Systematic palynological study of the Cortaderas Formation, (Mississippian) Río Blanco Basin, Argentina. Part One
Valeria Pérez Loinaze1
1División Paleobotánica, Museo Argentino de Ciencias Naturales "Bernardino Rivadavia". Av. Ángel Gallardo 470, 1405 Buenos Aires, Argentina. loinazev@macn.gov.ar
Abstract. Well preserved Mississippian palynological assemblages have been obtained from the Cortaderas Formation in the Río Blanco Basin, northwestern Argentina. The geological setting and stratigraphic provenance of the samples is provided. In this first part, fifty-two species are described from a total of seventy-eight species of spores recognized. Five new species are proposed: Verrucosisporites incompositus sp. nov., Verrucosisporites perbrevis sp. nov., Anapiculatisporites exiguus sp. nov., Anapiculatisporites expolitus sp. nov. and Dibolisporites insolitus sp. nov. Previous worldwide geographic and stratigraphic records of taxa are detailed.
Resumen. Estudio sistemático palinológico de la Formación Cortaderas, (Mississippiano), Cuenca R ío Blanco, Argentina. Parte I. Asociaciones palinológicas bien preservadas han sido obtenidas de la Formación Cortaderas, de la Cuenca de Río Blanco, noroeste de Argentina. Es presentado el marco geológico y la procedencia de las muestras palinológicas obtenidas. En esta primera parte, 52 especies son descriptas de un total de 78 especies de esporas reconocidas. Cinco nuevas especies son propuestas: Verrucosisporites incompositus sp. nov., Verrucosisporites perbrevis sp. nov., Anapiculatisporites exiguus sp. nov., Anapiculatisporites expolitus sp. nov. y Dibolisporites insolitus sp. nov. Se detalla la distribución geográfica mundial y estratigráfica de los diferentes taxones.
Key words. Argentina; Cortaderas Formation; Mississippian; Palynology; Systematics.
Palabras clave. Argentina; Formación Cortaderas; Mississippiano; Palinología; Sistemática.
Introduction
With the exception of the Australian microfloristic
associations, palynological assemblages from
Mississippian of Gondwana are poorly known, especially
in South America, where few records are available.
Bolivian palynomorphs of Late Devonian-
Mississippian age were reported by Vavrdová et al.
(1991, 1993, 1996), and a Mississippian assemblage
was described for the Retama Formation by Azcuy
and Ottone (1987). Daemon (1974, 1976) and Melo
and Loboziak (2000, 2003) have investigated rich palynofloras
from the Mississippian sequences of the
Parnaíba and Amazon Basin in Brazil. In the
Peruvian Ambo Formation, Parnaíba palynomorphs
have been recorded by Doubinger and Alvarez
Ramis (1980) and Azcuy and di Pasquo (2005, 2006).
Subsurface strata located in the Colombian Llanos
Orientales Basin were refered to Mississippian on the
basis of miospore assemblages studied by Dueñas
and Césari (2005, 2006).
In Argentina, only three formations have yielded
palynological assemblages. The earliest record was
reported by Sessarego and Césari (1989) in the
Calingasta-Uspallata Basin, where Mississippian
spores (Viséan) were recovered from the El Ratón
Formation. New findings for this unit were listed by
Amenábar and di Pasquo (2006). A rich palynoflora
was illustrated originally from the Malimán
Formation in the Rio Blanco Basin by Césari and
Limarino (1995), and more recently has been re-investigated
by Amenábar (2006), Pérez Loinaze
(2005), Rodríguez Amenábar and di Pasquo (2004),
Rodríguez Amenábar et al. (2003) and Amenábar et
al. (2006). Miospore assemblages from the overlying
Cortaderas Formation have been dated as
Mississippian by Limarino and Césari (1992) and
Pérez Loinaze and Césari (2003). Nevertheless, only
few a species have been systematicaly described for
this formation (Pérez Loinaze, 2005).
This contribution is part of the Ph.D. Thesis of the
author that comprises the study of the floristic
change which occurred during the Mississippian- Pennsylvanian interval in western Argentina. The
aim of this first part is to describe some of the
Mississippian spores from the Cortaderas Formation.
The next contribution will complete the descriptions
of spores and microplankton, increasing the palynologycal
knowledge of Mississippian sequences in
South America.
Geological setting
The Late Paleozoic Río Blanco Basin shows one of the more complete stratigraphic records of Carboniferous sedimentation in South America encompassing a large portion of Precordillera and Frontal Cordillera in northwest Argentina (La Rioja and San Juan provinces). Carboniferous sediments have been included in two different stratigraphic units: the Mississippian Angualasto Group (Limarino and Césari, 1992) and the Pennsylvanian Quebrada Larga Formation (Scalabrini Ortiz, 1973). In the current study area, the Angualasto Group is divided in the Malimán and Cortaderas formations in ascending stratigraphic order with both having their type localities in the Cortaderas Creek (figure 1).
Figure 1. Geological map and geographical location of the studied area, showing the different facies associations recognized in the
Cortaderas Formation / mapa geológico y ubicación geográfica del area de estudio, mostrando las diferentes asociaciones de facies reconocidas en la Formación Cortaderas.
The name Cortaderas Formation was introduced
by Scalabrini Ortiz (1970, 1973) to describe a thick (up
to 1160m) siliciclastic sequence composed of scarce
conglomerates, sandstones and shales. These rocks
partially correlate with the upper part of Volcán
Formation as was defined for the east by Furque
(1963). The Cortaderas Formation was originally
considered, based on stratigraphic correlations, to be
Pennsylvanian-Lower Permian by Scalabrini Ortiz
(1970; 1973). The later finding by Limarino y Césari
(1992) of palynological remains allows reassigning
the unit to the Mississippian. As shown in figure 1 Cortaderas Formation outcrops in the core of an anticline
structure resulting from the uplift of the Sierra
de Volcán during the Andean Orogeny (Miocene). In
the anticline core, diamictites and shales bearing
dropstones belonging to the Upper Member of
Cortaderas Formation are reported. According to
Cardó et al. (2001), the diamictites are overlain by
arkosic sandstones and conglomerates included in
the Quebrada Larga Formation (Pennsylvanian)
where Carrizo (1990) reported megafloristic remains
assignated to the Late Carboniferous NBG Biozone.
The Cortaderas Formation has been divided in
five facies associations (figure 2): 1. basal conglomerate
(FA1), 2. fining-upward cycles from conglomerates
to sandstones (FA2), 3. thickening-upward cycles
(FA3), 4. amalgamated sandstones (FA4) and 5.
interbedded shales and diamictites (FA5). FA1 (106
m in thick) comprises coarse-grained conglomerates,
fine-grained sandstones and green mudstones with
occasional thin coal beds. These rocks were interpreted
as deposited in fandelta environments (Limarino
and Caselli, 1992), including gravelly mouth bars deposits,
fine-grained sediments of overbank areas and
small swamps where organic-rich sediments accumulated.
FA2 is composed of fining-upward cycles (up to
4 m thick) from conglomerates in the base to finegrained
sandstones (or mudstones) at the tops, which
would have been deposited by high sinuosity rivers.
Figure 2. Schematic stratigraphic section of the Cortaderas
Formation in the Cortaderas Creek, showing the provenance of
palynological samples (BA Pal) / sección estratigráfica esquemática de la Formación Cortaderas en la quebrada Cortaderas, mostrando la ubicación de las muestras palinológicas (BA Pal).
FA3 (240 m thick) forms thickening-upward cycles composed of greenish grey fine- and very finegrained sandstone covered by medium-grained sandstones representing marginal marine environments. FA4 reaches 540 m in thickness, and is composed of a monotonous sequence of grey-yellow to red fine- and coarse-grained sandstones, probably deposited in shallow marine environment. Finally, the FA5, up to 106 m of thickness, consists of rhythmically stratified, greenish grey laminated claystones and siltstones, laminated mudstones with dropstones and different types of diamictites. All these sediments were interpreted as being formed in a glacimarine environment sporadically dominated by gravity flow processes.
Material and methods
The present paper is based on 50 palynological
samples obtained from the five described facies associations
(FA) in the Cortaderas Creek area (29º 45'S,
69º 02'W), western flank of the Sierra de Volcan, San
Juan Province (figure 1). The location of samples is
given in figure 2, which shows that associations FA1
and FA5 provided the majority of the material (16
samples and 27, respectively). The rest of the fossiliferous
levels are chiefly concentrated in fine-grained
shallow marine sediments.
Laboratory procedures for extraction and concentration
of palynomorphs followed conventional
practices. Carbonates and silicates were removed by
treatment with hydrochloric and hydrofluoric acids.
The palynological residues were sieved on a 17 µm
mesh and mounted in unstained glycerin jelly. The
photographs were taken with a Nikon Coolpix 995
adapted to an Olympus BX 51 binocular microscope.
All illustrated specimens are deposited in the
Palynological Collection of the Museo Argentino de
Ciencias Naturales "Bernardino Rivadavia" (BA
Pal) and are identified with England Finder coordinates.
The suprageneric classification of the spores follows
the classical scheme of Dettmann (1963).
