Comparison of traditional wild plant knowledge between aboriginal communities inhabiting arid and forest environments in Patagonia, Argentina

ARTICLE IN PRESS Journal of Arid Environments

Journal of Arid Environments 69 (2007) 695–715

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Comparison of traditional wild plant knowledge between aboriginal communities inhabiting arid and forest environments in Patagonia, Argentina A. Ladio, M. Lozada, M. Weigandt Laboratorio Ecotono, CONICET, Centro Regional Universitario Bariloche, Universidad Nacional del Comahue, Quintral 1250-8400-San Carlos de Bariloche-Rı´o Negro, Argentina Received 27 March 2006; received in revised form 15 November 2006; accepted 22 November 2006 Available online 18 January 2007

Abstract The Mapuche people have a deeply rooted tradition of wild plant use for their subsistence. In this study we evaluated whether plant selection by the Mapuche is influenced by environmental constraints. Selection patterns of wild plant richness were compared in two Mapuche communities sharing the same traditions and beliefs but inhabiting different environments. Similarities and differences in the use patterns of wild medicinal and edible plants were quantitatively analyzed by using concepts derived from ecological theory. The Cayulef community is settled in the NW Patagonian (herbaceous) steppe while the Curruhuinca community is established near the Andean temperate forest, 100 km away from each other. Free-listing and semi-structured interviews were conducted with 32 families in Cayulef and 39 in Curruhuinca corresponding to 80% and 40% of each population, respectively. Data were analyzed by means of the Jaccard similarity index and statistical tests. The main results of this study showed that the home environment is a determinant factor that influences which plant species are utilized, as is the anthropic environment (with its high exotic species richness) in each community. However, similar patterns were found with respect to plant families, ways of utilization, plant parts and life forms utilized irrespective of where the communities are settled. In addition, in both populations, wild plant richness was significantly higher for medicinal than for edible species. Our results showed that the use of wild plants is primarily based on the utilization of species belonging to the nearest ecological environments that have both therapeutic and dietary functions. r 2006 Published by Elsevier Ltd. Keywords: Andean forest; Ethnobotany; Patagonia; Steppe; Useful plant richness

Corresponding author. Tel.: +54 2944 430949; fax: +54 2944 422111.

E-mail addresses: [email protected], [email protected] (A. Ladio). 0140-1963/$ - see front matter r 2006 Published by Elsevier Ltd. doi:10.1016/j.jaridenv.2006.11.008

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1. Introduction Historically, the use of medicinal and edible plants has contributed to the subsistence of many aboriginal Mapuche communities from northwestern Patagonia (Citarella, 1995; Ladio and Lozada, 2000; Estomba et al., 2006). At present, the Mapuche people maintain this plant gathering tradition in spite of severe changes these populations have undergone. Despite being native to temperate forests, the majority of the Argentinean Mapuche communities live in stepparian regions at present, although a few reservations remain in the forest (Falaschi, 1996). In both environments, the traditional use of above and belowground plant biomass has provided food and medicine to these indigenous populations. Consequently, wild plant use and knowledge still constitute an important aspect of their cultural identity. The Patagonian region presents a noticeable environmental gradient which offers a wide variety of vegetation species within a range of 100 km. At one extreme lies the temperate rainforest (1500 mm) and adjacent to it the dry herbaceous steppe (300 mm). The abundant foliage and fruit from the forest as well as the spiny resinous plants from the steppe have provided valuable resources to these local communities (Rapoport et al., 1999, 2003). Since ancestral times, the use of wild plants in the Mapuche people reflects a strong interaction with their natural resources (Smith-Ramı´ rez, 1996; Ladio and Lozada, 2000). Long journeys in search of useful plants have been extensively documented in both archeological and historical studies in the Patagonian region (Nacuzzi and Pe´rez de Micou, 1984; Ladio and Lozada, 2000, 2004a). Plants from mesic and arid environments have been selectively gathered by these aboriginal inhabitants (Ladio and Lozada, 2004b). Our study compared the use of wild plants by two Mapuche populations, Cayulef and Curruhuinca, which share the same cultural identity, but live in significantly different environments, the steppe and the forest, respectively. This comparison might shed light on to our understanding of how the environment influences wild plant gathering traditions. We analyzed the plant characteristics and biogeographical origin of collected species, as well as selection patterns of use. Plant availability and richness play an important role in shaping ethnobotanical knowledge in human populations (Phillips and Gentry, 1993; Nolan, 1998). It has been proposed that the more readily available or ‘‘apparent’’ (visible) a species is, the more likely its use and cultural importance will be (Phillips and Gentry, 1993; Galeano, 2000; Albuquerque and Paiva de Lucena, 2005; Ladio and Rapoport, 2002). This hypothesis was mainly shown in humid tropical forests, whereas in arid areas, such as Patagonia, this trend remains unclear (Alburquerque et al., 2005; Almeida et al., 2005). Human influence over natural environments has created new plant gathering sites for medicinal and edible resources in which exotic weeds predominate (Dı´ az-Betancourt et al., 1999). In Patagonia, the initial spread of exotic species was found to start about three hundred years ago (Ezcurra and Brion, 2005). Consequently, the first uses of these plants might have occurred many generations ago. At present, the utilization of anthropic environments is common among many aboriginal and rural communities (e.g. Ladio and Lozada, 2004b). Anthropic areas, characterized by a high productivity rate, are well-known for their availability of useful plants (e.g. Rapoport and Drausal, 2001; Ogle et al., 2003; Stepp, 2004). Several studies have found that local people use medicinal plants from disturbed areas more frequently than from primary forests (Voeks, 1996; Caniago and Siebert, 1998;

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Frei et al., 2000; Stepp and Moerman, 2001). In addition, several authors have highlighted that medicinal species found in disturbed habitats are predominantly herbaceous plants, whereas those found in the primary forests are principally arboreal species (Voeks, 1996; Alburquerque et al., 2005). It has been proposed that medicinal flora is mainly constituted by herbs due to the high concentration of active secondary compounds of this life form (Bennett and Prance, 2000; Stepp and Moerman, 2001; Stepp, 2004). However, Almeida et al. (2005) have found that shrubs and trees from arid regions are relevant medicinal resources, containing bioactive compounds which have developed in these rigorous environments. When there is greater knowledge of the potential uses of a given species, a multifunctional and complementary utilization becomes enhanced (Nolan, 1998; Ogle et al., 2003). Moreover, wild plant knowledge, preserved in local food customs, is sometimes inseparable and incorporated into the traditional therapeutic system (Bennett and Prance, 2000; Ladio, 2002). Many of these species are used in multiple contexts, providing nutrients, adding to dietary variation and constituting worthy remedies for health problems (Etkin and Ross, 1982). These authors proposed that the overlapping uses of plants would extend the range of circumstances in which indigenous people benefit from plant constituents. The distinction between medicinal and edible categories may depend on the plant part used as well as to the amount of time required for its preparation (Etkin and Ross, 1982; Ladio, 2002). The use of underground or above-ground plant organs is also related to plant selection. Upon examining differences in foragers’ diets between savanna and rain forest landscapes, Gragson (1993) found that rainforest people rely particularly on aerial resources such as fruits, while savanna groups rely on herbs. In addition, underground plant organs seem to be particularly relevant to human populations settled in dry land environments (Gragson, 1997). In NW Patagonia, fruits are one of the most frequent resources in rainforests (Aizen and Ezcurra, 1998). On the other side, it is likely that, in communities inhabiting the arid steppe, herbs and underground parts would be more frequently consumed than fruit and aerial plant organs. Finally, if an ample resource offer is available, human communities orient their selection towards more significant and appreciated items, whereas under scarce resource conditions, the ecological niche widens to include a greater variety of species (Hardesty, 1975; Castro and Begossi, 1996). A similar trend prevails in the complementary use of different environments, a common pattern in Mapuche populations in a subsistence economy (Ladio and Lozada, 2004b). Since food and medicine are the categories of use with the greatest overlap, in the present study we explored whether a greater complementary use was found in the population inhabiting the rigorous steppe environment. The aim of the present study was to compare species richness and families of plants utilized by two Mapuche communities, Curruhuinca and Cayulef, and to analyze how these attributes are influenced by their home environments. (1) Firstly, we evaluated whether both communities select similar species and plant families (number of plants and taxonomical similarity). Significant dissimilarity of plants utilized was predicted. (2) Additionally, we analyzed whether the environments were similarly used by both communities. We expected to find a greater use of steppe species in Cayulef and a major use of forest plants in the Curruhuinca population, given that these correspond to their closest environment. (3) Moreover, we studied whether medicinal and edible species were gathered in certain preferential environments. (4) In addition, we compared the proportion

