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Journal of Ethnopharmacology 122 (2009) 251–260

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Journal of Ethnopharmacology

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Ethnobotanical review of the Mapuche medicinal flora: Use patterns on aregional scale

Soledad Molares, Ana Ladio ∗

INIBIOMA, Universidad Nacional del Comahue, Quintral, 1250-8400 San Carlos de Bariloche, Río Negro, Argentina

a r t i c l e i n f o

Article history:Received 6 October 2008Received in revised form 18 December 2008Accepted 4 January 2009Available online 8 January 2009

Keywords:PatagoniaMapucheKnowledge and use patterns of medicinalplants

a b s t r a c t

Aim of the study: This paper aimed to present a quantitative review of information on Mapuche ethnob-otany published for Argentina and Chile in the period 1955–2007.Materials and methods: Sixteen ethnobotanical articles were studied quantitatively by utilizing ethnob-otanical indices, non-parametric and multivariate tests.Results: A total of 505 medicinal species, 304 native (60%) and 196 exotic (39%) were registered. Ailmentswere treated with both native and exotic species, although native showed higher values of consensus(between studies) than exotic, and were more frequently used in all cases. The most common ailmentswere gastro-intestinal pains (26%). Most cures used herbs (56%). Native species were obtained mainly bygathering in forests and ecotones (40%), and exotics by gathering in anthropogenic environments (28%).Our results demonstrate the existence of a common, shared body of knowledge of the medicinal flora ata regional level, integrating ancestral knowledge with foreign knowledge accumulated over time. Finally,reflecting cultural erosion, modern articles cited significantly fewer native plants than older articles; atrend not found with exotic species.Discussion and conclusion: The information offered can be used as a guide for future work on promisingspecies for health care, and as background information for the development of bio-cultural conservationstrategies.

© 2009 Elsevier Ireland Ltd. All rights reserved.

1. Introduction

The Mapuche medicinal flora is based on profound, an extensiveand diverse knowledge of local plant varieties (e.g. Gusinde, 1917;Mösbach, 1992; Citarella, 1995; Estomba et al., 2006). Its richnessstems from complex, multifactorial reasoning, making connectionsbetween the different natural plant properties (morphological, eco-logical and uses) from a specific perspective and conception of theworld which is not only unifying in terms of nature, but whichis also full of symbolism (Erize, 1989; Citarella, 1995; Villagrán,1998).

The Mapuche people evolved from the cultural fusion of dif-ferent communities which originally lived in the cold temperateforests in the south of the Andes (Mösbach, 1992). These peoplewere horticulturalists and gatherers from the woodlands (Mösbach,1992). After the sixteenth century, they suffered great social andeconomic changes, and, as a result of extending their lands towardsthe south and the east, the driest areas of Argentina became partof their territory (Ladio, 2006b; Ladio and Lozada, 2009). Theynow became mounted shepherds, in general, living far from their

∗ Corresponding author. Tel.: +54 2944 430949; fax: +54 2944 422111.E-mail address: aladio@crub.uncoma.edu.ar (A. Ladio).

ancestral lands, but the use of medicinal plants continued to be anessential feature in their subsistence (Estomba et al., 2006; Ladioet al., 2007; Ladio and Lozada, 2008). This knowledge has primarilybeen registered in the chronicles of the first settlers who, in turn,enriched it with their own knowledge of plants from Europe andother regions of the world (Citarella, 1995; Massardo and Rozzi,1996).

At the present time, the Mapuche are considered the mostnumerous indigenous nation in southern South America (Labaríasand Antiqueo, 2008). As with other indigenous societies in theworld, in recent years it has been observed that the Mapuchechildren and youth preserve significantly less in the way of plantknowledge and practices than adults and the elderly (Ladio andLozada, 2004a; Estomba et al., 2006). Of the many factors involvedin this process of loss, the most relevant are the loss of their nativelanguage, a moving away from their particular perception of theworld (Ladio, 2002; Ladio, 2006b), market influences, globalisation,uprooting from their native lands (Ladio and Lozada, 2001; Estombaet al., 2006), the lack of access to traditional gathering environments(Aagesen, 1998; Ladio and Lozada, 2004a,b; Ladio, 2006a), the aban-donment of horticultural practices (Ladio, 2002; Eyssartier et al., inpress) and cattle transhumance (Ladio and Lozada, 2004b); theselast two are traditionally associated with the gathering of medicinalplants.

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In this work, we analyse ethnobotanical information forArgentina and Chile during the period 1955–2007, in order toidentify patterns and analyse differences on a regional scale. Ourtheoretical base coincides with that proposed by Boster (1985),which shows that although plant knowledge is not uniformly dis-tributed within a culture, this does not make a coherent culturaldescription impossible. Moerman et al. (1999) have put forwarda hypothesis on the existence of a “global pattern of humanknowledge on medicinal plants”, which would relate mainly tothe characteristics of local flora, and then to the transmission ofknowledge throughout space and time from the Palaeolithic Era.Subsequently, Leonti et al. (2003) refer to the existence of “commonselection criteria”, related to organoleptic perceptions and othercommon cross-cultural criteria.

Bibliographical studies on indigenous medicinal floras like thatof the Mapuche, which have a wide geographical distribution andhave been submitted to cultural changes over time, are complexapproximations. However, if with this kind of comparative studyit is possible to identify common patterns in spite of the diffi-culties (e.g. Toledo et al., 1995; Begossi et al., 2002; Albuquerqueet al., 2008), there is no doubt that we are observing a commonbody of knowledge and practices of the highest ethnomedicinal andethnopharmacological value.

