Agroforest Syst (2010) 80:131–152 DOI 10.1007/s10457-010-9315-x
Plant use and management in homegardens and swiddens: evidence from the Bolivian Amazon Evert Thomas • Patrick Van Damme
Received: 28 August 2009 / Accepted: 12 May 2010 / Published online: 25 May 2010 Springer Science+Business Media B.V. 2010
Abstract Amazonian plant management is perhaps nowhere as intense as in homegardens and swiddens. A quantitative ethnobotanical study was conducted in Indigenous Territory and National Park IsiboroSe´cure, Bolivia, to investigate plant use and management in homegardens and swiddens by local Yuracare´ and Trinitario ethnic groups. Ethnobotanical data of plants were obtained from 11 Yuracare´ and 11 Trinitario participants through semistructured interviews. A total of 151 different cultivated or tolerated species was recorded, accounting for 21% of all inventoried plants considered useful to local Yuracare´s and Trinitarios. The local use value of managed plants is almost twice that of wild plants. Managed plants score particularly higher than wild plants for medicinal, food and material applications. Most managed plants are herbs, followed by trees and shrubs. Nevertheless, managed trees have significantly higher overall use values than all other life forms. Managed trees tend to be particularly more appreciated as sources of food and materials, whereas herbaceous plants generally have a higher therapeutic value. Our results support observations made in literature that moderately humanized landscapes, and homegardens and swiddens in particular, are an
E. Thomas (&) P. Van Damme Laboratory of Tropical and Subtropical Agriculture and Ethnobotany, Ghent University, Coupure links 653, 9000 Ghent, Belgium e-mail:
[email protected]
important source of food and healing for forest people. Although people generally start managing plants in homegardens and swiddens because of their perceived usefulness, they are also favourable locations to experiment with the usefulness of (managed or wild) plants prevailing there. This particularly accounts for medicinal plants and it is argued that the use of managed plants in traditional medicine relates to (1) the high intensity of contact with theses species, and (2) their chemical defence strategy. To conclude, a number of policy recommendations are presented. Keywords Yuracare´ Trinitario Mojen˜o Resource availability theory Use value
Introduction All human societies owe their present day existence to a variety of ancestral accomplishments. Some of these are undoubtedly related to the long-standing management of botanical diversity. In the same way that intellectual diversity continuously stimulated cultural evolution (Kimmerer 2002), the raw material of botanical evolution has been manipulated to a varying extent throughout history. Some signs of plant management date back more than 30,000 years (Etkin 1998). Since then it has been a process developing in different societies around the globe as
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an adaptation strategy in an ever-changing world. In the Americas, at least 257 species were cultivated (138 species in lowland northern South America) when Columbus arrived in 1492 (Clement 1999a, b; Leon 1992). It has been argued that Amazonia has been overlooked as a cradle of crop diversity (Clement 1999b). More than one hundred plant species have been domesticated in the Neotropics, of which at least 52 in Amazonia. This is more than in any other region of the world (Bale´e 1994; Bru¨cher 1989; Clement 1999a). In addition, it is now clear that many crops were lost after contact (1492) as a consequence of post-contact Amazonian population decline. Therefore, it seems safe to assume that the Amazonian crop genetic heritage at contact was at least an order of magnitude greater than it is today (Clement 1999a). Plant management in Amazonia covers a continuum, ranging from gathering and protecting plants in wild populations, over deliberately tolerating plants in man-made habitats (also defined as disturbance habitats), to cultivating domesticates as well as non-domesticates. Amazonian plant management is perhaps nowhere as intense as in homegardens and swiddens (i.e. slash and burn cultivation areas). Homegardens have been one of man’s survival strategies since the Neolithic Period (8,500 BC). They have played a prominent role in the management and domestication of wild plants (Agelet et al. 2000; Soleri and Cleveland 1991). In homegardens, plant management is usually manifested as a combination of cultivated, domesticated and tolerated plants, whereas plant protection (basically referring to removing competing plants or pests to enhance the target plants’ chances of survival) mostly occurs in ‘‘natural’’ habitats (Miller and Nair 2006). However, it is often hard to define the exact management status of plants (Bennett 1992; Rival 2006), especially because some species are not restricted to a single category of resource management. Some species are cultivated in gardens at the same time as being protected in natural vegetation. Others occur as wild species in a natural habitat and can have been transplanted to a garden where they are considered cultivated (Bennett 1992; Van den Eynden 2004). Although most of these management techniques have been reported for many societies (e.g. Bennett 1992; Bonet and Valles 2002; Casas and Caballero 1995; Van den Eynden 2004), plant management can be quite variable, even within
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small communities. According to Van den Eynden (2004): ‘‘Management decisions are very individual and dynamic in time, as are use decisions. A plant managed by one individual is not necessarily managed by anyone else. Similarly, one particular plant species may be managed in different ways by different people, and its management may change in time.’’ In view of this high variability between those who manage plants and how they manage them, we will first describe how different management statuses have been interpreted in the present study. We distinguish between cultivated, tolerated and wild plants. Cultivated plants Cultivated or planted species are managed and cared for by people during their entire life cycle (Van den Eynden 2004). They are sown as seed, multiplied vegetatively by means of stem, root or other cuttings, or transplanted from natural habitats. Apart from interfering in their abundance and distribution, people also enhance their growth through irrigation, weeding, pruning, fertilization, insect control or protection against herbivores (Bennett 1992). A substantial number of cultivated plants depend (entirely) on human interference for propagation and survival. These plant species have been identified as domesticates since ‘‘the reproductive system of their populations has been so altered by sustained human intervention that the domesticated forms––genetically and/or phenotypically selected—have become dependent upon human assistance for their survival’’ (Harris 1989, p. 19). There also exist cultivated plant species that propagate naturally and hence are represented by wild populations. These species clearly do not (yet) satisfy the requirements to unmistakably be called domesticates. The average phenotypic variation of the part of the population modified by human selection may diverge from the range of variation found in the wild population, but the plants retain sufficient ecological adaptability to survive in the wild if human intervention ceases (Clement 1999a). They are often designated as semi-domesticates (Bale´e 1994; Clement 1999a). Hence, domesticated plants are largely, but not exclusively, a subgroup of cultivated plants: although most domesticated plants are cultivated, not all cultivated plants are necessarily
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domesticated (Van den Eynden 2004). According to Hawkes (1983), only about 200 of the 3000 crops used by humans worldwide have ever been domesticated1 (Clement 1999a). Tolerated plants Tolerated plant species are defined here as those species that are deliberately spared during weeding and land clearing activities for the benefits or usefulness they provide to humans. Tolerated species are not restricted to man-made landscapes. Since they do not depend on humans for reproduction, they can also prevail in natural disturbance (e.g. natural tree fall gaps, landslides…) or other habitats. Clement (1999a) called these plants ‘incipiently domesticated’ because their populations have been modified by human selection and intervention, but their average phenotype is still within the range of variation found in the wild population for the trait(s) subject to selection. Wild plants As opposed to ‘‘managed’’ plants, ‘‘wild’’ plants are defined as those plants that are not manipulated genotypically or phenotypically by humans in any way (Clement 1999a; Dufour and Wilson 1994). They can prevail both in anthropogenic and natural environments. Wild plants are defined by other authors (e.g. Van den Eynden 2004) as unmanaged plants that grow naturally outside habitats disturbed by humans, whereas weeds are unmanaged plants that grow spontaneously in man-made habitats. In fact, many weeds represent a status of ‘‘incidental coevolution’’. They have adapted to human disturbance environments, possibly undergoing genetic change, but without intentional human selection (Clement 1999a). The overall goal of this paper is to compare the relative utility of plants from anthropogenic and natural landscapes, and of plants with different growth forms, for indigenous people from the Amazon. To this end, and in light of the different management statuses described above, we describe 1
A more recent paper by Khoshbakht and Hammer (2008) puts the number of cultivated crop species worldwide at 7,000.
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the life form distribution, floristic composition and uses of plants managed in homegardens and swiddens by Yuracare´ and Trinitario people from the Bolivian Amazon. We investigate whether, and to what extent, plant management is growth-form dependent for different plant uses such as food and medicines. More specifically, we identify and discuss from a literature perspective, some of the factors that contribute to the human selection process of managed medicinal plant species. To conclude, we give an indication of the policy relevance of our findings.
Methods Research area Research was conducted in 32 different homegardens and swiddens from five indigenous communities that are located in Territorio Indı´gena Parque Nacional Isiboro-Se´cure (Indigenous Territory and National Park Isiboro––Se´cure, TIPNIS; Fig. 1). TIPNIS has been declared a protected area since 1965. Its 12,000 km2 surface area (between 16230 –16400 S and 65410 –65570 W) is cut in half by the still inaccurately defined frontier that separates the Bolivian departments of Beni and Cochabamba. TIPNIS is inhabited by Yuracare´s, Mojen˜os, Tsimane’ and Andean settlers, in relatively low population densities.
