Polyphasic taxonomy of Aspergillus section Fumigati and its teleomorph Neosartorya

RA Samson; S Hong; SW Peterson; JC Frisvad; J Varga

doi:10.3114/sim.2007.59.14

. 2007;59:147–203. doi: 10.3114/sim.2007.59.14

Polyphasic taxonomy of Aspergillus section Fumigati and its teleomorph Neosartorya

RA Samson ^1,^*, S Hong ², SW Peterson ³, JC Frisvad ⁴, J Varga ^1,⁵

PMCID: PMC2275200 PMID: 18490953

Abstract

The taxonomy of Aspergillus section Fumigati with its teleomorph genus Neosartorya is revised. The species concept is based on phenotypic (morphology and extrolite profiles) and molecular (β-tubulin and calmodulin gene sequences) characters in a polyphasic approach. Four new taxa are proposed: N. australensis N. ferenczii, N. papuaensis and N. warcupii. All newly described and accepted species are illustrated. The section consists of 33 taxa: 10 strictly anamorphic Aspergillus species and 23 Neosartorya species. Four other Neosartorya species described previously were not available for this monograph, and consequently are relegated to the category of doubtful species.

Keywords: Aspergillus section Fumigati, extrolite profiles, Neosartorya, phylogenetics, polyphasic taxonomy

INTRODUCTION

Aspergillus section Fumigati includes species characterised by uniseriate aspergilli, columnar conidial heads in shades of green and flask shaped vesicles (Raper & Fennell 1965). Teleomorphic species belonging to the “Aspergillus fischeri series” of the A. fumigatus group (Raper & Fennell 1965) were placed in the genus Neosartorya (family Trichocomaceae) by Malloch & Cain (1972). Section Fumigati includes more than 20 Neosartorya species and 10 anamorphic species (Pitt et al. 2000; Samson 2000; Horie et al. 2003; Hong et al. 2005, 2006, 2007).

Aspergillus fumigatus Fresenius is an ubiquitous filamentous fungus in the environment, and also an important human pathogen (Raper & Fennell 1965). Several Neosartorya species have been described as causal agents of human diseases including invasive aspergillosis, osteomyelitis, endocarditis and mycotic keratitis (Coriglione et al. 1990; Summerbell et al. 1992; Padhye et al. 1994; Lonial et al. 1997; Jarv et al. 2004; Balajee et al. 2005, 2006). All of the Neosartorya species produce heat-resistant ascospores that are frequently encountered in different food products (Gomez et al. 1994; Samson 1989; Tournas 1994). The several mycotoxins produced by these species may cause serious health hazard (Fujimoto et al. 1993; Frisvad & Samson 1990; Larsen et al. 2007). Some species also have valuable properties for mankind; e.g. N. fischeri strains produce fiscalins which effectively inhibit the binding of substance P to the human neurokinin receptor (Wong et al. 1993), while A. fumigatus strains produce pyripyropenes, potent inhibitors of acyl-CoA:cholesterol acyltransferase (Tomoda et al. 1994), the immunosuppressant restrictocins (Müllbacher & Eichner 1984), ribotoxins (Lin et al. 1995) and fumagillin that has amebicidal activity (McCowen et al. 1951). Neosartorya spinosa can be used for the complete enzymatic recovery of ferulic acid from corn residues (Shin et al. 2006).

Here we present an overview of the species belonging to Aspergillus section Fumigati based on analysis of macro- and micromorphology, extrolite profiles and β-tubulin, calmodulin, ITS and actin gene sequences of the isolates. We also describe four new homothallic Neosartorya species found in soil samples in Australia and Papua New Guinea using this polyphasic approach and list synonymies.

MATERIALS AND METHODS

Source of microorganisms

The fungi examined included type strains or representatives of all species available for examination in Aspergillus section Fumigati. Some atypical isolates collected in Australia and Papua New-Guinea were also examined to clarify their taxonomic status (Table 1).

Table 1.

Aspergillus section Fumigati isolates used in this study.

