We also mixed in photos of similar local plants to establish whether or not the potential interviewee was actually talking about cycads. In many cases, cycads are confused with palms, ferns, pine trees, and other plants, particularly in regions where substantial knowledge erosion has taken place. If possible, we had the potential interviewee show us what they thought was a cycad plant of their own volition—i. In addition, given the extensive work that has been done by biologists, it was usually easy for us to identify local cycad species, which are typically limited to a handful in any given locale, with little potential for confusion.
Any uncertainties are flagged Additional file 1 : Table S1. If we established that the potential interviewee was familiar with cycads, then with their permission, an open, exploratory interview usually commenced, focusing on its names, and if these were in an Indigenous language, we asked the interviewee or a family member, guide, or interpreter to write them phonetically. Names were also recorded on audio, if possible. Whether or not names were known, we asked whether any uses were known.
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In many cases, however, in-depth interviews were left for a later time, to allow the interviewee to collect their thoughts. Where very little was known, we asked the interviewee or they responded unsolicited who would possibly know more, thus a snowball sampling method of potential interviewees was used in some communities. In Teenek, Nahua, and certain other Indigenous communities, where we apprised community leaders of the nature of the research, they met to decide whether to grant FPIC, and if they did, thence accompanied us or let us go solo or with an interpreter to interview our target of ten pre-designated elderly cycad experts.
Each community was quite distinct in how it approached our research, and in some regions and contexts, particularly in non-Indigenous areas, it was not necessary to do more than inform local leaders of our presence and purpose and show letters of introduction from our research institutions. In some cases, local leaders evinced marginal or no interest in what we were doing and told us there was no need to garner their permission; in such contexts, we sought FPIC person by person. Interpreters were utilized in the case of interviewees who did not or preferred not to speak in Spanish.
We gave sufficient opportunity for interviewees to provide as much detail as they wished. In Mexico, though we did not find this to be the case, we opted to hide identities and localities to impede plant traffickers who access published data to find informants who can point them to cycad populations.
This is a serious problem in Mexico, a result of the extremely high value of cycads for domestic and foreign decorative use and is why exact geographical coordinates of population locations for most cycads should not be revealed in publications [ 4 , 5 ]. As an additional precaution, we carefully explained to collaborators and authorities, whom we assumed were unaware, that cycads are sometimes utilized for secret and even illicit purposes connected to ethnomedicine and narcotics [ 58 ].
Though we were never asked to destroy our data, in several cases potential interviewees—typically traditional healers and shamans—whom local authorities had identified and recommended to us did not wish to be interviewed once they learned the exact nature of our research. In NEMX and Honduras, we stayed in local homes over the course of days and in some cases weeks, affording us the opportunity to gain trust.
Where permitted, we visited and documented local cycad populations. Where possible, we participated in or observed cycad food preparation and consumption as well as religious events. In addition, we searched local landscapes, particularly graveyards, for iconographic representations and decorative uses. While some terms are generic and are applied widely to other plants, others are specific to cycads. We recorded locally used terms in 17 Indigenous languages as well as in Spanish and English.
Several terms combine roots from two languages. More ethnic groups 27 Indigenous, in addition to non-Indigenous mestizos have terms for cycads than are reflected in languages, because some groups no longer retain Indigenous languages, while other groups only retain names in Spanish or English.
Several ethnic groups visited—e. In some cases, this may have been due to insufficient fieldwork on our part, but in many instances, it was the fault of eroded cultural knowledge as well as non-transmission of knowledge between earlier inhabitants and in-migrants who had mixed with or supplanted them. In many instances, cycads were known only as palm trees palmas , palmitas , etc.
Intra-regional commonalities in cycad usage and knowledge are detectable within five NMCA regions, based on geographic proximity and shared cultural histories. Scant ethnobotanical data are available. No Spanish or Indigenous terms or knowledge relate WMX cycads directly to any current or prior food uses, though two common NEMX food-related terms for cycads— chamal and teosintle —used as recently as the s in Jalisco suggest potential former food uses.
In addition, Ceratozamia spp. Southward, food uses are mostly unknown to mestizo, Nahua, and Totonac peoples e. Across the entire region, cycads are often conceptualized as types of maize, friends of maize, and ancestors of maize. Our preliminary researches and inquiries suggest cycads are of minor importance to local Indigenous groups primarily Maya language group speakers. They are occasionally made into breads and medicines. Belief in cycads as maize ancestors is found here among the Chontal de Oaxaca, and alimentary uses are still common among Chinantec speakers.
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Numerous ceremonial and medicinal uses exist. Elaborate pilgrimages involving carrying leaf bundles from sacred Dioon merolae populations to local altars characterize Indigenous cultures in the Isthmus of Tehuantepec and nearby regions of both Chiapas and Oaxaca. In northern Central America, sparse data for Guatemala and Belize indicate potential connections to maize, and an overall association with camotillo poison. For El Salvador, we found virtually no ethnobotanical data on Zamia herrerae , the single species recorded there.
