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Ameghiniana

versión On-line ISSN 1851-8044

Ameghiniana v.44 n.3 Buenos Aires jul./sep. 2007

 

Palynology of the Chigua (Devonian) and Malimán (Carboniferous) formations in the Volcán Range, San Juan Province, Argentina. Part II. Cavate, pseudosaccate and cingulizonate spores

Cecilia R. Amenábar 1, Mercedes Di Pasquo1, Hugo A. Carrizo2 and Carlos L. Azcuy1

1Consejo Nacional de Investigaciones Científicas y Técnicas. Departamento de Geología, Facultad de Ciencias Exactas y Naturales. Ciudad Universitaria, Pabellón 2, 1º Piso, 1428 Buenos Aires, Argentina. amenabar@gl.fcen.uba.ar, medipa@gl.fcen.uba.ar, azcuy@ciudad.com.ar
2Fundación Miguel Lillo, Sección Paleobotánica, Miguel Lillo 251, 4000 San Miguel de Tucumán, Argentina. hugocarrizo5@yahoo.com.ar

Resumen. Palinología de las formaciones Chigua (Devónico) y Malimán (Carbonífero) en la Sierra del Volcán, Provincia de San Juan, Argentina. Parte II. Esporas cavadas, pseudocavadas y cingulizonadas. Se presenta el análisis sistemático de las asociaciones palinológicas obtenidas de sedimentitas de las formaciones Chigua y Malimán, Devónico medio y Mississippiano respectivamente, aflorantes en la sierra del Volcán, cuenca Río Blanco, oeste de Argentina. En esta segunda parte se ilustran 40 especies de esporas cavadas, pseudosacadas y cingulizonadas presentes en ambas formaciones. La especie Geminospora sp. cf. G. tuberculata var. tuberculata McGregor es registrada por primera vez en el Devónico medio de América del Sur. Las siguientes especies también son registradas por primera vez para el Mississippiano de América del Sur, Crassispora invicta Playford, Densosporites asperus Braman y Hills, Densosporites gracilis Smith y Butterworth, Densosporites secundus Playford y Satterthwait, Densosporites sp. cf. D. triangularis Kosanke, Gorgonispora sp. cf. G. crassa (Winslow) Higgs et al., Lophozonotriletes dentatus Hughes and Playford, Spelaeotriletes echinatus (Hacquebard) Utting, Velamisporites perinatus (Hughes y Playford) Playford. Se ilustran y en algunos casos se describen algunas especies (escolecodontes, paleomicroplancton y esporas) posiblemente retrabajadas presentes en la Formación Malimán.

Abstract. Palynological analysis of the Chigua and Malimán formations of Middle Devonian and Mississippian age respectively, is presented. They crop out in the Del Volcán Range at the Río Blanco Basin in western Argentina. In this second part, 40 indigenous species of cavate, pseudosaccate and cingulizonate spores from both units are illustrated. The species Geminospora sp. cf. G. tuberculata var. tuberculata McGregor is recorded in the Middle Devonian of South America for the first time. The following species are also recorded in the Mississippian of South America for the first time as Crassispora invicta Playford, Densosporites asperus Braman and Hills, Densosporites gracilis Smith and Butterworth, Densosporites secundus Playford and Satterthwait, Densosporites sp. cf. D. triangularis Kosanke, Gorgonispora sp. cf. G. crassa (Winslow) Higgs et al., Lophozonotriletes dentatus Hughes and Playford, Spelaeotriletes echinatus (Hacquebard) Utting, Velamisporites perinatus (Hughes and Playford) Playford. Some possibly reworked species (scolecodonts, paleomicroplankton and spores) from the Malimán Formation are illustrated and some of them are also described.

Key words. Palynology; Chigua Formation; Malimán Formation; Middle Devonian; Mississippian; San Juan Province; Argentina.
Palabras clave. Palinología; Formación Chigua; Formación Malimán; Devónico medio; Mississippiano; Provincia de San Juan; Argentina.

Introduction

In this paper a palynological analysis of Devonian and Carboniferous sediments of the Río Blanco Basin in the western Argentina is presented. The Río Blanco Basin is one of the Late Palaeozoic basins with the thickest Carboniferous deposits of Argentina, which overlies Devonian levels in angular unconformity. The palynomorphs studied herein come from samples obtained from the Chigua and Malimán formations that crop out at the La Cortadera creek in the western flank of the Volcán Range, Precordillera of San Juan. This analysis is presented in two parts. The paleomicroplankton from the Chigua Formation and acavate smooth and ornamented trilete spores from both units along with the geological setting, location of fertile samples in the outcrop scheme and material and methods were already presented in the first part (Amenábar et al., 2006). This second part deals mainly with the systematic and the stratigraphic distribution of selected cavate, pseudosaccate and cingulizonate spores of both units. In addition, selected paleomicroplankton species either indigenous, recycled or of doubtful origin, obtained from the Malimán Formation, are illustrated in figure 1. Comparisons with other similar microfloras and the stratigraphic importance of both Devonian and Mississippian microfloras are subject of other contribution (in preparation).

