Fungal Biodiversity in Salt Marsh Ecosystems

Abstract

This review brings together the research efforts on salt marsh fungi, including their geographical distribution and host association. A total of 486 taxa associated with different hosts in salt marsh ecosystems are listed in this review. The taxa belong to three phyla wherein Ascomycota dominates the taxa from salt marsh ecosystems accounting for 95.27% (463 taxa). The Basidiomycota and Mucoromycota constitute 19 taxa and four taxa, respectively. Dothideomycetes has the highest number of taxa, which comprises 47.12% (229 taxa), followed by Sordariomycetes with 167 taxa (34.36%). Pleosporales is the largest order with 178 taxa recorded. Twenty-seven genera under 11 families of halophytes were reviewed for its fungal associates. Juncus roemerianus has been extensively studied for its associates with 162 documented taxa followed by Phragmites australis (137 taxa) and Spartina alterniflora (79 taxa). The highest number of salt marsh fungi have been recorded from Atlantic Ocean countries wherein the USA had the highest number of species recorded (232 taxa) followed by the UK (101 taxa), the Netherlands (74 taxa), and Argentina (51 taxa). China had the highest number of salt marsh fungi in the Pacific Ocean with 165 taxa reported, while in the Indian Ocean, India reported the highest taxa (16 taxa). Many salt marsh areas remain unexplored, especially those habitats in the Indian and Pacific Oceans areas that are hotspots of biodiversity and novel fungal taxa based on the exploration of various habitats.

1. Introduction

Salt marsh ecosystems are known for their high productivity, exceeding primary production estimates of species rich ecosystems (e.g., tropical rainforests, coral reefs) [1]. The flora in salt marsh ecosystems is mainly composed of grasses, herbs, and shrubs and these are terrestrial organisms variously adapted to, or tolerant of, a semi-marine environment. Halophytes are a diverse group of plants that have a worldwide distribution, and grow in different climatic regions, wherein soils have high salinity levels [2]. Halophytes are common in temperate and Mediterranean climates, and fewer both in the tropics and at high latitudes [3,4,5,6]. The vegetation in these ecosystems shows the vertical zonation of different communities as tidal submergence decreases with increasing elevation, and species tolerance to changing gradient conditions. Salt marsh vegetation generally increases the attenuation of both tidal currents and waves as they pass over the vegetated area and immobilize elements with their sediments. Furthermore, halophytes serve as a natural buffer, protecting other shoreline ecosystems from human impacts and disturbances. The area provides a habitat and nursery for marine organisms [7]. Worldwide, salt marshes cover an area of 5,495,089 hectare in 43 countries [8].

There are over 500 species of salt marsh plants worldwide [9]. The families Amaranthaceae (subfamilies Chenopodiaceae, Salicornieae), Poaceae, Juncaceae, and Cyperaceae are the major vegetation in salt marsh ecosystems, while the minor components are Plumbaginaceae and Frankeniaceae [3], and are represented in Figure 1 and Figure 2. Salinity, latitude, region of the world, the frequency and duration of tidal flooding, substrate, oxygen and nutrient availability, surface elevation, competition among species, disturbance by wrack deposition are interacting factors that influence the species of halophytes in the salt marshes [10,11]. For example, Spartina alterniflora is a dominant grass from mid-tide to high-tide levels in temperate Eastern North America, while Puccinellia dominates in boreal and arctic marshes [10,11].

Figure 1.

Salt marsh ecosystems in UK (a–d) and Thailand (e–f). (b–d) Tidal grasses, Spartina townsendii (Poaceae) and Phragmites (Poaceae), dominate the salt marsh in UK (50°49′55.4″ N 0°58′25.1″ W; 51°43′03.1″ N 5°10′24.8″ W); (e) Spartina (Poaceae) (12°22′4.0″ N 99°59′6.7″ E) (f) and Suaeda (Amaranthaceae) (12°10′19.6″ N 99°58′20.3″ E) in tidal marsh areas in southern Thailand.

Figure 2.

Halophytes in salt marsh ecosystems: (a) flowering inflorescence of Spartina, (b) Phragmites, (c) Salicornia, (d) Typha, (e,f) Atriplex, and (g,h) Suaeda.

Major studies on halophytes focus on ecology and conservation [12,13,14]. One of these is the decomposition of vascular plant material wherein the detritus breakdown was reviewed in Pomeroy and Wiegert [15], Howarth and Hobbie [16], and Long and Mason [17]. The active decomposition processes in salt marsh ecosystems reflects to the relatively high rates of primary production. Three phases of plant decomposition were noted by Valiela et al. [18]. The early phase involves the leaching of soluble compounds, resulting in a fast rate of weight loss lasting for less than a month. Organic matter breakdown by microorganisms and continuous leaching of decayed products occurs in the second phase that lasts for a year. The last phase lasts for another year when there is a slow decay of refractory materials such as humates and fulvates [19].

The continuous breakdown of detritus into smaller fragments increases the surface-to-volume ratio and this is exposed to further microbial degradation. Bacteria and fungi are key decomposers in the salt marsh ecosystem that are essential for the transformation and recycling of nutrients through the environment. The colonization of fungi on standing dead halophytes commences during the early stages of decomposition before leaf fall to the salt marsh sediment surface [20,21]. The decomposition of the senescent tissues of halophytes by salt marsh fungi is brought about by the direct penetration of the host cell wall and the production of enzymes active in degrading lignocellulosic compounds, such as lignin, cellulose, and hemicellulose [22,23,24,25,26]. Bacterial communities are the major decomposers in the latter stage of decomposition [27,28]. Studies in salt marsh ecosystems not only consider microbial activity and the recycling of nutrients, but also bacterial [29,30] and fungal diversity [20,31,32].

The present review compiles the published data of fungi from halophytes, including their geographical distribution and host association. When compared to other fungal groups, salt marsh fungi are underexplored, and this review brings together the research efforts on these undiscovered habitats and plants. The pertinent literature from bibliographic databases (e.g., Scopus, Web of Science, Google Scholar) and published resources on salt marsh fungi documenting halophytes were compiled. Published works, wherein the documented fungal taxa were observed directly from halophytic substrates, are included (Table 1). The different host parts, living and dead, that are either partly or wholly submerged are documented, as well as drift plant portions washed up in salt marsh areas. Salt marsh fungi isolated using cultivation-dependent techniques were not included since it is not known if these fungi were actively growing and reproducing on the halophytes. The taxa were listed based on the recent outline of fungi and fungus-like taxa by Wijayawardene et al. [33]. Since previous works only listed the taxa and the hosts [34,35,36], here we include the plant parts where the fungus was observed, the location (country: state/province) where the host was collected, the life mode of the fungus, and the pertinent literature citations are included (Table 1). The accepted name of the host was based on the webpage of the World Flora Online consortium (http://www.worldfloraonline.org/; accessed on 10 May 2021), GrassBase (https://www.kew.org/data/grasses-db/sppindex.htm; accessed on 10 May 2021) and CRC World Dictionary of Grasses by Quattrocchi [37]. The graphs presented in the next sections summarizes the information from Table 1 and was developed using data visualization tools (Excel Office 365, Tableau Desktop Professional Edition 19.2.2).

Table 1.

Geographical distribution of salt marsh fungi recorded from various halophytes.