Systematic palaeontology
Anteturma PROXIMEGERMINATES Potonié 1970
Turma TRILETES Reinsch emend. Dettmann 1963
Suprasubturma ACAVATITRILETES Dettmann 1963
Subturma AZONONOTRILETES Luber emend. Dettmann
1963
Infraturma LAEVIGATI (Potonié and Kidston) Potonié
1956
Genus Punctatisporites Ibrahim emend. Potonié and Kremp 1954
Type species. Punctatisporites punctatus (Ibrahim) Ibrahim 1933.
Punctatisporites irrasus Hacquebard 1957 Figure 3.8
Figure 3. 1, Punctatisporites minutus Kosanke, BA Pal 5741-1: T49/2; 2, Retusotriletes anfractus Azcuy, BA Pal 5739-2: X27/0; 3, Retusotriletes sp. 2, BA Pal 5747: E44/4; 4, Punctatisporites planus Hacquebard, BA Pal 5741-1: E50/2; 5, Retusotriletes sp. 1, BA Pal
5741-1: C58/3; 6, Retusotriletes sp. cf. R. leptocetun Higgs, BA Pal 5748-2: G55/0; 7, Punctatisporites sp. 2, BA Pal 5743-1: C58/3; 8, Punctatisporites irrasus Hacquebard, BA Pal 5741-1: Y39/1; 9, 12, Cyclogranisporites sp. cf. C. firmus Jones and Truswell; 9, BA Pal
5754: T49/3; 12, BA Pal 5756: C51/0; 10, Punctatisporites sp. 1, BA Pal 5741-1: P32/4; 11, 14, Cyclogranisporites sp.; 11, BA Pal 5788-10:
K35/2; 14, BA Pal 5744-1: X41/0; 13, Verrucosisporites baccatus Staplin, BA Pal 5760-5: S29/4. Scale bar: 20 µm / escala gráfica: 20 µm.
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae straight, simple, length up to
two-thirds of spore radius. Exine laevigate, about 1-2
µm thick. Compression folds common, particularly
in the equatorial regions.
Dimensions. Equatorial diameter: 45 (55) 63 µm (26
specimens).
Comparisons. The specimens are very similar to the
originally described by Hacquebard (1957), but some
have smaller equatorial diameters. Punctatisporites
glaber Playford 1964 has more rigid exine and compression
folds are not common.
Previous records. Early Upper Carboniferous: Argentina,
Paganzo Basin, Lagares Formation (Menéndez
and Azcuy, 1969), Agua Colorada Formation
(Gutiérrez, 1988), Tarija Basin, Macharetí and
Mandiyutí Groups (di Pasquo, 2003; di Pasquo et al.,
2001). Lower Carboniferous-Upper Carboniferous:
Saudi Arabia (Clayton, 1995); USA (Felix and Burbridge,
1967). Lower Carboniferous: Alaska (Ravn,
1991); Canada (Hacquebard, 1957; Utting et al., 1989;
Utting, 1987a, 1987b); China (Zhu, 1993); Colombia
(Dueñas and Césari, 2005; 2006); Egypt (Kora, 1993);
Scotland (Neves and Ioannides, 1974); Ireland
(Owens et al., 1977). Devonian-Carboniferous boundary:
Ghana (Atta Peters and Yorke, 2003); Turkey
(Higgs et al., 2002).
Punctatisporites minutus Kosanke 1950 Figure 3.1
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae straight, simple, occasionally
with raised narrow lips, length three-quarter of spore
radius. Exine laevigate, about 1-1.2 µm thick.
Compression folds are common.
Dimensions. Equatorial diameter: 33 (36) 41 µm (12
specimens).
Comparisons. Punctatisporites minutus Kosanke 1950
resembles this material but always possesses simple
laesurae. Punctatisporites debilis Hacquebard 1957 differs
by its smaller size and infragranular exine.
Punctatisporites densiminutus Staplin 1960 has thicker
exine.
Previous records. Early Upper Carboniferous: China,
Namurian (Zhu, 1993); Great Britain (Smith and
Butterworth, 1967); Saudi Arabia (Clayton, 1995);
USA (Kosanke, 1950). Lower Carboniferous: Canada
(Utting, 1987a); Saudi Arabia (Clayton, 1995);
Scotland, (Love, 1960); USA (Clayton et al., 1998).
Devonian-Carboniferous boundary: Germany (Higgs
and Streel, 1984); Ireland (Clayton et al., 1982);
Turkey (Higgs et al., 2002).
Punctatisporites planus Hacquebard 1957 Figure 3.4
Description. Spores radial, trilete. Amb triangular
circular to subcircular. Laesurae straight, with raised narrow lips, length two-thirds of spore radius. Exine
laevigate, about 2-2.5 µm thick. Compression folds
are uncommon.
Dimensions. Equatorial diameter: 46 (60) 76 µm (15
specimens).
Comparisons. Punctatisporites aerarius Butterworth
and Williams 1958 differs in having microgranular
exine. Punctatisporites densiminutus Staplin 1960 has a
smaller equatorial diameter.
Previous records. Early Upper Carboniferous: Saudi
Arabia (Clayton, 1995). Lower Carboniferous: Saudi
Arabia (Clayton, 1995); Canada (Utting, 1987a,
1987b); USA (Clayton et al., 1998); Turkey (Higgs et
al., 2002). Devonian-Carboniferous boundary:
Germany (Higgs and Streel, 1984); Ireland (Clayton
et al., 1982). Upper Devonian: Libya (Coquel and
Moreau-Benoit, 1986). Devonian: Argentina, Chigua
Formation (Amenábar et al., 2006).
Punctatisporites sp. 1 Figure 3.10
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae straight, with raised narrow
lips, reaching almost the equatorial margin. Exine
laevigate, about 2-4 µm thick. Compression folds are
uncommon.
Dimensions. Equatorial diameter: 50 (61) 74 µm (30
specimens).
Comparisons. Punctatisporites solidus Hacquebard
1957 is distinguished by its prominent lips and
Punctatisporites punctatus Ibrahim 1933 differs in having
a thinner exine and simple laesurae.
Punctatisporites sp. 2 Figure 3.7
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae distinct, straight, with narrow
lips, length three-quarters of spore radius. Exine micropunctate,
about 2.5-4 µm thick.
Dimensions. Equatorial diameter: 66 (76) 90 µm (22
specimens).
Comparisons. Punctatisporites subtritus Playford and
Helby 1968 possesses a microgranular exine. Punctatisporites
ocellatus Sullivan 1964 and Punctatisporites
pseudofoveosus Azcuy 1975 differ in having simple
laesurae and smaller equatorial diameters. Punctatisporites
minutus Kosanke 1950 and Punctatisporites
punctatus Ibrahim 1933 are smaller in size and the latter
species has a thinner exine. Punctatisporites sp. 1
possesses a laevigate exine.
Infraturma RETUSOTRILETI Streel 1974
Genus Retusotriletes (Naumova) Streel 1964
Type species. Retusotriletes simplex Naumova 1953.
Retusotriletes anfractus Menéndez and Azcuy 1969 Figure 3.2
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae straight, slightly sinuous, with
raised narrow lips, length two-thirds of spore radius
with imperfect curvaturae. Exine laevigate,
about 1-1.5 µm thick. Compression folds are not
common.
Dimensions. Equatorial diameter: 31 (42) 48 µm (13
specimens).
Comparisons. Retusotriletes communis Naumova 1953
differs in having perfect curvaturae and a larger
equatorial diameter. Retusotriletes famenensis Naumova
1953 has prominent lips and perfect curvaturae.
Previous records. Early Upper Carboniferous:
Argentina, Paganzo Basin, Lagares Formation (Menéndez
and Azcuy, 1969), Jejenes Formation (González
Amicón, 1973; Gutiérrez and Césari, 1986), Agua Colorada
Formation (Menéndez and González Amicón,
1979; Gutiérrez, 1988; Vergel and Luna, 1992), Santa
Máxima Formation (Ottone, 1989), Tupe Formation
(Ottone and Azcuy, 1990), Tarija Basin, Macharetí
Group (di Pasquo, 2003). Lower Carboniferous: Perú,
Ambo Formation (Azcuy and di Pasquo, 2005).
Retusotriletes sp. cf. R. leptocentrum Higgs 1975 Figure 3.6
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae straight, slightly sinuous, extending
almost to equator, accompanied by thin lips.
The laesurae end in perfect curvaturae. Exine laevigate,
about 1.5-2 µm thick.
Dimensions. Equatorial diameter: 65-83 µm (7 specimens).
Comparisons. Retusotriletes leptocentrum Higgs 1975
resembles these specimens but differs by having simple
laesurae.
Retusotriletes sp. 1 Figure 3.5
Description. Spores radal, trilete. Amb circular to
subcircular. Laesurae straight to slightly sinuous, extending
almost to the equator, accompanied by thin
lips.The laesurae ends in curvaturae imperfectae.
Exine laevigate, about 1.5-3 µm thick.
Dimensions. Equatorial diameter: 36 (42) 67 µm (48
specimens).
Comparisons. Retusotriletes anfractus Menéndez and
Azcuy 1969 resembles these specimens but differs in
having a shorter and more sinuous laesurae not extending
to the equator.
