A revision of Chrysosporium and allied genera

C.A.N. van Oorschot
Centraalbureau voor Schimmelcultures, Baarn


Summary

       Chrysosporium Corda and those genera formerly placed in synonymy with Chrysosporium are revised, with the addition of Trichosporiella Kamyschko ex W. Gams & Domsch. The monograph provides keys to descriptions and illustrations of the taxa now recognised in Chrysosporium (22 species), Myceliophthora Cost. (8 species), Emmonsia Cif. & Montemartini (2 varieties), Zymonema de Beurmann & Gougerot (1 species), Trichosporiella (2 species) and Geomyces Traaen (3 varieties). The thallic and blastic modes of conidiogenesis and the rhexolytic dehiscence of conidia are considered. The relation to other hyphomycete genera, particularly the macroconidial genera Trichophyton Malmsten and Microsporum Gruby, are discussed. The known teleomorphs associated with the described species, belonging to the Gymnoascaceae, Onygenaceae, Ascosphaeraceae and Sordariaceae are listed. Hyphomycete genera which have been confused with those dealt with in the monograph are treated. An annotated check list of all excluded and doubtful species is given.

 

Introduction

     The genera dealt with in this monograph are subhyaline or lightly pigmented hyphomycetes belonging to the Moniliales (von Arx, 1974). The species in the group are mostly soil-borne and often keratinolytic and/or pathogenic. Two of the genera, Chrysosporium Corda and Myceliophthora Cost., are linked to the macroconidium-producing dermatophytes, Trichophyton Malmsten and Microsporum Gruby, by the type of (micro)conidium. Several species in this monograph have known associated teleomorphic genera (Table 1).
     The name Chrysosporium was introduced by Corda (in Sturm, 1833) for the single species, C. corii Corda. Saccardo (1901) placed Chrysosporium in synonymy with Sporotrichum Link and consequently the genus was practically forgotten. Hughes (1958) reintroduced Chrysosporium for C. corii and C. pannorum (Link) Hughes (= Geomyces pannorum (Link) Sigler & Carmichael), restricting Sporotrichum to species with coloured, broad hyphae without intercalary conidia. Sporotrichum was later found to be basidiomycetous when von Arx (1971) observed clamp connections in a strain of the type species, S. aureum Link ex S.F. Gray, and when Donk (1974) mentioned an associated Poria metamorphosa (Fuck.) Cooke teleomorph.
     Geomyces Traaen, Emmonsia Cif. & Montemartini, Myceliophthora Cost. and [p. 2] Blastomyces Gilchrist & Stokes (= Zymonema de Beurmann & Gougerot) were reduced to synonymy with Chrysosporium in a revision by Carmichael (1962), leaving the genus morphologically heterogeneous. A number of species attributed to the genus Aleurisma Link ex Fr. are strains of G. pannorum. The type species of Aleurisma, A. sporulosum Link ex Fr., was reidentified as Trichoderma sporulosum (Link ex Fr.) Hughes by Hughes (1958) (today T. polysporum (Link ex Pers.) Rifai). Another monograph of Chrysosporium was published by Dominik (1976) who broadened Carmichael's concept to include certain former species of Sepedonium Link ex Fr. Most of these Sepedonium species are here regarded as doubtful in the absence of clear descriptions and type material; S. spinosum Sacc. has been found to be a basidiomycete (Stalpers. pers. comm.). Since Carmichael (1962) and Dominik (1967) several new species have been described and new material has become available of which a number of strains could not be associated with any known species. A new taxonomic revision of Chrysosporium was therefore deemed overdue.
     The present revision includes those genera formerly regarded as synonymous to Chrysosporium, viz. Myceliophthora, Emmonsia, Zymonema, and Geomyces, with the addition of Trichosporiella Kamyschko ex W. Gams & Domsch, which shows a similar type of conidiogenesis to Myceliophthora, Emmonsia and Zymonema. Delimitation of taxa has been made primarily on the basis of colony and conidium morphology, with temperature resistance and ability to degrade keratin having been employed as additional characters.

 

Conidiogenesis

     Fig. 1 gives a schematic illustration of the thallic and blastic types of conidiogenesis encountered in the present group of fungi. In thallic conidiogenesis an undifferentiated cell is converted into a conidium, while in blastic conidiogenesis the initial is an extrusion of the conidiogenous cell (Cole & Samson, 1979). Holoblastic conidia are produced as extensions from a conidiogenous cell with all the wall layers of that cell being involved in the formation of the conidium wall (Kendrick, 1971). Blastic conidia occur scattered on undifferentiated hyphae (Fig. 1 A, B), or arise solitarily or in small clusters from lateral ampulliform swellings (Fig. 1. C, D). Thallic conidia are formed after conversion of all the wall layers of a terminal, lateral or intercalary segment of a fertile hypha (Fig. 1 E. F). Kendrick (1971) associated blastic development with conidium enlargement before delimitation by a septum; thallic development with enlargement after delimitation by a septum. Observations made of the present group of fungi support Cole & Samson (1979) in that clear blastic and thallic development cannot be differentiated on the basis of septation alone. Moreover, it was found that many intermediate cases of solitary conidia exist. It is considered useful to continue the use of the descriptive distinction of conidia attached with a narrow or a broad base to the subtending cell (von Arx. 1974). Terminal and lateral conidia attached with a broad base in the fungi under consideration have been referred to as chlamydospores (Hughes, 1953), aleuriospores (Vuillemin, [p. 3] 1911; Carmichael, 1962; Barron, 1968) and gangliospores (Subramanian, 1962). The term aleuriospore was first introduced by Vuillemin (1911) for spores which dehisce by the disintegration or fracture of a supporting cell. Use of the term subsequently became confused and it is therefore avoided here. In the studied group conidia borne on ampulliform swellings are obviously blastic, but the thallic or blastic nature of lateral conidia is often difficult to discern. Blastic conidia tend to be more constricted at the base than thallic conidia. Intercalary conidia are described as alternate when separated from one another by single, short, relatively empty hyphal segments (Sigler & Carmichael, 1976). The term chlamydospore was reserved by Kendrick (1971) for thick-walled, thallic, terminal or intercalary spores. Here the term is restricted to thallic, globose, hyaline, often thick-walled, usually intercalary resting cells (sensu Barron, 1968).

 fig. 1

Fig. 1
Conidium development. a. terminal blastic conidium: b. lateral blastic conidium: c. blastic conidium on ampulliform swelling: d. multiple conidium formation on ampulliform swelling; e. terminal holothallic conidium; f. lateral holothallic conidium.

     Large, globose, thick-walled propagules of Emmonsia, which are formed in vivo or at 37°C in vitro through the expansion of holoblastic conidia, are termed adiaspores (Emmons & Jellison, 1960). The cells produced by Zymonema in vivo or at 37°C in vitro have been variously referred to as sprout cells (Dodge, 1935), yeast-like cells (Garrison & Boyd, 1978) or budding cells (Carmichael, 1962; Emmons et al., 1977). The last term is employed in the present study. The cells concerned are liberated blastic conidia with germ tubes which enlarge to give rise to more or less spherical cells which in turn bud, producing secondary cells (Garrison & Boyd, 1978).
     The conidia of the fungi studied are released by fracture of the supporting cells just below the basal septum of the conidium (Fig. 2A). Following dehiscence the conidiogenous cell shows an opening in which no septum can be discerned (Fig. 2B), and the free conidium bears the minute remains of the formerly attached cell (Plate 2). [p. 4] Where the basal scar is narrow, as in Emmonsia and Trichosporiella, these remains are not clearly visible. This type of dehiscence is known as rhexolytic (Hughes. 1971; Cole & Samson, 1979). In all the genera except Geomyces this process resembles that of Endophragmia Duvernoy & Maire as illustrated by Carmichael in Kendrick (1971). However, the point of fracture in the present fungi is slightly lower. In Geomyces fracture occurs across the middle of the short hyphal segment connecting the alternate intercalary conidia. The microconidia of Trichophyton and Microsporum are released in a similar manner to the above genera. Galgoczy (1978) referred to total disintegration of the supporting cells of the microconidia but this does not seem to occur consistently.

 fig. 2

Fig. 2
Rhexolytic dehiscence of conidia. a. before, dehiscence, arrows indicating points of fracture. b. after dehiscence.

 

Trichophyton and Microsporum species with Chrysosporium-like microconidia

     Chrysosporium is related to the genera Trichophyton and Microsporum which have Chrysosporium-like conidia (microconidia) in addition to macroconidia. In Trichophyton the macroconidia are mostly clavate, smooth-walled and measure 8-50 x 4-8 µm, and in Microsporum they are mostly fusiform, with ornamented walls and measure 35-125 x 7-20 µm (Emmons et al., 1977). Macroconidium production often declines with repeated subculturing, leaving only the microconidia. Enriched media sometimes stimulate renewed macroconidium formation. On hair, skin and nails the fungi usually show only intercalary conidia, the macro- and microconidia appearing on agar media. The dermatophyte Epidermophyton Sabour also forms multicellular macroconidia but no microconidia.
     The identification of Trichophyton and Microsporum species on the basis of their microconidia is problematic. Most species occur as dermatophytes on man and animals. Many names were applied on the basis of the clinical aspect only; descriptions of the micromorphology of the fungus were often omitted or vague, and herbarium material was seldom preserved. Copious superfluous names were allotted to species of the two genera and the overall picture had become rather confused. Emmons et al. (1934; 1977), Ajello (1967) and Caprilli et al. (1971) revised Trichophyton and Microsporum, reducing a large number of species to synonymy. For the present study only those species accepted by these authors were examined.
     In Trichophyton several types of microconidium morphology could be distinguished: [p. 5]
(1) The microconidia of species such as Trichophyton vanbreuseghemii Rioux et al. or T. equinum (Matruchot & Dassonville) Gedoelst and many others are hyaline, smooth- and thin-walled, and are born laterally on straight, seldom branched hyphae.
(2) In the Trichophyton mentagrophytes (Robin) Blanchard complex and T. rubrum (Castell.) Sabour. two types of strain can be distinguished, each with different colony characters and conidia. Floccose strains tend to produce clavate, lateral, broad-based microconidia similar to the conidia of Chrysosporium; powdery or granular strains form subglobose to globose, blastic microconidia occasionally borne on ampulliform swellings and with narrow basal attachments. The latter type of conidiogenesis is similar to that seen in Myceliophthora.
(3) Trichophyton terrestre Duriee & Frey produces conidia which remain unicellular or become septate and grade into multiseptate macroconidia. Here the distinction between micro- and macroconidia is not clear.
(4) Trichophyton schoenleinii (Lebert) Langeron & Milochevitch, T. verrucosum Bodin and T. violaceum Sabour. often sporulate poorly or not at all in culture and are recognised primarily by the colony type and hyphae; this in contrast to Chrysosporium which produces repeatedly branched fertile hyphae on initial isolation.
     The anamorph of Arthroderma multifidum Dawson was considered to be a Chrysosporium by Padhye & Carmichael (1971), but the paratype strains CBS 419.71 and 420.71, and the dried type specimen IMI 94207, show pyriform to mostly clavate, rough-walled macroconidia which are 2-5-septate. The macroconidia with only 2-3 septa are broadly clavate, but where there are 4-5 septa the conidia are frequently irregular in shape. The form of these conidia resembles those of Trichophyton, while the wall ornamentation resembles that of Microsporum.
     Microsporum microconidia are borne laterally on little branched hyphae with normally several microconidia per hyphal cell. Microconidia are almost sessile, hyaline, clavate, smooth- and thin-walled, truncate, have broad basal scars and measure 4-7 x 2.5-3.5 µm. The microconidia vary little from species to species, the main difference being in size. The presence of macroconidia and the little differentiated fertile hyphae distinguish Microsporum and Trichophyton species from those of Chrysosporium.

 

Teleomorphs

     The ascomycetous teleomorphs associated with the species described in this monograph are either Gymnoascales (Gymnoascaceae, Onygenaceae or Ascosphaeraceae) or Sordariaceae (von Arx in Kendrick, 1979). The Gymnoascales have free or clustered ascogonia, sometimes surrounded by a spherical peridium of hyphae, small, spherical asci and small, 1-celled, subhyaline or lightly pigmented ascospores devoid of germpores. The type genus of the Sordariaceae, Sordaria Ces. & de Not., has ostiolate ascomata, elongate asci and larger, 1-celled, pigmented ascospores with a single germpore, but no associated anamorph. Table 1 illustrates that the distribution of the teleomorphs over these families does not exactly correlate [p. 6] to the division of the anamorph genera on the basis of conidium morphology.

 

Table 1. The known teleomorphs associated with the anamorph species described in this monograph.

 

ANAMORPH

TELEOMORPH

FAMILY OF TELEOMORPH

 

 

Chrysosporium merdarium

Gymnoascus uncinatus

Gymnoascaceae

 

Chrysosporium queenslandicum

Apinisia queenslandica

Gymnoascaceae

 

Chrysosporium anamorph of

Gymnoascus demonbreunii

Gymnoascaceae

 

Chrysosporium georgii

Arthroderma ciferrii

Gymnoascaceae

 

Chrysosporium anamorph of

Arthroderma curreyi

Gymnoascaceae

 

Chrysosporium anamorph of

Arthroderma cuniculi

Gymnoascaceae

 

Chrysosporium anamorph of

Pectinotrichum llanense

Gymnoascaceae

 

Chrysosporium anamorph of

Renispora flavissima

Gymnoascaceae

 

Chrysosporium anamorph of

Rollandina vriesii

Gymnoascaceae

 

Chrysosporium tropicum

Aphanoascus species

Onygenaceae

 

Chrysosporium keratinophilum

Aphanoascus fulvescens

Onygenaceae

 

Chrysosporium indicum

Aphanoascus terreus

Onygenaceae

 

Chrysosporium farinicola

Bettsia alvei

Ascosphaeraceae

 

Chrysosporium sulfureum

Bettsia species

Ascosphaeraceae

 

 

Myceliophthora anamorph of

Arthroderma tuberculatum

Gymnoascaceae

 

Myceliophthora anamorph of

Ctenomyces serratus

Gymnoascaceae

 

Myceliophthora thermophila

Thielavia heterothallica

? Sordariaceae

 

Myceliophthora fergusii

Corynascus thermophilus

? Sordariaceae

 

Myceliophthora anamorph of

Corynascus novoguineensis

? Sordariaceae

 

Myceliophthora anamorph of

Corynascus sepedonium

? Sordariaceae

 

 

Zymonema dermatitidis

Ajellomyces dermatitidis

Gymnoascaceae

 

 

Geomyces vinaceus

Pseudogymnoascus roseus

Gymnoascaceae

 

     Several species of Chrysosporium, such as C. merdarium, have gymnoascaceous teleomorphs belonging to the genera Gymnoascus Baran., Apinisia La Touche. Arthroderma Currey, Renispora Sigler et al., Rollandina Pat. and Pectinotrichum Varsavsky & Orr. These teleomorphs have ascoma walls consisting of interwoven hyphae which mostly have terminal appendages. The Onygenaceae associated with the three species C. tropicum Carmichael, C. keratinophilum Frey ex Carmichael and C. indicum (Randhawa & Sandhu) Garg belong to the genus Aphanoascus Zukal (= Anixiopsis sensu de Vries. 1969), which has an ascoma wall of a single layer of flattened polygonal cells. The sordariaceous Corynascus v. Arx and Thielavia Zopf teleomorphs associated with Myceliophthora anamorphs (Table 1) have ascoma walls composed of layers of pigmented adjoining flattened cells. The ascoma of Ctenomyces Eidam has a peridium of flattened cells and differentiated appendages. The teleomorphs associated with Myceliophthora species substantiate the close relationship between the Gymnoascaceae and Sordariaceae. Ctenomyces has small ascospores with no germpores and probably represents a reduced form of the sordariaceous Corynascus with larger ascospores showing two germpores. Similar relationships are also found in other reduced, rare ascomycetes and certain Sordariaceae.
     For example a) Leuconeurospora Malloch & Cain with its small, ridged, hyaline ascospores devoid of germpores (Malloch & Cain, 1970) probably represents a reduced gymnoascaceous form of the sordariaceous Neurospora Shear [p. 7] & Dodge which has larger, striate ascospores with 1 or 2 germpores (Muller & von Arx, 1973); b) Renispora Sigler et al. with its finely rough-walled, reniform or bacilliform ascospores devoid of germpores (Sigler et al., 1979) probably represents a reduced gymnoascaceous form of the sordariaceous Gelasinospora Dowding which has larger, reticulate or pitted, ellipsoid ascospores with one or several germpores (von Arx, 1973b). The Myceliophthora anamorphs of Arthroderma tuberculatum Kuehn and Ctenomyces serratus Eidam develop few ampulliform swellings, unlike of their Myceliophthora counterparts. However, the conidia strongly resemble those of M. vellerea (Sacc. & Speg.) van Oorschot and M. thermophila (Apinis) van Oorschot. The conidiogenous cells in M. fergusii (Klopotek) van Oorschot are mostly only slightly swollen and not globose, thus resembling the swollen side branches of Chrysosporium indicum and C. pannicola (Corda) van Oorschot & Stalpers.
     Bettsia alvei (Betts) Skou, the teleomorph of Chrysosporium farinicola (Burnside) Skou, stands out among the other ascomycetes associated with both Chrysosporium and Myceliophthora. The fruiting body is a single enlarged cell, the fertilised nutriocyte. The wall is homogeneous and not composed of cells or hyphae. Chrysosporium sulfureum van Oorschot & Samson produces sterile fruiting structures like those of Bettsia alvei.

 

Material and methods

     Phytone yeast extract agar (PYE; produced by BBL) was used as the standard medium for nearly all the strains studied (Carmichael, 1962). A few species did not show optimum sporulation on PYE and were therefore grown on cherry decoction, honey, 2% malt, hay infusion or Sabouraud's glucose (Difco) agars. A species was termed osmophilic if it grew well on media with high sugar concentrations, such as honey or malt with 20% glucose, and poorly on other media. Descriptions are based on cultures grown on PYE unless stated otherwise. Cultures were incubated in diffuse daylight at 24°C except where a higher temperature stimulated better sporulation; temperatures are then indicated. Growth temperatures were recorded between 0-50°C at 5°C intervals. Strains were regarded as thermotolerant when minima were well below 20°C and maxima between 40-50°C, and as meophilic when they grew between 10°C and 40°C with an optimum between 25°C and 35°C (Cooney & Emerson, 1964). Growth rate and colony appearance were noted after 14 days, or after 7 days if growth was fast; microscopic morphology was examined between 10 to 21 days. (Slide preparations were made using Shear's mounting fluid without addition of stains).
     Each species was also tested for ability to digest keratin by placing a number of fine, blond, prepubertal hairs on the agar 1 cm away from the point of inoculation. Pronounced denser growth on the hairs was recorded as positively keratinolytic, but no differentiation was made between mere penetration of the hair or marked degradation. [p. 8]

 

 

 

Key to the revised and related genera

 

1a.

 

Lateral conidia rare or absent

2

 

1b.

 

Lateral conidia abundant

4

 

 

2a.

 

Branching of fertile hyphae mostly verticillate

Geomyces

 

2b.

 

Branches of fertile hyphae more or less orthotropic

3

 

 

3a.

 

Intercalary conidia broader than the supporting hyphae, delimitation progressing basipetally

(Ovadendron)

 

3b.

 

Intercalary conidia not broader than the supporting hyphae, delimitation at random

(Malbranchea)

 

 

4a.

 

Fertile hyphae forming sporodochia on the natural substrate, isolated from grasses

(Beniowskia)

 

4b.

 

Fertile hyphae not forming sporodochia on the natural substrate, not isolated from grasses

5

 

 

5a.

 

Aerial hyphae mostly absent

Trichosporiella

 

5b.

 

Aerial hyphae present

6

 

 

6a.

 

Conidia supported on long stalks, markedly ornamented, often accompanied by a phialidic state

(Sepedonium)

 

6b.

 

Conidia sessile or on short stalks, not accompanied by a phialidic state

7

 

 

7a.

 

Ornamented macroconidia present

(Histoplasma)

 

7b.

 

Macroconidia absent

8

 

 

8a.

 

Conidia thallic, not borne on ampulliform swellings

Chrysosporium

 

8b.

 

Conidia blastic, often borne on ampulliform swellings

9

 

 

9a.

 

Conidia more than 5 µm in length; colonies spreading; aerial mycelium dense; not forming adiaspores or budding cells at 37°C

Myceliophthora

 

9b.

 

Conidia less than 5 µm in length; colonies restricted; aerial mycelium sparse; forming budding cells or adiaspores at 37°C

10

 

 

10a.

 

Colonies developing adiaspores at 37°C

Emmonsia

 

10b.

 

Colonies developing budding cells at 37°C

11

 

 

11a.

 

Cells at 37°C producing many buds with narrow connections

(Paracoccidioides)

 

11b.

 

Cells at 37°C producing 1 to 3 buds with broad connections

Zymonema

 

 

Descriptions of recognised taxa

 

Chrysosporium Corda

       Chrysosporium Corda in Sturm - Deutschl. Fl. (Pilze) III. Bd. 3, Heft 13: 85. 1833.
       Blastomyces Cost. & Roll - Bull. Soc. mycol. Fr. 4: 153. 1888.
       Rhinocladiella Kamyschko - Notul. syst. Inst, cryptog. Hord bot. petropol. 13: 165. 1960 (non Rhinocladiella Nannf. in Melin & Nannf. - Svenska Akogsvför. Tidskr. 32: 461. 1934) = Rhinocladiopsis Kamyschko - Notul. syst. Inst. cryptog. Horti. bot. petropol. 14: 243. 1961 (name
change). [p. 9]

       Teleomorph genera: Gymnoascus, Apinisia, Arthroderma, Pectinotrichum, Aphanoascus, Bettsia, Renispora, Rollandina.

Colonies usually spreading, white, sometimes cream-coloured, pale brown or yellow, felty and/or powdery. Hyphae mostly hyaline, smooth-walled, with irregular, more or less orthotropic branching. Fertile hyphae not or little differentiated. Terminal and lateral thallic conidia borne all over the hyphae, sessile or on short protrusions or side branches, subhyaline or pale yellow, thin- or thick-walled, subglobose, clavate, pyriform, or obovoid, 1-, rarely 2-celled, truncate. Intercalary conidia sometimes present, solitary, occasionally catenate, subhyaline or pale yellow, broader than the supporting hyphae, normally 1-celled, truncate at both ends. Chlamydospores occasionally present.