Species	Isolate No.^*	Source
A. brevipes	CBS 118.53^T	Soil, Australia
A. duricaulis	CBS 481.65^T	Soil, Buenos Aires, Argentina
A. fumigatiaffinis	IBT12703^T	Soil, U.S.A.
A. fumigatus	CBS 133.61^T = NRRL 163	Chicken lung, U.S.A.
A. fumisynnematus	IFM 42277^T	Soil, Venezuela
A. lentulus	CBS 117887^T = NRRL 35552 = KACC 41940	Man, U.S.A.
A. novofumigatus	IBT 16806^T	Soil, Ecuador
A. unilateralis	CBS 126.56^T	Rhizosphere, Australia
A. viridinutans	CBS 127.56^T	Rabbit dung, Australia
A. turcosus	KACC 42090 = IBT 27920	Air conditioner, Inchen, Korea
	KACC 42091^T = IBT 27921	Air conditioner, Seoul, Korea
	KACC 41955 = CBS 117265= IBT 3016	Car air conditioner, Seoul, Korea
N. assulata	KACC 41691^T	Tomato soil, Buyeo, Korea
N. aurata	CBS 466.65^T	Jungle soil, Brunei
N. aureola	CBS 105.55^T	Soil, Tafo, Ghana
N. australensis sp. nov	CBS 112.55^T = NRRL 2392 = IBT 3021	Garden soil, Adelaide, Australia
N. coreana	KACC 41659^T = NRRL 35590 = CBS 121594	Tomato soil, Buyeo, Korea
N. denticulata	CBS 652.73^T = KACC 41183	Soil under Elaeis guineensis, Suriname
	CBS 290.74 = KACC 41175	Acer pseudoplatanus, Netherlands
N. fennelliae	CBS 598.74^T	Eye ball of Oryctolagus cuniculus, U.S.A.
	CBS 599.74	Eye ball of Oryctolagus cuniculus, U.S.A.
N. ferencziisp. nov.	CBS 121594^T = IBT 27813 = NRRL 4179	Soil, Australia
N. fischeri	CBS 544.65^T = NRRL 181	Canned apples
N. galapagensis	CBS 117522^T = IBT 16756 = KACC 41935	Soil, Ecuador
	CBS 117521 = IBT 16763 = KACC 41936	Soil, Ecuador
N. glabra	CBS 111.55^T	Rubber scrab from old tire, Iowa, U.S.A.
N. hiratsukae	CBS 294.93^T	Aloe juice, Tokyo, Japan
N. laciniosa	KACC 41657^T = NRRL 35589 = CBS 117721	Tomato soil, Buyeo, Korea
N. multiplicata	CBS 646.95^T = IBT 17517	Soil, Mouli, Taiwan
N. nishimurae	IFM 54133 = IBT 29024	Forest soil, Kenya
N. nishimurae	CBS 116047	Cardboard, Netherlands
N. papuensissp. nov.	CBS 841.96^T = IBT 27801	Bark of Podocarpus sp. (Podocarpaceae), bark, Myola, Owen Stanley Range, Northern Province, Papua New Guinea
N. pseudofischeri	NRRL 20748^T = CBS 208.92	Human vertebrate, U.S.A.
N. quadricincta	CBS 135.52^T = NRRL 2154	Cardboard, York, U.K.
	CBS 107078	Soil, Korea
	CBS 100942	Fruit juice, Netherlands
	CBS 253.94	Canned oolong tea beverage, Japan (type strain of N. primulina)
N. spathulata	CBS 408.89^T	Soil under Alocasia macrorrhiza, Taiwan
N. spinosa	CBS 483.65^T	Soil, Nicaragua
N. stramenia	CBS 498.65^T	Soil from maple-ash-elm forest, Wisconsin, U.S.A.
N. tatenoi	CBS 407.93^T	Soil of sugarcane, Timbauba, Brazil
	CBS 101754	Fruit, Yunnan, China (type strain of N. delicata)
N. udagawae	CBS 114217^T	Soil, Brazil
	CBS 114218	Soil, Brazil
N. warcupiisp. nov.	NRRL 35723^T	Arid soil, Finder”s Range, Australia

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CBS = Centraalbureau voor Schimmelcultures, Utrecht, the Netherlands; IBT = Institute for Biotechnology, Lyngby, Technical University of Denmark; IFM = Institute for Food Microbiology (at present, the Research Center for Pathogenic Fungi and Microbial Toxicoses, Chiba University), Chiba, Japan; KACC = Korean Agricultural Culture Collection, Suwon, Korea; NRRL = Agricultural Research Service Culture Collection, Peoria, Illinois, U.S.A.; T = type strain.

Morphology and physiology

The strains (Table 1) were grown for 7 d as 3-point inoculations on Czapek agar, Czapek yeast autolysate agar (CYA), oat meal agar (OA) and malt extract agar (MEA) plates at 25 °C, and on CYA at 37 °C. For Neosartorya species Hay infusion agar and SNA agar have also been used for inducing the anamorphs (medium compositions in Samson et al. 2004). In some species e.g. N. tatenoi the anamorph could only be produced when growing the cultures at 30 or 37 °C on MEA + 40 % sucrose.

