Palm uses in northwestern South America: a quantitative review.

Introduction

The great and quantitatively dominant ethnobotanical importance of palms (Arecaceae) in comparison to other botanical families in tropical American forests is well documented (Prance et al., 1987; Pinedo-Vasquez et al., 1990; Phillips & Gentry, 1993; Galeano, 2000; Macia et al., 2001; Lawrence et al., 2005). Palms have great cultural and economic importance among rural indigenous and peasant populations in tropical America (Schultes, 1974; Bodley & Benson, 1979; Balick, 1984; Balslev & Barfod, 1987; Balde, 1988; DeWalt et al., 1999; Galeano, 2000; Macia, 2004; Paniagua-Zambrana et al., 2007; Brokamp et al., 2011), and numerous studies have described their essential role in covering basic needs for human subsistence, such as for food and house construction (L6pez-Parodi, 1988; Morcote-Rios et al., 1998; Coomes & Burt, 2001; Campos & Ehringhaus, 2003; Macia, 2004; PaniaguaZambrana et al., 2007). Their great importance is closely related to their ecological, morphological, physiological and bromatological characteristics, Palms are conspicuous and abundant in many different tropical forest types (Henderson et al., 1995; Macia & Svelming, 2005; Balslev et al., 2011) and they are distributed in all forest strata and soil types (Kahn & de Granville, 1992; Kahn & Henderson, 1999; Vormisto, 2002b; Balslev et al., 2010a; 2011). Palms have distinctive morphological and physiological characteristics: they have straight and generally un-branched stems, large pinnate leaves, a vascular system with living cells throughout the plant's lifespan, they produce many adventitious roots, and are highly durable (Balick, 1984; Tomlinson, 2006). Their fruits and seeds contain starch, essential amino acids, and oils that are rich in polyunsaturated fatty acids (Balick, 1984; Balre, 1988; Moraes et al., 1996; Olvera-Fonseca, 2004).

Their extensive use and relatively well known taxonomy have facilitated their utilisation as model organisms for analysing the influence of ecological and socioeconomic variables on interrelations between humans and plants in tropical American ecosystems (Uhl & Dransfield, 1987; Henderson, 1995; Henderson et al., 1995; Borchsenius et al., 1998; Moraes, 2004; Govaerts & Dransfield, 2005; Dransfield et al., 2008; Galeano & Bernal, 2010). Several studies have shown that humans tend to use to a higher degree those species that are widespread and conspicuous (Ruokolainen & Vormisto, 2000; Byg et al., 2006). However the most used species are not always the same as those with greatest ecological importance (Boom, 1986). Moreover, higher species diversity in one region does not necessarily imply a greater use by the local population (Byg et al., 2007). The most abundant species in an ecosystem are often used, although with different intensities, and some of the more important species are managed to obtain a better use of their products (Paniagua-Zambrana, 2005; Byg et al., 2006; Bernal et al., 2011). Palms are used more by human groups which have more limitations in their access to markets (Byg & Balslev, 2004; Macia, 2004; Byg & Balslev, 2006; Byg et al., 2007; Paniagua-Zambrana et al., 2007). Recent studies have shown that indigenous people usually posses greater ethnobotanical knowledge than other human groups (Campos & Ehringhaus, 2003; Byg & Balslev, 2004; De la Torre et al., 2008).

Despite the large number of publications on traditional use of palms in tropical America, at a local or regional scale, comparative studies presenting a general vision of the use of palms at a larger geographical scale, and comparing use patterns in different biogeographic regions, habitats, countries or human groups, have not been done so far. Here, we present an exhaustive revision of the use of palms in north-western South America, where an effort has been done to compile existing bibliographical references at the local scale and of limited diffusion. The specific aims of this compilation are: (a) to quantitatively evaluate the use of palms in north-western South America, comparing different ecoregions (the Amazon, Andes and Chocd) and countries (Colombia, Ecuador, Peru and Bolivia); (b) to compare palm use patterns in different ethnobotanical use-categories and subcategories; (c) to analyze differences in the use of palms among different human groups (indigenous, mestizos, afroamericans and colonos) and compare the knowledge between different indigenous groups; and (d) to identify the most important palm species for the local populations living in the tropical forests of the study region.

Methods

Study Area

We compiled ethnobotanical information for palms occurring in the Amazon and Andes of Colombia, Ecuador, Peru and Bolivia, and the Choc6 of Colombia and Ecuador (Fig. 1). The Amazon ecoregion was defined as the lowlands to the east of the Andes up to 1,000 m elevation (Rennet et al., 1990; Jorgensen & Ledn-Yfinez, 1999). Data for species existing in all broad forest types were included: both welldrained terra firme forests and floodplain forests, and poorly-drained swamp forests. The Andes ecoregion was defined as the montane forests on both slopes of the Andes above 1,000 m, including the forests of the inter-Andean valleys of Bolivia with lower precipitation (Beck et al., 1993). The Choc6 ecoregion was defined as the area of humid forests along the Pacific coast of Colombia and northern Ecuador.

[FIGURE 1 OMITTED]

Bibliographical Search and Data Organization

A thorough bibliographic revision was performed to search for international and national publications for each of the four countries, including ethnographical publications with data on the uses of palms, when species identification was clear. Three categories of publications were selected. The first included publications based on original data gathered from fieldwork, including scientific papers, books, monographs, book chapters, and graduate, masters and doctoral theses. The second category included review publications for which we checked that data had not been previously published, in order to avoid duplication of information. The third type included publications based on herbarium material which included ethnobotanical information that was not included in any publications (Borchsenius et al., 1998; Moraes, 2004; Moreno Suarez & Moreno Suarez, 2006).

A database was constructed in Microsoft Access. For each publication, the following information was included (when available): scientific name of the species as it was published, country, ecoregion, human group, assignation to categories and subcategories of use, plant part used, and vernacular names. To unify the nomenclature of the scientific names, the world checklist of palms was followed (Govaerts & Dransfield, 2005; Govaerts et al., 2006). The only exception was Cero[C],lon peruvianum, which was recently described (Galeano et al., 2008), and therefore was not included in the checklist. The three broad habitat types in the Amazon ecoregion were lumped because most bibliographical references did not specify which of the three regions they referred to. Four human groups were recognized: (a) Indigenous, original population of a particular geographic region; (b) Mestizo, population of mixed origin, born from a father and mother of different race, generally white-indigenous; (c) Afroamerican, population of black race descendant of African slaves brought to America; in the study area they only live in the Choc6 of Colombia and Ecuador; and (d) Colono, native to an ecoregion different to the one where they presently live due to recent migrations (e.g. Andean people living in the Amazon). Those use-reports where no indication of human group was mentioned were classified as "Not identified." The Ecuadorean Quichua and Shuar indigenous groups living in the transition between the Amazon and the Andes ecoregions were considered Amazonian groups because the majority of existing literature referring to them is from that ecoregion. Mixed populations of two or more indigenous groups were considered as a single group for analysis, but were not computed as a different ethnic group.

All uses recorded from the literature were classified in 10 ethnobotanical categories that were further divided into subcategories (Table 1). When the ethnobotanical information was not classifiable within the previous subcategories, it was assigned to the subcategory "Other". In the Medicinal and Veterinary category we also used the term "Not Specified" when a medicinal use description contained insufficient information to assign the use to one of the described subcategories. The vernacular names of all palm species cited in the bibliography were compiled, independent of the existence of ethnobotanical information.

Data Analysis

All data analysis were performed at the species level and thus the ethnobotanical information obtained for infraspecific taxa (i.e., subspecies or varieties) were lumped into the corresponding species. Ethnobotanical data recorded only at the genus level (460 use-reports) were excluded from the analysis, and seven references recorded palm uses only at this level (Acosta-Solis, 1948; Forero, 1980; Toumon et al., 1986; Salick, 1989; Fujisaka & White, 1998; Vasquez, 2000; Bussmann & Sharon, 2006).

To analyse the uses of palms in different ecoregions, countries and human groups, the term "palm use" for a given species was defined as the use associated to a use category and use subcategory for a specific plant part. To analyse the abundance of palm uses, the term "use-report" was defined as the palm use described previously in one bibliographical reference.

To quantify the relative importance of the different use categories, the percentage of useful species for each category with respect to the total number of species used per ecoregion or country was used. In the case of the use subcategories, the percentage of useful species in each subcategory with respect to the total species used in the associated category was used.

To have an estimation of the ethnobotanical knowledge that exists in both different countries and ecoregions with respect to the total number of palm species, the percentage of useful species was calculated following the catalogue of Pintaud et al. (2008). To have an estimation of the number of indigenous groups with ethnobotanical information with respect to the existing total number of indigenous groups in the study area, the percentage of indigenous groups was calculated following Lewis (2009). In these latter calculations, the linguistic variants for the denomination of the same indigenous group were not considered.

To identify the most important useful species in each ecoregion, the Relative Importance (RI) index was calculated: RI-NUC+NT, where NUC=number of use categories in which a given species is used, divided by the total number of use categories of the most versatile species; NT=number of total use subcategories in which the cited species is found, divided by the total number of use subcategories that the most versatile species obtained (Bennett & Prance, 2000; Albuquerque et al., 2006). The maximum RI value that a species could obtain was 2. This index values the importance of the different species as a function of their versatility, without considering data relative to the number of bibliographic citations (Cartaxo et al., 2010).

