalletti, L., Artabe, A., Morel, E., Brea, M., 1999. Biozonacio  n paleoflorística y cro-
noestratigrafía del Tria sico argentino. Ameghiniana 36, 419e451.
Spalletti, L.A., Artabe, A.E., Morel, E.M., 2003. Geological factors and evolution of
Southwestern gondwana triassic plants. Gondwana Res. 6, 119e134.
Stewart, W.N., Rothwell, G.W., 1993. Paleobotany and the Evolution of Plants, 2nd
ed. Cambridge University Press, Cambridge, p. 521.
Stipanicic, P.N., Marsicano, C.A., 2002. Le xico estratigrafico de la Argentina, Volu-
men VIII. Tri asico. Asociacion Geologica Argentina, Serie “B”. Dida ctica Com-
plement. 26, 370.
Stipanicic, P., Herbst, R., Bonetti, M., 1996. Floras tria sicas. Revision y
actualizacio  n de la obra paleobota nica de Kurtz en la República Argentina. In:
Stipanicic, P., Hünicken, M. (Eds.), Academia Nacional de Ciencias, Actas, vol.
11, pp. 127e184.
Taylor, T.T., Taylor, E.L., Krings, M., 2009. Paleobotany, the Biology and Evolution of
Fossil Plants, 2nd. ed. Elsevier, USA, p. 1230.
Troncoso, A., Herbst, R., 1999. Ginkgoales del Tria sico del norte de Chile. Rev. Geol.
Chile 26, 255e273.
Troncoso, A., Herbst, R., 2007. Cuatro flo rulas del Tria
sico Superior del centro sur de
Chile (Cerro Ranguilí, Cerro Quilvo, Cerro Gupo y Río Quille n). Ameghiniana 44,
661e672.
Troncoso, A., Basso, M., Herbst, R., 2002. Redescubrimiento de Sphenobaiera stein-
manni (Solms-Laubach) Anderson y Anderson (Pinophyta, Ginkgoales). In:
Congreso Argentino de Paleontología y Bioestratigrafía, 8, Corrientes, Anales,
p. 42.
Uliana, M.A., Biddle, K., Cerdan, J., 1989. Mesozoic extension and the formation of
Argentine sedimentary basins. In: Tankard, A.J., Balkwill, H.R. (Eds.), Extensional
Tectonics and Stratigraphy of the North Atlantic Margins, vol. 46. American
Association of Petroleum Geologists Memoir, pp. 599e613.
Van Konijnenburg-Van Cittert, J.H.A., 2010. The Early Jurassic male ginkgoalean
inflorescence Stachyopitys preslii Schenk and its in situ pollen. Scr. Geol. Spec.
Issue 7, 141e149, 2 pls.
Walkon, A.B., 1917. Mesozoic floras of Queensland. Part 1. The flora of the Ipswich
and Walloon series, (d) Ginkgolaes, (e) Cycadophyta, (f) coniferales. Publ. Geol.
Surv. Qld. 259, 1e51.
Watson, J., Lydon, S.J., Harrison, N.A.A., 1999. Consideration of the genus Ginkgoites
Seward and a redescription of two species from the Lower Cretaceous of Ger-
many. Cretac. Res. 20, 719e734.
Wang, X., Duan, S., Geng, B., Cui, J., Yang, Y., 2007. Schmeissneria: a missing link to
angiosperms? BMC Biomed. Cent. Evol. Biol. 7, 1e14. http://dx.doi.org/10.1186/
1471-2148-7-14.
Wang, Y., Guignard, G., Thevenard, F., Dilcher, D., Barale, G., Mosbrugger, V., Yang, X.,
Mei, S., 2005. Cuticular anatomy of Sphenobaiera huangii (Ginkgoales) from the
lower jurassic of Hubei, China. Am. J. Bot. 92, 709e721.
Webb, J.A., Mitchell, M.M., 2006. Stratigraphy and palaeoflora of the triassic Council
Trench formation, central Victoria. Proc. R. Soc. Vic. 118, 113e127.
Wcislo-Luraniec, E., 1992. A fructification of Stachyopitys preslii Schenk from the
lower Jurassic of Poland. Cour. Forschungsinstitut Senckenberg 147,
247e253.
Weber, R., 1968. Die fossile Flora der Rha €t-Lias-Übergangsschichten von Bayreuth
(Oberfranken) unter besonderer Berücksichtigung der Coenologie. Erlanger
Geol. Abh. 72, 73.
 R. Barboni, T.L. Dutra / Journal of South American Earth Sciences 63 (2015) 417e435 435

Wilberger, T.P., Dutra, T.L., Barboni, R., 2008. An alise da ultra-estrutura da epiderme References to the Supplementary material
em folhas de coníferas preservadas de modo autige ^nico no Tri assico do RS. In:
Simpo sio de Paleobota ^nicos e Palino logos, 12, Floriano polis. Boletim de Resu-
Artabe, A.E., Zamuner, A., Ganuza, D.G., Spalletti, L.A., 1995. Novedades en la tafo-
mos, p. 229. sica de la quebrada de La Cortaderita, Barreal, vol. 1. Reunio n del
flora tria
Wu, X., Yang, X., Zhou, Z., 2006. Ginkgoalean ovulate organs and seeds associated sico del Calla Sur, 2, Bahía Blanca, Actas, p. 7. Provincia de San Juan,
Tria
with Baiera furcata-type leaves from the Middle Jurassic of Qinghai Province,
Argentina.
