Review of Palaeobotany and Palynology 138 (2006) 31 42

www.elsevier.com/locate/revpalbo

Palynological evidence of Early Carboniferous sedimentation

in the Llanos Orientales Basin, Colombia
H. Duenas a, S.N. Cesari b,*
a
Academia Colombiana de Ciencias Exactas, Fsicas y Naturales Transversal 27 No. 39A-63 Bogota, Colombia
b
Museo Argentino de Cs. Naturales, B. Rivadavia, Av. Angel Gallardo 470, C1405DJR Buenos Aires, Argentina
Received 17 July 2004; received in revised form 21 September 2005; accepted 6 October 2005

Abstract

Early Carboniferous palynological assemblages from the SM-4 well located in the Llanos Orientales Basin constitute the only
definitively geological evidence of Carboniferous strata in this Basin. Stratigraphically significant species include: Anapiculatis-
porites concinnus, Apiculiretusispora mutiseta, Grandispora spiculifera, Indotriradites dolianitii morphon, Spelaeotriletes pre-
tiosus and Prolycospora rugulosa. The presence of scarce acritarchs indicates a shallow marine environment. A Tournaisian age,
probably reaching the Visean, is proposed for the interval 20102340 ft based on the presence of distinctive spore species with
previous records in the Visean and Tournaisian of Western Europe and Western Gondwana. This report increases the knowledge
about the distribution and composition of the Early Carboniferous palynological assemblages in Northern Gondwana.
D 2005 Elsevier B.V. All rights reserved.

Keywords: Lower carboniferous; Palynology; Colombia

1. Introduction summaries by Duenas (2002) and Duenas and Cesari

The South American Early Carboniferous palyno-
logical record is constrained to Parnaiba and Amazon
Basins in Brazil (Melo and Loboziak, 2000, 2003),
Ambo Formation in Peru (Azcuy and di Pasquo,
2005), Retama Formation in Bolivia (Azcuy and
Ottone, 1987) and Rio Blanco Basin in Argentina
(Cesari and Limarino, 1992, 1995). These assemblages
have mixed Euramerican/Gondwanan compositional
characters, but the similarities among some of them
are still uncertain. The discovery of Early Carbonifer-
ous assemblages in strata of the SM-4 well in the
Llanos Orientales Basin, Colombia, was reported in
* Corresponding author.
E-mail address: scesari@macn.gov.ar (S.N. Cesari).
0034-6667/$ - see front matter D 2005 Elsevier B.V. All rights reserved.
doi:10.1016/j.revpalbo.2005.10.002
(2003). The section of the SM-4 well, analyzed here, is
the only known reference to Early Carboniferous sedi-
ments in the whole Colombian Llanos Basin.
The aim of this paper is to contribute to the knowl-
edge of the composition and distribution of the Peri-
gondwanan palynofloras. The Early Carboniferous
assemblages are compared, for purposes of correlation,
with those that characterize palynostratigraphic units in
northern and western Gondwana. This study is part of a
palynological research programme on all the sedimen-
tary sequence of the well.
The palynological samples were collected from the
depth interval 20102340 ft, of the SM-4 well in the
Llanos Basin. This structural depression located in the
eastern part of Colombia is filled by a sedimentary
sequence clearly divisible on a regional scale into
three time units dated as Paleozoic, Cretaceous and
32 H. Duenas, S.N. Cesari / Review of Palaeobotany and Palynology 138 (2006) 3142
Tertiary. The published stratigraphic data from this vast
grass and forest-covered region are scarce; the major
part of the information corresponds to oil-companies
reports.
2. Geological setting
The Llanos Orientales Basin is a lowland area,
which covers some 250,000 km2 of the eastern part
of Colombia. It is bounded to the west, by the faulted
eastern limit of the Andean Eastern Cordillera (Fig. 1);
Fig. 1. Location map of the studied area.
to the north, by the Venezuelan Colombian border
which approximately corresponds to a structural high
known as the Arauca Arch; to the south-west, by the
Macarena Range and to the south-east and east by
outcrops of igneous and metamorphic rocks which
belong to the Guyana Shield (Fig. 1).
This is a northeastern trending structural depression,
deepest adjacent to the Eastern Cordillera. The sedi-
mentary sequence, which fills this depression, is clearly
divisible on a regional scale into three chronostrati-
graphic units, which have been dated as Paleozoic,
Cretaceous and Tertiary. These units are separated re-
spectively by well-known regional unconformities and
overlie a Precambrian crystalline and metamorphic
basement, which forms a structural platform that dips
gently (258) to the west. This stratigraphic sequence
partially outcrops, as faulted blocks, along the Eastern
Cordillera Foothills as well as in the Macarena Range.
All the stratigraphic information from the grass and
forest-covered Llanos foreland comes from seismic
and well data.
During Precambrian time, the vast area of the Llanos
Orientales Basin was a part of relatively immobile
pericratonic belt. At the end of the Precambrian, the
igneousmetamorphic basement subsided allowing the
accumulation of a thick sequence of Late Precambrian
and Paleozoic sediments. There are three different Pa-
leozoic depocenters in the Llanos Basin. The first one
runs parallel to the Llanos Foothills with more than
15,000 ft. of sedimentary rocks. The second one is
located on the southwest part of the Basin in the
Apiay area, where the seismic lines show the presence
of more than 20,000 ft. of Paleozoic strata. The third is
located at the northeastern part of the Basin with a
Paleozoic infill of more than 8500 ft. and related to
the Cambrian Basin, which covers this part of the
Llanos Basin and the Barinas Basin in Venezuela,
where more than 8500 ft. of Paleozoic stratigraphic
section has been reported (Feo-Codecido et al., 1984;
Sinanoglu, 1986). The Llanos Basin can be considered
as a major Paleozoic intracratonic Basin.
The Paleozoic section ranges in age from Cambrian
to Carboniferous. Correlation of these Paleozoic sedi-
ments with those outcropping in the Eastern Cordillera
is uncertain because only unmetamorphased sediments
Silurian have been reported in the Cordillera area. The
Paleozoic sediments have received very little attention
to the present, probably because the low hydrocarbon
potential of these strata has been wrongly assumed.
The stratigraphy of the Llanos Orientales reveals that
the Tertiary and Cretaceous sediments present clear
biostratigraphic and lithostratigraphic subdivisions
 H. Duenas, S.N. Cesari / Review of Palaeobotany and Palynology 138 (2006) 3142 33

