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Underutilized wild edible fungi and their undervalued ecosystem services in Africa

Abstract

Background

Edible fungi including wild mushrooms have been largely neglected and underutilized in Africa. Not only is the number of edible species unknown, but the critical role they play in human food and nutrition and the ecosystem services they provide have remained poorly understood and undervalued.

Methods

We reviewed the literature with the objective of providing a synthesis of knowledge on (1) the diversity of wild edible fungi of Africa with emphasis on mushrooms; (2) the ecosystem services provided by wild edible fungi; (3) anthropogenic threats to their diversity and utilization; and (4) opportunities for their conservation and cultivation.

Results

We identified a total of 480 species of wild edible mushrooms in 126 genera belonging to 60 fungal families across Africa. A total of, 249 species were mycorrhizal, 28 species were termitophilic and 203 species were saprophytic or parasitic. Wild edible mushrooms provide food that is high in digestible proteins, antioxidants and fibre but low in fats. They are also sources of income for rural populations. Almost all species play a role in nutrient recycling and hence the productivity of forests and agroecosystems. However, deforestation and land degradation are threatening the mushroom diversity in some regions of Africa.

Conclusion

The African continent is endowed with a tremendous diversity of neglected and underutilized edible wild mushrooms providing nutritious food for humans and playing a key role in the structure and functioning of native forests and woodlands. Deforestation and the loss of habitat are the greatest threats to edible wild species in Africa. The loss of indigenous knowledge can also potentially limit dietary choices and market opportunities. Therefore, we recommend national forestry research and development programs and international frameworks such as Reducing Emissions from Deforestation and Degradation (REDD +) to invest in the conservation, cultivation and valorisation of wild edible mushrooms to achieve sustainable forest management and the welfare of local communities.

Introduction

Edible wild fungi include boletes, bracket fungi, button mushrooms, chanterelles, coral mushrooms, jelly fungi, morels, oyster mushrooms, russula and smuts (Arora and Shepard 2008; Boa 2012; Degreef et al. 2016). In its broad sense the term mushroom includes fruiting-bodies of most of these macrofungi. Mushrooms are highly valued as food and also as sources of medicinal products. According to Boa (2004), poisonous species are relatively few, while fatal ones belong to a tiny minority. Recognition of edible and inedible mushrooms depends largely on folk taxonomy whereby traditional knowledge provides the tool for communication, and the passing of information from one generation to another (Tibuhwa 2012). Owing to non-exhaustiveness of the macrofungal research inventory and insufficient coverage of many parts of Africa, the actual proportion consumed is still unknown. Whatever the actual proportion is, indications are that the numbers are dwindling due to the loss of indigenous knowledge and destruction of habitats.

In Africa, edible fungi are mostly harvested from the wild (Boa 2004). Relatively little success has been achieved in cultivation and commercialization of edible wild mushrooms except in a few countries such as South Africa and Zimbabwe (Boa 2004; Ndifon 2022). Mushroom cultivation requires less land than livestock or crops and can be a cost-effective way to provide proteins for low-income households (Thomas and Vazquez 2022). Using Lactarius indigo as an example, Thomas and Vazquez (2022) show how the cultivation of mycorrhizal fungi combined with trees can provide dietary proteins exceeding that of extensive pastoral beef production. Although there are general prospects for commercialization, trade in mushrooms is also poorly developed in Africa. Despite being available in quantities more than local people can actually use, edible wild mushrooms are also commercially underexploited in some parts of Africa (Tibuhwa 2012; Milenge and de Kesel 2020. In many countries, mushrooms are consumed out of necessity for food. For example, in Malawi, Zambia and the Democratic Republic of Congo, mushrooms are consumed during the “hunger” months in the agricultural cycle when the main foods are scarce (Boa 2004; Degreef et al. 1997). The role of wild edible mushrooms in environmental health has also not attracted significant attention in research and development (R&D). In development projects, mushrooms are often cited as non-timber forest products (NTFPs), but their commercial exploitation is promoted without explicit consideration for their sustainable use and conservation. Mushrooms, and truffles which grow and fruit underground, are a particularly overlooked component of tree health. Many species play a vital role in sustaining the growth and productivity of native forest, woodlands and commercial plantations through their symbiotic associations with trees and termites (Boa 2004; Sileshi et al. 2010).

Most of the current research in Africa involves ethnomycological surveys to identify preferred species from the wild followed by some analysis of their nutritional value. The focus of such studies has been on species from tropical/sub-tropical woodlands and grasslands, while relatively little is known about species that occur in the drylands. For example, all known truffles of Africa are desert species producing subterranean fruiting bodies in mycorrhizal associations with desert plants (Thomas et al. 2019). While some information exists on the use of truffles by local populations in North Africa and Southern Africa, literature is scanty on the truffles of tropical Africa (Thomas et al. 2019). One of the co-authors (A. Mlambo) has a first-hand experience about an edible truffle Mackintoshia persica in Zimbabwe although it has not been reported in the literature. Current knowledge of the threats to edible mushrooms and their ecosystem service from anthropogenic and other factors is also severely limited. Whereas opportunities exist for conservation and cultivation of mycorrhizal mushrooms, lack of investment in research and development has limited generation of critical information in this area. Knowledge gaps also exist on the diversity and ecological functions of edible wild mushrooms in many countries in Africa (Tibuhwa et al. 2011, de Crop et al. 2012).

Saprobic mushrooms play critical ecological roles in nutrient recycling. Mycorrhizal fungi shape plant communities through mutualistic relationships (Martin et al. 2016). Termitophilic fungi may help in shaping the vegetation through their mutualistic association with termites, which in turn create substrate heterogeneity by altering soil properties (Bonachela et al. 2021; Sileshi et al. 2010). Over the years considerable knowledge has accumulated on the role of these associations in forest health (Anthony et al. 2022; Martin et al. 2016; Meidl et al. 2021; Pérez-Moreno et al. 2021). However, the critical roles that mushrooms play through their symbiotic associations with trees and termites have remained poorly understood by the general public and not fully integrated in international development frameworks such as REDD + . Imbalances in research and the lack of representation in the academic literature also create a bottleneck for evidence-based practice and policy for conservation and utilization of edible fungi in Africa. Therefore, the objective of this review is to provide a synthesis of the state of knowledge on (1) the diversity of wild edible fungi of Africa with emphasis on mushrooms; (2) the ecosystem services provided by wild edible fungi; (3) anthropogenic threats to their diversity and utilization; and (4) opportunities for their conservation and cultivation.