In this work, therefore, we have quantitatively characterized thebody of knowledge on medicinal plants which has accumulatedand been documented up to the present time, in botanical, ecolog-ical, biogeographical and cultural terms, as well as its irregularitiesthroughout this time. This involves work at a regional level, whichdiffers from the local scale, since it reflects properties and patternsnot observable on a small scale.

Our general working hypothesis is that in Mapuche plant use aregional pattern exists (due to the regional diversity of species) anda global one (common to other medicinal floras in the world), andtherefore we propose that:

(1) The Mapuche medicinal flora has selected and accumulateda wide variety of plant species, mostly native to Patagonia, and toa lesser extent, of exotic origin; (2) ethnobotanical studies revealsimilarities in species at a regional level, indicating a mass of sharedknowledge; (3) the native species used show high levels of consen-sus and use value, i.e., different ailments are treated with a singlespecies; (4) the main botanical families are similar to those belong-ing to the world medicinal flora. Just as in other medicinal floras,(5) the most common ailments are gastro-intestinal; (6) herbs andshrubs are the most important plant forms. Given the close con-tact of Mapuches in rural areas have with wild areas, it is alsoproposed that: (7) gathering in natural environments is the mainsource of supply, especially native woodlands and anthropogenicenvironments and to a lesser extent cultivation in home gardensand purchasing in markets. Finally, considering global processes ofcultural and biological diversity loss; (8) older studies will cite morespecies than modern ones.

1.1. Medical system: illness and medicinal plants in Mapuchecommunities

The traditional Mapuche medical system has suffered markedprocesses of syncretism and erosion (Citarella, 1995). Nevertheless,the concept of illness (“kutrán” en Mapuzungun) has not varied,and is intimately linked to the loss of balance between a per-son (“che”) and his surroundings (“mapu”). This imbalance canoccur on a physical, spiritual or psychological plane and is theresult of an inharmonious relationship with the surroundings, i.e.,imbalance with oneself, one’s family, one’s food, with the com-munity, the place where one lives, or with the different beingswhich form part of one’s cosmos (Citarella, 1995; Ladio and Lozada,2009).

This vulnerable state provokes the working of both naturaland supernatural forces, which cause illness (Citarella, 1995). Thecurative system includes the use of plants and other natural prod-ucts (mineral or animal), and specialists also have their place, forexample the “Machi”, a “shaman” responsible for establishing a con-nection with the protective spirits, who concentrates especially oncuring illnesses of a supernatural nature, and holds a position ofgreat prestige (Bacigalupo, 1995). In addition, specialists such as“parteras” (midwives), “yerbateros”, “meicas”, and “hueseros” cureillnesses caused by the effects of the cold, heat, food, knocks andfalls, plus some of magic origins.

We should highlight here the importance within Mapuchemedicine of home-based health care, practised by adults within thefamily (Idoyaga Molina, 2000). In addition, as in other rural popula-tions at the present time, other healing therapies are added to thesehome practices, which may be of a religious nature (evangelical andcatholic) or alternative medicine from cultural origins outside theregion (Idoyaga Molina, 2000). Official health care exists alongsidethese alternatives, its presence in rural areas depending on healthcentres and/or sporadic visits from doctors and nurses from thenearest hospitals (Arrue and Kalinsky, 1991).

2. Materials and methods

2.1. Study region

The articles analysed here are from studies carried out in ruralcommunities with Mapuche ancestry of Argentina and Chile, dis-tributed throughout an area extending approximately between 37◦

and 46◦ lat. S. These communities are established in different phyto-geographical provinces, namely: Patagonian, Monte, Central Chileand Sub-Antarctic (Fig. 1). However, the people do pass throughand gather plant species in environments where they are not set-tled, such as the High Andean (Fig. 1). The Patagonian and Monte(Argentina) provinces cover plateaus and low mountains, in dry,cold climates where annual rainfall varies between 100 and 270 mm(Fig. 1). In Patagonian province the dominant vegetation is shrubbyor grassy steppe, with bushes and/or species of cushion-plants(Fig. 1). The Monte vegetation consists of xerophytic, sammophyticand halophytic scrubs (Cabrera, 1971).

The Sub-Antarctic province (in both countries) extends overmountains and glacial valleys, in a humid, temperate climate, withannual rainfall of up to 2000 mm. The vegetation here is deciduousor evergreen forest with patches of, for example, meadows and peatbogs. In addition, in Chile, the Central Chilean province, which has aMediterranean type climate and annual rainfall of between 400 and900 mm, has vegetation which varies from woods of thorny speciesand open schlerophylls to spiny steppe scrub. The High Andeanprovince extends across the high border mountains, plateaus andrugged slopes, in a cold, dry climate, with vegetation consisting ofgrassy steppe or camephytes in cushion form (Cabrera, 1971). Thisphytogeographical heterogeneity implies a high degree of diversityof environments and microenvironments, ways of life, biologicalassociations and plant species.

The entire Argentine-Chilean Patagonia has been, and still is,subject to permanent human influence. During the past centuryactivities associated with forestry, cattle rearing and tourism havein some areas generated high levels of deforestation, contamina-tion and desertification (Paruelo and Aguilar, 2003; CONAF, 2008).Desertification in this region has caused disturbances of varyingdegrees and the progressive loss of ecological services (Paruelo andAguilar, 2003). The general result is a moving away from the naturalstate of the vegetation towards antropogenic environments withthe incorporation of exotic species, amongst other consequences(Rapoport and Brion, 1991; Correa, 1998; Ezcurra and Brion, 2005).

