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The final phase of tropical lowland conditions

in the axial zone of the Eastern Cordillera of
Colombia: Evidence from three palynological
records
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Journal of South American Earth Sciences 39 (2012) 157e169

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journal homepage: www.elsevier.com/locate/jsames

The nal phase of tropical lowland conditions in the axial zone of the Eastern
Cordillera of Colombia: Evidence from three palynological records
D. Ochoa a, b, *, C. Hoorn c, C. Jaramillo a, G. Bayona d, M. Parra e, F. De la Parra f
a

Smithsonian Tropical Research Institute, P.O. Box 0843, Balboa, 03092 Ancon, Panama
Department of Biological Sciences, East Tennessee State University e ETSU, Johnson City, TN 37614-14850, USA
c
Paleoecology and Landscape Ecology, Institute for Biodiversity and Ecosystem Dynamics (IBED), University of Amsterdam, Science Park 904, 1098 XH Amsterdam, Netherlands
d
Corporacin Geolgica ARES, Calle 44A #53-96, Bogot, Colombia
e
Department of Geological Sciences, Jackson School of Geosciences, University of Texas at Austin, Austin, TX 78712, USA
f
Instituto Colombiano del Petrleo - Ecopetrol, Km 7 va Piedecuesta-Bucaramanga, Colombia
b

a r t i c l e i n f o

a b s t r a c t

Article history:
Received 10 May 2011
Accepted 2 April 2012

Deformation of the Eastern Cordillera, as a double-verging thrust belt that separates the Magdalena
Valley from the Llanos Basin, is a dening moment in the history of the northern Andes in South America.
Here we examine the age and depositional setting of the youngest stratigraphic unit in three sectors of
the Eastern Cordillera: (i) the Santa Teresa Formation (western ank), (ii) the Usme Formation (southern
central axis), and (iii) the Concentracin Formation (northeastern central axis). These units were
deposited prior to the main Neogene deformation events. They represent the last preserved record of
lowland conditions in the Eastern Cordillera, and they are coeval with a thick syn-orogenic deposition
reported in the Llanos Basin and Magdalena Valley. Based on palynological data, we conclude that the
upper Usme Formation was deposited during the Bartonian-earliest Rupelian? (Late Eocene-earliest
Oligocene?); the Concentracin Formation was deposited during the Late Lutetian-Early Rupelian
(Middle Eocene to Early Oligocene), and the upper Santa Teresa Formation was accumulated during the
Burdigalian (Early Miocene). These ages, together with considerations on maximum post-depositional
burial, provide important time differences for the age of initial uplift and exhumation along the axial
zone and western foothills of the Eastern Cordillera. The switch from sediment accumulation to erosion
in the southern axial zone of the Eastern Cordillera occurred during the Rupelian-Early Chattian
(Oligocene, ca 30 to ca 26 Ma), and in the northeastern axial zone occurred prior to the latest ChattianAquitanian (latest Oligocene-Early Miocene ca 23 Ma). In contrast, in the western ank, the switch
occurred during the Tortonian (Late Miocene, ca 10 Ma). In addition, we detected a marine transgression
affecting the Usme and Concentracin formations during the Late Eocene; coeval marine transgression
has been also documented in the Central Llanos Foothills and Llanos Basin, as evidenced by the similarity
in oras, but not in the western foothills. Our dataset supports previous sedimentological, geochemical
and thermochronological works, which indicated that (i) deformation in the Eastern Cordillera was
a diachronous process, (ii) the sedimentation along the axial zone stopped rst in the south and then in
the north during the Oligocene, (iii) depositional systems of the axial zone and central Llanos Foothills
kept partly connected at least until the Late Eocene, and (iv) Miocene strata were only recorded in
adjacent foothills as well as the Magdalena and Llanos basins.
2012 Elsevier Ltd. All rights reserved.

Keywords:
Eastern Cordillera
Santa Teresa Formation
Usme Formation
Concentracin Formation
Andean orogeny
Late Eocene-Early Miocene

1. Introduction

* Corresponding author. Smithsonian Tropical Research Institute, P.O. Box 0843,

Balboa, 03092 Ancon, Panama.
E-mail addresses: dianita.ochoa@gmail.com, ochoalozano@goldmail.etsu.edu
(D. Ochoa), carina.hoorn@milne.cc (C. Hoorn), jaramilloc@si.edu (C. Jaramillo),
gbayona@cgares.org (G. Bayona), mparra@cgares.org (M. Parra), felipe.delaparra@
ecopetrol.com.co (F. De la Parra).
0895-9811/$ e see front matter 2012 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jsames.2012.04.010

The northern Andes Mountains are the result of a complex

interaction between the continental South American plate and the
oceanic Caribbean and Nazca plates (Fig. 1). One consequence of this
multistage orogeny was the asynchronous building of three
different mountain belts -the Western, Central, and Eastern
Cordilleras- throughout the Mesozoic and Cenozoic (Barrero, 1979;
Etayo et al., 1983; Villamil, 1999). Deformation of the Eastern

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D. Ochoa et al. / Journal of South American Earth Sciences 39 (2012) 157e169

Fig. 1. Geographical location of studied sections. Modied after Pardo-Trujillo (2004). CC Central Cordillera, WC Western Cordillera, UMV Upper Magdalena Valley,
SNSM Sierra Nevada de Santa Marta, SNCo Sierra Nevada del Cocuy, CatB Catatumbo Basin.

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D. Ochoa et al. / Journal of South American Earth Sciences 39 (2012) 157e169