(Duque-Caro, 1997; Germeraad et al., 1968; Muller et

al., 1987). In contrast, the Paleozoic sediments possess
neither a stratigraphic name nor lithostratigraphic sub-
divisions. However, seismic information shows that
these sediments can reach more than 20,000 feet in
thickness and that it is possible to subdivide them,
litostratigraphic subdivision of the Paleozoic was pro-
posed only in the wells Negritos-1 and the Heliera-1
and the name of Negritos Formation was proposed
(Ulloa et al., 1982). At present, the Paleozoic sequence
is considered as a unit unconformably underlying Cre-
taceous rocks and overlying the granitic basement of
the Guyana Shield.
The sedimentary sequence of the SM-4 well can be
divided in a lower unnamed Paleozoic section includ-
ing Late Devonian and Early Carboniferous sediments
that are separated by an unconformity from the Ter-
Fig. 2. Generalized scheme of the stratigraphical succession in the
tiary section represented by the Carbonera Formation SM-4 well, showing the position of the studied palynological section.
and Leon Formations (Notestein et al., 1944). The
Early Carboniferous strata found in the interval
but frequently is poor to moderate, and the palyno-
20102340 ft of the SM-4 well yielded assemblages
morphs are thermally mature and are dark brown to
dominated by terrestrial palynomorphs and since acri-
black.
tarchs are poorly represented, a shallow marine envi-
ronment was suggested (Duenas and Cesari, 2003).
4. Identified palynomorphs
Late Devonian palynological assemblages, presently
being study by the authors, are recognized below the
The identified palynomorphs (spores and acritarchs)
2340 ft. These Devonian palynofloras are character-
are listed below in alphabetical order.
ized by the presence of Hystricosporites spp., Ancyr-
ospora spp. and Teichertospora torquata among other
4.1. Spores
species.
At the end of the Early Carboniferous, there were
Anapiculatisporites concinnus Playford, 1962 (Plate
strong tectonic movements, characterized by block
I, 1)
faulting which produce the subdivision of the Andean
Anaplanisporites cf. A. denticulatus Sullivan, 1964
region in eastern and western provinces separated by
(Plate I, 2)
the ancestral Cordillera Central. The Permian was a
period of emergence. Widespread igneous ctivity took Remarks: The present specimens differ from the orig-
place and was particularly intense in the Central Cor- inal material in having a smaller diameter (2940
dillera area. It was during this time, that the basis for Am) and the sculpture projecting at the equatorial
the structural features of the Llanos Basin occurred. margin.
Permian sediments have not been reported in the Llanos
Apiculiretusispora multiseta (Luber) Butterworth and
area.
Spinner, 1967 (Plate I, 3)
Auroraspora macra Sullivan, 1968 (Plate I, 12)
3. Materials and methods
Auroraspora solisorta Hoffmeister et al., 1955 (Plate
I, 15)
Ten samples of cuttings were studied from the inter-
Bascaudaspora submarginata (Playford) Higgs et al.,
val 20102340 ft of borehole SM-4 (Fig. 2). Standard
1988 (Plate I, 10)
palynological preparation techniques using HF, HCl
Calamospora liquida Kosanke, 1950 (Plate I, 13)
