Comparing Wild and Cultivated Food Plant Richness Between the Arid American and the Mesoamerican Centers of Diversity, as Means to Advance Indigenous Food Sovereignty in the Face of Climate Change

Abstract

Climate change is aggravating agricultural crop failures, and the paucity of wild food harvests for Indigenous desert dwellers in Mexico and the U.S. This food production crisis challenges ongoing efforts by Indigenous communities in obtaining greater food security, prompting them to reconsider the value of traditional Indigenous food systems in both Mesoamerica and Arid America, two adjacent centers of crop diversity. While food production strategies in these two centers share many features, the food plant diversity in the Western Mesoamerican region appears to be greater. However, a higher percentage of plants in Arid America have adapted to water scarcity, heat, and damaging radiation. The phytochemical and physiological adaptations of the food plants to abiotic stresses in arid environments offer a modicum of resilience in the face of aggravated climate uncertainties. By comparing food plant genera comprising Western Mesoamerican and Arid American diets, we detected a higher ratio of CAM succulents in the wild and domesticated food plant species in the Arid American food system. We conclude that food plant diversity in the ancestral diets of both centers can provide much of the resilience needed to advance Indigenous food sovereignty and assure food security as climate change advances.

Full text

ORIGINAL RESEARCH
published: 20 June 2022
doi: 10.3389/fsufs.2022.840619
Frontiers in Sustainable Food Systems | www.frontiersin.org 1June 2022 | Volume 6 | Article 840619
Edited by:
Zakaria Kehel,
International Center for Agricultural
Research in the Dry Areas
(ICARDA), Morocco
Reviewed by:
Peter J. Jacques,
University of Central Florida,
United States
Daniel Pinero,
National Autonomous University of
Mexico, Mexico
*Correspondence:
Gary Paul Nabhan
gpnabhan@arizona.edu
Specialty section:
This article was submitted to
Crop Biology and Sustainability,
a section of the journal
Frontiers in Sustainable Food Systems
Received: 21 December 2021
Accepted: 29 April 2022
Published: 20 June 2022
Citation:
Nabhan GP, Colunga-GarcíaMarín P
and Zizumbo-Villarreal D (2022)
Comparing Wild and Cultivated Food
Plant Richness Between the Arid
American and the Mesoamerican
Centers of Diversity, as Means to
Advance Indigenous Food Sovereignty
in the Face of Climate Change.
Front. Sustain. Food Syst. 6:840619.
doi: 10.3389/fsufs.2022.840619
Comparing Wild and Cultivated Food
Plant Richness Between the Arid
American and the Mesoamerican
Centers of Diversity, as Means to
Advance Indigenous Food
Sovereignty in the Face of Climate
Change
Gary Paul Nabhan 1
*, Patricia Colunga-GarcíaMarín 2and Daniel Zizumbo-Villarreal2
1Desert Laboratory of Tumamoc Hill and Southwest Center, University of Arizona, Tucson, AZ, United States, 2Retired,
Mérida, México
Climate change is aggravating agricultural crop failures, and the paucity of wild food
harvests for Indigenous desert dwellers in Mexico and the U.S. This food production
crisis challenges ongoing efforts by Indigenous communities in obtaining greater food
security, prompting them to reconsider the value of traditional Indigenous food systems
in both Mesoamerica and Arid America, two adjacent centers of crop diversity. While food
production strategies in these two centers share many features, the food plant diversity in
the Western Mesoamerican region appears to be greater. However, a higher percentage
of plants in Arid America have adapted to water scarcity, heat, and damaging radiation.
The phytochemical and physiological adaptations of the food plants to abiotic stresses
in arid environments offer a modicum of resilience in the face of aggravated climate
uncertainties. By comparing food plant genera comprising Western Mesoamerican and
Arid American diets, we detected a higher ratio of CAM succulents in the wild and
domesticated food plant species in the Arid American food system. We conclude that
food plant diversity in the ancestral diets of both centers can provide much of the
resilience needed to advance Indigenous food sovereignty and assure food security as
climate change advances.
Keywords: Arid America, climate change, desert agriculture, diabetes, Indigenous food sovereignty,Mesoamerica,
traditional food systems, centers of biocultural diversity
INTRODUCTION
There is the smell of danger in the dry air: A recent analysis summarizing the U.S. Environmental
Protection Agency’s “Climate Change Impacts and Risk Analysis” concluded that by 2090, climate-
induced impacts on agriculture and 21 other natural resource-based sectors of the economy could
cost over $224 billion more per year to the U.S. economy alone, with impacts in Mexico and Canada
approaching similar levels (Martinech, 2018; Nuchatelli, 2019).

Nabhan et al. Mesoamerican and Arid American Food Richness
It is likely that because of water scarcity, increased heat,
and damaging solar radiation during flowering and fruiting—in
addition to land degradation—the yields of temperate-adapted
grains, vegetables and fruits will decline enough to pose food
security risks for most Indigenous and campesino farmers in the
Americas (Altieri and Nicholls, 2009; Nabhan, 2013, 2020). This
may be due to herbaceous crops like broccoli, common beans,
cowpeas, groundnuts, maize, rice, sorghum, tomato and wheat
hitting temperature thresholds near flowering times (Challinor
et al., 2005; Luo, 2011). In addition, climate catastrophes may
disrupt or set back the valiant efforts by Indigenous communities
to regain or retain food sovereignty (Peña et al., 2017; Mihesuah
and Hoover, 2019). We endorse and endeavor to advance the
definition of food sovereignty as stated in the 2007 Declaration of
Nyéléni, released at the first global forum on food sovereignty in
Mali (https://nyeleni.org/spip.php?article290) [accessed Feb 21,
2022]:
“Food sovereignty is the right of peoples to healthy and
culturally appropriate food produced through ecologically sound
and sustainable methods, and their right to define their food
and agriculture systems. It puts the aspirations and needs of
those who produce, distribute and consume food at the heart of
food systems and policies rather than the demands of markets
and corporations.”