Analysis for extrolites

Extrolites were analysed using the HPLC-diode array detection method of Frisvad & Thrane (1987, 1993) as modified by Smedsgaard (1997). Extrolites were analyzed from cultures grown on CYA, OA and YES agar using three agar plugs (Smedsgaard 1997).

Isolation and analysis of nucleic acids

Isolates used for the molecular studies were grown on 2 mL of malt peptone broth [10 % (v/v) malt extract (Brix 10) and 0.1 % (w/v) bacto peptone (Difco)], in 15 mL tubes. The cultures were incubated at 25 °C for 7 d. DNA was extracted from the cells using the Masterpure™ yeast DNA purification kit (Epicentre Biotechnol.) following the instructions of the manufacturer. Fragments containing the ITS region were amplified using primers ITS1 and ITS4 as described (White et al. 1990). Amplification of partial β-tubulin gene was performed using the primers Bt2a and Bt2b and methods of Glass & Donaldson (1995). Amplifications of the partial calmodulin and actin genes were as described (Hong et al. 2005, 2007). Sequencing reactions were performed with the Big Dye Terminator Cycle Sequencing Ready Reaction Kit and carried out for both strands. All the sequencing reactions were purified by gel filtration through Sephadex G-50 (Amersham Pharmacia Biotech, Piscataway, NJ) equilibrated in double-distilled water and analyzed on the ABI PRISM 310 Genetic Analyzer (Applied Biosystems). The complementary sequences were corrected with the MT Navigator software (Applied Biosystems). Unique ITS, β-tubulin, actin and calmodulin sequences were deposited in GenBank (http://www.ncbi.nlm.nih.gov) with accession numbers DQ534140, DQ534141 and EU20279-EU220287.

Data analysis

Sequence alignments were performed using CLUSTAL-X (Thompson et al. 1997) and improved manually. The neighbour-joining (NJ) method was used for the phylogenetic analysis. For NJ analysis, the data were first analysed using the Tamura-Nei distance calculation with gamma-distributed substitution rates (Tamura & Nei 1993), which were then used to construct the NJ tree with MEGA v. 3.1 (Kumar et al. 2004). A bootstrap analysis was performed with 1 000 replications to determine the support for each clade,.

PAUP v. 4.0 b10 software was used for parsimony analysis (Swofford 2002). Alignment gaps were treated as a fifth character state and all characters were unordered and of equal weight. Maximum parsimony analysis was performed for all data sets using the heuristic search option with random addition order (100 reps) and tree bisection-reconnection (TBR) branch-swapping algorithm. Branches of zero length were collapsed and all multiple, equally parsimonious trees were saved. The robustness of the trees obtained was evaluated by 1 000 bootstrap replications (Hillis & Bull 1993). Sequences from an A. clavatus isolate were used as outgroups in these experiments.

RESULTS AND DISCUSSION

Phylogenetic analysis

We examined the phylogenetic relatedness of species belonging to Aspergillus section Fumigati using sequence analysis of partial β-tubulin, calmodulin and actin genes including sequences of all known species. ITS sequences were determined from the new species and the species most closely related to them in the β-tubulin tree. The partial β-tubulin gene alignment included 453 characters. Among the polymorphic sites, 102 were found to be phylogenetically informative. The Neighbour-joining tree based on partial β-tubulin genes sequences is shown in Fig. 1. The topology of the tree is the same as one of the 419 maximum parsimony trees constructed by the PAUP programME (length: 465 steps, consistency index: 0.6710, retention index: 0.6467). The calmodulin data set included 549 characters with 85 parsimony informative characters. The Neighbour-joining tree shown in Fig. 2 has the same topology as one of the 9 maximum parsimony trees (tree length: 323, consistency index: 0.7585, retention index: 0.6422). The actin data set included 390 characters with 104 parsimony informative characters. The Neighbour joining tree shown in Fig. 3 has the same topology as one of the 312 maximum parsimony trees (tree length: 397, consistency index: 0.6675, retention index: 0.7130). The ITS data set included 501 characters with 26 parsimony informative characters. The Neighbour joining tree shown in Fig. 4 has the same topology as one of the 57 maximum parsimony trees (tree length: 77, consistency index: 0.7532, retention index: 0.7765).

Fig. 1. — Neighbour-joining tree based on β-tubulin sequence data of *Aspergillus* section *Fumigati*. Numbers above branches are bootstrap values. Only values above 70 % are indicated.

