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Palynology of the Lower Permian of Paraná Basin, Uruguay

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Palynology of the Lower Permian of Paraná Basin, Uruguay

Pedro R. Gutiérreza; María Lucía Balarinoa; Ángeles Berib
a
CONICET, Sección Paleopalinología, División Paleobotánica, Museo Argentino de Ciencias Naturales
“B. Rivadavia”, Argentina b Departamento de Paleontología, Facultad de Ciencias, Montevideo,
Uruguay

Online publication date: 05 November 2010

To cite this Article Gutiérrez, Pedro R. , Balarino, María Lucía and Beri, Ángeles(2010) 'Palynology of the Lower Permian
of Paraná Basin, Uruguay', Journal of Systematic Palaeontology, 8: 4, 459 — 502
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 Journal of Systematic Palaeontology, Vol. 8, Issue 4, December 2010, 459–502

Palynology of the Lower Permian of Paraná Basin, Uruguay

Pedro R. Gutiérreza∗, Marı́a Lucı́a Balarinoa and Ángeles Berib
a
CONICET, Sección Paleopalinologı́a, División Paleobotánica, Museo Argentino de Ciencias Naturales “B. Rivadavia”, Av. Ángel
Gallardo 470, C1405DJR Buenos Aires, Argentina; bDepartamento de Paleontologı́a, Facultad de Ciencias, Iguá 4225, 11400,
Montevideo, Uruguay
(Received 10 August 2009; accepted 17 December 2009)

The palynological contents of the Permian Tres Islas and Melo formations (Frayle Muerto, Mangrullo and Paso Aguiar
members), obtained from boreholes and outcrop in northeastern Uruguay, are reported. These Permian sediments of the
Paraná Basin yielded a total of 147 species, including the following new taxa: Archangelskiapollenites globocorpus gen.
et sp. nov., A. plicatus gen. et sp. nov., Satsangisaccites uruguaiensis sp. nov. and Platysaccus orientalis sp. nov. The
Downloaded By: [GutiérrezBuenos Aires, Pedro] At: 16:04 14 November 2010

palynomorphs found allow identification, in borehole 221, of two palynological assemblages: 221-I (140-225 m) and 221-II
(93-140 m); the former is characterised by the presence of Murospora torifera Ybert, Vallatisporites arcuatus Archangelsky &
Gamerro and Lundbladispora areolata Césari et al., whereas assemblage 221-II contains Corisaccites alutas Venkatachala &
Kar, Lueckisporites stenotaeniatus Menéndez, L. virkkiae Potonié & Klaus, L. angoulaensis Jardiné, Lunatisporites variesec-
tus Archangelsky & Gamerro, Striatoabieites anaverrucosus Archangelsky & Gamerro and Staurosaccites cordubensis
Archangelsky & Gamerro, among others. In borehole 201, two assemblages were also identified: 201-I (34 and 93 m) and
201-II (5 and 34 m). The former is typified by the presence of Cristatisporites chacoparanensis Ottone, C. lestai Archangel-
sky & Gamerro, Leiotriletes corius Kar & Bose, Murospora torifera, M. bicingulata Ybert, Vallatisporites arcuatus, V. russoi
Archangelsky & Gamerro, Illinites unicus Kosanke, Limitisporites hexagonalis Maheswari and Protohaploxypinus bhard-
wajii Foster among others. Assemblage 201-II is characterized by the presence of Brevitriletes levis (Balme & Hennelly)
Bharadwaj & Srivastava, Pakhapites fusus (Bose & Kar) Menéndez, P. ovatus (Bose & Kar) Garcı́a, Platysaccus papilio-
nis Potonié & Klaus, Protohaploxypinus limpidus (Balme & Hennelly) Balme & Playford, Striatopodocarpites cancellatus
(Balme & Hennelly) Hart and Polarisaccites bilateralis Ybert & Marques-Toigo. Based on the previous records of those taxa,
assemblages 221-I, 201-I and 201-II are assigned to the Early Permian (Asselian-Artinskian) and assemblage 221-II to the
upper part of the Early Permian (Artinskian-Kungurian). According to the stratigraphical schemes of Argentina and Brazil,
they are correlated with the Biozones Cristatisporites (assemblages 221-I, 201-I and 201-II) and Striatites (assemblage
221-II and El Barón) (Chacoparaná Basin), Fusacolpites fusus-Vittatina subsaccata (assemblages 221-I, 201-I and 201-II)
and Lueckisporites-Weylandites (assemblage 221-II and El Barón) (Paganzo Basin), and Vittatina costabilis (assemblages
221-I, 201-I and 201-II) and Lueckisporites virkkiae (assemblage 221-II and El Barón) (Paraná Basin).
Keywords: palynology; systematics; Lower Permian; Uruguay

Introduction holes, which led to the proposal of a palynostratigraphic

Neopalaeozoic sequences in Uruguay occupy an area of
94,000 km2. They are found in the northern region and
are part of the Paraná Basin (Fig. 1B). Outcrops spread
through 24,000 km2 in the Departments of Cerro Largo,
Tacuarembó, Rivera and Durazno (NE Uruguay). Else-
where they are found in the subsurface (Andreis et al. 1996).
This basin is continuous with the Chacoparanense Basin of
Argentina and the Paraná Basin of Brazil (Fig. 1A).
In the Chacoparanense Basin, palynomorph fossils from
Neopalaeozoic rocks are well known (Archangelsky &
Gamerro 1979; Vergel 1986, 1987b, 1998; Césari et al.
1995; Gutiérrez et al. 1997, 2002; Playford & Dino
2002). These previous authors studied palynological floral
assemblages of Pensylvannian-Permian age from bore-
scheme for the region (see Russo et al. 1980; Vergel 1993;
Césari et al. 1995; Archangelsky & Vergel 1996).
The Upper Palaeozoic strata of the Paraná Basin in Brazil
have yielded diverse and abundant fossils, including inver-
tebrates, vertebrates, plant remains and palynomorphs. The
palynomorphs from the Paraná Basin have been known
since the late 1960s, especially those from the coal beds of
the Rio Bonito Formation in Rio Grande do Sul and Santa
Catarina states. Papers that contribute to the taxonomy
and zonation schemes of this palaeoflora include Daemon
& Quadros (1970), Marques-Toigo (1991) and Souza &
Marques-Toigo (2001, 2003, 2005).
In Uruguay, the first palynomorphs from Neopalaeo-
zoic sediments were described by Martı́nez Macchiavello
(1963) from the San Gregorio and Tres Islas formations.

∗
Corresponding author. Email: prgutiérrez@macn.gov.ar

ISSN 1477-2019 print / 1478-0941 online

Copyright 
C 2010 The Natural History Museum
DOI: 10.1080/14772011003794546
http://www.informaworld.com
 460 P. R. Gutiérrez et al.

Figure 1. Palaeogeographical A, location of the study area;

geographical B, location of the boreholes DCLS 201
(DI.NA.MI.GE.) and DCLS 221 (DI.NA.MI.GE), and the locality
of El Barón.
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Afterwards, Marques-Toigo (1970, 1972, 1974) and Ybert

& Marques-Toigo (1971) extended the study of material
from the San Gregorio Formation. In 1980, palynological
studies started in Uruguay and a series of papers were subse-
quently published covering the systematics, biostratigraphy
and palaeoecology of the San Gregorio, Tres Islas and Melo
formations (Beri 1987, 1988, 2003; Beri & Daners 1995,
1998; Beri & Goso 1996, 1998; Beri et al. 2000, 2006; Beri
& Pecoits 2001; Gutiérrez et al. 2006). Finally, some publi-
cations by Argentina-based authors on Uruguayan material
have been carried out, including those of Vergel (1987a),
Anzótegui & Mautino (1992), Fasolo & Vergel (1994) and
Mautino et al. (1998a, b, c).
The present study provides systematic descriptions of
assemblages from boreholes 201 and 221, and an outcrop
at El Barón (Figs 1–4). This study complements the results
previously published by Beri & Daners (1995 [borehole
221], 1998 [borehole 201]) and Beri & Pecoits (2001 [El
Barón]).

