Journal of Ethnobiology 25(2): 155–174 Fall/Winter 2005

HUMAN SELECTION AND DOMESTICATION OF CHIA

(Salvia hispanica L.)

JOSEPH P. CAHILL
University of California, Riverside, Riverside, CA 92521

ABSTRACT.—The plant species Salvia hispanica L. has a long history of plant-

human interaction. This study defines the human-selected morphological
characters associated with domestication of S. hispanica and presents a review
of ethnobotanical and historical information pertaining to the human selection
forces that resulted in changes in morphology. The compiled ethnobotanical
information for S. hispanica is applied to the framework of the hypothetical
ecological-evolutionary continuum of plant-human interactions, highlighting the
contributions of ethnobotanical data to our understanding of plant domestication
processes.

Key words: Salvia, domestication, Mexico, ethnobotany, agriculture.

RESUMEN.—La planta Salvia hispanica L. tiene una larga historia de interacción
con los humanos. Este estudio define los caracteres morfológicos seleccionados
por el hombre, asociados con la domesticación de S. hispanica, y presenta una
revisión de la información etnobotánica e histórica relacionada con las fuerzas
humanas de selección que provocaron los cambios morfológicos. La información
etnobotánica compilada para S. hispanica se enmarca en el hipotético continuo
ecológico-evolutivo de las interacciones planta-hombre, destacándose las
contribuciones de los datos etnobotánicos para nuestra comprensión de los
procesos de domesticación de plantas.
RÉSUMÉ.—La plante nommée Salvia hispanica L. est utilisée depuis fort
longtemps par les humains. Dans cette étude, nous décrivons non seulement
les caractères morphologiques du S. hispanica liés à la domestication et
sélectionnés par les êtres humaines, mais nous faisons également une synthèse
des données ethnobotaniques et historiques se rapportant aux forces sélectives
exercés par les humains qui ont abouti à des modifications morphologiques. Les
données ethnobotaniques recueillies pour la S. hispanica sont discutées dans le
contexte hypothètique d’un continuum écologico-évolutif des interactions plante-
homme; ce qui contribue à faire ressortir l’importance des données ethnobota-
niques pour mieux saisir les processus de domestication des plantes.

INTRODUCTION

This paper examines traditional plant management techniques and direct

human selection as forces effecting morphological changes in the crop plant
Salvia hispanica L. from an ethnobotanical perspective. In pre-Columbian
Mesoamerica, the species known commonly as ‘‘chia’’ was a major commodity
and its seeds were valued for food, medicine, and oil (Cahill 2003). After Spanish
156 CAHILL Vol. 25, No. 2

contact and colonization, the level of cultivation of the species plummeted.

Limited cultivation and traditional medicinal, culinary, and religious uses persist
today; however, the origins of cultivation and the domestication process are
unknown (Cahill 2003). The ecological-evolutionary-economic model of plant
domestication developed by Harris and others is expanded upon by considering
data from ethnobotany and the genetics of human selected traits for S. hispanica
(Harris 1989; Hynes and Chase 1982; Rindos 1980; Weirsum 1997).
Ethnobotany has contributed considerably less than other scientific dis-
ciplines to models of plant domestication in part because ethnobotanical
knowledge related to human selection is less abundant than knowledge of plant
use. With respect to Salvia hispanica, knowledge of selection is also less abundant
because typically only one or a few people in a community cultivate the species.
Few studies of plant domestication include ethnobotanical data, and still fewer
correlate management techniques with morphological variation in the crops. An
exception is Nabhan’s classic investigation of Hopi blue maize, in which he
found a positive correlation between depth of planting and increased seed size
(Nabhan 1989). Casas and Caballeros found that local management methods
selected for certain phenotypes such as larger plants in the arborescent genus
Leucaena (Fabaceae) (Casas et al. 1996). However, these studies focused on
management and cultivation techniques within communities, and did not
encompass the full range of any species in terms of geographic and morphologic
diversity. Given the successful dispersal of useful plants, not all aspects of plant
domestication processes can be presumed to take place in one location. Thus, at
a given point in time and place the full continuum of the domestication process
cannot be observed. Application of the more generalized Harris model to
ethnobotanical data that covers the geographic, morphological, chronological,
and cultural diversity of Salvia hispanica reveals the strength of ethnobotany as
a tool in understanding domestication. A model of plant-people interactions for
S. hispanica based on ethnobotanical data can be used to develop testable
hypotheses about domestication that can be evaluated with genetic and other
data for the species.

Salvia hispanica AS AN EXAMPLE OF DOMESTICATION

For plant species, increased range of morphological and phenological

diversity, suppression of natural mechanisms for dispersal and protection,
gigantism, and other characteristics have been associated with domestication (De
Wet and Harlan 1975; Harlan 1975; Hawkes 1983; Schwanitz 1967). Salvia
hispanica, having a spectrum of domesticated varieties, is no exception. For the
purpose of this study, varieties have been categorized loosely as ‘‘wild,’’
‘‘primitive domesticates’’ (plants in cultivation that have been modified from
wild-type morphology), ‘‘full domesticates’’ (plants that lack dispersal mechan-
isms and are dependent on human cultivation for survival), and ‘‘advanced full
domesticates’’ (plants that exhibit human-selected traits and also lack dispersal
mechanisms). This paper expands on published descriptions of the morpholog-
ical details that distinguish wild and domesticated varieties of Salvia hispanica
(Cahill 2004; Cahill and Provance 2002).
Fall/Winter 2005 JOURNAL OF ETHNOBIOLOGY 157

With each mature plant capable of producing a thousand seeds or more, the
progeny of a single plant could supply a large portion of the next year’s crop.
This high fecundity combined with a short annual life cycle and a strong genetic
component to inheritance for some domesticated traits made human selection in
Salvia hispanica a powerful force in evolution of the crop (Cahill and Ehdaie 2005).
These biological characteristics considered in conjunction with ethnobotanical
data related to selection make S. hispanica an excellent candidate for the
examination of plant-people interactions leading to domestication (see Table 1).

