Marine Mycology: An Overview of Pathogens, and Secondary Metabolites
Introduction and History The golden age of marine mycology occurred from 1960-1990 with the research and discovery of most of the roughly 500 species of obligate marine fungi. Much of said research was conducted from 1980-2000; this 30 year period saw the report of nearly half of the marine fungal species currently known (Jones et al. 2009; Jones, 2011). That being said, marine fungi are vastly understudied and under rated in comparison to marine plants, animals, and other microorganisms; frequently they are omitted or only briefly referenced in marine biodiversity and ecology text books (Jones and Pang, 2012). The cladistics of marine fungi is currently in a state of flux, with new taxa being discovered as molecular techniques such as DNA and RNA analysis via polymerase chain reactions, and gel electrophoresis are implemented (Ald et al 2005). Even though fungus-like organisms such as oomycetes are not fungi, marine mycologists often study them as they perform similar functions, and until recently most had been classified as fungi based on their morphological similarities (Jones, 2011). These fungus-like organisms are eukaryotic, heterotrophic, zoospores, have chitin containing cell walls, and similar life cycles to fungi (Neuhauser et al. 2012). Conventionally terrestrial or freshwater species are also included in the marine fungal group as facultative species; this is due to their active ecological role in the marine, and estuarine environment. Here is broad, but accepted definition for obligate, and facultative marine fungi from Kohlmeyer, 1979 "obligate marine fungi are those that grow and sporulate exclusively in a marine or estuarine (brackish water) habitat; facultative marine are fungi from freshwater or terrestrial areas able to grow also in the natural marine environment". Due to the expansive range of organisms studied as, or in conjunction with marine fungi, it is common place to describe the entirety of the group as "marine derived". This umbrella-term is applicable based on these organisms presence within the marine habitat, and their morphological resemblance to, or similar ecological niche of true marine fungi (Jones, 2011). Marine Derived Fungal Phyla | Unikonts (Basal to Fungi) | True Fungi | Fungal-Like | Cryptomycota | Ascomycota | Oomycota | | Basidomycota | Hyphochytriomycota | Mesomycetozoea | Blastocladiomycota | Labyrinthulomycota | | Chytridiomycota | Phytomyxea | | Neocallimastigomycota | | Table 1

Table 1 The currently recognized phyla of fungi, and fungal-like organisms that occur in the marine environment (Jones, 2012; Neuhauser, 2012). Marine derived fungal organisms are ubiquitous in our world's oceans, but are usually found in tropical and subtropical regions. Water temperature is the most important physical factor determining fungal distribution within the marine environment (Hughes, 1986). Marine fungi have been found in the deep ocean, hydrothermal vent systems, hypersaline seas, anoxic environments, methane hydrate cold seeps (Bhadury et al. 2006; Bugni and Ireland 2004), Antarctic waters, and in Arctic waters (Fell, 2011). Marine derived fungi can be classified into three general ecological functions, saprophobic, parasitic, and mutualistic (Buée et al. 2009). Saprophobic fungi play a key role in nutrient cycling, especially in estuarine environments, and have been extensively studied in mangrove forests as well as on driftwood (Kohlmeyer and Kohlmeyer, 1986). Mutualistic relationships between endophytic fungi and their hosts has also been studied a great deal, as they represent a source of potentially bioactive secondary metabolites (Chen et al 2007). Parasitic and pathogenic marine fungi have been well studied as symptoms of the host organism, mainly because of mortality events affecting economically valuable species (Lightner and Sinderman, 1988).
Pathogens, Parasites, and Diseases Caused by Marine Derived Fungi An expanse of different marine fungi parasitize marine animals, plants, and algae with varying degrees of severity (Porter, 1986). A well explored facet of pathogenic fungi is their potent effect on mariculture systems, and ability to destroy entire crops of valuable food organism. Due to an economic interest in discovering the causative agent behind expensive mortality events, fungal pathogens affecting mariculture are well studied and documented. On the other hand, fungal agents causing disease in wild animals are less studied, but there are documented cases of mortality events in natural populations as well (Sarmiento‐Ramírez, et al. 2010).
