A Review on Biodiversity of Marine and Mangrove Fungi

ISSN: 2347-3215 Volume 1 Number 1 (2013) pp. 26-44 www.journals.excellentpublishers.com

A review on biodiversity of marine and mangrove fungi

T.Sivakumar*

Department of Microbiology, Kanchi Shri Krishna College of Arts and Science, Kilambi 631 551, Kancheepuram, Tamil Nadu, India

*Corresponding author e-mail:[email protected]

K E Y W O R D S A B S T R A C T

Biological diversity; Biological diversity refers the variability among living organisms from all Terrestrial; sources including terrestrial, marine and other aquatic ecosystem and marine; ecological complexes of which they are part. Biodiversity encompasses all aquatic ecosystem; life forms, ecosystems and ecological processes and acknowledges the Marine fungi; hierarchy at genetic, taxon and ecosystem level. The essential ingredients of Mangrove fungi; biodiversity are phenotypic flexibility genetic variation within populations Fungi in water; and ecotypic variations. Microbial biodiversity can be viewed from a variety Sediment; of perspectives, including physiological diversity, interaspecific genetic Fungi on diversity and phylogenetic diversity of species and higher taxa. Microbial different substrates. diversity represents the largest untapped reservoir of biodiversity for potential discovery of new biotechnological products, including new pharmaceuticals, new enzymes, new special chemicals or new organisms that carryout novel process. .

Introduction

Most early studies on fungi colonizing studies on manglicolous fungi. Recent mangroves were taxonomic and confined studies on inertial mangrove fungi have mainly to cataloguing fungi and describing provided information on (a) frequency of new taxa collected in a given area (Cribb occurrence (b) vertical zonation, (c) host and Cribb, 1955; Kohlmeyer and and substratum specificity, (d) succession, Kohlmeyer, 1964 1969, 1971, 1977; and (e) seasonal occurrence (Aleem, 1980; Kohlmeyer, 1966, 1969a, 1981, 1984, Jones et al., 1988; Hyde, 1988a, 1989c, 1985; Kohlmeyer and Schatz, 1985). Until 1990b, 1991; Leong et al., 1991; Poonyth recently, there have been few ecological et al., 1999). Of these, considerable effort

has been spent investigating the frequency work has been done on the taxonomy and of occurrence of manglicolous fungi (Jones ecology of mangrove swamp fungi in India and Tan, 1987; Borse, 1988; Hyde, (Padhye et al., 1967; Pawar, et al., 1963; 1988a,b, 1989a,b,c; Hyde and Jones, 1988; 1965 and Rai and Tewari, 1963). Jones et al., 1988; Jones and Kuthubutheen, 1989; Tan et al., 1989; Tan and Leong, Earliest studies on the ecology of 1990,1992). mangroves fungi Kohlmeyer (1969a) encountered 3 common species of marine Early studies on marine fungi on fungi in the mangrove habitat, namely mangroves have focused on taxonomy of Lulworthia spp (20% of all collections) marine fungi including descriptions of new Leptosphaeria australiensis (15%) and species and new genera, lists of fungi and Phoma species (10%). 100 Mangrove surveys. This includes the marine fungi species; only 8 have been examined for the occurring in mangrove environments. occurrence of marine fungi and the latest Excellent reviews and vast amounts of Island research was conducted in Bermuda information on marine fungi have appeared and collected 15 marine Ascomycetes, 1 in several texts. (Johnson and Sparrow, Basidiomycetes, and 6 Deuteromycetes 1961; Jones, 1976; Moss, 1986; Hyde and (Kohlmeyer and Kohlmeyer, 1977). Aleem Lee, 1995; Jones, 1995; Jones and Alias, (1980) reported that the Ascomycetes; 1996). For several accounts on various Halosphaeria viscidula, Rosellinia sp and aspects of marine fungi the following Torpedospora radiata were frequent on works, among others, are referable mangroves in sierra Leone and also found (Chinnaraj, 1993a; Jones and Tan, 1987; that the Mitosporic taxa. Cirrenalia Hyde and Jones, 1988; Hyde, 1990a; Hyde macrocephala, C. pygmea, C. tropicalis, et al., 1990; Leong et al., 1991; Kohlmeyer Periconia and Zalerion spp were abundant and Kohlmeyer, 1979; and Kohlmeyer et on mangrove wood. Kohlmeyar (1984) also al., 1995). Ecological studies on reported that L. australiensis was a manglicolous fungi are relatively recent i.e. common species of mangroves. Although from late 1980s onwards. A wealth of mangrove fungi of the West coast of India information is now available on different have been well studied, there have been ecological aspects of fungi in mangroves few studies on the East coast (Bay of including frequency of occurrence, vertical Bengal), despite the fact that its mangroves distribution, substrate preference, are more extensive compared to the west succession, seasonal occurrence and host coast. (Untawale, 1987). Hyde and Jones specificity. However these are mainly from (1988) observed the some fungi tend to South East Asia (Hyde and Lee, 1995; occur configuration at certain levels and Jones and Alias, 1996). also reported that the greater species diversity occurred at the mid littoral level. There are umpteen literatures on the ecology and taxonomy of soil fungi Most Booth (1971b) observed on occurrence and of the reports relate to the study of fungal taxonomy of aquatic fungi in saline flora from cultivated agricultural soils, habitats. Hyde (1989a) reported that the uncultivated soil, pasturelands and forest lignicolous materials were collected from 5 soils. However, little is known about the marine locations in Brunei; a rocky head microbial ecology of mangrove swamps. land, a sandy beach, a man made brackish During the past several years, considerable lake, a healthy mangrove and an oil -