were applied to the samples. Sample residues were
Calamospora cf. Calamospora nigrata (Naumova)
mounted on microscope slides in Canada balsam and
Allen, 1965 (Plate I, 5)
all slides are stored at the National Core Library,
Colombian Petroleum Institute, Bucaramanga (Colom- Remarks: The present specimens are smaller (4450
bia). The preservation is variable within the samples, Am) than those described by Naumova (1953).
34 H. Duenas, S.N. Cesari / Review of Palaeobotany and Palynology 138 (2006) 3142

Colatisporites decorus (Bharadwaj and Venkata- Crassispora maculosa (Knox) Sullivan, 1964 (Plate I,
chala) Williams in Neves et al., 1973 (Plate 17)
I, 14) Crassispora sp. (Plate I, 18)
 H. Duenas, S.N. Cesari / Review of Palaeobotany and Palynology 138 (2006) 3142 35

Remarks: The studied specimens differ from the known
species of the genus in the distal very fine and dense
sculpture.
Cristatisporites sp. (Plate I, 7)
Remarks: The scarce number of well preserved speci-
mens prevents a close comparison with known species
of this genus, characterized by a great variability be-
tween representatives of the same species.
Cymbosporites acutus (Kedo) Byvscheva, 1985 (Plate
I, 9)
Densosporites rarispinosus Playford, 1963 (Plate I,
6)
Discernisporites micromanifestus (Hacquebard) Sabry
and Neves, 1971 (Plate I, 4)
Endosporites sp. (Plate I, 16)
Remarks: The specimens resemble Diducites poljessi-
cus (Kedo) Van Veen, 1981 but do not have the two-
layered exoexine.
Grandispora spiculiferaPlayford, 1976 (Plate I, 8)
Indotriradites dolianitii (Daemon) Loboziak et al.,
1999 (Plate I, 11)
Indotriradites daemonii Loboziak et al., 1999 (Plate
II, 13)
Leiotriletes sp. (Plate II, 8)
Remarks: The specimens are treated at the generic level
only, because they are not clearly assignable to any
described species.
Prolycospora rugulosa (Butterworth and Spinner)
Turnau, 1978 (Plate II, 4)
Punctatisporites irrasus Hacquebard, 1957 (Plate II,
15)
Raistrickia cf. Raistrickia clavata Hacquebard emend.
Playford, 1964 (Plate II, 9)
Comparisons: The specimens are very dark in colour
not allowing a precise description of some morpho-
logical characters. Nevertheless, they seem co-spe-
cific with R. clavata by the varied sculpture.
Retusotriletes crassus Clayton, 1980 in Clayton et al.,
1980 (Plate II, 6)
Retusotriletes incohatus Sullivan, 1964 (Plate II, 7)
Retusotriletes mirabilis (Neville) Playford, 1978 (Plate
I, 3)
Rugospora flexuosa (Jushko) Streel in Becker et al.,
1974 (Plate II, 14)
Schopfites claviger Sullivan emend. Higgs et al., 1988
(Plate II, 5)
Spelaeotriletes pretiosus (Playford) Utting, 1987 (Plate
II, 11)
Spelaeotriletes sp. (Plate II, 16)
Remarks: According to the detailed revision of Higgs et
al. (1988) and Playford et al. (2001) of some different
species of Spelaeotriletes our specimens are distin-
guishable from the morphotaxa Spelaeotriletes triangu-
lus/Spelaeotriletes arenaceous by having diminutive
sculpture mostly of galeae, grana and coni and a slight-
ly more thicker exine at the equatorial margin forming a
characteristically narrow dark area.
Vallatisporites splendens Staplin and Jansonius, 1964
(Plate II, 1)
Vallatisporites hystricosus (Winslow) Byvscheva, 1985
(Plate II, 12)