Systematic palaeontology

Forty species of cavate, pseudosaccate and cingulizonate spores are presented in a parataxonomic scheme introduced by Potonié and Kremp (1954) modified by Potonié (1970) and others. The latest Spanish edition of the ICBN Code (Kiesling, 2002) is here adopted for the treatment of form taxa. Morphological terminology is mainly in accordance with the last glossary provided by Punt et al. (1994). Only species left in open nomenclature are described in detail whereas the main diagnostic features and/or remarks are given for previously known species that deserve a taxonomic and/or morphological discussion. Synonymy lists that have not been published yet are provided; otherwise they will be referred to where they were cited. The range and the worldwide geographic distribution of the species recognized in the Chigua and Malimán formations as shown in tables 1 and 2 are taken from selected systematic papers and others from illustrated lists of palynomorphs. Only the South American records are included in the text following the format presented in Part I. Selected palynologic works with illustrations are considered under the title Illustrated, while abstracts or non-illustrated works are mentioned as Listed. Illustrations of specimens are given in figures 4 to 7.
Finally, the figure 1 shows some species of scolecodonts, paleomicroplankton and spores found in the Malimán Formation. The specimens of the colonial algae Botryococcus braunii Kützing 1849 (figure 1.16) are here considered as indigenous. Other species, such as Maranhites brasiliensis Brito emend. Burjack and Oliveira 1989 (figure 1.11), Maranhites sp. cf. M. brasiliensis Brito emend. Burjack and Oliveira 1989 (figure 1.10), Hystricosporites sp. cf. H. gravis Balme 1988 (figure 1.13), Hystricosporites sp. cf. H. porrectus (Balme and Hassell) Allen 1965 (figure 1.19), Petaloferidium ancorum (Wicander and Loeblich) Sarjeant and Vavrdová 1997 (figure 1.20), Estiastra improcera Loeblich 1970 (figure 1.21), Synsphaeridium sp. (figure 1.5), Corystisporites sp. (figure 1.14), are here interpreted as undoubted reworked forms. They are added to other reworked species registered in the unit and recently discussed and illustrated by Amenábar (2006). The remainder forms showed in the figure 7, Tasmanites sp. (figure 1.6), Veryhachium sp. cf. V. trispinosum (Eisenack) Deunff 1954 (figure 1.9), Dictyotidium torosum Playford in Playford and Dring 1981 (figure 1.12) and other species left in open nomenclature such as, scolecodonts (figures 1.1-1.4), and the paleomicroplankton Dictyotidium sp. (figures 1.7, 1.8), Cymatiosphaera sp. (figures 1.17, 1.18) and Micrhystridium sp. (figure 1.15), could be either indigenous (Carboniferous) or reworked forms. The latter species are described after the spores. A discussion about their stratigraphical value will be treated in other contribution.
The samples studied are housed in the repository of the Palynology Laboratory, Department of Geology, University of Buenos Aires. The slide number is denoted by the prefix BAFC-Pl followed by the England Finder (EF) reference.


Figure 1.
Scolecodonts, palaeomicroplankton and spores from the Malimán Formation. Scale bar: 5, 12, 15, 20 = 10 µm (x 1000); 6-8, 17, 18, 21 = 15 µm (x 750); 1-4, 9-11, 13, 14, 16, 19 = 20 µm (x 500). Coordinates after EF (England Finder) graticule / escolecodontes, paleomicroplancton y esporas de la Formación Malimán. Escala gráfica: 5, 12, 15, 20 = 10 µm (x 1000); 6-8, 17, 18, 21 = 15 µm (x 750); 1-4, 9-11, 13, 14, 16, 19 = 20 µm (x 500). Las coordenadas corresponden a la reglilla EF (England Finder). 1-4, Scolecodonts. 1, BAFC-Pl 1503 (2): L31; 2, BAFC-Pl 1501 (1): G28/4; 3, BAFC-Pl 1501 (1): V56/2; 4, BAFC-Pl 1501 (1): Y47/0-4. 5, Synsphaeridium sp. Playford and Dring. BAFC-Pl 1504 (1): X55/1. 6, Tasmanites sp. BAFC-Pl 1506 (4): V44. 7-8, Dictyotidium sp. 7, BAFC-Pl 1506 (2): A22; 8, BAFC-Pl 1655 (2): G21/4. 9, Veryhachium sp. cf. V. trispinosum (Eisenack) Deunff 1954. BAFC-Pl 1506 (4): X55. 10, Maranhites sp. cf. M. brasiliensis Brito emend. Burjack and Oliveira. BAFC-Pl 1504 (2): Y42. 11, Maranhites brasiliensis Brito emend. Burjack and Oliveira. BAFC-Pl 1506 (2): R23/1-2. 12, Dictyotidium torosum Playford in Playford and Dring. BAFC-Pl 1504 (4): X60/2. 13, Hysticosporites sp. cf. H. gravis Balme. BAFC- Pl 1508 (4): P29/4. 14, Corystisporites sp. BAFC-Pl 1508 (4): A22/2. 15, Micrhystridium sp. BAFC-Pl 1504 (2): X55/1. 16, Botryococcus braunii Kützing. BAFC-Pl 1504 (3): S40/1. 17-18, Cymatiosphaera sp. Q, BAFC-Pl 1506 (2): V57; R, BAFC-Pl 1506 (4): Z39/4. 19, Hystricosporites sp. cf. H. porrectus (Balme and Hassell) Allen. BAFC-Pl 1508 (4): V23/4. 20, Petaloferidium ancorum (Wicander and Loeblich) Sarjeant and Vavrdová. BAFC-Pl 1501 (1): K36/1. 21, Estiastra improcera Loeblich. BAFC-Pl 1506 (1): C32/3.

Table 1. Worldwide range and geographical distribution of selected species recognized in the Chigua Formation. Eurasia (E), South America (SA), North America (NA), Africa (Af), Australia (Au) / distribución estratigráfica y geográfica mundial de las especies seleccionadas reconocidas en la Formación Chigua. Eurasia (E), Sud América (SA), Norte América (NA), África (Af), Australia (Au). References: for South American records see text and those for the rest of the world are mainly based on the following selected literature / Referencias: los registros de América del Sur figuran en el texto y aquellos del resto del mundo se basan principalmente en la siguiente literatura seleccionada: Allen, 1965; Richardson, 1965; Menéndez and Pöthe de Baldis, 1967; Owens, 1971; McGregor, 1973, 1977, 1979, 1984, 1996; McGregor and Canfield, 1976, 1982; Moreaut-Benoit, 1979, 1980; Loboziak and Streel, 1981; Paris et al., 1985; Richardson and McGregor, 1986; Turnau, 1986, 1996; Burjack et al., 1987; Balme, 1988; Loboziak et al., 1988, 1991, 1992, 1997a, 1997b; Pérez Leyton, 1990, 1991; Braman and Hills, 1992; Avchimovitch et al., 1993; Rodrigues et al., 1995; Limachi et al., 1996; Ottone, 1996; Ottone and Rossello, 1996; Vavrdová et al., 1996; Dino, 1999; Turnau and Racki, 1999; Loboziak and Melo, 2000, 2002; Melo and Loboziak, 2003; Rubinstein et al., 2005; di Pasquo, 2007a.

