Native Crop Diversity in northwest. Mexico and the u.s. Southwest as resources of same cohesive ecological and cultural region. Signifcant crop / weed introgression continues where indigenous agriculture persists, but native fields are being. Rapidly abandoned or converted.
Native Crop Diversity in northwest. Mexico and the u.s. Southwest as resources of same cohesive ecological and cultural region. Signifcant crop / weed introgression continues where indigenous agriculture persists, but native fields are being. Rapidly abandoned or converted.
Native Crop Diversity in northwest. Mexico and the u.s. Southwest as resources of same cohesive ecological and cultural region. Signifcant crop / weed introgression continues where indigenous agriculture persists, but native fields are being. Rapidly abandoned or converted.
Native Crop Diversity in Aridoamerica: Conservation of Regional Gene Pools
Author(s): Gary Paul Nabhan
Source: Economic Botany, Vol. 39, No. 4 (Oct. - Dec., 1985), pp. 387-399 Published by: Springer on behalf of New York Botanical Garden Press Stable URL: http://www.jstor.org/stable/4254790 . Accessed: 01/03/2014 15:04 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org. . New York Botanical Garden Press and Springer are collaborating with JSTOR to digitize, preserve and extend access to Economic Botany. http://www.jstor.org This content downloaded from 132.248.181.31 on Sat, 1 Mar 2014 15:04:38 PM All use subject to JSTOR Terms and Conditions Native Crop Diversity in Aridoamerica: Conservation of Regional Gene Pools' GARY PAUL NABHAN 2 Scholars have seldom considered the native crop diversity in northwest Mexico and the U.S. Southwest as resources of the same cohesive ecological and cultural region. The term Aridoamerica is introduced to describe this overlooked center of plant domestication and diversification, which is distinct from centers of Meso- america and the Mississippi Valley. To understand why certain of its landraces are unique, the systematic relationships and gene-pool relations of crops found prehistorically and protohi storically in Aridoamerica are reviewed. Signifcant crop/ weed introgression continues where indigenous agriculture persists, but native fields are being rapidly abandoned or converted. In planning in situ and ex situ conser- vation efforts to maintain this diversity, both cultural factors and plant population genetics must be considered. Native crops are defined here as domesticated plants cultivated prehistorically or protohistorically within a region by its indigenous cultures since prehistoric or protohistoric times. Although they need not be endemic to the region of concern, their landraces should have been grown long enough in the region to exhibit morphological or physiological adaptations to the soils and climates found there. The landraces may in fact be key ecological components of the distinctive agroeco- systems that native farmers have developed, given the climatic and edaphic con- straints in their area (Hernandez X., 198 1). Native crops are directly dependent upon management by humans; therefore, they have evolved in part under the influence of farming practices of particular cultures. As such, native crop diversity directly reflects a region's cultural diversity. It is awkward to view these resources merely as a set of genes that can be conserved simply by depositing them in a gene bank. If isolated from the folk science and traditional uses of the cultures that have nurtured them, they lose their historical cultural context. If isolated from cultural selection and natural selection exerted in their endemic agroecosystems where they have long evolved, their subsequent evolution may change in direction. If removed from native fields where introgression with wild relatives has continued for centuries, other genetic changes will occur. Therefore, in situ and ex situ conservation of native crops are much more complex than conservation of wild genetic resources. In the U.S. Southwest and northwestern Mexico, much of the land is arid. Indigenous agriculture persists there, in some places beyond where conventional modern agriculture is successful. In addition to the reason usually given for genetic conservation -to preserve for future generations genes that may make commercial IReceived 7 January 1985; accepted 17 May 1985. Presented at the Symposium on Ethnobotany of the Greater Southwest at the Twenty-fifth Annual Meeting of the Society for Economic Botany, Texas A&M University, College Station, TX, 11-13 June 1984; symposium organized and chaired by Dr. Robert A. Bye, Jr. 2 Office of Arid Lands Studies, University of Arizona, 845 N. Park Ave., Tucson, AZ 85721; and Native Seeds/SEARCH, 3950 W. New York Dr., Tucson, AZ 85745. Economic Botany, 39(4), 1985, pp. 387-399