Retusotriletes sp. 2 Figure 3.3
Description. Spores radial, trilete. Amb circular to
rounded subtriangular. Laesurae straight to slightly
sinuous, extending almost to the equator, accompanied
by thin lips. The laesurae terminate in perfect
curvaturae and are enhanced by thickenings of 1-2
µm high. Exine laevigate, about 0.8-1.5 µm thick.
Dimensions. Equatorial diameter: 33 (58) 53 µm (42
specimens).
Comparisons. Retusotriletes miriabilis (Neville)
Playford 1978 possesses a thicker exine and perfect
curvaturae enhanced by higher thickenings. Retusotriletes
famenensis Naumova 1953 differs by possessing
laesurae with prominent lips. Retusotriletes
sp. 1 presents imperfecta curvaturae.
Infraturma APICULATI Bennie and Kidston emend.
Potonié 1956
Subinfraturma GRANULATI Dybová and Jachowicz
1957
Genus Cyclogranisporites Potonié and Kremp 1954
Type species. Cyclogranisporites leopoldi (Kremp) Potonié and Kremp 1954.
Cyclogranisporites sp. Figure 3.11, 14
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae straight, with raised narrow
lips, length three-quarters of spore radius. Exine with
densely distributed grana (0.5-1.5 µm wide at base
and 0.3-0.5 µm high), circular to subcircular in plan
view. Exine about 0.8-2 µm thick. Compression folds
are common.
Dimensions. Equatorial diameter: 42 (53) 68 µm (26
specimens).
Comparisons. Cyclogranisporites pisticus Playford
1978 differs in having simple laesurae and grana (0.4-
2.5 µm wide at base and 0.2-0.3 µm high). Cyclogranisporites
commodus Playford 1964 resembles our
specimens but differs in the presence of simple laesurae
and smaller grana on the proximal face.
Cyclogranisporites multigranus Smith and Butterworth
1967 has shorter, simple laesurae and smaller grana
(0.5 µm wide at base and high). Cyclogranisporites
isostictus Balme 1988 differs in having simple laesurae,
an exine 2-3 µm thick and larger diameter (54-
85 µm). Cyclogranisporites fandersii Higgs 1996 possesses
simple laesurae, a thicker exine (2-5 µm) and
grana commonly fused to form sinuous to irregular
chains. Cyclogranisporites lasius (Waltz) Playford 1963
has simple laesurae, a thicker exine 1-3 µm and larger
diameter (50-88 µm). Cyclogranisporites firmus
Jones and Truswell 1992 has simple laesurae and a
thicker exine. Cyclogranisporites aureosus Mädler 1964
possesses a thicker exine and simple laesurae.
Cyclogranisporites sp. cf. C. firmus Jones and Truswell 1992 Figure 3.9, 12
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae simple, straight, length about
three-quarters of spore radius, usually indistinct.
Sculptural elements of 0.5-4 µm in basal width and
0.3 (0.5) 1 µm high, densely distributed, delineate a
fine negative reticulum, polygonal to subpolygonal
in plan view, up to 1 µm apart (typically 0.5 µm).
Exine about 3.5-5 µm thick. Secondary exinal folds
usually evident.
Dimensions. Equatorial diameter: 47 (59) 80 µm (34
specimens).
Comparisons. Cyclogranisporites firmus Jones and
Truswell 1992 is very similar to these specimens in its
thick exine and sculpture but differs in the smaller
height of the sculptural elements (0.2 µm). Cyclogranisporites
plicatus Pérez Loinaze and Césari 2004 possesses
a thinner exine. Cyclogranisporites pisticus Playford
1978 has a thinner exine (1.3-2.5 µm) and grana
up to 0.5 µm high. Verrucosisporites endicotensis (Anderson)
Backhouse 1988 possesses polygonal verrucae
and short rugulae among its sculptural elements. Verrucosisporites
depressus Winslow 1962 and Verrucosisporites
verrucosus (Ibrahim) Ibrahim 1933 differ in having
thinner exines (1.3 µm and 2-3 µm, respectively).
Subinfraturma VERRUCATI Dybová and Jachowicz
1957
Genus Verrucosisporites (Ibrahim) Smith and Butterworth 1967
Type species. Verrucosisporites verrucosus Ibrahim 1933.
Verrucosisporites baccatus Staplin 1960 Figure 3.13
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae simple, straight, usually open,
length up to two-thirds of spore radius. Proximal
surface laevigate. Distal surface and equatorial region
sculptured with verrucae and grana of 1-4 µm
wide at base and 1.5-4 µm high, subcircular in plan
view and with rounded to flattened apices, discrete
and irregularly distributed. Considerable areas of exine
may be without ornament. The density of sculptural
elements varies considerably among specimens.
Exine about 1.5-2.5 µm thick.
Dimensions. Equatorial diameter: 58 (67) 80 µm (13
specimens).
Previous records. Lower Carboniferous: Argentina,
Malimán Formation (Amenábar, 2006; Amenábar et
al., 2006; ); Alaska (Ravn, 1991); Canada (Staplin,
1960); Scotland (Love, 1960; Neves and Ioannides,
1974); Great Britain (Smith and Butterworth, 1967);
Poland (Filipiak, 2004).
Verrucosisporites congestus Playford 1963 Figure 4.1
Figure 4. 1, Verrucosisporites congestus Playford, BA Pal 5756: Y47/2; 2, 3, Verrucosisporites quasigobbettii Jones and Truswell; 2, BA
Pal 5756: X48/0; 3, BA Pal 5788-7: B46/3; 4-6, Verrucosisporites incompositus sp. nov.; 4, BA Pal 5788-10: M38/3; 5, BA Pal 5788-7: T43/3; 6, holotype/ holotipo, BA Pal 5790: N27/3; 7, 8, 13, Verrucosisporites perbrevis sp. nov.; 7, BA Pal 5730-4: O28/1; 8, holotype/ holotipo,
BA Pal 5757-5: K30/3; 13, BA Pal 5729-2: N45/0; 9, Verrucosisporites sp. 1, BA Pal 5788-11: N44/3; 10, 14, Verrucosisporites sp. 2, BA Pal
5729-2: N30/2; 10, Proximal surface/cara proximal; 14, Distal surface / cara distal; 11, Verrucosisporites sp. cf. V. nitidus Playford, BA Pal
5755: Y29/2; 12, Verrucosisporites cortaderensis Pérez Loinaze, BA Pal 5756: R49/4. Scale bar: 20 µm / escala gráfica: 20 µm.
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae simple, straight, length about
three-quarters of spore radius, usually indistinct because
of comprehensive verrucae. Sculpture strongly
developed, composed by large verrucae (6-18 µm
wide at base and 3-8 µm high), discrete, subcircular
amb and usually rounded apex, closely spaced, separated
by more or less regular and continuous negative
microreticulum. Exine about 2-4 µm thick.
Dimensions. Equatorial diameter: 66 (74) 82 µm (23
specimens).
Comparisons. Following the criteria of Turnau et al.
(1994), V. congestus is distinguished from V. depressus
Winslow 1962 and V. nitidus Playford 1964 by an arbitrary
separation on basis of the greater basal diameter
of its sculptural elements. Verrucosisporites depressus possesses verrucae up to 3 µm in basal diameter, V. nitidus among 3-8 µm and V. congestus has
greater than 8 µm.
Previous records. Lower Carboniferous: Argentina,
Río Blanco Basin, Malimán Formation (Césari and
Limarino, 1992; Amenábar, 2006), Cortaderas
Formation (Césari and Limarino, 1993; Pérez Loinaze
and Césari, 2003); Libya (Coquel and Moreau-Benoit,
1986); Brazil (Melo and Loboziak, 2000); Peru (Azcuy
and di Pasquo, 2005); Canada (Utting, 1987b; Varma,
1969); USA (Clayton et al. 1998); Scotland (Utting et
al., 1989). Upper Devonian: Turkey (Higgs et al.,
2002); Libya (Coquel and Moreau-Benoit, 1986).
Verrucosisporites cortaderensis Pérez Loinaze 2005 Figure 4.12
Dimensions. Equatorial diameter: 60 (65) 87 µm (50
specimens).
Previous records. Lower Carboniferous: Argentina,
Río Blanco Basin, Malimán and Cortaderas
Formations (Pérez Loinaze, 2005).