Application in Mariculture The aquaculture and mariculture industries have been subject to considerable economic loss via devastating mortality events of cultured organisms caused by fungal pathogens (Wang et al. 2007) Cultivation of aquatic organisms for food purposes is an essential socioeconomic industry that helps feed the world's population, and in some cases attempt to ease stress from overfishing our global oceans. Disease causing fungal organisms in the marine environment are more or less studied as functions of their host species and the economical importance of their survival, therefore a wealth of research surrounding fungal pathogens and crop species is available (Alderman and Polglase, 1986). Disease is the leading cause of damage to livestock in aquaculture, and fungal diseases are second only to bacterial ones (Ramaiah, 2006). Fungal agents can be extremely virulent, destroying harvests in a matter of weeks, in some cases. Coupled with the high mortality rates of many fungal diseases, there is severe lack of available preventative and therapeutic methods to reduce outbreaks, therefore the destruction of economically significant organisms is a persistent problem. Compounding this effect, the occurrence of severe fungal disease is on the rise as more farmers employ high density, intensive culture techniques. The ascomycete yeast Metschnikowia bicuspidata is known to cause disease in several cultured arthropod species. In the brine shrimp Artemia salina, this yeast exhibits an interesting and unique predatory behavior; it fires its acsospores as a method of infection rather than dispersal. The ascospores have a spear shaped head and are fired from the gastrointestinal system into the body cavity of the shrimp so the yeast can colonize the rest of the organism (Lachance et al., 1976). As it is not a picky pathogen and can infect many kinds of hosts, since 2001 M. bicuspidata has been devastating the culture populations of the crab Portunus trituberculatus in China, causing a fatal illness known as Milky disease. The infection causes abnormalities within the heart, muscles and hepatopancreas of the crab's body, and can kill a healthy adult crab in four days. (Wang et al. 2007). The oomycete Lagenidium spp. is another generalized parasite of marine animals, and generally infect the larval stages of crustaceans like Penaesus spp. (Lightner and Fontinae, 1973), the blue crab Callinectes sapidus (Couch, 1942), the Dungeness crab Cancer magister (Armstrong et al. 1976), the American lobster Homarus americanus (Nilson et al., 1976) and many others. Lagenidium callinectes not host specific, and can affect the protozoeal and mysis stages of Penaesus spp. larvae, as well as eggs. The efficiency with which this fungal agent kills penaeid larvae is astounding; once infected, 20-100% of the population die within 48-72 hours (Lightner 1988). The fungus spreads throughout the larval thallus, converting internal tissues to hyphae. When the larva is dead, Lagenidium callinectes grow sporangial discharge tubes through the carapace. The tubes form vesicles that in turn release zoospores capable of infecting other larval crustaceans (Lightner and Fontaine, 1973). Healthy eggs have a commensal, antifungal-producing bacterium coating the surface of the eggs, therein resisting fungal attack. Poorly managed cultivation systems, unsavory water quality, and other stressors can decrease the abundance of the helpful Alteromonas spp. and increase the susceptibility of the eggs and larvae to Lagenidium invasion (Fisher, 1983).
Fungi Affecting Animals in the Natural Environment The incubating eggs of sea turtles are also at great risk for fungal infection and mortality. In studies conducted from 2005-2012, eight important nesting sites in the Atlantic, Indian, and Pacific Oceans were surveyed for the presence of Fusarium spp. In all nesting sites and turtle species surveyed, eggs colonized by Fusarium spp. were found (Sarmiento‐Ramírez et al. 2014).
