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Article

Forest Type and Site Conditions Influence the Diversity and Biomass of Edible Macrofungal Species in Ethiopia

by
Gonfa Kewessa
1,2,3,
Tatek Dejene
1,4,
Demelash Alem
5,
Motuma Tolera
3,4 and
Pablo Martín-Pinto
1,*
1
Sustainable Forest Management Research Institute, University of Valladolid, Avda. Madrid 44, 34071 Palencia, Spain
2
Department of Forestry, Ambo University, Ambo P.O. Box 19, Ethiopia
3
Wondo Genet College of Forestry and Natural Resources, Hawassa University, Shashemene P.O. Box 128, Ethiopia
4
Ethiopian Forestry Development, Addis Ababa P.O. Box 3243, Ethiopia
5
Ethiopian Forest Development Bahir Dar Center, Bahir Dar P.O. Box 12627, Ethiopia
*
Author to whom correspondence should be addressed.
J. Fungi 2022, 8(10), 1023; https://doi.org/10.3390/jof8101023
Submission received: 26 August 2022 / Revised: 19 September 2022 / Accepted: 24 September 2022 / Published: 28 September 2022
(This article belongs to the Special Issue Fungal Taxonomy, Phylogeny, and Ecology: A Themed Issue Dedicated to Academician Wen-Ying Zhuang)
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Abstract

:
Ethiopian forests are rich in valuable types of non-wood forest products, including mushrooms. However, despite their nutritional, economic, and ecological importance, wild edible mushrooms have been given little attention and are rarely documented in Ethiopia. In this study, we assessed mushroom production levels in natural and plantation forests and the influence of climate and environmental variables on mushroom production. Sporocarps were sampled weekly from July to August 2019 at a set of permanent plots (100 m2) in both forest systems. We analyzed 63 plots to quantify sporocarp species’ richness and fresh weight as well as to elucidate the degree of influence of forest types and site conditions, including soil and climate. Morphological analyses were used to identify fungi. In total, we recorded 64 wild edible fungal species belonging to 31 genera and 21 families from the plots established in the natural and plantation forests. A significantly greater total number of edible fungi were collected from natural forests (n = 40 species) than from plantations. Saprotrophs (92.19%) were the dominant guild whereas ectomycorrhizal fungi represented only 6.25% of species. Ecologically and economically important fungal species such as Agaricus campestroides, Tylopilus niger, Suillus luteus, Tricholoma portentosum, and Morchella americana were collected. The sporocarp yield obtained from plantation forests (2097.57 kg ha−1 yr–1) was significantly greater than that obtained from natural forests (731.18 kg ha−1 yr–1). The fungal community composition based on sporocarp production was mainly correlated with the organic matter, available phosphorus, and total nitrogen content of the soil, and with the daily minimum temperature during collection. Accordingly, improving edible species’ richness and sporocarp production by maintaining ecosystem integrity represents a way of adding economic value to forests and maintaining biological diversity, while providing wood and non-wood forest products; we propose that this approach is imperative for managing Ethiopian forests.

1. Introduction

The Ethiopian highlands were once covered with dense natural high forests [1,2]. However, anthropogenic influences associated with the expansion of farming and human habitation have brought about deforestation and forest degradation [2], resulting in dramatic changes in the abundance and extent of native forest types. What remains is highly fragmented and frequently modified by non-native fast-growing trees such as Eucalyptus and Pinus species [3]. However, an important feature of these natural and plantation forests is their richness in valuable types of non-timber forest products (NTFPs) [4,5,6]. Various studies have revealed long lists of NWFPs, indicating their importance, contribution, and use by rural communities in Ethiopia [7,8]. If managed and conserved properly, NTFPs could potentially support the livelihoods of rural people by providing a source of income as well as food and medicine [7]. The most important NTFPs include honey, bees-wax, bamboo, coffee, spices, edible plant products such as fruits, seeds, and fodder, medicinal plants, various extractives and flavorings [7,9], and wild edible mushrooms [10,11].
More than 2000 fungi are known to produce edible sporocarps [12]. Edible wild mushrooms are not only a source of nutrition but also can be an important income generator [12,13], helping rural people to reduce their vulnerability to poverty and strengthen their livelihoods by providing a reliable source of revenue in many regions of the world [14,15,16]. In addition, edible mushrooms also form the basis of many manufactured products, including medicine, and are the focus of a new wave of tourism resulting from recreational programs linked to nature. Furthermore, wild mushrooms in general are important components of biodiversity in forest systems [17], playing an essential ecological role in forest communities. In particular, mycorrhizal associations have significant effects on nutrient and water uptake, growth, and plant survival [18,19], improving soil aeration and porosity [20] and resistance to pathogens [21], and also providing food for many organisms [22]. Saprotrophic fungi are essential for the decomposition of dead matter and, therefore, for nutrient cycling in forest ecosystems [23]. They are also important in the cycling of various elements, such as carbon, nitrogen, and oxygen [17].
There has been a tradition of ethnomycological usage among communities that dwell in Ethiopian forests [10,11,24]. However, despite all the benefits, wild mushrooms are the most neglected of Ethiopia’s NTFP resources. To date, there have been few studies of wild mushrooms and they are rarely documented [25,26], which may reflect that forest resource management has been primarily based on the production of wood products [8,9,27]. Wild mushroom formation is influenced by various factors, such as the forest type, host plant, nutritional status of the mycelium, and environmental factors [28,29,30]. Climatic variables also influence mushroom production because the development of fungal fruit bodies is dependent on soil temperature and the availability of surface water [31,32]. In addition, sporocarp composition is strongly determined by soil nutrients and chemical properties [33,34]. This is particularly the case for saprotrophic fungi, which are more dependent on their respective substrates than mycorrhizal fungi [35]. Forest management can, therefore, play a crucial part in shaping macrofungal communities because it can modify vegetation parameters such as tree density, canopy cover, understory plant communities, and soil conditions [30,33].
The negative impact of plantation forests is a common narrative in Ethiopia. Among these criticisms, the most cited is the lack of plant diversity in plantation forests [36,37]. However, our knowledge and understanding of fungal communities in plantation forests are limited compared to our knowledge of those in natural forests. Such information is essential to encourage forest management practices that include the conservation and production of valuable mushrooms through the adoption of a mycosilvicultural approach [38,39] for plantation and natural forest systems in Ethiopia. However, few studies have assessed the different environmental factors required for the production of wild edible mushrooms in these forest systems. To date, most have been performed at local scales and investigated a limited set of environmental variables. To gain a deeper understanding of the different environmental factors driving fungal communities as well as host plants, we should consider a wide range of variables from broad geographic areas. This should enable the development of strategies to manage different forests in different landscapes, to promote sporocarps production in the country. Thus, the main goal of this study was to assess the diversity of valuable macrofungal species in different areas of the country and compare the edible macrofungal species’ richness, sporocarp production, and community assemblages in native mixed forests and non-native fast-growing plantation forests. We hypothesized (1) that forest type is important for structuring macrofungal communities and the production of their sporocarps at small spatial scales in Ethiopian forest ecosystems. We expected that the macrofungal community would differ between the two forest types, resulting in an overall higher diversity value for study sites in natural forests because fungal diversity would be driven mainly by vegetation type, substrate availability, and other environmental variables [40,41,42]. However, the shorter rotation period of trees in plantation forests would also result in nutrient stress in these forests [43], which would favor only certain macrofungal species, such as ectomycorrhizal species [44]. Thus, we hypothesized that the overall sporocarp biomass produced in plantation forests would be higher than that in natural mixed forests, particularly those of potentially marketable species such as Suillus luteus. Differences in environmental variables such as edaphic and climate variables are the main driving forces for edible mushroom communities and, hence, would be important for the maintenance of their local production and diversity [45,46]. However, in the dry Afromontane region, water is scarce for a long period each year, implying that sporocarp development by some species would be limited. Thus, we also expected (2) that climate variables would govern fungal communities more than edaphic variables in our study areas [47]. When we set out, our specific objectives were: (i) to analyze the richness and production of sporocarps according to forest types, and (ii) to assess the influence of edaphoclimatic variables on taxa composition. The information generated from this study should highlight the economic value of edible mushrooms as NTFPs and encourage forest managers to manage these forests sustainably to promote sporocarp development and maintain the biological diversity of Ethiopian forests.

