The cultural and chronological context of early

Holocene maize and squash domestication in the

Central Balsas River Valley, Mexico
Anthony J. Ranerea,b,1, Dolores R. Pipernob,c, Irene Holstb, Ruth Dickaua, and José Iriarted
aDepartment of Anthropology, Temple University, Philadelphia, PA 19122; bSmithsonian Tropical Research Institute, Apartado Postal 0843-03092, Balboa,

Republic of Panama; cArchaeobiology Program, Department of Anthropology, National Museum of Natural History, Washington, DC 20560;
and dDepartment of Archaeology, School of Geography, Archaeology, and Earth Resources, University of Exeter, Exeter EX4 4QJ, United Kingdom

Edited by Jeremy A. Sabloff, University of Pennsylvania Museum of Archaeology and Anthropology, Philadelphia, PA, and approved January 23, 2009
(received for review December 10, 2008)

Molecular evidence indicates that the wild ancestor of maize is

presently native to the seasonally dry tropical forest of the Central
Balsas watershed in southwestern Mexico. We report here on
archaeological investigations in a region of the Central Balsas
located near the Iguala Valley in Guerrero state that show for the
first time a long sequence of human occupation and plant exploi-
tation reaching back to the early Holocene. One of the sites
excavated, the Xihuatoxtla Shelter, contains well-stratified depos-
its and a stone tool assemblage of bifacially flaked points, simple
flake tools, and numerous handstones and milling stone bases
radiocarbon dated to at least 8700 calendrical years B.P. As re-
ported in a companion paper (Piperno DR, et al., in this issue of
PNAS), starch grain and phytolith residues from the ground and
chipped stone tools, plus phytoliths from directly associated sed-
iments, provide evidence for maize (Zea mays L.) and domesticated
squash (Cucurbita spp.) in contexts contemporaneous with and Fig. 1. The study area in northern Guerrero.
stratigraphically below the 8700 calendrical years B.P. date. The
radiocarbon determinations, stratigraphic integrity of Xihuatoxt-
la’s deposits, and characteristics of the stone tool assemblages B.P. (20, 21), and maize phytoliths and pollen dated to 7300 cal
associated with the maize and squash remains all indicate that B.P. from San Andres, Tabasco, on the Caribbean coast (10, 22).
these plants were early Holocene domesticates. Early agriculture in These regions are well outside the present distributional range of
this region of Mexico appears to have involved small groups of the wild ancestor of maize, which, indeed, was not present in
cultivators who were shifting their settlements seasonally and their premaize records. Furthermore, a large corpus of archae-
engaging in a variety of subsistence pursuits. ological and paleoecological microfossil (phytolith, starch grain,
and pollen) research indicates that maize had spread south out
agricultural origins 兩 Mesoamerican preceramic 兩 maize domestication of Mexico earlier—arriving in Panama by 7600 cal B.P.—and by
6000 cal B.P. was well established in northwestern South Amer-

M exico was one of the world’s great centers for the inde-
pendent development of agriculture beginning 10,000
calendrical years B.P. (cal B.P.) (1–3). Among the many plants
ica (e.g., 9, 12, 23–26). This time frame accords with the
estimated ⬇9000 cal B.P. divergence of maize from teosinte
based on a molecular clock (7). It seems clear that documenting
that were domesticated there during the pre-Columbian era, the earliest history and cultural context of maize domestication,
none has received as much attention and been subject to as much together with the role more generally that tropical southwest
debate as corn, or maize (Zea mays L.), the most important crop Mexico played in agricultural origins, requires investigation of
of the Americas (4–12). Once thought to be a cultivar of the arid archaeological deposits in the Central Balsas watershed that date
Mexican highlands, molecular data now indicate that maize was back at least 9,000 years.
domesticated a single time and that a subspecies of teosinte Our field research was carried out within the Central Balsas
classified as Zea mays ssp. parviglumis (Iltis and Doebley) native watershed of northern Guerrero near the modern town of Iguala
to the tropical Central Balsas River Valley, Mexico, is its wild in an area drained by northern tributaries of the Balsas River
ancestor (Fig. 1) (7, 13). Other crop plants that have close wild (Fig. 1). Elevations vary from 700 to 900 m above sea level (asl)
relatives native to the Central Balsas region, and may thus have in the valley bottoms to 1,500–1,800 m asl at the summits of small
been domesticated there, include the silverseeded squash (Cu- ranges that separate the valleys. Rainfall varies from ⬇1,000 to
curbita argyrosperma Huber) and important tree crops (e.g.,
Leucaena spp., Spondias purpurea L.) (14–16). However, previ-
ous archaeological research there has focused on ceramic era Author contributions: A.J.R. and D.R.P. designed research; A.J.R., D.R.P., I.H., R.D., and J.I.
occupations, those dating no earlier than 4,000 years ago, long performed research; A.J.R., D.R.P., I.H., R.D., and J.I. analyzed data; and A.J.R. and D.R.P.
after maize and squash were domesticated and dispersed (17, wrote the paper.