Cordillera (EC), as a double-verging orogen with intermontane

basins along the axial zone (Cooper et al., 1995; Corts et al., 2006),
was of particular signicance in the geological history of northern
South America as it modied river drainages (Guerrero, 1997; Hoorn
et al., 1995, 2010), promoted vertical surface uplift in the last
3e5 m.y. (Gregory-Wodzicki, 2000; Mora et al., 2008b), and created
new habitats, such as pramos and cloud forests in the axial zone of
the EC (Hooghiemstra, 1984, 1988; Van der Hammen et al., 1973).
Tectono-sedimentological, geodynamic and thermochronological
studies have shown that the Eastern Cordillera in the Colombian
Andes was already tectonically active during the Early Paleocene
(Bayona et al., 2008; Parra et al., 2012) and the Late Eocene-Early
Oligocene (Bande et al., 2012; Bayona et al., 2008; Gmez et al.,
2005, 2003; Horton et al., 2010b; Mora et al., 2010a; Nie et al.,
2010; Parra et al., 2009a, 2009b; Saylor et al., 2011; Toro et al.,
2004). During the Late Neogene, the tectonic activity in the
Eastern Andes rapidly increased (Duque-Caro, 1990; Helmens and
Van der Hammen, 1994; Hooghiemstra and Van der Hammen,
1998; Hoorn et al., 1987; Mora et al., 2010b; Shephard et al., 2010;
Taboada et al., 2000), generating a major exhumation pulse of the EC
in the last w7 m.y. (Bayona et al., 2008; Corts et al., 2006; Mora
et al., 2010b). These pulses led to changes in regional climate
(Ehlers and Poulsen, 2009; Insel et al., 2009; Sepulchre et al., 2009)
and the sedimentary regimes in the adjacent lowlands which
resulted in the establishment and evolution of the Amazon River
(Figueiredo et al., 2009, 2010; Hoorn, 1994; Hoorn et al., 1995, 2010;
Hoorn and Wesselingh, 2010; Shephard et al., 2010) and increased
biodiversity in western Amazonia (Hoorn et al., 2010).
Both the onset of double-verging deformation of the EC and
consequent isolation of the Magdalena River Valley from the Llanos
region are critical to the development of the northern Andes. In
order to understand the nal phase of the sedimentation within the
EC, we have dated the youngest sediments preserved in three
different synclines located (1) along the western ank of the EC
(Santa Teresa Formation in the Guaduas Syncline), (2) in the
southern axial zone of the EC (Usme Formation in the Usme
Syncline, Bogot High Plain), and (3) in the northeastern axial zone
of the EC (Concentracin Formation in the Floresta Syncline, footwall of the Soapaga Fault) (Fig. 1). In addition, we have provided
temporal estimates of the switch from burial to exhumation
through the assessment of the magnitude of post-depositional
burial based on recently published paleothermal and thermochronometric data. Finally, we have reviewed several depositional
and kinematic models to evaluate the degree of connectivity
between the basins during the Late Eocene to Early Miocene.
2. Lithostratigraphic setting
The Late Eocene to Miocene evolution of the EC is recorded in
three different large synclines located across the EC (Table 1). The
youngest sediments, which are preserved at the core of each
syncline, also represent the youngest sediments preserved in the
EC, other than the Pliocene lake and alluvial sediments from the
Bogota High plain (Helmens, 1990; Helmens and Van der Hammen,
1994; Hooghiemstra and Van der Hammen, 1998; Hoorn et al.,
1987). All stratigraphic sections were measured near the area
where the type section of each formation was proposed (De Porta,
1974). In the western ank, we studied the youngest sediments of
the Guaduas Syncline, represented by the Santa Teresa Formation
(De Porta, 1966; De Porta, 1974). In the southern axial zone, we
examined the youngest strata of the Usme Syncline, represented by
the Usme Formation (De Porta, 1974; Hoorn et al., 1987). Lastly, in
the northeastern axial zone, we studied the youngest strata of the
Floresta Syncline, represented by the Concentracin Formation
(Rodrguez and Solano, 2000; Ulloa et al., 2001) (Fig. 1).

159

Table 1
Location of selected sedimentary sections.
Formation

Syncline Regional location

Coordinates

Samples

Latitude Longitude Analyzed

Santa Teresa
Guaduas Middle Magdalena 4.86 N
Fm.
Valley
Usme Fm.
Usme
Bogot High Plain
4.52 N
Concentracin Floresta Paz del Ro, Boyac 6.03 N
Fm.

74.63 W

74.15 W
72.76 W

16
16

The fossil-rich Santa Teresa Formation was dened by De Porta

(1966) and originally thought to be the Early-Middle Miocene La
Cira Formation (Raasveldt and Carvajal, 1957; Van der Hammen,
1958), which outcrops in the northern Middle Magdalena Valley
Basin. However, De Porta (1966) suggested a different nomination,
assigning the Santa Teresa name for the rocks outcropping in the
area of the Guaduas Syncline. The formation conformably overlies
the San Juan de Ro Seco Formation and is w475 m thick along the
Bogot-Cambao road. It has a lithofacies distinctive of quiet
lagoonal settings (De Porta, 1966), which is partially conrmed by
some forms of sapropelic organic matter (alginite and liptodetrinite), reported in the upper half of the unit (Gmez, 2001). Acosta
and Ulloa (2001) report lithofacies representative of shallow,
freshwater environments with channels lled with conglomeratic
and sandy lithologies, presence of several coal beds, abundant leaf
impressions, sh bones, gastropods and bivalves along the Bal
Creek in Cundinamarca. Pilsbry and Olsson (1936), in contrast,
suggested occasional presence of salty waters based on some
horizons with gastropods and mollusks favoring brackish waters
(Corbula). The age of the formation is uncertain although Oligocene
(De Porta and De Porta, 1962), Late Oligocene (Acosta and Ulloa,
2001), Oligocene-Early Miocene? (De Porta, 1966), and Early
Miocene ages (Nuttall, 1990) have been suggested based on malacological and palynological data. The formation broadly correlates
with the Colorado and La Cira formations in the Middle Magdalena
Valley Basin (De Porta, 1974; Van der Hammen, 1958).
The Usme Formation was proposed by Hubach (1957) and
Julivert (1963), it overlies the Regadera Formation. The type of
contact is variable, from unconformable in the eastern ank of the
Usme Syncline (Hoorn et al., 1987; Julivert, 1963) to conformable in
the western ank (Julivert, 1963; Montoya and Reyes, 2005). The
Usme Formation is unconformably overlain by Late Miocene? Pliocene alluvial, gravity-ow material and lacustrine deposits,
corresponding to the Marichuela Formation (De Porta, 1974; De
Porta, 2003; Helmens, 1990; Helmens and Van der Hammen,
1994; Hoorn et al., 1987; Roddaz et al., 2009; Toro et al., 2003).
The Usme Formation is w365 m thick and has been traditionally
subdivided into two members (Hoorn et al., 1987; Julivert, 1963).
The lower member consists dominantly of dark brown and graycolored claystones, siltstones, and shales. Some dark gray siltstones beds are intercalated with thin coal layers and abundant
plant remains. A general upward-coarsening trend in grain size is
evident in the lower member. Towards the upper part of this
member, some very ne to ne quartzitic sandstones with crossstratication are interbedded with bioturbated shales and siltstones levels (Hoorn et al., 1987; Montoya and Reyes, 2005). The
lower member has been interpreted as coastal plain deposits
incised by subtidal channels (Hoorn et al., 1987). The upper
member of the Usme Formation is dominated by multicolored
siltstones and shales, intercalated with yellowish massive to crossbedded sandstones varying from coarse sand to conglomeratic
lithologies (Hoorn et al., 1987). Towards the top of the section,
several coal and lignite layers with abundant plant material are
found. The upper member is interpreted as the product of deltaic to
intertidal settings, with sand bars, and interdigitated and

