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| Mean and range (95% HPD) divergence time estimations (Mya) of the suspected obligate troglobitic fungi.
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Karst caves are obviously characterized by darkness, constantly low temperature, high humidity, and oligotrophy. Previous studies revealed that Karst caves have a high and specific bio-diversity. A large number of troglobiont animals had been discovered and their evolution and speciation have been well investigated. However, the origin and evolutio...
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... times of the suspected obligate troglobitic fungi were different (Figure 3 and Table 2). The most recent divergence is Metapochonia variabilis, occurred in Miocene (7.2 Mya, 5.3- 9.2 Mya for 95% HPD), and the oldest is Gymnoascus exasperates in late Jurassic (157 Mya, 139-181 Mya for 95% HPD). ...
Citations
... Os estudos no ambiente cavernícola têm revelado uma riqueza e abundância fúngica de mais de 2.000 espécies com registros em todo mundo (Vanderwolf et al., 2013), inclusive no Brasil onde cerca de 125 espécies de fungos foram relatadas em cavernas do país (Alves et al. 2022). Os fungos cavernícolas são considerados cosmopolitas, sendo a maioria oriunda do ambiente externo, e se adaptam à dinâmica ecológica desse ambiente (Zhang et al., 2018). A presença de fungos nas cavernas é diretamente influen- (Lobo & Boggiani, 2013;Pedro & Bononi, 2007). ...
... Studies in the cave environment have revealed a fungal richness and abundance of more than 2,000 species worldwide (Vanderwolf et al., 2013), including in Brazil where around 125 species of fungi have been reported in the country's caves (Alves et al. 2022). Cave fungi are considered cosmopolitan, with the majority coming from the external environment and adapting to the ecological dynamics of this environment (Zhang et al., 2018). Fungi presence in caves is directly influenced by air currents arriving from the external environment (Lobato et al., 2009;Taylor et al., 2014), bats (Cunha et al., 2020;Ferreira et al., 2000a, b;Pereira et al., 2022) or even tourists and researchers (Zhelyazkova et al., 2020). ...
... Semelhante ao nosso estudo, Taylor rwolf et al., 2013;Cunha et al., 2020;Alves et al., 2022). Além disso, foi demonstrado que os fungos cavernícolas podem ser originários do ambiente externo (Zhang et al., 2018) e a visitação turística de cavernas pode auxiliar na entrada de espécies que ainda não estão presentes no ambiente (Zhelyazkova et al., 2020). ...
Fungos desempenham um papel ecológico importante nas cavernas, apesar de ainda serem um tesouro que precisa ser explorado no Brasil. Estudos micológicos em cavernas brasileiras têm revelado uma riqueza de fungos surpreendente e ainda desconhecida. O presente estudo teve como objetivo relatar a riqueza de espécies de Penicillium em caverna do Cerrado, Brasil. Fungos foram isolados do ar e do sedimento de oito pontos da caverna e identificados com base em características morfológicas e de sequências de ITS do rDNA. No total, foram contabilizadas 890 colônias (UFC), sendo 178 (100 = ar e 78 = sedimento) identificadas como Penicillium. Associando as duas ferramentas de identificação, isolados representativos foram identificados em 12 espécies pertencentes a cinco seções (Brevicompacta, Citrina, Lanata-Divaricata, Exilicaulis e Ramosum), sendo a seção Citrina a que apresentou o maior número de espécies (seis), com destaque para P. sumatraense com a maior abundância (ar = 25 e sedimento = 15). Para o nosso conhecimento, seis (50%) espécies estão sendo relatadas pela primeira vez em ambiente cavernícola, sendo a maioria (quatro) da seção Citrina. O relato da riqueza de espécies de Penicillium está entre as “primeiras pistas do mapa do tesouro” escondidas em cavernas brasileiras. O nosso estudo é um exemplo do grande número de espécies de fungos que são encontradas nas cavernas, sugerindo que estudos micológicos do ambiente cavernícola são importantes para inclusão de dados fúngicos no plano de manejo de cavernas com potencial turístico no Brasil.
... Cave ecosystems display unique environmental conditions that are unfavorable for the life and development of most living organisms [1]. They appear as highly zonal environments with characteristics defined by the ground waters, surrounding rocks, and karst morphology [2][3][4]. The conditions of subterranean facilities significantly differ from the surface and include, i.e., a constant temperature through the year (oscillating between 6 to 10 • C), high humidity (often close to saturation) [5], minimal to no light, restricted or absent air flow with the outside environment, oligotrophy, and sometimes elevated levels of CO 2 [6,7]. ...