Fig. 1 Location of the participating indigenous communities (El Carmen, Tres de Mayo, San Antonio, San Jose and Sanandita) in TIPNIS (grey area), Chapare Province (hatched area), Bolivia (map elaborated with DIVA-GIS; www.divagis.org)
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The Andean settlers, who are principally concentrated along unpaved roads where vegetation is mainly secondary or relict, represent about 13,000 individuals, in a proportion of 14:1 Quechua:Aymara (Rico Pareja et al. 2005). Most Mojen˜os, Yuracare´ and Tsimane’ people live more dispersed and deeper in the forest, on high river banks and in the direct vicinity of old growth forest (Thomas 2009b). During the Bolivian census of 1992 and 1993, less than 5,000 indigenous inhabitants were counted in TIPNIS (68% Mojen˜o, 26% Yuracare´ and 4% Tsimane’). Nearly all of these people (93%) lived in small communities, whereas 7% lived in isolated houses. Only four communities contained more than 200 persons and the largest had only 265 (Andina and Fabricano 1994). Over the years this situation seems to have remained more or less unchanged. Rico Pareja et al. (2005) reported that at the time of this research Mojen˜os and Yuracare´s were the main indigenous groups in TIPNIS, being distributed over 53 settlements, and representing some 900 families and 5,000 inhabitants. The present study was carried out in participation with Yuracare´ and Mojen˜o ethnic groups only. Four of the participating communities are situated near the geographical centre of TIPNIS, in the upstream area of Rio Ichoa and Rio Moleto. They are the Trinitario (a subgroup of the Mojen˜os) communities of San Jose de la Angosta and El Carmen de la Nueva Esperanza, the Yuracare´ community of San Antonio de Moleto and the mixed community of Tres de Mayo (Fig. 1). The remaining Yuracare´ village of Sanadita is located near the south-eastern margin of TIPNIS, on the banks of Rio Isiboro. These communities are basically sedentary, but displacement of houses or even entire settlements within a given area or to nearby locations is not uncommon. The research area is located at altitudes below 300 m.a.s.l., on the transition between the ultimate Andean foothills and the eastern Bolivian lowland plains. It is characterized by a warm subtropical climate with mean annual temperature and precipitation of about 27C and 4000 mm, respectively (Rico Pareja et al. 2005). Vegetation largely consists of undisturbed old-growth tropical forest, interspersed with small patches (generally \1 ha) of secondary forest in different stages of succession that are the result of small-scale swidden agriculture. More specifically, natural (climax) vegetation is composed of subandean evergreen terra firme forests, white-water
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floodplain forests, piedmont forest and preandean riverine successional shrub- and woodlands (Navarro 2002; Thomas 2009a, b). Human disturbance of vegetation is rather limited, although new swiddens are generally cleared in mature vegetation and only secondary forest or fallows in the direct vicinities of settlements are cleared somewhat more sporadically (rotation cycles of ±10–20 years were reported). The Yuracare´ speak an unclassified language. In origin, they were hunter-gatherers with limited practice of agriculture. TIPNIS is one of their original habitats (Querejazu 2005). The Trinitarios (a subgroup of the Mojen˜os) belong to the Arawak language family (Rico Pareja et al. 2005). In the Bolivian municipality of Moxos (Beni department), the Mojen˜os reached a high level of development in pre-Hispanic times. From the nineteenth century onwards, many Trinitarios started to migrate from Moxos in search of the Holy Land (Loma Santa) as a response to land pressure problems in their homeland area and ended up in TIPNIS. The participating Trinitario communities were established from the 1970s onwards. Local Trinitario people still actively engage in shamanistic curing rituals (Thomas 2009b; Thomas et al. 2009d). Local Yuracare´ and Trinitario villagers hold extensive knowledge of the forests that surround them for fulfilling subsistence needs (Thomas et al. 2009c). Their main economic activity is slash-and-burn subsistence cultivation in swiddens of principally rice, banana, cassava and maize, supplemented with fishing and to a lesser extent hunting of large rodents, wild swine, deer, birds, monkeys, etc. People breed domestic animals like pigs, chickens and ducks for consumption. Coca (Erythroxylum coca Lam.) is grown as a cash crop to a limited extent. The participating communities were characterized by low accessibility. After a 3–5 h drive from the nearest town of Isinuta in a loaded cargo truck or shared taxi, the nearest community was reached on foot after a 0.5–4 h walk, the furthermost after a 5–8 h walk, depending on the location of the last truck stop and weather, etc. Ethnobotany The results presented here are part of a comprehensive quantitative ethnobotanical inventory that took place between March 2004 and February 2006. A total of 888 different taxa was inventoried during transect,
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walk-in-the-woods and homegarden and swidden sampling (see Thomas 2009b; Thomas et al. 2007, 2009c). The term homegarden is used here in line with Perrault-Archambault and Coomes (2008, p. 112) who define it as ‘‘the peridomestic area belonging to the household where members plant and/or tend useful plants’’. This includes the patio—the area in front of the house where mainly ornamental and occasionally also food and medicinal plants are grown; the huerto—the garden per se, most often found behind or beside the house; and the puerto—the household’s land near the river, where family members go to bathe and canoes are launched (Perrault-Archambault and Coomes 2008, p. 112). Useful plant species growing in homegardens and swiddens were collected upon indication, and after obtaining permission of their owners. All collected plants were identified by the first author with the help of several international taxonomic specialists (see acknowledgements) and deposited in the national Bolivian herbaria of Cochabamba (BOLV) and La Paz (LPB). Plant use data was obtained from 22 household members (7 women and 15 men, 11 Yuracare´s and 11 Trinitarios) through a variety of ethnobotanical interviewing techniques discussed in Thomas et al. (2007), i.e. (1) in situ interviewing during transect, walk-in-the-woods, and homegarden and swidden sampling, and (2) ex situ interviewing using fresh plant specimens, dried specimens, and photographs. Information on plant species from homegardens and swiddens was first and foremost obtained from their owners (several of the 22 participants owned more than one of the 32 sampled homegardens and swiddens), and to the extent possible also from other participants. Plant uses were grouped into use categories according to Cook’s (1995) Economic Botany Data Collection Standard. However, in analogy with other ethnobotanical studies in tropical forest environments (see Thomas et al. 2009c), construction materials were classified separately from materials. The latter are categorized in most other studies as ‘‘(handi)crafts’’. The eight resulting use categories are: medicines, including human and veterinary medicines; food, including beverages; construction, i.e. all materials used in house construction, timber and species used for manufacturing canoes; materials, including (handi)crafts, hunting gears, dyes, hygienic substances, instruments, toys, etc.; fuel;
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social uses, including ritual and magical uses, smoking materials/drugs and other unspecified social uses; environmental uses, i.e. exclusively ornamentals in the present study; and poison, including fish poison. Out of a total of 884 taxa shown to local Trinitario and Yuracare´ participants, 732 were considered useful (Thomas et al. 2009c). The usefulness of each plant species s was assessed according to the simplified formula of Phillips and Gentry (1993a): n P
UVs ¼
Uis
i
ns
whereby Uis equals the number of uses of species s mentioned by informant i and ns is the number of informants interviewed on species s. This approach has the advantage that, given a sufficient number of participants interviewed, minor uses or even erroneous answers will only minimally influence use values (Phillips and Gentry 1993a). Use values were calculated for each use category (i.e. medicine, food, material…). The overall use value of a plant species represents the sum of all categorical use values for that species. The management status of plants in homegardens and swiddens, was characterised as cultivated, tolerated, or wild following the descriptions in the introduction, based on interviews with garden tenders and personal observation. In case the management status of a plant species was confusing because management practices differed between household members and/or ethnic groups, it was assigned to the highest hierarchical management status. The following hierarchical order was applied: wild, tolerated and cultivated. Hence, the management status of a plant that was tolerated in swiddens by one Yuracare´ participant and considered wild by another was classified as ‘tolerated’. For the present analysis, no distinction was made between management statuses based on ethnic affiliation. The highest hierarchical management status of both classifications made by Yuracare´s and Trinitarios was consistently assigned to each plant species. For example, if a plant was labelled as wild for Yuracare´s and as cultivated for Trinitarios, then its overall management status was set as cultivated. A comparison of the plant knowledge, use and management between Yuracare´s and Trinitarios is described elsewhere (Thomas 2009b).
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SPSS 12.0 was used for performing statistical tests on data, including t-tests, Mann–Whitney tests and Kruskal–Wallis tests.
