https://orcid.org/0000-0003-1824-5500
Figures
Abstract
Wild edible plants (WEPs) have long supported rural livelihoods and cultural practices in Bhutan, yet their use and associated knowledge are increasingly challenged by agricultural change, modernization, and limited documentation. This study documented the diversity, use patterns, and socio-ecological drivers of WEP use across Bhutanese communities in all 20 districts using purposive and snowball sampling of knowledgeable informants. Data were analyzed for species composition, life form, plant parts used, food categories, utilization trends, and ethnobotanical indices. A total of 114 WEP species belonging to 61 families were recorded. Urticaceae had the highest family importance value, herbs constituted the dominant life form (42%), and fruits were the most frequently used plant part (36.89%). Accessibility and availability were the main factors supporting continued use, whereas perceived ecological change, resource scarcity, land-use change, and limited knowledge constrained utilization. Fruits and vegetables dominated current use patterns, but responses also indicated declining knowledge transmission across generations. These findings highlight the continuing nutritional and cultural importance of WEPs in Bhutan and support the need for conservation, sustainable harvesting, and stronger integration of indigenous knowledge into education and policy.
Citation: Jamtsho T, Tobgay, Takchu U, Wangchuk P (2026) Wild edible plants of Bhutan: Diversity, use patterns, and motivations for consumption. PLoS One 21(8): e0355212. https://doi.org/10.1371/journal.pone.0355212
Editor: Mehmooda Munazir, Government College Women University Sialkot: GC Women University Sialkot, PAKISTAN
Received: October 9, 2025; Accepted: July 18, 2026; Published: August 28, 2026
Copyright: © 2026 Jamtsho et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Data Availability: All relevant data are within the manuscript.
Funding: The author(s) received no specific funding for this work.
Competing interests: The authors have declared that no competing interests exist.
Abbreviations: WEPs, Wild edible plants;; Masl, Meters above sea level; FAO, Food and Agriculture Organization; UV, Use Value; ICF, Informant Consensus Factor; RFC, Relative Frequency Citation; FIV, Family Importance Value; FL, Fidelity Level
Introduction
Wild edible plants (WEPs), commonly referred to as bush foods, have long been integral to traditional food systems worldwide. The Food and Agriculture Organization (FAO) defines WEPs as plants that grow naturally in self-sustaining populations within natural or semi-natural ecosystems without direct human intervention [1]. Historically, they have played a crucial role in sustaining communities during periods of food scarcity, conflict, and natural disasters [2,3]. Research further shows that WEP use patterns vary markedly across cultural and geographic settings [4], highlighting their diverse functions and values. Even today, WEPs remain essential for approximately one billion people globally, especially in tribal and rural communities [5]. The edible parts of WEPs include inflorescences, shoots, buds, fruits, nuts, seeds, leaves, bark, stems, and roots [6,7], which are consumed either raw or prepared as spices and vegetables to suit local culinary preferences [8,9]. Wild edible fruits are particularly valued for their richness in micronutrients, vitamins, and antioxidants [10,11], while wild spices and vegetables contribute energy and essential nutrients that support dietary diversity and nutrition security [12].
Despite these benefits, global WEP use and knowledge are declining. Key drivers include urbanization, land-use change, agricultural expansion, loss of indigenous knowledge, sociocultural transformation, and climate change [5]. This trend has raised concerns amid rising food insecurity and biodiversity loss. As the world population is projected to exceed 9 billion by 2050, requiring a 50% increase in food production, experts suggest that the domestication and sustainable utilization of WEPs and other neglected species will be critical to future food systems [13].
Within this global discourse, Bhutan presents a distinctive ecological and cultural landscape for the study of WEPs. Situated in the eastern Himalayas and straddling the Indo-Malayan and Palaearctic biogeographic realms, Bhutan lies between China and India, two of the world’s most populous countries. The country is characterized by dramatic altitudinal variation, from 150 meters above sea level (masl) in the subtropical south to more than 7,500 masl in the northern alpine regions, resulting in highly diverse topography, ecosystems, and ecofloristic zones [13,14]. This diversity, together with substantial summer monsoon precipitation, places Bhutan within one of the world’s ten biodiversity hotspots [10].
Bhutan’s national development is guided by the philosophy of Gross National Happiness (GNH), which emphasizes environmental preservation alongside socioeconomic well-being. As a result, 71% of the country’s land area remains under forest cover, and 51% is protected through national parks, wildlife sanctuaries, nature reserves, and biological corridors [14]. However, the Flora of Bhutan has documented just over 5,603 species of flowering plants spanning 220 families and 1,415 genera, indicating substantial scope for further exploration. Approximately 94% of these species are native, and about 105 are endemic [7]. Many of these plants, including lichens, are traditionally used as medicinal plants and bush foods [15], contributing to both nutritional health and indigenous healing systems [16].
Despite this richness, wild edible plants in Bhutan have received limited research attention, as reflected in the small number of publications and policies devoted to them. As in other parts of the world, rapid urbanization and agricultural intensification have contributed to declines in both WEP consumption and the transmission of ethnobotanical knowledge [5,16,17]. In Bhutan, these shifts are compounded by environmental degradation, changing social values, and declining interest among younger generations [18,19]. Consequently, Bhutan risks losing the cultural and biological heritage associated with WEPs.
Against this backdrop, a nationally integrated assessment is needed to move beyond descriptive inventories and examine how WEP diversity, habitat association, use patterns, and socio-ecological drivers interact in Bhutan. This study therefore characterizes WEP diversity across communities, evaluates the cultural importance of taxa and families using ethnobotanical indices, assesses use patterns across food categories and plant parts, and identifies factors associated with continuing, increasing, moderate, or declining use.
Guided by ethnobotanical theory and global patterns of WEP use, we hypothesize that WEP importance is unevenly distributed across taxa, that forest-associated and herbaceous species make major contributions to use, and that land-use change and agricultural transitions are among the principal forces shaping contemporary WEP utilization. By addressing these questions, the study seeks to provide an evidence base for conserving biocultural diversity and supporting community-based resource management strategies in Bhutan.
Methodology
Study site and informant selection techniques
Located in the eastern Himalayas, Bhutan lies between 26.7263° and 28.2466° N and 88.7495° and 92.1236° E. It is bordered by India to the east, west, and south, and by China to the north. Administratively, the country is divided into 20 districts and 205 gewogs (blocks) [20]. Following a reconnaissance survey, data were collected across all 20 districts using purposive and snowball sampling to identify informants with recognized knowledge of WEPs. The final sample comprised 192 informants, including 20 key informants (16 male and 4 female) and 172 additional community informants (110 male and 62 female).