1985, by the New York Botanical Garden, Bronx, NY 10458 This content downloaded from 132.248.181.31 on Sat, 1 Mar 2014 15:04:38 PM All use subject to JSTOR Terms and Conditions ECONOMIC BOTANY crop varieties less vulnerable to stresses and maladies-there are others worth considering with regard to native crops of this binational region: 1. Native crops may be critical to continued food production by indigenous cultures. This is particularly true for the most marginal lands, where such crops have superior adaptations to local peculiarities of climate and soil (Toledo et al., 1981). Within southwestern North America, certain crop landraces oftepary beans and cushaw squash outproduce commercial cultivars of related species. These ecotypes tolerate low soil moisture in areas receiving less than 70 mm seasonal rainfall, air temperatures reaching 49C and soils with high pH, and/or soluble salts content as high as 1,800 ppm (Nabhan, 1983; Nabhan et al., 1985). 2. Certain crops, though not necessarily high yielders, may be important in efficient utilization and conservation of chronically scarce resources such as water and nitrogen (Romney et al., 1978). Water, in particular, is in such short supply that it is subject to profound intra- and intercultural conflicts (US OTA, 1984). 3. Even if introduced and improved varieties yield better, there may be non- economic motives for honoring an "obsolete" native crop's right to exist (Ehren- feld, 1977). For example, native crops and special foods derived from them may be symbols of cultural identity, and, as such, may reinforce an indigenous com- munity's pride and persistence (Spicer, 1971). 4. Native crops may cumulatively provide a different and perhaps superior set of nutritional resources to indigenous communities than may be obtained through government food welfare programs or through trading posts (Calloway et al., 1974; Nabhan et al., 1985). Native Americans and "Mexican-Americans" currently suffer from high incidences of diabetes and other nutrition-related diseases (West, 1974). It may be that fiber-rich foods formerly more prevalent in their diets-a diversity of beans (Phaseolus), mucilaginous seeds such as chia (Salvia), conivari (Hyptis) and cacti (Opuntia)-served to flatten postprandial blood sucrose curves in the same manner that artificial insulin is used today, thereby reducing the side effects of the adult-onset diabetes syndrome (Ramos, 1980; Leeds, 1981). Given these reasons for conserving native crop resources in southwestern North America, several questions must be raised. What native crops were once found in the region? How are they geographically and culturally distributed? Which gene pools are diminished and in need of intervention before further depletion? What combination of in situ and ex situ measures will best maintain remaining genetic variation? ARIDOAMERICA: AN OVERLOOKED CENTER OF DIVERSITY Table 1 lists the native crops found in indigenous communities in just 4 states in the U.S./Mexico borderlands: Arizona, New Mexico, Sonora, and Chihuahua. As can be seen, at least 25 plant species in advanced stages of domestication have been cultivated in these states prehistorically or protohistorically. There is in- triguing evidence that a number of additional species were cultivated and/or genetically selected within the area of these states (Table 2), but the degree of their domestication remains unclear. The point is not that we should inventory domesticates state-by-state. On the contrary, it can be demonstrated that most of these crops' distributions ignore such political boundaries and are shared by cultures on both sides of the current 388 [VOL. 39 This content downloaded from 132.248.181.31 on Sat, 1 Mar 2014 15:04:38 PM All use subject to JSTOR Terms and Conditions NABHAN: NATIVE CROP DIVERSITY TABLE 1. DOMESTICATED PLANT SPECIES OF ARIDOAMERICA: PREHISTORIC AND HISTORIC GEOGRAPHY.a Meso- Miss. Crop species AZ NM CHIH. SON. amer. Valley Amaranthus cruentus* 0 0 x x 0 A. hypochondriacus* 0 x x x 0 Canavalia ensiformis* 0 Capsicum annuum* x x x x 0 Chenopodium berlandieri* x? x x 0 0 Cucurbita ficifolia x x x C. mixta* 0 0 x x 0 C. moschata* 0 0 x x 0 x C. pepo 0 0 0 x 0 0 