Results Yuracare and Trinitario plant management At least 151 plant species receive some form of management in Yuracare´ and/or Trinitario homegardens and swiddens (Table 1) where they are cultivated (105 species; 70%) or tolerated (46 species; 30%). The majority (75%) of cultivated plant species depend upon anthropogenic interference for survival and hence are labelled as domesticates. A number of cultivated, non-domesticated species (26 species) also occur naturally in the forest from where propagative (vegetative or seed) material is sometimes collected and (trans)planted in gardens. Fourteen percent of all cultivated useful plant species were encountered in just one homegarden each. Yuracare´s and Trinitarios weed their homegardens on an almost daily basis, basically to prevent invasion of pests such as insects, rodents and reptiles, hereby at the same time reducing snakebite incidents. However, this does not imply that homegardens are sterile environments that are completely free of weeds, colonizing or opportunistic (wild) plants. We counted 40 naturally occurring species that may deliberately be spared during yard cleaning, while others are removed. These spared species are therefore identified as being tolerated. This does not mean all Trinitario or Yuracare´ individuals spare these species at all times. Weeding does occur also among tolerated species, but generally at least part of the local plant population is left untouched. In addition, toleration is a highly personal experience: what is spared by some for its usefulness is simply considered a pest by others and removed. Some useful species of the original (primary or secondary) vegetation may be spared by Yuracare´s and/or Trinitarios during the establishment of settlements and/or while clearing forest for swidden cultivation. In all the swiddens we visited, the canopy consisted of at least one or more of the following multipurpose palm species: Socratea exorrhiza, Iriartea deltoidea, Euterpe precatoria, Attalea butyracea, and exceptionally the less abundant Bactris
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gasipaes or Jessenia bataua. Of these, only S. exorrhiza and I. deltoidea are considered tolerated species because the other species were said to be sowed or planted by at least one of our participants and therefore were assigned the hierarchically higher management status of ‘cultivated’. Numerous other useful (edible) species (e.g. Chrysophyllum venezuelanense, Pouteria spp., Pseudolmedia spp. or Inga spp.) may be spared during the establishment of new settlements or swiddens. However, the presence of these mostly rare (in terms of abundance in the entire landscape) species in homegardens or swiddens is far from consistent, and their occurrence seems to be highly dependent on the sites where settlements and/ or swiddens are established. In view of the haphazard occurrence of these species and the incompleteness of our data regarding which species are ‘typically’ spared and which ones are not, we classified them as wild species. Sometimes oversized trees are left behind on swiddens, simply because they are too labour-intensive to cut down. Although not respected by all, some conservation ethics come into play as well. A Trinitario elder explained that ‘‘one should not cut down mature trees of Hura crepitans, Ceiba pentandra, Dipterix micrantha or D. odorata since they are ‘‘the mothers of the forest’’ that keep the soil humid, hereby allowing other plants to grow. If these trees would be cut down systematically, ‘‘the forest would die’’. C. pentandra is a tree species with a particular spiritual connotation. Trinitarios believe that sorcerers learn their evil magic from this tree. Felling large individuals of this species is not done as it is believed to cause mischief. We have observed various swiddens that contained spared large-sized individuals of these ‘‘mother trees’’ that therefore are considered as tolerated species. Hence, we identified 46 different species as being tolerated by Yuracare´s and/or Trinitarios: 40 species that occur spontaneously in homegardens or swiddens and six species that are spared when clearing original forest vegetation. Although both are classified as ‘tolerated’, the main difference between plants tolerated in homegardens and swiddens, and plants that are spared during forest clearing is that the former are often typical disturbance species, whereas the latter are essential parts of the natural (climax) flora. As opposed to tolerated plants in already established homegardens and swiddens, plants spared during
t h h t t s v h t h s h
Zea mays L. (y/t)* (Poaceae) Sida rhombifolia L. (y/t) (Malvaceae)
Astrocaryum murumuru Mart. (y)* (Arecaceae)
Ochroma pyramidale (Cav. ex Lam.) Urb. (y/t) (Bombacaceae)
Erythroxylum coca Lam. (y/t)* (Erythroxylaceae)
Momordica charantia L. (y/t) (Cucurbitaceae)
Nicotiana tabacum L. (y/t)* (Solanaceae)
Euterpe precatoria Mart. (t)* (Arecaceae)
Zingiber officinale Roscoe (y/t) (Zingiberaceae)
Bixa orellana L. (y/t)* (Bixaceae)
Gynerium sagittatum (Aubl.) P.Beauv. (y/t) (Poaceae)
Mangifera indica L. (y/t) (Anacardiaceae)
Carica papaya L. (y/t)* (Caricaceae)
h t
Oryza sativa L. (y/t) (Poaceae)
t s
Lantana fiebrigii Hayek (y/t) (Verbenaceae)
Citrus reticulata Blanco (y/t) (Rutaceae)
Citrus sinensis (L.) Osbeck (y/t) (Rutaceae)
t t
Citrus maxima (Burm.) Merr. (y/t) (Rutaceae)
v
Cissus gongylodes (Baker) Planch. (t)* (Vitaceae)
t
Hura crepitans L. (t) (Euphorbiaceae)
t
t t
Genipa americana L. (y/t)* (Rubiaceae) Socratea exorrhiza (Mart.) H.Wendl. (Arecaceae)
t
t
Swietenia macrophylla King (y/t) (Meliaceae)
Theobroma cacao L. (y/t)* (Sterculiaceae)
s
Manihot esculenta Crantz (y/t)* (Euphorbiaceae)
Iriartea deltoidea Ruiz & Pav. (Arecaceae)
t
Jessenia bataua (Mart.) Burret (t)* (Arecaceae)
v
t
Attalea butyracea (Mutis ex L.f.) Wess.Boer (y/t) (Arecaceae)
t
t
Bactris gasipaes Kunth (y/t)* (Arecaceae)
Persea americana Mill. (y/t)* (Lauraceae)
t
Attalea phalerata Mart. ex Spreng. (y/t) (Arecaceae)
Ipomoea batatas (L.) Lam. (y/t)* (Convolvulaceae)
Habita
Species and family
x
x
x
x
x
