Received: 23 April 2020  |  Revised: 12 June 2020  |  Accepted: 15 June 2020
DOI: 10.1002/ppp3.10145  
R E V I E W  A R T I C L E
Unlocking plant resources to support food security and 
promote sustainable agriculture
Tiziana Ulian1  |   Mauricio Diazgranados1 |   Samuel Pironon2 |   Stefano Padulosi3 |   
Udayangani Liu1 |   Lee Davies2 |   Melanie-Jayne R. Howes2,4  |   James S. Borrell2  |   
Ian Ondo2 |   Oscar A. Pérez-Escobar2 |   Suzanne Sharrock5 |   Philippa Ryan2 |   
Danny Hunter3 |   Mark A. Lee2  |   Charles Barstow6 |   Łukasz Łuczaj7 |    
Andrea Pieroni8,9 |   Rodrigo Cámara-Leret10  |   Arshiya Noorani11 |   Chikelu Mba11 |   
Rémi Nono Womdim11 |   Hafiz Muminjanov11 |   Alexandre Antonelli2,12  |    
Hugh W. Pritchard1  |   Efisio Mattana1
1Royal Botanic Gardens Kew, Ardingly, UK
2 Societal Impact Statement
Royal Botanic Gardens Kew, Richmond, UK
3Alliance of Bioversity International and Biodiversity is essential to food security and nutrition locally and globally. By review-
CIAT, Rome, Italy ing the global state of edible plants and highlighting key neglected and underutilized 
4Institute of Pharmaceutical Science, Faculty species (NUS), we attempt to unlock plant food resources and explore the role of 
of Life Sciences & Medicine, King's College 
London, London, UK fungi, which along with the wealth of traditional knowledge about their uses and 
5Botanic Gardens Conservation practices, could help support sustainable agriculture while ensuring better protection 
International, Richmond, UK of the environment and the continued delivery of its ecosystem services. This work 
6Slow Food International, Bra, Italy
7 will inform a wide range of user communities, including scientists, conservation and 
Institute of Biology and Biotechnology, 
University of Rzeszów, Rzeszów, Poland development organizations, policymakers, and the public of the importance of biodi-
8University of Gastronomic Sciences, versity beyond mainstream crops.
Pollenzo/Bra, Italy
Summary 
9Department of Medical Analysis, Tishk 
International University, Erbil, Kurdistan, As the world's population is increasing, humanity is facing both shortages (hunger) 
Iraq and excesses (obesity) of calorie and nutrient intakes. Biodiversity is fundamental to 
10Department of Evolutionary Biology and 
Environmental Studies, University of Zurich, addressing this double challenge, which involves a far better understanding of the 
Zurich, Switzerland global state of food resources. Current estimates suggest that there are at least 7,039 
11Food and Agricultural Organization of the edible plant species, in a broad taxonomic sense, which includes 7,014 vascular plants. 
United Nations, Rome, Italy
12 This is in striking contrast to the small handful of food crops that provide the major-
Gothenburg Global Biodiversity Centre 
and Department of Biological and ity of humanity's calorie and nutrient intake. Most of these 7,039 edible species have 
Environmental Sciences, University of additional uses, the most common being medicines (70%), materials (59%), and envi-
Gothenburg, Göteborg, Sweden
ronmental uses (40%). Species of major food crops display centers of diversity, as pre-
Correspondence viously proposed, while the rest of edible plants follow latitudinal distribution patterns 
Tiziana Ulian, Royal Botanic Gardens Kew, 
Wellcome Trust Millennium Building, similarly to the total plant diversity, with higher species richness at lower latitudes. 
Wakehurst, Ardingly, West Sussex RH17 The International Union for Conservation of Nature Red List includes global conser-
6TN, UK.
Email: t.ulian@kew.org vation assessments for at least 30% of edible plants, with ca. 86% of them conserved 
This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, 
provided the original work is properly cited.
© 2020 The Authors, Plants, People, Planet © New Phytologist Foundation
Plants, People, Planet. 2020;2:421–445.  wileyonlinelibrary.com/journal/ppp3  |  421
422  |     ULIAN et AL.
Funding information
the Swedish Research Council; the Swedish ex situ. However, at least 11% of those species recorded are threatened. We highlight 
Foundation for Strategic Research; the Knut multipurpose NUS of plants from different regions of the world, which could be key 
and Alice Wallenberg Foundation
for a more resilient, sustainable, biodiverse, and community participation-driven new 
“green revolution.” Furthermore, we explore how fungi could diversify and increase 
the nutritional value of our diets. NUS, along with the wealth of traditional knowl-
edge about their uses and practices, offer a largely untapped resource to support 
food security and sustainable agriculture. However, for these natural resources to be 
unlocked, enhanced collaboration among stakeholders is vital.
K E Y W O R D S
crops, ex situ conservation, fungi, livelihoods, minor crops, neglected and underutilized 
species, plant diversity, sustainable agriculture
1  | INTRODUC TION the Green Revolution of the 1960s–1980s. This period of crop inten-
sification was also aided by developments in the use of chemical fer-
As the world's population is expected to reach 10 billion by 2050, hu- tilizers, irrigation techniques, and pesticides (Pingali, 2012). Although 
manity is increasingly facing a double burden of malnutrition, com- the intensification of agriculture led to reduced pressure on natural 
prising of a shortage of calories (hunger) at one end of the spectrum ecosystems (Godfray et al., 2010; Green, Cornell, Scharlemann, & 
and excess at the other one (obesity; Abarca-Gómez et al., 2017; Balmford, 2005), it created multiple unintended environmental con-
Alexandratos & Bruinsma, 2012; FAO, IFAD, UNICEF, WFP, & WHO, sequences such as water pollution, soil degradation, agrochemical 
2019). Addressing these challenges will require an increase of food runoff, increased susceptibility to pests and diseases, and biodiver-
production globally, which cannot be achieved by simply expand- sity loss (Pingali, 2012). Crop intensification also decreased dietary 
ing industrial agriculture through land conversion to the detriment diversity along with food cultures, and many traditional crops that 
of the surrounding environment and already declining biodiversity were important sources of critical micronutrients (such as iron, 
(Jacobsen, Sørensen, Pedersen, & Weiner, 2013; Padulosi, Heywood, provitamin A, and zinc) for poor communities were lost (Webb & 
Hunter, & Jarvis, 2011; Sunderland, 2011), and a shift to health- Eiselen, 2009). However, there is now increasing recognition given 
ier diets (Abarca-Gómez et al., 2017; FAO, IFAD, UNICEF, WFP, & to the importance of biodiversity for food and nutrition security, 
WHO, 2019). In addition, around 36% (by calorific value) of arable local livelihoods, and sustainable development (Bala, Hoeschle-
crops such as wheat, maize, and sorghum are consumed by live- Zeledon, Swaminathan, & Frison, 2006; FAO, 2019). Consequently, 
stock and this requires one-third of the total area currently utilized the benefits of using underutilized traditional crops, and exploring 
for arable farming (Cassidy, West, Gerber, & Foley, 2013; Herrero more sustainable production methods to grow mainstream crops, 
et al., 2013). Overall, 26% (3.4 billion ha) and 4% (0.5 billion ha) of the are being widely promoted (FAO & WHO, 2018).
Earth's ice-free surface is used for livestock grazing and livestock Neglected and underutilized species (NUS) include wild, domes-
feed production, respectively (Foley et al., 2011). This is a complex ticated, or semi-domesticated plants, whose potential to improve 
situation, as there is a need to ensure the sustainable production of people's livelihoods, as well as food security and sovereignty, is not 
safe and nutritious food, while protecting biodiversity, to allow the fully realized because of their limited competitiveness with com-
delivery of other goods and ecosystem services, which are directly modity crops in mainstream agriculture. Nevertheless, they are lo-
and indirectly critical for human well-being. Furthermore, it is neces- cally important to people and often adapted to unique climatic and 
sary to facilitate societal adaptation to climate-driven environmental environmental conditions (Padulosi et al., 2011). Bringing NUS into 
changes that can disrupt food production and people's livelihoods mainstream agriculture could strengthen the resilience and sustain-
(Alae-Carew et al., 2020; FAO, 2019; Jacobsen et al., 2013). ability of food production systems (FAO, 2018; Padulosi, Cawthorn, 
Of the thousands of plant species that have been cultivated since et al., 2019; Raneri, Padulosi, Meldrum, & King, 2019). In addition, 
agriculture began around 12,000 years ago, only about 200 have NUS often provide benefits beyond food, by virtue of being multi-
been extensively domesticated, leading to dependence on a narrow purpose. For instance, they often yield other useful products such 
range of genetic diversity of crops (Meyer, Duval, & Jensen, 2012; as timber, fibers, or medicines, and contribute to safeguarding bio-
Vaughan, Balazs, & Heslop-Harrison, 2007). Together, wheat, rice, cultural diversity (Cámara-Leret et al., 2019). Increasing the inherent 
and maize alone provide almost half of the world's food calorie in- value of wild species as NUS and the ecosystem services that na-
take, making our food supply extremely vulnerable (Reeves, Thomas, tive species can provide to surrounding environments (such as food 
& Ramsay, 2016). Plant-breeding programs narrowed the focus to sources for pollinators and birds, maintenance of water supply and 
large-seeded, high-yielding varieties of crops (Gruber, 2017), whose soils, and control of pests and diseases), will support biodiversity 
global production intensified (higher yield by unit of land area) during protection and provide cultural services (Díaz et al., 2020). Many 
ULIAN et AL.      |  423
NUS are referred to as “minor” or “orphan” crops because of their analyzed in this review, were not treated separately, as several stud-
limited role in larger agricultural production systems and have been ies are already available on their richness, global distribution, and 
“neglected” by agricultural researchers, plant breeders, and policy- conservation, for example in Castañeda-Álvarez et al. (2016) and 
makers alike. Some have been major crops in the past, but are now Milla (2020). Species from Diazgranados et al. (2020) that were also 
displaced by modern commercial varieties and this is especially the listed in Annex 5 of FAO (2015) were identified as “major food crops” 
case for many millets (which is a common term for a group of cereals in this review. Plant uses were classified according to the Level 1 of 
in the Panicodeae and Chlorideae grass subfamilies) and less well- Uses of the Economic Botany Data Collection Standard (Cook, 1995), 
known pulses such as lablab (Reed & Ryan, 2019). Many of these simplified to 10 categories, as in Diazgranados et al. (2020).