MATERIALS AND METHODS

The morphological descriptions presented are based on a collection of 42

accessions categorized as wild or domesticated from the known range of the
species (Cahill 2004). Recording of observations took place in common garden
and common greenhouse experiments in Riverside, California, designed to
eliminate effects of plasticity and to gauge genetic differences (Cahill and Ehdaie
2005; Cahill and Provance 2002). Herbarium voucher specimens were deposited
at the herbarium of the University of California, Riverside (UCR)1 and are
referenced in this study whenever possible. Field work was conducted during the
winter of 1998–1999 in Pachali, Sacatepequez, Guatemala (lat 14u389 N, long
90u389 W, elev. 2200 m), in the Tarascan village of Capacuaro, Michoacan, Mexico
(lat 19u339 N, long 102u039 W, elev. 2300 m) and in Santa Lucia, a rural region
near Buena Vista, Santa Rosa, Guatemala (lat 14u289 N, long 90u149 W, elev.
2120 m). Field work was also conducted at the end of 1999 in the Coran village of
Mesa de Nayar, Nayarit, Mexico (lat 22u139 N, long 104u419 W, elev. 1800 m) and
in Acatic, Jalisco (lat 20u469 N, long 102u539 W, elev. 1720 m). The information
presented is based on field interviews and participant observation when possible.
In each location, a list was compiled of individuals who reportedly had
knowledge of Salvia hispanica. Plants were collected and used during informant
interviews as described by Cahill (2003). In each location, about twenty
informants who volunteered information were re-interviewed using a more
extensive set of questions (Cahill 2003). While many informants had extensive
knowledge of uses of the plant, typically only one or two individuals had
knowledge related to collection or management of wild plants or cultivation of
the species in western Mexico. Information for those informants has been
provided by Cahill (2003) and they are referred to by name below.

COLLECTION FROM WILD POPULATIONS (GATHERING)

The descriptions of human manipulation of wild populations of Salvia

hispanica presented here roughly correlate to ‘pre-domestication’ components of
the domestication process continuum (e.g., Harris 1989; Rindos 1984). Wetlands
and areas along streams above 1800 m in pine or oak forest of Mesoamerica
constitute ideal habitat for wild populations of S. hispanica. In Santa Lucia,
Guatemala, a farmer called Lazaro del Cid and his family explained how the
seeds of S. hispanica (Cahill and Castillo 2948, UCR), known locally as chan, were
collected from the wild and mixed with seeds collected from wild Salvia tiliaefolia
 158
TABLE 1.—Continuum of plant-people interactions for Salvia hispanica.1
Food yielding Morphological/
system Human action Ecological effect physiological effect Inheritance Uses2–parts
Wild plant-food Collecting seed, Change in population None observed None observed M, C–all
procurement vegetative parts size
(foraging)
Selective seed Unintentional White-seeded Single genes; M, C–all
collecting dispersal by humans plants qualitative traits
Cultivation Selective seed Dispersal to new Change in seed coat Single genes; M–seed oil
(planting) collecting agricultural habitats patterns and colors, qualitative traits
loss of stem
pigmentation stripes
Selective seed Dispersal to new Increased branching Quantitative traits M–seed oil
collecting agricultural habitats
Natural germination/ Decreased competition None observed None observed M–seed oil
tolerance of
volunteers
CAHILL

Broadcasting–bare Increased population Decreased genetic Unknown M–seed oil

soil size diversity; uniform
germination
Broadcasting– Transformation of None observed None observed M–seed oil
roughing of soil existing vegetation
Covering seeds with Increased seed Increased seed size; Quantitative, many M–seed oil
soil survival uniform germination genes
rate
Intercropping Competition Increased plant height; Many genes; M–seed oil
decreased branching quantitative, traits
Burning Enhanced Increased plant height Many genes; M–seed oil
productivity quantitative, traits
Irrigation Enhanced Increased plant height Many genes; M–seed oil
productivity; less quantitative, traits
water stress
Vol. 25, No. 2
TABLE 1.—Continued.
Food yielding Morphological/
system Human action Ecological effect physiological effect Inheritance Uses2–parts

Agriculture Selective seed Decreased genetic Changes in seed coat Single or few genes; M, C–seed
(farming) collecting diversity patterns and colors qualitative traits
Fall/Winter 2005

Selective seed Decreased genetic Excessive stem and Several genes; M, C–seed
collecting diversity inflorescence qualitative traits
pigmentation; dark
blue flower color
Selective seed Decreased genetic Further increase in seed Many genes; M, C–seed
collecting diversity size quantitative traits
Selective seed Decreased genetic Foliage scent Unknown M, C–seed
collecting diversity
Irrigation, furrow Decreased water Gigantism in seed and Many genes; M, C–seed
planting stress, greater flower quantitative traits
productivity
Harvest whole plant Dependent on human Closed calyx (loss of Few genes; modifier M, C–seed
cultivation dispersal mechanism) genes; reinforcement
(domestication)
Threshing Unknown Decreased calyx Many genes; M, C–seed
pubescence quantitative traits
Cultivation in humid Adaptations to Varied ripening time Many genes; M, C–seed
environment tropical environment (extended harvesting); quantitative traits
JOURNAL OF ETHNOBIOLOGY

increased calyx
pubescence
Monocropping, Decreased Rapid growth; more Many genes; M, C–seed
mechanized competition from flowers per axil; apical quantitative traits
harvest weeds dominance; uniform
germination; uniform
height; uniform seed
maturation
Monocropping, Unknown Allelopathy Unknown M, C–seed
mechanized harvest
1
Sources: this work; Cahill and Ehdaie 2005; Cahill and Provance 2002; Hernández Gomez 1989, 1994.
159

