BIOTROPIA Vol. 29 No. 3, 2022: 272 - 282 DOI: 10.11598/btb.2022.29.3.1792 272 MACROFUNGAL DIVERSITY IN DIFFERENT VEGETATION COMPOSITIONS IN TEGHARI COMMUNITY FOREST, KAILALI, WEST NEPAL KAUSALYA JOSHI1, HARI SHARAN ADHIKARI1,*, HARI PRASAD ARYAL2 AND LAXMI JOSHI SHRESTHA1 1Department of Botany, Amrit Science College (Tribhuvan University), Kathmandu 44600, Nepal 2Central Department of Botany, Tribhuvan University, Kathmandu 44600, Nepal Received 1 August 2022/Accepted 18 October 2022 ABSTRACT Macrofungi are high-value forest resources that have functionally significant roles in the forest ecosystem. The macrofungal community of three different vegetation compositions, i.e., Sal (Shorea robusta) Forest, Tropical Deciduous Riverine Forest, and Tropical Evergreen Forest of Teghari Community Forest were investigated. Systematic random sampling was made where 60 plots (10 x 10 m) were laid in all different forest types (20 plots in each). A total of 102 macrofungi species were reported belonging to 36 families. Polyporaceae (17 species) was the largest family followed by Tricholomataceae (13 species) and saprophytic fungi were more frequent than mycorrhizal and parasitic fungi. The tropical evergreen forest was rich in macrofungi (59 species) followed by sal forest (40 species) and tropical deciduous riverine forest (38 species). Macrofungal diversity was directly related to surrounding host species. Similarly, increased soil moisture and canopy cover intensified the abundance of saprophytic fungi. The species richness was increased with increasing organic carbon, canopy, moisture, pH, and litter cover. However, soil nitrogen, phosphorus, and potassium were less significant in affecting species richness. Also, the disturbance was negatively correlated with the species richness of macrofungi. This study highlights the hidden diversity which is necessary for the conservation of macrofungi, to optimize forest ecosystem integrity and resilience against biotic and abiotic agents. Keywords: macrofungal diversity, sal forest, species richness, tropical evergreen forest, tropical riverine forest INTRODUCTION Biodiversity is simply defined as the presence of the total organism of a particular group at a particular time in a particular area. Conservation of these natural resources is the priority for ecosystem functioning as well as human welfare. Fungi are an enormous usly diverse group of organisms ranging from microscopic to macroscopic forms that grow mostly in the dead and decaying substrate. They appear in all seasons, mostly rainy season, wherever nutrient organic matters or decomposed products are easily available (Jha & Tripathi 2012). Macrofungi are a group of higher fungi that produce mature spore-bearing fruiting bodies, which are visible to the naked eye (Chang & Miles 1992). They are known to inhabit diverse kinds of habitats varying in the composition of their tree species and substrates. Based on ecology, they are parasitic or saprophytic or may show some mycorrhizal associations with vascular plants (Kumar & Sharma 2011). However, some macrofungi are neutral to the abundance of dominant tree species, in particular, habitat type (Zhang & Zak 1998). The relationship between the tree and fungal communities is reflected in host trees affecting fungal specialization and providing unique habitat availability and different resource quality. The composition and structure of aboveground vegetation are responsible for diverse macrofungi communities (Buee et al. 2011). *Corresponding author, email: aharisharan@gmail.com Macrofungal diversity in different vegetation composition in Kailali, West Nepal – Joshi et al. 273 Generally, macroscopic fruiting bodies of the fungi is called mushroom which can be epigeous or hypogeous and vary in different shape and sizes. They are fleshy, sub-fleshy, or sometimes leathery and woody and bear their fertile surface either on lamellae or lining the tubes, opening out through pores. The most suitable condition for the growth of carpophores depends upon the high humidity, nutritionally rich substrate, and warm atmospheric temperature (Dickinson & Lucas 1979). Similarly, other environmental conditions such as geographic location, light, and surrounding vegetation types also play a major role in the distribution of the macrofungi (Sibounnavong et al. 2008). Diversity-related studies are carried out in different forests but their relationship with higher plants was poorly explored except these studies such as Pradhan (2013), Baral et al. (2015), and Bhandari and Jha (2017). This study aimed to to optimize forest ecosystem integrity and resilience against biotic and abiotic agents, by looking at the effect of different vegetation characteristics and environmental factors on