Genet Resour Crop Evol (2010) 57:813–825
DOI 10.1007/s10722-009-9521-4
RESEARCH ARTICLE
Origin of agriculture and plant domestication
in West Mesoamerica
Daniel Zizumbo-Villarreal • Patricia Colunga-Garcı́aMarı́n
Received: 24 April 2009 / Accepted: 14 December 2009 / Published online: 2 February 2010
Ó Springer Science+Business Media B.V. 2010
Abstract Recent paleoecological, archaeobotanical
and genetic-molecular data are used to develop a
hypothesis on the where, when, how and whom of
plant domestication and the origin of agriculture in
west Mesoamerica, and the formation of the maize-
bean-squash multicrop milpa system and agro-food
system which formed the base for development of
ancient complex societies in this area. It is highly
likely that about 10,000 before present (BP) human
groups specializing in plant gathering and small game
hunting in the dry tropical forest of the Balsas-Jalisco
biotic morphotectonic province began the process of
plant domestication and agriculture, using fire as a
tool. Sympatric distribution of the putative wild
ancestral populations of maize, beans and squash
indicate the extreme northwest Balsas-Jalisco region
as a possible locus of domestication. Diffusion of
these domesticates to the rest of Mesoamerica would
have occurred via existing biological-cultural corri-
dors. The milpa agro-food system would have been
established between 7,000 and 4,400 calendar years
(cal) BP. The complex food technology developed in
the northwest Balsas-Jalisco region between 4,500
and 3,500 BP, much more complex than in other
areas at the time, also suggests this area as the origin
D. Zizumbo-Villarreal (&)
P. Colunga-Garcı́aMarı́n
Unidad de Recursos Naturales, Centro de Investigación
Cientı́fica de Yucatán, Calle 43 No 130. Col. Chuburná de
Hidalgo, 97200 Mérida, Yucatán, México
e-mail: zizumbodaniel@gmail.com
of the milpa agro-food system. Further archaeobo-
tanical research is needed to confirm this hypothesis.
Exploratory, collection and conservation efforts are
needed in these putative source populations, as well
as studies on their adaptation to climatic, edaphic and
biotic factors, before they are displaced by the
African grasses and pesticides forming part of the
region’s growing cattle industry.
Keywords Agriculture
Beans
Domestication

Maize
Mesoamerica
Squash
Introduction
One of the most significant events in human history
was the transformation from a hunting-gathering
economy to an agricultural economy (Smith 2005).
This change probably occurred independently in at
least six regions in the world, primarily tropical and
subtropical areas with high biological and cultural
diversity (Gepts 2008; Piperno and Pearsall 1998;
Sauer 1952). Along with the Middle East and north
China, Mesoamerica is one of the world’s primary
centers of domestication (Harlan 1971, 1995). It was
here that species such as maize (Zea mays L.), beans
(Phaseolus spp.) and squash (Cucurbita spp.) were
domesticated and integrated into a multi-crop system
known in the region today as milpa. It was this
system’s ecological and nutritional complementarity
that helped to support the development of highly
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complex societies in Mesoamerica (Gepts 2008).
Mesoamerica is most commonly defined as the
cultural area extending from the Santiago and Panuco
river basins in Mexico, south to the Central Valley
region of Costa Rica (Kirchhoff 1943).
Plant domestication is understood as the continuous
historic evolutionary process, driven by human selec-
tion, which produces fixation of a set of alleles that
provides wild populations with favorable human
consumption and cultivation phenotypes, but which
also diminishes or eliminates the capacity for survival
under natural conditions, making a domesticated
population dependent on humans (Harlan 1992). The
set of phenotypic traits determined by these alleles is
known as the domestication syndrome (Hammer
1984). Plants’ reproductive systems and genetic con-
stitution have favored and hindered domestication to
varying degrees. Plants that have responded positively
to the selection process and agricultural management
have been completely domesticated, while in others
only certain traits of the syndrome have been fixed,
leaving them semi-domesticated or in process (Gepts
2004). In agricultural systems under limiting environ-
mental conditions, humans have managed to maintain
some species in a semi-domesticated state as part of
their productive strategy (Colunga-Garcı́aMarı́n and
Zizumbo-Villarreal 1993).