varieties and species, along with a wealth of traditional knowledge 
about their use and cultivation, are being lost at an alarming rate (Díaz 
et al., 2020). Access to NUS is also important because domesticated 2.1 | Taxonomic diversity
legumes (Fernández-Marín et al., 2014), cereals (Hebelstrup, 2017), 
other crops that contribute to food security (Tamrat et al., 2020), Depending on authority, the total number of edible plants in the 
and fungi (Stojković et al., 2013) can vary in their nutritional, antioxi- world varies from 100s (Van Wyk, 2019) to >30,000 plants, includ-
dant, and other chemical content. This has potential implications for ing infraspecific taxa (French, 2019). These differences in numbers 
human health, which could be positive or negative, as for example are based on multiple factors, such as taxonomic rank (e.g., counting 
on the diversity of the human intestinal microbiome (e.g., Albenberg infraspecific taxa), accuracy (e.g., using reconciled taxonomy), and 
& Wu, 2014). precision (e.g., using a unique taxonomic backbone), as well as the 
As global biodiversity is rapidly declining, limiting our possibili- types of consumers and their diets. For example, using a conserva-
ties of finding new food sources (Díaz et al., 2020), and considering tive approach based on reported uses, RBG Kew has recorded to 
that most analyses lack information on the entire spectrum of food date 7,039 edible species, in a broad taxonomic sense, from 288 fam-
resources consumed across the world, an assessment of their current ilies and 2,319 genera, including 7,030 edible species of Bryophyta, 
distribution and conservation status to inform science-based policy Chlorophyta, Rhodophyta, and Tracheophyta (Diazgranados 
making has become urgent. In addition, the adverse impacts of cli- et al., 2020). Many more edible species are expected to be identified 
mate change on biodiversity, agricultural production, and food se- in the future, as under-documented regions, for example, tropical 
curity have made the conservation of food diversity and associated America and New Guinea, are better characterized (Cámara-Leret 
traditional knowledge a global priority (Corlett, 2016; Maxted, Ford- & Dennehy, 2019; Cámara-Leret, Paniagua-Zambrana, Balslev, & 
Lloyd, Jury, Kell, & Scholten, 2006; Vincent et al., 2013). Finally, as Macía, 2014). Recognizing variation within species (subspecies, lan-
intact habitats come under pressure from the increased demand for draces, etc.) is equally important. While Brassica oleracea is known to 
cropland worldwide (Tilman et al., 2017), ex situ plant conservation cover nine crops, the level of plant diversity in use can be obscured 
measures need accelerating (Larkin, Jacobi, Hipp, & Kramer, 2016), by the widespread use of a common name (e.g., “beans” apply to at 
as promoted in the UN Sustainable Development Goal (SDG) Target least 17 genera, 30 species, and thousands of varieties).
2.5 (https://susta inabl edeve lopme nt.un.org/). Vascular plants (Tracheophyta) are the most important for 
In this article, we (a) consider the global state of edible plants, human food, encompassing 272 families, 2,300 genera, and 7,014 
their taxonomic diversity, uses, distribution, and conservation sta- known species, that is, 2.0% of the total angiosperm species diver-
tus; and (b) explore untapped plant and fungi resources, by review- sity (347,298 accepted species; WCVP, 2020). Sixty percent of the 
ing the role of multipurpose NUS that could be adopted as potential vascular plant families include edible species, covering almost all the 
future food crops under a changing climate. major phylogenetic clades (Figure 1). The most diverse orders are 
Fabales (640 edible species), Malpighiales (550), Sapindales (465), 
Gentianales (444), and Rosales (395). The richest families (see Figure 
2  | THE GLOBAL STATE OF EDIBLE S1) are Fabaceae (i.e., beans, 625 edible species), Arecaceae (palms, 
PL ANTS AND MA JOR FOOD CROPS 325), Poaceae (grasses and includes cereals, 314), Malvaceae (mal-
low family, includes cacao, okra and durian, 257), and Asteraceae 
To assess the global diversity of edible plants we used the “World (sunflower and lettuce families, 251). With at least 100 edible spe-
Checklist of Useful Plant Species” data set, produced by the Royal cies, Ficus (figs) is the richest genus, followed by Diospyros (52), 
Botanic Gardens, Kew (Diazgranados et al., 2020). This data set in- Solanum (51), Garcinia (48), and Grewia (46). Most of the edible plants 
cludes 40,292 species with at least a documented human use and (97%) correspond to flowering plants, with 245 families, 2,235 gen-
was redacted by compiling plant uses and reconciling species names era, and 6,828 species. However, there is substantial variation in the 
using the taxonomic backbone of Kew's Plants of the World Online proportion of edible plants among non-flowering plant groups, for 
portal (http://www.plant softh eworl donli ne.org/) from 13 large example, 0.5% (six species) of Lycopodiopsida, 1.0% (109 species) 
datasets, listed in Diazgranados et al. (2020). Species with “human of Polypodiopsida, 3.9% (13 species) of cycads, 7.5% (47 species) of 
food” use in this list were extracted and analyzed in this review as Pinopsida, 8.9% (10 species) of Gnetopsida, and 100% (one species) 
“edible plants.” Crop wild relatives, although included in the list and of ginkgo.
424  |     ULIAN et AL.
F I G U R E  1   Phylogenetic distribution of edible plants from Diazgranados et al. (2020), and major food crops also listed in FAO (2015). A 
phylogeny of 448 vascular plant families was derived from the Spermatophyta supertree inferred from sequence data of 79,881 species by 
Smith and Brown (2018) by keeping one representative species per plant family. Presence/absence of edible plants and major food crops 
per family was drawn at the tips of the phylogeny using the R-package GGTREE (Yu, Smith, Zhu, Guan, & Lam, 2017). The rectangles at the 
tips of the phylogeny denote the presence of human food plants (orange) and major food crops (brown) in each family. Major plant clades are 
color-coded, except for clades with just a few families, indicated with numbers: 1. Chloranthales (1 family); 2. Ceratophyllales (1); 3. Proteales 
(4), Trochodendrales (1), Buxales (1) and Gunnerales (2); 4. Dilleniales (1 fam.); and 5. Berberidopsidales (2). Please see Figure S1 in the 
Supporting Information for the detailed tree with the names in the tips for all families
To understand the taxonomic distribution of intensively used where practices of domestication of those species are not known); 
food plants, we mapped the major food crop species listed by the or simply because it was not needed (e.g., high abundance of man-
FAO (2015) onto the phylogeny (Figure 1). Only 417 (5.9%) of edi- grove trees in the Rhizophoraceae, which provide food among other 
ble plant species from Diazgranados et al. (2020), belonging to 168 uses, may be sufficient for the local demand). Some families, such 
(7.2%) genera and 62 (21.5%) families, appear in the FAO (2015) list. as Acanthaceae and Phyllanthaceae, have a few species under cul-
The richest families in major crops are Rosaceae (e.g., apples; 51 spe- tivation but these are not used as food (e.g., as ornamental plants). 
cies/eight genera), Fabaceae (51 species/19 genera), Dioscoreaceae Lastly, 77 plant families have one or two edible species which are 
(i.e., yams; 41 species/one genus), Poaceae (27 species/16 genera), not crops.