... Fungi in the cave environments play important roles-mainly as parasites/pathogens, saprotrophs (decomposers), or mutualistic fungi (e.g., mycorrhizae), but also participants in geological processes [4,13,14]. Some fungal species are known to parasitize insects [15][16][17], while others may contribute to the inorganic nutrient pool replenishment by dissolving the bedrock [18]. ...
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.
... Earth's subsurface environments are isolated from phototrophic energy sources, which are characterized by oligotrophic condition and limitation in electron donors or electron acceptors (Jones et al., 2018). Microorganisms living in such environments are highly dependent on the oxidation of limited inorganic matter for energy (Zhang et al., 2018;Dong et al., 2020;Jones and Northup, 2021). Nevertheless, a large number of microbial cells is estimated to be 2 × 10 29 -6 × 10 29 in the terrestrial subsurface biosphere excluding those in soils (Magnabosco et al., 2018). ...
Subsurface karst caves provide unique opportunities to study the deep biosphere, shedding light on microbial contribution to elemental cycling. Although ammonia oxidation driven by both ammonia-oxidizing bacteria (AOB) and ammonia-oxidizing archaea (AOA) is well explored in soil and marine environments, our understanding in the subsurface biosphere still remained limited to date. To address this gap, weathered rock and sediment samples were collected from the Xincuntun Cave in Guilin City, an alkaline karst cave, and subjected to high-throughput sequencing and quantification of bacterial and archaeal amoA , along with determination of the potential nitrification rates (PNR). Results revealed that AOA dominated in ammonia oxidation, contributing 48–100% to the PNR, and AOA amoA gene copies outnumbered AOB by 2 to 6 orders. Nitrososphaera dominated in AOA communities, while Nitrosopira dominated AOB communities. AOA demonstrated significantly larger niche breadth than AOB. The development of AOA communities was influenced by deterministic processes (50.71%), while AOB communities were predominantly influenced by stochastic processes. TOC, NH 4 ⁺ , and Cl ⁻ played crucial roles in shaping the compositions of ammonia oxidizers at the OTU level. Cross-domain co-occurrence networks highlighted the dominance of AOA nodes in the networks and positive associations between AOA and AOB, especially in the inner zone, suggesting collaborative effort to thrive in extreme environments. Their high gene copies, dominance in the interaction with ammonia oxidizing bacteria, expansive niche breadth and substantial contribution to PNR collectively confirmed that AOA better adapted to alkaline, oligotrophic karst caves environments, and thus play a fundamental role in nitrogen cycling in subsurface biosphere.
... Information on the species richness of Aspergillus, Penicillium, and Talaromyces from tropical caves, such as those we have studied in Brazil, has revealed the presence of species mainly restricted to surface environments and with specialised ecological relationships (Cunha et al. 2020, Alves et al. 2022. The data reported here from Brazilian caves may help us to understand the origin of cave fungi (Zhang et al. 2018) and also support the argument that caves are a hotspot of mycological diversity, as is observed for obligate stygobitic and troglobitic species (Silva & Ferreira 2016). Our study revealed 26 species of Aspergillus, Penicillium, and Talaromyces in caves, eight of which were reported for the first time in cave environments (Aspergillus alvaroi sp. ...
The study of the Brazilian cave mycobiota has revealed a rich but highly diverse assemblage of fungi, with Aspergillus, Penicillium, and Talaromyces being the most frequently reported genera. The present study investigated the airborne fungi and fungi obtained from the bodies of bats, guano, and the soil/sediment from the caves Urubu (in the Atlantic Forest) and Furna Feia (in the Caatinga dryland forest) in the Northeast region of Brazil. Fungal strains were identified based on morphological features and multilocus phylogenetic analyses of ITS, beta-tubulin (BenA), calmodulin (CaM), and RNA polymerase II second largest subunit (RPB2) sequences. A total of 86 isolates were obtained, representing Aspergillus (34), Penicillium (20), Talaromyces (2), and 30 isolates belonging to other genera that will be reported on elsewhere. These isolates were identified as 18 Aspergillus, nine Penicillium, and one Talaromyces species. Eight of the species identified are reported for the first time from a cave environment. Four species showed unique morphological features and phylogenetic relationships, and are newly described. These include two new species of Aspergillus (A. alvaroi sp. nov. and A. guanovespertilionum sp. nov.), one of Penicillium (P. cecavii sp. nov.), and one of Talaromyces (T. potiguarorum sp. nov.). Our study increases the awareness and known richness of the Brazilian and global fungal diversity found in caves.