Fig. 2. — Neighbour-joining tree based on calmodulin sequence data of *Aspergillus* section *Fumigati*. Numbers above branches are bootstrap values. Only values above 70 % are indicated.

Fig. 3. — Neighbour-joining tree based on actin sequence data of *Aspergillus* section *Fumigati*. Numbers above branches are bootstrap values. Only values above 70 % are indicated.

Fig. 4. — Neighbour-joining tree based on ITS sequence data of selected species of *Aspergillus* section *Fumigati*. Numbers above branches are bootstrap values. Only values above 70 % are indicated.

The four Neosartorya isolates representing new species were found to be different from all known species of Aspergillus section Fumigati based on either their β-tubulin, calmodulin or actin gene sequences. However, one of them (NRRL 4179) had identical ITS sequences with N. denticulata (Fig. 4). This isolate was found to be closely related to a clade including N. fennelliae and N. denticulata on all other trees.

Possible synonymies of some species described previously have also been examined during this study. Based on multilocus sequence analyses Hong et al. (2007) discussed the synonymy of N. botucatensis, N. paulistensis and N. takaki with N. spinosa (Raper & Fennell) Kozak. (1972). N. spinosa and the synonyms have roughly circular arrangements of projections on the ascospore convex walls. N. spinosa produces echinulate ascospores with spines ranging from < 0.5 μm up to 5(-7) μm long with verruculose and small triangular projections or sometimes with circularly arranged projections.

N. otanii Takada, Y. Horie & Abliz (2001) was described on the basis of its rapid growth on Czapek and malt extract agars, lenticular ascospores with two widely separated equatorial crests, tuberculate or lobate-reticulate convex surface, and globose to broadly ellipsoidal conidia with a microtuberculate wall. The morphology of N. otanii resembles N. fennelliae, although Takada et al. (2001) reported small differences of the ascospore ornamentation, which was not confirmed in our SEM studies. The β-tubulin gene sequences of N. otanii (GenBank accession numbers AB201363 and AB201362) were identical with N. fennelliae (KACC 42228) (Fig. 5A). These N. fennelliae isolates produced ascospores after mating with the N. fennelliae type strains (data not shown). N. otanii is probably synonymous with N. fennelliae, but mating experiments with N. fennelliae and N. otanii are needed for its confirmation. These experiments could not be carried out because the ex type cultures of N. otanii were not available.

Fig. 5 — A. Neighbour-joining tree based on β-tubulin sequences showing the relationship of *N. otanii* and *N. fennelliae*. B. Neighbour-joining trees based on β-tubulin, calmodulin and actin sequence data of *Neosartorya* spp. showing the relationship of *N. primulina, N. quadricincta, N. tatenoi* and *N. delicata*.

Neosartorya primulina Udagawa, Toyaz. & Tsub. (1993) was characterised by its restricted growth on Czapek agar, chalky-buff ascomata, and lenticular ascospores with a very irregular ornamentation composed of several narrow crests and verrucose hemispheres. The ascospore ornamentation and anamorph morphology resembles those of N. quadricincta. Furthermore, the ex type culture (CBS 253.94) of N. primulina showed nearly identical sequences with strains of N. quadricincta for β-tubulin, calmodulin and actin genes (Fig. 5B). N. primulina is reduced to synonymy with N. quadricincta.

Neosartorya delicata H.Z. Kong (1997) was described based on its ellipsoid or nearly clavate vesicles, and ascospores with conspicuous spines, joining one spine to another by fairly prominent ridges and reticulate ornamentation, the ridges spreading to the equatorial crests. This species has identical ascospore morphology with N. tatenoi (Fig. 36), and both taxa were clustered into a clade in three gene trees (99.6 % in β-tubulin, 98.5 % in calmodulin and 97.3 % in actin gene sequences) (Fig. 5B). Therefore, we consider N. delicata as a synonym of N. tatenoi.

Fig. 36. — *Neosartorya tatenoi*. A-B. Colonies 14 d 25 °C. A. MEA. B. OA. C-E. Ascomata. F-G. Asci and ascospores. H. Ascospores. I. SEM of ascospores. J-L. Conidiophores. M. Conidia. Scale bars = 10 μm, except D = 30 μm, E = 15 μm̧ I = 1 μm.

Neosartorya nishimurae (Takada et al. 2001), N. indohii, N. tsurutae (Horie et al. 2003), N. takakii (Horie et al. 2001) and N. sublevispora (Someya et al. 1999) ex-type cultures were not available for this monograph of Aspergillus section Fumigati, and because we could not study them, they are listed as doubtful species.

