Cladosporium

Link, Mag. Gesell. naturf. Freunde, Berlin 7: 37. 1816 (1815).

For synonyms see Bensch et al. (2012).
Classification: Dothideomycetes, Dothideomycetidae, Capnodiales, Cladosporiaceae.
Type species: Cladosporium herbarum (Pers. : Fr.) Link. Lectotype: L 910.225-733. Epitype and ex-epitype culture: CBS H-19853, CPC 12100 = CBS 121621.
DNA barcodes (genus): LSU.
DNA barcodes (species): act and tef1; in a few cases tub2.

Ascomata pseudothecial, black to red-brown, globose, inconspicuous and immersed beneath stomata to superficial, situated on a reduced stroma, with 1(–3) short, periphysate ostiolar necks; periphysoids frequently growing down into cavity; ascomatal wall consisting of 3–6 layers of textura angularis. Pseudoparaphyses frequently present in mature ascomata, hyaline, septate, subcylindrical. Asci fasciculate, short-stalked or not, subsessile, bitunicate, obovoid to broad ellipsoid or subcylindrical, straight to slightly curved, 8-spored. Ascospores bi- to multiseriate, hyaline, obovoid to ellipsoid-fusiform, with irregular luminar inclusions, mostly thick-walled, straight to slightly curved, frequently becoming brown and verruculose in asci, at times covered in mucoid sheath. Dematiaceous hyphomycetes. In vivo: Mycelium internal or external, superficial; hyphae branched, septate, subhyaline to usually pigmented, smooth, sometimes slightly rough-walled to verruculose. Stromata absent to sometimes well-developed. Conidiophores mononematous, usually macronematous, solitary, fasciculate, in small to large fascicles, loosely to densely caespitose, usually erect, occasionally subdecumbent, decumbent or repent, straight to flexuous, unbranched or branched, continuous to septate, subhyaline to usually distinctly pigmented, smooth to verruculose, proliferation holoblastic, occasionally enteroblastic (after a period when growth has stopped and then resumed), usually sympodial, rarely monopodial (sometimes leaving coarse annellations from repeated enteroblastic proliferation). Conidiogenous cells integrated, terminal or intercalary, monoblastic or usually polyblastic, mostly sympodially proliferating, more or less cylindrical, geniculate-sinuous or nodulose, sometimes with unilateral swellings; conidiogenous loci usually conspicuous, protuberant, composed of a central convex dome surrounded by a more or less raised periclinal rim (coronate), thickened, refractive or barely to distinctly darkened; conidial formation holoblastic. Conidia solitary or catenate, in unbranched or branched acropetal chains, amero- to phragmosporous, shape and septation variable, usually subglobose, ovoid, obovoid, ellipsoid, fusiform, limoniform to cylindrical, aseptate or with several transverse eusepta, rarely with a single longitudinal septum, subhyaline to usually pigmented, smooth, verruculose, verrucose, echinulate, cristate; hila protuberant, coronate, with a central convex dome and raised periclinal rim, thickened, refractive to darkened; microcyclic conidiogenesis often occurring. In vitro: Stromata usually lacking. Conidiophores usually solitary, arising terminally or laterally from plagiotropous or ascending hyphae, often longer than in vivo. Micronematous conidiophores, lacking in vivo, are often formed in culture. Conidial chains often longer than in vivo (species with solitary conidia are often capable of forming conidial chains in culture).

Culture characteristics:

Colonies on SNA often grey olivaceous or olivaceous grey, reverse leaden grey or black, flat, velvety with fluffy or cottony patches, margin irregular or undulate, aerial mycelium loose diffuse or more abundantly formed, often with abundant submerged mycelium.

Optimal media and cultivation conditions: For morphological examinations SNA incubated under continuous near-ultraviolet light at 25 °C proved to be best suited to promote sporulation. The sexual morph can be induced by inoculating plates of 2 % WA onto which autoclaved stem pieces of Urtica dioica (European stinging nettle) are placed. Inoculated plates have to be incubated on the laboratory bench for 1 wk, after that period they have to be further incubated at 10 °C in the dark for 1–2 mo to stimulate sexual morph development.

Distribution:

Worldwide.

Hosts:

Asparagaceae, Asteraceae, Fabaceae, Myrtaceae, Orchidaceae, Poaceae and many other hosts, including fungi and insects.