2. Materials and Methods

2.1. Study Sites

The study was conducted in five dry Afromontane areas in Ethiopia: the Taragedam, Alemsaga, Banja, Wondo Genet, and Menagesha Suba forests (Figure 1). Comprehensive descriptions of the forests are provided in Table 1. Some pictures of the forest types where the sporocarps collection was carried out in the study areas are provided in Figure 2 and Figure 3.

2.2. Experimental Design and Sporocarp Sampling

In total, 63 sample plots were established at seven sites, nine plots in each of the natural and plantation forests described in [53,54,55]. The plantation forests are located in the Wondo Genet and Menagesha Suba areas, while the natural forests are found in the Wondo Genet, Taragedam, Banja, and Alemsaga areas. The plantations mainly constitute the Eucalyptus grandis, Pinus patula, and Pinus radiata tree species (Table 1). The main tree species in the dry Afromontane natural forests include Juniperus procera, Podocarpus falcatus, Hagenia abyssinica, and Olea africana in the Wondo Genet natural forest area [27]. The dominant tree species in the natural forests of Taragedam, Banja, and Alemsaga, meanwhile, include Maytenus obscura, Carissa edulis, Olea sp., Acacia abyssinica, Buddleja polystachya, Acacia nilotica, Albizia gummifera, Prunus africana, and Brucea antidysenterica. These trees serve as the main sources of timber for the country [27] and thus indicate a need for the sustainable management of these forests. Each plot in each forest type was rectangular in shape (2 m × 50 m) and covered an area of 100 m2. Within each of the selected forest stands, we studied three different blocks with three plots per block. The plots were established at least 500 m apart and were laid out randomly in the forests to avoid confounding spatial effects inherent to such a plot-based design [56,57] and to reduce environmental heterogeneity [58]. The plots were analyzed as independent samples as suggested by Ruiz-Almenara [59]. All fungal fruit bodies found were harvested weekly during the major rainy season through July and August 2019. Fresh weight measurements were taken in situ using a digital sensitive balance (SF-400) to determine the fruit body production in kilograms per hectare per year. The number of sporocarps of each species in each plot was also recorded. Specimens were photographed in the field, and their morphological features and ecological characteristics were noted to facilitate taxonomic identification in the laboratory [60]. Specimens of each macrofungal species were taken to the laboratory and dried to preserve as herbaria specimens, then used for species identification purposes.

2.3. Species Identification and Characterization

In this study, morphological analyses were used for taxa identification. In the laboratory, fruit body tissues and spores were examined using an Optika B-350PL microscope (Optika, Pontenarica, Italy). Monographs were also used for taxa identification [61,62,63,64,65,66,67,68,69]. Up-to-date fungal taxa names and authors’ names were obtained from the Mycobank database (http://mycobank.org (accessed on 1 November 2020). Trophic levels were assigned to species using the recent classification compiled by Põlme et al. [70]. Fungal species’ edibility classification was accomplished by assessing the commercial importance of the collected species [71,72].

2.4. Environmental Data Collection and Analysis

To relate the composition of edible fungal taxa to edaphic variables, soil samples were collected from each of the sample plots established in each forest. After clearing and removing plant matter and debris from the soil surface, five soil cores were extracted from the center and the four corners of each plot using an auger (2 cm radius, 20 cm depth, and 250 cm3). The soil cores collected from each plot were pooled, and a composite, relatively homogeneous subsample of approximately 500 g from each plot was placed in a plastic bag for analysis. The soil pH was determined by analyzing a soil:water (1:2.5) suspension with the aid of a pH meter [73]. The organic carbon (C) content of the soil was determined using wet digestion [74]. The Kjeldahl procedure was used to determine the total nitrogen (N) content of the soil samples [75]. Sodium bicarbonate (0.5 M NaHCO3) was used as an extraction solution to determine the available phosphorus (P) [76]. The soil analysis was conducted by Water Works Design and Supervision Enterprises, a laboratory service sub-process, the soil fertility section at Addis Ababa, and the Amhara Water Works Design and Supervision Works Enterprise at Bahir Dar, Ethiopia.
In addition to the soil samples, the following climate variables were obtained for each forest from nearby meteorological stations: daily, mean, minimum, and maximum temperatures (°C), total annual precipitation (mm), and the average temperature (°C) and precipitation (mm) values for July and August of 2019 (i.e., the sporocarp collection season).

2.5. Data Analysis

Data were transformed when necessary to achieve the parametric criteria of normality and homoscedasticity. Macrofungal data were normalized by rarefying the abundance data to the smallest number of macrofungi per plot. In addition, data from environmental variables were scaled using base R and used for subsequent statistical analyses. The sporocarp biomass (kg ha–1 yr–1) was estimated for each forest. Differences in sporocarp production levels across forests were assessed using linear mixed-effects models (LME) [77], where a block (a set of plots at the same site in each forest) was defined as random and the forest was defined as a fixed factor. LME analyses were used to prevent false positive associations due to the relatedness structure in the sampling. Tukey’s test was later used to check for significant differences (p ≤ 0.05) between forests when needed.
Relationships between sporocarp composition and environmental parameters were visualized using non-metric multidimensional scaling (NMDS) based on an absence and presence species data matrix and environmental scaled data. A permutation-based nonparametric MANOVA (PerMANOVA) [78] using the Euclidean distance was performed to analyze differences in sporocarp communities between forest types and across the five sites. Isolines were also plotted on the NMDS ordinations for rainfall using the ordisurf function. Correlations between NMDS axes scores with explanatory variables were assessed using the envfit function in R. To assess the influence of edaphic, climate, and location variables on the fungal community, we performed a Mantel test (Bray–Curtis distance) on the total species matrix and scaled environmental parameters.

3. Results

3.1. Edible Fungal Richness

In total, 64 edible fungal species belonging to 31 genera and 21 families were identified from the plots established in the natural and plantation forests. All these species belonged to the Basidiomycota, except for Morchella spp., which belonged to the Ascomycota. There was a significant difference in the total number of edible species found in each of the forest types (F = 7.23, p = 0.002). Forty edible species belonging to 22 genera and 15 families were found in the natural forests, while 16 species belonging to 16 genera and 12 families were found in plantation forests (Table 2). Eight edible species were common to both forest types.
Saprotrophs were the dominant guild (92.19%; n = 59 species) followed by ectomycorrhizal fungi (6.25%; n = 4 species), with other guilds comprising 1.56% of the species (Table 2). Ecologically and economically important edible fungal species such as Agaricus campestroides, Agaricus subedulis, Tylopilus niger, Suillus luteus, Tricholoma portentosum, Tricholoma saponaceum, Morchella americana, and Morchella anatolica were collected (Table 2).

3.2. Sporocarp Production

We found significant differences in the total edible sporocarp production between the two forest types (F = 4.293 p = 0.04; Figure 4A), with significantly greater mean sporocarp production levels in plantation forests (2097.57 kg ha−1 yr–1) than in natural forests (731.18 kg ha−1 yr–1).
The average sporocarp production levels of edible species differed significantly among the five studied sites (F = 9.24; p = 0.0001), with the highest mean production levels recorded in the Menagesha Suba forest (7.49 kg ha−1). This value was significantly higher than that of the Wondo Genet (p-MS _ p-WG = 0.004), Banja (p-MS _ p-Ba = 0.0001), Alemsaga (p-MS _ p-Al = 0.000), and Taragedam forests (p-MS _ p-TG = 0.001). Mean sporocarp production in Wondo Genet forests (3.29 kg ha−1) was also significantly higher than that in Alemsaga forests (p-WG _ p-Al = 0.02) but was not significantly different from that recorded for the Taragedam (p-WG _ p-TA = 0.775) and Banja forests (p-WG _ p-TG = 0.2626). Mean sporocarp production levels in Alemsaga, Banja, and Taragedam forests did not differ significantly (p > 0.05; Figure 3B).

3.3. Sporocarp Composition and Environmental Variables

The perMANOVA analyses indicated that the two forest types differed significantly in their sporocarp composition (F = 5.343, R2 = 0.14, p = 0.001; Figure 5A). Explanatory variables categorized as edaphic, climate, and location parameters were correlated with fungal community composition (p < 0.05; Table 3). Of these, a Mantel test confirmed that location variables had a significantly stronger aggregate effect on the sporocarp composition of both forest types (p = 0.000) than climate (p = 0.0001) or edaphic variables (p = 0.0001). The significance of each explanatory variable and their aggregated contribution to differences in sporocarp composition are shown in Table 3.
When sites were analyzed separately, environmental variables such as N, P, OC, Tmin, and latitude were significantly correlated (p < 0.05) with the composition of edible sporocarps in the study areas. Of these variables, the Mantel test confirmed that location variables had a significantly stronger aggregate effect on sporocarp composition (r = 0.6419; p = 0.0001) than climate (r = 0.4378; p = 0.0001) and edaphic variables (r = 0.3166; p = 0.0001) when analyzed by site (Figure 4B).