Gossypium hirsutum 0 0 0 0 0 x Helianthus annuus* x x x 0 Hordeum pusillum* Indigofera suffruticosa x x x Lagenaria siceraria 00 0 0 Nicotiana rustica 0 x x x 0 0 N. tabacum x x x x 0 Panicum sonorum* 0 x 0 Phaseolus acutifolius* 0 0 x 0 0 P. coccineus* x x 0 P. lunatus* 0 0 x x 0 P. polyanthus* x 0 P. vulgaris* 0 0 x 0 0 Physalis philadelphica* x x 0 Proboscidea parviflora* x Zea mays* 0 0 0 0 0 0 a Data derived from numerous published and unpublished records, available on request. Prehistoric period, indicated by a circle, refers to archaeological records predating 1492. Protohistoric period, indicated by an x, refers to archaeological and contact-time written documents, primarily from Jesuits or early explorers in Aridoamerica, postdating 1492. An asterisk behind the crop species binomial indicates that conspecific or cross-compatible congeneric wild plants are found within Aridoamerica. international border. Of course, no such border existed prehistorically. Its recent presence has hardly affected the distribution of native crops or of vegetation types within which their wild relatives are found (Fig. 1-3, based on vegetation types of Rzedowski, 1978). This point is belabored because most treatments of crop geography have in fact stopped at or near this border! At best, they consider the U.S. Southwest's farming traditions to be a crop and technology complex that invaded this "marginal agricultural area" as a package from Mesoamerica (Woodbury and Zubrow, 1979). George Carter's (1945) classic Plant Geography and Culture History in the Amer- ican Southwest dealt only with the U.S. Southwest, plus some 250 km2 of the Colorado River Delta in northwestern Mexico. Subsequent updates or summaries of distributions of native crop complexes by Winter (1974) and Ford (1981) have again dealt only with those tribes that are distributed north of the International Boundary. Dressler (1953) and others, in a similar manner, have dealt only with pre-Columbian cultivated plants of Mexico. Such limited views obscure the fact that this binational region is the hearth of endemic domesticates such as Panicum sonorum that have been cultivated in a 1985] 389 This content downloaded from 132.248.181.31 on Sat, 1 Mar 2014 15:04:38 PM All use subject to JSTOR Terms and Conditions ECONOMIC BOTANY TABLE 2. ADDITIONAL PLANT SPECIES THAT MAY HAVE BEEN CULTIVATED AND/OR CULTURALLY SELECTED IN ARIDOAMERICA.a Meso- Miss. Incipient domesticate AZ NM CHIH. SON. amer. Valley Allium sp. x Agave angustifolia x x x A. murpheyi ? x? Brassica campestris x x ? Cleome serrulata 0 x Dactyloctenium aegypticum x x Distichlis palmeri x x Hyptis suaveolens x x Jaltomata procumbens x x Nicotiana attenuata 0 N. trigonophylla 0 x Solanum jamesii x x I See footnote for Table 1. restricted area on both sides of the border but not in Mesoamerica or in the Mississippi Valley (Nabhan and de Wet, 1984). Whereas Ford (1981) and Doebley (1984) failed to recognize a truly domesticated form of this species with prehistoric and historic presence north of the border, others (Harlan, 1975) have erroneously assigned this plant to a Mesoamerican origin. Other crops such as tepary beans (Phaseolus acutifolius) and cushaw squash (Cucurbita mixta) may have noncentric origins stretching from Guatemala to Sonora, but have been considered strictly Mesoamerican in origin (Harlan, 1975). Most delimitations of a Mesoamerican center of crop origins extend from Mexico City or Durango southward to Honduras (Vavilov, 1951; Harlan, 1975). However, Zhukovsky (1975) and Zeven and de Wet (1982) define a Mexican and Central American region with northern limits exactly where the present day United States-Mexico boundary is situated (Fig. 1). It is remarkable that crop geographers would pretend that plants domesticated prehistorically were wise enough to anticipate where the Gadsden Purchase would finally place a political boundary in 1849! Notably, the term Mesoamerica is commonly used in another manner by Latin American geographers. It refers to a region with climate and vegetation that is predominantly tropical and mesic-intermediate between xeric and hydric in its access to water-that is located south of the extratropical dry lands and adjacent semiarid highlands of North America (Kirchhoff, 1954; West 1964). In an excellent but little-cited publication that is available in both Spanish and English, Dr. Jorge Leon (1979) details the distribution of Mesoamerican crop genetic resources. Leon notes that anthropologists