Table 2. Worldwide range and geographical distribution of selected species recognized in the Malimán Formation. Eurasia (E), South America (SA), North America (NA), Africa (Af), Australia (Au) / distribución estratigráfica y geográfica mundial de las especies seleccionadas reconocidas en la Formación Malimán. Eurasia (E), América del Sur (AS), América del Norte (AN), África (Af), Australia (Au). References: for South American records see text and those for the rest of the world are mainly based on the following selected literature / Referencias: los registros de América del Sur figuran en el texto y aquellos del resto del mundo se basan principalmente en la siguiente literatura seleccionada: Huges and Playford, 1961; Smith and Butterworth, 1967; Braman and Hills, 1977, 1992; Clayton et al., 1978; Turnau, 1978; McGregor, 1979; Paris et al., 1985; Richardson and McGregor, 1986; Azcuy and Ottone, 1987; Avchimovitch et al., 1988, 1993; Higgs et al., 1988; Playford, 1978, 1985, 1991; Utting, 1987; Playford and Satterthwait, 1988; Pérez Leyton, 1990, 1991; Loboziak et al., 1991, 1992, 1997a, 1997b, 1998, 2000; Ravn, 1991; Vavrdová et al., 1993, 1996; Césari and Limarino, 1995; Rubinstein et al., 1996; Díaz Martínez et al., 1999; Melo et al., 1999; Niemeyer and Rubinstein, 2000; Jäger, 2002; Loboziak and Melo, 2002; Melo and Loboziak, 2000, 2003; Iannuzzi and Pfefferkorn, 2002; Utting and Giles, 2004; Fasolo et al., 2006; Azcuy and di Pasquo, 2005, 2006; McLean et al., 2005; di Pasquo, 2007b; Souza, 2006.

Spores

Anteturma PROXIMEGERMINANTES Potonié 1970
Turma TRILETES Reinsch emend. Dettmann 1963
Suprasubturma ACAVATITRILETES Dettmann 1963
Infraturma MURORNATI Potonié and Kremp 1954

Genus Acinosporites Richardson 1965

Type species. Acinosporites acanthomammillatus Richardson 1965.

Acinosporites acanthomammillatus Richardson 1965
Figures 2.1-2.2

Occurrence. (Illustrated): Middle Devonian, Bolivia (McGregor, 1984; di Pasquo, 2007a), Brazil (Burjack et al., 1987). Late Devonian, Brazil (Loboziak et al., 1988, 1992). (Listed): Middle-Late Devonian, Brazil (Dino, 1999), Bolivia (Pérez Leyton, 1991; Limachi et al., 1996).

Subturma ZONOTRILETES Waltz emend. Potonié and Kremp 1954
Infraturma CINGULATI Potonié and Klaus emend. Dettmann 1963
Genus Lophozonotriletes Naumova emend. Potonié 1958

Type species. Lophozonotriletes lebedianensis Naumova 1953.


Figure 2.
Cavate, pseudosaccate and cingulizonate spores from the Chigua Formation.
Scale bar: 10, 11 = 20 µm (x 500), the rest = 15 µm (x 750). Coordinates after EF (England Finder) graticule / esporas cavadas, pseudosacadas y cingulizonadas de la Formación Chigua. Escala gráfica: 10, 11 = 20 µm (x 500), el resto = 15 µm (x 750). Las coordenadas corresponden a la reglilla EF (England Finder). 1-2, Acinosporites acanthomammillatus Richardson. BAFC-Pl 1656 (1): R35; 1, proximal face showing a labiate trilete mark / cara proximal mostrando la marca trilete labiada; 2, distal face showing the rugulae pattern / cara distal mostrando el patrón rugulado. 3-4, Cymbosporites catillus Allen. BAFC- Pl 1505 (2): G38/2. 5-6, 9, Geminospora lemurata Balme 1962 emend. Playford. 5, BAFC-Pl 1505 (2): X34/2; 6, BAFC-Pl 1505 (5): D28/2; 9, BAFC-Pl 1505 (2): X35/1. 7, Archeozonotriletes chulus var. chulus (Cramer) Richardson and Lister. BAFC-Pl 1505 (5): X47. 8, Geminospora sp. cf. G. tuberculata var. tuberculata McGregor. BAFC-Pl 1505 (2): P52/4. 10, Cymbosporites cyathus Allen. BAFC-Pl 1505 (3): C37/3. 11, Grandispora pseudoreticulata (Menéndez and Pöthe de Baldis) Ottone. BAFC-Pl 1656 (1): F24/1.

Lophozonotriletes dentatus Hughes and Playford 1961
Figure 3.1

Occurrence. First record for the Mississippian of South America.

Genus Gorgonispora Urban 1971

Type species. Gorgonispora convoluta (Butterworth and Spinner) Playford 1976.

Gorgonispora sp. cf. G. crassa (Winslow) Higgs, Clayton and Keegan 1988
Figure 3.2

Studied material. BAFC-Pl 1501 (1): M29/2.

Dimension (1 specimen). Equatorial diameter 57 µm.

Remarks. Gorgonispora crassa (Winslow) Higgs et al. 1988 presents slightly larger diameter 70-110 µm. Occurrence. First record for the Mississippian of South America.

Genus Knoxisporites Potonié and Kremp emend. Neves 1961

Type species. Knoxisporites hagenii Potonié and Kremp 1954.

Knoxisporites sp.
Figure 3.3

Studied material. BAFC-Pl 1506 (4): B30/2.

Description. Spore radial trilete, subcircular to triangular amb. Suture indistinct. Equatorial cingulum not clearly defined. Proximal surface laevigate, laesurae indistinct with broad labra up to 10 µm thickness. Distal surface laevigate bearing three muri arranged in a triangular pattern with the ends of the triangle joining the equator in the interradial positions. Muri 6-8 µm wide often slightly expanded at the equator or cingulum.
Dimension (1 specimen). Equatorial diameter 48.7 µm.
Comparison. K. triangularis Higgs et al. 1988 is very similar to Knoxisporites sp., but differs in having a larger diameter and a clearly defined cingulum.
Remarks. The poor preservation of the specimen does not allow an accurate taxonomic assignment.

