Smith (1967) reported remains of nine cactus species (Table 9.1) from archaeological excavations of prehistoric Mesoamerican sites in caves of the Tehuacán Valley. For caves at Guilá Naquitz, Smith (1986) reports stems, fruits, and seeds of Opuntia species in almost all of the strati-graphic zones studied, from nearly 12,000 years BP, as well as a gumball that could have come from a columnar cactus. Callen (1967) identified the following types of cactus remains in human copro lites of Tehuacán: (1) "Opuntia," which might represent some of the 18 species of this genus existing in the region (Arias et al. 1997); (2) "Lemaireo-cereus," which might represent some of the 13 species of columnar cacti of the genera Escontria, Myrtillocactus, Pachycereus, Polaskia, and Stenocereus (Casas et al. 1999a); and (3) "cactus tissue," from unidentified cacti. Callen (1967) further found that in the earliest coprolites from the El Riego phase (8,500-7,000 years BP), these types of cactus remains were a part of a wild food diet, along with Setaria spp. seeds, pochote (Ceiba parvifolia) roots, maguey (Agave spp.) leaves, and meat. In the Coxcatlán phase (7,000-5,500 years BP), stem tissue and fruits of "Opuntia" and "Lemaireocereus" were equally dominant materials. In the Abejas (5,500-3,300 years BP), Ajalpan (3,500-2,900 years BP), Santa María (2,900-2,200 years BP), Palo Blanco (2,200-1,300 years BP), and Venta Salada (1,300460 years BP) phases, findings suggest that consumption of "Lemaireocereus" stem tissue, fruits, and seeds were more important than products of "Opuntia"; and during the Ajalpan and Santa María phases, "Lemaireocereus" was the principal plant constituent of human diets.

The importance of cacti in Mesoamerican cultures can be recognized in pre-Columbian codices, which contain many toponymic glyphs referring to the names of cacti or their parts. Among the most famous are Tenochtitlán ("place of stony prickly pears" in Náhuatl), the original name of Mexico City, and Nochistlán ("place of prickly pears" in Náhuatl), in the state of Oaxaca. Historical information on utilization of cacti can be found in La Historia General y Natural de las Indias, published by Oviedo y Valdés in 1535. The Barberini Codex from 1552 (De la Cruz and Badiano 1964) includes information on medicinal utilization of Tlatocnochtli, a species of Opuntia, and a description of Teonochtli, identified as Stenocereus sp. by Bravo-Hollis (1978). The Florentino Codex (Sahagún 1970) contains a section dedicated to the description of the "diversity of tunas," which includes a list of variants of Opuntia species and their uses as edible fruits and stems. Estrada (1989) identified Cacanochnopalli (a Náhuatl term) as O. megarhiza, Tecolnochnopalli as O. streptacantha, Uitzocuitlapalli as Aporocactusflagelliformis, Nopalxochitl as

Epiphyllum ackermanii, Teonochtli as Hylocereus undatus, Peyotl as Lophophora williamsii (now commonly known as "peyote"), as well as several types of Tecomitl as Mammil-laria, Echinocactus, and Ferocactus species. The Florentino Codex also includes information on two columnar cacti, one of them called Netzolli, which is probably Escontria chiotilla, and Teunochtli, which could be a species of Stenocereus (Casas et al. 1999a). The books of Francisco Hernández in the 16th century describe several species of cacti utilized as medicine, among them several species of Opuntia, two columnar cacti identified as Myrtillocactus geometrizans, and a possible Stenocereus species called Teonochtli (Hernández 1959). The Geographic Relations of the XVI Century described the cultivation of Opuntia species for the production of cochineal and contains a reference to the columnar cactus Teonochtli, the "Relation of Acatlán" (Acuña 1985). Based on these sources, cacti were clearly utilized as food (fruits, young stems, and in some cases the flowers and seeds) and medicine (fruits, stems, and roots).