place the northern limits of Mesoamerica near the watershed divide between the Rio Panuco and Rio Santiago at about 22N, and the southern limits at about 11N in northeastern Costa Rica. His map, however, further limits the Mesoamerican center of crop diversity around the southern boundaries of the Sonoran and Chihuahuan Deserts (Fig. 2). This natural geographic boundary is roughly the southern limit of what Latin American geographers sometimes refer to as Aridoamerica, or as Norteamerica Arida (for those not wishing to infer that it includes the only arid zones in the Americas) (Kirchhoff, 1954). Actually, Kirchhoffs (1954) cultural regions of Ar- 390 [VOL. 39 This content downloaded from 132.248.181.31 on Sat, 1 Mar 2014 15:04:38 PM All use subject to JSTOR Terms and Conditions NABHAN: NATIVE CROP DIVERSITY Fig. 1. Delimitation of North American versus Central American and Mexican centers of crop origins as defined by Zeven and de Wet (1982). Base map follows Rzedowski's 1978 map of the vegetation of Mexico, extended into the southern U.S. idoamerica and Oasis America are combined in my revised concept of Aridoamer- ica, since I see a gradual transition in genetic resources available within the two. Kirchhoffs Oasis America roughly coincides with what was recently discussed as the Sonoran Desert Agricultural Region and its native crop complex (Nabhan and de Wet, 1984). However, a broader binational region of Aridoamerica may be more geographically cohesive as a proposed center of diversity for the following reasons: 1. It encompasses the major North American deserts within which indigenous agriculture shared many of the same crops. 2. Certain crops that were previously assigned to the Sonoran Desert Agricul- tural Region also extend beyond desert environments into adjacent semiarid and subtropical sierras. Though not arid in the true sense, these upland areas share the same pattern of evapotranspiration far exceeding precipitation, and their climates are controlled by the same air current patterns. 3. Certain cultural subfamilies, for example, the Sonoran branch of the southern Uto-Aztecan languages, include tribes that historically occupied lands in the So- noran Desert, Chihuahuan Desert fringe, and Sierra Madre Occidental (Miller, 1983). Apaches, the southernmost Athapaskans, also had a binational distribution that extends from Great Basin to Sonoran and Chihuahuan Desert areas, and they formerly ranged into the Sierra Madres as well. Yuman tribal distributions extend from the Mohave Desert, into the Sonoran Desert and adjacent uplands of Baja California, again on both sides of the international boundary. 1985] 391 This content downloaded from 132.248.181.31 on Sat, 1 Mar 2014 15:04:38 PM All use subject to JSTOR Terms and Conditions ECONOMIC BOTANY Fig. 2. Northern delimitation of Mesoamerican center of crop diversity and of Mesoamerican cultures as defined by Leon (1979). His Mesoamerica extends southward to northeastern Costa Rica, and in his written description, Mexico's Rios Panuco and Santiago are northern borders. In short, an Aridoamerican center of crop diversity is binational, including both deserts and adjacent semiarid uplands, as do cultural distributions in this region. It extends from roughly the southern Great Basin in Utah, around 38N, southward at least as far as the Tropic of Cancer, and perhaps farther south in the Chihuahuan Desert, to around 23N. Its farthest westward reaches prehistorically were the Salton Basin and Coachella Valley in California. It extended east to the Rio Grande (Rio Bravo) drainage, but farther south, I am not sure of its eastern limits. I hope that the provisional boundaries of an Aridoamerican center of native crop di- versity, as illustrated in Fig. 3, can be refined on the basis of criticism from archaeologists and plant geographers. Until more archaeobotanical work is ac- complished in northern Mexico, the geographic limits of this region's crop heritage will remain imprecise. GENE POOLS OF ARIDOAMERICAN CROPS The genetic diversity of crops within any region may be related to a number of factors, such as: (1) the antiquity and continuity of agriculture; (2) ecological (habitat) diversity; (3) cultural diversity; and (4) introgression of crops with their wild or weedy relatives (Harlan, 1975). Evidence for agriculture in Aridoamerica dates back to between 2000 and 1500 B.C. (Woodbury and Zubrow, 1979). After crops and farming technology emerged or were introduced, they were refined