macrofungal species composition and richness in the tropical region of western Nepal. MATERIALS AND METHODS Study Sites The study was carried out in three different vegetation patches within Teghari Community Forest in the tropical riverine belt of Kailali District, West Nepal (Fig. 1). The study area lies between latitudes from 28°50'45" N to 28°51'01" N and longitude 80°33 ̍ 3" E to 80°33'13" E, covering an area of 340 ha. The altitude range of the study area is 155 - 254 masl. Meteorological data of the Dhangadi Airport in the year 2019 was obtained from the Department of Hydrology and Meteorology, Government of Nepal which revealed that the study area is represented by a tropical climate and receives an average of 1,406.6 mm annual rainfall with the highest monthly rainfall happens in July (466.9 mm) and the lowest in May (5 mm). The highest monthly mean temperature happens in May (40.41°C) and the lowest in January (6.77 °C). Figure 1 Map of the study area BIOTROPIA Vol. 29 No. 3, 2022 274 Study Design The Teghari Community Forest was selected for the field study as it has three different forest types at the same elevation, i.e., Sal Forest, Tropical Riverine Deciduous Forest, and Tropical Evergreen Forest. Shorea robusta (Sal) is the dominant tree species in the Sal Forest which forms magnificent forest stand on the edges of the Godawari River. The tropical deciduous riverine forest is also located similarly and is mainly dominated by Acacia catechu and Dalbergia sissoo along with Bombax ceiba, Syzygium cumini, Adina cordifolia, Hollarrhena pubescens, Murraya koenginii, Aegle marmelos, and Semicarpus anacardium. The tropical evergreen forest lies on the Northwest side of Mahakali Highway and is dominated by Terminalia alata, Lagerstroemia parviflora, Terminalia bellerica, Ficus religiosa, Schleichera oleosa, Aegle marmelos, and Cassia fistula. Mallotus phlippensis is present all over the study area. In each forest type, rectangular plots of 10 × 10 m were established. The number of plots to be sampled were determined based on the spatial area of each forest. Field Sampling Detailed sampling of macrofungi diversity was made by applying a systematic random method within the period of June - October 2019, where plots were laid in each forest type. A total of 20 plots were laid in each forest type along with the two transects for maintaining an inter-plot distance of at least 20 m (Baral et al. 2015). Presence or absence data of macrofungal species were recorded in each plot. Biophysical variables, such as tree canopy cover, litter cover, and anthropogenic disturbances (trampling, fire grazing, non-degradable waste, etc.) were also recorded in each plot. Tree canopy cover and litter cover (in percentage) was estimated visually. For tree canopy cover, observation was made from the middle of each plot. Soil samples were collected at a depth of 15 cm from four corners and at the middle of each plot using a soil digger. The soil samples from each plot were mixed thoroughly. From the mixed soil sample, about 200 g of soil sample was taken and put in a zipper polythene bag. The soil samples were air-dried in shade for a week and stored in airtight plastic bags until laboratory analysis. The physiochemical parameters of soil, such as soil pH, moisture, organic carbon, nitrogen, potassium, and phosphorus were assessed using a standard soil analysis manual (Zobel et al. 1987). Macrofungal specimens were collected, preserved (dry), and taken to National Herbarium (KATH) in Lalitpur, Nepal. Collected specimens were studied based on their morphological characters and ecology with the help of several websites, such as https://www.mushroomexpert.com and http://www.indexfungorum.org. Finally, the identification of specimens was confirmed using relevant literature (Pacioni & Lincoff 1981; Adhikari 2014; Laessoe 2013) along with identification conducted by macrofungi expert. All of the collected macrofungal specimens were deposited in ASCOL herbarium, Amrit Science College, Kathmandu, Nepal. Data Analysis All data were entered in Microsoft Excel 2010 for further analysis. Pearson correlation method was used to know the effect of a different environmental variable on macrofungal diversity. Simpson’s Diversity Index (Simpson 1949) and Shannon-Wiener Index (Shannon & Weaver 1963) were also calculated. Regression analysis was performed using SPSS Version 20 and Microsoft Excel version 2010. Species composition of different macrofungi species along with different environmental components were evaluated by Canonical Correspondence Analysis (CCA). RESULTS AND DISCUSSION Macrofungal Diversity in Different Vegetation Composition A total of 102 macrofungi consisting of Ascomycetes-5 and Basidiomycetes-97 species were