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of native and exotic species utilized by both populations, and whether these patterns differed in each ecological environment. (5) We examined the proportion of trees, shrubs and herbs utilized in both communities and analyzed the relationship with the type of environment inhabited. (6) We also hypothesized that in Cayulef, settled in an area with less wild plant richness, a greater number of edible and medicinal species would be utilized, in a more multi-functional and complementary way than in the Curruhuinca population. (7) Plant parts utilized for medicinal and edible purposes were compared in both communities. Significant differences were also predicted. (8) Moreover, we analyzed whether the Cayulef community uses more subterranean biomass (root-diggers) and the Curruhuinca more aerial biomass. (9) Finally, considering the niche breadth analysis, we hypothesized that in the Curruhuinca population there would be a greater plant selection due to the higher plant richness available, whereas in the Cayulef population there would be a greater incorporation of species given the prevailing plant scarcity. 2. The Cayulef community The Cayulef community is located in northwestern Patagonia, in the Argentine province of Neuque´n (391700 S and 701590 W, Fig. 1). Phyto-geographically, the region belongs to the Patagonian steppe. The study area receives strong westerly winds and has a mean annual temperature of 8 1C, a mean maximum temperature of 14.1 1C, and a mean minimum temperature of 2.4 1C. Mean annual precipitation of 300 mm is concentrated in autumn and winter (March–September). Therefore, the combined increase in temperature and decrease in rainfall during spring and summer (October–February) causes a moderate water stress conditions for plant growth (Barros et al., 1983). Sandy soils are dominant and the vegetation cover is mostly made up of native shrubs and grasses such as: Mulinum spinosum (Cav.) Pers. (Apiaceae), Haplopappus pectinatus Cassini (Asteraceae), Senecio filaginoides DC. (Asteraceae) and Stipa speciosa Trinius et Ruprecht (Poaceae) (Cabrera, 1976; Cabrera and Willink, 1980). Anthropic environments are found around their dwellings, dirt roads and corrals, which are principally dominated by the exotic grass Bromus tectorum L. (Poaceae), and the exotic herbs: Erodium cicutarium (L.) L’Herit. (Geraniaceae), Plantago lanceolata L. (Plantaginaceae) and Stellaria media (L.) Villars. (Caryophyllaceae). At present, the area is intensively grazed by both caprine and ovine livestock that negatively impact the abundance of the preferred palatable species, some of which have edible and medicinal usages (Golluscio et al., 1998; Ladio, 2006). This aboriginal community is composed of 40 families, 32 of which were interviewed. It is settled in two locations; one called La Costa located near the Catan-lil river and the other one: El Salitral, 30 km away. The access to both areas is difficult due to poor conditions of the few existing dirt roads. Livestock raising is the primary source of income. Fuel wood, mainly collected from their surroundings, is the primary energy source for cooking and heating. They inhabit precarious single-room houses with almost no furniture on bare-ground. Most adults are illiterate or semi-literate and children attend nearby rural schools. A professional health care assistant infrequently visits the community. 3. The Curruhuinca community The Curruhuinca community is settled within the temperate Andean forest of north-western Patagonia, in the province of Neuque´n, Argentina (411100 S, 711200 W,

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Fig. 1. Map of the study site. Geographic location of the Cayulef and Curruhuinca communities in Neuque´n province, Argentina.

Fig. 1). The landscape is composed of mountains and valleys, and the soil is of volcanic origin. The climate is cool-temperate with snow in winter. The area receives humid westerly winds and has a mean annual temperature between 9.5 and 5.4 1C. The mean annual precipitation varies between 500 and 1500 mm and is concentrated in autumn and winter (March–September). In the region the main vegetation species are Austrocedrus chilensis (D. Don) Pic. Ser. Et Bizarri (Cupressaceae), Nothofagus dombeyi (Mirb.) Oerst. (Fagaceae), Nothofagus obliqua (Mirb.) Oerst. (Fagaceae) and N. alpina (Poepp. & Endl.) Oerst. (Fagaceae), and in higher elevations Nothofagus pumilio (Poepp. & Endl.) Krasser (Fagaceae). In the understory: Maytenus boaria Molina (Celastraceae), Aristotelia chilensis (Molina) Stuntz (Elaeocarpaceae), Berberis darwinii Hook. (Berberidaceae), Ribes magellanicum Poir. (Saxifragaceae), Buddleja globosa Hope (Buddlejaceae), and Fuchsia magellanica Lam. (Onagraceae) are commonly found (Cabrera, 1976). Anthropic environments around their homes and surroundings included some native bushes and herbs such as: Berberis spp., Osmorrhiza

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chilensis Hook. & Arn. (Apiaceae), and exotic herbs Holcus lanatus L. (Poaceae), Plantago lanceolata and Taraxacum officinale Weber ex F.H. Wigg. (Asteraceae). Highly anthropically disturbed plots exhibit a decline in the abundance of native bushes and an increase in the dominance of the exotic shrub Rosa rubiginosa L. (Rosaceae), which also has fleshy fruit. The Curruhuinca population is composed of (approximately) 100 families living under rudimentary socio-economic conditions without running water, sewage system, telephones, etc. This study was conducted in two settlements of the community: Pil-Pil and Quila Quina, comprising about 60 families. They inhabit precarious single-room houses with almost no furniture on bare-ground. Fuel wood, collected mainly from their surroundings, is the primary energy source for cooking and heating. The access to the city of San Martin de los Andes (50.000 inhabitants) is easy through a near paved road with regular transportation service (10–20 km away). Most adults are illiterate or semi-literate and children attend nearby rural schools where they receive the state endorsed bilingual education in Spanish and Mapudungum (Mapuche language). A professional health care assistant visits the community once a week and the nearest public hospital is located in San Martin de los Andes. Livestock (cows, sheep and horses) constitutes an important source of income. This animal husbandry highly influences plant species composition causing environmental degradation and negatively affecting the abundance of plants in the area (Veblen et al., 1992). In both communities, most dwellers carry out their own medical practices, but sometimes they consult the ‘‘machi’’ or ‘‘yerbatera’’ who is the most highly regarded ‘‘medical practitioner’’, especially when health problems are considered related to illnesses of the soul or supernatural factors. 4. Material and methods 4.1. Sampling and interview process In each population, 39 families (39% of the Curruhuinca population) and 32 families (80% of the Cayulef population) were randomly selected. We interviewed people over 20 years of age, and enquired about medicinal and edible plants utilized. The average age of the informants was 50715 years in Cayulef and 5974 years in Curruhuinca. Houses were selected using random numbers, and one member per family was interviewed in Spanish. Introductory talks, aimed at establishing contact with each interviewee, were done on previous days. Ethnobotanical data were obtained by means of free listings and semistructured interview methods (Alexiades, 1996; Albuquerque and Paiva de Lucena, 2004; Edwards et al., 2005). The participants seemed to be keen on answering the questions in the interview, which took about 2 h, because they were interested in documenting and preserving their knowledge about wild plants. We adopted this methodology given that both communities were interested in participating, and because data collection is simple and rigorous, it allows for statistical analysis, and it permits comparable surveys elsewhere. In the interview, we asked informants to provide medicinal and edible plants’ local names, their uses and utilized parts. Some of the basic questions asked were: ‘‘which plants do you use for food and which for illnesses? Which illnesses? Which plant parts do you use? Where do you collect them?’’. Interviewees mentioned species using folk names and then we used specimens’ vouchers and photographs to identify them properly. We supplemented this