polluted mangrove. Higher marine fungi incorporated in biogeographical maps by present were identified and their percentage Jones (1993), Kohlmeyer (1981, 1984), occurrence noted. There were significantly Kohlmeyer and Volkmann Kohlmeyer less diversity and number of fungi in the (1987) and Volkmann - Kohlmeyer and oil-polluted mangrove when compared to Kohlmeyer (1993). Based on the the healthy mangrove. Most attention to distribution in Atlantic Ocean, Indian data has concentrated on assessment of Ocean, South East Asia and Pacific fungal diversity, physiology and Ocean. Hyde and Lee (1995) revised biochemistry (Kohlmeyer and Kohlmeyer, geographical distribution of representative 1993). Hyde and Lee (1995) suggested that mangrove fungi (Halosarphaeria fibrosa, the diversity of marine fungi is greater in H. marina, Lignincola laevis and the tropics and attributed this to mangrove Lulworthia grandispora). Geographical and tree species richness. Jones et al. (1999) seasonal distribution of Asteromycetes recorded all marine fungi from Marine cruciatus, Stigmoidea marina and habitats can be designated as micro fungi Varicosporina ramulosa correlated with the micro habitat predictor model their growth patterns under different appear to the applicable in the marine temperature regimes (Boyd and environments. Kohlmeyer, 1982).

There have however been no efforts to Predominantly mangrove species: the study the marine fungi on mangroves until Ascomycetes Dactylospora haliotrepha, recently when systematic studies on Halorosellinia oceanica, Lignincola leavis, manglicolous fungi in India were initiated. Lulworthia grandispora, Saagaromyces A detailed investigation of fungi on abonnis and Verruculina enalia;the mangroves of west coast was made by Patil basidiomycete Halocyphina villosa and and Borse (1985a,b) and Chinnaraj anamorphic fungi Cirrenalia pygmea and (1993a,b). However vast tracts of Zalerion varium (Kohlmeyer 1984; Jones mangroves on the east coast remained and Alias, 1996; Sarma and Hyde, 2001; virtually unexplored except for the studies Abdel-Wahab and El- Sharouney, 2002; of Ravikumar and Vittal (1996). Jones and Abdel-Wahab, 2005). Other species are more characteristic of open ocean waters: Antennospora quadricornuta, Quantitative data on the occurrence of tropical marine fungi have been published A. salina, Periconia prolifica, Torpedospora radiata, or wood associated by Raghukumar (1973); Koch (1986); with sand: Corollospora maritima, Kohlmeyer (1984); Zainal and Jones (1984, Trichocladium melhae. Mangrove fungi of 1986). However all of these reports were on the east coast of India have been well driftwood in the sea, along with driftwood studied, there have been few studies on the on the mangrove floor or panels belonging east coast (Bay of Bengal), despite the fact to various timbers submerged near jetties. that its mangroves are more extensive compared to the west coast (Untawale, Marine fungi have been classified into three 1987). geographical groups by Kohlmeyer and Kohlmeyer (1979): i) cosmopolitan species; The marine fungi of Hong Kong and ii) temperate water species and iii) Thailand have been studied intensively species from tropical and subtropical over the past 15 years, and include not only waters. Mangrove fungi have been