... Fungi are a prominent group of microorganisms found in caves because of their important ecological functions, such as the cycling of organic matter, rock weathering, and symbiosis with other organisms, in addition to serving as food for cavernicolous fauna [4,5,[11][12][13]. The first report of fungi in caves was by Humboldt in 1794 [14] [apud 5,13,15,16]. However, until the mid-1980, studies in these environments were mainly focused on fauna and the discovery of pathogens such as Histoplasma capsulatum, Pseudogymnoascus destructans, and Trichophyton mentagrophytes [5,16]. ...
... The first report of fungi in caves was by Humboldt in 1794 [14] [apud 5,13,15,16]. However, until the mid-1980, studies in these environments were mainly focused on fauna and the discovery of pathogens such as Histoplasma capsulatum, Pseudogymnoascus destructans, and Trichophyton mentagrophytes [5,16]. ...
... A list of 1626 fungal species divided into 644 genera has been documented in underground environments worldwide [4]. However, it is estimated that there may be 50,000-100,000 fungal species in caves [16]. Several studies have reported the discovery of new species in caves [4,5,12,17,18]. ...
Caves are unique environments characterized by spatial limitations, partial or total absence of direct light, and scarcity of organic carbon and nutrients. Caves are shelters for a variety of adapted animals and microorganisms such as fungi, many of which are still unknown. Amphichorda is a fungal genus belonging to the family Bionectriaceae, which includes cave-dwelling and entomopathogenic species with biotechnological applications. In this study, a new fungal species was identified using morphological and multi-locus phylogenetic analyses of the ITS, LSU, and TEF loci, in the Gruta Velha Nova limestone cave located in the Southern Espinhaço Range, Monjolos, Minas Gerais, Brazil. During the exposure of potato dextrose agar plates to the cave environment, an insect from the family Rhaphidophoridae passed by and fed on the culture medium, resulting in three fungal isolates. Phylogenetic analyses showed that these isolates formed a clade distinct from all known species, leading us to introduce a new species, Amphichorda monjolensis, which may be associated with this insect. Here, we also proposed two new combinations for species of acremonium-like fungi in the Bionectriaceae: Bulbithecium globosisporum (synonym: Acremonium globosisporum) and Hapsidospora curva (synonym: Acremonium curvum). The discovery of A. monjolensis highlights the potential of caves as shelters for new species with significant biotechnological importance.
... Caves harbor a diverse array of parasitic and decomposing fungi. More than 1150 fungi species have been described from caves [7]. Furthermore, many novel compounds and bioactive substances were isolated from cave-derived microorganisms [8][9][10][11]. ...
... The racemic-2 (4 mg) was resolved as the optically-pure (+)-2 (1.7 mg, t R 5.9 min) and (− )-2 (1.6 mg, t R 9.5 min) by a FLM chiral NY column (5:95 n-hexane/2-propanol, 0.8 mL/min). Fr.6 (7.2 g) was separated by using VLC eluting with CH 2 Cl 2 /MeOH (50:1-2:1) to afford eight subfractions (Fr.6-1 to Fr. [6][7][8]. Fr.6-6 (0.18 g) was separated on a pre-TLC eluted with CH 2 Cl 2 /CH 3 OH (v/v, 25:1), then Fr.6-6-1 (0.11 g) was purified using a C 18 column (55:45 MeOH/H 2 O, 4 mL/min) to yield 1 (15.7 mg, t R 8.0 min). ...
... Additionally, the influences of external communities, as well as animal and human activities, on fungal communities differ among caves. For instance, Zhang et al. found that fungal communities in Karst caves were greatly influenced by fungal communities in external environments due to airflow exchange [21,45]. Meanwhile, bats and human activities might introduce foreign fungi and change the fungal community in caves [24,25,[46][47][48]. ...
As a unique ecosystem, Karst caves harbor an impressive diversity of specific fungi. However, the factors and mechanisms that shape fungal biodiversity in caves remain elusive. In this study, we explored the assembly patterns of fungal communities based on our previous research in eight representative Karst caves in Southwest China. Our results indicated that dispersal limitation plays a crucial role in shaping the overall fungal community as well as specific communities in rock, sediment, and water samples. However, “Undominated” processes contributed more than dispersal limitation in air samples. Interestingly, the dominant assembly processes varied between caves. Consistently, environmental selection had a minor impact on the assembly of fungal communities. Among the examined spatial and environmental variables, latitude, longitude, altitude, and temperature were found to significantly influence fungal communities irrespective of substrate type. These findings provide valuable insights into the ecological factors governing fungal community assembly in Karst caves.