and Malvaceae (21 species/eight genera). Interestingly, several edible 
species-rich families have very few major crops, or none. For exam-
ple, Arecaceae (325 species) has only six crop species, Apocynaceae 2.2 | Plant uses
(228 spp.) just one, and Phyllanthaceae (101 species) none. The rea-
sons for the low (or absent) domestication rate detected in some Edible plants often have additional uses, which may differ in the 
families may include: habit (e.g., tall trees/palms from the tropics; world as part of the existing cultural diversity. The most frequent 
parasitic families such as Balanophoraceae or Loranthaceae); high use is medicines (70% of species), followed by materials (59%), en-
toxicity (e.g., many Apocynaceae bear edible fruits, but all other vironmental uses (40%), gene sources (i.e., wild relatives of major 
parts are poisonous); habitat specificity (e.g., plants adapted to crops which may possess traits associated to biotic or abiotic resist-
harsh weather, making difficult to establish crops); low growth rate ance and therefore be valuable for breeding programs; Cook, 1995; 
(e.g., many woody plants); spatial distribution (e.g., plants from areas 32%), and animal food (30%; Figure 2a). The same general trend 
ULIAN et AL.      |  425
F I G U R E  2   Heat map showing the 
proportion of plant species in each 
additional use category for (a) edible plant 
species from Diazgranados et al. (2020) 
and (b) species of major food crops also 
listed in FAO (2015). The dendrogram 
represents a hierarchical clustering of 
the uses: clustered uses indicate closer 
proportion pattern, using the Euclidian 
distance for building the distance matrix 
and the “Complete-linkage” method 
for the hierarchical aggregation of the 
dendrogram
was identified for species of major crops, with 83% also reported plant diversity (Kier et al., 2009). Although a major hotspot of plant 
as “medicinal,” and with “gene source” having higher weight (70%) species richness, tropical Americas is under-represented in terms 
than in the full list of edible plants (Figure 2b). The link between of edible plants. This highlights a likely spatial bias in Diazgranados 
food and medicine is well documented (e.g., Iwu, 2016), and al- et al. (2020) toward other tropical, and better investigated, areas of 
ready demonstrated for plant-rich diets, such as the traditional the world, for which information is databased and accessible, such as 
Mediterranean diet (Willett et al., 1995). Livestock and wild animals Africa, which is well represented in the Plant Resources for Tropical 
can also make use of the medicinal properties of plants to improve Africa (PROTA) database (https://www.prota 4u.org/database).
or maintain their health, for example, to control internal parasites The native distribution of some of the major food crop plant 
(Villalba & Provenza, 2007; Villalba, Provenza, K Clemensen, Larsen, species from FAO (2015; Figure 3b) generally maps over Vavilov's 
& Juhnke, 2011). Indeed, the boundaries between foods, including centers of diversity (Vavilov, Vavylov, Vavílov, & Dorofeev, 1992), 
functional foods, medicine, and nutraceuticals are often blurred, at- that is, the Mediterranean, Middle East, and Central Asia (for wheat, 
tributed to certain phytochemicals in edible plants that have mecha- lentils, peas, artichokes, apples), Ethiopia/Eritrea highlands (for teff, 
nistic effects relevant to human health, independent of fundamental Arabica coffee, enset), India (for aubergines, pigeon pea, mangoes), 
nutrition (Howes, 2018b; Howes et al., 2020; Paradee et al., 2019). East Asia (for soybean, Asian rice, oranges, peaches), Mesoamerica, 
Certain edible plants and their constituents are associated with a and the Andes (for maize, chillies, common bean, tomatoes, pota-
reduced risk of some diseases. For example, there has been interest toes). However, there is a relatively low species richness in major 
in the role of cruciferous vegetables and turmeric (Curcuma longa) to food crops from the Malay Archipelago and high edible species rich-
reduce cancer risk (Howes, 2018a), while Perilla frutescens nutlets ness from parts of Sub-Saharan Africa. Additional centers of origin 
have been evaluated to provide protection against oxidative stress in have been proposed in recent years based on new archaeological ev-
some hepatic disorders (Paradee et al., 2019). This concept extends idence, such as West Africa for pearl millet and cowpea and Eastern 
to livestock and there is emerging evidence that the phytochemi- Sahel for sorghum (Fuller et al., 2014; Harlan, 1971; Purugganan & 
cal composition of animal feed can enhance meat and dairy prod- Fuller, 2009).
ucts, which may reduce the incidence of some diseases in humans There is a geographical spectrum to food plant domestication, 
(Provenza, Kronberg, & Gregorini, 2019). with total food plant richness mostly in the tropics and major do-
mestication events more scattered at mid-latitudes, following a 
global pattern associated with environmental and historical fac-
2.3 | Global distribution tors (Diamond, 2002). The proportion of highly domesticated spe-
cies increases from species-rich, forested, warm, and wet areas 
We found the native distribution of the large array of edible plant to drier climates, rugged terrains (i.e., mountainous areas exhib-
species documented in Diazgranados et al. (2020) to exhibit a clear iting high heterogeneity in environmental conditions), and large 
latitudinal gradient, with food plant species richness decreasing from human settlements developing agriculture (Lev-Yadun, Gopher, & 
low to high latitudes (Figure 3a), similarly to general patterns in total Abbo, 2000; Meyer et al., 2012; Vavilov et al., 1992). In contrast, 
426  |     ULIAN et AL.
F I G U R E  3   (a) Global species richness 
per country of 6,959 out of the 7,039 
edible species from Diazgranados 
et al. (2020). (b) Global species richness 
per country of 171 out of the 417 major 
food crops also listed in FAO (2015). 
While edible species richness decreases 
with increasing latitude, high richness 
in major food crops is mainly found in 
centers of domestication at mid-latitudes. 
Maps include species for which an IPNI 
ID (https://www.ipni.org/), as well as 
countries and sub-countries distribution 
data from the World Checklist of Selected 
Plant Families (WCVP, 2020), were 
available
wild, species-poor, cold, and flat areas of high latitudes contain few collections housed in botanic gardens are included, we find a sub-
highly domesticated plants. However, humans are now changing stantial representation of edible plant species conserved ex situ 
these spatial patterns in food supply, demand, and cultivation by ho- worldwide (Table 1). These results were achieved thanks to the 
mogenizing the distribution of both agro-biodiversity and biodiver- joint efforts of the international CGIAR genebanks (https://www.
sity in general (Baiser, Olden, Record, Lockwood, & McKinney, 2012; cgiar.org/), botanic gardens (https://www.bgci.org/), and interna-
Khoury et al., 2014). tional plant conservation networks, such as Kew's Millennium Seed 
Understanding better the global distribution of edible plants of- Bank Partnership (Liu, Breman, Cossu, & Kenney, 2018). However, 
fers an opportunity to identify future crops that are better adapted some food species might be missing from ex situ collections due 
to present and future climatic conditions, and whose plant material to incomplete data sets, geographic rarity, and having recalcitrant 
is locally accessible. This could improve food security by increas- (i.e., desiccation sensitive) seeds, such as some tropical fruit trees 
ing the cultivation of “climate smart” crops with fit-for-purpose (Li & Pritchard, 2009) and some priority crops on Annex 1 of the 
seed lots (Castillo-Lorenzo, Pritchard, Finch-Savage, & Seal, 2019) “International Treaty on Plant Genetic Resources for Food and 
that will produce food despite changing growing conditions (Borrell Agriculture” (FAO, 2009). More work is also needed to understand 
et al., 2020; Díaz et al., 2019; Pironon et al., 2019). and evaluate the functional and genetic diversity of ex situ collec-
tions, their potential for reintroduction efforts (Hay & Probert, 2013) 
and adaptability to future climate change (Borrell et al., 2020; 
2.4 | Conservation status and measures in place Fernández-Pascual, Mattana, & Pritchard, 2019).
The International Union for Conservation of Nature (IUCN) Red 
Previous studies on the comprehensiveness of the conservation of List (IUCN, 2020) includes species-level global conservation assess-
useful plants have highlighted that they are currently highly under- ments for at least 2,108 (30%) edible species listed in Diazgranados 
conserved, both ex situ and in situ (Castañeda-Álvarez et al., 2016; et al. (2020) and 1,811 of these (86%) are conserved ex situ (Table 2). 
Fielder et al., 2015; Khoury et al., 2019). However, when the Although most species (78%) are identified as Least Concern, at least 
ULIAN et AL.      |  427
TA B L E  1   Taxonomic representation of 
food plant species in ex situ conservation Total number conserved ex situ
Total number in the 
facilities worldwide as seeds or/and as Taxonomy reference list Seedsa  Unspecified Plantsb  Overall
living plants
Class 8 8 8 6 8 (100%)
Order 69 62 68 43 69 (100%)
Families 272 231 254 127 263 (97%)
Genera 2,300 1,573 1,834 556 2,016 (88%)
Species 7,014 3,810 4,789 1,100 5,454 (78%)
aAs populations/seed lots; 
bCould be limited to a few individuals. 
Sources: RBG Kew's MSB Partnership (https://www.kew.org/scien ce/our-scien ce/proje cts/banki 
ng-the-world s-seeds), Genesys (https://www.genes ys-pgr.org/) and Botanic Gardens Conservation 
International (https://tools.bgci.org/plant_search.php)
TA B L E  2   Current conservation status and ex situ conservation landraces, which may have unique climatic and environmental toler-
measures for food plant species ances, and upon which human communities may depend, might still 
be threatened. Therefore, future conservation priorities should re-
Total number of species flect assessments at the global level, and, for narrow distributed spe-
The IUCN cies, at the national level (Forest et al., 2018; Liu, Kenney, Breman, 
Red List Conserved Conserved & Cossu, 2019).
