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Microscopic fungi from the Aspergillus and Penicillium genera isolated from the rock surfaces of Dem? anovsk a Ice Cave and Dem? anovsk a Cave of Liberty, sevenday-old at 25 ? 1 C, top and bottom views of a colony on the PDA medium (1), CYA (2), MEA (3), and Czapek-Dox Agar (4): A-A. elegans, B-A. flavus, C-A. niger, DP. brevicompactum, E-P. crustosum.
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This paper is the first geomycological report regarding the fungal communities on rock surfaces in the Demänovská Ice Cave and the Demänovská Cave of Liberty, Slovakia. The samples were collected in June 2014 from five locations inside both caves by using sterile swabs wetted with physiological saline (0.85% NaCl). The density of epilithic fungi is...
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... morphological analysis of the sampled fungi was performed using macro-and microscopic observations of the colonies that had grown on the culture media (the morphology of hyphae, conidia and sporangia). For this purpose, plates were incubated in plastic boxes for 7 or 21 days in the dark at 25 § 1 C (Figures 2-4). The fungi were identified using diagnostic keys (Pitt 1979;Golubev 1981;Tanaka et al. 2001;Mukhin and Votintseva 2002;Frisvad and Samson 2004;Chilvers and Toit 2006;De Hoog et al. 2006;Lim et al. 2010;Krzysiak et al. 2011;Bensch et al. 2012;Woudenberg et al. 2013;Jung et al. 2014;Visagie et al. 2014;Kim et al. 2016). ...
Citations
... Fungi exhibit polymorphism, with mycelium in hyphae and unicellular forms, as well as intermediate forms as yeast-like fungi, reproducing via numerous spores and thriving on mineral, oligotrophic substrates by absorbing airborne nutrients (Ogórek, 2018a(Ogórek, , 2018b. Over 1150 species across 550 genera have been identified in underground environments, including cosmopolitan, saprotrophic, and opportunistic pathogenic species (Nováková, 2009;Vanderwolf et al., 2013a). ...
... Our previous speleomycological research of Demänovská Slobody Cave revealed higher species biodiversity of fungi inside the cave, including those on rock surfaces (Ogórek, 2018a), in the air (Ogórek, Fig. 2. The average number (CFU‧10 2 ± SD per 1 g) of fungi (A) cultured at 5, 24 and 37 o C from individual study sites of the Demänovská Slobody Cave (Ilocation outside the caves, from II to IXlocations inside the caves), and overall as the sum of all incubation temperatures as well as percentage of each fungal species contributing to the total fungi cultured from the samples collected outside (B) and inside (C) of the cave for all incubation temperatures. 2018b), especially species of keratinolytic and keratinophilic fungi compared to the external environment. ...
... Furthermore, Zhang et al. (2021) conducted a study on the mycobiota of 13 caves in China, revealing that the highest number of fungal strains were isolated from soil, surpassing those from rock, water, and organic litters. This is corroborated by both our earlier research (Ogórek, 2018a(Ogórek, , 2018bOgórek et al., 2022) and our ongoing investigations, highlighting that, among all the elements studied within the Demänovská Slobody Cave, the soil exhibited the highest diversity of fungal species. Our study aligning with recent research that highlights the critical role of microbial biochemical mechanisms in shaping underground soil ecosystems . ...
Our study unveils the complex interplay between fungal diversity and geochemical environments within the Demänovská Slobody Cave, marking the first comprehensive speleomycological and chemical investigation of its kind. Through meticulous analysis using both microscopic techniques and ITS region gene barcoding, we recorded 17 fungal species, including four previously unrecorded in subterranean settings (Cladosporium ramo-tenellum, Porostereum spadiceum, Rhexocercosporidium carotae, and Trichocladium nigrospermum), with Porostereum spadiceum as the most prevalent. These findings illuminate the crucial role of fungi in subterranean biogeo-chemical cycles, demonstrating how nutrient levels and pH significantly influence fungal populations. Namely, the levels of mineral nutrients (N, P, K, Mg, Ca) and organic matter in sediments within the cave were lower than in soil those outside it. Conversely, the pH values of the tested samples inside the cave were higher compared to those outside. Positive correlations were observed between the average number of fungi and the content of N, P, K, CO, and organic matter (p < 0.05). Conversely, the concentration of fungi decreased with increasing levels of Mg, Ca, and pH (p < 0.05). This research not only expands our understanding of cave ecosystems but also sets a new precedent for exploring the ecological impact of fungi in extreme environments.