       Type species: Chrysosporium merdarium (Link ex Grev.) Carmichael


Discussion

     The species of the genus are mostly isolated from soil and human or animal tissue, especially from keratinous areas of the body such as the skin, hair or nails. On initial isolation colony colour is most pronounced and sporulation most abundant, respectively becoming white and reducing or disappearing with repeated subculturing. Fertile hyphae are more branched than the infertile and tend to be of more constant width. A number of species produce aerial racquet hyphae which are hyaline, smooth- and thin-walled just as the main hyphae, but remain infertile. Submerged hyphae are occasionally thick-walled and contorted and seldom fertile. Conidia are subhyaline to pale yellow, having denser cell contents than the hyphae. Only the conidia of Chrysosporium lobatum Scharapov are darkly pigmented but this character alone does not warrant the erection of a separate genus. The rhexolytic nature of liberation is often quite clearly visible, such species as C. merdarium (Link ex Grev.) Carmichael or C. keratinophilum (D. Frey) Carmichael having marked remains of the subtending hyphal cell (Plate 2). Chlamydospores are not common, usually occurring in degenerate strains.
     Chrysosporium differs from Myceliophthora and Emmonsia by the lack of narrow-based blastoconidia frequently borne on ampulliform swellings; from Trichosporiella by the shape of the conidia and the development of abundant aerial mycelium; from Geomyces by the lack of differentiated fertile hyphae and alternate intercalary conidia separated by single, short, sterile cells; from Malbranchea by the broader intercalary conidia and the presence of terminal and lateral conidia; from Ovadendron Sigler & Carmichael by the solitary intercalary conidia developing at random; from Botryotrichum Sacc. & March. because the conidia are not spherical nor borne on branched conidiophores and lack phialides; from Sepedonium because of the absence of an associated phialidic state and large, brightly coloured, markedly ornamented, globose conidia; from Mycogone Link ex Chev. again because of the absence of an associated phialidic state and also because of the lack of ornamented, 2-celled conidia. The ascomycetes Apinisia graminicola La Touche and Shanorella spirotricha R.K. Benjamin have anamorphs which are mainly Malbranchea - like with chains of cylindrical intercalary conidia each separated by sterile hyphal cells. However, conidia [p. 10] occasionally do develop laterally on intercalary or sterile hyphal cells, thus suggesting certain affiliation to Chrysosporium. Sporotrichum is a basidiomycetous genus which requires revision. The few species which may remain in the genus (Stalpers, pers. comm.) all have known Phanerochaete Karst. or Poria Pers. ex S.F. Gray teleomorphs. Not all species form clamp-connections, but they do differ from Chrysosporium by the usually fast growth rate, the conidia borne on long, often bent side branches which are frequently in sympodial clusters, the intercalary conidia which are few in number and cylindrical, and the frequent development of large chlamydospores.
     Some strains of Arachniotus Schroet. species show scattered, poorly differentiated, intercalary conidia which intergrade with disarticulated hyphal segments. Where these conidia show some differentiation, in the form of dense cytoplasmic content or wall thickening, they resemble intercalary conidia in Chrysosporium and Malbranchea.

 

 

Key to the species

 

1a.

 

Intercalary conidia absent or rare

2

 

1b.

 

Intercalary conidia abundant

16

 

 

2a.

 

Intercalary conidia rare

3

 

2b.

 

Intercalary conidia absent

9

 

 

3a.

 

Conidia thick-walled

4

 

3b.

 

Conidia thin-walled

7

 

 

4a.

 

Conidia reddish-brown to dark brown

C. lobatum

 

4b.

 

Conidia subhyaline

5

 

 

5a.

 

Conidia reaching a maximum length of over 10 11 µm, smooth-walled to echinate or verrucose

6

 

5b.

 

Conidia not reaching a maximum length of over 10 µm, remaining smooth-walled

C. tropicum

 

 

6a.

 

Conidia subhyaline to pale yellow, mostly globose, often markedly verrucose

C. anamorph of Renispora flavissima

 

6b.

 

Conidia subhyaline, obovoid to clavate, smooth-walled or echinate

C. keratinophilum

 

 

7a.

 

Conidia more than 7 µm long

C. pannicola

 

7b.

 

Conidia less than 7 µm long

8

 

 

8a.

 

Conidia less than 3 µm broad, often cymbiform, borne on swollen side branches

C. indicum

 

8b.

 

Conidia more than 3 µm broad, not cymbiform, not borne on swollen side branches

C. carmichaelii

 

 

9a.

 

Conidia developing more or less synchronously, obovoid or subglobose, any sessile subglobose conidia echinate

10

 

9b.

 

Conidia not developing synchronously, often clavate, any sessile subglobose conidia smooth-walled

11

 

 

10a.

 

Synchronous development always marked: mature conidia borne on narrow side branches, basal scars narrow, conidia subhyaline

C. synchronum

 

10b.

 

Synchronous development not always marked: mature conidia sessile or borne on short, broad protrusions, basal scars wide, conidia mostly pale yellow [p. 11]

C .sulfureum

 

 

11a.

 

Conidia smooth-walled

12

 

11b.

 

Conidia echinulate

C. anamorph of Arthroderma curreyi

 

 

12a.

 

Conidia borne singly on orthotropic side branches, mostly clavate or obovoid

13

 

12b.

 

Conidia sessile or on short protrusions, mostly subglobose

14

 

 

13a.

 

Conidia more than 6 µm in length, sometimes 2 to 3-celled

C. anamorph of Arthroderma cuniculi

 

13b.

 

Conidia less than 6 µm in length, never 2 to 3-celled

C. anamorph of Pectinotrichum llanense

 

 

14a.

 

Conidia usually obovoid, borne on conical protrusions on much branched fertile hyphae

15

 

14b.

 

Conidia usually subglobose, sessile on little branched fertile hyphae

C. lucknowense

 

 

15a.

 

Optimum growth temperature 30°C.

C. anamorph of Rollandina vriesii

 

15b.

 

Optimum growth temperature 25°C

C. georgii

 

 

16a.

 

Lateral conidia absent

17

 

16b.

 

Lateral conidia present

18

 

 

17a.

 

Conidia thick-walled, hyphae showing irregularly swollen cells.

C. inops

 

17b.

 

Conidia thin-walled, hyphae devoid of irregularly swollen cells

C. anamorph of Gymnoascus demonbreunii

 

18a.

 

Intercalary conidia sometimes directly adjacent in chains of 3 or more

19

 

18b.

 

Intercalary conidia not in chains

21

 

 

19a.

 

Conidia smooth-walled, 1-celled

20

 

19b.

 

Conidia echinate, 1-, sometimes 2-celled

C. pseudomerdarium

 

 

20a.

 

Lateral or terminal conidia borne on long side branches of little-septate, locally swollen fertile hyphae

C. xerophilum

 

20b.

 

Lateral or terminal conidia mostly sessile on septate, regular fertile hyphae

C. farinicola

 

 

21a.

 

Conidia mostly sparsely echinate, colony yellow, rarely white or green, powdery

C. merdarium

 

21b.

 

Conidia smooth-walled, colony white, powdery

C. queenslandicum

 

 

1. Chrysosporium merdarium (Link ex Grev.) Carmichael - Fig. 3.

       Sporotrichum merdarium Link - Jb. Gewächsk. 1:176. 1818: ex Grev. - Scott. Crypt. Fl. 1:105. 1823 = Chrysosporium merdarium (Link ex Grev.) Carmichael - Can. J. Bot. 40:1160. 1962.
       Sporotrichum inquinatum Link - Jb. Gewächsk. 1:172. 1818: ex Link - Spec. Pl. 6:5. 1824.
       Sporotrichum scotophilum Link - Jb. Gewächsk. 1:180. 1818: ex Link - Spec. Pl. 6:16. 1824.
       Chrysosporium corii Corda in Sturm - Deutschl. Fl. (Pilze) III, Bd. 3, Heft 13:85. 1833 = Sporotrichum corii (Corda) Sacc. & Trav. - Syll. Fung. 20:872. 1911.
       ? Sporotrichum vellereum Sacc. & Speg. var. flavum Sacc. - Michelia 2:635. 1882.
       ? Blastomyces luteus Cost. & Roll. - Bull. Soc. mycol. Fr. 4:153. 1888.
       Chrysosporium merdarium (Link ex Grev.) Carmichael var. roseum W. Gams & Domsch - Nova Hedwiga 18:7. 1969.
       Chyrosporium verruculatum Scharapov - Nov. Syst. nit Rast. 15:143. 1978. [p. 12]

 fig. 3

Fig. 3
Chrysosporium merdarium. CBS 112.63

       Misapplied name: Aleurisma flavissimum Link sensu Vuill. Bull. Séanc. Soc. Sci. Nancy 3:154. 1911.

       Teleomorph: Gymnoascus uncinatus Eidam in Cohn - Beitr. Biol. Pfl. 3:292. 1880.

     Colonies attaining 30-35 mm diam in 14 days, initially white, later often bright yellow, pink or green, slightly fluffy, felty to powdery, up to 10 mm high in the centre and descending to 1 mm outward, sometimes with radial grooves; margin regular, defined; reverse bright yellow at the centre, paler yellow outward. Hyphae hyaline, thin-walled with relatively thicker septa, 1.5-4.5 µm wide; aerial hyphae mostly fertile; submerged hyphae mostly sterile, often contorted. Racquet hyphae present. Terminal and lateral conidia sessile or borne on short protrusions or side branches, solitary, frequently only separated from another intercalary conidium by a single hyphal cell, subhyaline to pale yellow, smooth-walled or sparsely echinate, thick-walled, obovoid to often subglobose, 1-celled, 4-10 x 3-6 µm, with wide basal scars (1.5-3 µm). Intercalary conidia slightly less abundant, either solitary or separated by a single hyphal cell or in pairs, cylindrical- to barrel-shaped, subhyaline to pale yellow, smooth- or sparsely echinate, 5-12 x 3-6 µm. Chlamydospores most common in degenerate cultures, subhyaline, smooth- and thick-walled, globose to subglobose, up to 11 µm diam. [p. 13]
     Growth temperatures: minimum 10°C, optimum 25°C, maximum 25-30°C. Not keratinolytic.


Material examined

Herbarium specimens. - Sporotricum merdarium Link, L, 4284:63, type, on dung, 1818, and slide DAOM 43321 from type material; Sporotrichum inquinatum Link and S. scotophilum Link, B, both type, collected by C.C. Ehrenberg, Germany, 1818; Chrysosporium corii Corda, PR 155415, on leather, Czechoslovakia, 1833, and slide DAOM 55541 from the type material.

Living strains with only the anamorph. - CBS 101.10 = UAMH 652 probably isolated from rat dung by P. Vuillemin; CBS 107.26 isolated by A. Nannizzi; CBS 112.63 = UAMH 649 isolated from plate contamination by J.W. Carmichael, Edmonton, Canada, 1959; CBS 387.68 isolated from wheat field soil by W. Gams, F.R.G., 1963; CBS 388.68 type strain of Chrysosporium merdarium var. roseum W. Gams & Domsch, isolated from soil by J.W. Veenbaas-Rijks, Netherlands; CBS 131.70 isolated from human nail by G.C.L. van de Ploeg, Netherlands, 1969; CBS 826.70 isolated from arable soil by W. Gams, Wageningen, Netherlands; CBS 162.72 isolated from greenhouse soil by H. Kaastra-Höweler, Naaldwijk, Netherlands; CBS 621.73 isolated from soil by J. Reid, Winnipeg, Canada; CBS 225.74 isolated from wheat field soil by W. Gams, F.R.G., 1974; CBS 475.76 and 477.76 isolated by H.M. Yusef, Egypt; CBS 665.78 = V-KMF 2121 type strain of Chrysosporium verruculatum Scharapov, isolated from mouse, Kirov U.S.S.R. 1974.

Living strains with the teleomorph and anamorph. - CBS 507.65 = IMI 111025 = ATCC 16006 isolated from soil by E. Varsaysky, Italy; CBS 408.72 = NRRL 3610 neotype of Gymnoascus uncinatus Eidam, isolated from dung by R.K. Benjamin, Peoria, U.S.A.


Discussion

     Hughes (1958) examined the type material of Sporotrichum merdarium Link and regarded it as a Sporotrichum. Vuillemin (1911) incorrectly identified an isolate of Chrysosporium merdarium from rat dung as Aleurisma flavissimum (cf p. 77). The strain CBS 101.10, which is presumed to be Vuillemin's isolate, is somewhat degenerate but does show typical C. merdarium conidia, as do the original drawings given by Vuillemin (1911). The type material of S. vellereum var. flavum is not preserved at PAD, but the drawing of the type material given by Vuillemin (1911) indicates that the species is probably identical to C. merdarium. The type material of Blastomyces luteus Cost. & Roll. is probably lost (Stafleu & Cowan, 1976), but the original descriptions and drawings (Costantin & Rolland, 1888) suggest synonymy to C. merdarium. Greville (1823) placed his S. sulphureum Grev. in synonymy with S. merdarium. However, the type material (PC HC-1800) shows only an Arthrographis species.
     Link (1824) separated the species Sporotrichum inquinatum, S. scotophilum and S. merdarium on the sole basis of the colour of their colonies, and under each name he listed an alternative name without citation, namely S. merdarium-album, S. merdarium-rubrum and S. merdarium-flavum respectively. Gams & Domsch (1969) separated Chrysosporium merdarium var. roseum on the basis of the bright pink colour of the colony on initial isolation and the uniform shape of the conidia. Scharapov (1978) only compared his species, C. verruculatum, in colour and colony to C. evolceanui (Rhandhawa & Sandhu) Garg and not to C. merdarium. The colony [p. 14] and conidial morphology of the type strain is identical to that of C. merdarium except in that the pale yellow conidia are more markedly rough-walled than most other C. merdarium strains.
     Most strains of Chrysosporium merdarium tend to loose their initial colouring (if any) and become white after repeated subculturing. Some strains even cease to sporulate normally and develop chlamydospores. Strains CBS 475.76 and 477.76 show poor growth at 30°C while all the other strains examined do not grow above 25°C.
     The anamorph of Gymnoascus uncinatus is identical to Chrysosporium merdarium in conidial morphology but the conidia are almost absent on PYE, yet abundant on hay infusion agar.

 fig. 4

Fig. 4
Chrysosporium pseudomerdarium. a. CBS 271.77: b. CBS 631.79.

 

2. Chrysosporium pseudomerdarium van Oorschot, sp. nov. - Fig. 4.

       Coloniae 7-20 mm diam. post 14 dies, plerumque albae, partim dilute flavae, coactae, modice floccosae, fere densae, partim ad 3 mm vel magis elevatae, plicatae: margo acuta, regularis, modice fimbriata; reversum primum cremeum, deinde obscure rubrobrunneum. Hyphae hyalinae, tenuitunicatae, 1-2 µm latae; hyphae aeriae fertiles, submersae steriles. Hyphae septum versus inflatae absunt. Conidia terminalla et lateralia, sessilia yel in protrusionibus brevibus vel ramulis lateralibus oriunda, solitaria vel [p. 15] breviter catenate, subhyalina, primum levia et tenuitunicata, deinde saepe echinulata et crassitunicata, subglobosa, pyriformia vel obovoidea, unicellularia, 2-6.5 x 1.5-5 µm, cicatrice basilari 0.5-1.5 µm. diam. Conidia intercalaria solitaria vel catenata, saepe alternantia, subhyalina, primum levia et tenuitunicata, nonnumquam deinde echinulate et inspissata, doliiformia, saturnoidera vel unilateraliter inflata, 3-6 x 3-6 µm diam. Chlamydosporae absunt.
       Typus CBS 631.79, isolatus ex arena litorali prope Katwijk in Neerlandia, a W. Gams, 1978.

     Colonies attaining 7-20 mm diam in 14 days, mostly white, locally pale yellow, felty, slightly fluffy, fairly dense, raised to at least 3 mm in parts, folded; margin defined, regular, slightly fimbriate: reverse initially cream-coloured, soon becoming dark red-brown. Hyphae hyaline, thin-walled, 1-2 µm wide; aerial hyphae fertile; submerged hyphae sterile. Racquet hyphae absent. Terminal and lateral conidia sessile or on short protrusions or side branches, solitary or in chains of up to 4, subhyaline, initially smooth- and thin-walled, later often becoming echinulate and thicker-walled, subglobose, pyriform or obovoid, 1-celled, 2-6.5 x 1.5-5 µm, with fairly narrow basal scars (0.5-1.5 µm). Intercalary conidia solitary or catenate, frequently alternate, subhyaline, initially smooth- and thin-walled, often remaining so or becoming echinulate and/or thicker-walled, barrel-shaped, saturnoid or unilaterally inflated, 3-6 x 3-6 µm. Chlamydospores absent.
     Growth temperatures: minimum 10°C, optimum 25°C, maximum 25-30°C. Slightly keratinolytic.


Material examined

CBS 271.77 = UAMH 871, isolated from lung of rodent by J.W. Carmichael, Canada, 1960: CBS 631.79 type strain and 633.79 and 636.79 all isolated from beach sand by W. Gams, Katwijk, Netherlands. 1978.


Discussion

     The species differs from Chrysosporium merdarium by the smaller, more densely echinate, often catenate conidia. Strain CBS 271.77 sometimes shows fertile hyphal branches which bend to alternate directions between conidia. Intercalary conidia in CBS 271.77 are occasionally 2-celled.

 

3. Chrysosporium carmichaelii van Oorschot, sp. nov. - Fig. 5.

     Coloniae post 14 dies 15-30 mm diam., albae, intricatae in medio ad circa 3 mm elevatae: margo acuta et regularis vel vaga et velde sulcata, parce fimbriata: reversum dilute cremeoflavum. Hyphae hyalinae, tenuitunicatae, aeriae plerumque fertiles, 1.5-2.5 µm latae, submersae steriles, ad 3 µm latae. Hyphae septum versus inflatae et aeriae et submersae adsunt. Conidia terminalia et lateralia, sessilia vel in protrusionibus brevibus oriunda, solitaria, subhyalina, levia vel exigue asperulata, tenuitunicata, obovoidea ad clavata, unicellularia, rarissime bicellularia, 3-6 x 3.0-3.5 µm, cicatrice basilari 1.5-2.0 µm diam. Conidia intercalaria parca, solitaria, subhyalina, levia et tenuitunicata, cylindrica, saturnoidea vel doliiformia, 3-6 x 1.5-3.0 µm, non multo latioria quam cellulae hyphales. Chlamydosporae absunt.
     Typus: CBS 643.79, isolatus a J.C. Easterling. Michigan. U.S.A., 1970. [p. 16]

 fig. 5

Fig. 5
Chrysosporium carmichaelii. CBS 643.79.

     Colonies attaining 15-30 mm diam in 14 days, white, felty, raised to about 3 mm at the centre; margin defined and regular or undefined and extremely indented, slightly fimbriate; reverse pale creamy yellow. Hyphae hyaline, thin-walled; aerial hyphae mostly fertile, 1.5-2.5 µm wide; submerged hyphae sterile, 1.5-3 µm wide, the wider hyphae being straighter and more regular. Racquet hyphae present in both the aerial and submerged mycelium. Terminal and lateral conidia sessile or on short protrusions, solitary, subhyaline, smooth- or sparsely rough-walled, thin-walled, obovoid to clavate, 1-celled, extremely rarely 2-celled, 3-6 x 3-3.5 µm, with broad basal scars (1.5-2 µm). Intercalary conidia less abundant, solitary, subhyaline, smooth and thin-walled, cylindrical, saturnoid or barrel-shaped, 3-6 x 1.5-3 µm, as wide as or slightly wider than the supporting hyphae. Chlamydospores absent.
     Growth temperatures: minimum 15°C, optimum 25°C, maximum 25-30°C. Not keratinolytic.


Material examined

CBS 383.77 = UAMH 2490 isolated from skin behind human ear by E.P. Hodges, New Orleans, U.S.A., 1965; CBS 639.79 = UAMH 1572 isolated from human sputum by G.F. Orr. 1963; CBS 641.79 = UAMH 2294 isolated from human sputum by H.W. Larsh, Missouri, U.S.A., 1965; CBS 642.79 = UAMH 3571 isolated from rabbit pellet by R. Watling, Scotland; CBS 643.79 = UAMH [p. 17] 3369 type strain, isolated by J.C. Easterling, Michigan, U.S.A., 1970; CBS 644.79 = UAMH 1715 isolated from sputum by J.W. Brandsberg, 1962; CBS 645.79 = UAMH 2857 isolated from soil by G.F. Orr, U.S.A., 1967.


Discussion

     Chrysosporium carmichaelii closely resembles C. merdarium because of the relative positioning of the terminal, lateral and intercalary conidia on the hyphae, but differs by the thinner colony, better growth at 30°C, and the thin-walled, relatively smaller conidia which tend to be clavate rather than subglobose. Strain CBS 645.79 shows some deviation in that it is able to grow, though poorly, at 35°C and in that the centre of the colony is pale yellow.

 fig. 6

Fig. 6
Chrysosporium queenslandicum. CBS 167.78.

 

4. Chrysosporium queenslandicum Apinis & Rees - Fig. 6.

       Chrysosporium queenslandicum Apinis & Rees - Trans. Br. mycol. Soc. 67:524. 1976.
       Chrysosporium crassitunicatum Kushwaha & Agrawal - Trans. Br. mycol. Soc. 68:464. 1977.
       Chrysosporium articulatum Scharapov - Nov. Syst. niz. Rast 15:146. 1978.
       Chrysosporium articulatum Scharapov var. minus Scharapov - Nov. Syst. niz. Rast. 15:146. 1978. [p. 18]

       Teleomorph: Apinisia queenslandica Apinis & Rees - Trans. Br. mycol. Soc. 67: 524. 1976.

     Colonies attaining 55-65 mm diam in 14 days, white. felty, slightly fluffy, up to 2-3 mm high and dense throughout, sometimes with concentric zones; margin defined, rarely regular; reverse pale creamy yellow, occasionally pinkish. Hyphae hyaline, thin-walled; aerial hyphae mostly fertile. 1.5-4.5 µm wide; submerged hyphae mainly sterile, often fragmenting, sometimes contorted, 1.5-6 µm wide, occasionally developing chains of swollen cells. Racquet hyphae present. Terminal and lateral conidia sessile or on short protrusions or side branches, solitary or rarely subtended by another conidium, subhyaline, smooth- and slightly thick-walled, obovoid, clavate or nearly ellipsoid, 1-celled, 3.5-9.5 x 3-6 µm, with broad basal scars (1-3 µm). Intercalary conidia fairly abundant, solitary, occasionally in series of 2-3, alternate, subhyaline, smooth- and slightly thick-walled, barrel-shaped or cylindrical, 5.5-11 x 2-4 µm. Chlamydospores absent.
     Growth temperatures: minimum 10°C, optimum 25°C, maximum 35-40°C. Keratinolytic.


Material examined

Living strains with only the anamorph. - CBS 167.78 = IMl 185320 type strain of Chrysosporium crassitunicatum, isolated from human hair buried in soil by R.K.S. Kushwaha and S.C. Agrawal. India, 1977; CBS 662.78 = VKM-F 2116 type strain of Chrysosporium articulatum, isolated from cool of Sicistae betulinae Pal. by V.M. Scharapov, Novosibirsk, U.S.S.R., 1974; CBS 663.78 = VKM-F 2117 type strain of Chrysosporium articulatum var. minus, isolated from wool of Apiodemus agrarius Pal. by V.M. Scharapov, Novosibirsk, U.S.S.R., 1974; CBS 629.79 = UAMH 1676 isolated from snake by L. Ajello, U.S.A., 1963.

       Living strain with the teleomorph and anamorph. - CBS 280.77 = IMl 121675 type strain of Chrysosporium queenslandicum, isolated from feathers of domestic fowl by A.E. Apinis & R.G. Rees, Cunnumala, Australia. 1965.