2
M–medicinal; C–culinary
160 CAHILL Vol. 25, No. 2

Vahl. The farmer stores the seed mixture in a glass jar, and combines portions of
the seed mixture with water to form a medicinal beverage. Collection of small
quantities of seed with no or minimal modification of environment is sustainable.
In Santa Lucia, the vigorous wild population grows along mountain streams and
has only wild type morphological characters, including short height (less than
1 m), striated stem pigmentation of anthocycanin, pubescent calyxes, open
calyxes, small black seeds (less than 12.0 mg/100 seeds), short inflorescence (less
than 15 cm), leaves and light blue corollas that are smaller than the domesticated
varieties, and bushy growth form.
Similarly, in the Tarascan villiage of Capacuaro, informants identified local
wild populations of Salvia hispanica as cueruni. The term cueruniquarani meaning
‘‘to gather chia’’ in Tarasco was also frequently mentioned and has been reported
by others (Gilberti 1983; Sandoval 1989). Because Tarascan use is limited to
medicinal use of small quantities of seeds, again there was no observable
morphological variation accompanying this pattern of use, nor were there any
apparent effects on population size.
In other wild populations, humans practice selective collection. For example,
the informant Silvino Lobatos Estrada of Mesa de Nayar described in detail the
technique for collection of seeds from the local wild population of Salvia hispanica
(Cahill 2979, UCR). The pine forest of the region contains a habitat, known locally
as ‘‘the swamp,’’ where this dense wild population flourishes. As with other wild
populations, dark-seeded plants are present; however, in Mesa de Nayar
a patchwork pattern exists with white-seeded wild plants, resulting in
a heterogeneous population with respect to this character. According to Silvino
Lobatos Estrada, selective collection of dark-seeded plants proceeds under the
premise that white seeds do not have the ‘‘strength’’ of darker ones. Whether the
‘‘strength’’ pertains to taste or some property of the culinary flour made from the
seeds is not clear. Since the flour forms the main ingredient in an atole beverage
consumed during a religious ceremony, the rationale behind the ‘strength’ of
dark seeds may have origins in superstition or religious belief. Though seasonal,
culinary usage requires collection of large quantities of dark seeds. This may
have the effect of selecting for an increase in the frequency of white-seeded plants
in the population.
Informants described the process by which white seeds are excluded from
collection. First, the seed color of an individual plant is determined by shaking
a small quantity of seed out of the calyxes. If the plant is white-seeded the seeds
are dropped to the ground. This shaking and dropping assists dispersal,
spreading seeds further from the parent plant, resulting in decreased seedling
competition for white-seeded plants. Apart from collection of seeds, informants
clearly indicated that no other techniques such as broadcasting, thinning,
burning, or weeding were employed.
I had assumed that white seeds were intentionally selected for because
certain fully domesticated varieties produced in the valley of Mexico consist
exclusively of white-seeded plants, suggesting direct human selection for white
seeds. Modest natural selection may occur in wild populations as white seeds
contrast with the dark soil, making white seeds more susceptible to predation.
However, human selection is undoubtedly more intense. The white-seeded
Fall/Winter 2005 JOURNAL OF ETHNOBIOLOGY 161

phenotype is rare in wild populations, with no other reported white-seeded

plants from wild populations other than Mesa de Nayar. Since geography,
altitude, and flowering time form reproductive barriers between wild types and
domesticated varieties, gene flow from domesticated to wild populations seems
an unlikely origin of white-seeded plants. The rise of white-seeded plants in wild
populations, ancestral to domesticated varieties, was likely a more complex
process as exemplified by the Mesa de Nayar population. There, approximately
60 percent of plants are white-seeded, a trait known to be recessive and
controlled by a single gene (Cahill and Provance 2002). This relatively high
percentage may be the result of the generations of collection of dark seeds that
left more white-seeded plants to disperse, artificially increasing fecundity for
white-seeded plants. A unique approach to further research would involve
a synthesis of population genetics and ethnobotany, beginning with an extensive
survey of population size, frequency of white-seeded plants, and intensity of
human collection.
In the case of wild populations of Salvia hispanica, ethnobotanical evidence
indicates only small quantities of plant material are collected. No evidence for or
information about management of wild populations other than that based on
seed collection, and no modern examples of reliance on the species as a primary
or secondary source of nourishment, have emerged. This may account for the
absence of ethnobotanical accounts of management techniques such as
broadcasting, burning, or weeding that have been reported for other gathered
wild species (see Table 1). The examples from S. hispanica provide documentation
of direct selective forces on wild populations and the resultant morphological
changes, offering important insights into beginnings of the domestication
process.

PRIMITIVE DOMESTICATES

Archaeological Evidence.—The seed crop Salvia hispanica has taken a simple path to
domestication, making it an excellent model for examining the plant domesti-
cation process. Unfortunately, little archaeobotanical evidence of chia has been
found (Gonzalez Quintero 1981, 1986; Hard and Roney 1998). S. hispanica is
a herbaceous annual with soft oily seeds, so there are no portions of the plant that
preserve particularly well, making it difficult to draw any conclusions regarding
domestication from this evidence. As a result, the time of earliest cultivation and
domestication of chia are unknown.
Historical Evidence.—Economic historians have suggested Salvia hispanica as
a staple food was as important as maize, and in some areas more important
(Harvey 1991; Hebert et al. 1995; Hunziker 1952; Perm and Carrasco 1974; Rojas
Rabiela 1988). The codices of sixteenth-century Mexico indicate that large areas of
agricultural land were devoted exclusively to chia cultivation. For example, 21 of
the 38 Aztec provincial states gave chia in annual tribute and separate records
from independent states such as Huexotzinco also list chia as tribute (Berdan and
Anawalt 1996; Perm and Carrasco 1974). Seeds from wild populations formed
a portion of this tribute, but the quantities recorded in tribute lists would have
162 CAHILL Vol. 25, No. 2

