Chiang Mai Journal of Science

Mini Review: Wetland Associated Fungi in Coastal Freshwater Ecosystems – A Focus on Poales-Dominated Coastal Wetlands in Thailand

1. INTRODUCTION

1.1 Wetlands
     Wetlands are dynamic ecosystems that contain soil, either covered by water or saturated by water. The water level can vary depending on the season, location, and vegetation, as these factors influence soil water availability and the release of nutrients for plant uptake. Water level, quality, and water flow mainly define the inhabitants of these environments [1]. The RAMSAR Convention provides a comprehensive definition, describing wetlands as encompassing marshes, fens, peatlands, and aquatic areas, whether natural or artificial, with water that may be static or flowing, and varying in salinity from freshwater to marine environments [2]. These ecosystems cover approximately 6% of the Earth's total land area and are classified based on water flow patterns, pH, soil characteristics, nutrient availability, and geographic location.
     The RAMSAR Convention recognizes six types of wetlands: Marine, Coastal, Estuarine, Lacustrine, Riverine, and Palustrine (Marshes). Lacustrine wetlands are associated with lakes and are fed with fresh water from a lake and nearby waterways. Riverine wetlands are situated along rivers and the banks of tributaries and are often referred to as floodplains. The water level, nutrient content, and water quality fluctuate with the water level of a nearby river [3]. Marshes show a relatively high-water table, lower peat content than bogs and swamps in riverine environments. Water level in marshes shifts with the season, and vegetation adapts to synchronize with seasonal water table changes [1,2,4]. The coastal freshwater wetlands are ecosystems that comprise a combination of riverine wetlands and marshes, situated in close proximity to the coastline, and are characterized by freshwater conditions [2–4].
     Marine and estuarine wetlands are primarily located along coastal regions and comprise ecosystems such as coastal lagoons, rocky shorelines, mangrove forests, and tidal marshes. [4,5]. These environments span gradients of elevation, from shallow subtidal depths that allow light penetration for photosynthetic organisms at the bottom and benthic plants. Towards the ocean, benthic algae and seagrasses dominate the vegetation but the landward niches are inhabited by organisms that can tolerate adverse environmental conditions such as salinity, water table, and pH fluctuations [6].

1.2 Evolution of Wetland Ecosystems
     Wetland ecosystems host distinct communities of flora and fauna, shaped by interactions between biotic and abiotic factors. Co-evolution between landscape geology and related organisms could be the underlying reason for this close association between wetlands and their resident organisms [7]. Geological and paleoecological evidence suggests that wetlands originated during the mid-Paleozoic era [8].
     The development of wetland types paralleled major events in plant evolution. The origin of vascular plants, such as shrubs in the Emsian age, and large trees in the Eifelian age, which led to the development of marshes and swamps in the mid-Devonian era. They are formed by holding soil particles tightly to develop and enable plant root systems [9]. The late Cretaceous saw the origin of the order Poales. At the end of the Paleogene era, those early monocotyledonous plants diversified into families such as Cyperaceae, Juncaceae, Poaceae, and Typhaceae [10]. The physiology and anatomical features of those families facilitated their colonization of wetland habitats. Towards the end of the Oligocene epoch, marshes and swamps were colonized by grasses and grass-like plants resembling present-day flora [7,8].

