Marine fungi

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Morphological diversity of fungi collected from a marine sponge species, Ircinia variabilis Morphological diversity of marine fungi.jpg
Morphological diversity of fungi collected from a marine sponge species, Ircinia variabilis

Marine fungi are species of fungi that live in marine or estuarine environments. They are not a taxonomic group, but share a common habitat. Obligate marine fungi grow exclusively in the marine habitat while wholly or sporadically submerged in sea water. Facultative marine fungi normally occupy terrestrial or freshwater habitats, but are capable of living or even sporulating in a marine habitat.

Contents

About 2,149 species of marine fungi have been described, within eleven phyla and 856 genera, although only about 64 species have been fully genetically sequenced. [2] Many species of marine fungi are known only from spores and it is likely a large number of species have yet to be discovered. [3] It has been estimated that less than 1% of all marine fungal species have been described, due to difficulty in targeting marine fungal DNA and difficulties that arise in attempting to grow cultures of marine fungi. [4] It is impracticable to culture many of these fungi, but their nature can be investigated by examining seawater samples and undertaking rDNA analysis of the fungal material found. [3]

Different marine habitats support very different fungal communities. Fungi can be found in niches ranging from ocean depths and coastal waters to mangrove swamps and estuaries with low salinity levels. [5] Marine fungi can be saprobic or parasitic on animals, saprobic or parasitic on algae, saprobic on plants, or saprobic on dead wood. [6]

There has been some debate as to what exactly a marine fungus should be defined as. A definition used previously was "individuals with a long-term presence and metabolic activities in a marine habitat." [7] A more commonly used definition now [8] [9] is from Ka-Lai et al. 2016: "any fungus that is recovered repeatedly from marine habitats because: 1) it is able to grow and/or sporulate (on substrata) in marine environments; 2) it forms symbiotic relationships with other marine organisms; or 3) it is shown to adapt and evolve at the genetic level or be metabolically active in marine environments." [10]

Overview

Terrestrial fungi play critical roles in nutrient cycling and food webs and can shape macroorganism communities as parasites and mutualists. Although estimates for the number of fungal species on the planet range from 1.5 to over 5 million, likely fewer than 10% of fungi have been identified so far. To date, a relatively small percentage of described species are associated with marine environments, with ~1,100 species retrieved exclusively from the marine environment. Nevertheless, fungi have been found in nearly every marine habitat explored, from the surface of the ocean to kilometers below ocean sediments. Fungi are hypothesized to contribute to phytoplankton population cycles and the biological carbon pump and are active in the chemistry of marine sediments. Many fungi have been identified as commensals or pathogens of marine animals (e.g., corals and sponges), plants, and algae. Despite their varied roles, remarkably little is known about the diversity of this major branch of eukaryotic life in marine ecosystems or their ecological functions. [11]

Fungi represent a large and diverse group of microorganisms in microbiological communities in the marine environment and have an important role in nutrient cycling. [12] They are divided into two major groups; obligate marine fungi and facultative marine fungi. [13] Obligate marine fungi are adapted to reproduce in the aquatic environment, while facultative marine fungi can grow in aquatic as well as terrestrial environments. [13] Marine fungi are called marine-derived fungi when their facultative or obligate state is not certain. [14]

Marine fungal species occur as saprobes, parasites, or symbionts and colonize a wide range of substrates, such as sponges, corals, mangroves, seagrasses and algae. [15] [16] [14] Factors that influence whether or not marine fungi are present in any particular location include the water temperature, its salinity, the water movement, the presence of suitable substrates for colonization, the presence of propagules in the water, interspecific competition, pollution and the oxygen content of the water. [5] Some marine fungi which have ventured into the sea from terrestrial habitats include species that burrow into sand grains, living in the pores. Others live inside stony corals, and may become pathogenic if the coral is stressed by rising sea temperatures. [3] [ self-published source? ] [17]

In 2011 the phylogeny of marine fungi was elucidated by analysis of their small subunit ribosomal DNA sequences. Thirty six new marine lineages were found, the majority of which were chytrids but also some filamentous and multicellular fungi. The majority of the species found were ascomycetous and basidiomycetous yeasts. [18] The secondary metabolites produced by marine fungi have high potential for use in biotechnological, medical and industrial applications. [19] [ better source needed ]

The ocean constitutes about 99% of the volume in the biosphere. Oceanic microbes comprise 70%–90% of the biomass of marine biota [20] [21] and are the main drivers of marine biogeochemical cycles. [22] They are responsible for determining the amount of carbon fixed during photosynthesis, the subsequent release of carbon back to CO2 through respiration, and ultimately the sequestration of carbon in the deep ocean over millennial scales. Present estimates of marine microbial biomass primarily rely on bacteria, archaea, and protists. Pelagic (planktonic) fungal biomass has been less studied and only superficially characterized, despite their recently documented ubiquitous presence and active participation in the marine carbon and nitrogen cycles. [23] [24] [25] [26] [27] [28] [29]

As of 2025, only a few studies on pelagic fungal biomass, restricted to distinct coastal areas, are available suggesting their potential importance in near-shore environments; however, an accurate representation of the vast open-ocean environment is still lacking. These available studies are based on different methods, each having their own limitations and therefore hampering a robust intercomparison of results. [30] Among the biomass of organisms/groups estimated in the ocean, fungi accounted for the highest uncertainty, ranging more than two orders of magnitude, [31] indicating the need for a direct estimate of fungal biomass in the ocean. There is need for large-scale sampling across biogeographical provinces in the open ocean to ensure precise quantification of pelagic fungi in the microbial biomass. [29]

Evolution

In contrast to plants and animals, the early fossil record of the fungi is meager. Since fungi do not biomineralise, they do not readily enter the fossil record. Fungal fossils are difficult to distinguish from those of other microbes, and are most easily identified when they resemble extant fungi. [32]

The earliest fossils possessing features typical of fungi date to the Paleoproterozoic era, some 2,400  million years ago (Ma). These multicellular benthic organisms had filamentous structures capable of anastomosis, in which hyphal branches recombine. [33] Other recent studies (2009) estimate the arrival of fungal organisms at about 760–1060 Ma on the basis of comparisons of the rate of evolution in closely related groups. [34]

Phylogenetic and symbiogenetic tree of living organisms, showing a view of the origins of eukaryotes and prokaryotes Tree of Living Organisms 2.png
Phylogenetic and symbiogenetic tree of living organisms, showing a view of the origins of eukaryotes and prokaryotes

For much of the Paleozoic Era (542–251 Ma), the fungi appear to have been aquatic and consisted of organisms similar to the extant Chytrids in having flagellum-bearing spores. [35] Phylogenetic analyses suggest that the flagellum was lost early in the evolutionary history of the fungi, and consequently, the majority of fungal species lack a flagellum. [36] Evidence from DNA analysis suggests that all fungi are descended from one common ancestor, at least 600 million years ago. It is probable that these earliest fungi lived in water, and had flagella. Fungi moved to land at about the same time as plants, about 460 million years ago, at least. [37] Although fungi are opisthokonts—a grouping of evolutionarily related organisms broadly characterized by a single posterior flagellum—all phyla except for the chytrids have lost their posterior flagella. [38]

The evolutionary adaptation from an aquatic to a terrestrial lifestyle necessitated a diversification of ecological strategies for obtaining nutrients, including parasitism, saprobism, and the development of mutualistic relationships such as mycorrhiza and lichenization. [39] Recent (2009) studies suggest that the ancestral ecological state of the Ascomycota was saprobism, and that independent lichenization events have occurred multiple times. [40]

The growth of fungi as hyphae on or in solid substrates or as single cells in aquatic environments is adapted for the efficient extraction of nutrients, because these growth forms have high surface area to volume ratios. [41] Hyphae are specifically adapted for growth on solid surfaces, and to invade substrates and tissues. [42] They can exert large penetrative mechanical forces; for example, many plant pathogens, including Magnaporthe grisea , form a structure called an appressorium that evolved to puncture plant tissues. [43] The pressure generated by the appressorium, directed against the plant epidermis, can exceed 8 megapascals (1,200 psi). [43] The filamentous fungus Paecilomyces lilacinus uses a similar structure to penetrate the eggs of nematodes. [44]