Results

Palm Use by Ecoregions and Countries

A total of 194 useful palm species (representing 63% of the potentially existing species in north-western South America), 2,395 different uses, and 6,141 use-reports were found in the revision of 255 bibliographical references, including 95 palm use monographs (Table 2; Appendix). The average (+ SD) number of different uses per species was 12.3 ([+ or -] 18.7) although great variability was observed between different species. Ethnobotanical information was recorded for 54 indigenous groups, which represents 49% of the total indigenous groups living in the study area (Lewis, 2009; Fig. 1).

The Amazon was the ecoregion with the highest values in all the variables compared: 134 useful species (90% of those potentially present), 82% of total different uses, 84% of total use-reports, an average ([+ or -] SD) of 14.7 ([+ or -] 20.0) uses per species, and 81% of total bibliographic references found (Table 2). Ethnobotanical information was found for 48% of all indigenous groups living in the area.

Comparing the Amazon ecoregion in each of the four countries (Colombia, Ecuador, Peru and Bolivia) independently also gave the highest values in all the variables analysed (Table 2). The highest number of useful species, different uses and bibliographical references was found in the Peruvian Amazon, but the number of uses per species was the lowest there. In the Ecuadorean Amazon we found both the highest percentage of useful species and the highest percentage of indigenous groups studied, while the lowest percentages for these variables were found in the Amazon of Colombia and Peru.

In the Andes and Choc6 ecoregions similar results were found for many of the studied variables, including number of palm uses, percentage of useful species, and number of bibliographic references (Table 2). However, even if the Andes ecoregion was slightly more diverse in useful palm species than the Choc6 ecoregion (68 vs. 52), the average of different uses per species was highest in Choc6 (6.7 vs. 5.1), where we found a higher number of use-reports (569 vs. 439). In general, the Andes was the second ecoregion in relative importance for Ecuador, Peru and Bolivia, whereas in Colombia the Choc6 was more important than the Andes. The Andean region of Bolivia was the best studied of the four countries since in the Andes of Colombia, Ecuador and Peru no information was recorded for any indigenous group.

Colombia was the country with the highest number of useful palm species (105) and the highest number of indigenous groups for which ethnobotanical information about palms has been published (23), although the proportion of indigenous groups studied with respect to the total groups for the country was moderate (49%), and inferior only to Peru (Table 2). However, in Colombia the percentage of useful species in relation to palm species richness was the lowest of the four countries (48%), as was the average number of uses per species (7.8 [+ or -]10.1). Colombia was the country with the second lowest number of bibliographical references referring to palm uses, but the number of different palm uses and use-reports was comparatively high, only surpassed by Ecuador.

In Ecuador, the number of useful species found was slightly lower than for Colombia (103 vs. 105). Nonetheless, it was the country with the highest values for many of the variables compared: the highest number of different uses (936), use-reports (2010), percentages of useful species in relation to the palm species richness of the country (79%), percentage of indigenous groups studied (83%) and number of bibliographical references (81), including 31 palm monographs (Table 2).

Peru had intermediate values for most analyzed variables (Table 2). It was the country with the second highest proportion of useful species relative to the total palm species number in the country (76%), and also with respect to the number of bibliographical references with information on palm uses (74). And while it was the country with the second highest number of indigenous groups with published ethnobotanical information about palms (18), the percentage of indigenous groups studied (38%) was the lowest of the four countries.

Bolivia had the lowest values in most of the analyzed variables except for the average number of uses per species, which was the highest of the four countries (10.6+ 14.7) (Table 2). The percentage of useful species in relation to the total palm species richness in the country (74%) and the percentage of indigenous groups with published ethnobotanical palm information (61%) were the second most important. Concerning the published bibliographical references, a great number of palm use monographs were registered compared to other countries.

Palms in Different Use Categories and Plant Parts Used

In general, the use categories and subcategories with most useful species were the same as those with most use-reports. The main exceptions to this were (a) the Agroforestry subcategory in the Environmental Uses category, where the number of use-reports was frequently higher than the number of species found (e.g. overall 48% of the species and 53% of the use-reports; in Colombia 48% vs. 94%), as well as (b) in the Firewood subcategory in the Fuel category (e.g. in Colombia, especially in the Choc6 ecoregion, 88% of the species and 94% of the use-reports). Therefore, to facilitate the interpretation of data in Table 3, only percentages of useful species for different use categories and subcategories are shown.

The main uses of palms in north-western South America were in the categories Human Food (70%), Utensils and Tools (66%), Construction (63%) and Cultural Uses (56%) (Table 3). The categories Animal Food (37%), Medicinal and Veterinary (35%), Environmental Uses (35%) and Fuel (22%) had the lowest numbers of useful species. In the initially proposed Toxic category, there were no use-reports. All parts of the palms had some ethnobotanical use, although the most used parts among all use categories (except in Construction) were fruits, stem and leaves (Table 4). For 9% of the use-reports, the plant part used was not indicated.

At the ecoregion level, the percentages of palm uses were higher in the lowlands (the Amazon and Chocr) than in the Andes for the majority of the use categories (Table 3). In the Amazon, the relative importance of the different use categories was similar to the general pattern previously described, with the only exception of Medicinal and Veterinary, palms, which were more important than Animal Food (46% vs. 43%). The Choco ecoregion had the same pattern of palm use as the Amazon, except in the Environmental Uses category, which was more prevalent (35% vs. 29%). In the Andes, the general pattern of palm use described above was also found, but with some notable exceptions: the Construction category had greater relative importance in the Andes compared to the lowlands, the Environmental Uses category, such as in Chocr, had more relevance than in the Amazon, and the Utensils and Tools category was less important in the Andes than in it was in the lowlands.

At the country level, the categories Human Food, Construction, Utensils" and Tools and Cultural Uses were, in this order of importance, the ones that presented the highest percentages of useful species, except in Colombia, where Utensils' and Tools" was the most important category (62%) and Human Food occupied the fourth position (52%) (Table 3). In Colombia and Peru, the greater relative importance of the Medicinal and Veterinary category was notable compared to Ecuador and Bolivia, although the latter country had the highest percentage of use-reports for this category. The categories of Animal Food and Environmental Uses had varying importance in the different countries, without a recognizable pattern. Finally, the categories of Fuel and Other Uses were the least important in all countries.

Human Food. Palm uses in the different subcategories of Human Food were similar for all ecoregions and countries (Table 3; Appendix). Over 90% of the species were used as food or snack and more than 25% were used to prepare fermented or unfermented drinks, such as leche or chicha especially in the Amazon. Preparation of oils for human consumption was very important throughout the study region, though most prominent in the Choc6 (34%) and Bolivia (33%). The use of palms for food additives was more prominent in the Amazon (11%) compared to other ecoregions, and at the country level in Bolivia and Colombia (14% and 11%, respectively). The palm parts most often used in this category were fruits (61%), palm heart (20%) and seeds (12%) (Table 4).

Utensils and Tools. In all ecoregions and countries, most species (77%) were used to make several objects for domestic use, such as hammocks, fans, carrying bags, baskets or mats (Table 3; Appendix). The second most important activity was the construction of tools for hunting and fishing (56%), including bows, arrows, harpoons and different types of traps, although this category had lower importance in the Andes (27%). The manufacturing of tools for cultivation in their fields (chacras or chagras) and homegardens was more important in the lowlands than in the Andes, especially in Ecuador (20%). The use of palm leaves for wrapping food or other objects was mostly recorded in the Amazon of Ecuador and Peru. Rope manufacture was less important, but uniform, for all ecoregions and countries, except in the Colombian Andes and Choc6. The subcategory Other uses had high values because many use-reports simply described the use as 'handicrafts' or 'ivory' (for instance, the use of Phytelephas seeds which were also used as handicrafts), and therefore could not be precisely assigned to a particular subcategory. The most important palm parts used for utensils and tools were the stem (30%), leaves (20%) and immature spear leaves (11%) (Table 4).

Construction. In this category, most species (>76%) were used for thatching houses and for temporal sheds in all ecoregions and countries (Table 3; Appendix). In second place was the use of palms in the construction of different house parts, such as beams, walls, floors or materials for the roof. The use of palms for construction of canoes was particularly relevant in the Choc6 (14%) and the Amazon (7%), and for construction of bridges in the Amazon (9%). In the subcategory Other Uses, many use-reports only mentioned 'construction', which is a general term, for which reason the use could not be assigned with precision to a particular subcategory. The most used palm parts were the leaves (53%) and the stem (36%) (Table 4).

Cultural Uses. The most important cultural use in all ecoregions and countries was for ritual purposes, including festivals and feasts, particularly in the Andes (62%), and among the countries in Ecuador (56%) and Bolivia (52%) (Table 3; Appendix). In the Amazon (57%) and particularly in Colombia (60%), the recreational use of palms for the manufacture of musical instruments and toys, and for the preparation of ashes from several palm parts to be used in the traditional consumption of tobacco (Nicotiana spp.) and coca leaves (Erythroxylum coca) were of great importance. The use of palms for personal adornment, such as necklaces, bracelets, armbands, pectorals or earrings, had great importance in the Amazon (51%), and at the country level in Ecuador (50%). In the manufacture of cloth and accessories, like hats or buttons, and in the preparation of cosmetics, the highest importance was recorded in the Amazon, and at the country level in Bolivia (37% and 48% for the first subcategory and 33% and 48% for the second, respectively). The use of palms to produce natural dyes was minor, but it was registered in all ecoregions and countries, and was of particular importance in the Choc6 (17%). The most used palm parts were the entire leaves (18%), seeds (17%), and the fruits (14%) (Table 4).