4. Discussion

In the most rural parts of Ethiopia, the local people are dependent on forest resources, either in the form of subsistence or as a cash income derived from NTFPs [7]. The collection of wild edible mushrooms by local people is a common practice, particularly in the southwestern parts of the country [11]. However, wild mushrooms are not considered important sources of food and medicine by rural communities in the northern part of the country [79]. This might be due to the continuing exodus of people from the countryside, which has meant that local communities are gradually losing their traditional knowledge, particularly about wild mushroom species. Furthermore, although a limited number of studies have reported the availability of wild mushrooms in Ethiopia and their importance as sources of food, medicine, and to some extent, income for local communities [10,11], information about the type of wild edible fungal species that are available, their potential production, and their status in different forest systems is scant [80]. This study is the first systematic survey focused on wild edible mushrooms, which was carried out in forests located in central and northern Ethiopia, where remnants of natural forests and plantations of exotic trees exist [81]. We collected a total of 64 edible fungal species from the study sites. The majority of the edible species (n = 40) were collected only from study sites in natural forests.
We collected wild edible species that have both economic and ecological significance belonging to the genera Morchella, Suillus, and Tylopilus in plantation forests and Tricholoma in natural forests [82]. In addition, we found some Agaricus species and Termitomyces, which are known to be used by rural people in the southwest part of the country [26,83], and a Schizophyllum species, which is eaten by local people in southern Ethiopia [10,24]. Of these species, Suillus luteus is consumed by local people and is also sold in markets at a good commercial price in different developing countries, along with other NTFPs [44]. Furthermore, these kinds of mushrooms could help to sustain communities during periods of food scarcity, serving as an important source of nutrients for local people [82]. The use of wild mushrooms by rural people as a food source during lean periods has been documented in Ethiopian ethnomycological literature. In most cases, these species are collected for subsistence use [84]. However, in some places, mushrooms can provide households with additional income when sold in the markets. For example, in local markets in the southern and southwestern parts of the country, Agaricus sp. and Termitomyces sp. are available occasionally in association with other vegetal products, which the local people sell to earn some extra money to supplement their household income [83]. Therefore, the conservation and development of these kinds of valuable species deserve special attention given their possible role in increasing food security and income generation to subsidize rural household economies. In addition, as mushroom collection from wild habitats is seasonal, maintaining some of these edible species by means of local small-scale cultivation practices or in private forest areas would be very remunerative. Therefore, a strategy is needed for the adoption and cultivation of important species from the wild, which will not only increase their utilization but also create new sources of income for rural people and contribute to food security.
Although we collected more edible species from plots in natural forests than in plantation forests, sporocarp production levels were higher in plots in plantation forests than in natural forests. The greater number of species but lower sporocarp production levels in natural forests is unsurprising given that almost all the species found were saprotrophic and they were mainly composed of singleton taxa, with a small number of frequent species, which is in agreement with the findings of previous studies [85]. The saprophytic fungal species collected were also characterized by low levels of biomass production, in accordance with Gassibe et al. and Mediavilla et al. [54,86]. Nonetheless, they are relevant for decomposition processes and ecosystem functioning [87], particularly in tropical forest systems such as the natural forests in this study, where decomposition is rapid [88]. This may reflect the accumulation of favorable substrates, which is likely to enhance the richness [89] of these systems. The conspicuous sporocarps produced by these saprophytic fungi may also have favored the collection of this particular fungal class, although basidiomycete mycelia are reported to be everywhere in forests [90]. Overall, the sporocarp yield obtained from plantation forests (2097.57 kg ha−1yr–1) was significantly higher than that from natural forests (731.18 kg ha−1yr–1). Interestingly, almost 25% of the species collected from plantation forests were marketable species and have economic significance, including Morchella sp., Suillus sp., and Tylopilus sp. [12]. These species were characterized by their high levels of biomass production. For example, in Peru and Mexico, Suillus and Morchella species are commercial NTFPs produced in plantation forests. They guarantee the economic performance of those forests [44,91] and the livelihoods of local communities [92], thus providing incentives for farmers to plant and manage more plantations in their surroundings. Furthermore, in Mexico, Morchella species are also exported to generate income [44]. Although the overall biomass produced by sites in this study was low, the most productive site (Menagesha Suba) had mean production levels per stand of 7.49 kg ha−1, which suggests the potential production levels of edible sporocarp species in forests with similar conditions in this area. This also provides a starting point in terms of broadening the management of forests for the production of NTFPs such as edible mushrooms in Ethiopia, depending on the location and type of forest.
Although natural forests produced lower sporocarp yields than plantation forests and had fewer marketable species, the overall yields and species were still valuably enhanced by plantations of exotic conifer species. A recent study [85] in the northern part of Ethiopia indicated that the overall land connectivity of natural forests with that of plantations provided important ectomycorrhizal species such as Tricholoma and Suillus in natural forest systems, indicating that such forest management activities could create important microniches with suitable resources and abiotic conditions to support more valuable mushroom species [93]. Thus, enrichment of natural forest systems through planting diverse tree species could potentially offer suitable habitats to enhance the richness and productivity of valuable edible species in natural forests in the study areas.
As we hypothesized, distinct fungal communities were observed in the two forest types. Fungal communities in natural forests were characterized by a large number of species, which may have been due to the greater spatial heterogeneity of the soil in natural forests compared with plantation forests. A heterogeneous soil environment and high rainfall levels create microhabitats in which saprotrophic species should be able to find the resources they require to survive in natural forests. The vegetation composition also impacts the composition of edible fungal species via the quantity and quality of the organic inputs, which mainly affect the saprotrophic community structure [94]. Most of the species found in the natural forests were associated with litter decomposition, which is typically characteristic of tropical forests [42]. However, some specific species such as Termitomyces and Tricholoma species were exclusively found in natural forests. The genus Termitomyces comprises a group of gilled mushrooms that have formed a termitophilic association with a particular family of termites, the Macrotermitinae (Isoptera), which are commonly found in Africa in places with a dry and humid climate [95]. This might be because our sampling sites in the natural forest areas were generally classified as dry Afromontane forest areas, characterized by high humidity and prolonged dry seasons [1]. These conditions might favor the occurrence and formation of distinct fungal community compositions in natural forest systems. Distinct fungal community compositions were also observed for plantation forests. Ectomycorrhizal fungi characterized the fungal composition of plantation forests along with some saprotrophic species. Tylopilus and Suillus species were site-exclusive species that were significantly more abundant in plantation forests comprised mainly Pinus than in other plantation forests. Some mushroom species, such as Agaricus campestroides, Coprinellus domesticus, Leucoagaricus holosericeus, Hygrophoropsis aurantiaca, and Leucocoprinus cepistipes, were common in both forest systems, indicating that these genera might be characterized as generalists.
Studies have shown that different environmental variables govern the composition of fungal species and that different fungal taxa are likely to respond to edaphic variables in different ways, depending on their characteristics [96,97] and, in turn, the composition of fungal communities is directly correlated with soil parameters [98]. In this study, organic matter, P, and N were significantly correlated with the whole edible fungal species community dataset. This is likely to be because organic matter influences the fungal community through its impact on the water-holding capacity of the soil and on nutrient availability [99]. Thus, organic matter may favor more fungal assembly in an area, particularly saprotrophic fungi. Furthermore, the finding that N was an important factor correlated with fungal taxa compositions is in accordance with previous studies [28,35,100] that noted the influence of N on fungal distribution patterns. These studies reported that fungi showed community specialization toward more N-rich soil sites. This might be because nitrogen can influence the formation of extraradical mycelium in the soil and play a vital role in sporocarp formation [101]. Other studies have also noted that fungal communities adapt to more nitrogen-rich sites [100,102]. Furthermore, the microclimate directly influences ecological processes and reflects subtle changes in ecosystem functioning, particularly in forests where the majority of the identified edible species are saprotrophic and depend on a suitable microclimate for their growth and production. In that context, a mosaic forest management scheme is needed that considers both timber production and edible wild mushroom production. Such as scheme must uphold the environment variables needed to create suitable habitats, with variable microclimates to promote diverse sporocarps. This could increase the value of Ethiopia’s remnant natural forests and provide incentives for forest owners to sustainably manage and conserve the forests’ resources in different forms.