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abandoned channels (Hoorn et al., 1987). The formation has been

dated by palynological analysis as Late Eocene to Early Oligocene
(Hoorn et al., 1987) or Middle Eocene to Middle Oligocene (Van der
Hammen, 1957); whereas as Middle to Late Oligocene based on the
presence of Globorotalia fohsi andina (Brgl, 1955). The Usme
Formation has been correlated with the Concentracin Formation
in the Floresta Basin (De Porta, 1974; Van der Hammen and Parada,
1958; Villamil, 1999) and the Upper Mirador Formation and Lower
Carbonera C8 member in the Foothills and Llanos basins (Cooper
et al., 1995; Pulham et al., 1997). The Upper Mirador consists of
dark shales interbedded with ne to coarse sandstones, and the
Lower Carbonera is composed of tabular sandstone interbedded
with dark gray mudstone, and aser and lenticular lamination (De
Porta, 1974; Parra et al., 2009a). These lateral differences in
lithology have been interpreted as facies changes between uvial
and marine-inuenced conditions (Cooper et al., 1995; Pulham
et al., 1997; Roddaz et al., 2009; Saylor et al., 2011).
The Concentracin Formation was proposed by Alvarado and
Sarmiento (1944) and conformably overlies the Picacho Formation (Rodrguez and Solano, 2000). It is w1 km thick and composed
of an upward-coarsening sequence of laminated mudstones interbedded with sandstones (Saylor et al., 2011). Oolitic ironstones,
plant fragments and bioturbation are common throughout the
section (Saylor et al., 2011). The sedimentary sequence is interpreted as having formed in a coastal plain environment with
lagoonal to partially closed estuarine conditions (Saylor et al., 2011;
Villamil, 1999). Oolitic ironstone up to 3 m thick, which is
commercially mined, occurs towards the base of the unit
(Kimberley, 1980; Saylor et al., 2011; Van Houten, 1967). Villamil
(1999) interpreted these oolitic layers as evidence of a Late
Eocene regional seaway that ooded northern South America.
Based on palynological data, the formation has been dated as
Middle Eocene to Late Oligocene (Hubach, 1957; Van der Hammen,
1957). Deposition of the Concentracin Formation is considered
coeval with the active stage of the Soapaga Fault system, which
affected only the northern axial zone of the cordillera (Saylor et al.,
2011). The unit correlates with the Usme Formation in the Bogot
High Plain area (De Porta, 1974; Van der Hammen and Parada,
1958), with the Carbonera and Len formations in the Catatumbo
Basin (De Porta, 1974; Van der Hammen, 1958), and with the Upper
Mirador and Lower Carbonera formations in the Foothills and Llanos basins (Cazier et al., 1995; Cooper et al., 1995; Santos et al.,
2008; Villamil, 1999) (Fig. 1).
3. Methods
A total of 41 samples from the three formations (Table 1) were
processed for palynological analysis following standard procedures
(Traverse, 2007). All samples were digested by using 10% HCl and
40% HF. Each sample was sieved using 10 mm and 100 mm meshes;
nally, permanent montages were prepared. Light microscopy was
used to examine the palynological content and at least 100 grains
were counted per slide. Morphological features were compared
with descriptions and illustrations from various resources (Dueas,
1980; Germeraad et al., 1968; Gonzalez, 1967; Hoorn et al., 1987;
Jaramillo et al., 2010; Jaramillo et al., 2007; Jaramillo and Dilcher,
2001; Jaramillo et al., 2011; Leidelmeyer, 1966; Lorente, 1986;
Muller et al., 1987; Van der Hammen, 1956; Van der Hammen and
Garca de Mutis, 1966) and taxonomical nomenclature followed
Jaramillo and Dilcher (2001).
The sections were dated using a maximum likelihood estimation based on the palynological zonation proposed by Jaramillo
et al. (2011), which has been time-calibrated using carbon
isotopes, radiometric dating, foraminifera and magneticstratigraphy data (Fig. 2). Maximum likelihood analysis follows

the procedure of Punyasena et al. (2012) (Appendix 1). Additionally,

a Marine Inuence Index (MI) was calculated using the ratio
proposed by Santos et al. (2008), in order to estimate the relation
between marine and terrestrial palynomorphs. The index was
estimated for each sample as MI M/T, where M is the total number
of marine palynomorphs and T is the total number of palynomorphs counted per sample (Rull, 2002; Santos et al., 2008). All
analyses were conducted using standard libraries from the program
R (R Development Core Team, 2009), which is a free-software,
developed by several contributors and administered by the R
Foundation for Statistical Computing (http://www.R-project.org).
In addition, the Stratigraph package was used to produce the
palynostratigraphic range charts (Green et al., 2010).
4. Palynological results
4.1. Santa Teresa Formation
Palynological matter was recovered only from the upper 266 m
of the Santa Teresa Formation. The palynoora includes a large
abundance of ferns spores (e.g. Psilatriletes and Laevigatosporites)
and palms (Mauritiidites). Fungal remains are common throughout
the entire section, also a 78.5 m thick interval with abundant
Pediastrum and Botryococcus algae was observed (from 549.74 m to
628.24 m) (Fig. 3). Neither mangrove elements nor marine palynomorphs were observed (Fig. 3). The most abundant palynomorphs included Mauritiidites franciscoi, Perisyncolporites
pokornyi, Magnaperiporites spinosus, Rhoipites guianensis, Psilamonocolpites, Laevigatosporites, Polypodiisporites and Psilatriletes
groups (Appendices 2 and 3).
The co-occurrence of Magnastriatites grandiosus (First Appearance Datum [FAD] at 451.2 m), M. spinosus (FAD at 456.79 m),
Concavissimisporites fossulatus (FAD at 470.7 m), and Bombacacidites muinaneorum (FAD at 468.71 m) (Fig. 4), as well as the
overall abundance of other key taxa, such as Bombacacidites brevis,
Echiperiporites akanthos, Mauritiidites franciscoi, Perisyncolporites
pokornyi, Ranunculacidites operculatus, Retitrescolpites? irregularis,
Spirosyncolpites spiralis, Striatriletes saccolomoides, Tetracolporopollenites maculosus and Tetracolporopollenites transversalis, indicates that this palynoora corresponds to the palynological zones
T-12 and T-13. These two zones have been dated as 16.1e23 Ma,
corresponding to the Aquitanian-Burdigalian, Miocene (Jaramillo
et al., 2011). This interpretation is supported by maximum likelihood analysis (Fig. 5), which indicates a high probability of correlation with the upper part of palynological zone T-12 Horniella
lunarensis and zone T-13 Echitricolporites maristellae (Burdigalian,
19e16 Ma).
4.2. Usme Formation
Good pollen recovery was found for most of the section, except
for the lower 116 m and the uppermost 60 m, where no pollen was
found. The palynoora is characterized by high abundances of fern
spores (e.g. Polypodiisporites and Laevigatosporites) and palms (e.g.
Mauritiidites) (Appendices 4 and 5). Four levels contain dinoagellate cysts and acritarchs (2140 m, 2141 m, 2152 m, and 2258 m).
Levels located towards the base of the upper member exhibit the
highest Marine Inuence Index values (at 2140 m, 2141 m and
2152 m, MI 0.05, 0.09 and 0.029, respectively), whereas marine
inuence was minimal at the top of the section, at 2258 m
(MI 0.012) (Fig. 3). Colonies of Pediastrum sp. were common
between 2140 m and 2142.6 m, and high frequencies of mangrove
elements (Rhizophoraceae and Pellicieraceae) occurred between
2258 m and 2299 m (mean 5.6%) (Fig. 3).

