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9.4. Aspergillus fumigatus Fresenius
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Aspergillus section Fumigati includes species characterised by uniseriate aspergilla with columnar conidial
heads in shades of blue-green and flask-shaped vesicles (Raper and Fennell, 1965). Teleomorphic species
belonging to the “Aspergillus fischeri series” of the A. fumigatus complex were originally placed in the genus
Neosartorya (family Trichocomaceae) by Malloch and Cain (1972). These have now been transferred to
Aspergillus and the section Fumigati now includes more than 60 species (Samson et al., 2007).
Although A. fumigatus sensu stricto is recognised as the major human pathogen within the complex, recent
phylogenetic studies have demonstrated that some human and animal infections may be caused by A. lentulus,
A. fumigatiaffinis, A. fumisynnematus, A. fischeri, A. udagawae and A. hiratsukae (Coriglione et al., 1990;
Summerbell et al., 1992; Padhye et al., 1994; Lonial et al., 1997; Jarv et al., 2004; Balajee et al., 2005a, 2006;
Barrs et al., 2013).
RG-2 organism
Morphological description: Colonies show typical blue-green surface pigmentation with a suede-like surface
consisting of a dense felt of conidiophores (Fig. 9.6). Conidial heads are typically columnar (up to 400 x 50 μm
but often much shorter and smaller) and uniseriate. Conidiophore stipes are short, smooth-walled and have
conical-shaped terminal vesicles which support a single row of phialides on the upper two thirds of the vesicle. Conidia are produced in basipetal succession forming long chains and are globose to subglobose (2.5-3.0
μm in diameter), green and rough-walled to echinulate (Fig. 9.6). Note: This species is thermotolerant and
grows at temperatures up to 55
Key features: Uniseriate and columnar conidial heads with the phialides limited to the upper two thirds of
the vesicle and curving to be roughly parallel to each other. Thermotolerant grows at temperatures up to
o
55
C. Note: Aspergillus lentulus differs by slower growth and absence of growth at 48oC (Yu et al., 2020).
Molecular identification: Sequence analysis of ITS is sufficient to identify to species complex level only.
Definitive identification requires sequence analysis of the β-tubulin, Cal and ACT genes (Balajee et al., 2005a;
Samson et al., 2007).
o
C.
10 ˜m 10 ˜m
Fig. 9.6. Aspergillus fumigatus culture and conidial head morphology. Note: Uniseriate row of phialides on the upper
two thirds of the vesicle.
Descriptions of Medical Fungi 26
Antifungal susceptibility: (Table 9.4).
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Table 9.4. Aspergillus fumigatus complex (Pfaller et al., 2013a; and Australian national data); MIC, *MEC μg/mL.
Antifungal No. ≤0.016 0.03 0.06 0.12 0.25 0.5 1 2 4 8 16 32 ≥64
AmB 1110 2 40 121 179 290 433 44 1
ISAV 344 1 1 6 42 188 83 20 1 2
VORI 1074 1 2 29 138 553 274 42 19 10 6
POSA 1005 44 88 147 419 258 34 13 1 1
ITRA 1110 8 20 53 149 542 298 28 4 1 1 6
ANID* 761 616 11 9 21 2 1 2
MICA* 761 653 89 13 4 2
9.5. Aspergillus lentulus Balajee & Marr
Aspergillus lentulus appears to be widely distributed in soil and is now well documented as a causative agent
of invasive aspergillosis in immunosuppressed patients. A. lentulus is part of the A. fumigatus complex
(Balajee et al., 2005b).
RG-2 organism
Morphological description: Colonies are suede-like to floccose, white with interspersed grey-green patches of
conidia (conidiation is slow to poor in most strains) (Fig. 9.7). Conidial heads are short columnar and uniserate.
Conidiophore stipes are smooth-walled, sometimes sinuous and are often constricted at the neck. Vesicles are
usually subglobose in shape. Conidia are globose to broadly ellipsoidal (2-3.2 μm in diameter), smooth to
finely roughened (Fig. 9.7).
Key features: Morphologically like A. fumigatus, but slower growing with poor conidiation and absence of
growth at 48
o
C (Yu et al., 2020).
Molecular identification: Aspergillus lentulus can be distinguished from other members of the section Fumigati
by sequence analysis of β-tubulin, Cal and ACT genes (Balajee et al., 2005a, 2005b; Samson et al., 2007).
Antifungal susceptibility: (Table 9.5).
Table 9.5. Aspergillus lentulus (Australian national data); MIC, *MEC μg/mL.