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information with field outings accompanied by some interviewees and key informants who helped us prepare the voucher herbarium. We also recorded the ecological site where these species belong to considering the following categories: Anthropic: an environment created by humans as orchards, fields, corrals, roads around dwellings; Forest: Andean temperate forests dominated by A. chilensis and deciduous Nothofagus forests; and Steppe: driest areas dominated by herbs and shrubs and with ‘‘mallines’’, often-swampy meadows used for cattle raising. Life forms were also evaluated and classified in: Herbs: non-woody plants as annuals, ferns, grasses and vines; Shrubs: woody multi-stemmed plants less than 5 m tall and Trees: singlestemmed woody plants greater than 5 m. Plant part were categorized in roots, rhizomes and bulbs, leaves, stems and flowers, and fruits and seeds. 4.2. Specimen identification Voucher plant collection and specimen identification was carried out by the authors and key informants. Plants mentioned by their local names were identified in their scientific nomenclature following Correa (1969, 1971, 1978, 1984, 1988, 1998, 1999) and Ezcurra and Brion (2005). Plants were also classified into native and exotic categories in accordance with their bio-geographical origin. The collected samples were stored in the Ecotono Laboratory herbarium. 4.3. Data analysis The first objective was evaluated by means of the Jaccard similarity index (Ho¨ft et al., 1999). This index is based on plant presence or absence in the community or data sets, while relating the number of species in common as a proportion of the total number of species, expressed as JI ¼ c=ða þ b þ cÞ  100, where c is the number of species in common, a is the number of unique species of the A community and b is the number of species solely of the B community. Species richness was compared through the Chi square w2 and Binomial tests (Agresti, 1996). From the second to ninth objective, data was analyzed by means of the Chi square w2 Test (po0.05). Additionally, binomial tests were used to evaluate hypothesis eighth and ninth (Agresti, 1996). 5. Results and discussion 5.1. Similarity in wild plant richness of edible and medicinal species utilized in both communities Although the two communities are settled in totally different environments, in both populations the total number of plants used and the type of use is similar. In the two communities, medicinal species are more frequently utilized (Cayulef: 77 and Curruhuinca: 68), then species for both purposes (medicinal and edible; Cayulef: 23 and Curruhuinca: 27) and lastly, those species used only for edible purposes (Cayulef: 10 and Curruhuinca: 15; Binomial test, po0.05, Table 1). However, in accordance with the Jaccard test, there is a low proportion of species in common (37% in total species). This was especially true for edible plants (24%). Other percentages were 40% for medicinal plants and among the

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Table 1 Medicinal and edible plants used by Cayulef community (Ca) and Curruhinca community (Cu) Life form

Plant part

Category of use

Community

Ecological environment

Origin

Acaena magellanica (Lam.) Vahl (abrojo) Acaena pinnatifida Ruiz & Pav. (pimpinela, abrojo) Acaena splendens Gillies ex Hook & Arn. (cepa caballo) Acaenna ovalifolia Ruiz & Pav. (cadillo) Acantholippia seriphioides (A. Gray) Mold. (tomillo del campo) Adesmia boronioides Hook.f. (paramela) Adiantum chilense Kaulf. (culandrillo) Allium cepa L. (cebolla) Allium sativum L. (ajo) Aloe sp. (aloe) Alstroemeria aurea Graham (amancay) Alstroemeria patagonica Philippi (liuto, chaquil) Anemona multifida Poir. (ane´mona) Apium australe Thouars (apio silvestre) Apium graveolens L. (apio) Araucaria araucana (Mol.) C. Koch (pehue´n, araucaria) Aristotelia chilensis (Molina) Stuntz (maqui) Arjona tuberosa Cav. (chaquil) Artemisa abrotanum L. (eter) Artemisa absinthium L. (ajenjo) Austrocactus patagonicus (Web.) Backeb. (tuna) Austrocedrus chilensis (D. Don.) Pic. Serm & Bizzarri (cipre´s) Azara microphylla Hook.f. (chin-chin) Azolla filiculoides Lam. (luchecito) Azorella monantha Clos (yareta, len˜a de piedra) Baccharis obovata Hook. & Arn (huautro) Baccharis sagittalis (Less) DC. (carqueja) Bacharis salicifolia (Ruiz & Pav.) Pers. (chilca) Balbisia gracilis (Meyen) Hunz.& Ariza (te santo, te de burro) Berberis buxufolia Lam. (michay, calafate) Berberis heterophylla Juss. ex. Poir. (michay) Blechnum penna marina (Poir) Kuhn (punque) Brassica rapa L. (repollo silvestre)

Rosaceae Rosaceae Rosaceae Rosaceae Verbenaceae Fabaceae Pteridaceae Liliaceae Liliaceae Liliaceae Amaryllidaceae Amaryllidaceae Ranunculaceae Apiaceae Apiaceae Araucariaceae Elaeocarpaceae Santalaceae Asteraceae Asteraceae Cactaceae Cupressaceae Flacourtiaceae Pteridaceae Apiaceae Asteraceae Asteraceae Asteraceae Ledocarpaceae Berberidaceae Berberidaceae Pteridaceae Cruciferae

Herb Herb Herb Herb Shrub Shrub Herb Herb Herb Herb Herb Herb Herb Herb Herb Tree Shrub Herb Herb Herb Herb Tree Shrub Herb Shrub Shrub Shrub Shrub Herb Shrub Shrub Herb Herb

1 1 2 2 2 2 2 1 1 2 1 1 2 1,2 1 3 2,3 1 2 2 2 2 2 2 1 2 2 2 2 2,3 3 2 2

Medicinal Medicinal Medicinal Medicinal Medicinal Medicinal Medicinal Both Medicinal Medicinal Medicinal Medicinal Medicinal Both Medicinal Edible Both Edible Medicinal Medicinal Edible Medicinal Medicinal Medicinal Both Medicinal Medicinal Medicinal Medicinal Both Edible Medicinal Edible

Cu Ca–Cu Ca Cu Ca Cu Ca Cu Ca–Cu Cu Cu Ca Cu Ca–Cu Ca Ca–Cu Ca–Cu Ca Ca–Cu Ca–Cu Ca Cu Cu Ca Ca Cu Ca–Cu Ca–Cu Ca Cu Ca Cu Ca

Steppe Steppe Forest Forest Steppe Steppe Steppe Anthropic Anthropic Anthropic Forest Steppe Steppe Forest Anthropic Forest Forest Steppe Anthropic Anthropic Steppe Forest Forest Steppe Steppe Forest Steppe Steppe Steppe Forest Steppe Forest Anthropic

N N N N N N N E E E N N N N E N N N E E N N N N N N N N N N N E E

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Family

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Species

2 2 1,2 2 2 2 2 2 2 2 2 2 1,2 2 2 1 2 3 2 1 1 2 2 2 3 2,3 2 2 2 1 2 2 2 2 1 2 2 2 3 2

Medicinal Medicinal Both Medicinal Medicinal Medicinal Medicinal Medicinal Medicinal Edible Medicinal Medicinal Both Edible Both Medicinal Medicinal Both Medicinal Both Edible Medicinal Medicinal Medicinal Edible Both Medicinal Medicinal Medicinal Medicinal Medicinal Medicinal Medicinal Both Medicinal Medicinal Medicinal Medicinal Edible Medicinal

Ca Ca–Cu Ca Ca Cu Ca Ca–Cu Ca Ca–Cu Ca–Cu Ca–Cu Ca Cu Cu Cu Ca–Cu Cu Cu Cu Cu Ca–Cu Ca–Cu Cu Ca–Cu Cu Ca Ca–Cu Ca Ca–Cu Ca–Cu Cu Ca–Cu Ca–Cu Cu Ca–Cu Ca–Cu Ca Cu Cu Ca

Steppe Anthropic Steppe Steppe Anthropic Steppe Forest Steppe Forest Anthropic Anthropic Anthropic Forest Anthropic Anthropic Anthropic Forest Anthropic Forest Anthropic Steppe Steppe Forest Forest Forest Steppe Steppe Steppe Anthropic Steppe Anthropic Steppe Forest Anthropic Forest Forest Steppe Steppe Forest Steppe