On Boavista Island, Cape Verde, a turtle nesting site with low hatch rates, Fusarium solani has recently been documented as a major pathogen of the loggerhead sea turtle eggs, Caretta caretta. The mortality rate of the infected nests being 83% in challenge experiments. It has been determined that the developing embryos come into contact with the pathogen in the sand, as the fungi was absent in samples taken from the nesting female's ovipositor and cloacal mucus (Sarmiento‐Ramírez et al. 2014). Interestingly enough, the presence F. solani has also been documented in asymptomatic nests, which suggests that other factors such as microclimatic condition, sand composition, and additional immunosuppression of developing eggs may also play a key role in the infection of nests (Sarmiento‐Ramírez et al. 2010). In subsequent genetic analyses, Sarmiento‐Ramírez et al, teased out two distinct species from Fisarium spp.: Fusarium falciforme and Fusarium keratoplasticum as the causative agents affecting sea turtle, specifically C. Caretta, nest mortality across the globe (Sarmiento-Ramierz et al. 2014). Figure 2. Dead C. caretta embryo infected by F. solani. (Sarmiento-Ramierz et al. 2014)
Just as in other pathogenic species of Fusarium spp.,
F. falciforme and F. keratoplasticum are virulent, opportunistic, and generalists. The optimal growth temperature of the two species of fungi and the critical incubation temperature for sexual determination in the eggs overlap significantly, therefore optimal conditions for egg development and fungal growth occur at the same temperature. This suggests an adaptation to the host environment and some degree of host specificity in these two pathogenic species (Sarmiento-Ramierz et al. 2014). There was also an observable effect of substrate type and tidal inundations on fungal infestation of the nests. Those nests laid in sand without frequent tidal inundations experienced a much lower mortality rate (21.4%), as opposed to those laid clay/silt substrates with or without frequent tidal inundations (71.8% and 78.4% respectively). It seems that nests exposed to tidal inundations and the presence of clay/silt are equally at risk for high mortality by Fusarium spp. This indicates that sea turtle egg mortality by fungal pathogens is only going to increase as anthropogenic destruction of nesting sites forces females to nest outside of their preferred location and substrate type (Sarmiento-Ramierz et al. 2014).
Applications of Marine Derived Fungi in Biotechnology Marine fungi are known to be an extremely bioactive group capable of producing secondary metabolites to interact with the organisms of the microbial world in which they live (Miller 1986). Terrestrial fungi have been utilized by the biotechnology industry since the advent of penicillin, and in the past two decades much research has been conducted on marine fungi concerning the extract and applications of these secondary metabolites. Marine derived fungi are emerging as a source for enzymes, anti-fouling, antibacterial, antiviral, anticancer, anti-inflammatory, and anti-plasmodial agents (Bhadury et al. 2006; Bugni and Ireland 2004). Marine fungi from extreme environments are also emerging as a promising supply of bioactive compounds, due to their ability to thrive in the harsh conditions in which they are found.
Antibacterial Agents from Marine Fungi It is common knowledge that humanity is facing an antibacterial crisis; bacteria are becoming resistant to common treatments and new drugs are not being developed fast enough (Coates et al. 2002). The observable evolution of bacterial strains and subsequent resilience to drug therapy is producing a growing number of diseases that are untreatable, and people are dying as a result of infections that used to be simple, and easily eradicated. Often times bacteria and fungi live in conjunction with one another, competing for the same resources; as a function of this, the two have been fighting chemical warfare with one another since the evolution of fungi (Janzen, 1977; Wicklow 1981). Therefore, it comes as no surprise that fungi and bacteria have developed chemical methods of deterring competition for resources with one another (Bennett 1998). Many researchers have screened species of marine derived fungi and found them to have antibiotic properties ranging from mildly to tremendously effective. This has lead to the belief that chemicals obtained from marine fungi, especially from those living in extreme environments, will become an excellent and invaluable source for the pharmaceutical industry to synthesize new, life saving antibiotics (Bhadury et al. 2006). In 2001, Cueto at al. collected the brown alga Rosenvingea sp. in the Bahamas Islands, and found an undescribed fungi, a member of Pestalotia spp. from its surface; they then isolated Pestalone and showed it to have significant antibiotic effects against several resistant strains of disease causing bacteria. Pestalone is a benzophenone compound, an aromatic ring with extended chromophore, and was isolated as a yellow crystal. The bioactive compound was shown to strongly affect vancomycin-resistant Enterococcus faecium and methicillin-resistant Staphylococcus aureus. Interestingly, Pestalotia spp. did not produce the antibacterial Pestalone when cultured alone in control experiments, only when the fungi and bacteria were allowed to grow together did Pestalotia spp. produce its antibacterial compound. The occurrence of Pestalone is a product of fungal biosynthesis in response to external stimulus, this was proven when Pestalone production could be induced by the addition of 1% ethanol after 24 hours of incubation. The induced production of Pestalone also disproves the notion that the antibacterial compound was created by the transformation of a bacterial metabolite by the fungus in the mixed fermentation experiment. The presence of the bacterial antagonists also increased the yield of Pestalone maufacture by the fungus. It is clearly described by the experiments conducted on the fungi Pestalotia spp that the biomixing of a microbe and its antagonist could lead to the discovery of new antibiotics and other products resultant of secondary metabolites. In conjunction with facilitating discovery of bioreactive agents, biomixing can also increase the yield of said compounds up to 400 fold (Cueto at al. 2001).