Methods

Through a comprehensive review of the literature, this synthesis attempts to integrate current knowledge from ethnomycological and field surveys, taxonomic studies, and studies on nutritional compositions conducted on edible wild fungi in African countries. The literature review considered all types of relevant studies including those published in peer-reviewed journals, book chapters, theses, dissertations and grey literature (reports and articles not formally published by commercial academic publishers) on the subject matter. We conducted the literature search using CAB index, Google Scholar, PubMed, and Scopus limiting the search to studies published in the English language covering continental Africa and all the island states. We focussed the search on studies on the diversity, use and valorization of edible wild mushrooms. Accordingly, in each search engine we used various combinations of the following key words: edible wild fungi, ethnomycology, mushrooms, mycorrhizal edible fungi, saprobic edible fungi, proximate composition, Termitomyces and Africa. In total we found 43 published studies focussing on ethnomycological surveys documenting the diversity of wild edible species (Additional file 1: Tables S1-S3 and References). We also found 17 studies providing proximate analysis and nutritional values, and four studies presenting comparable antioxidant contents. We confirmed the preferred species names by reference to the most up-to-date checklists of macrofungi published in Kinge et al. (2020), Ndifon (2022), Index Fungorum website (http://www.indexfungorum.org/names/names.asp) and Mycobank (https://www.mycobank.org/).

Synthesis

Diversity of edible wild mushrooms

The literature review revealed that the African continent holds a large diversity of wild edible fungi. In total 480 species in 126 genera belonging to 60 fungal families were confirmed to be edible across Africa (Table 1; Additional file 1: Table S1-S3). This only covers species identified as edible by local communities during ethnomycological surveys and confirmed by taxonomists to some degree. Obviously, the list is incomplete because those surveys have only covered a limited number of communities or parts of each country. The results are also limited by lack of nation-wide inventories in many countries. Therefore, the numbers are likely to increase as species are added when new surveys are reported as demonstrated by a recent study by Degreef et al. (2016) on mushrooms of Burundi and Rwanda. In addition, species new to science are added with increasing collection of specimens in areas that have not been covered in earlier taxonomic work. There is also a continuous influx of new species as taxa are revised, species are described, and records are added to the distribution of known species in Africa. With the application of molecular taxonomic tools, a number of new edible species have been added to genera such as Russula, Cantharellus and Amanita. For example, in 2007 Buyck and Sharp (2007) published first records of 15 Russula species including two new species (Russula termitaria and Russula terrena) in Zimbabwe. In 2008, Buyck et al. (2013) described two new species (Russula edulis and Russula prolifica) from Madagascar. In 2013, Buyck et al. (2013) described five new species of Cantharellus (Cantharellus afrocibarius, C. gracilis, C. humidicolus, C. miomboensis and C. tanzanicus) from the Zambezian miombo woodlands in tropical Africa. Buyck et al. (2013) rejected the recognition of Afrocantharellus as a separate genus initially elevated from a subgenus to genus by Tibuhwa et al. (2012). In 2016, de Kesel et al. (2016) described four new species of Cantharellus (Cantharellus guineensis, C. mikemboensis, C. pseudomiomboensis and C. stramineus) from the miombo woodlands. In 2019, Buyck et al. (2019) descried two new species (Cantharellus densilamellatus and C. tomentosoides) from the Democratic Republic of Congo and Tanzania. In the same year, Fraiture et al. (2019) described two new species of Amanita (Amanita bweyeyensis and Amanita harkoneniana) from Rwanda, Burundi and Tanzania and Madagascar. Mighell et al. (2021) described three new species of Amanita (Amanita minima, Amanita luteolamellata, and A. goossensfontanae) from the Guineo-Congolian rainforests of Cameroon. Some of these are edible (Additional file 1: Table S1-S3).

Table 1 Number of mycorrhizal edible wild mushroom genera and species reported r in different African countries and the most frequently reported species (details in Additional file 1: Tables S1–S3)

Based on their mode of nutrition, the edible fungal species can be categorized into three trophic groups, namely mycorrhizal, termitophilic and saprobic mushrooms. This distinction is very important because it reveals whether or not a mushroom can be cultivated on artificial media or not. It also helps in understanding their ecological functions and preferred habitats. For example, mycorrhizal mushrooms generally occur in forests and woodlands, while termitophilic mushrooms occur on termitaria (termite nests or mounds) in woodlands, grasslands or arable lands. Indeed, the occurrence and habitat of mycorrhizal mushrooms is entirely dependent on the host termite species (Koné et al. 2011; 2018). In terms of preference by consumers, mycorrhizal and termitophilic species are generally preferred to saprotrophic mushrooms by local population (Degreef et al. 2016).

In the following sections, these fungal groups will be briefly described, and an up-to-date list of species is provided in Additional file 1: Tables S1-S3. Since the culinary and gastronomic qualities of mushrooms may differ with species, the scientific name is an important guide for chefs and consumers. The Additional file 1: Tables S1-S3 should not be viewed as a mere compilation of known species but rather as a useful aid for communication in science and guiding research and development (R&D) efforts including cultivation and commercialization. For example, species that have been reported to be eaten in a number of countries may be considered priority species for R&D. Similarly, species that are symbiotically associated with valuable indigenous fruit trees may be given priority for research and development in agroforestry systems, afforestation and reforestation programmes.

Mycorrhizal species

The mushrooms in this category consist of the fruit bodies of fungi that form obligate symbiotic associations with higher plants (Hall et al. 2011). The fruit bodies include above-ground structures such as chanterelles, puffballs, coral mushrooms, and subterranean structures such as truffles. Mycorrhizal mushrooms account for the majority of edible wild mushrooms compared to their saprotrophic or termitophilic counterparts (Boa 2012;Bourdeaux et al. 2003; Degreef et al. 2020). In total 249 species of mycorrhizal fungi in 52 genera in 26 families have been reported to produce edible mushrooms in Africa, including Madagascar (Table 1). The majority (92%)of the species belong to the class Agaricomycetes and the remaining species (8%) belong to the class Pezizomycetes. Cantharellus congolensis was reported as edible in the largest number of countries (12 countries) followed by Amanita masasiensis (11 countries) and Russula celluata (10 countries) (Table 1; Additional file 1: Table S1). With 46 edible African species, the genus Russula is the most specious among the mycorrhyzal edible mushroom clades, followed by Cantharellus (38 species), Lactarius (36 species), and Amanita (31 species). The taxonomy of many mycorrhizal fungi is incomplete in Africa, and there are many problems in their classification even at the family and genus levels. For example, Cantharellaceae, Hydnaceae, Clavulinaceae and Sistotremataceae formerly treated as separate families in the order Cantharellales, were recently combined and redefined as members of the family Hydnaceae (Cao et al. 2021; Hibbett et al. 2014; Lawrey et al. 2016). Southern African truffles (Pezizales) formerly known to be Terfezia pfeilii and Choiromyces echinulatus under the family Terfeziaceae now belong to the genera Kalaharituber and Eremiomyces, respectively, in the family Pezizacea (Ferdman et al. 2005). Many clades not yet identified to the species level also exist in the literature. Therefore, the list of edible mycorrhizal species is by no means complete.

