[BoDD logo]


Google uses cookies
to display context-
sensitive ads on this
page. Learn how to
manage Google cookies
by visiting the

Google Technologies Centre

 ▼ ▼ ▼ ▼ ▼ ▼ ▼


 ▲ ▲ ▲ ▲ ▲ ▲ ▲

[BBEdit logo]





Fungi are non-photosynthesising, heterotrophic organisms that derive their energy from a saprophytic or parasitic existence. They are unicellular, amoeboid, or filamentous, never having the leaves, stems, and roots characteristic of higher plants. Reproduction occurs by sexual or asexual spore formation.

Mycologists now regard the fungi as being distinct from plants, and accordingly that they should be classified within their own kingdom, namely the Fungi or Mycetae (Talbot 1971, Cooper 1985, Holmes 1983). Recent treatments now segregate the fungi in the widest sense into three kingdoms, namely the Fungi (i.e. true fungi), the Chromista (i.e. the pseudofungi), and the Protozoa (i.e. the slime moulds). These three kingdoms are each further subdivided into phyla or divisions (Hawksworth et al. 1995):

Ascomycota (loosely: "ascomycetes")
Basidiomycota (loosely: "basidiomycetes")
Zygomycota (loosely: "zygomycetes")

This monograph includes a consideration of the pseudofungi and slime moulds where these micro-organisms have produced dermatological symptoms.

A further division within the kingdom Fungi comprises the mitosporic fungi or "fungi imperfecti" (formerly the Deuteromycotina or Deuteromycetes); fungi that are known only to reproduce asexually are classified therein. Strictly, taxonomic ranks within the mitosporic fungi should be prefixed with the term "form-", as in "form-genus", "form-class", "form-order", etc. This is because classification is based on outward appearance only - since characteristics concerned with sexual reproduction are unknown (Holmes 1983). On occasion, a fungus classified in the mitosporic fungi is persuaded to undergo sexual reproduction, and therefore to show its "perfect" state, whence it becomes evident that it should be reclassified within one of the other divisions, usually the Ascomycota or less commonly the Basidiomycota. When such an event occurs, the fungus has to be provided with a new name. It should be noted that both names remain valid and their usage is determined by the morphological state in which the fungus is being referred to. Contemporary terminology refers to the imperfect states as anamorphs, and to the perfect states as teleomorphs (see Domsch et al. 1980).

One of the purposes of considering the higher plants as species within genera belonging to families is to enable some predictions to be made as to the hazard that any particular plant is likely to present, on the basis that related plants are likely to contain similar phytochemicals. It should be evident, therefore, that classification of the fungi imperfecti into form-families does not produce any useful information concerning genetic relationships between individual form-species or even (but to a lesser extent) form-genera. This may be illustrated by the observation that the conidial (i.e. imperfect) forms of Nectria Fr. (fam. Nectriaceae) are distributed among 14 form-genera and four form-families of the fungi imperfecti (Talbot 1971). For this reason, the fungi imperfecti are considered below as individual form-species within form-genera in the division Mitosporic Fungi (which is included in its correct alphabetical position as if it were a family), with no attempt being made to classify into form-families. Many form-family names may be encountered in the literature, including Bactridiaceae, Conidiosporiaceae, Cryptococcaceae, Dematiaceae, Excipulaceae, Geotrichaceae, Helminthosporiaceae, Leptostromataceae, Melanconiaceae, Moniliaceae, Mucedinaceae, Sphaeropsidaceae, Sporobolomycetaceae, Stilbellaceae (or Stilbaceae), and Tuberculariaceae. These have not been used in this text.

Fungi have many economic uses and human contact with either the fungi or their secondary metabolites occurs commonly. Several species are often used for food, for example Agaricus bisporus Pilát (the cultivated mushroom), A. campestris L. ex Fr. (the field mushroom), and A. arvensis Schaeff. ex Secr. (the horse mushroom) - all belonging to the fam. Agaricaceae; the cep (Boletus edulis Bull. ex Fr., fam. Boletaceae); the Japanese "shiitake" (Lentinula edodes Pegler) and "matsutake" (Tricholoma matsutake - both of the fam. Tricholomataceae); Volvariella esculenta Singer, fam. Pluteaceae, a South-East Asian species; truffles belonging to the genera Tuber Mich., Terfezia Tul. (both of the fam. Terfeziaceae) and Stephensia Tul. & C. Tul. (syn. Elderia McLennan), fam. Humariaceae; and morels (Morchella St Amans, fam. Morchellaceae). A number of species are utilised in the manufacture of various foods and beverages. Certain Penicillium Link ex Fr., Rhizopus Ehrenb., Aspergillus Mich. ex Fr. (all fungi imperfecti) and Saccharomyces Meyen ex E. Hansen (fam. Saccharomycetaceae) species are valuable in this respect (see Talbot 1971).

Inhalation of fungal spores from various species can produce allergic alveolitis (McCombs 1972, Seaton & Morgan 1984). Towey et al. (1932) described a "coniosporiosis" of timber workers who developed a severe bronchial asthma after inhaling the spores of the mitosporic fungus Cryptostroma corticale Gregory & Walker (syn. Coniosporium corticale Ellis & Everhart) from infected maple (Acer L. spp., fam. Aceraceae) logs.

A number of medicinally used antibiotics are derived from fungi (Reynolds 1996, Domsch et al. 1980). Penicillins are derived from Penicillium chrysogenum Thom (syn. P. notatum Westling). Most of the more modern penicillins are produced semi-synthetically from 6-aminopenicillanic acid derived from the Penicillium culture. A related group of antibiotics, the cephalosporins, are derived from cultures of Acremonium chrysogenum W. Gams (syn. Cephalosporium acremonium Corda), the active nucleus in this group being 7-aminocephalosporanic acid. Fusidium coccineum Fuckel is the source of fusidic acid, an antibiotic with a steroid structure. Other less well known antibiotics include adicillin from Emericellopsis salmosynnematum Groskl. & Swift (provisionally classified in the fam. Trichocomaceae) and fusafungine from Fusarium lateritium Nees ex Link (teleomorph: Gibberella baccata Sacc., fam. Nectriaceae).

The pathogenicity of certain fungi, often manifesting itself as a skin disorder, is perhaps the commonest form of interaction between man and members of the fungus kingdom. This aspect is covered in the monographs below but interested readers are advised to consult an appropriate clinical mycology text for more detailed coverage. Fungal infections acquired from contact with plant material also occur fairly commonly. In particular, sporotrichosis (see Sporothrix schenckii Hektoen & Perkins) should be suspected in patients with subcutaneous lesions who handle thorny plants, timber, or sphagnum moss (Kaufman & Reiss 1985). Otherwise, both irritant and allergenic reactions have been reported following contact with various fungi or their extractives. No one group of compounds can be incriminated as being a characteristic cause of a fungus-induced dermatosis. Dermatological effects following deliberate or inadvertent ingestion of certain fungi have also been reported. These include ergotism from Claviceps purpurea Tul. (fam. Clavicipitaceae) and the disulfiram-like reaction following the ingestion of alcohol and ink cap fungi (Coprinus atramentarius Fr., fam. Coprinaceae).



This family of basidiomycetes comprises about 1340 species in 85 genera (Kirk et al. 2008). Fungi previously classified in the Lycoperdaceae (Hawksworth et al. 1995) are now regarded as belonging to this family (Kirk et al. 2008).

Agaricus bisporus Pilát

Workers in canneries who prepare mushrooms are subject to keratitis, lachrymation, and other ocular affections accompanied by such constitutional symptoms as vomiting and jaundice. Helvellic acid which becomes vaporised in the air when the mushrooms are washed in cold water is believed to be the cause (Schwartz et al. 1957), but a more likely culprit is p-hydroxymethyl phenylhydrazine (Chilton 1978).

Agaricus campestris L. ex Fr.

These mushrooms were reported to be the cause of contact dermatitis of the hands and face in a male mushroom grower. Patch tests implicated the mushroom rather than the nicotine-containing insecticide used (Hopkins 1952, 1953).

Inhalation of fungal particles from mushroom compost can produce mushroom worker's lung, a form of allergic alveolitis (Seaton & Morgan 1984).

Calvatia gigantea Lloyd
[syns Bovista gigantea Gray, Langermannia gigantea Rostk., Lycoperdon bovista L., Lycoperdon giganteum Batsch.]
Giant Puff Ball

The use of the spongy interior of this puff ball as a haemostat is noted by Wren (1975).

[Further information available but not yet included in database]

Lycoperdon fontanesii Durieu & Mont.

[Information available but not yet included in database]



Members of this family of ascomycetes were formerly classified in the Gymnoascaceae (Hawksworth et al. 1995).

Ajellomyces capsulatus McGinnis & Katz
[syn. Emmonsiella capsulata Kwon-Chung]
Anamorph: Histoplasma capsulatum Darling
[syns Coccidioides farciminosa Vuill., Cryptococcus farciminosus Rivolta & Micelloni ex Guég., Cryptococcus capsulatus Castell. & Chalm., Histoplasma capsulatum Darling var. farciminosum R.J.Weeks, A.A.Padhye & Ajello, Histoplasma farciminosum Redaelli & Cif., Torulopsis capsulata F.P.Almeida]

The spores of these organisms can cause the lung disease histoplasmosis if inhaled (Roberts et al. 1984). Most patients develop an hypersensitivity state that is readily demonstrated by skin tests (Larsh & Goodman 1985).

The disease may exhibit cutaneous symptoms such as erythema nodosum and multiforme. African histoplasmosis caused by Histoplasma duboisii Vanbreus. (syn. Histoplasma capsulatum var. duboisii Cif.) may present with painless nodules, abscesses, or ulcers, or acneiform papules (Roberts et al. 1984).

Histoplasma farciminosum Redaelli & Cif. is a cause of epizootic lymphangitis in animals (Ainsworth & Austwick 1973).

Ajellomyces dermatitidis McDonough & A.L. Lewis
Anamorph: Zymonema dermatitidis Gilchrist & W.R. Stokes
[syns Blastomyces dermatitidis Gilchrist & W.R. Stokes, Cryptococcus dermatitidis Lodder]

This organism can cause blastomycosis, also known as Gilchrist's disease, a systemic/cutaneous mycosis (Ainsworth & Austwick 1973, Larsh & Goodman 1985, Roberts et al. 1984).



Members of this family of ascomycetes were formerly classified in the Gymnoascaceae (Hawksworth et al. 1995).

Arthroderma Berk.

About 15 species of Arthroderma are known, all of which are keratophilic. These fungi commonly occur near animal burrows and bird nests, but a number are parasitic and hence pathogenic to man and other mammals causing the diseases known as ringworm.

These organisms are better known as their anamorphs of the form-genus Trichophyton Malmsten. In fact, most of the pathogenic Trichophyton species are considered under FUNGI - MITOSPORIC since their teleomorphs are as yet unknown.

Arthroderma benhamiae Ajello & Cheng
Anamorph: Trichophyton mentagrophytes Blanchard
[syns Trichophyton asteroides Sabouraud, Trichophyton granulosum, Trichophyton gypseum Bodin, Trichophyton interdigitale Priestley, Epidermophyton interdigitale Kesteven, Trichophyton quinckeanum MacLeod & Munde]

This organism is a common cause of ringworm in humans and animals (Ainsworth & Austwick 1973, Ajello & Padhye 1980, Roberts et al. 1984). It can also produce a vesicular form of athlete's foot (Sande & Mandell 1985).

Arthroderma simii Stockdale, Mackenzie, & Austwick
[syn. Epidermophyton simii Pinoy]
Anamorph: Trichophyton simii Stockdale, Mackenzie, & Austwick

This organism may cause ringworm in various animals and man (Ainsworth & Austwick 1973).

Arthroderma uncinatum Dawson & Gentles
Anamorph: Trichophyton ajelloi Ajello
[syn. Keratinomyces ajelloi Vanbreuseghem]

Skin infections caused by this organism have been observed rarely in animals and in isolated cases in man (Ainsworth & Austwick 1973, Domsch et al. 1980).

Nannizzia Stockdale

This genus of ascomycetes is closely related to Arthroderma Berk. and is similarly capable of producing the disease known as ringworm. Generally, these organisms are only infrequently found to be responsible for human ringworm infections, being more commonly found affecting livestock (Ainsworth & Austwick 1973, Ajello & Padhye 1980).

The organisms are better known as their anamorphs in the form-genus Microsporum Gruby, under which heading some are considered because their teleomorphs are as yet unknown.

