Quaternary Science Reviews 86 (2014) 49e62

Contents lists available at ScienceDirect

Quaternary Science Reviews

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

Early and Middle Holocene evidence for plant use and cultivation in
the Middle Cauca River Basin, Cordillera Central (Colombia)
Francisco J. Aceituno a, *, Nicolás Loaiza a, b
a
Grupo medioambiente y sociedad, Departamento de Antropología, Universidad de Antioquia, Calle 67 No 53-108, AA 1226 Antioquia, Colombia
b
Temple University Department of Anthropology, Philadelphia, PA 19119, USA

a r t i c l e i n f o a b s t r a c t

Article history: This paper presents the latest results of research done in the Colombian Andean region known as Middle
Received 5 July 2013 Cauca River Basin, an important location for the study of the origins of plant use and the dispersal of
Received in revised form domesticates throughout the Americas due to its geographical position in northwest South America. We
30 November 2013
discuss humaneenvironment interactions during Pleistocene/Holocene transition to middle Holocene
Accepted 13 December 2013
Available online
(ca 10,000e4000 BP), specifically humaneplant interaction and environmental factors that led to the
adoption of horticultural practices. Three lines of evidence are analyzed: archaeological stratigraphy,
lithic technology, and microbotanical remains. Our results suggest that early Holocene environmental
Keywords:
Colombia
stability allowed Middle Cauca settlers to use the diverse local resources for several millennia, altering
Cauca River Basin the local vegetation, and leading to the development of horticultural practices that included the use of
Early Holocene both local and foreign plants. These results inform the ongoing debate about the antiquity and nature of
Lithic technology plant domestication and dispersals in the Americas.
Starch grains Published by Elsevier Ltd.
Plants management

1. Introduction societies of the Intermediate Area, such as the origin of plant

Until recently, our understanding of preceramic occupation in
the humid montane forests of the Colombian Andes was limited
due to the lack of known archaeological sites dating to this period in
this region. With the implementation of cultural resource man-
agement in Colombia in the mid 1990’s, contract archeology started
to yield new information about the preceramic period in the Andes
and increased the meager data produced by previous studies
(Correal and van der Hammen, 1977; Correal, 1979, 1986; Herrera
et al., 1988; Salgado, 1988e1990; Gnecco and Salgado, 1989;
INCIVA, 1995e1996; INTEGRAL, 1997; Gnecco, 2000; Cano, 2001,
2004; Aceituno and Castillo, 2005; Otero and Santos, 2006;
Santos, 2008). The information presented here combines the lat-
est data from several early preceramic sites as well as a broader
archaeological contextualization using data published elsewhere
(e.g Aceituno, 2001; Aceituno et al., 2001; Aceituno and Castillo,
2005; Aceituno and Loaiza, 2006, 2007, 2008, 2010; Castillo and
Aceituno, 2006) that provide relevant information for under-
standing the evolution and adaptive changes in prehistoric
* Corresponding author. Tel./fax: þ57 211 83 59.
E-mail addresses: csfjace@antares.udea.edu.co, aceitunob@hotmail.com (F.
J. Aceituno).
cultivation and expansion of agriculture are two important evolu-
tionary developments that enabled the rise of sedentary villages
and the development of complex societies (e.g. Piperno and
Pearsall, 1998; Diamond, 2002; Dickau, 2005; Iriarte, 2009).
After the Pleistocene/Holocene transition and the end of the
Paleoindian period ca 10,000 BP, significant climatic changes
occurred that affected the flora and fauna of northern South
America, altering the conditions to which hunteregatherers
needed to adapt (Reichel-Dolmatoff, 1997 [1965]: 57). Although the
archaeological data for the period between 10,000 and 7000 BP is
very limited, it has been suggested that significant environmental
and cultural changes occurred that shaped early Holocene human
occupation (e.g. Herrera et al., 1988; Gnecco and Salgado, 1989;
Gnecco, 2000; Aceituno and Castillo, 2005; Aceituno, 2007;
Aceituno and Loaiza, 2007). The increase of archaeological sites
indicates that this was a period of expansion and population
growth along the Northern Andes (Aceituno, 2007; Aceituno and
Loaiza, 2007; Aceituno et al., 2013). The archaeological record
shows a high heterogeneity in lithic technology between regions,
including the emergence of stone tools used for preparing plant
foods. The new environmental conditions imposed by an increase
in temperature and rainfall (Marchant et al., 2002), and the sub-
sequent expansion of the humid tropical forests, led human groups
to make adaptive adjustments depending on the characteristics of

0277-3791/$ e see front matter Published by Elsevier Ltd.

http://dx.doi.org/10.1016/j.quascirev.2013.12.013
50 F.J. Aceituno, N. Loaiza / Quaternary Science Reviews 86 (2014) 49e62
the new ecosystems, amongst which we highlight the increased
usage of plant resources and the first manifestations of plant
cultivation in tropical and premontane forests (Cavelier et al.,
1995; Piperno and Pearsall, 1998; Gnecco, 2003; Mora, 2003;
Castillo and Aceituno, 2006; Piperno, 2006; Aceituno and Loaiza,
2007, 2008; Santos, 2008).
Within this context, this paper aims to report, review, and
summarize the archaeological record of the preceramic occupa-
tions in the Middle Cauca River Basin (MC), an inter-Andean region
located in the western central part of the Northern Andes
(Colombia), and suggest some explanations about adaptive pat-
terns to humid montane forest ecosystems for the period between
the early and middle Holocene. This period was marked by warmer
and wetter conditions than those of the Late Pleistocene, and the
spread of seasonal and humid tropical forests (Marchant et al.,
2004).
The Colombian Andes, due to its geographical position and high
environmental diversity, is an important region to investigate
major issues regarding the early human history of Northwest South
America, such as population dispersals, cultural diversity, and ad-
aptations of mobile huntereforagers and early cultivators. The
Colombian Andes links Panama and Ecuador where during the
early Holocene, there is evidence of ecosystems alteration and
intensive use of plants (e.g. Pearsall and Piperno, 1990; Piperno
et al., 1991; Piperno and Pearsall, 1998; Piperno et al., 2000;
Piperno and Stothert, 2003; Ranere and Cooke, 2003; Stothert
et al., 2003; Pearsall et al., 2004; Zarrillo et al., 2008). This pro-
vided the basis for later development of wide-spread swidden
horticulture in the middle Holocene, accompanied by a territorial
growth and population expansion (e.g. Cooke and Ranere, 1992;
Bruhns, 1994, p.83; Pearsall, 1994; Ranere and Cooke, 1995;
Piperno and Pearsall, 1998; Cooke, 2005). The MC region is
known to have an abundance of sites (<20) with occupations
reaching back to the Pleistocene/Holocene transition where plant
usage has been inferred (Herrera et al., 1988; INCIVA, 1995e1996;
INTEGRAL, 1997; Aceituno and Loaiza, 2007). The data discussed
in this paper helps bridge the gap between the other tropical re-
gions, because it provides a strategic location to investigate early
plant cultivation, and dispersal of various plant species in the
Neotropics, such as Zea mays (maize), Phaseolus vulgaris (beans),
Dioscorea spp. (yam) and Manihot esculenta Crantz (manioc). In
addition, the humid premontane and montane tropical forests of
the MC region represent environments that have little archaeo-
logical attention in the past in contrast to the lowland rainforest
(e.g. Bailey et al., 1989; Bauchet et al., 1991; Piperno and Pearsall,
1998) new information in terms early human history as well as
the origins of plant cultivation.
To investigate these issues, multiple lines of data were analyzed
within a broad stratigraphical context that the correlation of
archaeological evidence with paleoenvironmental conditions
within tightly restricted periods of time. Lithic tools are the main
technological indicator, from which starch grains were extracted for
analysis and identification of direct plant usage. Sediment columns
for pollen analysis were taken from walls at excavation sites and
enabled paleobotanical reconstruction as well as the identification
of potential economic plant species.
2. Regional setting
The Colombian Andes are comprised of three major cordilleras:
Occidental, Central, and Oriental. The Oriental and Central Cordil-
leras are separated by the Magdalena River, while the Central and
Occidental Cordilleras are separated by the Cauca River, a tributary
of the Magdalena which rejoins it in the Mompox Depression north
of the Central Coridllera(Fig. 1). The MC region roughly coincides
with the Eje Cafetero (the coffee belt), with elevations ranging from
ca 800 m asl (meters above sea level) at the river level in the north
end to ca 5300 m asl at the highest peaks in the Cordillera Central,
temperature and rainfall vary according to elevation.
So far, over 20 archaeological sites that date back to the early
Holocene have been found within the MC region in several inter-
Andean river valleys of the Central Cordillera (INCIVA, 1995e1996;
INTEGRAL, 1997; Tabares and Rojas, 2000; Cano, 2004, 2008;
Tabares, 2004; Aceituno and Loaiza, 2007; ; Aceituno and
Lalinde, 2011; Herrera et al., 2011; Restrepo, 2013). According to
the Holdridge classification (adapted to local environments by
Espinal, 1990), these sites are located in the very humid pre-
montane forest life-zone that ranges between 1000 and
2000 m asl, with average temperatures ranging between 18 and
24  C, and yearly rainfall from 2000 to 2750 mm (Espinal, 1990;
IGAC, 1998). In geological terms, the Central cordillera landscape
was formed under strong volcanic influence. Enormous mudflows
generated by the melting of ice sheets after volcanic eruptions
around four million years ago make up most of the base formation
(Tistl, 2006). During the height of the last glaciation (ca 20,000 e
cal. BP.), activity was shifted to volcanic explosions that covered
the region with successive ash falls that happened on average
every 3000e4000 years, which differentially eroded to create an
undulating landscape of smooth hills (Salomons, 1989; Tistl,
2006). Early settlers found a landscape similar to the present
one, with deep, well-drained, fertile soils (IGAC, 1998; Hermelin,
2001). Preceramic deposits were covered by more recent ash
falls that preserved most sites, burying them as much as 2 m
below the modern surface.
The data presented here come from the sites El Jazmín, La
Pochola, San Germán, and La Selva, all located between 1600 and
1677 m asl. We compare these sites with several others from similar
environmental settings, representing two different areas: the San
Eugenio Valley, El Antojo (lithic analysis) (INTEGRAL, 1997) and
Guayabito (palynological analysis) (INTEGRAL, 1997) (Fig. 1); and
from a nearby area ca 15 km north from the main sites in the MC,
Campoalegre site (INTEGRAL, 1997), and 39 El Recreo Cancha site
(Herrera et al., 2011).
A small description of each of the four sites follows:
 El Jazmin (1650 m asl): located on top of a wide ridge east of the
San Eugenio River. Preceramic deposits have been dated from
4715  45 BP to 10,120  70 BP (Aceituno and Loaiza, 2007)
(Table 1), and include handstones, grinding bases, notched axes/
hoes (over 20 in total), flake tools, and debitage. Both palyno-
logical and starch grain analyses have been done at this site.
 La Pochola (1677 m asl): located on top of a rounded terrace
ca 3 km south of El Jazmín but west of the San Eugenio River. The
preceramic lithic assemblage is composed of handstones,
grinding bases, flaked tools, and debitage. Four radiocarbon
dates refer to the preceramic deposits; 6743  45 BP,
6903  45 BP, 8680  55 BP and 9312  45 BP. (Aceituno and
Loaiza, 2007; Aceituno and Lalinde, 2011) (Table 1). Both paly-
nological and starch grain analyses have been done at this site.
 San German (1649 m asl): located less than one kilometer south
of La Pochola on the same side of the San Eugenio River, on the
top of a terrace overlooking the junction of two small streams
that join the San Eugenio River approximately 600 m to the
northeast. From a single small excavation unit, grinding and
flaked tools, as well as debitage, were recovered associated with
a date of 8136  65 BP (Aceituno and Loaiza, 2007) (Table 1).
Starch grain analysis was done at the site.
 La Selva (1600 m asl): located on a high terrace overlooking the
Cauca River Valley ca 20 km northwest from El Jazmin. Pre-
ceramic deposits, associated with four dates between
 F.J. Aceituno, N. Loaiza / Quaternary Science Reviews 86 (2014) 49e62 51