The Paraná Basin

The existence of Neopalaeozoic rocks in Uruguay was

established by Guillemain (1911), Llambı́as de Olivar
(1918) and Walther (1919), while its presence in the
subsurface has been known since boreholes were drilled
by ANCAP in 1956 (Mackinon 1969). These sequences
occupy an area of 94,000 km2 in the northern region of
Uruguay and are part of the Paraná Basin (Fig. 1A). The
outcrops extend for 24,000 km2 in the departments of Cerro
Largo, Tacuarembó, Rivera and Durazno (Andreis et al.
1996). This basin is continuous with the Chacoparaná Basin
of Argentina and with the Paraná Basin of Brazil and has
Figure 2. Columnar stratigraphic section of the core DCLS
also been termed the Northern Basin (“Cuenca Norte”, de 201 (DI.NA.MI.GE.) showing lithology, stratigraphical units, and
Santa Ana et al. 2006). sampling levels.
 Lower Permian palynology of Paraná Basin
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Figure 3. Columnar stratigraphic section of the core DCLS
221 (DI.NA.MI.GE.) showing lithology, stratigraphical units, and
sampling levels.
461
Figure 4. Stratigraphical section at El Barón: showing lithology
and sampling levels.
According to Andreis et al. (1996), four formational units
are found here: the San Gregorio, Tres Islas, Melo and
Yaguarı́ formations. The San Gregorio Formation (Ferrando
& Andreis 1982) is composed mainly of diamictites, tillites
and, to a lesser extent, conglomerates, although there are
also claystones, siltstones and occasional sandstones. They
are predominantly reddish and brownish in colour, but there
are also greyish tones. According to Andreis et al. (1996),
these beds were deposited in glacial, glacio-marine and
glacio-lacustrine environments. De Santa Ana et al. (2006)
distinguished the organic-rich, fine grained sediments at
the top of the San Gregorio Formation as the Cerro Pelado
Formation.
The overlying unit, the Tres Islas Formation (Ferrando
& Andreis 1982), is composed of fine to very coarse
 462 P. R. Gutiérrez et al.
sandstones of mainly brownish colour, with a variable
percentage of usually grey claystones and siltstones (see
Fig. 2; Andreis et al. 1996). It was deposited in fluvial and
deltaic environments (Andreis et al. 1996).
The third unit, the Melo Formation (Ferrando & Andreis
1986), consists mainly of sandstones and to a lesser extent
of finer sediments, conglomerates and calcareous levels (see
Figs 2–4), mostly grey and black in colour (Andreis et al.
1996). Ferrando & Andreis (1986, 1990) suggested that the
Melo Formation was deposited in wide and shallow bays
with lagoons. Bossi & Navarro (1988) recognized three
members, from base to top, the Frayle Muerto, Mangrullo,
and Paso Aguiar members. Other authors (Preciozzi et al.
1985; de Santa Ana 2004; de Santa Ana et al. 2006) have
considered these members to have formational status.
The uppermost unit of this sequence, the Yaguarı́ Forma-
tion (Bossi 1966), conformably overlies the Melo Forma-
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tion (Ferrando & Andreis 1986), and is composed mainly
of sandstones and to a lesser extent claystones and silt-
stones, deposited under oxic conditions in a prograding
fluvial system (Bossi et al. 1998).
Material and methods
Eleven levels in the DCLS 201 (DI.NA.MI.GE.) borehole
were sampled. This well was drilled by the “Dirección
Nacional de Minerı́a y Geologı́a” (DI.NA.MI.GE.) and is
located in the northeast of the Departamento de Cerro Largo
in northeastern Uruguay (Fig. 1). The well penetrated 96 m
of Permian sediments, including 32 m of siltstones and
sandstones with argillaceous matrix, referred to the Frayle
Muerto Member (lower section of the Melo Formation; see
Fig. 2); and 64 m of sandstones, siltstones and claystones-
coals referred to the Tres Islas Formation.
Seventeen levels within the DCLS 221
(“DI.NA.MI.GE”) well were sampled. Located in the
same area as DCLS 201 (Fig. 1), this well penetrated
Permian sediments referred to the Melo Formation,
including 35 m of siltstones and sandstones with argilla-
ceous matrix (Paso Aguiar Member), 18 m of claystones
(Mangrullo Member), and 103 m of siltstones and sand-
stones (Frayle Muerto Member). The granitic Precambrian
basement was reached at a depth of 240 m (see Fig. 3).
Additional samples were collected from outcrops at
Cañada El Barón (31◦ 58’32”S, 54◦ 05’10”W) located close
to the two wells (Fig. 1).
Each sample was processed following standard palyno-
logical methods (Wood et al. 1996). Samples were prepared
in the Departamento de Paleontologı́a de la Facultad de
Ciencias laboratories in Montevideo, Uruguay. Photographs
were made with an Olympus X-55 optical light microscope.
Slides are housed in the Facultad de Ciencias, Paleopaly-
nology Collection (FCPP), Uruguay. Approximately 400
sporomorphs were counted from each sample; the rela-
tive proportions of each genus identified are specified by
the lowest and highest value and the arithmetic mean, e.g.
(55.3–96.1%; mean 79.3%).
Systematic descriptions
The majority of species encountered are simply listed. For
species previously described from the Permian of Uruguay,
see Marques-Toigo (1970, 1974), Ybert & Marques-Toigo
(1971), Beri (1988), Beri & Daners (1995, 1998), Mautino
et al. (1998a, b, c), Beri et al. (2006) and Gutiérrez et al.
(2006). Full descriptions are given only for species that are
new, particularly abundant, significant, or which taxonomic
or morphological clarification is deemed necessary.
The suprageneric classification of spore-pollen paly-
nomorphs largely follows Potonié & Kremp (1954), with
later modifications by Playford & Dettmann (1996).
Morphometric terminology for bisaccate and bilateral
symmetrical pollen grains follows Playford & Dino (2002,
p. 242).
Definitions of terms adopted in the description of
morphological attributes of spores and pollen grains are
to be found in glossaries or explanatory notes furnished
by several authors (see Playford & Dino 2000a, p. 14).
Measurements of equatorial diameters are presented in the
format N1 (N2) N3, where N1 and N3 represent the mini-
mum and the maximum value measured respectively, and
N2 the arithmetic mean for all the specimens evaluated. The
number of specimens measured appears between brackets.
Measurements quoted for saccate pollen grains are in accor-
dance with the conventions adopted by Couper (1958),
Balme (1963), Singh (1971) and Burden & Hills (1989)
(see Playford & Dino 2000a, 2002). Under Distribution
we note primarily only the material described for Uruguay,
Argentina and Brazil.
Anteturma Proximegerminantes Potonié, 1970
Turma Triletes Reinsch, emend. Dettmann, 1963
Suprasubturma Acavatitriletes Dettmann, 1963
Subturma Azonotriletes Luber, emend. Dettmann, 1963
Infraturma Apiculati Bennie & Kindstom, emend.
Potonié, 1965
Subinfraturma Nodati Dybová & Jachowicz, 1957
Genus Anapiculatisporites Potonié & Kremp, 1954
Anapiculatisporites sp. A
(Fig. 7J)
Occurrence. DCLS wells 201, 221.
Description. Spores radial, trilete; amb triangular. Leasure
apparently simple, extending to the apices. Proximal face
laevigate. Distal face ornamented with low coni. Coni 1 µm
wide at base, 1 µm height.
 Lower Permian palynology of Paraná Basin
Dimensions. (2 specimens). Equatorial diameter, 37–
43 µm; exine, 1–1.5 µm thick.
Remarks. The poor state of preservation of both speci-
mens does not allow a more precise assignment.
Subinfraturma Baculati Dybová & Jachowicz, 1957
Genus Microbaculispora Bharadwaj, 1962
Microbaculispora sp. A
(Fig. 8B)
1995 Granulatisporites austroamericanus Archangelsky &
Gamerro; Beri & Daners: pl. 1, fig. 5.
Occurrence. DCLS well 221.
Description. Spores radial, trilete; amb triangular with
rounded apices and straight to slighly convex sides. Laesura
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simple, length two-thirds of spore radius. Exine 1–1.5 µm
thick; ornamented with small bacula (0.5–1 µm diameter,
1–1.5 µm high), 1–2 µm apart.
Dimensions. (1 specimen). Equatorial diameter, 34 µm.
Remarks. Differentiated from Microbaculispora gondwa-
nensis Bharadwaj (1962, p. 81) and M. villosa (Balme
& Hennelly) Bharadwaj (1962, p. 81) by its diameter
and smaller dimensions at the sculpture. Microbaculispora
tentula Tiwari (1965, p. 175, figs 35–37) has similar dimen-
sions, but has equidimensional bacula.
Infraturma Murornati Potonié & Kremp, 1954
Genus Convolutispora Hoffmeister, Staplin & Malloy,
1955
Convolutispora archangelskyi Playford & Dino, 2002
(Fig. 7M)
1995 Verrucosisporites cf. V. microtuberosus (Loose) Smith
& Butterworth; Beri & Daners: pl. 1, fig. 3.
1995 Convolutispora sp.; Beri & Daners: pl. 1, fig. 6.
1995 Convolutispora sp.; Césari et al.: 81, pl. 2, fig. 16.
?1998a Microreticulatisporites inaequalis Menéndez &
Azcuy; Mautino et al.: 73, pl. 1, fig. 17.
2002 Convolutispora sp. Dino et al.: pl. 1, fig. 4.
(for further synonymy see Playford & Dino 2002, p. 254)
Occurrence. DCLS well 221, El Barón.
Description. Spore radial, trilete. Amb circular to subcir-
cular. Laesura perceptible to distinct, straight; length
varying from three-quarters of spore radius to near-
equal; laesura flanked by narrowly coalescent sculpturing
elements (rugula). Exine 3.5 µm thick, including compre-
hensively developed sculpture: irregularly branching, flat-
topped, sinuosous rugulae, with interspersed verrucate and
less-frequent granulose. Rugulate (1–2 µm wide, 1–1.5 µm
high) spacing variable (up to 1–1.5 µm).
463
Dimensions. (6 specimens). Equatorial diameter, includ-
ing sculpture, 42(50.8)58 µm.
Remarks. Convolutispora archangelskyi Playford & Dino
generally has a much thinner and less densely sculp-
tured exine with rugulate prevalence among the verru-
cate elements than that of C. ordonezii Archangelsky &
Gamerro, (1979). Despite this, and following Playford &
Dino (2002), we accept that there is morphological intergra-
dation between the two species. Microreticulatisporites
inaequalis Menéndez & Azcuy, described by Mautino et al.
(1998a, p. 73, pl. 1, fig. 17) from Permian strata of the
Paraná Basin, Uruguay (Melo Formation sensu Andreis
et al. 1996), is similar in appearance to C. archangelskyi,
but was described as having a reticulate ornamentation.
However, illustrations of this material show a rugulate exine
with anastomosing and sinuous pattern, different from the
type material (Menéndez & Azcuy 1973, p. 57, pl. 1, figs
3–6). We therefore provisionally assign it to C. archangel-
skyi. Likewise, C. pintoi Dellazana (1976, pp. 4–6, pl. 1, figs
4–6), should also be assigned to C. archangelskyi, because
of the similarities in ornamentation.
Previous records. Lower Permian of Argentina
(Archangelsky & Gamerro 1979; Césari et al. 1995;
Dino & Playford 2002), Brazil (Dino et al. 2002), and
Uruguay (Beri & Daners 1995; Mautino et al. 1998a; Beri
et al. 2006).
Convolutispora candiotensis Ybert, 1975
(Fig. 7O)
Occurrence. DCLS wells 201, 221.
Description. Spore radial, trilete. Amb subcircular to
convexly subtriangular. Laesura perceptible, simple,
straight; length varying from four-fifths of spore radius.
Exine 3.5 µm thick, comprehensively rugulate sculpture
of sinuous, narrow, usually anastomosing rugulae (2–3 µm
longitude), giving a reticulate appearance. Rugulae up to
6 µm long (usually about 2–4 µm) and varying in width
(1–2 µm) spacing variable (about 3–5 µm).
Dimensions. (2 specimens). Equatorial diameter,
40(42.3)47 µm.
Previous records. Lower Permian of Brazil (Ybert 1975;
Quadros et al. 1996; Smaniotto et al. 2006). This is the first
record of this species from Uruguay.
Subturma Zonotriletes Waltz, 1935
Infraturma Auriculati Schopf, emend. Dettmann, 1963
Genus Stenozonotriletes Naumova, emend. Potonié, 1958
Stenozonotriletes sp. A
(Fig. 8E)
 464 P. R. Gutiérrez et al.
Occurrence. DCLS well 201.
Description. Spore radial, trilete, cingulate, subcircular
to convexly subtriangular. Laesura perceptible, simple,
straight; length varying from four-fifths to one spore
radius. Cingulum 2.5–5 µm thick, smooth. Exine smooth
to scabrate.
Dimensions. (15 specimens). Equatorial diameter,
33(44.1)56 µm; cingulum, 2.5(3.4)5 µm wide.
Suprasubturma Laminanititriletes Smith & Butterworth,
1967
Subturma Zonolaminatitriletes Smith & Butterworth,
1967
Infraturma Cingulicavati Smith & Butterworth, 1967
Genus Cristatisporites Potonié & Kremp, 1954
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Cristatisporites rollerii Ottone, 1989
(Fig. 8K)
Occurrence. DCLS well 201.
Description. Spore trilete, cavati, cingulizonate; amb
subtriangular. Central corpus sub-triangular to subcircular
in outline. Laesura with thin lips, slighly sinuous; extend-
ing to inner margin of cingulum. Proximal surface laevigate.
Distal surface with coni (wider bases and acuminate tips),
mamoid elements, and spines. Sculptural elements discrete
at bases forming verruca muronate, crests and irregular
walls with aserrate surface; sporadically delimits areoles of
variable form and size. In the zona the sculpture merges
forming radial crests that diminish in size towards the
external margin, where they disappear. Cingulum thin (1–3
µm wide), with acuminate coni and mammoid elements.
Translucent, wide zona, ranging from one-third to one-half
of the radius of the spore.
Dimensions. (4 specimens). Equatorial diameter, 45–60
µm.
Previous records. Pennsylvanian-Lower Permian of
Argentina (see Césari & Gutiérrez 2001; Gutiérrez &
Barreda 2006); Pennsylvanian of Brazil (Souza et al.
2003). This is the first record of the species in Uruguay.
Genus Densosporites Berry, emend. Butterworth et al.,
1964
Densosporites sp. A
(Fig. 8I)
Occurrence. DCLS well 201.
Description. Spore radial, trilete, cingulate, cavate. Amb
subcircular. Laesura straight, surrounded by narrow, smooth
and low lips, 1.0–1.5 µm wide; extending to inner margin
of cingulum. Cingulum with irregular margin. Exoexine
distinctive, cingulate (6 µm wide). Distal surface with coni
mammoid, evident at cingulum; coni bases have circular
outline (4–6 µm wide). Intexine thin, up to 1 µm thick,
showing partial detachment from exoexine.
Dimensions. (1 specimen). Equatorial diameter, 45 µm;
corpus diameter, 33 µm.
Densosporites sp. B
(Fig. 8R)
Occurrence. DCLS well 201.
Description. Spore radial, trilete, cingulate, cavate. Amb
subtriangular to subcircular. Laesura straight acompanied
by narrow and low lips, 1–2 µm wide; extending to
inner margin of cingulum. Exoexine distinctive, cingulate
(4 µm wide). Distal surface with coni bases circular
(2–3 µm wide). Intexine thin showing partial detachment
from exoexine.
Dimensions. (2 specimens). Equatorial diameter, 30–45
µm; corpus diameter, 30–35 µm.
Genus Kraeuselisporites Leschick, emend. Jansonius,
1962
Kraeuselisporites punctatus Jansonius, 1962
(Fig. 8U)
1974 Kraeuselisporites punctatus Marques-Toigo: 605,
pl. 2, figs 1–2.
Occurrence. DCLS wells 201, 221.
Description. Spore radial trilete, zonate, subtriangular to
subcircular in outline. Laesura distinctive, sinuous with
narrow and low lips extending to the equatorial margin
of the intexine, individually 1 to 2 µm wide. Distal surface
of exoexine punctuate with occasional small granules or
verrucae towards equatorial margin. Proximal face laevi-
gate or punctuate. Zona is generally smaller than one-third
to one-quarter of spore radius, with irregular margin and a
thickening exinal dark at its base.
Dimensions. (12 specimens). Overall equatorial diame-
ter, 50(65)90 µm; corpus diameter, 40(51.2)78 µm; zona,
4(6.8)10 µm wide.
Remarks. The species differs from Kraeuselisporites
apiculatus Jansonius (1962, p. 47, pl. 11, fig. 26), K.
spinosus Jansonius (1962, p. 47, pl. 11, fig. 22), K. sommeri
Cauduro (1970, p. 11, figs 30–36), and K. noduspiniferus
Cauduro (1970, p. 11, figs 37–42) by the virtual absence of
conspicuous distal ornamentation. ‘Kraeuselisporites punc-
tatus’ (non Jansonius 1962, p. 48, pl. 11, figs 2, 20) proposed
by Marques-Toigo (1974, p. 605, pl. 2, figs 1, 2), from the
 Lower Permian palynology of Paraná Basin
San Gregorio Formation, differs from the species described
here only by the characteristics of the trilete mark (sinuous
and raised) lips and, therefore, is likely to be conspecific.
Furthermore, Marques Toigo’s species is a homonym of a
name previously validly published. Vallatisporites puncta-
tus (Marques-Toigo) Souza et al. (2003, p. 55, pl. 3, fig. 1)
and V. sp. A of Playford & Dino (2000a, p. 20, pl. 4, fig.
7) are distinguished by having a vacuole in the equatorial
zone.
Previous records. Lower Triassic of Argentina and
Canada (Jansonius 1962; Jain 1968); Upper Permian of
Russia (Mangerud 1994). Lower Permian of Uruguay
(Marques-Toigo 1974; Beri & Goso 1996).
Turma Monoletes Ibrahim, 1933
Suprasubturma Acavatomonoletes Dettmann, 1963
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Subturma Azonomonoletes Luber, 1935
Genus Reticuloidosporites Pflug in Thomson & Pflug,
1953
Remarks. In this paper we follow Dettmann (1963) and
Balme’s (1970) definition of Reticuloidosporites, which
includes spores with elongate sculptural elements (verruca,
rugula, crista), widely separated, that do not give rise to a
negative reticulum.
Reticuloidosporites sp. A
(Fig. 9A)
Occurrence. DCLS well 221.
Description. Bilateral dilete spore. Amb broadly oval in
polar view, periphery slightly undulating. Laesura extend-
ing about three-quarters spore length. Exine (2–3 µm thick)
sculptured proximally and distally with flat, low and irreg-
ular ridges with rounded crest, giving appearance of nega-
tive reticulum. Sculptural elements about 2–3 µm wide,
1–1.5 µm high, and separated by sinuous channels less than
1–2 µm wide.
Dimensions. (3 specimens). Length, 32(34)35 µm;
breadth, 28(29.3)30 µm.
Anteturma Variegerminantes Potonié, 1970
Turma Saccites Erdtman, 1947
Subturma Monosaccites Chitaley, emend. Potonié &
Kremp, 1954
Infraturma Dipolsacciti Hart, emend. Dibner, 1971
Subinfraturma Apertacorpini Dibner, 1971
Genus Tuberisaccites Lele & Makada, 1972
Tuberisaccites tuberculatus (Maheswari) Lele & Makada,
1972
(Fig. 9J)
1988 Polarisaccites sp. Beri: 37, fig. 22.
465
Occurrence. DCLS wells 201, 221.
Description. Pollen grain radial, monosaccate. Amb
subcircular, corpus outline subcircular. Corpus with one
irregular to lobate protuberance in pole (35 µm × 28 µm).
Saccus narrow (3 to 6 µm wide).
Dimensions. (3 specimens in polar view). Overall diame-
ter, 62–87 µm; corpus diameter, 49–72 µm.
Remarks. The material illustrated by Beri (1988, fig. 22)
shows the diagnostic characteristics of this species.
Previous records. Permian of India and Africa (Mahesh-
wari 1969; Lele & Makada 1972); Lower Permian of
Uruguay (Beri 1988).
Infraturma Striasacciti Bharadwaj, 1962
Genus Meristocorpus Playford & Dino, 2000a
Meristocorpus sp. A
(Fig. 10k)
Occurrence. DCLS wells 201 and 221.
Description. Pollen grains bilateral, monosaccate, taeni-
ate. Amb transversely oval. Corpus distinct, outline subcir-
cular to longitudinally oval; proximal face bearing 7 to 9
transverse taeniae, each 4–8 µm wide. A distinct, semilu-
nar, subparallel fold is developed marginally on corpus.
Dimensions. (4 specimen in polar view). Overall breadth,
95(113.8)130 µm; overall length, 70(81.7)100 µm; corpus
breadth, 50(60.8)73 µm; corpus length, 60(62.7)65 µm.
Subturma Disaccites Cookson, 1947
Infraturma Dissaccitrileti Leschik, 1955
Genus Archangelskiapollenites nov.
Type species. Archangelskiapollenites globocorpus sp.
nov.
Diagnosis. Characterized by its spherical corpus with
occasional exinal folds by compression, mark dilete, amb
diploxilonoid to slightly diploxilonoid, and cappula rectan-
gular to quadrangular.
Etymology. In honour of Dr Sergio Archangelsky, Argen-
tinian palaeobotanist and palynologist.
Remarks. Differentiation of pollen assigned to the new
genus Archangelskiapollenites is difficult because the
grains do not possess some of the distinct morpho-
logic features associated with other Neopalaeozoic bisac-
cate pollen grains. Archangelskiapollenites is characterized
by its spherical corpus with occasional exinal folds by
 466 P. R. Gutiérrez et al.
compression, mark dilete, amb diploxilonoide to slightly
haploxilonoid, and cappula rectangular to quadrangular.
Limitisporites Leschik is distinguishable mainly by its
sacci being distally attached associated with a semilunar
corpus infold system, and a corpus that is not spheri-
cal. Scopulisporites Leschik (1956, p. 64), Pinuspollenites
Raatz ex Potonié (1958, p. 62), and Piceapollenites Potonié
(1931, p. 28) are very similar to Archangelskiapollenites,
but do not have a monolete-dilete mark over proximal face
of corpus. Triadispora Klaus (1964, p. 120) has central
corpus folds (secondary structures) and a trilete mark, but
is protobisaccate. Pityosporites Seward emend. Manum
(1960, p. 14) has a very narrow cappula and no proximal
mark. Podocarpidites Cookson ex Couper (1953, p. 35)
has an exine corpus that is finely granular with a marginal
crest and has no proximal mark. Valiasaccites Bose &
Kar (1966, p. 43) has two lateral ridges (taeniae sensu
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Jansonius & Hills 1979, card 3153), a monolete mark, and
sacci distally attached associated with a semilunar corpus
infold system. Walikalesaccites Bose & Kar (1966, p. 102),
differs in being strongly diploxylonoid, with sacci distally
attached near the distal pole, slightly curved, corpus central
vertically oval and mark dilete generally extending over
the full width of the corpus. Labiisporites Leschik emend.
Klaus (1963) is haploxilonoid and has a monolete mark.
Archangelskiapollenites globocorpus sp. nov.
(Fig. 11A, C-E)
2002 Limitisporites sp. Dino et al.: fig. 20.
Holotype. FCPP 152(A) M28/1 (Fig. 11E).
Paratypes. FCPP 152(A) R35/3 (Fig. 11A), 155(B) M49/0
(Fig. 11C), and 155(B) V35/2 (Fig. 11D).
Type locality and stratum. DSCL well 221; Mangrullo
Member; Melo Formation.
Diagnosis. Pollen grain bisaccate, haploxylonoid to
diploxylonoid, equatorial outline oval to subquadrangular.
Central corpus subcircular to transversely oval in outline,
spherical with occasionally semilunar transversal folds
by compression. Cappa with dilete mark. Sacci slightly
distally inclined, equally to slightly higher than corpus.
Sacci subcircular crescentic in outline, inflate, discrete, and
infrareticulate. Proximal attached sacci are equatorial to
subequatorial; distal attachment sacci are subequatorial and
straight. Cappula slightly narrow, rectangular.
Etymology. Latin, globus: ball, referring to spherical
shape of the corpus.
Occurrence. DCLS well 201, 221.
Description. Cappa with dilete mark extended, one-half to
one-fifth corpus breath, ca, one-third. Cappula with sides
parallel or expanded at lateral extremes; extending over
full length of corpus, breadth one-half to one-fifth corpus
breath, ca, one-third.
Dimensions. (35 specimens in polar view). Over-
all breadth, 62.5(81.1)102.5 µm; corpus breadth
32.5(52.3)70 µm, corpus length, 33.6(49.1)65.5 µm; ratio
corpus breadth:corpus length, 0.80(1.08)1.27; proximal
saccus breadth, 10(13.9)20 µm, distal saccus breadth,
22.5(29.7)42.5 µm, saccus length, 40(49.5)65.5 µm;
cappula breadth, 5(19.7)32.5 µm; dilete, 9(20.1)30 µm.
Remarks. Archangelskiapollenites globocorpus sp. nov. is
characterized by its globosus corpus, dilete mark, inflated
and discrete sacci, outline subquadrangular to oval, and amb
haploxylonoid to diploxylonoid. The material illustrated by
Dino et al. (2002, fig. 20) from the Permian of Brazil is
included in this species based on its general outline and
characteristics of the corpus and cappula.
Archangelskiapollenites plicatus sp. nov.
(Fig. 11F-G, I)
1995 Limitisporites sp. 3 Beri & Daners: pl. 2, fig. 2.
Holotype. FCPP 152(A) H35/1 (Fig. 11F).
Paratypes. FCPP 150(A) D42/0 (Fig. 11I), and 152(A)
O32/4 (Fig. 11G).
Type locality and stratum. DCLS well 221; Mangrullo
Member; Melo Formation.
Diagnosis. Pollen grain bisaccate, diploxylonoid, oval in
outline. Central corpus transversely oval in outline, spher-
ical, with semilunar transversal and longitudinal folds by
compression in both faces. Cappa with dilete mark. Sacci
subcircular crescentic in outline, inflated, discrete, slightly
distally inclined, and much higher than corpus. Proximal
attached sacci are equatorial to subequatorial, associated
with exinal folds, sometimes; distal attachment sacci are
subequatorial and straight, located near the central distal
face. Cappula rectangular extending over full length of
corpus.
Etymology. Latin, plicatus: folds, referring to presence of
folds in the corpus.
Occurrence. DCLS well 221, El Barón.
Description. Cappa with dilete mark extending one-half
to one-fifth corpus breath, ca one-third. Cappula slightly
narrow (ca one-half corpus breadth), rectangular with sides
parallel or expanded at lateral extremes; extending over full
 Lower Permian palynology of Paraná Basin
length of corpus, breadth of cappula is equal to one-quarter
to one-half corpus breath.
Dimensions. (34 specimens in polar view). Overall
breadth, 67.5(87.0)107.5 µm; corpus breadth 40(55.3)87.5
µm, corpus length, 32.5(47.2)62.5 µm; ratio corpus
breadth:corpus length, 0.83(1.18)1.79; proximal saccus
breadth, 12.5(17.1)30 µm, distal saccus breadth,
22.5(33.7)50 µm, saccus length, 32.5(54.5)70 µm; cappula
breadth, 10(19.5)30. µm; dilete, 12.5(19.8)30 µm. Ratio
cappula breadth:corpus breadth, 1:4 to 1:2.
Remarks. Archangelskiapollenites plicatus is character-
ized by its spherical corpus with exinal folds, dilete mark,
inflated and discrete sacci, outline oval, and amb diploxy-
lonoid. These characteristics are seen in the material illus-
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trated by Beri & Daners (1995, pl. 2, fig 2) as Limitisporites
sp. 3 from the Permian of Uruguay which is therefore
included in this species.
Genus Limitisporites Leschik, emend. Schaarschmidt,
1963
Remarks. In this paper, we follow Potonié (1958) and
Schaarschmidt (1963) in treating Jugasporites as a junior
synonym of Limitisporites.
Limitisporites amazonensis Playford & Dino, 2000a
(Fig. 10I)
Occurrence. DCLS well 221.
Description. Pollen grain bisaccate, dilete, lightly diploxy-
lonoid to haploxylonoid, with overall outline transversely
oval. Corpus distinctive, longitudinally to transversely
oval in outline. Mark distinctive, length 0.3–0.5 corpus
breadth. Sacci hemispherical, discrete, distal sacci attach-
ments comprising two well-defined longitudinal crescentic
folds (5–9 µm wide; ca 6.7), delimiting cappula 10–38 µm
wide (0.3–0.7 corpus breadth; ca 0.5). Sacci opposite, equal
or smaller than corpus.
Dimensions. (19 specimens in polar view). Overall
breadth, 69(84.7)105 µm; corpus breadth, 32(47.9)63
µm; corpus length, 45(51.4)60 µm; ratio corpus
breadth:corpus length, 0.61(0.93)1.22; proximal saccus
breadth, 25(33.5)47.5 µm; distal saccus breadth,
16(21.0)32.5 µm; saccus length, 46.3(54.6)65 µm.
Remarks. Our material is slightly more variable in corpus
shape and not so strongly diploxylonoid as typical Limi-
tisporites amazonensis (cf. Playford & Dino 2000a, p. 102,
pl. 5, figs 7–9).
467
Previous records. Pennsylvanian of Brazil (Playford &
Dino 2000a). This is the first record of this species in
Uruguay.
Limitisporites delasaucei (Potonié & Klaus)
Schaarschmidt, 1963
(Fig. 10G)
Occurrence. DCLS well 221.
Description. Pollen grain bisaccate diploxylonoid, over-
all outline transversely oval. Corpus distinctive, longitudi-
nally oval in outline, with broadly rounded ends. Mark on
proximal surface (monolete to dilete). Sacci hemispherical,
discrete, without lateral exoexinal bands; distal sacci attach-
ments comprising two well-defined longitudinal crescentic
folds (8 µm wide), delimiting cappula 6–25 µm wide (one-
half to one-fifth-corpus breadth). Sacci opposite, equal to
larger than corpus.
Dimensions. (2 specimens in polar view). Overall breadth,
72–85 µm; corpus breadth, 30–34 µm; corpus length,
42–55 µm; ratio corpus breadth:corpus length, 0.55–0.81;
proximal saccus breadth, 29–40 µm; distal saccus breadth,
23–28 µm; saccus length, 43–50 µm.
Previous records. Known widely, especially from the
Permian of Gondwanaland and Europe; also from the
Permian of Brazil (Dino et al. 2002). This is the first record
from Uruguay.
Limitisporites sp. cf. L. luandensis Bose & Maheswari,
1968
(Fig. 10O)
Occurrence. DCLS well 221.
Description. Pollen grain bisaccate dilete, diploxylonoid.
Corpus distinctive, subcircular to transversally elongate.
Sacci hemispherical, discrete; distal sacci attachments
comprising two well-defined longitudinal crescentic folds
(6–13 µm wide), delimiting cappula 13–25 µm wide
(0.6–0.3 corpus breadth). Sacci inflated, opposite, equal
to or larger than corpus.
Dimensions. (6 specimens in polar view). Overall breadth,
80(82.9)93 µm; corpus breadth, 40(42.8)48 µm; corpus
length, 30(41.7)50 µm; ratio corpus breadth:corpus length,
0.88(1.05)1.23; proximal saccus breadth, 28(35.4)40 µm;
distal saccus breadth, 15(23.8)32.5 µm; saccus length,
45(53.5)60 µm.
Remarks. Limitisporites luandensis Bose & Maheswari
(1968, p. 63) differs in being larger (overall breadth,
112–162 µm), but is otherwise identical.
Infraturma Disacciatrileti Leschik, emend. Potonié, 1958
Genus Alisporites Daugherty, emend. Jansonius, 1971
 468 P. R. Gutiérrez et al.
Remarks. Balme (1970) separated Alisporites and Falcis-
porites Leschik, emend. Klaus, 1963, on the presence of
a longitudinal sulcus in Falcisporites, a feature which
he did not observe in Alisporites. Jansonius (1971) re-
examined Daugherty’s type material, and concluded that
Alisporites includes bisaccate, non-taeniate and distally
sulcate pollen grains. Falcisporites is also distally sulcate
but can be distinguished from Alisporites by the saccus,
which has a much coarser reticulation, and the ‘distal
leptoma’ (cappula?) is delineated by distinct distal bladder
bases and a differential expansion of protosaccate exoexine
over the cappa along the longitudinal axis (Jansonius 1971,
p. 356). Satsangisaccites Bharadwaj & Srivastava (1969)
can be differentiated from Alisporites by the presence of a
narrow sulcus and sacci attachment being laterally contin-
uous.
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Alisporites sp. cf. A. opii Daugherty, emend. Jansonius,
1971
(Fig. 11M)
Occurrence. DCLS well 221.
Description. Pollen grain bisaccate, haploxylonoid to
slightly diploxylonoid, overall outline transversally oval.
Corpus distinct oval, longitudinally elongated. Sacci attach-
ments comprising to defined longitudinal crescentic folds
delimiting cappula (13–16 µm wide) and cappa.
Dimensions. (6 specimens). Overall breadth, 45(64.0)74
µm; corpus breadth 28(39.5)46 µm, corpus length,
43(53.0)62 µm; ratio corpus breadth:corpus length,
0.61(0.7)0.87; proximal saccus breadth, 14(14.7)15 µm,
distal saccus breadth, 26(26.5)27 µm; saccus length,
40(50.7)72 µm.
Remarks. The present material is smaller than Alisporites
opii Daugherty, emend. Jansonius, and includes diploxy-
lonoid specimens.
Alisporites rioclarensis Menéndez, 1976
(Fig. 11J)
1996 Alisporites australis de Jersey; Césari et al.: 54 (pars),
pl. 1, fig. 1.
Occurrence. DCLS well 221, El Barón.
Description. Pollen grain bisaccate, haploxylonoid with
transversally subrectangular outline. Corpus outline
discernible, typically sub-quadrate. Exoexine of corpus
thick, cappa poorly defined. Cappula exine thick, with
narrow sulcus poorly defined. Sacci opposite, semicircu-
lar in outline, discrete, slightly smaller than corpus. Sacci
attached equatorially to corpus. Exoexine of sacci finely
infrareticulate.
Dimensions. (13 specimens, in polar view). Overall
breadth, 54(70)93 µm; corpus breadth 30(41.2)61 µm,
corpus length, 35(41.4)53 µm; ratio corpus breadth:corpus
length, 0.79(1)1.15; proximal saccus breadth, 9(16.7)28
µm, distal saccus breadth, 20(28.6)37 µm, saccus height,
27(34.2)42 µm.
Remarks. Part of the material described as Alisporites
australis de Jersey, coming from the Yacimiento Los
Reyunos Formation (Césari et al. 1996, p. 54, pl. 1, fig.
1), could be referred to this species on the basis of sacci
and body shape. The central body shape, sacci insertion and
their features differentiate this form from known species of
Alisporites. We leave this species unnamed until further
samples are available, and a more detailed characterization
can be made.
Previous records. Lower Permian of Brazil (Menéndez
1976) and Argentina (Césari et al. 1996). This is its first
record in Uruguay.
Alisporites similis (Balme) Dettmann, 1963
(Fig. 11H)
1994 Alisporites australis de Jersey; Fasolo & Vergel: pl.
3, fig. 1.
1995 Alisporites australis de Jersey; Césari et al.: 54 (pars),
pl. 1, fig. 4.
1996 Alisporites sp. Césari et al.: pl. 1, fig. 7.
Occurrence. DCLS well 221, El Barón.
Description. Pollen grain bisaccate, haploxylonoid, amb
transversally oval. Corpus longitudinally oval. Sacci distally
inclined, cresentic in outline, much smaller than corpus.
Cappa oval in outline with marginal crest, which is devel-
oped between the proximal saccus bases and the cappa.
Cappula lightly convex-sided, extending full length of the
corpus, with narrow sulcus (2–5 µm wide).
Dimensions. (8 specimens). Overall breadth, 44(56.1)84
µm; corpus breadth, 22(31.2)47.5 µm, corpus length,
32(43.5)63 µm; ratio corpus breadth:corpus length,
0.58(0.7)0.85; distal saccus breadth, 10(14.9)17.5 µm;