x
x
Introdb
x
x
x
x
x
x
x
x
x x
x
x
x
x
x
Also wildc
c
c
c
c
c
c
c
t
c
c t
c
c
c
c
c
c
c
t
c
t
c
c
t
c t
c
c
c
c
c
c
Managd
Table 1 Plant species that were reported and observed to be cultivated or tolerated by Yuracare´ (y) and/or Trinitario (t) participants
Ma
Fo/Me/Ma
Fo/Me
Ma/Fo/Co/Me
Me/Soc
Me/Fo/Ma
Me/Fo
Ma/Me
Ma/Fo/Co
Fo Me/Ma
Fo/Me
Fo/Me
Fo/Me
Fo/Me
Fo/Me
Fo/Me
Fo/Me
Me
Fo/Me
Ma/Co/Fo/Me
Fo/Me
Fo/Me
Me/Ma/Co/Ti/P
Fo/Me/Ma/Fu Ma/Co/Fo/Me
Me/Co/Ti/Ma
Fo/Me
Ma/Fo/Co/Me
Ma/Fo/Co/Me
Ma/Fo/Co
Ma/Fo/Co/Me
Main usese
2.40
2.40
2.43
2.50
2.57
2.57
2.60
2.64
2.67
2.75 2.71
2.80
2.85
3.00
3.00
3.00
3.00
3.00
3.00
3.17
3.18
3.20
3.25
3.33
3.43 3.36
4.00
4.00
4.63
5.83
5.86
6.13
UVfs
9
5
7
4
7
7
5
11
12
4 7
5
8
2
1
1
1
2
3
6
11
5
8
12
8 14
4
3
8
6
8
8
Ngs
Agroforest Syst (2010) 80:131–152 137
123
123 t t t s h h
Heliocarpus americanus L. (y/t) (Tiliaceae)
Psidium guajava L. (y/t)* (Myrtaceae)
Rheedia gardneriana (Ruiz & Pav.) Planch. & Triana (y/t) (Clusiaceae)
Gossypium barbadense L. (y/t)* (Malvaceae)
Ocimum micranthum Willd. (y/t) (Lamiaceae)
Portulaca grandiflora Hook. (t) (Portulacaceae)
x
x
x
x
x
s
x
x
c
c
c
c
c
c
t
c
c
c
c
c c
t
t
x
c
t
c
Salmea scandens (L.) DC. (y/t) (Asteraceae)
x
x
Citrus cf. limetta Risso (y/t) (Rutaceae)
s
Pereskia sacharosa Griseb. (t) (Cactaceae)
x
s
h
Paspalum conjugatum P.J.Bergius (y/t) (Poaceae)
Capsicum chinense Jacq. (y/t)* (Solanaceae)
h
Kalanchoe pinnata (Lam.) Pers. (y/t) (Crassulaceae)
c
v
h
Eleutherine citriodora (Ravenna) Ravenna (t) (Iridaceae)
c
c
Cucurbita maxima Duchesne ex Lam. (y/t)* (Cucurbitaceae)
h
Arachis hypogaea L. (y/t)* (Fabaceae)
c
c
c
t
Jatropha curcas L. (y/t) (Euphorbiaceae)
x
s t
s
Datura suaveolens Willd. (y/t)* (Solanaceae)
c
h
h
Cymbopogon citratus (DC.) Stapf (y/t) (Poaceae)
x
Coix lacryma-jobi L. (y/t) (Poaceae)
h
Petiveria alliacea L. (y/t) (Phytolaccaceae)
c t
c
c
Ricinis communis L. (y/t) (Euphorbiaceae) Rheedia acuminata (Ruiz & Pav.) Planch. & Triana (y/t) (Clusiaceae)
s v
Crescentia cujete L. (y/t)* (Bignoniaceae) Dioscorea dodecaneura Vell. (t)* (Dioscoreaceae)
x
x
c
t
Citrus aurantifolia (Christm.) Swingle (y/t) (Rutaceae)
s
h
Saccharum officinarum L. (y/t) (Poaceae)
c
t
Lippia alba (Mill.) N.E.Br. (t) (Verbenaceae)
h
Chenopodium ambrosioides L. (y/t) (Chenopodiaceae)
x
t
t
h
t
Cecropia concolor Willd. (y/t) (Cecropiaceae)
x
x
Managd
t
t
Dipterix odorata (Aubl.) Willd. (t) (Fabaceae)
Also wildc
Cecropia polystachya Trecul (y/t) (Cecropiaceae)
t
Dipterix micrantha Harms (t) (Fabaceae)
Introdb
Sansevieria cf. trifasciata Prain (t) (Liliaceae)
Habita
Species and family
Table 1 continued
O/Me
Me
Ma/Me
Fo/Me/Ma
Fo/Me
Ma/Me
Me/O
Fo/Me
Fo/Me
Fo/Me
Me/Fu/O/Ma Fo/Ma
Fo/Me/Ma
Me
Fo/Fu/Me/Ma
Me/O
Me/O
Me
Me/O
Me
Fo/Me
Me/Fu/Ma
Me/Soc/O
Fo/Me
Me/O
Ma/Me Me/Fo
Fo/Me
Fo/Me
Me
Ma/Fu/Fo
Ti/Fo/Ma/Me/Co
Ti/Fo/Ma/Me/Co
Main usese
1.58
1.58
1.63
1.67
1.67
1.67
1.80
1.80
1.82
1.83
1.86 1.85
1.86
1.88
1.91
2.00
2.00
2.00
2.00
2.00
2.00
2.07
2.13
2.17
2.19
2.20 2.20
2.20
2.25
2.33
2.33
2.38
2.38
UVfs
12
12
8
9
12
9
15
5
11
6
14 13
7
8
11
1
7
8
8
3
6
15
15
6
16
5 5
5
4
6
6
8
8
Ngs
138 Agroforest Syst (2010) 80:131–152
s
Casearia pitumba Sleumer (y/t) (Flacourtiaceae)
h
Geophila repens (L.) I.M.Johnst. (y/t) (Rubiaceae)
l
s
Capsicum chacoense Hunz. (y/t) (Solanaceae)
Passiflora nigradenia Rusby (y/t) (Passifloraceae)
l
Passiflora nitida Kunth (y/t) (Passifloraceae)
h
h
Heliconia rostrata Ruiz & Pav. (y/t) (Heliconiaceae)
Hedychium coronarium J.Koenig (y/t) (Zingiberaceae)
h h
Colocasia esculenta (L.) Schott (y/t) (Araceae)
s s
Tephrosia vogelii Hook.f. (y/t) (Fabaceae) Phyllanthus brasiliensis (Aubl.) Poir. ssp. glaber (Pax & K.Hoffm.) G.L.Webster (y/t)* (Euphorbiaceae)
Scoparia dulcis L. (y/t) (Scrophulariaceae)
s
Urera baccifera (L.) Gaudich. (y/t) (Urticaceae)
s
Solanum mammosum L. (y/t) (Solanaceae)
h
t
Heliconia lingulata Ruiz & Pav. (y/t) (Heliconiaceae)
v
Lagenaria siceraria (Molina) Standl. (y/t)* (Cucurbitaceae)
Rollinia herzogii R.E.Fr. (y/t) (Annonaceae) t
s
t
h
Solanum morellifolium Bohs (y/t) (Solanaceae) Cereus cf. braunii Ca´rdenas (t) (Cactaceae)
Vernonia patens Kunth (y/t) (Asteraceae)
h
Alpinia zerumbet (Pers.) Burtt & R.M.Sm. (y/t) (Zingiberaceae)
Annona montana Macfad. (y)* (Annonaceae)
s h
Cyperus cf. corymbosus Rottb. (t) (Cyperaceae)
t
Anacardium occidentale L. (y/t)* (Anacardiaceae)
Coffea arabica L. (t) (Rubiaceae)
h t
Pothomorphe peltata (L.) Miq. (y/t) (Piperaceae) Pouteria nemorosa Baehni (y) (Sapotaceae)
t
Inga edulis Mart. (y/t)* (Fabaceae) h
s
Ananas comosus (L.) Merr. (y/t)* (Bromeliaceae)
h
p
Struthanthus acuminatus (Ruiz & Pav.) Blume (y/t) (Loranthaceae)
Musa sapientum L. (y/t) (Musaceae)
h
Geophila macropoda (Ruiz & Pav.) DC. (y/t) (Rubiaceae)
Mentha sp. (t) (Lamiaceae)
Habita
Species and family
Table 1 continued
x
x
x
x
x
x
x
Introdb
x
x
x
x
x
x
x
x
x
x
x
x
x x
x
x
Also wildc
c
c
c
t
c
c
t
c
t
c c
t
t
t
c
c
c
c
c
t
c
c
c
c
t c
c
c
c
c
t
t
Managd
Fo/Me/Ma
Fo/Me
Me/O
Me/Ma
Fo/Me
Fo/Me
O/Ma
Fo/Me
Me/Ma
P/Me/Ma P/Me
Me/Fo
Ma/Me
Fu/Me/Ma
Fo/Me/Ma
Me/O
Fo/Ma
Ma/Me
Me
Me
O/Me/Fo
Me
Fo/Me
Fo/Me/Ma
Me Fo
Fo/Me/Ma
Fo/Me
Fo/Fu
Fo
Me
Me/Ma
Main usese
1.14
1.17
1.17
1.17
1.18
1.20
1.20
1.20
1.21
1.23 1.21
1.25
1.25
1.29
1.30
1.33
1.33
1.33
1.33
1.36
1.38
1.40
1.40
1.40
1.50 1.50
1.50
1.50
1.50
1.50
1.57
1.57
UVfs
7
6
6
6
11
5
5
5
14
13 14
12
4
7
10
12
6
6
3
11
8
10
5
10
14 4
6
2
6
6
7
14
Ngs
Agroforest Syst (2010) 80:131–152 139
123
123 t h h s s f h h h
Cyathula prostrata (L.) Blume (t) (Amaranthaceae)
Pachyrhizus tuberosus (Lam.) Spreng. (y/t)* (Fabaceae)
Hibiscus rosa-sinesis L. (y/t) (Malvaceae)
Cajanus cajan (L.) Millsp. (y/t) (Fabaceae)
Polypodium decumanum Willd. (y/t) (Polypodiaceae)
Bidens pilosa L. (y/t) (Asteraceae)
Chamaesyce hirta (L.) Millsp. (t) (Euphorbiaceae)
Desmodium axillare (Sw.) DC. (t) (Fabaceae)
Syzygium malaccense (L.) Merr. L.M.Perry (y/t) (Myrtaceae) Tagetes patula L. (t) (Asteraceae) t
t h
Solenostemon scutellarioides (L.) Codd (y/t) (Lamiaceae)
Thevetia peruviana (Pers.) K.Schum. (t)* (Apocynaceae)
h
Solanum sp. (y/t) (Solanaceae)
Tamarindus indica L. (t) (Fabaceae)
t s
Rollinia mucosa (Jacq.) Baill. (y/t)* (Annonaceae)
