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.