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Fig. 1. Study region including the most important ecological provinces of Argentinean and Chilean Patagonia.

2.2. Database construction

The analysis of Mapuche medicinal plants started with theconstruction of a database obtained by compiling studies on eth-nobotany. The studies were selected according to the followingcriteria: (1) the study must have been carried out in communi-ties with Mapuche ancestry in the southern and central region ofArgentina and Chile, (2) the species lists have arisen from eth-nobotanical fieldwork, i.e., not from bibliographical revision orcompilation, (3) only spermatophytes and pteridophytes were con-sidered, (4) work done by chroniclers, naturalists and/or expeditionmembers whose species identification is doubtful or difficult torelate to others was not included, (5) conference, seminar or sym-posium summaries or reports were not included due to fact thatthey generally do not contain completed species lists, (6) specieslists without information on uses, etc., were not included. Of atotal of 41 studies considered, 16 articles obeying these crite-ria and were included in the analysis as independent surveys(Martinez-Crovetto, 1980; San Martín, 1983; Villagrán et al., 1983;Houghton and Manby, 1985; Montecino and Conejeros, 1985; Mezaand Villagrán, 1991; Mösbach, 1992; Citarella, 1995; Conticello etal., 1997; Funes, 1999; Kutschker et al., 2002; Estomba et al., 2005,2006; Ladio, 2005, 2006a; Ladio et al., 2007).

We are confident that this selection is a representative sampleof the total scientific literature available about Mapuche medic-inal flora during the period under study because they includedboth indexed papers (in Biological Abstracts, Web of Science and

Scielo) as well as local studies that usually were not published ininternational journals.

2.3. The information obtained

The following types of information were obtained from thesearticles: number of species, common names, mapuzungun (indige-nous) names, medicinal uses and parts used. The next step was toput the species into different categories of analysis. According tohow the plants were obtained, we considered the ecological envi-ronment where they had been collected and the cultivation and/orpurchase of medicinal species in the following way: Sub-Antarcticforest and/or ecotone (F–E), Patagonian steppe and/or Monte (S–M),wild anthropogenic environment such as grazing land, road vergesand land around houses, etc. (A–E), wet habitats (WH), High Andeanenvironment (HA), and finally, gardens with cultivated species,and plants purchased from herbalists or traders (C–B). Categoriesaccording to growth form are as follows: “tree”, “shrub”, “climbingplant”, “parasite and hemiparasite” and “herb”. According to bio-geographical origin: exotic or native (native: those plants whichgrow in central and/or southern Argentina and Chile, below 35◦

lat. S.), this categorization is based on the scientific judgementassumed by the Zuloaga and Morrone catalogue (1999). And finally,use categories: gastro-intestinal disorders (GI), genito-urologicaldisorders (GU), pain and inflammation (PI), respiratory diseases(RS), dermatological diseases (DE), gynaecological-obstetric dis-eases (GO), circulatory and heart disorders (CH), nervous system

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disorders (NS), cultural syndromes (CS), fever (F), others (OT), notspecified (N).

All scientific names (considering that the study surveys spannedmore than 50 years) were updated by consulting specialists andspecialized databases: the Zuloaga and Morrone catalogue (1999) ofthe Darwinion Institute (www.darwinion.gov.ar), the InternationalPlant Names Index (www.ipni.org) and the Ezcurra and Brioncatalogue (2005). The complete species list is not presented here,due to its length; please consult authors for further information.However, Table 1 shows the main species used for the most commonailments in Patagonia.

2.4. Data analysis

Richness, diversity and biogeographical origin of Mapuchemedicinal flora: Richness (S), or the number of species recorded,and their diversity (H′) were calculated both in total and separately,according to biogeographical origin. Native and exotic species rich-ness was compared using the Chi-square test (p < 0.05). Diversitywas estimated with the Shannon–Wiener index (H′), which consid-ers the richness and relative proportions of each species, and thencompares them with the Student test (p < 0.05) (Ladio and Lozada,2003). Following Begossi (1996) and Ladio and Lozada (2003), thedegree of homogeneity of use of native and exotic species was com-pared using the Pielou eveness test [J′0–1 = H′/log(S)]. In addition,species richness was compared in total, and between natives andexotics, for each growth habit and methods of obtaining, using thebinomial test (p < 0.05). We also analysed whether for each methodof obtaining plant samples, results varied in the proportions of spe-cific growth habits. This comparison was also carried out using theChi-square test (p < 0.05) (Agresti, 1996).

2.4.1. Patterns of similarityThe similarity of named species between studies was analysed

using the Jaccard Index (JI) (Höft et al., 1999), which is based onthe presence or absence of species on each list. Relating the num-ber of species in common to the total number, it is expressed asJI = (c/a + b + c) × 100, where c is the number of species in common,a is the number of species unique to author A and b is the numberof species named solely by author B. In order to attain a global viewof these similarities, the MDS (Analysis of Euclidian distances on amultidimensional scale) was used. This multivariate technique pro-vided us with a spatial arrangement of all the studies carried out,showing the relative positions of all the components, according totheir degree of similarity (Hair et al., 1998). The proportion of vari-ation described by this arrangement was measured by the value ofR2 (which varies between 0 and 1), and by the stress estimating the

degree of adjustment of the distances created, whose values shouldbe less than 0.1 for a good adjustment (Hair et al., 1998). For bothanalyses, the work of Ladio (2005) was excluded, since it consideredexotic species only.