1.3 Fungal Colonization of Wetlands
     Fungi may exploit available niches in developing wetlands, which offer nutrient-rich substrates, hosts, and favorable environmental conditions that support their establishment and evolution [11,12]. Fungal spores can reach wetlands through several pathways [13]. Wind disperses lightweight conidia and ascospores across long distances; some have been detected 10 km from their origin, with Fusarium graminearum spores found over 500 m above the Earth’s surface [14,15]. Water also serves as an effective dispersal vector from rivers, tributaries, rain, and ocean tides and transportation of spores into wetland environments. Water protects spores from desiccation, UV radiation, and extreme temperatures [15]. Certain fungi, like Ingoldian species, and marine fungi, have spore morphologies adapted for waterborne dispersal. For instance, Lignicola laevis is widely distributed in marine habitats, likely due to ocean currents aiding its spread [16]. Other taxa related to L. laevis have elaborately branched or appendaged spores, such as Flabellospora verticillata. Thus, fungi from upstream leaf litter or nearby vegetation can travel as propagules and colonize downstream wetlands [17].
     Fungi colonize both living and dead plant material. Wetlands often receive driftwood and herbaceous debris through water currents, supporting fungal thalli or reproductive structures [18]. Dispersal of plant propagules, such as seeds bearing endophytes, mycorrhizal fungi, or pathogens, can introduce fungi from other regions to wetlands [19]. Floating live plants can also transport fungal communities, which may establish in new wetland habitats upon settlement [16]. Animals, including humans, act as fungal dispersal agents. Biodiverse wetlands serve as feeding and breeding grounds for wildlife, and the movement of animals in and out facilitates fungal transfer. Migratory birds, for instance, have been shown to carry fungi like Cladosporium cladosporioides and Alternaria alternata between aquatic environments [20]. Human activities, including agriculture and trade can introduce new organisms to wetlands, some of which may harbor fungi that eventually become established [21].

1.4 Wetlands and Evolution of Fungi
     Fungi evolved alongside plants and wetlands, diversifying as decomposers, symbionts, and pathogens. True fungi with chitinous cell walls originated around 760 million years ago, and molecular clock estimates place the divergence of Ascomycota at 500–650 million years ago [22].
     Aero-aquatic fungi are a significant group in wetlands and aquatic habitats. These saprobes decompose submerged plant litter and produce conidia with adaptations for dispersal through air in freshwater environments. Although polyphyletic, their conidial forms evolved independently across evolutionary distinct lineages in response to aquatic conditions and selection pressures, a case of convergent evolution, as seen in Spirosphaera, Tricladium and Varicosporium [22–26]. Genera like Helicodendron and Pseudaegerita produce both large macroconidia and small microconidia. While microconidia rarely germinate, they are considered the true conidia, whereas macroconidia likely evolved as aquatic propagules [23,24]. The emergence of wetlands created new ecological opportunities, driving fungal adaptations and contributing to the structure and diversity of wetland fungal communities [22,24].

1.5 Wetlands in Thailand
     Thailand, a tropical Asian nation, hosts diverse flora and fauna, with wetlands playing a crucial role in supporting biodiversity and offering economic benefits such as eco-tourism and aquaculture [25]. The country's wetlands, comprising marine wetlands, marshes, estuaries, and mangrove swamps that cover about 36,600 km², or 7.5% of land area [27,28]. Important areas include: Thale Noi Non, Don Hoi Lot, the Mekong River, and coastal zones: Khao Sam Roi Yot and Pran Buri estuary [28]. However, these ecosystems face significant threats from resource overexploitation, including logging, land conversion for shrimp farming, poaching, and unsustainable agriculture, leading to the loss of nearly 2,000 km² of mangroves between 1961 and 1996 [29].
     Preserving and studying Thailand’s wetlands is essential not only for wildlife conservation but also for advancing our understanding of wetland-associated microbial diversity. Focused research on wetland fungi can clarify taxonomic relationships and community structures, contributing to a more comprehensive picture of the region’s microbial ecology.

2. FUNGUS-LIKE TAXA FOUND IN WETLANDS

2.1 Zoosporic Fungi and Chytridiomycota
     Wetlands are ecologically diverse habitats offering a wide range of niches for fungal colonization, supporting nearly all major fungal groups and fungus-like organisms [8]. These include zoosporic fungi (Myxomycota, and Oomycota), as well as higher fungi such as Ascomycota, Glomeromycota, Basidiomycota and Zygomycota which process multicellular organization and lack flagellated propagules [30]. Lichens, though present in wetlands and closely related life form to fungi, are not addressed in this review. [31].
     Zoosporic fungi are characterized by their motile, flagellated zoospores [32]. Chytridiomycota, one of the principal zoosporic fungal phyla along with Oomycota, is considered the oldest lineage of true fungi. Though unicellular, chytrids possess true fungal traits and are globally distributed [33–35]. In wetlands, they function primarily as parasites or saprobes, thriving in challenging conditions and tolerating broad pH ranges [36,37]. Saprobic chytrids have been recorded on decaying plant materials, including submerged wood, algae, and herbaceous tissues [30]. For examples Phlyctochytrium planicorne found to exist on filaments of green algae, and Micromyces sp. reported on shrimp chitin [33].