For the purposes of this discussion, agricultural
management is to be understood as a set of deliberate,
environmental modifications made by humans aimed
at increasing survival and biomass production in
selected plant species, and therefore meeting human
needs given prevailing environmental conditions.
Plant domestication and agriculture are temporally
continuous and interdependent processes. Over time,
humans have selected a group of local and introduced
species for subsistence, and developed knowledge
and techniques for their cultivation, transport, stor-
age, transformation, consumption and preservation.
This group of species and the knowledge and
techniques associated with them are known as an
agro-food system.
The present study aim was to integrate current
paleoecological, archaeobotanical and genetic-molec-
ular data to develop hypotheses on the where, when,
how and whom of plant domestication and agricul-
tural development in west Mesoamerica; by what
routes these domesticates and related knowledge and
technology were distributed; and how and where the
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Genet Resour Crop Evol (2010) 57:813–825
milpa multicrop system and associated agro-food
system came into existence. The answers to these
questions will be vital to locating the genetic
diversity nuclei so important to productivity, sustain-
ability and improvement in current agro-food sys-
tems, and to better understanding the material
foundations of complex cultures in Mesoamerica.
Initial human occupation
Human occupation of Mesoamerica began about
11,600 years before present (BP) (Dixon 1999).
These partially sedentary early arrivals survived by
fishing, hunting megafauna and collecting plants in
areas near the highland intermontane lake systems of
Chapala, Sayula-Zacoalco, Zacapu, Cuitzeo, Mexico
and Puebla-Valsequillo, (Aliphat 1980; Irish et al.
2000; Lorenzo and Mirabell 1986; Niederberger
1979; Tolstoy et al. 1977). Their principal hunting
implement was initially designed like a harpoon used
for hunting marine mammals, suggesting that they
had moved inland from the Pacific coast (Fig. 1).
A following migration into the area probably
occurred around 10,600 BP (Dixon 2001) from the
west and great plains of North America. These groups
implemented an ecological strategy based on small
game hunting and plant gathering. They processed
some plant species to make them edible or extract
certain elements; for instance, ground seeds from
Panicum spp. and Setaria spp.; cooked stems and leaf
bases from Agave spp.; stems and fresh fruits from
Opuntia spp.; seeds from Quercus spp. and Pinus
spp.; and fruits from Prosopis spp. They used
sharpened scrapers and points for gathering and
hunting, and stone technology for breaking, grinding
and cooking (Doebley 1984; Poinar et al. 2001;
Willis 1995). In addition to their use of fire, they
introduced the dog (Canis familiaris L.), domesti-
cated from multiple lineages in eastern Asia (Leonard
et al. 2002; Wayne et al. 2006). Use of dogs allowed
them to form small, highly mobile bands with
defensive capacity. They occupied rock shelters
along the banks of rivers communicating the inter-
montane valleys with the west coast. Technology
among these groups included thin bifaces, flint points,
river pebbles and stone blocks or platforms modified
for grinding (Flannery 1986; MacNeish 1967a; Mac-
Neish and Peterson 1962; Ranere et al. 2009).
Genet Resour Crop Evol (2010) 57:813–825
Fig. 1 Probable routes of initial human migrations into
Mesoamerica along river corridors: (1) Grande de Santiago-
Lerma, (2) Armerı́a-Tuxcacuesco, (3) Balsas, (4) Amacuzac
and (5) Mexcala-Nexpa-Atoyac. Main Pleistocenic fauna
Ecological conditions during initial domestication
and early agriculture
Paleoecological records for the Neovolcanic Trans-
verse Axis and the lowlands south of this axis, indicate
rises in temperature, rainfall and atmospheric CO2
concentrations between 12,000 and 9,000 BP, as well as
the presence of a long drought period before the rainy
season (Cunniff et al. 2008; Metcalfe 2006; Piperno
2006). Tropical flora began to displace the boreal
forests, and the lowland thorny bush vegetation was
taken over by tropical dry forest (TDF). The combina-
tion of these transformations produced a transition from
C3 to C4 grasses, growing populations of Panicum spp.,
Setaria spp., Tripsacum spp. and Zea spp., and
expansion of certain dicotyledonea families, such as
Chenopodiaceae, Amaranthaceae, Asteraceae, Cucur-
bitaceae and Solanaceae (Cunniff et al. 2008; Piperno
et al. 2007; Sage 1995). It was during this period that
the Balsas-Jalisco (BJ) biotic morphotectonic province
(Fig. 2) took form south of the Neovolcanic Transverse
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hunting sites in the Neovolcanic Transversal Axis: (A)
Chapala-Zacoalco-Sayula, (B) Valley of Mexico and (C)
Puebla-Valsequillo
Axis (Ferrusquı́a-Villafranca 1990). Its floral and
faunal distribution, structure and composition have
changed little during the last 9,000 years (Metcalfe
2006; Piperno 2006). Between 16,500 and 4,000 BP,
high eruptive activities and extensive lava flows from
the Colima Volcanic Complex (CVC) split the region
from north to south (Cortés et al. 2005; Capra and
Macı́as 2002), leaving an area to the northwest rich in
grasses, including: Tripsacum dactyloides (L.) L. var.