At the same time, we conceded Indigenous farmers on
several continents are now among the ranks of those being
called “climate refugees” (Li et al., 2021), and that even their
basic food security—their reliable access to a sufficient quantity
of affordable, nutritious food—has been placed at further risk
by climate change, neo-liberal economic globalization and the
Covid-19 pandemic.
Overall, the current rates of global climate change are gravely
impacting food production and security while increasing health
risks for all residents of North America. Additionally, climate
catastrophes and carbon emissions have generated more than
$820 billion per year in additional physical and mental health care
costs to U.S. and Mexican farm-based food system economies
(Crimmins et al., 2016; Ebi and Hess, 2020; De Alwis and Limaye,
2021).
We are especially concerned by these climatic challenges
to Indigenous communities in deserts, along coastlines or in
mountainous regions who are being differentially affected by
climate catastrophes and sea level rising. It is important to
note the tremendous strides Indigenous peoples in the U.S. and
Mexico have made in their social movements in affirming food
sovereignty to gain further control over their dietary options in a
globalized food system (Peña et al., 2017; Mihesuah and Hoover,
2019). We do not wish to see such initiatives compromised or
disrupted by climatic stresses or by economic pressures.
Nevertheless, the combination of the Covid pandemic, high
unemployment, rising gas prices, climate change and accelerating
globalization pressures on Indigenous food systems have reduced
dietary diversity in Indigenous pueblos at a critical time when
such a variety of nutritional sources is desperately needed.
One tangible example was recently documented (Nabhan and
Molina, 2021) during the 2022 phase of the Covid-19 pandemic,
which was also one of the hottest, driest years suffered in recent
memory by Indigenous communities in coastal Sonora, Mexico.
The Comcaac, or Seri communities of coastal Sonora, Mexico,
historically used between 95 and 100 wild native plants for fresh
foods and beverages. However, only 12 of those species continue
to be routinely harvested for food by the 1,500 inhabitants of
two fishing villages today, and by far less than half of their
population. Instead, they rely on an extremely limited inventory
of domesticated crops such as fresh fruits and vegetables that may
be found in just seven local grocery stores.
The Comcaac diet has lost 57% of the plant species richness
found in their historic diet. This includes considering all
purchased fruits and vegetables as well as the occasional wild
foraging of native fruits, seeds, tubers and shoots. Some plant
populations are no longer accessible due to the loss of aboriginal
lands, while others have been destroyed altogether by land
clearing. Additionally, other plant species have fallen out of
use due to the loss of traditions, acculturation and shifts in
livelihoods. Similar trends have occurred in other Indigenous
villages. That is why we are assisting them in a three-year
program where they discuss their aspirations for enhancing food
sovereignty, while we provide technical assistance to help achieve
food, water, and energy security.
The possibility that Indigenous cultures’ own ethnobotanical
resources can be used as an incentive for renewed agro-ecological
development and plant domestication was first proposed in
Mesoamerica by Colunga-GarcíaMarín and Zizumbo-Villarreal
(1993),Colunga-GarcíaMarín et al. (2007) and Zizumbo-
Villarreal et al. (2012, 2016). It prompted Mexico’s activist-
scholars to initiate thoughtful, thorough, and far-reaching
efforts to revive the original structure and composition of the
Mesoamerican farming and food plant gathering in ways that will
strengthen Indigenous food sovereignty initiatives (e.g., Calvo
and Rueda-Esquibel, 2015).
Similarly, Indigenous populations in the U.S. Desert
Southwest have self-initiated several programs to safeguard,
produce and prepare “native foods” from their own farming
and wild foraging traditions to deal with food security and
sovereignty issues (Edaakie and Enote, 1999; Tohono O’odham
Community Action., 2010; Mihesuah and Hoover, 2019). This
native farming and foraging revival began within Indigenous
communities well before climate change was recognized as a
stressor. Even then, it was clear that food security interventions
and cultural food sovereignty affirmations could be effective
in dealing with several historic issues that were both culturally
devastating and economically burdensome (Kuhnlein and
Receveur, 1996; Minnis, 2021).
Our objectives for this reappraisal and revival of healthful,
plant-based food systems that emerged in the pre-colonial era in
what is now Mexico and the Southwestern U.S. are that: (a) they
can reduce crop failures from climate-related water scarcity and
heat waves, and that (b) they may help restore food sovereignty
to Indigenous and campesino communities in ways that reinforce
their cultural identity, and reaffirm their rights to land, water,
seeds and local food processing options (Peña et al., 2017).
This issue is important for the future as most of Mexico’s
population now dwells in hot, dry climates, where the arid food-
producing landscapes dominate 60% of the national territory
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Nabhan et al. Mesoamerican and Arid American Food Richness
(Appendini and Liverman, 1994; Pontifes et al., 2018). Across
much of Latin America, temperature thresholds and drought
are beginning to limit the production of most maize and bean
varieties (Rodríguez-De Luque et al., 2016; Stiller et al., 2021).
Similar trends now occur in the Western U.S. (Nabhan, 2013).
We therefore wish to compare, contrast and, if possible, revive
elements of food systems in what we call the Mesoamerican
and Arid American centers of diversity. Our ultimate goal for
this article is to detail how restoring the broad diversity of
wild and cultivated plants once found in the ancestral diets
of these areas, originated in the Archaic or preceramic period
(Zizumbo-Villarreal et al., 2014, 2016) may enhance human
health, economic wellbeing, food security and resilience in the
face of climate change.
