Journal of South American Earth Sciences 19 (2005) 547562

www.elsevier.com/locate/jsames

The Triassic Santa Juana Formation at the lower Biobo River,

south central Chile
Sven N. Nielsen
Institut fur geologische Wissenschaften, Fachrichtung Palaontologie, Freie Universitat Berlin, Malteserstrasse 74-100, 12249 Berlin, Germany
Accepted 15 June 2005

Abstract
A sequence of Triassic rocks is exposed near the town of Concepcion, Chile. These clastic strata are interpreted as the deposits of rivers,
lakes, playas, and alluvial fans. The deposits comprise conglomerates, arkosic sandstones, and sand-, silt- and mudstones. Four facies
associations comprising eight sedimentary facies can be distinguished. Plant fossils from the sedimentary sequence of the Santa Juana
Formation indicate a Carnian age. The flora includes ferns (Gleichenites, Cladophlebis, Dictyophyllum, Thaumatopteris, Asterotheca,
Saportaea) and seed ferns (Kurtziana, Antevsia, Dicroidium), ginkgophytes (Sphenobaiera), cycads (Pseudoctenis), conifers (Heidiphyllum,
Telemachus, Rissikia), and gymnosperms of uncertain affinities (Linguifolium, Gontriglossa). Two new species are presented: Pseudoctenis
santajuanensis and Gontriglossa reinerae.
q 2005 Elsevier Ltd. All rights reserved.
Keywords: Chile; Flora; Sedimentology; South America; Triassic

Resumen
En las proximidades de la ciudad de Concepcion, Chile, se exponen una secuencia de rocas Triasicos. Estas capas clasticas son
interpretadas como depositos de ros, lagos, lagos efimeros (playa-lakes), y abanicos aluviales. Esta secuencia comprende fanglomerados,
areniscas arcosicas, areniscas, lutitas y arcillolitas. Pueden ser distinguidas cuatro asociaciones de facies que incluyen ocho facies
sedimentarias. En la Formacion Santa Juana se describen restos de plantas, las quales indican una edad Carnico para esta secuencia. Como
elementos florales se pueden distinguir helechos (Gleichenites, Cladophlebis, Dictyophyllum, Thaumatopteris, Asterotheca, Saportaea) y
helechos con semillas (Kurtziana, Antevsia, Dicroidium), ginkgophitas (Sphenobaiera), cicadeas (Pseudoctenis), conferas (Heidiphyllum,
Telemachus, Rissikia), y algunos gimnospermas de atribucion incierta (Linguifolium, Gontriglossa). Se describen dos nuevas especies:
Pseudoctenis santajuanensis y Gontriglossa reinerae.
q 2005 Elsevier Ltd. All rights reserved.

1. Introduction
This paper presents sedimentologic, stratigraphic, and
fossil data for a sequence of Triassic rocks exposed in the
Chilean Coastal Cordillera along the lower Biobo River
west of the town of Concepcion, Chile (Fig. 1). These rocks
were classified as the Santa Juana Formation by Ferraris
(1981), but no clear definition of the formation was
provided. Here, the name Santa Juana Formation is used

E-mail address: nielsen@zedat.fu-berlin.de

0895-9811/$ - see front matter q 2005 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jsames.2005.06.002

for all Triassic deposits in the area surrounding the village

of Santa Juana (Fig. 1). The clastic rocks of the Santa Juana
Formation (Ferraris, 1981) were dated as Late Triassic in
age by Steinmann (1921) and Jaworski (1922) on the basis
of fossil plants and marine invertebrates. Newly collected
plant fossils necessitate a reinvestigation of the flora
previously described by Steinmann (1921) and Tavera
Jerez (1960).
The rocks of the Santa Juana Formation were deposited
in a NW-SEtrending basin formed during the initial
breakup of Gondwana and the beginning of the Mesozoic
subduction along its western margin, possibly directly
related to the Gastre fault system (Rapela and Pankhurst,
1992). The Santa Juana Basin is one of a series of other

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S.N. Nielsen / Journal of South American Earth Sciences 19 (2005) 547562

Fig. 1. Geological setting and location of the study area with localities of vertical profiles and the found flora.

Triassic basins in Chile (Charrier, 1979) and Argentina

(Uliana and Biddle, 1988) that are generally thought to be
rift related (Suarez and Bell, 1992). During deposition,
highlands surrounding the basin consisted of Paleozoic
metasediments in the north and northwest and granitoid
rocks of the southern Coastal Batholith in the remaining
parts. The Santa Juana Formation unconformably overlies
these crystalline basement rocks (Gonzalez-Bonorino and
Aguirre, 1970). Consequently, in the north and northwest of
the basin, coarse conglomerates consisting of mainly
metamorphic rocks occur at the base, and granitoid debris
is prominent in the basal part of the central and southern
area. Higher in the sequence, as well as basinward, the
formation becomes progressively finer grained. Dominated
by rivers and lakes, terrestrial sedimentation was interrupted
by at least one short marine ingression. However, this
ingression is recognizable not by sedimentological changes
but solely through the occurrence of marine fossils.
In the study area, gentle folding and intense faulting of
the Santa Juana Formation, together with the scarcity of
stratigraphically valuable fossils and lack of marker
horizons, make stratigraphic correlations among the isolated
outcrops extremely difficult. The few reported volcanic
rocks (Bruggen, 1934; Abad and Figueroa, 2003) and
anthracite seams (Alfaro and Helle, 2000) have not proven

to be of stratigraphic value because of their weathering and

alteration. These same stratigraphic and structural complexities prevent any definitive thickness estimates for the
formation, though such estimates in the literature range
from 300350 m (Felsch, 1913) to 10,00011,000 m
(Tavera Jerez, 1960).

