Comparative Analysis of the Río Balsas and Tehuacán Models for the Origin of Maize

Author(s): Richard S. MacNeish and Mary W. Eubanks

Source: Latin American Antiquity , Mar., 2000, Vol. 11, No. 1 (Mar., 2000), pp. 3-20
Published by: Cambridge University Press

Stable URL: https://www.jstor.org/stable/1571668

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 ARTICLES

COMPARATIVE ANALYSIS OF THE RIO BALSAS AND TEHUACAN

MODELS FOR THE ORIGIN OF MAIZE

Richard S. MacNeish and Mary W. Eubanks

This paper examines the archaeological and biological evidence for shifts in human subsistence str
tion from hunting and foraging to maize agriculture as posited in the Rio Balsas, or lowland origi
Tehuacdn, or highland origin of maize, model. These are two different interpretations of the genetic
of maize, the archaeological evidence for plant exploitation, and the ecological evidence for paleo
change in the two regions. In contrast to Panama, where there is good evidence for progressive inten
disturbance by 10,000 B.P, horticulturalforest clearing by 8000 B.P, and slash-and-burn agricultu
dencefor Mesoamerica, where maize agriculture originated, fits a different picture of biocultural evolu
of Mexico, Guatemala, Belize, and probably Honduras, were apparently undisturbed, semi-evergre
B.P. New findings from experimental maize genetics, combined with the comprehensive archaeologica
Oaxaca, Tamaulipas, and the Valley of Mexico, support a highland Mesoamerican origin of maize.

Este articulo examina la evidencia arqueol6gica y biol6gica sobre los cambios ocurridos durante la tran
subsistencia de la caza y recoleccion a la agricultura del maiz en el Rio Balsas o modelo del origen del m
y el modelo de Tehuacdn del origen del maiz en las tierras altas. Estos modelos constituyen dos interp
evidencia genetica acerca del ancestro del maiz, la evidencia arqueologica de la explotaci6n de la pla
ica acerca del paleo-ambiente y cambio climdtico en estas dos regiones. En contraste con Panama, dond
sobre la intensificaci6n progresiva de perturbaciones humanas sobre la selva alrededor de 10.000 AP., l
horticultura alrededor de 8000 A. P, y la agricultura de tala y quema alrededor de 6000 A. R, la eviden
lugar del origen del maiz se adecua a un esquema de evoluci6n biocultural diferente. Las tierras ba
Belize y probablemente Honduras aparentemente eran selvas semideciduas no perturbadas alrededor
tados de un estudio experimental del cultivo del maiz han proporcionado ahora nueva evidencia sobre e
derivados de un cruce de teosinte (Zea diploperennis), una especie de diploide de las tierras altas m
de oriente (Tripsacum dactyloides), segregando asilaprogenie hibrida que es muyparecida a los restos ar
del maiz que se han recuperado de las cuevas secas de Tehuacdn. La evidencia comprehensiva que com
de genetica experimental del maiz con un esquema completo de la arqueologia de Tehuacdn, Oaxaca,
de Mexico, apoya la teoria del origen del maiz en las tierras altas de Mesoamerica.

T wo divergent and often hotly debated mod-

shells and borne in a spike composed of five t
els, centered around the question ofindividual
the ori- units arranged in a vertical row th
gin of maize (Zea mays ssp. mays), the upon maturity for natural disper
ters apart
maize ear
keystone crop plant upon which Mesoamerican bears hundreds of exposed kerne
civ-
ilization was founded, have been proposed
firm
to explain
rachis called a cob, and the entire seed-
the origins of agriculture in Mesoamerica. The ear
structure is enclosed in multiple leaf sheath
of maize is a unique morphological structure
husks.
unpar-
Maize has been so radically modified b
alleled in the botanical kingdom. In contrast to its
ficial selection that the plant is no longer cap
wild relatives, whose seeds are encased innaturally
hard outer reproducing itself and would go

Richard S. MacNeish * Andover Foundation for Archaeological Research, PO Box 83, Andover, M
Mary W. Eubanks * Department of Botany, Duke University, Durham, NC 27708-0338

Latin American Antiquity, 11(1), 2000, pp. 3-20

Copyright ? 2000 by the Society for American Archaeology

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4 LATIN AMERICAN ANTIQUITY [Vol. 11, No. 1, 2000

without human intervention to plant, protect, and to both hybrid versions, annual teosintes
According
harvest the seed. Although maize geneticists
andhave
domesticated maize derived from introgression
between an extinct maize and its wild ancestors. The
identified and mapped five major genes responsible
for this transformation to the maize ear (Doebley
debate et
about the origin of maize is important because
al. 1990; Galinat 1977,1985; Langham 1940; aMan-
clear understanding of the biological parameters
of maize evolution will enable better resolution of
gelsdorf 1947; Rogers 1950), how this anomalous
form originated is still a scientific puzzle. Specimens
the respective roles played by humans and environ-
that show a stepwise transition from a shattering
mental change in crop plant domestication and the
origins
spike to the many-rowed ear of cultivated maize haveof agriculture.
not been identified in the archaeological record,New
anddata from molecular systematics and exper-
experiments to reconstruct the maize progenitorimental
from plant breeding call for a reexamination of
crosses between maize and annual teosinte have current models of the origins of Mesoamerican agri-
culture. First, a nuclear DNA study of ribosomal
failed to recover pure segregating parental pheno-
types. What is known from archaeology is thatinternal
the transcribed spacer (ITS) sequences (Buck-
macroevolutionary event leading to this radical
ler and Holtsford 1996) indicates that maize diverged
either before or contemporaneous with Balsas
change in morphology occurred within the last
teosinte (Z. m. ssp. parviglumis) and Chalco teosinte
10,000 years, a short span of time for such profound
biological modification to occur solely under natural
(Z. m. ssp. mexicana) on the phylogenetic tree. Log-
selection. Another remarkable feature of maize evo- ically, for either of these annual teosintes to be a
lution is that subsequent introgression from back-progenitor of maize (Doebley 1990; Galinat 1977,
crossing with the wild ancestors and adaptive1985), they should appear earlier, rather than con-
radiation into new ecological niches gave rise to temporaneous with or later than maize. A second
extensive biodiversity as evidenced by the many dif-development is demonstration of cross compatibil-
ferent maize races that had appeared around 1750 ity between teosinte and Tripsacum. Eubanks (1995,
B.P. (Eubanks 1999a). 1997a) obtained fully fertile hybrids by crossing
One hypothesis, referred to here as the Rio Bal- diploid perennial teosinte (Zea diploperennis) and
sas (or lowland) model, argues that maize derivedEastern gamagrass (Tripsacum dactyloides). Some
from mutations in annual teosinte (Zea mays ssp. segregating progeny from her crosses are remarkably
parviglumis), which grows in the Balsas Riversimilar in ear morphology to the oldest archaeolog-
drainage in Guerrero, Mexico (Benz 1999; Doebley ical maize remains (Mangelsdorf et al. 1967a).
1990; Pipemo and Pearsall 1998; Smith 1995). This This paper reexamines how the basic tenets of the
view, based on molecular data from isozyme and Rio Balsas lowland and Tehuacan highland models
chloroplast DNA studies in the 1980s, showed thatfor the origin of maize agriculture in Mesoamerica
parviglumis teosinte is the species most closely fit the archaeological and botanical evidence in light
related to maize, and assumes phylogenetic descentof these recent developments, and discusses how
from the species with the greatest number of sharedthey may support, or fail to support, each of the pro-
genes. An alternate view, referred to as the Tehuacanposed schemes. For purposes of organizing the pre-
(or highland) model (Mangelsdorf et al. 1967a),sentation, the discussion is structured in the context
argues that maize originated in the highlands ofof the standard temporal framework for the Archaic
Mesoamerica as a result of hybridization between period of Mesoamerican prehistory, divided into the
two wild relatives of maize. The hybrid origin wasfollowing chronological phases: Stage I, before 7500
initially thought to have involved Tripsacum and an B.P.; Stage II, 7500-7000 B.P.; Stage III, 7000-6500
extinct wild maize (Mangelsdorf and Reeves 1939);B.P.; Stage IV, 6500-6000 B.P.; Stage V, 6000-5500
this view was later modified to attribute the origin B.P.; and Stage VI, 5500-4500 B.P.
of annual teosinte to a hybrid between diploid peren-
nial teosinte and maize in the early stages of domes-
Archaeological Models
tication (Mangelsdorf et al. 1981; Mangelsdorf 1983,
The Rio Balsas (or Lowland) Model
1986; Wilkes 1979), with domesticated maize evolv-
ing from subsequent introgressive hybridizationThe idea of a tropical Guatemalan hearth of agri-
between maize and the new created annual teosintes. culture originated with Vavilov (1952). Concomitant