18). Moreover, since the seminal studies of Flannery and Mac- The authors declare no conflict of interest.
Neish (1, 2, 19) in the central and southern Mexican highlands, This article is a PNAS Direct Submission.
there has been little systematic fieldwork on early human See Commentary on page 4957.
settlement and agricultural origins elsewhere in Mesoamerica. 1To whom correspondence should be addressed. E-mail: ranere@temple.edu.
Currently, the earliest evidence for Mexican maize consists of This article contains supporting information online at www.pnas.org/cgi/content/full/
maize cobs from Guilá Naquitz Cave, Oaxaca, dated to 6200 cal 0812590106/DCSupplemental.

5014 –5018 兩 PNAS 兩 March 13, 2009 兩 vol. 106 兩 no. 13 www.pnas.org兾cgi兾doi兾10.1073兾pnas.0812590106
 SEE COMMENTARY
Fig. 2. A view of the enormous boulder that formed the Xihuatoxtla Shelter.

1,400 mm annually with over 90% falling during the June-to- B.P.), but the reworking of the rock shelter deposits in the early
October rainy season. The potential vegetation of the region is 20th century has left them thoroughly mixed over most of the
tropical deciduous forest, seen today in remnants scattered site. At a fourth site, the Temaxcalapa Shelter, we recovered a
throughout the region. Vegetational reconstructions indicate large number of chipped stone artifacts from the eroded slope in
that a species-diverse seasonal tropical forest flourished there front of the shelter, including preceramic-aged spear points. We
after the Pleistocene ended and until systematic human distur- were unable to excavate this site (see SI Text, Site Descriptions for
bance began (27). The native vegetation contains legumes and additional information on these 4 sites). In a fifth site, the
many other plants that would have been important subsistence Xihuatoxtla Shelter, our excavations encountered undisturbed
resources, such as Pithecolobium dulce, Prosopis spp, Leucaena preceramic and ceramic deposits containing both chipped and
spp., Spondias spp., palms, Dioscorea spp., and others (27). ground stone tools, plant remains, and ceramics (see Table S1 for
Bedrock in the study area is predominantly limestone, although 14C dates from these sites).

extrusive igneous formations are found in the eastern portion of

the area. Both limestone and igneous formations provide caves Results
and overhangs that early occupants of the region used as shelters. The Xihuatoxtla Shelter (964 m asl) is formed by the overhang
The karstic landscape also resulted in the formation of a of an enormous boulder (17 ⫻ 16 ⫻ 14 m) and contains ⬇75 m2
number of permanent lakes near the shelters that offered a of protected floor space inside the dripline (Fig. 2 and Fig. S1).
stable and abundant supply of animal protein as well as diverse It is located 260 m from Barranca Xihuatoxtla near the point
plant foods (27). where the now-seasonal stream opens out onto a flat basin. The
Our survey focused on caves and rock shelters because they nearly 1-m-deep stratigraphic sequence (Fig. 3) is anchored by
were commonly used by ancient peoples and because they 4 radiocarbon dates on charcoal: (1) 1200 ⫾ 40 B.P. (1240–1000
provide better protection for perishable materials, such as plant cal B.P.) from layer B, 20–30 cm below surface, (2) 2790 ⫾ 40
remains, than open-air sites. We located 15 caves and rock B.P. (2970–2780 cal B.P.) from layer B, 30–40 cm below surface,
shelters that had been occupied sometime in prehistory, based (3) 4730 ⫾ 40 B.P. (5590–5320 cal B.P.) from layer C, 49 cm
on the stone and ceramic artifacts recovered from the site below surface, and (4) 7920 ⫾ 40 B.P. (8990–8610 cal B.P.) from
surfaces, and carried out excavations in 4 of these sites (Fig. 1). layer D, 65 cm below surface. The cultural deposits extended
Two of the sites, Cueva del Agua and El Abra 2, contained only another 35 cm below the 8990–8610 cal B.P. level, but we were
ceramic-aged deposits. A third site, El Abra, was occupied at unable to date any of the samples recovered from these earlier
least as early as 8330–8160 cal B.P. (14C AMS date of 7400 ⫾ 40 deposits. Though starch grains and phytoliths were well pre-