2.2 Oomycota
     Oomycota resemble true fungi in forming mycelium-like thalli and digesting substrates extracellularly, but they differ significantly in structure and genetics. Their coenocytic hyphae contains diploid nuclei and have cell walls composed of β-glucans and cellulose [38]. Primarily saprophytes and pathogens of plants, animals, and fungi, Oomycota inhabit diverse ecosystems including freshwater, marine, and terrestrial habitats [39]. In Korea, species such as Pythium diclinum, P. heterothallicum, P. inflatum, P. intermedium, and P. oopapillum were isolated from sediments and decaying plant matter in freshwater systems [39]. Studies in the Arctic ocean revealed that Oomycota account for about 6% of the eukaryotic microbial community, with diatom parasites like Olpidiopsis drebesii dominant in saline environments. Additionally, Pythiogeton species have been implicated in the die-back of Phragmites australis [41]. These findings highlight the ecological range and widespread occurrence of Oomycota in aquatic systems, including wetlands.

3. HIGHER FUNGI FOUND IN WETLANDS

3.1 Glomeromycota
     Glomeromycetes are true fungi residing in the rhizosphere, forming endomycorrhizal (Arbuscular Mycorrhizal Fungi: AMF) associations within plant root cells [30,42]. Characterized by aseptate hyphae and large multinucleate spores, they lack observed sexual reproduction and are difficult to culture, making metagenomics the primary approach for study. Although data on AMF in wetlands are limited, their presence underscores their ecological significance [43].
     In Thailand’s coastal freshwater wetlands, Typha species dominate, creating monocultures that influence AMF diversity. Studies on Phragmites australis in wetlands identified Glomus species, especially G. mosseae and G. fasciculatum, as dominant [44]. Coastal wetland conditions, such as flooding and hypoxia, impact AMF community structures. Santillán-Manjarrez et al. [45] reported population decline of AMF under anoxic stress. Meanwhile, Silvani et al. [46] identified 14 AMF species from Glomeraceae and Claroideoglomeraceae in similarly extreme environments using both molecular and morphological approaches. Given the fluctuating hydrology, oxygen limitation, and Poales-dominated vegetation of wetlands in Thailand, AMF in these ecosystems likely exhibit dynamic and stress-adapted population structures, warranting further investigation.

3.2 Yeasts
     Yeasts are a group of unicellular fungi that inhabit various habitats including aquatic ecosystems, including wetlands. Yeast stages are found in both Ascomycota and Basidiomycota. The vegetative cells of most yeasts are obovoid, ellipsoid, or spherical in shape; they are occasionally known to produce pseudohyphae depending on environmental conditions, while Mrakia aquatica forms tetraradiate propagules in nature [46]. Candida albicans, Mrakia aquatica, Papiliotrema laurentii, and Saccharomyces cerevisiae are among the most commonly reported yeast species from freshwater ecosystems, with Saccharomycetales being the most dominant order [25,48–50].
     Yeasts in aquatic ecosystems predominantly propagate asexually via budding, although sexual structures have occasionally been reported in Saccharomyces species [51]. In freshwater environments, yeasts are known to function as decomposers (e.g., Saccharomyces cerevisiae, and Mrakia aquatica), endophytic organisms (e.g., Rhodotorula), and as pathogenic organisms associated with aquatic fauna such as Candida species, Debaryomyces hansenii, and Metschnikowia bicuspidata) [49,52,53]. Although most yeast studies in freshwater environments have focused on sediments and aquatic fauna, in-depth taxonomic investigations of yeasts associated with freshwater wetlands remain limited. Therefore, a polyphasic taxonomic approach is recommended to better understand the yeast diversity in these habitats.