IUCN category (IUCN, 2020) ex situ (N) ex situ (%)
Extinct in the Wild 1 1 100
(EW) 3  | UNTAPPED PL ANT FOOD RESOURCES
Critically 23 17 74
Endangered (CR)
Beyond habitat destruction, many NUS are at risk of disappearing be-
Endangered (EN) 68 53 78 cause of changing cultural views and lack of documentation (National 
Vulnerable (VU) 142 94 66 Research Council, 2008). Promoting their role in food security calls for 
Near Threatened 64 53 83 coordinated approaches across plant science and food systems, from 
(NT) local to international levels (Baldermann et al., 2016), as actively pro-
Lower Risk/ 5 5 100 moted since 1988 by the International Centre for Underutilized Crops 
conservation 
dependent (LR/cd) (Tchoundjeu & Atangana, 2006). However, consolidated attention to 
Lower Risk/near 35 24 69 NUS has really only emerged in the last decade, as the fight against 
threatened (LR/nt) climate change and the need to make agricultural production systems 
Lower Risk/least 52 42 81 more diverse and environment resilient has accelerated (see Table S1 
concern (LR/lc) for a selection of projects/initiatives). The same trend is also evident 
Least Concern (LC) 1,656 1,468 89 for the limited pool of human and animal food crops, for which the chal-
Data Deficient (DD) 62 54 87 lenges of feeding a growing population with a limited pool of crops have 
Total 2,108 1,811 86 been highlighted (Lee, 2018; Lee, Davis, Chagunda, & Manning, 2017).
Sources: RBG Kew's MSB Partnership (https://www.kew.org/ There are many incentives and subsidies that tie countries into 
scien ce/our-scien ce/proje cts/banki ng-the-world s-seeds), Genesys the production of major crops (Hunter et al., 2019; Noorani, Bazile, 
(https://www.genes ys-pgr.org/), and Botanic Gardens Conservation Diulgheroff, Kahane, & Nono-Womdim, 2015) and which potentially 
International (https://tools.bgci.org/plant_search.php)
hinder conservation efforts (Kahane et al., 2013). Addressing NUS 
conservation and their sustainable production is critically import-
234 species (11%) are considered at risk of extinction (i.e., extinct ant if they are to compete in the marketplaces dominated by a few 
in the wild; critically endangered; endangered; or vulnerable). The commodity crops. An integrated conservation approach combines ex 
Botanic Gardens Conservation International (BGCI) ThreatSearch situ, in situ and on farm methods and ensures the effective mainte-
database (https://tools.bgci.org/threat_search.php) lists conserva- nance and use of genetic diversity, the knowledge associated with 
tion assessments at global, regional, and national level for at least this diversity and its transmission to future generations (Padulosi, 
3,893 (55%) of the species in our list, with most species (76%) iden- Bergamini, & Lawrence, 2012). The primary challenge is the priori-
tified as “Not Threatened” (Figure 4). Many major food crop species tization of model species for impact, to make the best use of limited 
are widespread; therefore, it is likely that their extinction risk will resources. Species selection should be driven by shared priorities in 
be relatively low. Nonetheless, specific plant populations, including terms of nutrition, climate adaptation, income generation, cultural 
428  |     ULIAN et AL.
F I G U R E  4   Conservation status 
for 3,893 edible plant species from 
Diazgranados et al. (2020), according 
to the BGCI ThreatSearch database 
(https://tools.bgci.org/threat_search.
php). One assessment per species was 
selected, giving priority to the most 
recent assessment with highest risk. 
Records without an assessment year were 
excluded
diversity, ecosystem health, and the urgency of the intervention due by major projects, international agencies (Table S1) and researchers 
to ongoing genetic erosion. Women, young people, and indigenous (references in Table 3). We highlighted (in bold text in Table 3) those 
groups must be active participants in all of these exercises, through which are not currently listed as major food crops by the FAO (2015), 
a participatory bottom-up process (Padulosi, Phrang, Phrang, & for example, the mesquites in the Americas, morama bean in Africa, 
Rosado-May, 2019), as carried out by Dansi et al. (2012) in Benin and, Akkoub in Asia, rocket in Europe and Pindan walnut in Oceania (Table 3). 
more recently, by FAO within its Future Smart Food Initiative in Asia In addition, considering the differences in nutritional properties of the 
(FAO, 2018). This bottom-up approach will help develop innovative organ types (Guil-Guerrero & Torija-Isasa, 2002), we also reported the 
methods and tools of wide applicability that could be applied to other edible parts of each species. When comparing the taxa listed in Table 3 
NUS. Success and failures in promoting “new” crops can be found with those reported by Diazgranados et al. (2020), we found an aver-
across many regions, for example the effective establishment of lupin age of five uses recorded per taxa, and a peak of 24 taxa with seven 
cultivation in Australia (Nelson & Hawthorne, 2000), or the negative uses (Figure S2). Examples of NUS with many uses include the baobab 
social and environmental impact in the Andes caused by the quinoa in Africa, which is known as the “tree of life,” whose leaves, flowers, fruit 
boom (McDonell, 2018). To strengthen the self-sufficiency of food pulp, and seeds are used as food and to make beverages; the bark, roots, 
and production systems in terms of climate resilience, agroecological and seeds are medicinal; the bark is used for making rope, roofing ma-
benefits (e.g., soil improvers and species' enhancers), food and nutri- terial and clothing; and the hard husk of the fruit is used as calabash 
tion security (e.g., species and varieties that build resilient, more nutri- (Chadare, Linnemann, Hounhouigan, Nout, & Van Boekel, 2008; National 
tious and healthy diets), and income generation (e.g., diversity to build Research Council, 2008; Ngwako, Mogotsi, Sacande, Ulian, Davis, et al., 
economic resilience), there is a need for both sustainable promotion 2019). The taro, originally from Asia and also cultivated in Oceania, has 
and integrated conservation. Sustainable promotion makes diversity edible leaves, flowers, and roots; the roots are also medicinal and used as 
a central feature of the food system (at both intra- and inter-specific an additive to render plastics biodegradable (Arora, 2014; Linden, 1996).
levels), thereby potentially avoiding what has happened in the Andes Therefore, NUS of plants, as well as many edible species of fungi 
with quinoa, where global demand is being met by a few mainstream (see Box 1), represent potentially low-hanging fruit for a more resil-
varieties, while hundreds of others are being marginalized (Zimmerer ient, sustainable, biodiverse, and community participation-driven new 
& Carney, 2019). Low levels of funding for the promotion of NUS, “green revolution,” equitable and fair to the environment and all mem-
like yams, amaranth, Bambara groundnut, or African leafy vegetables, bers of society.
represents a major challenge for most countries interested in their 
promotion. Economic incentives and subsidies to private companies 
for producing local crops or certification schemes to recognize bio- 4  | QUALIT Y OF FOOD RESOURCES IN A 
diversity-rich products, should be actively pursued and include the CHANGING CLIMATE
establishment of an international “NUS Fund” specifically dedicated 
to supporting their development (Padulosi, Cawthorn, et al., 2019). In the coming century, major challenges to agriculture and biodi-
It is with this vision in mind that we provide a selection of highly versity will be dominated by increased climate variation. Hence, re-
promising NUS of plants (wild, domesticated, or semi-domesticated) search needs to increase our knowledge on the biology and ecology 
from different regions of the world (Table 3), which have been targeted of many NUS to be able to synthesize the future impact of climate 
ULIAN et AL.      |  429
TA B L E  3   Selection of neglected and underutilized plants (NUS) that have been recommended in scientific papers or targeted by 
collaborative projects, networks or international agencies. Species in bold are not listed in FAO (2015). Scientific names are ordered 
alphabetically and according to Kew's Plants of the World Online portal (http://www.plant softh eworl donli ne.org).
Target of projects, 
included in 
Main regions priority lists or 
Plant of natural focus of efforts by 
Common part(s) Region(s) of occurrence or key international 
name(s) Scientific name Source used origin cultivation Key reference(s) agencies (Table S1) Number of uses
1. Slippery Abelmoschus 2 1 Asia, Oceania Oceania, Asia Solberg, Seta- 7
cabbage, manihot (L.) Waken, Paul, 
bele, abika Medik. Palaniappan, and 
Iramu (2018)
2. Baobab Adansonia 1 1,2,3,4 Africa Africa Hall, Rudebjer, 3; 9; 12 10
digitata L. and Padulosi 
(2013), Kahane 
et al. (2013), 
National Research 
Council (2008), 
Ngwako, Mogotsi, 
Sacande, Ulian, 
Davis, et al. (2019)
3. Ground Aegopodium 1 1 Europe Europe Łuczaj et al. (2012) 0
elder podagraria L.
4. Candlenut Aleurites 1 5 Asia, Oceania Asia, Oceania 12 10
moluccanus (L.) 
Willd
5. Amaranth Amaranthus 3 1,4 Americas, Americas, Arora (2014), Hall 3; 4; 5; 8; 9;10; A. caudatus = 7;  
L. (incl.: A. Asia, Africa, Asia, Africa, et al. (2013), 11; 12 A. hybridus = 7; 
caudatus L., Europe Europe Hernandez- A. spinosum = 7; 
A. hybridus Bermejo and León A. retroflexus = 4
subsp. quitensis (1992), Kahane 
(Kunth) Costea et al. (2013), 
& Carretero, A. Kasolo, 
spinosus L., A. Chemining'wa, 
retroflexus L.) G., and Temu, A. 
(2018), Li et al. 
