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Drawing on traditional knowledge to identify and describe ecosystem services associated with Northern Amazon’s multiple-use plants

Drawing on traditional knowledge to identify and describe ecosystem services associated with... INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT, 2016 VOL. 12, NOS. 1–2, 39–56 http://dx.doi.org/10.1080/21513732.2015.1136841 Special Issue: Traditional knowledge for sustainable forest management and provision of ecosystem services Drawing on traditional knowledge to identify and describe ecosystem services associated with Northern Amazon’s multiple-use plants a b Anthony R. Cummings and Jane M. Read Geospatial Information Sciences, School of Economic, Political and Policy Sciences, University of Texas at Dallas, Richardson, TX, USA; Department of Geography, Maxwell School of Citizenship and Public Affairs, Syracuse University, Syracuse, NY, USA ABSTRACT ARTICLE HISTORY Received 30 December 2014 Tropical multiple-use plants provide an array of ecosystem services, including providing food Accepted 23 December 2015 for wildlife, non-timber forest products (NTFPs) and commercial logs. As land-use and land- cover change (LUCC) continues in tropical forests, questions on whether the ecosystem EDITED BY services they provide can be sustained become more pressing. As particular species of plants John Parrotta are targeted for provisioning services, such as commercial logging, the implications for other KEYWORDS ecosystem services are often unknown. In this paper we drew on the traditional knowledge of Traditional knowledge; the Makushi and Wapishiana Amerindians of Southern Guyana, and multiple-use plants to multiple-use plants; gauge the ecosystem services that may be compromised as tropical forests undergo change. ecosystem services; wildlife An inventory of tree and palm species was classified into one or more of four resource-use food; Rupununi; Guyana classes: commercial timber, wildlife food, traditional uses or no known use. Species that intersected more than one resource use were defined as multiple-use species, and occupied four primary classes – wildlife food/commercial timber, commercial timber/traditional uses, wildlife food/traditional uses and wildlife food/commercial timber/traditional uses. Traditional knowledge allowed us to identify plant species and describe how they are used for provision- ing, cultural and supporting ecosystem services and shaping our understanding of the multiple dimensions of ecosystem services associated with a single species. Introduction Albert et al. 2014), examples from tropical forests are still lagging in the literature. The subject of identifying and mapping ecosystem The utilization of traditional knowledge (Posey services across the globe has gained substantial atten- 1992; Parrotta & Trosper 2012) to understand the tion recently. Scholars have established the defini- various types of ecosystem services associated with tions, conceptual framework and problems with tropical plant species and how these may be impacted understanding ecosystem services (e.g. Costanza by land-use and land-cover change (LUCC) and glo- et al. 1997; Daily 1997; Kareiva et al. 2011; bal climatic variability (Parrotta & Agnoletti 2007)is Cardinale et al. 2012; Albert et al. 2014), with various an area requiring pressing attention. The attention to systems emerging for their classification (see traditional knowledge and linkages to ecosystem ser- Millennium Ecosystem Assessment (MEA) 2005; vices is particularly critical as globalization and other Wu 2013). Ecosystem services and ecosystem func- forces reach previously self-sufficient rural areas and tions have been broadly described, while sophisti- result in a loss of traditional knowledge (Parrotta cated methods, models and data have been used to et al. 2009; Trosper et al. 2012). The urgency for better understand select ecosystem services (e.g. understanding the traditional knowledge-ecosystem Burkhard et al. 2012). Despite this progress, however, services connection is further amplified as the conse- the interactions between ecosystem services, as they quence of such loss on livelihoods and cultural and may originate from a single supplier (Furst et al. biological diversity remains poorly understood 2014) such as a tropical plant species, or how such (Parrotta et al. 2009). Furthermore, as Parrotta et al. ecosystem services relate directly to the well-being of (2009) noted, the general public lacks an appreciation human populations, remain under-explored. As the of traditional knowledge, and as a consequence its literature on ecosystem services continues to evolve, erosion may curtail the involvement of local and it seems necessary to understand localized examples indigenous communities in the development of forest of ecosystem services and how they relate to human and land-use policies that impact their livelihoods. As populations critical to the sustainability of such ser- such, scholars have called (e.g. Parrotta et al. 2009; vices. While past studies have covered examples of Sutherland et al. 2014) for greater recognition of the mapping ecosystem services in urban (e.g. Kareiva role indigenous and local people play in the et al. 2011) and developed world settings (e.g. CONTACT Anthony R. Cummings anthony.cummings@utdallas.edu © 2016 Informa UK Limited, trading as Taylor & Francis Group 40 A. R. CUMMINGS AND J. M. READ provisioning services, such as the use of plant parts management of biodiversity (see Mauro & Hardison for medicines (e.g. Forte 1996; van Andel 2000; Khan 2000; and the volume edited by Parrotta & Agnoletti 2007) and merging traditional knowledge with con- et al. 2013) and commercial logging (e.g. Polak 1992), have gained considerable attention. In a similar vein, ventional scientific knowledge (Sutherland et al. cultural services associated with a single species have 2014). In response to such calls, this paper explores the uses associated with plants from the forests of also been examined (see Forte 1996; González-Marín et al. 2012; Cámara-Leret et al. 2014). Of particular Guyana through the lens of forest-related traditional interest to this paper is the fact that individual species knowledge, and the guidance it can provide for forest management and landscape valuation efforts (e.g. of tropical plants are often associated with more than Castro et al. 2014; Paletto et al. 2015; Stara et al. one ecosystem services (similar to the notion of 2015) that have been gaining momentum in the Albert et al. 2014 that there may be multiple demands literature. from a single source for ecosystem services) and our A prominent feature of tropical forests is the host knowledge of such species and their uses is not well of multiple-use plant species they contain that have developed. direct associations with the four dimensions of eco- As tropical forests continue to be converted to system services (provisioning, regulating, supporting support human activities, concerns have arisen over and cultural services) identified by MEA (2005). At whether the ecosystem services they provide can be their most critical level, multiple-use plants provide sustained (e.g. Nepstad et al. 1999;Fearnside 2005). food for wildlife, non-timber forest products (NTFPs) Much of the recent assessments on the impacts of for forest dwelling and dependent peoples, and com- tropical forest area decline on ecosystem services mercial timber (Cummings 2013). The plant uses for have been on implications for carbon sequestration provisioning and supporting ecosystem services may and the hydrological cycle. Although these global- also be superimposed upon the cultural and regulat- level ecosystem services are important, the impacts ing services within a single species. The provisioning on forest-dwelling and dependent peoples, either ecosystem services associated with multiple-use directly through the removal of plants that provide plants, in particular the various demands from differ- important NTFPs or indirectly through impacts on ent groups of forest users on a particular species, have wildlife that they hunt, require equally urgent atten- been gaining attention in the literature (see for exam- tion. The direct impacts on wildlife are critical too, ple Klimas et al. 2007, 2012a, 2012b; Mwavu & as many species disperse seeds and fruits of forest Witkowski 2009). But this attention has often focused plants (Roosmalen 1985), thereby helping maintain on single well-known species (‘trophy species’), such the forest structure. Therefore, identifying and as Carapa guianensis – a species well-known for its studying multiple-use plants and how they are valuable timber and therapeutic oil (Klimas et al. impacted by LUCC is critical. In this regard, the 2007). Moreover, the efforts to probe and describe four parameters to be understood when managing the ecosystem services associated with multiple-use multiple-use plants – an understanding of their plants have made little mention of local-level man- multiplicity of uses and types of resources pro- agers, such as indigenous peoples, in the process of duced, abundance in different forest types, size- studying such species. Therefore, while previous stu- class distribution of populations and life history dies have provided valuable data on the ecology and characteristics (Peters 1996) – are critical in guiding distribution characteristics of multiple-use species our work. We pay particular attention to the first and how they fare under conditions of commercial two parameters suggested by Peters using a sample logging and NTFPs extraction, they introduce ques- of plants from Northern Amazonia. We argue that tions of how many more species exist, and the best there are many tropical multiple-use plant species, means for obtaining knowledge on the ecosystem and that when the knowledge of scientific experts is services they provide. combined with traditional knowledge a more com- In addition to only the prominent multiple-use prehensive appreciation of ecosystem services with species gaining attention in the literature, the diverse which they are associated can be derived. Such range of ecosystem services associated with a species knowledge can then be used for planning the sus- has also often been ignored. Single facet of plant uses, tainability of ecosystem services associated with for example their role in providing wildlife food, individual species and the landscapes in which NTFPs or commercial timber, separately, has been they are found (see Parrotta et al. 2009). Scholars commonly reported (see for example Peters 1996; such as Forte (1996), Parrotta and Trosper (2012) Klimas et al. 2007; Mwavu & Witkowski 2009). and Posey (1992) have drawn on traditional knowl- Scholars have examined the supporting ecosystem edge to various resource management-related ends. services associated with tropical plants (trees and Forte (1996), for example, utilized the knowledge of palms) through their fruits and seeds (e.g. Makushi women of the Rupununi, Southern Roosmalen 1985; Silvius & Fragoso 2003), while the Guyana, to describe how plants have been used for INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 41 processes that occur at temporal or spatial scales not medicinal purposes. Parrotta and Trosper (2012) easily observed by conventional scientific research. have highlighted the important knowledge that tra- ditional societies – indigenous and local commu- Peloquin and Berkes (2009) pointed out that indigen- ous and rural groups appeared to have developed nities – have used to manage their forests in order practices that allow them to live with complexity to sustain their livelihoods and cultures. The rapid changes in forested regions endanger not only indi- that are often missed by Western science-based socie- ties who seek to simplify ecosystems. genous and local communities that live within such As a consequence of the recognition of the value forests, but also the important ecosystem services that these forests may hold even before we are fully traditional knowledge has for humanity, scholars (e.g. Ens et al. 2015; Whyte et al. 2015) have been suggest- aware of them. Traditional knowledge of how indi- ing measures to incorporate traditional knowledge genous communities have used forests is therefore a critical step in understanding the impacts of LUCC into sustainability science. In recognition of the role within the context of sustainable forest management traditional knowledge must play in ecosystem con- servation and the protection of local livelihoods, this and preserving traditional lifestyles. paper seeks to bring the importance of traditional knowledge in helping shape our understanding of The value of traditional knowledge forests and their ecosystem services in landscapes that are vulnerable to change. In this regard our The value of traditional knowledge to ecosystem con- work will help inform modern-day approaches to servation, sustainable resource management and local sustainable forest management on how traditional livelihoods has been stressed by a number of scholars knowledge should be incorporated to protect the over the years (e.g Berkes et al. 2000; Mauro & livelihoods of local and forest-dependent peoples. Hardison 2000; Turner & Cocksedge 2001; LaRochelle & Berkes 2003; Houde 2007; Parrotta & Trosper 2012; de Freitas et al. 2015). Traditional Goals and objectives knowledge, which consists of the past and present beliefs, traditions, practices and views developed by With the foregoing discussion in mind, the goals of indigenous and local communities (Huntington this paper are twofold. First, drawing on the tradi- 2000), is a key to accessing, using and managing tional knowledge of the Makushi and Wapishiana ecosystem services (Cámara-Leret et al. 2014). Amerindians, we identified multiple-use plants spe- Cámara-Leret et al. (2014) further noted that tradi- cies of the Rupununi, Southern Guyana, landscape tional knowledge is diverse, dynamic and place-spe- (Figure 1). Secondly, we determined which uses, or cific, while Byg and Balslev (2004) noted that types of ecosystem services, are associated with each traditional knowledge can vary among individuals species. As a clear departure from previous studies, and communities. Phillips et al. (1994) posited that our analysis combined the literature on how plants the benefits local communities obtain from their sur- are used in Guyana with traditional knowledge of the rounding ecosystems are determined by the levels of Makushi and Wapishiana Amerindians to identify traditional knowledge about the classification, ecol- multiple-use plants sampled from the Rupununi. ogy, usefulness and management of natural resources. Our main objective is to demonstrate to policymakers Turner et al. (2000) noted that traditional knowledge the role traditional knowledge can play in sustainable is ‘fundamentally important in the management of forest management, with particular emphasis on con- local resources, in the husbanding of the world’s serving ecosystem services and protecting local liveli- biodiversity, and in providing locally valid models hoods. We focused primarily on species of plants that for sustainable living.’ Interest in traditional knowl- intersect the three main resource uses: commercial edge has been growing primarily as scholars recog- logging, traditional uses (NTFPs) and wildlife food, as nize the contribution it can make to the conservation the intersection of these uses in tropical forests will of biodiversity (Gadgil et al. 1993; Berkes et al. 2000), determine which species and the ecosystem services rare species (Colding 1998), protected areas with which they are associated will be vulnerable. (Johannes 1998) and to sustainable resource use in Where a single species had more than one use we general (Berkes 1999). Berkes et al. (2000) further described it as multiple-use (see Cummings 2013). noted that conservation biologists, ecological anthro- We used multiple-use classes to explore how plants pologists, ethnobiologists, other scholars and the are used and how these uses may be compromised by pharmaceutical industry, all share an interest in tra- changing landscapes. In this regard, this paper goes ditional knowledge for scientific, social or economic beyond previous efforts that provided a cursory view reasons. More recently, de Freitas et al. (2015) noted of the important resource-use intersections associated that traditional knowledge can provide insights into with an individual plant species (e.g. Klimas et al. studying ecological systems that are important to 2007, 2012a, 2012b), and offers a comprehensive local populations, and understanding ecological description of the ecosystem services associated with 42 A. R. CUMMINGS AND J. M. READ Figure 1. Study area. each species. Our work therefore contributes to the of traditional knowledge in sustainable forest man- ongoing discussion on valuing landscapes for the agement and valuation processes of tropical land- ecosystem services they provide. scapes and the impacts of deforestation beyond the We make the argument that through identifying current carbon sequestration focus. multiple-use plants within tropical forests, and utiliz- ing traditional knowledge to identify such plants, national-level forest managers will be better placed Methods to understand the impacts of forest-cover change on Study area forest-dwelling and dependent peoples, and be able to incorporate traditional knowledge in sustainable for- This study was completed in the tropical forest and est management practices. Our work contributes to savannah biome of the Rupununi, Southern Guyana. the call by Parrotta et al. (2009) for documenting The Rupununi study area lies between 0°50ʹ – 40° 49ʹ N traditional forest-related knowledge (TFRK) in colla- and 56°54ʹ – 59°55ʹ W and is approximately 350 km boration with the holders and users of such knowl- southwest of the capital of Guyana, Georgetown edge. As a consequence, our work establishes the (Figure 1). The study area is ecologically important, basis for ecological studies on the health and status supporting a high diversity of ground-living mammals, of plants. By identifying the threats that exist to the frugivorous primates and birds (Montambault & Missa ecosystem services associated with tropical multiple- 2002; Watkins et al. 2011). Of the ground-living mam- use plants, we hope to contribute to shaping our mals, reptiles and birds, more than 107 species are understanding of ecosystem services and the value hunted by the native Amerindians for food, with that is lost as tropical forest areas decline. Through eight species (Paca or labba (Cuniculus paca), agouti the process of combining traditional knowledge with (Dasyprocta leporina), white-lipped peccary (Tayassu conventional science, this study contributes to the pecari), collared peccary (Pecari tajacu), red brocket process of allowing for more