proximal saccus breadth, 20(25.3)28 µm.
Remarks. Alisporites similis (Balme) Dettmann resembles
A. haradensis Kumar (1973, p. 117, figs 115–116) in shape
but the latter is larger, has a fusiform sulcus, and lacks a
thickened exine. Vitreisporites itunensis Pocock (1970b, p.
86, pl. 18, figs 12, 14, 24) is very similar to A. similis, but
is larger. The material from the Melo Formation assigned
to A. australis de Jersey by Fasolo & Vergel (1994) might
be better included in A. similis based on body and sacci
shape, sacci insertion, thicker exinal, etc. Part of the mate-
rial described as A. australis de Jersey, coming from the
 Lower Permian palynology of Paraná Basin
Yacimiento Los Reyunos Formation (Césari et al. 1996, p.
54, pl. 1, fig. 4), could be referred to this species.
Previous records. Triassic-Cretaceous of Gondwanaland
(see Raine et al. 2005). Lower Permian of Uruguay (Fasolo
& Vergel 1994; Beri & Pecoits 2001).
Alisporites sp. A
(Fig. 10M)
Occurrence. DCLS well 221.
Description. Pollen grain bisaccate, diploxylonoid, over-
all outline transversally oval to sub-rectangular. Corpus
distinctive, longitudinally oval in outline. Longitudinal
sulcus poorly defined, extending to lateral margins of
corpus. Cappa delimited by slightly convex equatorial sacci
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attachment. Sacci crescentic, discrete, opposite or with
slight distal inclination, equal or larger than corpus.
Dimensions. (6 specimens in polar view). Overall breadth,
47(53)59 µm; corpus breadth, 34(37.6)45 µm; corpus
length, 28(36.2)46.5 µm; ratio corpus breadth:corpus
length, 0.93(1.05)1.23; proximal saccus breadth, 6.5(8)10
µm; distal saccus breadth, 20(23.5)29 µm; saccus length,
32(38)47.5 µm; sulcus breadth, 6(7.2)8 µm.
Remarks. Alisporites sp. A is distinguished from other
Alisporites species by the shape and size of its corpus and
grain, the ratio of sulcus breadth:corpus breadth, and the
sacci being opposite or with a slight distal inclination.
Genus Chordasporites Klaus, 1960
Chordasporites sp. cf. C. australiensis de Jersey, 1963
(Fig. 12D)
Occurrence. DCLS wells 221, 201, El Barón.
Description. Pollen grain bisaccate, slighly diploxylonoid,
overall outline transversally oval. Corpus distinctive,
subcircular in outline. Chorda (3–4 µm wide), with non-
sinuous margins, extending total breadth, or almost total
breadth of corpus. Sacci crescentic, opposite or with slight
distal inclination. Sacci with semicircular outline, approxi-
mately equal to two-thirds body in size.
Dimensions. (11 specimens in polar view). Overall
breadth, 73(96.4)135 µm; saccus length, 40(52.8)73 µm;
corpus breadth, 40(55.2)70 µm; corpus length, 40(55.0)75
µm; ratio corpus breadth:corpus length, 0.93(1.0)1.2; distal
saccus breadth, 26(38)50 µm; proximal saccus breadth,
20(25)30 µm.
Remarks. Chordasporites australiensis de Jersey differs
in having larger sacci, wider chorda, and a slightly smaller
size range. Our specimens are similar to those described by
469
Zavattieri (1991a, p. 20, pl. 7, fig. 7 as C. cf. australiensis)
from the Triassic of Argentina, but differ in the corpus being
longitudinally oval. C.? sp. (Gutiérrez et al. 2006) from the
San Gregorio Formation differs in being smaller (overall
breadth, 50–55 µm).
Genus Falcisporites Leschik, emend. Klaus, 1963
Falcisporites nuthallensis (Clarke) Balme, 1970
(Figs 11L, 12A)
1976 Alisporites indarraensis Segroves; Dellazana: 10, pl.
3, figs 6–8.
1976 Sulcatisporites ovatus (Balme & Hennelly)
Menéndez: 8, pl. 1, fig. 11.
2001 ?Alisporites australis de Jersey; Beri & Pecoits: fig.
3D.
(for full synonymy see Balme 1970, p. 389)
Occurrence. DCLS well 221, El Barón.
Description. Pollen grains bisaccate, haploxylonoid, over-
all outline transversally oval. Corpus distinct, oval, longitu-
dinally elongated, and with a thin marginal border. Cappa
exoexine thin, delimited by slightly convex saccus attach-
ment, and by associated folds. Cappa wide, about two-thirds
of corpus width. Sulcus very narrow, with parallel sides
and thin exoexine, defined by sacci straight distal attach-
ment. Sacci discrete, semicircular in outline, opposite or
with slight distal inclination, equal or smaller than corpus;
finely endoreticulate; attached saccus sub-equatorially to
proximal face.
Dimensions. (32 specimens in polar view). Overall
breadth, 40(66.1)94 µm; corpus breadth, 21(35.2)61 µm;
corpus length, 26(42.9)62 µm; ratio corpus breadth:corpus
length, 0.54(0.8)1.1; proximal saccus breadth, 6(17.3)38
µm; distal saccus breadth, 16(31.2)50 µm; saccus
length, 24(38)64 µm; cappa breadth, 10(23.5)41 µm;
sulcus breadth, 1(2.5)5 µm; ratio sulcus:corpus breadth,
1/45(1/20)1/8. Saccus heigth, 24(37)64 µm. Ratio saccus
height:corpus length, 0.65(0.80)1.05.
Remarks. Falcisporites nuthallensis (Clarke) Balme is
distinguished from Alisporites by its narrow sulcus, high
ratio sulcus:corpus breadth, corpus with thin marginal
border, and sacci opposite or with slight distal inclination
(Clarke 1965, p. 346, pl. 43, figs 1, 15; text-fig. 15; Balme
1970, p. 389, figs 15–18). F. nuthallensis (Clarke) Balme,
from the Triassic of Argentina (Zavattieri & Volkheimer
1992, p. 32, pl. 2, figs 5, 7), and F. stabilis Balme (1970, p.
387, pl. 16, figs 6–10), from the Permo-Triassic of Iran, are
distinguished by their greater sulcus:corpus breadth ratio
(ca 1:5, and 1:2–1:3 respectively). A. splendens (Leschik)
Foster (1979, p. 73, pl. 25, figs 9, 10) differs in having an
inconspicuous sulcus. Scheuringipollenites ovatus (Balme
& Hennelly) Foster (1975, p. 19, pl. 6, figs 5–6) is
 470 P. R. Gutiérrez et al.
distinguished by its poorly defined corpus and semi-
elliptical sacci in polar view. A. indarraensis Segroves,
described by Dellazana (1976, p. 10, pl. 3, figs 6–8), is
conspecific with the material described here. In A. indar-
raensis the corpus is circular to subcircular in polar view,
and the sacci, usually inflated, having strong distal inclina-
tion (Segroves 1969, p. 191, pl. 6, figs A-E). Sulcatisporites
ovatus, described by Menéndez (1976, p. 89, pl. 1, fig.
11), probably should also be assigned to F. nuthallensis.
Alisporites cf. A. splendens (Leschik) Foster, described by
Césari et al. (1996, p. 54, pl. 1, fig. 6), is very similar to
the Uruguayan material, nevertheless it is larger and exinal
folds on the proximal face and sulcus cannot be clearly
seen.
Previous records. Permian-Triassic of Gondwanaland
and Europe (see Balme 1970; Zavattieri 1992). Triassic
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of Argentina and Chile (Zavattieri 1987, 1991a, 1992;
Zavattieri & Volkheimer 1992; Zavattieri et al. 2003);
Permian of Brazil (Daemón & Quadros 1970; Dellazana
1976); Lower Permian of Uruguay (Beri & Pecoits 2001).
Genus Phrixipollenites Haskell, 1968
Phrixipollenites sp. A
(Fig. 12B-C, F)
Occurrence. DCLS well 221, El Barón.
Description. Pollen grain bisaccate. Corpus subcircular
in polar view, biconvex in equatorial view. Cappa thick
(2.0–3.5 µm thick), scabrate, breadth about that of corpus.
Cappula thinner than cappa. Sacci smaller than the
corpus, strongly pendant distally. Sacci inflated, round to
semicircular, irregularly shallow and infrareticulate with
radial arrangement.
Dimensions. (7 specimens in polar view). Overall breadth,
56(53)64 µm; corpus breadth 37(38.2)40 µm, corpus
length, 31(36.4)41 µm; ratio corpus breadth:corpus length,
0.93(1.05)1.20; maximum saccus breadth, 22–23 µm,
saccus length, 29(34.7)38 µm.
Remarks. Pollen grains resembling Phrixipollenites sp. A
are uncommon in Palaeozoic sediments. P. sp. from Trias-
sic of Argentina (Zavattieri & Volkheimer 1992, p. 38,
pl. 6, fig. 8, pl. 5, fig. 3) is distinguished by its trans-
versely oval corpus. Pityosporites communis Tschudy &
Kosanke (1966, p. 66, pl. 2, figs 32–33) from the Permian
of Texas appears similar to Phrixipollenites sp. A, but
has a less extensive cappa which is thinner and granulate,
bigger sacci, and an oval corpus. Some of the specimens
assigned to Pinuspollenites thoracatus by Balme (1970,
p. 400, fig. 11, pl. 15, figs 10–14) bear a general resem-
blance to Phrixipollenites sp. A, but differ in having wider
cappula and sacci abruptly differentiated from the corpus,
which are strongly inclinated distally and with a wider
cappula.
Genus Platysaccus Naumova ex Potonié & Kremp, 1954
Platysaccus orientales sp. nov.
(Fig. 12L, N)
Holotype. FCPP 155(B) C38/1 (Fig. 12N).
Paratype. FCPP 155(B) V44/0 (Fig. 12L).
Type locality and stratum. DCLS well 221; Mangrullo
Member, Melo Formation.
Diagnosis. Pollen grain bisaccate, amb strongly diploxy-
lonoid. Corpus well defined, subcircular to slightly longitu-
dinally oval. Cappula narrow, rectangular, margins paral-
lel, coinciding with total distal roots of sacci; extend-
ing over full length of corpus, breadth one-fifth that of
corpus. Sacci reniform in shape, distinctly inflated, discrete,
and much greater than semicircular in equatorial outline.
Saccus infrastructure fine to moderately coarse. Root sacci
straight.
Etymology. Due to its geographic provenance.
Occurrence. DCLS well 221.
Dimensions. (7 specimens in polar view). Overall breadth,
65(78.5)83 µm; corpus breadth 36(43.3)55 µm, corpus
length, 43(47.5)55 µm; ratio corpus breadth:corpus length,
0.80(0.91)1; proximal saccus breadth, 36(37.5)39 µm,
saccus length, 50(56.5)65 µm.
Remarks and comparison. Platysaccus orientalis sp. nov.
is characterized by its rectangular cappula, inflated, reni-
form and discrete sacci with straight root sacci, and amb
diploxylonoid. P. cacheutensis Jain (1968, p. 28, pl. 8, figs
105–107) differs from P. orientalis in having a distinctly
longitudinal elongate corpus and larger sacci. Specimens of
P. queenslandii de Jersey described by Jain (1968, p. 27, pl.
8, figs 102–104) appear distinguishable from P. orientalis
only by their root sacci and total amb. The cf. P. triassicus
(Mädler, 1964) from the Triassic of China (Ouyang Shu &
Norris 1999, p. 41, pl. 6, fig. 6) is very similar to P. orien-
talis. However, the lack of description prohibits detailed
comparison. Besides, P. triassicus (Maljavkina) Dunay &
Fisher (1979, p. 86, pl. 5, figs 14–15) is smaller, has a
narrower cappula with concave sides, longitudinally ellipti-
cal corpus and which is smaller than the sacci. P. katriensis
Kar (1968, p. 126, pl. 2, figs 52, 58) from the Permian of
India is slightly smaller and has a denser walled, vertically
elliptical corpus. P. queenslandii de Jersey, from the Triassic
of Argentina (Zavattieri 1992, p. 20, pl. 3, fig. 7), is similar
to P. orientalis but has exinal plicae associated with the
 Lower Permian palynology of Paraná Basin
distal insertion of the sacci, cappula with concave sides and
exinal bridges. Podocarpidites nous Sah & Jain (1965, p.
278, pl. 6, figs 113–114), from the Jurassic of India, shows
a superficial resemblance to Platysaccus orientalis, but is
slightly smaller and has a conspicuous marginal rim in the
central corpus. The same is true for Podocarpidites cf. P.
multesimus (Bolkhovitina) Pocock illustrated by Dettmann
(1963, p. 103, pl. 25, figs 13–14 only), but the reticulation
of the sacci is subdued and seems to show granulation in
the central corpus. Podocarpites arquatus (Kara-Muza in
Bolkhovitina) Haskell (Zavattieri 1992, p. 16, pl. 8, fig. 24)
is separated by the presence of cappula with concave sides,
sacci smaller than the body and a smaller overall size.
Platysaccus sp. A
(Fig. 12E)
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Occurrence. DCLS well 221.
Description. Pollen grain bisaccate, amb diploxylonoid,
shortly oval transversely. Corpus well defined, longitudi-
nally subhexagonal, and with thicker exine (3 µm wide).
Cappula narrow, rectangular, margins sub-parallel; extend-
ing over full length of corpus; breadth one-fifth that of
corpus. Sacci short and reniform, discrete, and semicircu-
lar in shape.
Dimensions. (1 specimen in polar view). Overall breadth,
43 µm; corpus breadth 24 µm, corpus length, 35 µm;
ratio corpus breadth:corpus length, 0.69; proximal saccus
breadth, 23 µm, saccus length, 47 µm.
Remarks. Platysaccus sp. A differs from other species of
Platysaccus in having a distinctly longitudinally elongate,
subhexagonal body, smaller sacci and a thicker exine of
the corpus. Triadispora suspecta Scheuring from Triassic
of France (Taugourdeau-Lantz 1974, pl. 2, fig. 4) is very
similar to P. sp. A, but lack of description prevents a more
precise comparison.
Genus Podocarpidites Cookson ex Couper, 1953
Podocarpidites sp. A
(Fig. 12J)
Occurrence. El Barón.
Description. Pollen grain bisaccate, amb slightly diploxy-
lonoid, oval transversely. Corpus dark in colour, well
defined, transversely elongate, and with thick exine
(1–2 µm wide). Cappa verrucose, verruca low and vermicu-
late in pattern. Cappula rectangular with margins subparal-
lel; extending over full length of corpus, breadth about half
that of corpus. Sacci smaller that corpus, discrete, semicir-
cular to weakly crescentic in polar view.
471
Dimensions. (1 specimen in polar view). Overall breadth,
72 µm; corpus breadth, 48 µm, corpus length, 38 µm;
saccus length, 50 µm.
Remarks. The species that comes closest to Podocar-
pidites sp. A is P. vermiculatus Kumar (1973, p. 119, figs
126–127), but this species differs in having a narrower
corpus and a roundly oval to subcircular amb.
Genus Pteruchipollenites Couper, 1958
Pteruchipollenites sp. cf. P. gondwanensis (Jain) Ottone
& Garcı́a, 1991
(Fig. 13A)
Occurrence. DCLS well 221.
Description. Pollen grain bisaccate, amb diploxylonoid,
non sulcate. Corpus subcircular to suboval in shape, verti-
cally elongate. Sacci about semicircular in shape, with bases
undifferentiated, gradually merging with corpus exine.
Sacci slightly inclined distally, finely reticulate.
Dimensions. (2 specimens in polar view). Overall breadth,
75–110 µm; corpus breadth, 53–70 µm; corpus length,
55–82 µm; cappula breadth, 15–27 µm; saccus length,
55–76 µm.
Remarks. The present material differs from the original
species description (Jain 1968, p. 21, pl. 4, figs 59–61)
mainly in the small sacci relative to the size of corpus.
Pterucipollenites gondwanesis (Jain) Ottone & Garcı́a
(Ottone et al. 1992, pl. 5, fig. 6) bears a strong resemblance
to this material.
Previous records. Triassic of Argentina (Ottone & Garcı́a
1991; Ottone et al. 1992).
Genus Satsangisaccites Bharadwaj & Srivastava, 1969
Satsangisaccites uruguaiensis sp. nov.
(Fig. 13B-C, E, H-I)
Holotype. FCPP 152(A) O32/0 (Fig. 13B).
Paratypes. FCPP 152(A) J36/2 (Fig. 13E), G55/1 (Fig.
13H), M44/4 (Fig. 13C); and 150(A) D43/1 (Pl. 7, Fig.
13I), C32/0.
Type locality and stratum. DCLS well 221 (120.5 m);
Melo Formation.
Diagnosis. Pollen grains bisaccate, diploxylonoid, corpus
subcircular or oval with a thin equatorial thickness. Sacci
laterally connected by exoexinal bands. Proximal sacci
attachment are subequatorial or equatorial, associated with
two crescentic, lateral exinal folds connected at the end
of the corpus. Distally, the sacci are inclined and attached,
 472 P. R. Gutiérrez et al.
sacci delimits sharply leaving a narrow, longitudinally elon-
gated sulcus; rarely with median groove. Sacci larger than
corpus, and distinctly greater than semicircular in equato-
rial outline.
Occurrence. DCLS well 221, El Barón.
Description. Pollen grain bisaccate, amb diploxylonoid,
overall outline, transversely oval. Corpus perceptible,
outline subcircular to longitudinally or transversely oval.
Corpus outline is marked by the presence of a thickened
wall along its equator. Corpus exine is thick, laevigate
and intramicroreticulate. Sacci are laterally connected by
distinct exoexinal bands (1–5 µm, ca 2.6 µm wide); prox-
imal sacci attachment is subequatorial or equatorial, asso-
ciated with two crescentic, lateral exinal folds (3–6 µm
wide), connected to the end of corpus; distally the sacci
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are inclined and attachment of the sacci leaving a narrow,
longitudinally elongated sulcus (sulcus wide, 2–5 µm, ca
3.7 µm; ratio sulcus wide:corpus breadth, 1:17–1:22, ca
1:11); rarely a median groove may be clearly seen. Sacci
larger than corpus (saccus breadth:corpus breadth, 1.0–1.2;
saccus length:corpus length, 1.0–1.6), and distinctly greater
than semicircular in equatorial outline.
Dimensions. (21 specimens in polar view). Overall
breadth, 59(84.0)98 µm; corpus breadth 26(40.6)52
µm, corpus length, 25(42.7)53 µm; ratio corpus
breadth:corpus length, 0.80(0.96)1.16; proximal saccus
breadth, 28(38.6)53 µm, distal saccus breadth, 18(25.1)35
µm, saccus length, 34(49.9)59 µm.
Comparison. Pityosporites nucleoparvus Anderson
(1977, p. 112, pl. 138, figs 1–2) is known only from its
brief diagnosis and its illustrations, thus precluding a
detailed comparison. It possibly differs from the present
species in having neither a narrow sulcus nor an exinal
band between the sacci. Satsangisaccites nidpurensis
Bharadwaj & Srivastava (1969, p. 132, pl. 27, figs 50–60)
is consistently larger than S. uruguaiensis (overall breadth,
107.5–152.5 µm), is haploxylonoid in outline, and has a
corpus that is rhomboidal or broadly vertically oval, with
its outline diffuse, only marked by the presence of folds
along its equator. S. triassicus Bharadwaj & Srivastava
(1969, p. 132, pl. 27, figs 61–66) is distinguished by its
transversely oval corpus, its diffuse haploxylonoid outline,
and its smaller size (overall breadth, 42.5–75 µm).
Satsangisaccites sp. A
(Fig. 13D)
Occurrence. El Barón.
Description. Pollen grain bisaccate, amb haploxylonoid,
overall outline transversely oval. Corpus transversely oval
in outline, thickened along its equator. Sacci are later-
ally connected by exoexinal bands (4–8 µm wide); prox-
imal sacci attachment are subequatorial, associated with
two crescentic, lateral exinal folds (2–3 µm wide), and
connected to the end of corpus; distally the sacci are slightly
inclined and sacci attachment delimits a narrow and longi-
tudinally elongated sulcus; median groove may be clearly
seen. Sacci larger than corpus and semicircular in equatorial
outline.
Dimensions. (1 specimen in polar view). Overall breadth,
60 µm; corpus breadth 30 µm, corpus length, 43 µm;
ratio corpus breadth:corpus length, 0.70; proximal saccus
breadth, 23 µm, distal saccus breadth, 15 µm, saccus
length, 38 µm.
Infraturma Striatiti Pant, 1954
Genus Hamiapollenites Wilson, 1962
Hamiapollenites sp. cf. bullaeformis (Samoilovich)
Jansonius, 1962
(Fig. 13L)
Occurrence. DCLS wells 201, 221, El Barón.
Description. Pollen grains bisaccate, taeniate, haploxy-
lonoid. Corpus subcircular to slightly transversely oval in
polar view; cappa divided transversely into 9 to 11 contin-
uous, subparallel taeniae, separated by narrow linear chan-
nels. Cappula bisected by central longitudinal exine band,
3–10 µm wide, thicker than remainder of distal exine. Sacci
much smaller than corpus, semicircular in outline; attached
equatorially to corpus.
Dimensions. (6 specimens in polar view). Overall breadth,
65(67.5)70 µm; corpus breadth, 35(43.3)50 µm; corpus
length, 35(44.3)55 µm; ratio corpus breadth:corpus length,
0.91(1.00)1.13; saccus length, 23 µm.
Remarks. Hamiapollenites bullaeformis (Samoilovich)
Jansonius, 1962 is very similar but slightly smaller. It
differs from H. fusiformis in that its sacci and corpus are
of clearly different size and shows fewer taeniae. Proto-
sacculina elliptica Menéndez (1976, p. 16, pl. 2, figs 3,
7), from the Permian of Brazil (Iratı́ Formation), is smaller
and has from 9 or 10 taeniae, and therefore is likely to be
synonymous.
Genus Lahirites Bharadwaj, 1962
Lahirites karanpuraensis Bharadwaj & Dwivedi, 1981
(Fig. 13K)
Occurrence. DCLS well 221.
Description. Pollen grain striate, bisaccate, diploxylonoid.
Central corpus dark, subcircular to oval transversely, with-
out an equatorial rim; proximally horizontally striated
 Lower Permian palynology of Paraná Basin
(number of striations, 8). Striations branched. Sacci hemi-
spherical; saccus attached along the full length of corpus;
straight at sides, convex in the middle. Distal sulcus straight
to biconvex.
Dimensions. (5 specimens in polar view). Overall breadth,
58(69.6)80 µm; corpus breadth, 38(43.4)50 µm, corpus
length, 33(37.5)43 µm; sulcus wide, 3–7 µm; saccus
length, 45(51.5)57 µm; distal saccus breadth, 25(33.3)40
µm.
Previous records. Permian of India (Bharadwaj &
Dwivedi 1981). This is the first record from Uruguay.
Genus Lueckisporites Potonié & Klaus, emend. Klaus,
1963
Lueckisporites angoulaensis Jardiné 1974
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(Fig. 14E)
1979 Lueckisporites densicorpus Archangelsky & figs 61–62), we believe this material should be included in
Gamerro: 450, pl. 9, figs 1–5.
Occurrence. DCLS well 221.
Description. Pollen grain bisaccate, strongly diploxy-
lonoid. Central corpus dark, subcircular. Cappa with two
horizontal taeniae. Taenia (6–8 µm wide) curved, having a
space at 3–5 µm wide in proximal pole. Sacci intrareticu-
late and hemipherical in polar view with a pronounced distal
inclination. Cappula about one-fifth breadth of corpus, rect-
angular, with parallel sides.
Dimensions. (2 specimens in polar view). Overall breadth,
60–75 µm; corpus breadth, 27–30 µm, corpus length,
28–30 µm; ratio corpus breadth:corpus length, 0.96–1.00;
saccus length, 38–42 µm; distal saccus breadth, 28–32 µm;
leptoma breadth, 8–10 µm.
Remarks. Lueckisporites densicorpus Archangelsky &
Gamerro (1979, p. 450, figs 1–5) presents the same char-
acteristics as L. angoulaensis (corpus with thick exine,
transversely oval, narrow cappula, outline strongly diploxy-
lonoide, similar size). The features used by Archangelsky &
Gamerro (1979, p. 450) to separate them (narrower central
ribs and lack of trilete mark) are not judged by us as suffi-
cient enough to create a new taxon. L. densus Cauduro
(1970, p. 20, figs 112–114) differs in having a circular
central corpus, a not so markedly diploxylonoid outline, and
a lower ratio of overall size:corpus than in L. angoulaensis.
Previous records. Lower Permian of Argentina
(Archangelsky & Gamerro 1979), Brazil (Souza &
Marques-Toigo 2005; Premaor et al. 2006), and Uruguay
(Beri & Daners 1995).
473
Lueckisporites singrauliensis Sinha, 1972
(Fig. 14J)
Occurrence. DCLS well 221, El Barón.
Description. Pollen grain strongly diploxylonoid, longi-
tudinally oval. Central corpus small, longitudinally oval
to circular, bearing two reniform taeniae, almost equal
in length to corpus breadth. Sacci more than half-circle,
proximally equatorially attached; distally inclined forming
narrow and straight cappula.
Dimensions. (14 specimens in polar view). Overall
breadth, 72(81.1)105 µm; corpus breadth, 36(48.8)68 µm,
corpus length, 29(37.5)56 µm; ratio corpus breadth:corpus
length, 1.00(1.19)1.42; saccus length, 38(58.5)90 µm.
Remarks. Despite the shape of the central corpus (more
elliptical than in the original; Sinha 1972, p. 187, pl. 5,
Lueckisporites singrauliensis Sinha because of similarities
in overall outline, size, form of the taeniae etc.
Previous records. Upper Permian of India (Sinha 1972).
This is the first record from Uruguay.
Genus Lunatisporites Leschik, emend. Scheuring, 1970
Lunatisporites sp. A
(Fig. 15B)
Occurrence. DCLS well 221.
Description. Pollen grains bisaccate, taeniate, strongly
diploxylonoid. Monolete geniculate, length ca 20% of
corpus width. Corpus subcircular; cappa featuring 4–6
transverse taeniae, about 5–10 µm wide. Cappula breadth
about one-quarter of corpus width, rectangular, usually
bordered by longitudinal intexinal folds asocciated with
the sacci attachment. Sacci with distal inclination.
Dimensions. (2 specimens in polar view). Overall breadth,
105–112 µm; corpus breadth, 51–53 µm, corpus length,
50–55 µm; ratio corpus breadth:corpus length, 0.93–1.06;
saccus length, 70–72 µm.
Lunatisporites sp. B
(Fig. 15F)
Occurrence. DCLS well 221.
Description. Pollen grains bisaccate, taeniate, strongly
diploxylonoid. Monolete geniculate, length ca half of
corpus wide. Corpus subcircular to transversely oval; cappa
featuring 6 transverse taeniae, about 10–20 µm wide.
Cappula breadth about half that of corpus wide, rectan-
gular. Sacci with strongly distal inclination.
 474 P. R. Gutiérrez et al.
Dimensions. (2 specimens in polar view). Overall breadth,
120–130 µm; corpus breadth, 79–84 µm, corpus length,
70–80 µm; ratio corpus breadth:corpus length, 0.98–1.20;
saccus length, 85–87 µm.
Genus Protohaploxypinus Samoilovich, emend. Morbey,
1975
Protohaploxypinus sp. cf. P. suchonensis (Sedova) Hart,
1964
(Fig. 15D)
Occurrence. El Barón.
Description. Pollen grains bisaccate, taeniate, haploxy-
lonoid, transversely oval. Corpus oval, with longitudi-
nal elongation. Cappa divided into 7 subparallel, trans-
verse taeniae of variable width (ca 2–4 µm), separated by
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narrow clefts. Cappula longitudinally rectangular (equal
to 1/4 of corpus breadth). Sacci greater than semicir-
cular in equatorial outline. Endoreticulum not clearly
discernible.
Dimensions. (1 specimen in polar view). Overall breadth,
42 µm; corpus breadth, 22 µm, corpus length, 37 µm; ratio
corpus breadth:corpus length, 0.60; saccus length, 28 µm.
Remarks. This form resembles greatly Protohaploxypinus
suchonensis (Sedova) Hart, 1964 in general morphology.
The Uruguayan form differs in having a slightly smaller
size than that observed in the original material (overall
breadth, 55–85 µm; corpus breadth, 30–44 µm).
Genus Rhizomaspora Wilson, 1962
Rhizomaspora divaricata Wilson, 1962
(Fig. 14D)
Occurrence. DCLS well 221.
Description. Pollen grain bisaccate, strongly diploxy-
lonoid. Corpus subcircular with exine 2 µm thick. Cappa
ornamentated with radiating ribs which are smooth. Sulcus
narrow, obscure. Sacci reniform, proximal attachment on
equator; distal attachment deeply inserted upon tube cell,
unattached parts often overlapping on free edges. Sacci
attachment with radial lineation.
Dimensions. (1 specimen in polar view). Overall breadth,
62 µm; corpus breadth, 23 µm, corpus length, 24 µm;
saccus length, 42 µm.
Previous records. Upper Permian of Oklahoma, USA
(Wilson 1962). Not previously recorded from Uruguay.
Genus Striatites Pant ex Jansonius & Hills, 1981
Striatites sp. A
(Fig. 15J)
Occurrence. DCLS wells 201, 221.
Description. Pollen grains bisaccate, taeniate, slightly
haploxylonoid. Corpus outline subcircular to transversely
oval. Corpus lined on the outside by an incomplete, all
round marginal proximal thickened wall. Cappa divided
into seven to twelve (generally ten) transverse taeniae,
simple to branched. Cappula defined by straight to convex
distal saccus attachment. Sacci semicircular in outline,
larger than the corpus. Saccus distal attachment full length,
straigth to slightly convex.
Dimensions. (10 specimens in polar view). Overall
breadth, 65(79.1)97 µm; sacci length, 45(51.7)60 µm;
corpus breadth, 45(55.8)70 µm; corpus length, 40(51.3)66
µm; ratio corpus breadth:corpus length, 0.91(1.03)1.15;
proximal saccus breadth, 20(24)27 µm; distal saccus
breadth, 12(15)19 µm.
Striatites sp. B
(Fig. 15E)
Occurrence. DCLS well 221, El Barón.
Description. Pollen grains bisaccate, taeniate, diploxy-
lonoid to slightly haploxylonoid. Corpus outline trans-
versely oval, darker than sacci. Cappa divided into 8 to
10 transverse taeniae, simple to branched. Cappula defined
by straight distal saccus attachment. Sacci crescentic to
semicircular in outline, larger than the corpus. Saccus
with distal attachment along full length of corpus, straight
to slightly convex; sometimes associated with exinal
folds.
Dimensions. (9 specimens in polar view). Overall breadth,
67(84.7)110 µm; sacci length, 37(50.6)65 µm; corpus
breadth, 36(54.6)72 µm; corpus length, 30(47.3)63 µm;
ratio corpus breadth:corpus length, 1.05(1.16)1.30; proxi-
mal saccus breadth, 30–31 µm.
Division Prasinophyta Round, 1971
Genus Leiosphaeridia Eisenack, emend. Downie &
Sarjeant, 1963
Leiosphaeridia sp. A
(Fig. 5D)
Occurrence. DCLS well 201.
Description. Vesicle spheroidal. Wall single-layered, thick
(2.5 µm wide), laevigate, folded. Folds ranging from a
single broad crescentic fold at vesicle margin.
Dimensions. (1 specimen). Diameter, 40 µm.
 Lower Permian palynology of Paraná Basin
Fungi
Genus Spheripollenites Couper, emend. Jansonius, 1962
Spheripollenites sp. A
(Fig. 5F)
Occurrence. DCLS wells 201, 221.
Description. Vesicle spheroidal, uniporate. Wall single-
layered, 1–1.5 µm wide, laevigate to infrapunctate, rarely
folded. Pore single, 0.5 µm diameter.
Dimensions. (10 specimens). Diameter, 20(22.5)28 µm.
Remarks. Spheripollenites stellarius (Leschik) Jain
(1968, p. 30, fig. 124) has a pore 8 µm in diameter. Inaper-
tisporites minutus (de Jersey) de Jersey (1963, p. 2, pl. 1,
figs 1–2) is similar, but shows a slightly smaller diameter
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(7–20 µm in diameter) and does not have a pore.
Microfloral characteristics
In borehole Cerrillada N◦ 221, samples were anal-
ysed between 91 and 225 m (Supplementary Data/
DI.NA.MI.GE. DCLS 221 WELL), corresponding to the
Melo Formation, more precisely to the Paso Aguiar (91 and
120 m), Mangrullo (120 and 138 m) and Fraye Muerto (138
and 225 m) members. Based on the distribution of the 109
identified species in this borehole, two assemblages can be
distinguished: 221-I (between 225 and 155 m) and 221-II
(between 140 and 91 m).
Assemblage 221-I (Frayle Muerto Member) is dominated
by trilete spores (38–92%; mean 66.3%), with subsidiary
Figure 5. A, Portalites gondwanensis Nahuys et al., 1968, FCPP
133, slide A, U43/4; B, C, Brazilea scisa (Balme & Hennelly)
Foster, 1975; B, FCPP 134, slide A, E31/1; C, FCPP 155, slide B,
D33/1; D, Leiosphaeridia sp. A, FCPP DCLS 201, 32 m, slide A,
X50/4; E, Botryococcus braunii Kützing, 1848, FCPP 153, slide
A, F28/2; F, Spheripollenites sp. A, FCPP 129, slide A, T52/0; G,
Leiosphaeridia talchirensis Lele & Karim, 1971, FCPP 134, slide
A, V36/1; H, Brazilea plurigenus (Balme & Hennelly) Foster,
1975, FCPP 134, slide A, L32/2. All x 350.
475
freshwater algae (3.5–45%; mean 18.3%), monosaccate
(3.1–11.1%; mean 7.2%) and bisaccate pollen grains
(0.7–8.9%; mean 5.5%), and rare striate (0.3–4; mean 1.7%)
and plicate pollen grains (0–7%; mean 0.3%). Among the
spores, lycophytes, especially Lundbladispora (27.3–71%;
mean 44.6%), and filicophytes (Punctatisporites: 3.3–25%;
mean 12.9%) are common; among the freshwater algae,
Botryococcus (2.8–43.3%; mean 17.1%) and Portalites
(0–1.7%; mean 1.1%) are frequent.
Some of the species found in this assemblage (e.g.,
Vallatisporites arcuatus, Portalites gondwanensis,
Scheuringipollenites barakarensis, Lundbladispora areo-
lata, Murospora torifera, Horriditriletes superbus, H.
ramosus, Anapiculatasporites sp. A, Leiotriletes directus,
L. corius, Caheniasaccites ovatus and Cristatisporites
inconstans) are also found in the upper levels (Assemblage
221-II).
Assemblage 221-II (Mangrullo and Paso Aguiar
members) is a palynoflora dominated by pollen grains, a
mixture of striate (9.6–53.3%; mean 33.1%), bisaccate non-
striate (5.0–42.8%; mean 24.8%), plicate (0.8–16.3%; mean
6.9%) and monosaccate (1.2–11.0%, mean 5.6%) forms.
There are also freshwater algae (0–79%; mean 22%) and
trilete spores (1.5–12.0%; mean 6.2%).
The predominant genera found are Botryococcus
(0–78%; mean 21.8%), Lunatisporites (0.8–12.6%; mean
6%), Limitisporites (0.4–18.2%; mean 5.3%), Lueck-
isporites (3.1–10%; mean 5%), Corisaccites (1.2–7.5%;
mean 4.6%), Protohaploxypinus (0.4–9%; mean 4.3%),
Vittatina (0.5–10%; mean 3.8%), Alisporites (0–6.9%;
mean 2.5%), Striatopodocarpites (0.4–4.1%; mean 2.4%),
Scheuringipollenites (0.4–5.2%; mean 2.1%), Platysaccus
(0–5.7%; mean 1.9%), Punctatisporites (0–2.1%; mean
1.7%), Weylandites (0–5.2%; mean 1.5%) and Lundbladis-
pora (0–4.8%; mean 1.4%).
Species that characterise this interval include:
Kraeuselisporites apiculata, Brazilea scissa,
Cannanoropollis mehtae, Limitisporites hexagonalis, L.
amazonensis, L. delasaucei, L. cf. luandensis, Alisporites
rioclarensis, A. parvus, A. similis, A. cf. opii, Falcisporites
nuthalensis, Archangelskiapollenites globocorpus, A.
plicatus, Vitreisporites pallidus, Polarisaccites bilater-
alis, Platysaccus leschikii, Pteruchipollenites gracilis,
P. cf. gondwanensis, Satsangisaccites uruguaiensis,
Scheuringipollenites medius, Mabuitasaccites crucistria-
tus, Corisaccites alutas, Hamiapollenites fusiformis, H.
cf. bullaeformis, Laharites karanpurensis, Lueckisporites
stenotaeniatus, L. latisaccus, L. singrauliensis, L. virkkiae,
L. angoulensis, Lunatisporites noviaulensis, L. variesectus,
Protohaploxypinus amplus, P. bardwajii, P. limpidus,
Staurosaccites cordubensis, Striatoabieites anaverruco-
sus, S. multistriatus, Striatopodocarpites octostriatus,
Marsupipollenites tiradiatus, Weylandites lucifer, W.
magmus, Vittatina corrugata, Pahkapites fusus, and P.
ovatus, among others.
 476 P. R. Gutiérrez et al.
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Figure 6. Correlation between the Upper Palaeozoic palynozones of the Brazilian Paraná Basin, Argentinian Paganzo Basin and Chaco-
paraná Basin, and the assemblages recognized in boreholes Paso de Melo 201, Cerrillada 221, and Barón. Abbreviations: PA Mb (Paso
Aguiar Member), Mg Mb (Mangrullo Member), FM Mb (Frayle Muerto Member).