The following species have been reported as causative organisms in human ringworm infections (Ainsworth & Austwick 1973, Domsch et al. 1980, Ajello & Padhye 1980):

Nannizzia cajetani Ajello
Anamorph: Microsporum cookei Ajello
Nannizzia fulva Stockdale
Anamorph: Microsporum fulvum Uriburu
Nannizzia grubyia Georg, Ajello, Friedman, & Brinkman
Anamorph: Microsporum vanbreuseghemii George, Ajello, Friedman, & Brinkman
Nannizzia gypsea Stockdale
Anamorph: Microsporum gypseum Guiart & Grigorakis
Nannizzia obtusa Dawson & Gentles
Anamorph: Microsporum nanum Fuentes
Nannizzia persicolor Stockdale
Anamorph: Microsporum persicolor Guiart & Grigorakis
[syn. Trichophyton persicolor Sabouraud] 



This family of basidiomycetes comprises about 16 species in 5 genera (Hawksworth et al. 1995).

Auricularia polytricha Sacc.
Black Tree Fungus, Mo-Er

This fungus is used as a texture food in many Szechwanese and Mandarin dishes. It is cultivated on oak (Quercus L., fam. Fagaceae) poles in China (Hawksworth et al. 1995).

Hammerschmidt (1980) described a patient in whom ingestion of a large quantity of the fungus produced a transient inhibition of platelet aggregation resulting in purpura and epistaxis.



This family is classified within the Basidiomycota. A distinguishing feature of members of this family is the presence of pores in place of gills on the fruiting bodies (mushrooms). Many of the boletes are edible (Grimes 1978).

Vetter (1993) reported that Boletus edulis Bull. ex Fr. (cep or penny bun) and Boletus luridus Schaeff. ex Fr. are selenium accumulators, the highest concentrations found being 3mg/100g (30 ppm). Whilst selenium is an essential trace nutrient for humans, excessive consumption may produce symptoms of poisonining (selenosis) that include hair loss and brittle nails (see Aster venustus M.E.Jones, fam. Compositae).

Suillus luteus S.F. Gray
[syn. Boletus luteus L. ex Fr.]
Slippery Jack

Two cases of cutaneous sensitisation to Boletus luteus were reported; one patient also showed a weakly positive reaction to Boletus edulis Bull. ex Fr. (Hellerström 1941). See also Lactarius deliciosus Fr., fam. Russulaceae and Ramaria flava Quélet, fam. Gomphaceae.



Macrophomina phaseolina Goid.
[syns Macrophoma phaseolina Tassi, Rhizoctonia bataticola E.J.Butler, Rhizoctonia lamellifera W.Small, Sclerotium bataticola Taubenh., etc.]

This organism has been reported to cause infection in humans, particularly in immunosuppressed patients. The infection may present as a cutaneous cellulitis or as an ocular keratitis (Schwartz & Kapila 2020).



This family of about 224 species in 23 genera is classified in the Ascomycota.

Claviceps Tul.

Members of this genus are pathogens of various grasses and cereals, forming sclerotia known commonly as ergots. Inadvertent ingestion of the sclerotia of certain Claviceps in flour prepared from infected cereal crops can produce gangrene of the extremities, often referred to as St Anthony's fire or ergotism (Trease & Evans 1966). Claviceps purpurea Tul., perhaps the most common species, is the original source of ergotamine, ergometrine, and other ergot alkaloids used in medicine (Talbot 1971, Robbers et al. 1996).



The family is classified in the Basidiomycota.

Coleosporium Lév.

The genus is one of the 100 or so genera of fungi known as rusts [see also Pucciniaceae below] and classified in the order Uredinales (Talbot 1971). The genus has also been classified in the family Melampsoraceae.

Coleosporium sonchi Lév. & Tul.
[syns Coleosporium sonchi-arvensis Lév., Uredo sonchi-arvensis Pers.]

Pammel (1911) lists this species as an irritant.



The family is classified in the Basidiomycota.

Serpula lacrimans Schröter
[syn. Merulius lacrimans Fr.]
Dry Rot

This well known wood-rotting fungus has also been classified in the family Meruliaceae.

Prosser White (1934) observed two adult males who suddenly developed an erythematous eruption on the face and exposed parts of the hands and arms following sweeping out a cellar containing a dry decaying wood. The eruptions subsided in 36 hours. He stated that the "dry rot" fungus was the probable irritant. It should be noted, however, that other species of wood-decaying fungus may be found in such a situation. Frankland & Hay (1951) also refer to allergic complaints from the dry rot fungus.



The family is classified in the Basidiomycota.

Coprinus atramentarius Fr.
Ink Cap

Although normally an edible fungus, poisoning occurs if alcohol is consumed up to 48 hours after ingestion. The symptoms are reddening of the face and difficulty in breathing, and resemble disulfiram poisoning. The compound responsible is coprine (Chilton 1978). The onset of symptoms varies from 20 minutes to 2 hours after consumption of alcohol (North 1967).



This is a family of about 812 species in 109 genera, classified in the Basidiomycota (Hawksworth et al. 1995).

Fomes fomentarius Kickx
[syn. Polyporus fomentarius L. ex Fr.]
Hoof Fungus, Tinder Fungus, Amadou, German Tinder, Oak Agaric, Surgeon's Agaric

Wren (1975), referring to Polyporus fomentarius, records that this fungus is prepared for use in folk medicine by being cut into slices, beaten, soaked in a solution of nitre [= KNO3 or NaNO3], and dried. It may then be used for arresting local haemorrhages, being applied with pressure to the affected part. Wren (1975) also notes that it may be used to treat ingrown toe-nails by being inserted between the nail and the flesh. It is not clear whether these medicinal uses are dependent upon the absorbency or other property of the fungus, or upon the properties of the nitre with which it has been impregnated. However, the oxidising properties of nitre would certainly contribute to its alternative use as a tinder material. According to Hawksworth et al. (1995), the fungus Phellinus ignarius Quél. (fam. Hymenochaetaceae) may also be soaked in nitre and used as a tinder material.



The family is classified in the Basidiomycota.

Hebeloma crustuliniforme Quélet
Poison Pie, Fairy-Cake Mushroom

This fungus smells of radish (North 1967). Whether or not isothiocyanates are responsible is unknown. See also Cruciferae.



The family is classified in the Ascomycota. It comprises about 185 species in 4 genera (Kirk et al. 2008).

Cladosporium Link ex Fr.

Cladosporium species, of which some 500 have been described, are amongst the most common air-borne fungi, and thus have a world wide distribution. They are particularly common on dying or dead plant material. Some have been identified with the teleomorphic genus Mycosphaerella Johanson, an ascomycete of the family Dothideaceae (Domsch et al. 1980).

Cladosporium species are occasionally reported from cutaneous, eye, and nail infections (McGinnis 1985).



This family is classifed in the Zygomycota. It comprises about 131 species in 11 genera (Hawksworth et al. 1995).

Basidiobolus haptosporus Drechsler
[syn. Basidiobolus meristosporus Drechsler]

This organism can produce a subcutaneous zygomycosis known as basidiobolomycosis. The disease usually occurs in children and is mainly seen in East Africa although cases elsewhere in the tropics have been reported (Tio et al. 1966, Greer & Friedman 1966, Roberts et al. 1984).

Conidiobolus coronatus Batko
[syns Entomophthora coronata Kevork., Conidiobolus coronatus Sriniv. & Thirum., Delacroixia coronata Kjøller]

This organism is typically associated with decaying insect remains in the soil (Talbot 1971).

Rhinoentomophthoromycosis, a zygomycosis also known as conidiobolomycosis, is produced by this organism. The disease is normally confined to the face, usually originating from the inferior turbinates of the nose from where it spreads slowly and progressively (Roberts et al. 1984). The disease occurs in the tropics (Greer & Rogers 1985, Domsch et al. 1980).

In a case described by Herstoff et al. (1978), the patient failed to respond with an inflammatory reaction when challenged with croton oil (see Croton tiglium L., fam. Euphorbiaceae). This observation is intriguing but difficult to interpret since no further information was provided regarding mode of application and effects on control subjects.



Ramaria flava Quélet
[syn. Clavaria flava Fr.]
Fairy Clubs

Two cases of cutaneous sensitisation to edible mushrooms (Boletus edulis Bull. ex Fr. and B. luteus L. ex Fr., fam. Boletaceae; Lactarius deliciosus Fr., fam. Russulaceae; and Clavaria flava) were reported. In one case hypersensitivity was also demonstrated after eating the mushrooms in question, fried. Boiling seems to destroy the antigenic effect of mushrooms. From a practical point of view, it is important to be familiar with the cases, as for example in differentiating between similar eruptions caused by sunlight or hypersensitivity to autumn flowers (Hellerström 1941).



This family of ascomycetes comprises 68 species in 11 genera (Hawksworth et al. 1995).

Gyromitra esculenta Fr.
Lorchel, Turban Fungus

The skin and the eyes may be irritated by handling the fungus (North 1967).

When damaged, the fungus releases monomethylhydrazine and methylformylhydrazine from stored hydrazones of ethanal [= gyromitrin], pentanal, and hexanal (Chilton 1978).



This family of ascomycetes comprises 84 species in 13 genera (Kirk et al. 2008).

Cladophialophora bantiana de Hoog, Kwon-Chung & McGinnis
[syns Cladosporium bantianum Borelli, Torula bantiana Sacc., Xylohypha bantiana McGinnis, A.A.Padhye, Borelli & Ajello]

This organism may cause cladosporiosis (Rogers 1980). It may cause subcutaneous phaeohyphomycosis (McGinnis 1983) and is the most frequently reported aetiological agent of cerebral phaeohyphomycosis (McGinnis 1985).

Cladophialophora carrionii de Hoog, Kwon-Chung & McGinnis
[syns Cladophialophora ajelloi Borelli, Cladosporium carrionii Trejos]

This form-species is a causative organism of chromoblastomycosis (Vollum 1977, Roberts et al. 1984, McGinnis 1985). See also Phialophora Medlar.

Exophiala dermatitidis De Hoog
[syns Hormiscium dermatitidis Kano, Phialophora dermatitidis Emmons, Wangiella dermatitidis McGinnis]

This organism is an important aetiological agent of chromoblastomycosis and of phaeohyphomycosis (Ajello et al. 1974, McGinnis 1978, Roberts et al. 1984). The fungus has most frequently been found in patients in Japan (McGinnis 1983, 1985). See also Phialophora Medlar.

Exophiala jeanselmei McGinnis & A.A.Padhye
[syns Phialophora jeanselmei Emmons, Phialophora gougerotii Borelli, Sporotrichum gougerotii Matr., Torula jeanselmei Langeron]

This organism is an occasional cause of mycetomas and chromomycosis in the USA (McGinnis 1978, Domsch et al. 1980, Roberts et al. 1984, McGinnis 1985, Padhye & Ajello 1985). See also Phialophora Medlar. According to the classification of Mackinnon et al. (1949), this organism produces the so-called Type III maduromycosis [see Madurella grisea Mackinnon, Ferrada, & Montemayer below].

Exophiala moniliae De Hoog


Exophiala spinifera McGinnis
[syn. Phialophora spinifera Nielsen & Conant]

These organisms have been reported as agents of phaeohyphomycosis causing subcutaneous cysts (McGinnis 1978, 1985). See also Phialophora Medlar.

Exophiala werneckii v. Arx
[syns Cladosporium werneckii Horta, Dematium werneckii Dodge, Pullularia werneckii de Vries, Aureobasidium mansonii Cooke]

This organism is the causative agent of tinea nigra, a superficial phaeohyphomycosis characterised by dark macular patches on the palms or palmar aspects of the wrists and fingers (Ajello & Padhye 1980, McGinnis 1985). The use of the term tinea to describe the disease is misleading since it is not a form of ringworm (Roberts et al. 1984, McGinnis et al. 1985).

There is a great deal of confusion in the literature concerning the taxonomy of Exophiala werneckii and its relationship with Microsporum mansonii Castellani. McGinnis (1979) attempted to clarify the situation and concluded that the confusion arose when a case of pityriasis versicolor was misdiagnosed as tinea nigra. The organism responsible was named Microsporum mansonii by Castellani and later renamed Aureobasidium mansonii by Cooke. However, Cooke was actually naming the causative organism of tinea nigra, so his name is now considered to be a synonym of Exophiala werneckii. Since Castellani's name actually referred to the causative organism of pityriasis versicolor, it is now considered to be a synonym of Malassezia furfur Baill.

