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50 ˜m
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Fig. 22.2. Colletotrichum coccodes sclerotia with setae and conidia.
Antifungal susceptibility: (Table 22.1).
Table 22.1. Colletotrichum spp. (Australian national data); MIC µg/mL.
Antifungal No. ≤0.016 0.03 0.06 0.125 0.25 0.5 1 2 4 8 16 32 ≥64
AmB 13 1 3 4 3 1 1
ISAV 4 2 2
VORI 13 1 1 4 5 2
POSA 13 1 2 5 4 1
ITRA 13 1 2 2 2 1 5
References: McGinnis (1980); Domsch et al. (2007); de Hoog et al. (2015).
DOI: 10.1079/9781800622340.0022
Descriptions of Medical Fungi 76
(Costantin) Batko
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Synonym: Entomophthora coronata (Costantin) Kevorkian
The species of the genus Conidiobolus produce characteristic multinucleate primary and secondary (replica-
tive) conidia on top of unbranched conidiophores. Each subspherical conidium is discharged as a result of
the pressure developed within the conidium, and each bears a more or less prominent papilla after discharge
(Ellis, 2005a). The genus contains 27 species, however only C. coronatus, C. incongruus and C. lamprauges
have been reported as causative agents of human and animal infection. A morphological identification key
for clinical isolates was given by Vilela et al. (2010), and an overview of the taxonomy of the genus by
Nie et al. (2020).
Conidiobolus coronatus is commonly present in soil and decaying leaves. It has a worldwide distribution
especially tropical rain forests of Africa. Human infections are usually restricted to the rhinofacial area.
However, there are reports of disseminated infection in immunocompromised patients. All human infections
have been confined to the tropics (Shaikh et al., 2016).
RG-2 organism
Morphological description: Colonies grow rapidly and are flat, cream coloured, glabrous, becoming radially
folded and covered by a fine, powdery, white surface mycelium and conidiophores (Fig. 23.1). The lid of the
petri dish soon becomes covered with conidia, which are forcibly discharged by the conidiophores. The
colour of the colony may become tan to brown with age. Conidiophores are simple forming solitary, terminal
conidia, which are spherical, 10-25 μm in diameter, single-celled and have a prominent papilla (Fig. 23.1).
Conidia may also produce hair-like appendages, called villae. Conidia germinate to produce either, (1) single
or multiple hyphal tubes that may also become conidiophores which bear secondary conidia, or (2) replicate
by producing multiple short conidiophores, each bearing a small secondary conidium.
Molecular identification: ITS sequencing is useful for identification of most clinical isolates.
© CAB International 2023. Descriptions of Medical Fungi (eds S. Kidd, C.Halliday and D. Ellis)
77
(a)
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(b)
(c)
10 ˜m
Fig. 23.1. Conidiobolus coronatus showing (a) culture with satellite colonies from forcibly discharged conidia,
(b) spherical conidia with hair-like appendages (villae) and (c) conidia with prominent papillae.
10 ˜m
References: McGinnis (1980); King (1983); Rippon (1988); Kwon-Chung and Bennett (1992); de Hoog et al.
(2015); Vilela and Mendoza (2018).
DOI: 10.1079/9781800622340.0023
Descriptions of Medical Fungi 78
Beyma) Z.U. Khan, Gene & Guarro
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Synonymy: Lecythophora hoffmannii (J.F.H. Beyma) W. Gams & McGinnis; Phialophora hoffmannii (J.F.H.
Beyma) Schol-Schwarz; Margarinomyces hoffmannii J.F.H. Beyma.
In response to recent changes in the nomenclature for pleomorphic fungi, Khan et al. (2013) have transferred all Lecythophora species to Coniochaeta. The genus Coniochaeta is morphologically similar to
Phialemonium. It also forms adelophialides, but in Coniochaeta, these conidiogenous cells show conspicuous
collarettes, and the colonies are usually pink salmon to dark brown, although discrete phialides like those of
Acremonium may be also present (Perdomo et al., 2011b). Coniochaeta hoffmannii has been associated with
cases of subcutaneous infections, keratitis, sinusitis, peritonitis and canine osteomyelitis. Coniochaeta muta-
bilis has been described to be the causative agent of human peritonitis, endocarditis, endophthalmitis and
keratitis (Perdomo et al., 2011b).
RG-1 organism
Morphological description: Colonies are flat, smooth, moist, pink to orange, with regular and sharp margin;
reverse pink (Fig. 24.1). Hyphae are narrow, hyaline, producing conidia laterally from small collarettes
directly on the hyphae, or from lateral cells which are sometimes arranged in dense groups; lateral cells
flask-shaped or nearly cylindrical. Collarettes are unpigmented, about 1.5 μm wide. Conidia are hyaline,
smooth, and thin walled, broadly ellipsoidal to cylindrical or allantoid, 3.0-3.5 x 1.5-2.5 μm, produced in
slimy heads (Fig. 24.1).