Taxon	Host Part	Life Mode	Hosts	Distribution	References
ASCOMYCOTA
DOTHIDEOMYCETES
Acrospermales
Acrospermaceae
Acrospermum graminum Lib.	–	–	Elymus pungens	UK	[38]
Asterinales
Morenoinaceae
Morenoina phragmitis J.P. Ellis	Living/decomposing leaf sheaths and stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40]
Botryosphaeriales
Botryosphaeriaceae
Botryosphaeria festucae (Lib.) Arx and E. Müll.	Living/decomposing leaf sheaths and stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40]
Macrophomina sp.	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Tiarosporella halmyra Kohlm. and Volkm.-Kohlm.	Senescent culms	Saprobic	Juncus roemerianus	USA: North Carolina	[42]
Phyllostictaceae
Guignardia spp.	Senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Phyllosticta sp.	–	Pathogenic	Spartina cynosuroides	USA: Maryland	[44]
Phyllosticta spartinae Brunaud	–	–	Spartina maritima	France	[45]
Phyllosticta suaedae Lobik	Leaves	–	Suaeda maritima	Russia	[46]
Capnodiales
Cladosporiaceae
Cladosporium algarum Cooke and Massee	–	–	Spergularia marina	–	[35]
	–	–	Suaeda maritima	–	[35]
Cladosporium allicinum (Fr.) Bensch, U. Braun and Crous	–	–	Elymus pungens	UK	[38]
Cladosporium cladosporioides (Fresen.) G.A. de Vries	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Leaves and roots	Saprobic	Spartina sp.	Canada: Bay of Fundy	[48]
Cladosporium herbarum (Pers.) Link	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
	Stem	Saprobic	Spartina townsendii	UK: England	[49]
	Leaves, stems, and roots	Saprobic	Spartina sp.	Canada: Bay of Fundy	[48]
Cladosporium macrocarpum Preuss	Leaves	Saprobic	Spartina sp.	Canada: Bay of Fundy	[48]
Cladosporium sphaerospermum Penz.	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Living/decomposing leaf sheaths and blades	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,41,50]
	–	Saprobic	Spartina patens	USA: Rhode Island	[36]
	–	Saprobic	Spartina sp.	Canada	[36]
Capnodiales genera incertae sedis
Mucomycosphaerella eurypotami (Kohlm., Volkm.-Kohlm. and O.E. Erikss.) Quaedvl. and Crous	Senescent leaves	Saprobic	Juncus roemerianus	USA: North Carolina	[51]
Mycosphaerellaceae
Fulvia fulva (Cooke) Cif.	Leaves and stems	Saprobic	Spartina sp.	Canada: Bay of Fundy	[48]
Micronectriella agropyri Apinis and Chesters	–	–	Elymus pungens	UK	[38]
	–	–	Puccinellia maritima	UK	[38]
	–	–	Spartina townsedii	UK	[38]
Mycosphaerella lineolata (Roberge ex Desm.) J. Schröt.	Living/decomposing leaf sheaths and stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40]
	–	–	Elymus pungens	UK	[38]
Mycosphaerella salicorniae (Auersw.) Lindau	–	–	Arthrocnemum subterminale	–	[35]
	–	–	Limonium sp.	–	[35]
	–	–	Sarcocornia perennis	–	[35]
	–	–	Salicornia fruticosa	–	[35]
	–	–	Salicornia procumbens	–	[35]
	–	–	Salicornia europaea	–	[35]
	–	–	Salicornia perennis	–	[35]
	–	–	Sarcocornia fruticosa	–	[35]
	Drying stalks and inflorescence	Saprobic	Salicornia sp.	India	[52]
	Dried inflorescences	Saprobic	Salicornia virginica	Bermuda	[35,53]
	–	Saprobic	Spartina marítima	Portugal: Alentejo, Lisbon	[54]
	–	–	Suaeda vermiculata	–	[35]
	Drying stalks and inflorescence	Saprobic	Suaeda sp.	India	[52]
Mycosphaerella spp.	–	–	Elymus pungens	UK	[38]
	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida, Mississippi	[43,55]
	Decaying leaves, leaf blades	Saprobic	Spartina alterniflora	Argentina: Buenos Aires; USA: Alabama, California, Georgia, Mississippi	[25,35,36,55,56,57,58]
	–	–	Spartina cf. densiflora	USA: California	[25,35]
	–	–	Spartina cf. pectinata	–	[35]
	–	–	Spartina sp.	Argentina: Buenos Aires; Canada	[35,36]
	Decaying leaf blades	Saprobic	Spartina foliosa	USA: California	[25]
	Leaf sheaths and blades, stem	Saprobic	Spartina marítima	Portugal: Alentejo, Lisbon, Centro	[54,59]
Mycosphaerella staticicola (Pat.) Dias	–	–	Armeria pungens	–	[35]
Mycosphaerella suaedae-australis Hansf.	–	–	Suaeda australis	–	[35]
Rivilata ius Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Tips of senescent, very old, and brittle leaves	Saprobic	Juncus roemerianus	USA: North Carolina	[60]
Septoria spp.	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
	Upper leaves, inflorescence, seeds	Saprobic	Spartina alterniflora	USA: Rhode Island	[61]
Septoria suaedae-australis Hansf.	Dead stems	Saprobic	Suaeda australis	South Australia	[62]
Sphaerulina albispiculata Tubaki	Sheath	Saprobic	Spartina marítima	Portugal: Alentejo, Lisbon	[54]
	Stem	Saprobic	Spartina marítima	Portugal: Alentejo	[63]
Sphaerulina orae-maris Linder	–	–	Ammophila arenaria	–	[35]
	Rhizome and root	Saprobic	Spartina densiflora	Argentina: Buenos Aires	[64]
	Leaf sheaths and blades, stem	Saprobic	Spartina marítima	Portugal: Alentejo, Lisbon, Algarve, Centro	[31,54,59,63]
Sphaerulina pedicellata T.W. Johnson	–	Saprobic	Spartina townsendii	–	[65]
	Attached culms, stems	Saprobic, parasitic	Spartina alterniflora	USA: Rhode Island	[20,61]
Sphaerulina sp.	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Dothideales
Saccotheciaceae
Aureobasidium sp.	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Pseudoseptoria donacis (Pass.) B. Sutton	Living/decomposing leaf blades and sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,50]
Selenophoma sp.	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Dothideaceae
Scirrhia annulata Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Senescent culms and leaves	Saprobic	Juncus roemerianus	USA: North Carolina	[66]
Dothideomycetes families incertae sedis
Eriomycetaceae
Heleiosa barbatula Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Senescent leaves	Saprobic	Juncus roemerianus	USA: North Carolina	[66]
Pseudorobillardaceae
Pseudorobillarda phragmitis (Cunnell) M. Morelet	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41,67]
Pseudorobillarda sp.	Dead stems	Saprobic	Spartina alterniflora	Canada	[36]
Dothideomycetes genera incertae sedis
Bactrodesmium atrum M.B. Ellis Lautitia danica (Berl.) S. Schatz	Living/decomposing stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[40]
	–	–	Elymus pungens	UK	[38]
	–	–	Puccinellia maritima	UK	[38]
Monodictys austrina Tubaki	Senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Monodictys castaneae (Wallr.) S. Hughes	Leaves	Saprobic	Spartina sp.	Canada: Bay of Fundy	[48]
Neottiosporina australiensis B. Sutton and Alcorn	Living/decomposing leaf blades and sheaths, stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40,50]
Neottiosporina sp.	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Otthia sp.	Senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Trichometasphaeria setulosa. (Sacc. and Roum.) Apinis and Chesters ined.	–	–	Elymus pungens	UK	[38]
Trichometasphaeria sp.	–	–	Elymus pungens	UK	[38]
Microthyriales
Microthyriaceae
Microthyrium microscopicum Desm.	–	–	Spartina patens	–	[68]
Microthyrium gramineum Sacc., E. Bommer and M. Rousseau	–	–	Elymus pungens	UK	[38]
Muyocopronales
Muyocopronaceae
Ellisiodothis inquinans (Ellis and Everh.) Theiss.	–	Saprobic	Spartina alterniflora	Argentina: Buenos Aires	[36]
Mytilinidiales
Mytilinidiaceae
Septonema secedens Corda	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Phaeotrichales
Phaeotrichaceae
Trichodelitschia bisporula (P. Crouan and H. Crouan) E. Müll. and Arx	–	–	Elymus pungens	UK	[38]
			Spartina townsendii	UK	[38]
Pleosporales
Amniculicolaceae
Neomassariosphaeria typhicola (P. Karst.) Y. Zhang ter, J. Fourn. and K.D. Hyde	–	–	Juncus roemerianus	–	[35]
	Decaying herbaceous stems	Saprobic	Spartina densiflora	Argentina: Buenos Aires	[64]
	–	Saprobic	Spartina spp.	Argentina: Buenos Aires	[32,35,36]
	–	Saprobic	Unidentified saltmarsh plants	USA: Mississippi	[58]
Camarosporiaceae
Camarosporium feurichii Henn.	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
Camarosporium palliatum Kohlm. and E. Kohlm.	–	–	Sarcocornia perennis	–	[35]
	–	–	Salicornia sp.	–	[35]
	–	–	Salicornia virginica	–	[35]
	–	Saprobic or perthophytic	Salt marsh plants	India: Maharashtra	[52]
	–	–	Suaeda vermiculata		[35]
Camarosporium roumeguerei Sacc.	–	–	Atripex halimus		[35]
	–	–	Atripex sp.		[35]
	–	–	Distichlis spicata		[35]
	Twigs	–	Salicornia europaea	France	[35,69]
	–	–	Sarcocornia fruticosa		[35]
	–	–	Salicornia sp.		[35]
	–	Saprobic or perthophytic	Salt marsh plants	India: Gujarat, Maharashtra, Tamil Nadu, Andhara Pradesh, West Bengal	[52]
	Leaf sheaths and blades, stem	Saprobic	Spartina maritima	Portugal: Algarve, Centro	[59]
	–	–	Suaeda maritima	–	[35]
Camarosporium salicorniae Hansf.	Twigs	–	Sarcocornia quinqueflora	South Australia	[62]
Camarosporium spp.	Living/decomposing leaf sheaths and stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40]
Camarosporium suaedae-fruticosae S. Ahmad	Dead branches	Saprobic	Suaeda vermiculata	Pakistan	[70]
Coniothyriaceae
Coniothyrium obiones Jaap	–	–	Atriplex portulacoides	–	[35]
	–	Saprobic	Salt marsh plants	India: Orissa	[52]
	Leaf sheaths and blades, stem	Saprobic	Spartina maritima	Portugal: Algarve	[59]
Coniothyrium spp.	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Cyclothyriellaceae
Massariosphaeria erucacea Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Senescent culms and leaves	Saprobic	Juncus roemerianus	USA: North Carolina	[66]
Massariosphaeria scirpina (G. Winter) Leuchtm.	–	Saprobic	Spartina sp.	USA: Florida, North Carolina	[71]
Massariosphaeria sp.	Living/decomposing stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[40]
Dictyosporiaceae
Dictyosporium oblongum (Fuckel) S. Hughes	Living/decomposing leaf blades and sheaths, stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40,50]
Dictyosporium pelagicum (Linder) G.C. Hughes ex E.B.G. Jones	Decomposing culms	Saprobic	Spartina alterniflora	USA: Rhode Island	[35,61]
	–	–	Spartina spp.	–	[32]
	Leaf sheaths and blades, stem	Saprobic	Spartina marítima	Portugal: Alentejo, Lisbon, Algarve, Centro	[54,59,63]
Jalapriya toruloides (Corda) M.J. D’souza, Hong Y. Su, Z.L. Luo and K.D. Hyde	Stems	Saprobic	Spartina sp.	UK	[72]
Didymellaceae
Ascochyta cf. arundinariae Tassi	Living/decomposing leaf blades and sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,50]
Ascochyta leptospora (Trail) Hara	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
Ascochyta salicorniae-patulae (Trotter) Melnik	–	Saprobic, parasitic	Salicornia spp.	Canada, Denmark, Germany, India, UK, USA	[52]
Ascochyta spp.	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
	Sheath	Saprobic	Spartina marítima	Portugal: Alentejo	[54]
Chaetasbolisia sp.	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Didymella glacialis Rehm	Living/decomposing leaf blades and sheaths, stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40,50]
Didymella glomerata (Corda) Qian Chen and L. Cai	Rhizome and basal area	Saprobic	Spartina densiflora	Argentina: Buenos Aires	[64]
Didymella spp.	Living/decomposing leaf blades and sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,50]
	–	Pathogenic	Spartina cynosuroides	USA: Louisiana	[44]
Epicoccum nigrum Link	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Inflorescence, upper leaves, seeds	Saprobic, parasitic	Spartina alterniflora	USA: Rhode Island, Connecticut, Virginia, Florida, North Carolina	[36,61,73,74]
Epicoccum sp.	–	–	Spartina alterniflora	–	[35]
Microsphaeropsis spp.	Living/decomposing leaf blades and sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,41,50]
Phoma herbarum Westend.	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
Phoma leveillei Boerema and G.J. Bollen	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
Phoma suaedae Jaap	Twigs, leaves, stems	Saprobic	Suaeda maritima, Suaeda sp.	Germany; India	[75]
	–	–	Suaeda maritima	–	[35]
Phoma spp.	–	–	Crithmum maritimum	–	[35]
	–	–	Atriplex portulacoides	–	[35]
	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Living/decomposing leaf blades and sheaths, stems	Saprobic	Phragmites australis	China: Hong Kong; Netherlands: Zeeland	[39,40,41,50]
	–	–	Salicornia europaea	–	[35]
	–	–	Spartina alterniflora	USA: North Carolina, Rhode Island	[20,35,36,61,73,74]
	–	Saprobic	Spartina patens	USA: Rhode Island	[36]
	–	Saprobic	Spartina sp.	Argentina: Buenos Aires; Canada; USA: Maine, South Carolina	[36,71]
	–	–	Spartina townsendii	UK: England	[35,49,65]
	Leaf sheaths and blades, stem	Saprobic	Spartina marítima	Portugal: Alentejo, Lisbon, Algarve, Centro	[54,59,63]
Paraboeremia putaminum (Speg.) Qian Chen and L. Cai	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
Stagonosporopsis salicorniae (Magnus) Died.	–	–	Salicornia europaea	–	[35]
	–	–	Salicornia patula	–	[35]
Didymosphaeriaceae
Didymosphaeria lignomaris Strongman and J.D. Mill.	Basal area of the sheath	Saprobic	Spartina densiflora	Argentina: Buenos Aires	[64]
	–	–	Spartina spp.	–	[32]
Julella herbatilis Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Senescent leaves	Saprobic	Juncus roemerianus	USA: North Carolina	[76]
Paraphaeosphaeria apicicola Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Senescent leaves	Saprobic	Juncus roemerianus	USA: North Carolina	[51]
Paraphaeosphaeria pilleata Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Senescent culms	Saprobic	Juncus roemerianus	USA: North Carolina	[77]
Paraphaeosphaeria michotii (Westend.) O.E. Erikss.	–	–	Elymus pungens	UK	[38]
	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
Pseudopithomyces atro-olivaceus (Cooke and Harkn.) G. Guevara, K.C. Cunha and Gené	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Pseudopithomyces chartarum (Berk. and M.A. Curtis) Jun F. Li, Ariyaw. and K.D. Hyde	Senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Pseudopithomyces maydicus (Sacc.) Jun F. Li, Ariyaw. and K.D. Hyde	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Spegazzinia tessarthra (Berk. and M.A. Curtis) Sacc.	Living leaves		Juncus roemerianus	USA: Florida	[43]
	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Tremateia halophila Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Lower and middle parts of senescent culms	Saprobic	Juncus roemerianus	USA: North Carolina	[78]
	–	Saprobic	Spartina marítima	Portugal: Alentejo, Lisbon	[54]
Lentitheciaceae
Halobyssothecium estuariae B. Devadatha, Calabon, K.D. Hyde and E.B.G. Jones	Dead culm	Saprobic	Phragmites australis	UK: Pembrokeshire	[79]
