Journal of Anthropological Archaeology 49 (2018) 161–172

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Journal of Anthropological Archaeology

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

The origins and early development of plant food production and farming in T
Colombian tropical forests
⁎
Francisco J. Aceitunoa, , Nicolás Loaizaa,b,c
a
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
c
ISA-INTERCOLOMBIA, Colombia

A R T I C L E I N F O A B S T R A C T

Keywords: This paper concentrates on archaeobotanical evidence for the adoption of plant cultivation in the forests in seven
Colombia regions of Colombia. We present a synthesis and explanation of the evidence we currently have for the process
Food production that involved the adoption of plant cultivation and the development of food production in this area. The use of
Plant cultivation locally available plant foods in these forests is evident by the Pleistocene/Holocene transition. By the Middle
Domestication
Holocene, exogenous plant domesticates were added, including maize, manioc, and possibly common beans. We
further explore available data on other proxies to discuss models to explain the transition from hunting and
gathering to horticulture.

1. Introduction
The origin of food production is one of the major landmarks in
human evolution (Watson, 2001; Winterhalder and Kennett, 2006).
Plants were domesticated in multiple regions around the globe, be-
tween five and ten, depending on the author (e.g. Harlan, 1971;
Vavilov, 1992: 126–128; Piperno and Pearsall, 1998; Piperno, 2011a,b;
Smith, 1998; Diamond, 2002; Zeder et al., 2006; Balter, 2007). One of
these regions – northwestern South America (Andean Colombia and
adjacent Ecuador) – is a belt of tropical forests with dissimilar tem-
perature and precipitation regimes. Piperno (2011a,b) has proposed
that this region was a center for plant domestication in South America.
Synergisms between socio-cultural, climatic, and environmental pro-
cesses at the late Pleistocene/early Holocene boundary and subse-
quently in the early and mid-Holocene, favored situations that en-
hanced the subsistence value of planting groups of vegetable foods,
rather than gathering them off the landscape (Gnecco, 2003; Gnecco
and Aceituno, 2004, 2006; Aceituno and Loaiza, 2014, 2015; Loaiza
and Aceituno, 2015; Morcote et al., 2014; Piperno, 2011a,b, 2017;
Dickau et al., 2015; Loaiza and Aceituno, 2015; Piperno, 1989; Santos
et al., 2015).
A growing number of taxonomic and genetic studies in different
parts of the lowland and premontane forests in the Neotropics show
that important food plants such as manioc (Manihot esculenta Crantz),
arrowroot (Maranta arundinacea L), cocoyam (Xanthosoma sagittifolium
(L) Schott), leren (Calathea allouia Lindl) and sweet potato (Ipomoea
batatas (L) Lam) were domesticated in different regions (e.g. Brücher,
1989; Piperno and Pearsall, 1998: 4–6; Iriarte, 2007, 2009; Rouiller
et al., 2013). Archaeological data recovered over the past three decades
have indicated that Colombia is a key region for understanding the
origin and dispersal of plant cultivation and domestication, as well as
their cultural evolutionary consequences throughout the Neotropics
(Cardale et al., 1989; Gnecco and Salgado, 1989; Gnecco and Mora,
1997; Morcote et al., 1998, 2014; Gnecco, 2000, 2003; Aceituno and
Loaiza, 2014; Santos et al., 2015). We present, analyze and discuss
several lines of information that elucidate the range of adaptive stra-
tegies adopted by people living in several regions of Colombian humid
forests. We show how they developed strategies during the early Ho-
locene and the middle Holocene, setting the scene for the transition
from lifeways combining hunting, fishing, and gathering to others that
increasingly relied upon a small number of domesticated vegetable
foods.
Human groups present on the Colombian landscape since at least
12,400 ± 160, years BP (Correal, 1986: 117) (all dates are un-
calibrated 14C, unless noted otherwise) responded to the strong en-
vironmental changes of this long period by developing plant manage-
ment practices that modified the regional and local vegetation and
included the use of cultigens whose initial domestication occurred
outside and even well beyond Colombia, such as manioc and maize
(Piperno, 2011a,b; 2017). These management strategies are the step-
pingstones for the origins of food production and signal the beginning
of the Archaic, a Holocene cultural period in which the development of

⁎
Corresponding author.
E-mail addresses: francisco.aceituno@udea.edu.co (F.J. Aceituno), nloaiza@temple.edu (N. Loaiza).