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MacNeish and Eubanks] MODELS FOR THE ORIGINS OF MAIZE 5

V_ 7500-7000 B.P.I

C e " bbea

Stre

-a1 a__

Boundaries Based On Current Evidence

Figure lb. In Stage II of the Rio Balsas model of origin
and domestication of maize, parviglumis teosinte mutated
- - - - Hypothesized Biogeographical Boundariesto become maize. No direct evidence or dates. Indirect
evidence shows Mesoamerican tropics with undisturbe
[\\\N\ Boundaries Inferred From Microfossil Datasemi-evergreen forest while Panama had small horticu
tural clearings.
Figure la. In Stage I of the Rio Balsas model of origin and
domestication of maize, parviglumis teosinte from the Rio
Balsas lowlands was the ancestor of maize based on there was similar exploitation and manipulation of
genetic (allozyme and cpDNA) studies of modern
otherteosinte.
plants that grew in this area that would also
No direct evidence or dates. Indirect evidence from
be domesticated, including beans (Phaseolus vul-
Mesoamerican tropics with undisturbed semi-evergreen
garis) (Gepts
forest while Panama has progressive intensification of et al. 1986), possibly sweet potato
human forest disturbance. (Ipomoea batatas), manioc (Manihot esculenta),
and squash, which evolved from Cucurbita sorovia
into C. argyrosperma.
with this concept and the "age-area" hypothesis pop-
ular at the time, Sauer (1952) predicted future
Palynological profiles are viewed as signaling
Mesoamerican research would show that agriculture
clearing of tropical forests and are interpreted as evi-
began on the wet Pacific slopes of Mexico wheredence for early agricultural activity during this tem-
many varieties of beans, maize, and squash grew.
poral stage in Panama (Piperno et al. 1991) and a
This hypothesis was revived by Pipero (1989), parallel
who development is proposed to have occurred
in Mesoamerica (Pipero and Pearsall 1993, 1998).
proposed that agriculture emerged in the Panaman-
ian tropical forest ca. 10,000-5000 years B.P. However,
Fol- no archaeological or paleoecological evi-
lowing the same theme, Pipero and Pearsall (1993,
dence has yet emerged from the Rio Balsas region
that supports early domestication of maize there
1998) have suggested there was a parallel develop-
ment in Mesoamerica in the Rio Balsas region. Cen-
(Benz 1999). In their search for early maize in 1959,
Garcia Cook and MacNeish selected Tehuacan
tral to this model for the emergence of Mesoamerican
agriculture is the hypothesis that the ancestor of
instead of Guerrero partly because, although they dis-
maize is a wild annual teosinte (Zea mays ssp. parvig-
covered huge shell mounds with probable long occu
lumis) species that grows in the Rio Balsas drainage
pations in Guerrero, there were few dry caves with
of Guerrero (Doebley 1990). Other proponents of preserved
this archaeobotanical remains. Pollen profile
view of the origin of maize include Benz (1994, from Lake Pet6n in Guatemala and Belize (Islebe e
1999), Blake et al. (1992), Pohl et al. (1996), Rust
al. 1996; Jones 1991, 1994) show "undisturbed sem
and Leyden (1994), Smith (1995), and Voorhies et
evergreen forests," rather than evidence of "pro
al. (1991). gressive intensification of human forest disturbanc
Stage I: Before 7500 B.R According to the Rio (Piperno and Pearsall 1998:305), as occurred i
Balsas model, during Stage I (Figure la) the annual Panama. Furthermore, Mesoamerica, unlike Centr
teosinte (Zea mays ssp. parviglumis) that is the puta- America, shows evidence of a younger Dryas even
tive progenitor of maize was growing in the tropi- with significant climatic changes that were not occur
cal lowlands of the Rio Balsas area, where it was ring in Panama. The present evidence suggests tha
harvested and possibly cultivated by human hunters Panama and the Mesoamerican lowlands are not
and gatherers. In addition to parviglumis teosinte, analogous situations.

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6 LATIN AMERICAN ANTIQUITY [Vol. 11, No. 1, 2000

Stage 111
'liE

Boundaries Based On Current Evidence Figure ld. In Stage IV of the Rio Balsas model of origin
and domestication of maize, maize had spread across
Isthmus of Tehuantepec to Olmec region of the Gulf coast
- - - - Hypothesized Biogeographical Boundaries
and reached Panama. Evidence consists of Zea pollen in
Panama; supposedly no maize in highlands as hypothesis
L\\\] Boundaries Inferred From Microfossil Data
relies on AMS dates of Tehuacan maize specimens rather
than 14C dates from well-documented stratigraphy.
Figure Ic. In Stage III of the Rio Balsas model of origin
and domestication of maize, early maize spread across
Isthmus of Tehuantepec to La Venta region of Tabasco the Tehuacan Valley (Flannery and MacNeish 1997;
and then moved southward toward Panama, which had
cleared forest areas with non-maize horticulture.
Mangelsdorf et al. 1967a). Available archaeological
evidence indicates that during Stage ml, the lowland
Stage II: 7500-7000 B.R Stage II (Figure Ib) is
Mesoamerican peoples apparently continued their
maritime orientation (Voorhies 1976; Wilkerson
projected as the era when Balsas teosinte was trans-
1975). This finding contrasts with Panama where
formed into maize via five mutations in the grain-
bearing structure. These mutant forms lost the traits
"small [non-maize] horticultural clearings in the for-
critical for their survival in nature and had to be est are indicated" (Pipemo and Pearsall 1998:305).
selected and preserved by humans who harvested and Stage IV: 6500-6000 B.R By Stage IV (Figure
Id), microfossil evidence from Cueva de los
eventually planted them to establish a convenient,
nutritious food supply. No archaeological remains Ladrones
of in Panama (Pipero 1985) suggested the
maize cobs or Zea microfossils have been reported
possibility that Zea (and perhaps other plants) trav-
eled southeast from the tropical lowlands of Guer-
from Guerrero during Stage II, and there is no paly-
rero on the Pacific coast of Mexico to Panama, where
nological evidence of forest disturbances that would
signal slash-and-bum agriculture during this period
forest clearing for agriculture may have occurred as
in the Mesoamerican tropics. Rather, the archaeo-early as 7000-5000 B.P. (Pipero and Pearsall
1998:292-293). However, north of Guerrero inten-
logical evidence from Veracruz, Belize, and lowland
Chiapas (Voorhies 1976; Wilkerson 1975) indicatessive archaeological site surveys in West Mexico in
that lowland peoples were developing a maritime the states of Jalisco, Nayarit, and Sinaloa have pro-
adaptation and were not engaged in forest-clearing
duced no lithic artifacts such as grinding stones used
activities as in Panama. in food preparation that would indicate incipient cul-
Stage III: 7000-6500 B.P. A premise of the Rio tivation activities during the Archaic (Benz 1999;
Balsas model is that Zea had not reached the high-Mountjoy 1998). To the south, maize does not appear
lands of Tehuacain, Oaxaca, and the Valley of Mex-in the pollen profiles from Laguna de Chantuto on
ico during Stage III (Figure Ic), but highlandthe Pacific coast of Chiapas, Mexico (Voorhies
microfossil evidence for Zea and maize cobs dated 1976). It is possible that maize was present by this
stage in Chiapas where Zea pollen was reported from
to this stage have been reported from the Oaxaca Val-
Santa Marta Cave by 6360 + 160 B.P. (uncalibrated)
ley (Flannery 1986:8; Schoenwetter and Smith 1986);
Zea and amaranth have been found in the open-air or about 6500 B.P. (Garcfa-Barcena 1976), but it was
not found at Santa Marta in an earlier context dated
site of Zohapilco (Tlapacoya) in the Valley of Mex-
ico (Niederberger 1979), and numerous macrobot- to 7310 ? 300 (uncalibrated) B.P. (MacNeish and
anical maize remains were excavated in dry caves Peterson
in 1962). The later Chantuto materials, and