Comment: Coniochaeta and Phialemonium species are poorly differentiated morphologically and are
difficult to identify. They may also be confused with poorly sporulating Fusarium or Acremonium species.
Molecular identification: Khan et al. (2013) used a combined sequence dataset of the ITS region, D1/D2,
ACT and β-tubulin genes to resolve the unique phylogenetic status of this species.
© CAB International 2023. Descriptions of Medical Fungi (eds S. Kidd, C.Halliday and D. Ellis)
79
10 ˜m
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10 ˜m
Fig. 24.1. Coniochaeta hoffmannii culture and hyphae with small collarettes (arrows) producing conidia.
Antifungal susceptibility: (Table 24.1).
Table 24.1. Coniochaeta hoffmannii very limited data (Australian national data); MIC μg/mL.
Antifungal
AmB 3 1 1 1
ISAV 2 1 1
VORI 3 1 1 1
POSA 3 1 1 1
ITRA 3 1 1 1
No. ≤0.016 0.03 0.06 0.125 0.25 0.5 1 2 4 8 16 32 ≥64
References: McGinnis (1978b); Perdomo et al. (2011b); Khan et al. (2013); de Hoog et al. (2015).
DOI: 10.1079/9781800622340.0024
Descriptions of Medical Fungi 80
25 Cryptococcus Vuill.
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The genus Cryptococcus is characterised by globose to elongate yeast-like cells or blastoconidia that reproduce
by narrow-necked budding. Pseudohyphae are absent or rudimentary. Most species are encapsulated,
although the extent of capsule formation depends on the medium (Fig. 25.1). Under certain conditions of
growth, the capsule may contain starch-like compounds, which are released into the medium by many strains.
Within tissue sections, mucicarmine or Alcian blue stains the capsule of Cryptococcus species to distinguish
it from other yeasts with similar morphologies. Historically the Cryptococcus genus was large, but many
species have been reclassified into other genera including Naganishia albida (Cryptococcus albidus) and
Papiliotrema laurentii (Cryptococcus laurentii).
On solid media the cultures are generally mucoid or slimy in appearance; red, orange or yellow carotenoid
pigments may be produced, but young colonies of most species are usually non-pigmented, and cream in
colour (Fig. 25.2). All Cryptococcus species produce urease and are non-fermentative. Nitrate may be assimilated
or not; inositol assimilated. The genus Cryptococcus differs from the genus Rhodotorula in its inositol
assimilation (Kurtzman et al., 2011).
Cryptococcosis is a chronic, subacute to acute pulmonary, systemic or meningitic disease, initiated by the
inhalation of infectious propagules (basidiospores and/or desiccated yeast cells) from the environment.
Primary pulmonary infections have no diagnostic symptoms and are usually subclinical. On dissemination,
the fungus usually shows a predilection for the central nervous system, however skin, bones and other visceral
organs may also become involved. Cryptococcus neoformans and C. gattii are the principle pathogenic
species. Naganishia albida and Papiliotrema laurentii have on occasion also been implicated in human infection
(Liu et al., 2015).
(a)
Fig. 25.1. (a) Bird seed agar plate showing brown colonies of Cryptococcus neoformans and white colonies of
Candida albicans, and (b) India ink preparation of a central spinal fluid specimen showing the characteristic wide
gelatinous capsule formed by C. neoformans.
© CAB International 2023. Descriptions of Medical Fungi (eds S. Kidd, C.Halliday and D. Ellis)
(b)
5 ˜m
81
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10 ˜m
Fig. 25.2. Cryptococcus neoformans culture and a basidium with basidiospores.
In a phylogenetic analysis, Hagen et al. (2015) proposed to recognise Cryptococcus neoformans (formerly
C. neoformans var. grubii) and Cryptococcus deneoformans (formerly C. neoformans var. neoformans) as
separate species. In addition, these authors also describe five genotypic clades of Cryptococcus gattii as new
species; C. gattii, C. bacillisporus, C. deuterogattii, C. tetragattii and C. decagattii (Hagen et al., 2015). This
proposal has not been universally accepted (Kwon-Chung et al., 2017). It has been recommended to continue
to use “Cryptococcus neoformans species complex” and “C. gattii species complex” for practical identification
purposes, rather than creating more species (Kwon-Chung et al., 2017).
Molecular identification: Requires ITS and/or D1/D2 sequencing, particularly for differentiating molecular
types (proposed species) within the species complex.
MALDI-ToF MS: Can provide reliable species and subspecies level identification, but its accuracy is dependent
on database quality (Arendrup et al., 2014).