Disease symptoms:

Leaf spots, leaf blight, discolorations, necrosis, or shot-hole symptoms, on stems and fruits, but also saprobic, endophytic or isolated from numerous substrates and environments, e.g. indoor environments, salterns and human and animal infections.

Notes:

The monophyletic genus Cladosporium is well characterised by the coronate structure of its conidiogenous loci and conidial hila, consisting of a central convex dome surrounded by a raised periclinal rim (David 1997, Braun et al. 2003). At the moment it comprises about 200 species. Cladosporium was previously extremely heterogeneous and encompassed 772 names assigned to this genus (Dugan et al. 2004). Heuchert et al. (2005) examined Cladosporium spp. dwelling on other fungi, and Schubert (2005) provided a comprehensive treatment of foliicolous species. Crous et al. (2007a) encompassed a series of papers dealing with a reassessment and new circumscription of Cladosporium s. str. and treatments of several cladosporioid genera. Bensch et al. (2012) published a taxonomic monograph of the genus Cladosporium which can be consulted for further information on the history and many other aspects of this genus.

Species delimitation in Cladosporium based on morphology alone is limited since many species have overlapping characters. Some key differential features have been identified and detailed in a series of monographic papers (Schubert et al. 2007, Zalar et al. 2007, Bensch et al. 2010, 2012). The most relevant differential morphological traits are the shape, width and complexity of conidiophores, the presence of ramoconidia, and the formation and ornamentation of conidia. However, given the overlapping of these features, and the need for standardisation using special culture media and scanning electron microscopy procedures, the use of a molecular approach should be mandatory for correct identification of the species in this complex fungal group (Sandoval-Denis et al. 2016).

Three different species complexes are recognised within the genus, mainly based on morphology, and used for practical purposes, the Cl. cladosporioides species complex characterised by mainly narrowly cylindrical or cylindrical-oblong, non-nodulose, mostly non-geniculate conidiophores and conidia with a quite variable surface ornamentation ranging from smooth to irregularly verrucose-rugose or rough-walled (reticulate or embossed stripes under SEM); the Cl. herbarum species complex with species mainly having nodulose conidiophores with conidiogenesis confined to swellings and verruculose, verrucose or echinulate conidia; and the Cl. sphaerospermum complex with its most remarkable feature of forming numerous globose or subglobose terminal and intercalary conidia in most of the species with a variable surface ornamentation and often poorly differentiated conidiophores (Bensch et al. 2012, 2015). Morphologically similar genera have been treated in Crous et al. (2007b).

Members of Cladosporiaceae: Cladosporium, Graphiopsis, Neocladosporium, Rachicladosporium, Toxicocladosporium, Verrucocladosporium.

References:

Braun et al. 2003 (sexual morph); Crous et al. 2007a, b (cladosporium-like genera); Schubert et al. 2007 (morphology, phylogeny Cl. herbarum complex); Zalar et al. 2007 (morphology, phylogeny Cl. sphaerospermum complex); Bensch et al. 2010 (morphology, phylogeny Cl. cladosporioides complex); Bensch et al. 2012 (morphology, phylogeny and key of all Cladosporium species); Bensch et al. 2015 (morphology, additions to the three species complexes); Sandoval-Denis et al. 2016 (morphology, phylogeny of clinical samples).
Bensch K, Braun U, Groenewald JZ, et al. (2012). The genus Cladosporium. Studies in Mycology 72: 1–401.
Bensch K, Groenewald JZ, Braun U, et al. (2015). Common but different: The expanding realm of Cladosporium. Studies in Mycology 82: 23–74.
Bensch K, Groenewald JZ, Dijksterhuis J, et al. (2010). Species and ecological diversity within the Cladosporium cladosporioides complex (Davidiellaceae, Capnodiales). Studies in Mycology 67: 1–94.
Braun U, Crous PW, Dugan FM, et al. (2003). Phylogeny and taxonomy of cladosporium-like hyphomycetes, including Davidiella gen. nov., the teleomorph of Cladosporium s. str. Mycological Progress 2: 3–18.
Crous PW, Braun U, Schubert K, et al. (eds) (2007a). The genus Cladosporium and similar dematiaceous hyphomycetes. Studies in Mycology 58: 1–253.
Crous PW, Braun U, Schubert K, et al. (2007b). Delimiting Cladosporium from morphologically similar genera. Studies in Mycology 58: 33–56.
David JC (1997). A contribution to the systematics of Cladosporium. Revision of the fungi previously referred to Heterosporium. Mycological papers 172: 1–157.
Dugan FM, Schubert K, Braun U (2004). Check-list of Cladosporium names. Schlechtendalia 11: 1–103.
Heuchert B, Braun U, Schubert K (2005). Morphotaxonomic revision of fungicolous Cladosporium species (hyphomycetes). Schlechtendalia 13: 1–78.
Sandoval-Denis M, Gené J, Sutton DA, et al. (2016). New species of Cladosporium associated with human and animal infections. Persoonia 36: 281–298.
Schubert K (2005). Morphotaxonomic revision of foliicolous Cladosporium species (hyphomycetes). Ph.D. dissertation. Mathematisch-Naturwissenschaftlich-Technischen Fakultät, Martin-Luther-University Halle-Wittenberg, Germany.
Schubert K, Groenewald JZ, Braun U, et al. (2007). Biodiversity in the Cladosporium herbarum complex (Davidiellaceae, Capnodiales), with standardisation of methods for Cladosporium taxonomy and diagnostics. Studies in Mycology 58: 105–156.
Zalar P, de Hoog GS, Schroers H-J, et al. (2007). Phylogeny and ecology of the ubiquitous saprobe Cladosporium sphaerospermum, with descriptions of seven new species from hypersaline environments. Studies in Mycology 58: 157–183.