to fit a diversity of local environments (Woosley, 1980). The diverse cultures in the region also applied distinct folk scientific and aesthetic criteria to crop varietal 392 [VOL. 39 This content downloaded from 132.248.181.31 on Sat, 1 Mar 2014 15:04:38 PM All use subject to JSTOR Terms and Conditions NABHAN: NATIVE CROP DIVERSITY Fig. 3. Delimitation of Aridoamerica as defined by geographical and cultural-agricultural factors. I have followed the vegetational limits of arid and semiarid zones to some extent, further limiting this center by excluding arid areas where there is no known prehistoric or protohistoric evidence of native agriculture. Southern and eastern boundaries will require further refinement. selection for color, taste, etc., such that Navajo blue flour corn looks remarkably different from Hopi blue flour corn grown just a few miles away from it. Intraspecific genetic variation within each crop species in Aridoamerica can also be evaluated from the species' geographic origins and interbreeding with species or varieties found within the region. Some crops were introduced after being domesticated elsewhere and have had no gene exchange with wild species in the region. These culturally allochthonous crops include: Lagenaria siceraria, the bottlegourd; Gossypium hirsutum var. punctatum, cotton; and the tobaccos, Nicotiana rustica, and later, Nicotiana tabacum. There are additional allochthonous crops that have wild relatives that barely enter this region. Limited gene exchange is theoretically possible between the wild and domesticated taxa, but it has not been documented. Canavalia ensiformis, the jack bean, has a close wild relative, C. brasiliensis, in Sinaloa. The degree of cross-compatibility and distributions of these taxa are poorly known. Pennington's (1982) ethnohistory of Eudeve agriculture argues that the nearly extinct Sonoran tribe once cultivated a Chenopodium species. If subsequent work confirms the presence of C. berlandieri var. nuttalliae in this region, as in the Mississippi Valley and Mesoamerica (Wilson, 1981), gene exchange with common, weedy C. ber- landieri varieties may have once been possible in Sonora. The interrelationships of Cucurbita pepo are also not as simple as many would think. Remains of Cucurbita pepo in the Ocampo Caves of Tamaulipas dating 1985] 393 This content downloaded from 132.248.181.31 on Sat, 1 Mar 2014 15:04:38 PM All use subject to JSTOR Terms and Conditions ECONOMIC BOTANY from 7000-5000 B.C. may represent the texana variety, now known to be more widespread than formerly assumed. Whether this variety was native to north- eastern Mexico and the adjacent United States is currently unresolved. After other C. pepo cultivated varieties were introduced, it may have exchanged genes with them. Until Hugh Wilson and his colleagues have reevaluated relationships among these taxa, it will remain difficult to interpret early C. pepo records on the north- eastern fringes of Aridoamerica. In published distributions of wild Phaseolus, no evidence is cited for either P. vulgaris or P. coccineus growing in arid northern Mexico. Wild varieties of both species have now been collected on the fringes of the Sierra Tarahumara in Chi- huahua. Together with my colleagues Jose Muruaga, Barney Burns, and Amadeo Rea, I have located populations of wild P. vulgaris near Yepachic and Balleza, and wild P. coccineus near Balleza and Laguna de Babicora. These newly located northern populations have the potential to hybridize with cultivated populations of domesticates, and the 2 species are cross-compatible. In contrast, Phaseolus lunatus var. silvester presently reaches Sinaloa, but no farther north. Introgression of wild genes from this taxon into domesticated limas or sievas is hardly a feasible explanation for the great varietal diversity of lima bean landraces found among the Pima, Hopi, and Pueblo Indians hundreds of kilometers to the north (Mackie, 1943). The Puebloan cultures of the Colorado Plateau may have harbored a secondary center of diversity for cultivated Phaseolus lunatus, far from its center of origin. Physalis philadelphica, the tomatillo or miltomate, may be the "tomato" men- tioned in early accounts of Eudeve agriculture (Pennington, 1982). Its wild variety has been undergoing selection and its seeds are saved at Zuni, far removed from other wild populations. The cultural geography of Physalis deserves further in- vestigation in Aridoamerica. For another set of allochthonous