https://doi.org/10.1016/j.jaa.2017.12.007
Received 17 January 2017; Received in revised form 5 December 2017
0278-4165/ © 2018 Elsevier Inc. All rights reserved.
F.J. Aceituno, N. Loaiza
plant domestication and cultivation as well as the insertion of horti-
culture as a principal player in the subsistence economy.
We use the expression food production in the sense of low-level food
production (Smith, 2001), defined as an economy that is a mixture of
practices characteristic of hunting and gathering and agricultural so-
cieties (Winterhalder and Kennett, 2006: 4). Horticulture should be
understood as a form of low-level food production that entails small-
scale planting of species that range from wild to domesticated (Piperno
and Pearsall, 1998: 6–7; Winterhalder and Kennett, 2006: 4). Following
these authors, domesticates are new varieties or species of plants or
animals created through artificial selection (Winterhalder and Kennett,
2006: 3). Domestication then is the process that leads to the evolution
by artificial selection of altered organisms that display features ad-
vantageous for human environments but that are disadvantageous for
surviving in the wild, and this last feature is also known as the do-
mestication syndrome (Allaby, 2014). Cultivation should be understood
as the “tending of plants, wild or domesticated” (Winterhalder and
Kennett, 2006: 4).
2. Archaeobotanical evidence of plant use during the Archaic
period
At the onset of the Holocene, field evidence shows a rise in regional
population as new areas were colonized (Aceituno et al., 2013). The
evidence to be discussed in this paper is located from south to north of
the Andean region of Colombia following the Cauca River Basin in
Popayan, Calima River Basin, middle Cauca River Basin (middle
Cauca), Medellin/Porce River Basin. Outside the Andes in the Amazon
basin the Peña Roja site is located. Finally we will refer to archae-
ological evidence in the Sabana de Bogota and middle Magdalena River
Basin (middle Magdalena) (Fig. 1).
2.1. Popayan
Popayan is located in the Central Cordillera in the upper Cauca
River Basin at about 1600 m above sea level (masl) in a Subandean
forest zone (Fig. 1) (Gnecco, 2000: 17). In this setting is located San
Isidro, a site dated between 10,050 ± 100 and 9530 ± 100 years BP
(Gnecco, 2000: 48, Gnecco, 2003) (Fig. 2) and characterized by thou-
sands of flaked chert and some obsidian artifacts There are retouched
and unretouched flakes, lanceolate bifaces and projectile preforms. In
addition to the knapped artifacts there are edge ground cobbles
(handstones), flat milling stones, cobbles with concave grooves, and a
ground stone axe (Gnecco, 2000: 60–62). Thousands of seeds were re-
covered in association with the artifacts. These included avocado
(Persea cf. americana), basul (Erythrina cf. edulis), Caryocar spp., Virola
spp., several palm types, highlighting Acrocomia (Piperno and Pearsall,
1998: 200; Gnecco, 2000: 67–69). Macrobotanical remains of Lagenaria
sp. were also recovered at San Isidro (Gnecco, 2003; Gnecco and
Aceituno, 2006: 93). Starch grains extracted from an edge ground
cobble were identified as cf. Xanthosoma/Ipomoea and/or Manihot and
Maranta (cf. arundinacea), as well as non-identified grasses and legumes
(Piperno and Pearsall, 1998: 200).
According to Gnecco (2000, 2003) San Isidro inhabitants were al-
ready practicing cultivation at the Pleistocene/Holocene transition. In
the pollen record the identification of colonizing plants (i.e. Plantago),
grasses and shrubs suggest plot preparation close to the site. Based on
the pollen and archaeobotanical records, Gnecco (2003) suggested that
San Isidro inhabitants practiced a form of agroecology (Rindos, 1984)
based on the selection and cultivation of non-local fruit trees and tubers
around 10,000 years BP.
2.2. Calima River Basin
North of Popayan in the Calima River Basin (Western Cordillera)
(Fig. 1) in a Subandean moist forest life zone and at 1750 masl, the sites
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Journal of Anthropological Archaeology 49 (2018) 161–172
of Sauzalito, El Recreo and El Pital are located. Radiocarbon dates from
these sites are between 9670 ± 100 years BP and 4090 ± 90 years BP
(Salgado, 1988–1990, Salgado, 1995) (Fig. 2). Stone tools found at
them, including milling stones and handstones, and axes/waisted hoes
suggest plant processing (Herrera et al., 1988; Salgado, 1988–1990).
The last-named is considered a diagnostic tool type for the region and
has been related to the removal of soils for plant cultivation, harvesting
tubers and roots, and the extraction of starchy hearths from palm trees
(Cardale et al., 1989; Gnecco and Salgado, 1989). From these sites ar-
chaeologists also recovered charred seeds from palm trees and avocado
along with phytoliths identified as palms, bamboos and arrowroot
Maranta sp. (Piperno, 1985; Piperno and Pearsall, 1998: 202).
It can be argued that the Calima River data revolutionized archae-
ological interpretations during the 1980s and 1990s) because they
suggested a model for early hunter-gatherers that stressed plants, as
opposed a heavy reliance on hunting as in the Sabana de Bogotá
(Gnecco and Salgado, 1989). This new model also put forward for the
first time the possibility of plant cultivation in early Holocene occu-
pations in Colombia.
2.3. Middle Cauca
Following the Cauca River Basin to the north, is found the middle
Cauca region (Central Cordillera) (Fig. 1). This region has one of the
most complete archaeological records in Colombia (26 sites excavated
and dated) and a long occupational sequence that dates back to the
Pleistocene/Holocene transition to the middle Holocene (Rojas and
Tabares, 2000; Rodríguez, 2002; Cano, 2004, 2008; Tabares, 2004;
Aceituno and Loaiza, 2007; Dickau et al., 2015). Most of these sites are
located between 1400 and 1700 masl in a premontane wet forest
(Espinal, 1985). Archaic occupations in the region are dated between
10,619 ± 66 and 4180 ± 70 years BP (Dickau et al., 2015) (Fig. 2).
The Middle Cauca lithic industry comprises flaked and ground tools.
Flaked tools were widely used, and some derived from a quartz quarry
site at El Antojo where thousands of flakes were collected, some of them
bifacially reduced (INTEGRAL, 1997; Aceituno and Loaiza, 2007:
77–78). Two bifacial projectile points: one at the site of El Mirador,
dated 9663 ± 83 years BP, and another at the 39 El Recreo Cancha site
dated between ca. 8500 and 8000 years BP (Herrera et al., 2011; Dickau
et al., 2015). Frequent handstones, milling stones, and waisted tools
probable axes/waisted hoes (Fig. 3: 1–3) were manufactured with local