Table 5. DNA barcodes of accepted Cladosporium spp.

Species	Isolates¹		GenBank accession numbers²			References
		ITS		*act*	*tef1*
Cl. acalyphae	CBS 125982^T	HM147994		HM148481	HM148235	Bensch et al. (2010)
Cl. aciculare	CBS 140488^T	KT600411		KT600607	KT600509	Bensch et al. (2015)
Cl. aggregatocicatricatum	CBS 140493^T	KT600448		KT600645	KT600547	Bensch et al. (2015)
Cl. alboflavescens	CBS 140690^T	LN834420		LN834604	LN834516	Sandoval-Denis et al. (2016)
Cl. allicinum	CBS 121624^NT	EF679350		EF679502	EF679425	Schubert et al. (2007)
Cl. allii	CBS 101.81^RS	JN906977		JN906996	JN906983	Bensch et al. (2012)
Cl. angulosum	CBS 140692^T	LN834425		LN834609	LN834521	Sandoval-Denis et al. (2016)
Cl. angustiherbarum	CBS 140479^T	KT600378		KT600574	KT600475	Bensch et al. (2015)
Cl. angustisporum	CBS 125983^T	HM147995		HM148482	HM148236	Bensch et al. (2010)
Cl. angustiterminale	CBS 140480^T	KT600379		KT600575	KT600476	Bensch et al. (2015)
Cl. antarcticum	CBS 690.92^T	EF679334		EF679484	EF679405	Schubert et al. (2007)
Cl. antropophilum	CBS 140685^T	LN834437		LN834621	LN834533	Sandoval-Denis et al. (2016)
Cl. aphidis	CBS 132182^ET	JN906978		JN906998	JN906985	Bensch et al. (2012)
Cl. arthropodii	CBS 124043^ET	JN906979		JN906998	JN906985	Bensch et al. (2012)
Cl. asperulatum	CBS 126340^T	HM147998		HM148485	HM148239	Bensch et al. (2010)
Cl. australiense	CBS 125984^T	HM147999		HM148486	HM148240	Bensch et al. (2010)
Cl. austroafricanum	CBS 140481^T	KT600381		KT600577	KT600478	Bensch et al. (2015)
Cl. austrohemisphaericum	CBS 140482^T	KT600382		KT600578	KT600479	Bensch et al. (2015)
Cl. basiinflatum	CBS 822.84^T	HM148000		HM148487	HM148241	Bensch et al. (2010)
Cl. chalastosporoides	CBS 125985^T	HM148001		HM148488	HM148242	Bensch et al. (2010)
Cl. chasmanthicola	CBS 142612^T	KY646221		KY646224	KY646227	Marin-Felix et al. (2017)
Cl. chubutense	CBS 124457^T	FJ936158		FJ936165	FJ936161	Schubert et al. (2009)
Cl. cladosporioides	CBS 112388^NT	HM148003		HM148490	HM148244	Bensch et al. (2010)
Cl. colocasiae	CBS 386.64^T	HM148067		HM148555	HM148310	Bensch et al. (2010)
Cl. colombiae	CBS 274.80B^T	FJ936159		FJ936166	FJ936163	Schubert et al. (2009)
Cl. crousii	CBS 140686^T	LN834431		LN834615	LN834527	Sandoval-Denis et al. (2016)
Cl. cucumerinum	CBS 171.52^ET	HM148072		HM148561	HM148316	Bensch et al. (2010)
Cl. cycadicola	CPC 17251^T	KJ869122		KJ869227	KJ869236	Crous et al. (2014)
Cl. delicatulum	CBS 126344^RS	HM148081		HM148570	HM148325	Bensch et al. (2010)
Cl. dominicanum	CBS 119415^T	DQ780353		EF101368	JN906986	Zalar et al. (2007), Bensch et al. (2012)
Cl. echinulatum	CBS 123191^RS	JN906980		JN906999	JN906987	Bensch et al. (2012)
Cl. exasperatum	CBS 125986^T	HM148090		HM148579	HM148334	Bensch et al. (2010)
Cl. exile	CBS 125987^T	HM148091		HM148580	HM148335	Bensch et al. (2010)
Cl. flabelliforme	CBS 126345^T	HM148092		HM148581	HM148336	Bensch et al. (2010)