In the 16th century, Oviedo y Valdés (1535) and Sahagún (1985) described how the harvest of fruits of Opuntia spp. and columnar cacti was crucial for subsistence of some pre-Columbian and post-Conquest peoples from northern and central Mexico. For example, indigenous people migrated during the summer from the coast of the Gulf of Mexico to the highlands of the northern plateau, looking for the fruits of platyopuntias. In this region, people stayed for two months, migrating from place to place consuming fruits.

Smith (1967) considered that species of Opuntia could have been among the first plants subject to human manipulation in the Tehuacán Valley, but no archaeological evidence exists. Apparently, Opuntia was cultivated in the 16th century for the production of cochineal (Opuntia ficus-indica, O. tomentosa var. hernandezii, and Nopalea cochenillifera). However, cultivation of Opuntia and columnar cacti for the production of fruits, as currently occurs, was not clearly recorded. The only document that indicates such cultivation is the book of Sahagún (1985), in which the wild variants are distinguished from others (presumably cultivated). This omission could be because the Spaniards did not consider fruits of cacti as important resources and therefore did not describe them, as was the case for many other plants cultivated by natives (Casas et al. 1999a). Such omission could also be explained if cultivation of these plants started more recently. Further studies can provide information on the changes of cacti under domestication, which would be helpful to estimate the antiquity of cactus domestication.

Ethnobotanical Information

According to the compendium of the Cactaceae of Mexico by Bravo-Hollis (1978; Bravo-Hollis and Sánchez-Mejorada 1991), about 850 cactus species occur within the Mexican territory and 420 in the Mesoamerican region. Ethno-botanical studies in the region have documented a total of 118 cacti species utilized by indigenous peoples (Table 9.2). Useful cacti include Opuntia species, epiphytes, as well as columnar, spherical, barrel, and shrubby cacti. Among the columnar cacti, nearly half of the species are uncultivated giant columnar cacti, some about 15 m tall, with slow vegetative growth, which flower only after decades (Casas et al. 1999a). However, 23 species of columnar cacti (Table 9.2) are 2 to 8 m high. They grow faster than the giant columnar cacti, and flower 6 to 8 years after seed germination (24 years after vegetative propagation); most of them reproduce vegetatively and are cultivated.

Cacti are used mainly for their fruits, which may be consumed both fresh and dried and are used to prepare jams (Table 9.2). With the exception of subfamily Pereskioideae, fruits of nearly all species of cacti are consumed by people (Bravo-Hollis 1978). Fruits of 83 species (Table 9.2) are the most commonly consumed, and it is possible to distinguish: (1) species producing sweet juicy fruits, which are "good quality fruits" and are commonly harvested; (2) species whose fruits are of "regular quality" and are collected only occasionally, because of the scarcity of individual plants or populations, tall branches, long or abundant spines, or lack of tastiness; and (3) species whose fruits do not contain juicy pulp and are consumed only during food scarcity. The main groups of cacti producing edible fruits are columnar cacti and Hylocereus species with fruits called pitayas and pitahayas, respectively (Chapter 11); some Mammillaria species with fruits called chilitos; and, most important, Opuntia species (Chapter 10), whose sweet fruits are called tunas and consumed fresh, and whose sour fruits are called xoconoztles (from the Náhuatl xocotl, meaning sour, and nochtli, prickly pear) and are utilized as greens, condiments (boiled or fried), or as an ingredient for several other dishes. Colunga (1984) identified as xoconoztles variants of the species Opuntia joconstle, O. lasiacantha, O. leucotricha, and O. streptacantha as well as the red variant jitomatilli of O. megacantha, which is utilized as a substitute for tomato, and the variant brevas of Opuntia robusta var. robusta, whose peel is consumed fried, resembling French fried potatoes.

From the useful species of cacti reported, the stems of 62 species are cut and fed (after removal of the spines) as fodder to domestic donkeys, cows, and goats (Table 9.2).

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