volcanic rocks obtained in streams (Aceituno and Loaiza, 2007; 2014;
2015; Loaiza and Aceituno, 2015).
Macrobotanical remains are scarce in middle Cauca sites, and
therefore archaeobotanical studies have been based mainly on pollen
and starch grain analysis. Even so, the sheer quantity of data underlines
the importance of this region for the understanding the evolution of
food production in the northern Andes. Tables 1 and 2 show wild and
domesticated taxa identified in archaeological sites dated from the early
to the middle Holocene. During the early Holocene, microbotanical
data on plant use suggest the earliest stages of a form of low-level food
production that entailed the selection, propagation, and protection of
some plants in areas close to settlements. On the other hand, the exo-
genous origin of maize (Fig. 4: d and k), Manihot cf. esculenta (Fig. 4: a,
e, m, and p) and Phaseolus cf. vulgaris (Fig. 4: l and n) confirms that
horticultural practices were well established by the middle Holocene
throughout the Neotropics. The remaining taxa identified are carbo-
hydrate rich tuberous plants, such as Xanthosoma (Fig. 4: f, h, and j), a
widespread genus across northern South America (Piperno and Pearsall,
1998: 165) that is well represented within the pollen record at the El
Jazmin site between ca. 9000 and 5000 years BP (Jaramillo and Mejía,
2000a; Aceituno and Loaiza, 2007: 84–86). That suggests these tubers
were likely a resource used since the first moments of food production.
Ipomoea (Fig. 4: g) – the genus to which the domesticated species I.
batatas (L) belongs – produces tuberous plants and has been identified
in the middle Cauca. Two wild possible ancestors of I. batatas, I. trifida
F.J. Aceituno, N. Loaiza
Fig. 1. Colombian geography and archaeological regions.
and I. triloba, are found in various regions of the Cordillera Central and
the Colombian Caribbean region (Roullier et al., 2013). Ipomoea cf
batatas has been reported at Middle Holocene in the Canaán site as it
was at the 39 El Recreo Cancha site (Dickau, 2008) (Table 2).
The Dioscorea situation (Fig. 4: c and o) is somewhat similar to that
of cocoyam. Tuber-producing yams have a wide distribution through
the Neotropics (Brücher, 1989: 19). Dioscorea pollen has been identified
at El Jazmin, Campoalegre and Guayabito sites (Jaramillo and Mejía,
2000a, 2000b; Aceituno, 2002), while starch grains from tools have
been recovered at La Pochola, El Jazmin and La Selva sites (Aceituno
and Loaiza, 2014). Of this genus only D. trifida was domesticated in the
Neotropics in the wide lowland area of northern Brazil, the Guyanas,
Suriname and southern Venezuela (Piperno and Pearsall, 1998: 117;
Piperno, 2011a). To date little is known of the domestication history of
this plant and we cannot assure if the starch grains identified belong to
a species ancestral to the domesticated D. trifida. However, it is likely
that the species used was part of the group of plants selected in the early
moments of food production.
With regard to common beans (Fabaceae: Phaseolus spp.) (Fig. 4: b
and i) we have been unable to determine if the archaeological speci-
mens are local or exotic (Aceituno and Loaiza, 2014). The early dates
suggest that it could be a local variety that remains unidentified but
that is not ancestral to the any of the domesticated species P. vulgaris
(common bean) and P. lunatus (lima bean) originated in Mexico and
Peru (Chacón, 2009; Chacón et al., 2005), around the same time the
local variety was being used in middle Cauca (Aceituno and Loaiza,
163
Journal of Anthropological Archaeology 49 (2018) 161–172
2014).
The pollen record corresponding to the early Holocene in the middle
Cauca region displays plant families typically associated with anthropic
alteration, such as Asteraceae, Melastomataceae, Poaceae and Cecropia
spp., among others (Jaramillo and Mejía, 2000a). Nonetheless, the
available data are not conclusive about the origin of this alteration and
we cannot assess if it was due to garden plot preparation, incidentally to
camp preparation as shown in ethnographic examples (e.g. Politis,
1995, 1996, 2007), natural causes or some combination of these sce-
narios.
As noted above, plant use in the middle Cauca increases by the mid-
Holocene. Towards. 7500/7000 years BP the pollen records from La
Pochola, El Jazmin and Campoalegre display clear evidence of vege-
tation alteration as suggested by the dramatic increase in the plant fa-
milies Melastomataceae and Asteraceae as well as the high taxa di-
versity displayed in the record. Associated with this increase the starch
grain evidence of Manihot cf. esculenta, maize and Phaseolus cf. vulgaris
strongly suggest the consolidation of a form of food production based
on horticulture.
Maize was not only identified through starch grains (Table 1), but
also from pollen, both at La Pochola dated 6743 ± 45 years BP
(Mercado, 2010) and El Jazmin dated between ca. 7000 and 5000 years
BP (Table 2) (Aceituno et al., 2001; Aceituno, 2002). Originally do-
mesticated in the Balsas River Basin of Mexico before 7900 years BP
(8700 cal. BP) (Matsuoko et al., 2002; Piperno et al., 2007, Piperno
et al., 2009). In any case, the incorporation of maize in the middle
F.J. Aceituno, N. Loaiza
Cauca to local practices played an important role in the consolidation of
horticultural strategies increasing the use of crops with higher return
rates in gardens near sites (Dickau, 2005; Aceituno and Loaiza, 2007,
2014).
Manihot spp. is another important taxa identified in the middle
Cauca region. Manioc was domesticated in Southwest Brazil in the
transitional zone between the Cerrado environment and the Amazon
Basin (Olsen and Schall, 1999; Arroyo-Kalin, 2010; Clement, 2010;
Fig. 3. Axes/Waisted Hoes: (1) El Jazmin Level 9 (code 433); (2) La Pochola level 16 (code 282); (3) El Jazmin (surface collection).
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Journal of Anthropological Archaeology 49 (2018) 161–172
Fig. 2. Radiocarbon dates from regions discussed.
Isendahl, 2011). The starch grains from the tools “mano 514” (older
than 7590 years BP), “mano 265” (dated 7080 ± 50 years BP) and
“mano” 762 (dated between 6903 ± 45 and 6743 ± 45 years BP)
(Table 2) display the distinctive characteristics of manioc (Aceituno and
Loaiza, 2014). Manihot spp. pollen was recovered from the Guayabito
site dated 4180 years BP (Table 2) (Jaramillo and Mejía, 2000b;
Aceituno, 2001, 2002; Aceituno et al., 2001) supporting the idea that
this plant was cultivated locally as early as the middle Holocene.
F.J. Aceituno, N. Loaiza Journal of Anthropological Archaeology 49 (2018) 161–172