documented, in which 100 species were identified up to species level and 2 species were identified up to genus level. Out of the 36 families, 17 species belonged to the Polypora- ceae family, 13 species to Tricholomataceae, 11 species to Marasmiaceae, 9 species to Agarica- ceae, 8 species to Coprinaceae, 4 species each to Russulaceae and Xylariaceae, 2 species representing each of the Cortinariaceae, Ento- Macrofungal diversity in different vegetation composition in Kailali, West Nepal – Joshi et al. 275 lomataceae, Fomitopsidaceae, Ganodermataceae, Hydnangiaceae, Podoscyphaceae, and Suillaceae family and the rest of the family was represented by single species only (Fig. 2). Tropical evergreen forest harbored the highest macrofungal diversity (59) in all three different substrates (Fig. 3), followed by Sal Forest (40) and tropical deciduous riverine forest (38). The maximum numbers of macrofungi were found growing on the soil, followed by wooden logs. The least number of macrofungal species were found growing on litters in all forest types. The present study relates to a study conducted in India where macrofungi were reported in various habitats, like wood, litter, and moist soil, among others (Nagaraju et al. 2014). As compared to litter and wood, the soil was the most important substrate for maintaining macrofungal diversity in all three forest types studied. In our study, higher number of macrofungi were grown on moist soil compared to those on litter and decaying wood. These findings resemble the previous findings of a study conducted by Bhandari and Jha (2017). Figure 2 Number of species with their respective family Figure 3 Distribution of macrofungi based on their habitat in different forests BIOTROPIA Vol. 29 No. 3, 2022 276 Based on the ecology of macrofungi, the maximum number of macrofungi were consisted of saprophytes, followed by mycorrhizal and parasitic macrofungi, while the least number belonged to termitophilous macrofungi. The mycorrhizal fungi serve as an extension of the plant root system, exploring soil far beyond the roots and transporting water and nutrients to the roots (Tapwal et al. 2013). The flourishing of carpophores is enhanced by litter accumulation and decomposition as well as the presence of extracellular microbial enzymes (Pushpa & Purushothama 2012). The rapid change in the weather and high response of mycelia were among the main factors for the increasing number of saprotrophic fungi (Pradhan et al. 2012). A similar result was obtained in the research of Topwal et al. (2013) and Dey et al. (2016). Higher species diversity in Basidiomycota compared to Ascomycota is probably contributed by a higher number of mycorrhizal species found on the soil as studies have shown that soil moisture and decaying litter facilitate many diverse macrofungi (Muller & Schmit 2007). Simpson’s Diversity Index (Table 1) was found to be the highest in Tropical Deciduous Riverine Forest and Tropical Evergreen Forest (0.91) in comparison to Sal Forest (0.88). Similarly, Shannon-Wiener Diversity Index was also found to be the highest in Tropical Evergreen Forest (2.91) followed by Tropical Deciduous Riverine Forest (2.77) and Sal Forest (2.53). The presence of diverse kinds of macrofungi communities is specifically related to the dominant tree species of the forest has been confirmed by many other studies (Straatsma & Krisai-Greilhuber 2003; Gates et al. 2011; O’Hanlon & Harrington 2011; Bhandari & Jha 2017; Collado et al. 2021; Kutszegi et al. 2021). Such variation may be attributed to microclimate conditions (Santos-Silva et al. 2011) and forest management practice (Kouki & Salo 2020). The high macrofungal diversity in the tropical evergreen forest is mainly related to high soil moisture and greater cover of tree species. The high diversity may be also due to suitable habitat, such as soil moisture, litter, and canopy cover which help to maintain sufficient moisture (Trudell & Edmonds 2004). A tropical deciduous riverine forest located on the water edges has a more open canopy and less humidity in the soil which creates a less suitable habitat for the growth of macrofungi. Also, thinning of trees caused a decrease in fruit-body production of the delicate and fragile macrofungi, but this effect varied greatly depending on the season, the macrofungi fruiting pattern and the levels of trees thinning (Luoma et al. 2004). Therefore, thinning and pruning, which are common silvicultural activities in the community forests of Nepal (Shrestha et al. 2010), might also affect the composition and abundance of macrofungi. Similarly, Sal Forest has a magnificent stand of tall trees and has a more open canopy in comparison to tropical evergreen forest. The growth of macrofungal species like Pycnoporus