Review of Palaeobotany and Palynology 257 (2018) 1–18

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Review of Palaeobotany and Palynology

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A Middle Triassic macroflora from southwestern Gondwana (Mendoza,

Argentina) with typical Northern Hemisphere elements: Biostrati-
graphic, palaeogeographic and palaeoenvironmental implications
Bárbara Cariglino a,⁎, Mariana Monti b, Ana María Zavattieri c
a
Museo Argentino de Ciencias Naturales “B. Rivadavia” (MACN), CONICET, Av. Ángel Gallardo 470, C1405DJR Buenos Aires, Argentina
b
YPF TECNOLOGÍA S.A. (Y-TEC), Avenida del Petróleo s/n, B1924CKU Berisso, Argentina
c
Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales (IANIGLA), CONICET, Av. Ruíz Leal s/n, Parque General San Martín, M5502IRA Mendoza, Argentina

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

Article history: A Triassic macrofloral assemblage is documented from the Quebrada de los Fósiles Formation, lower unit of the
Received 11 September 2017 Puesto Viejo Group (Mendoza, Argentina), southwestern Gondwana. The impression-compression flora incorpo-
Received in revised form 16 May 2018 rates fertile and sterile remains of lycophytes, sphenophytes, and the first seed ferns recorded from this unit.
Accepted 13 June 2018
Among the novel components, Ptilozamites, a typical Northern Hemisphere genus, is herein described for the
Available online 18 June 2018
first time from Gondwana. Ptilozamites longifolia sp. nov. consists of unipinnate leaves with an often forked rachis,
Keywords:
with long, linear to sub-rectangular pinnae attached by their whole bases. The veins are distinct, inserted directly
Triassic on the rachis, and run almost parallel to the margins, with few dichotomies occurring. The cuticle is thick, stomata
Puesto Viejo Group constituted by two sunken guard cells surrounded by 4–6 subsidiary cells fused to form a ring-like structure. The
South America association of Ptilozamites, Pleuromeia, Lepacyclotes, Equisetites and Neocalamites was common in many Middle
Ptilozamites and Late Triassic assemblages in Laurasia, whereas the Southern Hemisphere Triassic assemblages were charac-
Seed fern terized by the Dicroidium flora. A Middle Triassic age inferred from various elements constituting this macroflora
Lepacyclotes is also concordant with the recent radiometric dates assigned to the unit. The lithofacies associations indicate that
this flora grew and was deposited in the floodplains of a high-sinuosity fluvial system where volcanic ashes were
deposited episodically. This could have affected the development of the flora by favoring only those plants able to
cope with stressful environmental conditions, as exemplified by the fleshy stems and leaves with thick cuticles
present in Pleuromeia and Ptilozamites.
© 2018 Elsevier B.V. All rights reserved.

1. Introduction Cycadales, Ginkgoales, Gnetales and Coniferales (Artabe et al.,

The Early Triassic is marked by impoverished vegetation communi-
ties that recovered slowly after the end-Permian mass extinction. By the
Middle Triassic, however, multi-storey vegetation was re-established
and formed extensive forests across the globe, comprising abundant
seed ferns, cycads, ginkgoaleans, conifers, sphenophytes, lycophytes
and ferns.
Triassic palaeofloras are well-documented from several basins
in Argentina, which host the most stratigraphically complete re-
cords of southwestern Gondwana. Most plants in these assem-
blages are typical of the Dicroidium flora, which was dominated by
seed ferns (Corystospermales, Peltaspermales and Petriellales),
but also included representatives of Filicales, Bennetittales,
⁎ Corresponding author.
E-mail addresses: barichi10@gmail.com (B. Cariglino), marimontivaldes@gmail.com
(M. Monti), amz@mendoza-conicet.gob.ar (A.M. Zavattieri).
2001, 2007a and references therein).
The Triassic flora of the Puesto Viejo Group (San Rafael Block, Men-
doza province) has long been considered an exception, incorporating
only a few lycophytes and sphenophytes (Morel and Artabe, 1994;
Coturel et al., 2016), interpreted to reflect a small homogeneous wet-
land palaeoflora. However, recent work on the deposit has yielded
novel material, including abundant remains of seed fern foliage previ-
ously unknown in the Southern Hemisphere, and also new lycophyte
and sphenophyte strobili and stem fragments, suggesting that this
flora was much more developed and diverse than originally thought.
Furthermore, the presence of these new elements suggests that, con-
trary to long-held opinions (Bonaparte, 1981, 2002; Morel et al.,
2003), this flora developed during the Middle Triassic, refuting the
idea that it represents a “post-extinction recovery flora”, and making
its age accord with other lines of evidence (i.e., radiometric dating; see
Domeier et al., 2011 and Ottone et al., 2014). Last, and most interesting,
is the fact that this new assemblage shares more elements with those

https://doi.org/10.1016/j.revpalbo.2018.06.004
0034-6667/© 2018 Elsevier B.V. All rights reserved.
2 B. Cariglino et al. / Review of Palaeobotany and Palynology 257 (2018) 1–18
from the Northern Hemisphere than any other Triassic flora in
Argentina.
In this paper we describe and illustrate a new macroflora recov-
ered from three stratigraphic levels of the Quebrada de los Fósiles
Formation, lower unit of the Puesto Viejo Group. The specimens are
mostly impressions, but a few preserved as compressions yielded cu-
ticles diagnostic of seed ferns. We compare the composition of this
taphoflora with those from coeval localities both in Argentina and
rest of the world, and we examine the results in light of recently pub-
lished radiometric dates constraining the age of the formation.
Lastly, we discuss the palaeoenvironmental conditions in the basin,
based on both palaeofloristic and sedimentological data.
2. Geological setting
2.1. General context of the Puesto Viejo Group
The continental Triassic Puesto Viejo Group crops out in a small
depocenter located in the San Rafael Block (Fig. 1), Mendoza Province,
Argentina (34°70′ S, 68°20′ W), on the SSW margin of the main Triassic
accumulations of the Cuyo Basin (Ramos, 1993). The San Rafael Block
incorporates up to 1000 m of alluvial and fluvial sedimentary strata, to-
gether with rhyolitic pyroclastic flows, andesite lavas and basaltic shal-
low intrusives of the Puesto Viejo Group (González Díaz, 1964; Spalletti,
1994; Kokogián et al., 1999, 2001; Kleiman and Salvarredi, 2001;
Sepúlveda et al., 2007; Monti, 2014).
The Puesto Viejo Group is divided into two units (Stipanicic et al.,
2007), the Quebrada de los Fósiles Formation at the base and the Río
Seco de la Quebrada Formation at the top (Fig. 1). The tectono-
stratigraphic and sedimentological analyses made by Monti and
Franzese (2016) defined five depositional systems for the group:
1) debris-flow deposits, 2) channel-fill deposits, 3) floodplain deposits,
4) pyroclastic deposits, and 5) volcanic intrusives and effusives.
The lowermost Quebradada de los Fósiles Formation is characterized
by coarse epiclastic strata related to proximal alluvial-fan systems lo-
cated adjacent to the basin boundary. These levels were eventually
Fig. 1. Geographic location of the Quebrada de los Fósiles Formation. A. Triassic palaeogeographic map. B. Outcrops of the Puesto Viejo Group in the San Rafael Block. C. Geological map of
the Puesto Viejo Group in the study area.
covered by pyroclastic flows. Finer-grained strata of the upper section
of the Quebrada de los Fósiles Formation were extensively deposited to-
wards the centre of the basin. This section is interpreted to represent
meandering fluvial systems of low to high sinuosity that were also
interrupted by the emplacement of pyroclastic flows (Spalletti, 1994;
Monti and Franzese, 2016). The Río Seco de la Quebrada Formation ini-
tiates with coarse-grained deposits linked to the development of
braided to meandering low-sinuosity fluvial systems that were covered
by lava flows, and towards the top, by coarse-grained strata interpreted
to represent the distal sections of alluvial fans (González Díaz, 1972;
Monti and Franzese, 2016).
2.2. Lithofacies analysis of the Quebrada de los Fósiles Formation
The fossil material described herein derives from the type locality of
the Quebrada de los Fósiles Formation exposed at the homonymous
Quebrada de los Fósiles Creek (Fig. 1B). The section is ~ 200 m thick,
characterized by epiclastic and volcaniclastic sedimentation bracketed
at the base and the top by pyroclastic flow deposits (Fig. 2). Information
obtained from sedimentological and petrographic analyses (Monti,
2014) has been synthetized as simple lithofacies association descrip-
tions that circumscribe the main depositional processes (Table 1). The
lithofacies were divided according to their granulometry, sedimentary
structures and lithological composition, and the codes are modified
from Miall (1992) and Bridge (1993). For pyroclastic rocks the codes
used herein are based and modified from Branney and Kokelaar
(2002), whereas we developed our own classification for the volcanic
rocks (Monti and Franzese, 2016).
2.2.1. Epiclastic lithofacies associations
2.2.1.1. Traction-load dominated deposits.
a. Channels (CH): coarse-grained conglomerate lithofacies association
(Gt, Gmg, Gp; Table 1). These deposits are grouped into well-defined
lenticular bodies. The sedimentary succession (Fig. 2) starts with
 B. Cariglino et al. / Review of Palaeobotany and Palynology 257 (2018) 1–18 3