Fungi were considered to be part of the plant kingdom until the mid-20th century. By the middle of the 20th century, fungi were considered a distinct kingdom, and the newly recognized kingdom Fungi becoming the third major kingdom of multicellular eukaryotes with kingdom Plantae and kingdom Animalia, the distinguishing feature between these kingdoms being the way they obtain nutrition. [45]

Planktonic marine fungi

Marine plankton are organisms that drift in the ocean but are unable to actively propel themselves against currents. Planktonic marine fungi are found drifting in the marine water column in the form of spores, yeasts, mycelia, sporangia or other fungal propagules, in suspension or attached to particles, substrates or planktonic hosts. [46] [47]

Food webs and the mycoloop

Diagram of a mycoloop (fungus loop)
Parasitic chytrids can transfer material from large inedible phytoplankton to zooplankton. Chytrids zoospores are excellent food for zooplankton in terms of size (2-5 mm in diameter), shape, nutritional quality (rich in polyunsaturated fatty acids and cholesterols). Large colonies of host phytoplankton may also be fragmented by chytrid infections and become edible to zooplankton. Parasitic chytrids as a mycoloop.jpg
Diagram of a mycoloop (fungus loop)
Parasitic chytrids can transfer material from large inedible phytoplankton to zooplankton. Chytrids zoospores are excellent food for zooplankton in terms of size (2–5 μm in diameter), shape, nutritional quality (rich in polyunsaturated fatty acids and cholesterols). Large colonies of host phytoplankton may also be fragmented by chytrid infections and become edible to zooplankton.

The mycoloop is a trophic pathway in aquatic food webs where parasitic fungi, particularly chytrids, facilitate the transfer of nutrients and energy from large, inedible phytoplankton (algae) to zooplankton. This process enhances nutrient cycling and supports higher trophic levels in aquatic ecosystems.

Chytrids infect large, inedible phytoplankton, such as diatoms or cyanobacteria, and produce zoospores (free-living, motile spores, 2–5 μm in diameter). These zoospores are rich in nutrients like polyunsaturated fatty acids (PUFAs) and cholesterol, making them an excellent food source for zooplankton, such as Daphnia and rotifers. By consuming the zoospores or fragmented phytoplankton, zooplankton gain access to nutrients that would otherwise be unavailable from inedible phytoplankton, creating the trophic link called the mycoloop. In this manner, the mycoloop channels carbon, phosphorus, and other nutrients from phytoplankton to zooplankton, bypassing the limitations of inedible phytoplankton.

The mycoloop can influence phytoplankton blooms by reducing host populations (via parasitism) and supporting zooplankton growth, potentially stabilizing aquatic food webs. It can also influence the carbon cycle by altering carbon fluxes, reducing the sinking of large phytoplankton and redirecting carbon to higher trophic levels.

Roles of fungi in the marine carbon cycle
Roles of fungi in the marine carbon cycle by processing phytoplankton-derived organic matter. Parasitic fungi, as well as saprotrophic fungi, directly assimilate phytoplankton organic carbon. By releasing zoospores, the fungi bridge the trophic linkage to zooplankton, known as the mycoloop. By modifying the particulate and dissolved organic carbon, they can affect bacteria and the microbial loop. These processes may modify marine snow chemical composition and the subsequent functioning of the biological carbon pump. Roles of fungi in the marine carbon cycle.jpg
Roles of fungi in the marine carbon cycle
Roles of fungi in the marine carbon cycle by processing phytoplankton-derived organic matter. Parasitic fungi, as well as saprotrophic fungi, directly assimilate phytoplankton organic carbon. By releasing zoospores, the fungi bridge the trophic linkage to zooplankton, known as the mycoloop. By modifying the particulate and dissolved organic carbon, they can affect bacteria and the microbial loop. These processes may modify marine snow chemical composition and the subsequent functioning of the biological carbon pump.
Mycoloop links between phytoplankton and zooplankton
Chytrid-mediated trophic links between phytoplankton and zooplankton (mycoloop). While small phytoplankton species can be grazed upon by zooplankton, large phytoplankton species constitute poorly edible or even inedible prey. Chytrid infections on large phytoplankton can induce changes in palatability, as a result of host aggregation (reduced edibility) or mechanistic fragmentation of cells or filaments (increased palatability). First, chytrid parasites extract and repack nutrients and energy from their hosts in form of readily edible zoospores. Second, infected and fragmented hosts including attached sporangia can also be ingested by grazers (i.e. concomitant predation). Mycoloop links between phytoplankton and zooplankton.jpg
Mycoloop links between phytoplankton and zooplankton
Chytrid‐mediated trophic links between phytoplankton and zooplankton (mycoloop). While small phytoplankton species can be grazed upon by zooplankton, large phytoplankton species constitute poorly edible or even inedible prey. Chytrid infections on large phytoplankton can induce changes in palatability, as a result of host aggregation (reduced edibility) or mechanistic fragmentation of cells or filaments (increased palatability). First, chytrid parasites extract and repack nutrients and energy from their hosts in form of readily edible zoospores. Second, infected and fragmented hosts including attached sporangia can also be ingested by grazers (i.e. concomitant predation).

Algae and phytoplankton

Chytrid parasites of marine diatoms. (A) Chytrid sporangia on Pleurosigma sp. The white arrow indicates the operculate discharge pore. (B) Rhizoids (white arrow) extending into diatom host. (C) Chlorophyll aggregates localized to infection sites (white arrows). (D and E) Single hosts bearing multiple zoosporangia at different stages of development. The white arrow in panel E highlights branching rhizoids. (F) Endobiotic chytrid-like sporangia within diatom frustule. Bars = 10 mm. Chytrid parasites of marine diatoms.jpg
Chytrid parasites of marine diatoms. (A) Chytrid sporangia on Pleurosigma sp. The white arrow indicates the operculate discharge pore. (B) Rhizoids (white arrow) extending into diatom host. (C) Chlorophyll aggregates localized to infection sites (white arrows). (D and E) Single hosts bearing multiple zoosporangia at different stages of development. The white arrow in panel E highlights branching rhizoids. (F) Endobiotic chytrid-like sporangia within diatom frustule. Bars = 10 μm.

Marine fungi associated with algae are largely unexplored, despite their ecological role and potential industrial applications. For example, it has been shown that fungi associated with algae produce many bioactive secondary metabolites. [52] [53] [54] Algae derived fungi can be associated with a variety of algae, including brown (e.g., Agarum clathratum , Fucus sp., Laminaria sp., Sargassum sp.), green (e.g., Ulva sp., Enteromorpha sp., Flabellia sp.), or red (e.g. Chondrus sp., Dilsea sp., Ceramium sp.) algae. [55] [56] [57] [58] [59] [14]

Almost one-third of all known marine fungal species are associated with algae. [60] The most commonly described fungi associated with algae belong to the Ascomycota and are represented by a wide diversity of genera such as Acremonium , Alternaria , Aspergillus , Cladosporium , Phoma , Penicillium , Trichoderma , Emericellopsis , Retrosium , Spathulospora , Pontogenia and Sigmoidea . [56] [61] [58] [59] [62] [63] [64] [14]

Rhyzophydium littoreum is a marine chytrid, a primitive fungus that infects green algae in estuaries. It obtains nutrients from the host alga and produces swimming zoospores that must survive in open water, a low nutrient environment, until a new host is encountered. [65] Another fungus, Ascochyta salicorniae , found growing on seaweed is being investigated for its action against malaria, [66] a mosquito-borne infectious disease of humans and other animals.