Antifungal No. ≤0.016 0.03 0.06 0.125 0.25 0.5 1 2 4 8 16 32 ≥64
AmB 17 1 2 2 7 4 1
ISAV 7 1 3 3
VORI 16 4 4 5 3
POSA 15 4 3 7 1
ITRA 17 4 2 7 2 2
ANID* 9 7 1 1
MICA* 9 8 1
Descriptions of Medical Fungi 27
10 ˜m
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Fig. 9.7. Aspergillus lentulus culture and conidial head morphology.
10 ˜m
9.6. Aspergillus nidulans (Eidam) Winter
Aspergillus section Nidulantes; Gams et al. (1985) includes species with biserate conidial heads, brown
pigmented often short stipes and green conidia, cleistothecia are soft-walled, surrounded by Hülle cells,
ascospores are red to purple in colour. Section Nidulantes is one of the largest subgenera of the genus Aspergillus
and includes about 80 species. A. nidulans is a typical soil fungus with a worldwide distribution and has been
reported to cause disease in human and animals (Bastos et al., 2020). Several other species within the section
have also been reported as medical pathogens, A. sydowii, A. unguis, A. rugulovalvus and A. quadrilineatus.
RG-1 organism
Morphological description: Colonies are typically plain green in colour with dark red-brown cleistothecia
developing within and upon the conidial layer. Reverse may be olive to drab grey or purple-brown (Fig. 9.8).
Conidial heads are short columnar (up to 70 x 30 μm in diameter) and biseriate. Conidiophore stipes are
usually short, brownish and smooth-walled. Conidia are globose (3-3.5 μm in diameter) and rough-walled
(Fig. 9.8). Ascomata are spherical, purple, 100-200 μm in diameter, surrounded by a yellowish to cinnamon
layer of scattered hyphae bearing a dense aggregation of pale yellow, thick-walled, spherical to subspherical
Hülle cells (Fig. 9.9). Asci are 8-spored, spherical to subspherical, 7-12 μm in diameter. Ascospores are purple-red, smooth-walled, lenticular with two equatorial crests, 3.5-4.5 x 3.5-4.0 μm.
Key features: Conidial heads are short columnar and biseriate. Stipes are usually short, brownish and
smooth-walled. Conidia are globose and rough-walled.
Molecular identification: ITS sequencing is sufficient to identify to species complex only. A. nidulans can be
distinguished from other members of the section Nidulantes by sequence analysis of β-tubulin, Cal and ACT
genes.
Descriptions of Medical Fungi 28
10 ˜m
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10 ˜m
Fig. 9.8. Aspergillus nidulans culture and biseriate conidial head morphology. Note: The short, brownish conidiophore stipes.
20 ˜m 15 ˜m
Fig. 9.9. Aspergillus nidulans cleistothecium showing numerous reddish-brown ascospores and aggregations of thick-
walled, spherical to subspherical thick-walled Hülle cells.
Descriptions of Medical Fungi 29
Antifungal susceptibility: (Table 9.6).
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Table 9.6. Aspergillus nidulans complex (Australian national data); MIC, *MEC μg/mL.
Antifungal No. ≤0.016 0.03 0.06 0.125 0.25 0.5 1 2 4 8 16 32 ≥64
AmB 57 5 10 3 18 16 5
ISAV 12 3 8 1
VORI 56 1 3 12 29 9 1 1
POSA 53 3 8 10 17 13 1 1
ITRA 57 2 4 9 15 25 1 1
ANID* 35 18 10 5 2
MICA* 35 22 11 2
9.7. Aspergillus niger van Tieghem
The black aspergilli, Aspergillus section Nigri (Gams et al., 1985) includes species with uniseriate or biseriate
conidial heads, spherical to pyriform vesicles, smooth-walled stipes and black or near black coloured conidia.
This group contains about 26 species with Aspergillus niger being the most common species isolated. A. niger
can be isolated from all continents and is not very selective with respect to environmental conditions. A. niger
is one of the most common and easily identifiable species of the genus Aspergillus, with its white to yellow
mycelial culture surface later bearing black conidia. This species is very commonly found in aspergillomas
and is the most frequently encountered agent of otomycosis. It is also a common laboratory contaminant.
Other species within this group that have been linked to human and animal infection include A. acidus,
A. aculeatus, A. brasiliensis and A. tubingensis.
RG-1 organism
Morphological identification: Colonies consist of a compact white or yellow basal felt covered by a dense
layer of dark brown to black conidial heads (Fig. 9.10). Conidial heads are large (up to 3 mm by 15 to 20 μm
in diameter), globose, dark brown, becoming radiate and tending to split into several loose columns with age.