3.3 Ascomycota
     Ascomycota is the most abundant and diverse fungal phylum that showcase cosmopolitan distribution across ecosystems. A large proportion of fungi in wetlands are occupied by this group, and function as saprobes, endophytes, and pathogens in these environments. They are well-studied due to their ecological significance and their reproductive strategies including producing either asci-bearing sexual morphs or conidial asexual morphs, and the contribution to maintain ecosystem balance [25,30].
     Four major Ascomycota classes are commonly associated with plants and plant-derived substrates in aquatic habitats include Dothideomycetes, Eurotiomycetes, Leotiomycetes, and Sordariomycetes [49]. Dothideomycetes, the largest class, are defined by their diverse ascomata and bitunicate asci with fissitunicate dehiscence. Sordariomycetes, the second largest, are characterized by perithecial ascomata and inoperculate unitunicate asci. Eurotiomycetes are mostly represented by asexual forms, with cleistothecial ascomata and primitive asci reported in their sexual morphs. Leotiomycetes produce apothecial or cleistothecial ascomata. These classes are well represented in aquatic environments, including wetlands, and are frequently associated with Poales vegetation (Figure 1) [50,54–57].

     Freshwater fungi, defined as “all ascomycetes isolated from submerged or partially submerged substrates in aquatic environments” [58], are well represented in wetland ecosystems shaped by hydrological factors such as water source, volume, and flow dynamics [1]. Most ascomycetes in freshwater wetlands fall within this functional group and exhibit key evolutionary adaptations for aquatic life.
     Aquatic Dothideomycetes and Sordariomycetes develop spores with specialized features, including thick-walls (e.g., Hongkongmyces), appendages (e.g., Tetraploa, Menisporopsis), gelatinous sheaths (e.g., Phaeonectriella appendiculata), and hydrophobic surfaces (Annulatascus), that enhance survival, dispersal, and substrate attachment under submerged conditions (Figure 2) [25,59–62]. Additionally, genera such as Lophiostoma and Massarina demonstrate slow growth, spore dormancy, and delayed germination to endure nutrient-poor or fluctuating environments [63,64]. These traits enable freshwater fungi to persist in dynamic wetland systems and distinguish them from many terrestrial relatives.

3.4 Basidiomycota
     Basidiomycota, the second largest fungal phylum, is distinguished by well-developed fruiting bodies and specialized spore-producing hyphae. Most members lack asexual morphs and are commonly recognized as wood-decaying fungi. In freshwater wetlands, they function as saprobes on decomposing wood, typically in drier microhabitats [30]. Calabon et al. [50] documented 218 Basidiomycota species across approximately 100 genera in freshwater ecosystems, with Agaricomycetes and Ustilaginomycetes being the most prevalent classes. As an example, Tangthirasunun et al. [82] documented Chaetospermum artocarpi (MFLUCC 21-0536) from Phang Nga: Khao Lak coast of Thailand. Similar to Ascomycota, in-depth research on wetland-associated Basidiomycota remains limited, indicating a need for further studies to elucidate their ecological roles in tropical coastal freshwater wetlands.

4. ECOLOGICAL ROLES Of WETLAND ASSOCIATED FUNGI

     The contribution of fungi can be observed in various aspects of wetland ecosystems, where they play a crucial role in maintaining the functionality of these intricate and complex environments [25,30]. As decomposers, fungi contribute to nutrient cycling, while symbiotic fungi, including endophytes, ectomycorrhizal, and endomycorrhizal species support the plant physiology and health. Additionally, pathogenic fungi affecting on both plants and wetland-dwelling animals significantly influences the longevity and balance of these ecosystems [25,30,83,84].