random collections of drift material, but habitats sampled. Ecological variation over also the exposure and recovery of bait the temporal and spatial dimensions of the samples (exposure of bait in Hong Kong sample may augment diversity because of (Vrijmoed et al., 1986; Sadaba et al., 1995; the increased number of areas, habitats or Abdel- Wahab, 2000; Thailand: seasons included. Hyde (1990c) recognized Pilantanapak et al., 2005), collection of the different in the common species at drift and attached mangrove samples in study sites, a core group of fungi occurring Hong Kong (Abdel-Wahab and El- in the mangrove ecosystem. The Majority Sharouny, 2002; Jones and Vrijmoed, of the species Dactylospora haliotrepha, 2003), Thailand (Hyde et al., 1993; Leptosphaeria avicenniae were also Sakayaroj et al., 2004). Schmidt and reported from Brunei and other tropical Shearer (2004) analysed the geographical mangroves (Hyde 1989a). Alias et al. distribution data published on lignicolous (1995) reported that more than 60 fungal mangrove fungi, and found that different species can be recorded as common to oceans supported varying numbers. The mangrove ecosystems of the West Indo number of fungi at each site varied: Pacific region. Atlantic Ocean: 12-46 per site (14 sites: mean 25.6); Indian Ocean: 12-64 (14: 42.9) Chinnaraj (1993a,b) had earlier reported 63 and the Pacific Ocean: 17-87 (16: 44). The species of higher marine fungi from the Pacific Ocean has the highest recorded Andaman and Nicobar Islands, which are number of fungi, again the result of approximately 1000 km away from the repeated collections over many years: mainland on the East coast. Ravikumar and Hyde (1988c) in Brunei; Jones and Vittal (1996) reported 48 species belonging Kuthubutheen (1989), Alias et al. (1995), to 37 fungal genera on Rhizophora Tan et al. (1989) and Leong et al. (1991) in apiculate at Pichavaram. Singapore, and the greater diversity of mangrove tree species in this region. The As diverse vegetation exists in mangroves, paucity of marine fungi from the Atlantic it is considered as a major niche of fungal has been attributed to low mangrove tree repository. Mangrove fungal diversity is diversity, for example three in Florida dependent on the age of mangrove, mangroves and four in the Bahamas (Jones diversity of mangrove plant species and the and Abdel-Wahab, 2005; Jones and Puglisi, physicochemical features of mangrove 2006). However, more intensive collections habitat (temperature, salinity and tidal yielded 81 species for Florida mangroves range) (Hyde and Jones, 1988; Jones, from 250 collected samples (previously 2000). Twenty-eight mangrove tree species only 28: Jones and Puglisi, 2006) and 112 yielded 120 higher marine fungi (Hyde, for the Bahamas from 600 collected 1990b). Fifty-five mangroves and their samples, where only 31 had previously associates yielded about 200 higher marine been recorded (Jones and Abdel-Wahab, fungi (Jones and Alias, 1996). Rhizophora 2005). apiculata among the mangrove tree species harboured a maximum of 63 higher marine Diversity most simply can be expressed as fungi (Sarma and Vittal, 2000). species richness, that is the number of species (Magurran, 1988). However, since Among the different geographical richness increase in direct relation to locations; South East Asia has been number of individuals, area and variety of sampled most thoroughly (Hyde and Lee,

1995; Jones and Alias, 1996). There seem freshwater Hyphomycetes (Sridhar and to be no discernible difference between Kaveriappa, 1988). mangrove fungi reported in the subtropics as compared to those found in tropical Sarma and Vittal (2000) investigated the areas. Among 900 known marine fungi, fungal diversity of proproots, seedlings and 358 are recorded from the mangrove wood of Rhizophora apiculata and wood, ecosystem (Jones and Alias, 1996; Jones roots and pneumatophores of Avicennia spp and Mitchell, 1996). Out of 54 mangrove in deltaic mangroves of Godavari and tree species and 60 mangrove associate Krishna rivers in the east coast of India. plant species, up to 55 species have been The number of fungi recorded on proproots studied for fungi (Jones and Alias, 1996). (61) was much greater when compared to Studies on mangrove fungi from the Indian wood (24) and seedling (21). Ocean are limited compared to the Atlantic Ocean and Pacific Ocean and South East Prasannarai and Sridhar (2001) reported the Asian region. Although the Indian diversity of marine fungi on intertidal wood peninsula possesses about 6700 km2 of collected from 13 locations in the West mangroves (Natarajan, 1998) only a few coast of India was assessed out of 3327 studies dealt with fungal richness and wood samples scanned, of which, 72%, diversity in Gujarat (Borse et al., 2000; posses sporulating fungi. Altogether 88 Patil and Borse, 2001), Maharashtra (Borse, species belonging to 47 genera were 1988), Karnataka (Ananda and Sridhar, uncounted. The species richness and 2003), Tamil Nadu (Ravikumar and Vittal, diversity was highest in Islands than in 1996) and Andra Pradesh (Sarma and beaches and harbour locations. It has been Vittal, 2000). predicted that Islands adjacent to the West coast of India provide critical habitat for Chandralata (1999) and Raghukumar and marine fungi. Raghukumar (1998) reported adaptation and activity of terrestrial fungi under Borse (2002) reported that the distribution marine/ mangrove ecosystem as and substratum range of 166 species (13 facultatives or indwellers or residents. Labyrinthulomycota, 4 Chytridiomycota, Terrestrial fungi are common in mangrove 20 Oomycota, 1 excluded sp., 120 water and mud (Chowdhery et al., 1982; Ascomycota, 3 Basidiomycota and 23 Garg, 1983), mangrove leaves mitosporic fungi) of marine fungi recorded (Raghukumar et al., 1995), wood (Aleem, so far from India on animal substratum, 1980), standing senescent stems (Sadaba et driftwood, intertidal wood, algae, al., 1995), decomposing mangrove palm mangroves, sea grasses, salt marsh plants (Nypa fruticans) (Hyde and Alias, 2000). and as propagules in the sea foams samples. Terrestrial fungi in deep sea region of Maria and Sridhar (2002) studied the Arabian Sea were recovered (Raghukumar richness and diversity of filamentous fungi and Raghukumar, 1998). Seawater, sea on woody litter of mangrove along the foam and beach soil of Arabian Gulf Coast, West coast of India. Diversity of fungi in Saudi Arabia yielded terrestrial fungi, the roots of mangrove species of West typical marine and freshwater fungi coast of India (Ananda and Sridhar, 2002). (Bokhary et al., 1992). Sampling of the leaf litter from the Nethravathi mangroves, India revealed the occurrence of many