cinnarius and Scleroderma cepa was specifically recorded in Sal Forest. A similar finding was also reported by Prasad & Pokhrel (2017) at Amrite community forest, Kapilvastu District (Central Nepal), which might be due to the host specificity of macrofungi with particular plant species. The presence of specific macrofungi communities in the present study may be due to host preferences which were related to the findings by Ding et al. (2011) and Lang et al. (2011). Species Richness of Macrofungi and Different Environmental Variables The Canonical Correspondence Analysis (CCA) revealed the relationship between macrofungi species composition and environmental gradient (Table 2). The analysis results indicated the effective separation of species along the main gradient (Table 3). Table 1 Diversity indices of macrofungi in different vegetation stands Forest stands Simpson’s index Simpson’s diversity index Shannon-Wiener diversity index Sal Forest 0.12 0.88 2.53 Tropical Deciduous Riverine Forest 0.09 0.91 2.77 Tropical Evergreen Forest 0.09 0.91 2.91 Macrofungal diversity in different vegetation composition in Kailali, West Nepal – Joshi et al. 277 Table 2 Summary of the results of Canonical Correspondence Analysis Axes 1 2 3 4 Total inertia Eigenvalues 0.369 0.34 0.279 0.208 11.381 Species-environment correlations 0.886 0.903 0.945 0.88 Cumulative % variance of species data 3.2 6.2 8.7 10.5 Cumulative % variance of species-environment data 18.7 35.9 50 60.5 Sum of all canonical eigenvalues 1.978 Table 3 Relative importance of environmental variables and their significance (P value) on macrofungal species composition derived by using the Monte Carlo permutation test from the Canonical Corrrespondence Analysis with 9999 replications Environmental variable Abbreviation F P Organic carbon Orgcarb 0.914 0.682 pH pH 1.086 0.227 Moisture Moist 1.249 0.11 Nitrogen Nitro 1.057 0.347 Phosphorus Phosp 1.176 0.182 Potassium Potas 1.4 0.162 Litter Litter 1.33 0.046 Canopy cover Canop 1.033 0.36 Disturbances Distrb 1.209 0.052 Our study showed that environmental variables, such as moisture, pH, canopy, organic carbon, nitrogen, phosphorus, potassium, and anthropogenic disturbances had significant effect on the distribution and composition of macrofungi. Organic carbon, moisture, pH, canopy, litter, and macrofungal species richness were positively correlated and was comparable with the study of Bhandari and Jha (2017). This finding indicated that these environmental variables play a vital role in shaping macrofungi communities. However, other soil properties such as nitrogen, phosphorus, and potassium were negatively correlated with the species richness. There were no significant effects on the increasing disturbances with the abundance of macrofungi (Fig. 4). -0.6 1.0 -0 .8 0 .8 gea_lexpod_des sch_une mar_eus ter_pus rus_ceastr_tussui_tus pod_ata col_nis ple_ensscl_epa xyl_lon xyl_phagan_tum spo_lor hex_ida lac_ata mar_cus mic_pus pyc_nus hyg_lus tri_ora myc_ptaama_iae lep_ria pol_ius ale_tia leu_onipsi_ata omp_era aga_sis hel_ata aur_ata len_aju lep_pes pan_vus mar_lus cya_tus mar_lislac_nda ear_osa ent_num cla_ata can_ius mac_des lep_ata hyg_ica lae_eus rus_ria leu_tusdae_ina cre_lis aga_tus xyl_ila lyc_eumcop_tus gan_dum cop_ans oud_atalac_ina tra_ens cop_pus ter_zus dal_ica mar_ula rus_ans ent_tum rus_sps inf_bba cal_osa cre_lis psa_ata mar_ans pol_nus tra_lor con_ens mac_nii lep_nea pan_tus col_ila cli_mis cop_tus ple_ius mar_dus arm_ens ant_ina tra_ans ram_cta hex_uis fom_ius ter_nus hyg_eus sui_dus psa_mis psa_pes cli_sps pan_cus pos_ica cop_lis len_ina mic_pes PH Orgcarbo Moist Nitrogen phosph Potas Canop Litter Distrb Figure 4 CCA biplot representing the effect of environmental variables on macrofungal species composition BIOTROPIA Vol. 29 No. 3, 2022 278 Macrofungi species, like Pycnoporus cinnarius (pyc_ius), Scleroderma cepa (scl_epa), Agaricus arvensis (aga_sis), Tricholomopsis decora (tri_ora), Termitomyces microcarpus (ter_pus), Aleuria aurantia (ale_tia) and Lentinus sajorcaju (len_aju), etc. showed strong presence in the Sal Forest and were more resistant to disturbances. Tropical Deciduous Riverine Forest was dominated by the macrofungi species, like Ramaria stricta (ram_cta), Antrodia juniperina (ant_ina), Fomes fomentarius (fom_ius), Hexagonia tenuis (hex_uis), Trametes elegans (tra_ans), Clitocybe infundibuliformis (cli_mis), Lenzites betulina (len-ina), Panus fasciatus (pan_tus), Coltricia perennis (col_nis) and Microporus xanthopus (mic_pus), etc. which favored more open canopy. Similarly, Tropical Evergreen Forest harbored macrofungi species, like Macrolepiota rickenii (mac_nii), Marasmius haematocephalus (mar_lus), Daldinia concentrica (dal_ica), Marasmius androsaceus (mar_eus), Lepiota