Fonsecaea pedrosoi Negroni
[syns Acrotheca pedrosoi Fonseca & Leāo, Fonsecaea cladosporium R.E.Powell, Hormodendrum pedrosoi Brumpt, Phialophora pedrosoi Redaelli & Cif., Rhinocladiella pedrosoi Schol-Schwarz, Trichosporum pedrosoi Brumpt]

Both species are agents of chromoblastomycosis (Vollum 1978, McGinnis 1983, Roberts et al. 1984, McGinnis 1985). See also Phialophora Medlar.

The genera Fonsecaea Negroni and Cladosporium Link ex Fr. are difficult to differentiate (Rogers 1980).

Phialophora Medlar

Teleomorphic genera associated with Phialophora species include Pyrenopeziza Fuckel (fam. Dermateaceae), Mollisia P.Karst. (fam. Dermateaceae), Ascocoryne Groves & Wilson (fam. Helotiaceae), Coniochaeta Cooke (fam. Xylariaceae), and Gaeumannomyces v. Arx & Olivier (fam. Gnomoniaceae) (Domsch et al. 1980).

Several species of Phialophora are well recognised as aetiological agents of chromoblastomycosis and phaeohyphomycosis. Chromoblastomycosis (also known as chromomycosis) is a chronic infection caused by dark-coloured yeast-like fungi which involves the dermis and epidermis, and is characterised by a warty, often foul-smelling proliferation of the skin. Phaeohyphomycosis (also known as cystic chromomycosis) is an infection of the skin in which brown-pigmented fungi are present in a hyphal or pseudohyphal form. In subcutaneous tissues, cystic lesions are the most frequently recognised form of infection (Roberts et al. 1984). McGinnis et al. (1985) discuss the nomenclature of chromoblastomycosis and phaeohyphomycosis and give several obsolete synonyms for each disease.

Tschen et al. (1984) described two cases of chromomycosis in which pigmented elements of unidentified fungal material were found within or on embedded wood splinters associated with foreign body reactions. Similar cases have been described by Mehregan & Rudner (1980) and by Vollum (1977).

Pathogenic species include (Ajello et al. 1974, Vollum 1977, McGinnis 1978, 1983, Domsch et al. 1980, Roberts et al. 1984, McGinnis 1985):

Phialophora bubakii Schol-Schwarz
Phialophora gougerotii Borelli
[syn. Sporotrichum gougerotii Matruchot]
Phialophora hoffmannii Schol-Schwarz
[syn. Phialophora mutabilis Scol-Schwarz]
Phialophora parasitica Ajello, Georg, & Wang
Phialophora repens Conant
[syn. Cadophora repens Davidson]
Phialophora richardsiae Conant
[syn. Cadophora richardsiae Nannf.]
Phialophora verrucosa Medlar
[syns Phialophora americana Hughes, Cadophora americana Nannf.] 

Phialophora gougerotii has been reported to have caused eye infections (Laverde et al. 1973, McGinnis 1985).

Rhinocladiella Nannf.

This genus of 12-15 species includes some "cellar-fungi" such as R. cellaris M.B.Ellis and R. ellisii D. Hawksw.

Rhinocladiella aquaspersa Schell, McGinnis, & Borelli
[syn. Acrotheca aquaspersa Borelli]

This organism may rarely cause chromoblastomycosis (McGinnis 1983). See also Phialophora Medlar.

Rhinocladiella compacta De Hoog
[syns Fonsecaea compacta Carrión, Hormodendrum compactum Carrión, Phialophora compacta Redaelli & Cif., Rhinocladiella compacta Schol-Schwarz]

This species is a cause of chromoblastomycosis (Vollum 1978, McGinnis 1983, Roberts et al. 1984, McGinnis 1985), a chronic fungal infection of the skin and the subcutaneous tissue caused by traumatic inoculation of the fungus through the skin. See also Fonsecaea pedrosoi Negroni below.



This family of basidiomycetes comprises 443 species of wood-rotting fungi in 23 genera (Hawksworth et al. 1995).

Inonotus tabacinus G. Cunn.
[syns Cyclomyces tabacinus Pat., Mucronoporus tabacinus Ellis & Everh., Polyporus tabacinus Mont.]

Marriott et al. (2006) in Australia reported the isolation of Cyclomyces tabacinus from a large cystic lesion anterior to the ankle of a 75-year-old retired dairy farmer who was being treated with a corticosteroid for rheumatoid arthritis. The source of the infection was uncertain as the patient could not recall any lacerations, splinters, or puncture wounds that might have explained the infection. This species is normally encountered as a bracket fungus on tree trunks, logs, and stumps.



The family is classified in the Ascomycota. It comprises about 219 species in 4 genera (Hawksworth et al. 1995).

Leptosphaeria Ces. & de Not.

About 100 species are recognised in this genus; the organisms occur widely as plant pathogens (Hawksworth et al. 1995). The genus has been regarded by some authorities as belonging to the Pleosporaceae and by others as belonging to the Phaeosphaeriaceae.

Leptosphaeria senegalensis Baylet, Camain, & Segretain


Leptosphaeria tompkinsii El-Ani

Cases of mycetoma caused by these organisms have been reported from northern tropical West Africa (Padhye & Ajello 1985).



This family of ascomycetes, also known as the Lophotrichaceae, comprises some 11 genera and 61 species found in soil and dung (Hawksworth et al. 1995).

Pseudallescheria boydii McGinnis, A.A.Padhye & Ajello
[syns Petriellidium boydii Malloch, Allescheria boydii Shear]
Anamorph: Scedosporium apiospermum Sacc. ex Castell. & Chalm.
[syns Monosporium apiospermum Sacc., Aleurisma apiospermum Maire]

"White grain" mycetomas, maduromycosis (madura foot), and other fungal infections produced by this organism have been reported from Australia and West Africa (Domsch et al. 1980, Roberts et al. 1984, Padhye & Ajello 1985).


(Fungi Imperfecti)


Acremonium Link ex Fr.

Many of the organisms hitherto placed in the form-genus Cephalosporium Corda are now considered to be species of Acremonium Link ex Fr. The name Cephalosporium acremonium Corda referred to commonly in the medical literature is actually of uncertain application. The vast majority of reports referring to the production of the antibiotic cephalosporin C by C. acremonium refer to Acremonium chrysogenum W. Gams; the C. acremonium that may cause mycetomas is now called Acremonium kiliense Grütz (Domsch et al. 1980).

The following organisms have been identified as causes of mycetomas (Domsch et al. 1980, Padhye & Ajello 1985):

Acremonium falciforme Gams
[syn. Cephalosporium falciforme Carrión]
Acremonium kiliense Grütz
Acremonium recifei Gams
[syn. Cephalosporium recifei Leāo & Lôbo] 

Acremonium kiliense and Acremonium potroni auct. may cause eye infections (Rebell & Forster 1980).

The teleomorphic genera Nectria Fr. (fam. Nectriaceae), Emericellopsis Van Beyma (provisionally classified in the fam. Pseudoeurotiaceae), and Mycoarachis Malloch & Cain (provisionally classified in the fam. Cephalothecaceae) have been identified with certain Acremonium species (Domsch et al. 1980).

Acremonium chrysogenum W. Gams

This organism provides 7-aminocephalosporanic acid from which semi-synthetic cephalosporin-type antibiotics are produced. A disulfiram reaction (see also Coprinus atramentarius Fr., fam. Coprinaceae) has been reported to occur following the ingestion of alcohol and certain cephalosporin-type antibiotics. The reaction appears to occur only with those compounds possessing the methyltetrazolethiol group at C-3, for example latamoxef, cefoperazone, and cephamandole (Stockley 1985).

Rudzki & Rebendel (1984) observed 3 positive patch test reactions to cefradine among 20 pharmaceutical workers and nurses screened for sensitivity to this antibiotic.

Tuft (1975) described the occurrence of contact urticaria in a chemist working with cephalosporins. Epicutaneous challenge tests with cephalothin sodium and other cephalosporins (unnamed) produced whealing.

Alternaria Nees ex Fr.

Members of this form-genus of some 44 species are common plant parasites, causing leaf spot and stem rot diseases (Martin 1969). They are occasionally implicated as agents of phaeohyphomycosis (see also Phialophora Medlar) and have been associated with infections involving bone, cutaneous tissue, ears, eyes, and urinary tract. Only A. alternata Keissler appears to have been positively identified in this respect (McGinnis 1985).

Alternaria alternata Keissler
[syns Alternaria tenuis Nees, Torula alternata Fr.]

Campbell & White (1989) noted that this organism may cause localised tissue destruction, usually in the nasal region and presumably following germination of inhaled conidia, in patients with AIDS.

A crude physiological saline extract of Alternaria tenuis produced positive patch test reactions in 1 of 6 patients with seasonal eczematous dermatitis; a crude ether extract failed to elicit a response in that patient (Fujisawa et al. 1966).

The possible role of the spores of this and other Alternaria Nees ex Fr. species as a cause of allergic eczema in dogs ("summer itch") and horses is discussed by Allen (1945) and by Austwick (1966).

[Further information available but not yet included in database]

Arthrinium sphaerospermum Fuckel
[syn. Papularia sphaerosperma von Höhnel]

The "maladie de la cannes de Provence" occurs on the shoulders of workmen who carry bundles of reeds (Arundo donax L., fam. Gramineae) affected by this fungus (Mandoul et al. 1954).

Aspergillus Mich. ex Fr.

Aspergillus species occur commonly in soil, particularly in association with decaying plant material (Domsch et al. 1980). Perfect states of these organisms have been classified in a number of ascomycete genera including Eurotium Link ex Fr., Emericella Berk. & Br., and Neosartorya Malloch & Cain, all in the family Trichocomaceae (Austwick & Longbottom 1980, Domsch et al. 1980, Hawksworth et al. 1995).

At least 8 of the 160 or so species of Aspergillus that have been described are known to be pathogenic to humans and to animals. Their pathogenicity is wide ranging: infection may occur within or on the surfaces of affected organs; sensitisation may occur through exposure to spores, hyphae, or metabolites by inhalation, ingestion, or by contact; and toxicosis may occur following ingestion of metabolites such as the aflatoxins or the ochratoxins (Austwick & Longbottom 1980, Roberts et al. 1984).

Autoerythrocyte sensitisation, which apparently resulted from mould antigen absorption onto the red cell, was found responsible for recurrent urticaria (Shelley & Florence 1961). The patient was exquisitely sensitive to trichophytin with a cross-sensitivity to mould-derived antibiotics and the common saprophytes Aspergillus glaucus Link, Aspergillus flavus Link, and an unidentified Alternaria Nees ex Fr. species.

Aspergillus flavus Link ex Gray

This species is a common cause of mycotic ocular infection (Rebell & Forster 1980).

Aspergillus fumigatus Fresen.
[syn. Aspergillus nigrescens Robin]

This species is the main cause of aspergillus infection and allergy. It is often found as a cause of infection of the turbinate and ethmoid regions of the nose (Austwick & Longbottom 1980). A. fumigatus is also a major cause of exogenous keratomycosis (Rebell & Forster 1974).

Abou-Gabal & Malik (1978) determined that this organism is a cause of dermatitis in chickens.

Aspergillus terreus Thom

Aspergillus species, and especially A. terreus, are the commonest cause of fungal infections of the ear. The organisms have also been found in burn eschar and subcutaneous abscesses, and may invade the nails (Paldrok & Hollström 1952, Austwick & Longbottom 1980).

Aspergillus versicolor Tiraboschi

A crude physiological saline extract of this organism produced positive patch test reactions in a patient with seasonal eczematous dermatitis; a crude ether extract failed to elicit a response (Fujisawa et al. 1966).

Candida Berkhout

Yeasts belonging to this form-genus are non-pigmented and have the capacity to form pseudomycelium in culture. Some can also form true mycelium. Candida albicans Berkhout and Candida glabrata S.A.Mey. & Yarrow are commonly found as commensal organisms in the gut of man, animals, and birds (Roberts et al. 1984).

Some Candida species are capable of producing opportunistic infections known variously as candidiasis, candidosis, thrush, and moniliasis. The disease may be mild, severe, or chronic, affecting the skin, nails, and mucous membranes including the eyes (Rebell & Forster 1980, Cooper & Silva-Hutner 1985), and may become systemic in immunosuppressed patients (Roberts et al. 1984).

The following species may produce disease (Roberts et al. 1984):

Candida albicans Berkhout
[syns Monilia albicans Zopf, Oidium albicans Robin]
Candida glabrata S.A.Mey. & Yarrow
[syns Torulopsis glabrata Lodder & De Vries, Cryptococcus glabratus Anderson]
Candida krusei Berkhout
[syn. Saccharomyces krusei Castellani]
Candida parapsilosis Langeron & Talice
[syn. Monilia parapsilosis Ashf.]
Candida pseudotropicalis Basgal
[syns Endomyces pseudotropicalis Castellani, Mycocandida pseudotropicalis Cif. & Red.]
Candida stellatoidea Langeron & Guerra
[syn. Monilia stellatoidea Jones & Martin]
Candida tropicalis Berkhout
[syn. Oidium tropicalis Castellani] 

The vast majority of infections are caused by Candida albicans.