Key informants were selected on the basis of experience in identifying, collecting, and using WEPs, whereas additional informants were recruited to document more widely shared knowledge and to capture variation in knowledge transmission across age groups. Initial contacts were facilitated by district forest officials and local leaders, after which recruitment proceeded through community referral. The study employed non-interventional ethnobotanical interviews, home-based plant demonstrations, and guided field walks. Authorization and field clearance to conduct the study were granted by the Jime Khesar Strict Nature Reserve, Department of Forests and Park Services, Ministry of Agriculture and Forests, Royal Government of Bhutan (letter no. JKSNR/NCS-07/2022–2023/364, dated 16 March 2022). The Jigme Khesar Strict Nature Reserve coordinated and facilitated a field visit with relevant local authorities during data collection. All participants were informed about the purpose of the study, confidentiality procedures, the voluntary nature of participation, and their right to withdraw at any time without penalty, and prior informed consent was obtained from each participant before conducting the interviews.
Because the sampling design prioritized expertise rather than statistical balance, respondent numbers were uneven across districts. District-level patterns are therefore presented descriptively, whereas the main interpretation focuses on national-level patterns of diversity, utilization, and socio-ecological drivers.
Questionnaire design
The questionnaire was developed in two sections. The first section addressed the socio-cultural attributes of the informants, including age, gender, and native origin. The second section focused on the traditional knowledge of informants regarding the uses of wild edible plants, covering aspects such as local and vernacular names, life forms, utilized parts, food categories, methods of usage, and modes of consumption. Expert review and feedback were sought on the questionnaires to ensure face and content validity prior to data collection.
Data collection tools and procedures
Data were collected through semi-structured interviews, home-based plant demonstrations, and guided field walks. Interview schedules recorded local names, habitats, life forms, food categories, plant parts used, preparation methods, modes of consumption, and perceived changes in WEP use. Questions were prepared in English and, where necessary, administered in local languages to ensure accurate capture of local knowledge.
Home-based demonstrations were particularly useful for older informants with limited mobility, whereas other participants demonstrated collection and preparation practices in both household and field settings. Field surveys were conducted across all four seasons from March 2022 to December 2023, and specimens were collected in flowering or fruiting condition wherever possible. Photographs of live plants were taken, and specimens were processed using standard herbarium procedures recommended by Smith and Chinnappa [21]. Voucher specimens were deposited at the Jigme Khesar Strict Nature Reserve, Department of Forests and Park Services, Haa District, Bhutan. Taxonomic identification was verified against the Flora of Bhutan and other Himalayan references, and botanical names were cross-checked using World Flora Online, eFloras, and TROPICOS.
Data analysis
Data were analyzed at the national level. Species diversity, life forms, habitats, food categories, plant parts used, and utilization trends were summarized using frequencies and percentages in figures and tables. Explanations for changes in WEP use were coded into environmental, economic, sociocultural, agricultural and land-use, and human-wildlife conflict domains; because informants could provide multiple explanations, Table 5 reports coded response frequencies rather than counts of unique informants. Chi-square tests were used to examine the association between WEP food categories and utilization trends, and chi-square goodness-of-fit tests were used to evaluate the distribution of coded responses within motivation domains.
Ethnobotanical indices, including Use Value (UV), Informant Consensus Factor (ICF), Relative Frequency Citation (RFC), Family Importance Value (FIV), and Fidelity Level (FL), were calculated following established methods [22–25].
The Family Importance Value (FIV) was determined by calculating the percentage of informants who referenced each family. It was computed as follows:
where FC represents the count of informants mentioning the family, and N signifies the total number of informants involved in the study.
The Informant Consensus Factor was used to assess the consistency of knowledge regarding WEP use [26]. ICF is computed as follows:
Where ‘nur’ represents the number of use reports for a given use category, and ‘nt’ indicates the number of taxa used for that category across all informants. ICF values are typically low (close to 0) when plant selection is random or when information exchange among informants is limited. Conversely, ICF values approach 1 when communities apply well-defined selection criteria or actively share knowledge [27].
To provide quantitative assessments of the relative importance of individual plant species to the informants, the Use Value (UV) was computed objectively [28]. This calculation was performed using the formula:
UV = ΣU/n, where UV denotes the use value of a species, U represents the number of use reports mentioned by the respondents, and n signifies the total number of respondents interviewed.
The Relative Frequency Citation (RFC) measured the frequency with which specific species were reported. It was calculated as follows:
Where FC denotes the number of informants mentioning a particular WEP, and N represents the total number of informants surveyed. RFC values range from 0 to 1, with higher values indicating greater local importance of a species [25].
The Fidelity Level (FL) index was used to identify the extent to which a species was associated with a particular food category when the same species had multiple reported food uses. It is calculated as follows:
FL = (Np/N x 100). Here, Np represents the number of informants who cited a species for a particular food category, whereas N represents the total number of informants who cited that species for any food use. Higher FL values therefore indicate stronger agreement on the association of a species with a specific food category [24].
All ethnobotanical indices (RFC, FIV, UV, and FL) were calculated at the national level by aggregating responses across districts. This approach provided an overall assessment of WEP importance and use patterns in Bhutan. However, because the purposive and snowball sampling design produced uneven representation across districts, these indices were not further stratified by ecological region for inferential comparison. Such stratification would require balanced, systematically distributed sampling across regions to ensure statistical validity and comparability. The calculated indices are therefore interpreted as indicators of national-level patterns rather than region-specific differences.
Nevertheless, to explore whether citation values varied among ecological zones, the species citation data in Table 1 were matched to the ecological-zone categories reported in Table 2 and compared using a Kruskal-Wallis test. Because several zones were represented by very few species, the inferential comparison was restricted to the six zones with at least five recorded species (Ap, Bf, Brf, Cf, Stf, and Wbf).
Results and discussion
Demography and social structure
Demographic information was collected across districts to characterize the respondent profile and contextualize ethnobotanical knowledge of WEPs, following Kumar et al. [29]. The respondents were distributed across all districts, providing broad national coverage of WEP knowledge and practices. However, the number of respondents varied considerably, ranging from 38 in Mongar to 1 each in Gasa and Paro. This disparity likely reflects differences in population size, settlement patterns, and the availability of recognized knowledge holders across regions.