Other roles

Role of fungi in ocean carbon sequestration
This representation includes the traditionally neglected pelagic fungi, both parasitic and saprotrophic, highlighting the central role played by them, parasitic fungi in the mycoloop, and saprotrophic fungi as active contributors to the microbial loop. As depicted by this diagram, the activity of heterotrophic microbes, including pelagic fungi, has far-reaching global implications for fisheries (i.e., the amount of carbon that will ultimately flow to higher trophic levels) and climate change (i.e., the amount of carbon that will be sequestered in the ocean or respired back to CO2 and the release of other greenhouse gases; e.g., N2O). Role of fungi in ocean carbon sequestration.jpg
Role of fungi in ocean carbon sequestration
This representation includes the traditionally neglected pelagic fungi, both parasitic and saprotrophic, highlighting the central role played by them, parasitic fungi in the mycoloop, and saprotrophic fungi as active contributors to the microbial loop. As depicted by this diagram, the activity of heterotrophic microbes, including pelagic fungi, has far-reaching global implications for fisheries (i.e., the amount of carbon that will ultimately flow to higher trophic levels) and climate change (i.e., the amount of carbon that will be sequestered in the ocean or respired back to CO2 and the release of other greenhouse gases; e.g., N2O).

Apart from serving as a food source for zooplankton and other microorganisms in the microbial loop, marine planktonic fungi (and ocean pelagic fungi) play other, often understudied, roles in marine ecosystems, contributing to: [30]

In the marine environment, fungi have been detected in every explored habitat, from surface to deeper waters, from the coast to the open ocean, and from beaches to deep sediments. [11] [67] However, compared to their terrestrial counterparts, planktonic marine fungi, belonging to the mycoplankton, have been much less studied in terms of occurrence, biodiversity, dynamics, and contribution to ecosystem processes. [68] [69] Like terrestrial species, marine fungi are thought to contribute to organic matter degradation processes and nutrient cycling by acting as saprotrophic (i.e., decomposers) or parasitic organisms at different trophic levels. [70] (For example, fungal zoospores efficiently transfer organic matter from large, otherwise inedible, phytoplankton cells to zooplankton, in the process termed mycoloop.) While mycoplankton are thought to have significant impacts on ecosystems, these organisms remain poorly understood. [30] [71]

As of 2025, metagenomic analysis shows the widespread presence of pelagic fungi in the global ocean, yet their quantitative contribution to carbon stocks remains elusive, hindering their incorporation into biogeochemical models. [29]

Marine plants

Mangroves

Avicennia marina - white mangrove (8697515733).jpg
A white mangrove, Avicennia marina
Annabella australiensis.png
Annabella australiensis, a helotialean fungi found on the decaying wood of an Australian white mangrove [72]

The greatest number of known species of marine fungi are from mangrove swamps. [6] In one study, blocks of mangrove wood and pieces of driftwood of Avicennia alba , Bruguiera cylindrica and Rhizophora apiculata were examined to identify the lignicolous (wood-decaying) fungi they hosted. Also tested were Nypa fruticans , a mangrove palm and Acanthus ilicifolius , a plant often associated with mangroves. Each material was found to have its own characteristic fungi, the greatest diversity being among those growing on the mangrove palm. It was surmised that this was because the salinity was lower in the estuaries and creeks where Nypa grew, and so it required a lesser degree of adaptation for the fungi to flourish there. Some of these species were closely related to fungi on terrestrial palms. Other studies have shown that driftwood hosts more species of fungus than do exposed test blocks of wood of a similar kind. The mangrove leaf litter also supported a large fungal community which was different from that on the wood and living material. However, few of these were multicellular, higher marine fungi. [5]

Other plants

Spartinadensiflora.jpg
Spartina plant
Littoraria irrorata.jpg
The shell of this sea snail, Littoraria irrorata, is covered with the lichen Pyrenocollema halodytes

The sea snail Littoraria irrorata damages plants of Spartina in the coastal sea marshes where it lives, which enables spores of intertidal ascomycetous fungi to colonise the plant. The snail eats the fungal growth in preference to the grass itself. This mutualism between the snail and the fungus is considered to be the first example of husbandry among invertebrate animals outside the class Insecta. [73]

Eelgrass, Zostera marina , is sometimes affected by seagrass wasting disease. The primary cause of this seems to be pathogenic strains of the protist Labyrinthula zosterae , but it is thought that fungal pathogens also contribute and may predispose the eelgrass to disease. [74] [75]

Wood

Driftwood Driftwood Beach.jpg
Driftwood

Many marine fungi are very specific as to which species of floating and submerged wood they colonise. A range of species of fungi colonise beech, while oak supports a different community. When a fungal propagule lands on a suitable piece of wood, it will grow if no other fungi are present. If the wood is already colonised by another fungal species, growth will depend on whether that fungus produces antifungal chemicals and whether the new arrival can resist them. The chemical properties of colonizing fungi also affect the animal communities that graze on them: in one study, when hyphae from five different species of marine fungi were fed to nematodes, one species supported less than half the number of nematodes per mg of hyphae than did the others. [65]

Detection of fungi in wood may involve incubation at a suitable temperature in a suitable water medium for a period of six months to upward of eighteen months. [65]

Lichens

Lichen covered rocks Lichen rock.jpeg
Lichen covered rocks

Lichens are mutualistic associations between fungi, usually an ascomycete with a basidiomycete, [76] and an alga or a cyanobacterium. Several lichens, including Arthopyrenia halodytes , Pharcidia laminariicola , Pharcidia rhachiana and Turgidosculum ulvae , are found in marine environments. [6] Many more occur in the splash zone, where they occupy different vertical zones depending on how tolerant they are to submersion. [77] Lichen-like fossils have been found in the Doushantuo Formation in China dating back about 600 million years ago. [78]

Fungi from Verrucariales also form marine lichens with the brown algae Petroderma maculiforme, [79] and have a symbiotic relationship with seaweed like (rockweed) and Blidingia minima , where the algae are the dominant components. The fungi is thought to help the rockweeds to resist desiccation when exposed to air. [80] [81] In addition, lichens can also use yellow-green algae ( Heterococcus ) as their symbiotic partner. [82]

Lichen-like fossils consisting of coccoid cells (cyanobacteria?) and thin filaments (mucoromycotinan Glomeromycota?) are permineralized in marine phosphorite of the Doushantuo Formation in southern China. These fossils are thought to be 551 to 635 million years old or Ediacaran. [78] Ediacaran acritarchs also have many similarities with Glomeromycotan vesicles and spores. [83] It has also been claimed that Ediacaran fossils including Dickinsonia , [84] were lichens, [85] although this claim is controversial. [86] Endosymbiotic Glomeromycota comparable with living Geosiphon may extend back into the Proterozoic in the form of 1500 million year old Horodyskia [87] and 2200 million year old Diskagma . [88] Discovery of these fossils suggest that marine fungi developed symbiotic partnerships with photoautotrophs long before the evolution of vascular plants. However a 2019 study concluded through age estimations obtained by time calibrated phylogenies, and absence of unambiguous fossil data that the origins of lichens postdate the evolution of vascular plants. [89]

Not to be confused with lichens are Mycophycobiosis, similar to lichens in being a symbiosis of an algae and a fungus. In mycophycobiosis the algae forms the external, multicellular structure housing the fungus. The reproduction of both partners is always disjoint. [90]

Invertebrates

The American lobster (Homarus americanus), like many other marine crustaceans, incubates its eggs beneath its tail segments. Here they are exposed to water-borne micro-organisms including fungi during their long period of development. The lobster has a symbiotic relationship with a gram-negative bacterium that has anti-fungal properties. This bacterium grows over the eggs and protects them from infection by the pathogenic fungus-like oomycete Lagenidium callinectes . The metabolite produced by the bacterium is tyrosol, a 4-hydroxyphenethyl alcohol, an antibiotic substance also produced by some terrestrial fungi. Similarly, a shrimp found in estuaries, Palaemon macrodactylis , has a symbiotic bacterium that produces 2,3-indolenedione, a substance that is also toxic to the oomycete Lagenidium callinectes. [91]

Vertebrates

Salmon with fungal disease Grilse001.jpg
Salmon with fungal disease

Whales, porpoises and dolphins are susceptible to fungal diseases but these have been little researched in the field. Mortalities from fungal disease have been reported in captive orcas; it is thought that stress due to captive conditions may have been predisposing. Transmission among animals in the open sea may naturally limit the spread of fungal diseases. Infectious fungi known from orcas include Aspergillus fumigatus , Candida albicans and Saksenaea vasiformis . Fungal infections in other cetaceans include Coccidioides immitis , Cryptococcus neoformans , Loboa loboi , Rhizopus sp., Aspergillus flavus , Blastomyces dermatitidis , Cladophialophora bantiana , Histoplasma capsulatum , Mucor sp., Sporothrix schenckii and Trichophyton sp. [92]

Salmonids farmed in cages in marine environments may be affected by a number of different fungal infections. Exophiala salmonis causes an infection in which growth of hyphae in the kidneys causes swelling of the abdomen. A cellular response by the fish aims to isolate the fungus by walling it off. Fish are also susceptible to fungus-like oomycetes including Branchiomyces which affects the gills of various fishes, and Saprolegnia which attacks damaged tissue. [93]

Arctic marine fungi

Pennate diatom from an Arctic meltpond, infected with two chytrid-like [zoo-]sporangium fungal pathogens (in false-colour red). Scale bar = 10 mm. Pennate diatom infected with two chytrid-like fungal pathogens.png
Pennate diatom from an Arctic meltpond, infected with two chytrid-like [zoo-]sporangium fungal pathogens (in false-colour red). Scale bar = 10 μm.