Fig. 2. Sedimentological section of the Quebrada de los Fósiles Formation at the type locality. A–B. Tuffaceous mudstones (AMZ2), with FTrf lithofacies of floodplain environments. Note the
presence of organic matter (OM) and volcaniclastic material (GM). C–D. Tuffaceous mudstones (AMZ1), with FTrf lithofacies of floodplain environments. E–F. Massive tuffaceous
mudstones (FTm; GzD), with laminated shales (Fl) and laminated carbonate levels (Cl) lithofacies of floodplain environments.

thick deposits of conglomerates, conglomeratic sandstones and
coarse-grained sandstones in trough cross-stratified sets, massive
or plane-bedded strata. Internally, sets of approximately 1 m in
thickness and fining-upward arrangement are identified. The chan-
nels represented by these lithofacies progressively transition up-
wards to facies representative of a lower flow-regime.
b. Sand bars (LA): coarse sandstones lithofacies association (Sp, Sl,
Sh; Table 1; Fig. 2). These facies are represented by lenticular
bodies with erosive bases and flat tops, always associated with
channels.
c. Crevasse (SB1): fine sandstones lithofacies association (STm, St;
Table 1; Fig. 2). These packages have sharp erosive bases,
4 B. Cariglino et al. / Review of Palaeobotany and Palynology 257 (2018) 1–18
Table 1
Lithofacies associations established for the Quebrada de los Fósiles Formation.
(Modified from Monti and Franzese, 2016)
Lithofacies association Geometry
Gt/Gmg; Gp Lenticular with erosive bottom and flat
top
Sp/Sl/Sh Lenticular with erosive bottom and flat
top
STm; St Tabular with flat bottom and top
Gp/Sm; Gh/SI Lenticular, abrupt to erosive bottom and
gradational to flat top
Fl/Fr; Fsm/FTm Tabular
Cl/Fr/FTrf/Fsm/Fl/FTm/FTl Tabular
mTL Tabular
Bi/Pp Tabular and domelike
gradational to flat tops and lenticular geometry, attributed to dis-
charge channels.
d. Crevasse splays (SB2): coarse sandstones lithofacies association
(Gh/Sl and Gp/Sm; Table 1; Fig. 2). These deposits have tabular
geometry with flat bases and tops and they are internally consti-
tuted by a fining-upward arrangement (Ghosh, 1987; Clemente
and Pérez-Arlucea, 1993; Monti and Franzese, 2016).
Both lithofacies associations (SB1 and SB2) are scattered within the
mudrock-dominated intervals of the succession being relatively thin
bodies formed during flooding of the fluvial system.
Collectively, these epiclastic deposits represent the main channels of
the high-sinuosity river system with a typical fining-upwards arrange-
ment and deposits of discharge during flooding events (Collinson,
1986; Clemente and Pérez-Arlucea, 1993; Spalletti, 1994; Bridge,
2006; Monti and Franzese, 2016).
2.2.1.2. Suspension-load dominated deposits.
a. Floodplain (FF1): siltstones and mudstones lithofacies association (Fl,
Fr, Fsm, FTm; Table 1; Fig. 2). These deposits are mainly represented
by fine-grained sediments consisting of thick tabular beds generated
by subaqueous sedimentation in marshes and abandoned-channel
lakes. They comprise thick intervals dominated by massive and lam-
inated greenish–gray mudstones, siltstones and whitish tuffaceous
mudstones with well-developed palaeosols evidenced by fine and
dense root cast systems.
b. Shallow water bodies (FF2): calcareous and tuffaceous mudstones
and siltstones lithofacies association (Cl/Fr/FTrf/Fsm/Fl/Tm; FTm/
FTl; Table 1; Fig. 2). These consist of thick tabular bodies that accumu-
lated in low-gradient parts of the depositional system. These deposits
developed locally without laterally contacting with the sandy chan-
nel facies. The presence of laminated calcareous levels together
with the gradation of massive to laminated facies favors interpreta-
tion of a subaqueous depositional environment (Plint and Browne,
1994) associated with non-permanent, shallow lacustrine bodies
(FF2).
The Quebrada de los Fósiles Formation fossil beds are located in the
floodplain facies of a high-sinuosity fluvial system that laterally inter-
grades with shallow lacustrine deposits (Spalletti, 1994; Zavattieri
et al., 2003, Monti and Franzese, 2016). The three levels of interest con-
sist of tuffaceous mudstones (AMZ2; Fig. 2A, B), tuffaceous mudstones
and limestones (AMZ1; Fig. 2C, D), and laminated siltstones interbed-
ded with light yellow-orange bio-laminated calcareous layers (GzD;
Fig. 2E, F) with light-gray to whitish mudstones having fine and densely
Architectural element Environment
Channel deposits CH
Sandy bars LA
Crevasse splay deposits or lobes of expansion SB1 Meandering
fluvial systems
Deposits of crevasse channels, also small channels of discharge SB2
Suspension-load, subaquatic deposits in floodplain FF1
Shallow bodies of water and incipient development of paleosols and FF2
carbonate levels in floodplain with fall tuff levels
Pyroclastic flow deposits TL
Subvolcanic intrusions and rocks derived from unconsolidated rocks VI Volcanic activity
distributed root casts indicating development of incipient palaeosols.
These levels also yielded abundant silicified megaspores, ostracod im-
pressions and few fish scales.
2.2.2. Volcanic lithofacies associations
2.2.2.1. Pyroclastic deposits.
a. Air fall tuffs (TL): rhyolitic air fall tuff (mTL; Table 1; Fig. 2). These are
preserved as thin deposits within lacustrine levels of the floodplains.
b. Pyroclastic flows (TL): rhyolitic pyroclastic flow deposits (mTL;
Table 1; Fig. 2). These are well-developed throughout the study
area as tabular bodies that cap the upper part of the Quebrada de
los Fósiles Formation (Monti and Franzese, 2016). These deposits
were formed by the collapse of eruptive columns associated with
nearby explosive volcanic activity (McPhie et al., 1993; Branney
and Kokelaar, 2002).
2.2.2.2. Volcanic intrusive rocks.
a. Subvolcanic intrusions (VI). These consist of basaltic intrusives (Bi;
Table 1; Fig. 2) with dome-shaped architecture (Monti and
Franzese, 2016). The basic volcanic rocks belonging to the Río Seco
de la Quebrada Formation deformed the fine deposits of the
Quebrada de los Fósiles Formation as intrusive sub-volcanic struc-
tures. The common presence of peperites (Pp; Table 1; Fig. 2) sug-
gests that these were formed by contact of the intrusive bodies
with unconsolidated and wet sediments (McPhie et al., 1993).
2.3. Age of the Quebrada de los Fósiles Formation
The Quebrada de los Fósiles Formation type section rests with ero-
sional unconformity on upper Choiyoi volcanics of Permian age
(González Díaz, 1972). The unit starts and culminates with ignimbritic
strata (Fig. 2), being the uppermost ignimbrite the boundary with the
overlying Río Seco de la Quebrada Formation.
Initial radiometric 40K/40Ar dating of the upper ignimbrite from the
unit estimated an age of 236 ± 10 Ma (Valencio et al., 1975). More re-
cently, SHRIMP U–Pb radiometric dating obtained from juvenile mag-
matic zircons in the same ignimbrite indicated an age of 235.8 ± 2 Ma
(Ottone et al., 2014).
On the other hand, 40Ar/39Ar isotopic dating carried out by Domeier
et al. (2011) on volcaniclastic and ignimbritic rocks from the basal levels
of the Quebrada de los Fósiles Formation indicated an age of ~245 Ma.
Similarly, the basal ignimbrite has been dated using LA-ICPMS U–Pb,
yielding a late Anisian age (Monti in Sato et al., 2015). Thus, the
Quebrada de los Fósiles Formation accumulated from the Middle (late
 B. Cariglino et al. / Review of Palaeobotany and Palynology 257 (2018) 1–18
Anisian) to Late (early Carnian) Triassic, being mostly constrained to the
Ladinian (Cohen et al., 2013, updated) (Fig. 2).
3. Materials and methods
Detailed geological sections were measured through the Triassic
continental sequence of the Puesto Viejo Group (Monti, 2014; Monti
and Franzese, 2016), including the Quebrada de los Fósiles Formation
section presented here (Fig. 2) and from whence the fossil plants were
recovered. The new plant-bearing levels (AMZ1 and AMZ2, recovered
in 2013), and the original level (GzD) mentioned by González Díaz
(1966), were stratigraphically placed on the measured section.
Fossil plants are preserved as impressions and compressions of var-
iable quality. Macrofossil specimens were photographed using a Canon
Powershot S5IS (8.0 megapixel) digital camera. Detailed illustrations
were obtained under a strong unilateral light at various angles to high-
light diagnostic characters, with a Nikon Ds-Fi1-U2 digital camera at-
tached to a Nikon SMZ800 stereomicroscope. Images were later
improved in contrast and standardized in color using Adobe Photoshop
CS5.
Cuticles were recovered and prepared following standard proce-
dures. Fragments of foliage with organic matter were macerated in
Schulze's reagent (KCLO3 and 30% HNO3), then neutralized with 10%
KOH and consecutively rinsed with distilled water. The pieces of cuticle
obtained were analyzed under SEM (Philips XL30 TMP) and light mi-
croscopy (Olympus BX-51), the latter photographed with an attached
Olympus C5060 digital camera at the Museo Argentino de Ciencias
Naturales “B. Rivadavia”, Buenos Aires.
Every fossiliferous rock slab is housed at the Museum of Natural His-
tory of San Rafael in Mendoza province, under the prefix MHNSR Pb-CN
1–63.
4. Description of the flora
4.1. Division Sphenophyta
Order Equisetales Dumortier, 1829
Family indet.
Genus Neocalamites Halle, 1908 emend. Bomfleur et al., 2013
Type-species: Neocalamites lehmannianus (Göppert) Weber, 1968
Neocalamites suberosus (Artabe et Zamuner, 1991) emend.
Bomfleur et al., 2013
Plate I, 1–3
Rock sample number. MHNSR Pb-CN 15, MHNSR Pb-CN 19, MHNSR
Pb-CN 22.
Stratigraphic level. AMZ1.
Description. Specimens consist of straight, narrow stems, with inter-
nodes at least 14.5 cm long (incomplete) and 1.6 cm wide, and slightly
expanded nodes, ~ 2 cm wide (Plate I, 1, arrow). The length to width
ratio of the internodes is at least 9:1. The internodes' surface is charac-
terized by conspicuous dense ribbing that crosses the nodes either in a
continuous or an alternate manner (Plate I, 3). There are 31 to 44 longi-
tudinal ridges per cm of width. Partially preserved leaves are evident in
specimen MHNSR Pb-CN 15, where their broadened bases fuse in a con-
tinuous rim (Plate I, 3). A detached branch 0.5 mm wide that probably
belongs to the stem is preserved on its right side (Plate I, 2, “br”).
Remarks. These sphenophyte stems are attributable to Neocalamites
based on the presence of long internodes with numerous conspicuous
ribs on the surface, and slightly swollen nodes where both opposite
and alternate longitudinal ridges are expressed. The assignation to
N. suberosus is founded on the fused basal leaves although not constitut-
ing a distinct leaf sheath characteristic of Equisetites, and differing from
other known Neocalamites species in which leaves are usually free for
their entire length (Bomfleur et al., 2013). The long internodal size of
our specimens make them similar to a congeneric species found com-
monly in the Triassic of Argentina, N. carrerei (Zeiller) Halle, 1908.
5
However, the latter has up to a 100 narrow leaves and typical circular
traces in the nodal lines that are not evident in our specimens (Artabe
et al., 2007b; Morel et al., 2015; Bomfleur et al., 2013).
The presence of Neocalamites sp. cf. N. suberosus in this unit was re-
cently mentioned by Coturel et al. (2016) based on regularly preserved
material from older collections, consisting of stems, nodal diaphragms
and strobili, the latter apparently borne on a long stalk departing from
a node. Although the presence of the long stalk was admittedly ‘obscure’
due to its placement under the stem, these authors used the assumed
existence of the stalk as the defining character to assign the specimen
to Neocalamites. Although we agree with the presence of attached stro-
bili (‘oval bodies’) in their material (see Coturel et al., 2016, fig. 5A–C),
we argue that those fertile structures were either sessile or on very
short stalks at the nodes of the stem. The ‘elongated stalk’ is most poten-
tially a preservational artifact, as it does not show a constant width, nor
it can be followed along its entire length. Both indicated strobili are
placed at the same level of the nodes; the example on the left is on
the upper node, whereas the one on the right is slightly detached
from the lower node, further suggesting the lack of long-stalked strobili.
If our interpretation is confirmed, the specimen described by Coturel
et al. (2016) would more accurately fit under Equisetites. The other
illustrated (and different) specimen assigned by these authors to
Neocalamites cf. N. suberosus, on the other hand, has similar features to
our specimens (Coturel et al., 2016, fig. 5D), such as the carination and
the alternation of the ridges across the node. Unfortunately, the lack of
a complete specimen prevents additional comparisons.
Family Equisetaceae Michaux ex DeCandolle, 1804
Genus Equisetites Sternberg, 1833
Type-species: Equisetites münsteri Sternberg, 1833
Equisetites sp.
Plate I, 4–12
Rock sample number. MHNSR Pb-CN 39 a–b, MHNSR Pb-CN 40,
MHNSR Pb-CN 44, MHNSR Pb-CN 57a-b.
Stratigraphic levels. AMZ1, AMZ2.
Description. Specimens MHNSR Pb-CN 39 a–b, MHNSR Pb-CN 40
and MHNSR Pb-CN 44 are all articulate vegetative stems, at least
16.5 cm long and 2.9 cm wide, with numerous internodes ranging
from 3.5 to 14 mm in length, decreasing towards the base and apex of
the stems, their length to width ratios being less than 1, and nodes
wider (~ 3.5 cm) than the internodes (Plate I, 4–8). Long and thin
branches about 4.7 cm long are present on MHNSR Pb-CN 44 at the
right side of the stem (Plate I, 5). The commissural furrows are well-
defined. Small, circular branch scars 0.5 mm in diameter are arranged
as a single or double line around the node. In addition, larger circular
to ovoid scars, ~ 7.5 mm in diameter (Plate I, 4), are evident at the
node level in various parts of the axis, indicating that the stems ramified
into larger lateral branches.
MHNSR Pb-CN 57a-b (Plate I, 9–12) consists of the part and counter-
part of a poorly preserved, decorticated stem impression, 13.4 cm long
(incomplete) and 4.4 cm wide, with a smooth external surface apart
from widely spaced (1.6 mm) longitudinal ribs (Plate I, 11). The poor
preservational state precludes the identification of further features;
however, to the left of the stem, there are at least two immature strobili
which are closely associated with the stem (Plate I, 9–10). The strobili
are attached to a short, thick, leafless shoot, 5.2 mm long and 2.9 mm
wide, which further divides into two smaller branches 2.5 mm long
and 1.3 mm wide, each strobilus being borne on a single axis (Plate I,
10). Strobili are circular to slightly ovoid, ~ 8 mm in diameter, com-
pact, and composed of at least seven sporangiophore heads. Their ar-
rangement, although not clearly visible, seems to be in 3–4 whorls.
Only the pentagonal heads (= discs) of the sporangiophores are vis-
ible, although their arrangement and size are obscured by the
preservational state. A detached pair of pentagonal sporangiophore
heads about 9 mm in diameter occur a short distance from the at-
tached strobili, suggesting they potentially belong to the same spec-
imen (Plate I, 9, 12).
6 B. Cariglino et al. / Review of Palaeobotany and Palynology 257 (2018) 1–18