Such variation in respondent distribution is an expected outcome of purposive ethnobotanical sampling, which prioritizes knowledgeable informants over statistically uniform representation. Nevertheless, this imbalance limits the robustness of direct district-level comparisons. District-level variation is therefore presented descriptively rather than interpreted as a statistical ranking of WEP richness or use intensity. Within this sampling context, gender may also shape ethnobotanical knowledge through differences in livelihood roles, access to resources, education, and social networks [30]. Although efforts were made to include both sexes, the respondent pool was male-skewed, with males accounting for 66% (n = 126) and females 34% (n = 66) (Fig 1a). This pattern suggests that the documented knowledge may partly reflect gendered differences in access to WEP collection and use and contrasts with studies reporting a stronger association between women and wild plant knowledge [24,27].
Age distribution further showed that respondents aged 40–59 years constituted the largest group, representing 48% (n = 93) of the total sample (Fig 1b). Individuals in this age group, predominantly male, demonstrated comparatively high levels of WEP knowledge. This pattern may reflect the Bhutanese context, in which men are more actively involved in field collection of wild plants, whereas women are often more engaged in household food preparation and nearby resource gathering. Similar observations have been reported elsewhere [31].
WEP diversity and habitat distribution patterns
This ethnobotanical survey documented 114 WEP species across 61 families in 20 districts of Bhutan. Earlier studies from Bhutan focused mainly on wild edible fruits, non-wood forest products, or the ethnomedicinal properties of wild plants in specific regions. For example, Bajpai reported 120 WEPs from 63 families in southeastern Bhutan [32], whereas Yangdon recorded 52 wild edible plants from 47 genera and 35 families in eastern Bhutan [19]. The present study takes a broader approach by covering all major WEP food categories. Comprehensive information on family and botanical names, local names, life forms, plant parts used across food categories, modes of consumption, frequency citation (FC), and relative frequency citation (RFC) is presented in Table 1.
Among the recorded families, Lauraceae contained the highest number of species [6], followed by Rosaceae, Asparagaceae, and Asteraceae (5 each), while Fabaceae, Moraceae, and Urticaceae each contained 4 species (Table 3). Similar dominance of Moraceae, Rosaceae, Fabaceae, Lauraceae, Rutaceae, and Urticaceae has been reported in Uttarakhand, India [32], among the Gaddis, a tribal community of the western Himalaya [33], in Burji District, Ethiopia [22], in Chile [34], and in eastern Bhutan [19]. The remaining 56 families were represented by fewer than 3 species each (Tables 1 and 3). Details on frequency citation (FC), relative frequency citation (RFC), and family importance value (FIV) for each family are provided in Table 3.
The application of plant family use value in ethnobotanical studies enables the assessment of the significance of different plant families, facilitating hypothesis testing, statistical validation, and comparative analysis [35]. Urticaceae, with an FIV of 6.250%, represented by four plant species, emerges as the most valuable family for wild edibles. Following Urticaceae, Lauraceae, Rosaceae, Acanthaceae, Arecaceae, Asparagaceae, and Elaeagnaceae also contribute substantially, reflecting their importance in local diets and traditional practices (Table 3). The suitability of habitats, favorable environmental conditions, and frequent interactions with local communities likely explain the dominance of these families [36]. Consequently, the traditional use of these plant species is widely recognized among the inhabitants.
While these indices provide valuable insights into the relative importance of plant families at the national scale, a supplementary ecology-stratified comparison of species citation values was undertaken by linking the species records in Table 1 to the ecological-zone categories in Table 2. Citation values did not differ significantly among the six sufficiently represented ecological zones (Kruskal-Wallis χ² = 5.12, df = 5, p = 0.402; n = 107 species), although warm broadleaved forest (Wbf) showed the highest median citation value. A sensitivity analysis using four broader ecological groups yielded the same conclusion (χ² = 1.73, df = 3, p = 0.630). Within the limits of the present sampling design, ecological variation in citation values is therefore interpreted descriptively rather than as evidence of statistically distinct among-zone differences.
Beyond taxonomic importance, the habitat distribution of wild edible plants also reflects their ecological and functional diversity. These species occur across a wide range of environments, from lowland to highland regions, including forests, fallow lands, riverine vegetation, scrublands, open lands, wetlands, alpine meadows, and disturbed habitats. Notably, a substantial proportion (47.5%) of WEPs were collected from forest ecosystems, followed by abandoned fields and riverine vegetation (Fig 2a). In terms of life forms, herbs dominated (42%), followed by trees (34%), shrubs (15%), and climbers (9%) (Fig 2b), indicating a strong reliance on both understory and woody vegetation components (Table 1).
(a) Percentage distribution of habitats from which wild edible plants were collected, showing the predominance of general forest habitats, followed by sites adjacent to cultivated fields and riverine vegetation. (b) Life forms among recorded wild edible plants are dominated by herbs, followed by trees, shrubs, and climbers. (c) Distribution of species citation values across the six sufficiently represented ecological zones included in the inferential comparison. Boxes show the median and interquartile range, whiskers indicate the range, points represent individual species, and sample sizes are shown above each box. Citation values did not differ significantly among ecological zones (Kruskal-Wallis χ2 = 5.12, df = 5, p = 0.402). Abbreviations: Ap, alpine; Bf, blue pine forest; Brf, broad leaf forest; Cf, chir pine forest; Stf, subtropical forest; Wbf, warm broad leaf forest.
This ecological dependence has important implications for conservation. The findings of the present study indicate that WEP utilization is closely linked to habitat availability and is increasingly constrained by locally perceived environmental pressures. A large proportion of WEPs are harvested from forest habitats that are vulnerable to disturbance and degradation. In addition, respondents identified ecological change, access limitations, and disturbances (including fire and human or animal interference) as key factors influencing WEP availability (Table 5). These constraints are further reflected in utilization trends, as a notable proportion of respondents reported declining WEP use over time (Fig 5).
Collectively, these patterns indicate that WEP conservation in Bhutan is not merely a general or theoretical concern but a locally experienced issue shaped by habitat dependence, environmental change, and shifting use patterns [37–39]. Preserving and revitalizing WEPs therefore requires integrated strategies that combine conservation, sustainable harvesting, and protection of traditional ecological knowledge. Approaches such as agroforestry, seed banking, and community-based conservation programs offer practical pathways to strengthen the long-term resilience and sustainability of these resources [40,41]. In addition, integrating WEPs into contemporary food systems through education and sustainable commercialization may reinforce their cultural value while supporting biodiversity conservation.