Antifouling Agents from Marine Derived Fungi Biofouling of man-made structures in the marine environment is costly, both economically and environmentally; there is a growing need for natural antifouling agents and marine fungal derivatives are proving to be a hopeful source for new antifouling products. Biofouling is the growth of sessile organisms on the surface of structures like tanker ships, mariculture facilities, bridge pylons, and sea water pipelines in the marine environment (Dobretsov et al. 2006; Gerhart et al. 1988). Organisms like barnacles, bivalves, and algae grow on and damage the surfaces of these objects. To prevent their growth and subsequent repair costs, over the last few decades, antifouling coatings and paints have been implemented to inhibit the colonization of these organisms. Unfortunately many antifoulding agents such as tributyltin (TBT)-based compounds, are toxic and are causing damage to surrounding ecosystems (Alzieu 2000). Marine fungal isolates and secondary metabolites have been shown to have antifouling properties (Qi et al. 2009). A scant few studies have endeavored to test marine derived fungi for antifouling agents, but the preliminary results are positive in nature. Qi et al. conducted a study in 2009 that tested the marine derived fungi Cladosporium sp. and found it to have both micro and macro antifouling properties. They isolated a total of nine compounds, " two hexaketides, two 12-membered macrolides, three benzene-type compounds, and two diketopiperazines," (Qi et al. 2009). A ketide is organic compound containing adjacent methylene and carbonyl functional groups, and a hexaketide is a six membered ketide (Clark et al 1995). These isolates induced a dramatic degenerative response to three common bacterial fouling organisms, Loktanella hongkongensis Micrococcus luteus and Ruegeria sp. as well as in the larval forms of Balanus amphitritei, the acorn barnacle, and Bugular neritina, a bryozoan. These preliminary findings support the hypothesized ability of marine fungi as sources of bioactive antifouling agents in the future, as well as suggest fungi containing hexaketides are good sources for these agents and should be investigated (Qi et al. 2009).

Secondary Metabolites from Extremophile Marine fungi In recent years, much attention and research has been devoted to organisms that can survive in seemingly lethal environments due to the possible application of their isolates in biotechnology and industry; marine fungi found in these environments are an exception to this research trend. Researchers looking for marine fungi derived secondary metabolites is increasing in these environments, but is still scant (Demare et al. 2006; Galkievicz et al. 2012). Environments in which extremophilic marine fungi are found include, deep sea hydrothermal vent communities, and cold water methane seeps (Lai et al. 2007). The deep sea environment is unforgiving, with extreme hydrostatic pressures bearing down on organisms in the absence of light, and extremely cold temperatures. Despite this, microbial organisms seem to grow and thrive in some deep sea communities (Nagano and Nagahama, 2012). The application of products derived from extremophile marine fungi are many. These include enzymes that are cold tolerant, heat tolerant, active at immense hydrostatic pressures, and alkaline tolerant (Kumar and Takagi 1999; Marine fungi found in the deep sea are sources of secondary metabolites that are cold tolerant, and able to function at extremely high hydrostatic pressures, many of them are also alkaline tolerant. An interesting example of one such bioactive compound with possible uses in industry is a cold water, alkaline tolerant protease. Proteases are used in the dairy industry, whereas alkaline proteases are necessary in the detergent and leather industry (Ohgiya et al. 1999). One study found 221 fungal isolates from sediments in the Central Indian Basin at a depth of 5,000m and screened them for enzymes. Many grew and produced the desired alkaline tolerant proteases, but the most successful was Aspergillus ustu (NIOCC #20). The protease was not affected by the addition of metals Cu, Hg, Fe, Ni and Zn ions, and retained 10% of its activity at 2 ° C. Furthermore, the enzyme was active at combinations of 5°C, 30°C and 300 bar pressure, 50 bar pressure (Damare et al. 2006). These characteristics make this fungal isolate valuable in the detergent industry as it can be used as an additive for low temperature wash (Ohgiya et al. 1999).