Remarks. As observed by several authors (Meyen, 1987; Kon'no, since their boundaries are not clearly defined. The defining character
1962; Weber, 2005), the classification of isolated fossil arthrophyte veg- used to separate vegetative Neocalamites from Equisetites is the presence
etative stems preserved as impressions can often lead to confusion, of long, free leaves vs. reduced, adpressed to the stem leaves,
 B. Cariglino et al. / Review of Palaeobotany and Palynology 257 (2018) 1–18
respectively. Short leaves are seen at the side of specimen MHNSR Pb-
CN 40 (see Plate I, 8, black arrows), suggesting leaves are adpressed to
the stems but their observation is obscured by a preservational artifact.
Therefore, specimens MHNSR Pb-CN 39 a–b, 40, 44 are here assigned to
Equisetites based on their shared similarities, and differing from the
Neocalamites specimens previously described in the presence of re-
duced leaves. The placement of MHNSR Pb-CN 57 a–b under this
genus, on the other hand, should be cautious. The stem is badly
preserved and does not provide further clues, but the close presence
of strobili and detached sporangiophore heads similar to those
equisetalian-like strobili placed in Equisetites-Equisetostachys could sug-
gest its potential belonging to this genus (Kon'no, 1962; Kelber and van
Konijnenburg-van Cittert, 1998; Kustatscher et al., 2007). Contrary to
Neocalamites, in which the strobili are borne on long stalks, the repro-
ductive structures of Equisetites are either sessile or most commonly,
on short fertile shoots.
When compared to other Triassic Equisetites species, the fossils from
the Quebrada de los Fósiles Formation most resemble E. mougeotii
(Brongniart) Wills, 1910, E. arenaceous (Jaeger) Schenk emend. Kelber
et van Konijnenburg-van Cittert, 1998, E. aequecaliginosus Weber,
2005 and E. fertilis (Frenguelli) Frenguelli, 1944. In general terms, all
these species share almost the same vegetative features with our mate-
rial, such as large stems (‘giant horsetails’) with somewhat expanded
nodes, branches in verticils at the node level that leave an abscission
scar, short leaves fused for most of their length in a sheaths distally end-
ing in free acuminate teeth, and distinct commissural furrows. Some of
these species have been synonymized by various workers or are cur-
rently considered closely related (i.e., Weber, 2005), pointing to their
similar overall vegetative physiognomy. The main differences separat-
ing these species lie in their reproductive structures (Table 2). Although
this material could be assigned to E. fertilis taking into account the Ar-
gentinian provenance of the specimens, the lack of known fertile struc-
tures in the former leads us to compare our material to E. mougeotii. In
addition to the morphological resemblance of their vegetative parts,
both our material and E. mougeotii have fertile structures borne on a
non-articulated, leafless branch that divides into two smaller branches,
each carrying one strobilus (Kustatscher et al., 2007). The cones consist
of pentagonal sporangiophore heads arranged in whorls. Differences in
the size of the strobilus-bearing branches, number of whorls per strobi-
lus, and size and number of sporangiophore heads could be specific
traits. Even if we prefer to leave our specific assignation open until bet-
ter preserved material with its reproductive structures organically at-
tach appears, the presence of Equisetites at the Quebrada de los Fósiles
Formation is here also supported by the Equisetites-like strobili
described.
4.2. Division Lycophyta
Order Pleuromeiales Zimmermann, 1959
Family Pleuromeiaceae Potonié, 1901 (in Engler et Prantl)
Genus Pleuromeia Corda in Germar, 1852
Type-species: Pleuromeia sternbergii (Münster) Corda in Germar,
1852.
Pleuromeia sternbergii (Münster) Corda in Germar, 1852
Plate II, 1–10
Rock sample number. MHNSR Pb-CN 37, MHNSR Pb-CN 41, MHNSR
Pb-CN 43, MHNSR Pb-CN 49 a–b, MHNSR Pb-CN 53.
Stratigraphic levels. GzD, AMZ2.
Description. Unbranched stems at least 14 cm long and 6–10 cm
wide, with distinct leaf scars on the surface. Specimen MHNSR Pb-CN
Plate I. Sphenophyta from the Quebrada de los Fósiles Formation (Puesto Viejo Group, Mendoza, Argentina). 1. Neocalamites suberosus (MHNSR Pb-CN 19), black arrow points to expanded
node. 2–3. Neocalamites suberosus (MHNSR Pb-CN 15), black arrow (3) points to fused leaf bases, white arrow (3) points to ribs crossing the node in an alternate manner; “br” (2) for
branch. 4–5. Equisetites sp. (MHNSR Pb-CN 44), black arrows (5) points to lateral branches. 6. Equisetites sp. (MHNSR Pb-CN 39 a–b). 7–8. Equisetites sp. (MHNSR Pb-CN 40), black arrows
(8) point to the short leaves no longer adpressed to the stem. 9–12. Equisetites sp. (MHNSR Pb-CN 57 a–b), black arrow and white arrow (9) point to enlarged images in (10) and (12). Black
arrows in (10) point to smaller branches bearing one strobilus each, white arrow (10) points to (?)aborted strobilus. Scales. 1, 2, 5, 6, 7, and 9 = 10 mm; 3, 4, 8, 10, 11, and 12 = 5 mm.
7
49 a–b (Plate II, 1–6) is a smooth stem 14.5 cm long (incomplete) and
6.8 cm wide covered in spirally arranged leaf bases, phyllotaxis ~ 35°,
widely separated by 1.1 cm horizontally and 0.6 cm longitudinally
from each other. The leaf bases are clearly defined, ranging from oval–
circular, 11.6 mm by 9.2 mm at the top (Plate II, 1) to more transversely
lenticular, 23 mm by 9.5 mm, at the bottom (Plate I, 6). Leaf scars with
two well-defined triangular areas 3.5 mm by 3 mm (Plate II, 2–5) at
each side of a central area, where a circular leaf trace 1 mm in diameter
slightly protrudes from the surface. Another stem, MHNSR Pb-CN 53
(Plate II, 7–9), is 14 cm long (incomplete) and 9.6 mm wide, also cov-
ered in clearly defined leaf bases with the three-part leaf scars as previ-
ously described. In this specimen, however, there is evidence of a small,
circular ligule pit placed in the upper-middle part of the leaf scars (Plate
II, 8–9). Other examples of stems (MHNSR Pb-CN 37, 41, 43), although
more fragmentary and decorticated (Plate II, 10) bear the same size
and arrangement of leaf scars.
Remarks. Given that our fossils consist of compressed stems with
clearly defined leaf scars on the surface with no leaves attached, they
are comparable only to those Pleuromeia species that have had their
leaf bases well characterized. Among these, Pleuromeia jiachochengensis
Wang et Wang, 1982 differs with our specimens in that the former has
smooth stems covered by sparse, barely visible leaf scars. Pleuromeia
hataii Kon'no, 1973 has spirally arranged leaf scars with an elongated
rhombic outline, a central groove with two triangular areas at the
sides, the leaf trace slightly projecting from the surface, and a small cir-
cular ligule scar; all these features are shared with our specimens. How-
ever, P. hataii has distinct, large parichnos scars near the base of each
triangular area, which are lacking in our specimens. Parichnos scars
are also described in Pleuromeia rossica Neuburg, 1960, which differs
from the new material in the transversely oval shape of the areas at
the side of the leaf scar and their crowded arrangement on the stem.
Pleuromeia longicaulis (Burges) Retallack, 1975 shares the transversely
lenticular leaf bases, with two lateral triangular areas and a circular cen-
tral pit with a slightly projected leaf trace, the main difference being the
L:W ratio of the leaf scars and their closely spaced arrangement in the
Australian species. In addition, there are numerous creases on the
surface of the stem in P. longicaulis that are not evident in the fossils
from the Quebrada de los Fósiles Formation. Pleuromeia obrutschewii
Elias ex Krassilov and Zakharov, 1975 and Pleuromeia jokvazhica
Dobruskina, 1985 are very similar to Pleuromeia sternbergii in their veg-
etative features, differing only in size, and thus, considered as poten-
tially conspecific by some authors (Krassilov and Zakharov, 1975;
Retallack, 1997). The stems of Pleuromeia sternbergii are characterized
by having spirally arranged and widely spaced leaf bases, with distinct
rhombic leaf scars outlined by a peripheral rim, consisting of three
parts: a median area representing the leaf trace and two triangular
areas at each of the sides of the former. Evidence of a ligule placed on
the upper-centre of the leaf scar has been observed in P. sternbergii,