Table 1
Starch grains recovered in middle Cauca sites.

14
Site Tool C YBP Xanthosoma spp. Fabaceae Phaseolus spp. Ipomoea spp. Zea mays Dioscorea spp. Manihot spp.

Pch Handst 55 8095 ± 55

Pch Handst 79 8095 ± 55 1
Pch Handst 238 9047 ± 45/6903 ± 45 4 1 1 1
Pch Handst 324 6903 ± 45 2
Pch Handst 458 6903 ± 45 7 1 10
Pch Handst 23 6903 ± 45/6743 ± 45 3 1
Pch Base 665 6903 ± 45/6743 ± 45 9 31 18 3
Pch Handst 762 6903 ± 45/6743 ± 45 4 3
Pch Handst 733 6903 ± 45/6743 ± 45 3
Pch Handst 236 6743 ± 45 2
Pch Handst 1012 6743 ± 45/5922 ± 51 1 2 1
Pch Handst 2168 6743 ± 45/5922 ± 51 1 1
Pch Handst 2154 5922 ± 51
Jaz Base 315 > 7590 5
Jaz Base 514 > 7590 2 1 5
Jaz Handst 265 7080 ± 50 1 5 1
Jaz Handst 830 5625 ± 50 1
Jaz Handst 202 5625 ± 50/7528 ± 51 4 1
La Selva Milling stone 39 8712 ± 60 2 1
La Selva Axe/hoe 90 > 8712 ± 60 1 2 5
La Selva Axe/hoe 281 8712 ± 60 1

Pch: Pochola; Jaz: El Jazmín; handst: handstone.