t
h
Hymenocallis cf. tubiflora Salisb. (t) (Liliaceae)
t
s
Hibiscus acetosella Welwitsch ex Hiern (y/t) (Malvaceae)
Rollinia cf. boliviana R.E.Fr. (y/t) (Annonaceae)
h
Celosia argentea L. ‘Cristata’ (y/t) (Amaranthaceae)
Neea sp.1 (Nyctaginaceae)
h
Carludovica palmata Ruiz & Pav. (t) (Cyclanthaceae)
h
s s
Capsicum frutescens L. (y/t)* (Solanaceae) Capsicum pubescens Ruiz & Pav. (y/t) (Solanaceae)
h
s
Caesalpinia pulcherrima (L.) Sw. (y/t) (Fabaceae)
Musa velutina H.Wendl. & Drude (y/t) (Musaceae)
v
Dioscorea trifida L.f. (y/t)* (Dioscoreaceae)
Musa paradisiaca L. (y/t) (Musaceae)
s
Mirabilis jalapa L. (y/t) (Nyctaginaceae)
l
l
Salacia impressifolia (Miers) A.C.Sm. (t) (Hippocrataceae)
v
h
Heliconia stricta Huber (y/t) (Heliconiaceae)
Merremia macrocalyx (Ruiz & Pav.) O’ Donell (t) (Convolvulaceae)
h
Arachis pintoi Krapov. & W.C.Greg. (y/t) (Fabaceae)
Marsdenia macrophylla (Schult.) E.Fourn. (y) (Asclepiadaceae)
Habita
Species and family
Table 1 continued
x
x
x
x
x
x
x
x
x
x
Introdb
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Also wildc
t
t
t
c
c
c
c
t
c
c
c c
c
t
c
c
c
c
c
t
c
c
c
c
c
c c
c
c
c
c
t
c
Managd
Me
Me
Me
Me
Fo/Ma
Me/O
Fo
Me
Ma/Me
Fo
Fo Me/O
O/Me
Fo/Me
Fo
Fo/Me
Ma
Fo/Me/O
Fo/Me
Me
Ma
O
O
O/Fo/Ma
Ma
Fo/Me Fo
O
Fo
O/Me
Fo
Ma/Me/O
O/Me
Main usese
0.75
0.80
0.83
0.86
0.89
0.91
0.92
1.00
1.00
1.00
1.00 1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00 1.00
1.00
1.09
1.10
1.11
1.11
1.11
UVfs
4
5
6
7
9
11
12
2
4
3
4 2
6
4
4
12
1
11
2
1
3
2
6
11
1
4 6
6
11
10
9
9
9
Ngs
140 Agroforest Syst (2010) 80:131–152
t h h h h
Ceiba pentandra (L.) Gaertn. (y/t) (Bombacaceae)
Desmodium cf. adscendens (Sw.) DC. (t) (Fabaceae)
Aneilema umbrosum (Vahl) Kunth subsp. ovato-oblongum (P.Beauv.) J.K.Morton (t) (Commelinaceae)
Dichorisandra hexandra (Aubl.) Standl. (y/t) (Commelinaceae)
Oxalis juruensis Diels (t) (Oxalidaceae)
x
x
Introdb
x
x
x
x
x
x
x
x
x
x
x
x x
x
x
x
x
x
Also wildc
t
t
t
t
t
t
c
t
t
t
t
c
c
t t
t
t
c
c
t
t
Managd
Me
Me/O
Me
Me
Ma/Me
Me
Fo
Ma
Me
Fo/Me
Fo/Me
Me
Ma
Me Me/Ma
Fo/Me/O
Fo
Fo/Me
Fo/O
Me
Me
Main usese
0.11
0.22
0.22
0.33
0.33
0.38
0.43
0.43
0.44
0.44
0.50
0.50
0.60
0.63 0.60
0.64
0.67
0.71
0.75
0.75
0.75
UVfs
9
9
9
3
6
8
7
7
9
9
2
4
5
8 5
11
6
7
8
8
8
Ngs
Different habits of managed plants are: t tree, s shrub, h herb, v vine, l liana, f fern, p hemi-parasite
g
f
e
d
c
ns = number of participants interviewed per plant species
UVs = overall use value
Principal uses are given per species, whereby Ti timber, Fo food, Fu fuel, Ma material; Me medicine, P fish poison, O ornamental, Soc social uses
Management status, c cultivated, t tolerated
Managed plant species that also occur in wild vegetation are marked with ‘x’
Managed plants species that are introduced from the Old World are marked with ‘x’; the introduced origin of different species was obtained from the following references: Bennett and Prance 2000; Cardenas 1989; Clement 1999a and b; INIBAP 2001; Killeen et al. 1993; Mabberley 1990; Morton 1987; Mukherjee 1957; Raemaekers 2001; Steyermark 1963; Va´squez and Coimbra 2002; Zeven and Zhukovsky 1975
b
a
Although listed only once, two varieties of Gynerium sagittatum are cultivated: G. sagittatum var. subandinum is used for shafts of arrows used by children and G. sagittatum var. sagittatum used for shafts of arrows used by adults. Plants marked with ‘‘*’’ were (incipiently) domesticated or semi-domesticated in Amazonia at time of contact (1492) according to Clement (1999a, b; see introduction for meaning of terminology)
t h
Lycianthes asarifolia Bitter (y/t) (Solanaceae)
t
Morus alba L. var. indica (L.) Bureau (y/t) (Moraceae)
h
Bauhinia longicuspis Spruce ex Benth. (y) (Fabaceae)
Neea cf. spruceana Heimerl (y) (Nyctaginaceae)
Solanum mite Ruiz & Pav. (y/t) (Solanaceae)
s
Lindernia crustacea (L.) F.Muell. (y/t) (Scrophulariaceae) Laportea aestuans (L.) Chew (y) (Urticaceae)
h
h h
Wulffia baccata (L.f.) Kuntze (y/t) (Asteraceae)
Solanum americanum Mill. (y/t) (Solanaceae)
s
Casearia sylvestris Sw. (y) (Flacourtiaceae)
s
s
Porophyllum ruderale (Jacq.) Cass. (y/t) (Asteraceae)
s
h
Muntingia calabura L. (y/t) (Elaeocarpaceae)
Solanum exiguum Bohs (y/t) (Solanaceae)
t
Lantana camara L.(t) (Verbenaceae)
Euphorbia pulcherrima Willd. ex Klotzsch (y/t) (Euphorbiaceae)
h s
Eupatorium macrophyllum L. (y/t) (Asteraceae)
Habita
Species and family
Table 1 continued
Agroforest Syst (2010) 80:131–152 141
123
142
Agroforest Syst (2010) 80:131–152
Usefulness of managed and wild plants The mean overall use value of managed species (UV = 1.70 ± 1.08 SD) is nearly twice that of wild species (UV = 0.95 ± 0.80 SD) (Table 2), showing the superior usefulness of managed over wild species. Managed plant species have particularly higher use values for medicine, food, materials and environmental uses. On the contrary, wild species score significantly higher for construction and fuel (Table 2). Therefore, we will particularly focus on overall, medicine, food and material use values when performing analyses related to plant management. The differences in mean use values between managed and wild species are more pronounced when managed plants are separated into cultivated and tolerated plants (Fig. 2). The mean overall use value (all uses) (P \ 0.01, t-test) and food use value (P \ 0.01, Mann–Whitney) of cultivated plants are significantly higher than of tolerated species. Mean use values for medicine and material are equal for cultivated and
2,50
cultivated tolerated wild
2,00
Mean Use Value
forest clearing did not establish themselves naturally during a period of human occupation. They were already present when the land surface became subjected to clearing and subsequent (intensive) human management. A number of wild plant species that develop spontaneously in homegardens are not managed, in spite of their usefulness. Participants declared that there is no need to deliberately spare these plant species because ‘‘they are always there, they never disappear’’. When needed, they are easy to find in habitats under anthropogenic disturbance such as homegardens, trails, and/or young or recently abandoned swiddens.