Halobyssothecium obiones (P. Crouan and H. Crouan) Dayarathne, E.B.G. Jones and K.D. Hyde	Drift stems, attached and dead culms	Saprobic	Spartina alterniflora	India: Maharashtra, Tamil Nadu, Andhara Pradesh; USA: Maine, Rhode Island, Connecticut, Massachusetts, New Jersey, Maryland, Virginia, North Carolina, South Carolina, Florida, Mississippi, Texas	[20,35,52,61,71,74,80,81,82]
	–	–	Spartina cynosuroides	–	[35]
	Pod and rhizome	Saprobic	Spartina densiflora	Argentina: Buenos Aires	[64]
	–	Saprobic	Spartina patens	USA: Rhode Island	[36]
	Culms	Saprobic	Spartina sp.	UK: England, Hampshire	[79,83]
	Stem	Saprobic	Spartina townsendii	UK: Hampshire, Wales	[49,65]
	–	Saprobic	Spartina spp.	USA: New Jersey, South Carolina; Mississippi, Argentina: Buenos Aires	[32,35,36,58,84]
	Stem, leaf sheaths, and blades	Saprobic	Spartina marítima	Portugal: Alentejo, Lisbon, Algarve, Centro	[31,54,59,63]
	–	Saprobic	Unidentified saltmarsh plants	USA: Mississippi	[55,58]
	–	–	Elymus pungens	–	[35]
	–	–	Atriplex portulacoides	–	[35]
	–	–	Spartina townsendii	–	[35]
Halobyssothecium phragmitis M.S. Calabon, E.B.G. Jones, S. Tibell and K.D. Hyde	Dead culm and stem	Saprobic	Phragmites sp.	Sweden: Gotland	[85]
Halobyssothecium versicolor M.S. Calabon, E.B.G. Jones and K.D. Hyde	Dead stem	Saprobic	Atriplex portulacoides	UK: Hampshire	[85]
Keissleriella culmifida (P. Karst.) S.K. Bose	–	–	Elymus pungens	UK	[38]
Keissleriella linearis E. Müll. ex Dennis	Living/decomposing stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[40]
	Dead culm	Saprobic	Phragmites sp.	Sweden: Gotland	[85]
Keissleriella phragmiticola Wanas., E.B.G. Jones and K.D. Hyde	Culms	Saprobic	Phragmites australis	UK: Wales	[79]
Keissleriella rara Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Senescent culms	Saprobic	Juncus roemerianus	USA: North Carolina	[77]
Keissleriella spp.	Senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Lentithecium fluviatile (Aptroot and Van Ryck.) K.D. Hyde, J. Fourn. and Ying Zhang	Dead leaf sheaths	Saprobic	Phragmites australis	Belgium: East Flanders	[86]
	Living/decomposing leaf blades and sheaths, stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40,50]
Setoseptoria arundinacea (Sowerby) Kaz. Tanaka and K. Hiray.	–	–	Elymus pungens	UK	[38]
	Living/decomposing leaf blades and sheaths, stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40,50]
	–	Saprobic	Spartina sp.	USA: North Carolina, Florida	[71]
Setoseptoria phragmitis Quaedvl., Verkley and Crous	Culm	Saprobic	Phragmites sp.	Sweden: Södermanland	[87]
Towyspora aestuari Wanas., E.B.G. Jones and K.D. Hyde	–	–	Phragmites australis	UK: Wales	[88]
Leptosphaeriaceae
Leptosphaeria albopunctata (Westend.) Sacc.	–	–	Juncus maritimus	–	[35]
	–	–	Phragmites australis	–	[35]
	Attached culms	-	Spartina alterniflora	USA: Rhode Island	[35,36,61,71,73,80]
	–	–	Spartina spp.	Canada: Bay of Fundy; USA: New Jersey, South Carolina; Argentina: Buenos Aires	[35,36,48,89,90]
	Stem	Saprobic	Spartina townsendii	UK: Wales	[35,65]
Leptosphaeria australiensis (Cribb and J.W. Cribb) G.C. Hughes	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Pod	Saprobic	Spartina densiflora	Argentina: Buenos Aires	[64]
	–	–	Spartina spp.	–	[32]
Leptosphaeria culmifraga (Fr.) Ces. and De Not.	–	–	Elymus pungens	UK	[38]
Leptosphaeria littoralis Sacc.	–	–	Elymus pungens	UK	[38]
Leptosphaeria marina Ellis and Everh.	–	–	Juncus roemerianus		[35]
	–	Saprobic	Spartina alterniflora	USA: Maine, Rhode Island, Connecticut, New Jersey, Delaware, Virginia, North Carolina, South Carolina	[35,36,71,73,80]
	–	Saprobic	Spartina spp.	Canada; USA: New Jersey	[32,35,36,65,89,90,91]
	–	–	Spartina townsendii	UK	[35,38]
	Leaf sheaths and blades, stem	Saprobic	Spartina maritima	Portugal: Algarve	[31,59]
Leptosphaeria orae-maris Linder	–	–	Arundo donax	–	[35]
	–	Saprobic	Lysimachia maritima	USA: Massachusetts	[35,92]
	–	Saprobic	Spartina alterniflora	USA: Massachusetts, Rhode Island, North Carolina, Florida, Texas	[36,71,80,92]
	Rhizome	Saprobic	Spartina densiflora	Argentina: Buenos Aires	[64]
	–	–	Spartina spp.	–	[32]
	–	Saprobic	Spartina townsendii	UK	[35,65,93]
Leptosphaeria pelagica E.B.G. Jones	–	–	Elymus pungens	UK	[35,38]
	–	–	Puccinellia maritima	UK	[38]
	Decaying herbaceous stems, dead culms, decaying leaves	Saprobic	Spartina alterniflora	USA: Connecticut, Mississippi, Rhode Island; India: Goa, Karanataka	[20,36,52,55,73,94]
	–	Saprobic	Spartina densiflora	Argentina: Buenos Aires	[64]
	–	Saprobic	Spartina patens	USA: Rhode Island	[36]
	–	–	Spartina townsendii	UK	[38]
	–	–	Spartina spp.	UK	[32,65]
	Sheath	Saprobic	Spartina marítima	Portugal: Alentejo, Lisbon	[54]
	Stem	Saprobic	Spartina marítima	Portugal: Alentejo	[63]
Leptosphaeria peruvianae Speg.	Decaying stems	Saprobic	Sarcocornia perennis	Argentina: Buenos Aires; in temperate marine waters	[52]
Leptosphaeria spp.	Decaying leaves	Saprobic	Juncus roemerianus	USA: Mississippi	[55]
	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
	–	–	Spartina alterniflora	USA: Rhode Island	[74]
	Leaf sheaths and blades, stem	Saprobic	Spartina maritima	Portugal: Centro	[59]
Leptosphaeria suaedae Hansf.	Dead twigs	Saprobic	Suaeda australis	South Australia	[95]
Lindgomycetaceae
Arundellina typhae Wanas., E.B.G. Jones and K.D. Hyde	Dead stem	Saprobic	Typha sp.	UK: England	[96]
Lophiostomataceae
Lophiostoma semiliberum (Desm.) Ces. and De Not.	Living/decomposing stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[40]
Lophiostoma sp.	–	–	Elymus pungens	UK	[38]
Sigarispora arundinis (Pers.) Thambug., Qing Tian, Kaz. Tanaka and K.D. Hyde	Living/decomposing stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[40]
Massarinaceae
Helminthosporium sp.	Decaying leaf blades	Saprobic	Spartina alterniflora	USA: Georgia	[56]
Massarina carolinensis Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Senescent culms	Saprobic	Juncus roemerianus	USA: North Carolina	[77]
Massarina igniaria (C. Booth) Aptroot	Decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Massarina phragmiticola Poon and K.D. Hyde	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Massarina ricifera Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Lower parts of senescent culms, decaying leaves	Saprobic	Juncus roemerianus	USA: Alabama, Mississippi, North Carolina	[55,58,97]
Massarina spp.	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
Stagonospora abundata Kohlm. and Volkm.-Kohlm.	Senescent leaves and bracts	Saprobic	Juncus roemerianus	USA: Florida, Georgia, North Carolina	[98]
Stagonospora cylindrica Gunnell	Living/decomposing stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[40]
Stagonospora elegans (Berk.) Sacc. and Traverso	Living/decomposing leaf sheaths, stems, culms	Saprobic	Phragmites australis	Australis; Netherlands: Zeeland	[39,40,95]
Stagonospora epicalamia (Cooke) Sacc.	–	–	Phragmites australis	Australia	[95]
Stagonospora haliclysta Kohlm.	Leaf sheaths and blades, stem	Saprobic	Spartina maritima	Portugal: Algarve	[59]
Stagonospora spp.	Living and senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Living/decomposing leaf blades and sheaths, stems	Saprobic	Phragmites australis	China: Hong Kong; Netherlands: Zeeland	[39,40,41,50]
	Senescent and dead leaves/inflorescence, living and dead seeds, decaying leaf blades	Saprobic, pathogenic	Spartina alterniflora	Canada; USA: Maine, Rhode Island, Georgia, Connecticut, New Jersey, Virginia, Florida, North Carolina; Argentina: Buenos Aires	[35,36,56,73,74]
	–	Pathogenic	Spartina cynosuroides	USA: Maryland	[44]
	–	Saprobic	Spartina patens	USA: Rhode Island	[35,36]
	–	Saprobic	Spartina spp.	Canada	[35,36]
	Leaf sheaths and blades, stem, limb	Saprobic	Spartina marítima	Portugal: Alentejo, Lisbon, Algarve, Centro	[31,54,59]
Stagonospora suaedae Syd. and P. Syd.	Leaves	–	Suaeda marítima	Germany	[99]
Melanommataceae
Aposphaeria spp.	Living/decomposing leaf sheaths, stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40]
Bicrouania maritima (P. Crouan and H. Crouan) Kohlm. and Volkm.-Kohlm.	Dead stems	Saprobic	Atriplex portulacoides	India	[35,52]
Morosphaeriaceae
Helicascus kanaloanus Kohlm.	–	–	Spartina spp.	–	[32]
Neocamarosporiaceae
Neocamarosporium artemisiae Dayarathne and E.B.G. Jones	–	Saprobic	Artemisia maritima	Sweden: Bohuslän	[100]
Neocamarosporium maritimae Dayarathne and E.B.G. Jones	–	Saprobic	Artemisia maritima	Sweden: Bohuslän	[100]
Neocamarosporium obiones (Jaap) Wanas. and K.D. Hyde	–	–	Atriplex portulacoides	–	[35]
Neocamarosporium phragmitis D.N. Wanasinghe, E.B.G. Jones and K.D. Hyde	Decaying culms	Saprobic	Phragmites australis	UK	[101]
Neocamarosporium salicorniicola Dayar., E.B.G. Jones and K.D. Hyde	Dead stems	Saprobic	Salicornia sp.	Thailand	[102]
Periconiaceae
Periconia cookei E.W. Mason and M.B. Ellis	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Living/decomposing leaf blades and sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,50]
Periconia digitata (Cooke) Sacc.	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Periconia digitata (Cooke) Sacc.	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
Periconia echinochloae (Bat.) M.B. Ellis	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Periconia minutissima Corda	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
Periconia sp.	–	Saprobic	Unidentifed saltmarsh plants	USA: Mississippi	[58]
Phaeosphaeriaceae
Amarenomyces ammophilae (Lasch) O.E. Erikss.	–	–	Ammophila arenaria	–	[35]
	–	–	× Ammocalamagrostis baltica	–	[35]
	–	–	Uniola paniculata	–	[35]
Amphisphaeria culmicola Sacc.	Stem		Spartina townsendii	UK: England	[49]
Camarosporioides phragmitis W.J. Li and K.D. Hyde	Dead stem	Saprobic	Phragmites australis	Germany	[96]
Hendersonia culmiseda Sacc.	Living/decomposing leaf blades	Saprobic	Phragmites australis	Netherlands: Zeeland	[50]
	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
	–	–	Spartina townsendii	UK	[103]
Hendersonia spp.	Living/decomposing leaf blades and sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland; USA: Florida	[39,43,50]
Loratospora aestuarii Kohlm. and Volkm.-Kohlm.	Senescent culms	Saprobic	Juncus roemerianus	USA: North Carolina	[104]
Loratospora aestuarii Kohlm. and Volkm.-Kohlm.	–	Saprobic	Unidentified saltmarsh plants	USA: Mississippi	[58]
Ophiobolus littoralis (P. Crouan and H. Crouan) Sacc.	–	–	Elymus pungens	UK	[38]
Phaeoseptoria sp.	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
Phaeosphaeria anchiala Kohlm., Volkm.-Kohlm. and C.K.M. Tsui	Senescent leaves	Saprobic	Juncus roemerianus	USA: Florida, Georgia, Maryland, North Carolina, Virginia	[105]
Phaeosphaeria caricinella (P. Karst.) O.E. Erikss.	–	–	Spartina sp.	USA: Florida, North Carolina	[71]
Phaeosphaeria culmorum (Auersw.) Leuchtm.	Living/decomposing leaf blades and sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,50]
Phaeosphaeria eustoma (Fuckel) L. Holm	Living/decomposing leaf blades and sheaths, stems, culms	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40,50,95]
Phaeosphaeria fuckelii (Niessl) L. Holm	–	–	Elymus pungens	UK	[38]
Phaeosphaeria gessneri Shoemaker and C.E. Babc.	–	–	Spartina spp.	–	[32]
Phaeosphaeria halima (T.W. Johnson) Shoemaker and C.E. Babc.	Dead culms; Decaying leaves, leaf blades	Saprobic	Spartina alterniflora	India: Kerala; USA: California, Georgia, Mississippi, Vancouver, North Carolina	[25,35,52,55,56,57,58,71,80]
	Decaying leaf blades	Saprobic	Spartina densiflora	USA: California	[25]
			Spartina spp.		[32]
	Decaying leaves	Saprobic	Spartina foliosa	USA: California	[25]
	Leaf sheaths and blades, stem	Saprobic	Spartina maritima	Portugal: Algarve, Centro	[31]
Phaeosphaeria herpotrichoides (De Not.) L. Holm	–	–	Spartina patens	USA: North Carolina, Florida	[71]
Phaeosphaeria juncina (Auersw.) L. Holm	–	Saprobic	Juncus roemerianus	USA: Florida	[43]
Phaeosphaeria luctuosa (Niessl ex Sacc.) Y. Otani and Mikawa	Living/decomposing leaf sheaths, stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40]
	–	–	Elymus pungens	UK	[38]
Phaeosphaeria macrosporidium (E.B.G. Jones) Shoemaker and C.E. Babc.	Decaying stems	Saprobic	Spartina sp	UK: Wales, England	[65]
	Stem	Saprobic	Spartina marítima	Portugal: Lisbon	[54,63]
Phaeosphaeria microscopica (P. Karst.) O.E. Erikss.	–	–	Elymus pungens	UK	[38]
Phaeosphaeria neomaritima (R.V. Gessner and Kohlm.) Shoemaker and C.E. Babc.	–	–	Juncus maritimus	–	[35]
	–	–	Juncus roemerianus	–	[35]
	–	Saprobic	Juncus sp.	Canada; India: Maharashtra, Karnataka; USA: Virginia, North Carolina	[36,52,71,80]
	–	–	Spartina alterniflora	–	[35]
	–	Saprobic	Spartina spp.	Canada; USA: North Carolina, Virginia	[32,71,80]
	–	–	Spartina townsendii	UK	[35,93]
	Stem	Saprobic	Spartina marítima	Portugal: Alentejo	[63]
Phaeosphaeria nigrans (Roberge ex Desm.) L. Holm	–	–	Elymus pungens	UK	[38]
Phaeosphaeria olivacea Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Senescent leaves	Saprobic	Juncus roemerianus	USA: North Carolina, Mississippi	[58,76]
Phaeosphaeria pontiformis (Fuckel) Leuchtm.	–	–	Elymus pungens	UK	[38]
	Living/decomposing leaf blades and sheaths, stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40,50]
Phaeosphaeria roemeriani Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Mississippi, North Carolina	[55,58,60]
Phaeosphaeria spartinae (Ellis and Everh.) Shoemaker and C.E. Babc.	–	Saprobic	Spartina spp.	India: Kerala	[32,52]
	Decaying herbaceous stems and pod	Saprobic	Spartina densiflora	Argentina: Buenos Aires	[64]
	–	Saprobic	Spartina marítima	Portugal: Lisbon	[54]
Phaeosphaeria spartinicola Leuchtm.	–	Saprobic	Juncus sp.	India	[52]
	Dead leaves, decaying leaf blades	Saprobic	Spartina alterniflora	Mexico; USA: Alabama, California, Georgia, Mississippi; Canada: Nova Scotia, New Brunswick	[25,36,55,56,57,58]
	Pod, leaf blades	Saprobic	Spartina densiflora	Argentina: Buenos Aires; USA: California	[25,64]
	–	–	Spartina spp.	–	[32]
	Leaf blades	Saprobic	Spartina foliosa	USA: California	[25]
	Leaf sheaths and blades, stem, limb	Saprobic	Spartina marítima	Portugal: Alentejo, Lisbon, Algarve, Centro	[31,54,59,63]
Phaeosphaeria spp.	Living/decomposing leaf blades and sheaths, stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40,50]
	–	Saprobic	Spartina alterniflora	USA: Rhode Island	[74]
Sclerostagonospora sp.	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Septoriella phragmitis Oudem.	Living/decomposing leaf sheaths and stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40]