widespread inclusion deer (Mazama americana), white-tailed deer INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 43 controls), eight 4-kilometer-long transects were (Odocoileus virginianus), nine-banded armadillo established for sampling. (Dasypus novemcinctus) and red footed tortoise (Chelonoidis carbonaria)) comprising 69% of the A two-person team comprising an indigenous person with expertise in identifying plant species hunted animals (Read et al. 2010). Among these top (tree spotter) and a recorder inventoried all trees hunted species, all but the nine-banded armadillo have a strong reliance on the fruits of trees and palms in the greater than 25 cm diameter-at-breast-height (dbh) on transects. In cases where trees had an irregularity area for food (Roosmalen 1985). The area also supports or buttress at breast-height, diameter measurements the Cariban-speaking Makushi and Arawakan-speak- ing Wapishiana Amerindians (Colchester 1997)who were taken above these (Polak 1992). A minimum tree size of 25 cm dbh was chosen because tradi- have lived in the Rupununi for centuries (Forte 1996; tional knowledge suggested that most trees that Plew 2005). The Rupununi Amerindians depend on swidden provide food for wildlife produce fruits consistently agriculture, with hunting and fishing being major and vigorously at around this size. Sampling from 25 cm dbh also allowed for trees that are of interest parts of their subsistence livelihood practices. In recent times the Rupununi has been more accessible from to all resource users to be included. Commercial coastal Guyana and Brazil, meaning that the area’s species in Guyana are generally logged at stem historically protected natural resource base is becoming diameters >40 cm, while for traditional purposes, attractive for resource extraction activities, including including fuelwood, trees are harvested at various commercial logging and gold mining. Even though sizes depending on their intended uses. such changes could present communities with alterna- Furthermore, sampling at this minimum size tives to their subsistence livelihood activities allowed for plant identification errors to be reduced, (Communities of the North Rupununi 2006), it could as it assumed that tree spotters would be better able also have implications for overall forest equilibrium to identify larger trees than smaller ones, thereby (Uhl et al. 1991; Martini et al. 1994), impacting the resulting in a more accurate inventory. In addition survival of wildlife populations hunted by Amerindians to trees, mature palms (dbh of palms was not mea- and multiple-use plant species. Given the diversity of sured), with maturity defined by evidence that a ecosystem services that the forests of the Rupununi palm had borne fruits in the past (fruits, seeds or provide, and the fact that the area’sindigenouspopula- seedlings on the forest floor, empty bunches on tions retain knowledge of how plants are used, this area stems on the forest floor), were also inventoried. provided an excellent opportunity to study the various A 10-meter belt transect (see Figure 2) was used ways in which tropical plants are used, and begin to for inventorying all trees and palms within each understand how LUCC may impact these ecosystem transect’s width. For all plants that met the sampling services. In addition, scholars based at the Iwokrama criteria, inventory teams recorded: distance a plant International Center for Rainforest Conservation and Development (Iwokrama) have studied the ways in which indigenous peoples use plants in their forests (e.g. Forte 1996), providing a foundation for under- standing ecosystem services in the area. Sampling procedure and data collection Data on trees and palms were collected from 14 study sites distributed across the Rupununi land- scape (Figure 1) during the period July–December 2008. Study sites were a subset of communities and control areas participating in a larger project (see Luzar et al. 2011;Readetal. 2010;for details).As described in Read et al. (2010), study sites were selected based on their spatial distribution through- out the study area, their location with respect to vegetation types and topographic characteristics, proximity to other communities and willingness of community members to participate in the project. Figure 2. Schematic representation of the 10-meter belt The subset of study sites were the predominantly transect used for completing the inventory of trees and forested sites from those sampled by Luzar et al. palms. This schematic shows a 50-meter transect segment, (2011) and Read et al. (2010); at each study site (12 the right and left side of a transect’s imaginary centerline, Amerindian communities and two unhunted and representations of trees (circles). 44 A. R. CUMMINGS AND J. M. READ species. Wildlife and their food sources were sepa- was found along a transect; direction a plant was rated based on their importance to the Amerindian located off a transect’s imaginary centerline – right or left; distance away from a transect’s imaginary diet; hunted species were separated from unhunted species, such as frugivorous primates. centerline (offset), dbh (for trees); and local name. Distance markers were placed at 25 m intervals along transects to aid in estimating distances along a trans- Results ect. Where the local name of a tree was not known at the time of sampling, plant parts including leaves, Multiple-use species barks and fruits were collected and used to consult Our results showed that multiple-use plant groups the literature, elders and hunters to identify that tree. were the most important part of our sample, account- Trees were labeled as ‘unidentified’ in cases where ing for more than 81% of the 33,457 plants sampled. after consulting these sources a positive identification Plants were placed into one of four multiple-use was not possible. In essence, traditional knowledge of classes each describing the ecosystem services that indigenous tree spotters was utilized to identify each the groups within that class provide. The multiple- plant that met our sampling criteria. use classes were derived from the intersections of the three major resource uses: commercial timber, wild- Data analysis life food and traditional uses, and the resulting classes were used to describe the nature of plant groups. The Botanical names and resource-use classes four multiple-use classes are: The common name attached to each plant in the inven- tory was translated to a botanical name, where possible, (1) Commercial timber and wildlife food using the literature (including Roosmalen 1985;Polak (2) Commercial timber and traditional uses 1992;Forte 1996; van Andel 2000; Iwokrama 2008), and (3) Wildlife food and traditional uses in consultation with other researchers. Each plant was (4) Wildlife food, commercial timber and tradi- assigned to one or more of four major resource-use tional uses classes, based on traditional knowledge and the litera- ture of their uses in the Rupununi or elsewhere in Wildlife food/commercial timber/traditional uses Guyana for traditional and other purposes: Forty-one groups of plants were classified as wildlife food/commercial timber/traditional uses. Seventeen of (1) Commercial timber: genus or species of tree these species provide food for hunted wildlife (Table 1) actively logged for commercial timber in Guyana. while the remaining groups provide food for birds and (2) Wildlife food: genus or species of tree or palm frugivorous primates. It is possible that this class could that provide food for wildlife (hunted or not). In be expanded as in many instances one common name a few cases, data on the dispersal mode of a plant referred to more than one species. For instance, in family were used to determine animals that are Guyana crabwood generally refers to Carapa guianen- likely associated with a species in the inventory. sis, but the literature (e.g. Polak 1992; Fisch et al. 1995; (3) Traditional uses: genus or species of tree or palm Sanogo & Sacande 2007) and traditional knowledge used by Amerindians for medicinal purposes, suggested that C. procera DC may also be present in traditional weaponry, boat crafting, traditional the study area. As Fisch et al. (1995) noted the diffi- utensils, home construction, thatching, fuelwood, culty in separating these two species under normal making beverages, food, including utilizing same field conditions, local tree spotters may have refer- for economic purposes and other purposes. enced both species under the common name crab- (4) No known uses: at the time of analysis no wood. A similar situation may be applied to resource uses could be determined for this Kautaballi, which refers at least to two species, genus or species. Licania alba (Bernouli) Cuatrec and L. majuscula Based on this classification, multiple-use species were Harvey ex Gomont (Roosmalen 1985;Polak 1992). those that intersected resource-use classes 1–3. In We made no attempt to differentiate between these addition to hunted wildlife, traditional knowledge species during data collection, as the provision services and the literature (e.g. Roosmalen 1985) provided associated with them appear to transcend species. information on other mammals and birds, including The groups of plants in this class varied in terms of frugivorous primates that depend on the fruits of their range of uses, either as wildlife food sources or plants of the Rupununi for food. The resulting data- as traditional uses, while some may be more preferred base of plants included a list of tree and palm species, over others for commercial timber. In the case of a resource-use class for each species, the purpose for Carapa genus, this group has a limited range of uses which the species is used and, where appropriate, the as wildlife food, with only the hunted rodent species species of animals that depend on the fruits of that (Table 1) that disperse their seeds linked to their INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 45 Table 1. Plant groups in the commercial timber/traditional uses/wildlife food multiple-use class, and the species of wildlife associated with their fruits and seeds. Family/Species Common name Eaten by Bombacaceae Catostemma commune Sandwith Baromalli agouti, deer, tapir, howler* and spider monkeys*, bats* Pachira aquatica Aubl. Kanahia genus synzoochorous scatter-hoarding rodents, such as agoutis and acouchis Boraginaceae Cordia alliodora (Ruiz & Pav.) Oken Freijor genus endozoochorous (birds, monkeys) Burseraceae Protium decandrum (Aubl.) Marchand Kurokai agouti, marudi, powis (black curassow), deer, capuchin* Caryocaraceae Caryocar nuciferum L. Sawari nut tapir, deer, labba, spider monkey* Chrysobalanaceae Licania alba (Bernouli) Kautaballi agouti Cuatrec. and L. majuscula Harvey ex Gomont Parinari campestris Aubl. Burada macaws*, saki monkeys* Flacourtiaceae Laetia procera (Poepp.) Eichl. Warakairo endozoochorous: several birds and most primates Goupiaceae Goupia glabra Aubl. Kabukalli genus endozoochorous(howler and spider monkeys, birds) Guttiferae Calophyllum brasiliense Camb. Cassava mama genus endozoochorous (birds, monkeys) 1,2 Symphonia coccinea (Aubl.) Oken Manniballi family endozoochorous or synzoochorous (birds, monkeys) 1,2 Symphonia globulifera L.f. Manni family endozoochorous or synzoochorous(birds, monkeys) Humiriaceae Humiria balsamifera (Aubl.) A. St.-Hil. Tauroniro deer, peccaries, powis, marudi, tortoise, foxes*, macaws*, other birds* and monkeys* Lauraceae Licaria canella (Meisn.) Kosterm. Brown Silverballi family endozoochorous (specialized frugivores: toucans, cotinga, guans, spider and howler monkeys) Chlorocardium rodiei (Schomb.) Rohwer, H.G.Richt. Greenheart agouti, labba, and white lipped peccaries & van der Werff Ocotea canaliculata (Rich.) Mez; Aniba hypoglauca Silverballi family endozoochorous (specialized frugivores: toucans, cotinga, guans, Sandw. spider and howler monkeys) Lecythidaceae Eschweilera alata A.C. Smith Guava Skin monkeys* Kakaralli Lecythis corrugata Poit. Wina Kakaralli monkeys* Eschweilera decolorans Sandw. Smooth-Leaf labba, several monkeys* Kakaralli Eschweilera sagotiana Miers Common Black monkeys, birds* Kakaralli Eschweilera wachenheimii (Benoist) Sandw. Fine Leaf Kakaralli monkeys* Lecythis cf. chartacea Berg Monkey Pot – labba small Lecythis zabucajo Aublet Monkey Pot labba Leguminosae (Caesalpiniaceae) Hymenaea courbaril L. Locust agouti, labba, tapir, peccaries, monkeys* (saki, capuchin) Mora excelsa Benth. Mora peccaries, deer and tapir eat newly germinated seedlings Peltogyne venosa (Vahl) Benth. Purpleheart genus endozoochorous (especially spider monkeys) Peltogyne paniculata Benth. Purpleheart saka genus endozoochorous (especially spider monkeys) Leguminosae (Mimosaceae) Inga alba (Sw.) Willd. Maporakon agouti, labba, tapir, deer, powis, marudi, peccaries, parrots*, capuchin* and spider* monkeys Abarema jupunba (Willd.) Britton & Killip Huruasa endozoochorous spider monkeys Leguminosae (Papilionoideae) Dipteryx odorata (Aubl.) Willd. Tonka bean toucans* and other birds* Ormosia coccinea (Aubl.) Jackson Barakaro peccaries Swartzia leiocalycina Benth. Wamara genus zoochorous (bats, rodents, monkeys) Meliaceae Carapa guianensis Aubl., C. procera DC Crabwood agouti, labba, peccaries Moraceae Bagassa guianensis Aubl. Cowwood tapir, deer, peccaries, labba, agouti Brosimum guianense (Aubl.) Huber Letterwood endozoochorous spider monkeys and birds Myristicaceae Virola surinamensis (Rol.) Warb. Swamp Dalli monkeys, toucans and other birds Sapotaceae Manilkara bidentata (A.DC.) Chev. Bulletwood peccaries, tapir, deer, tortoise, labba, agouti, spider* and howler monkeys*, macaws* Pouteria cuspidata (A.DC.) Baehni Kokoritiballi agouti, red brocket deer, labba, monkeys* Pouteria guianensis Aubl. Asepoko agouti, red brocket deer, labba, spider* and howler monkeys*, capuchins* Simaroubaceae Quassia spp. Angelina Rock genus endozoochorous (spider and howler monkeys) Quassia simarouba L.f. Simarupa genus endozoochorous (spider and howler monkeys) *species not hunted – data from traditional knowledge and Roosmalen (1985) data from Roosmalen (1985) data from Forget and Hammond (2005) **the scientific names of hunted wildlife species are provided in the text. Species listed above are: howler monkey (Alouatta seniculus); spider monkey (Ateles paniscus); white saki (Pithecia pithecia); bats (Artibeus lituratus among other species); black saki (Chiropotes chiropotes); capuchin (Cebus apella, Cebus olivaceus); toucans (Ramphastos toco, Ramphastos vitellinus); and macaws (Ara ararauna, Ara macao, Ara nobilis). 46 A. R. CUMMINGS AND J. M. READ traditional knowledge suggested that the fruits of fruits (Forget et al. 1999; Forget & Jansen 2006). The Ormosia coccinea (Aubl.) Jackson are eaten by pecc- traditional value of Carapa spp. is, however, in little doubt; the seeds are widely utilized throughout aries, but Roosmalen (1985) noted that the genus is endozoochorous with birds mainly dispersing these Guyana and Amazonia (van Andel 2000; Shanley & seeds. In the case of Licania alba and L. majuscula Luz 2003; Sullivan & O’Regan 2003; Plowden 2004; Klimas et al. 2007; Costa-Silva et al. 2008) for its (Kautaballi) van Andel (2000) noted that the name given to L. alba by the Warrau people of the North valuable oil (see also Klimas et al. 2007). Similarly, West District (NWD) translates to ‘agouti food tree’ as Chlorocardium rodiei Schomb.) Rohwer, H.G.Richt. & van der Werff (greenheart) has only been asso- these rodents feed on the seeds of the tree. The tradi- tional uses of Eschweilera alata A.C. Smith (house ciated with hunted scatter-hoarding rodents posts), E. sagotiana (head straps and lashing materials, (Hammond et al. 1999) and peccaries (traditional knowledge). However, like the species in the Carapa house construction, and seeds used as bait in bird genus, C. rodiei is a primary target for commercial traps), E. wachenheimii (bark used for head straps and lashing materials and house construction), L. corrugata logging in Guyana (Zagt 1997; Ter Steege et al. 2002). Further, C. rodiei is listed by the International Union (house construction), Virola surinamensis (Rol.) Warb. for the Conservation of Nature and Natural Resource (used for making Arawak musical instruments and the (IUCN) as a threatened species (IUCN 2012). The sap is used for treating thrush in babies and also gargled traditional uses of C. rodiei are primarily medicinal, for tonsillitis and toothaches), Pouteria cuspidate (house with the seeds being used to treat parasitic worms construction) and Lecythis cf. chartacea Berg (lashing and skin diseases (Forte 1996). material) were noted by van Andel (2000)for their In contrast to the limited range of uses associated utility in the NWD. It seems likely that these species with plants in the Carapa genus as wildlife food are similarly utilized in the Rupununi, but this was not sources, Manilkara bidentata (A.DC.) Chev. may be revealed during data collection or through the literature described as a universal wildlife food source. It was (e.g. Forte 1996). suggested, in particular by traditional knowledge, that Sixteen groups in this class were included based solely all the hunted wildlife species depend on the fruits of on the data provided by Roosmalen (1985)ontheir M. bidentata. Furthermore, M. bidentata has long interaction with wildlife. Abarema jupunba (Willd.) been recognized as a commercial timber species Britton & Killip, Anibaspp.,Brosimumguianense (Polak 1992). As an NTFP, Amerindians have used (Aubl.) Huber, Calophyllum spp., Cordia alliodora balata, the gum derived from M. bidentata, to make (Ruiz & Pav.) Oken, Goupia glabra Aubl., Licaria canella utensils for storing water and beverages, and the (Meisn.) Kosterm. Laetia procera (Poepp.) Eichl, Ocotea fruits of M. bidentata are eaten (Forte 1996). spp Quassia simarouba, Pachira aquatica Au bl., Indeed, there was once a thriving trade in the sap of Peltogyne paniculata Benth., Peltogyne venosa (Vahl) M. bidentata from the Rupununi. Currently the Benth, Swartzia leiocalycina Benth. and Quassia spp. dependence on this species as an NTFP has waned, were listed at the genus or species level as being either with its consumption generally restricted to making synzoochorous (animals that bury or store seeds) or figurines of animals and common articles found in endozoochorous (dispersal of seeds after passing the Rupununi for the tourism market. through the gut of an animal). Among these, only However, in addition to the most abundant groups Pachira spp. was listed as being dispersed by hunted – Catostemma commune Sandwith, Parinari campes- wildlife (agoutis and acouchis). In two exceptional tris Aubl. Mora excelsa Benth. and Protium decan- cases, Roosmalen (1985)noted Symphonia coccinea drum (Aubl.) Marchand – being Aubl. and Symphonia globulifera L.f. to be dispersed at included in this class (Table 1), there were also groups with low abun- the