mexicanum De Wet et Harlan L; T. lanceolatum Rupr.
ex Fourn.; T. maizar Hernández et Randolph;
T. pilosum Scribner and Merr.; Zea diploperennis Iltis,
Doebley et Guzmán; Z. mays L. ssp. parviglumis Iltis et
Doebley; and Z. perennis (Hitchcock) Reeves et
Mangelsdorf (Eubanks 2001; Wilkes 2004).
Distribution of putative ancestral populations
Phylogenetic analyses using nuclear DNA have shown
that Z. mays ssp. parviglumis populations in the central
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Fig. 2 Biotic provinces (Ferrusquı́a-Villafranca 1990), and archaeological sites (Flannery 1986; Kelly 1980; MacNeish 1964;
Mountjoy 2006; Ranere et al. 2009)
and west BJ region could be the putative progenitors of
domesticated maize (Buckler et al. 2006; Fukunaga
et al. 2005; Matsuoka et al. 2002). Phylogenetic
relationships inferred from mitochondrial genes sug-
gest that wild populations of C. argyrosperma Huber
ssp. sororia Merrick et Bates in this region are the
putative source populations of domesticated varieties
(Sanjur et al. 2002). For the bean Phaseolus vulgaris
L., genetic-molecular research using nuclear DNA
suggests that wild populations of this species in the
lower Lerma-Santiago river basin and in the northwest
BJ region are the ancestral populations of domesti-
cated varieties (Gepts 1988; Kwak et al. 2009,
Zizumbo-Villarreal et al. 2009a) (Fig. 3).
The origin of agriculture
A growing carbon accumulation in paleoecological
records from the BJ region to Panama’s southern
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Genet Resour Crop Evol (2010) 57:813–825
coast between 10,000 and 7,000 BP seems to indicate
systematic use of fire by humans. There is also a
simultaneous increase in the presence and accumu-
lation of Zea genus grass pollens in the central-
western BJ (Piperno 2006; Piperno et al. 2007).
Continued use of fire in the TDF of the BJ region led
to a drastic decline in diversity, the dominance of tree
species with the capacity to sprout from the root
crown or stem and the appearance of grass patches
(Miller 1999; Miller and Kauffman 1998; Sánchez-
Velázquez et al. 2002). Eventually, teocintle, beans
and squash would begin to grow together in these
patches (Flannery 1986; Wilkes 2004).
The increased density of edible species promoted by
burning could have defined collecting rounds and the
location of seasonal shelters nearby. Rock shelter
distribution, their size and the archaeological remains
inside them in the central BJ region about 8,700 cal-
endar years (cal) BP indicate that they were occupied
for a number of weeks by small groups of humans.
Genet Resour Crop Evol (2010) 57:813–825
Fig. 3 Location of wild populations of putative ancestors to
plant species domesticated in Mesoamerica: (Cu) Cucurbita
argyrosperma Huber; (Pv) Phaseolus vulgaris L; (Zm) Zea
These groups made use of the surrounding resources by
gathering plants, hunting deer and smaller fauna, and
cultivating of maize and squash using tools such as fire,
bifacial scrapers and choppers, as well as handstones
and base stones for grinding (Ranere et al. 2009).
Plant domestication
Based on molecular clock results, the initial separa-
tion between wild and domesticated maize popula-
tions probably occurred &9,000 BP (Matsuoka et al.