and although the presence of parichnos scars is still not fully
ascertained, some authors considered the triangular areas at the sides
of the leaf trace as parichnos (Boureau, 1967; Kon'no, 1973; Pigg,
1992). Of all described species, our material can be assigned to
P. sternbergii based on the overall similarities in size, shape, and arrange-
ment of the leaf bases on the stems, the ligule pit observed in some of
the specimens, and also, by the distinguishable triangular areas at
each side of the median area containing the circular leaf trace, that
could be considered to constitute the ‘parichnos’ of other authors. The
potential presence of this species in this unit was mentioned by
Coturel et al. (2016) as cf. P. sternbergii based on regularly preserved
material. Here, we adhere to their assignation and confirm the existence
8 B. Cariglino et al. / Review of Palaeobotany and Palynology 257 (2018) 1–18
Table 2
Comparison of the fertile structures in Equisetites.
MHNSR Pb-CN 57 Equisetites sp.
Strobili 2 attached to a leafless, unarticulated fertile
attachment shoot that divides into two smaller branches,
branch 5.2 mm long and 2.9 mm wide, smaller
branches 2.5 mm long and 1.3 mm wide
Strobili features Circular to slightly oval, 8 mm in diameter
Sporangiophore 3–4? whorls, pentagonal heads, ~9 mm in
heads diameter
of Pleuromeia sternbergii in the Southern Hemisphere for the first time,
pointing to the wide distribution of this species during the Triassic.
Order Isoetales Prantl, 1874
Family Isoetaceae Reichenbach, 1828
Genus Lepacyclotes (Emmons) Retallack, 1997
Type-species: Lepacyclotes circularis Emmons, 1856
Lepacyclotes sp.
Plate II, 13–16; Plate III, 1–5
Rock sample number. MHNSR Pb-CN 1 a–b, MHNSR Pb-CN 34 a–b,
MHNSR Pb-CN 38, MHNSR Pb-CN 45, MHNSR Pb-CN 46, MHNSR Pb-
CN 54.
Stratigraphic level. GzD, AMZ2.
Description. Specimen MHNSR Pb-CN 34 a–b (Plate II, 13) is a par-
tially preserved axis ~90 mm inferred diameter (semi-covered and in-
complete), with at least two cycles of densely packed helically
arranged sporophylls. Largest sporophylls are 42 mm long and 8 mm
wide, with a distinct straight midline 1.8 mm thick. Specimen MHNSR
Pb-CN 1 a–b (Plate II, 14) is a compact stem preserved as a compression,
7.1 cm long (incomplete) and 3.2 cm wide. Densely inserted sporophylls
cover the apex; they are 12 mm long, 1.8–2 mm wide (Plate II, 14–15),
with straight margins and distally pointed. The stem is covered in disor-
dered, closely spaced, eye-shaped leaf scars, with a central vascular
trace (Plate II, 16). In addition, other rock slabs (MHNSR Pb-CN 38, 45,
46; Plate III, 1–5) host numerous isolated, tongue-shaped sporophylls,
38–47 mm long and 9–18 mm wide. The distal portion is wide, whereas
the proximal portion is narrow with a slightly expanded base. In one
well-preserved example, the sporangia boundary is visible (Plate III,
5). The apex is mucronate, with a blunt tip. The sporophylls are sym-
metrically divided by a straight distinct longitudinal midline, 1–2 mm
wide. A lateral wing ~ 4 mm wide was observed in only one specimen
(Plate III, 5).
Remarks. The two-cycle arrangement of sporophylls in MHNSR
Pb-CN 34 a–b shows the typical ‘pineapple slice-like structures’ de-
scribed as Lepacyclotes zeilleri (see Kustatscher et al. 2015, fig. 2e–g).
Sporophylls are tongue-shaped with a mucronate apex and a strong,
straight midline that divides them into two symmetrical halves in
both the articulated and isolated materials. No ligule or ligule scars
are clearly observable in our material; only a doubtful example of a
ligule 3 mm long and 1.3 mm wide is present in one of the many ex-
amples (Plate III, 3). The vertical midline symmetrically dividing a
central sporangium is mentioned by some authors as a diagnostic
feature for Lepacyclotes zeilleri (Moisan and Voigt, 2013). Even
though the borders of the sporangium are not well-delimited, the
distinct midline observed in our material would suggest its potential
assignment to L. zeilleri. On the other hand, the sporophylls of nu-
merous similar taxa (i.e., Tomiostrobus, Skilliostrobus, Cylostrobus)
also show sporangia with a longitudinal midline (see Grauvogel-
Stamm and Lugardon, 2001 for further details), indicating this char-
acter alone is not enough to define L. zeilleri. The absence of a ligule
Equisetites mougeotti Equisetites Equisetites aequecaliginosus Equisetites
arenaceous fertilis
2 attached to a leafless unarticulated 3 attached to a 1 attached to a short Unknown
branch that divides into two, branch 80 articulated peduncle bearing
slender articulated
mm long and 6.5 mm wide, smaller fused leaf-like lobes forming
leafy fertile branch
branches 22 mm long and 3.5 mm wide a sheath in the uppermost
node
Obovate, 40 mm long and 25–28 mm Obovate, 21–35 Cylindrical, 50–60 mm long Unknown
wide mm long and 17–22 and 25 mm wide, rounded
mm wide apex
5–6 whorls, pentagonal or hexagonal 7–9 whorls, 10–12 Hexagonal heads, 4–6 mm in Unknown
heads, 2.5–3.5 mm in diameter pentagonal or diameter
hexagonal heads,
2.5–3.5 mm in
diameter
in our specimens seems to be a common preservational issue, as no-
ticed in other sporophylls from Germany and China
(Grauvogel-Stamm and Lugardon, 2001; Yu et al., 2010; Kustatscher
et al., 2015). Several faint impressions of rounded corpuscles ~ 270
μm in diameter are evident along the midline of some of the sporo-
phylls and even spread on their surface (Plate III, 5), and could po-
tentially be spore impressions, but the preservational nature of
these fossils did not allow the recovery of in-situ spores.
The compact stem (=‘corm’) with densely inserted sporophylls cov-
ering most of the apex in MHNSR Pb-CN 1 a–b matches the diagnosis of
Lepacyclotes provided by Retallack (1997). In particular, the Italian
Lepacyclotes bechstaedtii Kustatscher et al., 2010 specimen (Kustatscher
et al., 2010, plate 6, fig. 1) has noticeable similarities in overall morphol-
ogy with our material. However, on closer inspection, the sporophylls in
MHNSR Pb-CN 1 a–b do not have the ‘spateolate’ shape of L. bechstaedtii,
being straight and ending in an acute tip, and most importantly, there are
no sterile leaves in our specimen, a key character used to identify
L. bechstaedtii. Unfortunately, the fragmentary nature of our specimen
does not provide details of the basalmost part of the stem, thus we can-
not ascertain the presence of a quadrilobate corm, preventing further
comparisons with the Italian species. Comparisons with Lepacyclotes
circularis Emmons, 1856 is more problematic because this species is de-
scribed from specimens preserved in dorsiventral view, with a detailed
count of the sporophylls in each whorl (‘rosette’), whereas our specimen
is preserved in lateral view; however, they differ in that L. circularis has
weakly serrated sporophylls with a blunt tip. Lepacyclotes convexus
(Brik) Retallack, 1997 on the other hand, has sub-rhombic sporophylls
with an obtusely pointed tooth, and Lepacyclotes ermayinensis (Wang)
Retallack, 1997 sporophylls are mucronate and less elongate triangular,
thus these species also differ from the Puesto Viejo material.
Many authors have already noted the close similarities among the
sporophylls of Triassic lycophytes (i.e., Sadovnikov, 1982; Dobruskina,
1985; Retallack, 1997; Grauvogel-Stamm and Lugardon, 2001; Yu
et al., 2010). The taxonomic position of the complex Lepacyclotes (=
Annalepis)–Tomiostrobus (=Skilliostrobus = Austrostrobus)–Cylostrobus
is still debated, with more of these genera being synonymized in time
owing to new discoveries that allow a better interpretation of their
structure (e.g., Lepacyclotes = Annalepis, Retallack, 1997; Kustatscher
et al., 2015, Tomiostrobus = Skilliostrobus = Austrostrobus, Sadovnikov,
1982; Retallack, 1975, 1997; Lepacyclotes = Tomiostrobus, Meng,
1998). It is clear that a full review of these taxa is still needed; hopefully,
more findings from new localities, such as the one presented here, will
help shed light on the real affinities of this group of lycophytes.
Indeterminate lycophyte stems
Plate II, 11–12
Rock sample number. MHNSR Pb-CN 33 a–b.
Stratigraphic level. AMZ2.
Description. Specimen MHNSR Pb-CN 33 a–b (Plate II, 11–12) is a
regularly preserved compressed stem of a lycophyte, at least 8 cm
 B. Cariglino et al. / Review of Palaeobotany and Palynology 257 (2018) 1–18 9