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MacNeish and Eubanks] MODELS FOR THE ORIGINS OF MAIZE 7

yu - r

Boundaries Based On Current Evidence Figure If. In Stage VI of the Rio Balsas model of origin
and domestication of maize, different races of maize
spread north into Thmaulipas and Zacatecas en route to
- - - - Hypothesized Biogeographical Boundaries
U.S. Southwest and extended south down the Andes.
Widespread subsistence use in Mesoamerica and Cen
L\\\X Boundaries Inferred From Microfossil Data
America. Evidence consists of well-dated cobs from
Tehuacan and Zea pollen from many sites from
Figure le. In Stage V of the Rio Balsas model of origin
Mesoamerica to northern South America.
and domestication of maize, rapid spread of maize into
the Mesoamerican highlands. Occurs in the Yucatan,
Belize, and perhaps rest of tropical lowlands. Evidence
The Tehuacan (or Highland) Model
consists of well-dated cobs from Tehuacan and Zea pollen
from Zacatecas, the Yucatan, Belize, and northern South
Large quantities of artifacts and ecofacts excavat
America.
from well-defined stratigraphic sequences dated b
possibly the materials from the tropical lowlandsradiocarbon
of determinations provide direct eviden
in support of the Tehuacan model. The paleobotan
Belize (MacNeish 1983), indicate a specialized adap-
ical remains for the six stages of the Tehuacan mod
tation to exploitation of the aquatic resources found
are summarized in Table 1 and described more fully
in marshes and river estuaries or along the seacoast,
but do not suggest an early manipulation of plants below.

in the fertile agricultural areas. Presently, there is lit-Stage I: Before 7500 B.P In the first stage (Fig
ure 2a), the earliest plants that people were manip
tle archaeological evidence for a connection between
ulating and domesticating in the Mesoamerica
Mesoamerica and Panama. No dated archaeological
highlands included pumpkins (Cucurbita pepo) an
and/or microfossil evidence are yet available to sup-
bottle gourd (Lagenaria siceraria). Gourd rinds we
port the diffusion purported to have occurred during
Stage IV, according to the Rio Balsas model. recovered at Guila Naquitz (Flannery 1986) in Zon
C (11,330-9865 cal B.P.) and in Zone B2
Stage V. 6000-5000 B.P In Stage V (Figure le),
maize was present in Mexico from the state(9945-8375
of cal B.P.) (Whitaker and Cutler 1986
Hidalgo to highland Chiapas, extended into Fartherthe north in Tamaulipas, gourd rinds occurred i
Zone G (8560-7960 cal B.P.) of Valenzuela Cave
Yucatan Peninsula (Pohl et al. 1990; Pohl etal. 1996),
Guatemala, and Honduras (MacNeish 1992), and (Whitaker et al. 1957), and remains of Cucurbita
pepo were even more common at this early tim
had apparently spread onto the Pacific coasts of both
Americas, down the Andes, and into Venezuela period (Smith 1997a, 1997b). At Guila Naquitz, a C
pepo rind occurred in Zone C (11,330-9865 cal B.P.
(Pearsall 1989). Even though maize was widespread,
its use in much of Central America, excepttwo
in seeds came from Zone B3 (10,250-8500 ca
Panama, appears to have been limited. B.P.), one seed from Zone B1 (8560-7930 cal B.P.)
and a seed and three peduncles from Zone B
Stage VI: 5500-4500 B.P By the final stage (Fig-
(8185-7655 cal B.P.) (Smith 1997b). One seed of C
ure If), according to the Rio Balsas model, maize
had spread throughout Central America and into pepo, originally classified as wild, occurred in Zon
South America. In the upper preceramic levelsXIV at (7800-7500 cal B.P.) of Coxcatlan Cave in
Puerto Marques, a site in a suburb of Acapulco, Tehuacan
are (Cutler and Whitaker 1967). In Tamaul
pas, two seeds of C. pepo recovered from Zone H
the remains of house floors with grinding stones sug-
gesting village agriculture (Brush 1965). (8800-8200 cal B.P.) and one from Zone G

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 Table 1. Paleobotanical Evidence for the Tehuacan Model.

Sites and Exavation Units by Region

Oaxaca Tehuacan Mexico Tamaulipas Archaeological Plant Remai
GN CB GS Cox SM Zop Val Ro LaP Years B.P. Lag Cp Trip Zmf Cap Av Ca Maize Cm
B 4700-4300 pg X
L3 4865-4465 14 16 pg 2
B 5200-4800 X X 15

Zop 5250-4850 pg
L8 5300-4900 2 15
L9 5450-4950 1 3pg X
LIO 5510-5010 1 2 X X
VIII 5940-4910 2 4s 4 4s 15
D 6160-5000 102
IX 6040-5080 2 3s 2 27 1
L4 6215-5743 2r
L5 6250-5750 3s
X 6300-5400 2r 2
P1 6800-6400 1 pg
E-F 6980-6100 33
P2 6810-6400 pg
XI 7750-6550 3s 32 25 Is
XII 7700-6500 2s 1 5 Is
XIII 7870-7040 Ir Is Is 18
C 8000-7000 2pg
C 7900-7100 1 2pg
XIV 7800-7500 Is 3s 2 2s
XV 7900-7700 3s 5
XVI 8830-7240 Is 3
B1 8560-7930 Is 4pg
G 8560-7960 2r Is
H 8800-8200 2s
B2 9945-8375 lr ls 9pg 6pg
B3 10250-8500 2s 13pg lpg
C 11330-9865 lr Is 2ng
Abbreviations Key: GN = Guild Naquitz; CB = Cueva Blanca; GS = Gheo Shih; Cox = Coxcatlan Cave; SM = San Marcos Cave; Zop = Zohapilco;
Romero's Cave; LaP = La Perra Cave; Lag = bottle gourd (Lagenaria siceraria); Cp = pumpkin (Cucurbita pepo);Trip = Tripsacum pollen; Zmf = Ze
(Capsium annuum); Av = avocado (Persea americana); Ca = squash (C. argyrosperma);Cm = C. moschata; Am = Amaranthus sp.; BS = black sapote
sapote (Casimiroa edulis); Pv = common bean (Phaseolus vulgaris); Chup = chupandilla (Cyrtocarpa prosera); Can = jack bean (Canavalia sp.); Tep =
pollen grain; X = number not reported; s = seed; r = rind.