necessitated cultivation. Human transportation of chia seeds and the

resulting dispersal to new habitats was perhaps the greatest ecological effect of
cultivation. Sandoval reviewed the sixteenth-century geography of S. hispanica
cultivation and concluded it was widely grown until the later part of the
sixteenth century, at which time it was displaced by European crops (Sandoval
1989). The origins of cultivation for this species remain unclear, though several
domesticated varieties with incremental degrees of human selected traits still
survive.
From 1575–1577, Fray Bernardino de Sahagún compiled the Florentine
Codex, working with Nahua informants in central Mexico who retained
memories of life prior to arrival of the Spanish. Sahagún described a method
of planting chia that inadvertently selects for larger seeds. The description and
illustrations of chia, subsequently recognized as Salvia hispanica by numerous
botanists, describes how sown chia seeds are ‘‘...covered over with the soil, just
thinly, just smoothed over with the foot.’’ (Sahagún 1950–1982, book 10, chapter
20, page 75). Smaller seeds do not contain enough stored resources to reach
the surface and are selected against. Though unintentional, increased seed
mass represents the first quantitative trait to be affected by human activities
associated with the species. Seed mass samples collected from common
garden and common greenhouse plants of different accessions showed seed
mass for individual plants and within accessions did not vary significantly
(Cahill and Ehdaie 2005). These results also indicate a significant difference
in seed mass between wild and domesticated accessions. Since each plant
is capable of producing thousands of seeds at maturity, the small increase
in seed size translates into markedly greater yields. Apart from the
essentially wild material collected in Pachali (see below), all other primitive
domesticated varieties in the current germplasm collection exhibit increased seed
size.
Ethnobotanical Evidence.—Ethnographic examples of plant manipulation are
presented that correlate to certain morphological characteristics. The examples
begin with the examination of the most morphologically primitive domesticated
varieties and the associated management techniques which have shed light on
the origins of cultivation. In Pachali, Department of Sacatepequez, Guatemala,
local farmers managed the growth of wild Salvia hispanica in fields of other crops,
including beans and maize (Cahill and Castillo 2970, UCR). After the bean
harvest, plants of S. hispanica were left standing to mature while all other species
had been cleared. The sporadic chia plants standing in stark contrast to the neatly
harvested and cleared field emphasize the value of the species and the effort put
into its cultivation. When asked if the seeds were planted, the response was an
emphatic no, and informants went on to explain that seedlings just appeared in
the same place year after year and were intentionally spared when weeding.
Plants do not spread within fields from year to year and do not appear on
margins or outside of the fields as weeds. Small quantities of seed are collected in
January and stored for use in medicinal beverages throughout the year. This
management by tolerance of volunteers has preserved many pre-Columbian
primitive domesticated varieties that formerly may have been grown as primary
Fall/Winter 2005 JOURNAL OF ETHNOBIOLOGY 163

crops. The material from Pachali has retained wild morphological characters, and
is indistinguishable from other Central American wild types when grown in
a common garden. Wild populations exist in the vicinity of Pachali, suggesting
that human or natural dispersal from a wild population gave rise to these plants
found in cultivation. With no human-selected traits and no actual sowing of
seeds or tillage, the Pachali material represents a very early stage of
domestication where plants are cultivated but no apparent modifications from
wild type morphology are observed. The presence of Salvia hispanica in this
cultivated setting seems largely dependent on the ecological effect of selective
weeding in the fields, without which S. hispanica would likely be out-competed
and eventually disappear from the fields. Management via cultivation continues
to play an important role in preserving varieties.
Planting of Salvia hispanica seeds in cultivated settings traditionally proceeds
by broadcasting seeds into the wind without soil preparation. Hernández Gómez
reported this technique in the settlements of Olinalá and Temalacacingo in the
Mexican state of Guerrero (Hernández Gómez 1989, 1994). Kelly reported the
identical technique in Ayotitlán, Jalisco (Kelly 1944). Generally, broadcasting
increases population size and density, resulting in greater productivity in
comparison to natural dispersal. Since later-germinating seeds are out-competed
in these dense cultivated plots, domesticated varieties have developed rapid and
uniform germination. Comparative observations made in germinators at
a constant 30uC showed wild types do not germinate uniformly and rapidly,
but sporadically over a 10-day period. Wild material from Mittlenam, Santa
Lucia, Sinaloa, Mexico, from the collection of the late Howard Scott Gentry best
exemplified this sporadic and delayed germination. In contrast, all primitive and
fully domesticated varieties germinated within two days. Several published
studies have described germination of 90 to 100 percent within 48 hours with no
difference between white and dark seeds, but these studies do not indicate
whether the seeds originated from domesticated, cultivated, or wild plants
(Labouriau and Agudo 1987; Naqvi et al. 1992).
The Mexican agronomist Rulfo provided invaluable accounts of traditional
cultivation techniques for Salvia hispanica, including preparation of soils by
‘‘roughing’’ (Rulfo 1937). Roughing represents the first significant increase in
human energy input into a system of cultivation. Disruption and loosening of the
soil surface provides an environment where more seeds are hidden, escaping
predation and facilitating establishment of seedling roots. While Rulfo described
roughing in conjunction with broadcasting, other traditional cultivation methods
build on roughing by covering seeds with soil.
The morphologically most primitive of these large-seeded domesticates has
a restricted range of cultivation centered near Merida in the Yucatan peninsula of
Mexico. In addition to larger seeds, the gene for white seed coat is present in the
gene pool. Apart from these two characters, the plants resemble typical wild
ecotypes. The presence of the gene for white seed coat in otherwise
morphologically primitive domesticated varieties suggests early human selection
for this trait. In the Florentine Codex, Sahagún described a variety of Salvia
hispanica known as iztac chia ‘white chia’, illustrated with white seeds (see
Figure 1) (Sahagún 1950–1982). Certainly, white seeds should not be considered
164 CAHILL Vol. 25, No. 2
Fall/Winter 2005 JOURNAL OF ETHNOBIOLOGY 165