Fig. 1. Maps of the Middle Cauca Region showing the location of the sites discussed in the paper.
52 F.J. Aceituno, N. Loaiza / Quaternary Science Reviews 86 (2014) 49e62

Table 1
Midde Cauca Early and Middle Holocene chronology of the sites referred in Fig. 1.
14
Site C years BPa Calibrated dateb 1 d Lab. code Material Reference

El Jazmin 10,120  70 10,019 BC: 9558 BC Ua-24497 Charcoal Aceituno and Loaiza, 2007
La Selva 9490  110 9221 BC: 8556 BC Beta-87188 Charcoal INCIVA, 1995e1996
La Pochola 9312  55 8640 BC: 8460 BC LTL4223A Charcoal Aceituno, 2010
La Pochola 9047  45 8290 BC: 8240 BC LTL5436A Charcoal Aceituno, 2010
El Jazmin 9020  60 8297 BC: 8208 BC Beta-95061 Charcoal INTEGRAL, 1997
La Selva 8712  60 7791 BC: 7605BC X23803 Charcoal Ranere, 2012, com. per.
La Selva 8704  56 7756 BC:7604 BC X23805 Charcoal Ranere, 2012, com. per.
La Selva 8680  60 7739 BC:7600 BC Ua-24498 Charcoal Aceituno and Loaiza, 2007
La Selva 8674  61 7734 BC:7599 BC X23804 Charcoal Ranere, 2012, com. per.
39 El Recreo Cancha 8550  60 7711 BC:7498 BC Beta-285871 Charcoal Herrera et al., 2011
39 El Recreo Cancha 8480  40 7587 BC:7498 BC Beta-290954 Charcoal Herrera et al., 2011
El Antojo 8380  90 7539 BC: 7344 BC Beta-93154 Charcoal INTEGRAL, 1997
San German 8136  65 7284 BC: 7051 BC CSIC 1987 Charcoal Aceituno and Loaiza, 2007
La Pochola 8095  55 7731 BC: 7602 BC Ua-24499 Charcoal Aceituno and Loaiza, 2007
39 El Recreo Cancha 8030  80 7303 BC: 6824BC Beta-283582 Charcoal Herrera et al., 2011
Guayabito 7990  100 7056 BC: 6768 BC Beta-95064 Charcoal INTEGRAL, 1997
La Selva 7685  110 6819 BC: 6344 BC Beta-87189 Charcoal INCIVA, 1995e1996
Campoalegre 7600  90 6587 BC: 6584 BC Beta-87730 Charcoal INTEGRAL, 1997
El Jazmin 7590  60 6492 BC: 6490 BC Beta-95602 Charcoal INTEGRAL, 1997
El Jazmin 7080  50 6012 BC: 5906 BC Ua-24496 Charcoal Aceituno and Loaiza, 2007
La Pochola 6903  45 5675 BC:5620 BC LTL4222A Charcoal Aceituno and Lalinde 2011
La Pochola 6743  45 5840 BC: 5730 BC LTL4221A Charcoal Aceituno and Lalinde 2011
La Selva 5825  70 4840 BC: 4514 BC Beta-87508 Charcoal INCIVA, 1995e1996
El Jazmin 5625  50 4501 BC: 4370 BC Ua-24495 Charcoal Aceituno and Loaiza, 2007
El Jazmin 4715  45 3629 BC: 3378 BC Ua-24494 Charcoal Aceituno and Loaiza, 2007
Campoalegre 4270  70 3011 BC: 2977 BC Beta-99859 Charcoal INTEGRAL, 1997
Guayabito 4180  70 2885 BC: 2836 BC Beta-95063 Charcoal INTEGRAL, 1997
a
All dates are given in uncalibrated 14C years BP unless noted.
b
Calibration done using Calib Rev. 5.0.1.

5825  70 BP and 9490  110 BP, contained handstones,
grinding bases, flake tools, and debitage (Aceituno and Loaiza,
2007) (Table 1). Both palynological and starch grain analyses
have been done at this site.
3. Materials and methods
Archaeological sites were excavated in arbitrary 5 cm levels. Soil
horizons were determined based on typical indicators such as color
and texture (Tobon and Perez, 2005; Tobon and Castaño, 2006). Soil
samples taken from defined horizons were separated into thick and
thin fraction using ASTM sieves. Thick fraction (>200 ASTM sieve)
was done using binocular petrographic microscope at a 10
magnification in order to determine soil texture and to identify
provenance and mineral transport (based on degree of roundness)
(Tobon and Perez, 2005; Tobon and Castaño, 2006). Thin fraction
(200 > ASTM sieve) was analyzed using X-ray diffraction to
determine the weathering degree of clays, which is related to the
climatic conditions they were exposed to (Tobon and Perez, 2005;
Tobon and Castaño, 2006).
Stone artifacts recovered in excavations were bagged, and tag-
ged according to archaeological context. Artifacts were classified
and, in general terms, 6 major types were defined: debitage flakes,
cores, handstones (Fig. 4B), milling stones (Fig. 4C) flake tools, and
axe/waisted hoes (Fig. 4A). Typical metric variables were described
for each tool, as well as raw materials, to facilitate correlation and
proper classification. Tool frequencies create cultural stratigraphy,
define the preceramic component of sites, and were key elements
in the selection process of datable materials.
Fifteen tools were analyzed from preceramic levels of La
Pochola, El Jazmin, San German II, and La Selva. Starch grains were
extracted in the lab following standard protocols (Loy, 1994;
Pearsall et al., 2004; Piperno, 2006; Torrence and Barton, 2006).
In all but two cases, grinding tools were selected for extraction
because these types of artifacts are more likely to have starch grains
embedded within their microtopography (Table 2).
Extraction from tools was done by a) humidifying the surface
with distilled water; b) extracting the grains with metallic probes;
and c) collecting the dissolved sediment in water vials perfectly
sealed to avoid sample contamination. Once the samples were
obtained, the starch grains were separated from the sediments
attached to stone tools. First, Sodium Hexameta-phosphate
(NaPO4)6, was added in a stir plate to disaggregate the sample.
The sample was then centrifuged at 2000 r.p.m. for 15 min to
concentrate the residue and later decanted. A solution of heavy
water prepared with Cesium Chloride (CsCl) with a density of
1.8 g/ml, was then added in order to separate the starch grains by
flotation (Dickau, 2005; Pagan et al., 2005; Piperno, 2006, p. 60).
The sample was centrifuged an additional five at 2000 r.p.m. to
allow the heavier particles to settle and the starch grains to float;
2 ml was removed from the floating surface and later deposited in
a separate test tube. Distilled water was added to the tube with
the floating section; the sample was stirred and centrifuged at
2000 r.p.m. for 15 min and decanted. This standard procedure was
repeated three times until the residue was concentrated at the
bottom of the test tube. The final residue was affixed to micro-
scope slides and observed under an Olympus CX-41 microscope, at
magnifications between 100 and 1000. The slides were
observed through a zigzag scan starting at the lower right corner.
Starch grains were described using metric and morphological
variables that can be used for taxonomical identification by com-
parison to reference specimens of archaeological starch grains (Loy,
1994; Perry, 2002; Dickau, 2005; Piperno, 2006; Torrence and
Barton, 2006; Babot, 2007). Identification of recovered archaeo-
logical starch grains was made using a regionally specific reference
collection of 59 wild and domesticated plant specimens kept at the
Archeology Laboratory of the Universidad de Antioquia, Medellin,
Colombia, as well as comparisons with digital images from the
University of Exeter Neotropical Archaeobotanical Reference
Collection (235 species), and numerous published sources (Piperno
and Holst, 1998; Piperno et al., 2000; Perry, 2002, 2004; Dickau,
2005; Pagan et al., 2005; Dickau et al., 2007; Holst et al., 2007;
 F.J. Aceituno, N. Loaiza / Quaternary Science Reviews 86 (2014) 49e62 53

Fig. 2. Stratigraphic profile from the El Jazmín site. See the text for a description of the layers.