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MacNeish and Eubanks] MODELS FOR THE ORIGINS OF MAIZE 9

Boundaries Based On Current Evidence

Figure 2b. In Stage II of the Tehuacan model of origin and
domestication of maize, early maize spread in highlands
from Valley of Mexico to Chiapas. Evidence consists of
- - -- -Hypothesized Biogeographical Boundaries
maize remains from Tehuacan and pollen from Oaxaca,
the Valley of Mexico, and Chiapas.
[\\\N Boundaries Inferred From Microfossil Data
Rockshelter, Zone C (some 11,000 years old) had
Figure 2a. In Stage I of the Tehuacan model of origin and
domestication of maize, natural hybrids between two Zea pollen grains, and at Gheo-Shih there were
two Zea grains at about 7000-6000 B.P. Predating
Tripsacum and perennial teosinte gave rise to novel phe-
notypes that evolved into maize. Evidence consists of and accompanying Zea pollen at Guila Naquitz was
experimentally reconstructed prototypes of early maize
produced by crossing Tripsacum and perennial teosinte. Tripsacum pollen-two grains in Zone C (some
Supporting evidence is Zea and Tripsacum pollen from 11,000 years old) and 26 grains in Zone B.
Guila Naquitz and Martinez Cave associated with 14C Stage II: 7500-7000 B.P. More tangible evidence
dates from 7630 to 10,500 B.P. and DNA fingerprinting
that shows maize has many of the same genes as for plant domestication exists in the Mesoamerican
Tripsacum. highlands in Stage II (Figure 2b). In Oaxaca, two Zea
pollen grains were present from Zone C at Geo-Shih,
(8560-7960 cal B.P.) of Valenzuela Cave are large and two Tripsacum pollen grains and a specimen of
enough to be considered domesticated (Cutler and Cucurbita pepo have been reported from Zone C of
Whitaker 1967; Smith 1997b). Cueva Blanca (7900-7100 cal B.P.) (Schoenwetter
Other evidence for plant manipulations in the and Smith 1986). The most abundant evidence for
Mesoamerican highlands at this time includes chile this stage, however, came from Coxcatlan Cave in
(Capsicum annuum) seeds found in Zone XVI the Tehuacan Valley (Mangelsdorfet al. 1967a; Smith
(8830-7240 cal B.P.), Zone XV (7900-7700 cal 1967). There, Zone XIII (7960-7860 cal B.P.) had
B.P.), and Zone XIV (7800-7500 cal B.P.) of Cox- Lagenaria siceraria rinds and seeds, Capsicum
catlan Cave; avocado (Persea americana) pits, and annuum seeds, Cucurbita argyrosperma, 17 cobs of
seeds of Cucurbita argyrosperma (originally named "wild" maize, and one of cultivated maize. Although
C. mixta) from Zones XIV and XV of Coxcatlan when first discovered this early form was referred to
Cave (MacNeish 1967). It should be noted, how- as "wild" maize, it was already distinct from teosinte
ever, that the avocado remains at Tehuacan may be and Tripsacum in its paired kernels, no longer pro-
intrusive from some tropical region where other plant tected from predators by being encased in hard fruit-
manipulations were taking place (Smith 1967). cases, and its inability to disperse its seed naturally.
Although there are no macroremains of maize Therefore, these specimens probably depict maize
from Stage I, and although Henry Irwin never com- in the early stages of cultivation by humans rather
pleted his study of the pollen profiles from Tehuacain than a truly wild form. Zone XII (7700-6500 cal B.P.)
(which were rumored to have excellent preservation had C. annuum, Persea americana, and five "wild"
with maize pollen throughout), the palynological corncobs, as well as possibly Cucurbita moschata
lacuna is filled by the pollen profile from Oaxaca and amaranth (Amaranthus leucocarpus). Zone XI
(Schoenwetter 1974; Schoenwetter and Smith 1986). (7650-6550 cal B.P.) of Coxcatlan Cave had 22
Guila Naquitz had eight grains of Zea pollen from "wild" and three cultivated corncobs, one rind of C.
Zone B (8185-7655 cal B.P.). At nearby Martinez moschata, two pits of black sapote (Diospyros dig-

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 10 LATIN AMERICAN ANTIQUITY [Vol. 11, No. 1, 2000

s65-6000 B.P.I

t C '.rbb. ,

W Carf bibean

IStage IVI
V

Boundaries Based On Current Evidence Figure 2d. In Stage IV of the Tehuacan model of origin
and domestication of maize in Mesoamerica, early race
of maize (Nal Tel and Chapalote) reached from Queretar
- - - - Hypothesized Biogeographical Boundaries
to Panama and extended into lowlands in the Olmec
region of Gulf of Mexico coast. Evidence consists of w
N\\,] Boundaries Inferred From Microfossil Data
dated early tripsacoid and Nal Tel/Chapalote cobs
Tehuacan and Zea pollen in Oaxaca, near La Venta i
Tabasco, Veracruz, and in Panama sites.
Figure 2c. In Stage III of the Tehuacan model of origin
and domestication of maize, early cultivated maize spread
in highlands, extending from Hidalgo to Central America
(6300-5400 cal B.P.) of Coxcatlan Cave in Tehua
and into lowlands of the Olmec region of the Gulf of
Mexico coast. Evidence consists of cobs from Coxcatlan, (Mangelsdorf et al. 1967a; Smith 1967). Th
Purr6n, and San Marcos caves in Tehuacin; dated cobs include two Lagenaria rinds, one cultivated and
and pollen from Oaxaca; dated pollen in the Valley of
Mexico, Chiapas; and Coxcatlan-like remains in Hidalgo
early tripsacoid maize cob, amaranth seeds, fi
and Tabasco. black sapote pits, and a cultivated chupandilla
Maize, teosinte, and amaranth occur in the Valley
Mexico
yna), one pit of white sapote (Casimiroa edulis), and (Niederberger 1979). The presence of ma
and metates suggests that maize agriculture h
a pit of cultivated chupandilla (Cyrtocarpaprocera).
reached
Again, it should be noted that black sapote, like avo- north to Hidalgo and south to highland H
duras
cado, is a tropical plant. As yet no evidence for Zea (Bullen and Plowden 1963).
or Tripsacum has been found in the pollen profilesStage V. 6000-5500 B.P Although Stage V (F
from Chiapas and the Valley of Mexico. ure 2e) also is poorly represented, there is evid
Stage III: 7000-6500 B.P. Stage III (Figure 2c)the variety of plants had increased. Relev
that
specimens appear mainly in Zone IX (6040-5808
is less well represented in the highlands by domes-
ticated plant remains at Tehuacian and Oaxaca. B.P.)
Zonesof Coxcatlan Cave (MacNeish 1967), and t
E and F (6980-6100 cal B.P.) from San MarcosOcampo
Cave levels in Romero, Valenzuela, and Ojo
Agua caves in Tamaulipas (Mangelsdorf et
had 27 "wild" maize cobs (Mangelsdorf et al. 1967a).
1967b). Zone IX had remains of gourd, avocad
In Zone B 1 at Guila Naquitz, dated to 9000-7500
cal B.P. (Flannery and MacNeish 1997:95), chile
therepepper, Cucurbita moschata, amaranth, b
were five primitive-looking maize specimens and white sapote, pits of cultivated chupandilla,
(Flan-
a mixture
nery 1986:8, Figure 1.2). Maize and/or teosinte and of 7 wild, 18 early cultivated, and 2
Tel-Chapalote
amaranth may be present in Playa 2 in the Valley of maize cob types. Tamaulipas
Mexico (Niederberger 1979), and Santa Martamainly
Cave Lagenaria and C. pepo remains (MacNe
in Chiapas may have maize pollen (MacNeish1958).
and The evidence is scarce, but it is possible
Peterson 1962). By this stage, maize had spread agricultural
out remains existed all over the highl
of the highlands to the Pacific coast (Voorhies and lowlands, and extended as far south as Pan
1976),
and into South America.
and down the highlands as far south as Panama
(Pipero 1989). Stage VI: 5500-4500 B.P. In final Stage VI (Fig-
Stage IV: 6500-6000 B.P. Stage IV (Figure
ure2d)
2f), the Archaic ended in Mesoamerica and incip-
ient
is the most poorly represented in the highlands. agriculture appeared everywhere. The word
Rel-
evant plant remains come only from Zone X is employed because the few isotopic (12-
incipient