Marine fungi have been observed as far north as the Arctic Ocean. Chytridiomycota, the dominant parasitic fungal organism in Arctic waters, take advantage of phytoplankton blooms in brine channels caused by warming temperatures and increased light penetration through the ice. These fungi parasitize diatoms, thereby controlling algal blooms and recycling carbon back into the microbial food web. Arctic blooms also provide conducive environments for other parasitic fungi. Light levels and seasonal factors, such as temperature and salinity, also control chytrid activity independently of phytoplankton populations. During periods of low temperatures and phytoplankton levels, Aureobasidium and Cladosporium populations overtake those of chytrids within the brine channels. [95]

Marine sediment

Ascomycota, Basidiomycota, and Chytridiomycota have been observed in marine sediments ranging in depth from 0 to 1740 meters beneath the ocean floor. One study analyzed subsurface samples of marine sediment between these depths and isolated all observable fungi. Isolates showed that most subsurface fungal diversity was found between depths of 0 to 25 meters below the sea floor with Fusarium oxysporum and Rhodotorula mucilaginosa being the most prominent. Overall, the ascomycota are the dominant subsurface phylum. [96] Almost all fungal species recovered have also been observed in terrestrial sediments with spore-sourcing indicating terrestrial origin. [96] [97]

Contrary to previous beliefs, deep subsurface marine fungi actively grow and germinate, with some studies showing increased growth rates under high hydrostatic pressures. Though the methods by which marine fungi are able to survive the extreme conditions of the seafloor and below is largely unknown, Saccharomyces cerevisiae shines some light onto adaptations that make it possible. This fungus strengthens its outer membrane in order to endure higher hydrostatic pressures. [96] [ clarification needed ]

Several sediment-dwelling marine fungi are involved in biogeochemical processes. Fusarium oxysporum and Fusarium solani are denitrifiers both in marine and terrestrial environments. [96] [98] Some are co-denitrifying, fixing nitrogen into nitrous oxide and dinitrogen. [97]  Still others process organic matter including carbohydrate, proteins, and lipids. Ocean crust fungi, like those found around hydrothermal vents, decompose organic matter, and play various roles in manganese and arsenic cycling. [11]

Sediment-bound marine fungi played a major role in breaking down oil spilled from the Deepwater Horizons disaster in 2010. Aspergillus,Penicillium, and Fusarium species, among others, can degrade high-molecular-weight hydrocarbons as well as assist hydrocarbon-degrading bacteria. [11]

Fungal dark matter

Microbial dark matter are microbial organisms microbiologists cannot culture in laboratories, due to lack of knowledge of the required growth conditions. Most microbial organisms fall in this category. [99] [100] Fungal dark matter refers specifically to the vast number of fungal species that are known only through environmental DNA (eDNA) sequencing but cannot be cultured, observed morphologically, or formally described under current taxonomic rules. [68] [101] These fungi, also called dark fungi or dark taxa, are detected via genetic material in soil, water, or other substrates, but their physical forms (e.g., fruiting bodies) or ecological roles remain largely unknown due to their resistance to cultivation and lack of visible structures. There is some analogy to cosmological dark matter, which is inferred but not directly observed, and are significant because they may dominate fungal diversity. [102] [103]

Human uses

Biomass processors

Medical

Marine fungi produce antiviral and antibacterial compounds as metabolites with upwards of 1,000 having realized and potential uses as anticancer, anti-diabetic, and anti-inflammatory drugs. [104] [105]

The antiviral properties of marine fungi were realized in 1988 after their compounds were used to successfully treat the H1N1 flu virus. In addition to H1N1, antiviral compounds isolated from marine fungi have been shown to have virucidal effects on HIV, herpes simplex 1 and 2, Porcine Reproductive and Respiratory Syndrome Virus, and Respiratory Syncytial Virus. Most of these antiviral metabolites were isolated from species of Aspergillus , Penicillium , Cladosporium , Stachybotrys , and Neosartorya. These metabolites inhibit the virus's ability to replicate, thereby slowing infections. [104]

Mangrove-associated fungi have prominent antibacterial effects on several common pathogenic human bacteria including, Staphylococcus aureus and Pseudomonas aeruginosa . High competition between organisms within mangrove niches lead to increases in antibacterial substances produced by these fungi as defensive agents. [106] Penicillium and Aspergillus species are the largest producers of antibacterial compounds among the marine fungi. [107]

Various deep-sea marine fungi species have recently been shown to produce anti-cancer metabolites. One study uncovered 199 novel cytotoxic compounds with anticancer potential. In addition to cytotoxic metabolites, these compounds have structures capable of disrupting cancer-activated telomerases via DNA binding. Others inhibit the topoisomerase enzyme from continuing to aid in the repair and replication of cancer cells. [105]