Perry and Flannery, 2007; Piperno and Dillehay, 2008; Rumold,
2010; Aceituno and Lalinde, 2011; Bonomo et al., 2011).
4. Results
volcanic glass and weathering of minerals. This horizon separates
the preceramic (IIIAb and below) component from the ceramic
component (B/C and above) (Fig. 3) (Tobon and Perez, 2005; Tobon
and Castaño, 2006).

A comparison of soil horizons between archaeological sites

suggest that El Jazmin’s profile can serve as the type profile for the
zone (Fig. 2). The major soil developments and events can be traced
in this sites 5 horizons (A, B/C, IIC, IIIAb, IIIB) that can be further
subdivided into different ash-falls that became the parent materials
for the soil formation process (Fig. 3). This also enabled the iden-
tification of a particular event that buried the preceramic compo-
nent and separated it from the posterior ceramic component
(Aceituno and Loaiza, 2010).
Mineralogical analysis, particularly of volcanic glass frequencies,
identified 2 major events that are clearly separated. These two
major events roughly coincide with the preceramic and ceramic
cultural components. IIIB and IIIAb horizons. According to the
volcanic glass record they represent an initial sedimentary event
(Fig. 3). IIC horizon is described as a reworked horizon due to loss of

Fig. 3. Stratigraphic distribution volcanic glass at the El Jazmín site.
Fig. 4. Lithic artifacts. A- Axes/waisted hoes: 1) El Jazmin Block III Level 20 (5 cm above
a 10,120  70 BP date), 2) El Jazmin Block III Level 14 (dated 5625  50 BP). B-
Handstones: 1) El Jazmin Block III Level 15 (dated between 7080  50 and
5625  50 BP), 2) La Pochola Block I Level 13 code 733 (dated between 6903  45 and
6743  45 BP. Starch grains extracted see Table 2), 3) San German Block I Level 9 (dated
8136  65). C- Milling Stones: 1) La Pochola Block I Level 12 (dated 6743  45), 2) El
Jazmin Block III Level 16 (dated between 7080  50 and 5625  50 BP).
54 F.J. Aceituno, N. Loaiza / Quaternary Science Reviews 86 (2014) 49e62

Table 2
Starch grains recovered from tools and associated dates.
14
Site Stone tool Assoc date C BP Phaseolus spp. Zea mays Dioscorea spp. Manihot spp. Unidentified Total

La Pochola Handstone 55 8095  55 0 43 43

La Pochola Handstone 79 8095  55 1 21 22
La Pochola Handstone 23 6903  45 & 6743  45 3 1 24 28
La Pochola Handstone762 6903  45 & 6743  45 4 24 28
La Pochola Handstone733 6903  45 & 6743  45 3 23 26
La Pochola Handstone 236 6743  45 2 16 18
El Jazmín Base 315 >7590 5 2 7
El Jazmín Base 514 >7590 2 1 5 1 9
El Jazmín Handstone 265 7080  50 5 1 76 82
El Jazmín Handstone 830 5625  50 1 27 29
El Jazmín Handstone 202 5625  50 & 7528  51 4 1 31 36
San Germán Handstone 8136  65 16 16
La Selva Base 39 8712  60 2 1 10 13
La Selva Chopper 90 >8712  60 1 2 5 11 20
La Selva Axe 281 8712  60 1 12 13
Total 22 11 5 13 337 390

Lithic artifacts are the earliest indicators of human occupations
in the valley, dating to 10,120  70 BP, from associated charcoal in El
Jazmin (Aceituno and Loaiza, 2007). There are a few lithic artifacts
in levels up to 10 cm below this dated association in the IIIB Hz, but
they remain undated. In the time interval between 9000 and
5000 BP, lithic artifacts appear in their the highest frequencies at all
sites, associated with an increase of grinding tools (Fig. 4B, C) and
the appearance of axes/waisted hoes (Fig. 4A). Frequencies start
dropping and lithic artifacts practically disappear towards
ca 4200 BP, when environmental conditions rapidly changed in the
valley due to a series of volcanic events documented between 5400
and 3100 BP (Salomons, 1989, p. 33).
A total of 372 starch grains were recovered from ten hand-
stones (La Pochola, El Jazmín and San German II), three grinding
bases (La Selva and El Jazmin), an axe/waisted hoe, and a chopper
(La Selva). No grains presented alteration due to heat (Henry et al.,
2009).
Twenty two oval/kidney shaped grains with lengths ranging
from 14.75 mm to 54.9 mm averaging 27.34 mm, falling well within
the length range values for modern common bean (P. vulgaris)
between ca 13 and ca 60 mm (Piperno and Dillehay, 2008) were
identified in nine artifacts from La Pochola, El Jazmin, and La Selva
(Table 2). They have no pressure facets, have discernible lamellae,
and a ragged longitudinal fissure (Fig. 5) (Piperno and Holst, 1998;
Dickau, 2005, p.186; Piperno and Dillehay, 2008).
Eleven starch grains extracted from two handstones from El
Jazmin and La Pochola were consistent with Z. mays (Table 2). These
grains are between 10 mm and 20 mm long, averaging 16.14 mm. This
fits within the average of modern maize starch, which is larger than
other Poaceae seed starch (Holst et al., 2007). They are irregularly
shaped with pressured facets on their sides. The hilum is open and
centric, and four of them display transverse fissures and one stel-
late fissure (Fig. 6) (Piperno and Holst, 1998; Perry, 2004; Dickau,
2005, p.185; Dickau et al., 2007; Holst et al., 2007).
Five triangular starch grains ranging between 17 and 34 mm
long, averaging 25.86 mm with cunate lamellae, and eccentric hilum
have been identified as Dioscorea spp., extracted from tools from La
Pochola, El Jazmin, and La Selva (Table 2) (Fig. 7). These features are
consistent with Dioscorea spp. starch grains described in the liter-
ature (Piperno and Holst, 1998; Dickau, 2005, p. 192; Pagan et al.,
2005).
A total of 13 bell shaped grains with lengths ranging from
13.01 mm to 20.75 mm, averaging 16.63 mm identified as Manihot
spp. starch grains (Fig. 8) were extracted from five stone tools, four
from El Jazmin and one from La Selva (Table 2). These values are
well within the ranges described in the literature for both
domesticated and wild varieties (Piperno, 2006, p. 53e57). Five of
the starch grains present longitudinal fissures, two present Y-sha-
ped fissures, 2 present wing-shaped fissures and two stellate fis-
sures, the remaining grains having no fissures.
5. Discussion
In all sites, the preceramic component is associated with an Ab
soil horizon that has slight variations (and subdivisions at El Jaz-
mín) depending on geomorphology and topography. This period
was dated between 10,120  70 BP and 4180  70 BP, with inter-
mediate dates shown in Table 1. At El Jazmín, the full period is
represented: the earliest date (10,120  70 BP) was obtained at the
stratigraphic boundary with the horizon below Ab, and the latest
date (4715  45 BP) was obtained at the boundary with the horizon
above Ab.
On a regional scale, the Ab Horizon represents an environ-
mentally stable period, lasting from the end of the Pleistocene
through Middle Holocene, even though there is evidence of some
volcanic activity during this time (Orozco, 2001). Stability is sug-
gested by the fact that three separate paleosols have been identified
at nearby Otun Lake, dated 8000, 6000, and 3000 BP, respectively,
suggesting that despite the ash falls, there were also periods of time
that lasted long enough to form stable and durable A Horizons (Toro
et al., 2001). Landscape stability is also supported by mineral
morphology, which reveals very low particle movement, inferred
by of the angular nature of the sediment particles: a phenomenon
also observed at other sites where mineralogy analysis has been
carried out, such as La Pochola, San German (Aceituno and Loaiza,
2007) and 39 El Recreo Cancha (Herrera et al., 2011). This enables
us to extend the argument of landscape stability to the entire
zone (Aceituno and Loaiza, 2007).
El Jazmin’s stratigraphy is representative of the formation pro-
cesses that acted upon the sites of the San Eugenio River basin. The
stratigraphic profile from Block 3 is divided into five soil horizons
(Hz) that may be partitioned into sub-horizons (subhz) (Fig. 2) The
A Hz extends between 5 cm and 55 cm below datum level (BDL)
and is divided into three sub-horizons, A1, A2, and A3, all associated
with ceramics, and disturbed by modern coffee cultivation. The B/C
Hz underneath the A Hz has the same parent material and extends
between 55 cm and 80 cm BDL. This horizon has low bioturbation
and at its bottom there is a discontinuity that marks the presence of
a different parent material. The IIC Hz, underneath B/C, lays buried
between 80 cm and 110 cm BDL and is the product of a major
depositional event: an ash flow with volcanic characteristics and
probably regional effects (following the volcanic emission
 F.J. Aceituno, N. Loaiza / Quaternary Science Reviews 86 (2014) 49e62
Fig. 5. Starch grains Phaseolus spp. a) La Pochola (762); b) El Jazmín (514); c) La Selva (90); def) modern Phaseolus vulgaris (code Larq 155).
chronology given by Orozco, 2001) that covered and preserved the
preceramic surfaces dealt with by this study. IIIAb Hz starts
ca110 cm and ends ca 155 cm BDL; it is divided into IIIAb1 subhz,
and IIIAb2 subhz. These two sub-horizons contain the archaeo-
logical and botanical evidence discussed in this paper and are dated
between 4715  45 BP at the intersection with II C on top and
10,120  70 BP at the intersection with IIIB at the bottom. III B Hz is
the deepest horizon buried from 155 cm to the bottom of the
excavation at 180 cm BDL and marks the base of the formation of
anthropic soils.
Middle Cauca lithic technology shares some features with other
Intermediate Area lithic industries from the same time period.
Broadly speaking, these include: 1) a decrease in bifacial technol-
ogy during the early and middle Holocene, 2) an increase of plant
processing tools such as handstones, milling bases, and axes during
the same time period, 3) the use of local materials for production,
and 4) the predominance of simple flaked tools. Early to middle
Holocene lithic technology dating to ca 10,000/4000 BP is charac-
terized by a high proportion of grinding tools, a low proportion of
flaked tools (and their by-products), and a number of
55
undetermined artifacts with different shapes and sizes. Grinding
tools consist of handstones and milling bases made out of coarse
grain andesite and dacite river cobbles and boulders (e.g. Correal
and van der Hammen, 1977; Correal, 1979, 1986; Lynch and
Pollock, 1981; Temme, 1982; Stothert, 1985; Salgado, 1988e1990;
Ardila and Politis, 1989; Barse, 1995; Cavelier et al., 1995; Ranere
and Cooke, 1995; Morcote et al., 1998; Gnecco, 2000; Aceituno
and Castillo, 2005; Cooke, 2005; Aceituno and Loaiza, 2007, 2008;
Ranere and López, 2007; Ranere, 2008; Santos, 2008).
Most of these characteristics are shared with other Colombian
regions, such as the use of local raw materials, the high frequencies
of grinding tools, the predominance of expedient flaked artifacts,
and the presence of axes or axes/waisted hoes (e.g. Salgado, 1988e
1990; Rodríguez, 1995; Aceituno and Castillo, 2005; Ranere and
López, 2007; Santos, 2008). Compared with most of these other
areas, MC sites have little evidence of tool making, except El Antojo.
This may be due to recurrent cleaning of living areas, off-site tool
making, and transportation of some tools made from higher quality
raw materials (Aceituno and Loaiza, 2007, p. 80). It is important to
note that recent findings in the MC include two stemmed projectile
56 F.J. Aceituno, N. Loaiza / Quaternary Science Reviews 86 (2014) 49e62