GEOGRAPHIC CONTEXT, MATERIALS AND
METHODS
For the purposes of this article, we compared two subareas of
the centers of diversity known as Mesoamerica and Aridoamerica
(Figure 1), two terms coined by Kirchhoff (2000) to describe the
two cultural areas of what is now central Mexico and part of
Central America, and Northern Mexico and the Southwestern
U.S. This Mesoamerican ethno-ecological region refers to
the megadiverse landscapes extending from central Mexico,
Guatemala, El Salvador, Honduras, Nicaragua, and part of Costa
Rica, where multiple cultures, languages and foodways flourished
in pre-Colonial eras, elements of which continue to persist today.
Here, we focus more narrowly on Western Mesoamerica along
the Pacific coast of Mexico to compare with the western reaches
of a considerably more arid region to its north.
The use of the term Aridoamerica is much more common
among Mexican scholars than among American ones. In this
paper, we have integrated Kirchoff ’s Oasisamerica (Kirchhoff,
1954) — with its nucleated set of irrigated agricultural
communities into the larger matrix of “seasonally dry-farmed”
(de temporal) communities of the cultural area which he
called Aridoamerica. To distinguish the units of Kirchhoff’s
Oasisamerica and Aridoamerica from our re-defined secondary
center of crop diversity, we use the new delineation of
“Arid America”. Our new geographic delineations of these
two centers were elaborated by our colleagues in Mexico
and the U.S. who contributed to an agroecological research
article complementary to this one (Nabhan et al., 2020),
based in Nabhan (1985). It integrates floristic, vegetational,
agroecological, ethnobotanical, anthropological and linguistic
factors first elaborated by Hernández-Xolocotzi (2013) to find the
most parsimonious fit of the boundaries of each region.
As defined here, both the Mesoamerican and Arid American
centers of crop diversity fall primarily within the larger
Neotropical phytogeographic region which biogeographer
Rzedowski considered to be “MegaMexico” that extended
into the U.S. Desert Southwest (Rzedowski, 1978). The Arid
American area is a secondary, binational center of crop diversity
that spans the Sonoran Desert, including Baja California’s deserts;
and to the east, the higher elevation Chihuahuan Desert, which
includes the Zacatecas-Potosí Desert. Its biocultural food system
has featured more floristic and cultural diversity than most other
desert regions in the Americas (Luque-Agraz et al., 2016).
To develop a full characterization of the Arid American center
of diversity, we have amplified and corrected Nabhan (1985)
initial inventory of domesticated crop plants to use as a point of
departure for the first published comparison of the archaic diets
of Mesoamerica and Arid America (Supplementary Table S1).
First, we determined what annual and perennial crops
dominated agriculture in each center during the pre-Colonial
era. Next, we documented which wild native plant species likely
preceded, and complemented or underpinned this domesticated
crop inventory from the late pre-Invasion era, through historic
“colonial” eras, rounding out the ancestral diets of “Indigenous”
or “Native American” communities of Arid America and
Western Mesoamerica.
To do so, we have drawn upon two ethnobotanies from each
of three Arid America subregions:
(1) Baja California, Aschmann (1959) for the Cochimí of the
central desert and Wilken-Robertson (2018) for the Kumeyaay
of the semi-arid foothills; (2) the mainland’s Sonoran Desert,
including Rea (1997) for the Upper Pima of the Arizona Uplands
and Felger and Moser (1985) for the hyper-Arid Gulf Coast
of Sonora; and (3) the Chihuahuan Desert, including Latorre
and Latorre (1977) for the Kickapoo of Coahuila and Texas, as
well as Solano-Picazo (2018) for the Wixarika (Huichol) of San
Luis Potosí. These have been supplemented by regional classics
such as Hodgson (2001) Food Plants of the Sonoran Desert and
Hernández-Sandoval et al. (1991) Plantas Utiles de Tamaulipas.
These six references yield a rather comprehensive view of the wild
or semi-cultivated food plants whose presence in these regions
predate Spanish Invasion, with a focus on nutritionally significant
genera that continue to be consumed by two or more Indigenous
cultures across the entire Arid American region.
Next, we will analyze which of these wild food plants
were crop relatives or congeners of the domesticated species
listed in the Supplementary Table S1 inventory. Our primary
hypothesis is that well before fully domesticated plants entered
their traditional cuisines, Indigenous communities were already
familiar with and gastronomically utilized several crop wild
relatives in their archaic diets (Zizumbo-Villarreal et al.,
2012; Contreras-Toledo et al., 2018; Riordan and Nabhan,
2019). Our secondary hypothesis is that these crop wild
relatives have characteristics that may make them ideal
food crops in a climate-changed world. Our reason for
focusing more on the wild food plant species rather than
on the Indigenous domesticated seeds crops of Western
Mesoamerica and Arid America is to minimize the risk of
cultural appropriation and legal enclosure of the particular
seed stocks that fall under the “farmers’ right” of specific
Indigenous communities. By and large, the rejuvenation of
the uses of widespread wild food plants does not pose
such a risk of violating the food sovereignty of any single
Indigenous community.
To compare the relative richness of crop wild relatives in
a local flora of Western Mesoamerica with one from Western
Arid America, we have selected the flora of the Sierra de
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Nabhan et al. Mesoamerican and Arid American Food Richness
FIGURE 1 | Aridamerican and Mesoamerican Centers of Diversity. Modified from Kirchhoff (1954, 2000),Nabhan (1985), and Nabhan et al. (2020).
Manantlán Biosphere Reserve on the Jalisco-Colima border
(Vásquez-García, 1995) and the flora of Cañón de Nacapule in the
Sierra de Aguaje, part of the Cajón del Diablo Biosphere Reserve
in coastal Sonora, Mexico (Felger et al., 2017).