Based on previous records of these species, it is possi-
ble to assign both assemblages to the Early Permian.
Assemblage 221-I probably indicates a slightly younger
age (end of Early Permian, beginning of Middle Permian?),
based on the presence of Lueckisporites stenotaenia-
tus, L. latisaccus, L. singrauliensis, L. virkkiae, L.
angoulensis, Lunatisporites noviaulensis, L. variesec-
tus, Laharites karanpurensis, Alisporites rioclarensis,
Striatoabieites anaverrucosus, Corisaccites alutas, Stri-
atopodocarpites octostriatus, Vitreisporites pallidus and
Falcisporites nuthalensis.
Compared with the palynostratigraphic schemes of
the Paraná (Brazil, see Souza & Marques-Toigo 2001,
2003, 2005), Chacoparaná (Argentina, see Russo et al.
1980; Césari et al. 1995; Gutiérrez et al. 2003, 2006)
and Paganzo basins (Césari & Gutiérrez 2001; Césari
2007) (Fig. 6), assemblage 221-I can be correlated with
the biozones of Vittatina costabilis (Vc, Paraná Basin),
Cristatisporites (Cr, Chacoparaná Basin) and Pakhapites
fusus-Vittatina subsaccata (FS, Paganzo Basin), as it
shares Vittatina costabilis, V. subsaccata, Lublandispora
riobonitensis and Potonieisporites novices with biozone
Vc, and Cristatisporites inconstans, Caheniasaccites
ovatus, Potonieisporites novicus, Vallatisporites arcuatus,
Horriditriletes superbus, H. ramosus, Anapiculatasporites
sp. A, Leiotriletes directus, L. corius, Lublandispora areo-
lata and Murospora torifera, among others, with biozone
Cr. Finally, it shares the presence of Vittatina subsaccata,
Pakhapites fusus and P. ovatus with biozone FS.
On the other hand, assemblage 221-II is corre-
lated with the biozones of Lueckisporites virkkiae
(Lv, Paraná), Striatites (S, Chacoparaná Basin) and
Lueckisporites-Weylandites (LW, Paganzo Basin). It shares
the presence of Lueckisporites virkkiae, L. stenotaeniatus,
L. angoulensis, Lunatisporites variesectus, Weylan-
dites lucifer, Marsupipollenites striatus, Staurosaccites
cordubensis and Vittatina costabilis with biozone Lv, and
Lueckisporites virkkiae, L. latisaccus, L. angoulensis, S.
cordubensis, Convolutispora archangelskyi, C. ordonezii,
Corisaccites alutas, Lunatisporites noviaulensis and Stri-
atoabieites anaverrucosus, among others, with biozone
S. Finally, the presence of Lueckisporites stenotaeniatus,
L. latisaccus, L. virkkiae, L. angoulensis, Lunatisporites
variesectus, Illinites unicus, Alisporites australis, Stria-
toabieites anaverrucosus, Corisaccites alutas, Staurosac-
cites cordubensis, Striatoabieites multistriatus, Weylandites
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Figure 7. A, Calamospora tener (Leschik) de Jersey, 1962, FCPP 152, slide A, N28/0; B, Leiotriletes corius Kar & Bose, 1967, FCPP
133, slide A, S54/4; C, Leiotriletes tiwarii Saxena, 1993, FCPP 133, slide A, S54/3; D, Leiotriletes directus Balme & Hennelly, 1956,
FCPP 134, slide A, G31/2; E, Punctatisporites gretensis Balme & Hennelly, 1956, FCPP 133, slide A, C57/0; F, Retusotriletes simplex
(Naumova) Potonié, 1958, FCPP 134, slide A, Q34/1; G, Brevitriletes levis (Balme & Hennelly) Bharadwaj & Srivastava, 1969, FCPP
133, slide A, E48/0; H, Cyclogranisporites gondwanensis Bharadwaj & Salujha, 1964, FCPP 129, slide A, O36/0; I, Granulatisporites
austroamericanus Archangelsky & Gamerro, 1979, FCPP 130, slide A, R38/1; J, Anapiculatisporites sp. A, FCPP 170, slide A, C39/1; K,
Brevitriletes cornutus (Balme & Hennelly) Backhouse, 1991, FCPP 134, slide A, F31/2; L, A. tereteangulus (Balme & Hennelly) Playford
& Dino, 2002, FCPP 131, slide A, K36/2; M, Convolutispora archangelskyi Playford & Dino, 2002, FCPP DCLS 221, 110 m, slide A,
D38/1, x750; N, Convolutispora ordonezii Archangelsky & Gamerro, 1979, FCPP 155, slide B, Q28/2; O, Convolutispora candiotensis
Ybert, 1975, FCPP 133, slide A, V36/0; P, Converrucosisporites micronodosus (Balme & Hennelly) Playford & Dino, 2002, FCPP 134,
slide A, F38/0; Q, Verrucosisporites andersonii (Anderson) Backhouse, 1988, FCPP 134, slide A, H51/4; R, Horriditriletes superbus
(Foster) Césari et al., 1995, FCPP 130, slide A, S31/4; S, Verrucosisporites microtuberosus (Loose) Smith & Butterworth, 1967, FCPP
150, slide A, J29/3; T, Horriditriletes uruguaiensis (Marques-Toigo) Archangelsky & Gamerro, 1979, FCPP 134, slide A, O37/4; U,
Converrucosisporites confluens (Archangelsky & Gamerro) Playford & Dino, 2002, FCPP 133, slide A, N57/0. All x 550.
 478 P. R. Gutiérrez et al.
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Figure 8. A, Horriditriletes ramosus (Balme & Hennelly) Bharadwaj & Salujha, 1964, FCPP 170, slide A, G37/0; B, Microbaculispora
sp. A, FCPP 146, slide A, V39/3; C, Murospora bicingulata Ybert, 1975, FCPP 134, slide A, X48/3; D, Murospora torifera Ybert, 1975,
FCPP 134, slide A, F51/0; E, Stenozonotriletes sp. A, FCPP 134, slide A, W37/3; F, Cristatisporites chacoparanensis Ottone, 1989,
FCPP 138, slide A, E34/2; G, Cristatisporites lestai Archangelsky & Gamerro, 1979, FCPP 138, slide A, F44/0; H, Cristatisporites
menendezii (Menéndez & Azcuy) Playford, 1978, FCPP 138, slide A, B51/4; I, Densosporites sp. A, FCPP 131, slide A, U34/4; J,
Gondisporites serrulatus Césari et al., 1995, FCPP 142, slide A, P51/2; K, Cristatisporites rollerii Ottone, 1989, FCPP 133, slide A,
F56/0; L, Cristatisporites inconstans Archangelsky & Gamerro, 1979, FCPP 165, slide A, P32/0; M, Vallatisporites russoi Archangelsky
& Gamerro, 1979, FCPP 134, slide A, Y43/0; N, Lundbladispora riobonitensis Marques-Toigo & Piccarelli, 1985, FCPP 165, slide A,
L36/3; O, Lundbladispora areolata Césari et al., 1995, FCPP 170, slide A, U40/3; P, Lundbladispora braziliensis (Pant & Srivastava)
Marques-Toigo & Pons, emend. Marques-Toigo & Piccarelli, 1985, FCPP 129, slide A, Y41/3; Q, Spelaeotriletes ybertii (Marques-Toigo)
Playford & Powis, 1979, FCPP 134, slide A, W46/3; R, Densosporites sp. B, FCPP 133, slide A, E37/3; S, Krauselisporites apiculatus
Jansonius, 1962, FCPP 130, slide A, X30/0; T, Vallatisporites arcuatus (Marques-Toigo) Archangelsky & Gamerro, 1979, FCPP 133,
slide A, C57/0; U, Krauselisporites punctatus Jansonius, 1962, FCPP 144, slide B, S28/2. All x 500.
 Lower Permian palynology of Paraná Basin 479
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Figure 9. A, Reticuloidosporites sp. A, FCPP 153, slide A, V35/0; B, Polypodiisporites mutabilis Balme, 1970, FCPP 153, slide A, F28/2;
C, Laevigatosporites vulgaris (Ibrahim) Alpern & Doubinger, 1973, FCPP 74, slide 2, E14/0; D, Barakarites rotatus (Balme & Hennelly)
Bharadwaj & Tiwari, 1964, FCPP 133, slide A, Y53/3; E, Cannanoropollis mehtae (Lele) Bose & Maheshwari, 1968, FCPP 140, slide
A, A34/3; F, Costatascyclus crenatus Felix & Burdbridge, emend. Urban, 1971, FCPP 150, slide A, D43/1; G, Cannanoropollis janakii
Potonié & Sah, 1960, FCPP 133, slide A, M54/3; H, Cannanoropollis densus (Lele) Bose & Maheshwari, 1968, FCPP 144, slide B, H31/4;
I, Plicatipollenites trigonalis Lele, 1964, FCPP 152, slide A, L35/4; J, Tuberisaccites tuberculatus (Maheshwari) Lele & Makada, 1972,
FCPP 133, slide A, B53/3; K, Potonieisporites novicus Bharadwaj, 1954, FCPP 131, slide A, W30/0; L, Potonieisporites brasiliensis
(Nahuys et al.) Archangelsky & Gamerro, 1979, FCPP 152, slide A, O32/3; M, Plicatipollenites malabarensis (Potonié & Sah) Foster,
1975, FCPP 140, slide A, B53/3; N, Potonieisporites neglectus Potonié & Lele, 1961, FCPP 144, slide B, V27/1; O, Potonieisporites
triangulatus Tiwari, 1965, FCPP 152, slide A, L36/1, x500; P, Plicatipollenites gondwanensis (Balme & Hennelly) Lele, 1964, FCPP
133, slide A, C56/4; Q, Potonieisporites magnus Lele & Karim, 1971, FCPP 155, slide B, F26/1. All x 360, except A and B x 800, and
C x 600.
 480 P. R. Gutiérrez et al.
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Figure 10. A, Caheniasaccites ovatus Bose & Kar, emend. Gutiérrez, 1993, FCPP 138, slide A, E49/0; B, Mabuitasaccites crucistriatus
(Ybert) Playford & Dino, 2000b, FCPP 152, slide A, M41/0; C, Latusipollenites quadrisaccatus Marques-Toigo, 1974, FCPP 133, slide A,
L55/0; D, Caheniasaccites densus Lele & Karim, emend. Gutiérrez, 1993, FCPP 144, slide B, C36/0; E, Striomonosaccites cicatricosus
Archangelsky & Gamerro, 1979, FCPP 150, slide A, J45/3; F, Colpisaccites granulosus Archangelsky & Gamerro, 1979, FCPP 152, slide
A, R35/0; G, Limitisporites delasaucei (Potonié & Klaus) Schaarschmidt, 1963, FCPP 152, slide A, F45/3; H, Stellapollenites talchirensis
Lele, 1965, FCPP 131, slide A, W41/4; I, Limitisporites amazonensis Playford & Dino, 2000a, FCPP 152, slide A, G36/1; J, Divarisaccus
stringoplicatus Ottone, 1991, FCPP 148, slide A, J41/1; K, Meristocorpus sp. A, FCPP 140, slide A, X41/3; L, Limitisporites vesiculosus
(Potonié & Klaus) Schaarschmidt, 1963, FCPP 144, slide B, O30/0; M, Alisporites sp. A, FCPP 152, slide A, P39/0; N, Limitisporites
rectus Leschik, 1956, FCPP 150, slide A, G34/3; O, Limitisporites sp. cf. Limitisporites luandensis Bose & Maheshwari, 1968, FCPP
153, slide A, N29/3; P, Limitisporites hexagonalis Bose & Maheshwari, 1968, FCPP 152, slide A, U39/0. All x 380, except B, F-G, I,
M-P x 500.
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Figure 11. A, C-E, Archangelskiapollenites globocorpus gen. et sp. nov.; A, paratype, FCPP 152, slide A, R35/3; C, paratype, FCPP 155,
slide B, M49/0; D, paratype, FCPP 155, slide B, V35/2; E, holotype, FCPP 152, slide A, M28/1; B, Alisporites australis de Jersey, 1962,
FCPP74, slide 2, D23/0; F-G, I, Archangelskiapollenites plicatus gen. et sp. nov.; F, holotype, FCPP 152, slide A, H35/1; G, paratype,
FCPP 152, slide A, O32/4; I, paratype, FCPP 150, slide A, D42/0; H, Alisporites similis (Balme) Dettmann, 1963, FCPP 150, slide A,
C43/2; J, Alisporites rioclarensis Menéndez, 1976, FCPP 152, slide A, K28/2; K, Alisporites parvus de Jersey, 1962, FCPP 152, slide A,
K43/2; L, Falcisporites nuthallensis (Clarke) Balme, 1970, FCPP 152, slide A, R37/3; M, Alisporites sp. cf. A. opii Daugherty, emend.
Jansonius, 1971, FCPP 152, slide A, Q35/0. All x 580.
 482 P. R. Gutiérrez et al.
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Figure 12. A, Falcisporites nuthallensis (Clarke) Balme, 1970, FCPP 152, slide A, R42/2; B-C, F, Phrixipollenites sp. A; B, FCPP 150,
slide A, G30/0; C, FCPP 152, slide A, L41/3; F, FCPP 152, slide A, L28/4; D, Chordasporites sp. cf. C. australiensis de Jersey, 1963,
FCPP 152, slide A, M38/1; E, Platysaccus sp. A, FCPP 155, slide B, U25/3; G, Platysaccus leschikii Hart, 1960, FCPP 155, slide B,
H37/1; H, K, Scheuringipollenites medius (Burjack) Dias-Fabrı́cio, 1981; H, FCPP74, slide 2, E13/0; K, FCPP 150, slide A, C35/1;
I, Platysaccus papilionis Potonié & Klaus, 1954, FCPP 152, slide A, O38/2; J, Podocarpidites sp. A, FCPP74, slide 2, S22/4; L, N,
Platysaccus orientalis sp. nov.; L, paratype, FCPP 155, slide B, V44/0; N, holotype, FCPP 155, slide B, C38/1; M, Pteruchipollenites
gracilis (Segroves) Foster, 1979, FCPP 150, slide A, H30/1. All x 600.
 Lower Permian palynology of Paraná Basin 483
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Figure 13. A, Pteruchipollenites sp. cf. Pteruchipollenites gondwanensis (Jain) Foster, 1979, FCPP 150, slide A, A43/1. B-C, E, H-I,
Satsangisaccites uruguaiensis sp. nov.; P., holotype, FCPP 152, slide A, O30/2; C, paratype, FCPP 152, slide A, M44/4; E, paratype, FCPP
152, slide A, J36/2; H, paratype, FCPP 152, slide A, G55/1; I, paratype, FCPP 150, slide A, D43/1; D, Satsangisaccites sp. A, FCPP74,
slide 2, D23/0; F, Scheuringipollenites circularis Césari et al., 1995, FCPP 150, slide A, C41/0; G, Scheuringipollenites barakarensis
(Tiwari) Tiwari, 1973, FCPP 144, slide B, J29/4; J, Lueckisporites virkkiae (Potonié & Klaus) Klaus, 1963, FCPP 152, slide A, B43/0; K,
Lahirites karanpuraensis Bharadwaj & Dwivedi, 1981, FCPP 155, slide B, B25/0; L, Hamiapollenites sp. cf. H. bullaeformis (Samoilovich)
Jansonius, 1962, FCPP 153, slide A, L41/0; M, Hamiapollenites fusiformis Marques-Toigo, emend. Archangelsky & Gamerro, 1979,
FCPP 152, slide A, N38/1; N, Vitreisporites pallidus (Reissinger) Nilsson, 1958, FCPP 155, slide B, F30/4; O, Corisaccites alutas
Venkatachala & Kar, 1966b, FCPP 152, slide A, K28/1. All x 550.
 484 P. R. Gutiérrez et al.
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Figure 14. A, C, Lueckisporites latisaccus Archangelsky & Gamerro, 1979; A, FCPP 152, slide A, K43/4; C, FCPP 152, slide A, G36/1;
B, Lueckisporites virkkiae (Potonié & Klaus) Klaus, 1963, FCPP 152, slide B, P37/1; D, Rhizomaspora divaricata Wilson, 1962, FCPP
152, slide A, Q38/2; E, Lueckisporites angoulaensis Jardiné, 1974, FCPP 152, slide A, M40/3; F, Protohaploxypinus limpidus (Balme &
Hennelly) Balme & Playford, 1967, FCPP 155, slide B, E38/0; G, L, Lueckisporites stenotaeniatus Menéndez, 1976; G, FCPP 152, slide
A, N40/3; L, FCPP 152, slide A, C37/1; H, Lunatisporites pellucidus (Goubin) Helby, emend. de Jersey, 1972, FCPP 133, slide A, R56/1;
I, Lunatisporites variesectus Archangelsky & Gamerro, 1979, FCPP 152, slide A, B32/3; J, Lueckisporites singrauliensis Sinha, 1972,
FCPP 152, slide A, P38/1; K, Lunatisporites noviaulensis (Leschik) de Jersey, 1979, FCPP 152, slide A, O39/0. All x 570.
 Lower Permian palynology of Paraná Basin 485
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Figure 15. A, C, Lunatisporites variesectus Archangelsky & Gamerro, 1979; A, FCPP 152, slide A, E42/3; C, FCPP 152, slide A, K41/0;
B, Lunatisporites sp. A, FCPP 152, slide A, T36/0; D, Protohaploxypinus sp. cf. P. suchoneneis (Sedova) Hart, 1964, FCPP74, slide 2,
T22/0; E, Striatites sp. B, FCPP 152, slide A, L36/2; F, Lunatisporites sp. B, FCPP 152, slide A, O38/0; G, Protohaploxypinus goraiensis
(Potonié & Lele) Hart, 1964, FCPP 130, slide A, N41/4; H, Protohaploxypinus amplus (Balme & Hennelly) Hart, 1964, FCPP 152, slide
A, O41/4; I, K, Staurosaccites cordubensis Archangelsky & Gamerro, 1979; I, FCPP 152, slide A, N42/0; K, FCPP 150, slide A, J31/4;
J, Striatites sp. A, FCPP 152, slide A, L33/2; L, Striatoabieites multistriatus (Balme & Hennelly) Hart, 1964, FCPP 150, slide A, C40/2;
M, Protohaploxypinus bhardwajii Foster, 1979, FCPP 152, slide A, O36/0. All x 550.
 486 P. R. Gutiérrez et al.
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Figure 16. A, Striatoabieites anaverrucosus Archangelsky & Gamerro, 1979, FCPP 152, slide A, H42/0; B, Striatopodocarpites octos-
triatus Hart, 1960, FCPP 150, slide A, J24/4; C, Striatopodocarpites cancellatus (Balme & Hennelly) Hart, 1963, FCPP 152, slide A,
J42/3; D, Polarisaccites bilateralis Ybert & Marques-Toigo, 1971, FCPP 155, slide B, P31/0; E, Illinites unicus Kosanke, 1950, FCPP
133, slide A, F46/0; F, Striatopodocarpites gondwanensis Lakhanpal et al., 1960, FCPP74, slide 2, S18/0; G, K, Vittatina subsaccata
Samoilovich, emend. Jansonius, 1962; G, FCPP 152, slide A, G43/0; K, FCPP 152, slide A, S42/0; H, Weylandites lucifer (Bharadwaj &
Salujha) Foster, 1975, FCPP 152, slide A, S35/4; I, Marsupipollenites striatus (Balme & Hennelly) Hart, 1965, FCPP74, slide 2, E43/1; J,
N, Vittatina corrugata Marques-Toigo, 1974; J, FCPP 152, slide A, T34/4; N, FCPP 150, slide A, C31/0; L, Vittatina costabilis (Wilson)
Tschudy & Kosanke, 1966, FCPP 150, slide A, C41/1; M, Weylandites magmus (Bose & Kar) Backhouse, 1991, FCPP 74, slide 2, V32/0;
O, Pakhapites fusus (Bose & Kar) Menéndez, 1971, FCPP 131, slide A, L35/1; P, Pakhapites ovatus (Bose & Kar) Garcı́a, 1996, FCPP
133, slide A, N43/0. Q, Marsupipollenites triradiatus Balme & Hennelly, 1956, FCPP 150, slide A, C42/0; R, Tetraporina punctata
(Tiwari & Navale) Kar & Bose 1976, FCPP 138, slide A, E31/1. All x 600.
 Lower Permian palynology of Paraná Basin
magmus and Vittatita subsaccata is shared with biozone LW
(earliest Permian).
Borehole DI.NA.MI.GE. DCLS 201 (Electronic Supple-
mentary Data/DI.NA.MI.GE. DCLS 201 WELL) pene-
trated about 89 m of Permian sedimentary rocks, assignable
to the Melo (Frayle Muerto Member, 6–32 m deep)
and Tres Islas (32–93 m) formations, yielding 98
species. Although most species are present throughout
the column (among them Kraeuselisporites punctatus,
Latusipollenites quadrisaccatus, Lueckisporites virkkiae,
Scheuringipollenites barakarensis, S. circularis, Vittatina
corrugata, Converrucosisporites confluens and Horrid-
itriletes uruguaiensis), it is possible to differentiate two
assemblages: 201-I and 201-II.
Assemblage 201-I (34–93 m) includes microflora domi-
nated by trilete spores (55.3–96.1%; mean 79.3%), accom-
panied by monosaccate (1–36.7%; mean 11.8%) and
Downloaded By: [GutiérrezBuenos Aires, Pedro] At: 16:04 14 November 2010
bisaccate pollen grains (1.8–9.3%; mean 4.3%). Other
groups are present in low proportions. Among the
spores, filicophytes and lycophytes, such as Punc-
tatisporites (1.9–81.3%; mean 43.3%), Lundbladispora
(0–65%; mean 20.4%), Cristatisporites (0–10.2%; mean
5.5%), Vallatisporites (0–6.7%; mean 2.9%), Leiotriletes
(0.5–2.5%; mean 1.1%) and Murospora (0.2–2.6%; mean
1.1%) are frequent. Also, pollen grains of gymnosperms
are found, such as Potonieisporites (0–5.1%; mean 1.2%),
Cannanoropollis (0–15.3%; mean 4%) and Scheuringipol-
lenites (0.3–1.4%; mean 1.1%).
Species typical of this interval include: Leiotriletes
corius, L. tiwari, Punctatisporites gretensis, Retusotriletes
simplex, Stenozonotriletes sp. A, Cristatisporites chaco-
paranensis, C. lestai, C. rollerii, Lundbladispora areolata,
Murospora bicingulata, M. torifera, Vallatisporites arcu-
atis, V. russoi, Caheniasaccites densus, Limitisporites vesis-
culosus, Protohaploxypinus bharadwajii, Illinites unicus
and Leiosphaeridia talchirensis.
Assemblage 201-II (6–34 m) is characterised by the
predominance of trilete spores (54–69%; mean 59.8%),
as well as monosaccate (9–23.3; mean 16.6%) and bisac-
cate pollen grains (7.3–14.2%; mean 11.7%). The spores
Lundbladispora (33.3–47.7%; mean 39.9%) and Punc-
tatisporites (7.8–12%; mean 10.2%) are frequent, as is
the bisaccate pollen grain Scheuringipollenites (3.8–5.9%;
mean 5%).
Species typical of this assemblage include Anapic-
ulatisporites tereteangulus, Cyclogranisporites gondwa-
nensis, Verrucosisporites andersonii, Brevitriletes levis,
Spelaeotriletes ybertii, Pakhapites ovatus, P. fusus,
Platysaccus papilionis, Polarisaccites bilateralis, Proto-
haploxypinus limpidus, Striatopodocarpites cancellatus
and Spheripollenites sp. A, among others.
Both assemblages are equivalent to the biozones
Vittatina costabilis (Vc, Paraná Basin), Cristatisporites
(Cr, Chacoparaná Basin) and Pakhapites fusus-Vittatita