Prasannaraj and Sridhar (2003) reported collected from driftwood and attached fungi from intertidal wood collected from decayed wood of mangrove trees. The four coastal locations of the West coast of holotype number of 15 taxa is from India. Of the 59 taxa identified, 43 Thailand and 33 are new records from the Ascomycetes, 3 Basidiomycetes and 13 country. anamorphic fungi. Hyde and Sarma (2006) Biodiversity and ecology of higher filamentous fungi on Detritus and live parts of mangrove Nypa fruticans in Brunei were examined vegetation have surveyed for the during 1999. Forty-six taxa were recorded occurrence of higher fungi. In recent past including 33 ascomycetes and 13 (up to 2000), 625 fungi encompassing 279 anamorphic taxa in 25 genera. Linocarpon Ascomycetes, 277 mitosporic fungi, 29 was the most species genus (6 species) Basiomycetes, 3 Chytridiomycetes, 3 followed by Aniptodera and Myxomycetes, 14 Oomycetes, 9 Astrosphaeriella (4 each). More diversity Thraustochytris and 12 Zygomycetes have was found on fronds than on leaves. been reported from mangrove forests Linocarpon appendiculatum, L. bipolaris, worldwide (Schmidt and Shearer, 2003). Neolinocarpon globosicarpum and Maria and Sridhar (2003) studied fungal Oxydothis nypae were more frequently diversity on decomposing biomass of five recorded on fronds than other fungi, while mangrove plant species from the South Linocarpon bipolaris (13.5%), West coast of India. Astrosphaeriella striatispora (12.2%), Trichocladium nypae (8.1%) and Linocarpon appendiculatum (8.1%) were Typical marine fungi were not dominant in more frequently recorded on leaves. root endophytes of coastal sand dunes An overview on the diversity and halophytes (Beena et al., 2000), roots of ecology of fungi colonizing litter of mangrove plant species (Ananda and mangroves in Bay of Bengal region Sridhar, 2002). The assemblage and (mangroves of Godavary and Krishna diversity of filamentous fungi on leaf and deltas of Andhra Pradesh, Pichavaram of woody litter accumulated on the floor of Tamil Nadu, and Andaman and Nicobar two mangrove forests (Nethravathi and Islands). A total number of 131 species Udyavara) in the South West coast of India. belonging to 77 genera have so far been In their study, yielded 78 taxa belonging to reported from the three regions. 32 ascomycetes and 46 mitosporic fungi Verruculina enalia showed highest (Ananda and Sridhar, 2004). Sridhar (2005) percentage occurrence at all the sites and attempted to deal with occurrence, on different hosts. The fungi exhibited distribution and diversity of filamrntous vertical zonation in their occurrence with fungi in mangrove ecosystem. more number occurring in the intertidal zone. While some fungi occurred throughout the tidal range many showed Jones et al. (2006) reported marine fungal affinity to a particular level. Ascomycetes diversity of Thailand was investigated and with immersed or semi-immersed fruit 116 Ascomycota, 3 Basidiomycota, 28 bodies occurred in water inundated niches anamorphic fungi, 7 Stramenopiles (Vittal and Sarma, 2006). recorded, with 30 tentatively identified. These species have primarily been

Sridhar and Maria (2006) studied that the Fungal biodiversity in freshwater, brackish pattern of colonization and diversity of and marine habitats were estimated based filamentous fungi on naturally deposited on reports in the literature. The taxonomic and deliberately introduced Rhizophora groups treated were those with species mucronata Lamk. wood during monsoon commonly found on submerged substrates and summer in a mangrove of southwest in aquatic habitats: Ascomycetes (exclusive India and compares overall occurrence with of yeasts), Basidiomycetes, three species co-occurrence. Among 17 Chytridiomycetes, and the non-fungal core-group fungi ( 10 %), Aigialus Saprolegniales in the Class Oomycetes. mangrovei, Cirrenalia pygmea, Lignincola Based on presence/absence data for a large laevis, Lulworthia grandispora, number and variety of aquatic habitats, Passeriniella mangrovei, Trichocladium about 3,000 fungal species and 138 linderi, Tirispora sp., Zalerion maritimum saprolegnialean species have been reported and Z. varium were highly dominant ( 20 from aquatic habitats. The greatest number %). On wood showing co-occurrence of of taxa comprise the Ascomycetes, three fungi, A. mangrovei, Cirrenalia including mitosporic taxa, and tropicalis, L. grandispora and T. linderi Chytridiomycetes. Taxa of Basidiomycetes were highly dominant core-group fungi. are, for the most part, excluded from Even though A. mangrovei, C. pygmea, C. aquatic habitats. The greatest biodiversity tropicalis, Halosarpheia cincinnatula, L. for all groups occurs in temperate areas, grandispora, P. mangrovei, Verruculina followed by Asian tropical areas (Shearer et enalia and Z. maritimum are typical marine al., 2007). or mangrove fungi, they were core-group fungi on deliberately introduced wood in Fungi in Marine, Mangrove water and monsoon season indicates their high sediment colonization activity on wood even under low salinity. Several terrestrial mitosporic Single species isolated from mangrove fungi (Alternaria, Arthrobotrys, soils, for example, that by Stolk (1955) on Aspergillus, Penicillium, Phoma and an Emericellopsis and Westerdykellaornata Tetracrium) were found particularly in from East Africa and the paper by Swart monsoon season, but none of them (1970) on a Pencillium from Australia. The belonged to core-group. thraustochytrids comprise 7 genera and 31 species of marine fungoid protists has been The distribution of fungi in Muthupettai found in estuarine, Littoral, and oceanic mangroves along the East coast of Tamil waters and sediments around the world. Nadu, India was studied in terms of species (Gaertner 1967a, 1968a,b; Bahnweg and diversity, seasonal variation, and frequency Sparrow, 1974). Most fungi recorded from of occurrence in five sampling stations at marine sediments use collected from two different seasons. In this study, total of coastal regions and are typical soil fungi, 118 species of fungi isolated, of which which are terrestrial in origin (Borut and maximum 94 species from sediment Johnson, 1962; Vrijmoed and Hughes, samples followed by water with 83 species. 1990). Ulken (1970, 1972) studied Among the fungal isolates Aspergillus was occurrence and physiology of lower fungi the common genus followed by from marine sediment in Brazil. Penicillium, Curvualria and Alternaria (Sivakumar et al., 2006).