Cl. flavovirens	CBS 140462^T	LN834440		LN834624	LN834536	Sandoval-Denis et al. (2016)
Cl. floccosum	CBS 140463^T	LN834416		LN834600	LN834512	Sandoval-Denis et al. (2016)
Cl. funiculosum	CBS 122129^T	HM148094		HM148583	HM148338	Bensch et al. (2010)
Cl. fusiforme	CBS 119414^T	DQ780388		EF101372	JN906988	Zalar et al. (2007), Bensch et al. (2012)
Cl. gamsianum	CBS 125989^T	HM148095		HM148584	HM148339	Bensch et al. (2010)
Cl. globisporum	CBS 812.96^T	HM148096		HM148585	HM148340	Bensch et al. (2010)
Cl. grevilleae	CBS 114271^T	JF770450		JF770473	JF770472	Crous et al. (2011)
Cl. halotolerans	CBS 119416^T	DQ780364		EF101397	JN906989	Zalar et al. (2007), Bensch et al. (2012)
Cl. herbaroides	CBS 121626^T	EF679357		EF679509	EF679432	Schubert et al. (2007)
Cl. herbarum	CBS 121621^ET	EF679363		EF679516	EF679440	Schubert et al. (2007)
Cl. hillianum	CBS 125988^T	HM148097		HM148586	HM148341	Bensch et al. (2010)
Cl. inversicolor	CBS 401.80^T	HM148101		HM148590	HM148345	Bensch et al. (2010)
Cl. ipereniae	CBS 140483^T	KT600394		KT600589	KT600491	Bensch et al. (2015)
Cl. iranicum	CBS 126346^T	HM148110		HM148599	HM148354	Bensch et al. (2010)
Cl. iridis	CBS 138.40^ET	EF679370		EF679523	EF679447	Schubert et al. (2007)
Cl. kenpeggii	CBS 142613^T	KY646222		KY646225	KY646228	Marin-Felix et al. (2017)
Cl. langeronii	CBS 189.54^NT	DQ780379		EF101357	JN906990	Zalar et al. (2007), Bensch et al. (2012)
Cl. licheniphilum	CBS 125990^ET	HM148111		HM148600	HM148355	Bensch et al. (2010)
Cl. limoniforme	CBS 140484^T	KT600397		KT600592	KT600494	Bensch et al. (2015)
Cl. longicatenatum	CBS 140485^T	KT600403		KT600598	KT600500	Bensch et al. (2015)
Cl. longissimum	CBS 140485^T	DQ780352		EF101385	EU570259	Zalar et al. (2007), Dugan et al. (2008)
Cl. lycoperdinum	CBS 574.78C^RS	HM148115		HM148604	HM148359	Bensch et al. (2010)
Cl. macrocarpum	CBS 121623^NT	EF679375		EF679529	EF679453	Schubert et al. (2007)
Cl. montecillanum	CBS 140486^T	KT600406		KT600602	KT600504	Bensch et al. (2015)
Cl. myrtacearum	CBS 126350^ET	HM148117		HM148606	HM148361	Bensch et al. (2010)
Cl. ossifragi	CBS 842.91^ET	EF679381		EF679535	EF679459	Schubert et al. (2007)
Cl. oxysporum	CBS 125991^RS	HM148118		HM148607	HM148362	Bensch et al. (2010)
Cl. paracladosporioides	CBS 171.54^T	HM148120		HM148609	HM148364	Bensch et al. (2010)
Cl. parapenidielloides	CBS 140487^T	KT600410		KT600606	KT600508	Bensch et al. (2015)
Cl. penidielloides	CBS 140489^T	KT600412		KT600608	KT600510	Bensch et al. (2015)
Cl. perangustum	CBS 125996^T	HM148121		HM148610	HM148365	Bensch et al. (2010)
Cl. phaenocomae	CBS 128769^T	JF499837		JF499881	JF499875	Crous & Groenewald (2011)
Cl. phlei	CBS 358.69^ET	JN906981		JN907000	JN906991	Bensch et al. (2012)
Cl. phyllactiniicola	CBS 126352^T	HM148150		HM148639	HM148394	Bensch et al. (2010)
Cl. phyllophilum	CBS 125992^ET	HM148154		HM148643	HM148398	Bensch et al. (2010)