5. Conclusions

Although our data were based on only one year of sampling during the rainy season in July and August 2019, we conclude that the richness of edible mushrooms is relatively higher in natural forests. The majority of these species are saprophytic and characterized by the production of small sporocarps. Although only a small number of edible species were recorded in plantation forests, sporocarp production levels in these forests were high. We also observed that some species specifically characterized each forest, and there was also a noticeable presence of valuable species in both forest types that could be potentially marketed in rural areas, providing forest managers and local people with supplementary incomes. Species such as Termitomyces and Tricholoma were exclusively found in natural forests, while Suillus luteus was found in plantation forests. However, environmental variables such as climate, spatial parameters, and edaphic parameters were found to affect the composition and, thus, sporocarp production in the study areas. In this regard, we found that the relative contributions of the spatial and climatic parameters were greater than those of the edaphic-specific variables, which significantly affected sporocarp production and species composition. However, a mosaic landscape, mixing natural and plantation forests, could provide timber and high levels of edible mushroom production. Accordingly, this could contribute to the conservation of remnant natural forests, which have high biodiversity values. Thus, a management scheme is needed that combines timber and mushroom production, which could provide economic and ecological benefits, especially in natural forest systems, which are characterized by excessive deforestation and forest degradation. Furthermore, the scheme should consider the influence of different environmental conditions, to create habitats that offer a suitable environment with variable microclimates for the promotion of diverse edible fungi and higher levels of sporocarp production, particularly in natural forests where the majority of the identified edible species were characterized by low levels of sporocarp production.

Author Contributions

Conceptualization, P.M.-P.; methodology, T.D., G.K. and P.M.-P.; formal analysis, P.M.-P., T.D. and G.K.; investigation, G.K., D.A. and T.D.; data curation, G.K. and D.A.; writing—original draft preparation, G.K.; writing—review and editing, P.M.-P., T.D. and M.T.; supervision, P.M.-P., T.D. and M.T.; project administration, P.M.-P.; funding acquisition, P.M.-P. All authors have read and agreed to the published version of the manuscript.

Funding

This work was supported by the projects SUSTIFUNGI_ET (Sustfungi_Eth: 2017/ACDE/002094) and MYCOPROED_ET (Mycoproed_Eth: 2019/ACDE/000921) and SUSTFUNGI_ET_III (Sustfungi_Eth_III: 2022/ACDE/000201), which were funded by the Spanish Agency for International Development and Cooperation. Gonfa Kewessa also received funds through the UVa 2021 call for predoctoral contracts, co-financed by Banco Santander.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

Not applicable.

Acknowledgments

We would like to express our gratitude to the people involved in the fieldwork.

Conflicts of Interest

The authors declare no conflict of interest.