... Numerous speleomycological studies conducted in recent years have indicated differences in fungal communities depending on the geographic location of the tested object, the location of the study within a given object, the season of the year, the external environment, and the presence of visitors and animals living in them. The presence of animals and the different seasons are among the most important determinants of the occurrence of fungi in underground ecosystems [14,26,[66][67][68][69]. This was partly confirmed by our previous study in the Brestovská Cave, which investigated the occurrence of psychrophilic and psychrotolerant airborne fungi in air samples [1] as well as the presence of keratinophilic and keratinolytic fungi [7]. ...
... The genus is commonly found in soil, especially in the rhizosphere [72] where it is usually beneficial for plants. Underground ecosystems are also enriched with other Penicillium spp., such as P. chrysogenum, P. camemberti, and P. commune, which are common in European caves [7,24,68,73], but they are also isolated in other parts of the world, e.g., P. chrysogenum was identified in cave soil samples in Russia, Korea, and Thailand, P. camemberti in Australia, and P. commune in northeastern USA [14]. Penicillium chrysogenum was isolated in our previous aeromycological studies of the Brestovská Cave from the air inside and outside of the cave [1]. ...
Soil and sediment host microorganisms are able to survive in extremely resource-limited environments. Therefore, more and more attention is being paid to cave sediments as a reservoir of microbiota. The aim of this study is the speleomycological evaluation of the culturable soil and sediment fungal communities in the Brestovská Cave. To explore the origins of fungi, speleomycological studies were conducted both inside and outside the cave under investigation. Additionally, two incubation temperatures (5 and 24 °C) were used to increase the species spectrum of isolated fungi. To achieve the most accurate species identification, we combined an assessment of morphological characteristics of the isolates with molecular sequencing (ITS, internal transcribed spacer). Twenty different species were found and the most frequent was Penicillium commune, fol-lowed by Trichosporiella cerebriformis and Pseudogymnoascus pannorum. To our knowledge, our study has enabled the first identification of fungal species such as Penicillium swiecicki, Cephalotrichum hinnuleum, Cosmpospora berkeleyana, Lecythophora hoffmannii, Ambomucor seriatoinflatus, and Mortierella minutissima in underground sites. Our data showed that the abundance and composition of the fungal community varied between the indoor and outdoor samples and thus from the entrance and less visited sites deeper in the cave.
... Opisthokonta is a taxonomic supergroup that includes various eukaryotic organisms, like fungi, animals, and some protists [71]. Many species of opisthokonts are found in caves [30,69], where they can exhibit important ecological roles as decomposers [72], predators [73], and symbionts [74]. Overall, Opisthokonta are significant contributors to the biodiversity and ecological intricacy of cave ecosystems. ...
... Fungi are particularly abundant in caves, where they can break down organic matter and recycle nutrients in the nutrient-poor environment. Some species of fungi form mutualistic relationships with other organisms (lichens), such as fungi, algae, or cyanobacteria [74]. Cercozoa are commonly found in soil and aquatic environments [89]. ...
The presence of microbial communities on cave walls and speleothems is an issue that requires attention. Traditional cleaning methods using water, brushes, and steam can spread the infection and cause damage to the cave structures, while chemical agents can lead to the formation of toxic compounds and damage the cave walls. Essential oils have shown promising results in disrupting the cell membrane of bacteria and affecting their membrane permeability. In this study, we identified the microorganisms forming unwanted microbial communities on the walls and speleothems of Petralona Cave using 16S and 18S rDNA amplicon sequencing approaches and evaluated the efficacy of essential oils in reducing the ATP levels of these ecosystems. The samples exhibited a variety of both prokaryotic and eukaryotic microorganisms, including Proteobacteria, Actinobacteria, Bacteroidetes, Chloroflexi, Firmicutes, the SAR supergroup, Opisthokonta, Excavata, Archaeplastida, and Amoebozoa. These phyla are often found in various habitats, including caves, and contribute to the ecological intricacy of cave ecosystems. In terms of the order and genu taxonomy, the identified biota showed abundances that varied significantly among the samples. Functional predictions were also conducted to estimate the differences in expressed genes among the samples. Oregano EO was found to reduce ATP levels by 87% and 46% for black and green spots, respectively. Consecutive spraying with cinnamon EO further reduced ATP levels, with reductions of 89% for black and 88% for green spots. The application of a mixture solution caused a significant reduction of up to 96% in ATP levels of both areas. Our results indicate that essential oils could be a promising solution for the treatment of microbial communities on cave walls and speleothems.