Plate II. Lycophyta from the Quebrada de los Fósiles Formation (Puesto Viejo Group, Mendoza, Argentina). 1–6. Pleuromeia sternbergii (MHNSR Pb-CN 49 a–b). 7–9. Pleuromeia sternbergii
(MHNSR Pb-CN 53), black arrows in (8) and (9) point to ligule pits. 10. Pleuromeia sternbergii (MHNSR Pb-CN 41). 11–12. Indeterminate lycophyte stem (MHNSR Pb-CN 33 a–b). 13.
Lepacyclotes sp. (MHNSR Pb-CN 34 a–b). 14–16. Lepacyclotes sp. (MHNSR Pb-CN 1 a–b). Scales. 1, 6, 7, 10, 12, 13, 14 = 10 mm; 2, 3, 4, 8, 15 = 5 mm; 5, 9, 11, 16 = 2 mm.

long and 3.3 cm wide, with a leafy apex. In the lower part of the stem
there are elongate oval leaf scars, 5–5.5 mm long and 1.9–2.5 mm
wide, arranged in a close spiral. Centrally located towards the upper
third of the scar there is a circular mark, 0.7 mm in diameter (Plate II,
11), here inferred to be the leaf trace. The upper part of the preserved
stem appears to be covered in microphylls (Plate II, 12), about 24 mm
long and 1 mm wide, with a distinct central midvein.
Remarks. The absence of distinctive features in this specimen pre-
vents confident assignation. However, the closed spiral arrangement
of the leaf bases, in addition to their elongate oval shape, differs from
10 B. Cariglino et al. / Review of Palaeobotany and Palynology 257 (2018) 1–18

Plate III. Lycophyta from the Quebrada de los Fósiles Formation (Puesto Viejo Group, Mendoza, Argentina). 1–5. Lepacyclotes sp. (MHNSR Pb-CN 38, 45, 46, 54), various disaggregated
tongue-shape sporophylls. Note the sporangia (large black arrow), the lateral wing (white arrow) and potential megaspore impressions (small black arrows) in (5); the white arrows
in (2) and (3) points to the strong midline dividing the sporophyll into two halves; the black arrow in (3) points the only evidence of ligule observed in this material. 6–8. Isolated mega-
sporophyll bearing clusters of megaspores (MHNSR Pb-CN 57 a–b). Scales. 1, 2, 3 = 10 mm; 4, 5, 7, 8 = 5 mm; 6 = 2 mm.

the typical pleuromeian-type of stems, and has a more lepidodendroid
affinity.
Indeterminate lycophyte reproductive structure
Plate III, 6–8
Rock sample number. MHNSR Pb-CN 57 a–b.
Stratigraphic level. AMZ1.
Description. Part and counterpart of a single, isolated sporophyll
bearing clusters of megaspores. Sporophyll is obovate with a wide trian-
gular apex, 8.2 mm long and 5 mm at its maximum width, length to
breadth ratio less than 2:1. Sporophyll bears numerous spherical mega-
spores, ~300 μm in diameter, clustered in triads and tetrads (Plate III,
6–8).
Remarks. Owing to the detached nature of the megasporophyll, and
since no further clarifying information could be obtained from the SEM
images of the megaspores, we cannot confidently determine its kinship.
However, its affinity probably lie with the lycophytes, being very similar
in size and shape to the elongate cuneate Cylostrobus megasporangia
bearing numerous trilete megaspores described by Helby and Martin
(1965, p. 391, fig. 4). Detached sporophylls described by Kon'no
(1973) and assigned to the fertile structures of Pleuromeia hataii Kon'no,
a Triassic lycophyte from Japan, are similar in the slightly elongated ob-
ovate shape, although considerably smaller than our specimen, being
only 2.5 mm long and 2 mm wide, and bearing megaspores 400 μm in
diameter. Last, isolated obovate sporophylls with a rounded to obtuse
apex assigned to Pleuromeia rossica Neuburg have been also described
by Naugolnykh (2013); however these sporophylls bear a small sinus
in the medial part, a feature not observed in our material.
4.3. Division Pteridospermophyta
Order indet.
Family indet.
Genus Ptilozamites Nathorst, 1878
Type-species: Ptilozamites heeri Nathorst, 1878
Ptilozamites longifolia sp. nov.
Plate IV, 1–12; Plate V, 1–12
Holotype. MHNSR Pb-CN 18 (Plate IV, 1; Plate V, 1–8)
Additional material. MHNSR Pb-CN 10a–b, MHNSR Pb-CN 12,
MHNSR Pb-CN 14, MHNSR Pb-CN 16, MHNSR Pb-CN 25, MHNSR Pb-
CN 27.
Repository. Palaeobotany Collection, Museo de Historia Natural de
San Rafael, San Rafael, Mendoza (MHNSR Pb-CN).
Type locality. Quebrada de los Fósiles Creek, type section (34°53′S,
68°23′W), Puesto Viejo Group, Mendoza, Argentina.
Stratigraphy. Puesto Viejo Group, Quebrada de los Fósiles Formation,
stratigraphic level AMZ1 (Fig. 2).
Age of the Formation. Middle to Late Triassic (late Anisian–early
Carnian).
Etymology. “longifolia” alluding to its long pinnae.
Species diagnosis. Unipinnate leaves with an often forked rachis.
Rachis stout and longitudinally striate. Pinnae sub-oppositely and later-
ally attached by their whole bases to the axis, both before and after the
dichotomy, at a 45° or higher degree angle, slightly separated from each
other, in some cases touching at their bases, but never imbricating.
Pinnae long, linear to sub-rectangular, length to width ratio equal or
 B. Cariglino et al. / Review of Palaeobotany and Palynology 257 (2018) 1–18 11