2. Previous models
The first stratigraphical model, provided by Tavera Jerez
(1960), distinguished the following three members in the
eastern part of the Santa Juana Formation, from base to top:
(1) the Quilacoya Member or lower continental section, (2)
the Unihue Member or marine section, and (3) the
Talcamavida-Gomero Member or upper limnic member.
The Quilacoya Member contains carbon seams, plant
remains, and freshwater bivalves. It shows no marine
influence, and its thickness has been estimated as 1500 m.
The Unihue Member contains marine mollusks, mainly the
bivalve Halobia, and has an estimated thickness of 2500
3000 m. The Talcamavida-Gomero Member contains plant
remains, freshwater bivalves, and conchostracans and has an
estimated thickness of 30003500 m. The western part of the
formation is composed of approximately 3000 m of

S.N. Nielsen / Journal of South American Earth Sciences 19 (2005) 547562

549

Fig. 2. Generalized facies distribution of the Santa Juana Formation including alluvial fan (first facies association, facies 1 and 2), alluvial plain (second facies
association, facies 3 and 4), lakes (third facies association, facies 57), and marine deposits (fourth facies association, facies 8). AD refer to sections in Fig. 3.
Not to scale.

conglomerates (Tavera Jerez, 1960). The ages of the three

members are given as possibly Early Carnian for the
Quilacoya Member, Carnian for the Unihue Member, and
Norian-Rhaetian for the Talcamavida-Gomero Member
(Tavera Jerez, 1960).
Cucurella (1978), measuring the uranium, copper, lead,
and zinc contents in these sediments, basically used
Taveras model and placed conglomerates as a fourth
member at the base of the unit. The thickness of the
conglomerates (member 1) is given as 200 m, member 2 is a
250 m thick continental succession, member 3 comprises
400 m of shallow marine rocks, and member 4 is constituted
by approximately 500 m of coarse-grained delta deposits
(Cucurella, 1978).
During fieldwork in 1997, sedimentological studies led to
a facies model that is not consistent with this lithostratigraphic model. Previous models therefore are here regarded
as too simple for the complex history of the Santa Juana
Basin. Similar successions of facies have been observed in
areas that belong to different members in Tavera Jerezs
(1960) and Cucurellas (1978) models. Features such as
grain size vary laterally and are related to transport
mechanisms and the relative distance of source rocks rather
than capable of distinguishing lithological units. At this
stage, it is preferred to distinguish only those sedimentary
facies in the Santa Juana Formation and to compile a still
generalized model of facies development for the basin
(Fig. 2) rather than present a stratigraphic model based on
assumptions but lacking any real stratigraphic data.
3. Sedimentary facies
3.1. Facies 1: granitoid debris
Facies 1 consists mainly of coarse quartz grains derived
from the granitoid basement. The amount of other

components that compose the fine-grained matrix varies

considerably. For the study area, the base of the sedimentary
cover on the granitoid basement (Fig. 3A) is difficult to
define because the transition from unweathered to
weathered granitoid basement and to unstratified granitoid
debris seems gradational. Upward in the section and
basinward, stratification becomes recognizable, and the
portion of finer-grained material increases. A transition to
facies 4, which sometimes occurs through facies 2 and/or
facies 3, takes place in a general fining- and thinningupward tendency. Even in coarse granitoid debris, fossilized
tree trunks up to 2 m long indicate colonization by
arboraceous plants. In finer-grained granitoid debris,
unidentifiable plant remains are common.
3.2. Facies 2: conglomerates and arkosic sandstones
The sedimentary strata overlying Paleozoic metamorphic
rocks are composed of conglomerates and arkosic sandstones, with bed thicknesses of several meters (Fig. 3B).
Beds with erosive bases are rare.
The weathered sediments have a yellowish to red color.
Scanning electron microscopy (SEM) examinations
revealed a high portion of unweathered idiomorphic
feldspar. The conglomerates are poorly sorted, with
predominantly angular to subrounded clasts that vary in
size from 2 to more than 20 cm. Most conglomerates are
supported by a heterogeneous sandy matrix. Only a few
clast-supported beds have been observed. Dominant largescale sedimentary features are continuous, and parallel
bedding planes are defined by color and grain-size
variations. Internally, the beds tend to be structureless,
chaotic, and disorganized. No bioturbation was observed.
Above the conglomerates and arkosic sandstones, finergrained sandstones, siltstones, and mudstones (facies

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S.N. Nielsen / Journal of South American Earth Sciences 19 (2005) 547562
Fig. 3. Selected lithosections from different localities (see Fig. 1, Table 2). For facies classification of A, B, and D, see Table 1. (A) Section near Quilacoya showing contact zone between granitoid basement and
sedimentary cover. (B) Section near Patagual showing basal part of sedimentary cover on metamorphic shales. (C) Section at the railway between Gomero and Buenuraqui. (D) Section near Santa Juana
illustrating black shale-sandstone alternation.

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551

3 and 4) become dominant, and plant debris becomes

common (mainly Heidiphyllum). This tendency of finingand thinning-upward beds corresponds to that of facies 1.
Facies 2 is interpreted as alluvial fan deposits. The poorly
sorted and poorly rounded coarse-grained sediments
indicate a short transport distance and therefore were
deposited relatively proximal to the basin margin. The
character of these sediments seems to indicate arid or
semiarid climate conditions, but the occurrence of few coal
seams suggests occasional deposition during more humid
intervals.

where few distinct sedimentary structures, mostly parallel

bedding and occasionally ripple marks, are preserved. Some
conglomerates with well-rounded, well-sorted clasts of
predominantly metamorphic origin occur. On the basis of
the occurrence of ripple marks and well-rounded conglomerates, as well as the relationship with other facies,
deposition is interpreted to have taken place near the shores
of lakes or river banks or on inactive, muddy, vegetated
overbank areas between sandy river channels.