Verrucosisporites papulosus Hacquebard 1957 Figure 5.8-9
Figure 5. 1, Anapiculatisporites austrinus Playford and Satterthwait, BA Pal 5740-1 : U54/4, 2, Anapiculatisporites austrinus Playford
and Satterthwait, BA Pal 5750-1: K26/4, 3, Apiculiretusispora microseta Ravn, BA Pal 5754: H25/3, 4-5, Anapiculatisporites exiguus sp.
nov., holotype / holotipo, BA Pal 5791-1: R56/1, 4, Proximal surface / cara proximal, 5, Distal surface / cara distal, 6, Apiculiretusispora microseta Ravn, BA Pal 5741-1: D29/2, 7, Anapiculatisporites kekiktukensis Ravn, BA Pal 5760-1: V48/3, 8-9, Verrucosisporites sp. 5,
BA Pal 5792: T38/1, 8, Distal surface / cara distal, 9, Poximal surface / cara proximal, 10, Apiculatisporis variornatus di Pasquo, Azcuy
and Souza, BA Pal 5761-4 : Z42/1, 11, Anapiculatisporites exiguus sp. nov., BA Pal 5741-1: Y47/1, 12, Anapiculatisporites exiguus sp.
nov., BA Pal 5741-1: R31/4, 13, Anapiculatisporites amplus Playford and Powis, BA Pal 5762-4: Z47/4, distal surface / cara distal, 14, Verrucosisporites sp. 3, BA Pal 5788-12: P51/4, 15, Verrucosisporites sp. 5, BA Pal 5729-2: L31/2, 16-19, Verrucosisporites sp. 4, 16, Proximal surface / cara proximal, 17, Anapiculatisporites semisentus Playford, BA Pal 5760-8: F52/4, 18, Anapiculatisporites kekiktukensis Ravn, BA Pal 5742-1: E24/4, 19, Distal surface / cara distal, BA Pal 5790: E24/4, 20, Anapiculatisporites semisentus Playford,
BA Pal 5762-6: V51/0. Scale bar: 20 µm / escala gráfica: 20 µm.
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae simple, straight, length threequarters
of spore radius, usually indistinct. Sculptural
elements 0.5-4 µm wide at base and about 0.5
µm high, densely distributed, polygonal to subpolygonal
in plan view, forming a more or less regular
and continuous negative microreticulum.
Sculptural elements are smaller on proximal face.
Exine about 2-2.5 µm thick.
Dimensions. Equatorial diameter: 60-68 µm (4 specimens).
Comparisons. Verrucosisporites depressus Winslow
1962 and Verrucosisporites verrucosus (Ibrahim) Ibrahim
1933 resemble the Argentinian specimens in
type of sculpture but differ in having similar size
sculptural elements on both proximal and distal surfaces.
Moreover, the latter species also differs in having
occasional rugulae.
Previous records. Lower Carboniferous: Argentina,
Río Blanco Basin (Amenábar et al., 2006, Césari and
Limarino, 1995); Canada, (Barss, 1967; Hacquebard,
1957; Playford, 1964; Utting, 1987 a and b; Varma,
1969); Bear Island, Arctic Ocean (Kaiser, 1970); China
(Kao, 1980). Upper Devonian: Libya (Coquel and
Moreau-Benoît, 1986).
Verrucosisporites quasigobbettii Jones and Truswell 1992 Figure 4.2-3
Description. Spores radial, trilete, originally spherical.
Amb circular to subcircular. Laesurae simple,
usually open, straight, length three-quarters of spore
radius. Sculpture composed of variably spaced, irregular,
coarse, discrete verrucae (3-12 µm wide and
2-6 µm high), sparcely distributed. Exine about 2-5
µm thick.
Dimensions. Equatorial diameter: 65 (71) 85 µm (16
specimens).
Comparisons. Verrucosisporites gobbettii Playford
1963 differs in having smaller sculpture.
The specimen illustrated by di Pasquo (2003) as V.
quasigobbettii (page 23, fig. L) seems to differ from the
original description in having more closely distributed
verrucaes. Since that material has not been described,
and the photographs reveal that is probably a
different species, this record is not considered valid.
Previous records. Upper Carboniferous: Argentina,
Paganzo Basin, Agua Colorada Formation (Gutiérrez
1988); Australia (Jones and Truswell, 1992; Playford
and Helby, 1968). Lower Carboniferous: Brazil,
Parnaíba Basin, Longa and Potí Formations, (Müller,
1962; Dino and Playford, 2002).
As was noted by Dino and Playford (2002), this
species was recorded in the Lower Carboniferous of
Bolivia by Lobo Boneta (1975), but this material is unsupported
by either photographs or any description
and it is therefore unreliable.
Jones and Truswell (1992) have recognized this
species in Australian sediments dated as Namurian
to Early Permian. These authors identified reworking
in the Australian samples. That could explain this
unusual extended record.
Verrucosisporites incompositus sp. nov. Figure 4.4-6
Holotype. BA Pal 5790: N27/3, figure 4.6.
Diagnosis. Spores radial, trilete. Amb circular to subcircular.
Laesurae simple, straight, length threequarters
of spore radius, usually indistinct. Sculpture
composed by verrucae 2 (5) 11 µm wide at base and
1.5 (4) 7 µm high, rounded or blunt apex, irregularly
and sparcely distributed.
Dimensions. Equatorial diameter: 55(73)83 µm (58
specimens).
Type locality and stratigraphic horizon. La Cortadera
Creek, San Juan Province, Argentina. Cortaderas
Formation, Viséan.
Derivation of name. From the latin incompositus, irregular,
not in order.
Comparisons. This species is distinguished from the
others included in the genus by its thicker exine and
the irregular and loose distribution of its sculptural elements.
Verrucosisporites quasigobbettii Jones and Truswell
1992 differs by having a more regularly distributed
ornament and a smaller number of sculptural elements
on each surface. Verrucosisporites gibberosus
and Verrucosisporites mesogrumosus (Kedo) Byvscheva
1985 resemble the Argentinian species but both possess
a laevigate proximal face. Verrucosisporites gobbettii
Playford 1963 differs in having smaller verrucae
and Verrucosisporites baccatus Jones and Truswell 1992
possesses smaller verrucae and a thinner exine.
Verrucosisporites perbrevis sp. nov. Figure 4.7-8, 13
Holotype. BA Pal 5757-5: K30/3, figure 4.8.
Diagnosis. Spores radial, trilete. Amb circular to subcircular.
Laesurae simple, straight, length three-quarter
of spore radius. Verrucae 1.5 (3) 7 µm wide at base
and 1 (1.2) 3 µm high, polygonal to subpolygonal in
plan view, closely spaced, separated by more or less
regular and continuous negative microreticulum.
Exine about 2.5 (3.5) 6 µm thick.
Dimensions. Equatorial diameter: 53(61)81 µm (88
specimens).
Type locality and stratigraphic horizon. La Cortadera
Creek, San Juan Province, Argentina. Cortaderas
Formation, Viséan.
Derivation of name. From the latin perbrevis, that is
very short.
Comparisons. This species is characterized by a thick
exine ornamented by polygonal verrucae in plan
view that display a great variability in size between
specimens. The more typical aspect is illustrated in figures 4.8 and 13 and the figure 4.7 shows a variant
with the widest verrucae. Verrucosisporites variotuberculatus
Sullivan 1968 possesses smaller verrucae on
the proximal face. Verrucosisporites endicotensis Ravn
1991 has a larger diameter and verrucae up to 1 µm
in basal width and height. Verrucosisporites basilisticus Jones and Truswell 1992 possesses verrucae up to 0.5
µm high and an exine 1-3 µm in thickness. Verrucosisporites
sifati (Ibrahim) Smith and Butterworth 1967
has a thinner exine. Verrucosisporites verrucosus (Ibrahim)
Ibrahim 1933 differs by having rugulae and coni
as subordinate elements and a thinner exine. Verrucosisporites
depressus Winslow 1962 and V. grandiverrucosis
(Kosanke) Smith et al. 1964 have smaller verrucae
and thinner exines. Verrucosisporites donarii Potonié and Kremp 1955 possesses sculpture composed
by verrucae and rugulae and a thinner exine.
Verrucosisporites sp. cf. V. nitidus Playford 1964 Figure 4.11
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae simple, straight, usually indistinct,
length three-quarters of spore radius. Sculpture
composed of verrucae (4-8 µm wide at base and 3-
7µm high), discrete, circular to subcircular in crosssection
and usually with a rounded apex, closely
spaced. Elements delineate a fine negative microreticulum.
Exine about 2-4 µm thick.
Dimensions. Equatorial diameter: 56 (65) 78 µm (36
specimens).
Comparison. Although the proposals of Turnau et al.
(1994) does not define a maximum height for the verrucae
of V. nitidus, the specimens in the present study
differ in having taller verrucae than those usually assigned
to this species.
Verrucosisporites sp. 1 Figure 4.9
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae simple, straight, length threequarters
of spore radius, usually indistinct. Sculptured
by verrucae 3-15 µm wide at base and 2.5-7 µm
high, rounded or irregular apex, irregularly distributed,
occasionally grana are present, 2-6 µm apart.
Exine about 5-6 µm thick.
Dimensions. Equatorial diameter: 62 (72) 87 µm (30
specimens).
Comparisons. Verrucosisporites quasigobbettii Jones
and Truswell 1992 possesses a thinner exine and occasional
pila. Verrucosiporites gibberosus (Hacquebard)
Higgs et al. 1988 has a laevigate proximal surface.
Verrucosisporites grumosus (Naumova) Sullivan
1964 and Verrucosiporites congestus Playford 1963 differ
in having more densely distributed verrucae.
Verrucosisporites sp. 2 Figure 4.10, 14
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae simple, straight, length threequarters
of spore radius. Sculpture composed by verrucae
(2-14 µm wide at base and 2-9 µm high); discrete,
irregularly distributed. The size of sculptural
elements varies considerably in the same specimen.