Mycorrhizal mushrooms are habitat-specific, preferring host plants in either woodlands, rainforests or deserts. Their capacity to form mycorrhizal associations has evolved convergently in 80 fungal lineages with over 30 plant lineages globally (Meidl et al. 2021). Most of the host plants are dominant trees in forests and woodlands (Meidl et al. 2021; Tedersoo and Brundrett 2017). In Africa, mycorrhizal fungi are distributed in open, gallery and rainforests of the Guineo-Congolian basin, Zambezian Miombo woodlands of East and South-Central Africa and Sudanian savannah woodlands. They are symbiotically associated mainly with members of the Caesalpionioidae, Sarcolaenaceae, Dipterocarpaceae, Asterpeiaceae, Phyllantaceae, Sapotaceae, Gnetaceae and Proteaceae (Bâ et al. 2012; Högberg 1982)). Edible mycorrhizal species in the families Pezizaceae, Pyronemataceae, Terfeziaceae, and Tuberaceae are predominantly associated with desert or drylands. Mycorrhizal association benefits the host plant through increased capacity for mineral nutrients and water uptake (Bâ et al. 2012; Thomas et al. 2019; Tsamba et al. 2015). For example, in desert conditions mycorrhizal associations have been reported to increase CO2 assimilation rates and water use efficiency, allowing the host plant to adapt to the harsh environmental conditions (Thomas et al. 2019). Mycorrhizal fungi also serve as extensions of the plant root system, increasing the volume of soil exploited by the plant root and transporting water and nutrients to the plant (Bourdeaux et al. 2003). The fungus, in exchange, gets simple sugars and vitamins from its host plant. Therefore, the fungi cannot live in the absence of the host plant and vice versa. In poor, dry tropical soils mycorrhizal associations are vital for plant growth and survival (Munyanziza 1994). Mycorrhizal fungi also develop an ecological succession in forest plantations (Termoshuizen 1991). Thus, mushrooms associated with seedlings may differ from those associated with adult trees. Some African indigenous fruit trees such as Uapaca kirkiana form symbiotic associations with many species in the genera Amanita, Cantharellus, Lactarius and Russula (Härkönen et al. 1995). Some of the dominant species in the miombo woodlands such as Afzelia, Berlina, Brachystegia, Julbernardia and Isoberlinia are associated with mycorrhizal mushrooms (Degreef et al. 2020; Högberg 1982; Munyanziza and Kuyper 1995). It is such associations that shape the predominance of certain native forests and woodlands.

Termitophilic species

Termitophilic mushrooms live in an obligate symbiotic association with fungus-growing termites of the subfamily Macrotermitinae. The fungus-growing termites consist of over 165 African species (Kambhampati and Eggleton 2000). A total of 28 edible species of termitophilic mushrooms, belong to the genus Termitomyces in the family Lyophyllaceae of the class Agaricomycetes, were recorded in Africa (Table 2). The list of species recorded as edible is probably a fraction of the total number of Termitomyces species on the continent as we have not included clades not yet identified at the species level. There are many instances of such cases in the literature (e.g., Koné et al. 2018; Sitotaw et al. 2020; Soro et al. 2019). For example, Koné et al. (2018) reported 7 unidentified species of Termitomyces sp from Côte d’Ivoire. Using molecular studies Osiemo et al. (2010) estimated the expected diversity of Termitomyces at about 41 lineages across Africa. Mycological surveys continue to identify undescribed species. For example, in a recent study in Côte d’Ivoire, Koné et al. (2018) identified two potentially new species to science. Termitomyces species are notably absent from Madagascar (Buyck 2008). Out of the species confirmed to be edible, Termitomyces striatus was reported as edible in 18 countries, followed by Termitomyces microcarpus (17 countries), Termitomyces clypeatus and Termitomyces schimperi were each reported to be edible in over 15 African countries (Additional file 1: Table S2). Molecular analyses suggest strong specificity of Termitomyces lineages to their host termite genera, namely Odontotermes, Macrotermes, Microtermes, and Pseudacanthotermes (Koné et al. 2018; Osiemo et al. 2010). The highest Termitomyces lineage diversity was found in Odontotermes across eastern and southern Africa (Osiemo et al. 2010). The termites cultivate the fungi in a garden, an assemblage of structures built from chewed up grass and wood inoculated with fungal spores. The fungi in turn break down the cellulose and lignin into a more nutritious compost that serves as termite food (Vesala et al. 2022). Each year, these fungi produce a crop of large mushrooms, which are a delicacy across most of Africa.

Table 2 Number of termitophilic and saprobic edible wild mushroom genera and species reported r in different African countries and the most frequently reported species (details in Additional file 1: Tables S1-S3)

Termitophilic mushrooms are often preferred over other mushrooms and cherished in many African countries for their taste, aroma, attractiveness, and their uses as substitutes for meat or fish, and medicinal values (Obodai et al. 2014; Sileshi et al. 2009). Termitomyces mushrooms were also reported in more countries (Additional file 1: Table S2) than most of the mycorrhizal (Additional file 1: Table S1) and saprobic species (Additional file 1: Table S3). Nutritionally, Termitomyces mushrooms have been reported to have higher protein contents than other edible mushrooms in Nigeria and Malawi (Masamba and Kazombo-Mwale 2010).

Saprobic or parasitic species

Saprobic edible mushrooms in Africa consist of over 203 species belonging to 73 genera in 33 families (Table 2). Schizophyllum commune was the species reported as edible in the largest number of countries (13) followed by Volvariella volvacea, Lentinus tuber-regium and Macrolepiota procera reported from 12, 11 and 10 countries, respectively (Table 2; Additional file 1: Table S3).