a compulsory character among the large-seeded primitive domesticates, but early

selection for the trait seems likely.
Material cultivated in Temalacacingo, Guerrero, Mexico, does not produce
white-seeded plants, but human selection for increased branching results in an
increased number of inflorescences per plant. It is also associated with large
seeds in this primitive domesticated variety. In his agronomic assessment trials,
Hernández Gómez found it to be more productive than many of the fully
domesticated varieties exhibiting greater numbers of human selected traits
(Hernández Gómez 1994). However, this variety fully disperses seeds, making it
difficult to harvest without affecting yield. In a common garden experiment at
the University of California, Riverside, plants of this variety became so heavy
with seeds the main stems broke, so plants were propped up. In Temalacacingo,
chia is reportedly still cultivated in tlacololes, plots that have been recently
slashed and burned, associated with maize (Hernández Gómez 1994). The
management techniques of burning, broadcasting, and covering seeds require
more labor than the simple tolerance for plants observed in Pachali. The degree
to which these management techniques are directed specifically at chia rather
than the associated crops deserves more investigation, since cultivated forms of
Salvia hispanica grow predominantly in mixed-crop settings today. In Tlalhuica in
1564, chia reportedly grew on the same irrigated plots of land with beans and
maize, sown between the maize (Rojas Rabiela 1988). In November of 1999, Cora
informant Reyes Camarena of Mesa de Nayar reported seeing mixed-crop
cultivation of Salvia hispanica with calabash (Cucurbita sp.) in the Sierra San
Andras and Santa Katarina. The erect stems of S. hispanica provide runners for
the gourd vines to climb. Whether directed at S. hispanica or not, the management
regime in such mixed-crop settings has produced quantitative morphological
variation.
From an ecological perspective, competition introduced by mixed cropping
could account for some quantitative morphological changes. Increased compe-
tition from crops of greater stature like maize could result in the increased height
seen in certain advanced fully domesticated varieties. The increased branching
observed in material from Temalacacingo could be the result of competition from
crops, decreased competition due to weeding, or a particular management
technique. Despite morphological changes that enhance productivity in
comparison to wild Salvia hispanica, primitive domesticated varieties have
largely been displaced by fully domesticated varieties with a greater number
of human selected traits. This process of displacement would have begun after
the appearance of full domesticates in pre-Columbian times. As a result,
primitive domesticated varieties are rarely encountered today and their survival
seems bleak, given their dependence on traditionally cultivated plots of land and
the general shift towards agricultural modernization in Mesoamerica.

FIGURE 1.—Illustrations of three Nahuatl varieties of Salvia hispanica from the sixteenth-
century Florentine Codex: iztac chian ‘white chia’ (top); mitzic chian, a type of black chia
(middle); and chian tzotzol ‘wrinkled chia’ or ‘chia that is ground in water’ (bottom).
(Source: Sahagún 1950–1982).
166 CAHILL Vol. 25, No. 2

FULL DOMESTICATES

Historical Evidence.—Francisco Hernández, writing from 1571–1577, described

how ‘‘[t]his plant chia grows wherever it seeds, principally in cultivated,
irrigated, and watery places’’ (Hernández 1959:207–210). This and other
sixteenth-century accounts stress the importance of water for successful
cultivation of chia. Over 9000 ha of land in the early 1500s were cultivated
under the traditional chinampa raised field ‘‘floating garden’’ system (Armillas
1971). In the chinampas, chia faced an extended growing season not affected by
drought (Coe 1964; Rojas Rabiela and Sanders 1985). Unique agricultural
techniques practiced in the chinampas may have worked as selective forces
upon morphology. Sandoval believes chia was sown directly and was not
grown in almacigos (seedling nurseries) or in chapines (muck cubes) (Sandoval
1989). The agricultural techniques applied to chia in the chinampas and
possible resulting morphological variation remain unresolved issues. While
chia constituted one of the major crops of the chinampas, after Spanish
conquest chia cultivation in chinampas ceased (Harvey 1991; Weaver 1981).
In contrast, Dr. Juan Jimenez-Osornio reported occasional cultivation of
Salvia hispanica in the modern chinampas system.2 The history of chia
cultivation in the wet soils of the chinampas is a reflection of the dependence
of ancestral wild ecotypes on wet soils. This wild heritage translates well to
the modified aquatic system of the chinampas. The ecological effect of
decreased or lacking water stress combined with the extended growing season
may have allowed for selection of traits associated with fully domesticated chia
varieties.
In the mid-sixteenth century, Sahagún described ‘‘iztac chian’’ or white
chia as ‘‘...that which is uprooted, which can be rubbed in the hands. I
uproot chia. I rub chia in my hands.’’ (Sahagún, 1950–1982). This technique of
hand threshing signals the development of the second major increase in
human energy input associated with Salvia hispanica. This laborious threshing
technique also requires harvesting and drying of the whole plant, a departure
from seed selection of wild and primitive domesticated varieties, which proceeds
by shaking seeds off the unharvested mature plants into textiles or baskets
(Castello Yturbide 1972; McVaugh 1943). In the tenth book of the Florentine
Codex, Sahagún described the world of the Aztec markets; a seller of wrinkled
chia describes how he sold ‘‘unthreshed’’ chia plants in markets, how ‘‘he
rubs it between his hands to clean it’’ (Sahagún 1950–1982). In all wild types
and primitive domesticated varieties, seeds fall from the plant with just
a brush or strong wind. The transport of harvested inflorescences to Aztec
markets suggests that non-shattering morphological forms already existed by
the 1500s. In addition, linguistic evidence of cultural knowledge of non-
shattering domesticated forms appears throughout Mesoamerica. Examples
include the reported seventeenth-century use of the term ti her, ti bak aaq,
which means ‘‘to thresh chia seeds’’ in the Mayan language Cakchiquel (Acuña
1983), and the more recent use of qui tu thohui inchonı́, which means ‘‘to
thresh chia’’ in the indigenous language Matlatzinca (Basalenque 1975; Sandoval
1989).
Fall/Winter 2005 JOURNAL OF ETHNOBIOLOGY 167