References: Barnett et al. (1983); Rippon (1988); Kwon-Chung and Bennett (1992); Casadevall and Perfect
(1998); Kwon-Chung (2011); McTaggart et al. (2011); de Hoog et al. (2015); Lockhart et al. (2016); Sugita
et al. (2017); Morales-Lopez and Garcia-Effron (2021).
25.1. Cryptococcus gattii complex (Vanbreus. & Takashio) Kwon-Chung & Boekhout
Synonymy: Filobasidiella bacillispora Kwon-Chung; Cryptococcus neoformans var. gattii Vanbreus &
Takashio.
Cryptococcus gattii complex has two serotypes (B and C) and was reclassified as a separate species
from C. neoformans in 2002 (Kwon-Chung et al., 2002). C. gattii generally has a more restricted geographi-
cal distribution than C. neoformans, causing human disease in climates ranging from temperate to tropical
Australia, Papua New Guinea, parts of Africa, India, Southeast Asia, Mexico, Brazil, Paraguay and Southern
Descriptions of Medical Fungi 82
on this medium.
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California, although recent infections have also been reported from Vancouver Island, Canada and in the
Pacific Northwest, USA (Pfaller and Diekema, 2010; Espinel-Ingroff and Kidd, 2015). C. gattii has a specific
ecological association with numerous species of Eucalyptus trees, although the Canadian isolates are associated
with a range of native non-Eucalyptus species (Kidd et al., 2007a,b). Historically considered a pathogen in
immunocompetent hosts, a recent review in Australia noted an increase in C. gattii infections in HIV-negative
immunocompromised patients (Chen et al., 2012). Cryptococcosis caused by C. gattii is often associated with
large mass lesions (cryptococcomas) in the lung and/or brain (Sorrell, 2001).
Canavanine glycine bromothymol blue (CGB) agar (Kwon-Chung et al., 1982) is the media of choice to differentiate C. gattii complex from C. neoformans complex (Fig. 25.3). This simple biotype test is based on the
ability of C. gattii isolates to grow in the presence of L-canavanine and to assimilate glycine as a sole carbon
source.
RG-2 organism
Culture: Colonies (SDA) cream coloured, smooth, mucoid, yeast-like colonies. Microscopy: Globose to ovoid
budding yeast-like cells 3.0-7.0 x 3.3- 7.9 μm.
India ink preparation: Positive - distinct, wide gelatinous capsules are present.
Dalmau plate culture: Budding yeast cells only. No pseudohyphae present.
Bird seed agar: Colonies turn dark brown in colour as colonies selectively absorb a brown pigment from this
media. Colonies are often more mucoid when compared to C. neoformans (Staib et al., 1989).
Canavanine-glycine-bromothymol blue (CGB) agar: Turns blue within 2-5 days.
Germ tube formation: Negative.
Physiological tests: (+ Positive, - Negative, v Variable, w Weak, s Slow, n not done).
Fermentation: Absent.
Descriptions of Medical Fungi 83
Growth reactions:
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Glucose + L-Sorbose – myo-Inositol +
Sucrose + L-Rhamnose + DL-Lactate –
Raffinose +,w D-Xylose + D-Gluconate +
Melibiose – L-Arabinose +,w 2-Keto-D-gluconate v
Galactose + D-Arabinose + D-Glucosamine v
Lactose – D-Ribose v N-Acetyl-D-glucosamine v
Trehalose + Glycerol – D-Glucuronate +
Maltose + Erythritol – Nitrate –
Melezitose + Ribitol v Urease +
Methyl-⍺-D-glucoside + Galactitol + 0.1% Cycloheximide –
Soluble starch + D-Mannitol + Growth at 37oC +
Cellobiose +,w D-Glucitol +
Key features: Encapsulated yeast; absence of pseudohyphae; growth at 37oC; positive hydrolysis of urea; negative
fermentation of sugars and positive assimilation of glucose, maltose, sucrose, galactose, trehalose, raffinose,
inositol, cellobiose, rhamnose, arabinose, melezitose and xylose, and negative assimilation of nitrate, lactose,
melibiose, erythritol and soluble starch; growth on bird seed (Guizotia abyssinica seed) or caffeic acid agar
colonies turn a dark brown colour; growth on CGB agar turning it blue within 2-5 days.
Antifungal susceptibility: (Table 25.1).
Table 25.1. Cryptococcus gattii complex (Australian national data with additional ISAV data from Espinel-Ingroff et al.
(2015b); MIC μg/mL. Note: All Cryptococcus species are intrinsically resistant to echinocandins.