Discussion

     The species resembles Chrysosporium tropicum Carmichael but is distinguished by the denser, higher colony, the abundance of cylindrical to barrel-shaped intercalary conidia and the associated teleomorph.
     The type strain of Chrysosporium articulatum agrees with the above description in all respects except in that the growth is less dense and in the occasional production of swollen terminal cells. Variety minus shows a pinkish reverse and the conidia are smaller on average but still fall within the range here given for C. queenslandicum. All the type strains listed above were isolated from chitinous sources, that is, feathers, human hair and wool. [p. 19]

 fig. 7

Fig. 7
Chrysosporium anamorph of Gymnoascus demonbreunii. CBS 122.67.

 

5. Chrysosporium anamorph of Gymnoascus demonbreunii Ajello & Cheng - Fig. 7.

       Gymnoascus demonbreunii Ajello & Cheng - Mycologia 59:692. 1976.

     Colonies attaining 25-35 mm diam in 14 days on hay infusion agar, white felty, flat, less than 1 mm high; margin defined, regular, slightly fimbriate; reverse pale brown. Hyphae hyaline, thin-walled, 0.5-3 µm wide; aerial hyphae fertile; submerged hyphae sterile. Racquet hyphae absent. Conidia terminal, never lateral, solitary, subhyaline, smooth- and thin-walled, pyriform to obovate, 1-celled, 6-9 x 5-6 µm, with wide basal scars (1.5-2.5 µm). Intercalary conidia most abundant, solitary or in series of 2-3 alternate conidia, subhyaline, smooth- and thin-walled, vase-shaped or inequilaterally swollen, 5-11 x 4.5-6 µm. Chlamydospores usually present in ageing cultures, hyaline, smooth- and fairly thick-walled, subglobose, globose or ellipsoid, 6-22 x 4-18 µm.
     Growth temperatures: minimum 10°C, optimum 20-25°C, maximum 30°C. Keratinolytic.


Material examined

CBS 121.67 = ATCC 18027 isolated from human by L. Ajello. Puerto Rico; CBS 122.67 = ATCC 18028 type strain, isolated from soil beneath starling roost by L. Ajello, Manteno. U.S.A.; CBS 123.67 = ATCC 18029 isolated from soil by L. Ajello, U.S.A.


Discussion

     The anamorph is unlike other known Chrysosporium species because it lacks [p. 20] lateral conidia. The shape of the intercalary conidia prevents its inclusion in Malbranchea. The conidia resemble those of C. merdarium, the anamorph of Gymnoascus uncinatus, but the walls are much thinner and smooth. Ovadendron sulphureo-ochraceum (van Beyma) Sigler & Carmichael also has series of intercalary conidia but these are delimited basipetally and are directly adjacent and barrel-shaped.
     Ajello & Cheng (1967) originally regarded Histoplasma capsulatum Darling as the anamorph of Gymnoascus demonbreunii. This was disproved by Kwon-Chung (1968) who showed that the fungus only has the kind of microconidia (terminal and lateral conidia) mentioned above, while one of the cultures originally described was actually a mixed culture of H. capsulatum and G. demonbreunii.

 fig. 8

Fig. 8
Chrysosporium tropicum. a. CBS 109.67; b. CBS 171.62.

 

6. Chrysosporium tropicum Carmichael - Fig. 8.

       Chrysosporium tropicum Carmichael - Can. J. Bot. 10: 1170. 1962.

       Teleomorph: Aphanoascus species. [p. 21]

     Colonies attaining 50-60 mm diam in 14 days, white, felty, powdery, most dense at the centre, less than 1 mm high; margin not well defined, fimbriate; reverse pale creamy yellow. Hyphae hyaline, thin-walled; aerial hyphae fertile, 0.5-3 µm wide; submerged hyphae sterile, 0.5-6 µm wide, the narrower occasionally being contorted. Racquet hyphae usually absent. Terminal and lateral conidia sessile or on short protrusions or side branches, solitary, subhyaline, smooth- and slightly thick-walled, obovoid to clavate, 1-celled, rarely 2-celled, 3.5-7.5 (10) x 3-4.5 µm, with wide basal scars (1.5-2 µm). Intercalary conidia not common, subhyaline, smooth- and slightly thick-walled, cylindrical to barrel-shaped, 3-4 x 6-10.5 µm, of the same width as or slightly wider than the supporting hypha. Chlamydospores (on oatmeal agar) subhyaline, smooth- and thick-walled, globose, 4.5-11.5 µm diam, with narrow basal scars.
     Growth temperatures: minimum 10°C, optimum 25-30°C, maximum 30-40°C. Keratinolytic.


Material examined

Living strains with only the anamorph. - CBS 277.30 = DAOM 41826 isolated by H.R. Whetzel; CBS 171.62 = IMI 94288 = UAMH 691 = JQMD 1087 = ATCC 14802 = IFO 7587 type strain, isolated from woollen cloth by W.L. White, Guadalcanal, 1945; CBS 109.67 isolated by K. Aschan-Oberg, Uppsala, Sweden; CBS 435.77 soil baited with hairs by H.M. Abdel-Fattah, Egypt.

Living strain with the teleomorph and anamorph. - CBS 763.79 isolated from polder South Flevoland by M.C.C. Elders, Netherlands, 1976.


Discussion

     Dominik (1967) regarded Chrysosporium lucknowense Garg as a synonym of C. tropicum, but the former has smaller, always lateral, conidia which are borne almost adjacent on the hyphae. Chrysosporium tropicum can be distinguished from C. keratinophilum (D. Frey) Carmichael by the smaller, smooth-walled conidia and the smaller number of intercalary conidia. Padhye (1969) tested the thermotolerance of numerous strains of C. tropicum and C. keratinophilum and found that the C. tropicum strains were able to grow at 45°C but the C. keratinophilum strains did not grow above 37°C. However, the C. tropicum strains listed above, of which the type strain was also examined by Padhye (1969), did not grow above 40°C.
     White et al. (1950) described seven isolates of fungi causing deterioration of woolen fabric and referred to these as 'Ctenomyces-like'. Carmichael (1962) reidentified one of these isolates as Chrysosporium pruinosum (Gilmann & Abbott) Carmichael (= Sporotrichum pruinosum Gilmann & Abbott), and described the remainder as C. tropicum. The species has been reported to develop yeast-like cells and to cause granulomatous skin-lesions when inoculated into white mice (Hubálek & Hornick, 1977).
     The teleomorph has only been seen in association with strain CBS 763.79. It probably represents a new species of Aphanoascus distinguished by the small ascospores. [p. 22]

 fig. 9

Fig. 9
Chrysosporium keratinophilum. a. CBS 476.76: b. CBS 480.76.

 

7. Chrysosporium keratinophilum D. Frey ex Carmichael - Fig. 9.

       Aleurisma keratinophilum D. Frey - Mycologia 51: 641. 1959. (without indication of type material)
       = Chrysosporium keratinophilum (D. Frey) ex Carmichael - Can. J. Bot. 40: 1157.1962.
         Chrysosporium kuzurovianum Scharapov - Nov. Syst. niz. Rast. 11.266. 1974.

       Teleomorph: Aphanoascus fulvescens (Cooke) Apinis - Mycopath. Mycol. appl. 35:99. 1968.

     Colonies attaining 40-55 mm diam in 7 days, white, cream or sulphur yellow, felty or powdery (especially in fresh isolates), dense at the center, raised to up to 1 mm; margin regular, not well-defined, somewhat fimbriate; reverse pale creamy yellow, occasionally pale reddish yellow. Hyphae hyaline, thin-walled; aerial hyphae fertile, 0.5-7.5 µm wide, submerged hyphae mostly sterile, 0.5-9 µm wide, the narrower often being a little contorted. Racquet hyphae present. Terminal and lateral conidia sessile or on short protrusions, solitary, occasionally subtended by another conidium, subhyaline, smooth-walled or echinate, thick-walled, obovoid to clavate, 1-celled, 3.5-22 x 3.5-11 µm, with marked variation in size in each strain, with wide basal scars (2.5-4 µm). Intercalary conidia less abundant (even rare in some strains), solitary, subhyaline, smooth-walled or echinate, thick-walled, cylindrical to barrel-shaped, 6-25 x 3.5-7 µm, of the same width as or slightly wider than the supporting [p. 23] hypha. Chlamydospores hyaline, smooth- and thick-walled, globose, up to 15 µm diam.
     Growth temperatures: minimum 10°C, optimum 25°C, maximum 30-40°C. Keratinolytic.


Material examined

Living strains with only the anamorph. - CBS 104.62 = IMI 91692 = ATCC 14803 = IFO 7584 type strain, isolated from soil, New Guinea, 1957; CBS 219.62 isolated from soil by T. Pinto Ribeiro, Portugal; CBS 913.70 isolated from rotting hoof by W. Gams and G.A. de Vries, Netherlands; CBS 476.76 isolated by H.M. Yusef, Egypt; CBS 480.76 isolated from soil by M.C.C. Elders, Netherlands; CBS 667.78 = VKM-F 2119 type strain of Chrysosporium kuzurovianum isolated from soil, Tomsk, U.S.S.R., 1969.

Living strains with the teleomorph and anamorph. - CBS 111.58 isolated from bear dung by R.F. Cain, Ontario, Canada, 1957; CBS 121.64 = ATCC 16542 isolated from soil by T. Pinto Ribeiro, Portugal, 1964; CBS 305.67 isolated from soil by A. von Klopotek. F.R.G.; CBS 392.67 = IMI 129854 = ATCC 14862 type strain of Anixiopsis reticulispora, isolated from soil by J.B. Routien, New Zealand, 1961; CBS 741. 68 isolated from chicken crest experimentally infected with Trichophyton gallinae (Megnin) Silya & Benham by H. Rieth, 1963; CBS 742.68 isolated by T. Benedek, U.S.A., 1964; CBS 743.68 isolated by W. Loeffler, Switzerland; CBS 316.76 isolated from dog fur by M.C.C. Elders, Netherlands; CBS 456.77 isolated from soil by R. Samson, Israel; CBS 457.77 and 458.77 isolated from soil by A.A. Padhye, U.S.A., 1974; CBS 459.77 isolated from agricultural soil by W. Verkerke, Netherlands.


Discussion

     Chrysosporium keratinophilum strains can be distinguished from C. tropicum by the larger conidia, occasionally the presence of more intercalary conidia and the often echinate conidial walls. Certain isolates, such as the type strain of C. kuzurovianum, are practically devoid of intercalary conidia, but the variation in size of the conidia disallow confusion with C. tropicum.
     De Vries (1969) attributed the anamorph of Aphanoascus (Anixiopsis) fulvescens
to the genus Chrysosporium but did not name the species. He separated Anixiopsis fulvescens var. stercoraria from var. fulvescens because the second named has larger ascospores and more regular reticulate wall ornamentation. The conidia of the two varieties are identical.
     Hubálek & Hornich (1977) intraperitoneally inoculated white mice with Chrysosporium keratinophilum. The fungus could be reisolated after two months and different strains caused marked splenomegaly, nodules on the liver and omentum, and an abscess on the intestine. Tissue sections showed hyphae, conidia and budding cells. Their results indicate that C. keratinophilum is potentially pathogenic to white mice. There have been few reports of direct isolation from animal tissue other than the hair. Krempl-Lamprecht (1965) reported occasional spontaneous infections of human nails; of the isolates listed above one came from a rotting hoof and another from a chicken's crest. [p. 24]

 fig. 10

Fig. 10
Chrysosporium xerophilum. CBS 153.67.

 

8. Chrysosporium xerophilum Pitt - Fig. 10.

       Chrysosporium xerophilum Pitt - Trans. Br. mycol. Soc. 49: 468. 1966.

     Colonies on cherry decoction agar attaining 85 mm diam in 7 days, white, powdery, flat; margin well-defined, more or less regular, slightly fimbriate; reverse cream-coloured. Colonies on PYE agar only attaining 35-40 mm diam in 7 days, white, thin, powdery, flat; margin ill-defined, more or less regular, slightly fimbriate; reverse cream-coloured. Hyphae hyaline, mostly thin-walled, 3-7.5 µm wide; aerial hyphae fertile, sometimes locally swollen, often devoid of septa at the bases of the branches; submerged hyphae mostly sterile or producing chlamydospores, occasionally slightly contorted. Racquet hyphae absent. Terminal and lateral conidia sometimes sessile, often on side branches of variable length and frequently originating from swollen parts of the main hyphae, solitary, subhyaline, smooth-walled, thin- or thick-walled, subglobose to obovoid or pyriform, 4-13 x 3-10 µm, with broad basal scars (1.5-4 µm). Intercalary conidia may or may not be abundant, solitary or in chains of 2-6, subhyaline, smooth-walled, either thin-walled, cylindrical and 7.5-12 x 3-4.5 µm, or thick-walled, subglobose to barrel-shaped and 6-16 [p. 25] x 6-8 µm. Chlamydospores subglobose to globose, smooth- and thick-walled, 17-30 x 17-27 µm.
     Growth temperatures: minimum 15°C, optimum 25°C, maximum 40°C. Slightly keratinolytic. Osmophilic.


Material examined

CBS 153.67 = UAMH 2368 type strain, isolated from spoiled high-moisture prunes by J.J. Pitt, New South Wales, Australia. 1965.


Discussion

     The species is marked by a great variation in conidial morphology under identical conditions of medium and temperature. The chains of thick-walled intercalary conidia resemble those of C. farinicola which is also osmophilic. Chrysosporium xerophilum is distinguished from C. farinicola because its fertile side branches frequently develop from swollen parts of the main hyphae and often lack basal septa. The morphology also somewhat resembles C. inops Carmichael but this has thin-walled catenate cells, a restricted, heaped and butyrous colony, and is not osmophilic.

 

9. Chrysosporium inops Carmichael - Fig. 11.

       Glenosporella dermatitidis Agostini - Atti. Ist. bot. Univ. Lab. crittogam. Pavia 4: 98. 1930 [non Chrysosporium dermatitidis (Gilchrist & Stokes) Carmichael - Can. J. Bot. 40: 1154. 1962] =
       Aleurisma dermatitidis (Agostini) Dodge - Medical Mycology p. 789. 1935.
       Chrysosporium inops Carmichael - Can. J. Bot. 40: 1156. 1962 (name change).

     Colonies attaining 0.5-10 mm diam in 14 days, cream-coloured, restricted, butyrous, tough, heaped, raised to 1-3 mm, occasionally with sparse aerial tufts of hyphae up to 1 mm in height; margin defined, irregular; reverse cream-coloured. Aerial and submerged hyphae rarely fertile, hyaline, thin-walled, mostly made up of chains of irregular, swollen, 4-13 µm wide cells which may in turn give rise to side branches, hyphae occasionally more regular and cylindrical, 1-4 µm wide. Racquet hyphae absent. Conidia terminal or intercalary along the regular hyphae, mostly solitary, subhyaline, smooth- and thick-walled, obovoid, ellipsoid or somewhat irregular in shape, 1-celled, 6.5-12 x 5-9 µm, mostly truncate with wide basal scars (2-4 µm). Chlamydospores absent.
     Growth temperatures: minimum 20°C, optimum 25°C, maximum 25-30°C. Not keratinolytic. [p. 26]

 fig. 11

Fig. 11
Chrysosporium inops. CBS 132.31.


Material examined

CBS 131.21 = IMI 96729 = UAMH 802 = IFO 7583 type strain, isolated from human skin by A. Agostini, Italy. 1930.


Discussion

     The type strain sporulates poorly but the morphological characters still resemble those depicted in the original diagrams given by Agostini (1930). The species probably reacts unfavourably to culturing on an agar medium such that conidium production is reduced. Carmichael's (1962) second isolate (UAMH 673) is no longer viable (Mrs. L. Northcott, pers. comm.). The fungus is unlike other Chrysosporium species because of its butyrous, restricted colony and the irregular hyphae and conidia. Single, intercalary, chlamydospore-like conidia are also found in poorly sporulating strains of the dematiaceous genus Humicola Traaen. [p. 27]

 fig. 12

Fig. 12
Chrysosporium farinicola. CBS 688.71.

 

10. Chrysosporium farinicola (Burnside) Skou. - Fig. 12.

       Ovularia farinicola Burnside - Pap. Mich. Acad. Sci. 8: 81. 1928 (as O. farinaecola) = Chrysosporium farinicola (Burnside) Skou - Friesia 11: 70. 1975.
       Chrysosporium fastidium Pitt - Trans. Br. mycol. Soc. 49: 467. 1966.

       Teleomorph: Bettsia alvei (Betts) Skou - Friesia 10: 7. 1972.

     Colonies on honey agar attaining 27-45 mm diam in 14 days, initially white, remaining white or becoming pale greenish yellow to pale brown, felty and slightly fluffy and powdery, most dense at the centre, occasionally with faint concentric zoning, up to 1 mm in height; margin defined, regular, very slightly fimbriate; reverse pale yellow. Hyphae hyaline, thin- to thick-walled, 2-8 µm wide, a single hypha often varying in width along its length, hyphae often producing irregularly swollen cells which may form singly or in chains and become thick-walled; aerial and submerged hyphae similar except in that the aerial produce relatively more conidia and the submerged more ascomata. Racquet hyphae absent. Terminal and lateral conidia sessile or on short, often conical, protrusions, solitary, subhyaline, smooth- and thick-walled, subglobose, obovoid or slightly pyriform, 1-celled, 6-11.5 x 4.5-9 µm, with relatively wide basal scars (1.5-5 µm). Intercalary conidia abundant on honey agar but not on malt + 40% sucrose agar, occasionally solitary, usually in chains (not always clearly distinguishable from the thick-walled, swollen, hyphal [p. 28] cells), subhyaline, smooth- and thick-walled, subglobose, short-cylindrical or barrel-shaped, 3-12 x 5-12 µm, of the same width as or slightly wider than the supporting hypha, truncate at both ends or with rounded tip. Chlamydospores absent.
     Growth temperatures: minimum 15°C, optimum 25°C, maximum 25-30°C. Not keratinolytic. Osmophilic.


Material examined

     Living strains with the teleomorph and anamorph.- CBS 137.22 isolated from honey comb by A. Maassen; CBS 223.30 and 224.30 isolated by A. Maurizio; CBS 154.67 = ATCC 18053 = UAMH 2369 = IMl 126288 type strain of Chrysosporium fastidium, isolated from prunes by J.I. Pitt, N.S. Wales, Australia, 1965; CBS 299.71 isolated from equipment for grading prunes by J.I. Pitt, Australia. 1970; CBS 688.71 = IMI 160840 neotype of Bettsia alvei, isolated from pollen in honey comb by J.P. Skou, Denmark.


Discussion

     Chrysosporium farinicola may be compared to C. sulfureum van Oorschot & Samson which produces similar fruiting bodies though these remained sterile. Chains of thick-walled, intercalary conidia are also seen in C. xerophilum which is also osmophilic but has a distinctive white, flat, spreading colony and longer lateral, non-septate hyphal branches bearing terminal and lateral conidia.
     The type material of Chrysosporium farinicola could not be traced, but the neotype of Bettsia alvei, CBS 688.71, shows a conidial morphology identical to that originally described by Burnside (1928) and Skou (1975) for Chrysosporium (Ovularia) farinicola and B. alvei respectively. The type strain of C. fastidium, CBS 154.67, produces sterile ascomata characteristic of B. alvei, thus substantiating synonymy with C. farinicola. Cross mating tests between all the strains examined failed to produce fertile ascomata on either honey, PYE or malt + 40% sucrose. CBS 688.71 is here proposed as the neotype strain for C. farinicola.

 

11. Chrysosporium sulfureum (Fiedl.) van Oorschot & Samson. comb. nov. - Fig. 13.

       Isaria sulfurea Fiedl. in Rabenh. Fungi eur. Ill, 60. 1859 (basionym).
       Isaria sulfurea Fiedl. var. ossicola Speg.- An. Mus. nac. Hist. nat. B. Aires 23: 120. 1912.

     Colonies attaining 10-15 mm diam in 14 days, pale creamy-yellow, felty or slightly powdery, 1-2 mm high, with a butyrous outer area devoid of aerial mycelium: margin defined, sometimes indented; reserve yellow. Hyphae hyaline to pale yellow, thin-walled, 1-4 µm wide, wall of the septa sometimes thickened; aerial hyphae mostly fertile and then more densely branched, submerged hyphae producing relatively few conidia but more fruiting bodies. Racquet hyphae absent. Terminal and lateral conidia sometimes developing semi-synchronously, sessile or on short [p. 29] conical protrusions, rarely on short side branches, solitary, 2-8 per conidiogenous cell, subhyaline to pale yellow, initially smooth- and thin-walled, becoming thick-walled and sometimes echinulate, subglobose, obovoid, occasionally clavate or pyriform, 1-celled, 3-8 x 3-7 µm, with broad or narrow basal scars (1-4 µm). Intercalary conidia absent. Chlamydospores absent.
     Growth temperatures: minimum 5°C, optimum 25°C, maximum 25-30°C. Not keratinolytic.

 fig. 13

Fig. 13
Chrysosporium sulfureum. a. CBS 632.79; b. CBS 634.79.


Material examined

Herbarium specimens. - Isaria sulfurea type, PAD, B. L, LP, isolated from bones, Doemitz, F.R.G.;
Isaria sulfurea var. ossicola, type, LP, no. 32799.

Living strains. - CBS 152.67 isolated from dead frog by H.A. van der Aa, Netherlands; CBS 632.79 isolated from cheese, St. Nectaire, France, 1974; CBS 634.79 = ETH-M 8821 isolated from cheese rind, received from E. Müller, Zürich, Switzerland, 1978. [p. 30]


Discussion

     CBS 634.79 and 152.67 produced immature ascomata on PYE, honey and hay infusion agars. These fruiting bodies were similar to those of Bettsia alvei, found in cultures of C. farinicola, but remained sterile in mating tests. The anamorph differs from C. farinicola by the absence of intercalary conidia, by the slightly smaller and sometimes echinate terminal and lateral conidia, and by being able to grow on media with low or high concentrations of sugar or salt. Strain CBS 632.79 does not produce fruiting bodies and often shows many wide fertile hyphae (4 µm).
     Strain CBS 634.79 was isolated as a contaminent from the rind of a manufactured 'formagelli' cheese. The same fungus has been reported from home made 'formagelli' cheese in Ticino, Switzerland, its presence having promoted sales (E. Müller, pers. comm.). The species has been isolated from bone, cheese and a dead frog, suggesting a preference for fatty or calcium-rich material.

 fig. 14

Fig. 14
Chrysosporium anamorph of Renispora flavissima. a. CBS 708.79; b. CBS 709.79. [p. 31]

  

12. Chrysosporium anamorph of Renispora flavissima Sigler et al. - Fig. 14.

       Teleomorph: R enispora flavissima Sigler et al. - Mycotaxon 10: 133. 1979.