Septoriella spp.	Decaying stems and leaf sheaths and blades, stems	Saprobic	Phragmites australis	China: Hong Kong; Netherlands: Zeeland	[39,40,41,50]
Septoriella thalassica (Speg.) Nag Raj	–	–	Distichlis spicata	–	[35]
			Distichlis spicata		[35]
Septoriella unigalerita Kohlm. and Volkm.-Kohlm.	Senescent leaves	Saprobic	Juncus roemerianus	USA: North Carolina	[98]
Septoriella vagans (Niessl) Y. Marín and Crous	–	–	Elymus pungens	UK	[38]
	–	–	Puccinellia maritima	UK	[38]
	–	Saprobic	Spartina alterniflora	USA: Rhode Island	[74]
Pleomassariaceae
Splanchnonema sp.	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Pleosporaceae
Alternaria alternata (Fr.) Keissl.	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Living/decomposing leaf blades and sheaths, stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,41,50]
	–	Saprobic	Spartina alterniflora	USA: North Carolina	[74]
	Leaves, stems, and roots	Saprobic	Spartina sp.	Canada: Bay of Fundy	[48]
Alternaria infectoria E.G. Simmons	–	–	Elymus pungens	UK	[38]
	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
Alternaria longissima Deighton and MacGarvie	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Alternaria maritima G.K. Sutherl.	Stem	Saprobic, pathogenic	Spartina townsendii	UK: England	[49]
Alternaria spp.	–	–	Atriplex portulacoides	–	[35]
	–	–	Juncus roemerianus	–	[35]
	–	–	Salsola kali	–	[35]
	Inflorescence and upper leaves	Saprobic, parasitic	Spartina alterniflora	USA: Rhode Island	[35,61]
	Culms	Saprobic	Spartina sp.	Thailand	This study
	–	–	Spartina townsendii	–	[35]
Bipolaris cynodontis (Marignoni) Shoemaker	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
Curvularia hawaiiensis (Bugnic. ex M.B. Ellis) Manamgoda, L. Cai and K.D. Hyde	Living and senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Curvularia protuberata R.R. Nelson and Hodges	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
	Senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Curvularia spp.	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	–	Saprobic	Spartina altrerniflora	USA: North Carolina	[74]
Curvularia tuberculata B.L. Jain	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Decorospora gaudefroyi (Pat.) Inderb., Kohlm. and Volkm.-Kohlm.	Stems	Saprobic	Atriplex sp.	UK: Portsmouth	[106]
	–	–	Atriplex portulacoides	–	[35]
	–	–	Sarcocornia perennis	–	[35]
	–	–	Sarcoconia fructicosa	–	[35]
	–	–	Salicornia europaea	–	[35]
	–	–	Salicornia sp.	–	[35]
	Leaf sheaths and blades, stem	Saprobic	Spartina maritima	Portugal: Algarve	[59]
	–	–	Suaeda maritima	–	[35]
Drechslera sp.	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Exserohilum rostratum (Drechsler) K.J. Leonard and Suggs	–	–	Distichlis spicata	–	[35]
	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Senescent and dead leaves	Saprobic	Spartina alterniflora	USA: Rhode Island, North Carolina, Florida	[35,36,73]
	–	–	Spartina spp.	–	[32]
Paradendryphiella arenariae (Nicot) Woudenb. and Crous	Decomposing culms	Saprobic	Spartina alterniflora	USA: Rhode Island	[35,61]
	–	–	Spartina spp.	–	[32]
Paradendryphiella salina (G.K. Sutherl.) Woudenb. and Crous	–	–	Atriplex portulacoides	–	[35]
	Decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	–	–	Puccinellia maritima	–	[35]
	–	–	Salicornia europaea	–	[35]
	Decomposing culms	Saprobic	Spartina alterniflora	USA: Rhode Island	[35,61]
	–	–	Spartina spp.	–	[32]
	–	–	Spartina townsendii	–	[35]
	Leaves and stems	Saprobic	Spartina sp.	Canada: Bay of Fundy	[48]
	–	–	Suaeda maritima	–	[35]
Pleospora abscondita Sacc. and Roum.	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
Pleospora pelagica T.W. Johnson	Decomposing culms; decaying leaf blades	Saprobic	Spartina alterniflora	India: Maharashtra, Kerala; USA: Georgia, Rhode Island, North Carolina, Florida	[35,36,52,56,71,73,74,80]
	Decaying leaf blades	Saprobic	Spartina densiflora	USA: California	[25]
		Saprobic	Spartina spp.	USA: South Carolina	[32,36]
			Typha sp.		[35]
Pleospora pelvetiae G.K. Sutherl.	–	Saprobic	Unidentifed saltmarsh plants	USA: Mississippi	[58]
Pleospora spp.	–	–	Salicornia virginica	–	[35]
	Dead leaves/culms	Saprobic	Spartina alterniflora	USA: Rhode Island	[61]
Pleospora spartinae (J. Webster and M.T. Lucas) Apinis and Chesters	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
	Decaying leaf blades	Saprobic	Spartina alterniflora	USA: Georgia	[56]
	Stem	Saprobic	Spartina spp.	Canada: Bay of Fundy	[32,48]
	–	–	Spartina townsendii	UK	[35,38,107]
Pleospora straminis Sacc. and Speg.	–	–	Elymus pungens	UK	[38]
Pleospora vagans Niessl var. vagans	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
	Dead culms	Saprobic	Spartina alterniflora	USA: Rhode Island	[73]
Pyrenophora tritici-repentis (Died.) Drechsler	–	–	Elymus pungens	UK	[38]
Stemphylium botryosum Wallr.	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
Stemphylium lycopersici (Enjoji) W. Yamam.	Living leaves	–	Juncus roemerianus	USA: Florida	[43]
Stemphylium maritimum T.W. Johnson	–	Saprobic	Spartina sp.	UK	[65]
Stemphylium spp.	–	–	Salsola kali	–	[35]
	Leaves	Saprobic	Spartina spp.	Canada: Bay of Fundy	[35,48]
Stemphylium vesicarium (Wallr.) E.G. Simmons	–	–	Elymus pungens	UK	[38]
	Living, senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	–	Saprobic	Lysimachia maritima	USA: Massachusetts	[92]
	–	Saprobic	Spartina alterniflora	USA: Rhode Island	[61]
	Glumes, rachis	–	Spartina townsendii	UK: England	[38,49]
			Spartina sp.	UK	[65]
Stemphylium triglochinicola B. Sutton and Piroz.	–	–	Triglochin maritima	Sweden: Västergötland	[35,87]
	Dead leaves and inflorescences	Saprobic	Triglochin sp.	India: Kerala; UK	[52,108]
Typhicola typharum (Desm.) Crous	Senescent and dead leaves	Saprobic, pathogenic	Spartina alterniflora	Canada; USA: Maine, Rhode Island, Connecticut, New Jersey, Virginia, North Carolina, Florida	[35,36,61,73,74]
	–	Saprobic	Spartina patens	USA: Rhode Island	[36]
	–		Spartina townsendii	UK	[38]
	–	Saprobic	Spartina spp.	Argentina: Buenos Aires; Canada; USA: Maine	[35,36]
	Stems	Saprobic	Spartina townsendii	UK: England	[35,49,65]
Pleosporales genera incertae sedis
Phialophorophoma litoralis Linder	Stem and sheath	Saprobic	Spartina marítima	Portugal: Alentejo, Lisbon	[54,63]
Phialophorophoma spp.	Living/decomposing leaf sheaths, stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40]
Pyrenochaeta sp.	Living leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Scolecobasidium humicola G.L. Barron and L.V. Busch	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Roussoellaceae
Cytoplea sp.	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Sporormiaceae
Preussia funiculata (Preuss) Fuckel	–	–	Spartina townsendii	UK	[38]
Preussia terricola Cain	–	–	Elymus pungens	UK	[38]
Sporormia longipes Massee and E.S. Salmon	–	–	Elymus pungens	UK	[38]
Sporormia sp.	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Sporormiella intermedia (Auersw.) S.I. Ahmed and Cain ex Kobayasi	–	–	Elymus pungens	UK	[38]
Sporormiella lageniformis (Fuckel) S.I. Ahmed and Cain	–	–	Spartina townsendii	UK	[38]
Sporormiella minima (Auersw.) S.I. Ahmed and Cain	–	–	Elymus pungens	UK	[38]
	–	–	Spartina townsendii	UK	[38]
Teichosporaceae
Teichospora striata (Kohlm. and Volkm.-Kohlm.) Jaklitsch and Voglmayr	Senescent leaves and inflorescences	Saprobic	Juncus roemerianus	USA: North Carolina, Virginia	[98]
Teichospora suaedae Speg.	Dead branches	Saprobic	Suaeda divaricata	Argentina: Mendoza	[109]
Testudinaceae
Verruculina enalia (Kohlm.) Kohlm. and Volkm.-Kohlm.	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Tetraplosphaeriaceae
Tetraploa aristata Berk. and Broome	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Torulaceae
Torula herbarum (Pers.) Link	Decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Trematosphaeriaceae
Halomassarina thalassiae (Kohlm. and Volkm.-Kohlm.) Suetrong, Sakay., E.B.G. Jones, Kohlm., Volkm.-Kohlm. and C.L. Schoch	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
EUROTIOMYCETES
Chaetothyriales
Herpotrichiellaceae
Rhinocladiella spp.	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Veronaea sp.	Decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Eurotiales
Aspergillaceae
Aspergillus fumigatus Fresen.	–	–	Elymus pungens	UK	[38]
Aspergillus nidulans (Eidam) G. Winter	–	–	Elymus pungens	UK	[38]
	–	–	Spartina townsendii	UK	[38]
Aspergillus niger Tiegh.	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Aspergillus spp.	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	–	–	Spartina townsendii	UK: England	[49]
Monascus purpureus Went	–	–	Elymus pungens	UK	[38]
Penicillium aurantiogriseum Dierckx	Leaves	Saprobic	Spartina sp.	Canada: Bay of Fundy	[48]
Penicillium brevicompactum Dierckx	Roots	Saprobic	Spartina sp.	Canada: Bay of Fundy	[48]
Penicillium chrysogenum Thom	Roots	Saprobic	Spartina sp.	Canada: Bay of Fundy	[48]
Penicillium lividum Westling	Leaves and stems	Saprobic	Spartina sp.	Canada: Bay of Fundy	[48]
Penicillium spp.	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
	–	–	Spartina townsendii	UK: England	[49]
Thermoascaceae
Thermoascus crustaceus (Apinis and Chesters) Stolk	–	–	Elymus pungens	UK	[38]
Paecilomyces spp.	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
	–	Saprobic	Salt marsh plants	India: Goa	[52]
Trichocomaceae
Thermomyces dupontii (Griffon and Maubl.) Houbraken and Samson	–	–	Elymus pungens	UK	[38]
Onygenales
Onygenaceae
Amauroascus albicans (Apinis) Arx	–	–	Elymus pungens	UK	[38]
Amauroascus albicans (Apinis) Arx	–	–	Spartina townsendii	UK	[38]
LECANOROMYCETES
Ostropales
Stictidaceae
Glomerobolus gelineus Kohlm. and Volkm.-Kohlm. Stictis sp.	Senescent culms	Saprobic	Juncus roemerianus	USA: North Carolina	[110]
	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
LEOTIOMYCETES
Helotiales
Amorphothecaceae
Amorphotheca resinae Parbery	Roots	Saprobic	Spartina sp.	Canada: Bay of Fundy	[48]
Calloriaceae
Cistella fugiens (W. Phillips) Matheis	Living/decomposing stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[40]
Helotiaceae
Cyathicula culmicola (Desm.) De Not.	–	–	Elymus pungens	UK	[38]
Helotium sp.	–	–	Elymus pungens	UK	[38]
Lachnaceae
Brunnipila palearum (Desm.) Baral	–	–	Elymus pungens	UK	[38]
	–	–	Spartina townsendii	UK	[38]
Lachnum controversum (Cooke) Rehm	–	–	Elymus pungens	UK	[38]
Lachnum spartinae S.A. Cantrell	Decaying leaf sheaths	Saprobic	Spartina alterniflora	USA: Georgia	[56,111]
	–	–	Spartina spp.	–	[32]
Mollisiaceae
Belonopsis atriella (Cooke) Lindau	–	–	Spartina cynosuroides	USA: Louisiana	[68,90,112]
Mollisia hydrophila (P. Karst.) Sacc.	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
Mollisia palustris (P. Karst.) P. Karst.	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
Trichobelonium kneiffii (Wallr.) J. Schröt.	Living/decomposing leaf sheaths, stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40]
Ploettnerulaceae
Cadophora melinii Nannf.	Leaves	Saprobic	Spartina sp.	Canada: Bay of Fundy	[48]
Sclerotiniaceae
Botrytis cinerea Pers.	Stem		Spartina townsendii	UK: England	[49]
	Leaves	Saprobic	Spartina sp.	Canada: Bay of Fundy	[48]
Monilia sp.	Decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Solenopeziaceae
Halenospora varia (Anastasiou) E.B.G. Jones	Senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Basal area of the sheath	Saprobic	Spartina densiflora	Argentina: Buenos Aires	[64]
	–	–	Spartina spp.	–	[32]
Helotiales genera incertae sedis
Cejpia hystrix (De Not.) Baral	–	–	Elymus pungens	UK	[38]
Dactylaria sp.	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Crocicreas gramineum (Fr.) Fr.	–	–	Elymus pungens	UK	[38]
Leotiales
Leotiales genera incertae sedis
Flagellospora sp.	Living leaves	–	Juncus roemerianus	USA: Florida	[43]
Rhytismatales
Rhytismataceae
Lophodermium arundinaceum (Schrad.) Chevall.	–	–	Elymus pungens	UK	[38]
	Living/decomposing leaf sheaths, stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40]
Thelebolales
Thelebolaceae
Thelebolus crustaceus (Fuckel) Kimbr.	–	–	Elymus pungens	UK	[38]
	–	–	Puccinellia maritima	UK	[38]
	–	–	Spartina townsendii	UK	[38]
ORBILIOMYCETES
Orbiliales
Orbiliaceae
Arthrobotrys conoides Drechsler	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Arthrobotrys sp.	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Orbilia junci Kohlm., Baral and Volkm.-Kohlm.	Tips of senescent leaves	–	Juncus roemerianus	USA: North Carolina	[113]
PEZIZOMYCETES
Pezizales
Pezizaceae
Belonium heteromorphum (Ellis and Everh.) Seaver	–	–	Spartina cynosuroides	USA: Louisiana	[68,114]
SACCHAROMYCETES
Saccharomycetales
Debaryomycetaceae
Debaryomyces hansenii (Zopf) Lodder and Kreger-van Rij	Decaying leaf blades	Saprobic	Spartina alterniflora	USA: Louisiana	[56]
Scheffersomyces spartinae (Ahearn, Yarrow and Meyers) Kurtzman and M. Suzuki	Decaying leaf blades	Saprobic	Spartina alterniflora	USA: Louisiana	[56]
Saccharomycetaceae
Kluyveromyces lactis (Stell.-Dekk.) Van der Walt	Decaying leaf blades	Saprobic	Spartina alterniflora	USA: Louisiana	[56]
SORDARIOMYCETES
Amphisphaeriales
Amphisphaeriaceae
Massariella sp.	–	–	Spartina townsendii	UK	[38]
Ommatomyces coronatus Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Lower parts of senescent culms	Saprobic	Juncus roemerianus	USA: North Carolina	[97]
Pestalotia sp.	Living, senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Apiosporaceae
Arthrinium arundinis (Corda) Dyko and B. Sutton	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
	Dead culms	Saprobic	Phragmites sp.	South Australia	[62]
Arthrinium phaeospermum (Corda) M.B. Ellis	Living/decomposing leaf blades and sheaths, stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40,50]
	–	Saprobic	Spartina patens	USA: Rhode Island	[61]
	Inflorescence and upper leaves	Saprobic	Spartina alterniflora	USA: Rhode Island	[36]
Arthrinium spp.	Living leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Nigrospora oryzae (Berk. and Broome) Petch	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Beltraniaceae
Beltrania querna Harkn.	Decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Hyponectriaceae
Phragmitensis ellipsoidea M.K.M. Wong, Goh and K.D. Hyde	Intertidal to aerial culms	Saprobic	Phragmites australis	China: Hong Kong	[115]