... Peter and Paul are in accordance with other mycological studies performed in caves and hypogean environments, with similar humidity and temperature conditions, indicating that these fungi have a particular affinity for environmental parameters prevailing in this type of cultural heritage site. The majority of documented culturable fungi have already been observed on these types of substrata (fresco, wood, and cave stone walls) in similar studies [55][56][57][58][59][60][61][62]. Several fungal species were isolated for the first time (e.g., Alternaria abundans, Penicillium freii, and P. samnosnianum from fresco; Aspergillus aureolatus and Blastobotrys niveus from iconostasis; As. pseudoglaucus, Beauveria pseudobassiana, Fusarium sporotrichioides, Mortierella alpina, and P. expansum from cave stone); however, fungi from these genera are known to colonize or be deposited on the surface of these cultural heritage substrata. ...
The mycobiome of the cave Church of Sts. Peter and Paul, housing the peculiar fresco painting of “The Bald-headed Jesus”, was analyzed via culture-dependent and -independent methods. Salt efflorescence, colored patinas, and biofilm, as well as biopitting, discolorations, and fruiting bodies of wood-decay fungi were observed on surfaces within the church. Microscopic analyses showed an abundance of fungal structures, i.e., conidiophores, conidia, chlamydospores, and ascospores. The estimated values of the contamination classified all surfaces as the “Danger zone”. A total of 24 fungi from 17 genera were determined as part of the culturable mycobiome, with a dominance of Ascomycota of genera Penicillium. Biodegradative profiles analyzed via plate assays demonstrated positive reactions for 16 isolates: most commonly acid production (8), followed by pigment production and ligninolytic activity (6), protein degradation (5), cellulolytic activity (3) and carbonate dissolution (2). Metabarcoding analysis showed a dominance of Ascomycota in all samples (79.9–99.7%), with high relative abundance documented for Hypoxylon fuscopurpureum on the iconostasis and unclassified Mycosphaerellaceae family within order Capnodiales on fresco and stone, as well as moderate relative abundance for unclassified Dothideomycetes, Botryolepraria lesdainii, Verrucaria sp. and Cladosporium sp. on stone walls. The used set of integrative methods pointed out species of genus Neodevriesia and H. fuscopurpureum as the main deteriogenic agents of fresco and iconostasis surfaces, respectively.
... Therefore, Slovak caves have been one of the great examples of fungal biodiversity studies due to their being important underground localities for Myotis spp. bats [27] and open tourist attractions [16,28,29]. ...
... During most studies of cave mycobiota in Slovak underground environments, the main focus has been on the diversity of microscopic fungi in cave air [16,27,28,30], rock surfaces [26,29] or bat guano [27]. However, from the evaluation of soil and sediment samples taken in and outside of the Harmanecká Cave, we have recently isolated a dermatophyte belonging to the Microsporum cookei clade with close affinities to Paraphyton cookei [31]. ...
Despite speleomycological research going back to the 1960s, the biodiversity of many specific groups of micromycetes in underground sites still remains unknown, including keratinolytic and keratinophilic fungi. These fungi are a frequent cause of infections in humans and animals. Since subterranean ecosystems are inhabited by various animals and are a great tourist attraction, the goal of our research was to provide the first report of keratinophilic and keratinolytic fungal species isolated from three caves in Tatra Mts., Slovakia (Brestovská, Demanovská Lǎdová and Demanovská Slobody). Speleomycological investigation was carried out inside and outside the explored caves by combining culture-based techniques with genetic and phenotypic identifications. A total of 67 fungal isolates were isolated from 24 samples of soil and sediment using Van-breuseghem hair bait and identified as 18 different fungal species. The study sites located inside the studied caves displayed much more fungal species (17 species) than outside the underground (3 species), and the highest values of the Shannon diversity index of keratinophilic and keratinolytic fungi were noted for the study sites inside the Demänovská Slobody Cave. Overall, Arthroderma quadrifidum was the most common fungal species in all soil and/or sediment samples. To the best of our knowledge, our research has allowed for the first detection of fungal species such as Arthroderma eboreum, Arthroderma insingulare, Chrysosporium europae, Chrysosporium siglerae, Keratinophyton wagneri, and Penicillium charlesii in underground sites. We also showed that the temperature of soil and sediments was negatively correlated with the number of isolated keratinophilic and keratinolytic fungal species in the investigated caves.