Plate I.

1. Anapiculatisporites concinnus Playford, 1962. Slide 2100-2130(2): M30/2, X1000.

2. Anaplanisporites cf.A. denticulatus Sullivan, 1964. Slide 2130-2160(2): H39/4, X750.
3. Apiculiretusispora multiseta (Luber) Butterworth and Spinner, 1967. Slide 2100-2130: T45, X750.
4. Discernisporites micromanifestus (Hacquebard) Sabry and Neves, 1971. Slide 2280-2310(2): H38/3, X750.
5. Calamospora cf. Calamospora nigrata (Naumova) Allen, 1965. Slide 2070-2100(1): M42, X750.
6. Densosporites rarispinosus Playford, 1963. Slide 2100-2130(1): J48/4, X1000.
7. Cristatisporites sp. Slide 2190-2220(1): C39, X750.
8. Grandispora spiculifera Playford, 1976. Slide 2100-2130(2): J49/4, X750.
9. Cymbosporites acutus (Kedo) Byvscheva, 1985. Slide 2100-2130(2): R33/2, X750.
10. Bascaudaspora submarginata (Playford) Higgs et al., 1988. Slide 2310-2340(1): J45, X750.
11. Indotriradites dolianitii (Daemon) Loboziak et al., 1999. Slide 2160-2190(1): V49, X750.
12. Auroraspora macra Sullivan, 1968. Slide 2070-2100(1): T41/3, X1000.
13. Calamospora liquida Kosanke, 1950. Slide 2040-2070(1): C38/1, X750.
14. Colatisporites decorus (Bharadwaj and Venkatachala) Williams in Neves et al., 1973. Slide 2100-2130(1): E39/4, X750.
15. Auroraspora solisorta Hoffmeister et al., 1955. Slide 2100-2130(2): J41/3, X750.
16. Endosporites sp. Slide 2100-2130(1): B33, X750.
17. Crassispora maculosa (Knox) Sullivan, 1964. Slide 2100-2130(1): Q36/4, X750.
18. Crassispora sp. Slide 2280-2310(1): Q34/4, X750.
36 H. Duenas, S.N. Cesari / Review of Palaeobotany and Palynology 138 (2006) 3142

Verrucosisporites irregularis Philips and Clayton, 1980 4.2. Acritarchs

(Plate II, 10)
Verrucosisporites nitidus Playford, 1964 (Plate II, Maranhites insulatus Burjack and Oliveira, 1989 (Plate
2) III, 1)
 H. Duenas, S.N. Cesari / Review of Palaeobotany and Palynology 138 (2006) 3142 37

Plate III.

1. Maranhites insulatus Burjack and Oliveira, 1989. Slide 2130-2160(1): S34/2, X750.
23. Gorgonisphaeridium cf. Gorgonisphaeridium winslowiae Staplin et al., 1965. Slide 2070-2100(1): O47/2. X1250, X1000.
4. Scolecodont. Slides 2190-2220(1): G33 X750.
5. Veryhachium pannuceum Wicander and Loeblich. Slide 2010-2040(1): D35, X1000.
6. Veryhachium europaeum Stockmans and Williere. Slide 2100-2130(2) L34/3, XX1000.