Use patterns of WEPs
Recorded WEPs spanned multiple food categories, but use was concentrated in fruits and vegetables rather than evenly distributed across all categories (Fig 3a; Table 1). Fruits accounted for the largest share of reported consumption modes (38%), followed by vegetables (23%), and fruit was also the single most frequently used plant part (36.89%) (Fig 3b). This pattern indicates that current WEP use in Bhutan is centered on a relatively small set of nutritionally and culturally important food types. The prominence of widely traded taxa further shows that WEPs contribute simultaneously to household consumption, local exchange, and culinary identity in Bhutan [42]. Examples include Cymbidium floribundum, Cinnamomum zeylanicum, Plectocomia himalayana, Phyllanthus emblica, Machilus edulis, Diplazium maximum, Houttuynia cordata, and Elatostema lineolatum [43]. Rather than functioning as marginal foods, these taxa form part of an active food system that links nutrition, markets, and local food practices [44,45]. Comparable evidence from broader reviews and market studies shows that WEPs often combine dietary, commercial, and cultural functions rather than serving only as fallback foods [4,46]. These representative market taxa are shown in Fig 4.
Trends in the usage of wild edible plants
Globally, forest resources and their products remain vital to the livelihoods of a large share of the world’s population [47]. In Bhutan, people from diverse communities collect wild edible plants for both nutritional and medicinal purposes. Several respondents expressed concern about declining abundance and availability of these plants in adjacent forests, noting that some now travel for more than an hour to collect them. More broadly, WEP consumption often increases under conditions of food scarcity, poverty, and extreme climatic events [48]. In many developing countries, limited access to adequate food supplies continues to drive reliance on wild edible plants [49].
Wild edible plants therefore remain important to food security in rural communities worldwide. Incorporating them into diets also aligns with the United Nations Sustainable Development Goal of ending hunger [50]. WEPs are recognized sources of vitamins, minerals, and other essential nutrients, providing sustenance and micronutrients to rural households throughout the year [5,16,17]. Numerous studies have demonstrated their importance for food and nutrition security [51,52]. However, the specific nutritional profiles of many reported WEPs remain unknown [53].
One study reported the mineral composition of five WEPs, with fiber content ranging from 2.7 to 12.5 g/100 g and protein content ranging from 1.8 to 6.8 g/100 g. Similarly, research showed that the rhizomes of Dioscorea species contain five times more protein and fiber than potatoes and sweet potatoes [54]. Despite these findings, the actual contribution of WEPs to daily dietary requirements remains insufficiently understood and warrants further investigation [28].
To this end, collecting wild edible plants has long been part of Bhutanese farming livelihoods [44]. This practice functions as a social safety net under variable agricultural conditions, as rural households draw on diverse WEPs to offset shortfalls in farm production [52]. Consequently, these plants contribute not only to subsistence but also to household income. Overall utilization trends are shown in Fig 5 and summarized in Table 4, where continuing use was the most frequently reported trajectory (46%).
Table 4 summarizes 206 categorical responses across food categories, comprising 56 increasing, 95 continuing, 45 declining, and 10 moderate responses. Use trajectories were mixed, with continuing use the most frequently reported pattern (46%), followed by declining use (28%), increasing use (22%), and moderate use (4%) (Fig 5). Because these data are categorical frequencies, a chi-square test of independence provided an appropriate inferential assessment of whether utilization trends varied across food categories. The test showed a significant association between WEP category and utilization trend (chi-square = 47.84, df = 33, p = 0.046), indicating that increasing, continuing, declining, and moderate use were not evenly distributed across categories. Respondents linked declining use to domesticated alternatives, environmental change, shifting food preferences, and reduced knowledge of species identification, particularly among younger generations [46,55]. At the same time, WEPs continue to function as a support resource for rural livelihoods in Bhutan [44].
Column totals in Table 4 show that fruits and vegetables together accounted for 60.71% of increasing-use responses (34/56), 75.79% of continuing-use responses (72/95), 68.89% of declining-use responses (31/45), and 90.0% of moderate-use responses (9/10). Vegetables dominated the increasing and continuing classes, whereas fruits dominated the declining class. These patterns indicate that the food categories most central to everyday WEP consumption are also those undergoing the clearest change. This interpretation is consistent with studies that similarly identify fruits and vegetables as dominant WEP categories and as especially sensitive to ecological and social change [56]; see also (58, 59) and [57,58]. Comparable concentration of use around fruits, leaves, and vegetable species has been reported in broader WEP syntheses and regional food-system studies from Africa, the Hindu Kush, and southern Europe [59–61]. These category-specific differences are illustrated in Fig 6.
Stacked bar charts show the frequency of responses (n) for four utilization trends: increasing, continuing, moderate, and declining use. Each bar represents a specific food category, with segment heights indicating the number of informant responses for each trend. The results highlight category-specific patterns in the persistence and decline of WEP use.
The significant category-level association therefore adds inferential support to the descriptive pattern: change in WEP use is concentrated in the most widely used food categories rather than dispersed across the entire repertoire. This suggests a reorganization of use within the core food system rather than a uniform retreat from wild foods. The observed decline in some categories is consistent with broader reports that shifts in access, preferences, and harvesting conditions can affect the persistence of everyday WEP use [62,63]. Similar reorganization of wild food use in response to social and environmental change has been documented in Spain and West Sumatra [64,65].
Taken together, the results suggest that traditional WEP knowledge is not disappearing uniformly but may be transmitted less securely where access declines and regular use is interrupted. In this sense, the reported decline reflects both ecological pressure and weakening opportunities for intergenerational learning. Comparable evidence from Guatemala showed that WEP knowledge was transmitted mainly through relatives, and that people who learned through family networks knew more plants and recognized them better than those learning through other pathways [66]. Comparable generational change linked to reduced forest access has also been documented among Mapuche communities in Chile [67].
Socio-ecological drivers of WEP use
A total of 922 coded responses were provided by 192 informants regarding factors that promote or constrain WEP use. Because Table 5 summarizes categorical response frequencies rather than continuous measurements, inferential assessment focused on chi-square goodness-of-fit tests within each domain. Domain totals were 187 environmental, 186 sociocultural, 184 agricultural and land-use, 182 economic, and 183 human-wildlife conflict responses (Table 5).
Responses within every domain were strongly uneven rather than uniformly distributed: environmental (chi-square = 200.43, df = 5, p < 0.001), sociocultural (chi-square = 92.12, df = 4, p < 0.001), agricultural and land-use (chi-square = 149.57, df = 6, p < 0.001), economic (chi-square = 295.98, df = 3, p < 0.001), and human-wildlife conflict (chi-square = 148.57, df = 5, p < 0.001). High availability dominated the environmental domain, local food preference, traditional practices, and medicinal value dominated the sociocultural domain, land-use change dominated the agricultural and land-use domain, free collection dominated the economic domain, and human disturbance dominated the human-wildlife conflict domain. These results show that WEP utilization in Bhutan is shaped by a limited number of dominant motivations and constraints rather than evenly distributed influences. These domain-level patterns are illustrated in Fig 7.