... They mentioned that the occurrence of black stains most likely results from spatially localized conditions promoting particular development or physiology of pigmented microorganisms. Other authors have also reported the presence of stain-forming fungi of Ascomycota (Dothideomycetes, Eurotiomycetes, Sordariomycetes) on black or green wall stains [21,26,27], while cave fungi in general can produce all types of pigments (grey, orange, purple, red, white, yellow) [27][28][29]. Additionally, green, brown or black spots have been attributed to the presence of unwanted photosynthetic microorganisms (Lampenflora) of which the growth has been promoted due to the introduction of lighting into the cave ecosystem for visiting reasons [13]. ...
... They mentioned that the occurrence of black stains most likely results from spatially localized conditions promoting particular development or physiology of pigmented microorganisms. Other authors have also reported the presence of stain-forming fungi of Ascomycota (Dothideomycetes, Eurotiomycetes, Sordariomycetes) on black or green wall stains [21,26,27], while cave fungi in general can produce all types of pigments (grey, orange, purple, red, white, yellow) [27][28][29]. Additionally, green, brown or black spots have been attributed to the presence of unwanted photosynthetic microorganisms (Lampenflora) of which the growth has been promoted due to the introduction of lighting into the cave ecosystem for visiting reasons [13]. ...
... However, besides the aesthetic appearance, bacteria may cause damage to prehistoric paintings by altering the mineral structure of the rock, since it was found able to produce inorganic and organic acids, neutral and acidic sugars that may degrade rock, speleothems and prehistoric paintings [13,30]. In general, inorganic acids are exclusively produced by bacteria, with some Firmicutes (Bacillus) being involved in calcite precipitation in caves which can induce damage in parietal markings (such as prehistoric human markings) [9,27,31]. The mineralization and precipitation of crystals by bacteria is initialized by locally changing the microenvironmental conditions (e.g., by increasing alkalinity supersaturation of CaCO3 in aqueous phase). ...
The present study concerns the serious issue of biodeterioration of the caves belonging to natural and cultural heritage sites due to the development of various microorganisms. Thus, a series of 18 essential oils (EOs) extracted from various Greek plants were evaluated in vitro (concentrations of 0.1, 0.2, 0.5, 1.0 and 5.0% v/v) against 35 bacterial and 31 fungi isolates (isolated from a Greek cave) and the antimicrobial activity was evident through the changes in optical density of microbial suspensions. In continuance, eight (8) representative bacterial and fungal isolates were further used to evaluate the minimum inhibitory concentration (MIC) and non-inhibitory concentration (NIC) values of the most effective EOs. According to the results, two EOs of Origanum vulgare were the most effective by inhibiting the growth of all the tested microorganisms at 0.1% (v/v), followed by that of Satureja thymbra which inhibited all bacterial isolates at 0.1% (v/v) and fungal isolates at 0.1, 0.2 and 0.5% (v/v) (depending on the isolate). The MIC ranged between 0.015-0.157 and 0.013-0.156 (v/v) for the bacterial and fungal isolates respectively, depending on the case. The current study demonstrated that conventional biocides may be replaced by herbal biocides with significant prospects for commercial exploitation.
... Fungi play important roles in cave ecosystems via biomineralization, decomposition, and by providing a food source for cave fauna (Northup & Lavoie 2001, Barton & Northup 2007, Nováková 2009. Fungi also secrete organic acids and enzymes involved in the biodegradation and remodeling of caves (Mehta et al. 1979, Ogórek 2018. ...
In this paper we describe a new species of cave fungus belonging to Sporocadaceae (Amphisphaeriales), collected from Gem Cave, Fumin County, Yunnan Province, China. Initial morphological observations confirmed that our fungal collection is a pestalotioid species. Phylogenetic analyses of combined internal transcribed spacer (ITS), β-tubulin (TUB) and translation elongation factor 1-alpha (TEF1α) gene sequence dataset confirmed that our fungus forms an independent branch within Neopestalotiopsis. Thus, we describe our fungus as a new species of Neopestalotiopsis based on both morphology and multigene phylogeny. This is the first-ever report of Neopestalotiopsis from a cave habitat. A full description, micrographs and a phylogenetic tree showing the placement of the new species are provided.