     Colonies attaining 30-40 mm diam in 14 days, pale yellow with a buff centre, felty and powdery, 0.5-2 mm high; margin regular, defined, slightly fimbriate; reverse reddish orange. Hyphae hyaline, thin-walled, 1.5-4.5 µm wide, septa sometimes thickened: aerial and submerged hyphae mostly fertile and then more branched. Racquet hyphae rare and showing only slight swellings. Terminal and lateral conidia sessile or on short protrusions or side branches, solitary, rarely subtended by another conidium, subhyaline to pale yellow, initially smooth- and thin-walled, becoming thick-walled and usually verruculose or markedly verrucose with projections reaching a maximum length of 1.5 µm, mostly globose, sometimes pyriform, 1-celled, 5-12 x 4-12 µm, with broad basal scars (1.5-3 µm). Intercalary conidia rare, sometimes alternate, subhyaline to pale yellow, initially smooth- and thin-walled, remaining so or becoming thick-walled and sometimes verruculose, barrel-shaped or urn-shaped, 1-celled, 6-8 x 5-8 µm. Chlamydospores absent.
     Growth temperatures: minimum 20°C, optimum 25°C, maximum 30-37°C. Keratinolytic.


Material examined

Living strains with the teleomorph and anamorph. - CBS 708.79 (-) and 709.79 (+) mating type strains of Renispora flavissima, isolated from soil in barn housing Myotis verlifer, by P.K. Gaur and R.W. Lichtwardt. S.E Kansas, U.S.A., 1977.


Discussion

     The anamorph differs from Chrysosporium sulfureum by the larger, more globose conidia which become prominently verrucose rather than echinate. Further, C. sulfureum develops semi-synchronously and does not form intercalary conidia. The type ornamentation resembles that of the macroconidia of Histoplasma capsulatum Darling, a species which develops additional microconidia at 24°C and budding cells at 37°C. Renispora flavissima shows little or no mycelial growth at 37°C, but no budding cells.

 

13. Chrysosporium georgii (Varsaysky & Ajello) van Oorschot, comb. nov. - Fig. 15.

       Trichophyton georgii Varsavsky & Ajello - Riv. Patol. veg., Padova III, 4: 357. 1964 (basionym).

       Teleomorph: Arthroderma ciferrii Varsaysky & Ajello - Riv. Patol. veg., Padova III, 4: 358. 1964. [p. 32]

 fig. 15

Fig. 15
Chrysosporium georgii. a. CBS 272.66: b. CBS 269.77.

     Colonies on malt 2% agar attaining 10-30 mm diam in 14 days, white, occasionally pale buff or pink, felty, fluffy or powdery, 1-3 mm high at the centre; margin not well defined, fimbriate; reverse cream-coloured. Hyphae hyaline, smooth-walled: aerial hyphae mostly fertile, thin-walled, 1.5-4.5 µm wide; submerged hyphae mostly sterile, 1.5-6 µm wide, the narrower being thin-walled and occasionally contorted, the broader thick-walled and straight. Racquet hyphae sometimes present. Terminal and lateral conidia borne on short, mostly conical, side branches, solitary, subhyaline, smooth- and thin-walled, subglobose, obovoid or clavate, 1-celled, rarely 2- or 3-celled, 3-8 x 2-3 µm, with relatively wide basal scars (0.5-4.5 µm). Intercalary conidia absent. Chlamydospores absent.
     Growth temperatures: minimum 10°C, optimum 25°C, maximum 30°C. Keratinolytic.


Material examined

Living strains with only the anamorph. - CBS 269.77 = UAMH 1887 isolated from soil by G.F. Orr, California, U.S.A., CBS 227.79 = UAMH 2535 isolated from opossum hair (Didelphys marsupialis) by W. Kaplan and L. Ajello, Georgia, U.S.A., 1958; CBS 625.79 = UAMH 1043 isolated from soil by [p. 33] G.F. Orr, California. 1961; CBS 646.79 = UAMH 2618 isolated as culture contaminant by P.M.D. Martin, Edmonton, Canada. 1966.

Living strain with the teleomorph and anamorph. - CBS 272.66 = UAMH 2534 type strain of Arthroderma ciferrii, isolated from soil by E. Varsaysky & L. Ajello, U.S.A., 1964.


Discussion

     Chrysosporium georgii may be distinguished from C. carmichaelii by the absence of intercalary conidia, and by the fact that lateral conidia are echinate and borne on shorter side branches in closer proximity. Only C. georgii is keratinolytic.
     The conidia are only rarely 2- or 3-celled and do not develop into the multicellular, thick-walled macroconidia typical of Trichophyton species. In the original article (Varsaysky & Ajello, 1964) it was also stated that no macroconidia were produced and yet the species was still placed in Trichophyton. The teleomorph, Arthroderma ciferrii, and the anamorph were based on the same type culture, CBS 272.66.

 fig. 16

Fig. 16
Chrysosporium lucknowense a. CBS 143.77: b. CBS 144.77.

 

14. Chrysosporium lucknowense Garg - Fig. 16.

       Chrysosporium lucknowense Garg - Mycopath. Mycol. appl. 30: 224. 1966. [p. 34]

     Colonies attaining 55 mm diam on sabouraud glucose agar in 14 days, cream-coloured, felty and fluffy, dense and 3-5 mm high; margin defined, regular, fimbriate; reverse pale yellow to cream-coloured. Hyphae hyaline, smooth- and thin-walled, little branched; aerial hyphae mostly fertile, closely septate 1-3.5 µm wide; submerged hyphae infertile, 1-4.5 µm wide, the thinner hyphae often being contorted. Racquet hyphae present. Terminal and lateral conidia mostly sessile or on short, frequently conical protrusions or short side branches, solitary but in close proximity to one another, 1-4 conidia developing on one hyphal cell, subhyaline, fairly thin- and smooth-walled, mostly subglobose, also clavate or obovoid, 1-celled, 2.5-11 x 1.5-6 µm, with broad basal scars (1-2 µm). Intercalary conidia absent. Chlamydospores absent.
     Growth temperatures: minimum 10°C, optimum 25-30°C, maximum 35°C. Keratinolytic.


Material examined

CBS 251.72 isolated from feather by G.F. Orr, California, U.S.A., 1972; CBS 143.77 = IMl 112798 type strain, isolated from soil by P. Tiagi, India, 1965; CBS 272.77 = UAMC 3675 isolated by A.A. Padhye, Atlanta, U.S.A., 1974.


Discussion

     The type strain was not seen to produce the rare 2- or 3-celled conidia nor the inconspicuously roughened conidia mentioned in the original description (Garg. 1966). The species differs from other Chrysosporium species, including those with 2- or 3-celled conidia, by the frequent septation of the fertile hyphae and the close proximity of the lateral conidia. In Geomyces the conidia are also close to one another but intercalary conidia predominate and the fertile hyphae show verticillate branching. Trichosporiella may be compared to C. lucknowense because of the lateral, often sessile conidia. However, the absence of aerial hyphae, the little septate and narrow fertile hyphae, and the narrow basal scars of the blastic conidia of Trichosporiella all exclude confusion.

 

15. Chrysosporium anamorph of Rollandina vriesii Apinis - Fig. 17.

       Teleomorph: Rollandina vriesii Apinis - Trans. Br. mycol. Soc. 55: 501. 1970.

     Colonies attaining 25-30 mm diam in 14 days, white, felty, powdery, quite dense, raised to 2-3 mm with an indentation at the centre; margin defined, regular, slightly fimbriate; reverse cream-coloured. Hyphae little branched, hyaline, thin-walled. 2-4.5 µm wide, aerial hyphae fertile, submerged hyphae sterile. Racquet hyphae rare. Terminal and lateral conidia mostly sessile or on short protrusions, sometimes on side branches, solitary or subtended by another conidium, subhyaline. Smooth- and [p. 35] thin-walled, obovoid, pyriform or clavate, 1-celled, 3-6 x 2-3 µm, with broad basal scars (1-2 µm). Intercalary conidia absent. Chlamydospores absent.
     Growth temperatures: minimum 20-25°C, optimum 30°C, maximum 30-35°. Keratinolytic.

 fig. 17

Fig. 17
Chrysosporium anamorph of Rollandina vriesii. CBS 138.66.


Material examined

Living strain with the teleomorph and anamorph. - CBS 407.71 type strain of Rollandina vriesii isolated from skin and lungs of lizard, Ameira species, by G.A. de Vries, Baarn, Netherlands, 1968.


Discussion

     The little branched fertile hyphae with lateral conidia resemble the microconidium-bearing hyphae of Trichophyton and Microsporum, The anamorph differs from Chrysosporium lucknowense because the conidia are narrower and sometimes subtend one another, and because the growth rate is slower. Chrysosporium georgii is very similar in conidium morphology, but differs in that [p. 36] more conidia are borne per conidiogenous cell, the optimum growth temperature is 25°C rather than 30°C and, lastly, in that the teleomorph is an Arthroderma species.

 fig. 18

Fig. 18
Chrysosporium anamorph of Arthroderma curreyi. CBS 138.26.

 

16. Chrysosporium anamorph of Arthroderma curreyi Berk. - Fig. 18.

       Teleomorph: Arthroderma curreyi Berk. - Outl. Br. Fungol., p. 357. 1860.

     Colonies on sabouraud glucose agar attaining 30-35 mm diam in 14 days, white, felty, with much aerial mycelium, sometimes grooved at the centre, raised to up to 4 mm; margin regular, defined, slightly fimbriate; reverse pale to bright yellow or pale-brownish locally. Aerial and submerged hyphae may be fertile, hyaline, smooth- and thin-walled, mainly developing short side branches from primary hyphae, 2.5-4.5 (6) µm wide. Racquet hyphae present. Terminal and lateral conidia sessile or on very short protrusions or side branches, solitary, subhyaline, smooth-walled or slightly echinulate, thin-walled, obovoid to clavate, usually 1-celled, rarely 2-celled, 3-6 x 2-3 µm, with wide basal scars (1-2 µm). Intercalary conidia absent. Chlamydospores absent.
     Growth temperatures: minimum 5°C, optimum 15-25°C, maximum 25-30°C. Keratinolytic.


Material examined

Living strains with the teleomorph and anamorph. - CBS 138.26 = ATCC 13550 isolated by A. Nannizzi; CBS 130.70 isolated from soil by J.H. van Emden, Netherlands, 1969. [p. 37]


Discussion

     The anamorph closely resembles Chrysosporium georgii, the anamorph of Arthroderma ciferrii, except in that the conidia are echinulate and more elongate. Arthroderma curreyi differs from A. ciferrii because the ascomata are yellowish rather than brown, and the peridial appendages all spiral rather than straight and tapering as well as spiral.
     There is no type material of Arthroderma curreyi at either K or CMI. The name Arthroderma was first proposed by Currey (1854) and later validated by Berkeley (1860) with A. curreyi as type species. The descriptions given by Currey (1854) and Berkeley (1860) are brief but the drawings given by the first author are sufficient to recognise the fungus.

 fig. 19

Fig. 19
Chrysosporium anamorph of Arthroderma cuniculi. CBS 492.71.

 

17. Chrysosporium anamorph of Arthroderma cuniculi Dawson - Fig. 19.

       Teleomorph: Arthroderma cuniculi Dawson - Sabouraudia 2: 187. 1963. [p. 38]

     Colonies on Sabouraud glucose agar attaining 35-60 mm diam in 14 days, white, felty to fluffy, dense throughout, richly sporulating, raised to 1-2 mm and slightly folded at the centre; margin regular, defined, markedly fimbriate; reverse bright orange-yellow to brown at the centre, pale sulphur yellow outward. Hyphae hyaline, thin-walled; aerial hyphae mostly fertile and 2-6 µm wide; submerged hyphae sterile, occasionally slightly thick-walled, sometimes contorted, 3-7.5 µm wide. Racquet hyphae present. Terminal and lateral conidia sessile or on short protrusions or side branches, solitary, hyaline, smooth- and thin-walled, obovoid or clavate, 1-celled, occasionally 2-3-celled. 3-8 (16) x 2-3 µm. with relatively wide basal scars (1-2 µm). Intercalary conidia absent. Chlamydospores absent.
     Growth temperatures: minimum 5°C, optimum 20-25°C, maximum 30°C. Keratinolytic.


Material examined

Living strains with the teleomorph and anamorph. - CBS 492.71 = IMl 19624 = ATCC 18442 paratype strain (-) and CBS 495.71 = IMl 96245 = ATCC 18444 paratype strain (+), isolated from soil and hair from rabbit burrows by C.O. Dawson, Scotland, 1963.


Discussion

     The anamorph of Arthroderma cuniculi resembles Chrysosporium tropicum which has an Aphanoascus teleomorph, but the conidia have thinner walls, are more elongate, sometimes 2- to 3-celled and the branching of the fertile hyphae is more markedly orthotropic. The conidial morphology of A. cuniculi is practically identical to that of Pectinotrichum llanense Varsavsky & Orr, except in that the conidia of that species are never 2- to 3-celled and do not reach more than 6.5 µm in length. The ascoma of the former has uncinately branched dumbbell-shaped peridial hyphae with tuberculate, spine-like or long flexuous appendages (Dawson, 1963: Varsavsky & Orr, 1971). The fungus has only been found in association with rabbit burrows or rabbit fur (Dawson, 1963).

 

18. Chrysosporium anamorph of Pectinotrichum llanense Varsavsky & Orr - Fig. 20.

       Teleomorph: Pectinotrichum llanense Varsavsky & Orr - Mycopath. Mycol. appl. 43: 231. 1971.

     Colonies on Czapek agar with human hair attaining 5-15 mm diam in 14 days, growth scattered, especially concentrated around the hairs, white, fluffy, locally powdery; margin undefined; reverse white. Hyphae mostly aerial, mostly fertile, hyaline, thin-walled. Racquet hyphae present. Terminal and lateral conidia sessile or on short protrusions or side-branches, solitary, subhyaline, smooth- and thin-walled, [p. 39] obovoid or clavate, 1-celled, 4-6.5 x 2-3 µm with wide basal scars (1-1.5 µm). Intercalary conidia absent. Chlamydospores absent.
     Growth temperatures: minimum 20°C, optimum 25°C, maximum 25-30°C. Keratinolytic.

 fig. 20

Fig. 20
Chrysosporium anamorph of Pectinotrichum llanense. a. CBS 882.71; b. CBS 804.71.


Material examined

Living strains with the teleomorph and anamorph. - CBS 493.71 isolated from human skin by G.A. de Vries, Netherlands; CBS 804.71 = ATCC 22224 isolated from soil by J.G. Wright, Argentina; CBS 882.71 = ATCC 18921 type culture isolated from soil by G.F. Orr, Venezuela.


Discussion

     See Chrysosporium anamorph of Arthroderma cuniculi.

 

19. Chrysosporium lobatum Scharapov - Fig. 21.

       Chrysosporium lobatum Scharapov - Nov. Syst. niz. Rast. 15: 144. 1978. [p. 40]

 fig. 21

Fig. 21
Chrysosporium lobatum. a. CBS 666.78; b. CBS 274.77

     Colonies attaining 30-35 mm diam in 14 days, initially white, becoming pale green with a pinkish purple centre and finally pale grey with a dark grey centre, felty, powdery, dense and 2-3 mm high at the centre; margin defined, indented, slightly fimbriate; reverse cream with a greeny black centre. Hyphae hyaline, thin-walled, with repeated, orthotropic branching when fertile, 0.5-3 µm wide; aerial hyphae mostly fertile; submerged hyphae sometimes fertile, frequently contorted. Raquet hyphae absent. Terminal and lateral conidia developing more or less synchronously, borne in close proximity to one another, sessile or on short side branches, solitary, initially hyaline, smooth- and thin-walled, becoming reddish-brown to dark brown, echinulate and thick-walled, subglobose to obovoid, 1-celled, 2-4 x 1.5-3.5 µm, with wide basal scars (0.5-1.5 µm). Intercalary conidia rare, initially hyaline, smooth- and thin-walled, becoming reddish-brown to dark brown, echinulate and thick-walled, barrel-shaped, 1-celled, 3-4 x 2-3 µm. Chlamydospores subhyaline, smooth and thick-walled, subglobose to globose, 5-8 µm diam.
     Growth temperatures: minimum 10°C, optimum 25-30°C, maximum 35°C. Keratinolytic.


Material examined

CBS 666.78 = VKM-F 2120 type strain isolated from mouse fur, Rostov, U.S.S.R., 1969; CBS 274.77 [p. 41]
= UAMH 2515 isolated from soil by E. Varsaysky, Italy, 1961; CBS 275.77 = UAMH 3413 isolated as spinal fluid plate contaminant, Edmonton, Canada. 1971; CBS 276.77 = UAMH 3434 isolated from feather of house sparrow (Passer domesticus) by Z. Hubalek, Valtice, Czechoslovakia, 1970; CBS 278.77 = UAMH 1233, isolated from scrapings from human leg by J.W. Carmichael, 1962; CBS 624.79 = UAMH 2258, isolated from skin crust on chicken by I. Alteras, Bucharest, Rumania, 1964.


Discussion

     Chrysosporium lobatum is unlike other species of the genus in that the conidia are darkly pigmented and the fertile hyphae show numerous short orthotropic branches particularly visible after liberation of the conidia. Orthotropic branching is rarely seen in the Hyphomycetes; certain strains of Monodictys Hughes show similar, then, hyaline branches, but these are not fertile. In Myriodontium Samson & Polonelli (1975) the hyphae bear numerous blastic conidia on short, narrow branches (denticles) which are especially prominent after release of the conidia. Myriodontium differs in that the fertile hyphae are broader, the blastic conidia hyaline, smooth- and thin-walled and borne on shorter branches.

 fig. 22

Fig. 22
Chrysosporium synchronum. CBS 640.79, a. young conidia; b. mature conidia.[p. 42]

 

20. Chrysosporium synchronum van Oorschot, sp. nov. - Fig. 22.

     Coloniae post 7 dies 80 mm diam., albae, floccosae ad intricatae, copiose sporulantes, ad 2 mm altae; margo irregularis; reversum cremeum. Hyphae hyalinae, tenuitunicatae; hyphae aeriae seu steriles, parce ramosae, 3-4.5 µm latae, seu fertiles, plus minusve ramosae, 1.5-3 (-6) µm latae. Hyphae septum versus inflatae absunt. Conidia terminalia ac lateralia simul formantur in protrusionibus brevibus vel ramulis lateralibus modice angustatis, singula, hyalina, primum tenuitunicata, sed demum fere crassitunicata, levia vel exigue echinulata, obovoidea ad pyriformia, unicellularia, raro cellula tenuitunicata basilari supportata, primum 2-6 x 1.5-4 µm, matura 7.5-11 x 4-5.5 µm, cicatrice basilari 0.5-1.0 µm diam. Conidia intercalaria et chlamydosporae absunt
     Typus CBS 640.79, isolatus ex Agarico bisporo in Canada, a J.W. Carmichael, 1963.

     Colonies attaining 80 mm diam in 7 days, white, fluffy to felty, sporulating richly, up to 2 mm high; margin irregular; reverse cream-coloured. Hyphae hyaline, thin-walled; aerial hyphae either sterile, little branched and 3-4.5 µm wide, or fertile, much branched, little septate and 0.5-2.5 µm wide; submerged hyphae sterile, either much branched and 1.5-3 µm wide, or less branched and 1.5-6 µm wide. Racquet hyphae absent. Terminal and lateral conidia developing synchronously, borne on short protrusions or side branches which may be slightly conical initially, becoming more elongate and of regular width; conidia solitary, hyaline, initially thin-walled, later fairly thick-walled, smooth-walled or slightly echinulate, obovoid to pyriform, 1-celled though occasionally subtended by a thin-walled basal cell, 2-6 x 1.5-4 µm when young, 7.5-11 x 4-5.5 µm at maturity, with narrow basal scars (0.5-1 µm). Intercalary conidia absent. Chlamydospores absent.
     Growth temperatures: minimum 5°C, optimum 20-30°C, maximum 35°C. Not keratinolytic.


Material examined

CBS 640.79  = UAMH 1607 type strain, isolated from cultivated mushroom, Agaricus bisporos, by J.W. Carmichael, Edmonton, Canada, 1963.


Discussion

     The species is intermediate between Chrysosporium and Myceliophthora. Young conidia normally have broad points of attachment as in most species of Chrysosporium, but they inflate during maturation finally resembling those of M. thermophila (Apinis) van Oorschot. However, as the conidia are not borne on ampulliform swellings the species is placed in Chrysosporium. The marked synchronous maturation of the conidia is otherwise only known in C. sulfureum and to a lesser extent in C. lobatum. Hyphal branches bearing young conidia may be compared to those of Chrysosporium georgii, and the little septate fertile hyphae to those of C. xerophilum. [p. 43]

 fig. 23

Fig. 23
Chrysosporium pannicola. a. CBS 116.63; b. CBS 848.73.

 

21. Chrysosporium pannicola (Corda) van Oorschot & Stalpers, comb. nov. - Fig. 23.

       Capillaria pannicola Corda - Icon. Fung. 1: 10. 1837 (basionym) = Sporotrichum pannicola (Corda) Rabenhorst - Deutschl. Kryptogfl. 1: 78. 1844.
       Trichophyton evolceanui Randhawa & Sandhu - Mycopath. Mycol. appl 20: 225. 1963 = Chrysosporium evolceanui (Randhawa & Sandhu) Garg - Sabouraudia 4: 262. 1966.

     Colonies attaining 20-38 mm diam in 14 days, white, felty, dense and fluffy at the centre and thinner outwards, raised to 3 mm centrally; margin defined, regular, slightly fimbriate; reverse pale creamy brown. Hyphae hyaline, rarely branched when sterile, repeatedly branched when fertile, occasionally becoming slightly thick-walled and/or disarticulating; aerial hyphae mostly fertile, thin-walled, 1.5-5.0 µm wide, frequently producing swollen conidiogenous cells which are hyaline, smooth- and thick-walled, subglobose or somewhat irregular in shape, up to 12 µm wide and often chlamydospore-like; submerged hyphae mostly sterile, 1.5-7 µm wide, the wider being thick-walled, regular and straight, the narrower thin-walled, occasionally contorted and sometimes having numerous short side branches. Racquet hyphae present. Terminal and lateral conidia sessile or on short, nearly conical protrusions, solitary, occasionally subtended by another conidium, subhyaline, initially smooth-walled, nearly all becoming echinulate at maturity, fairly thick-walled, obovoid to clavate, 1-celled, 6-11 x 3.5-4.5 µm, with fairly wide basal scars (1-3 µm). [p. 44] Intercalary conidia less abundant, solitary, subhyaline, smooth-walled or echinulate, fairly thick-walled, barrel-shaped to ellipsoid, 4-8 x 2-3 µm. Chlamydospores absent.
     Growth temperatures: minimum 20°C, optimum 25°C, maximum 35°C. Keratinolytic.


Material examined

Herbarium specimen. - Capillaria pannicola, type, PRM, on decayed cloth.

Living strains. - CBS 116.63 = ATCC 22400 type strain of Trichophyton evolceanui, isolated from soil by R.S. Sandhu, Meerut, India, 1961; CBS 848.73 isolated from dog by M. Hajsig, Zagreb. Yugoslavia; CBS 277.77 sent in by G.F. Orr, 1961.