Morphology and extrolite production

The atypical N. glabra isolate NRRL 4179 (Raper & Fennell 1965) produced asperfuran, aszonalenin, fumigaclavine, viridicatumtoxin, and fumigatins, extrolites common in N. fennelliae, but none of the extrolites produced by N. glabra. However, in contrast with the heterothallic N. fennelliae, this isolate is homothallic. It is closely related to N. denticulata based on phylogenetic analysis of sequence data, although their ascospore ornamentations are strikingly different (Figs. 21, 23). Ascospore ornamentation of NRRL 4179 is similar to that of the heterothallic N. fennelliae (Fig. 22) with equatorial crests much narrower, while N. denticulata has denticulate ascospores without equatorial crests. Isolate NRRL 4179 exhibited 72 % nuclear DNA relatedness to N. fennelliae and only 60 % relatedness to N. glabra isolates (Peterson 1992). This isolate also yielded different mtDNA and SmaI-digested repetitive DNA patterns from those of all the other Neosartorya strains examined (Rinyu et al. 2000). Hybridisation experiments were also carried out with Neurospora crassa mating type genes (the A idiomorph with about 6 kb flanking sequences, or the a idiomorph flanked by about 2 kb genomic DNA on either side) to the EcoRI digested DNA of several teleomorphic and asexual Aspergillus strains. Hybridisation to a 1.9 kb band was observed for both mating-type strains of N. fennelliae and isolate NRRL 4179 (Rinyu et al. 2000). Based on these observations, isolate NRRL 4179 seems to be closely related to N. fennelliae strains. These results are in agreement with those found using carbon source utilisation tests and isoenzyme analysis of these strains (Varga et al. 1997).

Fig. 21. — *Neosartorya denticulata*. A-B. Colonies 14 d 25 °C. A. OA. B. MEA. C-E. Ascomata. F-G. Asci and ascospores. H. Ascospores. I. SEM of ascospores. J-L. Conidiophores. M. Conidia. Scale bars = 10 μm, except D = 30 μm, E = 15 μm, I = 5 μm.

Fig. 23. — *Neosartorya ferencii*. A-B. Colonies 14 d 25 °C. A. OA. B. MEA. C-E. Ascomata. F-G. Asci and ascospores. H. Ascospores. I. SEM of ascospores. J-L. Conidiophores. M. Conidia. Scale bars = 10 μm, except D = 30 μm, E = 15 μm, I = 1 μm.

Fig. 22. — *Neosartorya fennelliae*. A-B. Colonies 14 d 25 °C. A. MEA. B-C. Crossing of mating types on MEA. D-E. Ascomata. F-G. Asci and ascospores. H. Ascospores. I. SEM of ascospores. J-L. Conidiophores. M. Conidia. Scale bars = 10 μm, except D = 30 μm, E = 15 μm, I = 1 μm.

Strain NRRL 35723 was isolated from soil in Australia, and produced compounds structurally related to wortmannin, aszonalenin, chromanols, tryptoquivalins and tryptoquivalons. This isolate was markedly different from all other known Neosartorya species in secreting a bluish pigment after 7 d incubation on MEA and CYA plates. The microtuberculate ascospore ornamentation of this isolate is similar to those of N. laciniosa, N. glabra and N. galapagensis (Hong et al. 2007). However, it grew more slowly on CYA than these species, and phylogenetic data also indicate that this isolate represents a new species.

CBS 112.55 was isolated from garden soil in Adelaide, Australia, and produced compounds similar to wortmannin and aszonalenin and some unique metabolites, while CBS 841.96 was isolated from Podocarpus bark in Papua New Guinea, and produced a compound related to wortmannins and some unique compounds the structures of which have not yet been elucidated (Table 2). The ascospore ornamentations of these isolates were microtuberculate, similarly to those of N. glabra and N. galapagensis. However, both isolates produced cream-coloured colonies on CYA in contrast with N. glabra which produces greyish green colonies. In phylogenetic analysis they were unrelated to any other Neosartorya species, justifying their treatment as new species. We propose four new homothallic and monotypic Neosartorya species; N. ferenczii (NRRL 4179), N. warcupii (NRRL 35723), N. australensis (CBS 112.55) and N. papuensis (CBS 841.96).

Table 2.

Extrolites produced by species assigned to Aspergillus section Fumigati.

