ARTÍCULOS ORIGINALES
New palynological data from the Malanzán Formation (Carboniferous), La Rioja Province, Argentina
Valeria Perez Loinaze1
1División Paleobotánica, Museo Argentino de Ciencias Naturales "Bernardino Rivadavia", Av. Angel Gallardo 470, 1405 Buenos Aires, Argentina. loinazev@macn.gov.ar
Abstract. Palynological analysis of the Malanzán Formation in the Paganzo Basin, northwestern Argentina, has yielded six well-preserved palynological assemblages. Fifty-nine species were determined: 37 spores, 19 pollen grains, and 3 algae. Thirty-two of these species have not been recorded so far from the Malanzán Formation, and all pollen species are illustrated for the first time. These assemblages are referred to Sub-Biozone A of the Raistrickia densa-Convolutispora muriornata Biozone, Serpukhovian in age. Moreover, the abundance of monosaccate pollen grains might constrain the Malanzán Formation to the late Serpukhovian.
Resumen. Nuevos datos palinológicos de la Formación Malanzán (Carbonífero), Provincia de la Rioja, Argentina. Nuevas asociaciones palinológicas han sido obtenidas de la Formación Malanzán (Cuenca Paganzo), noroeste de Argentina. En las seis muestras fértiles estudiadas para este trabajo, se identificaron un total de cincuenta y nueve especies de palinomorfos: 37 especies de esporas, 19 de granos de polen y 3 especies de algas. Treinta y dos de las especies de palinomorfos reconocidas no habían sido reportadas hasta el momento para la unidad. Además, se ilustran por primera vez los granos de polen de la Formación Malanzán. Las asociaciones estudiadas son referidas a la Biozona Raistrickia densa- Convolutispora muriornata, y más precisamente a la sub Sub-Biozona A, de edad serpukhoviana. Además, la abundancia de granos de polen monosacados podría restringir a la Formación Malanzán al Serpukhoviano tardío.
Key words. Argentina; Malanzán Formation; Carboniferous; Palynology.
Palabras clave. Argentina; Formación Malanzán; Carbonífero; Palinología.
Introduction
Thick Upper Palaeozoic sequences located in western and northwestern Argentina and constituting the Río Blanco, Calingasta-Uspallata and Paganzo basins, represent one of the most continuous sedimentary sequences of this age in Gondwana. During the past two decades, a large number of systematicpalynological and biostratigraphic studies of these Carboniferous and Permian sediments have been published (e.g., Archangelsky et al., 1996; Césari and Limarino, 2002; Césari and Gutiérrez, 2001; Gutiérrez et al., 2003; Gutiérrez and Barreda, 2006). Despite this extensive research on the Late Palaeozoic, some issues are still unresolved, including the lack of a detailed characterization of the latest Mississippian palynological content. Such study would allow a clear distinction between Mississippian and Pennsylvanian palynofloras; i.e., those preserved in glaciogene sedimentary sequences deposited during the "Middle Carboniferous glaciation" event of the latest Mississippian-earliest Pennsylvanian, thus providing an important tool for stratigraphic correlations. In this work, new palynological assemblages recovered from fine-grained sediments of the Malanzán Formation, deposited in a fjord environment (during a postglacial transgressive event) and in a deltaic environment, are described in detail. The assemblages contain abundant monosaccate and bisaccate pollen grains, which are identified and illustrated for the first time from this formation.
Geological setting
Located in the northwestern region of Argentina, the Paganzo Basin includes sequences referred to the Paganzo Group by Azcuy and Morelli (1970) and dated as Serpukhovian-Permian. In the Sierra de los Llanos area (figure 1), this group is represented, in ascending order, by the Malanzán, Loma Larga, Solca, and La Colina formations (Andreis et al, 1986; Net, 1998). The Malanzán Formation is overlain conformably by conglomerates, sandstones and some mudstone beds of the Loma Larga Formation, and lies on crystalline basement of the Cambrian Pacatala Complex (Page et al., 2002) and the Ordovician Chepes Complex (Pieters and Skinom, 1997).
Figure 1. Geological map and geographic location of the fossiliferous localities (modified from Net , 1999) / mapa geológico y ubicación
geográfica de las localidades fosilíferas (modificado de Net, 1999).
The Malanzán Formation was originally defined
by Furque (1968) to include Late Palaeozoic sequences
outcropping in the Sierra de los Llanos (Bracaccini,
1946, 1948). Later, Azcuy (1975a) subdivided the Malanzán
Formation into three members, named Divisoria,
Estratos Carbonosos and Conglomerados Amarillo
Verdosos-Violáceos (in ascending order). A different
classification was proposed by Andreis et al.
(1986), who restricted the name Malanzán Formation
to the basal member (the "Divisoria Member" of Azcuy,
1975a). Moreover, Andreis et al. (1986) defined the
Loma Larga Formation to incorporate Azcuy's (1975a)
Estratos Carbonosos and Conglomerados Amarillo
Verdosos-Violáceos members. More recently, Net
(1998) and Net and Limarino (1999) included only the
Estratos Carbonosos Member in the Loma Larga
Formation, the upper member being considered part
of the overlying Solca Formation, defined by Andreis et al. (1986).
In the studied area, the Malanzán Formation, up
to 345 m thick, is segregated into five sections based
on lithology, sedimentary structures, and cyclicity
(figure 2). (1) The lowermost section, 85 m thick, is
composed of coarse-grained conglomerates, breccias,
massive or cross-stratified pebble sandstones, diamictites
and some shales with dropstones. These deposits
were interpreted by Andreis et al. (1986) as
formed in alluvial fans. (2) Grey laminated mudstones
with dropstones (55 m thick) bearing megaclasts
up to 20 cm in diameter. Some beds of massive
diamictites, up to 1 m thick, and beds of ripple crosslaminated
sandstones are intercalated within the
laminated mudstones. These deposits were interpreted
by Limarino and Césari (1988) and Sterren and
Martínez (1996) as having accumulated in glaciolacustrine
environments; however, Limarino et al. (2002) suggested that the sediments represent the beginning of the postglacial "Middle Carboniferous" transgression in the Paganzo Basin. (3) Black shales
and mudstones without dropstones, up to 35 m thick,
including thin beds of marls and ripple cross-laminated
fine-grained sandstones. This interval is believed
to have been deposited during the maximum
post-glacial flooding event (Limarino et al., 1996).