By ca. 6900–6700 years BP at La Pochola site a large number of and gathering (Johnson, 1983; Posey, 1984; Balée, 1989, 2006; Balée
starch grains display the typical characteristics of common beans and Gely, 1989; Andrade, 1993; Politis, 1996, 2007; Rival, 1998, 2006;
(Phaseolus vulgaris) (Table 1) (Fig. 4: l and n). It has been estimated that Balée and Erickson, 2006; Winterhalder and Kennett, 2006: 4).
by this time beans had been domesticated in the centers of origin and
that therefore there is a good chance that the starch grains from these
2.5. Medellin/Porce River Basin
levels belong to the domesticated species. Nonetheless this is hard to
prove because the starch grains of domesticated and wild varieties of
North of middle Cauca, the Medellin/Porce basin in the Central
common beans are similar and therefore hard to discriminate through
Cordillera has produced ten Archaic sites that display important evi-
starch grains (Piperno and Dillehay, 2008).
dence of food production. In the South of this region (Medellin), three
In summary, the data from middle Cauca suggest that plants began
sites, La Morena, La Blanquita, and Casa Blanca, are located between
to be manipulated by humans in the early Holocene setting the foun-
2000 and 2100 masl, in an Andean life zone (Santos, 2010). The other
dations for the development and establishment of horticulture by the
sites are located further North (Porce) in a tropical forest life zone just
middle Holocene in which tuberous plants played a key role (i.e. the
below 1000 masl and named PIIIOI-52, PIIIOI-59, PIIIOI-40, PIIIOP-61,
preparation of small gardens where wild and domesticated species are
PII-021, PII-045 and PII-107. Lithic technology is similar to that of the
cultivated) (Piperno and Pearsall, 1998: 7; Winterhalder and Kennett,
Calima and middle Cauca regions, comprising handstones, milling
2006: 4; Harris, 2007: 23–24). An important feature of horticulture in
stones, axes, hoes, and unifacial tools. Besides these kinds of tools, there
tropical environments is the high diversity of plants that are cultivated
are thousands of flintknapped quartz tools (such as knives and scrapers)
nearby habitation sites as an economic strategy that also entails hunting
that have been associated with hunting and butchering activities

Table 2
Middle Cauca archaeological sites with associated archaeobotanical evidence and dates.

14
Site C YBP Archaeobotanical evidence Reference

Jaz ca. 9000–5000 Xanthosoma (p); Dioscorea (p) Aceituno and Loaiza (2007)
Jaz > 7590 Phaseolus spp. (s); Dioscorea spp. (s); Manihot spp. (s) Aceituno and Loaiza (2014)
Jaz ca. 7000–5000 Zea mays (p) Aceituno and Loaiza (2007)
Jaz 7080 ± 50 Zea mays (s); Manihot cf. esculenta. (s) Aceituno and Loaiza (2014)
Jaz 5625 ± 50 Manihot spp. (s) Aceituno and Loaiza (2014)
Campoalegre ca. 7600 Dioscorea spp. (p) Aceituno (2001)
Pch 8680 ± 55 Dioscorea spp. (s); Phaseolus spp. (s) Aceituno and Loaiza (2014)
Pch 8095 ± 55 Phaseolus spp. (s) Aceituno (2016)
Pch 9047 ± 45/6903 ± 45 Xanthosoma spp. (s), Fabaceae (s); Zea mays (s) Phaseolus spp. (s) Aceituno (2016)
Pch 6903 ± 45 Xanthosoma spp. (s), Fabaceae (s); Zea mays (s) Aceituno (2016)
Pch 6903 ± 45/6743 ± 45 Xanthosoma spp. Phaseolus cf vulgaris (s) Fabaceae (s); Zea mays (s); Dioscorea spp. (s); Manihot cf Aceituno and Lalinde (2011)
esculenta (s)
Pch 6743 ± 45 Zea mays (s, p) Aceituno and Lalinde (2011)
Pch 6743 ± 45/5922 ± 51 Xanthosoma spp. (s); Phaseolus cf vulgaris (s); Fabaceae (s) Aceituno (2016)
Arrayanes 6520 ± 90 Juglans nigra (m) Rodríguez (1997)
La Selva 8712 ± 60 Phaseolus spp. (s); Dioscorea spp. (s) Aceituno and Loaiza (2014)
Canaán ca.5600 Zea mays (s); Manihot cf esculenta (s); Calathea sp. (s); Ipomoea cf batatas (s) Dickau (2008)
Guayabito 4180 ± 70 Zea mays (p); Manihot esculenta (p); Passiflora spp. (p) Aceituno (2002)

p: pollen.
s: starch grain.
m: macrobotanical remain.

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F.J. Aceituno, N. Loaiza Journal of Anthropological Archaeology 49 (2018) 161–172

Fig. 4. (a) El Jazmín (code 514) milling stone, Manihot cf esculenta; (b) El Jazmín (code 514) Phaseolus spp.; (c) El Jazmín (code 514) Dioscorea spp.; (d) El Jazmín (code 202) Zea mays; (e)
El Jazmín (code 202) B2 level 16 milling stone, Manihot cf esculenta; (f) El Jazmín (code 30) A1 level 14, handstone, Xanthosoma spp.; (g) El Jazmín (code 265) D2 level 18 handstone,
Ipomoea spp.; (h) La Pochola (code 43) A2 level 19 handstone, Xanthosoma spp; i) La Pochola (code 104) Fabaceae; (j) La Pochola (code 665) B5 level 14 milling stone, Xanthosoma spp.;
(k) La Pochola (code 665) Zea mays; (l) La Pochola (code 762) A’5 level 13 handstone, Phaseolus cf vulgaris; (m) La Pochola (code 762) Manihot cf esculenta; (n) La Pochola (code 665)
Phaseolus cf vulgaris; (o) La Selva (code 90) level 10, scraper, Dioscorea spp.; (p) La Selva (code 90) Manihot spp.

according to the shape (Castillo and Aceituno, 2006; Otero and Santos,
2006: 74–86).
As in middle Cauca, the Medellin/Porce region cultural sequence is
long (Fig. 2), allowing an in-depth analysis of the adaptive behaviors
involved in the development of food production. This region has also
zooarchaeological and human remains, adding two more lines of evi-
dence. The early Holocene record is represented only by the identifi-
cation of Dioscorea spp. pollen at La Morena dated between
10,060 ± 60 and 9680 ± 60 years BP (Santos et al., 2015). For this
same time it has been suggested for the Porce subregion a broad
spectrum economy that used mostly riverine ecosystems since their
resources are richer and more predictable than inland resources
(Piperno and Pearsall, 1998: 74; Castillo and Aceituno, 2006). The
pollen record does not show evidence of anthropogenic alteration as-
sociated with plant use.
The middle Holocene (ca. 7700–4000 years BP) pollen record dis-
plays major changes that suggest variation in the economic strategies
adapted by Medellin/Porce region settlers. Towards ca. 7700 years BP
there is evidence of vegetation alteration associated with the clearing of
plots to favor the growth of food plants from the families Solanaceae,
Arecaceae, Annonaceae and Passiflorae, among others (Castillo and
Aceituno, 2006). Associated with the clearings, the record shows the
appearance of exogenous crops such as maize (pollen, phytoliths, and
starch grains) and Manihot spp. (starch grains), along with other plants

166
F.J. Aceituno, N. Loaiza Journal of Anthropological Archaeology 49 (2018) 161–172

Table 3
Medellín/Porce River sites with associated archaeobotanical evidence and dates.