Phragmitensis marina M.K.M. Wong, Poon and K.D. Hyde	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Physalospora citogerminans Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Lower and upper parts of senescent culms	Saprobic	Juncus roemerianus	USA: North Carolina	[116]
Sporocadaceae
Discostroma sp.	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
Pestalotiopsis juncestris Kohlm. and Volkm.-Kohlm.	Senescent involucral leaves and culms	Saprobic	Juncus roemerianus	USA: North Carolina	[117]
Pestalotiopsis planimi (Vize) Steyaert	–	–	Spartina alterniflora	USA: Rhode Island	[61]
Pestalotiopsis sp.	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Coronophorales
Ceratostomataceae
Melanospora sp.	Decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Microthecium fimicola (E.C. Hansen) Y. Marín, Stchigel, Guarro and Cano	–	–	Elymus pungens	UK	[38]
Microthecium levitum Udagawa and Cain	Dead leaves/culms	Saprobic	Spartina alterniflora	USA: Rhode Island	[61]
Coronophorales genera incertae sedis
Papulaspora halima Anastasiou	Living and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Papulosa amerospora Kohlm. and Volkm.-Kohlm.	Senescent culms	Saprobic	Juncus roemerianus	USA: North Carolina	[118]
Diaporthales
Diaporthaceae
Phomopsis spp.	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
	–	–	Spartina sp.	–	[71]
Gnomoniaceae
Gnomonia salina E.B.G. Jones (probably a nomen dubiumand possibly a Halosarpheia species)	–	Saprobic	Spartina alterniflora	USA: Connecticut	[36]
			Spartina spp.		[32,35]
	–	–	Spartina townsendii	UK	[35,65]
Diaporthales incertae sedis
Botryodiplodia sp.	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Glomerellales
Glomerellaceae
Colletotrichum sp.	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Plectosphaerellaceae
Stachylidium bicolor Link	Senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Hypocreales
Bionectriaceae
Acremonium spp.	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Clonostachys rosea (Link) Schroers, Samuels, Seifert and W. Gams	Leaves	Saprobic	Spartina sp.	Canada: Bay of Fundy	[48]
Fusariella obstipa (Pollack) S. Hughes	Decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Gliomastix spp.	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Hydropisphaera arenula (Berk. and Broome) Rossman and Samuels	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
Hydropisphaera erubescens (Roberge ex Desm.) Rossman and Samuels	Decaying leaf blades	Saprobic	Spartina alterniflora	USA: Georgia	[56]
	–	–	Spartina spp.	–	[32]
Clavicipitaceae
Atkinsonella hypoxylon (Peck) Diehl	–	–	Spartina cynosuroides	–	[68]
Claviceps purpurea (Fr.) Tul.	–	Saprobic	Phragmites australis	UK: England (Southampton Hampshire, Sussex, Oxon)	[119,120]
	Replaced seeds in the inflorescence, ovaries of the flowers	Saprobic, parasitic	Spartina alterniflora	USA: Rhode Island; Argentina	[36,61,68,73,121,122]
	–	Pathogenic	Spartina anglica	UK	[123]
	–	Saprobic, parasitic	Spartina cynosuroides	USA: New York, Florida, Mississippi	[44,68,121,124]
			Spartina patens	USA: Maryland, Mississippi	[44,68,124,125]
	–	–	Spartina townsendii	UK: England	[120,126]
	–	–	Spartina sp.	Argentina	[122]
Claviceps sp.	–	–	Spartina foliosa	USA: California	[127]
Metarhizium anisopliae (Metschn.) Sorokīn	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
Hypocreaceae
Cladobotryum sp.	Decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Gliocladium sp.	Senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Trichoderma citrinum (Pers.) Jaklitsch, W. Gams and Voglmayr	Leaves	Saprobic	Spartina sp.	Canada: Bay of Fundy	[48]
Trichoderma sp.	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Trichoderma viride Pers.	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Nectriaceae
Calonectria sp.	–	–	Elymus pungens	UK	[38]
Fusarium fujikuroi Nirenberg	–	Saprobic	Suaeda australis	South Australia	[62]
Fusarium graminearum Schwabe	Living/decomposing leaf sheaths, stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40]
Fusarium heterosporum Nees and T. Nees	–	–	Spartina maritima	–	[128]
Fusarium incarnatum (Desm.) Sacc.	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
Fusarium oxysporum Schltdl.	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
	Leaves and roots	Saprobic	Spartina sp.	Canada: Bay of Fundy	[48]
Fusarium poae (Peck) Wollenw.	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
Fusarium solani (Mart.) Sacc.	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Fusarium spp.	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Living/decomposing leaf sheaths, stems	Saprobic	Phragmites australis	China: Hong Kong; Netherlands: Zeeland	[39,40,41]
	Leaf sheaths and blades, stem	Saprobic	Spartina maritima	Portugal: Algarve	[59]
Gibberella sp.	–	Saprobic	Spartina alterniflora	Argentina: Buenos Aires	[36]
Nectria sp.	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Tubercularia pulverulenta Speg.	–	–	Sarcocornia perennis	–	[35]
	–	–	Salicornia europaea	–	[35]
	–	Saprobic	Unidentified saltmarsh plants	USA: Mississippi	[58]
	–	–	Sarcocornia fruticosa	–	[35]
Tubercularia sp.	Decaying leaf blades	Saprobic	Spartina alterniflora	USA: Georgia	[56]
Volutella ciliata (Alb. and Schwein.) Fr.	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
Sarocladiaceae
Sarocladium implicatum (J.C. Gilman and E.V. Abbott) A. Giraldo, Gené and Guarro	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
Sarocladium sp.	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Stachybotryaceae
Albifimbria verrucaria (Alb. and Schwein.) L. Lombard and Crous	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
Paramyrothecium roridum (Tode) L. Lombard and Crous	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Stachybotrys chartarum (Ehrenb.) S. Hughes	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Stachybotrys cylindrosporus C.N. Jensen	Decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Stachybotrys echinatus (Rivolta) G. Sm.	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Stachybotrys kampalensis Hansf.	Senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Stachybotrys nephrosporus Hansf.	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Stachybotrys spp.	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
	Decaying leaf blades	Saprobic	Spartina alterniflora	USA: Georgia	[56]
Striaticonidium cinctum (Corda) L. Lombard and Crous	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
Xepicula jollymannii (N.C. Preston) L. Lombard and Crous	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Hypocreales genera incertae sedis
Cephalosporium spp.	Dead leaves/culms	Saprobic	Spartina alterniflora	USA: Rhode Island	[61]
Lulworthiales
Lulworthiaceae
Cumulospora marina I. Schmidt	Dead culm	Saprobic	Phragmites australis	Iraq, Egypt, Germany, Thailand	[129]
	–	–	Spartina spp.	–	[32]
Halazoon fuscus (I. Schmidt) Abdel-Wahab, K.L. Pang, Nagah., Abdel-Aziz and E.B.G. Jones	Decaying rhizomes	Saprobic	Phragmites australis	France, Germany, Japan	[35,130]
	Rhizomes and culms	Saprobic	Phragmites sp.	Sweden	[87]
Halazoon melhae Abdel-Aziz, Abdel-Wahab and Nagahama	Decaying stem	Saprobic	Phragmites australis	Egypt: Port Said	[130]
Lulworthia floridana Meyers	–	Saprobic	Spartina alterniflora	USA: North Carolina, Rhode Island	[20,131]
Lulworthia medusa (Ellis and Everh.) Cribb and J.W. Cribb	–	–	Elymus pungens	UK	[38]
	–	Saprobic	Spartina cynosuroides	USA: New Jersey	[89,132]
	–		Spartina spp.	USA: New Jersey	[32,89]
	Stems	Saprobic	Spartina townsendii	UK: England (Wales); USA: Virginia, North Carolina, South Carolina, Florida, Texas	[38,49,71,72,89,132,133,134]
Lulworthia spp.	–	–	Elymus pungens	–	[35]
	–	–	Juncus roemerianus	–	[35,36]
	Dead culms	Saprobic	Spartina alterniflora	Argentina: Buenos Aires; USA: Rhode Island, North Carolina	[35,36,61,73,74]
	–	–	Spartina cynosuroides	–	[35]
	–	Saprobic	Spartina sp.	Argentina: Buenos Aires; Canada; USA: Maine, North Carolina	[36]
	–	–	Spartina townsendii	–	[35]
	Leaf sheaths and blades, stem	Saprobic	Spartina maritima	Portugal: Alentejo, Lisbon, Algarve, Centro	[31,54,59,63]
Moleospora maritima Abdel-Wahab, Abdel-Aziz and Nagah.	Decayed stems	Saprobic	Phragmites australis	Egypt: Port Said	[130]
Magnaporthales
Ceratosphaeriaceae
Ceratosphaeria sp.	Senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Magnaporthaceae
Buergenerula spartinae Kohlm. and R.V. Gessner	Lower stem and leaf sheath during the growth phase of the plant/living and dead; decaying leaf blades	Saprobic, parasitic	Spartina alterniflora	USA: Alabama, Rhode Island, Maine, New Hampshire, Connecticut, Mississippi, New Jersey, Virginia, North Carolina, Florida, Georgia	[20,35,36,55,56,58,61,73,74,82,92]
	Leaves	Saprobic	Spartina spp.	Canada: Bay of Fundy; USA: South Carolina; UK	[32,35,36,48,65] this study
	Leaf sheaths and blades, stem	Saprobic	Spartina maritima	Portugal: Alentejo, Lisbon, Algarve, Centro	[31,54,59]
Gaeumannomyces sp.	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Kohlmeyeriopsis medullaris (Kohlm., Volkm.-Kohlm. and O.E. Erikss.) Klaubauf, M.-H. Lebrun and Crous	Lower parts of senescent culms	Saprobic	Juncus roemerianus	USA: North Carolina	[97,135]
Utrechtiana roumeguerei (Cavara) Videira and Crous	Living/decomposing leaf blades and sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,50]
Pseudohalonectriaceae
Pseudohalonectria falcata Shearer	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Pseudohalonectria halophila Kohlm. and Volkm.-Kohlm.	Fragments of leaves and culms in the wrack	Saprobic	Juncus roemerianus	USA: North Carolina	[105]
Meliolales
Meliolaceae
Meliola arundinis Pat.	–	–	Phragmites australis	Australia: Queensland	[62]
Microascales
Halosphaeriaceae
Aniptodera chesapeakensis Shearer and M.A. Mill.	Dead leaves	Saprobic	Juncus roemerianus	USA: North Carolina	[35]
	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
	–	–	Spartina alterniflora	–	[35]
	–	–	Spartina spp.	–	[32]
	Leaf sheaths and blades, stem	Saprobic	Spartina maritima	Portugal: Alentejo, Algarve, Centro	[59,63]
Aniptodera juncicola Volkm.-Kohlm. and Kohlm.	Dead standing culms of	Saprobic	Juncus roemerianus	India: Kerala, West Bengal, Tamil Nadu; USA: North Carolina	[52,136]
Aniptodera phragmiticola O. K. Poon et K. D. Hyde	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Ceriosporopsis halima Linde	–	–	Arundo donax	–	[35]
	Submerged seeds	Saprobic	Spartina alterniflora	USA	[137]
	–	–	Spartina spp.	–	[32]
	–	–	Spartina townsendii	UK	[35,38]
	Stem	Saprobic	Spartina maritima	Portugal: Alentejo	[63]
Cirrenalia macrocephala (Kohlm.) Meyers and R.T. Moore	–	–	Ammophila arenaria	–	[35]
	Decaying culms	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Decomposing culms, submerged seeds	Saprobic	Spartina alterniflora	USA: Rhode Island	[35,61,137]
	–	–	Spartina spp.	–	[32]
	Stem	Saprobic	Spartina maritima	Portugal: Alentejo	[63]
Cirrenalia pseudomacrocephala Kohlm.	Senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Corollospora maritima Werderm.	Submerged seeds, decomposing culms	Saprobic	Spartina alterniflora	USA: Rhode Island	[20,35,61,137]
	–	–	Spartina spp.	–	[32]
	Stem	Saprobic	Spartina maritima	Portugal: Alentejo	[63]
	–	Saprobic	Unidentified saltmarsh plants	USA: Mississippi	[58]
Corollospora ramulosa (Meyers and Kohlm.) E.B.G. Jones and Abdel-Wahab	–	Saprobic	Unidentified saltmarsh plants	USA: Mississippi	[58]
	–	Saprobic	Zostera marina	USA: North Carolina	[74]
Haligena elaterophora Kohlm.	–	–	Spartina alterniflora	–	[35]
	–	–	Spartina tonwsendii	UK	[38]
	–	–	Spartina spp.	–	[32]
Halosarpheia culmiperda Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Lower parts of senescent culms	Saprobic	Juncus roemerianus	USA: North Carolina	[97]
Halosarpheia sp.	Stem	Saprobic	Spartina maritima	Portugal: Alentejo	[63]
Halosarpheia viscosa I. Schmidt ex Shearer and J.L. Crane	Decaying leaf blades	Saprobic	Spartina alterniflora	USA: Georgia	[56]
	–	Saprobic	Spartina maritima	Portugal: Lisbon	[54]
Halosphaeria appendiculata Linder	–	–	Arundo donax	–	[35]
Halosphaeria sp.	Submerged seeds	Saprobic	Spartina alterniflora	USA	[137]
Lautisporopsis circumvestita (Kohlm.) E.B.G. Jones, Yusoff and S.T. Moss	–	–	Arundo donax	–	[35]
Lignincola laevis Höhnk	–	–	Elymus pungens	–	[35]
	–	Saprobic	Spartina spp.	USA: North Carolina	[32,138]
	–	–	Spartina townsendii	–	[35]
	Stem	Saprobic	Spartina maritima	Portugal: Alentejo	[63]
Magnisphaera spartinae (E.B.G. Jones) J. Campb., J.L. Anderson and Shearer	–	–	Elymus farctus	–	[35]
	–	–	Elymus pungens	–	[35]
	Living/decomposing stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[40]
	–	Saprobic	Spartina alterniflora	USA: Rhode Island	[20,35,61]
	–		Spartina spp.		[32]
	–	Saprobic	Spartina patens	USA: Rhode Island	[36]
	Stem	Saprobic	Spartina townsendii	UK: Wales	[35,139]
	–		Typha sp.		[35]
Nais inornata Kohlm.	Decomposing culms	Saprobic	Spartina alterniflora	USA: Rhode Island	[20,35,61]
			Spartina spp.		[32]
Natantispora unipolaris K.L. Pang, S.Y. Guo and E.B.G. Jones	Dead stem	Saprobic	Phragmites australis	Taiwan: Nankunshen	[140]
Natantispora retorquens (Shearer and J.L. Crane) J. Campb., J.L. Anderson and Shearer	Leaf sheaths and blades, stem	Saprobic	Spartina maritima	Portugal: Alentejo, Lisbon, Algarve, Centro	[31,54,59,63]
Oceanitis unicaudata (E.B.G. Jones and Camp.-Als.) J. Dupont and E.B.G. Jones	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
	Stem	Saprobic	Spartina maritima	Portugal: Alentejo	[63]
Panorbis viscosus (I. Schmidt) J. Campb., J.L. Anderson and Shearer	Leaf sheaths and blades, stem	Saprobic	Spartina maritima	Portugal: Alentejo, Algarve	[59,63]
Remispora hamata (Höhnk) Kohlm.	–	–	Elymus pungens	UK	[35,38]
	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Living/decomposing leaf blades and sheaths, stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40,50]
	–	Saprobic	Phragmites sp.	Sweden	[87]
	Dead leaves	Saprobic	Spartina alterniflora	USA: Rhode Island, Maine, Florida	[20,35,36,61,73]
	–	Saprobic	Spartina patens	USA: Rhode Island	[36]
	–	Saprobic	Spartina sp.	USA: North Carolina; Argentina: Buenos Aires	[36,138]
	–	–	Spartina townsendii	–	[35]
	–	–	Typha sp.	–	[35]
Remispora trullifera Kohlm.	Leaf sheaths and blades, stem	Saprobic	Spartina maritima	Portugal: Centro	[59]
Tirispora unicaudata E.B.G. Jones and Vrijmoed	Stem	Saprobic	Spartina maritima	Portugal: Alentejo	[63]
Microascaceae
Scopulariopsis spp.	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Myrmecridiales