Bar charts represent the frequency of informant responses (n) across five domains: (a) environmental factors, (b) sociocultural drivers, (c) agricultural and land-use practices, (d) economic factors, and (e) human-wildlife conflict. Within each panel, bars correspond to specific explanations, with heights indicating the number of responses associated with each factor. Across domains, responses were unevenly distributed, with a limited number of dominant drivers accounting for most reported influences on WEP utilization.
Table 5 strengthens the interpretation of Table 4 by showing that change in WEP use is tied to a concentrated set of socio-ecological drivers. The prominence of local food preference, traditional practices, and medicinal value indicates that continued use in Bhutan is sustained not only by subsistence need but also by taste, cultural embeddedness, and the food-medicine interface. Similar patterns have been reported elsewhere, where the cultural significance of wild foods and the persistence of taste-based gathering practices help maintain use despite broader socioeconomic change [62,63]; see also [68]. The importance of taste, culinary memory, and food-medicine links in maintaining WEP use has likewise been emphasized in Italy and other Mediterranean settings [2,69].
At the same time, the strong signals for land-use change, environment-driven scarcity, and disturbance indicate that declining use is being shaped by both ecological pressure and changing livelihood conditions. This interpretation is consistent with studies showing that wild food use becomes less secure when access changes, habitats are modified, or harvesting conditions deteriorate [62,70]; see also [57,58]. In the present study, this is reflected most clearly in fruits and vegetables, which remained the dominant food categories but also accounted for most declining-use responses.
The dominance of free collection in the economic domain suggests that WEPs continue to provide a low-cost supplement to household diets and small-scale livelihoods, while the prominence of local food and traditional practices shows that their value is simultaneously economic and cultural. Comparative studies likewise note that wild foods can support both subsistence buffering and livelihood resilience, especially where cash resources are limited [71,72]. Overall, the statistical pattern supports interpreting WEP change in Bhutan as a reorganization within the most important food categories, driven by interacting ecological, sociocultural, and economic pressures rather than by simple abandonment of wild foods [73]; see also [74].
Preservation and governance of WEPs
Wild edible plants (WEPs) are recognized as an economical and nutritious food source that contributes substantially to the global food supply, particularly in Himalayan regions [1,75]. In Bhutan, rural communities use WEPs for household consumption and income generation through local marketing [16,76,77]. Their value also derives from their nutritional content, including proteins, vitamin B2, and vitamin C, which makes them viable supplements to plant-based diets [78,79]. Despite their role in food security, WEPs are often neglected and stigmatized as foods for poorer households [5]. However, several studies have noted declining accessibility of WEPs, because of habitat degradation, land-use and forest-management change, overexploitation, changing dietary preferences, rapid development, and erosion of traditional ecological knowledge [80–83], and similar trends have been reported in Bhutan [19,44].
Growing interest in local products and the commercialization of high-value or rare WEP species by farmers has led to unsustainable extraction and declining populations [71,72]. For example, unsustainable harvesting has pushed popular species such as Gnetum buchholzianum and G. africanum toward vulnerable status. Likewise, species of Dioscorea have shown declining populations in Nangkor Gewog, Zhemgang [84], while Pogostemon amaranthoides Benth. and Persea bootanica (Meisn.) Kosterm. have declined in Maenbi Gewog, Lhuntse, because of overexploitation and destructive collection practices such as branch cutting [45]. These cases highlight the need to control overexploitation, promote environmentally responsible harvesting methods, and prevent illegal collection of rare WEP species. The herb gardens established in Lingshi (4,000 masl) and Lingmithang (600 masl) for ex situ conservation of crops, medicinal plants, and aromatic plants demonstrate the value of similar approaches for important and rare WEPs [85]. Likewise, in situ conservation in the Yakpugang Community Forest in Kelikhar, Mongar District, eastern Bhutan, illustrates the value of diversifying these approaches nationwide (86). Long-term sustainability will therefore depend on coordinated off-site and on-site conservation, domestication, and management strategies involving government agencies, local communities, and individual resource users [16,19,71,84].
Numerous studies show that the preservation and utilization of WEPs extend beyond botanical or ecological concerns and play an important role in supporting the quality of life of rural communities in many developing regions. WEPs are particularly important during periods of food shortage and continue to form part of everyday diets for millions of people worldwide, providing essential minerals and nutrients [86]. Given their significance, further research is needed to update inventories and document their availability and use among Bhutanese communities. Involving local communities in conservation and management is also crucial because they are the primary custodians and users of these resources. WEP fairs and food festivals may further strengthen awareness, renew interest among younger generations, and encourage active community participation in resource management. In addition, cultivation and domestication through appropriate modern approaches will be important for maintaining future availability.
Conclusion
This national assessment recorded 114 WEP species in 61 families and shows that WEP use in Bhutan remains important but unevenly distributed. Herbs were the dominant life form, fruits were the most frequently used plant part, and fruits and vegetables remained the core food categories in current use. The results suggest that WEP use is being reconfigured rather than uniformly abandoned: continued use is supported by accessibility, local food traditions, and perceived medicinal value, whereas land-use change, perceived scarcity, and limited species knowledge constrain use. Pressure on both availability and knowledge transmission appears greatest where WEP use depends on forest habitats and younger generations are less engaged in plant identification and use. To support the long-term sustainability of WEPs, the study recommends:
- i. Integrating WEP knowledge into school curricula and community education programs to engage younger generations;
- ii. Supporting community-based conservation and sustainable harvesting initiatives;
- iii. Promoting value-added processing and marketing of WEPs to enhance livelihoods; and
- iv. Strengthening policy support for preserving indigenous knowledge systems.
Together, these actions can help sustain WEP use, preserve cultural identity, strengthen food and nutrition security, and provide a clearer foundation for future research and conservation planning.
Acknowledgments
The authors sincerely thank the forest officials and all respondents for their valuable support and cooperation during data collection. The authors also acknowledge the Jigme Khesar Strict Nature Reserve for granting field clearance and facilitating coordination of field visits and data collection. In addition, the authors are grateful to Yangchenphug Higher Secondary School for providing laboratory facilities for specimen preparation and plant taxonomic identification.