Discussion

     Randhawa & Sandhu (1963) placed the species in Trichophyton Malmsten on the basis of structures which they considered to be macroconidia. However, their type strain as well as their description and illustration show that the said macroconidia were actually irregular, swollen hyphal cells.
     Chrysosporium pannicola differs from C. tropicum by having a less dense colony, more markedly echinulate conidia and often swollen hyphal cells. The species also resembles C. indicum but may again be distinguished by the more echinulate and broader lateral conidia. After continued subculturing the conidia of C. pannicola may cease to be echinulate.
     The swollen conidiogenous cells may be regarded as rudimentary ampulliform swellings similar to those which develop in Myceliophthora fergusii (Klopotek) van Oorschot. As the lateral conidia have broad bases the species is classified in Chrysosporium, though it is regarded as intermediate between this genus and Myceliophthora.

 

22. Chrysosporium indicum (Randhawa & Sandhu) Garg.- Fig. 24.

       Trichophitton indicum Randhawa & Sandhu - Mycopath. Mycol. appl. 20: 227. 1963 [non Trichophyton indicum (Cast.) Nannizzi - Tratt. Micopat. umana 4: 186. 1934] = Chrysosporium indicum Garg - Sabouraudia 4: 262. 1966.

       Teleomorph: Apianoascus terreus (Randhawa & Sandhu) Apinis - Mycopath. Mycol. appl. 35: 99. 1968.

 fig. 24

Fig. 24
Chrysosporium indicum. a. CBS 117.63: h. CBS 472.76.

     Colonies attaining 40-45 mm diam in 14 days, white, felty, thin, denser and up to 2 mm high at the centre; margin defined, regular, fimbriate; reverse cream-coloured. Hyphae hyaline, smooth- and thin-walled, only rarely branched when sterile, more frequently branched when fertile; aerial hyphae fertile or infertile [p. 45] (sporulation often being poor), 1.5-5 µm wide; submerged hyphae mostly sterile, straight, 1-4.5 µm wide, the narrower often contorted. Racquet hyphae present. Terminal and lateral conidia sessile or on short protrusions or slightly swollen lateral branches, solitary, occasionally subtended by another conidium, subhyaline, smooth to slightly echinulate, thin-walled, obovoid to ellipsoid, often cymbiform, 1-celled, 3.5-7.5 x 1.5-3 µm, with fairly wide basal scars (1-1.5 µm). Intercalary conidia less abundant, solitary, subhyaline, smooth-walled to slightly echinulate, thin-walled, cylindrical to barrel-shaped, not always clearly distinguishable from disarticulating hyphal segments, 6-12 x 2-3.5 µm. Chlamydospores absent.
     Growth temperatures: minimum 20°C, optimum 25-30°C, maximum 40°C. Keratinolytic.


Material examined

Living strains with only the anamorph. - CBS 117.63 = ATCC 22401 = IMI 147544, type strain isolated from soil by R.S. Sandhu, India; CBS 472.76 = ATCC 26360 = UAMH 2733 isolated from soil by C. Magalhaes, Moçambique; CBS 181.77 = IMI 145695 isolated by H.F. Müller, F.R.G., 1969; CBS 134.78 and 168.78 both isolated from soil baited with hairs by H.M. Abdel-Fattah, Egypt, 1977.

Living strains with the teleomorph and anamorph. - CBS 503.63 isolated from forest soil, sent in by H.S. Randhawa, Darjeeling, India; CBS 504.63 isolated from garden soil sent by H.S. Randhawa, Delhi, India; CBS 342.64 = ATCC 16413 = IMl 108689 type culture of Keratinophyton terreum [p. 46] (Aphanoascus terreusi), isolated from lawn soil, sent in by H.S. Randhawa, India; CBS 379.72 isolated trom rabbit ground by De Buys Scott, Bloemtontein, South Africa.


Discussion

     The species was originally placed in Trichophyton Malmsten by Randhawa & Sandhu (1963) on account of the presence of swollen cells which they regarded as macroconidia. Similar cells can still be found in the type culture CBS 117.63. Neither this culture nor any of the other strains examined developed multi-septate macroconidia, characteristic for Trichophyton, on media such as malt or Sabouraud maltose agars which normally promote sporulation in Trichophyton species.
     Chrysosporium indicum is distinguished from other Chrysosporium species by the narrow, cymbiform terminal and lateral conidia which are frequently borne on slightly swollen lateral branches. Chrysosporium pannicola also shows this type of conidiogenesis but has larger, more echinate conidia.

 

Myceliophthora Cost.

       Myceliophthora Cost. - C. r. hebd. Séanc. Acad. Sci., Paris 114: 850. 1892.

       Teleomorph genera: Arthroderma, Ctenomyces, Thielavia, Corynascus.

     Colonies spreading, white, cream-coloured, pale yellow, pale brown or dark green, fluffy, felty, floccose or powdery. Hyphae hyaline, smooth-walled, septate, with irregular more or less orthotropic branching. Blastic conidia borne terminally or laterally all over the hyphae, sessile or on short protrusions or side branches, frequently with 1-4 blastic conidia on an ampulliform swelling, solitary, sometimes in chains of 2-4, subhyaline or pale yellow, pale brown, or reddish brown, smooth-walled or verruculose to verrucose, globose, subglobose, ellipsoid or pyriform, 1-celled, with narrow basal scars. Intercalary conidia absent. Chlamydospores absent. Thermotolerant.

     Type species: Myceliophthora lutea Cost.


Discussion

     The species of Myceliophthora nearly all produce ampulliform swellings, a feature absent in Chrysosporium. Colony growth is fast and dense, sporulation often being good between 30 and 40°C. Dehiscence of blastic conidia is rhexolytic, though the remains of the former supporting cell are distinguished with difficulty when the basal scar is very small.
     The morphology of Emmonsia and Zymonema species grown at 24°C resembles [p. 47] that of Myceliophthora except in that the blastic conidia are much smaller, unpigmented and thin-walled. However, Emmonsia and Zymonema are dimorphic, producing adiaspores and budding cells respectively at 35 to 40°C. In addition, the latter genera are pathogenic whilst Myceliophthora species, most commonly isolated from soil or mushroom compost, are sometimes keratinolytic but not pathogenic. Beniowskia sphaeroidea (Kalchbr. & Cooke) Mason may produce blastic conidia on ampulliform swellings but the fertile hyphae are much wider than those of Myceliophthora species and, further, B. sphaeroidea is a sporodochial fungus (Samson & Polonelli. 1978).
     Myceliophthora was placed in synonymy with Chrysosporium by Carmichael (1962) but was reintroduced by von Arx (1973).

 

 

Key to the species

 

1a.

 

Conidia verrucose

2

 

1b.

 

Conidia smooth-walled

6

 

 

2a.

 

Conidia thin-walled

M. anamorph of Corynascus sepedonium

 

2b.

 

Conidia thick-walled

3

 

 

3a.

 

Conidia subhyaline: ampulliform swellings rare

M. anamorph of Arthroderma tuberculatum

 

3b.

 

Conidia lightly pigmented; ampulliform swellings common

4

 

 

4a.

 

Conidia mostly less than 8 µm long

M. thermophila

 

4b.

 

Conidia mostly more than 8 µm long

5

 

 

5a.

 

Conidia reaching up to 23 µm in length, reddish brown

M. anamorph of Ctenomyces serratus

 

5b.

 

Conidia reaching up to 13 µm in length. pale yellow or pale brown

M. vellerea

 

 

6a.

 

Conidia globose or subglobose

M. anamorph of Corynascus novoguineensis

 

6b.

 

Conidia mostly obovoid or clavate

7

 

 

7a.

 

Ampulliform swellings usually globose; fertile hyphae not reaching 6 µm in width: colonies felty or powdery: optimum growth temperature 30°C

M. lutea

 

7b.

 

Ampulliform swellings usually elongate; fertile hyphae often reaching 6 µm in width; colonies floccose; optimum growth temperature 45°C

M. fergusii

 

 

23. Myceliophthora vellerea (Sacc. & Speg.) van Oorschot, comb. nov. - Fig. 25, Plate 1.

       Sporotrichum vellereum Sacc. & Speg. - Michelia 2: 287. 1882 (basionym).
       Chrysosporium asperatum Carmichael - Can. J. Bot. 40: 1153. 1962.
       Sporotrichum ochrokeratinophilum Matsushima - Icones Microf. a Mats. lect., p. 143. 1975. [p. 48]

 fig. 25

Fig. 25. Myceliophthora vellerea. a. CBS 169.62; b. CBS 478.76.

     Colonies on oatmeal agar attaining 35-40 mm diam in 14 days, initially white and remaining so or becoming pale brown, fluffy to felty, 1-2 mm in height; margin not well defined, ir+regular, slightly fimbriate; reverse cream-coloured. Hyphae thin-walled, 1-2.5 µm wide; aerial hyphae mostly fertile, the fertile being more densely branched; submerged hyphae sterile. Racquet hyphae absent. Conidia borne on short, narrow protrusions or side branches, or 1-2 borne on ampulliform swellings, solitary or in chains of 2-3 conidia separated by narrow, short hyphal segments, initially subhyaline and smooth- and thin-walled, normally becoming pale yellow or pale brown, verruculose and thick-walled, subglobose, pyriform or ellipsoid, 4-13 x 3-9 µm, mature conidia mostly 8-13 x 7-9 µm, with narrow basal scars (0.5-1.5 µm wide).
     Growth temperatures: minimum 10°C, optimum 20-25°C, maximum 30°C. Keratinolytic.


Material examined

Herbarium specimen. - Sporotrichum vellereum type (PD), on hairs of dog, Conegliano, Italy. 1878.

Living strains. - CBS 169.62 = ATCC 14801 = IMI 94289 = UAMH 560 = IFO 7582 type strain [p. 49] of Chrysosporium asperatum, isolated from soil, Alberta, Canada, 1956; CBS 487.76 and 479.76 isolated from soil by H.M. Yusef, Alexandria, Egypt.


Discussion

     The pale brown colour of the mycelium may be lost after repeated subculturing, conidium production occasionally being reduced. Condiogenesis may show some variation within a single strain in that conidia may or may not be subtended by a narrow hyphal segment. In young colonies smooth-walled, hyaline conidia are sometimes seen in series of up to three, interconnected by narrow hyphal segments. Side branches bearing single conidia are occasionally slightly swollen, a feature also seen in Chrysosporium pannicola and C. indicum.

 fig. 26

Fig. 26
Myceliophthora anamorph of Arthroderma tuberculatum. a. CBS 473.77; b. CBS 217.62.

 

24. Myceliophthora anamorph of Arthroderma tuberculatum Kuehn - Fig. 26.

       Chrysosporium tuberculatum (Kuehn) Dominik - Zesz. nauk. wyzsz. Szk. roln. Szczec. 24: 59. 1968.

       Teleomorph: Arthroderma tuberculatum Kuehn - Mycopath. Mycol. appl. 13: 190. 1960. [p. 50]

     Colonies attaining 40-50 mm diam in 14 days, white, fluffy, with much sterile aerial mycelium, 1-3 mm high; margin fairly defined, regular, fimbriate; reverse cream-coloured. Hyphae thin-walled, branching infrequent, 0.5-4.5 µm wide; aerial hyphae partly fertile, submerged hyphae sterile. Racquet hyphae absent. Conidia sessile or on short protrusions or borne singly on ampulliform swellings, solitary, subhyaline, initially smooth- and thin-walled, usually becoming verrucose and thick-walled, subglobose, pyriform or obovoid, occasionally with a medial indentation. 4-10 x 5-13 µm, with mostly narrow basal scars (1-2 µm).
     Growth temperature: minimum 10°C, optimum 25°C, maximum 30°C. Keratinolytic.


Material examined

Living strains with the teleomorph and anamorph. - CBS 217.62 isolated from soil by T. Pinto Ribeiro. Lisbon, Portugal; CBS 312.65 (+). 313.65 (-) and CBS 413.65 (cross between 312.65 and 313.65) all isolated from soil in bird cages by R.S. Pors, California, U.S.A., 1964; CBS 418.66 isolated by E. Varsavsky, Argentina. 1963; CBS 474.77 (+) = UAMH 873 type strain of Arthroderma tuberculatum, isolated from feathers of Turdus americanus, Illinois, U.S.A., 1954.


Discussion

     Unlike other Myceliophthora species the ampulliform swellings are quite rare and do not bear more than one conidium connected by a fairly broad or narrow base. The conidia themselves are almost identical to those of M. vellerea and also resemble those of the Myceliophthora state of Ctenomyces serratus Eidam. The measurements given by Kuehn (1960) for the conidia were 8.8-14.3 x 13.2-18.7 (24.2) µm, exceeding those given by Carmichael (1962) and those observed in the type and other strains listed above. It seems rather peculiar that the teleomorph is an Arthroderma; nearly all other known anamorphs of Arthroderma develop mostly smooth-walled, clavate or obovoid thallic (micro)conidia and are classified in Trichophyton or Chrysosporium.

 

25. Myceliophthora anamorph of Ctenomyces serratus Eidam - Fig. 27.

       Chrysosporium serratum (Eidam) Dominik - Zesz. nauk. wyzsz. Sek. roln. Szczec. 24: 59. 1967.

       Teleomorph: Ctenomyces serratus Eidam in Cohn - Beitr. Biol. Pfl. 3: 274. 1880.

     Colonies attaining 35-40 mm diam in 14 days, initially white, usually becoming pale brown and sometimes exuding pale brown droplets, felty, flat; margin irregular, not defined; reverse cream-coloured. Hyphae hyaline, smooth- and thin-walled, rarely branched when sterile, 1-3 µm wide, more branched when fertile; aerial hyphae mostly fertile; submerged hyphae sterile. Racquet hyphae absent. Conidia occasionally on short protrusions but usually 1-2 borne on ampulliform swellings, [p. 51] solitary or in series of up to 3 conidia separated by short, narrow hyphal segments, initially subhyaline, thin- and smooth-walled, soon becoming reddish brown, verrucose and thick-walled, ellipsoid, 5-23 x 3.5-12 µm, mature conidia usually 12-23 x 10.5-12 µm, with narrow basal scars (approx. 1 µm).
     Growth temperatures: minimum 10°C, optimum 30°C, maximum 30-35°C. Keratinolytic.

 fig. 27

Fig. 27
Myceliophthora anamorph of Ctenomyces serratus. a. CBS 187.61; b. CBS 223.64.


Material examined

CBS 187.61 = IMl 86199 = ATCC 15504 neotype of Ctenomyces serratus, isolated by D.M. Griffin from soil, Australia; CBS 544.63 = ATCC 15503 = NRRL A-11828 isolated by C. Dickinson from soil, England; CBS 221.64 = ATCC 15502 = NRRL A-13807 isolated by H.S. Randhawa from soil, India; CBS 222.64 (+) = ATCC 15536 and CBS 223.64 (-) = ATCC 15537 isolated from soil by E. Varsavsky, Argentina.


Discussion

     The anamorph of Ctenomyces serratus strongly resembles Myceliophthora [p. 52] vellerea but may be distinguished by its flatter, browner colony and the more elongate, darker conidia. The conidia differ from those of Arthroderma tuberculatum which are more globose, subhyaline and borne infrequently and singly on ampulliform swellings.
     The conidia attributed to Ctenomyces serratus by Eidam (1880) in the original description belonged to Arthroderma curreyi (Benjamin, 1956; Kuehn, 1958; Frey & Griffin, 1961). The anamorph was first accurately described by Frey & Griffin (1961) but not related to any hyphomycetous genus. Carmichael (1962) as well as Dominik (1967) regarded the anamorph as a Chrysosporium. Ctenomyces serratus is most commonly isolated from feathers of wild and domestic birds, as well as from soil by means of hair bait methods. The fungus showed no signs of pathogenicity after inoculation into guinea pigs or mice (Frey & Griffin, 1961).

 fig. 28

Fig. 28
a. Myceliophthora fergusii. CBS 406.69; b. M. lutea. CBS 379.76; c. M. thermophila. CBS 202.75. [p. 53]

 

26. Myceliophthora thermophila (Apinis) van Oorschot - Fig. 28c.

       Sporotrichum thermophilum Apinis - Nova Hedwigia 5: 74. 1962 = Chrysosporium thermophilum (Apinis) Klopotek - Arch. Mikrobiol. 98: 366. 1974 = Myceliophthora thermophila (Apinis) van Oorschot - Persoonia 9: 403. 1977.

       Teleomorph: Thielavia heterothallica Klopotek - Arch. Microbiol. 107: 223. 1976.


Discussion

     The fungus differs from the foregoing species by the dark colony and smaller, mostly obovate conidia.
     The species was fully described by van Oorschot (1977) and also by Klopotek (1974). Myceliophthora thermophila is known from pasture soil (Apinis, 1963), molding open-stack alfalfa ensilage, fir and spruce pulpwood chips (Semeniuk & Carmichael, 1966) and wheat straw compost (Yung-Chang & Hudson, 1967).

 

27. Myceliophthora lutea Cost. - Fig. 28b.

       Myceliophthora lutea Cost. - C. r. hebd. Séanc. Acad. Sci., Paris 114: 850. 1892 = Scopulariopsis lutea (Cost.) Tubaki - Nagaoa 5: 29. 1955 = Chrysosporium luteum (Cost.) Carmichael - Can. J. Bot. 40: 1158. 1962.


Discussion

     Myceliophthora lutea differs from M. thermophila by the yellow colony, smooth, subhyaline conidia and a maximum growth temperature of 40°C rather than 45°C. Full descriptions were given by van Oorschot (1977) and also Carmichael (1962). The fungus was originally described by Costantin (1892) as the 'vert de gris' disease of cultivated mushrooms. Other sources of isolation include soil, hay and dust.
     Even though the fungus isolated by Tubaki (1955) was isolated from cultivated mushrooms, the strain depicted shows the verticillate branching and series of alternate intercalary conidia characteristic of Geomyces pannorum.

 

28. Myceliophthora fergusii (Klopotek) van Oorschot - Fig. 28a.

       Chrysosporium fergusii Klopotek - Arch. Mikrobiol. 98: 366. 1974 = Myceliophthora fergusii (Klopotek) van Oorschot - Persoonia 9: 406. 1977.

       Teleomorph: Corynascus thermophilus (Fergus & Sinden) Klopotek - Arch. Mikrobiol. 98: 366. 1974. [p. 54]


Discussion

     Unlike Myceliophthora lutea, M. fergusii has a known associated teleomorph, Corynascus thermophilus. The anamorph differs from M. lutea in that the mycelium is pinkish-cream and floccose and the fungus is markedly thermophilic, with an optimum growth temperature of 45°C and a maximum of 55°C. Fertile hyphae are generally broad (up to 6 µm) compared to other species in the genus. Full descriptions were given by van Oorschot (1977), Klopotek (1974) and Hedger & Hudson (1970). Source of isolation is usually mushroom or wheat straw compost.

 fig. 29

Fig. 29
Myceliophthora anamorph of Corynascus novoguineensis. CBS 359.72.

 

29. Myceliophthora anamorph of Corynascus novoguineensis (Udagawa & Horie) v. Arx - Fig. 29.

       Teleomorph: Corynascus novoguineensis (Udagawa & Horie) v. Arx - Proc. K. ned. Akad. Wet., Ser. C., 76: 292. 1973.

     Colonies attaining 45-55 mm diam in 14 days, white, felty, slightly fluffy, 2-3 mm high at the centre; margin not well defined, fimbriate; reverse pale yellow to cream-coloured. Aerial hyphae mostly fertile, 1-3 µm wide; submerged hyphae sterile, 1.5-4 µm wide, occasionally slightly contorted. Racquet hyphae absent. Conidia [p. 55]sessile or on short protrusions, 1-3 often borne on globose or elongate, frequently catenate, ampulliform swellings, solitary, subhyaline, smooth- and thick-walled, usually globose, sometimes subglobose, rarely ellipsoid, 4-9 x 4-8 µm, with very narrow basal scars (up to 1 µm).
     Growth temperatures: minimum 15°C, optimum 25-30°C, maximum 40°C. Keratinolytic.


Material examined

Living strain with the teleomorph and anamorph. - CBS 359.72 type strain of Thielavia novoguineensis Udagawa & Horie, isolated from soil, New Guinea.


Discussion

     The globose conidia and the catenate nature of the ampulliform swellings distinguish this fungus from Myceliophthora fergusii, the anamorph of Corynascus thermophilus. Udagawa & Horie (1972) described the anamorph of C. novoguineensis as a Chrysosporium but the blastic ontogeny of the conidia and the abundant ampulliform swellings place the fungus in Myceliophthora.

 fig. 30

Fig. 30
Myceliophthora anamorph of Corynascus sepedonium. CBS 440.51. [p. 56]

 

30. Myceliophthora anamorph of Corynascus sepedonium (Emmons) v. Arx - Fig. 30.

       Chrysosporium sepedonium (Emmons) Dominik - Zesz. nauk. wyzsz. Szk. roln. Szcec. 24: 58. 1968.

       Teleomorph: Corynascus sepedonium (Emmons) v. Arx - Stud. Mycol. 8: 21. 1975.

     Colonies attaining 80-90 mm diam in 14 days, initially white, sometimes becoming pale brown, fluffy to felty, fairly dense throughout, 1-2 mm high; margin defined, regular, slightly fimbriate; reverse cream-coloured, occasionally slightly pale brown at the centre. Hyphae thin-walled; aerial hyphae fertile, 1-1.5 µm wide; submerged hyphae sterile, 1-3.5 µm wide. Racquet hyphae absent. Conidia borne singly on short, narrow protrusions or 1-2 may be borne on globose, frequently catenate ampulliform swellings, subhyaline to pale brown, initially smooth-walled and thin-walled, usually becoming verruculose and slightly thick-walled, mostly globose, occasionally obovoid, 4-11 (17) x 4-11 µm, with very narrow basal scars (up to 1 µm).
     Growth temperatures: minimum 10°C, optimum 20-30°C, maximum 40-45°C. Keratinolytic.


Material examined

Living strain with the teleomorph and anamorph. - CBS 340.33 type strain of Thielavia sepedonium Emmons, isolated from skin of human foot, U.S.A., 1932; CBS 415.48, isolated from cotton rope by W.L. White, India, 1944; CBS 541.50, isolated from pea seed by J.W. Groves, Canada; CBS 440.51 = IMl 45476 isolated from soil by J.H. Warcup, U.K., 1950; CBS 412.52 isolated from soil by P. Negroni, Argentina; CBS 295.56 isolated from buried cables by J.H. Hueck, Delft, Netherlands, 1955; CBS 387.65, isolated from soil, Baarn, Netherlands; CBS 632.67 = VKM-F 1142 type strain of Thielavia lutescens Kamyschko; CBS 111.69 = IMI 136625 isolated from soil by B.S. Mehrotra, India, 1968; CBS 213.74 isolated from sandy soil by J.A. von Arx, Senegal.


Discussion

     The conidial morphology strongly resembles that of the anamorph of Corynascus novoguineensis except in that the conidia are less thick-walled and verruculose. The anamorph of C. sepedonium was regarded as a Sepedonium by Emmons (1932) and Malloch & Cain (1973), and as a Chrysosporium by Mouchacca (1973) and Dominik (1967). The blastic conidia and ampulliform swellings place the fungus in Myceliophthora.