Animal Food. The highest percentage of species used for Animal Food were used as wildlife attractant for hunting (72%), particularly in the Amazon (79%), and among the countries in Ecuador (91%) and Colombia (76%) (Table 3; Appendix). However, in the Andes and Choc6 the use of palms as fodder had greater importance (57% and 50% respectively), and particularly in Bolivia (50%). The use of palms as fish bait had high values in the Amazon (30%) and in Peru (52%). The fruits were clearly the palm part most used (77%), followed by the seeds (5%) and leaves (4%) (Table 4).

Medicinal and Veterinary. Medicinal uses were found in all the proposed subcategories and were especially important in the Amazon, where the highest percentages were recorded for most subcategories (Table 3; Appendix). The highest percentage of medicinal species (56%) was registered for the treatment of ailments of the digestive system (e.g. stomach pains and diarrhoea), particularly in the Andes and the Amazon (55% and 54%, respectively) and, among the countries, for Peru (64%). The treatment of respiratory ailments, in particular colds and catarrh, were very important in the Amazon (39%) and Bolivia (48%). The use of palms to treat general common ailments of nonspecific character, such as headaches, general discomfort and body pains, was the subcategory with the third highest percentage of useful species (34%) in particular in the Amazon, and among the countries in Peru and Bolivia. Similarly, the treatment of infectious and parasitic diseases was most prominent in the Amazon (33%) and in Peru (42%). The percentages of palms used for skin and subcutaneous ailments were higher in the Amazon and Andes than in the Choc6, and among countries its use was highest in Bolivia (38%). The treatment of ailments and injuries of the muscular-skeletal system such as traumatisms, bone fractures, dislocations or sprains were more relevant in the Amazon (23%) and in Bolivia (33%). Palms were also used as antidotes against snakebites, scorpion stings and ant bites and stings, especially in the Amazon (21%), and in Colombia and Bolivia (24% in both cases). The percentage of palm species used for treating diseases of the reproductive system and for sexual health was higher in Choc6 (25%) than in the other ecoregions, and among countries in Peru and Bolivia. Palms were also used to treat less well defined diseases, such as aire, evil eye, and arrebato, especially in the Choc6 (25%) the Andes (18%), and in Bolivia (19%). For the treatment of blood and cardiovascular system ailments, the highest percentage of species was reported for the Choc6 (17%) and in Bolivia (14%). The percentage of species used in both diseases and ailments of the urinary system, like cystitis, and the treatment of problems relating to pregnancy, birth and puerperium was highest in the Andes in both cases (27% and 18%, respectively). For the treatment of dental problems, diseases of the endocrine system, metabolic and nutritional problems, and for veterinary use, medicinal palm species were only reported in the Amazon, and the greatest percentage was registered in Bolivia (10% in all cases, except in the subcategory Veterinary Uses which was 5%). For treating diseases and ailments of the nervous system, mental health and sensory system, the highest percentage of useful species was found in the Andes (9% in both cases) and in Ecuador (9% and 6%, respectively). The subcategory Other Uses included species with medicinal uses which could not be assigned to a described subcategory, for example for the treatment of cancer, hernia, or when the nature of an illness was not specified. The most used plant parts in popular medicine were the roots (31%), fruits (24%) and seeds (12%) (Table 4).

Environmental Uses. The main use in the category Environmental Uses" was as ornamental plants for all ecoregions and countries except Peru, with special importance in the Andes (68%) and among the countries in Ecuador (67%) (Table 3; Appendix). The use of palms in agroforestry systems with different degrees of management ranked second, particularly in the Amazon (58%) and in Peru (71%), where it was the most important use. The use of palms as natural barriers and to delimit properties was used in all ecoregions and especially in Bolivia and Peru. The

use of palms to improve soils was only registered in the Amazon of Peru and Bolivia. In this category the whole plant (65%) and the stem (21%) were mostly used (Table 4).

Fuel. The majority of the species were used for firewood in all countries and ecoregions, especially in the Amazon (85%) and among the countries in Bolivia and Colombia (89% and 88%, respectively) (Table 3; Appendix). The palms had notable importance as fire starters and as torches, candles, and lamps, particularly in the Chocr. Within the subcategory Other Uses, the use of palm leaves for burning and water-proofing canoes was important in the Amazon, particularly in Peru. The predominant parts used were the stem (52%), leaves (22%) and the fruits (8%) (Table 4).

Other Uses. The highest percentage of useful palms in all ecoregions and countries was related to the use of the larvae of the Rhyncophorus palmarum (Coleoptera) for human food (66% of total species), medicinal use, and as fish bait (Table 3; Appendix). These larvae develop mainly in rotting palm stems. The remaining uses are miscellaneous. The plant parts mostly used were the stem (65%), seeds (8%) and fruits (6%) (Table 4).

Palm Uses by Different Human Groups

Indigenous groups clearly used palms more prominently than other human groups. They presented the highest palm use values: number of useful species (129), different uses (1,555), use-reports (3,713), and higher average number of uses per species (12.1 [+ or ] 16.7), although they were also the best studied human group (166 bibliographical references) (Table 5). The Amazon was the ecoregion with the highest values in all countries and for all human groups, except for the mestizos in Ecuador. In the Chocr, the indigenous groups recorded higher values for all variables compared to the Andes of Colombia and Ecuador, although in Ecuador the differences between these ecoregions were small.

The mestizos were the second human group in terms of palm use values (Table 5). The Amazon was the ecoregion with the highest values, with the exception of Ecuador, where the Andes had a greater importance. Peru was the country with the highest number of bibliographical references. No use-report was found for mestizos in the Choc6 ecoregion or Colombia.

A greater number of useful pahns were registered for the afroamericans, when compared to colonos (Table 5). Colombia reported higher values than Ecuador for all the variables analyzed, and these values were similar to those registered for the Colombian indigenous groups of the same ecoregion, despite having a lower number of references.

The colonos presented the lowest values of all groups compared in all countries, except for the average number of uses per species, which was slightly higher than for afroamericans (Table 5). Most information was registered in the Amazon, and among the countries in Colombia and Ecuador.

Finally, it is worth mentioning that for all ecoregions and countries very high values were registered for unidentified human groups, since the bibliographical information was not precise. Curiously, they registered the highest number of useful species (170) (Table 5).

Palm Uses by Indigenous Groups

Ethnobotanical information concerning palms was found for 54 indigenous groups: 47 in the Amazon ecoregion, two in the Andes, and five in the Choc6 (Fig. 1; Table 2). There was great variation in the ethnobotanical knowledge of palms for the different indigenous groups, and the greatest knowledge was observed in Ecuador for all three ecoregions (Table 6). In general, the most studied indigenous groups were also those with the greatest observed ethnobotanical knowledge. For example, for some indigenous groups (e.g. Quichua, Huaorani or Shuar in Ecuador) many useful palm species, different uses and use-reports were found, but this may be because they were particularly well studied ([greater than or equal to]13 bibliographical references per group). However, other indigenous groups (e.g. Muinane of Colombia or Cocama of Peru) also had many useful palm species, different uses and use-reports, but these were described in just two bibliographical references.

A total of 1,933 vernacular palm names were registered (including orthographic variants), corresponding to 178 palm species. Of these names, 33% were in Spanish (158 species) and 67% in different indigenous languages (130 species).

Outstanding Useful Palm Species by Ecoregions

In general, the species with the highest relative importance values also had the highest number of palm uses, use-reports and bibliographical references (Table 7). Five species were found as the most important in all three ecoregions: Bactris gasipaes, Iriartea deltoidea, Oenocarpus bataua, O. mapora and Socratea exorrhiza. In the Amazon ecoregion, the most important genera were Astrocaryum, Attalea, Oenocarpus and Phytelephas, each with more than one species with the highest relative importance whereas in the Choc6 three of them, Attalea, Oenocarpus and Phywlephas, were among the most important. In contrast, in the Andes the genera Ceroxylon, Oenocarpus and Parajubaea were the most important. In the Amazon and Choc6 ecoregions, the most versatile species were used in all countries, but in the Andes the most important species did not have such a broad geographical range of use, with the exception of Bactris gasipaes, which was registered in all four countries.

Discussion

Areeaceae is probably the most important plant family in the Neotropics, in terms of use diversity and abundance. Palms are widely used for a great number of purposes throughout all ecoregions and by all human groups in north-western South America. The use of palms has been documented in several monographs with local or national scope (e.g. Balslev & Barfod, 1987; Bemal, 1992; Borchsenius et al., 1998) and in numerous cthnobotanical studies with diverse indigenous groups (e.g. Boom, 1986; Kronik, 2001; Macia, 2004), mestizos (e.g. Mejia, 1988; Stagegaard et al., 2002; Balslev et al., 2008), afroamericans (e.g. Galeano, 2000), and colonos (e.g. Flores Paitfin, 1998). The present quantitative revision underlines the great importance of comparative ethnobotanical studies at a regional geographic scale, and call attention to many different uses and species consistently shared between different human groups across the western Amazon, the Choc6 and the Andes ecoregions.