See also

References

  1. Paz, Z.; Komon-Zelazowska, M.; Druzhinina, I. S.; Aveskamp, M. M.; Shnaiderman, A.; Aluma, Y.; Carmeli, S.; Ilan, M.; Yarden, O. (30 January 2010). "Diversity and potential antifungal properties of fungi associated with a Mediterranean sponge". Fungal Diversity. 42 (1): 17–26. doi:10.1007/s13225-010-0020-x. S2CID   45289302.
  2. Jones, EB Gareth et al. Marine Fungi. Originally published May 17, 2019. Last updated February 27, 2025. Accessed April 2, 2025.
  3. 1 2 3 "Cool New Paper: Marine Fungi". Teaching Biology. 17 December 2011. Archived from the original on March 16, 2012.
  4. Gladfelter, Amy S.; James, Timothy Y.; Amend, Anthony S. (March 2019). "Marine fungi". Current Biology. 29 (6): R191 –R195. Bibcode:2019CBio...29.R191G. doi: 10.1016/j.cub.2019.02.009 . PMID   30889385.
  5. 1 2 3 E. B. Gareth Jones (2000). "Marine fungi: some factors influencing biodiversity" (PDF). Fungal Diversity. 4: 53–73.
  6. 1 2 3 Species of Higher Marine Fungi Archived 2013-04-22 at the Wayback Machine University of Mississippi. Retrieved 2012-02-05.
  7. Stal, Lucas J.; Cretoiu, Mariana Silvia (June 3, 2016). "Marine Fungi". The Marine Microbiome: An Untapped Source of Biodiversity and Biotechnological Potential. Springer. ISBN   9783319330006.
  8. Sen, Kalyani; Sen, Biswarup; Wang, Guangyi (2022). "Diversity, Abundance, and Ecological Roles of Planktonic Fungi in Marine Environments". Journal of Fungi. 8 (5): 491. doi: 10.3390/jof8050491 . PMC   9147564 . PMID   35628747.
  9. Mitchison-Field, Lorna; et al. (June 2, 2019). "Unconventional cell division cycles from marine-derived yeasts". Current Biology.
  10. Pang, Ka-Lai; et al. (September 2016). "'Marine fungi' and 'marine-derived fungi' in natural product chemistry research: Toward a new consensual definition". British Mycological Society. 30 (4): 163. Bibcode:2016FunBR..30..163P. doi:10.1016/j.fbr.2016.08.001.
  11. 1 2 3 4 5 Amend, Anthony; Burgaud, Gaetan; Cunliffe, Michael; Edgcomb, Virginia P.; Ettinger, Cassandra L.; Gutiérrez, M. H.; Heitman, Joseph; Hom, Erik F. Y.; Ianiri, Giuseppe; Jones, Adam C.; Kagami, Maiko; Picard, Kathryn T.; Quandt, C. Alisha; Raghukumar, Seshagiri; Riquelme, Mertixell; Stajich, Jason; Vargas-Muñiz, José; Walker, Allison K.; Yarden, Oded; Gladfelter, Amy S. (2019-04-30). "Fungi in the Marine Environment: Open Questions and Unsolved Problems" (PDF). mBio. 10 (2). doi: 10.1128/mBio.01189-18 . ISSN   2161-2129 . Retrieved 2025-08-31. CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  12. Tisthammer, K.H.; Cobian, G.M.; Amend, A.S. Global biogeography of marine fungi is shaped by the environment. Fungal Ecol. 2016, 19, 39–46.
  13. 1 2 Raghukumar, S. The marine environment and the role of fungi. In Fungi in Coastal and Oceanic Marine Ecosystems: Marine Fungi; Springer International Publishing: Cham, Switzerland, 2017; pp. 17–38.
  14. 1 2 3 4 Patyshakuliyeva, A., Falkoski, D.L., Wiebenga, A., Timmermans, K. and De Vries, R.P. (2020) "Macroalgae derived fungi have high abilities to degrade algal polymers". Microorganisms, 8(1): 52. doi : 10.3390/microorganisms8010052 . CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  15. Jones, E.B.G.; Suetrong, S.; Sakayaroj, J.; Bahkali, A.H.; Abdel-Wahab, M.A.; Boekhout, T.; Pang, K.-L. Classification of marine Ascomycota, Basidiomycota, Blastocladiomycota and Chytridiomycota. Fungal Divers. 2015, 73, 1–72.
  16. Debbab, A.; Aly, A.H.; Proksch, P. Mangrove derived fungal endophytes–a chemical and biological perception. Fungal Divers. 2013, 61, 1–27.
  17. Holmquist, G. U.; H. W. Walker & Stahr H. M. (1983). "Influence of Temperature, pH, Water Activity and Antifungal Agents on Growth of Aspergillus flavus and A. parasiticus". Journal of Food Science. 48 (3): 778–782. doi:10.1111/j.1365-2621.1983.tb14897.x.
  18. Richards, Thomas A.; Jones, Meredith D.M.; Leonard, Guy; Bass, David (15 January 2012). "Marine Fungi: Their Ecology and Molecular Diversity". Annual Review of Marine Science. 4 (1): 495–522. Bibcode:2012ARMS....4..495R. doi:10.1146/annurev-marine-120710-100802. PMID   22457985.
  19. Marine Fungi Retrieved 2012-02-06.
  20. Bar-On, Yinon M.; Phillips, Rob; Milo, Ron (2018-06-19). "The biomass distribution on Earth" (PDF). Proceedings of the National Academy of Sciences. 115 (25): 6506–6511. doi: 10.1073/pnas.1711842115 . ISSN   0027-8424. PMC   6016768 . PMID   29784790 . Retrieved 2025-09-01.
  21. Cavicchioli, Ricardo; Ripple, William J.; Timmis, Kenneth N.; Azam, Farooq; Bakken, Lars R.; Baylis, Matthew; Behrenfeld, Michael J.; Boetius, Antje; Boyd, Philip W.; Classen, Aimée T.; Crowther, Thomas W.; Danovaro, Roberto; Foreman, Christine M.; Huisman, Jef; Hutchins, David A.; Jansson, Janet K.; Karl, David M.; Koskella, Britt; Mark Welch, David B.; Martiny, Jennifer B. H.; Moran, Mary Ann; Orphan, Victoria J.; Reay, David S.; Remais, Justin V.; Rich, Virginia I.; Singh, Brajesh K.; Stein, Lisa Y.; Stewart, Frank J.; Sullivan, Matthew B.; van Oppen, Madeleine J. H.; Weaver, Scott C.; Webb, Eric A.; Webster, Nicole S. (2019). "Scientists' warning to humanity: microorganisms and climate change" (PDF). Nature Reviews Microbiology. 17 (9): 569–586. doi: 10.1038/s41579-019-0222-5 . ISSN   1740-1526. PMC   7136171 . PMID   31213707 . Retrieved 2025-09-01. CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License
  22. Falkowski, Paul G.; Fenchel, Tom; Delong, Edward F. (2008-05-23). "The Microbial Engines That Drive Earth's Biogeochemical Cycles" (PDF). Science. 320 (5879): 1034–1039. doi: 10.1126/science.1153213 . ISSN   0036-8075 . Retrieved 2025-09-01.
  23. Cunliffe, Michael; Hollingsworth, Anita; Bain, Cal; Sharma, Vikram; Taylor, Joe D. (2017). "Algal polysaccharide utilisation by saprotrophic planktonic marine fungi" (PDF). Fungal Ecology. 30: 135–138. doi: 10.1016/j.funeco.2017.08.009 . Retrieved 2025-09-01.
  24. Morales, Sergio E.; Biswas, Ambarish; Herndl, Gerhard J.; Baltar, Federico (2019-03-19). "Global Structuring of Phylogenetic and Functional Diversity of Pelagic Fungi by Depth and Temperature". Frontiers in Marine Science. 6. doi: 10.3389/fmars.2019.00131 . ISSN   2296-7745.
  25. Orsi, William D; Vuillemin, Aurèle; Coskun, Ömer K; Rodriguez, Paula; Oertel, Yanik; Niggemann, Jutta; Mohrholz, Volker; Gomez-Saez, Gonzalo V (2022-05-01). "Carbon assimilating fungi from surface ocean to subseafloor revealed by coupled phylogenetic and stable isotope analysis" (PDF). The ISME Journal. 16 (5): 1245–1261. doi: 10.1038/s41396-021-01169-5 . ISSN   1751-7362. PMC   9038920 . PMID   34893690 . Retrieved 2025-09-01.
  26. Baltar, Federico; Zhao, Zihao; Herndl, Gerhard J. (2021). "Potential and expression of carbohydrate utilization by marine fungi in the global ocean". Microbiome. 9 (1). doi: 10.1186/s40168-021-01063-4 . ISSN   2049-2618. PMC   8114511 . PMID   33975640.