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161

Fig. 2. Palynological zonation proposed for the Cenozoic of the Llanos and Llanos Foothills by Jaramillo et al. (2011) and Muller et al. (1987). Geologic time scale after Gradstein et al.
(2004).

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D. Ochoa et al. / Journal of South American Earth Sciences 39 (2012) 157e169

Fig. 3. Relative abundances of terrestrial plants, marine and fresh waters elements present in each unit. Marine Inuence (MI) Index values ranges from 0 (fully terrestrial) to 1
(fully marine). Palynological zones after Jaramillo et al. (2011). Vertical scales vary in each section.

The most common angiosperm taxa recovered include M. franciscoi, Echitriporites trianguliformis orbicularis, Longapertites proxapertitoides var. proxapertitoides, P. pokornyi, R. guianensis, T.
maculosus, T. transversalis, and the Psilamonocolpites and Retitricolporites groups.
Cicatricosisporites dorogensis and the Laevigatosporites, Polypodiisporites, and Psilatriletes groups dominated pteridophyte
spores composition. Other important taxa recovered include Arecipites regio, Cricotriporites guianensis, E. akanthos, Echitetracolpites?
tenuiexinatus, Foveotriporites hammenii, Lanagiopollis crassa,
Monoporopollenites annulatus, Poloretitricolpites absolutus, Spirosyncolpites spiralis, Syncolporites marginatus and Zonocostites
ramonae (Appendices 4 and 5). Echitriporites trianguliformis orbicularis occurs throughout the section, except for the uppermost
sample, at 2354 m. A single occurrence of C. fossulatus was also
found at 2254 m (Fig. 4).
The frequent occurrence of Echitriporites trianguliformis orbicularis, along with the Last Appearance Datum [LAD] of A. regio (at
2254 m), S. marginatus (at 2294 m) and Echitetracolpites? tenuiexinatus (at 2299 m), the FAD of E.chiperiporites akanthos (at 2254 m),
and the presence of P. absolutus (single occurrence at 2267 m)
(Fig. 4) indicates that up to meter 2299 the palynoora belongs to
palynological zone T-07 Echitriporites trianguliformis orbicularis,

dated as Late Eocene (38e33.9 Ma) (Jaramillo et al., 2009, 2011)

(Fig. 2). This dating is also supported by the maximum likelihood
analysis, which indicates a high probability of correlation with zone
T-07 Echitriporites trianguliformis orbicularis for the entire section,
except for the last sample (Fig. 5).
Only one sample with poor recovery (2354 m) was obtained
from the uppermost 60 m of section. Among other taxa, F. hammenii
was recovered in this sample. The LAD of F. hammenii is considered
as an important biostratigraphic event and was estimated at
33.24 Ma (Jaramillo et al., 2011). Based on the presence of this
taxon, we consider the upper sediments to be no younger than
early Oligocene (earliest Rupelian, ca 33 Ma). In consequence, we
suggest a Bartonian-earliest Rupelian? age (ca 38 to 33 Ma) for the
upper-lower to upper Usme Formation.
4.3. Concentracin Formation
Good pollen recovery was obtained in almost the entire
sequence except for the upper part of the section, where palynological recovery was poor (1211 me1440 m) (Appendices 6 and 7).
The palynooral assemblage is dominated by fern spores (C. dorogensis, Laevigatosporites, M. grandiosus, and Polypodiisporites) and
angiosperms including M. franciscoi, Psilamonocolpites medius, T.

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163

Fig. 4. Palynological zones established for each section. Arrows indicated the occurrence of key taxa used to dene the palynological zones. Note Santa Teresa Formation includes
from the upper part of zone T-12 to zone T-13, Usme Formation only includes zone T-07 and Concentracin Formation includes from zone T-06 to T-09. Vertical scales vary in each
section.

maculosus, T. transversalis, and L. crassa. Three peaks of mangrove

abundance (L. crassa, a Pellicieraceae) occur throughout the section
at 40.49 m (23.7% of the palynoora), 559.99 m (11.5%), and
between 774.49 m and 832 m (47%) (Fig. 3). Three intervals with
increased abundances of freshwater algae were found at 40.49 m
(0.7%), from 360 m to 391.49 m (0.6%) and from 715.49 m to
751.49 m (1.5%). Four intervals displaying high MI values were
found at 40.49 m (MI 0.13), at 560 m (MI 0.019), from 751.49 m
to 774.5 m (MI 0.02), and from 797.5 m to 832.49 m (MI 0.15)
(Fig. 3).
The palynoora indicates the presence of four palynological
zones as follows: zone T-06 Spinizonocolpites grandis from the base
to 116.9 m; zone T-07 Echitriporites trianguliformis orbicularis from
116.9 m to 735.5 m; zones T-08 Nothofagidites huertasii to T-09
Foveotricolporites etayoi from 735.5 m to 1210.5 m.
Palynological zone T-06 is identied by the LAD of S. grandis (at
116.9 m) (Fig. 4). In addition, it is supported by the co-occurrence of
C. dorogensis, Laevigatosporites catanejensis and R. guianensis, all of
which have their FADs at 40.49 m. Palynological zone T-07 is indicated by the LADs of Echitriporites trianguliformis orbicularis (at
735.5 m), Proxapertites magnus (at 391.4 m), and Racemonocolpites
facilis (at 371.9 m), and by FADs of Striatriletes saccolomoides (at
559.9 m) and Polypodiisporites usmensis (at 391.49 m) (Fig. 4). The
occurrence of Echitetracolpites? tenuiexinatus (at 334.49 m) also
supports this zone. Finally, zones T-08 N. huertasii to T-09 F etayoi are
noted by the FAD of M. grandiosus (at 1101.0 m), and the LAD of
Spinizonocolpites echinatus (at 797.49 m) (Fig. 4). Zone T-09 is also
supported by high abundances of C. dorogensis, occurring towards
the top of the section (see Appendices 6 and 7). These four palynological zones, also supported by the maximum likelihood analysis
(Fig. 5), indicate a Lutetian-Early Rupelian (Middle Eocene-Early
Oligocene, ca 39 to 31.5 Ma) age for the Concentracin Formation.