Saprobic and parasitic species do not need living hosts; they either grow on decaying material or they are plant pathogenic fungi. Those that grow on dead organic matter can be easily cultivated on artificial media ranging from straw, sawdust to entire logs. They can occur in agricultural land, in pastures, woodlands and forests growing on a variety of decaying material. Some saprobic mushrooms can also be pathogenic. The bracket fungi (also called shelf fungi) fall in this category. Members of the genera Armillaria, Ganoderma, Lignosus, Phellinus and Pycnoporus are some of the pathogenic decay fungi which produce edible brackets (Table 1). At least 12 species of bracket fungi are edible in Africa including 8 species of Armilaria, 4 of Ganoderma and 1 species of each of Lignosus, Phellinus and Pycnoporus (Additional file 1: Table S3). The genus Armillaria in Africa consists of several species, which may differ in host range as well as in pathogenicity (Strouts and Winter 1994). These fungi kill the inner bark and cambium and invade the wood. Armillaria is a potential threat to orchards of wild fruits, particularly where miombo woodlands have been cleared to provide plantation sites (Parker 1978). Ganoderma spp. develop their fruit-bodies close to the stem base or on roots (Parker 1978). By the time the fruit-bodies appear on the trunk, there is already a considerable rot developed in the trunk and the tree death takes place soon after. Piptoporellus baudonii is a recently described macrofungus plant pathogen that attacks cashew trees, Eucalyptus, cassava, Tectona, and some indigenous trees in southern regions of Tanzania and poses a serious threat to agroforestry and livelihood conditions in the area (Tibuhwa et al., 2020). Piptoporellus grows on the base of the tree extending its mycelia to the xylem and phloem vessels, leading to their clogging thus succumbing the attacked plant to complete wilting. Species belonging to the genus Pleurotus have been reported as parasitic on several trees and as the probable cause of plant diseases. The taxonomy of many species is incomplete and there are on-going changes. For example, some members of the genus Pleurotus are confused with Lentinus. According to Buyck (2008) Pleurotus tuber–regium (Rumph. Ex Fr.) Singer is now Lentinus tuber–regium [Fr.] Fr.) and Pleurotus sajor-caju is now Lentinus sajor-caju. Therefore, the number of species we quoted are likely to change as taxonomic revisions take place and new species are described.

Ecosystem services

The ecosystem services of edible wild mushrooms is rarely discussed in the literature, and probably little known to the general public. In the following sections, we will describe the ecosystem services following the Common International Classification of Ecosystem Services (CICES) developed by the European Environment Agency (Haines-Young and Potschin 2017). Accordingly, the ecosystem services of mushrooms fall under three categories: (a) provisioning services, (b) regulation and maintenance services, and (c) cultural services.

Provisioning services

Provisioning services cover all nutritional, non-nutritional and energetic outputs including food, medicine, feed and energy. Edible mushrooms are considered a popular delicacy all over Africa and are collected and consumed when in season (Boa 2004; Rammeloo and Walleyn 1993). Mushrooms have been reported to provide food with low fat (median: 3.2%), high fibre (median: 11.6% of dry weight) and rich in digestible protein and minerals (Tables 3, 4 and 5). In addition, nutriceuticals, a new class of compounds extracted from mushroom, are used as a dietary supplement or for therapeutic applications (Assemie and Abaya 2022; Shiuan 2004; Wandati et al. 2013). In some countries large quantities are consumed every year. For example, in northern Mozambique, average consumption of mushrooms was estimated at 72–160 kg per household per year. Average consumption of Termitomyces schimperi alone was estimated to be 30–35 kg per household per year (Boa 2004). Similarly, in Zimbabwe households eat up to 20 kg in a productive year (Boa 2004).

Table 3 Proximate composition (crude protein, fat, carbohydrate, fibre and ash content (in % on dry weight basis) of wild edible mushrooms from Africa Species
Table 4 Vitamin contents of wild edible mushrooms (in mg/100 g)
Table 5 Sodium (Na), potassium (K), Calcium (Ca), Magnesium (Mg), iron (Fe) and zinc (Zn) concentrations (in mg/100 g on dry mass basis) of wild edible mushrooms reported from different countries

Some edible mushrooms are also used in folk medicine in many African countries (El Enshasy et al. 2013; Thomas et al. 2019). For example, the Kalahari truffle (Kalaharituber pfeilii) endemic to Botswana and Namibia is known to possess strong curative powers by local peoples. San hunters carry dried pieces of its fruiting bodies to use as an oral antidote to the poison of their arrows (Thomas et al. 2019). In Botswana, dried and powdered fruiting bodies are also reportedly used to induce birth in humans and livestock (Khonga and Mogotsi 2007). In Uganda Volvariella speciosa and Podabrella microcarpa are used in traditional medicinal practice (Engola et al. 2006). According to an ethnomycological survey in Uganda, 45% of the respondents consumed Termitomyces microcarpus for its medicinal value, whereas only 5% consumed it just as food (Kabasa et al. 2006). In the same survey in Uganda, 70–87% of respondents mentioned that Termitomyces microcarpus is used as a nutritional supplement for individuals suffering from yellow fever, jaundice or lack of appetite (Kabasa et al. 2006). Among communities living around Ngorongoro and Serengeti National Park in Tanzania, Tibuhwa (2012) found that Termitomyces titanicus, T. letestui, T. eurhizus, and T. auranticus are used for the treatment of different intestinal problems including pain, ulcer, constipation, and stomach ache. Termitomyces microcarpus was used as an immune booster and is given to sick people to speed up recovery, and to breast-feeding mothers to induce lactation (Tibuhwa 2012). In Cameroon, Termitomyces titanicus is dried, mixed with pastry and fed to underweight children (Yongabi et al. 2004). In Ghana, Lentinus (syn. Pleurotus) tuber-regium has been used by traditional herbal doctors for treating underweight children, asthma and high blood pressure (Dzomeku 2009). In Tanzania, Ganoderma applanatum, Ganoderma tsugae, Ganoderma lucidum and Termitomyces microcarpus are the most popular medicinal mushrooms sold in different traditional markets. They are reported to improve health of long ill people, boost immunity, aphrodisiac in men and anti-tumour agents (Tibuhwa 2018). Recent reviews show that many mushrooms offer health benefits through immune modulation, anti‐cancer (Patel and Goyal 2012), anti-oxidant (Assemie and Abaya 2022; El Enshasy et al. 2013; Ferreira et al. 2009), cholesterol lowering and antibacterial (Alves et al. 2012) effects.

Several studies have reported analyses of the nutritional value of edible wild mushrooms in Africa (Tables 3, 4 and 5). Most reports demonstrate that mushrooms are rich in proteins, carbohydrates, minerals, vitamins and dietary fibers but low fat and oil contents (Dimopoulou et al. 2022; Rasalanavho et al. 2020). However, differences in analytical techniques limits comparison of results from different studies. Therefore, we refrain from directly comparing results of the different studies. Instead, we will focus on broad comparisons with staple crops. In this section, we will briefly review studies on proximate composition (proteins, carbohydrates, fats), vitamins and antioxidants and mineral elements reported from African mushrooms.