Studied on fungi of soil in mangrove nutrients from a discrete organic base in the vegetation and investigations on Indian sediment and produce a profusion of mangal soil were conducted by Pawar and hyphae which spread out and grow on the Thirumalachar (1966), Padhye et al. surface of sand grains producing ascocarps (1967), Rai et al. (1969), and Rai and strongly adding to them and this space Chowdhery (1975, 1976). In India single invading mode of life is one of the species isolated from mangrove soil by Rai characteristics of mycelial fungi and is well and Tewari (1963) on Preussia isolates, by known among fungi in terrestrial soils. Kirk Pawar et al. (1963) on a Monosporium, by (1983) reported that the flotation were Pawar et al. (1965) on a Cladosporium devised for the qualitative and quantitative and by Pawar et al. (1967) on Phoma study of marine higher fungi on sandy species. Pawar and Thirumalachar (1966) beach communities. Sample of the water, compared the growth of pure cultures of surface layer, sea foam, and sandy marine and terrestrial isolates of the same substrates and nearly fresh water body were species of soil fungi and concluded that examined marine fungi were readily most of the marine isolates grew batter on demonstrated in the water column and in sea water agar, then on a distilled water Neurton screen samples of the ocean medium whereas, the terrestrial isolates of surface, as well as sea foam and sand the same specie showed the reverse extracts. reaction. Hyde et al. (1987) reported the techniques Ulken (1970, 1972, 1975) isolated for obtaining sporulation of marine fungi, Phycomycetes from mangrove sediments in 65 Ascomycotina, 24 Deuteromycotina, ad Brazil and Hawaii, and Lee and Baker 3 Basidiomycotina known to sporulating (1972a,b, 1973) investigated soil micro culture. Jones (1993) revealed the marine fungi from a Hawaiian mangrove swamp. fungi appear to be distributed in relation to Nya (1976) revealed qualitative seawater temperature; arctic, temperate composition of marine fungi from 3 tropical; although others grew equally well biotopes, water at different depths, bottom over a range of temperature. sediments, and Macrophytes and the classes Christopherson et al. (1999) reported that a of Phycomycetes, Ascomycetes and total of 227 marine isolates of ubiquitous Deuteromycetes were recorded and the fungi were cultivated on different media total of 116 species of 41 genera with the and fungi were isolated include 18 different greatest variety of species observed in the fungal species from 8 Ascomycetes genera bottom sediments. The Macrophyte from animals, plants and sediments of mycoflora comprised 67 species of 28 Venezuelan waters (0 10m) including while in water 64 species of 27 general mangroves and lagoon areas. were found. Kohlmeyer and Kohlmeyer (1979) maintained that most of these fungi Occurrence of fungi on different are isolated from dormant propagules and substrata / tissue specificity (wood/twig, not from actively growing fungi and that pneumatopores, seedlings, leaves and isolation methods excluded almost all true roots marine inhabitat s. Soft rot in terrestrial known since the The arenicolous or sand inhabitating middle of nineteenth century and this type (Kohlmeyer and Kohlmeyer, 1979) draw of wood decay was later described and