5. Discussion of palynological data

5.1. Age
We found that the upper sediments of the Santa Teresa
Formation (corresponding to the uppermost 266 m) have a high
probability of belonging to the upper part of palynological zone T12 H. lunarensis and zone T-13 E. maristellae (Burdigalian, Early
Miocene, 19e16 Ma) (Figs. 4 and 5). Earlier stratigraphic, malacological and palynological studies considered the formation as
having accumulated during the Oligocene (De Porta and De Porta,
1962), Late Oligocene (Acosta and Ulloa, 2001), Oligocene to Early
Miocene? (De Porta, 1966) or Early Miocene (Nuttall, 1990).
De Porta and De Porta (1962) studied the palynological associations within the section and reported occurrences of Polypodiisporites usmensis, Psilamonocolpites, Mauritiidites and
Psilatriletes groups, and by using the relative abundance of each
group, they correlated the section with the Oligocene zones
proposed by Van der Hammen (1958). Van der Hammens zones are
based on abundance peaks of broad categories that represent
modications in the vegetation due to regional climatic changes.
Thus, each abundance cycle had a certain chronostratigraphic value
according to the climate variation. This methodology has been
described as inappropriate for biostratigraphic purposes (Jaramillo
et al., 2011; Moore et al., 1991), because it depends on factors such
as environmental settings during deposition, type of sediments
sampled, and it is also greatly affected by the closed sum mathematical effect (Kovach and Batten, 1994; Moore et al., 1991).
Furthermore, Van der Hammens approach did not use an external
age-dataset for calibration.
De Porta (1966) suggested an Oligocene to Early Miocene? age.
He established the Oligocene age by using the stratigraphic position

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D. Ochoa et al. / Journal of South American Earth Sciences 39 (2012) 157e169

Fig. 5. Stratigraphic age estimates per section calculated using the probabilistic
Maximum Likelihood approach based on the taxa abundance. Normalized likelihood
values are represented by color with higher and lower likelihood values symbolized by
green and blue, respectively. Inferior axis showing geologic time (Ma), vertical axis
representing depth and superior axis marking the palynological zones proposed by
Jaramillo et al. (2011). Vertical scales vary in each section. Samples are represented by
yellow bars.

of the section and previous palynological data as discussed above

(De Porta and De Porta, 1962), whereas the Early Miocene? age was
based on the mollusk association Anodontites laciranus, Hemisinus
waringi and Diplodon opocitonis described by Anderson (1929).

Likewise, Nuttall (1990) assigned an Early Miocene age after

reviewing the malacological material collected and described by
Anderson (1929). Although both studies assigned an Early Miocene
age to the fauna, they also discussed extensively the uncertainty of
the exact stratigraphic position of this association within the Guaduas Syncline. Furthermore, Nuttall (1990) pointed out that there is
no precise information indicating that the mollusk fauna actually
belongs to any level from the Guaduas structure. He also argued that
Butler (1939) tentatively located the fauna close to the core of the
Guaduas Syncline, and as a result, the mollusk assemblage has been
placed at the upper part of the Santa Teresa Formation. In addition to
the stratigraphic uncertainty, De Porta (1966) signaled that the
correlation with the Miocene La Cira Formation based on the
mollusk fauna was biased by the high frequency of endemic species
within the Santa Teresa Formation. Consequently, De Porta (1966)
and Nuttall (1990) concluded that although the fauna probably
could suggest an Early Miocene age, the certainty of the dating was
very low. The new palynological data presented herein supports
a Burdigalian, Miocene (19e16 Ma) age for the upper sediments
from the Santa Teresa Formation (Figs. 4 and 5). However, the lower
part of the Santa Teresa still needs to be dated.
The palynooral assemblage recovered from upper part of the
lower Usme Formation and lower part of the upper Usme Formation corresponds to palynological zone T-07 Echitriporites trianguliformis orbicularis of Jaramillo et al. (2011) (Figs. 4 and 5), which is
dated as Late Bartonian-Priabonian, Eocene (38e33.9 Ma) (Fig. 2).
The uppermost 60 m of the section did not provide conclusive
evidence supporting zone T-07, and it contains F hammenii, a taxon
whose extinction is estimated at 33.24 Ma (Jaramillo et al., 2011).
Therefore we consider the upper-lower and upper Usme Formation
as Bartonian to earliest? Rupelian (Late Eocene to earliest? Oligocene, 38e33 Ma) in age.
Van der Hammen (1957) dated the Usme section as Middle
Eocene to Middle Oligocene, based on the stratigraphic position of
the unit and occurrence peaks of Striatriletes susannae, a junior
synonym of C. dorogensis (Potonie and Johann, 1933). As discussed
before, age dating based on vegetation cycles may be biased
because it may signal ecological settings rather than chronological
events. Hoorn et al. (1987), following the palynological zonation
proposed by Muller et al. (1987), designated a Late Eocene-Early
Oligocene age by correlation with the Echiperiporites estelae and
Magnastriatites-Cicatricosisporites Zones (see Fig. 2). The latter zone
was supported by the co-occurrence of M. grandiosus and C dorogensis in the uppermost part of the Usme Formation. We carefully
reanalyzed the same palynological slides examined by Hoorn et al.
(1987) but never detected the presence of M. grandiosus. Moreover,
we did not nd M. grandiosus in additional samples from the rest of
the section. It is possible that large specimens of C. dorogensis
sometimes can be confused with small specimens of M. grandiosus,
and this could be the case in Hoorn et al.s (1987) work.
The palynoora recovered from the Concentracin Formation
ranges from the upper T-06 S. grandis to T-09 F. etayoi of Jaramillo
et al. (2011) (Figs. 4 and 5). Thus, we interpret the lower 1210 m
of the section as having accumulated during the Late Lutetian to
Early Rupelian (latest Middle Eocene to early Early Oligocene, 39 to
31.5 Ma). Because we did not recover pollen from the upper
w240 m, we were unable to directly establish the depositional age
for this segment. Nevertheless, we estimate the approximate age at
the top of the formation as Rupelian (ca 31.5 Ma), by assuming
constant sedimentation rates in the upper 240 m, and by using the
FAD of M. grandiosus within the section (1101.0 m) as age reference
(ca 33.67 Ma, according to Jaramillo et al., 2011).
Van der Hammen (1957) dated the Concentracin Formation as
extending from the Middle Eocene to the earliest Late Oligocene
based on vegetational abundance peaks. As in the case of the Usme