Morphology of Full Domesticates.—The group of full domesticates is defined by

their closed calyxes, larger seeds, variability in calyx pubescence, and gigantism.
Closed calyxes prevent seed dispersal and effectively prevent survival of fully
domesticated varieties outside of human cultivation. This character meets the
criterion for ‘‘full domestication’’ as described by Harlan (1975) or ‘‘agricultural
domestication’’ as described by others. None of the herbarium specimens of wild
populations or wild material sampled for this study exhibit any degree of closure
of calyxes. To date, the trait has been observed only in the fully domesticated
varieties of Salvia hispanica and has not been reported in wild populations or
related species of Salvia. Like other qualitative traits in the species, closure of
calyxes could have developed as the result of conscious human selection.
Fully domesticated varieties exhibited seed mass ranging from 15.0–
16.5 mg/100 seeds, a large increase in comparison to wild types, but only
a slight increase (though statistically significant) over primitive domesticated
varieties. Furthermore, successful selection for increased seed mass has a strong
genetic component demonstrating the potential for strong, rapid, and effective
human selection (Cahill and Ehdaie 2005). However, no correlation between seed
size and white color was observed among varieties.
Since dried calyx trichomes can induce sneezing, decreased calyx pubescence
relieves the hand thresher from incessant irritation. The popular Mexican
varieties chia poblana from Cuatepec, Puebla, and others grown in Sonora typify
fully domesticated varieties with decreased density of calyx pubescence. Fully
domesticated varieties from Central America, including those now cultivated in
Jutiapa, Guatemala, San Salvador, El Salvador, and throughout Nicaragua,
characteristically display an increase in the density and length of calyx
pubescence in comparison to wild populations. Material from Nicaragua exhibits
these traits to the greatest degree. For Salvia hispanica, an increase in the length
and density of calyx pubescence is associated with adaptation to a more humid
environment; in the absence of enhanced pubescence, precipitation causes the
mature dry calyxes to soak up water, which renders the enclosed mature nutlets
susceptible to hydration. Seeds release a polysaccharide mucilage upon
hydration, causing the ripe inflorescence to develop a sticky coating that hardens
upon drying, destroying the crop. The dry winters of Mexico do not present
a problem, but cultivation in more humid areas outside the natural range of the
species, such as Nicaragua, necessitates protection from precipitation. An
additional novel adaptation to a humid environment found in Nicaraguan
domesticates is lack of determinacy in timing of flowering and ripening of seeds.
In the common garden experiment mentioned above, individual plants flowered
continuously from December through May and retain leaves, suggesting
a selective trend away from annualism.
A compact inflorescence, defined by small space between glomeruli and
number of flowers per axil, appears in all fully domesticated varieties. Wild
populations of Salvia hispanica have great variation with respect to these traits. As
a general rule, however, the higher the elevation the more compact the
inflorescence becomes.
Some degree of gigantism occurred in all fully domesticated varieties
examined in a common garden setting (see Table 2).
168 CAHILL Vol. 25, No. 2

TABLE 2.—Results of common garden and green house experiments in Riverside,

California: quantitative variation in Salvia hispanica.
Primitive Full Advanced
Wild domesticate domesticate domesticate
Corolla length (mm) 4.5 6 0.5 4.6 6 0.1 6.1 6 0.3 9.2 6 0.2
Labium length (mm) 3.7 6 0.5 3.2 6 0.2 4.4 6 0.5 9.0 6 0.1
Leaf length (cm) 7.4 6 1.6 14.0 6 0.7 14.9 6 2.5 18.3 6 0.2
Leaf width (cm) 5.6 6 0.7 10.2 6 0.3 10.4 6 1.8 12.2 6 0.6
Height (m) 0.98 6 0.5 1.1 6 0.2 1.0 6 0.3 1.83 6 0.4
Apical inflorescence 9.5 6 2.6 9.4 6 1.1 9.7 6 1.3 40.2 6 2.3
length (cm)
Mean seed masses 10.0 6 1.3 14.0 6 0.5 16.0 6 0.8 16.0 6 0.2
(mg/100 seeds)
20 plants of each variety listed were grown in a common garden and common greenhouse. The above
values are means obtained by pooling values for all plants for each group, i.e., wild, primitive
domesticate, full domesticate, and advanced domesticate. Unless otherwise noted, the varieties tested
were from Mexico.
Wild types: Mittlenam, Santa Lucia, Sinaloa (lat 25u539 N, long 107u269 W); Chiepetlan, Guerrero (lat
17u469 N, long 98u349 W); Cerro Verde, Teotitlan, Oaxaca (lat 17u489 N, long 97u519 W); Barranca del
Cobre, Chihuahua (lat 27u039 N, long 107u889 W); Buena Vista, Santa Rosa, Guatemala (lat 14u289 N,
long 90u149 W); Chiantla, Huehuetenango, Guatemala (lat 15u559 N, long 91u459 W); Mesa de Nayar,
Nayarit (lat 22u139 N, long 104u419 W); Comanja, Michoacan (lat 19u419 N, long 101u359 W); and Oja de
Agua, Jalisco (lat 20u059 long 103u529 W).
Primitive domesticates: Merida, Yucatán (lat 21u039 N, long 89u549 W); Temalacacingo, Guerrero (lat
18u069 N, long 100u129 W); and Cuescomapa, Guerrero (lat 18u039 N, long 99u539 W).
Full domesticates: San Salvador, El Salvador (lat 13u429 N, long 89u199); Huehuetenango, Guatemala
(lat 15u329 N, long 91u479 W); Jutiapa, Guatemala (lat 13u479 N, long 90u119 W); Managua market,
Nicaragua (lat 12u159 N, long 86u279 W); Cuatepec, Puebla (lat 18u309 N, long 98u539 W).
Advanced full domesticates: Acatic, Jalisco (lat 20u469 N, long 102u539 W); Oaxaca city market, Oaxaca
(lat 17u019 N, long 96u499 W); Tuxtapec, Oaxaca (lat 18u189 N, long 96u219 W); Cordoba, Veracruz (lat
25u409 N, long 100u189); Guadalajara market, Jalisco (lat 20u409 N, long 103u199 W); Calvio, Agua
Calientes (lat 21u549 N, long 103u029 W); and Durango (lat 24u089 N, long 104u519 W).