2002). This coincides with accelerator mass spec-
trography (AMS) results for maize and squash starch
grains domesticated about 8,700 cal BP found in
Xihuatoxtla in the central BJ region (Ranere et al.
2009). Results reported by Piperno et al. (2009)
support a scenario in which maize was domesticated
in the low, humid portions of the Balsas river
watershed, where Z. mays ssp. parviglumis is native.
The size and morphology of maize starch grains and
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mays L. (Fukunaga et al. 2005; Gepts 1988; Kwak et al. 2009;
Matsuoka et al. 2002; Sanjur et al. 2002; Zizumbo-Villarreal
et al. 2009a)
phytoliths is indicative of human selection for the
teocintle glume architecture gene (tga1), a major
domestication gene in Zea which controls phenotypic
attributes key to efficient utilization of maize as a
grain (Doebley 2004; Piperno et al. 2009). This
selection increased starch quantity and quality, while
reducing grain glume size and hardness, leading to
naked grains with soft glumes (Wang et al. 2005). For
squash, phytolith size and morphology is indicative
of human selection pressure on the Hr gene, a major
domestication gene in Cucurbita which controls
crucial phenotypic attributes such as fruit hardness
and phytolith quantity (Piperno et al. 2002; 2009).
Leguminous starch grains found at Xihuatoxtla in
the central BJ region may correspond to beans
(P. vulgaris or P. lunatus) (Piperno et al. 2009:
supporting data) and suggest that bean may have been
domesticated in this region as early as 9,000 cal BP.
However, AMS-dated macro-remains of beans found
at Tehuacán suggest domestication by 2,285 cal BP,
while bean remains from Guilá Naquitz have been
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dated to 2,098 cal BP (Kaplan and Lynch 1999). In
beans, the domestication syndrome includes traits
that limit natural dispersion, such as loss of fruit
dehiscence, seed latency and photoperiod sensitivity
(Koinange et al. 1996).
Diffusion of domesticated species
In the early Holocene, the distribution of human
shelters in Mesoamerica and the similar technological
development of human groups in the area (MacNeish
1967a; MacNeish and Peterson 1962; Flannery 1986;
Ranere et al. 2009) suggest that rivers, which would
have been regular water sources during the long dry
season, functioned as biological and cultural disper-
sion corridors (Fig. 4). Diffusion of domesticated
Fig. 4 Distribution of human populations in Mesoamerica
during the Archaic period: (a) Sayula (Irish et al. 2000; Benz
2002); (b) Matanchén (Mountjoy et al. 1972); (c) Tlapacoya
(Tolstoy et al. 1977); (d) Valsequillo (MacNeish 1967b); (e)
Xihuatoxtla (Ranere et al. 2009); (f) Puerto Marqués (Brush
1965); (g) Tehuacán (MacNeish 1964); (h) Oaxaca (Flannery
1986); (i) Ocozocuautla (MacNeish and Peterson 1962); and (j)
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Genet Resour Crop Evol (2010) 57:813–825
plant species throughout Mesoamerica along these
river corridors can be inferred from the earliest dates
for the presence of maize. Paleoecological maize
pollen and phytolith records from the Gulf of Mexico
coast date from ca. 7,300 BP (Pohl et al. 2007),
suggesting diffusion along the Tehuantepec, Coatza-
coalcos and Grijalva rivers. Records for maize in the
Tehuacán and Oaxaca intermontane valleys dating to
about 6,300 BP imply diffusion along the Tepalcate-
pec-Balsas-Mexcala and Atoyac-Salado systems. The
presence of maize pollen in Belize by about 5,400 BP
(Pohl et al. 1996) suggests another route along the
Grijalva and Motagua rivers, while records from
Veracruz state dating to ca. 5,000 BP (Sluyter and
Domı́nguez 2006) indicate the Salado and Papaloapan
rivers also functioned as diffusion corridors (Fig. 5).