crops, continued introgression with wild rel- atives is more likely, and this process has probably contributed to landrace di- versity in the region. For example, Wilkes (1970) documented Nobogame teosinte introgression into maize varieties in the northern Tepehuan region of Chihuahua. This teosinte introgression has long been considered as a key factor for the presence of certain morphological traits found in prehistoric maize elsewhere in Aridoam- erica, but such influence is now subject to debate by corn geneticists. The bewildering diversity of morphological forms of Amaranthus hypochon- driacus and A. cruentus from the Warihio Indians may be due to introgression with A. hybridus (Sauer, in Nabhan, 1979a). Near the Sonora-Chihuahua border a few Warihio, Mayo, and Mountain Pima farmers still plant amaranths. Ama- ranthus hybridus is found in these fields in high densities (Nabhan, 1979a). More recently, I have encountered possible evidence of A. powellii introgression with A. cruentus grown by the Hopi at Lower Moenkopi, Arizona. Work in progress may confirm whether gene exchange is actually occurring in Hopi fields. It is intriguing that Jonathan Sauer (1977) has proposed a North American domesti- cation of A. hypochondriacus from A. powellii, on the basis of geographic as well as ethnohistoric data from Aridoamerica. Laura Merrick and I have also begun documenting introgression between wild Cucurbita and Pima landraces of squashes (C. mixta and possibly C. moschata) at Onavas, Sonora (Nabhan, 1984). This work will complement Merrick's more extensive biosystematic study of the Cucurbita sororia complex (Merrick and 394 [VOL. 39 This content downloaded from 132.248.181.31 on Sat, 1 Mar 2014 15:04:38 PM All use subject to JSTOR Terms and Conditions NABHAN: NATIVE CROP DIVERSITY Nabhan, 1984; Merrick, in prep.). It may be that this introgression is responsible for the rich varietal diversity of C. mixta cushaw squashes in Sonora and adjacent Sinaloa and Chihuahua, where silver-seeded, Taos-type, and marginless seed seg- regates appear together in populations that evolved within Indian fields. In addition, farmers from Onavas, Sonora, insist that wild chiltepines are con- tributing genes to their local cultivated chiles, some of which thereby become too hot to market! Cindy Baker and I have submitted samples of wild and cultivated Capsicum annuum from such settings to Dr. Steve Tanksley and Fernando Loasa of New Mexico State University. They hope to use electrophoresis as a tool to resolve whether chiltepine genes indeed have introgressed into chiles (Tanksley, 1983). Dr. Giles Waines of the University of California at Riverside is studying a similar story of potential introgression of wild and domesticated tepary beans (Phaseolus acutifolius), both of which occur on the same floodplains in Sonora, Chihuahua, and Arizona (Nabhan, 1979b). The role of introgression in the diversification of sunflower landraces after initial domestication also deserves further field study. Hopi dye sunflowers (Helianthus annuus) have long been grown in the same fields where Helianthus anomalus, a cross-compatible species (Rogers et al., 1982) is protected by the Hopi (Nabhan and Reichhardt, 1983). Wild Helianthus annuus is not common in Hopi fields, although it is abundant in those of other tribes, where it could potentially "swamp" the domesticate. Most important in defining Aridoamerica as a distinct center of diversity are its endemic domesticates. Panicum sonorum was domesticated entirely within this region, but its relationships with other taxa in the Panicum hirticaule complex are not clear (Nabhan and de Wet, 1984). Today, this domesticate is extremely rare. In the area inhabited by the prehistoric Hohokam, Hordeum pusillum ap- pears to have been culturally selected. Karen Adams, with the aid of Vorsila Bohrer, Robert Gasser, and Charles Miksicek, is in the process of studying this little-known prehistoric domesticate. Also, the basketry fiber plant, Proboscidea parviflora var. hohokamiana appears to have been domesticated in the last century in the northern Sonoran Desert, from which it was rapidly diffused to the Mohave Desert and Great Basin (Nabhan and Rea, in press). Peter Bretting (1982) and a team of Arizona scientists (Nabhan et al., 198 la) have published on the dynamics of this domestication and continue to study it. This domesticate no doubt ex- changes genes with wild P. parviflora var. parviflora throughout much of its range of