References

  1. Friis, B.; Demissew, S.; Breugel, P. Atlas of the Potential Vegetation of Ethiopia; The Royal Danish Academy of Sciences and Letters: Copenhagen, Denmark, 2010. [Google Scholar]
  2. Badege, B. Deforestation and Land Degradation in the Ethiopian Highlands: A Strategy for Physical Recovery. Northeast Afr. Stud. 2001, 8, 7–26. [Google Scholar]
  3. Bekele, M. Forest Plantations and Woodlots in Ethiopia. Afr. For. forum 2011, 1, 52. [Google Scholar]
  4. Adilo, M. The Contribution of Non-Timber Forest Products to Rural Livelihood in Southwest Ethiopia. Master’s Thesis, Wageningen University, Wageningen, The Netherlands, 2007. [Google Scholar]
  5. Seyoum, A. Economic value of Afromontane Forests of Sheka: Forest of Sheka. In Forest of Sheka; Fetene, M., Ed.; MEHBER: Addis Ababa, Ethiopia, 2007; pp. 183–218. [Google Scholar]
  6. Sultan, M. The Role of Non-Timber Forest Products to Rural Livelihoods and Forest Conservation: A Case Study at Harana Bulluk District Oromia National Regional State, Ethiopia. Master’s Thesis, Wondo Genet College of Forestry and Natural Resource, Hawassa University, Wondo Genet, Ethiopia, 2009. [Google Scholar]
  7. Lulekal, E.; Asfaw, Z.; Kelbessa, E.; Van Damme, P. Wild edible plants in Ethiopia: A review on their potential to combat food insecurity. Afrika Focus 2011, 24, 71–122. [Google Scholar] [CrossRef]
  8. Melaku, E.; Ewnetu, Z.; Teketay, D. Non-timber forest products and household incomes in Bonga forest area, southwestern Ethiopia. J. For. Res. 2014, 25, 215–223. [Google Scholar] [CrossRef]
  9. Asfaw, Z.; Tadesse, M. Prospects for sustainable use and development of wild food plants in Ethiopia. Econ. Bot. 2001, 55, 47–62. [Google Scholar] [CrossRef]
  10. Sitotaw, R.; Lulekal, E.; Abate, D. Ethnomycological study of edible and medicinal mushrooms in Menge District, Asossa Zone, Benshangul Gumuz Region, Ethiopia. J. Ethnobiol. Ethnomed. 2020, 16, 11. [Google Scholar] [CrossRef]
  11. Dejene, T.; Oria-de-Rueda, J.A.; Martín-Pinto, P. Edible wild mushrooms of Ethiopia: Neglected non-timber forest products. Rev. Fitotec. Mex. 2017, 40, 391–397. [Google Scholar] [CrossRef]
  12. Boa, E. Wild Edible Fungi: A Global Overview of Their Use and Importance to People; FAO: Rome, Italy, 2004; ISBN 9251051577. [Google Scholar]
  13. Barroetaveña, C.; La Manna, L.; Alonso, M.V. Variables affecting Suillus luteus fructification in ponderosa pine plantations of Patagonia (Argentina). For. Ecol. Manag. 2008, 256, 1868–1874. [Google Scholar] [CrossRef]
  14. De Román, M.; Boa, E. International Seminar on Ethnomycology Collection, Marketing and Cultivation of Edible Fungi in Spain. Micol. Apl. Int. 2004, 16, 25–33. [Google Scholar]
  15. Chang, Y.S.; Lee, S.S. Utilisation of macrofungi species in Malaysia. Fungal Divers 2004, 15, 15–22. [Google Scholar]
  16. Mau, J.; Chang, C.; Huang, S.; Chen, C. Antioxidant properties of methanolic extracts from Grifola frondosa, Morchella esculenta and Termitomyces albuminosus mycelia. Food Chem. 2004, 87, 111–118. [Google Scholar] [CrossRef]
  17. Chen, Y.; Yuan, Z.; Bi, S.; Wang, X.; Ye, Y.; Svenning, J.-C. Macrofungal species distributions depend on habitat partitioning of topography, light, and vegetation in a temperate mountain forest. Sci. Rep. 2018, 8, 13589. [Google Scholar] [CrossRef]
  18. Brundrett, M.C. Mycorrhizal associations and other means of nutrition of vascular plants: Understanding the global diversity of host plants by resolving conflicting information and developing reliable means of diagnosis. Plant Soil 2009, 320, 37–77. [Google Scholar] [CrossRef]
  19. Courty, P.-E.; Buée, M.; Diedhiou, A.G.; Frey-Klett, P.; Le Tacon, F.; Rineau, F.; Turpault, M.-P.; Uroz, S.; Garbaye, J. The role of ectomycorrhizal communities in forest ecosystem processes: New perspectives and emerging concepts. Soil Biol. Biochem. 2010, 42, 679–698. [Google Scholar] [CrossRef]
  20. Fernández-Toirán, L.; Agreda, T.; Olano, J. Stand age and sampling year effect on the fungal fruit body community in Pinus pinaster forests in central Spain. Can. J. Bot. 2006, 84, 1249–1258. [Google Scholar] [CrossRef]
  21. Martín-Pinto, P.; Vaquerizo, H.; Peñalver, F.; Olaizola, J.; Oria-De-Rueda, J.A. Early effects of a wildfire on the diversity and production of fungal communities in Mediterranean vegetation types dominated by Cistus ladanifer and Pinus pinaster in Spain. For. Ecol. Manag. 2006, 225, 296–305. [Google Scholar] [CrossRef]
  22. Yanagihara, H.; Kamo, K.; Imori, S.; Satoh, K. Bias-corrected AIC for selecting variables in multinomial logistic regression models. Linear Algebra Appl. 2012, 436, 4329–4341. [Google Scholar] [CrossRef]
  23. Ferris, R.; Peace, A.J.; Newton, A.C. Macrofungal communities of lowland Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karsten.) plantations in England: Relationships with site factors and stand structure. For. Ecol. Manag. 2000, 131, 255–267. [Google Scholar] [CrossRef]
  24. Tuno, N. Mushroom utilization by the Majangir, an Ethiopian tribe. Mycologist 2001, 15, 78–79. [Google Scholar] [CrossRef]
  25. Alemu, F. Assessment of wild mushrooms and wood decaying fungi in Dilla University, main campus, Ethiopia. Int. J. Adv. Res. 2013, 1, 458–467. [Google Scholar]
  26. Muleta, D.; Woyessa, D.; Teferi, Y. Mushroom consumption habits of Wacha Kebele residents, southwestern Ethiopia. Glob. Res. J. Agric. Biol. Sci. 2013, 4, 6–16. [Google Scholar]
  27. Kassa, H.; Campbell, B.; Sandewall, M.; Kebede, M.; Tesfaye, Y.; Dessie, G.; Seifu, A.; Tadesse, M.; Garedew, E.; Sandewall, K. Building future scenarios and uncovering persisting challenges of participatory forest management in Chilimo Forest, Central Ethiopia. J. Environ. Manag. 2009, 90, 1004–1013. [Google Scholar] [CrossRef] [PubMed]
  28. Gassibe, P.V.; Oria-de-Rueda, J.A.; Martín-Pinto, P.P. pinaster under extreme ecological conditions provides high fungal production and diversity. For. Ecol. Manag. 2015, 337, 161–173. [Google Scholar] [CrossRef]
  29. Bonet, J.A.; Fischer, C.R.; Colinas, C. The relationship between forest age and aspect on the production of sporocarps of ectomycorrhizal fungi in Pinus sylvestris forests of the central Pyrenees. For. Ecol. Manag. 2004, 203, 157–175. [Google Scholar] [CrossRef]
  30. Smith, J.E.; Molina, R.; Huso, M.M.; Luoma, D.L.; McKay, D.; Castellano, M.A.; Lebel, T.; Valachovic, Y. Species richness, abundance, and composition of hypogeous and epigeous ectomycorrhizal fungal sporocarps in young, rotation-age, and old-growth stands of Douglas-fir (Pseudotsuga menziesii) in the Cascade Range of Oregon, U.S.A. Can. J. Bot. 2002, 80, 186–204. [Google Scholar] [CrossRef]
  31. Bonet, J.A.; Palahí, M.; Colinas, C.; Pukkala, T.; Fischer, C.R.; Miina, J.; Martínez de Aragón, J. Modelling the production and species richness of wild mushrooms in pine forests of the Central Pyrenees in northeastern Spain. Can. J. For. Res. 2010, 40, 347–356. [Google Scholar] [CrossRef]
  32. Pinna, S.; Gévry, M.F.; Côté, M.; Sirois, L. Factors influencing fructification phenology of edible mushrooms in a boreal mixed forest of Eastern Canada. For. Ecol. Manag. 2010, 260, 294–301. [Google Scholar] [CrossRef]
  33. Santos-Silva, C.; Gonçalves, A.; Louro, R. Canopy cover influence on macrofungal richness and sporocarp production in montado ecosystems. Agrofor. Syst. 2011, 82, 149–159. [Google Scholar] [CrossRef]
  34. Straatsma, G.; Ayer, F.; Egli, S. Species richness, abundance, and phenology of fungal fruit bodies over 21 years in a Swiss forest plot. Mycol. Res. 2001, 105, 515–523. [Google Scholar] [CrossRef]
  35. Reverchon, F.; Del Ortega-Larrocea, P.M.; Pérez-Moreno, J. Saprophytic fungal communities change in diversity and species composition across a volcanic soil chronosequence at Sierra del Chichinautzin, Mexico. Ann. Microbiol. 2010, 60, 217–226. [Google Scholar] [CrossRef]
  36. Jaleta, D.; Mbilinyi, B.; Mahoo, H.; Lemenih, M. Eucalyptus expansion as relieving and provocative tree in Ethiopia. J. Agric. Ecol. Res. Int. 2016, 6, 1–12. [Google Scholar] [CrossRef]