family level synzoochorously or endozoochorously, but Forget and Hammond (2005) indicated that this dance. For example, Caryocar nuciferum L. fruits are genus is an important food source for the red brocket eaten throughout forested Guyana, with fruits and trees observed at various Rupununi homes during deer. The other groups are primarily of interest to fru- givorous primates and birds. Traditionally, Pachira spp. data collection, although trees were only observed at two study sites – Villages 18 and 23. and L. procera are utilized as fuelwood, while C. allio- In some instances the traditional uses of groups in dora, Calophyllum spp., G. glabra, Peltogyne spp.and Quassia spp. are used in house construction and for this class were well established, but not so their value as wildlife food. M. excelsa illustrates this point as tradi- making canoes. The bark of G. glabra has medicinal tional knowledge suggested that the fruits and seeds of applications, with the boiled bark being used to treat eczema andtheleaves andboiledbarkareusedforthe M. excelsa are eaten by paca and peccaries and that deer, tapir and peccaries browse on the newly germinated treatment of chickenpox and to relieve toothache (van seedlings. However, the literature, in particular Andel 2000). P. venosa is used in house construction, A. jupunba is used as a soap substitute (van Andel 2000) Roosmalen (1985), noted that M. excelsa is dispersed by water, that is, they are hydrochorous. Similarly, and Brosimum spp. isusedtomakebowsand walking INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 47 other genus including Cordia are also referred to by sticks. S. coccinea and S. globulifera are used for making this common name. This common name was therefore canoes in both the Rupununi and the NWD. Furthermore, van Andel (2000)pointedoutthatasa a point of uncertainty in our analysis for identification, but all of these species are used traditionally, primarily in medicinal product, the boiled bark of S. coccinea and S. house construction and for making paddles (Forte globulifera is used to treat abscesses and diarrhea and thrush in babies. The traditional uses of Licaria canella, 1996). Forte (1996)points out toothat Swartzia leioca- lycina is a much-favored fuelwood, while Quassia simar- Ocotea spp., Aniba spp., Swartzia leiocalycina and ouba is used by the Makushi in house construction. Quassia simarouba were noted by Forte (1996)based on the knowledge of the Makushi people (primarily women) of the North Rupununi. Ocotea spp., Aniba Commercial timber/wildlife food spp., Swartzia leiocalycina and Quassia simarouba are The commercial timber/wildlife food class contained all listed by Polak (1992) as major timber species. Licaria 20 groups of plants. These plants, even though they cannella is used in house construction, and the bark is provide food for hunted wildlife, are more important used medicinally and for tanning leather. Members of for the frugivorous primate population of the the Ocotea genus including Ocotea canaliculata (Rich.) Rupununi and birds, with only seven species linked Mez, (Iwokrama 2008) along with members of the to hunted wildlife (Table 2). Sixteen groups of plants Aniba genus including Aniba hypoglauca Sandw. are were included in this class based primarily on data commonly referred to as Silverballi, but members of from Roosmalen (1985) and are dispersed mostly by Table 2. Plant groups in the commercial timber and wildlife food class and the species of wildlife associated with their fruits and seeds. The relationship between these plants and hunted wildlife is highlighted. Eaten by Hunted Family/Species Common name Wildlife? Eaten by Bombacaceae Catostemma fragrans Benth. Sand Baromalli Yes agouti, red and gray brocket deer, tapir, bats, howler and spider monkeys* Burseraceae Trattinnickia rhoifolia Willd. Ulu No genus endozoochorous (especially birds) Guttiferae Calophyllum lucidum Kuruhara No genus endozoochorous Benth. Platonia insignis Mart. Kaslego Yes monkeys, peccaries* Lauraceae Aniba hypoglauca Sandw. Yellow Silverballi No family endozoochorous (specialized frugivores: toucans, cotinga, guans, spider and howler monkeys) Ocotea canaliculata (Rich.) Mez White Silverballi No family endozoochorous (specialized frugivores: toucans, cotinga, guans, spider and howler monkeys) Ocotea rubra Mez Determa No family endozoochorous (specialized frugivores: toucans, cotinga, guans, spider and howler monkeys) Ocotea splendens (Meisn.) Mez. Kereti Silvaballi No family endozoochorous (specialized frugivores: toucans, cotinga, guans, spider and howler monkeys) Lecythidaceae Eschweilera chartacea (O.Berg) Broad Leaf Kakaralli Yes parrots, macaws*; genus synzoochory: (scatter-hoarding rodents) Eyma and several monkey species Eschweilera pedicellata (Rich.) White Skin Kakaralli Yes genus synzoochorous: (scatter-hoarding rodents) and several monkey species Eschweilera subglandulosa (Steud. Black Kakaralli Yes macaws, parrots*; genus synzoochory: (scatter-hoarding rodents) ex O.Berg) Miers and several monkey species Lecythis confertiflora (A.C. Sm.) Werimiri Kakaralli No genus endozoochorous (spider monkeys) and synzoochorous (bats) S.A. Mori Leguminosae (Caesalpiniaceae) Mora gonggrijpii (Kleinhoonte) Morabukea Yes peccaries, deer and tapir eat newly germinated seedlings* Sandwith Leguminosae (Papilionoideae) Ormosia coutinhoi Ducke Korokororo No genus endozoochorous (birds) Swartzia benthamiana Miq. Itikiboroballi No genus zoochorous (bats, rodents, monkeys) Myristicaceae genus zoochorous (bats, rodents, monkeys) Iryanthera lancifolia Ducke Kerikowa No genus endozoochorous (specialized frugivores, such as toucans, guans, cotingas, monkeys) Iryanthera macrophylla (Benth.) Black No genus endozoochorous (specialized frugivores, such as toucans, Warb. Kerikowa guans, cotingas, monkeys) Virola michelii Heckel Hill Dalli No genus endozoochorous (specialized frugivores, such as toucans, guans, cotingas, monkeys) Sapotaceae Pouteria speciosa (Ducke) Baehni Suya Yes labba, agouti, peccaries, tapir* Verbenaceae Vitex stahelii Moldenke Hakiaballi No genus endozoochorous (spider and howler monkeys) data from Roosmalen (1985) data from Forget and Hammond (2005) *data from Rupununi traditional knowledge 48 A. R. CUMMINGS AND J. M. READ frugivorous primates, either endozoochorously or Eperua falcata Aubl., Tabebuia insignis (Miq.) synzoochorously. In a few instances, scatter-hoarding Sandwith, Tabebuia serratifolia (Vahl) G. Nicholson rodents also played a role in dispersing these seeds. and Pterocarpus rohrii Vahl; Polak 1992). It was also Furthermore, Forget and Hammond (2005) sug- suggested by traditional knowledge that Thyrsodium gested, based on data provided by Gayot (2000) and guianensis Sagot is a timber species. However, the Gayot et al. (2004), that Swartzia benthamiana Miq. data suggested that the traditional uses of the species is a food source for red and gray brocket deer. Forget in this multiple-use class are not universal. Some species are only used for traditional purposes in the and Hammond (2005) also suggested that the fruits of most of the groups in the Leguminosae family, a Rupununi, while others are only used in the NWD number of which are included in this class, are eaten and coastal Guyana (Table 3). The fact that the tradi- tional uses of these plants is not universal indicates by the wildlife species hunted in the Rupununi. The fact that most of the groups of plants in this class are the cultural value that different groups of people give of more importance to frugivorous primates than to plants, and is an area that requires further analysis in Guyana and where these species are found across hunted wildlife is critical, as primates play important ecological functions in maintaining the forest struc- the Neotropics. ture (Roosmalen 1985; Forget & Hammond 2005). The interactions between plants and animals is an Wildlife food/traditional uses area that requires further research as Forget and There were 39 groups of plants within the wi ldlife food/ Hammond (2005) suggested that the diet of the pri- traditional uses class (Table 4). The palms (eight spe- mates and ungulates (deer, tapir and peccaries) needs cies) dominated this class of multiple-use plants, and to be better understood. this indicates the critical role palms play in traditional practices and NTFPs extraction from tropical forests. Commercial timber/traditional uses The fruits of palm are eaten by people and the wildlife The commercial timber/traditional uses class (Table 3) and the leaves of many are harvested for thatching roofs and basketry making (Forte 1996). But like traditional was constituted of 11 groups of plants. Of these, seven groups are major timber species (Clathrotropis uses of plants in the commercial timber and traditional brachypetala (Tul.) Kleinhoonte, Clathrotropis macro- uses, not all uses appear universal. For example, the fruits of Oenocarpus bacaba Mart. and Oenocarpus carpa Ducke, Diplotropis purpurea (Rich.) Amshoff, Table 3. Plant groups in the commercial timber and traditional uses class, highlighting their traditional uses and the region of Guyana that these uses have been attributed. Family/Species Common name Traditional Uses Region Bignoniaceae Tabebuia serratifolia Ironwood used in house construction, to make 8–9-foot-long posts; bark used to All (Vahl) G. prevent scabies; young leaves used to preserve teeth Nicholson Tabebuia insignis White cedar house construction Rupununi (Miq.) Sandwith Leguminosae (Caesalpiniaceae) Eperua falcata Aubl. Wallaba used to make posts, shingles and bows; bark used medicinally to treat Rupununi internal injuries Leguminosae (Papilionoideae) Centrolobium Paurine used in house construction, to make posts; also paddles and good Village 19 has a paraense Tul. fuelwood conservation area to protect this species Clathrotropis Aromata – fine leaf bark scraped and applied to leishmaniasis parasite wounds to kill the NWD brachypetala (Tul.) parasite. Drinking the sap from the bark could be used to treat the bite Kleinhoonte from the water coral snake: Helicops angulata Clathrotropis Aromata bark scraped and applied to leishmaniasis parasite wounds to kill the NWD macrocarpa Ducke parasite. Drinking the sap from the bark could be used to treat the bite from the water coral snake: Helicops angulata Diplotropis purpurea Tatabu used to make canoes NWD (Rich.) Amshoff Pterocarpus rohrii Hill Corkwood used for making canoes, paddles, carved toys All Vahl Pterocarpus spp. Corkwood used for making canoes, paddles, carved toys All Meliaceae Tabebuia stenocalyx Water Cedar boards used in house construction, to make shingles, dugout canoes Rupununi Sprague & Stapf. Sapindaceae Thyrsodium Sand Mora fish poison NWD guianensis Sagot van Andel (2000), NWD data based on van Andel (2000), other data from Rupununi traditional knowledge and Forte (1996). INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 49 Table 4. Multiple-use tree species in the class wildlife food and traditional uses. Family/Species Common name Eaten by Traditional uses Region Anacardiaceae Anacardium giganteum Hancock ex Hubudi/wild cashew tapir, tortoise, deer, powis, peccaries, monkeys (capuchin, fruits eaten and used for making beverages NWD Engl. spider) Mangifera indica L. Mango domesticated animals, wildlife fruits eaten All Spondias mombin L. Plum tapir, agouti, labba, tortoises, monkeys (squirrel, howler, fruits eaten, various beverages, seeds used as fish bait, soot of roasted seeds used to treat All spider, capuchin), toucans** wounds from vampire bats, used to treat hemorrhaging post childbirth, bark and leaves used to treat sores and tea from bark used for colds Tapirira guianensis Aubl. Duka hanaqua, marudi, some monkeys, foxes** fuelwood Rupununi Annonaceae Duguetia calycina Benoist . Yari Yari genus zoochorous (monkeys, scatter-hoarders) bows, fishing rods NWD Unonopsis glaucopetala R.E.Fr. Rough Skin Arara endozoochorous (spider monkeys, cotingas) treating the bite of labaria (Bothrops asper) or fer-de-lance NWD Xylopia cayennensis Maas Black Maho genus endozoochorous (monkeys, birds) astringent tea for diarrhea; firewood; bark used for warashis straps; decoction also drunk by NWD; pregnant women as an aid to eventual easy delivery Rupununi Xylopia cf. surinamensis R.E. Fr. Kuyama* genus endozoochorous (monkeys, birds) astringent tea for diarrhea; firewood NWD; Rupununi Apocynaceae Geissospermum argenteum Woodson Sinkona tortoises; genus endozoochorous(monkeys, birds) bark boiled for treating malaria – source of quinine All Burseraceae Protium guianense (Aubl.) Marchand. Haiawa genus endozoochorous (toucans, cotingas, guans, spider gum used to cure diarrhea; mixed with cow fat and used to caulk boats in the absence of Rupununi monkeys, howler monkeys, tortoises, kinkajous, karamani gum (maitakin), gum used to anchor grater’s teeth firmly on board; burned as incense opossums) under the hammock of a sick person; fruits eaten Boraginaceae Cordia tetrandra Aubl. Grandma Cherry birds and monkeys** fruits used as glue for paper and making kites NWD Chrysobalanaceae Chrysobalanus icaco L. Fatpoke birds** fruits eaten All Ebenaceae Diospyros ierensis Britton Barabara tapir and birds fuelwood Rupununi Guttiferae Clusia grandiflora Splitg. Kufa genus endozoochorous birds sometimes monkeys used in the furniture industry All Tovomita schomburgkii Planch. & Awasokule genus endozoochorous house construction, fuelwood NWD Triana Vismia macrophylla Kunth Bloodwood genus endozoochorous (birds) latex used to treat skin fungi (ringworm, lota, ground itch) All Lecythidaceae Bertholletia excels Bonpl. Brazil Nut agouti, labba, peccaries and monkeys fruits eaten Rupununi Leguminosae (Caesalpiniaceae) Hymenaea oblongifolia Huber Locust -fine leaf agouti, labba, peccaries, monkeys (saki and capuchin) powdery pulp eaten and beverage made from bark All (Shimiri) Leguminosae (Mimosaceae) Inga huberi Ducke Fern Tree agouti, labba, tapir, deer, powis, marudi, peccaries, fruits of various Ingas eaten; fuelwood All monkeys (capuchin, spider), parrots Inga rubiginosa (Rich.) DC. Wild Tamarin agouti, labba, tapir, deer, powis, marudi, peccaries, fruits of various Ingas eaten; fuelwood All capuchin, spider, parrots Leguminosae (Papilionoideae) Swartzia arborescens (Aubl.) Pittier Bowwood genus zoochorous (bats, rodents, monkeys) used for making bows Rupununi Malpighiaceae Byrsonima aerugo Sagot Moreye tapir, deer, marudi, powis, chachalaca, labba, agouti, fruits eaten, fuelwood All peccaries, tortoises, parrots, hanaqua, toucan, parrots, macaw, savanna fox, monkeys (Continued) 50 A. R. CUMMINGS AND J. M. READ Table 4. (Continued). Family/Species Common name Eaten by Traditional uses Region Byrsonima stipulacea A.Juss Hicha tapir, deer, marudi, powis, chachalaca, peccaries, labba, fruits eaten, fuelwood All agouti, tortoises, hanaqua, toucan, parrots, macaw, savanna fox, monkeys Moraceae Cecropia peltata L. Congo pump marudi, hanaqua, capuchin monkeys leaves of some species used for making teas for treating kidney problems, back pain and high All blood pressure. Moraceae (Cecropiaceae) Pourouma guianensis Aubl. Baruma birds and monkeys** fuelwood, leaves used like sandpaper all Myrtaceae Myrciaria vismeifolia (Benth.) O.Berg Taparowballi most animals and birds fruits used to make beverage Rupununi Palmae Astrocaryum aculeatum G.Mey. Akayuru tapir, deer, peccaries, labba, agouti, squirrel, rats, fleshy endocarp eaten All Astrocaryum vulgare Mart. Awara agouti, labba, peccaries, tapir fleshy endocarp eaten All Attalea maripa (Aubl.) Mart. Kokerite tapir, agouti, labba, tortoise, deer, black curassow, fruits eaten, used to make beverages, and leaves used for thatching roofs All regions marudi, macaws, parrots Euterpe oleracea Mart. Manicole deer, peccaries, marudi, hanaqua, agouti, labba, tapir, fruits used to make beverages in the Rupununi, heart-of-palm eaten as a vegetable (cooked or All regions macaws, parrots raw) Manicaria saccifera Gaertn. Trulli** monkeys, scatter-hoarding rodents, such as agoutis and leaves used for thatching in NWD NWD acouchis Mauritia flexuosa L.f. Ite tapir, deer, peccaries, powis, labba, agouti, acouchis, leaves used for thatching in Rupununi, fruits eaten all over Guyana Thatch tortoises, macaws, parrots, monkeys (howler and spider) Rupununi, eaten all Oenocarpus bacaba Mart. Lu deer, marudi, powis, hanaqua, agouti, labba, tapir, fruits used to make beverage All peccaries, macaws, parrots Oenocarpus bataua Mart. Turu deer, peccaries, marudi and powis, hanaqua, agouti, fruits used to make beverage All labba, tapir, macaws, parrots Sapotaceae Pouteria caimito (Ruiz & Pav.) Radlk. Asepokoballi agouti, deer, labba, spider and howler monkeys, fruits eaten All capuchins Pradosia schomburgkiana (A.DC.) Kakarua tapir, deer, peccaries, monkeys fruits eaten, bark boiled with other species to treat tuberculosis All Cronquist Sterculiaceae Sterculia pruriens (Aubl.) K.Schum. Maho agouti, labba, rats lashing material for warashis, quakes, and other traditional utensils in NWD All Sterculia rugosa R. Br. Birdwood genus synzoochronous scatter-hoarding rodents, wattles used in house construction Rupununi capuchin and saki monkeys Violaceae Paypayrola longifolia Tul. Adebero labba, agouti, peccaries, monkeys, birds fuelwood Rupununi INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 51 bataua Mart. are eaten in the Rupununi, but it is not so 2008) within the study area. There were, however, for the fruits and heart-of-palm of Euterpe oleracea potentially more than 165 groups of plants in the sam- Mart. in this region, despite heart-of-palm being ple. It is acknowledged that trees in the forests of the exported from the NWD. Perhaps the most important Rupununi, like other Neotropical settings, are difficult palm, given that its fruits are eaten by people and wild- to identify (see Dinerstein 2013), and even though 3% life and the leaves are used for thatching roofs, is Attalea did not represent a large portion of our sample, it still maripa Mart. (Kokerite). A key food source for hunted referenced species that tree spotters had not worked rodent species (Silvius & Fragoso 2003), this was the with in the past. It is possible too that some species most abundant species in this class. In fact, to highlight were misidentified, especially in the process of transfer- the favorability of the fruits in the Rupununi, cut ring common names to botanical names. In many instances, the literature (e.g. Roosmalen 1985;Polak stumps of this palm were often observed in the land- scape, reflecting the desire of the cutter to gain access to 1992; van Andel 2000) on a number of species had the ripe fruit on palms that were deemed too tall. This one common name referring to multiple botanical names. Species identification remains a challenge in multiple-use class also included a number of tree spe- cies. Geissospermum spp. or Sinkona is probably the reconciling traditional