illustrated for fungi of marine habitats seedling of Rhizophora spp. Fell and (Kohlmeyer, 1958). Ritchie (1959) has Master (1973) reported that the senescence shown that common terrestrial fungi exist of leaf materials, they support a very there on submerged wood and other similar different fungal community, which are substrates. Kohlmeyer (1969a) observed responsible for their degradation. that among large collections, several fungi were encounted only in roots and stems of living Avicennia or Rhizophora and appear Rai et al. (1969) tested wood decaying to be host specific. Terrestrial fungi potential of twenty-two species of develop on roots and branches above the mangrove swamp fungi. The classified high tide line and on over lapping these fungi into groups based of on weight between marine and terrestrial species may loss; above 31% strong, between 28 and occur at the water air interface 37% - moderate and below 28% weak. (Kohlmeyer, 1969b). Lee and Baker (1973) Ravikumar and Purushothaman (1998) demonstrated some of the isolated fungi studied on lignicolous marine fungi in the derived from dormant propagules of Vellar estuary, Tamil Nadu and recorded 1 terrestrial species. species of Ascomycetces fungus Corollospora intermedia which are a new, Johnson (1967) noted that the lignicolous recorded from India. Few reports of fungi collected in the Neuse Newport endophytic fungi on substrates collected estuary were predominantly of terrestrial from the marine environment and these origin (73%) and majority of these were include leaves and droppers of mangrove from water of less than 18% salinity (81%) plants (Suryanarayanan et al., 1998; Abdel while only 12% were recovered from whole Wahab et al., 1999; Kumaresan and estuary i.e. 0- 34 %. Kohlmeyer and Suryanarayanan, 1999). Mitosporic fungi Kohlmeyer (1971) listed 51 marine and frequent inhabitants of leaves and it is Deuteromycetes and 36 of these are common to find non marine species such lignicolous and the majority belongs to as Cladosporium cladosporiodes and terrestrial genera (Alternaria, Pestalotiopsis species in the early stages of Camerosporium, Dedryphiella, Diplodia, incubation and Pencillium species and Humicola, Monodictys), while other Trichoderma species. exclusively marine (Cirrenalia, Orbimyces and Zalerion). The only extensive study of the terrestrial colonization of mangrove seedlings is that Ritchie (1968) submerged wood of 8 by Newell (1973, 1976) investigated the tropical trees in the Panama canal zone and succession of fungi on submerged seedlings examined wood sections and found green of Rhizophora mangle. Accordingly the heart (Ocotea rodiaei) and red mangrove marine fungi encounted in the mangrove (Rhizophora mangle) preactically fungus habitat live on roots, stems, and twigs free after about 6 weeks of exposure and submerged in water and their terrestrial one marine fungi Lulworthia species in counterparts inhabit leaves, stems, branches addition to some 20 Deuteromycetes of and upper parts of the roots above the water terrestrial origin was isolated Kohlmeyer surface (Kohlmeyer 1974; Kohlmeyer and (1969b) reported that the ascocarps of Kohlmeyer, 1979). Kohlmeyer and Keissleriella blepharospora develop Kohlmeyer (1977) examined host plants of between cork cells of roots or submerged Avicennia germinans (L.,), Conocarus

erecta L., Salicornia virginica L., Tamarix Kohlmeyar and Vittal (1986) studied gallica L., Thalassia testudinum Koenig marine fungi of the mangal in Belize and and the collections include 15 India, encounted indepandtly a common Ascomycetes, 1 Basidiomycetes and 6 Ascomycete (Lophiostoma sp.) growing at Deuteromycetes which are new records for the upper intertidal level on bark ad wood the Bermuda Islands. of mangrove trees. Mouzouras et al. (1988) reported the ability of microorganisms to Byrne and Jones (1975) reported that the 28 decay wood submerged in the sea and 42 microfungi which grew on wood blocks of marine fungi, belonging to the beach (Fugues sylvatica) and Scots pine Asomycotina and Deuteromycotina have (Pinus sylvestris) submerged in seawater at been shown to cause soft - rot decay of Port Erin, Langstone harbour and Newton wood while 3 Basidiomycotina caused Farers were recorded. Schneider (1971) white rot - decay. Pena et al. (1996) observed 23 species of fungi with the reported that the first contribution to the greatest number occurring on beach wood, knowledge of lignicolous marine fungi which also offered the best conditions for from Mardal plata, Argentina .10 species rapid development. Quantitative data on the were collected from submerged wood occurrence of tropical marine fungi have panels, intertidal wood and driftwood. been published by Kohlmeyer (1984) and Zainal and Jones (1984, 1986), however all Studies on marine and mangrove fungi of of their reports were on driftwood in the sea Indian ocean is quite recent and has been along with driftwood on the mangrove less well explored compared to the Atlantic floor. and Pacific Oceans (Koch, 1986; Borse, 1988, Zainal and Jones; 1986; Hyde, Marine fungi like Leptosphaeria species, 1988a; Steinke and Jones, 1993). Although Mycosphaerella species, and Cirrenalia Indian sandy beaches, mainland mangroves macrocephala were for the first time and some islands have been studied for recorded only by direct microscopic mangrove and marine fungi, there are few observation of the pneumatopores of quantitative studies (Borse, 1988; Avicenia officinalis from Indian mangrove Ravikumar and Vittal, 1996; Prasannarai habitat by Garg (1982). Hyde (1989c) and Sridhar, 1997, 2000-2001, 2001; Sarma reported that the decayed archids and Vittal, 2000, 2001; Sarma et al., 2001; Acrostichum speciosum (Mangrove fern) Maria and Sridhar, 2002). Sarma and Hyde were collected from Kampong Kapok (2001) have reviewed factors affecting the mangrove, Brunei and examined for higher frequency of occurrence of fungi in marine fungi (A new bitunicate mangroves. A few studies are available on ascomycete, Massarina acrostichi). the impact of incubation of lignocellulosic materials collected from different habitat on the occurrence of fungi (Hyde, 1992b; Hyde et al. (1990) investigated the Prasannrai and Sridhar, 1997). distribution of fungi on Sonnertia griffithi and showed that some fungi were common Marine substrata support different fungal on pneumatophores (Aigialus grandis and assemblages, for example the mangrove Massarina velatospora) while others were palm Nypa fruticans and woody tissue of common on twigs (Saccarcloella mangrove trees such as Rhizophora mangrovei and Savoryella longispora). apiculata and Avicennia marina. Typical