subsaccata (FS, Paganzo Basin). They share the
487
following taxa with these biozones: Vittatina subsac-
cata, Stellapollenites talchirensis, Converrucosisporites
confluens, Hamiapollenites fusiformis, Illinites unicus,
Lundbladispora riobonitensis, Potonieisporites novicus,
Spelaeotriletes ybertii, Protohaploxypinus goraiensis and
Cristatisporites inconstans. They also share with biozone
Cr the following species: C. inconstans, Converuco-
sisporites micronodosus, C. confluens, Vallatisporites
russoi, Lundbladispora areolata, Gondisporites serru-
latus, Leiotriletes tiwari, Brevitriletes levis, B. cornu-
tus, Convolutispora ordonezii, Horriditriletes superbus,
Hamiapollenites fusiformis, Illinites unicus, Mabuitasac-
cites crucistriatus, Striomonosaccites cicatricosus, Stri-
atopodocarpites cancellatus, Scheuringipollenites medius,
Vittatina subsaccata, V. costabilis, Pahkapites fusus and
Weylandites lucifer, among others.
Finally, Barakarites rotatus, Pakhapites fusus, Vittatina
subsaccata, Protohaploxypinus limpidus, Latusipollen-
ites quadrisaccatus, Cannanoropollis mehtae, Lund-
bladispora brasiliensis, L. riobonitensis, Cristatisporites
rolleri, C. menendezii, Striatoabieites multistriatus and
Marsupipollenites striatus are shared with biozone
FS.
The following taxa are added to the assemblage listed
by Beri and Pecoits (2001, with 30 species) from El
Barón: Convolutispora archangelskyi, Potonieisporites
novicus, Mabuitasaccites crucistriatus, Archangelskiapol-
lenites plicatus, Colpisaccites granulosus, Alisporites
rioclarensis, A. similis, Chordasporites cf. australiensis,
Falcisporites nuthallensis, Prixipollenites sp. A, Platysac-
cus papilionis, Podocarpidites sp. A, Satsangisaccites
uruguaiensis, S. sp. A, Scheuringipollenites circularis,
S. barakarensis, S. medius, Vitreisporites pallidus,
Polarisaccites bilateralis, Hamiapollenites cf. bullae-
formis, Lueckisporites latisaccus, L. singraulensis, L.
virkkiae, Protohaploxypinus limpidus, P. cf. suchonensis,
Striatites sp. B, Striatopodocarpites gondwanensis, S.
cancellatus, S. octostriatus, Marsupipollenites striatus,
Weylandites magmus and Brazilea plurigenus. This
microflora is correlated with assemblage 221-II and,
therefore, equivalent to the biozones of Lueckisporites
virkkiae (Paraná Basin), Striatites (Chacoparaná Basin)
and Lueckisporites-Weylandites (Paganzo Basin).
Acknowledgements
This research has been funded by the Universidad de
la República (research grant CSIC, I+D) and Agencia
Nacional de Promoción Cientı́fica y Técnica (research grant
ANPCYT-PICT 11817, 32693) awarded to Á. Beri and P.R.
Gutiérrez, respectively. These studies were carried out as
part of of the CSIC, I+D Project (“Estudio palinológico de
sedimentos del Paleozoico Superior en las cuencas Paraná
and Chacoparanense”), Facultad de Ciencias (Uruguay).
 488 P. R. Gutiérrez et al.
Note
Supplementary material can be viewed online.
References
Alpern, B. & Doubinger, J. 1973. Microfossiles organiques du
Paléozoı̈que. 6. Les Miospores Monoletes du Paléozoı̈que.
CIMP/CNRS, Paris, 103 pp.
Anderson, J. M. 1977. The biostratigraphy of the Permian and
Triassic. Part 3. A review of Gondwana Permian palynology
with particular reference to the Northern Karoo Basin, South
Africa. Memoirs of the Botanical Survey of South Africa, 41,
1–67.
Andreis, R. R., Ferrando, L. A. & Herbst, R. 1996. Terrenos
carbonı́feros y pérmicos de la República Oriental del Uruguay.
Pp. 309–390 in S. Archangelsky (ed.) El Sistema Pérmico en
Downloaded By: [GutiérrezBuenos Aires, Pedro] At: 16:04 14 November 2010
la República Argentina y en la R. O. del Uruguay. Academia
Nacional de Ciencias, Córdoba.
Anzótegui, L. M. & Mautino, L. R. 1992. Palinomorfos de la
Formación Melo (Pérmico Inferior) de Uruguay. 8◦ Simposio
Argentino de Paleobotánica y Palinologı́a, Corrientes 1991.
Asociación Paleontológica Argentina, Publicación Especial,
2, 11–14.
Archangelsky, S. & Gamerro, J. C. 1979. Palinologı́a del Paleo-
zoico Superior en el subsuelo de la Cuenca Chacoparanense,
República Argentina. I. Estudio sistemático de los palinomor-
fos de tres perforaciones de la provincia de Córdoba. Revista
Española de Micropaleontologı́a, 11, 417–478.
Archangelsky, S. & Gamerro, J. C. 1980. Palinomorfos
pérmicos del subsuelo de la Cuenca Colorado, en la
Plataforma del Mar, Argentina, Provincia Buenos Aires. Bole-
tim IG Instituto de Geociências USP, 11, 119–124.
Archangelsky, S., Gamerro, J. C. & Leguizamón, R. R. 1980.
Estudios palinológicos de las perforaciones YCF CO1, CO2
y CO3 (Paleozoico superior), Santiago Temple, provincia de
Córdoba. II Congreso Argentino de Paleontologı́a y Bioes-
tratigrafı́a y I Congreso Latinoamericano de Paleontologı́a,
Buenos Aires 1978, Actas, 4, 45–49.
Archangelsky, S. & Vergel, M. M. 1996. Capı́tulo 2. Cuenca
Chacoparanense. Paleontologı́a, bioestratigrafı́a y paleoe-
cologı́a. Pp. 40–44 in S. Archangelsky (ed.) El Sistema
Pérmico en la Republica Argentina y en la República Oriental
del Uruguay. Academia Nacional de Ciencias, Córdoba.
Azcuy, C. L. & Di Pasquo, M. 2000. Palynology of the Late
Carboniferous from the Tarija Basin, Argentina: A systematic
review of monosaccate pollen genera. Palaeontographica, B,
253, 107–137.
Azcuy, C. L., Di Pasquo, M. & Valdivia Ampuero, H. 2002. Late
Carboniferous miospores from the Tarma Formation, Pongo
de Mainique, Perú. Review of Palaeobotany and Palynology,
118, 1–28.
Backhouse, J. 1988. Late Jurassic and early Cretaceous palynol-
ogy of the Perth basin, Western Australia. Geological Survey
of Western Australia Bulletin, 135, 1–233.
Backhouse, J. 1991. Permian palynostratigraphy of the Collie
Basin, Western Australia. Review of Palaeobotany and Paly-
nology, 67, 237–314.
Balarino, M. L. & Gutiérrez, P. R. 2006. Palinologı́a de
la Formación Tasa Cuna (Pérmico Inferior), Córdoba,
Argentina: sistemática y consideraciones bioestratigráficas.
Ameghiniana, 43, 437–460.
Balme, B. E. 1963. Plant microfossils from the Lower Triassic of
Western Australia. Palaeontology, 6, 12–40.
Balme, B. E. 1970. Palynology of Permian and Triassic Strata
in the Salt Range and Surghar Range, West Pakistan.
Pp. 305–453 in B. Kummel & C. Teichert (eds) Stratigraphic
Boundary problems: Permian and Triassic of West Pakistan.
University of Kansas, Special Publication, 4.
Balme, B. E. 1979. Palynology of Permian-Triassic Bound-
ary beds at Kap Stosch, East Greenland. Meddelelser om
Grønland, 200(6), 4–37.
Balme, B. E. & Hennelly, J. P. F. 1956. Trilete sporomorphs from
Australian Permian Sediments. Australian Journal of Botany,
4, 240–260.
Balme, B. E. & Playford, G. 1967. Late Permian plant microfos-
sils from the Prince Charles Mountains, Antarctica. Revue de
Micropaléontologie, 10, 179–192.
Banerjee, M. 1988. Karharbari: a formation or biozone. The
Palaeobotanist, 36, 37–50.
Beri, A. 1987. Estudio perliminar del contenido palinológico de la
perforación 201 (Carbonı́fero Superior/Pérmico Inferior) del
NE del Uruguay. VII Simposio Argentino de Paleobotánica y
Palinologı́a; Buenos Aires, pp. 33–36.
Beri, A. 1988. Estudio sistematico del contenido palinologico de
una muestra de la Formacion Tres Islas (Permico Inferior) del
Uruguay. Paula Coutina, 3, 27–48.
Beri, A. 2003. Revisión del conocimiento paleontológico del Pale-
ozoico Tardı́o del Uruguay. Revista del Museo Argentino de
Ciencias Naturales, nueva serie, 5(2), 163–168.
Beri, A. & Daners, G. 1995. Palinologı́a de la perforación N.
221, Pérmico, R. O. del Uruguay. Revista Geociências, 14,
145–160.
Beri, A. & Daners, G. 1998. Estudio palinologico de la perfora-
cion 201 (DI.NA.MI.GE.), Pérmico, Uruguay. Boletin de
la Real Sociedad Espanola de Historia Natural, Sección
geológica, 94, 55–64.
Beri, A. & Goso, C. A. 1996. Análisis palinológico y estratigráfico
de la Fm. San Gregorio (Pérmico Inferior) en el área de los
cerros Guazunambi, Cerro Largo, Uruguay. Revista Española
de Micropaleontologı́a, 28, 211–223.
Beri, A. & Goso, C. A. 1998. Resultados palinológicos y
estratigráficos de la formación San Gregorio (Pérmico Infe-
rior), Uruguay. Geociências (Revista da Universidade de
Guarulhos), 3(6), 108–119.
Beri, A. & Pecoits, E. 2001. Palinologı́a y sedimentologı́a de
la Formación Melo (Pérmico) en la cañada “El Barón”,
departamento de Cerro Breadth, Uruguay. Ameghiniana, 38,
419–428.
Beri, A., Pecoits, E. & Tejera, L. 2000. Palinologı́a de la
Formación Tres Islas en la perforación 313/1, Pérmico Infe-
rior, Uruguay. Geociências (Revista da Universidade de
Guarulhos), 5(6), 71–79.
Beri, A., Gutiérrez, P. R., Cernuschi, F. & Balarino, M. L.
2006. Palinologı́a del Pérmico Inferior en la perforación CLS-
24 (Formación San Gregorio), departamento de Cerro Largo,
Uruguay. Parte 1: esporas, algas, prasinofitas y acritarcas.
Ameghiniana, 43, 227–244.
Bharadwaj, D. C. 1954. Einige neue Sporengattungen des
Saarkarbons. Neues Jahrbuch für Geologie und Paläontolo-
gie, 1954, 512–525.
Bharadwaj, D. C. 1962. The miospore genera in the coals of
Raniganj Stage (Upper Permian), India. The Palaeobotanist,
9, 68–106.
Bharadwaj, D. C. 1964. Potonieisporites Bhard., ihre Morpholo-
gie, Systematik and Stratigraphie. Fortschritte in der Geologie
von Rheinland und Westfalen, 12, 45–54.
 Lower Permian palynology of Paraná Basin
Bharadwaj, D. C. & Dwivedi, A. 1981. Sporae Dispersae of the
Barakar sediments from South Karanpura Coalfield, Bihar,
India. The Palaeobotanist, 27, 21–94.
Bharadwaj, D. C. & Salujha, S. K. 1964. Sporological study
of seam VIII in Raniganj Coalfield, Bihar (India). Part 1.
Description of Sporae Dispersae. The Palaeobotanist, 12,
181–215.
Bharadwaj, D. C. & Srivastava, S. C. 1969. Some new miospores
from Bharakar Stage, Lower Gondwana, India. The Palaeob-
otanist, 17, 220–229.
Bharadwaj, D. C. & Tiwari, R. S. 1964. On two monosaccate
genera from Barakar Stage of India. The Palaeobotanist, 12,
139–146.
Bharadwaj, D. C., Kar, R. K. & Navale, G. K. B. 1976.
Palynostratigraphy of Lower Gondwana deposits in Paraná
and Maranhao basins, Brazil. Biological Memoirs, 1, 56–
103.
Biswas, B. 1962. Stratigraphy of the Mahadeo, Langpar, Cherra
and Tura Formations, Assam, India. Bulletin of the Geologi-
cal, Mining and Metallurgical Society, 25, 1–48.
Downloaded By: [GutiérrezBuenos Aires, Pedro] At: 16:04 14 November 2010
Borge, O. & Pascher, A. 1913. Zygnemales. P. 151 in A. Pascher
(ed.) Die Süsswasser-Flora Deutschlands, Österreichs und
der Schweiz, 9. Verlag Gustav Fischer, Jena, 51 pp.
Bose, M. N. & Kar, R. K. 1966. Palaeozoic Sporae Dispersae
from Congo. I-Kindu-Kalima and Walikale Regions. Annales
de Musée Royal de l’Afrique Centrale, Série 8, Sciences
géologiques, 53, 1–238.
Bose, M. N. & Maheshwari, H. K. 1968. Palaeozoic Sporae
Dispersae from Congo. VII. Coal measures near Lake
Tanganyika, south of Albertville. Annales de Musee Royal
de l’Afrique Centrale, Serie 8, Annales de Musee Royal de
l’Afrique Centrale, Serie 8, 60, 1–116.
Bossi, J. 1966. Geologı́a del Uruguay. Colección Ciencias N◦
2. Departamento de Publicaciones de la Universidad de la
República. Montevideo.
Bossi, J. & Navarro, R. 1988. Geologı́a del Uruguay. Tomo I y
II. Departamento de Publicaciones de la Universidad de la
República. Montevideo, Uruguay.
Bossi, J., Ferrando, L. A., Montaña, J., Campal, N., Morales,
H., Gancio, F., Schipilov, A., Piñeyro, D. & Sprechmann, P.
1998. Carta geológica del Uruguay. Escala 1/500.000. Memo-
ria Explicativa. Geoeditores. Montevideo, Uruguay.
Burden, E. T. & Hills, L. V. 1989. Illustrated key to genera
of Lower Cretaceous terrestrial palynomorphs (exclud-
ing megaspores) of western Canada. American Associa-
tion of Stratigraphic Palynologists Contributions Series, 21,
1–147.
Butterworth, M. A., Alpern, B., Artüz, S., Jansonius, J.,
Kosanke, R. M., Richardson, J. B., Smith, A. V. H. &
Staplin, F. L. 1964. Densosporites (Berry) Potonié & Kremp
and related genera. V Congres International de Stratigraphie
et de Géologie du Carbonifere (Paris 1963), Compte Readu,
3, 1049–1057.
Cauduro, A. D. 1970. Lower Gondwana Miospores from Sâo
Sepé outcrop (Rio Grande do Sul, Brasil). Escola de Geolo-
gia, Universidade Federal de Rio Grande do Sul, Publicaçâo
Especial, 17, 1–34.
Cazzulo-Klepzig, M., Guerra-Sommer, M., Formoso, N. L.
& Calarge, L. M. 2002. Geochemical and palynological
evidence for the age determination of Permian coals, south-
ern Brazil. Journal of South American Earth Sciences, 15,
375–380.
Césari, S. N. 2007. Palynological biozones and radiometric data at
the Carboniferous-Permian boundary in Western Gondwana.
Gondwana Research, 11, 529–536.
489
Césari, S. N. & Bercowski, F. 1998. Palinologı́a de la Formación
Jejenes (Carbonı́fero) en la quebrada de Las Lajas, provincia
de San Juan, Argentina. Nuevas inferencias paleoambientales.
Ameghiniana, 34, 497–509.
Césari, S. N. & Gutiérrez, P. R. 1985. Microflora de la locali-
dad de Los Mogotes Colorados (Paleozoico superior), provin-
cia de La Rioja, República Argentina. Boletim Instituto de
Geociências, Universidade de Saõ Paulo, 15, 20–31.
Césari, S. N. & Gutiérrez, P. R. 2001. Palynostratigraphy of
the Upper Paleozoic sequences in central-western Argentina.
Palynology, 24, 113–146.
Césari, S. N. & Limarino, C. O. 2002. Palynology of glacial sedi-
ments from the Guandacol Formation (Middle Carboniferous)
in the Cerro Bola area, Paganzo Basin, Argentina. Alcheringa,
26, 159–176.
Césari, S. N., Archangelsky, S. & Seoane, L. V. 1995. Pali-
nologı́a del Paleozoico Superior de la perforación Las
Mochas, provincia de Santa Fé, Argentina. Ameghiniana, 32,
73–106.
Césari, S. N., Meza, J. C. & Melchor, R. N. 1996. Primer registro
palinológico de la Cuenca Pérmica Oriental (Fm. Yacimiento
Los Reyunos), Mendoza, Argentina. 13◦ Congreso Geológico
Argentino y 3◦ Congreso de Exploración de Hidrocarburos
(Mendoza), Actas, 5, 49–63.
Chandra, A. & Lele, K. M. 1980. Talchir miofloras from South
Rewa Gondwana Basin, India and their biostratigraphical
significance. Proceedings of the IV International Palynologi-
cal Conference (Lucknow 1976–77), 2, 117–151.
Clarke, R. F. A. 1965. British Permian saccate and monosulcate
miospores. Palaeontology, 8, 322–354.
Cookson, I. C. 1947. Plant microfossils from lignites of Kerguelen
Archipelago. Antarctic Research Expedition 1929–1931, 2,
127–142.
Couper, R. A. 1953. Upper Mesozoic and Cainozoic spores and
pollen grains from New Zealand. New Zealand Geological
Survey Paleontological Bulletin, 22, 1–77.
Couper, R. A. 1958. British Mesozoic microspores and pollen
grains. A systematic and stratigraphic study. Palaeontograph-
ica B, 103, 75–179.
Crosbie, Y. M. 1985. Permian palynomorphs from the Kuriwao
Group, Southland, New Zealand. New Zealand Geological
Survey Record, 8, 109–119.
Daemon, R. F. & Quadros, L. P. 1970. Bioestratigrafia de Neopa-
leozóico da Bacia do Paraná. 24th Congresso Brasileiro da
Geologia, Brasilia, 1970, 359–412.
De Jersey, N. J. 1962. Triassic spores and pollen grains from the
Ipswich Coalfield. Geological Survey of Queensland, Publi-
cation, 307, 1–20.
De Jersey, N. J. 1963. Jurassic spores and pollen grains from
the Marburg Sandstone. Geological Survey of Queensland,
Publication, 313, 1–15.
De Jersey, N. J. 1972. Triassic miospores from the Esk Beds.
Geological Survey of Queensland, Publication, 357, 40 pp.
De Jersey, N. J. 1979. Palynology of the Permian-Triassic tran-
sition in the western Bowen Basin. Geological Survey of
Queensland, Publication, 374, 1–39.
De Santa Ana, H. 2004. Análise Tectono-estratigráfica das
Secüências Permo-Triássicas e Juro-Cretáceas da Bacia
Chacoparanaense Uruguaia. Tesis de Doctorado, Rio Claro
(SP), UNESP, Brasil.
De Santa Ana, H., Goso, C. & Daners, G. 2006. Cuenca Norte:
Estratigrafı́a del Carbonı́fero-Pérmico. Pp. 147–208 in G.
Veroslavsky, M. Ubilla & S. Martinez (eds) Cuencas Sedi-
mentarias de Uruguay. Geologı́a, Paleontologı́a & Recursos
Naturales. DIRAC. Montevideo, Uruguay.
 490 P. R. Gutiérrez et al.
Dellazzana, J. G. 1976. Contribuiçao à palinologia da Formaçao
Iratı́ (Permiano), Rio Grande do Sul, Brasil. Ameghiniana, 13,
1–42.
Dettmann, M. E. 1963. Upper Mesozoic microfloras from south-
eastern Australia. Proceedings of the Royal Society of Victo-
ria, 77, 1–148.
Di Pasquo, M. 2002. The Crassispora kosankei-Cystoptychus
azcuyi palynozone of the Upper Carboniferous Tupambi
Formation, Tarija Basin, northern Argentina. Review of
Palaeobotany and Palynology, 118, 47–76.
Di Pasquo, M., Azcuy, C. L. & Starck, D. 2001. Palinologı́a de
la Formación San Telmo (Carbonı́fero Superior) en la sierra
de San Antonio, provincia de Salta, Argentina. Ameghiniana,
38, 85–98.
Di Pasquo, M., Azcuy, C. L. & Souza, P. A. 2003a. Pali-
nologı́a del Carbonı́fero Superior del Subgrupo Itararé en
Itaporanga, Cuenca Paraná, Estado de Sao Paulo, Brasil. Parte
1: sistemática de esporas y paleofitoplancton. Ameghiniana,
40, 277–296.
Di Pasquo, M., Azcuy, C. L. & Souza, P. A. 2003b. Pali-
Downloaded By: [GutiérrezBuenos Aires, Pedro] At: 16:04 14 November 2010
nologı́a del Carbonı́fero Superior del Subgrupo Itararé en
Itaporanga, Cuenca Paraná, Estado de Sâo Paulo, Brasil. Parte
2: sistemática de polen y significado paleoambiental y estrati-
grafico. Ameghiniana, 40, 297–313.
Dias, M. E. R. 1993. Palinologia do Grupo Itararé na porção
centro-sul do Rio Grande do Sul, Permiano da Bacia do
Paraná, Brasil. Pesquisas, 20, 119–131.
Dias-Fabricio, M. E. 1981. Palinologia da Formaçao Rio Bonito
na área de Gravataı́-Morungava, Rio Grande so Sul. Pesquisas,
14, 69–130.
Dibner, A. F. 1971. Cordaites pollen of Angaraland. Uchenye
Zapiski, Nauchno Issled Institut Geolgii Artsakhskij, 32,
5–66.
Dino, R. & Playford, G. 2002. Stratigraphic and palaeoenviron-
mental significance of a Pennsylvanian (Upper Carbonifer-
ous) palynoflora from the Piauı́ Formation, Parnaı́ba Basin,
northeastern Brazil. Paleontological Research, 6, 23–40.
Dino, R., Antonioli, L. & Braz, S. M. N. 2002. Palynological data
from the Trisidela Member of Upper Pedra de Fogo Formation
(“Upper Permian”) of the Parnaiba Basin, northeastern Brazil.
Revista Brasileira de Paleontologia, 3, 24–35.
Dolby, J. H. & Balme, B. E. 1976. Triassic palynology of the
Carnarvon Basin, Western Australia. Review of Palaeobotany
and Palynology, 22, 105–168.
Downie, C. & Sarjeant, W. A. S. 1963. On the interpretation
and status of some hystrichosphere genera. Palaeontology, 6,
83–96.
Dunay, R. E. & Fisher, M. J. 1979. Palynology of the Dockum
Group (Upper Triassic), Texas, U.S.A. Review of Palaeob-
otany and Palynology, 28, 61–92.
Dybová, S. & Jachowicz, A. 1957. Mikrospory górno-ślaskiego