(2018), National 
Research Council 
(1996), Tyagi 
et al. (2017)
6. Elephant Amorphophallus 2 5 Asia Asia Arora (2014), Raneri 2 6
foot yam paeoniifolius et al. (2019), Tyagi 
(Dennst.) et al. (2017)
Nicolson
7. Sugar Annona spp. 2 3 Americas, Americas, Hall et al. (2013), 3; 12 4 (15 species)
apple (incl.: A. Asia Asia, Oceania Kasolo et al. (2018), 
squamosa L., A. Kour et al. (2018), 
cherimola Mill., Hernandez-
A. crassiflora Bermejo and León 
Mart., A. (1992), Padulosi 
muricata L.). et al. (2011), Tyagi 
et al. (2017)
8. Araucarias Araucaria 1 4 Americas, Americas, 12 4 (6 species)
Juss. [incl.: A. Oceania Oceania
angustifolia 
(Bertol.) Kuntze, 
A. araucana 
(Molina) 
K.Koch, A. 
bidwillii Hook.].
(Continues)
430  |     ULIAN et AL.
TA B L E  3   (Continued)
Target of projects, 
included in 
Main regions priority lists or 
Plant of natural focus of efforts by 
Common part(s) Region(s) of occurrence or key international 
name(s) Scientific name Source used origin cultivation Key reference(s) agencies (Table S1) Number of uses
9. Estragon Artemisia 2 1 Europe, Asia Europe, Asia 2 3
dracunculus L.
10. Artocarpus altilis 2 3 Asia, Oceania Asia, Oceania Thomson, Doran, 3; 9;11; 12 10
Breadfruit (Parkinson) and Clarke (2018), 
Fosberg Tyagi et al. (2017)
11. Jackfruit Artocarpus 2 3 Asia Asia Arora (2014), Kour 3; 9; 11 7
heterophyllus et al. (2018), Li 
Lam. et al. (2018), Tyagi 
et al. (2017)
12. Asparagus Tourn. 1 6 Europe Europe Arora (2014), Łuczaj 4 (19 species)
Asparagus ex L. et al. (2012), Tyagi 
et al. (2017)
13. Peach Bactris gasipaes 3 3 Americas Americas Hernandez- 2; 3; 12 7
palm Kunth Bermejo and León 
(1992), Kahane 
et al. (2013), 
Raneri et al. (2019), 
Wickens, Haq, and 
Day (1989)
14. Common Bambusa vulgaris 3 6 Asia Africa, Asia, 14; 15 9
bamboo Schrad. ex Oceania, 
J.C.Wendl. Americas
15. Ackee Blighia sapida 3 3 Africa Africa, Dansi et al. (2012), 3 7
K.D.Koenig Americas Hall et al. (2013)
16. Kale Brassica oleracea 3 1 Asia Europe, 8 7
L. Americas
17. Pigeon Cajanus cajan (L.) 2 4 Asia Africa, Asia, FAO (2010) 3; 10; 12 9
pea Huth Americas
18. Carissa Carissa spinarum 1 1,3 Africa Africa Kour et al. (2018), 3 9
L. National 
Research Council 
(2006), Omondi 
et al. (2019)
19. Lagos Celosia argentea 1 1 Africa Africa, Asia Hall et al. (2013), 3; 10; 11 8
spinach L. National Research 
Council (2006)
20. Bulbous Chaerophyllum 1 1 Europe Europe Łuczaj et al. (2012) 2
chervil bulbosum L.
21. Quinoa, Chenopodium 1 4 Americas Asia, Kasolo et al. (2018), 3; 4; 8; 9 4 (5 species)
Goosefoots, L. [incl.: Americas, Arora (2014), 
Cañahua Chenopodium Africa, Li et al. (2018), 
quinoa Willd., Europe Li et al. (2018), 
C. pallidicaule Łuczaj et al. (2012), 
Aellen, C. Padulosi 
giganteum et al. (2011), Raneri 
D.Don, Blitum et al. (2019)
bonus-henricus 
(L.) Rchb.]
22. Chicory Cichorium intybus 1 1 Europe Europe Łuczaj et al. (2012), 12 6
L. National Research 
Council (1996)
(Continues)
ULIAN et AL.      |  431
TA B L E  3   (Continued)
Target of projects, 
included in 
Main regions priority lists or 
Plant of natural focus of efforts by 
Common part(s) Region(s) of occurrence or key international 
name(s) Scientific name Source used origin cultivation Key reference(s) agencies (Table S1) Number of uses
23. Spider Cleome gynandra 3 1 Africa Africa, Asia, Dansi et al. (2012), 2; 3; 9; 11; 12 7
plant L. Americas Hall et al. (2013), 
Kahane et al. (2013), 
Raneri et al. (2019)
24. Chaya Cnidoscolus 2 1 Americas Americas Raneri et al. (2019) 4; 9; 10 6
aconitifolius 
(Mill.) 
I.M.Johnst.
25. Jobs’ Coix lacryma- 3 4 Asia Cosmopolitan Hall et al. (2013), 12 8
tears jobi L. Li et al. (2018), 
Sanogo et al. (2019)
26. Taro Colocasia 1 1,2,5,6 Asia Asia, Oceania Arora (2014), Hall 3; 11 8
esculenta (L.) et al. (2013), 
Schott Kahane et al. (2013)
27. Jute Corchorus 2 1,3 Asia, Africa Africa, Asia, Dansi et al. 4; 11 7
mallow olitorius L. Americas (2012), Padulosi 
et al. (2011), Raneri 
et al. (2019)
28. Crotalaria L. 3 1 Africa Cosmopolitan 3; 11; 12 4 (23 species)
Rattlepods
29. Japanese Cucurbita 2 3 Americas Americas Hernandez-Bermejo 3
pie pumpkin argyrosperma and León (1992)
C.Huber
30. Squash Cucurbita 2 3 Americas Americas Hernandez-Bermejo 5
moschata and León (1992)
Duchesne
31. Swamp Cyrtosperma 2 5 Asia Oceania, Asia Arora (2014), 3; 8; 9 3
taro merkusii (Hassk.) Kahane 
Schott et al. (2013), Li 
et al. (2018), Tyagi 
et al. (2017)
32. Fonio Digitaria exilis 2 4 Africa Africa FAO (2010), Kahane 3; 4; 8; 9; 11; 12 D. exilis = 8
(Kippist) Stapf et al. (2013), D. iburua = 3
and D. iburua National Research 
Stapf Council (1996); 
Raneri et al. (2019)
33. Yams Dioscorea 3 5 Africa Africa, Asia, Arora (2014), Hall 3; 8; 11; 12 D. 
cayenensis Americas, et al. (2013) cayenensis = 05, 
subsp. rotundata Oceania D. polystachya: 
(Poir.) J.Miège, 3, D. 
D. polystachya dumetorum = 6, 
Turcz., D. D. bulbifera = 7
dumetorum 
(Kunth) Pax, D. 
bulbifera L.
34. Barnyard Echinochloa 1 4 Africa, Asia, Africa, Asia, Arora (2014), Li 3 5 (10 species)
grass P.Beauv. Americas Americas et al. (2018), Raneri 
et al. (2019), Tyagi 
et al. (2017)
(Continues)
432  |     ULIAN et AL.
TA B L E  3   (Continued)
Target of projects, 
included in 
Main regions priority lists or 
Plant of natural focus of efforts by 
Common part(s) Region(s) of occurrence or key international 
name(s) Scientific name Source used origin cultivation Key reference(s) agencies (Table S1) Number of uses
35. Finger Eleusine coracana 2 4 Asia Asia, Africa Arora (2014), 3; 4; 7; 8; 9; 11; 12 7
millet (L.) Gaertn. FAO (2010), Hall 
et al. (2013), Li 
et al. (2018), 
National Research 
Council (1996), 
Raneri et al. (2019), 
Tyagi et al. (2017)
36. Teff Eragrostis tef 2 4 Africa Africa Arora (2014), FAO 2; 3; 8; 9; 11; 12 7
(Zuccagni) (2010), National 
Trotter Research Council 
(1996), Wickens 
et al. (1989)
37. Rocket Eruca vesicaria 2 1 Europe, Asia Europe, Asia, Arora (2014), Raneri 1; 3; 9; 7
(L.) Cav. Americas et al. (2019)
38. Torch lily Etlingera spp. 3 1,2,3,4 Asia Asia, Africa, 12 2 (13 species)
[incl.: E. elatior Americas, 
(Jack) R.M.Sm., Oceania
E. hemisphaerica 
(Blume) 
R.M.Sm.].
39. Fagopyrum 2 4 Asia Asia, Europe Arora (2014), Hall 3; 7; 8; 9; 12 9
Buckwheat esculentum et al. (2013), Kasolo 
Moench et al. (2018), Li 
et al. (2018), Raneri 
et al. (2019), Tyagi 
et al. (2017)
40. Sycamore Ficus sycomorus 1 1 Africa, Asia Africa, Asia, Tyagi et al. (2017) 3; 12 8
fig L. Americas
41. Kokum Garcinia L. 1 3 Asia Asia Arora (2014), Kour 3; 6; 12 3 (548 species)
[incl.: G. indica et al. (2018), Li 
(Thouars) et al. (2018), Tyagi 
Choisy, G. et al. (2017)
parvifolia 
(Miq.) Miq., G. 
gummi-gutta (L.) 