fungi on N. fruticans included Lulworthia species were more common as Astosphaeriella striatispora, Linocarpon early colonizer. A relatively high appiculata, L. nypae, Oxydothis nypae and percentage of the fungi colonizing the test Trichocladium nypae, taxa never recorded blocks were mitosporic fungi. Seasonal from mangrove wood (Hyde and Nakagiri, occurrence and colonization of intertidel 1989; Hyde, 1992a; Hyde and Alias, 2000; wood and introduced teak wood panels by Pilantanapak et al., 2005). marine fungi was investigated in mangrove Harbour (West coast of India) of the 33 Hyde (1991) reported Phomopsis taxa encounted 29 taxa were found on mangrovei, coelomycetous fungus on intertidal wood and 15 species on teak. proproots of Rhizophora apiculata from The most frequently collected species on Ranong mangrove, Thailand. Kohlmeyer intertidal wood were Trichocladium and Kohlmeyer (1993) revealed the species, Zalenion varium, and T. comparison of the marine mycota of alopallonellum. Sarma and Vittal (2001) recently introduced Rhizophora species recorded the number of fungi on prop roots (Hawaii and Moorea) with that of long of R. apiculata (61) was much greater when established Rhizophora stands from the compared to wood (24) and seedlings, 21 Caribbean (Belize) and 43 species are from the developing substrate collected known from Rhizophora in Belize and only from the deltaic mangrove of Godavari and 7 and 21 species were collected from Krishna rivers in the East coast of India. Moorea and Oahu respectively. Borse et al. (2000) reported that the higher Ravikumar and Vittal (1996) reported on marine fungi in foam and intertidal wood the fungi colonizing different substrata of and dead submerged wood of Avicennia Rhizophora apiculata and R. mucronata marina from Daman coast. In this study 13 from Pichavaram mangroves of Tamil species of higher marine fungi (10 Nadu, East coast of India and concluded Ascomycetes,3 Deuteromycetes) were that different substrata of the same host recorded. Borse (2000) studied that the 83 plant are colonized by different frequently species (62 Ascomycetes, 3 occurring fungi. According to Hyde et al. Basidiomycetes, 18 Deuteromycetes) of (1990b) bark was an important factors in higher marine fungi from Maharashtra determing the mycota present on Coast (The Arabian sea) including 19 Rhizophora apiculata particularly when species as new records for the fungi of small diameter roots were examined. Maharashtra from the substrates include Young roots surrounded by bark were driftwood, intertidal wood harbour timber invariably colonized by Leptosphaeria sp., and dead submerged parts of the Lulworthia grandispora, Massarina mangroves. ramunculicola, phomopsis sp. and Rhizophila marina. Sarma and Vittal (2001) examined the decaying mangrove materials belonging to Poonyth et al. (1999) revealed split wood a host plants species collected from blocks of Bruguiera gymnorrhiza and Godavari and Krishna deltas, East coast of Rhizophora mucronata were submerged in India include, 65 Ascomycetes (74%), 1 the intertidal zone of 5 mangrove sites in Basidiomycetes and 22 Mitosporic fungi Maritrius of the blocks over regular (25%) (Including 6 Coelomycetes and 16 intervals and Cirrenalia pygmea and Hyphomycetes).

leaves of a spermatophyte (Thalassia) as Fungi on Seaweeds and Seagrasses well as in air vesicles of Sargassum sp.

