1. Introduction

Mushrooms are the fruiting bodies of some macro-fungi that produce spores [1]. They are found in the Phylum Basidiomycota and class Basidiomycetes. They are saprophytic and digest their food by secreting enzymes that degrade food extracellularly [2]. Mushrooms lack chlorophyll and cannot obtain energy from the sun through photosynthesis, as like green plants [3]. Instead, during the vegetative growth stage, mushroom mycelia secrete enzymes that break down compounds such as cellulose and lignin present in the substrate. The hyphae then absorb the degraded compounds, and the mycelium enlarges [3].

Mushrooms grow above the ground and mainly consist of two parts: mycelium and fruiting bodies [4]. The mycelium is composed of a tree-like structure called, hyphae, which remain within the substrate. The pileus of the fruiting body may be attached laterally (with no stem), or, if there is a stem, it is usually eccentric, and the gills are decurrent around it, i.e., pleurotoid. Mushroom gill attachment types vary; they can be adnate (gills widely attached to the stem), adnexed (gills narrowly attached to the stem), decurrent (gills running down the stem for some distance), emarginate (gills notched immediately before attaching to the stem), free (gills not attached to the stem), seceding (gills attached but breaking away from the stem at the margin), sinuate (gills smoothly notched and running briefly down the stem), or sub-decurrent (gills running briefly down the stem). The shape of the cap also varies. Different cap shapes include: campanulate (bell-shaped), conical (triangular-shaped), convex (outwardly rounded), depressed (with a low central region), flat (with a uniform top), infundibuliform (deeply depressed), ovate (shaped like half an egg), umbilicate (with a small deep depression), and umbonate (with a central bump or knob) [4].

Mushrooms are consumed as food [5] due to their pleasant taste and aroma. It has been widely employed in traditional medicine [6], bioremediation [7], and waste conversion [8, 9]. Edible mushrooms are invaluable sources of dietary nutrients (proteins, fibres, vitamins and minerals) and nutraceuticals (antioxidant and antitumor agents in the fight against cancer) [10]. However, deadly or poisonous mushrooms are known to produce toxic substances (e.g. alpha amanitin) that are harmful and have lethal effects when consumed [11]. Wild edible mushrooms have been collected from time immemorial, however, concerns have emerged regarding the consumption of unidentified mushrooms within the environment, and the inability of collectors to distinguish between edible and non-edible/poisonous ones.

In nature, mushrooms are widely dispersed and are the earliest form of fungi known to humans [12]. The numerous forests and agroforests in Nigeria remain the primary sources of wild marketable, edible, and medicinal mushrooms, following the dearth of mushroom cultivation industries. The rate of decline in mushroom genetic resources due to the growing agroforest industries, the current change in climate, rate of deforestation, bush burning, and overexploitation of both timber and non-timber products are threatening mushroom diversity [12, 13]. In Nigeria, mushrooms are often overlooked in many biodiversity studies compared to plants and animals [14]. Although an array of edible mushrooms consumed in Nigeria has been reported by Akpaja et al. [14-19], mushrooms are rarely studied, and are relatively underutilized in Nigeria [20].

Indigenous mushrooms are diverse, and morphological identification is insufficient for taxonomy [21] because it can be misleading. Therefore, identifying mushrooms requires a basic understanding of their macroscopic structures. The colour of the powdery print (which is called a spore print) has been used to help classify mushrooms. For most mushrooms, the spore print is formed when the cap is cut off and placed gill-side down (usually overnight). A powdery impression reflecting the shape of the gills (spore print) is formed [22, 23]. Morphological identification also utilizes the size, color, and consistency of the cap and stalk; mode of attachment of the gills to the stalk; spore color in mass; and chemical tests or reactions [22-25]. A general rise in mushroom consumption has been observed in Ile-Ife, but little is known about the diversity of edible mushrooms that could be found in the Ile-Ife. Hence, there is a need to survey, collect, identify, and catalogue the various wild edible mushrooms in the locality using morphological tools, and isolate pure cultures of selected mushrooms.