The use of palms is not random since their main uses are the same in different ecoregions and countries: palms are mostly used for human food, for manufacture of objects and utensils of domestic use, and for the construction of houses. This underlines their fundamental role in satisfying basic subsistence needs of rural indigenous and peasant population of north-western South America, in the same way that previous studies have demonstrated the importance of palms on local scales (Galeano, 2000; Narvaez et al., 2000; Gertsch et al., 2002; Campos 8,: Ehringhaus, 2003; Macia, 2004; Paniagua-Zambrana et al., 2007). Palms also have great importance in different cultural practices, which also confirms at regional scales the results of previous papers showing the cultural importance of palms for some ethnic groups in South America (Schultes, 1974; Bodley & Benson, 1979; Gertsch et al., 2002).

The enormous importance of palms in the Amazon can be explained by two complementary factors. On the one hand, their high species diversity allows access to a wide array of potential resources (Begossi, 1996; De la Torre et al., 2009; Brokamp et al., 2011), and on the other hand, the great diversity of indigenous groups favours a highly distinctive ethnobotanical knowledge (Campos & Ehringhaus, 2003). Although the Amazon was clearly the best studied ecoregion, ethnobotanical studies (that include palms) have so far only been conducted among less than 50% of the remaining indigenous groups. Likewise, in the Andes and the Choc6, ethnobotanical knowledge of palms is even more restricted, and for more than 50% of the species in both ecoregions, no uses have been documented.

We found that Ecuador is the best studied of the four countries in all ecoregions. There, and to a lesser degree in Bolivia, the percentage of useful palm species and the percentage of indigenous groups with documented palm uses were higher, which indicates that the use of palms is comparatively better documented than in Peru and Colombia. The high average number of uses per species recorded in Bolivia could be explained by the higher number of palm monographs from that country. Following this thinking, Peru and Colombia would be less known in palm ethnobotany than the two other countries. In Peru, the high number of indigenous groups for which we do not have ethnobotanical information underlines that the available data on palm uses remains incomplete. In Colombia a very low percentage of useful species was recorded, even though it is the country with the highest species richness. All this points to the need for more studies to complement the ethnobotanical knowledge on palms in all three ecoregions, but particularly in the Choc6 where a great richness of potentially useful species has been reported (Galeano & Bernal, 2010).

Previous studies suggested that indigenous people possess a greater knowledge about the uses of palms than mestizos or colonos possess in north-western South America (Campos & Ehringhaus, 2003; Byg & Balslev, 2004; Byg et al., 2007) and our paper reinforces this conclusion. This is the result of a complex set of interactions between diverse factors, including: (a) historical ones, since a long occupation of a territory facilitates the development of extensive ethnobotanical knowledge, (b) cultural ones, based on hundreds of years of orally transmitted traditional ecological knowledge, and (c) economic ones, in particular by the reduced degree of access to markets which mean they use palms for subsistence and are not able to purchase palm products substitutes (Alcorn, 1981; Balre, 1994; Byg & Balslev, 2004; Byg et al., 2007; Paniagua-Zambrana et al., 2007). Moreover, indigenous knowledge is highly differentiated, even between ethnic groups that occupy nearby geographical areas and share similar resources such as palms (Campos & Ehringhaus, 2003), or medicinal plants (Shepard, 2004; Collins et al., 2006). In general, the best studied indigenous groups had a richer and more diversified ethnobotanical knowledge. But, not only the number of publications existing for each group is important, so is the existence of monographs on palms that contribute to a greater degree to the number of useful species and different uses. This again underlines the need for more ethnobotanical studies focusing on palms, as information is nonexistent for over 50% of the indigenous groups in north-western South America.

The traditional knowledge of mestizos should not be undervalued, since it is equally diverse and even complementary to that of indigenous groups in several use categories (see also De la Torre et al., 2008). Frequently, mestizos have a long settlement history, which allow them to develop a profound ecological knowledge in their environment, which may be similar to those of various indigenous groups. In our study, the number of palm uses for mestizo people could probably be higher, because many publications do not mention explicitly the human group studied, and it is likely that many of these publications refer to mestizos.

The Colombian afroamericans, who have been better studied than the Ecuadorean afroamericans, had a similar level of knowledge of palms as did indigenous groups in the Choc6 ecoregion. This can be explained by the group's long history of residence and their prolonged contact with indigenous people in this region (Mendoza et al., 1995).

Some palm species have an enormous importance due to their large number of different uses. This uneven distribution in their uses has also been registered in previous studies (Campos & Ehringhaus, 2003; Macia, 2004; Byg et al., 2006; Paniagua-Zambrana et al., 2007; Balslev et al., 2010b). Such species are often trees that are relatively abundant in the different habitats, due to their wide ecological amplitude (Ruokolainen & Vormisto, 2000; Byg et al., 2006; Balslev et al., 2011). The preference for certain uses can be interpreted as the result of a number of factors, including easy accessibility to the species, larger quantities of resources available, and the potentially greater sustainability of their use under minimum management (Byg et al., 2006; Bernal et al., 2011). These multi-use species play a fundamental role in the local subsistence strategies and represent key cultural species (Garibaldi & Turner, 2004; Balslev et al., 2010b).

The unequal number of bibliographic references and monographs that refer to the different variables analyzed (ecoregions, countries, human groups, indigenous groups, and palm species), certainly limits the strength of the conclusions that can be drawn. Nevertheless, the variables with higher use-reports showed a more intense and diversified use of palms.

As a result of the experience gained in this palm use revision, we suggest a more precise ethnobotanical data collection that would include: (a) making an effort to identify plants to the species level; (b) writing vernacular names carefully and indicating the language in each case; (c) gathering information from different uses as completely as possible in order to subsequently classify uses within at least two levels of utility (category and subcategory); (d) noting the plant part used for each different use; (e) specifying the human group and/or ethnic group from which the information was gathered; (f) obtaining detailed geographical information of the study area, including forest types or habitats; and lastly, (g) in the case of medicinal species, writing precisely the medicinal indication, mode of preparation and ways of administration for each case.

Acknowledgements We kindly thank Joaquina Alban, Rodrigo Bernal, Roxanna Castaneda, Lucia de la Torre, Gloria Galeano, Eva Ledezma, and Laura Mesa for their assistance in searching for bibliographical references, Bob Allkin, Bill Baker and Anders Barfod for their help in the construction of our database, and Patricia Balvanera, Rodrigo Berual, Jamie Nicole Cotta, and Lucia de la Torte for their helpful comments on an earlier draft of the manuscript. This study was funded by European Union, 7'h Framework Programme (contract no. 212631) for which we are grateful. Henrik Balslev also acknowledges support from the Danish Council for Independent Research--Natural Sciences (grant no. 10-83348).

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DOI 10.1007/s12229-011-9086-8

Manuel J. Macia (1,5) * Pedro J. Armesilla (1) * Rodrigo Camara-Leret (1) * Narel Paniagua-Zambrana (2) * Soraya Villalba (3) * Henrik Balslev (4) * Manuel Pardo-de-Santayana (1)

(1) Departamento de Biologia, Area de Botanica, Universidad Autonoma de Madrid, Calle Darwin 2, ES28049 Madrid, Spain

(2) Herbario Nacional de Bolivia, Universidad Mayor de San Andres, Campus Universitario, Cota Cota calle 27, Casilla 10077--Correo Central, La Paz, Bolivia

(3) Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, United Kingdom

(4) Department of Biological Sciences. Ecoinformatics & Biodiversity Research Group, Aarhus University, Build. 1540, Ny Munkegade 114, DK-8000 Aarhus C, Denmark

(5) Author for Correspondence; e-mail: manuel.macia@uam.es

Table 1 Description of use categories and subcategories in the present
review

Use category     Use subcategory            Description

Animal Food      Fish Bait                  Bait for fishing

                 Fodder                     Food for domestic animals

                 Wildlife Attractant        Palms that provide food
                                            for mammals and whose
                                            location constitutes
                                            preferential areas for
                                            hunting

Construction     Bridges                    Materials to bridge
                                            watercourses

                 Houses                     Houses and other
                                            constructions such as
                                            temporary camps, animal
                                            yards

                 Thatch                     House thatching and other
                                            constructions

                 Transportation             Canoes, rafts, oars and
                                            other materials for
                                            sealing

                 Other                      Uses not classifiable
                                            within the previous
                                            subcategories, for example
                                            stems used as posts for
                                            telephone lines and
                                            gutters to transport water

Cultural Uses    Clothes and Accessories    Articles of clothing and
                                            accessories such as hats

                 Cosmetic                   Beauty products, including
                                            perfumes, oils, shampoo,
                                            and other hair care
                                            products

                 Dyes                       Dyeing of diverse
                                            materials (vegetables) and
                                            as body paint

                 Personal Adornment         Necklaces, bracelets,
                                            earrings, armbands,
                                            pectorals, anklets