Considerable areas of exine can be without ornament.
The density of sculptural elements varies considerably
among specimens. Exine about 1-1.5 µm
thick.
Dimensions. Equatorial diameter: 65-88 µm (5 specimens).
Comparisons. Verrucosisporites baccatus Staplin 1960
differs in having a smaller number of sculptural elements
on its exine. Verrucosisporites perverrucosus
(Loose) Smith 1971 has sculpture composed by verrucae
and rugulae.
Verrucosisporites sp. 3 Figure 5.14
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae simple, straight, length threequarters
of spore radius. Sculpture composed of verrucae
(3-10 µm wide at base and 2-5 µm high), discrete,
irregularly distributed, concentrated at the centre
of each surface, equatorial margin without ornament.
Exine about 2-2.5 µm thick.
Dimensions. Equatorial diameter: 52-70 µm (6 specimens).
Comparisons. Schopfites augustus Playford 1964 possesses
similar sculptural distribution but its proximal
face is laevigate. Verrucosisporites sp. 3 differs of
Verrucosisporites sp.1, Verrucosisporites sp. 2 and
Verrucosisporites sp. 4 by its characteristic disposition
of sculptural elements.
Verrucosisporites sp. 4 Figure 5.16, 19
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae simple or with thin lips,
straight, length three-quarters of spore radius. Sculpture
composed of verrucae (2-10 µm wide at base
and 1-9 µm high). The proximal surface possesses
smaller sculptural elements. Exine about 2-3.5 µm
thick.
Dimensions. Equatorial diameter: 60-68 µm (4 specimens).
Comparisons. Verrucosisporites italianensis Playford
and Helby 1968 and Verrucosisporites papulosus
Hacquebard 1957 closely compare with the
Argentinian specimens in the presence of smaller
verrucae in the proximal face but differ in the small
size of all sculpture. Verrucosisporites italianensis Ravn
1991 is moreover distinguished by possessing larger
equatorial diameter. Verrucosisporites sp. 4 differs of
Verrucosisporites sp. 1, Verrucosisporites sp. 2,
Verrucosisporites sp. 3 and Verrucosisporites sp. 5 by
presents smaller sculptural elements on proximal
surface.
Verrucosisporites sp. 5 Figure 5.15
Description. Spores radial, trilete. Amb circular to
subcircular. Laesuare simple, straight, length threequarters
of spore radius, usually indistinct. Sculpture
composed of verrucae 2.5-8 µm wide at base and 2-6
µm high, rounded apex, sparcely distributed. Coni
and grana are sometimes additionally present but in
minor proportions (1.5-5 µm wide at base and 2-7 µm
high), with rounded, acute or blunt apices. Exine
about 2.5-3 µm thick.
Dimensions. Equatorial diameter: 63-77 µm (6 specimens).
Comparisons. Verrucosisporites gobbettii Jones and
Truswell 1992 differs by having a more regular sculpture.
Verrucosisporites microtuberosus (Loose) Smith
and Butterworth 1967 resembles the Argentinian
specimens by the presence of verrucae, coni and
grana but differs in having smaller sculptural elements.
Verrucosisporites sp. 5 differs of Verrucosisporites
sp. 1, Verrucosisporites sp. 2, Verrucosisporites sp.
3 and Verrucosisporites sp. 4 by having cones and
grana between its sculptural elements.
Subinfraturma NODATI Dybová and Jachowicz 1957
Genus Apiculiretusispora Streel 1964
Type species. Apiculiretusispora brandtii Streel 1964.
Apiculiretusispora microseta Ravn 1991 Figure 5.3,6
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae straight to slightly sinuous
with thin lips, reaching almost to the equatorial margin.
Contact area laevigate. Distal surface and equatorial
margin sculptured by small coni with sharp or
rounded apices (0.5-1 µm wide at base and 0.5-1.2 µm
high), basal outline circular to subcircular. Exine
about 1-1.5 µm thick.
Dimensions. Equatorial diameter: 37 (43) 47 µm (39
specimens).
Comparisons. Apiculiretusispora microseta was originally
described by Ravn (1991) with laesurae simple.
However, some specimens described and illustrated
by Ravn (1991) (Plate 2, fig. 8-10) apparently have
thin lips as the material herein described.
Apiculiretusispora multiseta (Luber) Butterworth and
Spinner 1967 differs by having larger sculptural elements.
Apiculiretusispora nitida Owens 1971 has a
thicker exine in curvaturae area. Apiculiretusispora
coniferous (Kedo) Higgs 1996 has a larger equatorial
diameter.
Previous records. Lower Carboniferous: Alaska, Viséan (Ravn, 1991).
Genus Anapiculatisporites Potonié and Kremp 1954
Type species. Anapiculatisporites isselburgensis Potonié and Kremp 1954.
Anapiculatisporites amplus Playford and Powis 1979 Figure 5.13
Description. Spores radial, trilete. Amb subtriangular
with straight to slightly convex sides and rounded
apices. Laesurae straight with lips, occasionally
emphasized by exinal folding, reaching almost to the
equatorial margin. Proximal surface laevigate. Distal
surface sculptured with blunt or sharp coni (0.8-2.5 in
basal width and 1.5-4 high), circular to subcircular
bases, discrete, 0.5-4 µm apart. Sculptural elements
do not project equatorially on the interradial distal
area, but only in radial area. Exine about 1.2-1.5 µm
thick.
Dimensions. Equatorial diameter: 51-60 µm (2 specimens).
Comparisons. Although one specimen has a smaller
size than the originally described by Playford and
Powis (1979), it possesses the characteristic sculpture
of the Australian species.
The specimens described by Amenábar et al.
(2006) from the Malimán Formation display the
sculpture projecting in the interradial area not coinciding
with the original diagnosis.
Previous records. Lower Carboniferous: Argentina,
Calingasta-Uspallata Basin, El Ratón Formation (Sessarego
and Césari, 1989), Río Blanco Basin, Malimán
Formation (Amenábar, 2006; Amenábar et al., 2006).
Lower Carboniferous-Upper Carboniferous: Australia
(Playford and Powis, 1979; Dino and Playford, 2002).
Anapiculatisporites austrinus Playford and Satterthwait 1986 Figure 5.1-2
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae straight, reaching almost to the
equatorial margin, with lips of 1.8 µm high and 2.5
µm in overall width, ocassionally terminally bifurcated
forming imperfect curvaturae. Contact area laevigate,
distal surface and equatorial margin sculptured
with discrete coni (0.5-2.2 µm in basal width and
height), circular to subcircular bases, sharp or rounded
apex, 0.5-2 µm apart. Exine about 0.8-1.8 µm thick.
Dimensions. Equatorial diameter: 30 (39) 50 µm (11
specimens).
Comparisons. Some specimens possess smaller and
more densely distributed sculptural elements, but
though they are included within the variation described
by Playford and Satterthwait (1986). Anapiculatasporites
atheticus Neves and Ioannides 1974 is distinguished
by its larger sculpture. Anapiculatisprites
baccatus (Hoffmeister, Staplin and Malloy) Smith and
Butterworth 1967 and Anapiculatisporites centrosus
Higgs et al. 1988 differ by possessing more densely
distributed sculpture, and the latter species having
perfect curvaturae.
Previous records. Lower Carboniferous: Australia,
Viséan (Playford and Satterthwait, 1986).
A Chinese record of Yang (1999) from the Devonian-
Carboniferous boundary is considered with
caution because it was not described and the illustration
does not appear to support the assignation.
Anapiculatisporites kekiktukensis Ravn 1991 Figure 5.7, 18
Description. Spores radial, trilete. Amb subcircular
to rounded subtriangular. Laesurae straight to slightly
sinuous with lips 1.5-2 µm high and 0.5 µm in
basal width, occasionally emphasized by exinal folding,
reaching almost the equatorial margin. Proximal
surface psilate or with reduced sculpture. Distal surface
ornamented with minute cones with rounded
apices (0.8-1.5 µm in height and width) and subcircular
base. Exine about 1.5-3 µm thick.
Dimensions. Equatorial diameter: 41 (46) 70 µm (18
specimens).
Comparisons. Although Anapiculatisporites kekiktukensis possesses slightly thicker exine (4-5 µm), the specimens
studied in the present investigation have a similar
type and distribution of sculpture. Moreover, they
have an ill defined "crassitude" visible, as described
by Ravn (1991). Therefore, the present specific assignation
is considered the more appropiate. Apiculatasporites
microconus Richardson (in McGregor 1973) is
distinguished by its larger diameter (82-176 µm).
Previous records. Lower Carboniferous: Alaska, Viséan (Ravn, 1991).
Anapiculatisporites semisentus Playford 1971 Figure 5.17, 20
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae straight with lips 2 µm in
height and 1.5 µm in overall width, length about
three-quarters of spore radius. Contact area laevigate,
distal surface and equatorial margin sculptured
with minute grana (0.2-1 µm wide at base and up to
1 high), basal outline circular to subcircular, 0.2-2 µm
apart. Exine about 1.8-2.5 µm thick.
Dimensions. Equatorial diameter: 48(62)82 µm (10
specimens).