N N N N E N N E N E E N N E E E N E N E N N N N N N N N E N E N N E N N N N N N

703

Shrub Shrub Herb Herb Herb Herb Herb Herb Shrub Herb Herb Herb Shrub Herb Herb Herb Shrub Shrub Tree Herb Herb Shrub Shrub Tree Shrub Shrub Herb Shrub Herb Shrub Tree Herb Shrub Herb Herb Shrub Herb Herb Tree Herb

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Buddlejaceae Buddlejaceae Loasaceae Scrophulariaceae Asteraceae Callitrichaceae Gentianaceae Caryophyllaceae Solanaceae Chenopodiaceae Chenopodiaceae Chenopodiaceae Poaceae Asteraceae Asteraceae Apiaceae Coriariaceae Cucurbitaceae Asteraceae Apiaceae Apiaceae Rhamnaceae Winteraceae Proteaceae Empetraceae Ephedraceae Equisetaceae Equisetaceae Geraniaceae Apiaceae Myrtaceae Euphorbiaceae Solanaceae Apiaceae Rosaceae Onagraceae Rubiaceae Geraniaceae Proteaceae Asteraceae

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Budleja araucana Phil. (pan˜il) Budleja globosa Hope (pan˜il, matico) Caiophora patagonica (Speg.) Urban et Gilg. (ortiga) Calceolaria filicaulis Clos (cohete) Calendula officinalis L. (calendula) Callitriche lechleri (Hegelm.) Fasset (fen fen) Centaurium cachanlahuen (Molina) B. L. Rob. (cachanlahue) Cerastium arvense L. (nin˜a bonita) Cestrum parqui L’He´r. (palque huerta) Chenopodium album L. (quinhuilla) Chenopodium ambrosioides L. (paico macho) Chenopoudium multifidum L. (paico arrastrado) Chusquea culeou Desv. (colihue) Cichorium intybus L. (achicoria) Cirsium vulgare (Savi) Ten. (cardo negro) Conium maculatum L. (cicuta) Coriaria ruscifolia L. (deu) Cucurbita moschata Duch. (calabaza) Dasyphyllum diacanthoides (Less.) Cabrera (palo santo) Daucus carota L. (zanahoria) Diposis patagonica Skottsb. (yoco´n) Discaria spp. (chacai) Drymis winteri J.R.Forst. & G. Forst (canelo) Embothrium coccineum J.R. Forst. & G. Forst. (notro) Empetrum rubrum Vahl ex Willd. (mutilla) Ephedra ochreata Miers. (cuparra) Equisetum bogotense H.B.K. (limpia plata) Equisetum giganteum L. (cola de caballo) Erodium cicutarium (L.) L’He´rit. (alfilerillo) Eryngium paniculatum Cavanilles et Dom. (cupalla) Eucaliptus globulus Labill. (eucalipto) Euphorbia collina Phil. (pichoga) Fabiana imbricata Ruiz & Pav. (palo piche) Foeniculum vulgare Mill. (hinojo) Fragaria chiloensis (L.) Duchesne f. (frutilla) Fuchsia magellanica Lam. (chilco) Galium eriocarpum Bartling ex DC (yerba del pollo) Geranium chilense Aedo & Mun˜oz Garm. (core-core) Gevuina avellana Mol. (avellano) Gnaphalium andicola Phil. (vira vira)

704

Table 1 (continued ) Life form

Plant part

Category of use

Community

Ecological environment

Origin

Gnaphalium molleri Phil. (santa maria) Grindelia chiloensis (Corn.) Cabrera (boto´n de oro) Gunnera tinctoria (Mol.) Mirbel (nalca) Haplopapus glutinosus Cassini (baylahuen) Hipericum perforatum L. (yerba de San Juan) Hypochoeris radicata L. Juncus balticus Willd. var montanus (mallin) Lactuca serriola L. (lechuga salvaje) Larrea spp. (jarilla) Laurus nobilis L. (laurel) Lecanophora ameghinoi (Speg.) Speg. (huella) Libertia sp. (trique-trique) Lomatia hirsuta Lam. Diels ex J.F. Macbr. (radal) Luzula correae Barros (San Juanito) Maihueniopsis darwinii var. hickenii (Brit.) Kiesling (chupa sangre) Malus sp. Malus sylvestris Mill. (manzana silvestre) Malva neglecta Wallroth (malva arrastrada) Margiricarpus pinnatus (Lam.) Kuntze (yerba de la perdiz, perlilla) Marrubium vulgare L. (malvarrubia) Matricaria matricarioides (Less.) Porter ex Britton (manzanilla) Maytenus boaria Mol. (maite´n) Melissa officinalis L. (toronjil) Mentha pulegium L. (poleo) Mentha rotundifolia (L.) Huds. (yerba buena) Mentha sp. Mentha spicata L. (yerba buena) Mentha x piperita L. (menta negra) Mimulus glabratus Kunth (placa) Muehlenbeckia hastulata (Sm.) Johnst. (quineo) Mulinum spinosum (Cav.) Pers. (neneo) Mutisia decurrens Cav. (mutisia) Nassauvia axilaris (Lag.) Don hol(un˜a de gato) Nothofagus dombeyi (Mirb.) Oerst. (coihue)

Asteraceae Asteraceae Gunneraceae Asteraceae Hypericaceae Asteraceae Juncaceae Asteraceae Zygophylaceae Lauraceae Malvaceae Iridaceae Proteaceae Juncaceae Cactaceae Rosaceae Rosaceae Malvaceae Rosaceae Lamiaceae Asteraceae Celastraceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Lamiaceae Scrophulariaceae Polygonaceae Apiaceae Asteraceae Asteraceae Fagaceae

Herb Shrub Herb Shrub Herb Herb Shrub Herb Shrub Tree Shrub Herb Tree Shrub Herb Tree Tree Herb Shrub Herb Herb Tree Herb Herb Herb Herb Herb Herb Herb Shrub Shrub Herb Herb Tree

2 2 1,2 2 2 2 1 2 2 2 2 1 2 1 1 2,3 3 2 2 2 2 2 2 2 2 2 2 2 3 2 1 2 1 2

Medicinal Medicinal Both Medicinal Medicinal Both Medicinal Both Medicinal Medicinal Medicinal Medicinal Medicinal Medicinal Both Both Edible Medicinal Medicinal Medicinal Medicinal Medicinal Medicinal Medicinal Medicinal Medicinal Medicinal Both Edible Both Medicinal Medicinal Medicinal Medicinal

Ca Ca Ca–Cu Ca Cu Cu Ca Cu Ca–Cu Ca Ca Cu Ca–Cu Ca Ca Cu Ca Ca–Cu Ca–Cu Ca–Cu Ca–Cu Ca–Cu Ca–Cu Ca–Cu Ca–Cu Cu Ca Ca Ca Ca Ca–Cu Cu Ca Cu

Steppe Steppe Forest Forest Anthropic Anthropic Steppe Anthropic Steppe Anthropic Steppe Steppe Forest Steppe Steppe Forest Forest Anthropic Steppe Anthropic Anthropic Anthropic Anthropic Anthropic Anthropic Anthropic Anthropic Anthropic Steppe Steppe Steppe Forest Steppe Forest

N N N N E E N E N E N N N N N E E E N E E N E E E E E E N N N N N N

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2 2 2 2 2 2 2 1 2 2 2 3 3 3 2 1 1 2,3 3 2,3 2 3 2 2 3 2 2 2 2 2 2 2 2 2,3 2 1 1 2 1 2

Medicinal Medicinal Medicinal Medicinal Medicinal Edible Both Both Medicinal Medicinal Medicinal Edible Edible Edible Medicinal Edible Medicinal Both Edible Both Both Both Medicinal Medicinal Edible Edible Medicinal Medicinal Medicinal Medicinal Medicinal Edible Medicinal Both Medicinal Medicinal Medicinal Medicinal Both Medicinal