Proteins

Malnutrition due to protein deficiency is a major factor for the high mortality and morbidity in Africa. For example, in much of eastern Africa the normal diet is cereal-based while in central and western Africa it is often cassava-based. Animal protein is beyond the reach of many poor families in this region, which constitute a large proportion of the population. Mushrooms can provide an alternative source of proteins for such communities. In the studies we reviewed, the total protein content varied widely from a low of 2% in Termitomyces tyleranus to 60% in Lepista nuda (Table 3), and a median value of 19.4%. The values reported for some species are comparable with those reported for protein-rich foods such as soybean (12–40%), cowpea (22.5%) and lima bean (23.3%) seeds but much higher than the staple cereals such as maize (9.4%) and rice (7.3%). This indicates that mushroom protein can supplement the low protein diets of poor people (Nakalembe et al. 2015). Some studies (e.g., Degreef et al. 1997; Nakalembe and Kabasa 2013) have shown that mushroom protein contains most essential amino acids, including lysine which is limiting in cereals. For example, in one study in Uganda the lysine content was 6.7% of protein dry weight in Polyporus tenuiculus, 6.5% in Termitomyces microcarpus, 6.1% in Termitomyces globulus and 5.5% in Termitomyces eurrhizus. (Nakalembe and Kabasa 2013). Wild mushrooms could be a good source of proteins for malnourished children.

Fatty acids

The total fat content reported for edible mushrooms varied from a low of 0.2% in Cantharellus cibarius to a high of 14.3% in Clitocybe odora (Table 3) with a median value of 3.2% and comparable with those of maize (4.2%). However, mushroom fat consists of predominantly unsaturated fatty acids (Dimopoulou et al. 2022; Nakalembe and Kabasa 2013). In an analysis of various wild edible mushrooms in Uganda (Nakalembe and Kabasa 2013) and Zambia (Bourdeaux et al. 2003) unsaturated fatty acids dominated over saturated ones. In Zambia, the percentage of unsaturated fatty acids varied between 69 and 82% of the total fatty acids in all 12 species analysed, and the ratio of monounsaturated to saturated fatty acids was closer to those recognized as good for human consumption (Bourdeaux et al. 2003). Global analysis of edible mushroom also shows that unsaturated fatty acid levels are generally greater than saturated ones, regardless of the continent where the mushroom is cultivated or harvested (Sande et al. 2019).

Carbohydrates

Global analyses have shown that mushrooms are nutritionally well-balanced sources of carbohydrates and proteins, with low fat concentrations, making them very healthy foods (Sande et al. 2019). The carbohydrate content reported for the different species vary between 4.2 and 80.2% with a median value of 51.5% on dry weight basis (Table 3). The reported values for Auricularia auricular, Russula celluata, Termitomyces robustus, Lentinus squarrosulus, and Lentinus tuber-regium are over 75%. Carbohydrates are contained in the dry matter, mainly as chitin, polysaccharides and sugar alcohols. The presence of sugar alcohols makes mushrooms a good substitute for high energy sugar in diabetic patients.

Vitamins

Only a small number of studies (Musieba et al. 2013; Nakalembe et al. 2015; Egwim et al. 2011; Mshandete and Cuff 2007) have reported the vitamin contents for wild edible mushrooms in Africa. The limited number of studies (Table 4) indicate a good profile of vitamins, including thiamine (vitamin B1), riboflavin (vitamin B2), nicotinic acid (vitamin B3), ascorbic acid (vitamin C) and folic acid (vitamin B9). The values reported for thiamine ranged from 0.05 mg/100 g in Termitomyces clypeatus to 0.94 mg/100 g in Polyporus tenuiculus. The riboflavin, nicotinic acid and ascorbic acid contents did not vary widely among mushroom species (Table 4). On the other hand, the folic acid content varied widely from a low of 70 μg/100 g in Termitomyces tyleranus to 380 μg/100 g in Termitomyces clypeatus (Table 4). Folic acid deficiency is one of the commonest deficiencies worldwide affecting women on oral contraceptives, pregnant mothers, the elderly, alcoholics and children. Folic acid bioavailability in mushrooms is good compared to some vegetables. Thus, mushrooms hold tremendous potentials for alleviating deficiencies (Nakalembe et al. 2015).

Minerals

Many studies have reported the ash contents rather than individual minerals (Table 3). Although the ash content is an indicator of the mineral quantities in the different species, it is not adequate to determine individual mineral contents. From the few studies that reported individual nutrients, it is apparent that mushrooms contain reasonable amounts of sodium (Na), potassium (K), phosphorus (P), iron (Fe), calcium (Ca), magnesium (Mg) and copper (Cu) (Table 5). This indicates that mushrooms may play a crucial role in alleviating mineral deficiencies. Some minerals are present in smaller quantities than others, and the mineral contents also vary with the species of mushroom. Information on concentrations of other micronutrients such as copper (Cu), selenium (Se) and zinc (Zn) is scarce. The only studies we found were Rasalanavho et al. (2020) and Rasalanavho (2021) on wild edible mushrooms in South Africa, who found Boletus edulis to be rich in Se contributing 145–836% towards the recommended dietary allowances (RDA). They also found Suillus luteus to be rich in Fe contributing 162% towards the RDA, and Clavulina cristata and Helvella crispa to be rich in Cu and Zn contributing 186% and 193% towards the RDAs, respectively (Rasalanavho et al. 2019).

Antioxidants

Mushrooms are known to contain chemicals with antioxidant properties such as phenolic compounds, alkaloids, saponins, flavonoids, tannins, sterols, triterpenes, coumarins and cyanogenic glycosides (Egwim et al. 2011; Ehssan and Saadabi 2012). Antioxidant activity has been measured using different methods including total antioxidant activity, reducing power, DPPH assay, metal chelating and active oxygen quenching assay (Egwim et al. 2011; Wandati et al. 2013; Woldegiorgis et al. 2014). For example, Egwim et al. (2011) analysed the antioxidant activity of the ten wild edible mushrooms in Nigeria and found the highest antioxidant activity in Laccaria amethysta (Table 6). Wandati et al. (2013) characterized the total polyphenol and flavonoid contents and free radical activity in 10 edible wild mushrooms and two commercially grown mushrooms in Kenya and found total polyphenol values of 210–1614 mg gallic acid equivalent /100 g on dry weight basis. The flavonoid values were in the range of 214–1695 mg quercetin equivalent /100 g (Wandati et al. 2013). Similarly, Woldegiorgis et al. (2014) reported higher antioxidant properties in wild mushrooms than in cultivated mushrooms in Ethiopia. In Tanzania, Tibuhwa (2012) analyzed qualitative and quantitative values of antiradical and antioxidant contents in crude methanolic extracts of six Termitomyces species using DPPH 1, 1-diphenyl-2-picrylhydrazyl free radical as a substrate. The highest antiradical activity unit EAU515 1.48 was recorded in T. microcarpus and the lowest (EAU515 0.7) in T. eurhizus. The scavenging power was also found to be highest in T. microcarpus (EC50 < 0.1 mg/ml) and least in T. eurhizus (EC50 = 0.36 mg/ml) (Tibuhwa, 2012).