Fig. 6. Starch grains Zea mays. aeb) La Pochola (236); c) El Jazmín (265); def) modern Zea mays (code Larq 9).

points. Both were recovered in excavations and one of them made
from quartz has been dated between ca 8000 BP and ca 8500 BP.
(Herrera et al., 2011), while the other, made in black chert still re-
mains undated. This suggests that even though activities were
mainly focused on plant resources, hunting may still have played
some role in adaptive strategies.
MC lithic technology shares some major similarities to that of
other preceramic sites in Neotropical forests: much of it appears to
be aimed at the exploitation of diverse resources of the humid
forests, it is made with local materials, and it is highly expedient
(Ranere, 1980, 2008; Stothert, 1985; Bruhns, 1994:67; Ranere and
Cooke, 1995; Piperno and Pearsall, 1998; Rossen, 1998; Ranere
and López, 2007). Lithic assemblages are composed of tools made
on flakes that are either not retouched or are minimally retouched
as well as specialized artifacts for obtaining and processing plant
resources, such as axes/waisted hoes and grinding tools.
The association between lithic technology and microbotanical
evidence suggests that plant use was an important adaptive strat-
egy of the preceramic groups in the MC premontane forests. Starch
grains recovered from grinding tools and the edges of the axes/
waisted hoes, reinforce the idea that they were used for plant
processing.
Phaseolus is a genus of the Fabaceae family, whose two most
important species in the Americas are P. vulgaris (common bean)
and Phaseolus lunatus (lima bean). The wild varieties of common
bean have a wide distribution in the Americas, from Mexico to the
Andes of Peru, Bolivia and Argentina (Chacón et al., 2005; Chacon,
2009), preferring habitats with dry, warm temperatures, and an
altitude that varies by region, at elevations between ca 1000 m asl
to ca 2600 m asl in the Andean region (Piperno and Pearsall, 1998,
p.134). Wild Lima bean has a wide distribution between Meso-
america through north Argentina. South American varieties can be
 F.J. Aceituno, N. Loaiza / Quaternary Science Reviews 86 (2014) 49e62
Fig. 7. Starch grains Dioscorea spp. a) La Selva (90); b) El Jazmin (510); ced) modern Dioscorea trifida (code Larq 139).
found in the western slopes of the Ecuadorian Andes and northern
Peru between ca 400 and ca 2000 m asl (Chacon, 2009).
Recent phylogenetic studies indicate two major regions as in-
dependent domestication centers for P. vulgaris. The first one is a
region that encompasses portions of several Mexican States (i.e.
Oaxaca, Durango, and Jalisco), and studies have yet to determine if
it was domesticated only once or more times (Chacón et al., 2005,
Chacon, 2009). The second domestication center is located in the
Apurimac-Cuzco region of southern Peru, where a single domes-
tication event took place (Chacón et al., 2005; Chacon, 2009). In
the case of P. lunatus two independent domestication centers have
been identified in South America one in southern Ecuador, the
other one in northern Peru (Piperno and Dillehay, 2008). That
said, Chacon (2009) argues that since there are wild populations
of P. lunatus in the regions mentioned above but also in Meso-
america, this crop could have been originally domesticated in
Mesoamerica too.
According to Piperno and Dillehay (2008), starch grains from
wild ancestral species of both P. vulgaris and P. lunatus overlap in
size with domesticated species, obscuring species-level tax-
onomical identification. In their study of the Ñanchoc Valley, in
northern Peru, they further conclude that it is not possible to
distinguish between common and lima bean starch grains, nor to
determine if they belong to a wild or domesticated species. Piperno
and Dillehay (2008) base their assessment that their identified
Phaseolus dated ca 8080 and ca 6970 BP is a cultivar since the wild
Phaseolus distribution does not match with what they found. At this
point, we cannot give a precise taxonomical identification for the
archaeological Phaseolus starch grains from MC, since there are
several species and varieties that grow in Colombia, none of which
have been suggested as ancestral to the domesticated species.
Therefore all we can say is that by ca 8700 BP MC inhabitants were
using and processing with stone tools some wild species of Pha-
seolus that cannot be identified to the species level at this point and
that is unlikely related to either of the domesticated species.
Neither can we suggest whether or not it was cultivated or
collected, local or foreign.
57
Z. mays was originally domesticated in the Balsas River Basin of
southwestern Mexico before 7900 BP (8700 cal. BP) (Piperno et al.,
2007, 2009), and likely sometime around 9000 cal. BP based on
molecular evidence (Matsuoko et al., 2002). Much has been dis-
cussed about the importance of maize in prehispanic economies.
For the tropical forests the case has been made that maize spread
rapidly because it became a basic resource since its arrival, due to
its advantages for storage and transport less bountiful tropical
forests (Piperno and Pearsall, 1998, p.315). Whatever the case,
archaeobotanical data suggest that maize adapted well to the
environmental conditions of tropical rainforests. In the MC the
starch grain evidence suggests that maize was introduced, around
7000 BP, to local plant cultivation practices and was part of the
consolidation of this adaptive strategy that now encompassed the
use of higher return rate crops planted in gardens near campsites
(Dickau, 2005; Aceituno and Loaiza, 2007).
Dioscorea spp. starch grains indicate the use of a plant from this
genus for its carbohydrate rich rhizomes. To date, little is known
regarding the domestication process of this plant (Piperno, 2011).
Many Dioscorea species are cultivated in household gardens in the
Neotropics today (Brücher, 1989, p. 19), and it is likely that the same
applies to the past. Several edible species are commonly used by
indigenous and peasant societies in northern South America, i.e.
Dioscorea dodecaneura and Dioscorea trifoliata, and the domesti-
cated species Dioscorea trífida. The latter is the most important
member of the genus for human beings whose domestication lo-
cality has been hypothesized as a wide lowland area of northern
Brazil, the Guyanas, Suriname, and southern Venezuela (Piperno
and Pearsall, 1998, p.117; Piperno, 2011). MC starch grains are
amongst the oldest recovered from this genus, and with exisitng
the evidence species assessment is inconclusive at this time. An
extensive comparison is needed with other species in order to be
able to assess a precise taxonomical identification of the Dioscorea
starch grains recovered from the archaeological tools. Dioscorea
spp. pollen grains were identified in the column samples from sites
at El Jazmín, Campoalegre and Guayabito (Aceituno, 2002), indi-
cating the local availability of one still unidentified species that was
58 F.J. Aceituno, N. Loaiza / Quaternary Science Reviews 86 (2014) 49e62
Fig. 8. Starch grains Manihot spp. a) El Jazmín (265); b) El Jazmin (514); c) La Selva (90); def) modern Manihot esculenta Crantz (code Larq 2).
processed with stone tools by MC inhabitants. As with the beans,
we can only say that at least one species of Dioscorea was being
ground and chopped with stone tools around 7600 BP in the MC.
To determine if a particular starch grain can be identified as M.
esculenta Crantz, the domesticated variety of this genus, several
attributes need to be analyzed (Piperno, 2006). Comparatively,
domesticated Manihot starch grains tend to be larger in general
size, with an open and centric hilum, and stellated, Y-shaped and
cross-shaped fissures (Piperno, 2006, p.53). A highly diagnostic
feature for the domesticated variety is a higher frequency of pres-
sure facets on the distal end (Piperno, 2006, p.57). Wild varieties
tend to be smaller, with narrower pressure facets, lower percentage
of fissures (especially stellate-shaped fissure), and the hilum is
usually eccentric and closed (Piperno, 2006, p.53). Five of the MC
Manihot spp. starch grains display the features described by
Piperno (2006) as M. esculenta Crantz, and were extracted from
three handstones from El Jazmin dated between 7600 and 5600 BP
(Table 2). The rest of the grains do not display all the required
features for the domesticated variety, but can still be classified as
belonging to the Manihot genus. The evidence for the use of this