greater than 3:1, apex rounded to obtuse. Numerous distinct veins
inserted directly on the rachis, in some cases decurrent at the base, run-
ning almost parallel to the margins, reaching the apex with very few di-
chotomies. Cuticle thick, with irregular to isodiametric epidermis cells,
randomly arranged trichomes bases and monocyclic stomata. Stomata
constituted by two sunken guard cells surrounded by 4–6 (usually
5) subsidiary cells fused to form a ring-like thickening.
Description. The most complete specimen (MHNSR Pb-CN 18, holo-
type; Plate IV, 1 and Plate V, 1–8) shows the bifurcated axis but other-
wise pinnate leaf. The rachis is striated, 1 cm thick proximal to the
dichotomy and 0.8 cm distally. Pinnae are attached to the rachis both
proximally and distally to the dichotomy, inserted by their entire base.
A complete pinna is 9.5 cm long and 2.4 cm wide, sub-rectangular in
shape, with a rounded obtuse apex. Venation is conspicuous, with nu-
merous veins running almost parallel to the margins and reaching the
apex in a slightly radiating manner. Cuticle was obtained from two
samples; the light microscopy images (Plate V, 9–12) illustrate cuticle
recovered from an isolated rock sample bearing leaves with
macromorphological characters attributable to this species (destroyed
during the preparation process), whereas the SEM images (Plate V,
1–8) were obtained from processing some of the cuticles preserved in
specimen MHNSR Pb-CN 18. The epidermal cells are irregular to

Plate IV. Ptilozamites longifolia sp. nov. from the Quebrada de los Fósiles Formation (Puesto Viejo Group, Mendoza, Argentina). 1. Ptilozamites longifolia sp. nov. holotype (MHNSR Pb-CN
18), black arrow points to bifurcation of the axis. 2. Ptilozamites longifolia sp. nov. (MHNSR Pb-CN 14), black arrow points to bifurcation of the axis. 3. Ptilozamites longifolia sp. nov. (MHNSR
Pb-CN 10), note the almost parallel venation of the leaves. 4. Ptilozamites longifolia sp. nov. (MHNSR Pb-CN 12). 5. Ptilozamites longifolia sp. nov. (MHNSR Pb-CN 27). 6. Ptilozamites longifolia
sp. nov. (MHNSR Pb-CN 16). 7. Ptilozamites longifolia sp. nov. (MHNSR Pb-CN 25). 8–11. Ptilozamites longifolia sp. nov. (MHNSR Pb-CN 14) detail of axis bifurcation (8) and impression of
trichomes on the leaves (black arrows, 9–10). 12. Ptilozamites longifolia sp. nov. (MHNSR Pb-CN 12), detail of leaf base and attachment to stem. Scales. 1–7 = 10 mm; 9–12 = 5 mm.
12 B. Cariglino et al. / Review of Palaeobotany and Palynology 257 (2018) 1–18

Plate V. Ptilozamites longifolia sp. nov. from the Quebrada de los Fósiles Formation (Puesto Viejo Group, Mendoza, Argentina). 1–8. Cuticle under SEM (MHNSR Pb-CN 18), observe the
sunken stomata with thick ring-like structure constituted by the fused walls of the subsidiary cells, a potential trichome (white arrow in 7), and the elongate cells indicative of venation
(8). 9–12. Cuticle obtained from isolated foliage (sample destroyed) under light microscopy. Observe the thick ring-like structure and sunken stomata (9–10), a trichome base (11) and
polygonal shape of the cells (12). Scales. 1, 2, 4, 5, 9–12 = 20 μm; 3, 6, 8 = 50 μm; 7 = 10 μm.

polygonal, 19–32 μm in diameter, with straight anticlinal walls 4 μm
thick. The stomata are constituted by two guard cells with 4–6, most
commonly 5, radially arranged subsidiary cells. The guard cells are reni-
form (Plate V, 4–5), ~24 μm long and 12 μm wide. The subsidiary cells
are also irregular to polygonal, with their walls fused in a thick ring
that partially covers the guard cells and the stomatal pit (Plate V, 2–3,
9–10). The similarities in size and morphology featured in the cuticles
from two samples support the idea of a single, new species of
Ptilozamites at this locality. A forked axis and pinnae attached before
and after the rachis are also represented in specimen MHNSR Pb-CN
14 (Plate IV, 2), although the apices in these pinnae are more
rounded-acute. The other specimens attributed to this new species
(MHNSR Pb-CN 10, 12, 16, 25, 27; Plate IV, 3–12) are fragmentary leaves
with pinnae attached to thick, striated rachises, with an ~3:1 length to
width ratio and venation almost parallel to the margins.
Remarks. Originally erected by Nathorst (1878), the defining charac-
ters of Ptilozamites have long been discussed, with several emendations
of the genus proposed in time. To date, the genus encompasses
unipinnate, petiolate, commonly forked fronds, with pinnae of variable
morphology, from square, falcate, triangular to rounded or rhombic, lat-
erally attached by their entire base to a stout rachis. Recently, the diag-
nosis was emended to include also bipinnate leaves, based on material
from the Triassic of Greenland (Popa and McElwain, 2009). Other au-
thors, however, suggested maintaining the original unipinnate charac-
ter for Ptilozamites and the bipinnate one for Ctenozamites, using the
lateral vs. the adaxial attachment of the pinnae to the axis, respectively,
as further evidence for their generic separation (Zhou, 1981;
Kustatscher and van Konijnenburg-van Cittert, 2013). When the cuticle
is preserved, the presence of a strong ring-like thickening surrounding
the stomata is considered as one of the defining characters to differenti-
ate Ptilozamites from other morphologically similar taxa (see the thor-
ough review of Kustatscher and van Konijnenburg-van Cittert, 2007
for further details). This feature was firstly recognized by Antevs
(1914) and widely acknowledged by subsequent authors (Harris,
1926; Florin, 1933; Popa and McElwain, 2009; Kustatscher and van
Konijnenburg-van Cittert, 2007, 2013).
Our material has superficial similarities to some cycadophyte and
seed fern leaf remains, particularly with Pseudoctenis, Komlopteris and
Dicroidium species. Pseudoctenis wardii (Fontaine) Artabe, 1985 was de-
scribed for the Triassic Los Menucos Formation (Río Negro Province,
Argentina). The leaf is very similar in its elongated pinnae inserted to
the rachis by their entire, slightly decurrent base. Pinnae margins are
parallel, with a rounded to obtuse apex. The venation is sub-parallel,
with veins bifurcating sometimes more than once. However, this spe-
cies differs from our Ptilozamites in the lack of a forked rachis. Unfortu-
nately, no cuticle was recovered from P. wardii, precluding comparisons
at the micromorphological level. Komlopteris nordenskioeldii Barbacka,
1994, on the other hand, has a thick cuticle with stomata consisting of
5–7 subsidiary cells forming a ring around the pit, in a similar manner
as Ptilozamites. The main differences between these taxa are at the
macromorphological level; in Komlopteris, pinnae have slightly con-
stricted bases, unparallel margins (sometimes undulated), and a dis-
tinct midrib (alethopteroid venation). Moreover, Komlopteris does not
have a forked rachis. Finally, the material from the Quebrada de los
Fósiles is somewhat macromorphologically similar to the recently de-
scribed Dicroidium bandelii Hamad et al., 2017 from the Permian of
Jordan in its forked rachis, the once pinnate architecture, and the elon-
gate pinnae with near parallel venation. Besides some disparity in the
overall shape of the pinnae, the main difference to separate these taxa
is cuticular. Whereas in Ptilozamites the stomata are sunken, in
D. bandelii the stomata are superficial, with 2–4 subsidiary cells that
do not form a ring-like structure.
Although the overall shape, venation of the pinnae and in some
cases, the cuticular features in our fossils resemble to those of other
gymnosperms, the combination of both macro- and micromorphologi-
cal characters of the Quebrada de los Fósiles Formation specimens
 B. Cariglino et al. / Review of Palaeobotany and Palynology 257 (2018) 1–18 13