To further refine our retrodiction of the archaic diets in
each center, we give particular attention to those documented
among cultures of Arizona, Baja California and Sonora to
represent the Arid American food systems, and those of Jalisco
and Colima to represent the Western Mesoamerican food
systems. Because early ethnographies typically lack scientific
names for the wild plants included in them, we cross-
referenced their regional and Indigenous folk names with
updated scientific nomenclature from Martínez (1979),Inés-
Olaya (1991),Moerman (1998),Hodgson (2001), and Avitia-
García and Castillo-González (2002). We included domesticated
foods recorded in compendia such as Burns et al. (2000),
Dunmire (2004),Zizumbo-Villarreal and Colunga-GarcíaMarín
(2010), and those in the Supplementary Table S1 derived from
Nabhan (1985).
RESULTS
Characterizing the Annual Food Crop
Biodiversity of Arid American and Western
Mesoamerican Food Systems
Of some thirty species of annual or annualized perennials
domesticated as crops in the pre-Colonial era, each of the
two centers of crop diversity harbors roughly the same
number of these short cycle, warm-season food staples
(Supplementary Table S2): 19 species for Mesoamerica and 2O
for Arid America. It is abundantly clear that during pre-Invasion
eras, Mesoamerican cultures domesticated far more annual food
crops plants than did Arid American cultures. The mix of plant
families in these two sets of regional annual crops is much the
same: composites, cucurbits, grasses, legumes, as well as pseudo-
cereals from the amaranth family dominate the mixes.
Many of the Arid American cultivated plants were rigorously
selected by both weather and culture to be short-cycle crops,
maturing with dry seeds in as little as 36 to 55 days during the
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Nabhan et al. Mesoamerican and Arid American Food Richness
monsoon season of mid-summer to early fall, to “escape” drought
rather than to endure it. These crops mimic summer “ephemeral”
wildflowers in the Sonoran and Chihuahuan deserts in that they
quickly germinate, flower, set fruit and die before the late autumn
drought period sets in. Since most of these crops utilize the
C3 metabolic pathway, they require considerable soil moisture
each week they are alive but reduce cumulative consumptive
water use by maturing quickly. In contrast, most annual West
Mesoamerican crops are facultative perennials, which persist in
gardens and fields for at least seven months during the warm
season, but may persist and set fruit for 12 to 15 months before
senescence. Often, root fungi, other diseases, and pests terminate
their growth, not climatic constraints.
Over the last four millennia, no fewer than 22 domesticated
annual food crops were culturally dispersed from Mesoamerican
food systems into Arid American food systems (Burns et al., 2000;
Dunmire, 2004). Most of these require much more irrigation than
the desert-adapted food crops like tepary beans, jack beans, little
barley, and sagui or Sonoran panicgrass (Nabhan and de Wet,
1984).
Arid American cultures obtained many of their staple foods
from the south via group-to-group diffusion across a Uto-
Nahua linguistic continuum (Merrill et al., 2009). Probably, most
domesticated annuals or annualized perennials grown for food
in Arid America of the last several millennia began to diffuse
into the more northern, arid region along the Western Mexican
coastal trade routes beginning between 6000 and 5500 calibrated
years before present (Merrill et al., 2009; Mabry1). The Las
Capas site in the Western Tucson Basin has yielded three direct
radiocarbon dates on maize remains between 5,700 and 4,500
calibrated years before present [Vint, 2015, 2018; Mabry (see
text footnote 1)]. These are currently the oldest domesticated
crop dates in the Southwest U.S./Northwest Mexico region, but
archaeologists anticipate that early dates for perennial crops
will eventually be reported as new genetic and archaeological
methodologies are utilized in the region.
In a few cases, we can posit that because all Mesoamerican
food crops adapted to the wet Neotropics did not grow well in the
hot, dry lowlands of Arid America, another, more desert-adapted
set of congeners of similar utility were utilized. For example, the
domestication of tepary beans in Arid America appeared well
after common beans, lima and runner beans were domesticated
in Mesoamerica (Ford, 1981). We might hypothesize similar
processes of “relay domestication” into more northerly, arid
climes with Cucurbita, Solanum, Physalis, Jaltomata, among
other annual crop genera were recruited to play a similar
role in diets as their tropical counterparts (see Rodríguez and
Spooner, 1997; Louderback and Pavlik, 2018). The broad climatic
differences between the centers of diversity also influenced the
prevailing plant chemical defenses in the sets of annual crops
dominating each of the two cultural regions, a topic which we
will address elsewhere.
These New World domesticated annuals have remained
among the most important warm-season food crops in both
biocultural food systems, but recent severe heat waves, prolonged
1Mabry, J. (2022). Southwest Center, University of Arizona maize archaeologist,
October 5 2022. (personal communication).
drought and water scarcity associated with climate change are
now impacting them in several ways, such as extremely high
summer temperatures causing abortion of flowers and fruits
(Altieri and Nicholls, 2009; Nabhan, 2013). This is disconcerting,
since they have provided much of the calories and complex
carbohydrates to their Indigenous communities for the last three
millennia. In many ways, they have provided a now imperiled
structural matrix, or “backbone”, of the food systems involving
most cultures in Mesoamerica and Arid America up until the last
several decades.
Characterizing the Perennial Food Crop
Biodiversity of Historic Arid American and
West Mesoamerican Food Systems
The domestication of some perennial food plants was once
more difficult to discern than that of annual crops, but recent
methodological advances have revealed many more domesticated
perennials in North America than previously recognized
(Colunga-GarcíaMarín et al., 1986; Hernández-Xolocotzi, 1993;
Casas et al., 1999, 2002). Of 43 perennial food crop species, once
found between the two cultural regions since pre-Invasion eras,
34 of these perennial crop species continue to be found in West
Mesoamerica, while 26 perennial crop species continue to be
found in Arid America (Supplementary Table S3). In general,
there appears to be no sharp break, but is rather a genetic
continuum from wild to domesticated in most of these perennial
crops of Aridamerica and Mesoamerica than in the perennial
crops from Eurasia and North Africa that were introduced during
the Colonial era (Ezcurra2).