genus in MC is also reinforced by the identification of pollen grains
at the Guayabito site (Aceituno, 2002). As with beans and yams we
cannot be certain of the exact taxonomic group the starch grains
belong to, but we can say that some type of Manihot was being
ground in the MC by the middle Holocene.
Pollen column samples from El Jazmin, Campoalegre, and
Guayabito suggest that prior to any human evidence there are high
frequencies of cool and humid climate taxa such as Podocarpus,
Junglans nigra, Quercus and Cyathacea present (Jaramillo and Mejía,
2000a, 2000b; Aceituno, 2001). These types of plants are related to
the cooler conditions of the final Pleistocene and the climatic
amelioration of the early Holocene (Marchant et al., 2002). During
the early and middle Holocene, when there are clear human oc-
cupations, there are no strong indicators of deforestation or forest
clearance, but there are indicators of forest disturbances in the form
of pioneer elements in the pollen record represented by several
families such as Araceae, Melastomataceae, Dioscoraceae, Sapin-
daceae, Cecropiaceae, Rubiaceae, and grasses (i.e. Andropogon spp.).
 F.J. Aceituno, N. Loaiza / Quaternary Science Reviews 86 (2014) 49e62
Nowadays, all these taxa are very common in MC human disturbed
areas such as cultivation plots, roads, and rural living areas (Mer-
cado 2009, per. Com).
Also in the pollen record, we identified several genera with
edible species, such as Xanthosoma, Dioscorea and Manihot
(Aceituno, 2002). Xanthosoma appears in El Jazmin’s pollen record
by ca 9000 BP, strongly increasing towards ca 5000 BP; Dioscorea
pollen has been identified at the El Jazmin, Campoalegre and
Guayabito sites during the early Holocene (from ca 9000e7600 BP)
(Aceituno, 2002). Manihot spp. pollen was recovered from Guayabito
dating 4180 BP (Aceituno, 2001, 2002). It is clear that the sole
presence of these genera in the pollen record is not a clear indicator
of plant use, but for the case of Dioscorea and Manihot there is also
the starch grain evidence that reinforces the argument that at least
one species of each one of this genera was present in MC. Other
plants identified in the pollen record that have potential human use
for early settlers include palms (Bactris, Geonoma, Astrocaryum,
Socratea) (Jaramillo and Mejía, 2000a, 2000b) and for Guayabito site
Passiflora spp. with high frequencies at 4180 BP (Aceituno, 2001).
Z. mays pollen was identified at El Jazmin and dated between
ca 7000 and 5000 BP. At Guayabito maize pollen was recovered in a
sample along with Manihot spp. pollen dating to 4180 BP (Aceituno
et al., 2001; Aceituno, 2002). Maize pollen coincides chronologi-
cally with the starch grains, suggesting that there is some form of
cultivation at least of this plant species as an economical strategy
starting at ca 7000 BP.
The palynological record suggests small alterations to the plant
coverage at nearby sites, nonetheless, this evidence is not strong
enough to determine if the alteration was due to intentional
clearing or if it was due to the incidental alteration by human
settlement in a forested area as in the case of the Nukak, Awa or
Hoti (Politis, 1996, 2007).
The only domesticated plant identified up to this point is maize,
with dates similar to those in Panama of 6680  90 BP (Dickau et al.,
2007) and Ecuador 6600 BP (Piperno, 2009), indicating the rapid
dispersion of this crop (Dickau et al., 2007). In the MC there is a
report of maize starch grains identified and dated ca 5600 BP
(Dickau, 2007, 2008). In neighboring regions there are reports of
maize pollen and starch grains such as in Porce (Central Cordillera)
where both were evidenced and dated ca 6000 BP (Castillo and
Aceituno, 2006), and in the Calima region (Occidental Cordillera)
were pollen was dated en ca 6680 BP (Monsalve, 1985). So far, with
the identification of starch grains and pollen of Z. mays from the
middle Holocene we can suggest that active plant cultivation was
taking place as a subsistence strategy, but in no way does it reflect
substant change in ecological, economical, or technological strate-
gies. Prior to this we can only suggest that the use of plants, likely
local plants, was part of the economic activities of the MC in-
habitants, and according to the data presented here and elsewhere
this kind of plant exploitation extends the Pleistocene/Holocene
transition.
In the case of Manihot type starch grains identified as M. escu-
lenta Crantz have been described for contexts in Panama and dated
between ca 7000 and ca 5000 BP (Piperno, 2006, p.60; Dickau et al.,
2007), with ranges similar to those of MC, but as mentioned before,
we are not yet certain if we are dealing with the domesticated
species. Manihot spp. pollen has been identified in the Abeja site
(Colombia, Caqueta River Basin-Greater Amazon Basin) 10 cm
below a layer dated 4696  40 BP (Piperno and Pearsall, 1998, p.
263). Also in Colombia, in the Central Cordillera (about 200 Km
north of MC) Manihot spp. pollen was recovered from the 021 Site
dated between ca 6500 and 6000 BP (Castillo and Aceituno, 2006).
In the Zaña Valley of Peru Manihot starch grains were identified in
Las Pircas Phase (9800/7800 BP) (Rossen and Dillehay, 1999;
Clement, 2010; Rossen, 2011, p. 184). Continental data for Manihot
59
are scarce in terms of determining when this plant was domesti-
cated as well as in determining possible dispersion routes into
Northern South America, given the fact that it was likely domesti-
cated in Southwest Brazil (Mato Grosso, Rondonia and Acre states)
in the transition zone between the savanna (i.e the Cerrado) and
the south of the Amazonia Basin (Olsen and Schall, 1999; Arroyo-
Kalin, 2010; Clement, 2010; Isendahl, 2011). Panamanian, Colom-
bian and Peruvian data suggest that the domestication of this
important crop happened prior to ca7500 BP in the Brazilian region
mentioned above (Clement, 2010).
Evidence from Middle Cauca suggests that human groups
exploited and manipulated plants since the Pleistocene/Holocene
transition. The development of horticulture is suggested by the
presence of maize, and may be manioc, since the middle Holocene.
Nonetheless, we know little so far about production intensification
and preparation of cultivation plots, mainly because most of the
record comes from habitation sites. Pollen records suggest plant
coverage alteration due to human activities but, as mentioned
above, we have yet to assess if it is associated with gardening, plot
preparation for camps, or both activities.
6. Conclusions
The data presented in this paper are important to understand
early human settling of the humid montane forests. The archaeo-
logical evidence, along with 14C dating suggests the importance of
MC in the inhabiting of northern Andes since the Pleistocene/Ho-
locene transition (Aceituno et al., 2013). Relatively stable environ-
mental conditions, a soil enriched with volcanic ash, and access to
different life zones and altitudes, made MC a region with ideal
conditions for long term settlement of preceramic hunteregath-
erers groups.
Lithic technology from MC is oriented to plant usage. Hand-
stones, grinding bases and axes/waisted hoes are associated to the
earliest dates around 10,000 BP and remain present in the
archaeological record up to Late Holocene (Aceituno and Loaiza,
2007). The recovery of starch grains from the edges of lithic tools
indicates both, the use of plants and the importance of a
carbohydrate-rich diet, a basic requirement for the development of
human populations in tropical environments (Piperno and Pearsall,
1998, p.53e54). Several authors have stated the importance of this
kind of evidence for the study of plant usage in tropical environ-
ments where local conditions do not favor the preservation of
organic remains (Piperno and Pearsall, 1998; Perry, 2002, 2004;
Haslam, 2004; Piperno, 2006, 2009; Balter, 2007; Dickau et al.,
2007; Holst et al., 2007).
The four genera of plants identified using starch grain analysis
(Phaseolus, Dioscorea, Zea and Manihot) suggest plant use during
the early and middle Holocene. However, we have two main
unresolved issues regarding this genera, 1- at this point we’re
unable to determine if the Phaseolus, Dioscorea, and some of the
Manihot starch grains are local or foreign plants, 2- we cannot
assess if Phaseolus and Dioscorea were collected or cultivated. For
Phaseolus we have stated the difficulty to differentiate between
wild and domesticated species using starch grains. Based on the
age of the remains and the location within the tropics we