Species	Extrolites produced
Aspergillus brevipes	roquefortine C, meleagrin-like
Aspergillus duricaulis	pseurotin A, fumagillin, asperpentyn, duricaulic acid and asperdurin, pthalides, chromanols, cyclopaldic acid, 3-O-methylcyclopolic acid
Aspergillus fumigatiaffinis	auranthine, cycloechinuline, fumigaclavines, helvolic acid, neosartorin, palitantin, pyripyropenes A, E, O & S, tryptoquivaline, tryptoquivalone
Aspergillus fumigatus	fumagillin, fumitoxins, fumigaclavines A & C, fumitremorgins, gliotoxin, trypacidin, pseurotins, helvolic acid, pyripyropens, methyl-sulochrin, verruculogen, fumiquinazolines
Aspergillus fumisynnematus	neosartorin, pyripyropens, fumimycin
Aspergillus lentulus	cyclopiazonic acid, pyripyropenes A, E & O, terrein, auranthine, neosartorin
Aspergillus novofumigatus	aszonalenin, cycloechinuline, fiscalins, helvolic acid, neosartorin, palitantin, terrein, territrem B
Aspergillus turcosus	kotanins and several unique but not yet elucidated secondary metabolites
Aspergillus unilateralis	mycophenolic acid, other unique secondary metabolites
Aspergillus viridinutans	viriditoxin, 13-O-methylviriditin, phomaligin A, variotin, viriditin, wasabidienone B0, B1, viriditin, 4-acetyl-6, 8-dihydroxy-5-methyl-2-benzopyran-1-1 A
Neosartorya assulata	indole alkaloids and apolar metabolites
Neosartorya aurata	helvolic acid, yellow unidentified compounds
Neosartorya aureola	fumagillin, tryptoquivaline, tryptoquivalone, pseurotin A and viriditoxin (FRR 2269 also produces helvolic acid)
Neosartorya australensis	wortmannin-like, aszonalenin-like
Neosartorya coreana	aszonalenins
Neosartorya denticulata	gliotoxin, viriditoxin
Neosartorya fennelliae	asperfuran, aszonalenin, fumigaclavine, viridicatumtoxin
Neosartorya ferenczii	asperfuran, aszonalenin, fumigaclavine, viridicatumtoxin, gliotoxin-like, fumigatins, aszonalenin-like
Neosartorya fischeri	terrein, fumitremorgins A & C, tryptoquivaline A, trypacidin, TR-2, verruculogen, sarcin, aszonalenins, fischerin, neosartorin, fiscalins, helvolic acid
Neosartorya galapagensis	gregatins
Neosartorya glabra	asperpentyn, avenaciolide, wortmannin-like compound
Neosartorya hiratsukae	avenaciolide
Neosartorya laciniosa	aszonalenins, tryptoquivaline, tryptoquivalone
Neosartorya multiplicata	helvolic acid
Neosartorya papuensis	wortmannin-like
Neosartorya pseudofischeri	asperfuran, cytochalasin-like compound, fiscalin-like compound, pyripyropens, gliotoxin
Neosartorya quadricincta	quinolactacin, aszonalenins
Neosartorya spinosa	aszonalenins, 2-pyrovoylaminobenzamide, pseurotin
Neosartorya spathulata	xanthocillins, aszonalenins
Neosartorya stramenia	quinolactacin, avenaciolide
Neosartorya tatenoi	aszonalenins
Neosartorya udagawae	fumigatin, fumagillin, tryptoquivaline, tryptoquivalone
Neosartorya warcupii	wortmannin-like, aszonalenin-like, chromanols-like, tryptoquivaline-like and tryptoquivalone-like

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Identification

Traditionally the identification of members of section Fumigati were done using the colony patterns and the morphology of the conidiogenous structures, conidia, ascomata and ascsopores. Ascospore ornamentation has been studied by Scanning electron microscopy, but our studies have shown that different species have similar ascospore shape and surface structure. Several species such A. fumigatus, A. novofumigatus, fumigatiaffinis, A. fumisynnematus and A. lentulus show strong morphological resemblance and in the lightmicroscope these species can be difficult to be separated. The anamorphs of Neosartorya udagawae and N. fennelliae also show a similar morphology. Therefore we recommend that for a correct species identification, sequence analysis should be carried out. Our experience with sequencing the calmodin and β-tubulin gen revealed good species delimitation and recognition. All sequences of the ex type cultures of section Fumigati are available from specialised databases and also from GenBank.

List of accepted species belonging to Aspergillus section Fumigati

The list of known species of Neosartorya and anamorphic species from the section Fumigati (Horie et al. 2003; Hong et al. 2005, 2006, 2007) is still expanding. With the species proposed here, there are now 23 Neosartorya species (including four new taxa) and 10 Aspergillus species in this group, 33 species in total and they are illustrated below.