3.3. Facies 3: sandstones and siltstones

The soft, easily weathered nature of these rocks results in

poor natural exposures. Only one good section is available
near the village of Santa Juana (Fig. 3D), where a bypass
cuts these mudstones. However, some small outcrops also
exist on both banks of the Biobo River. On some surfaces,
conchostracans (Estheria forbesii Jones 1862 of Tavera
Jerez, 1960; Fig. 4(1)) are very abundant, and isolated plant
fragments and insect remains occur. The shales are rich in
organic carbon and partially bioturbated. These deposits
suggest accumulation in the subaquatic part of a fan delta or
talus deposits with occasional turbidity currents (facies 6).
Finely laminated black shales are interpreted as the product
of settling from suspension in relatively deep water, as
indicated by facies 6. However, some of the black shales
may have been deposited in abandoned channels or lakes,
leading to the still conditions preferred by conchostracans.
Assignment to one of the two rather different environmental
settings is based on the association with facies 6 (deeper
water) or the occurrence of conchostracans (still shallow
water).

Facies 3 consists mainly of yellowish to red, weathering,

interbedded sandstones and siltstones. Red-brown staining
is interpreted as due to oxidation of pyrite contained in
carbonaceous plant debris (Bell and Suarez, 1995). Some
siltstone layers are cross-laminated, whereas some sandstone bedding surfaces contain asymmetrical current ripple
marks. Root traces are also common. A rich flora was found
in silty layers near Patagual (upper part of Fig. 3B).
Steinkerns of freshwater bivalves, probably of the family
Unionidae, and plant remains occur at the railway cut near
Buenuraqui (Fig. 3C). At this location, the rocks are less
weathered and dark gray in color. Frequent fining-upward
cycles and occasional normally graded sandstones were also
observed in this section, as well as lenticular beds
interpreted as channel fills. This outcrop shows gentle
folding and normal faults with dislocation rates of less than
5 m.
The facies assemblage and upward-fining cycles
(Fig. 3C) indicate distal rivers and alluvial plain deposits.
The repeated succession of structures like ripple marks, root
horizons, and layers with plant debris (Fig. 3B, upper part)
suggests deposition in floodplain lakes that were later
colonized by rooting plants.
3.4. Facies 4: siltstones and mudstones
Facies 4 comprises silt- and mudstones that contain a rich
flora. A diverse association of ferns and gymnosperms was
recovered mainly from outcrops at the Cerro Calquinhue,

3.5. Facies 5: black shales

3.6. Facies 6: sandstones and siltstones

The coarse- to fine-grained sandstones representing
facies 6 usually occur alternating with the black shales of
facies 5 (Fig. 3D). Sedimentary structures like slumping,
fining-upward sequences, and reworked shale intraclasts are
common. These sandstones are interpreted as lacustrine
turbidites. Occasionally, load structures and flame structures
are present at contact planes with intercalated black shales
of facies 5. These structures, together with slumping, might

Fig. 4. (1) Conchostraca indet., height of photo 8 mm. (23) Halobia sp., scale bar 10 mm.

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S.N. Nielsen / Journal of South American Earth Sciences 19 (2005) 547562

Table 1
Facies classification (from Miall, 1996)
Facies code

Facies

Sedimentary structures

Interpretation

Gmm

Matrix supported,
massive gravel

Weak grading

Plastic debris flow (high-strength,

viscous)

Sr

Sand, very fine to

coarse

Ripple cross-lamination

Ripples (lower flow regime)

Sh

Sand, very fine to

coarse, may be
pebbly
Sand, fine to
coarse

Horizontal lamination parting or

streaming lineation

Plane-bed flow (critical flow)

Massive, or faint lamination

Sediment-gravity flow deposits

Fl

Sand, silt, mud

Fine lamination, very small ripples

Overbank, abandoned channel, or

waning flood deposits

Fsm

Silt, mud

Massive

Backswamp or abandoned channel

deposits

Fr

Mud, silt

Massive, roots, bioturbation

Root bed, incipient soil

Sm

be related to earthquake-induced liquefaction rather than

load-related fluid loss, but no unequivocal structures have
been observed.
3.7. Facies 7: massive sandstones
Facies 7 comprises massive sandstones with rare
intercalations of well-rounded conglomerates. The massive
sandstones contain sedimentary structures such as channel
fills, slumping, fining-upward sequences, convolution,
reworked shale intraclasts, and load casts. A few trunks of
possibly driftwood have been recorded. Near the village of
Santa Juana, this facies is exposed at both banks of the
Biobo River. Locally the massive sandstones have a dark
gray color and are rich in organic carbon. Some finer layers
contain unidentifiable plant remains. These massive
sandstones are interpreted as mass-flow deposits. On top
of these deposits, rivers cut channels and deposited
conglomerates. The erosive nature is indicated by reworked
finer sediment, such as shale intraclasts. However, the base
of this facies was not found, so an interpretation of the place
of deposition is not currently possible.
3.8. Facies 8: marine siltstones
Facies 8 contains siltstones superficially indistinguishable from those of facies 4 or 6. The abundant marine
bivalve Halobia (Fig. 4(23)) indicates a marine environment (e.g., Campbell, 1994). Bioturbation led to knobby
surfaces. Other marine fossils could not be found. However,
Jaworski (1922) and Tavera Jerez (1960) reported ammonoids (Arcestes, Trachyceras?), bivalves (Palaeoneilo,
Lima, Myoconcha, Phaenodesmia, Myophoria), and a
gastropod (Turbo). No section was measured due to the
small scale of the outcrops.