1,50
1,00
0,50
0,00 all uses
medicine
food
material
Use Category
Fig. 2 Comparison of mean categorical use values for all inventoried cultivated, tolerated and wild plant species; error bars represent standard errors of the mean; significantly equal mean use values are indicated by horizontal lines
tolerated species (P [ 0.01, t-test and Mann–Whitney). Hence, cultivated plants have an important food value in addition to their medicinal importance, whereas the value of tolerated plants is mainly medicinal. In addition to managed plants, a number of species typically grow wild in environments under a regime of anthropogenic disturbance. Many of these wild species, such as ruderal plants, agricultural weeds, or disturbance trees and shrubs (e.g. Piper spp., Urera spp. and Cecropia spp.) can also appear in natural environments (e.g. tree fall gaps or landslides), but their abundance/and or distribution is positively influenced by human activities. Since people spend most of their time in anthropogenic disturbance habitats, it is logical to expect that they come into contact more with the plants that grow there. The question that follows is whether these species are also
Table 2 Mean use values and standard deviations of wild and managed plants for different use categories UVs Wild (N = 733)
UVfo
UVma
UVc
0.95 ± 0.80 0.16 ± 0.33 0.18 ± 0.39 0.12 ± 0.35 0.22 – 0.47
Managed (N = 151) 1.70 – 1.08 Significance
UVme
***a
UVfu
UVe
UVso
UVp
0.24 – 0.43 0.01 ± 0.04 0.01 ± 0.08 0.003 ± 0.05
0.68 – 0.67
0.51 – 0.65
0.24 – 0.54
0.12 ± 0.42 0.04 ?0.12 0.07 – 0.23
0.04 ?0.13
0.02 ± 0.13
**b
**b
**b
**b
NSb
NSb
**b
**b
Highest values are marked in bold UVs overall use value, UVme medicinal component, UVfo food component, UVma materials component, UVc construction component, UVfu fuel component, UVe environmental uses (i.e. ornamental use) component, UVso social uses component, UVp poison component; NS not significant a
t-test
b
Mann–Whitney test
*** P\0.001; ** P\0.01
123
Agroforest Syst (2010) 80:131–152
143
Taxonomy, life form, origin and usefulness of managed plants The 151 managed plant species encountered in Yuracare´ and Trinitario homegardens belong to sixty botanical families. The best-represented families are: Solanaceae (14 species), Fabaceae (13), Arecaceae (8), Asteraceae (7), Poaceae (7), Rutaceae (5) and Euphorbiaceae (5). The majority of plants managed in homegardens are herbs (64 species, including 8 vines; 42%), followed by trees (46 species; 30%) and shrubs (35 species; 23%). Four lianas and one
medicinal fern species are cultivated and one hemiparasite (Struthanthus acuminatus) frequently growing on Citrus spp. is tolerated for its potent medicinal properties against sprains and fractures (Thomas and Vandebroek 2006). Thirty-three cultivated and two tolerated plant species (representing 31% of all cultivated and 23% of all managed species, respectively) are introduced from the Old World (Eurasia and Africa) (Table 1). In proportion, introduced plants make up around one-third of cultivated food or medicinal plants. As indicated in Table 1, managed plants serve many uses in TIPNIS. Taking cultivated and tolerated species together, the majority are used as herbal medicine (81%), followed by food (52%), material (34%), environmental uses (17%), fuel (15%), social uses (13%) and construction (13%) (Fig. 3). Three 140 120
Number of Species
considered more useful than wild species that grow in natural landscapes. To verify this, we made a comparison between the usefulness of wild plant species that typically grow in natural (N = 687 spp.) and anthropogenic (N = 55 spp.) landscapes, respectively. In the scope of this analysis we considered as anthropogenic landscapes: homegardens and swiddens (20 wild species); fallows and secondary forest (24); and ruderal environments such as roadsides or trails (11). Natural landscapes include old growth forest vegetation, as well as riverine vegetation. Analysis showed that the calculated mean overall use value of wild plants from natural environments is higher than that of wild plants from anthropogenic environments (Table 3), but this difference is not significant. However, wild plants from anthropogenic environments have a significantly higher mean medicinal use value. For food, construction and fuel, wild plants from anthropogenic environments have lower mean use values as compared with wild plants from natural environments (Table 3).
100
tolerated cultivated
42
80 13
60 40
20
80 66
4
20
7
32 21
15
0 medicine
food
material env. use
fuel
8
9
11
10
soc. use constr
2
poison
Use Category
Fig. 3 Numbers of managed species used per use category (plants can have multiple uses); managed plants are represented as the sum of cultivated and tolerated plants (env. use = environmental use; soc. use = social use)
Table 3 Mean use values and standard deviations of wild plants typically growing in anthropogenic and natural landscapes for different use categories UVs
UVme
UVfo
UVma
UVc
UVfu
UVe
UVso
UVp
Anthropogenic landscapes (N = 55)
0.81 ± 0.67 0.34 – 0.38 0.04 ± 0.20 0.16 ± 0.37 0.08 ± 0.40 0.02 ± 0.14 0.02 ± 0.07 0.02 ± 0.14 0 ± 0
Natural landscapes (N = 687)
0.96 ± 0.85 0.15 ± 0.33 0.19 – 0.40 0.12 ± 0.35 0.23 – 0.47 0.26 – 0.44 0.01 ± 0.04 0.01 ± 0.08 0.003 – 0.05
Significance
NSa
**a
**b
NSb
**b
**b
NSb
NSb
**b
Highest values are marked in bold UVs overall use value, UVme medicinal component, UVfo food component, UVma materials component, UVc construction component, UVfu fuel component, UVe environmental uses (i.e. ornamental use) component, UVso social uses component, UVp poison component, NS not significant a b
t-test Mann–Whitney test
** P \ 0.01
123
144
Agroforest Syst (2010) 80:131–152
species are used as fish poison. Similar proportions of plants per use category were obtained for cultivated species on their own, whereas most tolerated plant species are used in traditional medicine (91%) and as sources of materials (43%). The medicinal uses of plant species listed in Table 1 are given in Thomas and Vandebroek (2006). Among the 30 managed species with highest overall use values (for ns C3; Table 1), the majority are trees (18 species), followed by herbs (9 species). All eight managed palm species are included. The majority of the thirty species with highest medicinal use value are herbs (15 species), followed by shrubs (8 species), whereas among the top thirty most useful food plants, palms are again prominently present (5 species). The majority of the thirty most useful food species are trees (14 species). It is clear that homegardens and swiddens fulfil an important economic function for local people. In addition to the traditional food crops, which are used for subsistence only, almost all households grow coca (Erythroxylum coca) as a cash crop. Coca leaves are sold locally to Andean settlers of Quechua or Aymara descent who distribute it into established legal (coca leaf chewing) and illegal (cocaine production) market chains. Another economic plant that is cultivated by Yuracare´s and Trinitarios in homegardens and swiddens is the timber species mara (mahogany; Swietenia macrophylla King). This CITES appendix III species (www.cites.org) is the most valued commercial timber species in Bolivia (De Pourcq et al. 2009). At the time of research, the wild population of mara was (illegally) being depleted in the park by Yuracare´s and Trinitarios. Individuals from both ethnic affiliations typically formed little groups, generally composed of at least one (generally Quechua or Aymara) contractor who owned a chainsaw and fellow and/or neighbouring villagers (i.e. Yuracare´s, Trinitarios, or less
frequently Andean settlers). The contractors were paid to cut the trees, process them into commercial-sized boards on the spot, and transport the latter manually to the nearest truck stop. In response to perceived resource depletion on the one hand, and the high monetary value of the timber on the other, members of both ethnic groups started cultivating this species in homegardens and swiddens in recent years. These small-scale plantations were regarded as investments for the future: most adults declared that not they, but their children will be able to benefit from these trees. Some participants claimed to own hundreds of young trees. We have seen at least dozens. Propagation material is mostly obtained in the form of seedlings that are collected under the canopy of mother trees and transplanted in homegardens and swiddens. Also, small-scale seedbeds have been observed. These are laid out in dug plain soil under the branches of shrubs or small trees in order to provide shade and lower the impact of rain. Occasionally, seeds are germinated in individual containers. Relation between plant growth form and use value In relative terms, Trinitario and Yuracare´ participants use proportionally more managed herb species from their homegardens and swiddens than trees, shrubs or lianas. Yet, plant species with a tree habit have a significantly higher mean overall use value than herbs or shrubs (P \ 0.01, Kruskal–Wallis; Table 4). Overall use values of herbs and shrubs are equal. Of all managed plants, proportionally more herbs are used as herbal remedies (47%) than trees (27%), shrubs (23%) or lianas (2%). Also the mean medicinal use value of herbs is higher than that of other growth forms, but this difference is not statistically significant (Table 4). Of a total of 79 managed food plants, proportionally more species are trees (46%),
Table 4 Mean use values and standard deviations of main plant growth forms for different use categories Habit Tree (N = 46) Shrub (N = 35) Herb (N = 64)
UVme
UVs
UVfo a
2.32 – 1.39
0.50 ± 0.62
1.44 ± 0.76
a
1.44 ± 0.76
a
a
0.68 ± 0.62
a
0.82 ± 0.72
UVma
0.85 – 0.70
0.45 – 0.73 a
0.08 ± 0.24a
a
0.17 ± 0.38a
0.44 ± 0.63 0.30 ± 0.50
Highest values are marked in bold UVs overall use value, UVme medicinal component, UVfo food component, UVma materials component a
Indicates values that are equal (P [ 0.05; Mann–Whitney and Kruskal–Wallis tests)
123
Agroforest Syst (2010) 80:131–152
followed by herbs (29%) and shrubs (21%). In fact, 76% of all managed tree species have food applications: they produce aromatic leaves or flowers, fruits, seeds or palm hearts that can be eaten raw, cooked or processed into refreshing drinks, edible oil, or alcoholic beverages. Trees also have the highest food use value of all life forms (P \ 0.01, Kruskal–Wallis test). A total of 50 managed plants are used as sources of material, among which trees are best represented, followed by herbs and shrubs. Only one liana species is used for this purpose. Also the mean material use value of trees is significantly higher than that of herbs or shrubs (P = 0.03, Kruskal–Wallis; Table 4). In sum, plants managed in Yuracare´ and/or Trinitario homegardens are mostly herbs, followed by trees and shrubs. Nevertheless, trees have significantly higher overall use values than all other life forms. Most managed medicinal plants are herbs, whereas trees are the dominant life form among all managed edible and material species.