asserts their correct assignation to Ptilozamites, making this the first ev-
idence of the genus not only for Argentina, but also for the entire South-
ern Hemisphere.
Of the five known species (i.e., Ptilozamites heeri Nathorst, 1878,
Ptilozamites nilssonii Nathorst, 1871, Ptilozamites sandbergeri (Schenk)
Kustatscher and van Konijnenburg-van Cittert, 2007, Ptilozamites blasii
(Brauns) Nathorst, 1879, Ptilozamites tenuis Oishi, 1932; Table 3), our
fossils share most similarities with P. nilssonii, such as the presence of
the bifurcate axis (also present in P. tenuis) and the numerous, distinct
veins running almost parallel to the margins. However, the main differ-
ences are that our fossils have considerably larger, more lanceolate pin-
nae, and that P. nilssonii is the only species in which the typical ring-like
structure of the subsidiary cells is not well developed. Furthermore, the
pinnae in P. nilssonii are falcate and imbricate on the axis, in contrast to
the non-imbricate, long, sub-rectangular pinnae in our specimens. One
exception to the L:W = 3:1 of the pinnae from our material is
MHNSR Pb-CN 27, which has at least three similar leaves with no
obvious organic connection but in close association on the same
rock slab, where pinnae are slightly shorter (~ L:W = 2:1). Al-
though our specimen is comparatively similar to P. blasii (see
Kustatscher and van Konijnenburg-van Cittert, 2007, fig. 2a–d), in
our material the pinnae are slightly separated, whereas in the
European specimens the pinnae bases touch. The significantly
larger size, the sub-rectangular shape, and the L:W = 3:1 (or
greater) ratio of the pinnae, the slightly spaced (never imbricate),
sub-opposite arrangement on the axis, and the presence of epider-
mal cells with straight anticlinal walls and stomata with sunken
guard cells surrounded by a thick ring-like structure constituted
by the fused walls of five subsidiary cells, justify the assignation
of our fossils to a new species of Ptilozamites, here proposed as
Ptilozamites longifolia sp. nov.
The lack of reproductive organs attached to its foliage still prevents
establishing with certainty a higher-level taxonomy for Ptilozamites. In
an attempt to resolve the phylogenetic relationships of various seed-
bearing plants, including Ptilozamites, Vajda et al. (2017) applied Fourier
transformed infrared (FTIR) spectroscopy and hierarchical cluster anal-
ysis to obtain chemical signatures from fossil cuticles. Surprisingly, their
results suggested a closer phylogenetic relationship among Ptilozamites
nilssonii Nathorst, Nilssonia brevis Brongniart, and the bennettitaleans
Pterophyllum schenkii Zeiller and Otozamites boolensis Douglas, rather
than to other pteridosperms or to the cycadales.
5. Discussion
5.1. Biostratigraphic implications
The macroflora of the Quebrada de los Fósiles Formation was previ-
ously considered to represent the only South American example of a ‘re-
covery flora’ that has been widely recognized to characterize the
beginning of the Triassic Period in the Northern Hemisphere (Wang,
1996; Looy et al., 1999; Grauvogel-Stamm and Ash, 2005). This idea
stemmed from the impoverished character of the plant assemblage
comprising lycophytes and sphenophytes (‘Pure Pleuromeia assem-
blage’, Morel and Artabe, 1994; Morel et al., 2003; Spalletti et al.,
2003; Artabe et al., 2007a). Furthermore, other palaeontological content
recovered from the Puesto Viejo Group, including fossil tetrapods, in-
vertebrates and palynomorphs partially supported an Early Triassic or
even a latest Permian age for the base of the succession (Ottone and
García, 1991; Zavattieri and Papú, 1993; Zavattieri and Batten, 1996;
Bonaparte, 2002; Zavattieri et al., 2003; Stipanicic et al., 2007; Gallego
et al., 2009). However, based on radiometric dating (Valencio et al.,
1975; Domeier et al., 2011; Ottone et al., 2014; Monti in Sato et al.,
2015), the Puesto Viejo Group has been recently constrained to Middle
to Late Triassic age, challenging previous concepts solely based on the
fossil content.
The recovery of new well-preserved and readily identifiable vegeta-
tive and fertile remains of lycophytes and sphenophytes, and the dis-
covery of abundant pteridosperm fossils, radically change the long-
held view of a ‘pure’ Pleuromeia flora of Early Triassic age and point to
a younger plant assemblage, more in line with these radiometric dates.
Lycophytes and sphenophytes were commonly associated during
the Triassic in both hemispheres. As rapidly growing species with high
adaptability to different environmental conditions, they were common
elements with wide distributions, in many cases constituting the dom-
inant plants in their communities (Kon'no, 1973; Retallack, 1975, 1997;
Looy et al., 1999; Grauvogel-Stamm and Ash, 2005). With a prominent
representation during the Early Triassic, Pleuromeia rapidly declined
until its complete disappearance by the end of this period worldwide
(Pigg, 1992; Grauvogel-Stamm and Ash, 2005; Naugolnykh, 2013).
Lepacyclotes (= Annalepis), a close ally of Pleuromeia also found in
these assemblages, has been reported from various localities in
Europe, Asia, Australia and Argentina. The oldest record comes from
China, where specimens were recovered from Induan deposits (Meng,

Table 3
Comparison of the main features in Ptilozamites species.

Ptilozamites longifolia sp. nov. Ptilozamites nilssonii Ptilozamites Ptilozamites heeri Ptilozamites Ptilozamites blasii
tenuis sandbergeri

Forked Yes Yes Yes Possibly No No

axis
Pinnae Slightly spaced, not imbricated, sub-opposite Imbricated Slightly – Alternate Bases touch
insertion spaced or
closely
inserted,
alternate
Apex Rounded to obtuse Obtuse Rounded to Truncate or slightly Rounded Rounded
obtuse apically inclined
Venation Conspicuous, numerous veins almost parallel Numerous, distinct, 4–7 veins, Dichotomizing, veins Very thick, parallel Parallel to slightly
to margins, very few dichotomies almost parallel to parallel, delicate, parallel to radiating, decurrent at the
margins simple or slightly radiating base
forked
Pinnae Long, large, L:W = 3:1 in general, sub Falcate, length N Small, Crescentic to Short, elongate to Rhombic to slightly falcate,
rectangular, length N N width width parallel rectangular, length ≥ equidimensional, length N width
sided, width subrectangular to
length ≥ subquadrate, length ≥
width width
Cuticle Stomata with sunken guard cells surrounded Stomata with 6 – Stomata with sunken – Stomata with sunken guard
by a thick ring-like structure constituted by (5–7) subsidiary guard cells and cells and well-developed
the fused walls of 5 subsidiary cells, trichomes cells, ring-like well-developed ring-like structure, cell's
present, cell's anticlinal walls straight structure not ring-like structure, anticlinal walls undulating
well-developed papillae present
14 B. Cariglino et al. / Review of Palaeobotany and Palynology 257 (2018) 1–18

1998; Yu et al., 2010); however, their climax occurred during the Middle
Triassic (Dobruskina, 1995; Retallack, 1997; Meng, 1998; Kustatscher
et al., 2010, 2015; Moisan and Voigt, 2013) (Fig. 3).
Regarding the sphenophytes, Neocalamites had a worldwide distri-
bution ranging from the Permian to Cretaceous. However, their acme
was during the Triassic (Boureau, 1964; Bomfleur et al., 2013). Such is
the case with Neocalamites suberosus, known only from Southern Hemi-
sphere localities (Artabe and Zamuner, 1991; Brea and Artabe, 1999;
Holmes, 2000; Bomfleur et al., 2013). Equisetites, on the other hand,
has been reported from the Carboniferous to the Jurassic. Similar to
Neocalamites, the majority of its species are known from Triassic
deposits (Taylor et al., 2009). The Equisetites species considered for
comparisons in this work were all recovered from Upper Triassic strata,
with the exception of E. mougeotii, restricted to the Anisian (Frenguelli,
1944; Kelber and van Konijnenburg-van Cittert, 1998; Weber, 2005;
Kustatscher et al., 2007) (Fig. 3).
Among the novel additions to the flora is Ptilozamites, which had an
exclusive Northern Hemisphere distribution until this study. Species of
Ptilozamites have been described from lower Middle to predominantly
Upper Triassic assemblages in Europe and Asia (Zhou, 1981; Popa and
McElwain, 2009; Kustatscher and van Konijnenburg-van Cittert, 2007,
2013) (Fig. 3).
Although not useful as biostratigraphic indices, the new macrofloral
evidence (Fig. 3) is in line with the radiometric ages obtained for the
Puesto Viejo Group, and reinforces a latest Anisian–earliest Carnian
age for the lower unit (Quebrada de los Fósiles Formation). The pres-
ence of Neocalamites and Equisetites does not contradict this age, even
though these genera are not constrained to the Middle Triassic per se.
Lepacyclotes, on the other hand, has a strong presence in the Middle Tri-
assic, whereas Pleuromeia was more common during the Early Triassic.
Lastly, Ptilozamites longifolia sp. nov. is one of the earliest evidences for
the genus, concordantly to Ptilozamites cf. P. sandbergeri from the
Upper Anisian of Italy (Kustatscher and van Konijnenburg-van Cittert,
2007).