Of these different routes in Mesoamerica, the
Chanuto (Voorhies et al. 2002). Biocultural corridors during
the Archaic: (A) Chapala-Santiago-Matanchén; (B) Sayula-
Tuxcacuesco-Armerı́a-Tuxpan-Coahuayana; (C) Chapala-Cuit-
zeo-Lerma; (D) Chapala-Tepacatepec-Infiernillo, (E) Valley of
Mexico-Cuautla-Amacuzac-Mezcala-Atoyac; (F) Valsequillo-
Salado-Atoyac-Tehuantepec-Grijalva-Coastal Chiapas
Genet Resour Crop Evol (2010) 57:813–825
Fig. 5 Possible routes for early diffusion of plant species
domesticated in Mesoamerica: (A) Santiago-Lerma; (B) Tepal-
catepec-Balsas; (C) Mexcala-Amacuzac; (D) Atoyac1-Salado-
Atoyac2; (E) Mixteco-Verde-Atoyac2; (F) Tehuantepec;
Tepalcatepec-Balsas-Mexcala corridor was probably
a major route for human movement beginning in the
early Holocene (Benz 1999).
Domesticated maize also moved quickly into
Central and South America, as shown by its presence
in Panama by about 7,800–7,000 BP (Dickau et al.
2007), suggesting diffusion routes along the Grijalva,
Motagua and Chamelecón rivers and along the south
coast of Central America. Paleoecological data for
the presence of maize pollen, starch grains and
phytoliths in the Cauca and Ponce valleys of
Colombia by about 7,500 BP (Aceituno and Castillo
2005), and on the southwest coast of Ecuador around
the 6,200 cal. BP (Zarrillo et al. 2008) imply another
route along the western and central mountain ranges
of Colombia. Finally, maize and chili phytoliths and
starch grain records from grinding artifacts in the
Andean region by 6,000 BP indicate these two
domesticated species were probably diffused simul-
taneously (Perry et al. 2007).
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(G) Grijalva; (H) South coast; (I) Motagua; (J) Caribbean;
(K) Chamalecón; (L) Belize; (M) Coatzacoalcos; (N) Papaluapan;
and (O) Maya lowland
Agricultural intensification
Paleoecological records for the BJ region between
7,000 and 5,550 BP contain evidence of increased
levels of Asteraceae family weed species (typical of
short fallow systems), greater maize pollen accumu-
lation and decreases in carbon deposits (Piperno
2006; Piperno et al. 2007), suggesting agricultural
intensification. Between 7,000 and 6,000 cal BP, a
new tool kit begins to appear in the archaeological
record, including large, open stone ovens, wooden
tools such as levers, contracting stems and traps,
bifacial knives, plano-convex scrapers, bifacial stone
choppers, mortars, pestles and grinding handstones
and bases. Human shelter and camp distribution, size
and occupation show that macrobands (15–20 per-
sons) began to form during the rainy season to
cultivate and harvest plants (Flannery 1986; MacNe-
ish 1964), further suggesting agricultural intensifica-
tion. Cultivation areas were in the piedmont, on
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terraces close to valleys and plains near lakes
(MacNeish 1964; Ranere et al. 2009).
Morphological analyses of maize cobs dating to
6,300 BP indicate the presence of cobs with two to
four rows of grains, grains with reduced glumes and
total retention of seeds, meaning this crop had
become completely reliant on humans as a replace-
ment seed dispersal mechanism (Benz 2001; Piperno
and Flannery 2001). Molecular analyses using
archaic DNA show fixing of Teocintle branched 1
(Tb1), a gene that largely controls short branches
tipped with ears in domesticated maize (Jaenicke-
Després et al. 2003; Jaenicke-Després and Smith
2006).
Milpa multicropping
Between 6,000 and 5,000 BP in Mesoamerica,
semi-sedentary macrobands lived on valley terraces
in small circular or oval units, with small ovens for
cooking food, and common storage of agricultural
surplus (MacNeish 1964; Flannery 1986). Stone
bases are replaced by manos and metates for
grinding, and stone cups and pots appear in the
record (Flannery 1986; MacNeish 1964, 1967b).
These tools broadened the ability to transform
foods, involving their human users in the applica-
tion of new selection pressures on grain species.
During this period, cultivated plots may have been
established near habitation units by transport of
domesticated perennials such as red mombin and
agaves and the spontaneous arrival of weed species
like chilies, tomatoes (Physalis spp.), amaranth
(Amaranthus spp.) and cotton (Gossypium hirsutum
L.). Dogs were raised as food, and archaeological
and molecular studies suggest it was at this time
that the Xoloitzcuintle dog breed arose and was
raised as a food resource in the BJ region (Wayne
et al. 2006).