14
Site C YBP Archaeobotanical evidence Reference

PIIIOI-52 7730 ± 170 Zea mays (p, ph, s) Phaseolus trichocarpus (p) Otero and Santos (2006) and Santos et al.
(2015)
PII-021 ca.7500–6500 Aracaeae (ph); Zea mays (s), Poaceae (ph) Castillo and Aceituno (2006)
PIIIOI-40 6890 ± 40 Zea mays (ph) Otero and Santos (2012)
PII-021 ca.6500–5000 Zea mays (p), Manihot spp. (p, s), Smilax spp. (p) Amaranthus spp. (p); Annonaceae Castillo and Aceituno (2006)
(ph)
PII-107 ca. 5000 Zea mays (p, ph), Manihot spp. (p) Castillo and Aceituno (2006)
PIIIOP-61 4650 ± 70 Persea americana (m); Cucurbitaceae (m) Cardona et al. (2007)
PIIIOI-52 4170 ± 40 Zea mays (p, ph, s) Otero and Santos (2006)
PII-045 ca. 4200–3500 Amaranthus spp. (s), Manihot spp. (s), Zea mays (s); Ipomoea spp. (s); Cucurbitaceae Castillo and Aceituno (2006)
(ph)
La Morena 10.060 ± 60–4170 ± 50 Phaseolus sp. (p) Persea (p) Santos (2010)
La Morena 10.060 ± 60 Dioscorea (s, p) Santos et al. (2015)
9680 ± 60
La Morena 7080 ± 40 Zea mays (p, s) Dioscorea (p) Santos et al. (2015)
La Morena 7080 ± 40 Zea mays (s) Santos (2010)
4170 ± 50
Casa Blanca 4810 ± 70 Zea mays (p) Langebaek et al. (2000)

p: pollen.
s: starch grain.
m: macrobotanical remain.
ph: phytolith.

such as Smilax spp. (pollen), Amaranthus spp. (pollen) and Phaseolus
trichocarpus (pollen) (Table 3), taxa that may have been cultivated
(Castillo and Aceituno, 2006; Otero and Santos, 2006: 420; Santos
et al., 2015). All this botanical evidence suggests that, similar to the
middle Cauca, the Medellin/Porce inhabitants developed horticulture.
As mentioned above, the Porce subregion provides two other lines
of evidence that enhance our understanding of its Archaic occupations:
the human and faunal remains found at the 021 site, dated
7500–5500 years BP (Castillo, 1998: 45) and with the appearance of
pottery ca. 5000 years BP (Castillo and Aceituno, 2006). Women, men
and children were buried individually, in couples and/or in small
groups, covered with soils and rocks (Castillo, 1998: 47). About 14,000
animal bones were recovered accompanying the human burials, in a
mammal fauna dominated by paca (Cuniculus paca), Tome’s spiny rat
(Proechimys semispinosus), black agouti (Dasyprocta fuliginosa), Brazilian
porcupine (Coendou prehensilis) and nine-banded armadillo (Dasypus
novemcinctus), along with small amounts of carnivore, bird and reptile
bones (Castillo and Aceituno, 2006).
To sum up, archaeobotanical and zooarchaeological evidence sug-
gest that horticulture was the base of the economy: small scale culti-
vation complemented by the gathering and hunting of local resources.
This is also supported by trace elements and stable isotopic analysis on
the human remains. The values for Strontium (558 ppm), Manganese
(919 ppm), Barium (148 ppm), Vanadium (98 ppm), Zinc (371 ppm),
Cooper (9 ppm) and 13C (−25.09 and −24.62) suggest an elevated
consumption of green plants, low consumption of nuts and fiber, mild
to low meat consumption and very low fish, crustacean and mollusk
consumption (Beasley et al., 2013; Corti et al., 2013). The Ba/Sr cor-
relation index (−0.58) also suggests a terrestrial diet based on plants
with a low representation of aquatic resources (Trancho et al., 1996;
Castillo and Aceituno, 2006).
The act of burying the dead in a single place accompanied by of-
ferings of several animal species and lithic tools can be interpreted as a
cultural strategy for appropriating the landscape, creating a bond with
the ancestors that reinforced the rights of a particular group over the
local resources (Brown, 1995; Castillo and Aceituno, 2006). It has been
suggested that the use of pottery vessels (called Cancana) was mainly
ceremonial, i e, for festivities and social exchanges between neigh-
boring groups as a strategy to minimize conflict (Castillo and Aceituno,
2006).
2.6. Amazon River Basin
Outside the Colombian Andes, in the Caquetá River Basin in the
greater Amazon River Basin lies the Peña Roja site at 100 masl in a
tropical rain forest life zone and dated between 9920 and 8090 years BP
(Fig. 1) (Fig. 2) (Cavelier et al., 1995; Gnecco and Mora, 1997; Mora,
2003: 102; Mora and Gnecco, 2003, Morcote et al., 2014: 43). The lithic
assemblage comprises unifacial flakes, choppers, drills, handstones,
milling stones, hammer stones, hoes and anvils, manufactured on local
raw materials such as chert, quartz and igneous rocks (Cavelier et al.,
1995: 31–32). Thousands of charred seeds were recovered from this
Archaic occupation that included several palm and fruit trees such as
Anaueria brasiliensis, Astrocaryum aculeatum, Astrocaryum chambira, As-
trocaryum jauari, Bactris sp., Euterpe precatoria, Oenocarpus bacaba, Oe-
nocarpus bataua, Oenocarpus minor, Mauritia flexuosa, Parkia multijuga,
Inga spp., Passiflora quadrangularis, Brosimum guianense, Sacoglottis spp.
and Caryocar spp. among others (Morcote et al., 1998, Morcote et al.,
2014). Phytoliths from squash (Cucurbita spp.), bottle gourd (Lagenaria
siceraria) and leren were also identified. The first two phytolith-pro-
ducing taxa are exogenous plants that had to be brought to the region
for cultivation (Piperno and Pearsall, 1998: 204–205).
Starch grain analysis identified Xanthosoma spp. was also recovered
from two lithic tools (hoe and handstone) and dated between 8800 and
8730 years BP (Morcote et al., 2014: 44). The evidence suggests that
Peña Roja settlers were practicing selective management of palm and
fruit trees, as well as cultivation of other food plants (Cavelier et al.,
1995: 36–41; Morcote et al., 1998; Morcote et al., 2014).
2.7. The Sabana de Bogota and the Middle Magdalena River Basin
Two other regions in Colombia that have provided important but
incompletely described archaeobotanical data for this period, are the
Sabana de Bogota and the Middle Magdalena river basin (Fig. 1). These
two regions have not yet been identified as being as important for
discussions about the origin of food production. Nonetheless, we pro-
pose that both sites are relevant for understanding Archaic adaptations
in New World tropical forests across the Plate Pleistocene/early Holo-
cene boundary.
The Sabana de Bogota is located in the Cordillera Oriental at 2600
masl with 11 archaeological sites. This region has the longest occupa-
tional sequence in northwest South America and includes megafauna