Joseph et al. (1965) reported two cases of children seemingly with generalised allergic reactions to a Candida albicans infection. Presenting symptoms included febrile arthralgia, ecchymotic urticaria, bloody diarrhoea, and onychomycosis of the toes preceded by nasal congestion and pharyngitis. Oidomycin, an extract of Candida albicans, produced positive delayed intracutaneous test reactions in sensitised persons (Hedenius & Frisk 1955).

Cercospora apii Fr.

Emmons et al. (1957) described verrucous lesions of the face of a 12 year old Indonesian boy, from which this organism was isolated.

Cercosporella Sacc.

Members of this form-genus cause leaf-spot diseases of some plants (Martin 1969). An eruption of the wrists of women sorting and packing dried fruits from the Orient was attributed to a mould from this form-genus. The wrists rubbed against the sacking on the tables containing the fruit (Russ 1923).

Corynespora cassiicola Wei
[syn. Helminthosporium cassiicola Berk. & Curt.]

This organism has been reported to cause mycetomas (Padye & Ajello 1980).

Curvularia Boedijn

Curvularia species have been implicated in a number of opportunistic infections, and may particularly cause mycotic ulcers (Rebell & Forster 1980, McGinnis 1983, 1985). The following may be regarded as pathogenic:

Curvularia pallescens Boedijn
Curvularia senegalensis Subram.
Curvularia verruculosa Tandon & Bilgrami ex M.B.Ellis 

Other Curvularia species are considered under Cochliobolus Drechsler, fam. Pyrenophoraceae, in which genus the teleomorphs of Curvularia geniculata Boedijn and Curvularia lunata Boedijn are classified.

Cylindrocarpon tonkinense Bugnicourt

Mycotic keratitis caused by this species has been reported (Laverde et al. 1973, Rebell & Forster 1980).

Cyphellophora pluriseptata G.A.de Vries, Elders & Luykx

[Information available but not yet included in database]

The genus is regarded as an anamorphic member of the family Chaetothyriaceae (Kirk et al. 2008).

Drechslera Ito

Perfect states of Drechslera species have been classified in the genus Pyrenophora Fr., fam. Pleosporaceae.

Drechslera species have been implicated in a number of opportunistic infections (Rogers 1980, McGinnis 1985) and may cause phaeohyphomycosis (Roberts et al. 1984). See also Phialophora Medlar. The following species are recognised pathogens (McGinnis 1983, 1985):

Drechslera hawaiiensis M.B.Ellis
[syns Drechslera hawaiiensis Subram. & Jain, Helminthosporium hawaiiensis Bugnicourt]
Drechslera rostrata Richardson & Fraser
[syn. Helminthosporium rostratum Drechsler] 

Epidermophyton cruris Castellani & Chalm.

This organism may cause tinea cruris and tinea pedis in man (Hawksworth et al. 1995). According to Castellani & Chalmers (1919), a condition known in Ceylon (now, Sri Lanka) as "mango toe" was a streptococcal infection secondary to an infection between the toes caused by Epidermophyton cruris. This condition was known also as "dermatitis interdigitalis" and as "frieira".

Epidermophyton floccosum Langeron & Milochevitch
[syn. Epidermophyton inguinale Sabouraud]

This organism is a common causes of ringworm in human skin (Ajello & Padhye 1980, Roberts et al. 1984). See also Microsporum Gruby and Trichophyton Malmsten.

Fusarium Link ex Fr.

Over 142 species, varieties, and forms are recognised in this form-genus. Teleomorphic genera include Nectria Fr., Gibberella Sacc., and Calonectria de Not. of the family Nectriaceae, and Plectosphaerella Kleb. of the family Trichosphaeriaceae (Domsch et al. 1980).

Fusarium dimerum Pensig-Sacardo (syn. Fusarium episphaeria Snyder & Hansen), Fusarium nivale Ces., Rabenh., and Fusarium oxysporum Schlecht. emend Snyder & Hansen) are occasionally responsible for producing mycotic ulcers of the cornea (Rebell & Forster 1980).

Fusarium culmorum Sacc.
[syn. Fusisporium culmorum W.G.Sm.]

[Information available but not yet included in database]

Fusidium coccineum Fuckel
[syn. Ramularia coccinea Vestergr.]

This organism is the source of fusidic acid, an antibacterial antibiotic (Hawksworth et al. 1995, Reynolds 1996). A few isolated cases of contact sensitivity to topically applied sodium fusidate have been reported in the literature (Cronin 1980, Romaguera & Grimalt 1985). See also Mucor ramannianus Moller, fam. Mucoraceae.

Geomyces destructans Blehert & Gargas

[Information available but not yet included in database]

Hendersonula toruloidea Nattrass

This organism, a black or grey saprophytic mould that sometimes causes plant disease in tropical and subtropical regions, has been recognised as a cause of skin and nail infections almost identical with those produced by Trichophyton rubrum Sabouraud (Campbell et al. 1973, Roberts et al. 1984).

Lasiodiplodia theobromae Griffon & Maubl.
[syn. Botryodiplodia theobromae Pat.]

This organism may cause mycotic keratitis and onchomycosis (Laverde et al. 1973, McGinnis 1983).

Madurella grisea Mackinnon, Ferrada, & Montemayer


Madurella mycetomatis Brumpt

These organisms can cause "black grain" mycetomas (or maduromycosis) in Latin America and the USA (Roberts et al. 1984, Padhye & Ajello 1985).

According to the classification of Mackinnon et al. (1949), Type I maduromycosis is produced by Madurella americana Vuillemin, Madurella ikedai Gammel, and Madurella mycetomatis Brumpt whilst Type II maduromycosis is produced by Madurella grisea.

Malassezia furfur Baill.
[syns Monilia furfur Vuillemin, Pityrosporum furfur Emmons, Binford, & Utz, Pityrosporum orbiculare Gordon, Microsporon furfur Robin, Microsporum mansonii Castellani, Cladosporium mansonii Castellani, Malasezzia ovalis Acton & Panja, Pityrosporum ovale Castellani & Chalmers, Saccharomyces ovalis Bizzozero]

Tinea versicolor or pityriasis versicolor, a mild superficial dermatomycosis that usually manifests itself as slightly raised scaly patches on the neck and torso, is thought to be caused by this organism (Ajello & Padhye 1980, Cooper & Silva-Hutner 1985). The organism, a lipophilic yeast which is a member of the normal flora of the skin, becomes pathogenic when it transforms into a more mycelial form but the factors causing this are not clearly understood (Roberts et al. 1984). The complex polymorphic nature of the organism has only recently been recognised (Salkin & Gordon 1977).

The organism has been associated with blepharitis, dacryocystitis, dandruff, and seborrhoea (Benham 1947, Cooper & Silva-Hutner 1985).

Microsporum Gruby

This genus of about 15 species is responsible for ringworm in a variety of animals, but generally only infrequently or rarely in humans. Perfect states of these organisms are classified in the genus Nannizzia Stockdale (Ajello & Padhye 1980) under which heading are considered those with known teleomorphs. See also Epidermophyton Sabouraud and Trichophyton Malmsten.

The following Microsporum species have been reported to have caused ringworm infections in man (Ainsworth & Austwick 1973, Domsch et al. 1980, Roberts et al. 1984):

Microsporum canis Bodin
[syns Microsporum felineum Mewborn, Microsporum lanosum Sabouraud, Microsporum equinum Sabouraud]
Microsporum distortum Di Menna & Marples
Microsporum ferrugineum Ota 

Mycocentrospora acerina Deighton
[syn. Centrospora acerina Newhall]

This organism has been reported as a cause of systemic phaeohyphomycosis (Deighton & Mulder 1977, McGinnis 1983, Roberts et al. 1984, McGinnis 1985). See also Phialophora Medlar.

Myrothecium roridum Tode ex Steud.

Brian et al. (1948) observed positive patch test reactions to the culture filtrate of several strains of this fungus in 2 subjects.

Myrothecium verrucaria Ditm. ex Steud.
[syns Metarrhizium glutinosum S.Pope, Peziza verrucaria Alb. & Schw.]

The fungus has been isolated in the USA from deteriorated baled cotton and from Maryland soil. It causes rapid decomposition of cellulose. White & Downing (1947) showed that Metarrhizium glutinosum and Myrothecium verrucaria were one and the same organism.

Brian et al. (1947) experienced severe facial inflammation whilst processing large volumes of culture fluid during the isolation of glutinosin (an antifungal agent). The inflammatory activity was attributed to a volatile compound in the culture fluid. Self applied patch tests with the culture fluid produced positive reactions. Bowden & Schantz (1955) isolated three compounds possessing high dermatitic activity from a culture of this organism. Their structures were not elucidated.

Paecilomyces lilacinus Samson
[syn. Penicillium lilacinum Thom]

This organism is a significant cause of keratomycosis and endophthalmitis (Rebell & Forster 1980). It has also been isolated from chronic cutaneous facial lesions (Takayasu et al. 1977).

Paracoccidioides brasiliensis De Almeida

This organism can cause paracoccidioidomycosis, a systemic mycosis which may involve the skin and mucous membranes. An hypersensitivity state has been demonstrated in asymptomatic individuals (Larsh & Goodman 1985, Roberts et al. 1984).

Penicillium Link

Eczematous dermatitis has been attributed to contact with or inhalation of spores of Penicillium species (Bocobo et al. 1954).

Penicillium chrysogenum Thom
[syn. Penicillium notatum Westling]

Cultures of this organism are used to produce natural and semi-synthetic penicillin-type antibiotics (Domsch et al. 1980). Penicillins are well known as potent contact sensitisers, this having been demonstrated in humans and in guinea pigs using a maximisation test. See Cronin (1980) for a more complete review, and Rudzki & Rebendel (1984).

Penicillium citrinum Thom

A crude physiological saline extract of this species elicited positive patch test reactions in 2 of 3 patients with seasonal eczematous dermatitis. A crude ether extract failed to elicit a response (Fujisawa et al. 1966).

Penicillium griseofulvum Dierckx


Penicillium nigricans Bain. ex Thom

These organisms are the source of griseofulvin which is used in the oral treatment of a number of dermatomycoses (Domsch et al. 1980).

Griseofulvin may be an occasional photosensitiser, but clinically this rarely seems to be a problem (Cronin 1980).

Phoma glomerata Wollenw. & Hochapfel

Several cases of pathogenicity ascribable to this organism have been recorded, including granuloma of the foot, mycoses of the hand, otomycosis, etc. (Domsch et al. 1980).

Phoma herbarum Westend.
[syn. Phoma hibernica Grimes, O'Connor, & Cummins]

P. hibernica has been documented as a cause of phaeohyphomycosis involving subcutaneous tissue (McGinnis 1983, 1985). See also Phialophora Medlar.

Pyrenochaeta romeroi Borelli

This organism is known as a cause of "black grain" mycetomas (Roberts et al. 1984, Padhye & Ajello 1985).

Scopulariopsis Bain.

Organisms belonging to this form-genus have been associated with the teleomorphic genus Microascus Zukal, fam. Microascaceae (Domsch et al. 1980).

Binders and vendors of hay, straw, and rushes, those engaged in putting matting around bottles and in seating cane-bottomed chairs may be affected by dermatoconiosis. Prosser White (1934) speculated that Scopulariopsis koningii Vuill. might be responsible; the fungus occurs in litter, straw, and manure.

Scopulariopsis brevicaulis Bain.
[syn. Penicillium brevicaule Sacc.]

This organism may cause deep-seated gummose ulcers and onchomycosis (Domsch et al. 1980, Roberts et al. 1984).

Scopulariopsis fusca Zach

This has been named as the causative organism in cases of onchomycosis and dermatomycosis (Domsch et al. 1980).

Scytalidium Pesante

This genus comprises some 7 species of cosmopolitan distribution.

McGinnis et al. (1985) refer to a dermatomycosis caused by the Scytalidium synanamorph of Hendersonula toruloides Nattrass.

Scytalidium hyalinum C.Campbell & Mulder

This organism, a white mould, has been recognised as a cause of skin and nail infections almost identical with those produced by Trichophyton rubrum Sabouraud (Roberts et al. 1984).

Scytalidium lignicola Pesante

This organism may cause cutaneous phaeohyphomycosis (McGinnis 1983).