 

Emmonsia Cif. & Montemartini

       Emmonsia Cif. & Montemartini - Mycopath. Mycol. appl. 10: 314. 1959. [p. 57]

     Colonies at 24°C restricted or slightly spreading, white, rarely pale brown, felty or cottony, sometimes tufted. Hyphae hyaline, smooth- and thin-walled, branching irregular, more or less orthotropic. Blastic conidia borne terminally, mostly laterally all over the hyphae, sessile or on short protrusions or side branches, occasionally 1-2 blastic conidia borne on an ampulliform swelling, solitary or sometimes in chains of 2-3, hyaline, smooth-walled or echinate, subglobose, globose, pyriform or cuneiform, 1-celled, with very narrow basal scars. Intercalary conidia absent. Dimorphic, producing adiaspores at 37°C to 40°C in vitro and in vivo.

     Type species: Emmonsia parva (Emmons & Ashburn) Cif. & Montemartini

 

Discussion

     The conidiogenesis of Emmonsia resembles that of Myceliophthora, both genera producing blastic conidia laterally or terminally on the hyphae or on ampulliform swellings. However, Emmonsia differs in that the blastic conidia are smaller and hyaline rather than subhyaline or pigmented, the growth rate slower, colonies much less dense, and the strains dimorphic. The two varieties of Emmonsia parva are pathogenic to humans and particularly rodents, while the species of Myceliophthora are not pathogenic.
     At 24°C the pathogenic genera Emmonsia, Histoplasma Darling, Paracoccidioides Almeida and Zymonema show a similar morphology and grow poorly. Histoplasma produces undifferentiated, slightly echinulate blastic conidia which reach 2-4 µm diam and are borne laterally on the hyphae but not on ampulliform swellings. However, Histoplasma also develops macroconidia which are spherical, 8-14 µm diam, with finger-like ornamentation. At 37°C in vitro or in vivo Histoplasma produces budding cells with narrow connection points rather than adiaspores. Paracoccidioides forms smooth-walled conidia with no ampulliform swellings at 24°C, and spherical cells which bud multilaterally at 37°C in vitro or in vivo. Zymonema is almost identical to Emmonsia at 24°C except in the more restricted growth of the colony. At 37°C in vitro or in vivo Zymonema shows budding cells with broad connection points.
     Emmonsia parva var. parva and var. crescens cause adiaspiromycosis, a pulmonary mycosis characterised by growth of inhaled blastic conidia. The disease caused by Histoplasma species, histoplasmosis, is an intracellular mycosis which is also usually respiratory in origin but may soon spread to the lymphatic tissues, spleen, liver, adrenals, kidneys, skin, central nervous system etc. Paracoccidioides causes a chronic often fatal mycosis which is first evident in the lungs, subsequently disseminating to the visceral organs, the skin and lymphthic system. Zymonema is again first recognised as a respiratory infection which later affects the bones and skin.
     Emmonsia was placed in synonymy with Chrysosporium by Carmichael (1962) but this was refuted by von Arx (1973a) on the basis of the blastic conidia and thick-walled chlamydospore-like cells (adiaspores). [p. 58]

 fig. 31

Fig. 31
a. Emmonsia parva var. crescens. CBS 508.78: b. E. parva var. parva. CBS 177.60.

 

31. Emmonsia parva (Emmons & Ashburn) Cif. & Montemartini var. parva - Fig. 31b.

       Haplosporangium parvum Emmons & Ashburn - Publ. Hlth Rep., Wash. 57: 1719. 1942 = Emmonsia parva (Emmons & Ashburn) Cif. & Montemartini - Mycopath. Mycol. appl. 10: 314. 1959 = Chrysosporium parvum (Emmons & Ashburn) Carmichael var. parvum (Emmons & Ashburn) Carmichael - Can. J. Bot. 46: 1164. 1962.

     Colonies attaining 25-40 mm diam in 14 days, white, occasionally pale brown at the centre, felty, thin, sometimes tufted or with slight radial grooves, 1-2 mm in height; margin usually regular, slightly fimbriate; reverse cream-coloured to pale brown. Hyphae hyaline or rarely pale brown, occasionally locally swollen, 1-3 µm wide; aerial hyphae partly fertile, submerged hyphae sterile, a few contorted. Racquet hyphae absent. Conidia sessile or on short protrusions or thin side branches, 1-2 conidia may be borne on ampulliform swellings or on irregularly swollen hyphal cells, solitary or in chains of 2-3, hyaline, smooth-walled or echinulate, thin-walled, globose to somewhat cuneiform, 2-4 x 2-5 µm.
     Growth temperatures: minimum 5-10°C, optimum 20-25°C, maximum 40°C. Not keratinolytic.
     At 37°C the blastic conidia swell and the hyphae become irregularly swollen and [p. 59] contorted. At 40°C the blastic conidia develop into uninucleate adiaspores 10-25 µm diam with walls up to 2 µm thick.


Material examined

CBS 204.48 and 205.58 isolated from rodent lung by C.W. Emmons, U.S.A.; CBS 242.52 isolated from mouse lung, U.S.A.; CBS 178.60 = ATCC 14051 isolated from ground squirrel lung, Arizona, U.S.A.; CBS 273.77 = UAMH 2304 isolated by C.W. Emmons, Kansas, U.S.A.; CBS 475.77 = ATCC 10785 isolated from rodent lung, U.S.A., 1942.


Discussion

     The pulmonary mycosis caused by the species is known as adiaspiromycosis, or haplomycosis. It was initially only known in rodents, the first human case being reported in 1964 by Doby-Dubois et al. Patients with adiaspiromycosis normally also suffer from a primary disease such as tuberculosis, aspergillosis or silicosis (Emmons et al., 1977). There are no reports suggesting that the disease is contagious, the source of infection probably lying in the soil and air. Most authors agree that the adiaspores are actually enlarged blastic conidia which when inhaled enlarge in the alveoli or bronchioles of the host, thus hindering normal pulmonary functioning.
     Transmission electronmicroscopy has confirmed an earlier report (Emmons & Jellison, 1960) that certain strains have multinucleate adiaspores (Kodousek et al., 1972; Watts et al., 1975). Uninucleate adiaspores are produced by those strains which develop the structures at 40°C, multinucleate adiaspores are produced at 37°C. The two types are indistinguishable at 24°C, even so they were regarded as separate species by Emmons & Jellison (1960). Carmichael (1962) placed the species in Chrysosporium, separating them only as varieties. Those strains which develop multinucleate adiaspores are here classified as Emmonsia parva var. crescens comb. nov.

 

32. Emmonsia parva (Emmons & Ashburn). Cif. & Montemartini var. crescens (Emmons & Jellison) van Oorschot, comb. nov. - Fig. 31a.

       Emmonisia crescens Emmons & Jellison - Ann. N.Y. Acad. Sci. 89: 96. 1960 (basionym) = Chrysosporium parvum (Emmons & Ashburn) Carmichael var. crescens (Emmons & Jellison) Carmichael - Can. J. Bot. 40: 1164. 1962.

     Emmonsia parva var. crescens develops multinucleate adiaspores, 200-700 µm diam with up to 70 µm thick walls at 37°C, but does not grow at 40°C (Emmons et al. 1977).


Material examined

CBS 177.60 = ATCC 13704 type strain isolated from Arvicola terrestris lung by W. Jellison, Norway, [p. 60] 1959; CBS 642.77 isolated from human brain by L. Haley, Atlanta, U.S.A.; CBS 508.78 = ATCC 24951 cause of adiaspiromycosis, isolated by R. Kodousek; CBS 509.78 = ATCC 32539 and CBS 510.78 = ATCC 32540 isolated from lung of prairie polecat by K. Krivance, U.S.A., 1975.


Discussion

     CBS 642.77 is the first report of an Emmonsia from brain tissue rather than from lungs. The symptoms of the disease are identical in both varieties, in vivo distinction being on adiaspores size and the number of nuclei. Emmons & Jellison (1960), Emmons et al. (1977) and Watts et al. (1975) are of the opinion that the adiaspores do not germinate, reproduce or disseminate in the host tissue, while Kodousek & Hejtmánek (1972) reported that endosporulation may be induced in vitro in E. parva var. crescens. Emmons & Jellison (1960), Carmichael (1962) and other authors made no reference to the rough-walled nature of the adiaspores, but this was demonstrated with scanning electron microscopy by Hejtmánek (1976).

 

Zymonema de Beurmann & Gougerot

       Blastomyces Gilchrist & Stokes - J. exp. Med. 3: 76. 1898 (non Blastomyces Cost. & Rolland - Bull. Soc. mycol. Fr. 4: 153. 1888) = Zymonema de Beurmann & Gougerot - Trib. Med., Paris 42:503. 1909 (name change) = Blastomycoides Castell. - Proc. R. Soc. Med. 21: 451. 1928 (name change) = Gilchristia Redaelli & Cif. - J. trop. Med. Hyg. 37: 281. 1934 (name change).

       Teleomorphic genus: Ajellomyces.

     Colonies restricted, white, butyrous, with tufts of aerial hyphae. Hyphae hyaline, thin- and smooth-walled, with irregular, more or less orthotropic branching. Blastic conidia borne terminally or laterally all over the hyphae, sessile or on short branches, sometimes 1-2 on ampulliform swellings, hyaline, thin-walled, pyriform, usually subglobose or globose, 1-celled. Intercalary conidia absent. Chlamydospores absent. Dimorphic, producing a budding phase at 37°C or in vivo.

     Type species: Zymonema dermatitides (Gilchrist & Stokes) Dodge


Discussion

     Zymonema is compared to Emmonsia, Histoplasma and Paracoccidioides on page 57. Only the species of Histoplasma and Zymonema have known teleomorphs in the Gymnoascaceae. Histoplasma capsulatum Darling has an associated Emmonsielle capsulate Kwon-Chung, and Zymonema dermatitidis and Ajellomyces dermatitidis McDonough & Lewis. The + and - mating types of A. dermatitidis are equally pathogenic, while in E. capsulata the - mating type is a more frequent clinical isolate (Emmons et al., 1977; Kwong-Chung et al., 1974).
     Blastomycoides Castell. is illegitimate because it was proposed as a nomen provisorum [p. 61]and included the type species of Coccidioides Stiles, viz. C. immitis Rixford & Gilchrist. Gilchristia Redaelli & Cif. was based on G. dermatitides (Gilchrist & Stokes) Redaelli & Cif. (= Zymonema dermatitidis) making it an obligate synonym of the earlier Zymonema. Zymonema remains the earliest valid name for the genus.

 fig. 32

Fig. 32
Zynemona dermatitidis. CBS 674.68. a. Sabouraud glucose agar, 24ºC; b. Sabouraud glucose agar, 37ºC; c. Brain heart infusion agar, 37ºC.

33. Zymonema dermatitidis (Gilchrist & Stokes) Dodge - Fig. 32.

       Blastomyces dermatitidis Gilchrist & Stokes - J. exp. Med. 3: 76. 1898 (basionym) = Oidium dermatitidis (Gilchrist & Stokes) Ricketts - J. med. Res. 6: 373. 1901 = Cryptococcus dermatitidis Brumpt in Castell. & Chalmers - Man. trop. Med., ed. 2, p. 769. 1913 = Mycoderma dermatitidis (Gilchrist & Stokes) Brumpt - Précis Parasitol. p. 934. 1913 = Blastomycoides dermatitidis (Gilchrist & Stokes) Castell. - Am. J. trop. Med. 8: 383. 1928 = Geotrichum dermatitidis (Gilchrist & Stokes) Basgal - Contrib. Estudo Blastomycoses pulmon., p. 156. 1931 = Torulopsis dermatitidis (Gilchrist & Stokes) Almeida- Ann. Fac. Med., S. Paulo 9: 76, 1933 = Endomyces dermatitidis (Gilchrist & Stokes) Moore - Ann. Mo. bot. Gdn 20: 106. 1933 = Gilchristia dermatididis (Gilchrist & Stokes) Redaelli & Cif. - J. trop. Med. Hyg. 37: 281. 1934 = Zymonema dermatitidis (Gilchrist & Stokes) Dodge - Med. Mycol. p. 168. 1935 = Chrysosporium dermatitidis (Gilchrist & Stokes) Carmichael - Can. J. Bot. 40: 1154. 1962.
       Cryptococcus gilchristi Vuill. in Guéguen - Champ. Paras., p. 108. 1904 (name change) = Zymonema gilchristi (Vuill.) de Beurmann & Gougerot Trib. med., Paris 42: 503. 1909 = Mycoderma gilchristi (Vuill.) Jannin - Les Mycodermas, p. 71. 1913.
       Glenospora gammeli Poll. & Nannizzi in Blankenhorn & Gammel - J. clin. Invest. 4: 483. 1927 = Trichosporium gammeli (Poll. & Nannizzi) Dodge - Med. Mycol., p. 792. 1935.
       Blastomycoides tulanensis Castell. - Proc. R. Soc. Med. 21: 451. 1928 = Monosporium tulanense [p. 62] (Castell.) Agostini - J. trop. Med. Hyg. 35: 268. 1932 = Aleurisma tulanense (Castell.) Ota & Kawatsure - Arch. Derm. Syphil. 169: 156. 1933.
       Endomyces capsulatus Dodge & Ayers in Rewbridge, Dodge & Ayers - Am. J. Path. 5: 356. 1929 = Zymonema capsulatum (Dodge & Ayers) Dodge - Med. Mycol. p. 167, 1935.
       Glenospora brevis Castell. - J. trop. Med. Hyg. 36: 311. 1933 = Aleurisma breve (Castell.) Dodge - Med. Mycol., p. 790. 1935.
       Scopulariopsis americanus Ota - Jap. J. Derm. Urol. 28: 4. 1928.

       Teleomorph: Ajellomyces dermatitidis McDonough & Lewis - Mycologia 60: 77. 1968.

     Colonies on 4% malt agar at 24°C attaining 48-60 mm diam in 21 days, appearing white, felty with scattered fascicles of aerial hyphae at the centre, occasionally powdery; margin regular, defined; reverse cream. Hyphae 1-3 µm wide when fertile, 1-1.5 µm wide when infertile. Racquet hyphae absent. Conidia sessile or borne on narrow protrusions or short, thin side branches, occasionally 1-2 conidia borne on an ampuliform swelling, solitary or subtended by another conidium, smooth-walled, rarely slightly echinulate, thin-walled, subglobose or globose, occasionally pyriform, 2-7 x 2-4.5 µm, with very narrow basal scars (approx. 0.5 µm). Chlamydospores absent.
     Growth temperatures: minimum 5-10°C, optimum 25ºC, maximum 37°C. Not keratinolytic.
     At 37°C on 4% malt agar the colonies are heaped and produce budding cells which are hyaline, smooth- and thick-walled, irregular or globose to subglobose, up to 15 µm diam, daughter bud being connected to parent bud by a broad base (up to 4-5 µm).


Material examined

Living strains with only the anamorph. - CBS 126.33 received from A. Castellani; CBS 127.33 and 128.33 received from M. Moore, 1933; CBS 122.34 and 123.34 isolated from human patient by C. Bonne, U.S.A.; CBS 124.34 received from A. Castellani; CBS 134.36 received from N.F. Conant; CBS 125.51 isolated from man by F. Blank; CBS 514.74 isolated from human skin by G. van de Ploeg, Angola, 1974.

Living strains with the teleomorph and anamorph. - CBS 673.68 = ATCC 18187 (A) and CBS 674.68 = ATCC 18188 (a) monoascospore type cultures of Ajellomyces dermatitidis, isolated from man; CBS 886.70 (-) and 887.70 (+) received from K.J. Kwon-Chung, U.S.A.


Discussion

     Zymonema dermatitidis is the causal agent of Gilchrist's disease, also known as American blastomycosis or blastomycosis, reported from North and South America and Africa. The disease is a chronic granulomatous and suppurative illness which is first evident as a respiratory infection. Garrison & Boyd (1978) observed the germination of blastic conidia into 'yeast cells' in vitro, suggesting that the blastic conidium acts as the primary infective unit following inhalation. The disease may spread from the lungs to the bones, finally clearly marking the skin. The species is primarily recognised by the symptoms of the disease; no type cultures [p. 63] have been preserved for the synonyms listed above. The descriptions of the disease together with the appearance in vitro serve to confirm the synonymy of these names.
     Moore (1933) reported a supposed teleomorph producing naked, globose, 8-13 µm in diameter, 8-spored asci. Moore's descriptions and illustrations were used by Dodge (1935) for the new combination, Zymonema dermatitidis, thereby regarding Zymonema as an ascomycetous genus. McDonough & Lewis (1968) introduced the cleistothecial gymnoascaceous teleomorph, Ajellomyces dermatitidis, on the basis of mating experiments, suggesting that Moore may have mistaken chlamydospores containing globules for asci with ascospores.

 

Trichosporiella Kamyschko ex W. Gams & Domsch

       Trichosporiella Kamyschko - Botan. Materialy, Otd. Sporov. Rasten. 13: 167. 1960 (without indication of type material); ex W. Gams & Domsch - Nova Hedwigia 18: 19. 1969.

     Colonies slow-growing, submerged, cream-coloured, rarely with white aerial fascicles of hyphae, moist, yeast-like, tough. Hyphae submerged, rarely aerial, hyaline, smooth- and thin-walled, branching little and more or less orthotropically, insertion of branches often adjacent to the septa of the primary hypha. Blastic conidia borne terminally, mostly laterally all over the non-differentiated hyphae, sessile or on narrow, short protrusions, solitary, subhyaline, smooth- and thin-walled, globose, subglobose, rarely pyriform, 1-celled, with narrow basal scars. Intercalary conidia absent. Chlamydospores absent. Not keratinolytic or thermotolerant.

     Type species: Trichosporiella cerebriformis (de Vries & Kleine-Natrop) W. Gams

Discussion

     Trichosporiella is characterised by the moist, flat, tough colonies and the submerged fertile hyphae. The rare aerial fascicles are always sterile. Daylight normally promotes but is not always essential for sporulation.
     The genus Myriodontium Samson & Polonelli also produces lateral blastic conidia but differs in that these are borne on pronounced denticles and in that the colonies have a dense, floccose, aerial mycelium. The anamorph of Mycosylva Samson & Hintikka has a conidial morphology like that of Trichosporiella, but the mycelium develops concentric rings of aerial capitate synnemata. The simple type of conidial morphology exhibited by Trichosporiella resembles that of several species of Sporothrix Hektoen & Perkins ex Nicot & Mariat with lateral blastic conidia, particularly S. schenckii Hektoen & Perkins which also has flat, moist colonies (de Hoog, 1974). However, these Sporothrix species produce additional terminal clusters of sympodial conidia. [p. 64]

 

 

Key to the species

 

1a.

 

Conidia up to 8 µm diam; fertile hyphae little septate, not disarticulating

T. cerebriformis

 

1b.

 

Conidia up to 6 µm diam; fertile hyphae much septate, disarticulating at the septa

T. sporotrichoides

 

 

34. Trichosporiella cerebriformis (de Vries & Kleine-Natrop) W. Gams - Fig. 33.

 fig. 33

Fig. 33
Trichosporiella cerebriformis. CBS 344.53.

       Sporotrichum cerebriforme de Vries & Kleine-Natrop - Mycopath. Mycol. appl. 8: 159. 1957 = Trichosporiella cerebriformis (de Vries & Kleine-Natrop) W. Gams in v. Arx - Persoonia 6: 184. 1971.
       Trichosporiella hyalina Kamyschko - Botan. Materialy, Otd. sporov. Rasten. 13: 167. 1960: ex W. Gams & Domsch - Nova Hedwigia 18: 19. 1969.

     Colonies on malt 2% agar in daylight attaining 15-25 mm diam in 14 days, cream-coloured, submerged, moist, tough, sometimes with sparse fascicles of aerial hyphae up to 0.5-2 mm in height, locally raised at the centre to 2-3 mm, otherwise flat; margin usually regular; reverse white, pale yellowish to cream-coloured. Hyphae 1.5-3.5 µm wide; aerial hyphae when present fasciculate and sterile; submerged hyphae mostly fertile. Racquet hyphae rare. Conidia solitary, 1 to 2, rarely 3 borne per conidiogenous cell, seldom abundant, hyaline, smooth- and thin-walled, subglobose to globose or horizontally flattened, rarely pyriform, 4-9 x 4-8 µm, with narrow basal scars (less than 0.5-1 µm).
     Growth temperatures: minimum 10°C, optimum 25°C, maximum 25-30°C. [p. 65]


Material examined

CBS 344.53 = ATCC 12124 = IMI 59974 type strain isolated from human scalp by H.E. Kleine-Natrop, Kiel. F.R.G., 1952; CBS 135.68 = ATCC 22552 type strain of Trichosporiella hyalina, isolated from agricultural soil by W. Gams, F.R.G., 1963; CBS 243.68 = ATCC 2253 and CBS 244.68 = ATCC 2251 isolated from agricultural soil by W. Gams, F.R.G., 1962.


Discussion

     The Fusarium Link ex Fr. macroconidia described by Taylor (1970) for the type strain on malt agar, were not observed by the present author on either malt, cherry decoction, oatmeal, Sabouraud glucose or PYE agars. It seems probable that Taylor (1970) was dealing with a contaminated culture.

 fig. 34

Fig. 34
Trichosporiella sporotrichoides. a. CBS 671.74; b. CBS 557.77. [p. 66]

 

35. Trichosporiella sporotrichoides van Oorschot, sp. nov. - Fig. 34, Plate 3.

       Coloniae in agaro maltoso (2%) sub luce diurna 30-60 mm diam. post 14 dies, cremeae, submersae, tenaces, leves; margo regularis, fimbriata; reversum cremeum. Hyphae submersae, saepe fragmentantes, fertiles, 1-4.5 (-7) µm latae. Hyphae septum versus inflatae absunt. Conidia copiosa, solitaria, singula ad quina e cellulis conidiogenis oriunda, subhyalina, levia et tenuitunicata, plerumque, globosa, vel modice pyriformia, 3-6 x 3-4.5 µm, ad basim exigue truncata, cicatrice 0.3-1 µm diam.
       Typus CBS 671.74, isolatus e terra in Surinamia, a J.H. van Emden, 1974.

     Colonies on malt 2% in daylight attaining 30-60 mm diam in 14 days, cream-coloured, submerged, moist and tough, flat; margin regular and fimbriate; reverse cream-coloured. Hyphae submerged, often disarticulating at the septa, fertile, 1-4.5 (7.5) µm wide. Racquet hyphae absent. Conidia abundant, solitary, 1-5 borne per conidiogenous cell, subhyaline, smooth- and thin-walled, mostly globose, also slightly pyriform, 3-6 x 3-4.5 µm, truncate with narrow points of attachment (0.3-1 µm).
     Growth temperatures: minimum 10°C, optimum 20-25°C, maximum 25-30°C.


Material examined

CBS 671.74, type strain isolated from soil by J.H. van Emden, Surinam; CBS 152.76 isolated from forest soil under Picea abies by B. Söderström, Sweden; CBS 557.77 isolated from humus around moist roots by W. Gams, Sierra Nevada de Santa Marta, Columbia.