References
- 1. Thakur D, Sharma A, Uniyal SK. Why they eat, what they eat: patterns of wild edible plants consumption in a tribal area of Western Himalaya. J Ethnobiol Ethnomed. 2017;13(1):70. pmid:29233181
- 2. Ghirardini MP, Carli M, del Vecchio N, Rovati A, Cova O, Valigi F, et al. The importance of a taste. A comparative study on wild food plant consumption in twenty-one local communities in Italy. J Ethnobiol Ethnomed. 2007;3:22. pmid:17480214
- 3. Turner NJ, Łuczaj ŁJ, Migliorini P, Pieroni A, Dreon AL, Sacchetti LE, et al. Edible and Tended Wild Plants, Traditional Ecological Knowledge and Agroecology. Critical Reviews in Plant Sciences. 2011;30(1–2):198–225.
- 4. Motti R. Wild Edible Plants: A Challenge for Future Diet and Health. Plants (Basel). 2022;11(3):344. pmid:35161324
- 5. Duguma HT. Wild Edible Plant Nutritional Contribution and Consumer Perception in Ethiopia. Int J Food Sci. 2020;2020:2958623. pmid:32953878
- 6. Hankiso M, Warkineh B, Asfaw Z, Debella A. Ethnobotany of wild edible plants in Soro District of Hadiya Zone, southern Ethiopia. J Ethnobiol Ethnomed. 2023;19(1):21. pmid:37269005
- 7. Jamtsho T, Wangchuk P, Yeshi K, Tobgay, Sridith K. Riparian plant species of Sherichhu River in Eastern Bhutan: their diversity, distribution, and ethnobotanical uses. Trop Ecol. 2021;62(4):563–79.
- 8. Summers RW, Elliott DE, Urban JF Jr, Thompson R, Weinstock JV. Trichuris suis therapy in Crohn’s disease. Gut. 2005;54(1):87–90. pmid:15591509
- 9.
Akhtar F. Uncultivated food in the context of poor people’s livelihood: a Nayakrishi experience. In: Proceedings of the regional workshop on uncultivated foods and biodiversity, 2001.
- 10. Khan WI, Blennerhasset PA, Varghese AK, Chowdhury SK, Omsted P, Deng Y, et al. Intestinal nematode infection ameliorates experimental colitis in mice. Infect Immun. 2002;70(11):5931–7. pmid:12379667
- 11. Maroyi A. The gathering and consumption of wild edible plants in Nhema communal area, Midlands Province, Zimbabwe. Ecol Food Nutr. 2011;50(6):506–25. pmid:22077930
- 12. Shin T, Fujikawa K, Moe AZ, Uchiyama H. Traditional knowledge of wild edible plants with special emphasis on medicinal uses in Southern Shan State, Myanmar. J Ethnobiol Ethnomed. 2018;14(1):48. pmid:30016980
- 13.
Food, Nations AO o t U. The future of food and agriculture: trends and challenges. FAO. 2017.
- 14. Jamtsho T, Sridith K. Species composition of the vegetation along the Sherichhu River, lower montane area of Eastern Bhutan. Songklanakarin J Sci Technol. 2017;93(3).
- 15. Jigme J, Yangchen K. An ethnobotanical study of plants used in socio-religious activities in Bhutan. Asian J Ethnobiol. 2022;5(1).
- 16.
Tshering K, Thapa L, Matsushima K, Minami M, Nemoto K. Edible wild plants of Bhutan and their contribution to food and nutrition security. Regional Symposium on “Promotion of underutilized indigenous food resources for food security and nutrition in Asia and the Pacific”. FAO Regional Office for Asia and the Pacific. 2014.
- 17. Bellows AC, Raj S, Pitstick E, Potteiger MR, Diemont SAW. Foraging Wild Edibles: Dietary Diversity in Expanded Food Systems. Nutrients. 2023;15(21):4630. pmid:37960283
- 18. Ahmed S, Warne T, Stewart A, Byker Shanks C, Dupuis V. Role of Wild Food Environments for Cultural Identity, Food Security, and Dietary Quality in a Rural American State. Front Sustain Food Syst. 2022;6.
- 19. Yangdon P, Araki T, Rahayu YYS, Norbu K. Ethnobotanical study of wild edible fruits in eastern Bhutan. J Ethnobiol Ethnomed. 2022;18(1):27. pmid:35354474
- 20. Wangchuk P, Namgay K, Gayleg K, Dorji Y. Medicinal plants of Dagala region in Bhutan: their diversity, distribution, uses and economic potential. J Ethnobiol Ethnomed. 2016;12(1):28. pmid:27342801
- 21. Smith B, Chinnappa CC. Plant Collection, Identification, and Herbarium Procedures. Plant Microtechniques and Protocols. Springer International Publishing. 2015. p. 541–72.
- 22. Ashagre M, Asfaw Z, Kelbessa E. Ethnobotanical study of wild edible plants in Burji District, Segan Area Zone of Southern Nations, Nationalities and Peoples Region (SNNPR), Ethiopia. J Ethnobiol Ethnomed. 2016;12(1):32. pmid:27485265
- 23. Cao Y, Li R, Zhou S, Song L, Quan R, Hu H. Ethnobotanical study on wild edible plants used by three trans-boundary ethnic groups in Jiangcheng County, Pu’er, Southwest China. J Ethnobiol Ethnomed. 2020;16(1):66. pmid:33109239
- 24. Tareen NM, Rehman M, Shinwari ZK, Bibi T. Ethnomedicinal utilization of wild edible vegetables in district Harnai of Balochistan Province-Pakistan. Pak J Bot. 2016;48(3):1159–71.
- 25. Wang J, Ding X, Guo CA, Zhang X, Feng H, Yang H. An ethnobotanical study of wild edible plants used by the Tibetan in the Rongjia River Valley, Tibet, China. J Ethnobiol Ethnomed. 2023;19(49):2–16.
- 26. Heinrich M, Ankli A, Frei B, Weimann C, Sticher O. Medicinal plants in Mexico: healers’ consensus and cultural importance. Soc Sci Med. 1998;47(11):1859–71. pmid:9877354
- 27. Bhatia H, Sharma YP, Manhas RK, Kumar K. Traditionally used wild edible plants of district Udhampur, J&K, India. J Ethnobiol Ethnomed. 2018;14(1):73. pmid:30486847
- 28. Aryal KP, Poudel S, Chaudhary RP, Chettri N, Chaudhary P, Ning W, et al. Diversity and use of wild and non-cultivated edible plants in the Western Himalaya. J Ethnobiol Ethnomed. 2018;14(1):10. pmid:29378614
- 29.
Kumar A. Research methodology in social science. Sarup & Sons. 2002.
- 30. Pfeiffer JM, Butz RJ. Assessing cultural and ecological variation in ethnobiological research: the importance of gender. J Ethnobiol. 2005;25(2):240–78.