Gorgonisphaeridium cf. Gorgonisphaeridium winslo- 5. Age assessment and paleoenvironment

wiae Staplin, Jansonius and Pocock, 1965 (Plate
III, 2, 3) Despite the variable state of preservation, the assem-
Comparisons: The specimens resemble closely G. win- blages contain a number of stratigraphically significant
slowiae, but are smaller in diameter (2533 Am). species, including those with an Euramerican affinity

Plate II.

1. Vallatisporites splendens Staplin and Jansonius, 1964. Slide 2130-2160(2): G42, X750.
2. Verrucosisporites nitidus Playford, 1964. Slide 2220-2250(1): O49/3, X1000.
3. Retusotriletes mirabilis (Neville) Playford, 1978. Slide 2100-2130(1): M35/1, X750.
4. Prolycospora rugulosa (Butterworth and Spinner) Turnau, 1978. Slide 2010-2040(1): J32/3, X1000.
5. Schopfites claviger Sullivan emend. Higgs et al., 1988. Slide 2100-2130(1): E41/4, X750.
6. Retusotriletes crassus Clayton et al., 1980. Slide 2100-2130(2): F36/1, X750.
7. Retusotriletes incohatus Sullivan, 1964. Slide 2100-2130(1): L37/4, X750.
8. Leiotriletes sp. Slide 2160-2190(2): V36/3, X750.
9. Raistrickia cf. Raistrickia clavata Hacquebard emend. Playford, 1964. Slide 2100-2130(2): N32, X1000.
10. Verrucosisporites irregularis Philips and Clayton, 1980. Slide 2280-2310(1): K35/1, X750.
11. Spelaeotriletes pretiosus (Playford) Utting, 1987. Slide 2160-2190(1): H35, X750.
12. Vallatisporites hystricosus (Winslow) Byvscheva, 1985. Slide 2220-2250(1): G37/4, X750.
13. Indotriradites daemonii Loboziak et al., 1999. Slide 2130-2160(1): W43, X750.
14. Rugospora flexuosa (Jushko) Streel in Becker et al., 1974. Slide 2280-2310(1): L42/2, X750.
15. Punctatisporites irrasus Hacquebard, 1957, Slide 2100-2130(2): W44, X750.
16. Spelaeotriletes sp. Slide 2010-2040(1): F43, X750.
38 H. Duenas, S.N. Cesari / Review of Palaeobotany and Palynology 138 (2006) 3142
and those with Gondwanan previous records (Fig. 3).
Among the northern species, many are characteristic of
Early Carboniferous biozones (Clayton et al., 1977).
The middle-lower Tournaisian biozone Spelaeotriletes
pretiosusRaistrickia clavata (PC), shares the presence
of Punctatisporites irrasus, Retusotriletes incohatus
and Auroraspora macra with the Colombian assem-
blages. The upper Tournaisian biozone Schopfites cla-
vigerAuroraspora macra (CM) is characterized
Verrucosisporites nitidus, Apiculiretusispora multiseta,
Colatisporites decorus, species present in Colombia
too.
Grandispora spiculifera is a characteristic species of
Tournaisian sections in Euramerica, it names a Tour-
naisian Assemblage palynozone in northwestern Aus-
tralia (Playford, 1985) and occurs in latest Devonian
Visean associations from the Amazon Basin (Melo and
Loboziak, 2003) as well.
Anapiculatisporites concinnus, Schopfites claviger
and Prolycospora rugulosa, are taxa that characterize
the Visean in the northern hemisphere. Moreover, the
two later spores and Auroraspora solisorta have their
incoming in the Late Visean of Brazil (Melo and
Fig. 3. Chronostratigraphic distribution of selected species recognized in the SM-4 well, based on Clayton et al. (1977, 1980), Higgs et al. (1988),
Melo and Loboziak (2003) and Playford (1991).
Loboziak, 2003). Also, the Gondwanan species Indo-
triradites daemonii and I. dolianitii are usually pres-
ent in Visean strata of northern Brazilian basins
(Daemon, 1974; Melo and Loboziak, 2003), although
also occur occasionally in late Tournaisian strata
(Loboziak et al., 1999). Indotriradites dolianitii is
also recognized in the Visean of the Grand Erg
occidental, Algerian Sahara (Lanzoni and Magloire,
1969).
The occurrence in the assemblages of miospore
taxa such as Vallatisporites hystricosus and Rugos-
pora flexuosa may reflect reworking from Strunian
sediments.
A Tournaisian age, probably reaching the Visean, is
proposed for the interval 20102340 ft by the presence
of distinctive spore species with previous records in the