Strict anamorphic species:

Aspergillus brevipes Smith

Aspergillus duricaulis Raper & Fennell

Aspergillus fumigatiaffinis Hong, Frisvad & Samson

Aspergillus fumigatus Fresenius

= A. anomalus Pidoplichko & Kirilenko
= A. fumigatus var. acolumnaris Rai et al.
= A. fumigatus var. ellipticus Raper & Fennell
= A. fumigatus mut. helvola Rai et al.
= A. phialiseptus Kwon-Chung
= A. neoellipticus Kozakiewicz
= Aspergillus arvii Aho, Horie, Nishimura & Miyaji

Aspergillus fumisynnematus Horie, Miyaji, Nishimura, Taguchi & Udagawa

Aspergillus lentulus Balajee & Marr

Aspergillus novofumigatus Hong, Frisvad & Samson

Aspergillus turcosus Hong, Frisvad & Samson

Aspergillus unilateralis Thrower

≡ A. brevipes var. unilateralis (Thrower) Kozakiewicz

Aspergillus viridinutans Ducker & Thrower

= A. fumigatus var. sclerotiorum Rai, Agarwal & Tewari

Teleomorph species:

Neosartorya assulata Hong, Frisvad & Samson [anamorph: A. assulatus Hong, Frisvad & Samson]

Neosartorya aurata (Warcup) Malloch & Cain [anamorph: A. igneus Kozakiewicz]

Neosartorya aureola (Fennell & Raper) Malloch & Cain [anamorph: A. aureoluteus Samson & Gams]

Neosartorya australensis Samson, Hong & Varga, sp. nov.

Neosartorya coreana Hong, Frisvad & Samson [anamorph: A. coreanus Hong, Frisvad & Samson]

Neosartorya denticulata Samson, Hong & Frisvad [anamorph: A. denticulatus Samson, Hong & Frisvad]

Neosartorya fennelliae Kwon-Chung & Kim [anamorph: A. fennelliae Kwon-Chung & Kim]

= Neosartorya otanii Takada, Horie & Abliz [anamorph: A. otanii Takada, Horie & Abliz]

Neosartorya ferenczii Varga & Samson, spec. nov.

Neosartorya fischeri (Wehmer) Malloch & Cain [anamorph: A. fischeranus Kozakiewicz]

Neosartorya galapagensis Frisvad, Hong & Samson [anamorph: A. galapagensis Frisvad, Hong & Samson]

Neosartorya glabra (Fennell & Raper) Kozakiewicz [anamorph: A. neoglaber Kozakiewicz]

Neosartorya hiratsukae Udagawa, Tsubouchi & Horie [anamorph: A. hiratsukae Udagawa, Tsubouchi & Horie]

Neosartorya laciniosa Hong, Frisvad & Samson [anamorph: A. laciniosus Hong, Frisvad & Samson]

Neosartorya multiplicata Yaguchi, Someya & Udagawa [anamorph: A. muliplicatus Yaguchi, Someya & Udagawa]

Neosartorya papuensis Samson, Hong & Varga, sp. nov.

Neosartorya pseudofischeri Peterson [anamorph: A. thermomutatus (Paden) Peterson]

Neosartorya quadricincta (Yuill) Malloch & Cain [anamorph: A. quadricingens Kozakiewicz]

= Neosartorya primulina Udagawa, Toyazaki & Tsubouchi [anamorph: A. primulinus Udagawa, Toyazaki & Tsubouchi]

Neosartorya spinosa (Raper & Fennell) Kozakiewicz [anamorph: A. spinosus Kozakiewicz]

≡ Aspergillus fischeri var. spinosus Raper & Fennell 1965 (basionym)
= Sartorya fumigata var. verrucosa Udagawa & Kawasaki
= Neosartorya botucatensis Horie, Miyaji & Nishimura [anamorph: A. botucatensis Horie, Miyaji & Nishimura]
= Neosartorya paulistensis Horie, Miyaji & Nishimura [anamorph: A. paulistensis Horie, Miyaji & Nishimura]
? = Neosartorya takakii Horie, Abliz & Fukushima [anamorph: A. takakii Horie, Abliz & Fukushima]

Neosartorya spathulata Takada & Udagawa [anamorph: A. spathulatus Takada & Udagawa]

Neosartorya stramenia (Novak & Raper) Malloch & Cain [anamorph: A. paleaceus Samson & Gams]

Neosartorya tatenoi Horie, Miyaji, Yokoyama, Udagawa & Campos-Takagi [anamorph: A. tatenoi Horie, Miyaji, Yokoyama, Udagawa & Campos-Takagi]

= Neosartorya delicata Kong [anamorph: A. delicatus Kong]

Neosartorya udagawae Horie, Miyaji & Nishimura [anamorph: A. udagawae Horie, Miyaji & Nishimura]

Neosartorya warcupii Peterson, Varga & Samson, sp. nov.