Suprasubturma LAMINATITRILETES Smith and Butterworth 1967
Subturma ZONOLAMINATITRILETES Smith and Butterworth 1967
Infraturma CRASSITI Bharadwaj and Venkatachala emend. Smith and Butterworth 1967

Genus Crassispora Bharadwaj emend. Keegan and Penney 1978

Type species. Crassispora kosankei (Potonié and Kremp) Bharadwaj 1957 emend. Smith and Butterworth 1967.

Crassispora invicta Playford 1971
Figure 3.9

Occurrence. First record for the Mississippian of South America.

Crassispora kosankei (Potonié and Kremp) Bharadwaj 1957 emend. Smith and Butterworth 1967
Figures 3.7-38

Remarks. The studied specimen shows the same sculpture like the Peruvian specimens illustrated by Azcuy and di Pasquo (2005).

Occurrence. (Illustrated): Viséan, Peru (Azcuy and di Pasquo, 2005, 2006), Bolivia (Fasolo et al., 2006). For other records see Azcuy and di Pasquo (2006).

Crassispora scrupulosa Playford 1971 emend. Playford and Satterthwait 1988
Figure 3.5

Occurrence. (Illustrated): Mississippian, Bolivia (di Pasquo, 2007b). (Listed): Mississippian, Argentina (Césari and Limarino, 1992), Bolivia (Vavrdová et al., 1996).

Crassispora trychera Neves and Ioannides 1974
Figure 3.6

Occurrence. (Illustrated): Tournaisian-Viséan, Brazil (Loboziak et al., 1991; Melo and Loboziak, 2003). Viséan, Bolivia (di Pasquo, 2007b). (Listed): Mississippian, Brazil (Loboziak et al., 1998).

Crassispora sp.
Figures 3.10-3.13

Studied material. BAFC-Pl 1502 (1): K46, C32/1-2, W40/3; BAFC- Pl 1655 (2): B26/3; BAFC-Pl 1504 (1): E48.

Description. Spore radial trilete, subcircular to oval amb, irregular due to the folded exine. Laesurae indistinct, imperfect curvaturae observed in some specimens. Proximal face smooth to chagrinate when is present because is usually absent. Distal face ornamented with discrete coni and spines, 0.6 µm high and wide, distributed randomly on the surface, bald areas frequently present. Equatorial crassitude narrow and irregular in thickness (cingulizone?) and wide varing from 2 to 3.5 µm.
Dimensions (5 specimens). Equatorial diameter 58-68 µm.
Comparison. Crassispora trychera Neves and Ioannides 1974 has a larger and more diverse ornamentation.

Infraturma CINGULATI (Potonié and Klaus) Dettmann 1963

Genus Bascaudaspora Owens 1983

Type species. Bascaudaspora canipa Owens 1983.

Bascaudaspora submarginata (Playford) Higgs, Clayton and Keegan 1988
Figure 3.4

Occurrence. (Illustrated): Late Tournaisian-Viséan, Colombia (Dueñas and Césari, 2006), Viséan, Peru (Azcuy and di Pasquo, 2005, 2006), Bolivia (Fasolo et al., 2006), Brazil (Melo and Loboziak, 2000). (Listed): Mississippian, Brazil (Iannuzzi and Pfefferkorn, 2002). For other records see Azcuy and di Pasquo (2006).

Genus Bellispores Artüz emend. Sullivan 1964

Type species. Bellispores nitidus (Horst) Sullivan 1964.

Bellispores sp.
Figures 3.14-3.16

Studied material. BAFC-Pl 1508 (1): C30/1; BAFC-Pl 1508 (2): Q28/1.

Description. Spore radial trilete. Amb triangular with straight or concave sides and rounded apices. Trilete mark distinct, simple, straight with rays that reached the spore margin. Proximal face smooth (?) and distal chagrinate to microgranulate-apiculate. Thickenings well defined on the distal surface, wart-like, with circular bases and slightly irregular margins, 5.8-8.12 µm wide. The radial thickenings do not continue to the equator, locating at the polar region leaving the apices free. Proximal subequatorial thickening (? kyrtome) continuous bearing apiculate elements, undulate to crenulate margin, 4.5-7 µm in thickness.
Dimensions (2 specimens). Overall diameter 50-56 µm.
Comparisons. The conspecific species Bellispores bellus Artüz 1957 and Bellispores nitidus (Horst) Sullivan 1964 (in agreement with Smith and Butterworth 1967) have a laevigate distal face, radial thickenings that continue to the equator and punctuations, foveae and short grooves developed on the bands and in the inner boundary of the border. Ahrensisporites Potonié and Kremp 1954 has kyrtomes curving toward the distal surface.
Remarks. The poorly preserved specimen illustrated in figure 3.16 shows much attenuated features respect to the one well preserved.

Genus Densosporites Berry emend. Butterworth, Jansonius, Smith and Staplin in Staplin and Jansonius 1964

Type species. Densosporites covensis Berry 1937.