                 Recreational               Musical instruments, toys,
                                            ashes as additives to the
                                            consumption of tobacco and
                                            coca leaves

                 Ritual                     Uses related to
                                            myth-religious aspects,
                                            including festivals and
                                            feasts, construction of
                                            coffins, to drive away
                                            feared animals, sorcery

                 Other                      Uses not classifiable
                                            under the previous
                                            subcategories

Environmental    Agroforestry               Palms that are part of
Uses                                        agroforestry systems with
                 Fences                     different management
                                            degrees Delimitation of
                                            properties, barriers

                 Ornamental                 Palms cultivated for
                                            ornamental purposes

                 Soil Improvers             Fertilizers, edaphic
                                            protectors and agents
                                            against soil erosion

Fuel             Firewood                   Wood to make fire

                 Fire Starter               Combustion starters

                 Lighting                   Lamps, torches and candles

                 Other                      Uses not classifiable
                                            within the previous
                                            subcategories, for example
                                            for waterproofing canoes

Human Food       Beverages                  Elaboration of unfermented
                                            or fermented drinks

                 Food                       Edible, generally with
                                            little preparation

                 Food Additives             Ingredients used in the
                                            preparation and processing
                                            of foods

                 Oils                       Edible fats

Medicinal and    Blood and                  Anemia, cardiovascular
Veterinary       Cardio-vascular System     problems and ailments,
                                            cardiac diseases, varicose
                                            veins, hypertension,
                                            hypotension, haemorrhoids

                 Cultural Diseases and      Ailments or disorders of
                 Disorders                  magic-religious origin
                                            recognized by a specific
                                            culture, like mal aire

Use category     Use subcategory            Description (`bad air'),
                                            arrebato (`outburst'),
                 Dental Health              susto, huarana Caries,
                                            tooth pains, fillings,
                                            dental hygiene

                 Digestive System           Carminative, colics,
                                            flatulence, emetic,
                                            indigestion, purgative,
                                            gastric or intestinal
                                            ulcers, diarrhea,
                                            laxatives, liver and
                                            vesicular disorders,
                                            hepatitis

                 Endocrine System           Diabetes

                 General Ailments with      General ailments like body
                 Unspecific Symptoms        pains, general discomfort,
                                            weakness, headache, fever

                 Infections and             Malaria, leishmaniasis,
                 Infestations               measles, antihelminthic,
                                            louse, fleas, chiggers,
                                            scabies

                 Metabolic System and       Obesity, weight loss
                 Nutrition

                 Muscular-Skeletal          Rheumatism, twists,
                 System                     fractures, sciatic,
                                            lumbalgia

                 Nervous System and         Migraine, mental
                 Mental Health              disorders, epilepsy,
                                            paralysis, nervous
                                            disorders

                 Poisoning                  Snakebites, scorpion
                                            stings, rays, spiders,
                                            insects

                 Pregnancy, Birth and       Gestation, haemorrhage,
                 Puerperium                 childbirth, postnatal,
                                            lactation, abortive,
                                            postpartum

                 Reproductive System        Menstruation, fertility,
                 and Reproductive           venereal diseases,
                 Health                     prostrate, impotence,
                                            menopause, aphrodisiacs,
                                            contraceptives

                 Respiratory System         Flu, cold, loss of voice,
                                            bronchitis, pneumonia,
                                            expectorant, cough

                 Sensory System             Eye infections, cataracts,
                                            loss of sight or smell,
                                            deafness, ear infection

                 Skin and Subcutaneous      Acne, boils, eczemas,
                 Tissue                     bums, extraction of spines
                                            stuck on the skin

                 Urinary System             Diuretic, kidney stones,
                                            urinary incontinence,
                                            urinary infections,
                                            cystitis

                 Veterinary                 Treatment of diseases or
                                            ailments for domestic
                                            animals

                 Not Specified              Medicinal use or with
                                            pharmacological
                                            properties, but with
                                            insufficient information
                                            to assign to one of the
                                            described subcategories

                 Other                      Uses not classifiable
                                            within the previous
                                            subcategories, for example
                                            tumours, cancer,
                                            anaesthetic

Toxic            Fishing                    Fish poison

                 Hunting                    Poison for hunting

Utensils and     Domestic Utensils          Baskets, fans, hammocks,
Tools                                       bags, domestic furniture,
                                            air freshener

                 Hunting and Fishing        Bows, arrows, blowpipes,
                 Tools                      harpoons, fishing nets,
                                            hunting traps

                 Labour Tools               Agricultural or domestic
                                            tools like spinners,
                                            machetes and lubricants of
                                            these materials

                 Rope                       Manufacturing of ropes and
                                            moorings

                 Wrappers                   Wrappers for materials and
                                            foods

                 Other                      Uses not classifiable
                                            within the previous
                                            subcategories, for example
                                            insect repellents

Other Uses       Miscellaneous              Uses not classifiable
                                            within the previous
                                            categories. Indirect use
                                            of palms: insect larvae
                                            feeding on rotting stems
                                            used as food, medicine or
                                            bait

Table 2 Palm uses in the Amazon and Andes ecoregions of Colombia,
Ecuador, Peru and Bolivia, and in the Choco ecoregion of Colombia and
Ecuador

Country/Ecoregion    Useful     Palm uses    Palm use-reports
                     species

All countries        194        2,395        6,141
Amazon               134        1,972        5,144
Andes                68         344          439
Choco                52         347          569
Colombia             105        814          1,429
Amazon               70         615          1,049
Andes                18         35           39
Choco                38         225          341
Ecuador              103        936          2,010
Amazon               62         676          1,494
Andes                52         240          295
Choco                30         167          228
Peru                 96         785          1,390
Amazon               93         772          1,369
Andes                4          19           21
Bolivia              62         655          1,348
Amazon               54         603          1,267
Andes                13         77           84

Country/Ecoregion    Average [+ or -] SD   Percentage of useful
                     of palm uses per      species/Potential
                     species               total species

All countries        12.3 [+ or -] 18.7    63.2
Amazon               14.7 [+ or -] 20.0    89.9
Andes                5.1 [+ or -] 6.0      52.7
Choco                6.7 [+ or -] 7.3      49.0
Colombia             7.8 [+ or -] 10.1     47.5
Amazon               8.8 [+ or -] 10.6     67.6
Andes                1.9 [+ or -] 1.2      19.8
Choco                5.9 [+ or -] 5.7      43.2
Ecuador              9.1 [+ or -] 11.9     79.4
Amazon               10.9 [+ or -] 12.3    91.2
Andes                4.6 [+ or -] 5.3      82.5
Choco                5.6 [+ or -] 5.2      44.6
Peru                 8.2 [+ or -] 10.1     75.6
Amazon               8.3 [+ or -] 10.1     87.7
Andes                4.8 [+ or -] 3.4      9.1
Bolivia              10.6 [+ or -] 14.7    73.8
Amazon               11.2 [+ or -] 14.6    85.7
Andes                5.9 [+ or -] 6.7      43.3

Country/Ecoregion    Indigenous groups with        Bibliographical
                     ethnobotanical information    references
                     (Percentage of indigenous     (Palm monographs)
                     groups with info/Total
                     indigenous groups)

All countries        54 (49.1)                     255 (95)
Amazon               47 (47.5)                     202 (69)
Andes                2 (28.6)                      40 (27)
Choco                5 (83.3)                      38 (20)
Colombia             22 (48.9)                     63 (20)
Amazon               19 (48.7)                     41 (12)
Andes                --                            6 (5)
Choco                3 (75.0)                      25 (11)
Ecuador              10 (83.3)                     81 (31)
Amazon               7 (87.5)                      59 (17)
Andes                --                            20 (12)
Choco                3 (75.0)                      13 (10)
Peru                 18 (38.3)                     74 (28)
Amazon               18 (38.3)                     70 (26)
Andes                --                            4 (2)
Bolivia              11 (61.1)                     47 (22)
Amazon               10 (58.8)                     41 (17)
Andes                2 (100)                       11 (9)

Total number of species in each ecoregion and country was obtained
from Pintaud et al. (2008), and total number of indigenous groups from
Lewis (2009)

Table 3 Percentages of useful palm species by different use categories
and subcategories in tropical forests of north-western South America,
broken down by ecoregion and country. Total percentages of each
category (in bold) were calculated relative to the total usefid
species registered for each ecoregion and country. The percentages for
the different subcategories were calculated relative to the total
useful species registered in each of the categories by ecoregion and
country