2.4.2. Patterns of consensus and use valueAs a measure of consensus between authors on cited species

and families, cite frequency was estimated for species and family,taking the percentage of authors citing each species (or family) withrespect to the total number of authors [(no. of authors who cite thesp. i (or family i) × 100)/total no. of authors].

In addition, the average number of uses for each species (valueindicating total use of each species) was calculated with the indexUVs = �UVis/ns (Phillips and Gentry, 1993), where UVis = the num-ber of uses registered by an author i for the species s, and ns = thenumber of authors who mention the species s.

The consensus in information relating to each use categorywas evaluated using the Heinrich et al. (1998) consensus factorFic〈0–1〉 = nur − nt/nur − 1, where nur = total number of reports of theuse category in question, and nt = total number of species used inthis category.

2.4.3. Patterns in timeThe Spearman correlation test was used to analyze the rela-

tionship between articles in terms of species richness and time(publication date, or for Mösbach, date work was finished: 1955),with a significance level of 0.05. The sign of the coefficient obtainedindicates the direction of the relationship (positive or negative), andthe value indicates the strength, with larger absolute values indicat-ing stronger relationships. The work of Ladio (2005) was excludedas it analysed only exotic species.

3. Results and discussion

3.1. Richness, diversity and biogeographical origin of medicinalspecies

The Mapuche medicinal flora holds a vast, rich body of knowl-edge learned and transmitted within the culture throughout spaceand time. This is amply demonstrated by the 505 species recorded,of which 304 (60%) are native and 196 (39%) exotic. Five species (1%of data) could not be analysed according to their origin, since theyhad only been classified to a generic level, belonging to taxa of widegeographical distribution.

In accordance with our first hypothesis, the sum of speciesrichness of native plants is significantly greater than that ofexotic species (�2 = 70.22, p < 0.001). The diversity of native species

Table 1Consensus factor for medicinal plants (Fic) (Heinrich et al., 1998), and most frequently cited species for each category of medicinal use indicated. References: nur = total numberof reports for the use category in consideration and nt = total number of species used.

Categories nur % nt % Fic Species most frequently cited

Gastro-intestinal disorders 512 26 271 54 0.47 Buddleja globosa Hope, Mentha × piperita L., Artemisia absinthium L.,Mentha pulegium L., Chenopodium ambrosioides L.

Dermatological diseases 219 11 129 25.5 0.41 Buddleja globosa, Plantago lanceolata L., Erodium cicutarium (L.)LıHér. ex Aiton

Genito-urological disorders 161 8 97 19 0.40 Fabiana imbricata Ruiz & Pav., Equisetum bogotense Kunth, Gunneratinctorea Molina (Mirb.)

Circulatory and heart disorders 150 7 94 19 0.38 Ribes magellanicum Poir., Gunnera tinctoreaRespiratory diseases 171 8 108 21 0.37 Centaurium cachanlahuen (Molina) B.L. Rob., Marrubium vulgare L.Pain and inflammation 252 12 159 31 0.37 Centaurium cachanlahuen, Stellaria media (L.) CirilloFever 117 6 82 16 0.30 Aristotelia chilensis (Molina) Stuntz, Centaurium cachanlahuenCultural syndromes 143 7 109 21.5 0.24 Drimys winteri J.R. Forst. & G. Forst.Others 135 7 105 21 0.22 Drimys winteri (toxic, anti-tumor), Stellaria media (ophthalmic,

refreshing), Conium maculatum L. (toxic, ophthalmic)Gynaecological-obstetric diseases 80 4 64 13 0.20 Fuchsia magellanica Lam.Nervous system disorders 67 3 59 12 0.12 Laureliopsis philippiana (Looser) Schodde, Melissa officinalis L.

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(H′ = 5.7) is also significantly higher than exotics (H′ = 5.2) (T = 24.9,p < 0.005), i.e., proportionally, native species are much more com-monly used than those of foreign origin. On the other hand,the evenness index indicates that the distribution of frequenciesof native species (J′ = 0.998) is similar to that of exotic species(J′ = 0.990), showing that authors’ documentation of native andexotic species is homogenous.

Results show the importance of the duration of the contactbetween Mapuche communities and plants, allowing more or lessexperimentation and experience of using them. Since ancestraltimes native plants have formed part of the Mapuche ecological andcultural landscape, resulting in profound, diverse wisdom. Personalcontact with nature provides opportunities for learning and exper-imentation, and this knowledge is later recreated and recorded bymeans of the social memory and forms part of the group’s collectivebody of knowledge.

The high proportion of exotic species in the Mapuche medicinalflora (39%) should be highlighted. Similarly high values for foreignplants have also been observed in other South American communi-ties (Bennett and Prance, 2000; Begossi et al., 2002, etc.). This couldbe due to the global character of the use of certain medicinal speciesgrowing wild in Patagonia, cultivated in gardens and home gar-dens, or being sold in some way, which enjoy great prestige and areof proven worth, e.g. Mentha spp. (Lamiaceae), Plantago lanceolataL. (Plantaginaceae), Artemisia absinthium L. (Asteraceae), Erodiumcicutarium (L.) LıHér. ex Aiton (Geraniaceae), etc. In the Patagonianregion, most of these species have been around for some 300 years,having arrived with the European colonizers (Ezcurra and Brion,2005), and this is long enough ago for them to have been adoptedand valued, given local names and gathered and/or cultivated ashealing elements of notable significance.