clypeolaria (lep_ria), Macrolepiota rhacodes (mac_des), Lacrymaria lacrymabunda (lac_nda), Lepiota clypeolaria (lep_ria), Lycoperdon subcretaceum (lyc_eum), Cyathus striatus (cya_tus) Microporus xanthopus (mic_pus) and Podoscypha multizonata (pod_ata). Our present study also showed that the species found in the Tropical Deciduous Riverine Forest and Tropical Evergreen Forests were more similar than those in the Sal Forest. However, some macrofungi species, like Schizophyllum commune (sch_une), Polyporus arcularius (pol_ius), Termitomyces tylerianus (ter_nus), Microporus xanthopus (mic_pus), and Geastrum triplex (gea_lex) were found in all three forest types. The CCA biplot showed environmental variables and aboveground vegetation were the major components for determining macrofungi composition (Fig. 4). The overlapping of macrofungi species was due to a similar ecological niche. As we found in our present study, most of the soil fungi such as Geastrum triplex (gea_lex), Podocypha petaloides (pod_des), Suillus granulatus (sui_tus), and Russula species occurred toward the moisture. Our study also showed that the presence of thin canopy cover and low soil moisture seemed to enhance the growth of wood-inhabiting fungi, such as Spongipollis unicolar (spo_lar), Crepodotus mollis (cre_lis), Xylaria sp., Trametes elegans (tra_ans), Antrodia juniperina (ant_ina), Fomes fomentarius (fom_ius), Hexagonia tenuis (hex_uis), Micrporus vernicipes (mic_pes) and Lenzites betulina (len_ina), which were mostly presented opposite direction to the moisture and pH. Organic carbon was also one of the major components which control the distribution pattern of soil fungi. Macrofungi species, like Termitomyces sp., Rusulla sp., Macrolepiota rhacodes (mac_des), Omphalina umbellifera (omp_era), and Marasmiellus ramealis (mar_lis) were found predominantly toward the direction of organic carbon. In our present study, disturbances seemed to have a poor impact on macrofungi species composition. Most of the fleshy, soft, and gilled macrofungi, like Clitocybe sp., Psathyrella obtusata (psa_ata), Agaricus augustus (aga_tus), Lepiota clypeolaria (lep_ria) and Coprinus disseminates (cop_tes) were favored by the higher soil pH. Species richness of macrofungi increased with the increasing soil organic carbon, moisture, pH, litter coverage, and canopy coverage. Soil pH and organic carbon ranged from 4.06 to 7.07 and 0.79 to 5.67, respectively. Similarly, soil moisture, litter cover, and canopy cover ranged from 7.5 to 45.46%, 11 to 49%, and 15 to 95%, respectively. Among all environmental variables, organic carbon, soil moisture, soil pH, litter cover and canopy cover had the most significant positive relationship with macrofungi species richness (Fig. 5). Also, the species richness of macrofungi showed a weak positive relationship with disturbances. Soil nitrogen, phosphorus, and potassium were negatively correlated with macrofungi species richness but the result was statistically insignificant (Table 4). Species diversity of macrofungi depends on their particular habitat. Geographic location, elevation, temperature, the humidity of air and soil, light, surrounding flora, and anthropogenic activity greatly influence the growth and reproduction of macrofungi (Zervakis & Venturella 2007; Topwal et al. 2013). Soil moisture is one of the most important environmental factors responsible for affecting the growth of the macrofungi (Kropp & Albee 2002). Present findings also showed increasing species richness is affected by the increasing moisture content of the soil. This finding was similar to the research of Bhandari and Jha (2017). The fungal diversity studies in Greece and Sicily (Venturella & Zervakis 2000; Zervakis & Venturella 2002) confirmed that fungi require a certain level of moisture; rainfalls, Macrofungal diversity in different vegetation composition in Kailali, West Nepal – Joshi et al. 279 Figure 5 Correlation between macrofungi species richness and organic carbon, moisture, soil pH, canopy cover, and litter cover Notes: Each point in each figure represents a sampling plot; Total number of sample plots = 60. Less number of points in the figure may be due to the overlapping of the data among the plots. The fitted line is based on the linear regression model for sampling plots. Table 4 Pearson correlations between environmental variables and species richness of macrofungi Soil pH Organic carbon Moisture N P K Canopy cover Litter cover Disturbances Species richness 0.506** 0.519** 0.580** -0.074 -0.183 -0.14 0.513** 0.612** 0.034 Notes: * = Correlation is significant at P < 0.05 level; ** = Correlation is significant at P < 0.01 level. air humidity, and soil moisture, which are all significant factors. Canopy cover and