From palynological data, Gutiérrez and Limarino
(2001) suggested brackish conditions, with sedimentation
in palaeofjords, as previously inferred by Net
and Limarino (1999). New palynological samples
from this section are documented herein. (4) Largescale
cross-bedded sandstones reaching 50 m in
thickness, exhibiting megasets (up to 9 m thick) formed
by the progradation of Gilbert-type deltas (Sterren
and Martínez, 1996). Three palynological samples
were collected from this interval. (5) Interbedded
organic-rich mudstones and ripple cross-laminated
fine-grained sandstones (130 m thick) interpreted
as mouth bar deposits.
Figure 2. Schematic stratigraphic section of the Malanzán Formation,
showing the provenance of palynological samples (modified
from Net, 1999) / sección estratigráfica esquemática de la Formación
Malanzán, mostrando la ubicación de las muestras palinológicas (modificado
de Net, 1999).
Material and methods
The studied samples were obtained from the Malanzán Formation (Paganzo Basin) in La Rioja Province, Argentina (figures 1, 2). Three fossiliferous levels were sampled from outcrops of this formation at the La Chimenea locality in the Olta Valley; they consist of thin coal beds, interpreted as frontal bars of a delta, here designated as Section 4 (BA Pal 5811-13). Other three samples came from the Cuestita de la Herradura locality in the Malanzán valley, they consist of laminated mudstones without dropstones and are included in Section 3 (BA Pal 5814-16). All specimens illustrated herein (figures 3-6) are deposited in the palynological collection of the Museo Argentino de Ciencias Naturales "Bernardino Rivadavia" (BA Pal) and are slide-located by England Finder coordinates.
Figure 3. 1, Punctatisporites irrasus Hacquebard 1957 BA Pal 5811-1 : Y49/3; 2, Punctatisporites priscus Bharadwaj and Salujha 1965
BA Pal 5813-1 : X48/3; 3, Leiotriletes tiwarii (Saxena) Saxena 1993 1965 BA Pal 5811-2 : E50/4; 4, Retusotriletes sp. 2 BA Pal 5811-4
:P28/1; 5, Cyclogranisporites firmus Jones and Truswell 1992 BA Pal 5812-1 : B12/3; 6, Cyclogranisporites plicatus Perez Loinaze and
Césari 2004 BA Pal 5813-2 : H45/4; 7, Retusotriletes sp. 1 BA Pal 5811-4 : P28/2; 8, Punctatisporites sp. cf. P. glaber (Naumova) Playford
1962 BA Pal 5812-3 : W40/1; 9, Apiculiretusispora sp. cf. A. ralla (Menéndez and Azcuy) Menéndez and Azcuy 1971 BA Pal 5816-2 :
G32/3; 10, Anapiculatisporites sp. cf. A. concinnus Playford 1962 BA Pal 5816-2 : E45/3; 11, Cyclogranisporites rinconadensis Césari and
Limarino 2002 BA Pal 5811-4 : R40/2; 12, Verrucosisporites andersonii Backhouse 1988 BA Pal 5812-1 : M26/3; 13, Dibolisporites disfacies Jones and Truswell 1992 BA Pal 5813-1 : D23/2; 14, Granulatisporites sp. cf. G. austroamericanus Archangelsky and Gamerro 1979
BA Pal 5811-1 : K51/4; 15, Lophotriletes intermedius Azcuy 1975 BA Pal 5816-1 : V38/3; 16, Reticulatisporites asperidictyus Playford
and Helby 1968 BA Pal 5812-1 : D51/3; 17, Dictyotriletes cortaderensis Césari and Limarino 1987 BA Pal 5811-1 : N53/1; 18, Apiculiretusispora alonsoi Ottone 1989 BA Pal 5813-1 : S33/3; 19, Apiculatisporis variornatus di Pasquo et al. 2003 BA Pal 5815-3 : Z30/3;
20, Apiculatasporites caperatus Menéndez and Azcuy 1969 BA Pal 5813-2 : Q30/3; 21, Convolutispora muriornata Menéndez 1965 BA
Pal 5814- : J22/4; 22, Punctatisporites gretensis Balme and Hennelly 1956 BA Pal 5813-2 : Z55/3. Scale bar / escala gráfica: 20 μm.
The samples were processed using standard palynological techniques for extraction and concentration of palynomorphs. The organic residues were sieved using 20 μm mesh and later mounted in glycerine jelly on microscope slides.
Systematic palynology
The identified palynomorphs (spores, pollen grains and algae) are listed below in alphabetical order (* indicates first report from the Malanzán Formation). Only the species recognized for the first time in Argentina, and those with doubtful specific assignment, are described.