Also noteworthy are the high Shannon–Wiener diversity val-ues found for the Mapuche medicinal flora, which are similar toother ethnobotanical studies carried out in very diverse locationslike Mexico, Peru and Brazil (Begossi, 1996; Begossi et al., 2002).This reaffirms the importance of the Mapuche medicinal flora as areservoir of healing resources of world importance.

3.2. Similarity of medicinal species used, on a regional scale(Fig. 2)

On a regional scale, it can be seen that the medicinal plants of theMapuche form a solid nucleus. The MDS grouping analysis (Fig. 2,R2 > 0.99; stress = 0.061) shows a common group of species sharedby 12 of the 15 studies from different regions and times of study(group I: Martinez-Crovetto, 1980; Villagrán et al., 1983; Montecinoand Conejeros, 1985; Meza and Villagrán, 1991; Citarella, 1995;Conticello et al., 1997; Funes, 1999; Kutschker et al., 2002; Estombaet al., 2005, 2006; Ladio, 2006a; Ladio et al., 2007) and three groupswhich are separated from each other and from the main group, eachmade up of one sole species list (each of one single author) (groupII: Houghton and Manby, 1985; group III: Mösbach, 1992; group IV:San Martín, 1983).

Of the main group, the most alike are the studies of theseauthors: Estomba et al. (2005) and Estomba et al. (2006) (35% JI),Funes (1999) and Kutschker et al. (2002) (32% JI), and Martinez-Crovetto (1980) and Funes (1999) (31% JI). The work by San Martín(1983), Houghton and Manby (1985) and Mösbach (1992), are sep-arate from the rest. Of these, San Martín (1983) is the most similarto the main group, with a value of 23% for the Jaccard Index withrespect to the work of Citarella (1995) (23% IJ). The two studies,which are most different to each other, are Conticello et al. (1997)and Montecino and Conejeros (1985) (6.8%).

The separation between the studies of San Martín (1983),Houghton and Manby (1985) and Mösbach (1992) could be a reflec-tion of environmental or methodological differences between the

Fig. 2. MDS of the studies analysed (R2 > 0.99; stress = 0.061), showing four groupsof species lists = group I (Martinez-Crovetto, 1980; Villagrán et al., 1983; Montecinoand Conejeros, 1985; Meza and Villagrán, 1991; Citarella, 1995; Conticello et al.,1997; Funes, 1999; Kutschker et al., 2002; Estomba et al., 2005, 2006; Ladio, 2006a;Ladio et al., 2007); group II (Houghton and Manby, 1985); group III (Mösbach, 1992);group IV (San Martín, 1983).

communities under study. With respect to the environmental fac-tor, Ghirardini et al. (2007), on comparing lists of edible species indifferent populations of Italy, found that the species, which wereexclusive to the environments they were found in, were respon-sible for the separation of populations in a multivariable analysis.In our case, the study by Houghton and Manby (1985) was car-ried out in the scattered bushes of the transition zone, whilst inthe two other studies there is no exact description of the location,making accurate interpretation difficult. Nevertheless, differencesin the methodology employed, for example, in the selection of infor-mants, could be the most logical reason for such a separation. BothHoughton and Manby (1985) and Mösbach (1992) interviewed keyinformants who had specialized knowledge which included speciesused by expert herbalists in the treatment of magic-religious ill-nesses, whilst the rest of the studies included informants withwider, less specialized knowledge, concentrating more on domesticmedicine.

Without doubt, the uncertainty of the above-mentioned intro-duces a bias into this study which cannot be controlled, as inevitablyhappens in quantitative analyses of bibliographical revisions ofany kind (Avilés-Merens and Morales-Morejón, 2008). However,despite this, the synthesis provided by the MDS allows us to seeclearly that there is a body of knowledge shared by most of thearticles.

3.3. Agreement and use values of medicinal species and families

In accordance with our findings above, and supporting ourthird hypothesis, the native species present a high level of agree-ment between studies and high use values. The species withgreatest consensus were Gunnera tinctorea Molina (Mirb.) (Gun-neraceae) (93.8%), Buddleja globosa Hope (Buddlejaceae) (87.5%)and Aristotelia chilensis (Molina) Stuntz (Elaeocarpaceae) (87.5%)(Table 1). These species are abundant in woodlands and in thePatagonian wood-steppe ecotone, and are distributed through-out the entire Sub-Antarctic phytogeographical province, on bothsides of the Andes mountain range. General information on themain chemical components and medicinal properties of thesespecies can be found, for example, in Munoz et al. (2001). Inparticular, the anti-inflammatory and anti-oxidant uses of Bud-dleja globosa have recently been studied by Backhouse et al.(2008). On the medicinal qualities of Gunnera tinctorea, infor-mation has been offered by Bittner et al. (1994), and on thenutritional value, Petzold et al. (2006). As to the chemical

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composition and pharmacological action of Aristotelia chilensis,Miranda-Rottmann et al. (2002) and Stuntz et al. (1996) may beconsulted.

Next in the similarities ranking come the exotic species Men-tha × piperita L. (75%), Plantago lanceolata (75%) and Stellariamedia (L.) Cirillo (Caryophyllaceae), all of which are cosmopoli-tan in distribution and widely used in the medicinal floras ofthe world (Font Quer, 1962; Marzocca, 1997; Karousou et al.,2007).