Table 6 Antioxidant contents of wild edible mushrooms

The antioxidant properties of most edible mushrooms have not been well studied. Even where such studies exist (Table 6), the variables measured vary from study to study, and there is no consistent approach of reporting. Information is also lacking on the antimicrobial properties of many mushroom species. This highlights the need for urgent research using standardized and consistent methodology into novel compounds with nutritional and medicinal values in edible African mushrooms.

Regulation and maintenance services

Regulation and maintenance services constitute all the ways in which organisms mediate the physical, chemical and biological conditions that affect human health, safety or comfort. Fungi in general play an important role in plant health, tree regeneration and the structure and functioning of forest ecosystems (Meidl et al. 2021; Pérez-Moreno et al. 2021). For example, mycorrhizal fungi may promote tree growth, forest sustainability, biodiversity and contribute to mitigation of greenhouse gas emissions through the maintenance of forest biomass (Anthony et al. 2022; Martin et al. 2016; Pérez-Moreno et al. 2021). Mycorrhizal fungi provide these services by facilitating nutrient and water uptake through underground mycelial networks that connect trees and other plants, increase nutrient availability, and metabolites that enhance protection against pathogens and abiotic stress (Pérez-Moreno et al. 2021). In nutrient-limited environments, these fungi help trees in acquiring limited nutrients and water from the soil beyond the reach of tree roots. As the levels of bioavailable nutrients in forest soils are often too low to sustain plant growth, most trees rely on mycorrhizal fungal symbioses (Martin et al. 2016). This promotes tree establishment and growth. This has been demonstrated through controlled experiments, for example, in the establishment and growth of exotic pines and Australian acacias in Africa (Bâ et al. 2010).

The mycelia of mycorrhizal fungi are also considered to make a large contribution to nutrient uptake and cycling in many ecosystems. The mycelia extend from tree root surfaces to much greater distances through the soil than root hairs, and act as a secondary root system that facilitates the uptake of nutrients especially phosphorus, zinc and sulphur by the host. Mycorrhization nearly always results in an improved P nutrition and water status, which leads to a healthy plant growth (Godbold and Sharrock, 2003). There is also evidence of direct uptake of phosphorus from litter via mycorrhiza, such that the soil pathway is bypassed and an entirely closed cycle exists (Herrera et al. 1978). Ectomycorrhizal fungi are adapted to systems in which nitrogen is limiting, and all nutrients and water taken up by the plant goes through the fungus. As such mycorrhizal fungi have the potential to determine plant community structure (van der Heijden et al. 1998). Common associations of particular tree species for specific sites are strongly linked by the endemic mycorrhizal fungi in the area. Interconnection of trees of different species has been reported to be through common mycelia to facilitate transfer of carbon, nitrogen and phosphorus. It is this interconnectedness that sustains the existence of certain woodlands and forests. For example, the trees of the miombo woodland of central and southern Africa would not exist without their fungal partners (Boa 2004). Recent analyses (Suz et al. 2021) suggest that changes in ectomycorrhizas can lead to a tipping point in such forests and affect several ecosystem processes directly linked to human wellbeing.

Termitophilic fungi occur in forest land, woodlands, grassland and arable land in association with fungus-growing termites, which are keystone species of African and Asian tropics. They induce vegetation heterogeneity directly or indirectly through their nest-building and foraging activities, associated nutrient cycling and their interaction with mammalian herbivores and fire (Sileshi et al. 2010). The ecological success of these termites is linked to their obligate mutualism with Termitomyces fungi (Osiemo et al. 2010). Termitomyces symbionts produce cell wall degrading enzymes and provide protein-rich nutrition especially for the termite larvae and the queen (Vesala et al. 2022). This association between Termitomyces and termites is particularly important in savannas of Africa, where termitaria create islands of fertility (Sileshi and Arshad 2012) favourable for tree regeneration (Sileshi et al. 2010). A growing body of literature suggests that embracing termite-induced vegetation heterogeneity may increase the robustness of drylands (Bonachela et al. 2021; Sileshi et al. 2010). Termite control using fungicides targeted at the Termitomyces spp. (Sileshi et al. 2009) may have negative impacts on the ecosystem services of termites including production of edible mushrooms.

Saprotrophic fungi are integral parts of the soil microbial communities, which play a crucial role in the decomposition of wood and forest litter and recycling the nutrients. Saprotrophic fungi especially play a pivotal role by secretion of extracellular enzymes targeting the decomposition of insoluble remains of other biological entities including the complex components of plant debris, such as cellulose, lignocellulose and chitin, which makes them central in nutrient recycling (Hartl et al. 2012). Nutrient recycling in turn sustains the productivity of agricultural land and forests.

Cultural services

Cultural services include non-material and non-consumptive benefits that affect physical and mental states of people through spiritual enrichment, cognitive development, reflection, recreation, and aesthetic experience. Mushrooms play an important role in cultural practices (Morris 1992; Engola et al. 2006). For example, according to Engola et al. (2006) Termitomyces microcarpa (Podabrella microcarpa) has an important cultural significance in respect to traditional ceremonies in Rakai District of Uganda. According to Onguene and Kuyper (2019) some species are used in rituals. For example, Lentinus (Pleurotus) tuber-regium is used as a love charm. Some Termitomyces species are also used to drive away bad spirits in Cameroon (Onguene and Kuyper 2019).

Anthropogenic threats to diversity and utilization of wild mushrooms

Some studies have documented decline in certain wild edible mushroom species in Africa (Dube et al. 2021; Guissou et al. 2008; Onguene and Kuyper 2019). For example, in a survey conducted in Binga district of Zimbabwe, 92.9% of the respondents believed the availability of wild edible mushrooms has decreased (Dube et al. 2021). Focus group discussions with elders and women also indicated extinction of certain species (e.g., Termitomyces titanicus and Lactarius kabansus) due to loss of native woodlands, anthills and indigenous trees (Dube et al. 2021). In Burkina Faso, Guissou et al. (2008) documented decrease in the gathering and consumption of wild edible mushrooms apparently due to decline in mushroom populations as a result of disappearing forest habitats. Some species have also been listed as vulnerable by the International Union for Conservation of Nature (IUCN). For example, Cantharellus afrocibarius was assessed as vulnerable under criterion C2a (ii) of the IUCN Red List (http://iucn.ekoo.se/iucn/species_view/800901/). The most pressing threat is the destruction of its habitat through logging and clearance for agriculture (Jew et al. 2016; Syampungani et al. 2009).