cultivation. Although I will not review them here, incipient domesticates such as Jaltomata in Mesoamerica and southern Aridoamerica (Davis and Bye, 1982), and an Allium cultivated by the Papago open many new questions. At the same time, we must reconsider whether or not early historic cultivation of Agave (Robertson, 1972) and of a turnip or rutabaga-like native root (Pennington, 1982) resulted in distinct genotypes. There is intriguing archaeological evidence that southern Arizona Ho- hokam may have cultivated an undetermined Agave in Classic and Late Sedentary times, in habitats where wild agaves are not now found (Fish et al., in prep.). DRAINING OF GENE POOLS Of the above-mentioned native crops, Hordeum, Chenopodium, and Canavalia regional gene pools have dried up completely, i.e., these crops are extinct in indigenous communities of Aridoamerica. Locally adapted Panicum, Amaran- 1985] 395 This content downloaded from 132.248.181.31 on Sat, 1 Mar 2014 15:04:38 PM All use subject to JSTOR Terms and Conditions ECONOMIC BOTANY thus, Gossypium, and several species of the cultivated but perhaps not necessarily domesticated plants (e.g., Hyptis) are near extinction within the region. Regional ecotypes of chiles, maize, and sunflowers are being rapidly replaced by introduced, improved varieties. Other native crops are not threatened, but have certainly decreased in abundance as modem cash-crop monoculture has replaced indige- nous mixed cropping. Modem agricultural management practices are likely de- creasing the potential for introgression of crops with weeds, since more intensive tillage and herbicide use are commonplace in certain subregions. Let me emphasize that the oft-cited example of Green Revolution hybrids replacing local landraces hardly accounts for much of the genetic erosion of in- digenous cultivated plants in Aridoamerica. Most, but not all, of this erosion has occurred within the last century, and is the result of several interacting factors: 1. Acculturation/abandonment of farming traditions by indigenous populations. 2. Economic change/rural migration to cities. 3. Destruction of indigenous agricultural "habitats" via man-induced environ- mental change. 4. Usurpation of traditional farming areas or irrigation water by others. 5. Replacement of small-scale mixed cropping by mechanized farming of single (often exotic) crops. On the scale of villages and fields, there has been a decrease in the genetic variation found within certain crop species due to: 1. Fewer neighboring farmers growing a particular crop, resulting in a smaller population/gene pool overall. 2. Smaller populations of crops per field. 3. Less frequent planting of a crop or landrace. 4. Loss of the skills of seed selection and storage. 5. Change in exposure to cross-compatible weedy species. 6. Change in vulnerability to competition by weeds and consumption by pests (including introduced species). 7. Collapse of several landraces into one multiline gene pool. CONSERVATION OF EXTANT NATIVE-CROP DIVERSITY A number of ex situ and in situ conservation strategies for conserving extant genetic diversity have been proposed. One might pursue emergency funding for plant collecting in localities where rapid cultural or environmental change is occurring, hoping to salvage as many varieties as possible for placement in seed banks or botanical gardens. At the other extreme, perhaps, is the Iltis (1974) suggestion that we "freeze the genetic landscape." Iltis argues that scientists should urge politicans to negotiate the "deliberate exclusion of agricultural improve- ments" into "selected specific local genetic landscapes" where primitive landraces continue to exchange genes with adjoining weedy and wild populations. He sug- gests that genetic erosion and environmental destruction caused by "economic development" projects be controlled by creating biosphere reserves where indig- enous farmers would be subsidized to continue with their traditional agriculture. Wilkes (1971) proposed that "world genetic resource areas" be established where native agriculturalists would become curators of living collections of crossbreeding crops and weedy relatives. To preserve areas of maize-teosinte introgression, for 396 [VOL. 39 This content downloaded from 132.248.181.31 on Sat, 1 Mar 2014 15:04:38 PM All use subject to JSTOR Terms and Conditions NABHAN: NATIVE CROP DIVERSITY example, Wilkes suggests that only 5 carefully chosen 5 x 20 km strips would be needed. While both of these strategies