Spondylocladium Mart. ex Corda

Cross-sensitivity was observed between castor bean dust (Ricinus communis L., fam. Euphorbiaceae) and an unidentified species of Spondylocladium (Panzani 1962).

Sporothrix schenckii Hektoen & Perkins
[syn. Sporotrichum schenckii de Buerm. & Gougerot]

Although the teleomorph of this organism has been identified as Ophiostoma stenoceras Nannf. (syns Ceratocystis stenoceras C.Moreau, Ceratostomella stenoceras Robak), the continuing discussion (Domsch et al. 1980) about this relationship suggests that it may be premature to consider this to be unequivocal. The organism occurs widely in soil and on vegetable matter.

This fungus is the aetiological agent of a chronic subcutaneous lymphatic, or rarely respiratory, mycosis in man and other mammals which has been named sporotrichosis (Domsch et al. 1980, Roberts et al. 1984). An hypersensitivity state to sporotrichin, an experimental antigen derived from the organism, has been demonstrated in some asymptomatic individuals (Larsh & Goodman 1985, Roberts et al. 1984).

According to Schwartz et al. (1957), the reeds of Provence (Arundo; fam. Gramineae), used to make sieves, trellises, and ceilings, when piled in heaps carry a dry white mould (Sporotrichum [sic]) which produces dermatitis in workers who strip the stalks.

Sporotrichosis may result from puncture wounds caused by barberry thorns (see Berberis vulgaris L., fam. Berberidaceae). The disease should be suspected in all persons with subcutaneous lesions who handle thorny plants, timber, or sphagnum moss (Kaufman & Reiss 1985).

Sporothrix schenkii Hektoen & Perkins var. luriei Ajello & Kaplan

This organism is the causal agent of a cranial osteolysis in South Africa (Domsch et al. 1980).

Stachybotrys chartarum S.Hughes
[syn. Stachybotrys alternans Bonord.]

Mycotoxins found in the mycelium can produce stachybotryotoxicosis in both horses (see VETERINARY ASPECTS) and man (Domsch et al. 1980). Stachybotryotoxicosis is caused by the consumption of hay on which the mould has been growing and is characterised by severe lesions of the nose, mouth, throat, and gastrointestinal tract. Ether extracts of cultures of the mould have dermatitic properties (Bowden & Schantz 1955).

Stenella araguata Syd.
[syns Cladosporium castellani Borelli & Marcano, Cladosporium araguatum Arx]

This organism may cause tinea nigra, a superficial phaeohyphomycosis (McGinnis et al. 1985).

Taeniolella stilbospora Hughes
[syn. Torula stilbospora Corda]

This organism may cause a dermatomycosis (McGinnis 1983).

Trichophyton Malmsten

This is a genus of about 20 species, some of which are commonly found as causative agents of ringworm in humans (Ajello & Padhye 1980). Teleomorphs are as yet unknown for many of the pathogenic species, but those that are known belong to the genus Arthroderma Berk., fam. Arthrodermataceae under which heading they are considered.

Trichophytin, a commercially produced extract from Trichophyton mentagrophytes Blanchard (see Arthroderma benhamiae Ajello & Cheng, fam. Arthrodermataceae) and related dermatophytes, produced positive delayed intracutaneous test reactions in sensitised persons (Ramirez 1930).

Roberts et al. (1984) list the following species (in addition to those described in the monographs below) as being locally important causes of ringworm in man:

Trichophyton gourvilii Catanei
Trichophyton megninii Blanchard
[syn. Megatrichophyton megnini Neveu-Lemaire]
Trichophyton soudanense Joyeux
[syn. Langeronia soudanensis Vanbreuseghem]
Trichophyton yaoundei Cochet & Doby-Dubois (nomen nudum) 

Trichophyton concentricum Blanchard

Tinea imbricata is caused by this organism. This distinctive dermatophyte disease is restricted to Malaysia, the Pacific, the Far East, and parts of Central and South America (Roberts et al. 1984).

Trichophyton equinum Gedoelst

This organism is a common cause of ringworm in horses, but may also invade human skin (Ainsworth & Austwick 1973, Ajello & Padhye 1980).

Trichophyton erinacei A.A.Padhye & Carmichael
[syn. Trichophyton mentagrophytes Blanchard var. erinacei J.M.B. Smith & Marples]

This organism has been reported as a cause of ringworm in hedgehogs, rat, mouse, dog, and man (Ainsworth & Austwick 1973, Roberts et al. 1984).

Trichophyton gallinae Silva & Bentham
[syn. Microsporum gallinae Grigorakis]

Although usually described as a species of Microsporon, Ajello (1968) considers that the organism belongs to the genus Trichophyton. The species is recorded as being pathogenic to man (Ainsworth & Austwick 1974, Roberts et al. 1984).

Trichophyton rubrum Castellani
[syns Trichophyton purpureum Bang, Epidermophyton purpureum C.W. Dodge]

This organism is a common cause of ringworm of the skin and nails (Ajello & Padhye 1980, Roberts et al. 1984). It may also produce a hyperkeratotic form of athlete's foot (Sande & Mandell 1985).

Trichophyton schoenleinii Langeron & Milochevitch
[syn. Achorion schoenleinii Remak.]

Favus, a form of ringworm characterised by heavy cup-shaped crusts (scutulae) and hair invaded throughout its length by hyphae which do not fragment into arthrospores, is produced by this organism (Ainsworth & Austwick 1973, Ajello & Padhye 1980, Roberts et al. 1984).

Trichophyton tonsurans Malmsten

This organism is a common cause of epidemic tinea capitis in both children and adults (Ajello & Padhye 1980, Roberts et al. 1984).

Trichophyton verrucosum Bodin
[syns Trichophyton album Sabouraud, Trichophyton discoides Sabouraud, Trichophyton ochraceum Sabouraud]

This organism is a common cause of ringworm in cattle, but may also invade human skin (Ainsworth & Austwick 1973, Ajello & Padhye 1980).

Trichophyton violaceum Bodin

Tinea capitis in both children and adults may be caused by this organism (Ajello & Padhye 1980). The organism is found in the Mediterranean region in particular (Roberts et al. 1984).

Trichosporiella cerebriformis W. Gams
[syn. Sporotrichum cerebriforme de Vries & Kleine-Natrop]

This soil fungus has been isolated from a seborrhoeic-pityriasis-like lesion of the human scalp (Domsch et al. 1980).

Trichosporon beigelii Vuillemin
[syns Geotrichum beigelii Coudert, Micrococcus beigelii Mig., Pleurococcus beigelii Küchenm. & Rabenh., Sporotrichum beigelii Sacc. & Traverso, Trichosporon aquatile L.R.Hedrick & P.D.Dupont, Trichosporon cutaneum Ota, Trichosporon ovoides Behrend]

This organism, a yeast, is responsible for white piedra, an uncommon infection of the scalp, axillary, and facial hair (Ajello & Padhye 1974, Roberts et al. 1984) and may also occur as an opportunistic pathogen of mucous membranes or skin (Ajello & Padhye 1980, Cooper & Silva-Hutner 1985).

There has been some disagreement in the literature as to whether Trichosporon cutaneum or Trichosporon beigelii should be considered to be the correct name. do Carmo-Sousa (1970) believes Trichosporon cutaneum to be the correct name whereas most medical mycology texts refer to Trichosporon beigelii.

Veronaia audouinii Benedek
[syns Microsporum audouinii Gruby, Microsporum langeronii Vanbreuseghem, Microsporum rivalieri Vanbreuseghem]

This organism was formerly a common agent of epidemic tinea capitis in young children (Ajello & Padhye 1980).

Volutella cinerescens

This organism has been isolated from keratitis lesions (Rebell & Forster 1980).

Zygosporium masonii Hughes

Minato et al. (1973) investigated the cytotoxicity and skin irritancy of a number of natural and semi-synthetic cytochalasins (or zygosporins) derived from the culture filtrate of this fungus. The two properties were found not to be directly related to one another; the most potent irritants amongst the natural cytochalasins were zygosporins A and D.



This small family of ascomycetes comprises 38 species in 3 genera (Hawksworth et al. 1995).

Morchella esculenta St Amans
Common Morel

Industrial workers preserving this edible fungus developed keratoconjunctivitis and respiratory irritation (Pick 1927, Grant 1974).



This family of 122 species in 20 genera is classified in the Zygomycota (Hawksworth et al. 1995).

A systemic zygomycosis known as mucormycosis is known as a rare opportunistic infection in immunosuppressed or diabetic patients. Wounds covered with dressings that are contaminated with these organisms may also become infected. Mucormycosis may be caused by several species of the genera Absidia Tiegh., Mucor Mich. ex St.-Am., and Rhizopus Ehrenb. (Greer & Rogers 1985, Roberts et al. 1984).

Absidia corymbifera Sacc. & Trotter
[syns Absidia cornealis Dodge, Absidia italiana Dodge, Absidia ramosa Lendner, Mucor corymbifer F. Cohn]

This species has been recorded as causing mucormycosis (Ajello et al. 1976).

Mucor Mich. ex St.-Am.

The following species have been recorded as causing mucormycosis (Ajello et al. 1976):

Mucor pusillus Lindt
[syn. Mucor miehei Cooney & Emerson]
Mucor ramosissimus Samsutsewitsch 

Mucor mucedo Mich. ex St.-Am.

Various moulds have been suspected as causes of dermatitis, including Mucor mucedo (Prosser White 1934).

Mucor ramannianus Moller

This organism yields ramycin, an antibiotic that is identical with the better known fusidic acid (Hawksworth et al. 1995). See Fusidium coccineum Fuckel under FUNGI - MITOSPORIC.

Rhizopus Ehrenb.

The following species have been recorded as causing mucormycosis (Ajello et al. 1976):

Rhizopus arrhizus A.Fisch.
Rhizopus microsporus Tiegh.
Rhizopus oryzae Went & Prinsen Geerligs
Rhizopus rhizopodiformis Zopf
[syns Rhizopus chinensis Saito, Rhizopus cohnii Berlese & De Toni, Mucor rhizopodiformis F.Cohn] 

Rhizopus stolonifer Lind.
[syn. Rhizopus nigricans Ehrenb. ex Corda]

Prosser White (1934) lists Rhizopus nigricans as a mould that has been suspected as a cause of dermatitis.

Ajello et al. (1976) reviewed several cases of zygomycosis that had been attributed to this organism but doubted the identifications made, especially since the organism is incapable of growing at 37°C.



This large family of ascomycetes comprises some 13 genera and 634 species that are biotrophic, necrotrophic or saprobic on various plant tissues. They cause a variety of leaf spot, blotch and other diseases in plants (Hawksworth et al. 1995).

Cymadothea trifolii F.A. Wolf
[syns Sphaeria trifollii Pers., Mycosphaerella killianii Petr.]
Anamorph: Polythrincium trifolii Kunze

This organism is the cause of black blotch or sooty blotch disease in clover (Trifolium L. spp., fam. Leguminosae) (Hawksworth et al. 1995).

Clover infected with black blotch disease was incriminated on circumstantial evidence (Ames et al. 1994) as a cause of secondary photosensitisation in horses when it was noticed that a substantial proportion of the clover (red or white) in pastures where the horses had been grazing was infected with this organism. See also Trifolium pratense L., fam. Leguminosae.



Many members of this family have previously been classified in the Hypocreaceae. These organisms are of cosmopolitan distribution; many are plant parasites (Hawksworth et al. 1995).

Gibberella moniliformis Winel
Anamorph: Fusarium moniliforme Sheldon

This organism may cause mycotic ulcers of the cornea (Rebell & Forster 1980) and mycetomas (Padhye & Ajello 1985).

Nectria Fr.

Anamorphs are found in the form-genera Acremonium Link ex Fr., Fusarium Link ex Fr., Verticillium Nees ex Link, Myrothecium Tode ex Fr., and many others (Domsch et al. 1980).

Nectria haematococca Berk. & Br. var. brevicona Gerlach
Anamorph: Fusarium solani Sacc.
[syn. Fusisporium solani Mart.]

This organism is the most important cause of mycotic ulcers of the cornea (Rebell & Forster 1980). It may also cause mycetomas (Padhye & Ajello 1985).

Nectria haematococca Berk. & Br. var. haematococca
[syn. Hypomyces haematococcus Wollenw.]
Anamorph: Fusarium eumartii Carp.

Mycelial extracts of this organism caused toxic reactions on the skin of rabbits (Domsch et al. 1980).

Nectria inventa Pethybr.
Anamorph: Verticillium tenereum Link

Nectria inventa was isolated from a keratomycosis in one of 88 cases (Domsch et al. 1980).