Spores
Anapiculatisporites sp. cf. A. concinnus Playford 1962
(figure 3.10)
Apiculatasporites caperatus Menéndez and Azcuy 1969
(figure 3.20)
Apiculatisporis variornatus di Pasquo et al. 2003* (figure
3.19)
Apiculiretusispora alonsoi Ottone 1989* (figure 3.18)
Apiculiretusispora sp. cf. A. ralla (Menéndez and
Azcuy) Menéndez and Azcuy 1971 (figure 3.9)
Convolutispora muriornata Menéndez 1965 (figure
3.21)
Cristatisporites inconstans Archangelsky and Gamerro
1979* (figure 4.7)
Cristatisporites menendezii (Menéndez and Azcuy)
Playford 1978 (figure 4.13)
Cristatisporites rollerii Ottone 1989* (figure 4.2)
Cristatisporites stellatus (Azcuy) Gutiérrez and Limarino
2001 (figure 4.3)
Cristatisporites sp.* (figure 4.5)
Cyclogranisporites firmus Jones and Truswell 1992*
(figure 3.5)
Cyclogranisporites plicatus Perez Loinaze and Césari
2004* (figure 3.6)
Cyclogranisporites rinconadensis Césari and Limarino
2002* (figure 3.11)
Dibolisporites disfacies Jones and Truswell 1992* (figure
3.13)
Dictyotriletes cortaderensis Césari and Limarino 1987
(figure 3.17)
Endosporites sp. * (figure 4.11)
Grandispora sp.* (figure 4.10)
Granulatisporites sp. cf. G. austroamericanus Archangelsky
and Gamerro 1979* (figure 3.14)
Grossusporites microgranulatus (Menéndez and Azcuy)
Perez Loinaze and Césari 2004* (figure 4.16)
Indotriradites volkheimeri (Azcuy) Perez Loinaze 2008
(figure 4.9)
Leiotriletes tiwarii (Saxena) Saxena 1993 (=Leiotriletes
virkii)* (figure 3.3)
Lophotriletes intermedius Azcuy 1975a (figure 3.15)
Microreticulatisporites punctatus Knox 1950* (figure
4.15)
Punctatisporites gretensis Balme and Hennelly 1956*
(figure 3.22)
Punctatisporites irrasus Hacquebard 1957* (figure 3.1)
Punctatisporites priscus Bharadwaj and Salujha 1965*
(figure 3.2)
Punctatisporites sp. cf. P. glaber (Naumova) Playford
1962 (figure 3.8)
Raistrickia paganciana Azcuy 1975a (figure 4.1)
Raistrickia rotunda Azcuy 1975a (figure 4.14)
Raistrickia sp. cf. R. radiosa Playford and Helby 1968*
(figure 4.8, 12, 17)
Reticulatisporites asperidictyus Playford and Helby
1968 (figure 3.16)
Reticulatisporites magnidictyus Playford and Helby
1968* (figure 4.6)
Retusotriletes sp. 1* (figure 3.7)
Retusotriletes sp. 2* (figure 3.4)
Vallatisporites ciliaris (Luber) Sullivan 1964 (figure
4.4)
Verrucosisporites andersonii Backhouse 1988* (figure
3.12)
Figure 4. 1, Raistrickia paganciana Azcuy 1975 BA Pal 5814-3: W53/3; 2, Cristatisporites rollerii Ottone 1989 BA Pal 5812-1: P35/2; 3,
Cristatisporites stellatus (Azcuy) Gutiérrez and Limarino 2001 BA Pal 5812-1: L27/2; 4, Vallatisporites ciliaris (Luber) Sullivan 1964
BA Pal 5816-2: W34/1; 5, Cristatisporites sp. BA Pal 5812-2: Z52/4; 6, Reticulatisporites magnidictyus Playford and Helby 1968 BA Pal
5812-2: Q27/2; 7, Cristatisporites inconstans Archangelsky and Gamerro 1979 BA Pal 5811-2: T39/4; 8, 12, 17, Raistrickia sp. cf. R. radiosa Playford and Helby 1968; 8, BA Pal 5812-1: A28/2; 12, BA Pal 5816-5: K40/4, sculptural detail/ detalle de la escultura; 17, BA Pal
5811-5: A28/3; 9, Indotriradites volkheimerii Azcuy 1975 BA Pal 5814-2: X29/3; 10, Grandispora sp. BA Pal 5812-1: J23/4; 11,
Endosporites sp. BA Pal 5812-3: O38/2; 13, Cristatisporites menendezii (Menéndez and Azcuy) Playford 1978 BA Pal 5811-2: O53/3; 14, Raistrickia rotunda Azcuy 1975 BA Pal 5815-3: H48/4; 15, Microreticulatisporites punctatus Knox 1950 BA Pal 5816-3: Y28/2; 16,
Grossusporites microgranulatus (Menéndez and Azcuy) Perez Loinaze and Césari 2004 BA Pal 5816-3: G32/4. Scale bar: 20 μm except
12: 10 μm / escala gráfica: 20 μm con excepción de 12: 10 μm.
Pollen grains
Caheniasaccites densus Lele and Karin emend. Gutiérrez
1993* (figure 5.12)
Cannanoropollis densus (Lele) Bose and Maheshwari
1968 (figure 5.8)
Cannanoropollis janakii Potonié and Sah 1960 (figure 5.3)
Cannanoropollis mehtae (Lele) Bose and Maheshwari
1968 (figure 6.4)
Circumplicatipollis plicatus Ottone and Azcuy 1988 (figure
5.14)
Colpisaccites granulosus Archangelsky and Gamerro
1979* (figure 5.15)
Costatascyclus crenatus Felix and Burbridge emend.
Urban 1971* (figure 5.13)
Crucisaccites monoletus Maithy 1965 (figure 5.10)
Divarisaccus stringoplicatus Ottone 1991* (figure 5.6)
Limitisporites rectus Leschik 1956* (figure 5.2)
Peppersites sp. cf. P. ellipticus Ravn 1979* (figure 5.1, 4)
Plicatipollenites gondwanensis (Balme and Hennelly)
Lele 1964* (figure 5.16)
Plicatipollenites trigonalis Lele 1964* (figure 6.5)
Plicatipollenites malabarensis (Potonié and Sah) Foster
1975 (figure 5.11)
Potonieisporites barrelis Tiwari 1965* (figure 6.1)
Potonieisporites densus Maheshwari 1967* (figure 5.5)
Potonieisporites magnus Lele and Karim 1971 (figure
6.2)
Potonieisporites neglectus Potonié and Lele 1961* (figure
5.9)
Potonieisporites sp. cf. P. jayantiensis Lele and Karim
1971* (figure 5.7)
Figure 5. 1, 4, Peppersites sp. cf. P. ellipticus Ravn 1979; 1, BA Pal 5815-2: A42/2; 4, BA Pal 5816-1: T22/2; 2, Limitisporites rectus Leschik
1956 BA Pal 5816-2: Y56/1; 3, Cannanoropolis janakii Potonié and Sah 1960 BA Pal 5815-1: P21/3; 5, Potonieisporites densus
Maheshwari 1967 BA Pal 5811-2: R45/4; 6, Divarisaccus stringoplicatus Ottone 1991 BA Pal 5811-6: Z57/1; 7, Potonieisporites sp. cf. P.