A semi-cultivated perennial, Palmer’s saltgrass (Distichlis
palmeri) of the Colorado River delta, fell out of management in
Arid America but has since been revived (Yensen, 2008), while
another cereal, foxtail millet (Setaria parviflora), has apparently
disappeared altogether as a crop from both cultural regions
(Callen, 1967; Austin, 2006).
Often ignored by early archaeologists seeking out the origins
of domesticated crops, these plants do not exhibit morphological
divergence from their wild ancestors as dramatically as annual
domesticates do [Colunga-GarcíaMarín and Zizumbo-Villarreal,
1993; Casas et al., 2002; Louderback and Pavlik, 2017; Ezcurra
(see text footnote 2)]. However, these resilient sets of perennial
crops in Arid America and Mesoamerica survive drought and
heat by deep roots tapping into deep soil moisture, by sloughing
off branches during drought, or by extended dormancy. They
may also sequester far more carbon in the soils of multi-cropped
milpas and agroforestry-based orchards than do annual crops
originating in either biocultural food system.
In contrast to the diverse perennial assemblage associated with
the Mesoamerican center of diversity and its milpa fields and
solar/huerta (dooryard garden) agroecosystems, only the food
systems at the southernmost edge of Arid America center retain
much diversity of perennial food crops. North of the Rio Soto La
Marina, Rio Conchos, Rio Mayo and Rio Yaqui there were very
few tree crops until Spanish introduction of Old-World fruits and
nuts in the Colonial era (Burns et al., 2000; Dunmire, 2004).
2Ezcurra, E. (2021). University of California Riverside Department of Botany and
Plant Sciences, Desert Ecologist, September 10, 2021. (personal communication).
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Nabhan et al. Mesoamerican and Arid American Food Richness
Arid American perennial crops were largely limited to
succulent plants utilizing the Crassulacean Acid Metabolism
(CAM) pathway because of the biotic and abiotic stresses on
fruit trees historically posed by highly variable, scarce rainfall
as well as challenging heat, transpiration rates, and pests. These
CAM succulents include agaves and cacti. Importantly, 76% of
Arid American perennial crops utilize the CAM pathway, while
only 66% of Western Mesoamerican crops use the CAM pathway.
The fruit crops of both trees and vines using the C3 pathway
are far more vulnerable to drought stress and crop failure, unless
frequently irrigated.
The water-conserving CAM of cacti and succulents such as
agaves allowed them to produce more edible biomass on less
moisture than needed by C3 and C4 crop species from the tropics,
including maize (Nobel, 2009). We predict that climate change
will increasingly constrain yields of fruit and nut trees that use the
C3 photosynthetic pathway and thereby suffer high transpiration
rates. In our view, many fruit tree orchards in arid and semi-
arid/subtropical landscapes will need to be replaced with CAM
succulents such as agaves, prickly pears and columnar cacti if
their farmers are to economically weather climate change.
In short, cultivators in the Arid American food system once
relied on a relatively greater species richness of domesticated
succulent crops, as opposed to trees and woody vines, in their
cultivation of perennial food plants than did cultivators in the
wetter Mesoamerican food system.
There are both agroecological and human health reasons for
reviving and extending the cultivation of these CAM succulent
crops in dry lands (Leach and Sobolik, 2010). First, the water-
conserving Crassulacean Acid Metabolism of cacti and succulents
such as agaves allowed them to produce equal tonnages of edible
biomass using just half to one-sixth of the moisture needed to
provide the same yields by C3 and C4 crop species from the
tropics, including maize (Nobel, 2009).
Several agaves were independently brought into cultivation
on well over 200,000 hectares in northwestern Arid America
food systems on the edges of the Sonoran Desert. They included
Agave delameteri, A. murpheyi, A. phillipsiana,A. sanpedroensis,
A. verdensis and A. yavapaiensis (Fish et al., 1985; Hodgson,
2012; Hodgson and Salywon, 2013; Hodgson et al., 2018), while
cultivation of a seventh species remains debated (Nabhan et al.,
2019).
Toward the southeastern edges of Arid American food
systems where several giant agaves used in the production
of fermented pulque beverages went through the initial
phases of their domestication process [Ezcurra (see text
footnote 2)], these species have long been cultivated on
hundreds of thousands of hectares. These magueyal plantations-
with giant agaves often “alley-cropped with annuals—have
included A. americana, A. angustifolia, A. mapisiga, A.
lophantha and A. salmiana.” These succulent species have long
been cultivated and culturally dispersed in the Chihuahuan
Desert-Altiplano ecotone as much as in Mesoamerica itself
(Gentry, 1982).
The same may be true with the prickly pear species and
varieties that dominate the extensively managed nopaleras in the
Chihuahuan Desert-Altiplano ecotone. Some appear to include
hybrid clones derived from multiple lineages, including Opuntia
leucotricha, O. megacantha, O. streptacantha and O. tomentosa
(Griffith, 2004). Aside from this baffling array of clones that
historically were all lumped into Opuntia ficus-indica, there
may additional domesticated prickly pears that were separately
domesticated in Arid America, such as Opuntia durangensis
and possibly Opuntia robusta in the fluctuating border of
Mesoamerica and Arid America (Colunga-GarcíaMarín et al.,
1986).
To summarize, cultivators in the Arid American secondary
center of diversity relied on a greater percentage of succulent
crops than cultivators in the primary center of Mesoamerica. We
hypothesize that the traditionally-processed foods and beverages
from these succulent crops of Arid America comprised a greater
portion of Indigenous diets than they did in the Mesoamerican
center (Leach and Sobolik, 2010). They may need to do so again.