litter cover had also provided positive influence on macrofungal species richness in the studied area and a similar result was found by Baral et al. (2015). The result unveiled that canopy cover plays a vital role in increasing macrofungal diversity in forests which was also reinforced by other previous findings (Dighton et al. 1986; Bonet et al. 2004; Sysouphanthong et al. 2010; Santos-Silva et al. 2011). The reason is likely to be the presence of more substrate on the forest floor and high humidity which favor the growth of more fungal species (Lodge et al. 2004). Litter is an important component of every ecosystem and it constitutes the major source of organic matter in the soil. The removal of litter directly affects the diversity and growth of macrofungi (Eaton et al. 2004; Sayer 2006). When the forest floor is covered with layers of well-decomposed BIOTROPIA Vol. 29 No. 3, 2022 280 leaves, saprotrophic fungi are favored by this organic resource which maintains the temperature and moisture of the surrounding area (Fernandez-Toiran et al. 2006). Also, the abundance of macrofungal species is closely correlated with soil organic matter and other soil parameters (Zamora-Martinez & de Pascual-Pola 1995; Engola et al. 2007). The growth of saprophytic fungi was enhanced at pH 7 or 8; while the ectomycorrhizal species showed the peak growth at pH 5 or 6 (Yamanaka 2003). Soil pH was also a major abiotic component responsible for changing macrofungi communities and was found to be positively correlated with macrofungi species richness. The pH range of 5 to 6 favors the growth of soil fungi (Bhandari & Jha 2017; Pavithra et al. 2016; Zhang et al. 2016). In this study, it was observed that there were no direct relationships between macrofungi richness and other soil parameters, such as nitrogen, phosphorus, and potassium with macrofungi diversity. However, the importance of forest soil chemistry parameters in fungal species distributions has also been reported by Hansen (1988) and Ruhling & Tyler (1990). CONCLUSION The study area is rich in macrofungal diversity with species’ richest families being the Polyporaceae followed by Tricholomataceae, Marasmiaceae, Agaricaceae, and Coprinaceae. The presence of diverse kinds of vascular plants and different environmental conditions in different forest types have created unique habitat for the growth and development of a wide variety of macrofungi species. Species diversity is higher in moist and dense canopy forests such as Tropical Evergreen Forests and Sal Forest compared to that in the open and dry Tropical Deciduous Riverine Forests. Soil moisture, organic carbon, soil pH, litter cover, and tree canopy cover are the most important variables affecting macrofungal diversity. ACKNOWLEDGMENTS The authors are grateful to the Forest Users Groups of Teghari Community Forest and Mr. Yagya Raj Bhatta for their support and cooperation in field. We are also thankful to the National Herbarium and Plant Laboratories (KATH) for the specimen identification which had been crucial for this study. Finally, we thank anonymous reviewers and editors for their valuable comments and suggestions on the earlier versions of the manuscript. REFERENCES Adhikari MK. 2014. Mushrooms of Nepal. 2nd edition. Durrieu G, Cotter VT, Editors. Kathmandu (NP): KS Adhikari Nepal. Baral S, Thapa-Magar KB, Karki G, Devkota S, Shrestha BB. 2015. Macrofungal diversity in community- managed Sal (Shorea robusta) forests in central Nepal. Mycol 6: 151-7. DOI: https://doi.org/ 10.1080/21501203.2015.1075232 Bhandari B, Jha SK. 2017. Comparative study of macrofungi in different patches of Boshan community forest in Kathmandu, Central Nepal. Bot Orient J of Plant Sci. 11: 43-8. DOI: https://doi.org/10.3126/botor.v11i0.21032 Bonet JA, Fischer CR, Colinas C. 2004. 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 203: 157-75. DOI: https://doi.org/ 10.1016/j.foreco.2004.07.063 Buée M, Maurice JP, Zeller B, Andrianarisoa S, Ranger J, Courtecuisse R, Le Tacon F. 2011. Influence of tree species on richness and diversity of epigeous fungal communities in French temperate forest stand. Fun Ecol 4: 22-31. DOI: https://doi.org/ 10.1016/j.funeco.2010.07.003 Chang ST, Miles PG. 1992. Mushroom biology: A new discipline. Mycol 6: 64-5. DOI: https://doi.org/ 10.1016/S0269-915X(09)80449-7 Collado E, Bonet JA, Alday JG, de Aragón JM, de-Miguel S. 2021. Impact of forest thinning on aboveground macrofungal community composition and diversity in Mediterranean pine stands. Ecol Indi 133: 1-9. DOI: https://doi.org/10.1016/j.ecolind.2021. 