ANTHROPOLOGY

Fig. 3. Stratigraphy of units 1 and 2 in the Xihuatoxtla Shelter.

Ranere et al. PNAS 兩 March 13, 2009 兩 vol. 106 兩 no. 13 兩 5015
served in stone tool residues, and phytoliths commonly occurred
in the shelter sediments, pollen grains and plant macrofossils
other than wood charcoal were scarce (28). No faunal remains
were preserved. The sedimentary sequence at Xihuatoxtla can
be divided into 5 major and discrete stratigraphic units (Fig. 3).
Layer A (⬇0–10/15 cm below surface) is a 10- to 15-cm-thick
deposit of very compact tan silt that contained numerous
pre-Columbian potsherds (n ⫽ 414) and a modest number (n ⫽
26) of lithic artifacts, primarily debitage but including a small
tanged obsidian point and an obsidian blade fragment. A small
number of bottle glass fragments (n ⫽ 3) are the only indication
of recent disturbance (all counts given here and below are for
units 1 and 2, contiguous 1 m2 blocks).
Layer B (⬇10/15–45/50 cm b.s.) is a 30- to 40-cm-thick deposit
of clayey silt with small amounts of angular roof-fall that changes
from gray/brown to brown in color with depth. Sherd densities
vary from 110 sherds per 5-cm level per square meter at the top
of the layer to 25 sherds per 5-cm level per square meter at the
bottom. Lithic densities averaged 27 specimens per 5-cm level
per square meter. Obsidian artifacts included 12 prismatic blade
fragments, 1 tanged point, 1 tabular wedge (piece esquillée), and
9 flakes. Other chipped stone artifacts of local materials included
1 steep scraper, 1 flake knife, 1 graver, 1 chopper, and 1 small
bipolar core. Also recovered from the layer were 3 handstones
(manos, small implements used for grinding plant and possibly
other kinds of materials), 2 milling stone base fragments, and 1
shaped metate fragment. Fig. 4. Handstones and milling stone bases from the preceramic layers in the
Layer C (⬇45/50–60 cm b.s.) is an 8- to 12-cm-thick deposit Xihuatoxtla Shelter. (A) Small handstone (318e) from layer E that yielded 80
of large angular roof-fall blocks (cobble sized) in a matrix of maize starch grains, maize cob phytoliths, and 29 squash phytoliths. (B)
gray/brown clayey silt. The density of lithic artifacts (119 per Complete milling stone base (316d) from layer D that yielded 68 maize starch
5-cm level per square meter), absence of obsidian blades, and the grains as well as 4 yam (Dioscorea sp.), 3 legume, and 1 Marantaceae starch
AMS 14C date of 4730 ⫾ 40 B.P. (5590–5320 cal B.P.) indicate grains. (C) Handstone fragment (318d) from layer E that yielded 22 maize
that this layer dates to the preceramic, although there is some starch grains, maize cob phytoliths, and 28 squash phytoliths. (D) Handstone
(322c) from layer E that yielded 11 maize starch grains, maize cob phytoliths,
intrusion of sherds from the ceramic occupation above (n ⫽ 36).
and 7 squash phytoliths. (E) Small handstone (365a) from layer C that yielded
The chipped stone tools recovered from layer C include 2 scraper 24 maize starch grains and maize cob phytoliths. (F) Handstone (319d) from
planes, 1 borer, 1 flake knife, 1 bifacial core, 2 bipolar cores, 1 layer E that yielded 8 maize starch grains, maize cob phytoliths, and 37 squash
small obsidian biface fragment, and 4 used flakes. There were phytoliths. (G) Slab milling stone fragment (316c) from layer D that yielded 2
also 6 handstones (see Fig. 4 and Fig. S2 A) and 1 possible milling maize starch grains, maize cob phytoliths, and 29 squash phytoliths.
stone base fragment recovered from layer C.
Layer D (⬇60–70 cm b.s.) is an 8- to 10-cm-thick deposit of
brown clayey silt with some angular roof-fall blocks. There were ments but where they occur they are aligned in parallel to each
251 chipped stone artifacts recovered (63 per 5-cm level per m2) other, suggesting a back and forth motion. An exception is the
and no ceramics. The chipped stone tools recovered from layer complete milling stone recovered from layer D (316d) that
D include 1 cobble spall chopper, 1 concave scraper, 2 gravers, exhibited randomly oriented striations near the edges of a