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MacNeish and Eubanks] MODELS FOR THE ORIGINS OF MAIZE 11

m.

3r961L

Boundaries Based On Current Evidence Figure 2f. In Stage IV of the Tehuacan model of origin
and domestication of maize, maize races spread north
from Tamaulipas and Zacatecas and traveled south down
- - - - Hypothesized Biogeographical Boundaries
the Andes. Maize used on subsistence level in
Mesoamerica and Central America. Evidence consists of
[\\\,1 Boundaries Inferred From Microfossil well-dated
Data cobs from Tehuacan and Zea pollen from
many sites in Mesoamerica, Central America, the north-
Figure 2e. In Stage V of the Tehuacan model of origin
ern and
Andes, and the north Pacific coast of South America.
domestication of maize, early races of maize reached
Tamaulipas, the Maya lowlands, and Colombia in South
America. Evidence consists of well-dated cobs from
related to this annual teosinte. Subsequent to these
Tehuacan and Zea pollen from Zacatecas, Yucatan,
Belize, and northern South America. studies, advances in molecular techniques have
shown there are limitations to isozyme data for
13C) tests on skeletons (Farnsworth et al. 1985)addressing evolutionary questions because of cryp-
reflect diets that were not dependent on maize tic
or allelic heterogeneity (Murphy et al. 1996). DNA
other domesticated plants. Jack beans (Canavalia sequencing and fingerprinting methods that are more
widely employed now reveal recessive as well as
ensiformis) and tepary beans (Phaseolus acutifolius)
were added to the larder in Zone VIm (5940-4910 dominant alleles, and are thus better for determin-
cal B.P.) of Coxcatlan Cave (Smith 1967). Duringing phylogenetic relationships.
this period, recognizable maize races adapted to dif- The second line of molecular evidence seen as
ferent environments appeared. One of these races,
supporting the Rio Balsas hypothesis was chloroplast
DNA (cpDNA) analysis (Doebley, Renfroe, and
Chapalote, was spreading northward into the U.S.
Southwest, where it evolved into a distinct form well
Blanton 1987), which partially complemented the
adapted to the arid conditions there (Galinat and
isozyme data. The cpDNA data, however, did not
Gunnerson 1963; Upham et al. 1987). divide highland Chalco teosinte (Z m. spp. mexi-
cana) and lowland Balsas teosinte (Z. m. spp. parvig-
Genetic Evidence
lumis) into separate subspecies. Thus the cpDNA
evidence indicated that if an annual teosinte is the
The Teosinte Hypothesis
progenitor of maize, highland teosinte from the Val-
ley of Mexico, which Galinat (1977) postulated to
In the 1980s, studies of 13 isozyme systems in maize
and teosinte (Doebley et al. 1984, 1985; Doebley,
be its ancestor, is equally as likely to be maize's pre-
Goodman, and Stuber 1987; Goodman and Stuber cursor as Balsas teosinte. Doebley (1990) attributed
1983; Mastenbroek et al. 1981; Smith et al. 1984,
this inability of the cpDNA data to resolve the two
1985; Stuber and Goodman 1983), indicated that annual teosintes into subspecies as due to introgres-
Balsas teosinte (Z. m. ssp. parviglumis) is most
sion via hybridization or to lineage sorting (reten-
closely related to maize. No comparable data tion
for of alleles inherited from a common ancestor).
Tripsacum, the other close relative of maize, Chloroplast
were DNA analysis of Tripsacum confirmed
that Zea and Tripsacum are closely related. Since
reported. Doebley's (1990) idea that the parviglumis
teosinte growing in the Rio Balsas drainage iscpDNA
the which is highly conserved in evolution does
ancestor of maize was partially based on these
not detect introgression and hybridization (Dowling
isozyme data that showed maize is most closely
et al. 1996; Rieseberg and Soltis 1991), DNA fin-

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12 LATIN AMERICAN ANTIQUITY [Vol. 11, No. 1, 2000