Comparisons. Massa et al. 1980 combined this species
with the genus Apiculiretusispora. This is characterized
by a laevigate proximal surface, sculpture
composed by grana, coni and/or spines and laesurae
terminally bifurcated forming imperfect or perfect
curvaturae. Playford (1971) described Anapiculatisporites
semisentus and remarked that some specimens
possess imperfect curvaturae, this character being
noted on some specimens in the present study.
Considering that Playford's specimens as well as the
Argentinian specimens only occasionally show imperfect
curvaturae is preferred to maintain this species
in the genus Anapiculatisporites.
Previous records. Lower Carboniferous: Argentina,
Río Blanco Basin, Malimán Formation (Césari and Limarino,
1995; Rodríguez Amenábar et al., 2003; Amenábar,
2006; Amenábar et al., 2006); Australia (Playford,
1971, 1978; Playford and Satterthwait, 1986);
Brazil, Poti Formation (Melo and Loboziak, 2000);
Bolivia, Kaka Formation (Fasolo et al., 2006); Peru,
Ambo Formation (Azcuy and di Pasquo, 2005).
Anapiculatisporites exiguus sp. nov. Figure 5.4-5, 11-12
Holotype. BA Pal 5791-1 : R56/1, figure 5.4-5.
Diagnosis. Spores radial, trilete. Amb circular to subcircular.
Laesurae straight to slightly sinuous, simple
or with thin lips, length about three-quarters of spore
radius, occasionally emphasized by exinal folding.
Contact area laevigate, distal surface and equatorial
margin sculptured with minute grana of 0.8-1 µm
width and 0.5-0.8 µm height, 0.2-1 µm apart. Exine
about 0.8 (1) 2 µm thick.
Dimensions. Equatorial diameter: 33 (48) 57 µm (82
specimens).
Type locality and horizon. The Cortadera Creek, San
Juan Province, Argentina. Cortaderas Formation, Viséan.
Derivation of name. From the latin exiguus, that is
small in size.
Comparisons. Anapiculatisporites kekiktukensis Ravn
1991 differs by its thicker exine and sculpture composed
by slightly larger cones. Anaplanisporites denticulatus Sullivan 1964 possesses some taller elements,
up to 3 µm high and verrucae arranged in a
quasi-concentric manner.
Anapiculatisporites expolitus sp. nov. Figure 6.8,12
Figure 6. 1-2, Anapiculatisporites sp. 4; 1, BA Pal 5739-2: X29/4; 2, Proximal surface / cara proximal, BA Pal 5738-3: C31/0; 3, Raistrickia rotunda Azcuy, BA Pal 5762-2: H43/2; 4, Raistrickia paganciana Azcuy, BA Pal 5761-5: B52/1; 5, 18, Dibolisporites malimanensis Pérez
Loinaze; 5, BA Pal 5754: A53/1; 18, BA Pal 5775-2: T47/1; 6, Dibolisporites disfacies Jones and Truswell, BA Pal 5733-5: V42/3; 7, 13- 17, Dibolisporites insolitus sp. nov.; 7, BA Pal 5760-1: K53/2; 13, Proximal surface / cara proximal, BA Pal 5760-1: M28/3; 14-15, holotype
/ holotipo, BA Pal 5760-5: B29/2; 16-17, Detail of sculpture / detalle de la escultura; 16, BA Pal 5760-1: M28/3; Q, BA Pal 5760-6:
H53/2; 8, 12, Anapiculatisporites expolitus sp. nov.; 8, BA Pal 5744-1: Y35/3, holotype / holotipo; 12, BA Pal 5742-2: M37/0; 9, Dibolisporites sp. 1, BA Pal 5773-1: E44/3; 10-11, Anapiculatisporites sp. 1., BA Pal 5760-1: S27/4; 10, Proximal surface / cara proximal; 11, Distal surface / cara distal; 19, Anapiculatisporites sp. 2, BA Pal 5760-1: E28/3; 20, Anapiculatisporites sp. 3, BA Pal 5733-5: K36/1.
Scale bar: 10 µm, except P and Q: 10 µm / escala gráfica: 20 µm, con excepción de P and Q: 10 µm.
Holotype. BA Pal 5744-1: Y35/3, figure 6.8.
Diagnosis. Spores radial, trilete. Amb circular to sub-
circular. Laesurae straight, simple, length about
three-quarters of spore radius. Proximal surface laevigate.
Distal surface sculptured with spines (0.8-1
µm wide and 3-5 µm high), sparsely distributed, 2.5-
5 µm apart. Sculptural elements do not project equatorially.
Exine about 1.5-2 µm thick.
Dimensions. Equatorial diameter: 59 (69) 90 µm (13
specimens).
Type locality and horizon. The Cortadera Creek, San
Juan Province, Argentina. Cortaderas Formation,
Viséan.
Derivation of name. From the latin expolitus, that is
smooth.
Comparisons. The studied specimens are referred to
Anapiculatisporites, which is characterized by specimens
with circular to subtriangular amb, proximal
surface laevigate and distal surface sculptured. Spinositriletes
is a similar taxon but was originally described
by Ravn (1991) to include triangular forms
sculptured with spines only in distal surface, that do
not project in the equatorial area.
Anapiculatisporites sp. 1 Figure 6.10-11
Description. Spores radial, trilete. Amb subcircular
to rounded subtriangular. Laesurae straight, with
lips 1.5-2 µm wide and 0.8-1 µm height, reaching almost
to the equatorial margin, occasionaly terminally
bifurcated in imperfect curvaturae. Contact area
laevigate. Distal surface and equatorial margin sculptured
with baculae and spines (0.8-3 µm wide and 2-
6 µm high), occasionally fusing to form short crests.
Spines with sharp apex and sometimes recurved,
baculae with blunt or sharp apex. Exine about 2-2.5
µm thick.
Dimensions. Equatorial diameter: 62 (70) 84 µm (13
specimens).
Comparisons. Aneurospora critia Turnau and Jakubowska
1989 differs by having an equatorial thickening
and sculpture composed by shorter spines.
Anapiculatisporites sp. 2 Figure 6.19
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae straight, with lips about 4-4.5
µm wide and 2.5 µm high, reaching almost to the
equatorial margin. Contact area laevigate, distal surface
and equatorial margin ornamented by minute
spines with sharp or blunt apices (0.3-0.05 µm wide
and 1-2.5 µm high), comprehensively distributed.
Exine about 1.5-2 µm thick.
Dimensions. Equatorial diameter: 68-85 µm (3 specimens).
Comparisons. Anapiculatisporites semisentus Playford
1971 differs with its sculpture composed mainly of
coni. Apiculatisporites adavalensis de Jersey 1966 resembles
these specimens with its ornament composed
of spines but they are present on proximal and
distal surfaces. Anapiculatisporites triangularis (Kosanke)
Potonié and Kremp 1955 has smaller ornamentation
elements.
Anapiculatisporites sp. 3 Figure 6.20
Description. Spores radial, trilete. Amb circular to
subcircular. Compression folds are common.
Laesurae usually indistinct, simple. Contact area laevigate.
Distal surface and equatorial margin ornamented
with short spines (1-2 µm wide and 2-3.5 µm
hight), sparsely distributed, 1-4 µm apart. Exine
about 0.5-0.8 µm thick.
Dimensions. Equatorial diameter: 74-92 µm (5 specimens).
Comparisons. Anapiculatisporites semisentus Playford
1971 differs by having sculpture composed of coni
and grana. Apiculatisporites adavalensis de Jersey 1966
has an exine ornamented on the proximal and distal
surfaces with spines. The sculpture does not project at
the equatorial margin in Anapiculatisporites expolitus.
Anapiculatisporites sp. 4 Figure 6.1-2
Description. Spores radial, trilete. Amb subtriangular
to subcircular. Laesurae straight, with lips about
1-1.5 µm wide and high, reaching almost to the equatorial
margin. Contact area laevigate, distal surface
and equatorial margin sculptured with coni and
grana (1-2 µm basal width and 0.5-4 µm high),
sparsely distributed, 1-5 µm apart. The sculptural elements
frequently basally fused into short, straight
and occasionally branched alignments. Exine about
0.8-1.5 µm thick.
Dimensions. Equatorial diameter: 42-55 µm (7 specimens).
Comparisons. Anapiculatisporites redactus Playford
1978 is similar by having an ornament composed by
spines frequently devoid of tips, resembling coni and
irregularly distributed, but differs because they are
not basally fused in short lines.
Genus Dibolisporites Richardson emend. Playford 1976
Type species. Dibolisporites echinaceous (Eisenack) Richardson 1965.
Dibolisporites malimanensis Pérez Loinaze 2005 Figure 6.5, 18
Remarks. The specimens here studied differ from
those originally described by having larger ornamentation
elements (0.8-4 µm wide and 1-6 µm high).
Dimensions. Equatorial diameter: 35 (46) 76 µm (71
specimens).
Comparisons. Amenábar et al. (2006) illustrated as
Dibolisporites microspicatus specimens from the
Malimán Formation that resemble the species originally
defined by Pérez Loinaze (2005) from that unit
and the Cortaderas Formation. Unfortunately,
Amenábar et al. (2006) did not provide a description
and therefore prevents any detailed comparison.