By 5,500 BP, the alleles for four rows of grains on
maize cobs had fixed; archaeobotanical remains
indicate the presence of cobs with 8–12 rows,
although the alleles for this trait had not yet fixed.
In-field selection for higher numbers of rows sub-
stantially increased maize’s genetic productivity
(Jaenicke-Després et al. 2003; Jaenicke-Després and
Smith 2006).
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Genet Resour Crop Evol (2010) 57:813–825
Development of the agro-food system
The traits involved in the compact architecture of the
maize plant had fixed by about 4,400 BP, including
apical dominance, fewer stalks, one to two cobs on
the central axis and cobs with 12–16 rows (Jaenicke-
Després et al. 2003; Jaenicke-Després and Smith
2006). Ceramics appear in the archaeological record
in the form of comals, cups and simple pots (Brush
1965; Mac Neish 1976b; Flannery 1994), which
would further open the possibilities for food trans-
formation and introduce new selection pressures. It is
during this period that human selection begins to
change the alleles determining protein and starch
quality (pbf and SuI) (Jaenicke-Després et al. 2003;
Jaenicke-Després and Smith 2006), suggesting a
focus on cooking traits. By 4,000 BP, inhabitants of
the intermontane valleys cultivated crops in agrohab-
itats including piedmont terraces, valley terraces,
plains along lakes and rivers and house gardens
(Flannery 1986; MacNeish 1964). A utilitarian
ceramic tradition developed and diversified between
4,500 and 3,500 BP that was related to food harvest,
transport, storage, transformation and consumption.
By 3,500 BP, the Capacha culture had developed in
Colima state and Mascota in southern Jalisco, with a
ceramic corpus that included food processing tools
for soaking, cooking and steaming, as well as
fermentation and possibly distillation of alcoholic
beverages (Kelly 1980; Mountjoy 2006; Zizumbo-
Villarreal et al. 2009b) (Fig. 2). This ceramic diver-
sification reflects greater complexity in the methods
used for food transformation and in the selection
pressures on domesticated species. Between 3,000
and 2,000 BP, maize with a high frequency of the suI-
M2 allele, implied in the amount and quality of floury
starch, appears in the archaeological record (Jae-
nicke-Després and Smith 2006), suggesting selection
linked to tortilla production.
Discussion
Archaeological evidence suggests that agriculture and
plant domestication in west Mesoamerica were
probably initiated by small, highly mobile groups of
humans from a Clovis cultural tradition who gathered
plants, hunted small animals and seasonally inhabited
Genet Resour Crop Evol (2010) 57:813–825
small rock shelters near rivers between inland lake
systems and the Pacific coast (Flannery 1986; Mac-
Neish 1967a; MacNeish and Peterson 1962; Ranere
et al. 2009). These archaeological data do not support
the hypothesis of Sauer (1952) and Harlan (1995) that
domestication was initiated by semi-sedentary fish-
erfolk living near lake systems who initiated agricul-
ture and domestication by transporting weedy
ancestral populations of domesticated maize, squash
and beans to plots near habitation sites, where they
would care for and harvest them. Evidence for more
permanent settlements with homegardens in the BJ
region, Oaxaca and Tehuacan appear rather later than
domestication and agriculture (Flannery 1986; Rane-
re et al. 2009).
Paleoecological, ecophysiological and molecular
genetic data suggest that plant domestication and
agriculture began in the TDF of the BJ region under
conditions of warm temperatures and annual rainfall
near 1,000 mm, and from selection and management
of the tropical grass Zea mays ssp. parviglumis. It is
unlikely that domestication and agriculture arose in
the intermontane valleys under temperate, semi-dry
conditions in xerophytic bush vegetation through
cultivation of the hybrid Zea diploperennis 9
Tripsacum dactyloides, as suggested by MacNeish
and Eubanks (2000) and Eubanks (2001, 2002).