... A variety of typical soil fungi occupying different habitats and soil types (Domsch et al., 2007, and literature therein) have been isolated from the Tabernas profiles. Some of these fungi were found to penetrate the soil to a depth of 50− 60 cm and were recorded in specific underground environments such as caves and mines (Ogorek et al., 2017;Ogorek, 2018). Many of dominant and abundant species in the studied profiles are adapted to survive under extreme abiotic conditionshigh UV and solar radiation, high temperatures, low water availability, low organic matter content, and limited aeration. ...
We studied the effect of plant canopy on the distribution of microfungi through 0-30-cm soil profiles under canopies of three plant species and in their adjacent interspace at sun-exposed plain and north-facing slope in the Tabernas Desert, Spain. Using the soil dilution plate method, 142 species from 68 genera were isolated. Melanin-containing fungi dominated in the topsoil layers because of high temperatures and dryness. By contrast, light-colored fungi generating a high amount of small thin-walled conidia substantially increased their abundance in the deeper soil layers – thermotolerant Aspergillus spp. in the 1-5-cm layers and mesophilic Penicillium spp. in the 10-20-cm layers. The significant increase in density of microfungal isolates at 1-10 cm-depth could be associated with the deposition of conidia of aspergilli and penicillii caused by shallow water infiltration in the fine-grained soil and their survival at these depths due to more appropriate edaphic conditions (mainly higher water availability and root concentration). Generally, the effect of plant canopy on microfungal communities was only weakly expressed in higher variations of isolate densities and species richness through the profiles, and in reduced abundance of melanized species with protective multicellular spore morphology in the topsoil layers; at the same time, soil depth strongly influenced all community characteristics. The study highlights the remarkable role of edaphic factors (temperature, water regime, and soil texture) in the distribution of microfungi through soil depth in semi-arid areas.
... Alternaria abundans Pusz et al. (2015), Ogórek et al. (2017Ogórek et al. ( , 2018, Ogórek (2018a) Al. alternata Y Y Connell and Staudigel (2013), Vanderwolf et al. (2013), Ogórek et al. (2014aOgórek et al. ( , b, c, 2016b, Man et al. (2015), Pusz et al. (2015Pusz et al. ( , 2017Pusz et al. ( , 2018a, Kokurewicz et al. (2016), Jiang et al. (2017a, b), Zhang et al. (2017), Nováková et al. (2018), Pusz et al. (2018a), Popkova and Mazina (2019) Al. alternariae Ogórek et al. (2013) Al. atra Vanderwolf et al. (2013) Al. botrytis Vanderwolf et al. (2013), Kokurewicz et al. (2016), Pusz et al. (2018a, b) Al. brevicolla Vanderwolf et al. (2013) Al. chartarum Vanderwolf et al. (2013) Al. humicola Vanderwolf et al. (2013) Al. infectoria Connell and Staudigel (2013) Al. longipes Nováková et al. (2018) Al. mali Y Zhang (2019) Al. mouchaccae Vanderwolf et al. (2013) Al. oudemansii Vanderwolf et al. (2013) Al. tenuissima Y Y Vanderwolf et al. (2013), Popović et al. (2015), Pusz et al. (2015), Zhang et al. (2017), Nováková et al. (2018) Alternaria sp. Y Vanderwolf et al. (2013), Martin-Sanchez et al. (2014), Popović et al. (2015), Belyagoubi et al. (2018), Bercea et al. (2018), Nováková et al. (2018), Leplat et al. (2018), Zhang (2019) Amauroascus Amauroascus albicans Vanderwolf et al. (2013) Am. kuehnii Vanderwolf et al. (2013) Amauroascus sp. ...
... gibbosaVanderwolf et al. (2013) Tra. hirsuta YVanderwolf et al. (2013),Man et al. (2015),Ogórek et al. (2017Ogórek et al. ( , 2018a Tra. ochraceaVanderwolf et al. (2013) Tra. ...