Some of the leaf inhabiting saprobes The rhizoid of Phycomycetous fungi and occur on rhizomes of the same host as well; the ectoplasm that elements of for instance, Lulworthia species, on Thraustochytrids penetrate the host cells rhizomes of Zostera marina (Kohlmeyer, and draw nutrients and they may also be 1963) and Thalassia testudrinum, endoparasites (Chandralata, 1986, 1987). (Kohlmeyer and Kohlmeyer 1971). Tubaki Recent work on Juncus roemerianus and Asano (1965) recorded 16 imperfect (Kohlmeyer and Kohlmeyer, 1995, 1996; fungi on seaweeds most of them as Kohlmeyer et al., 1995, 1996, 1997) and on saprophytes like Dendryphiella qrenaria, Phragmites australis (Poon and Hyde, D. salina, Alternaria maritima and 1998) have resulted in the discovery of a Monodictys austrina. Leptosphaeria sp., number of species new to science. Pleospora sp., and Sphaerulina pedicellata are all very common on Spartina culms by Fungi on Sea foams Johnson and Howard (1968). The sporadic occurrence of algicolous fungi may be Kohlmeyer (1966) found that the fungal explained by antibiotic substances spores in foam, which is a good indication produced by healthy algae (Sieburth, 1968). of the mycota present in the sand of a particular beech. Kohlmeyer (1966) Meyers (1969) has described a second reported that the propagules of marine species of Lindra from Thalassia species, fungi contained in foam are deposited by namely Lindra marinera and suggests it the waves on washed up substrates, such may be active in the degradation of as seagrasses, algae or animal remains. Thalassia leaves. Meyers et al. (1970) Appendaged Ascospores and Basidiospores suggest that yeast my be active in the , and tetra radiate conidia, are regularly degradation of Spartina as well as a found in sea foam along sandy beaches, number of fungi imperfect like Fusarium, mostly together with algae and protozoa Phoma, and Nigrospora sp. Kohlmeyer and (Schlichting, 1971). Kohlmeyer (1966) Kohlmeyer (1971) have reported examined 5 Ascomycetes, one Varicosporina ramulosa, Halocyphina Basidiomycates, and 9 fungi Imperfecti villosa, from Spartina alterniflora, Loisel, from foam samples collected along shores. from decaying leaves of Claviceps purpurea. Jones (1976) and Kohlmeyer and Conidial fungi on sea foams and leaves Kohlmeyer (1979) have revised the have been explored from various parts of occurrence of marine fungi on algae and the world by Bandoni (1972), Barlocher seaweed though these substrates have not and Kendrick (1977), Crane (1968), been studied as intensively as fungi on Conway (1969), Descals et al. (1977), wood tissue. Stanley (1991). Recent Dudka (1974), Durrieu (1970), Dyko reviews of litereature on algicolous fungi (1978), Gonczol (1975), Greathead (1961), have been published by Andrews (1976), Ingold (1975), Iqbal and Webster Jones (1976) and Kohlmeyer (1974). (1973,1977), Iqbal (1974), Marvanova Kohlmeyer and Kohlmeyer (1979) reported (1972), Marvanova et al. (1967), Nilsson that the Lindra thalassiae is the most (1958) and Nawawi (1973, 1975). Presence indiscriminate species of all, as it occurs in of fungal spores in foam in aquatic

environments has long been recognized by 1979; Boyd and Kohlmeyer, 1982). Kohlmeyer and Kohlmeyer (1979), Kirk Microscopic counts of fungi, algae, (1983) and also systamatic investigation protozoa and in liquefied sea foam fungi in sea foam was carried out on a demonstrated the potential of the simple sandy beach in Viginia. and direct approach information of the ecological roles, seasonal and geographic Recently, Bandoni (1972) reported the distribution of marine microorganisms occurrence of some of these conidial fungi (Kohlmeyer and Kohlmeyer, 1979). New in terrestrial habitats and 40 conidial fungi geographical limits for several species, the were identified and assigned to 20 genera. seasonal distribution of tropical V. Kirk et al. (1973) reported that the ramulosa and the vertical zonation of collection ,identification of ecological Corollospora species, deposited by sea groups of lignicolous, arenicolous, foam were less active physiologically graminicolous and endocommansalic fungi than geo fungi and calcicolous marine within the sea foam ,wood and marsh grass species in the upper 30cm of the strand plants. The role of conidial fungi in line. (Kirk, 1983). A new marine processing of an aquatic litter, energy flow, Ascomycte, Lindra obtusa and its productivity and experimental aspects of anamorph, Anguillospora marina isolated these fungi have been worked out by from sea foam samples on some shores of Barlocher and Kendrick (1974, 1976), Japan and appear to be adapted to marine Suberkropp and Klug (1976) and habitats was described by Nakagiri and Suberkropp and Thomas (1984). Ingold Tubaki (1983). (1975) reported that conidia of most of them accumulate in foam, which acts as a Murthy and Manoharachary (1981), spore trap, and the examination of a foam Manoharachary and Madhusudhan Rao samples can very quickly give a picture of (1983), Madhusudhan Rao and the species present in a particular stream Manoharachary (1984), Sridhar and above the point of collection . The Kaveriappa (1982) and Subramanian and ecological studies of concerning conidial Bhat (1981) have studied the conidial fungi fungi have emphasized their role in associated with foam and submerged processing litter, energy flow, and leaves. Sridhar and Kaveriappa (1982) have productivity in aquatic ecosystems studied the conidial fungi associated with (Suberkropp and Klug, 1976). foam and submerged leaves. Mahusuhan Conidiogenesis has been worked Rao and Manocharachary (1984) have thoroughly in some conidial fungi reported the association of conidial fungi colonizing submerged leaves and foam by on diversified submerged leaves. Descals et al. (1977) and Ingold (1975). Patil (2003) reported that the submerged Seafoam generally contains the highly leaves of Memmeaylon umbellaun and distinctive spores of Corollospora sp., Mangifera indica were colonized by Carbosphaerella sp., Varicosporina species of Alastospora, Beltrana, ramulosa in tropical and Asteromyces Flagellospora, Ingolidiella, Lemonniera, cruciatus in temperate areas, Alternaria and Lunulospora, Monosporella and other terrestrial and marine fungi Triselophorus along with these fungal depending upon climate and local forms of species of Actinospora, conditions, (Kohlmeyer and Kohlmeyer, Angiullospora, Cameroporium,

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