strongly suggest that they do not represent ancestral forms of
domesticated species. For Dioscorea we need to increase our
starch grain reference collection to determine its status, but also
compare it to regional species. The identification of Z. mays
starch grains and pollen is important because it indicates the
presence of this important crop in MC starting at ca 7000 BP and
during middle Holocene. Some of the Manihot starch grains have
clear indicators of belonging to the domesticated species while
others to wild species.
60 F.J. Aceituno, N. Loaiza / Quaternary Science Reviews 86 (2014) 49e62
Middle Cauca data support the idea that plant resources played
a key role in human economies in the Neotropics since the Pleis-
tocene/Holocene transition (Piperno, 2011). Nonetheless, the cur-
rent availability of microbotanical data do not allow us to conclude
if the plant genera used before the middle Holocene were being
cultivated, as it has been argued for Panama, Ecuador, and Peru,
where multiproxy evidences suggest systematic cultivation of small
plots from ca 9000 BP (Piperno and Pearsall, 1998, p.4; Piperno,
2011). The introduction of the foreign species maize and manioc
from ca 7000 BP clearly suggests that cultivation became a part of
the local strategies of food production for the inhabitants of MC
since the middle Holocene.
Finally it is important to state the important role played by sub-
Andean valleys in the rapid dispersion of cultigens across the
Neotropics, an idea that has also been suggested for other envi-
ronments based on data from Panama, Ecuador, and Peru (Piperno
and Pearsall, 1998; Rossen and Dillehay, 1999; Piperno, 2006, 2009;
Dickau et al., 2007; Piperno and Dillehay, 2008; Rossen, 2011). The
Middle Cauca River Valley of Colombia is a promising region for
studying the early use, cultivation, dispersal, and possibly the
domestication of plants because of the amount of sites identified
with Pleistocene/Holocene transition deposits containing both
microbotanical remains and a stone tool technology directed to
plant use in good stratigraphic contexts.
Acknowledgments
We want to thank the anonymous reviewers for the com-
ments and kind suggestions. This research project was financed
by the Vicerrectoría de Investigación from the Universidad de
Antioquia (UdeA - CODI) and by the Instituto Colombiano de
Antropología e Historia (ICANH). Thanks to all the people that
helped in this research project. Nicolás Loaiza wishes to thank
the Wenner-Gren Foundation for the Wadsworth International
Fellowship awarded to him from 2006 to 2009, Colciencias for
the Francisco Jose de Caldas Fellowship awarded in 2011, as well
Temple University for its support. Thank you to Ruth Dickau for
comments on starch identifications and suggestions on earlier
drafts of the manuscript.
References
Aceituno, F.J., 2001. Ocupaciones tempranas del bosque tropical subandino en la
Cordillera Centro-Occidental de Colombia. Unpublished PhD dissertation. Fac-
ultad de Geografía e Historia, Universidad Complutense de Madrid, Madrid.
Aceituno, F.J., 2002. Interacciones fitoculturales en el Cauca medio durante el Hol-
oceno temprano y medio. Arqueol. Area Intermed. 4, 89e113.
Aceituno, F.J., 2007. Poblamiento y variaciones culturales en la región Andina del
noroccidente de Suramérica en la transición Pleistoceno/Holoceno. In:
Bayón, C., Pupio, A., González, I., Flegenheimer, N., Frére, M. (Eds.), Arqueología
de las Pampas. Sociedad Argentina de Antropología, Buenos Aires, pp. 15e38.
Aceituno, F.J., Castillo, N., 2005. Strategies of Mobility in the Middle Range of
Colombia. Before Farming 2005/2 article 2.
Aceituno, F.J., 2010. Nuevos datos para la arqueología temprana del Cauca Medio: La
Pochola, un sitio precerámico en la Cuenca del río San Eugenio. Cordillera
Central (Colombia). Universidad de Antioquia (Unpublished manuscript).
Aceituno, F.J., Lalinde, V., 2011. Residuos de almidones y el uso de plantas durante el
Holoceno medio en el Cauca medio. Caldasia 33 (1), 1e20.
Aceituno, F.J., Loaiza, N., 2006. Una aproximación ecológica al poblamiento del
Cauca medio entre el Pleistoceno final y el Holoceno medio. In: López, C.,
Cano, M. (Eds.), Cambios ambientales en perspectiva histórica: Ecorregión Eje
Cafetero, vol. II. Universidad Tecnológica de Pereira, Pereira (Colombia),
pp. 42e55.
Aceituno, F.J., Loaiza, N., 2007. Domesticación del bosque en el Cauca Medio
colombiano entre el Pleistoceno Final y el Holoceno Medio. British Archaeo-
logical Reports, International Series 1654. Archaeopress, Oxford.
Aceituno, F.J., Loaiza, N., 2008. Rastreando los orígenes de la agricultura en la ver-
tiente oriental del Cauca medio. In: López, C., Ospina, G. (Eds.), Ecología His-
tórica: Interacciones Sociedad Ambiente a Distintas Escalas Socio Temporales.
Universidad Tecnológica de Pereira-Universidad del Cauca-Sociedad Colombi-
ana de Arqueología, Pereira, pp. 68e73.
Aceituno, F.J., Loaiza, N., 2010. Estructura interna y movilidad en el valle del río San
Eugenio en la Cordillera Central de Colombia. Arqueol. Area Intermed. 8, 83e
120.
Aceituno, F.J., Loaiza, N., Delgado, M.E., Barrientos, G., 2013. The initial human
settlement of Northwest south America during the Pleistocene/Holocene
transition: synthesis and perspectives. Quat. Int. 301, 23e33.
Aceituno, F.J., Treserras, J.J., Jaramillo, A., Loaiza, N., Vélez, L., 2001. Identificación de
plantas alimenticias en el Cauca medio durante el Holoceno temprano y medio.
Bol. Antropol. 15 (32), 51e72.
Ardila, G., Politis, G., 1989. Nuevo datos para un viejo problema. Bol. Mus. Oro 23, 3e45.
Arroyo-Kalin, M., 2010. The Amazonian Formative: crop domestication and
anthropogenic soils. Diversity 2, 473e504. Doi:10.3390/d2040473.
Babot, P., 2007. Granos de almidón en contextos arqueológicos: posibilidades y
perspectivas a partir de casos del Noroeste Argentino. In: Marconetto, B.,
Babot, P., Oliszewski, N. (Eds.), Paleoetnobotánica del Cono Sur. Estudios de
casos y propuestas metodológicas. Ferreyra Editor, Córdoba, pp. 95e125.
Bailey, R., Head, G., Jenike, M., Owen, B., Rechtman, R., Zechenter, E., 1989.
Hunting and gathering in tropical rain forest: is it possible? Am. Anthropol.
91, 59e81.
Barse, W.P., 1995. El período arcaico en el Orinoco y su contexto en el norte de Sud
América. In: Cavelier, I., Mora, S. (Eds.), Ambito y ocupaciones tempranas de la
América Tropical. Fundación Erigaie- Instituto Colombiano de Antropología,
Bogotá, pp. 99e114.
Balter, M., 2007. Seeking agriculture’s ancients roots. Science 316, 1830e1835.
Bauchet, S., McKey, D., De Garine, I., 1991. Wild yams revisited: is independence
from agriculture possible for rain forest hunteregatherers? Hum. Ecol. 19 (2),
213e243.
Bonomo, M., Aceituno, F.J., Politis, G., Pochettino, M.L., 2011. Pre-hispanic horticul-
ture in the Paraná Delta (Argentina): archaeological and historical evidence.
World Archaeol. 43 (4), 554e575.
Bruhns, K., 1994. Ancient South America. Cambridge World Archaeology. Cambridge
University Press.
Brücher, H., 1989. Useful Plant of Neotropical Origin and Their Wild Relatives.
Springer, Berlin.
Cano, M., 2001. Reconocimientos arqueológicos en los municipios de Marsella,
Dosquebradas y Pereira (Risaralda). In: González, V., Barragán, C. (Eds.),
Arqueología preventiva en el eje cafetero. Reconocimiento y rescate arqueoló-
gico en los municipios jurisdicción del fondo para la reconstrucción del eje
cafetero, FOREC. Serie ordenamiento territorial y reconstrucción del eje cafe-
tero, Patrimonio arqueológico en el ordenamiento territorial, vol. XI. ICANH,
Bogotá, pp. 39e50.
Cano, M., 2004. Los primeros habitantes de las cuencas medias de los ríos Otún y
Consota. In: López, C., Cano, M. (Eds.), Cambios ambientales en perspectiva
histórica ecorregión del eje cafetero, vol. I. Universidad Tecnológica de Pereira,
Programa Ambiental GTZ, Pereira, pp. 68e91.
Cano, M., 2008. Evidencias precerámicas en el municipio de Pereira: efectos del
vulcanismo y colonización temprana de los bosques ecuatoriales en el abanico
fluvio-volcánico Pereira-Armenia. In: López, C., Ospina, G. (Eds.), Ecología His-
tórica: interacciones sociedad ambiente a distintas escalas sociotemporales.
Universidad Tecnológica de Pereria-Universidad del Cauca-Sociedad Colo2mbi-
ana de Arqueología, Pereira, pp. 84e89.
Castillo, N., Aceituno, F.J., 2006. El bosque domesticado, el bosque cultivado: Un
proceso milenario en el valle medio del rio Porce en el noroccidente Colom-
biano. Lat. Am. Antiq. 17, 561e578.