Figure 3.
Cavate, pseudosaccate and cingulizonate spores from the Malimán Formation. Scale bar: all 15 µm (x 750), except 12: 10 µm (x 1000). Coordinates after EF (England Finder) graticule / esporas cavadas, pseudosacadas y cingulizonadas de la Formación Malimán. Escala gráfica: todos 15 µm (x 750), excepto 12: 10 µm (x 1000). Las coordenadas corresponden a la reglilla EF (England Finder). 1, Lophozonotriletes dentatus Hughes and Playford. BAFC-Pl 1655 (2): Y29. 2, Gorgonispora sp. cf. G. crassa (Winslow) Higgs, Clayton and Keegan. BAFC-Pl 1501 (1): M29/4. 3, Knoxisporites sp. BAFC-Pl 1506 (4): A30/4. 4, Bascaudaspora submarginata (Playford) Higgs, Clayton and Keegan. BAFC-Pl 1506 (4): A39/2. 5, Crassispora scrupulosa Playford emend. Playford and Satterthwait. BAFC-Pl 1508(4): F23/2-4. 6, Crassispora trychera Neves and Ioannides. BAFC-Pl 1508(2): F49/4. 7-8, Crassispora kosankei (Potonié and Kremp) Bharadwaj emend. Smith and Butterworth. BAFC-Pl 1504 (1): E30/3; 7, Proximal focus / foco proximal; 8, Distal focus / foco distal; 9, Crassispora invicta Playford. BAFC-Pl 1508 (1): W36. 10-13, Crassispora sp.; 10-12, BAFC-Pl 1502 (1): K46; 12, Detail of the ornamentation in the margin / detalle de la ornamentación en el margen; 13, BAFC-Pl 1655 (2): B26/3. 14-16, Bellispores sp.; 14-15, BAFC-Pl 1508 (1): C30/1; 14, Proximal focus showing the kyrtome / foco proximal mostrando el kirtoma; 15, Distal focus showing warts / foco distal mostrando verrugas; 16, BAFC-Pl 1508 (1): Q28/1. 17-18, Densosporites anulatus (Loose) Schopf, Wilson and Bentall. 17, BAFC-Pl 1506 (3): Y48; 18, BAFC-Pl 1506 (1): C37/1. 19-20, Densosporites gracilis Smith and Butterworth. 19, BAFC-Pl 1506 (1): H34; 20, BAFC-Pl 1506 (4): J56.

Densosporites anulatus (Loose) Schopf, Wilson and Bentall 1944
Figures 3.17-3.18

Occurrence. (Illustrated): Viséan, Peru (Azcuy and di Pasquo, 2005, 2006). (Listed): Mississippian, Bolivia (Fasolo et al., 2006), Brazil (Iannuzzi and Pfefferkorn, 2002); Pennsylvanian, Brazil (Souza, 2006). For other records see Azcuy and di Pasquo (2006).

Densosporites asperus Braman and Hills 1977
Figures 4.1-4.2

Remarks. Cristatisporites alpernii Staplin and Jansonius 1964 differs from this species in having an apiculate sculpture with some setose tips on the distal surface and a trilete mark that extends up to the central body.
Occurrence. First record for the Mississippian of South America.

Densosporites gracilis Smith and Butterworth 1967
Figures 3.19-3.20

Remarks. The specimens agree with the original description, but exhibit a slightly narrower cingulum (9.3 µm wide that is about 15 % of total spore diameter). ?Stenozonotriletes bracteolus (Butterworth and Williams) Smith and Butterworth 1967 has a granular ornamentation.
Occurrence. First record for the Mississippian of South America.

Densosporites regalis (Bharadwaj and Venkatachala) Smith and Butterworth 1967
Figures 4.7, 4.9

1984. Densosporites spitsbergensis Playford 1963; Higgs and Clayton, p. 24, pl. 2, fig. 17.
1988. Densosporites spitsbergensis Playford 1963; Higgs et al., p. 78-79, pl.15, figs. 13-14.
1991. Densosporites spitsbergensis Playford 1963; Utting, p. 98, pl. 4.2, fig. 15.
2000. Densosporites spitsbergensis Playford 1963; Melo and Loboziak, p. 153, pl. 2, fig. 11.
2003. Densosporites spitsbergensis Playford 1963; Melo and Loboziak, p. 174, pl. 3, fig. 1.

Remarks. The specimens here illustrated as well as the above cited show the same intraspecific variation according to Smith and Butterworth, 1967. Occurrence. (Illustrated): Viséan, Brazil (Melo and Loboziak, 2000, 2003). (Listed): Mississippian, Bolivia (Vavrdová et al., 1996; Díaz Martínez et al., 1999), Brazil (Iannuzzi and Pfefferkorn, 2002).

Densosporites secundus Playford and Satterthwait 1988
Figures 4.5-4.6

1960. Spinozonotriletes ? exiguus Staplin, p. 22, pl. 4, figs. 27-28.

Remarks. From the original diagnosis of this species given by Playford and Satterthwait (1988), some other features are evidenced here like the presence of some discrete biform elements between coni and spinae at the equatorial zone both may be coalescent in pairs or three elements.
Comparison. In agreement with Playford and Satterthwait (1988), Densosporites aculeatus Playford (1963) is distinguished from Densosporites secundus by having a wider cingulum (1/2 vs. 1/3 - 1/4 of central body) and the absence of labra.
Occurrence. First record for the Mississippian of South America.

Densosporites spinifer Hoffmeister, Staplin and Malloy 1955
Figures 4.3-4.4, 4.8

Occurrence. (Listed): Mississippian, Peru (Iannuzzi and Pfefferkorn, 2002).

Densosporites sp. cf. D. triangularis Kosanke 1950
Figures 4.10-4.12, 4.16

Studied material. BAFC-Pl 1655 (1): L25/1; BAFC-Pl 1655 (2): O28, F31/2, X28/2; BAFC-Pl 1503 (1): B35/3; BAFC-Pl 1506 (3): Z41; BAFC-Pl 1506 (5): K23, G26/4, Y35/2, Q28/3, U31/1, D24/3, Y36/3, B40.

Description. Spore radial trilete, amb subtriangular to oval, often with one apex more pronounced than the other two. Margin smooth to serrate. Laesurae not visible. Central area granulate and vermiculate, 2.3-4.6 µm wide, some elements with fused bases forming short rugulae. Cingulum well defined with cuniculus somewhat visible, width 6-8 µm (aprox. 1/8 of spore diameter or 12% of the spore ratio) with coni and subordinate verrucate, 1.2-2.3 µm high and wide, some coalescent mainly in its inner portion. In some specimens the cingulum shows an irregular thickness decreasing to the amb.
Dimensions (14 specimens). Equatorial diameter 49-69 µm.
Remarks. In all specimens the proximal surface is not visible. Scarce forms (figure 4.16) show transitional features to D. spinifer Hoffmeister et al. 1955. Comparison. D. triangularis Kosanke 1950 has a broader cingulum (more than 40% of the spore diameter) with foveolate dissections. Cymbosporites cyathus Allen 1965 has a lipped laesurae, inner border of the cingulum (or patina) less defined (cuniculus absent) and a dense ornamentation of verrucae, coni and rugulae outside the contact areas.
Occurrence. (Listed): Pennsylvanian, Brazil (Souza, 2006).