3.9. Facies associations

The sedimentary rocks of the Santa Juana Formation
have been subdivided into eight facies (Fig. 2) grouped into
four facies associations (Fig. 3, Table 1). The first facies
association comprises granitoid debris (facies 1) and alternating conglomerates and arkosic sandstones (facies 2).
These two facies are interpreted as deposits of alluvial fans.
The second facies association contains sandstones to
siltstones (facies 3) and siltstones to mudstones (facies 4),
interpreted as alluvial deposits. The third facies association
comprises black shales containing conchostracans (facies 5,
Fig. 4(1)) alternating with sand- to siltstones (facies 6) and
locally massive sandstones (facies 7). These deposits are
interpreted as accumulating in a sublacustrine setting with
occasional turbidity currents and mass flows. The fourth
facies association consists of marine mudstones, sandstones,
and bioturbated siltstones (facies 8) that contain the
abundant bivalve Halobia sp. (Fig. 4(23)).

4. The paleoflora
Descriptions of fossil plant remains from the Santa Juana
Formation, mostly without figures, have been published by
Steinmann (1921) and Tavera Jerez (1960). The flora of the
Santa Juana Formation is a typical Gondwana Upper
Triassic Dicroidium flora, similar to the Molteno flora of
southern Africa (Anderson and Anderson, 1983, 1984,
1989). However, in contrast to the Molteno flora, the Santa
Juana flora is dominated by Heidiphyllum and Pseudoctenis
with Dicroidium as a subordinate component.
Locality 1 (Fig. 1, Table 2) comprises two outcrops at the
Cerro Calquinhue that were cut by a forest road. The
sediments have been deposited by rivers in an alluvial plain
(facies 4). Locality 2 (Fig. 1, Table 2) is located along

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553

Table 2
Coordinates of vertical profiles and flora. Map: data taken from map, GPS: measured by GPS
Section near Quilacoya (Fig. 3A)
Section near Patagual (Fig. 3B)
Section between Gomero and Buenuraqui (Fig. 3C)
Section near Santa Juana (Fig. 3D)
Locality 1 at the Cerro Calquinhue
Locality 2 between Quilacoya and Chillancito
Locality 3 near Patagual

S37803 0 /W073857 0 (Map)

S37801,307 0 /W073801,303 0 (GPS)
S37812,138 0 /W072847,665 0 (GPS)
S37810,270 0 /W072857,406 0 (GPS)
S37805,595 0 /W072853,534 0 (GPS)
S37804,710 0 /W072856,328 0 (GPS)
S37801,5 0 /W073801,5 0 (Map)

Fig. 5. (1) Gleichenites sp., Cp 1299. (2) Dictyophyllum fuenzalidai Herbst (2000), Cp 1277. (3) Thaumatopteris rothii Frenguelli 1941, Cp 1284. (4)
Cladophlebis sp., Cp 1270. (5) Dictyophyllum tenuifolium (Stipanicic and Menendez 1949) Bonetti and Herbst 1964, Cp 1291. (6) Asterotheca fuchsii
(Schimper ex Zeiller 1875) Herbst 1998, Cp 1282. (7) Saportaea dichotoma (Frenguelli, 1942) Stipanicic and Bonetti (1965), Cp 1297. Scale bar for all figures
is 10 mm.

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S.N. Nielsen / Journal of South American Earth Sciences 19 (2005) 547562

Fig. 6. (1) Kurtziana cacheutensis (Kurtz 1921) Frenguelli 1941, Cp 1287. (2) Antevsia (?) sp., UdeC. (3) Dicroidium crassinervis (Geinitz 1876) Anderson and
Anderson (1983) forma trilobitum (Johnston 1886) Anderson and Anderson (1983), Cp 1285. (4) Dicroidium odontopteroides (Morris 1845) Gothan 1912
forma lineatum (Tenison-Woods 1883) Anderson and Anderson (1983), UdeC. (5) Dicroidium elongatum (Carruthers 1872) Archangelsky 1968 forma
remotipinnulum Anderson and Anderson (1983), Cp 1298. (6) Seed of Dicroidium, UdeC. (7) Dicroidium elongatum (Carruthers 1872) Archangelsky 1968

S.N. Nielsen / Journal of South American Earth Sciences 19 (2005) 547562

the road from Quilacoya to Chillancito, where the marine

facies (facies 8) is exposed in a small outcrop. Locality 3
(Fig. 1, Table 2) is situated along the road from the Biobo
River to Coronel near the village of Patagual. Similar to
locality 1, the exposed sediments have been deposited by
rivers in an alluvial plain (facies 3).
4.1. Material
Material is stored at the following sites: (Cp) Geologisch-Palaontologisches Institut und Museum, Abteilung
Palaobotanik, Westfalische Wilhelms-Universitat Munster,
Germany; (UdeC) Santa Juana Collection, Departamento
Ciencias de la Tierra, Universidad de Concepcion, Chile.
UdeC specimens are not numbered.
4.2. Ferns
Gleichenites sp.
(Fig. 5(1))
Material examined: Cp 1299.
Locality: 1.
Description: Axis 62 mm long. Pinnae 5060 mm long.
Central vein of pinnules 5 mm long.
Stratigraphic range: Triassic-Middle Jurassic.
Distribution: Gleichenites is known worldwide.
Systematic position: Gleicheniaceae.
Remarks: This specimen shows similarities with Gleichenites gallegoii Herbst, 1996 from Argentina and Gleichenites
sp. from the Santa Juana Formation (Herbst, 1996). It is
believed to represent a new species because of its differing
pinnule margins. However, characters are too few to allow a
satisfactory diagnosis. Similarities exist with the genus
Todites, especially Todites goeppertiana (Munster 1846)
Krasser 1922, as figured by Harris (1931, Fig. 6C/D) from
eastern Greenland. However, to prove its position relative to
Todites, sporangia of the Osmundaceae must be present.
Cladophlebis sp.
(Fig. 5(4))
Material examined: Cp 1270, Cp 1271.
Locality: 2.
Description: Terminal part of pinna, 60 mm long and
15 mm wide.
Systematic position: Cladophlebis is regarded as belonging to the Osmundales.
Remarks: Steinmann (1921) reported Cladophlebis
roessertii (Presl 1838) Saporta 1873 (ZGleichenites
roessertii (Presl 1838) Kryshtofovich 1912) and Cladophlebis australis (Morris 1845) Halle 1913 from the Santa
Juana Formation. The specimen figured cannot be attributed