The autoecological traits and altitude distribution
of putative wild ancestral populations for maize and
beans suggest that agriculture could have begun at
intermediate elevations (600–1,600 masl), including
at the oak forest ecotone (Buckler et al. 2006; Kwak
et al. 2009). This would have enabled human groups
to cultivate plants, while making relatively short
excursions to the coast or intermontane lakes (&40–
80 km), thus reinforcing the biocultural corridors
developed during the early Holocene.
The sympatric geographic distribution of the
putative ancestral populations of maize, beans and
squash in the northwest BJ region suggest that their
cultivation and domestication could have begun in
that area. Early dates for maize and squash place the
beginning of this process around 10,000 BP, making
it roughly contemporaneous with the beginning of
domestication in the Old World (Piperno et al. 2009).
The presence of starch grains from domesticated
maize and squash, as well as the presumed presence
of beans, in grinding stones by 9,000 BP (Piperno
et al. 2009) indicates that these species were
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simultaneously harvested and consumed and may
have been contemporaneously domesticated. This
possibility is reinforced by records of early and
simultaneous diffusion of maize and chili to South
America (Perry et al. 2007), although a number of
authors have suggested that maize, beans and squash
were domesticated in different regions and periods
(Harlan 1995; Kwak et al. 2009).
According to paleoecological data, clearing and
fire were commonly used between 10,000 and
7,000 cal BP. Initially, fire was probably a tool for
hunting, but probably also became the main strategy
to produce harvests and a strong selective force for
plant species that would become domesticated, a
scenario suggested for other regions in the world
(Lewis 1972; Zong et al. 2007). During the Archaic
period, at least 220 hunter-gatherer groups in what is
today west Mexico and the United Stated used fire to
establish grasslands, guide and enclose animals and
augment forage and grain species production (Parker
2002; Stewart et al. 2002; Williams 2003). Zea,
Phaseolus and Cucurbita species are preadapted to
fire. This element favors colonization and establish-
ment in Zea species, and cyclical fire disturbance
helps them maintain populations (Sánchez-Velázquez
et al. 2002). Fire also helps Phaseolus, Cucurbita and
Capsicum species to recolonize fire-disturbed areas
since their seeds exit latency when exposed to
temperatures greater than 60°C (Rolston 1978;
Degreef et al. 2002). Arboreal species such as
Leucaena spp.; Psidium spp., Prosopis spp. and
Spondias purpurea L. are also preadapted to fire
because of their ability to sprout from the stem base
and root crown. The same holds true for agaves with
the capacity for vegetative propagation because they
can emit root-shoots. Under these circumstances,
humans may have taken advantage of partial removal
of vegetation by fire to cultivate desired grains and
seeds, while the sprouting ability of wild bush species
and humans’ technological inability to eliminate
perennial plant roots dictated use of an agricultural
system with a long fallow period in which both fire-
resistant annuals and perennials were domesticated
and incorporated.
Changes in the maize tga 1 gene were initially
focused on the cob and grain, in an effort to produce
naked, detachable grains (Piperno et al. 2009).
Selection for infructescences that retain ripe grains
increased the likelihood of harvest in the field, while
123
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selection for naked grains facilitated processing by
grinding. Archaeological dates for maize from
Xihuatoxtla (Piperno et al. 2009) do not support the
hypothesis of Harlan (1995), Iltis (2000) or Smalley
and Blake (2003) that initial selection of maize was
linked to use of the stalk and unripe cobs as greens
and as a sugar source in alcoholic beverage produc-
tion. Archaeological records also indicate frequent
use of cooked agave leaf bases and Spondias fruit
about 9,000 BP (Callen 1967; Flannery 1986; Smith
1986), suggesting selection for high sugar content
variants in both plants. These sugar sources became
extremely important in alcoholic beverage production
in the BJ region (Bruman 2000; Colunga-Gar-
cı́aMarı́n and Zizumbo-Villarreal 2007).
Initial domestication of grasses was therefore
linked to ease of harvest and processing. Given that
technological development was similar among human
groups at the time, domestication could have
occurred over a wide area, both in the intermontane
valleys where Setaria parviflora (Poiert) Kerguélen
could have been domesticated (Austin 2006; Callen
1967), and in the lowlands where Z. parviglumis was
domesticated (Ranere et al. 2009). Fixation of
maize’s Tb1 and tga1 genes by about 6,400 BP
suggests that selection continued to be linked to ease
of harvest (e.g. total retention of seeds on the cob,
rachis with two spikelets with two rows of grains) and
processing (e.g. naked grains with short glumes)
(Doebley 2006; Dorweiler et al. 1993; Wang et al.