jayantiensis Lele and Karim 1971 BA Pal 5811-4: Q48/0; 8, Cannanoropolis densus (Lele) Bose and Maheshwari 1968 BA Pal 5816-2:
Q49/3; 9, Potonieisporites neglectus Potonié and Lele 1961 BA Pal 5815-1: J32/4; 10, Crucisaccites monoletus Maithy 1965 BA Pal 5813-
1: U21/2; 11, Plicatisporites malabarensis (Potonié and Sah) Foster 1975 BA Pal 5813-2: Z25/3; 12, Caheniasaccites densus Lele and
Karin emend. Gutiérrez 1993 BA Pal 5813-2: N39/4; 13, Costatascyclus crenatus Felix and Burbridge emend. Urban 1971 BA Pal 5813-1:
Y18/4; 14, Circumplicatipollis plicatus Ottone and Azcuy 1988 BA Pal 5811-5: U51/2; 15, Colpisaccites granulosus Archangelsky and
Gamerro 1979 BA Pal 5815-1: Y34/4; 16, Plicatipollenites gondwanensis (Balme and Hennelly) Lele 1964 BA Pal 5811-2: D62/3. Scale bar
/ escala gráfica: 20 μm.
Algae
Tetraporina punctata (Tiwari and Navale) Kar and
Bose 1976 (figure 6.3)
Botryococcus sp. (figure 6.6)
Navifusa sp. (figure 6.7)
Figure 6. 1, Potonieisporites barrelis Tiwari 1965 BA Pal 5815-1: R24/4; 2, Potonieisporites magnus Lele and Karim 1971 BA Pal 5816-
1: X42/3; 3, Tetraporina punctata (Tiwari and Navale) Kar and Bose 1976 BA Pal 5811-5: U60/1; 4, Cannanoropolis mehtae (Lele) Bose
and Maheshwari 1968 BA Pal 5813-1: C33/1; 5 , Plicatipollenites trigonalis Lele 1964 BA Pal 5812-1: R57/4; 6, Botryococccus sp. BA Pal
5815-1: M37/4; 7, Navifusa sp. BA Pal 5816-1: U54/3. Scale bar / escala gráfica: 20 μm.
Anteturma PROXIMEGERMINANTES Potonié 1970
Turma TRILETES Reinsch emend. Dettmann 1963
Suprasubturma ACAVATITRILETES Dettmann 1963
Subturma AZONOTRILETES Luber emend. Dettmann 1963
Infraturma LAEVIGATI Bennie and Kidston emend. Potonié 1956
Infraturma RETUSOTRILETI Streel 1964Genus Retusotriletes Naumova emend.
Streel 1964
Type species. Retusotriletes simplex Naumova 1953.
Retusotriletes sp. 1
Figure 3.7
Description. Spore radial, trilete. Amb subcircular.
Laesurae straight, with raised narrow lips, length
three-quarters of spore radius, with imperfect curvaturae.
Exine laevigate, about 0.8 μm thick.
Dimensions. Equatorial diameter: 40 μm (1 specimen).
Comparisons. Retusotriletes anfractus Menéndez and
Azcuy 1969 differs by its sinuous laesurae. Retusotriletes
tenuis Menéndez 1965 has granular exine. Retusotriletes simplex Naumova 1953 is smaller (equatorial
diameter, 30-35 μm). Retusotriletes indignus Azcuy
1975a differs by its thicker exine.
Retusotriletes sp. 2
Figure 3.4
Description. Spore radial, trilete. Amb circular. Laesurae
straight, with raised narrow lips, length threequarters
of spore radius, with perfect curvaturae.
Exine laevigate, about 0.8 μm thick.
Dimensions. Equatorial diameter: 42 μm (1 specimen).
Comparisons. Retusotriletes minutus Butterworth and
Mahdi 1982 differs by its thicker exine. Retusotriletes sp. 1 possesses imperfect curvaturae.
Infraturma APICULATI Bennie and Kidston emend. Potonié 1956
Subinfraturma GRANULATI Dybová and Jachowicz 1957Genus Anapiculatisporites Potonié and Kremp 1954
Type species. Anapiculatisporites isselburgensis Potonié and Kremp 1954.
Anapiculatisporites sp. cf. A. concinnus Playford 1962
Figure 3.10
Remarks. Following such authors as Dueñas and Césari (2005), it is preferred to retain this species in the genus Anapiculatisporites, rejecting Ravn's (1991) proposal. Anapiculatisporites argentinensis Azcuy 1975a can be considered conspecific with A. concinnus, as suggested by such authors as Jones and Truswell (1992) and Stephenson (2004).
Genus Lophotriletes Naumova emend. Potonié and Kremp 1954
Type species. Lophotriletes gibbosus (Ibrahim) Potonié and Kremp 1954.
Lophotriletes intermedius Azcuy 1975a
Figure 3.15
Remarks. The exine of the studied specimens is slightly thinner than was reported for this species by Azcuy (1975a).
Genus Apiculiretusispora Streel 1964
Type species. Apiculiretusispora brandtii Streel 1964.
Apiculiretusispora sp. cf. A. ralla (Menéndez and Azcuy) Menéndez and Azcuy 1971
Figure 3.9
Description. Spore radial, trilete. Amb subcircular.
Laesurae straight, simple, extending almost to equator,
ending in imperfect curvaturae. Proximal face
laevigate. Distal face sculptured irregularly and
sparsely with grana and coni (0.5-1.5 μm in basal
width and height). Exine 1.5 μm thick.
Dimensions. Equatorial diameter: 42 μm (1 specimen).
Comparisons. Since only one poorly preserved specimen
was found, the specific assignment is made reservedly.
Distribution. This species was previously recorded
in Pennsylvanian sequences of Argentina and Brazil
(e.g., Ottone, 1989, 1991; García, 1995; Longhim et al.,
2002; di Pasquo, 2003; di Pasquo et al., 2003).
Infraturma BACULATI Dybová and Jachowicz 1957
Genus Raistrickia Schopf, Wilson and Bentall emend. Potonié and Kremp 1954
Type species. Raistrickia grovensis Schoff, Wilson and Bentall 1944.
Raistrickia sp. cf. R. radiosa Playford and
Helby 1968
Figure 4.8, 12, 17
Description. Spores radial, trilete. Amb subcircular
to circular. Laesurae straight, simple, length threequarters
of spore radius. Exine with densely distributed
sculptural elements (1-2 μm in basal width, 1.5-
3 μm high) having straight sides and blunt, rounded,
or expanded apices, and occasionally fungiform elements.