Tournaisian and Visean of Western Europe and Western
Gondwana. It is difficult to define accurately the limits
because of the continuous presence of several species
through the sequence, probably due to reworking or
caving (Fig. 4). In addition, the high degree of thermal
maturity of many palynomorphs avoids an accurate
identification. Nevertheless, it is remarkable the ab-
 H. Duenas, S.N. Cesari / Review of Palaeobotany and Palynology 138 (2006) 3142
Fig. 4. Distribution chart of selected miospore taxa in well SM-4.
sence of the characteristic Visean species: Indotrira-
dites morphon, Anapiculatisporites concinnus and Pro-
lycospora rugulosa below the 2250 ft. Moreover, the
StrunianEarly Tournaisian species Rugospora flex-
uosa and Vallatisporites hystricosus were identified
only below the 2220 ft.
All the studied interval contains acritarchs and pra-
sinophyceae in variable proportions and the section
21902310 ft also contains scolecodonts (Plate III, 4).
Some of the marine species seem to be autochthonous
but many are diagnostic taxa of older associations, such
as Umbellasphaeridium sp., Veryhachium pannuceum
Wicander and Loeblich (Plate III, 5), V. europaeum
Stockmans and Williere (Plate III, 6) or Crassiangulina
tessellita Jardine et al., therefore are considered
reworked. If we assume that Maranhites insulatus,
Gorgonisphaeridum cf. winslowiae and scolecodonts
are autochthonous it is possible to suggest a shallow
marine environment for the sequence.
39
6. Other Perigondwanan Early Carboniferous
palynological records
Melo and Loboziak (2000) described Visean assem-
blages from the Parnaiba Basin, Brazil, characterized,
among other species, by the presence of Bascaudaspora
submarginata, Apiculiretusispora multiseta, Colatispor-
ites decorus, Grandispora spiculifera, Indotriradites
daemonii, I. dolianitii, Vallatisporites splendens and
Verrucosisporites nitidus which are shared with the
Colombian associations.
Recently, Melo and Loboziak (2003) revised the
miospore biostratigraphy of the DevonianEarly Car-
boniferous interval from the Amazon Basin, northern
Brazil. They proposed 17 interval zones based on se-
lected Euramerican and Western Gondwanan species,
which were correlated with zones of Western Europe
and the Old Red Sandstone Continent. The Tournaisian
section in the Amazon Basin corresponds principally to
40 H. Duenas, S.N. Cesari / Review of Palaeobotany and Palynology 138 (2006) 3142
the Oriximina Formation from which three miospore
assemblages were established. The highest Tournaisian
(Interval Zone PD) assemblage is defined by the ap-
pearance of Spelaeotriletes pretiosus and Colatisporites
decorus accompanied by rare specimens of Raistrickia
clavata, which characterizes the PC Biozone in Britain.
The Visean interval (Interval Zone Mag) in the
Amazon Basin corresponds principally to the Faro
Formation. It is characterized by the incoming of Cor-
dylosporites magnidictyus and the occurrence of long-
ranging species: Auroraspora macra, Colatisporites
decorus, Grandispora spiculifera, Verrucosisporites
nitidus among others. Auroraspora solisorta, Schop-
fites claviger and Indotriradites dolianitii Morphon
appear restricted to the Mag Zone in the Amazon
Basin. Prolycospora rugulosa, a species that continues
into the Late Carboniferous appears in this zone. De-
spite the absence of C. magnidictyus in the Colombian
palynofloras, numerous species are shared with this
Brazilian interval.
Recently, Azcuy and di Pasquo (2005) described an
Early Carboniferous palynoflora from the Ambo For-
mation, Peru. The Peruvian association is referred to the
Late Visean Cordylosporites magnidictyus Palynozone,
defined in the Amazon Basin, Brazil. The common
species shared with the Colombian assemblages are:
Auroraspora macra, Bascaudaspora submarginata,
Colatisporites decorus, Retusotriletes crassus, Retuso-