Advanced Full Domesticates.—While the fully domesticated varieties described

above are rare, a second group of fully domesticated varieties are more common.
They exhibit an even greater number of human-selected traits. These advanced
fully domesticated varieties are more uniform with respect to morphology and
genetic diversity and can be found in cultivation throughout Mexico (Cahill
2003). Three principal morphological characteristics separate this group from the
other fully domesticated varieties described above: increased inflorescence
length, apical dominance, and a further increase in plant height. An increase in
apical inflorescence length from about 6 cm to 40 cm represents the first
quantitative characteristic that presumably resulted from direct human selection.
Longer inflorescence would increase yield, but the character always appears in
conjunction with apical dominance so that only the inflorescence closest to the
apex has increased length, and secondary branches and inflorescences remain
normal or slightly stunted. There are clearly genetic factors controlling the
quantitative traits of apical dominance and plant height observed in common
garden and common greenhouse settings with well spaced plants. Hernández
Gómez has shown competition from close spacing of plants can also effect these
traits to some extent (Hernández Gómez 1989).
Fall/Winter 2005 JOURNAL OF ETHNOBIOLOGY 169

Advanced fully domesticated varieties traditionally grown in parts of Jalisco

and the central valley of Mexico produce considerably more anthocyanin
pigmentation on the stems and calyxes. In contrast to wild ecotypes with striated
stem pigmentation, stems of these varieties have an unbroken purple pattern that
extends to the calyxes. Some advanced fully domesticated varieties produce pure
purple calyxes, while in others pigmentation only occurs on the outer half of each
calyx. Observation of wild populations in the Sierra Madre Occidental indicate
some populations have a few plants that express slight purple shading on stems
and calyxes under stress conditions towards the end of the life cycle, but not
nearly to the degree seen in advanced fully domesticated varieties with purple
calyxes. This may be an example of aesthetic selection, but the reasons for
selection for excess anthocyanin pigmentation remain a mystery. In Amaranthus,
pigmented varieties had associations with pre-Columbian rituals. Given the
religious and cultural uses of Salvia hispanica, selection for religious use remains
a possibility, but no explicit accounts correlating any use with pigmented
varieties have been uncovered (Cahill 2003).
A shift towards use as a nutritionally reliable subsistence food and as a seed oil
accompanies cultivation of domesticated chia. Today, monocropping of the
advanced domesticated varieties has almost completely displaced the traditional
intercropping system of the primitive domesticated varieties and the non-advanced
full domesticates. It has led to a loss of genetic diversity over a wide geographic area.
Numerous studies have implicated cultivation and agriculture as causative
factors in the development of allelopathy (Einhellig 1996; Seigler 1996). The
beneficial chemical and ecological characteristic of allelopathy has accompanied
advanced full domestication. Preliminary experiments with aqueous root
leachates of the morphologically advanced domesticates of Salvia hispanica
applied to Echinochloa and Amaranthus seeds show statistically significant
inhibition of germination of Echinochloa seeds, but not for wild or any of the
other domesticated varieties tested (see Table 3). Further experimentation is
needed, both in isolating chemical compounds involved and identifying common
weeds associated with Salvia hispanica monocrop systems in Mexico.

Mechanized Agriculture.—In recent decades, mechanized agriculture has come to

dominate production of morphologically advanced fully domesticated forms of
Salvia hispanica. Mechanized agricultural techniques may act as selective forces
further reinforcing the morphological features of domestication. Uniform
morphology of numerous characters, particularly height and determinacy, is
found in varieties grown under mechanized agriculture, although it is not
completely certain that these traits are due to mechanized agriculture. In the
common garden plots, each Mexican advanced domesticated variety flowered
and matured at the same time and all of the inflorescences on each plant flowered
and matured at the same time in early October and set seed about 2–3 weeks
later, though the timing varied among the varieties. As plants began to flower,
leaves became necrotic and abscised, leaving a stem and ripe inflorescence. The
lack of leaves facilitates harvesting, particularly in mechanical settings. Wild
populations in Mexico also drop leaves as flowering begins, but individual plants
and inflorescences on the same plant do not synchronously flower.
170 CAHILL Vol. 25, No. 2

TABLE 3.—Allelopathic activity among wild and domesticated varieties of Salvia hispanica.
Amaranthus Echinochloa
Mean radical % radical Mean radical % radical
Treatments growth (mm) growth growth (mm) growth
Wild 1 10.7 130.4 16.6 102.4
Wild 2 10.4 126.8 17.0 104.9
Wild 3 12.3 100.8 21.9 106.8
Wild 4 13.6 111.4 18.9 91.2
Primitive domesticate 1 10.2 124.3 16.8 103.7
Primitive domesticate 2 8.3 101.2 19.7 121.6
Full domesticate 9.0 109.7 17.7 109.2
Advanced full domesticate 8.7 108.5 12.8 79.0*
Control 8.2 100.0 16.2 100.0
Aqueous extracts of Salvia hispanica roots were screened for activity against representative monocot
(Echinochloa crus-galli) and dicot (Amaranthus hypochondriachus) weeds. The aqueous extracts were
applied with a micropipette to a 3.5 cm2 disk of filter paper in a 5.5 cm2 diameter Petri dish. After
evaporation of the solvent, the disk was wetted with 0.2 cm3 of distilled water. Ten seeds of either E.
crus-galli or A. hypochondriachus were placed on the disk. The seeds were incubated in a germinator at
30uC for 24 hours. Root lengths were measured to determine mean root length. A randomized
complete block design with four repetitions was employed for each of the eight accessions of S.
hispanica and a control. The experiment was replicated twice to screen against the representative
monocot and dicot. An analysis of variance (ANOVA) was performed on resultant data. Of the
varieties of S. hispanica tested, only one domesticated variety exhibited significant activity, resulting in
inhibition of germination of the assay weeds (see * above). These preliminary results were intriguing
for two reasons. First, the results were opposite of what was expected, since most domesticated plants
lack allelopathic activity, but the trait is retained in wild relatives. Second, inhibition of germination
only appeared in the representative monocot. When allelopathic activity is identified in plant species,
it usually acts on both monocots and dicots (Einhellig 1996; Seigler 1996). Unless otherwise noted, the
varieties tested were from Mexico; for coordinates see Table 2: Wild 1: Mittlenam, Santa Lucia,
Sinaloa; Wild 2: Cerro Verde, Teotitlan, Oaxaca; Wild 3: Sierra Saguaribo, Durango (lat 25u349 N, long
105u419 W); Wild 4: Buena Vista, Santa Rosa, Guatemala; Primitive domesticate 1: Temalacacingo,
Guerrero; Primitive domesticate 2: Cuescomapa, Guerrero; Full domesticate: Cuatepec, Puebla;
Advanced full domesticate: Acatic, Jalisco.