2. Materials and methods

2.1. Study area

The three Local Government Areas within Ile-Ife metropolis were selected for this study, these include: Ife Central (Latitude 7°33’N and Longitude 4°32’E); Ife North (Latitude 7° 30' 59.99" N and Longitude 4° 26 59.99" E); and Ife South (Latitude 7°12′N and Longitude 4°36’E), respectively. Two towns within each LGA were purposively surveyed due to prevalent agricultural and agroforestry activities: Ipetumodu and Asipa (Ife North), Olode and Odemuyiwa (Ife South), while Obafemi Awolowo University Teaching and Research Farm, a field within the University Campus (including the botanical garden), and Kajola village (Ife Central) LGA. A map of the study area is presented in Fig. 1.

Figure 1. Map of Ife central, Ife north and Ife south local government area of Osun state.

2.2. Sample collection

Survey and collection started from Ife North, through Ife South and then Ife Central LGA. Samples were randomly collected from two farms within each LGA. Systematic and periodic surveys based on information provided by the farmers and rainfall patterns were carried out on farms and gardens in each LGA. While farms and gardens in Ife Central LGA were surveyed daily due to their proximity to the research station, those in Ife North LGA were visited four times on a weekly basis, and surveys for wild edible mushrooms were carried out twice in Ife South, considering the available resources for the study. The materials used for the sample collection included a sharp knife, scissors, sample bags, a digital camera, and a data book. Pictures of the mushroom samples were taken in situ with the digital camera before and after collection. Observations of the substrate and environmental condition from which the mushroom samples were collected. Collections were carried out during the rainy season from May to October, during the day, following reported method [23].

2.3.         Mushroom identification

Samples of mushrooms collected from the field were identified in situ using a Mushroom Chart Book by Anna Del Conte and Laessoe [24], and an Android application: ‘The Mushroom Identifier’ version 2.29, to confirm the scientific name, family, and genus of each sample.

2.4.         Morphological classification of collected mushrooms

The morphological features such as the shape of the pileus, pileal attachment to the stipe, colour of the fruiting body, gill arrangement and attachment, and shape of the stipe were observed and compared with those in the Mushroom Chart Book for identification [24, 25].

2.5.         Spore print determination

Spore prints of the mushroom samples were determined by placing the pileus with their hymenium layer facing down on a white and dark surface for 3 to 6 h for the print of the spheres to be revealed on the collecting surface, and the imprinted colours were recorded.

2.6.         Mushroom tissue culture

Two centimeters of the identified mushroom’s pileus was surface sterilized and blotted with a pair of sterile paper towels.  The blotted tissues were aseptically transferred onto Potato Dextrose Agar (PDA, HiMedia, 39 g/L) media in a laminar flow and incubated for 7 days at room temperature. Pure cultures were obtained via single sporing and stored in slant bottles for further analysis.

3.   Results and discussion

A total of 23 different species belonging to 19 different genera of wild edible mushrooms were identified (some, up to the species level and some to the genera phenological level) across the surveyed locations. Three species of Termitomyces and two species of Pleurotus were identified. Since the study was conducted during the rainy period in the surveyed region, our results are similar to those of Aminuzzaman et al. [26] and Shah et al. [27]. The identified mushrooms are presented in Tables 1-3, while in-situ pictures of the mushrooms, spore prints, and tissue cultures of selected mushrooms are presented in Figs. 2-5, respectively.

Table 1. Mushroom samples identified at Ife central local government area.