Our results show that 60% of the species and 80% of the fami-lies registered are used for more than two different ailments (usevalue higher than two). The species with the highest use valueswere the natives Schinus latifolius (Gill. ex Lindl.) Engler (Anac-ardiaceae) (UVi = 5), Luma chequen (Molina) A. Gray. (Myrtaceae)(UVi = 5), Acantholippia seriphioides (A. Gray) Moldenke (Verbe-naceae) (UVi = 4), Grindelia chiloensis (Cornel.) Cabrera (Asteraceae)(UVi = 4) and Plantago australis Lam. (Plantaginaceae) (UVi = 4). Asindicated above, these data again reflect the importance of timespent in contact with native species, generating more intenseexperimentation with medicinal properties, leading to greater mul-tiplicity of application of resources. In addition, the multiple usesof plant species (in contrast to having a ‘specific’ use) in this tradi-tional medical system could reflect a certain capacity for resiliencein these local communities, since they adapt their knowledge anduse of certain species to different situations and contexts of illness(Ladio and Lozada, 2009). Patterns detected at a regional level inthis study reveal the importance of complementing this work withexhaustive pharmacological studies, particularly of those specieswith a wide spectrum of uses, given how little we know of theirmechanisms of action, levels of toxicity, among others.

In accordance with our fourth hypothesis, the botanical fam-ilies with greatest consensus between studies coincide withthose, which are most significant at a global level, althoughthere are local particularities. Of a total of 127 families, themost frequent were: Asteraceae (14%), Rosaceae (5.4%), Lami-aceae (6%), Solanaceae (4.7%), Apiaceae (4.2%), Polygonaceae(2.5%) and Fabaceae (2.4%). Asteraceae is the most frequentlyappearing family, as in the Holarctic and Neotropical medici-nal floras (Moerman et al., 1999; Leonti et al., 2003). Lamiaceaeand Fabaceae, on the one hand, follow the Neotropical pattern(Leonti et al., 2003) more closely than the Holarctic one. Api-aceae, in contrast, is important in the Holarctic medicinal florasbut not in the Neotropical ones. It is curious, on the other hand,that Polygonaceae, with a worldwide distribution of genera andspecies, presents high frequencies in this study, but is not foundamongst the most frequently mentioned medicinal families in otherregions.

These similarities and differences seem to indicate complexpatterns of use, where historical (Moerman et al., 1999), ecolog-ical (Ladio et al., 2007), phytochemical and organoleptic (Leontiet al., 2003) factors interact in the selection of medicinal species.Since the presence of active secondary metabolites is a cru-cial point, it is not surprising that Asteraceae, which has thecharacteristic of including a high number of species with notewor-thy pharmacological and organoleptic properties (Gottlieb, 1982;Milliken and Albert, 1999), should be the dominant family. More-over, Moerman (1998) has suggested that Asteraceae dominatetraditional pharmacopoeias as medicinal resources because theirtendency to grow in disturbed habitats near residential areas makesit easier to obtain information about them. On the other hand,it is interesting to note that Asteraceae, Fabaceae, Apiaceae andSolanaceae (as well as Poaceae) are, in all the Patagonian regionof Argentina, the most important families in terms of species rich-ness (Correa, 1998), thus revealing a possible correlation betweenregional flora and the ethnobotanical selection of plant fami-lies.

3.4. Main health ailments and use of medicinal plants

The authors cite a total of 2007 plant uses. In 1% of these, infor-mation is omitted on the uses. As proposed in our fifth hypothesis,gastro-intestinal problems are the most frequently encountered(26%) and present the highest level of consensus between studies(Fic = 0.47, Table 1), whilst ailments connected with the nervous sys-tem are the least frequent (3%) with the lowest consensus (Fic = 0.12,Table 1, Fig. 3). This highest frequency for the gastro-intestinalcategory has been registered in numerous studies carried out indifferent human groups throughout the world (e.g., Milliken andAlbert, 1997; Ankli et al., 1999; Schlage et al., 2000; Begossi et al.,2002).

The high citing frequency of gastro-intestinal uses could be dueto the kind of food and general state of hygiene in the Mapuchepopulations of the region, plus the fact that these ailments are con-sidered ‘of minor importance’ and are not dealt with in the healthcentres. During the last century, the traditional diet has undergonechanges, resulting in an increase in the consumption of refined flourand meat, and a decrease in the consumption of wild and culti-vated vegetables, leading to an unbalanced diet, low in fibre content(Ferrari et al., 2004). On the other hand, many of these communitieshave no supply of clean drinking water, their wells being situatedclose to latrines, or shared with farm animals, thus increasing therisk of ingesting pathogens.

At the other end of the scale, it is notable that cultural syndromespresented a low frequency of citing by authors (3%), and a lowconsensus index (Table 1). This low consensus of culture-boundsyndromes has also been found in other ethnobotanical studies(Heinrich et al., 1998; Leonti et al., 2001), possibly indicating thatthese ailments, or the traditional way of categorising them are beinglost.

Nevertheless, Campos et al. (1997) recorded 39 medicinalspecies to the south of Chile for treating ailments of a culturalnature, of which the most important were Chenopodium spp.(Chenopodiaceae), Aristotelia chilensis and Luma apiculata (DC.)Burret (Myrtaceae), amongst others. The central role played bythese cultural ailments, like the “aire”, “enfriamiento”, “susto”, etc.,in popular medicine cannot pass without comment, not only in

Fig. 3. Percentage of use reports (nur) per category of medicinal uses: GI, gastro-intestinal disorders; GU, genito-urological disorders; PI, pain and inflammation; RS,respiratory diseases; DE, dermatological diseases; GO, gynaecological-obstetric dis-eases; CH, circulatory and heart disorders; NS, nervous system disorders; CS, culturalsyndromes; F, fever; OT, others.