China. Rev. Palaeobot. Palynology 138, 209e225.
Barone-Nugent, E.-D., McLoughlin, S., Drinnan, A.N., 2003. New species of Rochip-
Yang, X.-J., Friis, E.M., Zhou, Z.-Y., 2008. Ovule-bearing organs of Ginkgo ginkgoidea
teris from the Upper Triassic of Australia. Rev. Palaeobot. Palynol. 123, 273e287.
(Tralau) comb. nov., and associated leaves from the Middle Jurassic of Scania,
Cantrill, D.J., Poole, I., 2013. The Vegetation of Antarctica Through Geological Time.
South Sweden. Rev. Palaeobot. Palynology 149, 1e17.
Cambridge University Press, p. 490.
Zavattieri, A.M., 2002. Anexo 4. Microfloras. In: Stipanicic, P.N., Marsicano, C.A. ~ os gimnospe rmicos de la Formacio  n Llantenes
Crisafulli, A., Herbst, R., 2010. Len
(Eds.), Lexico Estratigrafico de la Argentina. Volumen VIII. Tri asico. Asociacion
(Tri
asico Superior), Provincia de Mendoza. Argentina. Gӕa J. Geoscience 6,
Geolo gica Argentina, Buenos Aires, Serie “B” (Dida ctica y Complementaria), vol.
14e20.
26, pp. 318e321.
Crisafulli, A., Herbst, R., 2011. La flora tri asica del Grupo El Tranquilo, Provincia de
Zerfass, H., Lavina, E.L., Schultz, C.L., Garcia, A.J.V., Faccini, U.F., Chemale Jr., F., 2003. ~ os fo
siles, 48, pp. 275e288. Ameghiniana.
Santa Cruz (Patagonia): len
Sequence stratigraphy of continental Triassic strata of Southernmost Brazil: a tica y paleo-
Ganuza, D.G., Zamuner, A., Artabe, A.E., Spalletti, L.A., 1998. Sistema
contribution to Southwestern Gondwana palaeogeography and palaeoclimate. sica de Hilario-Agua de los Pajaritos (Formacio  n El
ecología de la flora tria
Sediment. Geol. 161, 85e105. zar), Provincia de San Juan, Argentina. Ameghiniana 35, 271e283.
Alca
Zerfass, H., Chemale Jr., F., Schultz, C.L., Lavina, E., 2004. Tectonics and sedi-
Goswami, S., Das, M., Guru, B.C., 2010. Paleonvironments in the Mahanadi Basin:
mentation in southern south america during triassic. Sediment. Geol. 166,
inferences from Mesozoic plants and ichnofossil diversity. Ecoscan 4, 7e14.
265e292.
Pal, P.K., 1984. Triassic plant megafossils from the Tiki formation, south rewa
Zhou, Z.Y., 1997. Mesozoic ginkgoalean megafossils: a systematic review. In: Hori, T.,
gondwana Basin, India. Palaeobotanist 32, 259e267.
Ridge, R.W., Tulecke, W., Del Tredici, P., Tre mouillaux-Guiller, J., Tobe, H. (Eds.),
Prasad, M., Lele, K., 1984. Triassic ginkgolean wood from the South Rewa Gondwana
Ginkgo Bilobada Global Treasure from Biology to Medicine. Springer Verlag,
basin, India. Rev. Palaeobot. Palynology 40, 387e397.
Tokyo, pp. 183e206.
Srivastava, S.C., 1983. Sidhiphyllites: a new ginkgophytic leaf genus from the Triassic
Zhou, Z.Y., 2009. An overview of fossil Ginkgoales. Palaeoworld 18, 1e22.
of Nidpur, India. Palaeobotanist 32, 20e25.
Zhou, Z.Y., Wu, X., 2006. The rise of Ginkgoalean plants in the early Mesozoic: a data € bler, R., Boppre, N., Fisher, O.W., 2009. Drei neue fossile Ho €lzer der
Süss, H., Ro
analysis. Geol. J. 41, 363e375. http://dx.doi.org/10.1002/gj.1049.
Morphogattung Primoginkgoxylon gen. nov. aus der Trias von Kenia. Feddes
Zhou, Z.Y., Zhang, B., Wang, Y., Guignard, G., 2002. A new karkenia (Ginkgoales)
Repert. 120, 273e292. http://dx.doi.org/10.1002/fedr.200911108.
from the jurassic Yima formation, Henan, China and its megaspore membrane
Torres, T., Philippe, M., 2002. Nuevas especies de Agathoxylon y Baieroxylon del Lías
ultrastructure. Rev. Palaeobot. Palynology 120, 91e105.  n del registro paleoxilolo
gico en el Jur
de La Ligua (Chile) con una evaluacio asico
Zhou, Z.Y., Quan, C., Liu, Y.-S., 2012. Tertiary Ginkgo ovulate organs with associated rica. Rev. Geol. do Chile 29, 3e19. http://dx.doi.org/10.4067/S0716-
de Sudame
leaves from North Dakota, U.S.A., and their evolutionary significance. Int. J. Plant
02082002000200001.
Sci. 173, 67e80.