4.1 Nutrient Recycling
     Nutrient cycling in wetlands is largely driven by saprobic fungi that decompose dead vegetation, including submerged, moist, or dry plant materials [25]. Woody debris, particularly abundant in tropical wetlands, supports fungi capable of all three wood decay types including white rot, brown rot, and soft rot, that depending on enzymatic capacity of each fungal group [30,85,86]. White rot fungi degrade both cellulose and mainly lignin, while brown and soft rot primarily target cellulose, leaving lignin intact [86]. Soft rot typically occurs in moist wood under aquatic conditions, producing dense dark cores that are later colonized by white or brown rot fungi in a succession process [88,89].
     During the soft rot process, the responsible fungi secrete extracellular enzymes such as endoglucanases, xylanases, and pectinases, which primarily degrade cellulose and hemicellulose compounds while leaving lignin largely intact [90]. This enzymatic activity results in cavity formation or erosion channels in woody substrates. However, in herbaceous materials, soft rot fungi are capable of rapidly utilizing the majority of the substrate [50,84,91–93]. Aquatic fungi commonly exhibit characteristics of soft rot fungi, as they predominantly colonize moist and herbaceous plant material. For example, Lunulospora curvula, Longipedicellata aptrootii, Pseudoastrosphaeriella africana, and Halobyssothecium unicellulare are known to inhabit Poales host materials in aquatic environments [71,77,94]. In such ecosystems, soft rot fungi play a critical role by making nutrients trapped in substrates rapidly available to other consumers, such as aquatic invertebrates like shrimps, thereby supporting ecosystem balance and longevity [50,84,95].
     Thailand’s coastal freshwater wetlands are dominated by Poales belonging to Cyperaceae, Poaceae, and Typhaceae. These monocotyledonous plants are generally relatively soft, and fast-decomposing herbaceous substrates that support fungal communities distinct from those on wood [30]. Freshwater discomycetes, for example members of Hymenoscyphus, and Lachnum, are frequently isolated from decomposing grasses and sedges [84]. Larger Poales leaves can trap aero-aquatic fungal spores via their bristled surfaces even before senescence. The location where leaf litter accumulates influences fungal colonization, with submerged litter decomposed mainly by aquatic fungi such as Ingoldian hyphomycetes (including Halosphaeriaceae and Pleosporaceae), which are predominantly associated with herbaceous substrates as non-lignin decomposers, whereas litter in drier habitats is primarily colonized by soil-associated Ascomycota and Basidiomycota [96,97]. These fungi involved in decomposition recycle essential nutrients such as carbon, phosphorus, potassium, and magnesium, thereby promoting wetland stability and longevity [30,98].
     Fungi also form symbiotic associations with wetland plants, particularly as endophytes. These fungi, including Trichoderma, Cladosporium, and various Fusarium species, colonize healthy plant tissues without causing physiological harm, offering benefits such as stress resistance. For instance, Cladosporium tenuissimum has been shown to enhance drought tolerance in the wetland species Alternanthera philoxeroides [99,100].

4.2 Endophytic and Mycorrhizal Relationships
     Wetland-associated endophytes disperse in the ecosystem by two main pathways. They may shift from a symbiotic to a saprobic or pathogenic lifestyle depending on host condition, producing sexual or asexual propagules for dispersal [25,87,101]. Alternatively, endophytes can be transmitted through seeds and vegetative tissues. Ingoldian hyphomycetes such as Lunulospora curvula and Tricladium splendens are known to spread via host tissues such as floating leaves, twigs, seeds, and fruits [102]. In Thailand’s coastal freshwater wetlands, seed-mediated dispersal is especially significant, as lightweight monocot seeds carrying endophytes and are easily transported by wind and water [93,100].
     Mycorrhizal fungi represent another vital symbiotic group in wetlands. They are broadly classified as ectomycorrhizal (mostly basidiomycetes) and endomycorrhizal (primarily Glomeromycota). Ectomycorrhizal fungi form external sheaths around roots without penetrating the cortex, while endomycorrhizal fungi colonize root cortical cells without causing damage. These mutualistic interactions enhance plant nutrient uptake, particularly phosphorus, in exchange for host-derived carbohydrates [42]. Although mycorrhizal associations occur in wetlands, waterlogged conditions limit ectomycorrhizal colonization, confining it to drier microhabitats or the dry season [43]. Endomycorrhizal fungi, including Glomus, Funneliformis, and Rhizophagus, have been widely reported in wetland plants, especially non-monocotyledonous hosts. Despite oxygen limitations in flowing-water wetlands, endomycorrhizal fungi remain ecologically important for supporting monocot plants in such environments [103,104].