du Bassin de Morondava (Madagascar). Revue de l’Institut
karbonu produktywnego. Prace Instytut Geologiczny, 23,
1–328.
Erdtman, G. 1947. Suggestions for the classification of fossil and
recent pollen grains and spores. Svensk Botanisk Tidskrift, 41,
104–114.
Evitt, W. R. 1963. A discussion and proposals concerning fossil
dinoflagellates, hystrichospheres, and acritarchs. I. Proceed-
ings of the National Academy of Sciences of the United States
of America, 49, 158–164.
Falcon, R. M. S. 1978. New Permian miospores from Rhodesia:
Arnoldia Rhodesia, 8, 10–32.
Farabee, M. J. & Canright, J. E. 1986. Stratigraphic palynology
of the lower part of the Lance Formation (Maestrichtian) of
Wyoming. Palaeontographica B, 199, 1–89.
Farabee, M. J., Taylor, E. L. & Taylor, T. N. 1991. Late
Permian palynomorphs from the Buckley Formation, central
Transantarctic Mountains, Antarctica. Review of Palaeob-
otany and Palynology, 69, 353–368.
Fasolo, Z. & Vergel, M. 1994. Palinologı́a de la perforación Fraile
Muerto (FM2), Formación Tres Islas (Pérmico Inferior) de la
Cuenca Paraná de Uruguay. Acta Geológica Leopoldesiana,
39, 679–689.
Ferrando, L. A. & Andreis, R. R. 1982. Correlación entre
las columnas estratigráficas paleozoicas de Uruguay y Rio
Grande del Sur, Brasil. (Resumen). Proyecto N◦ 42 IUGS-
UNESCO. Boletı́n, 5, 34–35. Montevideo, Uruguay.
Ferrando, L. A. & Andreis, R. R. 1986. Nueva estratigrafı́a
en el Gondwana de Uruguay. 1◦ Congreso Latinoamericano
de Hidrocarburos, Buenos Aires, Actas, 295–323. ARPEL,
Argentina.
Ferrando, L. A. & Andreis, R. R. 1990. La glaciación
gondwánica en Uruguay, extremo austral de la cuenca intra-
cratónica de Paraná. Proyecto N◦ 211. IUGS-UNESCO.
Buenos Aires. Argentina, pp. 72–76.
Felimean C. J. & Burbridge, P. P. 1967. Palynology of the
Springer Formation of Southern Oklahoma, U.S.A. Palaeon-
tology, 10, 349–425.
Felix, C. M., Premaor, E., Hermany, G. & Souza, P. A. 2005.
Análise palinotaxonômica e bioestratigráfica da Subturma
Monosaccites na Bacia do Paraná, Brasil. I: Plicatipollen-
ites Lele 1964 e Crucisaccites Lele & Maithy 1965. Revista
Brasileira de Paleontologia, 9, 63–72.
Foster, C. B. 1975. Permian plant microfossils from the Blair
Athol Coal Measures, central Queensland, Australia. Palaeon-
tographica B, 154, 121–171.
Foster, C. B. 1979. Permian plant microfossils of the Blair Athol
Coal Measures, Baralaba Coal Measures, and basal Rewan
Formation of Queensland. Geological Survey of Queensland,
Publication, 372, 1–244.
Gamerro, J. C. & Archangelsky, S. 1981. Hallazgos de pali-
nomorfos pérmicos en sedimentitas de la Formación Piedra
Shotle, Estancia la Casilda y en la perforación Cañadón
Pastos Blancos (YPF.Ch.CPB.es-1), Chubut, Argentina. 2◦
Congresso Latino-Americano de Paleontologı́a, Porto Alegre
1978, Actas, 1, 169–179.
Garcı́a, G. B. 1995. Palinologı́a de la Formación El Imperial,
Paleozoico Superior, Cuenca San Rafael, Argentina. Parte I:
esporas. Ameghiniana, 32, 315–339.
Garcı́a, G. B. 1996. Palinologı́a de la Formación El Imperial, Pale-
ozoico Superior, Cuenca San Rafael, República Argentina.
Parte II: granos de polen, incertae sedis, acritarcas. Amegh-
iniana, 33, 7–33.
Goubin, N. 1965. Description et répartition des principaux
Pollenites Permiens, Triasiques et Jurassiques des sondages
Français du Pétrole, 20, 1415–1461.
Guillemain, C. 1911. Zur Geologie Uruguays. Zeitschrift
Deutscher Geologischer Gesselschafte, 63, 202–220.
Gutiérrez, P. R. 1993. Palinologı́a de la Formación Agua Colorada
(Carbonı́fero Superior), sierra de Famatina, provincia de
La Rioja, Argentina. I. Granos de polen. Ameghiniana, 30,
163–212.
Gutiérrez, P. R. & Barreda, V. D. 2006. Palinologı́a de la
Formación El Trampeadero (Carbonı́fero Superior), La Rioja,
Argentina: significado bioestratigráfico. Ameghiniana, 43,
71–84.
Gutiérrez, P. R. & Césari, S. N. 1989. Nuevas microfloras de
la Formación Lagares (Carbonı́fero), provincia de La Rioja,
República Argentina. Ameghiniana, 25, 85–96.
 Lower Permian palynology of Paraná Basin
Gutiérrez, P. R. & Césari, S. N. 2000. Palinologı́a de la
Formación Bajo de Véliz (Pérmico Inferior), San Luis,
Argentina: revisión sistemática y consideraciones bioes-
tratigráficas. Ameghiniana, 37, 439–462.
Gutiérrez, P. R. & Limarino, C. O. 2001. Palinologı́a de
la Formación Malanzán (Carbonı́fero Superior), La Rioja,
Argentina: nuevos datos y consideraciones paleoambientales.
Ameghiniana, 38, 99–118.
Gutiérrez, P. R., Césari, S. N. & Archangelsky, S. 1997.
Deusilites tenuistriatus sp. nov. (acritarca) en el Pérmico Infe-
rior de la Cuenca Chacoparanense (Argentina). Ameghiniana,
34, 247–250.
Gutiérrez, P. R., Césari, S. N. & Archangelsky, S. 2002. Revisión
palinológica del pozo Árbol Blanco (YPF.SE.AB.x-1) (Pale-
ozoico Superior), Santiago del Estero, Cuenca Chacopara-
nense, Argentina. 8◦ Congreso Argentino de Paleontologı́a y
Bioestratigrafı́a, Corrientes, Abstracts, 72.
Gutiérrez, P. R., Di Pasquo, M. & Vergel, M. 2003. Palinoestrati-
grafı́a del Carbonı́fero-Pérmico de la Argentina: estado actual
del conocimiento. Revista del Museo Argentino de Ciencias
Downloaded By: [GutiérrezBuenos Aires, Pedro] At: 16:04 14 November 2010
Naturales, nueva serie, 5, 185–196.
Gutiérrez, P. R., Beri, A., Balarino, M. L. & Cernuschi, F. 2006.
Palinologı́a del Pérmico Inferior en la perforación CLS-24
(Cerro Largo Sur), departamento de Cerro Largo, Uruguay.
Parte II: granos de polen. Ameghiniana, 43, 611–635.
Hacquebard, P. A. 1957. Plants spores in coal from the Horton
Group (Mississippian) of Nova Scotia. Micropaleontology, 3,
301–324.
Hart, G. F. 1960. On spores and pollens from Permian deposits
of the Donets Basin. Witwatersrand University Press, Johan-
nesburg, 9 pp.
Hart, G. F. 1963. Microflora investigation of the Lower Coal
Measures (K2); Ketewaka-Mchuchuma Coalfield. Bulletin
Geological Survey Tanganyika, 30, 1–60.
Hart, G. F. 1964. A review of the classification and distribu-
tion of the Permian miospore: Disaccate Striatiti. 5◦ Congres
International du Stratigraphie et de Geologie du Carbonifere,
Compte Rendu, 3, 1171–1199.
Hart, G. F. 1965. The systematics and distribution of Permian
miospores. Witwatersrand University Press, Johannesburg,
432 pp.
Haskell, T. R. 1968. Saccate pollen grains from the Lower
Cretaceous of the Great Artesian Basin, Australia. Univer-
sity of Queensland, Department of Geology Papers, 6,
211–243.
Helby, R. 1973. Review of Late Permian and Triassic palynology
of New South Wales. Geological Society of Australia, Special
Publication, 4, 141–155.
Hemer, D. & Nygreen, P. 1967. Algae, acritarchs and other micro-
fossils incertae sedis from the Lower Carboniferous of Saudi
Arabia. Micropaleontology, 13, 183–194.
Høeg, O. A. & Bose, M. N. 1960. The Glossopteris flora of the
Belgian Congo with a note on some fossil plants from the
Zambesi basin (Mozambique). Annales de Musée Royal de
l’Afrique Centrale, Série 8, Sciences géologiques, 32:1–107.
Hoffmeister, W. S., Staplin, F. L. & Malloy, R. E. 1955. Missis-
sippian plant spores from the Hardinsburg Formation of Illi-
nois and Kentucky. Journal of Paleontology, 29, 372–399.
Ibrahim, A. C. 1933. Sporenformen des Aegirhorizonts des Ruhr-
Reviers. Konrad Triltsch, Würzburg, 47 pp.
Iversen, J. & Tröels-Smith, J. 1950. Pollenmorfologiske defini-
otioner og typer. Danmarks Geologiske Unders, 4, 1–54.
Jain, R. K. 1968. Middle Triassic pollen grains and spores from
Minas de Petroleo Beds of the Cacheuta Formation (Upper
Gondwana), Argentina. Palaeontographica B, 122, 1–47.
491
Jansonius, J. 1962. Palynology of Permian and Triassic sedi-
ments, Peace River area, western Canada. Palaeontographica
B, 110, 35–98.
Jansonius, J. 1971. Emended diagnosis of Alisporites Daugherty
1941. Pollen et Spores, 13, 349–357.
Jansonius, J. & Hills, L. V. 1976. Genera file of fossil spores.
Special Publication, Department of Geology, University of
Calgary, Canada, Card 1300.
Jansonius, J. & Hills, L. V. 1979. Genera file of fossil spores.
Special publication, Department of Geology, University of
Calgary, Canada, Card 3153.
Jansonius, J. & Hills, L. V. 1981. Genera file of fossil spores.
Special Publication, Department of Geology, University of
Calgary, Canada, Cards 2746 and 3916.
Jardiné, S. 1974. Microflores des Formations du Gabon attribuées
au Karroo. Review of Palaeobotany and Palynology, 17,
75–112.
Jones, M. J. & Truswell, E. M. 1992. Late Carboniferous and
Early Permian palynostratigraphy of the Joe Joe Group, south-
ern Galilee Basin, Queensland, and implications for Gond-
wanan stratigraphy. Bureau of Mines and Mineral Resources,
Journal of Australian Geology and Geophysics, 13, 143–
185.
Kar, R. K. 1968. Palynology of the Barren Measures sequence
from Jharia Coalfield, Bihar, India. 2. General palynology.
The Palaeobotanist, 16, 115–140.
Kar, R. K. & Bose, M. N. 1967. Palaeozoic Sporae Disper-
sae from Congo. III. Assise des schistes noirs de la Lukuga.
Annales de Musee Royal de l’Afrique Centrale, Serie 8,
Sciences Géologiques, 54, 2–84.
Kar, R. K. & Bose, M. N. 1976. Palaeozoic Sporae Dispersae
from Zaı̈re (Congo). XII. Assise a couches de houille from
Greinerville region. Annales de Musee Royal de l’Afrique
Centrale, Serie 8, Sciences Géologiques, 77, 21–133.
Klaus, W. 1960. Sporen der Karnischen Stufe der ostalpinen Trias.
Jahrbuch der Geologischen Bundesanstalt, Sonderband, 5,
107–184.
Klaus, W. 1963. Sporen aus dem südalpinen Perm. Jahrbuch der
Geologischen Bundensanstalt, 106, 229–363.
Klaus, W. 1964. Zur sporenstratigraphischen Einstufung von
gipsführenden Schichten in Bohrungen. Erdöl Zutschvift
Bohr-und Fördertechik, 4, 119–132.
Kosanke, R. M. 1950. Pennsylvanian spores of Illinois and their
use in correlation. Illinois State Geological Survey, Bulletin
74, 128 pp.
Kumar, P. 1973. The Sporae Dispersae of Jabalpur Stage, Upper
Gondwana, India. The Palaeobotanist, 20, 91–126.
Kützing, F. T. 1843. Phycologia generalis oder, Anatomie Physi-
ologie und Systemkunde der Tange. Brockhaus, Leipzig, 458
pp.
Kützing, F. T. 1849. Species algarum. Brockhaus, Leipzig, 922
pp.
Lakhanpal, R. N., Sah, S. C. D. & Dube, S. N. 1960. Further
observations on plant microfossils from a carbonaceous shale
(Krols) near Naini Tal, with a discussion on the age of the
beds. The Palaeobotanist, 7, 111–120.
Lele, K. M. 1964. Studies in the Talchir flora of India 2. Resolution
of the spore genus Nuskoisporites Pot. & Kl. The Palaeob-
otanist, 12, 147–168.
Lele, K. M. 1965. Studies in the Talchir flora of India. 3. Stel-
lapollenites - a new monosaccate pollen genus from the South
Rewa Gondwana basin. The Palaeobotanist, 13, 109–113.
Lele, K. M. 1975. Studies in the Talchir Flora of India. 10. Early
and Late Talchir microfloras from the West Bokaro Coalfield,
Bihar. The Palaeobotanist, 22, 219–235.
 492 P. R. Gutiérrez et al.
Lele, K. M. & Karim, R. 1971. Studies in the Talchir flora of
India. 6. Palynology of the Talchir Boulder Beds in Jayanti
Coalfield, Bihar. The Palaeobotanist, 13, 52–69.
Lele, K. M. & Makada, R. 1972. Studies in the Talchir Flora of
India. 7. Palynology of the Talchir Formation in the Jayanti
Coalfield, Bihar. Geophytology, 2, 41–73.
Leschik, G. 1955. Die Keuperflora von Neuewelt bei Basel. II.
Die Iso-und Mikrosporen. Schweizerische Paläontologische
Abhandlungen, 72, 1–70.
Leschik, G. 1956. Sporen aus dem Salzton des Zechsteins von
Neuhof (bei Fulda). Palaeontographica B, 100, 122–142.
Lindgren, S. 1980. Algal microfossils of the form genus Tetrapo-
rina from Upper Cretaceous clays, southern Sweden. Review
of Palaeobotany and Palynology, 30, 333–359.
Lindström, S. 1995. Early Permian palynostratigraphy of the
northern Heimefrontfjella mountain-range, Dronning Maud
Land, Antarctica. Review of Palaeobotany and Palynology,
89, 359–415.
Lindström, S. 1996. Late Permian palynology of Fossilryggen,
Vestfjella, Dronning Maud Land, Antarctica. Palynology, 20,
Downloaded By: [GutiérrezBuenos Aires, Pedro] At: 16:04 14 November 2010
15–48.
Llambias de Olivar, A. 1918. Informe acerca de la investi-
gación geológica efectuada en la República con el Dr. Arro-
jado Lisboa. Boletı́n Instituto de Geologı́a y Perforaciones, 3,
20–42.
Longhim, M. E., Souza, P. A. & Rohn, R. 2002. Palinologia
do Grupo Itararé na região de Salto (Carbonı́fero Superior),
Estado de São Paulo, Brasil. Parte 1- Palinologia Sistemática.
Revista Universidade Guarulhos, Geociências, 7, 43–60.
Luber, A. A. 1935. Les types petrographiques des charbons
fossiles du Spitsbergen Chimie. Combustible Solide, 5,
186–195.
Mackinon, J. 1969. Informe sobre estratigrafı́a del Gondwana en
la región noroeste de Uruguay. 1st International Symposium
Gondwana, Geology & Stratigraphy (Mar del Plata, 1967),
IUGS, Geology, 2, 973–983.
MacRae, C. S. 1988. Palynostratigraphic correlation between
the Lower Karoo sequence of the Waterberg and Pafuri
coal-bearing basins and the Hammanskraal plant macrofossil
locality, Republic of South Africa. Republic of South Africa
Memoir of the Geological Survey, 75, 1–217.
Mädler, K. 1964. Die geologische Verbreitung von Sporen und
Pollen in der Deutschen Trias. Beihefte zum Geologischen
Jahrbuch, 65, 1–147.
Maheshwari, H. K. 1967. Studies in the Glossopteris flora of
India—29. Miospore assemblage from the Lower Gondwana
exposures along Bansloi River in Rajmahal Hills, Bihar. The
Palaeobotanist, 15, 258–280.
Maheshwari, H. K. 1969. Palaeozoic Sporae Dispersae from
Congo. X. Microfossils from a cliff section at the confluence
of Lufupa and Mushyashya rivers, South Katanga. Annales
de Musee Royal de l’Afrique Centrale, Serie 8, Sciences
Géologiques, 63, 115–193.
Mandal, J. & Maithy, P. K. 1981. Palaeobotany of Tattitola Bed
in Pachwara Coalfield, Santhal Pargana, Bihar with remarks
on the age of the bed. The Palaeobotanist, 27, 12–20.
Mangerud, G. 1994. Palynostratigraphy of the Permian and
lowermost Triassic succession, Finnmark Platform, Barents
Sea. Review of Palaeobotany and Palynology, 82, 317–349.
Manum, S. 1960. On the genus Pityosporites Seward 1914, with
a new description of Pityosporites antarcticus Seward. Nytt
magasin for botanic, 8, 11–15.
Marques-Toigo, M. 1970. Anabaculites nov. gen., a new miospore
genus from San Gregorio Formation of Uruguay. Ameghini-
ana, 7, 79–82.
Marques-Toigo, M. 1972. Ammonoids vs. pollen and the
Carboniferous or Permian age of San Gregorio Formation
of Uruguay, Paraná basin. Anais da Academia Brasileira da
Ciências, 44 (suplemento), 237–241.
Marques-Toigo, M. 1974. Some new species of spores and
pollens of Lower Permian age from the San Gregorio Forma-
tion in Uruguay. Anais da Academia Brasileira da Ciências,
46, 601–616.
Marques-Toigo, M. 1991. Palynobiostratigraphy of the southern
Brazilian Neopalaeozoic Gondwana sequence. Pp. 503–515
in H. Ulbrich & A. C. Rocha-Campos (eds) Gondwana Seven
Proceedings, Instituto de Geociências, Universidade de São
Paulo.
Marques-Toigo, M. & Pons, M. E. 1976. Estudo palinológico do
furo de sondagem P7 Malha Oeste da bacia carbonı́fera do
Iruı́, RS, Brasil. 28th Congresso Brasleiro da Geologia, Porto
Alegre, Annais 1974, 277–288.
Marques-Toigo, M. & Piccarelli, A. T. 1985. On the morphol-
ogy and botanical affinities of Lundbladispora Balme 1963 in
the Permian of the Paraná Basin, Brazil. Boletim Instituto de
Geociencias, Universidade de São Paulo, 15, 46–52.
Marques-Toigo, M., Dias-Fabricio, M. E., Guerra-Sommer,
M., Cazzulo-Klepzig, M. & Piccoli, A. E. M. 1990. Aflo-
ramentos da área de Trombudo Central, Permiano Infe-
rior, Santa Catarina: palinologia, icnologı́a e sedimentologia.
11th Congresso Brasleiro da Paleontologia, Curitiba, Annais,
1989, 125–150.
Martı́nez-Macchiavello, I. C. 1963. Microesporomorfos tipos
contenidos en el glacial en la base del Sistema de Gondwana
en Uruguay. Boletim da Universidade de Paraná, Geologia,
10, 1–4.
Mautino, L. R., Anzótegui, L. M. & Vergel, M. M. 1998a. Pali-
nologı́a de la Formación Melo (Pérmico Inferior) en Arroyo
Seco, Departamento Rivera, República Oriental del Uruguay.
Parte IV: esporas. Ameghiniana, 35, 67–79.
Mautino, L. R., Vergel, M. M. & Anzótegui, L. M. 1998b.
Palinologı́a de la Formación Melo (Pérmico Inferior) en
Arroyo Seco, departamento Rivera, Uruguay. Parte V: Granos
de polen, acritarcas e incertae sedis. Ameghiniana, 35,
299–314.
Mautino, L. R., Vergel, M. M. & Anzótegui, L. M. 1998c. Pali-
nologı́a de la Formación Melo (Pérmico inferior) en Arroyo
Seco, Departamento Rivera, República Oriental del Uruguay,
Parte III. Especies nuevas. Revista Española de Micropaleon-
tologı́a, 30, 107–110.
Menéndez, C. A. 1971. Estudio Palinológico del Pérmico de
Bajo de Vélez, provincia de San Luis. Revista del Museo
Argentino de Ciencias Naturales “Bernardino Rivadavia” e
Instituto Nacional de Investigación de las Ciencias Naturales,
Paleontologı́a, 1, 263–306.
Menéndez, C. A. 1976. Contenido palinológico de estratos
pérmicos con “Mesosaurus” de Rio Claro, Sâo Paulo,
Brasil. Revista del Museo Argentino de Ciencias natu-
rales “Bernardino Rivadavia” e Instituto Nacional de
Investigación de las Ciencias Naturales, Paleontologı́a, 2,
1–30.
Menéndez, C. A. & Azcuy, C. L. 1969. Microflora carbónica
de la localidad de Paganzo, Provincia de la Rioja, Parte I.
Ameghiniana, 6, 77–97.
Menéndez, C. A. & Azcuy, C. L. 1971. Microflora carbónica
de la localidad de Paganzo, Provincia de la Rioja, Parte II.
Ameghiniana, 8, 25–36.
Menéndez, C. A. & Azcuy, C. L. 1973. Microflora carbónica
de la localidad de Paganzo, Provincia de La Rioja, Parte III.
Ameghiniana, 10, 51–71.
 Lower Permian palynology of Paraná Basin
Mildenhall, D. C. 1994. Palynological reconnaissance of early
Cretaceous to Holocene sediments,Chatham Islands, New
Zealand. Institute of Geological and Nuclear Sciences Mono-
graph, 7, 1–206.
Morbey, S. J. 1975. The palynostratigraphy of the Rhaetian Stage,
Upper Triassic in the Kendelbachgraben, Austria. Palaeonto-
graphica B, 152, 1–75.
Muff, R., González, M. E., Vergel, M. M., Herbst, R. &
Fasolo, Z. 1999. Palynological investigations and biostrati-
graphic correlations of Permian sedimentary rocks from east-
ern Paraguay. Newsletters on Stratigraphy, 37, 21–35.
Nahuys, J., Alpern, B. & Ybert, J. P. 1968. Estudo palinológico
e petrográfico de alguns carvôes do sul do Brasil. Instituto
Tecnológico do Rio Grande do Sul, Porto Alegre, Boletim, 46,
1–61.
Naumova, S. N. 1953. Sporovo-pyltsevye kompleksy verkhnego
devona Russkoi platformy I ikh znachenie dlya stratigrafii
(Complexes sporo-polliniques du Dévonien supérieur de la
plateforme russe et leur valeur stratigraphique). Trudy Insti-
tuta Geologischeskikh Nauk, Akademiya Nauk SSSR 143,
Downloaded By: [GutiérrezBuenos Aires, Pedro] At: 16:04 14 November 2010
Seriya Geologischeskaya, 60, 1–204 (in Russian).
Neves, R. & Owens, B. 1966. Some Namurian camerate
miospores from the English Pennines. Pollen et Spores, 8,
337–360.
Nilsson, T. 1958. Uber das Vorkommen eines mesozoischen
Sapropelgesteins in Schonen. Lunds Universitets Arsskrift,
54, 1–112.
Ottone, E. G. 1989. Palynoflores de la Formation Santa Máxima,
Paléozoı̈que supérieur, République Argentine. Palaeonto-
graphica B, 213, 89–187.
Ottone, E. G. 1991. Palynologie du Carbonifère supérieur de la
coupe de Mina Esperanza, Bassin Paganzo, Argentine. Revue
de Micropaléontologie, 34, 118–135.
Ottone, E. G. & Azcuy, C. L. 1990. Datos palinológicos de
la Formación Guandacol (Carbonı́fero) en la quebrada La
Delfina, provincia de San Juan, Argentina. Ameghiniana, 26
[for 1989], 191–208.
Ottone, E. G. & Garcı́a, G. B. 1991. A Lower Triassic miospore
assemblage from the Puesto Viejo Formation, Argentina.
Review of Palaeobotany and Palynology, 68, 217–232.
Ottone, E. G., Álvarez, P. P. & Benoit, S. V. 1992. Late Trias-
sic plant microfossils from the Rancho de Lata Formation,
Main Cordillera, Argentina. Micropaleontology, 38, 261–
278.
Ouyang Shu & Norris, G. 1999. Earliest Triassic (Induan) spores
and pollen from the Junggar Basin, Xinjiang, northwestern
China. Review of Palaeobotany and Palynology, 106, 1–56.
Pascher, A. 1914. Über Flagellaten und Algen. Veröffentlichung
Deutsche Botanische Gesellschaft, 32, 136–160.
Pant, D. D. 1954. Suggestions for the classification and nomencla-
ture of fossil spores and pollen grains. The Botanical Review,
20, 33–60.
Pant, D. D. 1955. On two new disaccate spores from the Bacchus
Marsh Tillite, Victoria (Australia). Annals and Magazine of
Natural History, 8, 757–764.
Papú, O. 1991. Contribución a la palinologı́a estratigráfica de la
Formación Malargüe, Cretácico Superior, sur de la provin-
cia de Mendoza, Argentina. 1: especies terrestres y de aguas
continentales. Ameghiniana, 27 [for 1990], 289–303.
Pérez Loinaze, V. & Césari, S. 2004. Palynology of the
Estratos de Mascası́n, Upper Carboniferous, Paganzo Basin,
Argentina: systematic description and stratigraphic considera-
tions. Revista Española de Micropaleontologı́a, 36, 407–438.
Playford, G. 1965. Plant microfossils from Triassic sediments
near Poatina, Tasmania. Journal of the Geological Society of
Australia, 12, 173–210.
493
Playford, G. 1978. Lower Carboniferous spores from the
Ducabrook Formation, Drummond Basin, Queensland.
Palaeontographica B, 167, 105–160.
Playford, G. 1990. Proterozoic and Paleozoic Palynology of
Antarctica: A Review. Pp. 51–70 in T. N. Taylor & E. L.
Taylor, (eds) Antarctic Paleobiology: Its Role in the Recon-
struction of Gondwana, Springer, New York.
Playford, G. & Dettmann, M. E. 1996. Spores. Pp. 227–260
in J. Jansonius & D. C. McGregor (eds) Palynology: Prin-
ciples and Applications, Volume I. American Association of
Stratigraphic Palynologists Foundation, Dallas.
Playford, G. & Dino, R. 2000a. Palynostratigraphy of upper
Palaeozoic strata (Tapajós Group), Amazonas Basin, Brazil:
Part One. Palaeontographica B, 255, 1–46.
Playford, G. & Dino, R. 2000b. Palynostratigraphy of upper
Palaeozoic strata (Tapajós Group), Amazonas Basin, Brazil:
Part two. Palaeontographica B, 255, 87–145.
Playford, G. & Dino, R. 2002. Permian Palynofloral assem-
blages of the Chaco-Paraná Basin, Argentina: Systematics
and Stratigraphic significance. Revista Española de Micropa-
leontologı́a, 34, 235–288.
Playford, G. & Powis, G. D. 1979. Taxonomy and distribution
of some trilete spores in Carboniferous strata of the Canning
Basin, Western Australia. Pollen et Spores, 21, 371–394.
Playford, G., Dino, R. & Marques-Toigo, M. 2001. The Upper
Paleozoic miospore genus Spelaeotriletes Neves and Owens,
1966, and constituent Gondwanan species. Journal of South
American Earth Sciences, 14, 593–608.
Pocock, S. A. J. 1970a. Palynology of the Jurassic sediments of
Western Canada. Part 1. Terrestrial species. Palaeontograph-
ica B, 130, 12–72.
Pocock, S. A. J. 1970b. Palynology of the Jurassic sediments
of Western Canada. Part 1 (continued). Terrestrial species.
Palaeontographica B, 130, 73–136.
Pons, M. E. 1977. Estudio palinológico do Sub-grupo Itararé na
“Coluna White”, Permiano Inferior, Santa Catarina, Brasil. I
parte. Ameghiniana, 13 [for 1976], 109–125.
Pons, M. E. 1979. Estudo palinológico do Sub-grupo Itararé, na
“Coluna White” Permiano Inferior, Santa Catarina, Brasil. III
Parte. Ameghiniana, 14 [for 1977], 87–99.
Potonié, R. 1931. Pollenformen der miozänen Braunkohlen.
Sitzungsberichte der Gesellschaft Naturforschender Freunde
zu Berlin, 52, 24–36.
Potonié, R. 1956. Synopsis der Gattungen der Sporae Dispersae.
I. Teil: Sporites. Geologisches Jahrbuch Beihefte, 23, 1–103.
Potonié, R. 1958. Synopsis der Gattungen der Sporae Dispersae.
Teil 2. Sporites (Nachträge), Saccites, Aletes, Praecolpates,
Polyplicates, Monocolpates. Geologisches Jahrbuch Beihefte,
31, 1–114.
Potonié, R. 1960. Synopsis der Gattungen der Sporae Disper-
sae. Teil III. Nachträge Sporites, Fortsetzung Pollenites. Mit
Generalregister zu Teil I-III. Geologisches Jahrbuch Beihefte,
39, 1–189.
Potonié, R. 1965. Fossile Sporae in situ. Forschungsberichte des
Landes Nordrhein-Westfalen, 1483, 1–74.
Potonié, R. 1970. Synopsis der Gattungen der Sporae Dispersae.
Teil V. Nachträge zu allen Gruppen (Turmae). Geologisches
Jahrbuch Beihefte, 87, 1–222.
Potonié, R. & Klaus, W. 1954. Einige Sporengattungen des
alpinen Salzgebirges. Geologisches Jahrbuch, 68, 517–546.
Potonié, R. & Kremp, G. 1954. Die Gattungen der paläozois-
chen Sporae Dispersae und ihre Stratigraphie. Geologisches
Jahrbuch, 69, 111–194.
Potonié, R. & Lele, K. M. 1961. Studies in the Talchir flora of
India-Sporae Dispersae from the Talchir beds of South Rewa
Gondwana basin. The Palaeobotanist, 8, 22–37.
 494 P. R. Gutiérrez et al.
Potonié, R. & Sah, S. C. D. 1960. Sporae Dispersae of the lignites
from Cannanore Beach on the Malabar Coast of India. The
Palaeobotanist, 7, 121–135.
Premaor, E., Fischer, T. V. & Souza, P. A. 2006. Palinologı́a fa
Formaçâo Irati (Permiano Inferior da Bacia do Paraná), em
Montividiu, Goiás, Brasil. Revista del Museo Argentino de
Ciencias Naturales, nueva serie, 8, 221–230.
Price, P. L. 1983. A Permian palynostratigraphy for Queensland.
Pp. 155–211 in Proceedings of the Symposium on the Permian
geology of Queensland, Brisbane 1982. Geological Society of
Australia, Queensland Division, Brisbane.
Preciozzi, F., Spoturno, J., Heinzen, W. & Rossi, P. 1985.
Carta Geológica del Uruguay y Memoria Explicativa a escala
10500.00. Dirección Nacional de Minerı́a y Geologı́a. Monte-
video, Uruguay.
Quadros, L. P. de, Marques-Toigo, M. & Cazzulo-Klepzig, M.
1996. Catálogo de esporos e pólen fósseis do Paleozóico.
Boletim de Geociências da Petrobras, 9, 1–151.
Raine, J. I., Mildenhall, D. C. & Kennedy, E. M. 2005.
New Zealand fossil spores and pollen: an illustrated
Downloaded By: [GutiérrezBuenos Aires, Pedro] At: 16:04 14 November 2010
catalogue. Institute of Geological and Nuclear Sciences
Information Series, 68, version 1. http://www.gns.cri.nz/
what/earthhist/fossils/spore pollen/catalog/index.htm.
Raine, J. I., Mildenhall, D. C. & Kennedy, E. M. 2006. New
Zealand fossil spores and pollen: an illustrated catalogue. 2nd
edition. GNS Science Miscellaneous Series, 4. http://www.
gns.cri.nz/what/earthhist/fossils/spore pollen/catalog/index.
htm.
Ravn, R. L. 1979. An introduction to the stratigraphic palynology
of the Cherokee Group (Pennsylvanian) coals of Iowa. Iowa
Geological Survey, Technical Paper, 6, 117 pp.
Richardson, J. B. 1965. Middle Old Red Sandstone spore assem-
blages from the Orcadian Basin northeast Scotland. Palaeon-
tology, 7, 559–605.
Rigby, J. F. & Hekel, H. 1977. Palynology of the Permian
sequence in the Springsure anticline, central Queensland.
Geological Survey of Queensland Publication, Palaeontolog-
ical Papers, 37, 1–76.
Rojo, L. D. & Zavattieri, A. M. 2005. Estudio microfloristico de
las Formaciones Potrerillos y Cacheuta (Triásico) en el sur del
cerro Cacheuta, Mendoza, Argentina. Parte 1. Ameghiniana,
42, 3–20.
Round, F. E. 1971. The taxonomy of the Chlorophyta II. British
Phycological Journal, 6, 235–264.
Russo, A., Archangelsky, S. & Gamerro, J. C. 1980. Los
depósitos suprapaleozoicos en el subsuelo de la Llanura
Chaco-Pampeana, Argentina. 2nd Congreso Argentino de
Paleontologı́a y Bioestratigrafı́a y 1st Congreso Latinoamer-
icano de Paleontologı́a, Buenos Aires, 1978, 4, 157–