Exine 1.5-2 μm thick.
Dimensions. Equatorial diameter: 53(63)72 μm (17
specimens).
Comparisons. Raistrickia radiosa Playford and Helby
1968 differs from the studied specimens in having reduced
sculptural elements on the proximal face.
Distribution. This species was recovered from palynoflora
of Australia (Playford and Helby 1968) and
Argentina (Gutiérrez and Césari, 1989; di Pasquo,
2003).
Genus Reticulatisporites Ibrahim emend. Potonié and Kremp 1954
Type species. Reticulatisporites reticulatus (Ibrahim) Ibrahim 1933.
Reticulatisporites magnidictyus Playford and Helby 1968
Figure 4.6
1991 Reticulatisporites riverosii Ottone, pp. 126-127, plate 1, figure 10, plate 2, figure 10.
Remarks. Examination of the original specimens used
by Ottone (1991) for erecting Reticulatisporites riverosii indicates that they possess all the diagnostic characteristics
of Reticulatisporites magnidictyus. Dino and
Playford (2002a) noted that this species tends to have
larger lumina in proximal face. The Argentinean
spores have a considerable variation, with specimens
showing similar size of lumina in both faces, and others
having larger lumina in the proximal face.
Accordingly, although Ottone did not recognize differences
between proximal and distal in his specimens,
Reticulatisporites riverosii is considered conspecific.
Reticulatisporites magnidictyus, originally described
from the Australian Italia Road Formation by
Playford and Helby (1968), is considered an important
biostratigraphic marker, since its oldest records
are late Visean. However, the species has been recorded
in younger sediments (late Serpukhovian-Pennsylvanian)
by Limarino and Gutiérrez (1990), Ottone
(1991), Gutiérrez and Barreda (2006) and Perez Loinaze
(2007).
Subturma ZONOTRILETES Waltz in Luber and Waltz 1938
Infraturma CINGULICAVATI Smith and Butterworth 1967Genus Cristatisporites Potonié and Kremp emend. Butterworth, Jansonius, Smith and Staplin 1964
Type species. Cristatisporites indignabundus (Loose) Potonié and Kremp 1954.
Cristatisporites sp.
Figure 4.5
Description. Spore radial, trilete, cavate, cingulizonate.
Amb subtriangular. Inner body subtriangular.
Laesurae straight, with narrow lips, extending almost
to the equatorial margin. Proximal face laevigate.
Central body with a narrow cingulum. Central area of
the distal face sculptured with grana and sharp coni
and spinae (3-10 μm high, 1.5-5 μm in basal width),
frequently fused basally to form short cristae, often
surmounted by a small spinus. Sculptural elements
progressively diminished polewards. Translucent
zona about 1/4 of spore radius, outer margin entire to
slightly irregular, sculptured with sparsely distributed
coni and spinae up to 5 μm high.
Dimensions. Equatorial diameter: 70 μm (1 specimen).
Comparisons. In Cristatisporites indignabundus (Loose)
Staplin and Jansonius 1964, the margin of the central
body bears setose spinae up to 2.5 μm high. Cristatisporites
lestai Archangelsky and Gamerro 1979 and
C. longispinosus Menéndez 1971 differ by having a
markedly irregular zona. Moreover, in the former,
the sculpture is dominantly verrucose. Cristatisporites
solaris (Dias-Fabrício) Picarelli and Dias-Fabrício
1990 has a smaller diameter and dentate margin
zona. In Cristatisporites rollerii Ottone 1989, the zonal
width is approximately 1/2-1/3 of the spore radius,
and the sculptural elements are differently distributed
on the body.
Genus Indotriradites (Tiwari) Foster 1979
Type species. Indotriradites korbaensis Tiwari 1964.
Indotriradites volkheimeri (Azcuy) Perez Loinaze 2008
Figure 4.9
Dimensions. Equatorial diameter: 60, 68 μm (2 specimens).
Remarks. Krauselisporites volkheimeri Azcuy 1975b was recently referred to Indotriradites by Perez Loinaze
(2008) in accordance with the emendation of
Krauselisporites by Scheuring (1974).
Suprasubturma PSEUDOSACCITITRILETES Richardson 1965
Infraturma MONOPSEUDOSACCITI Smith and Butterworth 1967Genus Grandispora Hoffmeister, Staplin and Malloy emend. Neves and Owens 1966
Type species. Grandispora spinosa Hoffmeister, Staplin and Malloy 1955.
Grandispora sp.
Figure 4.10
Description. Spores radial, trilete, cavate. Amb triangular
to rounded subtriangular. Laesurae straight, with
narrow lips reaching almost to the equatorial margin.
Intexine distinct, outline conformable with amb, about
2/3 of spore radius. Proximal surface laevigate. Distal
surface sculptured with coni and grana, rounded in lateral
view, and occasionally with irregularly distributed
biform elements (2-6 μm high, 2-5 μm in basal width).
Equatorial region with close-spaced coni and bacula (2-
5 μm high and 5-10 μm in basal width).
Dimensions. Equatorial diameter: 56, 91 μm (2 specimens).
Comparison. The above specimens differ from previously
described species of Grandispora in that the distal
sculptural elements are smaller than those in the
equatorial region. Jayantisporites Lele and Makada
1972 possesses a pseudozona, formed by basally
fused sculptural elements. However, this genus differs
from the Malanzán specimens in having basally
fused, biform sculptural elements, thus forming
cristate ridges on the distal face. Grandispora uncata (Hacquebard) Gupta 1969 is similar to these specimens,
but both its proximal and distal faces are
sculptured, and more densely so in the equatorial region.
Spinozonotriletes sp. 1 Azcuy 1975b differs in
having larger sculptural elements (8-23 μm).
Genus Endosporites Wilson and Coe emend. Bharadwaj 1965
Type species. Endosporites ornatus Wilson and Coe 1940 (designated by Schopf et al., 1944).
Endosporites sp.
Figure 4.11
Description. Spores radial, trilete, cavate, amb subcircular.
Laesurae straight, simple, reaching almost
to the equatorial margin. Intexine laevigate, outline
conformable with amb. Exoexine microgranular,
thicker than intexine.