S/N	Scientific names of mushrooms	Common/Local Name/Indigenous Name	Substrate	Location
1	Auricularia sp.	Jelly ear mushroom/etí ológbò	Tree trunk	OAU botanical garden
2	Armillaria sp.	Honey cap mushroom	Littered leaves	OAU Campus lawn (Faculty of Education)
3	Clitocybe sp.	Funnel cap Mushroom	Littered leaves	OAU Campus lawn (Faculty of Education)
4	Favolus sp	Honeycomb mushroom/ Olú awọ	Dead tree	OAU botanical garden
5	Hypholoma sp.	Olú wòwó	Decayed tree	OAU botanical garden
6	Lepiota sp.	Olú kòtòpó	Dead tree	Lawn close to OAU gate
7	Laccaria sp.	Amytheist Mushroom	Heap of mulch	OAU Campus lawn (Social sciences)
8	Marasmius oreades	Scotch Bonnet	Decomposed woods	OAU botanical garden
9	Macrolepiota procera	Shaggy Parasol	Soil (loam)	OAU botanical garden
10	Parasola conopilus	Brittle storm mushroom	Soil (loam)	OAU teaching and research farm
11	Pleurotus florida	Olú igi	Dead tree	OAU teaching and research farm, Kajola village
12	Pleurotus pulmonarius	Olú igi	Littered leaves	OAU Campus lawn (Education and social sciences)
13	Termitomyces globulus	Olú ewè/ Olú ògògó	Soil (loam)	Ife Market
14	Russula cyanoxantha	Charcoal burner mushroom	Soil (loam)	OAU botanical garden
15	Volvopluteus gloeicephalus	Field Mushroom	Grass, Soil covered with Straws	OAU teaching & research farm, Kajola village

Table 2. Mushroom samples identified at Ife north local government area.

S/N	Scientific names of mushrooms	Common/Local Name/ Indigenous Name	Substrate	Location
1	Coprinus sp.	Olú ihá or Olú ekù	Palm oil chaff/Spadix	Asipa and Edun-abon
2	Clitocybula lacerata	Olú igi	Tree trunk	Asipa and Edun-abon
3	Lentinus squalrosulus	Olú igi	Tree trunk	Asipa
4	Termitomyces microcarpus	Olú ọrán	Soil (loam)	Asipa
5	Termitomyces clypeatus	Tàkèlè	Soil (loam)	Asipa

 

 

Table 3. Mushroom samples identified at Ife south local government area.

 

S/N	Scientific names of mushrooms	Common/Local Name/Indigenous Name	Substrate	Location
1	Lycoperdon sp.	Puffballs/Olú- ẹyin	Soil (loam)	Olode
2	Marasmiellus sp.	Olú jànpá	Oil Palm branch	Olode and Odemuyiwa village
3	Termitomyces clypeatus	Tàkèlè	Soil (loam)	Olode
4	Termitomyces microcarpus	Olú ọrán	Soil (loam)	Olode

Figure 3. Spore prints of selected mushroom fruiting bodies.

(A. Marasmiellus sp. B. Lepiota sp. C. Lentinus squalrosulus D. Armillaria sp. E. Panellus sp. F. Hypholoma sp.   G. Volvopluteus gloeicephalus H. Coprinus sp.  I. Pleurotus pulmonarius J. Clitocybe sp).

Coprinus sp., Clitocybula lacerata, Termitomyces clypeatus, Termitomyces microcarpus, and Lentinus squalrosulus were identified in Ife North LGA. In addition, Termitomyces clypeatus, T. microcarpus, Lycoperdon sp., and Marasmiellus sp., were collected from the Ife South LGA. Furthermore, Auricularia sp., Volvopluteus gloeicephalus, Termitomyces globulus, Laccaria sp., Favolus sp., Hypholoma sp., Panellus sp., Clitocybe sp., Armillaria sp., Russula sp., Macrolepiota procera, and Parasola conopilus were collected in Ife Central LGA (Table 4). More mushroom species were collected in Ife Central LGA compared to other LGAs, which is attributable to the frequency of visits and surveys in the LGA. The different species of mushrooms identified in Ife Central LGA indicated the rich biodiversity available in the LGA.

 

Table 4. Summary of morphological features of different mushrooms collected across the three local government areas surveyed.