Conclusion and Discussion
Gaps in Our Knowledge and Prospects for the Future The past two decades have seen a sharp decline in the number of practicing mycologists and labs conducting research, and applied research is favored over the fundamental. The study of their ecological functions has taken a back seat to the pursuance of their possible economic uses. This is in part due to the difficulty faced by researchers in gaining funding for projects, it is simply easier to receive grant money if the proposed project has an economically based goal in mind. Thus, the number of active mycologists and labs delving into the fundamental study of marine fungi has rapidly decreased since the bloom of information discovered in the 1980's and 90's. The inclusion of fungal-like organisms in marine mycology books may in part be due to how recently the taxonomy of marine fungi has been revised, many of the scientist active today came up under the old system of higher and lower marine fungi. The slow pace at which the fundamental research of marine fungi has progressed can be explained by how difficult it can be to culture these organisms in the lab, with certainty that terrestrial spores have not contaminated the samples. This problem was solved by the advent and use of molecular techniques to analyze samples without needing to culture them. Molecular genetic techniques aided in the discovery of new species of marine fungi without researchers having gained the fundamental knowledge of their ecology, or how to culture them (Pang and Jones 2012). This issue has two sides: general knowledge of the number of marine fungi, clades, and their evolutionary relatedness to one another has increased; on the other hand, critical discoveries are potentially being glossed over due to this lack of knowledge. A prime example was presented previously in this paper, Pestalone would not have been discovered if researchers had not known its dietary needs or how to culture it. It also plainly states that fungi and bacteria are ecologically linked, and more fundamental scientific research on the ecology of fungi species can facilitate the discovery of economically and medically important products. It remains to be seen how increasing anthropogenic stress on the world's oceans will affect marine fungi, but if the mortality of Gorgonian sea fans (Alker et al. 2001) and sea turtle eggs are any standard to go by, I believe we will see many more instances of fungal species causing problems in the near future. As fungi are temperature dependent and have an affinity for warmer waters, it only seems a matter of time before another fungal epizootic strikes tropical ecosystems.
Final Thoughts Presented as a review are two facets of marine fungi, the destructive, and the constructive. A broad survey of the known types of parasitism by marine derived fungi (with the omission of algal and plant hosts) and their effects in our world today. Also omitted was the well known relationship between saprobic estuarine fungi, their mangrove or salt marsh hosts and nutrient cycling, as that topic may be better suited for an ecological discussion. The destructive nature of many known species of marine derived fungi is contrasted with the prospective emergence of useful drugs, and products isolated from marine fungal agents. It should not go without mention that three books greatly expanded my knowledge on the subject. Two of these volumes were published in the 80's (The Biology of Marine Fungi and Marine Mycology the Higher Fungi) and reflect the previous generation of marine mycologists, their work, and what they passed on to the generation of mycologist today. In conjunction, the final volume was published in 2012 (Marine Fungi and Fungi and Fungal-like Organisms) and presented a wonderful dissimilarity between new discoveries and somewhat outdated foundational knowledge coming together in one field.