Cueva de la Perra a. Cueva del Armadillo. Ojo de Agua cave c. Dioon angustifolium : 2 sclerotesta fragments, with Lagenaria and Cucurbita remains unexamined remains may include other Dioon fragments. El Riego Cave h. Escalera al Cielo i. There were 14 bases, 31 bracts, and seeds of this plant. Since of these acornlike seeds fill a liter, 8.
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But with maize only appearing in very small quantities, MacNeish concluded that La Perra had functioned as a seasonal wild plant foraging camp. Given cycad longevity, it is likely that the same populations and possibly some of the same plants were harvested continuously over this time frame. A question unanswerable until lithic or ceramic implements at La Perra are analyzed is whether or not the toxic starches of cycad seeds were processed for food, or whether the apparently innocuous sarcotestas alone were used they still are processed into tortillas in other parts of NEMX at present, e.
Purportedly small Dioon populations near the rock shelters would not have allowed them to occupy dietary roles as prominent as those in the Sierra de Tamaulipas [ 19 ] but the remains are important nevertheless as they show yet another area where cycads and early maize existed in the same cultural context.
While European chroniclers often recorded Zamia spp. The voluminous Florentine Codex [ 64 ], an ethnography relying largely on Nahua informants from central Mexico, contains no obvious references to cycads, and only a few vague potential references, e. All other written historical ethnobotanical data we have found for Mexico refer to the chamal cycads [ 67 , 68 ]. From the s onward, they were mentioned as important foods for indigenous people and starvation fare for non-indigenous people in NEMX.
Cora people utilize turhaa leaves from wild populations to decorate churches and churchyards in the Sierra del Nayar region during Holy Week Semana Santa. Teocentli tepecintle , Ceratozamia fuscoviridis leaf crown and cone on outside wall of colonial church, Huayacocotla, Veracruz, Close-up of dameu Dioon edule cones and leaves in Tancoyol, While we have examined but a very small portion of archival and published accounts potentially containing cycad references in Mexico, in Honduras we have studied all available land titles and other documents across the remote northeastern part of the country where culturally significant tiusinte cycads have been found [ 51 , 52 , 69 ].
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Extrapolating from our exhaustive Honduran research, we think it safe to assume fine-grained archival research elsewhere in MNCA where cycads have been prominent as staple foods and in religious contexts will yield similar results. Where cycads are still eaten and appreciated, oral histories typically recalled richer, more nuanced traditions and knowledge than those that persist today. Where cycads are no longer eaten but are still utilized for other purposes, particularly in Zapotec and Teenek contexts, cycad consumption often evoked distant shameful memories of dire poverty and lack of maize among elderly first-language speakers.
Teenek interviewees who as children had been forced to gather and eat cycad seeds during famines in some instances terminated interviews with us or cried over the long-buried memories of the suffering associated with the practice. The same can be said in Oaxaca, where Chinantec and to a lesser extent Mazatec speakers consume and value cycad foods at present, Chontal de Oaxaca value them as a maize food of the ancestors, while Zapotec people value them for religious reasons but excoriated them in communities we visited as famine foods. Across all MNCA cycad-eating regions, consumption has diminished considerably over the past century as modern transportation has brought communities closer to market centers and maize has become regularly and predictably available for purchase.
In the post-World War II period, cycads and numerous other wild foods were all but abandoned. Even in areas where strong connections between cycads and maize continue to exist and cycad eating is not stigmatized e.
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Mestizo practices have been derived in large part from Indigenous practices via knowledge transmission within mixed Indigenous-mestizo families and communities. However, non-Indigenous ingredients of European derivation are also incorporated into mestizo recipes, particularly in Honduras. As discussed below, starch detoxification is centrally important to cycad consumption. However, as noted above, the sweet sarcotesta is also eaten. It is consumed as a raw snack or in preserves jellies in Mexico, a practice unknown in Honduras. This nearly vanished food is still prepared by a few of the most impoverished residents of the Sierra Teenek region.
Additionally, in Sonora, extraction of Dioon stem starch for use in sotol alcohol resulted in the complete destruction of populations. In Honduras, we did not uncover accounts of stem starch use. In NEMX and Honduras, detoxification methods for seed starch favor the use of ash ceniza from a variety of trees [ 52 , 72 , 73 ], rather than lime cal.
In these two regions, except among the Tolupan and some Teenek, detoxification with ash is regarded as producing flour or meal superior to that produced using lime. Nowhere else did we find evidence of ash ever being used, lime predominating instead. Across the region, other methods, including boiling, baking on coals, and leaching in water and sunlight, are occasionally employed as well.
The critical moment in cycad starch detoxification is the point when the cook applies a test to determine that the starch is toxin-free. Food derived from incompletely detoxified cycad starch produces effects ranging from stomachaches, nausea and vomiting to death, the latter mentioned as the result of Zamia stem consumption in a NEMX Nahua community. Part of a large batch prepared by local women for sale at the tiusinte fair Feria del Teocinte.
Such products include either maize or wheat flour, and are baked in outdoor clay-brick ovens. They are known only from Gualaco.