  27. Breyer, Eva; Zhao, Zihao; Herndl, Gerhard J.; Baltar, Federico (2022-09-01). "Global contribution of pelagic fungi to protein degradation in the ocean". Microbiome. 10 (1). doi: 10.1186/s40168-022-01329-5 . ISSN   2049-2618. PMC   9434897 . PMID   36050758.
  28. Peng, Xuefeng; Valentine, David L. (2021-03-17). "Diversity and N2O Production Potential of Fungi in an Oceanic Oxygen Minimum Zone". Journal of Fungi. 7 (3): 218. doi: 10.3390/jof7030218 . ISSN   2309-608X. PMC   8002691 . PMID   33802893.
  29. 1 2 3 Breyer, Eva; Stix, Constanze; Kilker, Sophie; Roller, Benjamin R.K.; Panagou, Fragkiski; Doebke, Charlotte; Amano, Chie; Saavedra, Daniel E.M.; Coll-García, Guillem; Steger-Mähnert, Barbara; Dachs, Jordi; Berrojalbiz, Naiara; Vila-Costa, Maria; Sobrino, Cristina; Fuentes-Lema, Antonio; Berthiller, Franz; Polz, Martin F.; Baltar, Federico (2025). "The contribution of pelagic fungi to ocean biomass". Cell. 188 (15): 3992–4002.e13. doi: 10.1016/j.cell.2025.05.004 . CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License
  30. 1 2 3 4 Breyer, Eva; Baltar, Federico (2023). "The largely neglected ecological role of oceanic pelagic fungi". Trends in Ecology & Evolution. 38 (9): 870–888. doi: 10.1016/j.tree.2023.05.002 . Retrieved 2025-08-28. CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  31. Bar-On, Yinon M.; Milo, Ron (2019). "The Biomass Composition of the Oceans: A Blueprint of Our Blue Planet". Cell. 179 (7): 1451–1454. doi: 10.1016/j.cell.2019.11.018 . Retrieved 2025-09-01.
  32. Donoghue MJ; Cracraft J (2004). Assembling the Tree of Life. Oxford (Oxfordshire), UK: Oxford University Press. p. 187. ISBN   978-0-19-517234-8.
  33. Bengtson, Stefan; Rasmussen, Birger; Ivarsson, Magnus; Muhling, Janet; Broman, Curt; Marone, Federica; Stampanoni, Marco; Bekker, Andrey (24 April 2017). "Fungus-like mycelial fossils in 2.4-billion-year-old vesicular basalt". Nature Ecology & Evolution. 1 (6): 0141. Bibcode:2017NatEE...1..141B. doi:10.1038/s41559-017-0141. hdl: 20.500.11937/67718 . ISSN   2397-334X. PMID   28812648. S2CID   25586788.
  34. Lucking R, Huhndorf S, Pfister D, Plata ER, Lumbsch H (2009). "Fungi evolved right on track" . Mycologia. 101 (6): 810–822. doi:10.3852/09-016. PMID   19927746. S2CID   6689439.
  35. James TY; et al. (2006). "Reconstructing the early evolution of Fungi using a six-gene phylogeny". Nature. 443 (7113): 818–22. Bibcode:2006Natur.443..818J. doi:10.1038/nature05110. PMID   17051209. S2CID   4302864.
  36. Liu YJ, Hodson MC, Hall BD (2006). "Loss of the flagellum happened only once in the fungal lineage: phylogenetic structure of Kingdom Fungi inferred from RNA polymerase II subunit genes". BMC Evolutionary Biology. 6 (1): 74. doi: 10.1186/1471-2148-6-74 . PMC   1599754 . PMID   17010206.
  37. "CK12-Foundation". flexbooks.ck12.org. Retrieved 2020-05-19.
  38. Steenkamp ET, Wright J, Baldauf SL (January 2006). "The protistan origins of animals and fungi". Molecular Biology and Evolution. 23 (1): 93–106. doi: 10.1093/molbev/msj011 . PMID   16151185.
  39. Taylor and Taylor, pp. 84–94 and 106–107.
  40. Schoch CL; Sung G-H; López-Giráldez F; et al. (2009). "The Ascomycota tree of life: A phylum-wide phylogeny clarifies the origin and evolution of fundamental reproductive and ecological traits". Systematic Biology. 58 (2): 224–39. doi: 10.1093/sysbio/syp020 . PMID   20525580.
  41. Moss ST (1986). The Biology of Marine Fungi. Cambridge, UK: Cambridge University Press. p. 76. ISBN   978-0-521-30899-1.
  42. Peñalva MA, Arst HN (September 2002). "Regulation of gene expression by ambient pH in filamentous fungi and yeasts". Microbiology and Molecular Biology Reviews. 66 (3): 426–46, table of contents. doi:10.1128/MMBR.66.3.426-446.2002. PMC   120796 . PMID   12208998.
  43. 1 2 Howard RJ, Ferrari MA, Roach DH, Money NP (December 1991). "Penetration of hard substrates by a fungus employing enormous turgor pressures". Proceedings of the National Academy of Sciences of the United States of America. 88 (24): 11281–4. Bibcode:1991PNAS...8811281H. doi: 10.1073/pnas.88.24.11281 . PMC   53118 . PMID   1837147.
  44. Money NP (1998). "Mechanics of invasive fungal growth and the significance of turgor in plant infection". Molecular Genetics of Host-Specific Toxins in Plant Disease: Proceedings of the 3rd Tottori International Symposium on Host-Specific Toxins, Daisen, Tottori, Japan, August 24–29, 1997. Netherlands: Kluwer Academic Publishers. pp. 261–271. ISBN   978-0-7923-4981-5.
  45. "Fungus - Evolution and phylogeny of fungi". Encyclopedia Britannica. Retrieved 2020-05-19.
  46. Wang, Guangyi; Wang, Xin; Liu, Xianhua; Li, Qian (2012). "Diversity and Biogeochemical Function of Planktonic Fungi in the Ocean". Biology of Marine Fungi. Vol. 53. Berlin, Heidelberg: Springer Berlin Heidelberg. p. 71–88. doi:10.1007/978-3-642-23342-5_4. ISBN   978-3-642-23341-8 . Retrieved 2025-08-31.
  47. Peng, Xuefeng; Amend, Anthony S.; Baltar, Federico; Blanco‐Bercial, Leocadio; Breyer, Eva; Burgaud, Gaëtan; Cunliffe, Michael; Edgcomb, Virginia P.; Grossart, Hans‐Peter; Mara, Paraskevi; Masigol, Hossein; Pang, Ka‐Lai; Retter, Alice; Roberts, Cordelia; van Bleijswijk, Judith; Walker, Allison K.; Whitner, Syrena (2024). "Planktonic Marine Fungi: A Review". Journal of Geophysical Research: Biogeosciences. 129 (3). doi: 10.1029/2023JG007887 . ISSN   2169-8953. CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License
  48. Kagami, M., Miki, T. and Takimoto, G. (2014) "Mycoloop: chytrids in aquatic food webs". Frontiers in microbiology, 5: 166. doi : 10.3389/fmicb.2014.00166 . CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 3.0 International License.
  49. Gutierrez MH, Jara AM, Pantoja S (2016) "Fungal parasites infect marine diatoms in the upwelling ecosystem of the Humboldt current system off central Chile". Environ Microbiol, 18(5): 1646–1653. doi : 10.1111/1462-2920.13257.
  50. Frenken, T., Alacid, E., Berger, S.A., Bourne, E.C., Gerphagnon, M., Grossart, H.P., Gsell, A.S., Ibelings, B.W., Kagami, M., Küpper, F.C. and Letcher, P.M. (2017) "Integrating chytrid fungal parasites into plankton ecology: research gaps and needs". Environmental microbiology, 19(10): 3802–3822. doi : 10.1111/1462-2920.13827. CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  51. Hassett, BT; Gradinger, R (2016). "Chytrids dominate arctic marine fungal communities". Environ Microbiol. 18 (6): 2001–2009. Bibcode:2016EnvMi..18.2001H. doi:10.1111/1462-2920.13216. PMID   26754171.
  52. Overy, D.P.; Bayman, P.; Kerr, R.G.; Bills, G.F. An assessment of natural product discovery from marine (sensu stricto) and marine-derived fungi. Mycology 2014, 5, 145–167.
  53. Flewelling, A.J.; Currie, J.; Gray, C.A.; Johnson, J.A. Endophytes from marine macroalgae: Promising sources of novel natural products. Curr. Sci. 2015, 109, 88–111.