555

to one of these species; a more precise identification is not

possible due to the fragmentary nature of the material.
Dictyophyllum fuenzalidai Herbst (2000)
(Fig. 5(2))
Material examined: Cp 1277.
Locality: 1.
Description: Specimen, 63 mm long showing an axis
with lateral veins positioned at distances of 11 mm.
Distribution: Chile.
Systematic position: Dipteridaceae.
Remarks: Steinmann (1921) and Tavera Jerez (1960)
reported Clathropteris platyphylla (Goeppert 1842) Brongniart 1849 from the Santa Juana Formation, which seems
very similar. However, though the venation is not clear, it is
regarded as Dictyophyllum fuenzalidai Herbst (2000), as
described from near Copiapo (Herbst, 2000).
Dictyophyllum tenuifolium (Stipanicic and Menendez
1949) Bonetti and Herbst 1964
(Fig. 5(5))
Material examined: Cp 1290, Cp 1291, Cp 1292.
Locality: 1.
Description: Frond fragment 120 mm long and 30 mm
wide. Axis of specimen 105 mm long.
Stratigraphic range: Carnian.
Distribution: Argentina.
Systematic position: Dipteridaceae.
Remarks: This species shows similarities to Dictyophyllum ellenbergii Greber 1960 (see Fabre and Greber, 1960;
Anderson and Anderson, 1983) from the Molteno Formation, southern Africa.
Thaumatopteris rothii Frenguelli 1941
(Fig. 5(3))
Material examined: Cp 1284.
Locality: 1.
Description: Specimen with 75 mm long axis. Pinnae
approx. 70 mm long and 13 mm wide. Venation is poorly
preserved.
Stratigraphic range: Late TriassicEarly Liassic.
Distribution: Chile, Argentina.
Systematic position: Dipteridaceae.
Remarks: Herbst (1992) regards Thaumatopteris as a
subgenus of Dictyophyllum. Because venation is poorly
preserved, the identification remains uncertain.
Asterotheca fuchsii (Schimper ex Zeiller 1875) Herbst
1998
(Fig. 5(6))
Material examined: Cp 1282.
Locality: 1.
Description: Frond fragment approximately 43 mm long
and 28 mm wide. Fragment shows second- and third-order

3
subsp. argentinum (Kurtz 1921) Anderson and Anderson (1983) or Sphenobaiera pontifolia Anderson and Anderson (1989), Cp 1295, on holotype of
Gontriglossa reinerae sp. nov. (8) Sphenobaiera africana (Baldoni 1980) Anderson and Anderson (1989), Cp 1298, same slab as Fig. 6(5). (911)
Pseudoctenis santajuanensis sp. nov. (9) Apical portion of frond, Cp 1300. (1011) Holotype, Cp1301. Scale bar for all figures is 10 mm.

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S.N. Nielsen / Journal of South American Earth Sciences 19 (2005) 547562

pinnules. Third-order pinnules (4 mm long and 2 mm wide)

have well-rounded tips with a marked midrib.
Stratigraphic range: Triassic.
Distribution: Chile, Argentina, Australia.
Systematic position: Asterothecaceae.
Remarks: Even if there are no synangia preserved, this
specimen is regarded as Asterotheca fuchsii because of its
pinnule morphology. However, there is a possibility that this
frond fragment belongs to Cladophlebis (R. Herbst, pers.
comm.)
Saportaea dichotoma (Frenguelli, 1942) Stipanicic and
Bonetti (1965).
(Fig. 5(7))
Material examined: Cp 1297.
Locality: 1.
Description: Frond fragment 80 mm long and 80 mm
wide. Lamina deeply segmented. Veins not preserved, only
prominent midrib present.
Stratigraphic range: uppermost Ladinian.
Distribution: Argentina.
Systematic position: Saportaea is regarded as belonging
to the Polypodiales (Anderson and Anderson, 1993).
Remarks: Assignment after the definition provided by
Frenguelli (1942) and Stipanicic and Bonetti (1965) rather
than in the broader sense of Anderson and Anderson (1989).
4.3. Seed ferns
Kurtziana cacheutensis (Kurtz 1921) Frenguelli 1941
(Fig. 6(1))
Material examined: Cp 1287.
Locality: 1.
Description: Frond 80 mm!18 mm. Tip and base
missing. Venation usually twice forking.
Stratigraphic range: Late Triassic.
Distribution: Argentina, Southern Africa.
Systematic position: This species is regarded as a
peltasperm (Anderson et al., 1998) or a cycad (Ganuza
et al., 1998).
Antevsia (?) sp.
(Fig. 6(2))
Material examined: UdeC.
Locality: 2.
Description: Maximum width of specimen 15 mm.
Stratigraphic range: Because Antevsia is the male
fructification of the peltasperms, an occurrence from Late
Permian to Late Triassic is presumed.
Systematic position: Peltaspermaceae.
Remarks: This specimen is on the back side of a slab with
Dicroidium odontopteroides.
Dicroidium crassinervis (Geinitz 1876) Anderson and
Anderson (1983)
forma trilobitum (Johnston 1886) Anderson and
Anderson (1983)