Myrmecridiaceae
Myrmecridium schulzeri (Sacc.) Arzanlou, W. Gams and Crous	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
Ophiostomatales
Ophiostomataceae
Sporothrix sp.	Senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Phomatosporales
Phomatosporaceae
Phomatospora bellaminuta Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Lower parts of senescent culms	Saprobic	Juncus roemerianus	USA: North Carolina	[116]
Phomatospora berkeleyi Sacc.	Living/decomposing leaf blades and sheaths, stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40,50]
Phomatospora dinemasporium J. Webster	Decaying stems and leaf sheaths, stems	Saprobic	Phragmites australis	China: Hong Kong; Netherlands: Zeeland	[40,41]
	Dead leaves	Saprobic	Phragmites sp.	South Australia	[62]
	–	–	Spartina townsendii	UK	[38]
Phomatospora phragmiticola Poon and K.D. Hyde	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Phomatospora spp.	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Living/decomposing leaf sheaths, stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40]
Phyllachorales
Phyllachoraceae
Phyllachora graminis (Pers.) Fuckel	–	–	Elymus pungens	UK	[38]
	–	Saprobic, pathogenic	Spartina alterniflora	USA: Massachusetts	[44]
	–	–	Spartina cynosuroides	–	[68]
Phyllachora cynodontis Niessl.	–	Saprobic, pathogenic	Spartina alterniflora	USA	[68]
	–	Saprobic, pathogenic	Spartina foliosa	USA: California	[44,112,141]
Phyllachora paludicola Kohlm. and Volkm.-Kohlm.	Dead leaves (lower half of standing culms)	Saprobic	Spartina alterniflora	USA: Florida, Georgia, North Carolina, Maryland, Delaware	[142]
Phyllachora sylvatica Sacc. and Speg.	–	Saprobic	Spartina patens	USA: South Carolina	[141]
Savoryellales
Savoryellaceae
Savoryella paucispora (Cribb and J.W. Cribb) J. Koch	–	–	Elymus pungens	–	[35]
	–	–	Juncus roemerianus	–	[35]
	–	–	Spartina alterniflora	–	[35]
	–	–	Spartina sp.	–	[35]
	–	–	Spartina townsendii	–	[35]
Sordariales
Chaetomiaceae
Achaetomium sp.	Decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Chaetomium elatum Kunze	–	–	Elymus pungens	UK	[38]
	–	–	Puccinellia maritima	UK	[38]
	–	–	Spartina townsendii	UK	[38]
	–	–
Chaetomium globosum Kunze	–	–	Elymus pungens	UK	[38]
	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
	–	–	Puccinellia maritima	UK	[38]
	–	–	Spartina townsendii	UK	[38]
Chaetomium spirale Zopf	–	–	Elymus pungens	UK	[38]
Chaetomium thermophilum La Touche	–	–	Elymus pungens	UK	[38]
	–	–	Puccinellia maritima	UK	[38]
	–	–	Spartina townsendii	UK	[38]
Chaetomium sp.	Stem	Saprobic	Typha sp.	UK	This study
Corynascus sepedonium (C.W. Emmons) Arx	–	–	Elymus pungens	UK	[38]
	–	–	Puccinellia maritima	UK	[38]
	–	–	Spartina townsendii	UK	[38]
Dichotomopilus funicola (Cooke) X.Wei Wang and Samson	–	–	Elymus pungens	UK	[38]
	–	–	Spartina alterniflora	USA: Rhode Island	[61]
	–	–	Spartina townsendii	UK	[38]
Dichotomopilus indicus (Corda) X.Wei Wang and Samson	–	–	Elymus pungens	UK	[38]
Humicola sp.	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Thermothielavioides terrestris (Apinis) X. Wei Wang and Houbraken	–	–	Elymus pungens	UK	[38]
	–	–	Puccinellia maritima	UK	[38]
Trichocladium constrictum I. Schmidt	Stem	Saprobic	Spartina maritima	Portugal: Alentejo	[63]
Trichocladium crispatum (Fuckel) X. Wei Wang and Houbraken	–	–	Elymus pungens	UK	[38]
	–	–	Spartina townsendii	UK	[38]
Lasiosphaeriaceae	–	–
Schizothecium hispidulum (Speg.) N. Lundq.	Living/decomposing leaf sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39]
Zopfiella latipes (N. Lundq.) Malloch and Cain	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Sordariaceae
Neurospora calospora (Mouton) Dania García, Stchigel and Guarro	–	–	Elymus pungens	UK	[38]
Sordaria fimicola (Roberge ex Desm.) Ces. and De Not.	–	–	Elymus pungens	UK	[38]
	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
	–	–	Puccinellia maritima	UK	[38]
	–	–	Spartina townsendii	UK	[38]
Sordariomycetes families incertae sedis
Koorchaloma galateae Kohlm. and Volkm.-Kohlm.	Senescent culms	Saprobic	Juncus roemerianus	USA: North Carolina	[117]
Koorchaloma spartinicola V.V. Sarma, S.Y. Newell and K.D. Hyde	Decaying leaf blades	Saprobic	Spartina alterniflora	USA: Georgia	[56]
Koorchaloma sp.	Decaying leaf blades	Saprobic	Spartina alterniflora	USA: Georgia	[56]
Lautospora simillima Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Lower parts of senescent, soft culms	Saprobic	Juncus roemerianus	USA: North Carolina	[78]
Sordariomycetes genera incertae sedis
Aquamarina speciosa Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Senescent culms		Juncus roemerianus	USA: Georgia, North Carolina, Virginia	[77]
Aropsiclus junci (Kohlm. and Volkm.-Kohlm.) Kohlm. and Volkm.-Kohlm.	Senescent culms	Saprobic	Juncus roemerianus	USA: North Carolina	[143]
Zalerion maritima (Linder) Anastasiou	Basal area of the sheath	Saprobic	Spartina densiflora	Argentina: Buenos Aires	[64]
	–	–	Spartina spp.	–	[32]
Ellisembia sp.	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Torpedosporales
Juncigenaceae
Juncigena adarca Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Senescent leaves	Saprobic	Juncus roemerianus	USA: North Carolina	[76]
Moheitospora adarca (Kohlm., Volkm.-Kohlm. and O.E. Erikss.) Abdel-Wahab, Abdel-Aziz and Nagah	Stems	Saprobic	Juncus roemerianus	USA	[130]
Moheitospora fruticosae Abdel-Wahab, Abdel-Aziz and Nagah.	Decayed stems	Saprobic	Suaeda vermiculata	Egypt: Alexandria	[130]
Torpedospora radiata Meyers	–	Saprobic	Unidentified saltmarsh plants	USA: Mississippi	[58]
Tracyllalales
Tracyllaceae
Tracylla spartinae (Peck) Tassi	–	Saprobic, pathogenic	Spartina patens	USA: Mississippi	[44,68]
Xylariales
Diatrypaceae
Cryptovalsa suaedicola Spooner	Dead twigs	Saprobic	Suaeda vermiculata	UK: Great Britain	[144]
Halocryptovalsa salicorniae Dayar. and K.D. Hyde	Dead stem	Saprobic	Salicornia sp.	Thailand: Prachuap Khiri Khan	[145]
Xylariaceae
Anthostomella atroalba Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Senescent leaves	Saprobic	Juncus roemerianus	USA: North Carolina	[60]
Anthostomella lugubris (Roberge ex Desm.) Sacc.	–	–	Elymus pungens	UK	[38]
Anthostomella phaeosticta (Berk.) Sacc.	–	–	Elymus pungens	UK	[38]
Anthostomella poecila Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Lower and upper parts of senescent culms, decaying leaves	Saprobic	Juncus roemerianus	USA: Alabama, Mississippi, North Carolina	[55,58,116]
Anthostomella punctulata (Roberge ex Desm.) Sacc.	Living/decomposing leaf blades and sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,50]
Anthostomella semitecta Kohlm., Volkm.-Kohlm. and O.E. Erikss.	Senescent culms	–	Juncus roemerianus	USA: North Carolina	[116]
Anthostomella spissitecta Kohlm. and Volkm.-Kohlm.	Leaf sheaths of senescent culms	Saprobic	Spartina alterniflora, S. densiflora.	USA: Connecticut, Florida, North Carolina, Rhode Island; Argentina: Buenos Aires	[32]
	–	–	Spartina sp.	–	[32]
	Leaf sheaths and blades, stem	Saprobic	Spartina maritima	Portugal: Algarve	[59]
Anthostomella spp.	–	–	Elymus pungens	UK	[38]
	–	Saprobic	Spartina alterniflora	USA: Connecticut, Florida, North Carolina, Rhode Island; Argentina	[36,61]
	–	–	Spartina townsendii	UK	[38]
Anthostomella torosa Kohlm. and Volkm.-Kohlm.	Senescent culms (restricted to short culms)	Saprobic	Juncus roemerianus	USA: North Carolina	[32]
Geniculosporium sp.	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Rosellinia sp.	Dead leaves/culms	Saprobic	Spartina alterniflora	USA: Rhode Island	[61]
Virgaria nigra (Link) Nees	Senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Zygosporiaceae
Zygosporium gibbum (Sacc., M. Rousseau and E. Bommer) S. Hughes	Decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Zygosporium masonii S. Hughes	Decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Zygosporium sp.	Decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Xylariales genera incertae sedis
Circinotrichum maculiforme Nees	Decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Xylariomycetidae family incertae sedis
Cainiaceae
Atrotorquata lineata Kohlm. and Volkm.-Kohlm.	Senescent culms	Saprobic	Juncus roemerianus	USA: North Carolina	[104]
		Saprobic	Unidentified saltmarsh plant	USA: Mississippi	[58]
Ascomycota genera incertae sedis
Asteromyces cruciatus C. Moreau and Moreau ex Hennebert	–	–	Agropyron sp.	–	[35]
	–	–	Ammophila arenaria	–	[35]
	–	–	Spartina spp.	–	[32,35]
	–	Saprobic	Zostera sp.	USA: California	[74]
Cremasteria cymatilis Meyers and R.T. Moore Nomen dubium	Senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Cytoplacosphaeria phragmiticola Poon and K.D. Hyde	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Cytoplacosphaeria rimosa (Oudem.) Petr.	Living/decomposing leaf sheaths, stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40]
Cytosporina sp.	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Didymosamarospora euryhalina T.W. Johnson and H.S. Gold	Culms	Saprobic	Juncus roemerianus	USA: North Carolina	[146]
Haplobasidion lelebae Sawada ex M.B. Ellis	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Hymenopsis chlorothrix Kohlm. and Volkm.-Kohlm.	Senescent culms	Saprobic	Juncus roemerianus	USA: North Carolina	[147]
Hyphopolynema juncatile Kohlm. and Volkm.-Kohlm.	Senescent leaves	Saprobic	Juncus roemerianus	USA: North Carolina	[148]
Kolletes undulatus Kohlm. and Volkm.-Kohlm.	Senescent leaves and culms	Saprobic	Juncus roemerianus	USA: North Carolina	[105]
Minimidochium parvum Cabello, Aramb. and Cazau	Leaves	Saprobic	Distichlis spicata	Argentina: Buenos Aires	[47]
Monodictys pelagica (T. Johnson) E.B.G. Jones	–	–	Juncus sp.	–	[35]
	Decomposing culms	Saprobic	Spartina alterniflora	USA: Rhode Island	[20,35,61,73]
	–	–	Spartina spp.	–	[32]
Neottiospora sp.	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Octopodotus stupendus Kohlm. and Volkm.-Kohlm.	Dead leaves (lower half of standing culms)	Saprobic	Spartina alterniflora	USA: North Carolina	[142]
Pycnodallia dupla Kohlm. and Volkm.-Kohlm.	Senescent inflorescences (involucral leaves and branchlets)	Saprobic	Juncus roemerianus	USA: North Carolina	[147]
Sphaeronaema sp.	Senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Stauronema sp.	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Tetranacriella papillata Kohlm. and Volkm.-Kohlm.	Senescent leaves	Saprobic	Juncus roemerianus	USA: North Carolina	[117]
Tetranacrium sp.	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
Zythia spp.	Living, senescent, and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Psammina sp.	Senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
BASIDIOMYCOTA
AGARICOMYCETES
Agaricales
Niaceae
Merismodes bresadolae (Grelet) Singer	Living/decomposing stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[40]
Nia globispora Barata and Basilio	Stem	Saprobic	Spartina maritima	Portugal: Alentejo	[63]
Nia vibrissa R.T. Moore and Meyers	Old stem	Saprobic	Spartina alterniflora	USA: North Carolina	[35,149]
	–	Saprobic	Spartina spp.	USA: North Carolina	[32,150]
	Stem	Saprobic	Spartina maritima	Portugal: Alentejo	[63]
AGARICOSTILBOMYCETES
Agaricostilbales
Chionosphaeraceae
Stilbum sp.	Decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
BARTHELETIOMYCETES
Sebacinales
Sebacinaceae
Chaetospermum camelliae Agnihothr.	Decaying stems and leaf sheaths	Saprobic	Phragmites australis	China: Hong Kong	[41]
MICROBOTRYOMYCETES
Sporidiobolales
Sporidiobolaceae
Sporobolomyces roseus Kluyver and C.B. Niel	Leaves	Saprobic	Spartina sp.	Canada: Bay of Fundy	[48]
Sporobolomyces spp.	Living/decomposing leaf blades and sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,50]
PUCCINIOMYCETES
Pucciniales
Pucciniaceae
Puccinia distichlidis Ellis and Everh.	–	–	Distichlis spicata	USA	[151]
Puccinia magnusiana Körn.	Living/decomposing leaf blades and sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,50]
Puccinia phragmitis (Schumach.) Tul.	Living/decomposing leaf blades and sheaths	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,50]
Puccinia sparganioidis Ellis and Barthol.	–	Saprobic, parasitic	Spartina alterniflora	USA: Maine, New Hampshire, Massachusetts, Rhode Island, Delaware, Virginia, North Carolina, Florida, Mississippi	[36,44,68,73,152]
	–	–	Spartina cynosuroides	USA: New Jersey, Delaware, Maryland, South Carolina, Florida, Louisiana	[44,68,153]
	–	Saprobic, pathogenic	Spartina patens	USA: Connecticut, Maryland, New Jersey, New York	[44,68,153]
Uromyces acuminatus Arthur	–	Saprobic, pathogenic	Spartina alterniflora	USA: Maine, New Hampshire, Massachusetts, Connecticut, New York, New Jersey, Delaware, Maryland, Florida	[44,68,152]
	–	Saprobic, pathogenic	Spartina cynosuroides	USA: Florida	[44,68,153]
	–	Saprobic	Spartina patens	USA: Connecticut, Delaware, Florida, Maine, Maryland, Massachusetts, New Hampshire, New Jersey,	[44,68]
Uromyces argutus F. Kern	–	Saprobic, pathogenic	Spartina alterniflora	France; USA: Florida	[44,68,152]
Uromyces salicorniae (DC.) de Bary	–	–	Salicornia sp.	South Australia	[95]
Pucciniales genera incertae sedis
Aecidium suaedae Thüm.	Leaves	–	Suaeda verae	Egypt	[154]
TREMELLOMYCETES
Tremellales
Tremellaceae
Tremella spicifera Van Ryck., Van de Put and P. Roberts	Living/decomposing leaf sheaths and stems	Saprobic	Phragmites australis	Netherlands: Zeeland	[39,40]
USTILAGINOMYCETES
Ustilaginales
Ustilaginaceae
Tranzscheliella distichlidis (McAlpine) Vánky	–	Pathogenic	Distichlis spicata	Australia: Victoria	[155]
Ustilaginales genera incertae sedis
Parvulago marina (Durieu) R. Bauer, M. Lutz, Piątek, Vánky and Oberw.	–	–	Eleocharis parvula	Finland, France, Germany, UK, Norway, Sweden	[156]
Urocystidales
Urocystidaceae
Flamingomyces ruppiae (Feldmann) R. Bauer, M. Lutz, Piątek, Vánky and Oberw.	–	Parasitic	Ruppia maritima	France	[156]
MUCOROMYCOTA
MUCOROMYCETES
Mucorales
Choanephoraceae
Blakeslea trispora Thaxt.	Senescent and decaying leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
Mucoraceae
Mucor sp.	Senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]
	Roots	Saprobic	Spartina sp.	Canada: Bay of Fundy	[48]
Rhizopodaceae
Rhizopus stolonifer (Ehrenb.) Vuill.	Stems	Saprobic	Spartina townsendii	UK: England	[49]
Syncephalastraceae
Syncephalastrum racemosum Cohn ex J. Schröt.	Living and senescent leaves	Saprobic	Juncus roemerianus	USA: Florida	[43]