2005). Simultaneously, the high eruptive activities in
CVC between 7,300 and 6,300 BP might have caused
environmental changes related to the metal content of
local soils that may have been important in maize
domestication (Cortés et al. 2005; Ville-Calzada et al.
2009).
Diffusion of the maize-beans-squash domesticates
suite most likely occurred via existing biocultural
corridors, i.e. rivers, since these would have been
dependable year round food and water sources for
humans and animals, particularly during the long dry
season. The diffusion of domesticated maize and
squash began when they still exhibited low genetic
differentiation with wild varieties. As a result, their
transport to areas lacking wild populations could have
accelerated the fixation of domestic traits and the
disappearance of wild traits. Environmental and
cultural conditions in each region would have
promoted initial diversification.
123
Genet Resour Crop Evol (2010) 57:813–825
Agricultural intensification began to occur
between 7,000 and 5,000 BP as humans actively
worked soils using wooden tools such as levers,
which helped to remove perennial plants, and extract
and arrange rocks to improve soil conditions. This
technique produces a heterogeneous land surface in
which maize, squash and bean seeds are sown
together simulating their natural growth after fire
disturbance in the BJ region. The structuring of the
milpa multi-crop agricultural system occurred
through joint sowing of one or various seeds of these
three species in the same ‘‘microsite’’ within a burned
area, which would have been previously prepared by
removal of roots and rocks using wooden levers.
Additional structuring would have been accom-
plished through individual care and harvest of each
plant. Under these circumstances, these three species
would have been subject to concurrent natural and
human selection pressures that could have led them to
a state of ecological adaptation and complementarity.
This does not coincide with the scenarios proposed
for other regions such as the Middle East, where
preparation of homogeneous planting areas using
draft animals and crop rows allowed use of broadcast
sowing and harvest without human selection of
individual plants. Fixing of the alleles involved in
maize plant architecture by 4,400 cal. BP suggests
that this multicrop system was established by this
time.
Grinding of maize, squash and possibly beans
under domestic conditions suggest the presence of a
selective process that could have produced food
complementarity. Fixation of the alleles involved in
maize protein and starch quality by 4,400 BP may
indicate that this complementarity had been attained
by this time. In the BJ region, agro-food technology
had become highly complex between 4,500 and 3,500
BP, as reflected in the use of pots for steaming, and
probably also distilling, not found in other regions of
Mesoamerica at this time (Zizumbo-Villarreal et al.
2009b). In other words, in this region at this time, the
agro-food system was complete, suggesting that the
milpa system originated here.
Despite the highly complex cultural development
in the BJ region during the rise of plant domestication
and agriculture in Mesoamerica, very few archaeo-
botanical studies have been done there (Benz 2002,
Ranere et al. 2009). New research will be vital to
better understanding the initial development of
Genet Resour Crop Evol (2010) 57:813–825
human societies in Mesoamerica. Study of the TDF in
the BJ region is particularly urgent since almost 80%
of this vegetation type has disappeared to date, and,
of the TDF zones in Mesoamerica, it suffers the
highest deforestation rate and is at the highest risk of
disappearing (Janzen 1988; Kauffman et al. 2003;
Trejo and Dirzo 2000). Native grasses in the region
have experienced strong selective pressure since the
introduction of cattle in the 16th Century. This has
accelerated during the last 30 years as cattle ranching
has grown rapidly and African grasses have been
introduced and managed with pesticides, putting
many of the putative wild source populations,
particularly those of Z. parviglumis, at serious risk
(Houghton et al. 1991; Wilkes 2007). In situ conser-
vation programs at reserves such as Manantlán-Cerro
Grande and Zicuirán-Infiernillo need to be reinforced
and broadened to include nearby areas and human
communities, collection programs need to be stepped
up and adaptive studies of climatic, edaphic and
biotic factors done before these wild ancestral of
domesticated species disappear.
Acknowledgments The authors thank the CONACYT and
CICY for sabbatical scholarships and P. Gepts for his
hospitality at UC-Davis.
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