(Fig. 6(3))
Material examined: Cp 1285.
Locality: 1.
Description: Frond forked, pinnate, medium sized
(100 mm long and 20 mm wide). Three-lobed pinnae.
Stratigraphic range: LadinianNorian.
Distribution: Argentina, Southern Africa, Antarctica,
Australia.
Systematic position: Umkomasiaceae.
Dicroidium odontopteroides (Morris 1845) Gothan 1912
forma lineatum (Tenison-Woods 1883) Anderson and
Anderson (1983)
(Fig. 6(4))
Material examined: UdeC.
Locality: 2.
Description: Frond 150 mm long, 40 mm wide.
Stratigraphic range: LadinianCarnian.
Distribution: Argentina, Brazil, Southern Africa, India,
Australia.
Systematic position: Umkomasiaceae.
Remarks: This specimen has previously been
figured by Bandel et al. (1997, Fig. 2b). It is associated
with marine bivalves (Halobia), Linguifolium, and Antevsia.
Dicroidium elongatum (Carruthers 1872) Archangelsky
1968
forma remotipinnulum Anderson and Anderson (1983)
(Fig. 6(5))
Material examined: Cp 1298.
Locality: 1.
Description: Frond forked, approximately 40 mm, pinnae
linear.
Stratigraphic range: LadinianCarnian.
Distribution: Southern Africa, Australia, Antarctica.
Systematic position: Umkomasiaceae.
Remarks: This specimen is located between some
Sphenobaiera africana and is poorly preserved.
Seed of Dicroidium
(Fig. 6(6))
Material examined: Cp 1303, Cp 1276.
Locality: 1, 3.
Description: Diameter of seed 13 mm.
Systematic position: Umkomasiaceae.
Remarks: Dicroidium seed similar to that
figured by Anderson and Anderson (1983). This
specimen has previously been figured by Bandel et al.
(1997, Fig. 2c).
4.4. Seed fern or ginkgophyte
Dicroidium elongatum (Carruthers 1872) Archangelsky
1968
subsp. argentinum (Kurtz 1921) Anderson and Anderson
(1983)

S.N. Nielsen / Journal of South American Earth Sciences 19 (2005) 547562

or Sphenobaiera pontifolia Anderson and Anderson

(1989)
(Fig. 6(7))
Material examined: Cp 1295, Cp 1296.
Locality: 1.
Description: This specimen is approximately 70 mm
long and 10 mm wide.
Stratigraphic range: LadinianCarnian (D. elong. arg.);
LadinianNorian (S. pontifolia).
Distribution: Argentina, Southern Africa, Antarctica
(D. elong. arg.); Argentina, Brazil, Southern Africa,
Antarctica, Australia (S. pontifolia).
Systematic position: Ginkgoopsida.
Remarks: Dicroidium elongatum argentinum and Sphenobaiera pontifolia are indistinguishable without preserved
cuticles (Anderson and Anderson, 1989) or more complete
specimens; therefore, exact determination of this specimen
is impossible.
4.5. Ginkgophytes
Sphenobaiera africana (Baldoni 1980) Anderson and
Anderson (1989)
(Fig. 6(8))
Material examined: Cp 1298.
Locality: 1.
Description: The bigger frond is 85 mm to 15 mm.
Lamina fork three to four times into 816 small linear
segments. Base, tips, and venation are not preserved.
Stratigraphic range: Carnian.
Distribution: Argentina, Southern Africa, Australia.
Systematic position: Ginkgoales.
4.6. Cycads
Pseudoctenis santajuanensis sp. nov.
(Fig. 6(911)
Material examined: Cp 1301 (holotype), Cp 1287, Cp
1300, Cp 1302, Cp 1304, Cp 1306, Cp 1307 (paratypes).
Locality: 1.
Diagnosis: Pinnate frond. Pinnae perpendicularly positioned to the rachis, widely spaced, attached to the upper
surface of the pinna rachis. Pinnae slightly constricted at the
base, widening toward the apex; apex truncate. Pinnae up to
50 mm long and 5 mm wide. Venation dense, up to 25 veins
per 10 mm; veins running more or less parallel, forking once
or twice.
Holotype: Specimen Cp 1301.
Type locality: Cerro Calquinhue, near Quilacoya.
Stratigraphic range: AnisianCarnian.
Systematic position: Cycadales.
Name derivation: After the Santa Juana Formation.
Remarks: This species differs from other species of
Pseudoctenis in having perpendicularly positioned pinnae
with conspicuously constricted bases, in the shape of its
pinna apices, and in its size. The most similar species is