Increase in population has driven expansion of agriculture and mining into forest land, and this is now a major driver of deforestation and decline in biodiversity of mycorrhizal fungi (Thomas and Vazquez 2022). Many forests and woodlands where mushrooms used to thrive are currently undergoing severe degradation and fragmentation due to human activity including land clearing for agriculture, conversion to other land uses, charcoal burning and extraction of timber. A growing body of research is showing that the miombo woodlands in southern and central Africa are being degraded due to clear-felling for agriculture, charcoal production and logging (Jew et al. 2016; Syampungani et al. 2009). In some countries, high human utilisation has resulted in decreases in species richness, abundance, diversity and loss of large trees and key miombo species (Jew et al. 2016). Some tree species such as Pterocarpus angolensis are now listed as Threatened in the IUCN Red List (IUCN). Many of the tree species are slow-growing and have very low recruitment levels. When host trees are felled, their fungal partners become particularly vulnerable (Degreef et al. 2020). When forests are cleared and the mycorrhizal tree species dies, the associated mycorrhizal fungi die. Without the right host, the fungus also does not reach stages of mushroom production. As agricultural operations disturb mycelial networks some mushrooms are also lost. Consequently, deforestation often leads to decline in mycorrhizal fungi and therefore disappearance of the mushrooms (Bourdeaux et al. 2003; Degreef et al. 2020). For example, as soon as miombo woodlands were converted to crop fields or pasture lands, the typical ectomycorrhizal mushrooms were lost irreversibly (Degreef et al. 2016). Fallowing creates land mosaics characterized by vegetation communities at different stages of succession (Tsamba et al. 2015). Mycorrhiza formation and mushroom production are also highly sensitive to soil disturbance including cultivation, weeding and the use of fungicides, herbicides and fertilizers (Amaranthus 1992). Overgrazing can also lead to decline in some mycorrhizal mushrooms. For example, the quantities of Kalahari truffles (Kalaharituber pfeilii) harvested have been declining where livestock have been concentrated (Trappe et al. 2008).

In the case of the termitophilic mushrooms, the greatest threat is the indiscriminate use of termite control practices such as mound destruction, use of insecticides including persistent organic pollutants (Sileshi et al. 2009). Some termite control practices involve fungicides targeting the Termitomyces spp. (Rouland-Lefevre and Mora 2002). This may drastically reduce both termites and fungi and ultimately decline in production of edible mushrooms. The conversion of forest lands to agriculture is also likely to have an effect on Termitomyces mushrooms as agricultural practices often negatively affect termitaria (Sileshi et al. 2010). Saprobic mushroom species need a constant supply of suitable organic matter to sustain production in the wild. Clearing of forests for agriculture and bush burning removes the organic matter necessary for the mushroom growth, and this can limit their production. Frequent bushfires that decimate suitable substrates (e.g., leaf litter and rotting trunks) also pose threats to saprobic mushrooms (Degreef et al. 2020).

The loss of indigenous knowledge also poses challenges to the conservation and sustainable utilization of edible mushrooms. Much of the knowledge on edible mushrooms remains with the local people, while scientific information on their uses and products remains inadequate. Many communities in Africa have an elaborate knowledge of the association between termites and edible mushrooms (Sileshi et al. 2010), and their nutritional and medicinal values (Kabasa et al. 2006; Oso 1975; Yongabi et al. 2004). Places where specific mushrooms can be found in abundance are often determined using indigenous knowledge and experience (Boa 2004). For example, desert truffles are subterranean, and can only be collected by people with traditional knowledge and expertise (Thomas et al. 2019; Trappe et al. 2008). Thomas et al. (2019) describes some unusual practices used for collecting truffles in rural Morocco. Ethnomycological surveys (e.g., Tibuhwa 2013; Mlambo and Maphosa 2022) show that women possess vast knowledge of mushroom folk taxonomy, biology and ecology and are therefore the principal knowledge transmitters. A knowledge of potential collecting sites and habitat preferences for edible species is an advantage to a collector (Boa 2012). Such knowledge often resides among older women. However, the knowledge and skills are lost when the older generation dies or a community breaks down due to war or natural disasters. When people move to cities, they also lose their confidence in recognising mushroom species or acquire prejudices about mushrooms (Boa 2012). Thus, the loss of knowledge could limit market opportunities or dietary choices.

Species that are highly prized in one community may be ignored due to lack of local knowledge on their edibility. For example, Boletus edulis growing in Malawi are not collected and eaten or sold locally (Boa 2012), while it is reported as edible only in Mozambique and Zimbabwe (Additional file 1: Table S1). Outside Africa, Boletus edulis is highly appreciated due to its high nutritional value and its exceptional flavour (Dimopoulou et al. 2022).

Opportunities for conservation and cultivation

Sustainable management of forests and woodlands, and protection of termitaria is an important step in the conservation of edible mushrooms and the long-term sustainability. In that sense, the REDD + and Clean Development Mechanism (CDM) frameworks provide an opportunity for international conservation initiatives to support mushroom conservation. For example, national REDD + demonstrations on sustainable forest management could include training and demonstrations on the linkages between mushrooms and trees. Such projects may also support value addition and commercialization of wild mushrooms to generate revenue by communities. According to Tibuhwa (2013), mushroom mongers in Tanzania earn up to $400–900 annually. This earning contributes significantly to the rural economy (Bourdeaux et al. 2003; Tibuhwa 2013). The woodland ecosystem delivers substantial amounts (100–300 kg/ha/year) of edible fungi (all taxa), but this function is annihilated by large-scale charcoal production. The cash generated from charcoal conversion does not outweigh the income accumulated from mushroom harvesting (Milenge and de Kesel 2020).

While a large proportion of the rural households may be involved in collecting mushrooms from the wild, very little efforts have been made to cultivate them. For example, in Zambia over 42% of households collect mushrooms compared to 29% who collect edible fruits, roots, tubers and bulbs from the forest (ILUA 2016). Mushroom collecting is done during the rainy season usually by women and children (Boa 2012; Mlambo and Maphosa 2017; Oso 1975; Wandati et al. 2013). While wild harvesting may be sustainable for local consumption and sale, there is a danger of overharvesting (Hall et al. 2003) if it is done for commercial purposes. Successful cultivation of native wild edible mushrooms can provide a guaranteed supply and uniformity of product for commercial production especially for the canning industry. Saprobic mushrooms hold considerable potential for cultivation as they can be grown on artificial media (Boa 2004; Buyck 2008). However, their cultivation on a commercial scale has been achieved in hardly any African country (Mlambo and Maphosa 2022). The cultivation of saprobic mushrooms offers prospects for using agricultural wastes such as wheat and rice straws, bean and soya bean stalks, maize stover, cotton husks, banana pseudostems and other lignocellulosic products as growing media (Mlambo and Maphosa 2022). The spent mushroom substrate (residues used for mushroom cultivation) can be used for soil application as a fertilizer or as animal feed. There is a huge potential for cultivation under smallholder farmers especially in the cooler parts of East Africa and West Africa (Degreef et al. 2016; Mlambo and Maphosa 2022). However, general lack of understanding of the trophic relationships, biotic, edaphic and climatic requirements mycorrhizal mushroom (Hall et al. 2003) and lack of low-cost approaches to the cultivation has hindered progress (Mlambo and Maphosa 2022). Potential research areas for saprophytic mushrooms include (1) search for candidate species, (2) determination of optimum growth conditions, and (3) post-harvest management.