4.3 Fungal Pathogens in Wetlands
     Fungal pathogens play an important role in ecosystems by regulating natural populations and exerting selective pressures on both plants and animals. Wetland ecosystems are no exception to fungal pathogens. These fungi infect healthy organisms and induce adverse physiological conditions that can be detrimental to the host’s health [105]. Although studies on fungal pathogens in wetland ecosystems are limited, numerous reports document pathogenic genera associated with members of the order Poales [106]. Ascomycota members, including Colletotrichum, Nigrospora, Pestalotiopsis, and Lasiodiplodia, have been isolated from grasses, while rust pathogens have been frequently reported from grasses and other related host plants [25,97].
    Other inhabitants of wetlands, such as animals, are affected by fungal pathogens in these ecosystems. For example, Candida species are known to infect fish and disperse through freshwater bodies by using them as vectors. Veronaea botryosa has been reported in freshwater-associated amphibians in North America and has been documented at pandemic levels [25,107]. Accordingly, wetland-associated fungi, including saprobes, endophytes, mycorrhizal fungi, and fungal pathogens, play a crucial role in maintaining ecosystem balance in these environments.

5. INDUSTRIAL AND ECOLOGICAL UTILIZATION OF WETLAND FUNGI

5.1 Wastewater Treatment and Constructed Wetlands
     Wetland ecosystems are defined by their close association with water, which shapes both their structure and resident biota. Consequently, many fungi inhabiting wetlands are aquatic or possess aquatic adaptations [25]. These features make them particularly suitable for applications in bioremediation and wastewater treatment, alongside their host plants [108]. Bioremediation involves the use of living organisms, primarily fungi and bacteria to break down hazardous substances into less harmful forms [108,109]. Early fungal applications in wastewater treatment focused on decolorizing textile effluents. Candida tropicalis was found effective in degrading dyes under acidic conditions [110], highlighting the potential of fungi from tannin-rich or blackwater wetlands for water purification [37]. Fungi, as saprobes, secrete extracellular enzymes like laccases that oxidize aromatic compounds. Notable laccase producers include Basidiomycota such as members in genus Coprinus and Trametes, and ascomycetes like Melanocarpus albomyces, species well-adapted to aquatic and wetland habitats. Their ability to live in wetland or aquatic ecosystems makes these organisms suitable for use in bioremediation processes [111].
     Aquatic hyphomycetes rapidly colonize leaf litter and other herbaceous materials in freshwater ecosystems such as wetlands. As previously noted, the decomposition activity of these fungi contributes to maintaining water quality and ensures that nutrients are readily available for higher trophic levels in the aquatic food web [108]. For example, Mirabile et al. [112] reported that aquatic hyphomycetes play a key role in leaf litter decomposition in Italian freshwater ecosystems, not only removing organic impurities from the water but also positively correlating with increased populations of macrozoobenthic grazers. This heterogeneous group of benthic invertebrates serves as an important food source for fish populations in freshwater habitats, thereby facilitating stable and balanced fish communities. These findings highlight the direct and indirect ecological importance of aquatic fungi in maintaining ecosystem health in wetlands [32].
     The natural purification capabilities of wetlands have inspired the development of constructed wetlands for wastewater treatment, where fungi native to such ecosystems contribute to pollutant removal [113]. Among these, Arbuscular Mycorrhizal Fungi (AMF) play a key role. AMF colonization in constructed wetlands can occur through plant propagules, water, wind, animals, or artificial introduction, often requiring minimal intervention [114]. Stable AMF-host associations are critical for bioremediation efficacy. Most host plants in constructed wetlands, including Carex, Typha, and other Poales, support AMF colonization. Phosphorus availability influences these interactions, while low phosphorus limits AMF colonization [115], phosphorus-rich wastewater facilitates it. Intern, AMF and other wetland-adapted fungi are integral to the success of constructed wetlands, underscoring their value as biological agents in environmental remediation.