References
Ald S.M. Adl, A.G.B. Simpson, M.A. Farmer, R.A. Andersen, O.R. Anderson, J.R. Barta, S.S. Bowser, G. Brugerolle, R.A. Fensome, S. Fredericq, T.Y. James, S. Karpov, P. Kugrens, J. Krug, C.E. Lane, L.A. Lewis, J. Lodge, D.H. Lynn, D.G. Mann, R.M. McCourt, L. Mendoza, Ø Moestrup, S.E. Mozley-Standridge, T.A. Nerad, C.A. Shearer, A.V. Smirnov, F.W. Spiegel, M.F.J.R. Taylor (2005). The new higher level classification of eukaryotes with emphasis on the taxonomy of protists. Journal of Eukaryotic Microbiology, 52:399–451
Alderman, D.J. and Polglase, J.L. (1986) Are fungal diseases significant in the marine environment? In: The Biology of Marine Fungi, Moss, S. T. (Ed.). (1986). London, Cambridge University Press. 189-198
Alker, A. P., Smith, G. W., & Kim, K. (2001). Characterization of Aspergillus sydowii (Thom et Church), a fungal pathogen of Caribbean sea fan corals. Hydrobiologia, 460(1-3), 105- 111.
Alzieu C (2000) Impact of tributyltin on marine invertebrates. Ecotoxicology 9:71–76.
Bhadury, P., Mohammad, B., & Wright, P. (2006). The current status of natural products from marine fungi and their potential as anti-infective agents. Journal of Industrial Microbiology and Biotechnology, 33(5), 325-337.
Buée M, Reich M, Murat C, Morin E, Nilsson RH, Uroz S, Martin F,(2009). 454 Pyrosequencing analyses of forest soils reveal an unexpectedly high fungal diversity. New Phytol. 184:449-456.
Bugni TS, Ireland CM (2004) Marine-derived fungi: a chemically and biologically diverse group of microorganisms. Nat Prod Rep 21:143–163.
Clark, W. D., & Crews, P. (1995). A novel chlorinated ketide amino acid, herbamide A, from the marine sponge dysidea herbacea. Tetrahedron Letters, 36(8), 1185-1188.
Coates A, Hu Y, Bax R, Page C (2002) The future challenges facing the development of new antimicrobial drugs. Nat Rev Drug Discov 1:895–910
Cueto M, Jensen PR, Kauffman C, Fenical W, Lobkovsky E, Clardy J (2001) Pestalone, a new antibiotic produced by a marine fungus in response to bacterial challenge. J Nat Prod 64:1444–1446
Damare, S., Raghukumar, C., Muraleedharan, U. D., & Raghukumar, S. (2006). Deep-sea fungi as a source of alkaline and cold-tolerant proteases. Enzyme and Microbial Technology, 39(2), 172-181.
Dobretsov S, Dahms HU, Qian PY (2006) A review: inhibition of biofouling by marine microorganisms and their metabolites. Biofouling 22:43–54.
Fell, J. (2012). 6 Yeasts in marine environments. Marine Fungi: and Fungal-like Organisms, Jones, E. (Ed.) & Pang, K. (Ed.) (2012). Berlin, Boston., 91-102
Fisher, W. S. (1983). Eggs of Palaemon macrodactylus: III. Infection by the fungus, Lagenidium callinectes. The Biological Bulletin, 164(2), 214-226.
Galkievicz JP, Stellick SH, Gray MA, Kellogg CA (2012) Cultured fungal associates from the deep-sea coral Lophelia pertusa. Deep Sea Res I 67:12–20
Gerhart DJ, Rittschof D, Mayo SW (1988) Chemical ecology and the search for marine antifoulants: studies of a predator–prey symbiosis. J Chem Ecol 14:1905–1910.
Hughes, GC. Biogeography and marine fungi. In: The biology of marine fungi. Moss, S. T. (Ed.). (1986). Cambridge, Cambridge University Press: 275-295.
Jones, E.B.G. (2011). Fifty years of marine mycology. Fungal diversity, 50(1), 73-112.
Jones, E.B.G., Sakayaroj, J., Suetrong, S., Somrithipol, S. and Pang, K.L. (2009). Classification of marine Ascomycota, anamorphic taxa and Basidiomycota. Fungal Diversity 35: 1-187.