taneously
gerprinting and sequence data are necessary for accumulating mutations in 5 different
resolving reticulate phylogenetic histories between
genes related to the plant's ability to reproduce and
closely related species (McDade 1992; Rieseberg
surviveet
in nature seems unlikely. This may explain
al. 1991; Soltis et al. 1991). Thus, the cpDNA
whystud-
no one has experimentally replicated the suite
ies may not be sufficient to elucidate the origin of
of mutations required to convert teosinte into maize
maize question, and other lines of evidence are
via recurrent selection or mutagenesis. Nor is it sur-
important for investigating biocultural evolution
prising that the mutation search, led by George Bea-
related to the origin and evolution of maize.dle in 1970, failed to recover any mutant ears among
natural
Teosinte as the progenitor of maize is the mostteosinte populations in Mexico (Mangels-
parsimonious hypothesis, and therefore the one
dorfmost
1974), or any forms with reduced, soft glumes
or other
favored by reductionist science. It avoids the issue macromutations converting teosinte into
of extinct ancestral taxa, one of Beadle's maize
(1939)among the numerous maize-teosinte hybrids
observed
strongest objections to the hybridization thesis (Man-during the field expedition.
gelsdorf and Reeves 1939). However, this rationale
An alternative explanation of how the teosinte ear
may be flawed because, although the simplest
wasexpla-
transformed into the maize ear is the catastrophic
nation is useful heuristically in laboratorysexual transmutation theory (CSTT) (Iltis 1983)
science
where variables can be carefully controlled,which proposes that a sex change transformed the
it may
male flowers
not accurately reflect natural phenomena subject to of the apical tassel to female flowers
multiple biotic and abiotic variables that in change
the leaf axils. This putative transformation is sup-
through time and space. The teosinte hypothesis
posed to have been induced by a hormonal switch
assumes that selection for mutations in five major
turned on by pathogens or environmental change. If
genes of teosinte gave rise to maize. true, it should be possible to replicate such a trans-
formation experimentally, but no corroborative
In general, mutations are rare; most newly formed
mutations are deleterious and do not surviveexperimental
(Lewin data have been reported. On the other
1997). Stadler's (1951) data on mutation frequencies
hand, Tripsacum, which bears male and female flow-
ers on the
for 7 genes provide a baseline for the frequency of same spike, has a genetic mechanism that
mutations in maize. Spontaneous mutation rates the switch from male to female. In early
regulates
ranged from 492 mutants out of a million development
gametes in teosinte and Tripsacum there are two
in the red color (R) gene; 106 out of a millionfemale
for the
flowers, one of which is normally aborted and
only
inhibitor (I) of R; 11 out of a million for the a single grain is produced. However, the genetic
purple
aleurone (Pr) gene; 2.4 out of a million for switch
starchythat regulates the transition from male to
(Su) endosperm; 2.2 out of a million for the yellow
female in Tripsacum can prevent abortion of the sec-
(Y) color gene; 1.2 out of a million for normal ker- flower and two kernels will develop in a
ond female
single
nels (Sh), and 0 out of a million for the waxy cupule (Dewald et al. 1987; Li et al. 1997).
(Wx)
gene). Since spontaneous mutations are soSince
infre-
Tripsacum has a gene for the distinctive trait
quent, geneticists and plant breeders routinely use kernels in each cupule of maize, it is plau-
of paired
sible that
mutagenic agents such as chemicals and radiation to this key trait of domesticated maize was
inherited from Tripsacum.
increase the frequency over spontaneous mutation
rates in order to recover variant forms that can be
The Hybrid Origin Hypothesis
used in genetic analysis and selection of new vari-
eties. Another method of inducing mutagenesis inA hypothesis of how, when, and where maize first
maize is transposon tagging in which a maize line isappeared and subsequently evolved into its domes-
crossed with one containing transposable elementsticated form should be consistent with both the
(i.e., jumping genes). When a transposable elementarchaeological record and experimental evidence.
is inserted into a gene, it causes a mutation. TheAlthough excavated over 30 years ago, the most com-
mutation frequency for different mutator lines variesplete prehistoric record of maize evolution is still the
from 1 in a thousand to 1 in a million (Chomet 1994).24,186 maize specimens excavated in dry caves in
To find a single mutation using a highly mutagenicthe Valley of Tehuacan (Mangelsdorf et al. 1967a).
transposable element line, a breeder needs to screenThese macrobotanical remains provide a well-
100,000 progeny. Therefore, the probability of simul-defined evolutionary sequence covering a period of

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MacNeish and Eubanks] MODELS FOR THE ORIGINS OF MAIZE 13

pollinations
6,500 years, and among them are the oldest pre- between teosinte and Tripsacum
(Tantravahi
served cobs yet found. Notable features of the earli- 1968), no viable hybrid plants were
est cobs include: remarkable uniformity obtained
in size, before the advent of diploid perennial
ranging from 1.9 to 3 cm long; 4 to 8 rows teosinte
of ker- (Zea diploperennis), discovered on the
threshold
nels; 2 kernels per cupule; long, papery glumes par- of extinction in the mountains of Jalisco,
Mexico,
tially covering the kernels; the cob axis formed byin the late 1970s (Iltis et al. 1979). While
conducting
individual cup-like units fused together at the ends cytological and crossing studies of the
newmale
and sides; and bisexual inflorescences with the diploid perennial teosinte, Eubanks (1987)
flowers above the kernel-bearing cob. The observed
missingstriking similarities between the chromo-
some architecture
piece of the evolutionary puzzle is how the maize ear ofZ. diploperennis and Tripsacum.
Both
metamorphosed into its unique multiple-rowed taxa have diminutive terminal knobs (a term
struc-
coined by McClintock [1929] to describe darkly
ture. This question has inspired various explanations
staining heterochromatic areas that resemble black
of how this botanical anomaly arose, and archaeol-
ogists have focused on these different viewsbeads on a string); whereas the other Zeas (maize
to guide
their interpretations ofbiocultural evolution.and
Didannual
this teosintes) have large internal knobs. Fur-
thermore,
change result from descent from a single ancestor as although the two genera have different
in the Rio Balsas model, or did it arise through
base chromosome numbers (in Tripsacum x= 18 and
in Zea x= 10), the total length of Tripsacum dacty-
hybridization between two or more wild progenitors
as in the Tehuacan model? loides chromosomes, 492.5 pm (Chandravadana et
Discovery of the diploid perennial teosinte led toal. 1971), closely approximates the total length of
Zea diploperennis chromosomes, 502.7 pm
a breakthrough in experimental plant breeding that
reproduced forms closely resembling the early (Pasupuleti and Galinat 1982). In contrast, the total
archaeological maize from Tehuacan. These results,
length of maize chromosomes is 552.7 lpm (Rhoades
1950), and the total length of annual teosinte chro-
along with new evidence from molecular biology, are
mosomes is 552.0 pm (Longley 1941). Thus the
compatible with the Tehuacan model of a hybrid ori-
annual species have genomes that are approximately
gin of maize in most respects. A significant finding
that is compatible with the Rio Balsas model, how-
ten percent greater in size than their perennial taxa.
ever, is that a teosinte species, rather than extinct wild Their similarities in chromosome architecture led
maize, is an ancestor of maize. Another important
Eubanks to believe it would be possible to cross Trip-
sacum with the diploid perennial teosinte. Her cross-
revelation to guide reinterpretation and future archae-
ing experiments have yielded fully fertile, hybrid
ological exploration is that "wild" maize was prob-
plants (1995, 1997a), some of which produced ears
ably a perennial plant adapted to high elevations and
bearing key morphological features in common with
capable of surviving extended periods of drought. It
the oldest maize cobs from Tehuacain (Mangelsdorf
reproduced asexually from rootstocks and stem cut-
tings, as well as sexually from seed; thus it could sur-et al. 1967a). Like the Tehuacan specimens, these
vive and reproduce despite overharvesting by
hybrid forms have fused rachis segments with paired
kernels partially exposed by reduced outer glumes,
humans or years of extreme fluctuations in environ-
and many are bisexual, with the male flowers above
mental conditions that might reduce or prevent seed
production. It is therefore possible that the earliestthe female flowers on the same spike, features
derived from their Tripsacum parent (Eubanks 1995,
cultivation and transport of maize did not fit the stan-
dard paradigm of planting and harvesting seed, but1997a; Galinat 1999). Thus, the critical steps in the
may have involved digging roots or transplanting transformation from the single-rowed spike of the
wild relatives to the multi-rowed maize ear have been
cuttings, a technique that might have diffused into
experimentally reconstructed in segregating prog-
the highlands from manioc cultivators in the tropi-
cal lowlands. eny of Tripsacum-Z.diploperennis hybrid plants.
These recreated prototypes of early maize provide
Teosinte was believed by some scientists to be the
ancestor of maize because crosses between maize the missing link in its evolutionary history. Further-
and teosinte produce viable offspring and pollina-
more, comparison of molecular profiles of Z. diplo-
tions between Tripsacum and maize generally perennis-Tripsacum
pro- hybrids, their parents, primitive
duce sterile progeny. Despite literally thousands of races, and annual teosintes show that many of
maize