knowledge with conventional most significant. A medicinal plant, the bark of this science, and traditional knowledge provides an avenue tree is used to treat malaria and the fruits are eaten by for dealing with this challenge. It is therefore critical, as tortoises. It was suggested (van Andel 2000)that Mauro and Hardison (2000) pointed out, to preserve Sinkona is utilized in coastal Guyana, with reference traditional knowledge. Identifying all species of plants to Cinchona spp. This is an area of debate in this analysis with ecological and cultural significance is an area that as tree spotters at Iwokrama suggested that this refer- scientists must work with indigenous peoples, such as ence may be to trees in the genus Geissospermum and the tree spotters involved in this study, for clarification. not Cinchona.Additionally, Myrciaria vismeifolia In this way, traditional knowledge can play a much (Benth.) O.Berg is used to make a beverage, while the more prominent role in guiding forest management fruits of Byrsonima species are eaten throughout the decision-making processes. The difficulties of identify- Rupununi and Guyana, and traditional knowledge sug- ing plants aside, tree spotters have worked extensively gested that hunted wildlife also eat these fruits. The with the major timber and well-known species, such as fruits of M. vismeifolia are eaten by most animals and Manilkara bidentata, Mora excelsa, Carapa guianensis birds, including hunted wildlife species. Pouteria cai- Aubl. and Inga alba (Sw.) Willd. in the logging industry mento (Ruiz & Pav.) Radlk. and Pradosia schomburgki- and their knowledge of plants has been relied upon in ana (A.DC.) Cronquist are eaten by hunted wildlife, other settings. This fact gives us confidence in their especially the hunted rodents and deer species. knowledge they shared for this study. The large sample Anacardium giganteum Hancock ex Engl., Mangifera size utilized in this study, and sampling larger plants, indica L. and Spondias mombin L. are popular with coupled with the TFRK of indigenous tree spotters Amerindians and throughout Guyana. M. indica,a allowed us to establish that multiple-use plant species cultivated species, was observed on old farms through- are the foundation of the Rupununi forested ecosystem. out the study area, and is a delicacy throughout Guyana. Multiple-use plant species and the ecosystem services S. mombin fruits are sold at local markets and used associated with them have therefore allowed for a throughout Guyana as well for making a beverage. greater appreciation of what is at stake as forest areas Protium guianense (Aubl.) Marchand fruits are eaten undergo change due to LUCC processes and perhaps by a number of the hunted wildlife species and people. larger processes of climatic variability. The gum of the Protium trees, which has a camphor- like odor, is used in several ways by Rupununi Multiple-use classes Amerindians, including treating diarrhea and is burned as incense under the hammock of a sick person. This This analysis showed that multiple-use trees and multiple-use class, despite not being the most abundant, palms dominate the forests of the Rupununi. Four is important to the Amerindian culture, and requires dimensions of multiple-use plants were identified: additional examination to better understand how these wildlife food/commercial timber/traditional uses, com- species will be affected by logging and other commercial mercial timber/wildlife food, commercial timber/tradi- exploitation within their habitats. tional uses and wildlife food/traditional uses. Each multiple-use class portrays a different facet of tropical forest utilization, ecosystem services and indeed Discussion and conclusions potential for resource-use conflicts. Because of the different demands on the various multiple-use Species composition classes, the groups of plants in each class will be All 165 groups of plants in our sample were recorded by impacted differently by LUCC. Based on our classifi- previous studies (e.g. Clarke et al. 2001; Iwokrama cation, and the relationships that have been 52 A. R. CUMMINGS AND J. M. READ better off for adopting traditional knowledge in established between plants and people, and plants informing policy positions. These positions could and wildlife (e.g. Roosmalen 1985; Forte 1996; van Andel 2000; Forget & Hammond 2005; Cummings include guiding processes of plant removal for tim- ber, and for also deciding on spaces to be preserved 2013), we argue that the removal of trees in the class for traditional activities. While other studies have wildlife food, commercial timber and traditional uses, for example, will have implications for wildlife and focused on the gross-scale importance of tropical forests for regulating services such as carbon seques- people. For instance, wildlife perform ecosystem tration (see for example Houghton 2005), this study functions and supporting services for plants (see Roosmalen 1985; Forget & Hammond 2005) while was able to describe finer-scale detail for ecosystem services associated with tropical plants. As a conse- plants are associated with provisioning services quence we are contributing to establishing for forest (such as traditional medicines) and cultural services of Amerindians (see Forte 1996; van Andel 2000). managers the value that may be lost as tropical forest Since these plants are connected to each of the main areas decline due to LUCC and indeed larger pro- cesses such as climatic variability. As Parrotta and resource-use classes identified in the Rupununi – commercial logging, provision of wildlife food and Agnoletti (2007) pointed out, traditional knowledge traditional uses – a lack of sound management prac- of rural communities, in our case the Makushi and tices for these species will have far-reaching conse- Wapishiana Amerindians, has served to sustain the quences for local populations and indeed forest goods and services they need for livelihood security, ecosystem processes. Forest managers accessing tradi- and as such measures should be taken to preserve tional knowledge on how such plants are used can be traditional knowledge. Activities such as deforesta- a critical step in preventing impacts on indigenous tion as it relates to gold mining, commercial logging people’s livelihood and wildlife. and large-scale farming that are features of tropical landscapes may remove plants while bringing with them change, such as socioeconomic pressures, that Synthesis and future directions may erode traditional knowledge. As such, we concur with the position of Parrotta and Agnoletti (2007) The objectives of this paper were to identify tropical and others who suggest that efforts are required to multiple-use plants and describe the ecosystem ser- arrest the decline of traditional knowledge in indi- vices associated with these in order to begin the genous people-produced landscapes. process of understanding what value is lost as tropical In essence then, we argue that the identification of forest areas decline. Our analysis has shown that multiple-use species using traditional knowledge is there are many multiple-use plants in tropical forests critical, because it provides knowledge of the goods that require special attention for ecological studies. and services that are kept within tropical landscapes. By exploring multiple-use plants through the lens of Furthermore, while we agree with the position of how they are used as commercial timber, wildlife Sutherland et al. (2014) that traditional knowledge food and traditional purposes, and by drawing on ‘. . .can provide complementary perspectives . . .often traditional knowledge to identify these plants and lacking in conventional science’, we emphasize that define their importance, we suggest that as tropical traditional knowledge can provide the basis for scien- forest areas decline the potential impact may be bet- tific inquiry into questions of both landscape ecology ter appreciated. and economics. Our work suggests that traditional Our analysis would not, however, have been pos- knowledge can provide insights to scientists on how sible without the availability and accessibility of tra- to fully grasp the impacts of LUCC, beyond the ditional knowledge. In fact, without the utilization of typical measurements of area loss (e.g. Kim et al. traditional knowledge in two dimensions – spotting 2015) or implications for the carbon cycle (Melillo (identifying) plants in the forest and describing how et al. 1996; Baccini et al. 2012; Zarin 2012; Kim et al. these plants are used for traditional purposes, wildlife 2015). While these areas are important, our work food and commercial timer (Tables 1–4) – the value suggests that attention needs to be paid to the of our study would have been significantly reduced. impacts of LUCC on the multiple roles forests and In this sense, this paper is a direct response to the the individual species of plants within them play for calls from scholars such as Sutherland et al. (2014), humanity and wildlife. Traditional knowledge pro- Fazey et al. (2006) and Raymond et al. (2010) for the vides that basis of developing such knowledge. greater use of local and traditional knowledge along- Furthermore, more indirectly, and perhaps more sub- side conventional knowledge for making decisions tly in the immediate term, are the impacts of climatic about biodiversity and natural resources. While we variability on the services traditional knowledge can view our study as a part of the process for beginning inform us about. Scholars and forest managers alike to value tropical landscapes for the ecosystem services need to better understand the conditions that lead to they provide, we do make the case that forest man- overexploitation of forests as this will help them agement decision-making processes are going to be INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 53 addressing pressing ecological questions on the better appreciate the value that is lost to humanity as impacts of forest change. Furthermore, as more forest areas decline. Such an understanding will lead to a clearer picture of the value that is lost to human- work unfolds on how plants are used by rural communities through the examples of González- ity and wildlife as forest area is lost. By responding to Marín et al. (2012) in Mexico, Khan et al. (2013) the call of Parrotta et al. (2009) for documenting TFRK, our paper has established, through multiple- in Pakistan, Klimas et al. (2007, 2012a, 2012b)in Amazonia and Mwavu and Witkowski (2009)in use plants, some of the complexities and nuances that Africa, our work contributes to setting up species are associated with TFRK that must be accounted for in the process of valuing the losses to humanity as and their importance for more in-depth analysis on how changes in landscape can impact ecosystem tropical forest areas decline. In this sense, traditional services and landscape sustainability. In the knowledge on tropical landscapes can serve both as a basis for understanding the value of landscapes Guyana setting, the work of Forte (1996) and van change, but also for the justification for investment Andel (2000) established the foundation for describing the relationship between indigenous peo- in more robust conservation and management strate- gies within these landscapes. ples and plants. Our work builds on these past As the literature on multiple-use species continues efforts by providing insights into plant taxonomy to evolve, the interaction between the various (Mauro & Hardison 2000) and ecosystem services resource uses in tropical forests, such as wildlife of individual plants, and provides the basis for food sources, commercial timber and traditional further spatial and socioeconomic analysis. In addi- uses, will require urgent attention. The conflicts tion, our work goes beyond previous studies, such between wildlife food sources and commercial log- as Scarpa (2009), whose data were based on pub- ging, for example, have only been reported in a few lished studies, Singh et al. (2015), Klimas et al. instances (Plumptre 1995; Mwavu & Witkowski (2007, 2012a, 2012b) who have considered single 2009). Mwavu and Witkowski (2009) identified 15 species, or McPherson (2014), where species distri- such multiple-use species in Budongo Forest bu tion models were derived from museum samples, Reserve, Uganda, while the present study suggests by providing a wider list of plants, through direct that more than approximately 61 such species are in interactions with the holders of TFRK as suggested this potential resource-use conflict area and should by Parrotta et al. (2009). therefore gain the attention of more detailed ecologi- While our work provides a list of plants and how cal studies. The intersection between wildlife food they are used for various ecosystem services, data on and commercial logging in tropical plant species is the current status of such species are not presented in critical, especially in those communities that are mov- the paper, including the abundance of species and the ing toward commercial logging as an alternative for level of dependence that local people have on each supporting their livelihoods. As various sectors of a both at the scale of the study area and at individual community embrace new technology at different study sites. The status of the ecosystem services rates, this change can potentially lead to local-scale associated with individual plant species is perhaps resource-use conflicts, where logging may place pres- the most pressing concern, as this will provide sure on the very food source hunted wildlife rely insights into the sustainability of local livelihood upon. practices and larger sustainable forest management Besides the species of wildlife that are hunted in questions. Traditional knowledge will be critical in the Rupununi, the needs of frugivorous primates addressing these concerns and helping craft immedi- should also be included in future analysis. For exam- ate changes in management practices to deal with the ple, Forget and Hammond (2005) pointed out that impacts of LUCC under various scenarios of resource even carnivorous species, such as coati living in the exploitation. forests of the Guiana Shield, may eat the fruits and Finally, in following up on the prescription for parts of trees in their habitat. Therefore, as suggested dealing with the hindrances to sustainable forest by Peters (1996) and Forget and Hammond (2005), management (Peters 1996), future research in the understanding which plant species provide food for Rupununi, and indeed areas where tropical forests animals, their demographics and how they respond to are continuing to decline, should focus on the spa- disturbance (such as commercial logging) is critical, tial distribution of multiple-use species and in the context of both the study area and the wider resource extraction activities. The list provided is Neotropics. With the high number of multiple-use this study can serve as the basis for local scale species in the forests of the Rupununi, it would (Rupununi) to larger scale – Guyana and the seem likely that there are also high numbers else- wider Neotropics – analysis of the implications of where in Guyana and the wider Neotropics. LUCC on the ecosystem services associated with Through embracing traditional knowledge, our multiple-use plants. Furthermore, through the list studyprovidesscientistswithastronger basisfor of plants provided in this paper, further studies can 54 A. R. CUMMINGS AND J. M. READ Baccini A, Goetz SJ, Walker WS, Laporte NT, Sun M, be established to determine the contribution LUCC Sulla-Menashe D, Hackler J, Beck PSA, Dubayah R, is making to the erosion of TFRK in indigenous Friedl MA, et al. 2012. Estimated carbon dioxide emis- peoples-produced landscapes of Guyana and sions from tropical deforestation improved by carbon- beyond. Specific questions on where exactly across density maps. Nat Clim Chang. 2:182–185. a species range it may be vulnerable to overexploi- Berkes F. 1999. Sacred ecology. Traditional ecological tation, and the magnitude that services associated knowledge and resource management. Philadelphia: Taylor & Francis. with each may be compromised as a result, may be Berkes F, Colding J, Folke C. 2000. Rediscovery of tradi- more precisely answered by drawing on traditional tional ecological knowledge as adaptive management. knowledge to identify the ecosystem services asso- Ecological Appl. 10:1251–1262. ciated with them. By answering such questions, Burkhard B, De Groot R, Costanza R, Seppelt R, Jørgensen efforts can be made to protect the livelihoods of SE, Potschin M. 2012. Solutions for sustaining natural forest-dwelling and dependent peoples, and as a capital and ecosystem services. Ecol Indic. 21:1–6. Byg A, Balslev H. 2004. Factors affecting local knowledge of result the traditional knowledge associated with palms in Nangaritza valley in south-eastern Ecuador. J their landscapes. We suggest that traditional knowl- Ethnobiol. 24:255–278. edge will continue to be critical in dealing with Cámara-Leret R, Paniagua-Zambrana N, Balslev H, Barfod many of the challenges surrounding the sustainable A, Copete JC, Macía MJ. 2014. Ecological community management of tropical forests, sustaining the eco- traits and traditional knowledge shape palm ecosystem services in northwestern South America. For Ecol system services that humanity derives from these Manage. 334:28–42. global resources and indeed protecting the liveli- Cardinale BJ, Duffy JE, Gonzalez A, Hooper DU, Perrings hoods of forest-dwelling and dependent peoples. C, Venail P, Narwani A, Mace GM, Tilman D, Wardle DA, et al. 2012. Biodiversity loss and its impact on humanity. Nature. 486:59–67. Note Castro AJ, Verburg PH, Martín-López B, Garcia-Llorente M, Cabello J, Vaughn CC, López E. 2014. Ecosystem service 1. The term groups of plants is used in this paper to trade-offs from supply to social demand: a landscape-scale refer to plants that were identified to the genus or spatial analysis. Landsc Urban Plan. 132:102–110. species level. In many instances a common name Clarke HD, Funk VA, Hollowell T. 2001. Using checklists and may have referred to more than one plant species collections data to investigate plant diversity. I: a compara- (see examples of this occurrence in Polak 1992; tive checklist of the plant diversity of the Iwokrama forest. Roosmalen 1985; van Andel 2000). Since this analysis Guyana: Botanical Research Institute of Texas. refers to genus and species, it seemed more efficient Colchester M. 1997. Guyana: fragile frontier. to use the term group to refer to them both. Gloucestershire (UK): World Rainforest Movement, Forest Peoples Programme. Colding J. 1998. Analysis of hunting options by the use of Acknowledgments general food taboos. Ecol Modell. 110:5–17. Communities of the North Rupununi. 2006. By-laws for I acknowledge and thank various members of the Moses the management of natural resources by the commu- family, Persaud, Seigmond, Carro, Ricky and Danford, Steve nities of the North Rupununi. Final Report from Andries and Benedict Joseph. At Project Fauna Drs Jose Government of Guyana/United Nations Development Fragoso, Han Overman and Jeff Luzar, and the field teams Programme Sub Project, Promoting Community supported data collection. The Guyana Environmental Participation in the Management of Natural Resources Protection Agency (EPA) in conjunction with the Ministry and Environment. Georgetown (Guyana): Ministry of of Amerindian Affairs granted permission for this research, Amerindian Affairs. while the villages across the Rupununi allowed us to work in Costanza R, d’Arge R, De Groot RS, Farber S, Grasso M, their forests. Hannon B, Limburg K, Naeem S, O’Neill RV, Paruelo J, et al. 1997. The value of the world’s ecosystem services and natural capital. Nature. 387:253–260. Disclosure statement Costa-Silva JH, Lima CR, Silva EJR, Araujo AV, Fraga MCCA, Ribeiro E Ribeiro A, Arruda AC, Lafayette No potential conflict of interest was reported by the authors. SSL, Wanderley AG. 2008. Acute and subacute toxicity of the Carapa guianensis Aublet (Meliaceae) seed oil. J Ethnopharmacol. 116:495–500. Funding Cummings AR. 2013. For logs, for traditional purposes and for food: identification of multiple-use plant species of Funding for this project was made possible by the US Northern Amazonia and an assessment of factors asso- National Science Foundation BE/CNH Grant 0837531. ciated with their distribution (Dissertations e ALL). The project was led by Dr. Jose Fragoso. Paper 17. Available from: http://surface.syr.edu/etd/17. Daily GC. 1997. Nature’s services. Covelo California: Island Press. 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Drawing on traditional knowledge to identify and describe ecosystem services associated with Northern Amazon’s multiple-use plants