Mushroom	Colour	Cap Shape	Annulus	Stipe (Attachment)	Volva	Gills	Gill Attachment	Spore Print	Substrate
Auricularia sp.	Red	Folded	Absent	Absent	Absent	Pores	Free	Yellow	Tree trunk
Armillaria sp.	Brown	Convex	Absent	Present (Central)	Absent	Equal	Free	White	Littered leaves
Clitocybula lacerata	White	Infudibuliform	Absent	Present (Excentric)	Absent	Equal	Decurrent	White	Tree trunk
Clitocybe sp.	Creamy	Infudibuliform	Absent	Present (Central)	Absent	Varying length	Decurrent	Creamy	Littered leaves
Coprinellus sp.	Brown	Convex	Absent	Present (Central)	Absent	Equal	Adnexed	Black	Oilpalm spadix
Favolus sp.	White	Reniform	Absent	Present (Short& Lateral)	Absent	Maze-like	Decurrent	White	Dead Tree
Hypholoma sp.	Orange	Convex	Absent	Present (Central)	Absent	Forked	Adnate	Clayey	Tree trunk
Lentinus squalrosulus	White	Infudibuliform	Absent	Present (Lateral)	Absent	Equal	White	White (Smoothy)	Tree trunk
Laccaria sp.	Brown	Convex	Absent	Present (Central)	Absent	Equal	Adnate	White	Heap of Mulch
Lepiota sp.	White	Spherical	Absent	Present (Central)	Absent	Equal	Adnate	White	Dead Tree
Lycoperdon sp.	White	Ball	Absent	Present (Short & Central)	Absent	Pores	Free	White	Soil
Marasmiellus sp.	White	Flat	Absent	Present (Central)	Absent	Unequal	Notched	White	Palm Tree
Macrolepiota procera	Brown	Umbonate	Present	Present (Long & Central)	Absent	Equal	Free	White	Soil
Marasmius oreades	Brown	Umbonate	Absent	Present (Central)	Absent	Wide	Adnexed	Creamy-white	Decomposed Woods
Panellus sp.	White	Reniform	Absent	Present (Lateral)	Absent	Forked	Decurrent	White	Chopped Tree
Pleurotus florida	White	Spherical	Absent	Present (Excentric)	Absent	Equal	Decurrent	White	Dead Tree
Pleurotus pulmonarius	Buffy-Cream	Infudibuliform	Absent	Present (Excentric)	Absent	Equal	Decurrent	White	Littered leaves
Parasola conopilus	Creamy	Campanulate	Absent	Present (Central)	Absent	Equal	Notched	Creamt	Soil
Termitomyces clypeatus	White	Depressed	Absent	Present (Lateral)	Absent	Equal	Free	White	Soil
Termitomyces globulus	Dark-grey	Campanulate	Absent	Present (Long & Central)	Absent	Crowded	Sinuately Notched	White	Soil
Termitomyces microcarpus	White	Flat	Absent	Present (Central)	Absent	Equal	Free	White	Soil
Russula cyanoxantha	Black	Depressed	Absent	Present	Absent	Equal, Crowded, Flexible	Notched	White	Soil
Volvopluteus gloeicephalus	White  (pink gills)	Flat	Absent	Present (Central)	Absent	Widely Spaced	Free	Pink	Grass, straws.

The presence of Termitomyces spp. in the three LGAs suggested the relative abundance of this genus across the studied locations. Termitomyces spp. are associated with ant hills in a mycorrhizal/saprophytic relationship which is present in the studied locations. In addition, Oso [28] and Alabi [29] reported that Termitomyces clypeatus, T. globulus, and T. microcarpus are significant to the Yoruba culture and are commonly found in the southwestern states of Nigeria, which host the survey location of the present study.

Similarly, Lentinus squalrosulus has been reported in some states of Nigeria, such as Oyo, Osun, Ondo, and Bayelsa [19]. However, Coprinus sp. was reported in the Delta and Edo states (South-south region of Nigeria) [19, 20]. Gbolagade et al. [19] reported the presence of Lycoperdon sp. in Oyo, Ogun and Edo states while Auricularia sp. has been reported to be found in the South Western states of Nigeria such as Oyo, Ogun, Ekiti and Ondo [19] and in some Southern states of Nigeria such as Delta, Edo and Bayelsa [19, 20]. Volvopluteus sp. has been reported to grow in Edo and Delta states [20].