Previous records. Lower Carboniferous: Argentina,
Río Blanco Basin, Malimán and Cortaderas Formations
(Pérez Loinaze, 2005).
Dibolisporites disfacies Jones and Truswell 1992 Figure 6.6
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae straight, with lips about 2.5 µm
high, extending to or almost to the equatorial margin.
Contact area laevigate. Distal surface and equatorial
margin ornamented with biform sculptural elements.
Each element composed of verrucae with
rounded apex (1.5-2 µm in basal wide and 1-1.5 µm
high), rounded in plan view, surmounted by a simple
blunt spine. Sculptural elements discrete, 0.2-0.5
µm apart. Exine about 1-1.5 µm thick.
Dimensions. Equatorial diameter: 41 (46) 57 µm (12
specimens).
Comparisons. Dibolisporites disfacies, as originally described
by Jones and Truswell (1992) possesses laevigate
and hyaline proximal surface, frequently missing,
or with membranous and undulating labra. Some of the
specimens from Cortaderas Formation do not display
this fragile aspect of the proximal surface, but the rest of
characteristics are very similar to the Australian species.
Previous records. Upper Carboniferous: Argentina,
El Trampeadero Formation (Gutiérrez and Barreda,
2006), Tarija Basin, Mandiyutí and Macharetí Group
(di Pasquo et al., 2001; di Pasquo et al., 2003); Australia
(Powis, 1984; Jones and Truswell, 1992), Oman (Besems
and Schurmanm, 1987). Lower Permian: Antarctica
(Lindström, 1995); Oman and Saudi Arabia
(Stephenson and Filatoff, 2000; Stephenson and Osterloff,
2002; Stephenson et al., 2003; Stephenson, 2004;
Besems and Schuurmanm, 1987).
Dibolisporites insolitus sp. nov. Figure 6.7, 13-17
Holotype. BA Pal 5760-5 : B29/2, figure 6.14-15
Diagnosis. Spores radial, trilete. Amb circular to subcircular.
Laesurae straight, length about three-quarters
of spore radius, accompanied by thin lips, usually
indistinct because of densely distributed sculpture.
Exine ornamented by short crests, coni, baculae
and verrucae, in different proportions among
specimens, 3-13 µm wide and 3-8 µm high, irregularly
distributed, closely spaced or apart up to 10 µm.
Usually, the apical region of sculptural elements is
modified by subsidiary surmounted minute grana or
coni. Exine about 2-3 µm thick.
Dimensions. Equatorial diameter: 45 (68) 95 µm (14
specimens).
Type locality and horizon. The Cortadera Creek, San
Juan Province, Argentina. Cortaderas Formation,
Viséan.
Derivation of name. From the latin insolitus, that is
uncommon.
Comparisons. Dibolisporites montuosus Playford 1976
resembles the specimens here studied but differs by
its more densely distributed ornament and by having
processes apically modified by more varied subsidiary
ornament of verrucae, grana, coni or pila.
Dibolisporites sp. 1. Figure 6.9
Description. Spores radial, trilete. Amb subtriangular
to subcircular. Laesurae straight, length about
three-quarters of spore radius, accompanied by thin
lips. Contact area laevigate, distal surface and equatorial
margin with elongated, subcylindrical sculptural
elements, sometimes faintly noded in lateral
view, and frequently biform (1.5-4 µm wide and 3-8
µm high). Each element is composed by a conus or
baculum, with subcircular to subpolygonal bases and
end in a spine. This projection can be simple or bifurcated.
Sometimes spines or baculae with expanded
apices are present. Sculptural elements 0.5-5 µm
apart. Exine about 0.8-2 µm thick.
Dimensions. Equatorial diameter: 48 (58) 78 µm (12
specimens).
Comparisons. Dibolisporites malimanensis differs by
its smaller and more densely distributed ornament,
membranous proximal surface and less variable
sculpture. Raistrickia intonsa Playford 1978 has a smaller
diameter (22-46 µm) and the ornamentation elements
are not biform.
Genus Apiculatisporis Ibrahim emend. Potonié and Kremp 1954
Type species. Apiculatisporis grovensis Schoff, Wilson and Bentall 1944.
Apiculatisporis variornatus di Pasquo, Azcuy and Souza 2003 Figure 5.10
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae usually indistinct, straight,
simple, length about three-quarters of the spore radius.
Ornament composed of spines with round
apices, coni and baculae in variable proportions
among individual specimens; sometimes clavae are
present in minor proportions (1-3 µm wide and 2- 4.5
µm high), up to 2 µm apart. Exine about 2 µm thick.
Dimensions. Equatorial diameter: 38-67 µm (4 specimens).
Previous records. Upper Carboniferous: Argentina,
Paganzo Basin, Guandacol and Tupe Formations
(Ottone, 1991), Jejenes Formation (Césari and Bercowski,
1997), San Rafael Basin, El Imperial Formation
(García, 1995); Brazil, Paraná Basin (di Pasquo et
al., 2003; Lima et al., 1983; Souza, 2006).
Infraturma BACULATI Dybová and Jacchowicz 1957
Genus Raistrickia (Schoff, Wilson and Bentall) Potonié and Kremp 1954
Type species. Raistrickia grovensis Schoff, Wilson and Bentall 1944.
Raistrickia paganciana Azcuy 1975 Figure 6.4
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae usually indistinct. Exine ornamented
with prominent sculptural elements with
parallel sides and blunt or rounded apices (5-11 µm
wide and 4-7 µm high), width/length ratio 1/1 or
like verrucae irregularly distributed, usually superimposed.
Exine about 2 µm thick.
Dimensions. Equatorial diameter: 37-47 µm (5 specimens).
Comparisons. The specimens described here have
the same characteristic sculpture of R. paganciana, but
sometimes display a smaller diameter. They seem to
be cospecific with the original material illustrated by
Azcuy (1975, pp. 106, plate XVI, 106, and pp. 59, plate
XVII, 109).
The specimen illustrated by di Pasquo et al. 2001
(pp. 91, Fig. 3, C), as R. paganciana, from San Telmo
Formation, apparently has sculpture composed only
by verrucae with rounded profiles that are unlike
the material originally described by Azcuy
(1975). Unfortunately, these specimens were not
described, therefore a more detailed comparison is
not possible.
Previous records. Upper Carboniferous: Argentina,
Paganzo Basin, Malanzán Formation (Azcuy, 1975),
Guandacol and Tupe Formations (Ottone, 1991), Tarija
Basin, Mandiyutí Group (di Pasquo, 2003); Brazil,
Paraná Basin (Souza et al., 1993; Souza, 2006). Lower
Permian: Brazil, Paraná Basin (Dias Fabricio, 1993).
Raistrickia rotunda Azcuy 1975 Figure 6.3
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae usually indistinct because of
densely distributed sculpture. Exine ornamented
with discrete, club or mushroom-shaped processes
(3-7 µm wide and 3-5 µm high), some coni are often
additionally present in minor proportions. Exine
about 2-3 µm thick.
Dimensions. Equatorial diameter: 45-60 µm (2 specimens).
Comparisons. Raistrickia clavata Playford 1964 resembles
R. rotunda, however they differ because the
former has a smaller number of sculptural elements
on proximal surface and verrucae as subordinate elements.
Raistrickia pinguis Playford 1971 has only a
small proportion of club or mushroom-shaped
processes.
Previous records. Upper Carboniferous: Argentina,
Paganzo Basin, Malanzán Formation (Azcuy, 1975),
Lagares Formation (Morelli et al., 1984), Tupe
Formation (Césari, 1986; Ottone, 1991), Santa
Máxima Formation (Ottone, 1989), Agua Negra
Formation (Vergel and Luna, 1992), Guandacol
Formation (Ottone, 1991); Tarija Basin, Mandiyutí
Group (di Pasquo, 2003); Brazil, Paraná Basin (Lima et al., 1983; Souza et al., 1993; 1997; Souza, 1997; 2006).
Lower Permian: San Rafael Basin, El Imperial
Formation (García, 1995).
Infraturma MURORNATI Potonié and Kremp 1954
Genus Convolutispora Hoffmeister, Staplin and Malloy 1955
Type species. Convolutispora florida Hoffmeister, Staplin and Malloy 1955.
Convolutispora ampla Hoffmeister, Staplin and Malloy 1955 Figure 7.1
Figure 7. 1, Convolutispora ampla Hoffmeister, Staplin and Malloy, BA Pal 5762-8: Z38/4; 2, Convolutispora sp. cf. C. sp. 1 Menéndez
and Azcuy, BA Pal 5760-1: E44/2; 3, Convolutispora sp. cf. C. sculptilis Felix and Burbridge, BA Pal 5760-1: C28/3; 4, Ductilispora corrugata Pérez Loinaze, BA Pal 5794: L44/0; 5, Convolutispora sp., BA Pal 5761-5: Q44/3; 6, Ductilispora longuitudinalis Pérez Loinaze,
BA Pal 5754: O59/2; 7, Ductilispora circularis Pérez Loinaze, BA Pal 5788-12: D40/3; 8-11, Reticulatisporites magnidictyus Playford and
Helby; 8, MEB; 9, BA Pal 5760-1:132.8/25.5; 10, BA Pal 5754: Z50/1; 11, Detail of muri / detalle de los muros; 12, Dictyotriletes sp. cf. D. propius Playford, BA Pal 5764-4: G51/3; 13-14, ?Ductilispora sp.; 13, BA Pal 5754: U41/1; 14, MEB. Scale bar: 20 µm, except K: 10 µm / escala gráfica: 20 µm, con excepción de K: 10 µm.