Dimensions. Equatorial diameter: 42-51 μm, intexine
diameter: 35-39 μm (6 specimens).
Comparisons. Endosporites parvus Menéndez 1965
possesses microgranular intexine and is smaller
(equatorial diameter, 32 μm; intexine diameter, 15.5-
18.5 μm). Endosporites translucidus Menéndez 1965
and Endosporites sp. of Gutiérrez and Limarino (2001)
differ in having granular intexine. Endosporites sp. of
Césari and Vázquez Nístico (1988) has laevigate and
strongly folded exoexine. Endosporites sp. A of Azcuy
(1975b) features a thicker exoexine and microgranular
intexine. Endosporites rhytidosaccus Menéndez and
Azcuy 1973 and Endosporites minutus Hoffmeister,
Staplin and Malloy 1955 have microgranular and
folded intexine.
Anteturma VARIEGERMINANTES Potonié 1970
Turma SACCITES Erdtman 1947
Subturma MONOSACCITES Chitaley emend. Potonié and Kremp 1954
Infraturma VESICULOMONORADITI Pant 1954Genus Costatascyclus Felix and Burbridge emend. Urban 1971
Type species. Costatascyclus crenatus Felix and Burbridge emend. Urban 1971.
Costatascyclus crenatus Felix and Burbridge emend. Urban 1971
Figure 5.13
Description. Pollen grains bilateral, monosaccate.
Amb transversely oval with crenulate margin. Corpus
outline subcircular to oval in polar view, denser
than saccus. Laesurae rarely perceptible. Saccus attached
to corpus distally and proximally; finely endoreticulate,
with coarse radial folds.
Dimensions. Overall breadth: 86(121)152 μm, overall
length: 60(79)100 μm, corpus length: 48(62)72 μm,
corpus breadth: 40(58)75 μm (32 specimens).
Comparisons. Potonieisporites balmeii (Hart) Segroves
1969 differ in having a pair of distinctive folds parallels
the corpus margin.
Previous records. Pennsylvanian: Brazil, Paraná Basin,
Itararé Subgroup (Souza, 2000; Souza et al., 2003);
Parnaíba Basin, Piauí Formation (Dino and Playford,
2002b); Amazonas Basin, Tapajós Group (Playford
and Dino, 2000); U.S.A. (Felix and Burbridge, 1967;
Urban, 1971; Ravn, 1979, 1986). Lower Permian: Uruguay,
San Gregorio Formation (Gutiérrez et al., 2006).
Subinfraturma DISACCITES Cookson 1947
Infraturma DISACCITRILETES Leschik 1956
Genus Colpisaccites Archangelsky and Gamerro 1979
Type species. Colpisaccites granulatus Archangelsky and Gamerro 1979.
Colpisaccites granulosus Archangelsky and Gamerro 1979
Figure 5.15
Comparisons. Compared to the material described
originally by Archangelsky and Gamerro (1979), the
present specimens are larger, having the following
dimensions: overall breadth, 125(145)170 μm; overall
length, 103(123)130 μm; corpus length, 105(117)150
μm; corpus breadth, 81(103)111 μm. However, the
Malanzán specimens are similar in size to those described
by Gutiérrez (1993), who noted that size differences
can be interpreted as infraspecific variability.
Among the three morphotypes of Colpisaccites
granulosus identified by Gutiérrez (1993), two (A and
B) were recognized during the present study. These
morphotypes differ in the degree of separation of the
exine (more marked in A).
Previous records. Pennsylvanian: Argentina, Paganzo
Basin, Agua Colorada Formation (Limarino and
Gutiérrez, 1990; Gutiérrez, 1993). Lower Permian:
Argentina, San Rafael Basin, Yacimiento Los Reyunos
Formation (Césari et al., 1996), Chacoparaná
Basin (Archangelsky and Gamerro, 1979); Uruguay,
San Gregorio Formation (Gutiérrez et al., 2006).
Genus Peppersites Ravn 1979
Type species. Peppersites ellipticus Ravn 1979.
Peppersites sp. cf. P. ellipticus Ravn 1979
Figure 5.1, 4
Description. Pollen grains bilateral, monosaccate.
Amb transversely oval to elliptical. Corpus outline
(in polar view) closely conformable, dark, without
folds. Laesurae rarely discernible. Saccus finely
infrareticulate, approximately 1/10 of central
body.
Dimensions. Overall breadth: 135-166 μm, overall
length: 102-148 μm, corpus length: 115-154 μm, corpus
breadth: 84-125 μm, saccus breadth: 6-12 μm (5
specimens).
Comparisons. This monotypic pollen genus is characterized
chiefly by its large corpus. The specimens
studied differ from Peppersites ellipticus in having a
corpus slightly narrower than the overall breadth.
Previous records. Pennsylvanian: Brazil, Amazonas
Basin, Tapajós Group (Playford and Dino, 2000);
U.S.A. (Ravn, 1979, 1986; Ravn and Fitzgerald,
1982).
Characteristics of the palynological assemblages
The palynomorphs recovered from the Malanzán
Formation are generally well-preserved and moderately
diverse. Relative abundance of pollen grains is
comparable between Section 3 (13-29 %) and Section
4 (12-24 %). The marine elements and algae species
are very scarce, representing less than 1% of the total
assemblage, and were recognized only in Section 4.
In his study of six samples of the Malanzán Formation
in the Sierra de los Llanos area, Azcuy (1975a,
b) recognized 19 spore species and two pollen species.
Based on those identifications, Azcuy suggested
an early Namurian age for the formation. More recently,
Gutiérrez and Limarino (2001) studied black
shale samples of this unit (from Section 3 of the present
study), providing a more complete palynological
inventory, with the identification of 41 species of
spores and 11 of pollen grains (the latter unillustrated).
These authors postulated a late Namurian-early
Westphalian for the association and described marine
elements for the first time from the unit. The palynological
assemblages of the Malanzán Formation
were referred by Césari and Gutiérrez (2001) to the Raistrickia densa-Convolutispora muriornata Biozone
(Serpukhovian-Stephanian).
The studied assemblages comprise a total of 37
spore species (all trilete) distributed among 23 genera.