Genus Cristatisporites Potonié and Kremp 1954 emend. Butterworth, Jansonius, Smith and Staplin in Staplin and Jansonius, 1964

Type species. Cristatisporites indignabundus (Loose) Potonié and Kremp, 1954.

Cristatisporites indignabundus (Loose) Potonié and Kremp 1954 emend. Staplin and Jansonius 1964
Figure 4.17

Occurrence. (Illustrated): Viséan, Peru (Azcuy and di Pasquo, 2005, 2006), Bolivia (Fasolo et al., 2006); Pennsylvanian, Brazil (Souza, 2006). For other records see Azcuy and di Pasquo (2006).

Cristatisporites indolatus Playford and Satterthwait 1988
Figure 4.15

1992. Cristatisporites sp. cf. C. indolatus Playford and Satterthwait
1988; Césari and Limarino, p. 47, pl. 1, fig. 6.

Occurrence. (Illustrated): Mississippian, Argentina (Césari and Limarino, 1992). (Listed): Mississippian, Argentina (Pérez Loinaze and Césari, 2003).

Cristatisporites matthewsii Higgs, Clayton and Keegan 1988
Figures 4.18-4.20

Occurrence. (Illustrated): Late Devonian, Brazil (Loboziak et al., 1997a).

Cristatisporites peruvianus Azcuy and di Pasquo 2005
Figures 4.13-4.14

Occurrence. (Illustrated): Viséan, Peru (Azcuy and di Pasquo, 2005, 2006), Bolivia (di Pasquo, 2007b).

Genus Kraeuselisporites Leschik 1955 emend. Azcuy and di Pasquo 2005

Type species. Kraeuselisporites dentatus Leschik 1955.


Figure 4.
Cavate, pseudosaccate and cingulizonate spores from the Malimán Formation. Scale bar: all 15 µm (x 750), except 3-4: 10 µm (x 1000). Coordinates after EF (England Finder) graticule / esporas cavadas, pseudosacadas y cingulizonadas de la Formación Malimán. Escala gráfica: todos 15 µm (x 750), excepto 3-4: 10 µm (x 1000). Las coordenadas corresponden a la reglilla EF (England Finder). 1-2, Densosporites asperus Braman and Hills. 1, BAFC-Pl 1655(2): E39/3; 2, BAFC-Pl 1655(2): M36/1. 3-4, 8, Densosporites spinifer Hoffmeister, Staplin and Malloy. 3-4, BAFC-Pl 1508 (3): S45; 8, BAFC-Pl 1506(3): W28/1. 5-6, Densosporites secundus Playford and Satterthwait. 5, BAFC-Pl 1504(1): B42/3; 6, BAFC-Pl 1655(1): A31/3. 7, 9, Densosporites regalis (Bharadwaj and Venkatachala) Smith and Butterworth. 7, BAFC- Pl 1504(2): M37/1; 9, BAFC-Pl 1655 (2): P53. 10-12, 16, Densosporites sp. cf. D. triangularis Kosanke. 10, BAFC-Pl 1506(5): Y35/2; 11, BAFC-Pl 1506(5): E26/4; 12, BAFC-Pl 1655(1): SEM photo, the same specimen as 11 / foto tomada con microscopio electrónico de barrido (MEB), el mismo ejemplar que 11; 16, BAFC-Pl 1655(2): F31. 13-14, Cristatisporites peruvianus Azcuy and di Pasquo. 13, BAFC-Pl 1506(4): Z50; 14, BAFC-Pl 1503(1): W28/1. 15, Cristatisporites indolatus Playford and Satterthwait. BAFC-Pl 1506(5): D50/1. 17, Cristatisporites indignabundus (Loose) Potonié and Kremp emend. Staplin and Jansonius. BAFC-Pl 1506 (2): K24/4. 18-20, Cristatisporites matthewsii Higgs, Clayton and Keegan. 18, BAFC-Pl 1506 (5): G23; 19, BAFC-Pl 1506 (1): X21/1; 20, BAFC-Pl 1655 (2): B40/1.

Kraeuselisporites sp. cf. K. mitratus Higgs 1975
Figure 5.1

Studied material. BAFC-Pl 1504(3): Z52.

Remarks. This species is comparable with K. mitratus Higgs 1975 but the absence of more specimens does not allow a more accurate assignment.
Occurrence. (Illustrated): Viséan, Peru (Azcuy and di Pasquo, 2005, 2006), Brazil (Melo and Loboziak, 2003). (Listed): Viséan, Bolivia (Fasolo et al., 2006). For other records see Azcuy and di Pasquo (2006).

Genus Vallatisporites Hacquebard 1957

Type species. Vallatisporites vallatus Hacquebard 1957.

Vallatisporites pusillites (Kedo) Dolby and Neves 1970
Figure 5.2

Occurrence. (Listed): Mississippian, Bolivia (Vavrdová et al., 1996). Late Devonian, Brazil (Loboziak et al., 1997a, 1997b; Limachi et al., 1996).

Infraturma PATINATI Butterworth and Williams 1958

Genus Archaeozonotriletes (Naumova) Allen 1965

Type species. Archaeozonotriletes variabilis (Naumova) Allen 1965.

Archaeozonotriletes chulus var. chulus (Cramer) Richardson and Lister 1969
Figure 5.7

Remarks. These spores are atributed to A. chulus var. chulus, for its cingulate/patinate character, thin proximal exine and absence of ornamentation. Occurrence. (Illustrated): Early-Middle Devonian, Brazil (Melo and Loboziak, 2003; Rubinstein et al., 2005), Bolivia (McGregor, 1984).

Genus Cymbosporites Allen 1965

Type species. Cymbosporites cyathus Allen 1965.

Cymbosporites catillus Allen 1965
Figures 2.3-2.4

Occurrence. (Illustrated): Middle-Late Devonian, Argentina (Ottone, 1996), Brazil (Burjack et al., 1987; Loboziak et al., 1988, 1992, 1997a; Loboziak and Melo, 2000, 2002; Melo and Loboziak, 2003), Bolivia (Pérez Leyton, 1990; di Pasquo, 2007a). (Listed): Middle Devonian, Brazil (Dino, 1999). Middle-Late Devonian, Bolivia (Pérez Leyton, 1991; Vavrdová et al., 1996). Late Devonian, Chile (Rubinstein et al., 1996).