Discussion

     The species differs from Trichosporiella cerebriformis by the faster growth rate, broader, often disarticulating, more frequently septate fertile hyphae and the smaller conidia. The resemblance to Sporothrix strains lacking sympodial conidia is striking. The conidial morphology resembles that of certain strains of the Trichophyton mentagrophytes (Robin) Blanchard complex, but these have powdery or granular aerial mycelium with hyaline rather than subhyaline catenate conidia (microconidia) in addition to macroconidia, ampulliform swellings and frequently antler hyphae.
     Strain CBS 152.76 has a slower growth rate of only 5 mm in 14 days and rarely develops ampulliform swellings which each bear a single conidium. The ampulliform swellings in Myceliophthora and Emmonsia are more abundant and normally bear 2 to 4 blastoconidia.

 

Geomyces Traaen

       Geomyces Traaen - Nytt Mag. Naturvid. 52: 28. 1914.
       Glenosporella Nannizzi in Agostini - Atti. Ist. Bot. crittoz. Lab. Pavia 4: 98. 1931.

       Teleomorphic genus: Pseudogymnoascus [p. 67]

     Colonies only slightly spreading, initially white but soon becoming pale brown, dark brown, grey, reddish or yellow, felty or powdery, often with scattered tufts of aerial hyphae. Hyphae hyaline or pale yellow, thin-walled, narrow, branching of sterile hyphae more or less orthotropic and usually not correlated with the septation of the supporting hypha, branching of fertile hyphae more frequent, at acute angles, often once or twice verticillate with 2-4 branches per whorl, the secondary or tertiary branches arising just below the septa of the primary or secondary branch respectively. Thallic conidia normally borne terminally on verticillate branches, intergrading with intercalary conidia, or borne laterally and solitary on short protrusions or short side branches, subhyaline, pale yellow or pale greenish, smooth-walled or echinulate, thin-walled to slightly thick-walled, cuneiform, obovoid, ellipsoid or clavate, 1-celled, truncate with wide basal scars. Intercalary conidia borne on the outer branches of the verticillate hyphae, alternate, separated by short, mostly more broad than long sterile hyphal segments, in series of 2-4, subhyaline, pale greenish or pale yellow, smooth-walled or echinulate, thin-walled to slightly thick-walled, barrel-shaped, 1-celled, broader than the supporting hypha, truncate at both ends. Not thermotolerant, frequently keratinolytic.

     Lectotype species: Geomyces auratus Traaen


Discussion

     Geomyces differs from Chrysosporium in that the fertile hyphae show repeated verticillate branching. Fertile side branches are septate at their bases and arise at acute angles just below the septa of the main hypha. An additional septum may arise below the junction of the branches, forming a characteristic triangular cell. The fascicles of hyphae seen in culture adhere closely and separate with difficulty in a slide preparation. Preponderance of tall fascicles (3-5 mm) is often a sign of degeneration after repeated subculturing, the fasciculate hyphae usually being sterile. More virulent or fresh isolates tend to be more powdery with relatively shorter fascicles of fertile hyphae or semi-macronematous fertile hyphae ramifying in divergent directions.
     Malbranchea also has chains of intercalary conidia but these are cylindrical and equal in width to the fertile hyphae which do not branch verticillately. The intercalary conidia of Ovadendron Sigler & Carmichael are wider than the supporting hyphae but the fertile hyphae arise as short, coiled, lateral branches which develop conidia in a more or less basipetal succession (Sigler & Carmichael, 1976).
     Traaen (1914) described four species when he introduced the genus; only the type strain of Geomyces auratus remains, no herbarium material has been preserved. This strain was selected by Sigler & Carmichael

 fig. 35

Fig. 35
Geomyces pannorum var. pannorum. a. CBS 360.66; b. CBS 378.76; c. CBS 471.76.

 

36. Geomyces pannorum (Link) Sigler & Carmichael var. pannorum Fig. 35.

       Sporotrichum pannorum Link - Linn. Spec. Pl. 6: 13. 1824 = Chrysosporium pannorum (Link) Hughes - Can. J. Bot. 36: 749. 1958 = Geomyces pannorum (Link) Sigler & Carmichael - Mycotaxon 4: 377. 1976.
       Trichophyton albisicans Nieuwenhuis - Tijdschr. Geneesk. Ned. Indië 48: 49. 1908 = Atrichophyton albisicans (Nieuwenhuis) Castell. & Chalmers - Man. trop. Med., ed. 3, p. 1008. 1910 = Glenospora albisicans (Nieuwenhuis) Ota Ann. Parasit. hum. comp. 3: 84. 1925 = Glenosporella albisicans (Nieuwenhuis) Nannizzi in Agostini - Atti Ist. bot. Univ. Lab. Crittogam. Pavia 4: 98. 1931 = Aleurisma albisicans (Nieuwenhuis) C.W. Dodge - Med. Mycol., p. 788. 1935.
       Botrytis terrestris Jensen - Bull. Cornell agric. Exp. Stn 315: 489. 1912.
       Ciliciopodium hyalinum Daszewska - Bull. Soc. bot. Genève 2: 304. 1912.
       Corethropsis hominis Vuill. in Spillman & Jannin - Bull. Soc. fr. Derm. Syphil. 24: 227. 1913 = Torulopsis hominis (Vuill.) Castell. & Jacono - J. trop. -Med. Hyg. 36: 297. 1933 = Torulopsis hominis (Vuill.) Almeida - Annali Fac. Med. Sâo Paulo 9: 76. 1933.
       Geomyces vulgaris Traaen - Nytt Mag. Naturvid. 52: 29. 1914.
       Geomyces auratus Traaen - Nytt Mag. Naturvid. 52: 30. 1914.
       Geomyces cretaceus Traaen - Nytt Mag. Naturvid. 52: 31. 1914.
       Sporotrichum carnis Brooks & Hansf. - Trans. Br. mycol. Soc. 8: 131. 1923 = Aleurisma carnis (Brooks & Hansf.) Bisby - Trans. Br. mycol. Soc. 27: 111. 1944.
       Aleurisma lugdunense Vuill. in Massia & Grigorakis - C.r. Séanc. Soc. biol. 91: 1381. 1924. [p. 69]
       Aleurisma guillermondi Grigorakis - C.r. Seanc. Soc. biol. 97: 493. 1927.
       Sporotrichum cejpii Fassatiova - Preslia 25: 274. 1953.
       Chrysosporium verrucosum Tubaki - Spec. Pubis. Seto mar. biol. Lab., Biol. Res. J.A.R.E. 14: 6. 1961.

       Misapplied names: Sporotrichum parvulum Pass. sensu Redaelli - I Mycet. Assoc. Microb., p. 15. 1925 = Rhinocladium parvulum (Pass.) Grandinetti - Contrib. est. Esporotricose Sâo Paulo, 1934.
       Scopulariopsis lutea (Cost.) Tubaki sensu Tubaki - Nagaoa 5: 29. 1955.

     Colonies attaining 25-40 mm diam in 14 days, initially white, later becoming pale to dark grey, buff or brown, slightly felty to powdery, often with scattered tufts of aerial hyphae, up to 2 mm high at the centre, sometimes with radial ridges; margin defined, regular or locally indented, reverse cream-coloured, pale yellow to yellowish brown with a reddish brown to brown centre. Hyphae hyaline, smooth-walled, 0.5-3 µm wide; aerial hyphae mostly fertile; submerged hyphae sterile. Racquet hyphae absent. Terminal and lateral conidia subhyaline, occasionally pale greenish to yellowish, initially smooth- and thin-walled, sometimes becoming slightly echinulate or verruculose and thicker-walled, obovoid, ellipsoid, cuneiform or a little clavate, 3-6 x 2-4 µm, with wide basal scars (1-1.5 µm). Intercalary conidia alternate, in series of 1-3, subhyaline, occasionally pale greenish to yellowish, initially smooth-and thin-walled, usually becoming echinulate or a little verruculose and thicker-walled, barrel-shaped, 3-6 x 2-5 µm. Chlamydospores absent.
     Growth temperatures: minimum 5 °C, optimum 20-25°C, maximum 25°C. Not keratinolytic.


Material examined

Herbarium specimen. - Sporotrichum pannorum Link. B, on rotten cloth, collected by C.G.
Ehrenberg, F.R.G., 1824.

Living strains. - CBS 105.13 type strain of Botrytis terrestris Jensen, isolated from potato field soil, New York, U.S.A., 1911; CBS 106.13 type strain of Ciliciopodium hyalinum Daszewska, isolated from peat, Switzerland, 1912; CBS 108.14 type strain of Geomyces auratus Traaen, isolated from soil, Norway; CBS 193.26 = ATCC 7153 received from P. Redaelli as Sporotrichum parvulum Pass.; CBS 359.29 received from M. Engelhardt, F.R.G.; CBS 101.31 isolated from dromedary skin by E.A.R.T. Baudet; CBS 102.31 received from E.A.R.T. Baudet; CBS 133.31 received from G. Pollaci; CBS 134.32 isolated from wood pulp by H. Robak, Norway; CBS 112.35 received from N.C.T.C., U.K.; CBS 125.42 isolated rom Lathyrus root, Netherlands; CBS 106.44. isolated by H. Killian, Poland; CBS 122.44 isolated from slate slopes by S. Blumer, Switzerland; CBS 163.48 isolated from germinating fern spores by J. van Holder, Belgium; CBS 103.52 isolated from flax by A. Kaars-Sijpestein, Netherlands; CBS 103.53 isolated from human eye, Netherlands; CBS 104.53 isolated from child's scalp, Netherlands; CBS 105.53 isolated from human, Netherlands; CBS 151.53 isolated from human by J.J. Zoon, Netherlands; CBS 101.54 and 102.54 isolated in F.R.G.; CBS 362.58 = ATCC 22086 type strain of Sporotrichum cejpii Fassatiová, isolated from insect, 1953; CBS 158.59 = DAOM 33738, isolated from Tsuga heterophylla, British Columbia. 1951; CBS 120.63 isolated from soil by E. Küster; CBS 105.64 isolated from acidic soil by G.L. Hennebert, 1963; CBS 107.65 and 807.68 isolated from agricultural soil by W. Gams, F.R.G., 1964; CBS 657.70 received from R.H. Haskins, Saskatchewan, Canada; CBS 226.74 isolated from dead Acer leaf by H.A. van der Aa, Netherlands; CBS 378.76 = ATCC 11501 = IMI 23281 type strain of Sporotrichum carnis Brooks & Hansf., isolated from frozen meat by H.A. Dade, 1923; CBS 471.76 isolated from sandy soil by G.S. Taylor, U.K.; CBS 182.77 = IFO 8279 and CBS 183.77 = IFO 8280 received from K. Tubaki as [p. 70] Chrysosporium verrucosum Tubaki, isolated in Japan, 1965; CBS 270.77 = UAMH 3004 isolated by R.A.A. Morrall, Saskatchewan, Canada, 1968; CBS 478.78 isolated by O.L. Rudakov, U.S.S.R.


Discussion

     From the original description and drawing given by Tubaki (1961) for Chrysosporium verrucosum, it is clear that the fungus he dealt with was a Geomyces pannorum. The type strain of C. verrucosum has been lost (K. Tubaki, pers. comm.) but two strains, CBS 182.77 and 183.77, identified by Dr. Tubaki as C. verrucosum showed the conidial morphology and cultural characteristics of G. pannorum.
     Redaelli (1925) identified a culture of Geomyces pannorum, CBS 193.26, as Sporotrichum parvulum Pass. Grandinetti (1943) probably based the combination Rhinocladium parvulum (Red.) Grandinetti on the same strain. The type material of S. parvulum Pass. could not be traced; the brief original description in Brunaud (1888) does not clarify the identity of the species. The fungus depicted by Tubaki (1955) as Scopulariopsis lutea (Cost.) Tubaki shows the verticillate branching and series of alternate intercalary conidia characteristic of G. pannorum. For some of the synonyms listed above there is no available authentic material, but their descriptions and drawings in the original articles clearly point to G. pannorum.
     Geomyces pannorum is commonly encountered in soil varying in pH from 3.5-8.0 (Williams & Pugh, 1974), and also as a human or animal pathogen. In man the fungus may attack the nails (Krempl-Lamprecht, 1965; Schönborn & Schmorange, 1970) or cause skin lesions (Massia & Grigorakis, 1924; Grigorakis, 1927; Krempl-Lamprecht, 1965). Pugh & Evans (1970) isolated the species from bird feathers.
     From the material examined above and numerous additional freshly isolated strains, it is apparent that the variation in colony colour from shades of grey to brown cannot be correlated to the presence or absence or degree of roughness of the conidium walls. For this reason three of Traaen's strains are placed in synonymy with G. pannorum. However, strains with yellow colonies tend to produce longer series of alternate conidia, conidia being shorter than in most of the grey or brown strains. In addition the yellow strains may show rough-walled fertile hyphal branches. Strains with reddish colonies are always associated with the Pseudogymnoascus roseus teleomorph but are identical to G. pannorum strains in conidial morphology. The differences shown by the yellow and red strains do not warrant species distinction and they are therefore described as varieties.

 

37. Geomyces pannorum (Link) Sigler & Carmichael var. asperulatus (Sigler & Carmichael) van Oorschot, comb. nov. - Fig. 36.

       ? Geomyces sulphureus Traaen - Nytt Mag. Naturvid. 52: 30. 1914.
       Geomyces asperulatus Sigler & Carmichael - Mycotaxon 4: 376. 1976 (basionym). [p. 71]

 fig. 36

Fig. 36
Geomyces pannorum var. asperulatus. CBS 628.79.

     Colonies attaining 10-25 mm diam in 2 weeks, initially white or pale pink, usually becoming pale brown at the centre and ochre yellow in a clearly marked outer zone, rarely retaining a few pink patches, felty and slightly powdery, radiately convoluted, maximally 1.5 mm high at the centre; margin defined, locally indented; reverse black at the centre and reddish brown outward. Hyphae hyaline to pale yellow, smooth-walled if infertile but sometimes rough-walled if fertile, thin-walled, 0.5-1.5 µm wide; aerial hyphae fertile; submerged hyphae less fertile. Racquet hyphae absent. Terminal and intercalary conidia in series of 1-4, pale to bright yellow, echinulate at maturity, thick-walled, 2-3 x 3-4.5 µm, with wide basal scars (1-3 µm); terminal conidia obovoid to ellipsoidal, intercalary conidia alternate and barrel-shaped. Chlamydospores sometimes developing in the aerial mycelium, solitary, brown, smooth- and thick-walled, 4 x 3.5 µm.
     Growth temperatures: minimum 5°C, optimum 20-25°C, maximum 25-30°C. Not markedly keratinolytic.


Material examined

CBS 574.68 isolated by D.S. Malla, Denmark; CBS 124.77 = UAMH 183 type strain, isolated from soil by F. Raymond, Massachussetts, U.S.A., 1976, CBS 626.79 isolated from beach sand by W. Gams, Netherlands, 1978; CBS 627.79 = DAOM 167041 sent in by J.D. Bissett, Ottawa, Canada; CBS 628.79 isolated from meadow soil by G.J. Bollen, Netherlands, 1978.


Discussion

The type strain does not show rough-walled fertile hyphae, but this feature is prevalent in CBS 626.79 and CBS 628.79. The type material of Geomyces [p. 72] sulphureus Traaen has been lost but as the fungus was described as being yellow it may well have been a G. pannorum var. asperulatus.

 

38. Geomyces pannorum (Link) Sigler & Carmichael var. vinaceus (Dal Vesco) van Oorschot, comb. nov.

       Geomyces vinaceus Dal Vesco - Allionia 3: 14. 1957.

       Teleomorph: Pseudogymnoascus roseus Raillo - Zentbl. Bakt. Parasitkde, Abt. II. 78: 520. 1929.

     Colonies attaining 15-16 mm diam in 14 days, initially white, becoming reddish brown or red, felty to powdery, locally raised to 5 mm, occasionally with radial grooves; margin regular, defined; reverse cream-coloured and pale orange-brown at the centre, becoming dark brown with age. A reddish brown pigment is exuded into the medium at the base of the mycelium. Hyphae hyaline, smooth- and thin-walled, 0.8-2.5 µm wide, aerial hyphae mostly fertile, submerged hyphae mostly infertile. Racquet hyphae absent. Terminal and intercalary conidia hyaline, smooth-walled or occasionally slightly echinulate, thin-walled, 3-5 x 2-3.5 µm, with wide basal scars (1-2 µm wide); terminal conidia obovoid, somewhat cuneiform or ellipsoid, intercalary conidia barrel-shaped. Chlamydospores absent.
     Growth temperatures: minimum 5°C, optimum 20-25°C, maximum 25°C. Not markedly keratinolytic.


Material examined

Living strains with the teleomorph and anamorph. - CBS 390.54 isolated from soil by F.L. Raymond, Massachussetts. U.S.A. 1949; CBS 261.61 isolated from soil by C.A. Ghillini, Italy, 1960; CBS 319.62 isolated from soil by G. Dal Vesco, Italy; CBS 320.62 type strain of Geomyces vinaceus isolated from soil by G. Dal Vesco, Italy; CBS 395.65 = IMI 114651 neotype strain of Pseudogymnoascus roseus Raillo, isolated from swamp soil by A.E. Apinis, England, 1959; CBS 387.69 isolated from pine soil by P. Widden, Alberta, Canada; CBS 722.69 = TRTC 45536 type strain of Gymnoascus rhousiogongylinus Wener & Cain, isolated from forest soil by H.M. Wener, Ontario, Canada, 1967; CBS 605.70 isolated from soil by K.H. Domsch, F.R.G.; CBS 691.70 isolated from alpine soil by J. Bisset, Alberta, Canada, 1968; CBS 310.76 isolated from soil by S.C. Agrawal, Saugar, India.


Discussion

     The type strain of Arachniotus hebridensis Apinis, CBS 360.66, today only develops a Geomyces pannorum conidial state, sporulating poorly and remaining white. The peridial hyphae described by Apinis (1964), were smooth and hyaline, forming a loose network; asci were subglobose to elliptical, yellowish and 3.2-4.8 x 1.8-3.0 µm. The conidial state was not clearly described and not depicted but the shape and sizes given agree with those still produced by the type strain. It seems likely that Apinis was dealing with a Pseudogymnoascus roseus with a G. pannorum var. vinaceus anamorph. [p. 73]

 

Excluded generic names

Aleurisma Link - Mag. Ges. naturf. Fr., Berl. 3: 19. 1809; ex Fr. - Syst. mycol. 3: 453. 1832.
  Type species: Aleurisma sporulosum Link - Mag. Ges. naturf. Fr., Berl. 3: 19. 1809; ex Fr. - Syst. mycol. 3: 453. 1832 = Sporotrichum sporulosum (Link ex Fr.) Link - Spec. Pl. 6: 16. 1924 = Trichoderma sporulosum (Link ex Fr.) Hughes - Can. J. Bot. 36: 820. 1958.
  The type specimen from B was reidentified by Hughes (1958) as Trichoderma sporulosum (Link ex Fr.) Hughes [today known as T. polysporum (Link ex Pers.) Rifai].
  Specific names: albisicans, apiospermum, arloini, benigna, brasiliensis, breve, bulbusorum, carnis, castellani, dermatitidis, erubescens, flavissimum, granulosum, guilliermondi, inspersum, keratinophilum, lugdunense, macrosporum, metaeuropeum, metamericanum, molardi, roseum, saccharinum, salmoneum, tulanense, vuillemini.

  Blastomyces Cost. & Rolland - Bull. Soc. mycol. Fr. 4: 153. 1888.
  Type species: Blastomyces luteus Cost. & Rolland.
  The herbarium material of Costantin is unknown (Stafleu & Cowan, 1976); the type specimen of Blastomyces luteus is not kept at PC. The descriptions and drawings given by Costantin & Rolland (1888) are almost certainly of Chrysosporium merdarium and hence Blastomyces Cost. & Rolland is considered synonymous to Chrysosporium Corda.

  Blastomyces Gilchrist & Stokes - J. exp. Med. 3: 76. 1898.
  Type species: Blastomyces dermatitidis Gilchrist & Stokes.
  There is no type material but the species was described for the well known fungus which causes Gilchrist's disease or North American Blastomycosis etc. The genus Blastomyces Gilchrist & Stokes is a homonym of Blastomyces Cost. & Rolland. The type species was placed in Zymonema by Dodge (1935) (see Z. dermatitidis in text).
  Specific names: brasiliensis, dermatitidis.

  Blastomycoides Castell. - Proc. Soc. Med. 21: 451. 1928 (nomen provisorium).
  Type species: Blastomyces dermatitidis Gilchrist & Stokes - J. exp. Med. 3: 76. 1898 = Zymonema dermatitidis (Gilchrist & Stokes) Dodge - Med. Mycol., p. 160. 1935 = Blastomycoides dermatitidis (Gilchrist & Stokes) Castell.
  The type species was described for the agent of Gilchrist's disease (see Zymonema dermatitidis in text).

  Gilchristia Redaelli & Cif. - J. trop. Med. Hyg. 37: 281. 1934.
  Only species: Blastomyces dermatitidis Gilchrist & Stokes - J. exp. Med. 3: 76. 1898 = Gilchristia dermatitidis (Gilchrist & Stokes) Redaelli & Cif. - J. trop. Med. Hyg. 37: 281. 1934 = Zymonema dermatitidis (Gilchrist & Stokes) Dodge - Med. Mycol., p. 168. 1935. [p. 74]
     There is no type material but the species was described for the agent of Gilchrist's disease. The genus is synonymous to Zymonema (see text).

  Glenosporella Nannizzi in Agostini - Atti. Ist. Bot. crittogam. Lab. Pavia 4: 98. 1931.
  Type species: Trichophyton albisicans Nieuwenhuis - Tijdschr. Geneesk. Ned. Indie 48: 49. 1908 = Glenosporella albisicans (Nieuwenhuis) Nannizzi.
  Nieuwenhuis (1908) described this species for a tropical skin disease 'tinea albigena'. Today there is no type material but Ota (1925) clearly described and illustrated a fungus isolated and identified by Nieuwenhuis as Trichophyton albisicans from a patient suffering from tinea albigena. Although Nieuwenhuis's illustrations are not clear, those given by Ota almost certainly, represent Geomyces pannorum var. pannorum. The genus Glenosporella is thus regarded as synonymous to Geomyces.
  Specific names: albisicans, dermatitidis.

  Rhinocladiella Kamyschko - Notul. syst. Inst. cryptog. Horti bot. petropol. 13. 165. 1960.
  Only species: Rhinocladiella vesiculosa Kamyschko.
  Rhinocladiella Kamyschko is a homonym of Rhinocladiella Nannf. (1934) and hence Kamyschko (1961) proposed the new name Rhinocladiopsis Kamyschko, but failed to make a new combination. The type material could not be obtained from Leningrad but the description and illustration of the type species are such that the genus may be considered synonymous to Chrysosporium.

  Rhinocladiopsis Kamyschko - Notul. syst. Inst. cryptog. Horti. bot. petropol. 14: 243. 1961.
  See Rhinocladiella above.