This family, which is classified in the Ascomycota, is quite closely related to the Ajellomycetaceae, the Arthrodermataceae, and to the Gymnoascaceae. It comprises 134 species in 23 genera (Kirk et al. 2008).

Coccidioides immitis G.W.Stiles
[syns Aleurisma immite Bogliolo & J.A.Neves, Blastomycoides immitis Castell., Geotrichum immite A.Agostini, Zymonema immitis Mello]

The vegetative phase of this fungus is highly infective and can cause coccidioidomycosis, a systemic mycosis which may involve the skin and bone as well as the lungs (Domsch et al. 1980). It is endemic to arid regions of North and South America (Ainsworth & Austwick 1973, Roberts et al. 1984). Most patients develop a hypersensitivity state that is readily demonstrated by intradermal coccidioidin skin tests (Larsh & Goodman 1985).

[Further information available but not yet included in database]

Nannizziopsis vriesii Currah
[syns Arachnotheca vriesii Samson, Rollandina vriesii Apinis]
Anamorph: Chrysosporium guarroi M.L.Abarca et al.

[Information available but not yet included in database]



The family is classified in the Ascomycota. It comprises 2 species in 1 genus (Hawksworth et al. 1995).

Piedraia hortae Fonseca & Leāo
Anamorph: Trichosporon hortae Brumpt

This organism is responsible for black piedra, a fungus infection of scalp (and occasionally axillary and pubic) hair which occurs primarily in tropical areas of the world (Ajello & Padye 1980, Roberts et al. 1984).



The family is classified in the Ascomycota. It comprises about 149 species in 15 genera, and includes organisms previously classified in the Pyrenophoraceae (Hawksworth et al. 1995).

Cochliobolus Drechsler

The anamorphic form-genus Curvularia Boedijn, which comprises some 35 species of mostly tropical and subtropical facultative plant parasites, has been identified with the teleomorphic genus Cochliobolus (Domsch et al. 1980). See also Curvularia Boedijn under FUNGI - MITOSPORIC.

Cochliobolus geniculatus R. Nelson
[syn. Pseudocochliobolus geniculatus Tsuda et al.]
Anamorph: Curvularia geniculata Boedijn
[syn. Helminthosporium geniculatum Tracy & Earle]


Cochliobolus lunatus R. Nelson & Haasis
[syn. Pseudocochliobolus lunatus Tsuda et al]
Anamorph: Curvularia lunata Boedijn
[syn. Helminthosporium curvulum Sacc.]

These two species have been reported to cause mycetomas. Curvularia geniculata has affected dogs in the USA whilst Curvularia lunata has affected humans in Senegal and Sudan (Padhye & Ajello 1985).

Cochliobolus spicifera R. Nelson
Anamorph: Curvularia spicifera Boedijn
[syns Drechslera spicifera v. Arx, Helminthosporium spiciferum Nicot, Brachycladium spiciferum Bainier]

McGinnis (1985) notes that this species is a recognised pathogen.

Leptosphaerulina chartarum Cec.Roux
Anamorph: Pithomyces chartarum M.B.Ellis
[syns Piricauda chartarum R.T.Moore, Sporidesmium bakeri Syd. & P.Syd., Sporidesmium chartarum Berk. & M.A.Curtis]

Exposure to the spores of, and ingestion of grass contaminated with the mitosporic fungus Pithomyces chartarum causes facial eczema in sheep and cattle, photosensitisation of the skin, liver lesions, and a raised phylloerythrin concentration in the blood. These toxic effects are attributable to the sporidesmins produced by the organism (Russell 1960, Leach & Tulloch 1971, Ainsworth & Austwick 1973, Domsch et al. 1980).

[Further information available but not yet included in database]



This family of "rust" fungi is classified in the order Uredinales of the Basidiomycota. Rusts are of great importance as pathogens of higher plants, especially grasses and cereals.

Endophyllum kaernbachii F.Stevens & Mendiola
Anamorph: Aecidium kaernbachii Henn.
[syn. Aecidium rechingeri Bubak]

McKenzie & Gerlach (1988) described a bullous skin reaction associated with an erythematous rash and a burning sensation after handling beach morning glory (Ipomoea pes-caprae R.Br. subsp. brasiliensis van Ooststr., fam. Convolvulaceae) infected with this rust fungus. Hyperpigmentation of the affected skin persisted for 15-18 months. The reaction was believed to be an example of a parasitophytophotodermatitis.

Puccinia Pers. ex Pers.

Smuts and rusts have been suspected of producing skin irritation, stomatitis, and inflammation of mucous membranes (Pammel 1911, Steyn 1934, Hurst 1942). Pammel (1911) noted that considerable irritation of the nose and throat is experienced when cereals infected with rusts are threshed. See also Ustilago Roussel, fam. Ustilaginaceae and Coleosporium Lév., fam. Coleosporiaceae.

Puccinia acetosa Barclay
[syn. Uromyces apiculatus F. Strauss in Fries]

Steyn (1934), citing Fröhner (1919), listed Uromyces apiculatus as being capable of causing inflammation of the skin.

Puccinia arundinacea Hedw.

Steyn (1934), citing Fröhner (1919), listed this species as being capable of causing inflammation of the skin.

Puccinia coronata Corda
Crown Rust, Oat Rust

Steyn (1934), citing Fröhner (1919), listed this species, which infects oats (Avena sativa L., fam. Gramineae) as being capable of causing inflammation of the skin.

Puccinia graminis Pers.
Wheat Rust, Black Rust, Stem Rust

The fungus can attack several species of grasses and cereals, including wheat (Triticum L., fam. Gramineae). Wheat rust has been listed as a possible cause of dermatitis (Steyn 1934 citing Fröhner (1919), Schwartz et al. 1957) as well as asthma (Cadham 1924).

Puccinia straminis Fuckel

Steyn (1934), citing Fröhner (1919), listed this species as being capable of causing inflammation of the skin.



This is a family of pseudofungi classified in the Oomycota. It comprises 183 species in 10 genera (Hawksworth et al. 1995).

Pythium insidiosum De Cock, L.Mend., A.A.Padhye, Ajello & Kaufman

[Information available but not yet included in database]



Lactarius deliciosus S.F. Gray
Saffron Milk Cap, Orange Agaric

A case of cutaneous sensitisation to the edible mushrooms Boletus luteus L. ex Fr., fam. Boletaceae; Lactarius deliciosus Fr., and Ramaria flava Quél., fam. Gomphaceae was reported. Hypersensitivity was also demonstrated after eating the mushrooms in question, fried. Boiling seems to destroy the antigenic effect of mushrooms. From a practical point of view, it is important to be familiar with the cases, as for example in differentiating between similar eruptions caused by sunlight or hypersensitivity to autumn flowers (Hellerström 1941).



This family of some 159 species of yeast fungi in 28 genera is classified in the Ascomycota (Hawksworth et al. 1995). The family includes the commercially important fermentative yeasts used by brewers, bakers, and wine-makers.

Meyerozyma guilliermondii Kurtzman & M. Suzuki
[syn. Pichia guilliermondii Wickerham]
Anamorph: Candida guilliermondii Langeron & Guerra
[syn. Endomyces guilliermondii Castellani]

This organism may be found as an opportunistic pathogen in man (Roberts et al. 1984).

Saccharomyces cerevisiae Meyen ex E. Hansen
Baker's Yeast

This yeast, which is used in baking, brewing, and wine making, has been reported to cause dermatitis. Brewery workers who scrape off masses of yeasts adhering to fermentation casks with their finger-nails develop crusted excrescences beneath the nails and destruction of the nail-plates (Prosser White 1934).

This organism is responsible for occasional cases of thrush and vulvovaginitis (Cooper & Silva-Hutner 1985).



This family comprises just 1 species in 1 genus classified in the Zygomycota (Hawksworth et al. 1995).

Saksenaea vasiformis Saksena

Ajello et al. (1976) described a case of zygomycosis caused by this organism in cranial and facial wounds sustained in an automobile accident.



This family comprises some 42 species of lignicolous fungi in 4 genera (Hawksworth et al. 1995).

Schizophyllum commune Fr.

This organism may cause a subcutaneous infection described as a hyalohyphomycosis (McGinnis et al. 1985). Kligman (1950) investigated a case of onychomycosis associated with an itching rash involving the large toes of both feet in a 30 year old male and concluded that it was probably caused by S. commune.



This family, which has been divided into 33 genera and 225 species (Hawksworth et al. 1995), is classified in the Ascomycota. It is an important family from an economic point of view since its members are responsible for much damage to fruit, vegetables, and ornamental plants.

The grey mould Botrytis cinerea Pers. ex Nocca & Balb. is a particularly well known organism which is in fact a form-species complex connected to several teleomorphs classified in the Sclerotiniaceae. These include Botryotinia fuckeliana Whetzel, Botryotinia convoluta Whetzel, Sclerotinia draytonii Buddin & Wakef., Botryotinia pelargonii Røed, and others (Domsch et al. 1980).

Sclerotinia sclerotiorum de Bary
[syn. Peziza sclerotiorum Lib.]
Pink Rot

Celery (Apium graveolens L., fam. Umbelliferae) infected with this fungus contains the phototoxic furocoumarins bergapten and xanthotoxin, and can therefore elicit photodermatitis (see Umbelliferae). Austad & Kavli (1983) reported the occurrence of phototoxic contact dermatitis of the hands and forearms of 11 celery harvesters in Norway. The conditions favouring an epidemic include diseased celery, a sunny day following wet weather, and unprotected harvesters.



This family of ascomycetes comprises 6 species in 5 genera (Hawksworth et al. 1995). All members of this family have previously been classified in the Zopfiaceae.

Neotestudina rosatii Segretain & Destombes
[syn. Zopfia rosatii D.Hawksworth & C.Booth]

This organism was originally isolated from severe "white grain" mycetomas in man in Somalia, other African countries, and in Australia (Domsch et al. 1980, Padhye & Ajello 1985).



This family of ascomycetes comprises 170 species in 26 genera (Hawksworth et al. 1995).

Emericella nidulans Vuillemin
Anamorph: Aspergillus nidulans Winter

Cases of "white grain" mycetomas caused by this organism have been described from Senegal, Sudan, and Tunisia (Roberts et al. 1984, Padhye & Ajello 1985). Onchomycosis and mandibular periostitis associated with this organism have also been described (Domsch et al. 1980).



This family of ascomycetes comprises 65 species in 11 genera (Kirk et al. 2008).

Khuskia oryzae H.J.Huds.
Anamorph: Nigrospora sphaerica E.W.Mason
[syns Monotospora oryzae Berk. & Broome, Nigrospora gossypii Jacz., Nigrospora oryzae Petch, Trichosporum sphaericum Sacc.]

[Information available but not yet included in database]



This family of "smut" fungi is classified in the Basidiomycota. It includes a number of important crop plant pathogens.

Filobasidiella neoformans Kwon-Chung
[syn. Saccharomyces neoformans Sanfelice]
Anamorph: Cryptococcus neoformans Vuillemin
[syn. Torulopsis neoformans Redaelli]

This organism is best known as its anamorphic form, the teleomorph having only recently (Kwon-Chung 1976) been described. It lives naturally on soil contaminated with bird droppings. The organism can produce a diffuse pulmonary infection if inhaled, leading to meningitis and followed by localised abscesses or granulomas (cryptococcoma or toruloma) in the lungs, lymph nodes, bones, and skin (Cooper & Silva-Hutner 1985). The disease is known as cryptococcosis. Skin involvement may occur in 10-15% of cases (Roberts et al. 1984). An hypersensitivity state has been demonstrated in some individuals suffering from the disease (Larsh & Goodman 1985).

Tranzscheliella hypodytes Vánky & McKenzie
[syns Caeoma hypodytes Schltdl., Cintractia hypodytes Maire, Uredo hypodytes Pers., Ustilago hypodytes Fr.]

Ustilago hypodytes, a common cause of stem smut in grasses, was incriminated in "la maladie des cannes" — reed dermatitis (Arundo donax L., fam. Gramineae); positive epicutaneous tests to the fungus were observed in rabbits (Gerbaud 1885). This represents an early use of the epicutaneous test.

Ustilago Roussel

The genus Ustilago is one of a number of genera of fungi known as smuts which are classified in the order Ustilaginales. Two families of smut fungi are recognised, namely Tilletiaceae and Ustilaginaceae (Talbot 1971).

A disorder resembling acrodynia occurs following ingestion of the corn-smut Ustilago (Watt & Breyer-Brandwijk 1962).