Use category/Subcategory            Total    Ecoregion

                                             Amazon    Andes    Choco

Human Food                          69.6     76.9      57.4     61.5
Food                                95.6     95.1      97.4     93.8
Beverages                           41.5     44.7      25.6     34.4
Oils                                20.0     20.4      10.3     34.4
Food Additives                      8.9      10.7      5.1      6.3
Utensils and Tools                  65.5     73.9      32.4     57.7
Domestic                            77.2     81.8      86.4     53.3
Hunting and Fishing                 55.9     63.6      27.3     46.7
Labour Tools                        17.3     17.2      4.5      20.0
Wrappers                            13.4     16.2      4.5      --
Rope                                11.8     9.1       9.1      13.3
Other                               34.6     26.3      18.2     66.7
Construction                        63.4     70.1      48.5     55.8
Thatch                              83.7     86.2      87.9     82.8
Houses                              63.4     60.6      57.6     58.6
Transportation                      8.9      7.4       3.0      13.8
Bridges                             8.1      8.5       3.0      3.4
Other                               21.1     25.5      3.0      17.2
Cultural Uses                       55.7     59.0      42.6     34.6
Ritual                              49.1     40.5      62.1     61.1
Recreational                        45.4     57.0      13.8     33.3
Personal Adornment                  39.8     50.6      6.9      16.7
Cloth and Accessories               34.3     36.7      24.1     27.8
Cosmetic                            25.0     32.9      13.8     11.1
Dyes                                9.3      8.9       3.3      16.7
Other                               9.3      12.7      --       --
Animal Food                         36.6     42.5      20.6     19.2
Wildlife Attractant                 71.8     78.9      42.9     50.0
Fodder                              38.0     31.6      57.1     50.0
Fish Bait                           26.8     29.8      --       20.0
Medicinal and Veterinary            35.1     45.5      16.2     23.1
Digestive System                    55.9     54.1      54.5     33.3
Respiratory System                  38.2     39.3      18.2     16.7
General Ailments with               33.8     37.7      --       8.3
  Unspecified Symptoms
Infections and Infestations         30.9     32.8      18.2     --
Skin and Subcutaneous Tissue        26.5     27.9      27.3     8.3
Muscular-Skeletal System            22.1     23.0      9.1      8.3
Poisoning                           19.1     21.3      9.1      --
Reproductive System and             16.2     14.8      18.2     25.0
  Reproductive Health
Cultural Diseases and Disorders     14.7     13.1      18.2     25.0
Blood and Cardio-Vascular           11.8     9.8       --       16.7
  System
Urinary System                      8.8      8.2       27.3     8.3
Pregnancy, Birth and                8.8      9.8       18.2     --
  Puerperium
Dental Health                       7.4      8.2       --       --
Endocrine System                    7.4      8.2       --       --
Nervous System and Mental           7.4      6.6       9.1      --
  Health
Metabolic System and                4.4      4.9       --       --
  Nutrition
Sensory System                      4.4      1.6       9.1      8.3
Veterinary                          2.9      3.3       --       --
Not Specified                       35.3     36.1      36.4     25.0
Other                               5.9      6.6       --       --
Environmental Uses                  34.5     29.4      36.8     34.6
Ornamental                          62.7     57.5      68.0     55.6
Agroforestry                        47.8     57.5      40.0     33.3
Fences                              34.3     37.5      36.0     22.2
Soil Improvement                    4.5      7.5       --       --
Fuel                                22.2     24.6      17.6     9.6
Firewood                            72.1     84.8      58.3     40.0
Fire Starter                        23.3     21.2      16.7     20.0
Lighting                            18.6     9.1       25.0     40.0
Other                               9.3      12.1      --       --
Other uses                          22.7     29.1      8.8      17.3

Use category/Subcategory            Country

                                    Colombia    Ecuador    Peru

Human Food                          52.4        64.1       75.0
Food                                96.4        95.5       94.4
Beverages                           32.7        37.9       34.7
Oils                                27.3        13.6       12.5
Food Additives                      10.9        4.5        4.2
Utensils and Tools                  61.9        58.3       56.3
Domestic                            64.6        75.0       81.5
Hunting and Fishing                 63.1        65.0       48.1
Labour Tools                        9.2         20.0       13.0
Wrappers                            7.7         15.0       13.0
Rope                                13.8        10.0       9.3
Other                               43.1        30.0       16.7
Construction                        56.2        63.1       67.7
Thatch                              76.3        86.2       83.1
Houses                              44.1        47.7       66.2
Transportation                      5.1         7.7        6.2
Bridges                             1.7         4.6        6.2
Other                               32.2        4.6        9.2
Cultural Uses                       55.2        48.5       37.5
Ritual                              44.8        56.0       16.7
Recreational                        60.3        20.0       25.0
Personal Adornment                  32.8        50.0       25.0
Cloth and Accessories               22.4        24.0       36.1
Cosmetic                            10.3        26.0       33.3
Dyes                                3.4         10.0       8.3
Other                               5.2         8.0        13.9
Animal Food                         20.0        44.7       23.9
Wildlife Attractant                 76.2        91.3       30.4
Fodder                              19.0        21.7       26.1
Fish Bait                           28.6        10.9       52.2
Medicinal and Veterinary            27.6        31.1       34.4
Digestive System                    31.0        37.5       63.6
Respiratory System                  20.7        31.3       18.2
General Ailments with               6.9         18.8       42.4
  Unspecified Symptoms
Infections and Infestations         20.7        15.6       42.4
Skin and Subcutaneous Tissue        6.9         21.9       12.1
Muscular-Skeletal System            10.3        12.5       15.2
Poisoning                           24.1        3.1        9.1
Reproductive System and             6.9         9.4        21.2
  Reproductive Health
Cultural Diseases and Disorders     10.3        12.5       3.0
Blood and Cardio-Vascular           6.9         3.1        9.1
  System
Urinary System                      13.8        6.3        6.1
Pregnancy, Birth and                3.4         12.5       12.1
  Puerperium
Dental Health                       6.9         6.3        3.0
Endocrine System                    --          --         9.1
Nervous System and Mental           --          9.4        6.1
  Health
Metabolic System and                3.4         --         --
  Nutrition
Sensory System                      --          6.3        --
Veterinary                          --          --         3.0
Not Specified                       27.6        43.8       24.2
Other                               --          6.3        6.1
Environmental Uses                  25.7        32.0       25.0
Ornamental                          48.1        66.7       37.5
Agroforestry                        48.1        39.4       70.8
Fences                              18.5        27.3       41.7
Soil Improvement                    --          --         8.3
Fuel                                16.2        27.2       7.3
Firewood                            88.2        75.0       42.9
Fire Starter                        11.8        21.4       28.6
Lighting                            --          21.4       --
Other                               --          10.7       28.6
Other uses                          21.9        14.6       28.1

Use category/Subcategory            Country

                                    Bolivia

Human Food                          67.7
Food                                90.5
Beverages                           40.5
Oils                                33.3
Food Additives                      14.3
Utensils and Tools                  53.2
Domestic                            90.9
Hunting and Fishing                 39.4
Labour Tools                        9.1
Wrappers                            9.1
Rope                                12.1
Other                               24.2
Construction                        56.5
Thatch                              80.0
Houses                              62.9
Transportation                      2.9
Bridges                             5.7
Other                               20.0
Cultural Uses                       50.0
Ritual                              51.6
Recreational                        41.9
Personal Adornment                  32.3
Cloth and Accessories               48.4
Cosmetic                            48.4
Dyes                                6.5
Other                               12.9
Animal Food                         35.5
Wildlife Attractant                 63.6
Fodder                              50.0
Fish Bait                           9.1
Medicinal and Veterinary            33.9
Digestive System                    42.9
Respiratory System                  47.6
General Ailments with               47.6
  Unspecified Symptoms
Infections and Infestations         9.5
Skin and Subcutaneous Tissue        38.1
Muscular-Skeletal System            33.3
Poisoning                           23.8
Reproductive System and             19.0
  Reproductive Health
Cultural Diseases and Disorders     19.0
Blood and Cardio-Vascular           14.3
  System
Urinary System                      14.3
Pregnancy, Birth and                9.5
  Puerperium
Dental Health                       9.5
Endocrine System                    9.5
Nervous System and Mental           --
  Health
Metabolic System and                9.5
  Nutrition
Sensory System                      4.8
Veterinary                          4.8
Not Specified                       33.3
Other                               --
Environmental Uses                  37.1
Ornamental                          52.2
Agroforestry                        52.2
Fences                              43.5
Soil Improvement                    4.3
Fuel                                14.5
Firewood                            88.9
Fire Starter                        11.1
Lighting                            22.2
Other                               --
Other uses                          14.5

Table 4 Percentages of use-reports for the different palm parts used
in each category in tropical forests of north-western South America

Plant part        Human Food    Utensils and Tools    Construction

Fruit             60.6          2.4                   --
Stem              0.8           30.0                  36.1
Entire leaf       0.1           20.2                  53.4
Seed              11.6          4.4                   --
Palm heart        19.5          0.5                   --
Root              0.2           3.2                   --
Entire plant      --            --                    --
Spear leaf        --            10.6                  0.1
Petiole           --            5.1                   0.5
Leaf rachis       --            4.8                   0.2
Leaf sheath       --            3.1                   --
Bract             --            1.7                   --
Inflorescence     0.3           0.7                   --
Flower            0.6           --                    --
Spine             --            0.4                   --
Infructescence    <0.1          0.2                   --
Not specified     6.3           12.6                  9.7

Plant part        Cultural    Animal Food    Medicinal and Veterinary

Fruit             13.7        76.5           23.8
Stem              7.7         0.9            4.0
Entire leaf       17.6        4.3            4.5
Seed              17.4        5.2            11.6
Palm heart        3.1         2.2            9.4
Root              2.5         --             31.3
Entire plant      7.4         --             0.4
Spear leaf        8.3         --             --
Petiole           1.3         --             0.2
Leaf rachis       1.3         --             0.2
Leaf sheath       0.5         --             0.2
Bract             3.5         --             --
Inflorescence     2.9         0.9            1.0
Flower            0.6         0.9            2.4
Spine             1.3         --             1.4
Infructescence    0.3         --             0.2
Not specified     10.5        9.1            9.4