167
F.J. Aceituno, N. Loaiza
butchering in the Pleistocene/Holocene transition to food production in
the upper Holocene, dated from ca. 12,500 to 3000 year BP (Fig. 1)
(Fig. 2) (Correal and van der Hammen, 1977; Hurt et al., 1977; Correal,
1982, 1986, 1990, 1993; Ardila, 1984; Groot, 1992, 1995). Correal
(1986: 123–125) interprets that faunal remains associated to Abriense
technology (unifacial retouched flakes belong to the edge-trimmed
tradition) suggests that hunting played an important role in the
economy and was complemented with the gathering of snails, crabs,
tubers and seeds. Recent lithic use-wear analysis supports the idea that
Abriense technology was used on a broad spectrum of resources that
included wood, bone and leather working, as well as tools that had
bamboo phytoliths on them, while only one was identified as a butch-
ering tool (Nieuwenhuis, 2002: 54–55, 61, 65, 67). The presence of a
wide variety of faunal remains that include species from lower regions
also supports the idea of a broad spectrum economy that not only fo-
cused on major game but also on hunting minor species (Correal, 1990:
257). The continuous presence of guinea pig (Cavia porcellus) and the
focus on this species over other larger mammals such as white tailed
deer (Odocoileus virginianus) by the middle Holocene species has led to
the hypothesis that guinea pig domestication may have taken place on
the Sabana de Bogota between ca 3600 and 2500 years BP (Correal and
Pinto, 1983; Correal, 1986: 125; Delgado, 2016).
As the Holocene advanced, lithic technology transformed in-
corporating an increasing amount of tools focused on plant processing,
such as handstones, anvils, hammers and milling stones (Ardila, 1984;
Correal, 1990: 257; Correal, 1987; Groot, 1992: 6; Groot, 1995: 54). At
the Checua site edge ground cobbles were recovered along with milling
and hammer stones dated ca. 8500 years BP, that suggest root and tuber
processing activities (Groot, 1995: 54; Groot, 1992: 6). By ca.
3800 years BP there is macrobotanical evidence of oca (Oxalis tuberosa)
and squash (Cucurbita pepo) according to Correal (1990: 260) and by ca.
3300 years BP, avocado, maize and sweet potato (Correal, 1990: 261),
suggesting that food production practices based on plant cultivation
since the onset of the Holocene. The increase of faunal diversity in
several sites along with the evidence from Checua suggest an expansion
of the resources used that had a minimum presence in Pleistocene le-
vels, such as the plants that increase in frequency as the Holocene ad-
vances.
The Middle Magdalena (middle Magdalena River Basin) is located
between the Cordillera Central and Oriental, between 100 and 200 masl
in a tropical rain forest environment (Fig. 1). It constitutes another
region that, even though it has yet to have substantive archaeobotanical
data on food production, it is relevant because of the time frame of the
seven sites, dated between 10,400 ± 60 and 3130 ± 70 years BP
(López, 1995, 1999, 2008; Otero and Santos, 2002) (Fig. 2), and also
the geographical setting. The lithic tools are composed of several fish-
tail bifacial projectile points, plano-convex scrapers, and many other
tools that have been associated with Paleoindian adaptations (López,
1999: 100). The fact that this technology remains the same into the
upper Holocene, has lead for the argument that it was a broad spectrum
adaptation based on the wide diversity of riverine resources such as
reptiles, birds, water mammals, fish, as well as inland plants and ani-
mals (Otero and Santos, 2002). To date, evidence for plant use is limited
to a few starch grains recovered from tools (Nieuwenhuis, 2002: 88–89)
with a handful of lithic tools focused on plant processing (Otero and
Santos, 2002). Lithic use-wear analysis suggest that the stone tools were
used mainly to process fish, leather, wood, bone, and marginally other
plants (Nieuwenhuis, 2002: 106).
3. Discussion
All the data presented above suggest low-level food production
economies that entail strategies such as ecosystem alteration, resource
management and plant cultivation during the Holocene, a geological
epoch that brought large scale environmental changes affecting plant
and animal communities alike as well as unprecedented anthropic
168
Journal of Anthropological Archaeology 49 (2018) 161–172
alterations (Sthal, 2016). Collectively, the archaeological evidence from
Colombia supports the worldwide hypothesis that even though some
plants and animals were used and domesticated since the Pleistocene/
Holocene transition, the development of low-level food production
economies is a Holocene adaptation (Pringle, 1998; Richerson et al.,
2001; Bettinger et al., 2009).
The progressive increase in archaeological evidence from ca.
10,500 years BP in several regions suggest the territorial movements
and expansion into other areas through the Colombian Andes. This si-
tuation is similar to what happens in the archaeological records of
adjacent regions such as Panama and Ecuador (Cooke, 1992, 2005;
Piperno and Pearsall, 1998: 187; Piperno et al., 2000; Piperno, 2006,
2011a,b; Dickau et al., 2007; Pagán-Jiménez et al., 2016).
The archaeobotanical component (macrobotanical remains, pollen,
starch grains and phytoliths) shows that carbohydrate rich tuberous
plants are a constant in the record of these regions. The data analyzed
not only suggest the use of specific plant resources but goes further; in
Popayan (Gnecco, 2000, 2003) and the Amazon basin (Morcote et al.,
1998, 2014) researchers have interpreted the data as evidence of early
anthropic alterations as a set of strategies of landscape management
and control of natural resources.
In general terms, the taxa identified through macro and micro-
botanical analysis, along with the evidence of anthropic alteration in
some regions during the early Holocene, suggest that early inhabitants
were practicing small scale cultivation and plant management as a
strategy to increase the carrying capacity of local ecosystems, allowing
for an increased control of local resources to ensure the procurement of
basic resources. The Peña Roja site is a good example: Cucurbita sp. and
bottle gourd phytoliths show that forest dwellers were cultivating both
crops and were also using several kinds of palms, as the thousands of
palm seeds recovered inform. Taken as a whole this suggest that people
were altering the distribution of natural resources in some sort of forest
management that may have further implications on mobility and de-
mographic growth.
For Popayan Gnecco defends the idea of early cultivation and
management based on taxa such as arrowroot, avocado, and others such
as Xanthosoma or Ipomoea (Gnecco, 2000, 2003; Gnecco and Aceituno,
2004, 2006), along with the presence of allopatric species in the pollen
record that have been interpreted as brought together by anthropogenic
action. For middle Cauca the alteration evidence are weaker, but the
taxa identified dating to the early Holocene (e.g. Fabaceae, Xanthosoma,
Dioscorea, Phaseolus) that endure through the middle Holocene, suggest
that selection through cultivation was a common strategy in the region
(Aceituno and Loaiza, 2014).
Assuming that all these Holocene adaptations included resource
control and management, we need to ask: which were the environ-
mental and/or cultural pressures that triggered the adaptive responses
that were a steppingstone for the development of food production,
domestication and installment of agriculture? The climatic changes
brought by the Pleistocene/Holocene transition included increased
rainfall that lasted up to the Holocene Climatic Optimum
(7000–5000 years BP) favoring the rapid and widespread expansion of
humid forests and decline of open habitats, which severely affected
fauna, especially large mammals (Piperno, 2017; Piperno and Pearsall,
1998: 106–107; Vivo and Carmignotto, 2004). Punctuated and/or cy-
clical events of different intensities such as earthquakes, volcanic ac-
tivity, tsunamis, ENSO, anthropic alterations, among others, need to be
added to climatic tendencies that affected the distribution and relative
abundance of flora and fauna species during the Holocene (Sthal,
2016). The lack of regional scale studies that provide information on
this topic prevents us from assessing how these continental scale
changes behaved in local scales and how they affected the distribution
of resources in the diverse environments of Colombia.
Optimal Foraging models has been a powerful theoretical tool to
explain the adaptive chances that lead some human societies towards
food production (Winterhalder and Kennett, 2006; Piperno, 2011,
F.J. Aceituno, N. Loaiza
Piperno, 2017; Cooding and Bird, 2016; Stiner and Kuhn, 2016). One of
its models Diet Breadth states that when return rates fall lower ranked
resources will be included in the diet, this may include innovations to
make them stable in the long run that allow individuals to maintain
stable return rates (Cooding and Bird, 2016). Demographic pressures
and resource depression have been the most used factors to explain why
human societies started to expand their diets at the Pleistocene/Holo-
cene transition as a prelude to the development food production
(Piperno, 1989; Smith and Winterhalder, 1992; Piperno and Pearsall,
1998; Richerson et al., 2001; Winterhalder and Kennett, 2006; Piperno,
2011a; Bird et al., 2016; Cooding and Bird, 2016; Stiner and Kuhn,
2016).
Resource depression has been wildly used as the major pressure that
drove human populations for the adaptations that led to the develop-
ment of food production. In this sense megafaunal extinction has been
assumed as the major change that triggered the adoption of lower
ranked resources i.e. plants and small animals (Bird et al., 2016). The
scarce archaeological evidence of megafauna in Colombia does not
suggest that human groups were specialized megafauna hunters
(Guitérrez (2010)). Megafauna hunting evidence has been found in
Tibito (Sabana de Bogota), were mastodon (Haplomastodon and Cu-
vieronius), horse (Equus sp.) and white tailed deer dated to
11,740 ± 110 BP were recovered (Correal, 1982). Other sites with