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Fig. 4. Comparison between numbers of native and exotic species in relation totherapeutic uses. GI, gastro-intestinal disorders; GU, genito-urological disorders; PI,pain and inflammation; RS, respiratory diseases; DE, dermatological diseases; GO,gynaecological-obstetric diseases; CH, circulatory and heart disorders; NS, nervoussystem disorders; CS, cultural syndromes; F, fever; OT, others.

the Patagonian region, but also for many Latin American peoples(Foster, 1953). Perhaps their poor representation in our results maybe explained by the authors’ categorization, where traditional con-cepts have been replaced by those from western medicine. Onceagain, the information brought to us by different authors presentsus with a challenge in understanding the selection of medicinalplants on a regional scale.

3.5. The biogeographical origin of species and different ailments

In the Mapuche medicinal flora, different ailments seem to betreated with both native and exotic plants (Fig. 4). However, inaccordance with the patterns found above, native species weremore frequently used than exotic ones, showing the greater culturalimportance of the former. Only in the cases of nervous system dis-orders and respiratory diseases were the differences not significant(Binomial test, p > 0.05, Fig. 4). In these categories, the exotic speciesused may be good for the treatment of modern illnesses, for whichthere is no equivalent amongst the native flora, or perhaps theyare more efficient in the treatment of these ailments (Albuquerque,2006). In addition, as suggested by Estomba et al. (2006), they mayhave a more pleasant taste, as in the case of Eucalyptus spp. (Myr-taceae), Mentha spp. or Rosmarinus officinalis L. (Lamiaceae), etc. It isalso true that the strong and bitter tasting species in the indigenoustaxonomy are considered to have therapeutic properties (e.g. Brett,1998; Pieroni et al., 2002; Pieroni and Torry, 2007). Examples ofexotic species with these characteristics are: Artemisia absinthium,Marrubium vulgare L. (Lamiaceae), and Tanacetum vulgare L. (Aster-aceae).

In agreement with our results, Leonti et al. (2001) found that agreat number of species employed by the Popoluca of Mexico in thetreatment of respiratory diseases are of exotic origin, and that thisis because in Pre-Hispanic times these ailments were less frequentthan now, having increased and become more widespread with thearrival of the Spanish Colonisers.

3.6. Growth habits and biogeographical origin of medicinal plants

As proposed in our sixth hypothesis, medicinal species aremainly herbs (56%), followed by shrubs (29%) and trees (13.5%), andto a lesser extent, climbing plants (1.4%) and parasites and hemi-parasites (0.5%) (Fig. 5). This would seem to be related in someway to the relative abundance of the different growth habits insome areas of Patagonian region. For example, Ezcurra and Brion(2005) showed that herbs (82%) and shrubs (11%) are the maincomponents of Nahuel Huapi National Park flora. Ferreyra et al.(2005) also demonstrated the predominance of herbs, followed byshrubs, in the upper mountain regions. Flores (2006) found that the

Fig. 5. Native and exotic: proportion of species in relation to growth habits.

understory vegetation of lenga (Nothofagus pumilio (Poepp. & Endl.)Krasser, Nothofagaceae) forest in the central and southern areas ofPatagonia consisted principally of herbs.

Growth habit varied significantly according to biogeographicalorigin (�2 = 93.5, p < 0.005). The trees, shrubs, climbing plants andparasites used are mainly native species (Binomial test, p < 0.005),whilst the herbs can be native or exotic (Binomial test, p = 0.45).

In general terms, the inclusion of exotic herbs in traditionalmedicinal floras is common in many parts of the world (Bennettand Prance, 2000; Stepp and Moerman, 2001; Stepp, 2004;Albuquerque et al., 2005, 2008). These plants are widely availablein the environment, being both abundant and easily visible (Voeks,2004), and on the other hand, they tend to be rich in medicallyvaluable components. These substances are products of secondarymetabolism and normally function as part of the plant’s defencesystem (Stepp and Moerman, 2001; Voeks, 2004).

3.7. Biogeographical origin of species in relation to gatheringenvironments and growth habits

Our results show that the Mapuche medicinal flora dependsmainly on gathering practices in wild areas, and to a lesser extenton cultivation and purchase in markets (Chi-square test = 696.7,p < 0.05, Fig. 6). As indicated in our seventh hypothesis, the great-est richness is found in woodlands and ecotones (F–E) (40%), withtrees, shrubs, herbs, parasites–hemiparasites and climbing plants,

Fig. 6. Native and exotic: proportion of species in relation to gathering environmentsand growth habits. F–E, Sub-Antarctic forest and/or ecotone; S–M, Patagonian steppeand/or monte; A–E, antropic environment with exotic and native species; C–B, gar-dens with cultivated species, and purchased exotic species; WH, wet habitats; HA,High Andean environment.

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which are mainly native species (Binomial test, p < 0.005, Fig. 6).The greater use made of these ecosystems seems to reflect, on theone hand, their greater structural complexity and richness in com-parison with other environments, and their greater cultural value,since the original ancestral territory of the present-day Mapucheswere the temperate forests of Argentina and Chile (Mösbach, 1992).Anthropogenic environments (A–E) follow in terms of speciesrichness (28%) and are composed mainly of exotic herb species(Binomial test, p < 0.005). These plants seem to be more oppor-tunistic than natives in environments suffering some degree ofdisturbance (Damascos et al., 1999).

On the other hand, it can be seen that at a regional level fewmedicinal species come from cultivation or purchase (C–B), addingup to only 14% of total richness, and these are mainly trees, shrubsor exotic herbs. The steppe–Monte (S–M) includes herbs and nativeshrubs, contributing 11.5%. Finally, the wet (WH) and High Andean(HA) environments basically contribute native herbs, which in totaladd up to 7% of the total richness (Binomial test, p < 0.005).