are motivated by a true concern about the alarming rate of genetic erosion, they have been criticized for being somewhat inoperable and ethnocentric. By relying solely on ex situ conservation measures, we make the collected material vulnerable to (1) decreased population variation and genetic shifts due to inbreeding, (2) the effects of seed storage, occasional, recurrent grow- outs, and (3) human errors in sampling and handling (Roos, 1980, 1984). Even with better handling of germplasm in seed banks, we cannot duplicate the "dy- namic evolutionary potential" of crops still found in their cradles of origin (Iltis, 1974). But by relying solely on in situ conservation measures, we would be fighting the tide of acculturation, assimilation, and economic change that affect virtually every human population on the planet. Today, it is inevitable that a sizeable portion of any indigenous community will want to seek opportunities other than those traditionally open to them in their village, even if it means foresaking part or all of their agricultural heritage. "Freezing" an agroecosystem may not even be possible, given that cultural and environmental changes will continue regardless of intentional efforts to stop or slow them. Instead, it may be possible to combine selected in situ and ex situ conservation measures in a dynamic way. Each crop or landrace may be suffering from a different rate of genetic erosion. The rarer a landrace or crop complex has become in a village, the more important it is to consider "rescue" techniques to assure that some seeds are conserved ex situ. When a landrace is still grown by a number of families in different villages, greater effort should be exerted to encourage at least a few growers to conserve it in situ. Various combinations of in situ and ex situ crop conservation are now being attempted by a number of organizations, including the nonprofit Native Seeds/ SEARCH organization based in Tucson. When Native Seeds/SEARCH staff makes field collections, farmers who donate seeds are usually asked if they have enough surplus supply to spare and if there are other kinds of native seeds that they formerly grew which we might help obtain for them. In general, conservation measures will be most effective when communities of native farmers are cognizant of, and involved in, their planning and implemen- tation. They should become aware that reciprocal exchanges of seeds with gene banks are possible and that other options are available as well. Farmers who donate seeds to a gene bank or botanical garden must be informed how to gain access to subsamples of seed increases held in gene banks and botanical gardens, in case they happen to lose their remaining seeds. They must know the reasons that others are interested in these seeds, and their own efforts to propagate them must be reinforced. In this respect, it is heartening that agricultural education programs on Indian reservations now include information from tribal elders on traditional planting techniques, seed saving, and selection (Bingham and Bingham, 1979; Nabhan et al., 1981 b). In the future, indigenous foods, seeds, and farming practices should be further encouraged as part of cultural revival movements, tribal health, and gardening projects, and educational outreach programs. New cultural and economic incen- tives for diversified, regionally adapted agriculture must be considered, particu- larly when extant incentives for growing certain native crops no longer are effective. 1985] 397 This content downloaded from 132.248.181.31 on Sat, 1 Mar 2014 15:04:38 PM All use subject to JSTOR Terms and Conditions ECONOMIC BOTANY ACKNOWLEDGMENTS Research on native crop/weed introgression in Aridoamerica has been supported by grants from the Wenner-Grenn Foundation and the National Science Foundation Anthropology and Linguistics Sections (BNS-8317190), through the University of Arizona. Research on the causes of genetic erosion and conservation strategies applicable in this region has been aided by C.S. Fund and Tides Foundation support to Native Seeds/SEARCH. I thank Mahina Drees, Karen Reichhardt, Barney Burns, Laura Merrick, Robert Bye, Charles Miksicek, and Garrison Wilkes for helpful discussion. LITERATURE CITED Bingham, S., and J. Bingham. 1979. Navajo Farming. 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Impact of Plant Breeding On The Genetic Diversity of Cultivated Strawberry As Revealed by Expressed Sequence Tag-Derived Simple Sequence Repeat Markers