Discussion Plants growing in homegardens and swiddens Homegardens and swiddens are among the most prominent regimes of human disturbance. Approximately 21% of all useful plant species (151 out of 732 species) inventoried are managed in Yuracare´ and Trinitario homegardens and swiddens. These numbers are comparable to those found by Lamont et al. (1999) (161 species) and Padoch and De Jong (1991) (168 species) in respectively 51 and 21 gardens in the Peruvian Amazon. Recently, Perrault-Archambault and Coomes (2008) recorded 45–161 different species per village during a large scale inventory of 300 homegardens in a northern Amazon region from Peru. High species numbers in the relatively few Yuracare´ and Trinitario communities we surveyed might be explained by their high degree of isolation. Studies on homegardens in many areas indicate that species diversity is greater in remote villages, where homegardens are an important source of subsistence products, because markets for products are unavailable (Fernandes and Nair 1986). On the other hand, the relatively high diversity of Trinitario and Yuracare´ homegardens and swiddens might also be due to the fact that they represent a combination of plants
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brought together through the cumulative action of various ethnic groups (i.e. Yuracare´s, Trinitarios, Andean settlers) with distinct provenance, contact history, lifestyle, cultural heritage and customs, and hence, to some extent, different managed plant species (see Thomas 2009b; cf. Perrault-Archambault and Coomes 2008; Wezel and Ohl 2005; Zaldivar et al. 2002). For example, we have demonstrated elsewhere that Trinitarios rely more on managed plants as sources for herbal medicines and food than Yuracare´s who use proportionally more wild species in these use categories. Trinitarios also accredit higher overall and medicinal use values to managed plants than Yuracare´s and they use more than three times as many introduced species as medicines (Thomas 2009b). An in-depth discussion of factors influencing differences in plant use, knowledge and management of Yuracare´s and Trinitarios is given in Thomas (2009b). An example that demonstrates the exchange of managed plants between different ethnic groups in TIPNIS is provided by Porophyllum ruderale. This aromatic herb is typically used in the Bolivian Andes as a flavouring agent in food as well as for its medicinal properties. Therefore, it was most likely introduced in TIPNIS by Andean settlers who passed it onto Yuracare´s (Thomas 2009b). In Amazonian homegardens there appears to exist a preference for managing species that belong to certain plant families. Arecaceae, Solanaceae, Fabaceae and Rutaceae were recorded by Lamont et al. (1999) as the most important families in three Peruvian villages. These families belong to the top six of families in the present study. The most important family in terms of species within the toptwenty highest overall use value in our study is undoubtedly the palm family (Arecaceae). Palms have the well-established reputation of being highly useful plants for local people in Amazonia (Macı´a 2004). They provide edible fruits, seeds, oils, palm hearts, fibres, thatch, construction materials, domestic artefacts, tools for traditional hunting and fishing, medicines and other minor products (Thomas 2009b). In a study among the Huaorani of Ecuador, eleven palm species showed the highest use values within the 30 most valued woody plants (Macı´a 2001). The same trend is observed in many other ethnobotanical studies throughout Latin America (e.g. Phillips et al. 1994; Stagegaard et al. 2002), including the present one (Thomas et al. 2009c).
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Cultivation of Swietenia macrophylla as observed in Trinitario and Yuracare´ homegardens and swiddens is clearly not restricted to our study area and similar responses to resource depletion have been reported among Amazon-based Riberen˜o people from Peru. After depletion of major timber species (including S. macrophylla), Riberen˜os actively and autonomously engaged in planting, managing and protecting individuals of these species, leading to renewed healthy populations (Pin˜edo-Vasquez et al. 2002). Also Paz (1991) mentioned that Yuracare´s from the Rio Chapare region engaged in planting seven new seedlings for every S. macrophylla adult they felled. As we have personally been able to witness, cultivation of S. macrophylla through sowing and transplanting seedlings is quite successful among the Trinitarios and Yuracare´s that participated in the present study. This example immediately illustrates the potential of homegardens for countering local resource depletion. The proportion of cultivated (70%) and tolerated (30%) plants managed in Yuracare´ and Trinitario homegardens and swiddens is in agreement with observations made elsewhere. In a study of 30 Mexican homegardens, Blanckaert et al. (2004) found that 68% of species were cultivated and 22% tolerated. Van den Eynden (2004) calculated that on average 82% of all species occurring in 42 villages in southern Ecuador were cultivated while 14% were tolerated. Our data also corroborate the notion that plant management in homegardens is highly variable (Ban and Coomes 2004; Coomes and Ban 2004; Perrault-Archambault and Coomes 2008; Wezel and Ohl 2005). In the present study, 14% of all cultivated species were encountered in only one homegarden. This percentage is conservative in comparison with large-scale homegarden surveys. In her study on the edible plants of Southern Ecuador, Van den Eynden (2004) noticed that about 60% of all edible plants were only used or known in one of the 42 villages she investigated. Studies in Amazonian homegardens have shown that around one-third of species are found exclusively in one garden (Padoch and De Jong 1991; Perrault-Archambault and Coomes 2008). Usefulness of managed and wild plants Managed plants in Yuracare´ and Trinitario homegardens are most often used as sources of medicine and
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food, followed by materials and environmental uses (i.e. ornaments). These appear to be the main reasons why people manage plants in homegardens all over the world (Agelet et al. 2000; Bale´e 1994; Blanckaert et al. 2004; Finerman and Sackett 2003; Lamont et al. 1999; Miller and Nair 2006; Perrault-Archambault and Coomes 2008; Trinh et al. 2003; Van Den Eynden 2004; Vogl and Vogl-Lukasser 2003; Wezel and Ohl 2005; Zaldivar et al. 2002). This is unsurprising as one could expect usefulness to be the force that drives plant management (Clement 1999a). Nonetheless, far from all useful plant species are managed and many other useful species are obtained exclusively from natural habitats (see Thomas et al. 2009c). We agree with Bennett (1992) that reasons that impede people from managing or cultivating wild useful species can be related to (1) difficulties in plant establishment; (2) slow growth; (3) the fact that common and abundant species do not warrant protection; and (4) their minor actual importance. However, such arguments should ultimately be based on explanations by the local people themselves and we would in this respect recommend future studies to focus on participatory exploration of people’s decision-making processes and criteria. The fact that higher medicinal and food use values were obtained for managed plants than for wild plants, together with our observation that wild plants that typically grow in anthropogenic landscapes have a higher mean medicinal use value than wild species that typically grow in natural landscapes, supports the growing consensus on the importance of disturbed landscapes, and particularly homegardens and swiddens, in the provision of medicinal remedies (Bale´e 1994; Begossi et al. 2002; Etkin 2002; Frei et al. 2000; Gavin 2009; Posey 1984; Stepp and Moerman 2001; Voeks 1996, 2004) and (non-staple) food (Styger et al. 1999; Toledo et al. 1995; Van den Eynden 2004). The usefulness of moderately humanized landscapes as sources of medicinal plants is often explained by the existence of a causal link between the frequency and/or intensity of contact with certain species and their utility. As a product of human creation, the anthropogenic environment is most salient, most familiar and most accessible and therefore most likely to be learned, named and used (Brown 1985; Gavin 2009; Phillips and Gentry 1993b; Thomas et al. 2008, 2009a, b, 2009c; Turner 1988; Voeks 2004).