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D. Ochoa et al. / Journal of South American Earth Sciences 39 (2012) 157e169

Formation, he based the Oligocene age mainly on high occurrences

of Striatriletes susannae, a junior synonym of C. dorogensis (Potonie
and Johann, 1933). Van der Hammen conrmed this age by using
a coal seam from the Usme Formation, which was previously dated
as Middle Oligocene also on the basis of a peak in C. dorogensis (Van
der Hammen, 1957). Given that non-independent data were used to
constrain the chronostratigraphic frame in both the Usme and
Concentracin formations, the dating may have relied on circular
reasoning. Nevertheless, the age proposed by Van der Hammen is
similar to the age proposed here.
The dating established herein implies a revision of the traditional chronostratigraphic correlations for the Eastern Cordillera. In
particular, the Usme and Concentracin formations have generally
been considered as coeval units (Gmez et al., 2005; Van der
Hammen, 1957, 1958), deposited during the Middle EoceneMiddle Oligocene. As shown here, the upper boundaries of the
two formations are not coeval (see Figs. 4 and 5). The uppermost
part of the Usme Formation is dated as Bartonian-earliest Rupelian?
(33.9 to ca 33 Ma), whereas the youngest dated strata of the Concentracin Formation are at least Early Rupelian (ca 31.5 Ma).
Nevertheless, lithological features in both formations indicate that
they share similar depositional settings (Hubach, 1957) and periods
of marine inuence (see Section 8).
5.2. Paleoenvironments
The pollen association from all three sites is indicative of
lowland regions (e.g. Mauritiidites, P. pokornyi, L. crassa, R. guianensis, Spirosyncolpites spiralis). No pollen or spore species typical
of Andean or pramo vegetation (Hooghiemstra et al., 1993;
Hooghiemstra and Ran, 1994; Van der Hammen et al., 1973;
Wijninga, 1996) were found.
The palynooral assemblage recovered from the Santa Teresa
Formation is typical for wetlands and swampy areas and, mostly
restricted to freshwaters conditions, as neither mangrove elements
nor marine palynomorphs were found. Furthermore, a freshwater
(lacustrine) interval was also inferred from high abundances of
both Botryococcus sp. and Pediastrum sp. (549.74 me628.24 m)
(Fig. 3). We agree with the conclusions of previous works that the
upper part of this unit was formed in a lacustrine setting (Acosta
and Ulloa, 2001; De Porta, 1966). De Porta (1966) suggested that
the sedimentary sequence was formed in a lagoonal setting but
with occasional connections to the sea based on the presence of
brackish-water mollusks and gastropods reported by Anderson
(1929). However, Nuttall (1990) revisited the fauna described by
Anderson, and he concluded that neither of the two reviewed
genera ascribed to the section (Pachydon and Verena) is a denitive
indicator of brackish waters. Moreover, living species from the
genus Verena are salinity intolerant and therefore restricted to
freshwater environments (Nuttall, 1990). Consequently, there is no
evidence supporting a connection to the sea.
Similar lacustrine conditions have been reported for the upper
Mugrosa (Middle Magdalena Valley) and Barzalosa formations
(Upper Magdalena Valley). The upper part of the Early Miocene
Mugrosa Formation (Caballero et al., 2010; Hubach, 1957) is a thick
lacustrine deposit (Nuttall, 1990), which might correlate with the
lacustrine levels of Santa Teresa. Likewise, the Barzalosa Formation
(De Porta, 1966) consists of multicolored mudstones interbedded
with some conglomeratic and gravel levels and contains thick
lacustrine deposits characterized by blue and green claystones of
Early Miocene age (De Porta, 1966). On the other side of the EC, the
Early Miocene C2 member of the Carbonera Formation was
continuously deposited in the easternmost ank of the EC (Parra
et al., 2009a) and in the Llanos Basin, east of the EC (Cooper et al.,
1995). The C2 accumulated in marine-inuenced coastal plain