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Abstract

INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT, 2016 VOL. 12, NOS. 1–2, 39–56 http://dx.doi.org/10.1080/21513732.2015.1136841 Special Issue: Traditional knowledge for sustainable forest management and provision of ecosystem services Drawing on traditional knowledge to identify and describe ecosystem services associated with Northern Amazon’s multiple-use plants a b Anthony R. Cummings and Jane M. Read Geospatial Information Sciences, School of Economic, Political and Policy Sciences, University of Texas at Dallas, Richardson, TX, USA; Department of Geography, Maxwell School of Citizenship and Public Affairs, Syracuse University, Syracuse, NY, USA ABSTRACT ARTICLE HISTORY Received 30 December 2014 Tropical multiple-use plants provide an array of ecosystem services, including providing food Accepted 23 December 2015 for wildlife, non-timber forest products (NTFPs) and commercial logs. As land-use and land- cover change (LUCC) continues in tropical forests, questions on whether the ecosystem EDITED BY services they provide can be sustained become more pressing. As particular species of plants John Parrotta are targeted for provisioning services, such as commercial logging, the implications for other KEYWORDS ecosystem services are often unknown. In this paper we drew on the traditional knowledge of Traditional knowledge; the Makushi and Wapishiana Amerindians of Southern Guyana, and multiple-use plants to multiple-use plants; gauge the ecosystem services that may be compromised as tropical forests undergo change. ecosystem services; wildlife An inventory of tree and palm species was classified into one or more of four resource-use food; Rupununi; Guyana classes: commercial timber, wildlife food, traditional uses or no known use. Species that intersected more than one resource use were defined as multiple-use species, and occupied four primary classes – wildlife food/commercial timber, commercial timber/traditional uses, wildlife food/traditional uses and wildlife food/commercial timber/traditional uses. Traditional knowledge allowed us to identify plant species and describe how they are used for provision- ing, cultural and supporting ecosystem services and shaping our understanding of the multiple dimensions of ecosystem services associated with a single species. Introduction Albert et al. 2014), examples from tropical forests are still lagging in the literature. The subject of identifying and mapping ecosystem The utilization of traditional knowledge (Posey services across the globe has gained substantial atten- 1992; Parrotta & Trosper 2012) to understand the tion recently. Scholars have established the defini- various types of ecosystem services associated with tions, conceptual framework and problems with tropical plant species and how these may be impacted understanding ecosystem services (e.g. Costanza by land-use and land-cover change (LUCC) and glo- et al. 1997; Daily 1997; Kareiva et al. 2011; bal climatic variability (Parrotta & Agnoletti 2007)is Cardinale et al. 2012; Albert et al. 2014), with various an area requiring pressing attention. The attention to systems emerging for their classification (see traditional knowledge and linkages to ecosystem ser- Millennium Ecosystem Assessment (MEA) 2005; vices is particularly critical as globalization and other Wu 2013). Ecosystem services and ecosystem func- forces reach previously self-sufficient rural areas and tions have been broadly described, while sophisti- result in a loss of traditional knowledge (Parrotta cated methods, models and data have been used to et al. 2009; Trosper et al. 2012). The urgency for better understand select ecosystem services (e.g. understanding the traditional knowledge-ecosystem Burkhard et al. 2012). Despite this progress, however, services connection is further amplified as the conse- the interactions between ecosystem services, as they quence of such loss on livelihoods and cultural and may originate from a single supplier (Furst et al. biological diversity remains poorly understood 2014) such as a tropical plant species, or how such (Parrotta et al. 2009). Furthermore, as Parrotta et al. ecosystem services relate directly to the well-being of (2009) noted, the general public lacks an appreciation human populations, remain under-explored. As the of traditional knowledge, and as a consequence its literature on ecosystem services continues to evolve, erosion may curtail the involvement of local and it seems necessary to understand localized examples indigenous communities in the development of forest of ecosystem services and how they relate to human and land-use policies that impact their livelihoods. As populations critical to the sustainability of such ser- such, scholars have called (e.g. Parrotta et al. 2009; vices. While past studies have covered examples of Sutherland et al. 2014) for greater recognition of the mapping ecosystem services in urban (e.g. Kareiva role indigenous and local people play in the et al. 2011) and developed world settings (e.g. CONTACT Anthony R. Cummings anthony.cummings@utdallas.edu © 2016 Informa UK Limited, trading as Taylor & Francis Group 40 A. R. CUMMINGS AND J. M. READ provisioning services, such as the use of plant parts management of biodiversity (see Mauro & Hardison for medicines (e.g. Forte 1996; van Andel 2000; Khan 2000; and the volume edited by Parrotta & Agnoletti 2007) and merging traditional knowledge with con- et al. 2013) and commercial logging (e.g. Polak 1992), have gained considerable attention. In a similar vein, ventional scientific knowledge (Sutherland et al. cultural services associated with a single species have 2014). In response to such calls, this paper explores the uses associated with plants from the forests of also been examined (see Forte 1996; González-Marín et al. 2012; Cámara-Leret et al. 2014). Of particular Guyana through the lens of forest-related traditional interest to this paper is the fact that individual species knowledge, and the guidance it can provide for forest management and landscape valuation efforts (e.g. of tropical plants are often associated with more than Castro et al. 2014; Paletto et al. 2015; Stara et al. one ecosystem services (similar to the notion of 2015) that have been gaining momentum in the Albert et al. 2014 that there may be multiple demands literature. from a single source for ecosystem services) and our A prominent feature of tropical forests is the host knowledge of such species and their uses is not well of multiple-use plant species they contain that have developed. direct associations with the four dimensions of eco- As tropical forests continue to be converted to system services (provisioning, regulating, supporting support human activities, concerns have arisen over and cultural services) identified by MEA (2005). At whether the ecosystem services they provide can be their most critical level, multiple-use plants provide sustained (e.g. Nepstad et al. 1999;Fearnside 2005). food for wildlife, non-timber forest products (NTFPs) Much of the recent assessments on the impacts of for forest dwelling and dependent peoples, and com- tropical forest area decline on ecosystem services mercial timber (Cummings 2013). The plant uses for have been on implications for carbon sequestration provisioning and supporting ecosystem services may and the hydrological cycle. Although these global- also be superimposed upon the cultural and regulat- level ecosystem services are important, the impacts ing services within a single species. The provisioning on forest-dwelling and dependent peoples, either ecosystem services associated with multiple-use directly through the removal of plants that provide plants, in particular the various demands from differ- important NTFPs or indirectly through impacts on ent groups of forest users on a particular species, have wildlife that they hunt, require equally urgent atten- been gaining attention in the literature (see for exam- tion. The direct impacts on wildlife are critical too, ple Klimas et al. 2007, 2012a, 2012b; Mwavu & as many species disperse seeds and fruits of forest Witkowski 2009). But this attention has often focused plants (Roosmalen 1985), thereby helping maintain on single well-known species (‘trophy species’), such the forest structure. Therefore, identifying and as Carapa guianensis – a species well-known for its studying multiple-use plants and how they are valuable timber and therapeutic oil (Klimas et al. impacted by LUCC is critical. In this regard, the 2007). Moreover, the efforts to probe and describe four parameters to be understood when managing the ecosystem services associated with multiple-use multiple-use plants – an understanding of their plants have made little mention of local-level man- multiplicity of uses and types of resources pro- agers, such as indigenous peoples, in the process of duced, abundance in different forest types, size- studying such species. Therefore, while previous stu- class distribution of populations and life history dies have provided valuable data on the ecology and characteristics (Peters 1996) – are critical in guiding distribution characteristics of multiple-use species our work. We pay particular attention to the first and how they fare under conditions of commercial two parameters suggested by Peters using a sample logging and NTFPs extraction, they introduce ques- of plants from Northern Amazonia. We argue that tions of how many more species exist, and the best there are many tropical multiple-use plant species, means for obtaining knowledge on the ecosystem and that when the knowledge of scientific experts is services they provide. combined with traditional knowledge a more com- In addition to only the prominent multiple-use prehensive appreciation of ecosystem services with species gaining attention in the literature, the diverse which they are associated can be derived. Such range of ecosystem services associated with a species knowledge can then be used for planning the sus- has also often been ignored. Single facet of plant uses, tainability of ecosystem services associated with for example their role in providing wildlife food, individual species and the landscapes in which NTFPs or commercial timber, separately, has been they are found (see Parrotta et al. 2009). Scholars commonly reported (see for example Peters 1996; such as Forte (1996), Parrotta and Trosper (2012) Klimas et al. 2007; Mwavu & Witkowski 2009). and Posey (1992) have drawn on traditional knowl- Scholars have examined the supporting ecosystem edge to various resource management-related ends. services associated with tropical plants (trees and Forte (1996), for example, utilized the knowledge of palms) through their fruits and seeds (e.g. Makushi women of the Rupununi, Southern Roosmalen 1985; Silvius & Fragoso 2003), while the Guyana, to describe how plants have been used for INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 41 processes that occur at temporal or spatial scales not medicinal purposes. Parrotta and Trosper (2012) easily observed by conventional scientific research. have highlighted the important knowledge that tra- ditional societies – indigenous and local commu- Peloquin and Berkes (2009) pointed out that indigen- ous and rural groups appeared to have developed nities – have used to manage their forests in order practices that allow them to live with complexity to sustain their livelihoods and cultures. The rapid changes in forested regions endanger not only indi- that are often missed by Western science-based socie- ties who seek to simplify ecosystems. genous and local communities that live within such As a consequence of the recognition of the value forests, but also the important ecosystem services that these forests may hold even before we are fully traditional knowledge has for humanity, scholars (e.g. Ens et al. 2015; Whyte et al. 2015) have been suggest- aware of them. Traditional knowledge of how indi- ing measures to incorporate traditional knowledge genous communities have used forests is therefore a critical step in understanding the impacts of LUCC into sustainability science. In recognition of the role within the context of sustainable forest management traditional knowledge must play in ecosystem con- servation and the protection of local livelihoods, this and preserving traditional lifestyles. paper seeks to bring the importance of traditional knowledge in helping shape our understanding of The value of traditional knowledge forests and their ecosystem services in landscapes that are vulnerable to change. In this regard our The value of traditional knowledge to ecosystem con- work will help inform modern-day approaches to servation, sustainable resource management and local sustainable forest management on how traditional livelihoods has been stressed by a number of scholars knowledge should be incorporated to protect the over the years (e.g Berkes et al. 2000; Mauro & livelihoods of local and forest-dependent peoples. Hardison 2000; Turner & Cocksedge 2001; LaRochelle & Berkes 2003; Houde 2007; Parrotta & Trosper 2012; de Freitas et al. 2015). Traditional Goals and objectives knowledge, which consists of the past and present beliefs, traditions, practices and views developed by With the foregoing discussion in mind, the goals of indigenous and local communities (Huntington this paper are twofold. First, drawing on the tradi- 2000), is a key to accessing, using and managing tional knowledge of the Makushi and Wapishiana ecosystem services (Cámara-Leret et al. 2014). Amerindians, we identified multiple-use plants spe- Cámara-Leret et al. (2014) further noted that tradi- cies of the Rupununi, Southern Guyana, landscape tional knowledge is diverse, dynamic and place-spe- (Figure 1). Secondly, we determined which uses, or cific, while Byg and Balslev (2004) noted that types of ecosystem services, are associated with each traditional knowledge can vary among individuals species. As a clear departure from previous studies, and communities. Phillips et al. (1994) posited that our analysis combined the literature on how plants the benefits local communities obtain from their sur- are used in Guyana with traditional knowledge of the rounding ecosystems are determined by the levels of Makushi and Wapishiana Amerindians to identify traditional knowledge about the classification, ecol- multiple-use plants sampled from the Rupununi. ogy, usefulness and management of natural resources. Our main objective is to demonstrate to policymakers Turner et al. (2000) noted that traditional knowledge the role traditional knowledge can play in sustainable is ‘fundamentally important in the management of forest management, with particular emphasis on con- local resources, in the husbanding of the world’s serving ecosystem services and protecting local liveli- biodiversity, and in providing locally valid models hoods. We focused primarily on species of plants that for sustainable living.’ Interest in traditional knowl- intersect the three main resource uses: commercial edge has been growing primarily as scholars recog- logging, traditional uses (NTFPs) and wildlife food, as nize the contribution it can make to the conservation the intersection of these uses in tropical forests will of biodiversity (Gadgil et al. 1993; Berkes et al. 2000), determine which species and the ecosystem services rare species (Colding 1998), protected areas with which they are associated will be vulnerable. (Johannes 1998) and to sustainable resource use in Where a single species had more than one use we general (Berkes 1999). Berkes et al. (2000) further described it as multiple-use (see Cummings 2013). noted that conservation biologists, ecological anthro- We used multiple-use classes to explore how plants pologists, ethnobiologists, other scholars and the are used and how these uses may be compromised by pharmaceutical industry, all share an interest in tra- changing landscapes. In this regard, this paper goes ditional knowledge for scientific, social or economic beyond previous efforts that provided a cursory view reasons. More recently, de Freitas et al. (2015) noted of the important resource-use intersections associated that traditional knowledge can provide insights into with an individual plant species (e.g. Klimas et al. studying ecological systems that are important to 2007, 2012a, 2012b), and offers a comprehensive local populations, and understanding ecological description of the ecosystem services associated with 42 A. R. CUMMINGS AND J. M. READ Figure 1. Study area. each species. Our work therefore contributes to the of traditional knowledge in sustainable forest man- ongoing discussion on valuing landscapes for the agement and valuation processes of tropical land- ecosystem services they provide. scapes and the impacts of deforestation beyond the We make the argument that through identifying current carbon sequestration focus. multiple-use plants within tropical forests, and utiliz- ing traditional knowledge to identify such plants, national-level forest managers will be better placed Methods to understand the impacts of forest-cover change on Study area forest-dwelling and dependent peoples, and be able to incorporate traditional knowledge in sustainable for- This study was completed in the tropical forest and est management practices. Our work contributes to savannah biome of the Rupununi, Southern Guyana. the call by Parrotta et al. (2009) for documenting The Rupununi study area lies between 0°50ʹ – 40° 49ʹ N traditional forest-related knowledge (TFRK) in colla- and 56°54ʹ – 59°55ʹ W and is approximately 350 km boration with the holders and users of such knowl- southwest of the capital of Guyana, Georgetown edge. As a consequence, our work establishes the (Figure 1). The study area is ecologically important, basis for ecological studies on the health and status supporting a high diversity of ground-living mammals, of plants. By identifying the threats that exist to the frugivorous primates and birds (Montambault & Missa ecosystem services associated with tropical multiple- 2002; Watkins et al. 2011). Of the ground-living mam- use plants, we hope to contribute to shaping our mals, reptiles and birds, more than 107 species are understanding of ecosystem services and the value hunted by the native Amerindians for food, with that is lost as tropical forest areas decline. Through eight species (Paca or labba (Cuniculus paca), agouti the process of combining traditional knowledge with (Dasyprocta leporina), white-lipped peccary (Tayassu conventional science, this study contributes to the pecari), collared peccary (Pecari tajacu), red brocket process of allowing for more widespread inclusion deer (Mazama americana), white-tailed deer INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 43 controls), eight 4-kilometer-long transects were (Odocoileus virginianus), nine-banded armadillo established for sampling. (Dasypus novemcinctus) and red footed tortoise (Chelonoidis carbonaria)) comprising 69% of the A two-person team comprising an indigenous person with expertise in identifying plant species hunted animals (Read et al. 2010). Among these top (tree spotter) and a recorder inventoried all trees hunted species, all but the nine-banded armadillo have a strong reliance on the fruits of trees and palms in the greater than 25 cm diameter-at-breast-height (dbh) on transects. In cases where trees had an irregularity area for food (Roosmalen 1985). The area also supports or buttress at breast-height, diameter measurements the Cariban-speaking Makushi and Arawakan-speak- ing Wapishiana Amerindians (Colchester 1997)who were taken above these (Polak 1992). A minimum tree size of 25 cm dbh was chosen because tradi- have lived in the Rupununi for centuries (Forte 1996; tional knowledge suggested that most trees that Plew 2005). The Rupununi Amerindians depend on swidden provide food for wildlife produce fruits consistently agriculture, with hunting and fishing being major and vigorously at around this size. Sampling from 25 cm dbh also allowed for trees that are of interest parts of their subsistence livelihood practices. In recent times the Rupununi has been more accessible from to all resource users to be included. Commercial coastal Guyana and Brazil, meaning that the area’s species in Guyana are generally logged at stem historically protected natural resource base is becoming diameters >40 cm, while for traditional purposes, attractive for resource extraction activities, including including fuelwood, trees are harvested at various commercial logging and gold mining. Even though sizes depending on their intended uses. such changes could present communities with alterna- Furthermore, sampling at this minimum size tives to their subsistence livelihood activities allowed for plant identification errors to be reduced, (Communities of the North Rupununi 2006), it could as it assumed that tree spotters would be better able also have implications for overall forest equilibrium to identify larger trees than smaller ones, thereby (Uhl et al. 1991; Martini et al. 1994), impacting the resulting in a more accurate inventory. In addition survival of wildlife populations hunted by Amerindians to trees, mature palms (dbh of palms was not mea- and multiple-use plant species. Given the diversity of sured), with maturity defined by evidence that a ecosystem services that the forests of the Rupununi palm had borne fruits in the past (fruits, seeds or provide, and the fact that the area’sindigenouspopula- seedlings on the forest floor, empty bunches on tions retain knowledge of how plants are used, this area stems on the forest floor), were also inventoried. provided an excellent opportunity to study the various A 10-meter belt transect (see Figure 2) was used ways in which tropical plants are used, and begin to for inventorying all trees and palms within each understand how LUCC may impact these ecosystem transect’s width. For all plants that met the sampling services. In addition, scholars based at the Iwokrama criteria, inventory teams recorded: distance a plant International Center for Rainforest Conservation and Development (Iwokrama) have studied the ways in which indigenous peoples use plants in their forests (e.g. Forte 1996), providing a foundation for under- standing ecosystem services in the area. Sampling procedure and data collection Data on trees and palms were collected from 14 study sites distributed across the Rupununi land- scape (Figure 1) during the period July–December 2008. Study sites were a subset of communities and control areas participating in a larger project (see Luzar et al. 2011;Readetal. 2010;for details).As described in Read et al. (2010), study sites were selected based on their spatial distribution through- out the study area, their location with respect to vegetation types and topographic characteristics, proximity to other communities and willingness of community members to participate in the project. Figure 2. Schematic representation of the 10-meter belt The subset of study sites were the predominantly transect used for completing the inventory of trees and forested sites from those sampled by Luzar et al. palms. This schematic shows a 50-meter transect segment, (2011) and Read et al. (2010); at each study site (12 the right and left side of a transect’s imaginary centerline, Amerindian communities and two unhunted and representations of trees (circles). 44 A. R. CUMMINGS AND J. M. READ species. Wildlife and their food sources were sepa- was found along a transect; direction a plant was rated based on their importance to the Amerindian located off a transect’s imaginary centerline – right or left; distance away from a transect’s imaginary diet; hunted species were separated from unhunted species, such as frugivorous primates. centerline (offset), dbh (for trees); and local name. Distance markers were placed at 25 m intervals along transects to aid in estimating distances along a trans- Results ect. Where the local name of a tree was not known at the time of sampling, plant parts including leaves, Multiple-use species barks and fruits were collected and used to consult Our results showed that multiple-use plant groups the literature, elders and hunters to identify that tree. were the most important part of our sample, account- Trees were labeled as ‘unidentified’ in cases where ing for more than 81% of the 33,457 plants sampled. after consulting these sources a positive identification Plants were placed into one of four multiple-use was not possible. In essence, traditional knowledge of classes each describing the ecosystem services that indigenous tree spotters was utilized to identify each the groups within that class provide. The multiple- plant that met our sampling criteria. use classes were derived from the intersections of the three major resource uses: commercial timber, wild- Data analysis life food and traditional uses, and the resulting classes were used to describe the nature of plant groups. The Botanical names and resource-use classes four multiple-use classes are: The common name attached to each plant in the inven- tory was translated to a botanical name, where possible, (1) Commercial timber and wildlife food using the literature (including Roosmalen 1985;Polak (2) Commercial timber and traditional uses 1992;Forte 1996; van Andel 2000; Iwokrama 2008), and (3) Wildlife food and traditional uses in consultation with other researchers. Each plant was (4) Wildlife food, commercial timber and tradi- assigned to one or more of four major resource-use tional uses classes, based on traditional knowledge and the litera- ture of their uses in the Rupununi or elsewhere in Wildlife food/commercial timber/traditional uses Guyana for traditional and other purposes: Forty-one groups of plants were classified as wildlife food/commercial timber/traditional uses. Seventeen of (1) Commercial timber: genus or species of tree these species provide food for hunted wildlife (Table 1) actively logged for commercial timber in Guyana. while the remaining groups provide food for birds and (2) Wildlife food: genus or species of tree or palm frugivorous primates. It is possible that this class could that provide food for wildlife (hunted or not). In be expanded as in many instances one common name a few cases, data on the dispersal mode of a plant referred to more than one species. For instance, in family were used to determine animals that are Guyana crabwood generally refers to Carapa guianen- likely associated with a species in the inventory. sis, but the literature (e.g. Polak 1992; Fisch et al. 1995; (3) Traditional uses: genus or species of tree or palm Sanogo & Sacande 2007) and traditional knowledge used by Amerindians for medicinal purposes, suggested that C. procera DC may also be present in traditional weaponry, boat crafting, traditional the study area. As Fisch et al. (1995) noted the diffi- utensils, home construction, thatching, fuelwood, culty in separating these two species under normal making beverages, food, including utilizing same field conditions, local tree spotters may have refer- for economic purposes and other purposes. enced both species under the common name crab- (4) No known uses: at the time of analysis no wood. A similar situation may be applied to resource uses could be determined for this Kautaballi, which refers at least to two species, genus or species. Licania alba (Bernouli) Cuatrec and L. majuscula Based on this classification, multiple-use species were Harvey ex Gomont (Roosmalen 1985;Polak 1992). those that intersected resource-use classes 1–3. In We made no attempt to differentiate between these addition to hunted wildlife, traditional knowledge species during data collection, as the provision services and the literature (e.g. Roosmalen 1985) provided associated with them appear to transcend species. information on other mammals and birds, including The groups of plants in this class varied in terms of frugivorous primates that depend on the fruits of their range of uses, either as wildlife food sources or plants of the Rupununi for food. The resulting data- as traditional uses, while some may be more preferred base of plants included a list of tree and palm species, over others for commercial timber. In the case of a resource-use class for each species, the purpose for Carapa genus, this group has a limited range of uses which the species is used and, where appropriate, the as wildlife food, with only the hunted rodent species species of animals that depend on the fruits of that (Table 1) that disperse their seeds linked to their INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 45 Table 1. Plant groups in the commercial timber/traditional uses/wildlife food multiple-use class, and the species of wildlife associated with their fruits and seeds. Family/Species Common name Eaten by Bombacaceae Catostemma commune Sandwith Baromalli agouti, deer, tapir, howler* and spider monkeys*, bats* Pachira aquatica Aubl. Kanahia genus synzoochorous scatter-hoarding rodents, such as agoutis and acouchis Boraginaceae Cordia alliodora (Ruiz & Pav.) Oken Freijor genus endozoochorous (birds, monkeys) Burseraceae Protium decandrum (Aubl.) Marchand Kurokai agouti, marudi, powis (black curassow), deer, capuchin* Caryocaraceae Caryocar nuciferum L. Sawari nut tapir, deer, labba, spider monkey* Chrysobalanaceae Licania alba (Bernouli) Kautaballi agouti Cuatrec. and L. majuscula Harvey ex Gomont Parinari campestris Aubl. Burada macaws*, saki monkeys* Flacourtiaceae Laetia procera (Poepp.) Eichl. Warakairo endozoochorous: several birds and most primates Goupiaceae Goupia glabra Aubl. Kabukalli genus endozoochorous(howler and spider monkeys, birds) Guttiferae Calophyllum brasiliense Camb. Cassava mama genus endozoochorous (birds, monkeys) 1,2 Symphonia coccinea (Aubl.) Oken Manniballi family endozoochorous or synzoochorous (birds, monkeys) 1,2 Symphonia globulifera L.f. Manni family endozoochorous or synzoochorous(birds, monkeys) Humiriaceae Humiria balsamifera (Aubl.) A. St.-Hil. Tauroniro deer, peccaries, powis, marudi, tortoise, foxes*, macaws*, other birds* and monkeys* Lauraceae Licaria canella (Meisn.) Kosterm. Brown Silverballi family endozoochorous (specialized frugivores: toucans, cotinga, guans, spider and howler monkeys) Chlorocardium rodiei (Schomb.) Rohwer, H.G.Richt. Greenheart agouti, labba, and white lipped peccaries & van der Werff Ocotea canaliculata (Rich.) Mez; Aniba hypoglauca Silverballi family endozoochorous (specialized frugivores: toucans, cotinga, guans, Sandw. spider and howler monkeys) Lecythidaceae Eschweilera alata A.C. Smith Guava Skin monkeys* Kakaralli Lecythis corrugata Poit. Wina Kakaralli monkeys* Eschweilera decolorans Sandw. Smooth-Leaf labba, several monkeys* Kakaralli Eschweilera sagotiana Miers Common Black monkeys, birds* Kakaralli Eschweilera wachenheimii (Benoist) Sandw. Fine Leaf Kakaralli monkeys* Lecythis cf. chartacea Berg Monkey Pot – labba small Lecythis zabucajo Aublet Monkey Pot labba Leguminosae (Caesalpiniaceae) Hymenaea courbaril L. Locust agouti, labba, tapir, peccaries, monkeys* (saki, capuchin) Mora excelsa Benth. Mora peccaries, deer and tapir eat newly germinated seedlings Peltogyne venosa (Vahl) Benth. Purpleheart genus endozoochorous (especially spider monkeys) Peltogyne paniculata Benth. Purpleheart saka genus endozoochorous (especially spider monkeys) Leguminosae (Mimosaceae) Inga alba (Sw.) Willd. Maporakon agouti, labba, tapir, deer, powis, marudi, peccaries, parrots*, capuchin* and spider* monkeys Abarema jupunba (Willd.) Britton & Killip Huruasa endozoochorous spider monkeys Leguminosae (Papilionoideae) Dipteryx odorata (Aubl.) Willd. Tonka bean toucans* and other birds* Ormosia coccinea (Aubl.) Jackson Barakaro peccaries Swartzia leiocalycina Benth. Wamara genus zoochorous (bats, rodents, monkeys) Meliaceae Carapa guianensis Aubl., C. procera DC Crabwood agouti, labba, peccaries Moraceae Bagassa guianensis Aubl. Cowwood tapir, deer, peccaries, labba, agouti Brosimum guianense (Aubl.) Huber Letterwood endozoochorous spider monkeys and birds Myristicaceae Virola surinamensis (Rol.) Warb. Swamp Dalli monkeys, toucans and other birds Sapotaceae Manilkara bidentata (A.DC.) Chev. Bulletwood peccaries, tapir, deer, tortoise, labba, agouti, spider* and howler monkeys*, macaws* Pouteria cuspidata (A.DC.) Baehni Kokoritiballi agouti, red brocket deer, labba, monkeys* Pouteria guianensis Aubl. Asepoko agouti, red brocket deer, labba, spider* and howler monkeys*, capuchins* Simaroubaceae Quassia spp. Angelina Rock genus endozoochorous (spider and howler monkeys) Quassia simarouba L.f. Simarupa genus endozoochorous (spider and howler monkeys) *species not hunted – data from traditional knowledge and Roosmalen (1985) data from Roosmalen (1985) data from Forget and Hammond (2005) **the scientific names of hunted wildlife species are provided in the text. Species listed above are: howler monkey (Alouatta seniculus); spider monkey (Ateles paniscus); white saki (Pithecia pithecia); bats (Artibeus lituratus among other species); black saki (Chiropotes chiropotes); capuchin (Cebus apella, Cebus olivaceus); toucans (Ramphastos toco, Ramphastos vitellinus); and macaws (Ara ararauna, Ara macao, Ara nobilis). 