 

Doubtful or excluded species

  albicans . - Endomyces albicans Okabe - Centbl. Bakt. Parasitkde, Abt. I.3: 186. 1929 = Zymonema albicans (Okabe) Dodge - Med. Mycol., p. 174. 1935.
  Endomyces albicans Okabe is a homonym of E. albicans (Robin) Vuill. which is today known as Candida albicans (Robin) Berkhout. There is no type material and it is not possible to identify the fungus from the original article.

  albo-luteolum. - Sepedonium albo-luteolum Sacc. & March. in March.- Bull. Soc. Roy. Bot. Belg. 24: 62. 1885 = Chrysosporium albo-luteolum (Sacc. & March.) Dominik - Zesz. nauk. wyzsz. Szk. roln. Szczec. 24: 45. 1967.
  The type material is not at PAD and was not referred to by Dominik (1967). The brief description given by Saccardo & Marchal and the lack of an illustration do not allow recognition of the species.

  album. - Zymonema album Dodge - Med. Mycol., p. 175. 1935. [p. 75]
  The description given by Dodge was probably based on that given for a Monilia species by Bianchi & Nino (1928) and it is possible that Dodge did not view the original material. The insufficient information and lack of material prevent classification.

  alvarezsotoi. - Monilia alvarezsotoi Mazza & Nino in Mazza et al. - Boln Inst. Clin. quir., Univ. B. Aires 6: 180. 1931 = Zymonema alvarezsotoi (Mazza & Nino) Dodge - Med. Mycol., p. 177. 1935 = Mycotorula alvarersotoi (Mazza & Nino) Nino - Boln. Inst. Clin. quir., Univ. B. Aires 14: 591. 1938.
  This species is today recognised as a synonym of Candida albicans (Robin) Berkhout (Lodder,. 1970).

  apiospermum. - Monosporium apiospermum Sacc. Annls mycol. 9: 254. 1911 = Scedosporium apiospermum (Sacc.) Sacc. in Dodge-Med. Mycol., p. 838. 1935 = Aleurisma apiospermum (Sacc.) Maire in Montpellier & Gouillon - Bull. Soc. Path. exot. 14: 290. 1921.
  The species is today recognised as a Scedosporium state of Petriellidium boydii (Shear) Malloch (v. Arx, 1973c).

  arlioni. - Achorion arlioni Busquet. - Thèse, Lyon, 1890; Ann. Micrographie 3: 11. 1891 = Aleurisma arlioni (Busquet) Vuill. in Massia & Grigorakis - C.r. séanc. Biol. 91: 1381. 1924.
  There appears to be no type material but the macro- and microconidia described and depicted by Busquet (1890) belong to a Trichophyton, probably T. mentagrophytes,

  barabicum. - Chrysosporium barabicum Scharapov - Nov. Syst. niz. Rast. 11: 268. 1974.
  The type strain was requested from VKM and the Central Siberian Botanical Garden, USSR, but not received. The diagrams and descriptions given by Scharapov (1974) show the fungus to be similar to Chrysosporium keratinophilum except in that the conidia are pyriform to subglobose and 9.7-11.2 x 11.2-13.2 µm, while C. keratinophilum conidia are obovoid to clavate and 3.5-22 x 3.5-11 µm.

  benigna. - Aleurophora benigna Magalhâes - Brasil Med. 30: 369. 1916 = Aleurisma benigna (Magalhâes) Vuill. - Champ. Paras. Myc. Homme, p. 114. 1931.
  The lack of herbarium material, illustrations and incomplete description make this a nomen dubium.

  bonaerense. - Endomyces bonaerensis Greco - Argent. med. 1908. = Zymonema bonaerense (Greco) Dodge - Med. Mycol., p. 175. 1935.
  The asci described in the original article were regarded with some doubt by Dodge (1935) and he provisionally placed the species in Zymonema. [p. 76]

  brasiliense. - Zymonema brasiliense Splendore - Bull. Soc. Pathol. exot. 5: 317. 1912 = Coccidioides brasiliensis (Splendore) Almeida - Ann. Fac. Med. Sâo Paulo 4: 95. 1929 = Paracoccidioides brasiliensis (Splendore) Almeida - C.r. Soc. Biol. 105: 316. 1930 = Blastomyces brasiliensis (Splendore) Conant & Howell - J. invest. Dermat. 5: 357. 1942 = Aleurisma brasiliensis (Splendore) Aroeia Neves & Bogliolo - Mycopath. Mycol. appl. 5: 140. 1951.
  The fungus is the agent of South American blastomycosis, Brazilian blastomycosis or Paracoccidiomycosis and is today known as Paracoccidioides brasiliensis. Conidial morphology at 24°C in culture is almost identical to Emmonsia parva var. parva, E. parva var. crescens and Zymonema dermatitidis. However at 37°C in vitro or in vivo the pathogen may be distinguished by the thinner walled, spherical or elliptical cells which may bud at any point on the surface without distortion of the original shape (Emmons et al., 1977). Similar cells are known in the pathogenic fungus Cryptococcus neoformans (Sanfelice) Vuill. which has a basidiomycetous Filobasidiella Kwon-Chung teleomorph with terminal or lateral groups of basidia and catenate basidiospores (Emmons et al., 1977).

  brasiliensis. - Emmonsia brasiliensis Batista et al. - Rev. Fac. Med. Univ. Ceara 3: 52. 1963.
  The type strain, CBS 255.65, is a Sporotrichum pruinosum Gilman & Abbott (Padhye & Carmichael, 1968).

  bucalis. - Monilia bucalis Nino & Puglisi - Semana Med. 34: 22. 1927 = Zymonema bucalis (Nino & Puglisi) Dodge - Med. Mycol. p. 176. 1935.
  The species was described for a human mouth pathogen and the original illustration shows a budding and a mycelial state. Redaelli & Ciferri (1942) placed the species in synonymy with Mycotorula albicans, today known as Candida albicans (Robin) Berkhout (Lodder, 1970), but they failed to refer to any type material. Correct identity remains uncertain.

  bulbusorum. - Aleurisma bulbusorum Link - Mag. Ges. naturf. Fr., Berl. 7: 38. 1815.
  The type specimen at B was examined by Hughes (1958) and found to consist of bulb scales bearing white pustules consisting of aggregations of mineral matter. Hughes concluded that Link's observations were based on this material.

  casteIlanii. - Acladium castellanii Pinoy in Castell. - Brit. med. J. 2: 486. 1916 = Pseudomicrosporon castellanii (Pinoy) Craik - J. trop. Med. Hyg. 26: 184. 1923 = Aleurisma castellanii (Pinoy) Vuill. - Champ. Paras. Myc. Homme, p. 115. 1931.
  Neither the diagram nor the description in the original article clarify the identity of the fungus in the absence of type material.

  ciferrina. - Emmonsia ciferrina Thirum. et al. - Mycopath. Mycol. appl. 26: 330. 1965.
  The type strain, CBS 363.65, was reidentified as Sporotrichum pruinosum Gilman & Abbott by Padhye & Carmichael (1968). [p. 77]

  crateriforme . - Endomyces crateriforme Hudelo et al. - C. r. Acad. Sci. 170: 1087. 1920 = Zymonema crateriforme (Hudelo et al.) Dodge - Med. Mycol., p. 174. 1935 = Sporotrichum crateriforme (Hudelo et al.) Vuill. - Champ. Paras. Myc. Homme, p. 70. 1931.
  There is probably no original material. Muller (1965) regarded the species as a nomen dubium but did suggest that it may be an Endomycopsis Dekker.

  cruzi. - Endomyces cruzi de Mello & Paes - Arg. Hig. Pat. Exot. 6: 60. 1918 = Zymonema cruzi (de Mello & Paes) Dodge - Med. Mycol. p. 176. 1935.
  The pathogen was isolated from a human suffering from bronchitis and asthma. The original description leaves the identification doubtful.

  degenerans. - Cryptococcus degenerans Vuill. - Rev. Ren. Sci. Pures. Appl. 12: 740. 1901 = Zymonema degenerans (Vuill.) de Mello & Paes - Hig. Pat. Exot. 6: 280. 1918 = Torulopsis degenerans (Vuill.) Almeida - Annaes Fac. Med. Sâo Paulo 9. 10. 1933.
  The basionym was introduced for the causal agent of a blastomycosis. There is no type material and the original description does not clarify the identity of the fungus.

  denticolum. - Chrysosporium keratinophilum (D. Frey) Carmichael var. denticolum C. Moreau - Mycopath. Mycol. appl. 37: 37. 1969 = Basipetospora denticola (C. Moreau) C. Moreau - Bull. Soc. mycol. Fr. 87: 43. 1971.
  There is no type material (Moreau, pers. comm.). A strain received from Moreau as Basipetospora denticola (CBS 132.78), which had been isolated from the same source, was reidentified as Scopulariopsis candida (Guéguen) Vuill.

  dimorphum. - Chrysosporium dimorphum Matsushima Icones Microf. a Mats. lect., p. 29. 1975.
  The type strain, CBS 167.79, was reidentified as Sporotrichum dimorphosporum v. Arx.

  erubescens. - Aleurisma erubescens Nees - Syst. fung. p. 51. 1817; ex Fries Syst. mycol. 3: 453. 1832 = Sporotrichum erubescens (Nees) Link - Jahrb. Gewachsk. 1: 178. 1818.
  Herbarium material from B included two collections made by Ehrenberg which may well be the type specimens used by Nees. Only sterile mycelium remains. Link (1824) placed Aleurisma erubescens in synonymy with Sporotrichum sporulosum which is today regarded as a Trichoderma polysporum (Link ex Pers.) Rifai.

  flavissimum. - Aleurisma flavissimum Link - Mag. Ges naturf. Fr. Berl. 7. 38. 1815; ex Steudel - Nomencl. bot., Berl. p. 63. 1824.
  Hughes's mount from the type specimen (DAOM 55593) consists of merely alga-like cells (Carmichael, 1962). Saccardo (1886) and Vuillemin (1911) listed Aleurisma flavissimum as a synonym of Sporotrichum laetum Link. The type specimen of S. laetum from B consists of unidentifiable pale brown to hyaline, globose, 2-4 µm diam spores. [p. 78]

  flavissimum. - Sporotrichum flavissimum Link - Mag. Ges. naturf. Fr, Berl. 7: 34. 1818; ex Steudel - Nomencl. bot., p. 401. 1824.
  This species was incorrectly listed as a synonym of Aleurisma flavissimum Link by both Saccardo (1886) and Vuillemin (1911). Vuillemin's isolate, CBS 101.10, which he labelled A. flavissimum, is actually a Chrysosporium merdarium. The type material of Sporotrichum flavissimum from B shows spores of a Leucogyrophana species (J.A. Stalpers, pers. comm.).

  foetidum. - Chrysosporium foetidum Apinis & B. Clark - Trans. Br. mycol. Soc. 63: 263. 1974.
  The mating type strains, CBS 453.75 and CBS 452.75, of the teleomorph Neoxinophila foetida Apinis & B. Clark only show a few isolated microconidia. In the original description the anamorph was attributed micro- as well as macroconidia similar to those of Trichophyton. It is therefore likely that N. foetida has a Trichophyton anamorph, macroconidium production having disappeared.

  fusca. - Myceliophthora fusca Doyer - Meded. phytopath. Lab. Willie Commelin Scholten 10: 32. 1927.
  The type strain, CBS 190.25, was reidentified by von Arx (1973a) as Ptychogaster rubescens Boud.

  granulosum. - Aleurisma granulosum Mart. - Flora crypt. Erlang., p. 335. 1817 = Sporotrichum granulosum (Mart.) Link - Jahrb. Gewachsk. 1: 183. 1818 = Collarium granulosum (Mart.) Rabenh. Deutschl. Krypt. Flora 1: 70. 1844 (non C. granulosum Fr. - Syst. mycol. 3: 441. 1832) = Coccotrichum martii Link - Linn. Spec. Plant. 4: 26. 1824 (name change).
The original description is vague and type material is not preserved at M.

  harteri. - Cryptococcus harteri Verdun - Précis Parasitol. 1912 = Atelosaccharomyces harteri (Verdun) de Beurmann & Gougerot - reference not located. 1913 = Parasaccharomyces harteri (Verdun) de Mello et al. - Arg. Hig. Pat. Exot. 6: 33. 1918 = Myceloblastenon harteri (Verdun) Ota - Jap. J. Derm. Urol. 28: 4. 1928 = Monilia harteri (Verdun) Vuill. - Champ. Paras. Myc. Homme, p. 85. 1931 = Zymonema harteri (Verdun) Dodge - Med. Mycol., p. 178. 1935 = Torulopsis harteri (Verdun) Almeida - Annaes Fac. med. Sâo Paulo 9: 10. 1933.
  The fungus was described as the causal agent of a case of generalised blastomycosis (Dodge, 1935). Dodge placed his Zymonema harteri in synonymy with Mycotorula albicans (today Candida albicans; Lodder, 1970) but it is unlikely that Dodge saw the original material.

  hirundo. - Chrysosporium hirundo Scharapov - Nov. Syst. niz. Rast. 15: 14. 1978.
  The type strain, CBS 668.78, was seen to be a Malbranchea species, most closely resembling the Malbranchea state of Uncinocarpus reesii Sigler & Orr.

  histosporocellularis . - Zymonema histosporocellularis Haberfeld - Tesis., Sâo Paulo, 1919 = Mycoderma histosporocellularis (Haberfeld) [p. 79] Neveu-Lemaire - Précis Parasitol. 70. 1921 = Coccidioides histosporocellularis (Haberfeld) da Fonseca in Zeissler. 1923 = Lutziomyces histosporocellularis (Haberfeld) Fonseca Filha. 1939.
  Emmons et al. (1977) listed this species as a synonym of Paracoccidioides brasiliensis. The present author has not had access to the original description but presumably Emmons et al. based their synonymy on the symptoms of the disease, type material almost certainly being non-existent.

  immitis. - Coccidioides immitis Rixford & Gilchrist - Johns Hopkins Hosp. Rept. 1. 1896 = Zymonema immitis (Rixford & Gilchrist) de Mello & Fernandes - Arg. Hig. Pat. Exot. 6. 1928 = Blastomycoides immitis (Rixford & Gilchrist) Castell. - Amer. J. trop. Med. 8: 381. 1928.
  The pathogen is today recognised as Coccidioides immitis, the cause of coccidiomycosis (Emmons et al., 1977).

  inflata. - Myceliophthora inflata Burnside - Pap. Mich. Acad. Sci. 8: 82. 1928 = Paecilomyces inflatus (Burnside) Carmichael - Can. J. Bot. 40: 1148. 1962.
  The species was placed in Paecilomyces after examination of the type strain, CBS 259.39.

  inspersum. - Aleurisma inspersum Link - Mag. Ges. naturf. Fr., Berl. 3: 38. 1809.
  Link (1824) placed Aleurisma inspersum in synonymy with Sporotrichum sporulosum Link which is today recognised as a Trichoderma (Hughes, 1958). The author has not had access to any type material.

  kreiselii. - Chrysosporium kreiselii Dominik - Zesz. nauk. wyzsz. Szk. roln. Szczec. 19: 135. 1965 (without reference to type material).
  Though the descriptions and drawings given by Dominik (1965) only indicate unicellular rough- and thin-walled, 4-9 x 2-3 µm microconidia, the photographs in the same article show additional 2- and 3-celled, apparently smooth-walled macroconidia resembling those of Trichophyton terrestre Durie & D. Frey.

  lanuginosus. - Blastomycoides lanuginosus Castell. - Brit. J. Derm. Syphil. 42: 372. 1930 = Glenospora lanuginosa (Castell.) Agostini - J. trop. Med. Hyg. 34. 287. 1931 = Trichosporium gammelii var. lanuginosus (Castell.) Dodge - Med. Mycol., p. 792. 1935.
  The fungus was originally described for a human blastomycosis which sporulated poorly in pure culture. Insufficient information prevents definite identification as Zymonema dermatitides the cause of North American blastomycosis.

  lignorum. - Chrysosporium lignorum Bergmann & Nilsson Dept. Forest. Prod. Roy. Coll. Forest., Stockholm, Res. notes 53: 28. 1966 (without latin diagnosis).
  W. Gams reidentified the type specimen, CBS 671.71, as Sporotrichum pulverulentum Novobranova, today known to have a Phanerochaete teleomorph (Stalpers, 1978).

  lipsiense. - Sporotrichum lipsiense Benedek - Derm. Wschr. 83: 1735. 1926 = Rhinocladium lipsiense (Benedek) (Brumpt, Précis Parasitol., ed 6: 1836. 1949. [p. 80]
  There does not appear to be any type material. From the original description it seems possible that the fungus was a Geomyces pannorum var. pannorum.

  macrosporum. - Aleurisma macrosporum Link - Mag. Ges. naturf. Fr., Berl. 3: 38. 1809;
  The type material was not obtained from B. Link (1824) regarded the species as synonymous to Sporotrichum fructigena (Pers.) Link of which the type material (in L) has been reidentified as Monilia fructigena (Pers.) Pers. (Hughes, 1958).

  metamericana. - Glenospora metamericana Castell. - J. trop. Med. Hyg. 36: 310. 1933 = Aleurisma metamericanum (Castell.) Dodge - Med. Mycol., p. 790. 1935.

  metaeuropeum. - Glenospora metaeuropeum Castell. - Med. Press Circular 136: 440. 1933; J. trop. Med. Hyg. 136: 309. 1933 = Aleurisma metaeuropeum (Castell.) Dodge - Med. Mycol., p. 770. 1935.

  molardi. - Endomyces molardi Salvat & Fontoynant - Bull. Soc. Path. exot.. 15: 320. 1922 = Zymonema molardi (Salvat & Fontoynant) Dodge - Med. Mycol., p. 173. 1935.
  From the original description and illustration it appears that Salvat & Fontoynant (1922) were probably dealing with a Candida albicans.

  olivaceum. - Botrytis olivacea Link - Linn. Spec. Plant. 1: 18. 1824 = Sporotrichum olivaceum (Link) Fr. - Syst. mycol. 3: 417. 1830 = Chrysosporium olivaceum (Link) Taylor - Mycologia 62. 815. 1970.
  The type specimen from B no longer shows any fungal material. Taylor (1970) examined a slide from the type specimen, DAOM 49583, and described 'aleuriospores' as well as thin-walled spores. The present author could only find thin-walled spores in the same slide, and these somewhat resembled phialospores. The species is regarded as doubtful.

  osteophilum. - Sepedonium osteophilum Bon. - Handb. allgem. Mycol. p. 100. 1851 = Chrysosporium osteophilum (Bon.) Dominik - Zesz. nauk. wyzsz. Szk. roln. Szczec., 24: 45. 1968.
  Herbarium material belonging to Bonorden is not known today (Stafleu & Cowan, 1976). The description given by Bonorden (1851) is very brief; the accompanying drawing could represent a Sepedonium, the conidia being pigmented, very rough-walled with narrow basal attachments. However, no sizes are given and no mention is made of an accompanying phialidic state. The true position of the species remains doubtful.

  pollacii. - Chrysosporium pollacii Taylor - Mycologia 62: 821. 1970.
  The type strain, CBS 270.28, is no longer in the CBS collection. The author was not able to obtain the herbarium material from Montana. Taylor (1970) compared the species to Chrysosporium tropicum and C. luteum (= Myceliophthora lutea) but classified it separately on account of an inability to degrade keratin, the narrow conidium bases and the lack of pigmentation.[p. 81]

  pruinosum, Sporotrichum pruinosum Gilman & Abbott - Iowa State Coll. J. Sci. 1: 36. 1927 = Chrysosporium pruinosum (Gilman & Abbott) Carmichael - Can. J. Bot. 40: 1166. 1962.
  The type specimen, CBS 129.27, is a Sporotrichum (Stalpers, 1978) and the species has been shown to have a basidiomycetous teleomorph, Phanerochaete chrysosporium Burds., by Burdsall & Eslyn (1974).

  roseum. - Aleurisma roseum Link - Mag. Ges. naturf. Fr., Berl. 3: 38.1809.
  The type material could not be obtained from B. Link (1824) placed the species as a synonym of Sporotrichum sporulosum which is today known as Trichoderma sporulosum (Hughes, 1958).

  saccharinum. - Aleurisma saccharinum Link - Mag. Ges. naturf. Fr., Berl. 3: 38. 1809.
  Link (1824) regarded the species as a synonym of the earlier Sporotrichum agaricinum Link ex Link. Herbarium material from Link's collection received from B consists of two samples, each marked Aleurisma saccharinum, A. agaricinum and Sporotrichum agaricinum. It is probable that Link based his diagnoses on this collection. The only fungal material present in the samples was identified as Aspergillus glaucus by R.A. Samson and J. Stalpers (pers. comm.).

  salmoneum. - Aleurisma salmoneum Vuill. - C.r. hebd. Séanc Acad. Sci., 189: 407. 1929.
  Lack of herbarium material and the brief description leave this species doubtful.

  sepedonioides. - Monosporium sepedonioides Harz - Bull. Soc. imp. Moscou 44: 105. 1871 = Sepedonium sepedonioides (Harz) Rabenh. - Deutsch. Krypt. Fl. 1: 222. 1907 = Chrysosporium sepedonioides (Harz) Dominik - Zesz. nauk. wyzsz. Szk. rolp. Szczec. 24: 57. 1967.
  The type material could not be obtained from either L or M. The drawing given by Harz cannot be used to identify the species or genus with certainty and it therefore remains doubtful.

  spinosum. - Sepedonium spinosum Sacc. - Michelia 2: 635. 1882 = Chrysosporium spinosum (Sacc.) Dominik Zesz. nauk. wyzsz. Szk. roln. Szczec. 24: 59. 1967.
  The type specimen from PAD shows capitate cystidia and clamp connections. Stalpers (pers. comm.) has identified this material as Resinicium bicolor (Alb. & Schw. ex Fr.) Parm.

  spinulosum. - Chrysosporium spinulosum Negroni - Boln Acad. nac. Med., Buenos Aires 41: 534. 1963 (with no indication of type material).
  The original strain has been lost and no herbarium material was prepared (Negroni, pers. comm.). The diagnosis of the species was not illustrated but the fungus described resembles Chrysosporium farinicola and C. sulfurea. [p. 82]

  suIphurea. - Myceliophthora sulphurea Goddard - Bot. Gaz. 56: 263. 1913.
  The type material could not be located. Correct placement of the species is uncertain since the original drawings show different kinds of conidia which could belong to Chrysosporium, Trichosporiella and to a yeast.

  Vuillemini. - Aleurisma vuillemini Grigorakis et al. - C.r. Soc. Biol. 95: 650. 1926.
  There is no type material at PC and the description does not serve to identify the species with certainty.

 

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Acknowledgements

  The author would like to express her gratitude to Dr. G.S. de Hoog for his guidance during the research work and thorough criticism of the manuscript. She is indebted to Dr. J.A. von Arx, Prof. Dr. K. Verhoeff, Dr. W. Gams, Dr. R.A. Samson and Dr. G.A. de Vries for their valuable comments on the manuscript. Dr. J.W. Carmichael is thanked for very kindly providing numerous UAMH isolates. Miss I. ten Hoedt is acknowledged for her technical assistance. [p. 87]

 Plate 1

Plate 1
Myceliophthora vellerea. CBS 479.76. Verruculose conidium and ampulliform swellings. Marker indicates 5 µm.

 Plate 2

Plate 2
Chrysosporium keratinophilum. CBS 122.79. Top left and bottom right conidia show remains of subtending cell. Marker indicates 10 µm.

 Plate 3

Plate 3
Trichosporiella sporotrichoides. CBS 671.74. Marker indicates 5 µm.