173.
Sah, S. C. D. & Jain, K. P. 1965. Jurassic spores and pollen grains
from the Rajmahal Hills, Bihar, India: with a discussion on the
age of the Rajmahal Intertrappean Beds. The Palaeobotanist,
13, 264–290.
Samoilovich, S. R. 1953. [Pollen and spores from the
Permian deposits of the Cherdyn’ and Aktyubinsk areas,
Cis-Urals]. Trudy Vsesoiuznyi Nauchno-issledovatel’skii
Geologo-razvedochnyi Institut, new series, Leningrad, 75,
5–57 [in Russian; translated by M.K. Elias (1961. Oklahoma
Geological Survey Circular, 56, 7–103)].
Saxena, R. K. 1993. New names for some palynofossil later
homonyms from India. Geophytology, 23, 195–196.
Schaarschmidt, F. 1963. Sporen und Hystrichosphaerideen aus
dem Zechstein von Büdingen in der Wetteran. Palaeonto-
graphica B, 113, 38–91.
Scheuring, B. W. 1970. Palynologische und palynostrati-
graphische Untersuchungen des Keupers im Bölchentunnel
(Solothurner Jura). Schweizerische Paläontologische Abhand-
lungen, 88, 1–199.
Schopf, J. M., Wilson, L. R. & Bentall, R. 1944. An anno-
tated synopsis of Paleozoic fossil spores and the definition
of generic groups. Illinois State Geological Survey Report of
Investigations, 91, 1–73.
Segroves, K. L. 1967. Cutinized microfossils of probable nonvas-
cular origin from the Permian of Western Australia. Micropa-
leontology, 13, 289–305.
Segroves, K. L. 1969. Saccate plant microfossils from the Permian
of Western Australia. Grana Palynologica, 9, 174–227.
Singh, C. 1971. Lower Cretaceous microfloras of the Peace River
area, northwestern Alberta. Research Council of Alberta,
Bulletin, 28, 1–542.
Sinha, V. 1972. Sporae Dispersae from Jhingurdah Seam,
Singrauli Coalfield (M.P.), India. The Palaeobotanist, 19,
175–201.
Smaniotto, L. P., Fisher, T. V., Souza, P. A. & Ianuzzi, R. 2006.
Palinologia do Morro do Papaléo, Marina Pimentel (Permiano
inferior, Bacia do Paraná), Rio Grande Sul, Brasil. Revista
Brasileira de Paleontologia, 9, 311–322.
Smith, A. H. V. & Butterworth, M. A. 1967. Miospores in the
coal seams of the Carboniferous of Great Britain. Special
Papers in Palaeontology, 1, 1–324.
Somers, G. 1952. A preliminary study of the fossil spore content of
the Lower Jubilee seam of the Sydney coalfield, Nova Scotia.
Nova Scotia Research Foundation, Halifax, 30 pp.
Souza, P. A. & Callegari, L. M. 2004. An Early Permian paly-
noflora from the Itararé Subgroup, Paraná Basin, Brazil.
Revista Española de Micropaleontologı́a, 36, 439–450.
Souza, P. A. & Marques-Toigo, M. 2001. Zona Vittatina: marco
palinobioestratigrafico do Permiano Inferior da Bacia do
Paraná. Ciência Ténica Petróleo, 20, 153–159.
Souza, P. A. & Marques-Toigo, M. 2003. An overview on the
palynostratigraphy of the Upper Paleozoic strata of the Brazil-
ian Paraná Basin. Revista de Museo Argentino de Ciencias
Naturales, nueva serie, 5, 205–214.
Souza, P. A. & Marques-Toigo, M. 2005. Progress on the paly-
nostratigraphy of the Permian strata in Rio Grande do Sul
State, Paraná Basin, Brazil. Anais da Academia Brasileira de
Ciencias, 77, 353–365.
Souza, P. A., Saad, A. R. & Lima, M. R. 1997. Palinologı́a
dos carvôes paleozicos de Estado do Sâo Paulo, Brasil. II-O
Carvâo de Monte Mor. Revista do Instituto Geológico, Sâo
Paulo, 18, 7–21.
Souza, P. A., Petri, S. & Dino, R. 2003. Late Carboniferous paly-
nology from the Itararé Subgroup (Paraná Basin) at Araçoiaba
da Serra, São Paulo State, Brazil. Palynology, 27, 39–74.
Srivastava, S. C. 1970. Miofloral investigations in some coals
of Talcher Coalfield (Orissa) India. The Palaeobotanist, 18,
154–166.
Stapleton, R. P. 1978. Microflora from a possible Permo-Triassic
transition in South Africa. Review of Palaeobotany and Paly-
nology, 25, 253–258.
Stephenson, M. H. 2004. Early Permian spores from Oman
and Saudi Arabia. Pp. 185–215 in M. I. Al-Husseini (ed.)
Carboniferous, Permian and Early Triassic Arabian Stratig-
raphy, GeoArabia Special Publication, 3, Gulf PetroLink,
Bahrain.
Stephenson, M. H. & Filatoff, J. 2000a. Correlation of
Carboniferous-Permian palynological assemblages from
Oman and Saudi Arabia. Pp. 168–191 in S. Al-Hajri &
B. Owens (eds) Stratigraphic Palynology of the Palaeozoic
 Lower Permian palynology of Paraná Basin
of Saudi Arabia, GeoArabia Special Publication, 1, Gulf
PetroLink, Bahrain.
Stephenson, M. H. & Filatoff, J. 2000b. Description and corre-
lation of Late Permian palynological assemblages from the
Khuff Formation, Saudi Arabia and evidence for the dura-
tion of the pre-Khuff Hiatus. Pp. 192–215 in S. Al-Hajri
& B. Owens, (eds) Stratigraphic Palynology of the Palaeo-
zoic of Saudi Arabia, GeoArabia Special Publication, 1, Gulf
PetroLink, Bahrain.
Stephenson, M. H. & Osterloff, P. L. 2002. Palynology of
the deglaciation sequence represented by the Lower Permian
Rahab and Lower Gharif Members, Oman. AASP, Contribu-
tion Series, 40, 1–41.
Stephenson, M. H., Osterloff, P. L. & Filatoff, J. 2003. Palyno-
logical biozonation of the Permian of Oman and Saudi Arabia:
progress and challenges. GeoArabia, 8, 467–496.
Streel, M. 1964. Une association de spores de Givétien inférieur
de la Vesdre, à Goé (Belgique). Annales de la Société
Géologique de Belgique, 87, 1–30.
Thomson, P. W. & Pflug, H. D. 1953. Pollen und Sporen des
Downloaded By: [GutiérrezBuenos Aires, Pedro] At: 16:04 14 November 2010
mitteleuropaischen Tertiars. Palaeontographica B, 94, 1–138.
Tiwari, R. S. 1965. Miospore assemblage in some coals of Barakar
Stage (Lower Gondwana) of India. The Palaeobotanist, 13,
168–214.
Tiwari, R. S. 1973. Scheuringipollenites, a new name for the
Gondwana sporomorphs so far assigned to “Sulcatisporites
Leschik 1955”. Senckenbergiana Lethaea, 54, 105–
117.
Tiwari, R. S. & Navale, G. K. B. 1967. Pollen and spore assem-
blage in some coals of Brazil. Pollen et Spores, 9, 583–
605.
Tiwari, R. S. & Rana, V. 1981. Sporae Dispersae of some Lower
and Middle Triassic sediments from Damodar Basin, India.
The Palaeobotanist, 27, 190–220.
Taugourdeau-Lantz, J. 1974. Premiere étude des spores du Trias
Moyen de Gabia (Bordure sud de la Montagne Noire, France).
Review of Palaeobotany and Palynology, 17, 149–159.
Tschudy, R. H. & Kosanke, R. M. 1966. Early Permian vesiculate
pollen from Texas, U.S.A. The Palaeobotanist, 15, 59–71.
Urban, J. B. 1971. Palynology of the Independence Shale of Iowa.
Bulletins of American Paleontology, 60, 103–189.
Venkatachala, B. S. & Kar, R. K. 1966a. Divarisaccus gen. nov.,
a new saccate pollen genus from the Permian sediments of
India. The Palaeobotanist, 15, 102–106.
Venkatachala, B. S. & Kar, R. K. 1966b. Corisaccites gen. nov.,
a new saccate pollen genus from the Permian of Salt Range,
West Pakistan. The Palaeobotanist, 15, 107–109.
Vergel, M. M. 1986. Palinologı́a del Paleozoico Superior en
la perforación YPF J1 (Josefina), provincia de Santa Fe,
Argentina. I. Anteturma Proximegerminantes. Ameghiniana,
23, 141–153.
Vergel, M. M. 1987a. Contenido palinológico de la Formación
Melo (Paleozoico Superior) de Melo, Uruguay. 10◦
Congreso Geológico Argentino (Tucumán), Actas, 3, 117–
120.
Vergel, M. M. 1987b. Palinologı́a del Paleozoico Superior en
la perforación YPF SF J (Josefina), Provincia de Sante Fe,
Argentina. II. Anteturma Variegerminantes, Grupo Acritar-
cha, e Incertae Sedis. Ameghiniana, 24, 67–80.
Vergel, M. M. 1993. Palinoestratigrafı́a de la secuencia neopa-
leozoica en la Cuenca Chacoparanense, Argentina. 12th
Congrès International de la Stratigraphie et Géologie du
Carbonifère et Permien (Buenos Aires 1991), Comptes
Rendus, 1, 201–212.
495
Vergel, M. M. 1998. Palinologı́a del Paleozoico Superior
(Formación Sachayoj) en tres perforaciones de la Subcuenca
de Alhuampa, Cuenca Chacoparanense (Argentina), Parte 1:
esporas. Ameghiniana, 35, 387–403.
Veroslavsky, G., Daners, G. & de Santa Ana, H. 2003.
Rocas sedimentarias pérmicas de la plataforma continental
uruguaya: el prerift de la Cuenca de Punta del Este. Geogac-
eta, 34, 203–206.
Volkheimer, W. & Papu, O. H. 1993. Una microflora del Triásico
Superior de la Cuenca de Malargüe, localidad Llantenes,
Provincia de Mendoza, Argentina. Ameghiniana, 30,
93–100.
Walther, K. 1919. Lı́neas fundamentales de la estructura
geológica de la República Oriental del Uruguay. Revista del
Instituto Nacional de Agronomı́a, Serie 2, 3, 1–186.
Waltz, J. E. 1935 in Luber, A. A. & Waltz, J. E. 1938
(eds). Classification and stratigraphical value of spores
and some Carboniferous coal deposits in the U.S.S.R.,
Trudy Tsentral’nogo Nauchno-Issledovatel’skogo Geologo-
Razvedochnogo Instituta 105, 1–45 (in Russian).
Weiss, R. H. 2001. Middle to Late Permian microfloral assem-
blages from the Ruhuhu and Selous basins, Tanzania. Pp.
497–529 in R. H. Weiss (ed.) Contributions to Geology and
Palaeontology of Gondwana -In Honour of Helmut Wopfner.
Geological Institute, University of Cologne, Cologne.
Wilson, L. R. 1962. Permian plant microfossils from the Flower-
pot Formation, Greev County, Oklahoma. Oklahoma Geolog-
ical Survey, Circular, 49, 1–50.
Ybert, J. P. 1975. Etude des miospores du Bassin Houiller de
Candiota-Hulha Negra, Rio Grande do Sul, Bresil. Pesquisas,
5, 181–226.
Ybert, J. P. & Marques-Toigo, M. 1971. Polarisaccites nov. gen.
Pollen et Spores, 12, 468–481.
Zavattieri, A. M. 1986. Estudio palinologico de la Formación
Potrerillos (Triásico) en su localidad tipo, Cuenca Cuyana
(Provincia de Mendoza, Argentina). Parte 1: Esporas triletes
y monoletes. Revista Española de Micropaleontologı́a, 18,
247–294.
Zavattieri, A. M. 1987. Estudio palinológico de la Formación
Potrerillos (Triásico) en su localidad tipo, Cuenca Cuyana
(provincia de Mendoza, Argentina). Parte 2: Granos de
polen. Aspectos estadı́sticos. Correlación palinoestratigráfica.
Revista Española de Micropaleontologı́a, 19, 173–213.
Zavattieri, A. M. 1991a. Granos de polen de la Formación
Las Cabras (Triásico), en su localidad Tipo, Provincia de
Mendoza, Argentina. Parte 1. Ameghiniana, 28, 3–29.
Zavattieri, A. M. 1991b. Palinologı́a de la Formación Las Cabras
(Triásico) en su localidad tipo, Cuenca Cuyana (provincia de
Mendoza, Argentina). Parte 2. Ameghiniana, 28, 205–224.
Zavattieri, A. M. 1992. Palinologia de la Formación El Tranquilo
(Triasico), Provincia de Santa Cruz, Argentina. Ameghiniana,
29, 305–314.
Zavattieri, A. M. & Rojo, L. D. 2005. Estudio microflorı́stico de
las Formaciones Potrerillos y Cacheuta (Triásico) en el sur del
cerro Cacheuta, Mendoza, Argentina. Parte 2. Ameghiniana,
42, 513–534.
Zavattieri, A. M. & Volkheimer, W. 1992. Granos de polen
sacados (Saccites) de la Formación Potrerillos (Triásico) en
la Localidad de Divisadero Largo, provincia de Mendoza,
Argentina. Ameghiniana, 29, 27–44.
Zavattieri, A. M., Herbst, R. & Muñoz Bravo, J. 2003.
Microflora de la Formación Panguipulli (Triásico Superior),
en Licán Ray, Lago Calafquén, 10◦ Región, Chile. Ameghini-
ana, 40, 585–600.
 496 P. R. Gutiérrez et al.
Appendix 1. List of taxa recorded from CDLS wells 201 and 221,
and from the locality El Barón, with notes on their distribution. For
detailed stratigraphic distribution within individual DCLS well see
Supplementary Data
Anteturma Proximegerminantes R. Potonié, 1970
Turma Triletes Reinsch, emend. Dettmann, 1963
Suprasubturma Acavatitriletes Dettmann, 1963
Subturma Azonotriletes Luber, emend. Dettmann, 1963
Infraturma Laevigati Bennie & Kidston, emend. R. Potonié, 1956
Genus Calamospora Schopf, Wilson & Bentall, 1944
Calamospora tener (Leschik) de Jersey, 1962 (Fig. 7A)
DCLS wells 201, 221. Well-known from Triassic-Jurassic
rocks of Europe, Greenland, Canada, Australia, Antarctica,
Chile and Argentina (see Zavattieri 1986; Ottone & Garcı́a
1991; Zavattieri et al. 2003). First record in Uruguay.
Genus Leiotriletes Naumova, emend. Potonié & Kremp, 1954
Leiotriletes corius Kar & Bose, 1967 (Fig. 7B)
DCLS wells 201, 221. Permian of Africa (Høeg & Bose
1960; Kar & Bose 1967); Lower Permian of Uruguay (Beri
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et al. 2006); Pennsylvanian of Argentina (Césari & Gutiérrez
1985, 2001; Gutiérrez & Césari 1989).
Leiotriletes directus Balme & Hennelly, 1956 (Fig. 7D)
DCLS wells 201, 221. Widely known from Permian to
Jurassic strata of the world (see Raine et al. 2005). Lower
Permian of Uruguay (Mautino et al. 1998a; Beri et al.
2006). Argentina: Pennsylvanian-Lower Permian (Gutiérrez
& Césari 2000; Césari & Gutiérrez 2001; di Pasquo et al.
2001) and Triassic (Rojo & Zavattieri 2005).
Leiotriletes tiwarii Saxena, 1993 (Fig. 7C)
DCLS well 201. Known widely from Permian to Jurassic
strata of world (see Raine et al. 2005). Lower Permian of
Uruguay (Beri 1987; Mautino et al. 1998a; Beri & Goso
1996; Beri et al. 2006) and Brazil (Quadros et al. 1996).
Saxena (1993) replaced Leiotriletes virkki (Tiwari: 170–171,
pl. 1, figs 2–3) with Leiotriletes tiwarii, a name previously
proposed by Biswas (1962: 43, pl. 9, fig. 53).
Genus Punctatisporites Ibrahim, emend. Potonié & Kremp, 1954
Punctatisporites gretensis Balme & Hennelly, 1956
(Fig. 7E)
DCLS well 201. Widely known from Early Permian strata
from Gondwanaland (see Foster 1975; Rigby & Heckel
1977). Lower Permian of Uruguay (Beri 1988; Beri & Daners
1995, 1998; Beri & Goso 1996; Mautino et al. 1998a; Beri
et al. 2006).
Infraturma Retusotrileti Streel, 1964
Genus Retusotriletes Naumova, emend. Streel, 1964
Retusotriletes simplex Naumova, 1953 (Fig. 7F)
DCLS well 201. Widely known from Palaeozoic strata from
Worldwide; Lower Permian from Uruguay (Beri et al. 2006).
Subinfraturma Granulati Dybová & Jachowicz, 1957
Genus Cyclogranisporites Potonié & Kremp, 1954
Cyclogranisporites gondwanensis Bharadwaj & Salujha, 1964
(Fig. 7H)
DCLS well 201. Permian from Africa, India, Antarctica and
Australia (Bharadwaj 1962; Bharadwaj & Salujha 1964;
Tiwari 1965, Bose & Kar 1966; Balme & Playford 1967;
Srivastava 1970; Sinha 1972; Anderson 1977; Rigby &
Heckel 1977; Foster 1979; Bharadwaj & Dwivedi 1981;
Mandal & Maithy 1981; Tiwari & Rana 1981; Crosbie
1985; Banerjee 1988). Lower Permian of Brazil (Ybert
1975). This is the first record in Uruguay.
Genus Granulatisporites Ibrahim, emend. Potonié & Kremp, 1954
Granulatisporites austroamericanus Archangelsky &
Gamerro, 1979 (Fig. 7I)
DCLS well 201. Permian from South America (see Playford
& Dino 2002). Lower Permian from Uruguay (Mautino
et al. 1998a; Beri et al. 2000, 2006).
Subinfraturma Nodati Dybová & Jachowicz, 1957
Genus Anapiculatisporites Potonié & Kremp, 1954
Anapiculatisporites tereteangulus (Balme & Hennelly) Play-
ford & Dino, 2002 (Fig. 7L)
DCLS well 201. Permian Gondwana strata (Balme &
Hennelly 1956; Balme 1970; Anderson 1977; Rigby &
Heckel 1977; Foster 1979; Backhouse 1991; Lindström
1995; Stephenson 2004). Lower Permian of Uruguay
(Mautino et al. 1998a; Beri & Pecoits 2001; Beri et al.
2006).
Genus Brevitriletes Balme & Hennelly, emend. Bharadwaj &
Srivastava, 1969
Brevitriletes cornutus (Balme & Hennelly) Backhouse, 1991
(Fig. 7K)
DCLS well 201. Permian from Africa and Australia (see
Stephenson 2004; Raine et al. 2005). Late Carboniferous-
Early Permian from Argentina, Brazil, Saudi Arabia and
Oman (Ybert 1975; Archangelsky & Gamerro 1979;
Quadros et al. 1996; Vergel 1998; Gutiérrez & Césari 2000;
Césari & Gutiérrez 2001; Cazzulo-Klepzig et al. 2002;
Stephenson 2004). Lower Permian from Uruguay (Beri
1987; Beri et al. 2006).
Brevitriletes levis (Balme & Hennelly) Bharadwaj & Srivas-
tava, 1969 (Fig. 7G)
DCLS well 201. Permian of Gondwanaland (Raine et al.
2005). Pennsylvanian-Lower Permian of Argentina and
Brazil (Menéndez & Azcuy 1969, 1971; Ybert 1975; Souza
et al. 1997, 2003; Vergel 1998; Césari & Gutiérrez 2001;
Dino & Playford 2002; di Pasquo et al. 2003a; Balarino &
Gutiérrez 2006); Lower Permian of Brazil (Smaniotto et al.
2006) and Uruguay (Fasolo & Vergel 1994; Mautino et al.
1998a; Beri et al. 2006).
Subinfraturma Verrucati Dybová & Jachowicz, 1957
Genus Converrucosisporites Potonié & Kremp, 1954
Converrucosisporites confluens (Archangelsky & Gamerro)
Playford & Dino, 2002 (Fig. 7U)
DCLS well 201. Lower Permian of Gondwanaland (Back-
house 1991; Lindström 1995; Stephenson 2004). Lower
Permian of Argentina (Archangelsky & Gamerro 1979;
Gamerro & Archangelsky 1981; Césari et al. 1995; Césari
& Gutiérrez 2001; Playford & Dino 2002), Brazil (Souza
& Calegari 2004) and Uruguay (Verolasky et al. 2003; Beri
et al. 2006).
Converrucosisporites micronodosus (Balme & Hennelly)
Playford & Dino, 2002 (Fig. 7P)
DCLS well 201. Lower Permian of Gondwanaland (Raine
et al. 2005); Triassic of Australia (de Jersey 1979). Lower
Permian of Argentina (Archangelsky & Gamerro 1979;
Gamerro & Archangelsky 1981; Playford & Dino 2002),
Brazil (Quadros et al. 1996) and Uruguay (Beri 1987, 1988;
Beri & Daners 1998; Mautino et al. 1998a; Beri et al.
2006).
Genus Verrucosisporites Ibrahim, emend. Smith & Butterworth,
1967
Verrucosisporites andersonii (Anderson) Backhouse, 1988
(Fig. 7Q)
DCLS well 201. Namurian-Lower Permian of Australia
(Price 1983; Backhouse 1988, 1991; Jones & Truswell
1992); Lower Permian of Africa (Anderson 1977), Antarc-
tica (Lindström 1995), Oman and Saudi Arabia (Stephenson
& Filatoff 2000a; Stephenson & Osterloff 2002; Stephenson
et al. 2003; Stephenson 2004). Pennsylvanian of Argentina
 Lower Permian palynology of Paraná Basin
(Garcı́a 1995; Césari & Gutiérrez 2001). This is the first
record in Uruguay.
Verrucosisporites microtuberosus (Loose) Smith & Butter-
worth, 1967 (Fig. 7S)
DCLS well 221. Carboniferous of the USA and Europe
(Smith & Butterworth 1967; Ravn 1979); Lower Permian
of Brazil (Quadros et al. 1996; Souza & Calegari 2004) and
Uruguay (Beri & Daners 1995).
Subinfraturma Baculati Dybová & Jachowicz, 1957
Genus Horriditriletes Bharadwaj & Salujha, 1964
Horriditriletes ramosus (Balme & Hennelly) Bharadwaj &
Salujha, 1964 (Fig. 8A)
DCLS wells 221, 201. Permian of Australia, India, Africa,
Brazil and Bolivia (Foster 1979; Quadros et al. 1996; Souza
& Calegari 2004; Raine et al. 2005; Smaniotto et al. 2006).
Triassic of Argentina (Zavattieri 1986; Volkheimer & Papú
1993), Lower Permian of Uruguay (Fasolo & Vergel 1994;
Beri et al. 2006).
Horriditriletes superbus (Foster) Césari, Archangelsky &
Villar de Seoane, 1995 (Fig. 7R)
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DCLS wells 221, 201. Lower Permian of Australia (Foster
1979; Mildenhall 1994), Brazil (Smaniotto et al. 2006),
Argentina (Césari et al. 1995; Vergel 1998) and Uruguay
(Mautino et al. 1998a).
Horriditriletes uruguaiensis (Marques-Toigo) Archangelsky
& Gamerro 1979 (Fig. 7T)
DCLS well 201. Lower Permian of Gondwanaland (Stephen-
son 2004); Pennsylvanian-Lower Permian from Brazil and
Argentina (Pons 1977; Archangelsky & Gamerro 1979;
Marques-Toigo 1991; Césari et al. 1995; Souza et al.
1997; Vergel 1998; Gutiérrez & Césari 2000; Césari &
Gutiérrez 2001; Gutiérrez & Limarino 2001; Dino & Play-
ford 2002; Smaniotto et al. 2006). Lower Permian of
Uruguay (Marques-Toigo 1974; Beri & Goso 1996, 1998;
Mautino et al. 1998a; Beri et al. 2006).
Infraturma Murornati Potonié & Kremp, 1954
Genus Convolutispora Hoffmeister, Staplin & Malloy, 1955
Convolutispora ordonezii Archangelsky & Gamerro, 1979
(Fig. 7N)
DCLS wells 201, 221. Pennsylvanian of northern Argentina
and eastern Paraguay (Archangelsky & Gamerro 1979;
Césari & Gutiérrez 1985; Vergel 1986; Ottone 1989; Césari
et al. 1995; Césari & Bercowski 1997; Muff et al. 1999;
Playford & Dino 2002). Lower Permian of Uruguay (Beri &
Daners 1995; Beri & Goso 1996; Mautino et al. 1998a; Beri
& Pecoits 2001; Beri et al. 2006).
Subturma Zonotriletes Waltz, 1935
Infraturma Auriculati Schopf, emend. Dettmann, 1963
Genus Murospora Somers, 1952
Murospora bicingulata Ybert, 1975 (Fig. 8C)
DCLS well 201. Lower Permian of Brazil (Ybert 1975;
Quadros et al. 1996) and Uruguay (Beri et al. 2000, 2006).
Murospora torifera Ybert, 1975 (Fig. 8D)
DCLS wells 201, 221. Lower Permian of Brazil (Ybert 1975)
and Uruguay (Beri & Goso 1996; Veroslavsky et al. 2003;
Beri et al. 2006).
Suprasubturma Laminanititriletes Smith & Butterworth, 1967
Subturma Zonolaminatitriletes Smith & Butterworth, 1967
Infraturma Cingulicavati Smith & Butterworth, 1967
Genus Cristatisporites Potonié & Kremp, 1954
Cristatisporites chacoparanaensis Ottone, 1989 (Fig. 8F)
DCLS well 201. Pennsylvanian-Lower Permian of Argentina
(Archangelsky & Gamerro 1979; Césari et al. 1995; Césari
& Gutiérrez 2001); Lower Permian of Uruguay (Fasolo &
Vergel 1994; Beri et al. 2006).
497
Cristatisporites inconstans Archangelsky & Gamerro, 1979
(Fig. 8L)
DCLS wells 201, 221. Pennsylvanian-Lower Permian of
Argentina (Archangelsky & Gamerro 1979; Césari et al.
1995; Césari & Gutiérrez 2001; Playford & Dino 2002)
and Brazil (Quadros et al. 1996; Souza et al. 2003); Lower
Permian of Uruguay (Mautino et al. 1998a; Veroslavsky
et al. 2003; Beri et al. 2006).
Cristatisporites lestai Archangelsky & Gamerro, 1979 (Fig.
8G)
DCLS well 201. Pennsylvanian-Lower Permian of Argentina
(Archangelsky & Gamerro 1979; Césari et al. 1995; Césari
& Gutiérrez 2001; Playford & Dino 2002) and Brazil
(Quadros et al. 1996; Souza et al. 2003); Lower Permian
of Uruguay (Beri et al. 2006).
Cristatisporites menendezii (Menéndez & Azcuy) Playford,
1978 (Fig. 8H)
DCLS well 201. Pennsylvanian of Argentina (Césari &
Gutiérrez 2001; Gutiérrez & Limarino 2001), and Brazil
(Quadros et al. 1996; Souza et al. 1997); Pennsylvanian-
Lower Permian of Brazil (Souza et al. 2003) and Oman
(Stephenson & Filatoff 2000a); Lower Permian of Uruguay
(Beri et al. 2006).
Genus Gondisporites Bharadwaj, 1962
Gondisporites serrulatus Césari, Archangelsky & Seoane,
1995 (Fig. 8J)
DCLS well 201. Lower Permian of Argentina (Césari et al.
1995) and Uruguay (Mautino et al. 1998a).
Genus Kraeuselisporites Leschick, emend. Jansonius, 1962
Kraeuselisporites apiculatus Jansonius, 1962 (Fig. 8S)
DCLS wells 201, 221. Upper Permian-Lower Trias-
sic of Canada and Russia (Jansonius 1962; Balme
1979; Mangerud 1994); Permian of Gabon (Jardiné
1974) and Brazil (Dı́as-Fabricio 1981; Quadros et al.
1996); Lower Permian of Uruguay (Beri & Goso
1996).
Genus Lundbladispora Balme ex Playford, 1965
Lundbladispora areolata Césari, Archangelsky & Seoane,
1995 (Fig. 8O)
DCLS wells 201, 221. Lower Permian from Argentina
(Césari et al. 1995) and Uruguay (Beri et al. 2006).
Lundbladispora braziliensis (Pant & Srivastava) Marques-
Toigo & Pons, emend Marques-Toigo & Piccarelli, 1985 (Fig.
8P)
DCLS wells 201, 221. Pennsylvanian-Lower Permian of
Argentina, Brazil, Oman and Saudi Arabia (Gutiérrez &
Césari 1989; Césari & Gutiérrez 2001; Playford & Dino
2002; Stephenson 2004), and Lower Permian from Uruguay
(Beri & Goso 1996, 1998; Beri et al. 2006).
Lundbladispora riobonitensis Marques-Toigo & Piccarelli,
1985 (Fig. 8N)
DCLS wells 201, 221. Pennsylvanian-Lower Permian of
Argentina (Césari & Gutiérrez 2001) and Lower Permian
of Uruguay (Beri et al. 2006).
Genus Vallatisporites Hacquebard, 1957
Vallatisporites arcuatus (Marques-Toigo) Archangelsky &
Gamerro, 1979 (Fig. 8T)
DCLS wells 201, 221. Pennsylvanian-Lower Permian of
Argentina, Brazil, Oman and Saudi Arabia (Archangelsky
& Gamerro 1979; Césari et al. 1995; Quadros et al. 1996;
Souza et al. 1997; Césari & Gutiérrez 2001; Playford &
Dino 2002; Stephenson 2004), Lower Permian of Uruguay
(Marques-Toigo 1974; Beri 1988; Beri & Daners 1995,
1998; Beri & Goso 1996; Mautino et al. 1998a; Beri et al.
2006).
 498 P. R. Gutiérrez et al.
Vallatisporites russoi Archangelsky & Gamerro, 1979 (Fig.
8M)
DCLS well 201. Pennsylvanian-Lower Permian of Argentina
(Archangelsky & Gamerro 1979; Césari et al. 1995; Césari
& Gutiérrez 2001; Playford & Dino 2002); Lower Permian
of Uruguay (Beri 1988; Beri & Goso 1996; Mautino et al.
1998a; Veroslavsky et al. 2003; Beri et al. 2006).
Suprasupturma Pseudosaccitriletes Richardson, 1965
Infraturma Monopseudosacciti Smith & Butterworth, 1967
Genus Spelaeotriletes Neves & Owens, 1966
Spelaeotriletes ybertii (Marques-Toigo) Playford & Powis,
1979 (Fig. 8Q)
DCLS well 201. Pennsylvanian-Lower Permian of Brazil
and Argentina (Césari & Gutiérrez 2001; Dino & Playford
2002; Souza et al. 2003) and Uruguay (Marques-Toigo 1970;
Playford et al. 2001).
Turma Monoletes Ibrahim, 1933
Suprasubturma Acavatomonoletes Dettmann, 1963
Subturma Azonomonoletes Luber, 1935
Genus Laevigatosporites Ibrahim, 1933
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Laevigatosporites vulgaris Ibrahim, 1933 (Fig. 9C)
DCLS well 201. This species is well known from Pennsylva-
nian, Permian and Triassic palynofloral assemblages (Foster
1979; Raine et al. 2005). Pennsylvanian of Argentina (Césari
& Gutiérrez 2001; Playford & Dino 2002). Permian of
Brazil (Playford & Dino 2000a). Lower Permian of Uruguay
(Fasolo & Vergel 1994).
Genus Polypodiisporites Potonié & Gelletich ex Potonié, 1956
Polypodiisporites mutabilis Balme, 1970 (Fig. 9B)
DCLS well 221. Upper Permian (Chhidru Formation)
of West Pakistan (Balme 1970); Permian of Tanzania
(Weiss 2001); Lower Permian of Uruguay (Mautino et al.
1988a).
Anteturma Variegerminantes Potonié, 1970
Turma Saccites Erdtman, 1947
Subturma Monosaccites Chitaley, emend. Potonié & Kremp, 1954
Infraturma Dipolsacciti Hart, emend. Dibner, 1971
Subinfraturma Apertacorpini Dibner, 1971
Genus Barakarites Bharadwaj & Tiwari, 1964
Barakarites rotatus (Balme & Hennelly) Bharadwaj & Tiwari,
1964 (Fig. 9D)
DCLS well 201. Known widely from Gondwanan Permian
strata (Foster 1979; Backhouse 1991; Playford & Dino
2000b; Stephenson & Filatoff 2000a). Argentina: Lower
Permian (Gutiérrez & Césari 2000; Césari & Gutiérrez
2001; Balarino & Gutiérrez 2006). This is the first record in
Uruguay.
Genus Cannanoropollis Potonié & Sah, 1960
Cannanoropollis densus (Lele) Bose & Maheshwari, 1968
(Fig. 9H)
DCLS wells 201, 221. Known widely from Gondwanan
Pennsylvanian-Permian strata (see Foster 1979; Gutiérrez
1993). Argentina: Pennsylvanian-Lower Permian (Gutiérrez
1993; Azcuy & di Pasquo 2000; Césari & Gutiérrez 2001).
Lower Permian of Uruguay (Beri 1988; Beri & Goso 1996;
Mautino et al. 1998b; Gutiérrez et al. 2006).
Cannanoropollis janakii Potonié & Sah, 1960 (Fig. 9G)
DCLS wells 201, 221. Known widely from Gondwanan
Permian strata (see Foster 1979; Gutiérrez 1993; Dino
& Playford 2000b). Pennsylvanian-Lower Permian of
Argentina (Gutiérrez 1993; Azcuy & di Pasquo 2000). Lower
Permian of Uruguay (Mautino et al. 1998b; Gutiérrez et al.
2006).
Cannanoropollis mehtae (Lele) Bose & Maheswari, 1968 (Fig.
9E)
DCLS wells 201, 221. Known widely from Gondwanan
Permian strata (Gutiérrez 1993). Pennsylvanian-Lower
Permian of Argentina (Gutiérrez & Césari 2000; Césari &
Gutiérrez 2001; Balarino & Gutiérrez 2006). Lower Permian
of Uruguay (Fasolo & Vergel 1994; Veroslavsky et al. 2003;
Gutiérrez et al. 2006).
Genus Costatascyclus Felix & Burbridge, emend. Urban, 1971
Costatascyclus crenatus Felix & Burdbridge, emend. Urban,
1971 (Fig. 9F)
DCLS wells 201, 221. Pennsylvanian of USA (Felix &
Burbridge 1967; Ravn 1979); Uppermost Carboniferous-
Lower Permian of Brazil (Playford & Dino 2000b; Souza
et al. 2003); Lower Permian of Uruguay (Beri 1988;
Gutiérrez et al. 2006). Caheniasaccites ovatus Bose & Kar,