Cu Ca–Cu Ca–Cu Cu Ca–Cu Ca–Cu Cu Ca Ca–Cu Ca–Cu Ca–Cu Cu Cu Cu Ca–Cu Cu Ca Ca Cu Cu Ca–Cu Cu Ca–Cu Ca–Cu Cu Ca–Cu Ca–Cu Ca–Cu Ca Ca–Cu Ca–Cu Ca–Cu Ca Ca Ca Ca Ca Cu Cu Ca

Forest Anthropic Forest Forest Steppe Forest Anthropic Steppe Anthropic Anthropic Anthropic Anthropic Anthropic Anthropic Forest Anthropic Steppe Steppe Anthropic Forest Steppe Anthropic Anthropic Anthropic Anthropic Anthropic Anthropic Anthropic Steppe Anthropic Forest Forest Steppe Steppe Steppe Steppe Steppe Forest Anthropic Steppe

N E N N N N E N E E E E E E N E N N N N E E E E E E E E N E E N N N N N N N E N

705

Tree Herb Herb Shrub Herb Herb Herb Shrub Herb Herb Herb Tree Tree Tree Herb Herb Herb Shrub Shrub Shrub Herb Shrub Shrub Shrub Shrub Herb Herb Shrub Shrub Shrub Tree Herb Shrub Shrub Herb Herb Herb Shrub Herb Herb

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Fagaceae Lamiaceae Apiaceae Thymelaceae Oxalidaceae Oxalidaceae Apiaceae Poaceae Plantaginaceae Plantaginaceae Polygonaceae Rosaceae Rosaceae Rosaceae Santalaceae Crucifereae Amaryllidaceae Saxifragaceae Saxifragaceae Saxifragaceae Crucifereae Rosaceae Rosaceae Lamiaceae Rosaceae Polygonaceae Polygonaceae Rutaceae Salicaceae Lamiaceae Caprifoliaceae Apiaceae Anacardiaceae Anacardiaceae Asteraceae Iridaceae Iridaceae Solanaceae Solanaceae Asteraceae

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Notopahgus obliqua (Mirb.) Oerst. (roble pellı´ n) Origanum vulgare L. (ore´gano) Osmorhiza chilensis Hook & Arn. (cacho de cabra) Ovidia andina (Poepp. & Endl.) Meisn. (pillo-pillo) Oxalis adenophylla Gill. ex Hook & Arn. (culle colorado) Oxalis valdiviensis Barneoud (culle, vinagrillo) Petroselinum sativum Hoffm. (peregil) Phragmites australis (Cav.) Trin. ex Steud. (carrizo) Plantago lanceolata L. (siete venas) Plantago major L. (llante´n) Polygonum aviculare L. (sanguinaria) Prunus cerasus L. (guindo) Prunus domestica L. (ciruelo) Prunus sp. Quinchamalium chilense Mol. (quinchamalı´ ) Raphanus sativus L. (nabo silvestre) Rhodophiala elwessii (C. H. Wright) Traub (cebolleta, ajo del diablo) Ribes cucullatum Hook. & Arn. (parrillita) Ribes grossularia L. (grosella) Ribes magellanicum Poir. (parrilla) Rorippa nasturtium-aquiaticum (L.) Hayek (berro) Rosa rubiginosa L. (rosa mosqueta) Rosa sp. (rosa) Rosmarinus officinalis L. (romero) Rubus idaeus L. (frambuesa) Rumex acetosella L. (acederilla) Rumex crispus L. (romasa) Ruta graveolens L. (ruda) Salix fragilis L. (sauce) Salvia officinalis L. (salvia) Sambucus nigra L. (sauco) Sanicula graveolens Poepp. ex D.C. (cilantro silvestre) Schinus johnstonii Barkley (molle blanco) Schinus o’donelli Barkley (molle) Senecio filaginoides DC. (mata mora) Sisyrinchium arenarium Poepp. (chaleo rosado) Sisyrinchium sp. (triqui trique) Solanum crispum Ruiz & Pav. (natre) Solanum tuberosum L. (papa) Solidago chilensis Meyen (vara amarilla)

706

Table 1 (continued ) Life form

Plant part

Category of use

Community

Ecological environment

Origin

Stellaria media (L.) Villars. (capiquı´ , quilloi-quilloi) Tanacetum balsamita L. (yerba San Pedro) Taraxacum officinale Weber (diente de leo´n) Thymus vulgaris L. (tomillo) Tragopogon dubuis Scop. (salsifı´ ) Tristagma patagonicum (Bak.) Traub. (chaleo) Tristeryx corymbosus (L.) Kuijt (quintral) Urtica urens L. (ortiga chica) Valeriana carnosa Sm. (n˜anculahuen) Verbascum thapsus L. (tabaco de indio) Viola maculata Cav. (violeta amarilla) Xanthium spinosum L. (cepa caballo)

Caryophyllaceae Asteraceae Asteraceae Lamiaceae Asteraceae Liliaceae Loranthaceae Urticaceae Valeraniaceae Scrophulariaceae Violaceae Asteraceae

Herb Herb Herb Shrub Herb Herb Shrub Herb Herb Herb Herb Herb

2 2 2 2 3 1 2 2 1 2 2 2

Medicinal Medicinal Both Medicinal Medicinal Both Medicinal Medicinal Medicinal Medicinal Medicinal Medicinal

Ca–Cu Ca–Cu Ca–Cu Ca–Cu Ca Ca Cu Ca–Cu Ca–Cu Ca Cu Ca

Anthropic Anthropic Anthropic Anthropic Steppe Steppe Forest Anthropic Forest Steppe Forest Steppe

E E E E E N N E N E N E

Life form: (Herb) non-woody plants as annuals, ferns, grasses and vines, (Shrub): woody multi-stemmed plants less than 5 m tall, (Tree): single-stemmed woody plants greater than 5 m. Plant part: (1) roots, ryzomes and bulbs, (2) leaves, stems and flowers, (3) fruits and seeds. Ecological environment: (Anthropic) created by humans as orchard, fields, corrals, roads around dwellings, (Forest) Andean temperate forests dominated by Austrocedrus chilensis and deciduous Nothofagus forests, (Steppe): driest areas dominated by herbs and shrubs and with ‘‘mallines’’, humid meadows used for cattle raising. Origin: (N) native, (E) exotic.

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species utilized for both purposes, edible and medicinal, 35% (Table 1). As hypothesized, this dissimilarity can be mainly attributed to plant richness differences given that both communities are settled in distinct ecological environments. In contrast, both communities share a total of 63% of useful plant families (Jaccard index). Within the edible plants, Rosaceae, Araucanaceae, Berberidaceae, Asteraceae and Cruciferae are the most frequently used families (60% in common). Regarding the medicinal species, they mainly use Asteraceae, Lamiaceae and Rosaceae families (54% in common), and in terms of edible and medicinal purposes, Apiaceae, Asteraceae, Rosaceae and Oxalidaceae are mainly utilized (83% in common, Table 1). This pattern shows a significant similarity of families selected for medicinal and edible purposes in both communities. Accordingly, Moerman (1996) and Leonti et al. (2003) found striking similarities in the selection of these medicinal families (e.g. Asteraceae, Lamiaceae, Apiaceae, etc.), when they compared plant use in widely separated human populations. They suggested the existence of a global pattern of wild plant knowledge. However, when comparing the utilization of edible and medicinal families in each community, we found, by means of the Jaccard test, only 33% in common in Cayulef and 27% in Curruhuinca, showing a clear distinction at this level of analysis. Gottlieb (1982) and Gottlieb et al. (2002) have attributed these differences to distinct chemo-taxonomical origins of the families used by Brazilian aboriginal communities. This author has proposed that people select a plant for each purpose, food or medicine, considering differences in secondary chemical compounds particular to each family that lead to distinct patterns of use. 5.2. Environments and plant use categories Independently from where they are settled, both Mapuche communities use three gathering environments: the forest, the steppe, and anthropic or secondary environments. This finding shows an ample exploration of ecological gathering sites, as found in other Mapuche communities (Ladio and Lozada, 2000, 2004a, b), as well as other region of the world (Frei et al., 2000; Benz et al., 1994). In accordance with our second hypothesis, gathering habitats were utilized differently by the two populations (w2 , po0.05). The Cayulef people utilized more species belonging to the steppe, while the Curruhuinca people utilized twice as many species from the forest and their dwellings (Fig. 2a and b). This pattern highlights the relative importance the environment plays in their present plant use, in comparison with their cultural tradition. Moreover, this selection pattern could also be related to the hypothesis which links plant use with its availability, which offers more possibilities for local people to experiment with these resources (Phillips and Gentry, 1993). Our study was based on plant richness, a direct way to analyze plant use and species diversity, however, use value and cultural importance indices among the informants were not measured. The results also show the relevant role that anthropic environments have on useful resource gathering in both communities (Fig. 2a and b), as found in other studies (Voeks, 1996; Bennett and Prance, 2000; Voeks, 2004; Stepp, 2004; Almeida et al., 2005). When considering the category of use of plants gathered in each environment, differential selection patterns between both communities were observed. In Cayulef, the most frequently used species for medicine or food were indistinctly gathered in the three environments (w2 , p40.05, Fig. 2a). In contrast, the Curruhuinca people collected more