Roxb., G. morella 
(Gaertn.) Desr., 
G. binucao 
(Blanco) Choisy]
42. Ginkgo Ginkgo biloba L. 1 4 Asia Asia Arora (2014) 4
43. Goeppertia allouia 2 5 Americas Americas Hernandez-Bermejo 3; 12 2
Arrowroot (Aubl.) Borchs. and León (1992)
& S.Suárez
44. Akkoub Gundelia 1 6 Asia Asia 3; 12 3
tournefortii L.
45. Roselle Hibiscus 2 1,2 Africa Africa, Asia, Arora (2014), Kasolo 3; 12 8
sabdariffa L. Oceania et al. (2018), Li 
et al. (2018), Tyagi 
et al. (2017)
(Continues)
ULIAN et AL.      |  433
TA B L E  3   (Continued)
Target of projects, 
included in 
Main regions priority lists or 
Plant of natural focus of efforts by 
Common part(s) Region(s) of occurrence or key international 
name(s) Scientific name Source used origin cultivation Key reference(s) agencies (Table S1) Number of uses
46. Sea Hippophae 2 3 Asia Asia, Europe Arora (2014), Łuczaj 0
buckthorn rhamnoides L. et al. (2012), 
Padulosi 
et al. (2011)
47. Hop Humulus lupulus 1 1 Europe Europe Łuczaj et al. (2012) 12 5
L.
48. Lablab Lablab purpureus 2 4 Africa Africa, Asia Arora (2014), 3; 9; 11;12 7
(L.) Sweet Kahane 
et al. (2013), 
National Research 
Council (1996), 
Tyagi et al. (2017)
49. Lupin Lupinus mutabilis 2 4 Americas, Americas, Hernandez-Bermejo 3; 4; 12 L. mutabilis = 7
Sweet and L. Africa, Africa, and León (1992) L. albus = 8
albus L. Europe, Europe
50. Macadamia 2 3 Oceania Africa, Asia, Arora (2014), Kasolo 2
Macadamia tetraphylla Oceania et al. (2018), Tyagi 
L.A.S.Johnson et al. (2017)
51. Mallow Malva Tourn. 1 1 Europe, Asia Europe, Asia Arora (2014), Łuczaj 3 4 (4 species)
ex L. et al. (2012)
52. Microseris 1 1 Oceania Oceania 12 0
Microseris D.Don. [incl.: 
(several M. scapigera 
common Sch.Bip., M. 
names) lanceolata 
(Walp.) Sch.
Bip.].
53. Moringa Moringa oleifera 2 1,2,3,4 Africa, Asia Asia, Africa, Arora (2014), 3; 8; 9; 11; 12 M. oleifera = 10
Lam. and M. Americas Kahane M. 
stenopetala et al. (2013), Kasolo stenopetala = 7
(Baker f.) Cufod. et al. (2018), Kour 
et al. (2018), Li et al. 
(2018), National 
Research Council 
(2006), Padulosi 
et al. (2011), Raneri 
et al. (2019), Tyagi 
et al. (2017)
54. African Oldeania alpina 1 6 Africa Africa 12 6
Alpine (K.Schum.) 
bamboo Stapleton
55. Ostrich Onoclea 1 1 Europe, Asia, Europe, Asia, Łuczaj et al. (2012) 12 2
fern struthiopteris Americas Americas
Roth
56. Oca Oxalis tuberosa 3 5 Americas Americas Hernandez-Bermejo 3; 9; 12 2
Molina and León (1992)
57. Pandanus 3 3 Asia, Oceania Asia, Oceania Thomson, Cruz-de 6
Pandanus, tectorius Hoyos, Cummings, 
screwpine Parkinson ex and Schultz (2016)
Du Roi
(Continues)
434  |     ULIAN et AL.
TA B L E  3   (Continued)
Target of projects, 
included in 
Main regions priority lists or 
Plant of natural focus of efforts by 
Common part(s) Region(s) of occurrence or key international 
name(s) Scientific name Source used origin cultivation Key reference(s) agencies (Table S1) Number of uses
58. Proso Panicum 2 4 Asia, Europe, Asia, Europe, Arora (2014), 3; 12 P. miliaceum = 5, 
millet miliaceum L. and Oceania Oceania Kahane P. 
P. decompositum et al. (2013), Li decompositum  
R.Br. et al. (2018), Raneri = 0
et al. (2019), Tyagi 
et al. (2017)
59. Guarana Paullinia cupana 1 3,4 Americas Americas Hernandez-Bermejo 3; 12 4
Kunth and León (1992)
60. Perilla Perilla frutescens 2 1,4 Asia Asia Arora (2014), Li 3; 12 5
(L.) Britton et al. (2018), Tyagi 
et al. (2017)
61. Tepary Phaseolus 2 4 Americas Americas Hernandez-Bermejo 4; 12 5
bean acutifolius and León (1992)
A.Gray
62. Runner Phaseolus 2 4 Americas Americas, Asia Arora (2014), 12 6
bean coccineus L. Hernandez-
Bermejo and 
León (1992), Tyagi 
et al. (2017)
63. Moso Phyllostachys 1 6 Asia Asia Arora (2014), Li et al. 4
bamboo edulis (Carrière) (2018)
J.Houz.
64. Plinia rivularis 2 3 Americas Americas Hernandez-Bermejo 3; 12 P. rivularis = 0, 
Jaboticaba (Cambess.) and León (1992) P. cauliflora = 2
Rotman and P. 
cauliflora (Mart.) 
Kausel
65. Zapote Pouteria 1 3 Americas Americas Hernandez-Bermejo 3; 12 2
sapota (Jacq.) and León (1992)
H.E.Moore & 
Stearn
66. Mesquite Prosopis L. [incl.: 3 4 Americas, Americas, Arora (2014), 3; 12 7 (14 species)
P. alba Griseb., Africa, Asia Africa, Asia Wickens 
P. chilensis et al. (1989)
(Molina) Stuntz, 
P. juliflora (Sw.) 
DC.].
67. Bracken Pteridium 1 1 Cosmopolitan Cosmopolitan Liu, Wujisguleng, 12 7
aquilinum L. and Long (2012)
Kuhn s.l.
68. Oak Quercus L. 1 3 Europe, Asia, Europe, Asia, Łuczaj et al. (2012) 12 4 (9 species)
Africa Africa, 
Americas
69. Rasberry Rubus L. [incl.: 3 3 Asia, Oceania Asia, Oceania Arora (2014), Łuczaj 3; 12 3 (36 species)
R. hawaiensis et al. (2012)
A.Gray, R. 
macraei A.Gray, 
R. rosifolius Sm., 
R. parvifolius L.].
(Continues)
ULIAN et AL.      |  435
TA B L E  3   (Continued)
Target of projects, 
included in 
Main regions priority lists or 
Plant of natural focus of efforts by 
Common part(s) Region(s) of occurrence or key international 
name(s) Scientific name Source used origin cultivation Key reference(s) agencies (Table S1) Number of uses
70. Sorrels Rumex acetosa 1 1 Europe, Asia, Europe, Asia, Łuczaj et al. (2012) 12 R. acetosa = 4, 
L. and R. Americas Americas R. 
lapponicus lapponicus = 0
(Hiitonen) 
Czernov
71. Sagittaria 1 1 Europe Europe, Asia Arora (2014) 2 (5 species)
Arrowhead Ruppius ex L.
72. Elder Sambucus nigra 1 2,3 America, America, Łuczaj et al. (2012) 12 S. nigra = 5, S. 
L., S. canadensis Africa, Africa, canadensis = 5
L. Europe, Europe, 
Americas Americas
73. Santalum 1 3 Oceania Oceania Arora (2014) 3; 12 6
Quandong acuminatum 
(R.Br.) A.DC.
74. Marula Sclerocarya 1 3 Africa Africa Kahane et al. (2013), 3; 11; 12 10
birrea (A.Rich.) National Research 
Hochst. Council (2008), 
Ngwako, Mogotsi, 
Sacande, Ulian, and 
Mattana (2019)
75. Common Scolymus 1 6 Europe Europe Łuczaj et al. (2012) 3; 12 2
golden hispanicus L.
thistle
76. Black Scorzonera 2 5 Europe Europe 8; 12 4
salsify hispanica L.
77. False Sesamum 1 1 Africa Africa Dansi et al. (2012) 3 6
sesame sesamoides 
(Endl.) Byng & 
Christenh.
78. Foxtail Setaria italica (L.) 2 4 Asia, Europe Asia, Europe Arora (2014), 2; 3; 8; 12; 7
millet P.Beauv. Kahane 
et al. (2013), Li 
et al. (2018), Raneri 
et al. (2019)
79. Bladder Silene vulgaris 1 1 Europe, Europe Łuczaj et al. (2012) 2
campion (Moench) Americas
Garcke
80. Cardus Silybum 1 6 Europe, Asia Europe, Asia, Łuczaj et al. (2012) 4
marianus marianum (L.) Africa, 
Gaertn. Americas
81. Mustard Sinapis L. 1 1,4 Europe Europe, Asia, Arora (2014) 7 (S. alba)
Africa, 
Americas
82. Yacon Smallanthus 2 5 Americas Americas Hernandez-Bermejo 3; 9; 12 2
sonchifolius and León (1992)
(Poepp.) H.Rob.