557

P. azcaratei Herbst and Troncoso (2000), from which it

differs by its wider-spaced pinnae and less dense venation
(28 veins per 10 mm in P. azcaratei). Herbst and Troncoso
(2000) mention additional material of P. azcaratei from the
Santa Juana Formation; however, it might prove to belong to
P. santajuanensis.
cf. Pseudoctenis fissa Du Toit 1927
(Fig. 7(1))
Material examined: Cp 1274.
Locality: 3.
Description: Fragment of frond 32 mm long, 16 mm
wide. Venation poor, moderately spaced.
Stratigraphic range: AnisianCarnian.
Distribution: Argentina, Southern Africa.
Systematic position: Cycadales.
Remarks: This specimen is placed within P. fissa even if
considerable doubt remains because the tip is unforked,
which seems untypical for P. fissa (see Anderson and
Anderson, 1989). However, there is not sufficient material
available for a safe determination.
4.7. Conifers
Heidiphyllum elongatum (Morris 1845) Retallack 1981
(Fig. 7(2))
Material examined: Cp 1300, Cp 1302, Cp 1303, Cp
1304.
Locality: 1.
Description: Leaf relatively large (170 mm long, 15 mm
wide), narrowly elliptic. Veins well spaced, forking once or
twice.
Stratigraphic range: AnisianRhetian.
Distribution: Gondwana.
Systematic position: Voltziaceae.
Telemachus elongatus Anderson (1978)
(Fig. 7(3))
Material examined: Cp 1300, Cp 1305.
Locality: 1.
Description: Specimen, 40 mm long and 20 mm wide.
Stratigraphic range: AnisianRhetian.
Distribution: Chile, Southern Africa, New Zealand,
Antarctica.
Systematic position: Voltziaceae.
Remarks: Anderson (1978) and Anderson and Anderson
(1989) proposed that Telemachus as a female cone and
Heidiphyllum leaves are part of the same plant. Even if there
remained some doubt, the coexistence at least 14 localities
worldwide (Axsmith et al., 1998) serves as additional
evidence.
Rissikia media (Tenison-Woods 1883) Townrow 1967
(Fig. 7(4))
Material examined: Cp 1275.
Locality: 3.

558

S.N. Nielsen / Journal of South American Earth Sciences 19 (2005) 547562

Fig. 7. (1) cf. Pseudoctenis fissa Du Toit 1927, Cp 1274. (2) Heidiphyllum elongatum (Morris 1845) Retallack 1981, Cp 1302. (3) Telemachus elongatus
Anderson (1978), Cp 1300. (4) Rissikia media (Tenison-Woods 1883) Townrow 1967, Cp 1275. (5) Linguifolium steinmannii (Solms-Laubach 1899)
Frenguelli 1941, Cp 1288. (69) Gontriglossa reinerae sp. nov. (6) Counterpart of holotype, Cp 1296. (7) Holotype, Cp 1295. (89) Paratype, Cp 1294. Scale
bar for all figures is 10 mm.

Description: Foliage shoot (30 mm long, 7 mm wide).

Pinnae linear.
Stratigraphic range: LadinianCarnian.
Distribution: Gondwana.
Systematic position: Podocarpaceae.
4.8. Gymnosperms incertae sedis
Linguifolium steinmannii (Solms-Laubach 1899)
Frenguelli 1941
(Fig. 7(5))
Material examined: Cp 1288, Cp 1289.

Locality: 1.
Description: Leaf 75 mm long and 14 mm wide, base
and tip missing. Veins well spaced (12 per 10 mm),
spreading relatively widely from midrib, arching
gently.
Stratigraphic range: LadinianRhetian.
Distribution: Chile, Argentina, Australia, New Zealand.
Systematic position: This species is now regarded as a
ginkgoopsid (Anderson and Anderson, 1993).
Remarks: After recent reports, it seems possible that
Linguifolium also lived in the northern hemisphere (e.g., in
Germany; Kelber, 1998).

S.N. Nielsen / Journal of South American Earth Sciences 19 (2005) 547562

Gontriglossa reinerae sp. nov.

(Fig. 7(69))
Material examined: Cp 1293 (paratype), Cp 1294
(paratype), Cp 1295 (holotype), Cp 1296 (holotype
counterpart).
Locality: 1.
Diagnosis: Leaf relatively large (approx. 150 mm long
and 70 mm wide), presumably elliptical. Base and apex
unknown. Veins frequently anastomosing, arching very
steeply (10208 from midrib).
Holotype: Cp 1295, Cp 1296 (counterpart).
Type locality: Cerro Calquinhue, near Quilacoya.
Stratigraphic range: LadinianCarnian.
Systematic position: Gontriglossa is regarded as belonging to the Gnetopsida (Anderson et al., 1998).
Name derivation: For my friend and colleague Janet
Zulauf, who at the time of description still held her name of
birth Reiner.
Remarks: Even if its leaf-base and cuticle are unknown,
this species is assigned to the genus Gontriglossa because it
is generally assumed that the Glossopteridales became
extinct at the PermianTriassic boundary.
Discussion: This species is larger than most other species
of Gontriglossa, such as the South African G. verticillata
(Thomas) (Anderson and Anderson, 1989) and the
Australian G. moribunda (Johnston) (Holmes, 1992).
Similarly large species are the Australian G. lacerata
(Holmes), G. nymboidensis (Holmes), and G. grandis
(Walkom) (Holmes, 1992), which have a less steep angle
between the midrib and veins. This angle is 65708 in

559

G. grandis and 20308 in G. nymboidensis and G. lacerata

but only 10208 in G. reinerae. In G. nymboidensis and
G. lacerata, the veins quickly curve outward, reaching an
angle of 708 to the midrib, whereas the veins in G. reinerae
remain always steeper and maintain a maximum angle of
approximately 508 to the midrib.