Cl. pini-ponderosae	CBS 124456^T	FJ936160		FJ936167	FJ936164	Schubert et al. (2009)
Cl. pseudiridis	CBS 116463^T	EF679383		EF679537	EF679461	Schubert et al. (2007)
Cl. pseudochalastosporioides	CBS 140490^T	KT600415		KT600611	KT600513	Bensch et al. (2015)
Cl. pseudocladosporioides	CBS 125993^T	HM148158		HM148647	HM148402	Bensch et al. (2010)
Cl. psychrotolerans	CBS 119412^T	DQ780386		EF101365	JN906992	Zalar et al. (2007), Bensch et al. (2012)
Cl. puyae	CBS 274.80A^T	KT600418		KT600614	KT600516	Bensch et al. (2015)
Cl. ramotenellum	CBS 121628^T	EF679384		EF679538	EF679462	Schubert et al. (2007)
Cl. rectoides	CBS 125994^T	HM148193		HM148683	HM148438	Bensch et al. (2010)
Cl. rhusicola	CBS 140492^T	KT600440		KT600637	KT600539	Bensch et al. (2015)
Cl. ruguloflabelliforme	CBS 140494^T	KT600458		KT600655	KT600557	Bensch et al. (2015)
Cl. rugulovarians	CBS 140495^T	KT600459		KT600656	KT600558	Bensch et al. (2015)
Cl. salinae	CBS 119413^T	DQ780374		EF101390	JN906993	Zalar et al. (2007), Bensch et al. (2012)
Cl. scabrellum	CBS 126358^T	HM148195		HM148685	HM148440	Bensch et al. (2010)
Cl. silenes	CBS 109082^T	EF679354		EF679506	EF679429	Schubert et al. (2007)
Cl. sinuosum	CBS 121629^T	EF679386		EF679540	EF679464	Schubert et al. (2007)
Cl. soldanellae	CBS 132186^NT	JN906982		JN907001	JN906994	Bensch et al. (2012)
Cl. sphaerospermum	CBS 193.54^NT	DQ780343		EF101380	EU570261	Zalar et al. (2007), Dugan et al. (2008)
Cl. spinulosum	CBS 119907^T	EF679388		EF679542	EF679466	Schubert et al. (2007)
Cl. subcinereum	CBS 140465^T	LN834433		LN834529	LN834617	Sandoval-Denis et al. (2016)
Cl. subinflatum	CBS 121630^T	EF679389		EF679543	EF679467	Schubert et al. (2007)
Cl. subtilissimum	CBS 113754^T	EF679397		EF679551	EF679475	Schubert et al. (2007)
Cl. subuliforme	CBS 126500^T	HM148196		HM148686	HM148441	Bensch et al. (2010)
Cl. succulentum	CBS 140466^T	LN834434		LN834618	LN834530	Sandoval-Denis et al. (2016)
Cl. tenellum	CBS 121634^T	EF679401		EF679555	EF679479	Schubert et al. (2007)
Cl. tenuissimum	CBS 125995^ET	HM148197		HM148687	HM148442	Bensch et al. (2010)
Cl. tuberosum	CBS 140693^T	LN834417		LN834601	LN834513	Sandoval-Denis et al. (2016)
Cl. uredinicola	ATCC 46649	AY251071		HM148712	HM148467	Braun et al. (2003), Bensch et al. (2010)
Cl. variabile	CBS 121635^ET	EF679402		EF679556	EF679480	Schubert et al. (2007)
Cl. varians	CBS 126362^T	HM148224		HM148715	HM148470	Bensch et al. (2010)
Cl. velox	CBS 119417^T	DQ780361		EF101388	JN906995	Zalar et al. (2007), Bensch et al. (2012)
Cl. verrucocladosporioides	CBS 126363^T	HM148226		HM148717	HM148472	Bensch et al. (2010)
Cl. versiforme	CBS 140491^T	KT600417		KT600613	KT600515	Bensch et al. (2015)
Cl. welwitschiicola	CBS 142614^T	KY646223		KY646226	KY646229	Marin-Felix et al. (2017)
Cl. xanthochromaticum	CBS 140691^T	LN834415		LN834599	LN834511	Sandoval-Denis et al. (2016)
Cl. xylophilum	CBS 125997^T	HM148230		HM148721	HM148476	Bensch et al. (2010)