83. Smilax excelsa 1 6 Europe, Asa, Europe, Asa, Tyagi et al. (2017) 12 S. excelsa = 0, 
Greenbriers L., S. glyciphylla Oceania Oceania S. 
J.White, S. ferox glyciphylla = 3, 
Wall. ex Kunth S. ferox = 0
(Continues)
436  |     ULIAN et AL.
TA B L E  3   (Continued)
Target of projects, 
included in 
Main regions priority lists or 
Plant of natural focus of efforts by 
Common part(s) Region(s) of occurrence or key international 
name(s) Scientific name Source used origin cultivation Key reference(s) agencies (Table S1) Number of uses
84. Tomato Solanum 1 3 Americas Americas, Hernandez-Bermejo 3 3
tree betaceum Cav. Asia, Oceania and León (1992)
85. African Solanum L. [incl.: 3 3 Africa, Asia, Africa, Asia, Kahane et al. (2013), 3; 11; 12 4 (51 species)
eggplant S. aethiopicum Americas, Americas, National Research 
and other L., S. quitoense Oceania Oceania Council (2006)
names Lam., S. 
sessiliflorum 
Dunal, S. 
ellipticum R.Br., 
S. juzepczukii 
Bukasov, S. 
curtilobum Juz. 
& Bukasov, S. 
glaucescens 
Zucc.]
86. African Solanum scabrum 3 1 Africa Cosmopolitan 3; 11 6
black Mill.
nightshade
87. African Sphenostylis 2 4,5 Africa Africa, Asia Arora (2014), Dansi 3; 8; 9; 11 5
yam bean stenocarpa et al. (2012), 
(Hochst. ex National Research 
A.Rich.) Harms Council, (1996), 
Tyagi et al. (2017)
88. Jocote Spondias spp. 3 3 Americas Americas Hernandez-Bermejo 3; 12 6 (6 species)
(incl.: S. and León (1992)
purpurea L., 
S. mombin 
L., S. dulcis 
Parkinson)
89. Marsh Stachys palustris 1 1 Europe, Asia Europe, Asia, Arora (2014), Łuczaj 12 0
woundwort L., S. tymphaea Americas et al. (2012), Tyagi 
et al. (2017)
90. Malay Syzygium 2 3 Asia, Asia, Oceania Thomson 6
apple malaccense Oceania, et al. (2018)
(L.) Merr. & 
L.M.Perry
91. Pindan Terminalia 1 4 Asia, Asia, Oceania Arora (2014), Tyagi 3; 12 6 (21 species)
walnut L. (incl.: T. Oceania, et al. (2017)
cunninghamii 
C.A.Gardner)
92. Salsify Tragopogon L. 1 6 Europe, Asia Europe, Asia, Arora (2014), Tyagi 3 (T. porrifolius)
Americas, et al. (2017)
Oceania
93. Buffalo Trapa L.(incl.: 1 4 Europe, Asia, Europe, Asia, Arora (2014), Turner 12 T. napans = 6
nut T. natans L., T. Africa Africa et al. (2011), Tyagi T. japonica = 0
japonica Flerow) et al. (2017)
94. African Treculia africana 1 3 Africa Africa Kasolo et al. (2018) 2; 3; 12 7
breadfruit Decne. ex 
Trécul
95. Snake Trichosanthes 2 3 Asia Asia Arora (2014), Li 5
gourd cucumerina L. et al. (2018), Tyagi 
et al. (2017)
(Continues)
ULIAN et AL.      |  437
TA B L E  3   (Continued)
Target of projects, 
included in 
Main regions priority lists or 
Plant of natural focus of efforts by 
Common part(s) Region(s) of occurrence or key international 
name(s) Scientific name Source used origin cultivation Key reference(s) agencies (Table S1) Number of uses
96. Morama Tylosema 2 4 Africa Africa (Mogotsi, Sacande, 8; 12 6
bean esculentum et al., 2019)
(Burch.) 
A.Schreib.
97. Bulrush Typha orientalis 1 6 Europe, Asia, Cosmopolitan Turner et al. (2011) 3; 12 T. orientalis = 5, 
C.Presl, T. Americas  T. 
domingensis dominigensis  
Pers., T latifolia = 8,  
L. T. latifolia = 7
98. Ulluco Ullucus tuberosus 2 5 Americas Americas Hernandez-Bermejo 3; 9; 12 2
Caldas and León (1992)
99. Nettle Urtica dioica L. 1 1 Europe, Asia, Europe, Asia, (Łuczaj et al., 2012) 12 U. dioica = 7, U. 
and U. massaica Africa Africa massaica = 3
Mildbr.
100. Small Vaccinium 1 3 Europe, Asia, Europe, Asia, 12 3 (20 species)
cranberry spp. (incl.: V. Americas Americas
oxycoccos L., 
V. floribundum 
Kunth, V. 
praestans Lamb.)
101. Vigna subterranea 2 4 Africa Africa, Asia, Arora (2014), Dansi 2; 3; 4; 5; 8; 9; 5
Bambara (L.) Verdc. Oceania et al. (2012), 11; 12
groundnut FAO (2010), Hall 
et al. (2013), 
Kahane et al. (2013), 
National Research 
Council, (1996), 
Padulosi 
et al. (2011), Raneri 
et al. (2019)
102. Shea Vitellaria 1 3 Africa Africa National Research 3; 11; 12 10
tree paradoxa Council (2006)
C.F.Gaertn.
Source: 1, mainly wild; 2, mainly cultivated; 3, wild and cultivated. Edible parts: 1, leaves; 2, inflorescences/flowers; 3, fruits; 4, seeds; 5, roots/tubers; 
6, stems/shoots. Number of uses according to Diazgranados et al. (2020); details on the type of uses for each species are reported in Diazgranados 
et al. (2020); 0, species not listed. When the common name of the NUS corresponds to more than one specific epithet, the number of uses is here 
reported as an average of the species listed in Diazgranados et al. (2020)
change over the century, including how climate change might im- least one month of the year and humans use 70% of available fresh 
pact the quality and nutritional value of edible species (Borrell water for agricultural purposes, the monitoring of water irrigation 
et al., 2020). Studies in this research area have mainly focused on systems is a recommended strategy to help conserve water (Green 
established domesticated edible crops. For example, under future et al., 2018).
climate-scenario drought stress conditions, Hummel et al., 2018 re- Although future drought conditions have been suggested to 
ported that iron levels in beans (Phaseolus vulgaris) decreased, while increase protein levels in the legume species P. vulgaris (Hummel 
levels of protein, zinc, lead, and phytic acid increased. This study also et al., 2018), in contrast, increased CO2 levels were found to reduce 
revealed that bean nutritional quality and yields were reduced under protein levels and increase omega-3 fatty acid levels in mung bean 
future predicted drought conditions, leading the authors to con- (Vigna mungo; Ziska, Epstein, & Schlesinger, 2009). Environmental 
clude, with supportive data from crop modeling, that current bean factors may also impact on the nutritional quality of edible nuts, 
growing areas in south-eastern Africa could become unsuitable by including almonds, pistachios, and walnuts. For example, in 29 
2050. Given the predicted impact of future drought conditions on different cultivars, protein, phytosterol, and mineral content were 
crops and, as 66% of people live with severe water scarcity for at affected, suggesting that climate change may also compromise 
438  |     ULIAN et AL.
BOX 1 Fungi as food resources
Beyond the few species that are used in biotechnology for the production of pharmaceuticals, industrial enzymes and plastics (Howes 
et al., 2020; Prescott et al., 2018), the vast majority of fungi are underutilized. However, those in mainstream agriculture have an 
estimated annual market value of more than US$62 billion by 2023 (Knowledge Sourcing Intelligence LLP, 2017). As edible fungi are 
sources of fiber, selenium, potassium, copper, zinc, B group vitamins, and are one of the only non-animal sources of dietary forms of 
vitamin D, a deficiency of which is a risk factor for rickets in children (World Health Organization, 2019), the potential future use of ne-
glected fungi is considerable. Indeed, during their growth stage and post-harvest, mushrooms exposed to sunlight or controlled levels 
of UV radiation had increased concentrations of vitamin D2 (Cardwell, Bornman, James, & Black, 2018). The impact of UV radiation on 
the vitamin D content of mushrooms could be evaluated further as a strategy to enhance availability of dietary vitamin D, especially 
in regions where rickets or osteomalacia are health risks.
Around 2% of fungi form mutualistic mycorrhizal relationships with plants (Suz et al., 2018). Within these mutualistic relationships, 
the plant provides sugars in exchange for minerals and nutrients from the fungus. While some mycorrhizal fungi are often the most 
desirable fungi for consumption, they elude efforts, with a few exceptions, to be cultivated commercially (Boa, 2004). These desir-
able mycorrhizal species are instead foraged from the wild, based on distinct cultural practices. However, it is unknown if the impact 
of foraging on wild populations can be sustained into the future, where harvesting is likely to increase. Currently of concern is the 
Kalahari truffle (Kalaharituber pfeilii), which is sold in local markets in southern Africa, with a rapidly increasing commercial harvesting 
(Mogotsi, Tiroesele, et al. (2019) and references therein). In contrast, saprotrophic fungi are well suited to commercial myco-culture, 
and up to 200 species are known to be cultivated around the world. Over 85% of cultivated mushroom species belong to just five 
genera: Agaricus (button, portobello, and chestnut mushrooms), Lentinula (shiitake), Pleurotus (oyster mushrooms), Auricularia (jelly and 
wood ear fungi), and Flammulina (Enokitake; Royse, Baars, & Tan, 2017).