5 flake knives, and 1 biface fragment. Also recovered were 2 shallow depression 11 cm in diameter. The use of unmodified
handstones, and 1 complete and 2 fragmentary milling stone river cobbles as handstones and unmodified river boulders or
bases (see Fig. 4 and Fig. S2 B and C). stone slabs as milling stone bases is a common pattern observed
Layer E (⬇70–95 cm b.s.) is an 8- to 25-cm-thick deposit of in early Holocene sites in the semiarid Mexican highlands in the
brown clayey silt with more angular roof-fall blocks than layer D. Tehuacan Valley (29) and at Guilá Naquitz in Oaxaca (1).
It rests on eroded bedrock (layer F) that slopes from 78 to 95 cm Similar ground stone tools were also recovered in early Holocene
below surface moving from south to north (see Fig. 3). There contexts at the Santa Marta Rock Shelter (30), a site in the
were 179 chipped stone artifacts recovered (30 per 5-cm level per Central Chiapas Depression that sits at an elevation of 860 m in
square meter), including the base of a stemmed, indented base a zone of tropical deciduous forest.
point, the distal end of a thick lanceolate point (Fig. 5), 1 graver, Chipped stone tool manufacturing at Xihuatoxtla was an
1 spokeshave, 1 flake knife, 9 used flakes, 5 thinning flakes with important activity during the preceramic period and continued
ground platforms, 2 core fragments, and 3 bipolar cores. Four into the ceramic period, albeit with less intensity (Table S2). A
handstones were recovered from this earliest occupation layer wide range of raw materials was used, including chalcedony,
(Fig. 4 and Figs. S2D and S3). chert, quartzite, rhyolite, basalt, sandstone, and obsidian. All but
Nearly all of the handstones and milling stone bases (Fig. 4) the obsidian appear to be locally available. At the base of the
used for plant processing consisted of river cobbles and boulders sequence in layer E (⬇75–95 cm below surface), evidence for
that were modified only by use. One milling stone base fragment bifacial reduction in the production of projectile points is
came from a stone slab (Fig. S2C) rather than a river boulder that indicated by thinning flakes oftentimes containing ground plat-
was also modified only by use. Wear polish can occur on one or forms. The preparation of the edge of a spear point preform by
both faces of a handstone and on the tool edges as well. One grinding is a way of increasing platform strength and allowing
handstone from layer E (318e) appears to have seen consider- larger thinning flakes to be removed. Bifacial thinning flakes
able use as a pestle. A number of the milling stone fragments with ground platforms are characteristic of Paleoindian and
exhibit polish on their flat or slightly concave surfaces as well. Early Archaic periods (before ⬇8000 cal years B.P.) (31), and
Striations are rare on both handstones and milling stone frag- their appearance in layer E is consistent with its estimated age

5016 兩 www.pnas.org兾cgi兾doi兾10.1073兾pnas.0812590106 Ranere et al.

 SEE COMMENTARY
As reported in a companion paper (28), starch grains identi-
fied as maize were recovered, often in large numbers, from most
artifacts identified as grinding stones found throughout the
Xihuatoxtla stratigraphic sequence. All 8 of the grinding tools
and both of the chipped stone tools examined for starch grains
in levels contemporary with or below the 14C date of 7920 ⫾ 40
B.P. (8990–8610 cal years B.P.) yielded maize starch. Maize
phytoliths were recovered from both sediments and stone tools
throughout the sequence, including those beneath and from the
level that produced the 7920 ⫾ 40 14C B.P. date. Phytoliths
identified as domesticated Cucurbita, possibly C. argyrosperma,
were also found in direct association with the 7920 ⫾ 40 14C B.P.
date at Xihuatoxtla and in levels below (see ref. 28 for details on
starch grain and phytolith recovery and identification). Thus, the
evidence firmly indicates that maize and squash were domesti-
cated by 8990–8610 cal years B.P., the earliest date yet recorded
for maize.