¹ATCC: American Type Culture Collection, Virginia, USA; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CPC: Culture collection of Pedro Crous, housed at Westerdijk Fungal Biodiversity Institute. ^{T, ET, NT}and ^RS indicate ex-type, ex-epitype, ex-neotype and reference strains, respectively.

²ITS: internal transcribed spacers and intervening 5.8S nrDNA; act: partial actin gene; tef1: partial translation elongation factor 1-alpha gene.

Bensch K, Braun U, Groenewald JZ, et al. (2012). The genus Cladosporium. Studies in Mycology 72: 1–401.
Bensch K, Groenewald JZ, Braun U, et al. (2015). Common but different: The expanding realm of Cladosporium. Studies in Mycology 82: 23–74.
Bensch K, Groenewald JZ, Dijksterhuis J, et al. (2010). Species and ecological diversity within the Cladosporium cladosporioides complex (Davidiellaceae, Capnodiales). Studies in Mycology 67: 1–94.
Braun U, Crous PW, Dugan FM, et al. (2003). Phylogeny and taxonomy of cladosporium-like hyphomycetes, including Davidiella gen. nov., the teleomorph of Cladosporium s. str. Mycological Progress 2: 3–18.
Crous PW, Groenewald JZ (2011). Why everlastings don’t last. Persoonia 26: 70–84.
Crous PW, Shivas RG, Quaedvlieg W, et al. (2014). Fungal Planet description sheets 214–280. Persoonia 32: 184–306.
Crous PW, Tanaka K, Summerell BA, et al. (2011). Additions to the Mycosphaerella complex. IMA Fungus 2: 49–64.
Dugan FM, Braun U, Groenewald JZ, et al. (2008). Morphological plasticity in Cladosporium sphaerospermum. Persoonia 21: 9–16.
Marin-Felix Y, Groenewald JZ, Cai, L, et al. (2017). Genera of phytopathogenic fungi: GOPHY 1. Studies in Mycology xxxx.
Sandoval-Denis M, Gené J, Sutton DA, et al. (2016). New species of Cladosporium associated with human and animal infections. Persoonia 36: 281–298.
Schubert K, Greslebin A, Groenewald JZ, et al. (2009). New foliicolous species of Cladosporium from South America. Persoonia 22: 111–122.
Schubert K, Groenewald JZ, Braun U, et al. (2007). Biodiversity in the Cladosporium herbarum complex (Davidiellaceae, Capnodiales), with standardisation of methods for Cladosporium taxonomy and diagnostics. Studies in Mycology 58: 105–156.
Zalar P, de Hoog GS, Schroers H-J, et al. (2007). Phylogeny and ecology of the ubiquitous saprobe Cladosporium sphaerospermum, with descriptions of seven new species from hypersaline environments. Studies in Mycology 58: 157–183.