megafaunal evidence associated with artifacts are El Totumo (Cordil-
lera Oriental) and La Pileta (Santander) (Correal, 1993); Toro, between
Calima and Middle Cauca in the Cauca River Basin (Rodriguez, 2002, p.
29), and Yumbo, south of Toro also in the Cauca River Basin
(Rodriguez, 2002, p. 33). Unfortunately, none of these sites have un-
disputable evidence of human activities nor have they been dated, and
stratigraphy has only been reported for the first two.
It is likely that the shifts in density and distribution of small fauna
during the Pleistocene/Holocene transition had a higher impact on
human populations than megafaunal extinction. South America went
from 20 mammalian orders in the Pleistocene to 12 in current times,
which means a 40% reduction (Vivo and Carmignotto, 2004). Holocene
mammals are usually small and solitary and because of the expansion of
humid forests in this time frame there is an associated expansion of
fauna that has arboreal locomotion (Piperno and Pearsall, 1998: 62;
Vivo and Carmignotto, 2004). Human groups must have been sensitive
to the changes in densities of minor fauna. The absence of studies
dealing with small fauna in Colombia difficult makes it hard to assess
the real weight of this variable, because paleontological studies have
centered mostly on megafauna. As an example, the archaeological re-
cord from Sabana de Bogotá, despite the extinction of horse and mas-
todon, suggests that the area had wide terrestrial and aquatic resources
that help explaining the continuity in human occupations through the
Holocene.
There is evidence on territorial expansion into other areas without
since the Pleistocene/Holocene transition in Colombia (Aceituno et al.,
2013), but there is no clear evidence that there was population grouth
enough to cause the demographic pressures required to make an impact
on resources. The estimation of this variable requires more detailed
paleoecological and archaeological information that is unavailable at
the time.
Rather than a single factor, we suspect that a combination of them
including megafaunal extinction (where present), redistribution of
ecological communities (plants and animals), demographic growth,
territorial expansion, among others, were the trigger to develop new
behaviors, such as the inclusion of other resources into the diet. In turn,
this led to the selective pressures over those resources and the alteration
of niches allowing for greater control over some animal and plant po-
pulations. Is in this scenario where plant cultivation emerges as an
adaptive strategy with ecological consequences that benefited human
populations, including the increase in carrying capacity, reduction of
resource acquisition costs and increase in predictability. The anthro-
pogenic alteration of ecosystems allowed for environmental conditions
169
Journal of Anthropological Archaeology 49 (2018) 161–172
that favored cultivation and the domestication process. These areas
would be the scenarios were new selective pressures would act upon the
phenotypic plasticity of plants and in some case developing domes-
ticates (Piperno, 2017).
Plant cultivation was developed in times of environmental and cli-
matic changes as adaptive responses to the new conditions that affected
resource distribution and availability. The inclusion of a wider diversity
of plants into the diet was a risk minimizing strategy that aimed to
guarantee resource availability in different parts of the year as well as
to provide a form of bait to procure faunal resources (e.g. Linares, 1976;
Dufour, 1990: 54). The high diversity of plant taxa, along with the
evidence for alteration and the presence of foreign plants reinforce the
idea of risk minimizing and broad spectrum economies.
The appearance of foreign domestic crops (manioc, maize and likely
common beans) between ca. 7700 and 6500 years BP, further supports
that cultivation was part of an adaptive strategy developed by popu-
lations in Colombia during the early Holocene, and maintained through
the middle Holocene. Furthermore, it supports the idea of early dis-
persal of crops and horticulture through the Neotropics, a food pro-
duction strategy that also involved the tenure of a wide variety of plants
including fruit trees and small plants (Piperno and Pearsall, 1998: 7;
Harris, 2007: 23–24). In spatial terms this would also mean a higher
dependency on cultivated plots, which in turn would reduce mobility
and increase territorial control. Nonetheless, this has regional differ-
ences: while in the middle Cauca there are no clear differences on site
sizes through time, in the Porce region there is a high investment on site
preparation that involved chipped stone floors for habitation areas as
well as a specific place to bury the dead between ca. 7700 and
5500 years BP.
The archaeobotanical data recovered during the last 30 years
highlight the role of Colombia in the domestication of plants, since the
macro region has been proposed as the domestication center for several
crops including cocoyam, leren, arrowroot and sweet potato (Piperno
and Pearsall, 1998: 164–165) within region D1 (Piperno, 2011a). Al-
though, the exact location of where domestication took place remains
unknown, microbotanical analysis done on archaeological sites have
been able to identify all of the genera that those crops belong to dating
back to the early and middle Holocene. Xanthosoma has been identified
in tools in the Amazon Basin at ca. 8800 year BP (Morcote et al., 2014:
44), middle Cauca in several tools dating between ca. 9000 and
5500 years BP, as well as in the pollen record at El Jazmin between ca.
9000 and 5000 years BP (Jaramillo and Mejía, 2000a; Aceituno and
Loaiza, 2007: 84–86; Aceituno and Lalinde, 2011; Aceituno, 2016), and
a tentative identification for Popayan dated ca. 9500 years BP sup-
porting this idea (Piperno and Pearsall, 1998: 200). The genus Maranta
was identified through phytoliths from soils in Calima dated between
ca. 9000 and 4500 years BP (Piperno, 1985; Piperno and Pearsall, 1998:
202), and tentatively in phytoliths from a tool residue at Popayan
dating ca. 9500 years BP (Piperno and Pearsall, 1998: 200). Calathea
has been identified in a tool from middle Cauca dated ca. 5600 years BP
(Dickau, 2008), and from phytoliths from soils in the Amazon basin
dating ca. 8800 years BP (Piperno and Pearsall, 1998: 204–205). Ipo-
moea from starch grains in the Middle-Cauca, dated ca. 7000 years BP
and Medellin/Porce region dated between ca. 4200 and 3500 years BP
(Castillo and Aceituno, 2006). As mentioned previously taxonomical
assessment has only been determined to genus, and in some cases is just
tentative, this is due to the fact that microfossil comparative collections
need to be improved to reach more precision and that sometimes the
level of species and assessment of wild vs. domesticated cannot be
reached.
4. Conclusions
Archaeobotanical data published over the past 30 years have posi-
tioned Colombia as an independent center for the origin of plant use
and domestication (Piperno and Pearsall, 1998: 165; Piperno, 2011;
F.J. Aceituno, N. Loaiza
Piperno et al., 2017). Several species have been suggested as being
brought under domestication in this part of the world including ar-
rowroot, cocoyam, leren and sweet potato, among others. All this bo-
tanical evidence suggests that the food production strategy of the in-
habitants of different regions responded to specific local agroecologies.
Food production in forests in broken terrain would have been more
microzonal than in wide colluvial valleys, volcanic basins or along
coasts.
The data discussed above show that since the Pleistocene/Holocene
transition human groups altered local environments in order to increase
human control over natural resources especially useful plants that
present a disperse distribution in tropical forests. Following the opti-
mization principle, cultural selection favored behaviors that enhanced
prediction and control and decreased procurement time. The alteration
in distribution and diversity of plants may have also served as a way to
increase the predictability on animal behavior, increasing the chances
of encounter. The result of combining all these strategies is the devel-
opment of food producing economies. By the middle Holocene, horti-
culture consolidates all of the strategies as the main form of food pro-
duction that extends to most of the regions discussed in the previous
sections. The development and implementation of horticultural prac-
tices must have altered population growth, mobility, and territoriality
in different ways in each region paving the way for the establishment of
agricultural societies, but those effects are beyond the scope of this
paper.
The progressive increase in use, control and management of plants
is associated to the environmental changes during the Pleistocene/
Holocene transition that affected resource distribution and structure.
The increased dependence on plants along with the strategies devel-
oped to manipulate them, are part of the behavioral adaptations that
human societies have to ensure survival. Cultural adjustments have an
impact in local ecosystems just as the latter have in the former, creating
codependency in this new ecological relation leading in some cases to
domestication. The arrival of foreign resources towards the middle
Holocene adopted into local practices reinforces the idea that native
adaptations played a fundamental role in the development of food
production strategies in Colombia.
Acknowledgements
We want to thank the anonymous reviewers for their valuable
comments and suggestions. NL wants to thank Wenner-Gren
Foundation for the Wadsworth International Fellowship and
Colciencias for the Beca Francisco Jose de Caldas, both aimed to sup-
port his doctoral studies. Both authors want to thank Vicerrectoría de
Investigación (Universidad de Antioquia) and the Research Group
Medioambiente y Sociedad for their support.
Appendix A. Supplementary material
Supplementary data associated with this article can be found, in the
online version, at http://dx.doi.org/10.1016/j.jaa.2017.12.007.
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