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14 LATIN AMERICAN ANTIQUITY [Vol. 11, No. 1, 2000

teosintes. Therefore, it is possible wherever Zea

the same Tripsacum alleles observed in the hybrids
pollen
also are found in the ancient indigenous maize raceshas been found it could actually be pollen of
a Tripsacum-teosinte
Chapalote, Nal Tel, and Pollo, but are not found in hybrid (Eubanks 1997b).
the annual teosintes from Chalco, Rio Balsas, and
Microfossils are important paleoecological indica-
Guatemala (Eubanks 1999b) as would be expected tors but other corroborative evidence is needed to
if annual teosinte is the ancestor of maize. augment inferences about early agriculture that
DNA evidence from internal transcribed spacer depend on presence of maize and/or its wild relatives.
(ITS) sequence comparisons indicates maize Since Eubanks (1995, 1997a) has demonstrated that
appeared either just before, or contemporaneouslyhybrids between perennial teosinte and Tripsacum
with, the Balsas and Chalco races of teosinte (Buck- resemble reconstructed prototypes of primitive
ler and Holtsford 1996), and the authors speculatemaize, and the pollen of these hybrids is indistin-
that the domestication of maize was a terminal Pleis- guishable from maize and teosinte (1997b), her sug-
tocene/early Holocene event, perhaps without humangestion that "wild" maize was a natural hybrid
manipulations. If correct, it is highly improbable that between teosinte and Tripsacum is compatible with
Balsas or Chalco teosintes are the ancestor of maize, the palynological data.
because the time frame seems too condensed for Another limitation of the Rio Balsas model may
be the rationale that the early picture in Mesoamer-
accumulation of the required mutations. In contrast,
DNA studies indicate maize, along with the Balsas
ica parallels developments in Panama. The well-doc-
and Chalco races of teosinte, derived from natural
umented sequence in Panama is based upon abundant
hybridization between two of the more primitivestratified cave materials (Ranere 1980) that contain
taxa. Other molecular data lend support to the
strong hints of early plant manipulations, as evi-
denced by grinding stones and metates, and are asso-
hybridization model. Talbert et al. (1990) showed that
ciated with pollen profiles that indicate clearing of
Tripsacum andersonii, which grows in Central and
the forest from 5,000 to 10,000 years B.P. At pre-
northern South America, is a natural hybrid between
maize and Tripsacum, a finding that demonstrates
sent, there is no comparable evidence from lowland
introgression between Tripsacum and Zea occursMesoamerica
in for a similar type of forest clearing dur-
nature. There also is molecular evidence indicating
ing that early period, or of plant manipulations that
might lead to maize domestication and the develop-
perennial teosinte genes may have introgressed into
South American maize in prehistory. It was reported
ment of agricultural practices. In contrast, the empir-
ical data for the Mesoamerican lowlands-coastal
that archaeological maize specimens from Peru had
an allele of the adh2 gene that is identical to one Guerrero
in (Puerto Marques) (Brush 1965), Pacific
perennial teosinte (Goloubinoff et al. 1993). coast Chiapas (Laguna de Chantuto) (Voorhie
1976), Belize (the Lowe-ha, Sand Hill, Orange Walk
Discussion
and Belize phases) (MacNeish 1983), and the early
Presently direct evidence from archaeology and coastalArchaic
pale- in Veracruz (Wilkerson 1975)-indi-
oecology that supports maize agriculture duringcate
the exploitation of maritime and aquatic resource
first two stages (from before 7500 B.P. to 7000 and
B.P.)little concern with plant manipulations. In all o
according to the Rio Balsas model has not been
the phases prior to 6000 B.P., no manos and metate
used in maize preparation have been found, and only
recovered. In the later stages, the evidence is repre-
a few scraper planes or pulpers for working plan
sented mainly by Zea pollen that is not accompanied
material have been excavated. There also is no sup
by solid dates, and that occurs without much asso-
ciated contextual archaeological remains. Pollenport
and for the assumption that domesticated maize an
phytoliths are collectively referred to as Zea micro-
other domesticated plants diffused through the trop
ical lowlands from Mesoamerica to Panama. The
fossils because it is difficult and generally not pos-
only evidence for such a diffusion into the area
sible to distinguish maize and teosinte pollen grains
between Mesoamerica and Panama are grinding
and phytoliths to the species level of identification
(Dunn 1983; Eubanks 1997b; Lippi et al. 1984; stones and scraper planes found in Chiapas at Santa
Piperno and Pearsall 1993; Roosevelt 1984; Rovner Marta (MacNeish and Peterson 1962), and the sur-
1999). Pollen grains of Tripsacum-teosinte hybrids face collection of Archaic materials at La Esperanza
in western Honduras (Bullen and Plowden 1963).
also are difficult to distinguish from maize and annual

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MacNeish and Eubanks] MODELS FOR THE ORIGINS OF MAIZE 15

Another problem with the Rio Balsas model may

be its assumption that early maize, Tripsacum, and
teosinte did not exist in the Mesoamerican highlands
before 3500 B.C. (Fritz 1994; Long et al. 1989).
Flannery and MacNeish (1997) argue that the
Tehuacan specimens used for theAMS dates of Long
et al. (1989) were contaminated with bedacryl, a
polymethylacrylate frequently used by conservators
to preserve fragile botanical materials. The speci-
mens were pretreated with hydrochloric acid before
I
t.

dating, but not with organic solvents that are required i

fI

for bedacryl removal (Long et al. 1989:1036); it is II

possible that the more recent dates may have been

skewed by the preservative. Another possible expla-
nation for the incongruity between the AMS dates
and the 14C dates could be accidental mixing of spec-
imens during the many times they were moved and
handled since originally excavated in the early 1960s.
The highland origin of maize model as presented here
thus assumes that MacNeish's earlier 14C dates are
valid, and that maize was present at Tehuacain around
7600 B.P. The debate about the dates of the earliest
maize remains is important, but not necessarily crit-
ical to the discussion, because Tripsacum and Zea
pollen have been identified from Oaxaca that date in
the 10,000 B.P. range, and maize cobs from Oaxaca
date to Zone B 1 at Guila Naquitz, around 6500 B.P.
The Tehuacain model seems more plausible than
the Rio Balsas model because it has better support
from existing archaeological and biological evi-
dence. One reason a highland origin of maize is pro-
jected is that diploid perennial teosinte is adapted to
a highland rather than a lowland environment (Iltis
et al. 1979), and Tripsacum grows sympatrically to
perennial teosinte in the same habitat (Vaizquez G.
Figure
et al. 1995). Specimens resembling the oldest 3. An ear resembling archaeological specime
maize
early cultivated maize from the Valley of Tehuac
ears at Tehuacain (Mangelsdorf et al. 1967a) can be
Mexico. Derived experimentally from intercrosse
recovered from segregating progeny from crosses diploperennis hybrid progeny with
Tripsacum-Z.
Tripsacum
between perennial teosinte and Tripsacum. When as the maternal parent throughout the descent
line. Length from base of ear to tip of staminate spike is
hybrid plants cross-pollinate, if the pollen11recipient
cm.
has a Tripsacum cytoplasm (i.e., Tripsacum is the
maternal parent of the hybrid), segregating maize and its evolutionary sequence as observed in
intercross
Tehuacainand
progeny produce ears with soft, reduced glumes archaeobotanical record.
Based
exposed kernels that resemble early cultivated on the experimental findings reported here,
maize
it is proposed
from Tehuacain (Figure 3, referred to as "early cul- that wherever populations of diploid
perennial teosinte and Tripsacum were growing in
tivated maize" in Figure 4). Segregating backcrosses
produce phenotypes similar to tripsacoid close
maizeproximity,
and because they are wind-pollinated,
wild-type cobs described and illustratednatural
among hybrids could have occurred. When hybrids
maize specimens at Tehuacan. Thus, these experi- with either parent, the resulting prog-
backcrossed
mental crosses appear to reconstruct theeny
origin
reverted
of to the parental phenotype. However,

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16 LATIN AMERICAN ANTIQUITY [Vol. 11, No. 1, 2000

1 *~ ca L~ ~-

t I
X
zEm - ?