Discussion
Although Mexico played a central role in New World agricul-
tural origins, contributing a large number of important crop
plants in addition to maize (9), little research directed toward
understanding early human occupation and agricultural evolu-
tion has been carried out in its lowland tropical regions. Our
investigations in northern Guerrero show that this region of
tropical southwest Mexico supported persistent human occupa-
tion beginning by at least 9,000 years ago and possibly a few
thousand years earlier, if some of the stone tools recovered from
the surface of the Temaxcalapa Shelter turn out to indeed be
Clovis in age. The archaeobotanical (28) and stone tool evidence
from Xihuatoxtla indicates that plant exploitation and cultiva-
tion were important activities from the earliest stages of the site’s
occupation.
Xihuatoxtla is a small shelter that appears to have been
repeatedly visited by small groups of people who stayed for
significant periods of time (several weeks or more) based on the
nature and density of materials left at the site. During the
preceramic period, these groups likely shifted locations season-
ally. Our data on early settlements from the Central Balsas
Fig. 5. Bifacial points and preforms from north central Balsas Valley sites. (A) region are insufficient to document this pattern in any detail.
‘‘Pedernales’’ point base from Xihuatoxtla, layer E. (B) Point from Xihuatoxtla, Nonetheless, the evidence we do have suggests that preceramic
layer E (distal end). (C) Point base from El Abra. (D) Reworked stemmed point groups were occupying different settings and were engaged in
from Temaxcalapa. (E) Preform fragment from Temaxcalapa. (F) Point base different subsistence pursuits. The much larger El Abra Shelter,
from Temaxcalapa. (G) Preform base from Temaxcalapa. only 7 km distant from Xihuatoxtla and occupied over much of
the same time span, included both small and large mammal bone
in the limited sample of undisturbed deposits that were exca-
(older than ⬇9000 cal years B.P.) The basal half of a stemmed, vated in that site. The near absence of grinding stones at this site
indented base point and the distal end of a diamond cross- suggests that activities were directed more toward hunting than
sectioned point were also recovered from this layer (Fig. 5). The plant exploitation and cultivation. The surface collections from
point base is similar to single examples classified as Pedernales the Temaxcalapa Shelter, located at a higher elevation (1,350 m
points from the Guilá Naquitz site in Oaxaca and from the asl) than Xihuatoxtla (964 m asl) and in more broken terrain
Tehuacan Valley in contexts dated between ⬇9510 and 7742 cal (Fig. 1), contained Archaic point types (Fig. 5) and other chipped
years B.P. (1, 29). Evidence of bifacial reduction continued in stone tools (n ⫽ 38) and flakes (n ⫽ 128) but no grinding tools,
layer D (⬇60–75 cm below surface) with the recovery of hinting again at a different set of activities carried out there than
additional thinning flakes and also one biface preform fragment. at Xihuatoxtla.
Our paleoecological data from nearby lakes (Fig. 1) indicate
Bifacial flaking occurred in layer C and in layer B associated with
that utilization of resource-rich lacustrine environments was an
ceramics, albeit at reduced levels and with no platform prepa-
early and important component of subsistence and probably
ration by grinding. seasonal settlement cycles (27). At Laguna Tuxpan, phytolith
Of equal or greater importance in stone tool manufacturing data indicate that humans were exploiting lake edge settings and
was core reduction for flakes that could be used with little or no modifying the local vegetation, including with fire, during the
modification. Cores were reduced opportunistically or by bipolar early Holocene. Phytoliths of maize and domesticated Cucurbita
methods. The resulting flakes were used as knives, scrapers,
ANTHROPOLOGY

plus Zea pollen were also recovered and minimally dated to 6000
borers, spokeshaves, and gravers and represented the majority of cal years B.P., with a likely actual early Holocene age (27). At the
the tools recovered in the preceramic deposits. Tools recovered Ixtacyola lake bed, chipped stone artifacts occurred in lake edge
from ceramic-aged deposits were mainly made on obsidian sediments to a depth of 4.3–4.4 m, 10 cm below a date of 6290 ⫾
prismatic blades, although some tools and the bulk of the 40 14C years B.P. (7280–7170 cal B.P.) (27). These tools are
chipping debris were of local nonobsidian materials. accompanied by a major increase in pollen disturbance indica-