5.2 Wetland Fungi as Bioindicators
     Aquatic hyphomycetes, common in wetland ecosystems, are highly sensitive to environmental factors, making them effective bioindicators of ecosystem health [116]. Their populations respond markedly to changes in water quality, pollutants, and temperature fluctuations [25,56,116]. A redundancy analysis (RDA) by Solé et al. [117] demonstrated that species such as Anguillospora longissima, Clavatospora longibrachiata, Clavariopsis aquatica, Flagellospora curvula, Heliscus lugdunensis, Tumularia aquatica, and Lemonniera aquatica experienced significant population declines in freshwater rivers contaminated with organic matter, heavy metals, sulfates, and nitrates. These species' absence or reduction serves as a clear indicator of freshwater pollution.
     Bai et al. [118] analyzed fungal communities in three polluted freshwater reservoirs using ITS sequencing and chemical profiling. Ecosystems with anthropogenic disturbances and high fertilizer residues showed a notable decline in fungal diversity and abundance, although Schizosaccharomyces showed increased relative abundance. These findings, supported by recent studies including Siriarchawatana et al. [119], emphasize the role of aquatic fungi, including those in freshwater wetlands as reliable bioindicators for monitoring ecological integrity and pollution levels.

6. THREATS, CHALLENGERS And MODERN CONSERVATION For FUNGAL FAUNA In COASTAL FRESHWATER WETLANDS

6.1 Threats and Challengers
     Wetland ecosystems are vital for environmental sustainability and the well-being of adjacent communities. However, expanding human settlements and unsustainable land use have placed these ecosystems under increasing pressure. Anthropogenic drivers, including climate change, invasive species, and habitat disturbance are negatively impacting wetland biodiversity globally [120]. Targeted conservation strategies are essential to protect wetland habitats and their inhabitants, including microfungi [25].
     In tropical countries such as Thailand, improper agricultural expansion poses one of the greatest threats to freshwater wetlands. Wetland water is frequently diverted for crop irrigation, reducing water levels and altering hydrological balance. Land clearing for cultivation leads to biodiversity loss, particularly among wetland flora and fauna [121,122]. As a result, fungi that depend on these plants and animals, whether as pathogens, endophytes, symbionts, or saprobes are also adversely affected [120]. Intensive agriculture further introduces chemical pollutants into wetlands through the use of fertilizers, pesticides, and fungicides. These substances often enter wetlands from surface runoff, disrupting nutrient dynamics and causing eutrophication [123,124]. Fungicide residues, in particular, can disturb aquatic fungal communities, diminishing diversity and ecological function [125].
     The introduction of invasive plant species also contributes to changes in fungal community structures. Non-native species can outcompete native flora, indirectly displacing specialized fungi adapted to native hosts. In contrast, fungi capable of utilizing invasive plant material may proliferate, potentially leading to long-term dominance and loss of fungal diversity [25,120,126]. For instance, in Thailand’s coastal wetlands, invasive Poales species outcompete native Nymphaeaceae, shifting fungal associations and community dynamics.
     Climate change is an overarching threat, with rising global temperatures affecting wetland hydrology and fungal physiology. Aquatic fungi are particularly sensitive to thermal stress, and elevated temperatures can reduce their diversity and metabolic function [25,127]. Furthermore, polar ice melt is expected to alter oceanic and riverine flow patterns, disrupting nutrient delivery and water distribution in wetland systems [120,128]. These compounded impacts pose significant risks to fungal diversity and ecological stability in freshwater coastal wetlands.