Species of Pleurotus are in high demand for consumption and are widely spread across the area surveyed in the present study. Pleurotus spp. are repositories of great nutritional value, such as protein, essential amino acids and dietary fibre, and are poor in fat content but enriched with excellent fatty acid content [30, 31]. This mushroom species has also been reported in Ondo, Oyo, Ogun, Ekiti, and some Southern parts of Nigeria, such as Bayelsa and Delta states [16, 18-20].

Species of Favolus, Hypholoma, Panellus, Clitocybe, Armillaria, Russula, Macrolepiota, and Parasola are not commonly harvested mushroom species, and information about their diversity in Nigeria is scarce. This could be attributed to the low rate of spread of their sporocarp in the wild [20]. In addition, the preference of collectors and consumers for the common and widespread edible mushrooms (Pleurotus, Lentinus, and Termitomyces) in the immediate environment may also result in the neglect of these species. However, Macrolepiota spp. has been reported by Osemwegie et al. [23] to be found in some major ecological zones of Nigeria, such as the South West and South-South [19, 20].

Spore prints of the selected mushroom species (Fig. 4) were collected for the laboratory identification. The colours were diverse and included white, black, pink, rusty brown, and creamy white.  These colours are same as those for some of the wild edible mushrooms reported by [22, 24, 25]. The tissues of the identified mushrooms were inoculated on PDA, however, only a few species grew on the media. While mushrooms are classified ecologically as saprophytic, mycorrhizal or parasitic [32], and saprophytic mushrooms are often easily cultivated on growth media.

Figure 4. Culture plates of the selected mushroom species on potato dextrose agar.

the present study, only a few species were successfully cultured on growth media, indicating that some of the collected mushrooms may fall within the mycorrhizal category, thus, requiring additional growth conditions. The culture plates of the tissues grown on the growth medium are presented in Fig. 5.   In-situ images of the mushroom, spore print, and tissue culture plates are presented in Fig. 5a-5c. These include Coprinus sp., Laccaria sp., Lentinus squalrosulus, Panellus sp., Pleurotus florida, and Marasmiellus sp. Bankole and Salami [6] and Son et al. [33] had reported the growth of different mushroom species on PDA. It would be valuable for further studies to investigate the growth conditions of indigenous wild edible mushrooms for cultivation and domestication purposes.

Figure 5a. Selected fruiting bodies were identified based on their sporophores, spore prints, and culture.

Figure 5b. Selected fruiting bodies were identified, showing their sporophores and spore prints.

Figure 5c. Selected fruiting bodies were identified showing their sporophores and culture.

4.   Conclusions

Twenty-three species of wild edible mushrooms species were collected and identified, and a catalogue of the collected mushroom species was developed. This would provide a guide for collectors while hunting mushrooms in Ile-Ife. This is the first survey and biodiversity study of edible mushrooms in the Ile-Ife metropolis. However, further study on mushroom biodiversity would be invaluable in providing useful information about more edible mushrooms available in the region and developing a substantive catalogue that would aid the proper identification of edible mushrooms.

Authors’ contributions

Conceptualization, data curation, formal analysis, investigation, methodology, supervision, validation, writing – review & editing, F.A.B.; Investigation, formal analysis, writing–original draft, writing – review & editing, R.O.A.; Methodology, investigation, methodology, supervision, validation, writing – review & editing, A.O.S.

Acknowledgements

The authors acknowledge the developers of the mushroom identifier app: www.googleplay.com/The-mushroom-identifier, which was used for field identification of the mushrooms.

Funding

This research received no external funding.

Availability of data and materials

All data will be made available on request according to the journal policy.

Conflicts of interest

The authors declare no conflict of interest.

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