2. Taxonomic Classification of Salt Marsh Fungi

2.1. Phyla

Calado and Barata [34] documented 332 taxa associated with Juncus roemerianus, Phragmites australis, and Spartina spp. In this review, we list 486 taxa that belong to three phyla (Ascomycota, Basidiomycota, Mucoromoycota) (Table 1, Figure 3) and selected species are illustrated in Figure 4. Ascomycota dominates the taxa from salt marsh ecosystems, accounting for 95.27% (463 taxa). Nineteen species in twelve genera (Aecidium, Chaetospermum, Falmingomyces, Merismodes, Nia, Parvulago, Puccinia, Sporobolomyces, Stilbum, Tranzscheliella, Tremella, Uromyces) belong to Basidiomycota (3.91%), while Mucoromycota account for 0.82% (four species) of the salt marsh fungi.

Figure 3.

The distribution of salt marsh fungi among three fungal phyla.

Figure 4.

Salt marsh fungi. (a,b) Halobyssothecium obiones from Atriplex portulacoides; (c,d) Halobyssothecium phragmites from culms of Phragmites sp.; (e,f) Buergenerula spartinae from culms of Spartina sp.; (g,h) Chaetomium sp. from stem of Typha sp.; (i,j) Alternaria sp. from culms of Spartina sp. Scale bars: (a,g) = 500 µm; (b,d,f,h,j) = 20 µm; (c,i) = 200 µm; (e) = 100 µm.

2.2. Class

Salt marsh fungi are distributed into 17 classes (Table 1, Figure 5). Dothideomycetes has the highest number of taxa, which comprises 47.12% (229 taxa), followed by Sordariomycetes with 167 taxa (34.36%). Twenty-one species (in 20 genera) can be referred to as Ascomycota genera incertae sedis. The Ascomycetes with the least number of species include Leotiomycetes (21 species, 4.32%), Eurotiomycetes (16 species, 3.29%), Orbiliomycetes (3 species, 0.62%), Saccharomycetes (3 species, 0.62%), Lecanoromycetes (2 species, 0.41%), and Pezizomycetes (1 species, 0.21%).

Figure 5.

The distribution of salt marsh fungi in different fungal classes.

Seven classes represent the Basidiomycota (Figure 5). Puccinomycetes has the highest number of taxa documented (eight species, three genera) followed by Agaricomycetes (three species, two genera), Ustilaginomycetes (three species, three genera), and Microbotryomycetes (two taxa, one genus). Agaricostilbomycetes, Bartheletiomycetes, and Tremellomycetes have one representative taxon each.

The Mucoromoycota account for the taxa Blakeslea trispora, Mucor sp., Rhizopus stolonifera, and Syncephalastrum racemosum [43,48,49].

2.3. Orders

Salt marsh fungi recorded from different halophytes were distributed among 48 orders (Table 1, Figure 6). The Pleosporales is the largest order, with 178 taxa recorded followed by Hypocreales (41), Microascales (26), Capnodiales (22), Helotiales (18), Xylariales (17), Sordariales (16), Amphisphaeriales (15), and Eurotiales (13). The remaining 41 orders have less than 10 species (Table 1, Figure 5). Forty-two taxa belong to incertae sedis (Ascomycota genera incertae sedis: 21; Dothideomycetes families incertae sedis: 11; Sordariomycetes families incertae sedis: 9; Xylariomycetidae family incertae sedis: 1).

Figure 6.

The distribution of salt marsh fungi in major fungal orders.

2.4. Families

A total of 108 families and 12 incertae sedis were recorded to be associated with salt marsh fungi (Table 1, Figure 7). Phaeosphaeriaceae and Pleosporaceae account for the largest families with 34 and 31 taxa recorded, respectively. Thirteen families have ten or more than taxa and include Nectriaceae (25), Halosphaeriaceae (25), Didymellaceae (17), Mycosphaerellaceae (14), Lentitheciaceae (13), Massarinaceae (13), Chaetomiaceae (12), Xylariaceae (11), Didymosphaeriaceae (10), Leptosphaeriaceae (10), and Aspergillaceae (10). The remaining 95 families have less than ten species recorded. Forty-four taxa are placed as incertae sedis, wherein 21 of these belong to Ascomycota genera incertae sedis.

Figure 7.

The distribution of salt marsh fungi among major fungal families.

3. Diversity of Fungi in Halophytes

Twenty-seven genera under 11 families (Amaranthaceae, Apiaceae, Caryophyllaceae, Compositae, Juncaceae, Juncaginaceae, Plumbaginaceae, Poaceae, Poaceae, Primulaceae, Ruppiaceae, Typhaceae, Zosteraceae) of halophytes were reviewed for its fungal associates (Table 1, Figure 8). Halophytic species are represented in Figure 1 and Figure 2.

Figure 8.

The number of taxa observed from different hosts in salt marsh ecosystems.

3.1. Amaranthaceae

Six genera (Arthrocnemum, Atriplex, Salicornia, Salsola, Sarcocornia, Suaeda) represent the Amaranthaceae. Suaeda and Salicornia are the most studied hosts in Amaranthaceae. Ascomycota account for 96.30% of the 52 taxa recorded in Amaranthaceae (Figure 9, Table 1). Two Pucciniomycetes species, Aecidium suaedae [154] and Uromyces salicorniae [95], represent Basidiomycota. The taxa in Amaranthaceae represent three classes wherein Dothideomycetes accounts for 85.19% (46 taxa), followed by Sordariomycetes with six taxa reported.

Figure 9.

The number of taxa observed from Amaranthaceae.

Fungi associated with Suaeda total 18 taxa. Dothideomycetes was represented by 14 taxa (77.78%), while three taxa were Sordariomycetes (Cryptovalsa suaedicola [144], Fusarium fujikuroi [62], Moheitospora fruticosae [130]) and one taxon of Pucciniomycetes (Aecidium suaedae [154]).

A total of 14 taxa were documented in Salicornia. Eleven of these belong to Dothideomycetes (Pleosporales: 10; Capnodiales: 1), followed by Sordariomycetes (two taxa: Halocryptovalsa salicorniae [145], Tubercularia pulverulenta [35]), and Pucciniomycetes (one taxon: Uromyces salicorniae [95]).

Fungi from Atriplex total 11 taxa (10 genera) and all of these belong to Pleosporales (Dothideomycetes). Sarcocornia harbors seven taxa (six Dothideomycetes, one Sordariomycetes). Only two taxa (Alternaria spp., Stemphylium spp.) and a single taxon (Mycosphaerella salicorniae) were reported from Salsola [35] and Arthrocnemum [35], respectively.

3.2. Poaceae

The association of fungi with grasses have been documented and most of the host plants are members of Poaceae. Ten genera of salt marsh grasses under Poaceae are included in this review wherein Spartina is the most studied of halophytic hosts for direct observation of marine fungi. In addition to Spartina, salt marsh grasses such as Phragmites and Distichlis were well studied also for their fungal associates.

Salt marsh fungi are not well-documented from grasses such as Spartina anglica, S. pectinata, Spergularia marina, Uniola paniculata, Elymus farctus, × Ammocalamagrostis baltica, and Agropyron sp. with one taxon recorded for each host [35]. Furthermore, there are few studies on the fungal composition of Arundo donax (4 taxa) [35] and Ammophila arenaria (four taxa). Marram grass (Ammophila arenaria) is more common in sand dunes and supports quite a diverse fungal community [157,158], while arbuscular mycorrhizal fungi (AMF) play a key role in the establishment, growth, and survival of plants [159].

3.2.1. Distichlis spicata

Ascomycota dominates the taxa associated with Distichlis spicata (93.55%) wherein 16 and 13 species are members of Dothideomycetes and Sordariomycetes, respectively. Pleosporalean taxa constitute the majority of fungi associated with D. spicata (14 species), followed by Hypocreales with nine species recorded. Puccinia distichlidis and Tranzscheliella distichlidis represent the Basidiomycota. A total of 26 genera were recorded as associates of D. spicata and were mostly observed on senescent and decaying leaves.

3.2.2. Elymus pungens

Sixty-seven taxa were recorded in Elymus pungens and belong to Ascomycota. Most of the taxa belong to Dothideomycetes (32 taxa), followed by Sordariomycetes (21 taxa), Leotiomycetes, and Eurotiomycetes (6 taxa) (Table 1, Figure 10).

Figure 10.

The distribution of fungal taxa associated with Elymus pungens.

3.2.3. Puccinellia maritima

A total of 12 taxa (six Sordariomycetes; the following five Dothideomycetes: Micronectriella agropyri, Lautitia danica, Leptosphaeria pelagica, Septoriella vagans, Paradendryphiella salina; one Leotiomycetes: Thelebolus crustaceus) were recorded in Puccinellia maritima [38]. All the taxa from Sordariomycetes belong to Sordariales (Chaetomium elatum, C. globosum, C. thermophilum, Corynascus sepedonium, Thermothielavioides terrestris, Sordaria fimicola) [38].

3.2.4. Spartina

A total of 149 taxa (141 Ascomycota, 6 Basidiomycota, 2 Mucoromycota) were recorded in Spartina. The majority of the taxa belong to Dothideomycetes (70 taxa), followed by Sordariomycetes (59 taxa). Pleosporaceae and Halosphaeriaceae dominate the fungi documented in Spartina with 19 and 17 taxa recorded, respectively. Spartina alterniflora, S. maritima, and Spartina × townsendii harbor 79, 46, and 49 taxa, respectively (Figure 11, Table 1). A total of 78 taxa were recorded in the unidentified Spartina species. The identification of the Spartina species can be challenging, wherein species are morphologically similar.

Figure 11.

The distribution of fungal taxa associated with Spartina.

Halobyssothecium obiones was recorded from six species of Spartina (S. alterniflora [20,35,52,61,71,74,80,81,82], S. cynosuroides [35], S. densiflora [64], S. maritima [31,54,59,63], S. patens [36], S. townsendii [49,65], and the unidentified Spartina sp. [32,35,36,58,84]), while six Spartina spp. harbors unidentified Mycosphaerella species. Six species (Leptosphaeria pelagica, Lulworthia spp., Phaeosphaeria halima, Phaeosphaeria spartinicola, Phoma spp., Stagonospora spp.) were recorded in five different hosts. The unidentified Spartina species harbors 28 unique species. Amongst the taxa found in Spartina, 32 species can only be found in S. alterniflora, while S. maritima harbors 21 unique species, the most intensively surveyed species.

3.2.5. Phragmites

A total of 138 taxa have been documented in Phragmites (Figure 12, Table 1). Most of the taxa belong to Ascomycota (131 taxa), while six taxa represent the Basidiomycota. Dothideomycetes dominates half of the taxa in Phragmites (71 taxa, 51.45%) followed by Sordariomycetes (44 taxa, 31.88%), Leotiomycetes (6 taxa, 4.35%), Ascomycota genera incertae sedis (5 taxa, 3.62%), Eurotiomycetes (3 taxa, 2.17%), Orbiliomycetes (2 taxon, 1.45%), and Pucciniomycetes (1 taxa, 1.45%). One taxon each were recorded to Agaricomycetes [40], Bartheletiomycetes [41], Lecanoromycetes [39], Microbotryomycetes [39,50], and Tremellomycetes [39,40]. Pleosporalean taxa accounts for the highest number of fungi associated with Phragmites (42.75%, 59 taxa).