- 31. Cheng Z, Lu X, Lin F, Naeem A, Long C. Ethnobotanical study on wild edible plants used by Dulong people in northwestern Yunnan, China. J Ethnobiol Ethnomed. 2022;18(1):3. pmid:35062974
- 32. Bajpai AB, Koranga N, Srivastava N, Koshal AK, Rana KPS. “Diversity of Wild Edible Plants in the Kotla Valley in Uttarkashi, Uttarakhand, India”. SCIENTIFICTEMPER. 2022;13(2):341–7.
- 33. Thakur A, Singh S, Puri S. Exploration of wild edible plants used as food by Gaddis-a tribal community of the Western Himalaya. Sci World J. 2020;2020(1).
- 34. León-Lobos P, Díaz-Forestier J, Díaz R, Celis-Diez JL, Diazgranados M, Ulian T. Patterns of Traditional and Modern Uses of Wild Edible Native Plants of Chile: Challenges and Future Perspectives. Plants (Basel). 2022;11(6):744. pmid:35336626
- 35. Gakuubi MM, Wanzala W. A survey of plants and plant products traditionally used in livestock health management in Buuri district, Meru County, Kenya. J Ethnobiol Ethnomed. 2012;8(39):1–20.
- 36. Rehman S, Iqbal Z, Qureshi R, Shah GM, Afzal A, Rahman KU. Quantitative ethnomedicinal survey of wild edible fruits used by the indigenous community in North Waziristan, Khyber Pakhtunkhwa, Pakistan. Ethnobot Res Appl. 2024;28:1–20.
- 37. Oluoch WA, Borgemeister C, de Deus Vidal Junior J, Fremout T, Gaisberger H, Whitney C, et al. Predicted changes in distribution and richness of wild edible plants under climate change scenarios in northwestern Kenya. Reg Environ Change. 2024;24(1).
- 38. Oncini F, Hirth S, Mylan J, Robinson CH, Johnson D. Where the wild things are: How urban foraging and food forests can contribute to sustainable cities in the Global North. Urban Forestry & Urban Greening. 2024;93:128216.
- 39. Shumsky SA, Hickey GM, Pelletier B, Johns T. Understanding the contribution of wild edible plants to rural social-ecological resilience in semi-arid Kenya. E&S. 2014;19(4).
- 40. Borelli T, Hunter D, Powell B, Ulian T, Mattana E, Termote C, et al. Born to Eat Wild: An Integrated Conservation Approach to Secure Wild Food Plants for Food Security and Nutrition. Plants (Basel). 2020;9(10):1299. pmid:33019632
- 41. Oduor F, Kaindi DM, Abong G, Thuita F, Termote C. Community-Based Conservation Strategies for Wild Edible Plants in Turkana County, Kenya. Conservation. 2024;5(1):1.
- 42. Yangchen U. Wild vegetable diversity and their contribution to household income at Pat-shaling Gewog, Tsirang. Bhutan Journal of Natural Resources and Development. 2017;4(1):1–10.
- 43. Dorji R, Gyeltshen C, Phuentsho, Wangmo KU, Dorji E. Enumeration of edible and useful ferns (Pteridophytes) in the Bhutan Himalayas. Korean J Pl Taxon. 2022;52(4):235–45.
- 44. Bajgai RC, Bajgai Y, Johnson SB. The presence of wild edible plants and determinants influencing their harvest, consumption, and conservation in south eastern Bhutan. PLoS One. 2023;18(10):e0285936. pmid:37816046
- 45. Dema U, Dolkar J. Diversity and use of wild edible plants in Maenbi Gewog, Lhuentse Dzongkhag. Sherub Doenme: The Research Journal of Sherubtse College. 2022;15(1).
- 46. Łuczaj Ł, Zovkokončić M, Miličević T, Dolina K, Pandža M. Wild vegetable mixes sold in the markets of Dalmatia (southern Croatia). J Ethnobiol Ethnomed. 2013;9:2. pmid:23286393
- 47.
Department AOotUNF. The state of world fisheries and aquaculture. Food and Agriculture Organization of the United Nations. 2018.
- 48. Asfaw A, Lulekal E, Bekele T, Debella A, Tessema S, Meresa A, et al. Ethnobotanical study of wild edible plants and implications for food security. Trees, Forests and People. 2023;14:100453.
- 49. Mukhtar E, Suwardi AB, Nzi S, Syamsuardi TH, M E. The diversity of wild edible fruit plants and traditional knowledge in West Aceh region, Indonesia. J Med Plants Stud. 2016;7(4):285–90.
- 50. Singh A, Dubey RK, Bundela AK, Abhilash PC. The Trilogy of Wild Crops, Traditional Agronomic Practices, and UN-Sustainable Development Goals. Agronomy. 2020;10(5):648.
- 51. Konsam S, Thongam B, Handique AK. Assessment of wild leafy vegetables traditionally consumed by the ethnic communities of Manipur, northeast India. J Ethnobiol Ethnomed. 2016;12(9):1–15.
- 52. Zhang L, Chai Z, Zhang Y, Geng Y, Wang Y. Ethnobotanical study of traditional edible plants used by the Naxi people during droughts. J Ethnobiol Ethnomed. 2016;12(1):39. pmid:27619157
- 53. Grivetti LE, Ogle BM. Value of traditional foods in meeting macro- and micronutrient needs: the wild plant connection. Nutr Res Rev. 2000;13(1):31–46. pmid:19087432
- 54. Bhandari MR, Kasai T, Kawabata J. Nutritional evaluation of wild yam (Dioscorea spp.) tubers of Nepal. Food Chem. 2003;82(4):619–23.
- 55. Serrasolses G, Calvet-Mir L, Carrió E, D’Ambrosio U, Garnatje T, Parada M. A matter of taste: Local explanations for the consumption of wild food plants in the Catalan Pyrenees and the Balearic Islands. Econ Bot. 2016;70:176–89.
- 56. Ojelel S, Mucunguzi P, Katuura E, Kakudidi EK, Namaganda M, Kalema J. Wild edible plants used by communities in and around selected forest reserves of Teso-Karamoja region, Uganda. J Ethnobiol Ethnomed. 2019;15(1):1–14.
- 57. Ladio AH, Lozada M. Patterns of use and knowledge of wild edible plants in distinct ecological environments: a case study of a Mapuche community from northwestern Patagonia. Biodivers Conserv. 2004;13:1153–73.