Eleven genera and 19 species of pollen grains are
identifiable; of these, bilateral or radial monosaccate
pollen grains (17 species) predominate, whereas only
two species are bisaccate. As in Azcuy´s (1975a, b)
pioneering work on the formation, taeniate pollen
grains were not recognized in the current study.
However, Gutiérrez and Limarino (2001) have identified
a specimen of Striatites sp. in a sample collected
from the here-designated Section 3.
Thirty-five species are identified herein for the
first time from the Malanzán Formation; viz., Apiculatisporis variornatus, Apiculiretusispora alonsoi,
Caheniasaccites densus, Colpisaccites granulosus, Costatascyclus
crenatus, Cristatisporites inconstans, C. rollerii,
Cyclogranisporites firmus, C. plicatus, C. rinconadensis,
Dibolisporites disfacies, Divarisaccus stringoplicatus,
Grossusporites microgranulatus, Leiotriletes
tiwarii, Limitisporites rectus, Microreticulatisporites
punctatus, Plicatipollenites gondwanensis, P. trigonalis,
Potonieisporites barrelis, P. densus, P. neglectus,
Punctatisporites gretensis, P. irrasus, P. priscus,
Reticulatisporites magnidictyus, Verrucosisporites andersonii,
Granulatisporites sp. cf. G. austroamericanus, Peppersites sp. cf. P. ellipticus, Potonieisporites sp. cf.
P. jayantiensis, Raistrickia sp. cf. R. radiosa, Cristatisporites sp., Endosporites sp. Grandispora sp., Retusotriletes sp. 1, and Retusotriletes sp. 2. This increases
appreciably the number of reported Malanzán
species.
Organic-walled microfossils identified as Tetraporina
punctata are referred to the charophycean algal
Family Zygnemataceae, which is interpreted by several
authors (e.g., Colbath and Grenfell, 1995; Grenfell,
1995; Geel and Grenfell, 1996) as a freshwater inhabitant.
On the other hand, the presence of Navifusa is a likely indicator of marine environments, and
Botryococcus occurs in habitats of variable salinity as
well as in freshwater or brackish environments (Guy-
Ohlson, 1992; Brenner and Foster, 1994; Batten and
Grenfell, 1996). The presence of these palynomorphs
reinforces the interpretation of a transitional, fjordtype
depositional environment, as previously suggested
(Buatois and Mangano, 1995; Gutiérrez and
Limarino, 2001; Limarino et al. 2002).
Comparison and age of the association
Several spore-pollen species typically found in
Mid Carboniferous sequences of Argentina and
Brazil occur in the studied samples. The most important
are Apiculatisporis variornatus, Apiculatasporites
parviapiculatus, Apiculiretusispora alonsoi, Convolutispora
muriornata, Raistrickia densa, Reticulatisporites passaspectus,
Plicatipollenites malabarensis, P. trigonalis, P.
gondwanensis, Potoniesporites barrelis, P. brasiliensis, P.
densus, P. magnus, P. neglectus, Circumplicatipollis plicatus,
and Crucisaccites monoletus. The presence of
pollen grains in the Malanzán Formation supports an
age not older than Serpukhovian, because the first
worldwide records of pollen grains occur in this uppermost
Mississippian stage (Brugman et al., 1985;
Loboziak and Clayton, 1988; Clayton et al., 1990; Zhu,
1993; Clayton, 1995).
Based on its palynological content, the Malanzán
Formation is referred to the Raistrickia densa-Convolutispora
muriornata Biozone (DM), proposed for western
Argentina by Césari and Gutiérrez (2001). Furthermore,
the scarcity of taeniate pollen grains would
suggest a restriction to the Sub-Biozone A of the DM
Biozone, dated as Serpukhovian (Césari and Gutiérrez,
2001; Gutiérrez et al., 2003).
The studied palynoflora shows strong similarities
to the assemblages described from the Agua
Colorada, Guandacol and El Trampeadero formations;
however, only broad similarities were found
with those reported from the Lagares and Jejenes
formations (figure 7). The palynoflora of the lower
member of the Agua Colorada Formation at the Las
Gredas locality (Limarino and Gutiérrez, 1990)
shares with the Malanzán assemblages stratigraphically
important spore species such as Convolutispora
muriornata, Reticulatisporites asperidictyus
and R. magnidictyus, together with an important
complement of pollen species (figure 7, 1A).
Assemblages from the upper part of the Agua
Colorada Formation differ in containing taeniate
pollen grains (Gutiérrez, 1993). The Guandacol Formation's
assemblages (Césari and Vázquez-Nístico,
1988; Ottone and Azcuy, 1990; Ottone, 1991; Césari
and Limarino, 2002; Perez Loinaze, 2007) share
with those of the Malanzán Formation a substantial
number of spore species. These are Anapiculatisporites
concinnus, Apiculatisporis variornatus,
Apiculiretusispora alonsoi, Convolutispora muriornata,
Cyclogranisporites rinconadensis, C. firmus, Dibolisporites
disfacies, Raistrickia densa, Reticulatisporites asperidictyus,
and R. magnidictyus (figure 7). The palynoflora
described by Gutiérrez and Barreda (2006)
from the El Trampeadero Formation (La Cébila
Creek, La Rioja Province) shares several spore
species with the Malanzán assemblages, including Convolutispora muriornata, Raistrickia densa, Dibolisporites
disfacies, Reticulatisporites asperidictyus, R.
magnidictyus, and many pollen species (figure 7).
Only a few species are shared between the
Malanzán and the lower part of the Lagares
Formation: Apiculatasporites caperatus, Apiculiretusispora
ralla, Cannanoropollis densus, and C. mehtae (Menéndez and Azcuy, 1969, 1971, 1973; Azcuy and
Gutiérrez, 1983; see figure 7). On the other hand, Convolutispora muriornata and Anapiculatisporites
concinnus (=Anapiculatisporites argentinensis) are the
only two species in common between the former
unit and the middle and upper sections of the
Lagares Formation (Césari and Gutiérrez, 1985; Gutiérrez
and Césari, 1989). In addition, as shown in
figure 7 and by reference to González-Amicón
(1973), Gutiérrez and Césari (1987), and Césari and
Bercowski (1997), six species are shared with the
palynofloras reported from the lower sections of
the Jejenes Formation (e.g., Apiculatisporis variornatus,
Grossusporites microgranulatus and Reticulatisporites
asperidictyus); and a further seven species
from the upper section (Apiculatasporites caperatus,
Apiculiretusispora ralla, Cannanoropollis densus,
Convolutispora muriornata, Microreticulatisporites
punctatus, Plicatipollenites malabarensis and Punctatisporites
gretensis).