  37. Teketay, D. The ecological effects of Eucalyptus: Ground for making wise and informed decision. In Proceedings of the Eucalyptus Dilemma, Addis Ababa, Ethiopia, 15 November 2000. [Google Scholar]
  38. Castellano, M.; Molina, R. Mycorrhizae. In The Biological Component: Nursery Pests and Mycorrhizal; Landis, T.D., Tinus, R.W., Mc Donald, S.E., Barnett, J.P., Eds.; The Container Tree Nursery Manual. Agriculture Handbook 674, Forest Service; USDA: Washington, DC, USA, 1989; Volume 5, 171p. [Google Scholar]
  39. Trappe, J. Selection of fungi for ectomycorrhizal inoculation in nurseries. Annu. Rev. Phytopatho. 1977, 15, 203–222. [Google Scholar] [CrossRef]
  40. Glassman, S.I.; Wang, I.J.; Bruns, T.D. Environmental filtering by pH and soil nutrients drives community assembly in fungi at fine spatial scales. Mol. Ecol. 2017, 26, 6960–6973. [Google Scholar] [CrossRef] [PubMed]
  41. Li, P.; Li, W.; Dumbrell, A.J.; Liu, M.; Li, G.; Wu, M.; Jiang, C.; Li, Z. Spatial Variation in Soil Fungal Communities across Paddy Fields in Subtropical China. mSystems 2020, 5, e00704-19. [Google Scholar] [CrossRef] [PubMed]
  42. Tedersoo, L.; Bahram, M.; Polme, S.; Koljalg, U.; Yorou, N.S.; Wijesundera, R.; Ruiz, L.V.; Vasco-Palacios, A.M.; Thu, P.Q.; Suija, A.; et al. Global diversity and geography of soil fungi. Science 2014, 346, 1256688. [Google Scholar] [CrossRef]
  43. Kranabetter, J.M.; Friesen, J.; Gamiet, S.; Kroeger, P. Ectomycorrhizal mushroom distribution by stand age in western hemlock—Lodgepole pine forests of northwestern British Columbia. Can. J. For. Res. 2005, 35, 1527–1539. [Google Scholar] [CrossRef]
  44. Pérez-Moreno, J.; Martínez-Reyes, M.; Yescas-Pérez, A.; Delgado-Alvarado, A.; Xoconostle-Cázares, B. Wild mushroom markets in central Mexico and a case study at Ozumba. Econ. Bot. 2008, 62, 425–436. [Google Scholar] [CrossRef]
  45. Collado, E.; Bonet, J.A.; Camarero, J.J.; Egli, S.; Peter, M.; Salo, K.; Martínez-Peña, F.; Ohenoja, E.; Martín-Pinto, P.; Primicia, I.; et al. Mushroom productivity trends in relation to tree growth and climate across different European forest biomes. Sci. Total Environ. 2019, 689, 602–615. [Google Scholar] [CrossRef] [PubMed]
  46. Martínez de Aragón, J.; Bonet, J.A.; Fischer, C.R.; Colinas, C. Productivity of ectomycorrhizal and selected edible saprotrophic fungi in pine forests of the pre-Pyrenees mountains, Spain: Predictive equations for forest management of mycological resources. For. Ecol. Manag. 2007, 252, 239–256. [Google Scholar] [CrossRef]
  47. Newsham, K.K.; Hopkins, D.W.; Carvalhais, L.C.; Fretwell, P.T.; Rushton, S.P.; O’Donnell, A.G.; Dennis, P.G. Relationship between soil fungal diversity and temperature in the maritime Antarctic. Nat. Clim. Chang. 2016, 6, 182–186. [Google Scholar] [CrossRef]
  48. Worku, M. The Role of Forest Biodiversity Conservation Practices for Tourism Development in a Case of Tara Gedam Monastery, South Gonder Zone, Ethiopia. J. Ecosyst. Ecography 2017, 7, 2–7. [Google Scholar] [CrossRef]
  49. Esubalew, E.; Giday, K.; Hishe, H.; Goshu, G. Carbon stock of woody species along Altitude gradient in Alemsaga Forest, South Gondar, North Western Ethiopia. Agric. J. IJOEAR 2019, 5, 13–21. [Google Scholar]
  50. Abere, F.; Belete, Y.; Kefalew, A.; Soromessa, T. Carbon stock of Banja forest in Banja district, Amhara region, Ethiopia: An implication for climate change mitigation. J. Sustain. For. 2017, 36, 604–622. [Google Scholar] [CrossRef]
  51. Alem, D.; Dejene, T.; Oria-de-Rueda, J.A.; Geml, J.; Castaño, C.; Smith, J.E.; Martín-Pinto, P. Soil fungal communities and succession following wildfire in Ethiopian dry Afromontane forests, a highly diverse underexplored ecosystem. For. Ecol. Manag. 2020, 474, 118328. [Google Scholar] [CrossRef]
  52. Jemal, A.; Getu, E. Diversity of butterfly communities at different altitudes of Menagesha-suba state forest, Ethiopia. J. Entomol. Zool. Stud. 2018, 6, 2197–2202. [Google Scholar]
  53. Dejene, T.; Oria-de-Rueda, J.A.; Martín-Pinto, P. Fungal diversity and succession under Eucalyptus grandis plantations in Ethiopia. For. Ecol. Manag. 2017, 405, 179–187. [Google Scholar] [CrossRef]
  54. Gassibe, P.V.; Fabero, R.F.; Hernández-Rodríguez, M.; Oria-de-Rueda, J.A.; Martín-Pinto, P. Fungal community succession following wildfire in a Mediterranean vegetation type dominated by Pinus pinaster in Northwest Spain. For. Ecol. Manag. 2011, 262, 655–662. [Google Scholar] [CrossRef]
  55. Hernández-Rodríguez, M.; Oria-de-Rueda, J.A.; Martín-Pinto, P. Post-fire fungal succession in a Mediterranean ecosystem dominated by Cistus ladanifer L. For. Ecol. Manag. 2013, 289, 48–57. [Google Scholar] [CrossRef]
  56. Hiiesalu, I.; Bahram, M.; Tedersoo, L. Plant species richness and productivity determine the diversity of soil fungal guilds in temperate coniferous forest and bog habitats. Mol. Ecol. 2017, 26, 4846–4858. [Google Scholar] [CrossRef]
  57. Rudolph, S.; Maciá-Vicente, J.G.; Lotz-Winter, H.; Schleuning, M.; Piepenbring, M. Temporal variation of fungal diversity in a mosaic landscape in Germany. Stud. Mycol. 2018, 89, 95–104. [Google Scholar] [CrossRef]
  58. O’Hanlon, R.; Harrington, T.J. Macrofungal diversity and ecology in four Irish forest types. Fungal Ecol. 2012, 5, 499–508. [Google Scholar] [CrossRef]
  59. Ruiz-Almenara, C.; Gándara, E.; Gómez-Hernández, M. Comparison of diversity and composition of macrofungal species between intensive mushroom harvesting and non-harvesting areas in Oaxaca, Mexico. PeerJ 2019, 7, e8325. [Google Scholar] [CrossRef] [PubMed]
  60. Adeniyi, M.; Adeyemi, Y.; Odeyemi, Y.; Odeyemi, O. Ecology, diversity and seasonal distribution of wild mushrooms in a Nigerian tropical forest reserve. Biodiversitas J. Biol. Divers 2018, 19, 285–295. [Google Scholar] [CrossRef]
  61. Antonin, V. Monograph of Marasmius, Gloiocephala, Palaeocephala and Setulipes in Tropical Africa. Fungus Fl Trop Afr. 2007, 1, 177. [Google Scholar]
  62. Hama, O.; Maes, E.; Guissou, M.; Ibrahim, D.; Barrage, M.; Parra, L.; Raspe, O.; De Kesel, A. Agaricus subsaharianus, une nouvelle espèce comestible et consomméeau Niger, au Burkina Faso et en Tanzanie. Crypto. Mycol. 2010, 31, 221–234. [Google Scholar]
  63. Pegler, D.N. Studies on African Agaricales: II *. Kew Bull. 1969, 23, 219–249. [Google Scholar] [CrossRef]
  64. Morris, B. An annotated check-list of the macrofungi of Malawi. Kirkia 1990, 13, 323–364. [Google Scholar]
  65. Pegler, D.N. Studies on African Agaricales: 1. Kew Bull. 1968, 21, 499–533. [Google Scholar] [CrossRef]
  66. Pegler, D.; Rayner, R. A contribution to the Agaric flora of Kenya. Kew Bull. 1969, 23, 347–412. [Google Scholar] [CrossRef]
  67. Rammeloo, J.; Walleyn, R. The edible fungi of Africa South of the Sahara: A literature survey. Scr. Bot. Belg. 1993, 5, 62. [Google Scholar]
  68. Ryvarden, L.; Piearce, G.D.; Masuka, A.J. An Introduction to the Larger Fungi of South Central Africa; Baobab Books: Harare, Zimbabwe, 1994. [Google Scholar]
  69. Singer, R. Marasmius. Fl. Icon. Champ. Congo 1965, 14, 253–278. [Google Scholar]
  70. Põlme, S.; Abarenkov, K.; Nilsson, R.H.; Lindahl, B.D.; Clemmensen, K.E.; Kauserud, H.; Nguyen, N.; Kjøller, R.; Bates, S.T.; Baldrian, P.; et al. FungalTraits: A user-friendly traits database of fungi and fungus-like stramenopiles. Fungal Divers 2020, 105, 1–16. [Google Scholar] [CrossRef]
  71. Gerhardt, E.; Vila, J.; Llimona, X. Mushroom from Spain and Europe: Manual for Identification; Omega SA Editions: Barcelona, Spain, 2000; ISBN 84-282-1120-5. [Google Scholar]
  72. Moreno, G.; Manjón, J.L. Guide to Fungi of the Iberian Peninsula; Omega SA Editions: Barcelona, Spain, 2010. [Google Scholar]
  73. Reeuwijk, L. Procedures for Soil Analysis, 6th ed.; International Soil Reference and Information Centre: Wageningen, The Netherlands, 2002. [Google Scholar]
  74. Walkley, A.; Black, I.A. An examination of the digestion method for determining soil organic matter and a proposed modification of the chromic acid titration method. Soil Sci. 1934, 34, 29–38. [Google Scholar] [CrossRef]