108340 Dey S, Paul M, Sarma GC, Sarma TC. 2016. Occurrence of macrofungi in Garbhanga Reserve Forest, Kamriup District, Assam (India). Int J Adv Res 4: 166-74. DOI: http://dx.doi.org/10.21474/ IJAR01/633 Dickinson C, Lucas J. 1979. The encyclopedia of mushrooms. New York (US): GP Putnam's Sons. Dighton J, Poskitt JM, Howard DM. 1986. Changes in occurrence of basidiomycete fruit bodies during Macrofungal diversity in different vegetation composition in Kailali, West Nepal – Joshi et al. 281 forest stand development: with specific reference to mycorrhizal species. Trans Br Mycol Soc 87: 163-71. DOI: https://doi.org/10.1016/S0007- 1536(86)80017-1 Ding B, Yin Y, Zhang F, Li Z. 2011. Recovery and phylogenetic diversity of culturable fungi associated with marine sponge’s Clathrina luteoculcitella and Holoxea sp. in the South China Sea. Marine Biotech 13: 713-21. DOI: https://doi.org/10.1007/s10126-010-9333-8 Eaton RJ, Barbercheck M, Buford M, Smith W. 2004. Effects of organic matter removal, soil compaction, and vegetation control on collembolan populations. Pedobiol 48: 121-8. DOI: https://doi.org/10.1016/j.pedobi.2003.10.001 Engola APO, Eilu G, Kabasa JD, Kisovi L, Munishi PKT, Olila D. 2007. Ecology of edible indigenous mushrooms of the Lake Victoria Basin (Uganda). Res J of Biol Sci 2: 62-8. DOI: https://medwelljournals.com/abstract/?doi=rjbsci .2007.62.68 Fernández-Toirán LM, Ágreda T, Olano JM. 2006. Stand age and sampling year effect on the fungal fruit body community in Pinus pinaster forests in central Spain. Bot 84: 1249-58. DOI: https://doi.org/ 10.1139/b06-087 Gates GM, Mohammed C, Wardlaw T, Ratkowsky DA, Davidson NJ. 2011. The ecology and diversity of wood-inhabiting macrofungi in a native Eucalyptus oblique forest of southern Tasmania, Australia. Fung Ecol 4: 56-67. DOI: https://doi.org/ 10.1016/j.funeco.2010.07.005 Hansen PA. 1988. Prediction of macrofungal occurrence in Swedish beech forests from soil and litter variable models. Veget 78: 31-44. DOI: https://doi.org/10.1007/BF00045637 Jha SK, Tripathi NN. 2012. Diversity of macrofungi in Shivapuri national park of Kathmandu valley, Nepal. Biol For An Intl J 4: 27-34. ISSN No. (Print): 0975-1130 ISSN No. (Online): 2249-3239 Kouki J, Salo K. 2020. Forest disturbances affect functional groups of macrofungi in young successional forests–harvests and fire lead to different fungal assemblages. For Ecol Manag 463: 118039. DOI: https://doi.org/10.1016/ j.foreco.2020.118039 Kropp BR, Albee S. 1996. The effects of silvicultural treatments on occurrence of mycorrhizal sporocarps in a Pinus contorta forest: a preliminary study. Biol Cons 78: 313-8. DOI: https://doi.org/10.1016/S0006-3207(96)00140-1 Kumar S, Sharma YP. 2011. Diversity of wild mushrooms from Jammu and Kasmir (India). In Proceedings of the 7th International Conference on Mushroom Biology and Mushroom Products (ICMBMP7). 4 October 2011. p. 568-77. DOI: https:// www.researchgate.net/ publication/224946419 Kutszegi G, Siller I, Dima B, Merényi Z, Varga T, Takács K, ..., Ódor P. 2021. Revealing hidden drivers of macrofungal species richness by analyzing fungal guilds in temperate forests, West Hungary. Comm Ecol 22(1): 13-28. DOI: https://doi.org/10.1007/ s42974-020-00031-6 Laessoe T. 2013. Mushrooms and toadstools. London (GB): Dorling Kindersley Book. Lang C, Seven J, Polle A. 2011. Host preferences and differential contributions of deciduous tree species shape mycorrhizal species richness in a mixed central European forest. Mycor 21: 297-308. DOI: https://doi.org/10.1007/s00572-010-0338-y Lodge DJ, Ammirati JF, O’Dell TE, Mueller GM, Huhndorf SM, Wang CJ, ..., Czederpiltz DL. 2004. Terrestrial and lignicolous macrofungi. In: Biodiversity of Fungi: Inventory and Monitoring Methods. Mueller GM, Bills GF, Foster MS, editors. New York (US): Academic Press. p. 127- 58. DOI: https://doi.org/10.1016/B978- 012509551-8/50011-8 Luoma DL, Eberhart J, Molina R, Amaranthus MP. 2004. Response of ectomycorrhizal fungus sporocarp production to varying levels and patterns of green- tree retention. For Ecol Manag 202: 337-54. DOI: https://doi.org/10.1016/j.foreco.2004.07.041 Mueller GM, Schmit JP. 2007. Fungal biodiversity: What do we know? What can we predict?. Biodivers Conserv 16: 1-5. DOI: https://doi.org/10.1007/ s10531-006-9117-7 Nagaraju D, Kumar S, Kunwar IK, Manoharachary C. 2014. Fungi occurring on diversified habitats around some sanctuaries and water-bodies of Telangana and Andhra Pradesh, India. J Exp Biol 2: 24-8. ISSN: 2320-8694 URL: http://www.jebas.org/00200520102014/N... O'Hanlon R, Harrington TJ. 2011. Diversity and distribution of mushroom-forming fungi (Agaricomycetes) in Ireland. In Biology and Environment: Proceedings of the Royal Irish Academy 111: 117-33. DOI: https://doi.org/ 10.3318/BIOE.2011.111.10 Pacioni G, Lincoff G. 1981. Simon and Shuster's Guide to Mushrooms. New York (US): Simon and Schuster. Pavithra M, Sridhar KR, Greeshma AA, Karun NC. 2016. Spatial