Figure 4. Putative evolutionary trajectory of

cross pollinating Tripsacum dactyloides and
sequent segregating generations that trace
vated" maize. Approximate lengths of spec
dactyloides, 10 cm; primitive maize prototyp

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MacNeish and Eubanks] MODELS FOR THE ORIGINS OF MAIZE 17

when hybrids crossed with other hybrids,replacement

some of for the Tehuacain model, as suggested
by advocates
the recombinant offspring produced ears with four of the Rio Balsas model, we propose
to eight rows of paired kernels in reducedthat domestication of lowland tropical plants
cupules
that exposed the grain, making it easy to occurred
remove as a parallel, dichotomous development
within
from the cob (Figure 4). This type of ears would a greater Mesoamerican interaction sphere
have
provided an attractive food source for humans. As the highlands and tropics. We argue that
connecting
humans dispersed early maize into different the Tehuacan highland model, which incorporates
habitats,
evidence
in addition to artificial selection for culturally pre- from the oldest, most complete archaeo-
ferred types, natural selection for new ecological
logical record of maize evolution and domestication,
and is now supported by new information from
races occurred, and gene pool diversity rapidly
expanded through introgression with other experimental
popula- plant breeding and molecular genetics,
tions of parental taxa and/or other hybrids. is the more
Within a probable model for the origin of maize.
Through comparative genomics of teosinte and
few millennia, maize was transformed by artificial
selection and adaptation to new habitats Tripsacum
into the taxa, segregating teosinte X Tripsacum
genetically diverse and highly productive crop plantand primitive maize races, it should be pos-
hybrids,
that was the cornerstone of New World civilizations.
sible to more accurately trace the ancestry of the
There is agreement between the Rio Balsas
maizeand
family and test the hypothesis that annual
teosinte
Tehuacan models in Stage V (6000-5000 B.P.) when species, and possibly some Tripsacum
maize was found in Mexico from the state of Hidalgo
species, arose via hybridization events between their
perennial ancestors. Molecular analysis of Zea and
to highland Chiapas, and extended into Guatemala,
Honduras (MacNeish 1992), and theYucatanTripsacum
Penin- taxa will not only aid in tracing the evo-
sula (Pohl et al. 1990, 1996). The Tehuacainlutionary
model, history of the maize and teosinte racial
complexes,
however, emphasizes that in much of Central Amer- but the findings can be integrated with
ica, except in Panama, maize use was limited. Maizefrom the archaeological record to elucidate
evidence
also apparently spread onto the Pacific coaststhe
of interrelationship
both of the origin and evolution of
Americas, down the Andes, and into Venezuela
maize with the transition from hunting-and-gather-
(Pearsall 1989) at this time. ing subsistence strategies to agricultural economies.
Anthere
In the final stage (5500-4500 B.P.), although important goal for new archaeological-botanical
is agreement between the two models, there research will be to determine whether there was a
is dis-
agreement in a basic assumption each makes.
single The
or multiple origin(s) of maize and to link routes
Rio Balsas model assumes that maize was important
of human movements with maize dispersals. Promis-
all over Central America and South America;ing
in areas
con-for more intensive archaeological recon-
trast, the Tehuacan model postulates that maize wasinclude Archaic caves on the tropical flanks
naissance
relatively unimportant south of Mesoamerica (i.e.,
of the big bend of the Rio Balsas, the Zacapa Valley
in Nicaragua and Costa Rica) until it reached
in Guatemala, the eastern slopes of the Sierra Madre
in Veracruz and Tabasco, as well as the mountains
Panama, where it flourished, as it did in northern
of Belize, Honduras, and Guatemala.
South America. Also, according to the Tehuacan
model, maize occurred throughout the southern
In part
addition to providing a model system for char-
of Mesoamerica, moved northward into the acterizing
states ofthe evolutionary mechanisms of crop plant
Tamaulipas and Zacatecas in Mexico, and spread and domestication, intercalation of the
speciation
north into the U.S. Southwest. evidence from these experimental prototypes of early
maize with the archaeological and paleoecological
Conclusion
record provides a unique opportunity for under-
Although aspects of the Rio Balsas model may be
standing how the evolution of maize is intricately
untenable in regard to the origin of maize, this interwoven
does with culture history and environmental
change. This is no trivial academic exercise. Maize
not preclude plant domestication and agricultural
is one of 3 grains, along with wheat and rice, that
development in the tropics. The evidence of lowland
provide the cornerstone of the world food supply
tropical plants, including avocados, black sapotes,
and beans imported into the highlands, suggests today.
that With the rapid acceleration of human popu-
lation
such developments took place. Rather than a rival or growth, reduction in amount of land available

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18 LATIN AMERICAN ANTIQUITY [Vol. 11, No. 1, 2000

Dewald, C. L., B. L. Burson, J. M. J. De Wet, and J. R. Harlan

for agriculture, and global climate change, there is
1987 Morphology, Inheritance, and Evolutionary Significance
serious concern about the ability of agriculture to of
feed
Sex Reversal in Tripsacum dactyloides (Poaceae). Amer-
the projected population of 12 billion humans on icantheJournal of Botany 74:1055-1059.
Doebley, J.
planet by 2050. The more we know about important
1990 Molecular Evidence and the Evolution of Maize. Eco-
crop plants, their relationships to their wild relatives,
nomic Botany 44 (suppl.):6-27.
their origins, their ecological adaptations, and howJ. F., M. M. Goodman, and C. W. Stuber
Doebley,
1984
and under what conditions humans exploited Isoenzymatic
and Variation in Zea (Gramineae). Systematic
Botany 9:203-218.
altered them in the early stages of domestication, the
1985 Isozyme Variation in the Races of Maize from Mexico.
greater our chances will be for identifying beneficial
American Journal of Botany 72:629-639.
1987
genes from wild plants and possibly recovering Patterns of Isozyme Variation between Maize and Mex-
lost
ican Annual Teosinte. Economic Botany 41:234-246.
ancient alleles useful for development of more pro-
Doebley, J. F., W. Renfroe, and A. Blanton
ductive varieties adapted to a wider range of 1987
envi-
Restriction Site Variation in the Zea chloroplast Genome.
ronmental conditions in the future. Genetics 117:139-147.
Doebley, J. F, A. Stec, J. Wendel, and M. Edwards
1990 Genetic and MorphologicalAnalysis of a Maize-teosinte
Acknowledgments. We are grateful to Jane Libby for her untir-
F1 Population: Implications for the Origin of Maize. Pro-
ing and able editorial assistance. A special thank you to Walton
C. Galinat and H. Garrison Wilkes for their helpful comments ceedings of the NationalAcademy of Sciences 87:9888-9892.
Dowling, T. E., C. Moritz, J. D. Palmer, and L. H. Rieseberg
on the first draft of the manuscript. We are especially indebted
1996 Nucleic Acids III: Analysis of Fragments and Restric-
to the valuable insights of several anonymous reviewers and
tion Sites. In Molecular Systematics, 2nd ed., edited by D.M.
their suggestions for focusing the manuscript more on content
Hillis, C. Moritz, and B.K. Mable, pp. 249-320. Sinauer
than on controversy. M. W. Eubanks gratefully acknowledges
Associates, Sunderland, Massachusetts.
research support under National Science Foundation grants
Dunn,no.
M. E.
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