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a

Cayulef- Steppe 60 Category 15 Both edible and medicinal

50

Medicinal

Species richness

Edible

40

30

38

4 26

20

4 13

10

0

3

4

3

Anthropic

Steppe

Forest

b

Curruhuinca- Forest 60 Category 17

50

Both edible and medicinal

Species richness

Medicinal

40

Edible

10

28

30

25

20 15

10 9 5

0 Anthropic

Steppe

Forest

Fig. 2. Utilized plants in the Cayulef (a) and Curruhuinca (b) communities according to gathering environments and category of use.

edible and medicinal species both from disturbed areas and from the forest and significantly fewer species from the steppe, particularly for food use (w2 , po0.05). It is interesting to note the high proportion of utilized medicinal plants from anthropic

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environments. This fact not only agrees with many other studies (e.g. Voeks, 2004; Stepp, 2004), but it might also be related to a greater acculturation process in this community, settled near an urban center, in comparison with the more isolated Cayulef population. Accordingly, when indigenous people become more integrated into the market economy, wild plant use is endangered (Reyes-Garcı´ a et al., 2005). In addition, for the Curruhuinca people, the steppe was mainly used to collect medicinal plants (Fig. 2b), indicating that this arid environment, although distant, still offers important medicinal resources that cannot be replaced by forest or anthropic species. The relevance of drylands for collecting medicinal resources has also been described in other arid zones (Almeida et al., 2005; Albuquerque and Paiva de Lucena, 2005). 5.3. Environments and biogeographical origin of selected plants Considering the biogeographical origin of all the utilized plants, the Cayulef people tend to use more native plants and less exotic ones than the Curruhuinca people, although these differences reached a marginal level of significance (w2 , p ¼ 0:07). This tendency could be due to the conspicuous western influence that the Curruhuinca people receive, as mentioned above, as well as the historic presence of introduced species in the Patagonian region (Ezcurra and Brion, 2005). For both communities, exotic species use was principally related to anthropic environments (w2 , po0.05), as in other Mapuche communities (Ladio and Lozada, 2000, 2001) and as in many indigenous and rural populations worldwide (Bennett and Prance, 2000; Stepp, 2004; Alburquerque et al., 2005). Particularly, in the Curruhuinca population, exotic plant richness from disturbed environments was twice as much as in the Cayulef community (Fig. 3a and b). In contrast, both communities use more native species from the steppe and the forest (po0.05, Fig. 3a and b). However, the Cayulef people use significantly more native plants from the steppe than the Curruhuinca population, whereas the latter uses more than twice as many native forest species (w2 , po0.05, Fig. 3a). This result shows that native plant use is particularly influenced by the type of environment each community inhabits and the resource availability hypothesis (Phillips and Gentry, 1993; Almeida et al., 2005). The Andean forest presents a greater abundance and richness of useful species than the steppe given its precipitation gradient. For example, the forest offers more than 3 tons/ha of edible resources, whereas the steppe around 0.5 ton/ha (Dı´ az-Betancourt et al., 1999; Ladio, 2002). 5.4. Life forms and plant biogeographical origin Both populations utilize more herbs than bushes or trees for medicinal and edible purposes, irrespective of their home environment (w2 , po0.05). Diverse studies have shown the importance that herbs have as nutrient contributors of carotene, vitamin C, calcium and iron (Nordeide et al., 1996; Gonza´lez et al., 2003) as well as medicinal components, particularly in tropical environments (Voeks, 1996; Voeks, 2004). However, in relation to this latter medicinal use, a different pattern was observed in a dry region (Almeida et al., 2005). Our findings are not in accordance with their results in which bushes and trees are more frequently used for medicinal purposes. The utilization of trees in both communities is restricted to a few species with traditional importance in the Mapuche culture, both for edible and medicinal purposes (e.g. Araucaria araucana (Mol.) C. Koch (Araucariaceae) seeds for food, M. boaria leaves as medicine, etc.). This trend coincides with other studies

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Cayulef - Steppe 60 Origin Native

Species richness

50

50

Exotic

40

30

32

20 18

10 7

0 Anthropic

b

Steppe

Forest

Curruhuinca - Forest 60 Origin Native

Species richness

50

51

Exotic

40 38

30

20 16

10

0

3

Anthropic

Steppe

Forest

Fig. 3. Biogeographical origin of plants used in Cayulef (a) and Curruhuinca (b) in relation to the different gathering environments.

which have shown that trees, restricted to primary forests, are only used to treat particular ailments (Caniago and Siebert, 1998). The Cayulef people seem to use more native herbs and shrubs (w2 , po0.05, Fig. 4a), and the Curruhuinca people more exotic herbs and native shrubs (w2 , po0.05, Fig. 4b). Accordingly, Almeida et al. (2005) have found that local people in Brazil prefer to use native shrubs in arid zones. Moreover, weeds from disturbed and secondary areas have been frequently mentioned in several ethnobotanical studies (Bennett and Prance, 2000;

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711

Cayulef - Steppe 80 Origin Exotic

33

Native

Species richness

60

40 5

36

29

20

4

0

Trees

b

Shrubs

Herbs

Currhuinca - Forest 70 Origin

60

39

Exotic Native

Species richness

50 40 30

8

20 10

24

24

Shrubs

Herbs

6 9

0 Trees

Fig. 4. Utilized plants in the Cayulef (a) and Curruhuinca (b) communities according to life forms and origin.

Stepp, 2004; Voeks, 2004). This pattern could also show the importance of weedy greens as useful and versatile resources, which were probably introduced as food at first and then, mostly employed in therapeutic usages (Bennett and Prance, 2000; Leonti et al., 2006). 5.5. Above-ground versus underground plant selection Both communities utilized similar above-ground or underground parts, either for medicinal or food use (w2 , p40.05). Irrespective of which species were selected, both communities utilized more aerial organs, i.e. leaves, bark and stems for medicinal purposes