The cultivation of fungi represents an opportunity to develop valuable new crops that require low resource inputs, create little waste 
(SureHarvest, 2017), are sustainable, and can be tailored to local cultural preferences. Cultivation can be at the domestic and commu-
nity level (Martínez-Carrera et al., 1998) and has the potential to be scaled up commercially (Zhang, Geng, Shen, Wang, & Dai, 2014). 
Importantly, new species are being brought into cultivation (Rizal et al., 2016; Thongklang, Sysouphanthong, Callac, & Hyde, 2014) 
and these have economic potential beyond the value of a few internationally grown strains (Hyde et al., 2019). For example, within 
the genus Termitomyces, species such as T. microcarpus and T. clypeatus are consumed across Africa and Asia (Boa, 2004) and bring-
ing species from this genus into cultivation could be a desirable cash crop for local communities. Myco-agriculture is most diverse in 
China, with over 100 species of the 1,789 reported edible species already in cultivation and around 60% in commercial production 
(Fang et al., 2018; Zhang et al., 2014).
Finally, mycorrhizal fungal associations can also improve the nutritional quality of the edible parts of plant crops. For instance, mycor-
rhizal fungi inoculation of strawberries can increase the levels of anthocyanins and phenolic compounds, and in tomatoes can increase 
the levels of P, N, and Cu and flavour compounds (Torres, Antolín, & Goicoechea, 2018). More research is needed to understand the 
promising role that mycorrhizal fungi play in the nutritional value of edible plants, including NUS, particularly in the context of strate-
gies to produce nutritious crops in a changing climate.
nutritional value in this food group (Rabadán, Álvarez-Ortí, & of human health. For instance, extreme environmental conditions 
Pardo, 2019). Together, these findings suggest that different climatic (late season cultivation) have been shown to increase phenolic 
factors could mediate contrasting effects on the nutritional value and vitamin C content in some broccoli cultivars (Vallejo, Tomas-
of crops, and this should be considered, separately for each spe- Barberan, & García-Viguera, 2003). Higher CO2 levels also increased 
cies, with respect to NUS. Although some studies conclude that vitamin C and antioxidant capacity in lettuce, celery, and Chinese 
elevated atmospheric CO2 reduces protein and mineral content in cabbage, although other nutrients (micro- and macro-) decreased 
vegetables, CO2 can enhance vegetable yield and concentrations of (Leisner, 2020). Thus, certain phytochemicals relevant to health in 
soluble saccharides, phenolic compounds, including flavonoids, and crop plants may be positively influenced by environmental changes, 
vitamin C, in addition to the antioxidant capacity (Dong, Gruda, Lam, while levels of some essential macro- and micro-nutrients may be 
Li, & Duan, 2018). Furthermore, flavonol and anthocyanin levels in negatively affected. In view of the emerging research that suggests 
fruits may be increased by changes in expression of hydroxylases in that certain environmental factors could negatively impact on the 
response to environmental conditions, including water deficits and nutritional quality of food, the potential consequences for human 
UVB radiation (Martínez-Lüscher et al., 2014). health in the long-term are concerning, particularly against the 
The impact of emerging environmental stresses on biologically backdrop of the global scale of malnutrition, which includes pro-
active chemicals of edible plants is important from the perspective tein-energy, vitamin and mineral deficiencies (De Onis, Monteiro, 
ULIAN et AL.      |  439
Akré, & Glugston, 1993; Green et al., 2018; https://www.who.int/ 5  | CONCLUSIONS
news-room/fact-sheet s/detai l/malnu trition). While biofortification 
could be one approach to mitigate the impact of climatic changes on In this article, we provide an overview of the global state of edible 
food nutritional status (Green et al., 2018), more extensive scrutiny plants, highlighting their diversity, and distribution among vascular 
of the nutritional quality of crops, including NUS, in the context of plant families from around the world. We emphasize that this di-
predicted environmental challenges, should be aligned with other versity stands in striking contrast with the few hundred food crops, 
strategies for food security. In circumstances where saccharide lev- originating from main domestication centers, that mainstream ag-
els increase in edible species in response to climate factors (Dong riculture currently relies on. By integrating the other uses, we also 
et al., 2018), the consequences should be considered in the context highlight the additional ecosystem services these plants provide 
of providing energy as a source of calories in both undernutrition that are important for people's livelihoods and wellbeing (Díaz 
(such as in wasting and being underweight) and obesity, with the lat- et al., 2020). While more work is needed to assess the actual con-
ter associated with increased risk of certain non-communicable dis- servation status of edible plants, ex situ conservation (and particu-
eases (https://www.who.int/news-room/fact-sheet s/detai l/malnu larly seed banking) is already playing an important role in preserving 
trition). them. However, information on the functional and genetic diversity 
Potential strategies to ameliorate the effects of climate change of stored seed collections is limited and alternative ex situ conser-
on food security in the future include greater understanding of the vation approaches, such as cryopreservation, need to be developed 
global distribution of edible plants and by creating more diverse for those species with non-bankable seeds (Li & Pritchard, 2009).
and climate-resilient agricultural production systems (see Table We highlight key NUS of edible plants with the potential to im-
S1). In addition, improved knowledge of naturally stress-resistant prove the quality, resilience, and self-sufficiency of food production, 
plants and their broader cultivation would enable agriculture, and while deploying a more sustainable local food supply. We also con-
the human diet, to be diversified as one strategy for global food sider the importance of fungi, which could enhance the nutritional 
security in the changing environment (Zhang, Li, & Zhu, 2018), value of foods, through the provision of beneficial vitamins and min-
especially when aligned with methods to maintain the genetic erals, and which have potential to be developed into valuable and 
diversity of crops (e.g., seed banking; Borrell et al., 2020). More sustainable crops.
research on elucidating the genes and processes that underlay However, before NUS can become successful crops of the fu-
the mechanisms for climate-resilience of edible species could also ture, many knowledge gaps need to be filled relating to their biology 
underpin future strategies to mitigate environmental challenges and ecology. In addition, research efforts are needed on understand-
that threaten food security (Dhankher & Foyer, 2018). Indeed, ing the impacts of climate change on NUS, to enable the develop-
a multi-faceted approach integrating physiology, genomics, and ment of effective and sustainable agricultural practices for future 
climate modeling has been proposed as important to develop a climate conditions (Turner et al., 2011; Ulian, Pritchard, Cockel, & 
sustainable future food supply considering global climate change Mattana, 2019). Although methods and tools developed by farm-
(Leisner, 2020). ers and researchers for the cultivation of major crops can be easily 
To address the impact of climate change on nutritional security adapted to improve the cultivation of NUS, these should be inte-
in the future, a model has been described (Fanzo, Davis, McLaren, & grated with local traditional knowledge on uses and practices to help 
Choufani, 2018) to increase net nutrition in the food chain under cli- protect the environment and promote the conservation of biodiver-
mate change. This model encompasses agriculture practices to cul- sity (Casas et al., 2016; Horlings & Marsden, 2011; Patel, Sharma, & 
tivate improved varieties, and new production locations to minimize Singh, 2020). To further aid the development of NUS as future crops, 
loss of biodiversity, through to processing, distribution, marketing research programs need to be strengthened and the necessary re-
(including promotion of food benefits), and consumption strategies search infrastructure put in place, including addressing shortages 
to maximize nutrition availability for vulnerable groups. A positive in relevant fields (FAO, 2019). This will require improved mech-
correlation between high agricultural diversity and high nutrient anisms for exchanging information rapidly and effectively, as well 
production, irrespective of farm cultivation size, has been suggested as increased awareness of the importance of crop diversity among 
from global examination of food commodities (Herrero et al., 2017), and between stakeholder groups. One way this could be achieved 
indicating that one strategy to protect availability of nutrients may is through participatory decision-making processes (Padulosi 
be through promoting agricultural diversity, and therefore, dietary et al., 2011) and by putting in place effective legal and policy frame-
range to support health. works (FAO, 2019; Noorani et al., 2015) that are accompanied by 
Emerging evidence shows climate change impacts not only on economic incentives and subsidies to support the development of 
food quality, nutrition, safety (Borrell et al., 2020), and cost, but also NUS (Padulosi, Cawthorn, et al., 2019).
on the ability to transport food from “farm to fork,” thus, for many Biodiversity offers a largely untapped resource to support our 
communities, restricting their access to an adequate dietary range planet and improve our lives and has the potential to “end hunger, 
(Fanzo et al., 2018). These factors combined will limit the availability achieve food security and improve nutrition and promote sustain-
of nutrients with potentially serious consequences for the health of able agriculture,” as articulated in the UN SDG 2, through the de-
humanity. velopment of climate-resilient crops and the more widespread use 
440  |     ULIAN et AL.
of localized crop species (Antonelli, Smith, & Simmonds, 2019), such changes on the yield and nutritional quality of fruits, nuts and seeds: 
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