  75. Kim, J.; Kreller, C.R.; Greenberg, M.M. Preparation and Analysis of Oligonucleotides Containing the C4‘-Oxidized Abasic Site and Related Mechanistic Probes. J. Org. Chem. 2005, 70, 8122–8129. [Google Scholar] [CrossRef]
  76. Kim, H.T. Soil Sampling, Preparation and Analysis; CRC Press: Boca Raton, FL, USA, 1996; pp. 139–145. [Google Scholar]
  77. Pinheiro, J.; Bates, D.; DebRoy, S.; Sarkar, D.; Team, R.C. Nlme: Linear and Nonlinear Mixed Effects Models. R Packag. Version 3.1–126. 2016. Available online: http://CRAN>R-project.org/package=nlme (accessed on 1 August 2022).
  78. Anderson, M.J. A new method for non parametric multivariate analysis of variance. Austral Ecol. 2001, 26, 32–46. [Google Scholar] [CrossRef]
  79. Zeleke, G.; Dejene, T.; Tadesse, W.; Agúndez, D.; Martín-Pinto, P. Ethnomycological Knowledge of Three Ethnic Groups in Ethiopia. Forests 2020, 11, 875. [Google Scholar] [CrossRef]
  80. Sitotaw, R.; Mulat, A.; Abate, D. Morphological and molecular studies on Termitomyces species of Menge District, Asossa Zone, Northwest Ethiopia. Sci. Technol. Arts Res. J. 2015, 4, 49–57. [Google Scholar] [CrossRef]
  81. Faye, M.D.; Weber, J.C.; Abasse, T.A.; Boureima, M.; Larwanou, M.; Bationo, A.B.; Diallo, B.O.; Sigué, H.; Dakouo, J.M.; Samaké, O.; et al. Farmers’ preferences for tree functions and species in the west African sahel. For. Trees Livelihoods 2011, 20, 113–136. [Google Scholar] [CrossRef]
  82. Sundriyal, M.; Sundriyal, R. Wild edible plants of the Sikkim Himalaya: Nutritive values of selected species. Econ. Bot. 2001, 55, 377–390. [Google Scholar] [CrossRef]
  83. Abate, D. Wild mushrooms and mushroom cultivation efforts in Ethiopia. WSMBMP Bull. 2014, 11, 1–3. [Google Scholar]
  84. Yehuala, K. Contribution of Small Scale Mushroom Production for Food Security in the Amhara. In Proceedings of the Experience Sharing and Revitalization Workshop of Amhara Region Food Security Network, Bahir Dar, Ethiopia, 26–27 August 2010. [Google Scholar]
  85. Alem, D.; Dejene, T.; Oria-de-Rueda, J.A.; Martín-Pinto, P. Survey of macrofungal diversity and analysis of edaphic factors influencing the fungal community of church forests in Dry Afromontane areas of Northern Ethiopia. For. Ecol. Manag. 2021, 496, 119391. [Google Scholar] [CrossRef]
  86. Mediavilla, O.; Oria-de-Rueda, J.A.; Martín-Pinto, P. Changes in sporocarp production and vegetation following wildfire in a Mediterranean Forest Ecosystem dominated by Pinus nigra in Northern Spain. For. Ecol. Manag. 2014, 331, 85–92. [Google Scholar] [CrossRef]
  87. Deacon, J. Fungal Biology, 4th ed.; Blackwell Publishing Oxford Press: Malden, MA, USA, 2006. [Google Scholar]
  88. Powers, J.S.; Montgomery, R.A.; Adair, E.C.; Brearley, F.Q.; DeWalt, S.J.; Castanho, C.T.; Chave, J.; Deinert, E.; Ganzhorn, J.U.; Gilbert, M.E.; et al. Decomposition in tropical forests: A pan-tropical study of the effects of litter type, litter placement and mesofaunal exclusion across a precipitation gradient. J. Ecol. 2009, 97, 801–811. [Google Scholar] [CrossRef]
  89. Wardle, D.A.; Bardgett, R.; Klironomos, J.; Setala, H.; van der Putten, W.; Wall, D. Ecological Linkages Between Aboveground and Belowground Biota. Science 2004, 304, 1629–1633. [Google Scholar] [CrossRef]
  90. Cairney, J.W.G.; Bastias, B.A. Influences of fire on forest soil fungal communitiesThis article is one of a selection of papers published in the Special Forum on Towards Sustainable Forestry—The Living Soil: Soil Biodiversity and Ecosystem Function. Can. J. For. Res. 2007, 37, 207–215. [Google Scholar] [CrossRef]
  91. Toivanen, T.; Markkanen, A.; Kotiaho, J.S.; Halme, P. The effect of forest fuel harvesting on the fungal diversity of clear-cuts. Biomass Bioenergy 2012, 39, 84–93. [Google Scholar] [CrossRef]
  92. Elizabeth, M.-E.; Felipe, R.-S.; Maribel, I.-M. Conocimiento Popular Acerca De La K’Allampa De Pino (Suillus luteus (L) Roussel) En La Localidad De Alalay, Mizque (Cochabamba, Bolivia): Un Ejemplode Dilogo De Saberes. Revista Etnobiologia 2018, 16, 76–86. [Google Scholar]
  93. Gómez-Hernández, M.; Williams-Linera, G. Diversity of macromycetes determined by tree species, vegetation structure, and microenvironment in tropical cloud forests in Veracruz, Mexico. Botany 2011, 89, 203–216. [Google Scholar] [CrossRef]
  94. Read, D.; Perez-Moreno, J. Mycorrhizas and nutrient cycling in ecosystems—A journey towards relevance? New Phytol. 2003, 157, 475–492. [Google Scholar] [CrossRef]
  95. Kirk, P.; Cannon, P.; Minter, D.; Stalpers, J. Dictionary of the Fungi, 10th ed.; The Centre for Agriculture and Bioscience International (CABI): Wallingford, UK, 2008. [Google Scholar]
  96. Crowther, T.W.; Stanton, D.W.G.; Thomas, S.M.; A’Bear, A.D.; Hiscox, J.; Jones, T.H.; Voříšková, J.; Baldrian, P.; Boddy, L. Top-down control of soil fungal community composition by a globally distributed keystone consumer. Ecology 2013, 94, 2518–2528. [Google Scholar] [CrossRef]
  97. Koide, R.T.; Fernandez, C.; Malcolm, G. Determining place and process: Functional traits of ectomycorrhizal fungi that affect both community structure and ecosystem function. New Phytol. 2014, 201, 433–439. [Google Scholar] [CrossRef] [PubMed]
  98. Cozzolino, V.; Di Meo, V.; Monda, H.; Spaccini, R.; Piccolo, A. The molecular characteristics of compost affect plant growth, arbuscular mycorrhizal fungi, and soil microbial community composition. Biol. Fertil. Soils 2016, 52, 15–29. [Google Scholar] [CrossRef]
  99. Harrington, T.J. Relationships between macrofungi and vegetation in the burren. Biol. Environ. 2003, 103, 147–159. [Google Scholar] [CrossRef]
  100. Kranabetter, J.M.; Durall, D.M.; MacKenzie, W.H. Diversity and species distribution of ectomycorrhizal fungi along productivity gradients of a southern boreal forest. Mycorrhiza 2009, 19, 99–111. [Google Scholar] [CrossRef] [PubMed]
  101. Trudell, S.A.; Edmonds, R.L. Macrofungus communities correlate with moisture and nitrogen abundance in two old-growth conifer forests, Olympic National Park, Washington, USA. Can. J. Bot. 2004, 82, 781–800. [Google Scholar] [CrossRef]
  102. Toljander, J.F.; Eberhardt, U.; Toljander, Y.K.; Paul, L.R.; Taylor, A.F.S. Species composition of an ectomycorrhizal fungal community along a local nutrient gradient in a boreal forest. New Phytol. 2006, 170, 873–884. [Google Scholar] [CrossRef] [PubMed]
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Figure 1. Map showing the locations of forests in which the study plots were located.
Figure 1. Map showing the locations of forests in which the study plots were located.
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Figure 2. Plantation of Pinus patula where sporocarps collection was carried out in Wondo Genet (photo credit: Tatek Dejene).
Figure 2. Plantation of Pinus patula where sporocarps collection was carried out in Wondo Genet (photo credit: Tatek Dejene).
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Figure 3. Dry Afromontane forests in (A) Wondo Genet, (B) Taragedam, (C) Alemsaga, and (D) Banja where sporocarps collections were conducted (photo credits: 3A, Tatek Dejene; 3B–3D, Demelash Alem).
Figure 3. Dry Afromontane forests in (A) Wondo Genet, (B) Taragedam, (C) Alemsaga, and (D) Banja where sporocarps collections were conducted (photo credits: 3A, Tatek Dejene; 3B–3D, Demelash Alem).
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Figure 4. Production of sporocarps at five study sites in Ethiopia according to the forest type (A) and site (B). Data are presented as boxplots showing the maximum and minimum values. The bar in the box is the standard deviation of the mean. Values with the same letter are not significantly different. Msuba, Menagesha Suba; WG, Wondo Genet.
Figure 4. Production of sporocarps at five study sites in Ethiopia according to the forest type (A) and site (B). Data are presented as boxplots showing the maximum and minimum values. The bar in the box is the standard deviation of the mean. Values with the same letter are not significantly different. Msuba, Menagesha Suba; WG, Wondo Genet.