Ranere et al. PNAS 兩 March 13, 2009 兩 vol. 106 兩 no. 13 兩 5017
tors and charcoal in the paleoecological records. The paleoeco- and the deposits screened through 1/4- and 1/16-inch screens. Major artifacts
logical data suggest that maize and squash were being planted in were measured in situ, and those that were potentially used for plant pro-
the productive soils near lake edges that were exposed during the cessing were immediately sealed in plastic bags to avoid any possibility of
dry season as lake levels fell. These practices would have contamination of plant residues. Sediment samples from the immediate vi-
provided attractive yields for minimal effort. cinity of potential plant processing tools were taken and cataloged along with
Our data from the Central Balsas provide a firm cultural and the associated tool. A 20 ⫻ 20-cm-column sample was removed from the north
chronological context for early domesticated maize and a species wall of unit 1 to a depth of 1 m for the purpose of extracting plant microfos-
of squash in their postulated Mexican homelands. They also add sils—phytoliths, pollen, and starch grains—if present.
to the increasing archaeological and molecular evidence attest-
ing to a primary role of seasonal tropical forests from Mexico to ACKNOWLEDGMENTS. We thank 3 anonymous reviewers for their helpful
comments on the manuscript. We also thank the Instituto Nacional de Antro-
Brazil in the origins and dispersals of New World domesticated
pología e Historia for permission to carry out the research and Froylan Cuenca,
crops (e.g., 9, 15, 23, 26, 32). the Xihuatoxtla Shelter landowner, for his support. This research was funded
by the National Science Foundation (Grant BSC 0514116), the National Geo-
Materials and Methods graphic Society (Grant 7748 – 04), the Wenner-Gren Foundation (Grant 7299),
Four contiguous 1-m2 units were opened at one edge of the Xihuatoxtla the Smithsonian National Museum of Natural History, Smithsonian Tropical
Shelter. The excavations were largely carried out with hand tools in 5-cm levels Research Institute, and the College of Liberal Arts, Temple University.