Smuts and rusts have been suspected of producing skin irritation, stomatitis, and inflammation of mucous membranes (Hurst 1942). See also Coleosporium Lév., fam. Coleosporiaceae and Puccinia Pers. ex Pers., fam. Pucciniaceae. Irritant properties have been ascribed to the following species (Pammel 1911, Barjaktarović & Bogdanović 1933, Preininger 1937/38):

Ustilago avenae Rostrup
[syn. Ustilago avenae Jensen]
Ustilago jensenii Rostrup
[syn. Ustilago hordei Lagerh.]
Ustilago neglecta Neissl
Ustilago nuda Rostrup
[syns Ustilago nuda Kell. & Swingle, Ustilago tritici Rostrup, Ustilago tritici Jensen]
Ustilago utriculosa Tul.
Ustilago zeae Unger
[syn. Ustilago maydis Corda] 

Ustilago bromivora A. Fisch. v. Waldh.

This smut, which frequently affects the prairie grass Bromus unioloides Kunth (syn. Bromus catharticus Vahl), has been reported to cause severe hay fever and to raise wheals on any part of the body that may be scratched. One smut body in a glass of water produced a solution that gave rise to large wheals when applied to broken skin (Cleland 1943).


The airborne spores of a number of moulds are known to have a causal role in many cases of atopic and seasonal eczemas, rhinitis, and asthma. Most commonly implicated are species of Alternaria Nees ex Fr., Aspergillus Mich. ex Fr., and Penicillium Link ex Fr. (Feinberg 1939, Tuft et al. 1950, Bocobo et al. 1954, Gomez Orbaneja & Quiñones Caravia 1953, Jillson & Adami 1955, Storck 1955, Jillson 1957, Strauss & Kligman 1957, Prince et al. 1960, Ofuji et al. 1961, Rajka 1963, Watanabe & Fujisawa 1965, Fujisawa et al. 1966). Crude extracts of moulds produced eczematous changes by patch test and also by inhalation. Type I hypersensitivity reactions can occur from inhalation of fungi (Bruce 1963).


As in man, fungi may affect animals either by contact (including subcutaneous inoculation), by ingestion, or by inhalation. Many of the pathogenic fungi can produce the same diseases in both humans and livestock. Interested readers are referred to specialised texts on the topic such as that produced by Ainsworth & Austwick (1973).

Perhaps most commonly encountered are ringworm infections which are caused by members of the form-genera Microsporum Gruby, Trichophyton Malmst., and Epidermophyton Sabouraud. Consumption of hay contaminated with the mitosporic fungus Stachybotrys chartarum Hughes (syn. Stachybotrys alternans Bonord.) produces a disease in horses known as stachybotryotoxicosis. It is characterised by ulceration in the nose, mouth, throat, and gastrointestinal tract (Brian et al. 1947, Domsch et al. 1980).

Pammel (1911) described the production of inflammation of skin and mucous membranes in animals after contact with the "rape fungus" (Alternaria exitiosa Jørstad, syn. Sporidesmium exitiosum Kühn) which may infect brassicas (fam. Cruciferae).

Sporotrichosis (see Sporothrix schenkii Hektoen & Perkins under FUNGI - MITOSPORIC), histoplasmosis (see Ajellomyces capsulatus McGinnis & Katz, fam. Ajellomycetaceae), blastomycosis (see Ajellomyces dermatitidis McDonough & Lewis, fam. Ajellomycetaceae), candidiasis (see Candida albicans Berkhout under FUNGI - MITOSPORIC) and other fungal diseases known in man are also found in livestock. Diseases more typically associated with animals include the respiratory diseases adiaspiromycosis caused by Emmonsia parva Ciferri & Montemartini (syn. Haplosporangium parvum Emmons & Ashburn) and Emmonsia crescens Emmons & Jellison; rhinosporidiosis caused by Rhinosporidium seeberi Seeber; and equine nasal granuloma caused by Conidiobolus coronatus Batko, fam. Entomophthoraceae. Miscellaneous fungal infections of animals include hyphomycosis produced by Hyphomyces destruens, a fungus of uncertain taxonomic position (Ainsworth & Austwick 1973).

A number of organisms are capable of producing diseases such as mycotic mastitis and mycotic abortion (Ainsworth & Austwick 1973).