165

environments (Cooper et al., 1995) and contains both marine

palynomorphs and mollusks (Gmez et al., 2009; Jaramillo et al.,
2011). The pattern suggests that Santa Teresa, Barzalosa and the
upper Mugrosa formations were part of a large lacustrine system
along the Magdalena River Basin, whereas the Llanos Basin was
already separated from the Magdalena Basin and part of a different
drainage basin. This conclusion is also supported by the detrital
zircon analysis of Horton et al. (2010b), who found a signicant
absence of MesozoiceCenozoic zircons in Middle and Late Miocene
samples from the Guayabo and Corneta formations (Llanos Basin);
this was attributable to the effective topographic barrier created by
the EC by the Middle Miocene, which separated the Central
Cordillera source area from the Llanos Basin. Our data suggest that
this barrier was already present by the Early Miocene. However,
given that the Santa Teresa palynological record contains abundant
oristic elements from lowland regions and none from Andean or
pramo vegetation, this topographic barrier apparently did not
reach elevations higher than 1000 m by the Early Miocene.
The Usme palynoora, with common presence of mangrove and
marine elements (Fig. 3) together with high abundances of lowland
terrestrial oras, suggests that accumulation took place in coastal
plains with tidal inuence, as had been suggested herein and
previously (Hoorn et al., 1987; Montoya and Reyes, 2005). A welldened marine ooding event in the middle part of the formation (2130 me2160 m) is indicated by the MI index (Fig. 3). A
similar marine ooding event in the Late Eocene has already been
identied in the Central Llanos and Llanos Foothills basins (Santos
et al., 2008). The presence of Late Campanian-Maastrichtian palynomorphs (Buttinia andreevi and Syndemicolpites typicus) within
the recovered palynoora (Appendices 4 and 5) indicates an active
erosional recycling from Campanian-Maastrichtian rocks (Guadalupe-Guaduas formations?).
The palynoora of the Concentracin Formation is similar to
that of the Usme Formation having moderate abundances of both
mangrove and marine elements (Fig. 3), together with high abundances of terrestrial lowland oras. This corroborates previous
suggestions that accumulation took place in coastal plains with
tidal inuence (Saylor et al., 2011; Villamil, 1999). There are three
distinctive marine intervals (Fig. 3). The rst event occurs within
zone T-06 (40.49 m) during the Middle Eocene (Bartonian), the
second within zone T-07 (560 m) during the Late Eocene (Priabonian) and third within the uppermost T-07 to the lower T-08 zone
(751.49 me832.49 m), during the latest Eocene to Early Oligocene
(latest Priabonian to Early Rupelian) (Fig. 3). A Middle Eocene
marine event that has been identied in the central Llanos Foothills
(Jaramillo and Dilcher, 2001), in the middle of the Mirador
Formation, can be correlated with the rst marine interval
(40.49 m). Another ooding event during the Late Eocene can be
correlated with the event observed in Usme, which is registered
throughout the Llanos and Llanos Foothills basins (Santos et al.,
2008). Our results agree with previous MI values reported by
Santos et al. (2008) for the Paz del Ro area (MI < 0.02) (Fig. 3).
Santos et al. (2008) proposed that a NWeSE marine incursion
entered via the Lower Magdalena Valley and inundated the basin
up to the Central Llanos Foothills. In this way, marginal marine and
estuarine settings were established along the northern axial zone
of the Eastern Cordillera. Our results support their interpretation.
The Early Rupelian (Early Oligocene) ooding corresponds to
a maximum ooding surface that has been identied at several
sites in Colombia and Ecuador including the lower Orteguaza
Formation in the Putumayo Basin (Christophoul et al., 2002;
Jaramillo et al., 2011; Osorio et al., 2002). This ooding has been
identied in the southern part of the Upper Magdalena Valley
(Neiva Subbasin), the Putumayo Basin (Osorio et al., 2002) to the
north (Fig. 1), and the Maraon Basin (Christophoul et al., 2002).

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6. Overburden
Vitrinite reectance (Ro) data in strata from the Santa Teresa
Formation in the Guaduas Syncline show Ro values of 0.54e0.60%
(Gmez, 2001; Gmez et al., 2003), which represent maximum
temperatures of w95e105  C for heating rates of 2e10  C,
according to the kinetic model of Burnham and Sweeney (1989).
In the absence of evidence of other heating mechanisms, we
calculate that 2.5e4.5 km of overburden, with a thermal gradient of
20e30  C/km and a surface temperature of 25  C -similar to
present-day values- would account for the observed Ro values.
There are no robust constraints on the time associated with the
accumulation of such eroded overburden. However, Gmez (2001)
calculated a concordant apatite-ssion track (AFT) age of
12.1  2.3 Ma (1s error) from sandstones of the Oligocene San Juan
de Ro Seco Formation [Ro 0.64%; Temp 115  C], in the Guaduas
Syncline, by using 20 apatites with average chlorine content of
0.34%. An in-depth analysis of these data shows that a kinetic
population of 19 apatites with chlorine content of 0.1% has an AFT
age of 8.1  2.3 Ma. We thus interpret this cooling age of 6e10 Ma in
samples that attained temperatures for full thermal resetting of
uorapatites as a reasonable approximation for the age of end of
burial and accumulation.
In the Usme Syncline, a Ro value of 0.27% in the Usme Formation
(Mora et al., 2008b) documents burial temperatures lower than
50  C. Using similar surface temperature and geothermal parameters as for the Guaduas Syncline, these paleotemperatures correspond to up to w1 km of overburden. Apatite (UeTh)/He
thermochronology (closure temperature w70  C) in two sandstone samples from the lower part of the Bogot Formation along
its western limb shows highly reproducible ages (samples C540 and
D937, palynologically dated as Middle/Late Paleocene and Early
Eocene, respectively; see Bayona et al., 2010), based on 4 aliquots
per sample, of 30.7  1.8 Ma and 26.4  1.6 Ma (Bayona et al., 2010).
These data reveal ongoing exhumation in the western limb of this
syncline by Late Eocene-Oligocene times. We believe that ca
30e26 Ma is also a maximum age for the switch from sediment
accumulation and burial to non-deposition in the core of the
syncline, where the Usme strata are preserved.
Paleotemperature estimates based on Ro and AFT data for the
Cenozoic units in the Floresta area are highly variable (Mora et al.,
2010a), indicating a greater magnitude of burial the farther to the
north in the direction of the El Cocuy area (see Fig. 1). In the north,
at w6 200 , the Concentracin Formation has a Ro value of 0.80%,
corresponding to a temperature of 140e150  C, whereas 30 km to
the southwest Ro is 0.71% (temperature w115e130  C). Based on
the considerations cited above, these estimates represent burial
exceeding 4e6 km in the north and 1 km less in the southern area
sampled. Thermal histories derived from Ro and AFT data in both
areas are consistent with an onset of exhumation occurring ca
23e20 Ma (Mora et al., 2010a).
7. Stepwise cessation in sediment preservation along the
Eastern Cordillera
The transition from burial to exhumation occurred rst in the
southern central axial zone of the Eastern Cordillera (Usme
Formation) during the Rupelian-Early Chattian (ca 30eca 26 Ma).
Subsequently, sediment accumulation stopped along the northeastern axial zone of the Eastern Cordillera (Concentracin
Formation) not before the Late Chattian (ca 23 Ma); and nally, it
was not until the Late Miocene (ca 10e6 Ma) that exhumation was
initiated along the western ank of the Eastern Cordillera (Santa
Teresa Formation). The observed age differences between the Ro/
thermochronology estimates and the reported palynological dating