  54. Miao, F.; Zuo, J.; Liu, X.; Ji, N. Algicidal activities of secondary metabolites of marine macroalgal-derived endophytic fungi. J. Oceanol. Limnol. 2019, 37, 112–121.
  55. Gnavi, G.; Garzoli, L.; Poli, A.; Prigione, V.; Burgaud, G.; Varese, G.C. The culturable mycobiota of Flabellia petiolata: First survey of marine fungi associated to a Mediterranean green alga. PLoS ONE 2017, 12, e0175941.
  56. 1 2 Kohlmeyer, J.; Volkmann-Kohlmeyer, B. Illustrated key to the filamentous higher marine fungi. Bot. Mar. 1991, 34, 1–61.
  57. Stanley, S.J. Observations on the seasonal occurrence of marine endophytic and parasitic fungi. Can. J. Bot. 1992, 70, 2089–2096.
  58. 1 2 Zuccaro, A.; Schoch, C.L.; Spatafora, J.W.; Kohlmeyer, J.; Draeger, S.; Mitchell, J.I. Detection and identification of fungi intimately associated with the brown seaweed Fucus serratus. Appl. Environ. Microbiol. 2008, 74, 931–941.
  59. 1 2 Lee, S.; Park, M.S.; Lee, H.; Kim, J.J.; Eimes, J.A.; Lim, Y.W. Fungal diversity and enzyme activity associated with the macroalgae, Agarum clathratum. Mycobiology 2019, 47, 50–58.
  60. Balabanova, L.; Slepchenko, L.; Son, O.; Tekutyeva, L. Biotechnology potential of marine fungi degrading plant and algae polymeric substrates. Front. Microbiol. 2018, 9, 15–27.
  61. Stanley, S.J. Observations on the seasonal occurrence of marine endophytic and parasitic fungi. Can. J. Bot. 1992, 70, 2089–2096. [Google Scholar] [CrossRef]
  62. Zuccaro, A.; Summerbell, R.C.; Gams, W.; Schroers, H.-J.; Mitchell, J.I. A new Acremonium species associated with Fucus spp., and its affinity with a phylogenetically distinct marine Emericellopsis clade. Stud. Mycol. 2004, 50, 283–297.
  63. Kohlmeyer, J.; Kohlmeyer, E. Marine Mycology: The Higher Fungi; Elsevier: Amsterdam, the Netherlands, 2013.
  64. Wainwright, B.J.; Bauman, A.G.; Zahn, G.L.; Todd, P.A.; Huang, D. Characterization of fungal biodiversity and communities associated with the reef macroalga Sargassum ilicifolium reveals fungal community differentiation according to geographic locality and algal structure. Mar. Biodivers. 2019, 49, 2601–2608.
  65. 1 2 3 Moss, Stephen T. (1986). The biology of marine fungi. CUP Archive. pp. 65–70. ISBN   9780521308991.
  66. Osterhage C.; R. Kaminsky; G. König & A. D. Wright (2000). "Ascosalipyrrolidinone A, an Antimicrobial Alkaloid, from the Obligate Marine Fungus Ascochyta salicorniae". Journal of Organic Chemistry. 65 (20): 6412–6417. doi:10.1021/jo000307g. PMID   11052082.
  67. Grossart, Hans-Peter; Van den Wyngaert, Silke; Kagami, Maiko; Wurzbacher, Christian; Cunliffe, Michael; Rojas-Jimenez, Keilor (2019). "Fungi in aquatic ecosystems". Nature Reviews Microbiology. 17 (6): 339–354. doi: 10.1038/s41579-019-0175-8 . ISSN   1740-1526 . Retrieved 2025-08-31.
  68. 1 2 Grossart, Hans-Peter; Rojas-Jimenez, Keilor (2016). "Aquatic fungi: targeting the forgotten in microbial ecology". Current Opinion in Microbiology. 31: 140–145. doi:10.1016/j.mib.2016.03.016 . Retrieved 2025-08-31.
  69. Wang, Yaqiong; Sen, Kalyani; He, Yaodong; Xie, Yunxuan; Wang, Guangyi (2019). "Impact of environmental gradients on the abundance and diversity of planktonic fungi across coastal habitats of contrasting trophic status". Science of The Total Environment. 683: 822–833. doi:10.1016/j.scitotenv.2019.05.204 . Retrieved 2025-08-31.
  70. Burgaud G, Edgcomb VP, Hassett BT et al (2022) "Marine fungi". In: Stal LJ and Cretoiu MS (ed) The marine microbiome: The microbiomes of humans, animals, plants, and the environment. Berlin, pp 243–295
  71. Banchi, Elisa; Manna, Vincenzo; Muggia, Lucia; Celussi, Mauro (2024). "Marine Fungal Diversity and Dynamics in the Gulf of Trieste (Northern Adriatic Sea)" (PDF). Microbial Ecology. 87 (1). doi: 10.1007/s00248-024-02394-z . ISSN   0095-3628. PMC   11133070 . PMID   38806848 . Retrieved 2025-08-28. CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License
  72. Fryar, S.C., Haelewaters, D. and Catcheside, D.E. (2019) "Annabella australiensis gen. & sp. nov.(Helotiales, Cordieritidaceae) from South Australian mangroves". Mycological Progress, 18(7): 973–981. doi:10.1007/s11557-019-01499-x.
  73. Silliman B. R. & S. Y. Newell (2003). "Fungal farming in a snail". PNAS. 100 (26): 15643–15648. Bibcode:2003PNAS..10015643S. doi: 10.1073/pnas.2535227100 . PMC   307621 . PMID   14657360.
  74. Disease Analysis in San Juan Archipelago Archived 2013-09-12 at the Wayback Machine Friday Harbor Laboratories Seagrass Lab. Retrieved 2012-02-06.
  75. Short, Frederick T.; Robert G. Coles (2001-11-06). Global seagrass research methods. Elsevier. p. 414. ISBN   9780080525617.
  76. Spribille, Toby; Tuovinen, Veera; Resl, Philipp; Vanderpool, Dan; Wolinski, Heimo; Aime, M. Catherine; Schneider, Kevin; Stabentheiner, Edith; Toome-Heller, Merje; Thor, Göran; Mayrhofer, Helmut; Johannesson, Hanna; McCutcheon, John P. (29 July 2016). "Basidiomycete yeasts in the cortex of ascomycete macrolichens". Science. 353 (6298): 488–492. Bibcode:2016Sci...353..488S. doi:10.1126/science.aaf8287. PMC   5793994 . PMID   27445309.
  77. Freshwater and marine lichen-forming fungi Retrieved 2012-02-06.
  78. 1 2 Yuan X, Xiao S, Taylor TN (2005). "Lichen-like symbiosis 600 million years ago". Science. 308 (5724): 1017–1020. Bibcode:2005Sci...308.1017Y. doi:10.1126/science.1111347. PMID   15890881. S2CID   27083645.
  79. Sanders, W. B.; Moe, R. L.; Ascaso, C. (2004). "The intertidal marine lichen formed by the pyrenomycete fungus Verrucaria tavaresiae (Ascomycotina) and the brown alga Petroderma maculiforme (Phaeophyceae): thallus organization and symbiont interaction – NCBI". American Journal of Botany. 91 (4): 511–22. doi:10.3732/ajb.91.4.511. hdl: 10261/31799 . PMID   21653406.
  80. "Mutualisms between fungi and algae – New Brunswick Museum". Archived from the original on 18 September 2018. Retrieved 4 October 2018.
  81. Miller, Kathy Ann; Pérez-Ortega, Sergio (May 2018). "Challenging the lichen concept: Turgidosculum ulvae – Cambridge" . The Lichenologist. 50 (3): 341–356. doi:10.1017/S0024282918000117. S2CID   90117544. Archived from the original on 7 October 2018. Retrieved 7 October 2018.
  82. Rybalka, N.; Wolf, M.; Andersen, R. A.; Friedl, T. (2013). "Congruence of chloroplast – BMC Evolutionary Biology – BioMed Central". BMC Evolutionary Biology. 13: 39. doi: 10.1186/1471-2148-13-39 . PMC   3598724 . PMID   23402662.
  83. Retallack G.J. (2015). "Acritarch evidence of a late Precambrian adaptive radiation of Fungi" (PDF). Botanica Pacifica. 4 (2): 19–33. doi: 10.17581/bp.2015.04203 . Archived (PDF) from the original on 22 December 2016. Retrieved 22 December 2016.
  84. Retallack GJ. (2007). "Growth, decay and burial compaction of Dickinsonia, an iconic Ediacaran fossil". Alcheringa: An Australasian Journal of Palaeontology. 31 (3): 215–240. Bibcode:2007Alch...31..215R. doi: 10.1080/03115510701484705 . S2CID   17181699.