Figure 12.

The distribution of fungal taxa associated with Phragmites.

Phragmites australis harbors diverse fungi that totals to 137 taxa (101 genera) [39,40,41,50,79,115]. Seven species (Arthrinium arundinis [62], Halazoon fuscus [87], Halobyssothecium phragmitis [85], Keissleriella linearis [85], Phomatospora dinemasporium [62], Remispora hamata [87], Setoseptoria phragmitis [87]) were recorded in unidentified Phragmites species.

3.3. Juncaceae

Juncus roemerianus, J. maritimus, and an unidentified Juncus species represent Juncaceae. Salt marsh fungi are diverse in Juncus and dominated by Ascomycota, which constitutes 97.58% of the 165 reported taxa (Figure 13, Table 1). Stilbum sp. represented the Basidiomycota, while three taxa (Blakeslea trispora, Mucor sp., Syncephalastrum racemosum) of Mucoromycota were recorded. Dothideomycetes and Sordariomycetes account for the highest number of Juncus-associated fungi with 72 (43.64%) and 64 (38.79%) taxa documented.

Figure 13.

The distribution of fungal taxa associated with Juncus.

Juncus roemerianus has been extensively studied for its associates with 162 documented taxa [32,42,43,60,66,76,77,78,97,98,104,105,110,116,117,118,135,147,148]. Few species were reported to Juncus maritimus that harbor only two taxa (Leptosphaeria albopunctata, Phaeosphaeria neomaritima) [35]. Phaeosphaeria neomaritima [36,52,71,80], P. spartinicola [52], and Monodictys pelagica [35] were observed in an unidentified species of Juncus.

Phragmites australis harbors diverse fungi that totals to 137 taxa (101 genera) [39,40,41,50,79,115]. Seven species (Arthrinium arundinis [62], Halazoon fuscus [87], Halobyssothecium phragmitis [85], Keissleriella linearis [85], Phomatospora dinemasporium [62], Remispora hamata [87], Setoseptoria phragmitis [87]) were recorded in unidentified Phragmites species.

3.4. Other Families

Few reports on salt marsh fungi are from the following hosts: Apiaceae: Crithmum maritimum (one taxon: Phoma sp.), Typhaceae: Typha spp. (five taxa: Arundellina typhae, Chaetomium sp., Magnisphaera spartinae, Pleospora pelagica, Remispora hamata); Compositae: Artemisia maritima (two taxon: Neocamarosporium artemisiae, N. maritimae); Caryophyllaceae: Spergularia marina (one taxon: Cladosporium algarum); Plumbaginaceae: Limonium sp. (one taxon: Mycosphaerella salicorniae); Armeria pungens (one taxon: Mycosphaerella staticicola); Juncaginaceae: Triglochin sp. and T. maritima (one taxon: Stemphylium triglochinicola); Primulaceae: Lysimachia maritima (two taxa: Leptosphaeria orae-maris, Stemphylium vesicarium); Ruppiaceae: Ruppia maritima (one taxon: Flamingomyces ruppiae); and Zosteraceae: Zostera marina (one taxon: Corollospora ramulosa) and Zostera sp. (Asteromyces cruciatus). Alva et al. [160] report Penicillium chrysogenum as an endophyte from Zostera japonica.

Fourteen taxa were documented from unidentified salt marsh plants. All of the taxa belong to Ascomycota (seven Dothideomycetes, five Sordariomycetes, one Eurotiomycetes). Pleosporalean taxa from six families account for half of the taxa (the following seven species: Camarosporium palliatum, C. roumeguerei, Coniothyrium obiones, Halobyssothecium obiones, Periconia sp., Loratospora aestuarii, Pleospora pelvetiae).

4. Geographical Distribution of Salt Marsh Fungi

The salt marsh fungi reported are from countries of three major oceans, as documented in Figure 14. The Atlantic Ocean consists of 12 countries, wherein the USA had the highest number of species recorded (232 taxa) followed by the UK (101 taxa), the Netherlands (74 taxa), and Argentina (51 taxa). China had the highest number of salt marsh fungi in the Pacific Ocean with 165 taxa reported, while in the Indian Ocean, India reported the highest taxa (16 taxa). Most of the biodiversity studies documenting salt marsh fungi in the Atlantic Ocean are mostly from the USA and the UK and this reflects the high number of taxa [32,36,38,49,61]. China ranked second with the most number of salt marsh fungal taxa, mainly due to the biodiversity study in Phragmites australis conducted by Poon et al. [41].

Figure 14.

The number of salt marsh fungi reported in the Pacific, Atlantic, and Indian Oceans.

The geographical distribution of salt marsh fungi and the different halophytes are presented in Figure 15. The fungi associated with salt marsh grass Phragmites australis have been studied in different countries (Australia, Belgium, Egypt, France, Germany, China, Iraq, Japan, the Netherlands, South Australia, Thailand). Spartina alterniflora was recorded in countries along the Atlantic (Argentina, Canada, France, USA) and the Indian Ocean (India), but lacks data from countries in the Pacific Ocean.

Figure 15.

Map of countries showing the global distribution of fungal diversity studies in halophytes. The different color of each pie chart represents the hosts, and the angle measured the number of their fungal associates.

United States of America

Most of the studies of halophytes-associated fungi were concentrated on the United States of America (USA) (Figure 16). Table 1 lists the salt marsh fungi in 20 states. Florida has been the frequently studied, wherein seven hosts (Juncus roemerianus: 108 taxa; Spartina × townsendii: 1; Spartina alterniflora: 16; Spartina cynosuroides: 3; Spartina densiflora: 1; Spartina patens: 2; Spartina spp.: 3) were observed for salt marsh fungi. Six hosts were studied in North Carolina, wherein Juncus roemerianus harbored the highest number of fungi (48 taxa). In Rhode Island, Spartina alterniflora accounts for the highest number of fungi, with 41 taxa recorded.

Figure 16.

Map of the United States of America (USA) showing the distribution of fungal diversity studies of halophytes in different states. The different color of each pie chart represents the hosts, and the angle measured the number of their fungal associates.

5. Conclusions and Future Perspectives

Most studies of fungi on salt marsh plants are from Spartina, Juncus, and Phragmites, probably due to the huge biomass generated by these taxa. The mycota of less bulky halophytes (e.g., Limonium, Triglochin, Uniola) and litter from the surrounding sea grass beds washed off to marsh areas (e.g., Zostera japonica, Z. marina, Z. noltii) are also less represented, or these hosts are yet to be explored. The checklist presented in the current study updates the list of Calado and Barata [34] and the inclusion of fungi associated with rarely studied halophytes record 486 taxa worldwide. Ascomycota dominate the taxa (463 taxa) and are comprised mostly of Dothideomycetes with their ability to eject their ascospores forcibly and widely, spore type, the formation of ascomata or ascostromata under a clypeus or just immersed in thin leaves, and an ability to decompose lignocellulose substrates [57,161]. Meyers et al. [162] showed that salt marsh yeasts and the ascomycete, Buergenerula spartinae, produce degradative enzymes and utilize simple carbon and nitrogen compounds. The yeast, Pichia spartinae, produces β-glucosidase and other degradative enzymes. Gessner [74] demonstrated that a number of salt marsh fungi isolated from Spartina alterniflora, Zostera sp., and Z. marina produced enzymes capable of degrading cellulose, cellobiose, lipids, pectin, starch, tannic acid, and xylan and, thus, play a key role in the degradation of storage and structural compounds. Salt marsh fungi might possess high biotransformation and metabolic abilities, which could be related to their ecology.

Basidiomycota (19 taxa) and Mucoromycota (4 taxa) are poorly represented in salt marsh ecosystems as they are in other marine habitats [163]. There are no records of Chytridiomycota listed in the present work and only a few authors detected this group, and other basal fungal lineages, in salt marsh ecosystems using molecular analysis [164,165,166,167]. These groups are worth exploring to determine the overall fungal communities in the salt marsh ecosystems. Many chytrids and other basal fungi are more challenging to cultivate and require different isolation methods (e.g., baiting techniques in liquid culture) than the saprobes, methods that have rarely been applied in the study of saltmarsh plants. When appropriate techniques are used, chytrids and other zoosporic organisms have been reported. For example, the fungal-like organism Phytophthora inundata has been recovered from the halophilic plants Aster tripolium and Salicornia europaea, while P. gemini and P. chesapeakensis occur on Zostera marina, and Salisapilia nakagirii on the decaying litter of Spartina alterniflora (www.marinefungi.org; accessed on 10 May 2021, [163]). Marine chytrids have been isolated from substrates such as seaweeds and mangrove leaves [163].

The taxa listed are mostly saprobes and these can be attributed to the inclusion of salt marsh fungi observed directly from the different host parts, which are mostly submerged decaying substrates. When compared to saprobic fungi in halophytes, few studies have been carried out on the diversity of endophytes and pathogens and their interaction in the salt marsh ecosystems. Surveys on endophytic fungi from halophytes using cultivation-dependent methods coupled with molecular approaches, showed that endophytes were dominated by Ascomycota and a few belonged to Basidiomycota and Zygomycota [168,169,170,171,172,173,174,175]. Pathogenic fungi from salt marsh ecosystems are poorly documented but play a significant role in the dynamics of the ecosystem [176,177,178]. For example, Govers et al. [179] reported that the fungal-like organisms Phytophthora gemini and P. inundata caused widespread infection of the common seagrass species, Zostera marina (eelgrass), across the northern Atlantic and Mediterranean that threatened the conservation and restoration of vegetated marine coastal systems. Likewise, Claviceps purpurea affects the viability of Spartina townsedii in south coast UK salt marshes. Fisher et al. [180] noted that Cl. purpea in the Alabama and Mississippi coastlines rendered the seeds of one of the primary salt marsh grasses sterile. Raybold et al. [181] recorded epidemics of C. purpurea on Spartina anglica in Poole Harbor (UK) and that ergot growth was detrimental to seed production. These underexplored fungal groups are worthy to be explored for their ecological and biotechnological importance.

This shows how salt marsh fungal studies were concentrated in countries in the Atlantic Ocean specifically the USA (232 taxa) and the UK (101 taxa). Many salt marsh areas remain unexplored, especially those in the Indian and Pacific Oceans, and these areas are hotspots of biodiversity and novel fungal taxa based on the exploration of various habitats [85,100,163,182,183,184,185,186,187]. Recently, novel species were isolated in halophytes [85,100,145] and further taxa remain to be discovered, isolated, and sequenced, while vast areas worldwide have yet to be surveyed. For example, salt marsh plants are immensely numerous, diverse, and common along the south-east coast of Australia, yet little is known of their fungal associates [188].

The salt marsh vegetation and its fungal associates are adapted to salt stress and inundation and are subjected to extreme environmental conditions such as being periodically wet to different lengths of time leading to drying out at low tides and exposure to high temperatures and drying out at midday. Many are well adapted to prevailing conditions by their fleshy leaves (Suaeda australis), others can tolerate high flooding.

Few data are currently available on the specificity of fungi on their salt marsh hosts. Figure 17 shows the number of fungal taxa recorded from the three commonly studied hosts, Juncus, Phragmites, and Spartina, wherein there is little overlap in the species composition. One of the common species on Spartina plants is undoubtedly Halobyssothecium obiones, while Leptosphaeria pelagica is common. A common ascomycete on Atriplex portulacoides and Suaeda maritima is Decorospora gaudefroyi. Host plants that have been little surveyed for fungi are Limonium vulgare (sea lavender) and Atriplex portulacoides (sea purslane), yet they do support a number of taxa, e.g., Neocamarosporium obiones and Amarenomyces ammophilae. The fungal community reported on Juncus roemerianus in the salt marsh at North Carolina is significantly different from those on Spartina and Phragmites. It remains to be seen if this is due to the host plant or its geographical location.

Figure 17.

Venn diagram showing the association of salt marsh fungi from commonly studied halophytes.

Another groups of fungi that have not been fully studied in the salt marsh habitat are yeasts, as these also require specific techniques for their isolation from the water column or from plant tissue. Spencer et al. [189] recovered a number of yeasts from the vicinity of Spartina townsendii, as follows: very numerous Cryptococcus spp.; Trichosporon cutaneum; Trichosporon pullulans; the relatively rare species, Metschnikowia bicuspidata and Cryptococcus flavus; and Saturnospora ahearnii [190]. Although marine yeasts are common in sea water and deep seawater vents [163], their large-scale sampling in salt marshes remains a challenge for the future.

Currently, the salt marsh ecosystem has been threatened both by global warming and human activity. Sea-level rises brought about by climate change alter the location and character of the land–sea interface wherein salt marsh vegetation moves upward and inland. The increase in the sea level may not lead to the loss of coastal marshes, but the resiliency will depend on the ability of halophytes to migrate upland. Susceptible areas are organogenic marshes and areas where sediment is limited, potentially leading to catastrophic shifts and marsh loss. In this paper, a total of 57 plant taxa under 27 genera were reviewed for their fungal associates. The halophytes included here are only approximately 11% of the total number of species of salt marsh plants worldwide. Thus, many salt marsh fungi await discovery with wider host plant sampling and the use of a wider range techniques for their isolation. For this reason, it is imperative to study the halophytic fungi to document not just biodiversity but also to discover novel taxa restricted only to this kind of habitat.

Author Contributions

Conceptualization: M.S.C., K.D.H. and E.B.G.J.; methodology: M.S.C., K.D.H. and E.B.G.J.; formal analysis and investigation: M.S.C., K.D.H. and E.B.G.J.; resources: K.D.H. and E.B.G.J.; writing—original draft preparation, M.S.C.; writing—review and editing, K.D.H., E.B.G.J. and I.P.; supervision, K.D.H. and E.B.G.J.; funding acquisition, K.D.H. and E.B.G.J. All authors have read and agreed to the published version of the manuscript.

Funding

K.D.H. thanks the Thailand Research Fund for the grant entitled “Impact of climate change on fungal diversity and biogeography in the Greater Mekong Subregion” (Grant No. RDG6130001). E.B.G.J. is supported under the Distinguished Scientist Fellowship Program (DSFP), King Saud University, Kingdom of Saudi Arabia.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

Not applicable.

Conflicts of Interest

The authors declare no conflict of interest.