Jensen PR, Fenical W. Secondary metabolites from marine fungi. In: Hyde KD, editor. Fungi in marine environments, Fungal Diversity Res. Ser. 2002;7:293–315.
Kohlmeyer, J, and Kohlmeyer E. (1979). Introdution. In: Marine mycology. The higher fungi. Academic Press, Inc., 1-5.
Kumar GC, Takagi H. Microbial alkaline proteases: from bioindustrial viewpoint. Biotechnol Adv 1999;17:561–94.
Lachance MA, Miranda M, Miller MW & Phaff HJ (1976) Dehiscence and active spore release in pathogenic strains of yeast Metschnikowia bicuspidata var Australis – possible predatory implication. Can J Microbiol 22: 1756–1761
Lai, X., Cao, L., Tan, H., Fang, S., Huang, Y., & Zhou, S. (2007). Fungal communities from methane hydrate-bearing deep-sea marine sediments in south china sea. Isme j, 1(8), 756- 762.
Liberra K, Lindequist U. Marine fungi-a prolific resource of biologically active natural products. Pharmazie 1995;50:583–8.
Lightner, DV, Fontaine, CT. (1973). A new fungus disease of the white shrimp penaeus setiferus. Journal of Invertebrate Pathology, 22(1), 94-99.
Lightner, DV, & Sinderman, C. J. (1988). Larval mycosis of penaeid shrimp. Disease diagnosis and control in North American marine aquaculture., (Ed. 2), 58-63.
Nagano Y, Nagahama T (2012) Fungal diversity in deep-sea extreme environments. Fungal Ecol 5:463–471
Neuhauser S, Glockling S, Leano S, Lilje O. (2012) An introduction to fungus-like microorganisms. Marine Fungi. and Fungal-like Organisms. Jones, E. (Ed.) & Pang, K. (Ed.) (2012). Berlin, Boston., 137-152.
Ohgiya S, Hoshino T, Okuyama H, Tanaka S, Ishizaki K. (1999). Biotechnology of enzymes from cold-adapted microorganisms. Biotechnological applications of cold-adapted organisms. Springer-Verlag; 1999. p. 17–34.
Pang, KL, Jones, E.B.G. (2012) Epilogue: importance and impact of marine mycology. In: Marine Fungi. and Fungal-like Organisms. Jones, E. (Ed.) & Pang, K. (Ed.) (2012). Berlin, Boston., 509-517
Porter D. (1986). Mycoses of marine organisms. In: The biology of marine fungi. Moss, S. T. (Ed.). (1986). London, Cambridge University Press. 141-153.
Ramaiah, N. (2006). A review on fungal diseases of algae, marine fishes, shrimps and corals. Indian Journal of Marine Sciences, 35(4), 380-387
Sarmiento‐Ramírez, J. M., Abella, E., Martín, M. P., Tellería, M. T., López‐Jurado, L. F., Marco, A., & Diéguez‐Uribeondo, J. (2010). Fusarium solani is responsible for mass mortalities in nests of loggerhead sea turtle, Caretta caretta, in Boavista, Cape Verde. FEMS microbiology letters, 312(2), 192-200.
Sindermaun, C. J. 1974. Diagnosis and control of mariculture diseases in the United States. Technical Series, No. 2., National Marine Fisheries SnVice, U.S. Dept. of Commerce, Washington, DC.
Sindennann, C. J. 1977. Disease Diagnosis and Control in North American Aquaculture. Elsevier, New York.
Qi, S., Xu, Y., Xiong, H., Qian, P., & Zhang, S. (2009). Antifouling and antibacterial compounds from a marine fungus cladosporium sp. F14. World Journal of Microbiology and Biotechnology, 25(3), 399-406.
Yu VB, Verigina NS, Sova VV, Pivkin MV, Zvyagintseva TN.(2003). Filamentous marine fungi as producers of O-glucosyl hydrolases: _-1,3 glucanase from Chaetomium indicum. Biotechnol, 5:349–59.