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(w2 , p ¼ 0:28), and more leaves, fruit and seeds for food (w2 , p ¼ 0:29). It is interesting to note the high proportion of leaf use in both communities. These results might show a similar selection pattern in both communities regardless of their home environments supporting the evidence that bioactive substances are principally concentrated in leaves (Almeida et al., 2005; Stepp, 2004; Voeks, 2004). As was predicted in our eighth hypothesis, seed and fruit producing species were more frequently cited in the Curruhuinca community than in Cayulef (17 vs. 8) for edible use, although these differences reached marginal significances (Binomial test, p ¼ 0:054). This result shows a differential pattern of plant part used in relation to habitat selection, as proposed by Gragson (1997). 5.6. Niche breadth and complementary use The Cayulef people use a number of useful plants (100 spp.) similar to that used by the Curruhuinca dwellers (96 spp.), (Binomial test, p ¼ 0:83). This indicates that both communities have a similar niche breadth, a result that disagrees with our last hypothesis, probably due to the fact that most resources utilized by both communities were medicinal plants, while the niche breadth concept has been mostly applied for food selectivity. However, when we considered only the edible species, no significant differences were found (Binomial test, p ¼ 0:424). In addition, the Cayulef population has a similar proportion of complementary use of edible and medicinal plants (70%) with respect to the Curruhuinca community (64%), (Binomial test, p ¼ 0:39). This result also diverges from our hypothesis, and could be related to the eroding process that wild plant knowledge and use is presently suffering in the Mapuche communites of Northwestern Patagonia, Argentina (Ladio and Lozada, 2000, 2003, 2004a, b). Selection patterns are sensitive (vulnerable) to the decay of wild plant richness known by the people and therefore this might explain the deviation from expected gathering patterns. Considering all plants listed by both Mapuche communities, food plants significantly overlap with medicinal species; indeed, 65% of the total plants identified as food were also cited for medicinal purposes. However, these plant species could be used in different ways according to different contexts, i.e. diverse attributes of a plant can be defined depending on the context in which it is utilized, as has been proposed by Etkin (1994) and Etkin and Ross (1982). In a therapeutic context, when a plant is explicitly ‘‘non-food’’, a particular plant part is used, and is ingested alone, or with other plants as part of a composite medicine, or added to the food of ailing individual. If the same plant in a dietary context is explicitly ‘‘non-medicinal’’, it is part of a meal and is consumed by all members of the family group. For a thorough understanding of the significance of this characteristic, further studies on wild plant use should be conducted focusing how this motivational aspect might influence the selection pattern of each plant in a certain context. 6. Conclusions The main results of this study show that the environment is a fundamental factor influencing the selection of plant species. However, it is interesting to note the many shared selection patterns that are found in both communities, for example at the level of plant families, ways of utilization, plant part and life forms utilized. It is well known how culture influences people’s values, perceptions and patterns of action (e.g. Cole, 1999). Accordingly, the observed similarities must be related to their shared cultural background.

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Despite the prevalent acculturation process these communities are enduring, the described gathering patterns are still preserved as important components of their traditional Mapuche heritage. In both Mapuche populations, the use of wild plants is primarily based on the utilization of medicinal, native and exotic species belonging to their home surroundings, showing a conspicuous environmental and anthropogenic influence. Both communities use native and exotic plants in a similar proportion. This fact might also indicate that the Mapuche populations utilize exotic species regardless of which environment they inhabit. The Mapuche people have an extensive history of use of invasive exotic species (e.g. Kutschker et al., 2002; Estomba et al., 2006). At present, traditional plant remedies used by both rural and suburban Mapuche communities are suffering a syncretic process due to the prevailing occidental medicinal practices (Citarella, 1995). A thorough documentation of this ancestral wild plant wisdom might help diminish the eroding process of native flora knowledge in these aboriginal communities. Acknowledgments This research was supported by the Consejo Nacional de Investigaciones Cientı´ ficas y Te´cnicas, the Universidad Nacional del Comahue of Argentina, and the FONCYT (grant PICT #: 03593). We are thankful for the revisions of the English manuscript by Andy Schwartz and Damasia Lozada. We also wish to thank to two anonymous reviewers for their valuable comments and suggestions on the manuscript. Special gratitude is expressed to the families from Curruhuinca and Cayulef for their kind hospitality. References Agresti, A., 1996. An Introduction to Categorical Data Analysis. Wiley, New York. Aizen, M., Ezcurra, C., 1998. High incidence of plant–animal mutualisms in the woody flora of the temperate forest of southern South America: biogeographical origin and present ecological significance. Ecologı´ a Austral 8, 217–236. Albuquerque, U.P., Paiva de Lucena, R.F., 2004. Me´todos e Te´cnicas na Pesquisa Etnobota´nica. Nupeea, Recife, Brazil. Albuquerque, U.P., Paiva de Lucena, R.F., 2005. Can apparency affect the use of plants by local people in tropical forest? Interciencia 30 (8), 506–511. Alburquerque, U.P., Cavalcanti Andrade, L., Oliveira de Silva, A.C., 2005. Use of plant resources in seasonal dry forest (Northeastern Brazil). Acta Botanica Brassilica 19 (1), 27–38. Alexiades, M.N., 1996. Selected Guidelines for Ethnobotanical Reserach: A Field Manual. The New York Botanical Garden, Bronx, New York. Almeida, C.F.C.B.R., Lima e Silva, T.C., de Amorin, E.L.C., Maia, M.B.S., Albuquerque, U.P., 2005. Life strategy and chemical compostion as predictors of the selection of medicinal plants from the Caatinga (Northeast Brazil). Journal of Arid Environments 62, 127–142. Barros, V., Cordon, V., Moyano, C., Mendez, R., Forguera, J., Picio, O., 1983. Cartas de precipitacio´n de la zona oeste de las provincias de Rı´ o Negro y Neuque´n. Primera contribucio´n. U.N.C. CONICET. Facultad de Ciencias Agrarias, Cinco Saltos. Bennett, B.C., Prance, G.T., 2000. Introduced plants in the indigenous pharmacopoeia of Northern South America. Economic Botany 54 (1), 90–120. Benz, B.F., Santana, F.M., Pineda, R.L., Cevallos, J.E., Robles, L.H., de Niz, D.L., 1994. Characterization of mestizo plant use in the Sierra de Manantlan, Jalisco-Colima, Mexico. Journal of Ethnobiology 14 (1), 23–41. Cabrera, A.L., 1976. Regiones Fitogeogra´ficas. Argentinas. Fac. 1. Enciclopedia Arg. de la Agricultura y Jardinerı´ a. Tomo II. Editorial Acme S.A.C.I., Buenos Aires.