On the other hand, cultivation of ectomycorrhizal and termitophilic mushrooms is challenging because they do not grow on artificial media. In addition to their requirement for a host tree, mycorrhizal mushrooms also have unique interdependences on other microorganisms that grow within the mushroom’s fungal tissues. These additional symbiotic relationships further complicate artificial cultivation, and thus there are no successful examples of domesticated ectomycorrhizal mushrooms. Therefore, wild harvesting has been the only practical option. However, opportunities exist for cultivation of ectomycorrhizal mushrooms in agroforestry systems and plantations where tree roots are deliberately infected with the desired ectomycorrhizal fungus (Thomas and Vazquez 2022). This has been successfully done with truffles managed under controlled conditions in orchards in the USA and Italy (Boa 2004). There are also many examples of opportunistic production and utilization of ectomycorrhizal mushrooms growing in plantations of exotic pines (Pinus spp.) and eucalypts (Eucalyptus spp.) and agroforestry arrangement in Africa (Boa 2012; Buyck 2008; Rasalanavho 2021). Highly prized species such as Boletus edulis, Suillus and Rhizopogon species are harvested commercially from pine plantations in Madagascar, Malawi and South Africa (Boa 2012; Buyck 2008; Rasalanavho 2021). Russula and Cantharellus, the most commonly consumed and economically important mushroom genera in Madagascar, were also found in abundance in plantations of Eucalyptus robusta (Buyck 2008). This highlights that such plantations remain underutilized for production of edible mycorrhizal mushrooms. A mixture of trees of different species and age group need to be inoculated with the right spectrum of fungi. However, our knowledge of the right combination is very limited. This is a fertile area for future research.

Limitations of this review

Although we tried to be as exhaustive as possible in our search for the literature, we probably have not captured all the relevant information especially from the grey literature. We depended on Google Scholar when searching for grey literature because it can find much grey literature and specific, known studies (Haddaway et al. 2015). Google Scholar is also known to identify the majority of literature identified using Web of Science (Haddaway et al. 2015). However, the various search engines are known to miss some important literature. We also did not search for literature published in languages other than English. Therefore, we do not claim the list of species is complete as reports from West and Central Africa often published in French may have been missed.

In addition, the taxonomy of many species is incomplete, and our list of species has not covered taxa that have not been identified to the species level. In many ethnomycological studies (e.g., Bourdeaux et al. 2003; Sitotaw et al. 2020; Soro et al. 2019), a number of wild edible mushrooms were indicated as “sp.” awaiting identification to the species. For example, Bourdeaux et al. (2003) reported 2 unidentified Lactarius sp and one for each of Macrolepiota, Pleurotus, Psathyrella, Russula and Xerocomus sp from Zambia. Similarly, Soro et al. (2019) reported 2 unidentified Agaricus sp, 3 Auricularia sp, 2 Cookeina sp and 2 Lycoperdon sp from Côte d’Ivoire. We have not included such species in our list due to the uncertainty about their identity and to avoid double counting when enumerating the diversity. Detailed ethnomycological studies and taxonomic work have been conducted only in a small number of African countries. Even in countries where such surveys exist, they do not cover the entire country. We strongly believed that many species new to science are likely to be added as more ethnomycological surveys are conducted and taxonomic revisions are made. Therefore, the number of species we quoted in Table 1 and Additional file 1: Tables S1-S3 are likely to represent only a fraction of the wild edible mushroom diversity.

Conclusions and recommendations

It is concluded that the African continent holds a great diversity of neglected and underutilized edible wild mushroom. It is also concluded that the various species provide ecosystem services that have remained undervalued. With the increasing emphasis on health foods, prevention and natural cures for human diseases, mushrooms are gaining importance in global trade, and this presents opportunities for cultivation and value-addition. Mushrooms collected from the wild can never be uniform and reliable as production can vary considerably from year to year, and this can severely limit opportunities for commercial exploitation. Therefore, investment in research and development is urgently needed on the cultivation of African mushrooms to ensure quality and consistent supply of raw materials. We strongly recommend integration of mycorrhizal mushroom production with commercial tree plantations, agroforestry and domestication of indigenous fruit trees. Such integration can provide opportunities to diversify household income, nutrition and food security while also providing climate change adaptation and mitigation benefits of trees. Therefore, we recommend national forestry research and development programs and international frameworks such as Reducing Emissions from Deforestation and Degradation (REDD +) to invest in the conservation, cultivation and valorisation of wild edible mushrooms to achieve sustainable forest management and the welfare of local communities. An emerging issue from the review is the poor linkage between studies focusing on analysis of nutritional and medicinal values and taxonomic work. In some cases, species were either misidentified or outdated names were used when reporting proximate composition and mineral content of mushrooms. Therefore, we strongly recommend investment in ethnomycological surveys and taxonomic work to support conservation and utilization of wild edible mushrooms.

Availability of data and materials

The datasets used and/or analysed during the current study are available as Additional file Online Materials.

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Acknowledgements

The authors would like to thank Addis Ababa University, University of Dar es Salaam and Lupane State University for the support provided to the co-authors.

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GWS, DDT and AM conceived the study. GWS led the review and write up of the manuscript with inputs from DDT and AM. All authors contributed critically to the drafts and finalized the text. All authors read and approved the final manuscript.

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Additional file 1: Table S1.

Edible wild ectomycorrhizal mushrooms and reported from different countries in Africa. Table S2. Edible wild termitophilic mushrooms and confirmed reports from different countries in sub-Saharan Africa. Table S3. Edible wild saprobic mushrooms and confirmed reports from different countries in sub-Saharan Africa.

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Sileshi, G.W., Tibuhwa, D.D. & Mlambo, A. Underutilized wild edible fungi and their undervalued ecosystem services in Africa. CABI Agric Biosci 4, 3 (2023). https://doi.org/10.1186/s43170-023-00145-7

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