Conidiophore stipes are smooth-walled, hyaline or turning dark towards the vesicle. Conidial heads are
biseriate with the phialides borne on brown coloured metulae, that are twice as long as the phialides
(Fig. 9.10). Conidia are globose to subglobose (3.5-5 μm in diameter), dark brown to black and rough-walled.
Key features: Conidial heads are dark brown to black, radiate and biseriate with metulae twice as long as the
phialides. Conidia brown and rough-walled.
Molecular identification: In Aspergillus section Nigri, all species can be distinguished from each other using
Cal sequence data, and all except one can be distinguished using β-tubulin sequence data. ITS sequencing is
not recommended for definitive species identification (Samson et al., 2007).
Antifungal susceptibility: (Table 9.7).
Table 9.7. Aspergillus niger complex (Pfaller et al., 2013a; and Australian national data); MIC, *MEC μg/mL.
Antifungal No. ≤0.016 0.03 0.06 0.125 0.25 0.5 1 2 4 8 16 32 ≥64
AmB 185 16 31 33 81 21 3
ISAV 46 1 3 2 17 18 3 2
VORI 18 1 4 9 10 24 79 51 3 1
POSA 17 0 2 8 19 28 61 50 2
ITRA 185 1 4 17 26 85 47 3 1 1
ANID* 128 89 24 6 2 1
MICA* 128 58 45 18 2 5
Descriptions of Medical F
ungi 30
10 ˜m
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10 ˜m
Fig. 9.10. Aspergillus niger culture and conidial head morphology. Note: Conidial heads are biseriate, large, globose,
dark brown, becoming radiate with the phialides borne on metulae.
9.8. Aspergillus terreus Thom
Aspergillus section Terrei (Gams et al., 1985); Aspergillus terreus complex includes species with biseriate,
columnar conidial heads in shades of buff to brown (Raper and Fennell, 1965). The most important species
of this section is A. terreus, which is ubiquitous in the environment (Samson et al., 2011). Aspergillus terreus
occurs commonly in soil and is occasionally reported as a pathogen of humans and animals. Two other
species have been reported as medical pathogens, A. alabamensis and A. niveus.
RG-2 organism
Morphological identification: Colonies are typically suede-like and cinnamon-buff to sand-brown in colour
with a yellow to deep dirty brown reverse (Fig. 9.11). Conidial heads are compact, columnar (up to 500 x
30-50 μm in diameter) and biseriate. Conidiophore stipes are hyaline and smooth-walled (Fig. 9.11). Conidia
are globose to ellipsoidal (1.5-2.5 μm in diameter), hyaline to slightly yellow and smooth-walled.
Key features: Cinnamon-brown cultures, conidial heads biseriate with metulae as long as the phialides.
Molecular identification: Aspergillus terreus can be distinguished from other members of the section Terrei
by sequence analysis of β-tubulin, Cal and ACT genes. ITS sequencing is sufficient to identify to species
complex level only.
Descriptions of Medical Fungi 31
10 ˜m
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Fig. 9.11. Aspergillus terreus culture and conidial head morphology. Note: Conidial heads are biseriate and tightly
columnar.
Antifungal susceptibility: (Table 9.8).
Table 9.8. Aspergillus terreus complex (Pfaller et al., 2013a; and Australian national data); MIC, *MEC μg/mL.
Antifungal No. ≤0.016 0.03 0.06 0.125 0.25 0.5 1 2 4 8 16 32 ≥64
AmB 168 1 3 11 37 78 36 2
ISAV 43 1 7 20 8 6 1
VORI 166 5 25 86 38 7 4 1
POSA 156 2 14 23 70 36 7 4
ITRA 169 5 3 19 53 78 10 1
ANID* 128 89 24 6 2 1
MICA* 128 58 45 8 2 5
DOI: 10.1079/9781800622340.0009
Descriptions of Medical Fungi 32
10 Aureobasidium pullulans (de
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Bary) Arnaud
Aureobasidium pullulans has a worldwide distribution and is usually isolated as a saprophyte, occasionally
from skin and nails. However, it has also been reported as a rare causative agent of phaeohyphomycosis,
mycotic keratitis and peritonitis in patients on continuous ambulatory peritoneal dialysis (CAPD).
RG-1 organism
Morphological description: Colonies are fast growing, smooth, soon covered with slimy masses of conidia,
cream or pink becoming brown or black (Fig. 10.1). Hyphae are hyaline and septate, frequently becoming
dark brown with age and forming chains of one- to two-celled, thick-walled, darkly pigmented arthroconidia.