  • Abou-Gabal M and Malik G (1978) The role of Aspergillus fumigatus Fresenius in dermatitis of chickens. Mykosen 21(8): 271-276
  • Ainsworth GC and Austwick PKC (1973) Fungal Diseases of Animals, 2nd edn. Farnham Royal, England: Commonwealth Agricultural Bureaux.
  • Ajello (1968)
  • Ajello L and Padhye A (1980) Dermatophytes and the agents of superficial mycoses. In: Lennette EH et al. (Eds) Manual of Clinical Microbiology, 3rd edn. pp. 541. Washington, DC: American Society for Microbiology.
  • Ajello L, Georg LK, Steigbigel RT and Wang CJK (1974) A case of phaeohyphomycosis caused by a new species of Phialophora. Mycologia 66(3): 490-498.
  • Ajello L, Dean DF and Irwin RS (1976) The zygomycete Saksenaea vasiformis as a pathogen of humans with a critical review of the etiology of zygomycosis. Mycologia 68(1): 52-62.
  • Allen T (1945) Some observations on the relationship of Alternaria tenuis to canine eczema. Journal of the American Veterinary Medical Association 106(816): 163-164
  • Ames T, Angelos J, Gould S, Kullot D, Jergens T, Voller K, Moberg R, Ruth G, O'leary T, Murphy M (1994) Secondary photosensitization in horses eating Cymodothea trifolli infested clover. 37th Annual Meeting, American Association of Veterinary Laboratory Diagnosticians, Abstracts. Grand Rapids, Michigan. p. 45
  • Austad J and Kavli G (1983) Phototoxic dermatitis caused by celery infected by Sclerotinia sclerotiorum. Contact Dermatitis 9: 448-451.
  • Austwick PKC (1966) The role of spores in the allergies and mycoses of man and animals. In: Madelin MF (Ed.) The Fungus Spore. pp. 321. London: Butterworths.
  • Austwick PKC and Longbottom JL (1980) Medically important Aspergillus species. In: Lennette EH, Balows A, Hausler WJ and Truant JP (Eds) Manual of Clinical Microbiology, 3rd edn. pp. 620-xxx. Washington, DC: American Society for Microbiology
  • Barjaktarović SS, Bogdanović SB (1933) Untersuchungen über die Wirkung des Maisbrandes (Ustilago maidis). [Studies on the effect of the maize ergot (Ustilago maidis)]. Archiv für Experimentelle Pathologie und Pharmakologie 173(1): 381-387 [doi] [url]
  • Benham RW (1947) Biology of Pityrosporum ovale. In: Nickerson WJ (Ed.) Biology of Pathogenic Fungi. p. 63. Waltham, MA: Chronica Botanica
  • Bocobo FC, Curtis AC, Block WD and Stubbart FJ (1954) Studies on fungi encountered in the atmosphere. II. Production of dermatitis in guinea pigs by crude ether-soluble extracts of Alternaria, Hormodendrum, Penicillium and Aspergillus. Journal of Investigative Dermatology 23(6): 489-496.
  • Bowden JP, Schantz EJ (1955) The isolation and characterization of dermatitic compounds produced by Myrothecium verrucaria. Journal of Biological Chemistry 214(1): 365-372 [url] [pmid]
  • Brian PW, Curtis PJ, Hemming HG (1947) Glutinosin: a fungistatic metabolic product of the mould Metarrhizium glutinosum S. Pope. Proceedings of the Royal Society of London. Series B, Biological Sciences 135(878): 106-132 [doi] [doi-2] [url] [url-2]
  • Brian PW, Hemming HG and Jefferys EG (1948) Production of antibiotics by species of Myrothecium. Mycologia 40(3): 363-368.
  • Bruce RA (1963) Bronchial and skin sensitivity in asthma. International Archives of Allergy 22: 294.
  • Cadham FT (1924) Asthma due to grain rusts. Journal of the American Medical Association 83: 27.
  • Campbell & White (1989)
  • Campbell CK et al. (1973) Fungal infection of skin and nails by Hendersonula toruloidea. British Journal of Dermatology 89: 45.
  • Castellani A and Chalmers AJ (1919) Manual of Tropical Medicine. 3rd edn. London: Baillière, Tindall and Cox
  • Chilton WS (1978) Chemistry and mode of action of mushroom toxins. In: Rumack BH and Salzman E (Eds) Mushroom Poisoning: Diagnosis and Treatment. pp. 87. Florida: CRC Press, Inc.
  • Cleland JB (1943) Plants, including fungi, poisonous or otherwise injurious to man in Australia. IV. Medical Journal of Australia ii: 161.
  • Cooper BH (1985) Taxonomy, classification, and nomenclature of fungi. In: Lennette EH, Balows A, Hausler WJ and Shadomy HJ (Eds) Manual of Clinical Microbiology, 4th edn. pp. 495-499. Washington, DC: American Society for Microbiology
  • Cooper BH and Silva-Hutner M (1985) Yeasts of medical importance. In: Lennette EH, Balows A, Hausler WJ and Shadomy HJ (Eds) Manual of Clinical Microbiology, 4th edn. pp. 526-541. Washington, DC: American Society for Microbiology
  • Cronin E (1980) Contact Dermatitis. Edinburgh: Churchill Livingstone [WorldCat]
  • Deighton FC and Mulder JL (1977) Mycocentrospora acerina as a human pathogen. Trans. Br. Mycol. Soc. 69: 326.
  • do Carmo-Sousa L (1970) Genus 11. TRICHOSPORON Behrend. In: Lodder L (Ed.) The Yeasts. A taxonomic study, 2nd edn. pp. 1309. Amsterdam: North-Holland Publishing Co.
  • Domsch KH et al. (1980) Compendium of Soil Fungi. Vol. 1 & 2. London: Academic Press.
  • Emmons CW, Joe L-K, Tjoei Eng N-I, Pohan A, Kertopati S and van der Meulen A (1957) Basidiobolus and Cercospora from human infections. Mycologia 49(1): 1-10.
  • Feinberg SM (1939) Seasonal atopic dermatitis. The role of inhalant atopens. Archives of Dermatology and Syphilology 40: 200.
  • Frankland AW and Hay MJ (1951) Dry rot as a cause of allergic complaints. Acta Allerg. IV: 186. Cited by Rajka (1963)
  • Fröhner E (1919) Lehrbuch der Toxicologie. Stuttgart: Verlag von Ferdinand Enke
  • Fujisawa S et al. (1966) Eczematous dermatitis produced by air borne molds. Archives of Dermatology 94: 413.
  • Gerbaud (1885) La maladie des cannes. Dermatose des ouvriers cannissiers (observations et expériences) [Cane-induced disease. Dermatitis in cane workers (observations and experiences)]. Annales de Dermatologie et de Syphiligraphie, Deuxième Série 6(11): 678-680
  • Gomez Orbaneja J, Quiñones Caravia PA (1953) La alergia a inhalantes (hongos) en el eczema de las manos; tipos clinicos. [Allergy to inhalants (fungi) in eczema of the hands; clinical types]. Actas Dermo-Sifiliográficas 44(8): 610-619
  • Grant WM (1974) Toxicology of the Eye. Drugs, chemicals, plants, venoms, 2nd edn. Springfield, IL: Charles C Thomas [WorldCat]
  • Greer DL and Rogers AL (1985) Agents of zygomycosis (phycomycosis). In: Lennette EH, Balows A, Hausler WJ and Shadomy HJ (Eds) Manual of Clinical Microbiology, 4th edn. pp. 575-583. Washington, DC: American Society for Microbiology
  • Greer DL and Friedman L (1966) Studies on the genus Basidiobolus with reclassification of the species pathogenic to man. Sabouraudia 4: 231.
  • Grimes GL (1978) Principles of mushroom identification. In: Rumack BH and Salzman E (Eds) Mushroom Poisoning: Diagnosis and Treatment. pp. 3. Florida: CRC Press Inc.
  • Hammerschmidt DE (1980) Szechwan purpura. New England Journal of Medicine 302: 1191.
  • Hawksworth DL, Kirk PM, Sutton BC, Pegler DN (1995) Ainsworth & Bisby's Dictionary of the Fungi, 8th edn. Wallingford: CAB International
  • Hellerström S (1941) Sensitization to edible mushrooms. Acta Dermato-Venereologica 22: 331.
  • Herstoff JK et al. (1978) Rhinophycomycosis entomophthorae. Archives of Dermatology 114: 1674.
  • Holmes S (1983) Outline of Plant Classification. London: Longman Group Ltd.
  • Hopkins HH (1952) Mushroom dermatitis. Md St. Med. J. 1: 504.
  • Hopkins HH (1953) Mushroom dermatitis. Report of a case. AMA Archives of Dermatology and Syphilology 67: 632.
  • Hurst E (1942) The Poison Plants of New South Wales. Sydney: NSW Poison Plants Committee [WorldCat] [url]
  • Jillson OF (1957) Allergic dermatitis produced by pathogenic and saprophytic fungi. Annals of Allergy 15: 14.
  • Jillson OF and Adami M (1955) Allergic dermatitis produced by inhalant molds. AMA Archives of Dermatology 72: 411.
  • Joseph R, Vialatte J, Job JC, Ribierre M, Brunet D, Jean N (1965) A propos de deux observations d'allergie a Candida albicans chez l'enfant, associant a des manifestations articulaires des troubles respiratoires, digestifs et cutanés. [Two cases of allergy to Candida albicans in children, associated with joint manifestations, respiratory, digestive and cutaneous symptoms]. Revue Française d'Allergie 5(1): 37-39 [doi] [url] [pmid]
  • Kaufman L and Reiss E (1985) Serodiagnosis of fungal diseases. In: Lennette EH, Balows A, Hausler WJ and Shadomy HJ (Eds) Manual of Clinical Microbiology, 4th edn. pp. 924-944. Washington, DC: American Society for Microbiology
  • Kirk PM, Cannon PF, Minter DW, Stalpers JA (2008) Ainsworth & Bisby's Dictionary of the Fungi, 10th edn. Wallingford: CAB International
  • Kligman AM (1950) A basidiomycete probably causing onychomycosis. Journal of Investigative Dermatology 14(1): 67-70.
  • Kwon-Chung KJ (1976) A new species of Filobasidiella, the sexual state of Cryptococcus neoformans B and C serotypes. Mycologia 68(4): 942-946.
  • Larsh HW and Goodman NL (1985) Fungi of systemic mycoses. In: Lennette EH, Balows A, Hausler WJ and Shadomy HJ (Eds) Manual of Clinical Microbiology, 4th edn. pp. 542-553. Washington, DC: American Society for Microbiology
  • Laverde S et al. (1973) Mycotic keratitis; 5 cases caused by unusual fungi. Sabouraudia 11: 119.
  • Leach CM and Tulloch M (1971) Pithomyces chartarum, a mycotoxin-producing fungus, isolated from seed and fruit in Oregon. Mycologia 63(5): 1086-1089.
  • Leonardi G (1954) Spore di lichene boschivo (Palmeria caperata) causa di allergosi cutanea. Folia Allerg. 1: 219.
  • Mackinnon JE et al. (1949) Investigaciones sobre las maduromicosis y sus agentes. An. Fac. Med. Univ. Montevideo 34: 231. Cited by Neuhauser (1955)
  • Mandoul R et al. (1954) La maladie de la canne de Provence en Algérie. Bull. Soc. Path. Exot. 47: 572.
  • Marriott D, Kwong T, Harkness J, Ellis D (2006) Cyclomyces tabacinus as a cause of deep tissue infection: the first case report. Mycoses 49(2): 147-149 [doi] [url] [url-2] [pmid]
  • Martin H (Ed.) (1969) Insecticide and Fungicide Handbook, 3rd edn. Oxford: Blackwell Scientific.
  • McCombs RP (1972) Diseases due to immunologic reactions in the lungs. New England Journal of Medicine 286: 1186 & 1245.
  • McGinnis MR (1978) Human pathogenic species of Exophiala, Phialophora, and Wangiella. Scient. Publs Pan-Am. Hlth Org. 356: 37.
  • McGinnis MR (1979) Taxonomy of Exophiala werneckii and its relationship to Microsporum mansonii. Sabouraudia 17: 145.
  • McGinnis MR (1983) Chromoblastomycosis and phaeohyphomycosis: new concepts, diagnosis, and mycology. Journal of the American Academy of Dermatology 8: 1.
  • McGinnis MR (1985) Dematiaceous fungi. In: Lennette EH, Balows A, Hausler WJ and Shadomy HJ (Eds) Manual of Clinical Microbiology, 4th edn. pp. 561-574. Washington, DC: American Society for Microbiology
  • McGinnis MR et al. (1985) Mycotic diseases. A proposed nomenclature. Int. J. Derm. 24: 9.
  • McKenzie EHC, Gerlach WWP (1988) Morning-glory, parasitophytophotodermatitis and Gods. Mycologist 2(4): 158-159
  • Mehregan AH, Rudner EJ (1980) Implantation dermatosis: wood splinter with fungus contamination. Journal of Cutaneous Pathology 7(5): 330-331 [doi] [url] [url-2] [pmid]
  • Minato H, Katayama T, Matsumoto M, Katagiri K, Matsuura S, Sunagawa N, Hori K, Harada M, Takeuchi M (1973) Structure-activity relationships among zygosporin derivatives. Chemical and Pharmaceutical Bulletin 21(10): 2268-2277
  • North PM (1967) Poisonous Plants and Fungi in Colour. London: Blandford Press [WorldCat]
  • Ofuji S et al. (1961) Studies on the role of airborne fungi in several skin diseases. 2. Studies on the seasonal variations of the mould counts in the Kyoto area. Acta Derm., Kyoto 56: 205. Cited by Fujisawa et al. (1966)
  • Padhye AA and Ajello L (1985) Fungi causing eumycotic mycetomas. In: Lennette EH, Balows A, Hausler WJ and Shadomy HJ (Eds) Manual of Clinical Microbiology, 4th edn. pp. 554-560. Washington, DC: American Society for Microbiology
  • Paldrok H and Hollström E (1952) Onychomycosis due to Aspergillus terreus Thom. Acta Dermato-Venereologica 32(Suppl. 29): 255-260
  • Pammel LH (1911) A Manual of Poisonous Plants. Chiefly of North America, with Brief Notes on Economic and Medicinal Plants, and Numerous Illustrations. Cedar Rapids, IA: Torch Press [WorldCat] [url] [url-2]
  • Panzani R (1962) Étude de l'allergie entre la graine de ricin et spondylocladium. Int. Archs Allergy Appl. Immun. 21: 288.
  • Pick L (1927) Augen- und Schleimhauterkrankungen durch Morchelausdunstungen (gewerbliche Massenerkrankung). Z. Augenheilk. 61: 325.
  • Preininger T (1937/8) Durch Maisbrand (Ustilago maydis) bedingte Dermatomykose. Arch. Derm. Syph. 176: 109.
  • Prince HE et al. (1960) The importance of inhalant mold sensitivity in atopic dermatitis. Annals of Allergy 18: 186.
  • Prosser White R (1934) The Dermatergoses or Occupational Affections of the Skin, 4th edn. London: HK Lewis [doi] [WorldCat] [url] [url-2]
  • Rajka G (1963) Studies in hypersensitivity to molds and staphylococci in prurigo Besnier (atopic dermatitis). Acta Dermato-Venereologica 43(Suppl. 54): 1-139.
  • Ramirez MA (1930) Trichophytin sensitization; case report. Med. J. Rec. 132: 382.
  • Rebell GC and Forster RK (1980) Fungi of keratomycosis. In: Lennette EH, Balows A, Hausler WJ and Truant JP (Eds) Manual of Clinical Microbiology, 3rd edn. pp. 553-561. Washington, DC: American Society for Microbiology
  • Reynolds JEF (Ed.) (1996) Martindale. The Extra Pharmacopoeia. 31st edn. London: Royal Pharmaceutical Society [WorldCat]
  • Robbers JE, Speedie MK and Tyler VE (1996) Pharmacognosy and Pharmacobiotechnology. Baltimore: Williams & Wilkins.
  • Roberts SOB, Hay RJ and Mackenzie DWR (1984) A Clinician's Guide to Fungal Disease. New York: Marcel Dekker, Inc.
  • Rogers AL (1980) Opportunistic and contaminating saprophytic fungi. In: Lennette EH, Balows A, Hausler WJ and Truant JP (Eds) Manual of Clinical Microbiology, 3rd edn. pp. 654-668. Washington, DC: American Society for Microbiology
  • Romaguera C and Grimalt F (1985) Contact dermatitis to sodium fusidate. Contact Dermatitis 12: 176.
  • Rudzki E and Rebendel P (1984) Contact sensitivity to antibiotics. Contact Dermatitis 11: 41.
  • Russ C (1923) A skin eruption due to a mould. Lancet i: 77.
  • Russell DW (1960) Sporidesmolide I, a metabolic product of Sporidesmium bakeri Syd. Biochimica et Biophysica Acta 45: 411-412 [doi] [pmid]
  • Salkin IF and Gordon MA (1977) Polymorphism of Malassezia furfur. Can. J. Microbiol. 23: 471.
  • Sande MA and Mandell GL (1985) Antimicrobial agents. Antifungal and antiviral agents. In: Gilman AG, Goodman LS, Rall TW and Murad F (Eds) Goodman and Gilman's The Pharmacological Basis of Therapeutics, 7th edn. pp. 1219-1239. New York: Macmillan Publishing Company.
  • Schwartz L, Tulipan L, Birmingham DJ (1957) Irritant plants and woods. In: Occupational Diseases of the Skin. 3rd edn, pp. 636-672. London: Henry Kimpton [doi] [WorldCat] [url] [url-2]
  • Seaton A and Morgan WKC (1984) Hypersensitivity pneumonitis. In: Morgan WKC and Seaton A (Eds) Occupational Lung Diseases, 2nd edn. pp. 564-608. Philadelphia: WB Saunders Co.
  • Shelley WB and Florence R (1961) Chronic urticaria due to mold hypersensitivity. A study in cross-sensitization and autoerythrocyte sensitization. Archives of Dermatology 83: 549.
  • Steyn DG (1934) The Toxicology of Plants in South Africa together with a consideration of poisonous foodstuffs and fungi. South Africa: Central News Agency
  • Stockley IH (1985) Disulfiram-type reaction with some cephalosporins. Pharm. J. 234: 239.
  • Storck H (1955) Ekzem durch Inhalation. Schweiz. Med. Wschr. 85: 608.
  • Strauss JS and Kligman AM (1957) The relationship of atopic allergy and dermatitis. AMA Archives of Dermatology 75: 806.
  • Takayasu SM, Akagi M and Shimizu Y (1977) Cutaneous mycosis caused by Paecilomyces lilacinus. Archives of Dermatology 113: 1687-1690
  • Talbot PHB (1971) Principles of Fungal Taxonomy. London: Macmillan Press Ltd.
  • Tio TH et al. (1966) Subcutaneous phycomycosis. Archives of Dermatology 93: 550.
  • Towey JW et al. (1932) Severe bronchial asthma apparently due to fungus spores found in maple bark. Journal of the American Medical Association 99: 453.
  • Trease GE, Evans WC (1966) A Textbook of Pharmacognosy, 9th edn. London: Baillière, Tindall and Cassell [WorldCat]
  • Tschen JA, Knox JM, McGavran MH and Duncan C (1984) Chromomycosis. The association of fungal elements and wood splinters. Archives of Dermatology 120: 107-108.
  • Tuft L (1975) Contact urticaria from cephalosporins. Archives of Dermatology 111: 1609.
  • Tuft L et al. (1950) Atopic dermatitis I. An experimental clinical study of the role of inhalant allergens. Journal of Allergy 21: 181.
  • Vetter J (1993) Selenium content of some higher fungi. Acta Alimentaria 22(4): 383-387
  • Vollum DI (1977) Chromomycosis: a review. British Journal of Dermatology 96: 454.
  • Watanabe S and Fujisawa S (1965) Eczematous hypersensitivity to molds. J. Med. Mycol. 6: 276.
  • Watt JM, Breyer-Brandwijk MG (1962) The Medicinal and Poisonous Plants of Southern and Eastern Africa. Being an account of their medicinal and other uses, chemical composition, pharmacological effects and toxicology in man and animal, 2nd edn. Edinburgh: E & S Livingstone [doi] [WorldCat] [url] [url-2]
  • White JC (1887) Dermatitis Venenata: an account of the action of external irritants upon the skin. Boston: Cupples and Hurd [doi] [WorldCat] [url] [url-2]
  • White WL and Downing MH (1947) The identity of "Metarrhizium glutinosum". Mycologia 39(5): 546-555.
  • Wren RC (1975) Potter's New Cyclopaedia of Botanical Drugs and Preparations. (Re-edited and enlarged by Wren RW). Bradford, Devon: Health Science Press [WorldCat] [doi] [url] [url-2]

Richard J. Schmidt

[Valid HTML 4.01!]

[2D-QR coded email address]
[2D-QR coded url]