Antifungal No. ≤0.016 0.03 0.06 0.125 0.25 0.5 1 2 4 8 16 32 ≥64
AmB 230 1 3 18 52 57 41 55 3
FLU 230 1 4 16 42 73 58 31 4 1
ISAV 430 106 109 93 90 25 7
VORI 230 21 43 72 41 26 5
POSA 168 20 13 32 57 41 9 1 1
ITRA 230 11 22 57 92 45 3
5FC 230 3 12 53 93 54 11 4
25.2. Cryptococcus neoformans complex (Sanfelice) Vuillemin
Synonymy: Filobasidiella neoformans Kwon-Chung; Cryptococcus neoformans var. neoformans.
This species has two varieties: C. neoformans var. grubii (serotype A) and C. neoformans var. neoformans
(serotype D).
C. neoformans var. grubii: Has a worldwide distribution, causing 95% of all C. neoformans infections. It has
been isolated from various sources in nature and is noted for its association with accumulations of avian
guano, especially with pigeon excreta. The fungus has also been isolated from the dung of caged birds including canaries, parrots and budgerigars. Other environmental isolations of C. neoformans var. grubii include
rotting vegetables, fruits and fruit juices, wood, dairy products and soil.
C. neoformans var. neoformans: Has a more restricted distribution with infections being more prevalent in
Europe, including France, Italy and Denmark, where it accounts for 30% of isolates. Moreover, C. neofor-
mans var. neoformans infections are more strongly correlated with older patients, the skin, and the use of
corticosteroids (Franzot et al., 1999).
Descriptions of Medical Fungi 84
RG-2 organism
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Culture: Colonies (SDA) cream coloured, smooth, mucoid, yeast-like colonies.
Microscopy: Globose to ovoid budding yeast-like cells 3.0-7.0 x 3.3-7.9 μm.
India ink preparation: Positive - distinct, wide gelatinous capsules are present.
Dalmau plate culture: Budding yeast cells only. No pseudohyphae present.
Bird seed agar: Colonies turn dark brown in colour as colonies selectively absorb a brown pigment from this
media (Staib et al., 1989).
Canavanine-glycine-bromothymol blue (CGB) agar: Does not grow on this medium and leaves the colour unchanged.
Creatinine dextrose bromothymol blue thymine (CDBT) agar: C. neoformans var. neoformans grows
as bright red colonies, turning the medium a bright orange after 5 days. No colour change is observed for
C. neoformans var. grubii (Irokanulo et al., 1994).
Germ tube formation: Negative.
Physiological tests: (+ Positive, – Negative, v Variable, w Weak, s Slow, n not done).
Fermentation: Absent.
Growth reactions:
Glucose + L-Sorbose – myo-Inositol +
Sucrose + L-Rhamnose + DL-Lactate –
Raffinose +,w D-Xylose + D-Gluconate +
Melibiose – L-Arabinose +,w 2-Keto-D-gluconate v
Galactose + D-Arabinose + D-Glucosamine v
Lactose – D-Ribose v N-Acetyl-D-glucosamine v
Trehalose + Glycerol – D-Glucuronate +
Maltose + Erythritol – Nitrate –
Melezitose + Ribitol v Urease +
Methyl-⍺-D-glucoside + Galactitol + 0.1% Cycloheximide –
Soluble starch + D-Mannitol + Growth at 37oC +
Cellobiose +,w D-Glucitol +
Key features: Encapsulated yeast; absence of pseudohyphae; growth at 37oC; positive hydrolysis of urea;
negative fermentation of sugars and positive assimilation of glucose, maltose, sucrose, galactose, trehalose,
raffinose, inositol, cellobiose, rhamnose, arabinose, melezitose and xylose, and negative assimilation of
nitrate, lactose, melibiose, erythritol and soluble starch; growth on bird seed (Guizotia abyssinica seed) or
caffeic acid agar colonies turn a dark brown colour; does not grow on CGB agar (no colour change).
Antifungal susceptibility: (Table 25.2).
Table 25.2. Cryptococcus neoformans complex (Australian national data with additional ISAV data from Espinel-Ingroff
et al., 2015b); MIC μg/mL. Note: All Cryptococcus species are intrinsically resistant to echinocandins.
Antifungal No. ≤0.016 0.03 0.06 0.125 0.25 0.5 1 2 4 8 16 32 ≥64
AmB 397 2 5 17 85 59 111 10 9 9
FLU 398 1 1 17 44 78 102 11 0 34 9 2
ISAV 494 178 179 10 6 22 7 2
VORI 359 85 90 103 59 21 1
POSA 314 17 46 70 104 66 11
ITRA 398 18 59 111 157 46 6 1
5FC 398 1 1 7 31 72 98 120 58 7 1 2
DOI: 10.1079/9781800622340.0025
Descriptions of Medical Fungi 85
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