Plant part        Environmental    Fuel    Other Uses    Total

Fruit             1.1              7.8     6.0           25.1
Stem              20.7             52.3    64.7          18.1
Entire leaf       5.2              21.9    1.3           16.7
Seed              2.9              7.8     8.2           7.9
Palm heart        0.4              --      --            7.1
Root              -                --      0.4           3.9
Entire plant      65.3             --      --            3.6
Spear leaf        --               0.8     4.7           3.1
Petiole           --               0.8     0.9           1.2
Leaf rachis       --               --      --            1.1
Leaf sheath       --               3.1     0.9           0.8
Bract             --               --      --            0.7
Inflorescence     0.4              --      --            0.7
Flower            0.4              --      --            0.5
Spine             --               --      0.4           0.4
Infructescence    0.4              --      --            0.1
Not specified     3.3              5.5     12.5          9.0

Table 5 Use of palms by different human groups in tropical forests of
north-western South America. For  some ecoregions and countries no
data was available

Human group/    Ecoregion          Useful     Palm     Palm use-
Country                            species    uses     reports

Indigenous      Total              129        1,555    3,713

Colombia        All ecoregions     74         574      926
                Amazon             59         513      823
                Andes              1          1        1
                Choco              26         87       102
Ecuador         All ecoregions     78         770      1,704
                Amazon             59         656      1,448
                Andes              23         95       109
                Choco              24         110      149
Peru            All ecoregions/    47         278      402
                  Amazon
Bolivia         All ecoregions     33         397      716
                Amazon             32         385      694
                Andes              3          25       25

Mestizo         Total              49         215      304

Ecuador         All ecoregions     15         28       30
                Amazon             5          6        5
                Andes              11         24       25
Peru            All ecoregions     35         163      239
                Amazon             34         155      226
                Andes              2          11       13
Bolivia         All ecoregions     10         35       35
                Amazon             9          29       29
                Andes              2          6        6

Afroamerican    Total              24         82       90

Colombia        All ecoregions/    23         77       84
                  Choco
Ecuador         All ecoregions/    3          6        6
                  Choco

Colono          Total              15         56       61

Colombia        All ecoregions/    10         33       33
                  Amazon
Ecuador         All ecoregions     8          14       14
                Amazon             6          9        9
                Andes              I          I        I
                Choco              1          4        4
Peru            All ecoregions/    4          9        9
                  Amazon
Bolivia         All ecoregions/    2          5        5
                  Amazon

Not identified  Total              170        1,166    2,012

Colombia        All ecoregions     82         293      393
                Amazon             51         173      199
                Andes              18         35       39
                Choco              29         113      156
Ecuador         All ecoregions     61         242      273
                Amazon             19         36       43
                Andes              38         155      160
                Choco              16         63       74
Peru            All ecoregions     87         531      750
                Amazon             85         523      742
                Andes              2          8        8
Bolivia         All ecoregions     56         382      596
                Amazon             48         340      543
                Andes              10         47       53

Human group/    Ecoregion          Uses [+ or -] SD      References
Country                            per species

Indigenous      Total              12.1 [+ or -] 16.7    166

Colombia        All ecoregions     7.8 [+ or -] 9.8      48
                Amazon             8.7 [+ or -] 10.3     37
                Andes              1.0 [+ or -] 0.0      1
                Choco              3.3 [+ or -] 2.9      13
Ecuador         All ecoregions     9.9 [+ or -] 11.9     67
                Amazon             11.1 [+ or -] 12.2    57
                Andes              4.1 [+ or -] 3.0      10
                Choco              4.6 [+ or -] 3.8      8
Peru            All ecoregions/    5.9 [+ or -] 5.8      29
                  Amazon
Bolivia         All ecoregions     12.0 [+ or -] 13.3    30
                Amazon             12.0 [+ or -] 12.2    28
                Andes              8.3 [+ or -] 11.0     3

Mestizo         Total              4.4 [+ or -] 4.5      30

Ecuador         All ecoregions     1.9 [+ or -] 1.8      4
                Amazon             1.0 [+ or -] 0.0      2
                Andes              2.2 [+ or -] 2.0      3
Peru            All ecoregions     4.7 [+ or -] 4.5      22
                Amazon             4.6 [+ or -] 4.1      20
                Andes              5.5 [+ or -] 4.9      2
Bolivia         All ecoregions     3.5 [+ or -] 2.1      4
                Amazon             3.2 [+ or -] 2.1      3
                Andes              3.0 [+ or -] 1.4      1

Afroamerican    Total              3.4 [+ or -] 2.7      7

Colombia        All ecoregions/    3.3 [+ or -] 2.4      5
                  Choco
Ecuador         All ecoregions/    2.0 [+ or -] 1.7      2
                  Choco

Colono          Total              3.7 [+ or -] 2.7      12

Colombia        All ecoregions/    3.3 [+ or -] 1.6      3
                  Amazon
Ecuador         All ecoregions     1.8 [+ or -] 1.0      7
                Amazon             1.5 [+ or -] 0.5      5
                Andes              1.0 [+ or -] 0.0      1
                Choco              4.0 [+ or -] 0.0      1
Peru            All ecoregions/    2.3 [+ or -] 0.5      1
                  Amazon
Bolivia         All ecoregions/    2.5 [+ or -] 0.7      1
                  Amazon

Not identified  Total              6.9 [+ or -] 10.1     86

Colombia        All ecoregions     3.6 [+ or -] 3.9      21
                Amazon             3.4 [+ or -] 3.3      11
                Andes              1.9 [+ or -] 1.2      6
                Choco              3.9 [+ or -] 3.1      12
Ecuador         All ecoregions     4.0 [+ or -] 5.0      18
                Amazon             1.9 [+ or -] 2.7      6
                Andes              4.1 [+ or -] 4.6      9
                Choco              3.9 [+ or -] 5.0      9
Peru            All ecoregions     6.1 [+ or -] 6.9      30
                Amazon             6.2 [+ or -] 7.0      28
                Andes              4.0 [+ or -] 2.8      2
Bolivia         All ecoregions     6.8 [+ or -] 8.0      18
                Amazon             7.1  [+ or -] 8.2     15
                Andes              4.7 [+ or -] 5.3      7

Table 6 Use of palms by the different indigenous groups living in the
tropical forests of north-western  South America

Indigenous        Country             Useful     Palm    Palm use-
group                                 species    uses    reports

Amazon
  Quichua (also   Ecuador             44         243     387
    in Andes)
  Huaorani        Ecuador             43         337     500
  Shuar(also      Ecuador             40         186     305
    in Andes)
  Mumane          Colombia            36         183     183
  Secoya          Ecuador/Peru        29         93      105
  Cofdn           Ecuador             26         99      127
  Cocama          Peru                25         59      59
  Tacana          Bolivia             23         205     262
  Siona           Colombia/Ecuador    22         39      40
  Shipibo-        Peru                20         55      63
    Combo
  Tikuna          Colombia            19         77      80
  Tsimane/        Bolivia             18         108     121
    Mosetene
  Cubeo           Colombia            18         45      46
  Achuar          Ecuador/Peru        18         41      49
  Huitoto         Colombia/Peru       17         62      62
  Chayahuita      Peru                17         26      26
  Mirana          Colombia            16         63      67
  Matsigenka      Peru                16         31      31
  Nukak           Colombia            15         120     168
  Bora            Colombia/Peru       14         58      66
  Matse           Peru                14         36      36
  Chacobo         Bolivia             13         36      37
  Quechua/        Bolivia             12         73      73
    Tacana
  Aguaruna        Peru                II         36      40
  Yucarare/       Bolivia             10         69      69
    Trinitario
  Andoque         Colombia            10         28      28
  Yucuna          Colombia            10         20      20
  Puinave         Colombia            9          18      18
  Yagua           Peru                9          15      15
  Ese Eja         Bolivia/Peru        9          14      14
  Tsimane         Bolivia             8          39      43
  Mosetene        Bolivia             7          26      29
  Siona-Secoya    Colombia/Ecuador    6          11      11
  Bora-Ocaina-    Peru                5          21      21
    Huitoto
  Siriono         Bolivia             5          16      26
  Orejon          Peru                5          7       7
  Guayabero       Colombia            3          18      18
  Curripaco       Colombia            3          8       8
  Yuracare        Bolivia             3          8       9
  Ocaina          Peru                3          5       5
  Omagua          Peru                3          4       4
  Yanesha         Peru                2          8       8
  Ashaninka       Peru                2          3       3
  Desano          Colombia            2          2       2
  Makuna          Colombia            2          2       2
  Piapoco         Colombia            1          2       2
  Trinitario      Bolivia             l          2       2
  Tukano          Colombia            1          2       2
  Cashibo         Peru                I          1       I
  Coreguaje       Colombia            1          1       1
  Sikuani         Colombia            I          I       1
  Araona          Bolivia             1          1       1
Andes
  Leco            Bolivia             2          24      24
  Quechua         Bolivia             l          I       1
Choco
  Awa             Colombia/Ecuador    18         74      84
  Chachi          Ecuador             15         70      87
  Tsachila        Ecuador             13         27      28
  Embera          Colombia            6          9       9
  Waunan          Colombia            1          8       8