The arthroconidia are only of secondary importance in recognising members of this genus. Conidia are produced synchronously in dense groups from indistinct scars or from short denticles on undifferentiated, hyaline to sub-hyaline hyphae. Conidia are hyaline, smooth-walled, single-celled, ellipsoidal but of very variable
shape and size (8-12 x 4-6 μm), often with an indistinct hilum (a mark or scar at the point of attachment)
(Fig. 10.1). Temperature: optimum 25
Key features: Slimy masses of conidia produced simultaneously from the vegetative hyphae, which may also
form chains of darkly pigmented, thick-walled arthroconidia.
Molecular identification: Recommended barcoding genes are TEF-1 and D1/D2 domains (de Hoog et al.,
2015).
Antifungal susceptibility: (Table 10.1).
o
C; maximum 35-37oC.
Table 10.1. Aureobasidium pullulans (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 15 1 6 4 4
ISAV 6 3 2 1
VORI 15 2 6 6 1
POSA 14 3 8 3
ITRA 15 3 9 3
References: Hermanides-Nijhof (1977); McGinnis (1980); Domsch et al. (2007); Zalar et al. (2008);
Najafzadeh et al. (2014); de Hoog et al. (2015, 2020).
© CAB International 2023. Descriptions of Medical Fungi (eds S. Kidd, C.Halliday and D. Ellis)
33
20 ˜m
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Fig. 10.1. Aureobasidium pullulans culture and microscopy showing darkly pigmented arthroconidia and the presence
of numerous hyaline conidia which are produced on short denticles.
DOI: 10.1079/9781800622340.0010
Descriptions of Medical Fungi 34
Synonymy: Basidiobolus meristosporus Drechsler; Basidiobolus heterosporus Srinivasan & Thirumalachar;
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Basidiobolus haptosporus Drechsler.
Basidiobolus ranarum is commonly present in decaying fruit and vegetable matter, and as a commensal in
the intestinal tract of frogs, toads and lizards. It has been reported from tropical regions of Africa and Asia
including India, Indonesia and Australia. B. ranarum is a known cause of subcutaneous and gastrointestinal
entomophthoromycosis often as a result of traumatic inoculation or a barrier break (Ellis, 2005a; Vilela and
Mendoza, 2018).
RG-2 organism
Morphological description: Colonies are moderately fast growing at 30
o
C, flat, yellowish-grey to creamygrey, glabrous, becoming radially folded and covered by a fine, powdery, white surface mycelium. Satellite
colonies are often formed by germinating conidia ejected from the primary colony (Fig. 11.1). Microscopic
examination usually shows the presence of large vegetative hyphae (8-20 μm in diameter) forming numerous
round (20-50 μm in diameter), smooth, thick-walled zygospores that have two closely appressed beak-like
appendages. The production of “beaked” zygospores is characteristic of the genus (Fig. 11.1). Two types of
asexual conidia are formed, although isolates often lose their ability to sporulate with subculture. Special
media incorporating glucosamine hydrochloride and casein hydrolysate may be needed to stimulate sporulation (Shipton and Zahari, 1987). Primary conidia are globose, one-celled, solitary and are forcibly discharged
from a sporophore. The sporophore has a distinct swollen area just below the conidium that actively participates in the discharge of the conidium (Fig. 11.1). Secondary (replicative) conidia are clavate, one-celled and
are passively released from a sporophore. These sporophores are not swollen at their bases. The apex of the
passively released spore has a knob-like adhesive tip. These spores may function as sporangia, producing
several sporangiospores.
Molecular identification: ITS sequencing is useful for identification of most clinical isolates. In the past,
pathogenic isolates of Basidiobolus were identified as either B. ranarum, B. meristosporus or B. haptosporus,
until all three species were found to be identical by rDNA RFLP analysis (Kwon-Chung and Bennett, 1992).
However, recent sequence data may suggest the need to re-establish several species (Bshabshe et al., 2020;
Al-Hatmi et al., 2021). Many sequences in GenBank are misidentified, so comparison to sequences from Type
cultures is important.
Key features: The genus Basidiobolus is distinguished primarily by the morphology and development of
forcibly discharged conidia and zygospores, both of which must be observed for reliable identification to
genus and species level (King, 1983).
References: Strinivasan and Thirumalachar (1965); Greer and Friedman (1966); Dworzack et al. (1978);
Rippon (1988); Kwon-Chung and Bennett (1992); Davis et al. (1994); Vikram et al. (2012); Shaikh et al.
(2016); Hung et al. (2020b).
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