In general, native species are obtained through gathering, whilstexotics are obtained through gathering, cultivation and purchase. Incontrast to the results found in this study, many authors have sug-gested that communities use anthropogenic environments morethan primary environments in order to obtain medicinal plants(Bennett and Prance, 2000; Frei et al., 2000; Stepp, 2004). How-ever, such findings (which also coincide in part with others fromthe Patagonian region) could be put down to a question of scale. Ata local level, the environments, which are most accessible and mostmodified by the inhabitants, are most used, but at a regional level,the most used are the most pristine. These ideas should be exploredin greater detail, taking into account cross-cultural comparisonson a broader scale. Nevertheless, as established by other authorsfor other cultural contexts (Begossi et al., 2002), the importanceof introduced plants in Mapuche medicine would help to preventthe overuse of native species, and so these practices could be ofconsiderable consequence in terms of conservation.

3.8. Patterns of variation with time in ethnobotanical studies inPatagonia

As put forward in our eighth hypothesis, ethnobotanical stud-ies reveal a decrease over time in the richness of Patagoniannative plants in the communities studied (Spearmannative = −0.537,p < 0.05, Fig. 7). Nevertheless, there seems to be no evidenceof a pattern of loss in terms of the total number of plants(Spearmantotal = −0.315, p > 0.05, Fig. 7), nor in the case of exoticspecies (Spearmanexotic = −0.094, p > 0.05, Fig. 7).

Fig. 7. Correlation between time and cited species per author (n = 15), divided into(�) total number of species, (�) exotic species and (©) native species.

With respect to the loss of use of native plants, the inaccessi-bility of ancestral environments for plant gathering, particularlywoodlands which are included in national parks or private prop-erty, could be playing a crucial role in this. The richness of exoticspecies, in contrast, is generally on the increase, due partly to thegrowing abundance of anthropogenic surroundings (Ezcurra andBrion, 2005), an increase in publicity proclaiming their virtues(e.g. Rapoport et al., 1997, 2001; Kutschker et al., 2002), and theirincreasing availability from herbalists (Pochettino et al., 2008).

These tendencies could be interpreted in terms of culturalerosion, a worldwide phenomenon amongst indigenous peoples(Quilan and Quilan, 2007), or rather, in terms of cultural adaptation(Pilgrim et al., 2007), based on innovation and the reconstructionof traditional knowledge, where local knowledge is hybridised in adynamic way with global, according to new social, economic, polit-ical and ecological phenomenon affecting rural populations (Brodt,2002; Sears et al., 2007). This could be true for Patagonia, wheremost Mapuche populations live in contact with urban centres,which could have fomented interaction with the market society,thus increasing exchanges of knowledge, experience, technologyand customs. In this sense, the incorporation of exotic species willnot be considered in terms of erosion, but rather as a strategy todiversify the stock of medicinal species, thus increasing the rangeof alternative medicines and taking more care of the environment(Albuquerque, 2006; Albuquerque et al., 2008).

4. Conclusions

In general, in ethnobotany, information is collected from localcommunities on a small scale, such as individuals or families. Acomplex question is how to use this local information to make gen-eral analyses or propositions that go beyond the local communityand can be related to global issues such as plant knowledge erosionand native plant management. This study shows the importance ofregional analyses, providing a description of particular tendenciesor characteristics, which may differ from those on a local scale.

This study, at a regional level, demonstrates that the Mapuchemedicinal flora consists of a body of traditional and non-traditionalknowledge of great richness in both native and exotic plant species(over 500 species). As mentioned by other authors, traditional plantmedicines are made up of families and species which have not beenchosen by chance, but by complex processes of reasoning whichhave integrated the empirical with the symbolic, founded on his-torical, cognitive, botanical and chemical concepts (Messer, 1991;Gottlieb et al., 1995). Therefore, all these species would seem to bethe result of a complete selection process, maintained and recreatedby cultural transmission. Interestingly, our results coincide in manyaspects with those observed in other parts of the world, where gen-era and families are used to treat similar health ailments, and thiscould be understood as the result of common patterns of botani-cal knowledge at a world level (Moerman et al., 1999; Leonti et al.,2003).

On the other hand, this regional scale research is based on stud-ies were carried out over a long period of time. This has allowedus to prove, indirectly, that the body of medical knowledge onthe Mapuche is not static, but has been subject to processes ofchange, involving the loss of traditional knowledge (e.g. Ladio,2001; Estomba et al., 2006), but also the assimilation of innovationswhich have generated a kind of hybrid knowledge (sensu Sears etal., 2007). In other words, as proposed by Pilgrim et al. (2007), ithas in somehow responded in a flexible way to both environmen-tal and socio-cultural changes; however, this has not prevented theloss of cultural diversity in the process. Consequently, we believethat only by means of greater revaluation of this knowledge withinlocal communities can the processes of bio-cultural deterioration

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be stopped. It is therefore fundamental to allow access to, and pro-mote conservation of wild environments, particularly woodlandand the wood-steppe ecotone, in order to maintain the Mapuchemedicinal flora, which we have seen depends mainly on plant gath-ering there.

Acknowledgements

This research has been carried out thanks to the support ofthe Consejo Nacional de Investigaciones Científicas y Técnicas de laArgentina, along with grants provided by the Universidad Nacionaldel Comahue (B125), and the FONCYT (Pict 13593).

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