emend. Archangelsky & Gamerro illustrated by Beri (1988,
p. 37, fig. 21), from the Lower Permian of Uruguay, is
included in this species.
Genus Plicatipollenites Lele, 1964
Plicatipollenites gondwanensis (Balme & Hennelly) Lele,
1964 (Fig. 9P)
DCLS wells 201, 221. Pennsylvanian-Permian strata of
Gondwanaland (Gutiérrez 1993; Césari et al. 1995; Césari &
Gutiérrez 2001; Playford & Dino 2000a; Felix et al. 2006).
Permian of Brazil (Cauduro 1970; Menéndez 1976; Quadros
et al. 1996); Lower Permian of Uruguay (Beri & Goso 1996;
Mautino et al. 1998b).
Plicatipollenites malabarensis (Potonié & Sah) Foster, 1975
(Fig. 9M)
DCLS wells 201, 221. Pennsylvanian-Permian strata of
Gondwanaland (Gutiérrez 1993; Quadros et al. 1996; Play-
ford & Dino 2000a; Felix et al. 2006). Lower Permian of
Uruguay (Beri & Goso 1996; Beri & Daners 1998; Mautino
et al. 1998b; Beri et al. 2000; Gutiérrez et al. 2006).
Plicatipollenites trigonalis Lele, 1964 (Fig. 9I)
DCLS well 221. Pennsylvanian-Permian of Argentina and
Brazil (Gutiérrez 1993; Césari et al. 1995; Quadros et al.
1996; Azcuy & di Pasquo 2000; Playford & Dino 2000a;
Césari & Gutiérrez 2001; Pérez Loinaze & Césari 2004;
Felix et al. 2006; Premaor et al. 2006). This is the first
record from Uruguay.
Genus Potonieisporites Bhardwaj, emend. Bharadwaj, 1964
Potonieisporites brasiliensis (Nahuys et al.) Archangelsky &
Gamerro, 1979 (Fig. 9L)
DCLS wells 201, 221. Pennsylvanian-Permian of Argentina
and Brazil (Nahuy et al. 1968; Gutiérrez 1993; Quadros et al.
1996; Azcuy & di Pasquo 2000; Playford & Dino 2000a,
2002; Stephenson & Filatoff 2000a; Césari & Gutiérrez
2001). Lower Permian of Uruguay (Beri & Goso 1996; Beri
& Daners 1998; Beri & Pecoits 2001; Gutiérrez et al. 2006).
Potonieisporites magnus Lele & Karim, 1971 (Fig. 9Q)
DCLS wells 201, 221. Pennsylvanian-Permian of Argentina
and Brazil (Gutiérrez 1993; Azcuy & di Pasquo 2000;
Gutiérrez & Césari 2000; Césari & Gutiérrez 2001; Souza
et al. 2003). Lower Permian of Uruguay (Gutiérrez et al.
2006).
Potonieisporites neglectus Potonié & Lele, 1961 (Fig. 9N)
DCLS well 221. Pennsylvanian-Permian of Gondwana
(Gutiérrez 1993; Quadros et al. 1996; Playford & Dino
2000b). Lower Permian of Uruguay (Veroslavsky et al.
2003).
Potonieisporites novicus Bharadwaj, 1954 (Fig. 9K)
DCLS wells 201, 221, El Barón. Widely reported from
strata ranging in age from Namurian to Early Permian of
Gondwana (Gutiérrez 1993; Quadros et al. 1996; Play-
ford & Dino 2000b). Pennsylvanian-Lower Permian of
 Lower Permian palynology of Paraná Basin
Argentina (Azcuy & di Pasquo 2000; Gutiérrez & Césari
2000; Césari & Gutiérrez 2001). Lower Permian of Uruguay
(Beri & Goso 1996; Mautino et al. 1998b; Gutiérrez et al.
2006).
Potonieisporites triangulatus Tiwari, 1965 (Fig. 9O)
DCLS wells 201, 221. Pennsylvanian and Permian of
Argentina, Brazil and India (see Tiwari 1965; Ottone &
Azcuy 1990; Gutiérrez 1993; Playford & Dino 2000b). This
is the first record in Uruguay.
Subinfraturma Clausicorpini Dibner, 1971
Genus Caheniasaccites Bose & Kar, emend. Archangelsky &
Gamerro, 1979
Caheniasaccites densus Lele & Karim, emend. Gutiérrez, 1993
(Fig. 10D)
DCLS well 201. Pennsylvanian-Permian of Argentina and
Brazil (see Gutiérrez 1993; Gutiérrez & Césari 2000; Césari
& Gutiérrez 2001; Playford & Dino 2002). Lower Permian
of Uruguay (Gutiérrez et al. 2006).
Caheniasaccites ovatus Bose & Kar, emend. Gutiérrez, 1993
(Fig. 10A)
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DCLS wells 201, 221. Widespread in Gondwana succes-
sions of Late Carboniferous-Permian age (Gutiérrez 1993;
Gutiérrez & Césari 2000; Playford & Dino 2000a; Césari
& Gutiérrez 2001). Lower Permian of Uruguay (Gutiérrez
et al. 2006).
Subinfraturma Bilateralsaccini Dibner, 1971
Genus Latusipollenites Marques-Toigo, 1974
Latusipollenites quadrisaccatus Marques-Toigo, 1974 (Fig.
10C)
DCLS well 201. Pennsylvanian-Permian of Argentina and
Brazil (Dias-Fabricio 1981; Dı́as 1993; Gutiérrez & Césari
2000; Césari & Gutiérrez 2001; Balarino & Gutiérrez 2006;
Gutiérrez et al. 2006). Lower Permian of Uruguay (Marques-
Toigo 1974; Gutiérrez et al. 2006).
Subinfraturma Amphisaccini Dibner, 1971
Genus Divarisaccus Venkatachala & Kar, 1966a
Divarisaccus stringoplicatus Ottone, 1991 (Fig. 10J)
DCLS well 221. Pennsylvanian of Argentina and Brazil
(Ottone 1991; Césari & Limarino 2002; Longhim et al. 2002;
Souza et al. 2003); Lower Permian of Brazil (Smaniotto et al.
2006). This is the first record in Uruguay.
Genus Stellapollenites Lele, 1965
Stellapollenites talchirensis Lele, 1965 (Fig. 10H)
DCLS well 201. Permian of India and Africa (Lele 1965;
Bose & Kar 1966; Chandra & Lele 1980). Lower Permian
of Brazil (Marques-Toigo & Pons 1976; Dias 1993; Quadros
et al. 1996) and Uruguay (Gutiérrez et al. 2006).
Infraturma Striasacciti Bharadwaj, 1962
Genus Mabuitasaccites Bose & Kar, 1966
Mabuitasaccites crucistriatus (Ybert) Playford & Dino, 2000b
(Fig. 10B)
DCLS wells 221, 201, El Barón. Permian of Brazil
and Argentina (Daemón & Quadros 1970; Ybert 1975;
Archangelsky & Gamerro 1979; Dias 1993; Césari et al.
1995; Quadro et al. 1996; Playford & Dino 2000b; Balarino
& Gutiérrez 2006). Lower Permian of Uruguay (Gutiérrez
et al. 2006).
Genus Striomonosaccites Bharadwaj, 1962
Striomonosaccites cicatricosus Archangelsky & Gamerro,
1979 (Fig. 10E)
DCLS wells 201, 221, El Barón. Permian of Brazil and
Argentina (Daemón & Quadros 1970; Archangelsky &
Gamerro 1979; Dias 1993; Césari et al. 1995). Lower
Permian of Uruguay (Beri & Daners 1998; Mautino et al.
1998b; Gutiérrez et al. 2006).
499
Subturma Disaccites Cookson, 1947
Subturma Dissaccitrileti Leschik, 1955
Genus Colpisaccites Archangelsky & Gamerro, 1979
Colpisaccites granulosus Archangelsky & Gamerro, 1979
(Fig. 10F)
DCLS well 221, El Barón. Pennsylvanian-Lower Permian
of Argentina (Archangelsky & Gamerro 1979; Césari et al.
1995, 1996; Césari & Gutiérrez 2001; Balarino & Gutiérrez
2006). Pennsylvanian of Brazil (Playford & Dino 2000a).
Lower Permian of Uruguay (Beri & Daners 1995; Mautino
et al. 1998b; Gutiérrez et al. 2006).
Genus Limitisporites Leschik, emend. Schaarschmidt, 1963
Limitisporites hexagonalis Bose & Maheshwari, 1968
(Fig. 10P)
DCLS well 201, 221. Widely known from Pennsylvanian-
Permian of Gondwanaland sequences. In Argentina
(Gutiérrez & Césari 2000; Césari & Gutiérrez 2001; di
Pasquo et al. 2001; Pérez Loinaze & Césari 2004); Permian
of Uruguay (Fasolo & Vergel 1994; Beri & Daners 1995;
Mautino et al. 1988b; Beri & Pecoits 2001). From the
observed features in the illustrations of the material referred
to Limitisporites rectus Leschik by Beri & Daners (1995, pl.
2, fig. 4) from the Lower Permian of Uruguay, this material
is here referred to L. hexagonalis.
Limitisporites rectus Leschik 1956 (Fig. 10N)
DCLS wells 201, 221. Widely known from Pennsylvanian-
Permian of Gondwanaland. Pennsylvanian of Argentina
(Gutiérrez & Césari 2000; Césari & Gutiérrez 2001; di
Pasquo et al. 2001; Pérez Loinaze & Césari 2004; Balar-
ino & Gutiérrez 2006) and Lower Permian of Brazil (Souza
& Calegari 2004) and Uruguay (Mautino et al. 1988b; Beri
& Pecoits 2001, Gutiérrez et al. 2006).
Limitisporites vesiculosus (Potonié & Klaus) Schaarschmidt,
1963 (Fig. 10L)
DCLS well 201. Permian of Brazil (Dino et al. 2002; Souza
et al. 2003; Souza & Callegari 2004). This is the first record
from Uruguay. From the features observed in the illustrations
of material referred to Limitisporites delasaucei (Potonié &
Klaus) Schaarschmidt by Dino et al. (2002, fig. 19), this is
better placed in L. vesiculosus.
Infraturma Disacciatrileti Leschik, emend. Potonié, 1958
Genus Alisporites Daugherty, emend. Jansonius, 1971
Alisporites australis de Jersey, 1962 (Fig. 11B)
DCLS well 221, El Barón. Permian-Lower Jurassic of Gond-
wana (Foster 1979). Argentina: Permian-Jurassic (Jain 1968;
Zavattieri 1987, 1991b, 1992; Papú 1991; Ottone et al. 1992;
Zavattieri & Volkheimer 1992; Césari et al. 1996; Césari &
Gutiérrez 2001; Balarino & Gutiérrez 2006). Lower Permian
of Uruguay (Fasolo & Vergel 1994; Beri & Pecoits 2001).
Based on the features observed in Sulcatisporites ovatus
(Balme & Hennelly) Bharadwaj illustrated by Russo et al.
(1980, pl. 1, fig. 8) from the Chacoparaná Basin, this material
is likely to be A. australis.
Alisporites parvus de Jersey, 1962 (Fig. 11K)
DCLS well 221. Triassic-Jurassic of Gondwanaland (Raine
et al. 2006); Triassic of Argentina (Zavattieri & Rojo 2005).
This is the first record from Uruguay.
Genus Platysaccus Naumova ex Potonié & Kremp, 1954
Platysaccus papilionis Potonié & Klaus, 1954 (Fig. 12I)
DCLS wells 201, 221, El Barón. Widely known from
Pennsylvanian-Jurassic sequences of Gondwanaland and
Euroamerica (see Zavatieri 1987). In Argentina it has been
recorded from the Pennsylvanian-Lower Permian (Césari &
Gutiérrez 2001) and Triassic (Jain 1968; Zavattieri 1987).
Lower Permian of Brazil (Quadros et al. 1996) and Uruguay
 500 P. R. Gutiérrez et al.
(Beri & Daners 1995; Gutiérrez et al. 2006). The material
illustrated as Platysaccus vesiculosus Schaarschmidt by Beri
& Daners (1995, pl. 2, fig. 1) from the Lower Permian of
Uruguay is included in P. papilionis on account of its char-
acteristics amb, form of the corpus, absence of folds, etc.
Platysaccus leschikii Hart, 1960 (Fig. 12G)
DCLS well 221. Widely known from Permian-Triassic
sequences of Gondwanaland and Euroamerica (Lindström
1995, 1996). Lower Permian of Uruguay (Fasolo & Vergel
1994).
Genus Pteruchipollenites Couper, 1958
Pteruchipollenites gracilis (Segroves) Foster, 1979 (Fig. 12M)
DCLS wells 201, 221. Permian of Africa, Antartic and
Australia (Foster 1979; Lindström 1995). Pennsylvanian-
Lower Permian of Argentina (Gutiérrez & Césari 2000;
Césari & Gutiérrez 2001; Balarino & Gutiérrez 2006). Trias-
sic of Argentina (Ottone & Garcı́a 1991). Lower Permian of
Uruguay (Fasolo & Vergel 1994; Gutiérrez et al. 2006).
Genus Scheuringipollenites Tiwari, 1973
Scheuringipollenites barakarensis (Tiwari) Tiwari, 1973 (Fig.
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13G)
DCLS wells 201, 201, El Barón. Permian of Gondwana (see
Lidström 1995, 1996). Uppermost Carboniferous of Brazil
(Longhim et al. 2002); Permian of Argentina (Césari &
Gutiérrez 2001; Balarino & Gutiérrez 2006); Lower Permian
of Uruguay (Gutiérrez et al. 2006).
Scheuringipollenites circularis Césari, Archangelsky &
Seoane, 1995 (Fig. 13F)
DCLS wells 201, 221, El Barón. Lower Permian of Argentina
(Césari et al. 1995; Césari & Gutiérrez 2001; Balarino
& Gutiérrez 2006), and Uruguay (Mautino et al. 1998b;
Gutiérrez et al. 2006).
Scheuringipollenites medius (Burjack) Dı́as-Fabricio, 1981
(Fig. 12H, K)
DCLS wells 201, 221, El Barón. Lower Permian of Argentina
(Césari et al. 1995, 1996; Césari & Gutiérrez 2001; Balar-
ino & Gutiérrez 2006), Brazil (Dias-Fabricio 1981; Quadros
et al. 1996) and Uruguay (Gutiérrez et al. 2006).
Genus Vitreisporites Leschik, emend. Jansonius, 1962
Vitreisporites pallidus (Reissinger) Nilsson, 1958 (Fig. 13N)
DCLS well 221, El Barón. Permian to Cretaceous (see
Farabee & Canright 1986; Lindström 1996; Raine et al.
2006). This is the first record from Uruguay.
Infraturma Striatiti Pant, 1954
Genus Corisaccites Venkatachala & Kar, 1966b
Corisaccites alutas Venkatachala & Kar, 1966b (Fig. 13O)
DCLS well 221. Permian of Gondwana (see Playford & Dino
2000b). Permian of Africa (Anderson 1977; Weiss 2001),
Oman (Stephenson & Filatoff 2000b), Brazil (Cauduro 1970;
Daemon & Quadros 1970; Playford & Dino 2000b; Dino
et al. 2002) and Argentina (Archangelsky & Gamerro 1980;
Playford & Dino 2002). This is the first record from Uruguay.
Some specimens assignable to this species (corpus with two
well-defined taeniae and with narrow cappula) were included
by Anderson (1977, p. 125, pl. 180, figs 18–19, 21–22; pl.
181, fig. 3–4, 7, 9 only) in Lueckisporites nyakapenden-
sis Hart and L. welkomensis Anderson (1977, p. 126; pl.
182, figs 2, 6–9 only). The specimens described by Cauduro
(1970, pp. 20–21, pl. 16, figs 112–114) as L. densus, from the
Rio Bonito Formation of Brazil, are identifiable as Corisac-
cites alutas. The material illustrated as C. sp. by Archangel-
sky & Gamerro (1980, pl. 2, fig. 5), from the Lower Permian
of Colorado Basin (Argentina), could be referred to C. alutas
on the basis of the narrow sulcus, body shape, and sacci
ouline.
Genus Hamiapollenites Wilson, 1962
Hamiapollenites fusiformis Marques-Toigo, emend.
Archangelsky & Gamerro, 1979 (Fig. 13M)
DCLS wells 201, 221, El Barón. Uppermost Carboniferous-
Lower Permian of Brazil (Dellazana 1976; Pons 1979;
Marques-Toigo et al. 1990; Dı́as 1993; Quadros et al. 1996;
Playford & Dino 2000b) and Argentina (Archangelsky &
Gamerro 1979; Vergel 1987b; Césari et al. 1995; Gutiérrez &
Césari 2000; Césari & Gutiérrez 2001; Balarino & Gutiérrez
2006). Lower Permian of Uruguay (Marques-Toigo 1974;
Vergel 1987a; Beri 1988; Beri & Daners 1995).
Genus Lueckisporites Potonié & Klaus, emend. Klaus, 1963
Lueckisporites latisaccus Archangelsky & Gamerro, 1979
(Fig. 14A, C)
DCLS well 221, El Barón. Lower Permian of Brazil
(Marques-Toigo & Souza 2005; Premoar et al. 2006) and
Argentina (Archangelsky & Gamerro 1979; Césari et al.
1995; Césari & Gutiérrez 2001). This is the first record from
Uruguay. In contrast to Premoar et al. (2006, p. 226), we
agree with the criteria used by Archangelsky & Gamerro
(1979, p. 451) for separating Luekisporites latisaccus from
L. nyakapendensis Hart (1960), including absence of striae
on cappa, wide average striae and presence of monolete
mark.
Lueckisporites stenotaeniatus Menéndez, 1976 (Fig. 14G, L)
DCLS well 221, El Barón. Lower Permian of Brazil
(Menéndez 1976; Souza & Marques-Toigo 2005; Premaor
et al. 2006), Argentina (Archangelsky & Gamerro 1979;
Césari et al. 1995, 1996; Césari & Gutiérrez 2001) and
Uruguay (Beri & Daners 1995; Mautino et al. 1998b; Beri
et al. 2000).
Lueckisporites virkkiae Potonié & Klaus, emend. Clarke, 1965
(Figs 13J, 14B)
DCLS well 201, 201, El Barón. Typical of the Permian
globally (Raine et al. 2006). Lower Permian of Argentina
(Archangelsky & Gamerro 1979; Césari et al. 1996; Césari
& Gutiérrez 2001) and Brazil (Dellazana 1976; Menéndez
1976; Quadros et al. 1996; Premaor et al. 2006). This is the
first record from Uruguay.
Genus Lunatisporites Leschik, emend. Scheuring, 1970
Lunatisporites noviaulensis (Leschik) de Jersey, 1979 (Fig.
14K)
DCLS well 221. Permian-Triassic of Gondwana and Europe
(Clarke 1965; Balme 1970, 1979; Jardiné 1974; de Jersey
1979; Raine et al. 2005). Permian of Brazil (Quadros et al.
1996). This is the first record from Uruguay.
Lunatisporites pellucidus (Goubin) Helby, emend. de Jersey
1972 (Fig. 14H)
DCLS well 201. Permian-Triassic of Gondwana (Balme
1970, 1979; de Jersey 1972, 1979; Raine et al. 2005). Lower
Permian of Brazil (Nahuys et al. 1968). This is the first
record from Uruguay.
Lunatisporites variesectus Archangelsky & Gamerro, 1979
(Figs 14I, 15A, C)
DCLS wells 201, 221, El Barón. Late Pennsylvanian-Lower
Permian of Argentina (Archangelsky & Gamerro 1979;
Césari et al. 1995, 1996; Césari & Gutiérrez 2001; Balar-
ino & Gutiérrez 2006). Other South American reports are
from the Permian in the Paraná Basin of Brazil and Paraguay
(Playford & Dino 2002; Premaor et al. 2006). Lower Permian
of Uruguay (Beri & Daners 1995; Beri et al. 2000; Beri &
Pecoits 2001).
Genus Protohaploxypinus Samoilovich, emend. Morbey, 1975
Protohaploxypinus amplus (Balme & Hennelly) Hart, 1964
(Fig. 15H)
 Lower Permian palynology of Paraná Basin
DCLS wells 201, 221, El Barón. Known widely from
Permian strata of Gondwanaland (Foster 1979; Backhouse
1991; Farabee et al. 1991; Lindström 1995). Lower Permian
of Brazil (Daemón & Quadros 1970; Ybert 1975; Dellazana
1976; Pons 1977; Quadros et al. 1996; Souza et al. 1997;
Playford & Dino 2000b; Souza & Callegari 2004; Premaor
et al. 2006). Uppermost Pennsylvannian-Lower Permian of
Argentina (Césari et al. 1995; Gutiérrez & Césari 2000;
Césari & Gutiérrez 2001; Balarino & Gutiérrez 2006). Lower
Permian of Uruguay (Beri & Daners 1995; Mautino et al.
1988b).
Protohaploxypinus bharadwajii Foster, 1979 (Fig. 15M)
DCLS well 201, 221. Permian of India, Australia and Antarc-
tica (Foster 1979; Playford 1990). Lower Permian of Brazil
(Playford & Dino 2000b; Premaor et al. 2006) and Uruguay
(Fasolo & Vergel 1994; Gutiérrez et al. 2006).
Protohaploxypinus goraiensis (Potonié & Lele) Hart, 1964
(Fig. 15G)
DCLS well 201. Lower Permian of Africa (Stapleton 1978),
Brazil (Quadros et al. 1996) and Uruguay (Beri & Daners
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1995; Mautino et al. 1998b; Gutiérrez et al. 2006).
Protohaploxypinus limpidus (Balme & Hennelly) Balme &
Playford, 1967 (Fig. 14F)
DCLS well 201, 221, El Barón. Reported widely from Gond-
wanaland (Anderson 1977; Foster 1979; Farabee et al. 1991).
Lower Permian of Argentina (Gutiérrez & Césari 2000;
Césari & Gutiérrez 2001), Brazil (Marques-Toigo 1972;
Marques-Toigo et al. 1990; Dias 1993; Quadros et al. 1996)
and Uruguay (Beri & Daners 1995; Veroslavsky et al. 2003;
Gutiérrez et al. 2006).
Genus Staurosaccites Dolby in Dolby & Balme, 1976
Staurosaccites cordubensis Archangelsky & Gamerro, 1979
(Fig. 15I, K)
DCLS well 221, El Barón. Lower Permian of Argentina
(Archangelsky & Gamerro 1979; Césari et al. 1996; Césari
& Gutiérrez 2001; Playford & Dino 2002), Brazil (Daemón
& Quadros 1970; Menéndez 1976; Quadros et al. 1996;
Premaor et al. 2006) and Uruguay (Beri & Daners 1995).
The material illustrated as Staurosaccites sp. by Archangel-
sky & Gamerro (1980, pl. 2, fig. 11), from the Lower
Permian of the Colorado Basin (Argentina), could be
referred to S. cordubensis on the basis of corpus and
sacci shape, overal ouline, and a monolete mark in the
corpus.
Genus Striatoabieites Zoricheva & Sedova ex Sedova, emend.
Hart, 1964
Striatoabieites anaverrucosus Archangelsky & Gamerro, 1979
(Fig. 16A)
DCLS well 221, El Barón. Lower Permian of Argentina
(Archangelsky & Gamerro 1979; Césari & Gutiérrez 2001;
Playford & Dino 2002) and Uruguay (Beri & Daners 1995;
Beri & Pecoits 2001).
Striatoabieites multistriatus (Balme & Hennelly) Hart, 1964
(Fig. 15L)
DCLS wells 201, 221, El Barón. Permian-Triassic? of Gond-
wanaland (Playford & Dino 2002; Raine et al. 2005). Lower
Permian of Argentina (Césari et al. 1995, 1996; Césari &
Gutiérrez 2001; Playford & Dino 2002), Brazil (Souza &
Calegari 2004) and Uruguay (Beri & Daners 1995; Beri &
Goso 1996; Mautino et al. 1998b; Beri & Pecoits 2001;
Gutiérrez et al. 2006).
Genus Striatopodocarpites Zoricheva & Sedova ex Sedova,
emend. Hart, 1964
Striatopodocarpites cancellatus (Balme & Hennelly) Hart,
1963 (Fig. 16C)
501
DCLS wells 201, 221, El Barón. Permian of Gondwanaland
(Raine et al. 2005). Lower Permian of Brazil (Premaor et al.
2006) and Argentina (Césari et al. 1996).
Striatopodocarpites gondwanensis Lakhanpal, Sah & Dube,
emend. Hart, 1964 (Fig. 16F)
DCLS wells 201, 221, El Barón. Permian of India (Lakhan-
pal et al. 1960; Tiwari 1965; Maheshwari 1967; Kar 1968;
Sinha 1972), Australia (Segroves 1969; Foster 1975, 1979)
and Africa (Jardiné 1974; MacRae 1988). This is the first
record from Uruguay.
Striatopodocarpites octostriatus Hart, 1960 (Fig. 16B)
DCLS well 221, El Barón. Permian of Africa (Hart 1960;
Weiss 2001). This is the first record from Uruguay.
Infraturma Circumstriatiti Lele & Makada, 1972
Genus Illinites Kosanke, emend. Jansonius & Hills, 1976
Illinites unicus Kosanke, 1950 (Fig. 16E)
DCLS well 201. Known widely from the Pennsylvanian and
Permian globally (Playford & Dino 2000b; Azcuy et al.
2002). This is the first record from Uruguay.
Subturma Polysaccites Cookson, 1947
Genus Polarisaccites Ybert & Marques-Toigo, 1971
Polarisaccites bilateralis Ybert & Marques-Toigo, 1971 (Fig.
16D)
DCLS well 201, 221, El Barón. Lower Permian of Uruguay
(Ybert & Marques-Toigo 1971; Marques-Toigo 1972),
Argentina (Archangelsky & Gamerro 1979; Vergel 1987b;
Césari et al. 1995) and Brazil (Ybert 1975; Dias-Fabricio
1981; Dias 1993; Playford & Dino 2000b). Some of the
material described as Gnetaceapollenites ovatus Anderson
(1977: 132, pl. 187, figs 3–5) may be referred to this
species.
Turma Plicates Naumova, emend. Potonié, 1960
Subturma Costates Potonié, 1970
Genus Vittatina Luber ex Samoilovich, emend. Wilson, 1962
Vittatina corrugata Marques-Toigo, 1974 (Fig. 16J, N)
DCLS wells 201, 221. Lower Permian of Brazil (Quadros
et al. 1996; Souza & Calegari 2004), Uruguay (Marques-
Toigo 1974; Beri & Daners 1996; Beri & Goso 1996, 1998;
Beri 1988; Mautino et al. 1998b; Beri et al. 2000; Beri
& Pecoits 2001). The material described as Tiwarispora
simplex (Tiwari) Maheshwari & Kar by Mautino et al.
(1988b, p. 309, pl. 3, fig. 7), from the Melo Formation,
is referred to this species.
Vittatina costabilis Wilson, emend. Tschudy & Kosanke, 1966
(Fig. 16L)
DCLS wells 201, 221, El Barón. Permian of Africa, Oman
(Stapleton 1978; Stephenson & Filatoff 2000a); Permian
of Brazil (Quadros et al. 1996; Playford & Dino 2000b;
Souza & Calegari 2004; Premaor et al. 2006; Smaniotto
et al. 2006); Lower Permian of Argentina (Césari et al.
1995; Césari & Gutiérrez 2001) and Uruguay (Beri & Daners
1995; Beri & Goso 1996; Beri 1988; Mautino et al. 1998b).
The material assigned to Vittatina latericostata Menéndez
by Mautino et al. (1998b, p. 309, pl. 3, fig. 3), from the
Melo Formation, is transferred to this species because of
the characteristic amb, number of teniae, presence of folds,
etc.
Vittatina subsaccata Samoilovich, 1953 (Fig. 16G, K)
DCLS wells 201, 221, El Barón. Typical of the Permian of
Russia and Canada (Jansonius 1962), it is also found in the
Lower Permian of Argentina (Césari et al. 1995; Gutiérrez &
Césari 2000; Césari & Gutiérrez 2001; Balarino & Gutiérrez
2006), Brazil (Playford & Dino 2000b; Quadros et al. 1996)
and Uruguay (Beri 1987; Beri & Daners 1995; Mautino et al.
1998b; Gutiérrez et al. 2006).
 502 P. R. Gutiérrez et al.
Genus Weylandites Bharadwaj & Srivastava, 1969
Weylandites lucifer (Bharadwaj & Salujha) Foster, 1975 (Fig.
16H)
DCLS wells 201, 221, El Barón. Permian of Gondwana
(Foster 1975, 1979; Macrae 1988; Raine et al. 2005); Lower
Permian of Argentina (Césari et al. 1995; Césari & Gutiérrez
2001), Brasil (Menéndez 1976; Quadros et al. 1996; Premaor
et al. 2006) and Uruguay (Fasolo & Vergel 1994; Beri &
Daners 1995; Mautino et al. 1998b; Beri & Pecoits 2001;
Gutiérrez et al. 2006).
Weylandites magmus (Bose & Kar) Bharadwaj & Dwivedi,
1981 (Fig. 16M)
DCLS wells 201, 221, El Barón. Permian of Gondwana
(Bhradwaj & Dwivedi 1981; Backhouse 1991; Raine et al.
2005); Lower Permian of Argentina (Césari et al. 1996;
Césari & Gutiérrez 2001; Balarino & Gutiérrez 2006) and
Uruguay (Mautino et al. 1998b; Beri & Pecoits 2001;
Gutiérrez et al. 2006). The material described as Vittatina
densa Anderson (1977, p. 107, pl. 132, figs 1–39), from
the Permian of South Africa, should be referred to this
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species.
Subturma Praecolpates Potonié & Kremp, 1954
Genus Marsupipollenites Balme & Hennelly, emend. Balme,
1970
Marsupipollenites striatus (Balme & Hennelly) Foster, 1975
(Fig. 16I)
DCLS wells 201, 221, El Barón. Permian of Gondwana
(Raine et al. 2005); Lower Permian of Argentina (Archangel-
sky & Gamerro 1979; Césari et al. 1995, 1996; Gutiérrez
& Césari 2000; Césari & Gutiérrez 2001), Brazil (Nahuys
et al. 1968; Dı́as-Fabricio 1981; Quadros et al. 1996;
Premaor et al. 2006) and Uruguay (Beri & Daners 1995;
Mautino et al. 1998b; Beri & Pecoits 2001; Gutiérrez et al.
2006).
Marsupipollenites triradiatus Balme & Hennelly, 1956 (Fig.
16Q)
DCLS well 221. Permian of Gondwana (de Jersey 1979;
Raine et al. 2005). Lower Permian of Brazil (Quadros et al.
1996; Premaor et al. 2006) and Uruguay (Mautino et al.
1998b; Beri & Pecoits 2001; Gutiérrez et al. 2006).
Turma Monocolpates Iversen & Troels-Smith, 1950
Subturma Striaticolpites Bose & Kar, 1966
Infraturma Monostriocolpites Bose & Kar, 1966
Genus Pakhapites Hart, 1965
Pakhapites fusus (Bose & Kar) Menéndez, 1971
(Fig. 16O)
DCLS wells 201, 221. Permian of Gondwanaland (Bose
& Kar 1966, 1967; Stapleton 1978; Foster 1979; Chan-
dra & Lele 1980; Playford & Dino 2000b; Gutiérrez &
Césari 2000). Lower Permian of Argentina (Menéndez 1971;
Césari et al. 1995, 1996; Gutiérrez & Césari 2000; Césari &
Gutiérrez 2001; Playford & Dino 2002; Balarino & Gutiérrez
2006), Brazil (Bharadwaj et al. 1976; Menéndez 1976; Dı́as-
Fabricio 1981; Marques-Toigo et al. 1990; Quadros et al.
1996) and Uruguay (Fasolo & Vergel 1994; Gutiérrez et al.
2006).
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Pakhapites ovatus (Bose & Kar) Garcı́a, 1996 (Fig. 16P)
DCLS wells 201, 221. Permian of Gondwana (Bose & Kar
1966; Kar & Bose 1967; Jardiné 1974). Lower Permian of
Argentina (Césari & Gutiérrez 2001), Brazil (Quadros et al.
1996) and Uruguay (Gutiérrez et al. 2006).
Division Chlorophyta Pacher, 1914
Class Clorophyceae Kützing, 1843
Genus Botryococcus Kützing, 1849
Botryococcus braunii Kützing, 1849 (Fig. 5E)
DCLS wells 201, 221. Widely known from Permian to
Quaternary strata of the world. Lower Permian of Uruguay
(Beri & Daners 1995, 1998; Mautino et al. 1998a; Beri et al.
2000, 2006).
Class Zygmantaceae Round, 1971
Order Zygnematales Borge & Pascher, 1913
Genus Tetraporina Naumova ex Naumova, emend. Lindgren,
1980
Tetraporina punctata (Tiwari & Navale) Kar & Bose, 1976
(Fig. 16R)
DCLS well 201. Pennsylvanian of Argentina (Césari et al.
1995; Césari & Gutiérrez 2001; Gutiérrez & Limarino
2001). Lower Permian of Uruguay (Beri et al. 2006).
Grupo Acritarcha Evitt, 1963
Subgrupo Schizomorphitae Segroves, 1967
Genus Brazilea Tiwari & Navale, 1967
Brazilea scissa (Balme & Hennelly) Foster, 1975 (Fig. 5B, C)
DCLS wells 201, 221. Pennsylvanian-Lower Permian of
Argentina (Archangelsky & Gamerro 1979; Césari et al.
1995, 1996; Césari & Gutiérrez 2001; Gutiérrez & Limarino
2001); Lower Permian of Brazil (Smaniotto et al. 2006) and
Uruguay (Beri et al. 2006).
Brazilea plurigenus (Balme & Hennelly) Foster, 1979 (Fig.
5H)
DCLS wells 201, 221, El Barón. Pennsylvanian-Lower
Permian of Argentina (Césari et al. 1995; Gutiérrez &
Limarino 2001), Lower Permian of Uruguay (Beri et al.
2006).
Division Prasinophyta Round, 1971
Genus Leiosphaeridia Eisenack, emend. Downie & Sarjeant, 1963
Leiosphaeridia talchirensis Lele & Karim 1971 (Fig. 5G)
DCLS well 201. Permian of India (Lele & Karim 1971; Lele
1975). Lower Permian of Brazil (Smaniotto et al. 2006)
and Uruguay (Beri et al. 2006). The material described as
Leiosphaeridia sp. by Beri et al. (2006, p. 241, fig. 4O;
San Gregorio Formation, Uruguay) and by Smaniotto et al.
(2006, p. 318–319, fig. 5P; Lower Permian of Brazil), is
included in L. talchirensis Lele & Karim based on the nature
of the exina (levigate, folded with a single broad crescentic
fold at vesicle margin, dimenions, etc.).
Fungi
Genus Portalites Hemer & Nygreen, 1967
Portalites gondwanensis Nahuys, Alpern & Ybert, 1968 (Fig.
5A)
DCLS wells 201, 221. Pennsylvanian of Argentina (Césari
& Gutiérrez 2001); Lower Permian of Uruguay (Beri et al.
2006).