  85. Retallack GJ. (1994). "Were the Ediacaran Fossils Lichens?". Paleobiology. 20 (4): 523–544. Bibcode:1994Pbio...20..523R. doi:10.1017/s0094837300012975. JSTOR   2401233. S2CID   129180481.
  86. Switek B (2012). "Controversial claim puts life on land 65 million years early" . Nature. doi:10.1038/nature.2012.12017. S2CID   130305901. Archived from the original on 1 January 2013. Retrieved 2 January 2013.
  87. Retallack, G.J.; Dunn, K.L.; Saxby, J. (2015). "Problematic Mesoproterozoic fossil Horodyskia from Glacier National Park, Montana, USA". Precambrian Research. 226: 125–142. Bibcode:2013PreR..226..125R. doi:10.1016/j.precamres.2012.12.005.
  88. Retallack, G.J.; Krull, E.S.; Thackray, G.D.; Parkinson, D. (2013). "Problematic urn-shaped fossils from a Paleoproterozoic (2.2 Ga) paleosol in South Africa". Precambrian Research. 235: 71–87. Bibcode:2013PreR..235...71R. doi:10.1016/j.precamres.2013.05.015.
  89. Nelsen, Matthew P.; Lücking, Robert; Boyce, C. Kevin; Lumbsch, H. Thorsten; Ree, Richard H. (January 2020). "No support for the emergence of lichens prior to the evolution of vascular plants" . Geobiology. 18 (1): 3–13. Bibcode:2020Gbio...18....3N. doi:10.1111/gbi.12369. ISSN   1472-4677. PMID   31729136. S2CID   208034624.
  90. Selosse, Marc-André; Le Tacon, François (2001). "Les stratégies symbiotiques de conquête du milieu terrestre par les végétaux". L'Année Biologique (in French). 40: 3–20. doi:10.1016/s0003-5017(01)72082-x.
  91. Gil-Turnes, M. Sofia & William Fenical (1992). "Embryos of Homarus americanus are Protected by Epibiotic Bacteria". The Biological Bulletin. 182 (1): 105–108. doi:10.2307/1542184. JSTOR   1542184. PMID   29304709.
  92. Gaydos, Joseph K.; Balcomb, Kenneth C.; Osborne, Richard W.; Dierauf, Leslie. A Review of Potential Infectious Disease Threats to Southern Resident Killer Whales (Orcinus orca) (PDF) (Report). Archived from the original (PDF) on 2016-03-04. Retrieved 2012-09-28.
  93. Fungal infections of farmed salmon and trout Archived 2012-10-24 at the Wayback Machine Retrieved 2012-02-06.
  94. Kilias, Estelle S.; Junges, Leandro; Šupraha, Luka; Leonard, Guy; Metfies, Katja; Richards, Thomas A. (2020). "Chytrid fungi distribution and co-occurrence with diatoms correlate with sea ice melt in the Arctic Ocean". Communications Biology. 3 (1): 183. doi:10.1038/s42003-020-0891-7. PMC   7174370 . PMID   32317738. S2CID   216033140. CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  95. Hassett, B. T.; Gradinger, R. (2016). "Chytrids dominate arctic marine fungal communities". Environmental Microbiology. 18 (6): 2001–2009. Bibcode:2016EnvMi..18.2001H. doi:10.1111/1462-2920.13216. ISSN   1462-2920. PMID   26754171.
  96. 1 2 3 4 Rédou, Vanessa; Navarri, Marion; Meslet-Cladière, Laurence; Barbier, Georges; Burgaud, Gaëtan (2015-05-15). Brakhage, A. A. (ed.). "Species Richness and Adaptation of Marine Fungi from Deep-Subseafloor Sediments". Applied and Environmental Microbiology. 81 (10): 3571–3583. Bibcode:2015ApEnM..81.3571R. doi:10.1128/AEM.04064-14. ISSN   0099-2240. PMC   4407237 . PMID   25769836.
  97. 1 2 Mouton, Marnel; Postma, Ferdinand; Wilsenach, Jac; Botha, Alfred (August 2012). "Diversity and Characterization of Culturable Fungi from Marine Sediment Collected from St. Helena Bay, South Africa". Microbial Ecology. 64 (2): 311–319. Bibcode:2012MicEc..64..311M. doi:10.1007/s00248-012-0035-9. ISSN   0095-3628. PMID   22430506. S2CID   9724720.
  98. Shoun, H.; Tanimoto, T. (1991-06-15). "Denitrification by the fungus Fusarium oxysporum and involvement of cytochrome P-450 in the respiratory nitrite reduction". The Journal of Biological Chemistry. 266 (17): 11078–11082. doi: 10.1016/S0021-9258(18)99130-1 . ISSN   0021-9258. PMID   2040619.
  99. Filee, J.; Tetart, F.; Suttle, C. A.; Krisch, H. M. (2005). "Marine T4-type bacteriophages, a ubiquitous component of the dark matter of the biosphere". Proceedings of the National Academy of Sciences. 102 (35): 12471–12476. Bibcode:2005PNAS..10212471F. doi: 10.1073/pnas.0503404102 . ISSN   0027-8424. PMC   1194919 . PMID   16116082.
  100. University of Tennessee at Knoxville (25 September 2018). "Study: Microbial dark matter dominates Earth's environments". Eurekalert! (Press release). doi:10.1128/mSystems.00055-18 . Retrieved 26 September 2018.
  101. Lücking, Robert; Hawksworth, David L. (2018). "Formal description of sequence-based voucherless Fungi: promises and pitfalls, and how to resolve them". IMA Fungus. 9 (1): 143–165. doi: 10.5598/imafungus.2018.09.01.09 . ISSN   2210-6359. PMC   6048566 . PMID   30018876 . Retrieved 2025-08-28.
  102. Nilsson, R. Henrik; Ryberg, Martin; Wurzbacher, Christian; Tedersoo, Leho; Anslan, Sten; Põlme, Sergei; Spirin, Viacheslav; Mikryukov, Vladimir; Svantesson, Sten; Hartmann, Martin; Lennartsdotter, Charlotte; Belford, Pauline; Khomich, Maryia; Retter, Alice; Corcoll, Natàlia; Gómez Martinez, Daniela; Jansson, Tobias; Ghobad-Nejhad, Masoomeh; Vu, Duong; Sanchez-Garcia, Marisol; Kristiansson, Erik; Abarenkov, Kessy (2023-04-10). "How, not if, is the question mycologists should be asking about DNA-based typification". MycoKeys. 96: 143–157. doi: 10.3897/mycokeys.96.102669 . ISSN   1314-4049. PMC   10194844 . PMID   37214179.
  103. Naqvi, Syed Atif Hasan; Abbas, Aqleem; Hasnain, Ammarah; Bilal, Zeshan; Hakim, Fahad; Shabbir, Muhammad; Amin, Ahsan; Iqbal, Muhammad Umer (2025). "Advancing fungal phylogenetics: integrating modern sequencing, dark taxa discovery, and machine learning". Archives of Microbiology. 207 (9). doi:10.1007/s00203-025-04392-2. ISSN   0302-8933 . Retrieved 2025-08-28.
  104. 1 2 Moghadamtousi, Soheil; Nikzad, Sonia; Kadir, Habsah; Abubakar, Sazaly; Zandi, Keivan (2015-07-22). "Potential Antiviral Agents from Marine Fungi: An Overview". Marine Drugs. 13 (7): 4520–4538. doi: 10.3390/md13074520 . ISSN   1660-3397. PMC   4515631 . PMID   26204947.
  105. 1 2 Deshmukh, Sunil K.; Prakash, Ved; Ranjan, Nihar (2018-01-05). "Marine Fungi: A Source of Potential Anticancer Compounds". Frontiers in Microbiology. 8: 2536. doi: 10.3389/fmicb.2017.02536 . ISSN   1664-302X. PMC   5760561 . PMID   29354097.
  106. Zhou, Songlin; Wang, Min; Feng, Qi; Lin, Yingying; Zhao, Huange (December 2016). "A study on biological activity of marine fungi from different habitats in coastal regions". SpringerPlus. 5 (1): 1966. doi: 10.1186/s40064-016-3658-3 . ISSN   2193-1801. PMC   5108748 . PMID   27933244.
  107. Xu, Lijian; Meng, Wei; Cao, Cong; Wang, Jian; Shan, Wenjun; Wang, Qinggui (2015-06-02). "Antibacterial and Antifungal Compounds from Marine Fungi". Marine Drugs. 13 (6): 3479–3513. doi: 10.3390/md13063479 . ISSN   1660-3397. PMC   4483641 . PMID   26042616.

Further reading