46 A. R. CUMMINGS AND J. M. READ traditional knowledge suggested that the fruits of fruits (Forget et al. 1999; Forget & Jansen 2006). The Ormosia coccinea (Aubl.) Jackson are eaten by pecc- traditional value of Carapa spp. is, however, in little doubt; the seeds are widely utilized throughout aries, but Roosmalen (1985) noted that the genus is endozoochorous with birds mainly dispersing these Guyana and Amazonia (van Andel 2000; Shanley & seeds. In the case of Licania alba and L. majuscula Luz 2003; Sullivan & O’Regan 2003; Plowden 2004; Klimas et al. 2007; Costa-Silva et al. 2008) for its (Kautaballi) van Andel (2000) noted that the name given to L. alba by the Warrau people of the North valuable oil (see also Klimas et al. 2007). Similarly, West District (NWD) translates to ‘agouti food tree’ as Chlorocardium rodiei Schomb.) Rohwer, H.G.Richt. & van der Werff (greenheart) has only been asso- these rodents feed on the seeds of the tree. The tradi- tional uses of Eschweilera alata A.C. Smith (house ciated with hunted scatter-hoarding rodents posts), E. sagotiana (head straps and lashing materials, (Hammond et al. 1999) and peccaries (traditional knowledge). However, like the species in the Carapa house construction, and seeds used as bait in bird genus, C. rodiei is a primary target for commercial traps), E. wachenheimii (bark used for head straps and lashing materials and house construction), L. corrugata logging in Guyana (Zagt 1997; Ter Steege et al. 2002). Further, C. rodiei is listed by the International Union (house construction), Virola surinamensis (Rol.) Warb. for the Conservation of Nature and Natural Resource (used for making Arawak musical instruments and the (IUCN) as a threatened species (IUCN 2012). The sap is used for treating thrush in babies and also gargled traditional uses of C. rodiei are primarily medicinal, for tonsillitis and toothaches), Pouteria cuspidate (house with the seeds being used to treat parasitic worms construction) and Lecythis cf. chartacea Berg (lashing and skin diseases (Forte 1996). material) were noted by van Andel (2000)for their In contrast to the limited range of uses associated utility in the NWD. It seems likely that these species with plants in the Carapa genus as wildlife food are similarly utilized in the Rupununi, but this was not sources, Manilkara bidentata (A.DC.) Chev. may be revealed during data collection or through the literature described as a universal wildlife food source. It was (e.g. Forte 1996). suggested, in particular by traditional knowledge, that Sixteen groups in this class were included based solely all the hunted wildlife species depend on the fruits of on the data provided by Roosmalen (1985)ontheir M. bidentata. Furthermore, M. bidentata has long interaction with wildlife. Abarema jupunba (Willd.) been recognized as a commercial timber species Britton & Killip, Anibaspp.,Brosimumguianense (Polak 1992). As an NTFP, Amerindians have used (Aubl.) Huber, Calophyllum spp., Cordia alliodora balata, the gum derived from M. bidentata, to make (Ruiz & Pav.) Oken, Goupia glabra Aubl., Licaria canella utensils for storing water and beverages, and the (Meisn.) Kosterm. Laetia procera (Poepp.) Eichl, Ocotea fruits of M. bidentata are eaten (Forte 1996). spp Quassia simarouba, Pachira aquatica Au bl., Indeed, there was once a thriving trade in the sap of Peltogyne paniculata Benth., Peltogyne venosa (Vahl) M. bidentata from the Rupununi. Currently the Benth, Swartzia leiocalycina Benth. and Quassia spp. dependence on this species as an NTFP has waned, were listed at the genus or species level as being either with its consumption generally restricted to making synzoochorous (animals that bury or store seeds) or figurines of animals and common articles found in endozoochorous (dispersal of seeds after passing the Rupununi for the tourism market. through the gut of an animal). Among these, only However, in addition to the most abundant groups Pachira spp. was listed as being dispersed by hunted – Catostemma commune Sandwith, Parinari campes- wildlife (agoutis and acouchis). In two exceptional tris Aubl. Mora excelsa Benth. and Protium decan- cases, Roosmalen (1985)noted Symphonia coccinea drum (Aubl.) Marchand – being Aubl. and Symphonia globulifera L.f. to be dispersed at included in this class (Table 1), there were also groups with low abun- the family level synzoochorously or endozoochorously, but Forget and Hammond (2005) indicated that this dance. For example, Caryocar nuciferum L. fruits are genus is an important food source for the red brocket eaten throughout forested Guyana, with fruits and trees observed at various Rupununi homes during deer. The other groups are primarily of interest to fru- givorous primates and birds. Traditionally, Pachira spp. data collection, although trees were only observed at two study sites – Villages 18 and 23. and L. procera are utilized as fuelwood, while C. allio- In some instances the traditional uses of groups in dora, Calophyllum spp., G. glabra, Peltogyne spp.and Quassia spp. are used in house construction and for this class were well established, but not so their value as wildlife food. M. excelsa illustrates this point as tradi- making canoes. The bark of G. glabra has medicinal tional knowledge suggested that the fruits and seeds of applications, with the boiled bark being used to treat eczema andtheleaves andboiledbarkareusedforthe M. excelsa are eaten by paca and peccaries and that deer, tapir and peccaries browse on the newly germinated treatment of chickenpox and to relieve toothache (van seedlings. However, the literature, in particular Andel 2000). P. venosa is used in house construction, A. jupunba is used as a soap substitute (van Andel 2000) Roosmalen (1985), noted that M. excelsa is dispersed by water, that is, they are hydrochorous. Similarly, and Brosimum spp. isusedtomakebowsand walking INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 47 other genus including Cordia are also referred to by sticks. S. coccinea and S. globulifera are used for making this common name. This common name was therefore canoes in both the Rupununi and the NWD. Furthermore, van Andel (2000)pointedoutthatasa a point of uncertainty in our analysis for identification, but all of these species are used traditionally, primarily in medicinal product, the boiled bark of S. coccinea and S. house construction and for making paddles (Forte globulifera is used to treat abscesses and diarrhea and thrush in babies. The traditional uses of Licaria canella, 1996). Forte (1996)points out toothat Swartzia leioca- lycina is a much-favored fuelwood, while Quassia simar- Ocotea spp., Aniba spp., Swartzia leiocalycina and ouba is used by the Makushi in house construction. Quassia simarouba were noted by Forte (1996)based on the knowledge of the Makushi people (primarily women) of the North Rupununi. Ocotea spp., Aniba Commercial timber/wildlife food spp., Swartzia leiocalycina and Quassia simarouba are The commercial timber/wildlife food class contained all listed by Polak (1992) as major timber species. Licaria 20 groups of plants. These plants, even though they cannella is used in house construction, and the bark is provide food for hunted wildlife, are more important used medicinally and for tanning leather. Members of for the frugivorous primate population of the the Ocotea genus including Ocotea canaliculata (Rich.) Rupununi and birds, with only seven species linked Mez, (Iwokrama 2008) along with members of the to hunted wildlife (Table 2). Sixteen groups of plants Aniba genus including Aniba hypoglauca Sandw. are were included in this class based primarily on data commonly referred to as Silverballi, but members of from Roosmalen (1985) and are dispersed mostly by Table 2. Plant groups in the commercial timber and wildlife food class and the species of wildlife associated with their fruits and seeds. The relationship between these plants and hunted wildlife is highlighted. Eaten by Hunted Family/Species Common name Wildlife? Eaten by Bombacaceae Catostemma fragrans Benth. Sand Baromalli Yes agouti, red and gray brocket deer, tapir, bats, howler and spider monkeys* Burseraceae Trattinnickia rhoifolia Willd. Ulu No genus endozoochorous (especially birds) Guttiferae Calophyllum lucidum Kuruhara No genus endozoochorous Benth. Platonia insignis Mart. Kaslego Yes monkeys, peccaries* Lauraceae Aniba hypoglauca Sandw. Yellow Silverballi No family endozoochorous (specialized frugivores: toucans, cotinga, guans, spider and howler monkeys) Ocotea canaliculata (Rich.) Mez White Silverballi No family endozoochorous (specialized frugivores: toucans, cotinga, guans, spider and howler monkeys) Ocotea rubra Mez Determa No family endozoochorous (specialized frugivores: toucans, cotinga, guans, spider and howler monkeys) Ocotea splendens (Meisn.) Mez. Kereti Silvaballi No family endozoochorous (specialized frugivores: toucans, cotinga, guans, spider and howler monkeys) Lecythidaceae Eschweilera chartacea (O.Berg) Broad Leaf Kakaralli Yes parrots, macaws*; genus synzoochory: (scatter-hoarding rodents) Eyma and several monkey species Eschweilera pedicellata (Rich.) White Skin Kakaralli Yes genus synzoochorous: (scatter-hoarding rodents) and several monkey species Eschweilera subglandulosa (Steud. Black Kakaralli Yes macaws, parrots*; genus synzoochory: (scatter-hoarding rodents) ex O.Berg) Miers and several monkey species Lecythis confertiflora (A.C. Sm.) Werimiri Kakaralli No genus endozoochorous (spider monkeys) and synzoochorous (bats) S.A. Mori Leguminosae (Caesalpiniaceae) Mora gonggrijpii (Kleinhoonte) Morabukea Yes peccaries, deer and tapir eat newly germinated seedlings* Sandwith Leguminosae (Papilionoideae) Ormosia coutinhoi Ducke Korokororo No genus endozoochorous (birds) Swartzia benthamiana Miq. Itikiboroballi No genus zoochorous (bats, rodents, monkeys) Myristicaceae genus zoochorous (bats, rodents, monkeys) Iryanthera lancifolia Ducke Kerikowa No genus endozoochorous (specialized frugivores, such as toucans, guans, cotingas, monkeys) Iryanthera macrophylla (Benth.) Black No genus endozoochorous (specialized frugivores, such as toucans, Warb. Kerikowa guans, cotingas, monkeys) Virola michelii Heckel Hill Dalli No genus endozoochorous (specialized frugivores, such as toucans, guans, cotingas, monkeys) Sapotaceae Pouteria speciosa (Ducke) Baehni Suya Yes labba, agouti, peccaries, tapir* Verbenaceae Vitex stahelii Moldenke Hakiaballi No genus endozoochorous (spider and howler monkeys) data from Roosmalen (1985) data from Forget and Hammond (2005) *data from Rupununi traditional knowledge 48 A. R. CUMMINGS AND J. M. READ frugivorous primates, either endozoochorously or Eperua falcata Aubl., Tabebuia insignis (Miq.) synzoochorously. In a few instances, scatter-hoarding Sandwith, Tabebuia serratifolia (Vahl) G. Nicholson rodents also played a role in dispersing these seeds. and Pterocarpus rohrii Vahl; Polak 1992). It was also Furthermore, Forget and Hammond (2005) sug- suggested by traditional knowledge that Thyrsodium gested, based on data provided by Gayot (2000) and guianensis Sagot is a timber species. However, the Gayot et al. (2004), that Swartzia benthamiana Miq. data suggested that the traditional uses of the species is a food source for red and gray brocket deer. Forget in this multiple-use class are not universal. Some species are only used for traditional purposes in the and Hammond (2005) also suggested that the fruits of most of the groups in the Leguminosae family, a Rupununi, while others are only used in the NWD number of which are included in this class, are eaten and coastal Guyana (Table 3). The fact that the tradi- tional uses of these plants is not universal indicates by the wildlife species hunted in the Rupununi. The fact that most of the groups of plants in this class are the cultural value that different groups of people give of more importance to frugivorous primates than to plants, and is an area that requires further analysis in Guyana and where these species are found across hunted wildlife is critical, as primates play important ecological functions in maintaining the forest struc- the Neotropics. ture (Roosmalen 1985; Forget & Hammond 2005). The interactions between plants and animals is an Wildlife food/traditional uses area that requires further research as Forget and There were 39 groups of plants within the wi ldlife food/ Hammond (2005) suggested that the diet of the pri- traditional uses class (Table 4). The palms (eight spe- mates and ungulates (deer, tapir and peccaries) needs cies) dominated this class of multiple-use plants, and to be better understood. this indicates the critical role palms play in traditional practices and NTFPs extraction from tropical forests. Commercial timber/traditional uses The fruits of palm are eaten by people and the wildlife The commercial timber/traditional uses class (Table 3) and the leaves of many are harvested for thatching roofs and basketry making (Forte 1996). But like traditional was constituted of 11 groups of plants. Of these, seven groups are major timber species (Clathrotropis uses of plants in the commercial timber and traditional brachypetala (Tul.) Kleinhoonte, Clathrotropis macro- uses, not all uses appear universal. For example, the fruits of Oenocarpus bacaba Mart. and Oenocarpus carpa Ducke, Diplotropis purpurea (Rich.) Amshoff, Table 3. Plant groups in the commercial timber and traditional uses class, highlighting their traditional uses and the region of Guyana that these uses have been attributed. Family/Species Common name Traditional Uses Region Bignoniaceae Tabebuia serratifolia Ironwood used in house construction, to make 8–9-foot-long posts; bark used to All (Vahl) G. prevent scabies; young leaves used to preserve teeth Nicholson Tabebuia insignis White cedar house construction Rupununi (Miq.) Sandwith Leguminosae (Caesalpiniaceae) Eperua falcata Aubl. Wallaba used to make posts, shingles and bows; bark used medicinally to treat Rupununi internal injuries Leguminosae (Papilionoideae) Centrolobium Paurine used in house construction, to make posts; also paddles and good Village 19 has a paraense Tul. fuelwood conservation area to protect this species Clathrotropis Aromata – fine leaf bark scraped and applied to leishmaniasis parasite wounds to kill the NWD brachypetala (Tul.) parasite. Drinking the sap from the bark could be used to treat the bite Kleinhoonte from the water coral snake: Helicops angulata Clathrotropis Aromata bark scraped and applied to leishmaniasis parasite wounds to kill the NWD macrocarpa Ducke parasite. Drinking the sap from the bark could be used to treat the bite from the water coral snake: Helicops angulata Diplotropis purpurea Tatabu used to make canoes NWD (Rich.) Amshoff Pterocarpus rohrii Hill Corkwood used for making canoes, paddles, carved toys All Vahl Pterocarpus spp. Corkwood used for making canoes, paddles, carved toys All Meliaceae Tabebuia stenocalyx Water Cedar boards used in house construction, to make shingles, dugout canoes Rupununi Sprague & Stapf. Sapindaceae Thyrsodium Sand Mora fish poison NWD guianensis Sagot van Andel (2000), NWD data based on van Andel (2000), other data from Rupununi traditional knowledge and Forte (1996). INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 49 Table 4. Multiple-use tree species in the class wildlife food and traditional uses. Family/Species Common name Eaten by Traditional uses Region Anacardiaceae Anacardium giganteum Hancock ex Hubudi/wild cashew tapir, tortoise, deer, powis, peccaries, monkeys (capuchin, fruits eaten and used for making beverages NWD Engl. spider) Mangifera indica L. Mango domesticated animals, wildlife fruits eaten All Spondias mombin L. Plum tapir, agouti, labba, tortoises, monkeys (squirrel, howler, fruits eaten, various beverages, seeds used as fish bait, soot of roasted seeds used to treat All spider, capuchin), toucans** wounds from vampire bats, used to treat hemorrhaging post childbirth, bark and leaves used to treat sores and tea from bark used for colds Tapirira guianensis Aubl. Duka hanaqua, marudi, some monkeys, foxes** fuelwood Rupununi Annonaceae Duguetia calycina Benoist . Yari Yari genus zoochorous (monkeys, scatter-hoarders) bows, fishing rods NWD Unonopsis glaucopetala R.E.Fr. Rough Skin Arara endozoochorous (spider monkeys, cotingas) treating the bite of labaria (Bothrops asper) or fer-de-lance NWD Xylopia cayennensis Maas Black Maho genus endozoochorous (monkeys, birds) astringent tea for diarrhea; firewood; bark used for warashis straps; decoction also drunk by NWD; pregnant women as an aid to eventual easy delivery Rupununi Xylopia cf. surinamensis R.E. Fr. Kuyama* genus endozoochorous (monkeys, birds) astringent tea for diarrhea; firewood NWD; Rupununi Apocynaceae Geissospermum argenteum Woodson Sinkona tortoises; genus endozoochorous(monkeys, birds) bark boiled for treating malaria – source of quinine All Burseraceae Protium guianense (Aubl.) Marchand. Haiawa genus endozoochorous (toucans, cotingas, guans, spider gum used to cure diarrhea; mixed with cow fat and used to caulk boats in the absence of Rupununi monkeys, howler monkeys, tortoises, kinkajous, karamani gum (maitakin), gum used to anchor grater’s teeth firmly on board; burned as incense opossums) under the hammock of a sick person; fruits eaten Boraginaceae Cordia tetrandra Aubl. Grandma Cherry birds and monkeys** fruits used as glue for paper and making kites NWD Chrysobalanaceae Chrysobalanus icaco L. Fatpoke birds** fruits eaten All Ebenaceae Diospyros ierensis Britton Barabara tapir and birds fuelwood Rupununi Guttiferae Clusia grandiflora Splitg. Kufa genus endozoochorous birds sometimes monkeys used in the furniture industry All Tovomita schomburgkii Planch. & Awasokule genus endozoochorous house construction, fuelwood NWD Triana Vismia macrophylla Kunth Bloodwood genus endozoochorous (birds) latex used to treat skin fungi (ringworm, lota, ground itch) All Lecythidaceae Bertholletia excels Bonpl. Brazil Nut agouti, labba, peccaries and monkeys fruits eaten Rupununi Leguminosae (Caesalpiniaceae) Hymenaea oblongifolia Huber Locust -fine leaf agouti, labba, peccaries, monkeys (saki and capuchin) powdery pulp eaten and beverage made from bark All (Shimiri) Leguminosae (Mimosaceae) Inga huberi Ducke Fern Tree agouti, labba, tapir, deer, powis, marudi, peccaries, fruits of various Ingas eaten; fuelwood All monkeys (capuchin, spider), parrots Inga rubiginosa (Rich.) DC. Wild Tamarin agouti, labba, tapir, deer, powis, marudi, peccaries, fruits of various Ingas eaten; fuelwood All capuchin, spider, parrots Leguminosae (Papilionoideae) Swartzia arborescens (Aubl.) Pittier Bowwood genus zoochorous (bats, rodents, monkeys) used for making bows Rupununi Malpighiaceae Byrsonima aerugo Sagot Moreye tapir, deer, marudi, powis, chachalaca, labba, agouti, fruits eaten, fuelwood All peccaries, tortoises, parrots, hanaqua, toucan, parrots, macaw, savanna fox, monkeys (Continued) 50 A. R. CUMMINGS AND J. M. READ Table 4. (Continued). Family/Species Common name Eaten by Traditional uses Region Byrsonima stipulacea A.Juss Hicha tapir, deer, marudi, powis, chachalaca, peccaries, labba, fruits eaten, fuelwood All agouti, tortoises, hanaqua, toucan, parrots, macaw, savanna fox, monkeys Moraceae Cecropia peltata L. Congo pump marudi, hanaqua, capuchin monkeys leaves of some species used for making teas for treating kidney problems, back pain and high All blood pressure. Moraceae (Cecropiaceae) Pourouma guianensis Aubl. Baruma birds and monkeys** fuelwood, leaves used like sandpaper all Myrtaceae Myrciaria vismeifolia (Benth.) O.Berg Taparowballi most animals and birds fruits used to make beverage Rupununi Palmae Astrocaryum aculeatum G.Mey. Akayuru tapir, deer, peccaries, labba, agouti, squirrel, rats, fleshy endocarp eaten All Astrocaryum vulgare Mart. Awara agouti, labba, peccaries, tapir fleshy endocarp eaten All Attalea maripa (Aubl.) Mart. Kokerite tapir, agouti, labba, tortoise, deer, black curassow, fruits eaten, used to make beverages, and leaves used for thatching roofs All regions marudi, macaws, parrots Euterpe oleracea Mart. Manicole deer, peccaries, marudi, hanaqua, agouti, labba, tapir, fruits used to make beverages in the Rupununi, heart-of-palm eaten as a vegetable (cooked or All regions macaws, parrots raw) Manicaria saccifera Gaertn. Trulli** monkeys, scatter-hoarding rodents, such as agoutis and leaves used for thatching in NWD NWD acouchis Mauritia flexuosa L.f. Ite tapir, deer, peccaries, powis, labba, agouti, acouchis, leaves used for thatching in Rupununi, fruits eaten all over Guyana Thatch tortoises, macaws, parrots, monkeys (howler and spider) Rupununi, eaten all Oenocarpus bacaba Mart. Lu deer, marudi, powis, hanaqua, agouti, labba, tapir, fruits used to make beverage All peccaries, macaws, parrots Oenocarpus bataua Mart. Turu deer, peccaries, marudi and powis, hanaqua, agouti, fruits used to make beverage All labba, tapir, macaws, parrots Sapotaceae Pouteria caimito (Ruiz & Pav.) Radlk. Asepokoballi agouti, deer, labba, spider and howler monkeys, fruits eaten All capuchins Pradosia schomburgkiana (A.DC.) Kakarua tapir, deer, peccaries, monkeys fruits eaten, bark boiled with other species to treat tuberculosis All Cronquist Sterculiaceae Sterculia pruriens (Aubl.) K.Schum. Maho agouti, labba, rats lashing material for warashis, quakes, and other traditional utensils in NWD All Sterculia rugosa R. Br. Birdwood genus synzoochronous scatter-hoarding rodents, wattles used in house construction Rupununi capuchin and saki monkeys Violaceae Paypayrola longifolia Tul. Adebero