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae simple, straight, usually indistinct
because to densely distributed ornament. Exine
compressively ornamented with vermiculate, anastomosing
ridges (1-2 µm wide at base and 1-1.2 µm
high). Exine about 1-1.5 µm thick.
Dimensions. Equatorial diameter: 48-59 µm (2 specimens).
Comparisons. Convolutispora finis Love 1960 differs
by having thinner ridges and athicker exine.
Previous records. This species, originally described
from the Upper Mississippian in the USA, is a common
component in palynofloras from the Viséan-
Namurian in Great Britain and USA. (Playford and
Satterthwait, 1985) and also has younger records
from the Westphalian (Braman and Hill, 1977;
Jachowicz, 1972); in Australia is known from
Tournaisian-Viséan sequences (Playford 1971, 1976,
1978; Playford and Satterthwait, 1985).
Convolutispora sp. cf. C. sp. 1 Menéndez and Azcuy 1973 Figure 7.2
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae straight, simple, length threequarters
of the spore radius. Exine ornamented with
vermiculate, sinuous, anastomosing and branching
muri (1.5-4 µm wide at base and 1-2 µm high), rounded
in profile, occasionally forming lumina of 2-4 µm
in the largest axis. Exine about 1.5 µm thick.
Dimensions. Equatorial diameter: 48-52 µm (2 specimens).
Comparisons. Convolutispora sp. 2 Menéndez and
Azcuy 1973 possesses straighter muri which anastomose
more frequently enclosing lumina.
Convolutispora sp. cf. C. sculptilis Felix and Burbridge 1967 Figure 7.3
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae simple, straight, usually indistinct
because of densely distributed ornament. Exine
sculptured with prominent, sinuous muri (3-5 µm
wide and 2-3 µm high), occasionally forming lumina
of 4-6 µm in the longer axes.
Dimensions. Equatorial diameter: 44-56 µm (2 specimens).
Comparisons. Convolutispora sculptilis as originally
described by Felix and Burbridge (1967) has taller
muri (3-6 µm). Convolutispora vermiformis Hughes
and Playford 1961 differs by having wider and taller
muri enclosing larger lumina.
Previous records. Lower Carboniferous-Upper
Carboniferous: USA (Felix and Burbridge, 1967).
Upper Carboniferous: Argentina, Paganzo Basin,
Malanzán Formation (Azcuy, 1975).
Convolutispora sp. Figure 7.5
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae simple, straight, usually indistinct
because of densely distributed ornament. Exine
ornamented with verrucae up to 5 µm wide and 2-3
µm high, short vermiculae, occasionally branching
and well rounded in profile and rugulae up to 2.5 µm
high. Sculptural elements densely distributed.
Dimensions. Equatorial diameter: 50-60 µm (2 specimens).
Comparisons. Convolutispora florida Hoffmeister,
Staplin and Malloy 1955 resembles the specimens described
here by having verrucae and short muri, but
differs in the larger size of the muri. Convolutispora
jugosa Smith and Butterworth 1967 and Convolutispora
insulosa Playford 1978 have a larger proportion
of rugulae between the sculptural elements.
Genus Reticulatisporites Ibrahim emend. Potonié and Kremp 1954
Type species. Reticulatisporites reticulatus (Ibrahim) Ibrahim 1933.
Reticulatisporites magnidictyus Playford and Helby 1968 Figure 7.8-11
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae straight, simple, length threequarters
of the spore radius. Exine ornamented with
a prominent perfect reticulum composed by muri (1-
3.5 µm wide at base and 3-17 µm high), enclosing
large polygonal lumina (8-28 µm in largest axes).
Occasionally muri are radially elongated on proximal
surface. Muri taper from relatively broad base to
sharp diaphanous tops. At the intersections of muri,
spine-like projections appear arising from the base of
muri. In specimens which are degraded, the membranous
part of muri may be eroded or missing, only
the more resistant bases and projections can be observed.
The floor of lumina is laevigate to microgranulate.
Exine about 2.5-6 µm thick.
Dimensions. Equatorial diameter: 61 (74) 97 µm (49
specimens).
Comparisons. Melo and Loboziak (2000) suggested
to included this species in the genus Cordylosporites
Playford and Satterthwait, which according to its
original diagnosis is characterized by having a perfectly
to imperfectly reticulate exine with distinctive
supramural projections as bacula, pila or verrucae.
As noted by Playford (1991) this species does not
possess these supramural projections and therefore,
it is here considered more appropriate for its inclusion
in the genus Reticulatisporites. Frequently, the
basal part of degraded muri may be misinterpreted
as supramural projections.
Previous records. Upper Carboniferous: Argentina,
Paganzo Basin, Agua Colorada Formation (Gutíerrez,
1988); Australia (Playford and Helby, 1968).
Lower Carboniferous: Brazil, Parnaíba Basin (Melo
and Loboziak, 2000); Amazonas Basin (Daemon and
Contreiras, 1971; Daemon, 1974; Loboziak et al. 1998);
Bolivia, Saipurú Formation (Suárez-Soruco and
Lobo-Boneta 1983; Suárez-Soruco and Lopez, 1983), Tupambi and Itacua Formations (Lobo-Boneta, 1975;
Rocha Campos and Archangelsky, 1985); Perú
(Azcuy and di Pasquo, 2005); Saudi Arabia (Clayton,
1995); Algeria (Lanzoni and Mangloire, 1969; Attar et
al., 1980); Libya (Massa et al., 1980).
Genus Dictyotriletes Naumova emend. Smith and Butterworth 1967
Type species. Dictyitriletes bireticulatus (Ibrahim) Potonié and Kremp 1954.
Dictyotriletes sp. cf. D. propius Playford 1978 Figure 7.12
Description. Spores radial, trilete. Amb circular to
subcircular. Laesurae straight, simple. Exine ornamented
with sinuous, thin muri (1.5-2 µm wide and
1.5 µm high) enclosing irregular to polygonal lumina
(5-7 µm in largest axes). Exine about 1 µm thick.
Dimensions. Equatorial diameter: 65 µm (1 specimen).
Comparisons. Dictyotriletes propius resembles the
specimens studied here by having sinuous muri enclosing
irregular reticulum, which frequently end
freely. Dictyotriletes densoreticulatus Potonié and
Kremp 1955 differs in having straighter muri.
Genus Ductilispora Pérez Loinaze 2005
Type species. Ductilispora circularis Pérez Loinaze 2005.
Ductilispora circularis Pérez Loinaze 2005 Figure 7.7
Dimensions. 38(51)58 µm (13 specimens).
Previous records. Lower Carboniferous: Argentina,
Río Blanco Basin, Malimán and Cortaderas Formations
(Pérez Loinaze, 2005).
Ductilispora longitudinalis Pérez Loinaze 2005 Figure 7.6
Dimensions. Equatorial diameter: 46 (59) 76 µm (12
specimens).
Previous records. Lower Carboniferous: Argentina,
Río Blanco Basin, Malimán and Cortaderas Formations
(Pérez Loinaze, 2005).
Ductilispora corrugata Pérez Loinaze 2005 Figure 7.4
Dimensions. Equatorial diameter: 41 (56) 70 µm (14
specimens).
Previous records. Lower Carboniferous: Argentina,
Río Blanco Basin, Malimán and Cortaderas Formations
(Pérez Loinaze, 2005).
? Ductilispora sp. Figure 7.13-14
Description. Spores radial, trilete. Amb circular to
subcircular, margin irregular. Laesurae usually indistinct.
Exine smooth, thin, approximately 0.8-1.5
µm wide, exine ornamented with prominent folds
or thickenings.
Dimensions. 75 (133) 192 µm (16 specimens).
Comparisons. The studied specimens are referred
with doubt to Ductilispora they have a sculptured
proximal surface. Rugospora Neves and Owens 1966
has a finer ornament and is cavate. Plicatispora Higgs
et al. 1988 is characterized by having curvaturae and
a distal surface sculptured by folds and fine muri.
Knoxisporites Potonié and Kremp 1954 presents distal
thickenings and a uniform cingulum.
Acknowledgments
Thanks are due to S.N. Césari for her critical comments, useful suggestions and invaluable help during all the development of this work, to O. Limarino for critical reading on early drafts of the manuscript and providing valuable advice, to E. Vera for reading the manuscript. Funding for this work was provided by the Agencia Nacional de Promoción Científica y Técnica, PICT 20752. The author is indebted to B. Owens and G. Clayton for their meticulous revisions of the manuscript as reviewers.
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Recibido: 10 de agosto de 2006.
Aceptado: 18 de setiembre de 2007.