Cavelier, I., Rodríguez, C., Herrera, L., Morcote, G., Mora, S., 1995. No solo de la caza
vive el hombre: Ocupación del bosque amazónico, Holoceno temprano. In:
Cavelier, I., Mora, S. (Eds.), Ámbito y Ocupaciones Tempranas de la América
Tropical. Fundación Erigaie, Instituto Colombiano de Antropología, Bogotá,
pp. 27e44.
Chacón, M.I., Pickersgill, B., Debouck, D.G., 2005. Domestication patterns in com-
mon bean (Phaseolus vulgaris L.) and the origin of the mesoamerican and an-
dean cultivated races. Theor. Appl. Genet. 110, 432e444.
Chacon, M.I., 2009. Darwin y la domesticación de plantas en las Américas: el caso
del maíz y el fríjol. Acta Biol. Colomb. 14 (4s), 351e364.
Clement, C., 2010. Origin and domestication of native Amazonian crops. Diversity 2,
72e106.
Cooke, R., 2005. Prehistory of native Americans on the Central American land
bridge: colonization, dispersal and divergence. J. Archaeol. Res. 13 (2),
129e183.
Cooke, R., Ranere, A., 1992. Prehistoric human adaptations to the seasonally dry
forests of Panama. World Archaeol. 24 (1), 116e133.
Correal, G., 1979. Investigaciones arqueológicas en los abrigos rocosos de Nemocón
y Sueva. Fundación de Investigaciones Arqueológicas Nacionales, Banco de la
República, Bogotá.
Correal, G., 1986. Apuntes sobre el medio ambiente pleistocénico y el hombre
prehistórico en Colombia. In: Bryan, A.L. (Ed.), New Evidence for the Pleistocene
Peopling of the Americas. Center for the Study of the Early Man, University of
Main, Orono, pp. 115e131.
Correal, G., van der Hammen, T., 1977. Investigaciones Arqueológicas en los Abrigos
Rocosos del Tequendama. Biblioteca Banco Popular, Bogotá.
Diamond, J., 2002. Evolution, consequences and future of plant and animal
domestication. Nature 418 (6898), 700e707.
Dickau, R., 2005. Resource Use Crop Dispersals, and the Transition to Agriculture in
Prehistoric Panama: Evidence From Starch Grains and Macroremains. Doctoral
Dissertation. Temple University, Philadelphia.
 F.J. Aceituno, N. Loaiza / Quaternary Science Reviews 86 (2014) 49e62
Dickau, R., 2007. Uso de recursos y producción de alimentos durante el precerámico
temprano del oeste de Panamá: nueva evidencia de análisis de granos de
almidón. Rev. Arqueol. Área Intermed. 7, 71e88.
Dickau, R., 2008. El uso de maíz y cultígenos de raíces en el precerámica de Panamá
y Colombia: evidencia de almidones en sitios húmidos subtropicales pre-
montanos. In: López, C., Ospina, G. (Eds.), Ecología Histórica: Interacciones
sociedad ambiente a distintas escalas socio temporales. Universidad Tecno-
lógica de Pereira-Universidad del Cauca-Sociedad Colombiana de Arqueología,
Pereira, pp. 60e67.
Dickau, R., Ranere, A., Cooke, R., 2007. Starch grain evidence for the preceramic
dispersals of maize and root crops into tropical dry and humid forest of Panama.
PNAS 14 (9), 3651e3656.
Espinal, L.S., 1990. Zonas de vida de Colombia. Universidad Nacional de Colombia,
Medellín.
Gnecco, C., 2000. Ocupación temprana de bosques tropicales de montaña. Uni-
versidad del Cauca, Popayán.
Gnecco, C., 2003. Agrilocalities during the Pleistocene/Holocene transition in
Northern South America. In: Salemme, M., Flegenheimer, N., Miotti, L. (Eds.),
The South Winds Blow, Ancient Evidence of Paleo South Americans, Corvalis:
Center for the Study of the First Americans. Texas A and M University Press,
Texas, pp. 7e12.
Gnecco, C., Salgado, H., 1989. Adaptaciones precerámicas en el suroccidente de
Colombia. Bol. Mus. Oro 24, 35e55.
Haslam, M., 2004. The decomposition of starch grain in soils: implications for
archaeological residues analyses. J. Archaeol. Sci. 31 (12), 1715e1734.
Henry, A., Hudson, G., Holly, F., Piperno, D., 2009. Changes in starch grain mor-
phologies from cooking. J. Archaeol. Sci. 36, 915e922.
Hermelin, M., 2001. Las cenizas volcánicas en el paisaje del noroccidente de
Colombia. In: VVAA Suelos del eje cafetero. Facultad de Ciencias Ambientales-
Universidad Tecnológica de Pereira, Cooperación Alemana al Desarrollo, Per-
eira, pp. 16e25.
Herrera, L., Bray, W., Cardale, M., Botero, P., 1988. Nuevas fechas de radiocarbono
para el precerámico en la Cordillera Occidental de Colombia. In: Paper Pre-
sented at the 46th International Congress of Americanists, Amsterdam.
Herrera, L., Peña, O., Calderón, D., Mendoza, L., Sepúlveda, L., Peña, J., Rodríguez, D.,
Acosta, C., 2011. Informe bimestral de actividades (noviembreediciembre 2010).
Proyecto de Rescate Arqueológico Aerocafé, Manizales (Unpublished
manuscript).
Holst, I., Moreno, J.E., Piperno, D., 2007. Identification of teosinte, maize and Trip-
sacum in Mesoamerica by using pollen, starch grains and phitoliths. PNAS 104
(45), 17608e17613.
IGAC, 1998. Paisajes Fisiográficos. ORAM, Bogotá.
INCIVA, 1995e1996. Proyecto de rescate arqueológico, gasoducto de Occidente,
Mariquita-Yumbo. ECOPETROL, Bogotá (Unpublished manuscript).
INTEGRAL, 1997. Arqueología de rescate: vía alterna de la troncal de Occidente río
Campoalegre-Estadio Santa Rosa de Cabal. Medellín. INTEGRAL S.A., Ministerio
de Transporte, Instituto Nacional de Vías, Medellín (Unpublished manuscript).
Iriarte, J., 2009. Narrowing the gap: exploring the diversity of early food-production
economies in the Americas. Curr. Anthropol. 5 (5), 677e680.
Isendahl, C., 2011. The domestication and early spread of manioc (Manihot esculenta
Crantz): a brief synthesis. Lat. Am. Antiq. 22 (4), 452e468.
Jaramillo, A., Mejía, J.C., 2000a. Análisis palinológico de los yacimientos el Jaz-
mín y Guayabito. Departamento de Risaralda, Medellín (Unpublished
manuscript).
Jaramillo, A., Mejía, J.C., 2000b. Análisis palinológico del yacimiento Campoalegre.
Departamento de Risaralda. Informe inédito, Medellín (Unpublished
manuscript).
Loy, T., 1994. Methods in the analysis of starch residues on prehistoric stone tools.
In: Hather, J.G. (Ed.), Tropical Archaeobotany Applications and New De-
velopments. One World Archaeology. Routledge, London, pp. 86e111.
Lynch, T., Pollock, S., 1981. La arqueología de la cueva Negra de Chobshi. Misc.
Antropol. Ecuat. 1, 92e119.
Marchant, R., Behling, H., Berrío, J.C., Cleef, A., Duivenvoorden, J., Hooghiemstra, H.,
Kuhry, P., Melief, R., Schreve-Brinkman, E., van Geel, B., van der Hammen, T., van
Reenen, G., Wille, M., 2002. Pollen-based biome reconstructions for Colombia at
3000, 6000, 9000, 12000, 15,000 and 18,000 14C yr ago: Late Quaternary
tropical vegetation dynamics. J. Quat. Sci. 17, 113e129.
Marchant, R., Boom, A., Behling, H., Hooghiemstra, H., Melief, B., van Geel, B., van
der Hammen, T., Wille, M., 2004. Colombian vegetation at the Last Glacial
Maximum: a comparison of model and pollen-based biome reconstructions.
J. Quat. Sci. 19, 721e732.
Matsuoko, Y., Vigouroux, Y., Goodman, M.M., Sanchez, J., Buckler, E., Doebley, J.,
2002. A single domestication for maize shown by multilocus microsatellite
genotyping. Proc. Nat. Acad. Sci. 99 (9), 6080e6084.
Mora, S., 2003. Early Inhabitants of the Amazonian Tropical Rain Forest a Study of
Humans and Environmental Dynamics. University of Pittsburgh, Department of
Anthropology, Pittsburg. University of Pittsburgh Latin American Archaeology
Reports No 3.
Monsalve, J.G., 1985. A Pollen Core From the Hacienda Lusitania, 4. Pro Calima,
pp. 40e44.
Morcote, G., Cabrera, G., Mahecha, D., Franky, C., Cavelier, I., 1998. Las palmas entre
los grupos cazadores-recolectores de la amazonía colombiana. Caldasia 20 (1),
57e74.
Olsen, K.M., Schall, B., 1999. Evidence of origin of cassava: phylogeography of
Manihot esculenta. PNAS 96, 5586e5591.
61
Orozco, J.I., 2001. Las cenizas volcánicas en el territorio de Pereira y sus alrededores.
In: VVAA Suelos del eje cafetero. Facultad de Ciencias Ambientales-Universidad
Tecnológica de Pereira, Cooperación Alemana al Desarrollo, Pereira, pp. 9e15.
Otero, H., Santos, G., 2006. Las ocupaciones prehispánicas del cañón del río Porce.
Prospección rescate y monitoreo arqueológico. Proyecto hidroeléctrico Porce III
e Obras de Infraestructura. Universidad de Antioquia-Empresas Públicas de
Medellín, Medellín (Unpublished manuscript).
Pagan, J.R., Rodríguez, M.A., Chanlatte, L.A., Narganes, Y., 2005. La temprana intro-
ducción y uso de algunas plantas domesticas, silvestres y cultivos en las Antillas
precolombinas. Una primera revaloración desde la perspectiva del arcaico de
Vieques y Puerto Rico. Diál. Antropol. 10, 7e33.
Pearsall, D.M., 1994. Investigating New World tropical agriculture: contributions
from phytolith analysis. In: Hather, J.G. (Ed.), Tropical Archaeobotany Applica-
tions and New Developments. One World Archaeology. Routledge, London,