Indigenous        Uses [+ or -] SD      References
group             per species

Amazon
  Quichua (also   5.5 [+ or -] 6.8      23
    in Andes)
  Huaorani        7.8 [+ or -] 6.8      13
  Shuar(also      4.7 [+ or -] 4.7      15
    in Andes)
  Mumane          5.1 [+ or -] 3.8      2
  Secoya          3.2 [+ or -] 2.1      6
  Cofdn           3.8 [+ or -] 2.8      5
  Cocama          2.4 [+ or -] 1.4      2
  Tacana          8.9 [+ or -] 8.1      5
  Siona           1.8 [+ or -] 1.9      4
  Shipibo-        2.8 [+ or -] 1.5      4
    Combo
  Tikuna          4.1 [+ or -] 2.7      5
  Tsimane/        6.0 [+ or -] 2.9      3
    Mosetene
  Cubeo           2.5 [+ or -] 2.8      4
  Achuar          2.3 [+ or -] 1.4      7
  Huitoto         3.6 [+ or -] 3.6      9
  Chayahuita      1.5 [+ or -] 0.9      l
  Mirana          3.9 [+ or -] 2.9      4
  Matsigenka      1.9 [+ or -] 1.0      1
  Nukak           8.0 [+ or -] 5.2      3
  Bora            4.1 [+ or -] 2.9      10
  Matse           2.6 [+ or -] 1.5      1
  Chacobo         2.8 [+ or -] 1.5      2
  Quechua/        6.1 [+ or -] 3.3      1
    Tacana
  Aguaruna        3.3 [+ or -] 2.0      2
  Yucarare/       6.9 [+ or -] 3.4      2
    Trinitario
  Andoque         2.8 [+ or -] 1.3      2
  Yucuna          2.0 [+ or -] 1.1      4
  Puinave         2.0 [+ or -] 1.4      4
  Yagua           1.7 [+ or -] 0.9      3
  Ese Eja         1.6 [+ or -] 0.7      2
  Tsimane         4.9 [+ or -] 3.9      5
  Mosetene        3.7 [+ or -] 2.3      4
  Siona-Secoya    1.8 [+ or -] 0.8      2
  Bora-Ocaina-    4.2 [+ or -] 2.0      1
    Huitoto
  Siriono         3.2 [+ or -] 4.9      3
  Orejon          1.4 [+ or -] 0.9      2
  Guayabero       6.0 [+ or -] 1.7      1
  Curripaco       2.7 [+ or -] 2.9      2
  Yuracare        2.7 [+ or -] 2.1      3
  Ocaina          1.7 [+ or -] 1.2      2
  Omagua          1.3 [+ or -] 0.6      1
  Yanesha         4.0 [+ or -] 2.8      2
  Ashaninka       1.5 [+ or -] 0.7      1
  Desano          1.0 [+ or -] 0.0      1
  Makuna          1.0 [+ or -] 0.0      I
  Piapoco         2.0 [+ or -] 0.0      1
  Trinitario      2.0 [+ or -] 0.0      1
  Tukano          2.0 [+ or -] 0.0      1
  Cashibo         1.0 [+ or -] 0.0      I
  Coreguaje       1.0 [+ or -] 0.0      I
  Sikuani         1.0 [+ or -] 0.0      1
  Araona          1.0 [+ or -] 0.0      1
Andes
  Leco            12.0 [+ or -] 12.7    2
  Quechua         1.0 [+ or -] 0.0      1
Choco
  Awa             4.1 [+ or -] 3.4      8
  Chachi          4.9 [+ or -] 3.5      6
  Tsachila        2.1 [+ or -] 1.1      3
  Embera          1.5 [+ or -] 1.2      2
  Waunan          8.0 [+ or -] 0.0      1

Table 7 Useful palms with high relative importance value index in
different ecoregions of tropical forests  of north-western South
America

Species per ecoregion             Relative      Palm    Palm use-
                                  Importance    uses    reports
Amazon
  Bactris gasipaes                2.0           76      414
  Euterpe precatoria              2.0           89      358
  Oenocaipus bataua               2.0           107     544
  Attalea phalerata               1.9           78      227
  Mauritia flexuosa               1.9           95      381
  Attalea maripa                  1.7           61      136
  Iriartea deltoidea              1.7           70      283
  Oenocarpus mapora               1.7           50      175
  Socratea exorrhiza              1.7           63      236
  Astrocaryum chambira            1.6           60      255
  Astrocaryum murumuru            1.6           53      103
  Attalea butyracea               1.6           37      85
  Astrocaryum aculeatum           1.5           39      65
  Phytelephas macrocarpa          1.5           35      118
  Phytelephas tenuicaulis         1.4           30      72
Andes
  Bactris gasipaes                2.0           34      45
  Oenocarpus bataua               1.6           22      38
  Iriartea deltoidea              1.4           18      24
  Attalea phalerata               1.3           21      21
  Wettinia maynensis              1.3           17      26
  Socratea exorrhiza              1.1           10      15
  Ceroxylon echinulatum           1.0           8       14
  Oenocarpus mapora               1.0           10      12
  Ceroxylon ventricosum           0.9           7       7
  Parajubaea sunkha               0.9           16      16
  Phytelephas aequatorialis       0.9           8       8
  Prestoea ensiformis             0.9           9       9
  Dictyocaryum lamarckianum       0.8           9       11
  Parajubaea torallyi             0.8           12      13
Choco
  Cocos nucifera                  2.0           30      48
  Bactris gasipaes                1.6           24      46
  Wettinia quinaria               1.6           20      32
  Astrocaryum standleyanum        1.4           27      54
  Iriartea deltoidea              1.4           17      32
  Euterpe oleracea                1.3           17      40
  Oenocarpus bataua               1.2           18      36
  Phrtelephas aequatorialis       1.2           14      20
  Oenocarpus mapora               1.1           14      20
  Attalea colenda                 1.0           12      21
  Socratea exorrhiza              1.0           14      21
  Manicaria saccifera             0.9           10      20
  Attalea cuatrecasana            0.8           10      13
  Geonoma cuneata                 0.8           8       10
  Phytelephas seemannii           0.8           7       13
  Synechanthus warscewiczianus    0.8           7       8

Species per ecoregion             Countries     References

Amazon
  Bactris gasipaes                C, E, P, B    109
  Euterpe precatoria              C, E, P, B    91
  Oenocaipus bataua               C, E, P, B    117
  Attalea phalerata               P, B          28
  Mauritia flexuosa               C, E, P, B    101
  Attalea maripa                  C, E, P, B    31
  Iriartea deltoidea              C, E, P, B    79
  Oenocarpus mapora               C, E, P, B    51
  Socratea exorrhiza              C, E, P, B    69
  Astrocaryum chambira            C, E, P       68
  Astrocaryum murumuru            C, E, P, B    24
  Attalea butyracea               C, E, P, B    26
  Astrocaryum aculeatum           C, P, B       17
  Phytelephas macrocarpa          C, E, P, B    47
  Phytelephas tenuicaulis         C, E, P       18
Andes
  Bactris gasipaes                C, E, P, B    8
  Oenocarpus bataua               E, S          6
  Iriartea deltoidea              C, E          4
  Attalea phalerata               B             I
  Wettinia maynensis              E             5
  Socratea exorrhiza              E             3
  Ceroxylon echinulatum           C, E, P       4
  Oenocarpus mapora               E             2
  Ceroxylon ventricosum           E             1
  Parajubaea sunkha               B             3
  Phytelephas aequatorialis       E             2
  Prestoea ensiformis             E             1
  Dictyocaryum lamarckianum       C, E, B       6
  Parajubaea torallyi             B             3
Choco
  Cocos nucifera                  C, E          11
  Bactris gasipaes                C, E          13
  Wettinia quinaria               C, E          13
  Astrocaryum standleyanum        C, E          18
  Iriartea deltoidea              C, E          14
  Euterpe oleracea                C, E          12
  Oenocarpus bataua               C, E          15
  Phrtelephas aequatorialis       E             5
  Oenocarpus mapora               C, E          8
  Attalea colenda                 C, E          8
  Socratea exorrhiza              C, E          10
  Manicaria saccifera             C             9
  Attalea cuatrecasana            C             5
  Geonoma cuneata                 C, E          5
  Phytelephas seemannii           C             7
  Synechanthus warscewiczianus    E             5

Country abbreviations

C Colombia, E Ecuador, P Peru, B Bolivia

Article Details

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Title Annotation:	Part 1
Author:	Macia, Manuel J.; Armesilla, Pedro J.; Camara-Leret, Rodrigo; Paniagua-Zambrana, Narel; Villalba, So
Publication:	The Botanical Review
Article Type:	Report
Geographic Code:	4EUDE
Date:	Dec 1, 2011
Words:	18817
Previous Article:	Disturbance and resilience in tropical American palm populations and communities.
Next Article:	Appendix.
Topics:	Ecosystems Indigenous peoples Palms Environmental aspects Physiological aspects Palms (Plants) Environmental aspects Physiological aspects Racially mixed people Rain forests Environmental aspects Valuation