- 58. Sõukand R. Perceived reasons for changes in the use of wild food plants in Saaremaa, Estonia. Appetite. 2016;107:231–41. pmid:27521164
- 59. Abdullah A, Khan SM, Pieroni A, Haq A, Haq ZU, Ahmad Z, et al. A Comprehensive Appraisal of the Wild Food Plants and Food System of Tribal Cultures in the Hindu Kush Mountain Range; a Way Forward for Balancing Human Nutrition and Food Security. Sustainability. 2021;13(9):5258.
- 60. Sansanelli S, Ferri M, Salinitro M, Tassoni A. Ethnobotanical survey of wild food plants traditionally collected and consumed in the Middle Agri Valley (Basilicata region, southern Italy). J Ethnobiol Ethnomed. 2017;13(1):50. pmid:28874202
- 61. Sardeshpande M, Shackleton C. Wild Edible Fruits: A Systematic Review of an Under-Researched Multifunctional NTFP (Non-Timber Forest Product). Forests. 2019;10(6):467.
- 62. Pieroni A. Evaluation of the cultural significance of wild food botanicals traditionally consumed in northwestern Tuscany, Italy. J Ethnobiol. 2001;21(1):89–104.
- 63. Schunko C, Grasser S, Vogl CR. Explaining the resurgent popularity of the wild: motivations for wild plant gathering in the Biosphere Reserve Grosses Walsertal, Austria. J Ethnobiol Ethnomed. 2015;11:55. pmid:26122103
- 64. Aceituno-Mata L, Tardío J, Pardo-de-Santayana M. The persistence of flavor: Past and present use of wild food plants in Sierra Norte de Madrid, Spain. Frontiers in Sustainable Food Systems. 2021;4:610238.
- 65. Pawera L, Khomsan A, Zuhud EAM, Hunter D, Ickowitz A, Polesny Z. Wild Food Plants and Trends in Their Use: From Knowledge and Perceptions to Drivers of Change in West Sumatra, Indonesia. Foods. 2020;9(9):1240. pmid:32899857
- 66. Turreira-García N, Theilade I, Meilby H, Sørensen M. Wild edible plant knowledge, distribution and transmission: a case study of the Achí Mayans of Guatemala. J Ethnobiol Ethnomed. 2015;11:52. pmid:26077151
- 67. Barreau A, Ibarra JT, Wyndham FS, Rojas A, Kozak RA. How Can We Teach Our Children if We Cannot Access the Forest? Generational Change in Mapuche Knowledge of Wild Edible Plants in Andean Temperate Ecosystems of Chile. Journal of Ethnobiology. 2016;36(2):412–32.
- 68. Serrasolses G, Calvet-Mir L, Carrió E, D’Ambrosio U, Garnatje T, Parada M, et al. A Matter of Taste: Local Explanations for the Consumption of Wild Food Plants in the Catalan Pyrenees and the Balearic Islands1. Econ Bot. 2016;70(2):176–89.
- 69. Guarrera PM, Savo V. Wild food plants used in traditional vegetable mixtures in Italy. J Ethnopharmacol. 2016;185:202–34. pmid:26944238
- 70. Kidane L, Kejela A. Food security and environment conservation through sustainable use of wild and semi-wild edible plants: a case study in Berek Natural Forest, Oromia special zone, Ethiopia. Agric & Food Secur. 2021;10(1).
- 71. Uprety Y, Poudel RC, Shrestha KK, Rajbhandary S, Tiwari NN, Shrestha UB, et al. Diversity of use and local knowledge of wild edible plant resources in Nepal. J Ethnobiol Ethnomed. 2012;8:16. pmid:22546349
- 72. Hickey GM, Pouliot M, Smith-Hall C, Wunder S, Nielsen MR. Quantifying the economic contribution of wild food harvests to rural livelihoods: A global-comparative analysis. Food Policy. 2016;62:122–32.
- 73. Schunko C, Vogl CR. Organic farmers use of wild food plants and fungi in a hilly area in Styria (Austria). J Ethnobiol Ethnomed. 2010;6:17. pmid:20565945
- 74. Pardo-de-Santayana M, Tardío J, Morales R. The gathering and consumption of wild edible plants in the Campoo (Cantabria, Spain). Int J Food Sci Nutr. 2005;56(7):529–42. pmid:16503563
- 75. Shukla V, Pala NA, Manohar A, Puri A, Chakravorty S. Bioprospecting wild plant resources to ensure food and nutritional security. Recent trends & advances in food science & post harvest technology.
- 76.
Dorji Y. Women’s roles in wild yam, conservation, management and use in Bhutan. Gender and biodiversity management in the Greater Himalayas. ICIMOD. 2012. p. 27–51.
- 77. Jigme J, Yangchen K. Ethnobotanical study of the wild edible plants used by the indigenous people of Merak in Bhutan. Asian J Ethnobiol. 2023;6(1).
- 78. Mishra A, Swamy SL, Thakur TK, Bhat R, Bijalwan A, Kumar A. Use of Wild Edible Plants: Can They Meet the Dietary and Nutritional Needs of Indigenous Communities in Central India. Foods. 2021;10(7):1453. pmid:34201552
- 79. Fratianni A, Albanese D, Ianiri G, Vitone C, Malvano F, Avino P, et al. Evaluation of the Content of Minerals, B-Group Vitamins, Tocols, and Carotenoids in Raw and In-House Cooked Wild Edible Plants. Foods. 2024;13(3):472. pmid:38338607
- 80. Oluoch WA, Whitney CW, Termote C, Borgemeister C, Schmitt CB. Integrated participatory approach reveals perceived local availability of wild edible plants in northwestern Kenya. Hum Ecol. 2023;51(1):59–74.
- 81. Steel EA, Bwembelo L, Mulani A, Siamutondo ALM, Banda P, Gumbo D, et al. Wild foods from forests: Quantities collected across Zambia. People and Nature. 2022;4(5):1159–75.
- 82.
Oluoch WA. Availability and sustainable harvesting of wild edible plants in Turkana County, Kenya. Universitäts-und Landesbibliothek Bonn. 2024.
- 83.
Nepal TK. Keeping the local tradition alive: the status of local traditional knowledge in Dorokha, Samtse, Bhutan. 2023.
- 84. Ugyen J, Gurung T. Assessment of yam (Dioscorea spp.) diversity at community level in Nangkor gewog under Zhemgang dzongkhag. Journal of Bhutan Studies. 2015;33:55–73.
- 85. Matsushima KI, Minami M, Nemoto K. Use and conservation of edible wild plants in Bhutan. J Fac Agr Kyushu U. 2012;48(1–2).
- 86. Pieroni A, Sõukand R. The disappearing wild food and medicinal plant knowledge in a few mountain villages of North-Eastern Albania. J Appl Bot Food Qual. 2017;90:58–67.