... Bats play a key role as sources of energy input into these subterranean environments through the deposition of guano (Pimentel et al., 2022), to the point of potentially bioengineering cave systems (Piló et al., 2023). Many organisms in these environments are functionally phylogenetically unique and have a high degree of endemism (Chagas & Bichuette, 2018;Zhang et al., 2018), relying on the injection of energy from the outside environment (Ferreira, 2019). The guano deposits in bat caves have also been linked to increases in the species richness and diversity of fungi (Cunha et al., 2020). ...
Bats frequently inhabit caves and other subterranean habitats and play a critical role in subterranean food webs. With escalating threats to subterranean ecosystems, identifying the most effective measures to protect subterranean‐roosting bats is critical. We conducted a meta‐analysis to evaluate the effectiveness of conservation and management interventions for subterranean‐roosting bats. We used network analyses to determine to what extent interventions for bats overlap those used for other subterranean taxa. We conducted our analyses with data extracted from 345 papers recommending a total of 910 conservation interventions. Gating of roost entrances was applied to preserve bat populations in 21 studies, but its effectiveness was unclear. Habitat restoration and disturbance reduction positively affected bat populations and bat behavior, respectively, in ≤4 studies. Decontamination was assessed in 2 studies and positively affected bat populations, particularly in studies focused on reducing fungal spores associated with white‐nose syndrome in North America. Monitoring of bat populations as an effective conservation strategy was unclear and infrequently tested. Only 4% of bat studies simultaneously considered other subterranean organisms. However, effective interventions for bat conservation had similarities with all other organisms. If other subterranean organisms are considered when applying interventions to conserve bats, they might also benefit.
... Vectors can be plant debris transported by water or wind into caves, airflow transporting particulate material from the epigean environment, and the activity of animals, such as bats, using caves as shelter and depositing feces on the cave floor (Ogorek et al. 2016;Zhang et al. 2017b;Cunha et al. 2020). According to Zhang et al. (2018), new species of fungi found in karst caves in China are unlikely troglobitic organisms but rather travelers from the epigean environment. They have used molecular dating evidence and stated that the geological age of caves is too short for fungal speciation. ...
Interest in cave fungal diversity is flourishing because it may represent a reservoir of new species and metabolites. However, the mycobiota remains poorly studied in the underground environment, especially in neotropical regions. During surveys that aimed to investigate the fungal diversity in quartzite and limestone caves in the Southern Espinhaço Mountain in Brazil, six Chaetomiaceae isolates were obtained from different cave substrates. Five taxonomical novelties of Chaetomiaceae in Brazilian caves were discovered based on phylogenetic analyses using DNA sequences from the ITS, LSU, TUB, and RPB2 genes. Chaetomium meridionalense, Pseudohumicola alba, and Pseudohumicola lutea are new species found in Gruta da Extração and Gruta Velha Nova caves. Parahumicola is introduced as a new genus representing a novel phylogenetic lineage with unique morphological characteristics in the family Chaetomiaceae. This new monotypic genus is typified by P. guana, which was found in a bat guano sample in the Gruta Monte Cristo cave. Furthermore, this is the first report of Collariella bostrychodes in a neotropical cave. Overall, these findings emphasise that Brazilian caves constitute an untapped source of fungal resources.
... Previous studies have suggested that some of the fungal communities in caves were introduced from the outside [11][12][13][14]. They have been reported to be affected by external factors, such as airflow, water movement, and visitors [11,12,15,16]. Among these, cave-inhabiting bats are estimated to be one of the most important factors in the inflow of fungal spores [17,18]. ...
Pseudogymnoascus is a psychrophilic fungus, which is a genus widely distributed in cold regions around the world. Recently, the presence of Pseudogymnoascus destructans (Pd), the causative agent of white-nose syndrome (WNS) belonging to Pseudogymnoascus, has been reported in neighboring countries of Korea. However, no investigation on Pd has been reported in Korea. In this study, cave-inhabiting bats and their habitats were investigated in terms of the diversity of cave fungi, and we tried to confirm the presence of Pd. Three caves suspected of hosting Pd were selected, and 83 environmental and 53 bat samples were collected. A total of 154 fungal strains belonging to 31 different genera were isolated, and 20 of 154 were confirmed to belong to Pseudogymnoascus. Pd-diagnostic PCR was performed to check whether Pd was present in the isolated Pseudogymnoascus, and seven positives were confirmed. However, phylogenetic analyses revealed that no isolates belonged or were closely related to the clade with Pd. Although samples were collected from limited areas, undescribed Pseudogymnoascus species were isolated, and it was confirmed that Korean isolates were distributed in various clades. In conclusion, it is hypothesized that Korean Pseudogymnoascus presents high diversity.