(Section 5.1) exposes the difference between the preserved sediments and those that should have been accumulated but were lost
by erosion, thus providing further evidence that the youngest
preserved sediments may not represent the last deposited
sediments.
The Ro/thermochronology ages reported herein agree with
previous thermochronological, structural, and sedimentological
data (e.g. detrital zircon UePb ages, apatite-ssion tracks, and
paleocurrents data), which indicate that initial exhumation of the
Eastern Cordillera took place during the Late Eocene-Early Oligocene (see Fig. 10 in Horton et al., 2010b) (Bande et al., 2012; Bayona
et al., 2008; Horton et al., 2010b; Mora et al., 2010a; Moreno et al.,
2011; Nie et al., 2010; Parra et al., 2009b; Saylor et al., 2011; Toro
et al., 2004). Evidence from the Usme Syncline (i.e. UeTh/He in
apatite and lithological features) indicates onset of exhumation by
the end of the Eocene (Bayona et al., 2010). All these sets of data
combined suggest that sedimentation indeed ceased in the
southern axial zone of the Eastern Cordillera by the Early Oligocene. Our data also support the hypothesis of a heterogeneous
Neogene uplifting process across the EC (Bande et al., 2012; Bayona
et al., 2008; Horton et al., 2010a; Mora et al., 2008a, 2010a, 2006,
2010b; Moreno et al., 2011; Parra et al., 2009b), being older in
the axial zone and having younger phases at the eastern and
western anks, and being older in the south than the north of the
axial EC.
8. Regional implications
Three different tectonic scenarios have been proposed to explain
the Cenozoic tectonic development of the axial zone of the EC and
adjacent Magdalena and Llanos basins to the west and east,
respectively. According to the rst model, the EC corresponds to
a large foreland basin system, including what are now the Llanos
and the Magdalena Valley basins (Cooper et al., 1995; Villamil,
1999). Initial breakup of the foreland basin occurred during the
Late Oligocene, with a complete segmentation by the Middle
Miocene. The second model considers a fragmentation of a continuous Paleocene foreland basin during the Middle-Late Eocene, with
complete isolation of the Llanos from the Magdalena Basin occurring by the Early-Middle Oligocene (Gmez et al., 2005; Horton
et al., 2010a, 2010b; Moreno et al., 2011; Nie et al., 2010; Parra
et al., 2009a; Saylor et al., 2011). A third model proposes that
a syn-orogenic basin to the east of the Central Cordillera was
interrupted by several localized uplifts throughout the EC (Bayona
et al., 2008; Fabre, 1987; Sarmiento-Rojas, 2001; Shagam et al.,
1984). These intrabasinal uplifts, which would give rise to
multiple intermontane basins, are the result of latest CretaceousMiddle Eocene events of tectonic deformation.
The biostratigraphic data presented here indicate that the last
preserved record of sedimentation along the present EC are not
coeval. Sediments were accumulated and preserved until the latest
Bartonian-earliest Rupelian? (Late Eocene-earliest Oligocene?) in
the southern central region (Usme Formation) and at least until the
Early Rupelian (Early Oligocene) in the northeastern central region
(Concentracin Formation), long before the major latest MiocenePliocene uplift of the EC (Van der Hammen et al., 1973). In
contrast, the western foothills (Santa Teresa Formation) and eastern
foothills (Carbonera Formation) (Parra et al., 2009a) continued
accumulating sediments during the Miocene. These marked time
differences of the youngest preserved units across the EC suggest
that a single foreland that covered the Magdalena Valley, EC and the
Llanos Basin was disrupted in the earliest Oligocene prior to the
onset of major Late Neogene exhumation pulses, in contrast to the
simpler model that was proposed during the 1990s (Cooper et al.,
1995; Villamil, 1999).

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D. Ochoa et al. / Journal of South American Earth Sciences 39 (2012) 157e169

Our palynological results indicate that during the Late Eocene

the Usme and Concentracin formations were part of the Central
Llanos Foothills and Llanos Basin, as evidenced by the similarity in
oras and the ooding event observed across this basin (Fig. 3),
suggesting that depositional systems of the Llanos-EC basin were
still connected. Growth strata in the western ank of the EC and
Magdalena Basin that record west-verging deformation of the
western ank of the EC (Corts et al., 2006; Gmez et al., 2003)
mark the westernmost boundary of depositional systems within
the Llanos-EC basin. The connection of this large depositional
system between the EC and the Llanos probably was broken during
the earliest Oligocene, as has been suggested by the models of
Gmez et al. (2005), Parra et al. (2009a), Horton et al. (2010b) and
Saylor et al. (2011). This is further supported by the similarity
between depositional environments of the EC, Llanos Foothills, and
Magdalena Valley basins until the Late Eocene (Parra et al., 2009a;
Saylor et al., 2011).

9. Conclusions
Biostratigraphic data from the youngest sediments preserved
in all three synclines indicate different ages associated with the
uppermost formations in each stratigraphic sequence. We estimated the age of the sediments from the Usme Formation (Usme
Syncline) as Bartonian-earliest Rupelian? (38e33 Ma), from the
Concentracin Formation (Floresta Syncline) as Late LutetianEarly Rupelian (39e31.5 Ma), and uppermost Santa Teresa rocks
(Guaduas Syncline) as Burdigalian (19e16 Ma). In addition, the
combination of biostratigraphic, thermochronometric, and paleothermometric data suggests that cessation of sediment accumulation in the EC was diachronous. In the southern axial zone of
the Eastern Cordillera (Usme Syncline), the shift from sediment
accumulation and burial to non-deposition occurred during the
Late Rupelian-Early Chattian (ca 30e26 Ma), whereas in the
northeastern axial zone of the Eastern Cordillera (Floresta
Syncline) it was estimated as latest Chattian-Aquitanian (ca
23e20 Ma); and in the western ank (Guaduas Syncline) as
Tortonian-Messinian (10e6 Ma). These estimates agree with dates
of published ages of exhumation of the EC, suggesting that the
process of uplifting of the EC is very complex and happened in
several steps. Finally, the palynoora indicates the exclusive
presence of lowland plant communities in all three areas,
including the Burdigalian (Early Miocene) Santa Teresa Formation, suggesting that the EC still was less than 1000 m in elevation
by the Early Miocene. Lowland conditions in the area of the
present EC gradually ended between the Oligocene and the Late
Miocene.

Acknowledgments
We thank the Colombian Petroleum Institute, the Smithsonian
Tropical Research Institute, the Corporacin Geolgica Ares, and
Colciencias for their nancial, logistic, and technical support.
Thanks to Andrs Pardo (Universidad de Caldas) for the palynological analysis of most of the samples of the Concentracin
Formation. We specially thank J. Saylor, B. Horton, J. Kellogg and an
anonymous reviewer for their helpful comments. Natasha Atkins
improved readability of the manuscript.
Appendix A. Supplementary material
Supplementary material associated with this article can be
found, in the online version, at doi:10.1016/j.jsames.2012.04.010.

167

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