Characterizing the Wild Food Plant
Biodiversity of Arid America and Western
Mesoamerica
In addition to domesticated plants in both Arid American
and Mesoamerican centers of crop diversity, Indigenous
communities in Arid America have continued to forage for a
significant number of food and beverage sources. These include
over 235 wild desert plant species from at least 125 genera and
60 families on monocots and dicots (Hernández-Sandoval et al.,
1991; Hodgson, 2001).
While it is beyond our current capabilities to make a
similar estimate of widely used food and beverage plants from
Mesoamerica, we are relatively certain that the ethnographically
documented inventory of wild foods species is more numerous
for Western Mesoamerica than for Western Arid America
(Mapes and Basurto, 2016). If only for its greater surface
area, rainfall, and floristic diversity, Western Mesoamerica most
likely has far more species, genera and families of food and
beverage plants.
It is likely that Supplementary Table S4 is a relatively modest
underestimate of the total number of the characteristic food and
beverage plants of historic Arid American food systems, since
we have not included all microendemics or famine foods—only
those known by two or more cultures in the region (Hernández-
Sandoval et al., 1991; Hodgson, 2001; Minnis, 2021).
While 225 food and beverage plants are certainly enough
to draw upon through local harvesting and intra-region trade,
there are some remarkable patterns evident within this Arid
American food system. For example, nearly a fourth, or 23%, of
the characteristic wild food and beverage plant species in Arid
America are succulents that utilize the CAM pathway for high
water use efficiency photosynthesis. We do not know of any
comparably high estimate for any other biocultural region in
the world.
Indigenous inhabitants of the Arid American center
historically drew upon a particularly high number of edible
species in the genera of Agave,Amaranthus, Atriplex,
Cylindropuntia, Echinocereus, Ferocactus, Opuntia, Physalis,
Quercus, Randia, and Salvia. That five of these ten genera are
CAM succulents and another two are drought hardy trees
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Nabhan et al. Mesoamerican and Arid American Food Richness
suggests that these food plants may help form a basis for climate-
resilient food security in the future (Nabhan et al., 2020). As we
will document in later discussions, these genera are particularly
rich in bio-active compounds that can potentially reduce diseases
and maladies of oxidative stress. In addition, the plants from
five succulent genera have also been utilized to elaborate the
probiotic beverages we and our colleagues have discussed in
other papers (e.g., Ojeda-Linares et al., 2021).
In general, Supplementary Table S4 makes it abundantly
clear that Arid American food systems before the Spanish
Invasion may not have been as rich in wild species as
Mesoamerican food systems. However, neither were they
impoverished, or lacking a variety of plants that provided a
diverse array of nutrients. This species richness of food plants
is evident in 28 genera of the crop wild relatives compiled for
Arid America and Northwestern Mesoamerica in another paper
of ours using the same data base (Nabhan et al., in press).
It determined that there are at least 28 genera and 43 wild
species of crop wild relatives that have been ethnographically
documented as food plants in Northwestern Mesoamerica,
compared to 21 genera and 48 wild species in Arid America.
There are clearly many species of crop wild relatives in each
region whose use as food probably predated the presence of
their domesticated congeners (Zizumbo-Villarreal and Colunga-
GarcíaMarín, 2010), which could again be utilized as foods in
the future (Contreras-Toledo et al., 2018; Riordan and Nabhan,
2019).
In short, certain domesticated species never really “replaced”
or “made obsolete” their wild congeners as foods, especially
during periods of drought or famine (Mapes and Basurto, 2016;
Minnis, 2021). To this day, the popularity of the wild foods such
as amaranth greens, chiltepín peppers, wild grapes and plums,
and prickly pear cactus fruits in Arid America has not waned.
Many of the crop wild relatives have the potential to provide more
yield stability under stressful climatic conditions than do their
domesticated congeners. They are also ideal to use as rootstock,
trap crops for pests in hedgerows, and as pollinator attractants in
orchards (Riordan and Nabhan, 2019).
Supplementary Table S5 compares the richness of crop wild
relatives in two local floras, each a representative “surrogate”
for its center of diversity. We compared two local floras to the
crop wild relatives lists in Contreras-Toledo et al. (2018) and
Riordan and Nabhan (2019). Of course the comparability of these
two local floras is not geographically optimal, but the lists are
taxonomically up to date, and comprehensive. One of the floras is
derived from the Sierra de Manantlán of Jalisco and Colima, not
far from the putative “cradle” of Mesoamerican domestication of
maize and beans in the Rio Balsas watershed (Vásquez-García,
1995, with updates on file at the biosphere reserve). The other
is derived from the Sierra El Aguaje of coastal Sonora, on the
Sonoran Desert ecotone with semi-arid subtropical thornscrub.
The Sierra del Aguaje lies within the 530,000 ha Guaymas region,
and harbors roughly 700 vascular plant species (Felger et al.,
2017). In contrast, the Sierra de Manantlán area—surrounding
the biosphere of the same name—covers less than a fourth of
the area of the Guaymas region, 140,000 ha, yet harbors at least
2,770 vascular plant species. That is nearly four times the species
richness of the desert region. As one might expect, the Sierra de
Manantlán in Mesoamerica is home to many more genera, 45, of
wild relatives than the number of genera represented in the Sierra
del Aguaje in Arid America, 17.
Remarkably, the Sierra de Manantlán conserves in situ over
330 species of crop wild relatives compared to the 30 species
in the Sierra del Aguaje reserve. We tentatively project that the
Mesoamerican biosphere reserve harbors ten times the number
of species than the Arid American biosphere reserve, even though
the latter is roughly four times larger in land area. These trends
suggest three patterns: (1) Mesoamerican farmers had far more
opportunities to recruit and domesticate food plant species from
local wild floras; (2) the constraints on those opportunities in
desert areas may have encouraged Arid American farmers to
actively seek out crops first domesticated in more tropical climes;
or (3) the Arid American cultivators sought to diversify the
number of locally adapted landraces of the few drought-tolerant
species they brought into cultivation that were derived from their
own regional flora.