Cymbosporites cyathus Allen 1965
Figure 2.10

Occurrence. (Illustrated): Middle-Late Devonian, Brazil (Burjack et al., 1987; Loboziak et al., 1988, 1992; Melo and Loboziak, 2003), Bolivia (Pérez Leyton, 1991; Vavrdová et al., 1996; di Pasquo, 2007a). (Listed): Middle-Late Devonian, Brazil (Loboziak et al., 1997a; Dino, 1999).

Suprasubturma PSEUDOSACCITRILETES Richardson 1965
Infraturma MONOPSEUDOSACCITI Smith and Butterworth 1967

Genus Auroraspora Hoffmeister, Staplin and Malloy 1955

Type species. Auroraspora solisorta Hoffmeister, Staplin and Malloy 1955.

Auroraspora macra Sullivan 1968
Figures 5.9, 5.14

Occurrence. (Illustrated): Late Tournaisian-Viséan, Colombia (Dueñas and Césari, 2006), Viséan, Peru (Azcuy and di Pasquo, 2005, 2006), Bolivia (di Pasquo, 2007b), Brazil (Loboziak et al., 1992, 1997a; Melo and Loboziak, 2003). (Listed): Late Devonian, Brazil (Burjack et al., 1987; Loboziak et al., 1997b). Late Devonian- Mississippian, Bolivia (Pérez Leyton, 1991; Vavrdová et al., 1996), Chile (Rubinstein et al., 1996). Mississippian, Brazil and Bolivia (Iannuzzi and Pfefferkorn, 2002). For other records see Azcuy and di Pasquo (2006).

Auroraspora solisorta Hoffmeister, Staplin and Malloy 1955
Figure 5.15

Occurrence. (Illustrated): Late Tournaisian-Viséan, Colombia (Dueñas and Césari, 2006), Mississippian, Brazil (Loboziak et al., 1992; Melo and Loboziak, 2003), Bolivia (di Pasquo, 2007b). (Listed): Late Devonian, Bolivia (Vavrdová et al., 1996). Late Devonian- Mississippian, Chile (Rubinstein et al., 1996). Mississippian, Brazil (Loboziak et al., 1998; Iannuzzi and Pfefferkorn, 2002).

Genus Geminospora Balme 1962 emend. Playford 1983

Type species. Geminospora lemurata Balme 1962.

Geminospora lemurata Balme 1962 emend. Playford 1983
Figures 2.5-2.6, 2.9

Occurrence. (Illustrated): Middle-Late Devonian, Argentina (Ottone, 1996), Brazil (Burjack et al., 1987; Loboziak et al., 1988, 1992, 1997a; Rodrigues et al., 1995; Dino, 1999; Loboziak and Melo, 2000, 2002; Melo and Loboziak, 2003), Bolivia (Ottone and Rosello, 1996; di Pasquo, 2007a). (Listed): Middle- Late Devonian, Bolivia (Pérez Leyton, 1991; Limachi et al., 1996; Vavrdová et al., 1996), Argentina (Rodríguez Amenábar et al., 2003). Late Devonian, Brazil (Loboziak et al., 1997b).

Geminospora sp. cf. G. tuberculata var. Tuberculata
McGregor 1996
Figure 2.8

Studied material. BAFC-Pl 1505 (2): P52/0-4.

Dimension (1 specimen). Equatorial diameter 45.2 µm.
Remarks. The lack of additional specimens does not allow a more precise assignment.
Occurrence. First record for the Middle Devonian of South America.

Genus Grandispora Hoffmeister, Staplin and Malloy 1955

Type species. Grandispora spinosa Hoffmeister, Staplin and Malloy 1955.

Grandispora debilis Playford 1971
Figure 5.7

Remarks. The specimens here studied are slightly larger (81.2-87 µm) than the original material. Occurrence. (Illustrated): Viséan, Peru (Azcuy and di Pasquo, 2005, 2006), Bolivia (Fasolo et al., 2006). (Listed): Mississippian, Argentina (Sessarego and Césari, 1989; Césari and Limarino, 1992; Rodríguez Amenábar et al., 2003; Pérez Loinaze and Césari, 2003). For other records see Azcuy and di Pasquo (2006).

Grandispora notensis Playford 1971
Figure 5.10

Occurrence. (Illustrated): Mississippian, Bolivia (di Pasquo, 2007b). (Listed): Late Devonian, Bolivia (Vavrdová et al., 1996).

Grandispora pseudoreticulata (Menéndez and Pöthe de Baldis) Ottone 1996
Figure 2.11

Occurrence. (Illustrated): Middle Devonian, Paraguay (Menéndez and Pöthe de Baldis, 1967). Middle-Late Devonian, Argentina (Ottone, 1996), Bolivia (Ottone and Rosello, 1996; di Pasquo, 2007a). (Listed): Middle Devonian, Argentina (Rodríguez Amenábar et al., 2003).

Grandispora spiculifera Playford 1976
Figures 5.12-5.13

Occurrence. (Illustrated): Late Tournaisian-Viséan, Colombia (Dueñas and Césari, 2006), Mississippian, Brazil (Loboziak et al., 1991, 1992; Melo et al., 1999; Melo and Loboziak, 2000, 2003). (Listed): Mississippian, Brazil (Loboziak et al., 1998; Iannuzzi and Pfefferkorn, 2002), Argentina (Pérez Loinaze and Césari, 2003).

Genus Spelaeotriletes Neves and Owens 1966

Type species. Spelaeotriletes triangulus Neves and Owens 1966.

Spelaeotriletes arenaceous Neves and Owens 1966
Figures 5.4-5.5

Remarks. The specimens here studied are smaller (54.5-75.4 µm) than the original material. Occurrence. (Illustrated): Late Tournaisian-Pennsylvanian, Brazil (Loboziak et al., 1991; Melo et al., 1999; Melo and Loboziak, 2000, 2003). Mississippian, Bolivia (Azcuy and Ottone, 1987; Fasolo et al., 2006). (Listed): Mississippian, Brazil (Loboziak et al., 1998, 2000; Iannuzzi and Pfefferkorn, 2002).