5. Biogeography and ecology

The flora of the Santa Juana Formation comprises mainly
elements that have a wide distribution in the Late Triassic of
Gondwana (Table 3). Close affinities exist throughout
Gondwana, especially in Argentina, southern Africa,
and Australia, where the flora are best known. Similarly
rich floras probably could be found at other localities; lack
of sampling in other areas appears to be the main reason the
Molteno flora stands out. The nongymnosperms of the
Molteno Formation are currently under revision (Anderson
and Anderson, in prep.), but a comparison is not possible at
the moment.
The ecology of Triassic plants has been discussed by
Anderson et al. (1998). While ferns grew in a wide variety
of habitats, other forms lived in more ecologically restricted
areas. Heidiphyllum, Rissikia, and Kurtziana preferred wet
areas on floodplains; Pseudoctenis, Saportaea, and
Dicroidium grew in forests to woodlands; Sphenobaiera,
Linguifolium, and Gontriglossa preferred water margins. This
pattern also seems to apply to the Santa Juana Formation. It
is therefore obvious that during the Late Triassic, at least

Table 3
Distribution of the plants found in the Santa Juana Formation
Species

Ch

Ar

Br

Mt

Au

Ts

NZ

Ind

Ant

Gleichenites sp.
Cladophlebis sp.
Dictyophyllum fuenzalidai
Dictyophyllum tenuifolium
Thaumatopteris rothii
Asterotheca fuchsii
Saportaea dichotoma
Kurtziana cacheutensis
Dicroidium crassinervis forma
trilobitum
Dicroidium odontopteroides
forma lineatum
Dicroidium elongatum remotipinnulum
D. elong. arg. / Sph. pontif.
Sphenobaiera Africana
Pseudoctenis gracipinnata
Pseudoctenis fissa
Heidiphyllum elongatum
Rissikia media
Linguifolium steinmannii
Gontriglossa reinerae

(*)
(*)
*
(*)

(*)
(*)

(*)

(*)

(*)

(*)

(*)

(*)

(*)

*
*

(*)

*
*
*

*
*
*
*
*
*

(*)
(*)
(*)
*
*

*/*
*
*
*
*
*
(*)
(*)

*
*
*

*
(?)

*
*
*
(*)

*
*
*
*
*
*
*
(?)

*
(*)

(*)

(*)
(*)
(?)

(?)

*
*

(*)

(?)

*
*
*

*
(?)

*
(*)

*
*
*

(?)

Ch: Chile (excluding this work); Ar: Argentina; Br: Brazil; Mt: Molteno Formation, Southern Africa; Au: East Australia; Ts: Tasmania; NZ: New Zealand; Ind:
India; Ant: Antarctica. *Occurrence of species; (*) occurrence of genus; (?) supposed occurrence of genus.

560

S.N. Nielsen / Journal of South American Earth Sciences 19 (2005) 547562

Table 4
Known stratigraphic range of the found taxa (derived mainly from Anderson and Anderson, 1983, 1989)

Black: range of species, grey: range of genera.

southern Gondwana had a very homogeneous flora that

reflects environments rather than geographical areas.

6. Stratigraphy
The stratigraphic ranges of several of the plant taxa
encountered (Table 4) are still unsatisfactorily known
because some are only known from a few, poorly dated
localities. Species such as Saportaea dichotoma, Kurtzia
cacheutensis, and Sphenobaiera africana are only known
from one dated locality each, and the presented
stratigraphic range certainly does not represent reality.

Therefore, no good age assessment based on the paleoflora

can be made for the Santa Juana Formation. However,
regarding the known ranges presented in Table 4, a
Carnian age seems most plausible. The nonmarine
invertebratesbivalves, conchostracans, and insectsdo
not provide good stratigraphical evidence. The marine
bivalve genus Halobia is a good index fossil, but the
material reported here probably represents a new species.
Other marine invertebrates, especially ammonoids, might
give a more exact age than currently is possible. Still,
further work, including palynological investigations and
absolute dating of volcanic rocks, is needed for a good age
assessment.

S.N. Nielsen / Journal of South American Earth Sciences 19 (2005) 547562

7. Conclusions
The succession of thick, terrigenous, clastic sediments of
the Santa Juana Formation represents sedimentation in an
actively subsiding basin. The sediments were derived
predominantly from Upper Paleozoic plutonic and metamorphic rocks. Deposition was likely to have been in a basin
directly related to the Gastre fault system of Rapela and
Pankhurst (1992). The Santa Juana Formation was
deposited on an alluvial braidplain that was superimposed
by a large-scale braided river system with lakes, playa lakes,
and alluvial fans. The intercalation of alluvial and fluvial
sediments contradicts a climatic model that assumes a
change from a semiarid to a more temperate climate regime.
It is more likely that these alluvial fluvial intercalations
reflect tectonic and seasonal influences on sedimentation. A
rich paleoflora is described, dating the Santa Juana
Formation as probably Carnian. Few nonmarine invertebrates support the environmental reconstructions. A
marine horizon yields only the bivalve Halobia, but other
marine invertebrates reported in the literature might give
better ages for the Santa Juana Formation in the future.

Acknowledgements
Klaus Bandel (Geologisch-Palaontologisches Institut,
Universitat Hamburg, Germany) is thanked for field
support and discussions about the Santa Juana Formation.
Klaus Reicherter (Institut fur Geophysik, Universitat
Hamburg, Germany) is thanked for discussions and
critical reviews of an early draft of the manuscript.
Heidi Anderson (Botanical Research Institute, Pretoria,
South Africa) and Rafael Herbst (PRINCEPA-CONICET,
Corrientes, Argentina) made valuable comments on the
determination of the plants. Paulina Vasquez (Institut fur
Angewandte Geowissenschaften, TU Berlin, Germany)
corrected the Spanish abstract. Revisions by Brian Currie
(Department of Geology, Miami University, USA) and
Heidi Anderson significantly improved this work. Field
study was financially supported by the Emmy and Alfred
B. Steffens Memorial Fund of the University of
Hamburg.

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