labba, agouti, peccaries, monkeys, birds fuelwood Rupununi INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 51 bataua Mart. are eaten in the Rupununi, but it is not so 2008) within the study area. There were, however, for the fruits and heart-of-palm of Euterpe oleracea potentially more than 165 groups of plants in the sam- Mart. in this region, despite heart-of-palm being ple. It is acknowledged that trees in the forests of the exported from the NWD. Perhaps the most important Rupununi, like other Neotropical settings, are difficult palm, given that its fruits are eaten by people and wild- to identify (see Dinerstein 2013), and even though 3% life and the leaves are used for thatching roofs, is Attalea did not represent a large portion of our sample, it still maripa Mart. (Kokerite). A key food source for hunted referenced species that tree spotters had not worked rodent species (Silvius & Fragoso 2003), this was the with in the past. It is possible too that some species most abundant species in this class. In fact, to highlight were misidentified, especially in the process of transfer- the favorability of the fruits in the Rupununi, cut ring common names to botanical names. In many instances, the literature (e.g. Roosmalen 1985;Polak stumps of this palm were often observed in the land- scape, reflecting the desire of the cutter to gain access to 1992; van Andel 2000) on a number of species had the ripe fruit on palms that were deemed too tall. This one common name referring to multiple botanical names. Species identification remains a challenge in multiple-use class also included a number of tree spe- cies. Geissospermum spp. or Sinkona is probably the reconciling traditional knowledge with conventional most significant. A medicinal plant, the bark of this science, and traditional knowledge provides an avenue tree is used to treat malaria and the fruits are eaten by for dealing with this challenge. It is therefore critical, as tortoises. It was suggested (van Andel 2000)that Mauro and Hardison (2000) pointed out, to preserve Sinkona is utilized in coastal Guyana, with reference traditional knowledge. Identifying all species of plants to Cinchona spp. This is an area of debate in this analysis with ecological and cultural significance is an area that as tree spotters at Iwokrama suggested that this refer- scientists must work with indigenous peoples, such as ence may be to trees in the genus Geissospermum and the tree spotters involved in this study, for clarification. not Cinchona.Additionally, Myrciaria vismeifolia In this way, traditional knowledge can play a much (Benth.) O.Berg is used to make a beverage, while the more prominent role in guiding forest management fruits of Byrsonima species are eaten throughout the decision-making processes. The difficulties of identify- Rupununi and Guyana, and traditional knowledge sug- ing plants aside, tree spotters have worked extensively gested that hunted wildlife also eat these fruits. The with the major timber and well-known species, such as fruits of M. vismeifolia are eaten by most animals and Manilkara bidentata, Mora excelsa, Carapa guianensis birds, including hunted wildlife species. Pouteria cai- Aubl. and Inga alba (Sw.) Willd. in the logging industry mento (Ruiz & Pav.) Radlk. and Pradosia schomburgki- and their knowledge of plants has been relied upon in ana (A.DC.) Cronquist are eaten by hunted wildlife, other settings. This fact gives us confidence in their especially the hunted rodents and deer species. knowledge they shared for this study. The large sample Anacardium giganteum Hancock ex Engl., Mangifera size utilized in this study, and sampling larger plants, indica L. and Spondias mombin L. are popular with coupled with the TFRK of indigenous tree spotters Amerindians and throughout Guyana. M. indica,a allowed us to establish that multiple-use plant species cultivated species, was observed on old farms through- are the foundation of the Rupununi forested ecosystem. out the study area, and is a delicacy throughout Guyana. Multiple-use plant species and the ecosystem services S. mombin fruits are sold at local markets and used associated with them have therefore allowed for a throughout Guyana as well for making a beverage. greater appreciation of what is at stake as forest areas Protium guianense (Aubl.) Marchand fruits are eaten undergo change due to LUCC processes and perhaps by a number of the hunted wildlife species and people. larger processes of climatic variability. The gum of the Protium trees, which has a camphor- like odor, is used in several ways by Rupununi Multiple-use classes Amerindians, including treating diarrhea and is burned as incense under the hammock of a sick person. This This analysis showed that multiple-use trees and multiple-use class, despite not being the most abundant, palms dominate the forests of the Rupununi. Four is important to the Amerindian culture, and requires dimensions of multiple-use plants were identified: additional examination to better understand how these wildlife food/commercial timber/traditional uses, com- species will be affected by logging and other commercial mercial timber/wildlife food, commercial timber/tradi- exploitation within their habitats. tional uses and wildlife food/traditional uses. Each multiple-use class portrays a different facet of tropical forest utilization, ecosystem services and indeed Discussion and conclusions potential for resource-use conflicts. Because of the different demands on the various multiple-use Species composition classes, the groups of plants in each class will be All 165 groups of plants in our sample were recorded by impacted differently by LUCC. Based on our classifi- previous studies (e.g. Clarke et al. 2001; Iwokrama cation, and the relationships that have been 52 A. R. CUMMINGS AND J. M. READ better off for adopting traditional knowledge in established between plants and people, and plants informing policy positions. These positions could and wildlife (e.g. Roosmalen 1985; Forte 1996; van Andel 2000; Forget & Hammond 2005; Cummings include guiding processes of plant removal for tim- ber, and for also deciding on spaces to be preserved 2013), we argue that the removal of trees in the class for traditional activities. While other studies have wildlife food, commercial timber and traditional uses, for example, will have implications for wildlife and focused on the gross-scale importance of tropical forests for regulating services such as carbon seques- people. For instance, wildlife perform ecosystem tration (see for example Houghton 2005), this study functions and supporting services for plants (see Roosmalen 1985; Forget & Hammond 2005) while was able to describe finer-scale detail for ecosystem services associated with tropical plants. As a conse- plants are associated with provisioning services quence we are contributing to establishing for forest (such as traditional medicines) and cultural services of Amerindians (see Forte 1996; van Andel 2000). managers the value that may be lost as tropical forest Since these plants are connected to each of the main areas decline due to LUCC and indeed larger pro- cesses such as climatic variability. As Parrotta and resource-use classes identified in the Rupununi – commercial logging, provision of wildlife food and Agnoletti (2007) pointed out, traditional knowledge traditional uses – a lack of sound management prac- of rural communities, in our case the Makushi and tices for these species will have far-reaching conse- Wapishiana Amerindians, has served to sustain the quences for local populations and indeed forest goods and services they need for livelihood security, ecosystem processes. Forest managers accessing tradi- and as such measures should be taken to preserve tional knowledge on how such plants are used can be traditional knowledge. Activities such as deforesta- a critical step in preventing impacts on indigenous tion as it relates to gold mining, commercial logging people’s livelihood and wildlife. and large-scale farming that are features of tropical landscapes may remove plants while bringing with them change, such as socioeconomic pressures, that Synthesis and future directions may erode traditional knowledge. As such, we concur with the position of Parrotta and Agnoletti (2007) The objectives of this paper were to identify tropical and others who suggest that efforts are required to multiple-use plants and describe the ecosystem ser- arrest the decline of traditional knowledge in indi- vices associated with these in order to begin the genous people-produced landscapes. process of understanding what value is lost as tropical In essence then, we argue that the identification of forest areas decline. Our analysis has shown that multiple-use species using traditional knowledge is there are many multiple-use plants in tropical forests critical, because it provides knowledge of the goods that require special attention for ecological studies. and services that are kept within tropical landscapes. By exploring multiple-use plants through the lens of Furthermore, while we agree with the position of how they are used as commercial timber, wildlife Sutherland et al. (2014) that traditional knowledge food and traditional purposes, and by drawing on ‘. . .can provide complementary perspectives . . .often traditional knowledge to identify these plants and lacking in conventional science’, we emphasize that define their importance, we suggest that as tropical traditional knowledge can provide the basis for scien- forest areas decline the potential impact may be bet- tific inquiry into questions of both landscape ecology ter appreciated. and economics. Our work suggests that traditional Our analysis would not, however, have been pos- knowledge can provide insights to scientists on how sible without the availability and accessibility of tra- to fully grasp the impacts of LUCC, beyond the ditional knowledge. In fact, without the utilization of typical measurements of area loss (e.g. Kim et al. traditional knowledge in two dimensions – spotting 2015) or implications for the carbon cycle (Melillo (identifying) plants in the forest and describing how et al. 1996; Baccini et al. 2012; Zarin 2012; Kim et al. these plants are used for traditional purposes, wildlife 2015). While these areas are important, our work food and commercial timer (Tables 1–4) – the value suggests that attention needs to be paid to the of our study would have been significantly reduced. impacts of LUCC on the multiple roles forests and In this sense, this paper is a direct response to the the individual species of plants within them play for calls from scholars such as Sutherland et al. (2014), humanity and wildlife. Traditional knowledge pro- Fazey et al. (2006) and Raymond et al. (2010) for the vides that basis of developing such knowledge. greater use of local and traditional knowledge along- Furthermore, more indirectly, and perhaps more sub- side conventional knowledge for making decisions tly in the immediate term, are the impacts of climatic about biodiversity and natural resources. While we variability on the services traditional knowledge can view our study as a part of the process for beginning inform us about. Scholars and forest managers alike to value tropical landscapes for the ecosystem services need to better understand the conditions that lead to they provide, we do make the case that forest man- overexploitation of forests as this will help them agement decision-making processes are going to be INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 53 addressing pressing ecological questions on the better appreciate the value that is lost to humanity as impacts of forest change. Furthermore, as more forest areas decline. Such an understanding will lead to a clearer picture of the value that is lost to human- work unfolds on how plants are used by rural communities through the examples of González- ity and wildlife as forest area is lost. By responding to Marín et al. (2012) in Mexico, Khan et al. (2013) the call of Parrotta et al. (2009) for documenting TFRK, our paper has established, through multiple- in Pakistan, Klimas et al. (2007, 2012a, 2012b)in Amazonia and Mwavu and Witkowski (2009)in use plants, some of the complexities and nuances that Africa, our work contributes to setting up species are associated with TFRK that must be accounted for in the process of valuing the losses to humanity as and their importance for more in-depth analysis on how changes in landscape can impact ecosystem tropical forest areas decline. In this sense, traditional services and landscape sustainability. In the knowledge on tropical landscapes can serve both as a basis for understanding the value of landscapes Guyana setting, the work of Forte (1996) and van change, but also for the justification for investment Andel (2000) established the foundation for describing the relationship between indigenous peo- in more robust conservation and management strate- gies within these landscapes. ples and plants. Our work builds on these past As the literature on multiple-use species continues efforts by providing insights into plant taxonomy to evolve, the interaction between the various (Mauro & Hardison 2000) and ecosystem services resource uses in tropical forests, such as wildlife of individual plants, and provides the basis for food sources, commercial timber and traditional further spatial and socioeconomic analysis. In addi- uses, will require urgent attention. The conflicts tion, our work goes beyond previous studies, such between wildlife food sources and commercial log- as Scarpa (2009), whose data were based on pub- ging, for example, have only been reported in a few lished studies, Singh et al. (2015), Klimas et al. instances (Plumptre 1995; Mwavu & Witkowski (2007, 2012a, 2012b) who have considered single 2009). Mwavu and Witkowski (2009) identified 15 species, or McPherson (2014), where species distri- such multiple-use species in Budongo Forest bu tion models were derived from museum samples, Reserve, Uganda, while the present study suggests by providing a wider list of plants, through direct that more than approximately 61 such species are in interactions with the holders of TFRK as suggested this potential resource-use conflict area and should by Parrotta et al. (2009). therefore gain the attention of more detailed ecologi- While our work provides a list of plants and how cal studies. The intersection between wildlife food they are used for various ecosystem services, data on and commercial logging in tropical plant species is the current status of such species are not presented in critical, especially in those communities that are mov- the paper, including the abundance of species and the ing toward commercial logging as an alternative for level of dependence that local people have on each supporting their livelihoods. As various sectors of a both at the scale of the study area and at individual community embrace new technology at different study sites. The status of the ecosystem services rates, this change can potentially lead to local-scale associated with individual plant species is perhaps resource-use conflicts, where logging may place pres- the most pressing concern, as this will provide sure on the very food source hunted wildlife rely insights into the sustainability of local livelihood upon. practices and larger sustainable forest management Besides the species of wildlife that are hunted in questions. Traditional knowledge will be critical in the Rupununi, the needs of frugivorous primates addressing these concerns and helping craft immedi- should also be included in future analysis. For exam- ate changes in management practices to deal with the ple, Forget and Hammond (2005) pointed out that impacts of LUCC under various scenarios of resource even carnivorous species, such as coati living in the exploitation. forests of the Guiana Shield, may eat the fruits and Finally, in following up on the prescription for parts of trees in their habitat. Therefore, as suggested dealing with the hindrances to sustainable forest by Peters (1996) and Forget and Hammond (2005), management (Peters 1996), future research in the understanding which plant species provide food for Rupununi, and indeed areas where tropical forests animals, their demographics and how they respond to are continuing to decline, should focus on the spa- disturbance (such as commercial logging) is critical, tial distribution of multiple-use species and in the context of both the study area and the wider resource extraction activities. The list provided is Neotropics. With the high number of multiple-use this study can serve as the basis for local scale species in the forests of the Rupununi, it would (Rupununi) to larger scale – Guyana and the seem likely that there are also high numbers else- wider Neotropics – analysis of the implications of where in Guyana and the wider Neotropics. LUCC on the ecosystem services associated with Through embracing traditional knowledge, our multiple-use plants. Furthermore, through the list studyprovidesscientistswithastronger basisfor of plants provided in this paper, further studies can 54 A. R. CUMMINGS AND J. M. READ Baccini A, Goetz SJ, Walker WS, Laporte NT, Sun M, be established to determine the contribution LUCC Sulla-Menashe D, Hackler J, Beck PSA, Dubayah R, is making to the erosion of TFRK in indigenous Friedl MA, et al. 2012. Estimated carbon dioxide emis- peoples-produced landscapes of Guyana and sions from tropical deforestation improved by carbon- beyond. 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Analysis of hunting options by the use of Acknowledgments general food taboos. Ecol Modell. 110:5–17. Communities of the North Rupununi. 2006. By-laws for I acknowledge and thank various members of the Moses the management of natural resources by the commu- family, Persaud, Seigmond, Carro, Ricky and Danford, Steve nities of the North Rupununi. Final Report from Andries and Benedict Joseph. At Project Fauna Drs Jose Government of Guyana/United Nations Development Fragoso, Han Overman and Jeff Luzar, and the field teams Programme Sub Project, Promoting Community supported data collection. The Guyana Environmental Participation in the Management of Natural Resources Protection Agency (EPA) in conjunction with the Ministry and Environment. Georgetown (Guyana): Ministry of of Amerindian Affairs granted permission for this research, Amerindian Affairs. while the villages across the Rupununi allowed us to work in Costanza R, d’Arge R, De Groot RS, Farber S, Grasso M, their forests. 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Journal

International Journal of Biodiversity Science, Ecosystem Services & ManagementTaylor & Francis

Published: Jan 2, 2016

Keywords: Traditional knowledge; multiple-use plants; ecosystem services; wildlife food; Rupununi; Guyana

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