and temporal heterogeneity of macrofungi in the protected forests of Southern India. J Agri Tech 12: 105-24. ISSN 1686-9141 Pradhan BA. 2013. Comparative study on the predictive ability of the decision tree, support vector machine and neuro-fuzzy models in landslide susceptibility mapping using GIS. Comput Geosci 51:350-65. DOI: https://doi.org/10.1016/j.cageo.2012.08.023 Pradhan P, Dutta AK, Paloi S, Roy A, Acharya K. 2016. Diversity and distribution of macrofungi in the eastern himalayan ecosystem. EurAsian J Biosci BIOTROPIA Vol. 29 No. 3, 2022 282 10: 1-12. DOI: http://dx.doi.org/10.5053/ ejobios.2016.10.0.1 Prasad D, Pokhrel B. 2017. Mushroom Diversity of Amrite Community Forest, Kapilvastu, Nepal. Himal Biodiv 5: 84-91. DOI: https://doi.org/ 10.3126/hebids.v5i1.36158 Pushpa H, Purushothama KB. 2012. Biodiversity of mushrooms in and around Bangalore (Karnataka), India. Amer-EurAsian J Agri Env Sci 12:750-9. ISSN: 1818-6769 URL: http://www.idosi.org/ aejaes/jaes12(6)12/10.pdf Rühling Å, Tyler G. 1990. Soil factors influencing the distribution of macrofungi in oak forests of southern Sweden. Ecogr 13: 11-8. DOI: https://doi.org/10.1111/j.1600- 0587.1990.tb00584.x Santos-Silva C, Gonçalves A, Louro R. 2011. Canopy cover influence on macrofungal richness and sporocarp production in montado ecosystems. Agro. Sys 82: 149-59. DOI: https://doi.org/ 10.1007/s10457-011-9374-7 Sayer EJ. 2006. Using experimental manipulation to assess the roles of leaf litter in the functioning of forest ecosystems. Biol Rev 81: 1-31. DOI: https://doi.org/10.1017/S1464793105006846 Shannon СВ, Weaver W. 1963. The Mathematical Theory of Communication. Urbana (US): The University of Illinois Press. Shrestha UB, Shrestha BB, Shrestha S. 2010. Biodiversity conservation in community forests of Nepal: Rhetoric and reality. Int J Biod Cons 2: 98-104. DOI: https://doi.org/10.5897/IJBC.9000016 Sibounnavong P, Cynthia CD, Kalaw SP, Reyes RG, Soytong K. 2008. Some species of macrofungi at puncan, carranglan, nueva ecija in the Philippines. J of Agri Tech 4: 105-15. http://www.ijat-aatsea. com/pdf/Nov_v4_n2_08/11%20IJAT2008-42-R. pdf Simpson EH. 1949. Measurement of diversity. Nature 163(4148): 688. DOI: https://doi.org/10.1038/ 163688a0 Straatsma G, Krisai-Greilhuber I. 2003. Assemblage structure, species richness, abundance, and distribution of fungal fruit bodies in a seven-year plot-based survey near Vienna. Mycol Res 107: 632-40. DOI: https://doi.org/10.1017/ S0953756203007767 Sysouphanthong P, Thongkantha S, Zhao R, Soytong K, Hyde KD. 2010. Mushroom diversity in sustainable shade tea forest and the effect of fire damage. Biodivers Conserv 19: 1401-15. DOI: https://doi.org/10.1007/s10531-009-9769-1 Tapwal A, Kumar R, Pandey S. 2013. Diversity and frequency of macrofungi associated with wet ever green tropical forest in Assam, India. Biodiversitas 14(2): 73-8. https://doi.org/10.13057/biodiv/ d140204 Trudell SA, Edmonds RL. 2004. Macrofungus communities correlate with moisture and nitrogen abundance in two old-growth conifer forests, Olympic National Park, Washington, USA. Can J Bot 82: 781-800. DOI: https://doi.org/10.1139/ b04-057 Venturella G, Zervakis G. 2000. Comparative evaluation of macro-mycetes diversity in Sicily and Greece. Bot Chron 13: 419-28. Yamanaka T. 2003. The effect of pH on the growth of saprotrophic and ectomycorrhizal ammonia fungi in-vitro. Mycol 95: 584-9. DOI: https://doi.org/10.1080/15572536.2004.11833062 Zamora-Martínez MC, de Pascual-Pola CN. 1995. Natural production of wild edible mushrooms in the Southwestern Rural Territory of Mexico City, Mexico. For Eco Manag 72: 13-20. DOI: https://doi.org/10.1016/0378-1127(94)03450-B Zervakis G, Venturella G. 2002. Mushroom breeding and cultivation enhances ex-situ conservation of Mediterranean Pleurotus taxa. In: Managing plant genetic diversity. Proceedings of An International Conference, Kuala Lumpur, Malaysia, 12-16 June 2000. p. 351-8. DOI: https://doi.org/10.1079/ 9780851995229.0351 Zervakis GI, Venturella G. 2007. Adverse effects of human activities on the diversity of macrofungi in forest ecosystems. Bocconea 21:77-84.ISSN 1120- 4060 (printed) ISSN 2280-3882 (online) Zhang Q, Zak J. 1998. Potential physiological activities of fungi and bacteria in relation to plant litter decomposition along a gap size gradient in a natural subtropical forest. Microb 35: 172-9. DOI: https://doi.org/10.1007/s002489900071 Zhang T, Wang NF, Liu HY, Zhang YQ, Yu LY. 2016. Soil pH is a key determinant of soil fungal community composition in the ny-ålesund region, Svalbard (high arctic). Front Microbiol 7: 227. DOI: https://doi.org/10.3389/fmicb.2016.00227 Zobel DB, Jha PK, Behan MJ, Yadav UKR. 1987. A practical manual for ecology. Kathmandu (NP): Ratna Book Distributors.