Karst caves are characterized by darkness, low temperature, high humidity, and oligotrophic organisms due to its relatively closed and strongly zonal environments. Up to now, 1626 species in 644 genera of fungi have been reported from caves and mines worldwide. In this study, we investigated the culturable mycobiota in karst caves in southwest China. In total, 251 samples from thirteen caves were collected and 2344 fungal strains were isolated using dilution plate method. Preliminary ITS analyses showed that these strains belonged to 610 species in 253 genera. Among these species, 88.0% belonged to Ascomycota, 8.0% Basidiomycota, 1.9% Mortierellomycota, 1.9% Mucoromycota, and 0.2% Glomeromycota. The majority of these species have been previously known from other environments, and some of them are known as mycorrhizal or pathogenic fungi. About 52.8% of these species were discovered for the first time in karst caves. Based on morphological and phylogenetic distinctions, 33 new species were identified and described in this paper. Meanwhile, one new genus of Cordycipitaceae, Gamszarea, and five new combinations are established. This work further demonstrated that Karst caves encompass a high fungal diversity, including a number of previously unknown species.
Taxonomic novelties: New genus: Gamszarea Z.F. Zhang & L. Cai; Novel species: Amphichorda cavernicola, Aspergillus limoniformis, Aspergillus phialiformis, Aspergillus phialosimplex, Auxarthron chinense, Auxarthron guangxiense, Auxarthronopsis globiasca, Auxarthronopsis pedicellaris, Auxarthronopsis pulverea, Auxarthronopsis stercicola, Chrysosporium pallidum, Gamszarea humicola, Gamszarea lunata, Gamszarea microspora, Gymnoascus flavus, Jattaea reniformis, Lecanicillium magnisporum, Microascus collaris, Microascus levis, Microascus sparsimycelialis, Microascus superficialis, Microascus trigonus, Nigrospora globosa, Paracremonium apiculatum, Paracremonium ellipsoideum, Paraphaeosphaeria hydei, Pseudoscopulariopsis asperispora, Setophaeosphaeria microspora, Simplicillium album, Simplicillium humicola, Wardomycopsis dolichi, Wardomycopsis ellipsoconidiophora, Wardomycopsis fusca; New combinations: Gamszarea indonesiaca (Kurihara & Sukarno) Z.F. Zhang & L. Cai, Gamszarea kalimantanensis (Kurihara & Sukarno) Z.F. Zhang & L. Cai, Gamszarea restricta (Hubka, Kubátová, Nonaka, Čmoková & Řehulka) Z.F. Zhang & L. Cai, Gamszarea testudinea (Hubka, Kubátová, Nonaka, Čmoková & Řehulka) Z.F. Zhang & L. Cai, Gamszarea wallacei (H.C. Evans) Z.F. Zhang & L. Cai.
... In the present investigation twenty fiveairborne fungal species were isolated from cave environment. These fungal species were identified such as Aspergillus species, Penicilliumsp, Fusarium sp etc. Aspergillus species are most commonly found in cave environment.Generally, Cladosporium cladosporioides and Aspergillus flavus were the most frequently cultured species from the Demanovska Ice Cave and the Demanovska a Cave of Liberty, respectively 27 .Most microscopic fungi are transferred in the bioaerosol form with air currents to underground sites from the external environment, as evidenced by the fact of isolation of the majority of fungi from the twilight zone 28,29 . On the other hand, the fewest fungi usually occur in the dark zone in which total darkness and constant low temperature prevail 26 . ...
The aim of present study was to isolate and characterize the fungal species from the air samples obtained from the Western Ghats Cave of Tirunelveli district. Airborne fungi occur in indoor cave were assessed during the study period using conventional methods. This was determined through air quality sampling using the open plate techniques. 25 species of sporulating mesophilous fungi were isolated from the air samples collected at the entrance, twilight and dark zones of two caves. Fungal genus were isolated from the air sample such as Aspergillus sp, Penicillium sp,Cladosporium sp,Fusariumsp,Curvularia clavata, Trichoderma sp,Mucor sp,Rhizopus sp,and Alternaria sp.Cave roosts fungi were constant temperature around 21.87 0 C to 28 0 C and humidity 53% to 74.89% were recorded.The role of airborne fungi identified in the study on controlling pathogenic microbes and in turn their impacts on bat roost microclimate were discussed.