6.2 Conservation
     The conservation of wetland-dwelling fungi is best achieved through the protection and sustainable management of their habitats. The current global framework allows for coordinated responses to environmental challenges, including the preservation of coastal freshwater wetlands, critical ecosystems that support fungal biodiversity [129]. International organizations such as the Ramsar Convention, IUCN, and WWF actively contribute to wetland conservation, indirectly safeguarding fungal communities [120,130]. National governments, particularly in countries like Thailand, can play a pivotal role by formulating conservation policies and raising public awareness about wetland protection [129,131]. Such efforts not only conserve biodiversity but also support local livelihoods that rely on wetlands for fisheries, agriculture, and eco-tourism [27,28].
     The different conservation strategies, such as in-situ and ex-situ conservation methods, enable effective and efficient fungal conservation. The conservation of fungi by protecting the environment in which they live, following the in-situ strategy, ensures the protection of the entire biome along with the fungi [134].
     The fungal culture collections in the world, such as the China General Microbiological Culture Collection (CGMCC), Guizhou Academy of Agricultural Sciences (GZAAS), National Culture Collection of Pathogenic Fungi (NCCPF), and Mae Fah Luang University Culture Collection (MFLUCC), are examples in ex-situ conservation of fungi by cataloguing, preserving, and making fungal strains available for further research [135, 136].
     Research in fungal ecology, taxonomy, and applied mycology provides essential guidance for conservation planning. Detailed studies help identify priority areas for protection, inform management strategies, and support the sustainable use of freshwater wetland resources (25,120). Fungal taxonomic research, in particular, is fundamental to conservation efforts, as it reveals the richness and ecological roles of fungal communities, enabling more targeted and effective protection of wetland biodiversity.

7. RESEARCH GAPS AND FUTURE PROSPECTIVE

     Although knowledge of freshwater wetlands is expanding, many aspects remain underexplored, particularly in relation to fungal diversity. Phang et al. [132] and Calabon et al. [25] noted the limited investigation of specific substrates in aquatic environments, including freshwater-associated grasses and other Poales, which dominate coastal wetlands in Thailand. These plants offer promising opportunities for detailed fungal taxonomic studies. Additionally, the ecological roles and phylogenetic significance of fungal morphological traits, such as appendages and mucilaginous sheaths, remain insufficiently understood in freshwater fungi [25]. Most taxa are known only from a single morph, and understudied habitats may reveal connections between sexual and asexual forms, alongside new molecular data for insufficiently characterized species [30].
     Wetlands provide a range of microhabitats that support fungal specialization and diversification [60,61]. In freshwater coastal wetlands, the same plant species can present multiple fungal niches including submerged, aerial, or intermediate, each influenced by fluctuating water levels. These zones expose fungi to distinct environmental stresses and ecological opportunities [30,133]. However, vertical stratification of fungal communities in such habitats remains insufficiently studied. Filling this knowledge gap could lead to the discovery of novel taxa and illuminate unique ecological interactions between fungi and their host substrates.

ACKNOWLEDGEMENTS

     Amuhenage T. Bhagya would like to thank Mae Fah Luang University for Partial Scholarship for the doctoral degree program under GR-ST-PS-65-18, Mushroom Research Foundation, and the National Research Council of Thailand (NRCT) grant, “Total fungal diversity in a given forest area with implications towards species numbers, chemical diversity, and biotechnology” under N42A650547 for the provided support. Chayanard Phukhamsakda was funded by the Alexander von Humboldt (AvH) foundation for a Fellowship of Experienced Researchers stipend. Kevin D. Hyde and Gareth Jones thank the Distinguished Scientist Fellowship Program (DSFP), King Saud University, Kingdom of Saudi Arabia for funding. Authors like to thank National Parks, forest parks, Botanic Gardens and Arboreta for facilitating the research project by providing permission to collect and process samples (No. 0907.4/23579, book no. 02/020 and book no. 02/001).

AUTHOR CONTRIBUTIONS

     Amuhenage T. Bhagya: Writing - Original draft preparation. Chayanard Phukhamsakda: Supervision, Reviewing and Editing, Project administration. E. B. Gareth Jones: Supervision, Reviewing and Editing, Funding acquisition. Kevin D. Hyde: Supervision, Reviewing and Editing, Funding acquisition.

CONFLICT OF INTEREST STATEMENT

     The authors declare no conflict of interest.

FUNDING

     Mushroom Research Foundation, doctoral degree program under GR-ST-PS-65-18 and National Research Council of Thailand (NRCT) grant under N42A650547.

REFERENCES

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