triletes incohatus, Schopfites claviger and Spelaeotri-
letes pretiosus. However, the absence of C.
magnidictyus, Anapiculatisporites austrinus, Dibolis-
porites microspicatus, Schopfipollenites ellipsoides
among others, in Colombia prevents a closer correlation.
Clayton et al. (2000) described latest Devonian and
Early Carboniferous assemblages from two wells in
Saudi Arabia. They recognized three assemblages,
which were, correlated with the RT (Prolycospora
rugulosaSpelaeotriletes triangulus) Biozone, original-
ly defined in Libya (Loboziak and Clayton, 1988): the
Indotriradites explanatus Assemblage, the Verruciretu-
sispora famenensis Assemblage and Retispora lepido-
phyta Assemblage. The Indotriradites explanatus
Assemblage was considered to be Tournaisian in age
and would be similar in composition to bPalynozone
12Q of Grignani et al. (1992) described in Libya. The
palynofloras from the RT Biozone are poorly preserved
and include Aratrisporites saharaensis, Prolycospora
rugulosa, Spelaeotriletes arenaceous among others.
Clayton et al. (2002) pointed out the strong differences
in miospore assemblages during Visean times between
North Africa and Western Europe. They characterized
the Visean in Algeria, Saudi Arabia, Libya and Syria
(Ravn et al., 1994) by the presence of A. saharensis
Loboziak, Clayton and Owens, Radiizonates arcuatus
Loboziak Playford and Melo, Spelaeotriletes owensii
Loboziak and Alpern and Vallatisporites agadesi Lobo-
ziak and Alpern. In the Colombian palynofloras none of
these species is registered, although they occur in the
Amazon Basin of Brazil.
Coquel et al. (1995) described palynological assem-
blages from northern Niger that show close resem-
blance to those from the ViseanNamurian of the
Saharan Platform. They proposed that during the
ViseanNamurian, the North African margin constitut-
ed a well-defined microfloristic province, clearly dis-
tinguished from Laurasia and Australia.
A southern record in Africa was recently described
by Atta-Peters and Anan-Yorke (2003) for Ghana.
They described a Tournaisian assemblage from the
Takoradi Shale Formation characterized by the pres-
ence of Auroraspora macra, Cordylosporites marciae,
Spelaeotriletes balteatus, Vallatisporites vallatus, and
Verrucosisporites nitidus among other species.
7. Conclusions
This report increases the knowledge about the dis-
tribution and composition of Early Carboniferous pal-
ynological assemblages in the Perigondwanic area. The
palynological assemblages provided the only evidence
for Early Carboniferous strata in the Colombian Llanos
Orientales Basin and constituted a new record for
northern South America.
The dominant species in the Llanos Basin consist of
morphologically simple, smooth (Retusotriletes spp.) or
apiculate (Apiculiretusispora) retusoids as well as
forms showing zonate structures and representatives
of the densospore group (Densosporites, Indotriradites,
Vallatisporites). In spite of the absence of some zonal
data like Cordylosporites magnidictyus or Aratrispor-
ites saharensis, a probable regional correlation could be
proposed with the Brazilian PD and Mag Zones. This
later zone, as well as our assemblages, contains numer-
ous long-ranging species persisting from the Strunian.
A predominance of Euramerican species together with
some Gondwanan species like Indotriradites dolianiti
morphon is recognized in both regions. A probable Late
TournaisianVisean age is proposed for the studied
interval of the SM-4 well.
Acknowledgements
The authors are indebted to Dr. Geoffrey Playford by
his valuable comments about some specific identifica-
 H. Duenas, S.N. Cesari / Review of Palaeobotany and Palynology 138 (2006) 3142
tions and to Dr. B. Owens, Dr. E. Turnau and Dr. J.
Utting by their suggestions as reviewers. This is a
contribution to the IGCP Project 471.
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