5.2. Palaeogeographic and palaeoenvironmental implications

The merger of landmasses into a single supercontinent (Pangaea)

and the absence of ice caps at the poles caused the reduction from
four floral realms in the late Paleozoic to two in the Mesozoic (Wing
and Sues, 1992). The ameliorated climatic conditions of the Triassic de-
creased the endemism and allowed the migration and expansion of the
geographic ranges of many species towards higher latitudes, some
reaching a cosmopolitan distribution (Dobruskina, 1987; Meyen,
1987; Artabe et al., 2001; Spalletti et al., 2003; Kustatscher et al., 2017).
When compared to other Middle and Late Triassic assemblages in
Argentina, the flora from the Quebrada de los Fósiles Formation is
unique in that it includes Ptilozamites (until now a genus exclusive
to the Northern Hemisphere), and abundant pleuromeids and sphe-
nopsids. Other typical elements of the Dicroidium flora (such as
corystosperms and peltasperms) common to the various Triassic
floras from Argentina and other Gondwanan regions are absent
(Anderson and Anderson, 1985, 1989, 1993, 2003; Pole and Raine,
1994; Artabe et al., 2003, 2007a, 2007b; Holmes, 2000, 2003;
Holmes and Anderson, 2005a, 2005b, 2007, 2013; Bomfleur and
Kerp, 2010; Chatterjee et al., 2013 among others). Nonetheless, sim-
ilar coeval plant assemblages have been described from some North-
ern Hemisphere localities (Fig. 4). Thus, an adequate explanation for
this unexpected disjunctive distribution of the Quebrada de los
Fósiles plant assemblage could lie in the local environmental condi-
tions in which the flora developed.
In Argentina, Neocalamites and Equisetites are represented by several
species (Zamuner et al., 2001), but the record of Pleuromeia is rather
poor, with a few fragmentary specimens assigned only at the genus
level (Labudia et al., 1992). These plants grew near fluvial systems,
and as ecological opportunists, they were fast and extensive colonizers
Fig. 3. Biostratigraphic distribution of main taxa (see text for further details). References:
(1, 5) Boureau, 1964; (2) Bomfleur et al., 2013; (3, 4) Zamuner et al., 2001, Artabe et al.,
2007b; (6) Kustatscher et al., 2007; (7) Kelber and van Konijnenburg-van Cittert, 1998;
(8) Weber, 2005; (9) Frenguelli, 1944; (10) Pigg, 1992; (11) Grauvogel-Stamm and
Lugardon, 2001; (12) Wang and Wang, 1982; (13) Kon'no, 1973; (14) Naugolnykh,
2013; (15) Retallack, 1975; (16) Krassilov and Zakharov, 1975; (17) Yu et al., 2010; (18)
Retallack, 1997; (19) Kustatscher et al., 2010; (20) Dobruskina, 1982; (21) Wang, 1991;
(22) Moisan and Voigt, 2013; (23) Kustatscher and van Konijnenburg-van Cittert, 2007.
of moist environments, with tolerance to various environmental
stresses, such as salinity or dryness (i.e., Retallack, 1975; Wang and
Wang, 1982; Kustatscher et al., 2010; Naugolnykh, 2009, 2013). Their
association in the Quebrada de los Fósiles Formation is indicative of veg-
etation growing close to shallow water bodies, coincident with the evi-
dence from the lithofacies (Fig. 2, Table 1) and the presence of
conchostracans and thin stomatolitic-like algal beds in the succession.
The record of Lepacyclotes in Triassic floras was probably more geo-
graphically extensive than previously acknowledged (Fig. 4). The recent
finding of more complete specimens has allowed the re-assignment of
various taxa to the same species (i.e., Kustatscher et al., 2015). We ex-
pect that with further discoveries, a better understanding of these
plants will prove their kinship and thus extend their record to other Tri-
assic assemblages worldwide.
The new Ptilozamites species represents the first evidence of the
genus for the entire Southern Hemisphere. As is the case with our spec-
imens, the European and Asian Ptilozamites species are characterized by
thick cuticles with sunken stomata and trichomes, features interpreted
to represent adaptations to reduce the loss of water in stressful or xero-
phytic environments. Some authors suggested it grew in coastal
 B. Cariglino et al. / Review of Palaeobotany and Palynology 257 (2018) 1–18
environments as a shrubby plant, next to halophytic lycophytes, such as
Lepacyclotes (= Annalepis) (Kustatscher and van Konijnenburg-van
Cittert, 2005; Kustatscher et al., 2010).
As episodes of intense magmatism driven by extensional rifting
were occurring in southwestern Gondwana during the Middle Triassic
(McKay et al., 2015), a feasible explanation for the presence of this un-
usual plant assemblage from the Quebrada de los Fósiles Formation
could relate to the adverse microenvironmental conditions created by
the volcanic episodes that occurred during the deposition of this unit.
The tuffaceous lithofacies (FF2, Table 1) linked to shallow water bodies
in which our plant assemblage was preserved suggests local events con-
trolled by tectonic and volcanic activity (Monti and Franzese, 2016) oc-
curred at the time that this flora was developing. The protective, thick
cuticle of Ptilozamites, and the presence of Lepacyclotes and Pleuromeia
(opportunistic taxa) in the same tuffaceous lithofacies, add to the idea
of altered environmental conditions at the site (Pfefferkorn, 1999;
Grauvogel-Stamm and Ash, 2005).
In this sense, the seasonally sub-humid climatic conditions inferred
from the clastic deposits of the Quebrada de los Fósiles Formation
(Spalletti, 1994; Kokogián et al., 2001; Tassi et al., 2013) are somewhat
hindered by the volcanism affecting the area. Even if not happening cat-
astrophically, the mild but continuous (or occurring episodically in
short lapses of time) release of volcanic gases and ash can produce
changes in the atmosphere, water bodies and soils that impede the nor-
mal development of vegetation (Sadler and Grattan, 1999; Robock,
2000, 2004). In this scenario, only species with a fast growth rate
(i.e., pioneer taxa) or with certain features (i.e., thick cuticles, fleshy
stems) seem to be able to establish at the site (Fig. 5).
The autochthonous nature of the plant assemblage at the Quebrada de
los Fósiles Formation is indicated by the preponderance of delicately pre-
served leafy shoots with intact leaf whorls belonging to sphenophyte's
and lycophyte's remains, strobili and sporophylls in close association,
and the large size and branching of the seed fern foliage. Most impor-
tantly, fossils are overlapping in single bedding planes, suggesting that
there was in situ accumulation of the plant assemblage.
The lack of pteridosperm foliage other than that of Ptilozamites is
nonetheless remarkable. Their absence as a consequence of taphonomic
influences is unlikely; if the thick cuticle in Ptilozamites provided it a
higher chance of being preserved, the same should have applied, for ex-
ample, for peltaspermalean foliage, which also had a thick cuticle
(Townrow, 1960; Barbacka, 1991). Most likely, Ptilozamites was part of
the only vegetation growing at the site, in the floodplains of the fluvial
Fig. 4. Generalized palaeogeographic distribution of plant assemblages comparable with the flora at the Quebrada de los Fósiles Formation. 1. San Rafael Block. 2. North America. 3. Europe.
4. Russia. 5. Kyrgystan. 6. China. 7. Japan + Korea. 8. Iran. 9. Australia. References: those in text and see also Berry, 1912; Kimura et al., 1982; Boersma and van Konijnenburg-van Cittert,
1991; Holmes, 2001; Shchervakov, 2008; Pattemore, 2016.
15
system, alongside Pleuromeia, Lepacyclotes, Equisetites and Neocalamites.
Collectively, all the evidence shows that the plant assemblage recovered
from the Quebrada de los Fósiles Formation represents an autochtho-
nous riverine flora, growing intermittently depending on the volcanic
activity in the area (Fig. 5).
6. Conclusions
The floral assemblage from the Quebrada de los Fósiles Formation of
the Puesto Viejo Group consists of impression-compression fossils of
fertile and sterile remains of lycophytes, sphenophytes, and seed ferns.
Among the novel additions to the diversity of this flora is Ptilozamites,
a typical Northern Hemisphere genus, here recorded for the first time
in Gondwana. The association of Ptilozamites, Pleuromeia, Lepacyclotes,
Equisetites and Neocalamites is unique when compared to other South-
ern Hemisphere Triassic assemblages, where Dicroidium flora elements
are dominant. However, this association of taxa is common in many Tri-
assic localities in Laurasia.
A potential explanation for this disjunctive distribution relates to the
environmental conditions under in which these plants developed. The
lithofacies associations suggest the parent vegetation grew and was de-
posited in the floodplains of a high-sinuosity fluvial system. The preser-
vation of the macroflora in fine-grained tuffaceous sediments points to
volcanic events occurring episodically, which probably affected the at-
mosphere and soils. This could have affected the development of the
flora by favoring only those plants able to cope with altered environ-
mental conditions, such as the fleshy stems and/or leaves with thick cu-
ticles present in Pleuromeia and Ptilozamites.
Acknowledgments
We want to express sincere gratitude to M. Carrizo, M.L. Balarino,
L.C.A. Martínez, and S. Mirabelli for their invaluable help in preparation
techniques. Thanks are also due to F. Tricárico for SEM assistance, and
P.R. Gutiérrez for comments and discussions on this material. Acknowl-
edgment is extended to both Eugenia and Victoria Zavattieri for art
work of Fig. 5. Valuable comments and suggestions by the two re-
viewers, E. Kustatscher and S. McLoughlin, greatly improved the final
version of the manuscript. This work was financially support by PICT
2011-2546, PICT 2012-1637, and PICT 2016-0431 financed by the
Agencia Nacional de Promoción Científica y Técnica (ANPCyT),
Argentina.
16 B. Cariglino et al. / Review of Palaeobotany and Palynology 257 (2018) 1–18
Fig. 5. Hypothetical reconstruction of the Quebrada de los Fósiles Formation palaeoenvironment. A. Pleuromeia, B. Sphenophytes, C. Ptilozamites, D. Lepacyclotes.
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