DISCUSSION
As explicitly stated earlier, our goal is to detail similarities
and differences in the composition of the archaic diets of
Indigenous communities in two adjacent centers of crop
diversity, Northwestern Mesoamerica, and Arid America.
Understanding the differences in the diversity of wild and
domesticated food plants in these two centers may help
broaden our perspective on how to gain more food system
resilience in the face of climate change, especially for Indigenous
communities who have valiantly struggled to maintain their food
sovereignty options.
Our results echo the distinctions between a primary and
secondary center of crop diversity highlighted by many crop
scientists, including Kumar (2016). As a primary center,
Mesoamerica has greater domesticated crop diversity and greater
diversity of crop wild relatives used as foods, with a greater
species richness of wild food plants overall. As a secondary center,
Arid America has relative lower crop diversity, especially for
domesticated perennials and crop wild relatives. Nevertheless,
the diets of Arid America harbor a high number of drought-
and heat-adapted food plants, especially wild and domesticated
succulents like agaves and cacti, as well as many wild “famine
foods” still in use.
Our documentation and analysis indicate that currently
the ancestral diets of both Arid America and Northwestern
Mesoamerica have an undervalued diversity of food and beverage
plants that are already cultural acceptable and accessible to
many Indigenous communities. These plants also demonstrate
superb adaptations to the stresses of water scarcity, heat and
damaging solar radiation that will become more valuable to
human wellbeing as climate change proceeds. There is an
extremely high ratio of water-conserving succulent plants with
CAM photosynthetic pathways compared to plants utilizing
the C3 and C4 photosynthetic pathways found in archaic Arid
American diets which have fallen out of daily cultural use. In
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Nabhan et al. Mesoamerican and Arid American Food Richness
contrast, more of the considerable species richness in CAM
plants historically found in Mesoamerican diets remain utilized
in contemporary Mexican diets to this day.
Food systems scholars express concern that the many
technological advances in food harvesting, storage, processing
technologies and medical care now function as disincentives
for embracing or reviving the labor-intensive foraging, farming
and food processing practices of the past. We do not deny
that there remain many formidable social, ecological, and/or
economic disincentives that keep contemporary Indigenous
communities from fully reviving some of their traditions of plant
food procurement.
Nevertheless, there are now many agricultural, nutritional,
medical, and even spiritual or cultural reasons for doing so. The
possibilities for “rebirthing,” “reviving” or “restoring” these foods
and beverages derived from ancestral diets are not inevitably
beyond affordability in Indigenous communities today.
This is particularly evident in communities where
philanthropic or governmental subsidies for producing or
harvesting healthy (including “native”) foods are offered to
Indigenous communities in either Mexico or the U.S. For
example, the USDA Women, Infants and Children (WIC)
food program subsidizes the collection or propagation of
certain native foods, as does the Comisión Nacional Forestal
(CONAFOR) in Mexico (https://www.gob.mx/conafor/articulo
s/artesanias-sonorenses-herencia-de-los-seris?idiom=es; https:/
/www.conafor.gob.mx/EstudiosRegionalesForestales/EstudioRe
gionalForestal_UMAFOR0302_PSSG.pdf.; Segura-Aguilar et al.,
2020) [accessed Feb. 21, 2022].
As Colunga-GarcíaMarín et al. (2007) have proposed
elsewhere, for many Indigenous communities, their most
healthful and secure “future may be ancestral”. As Nabhan and
colleagues have proposed, “the cultivation of many of these
Indigenous foods of Arid America is needed in newly designed
or renovated agroforestry systems to address the emerging food
security and agricultural crises” due to climate change (Nabhan,
2020; Nabhan et al., 2020) triggered by climatic changes and
the pandemic. The diversity of these regionally adapted food
crops—when planted in perennial-dominated polycultures—
may restore land health, especially soil moisture holding capacity,
while reducing crop consumptive water use and providing yield
stability in the face of climatic uncertainty (Nabhan et al., 2020).
CONCLUSIONS
We have validated our primary hypothesis that prior to
when domesticated crop plants entered their food production
strategies, Indigenous communities were already familiar with
and gastronomically utilized numerous crop wild relatives that
could be used in their archaic diets. Our secondary hypothesis
is potentially valid, but remains to be fully tested, accepted,
or rejected: that many of these wild congeners of crops—as
opposed to domesticated crop seedstocks— have agroecological
and nutritional value for enhancing Indigenous food security
in a future hotter, drier world. Nervertheless, as the recent
documentary film Gather (sponsored by the First Nations
Development Institute) illustrates, there is already a growing
movement among Native Americans to utilize community-based
wild harvesting to reclaim their spiritual, political, and
cultural identities through food sovereignty (https://www.
nativefoodsystems.org/).
To advance their food plant options for integrating into
Indigenous food sovereignty initiatives to weather climate
change, we have assembled the first ethnographically
documented inventory of food plant diversity in two adjacent
centers of biocultural diversity in the Americas. We have
concluded that the following geographic patterns in food
systems emerged prior to the Spanish Invasion of Arid America
and Mesoamerica, and that many of their features are still
viable agroecological assets or food sovereignty strategies for
agriculture and wild plant foraging today:
1. Both centers still harbor a significant diversity of plant foods
with a wide variety of plant growth forms such as trees,
herbs, vines, succulents, and perennials. These food plants
employ a broad array of phytochemical and physiological
adaptations for producing food in hot, dry climates. These
adaptive strategies may become even more important to
Indigenous communities as they struggle to survive the hotter,
drier climates that humankind is increasingly facing with
global change.