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A similar pattern is observed with respect to the medicinal uses of introduced plants. Introduced plants play an important role in Yuracare´ and Trinitario homegardens. At first, one might be surprised by the fact that nearly one-third of cultivated species are introduced. However, the literature describes plenty of cases for indigenous communities that corroborate the same trend (Bale´e 1994; Begossi et al. 2002; Eyssartier et al. 2009; Janni and Bastien 2004; Hanazaki et al. 2006). According to Voeks (2004), intercultural contacts with European settlers some five centuries ago led to an early but systematically underestimated (intentional and accidental) floristic homogenization. Useful plants from the Old World were actively and passively distributed over the New World tropics and subtropics as a consequence of colonial horticultural endeavours (Bennett and Prance 2000). By the mid 19th century, exotic fruit trees were fully incorporated into homegardens along the Amazon (Miller and Nair 2006). The Amazonian societies themselves contributed considerably to this floristic homogenization through the active exchange of plant material (Anderson and Posey 1989; Bennett 1992; Milliken and Albert 1997; Phillips and Gentry 1993b; Perrault-Archambault and Coomes 2008). Harris (1998, quoted in Bennett and Prance (2000) noted that ‘‘for many millennia, people of different cultural traditions living in different geographical regions have obtained useful plants from each other in a long drawn-out process of cross-cultural exchange… ‘‘Most Old World plants were introduced originally as foods or ornamentals and relatively few for their medicinal value. Through an ongoing process of ethnomedical experimentation, the medicinal power of many ornamentals and food plants were and most likely still are being ‘‘discovered’’ (Bennett and Prance 2000). The fact that the dichotomy between food and medicines is largely absent among (South American) indigenous and rural populations (Bennett and Prance 2000; Etkin 1994; Voeks 2004), may have played a beneficial role in the discovery of new herbal medicines. The results from this study largely confirm the previous arguments since 79% of all introduced cultivated plants with a food use are also used medicinally, whereas eight out of ten (80%) introduced cultivated ornamentals are used in traditional medicine. For native cultivates these values
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are 63 and 73%, respectively. Only two introduced cultivated plants with a local medicinal value have no additional ornamental or food use value, whereas for native cultivated plants this amounts to 15 species (i.e. 6 and 21% of all introduced and native cultivates, respectively). Hence, it seems that for native plants, the main reason for cultivation is less related to their ornamental or food value as compared to introduced plants. This result suggests that it is likely that relatively few species were purposefully introduced for their medicinal properties, whereas the medicinal applications of plants that were initially introduced as ornamentals and food species have gradually been learned and developed, partly due to a high intensity of contact with these species. Relation between growth form and use of managed plants Apparently, there exists a rough correlation between the life form of managed plants and their utility. Plants with a tree habit are more likely to be managed for their overall and food usefulness, whereas herbs are more popular as medicines. The fact that trees possess the highest mean overall use value of all managed life forms is basically due to their more complex habit and multipurpose nature: apart from fulfilling human needs for food and medicine they also contribute considerably as sources of materials and fuel (Tardio and Pardo-de-Santayana 2008). Also from a historical perspective, there seems to have existed a preference for managing plants with a tree habit in Amazonia. Sixty-eight percent of the 138 species of plants that were identified by Clement (1999a) to have been under cultivation or management at the time of European arrival in Amazonia, were trees or woody perennials. Of all services provided, in TIPNIS the most important use of trees is for food (mostly fruits). This might be due to a number of reasons. First, people’s interest in having diverse food sources spread throughout the year, might stimulate them to manage trees with staggered fruiting periods, as opposed to annuals which tend all to produce at once. Second, trees are long-lived and hence require less labour as compared to herbaceous food producing plants that need much more care (sowing, weeding, etc.). Third, food harvest quantity is in most cases considerably
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higher for individual trees as compared to herbs. These observations are corroborated by information from literature (Miller and Nair 2006). In a study on the use and management of edible non-crop plants in southern Ecuador, Van den Eynden (2004) demonstrated people’s preference for managing trees with edible fruits, as compared to other life forms. Fruit trees are also the most frequent plants in Peruvian and Spanish homegardens (Lamont et al. 1999; Agelet et al. 2000), while in Senegalese fields, indigenous trees are mainly managed for their fruits (Lykke 2000). By contrast, managed herbs in Yuracare´ and Trinitario homegardens have the highest mean medicinal use value of all life forms. We have demonstrated elsewhere that although Yuracare´s and Trinitarios use a higher number of woody species in traditional medicine, herbs are significantly overrepresented in their local pharmacopoeia (comprising both wild and managed plants; Thomas 2009b). We additionally showed that for the entire plant inventory the medicinal use value of trees was the lowest, whereas that of herbs scored highest of all life forms (Thomas et al. 2009c). The popularity of managing herbs for therapeutic uses might not be surprising. Plants in general owe their medicinal properties in humans to the chemical defence mechanism they apply against predators that attack them: mammals, insects, bacteria, fungi, and the like (Voeks 2004). Plants are known to use two main types of chemical defence strategies against predators (Stepp and Moerman 2001). According to a first scenario (apparency theory; Feeny 1976), plants reduce digestibility by producing and storing metabolically and generally non-toxic, inactive quantitative defences such as tannins and lignins. Mostly slowly growing plants with relatively long-lived leaves such as trees or shrubs make use of this strategy. The therapeutic value of such high molecular weight molecules in humans is rather low. By contrast, the products of a second plant defence mechanism have a higher probability of evocating biomedical effects in humans. These are low molecular weight secondary metabolites, such as alkaloids, terpenoids and cardiac glycosides. Opportunistic, rapidly colonizing and short-lived plant species such as herbs and pioneer species, tend to rely preferentially on such toxic and highly bioactive qualitative compounds (resource availability theory; Coley et al. 1985).
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Conclusions Homegardens and swiddens fulfil a central role in the livelihoods of contemporary Yuracare´s and Trinitarios from TIPNIS. In addition to representing the main staple food source, as well as supplying local people with numerous important goods and services such as medicines, food additives, snack foods, materials, environmental services, etc., they are also the ultimate loci for experimentation with plant management and use. Decisions to manage plants may be based on a variety of reasons such as to (1) increase their abundance or availability (e.g. after resource depletion in natural vegetation through overharvesting); (2) enhance their productivity; or (3) improve particular plant characteristics (e.g. plant selection based on nutritional value, content in bioactive compounds, etc.). Plant management seems to be partly growth form dependent for different plant uses in TIPNIS whereby trees seem more likely to be managed in Yuracare´ and Trinitario homegardens for their food usefulness as compared to other life forms, whereas herbs are more popular for use in traditional medicine. Trees owe their higher overall usefulness to their more complex habit and multipurpose nature, while the higher medicinal value of herbaceous plants in herbal medicine might partly relate to their chemical defence strategy. However, although people generally start managing plants in homegardens and swiddens because of their perceived usefulness, they also seem to be favourable locations to experiment with the usefulness of (managed or wild) plant species prevailing there. Our research suggests that as a consequence of a higher intensity of contact, species in homegardens and swiddens may gradually be attributed new uses through an ongoing process of experimentation. In line with these findings, we have reason to believe that the human selection process of managed medicinal plant species in TIPNIS is guided by both (1) the high intensity of contact with these species and (2) their chemical defence strategy. There are several indications pointing to the link between the frequency and/or intensity of contact with certain species and their utility in traditional medicine. First, wild plant species from disturbance habitats have a higher medicinal use value than wild species from natural environments. People spend most of their time in disturbance habitats and are therefore more
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likely to learn about the properties of plants that typically grow there. Also the fact that nearly all tolerated plants are used in traditional medicine might support the correlation between intensity of contact with and usefulness of plants. After all, it is likely that plants that are currently tolerated, used to be wild species typically prevailing in disturbance landscapes. As people progressively learned about the medicinal properties of these species––through increased contact––they gradually started managing them. Finally, our observation that most cultivated species that were initially introduced for their food or ornamental value have an additional medicinal use value likewise suggests that people gradually learned about their medicinal properties through an increased contact with these species as compared to plants from wild habitats. The higher usefulness of plants with herbaceous growth forms in traditional medicine seems, on the other hand, to have a phytochemical basis as they mainly rely on toxic and highly bioactive qualitative compounds such as alkaloids for their defence against predators. Hence we agree with Voeks (1996, 2004) that moderately humanized landscapes are an important source of healing for forest people, particularly because disturbance pharmacopoeias combine optimal foraging features with the natural distribution of promising plant-derived compounds. To conclude, a number of policy recommendations can be drawn from our investigation. •
•
The fact that different types of plant uses predominate in anthropogenic versus natural environments and that homegardens and swiddens can only fulfil part of the subsistence requirements of people in TIPNIS points to the livelihood importance of maintaining indigenous communities’ access to natural landscapes (also see Gavin 2009; Thomas et al. 2009c). Moreover, natural landscapes are important as a source of propagative material of managed, non-domesticated species. From a perspective of food security and risk management, it is important to conserve and promote the continuing use of the wide diversity of species that is currently managed by indigenous people from TIPNIS and to include diverse species in (agro)forestry support programmes.
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•
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Given that homegardens and swiddens vary widely, and management and use of plants is concentrated on certain individuals or families, there is a need for individual diagnoses and differentiated approaches between different garden and swidden tenders. This can then provide an indication of (1) the most important or promising species on which (agro)forestry and conservation programmes should focus; and (2) the potential scope of people’s practices as a tool for biodiversity conservation and (agro)forestry programmes. The fact that plants with different growth forms have differentiated usefulness points to the importance of valuing, supporting and promoting traditional agroforestry systems with diverse structures. Furthermore, it stresses the importance of valuing local people as researchers and focussing on anthropogenic environments when looking for plants with possible pharmaceutical applications.
Acknowledgements The present research was financed by a doctoral research grant of the Bijzonder Onderzoeksfonds (BOF) of Ghent University to Evert Thomas (Grant Number: B/03801/01 FONDS IV 1). Logistic support in Bolivia was provided by the Centre of Biodiversity and Genetics and the Herbarium Martin Cardenas of the Universidad Mayor de San Simon in Cochabamba. We are grateful to Reynaldo Berdeja, Kim Torfs, Jamie De Munk, Anouk Floren, Jurgen Ceuppens, Bert Wallyn and Olivier Beck for collaboration during data collection. Special thanks are due to all inhabitants of the indigenous communities San Jose de la Angosta, San Antonio, El Carmen de la Nueva Esperanza, Tres de Mayo and Sanandita for their kind assistance in this project. We are indebted to the professional botanists who identified several collections. They are C. Berg, I. Jime´nez, R. Liesner, J. Lombardi, P. Maas, M. Moraes, M. Nee, H. Rainer, R. Swennen, C. Taylor and J. Wood. Thanks also go to Ina Vandebroek, Paul Goetghebeur and two anonymous reviewers for commenting on earlier drafts of this article.
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