ARTICLE IN PRESS 714

A. Ladio et al. / Journal of Arid Environments 69 (2007) 695–715

Cabrera, A.L., Willink, A., 1980. Biogeografı´ a de Ame´rica Latina. Monografı´ a no. 13. Washington, DC. Secretarı´ a General de la Organizacio´n de los Estados Americanos, Washington, DC. Caniago, I., Siebert, S., 1998. Medicinal plant ecology, knowledge, and conservation in Kalimantan, Indonesia. Economic Botany 52 (3), 229–250. Castro, F., Begossi, A., 1996. Fishing at Rio Grande (Brazil): ecological niche and competition. Human Ecology 24 (3), 401–411. Citarella, L., 1995. Medicinas y culturas en la Araucanı´ a. Editorial Sudamericana, Santiago de Chile. Cole, M., 1999. Psicologı´ a Cultural. Una disciplina del pasado y del futuro. Ediciones Morata, Madrid. Correa, M.N., 1969, 1971, 1978, 1984, 1988, 1998 and 1999. Flora Patago´nica. Partes 1,2,3,4,5, 6 y 7. Coleccio´n Cientı´ fica del Instituto Nacional de Tecnologı´ a Agropecuaria. Bs. As. Dı´ az-Betancourt, M.E., Ghermandi, L., Ladio, A.H., Lo´pez Moreno, I.R., Raffaele, E., Rapoport, E.H., 1999. Weeds as a source for human consumption. A comparison between tropical and temperate Latin America. Revista Biologı´ a Tropical 47 (3), 329–338. Edwards, S., Nebel, S., Heinrich, M., 2005. Questionnaire surveys: methodological and epistemological problems for field-based ethnopharmacologists. Journal of Ethnopharmacology 100, 30–36. Estomba, D., Ladio, A., Lozada, M., 2006. Medicinal wild plant knowledge and gathering patterns in a Mapuche community from North-western Patagonia. Journal of Ethnopharmacology 103, 109–119. Etkin, N.L., 1994. Consuming a therapeutic landscape: a multicontextual framework for assessing the health significance of human-plant interactions. In: Flagler, J., Poincelot, R.G. (Eds.), People—Plant Relationships: Setting Research Priorities. Haworth Press, pp. 61–81. Etkin, N.L., Ross, P., 1982. Food as medicine and medicine as food. An adaptative framework for the interpretation of plant utilization among the Hausa of Northern Nigeria. Social Science and Medicine 16, 1559–1573. Ezcurra, C., Brion, C., 2005. Plantas del Nahuel Huapi. Cata´logo de la Flora Vascular del Parque Nacional Nahuel Huapi, Argentina. Universidad Nacional del Comahue, Red Latinoamericana de Bota´nica, S.C. de Bariloche, Argentina. Falaschi, C.O., 1996. Defensa y reivindicacio´n de tierras indı´ genas. Informe final proyecto especial de Investigacio´n y Extensio´n. Universidad Nacional del Comahue y Asociacio´n por los derechos humanos, Neuque´n, Argentina. Frei, B., Sticher, O., Heinrich, M., 2000. Zapotec and Mixe use of tropical habitats for securing medicinal plants in Mexico. Economic Botany 54 (1), 73–81. Galeano, G., 2000. Forest use at the Pacific Coast of Choco, Colombia: a quantitative approach. Economic Botany 54 (3), 358–376. Golluscio, R.A., Deregibus, V.A., Paruelo, J.M., 1998. Sustainability and range management in the Patagonian steppes. Ecologı´ a Austral 8, 265–284. Gonza´lez, A., Janke, R., Rapoport, E., 2003. Valor nutricional de las malezas comestibles. Ciencia Hoy 13 (76), 41–47. Gottlieb, O., 1982. Ethnopharmacology versus chemosystematics in the search for biologically active principles in plants. Journal of Ethnopharmacology 6, 227–238. Gottlieb, O., Borin, M.R., de, M.B., de Brito, N.R.S., 2002. Integration of ethnobotany and phytochemistry: dream or reality? Phytochemistry 60, 145–152. Gragson, T.L., 1993. Human foraging in lowland South America: patterns and process of resource procurement. Research in Economic Anthropology 14, 107–138. Gragson, T.L., 1997. The use of underground plant organs and its relation to habitat selection among the Pume´ Indians of Venezuela. Economic Botany 51 (4), 377–384. Hardesty, D.L., 1975. The niche concept: suggestions for its use in human ecology. Human Ecology 3, 71–85. Ho¨ft, M., Barik, S.K., Lykke, A.M., 1999. Quantitative Ethnobotany. Applications of multivariate and statistical analyses in ethnobotany. People and Plant Working Paper. Kutschker, A., Menoyo, H., Hechem, V., 2002. Plantas Medicinales de Uso popular en comunidades del Oeste de Chubut INTA. Esquel. GTZ. Bariloche, Argentina. Ladio, A.H., 2002. Las plantas comestibles en el noroeste patago´nico y su utilizacio´n por las poblaciones humanas: una aproximacio´n cuantitativa Tesis doctoral. Universidad Nacional del Comahue, Bariloche, Argentina. Ladio, A.H., 2006. Uso y conservacio´n de plantas silvestres con o´rganos subterra´neos comestibles en comunidades Mapuche de la estepa patago´nica argentina. In: Alburquerque, U.P., Andrade Maris, J.F.,

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Almeida, C.B.R. (Eds.), To´picos em conservacˆa˘o e etnobota˘nica de plantas comestı´ veis. Recife, Brazil, pp. 53–72 (in press). Ladio, A.H., Lozada, M., 2000. Edible wild plant use in a Mapuche community of northwestern Patagonia. Human Ecology 28 (1), 53–71. Ladio, A.H., Lozada, M., 2001. Non-timber forest product use in two human populations from NW Patagonia: a quantitative approach. Human Ecology 29 (4), 367–380. Ladio, A.H., Lozada, M., 2003. Comparison of wild edible plant diversity and foraging strategies in two aboriginal communities of northwestern Patagonia. Biodivesity and Conservation 12, 937–951. Ladio, A.H., Lozada, M., 2004a. Patterns of use and knowledge of wild edible plants in distinct ecological environments: a case study of a Mapuche community from Nothwestern Patagonia. Biodiversity and Conservation 13, 1153–1173. Ladio, A.H., Lozada, M., 2004b. Summer cattle transhumance and wild edible plant gathering in a Mapuche community of NW Patagonia. Human Ecology 32 (2), 225–240. Ladio, A.H., Rapoport, E.H., 2002. Variacio´n estacional de la biomasa y riqueza de plantas silvestres comestibles en baldı´ os suburbanos de Bariloche (Parque Nacional Nahuel Huapi, Patagonia, Argentina). Vida Silvestre Neotropical 11 (1–2), 33–41. Leonti, M.S., Ramirez, R., Sticher, O., Heinrich, M., 2003. Medicinal flora of the Popoluca, Mexico: a botanical systematical perspective. Economic Botany 57 (2), 218–230. Leonti, M.S., Nebel, S., Rivera, D., Heinrich, M., 2006. Wild gathered food plants in the European Mediterranean: a comparative analysis. Economic Botany 60 (2), 130–142. Moerman, D.E., 1996. An analysis of the food plants and drug plants of native North America. Journal of Ethnopharmacology 52, 1–22. Nacuzzi, L.R., Pe´rez de Micou, C., 1984. Los recursos vegetales de los cazadores de la Cuenca del Rı´ o Chubut. Cuadernos del Instituto Nacional de Antropologı´ a 10, 407–423. Nolan, J., 1998. The roots of tradition: social ecology, cultural geography, and medicinal plant knowledge in the Ozark-Ouachita highlands. Journal of Ethnobiology 18 (2), 249–269. Nordeide, M.B., Hatløy, A., Følling, M., Lied, E., Oshaug, A., 1996. Nutrient composition and nutritional importance of green leaves and wild food resources in an agricultural district, Koutiala, in Southern Mali. International Journal of Food Sciences and Nutrition 47, 455–468. Ogle, B.M., Tuyet, H.T., Duyet, H.N., Dung, N.N.X., 2003. Food, feed or medicine: the multiple functions of edible wild plants in Vietnam. Economic Botany 57 (1), 103–117. Phillips, O.L.B., Gentry, A.H., 1993. The useful plants of Tamborata, Peru´. I: statistical hypothesis tests with a new quantitative techniques. Economic Botany 47, 15–32. Rapoport, E.H., Drausal, B.S., 2001. Edible plants. In: Levin, S. (Ed.), Encyclopaedia of Biodiversity, vol. 2. Academic Press, New York, USA, pp. 375–382. Rapoport, E.H., Ladio, A.H., Sanz, E., 1999. Plantas nativas comestibles de la Patagonia andina argentinochilena. Parte I. Ediciones Alternatura. National Geographic Society, Bariloche, Argentina. Rapoport, E.H., Ladio, A.H., Sanz, E., 2003. Plantas nativas comestibles de la Patagonia andina argentinochilena. Parte II. Ediciones Alternatura. Secretarı´ a de Extensio´n Universitaria. UNC. Fundacio´n Antorchas. Bariloche, Argentina. Reyes-Garcı´ a, V., Vadez, V., Huanta, T., Leonard, W., Wilkie, D., 2005. Knowledge and consumption of wild plants: a comparative study in two Tsimane’s villages in the Bolivian Amazon. Ethnobotany Research and Applications 3, 201–207. Smith-Ramı´ rez, C., 1996. Algunos Usos Indı´ genas Tradicionales de la Flora del Bosque Templado. In: Armesto, J., Villagra´n, C., Arroyo, M.K. (Eds.), Ecologı´ a de los bosques nativos de Chile. Editorial Universitaria, Santiago, Chile, pp. 389–404. Stepp, J.R., 2004. The role of weeds as sources of pharmaceuticals. Journal of Ethnopharmacology 92, 163–166. Stepp, J.R., Moerman, D., 2001. The importance of weeds in ethnopharmacology. Journal of Ethnopharmacology 75, 19–23. Veblen, T.T., Mermoz, M., Martı´ n, C., Kitzberger, T., 1992. Ecological impacts of introduced animals in Nahuel Huapi National Park, Argentina. Conservation Biology 6, 71–83. Voeks, R.A., 1996. Tropical forest healers and habitat preference. Economic Botany 50, 381–400. Voeks, R.A., 2004. Disturbance Pharmacopoeias: medicine and myth from the Humid Tropics. Annals of the Association of American Geographers 94 (4), 868–888.