2. While the Mesoamerican center is much more floristically
diverse, with greater crop diversity and species richness in
wild food plants than in Aridamerica, the latter region has an
unusually high percentage of endemic succulent food plants
which have been elaborated into prebiotic foods and probiotic
beverages for millennia. Renewed domestication, agricultural
production and processing of these healthful native foods and
beverages should be an explicit goal in redesigning the food
systems in both centers of crop diversity.
3. Given that many Indigenous communities have taken it upon
themselves to reintegrate these plants into their contemporary
diets as a strategy toward achieving greater food sovereignty,
agricultural scientists, educators and policymakers should
offer both technical and financial support whenever requested
to help them achieve their explicit desires and aspirations.
4. Far from food systems of “Mexican origin” being dominated
by maize, beans, and squashes (e.g., Calvo and Rueda-
Esquibel, 2015, who include these three crops in 57% of
their recipes), the ancestral diets in both centers have long
benefitted from employing an astonishingly broad diversity
of food plants. These benefits are no less relevant to
Indigenous food sovereignty today that they were historically.
Furthermore, most maize varieties, as well as many beans,
and cucurbits are hitting their temperature thresholds and
water deficit limits in most American landscapes (Altieri and
Nicholls, 2009; Nabhan, 2013). Clearly, food production and
diets in the “new climatic normal” will have to employ a set of
food crops far more diverse and different than those employed
in conventional agriculture at this moment in time (Nabhan,
2020).
Finally, we wish to emphasize that many of the initial efforts
to revive the original structure and composition of ancestral
Mesoamerican and Arid American diets have already emerged
from Indigenous communities themselves (e.g., Kavena, 1980;
Wolfe et al., 1985; Edaakie and Enote, 1999; Mihesuah, 2005;
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Nabhan et al. Mesoamerican and Arid American Food Richness
Tohono O’odham Community Action., 2010; Patchell and
Edwards, 2013; Peña et al., 2017; Mihesuah and Hoover, 2019).
Well before many of these articles or books were published by
authors of Indigenous heritage, their communities in Mexico and
the U.S. were quietly reintegrating traditional foods long a part of
Mesoamerican and Arid American diets into their contemporary
feasts and health care strategies. These community-based efforts
“go far beyond” the mere achievement of food security in terms
of reliable access to affordable, nutritious food.
As noted in the 2007 Declaration of Nyéléni, a broad multi-
cultural coalition wishes to place “the aspirations and needs of
those who produce, distribute and consume food at the heart of
food systems and policies rather than the demands of markets
and corporations.” We wish to support that intent by promoting
participatory research and on-ground technical assistance of the
kind we have begun with CONACYT support to advance climate-
friendly food, water and energy solutions in two Indigenous
Comcaac communities in costal Sonora, Mexico.
In essence, the revival of Mesoamerican and Arid American
agricultural and foraging traditions is not a top-down or
exogeneous pressure toward “reverse-engineering” dietary
change, but an Indigenous “grassroots community-based
movement toward true food sovereignty (Patchell and Edwards
2013; Peña et al., 2017).” Our research only validates many of the
tenets that underlie this Indigenous movement; it points to the
nutritional benefits of often-forgotten wild and cultivated food
plants. Conserving the wild plants, protecting the traditional
crop landraces, and safeguarding the traditional ecological,
gastronomic, and agricultural knowledge of Indigenous Nations
in Arid America and Mesoamerica will be as critical as
policy reforms to foster their food justice. As climate change
differentially threatens Indigenous communities in many ways,
advancing and safeguarding food sovereignty and the wild food
plants that may contribute to it will be paramount.
DATA AVAILABILITY STATEMENT
The original contributions presented in the study are included
in the article/Supplementary Material, further inquiries can be
directed to the corresponding author/s.
AUTHOR CONTRIBUTIONS
GN, PC-GM, and DZ-V conceived together the article in Merida,
Yucatan, Mexico and did multiple revisions. GN elaborated the
first draft. All authors contributed to the article and approved the
submitted version.
ACKNOWLEDGMENTS
We wish to thank our many Indigenous collaborators on
dietary studies over the last four decades, most recently Luis
Eduardo Molina of the Comcaac community of Sonora, who
assisted GN with establishing baselines for monitoring dietary
changes and advancing food sovereignty through time in two
Indigenous villages, and the families from Zapotitlán, Jalisco,
who shared their deep knowledge of their ancestral foodways
with PC-G and DZ-V. We also wish to acknowledge our
collaborators from many cultures including A. Mellado, E.
Barnett, M. L. Astorga, M. Estrella Astorga, D. Lewis, J. Ascencio,
F. Kabotie, H. Dukepoo, L. Noriega, J. Martínez-Castillo, O.
Vargas-Ponce, G. Carrillo-Galván, A. Flores-Silva, I. Torres-
García, A. Casas, C.J. Figueredo, S. Rangel-Landa, A. Delgado,
D. Cabrera-Toledo; X. Aguirre-Dugua, L. Eguiarte, R. Bye, E.
Linares, D, A. Mellado, E. Barnett, R. Felger, C. Marlett, E.
Ezcurra, E. Riordan, A. Búrquez-Montijo, L. Smith Monti, E.
Riordan, B. Wilder, J. Mabry, C. Khoury, T. Crews, and J.
Aronson. Barbara Kuhns graciously assisted with edits of the
near-final draft. GN acknowledges long-term funding from the
W.K. Kellogg Foundation, and initial funding many decades
ago on three grants from the National Science Foundation. GN
acknowledges the W.K. Kellogg Foundation Endowment for his
position at the Southwest Center which covered page charges as
well as travel to Arizona for PC-GM and DZ-V, and to Yucatán
for GN.
SUPPLEMENTARY MATERIAL
The Supplementary Material for this article can be found
online at: https://www.frontiersin.org/articles/10.3389/fsufs.
2022.840619/full#supplementary-material
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