pp. 115e138.
Pearsall, D.M., Piperno, D.R., 1990. Antiquity of maize cultivation in Ecuador:
summary and reevaluation of the evidence. Am. Antiq. 55 (2), 324e337.
Pearsall, D.M., Chandler-Ezell, K., Zeidler, J.A., 2004. Maize in ancient Ecuador: re-
sults of residue analysis of stone tools from the Real Alto site. J. Archaeol. Sci. 31,
423e442.
Perry, L., 2002. Starch granule size and the domestication of Manioc (Manihot
esculenta) and sweet potato (Ipomea batatas). Econ. Bot. 56 (4), 335e349.
Perry, L., 2004. Starch analyses reveal the relationship between tool type and
function: an example from Orinoco valley of Venezuela. J. Archaeol. Sci. 31,
1069e1081.
Perry, L., Flannery, K., 2007. Precolumbian use of chili peppers in the valley of
Oaxaca, México. PNAS 104 (29), 11905e11909.
Piperno, D.R., 2006. Identifying manioc (Manihot esculenta Crantz) and other crops
in Pre-Columbian tropical America through starch grain analysis a case study
from Central Panama. In: Melinda A. Zeder, M.A., Bradley, D.G., Emshwille, E.,
Smith, B.D. (Eds.), Documenting Domestication New Genetic and Archaeological
Paradigms. University of California Press, Berkeley, pp. 46e67.
Piperno, D.R., 2009. Crop plants with phytoliths (and starch grains) in Central and
South America: a review and an update of the evidence. Quat. Int. 193, 146e159.
Piperno, D.R., 2011. The origins of plant cultivation and domestication in the new
world tropics. Curr. Anthropol. 52 (4), 453e470.
Piperno, D.R., Dillehay, T., 2008. Starch grains on human teeth reveal early broad
crop diet in northern Peru. PNAS 105 (50), 19622e19627.
Piperno, D.R., Holst, I., 1998. The presence of starch grains on prehistoric stone tools
from de humid Neotropics: indications of early tuber use and agriculture en
Panamá. J. Archaeol. Sci. 25, 765e776.
Piperno, D., Moreno, J.E., Iriarte, J., Holst, I., Lachniet, M., Jones, J.G., Ranere, A.J.,
Castanzo, R., 2007. Late Pleistocene and Holocene environmental history of
the Iguala valley, Central Balsas watershed of Mexico. PNAS 104 (29), 11874e
11881.
Piperno, D.R., Pearsall, D., 1998. The Origins of Agriculture in the Lowland Neo-
tropics. Academic Press, San Diego.
Piperno, D.R., Stothert, K.E., 2003. Phytolith evidence for early Holocene Cucurbita
domestication in Southwest Ecuador. Science 299, 1054e1057.
Piperno, D.R., Bush, M., Colinvaux, P., 1991. Paleoecological perspectives on human
adaptation in Central Panama in the Pleistocene. Geoarchaeology 6, 210e226.
Piperno, D.R., Ranere, A.J., Holst, I., Hansell, P.K., 2000. Starch grains reveal early root
crop horticulture in the Panamanian tropical forest. Nature 407, 894e897.
Piperno, D.R., Ranere, A.J., Holst, I., Iriarte, J., Dickau, R., 2009. Starch grain and
phytolith evidence for early ninth millennium B.P. maize from the central Balsas
river valley, Mexico. PNAS 106 (13), 5019e5024.
Politis, G., 1996. Moving to produce: Nukak mobility and settlement patterns in
Amazonia. World Archaeol. 27 (3), 492e511.
Politis, G., 2007. Nukak: Ethnoarchaeology of an Endangered Amazonian People.
Left Coast, California.
Ranere, A.J., 1980. Preceramic shelters in the Talamancan range. In: Linares, O.,
Ranere, A.J. (Eds.), Adaptive Radiations in Prehistoric Panama, Peabody Museum
Monographs, 5. Harvard University, Cambridge Massachusetts, pp. 16e43.
Ranere, A.J., 2008. Lower Central America. In: Pearsall, D.M. (Ed.), Encyclopedia of
Archaeology. Academic Press, New York, pp. 192e209.
Ranere, A.J., Cooke, R., 1995. Evidencias de ocupación humana en Panamá a post-
rimerías del Pleistoceno y a comienzos del Holoceno. In: Cavelier, I., Mora, S.
(Eds.), Ámbito y Ocupaciones Tempranas de la América Tropical. Fundación
Erigaie, Instituto Colombiano de Antropología, Bogotá, pp. 5e26.
Ranere, A.J., Cooke, R., 2003. Late Glacial and early Holocene occupation of Central
American tropical forests. In: Mercader, J. (Ed.), Under the Canopy: the
Archaeology of Tropical Rain Forests. Rutgers University Press, New Brunswick,
NJ, pp. 219e248.
Ranere, A.J., López, C.E., 2007. Cultural diversity in Late Pleistocene/Early Holocene
populations in Northwest South America and Lower Central America. Int. J.
South Am. Archaeol. 1, 25e31.
Reichel-Dolmatoff, G., 1997. [1965]. Arqueología de Colombia: un texto intro-
ductorio, second ed. Biblioteca familiar colombiana. Presidencia de la República,
Bogotá.
Restrepo, C.A., 2013. Informe de Monitoreo Arqueológico en el área de influencia del
Proyecto de Desarrollo Vial Armenia Pereira Manizales, Autopista de Café.
Instituto Nacional de Vías. Autopistas de Café S. A. Documento elaborado para
INCO-AUTOPISTAS DEL CAFÉ S.A. (Unpublished manuscript).
Rodríguez, C., 1995. Asentamientos de los bosques subandinos durante el Holoceno
medio. In: Cavelier, I., Mora, S. (Eds.), Ámbito y Ocupaciones Tempranas de la
62 F.J. Aceituno, N. Loaiza / Quaternary Science Reviews 86 (2014) 49e62
América Tropical. Fundación Erigaie, Instituto Colombiano de Antropología,
Bogotá, pp. 115e123.
Rossen, J., 1998. Unifaces in early Andean cultura history: the Nanchoc lithic
tradition of northern Perú. Andean Past 5, 241e299.
Rossen, J., 2011. Preceramic plant gathering, gardening, and farming. In: Dillehay, T.
(Ed.), From Foraging to Farming in the Andes: New Perspectives on Food Production
and Social Organization. Cambridge University Press, Cambridge, pp. 177e192.
Rossen, J., Dillehay, T., 1999. La colonización y el asentamiento del norte de Perú:
innovación, tecnología y adaptación en el valle de Zaña. Bol. Arqueol. PUCP 3,
121e139.
Rumold, C.U., 2010. Illuminating Women’s Work and the Advent of Plant Cultivation
in the Highland Titicaca Basin of South America: New Evidence from Grinding
Tool and Starch Grain Analysis. Unpublished PhD dissertation. University of
California, Santa Barbara.
Salgado, H., 1988e1990. Asentamientos precerámicos en el alto medio río Calima,
cordillera Occidental, Colombia. Cespedesia 57e58, 139e162.
Salomons, J.B., 1989. Paleoecology of volcanicsoils in the colombian Central
Cordillera (Parque Nacional de los Nevados). In: Van der Hammen, T., Díaz-
Piedrahita, S., Alvarez, V.J. (Eds.), Studies on Tropical Andean Ecosystems, La
Cordillera Central colombiana transecto Parque de los Nevados, vol. 3. J. Cramer,
Vaduz, pp. 15e216.
Santos, G., 2008. Cazadores-Recolectores y horticultores del Holoceno temprano y
medio en la cuenca baja del Porce. In: López, C., Ospina, G. (Eds.), Ecología
Histórica: Interacciones Sociedad Ambiente a Distintas Escalas Socio Tempo-
rales. Universidad Tecnológica de Pereira-Universidad del Cauca-Sociedad
Colombiana de Arqueología, Pereira, pp. 74e77.
Stothert, K., 1985. The preceramic Las Vegas culture of coastal Ecuador. Am. Antiq.
50 (3), 613e637.
Stothert, K.E., Piperno, D.R., Andres, T.C., 2003. Terminal Pleistocene/Early Holocene
human adaptation in coastal Ecuador: the Las Vegas evidence. Quat. Int. 109e
110, 23e43.
Tabares, D., Rojas, E., 2000. Aportes para una historia en construcción: arqueología
de rescate en la doble calzada Manizales-Pereira-Armenia. INVIAS-CISAN,
Bogotá (Unpublished).
Tabares, D., 2004. Fase I: Prospección río Campoalegre, mundo arcaico en la región
del Cauca medio, Colombia. Fundación de Investigaciones Arqueológicas
Nacionales, Bogotá (unpublished).
Temme, M., 1982. Excavaciones en el sitio precerámico de Cubilán (Ecuador). Misc.
Antropol. Ecuat. 2, 135e164.
Tistl, M., 2006. La formación geológica del paisaje en el piedemonte del eje cafetero
colombiano. In: López, C., Cano, M. (Eds.), Cambios ambientales en perspectiva
histórica: Ecorregión Eje Cafetero, vol. II. Universidad Tecnológica de Pereira,
Pereira (Colombia), pp. 77e92.
Tobon, J.I., Perez, N., 2005. Análisis mineralógico y textural, sitios La Selva y El
Jazmin (Unpublished report).
Tobon, J.I., Castaño, C.P., 2006. Caracterización mineralógica de los suelos de San
Germán y La Pochola (Unpublished report).
Toro, G., Hermelín, M., Poupeau, G., 2001. Depósitos de los últimos 40.000 años BP
en el departamento de Risaralda, Colombia. In: VVAA Suelos del eje cafetero.
Facultad de Ciencias Ambientales-Universidad Tecnológica de Pereira, Cooper-
ación Alemana al Desarrollo, Pereira, pp. 26e31.
Torrence, R., Barton, H., 2006. Looking ahead. In: Torrence, R., Barton, H. (Eds.),
Ancient Starch Research. Left Coast Press, Walnut Creek, pp. 217e223.
Zarrillo, S., Pearsall, D.M., Raymond, J.S., Tisdale, M.A., Quon, D.J., 2008. Directly
dated starch residues document early formative maize (Zea mays L.) in tropical
Ecuador. PNAS 105 (13), 5006e5011.