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Figure 5. Non − metric multidimensional scaling (NMDS) ordination graph with fitted explanatory variables based on dissimilarities calculated using the Euclidean distance of sporocarp composition from plots in the two forest types (A) and by study sites (B), with rainfall displayed as isolines (fitted LOESS model R2 = 0.86). Arrows represent the environmental variables that were most significantly (p < 0.005) related to ordination. The five ellipses indicate the plots in the five study areas (Msuba, Menagesha Suba; WG, Wondo Genet). Explanatory variables are shown in blue: OM, organic matter; Tmin, minimum daily temperature; N, total nitrogen; P, available phosphorus; OC, organic carbon.
Figure 5. Non − metric multidimensional scaling (NMDS) ordination graph with fitted explanatory variables based on dissimilarities calculated using the Euclidean distance of sporocarp composition from plots in the two forest types (A) and by study sites (B), with rainfall displayed as isolines (fitted LOESS model R2 = 0.86). Arrows represent the environmental variables that were most significantly (p < 0.005) related to ordination. The five ellipses indicate the plots in the five study areas (Msuba, Menagesha Suba; WG, Wondo Genet). Explanatory variables are shown in blue: OM, organic matter; Tmin, minimum daily temperature; N, total nitrogen; P, available phosphorus; OC, organic carbon.
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Table
Table 1. Comprehensive descriptions of the forests and the study sites.
Table 1. Comprehensive descriptions of the forests and the study sites.
ForestGeographical CoordinatesAltitude
(m asl)		MAP
(mm)		MAT
(°C)		Vegetation TypesReference
Taragedam12°06′59″–12°07′25″ N and 37°46′14″–37°47′02″ E2062–2457130020.4Natural forests[48]
Alemsaga11°54′30″–11°56′00″ N and
37°55′00″–37°57′00″ E		2100–2470148416.4Natural forests[49]
Banja10°57′17″–11°03′05″ N and
36°39′09″–36°48′25″ E		1870–25701215.317.7Natural forests[50]
Wondo Genet7°06′–7°07′ N and 38°37′–38°42′ E1600–2580121020Natural forests and Pinus patula and Eucalyptus grandis plantation forests[51]
Menagesha Suba8°56′–9°03′ N and 38°28′–38°36′ E2200–3385110016Pinus radiata plantation forests[52]
Note: MAP, mean annual precipitation; MAT, mean annual temperature.
Table
Table 2. List of wild edible fungal species collected from study sites in natural and plantation forests, Ethiopia.
Table 2. List of wild edible fungal species collected from study sites in natural and plantation forests, Ethiopia.
SpeciesFamilyTNP
Agaricus augustus Fr.AgaricaceaeSx
Agaricus bitorquis (Quél.) Sacc.AgaricaceaeSx
Agaricus campestris L.AgaricaceaeSx
Agaricus campestroides Heinem. and Gooss. -Font.AgaricaceaeSxx
Agaricus moelleri WasserAgaricaceaeSx
Agaricus murinaceus Bull.AgaricaceaeSx
Agaricus subedulis Heinem.AgaricaceaeSxx
Ampulloclitocybe clavipes (Pers.) Redhead, Lutzoni, Moncalvo and VilgalysHygrophoraceaeSx
Armillaria heimii PeglerPhysalacriaceaePx
Auricularia auricula-judae (Bull.) Quél.AuriculariaceaeSx
Calvatia cyathiformis (Bosc) MorganLycoperdaceaeSx
Calvatia gigantea (Batsch) LloydLycoperdaceaeSx
Calvatia subtomentosa Dissing and M. LangeLycoperdaceaeS x
Clitocybe carolinensis H.E. Bigelow and HeslerTricholomataceaeSx
Clitocybe cistophila Bon and ContuTricholomataceaeSx
Clitocybe foetens MelotTricholomataceaeSx
Clitocybe fragrans (With.) P. Kumm.TricholomataceaeSx
Clitocybe geotropa (Bull. ex DC.) Quél.TricholomataceaeSx
Coprinellus domesticus (Bolton) Vilgalys, Hopple and Jacq. JohnsonPsathyrellaceaeSxx
Coprinopsis nivea (Pers.) Redhead, Vilgalys and MoncalvoPsathyrellaceaeSx
Coprinus comatus (O.F. Müll.) Pers.AgaricaceaeSx
Coprinus lagopus (Fr.) Fr.PsathyrellaceaeSx
Coprinus micaceus (Bull.) Fr.PsathyrellaceaeSx
Coprinus niveus (Pers.) Fr.PsathyrellaceaeSx
Craterellus ignicolor (R.H. Petersen) Dahlman, Danell and SpataforaHydnaceaeSx
Crepidotus applanatus (Pers.) P. Kumm.CrepidotaceaeSx
Crepidotus mollis (Schaeff.) StaudeCrepidotaceaeSx
Gymnopilus pampeanus (Speg.) SingerStrophariaceaeS x
Hygrocybe chlorophana (Fr.) WünscheHygrophoraceaeSx
Hygrophoropsis aurantiaca (Wulfen) MaireHygrophoropsidaceaeSxx
Laetiporus sulphureus (Bull.) MurrillLaetiporaceaeSx
Lentinellus cochleatus (Pers.) P. Karst.AuriscalpiaceaeSx
Lepista sordida (Schumach.) SingerTricholomataceaeS x
Lepista sordida var. lilacea (Quél.) BonTricholomataceaeS x
Leucoagaricus americanus (Peck) VellingaAgaricaceaeSx
Leucoagaricus holosericeus (J.J. Planer) M.M. MoserAgaricaceaeSxx
Leucoagaricus leucothites (Vittad.) WasserAgaricaceaeSxx
Leucoagaricus purpureolilacinus HuijsmanAgaricaceaeSx
Leucoagaricus rubrotinctus (Peck) SingerAgaricaceaeSxx
Leucocoprinus birnbaumii (Corda) SingerAgaricaceaeSx
Leucocoprinus cepistipes (Sowerby) Pat.AgaricaceaeSxx
Lycoperdon perlatum Pers.LycoperdaceaeS x
Lycoperdon umbrinum Pers.LycoperdaceaeS x
Macrolepiota africana (R. Heim) Heinem.AgaricaceaeS x
Macrolepiota procera (Scop.) SingerAgaricaceaeSx
Morchella cf americana Clowez and C. MatherlyMorchellaceaeS x
Morchella anatolica Isiloglu, Spooner, Alli and SolakMorchellaceaeS x
Omphalotus illudens (Schwein.) Bresinsky and BeslOmphalotaceaeS x
Pholiota aurivella (Batsch) P. Kumm.StrophariaceaeSx
Pleurotus luteoalbus BeeliPleurotaceaeSx
Pleurotus populinus O. Hilber and O.K. Mill.PleurotaceaeSx
Pleurotus pulmonarius (Fr.) Quél.PleurotaceaeSx
Polyporus badius (Pers.) Schwein.PolyporaceaeS x
Polyporus tenuiculus (P. Beauv.) Fr.PolyporaceaeS x
Polyporus tuberaster (Jacq. ex Pers.) Fr.PolyporaceaeS x
Schizophyllum commune Fr.SchizophyllaceaeS x
Suillus luteus (L.) RousselSuillaceaeEM x
Termitomyces clypeatus R. HeimLyophyllaceaeSx
Termitomyces microcarpus (Berk. and Broome) R. HeimLyophyllaceaeSx
Termitomyces robustus (Beeli) R. HeimLyophyllaceaeSx
Termitomyces schimperi (Pat.) R. HeimLyophyllaceaeSx
Tricholoma portentosum (Fr.) Quél.TricholomataceaeEMx
Tricholoma saponaceum (Fr.) P. Kumm. TricholomataceaeEMx
Tylopilus niger (Heinem. and Gooss. -Font.) WolfeBoletaceaeEM x
Note: T, trophic groups; S, saprotrophic; P, parasitic; EM, ectomycorrhizal; N, natural forest; P, plantation forest.
Table
Table 3. Significance of explanatory variables for sporocarp composition in the two forest types. The numbers in bold indicate a highly significant effect of environmental variables (p < 0.001).
Table 3. Significance of explanatory variables for sporocarp composition in the two forest types. The numbers in bold indicate a highly significant effect of environmental variables (p < 0.001).
SourcesContributionVariablesPseudo-Fp-Values
Edaphic variables32.33%OM0.28240.001
N0.30570.001
P0.41840.001
OC0.31220.001
Climate60.76%Elevation0.07620.088
Tmin0.52170.001
Spatial factors64.19%Latitude0.77770.001
Note: OM, organic matter; N, total nitrogen; P, available phosphorus; OC, organic carbon; Tmin, minimum daily temperature.
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Kewessa, G.; Dejene, T.; Alem, D.; Tolera, M.; Martín-Pinto, P. Forest Type and Site Conditions Influence the Diversity and Biomass of Edible Macrofungal Species in Ethiopia. J. Fungi 2022, 8, 1023. https://doi.org/10.3390/jof8101023

AMA Style

Kewessa G, Dejene T, Alem D, Tolera M, Martín-Pinto P. Forest Type and Site Conditions Influence the Diversity and Biomass of Edible Macrofungal Species in Ethiopia. Journal of Fungi. 2022; 8(10):1023. https://doi.org/10.3390/jof8101023

Chicago/Turabian Style

Kewessa, Gonfa, Tatek Dejene, Demelash Alem, Motuma Tolera, and Pablo Martín-Pinto. 2022. "Forest Type and Site Conditions Influence the Diversity and Biomass of Edible Macrofungal Species in Ethiopia" Journal of Fungi 8, no. 10: 1023. https://doi.org/10.3390/jof8101023

APA Style

Kewessa, G., Dejene, T., Alem, D., Tolera, M., & Martín-Pinto, P. (2022). Forest Type and Site Conditions Influence the Diversity and Biomass of Edible Macrofungal Species in Ethiopia. Journal of Fungi, 8(10), 1023. https://doi.org/10.3390/jof8101023

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