Informant Jesus de Latorre Guiterrez of Acatic, Jalisco, described his method

for mechanical planting of Salvia hispanica. He plows furrows approximately
8 cm apart. On the ridges between the furrows, the machinery punches holes
about 2.5 cm in diameter, 5 cm deep, and 50 cm apart. He then drops several
seeds in each hole which is left open. The mesic microenvironment in the holes
facilitates germination and within a few days the seedlings emerge. With other
planting techniques, hydrated seeds tend to dry out, inhibiting germination and
increasing predation by birds and ants. Approximately 10–20 seeds are placed in
each hole, but only 3–5 plants per hole survive to maturity. Seedling competition
likely selects for larger seed mass and rapid germination, even though the seeds
are not covered with soil as with many other traditional cultivation techniques.
Two distinct varieties of morphologically advanced full domesticated
varieties are found in the Acatic region. The predominant variety, known locally
as chia pinta, forms heterogeneous populations. The majority of plants have gray
seeds with black marbling and lack stem pigmentation; a minority of plants may
be white-seeded, and a few have the striated stem pigmentation typical of wild
populations. Individuals of the second unnamed variety occur sporadically
among the vast fields of chia pinta, but produce only dark seeds and solid purple
Fall/Winter 2005 JOURNAL OF ETHNOBIOLOGY 171

pigmentation on stems and calyxes. This contrasts to the striated stem

pigmentation of wild types and the absence of stem pigmentation found in
nearly all advanced full domesticated varieties. Additionally, it flowers and
seeds slightly later than chia pinta. This second variety is distinct and has bred
true in three generations of experiments. Nabhan has stated, ‘‘many traditional
farmers sow multi-line mixtures of the same crop as a kind of insurance’’
(Nabhan 1989:73). This may have initially been the case in Acatic, as farmers
attempted to increase productivity. However, none of the informants were aware
of multiple varieties in their fields, and insisted they planted only chia pinta. With
mechanized planting and harvesting, nearly all of the traditional agricultural
knowledge associated with the crop has vanished. Each farmer saves seed from
year to year and does not buy seed or select individual plants for improved traits.
The only selective forces acting today on advanced full domesticates seem to be
those stemming from the use of mechanical sowing and harvesting equipment.
Ethnobotanical knowledge associated with direct selection methods and
traditional management techniques and how they might act as selection factors
in advanced full domesticated varieties has been lost with the conversion to
mechanized agriculture in the commercial production centers of Salvia hispanica
in Acatic and Jutiapa.

CONCLUSIONS

This investigation of Salvia hispanica demonstrates that a detailed ethno-

botanical study of wild and cultivated populations over the entire range of
a species can offer a more comprehensive understanding of the human selective
forces that set into motion the domestication process than one restricted to
a single community or region. While such studies can be more difficult to carry
out, from a biological or morphological perspective the benefits outweigh the
costs and the resulting data may encompass the entire continuum or stages of
interactions leading to domestication.
An evolutionary process for Salvia hispanica governed primarily by human
selection has emerged from this more comprehensive single species approach.
Whether intentional or unintentional, each instance of human selection could be
placed within the context of the ecosystem or agricultural management regimes
of the Harris continuum (see Table 1). Not all of the categories of Harris’s more
generalized continuum could be related to the data for a single species. Data that
fits in many of the cultivation and agriculture categories overlap, supporting the
concept of a continuum rather than distinct steps. In contrast, clear distinctions in
morphology, such as larger seed mass and non-shattering inflorescence,
specifically correlate with changes in human agricultural practices, suggesting
plant-human interaction for S. hispanica may be a stepwise process rather than
a continuum. While the ethnobotanical data presented in this paper by no means
offer a complete picture of the domestication process of S. hispanica, the
information does include known instances of selection, providing much needed
documentation of plant-human interactions.
Bye provided ethnobotanical evidence that human management of weedy/
wild populations results in quantitative morphological variation; management
172 CAHILL Vol. 25, No. 2

resulting in qualitative morphological variation, such as seed color, appears to

be less common (Bye 1981). Results from the analysis of Salvia hispanica
data suggest selection of qualitative traits occurred prior to incorporation of
the species into agriculture settings. The selection against and frequency of
white-seeded plants in the wild population of Mesa de Nayar best support
this conclusion. Unlike Bye’s weedy species, more intentional or intense
selection of quantitative traits may be required for S. hispanica. Information
collected in Pachali supports this idea in that S. hispanica is not weedy within or
at the edges of that agricultural setting and does not persist without attentive
human care.
The ethnobotany of Salvia hispanica can be viewed as a hypothetical
framework for the process of domestication that can be tested for all the
domesticated seed crops in Lamiaceae. Through ethnobotanical studies of wild
ancestors and primitive forms of related species, ethnobotany will continue to
provide answers, as it has in this study, to questions related to the genesis of
plant-people interactions.

NOTES
1All voucher herbarium specimens cited herein are deposited in the herbarium of the
University of California, Riverside.

2Interview with Dr. Juan Jimenez-Osornio professor at Universidad Autónoma de

Yucatán, in Riverside, California, September 18, 2002.

ACKNOWLEDGMENTS

I wish foremost to thank Dr. Arturo Gómez-Pompa for five valuable years of guidance
and assistance during my graduate studies. I also thank J. Giles Waines, Eugene Anderson,
and Blanca Hernández Bautista. I am indebted to the farmers and native people in the
areas of study for their willingness and excitement in relating information.

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