1. Flannery K (1986) Guilá Naquitz (Academic, Orlando). 18. Smith BD (1997) Reconsidering the Ocampo caves and the era of incipient cultivation
2. MacNeish RS (1967) in Environment and Subsistence, Vol 1: The Prehistory of the in Mesoamerica. Lat Am Antiq 8:342–383.
Tehuacan Valley, ed Byers D (Univ Texas Press, Austin), pp 290 –309. 19. Byers DS (1976) The Prehistory of the Tehuacan Valley, Vol 1: Environment and
3. Smith BD (1997) The initial domestication of Cucurbita pepo in the Americas 10,000 Subsistence (Univ Texas Press, Austin).
years ago. Science 276:932–934. 20. Piperno DR, Flannery KV (2001) The earliest archaeological maize (Zea mays L.) from
4. Beadle GW (1977) in Origins of Agriculture, ed Reed CE (Mouton, The Hague), pp Highland Mexico: New accelerator mass spectrometry dates and their implications.
615– 635. Proc Natl Acad Sci USA 98:2101–2103.
5. Iltis H (2000) Homeotic sexual translocations and the origin of maize (Zea mays, 21. Benz B (2001) Archaeological evidence of teosinte domestication from Guilá Naquitz,
Poaceae): A new look at an old problem. Econ Bot 54:7– 42. Oaxaca. Proc Natl Acad Sci USA 98:2104 –2106.
6. Mangelsdorf PC (1974) Corn: Its Origins, Evolution, and Improvement (Harvard Univ 22. Pohl MED, Piperno DR, Pope KO, Jones JG (2007) Microfossil evidence for pre-
Press, Cambridge). Columbian maize dispersals in the neotropics from San Andrés, Tabasco, Mexico. Proc
7. Matsuoka Y, et al. (2002) A single domestication for maize shown by multilocus Natl Acad Sci USA 104:6870 – 6875.
microsatellite genotyping. Proc Natl Acad Sci USA 99:6080 – 6084. 23. Aceituno FJ, Castillo N (2005) Mobility strategies in Colombia’s middle mountain range
8. Bennetzen J, et al. (1991) Genetic evidence and the origin of maize. Lat Am Antiq between the early and middle Holocene. Before Farming 2005/2, article 2. Available at
12:84 – 86. www.waspress.co.uk/journals/beforefarming/journal㛭20052/abstracts/index.php.
9. Piperno DR, Pearsall DM (1998) The Origins of Agriculture in the Lowland Neotropics 24. Bush MB, Piperno DR, Colinvaux PA (1989) A 6,000 year history of Amazonian maize
(Academic, San Diego). cultivation. Nature 340:303–305.
10. Pope KO, et al. (2001) Origin and environmental setting of ancient agriculture in the 25. Pearsall DM, Chandler-Ezell K, Zeidler JA (2004) Maize in Andean Ecuador: Results of
lowlands of Mesoamerica. Science 292:1370 –1373. residue analysis of stone tools from the Real Alto Site. J Arch Sci 31:423– 442.
11. Benz, BF (2006) in Histories of Maize: Multidisciplinary Approaches to the Prehistory, 26. Piperno DR, Ranere AJ, Holst I, Hansell P (2000) Starch grains reveal early root crop
Linguistics, Biogeography, Domestication and Evolution of Maize, eds Staller J, Tykot horticulture in the Panamanian tropical forest. Nature 407:894 – 897.
R, Benz B (Academic, San Diego), pp 9 –20. 27. Piperno DR, et al. (2007) Late Pleistocene and Holocene environmental history of the
12. Zarillo S, Pearsall DM, Raymond JS, Tisdale MA, Quon DJ (2008) Directly dated starch Iguala Valley, Central Balsas Watershed of Mexico. Proc Nat Acad Sci USA 104:11874 –
residues document early formative maize (Zea mays L.) in tropical Ecuador. Proc Natl 11881.
Acad Sci USA 105:5005–5011. 28. Piperno DR, Ranere AJ, Holst I, Dickau R, Iriarte J (2009) Starch grain and phytolith
13. Doebley J (2004) The genetics of maize evolution. Ann Rev Genet 38:37–59. evidence for early ninth millennium B.P. maize from the Central Balsas River Valley,
14. Sanjur OI, Piperno DR, Andres TC, Wessel-Beaver L (2001) Phylogenetic relationships Mexico. Proc Natl Acad Sci USA 106:5019 –5024.
among domesticated and wild species of Cucurbita (Cucurbitaceae) inferred from a 29. MacNeish RS (1976) in The Prehistory of the Tehuacan Valley, Vol 3: Lithics (Univ Texas
mitochondrial gene: Implications for crop plant evolution and areas of origin. Proc Natl Press, Austin).
Acad Sci USA 99:535–540. 30. MacNeish RS, Peterson FA (1962) The Santa Marta Rock Shelter, Ocozocoautla,
15. Miller A, Schaal B (2005) Domestication of a Mesoamerican cultivated fruit tree, Chiapas, Mexico. Papers of the New World Archaeology Foundation 14 (Brigham
Spondias purpurea. Proc Natl Acad Sci USA 102:12801–12806. Young Univ, Provo, UT).
16. Hughes CE, et al. (2007) Serendipitous backyard hybridization and the origins of crops. 31. Ranere AJ (2006) in Paleoindian Archaeology: A Hemispheric Perspective, eds Morrow
Proc Natl Acad Sci USA 104:14389 –14394. J, Gnecco C (Univ Florida Press, Gainesville), pp 69 – 85.
17. Niederberger C, Rayna Robles RM (2002) The Archaeological Past of Guerrero (original 32. Dillehay TD, Rossen J, Andres TC, Williams DE (2007) Preceramic adoption of peanut,
title in Spanish) (Consejo Nacional para la Cultura y las Artes, Mexico City). squash and cotton in northern Peru. Science 316:1890 –1893.

5018 兩 www.pnas.org兾cgi兾doi兾10.1073兾pnas.0812590106 Ranere et al.