Spelaeotriletes echinatus (Hacquebard) Utting 1987
Figure 5.6

Occurrence. First record for the Mississippian of South America.

Spelaeotriletes obtusus Higgs 1975
Figure 5.3

Occurrence. (Illustrated): Late Famennian-Tournaisian, Brazil (Melo and Loboziak, 2003). (Listed): Late Devonian, Brazil (Loboziak et al., 1997b). Mississippian, Bolivia (Vavrdová et al., 1996).

Genus Velamisporites Bharadwaj and Venkatachala 1962

Type species. Velamisporites rugosus Bharadwaj and Venkatachala 1962.

Velamisporites perinatus (Hughes and Playford) Playford 1971
Figures 5.8, 5.11

Occurrence. First record for the Mississippian of South America.

Genus Colatisporites Williams in Neves, Gueinn, Clayton, Ioannides, Neville and Kruszewska 1973

Type species. Colatisporites decorus (Bharadwaj and Venkatachala) Williams in Neves, Gueinn, Clayton, Ioannides, Neville and Kruszewska 1973.

Colatisporites decorus (Bharadwaj and Venkatachala) Williams in Neves, Gueinn, Clayton, Ioannides, Neville and Kruszewska 1973
Figures 5.16-5.17

Occurrence. (Illustrated): Late Tournaisian-Viséan, Colombia (Dueñas and Césari, 2006), Viséan, Brazil (Melo and Loboziak, 2003), Bolivia (di Pasquo, 2007a), Peru (Azcuy and di Pasquo, 2005, 2006). (Listed): Mississippian, Brazil (Loboziak et al., 1992, 1998, 2000; Iannuzzi and Pfefferkorn, 2002). For other records see Azcuy and di Pasquo (2006).

Paleomicroplankton

Micrhystridium sp.
Figure 1.15

Description. Vesicle subcircular in outline, wall psilate and thin with at least fifteen discrete, heteromorphic processes with broad bases that sharply tapered to the end with simple apices or bearing a little spine.
Processes interior hollow and in free communication with the vesicle cavity. No excystment structure observed.
Dimensions (1 specimen). Overall diameter 35 µm, diameter of vesicle 27 µm, basal breath of processes 2-3 µm, length of processes 4.6 - 7 µm.

Cymatiosphaera sp.
Figures 1.17-1.18

Description. Vesicle subspherical to polyhedral in outline. Vesicle surface granular, divided into around 12 polygonal fields, generally pentagonal; muri 2 to 3.5 µm height, luminae 7-15 µm in diameter.

Dimensions (2 specimens). Overall diameter 46-58 µm.

Comparison. Cymatiosphaera apiaria Ottone 1996 has larger fields with a fine reticulate sculpture inside.

Dictyotidium sp.
Figures 1.7-1.8

Description. Vesicle circular to subcircular in outline, deformed by frequent folds. Vesicle surface divided into polygonal to subcircular psilate fields that formed a well defined reticulum. Muri are psilate and narrow. No excystment structure is observed. Dimensions (2 specimens). Overall diameter 55-63 µm, muri 1 µm height, fields 2.3-3.6 µm in diameter. Wall 1.2 µm thick.

Comparisons. Dictyotidium sp. is similar to D. torosum Playford in Playford and Dring 1981 but differs in having a thinner and folded wall. D. prolatum Playford in Playford and Dring 1981 is smaller, has a thinner wall and slightly sinuous muri.


Figure 5.
Cavate, pseudosaccate and cingulizonate spores from the Malimán Formation.
Scale bar: all 15 µm (x 750). Coordinates after EF (England Finder) graticule / esporas cavadas, pseudosacadas y cingulizonadas de la Formación Malimán. Escala gráfica: todos 15 µm (x 750). Las coordenadas corresponden a la reglilla EF (England Finder). 1, Kraeuselisporites sp. cf. K. mitratus Higgs. BAFC-Pl 1504 (3): Z52. 2, Vallatisporites pusillites (Kedo) Dolby and Neves. BAFC-Pl 1655(1): Z30/4. 3, Spelaeotriletes obtusus Higgs. BAFC-Pl 1501 (1): H33. 4-5, Spelaeotriletes arenaceous Neves and Owens. 4, BAFC-Pl 1508 (5): E46; 5, BAFC-Pl 1655 (2): T57/1. 6, Spelaeotriletes echinatus (Hacquebard) Utting. BAFC-Pl 1501 (1): B26. 7, Grandispora debilis Playford. BAFC-Pl 1508 (3): K28/1. 8, 11, Velamisporites perinatus (Hughes and Playford) Playford. 8, BAFC-Pl 1506 (3): U41/2; 11, BAFC-Pl 1501 (1): Q31/1. 9, 14, Auroraspora macra Sullivan. 9, BAFC- Pl 1508 (1): J58/3; 14, BAFC-Pl 1504 (3): D33/1. 10, Grandispora notensis Playford. BAFC-Pl 1508 (4): Z51/1. 12-13, Grandispora spiculifera Playford. BAFC-Pl 1508 (1): Z27/3. 15, Auroraspora solisorta Hoffmeister, Staplin and Malloy. BAFC-Pl 1501 (2): V28/3. 16-17, Colatisporites decorus (Bharadwaj and Venkatachala) Williams in Neves, Gueinn, Clayton, Ioannides, Neville and Kruszewska. 16, BAFC-Pl 1506 (1): X28; 17, BAFC-Pl 1501 (1): R38/3.

Acknowledgements

The authors are grateful to G. Holfeltz for preparing the palynological samples. This research was supported with funds from Agencia Nacional de Promoción Científica y Tecnológica PICTR 00313/03, from the University of Buenos Aires UBACYT X 136 and the Consejo Nacional de Investigaciones Científicas y Técnicas PIP 5518 CONICET. This is a contribution to the International Geological Correlation Program 471 (IGCP-471).

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Recibido: 7 de noviembre de 2005.
Aceptado: 17 de julio de 2007.