Figure 7. Stratigraphic distribution of the identified species/ distribución
estratigráfica de las especies identificadas: 1= Agua Colorada
Formation / Formación Agua Colorada, A- Lower member / miembro
inferior (Azcuy et al., 1982; Limarino et al., 1984; Limarino and
Gutiérrez, 1990; Vergel and Luna, 1992, Gutiérrez, 1993; Vergel et al., 1993), B- Middle and upper sections / sección media y superior (Menéndez, 1965; Menéndez and González-Amicón, 1979;
Gutiérrez, 1993); 2= Jejenes Formation / Formación Jejenes, ALower
section / sección inferior (Césari and Bercowski, 1997), BMiddle
and upper sections / sección media y superior (González-
Amicón, 1973; Gutiérrez and Césari, 1987); 3= Lagares Formation
/ Formación Lagares, A- Lower section / sección Inferior (Menéndez
and Azcuy, 1969, 1971, 1973; Azcuy and Gutiérrez, 1983), BMiddle
and upper sections / sección media y superior (Césari and
Gutiérrez, 1985, Morelli et al., 1984; Gutiérrez and Césari, 1989); 4=
Guandacol Formation / Formación Guandacol (Césari and Vázquez-
Nístico, 1988; Ottone and Azcuy, 1990; Ottone, 1991; Césari and
Limarino, 2002; Perez Loinaze, 2007); 5= Trampeadero Formation
/ Formación Trampeadero (Barreda, 1986; Moreno, 1993; Bossi, 1999;
Gutiérrez and Barreda, 2006).
The palynological assemblages recovered from
the Guandacol and Trampeadero formations and
those from the lower part of the Jejenes, Lagares and
Agua Colorada formations were included in the Subzone
A of the DM Biozone, characterized by the presence
of Convolutispora muriornata and Raistrickia densa and the absence of striate pollen grains (Césari and
Gutiérrez, 2001).
It is noteworthy that Reticulatisporites magnidictyus was widely reported in the late Viséan, but in Argentina
occurs in younger palynofloras (Serpukhovian)
belonging to Subzone A of the DM Biozone (e.g., Limarino
and Gutiérrez, 1990; Ottone, 1991; Gutiérrez
and Barreda, 2006; Perez Loinaze, 2007). Other species
that may also have stratigraphic significance are Reticulatisporites asperidictyus and Dibolisporites disfacies recently identified in assemblages referred to the
DM Biozone (e.g., Gutiérrez and Limarino, 2001; Gutiérrez
and Barreda, 2006; Perez Loinaze, 2007).
Souza (2006) proposed two new interval biozones
for Pennsylvanian deposits of the northeastern
Paraná Basin. The older, the Ahrensisporites cristatus Biozone (dated as late Bashkirian to Kasimovian),
shares with the Malanzán palynoflora many species,
such as Anapiculatisporites concinnus (=Anapiculatisporites
argentinensis), Cristatisporites menendezii, Raistrickia
rotunda, R. paganciana, Convolutispora muriornata,
and some pollen species, but differs in its content of
taeniate pollen grains (Protohaploxypinus).
Jones and Truswell (1992) identified five Oppelzones
in the Joe Joe Group of the Galilee Basin, Australia.
The oldest, the Verrucosisporites basiliscutis Oppel-
zone dated as early Namurian, shares with the
Malanzán palynoflora only two species (i.e., Cannanoropollis janakii and Cyclogranisporites firmus).
The slightly younger Brevitriletes leptoacaina Oppelzone
(Namurian-?early Westphalian) has only one
species (D. disfacies) in common with the Malanzán
assemblages. Both Australian Oppel-zones are characterized,
as the Malanzán palynoflora, by the complete
absence of taeniate pollen grains.
The base of the Australian Spelaeotriletes ybertii Assemblage (Kemp et al., 1977) and the Diatomozonotriletes
birkheadensis Assemblage (Powis, 1984)
are marked by the first appearance of monosaccate
pollen grains. However, occurrence of taeniate
pollen grains in these Australian Spelaeotriletes ybertii and Diatomozonotriletes birkiadensis assemblages
preclude their correlation with the Malanzán palynoflora.
Conclusions
A total of 37 spore and 19 pollen species have been
identified in the analysed samples of the Malanzán Formation, as exposed at the La Chimenea locality in
the Olta valley and at the Cuestita de la Herradura locality
in the Malanzán valley. Thirty-five species of
these species have not been recovered so far from the
Malanzán Formation. Furthermore, pollen grains are
illustrated for the first time for this unit.
The co-occurrence of the stratigraphically significant
spore species as Convolutispora muriornata and
Raistrickia densa, and a great number of pollen species,
clearly indicate a Serpukhovian age for the Malanzán
Formation, and confirms (apropos of Césari
and Gutiérrez, 2001) its attribution to the Sub-Biozone
A of the Raistrickia densa-Convolutispora muriornata Biozone. The Malanzán palynoflora can be correlated
with those described from the Guandacol and El
Trampeadero formations, and with those from the
lower parts of the Jejenes, Lagares and Agua Colorada
formations.
The scarce presence of Navifusa sp., together with
brackish water algae and the predominance of terrestrial
palynomorphs testified the interpretation of the
Malanzán Formation as deposited in a fjord environment.
Acknowledgements
Thanks are due to S.N. Césari for her critical comments and useful suggestions during all phases of this work; O. Limarino for his invaluable help; and E. Vera for reading the manuscript. Thanks are extended to P.R. Gutiérrez and G. Playford , for their enriching comments and suggestions as referees. Funding for this work was provided by the Agencia Nacional de Promoción Científica y Técnica, PICT 20752 and CONICET PIP 5723.
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Recibido: 11 de noviembre de 2008.
Aceptado: 10 de mayo de 2009.