Doubtful species:

Neosartorya sublevispora Someya, Yaguchi & Udagawa [anamorph: A. sublevisporus Someya, Yaguchi & Udagawa]

Neosartorya indohii Horie [anamorph: A. indohii Horie]

Neosartorya tsurutae Horie [anamorph: A. tsurutae Horie]

Neosartorya nishimurae Takada, Horie & Abliz [anamorph: A. nishimurae Takada, Horie & Abliz]

Aspergillus brevipes Smith, Trans. Br. mycol. Soc. 35: 241. 1952. Fig. 6.

Type: CBS 467.91, from soil, New South Wales, Australia

Other no. of the type: ATCC 16899; CBS 118.53; IFO 5821; IMI 16034; IMI 51494; NRRL 2439; WB 4772 = IBT 22571; WB 4078 = IBT 22572

Description

Colony diam (7 d): CYA25: 12-15 mm; MEA25: 30-34 mm; YES25: 23-25 mm; OA25: 28-33 mm; CYA37: 16-19 mm; CREA: weak growth, no acid production

Colony colour: purple red

Conidiation: abundant

Reverse colour (CZA): dull yellow turning to reddish brown

Colony texture: velutinous

Conidial head: short columnar

Stipe: 15-50 (-100) μm, occasionally septate, heavy walled

Vesicle diam, shape: 10-18 μm, pear shaped

Conidium size, shape, surface texture: 2.8-3.5 μm, globose, spinulose

Cultures examined: CBS 467.91; WB 4772; WB 4078; CBS 118.523 = IBT 3051, all from the same original source

Diagnostic features: short heavy walled stipes, finely spinulose conidia, purple red colony colour, coloured vesicles and phialides and dark blue conidia; characterised by its vesicles borne at an angle to the stipe, as in A. viridinutans and A. duricaulis

similar species: A. duricaulis

Distribution: Australia

Ecology and habitats: soil

Extrolites: Roquefortine C, cf. meleagrin, red metabolite (not structure elucidated)

Pathogenicity: not reported

Note: previous reports on viriditoxin production of A. brevipes (Weisleder & Lillehoj 1971; Cole & Cox 1981) were based on studies of a mixed culture of A. brevipes and A. viridinutans (Peterson SW, pers. comm.)

Aspergillus duricaulis Raper & Fennell, The genus Aspergillus, 249. 1965. Fig. 7.

Type: CBS 481.65, from soil, Buenos Aires, Argentina

Other no. of the type: ATCC 16900; IMI 172282; JCM 01735; IBT 23177; NRRL 4021; VKM F-3572; WB 4021

Description

Colony diam (7 d): CYA25: 21-25 mm; MEA25: 20-22 m; YES25: 40-44 mm; OA25: 40-44 mm, CYA37: 21-25 mm, CREA: poor growth, no acid production

Colony colour: lily green to slate olive

Conidiation: heavy in central areas

Reverse colour (CZA): colourless to pinkish drab

Colony texture: velutinous

Conidial head: loosely columnar

Stipe: 5-50 × 3.5-5.5 μm, smooth thick walled

Vesicle diam, shape: 7-14 μm, flask shaped

Conidium size, shape, surface texture: (2.8-)3-3.3(-3.3) μm, globose, echinulate

Cultures examined: IMI 172282 = IBT 23177; CBS 481.65

Diagnostic features: echinulate conidia and weakly coloured reverse on CYA distinguish it from other anamorphic species

Similar species: A. brevipes

Distribution: Argentina

Ecology and habitats: soil

Extrolites: pseurotin A, fumagillin (found here), asperpentyn (Muhlenfeld & Achenbach 1988), duricaulic acid and asperdurin (Achenbach et al. 1985a), phthalides and chromanols (Achenbach et al. 1982a, 1985b), cyclopaldic acid and 3-O-methylcyclopolic acid (Brillinger et al. 1978; Achenbach et al. 1982b)

Pathogenicity: not reported

Aspergillus fumigatiaffinis Hong, Frisvad & Samson, Mycologia 97: 1326. 2005. Fig. 8.

Type: CBS 117186, from soil, Socorro County, Sevilleta National Wildlife Refuge, New Mexico, U.S.A..

Other no. of the type: KACC 41148; IBT 12703

Description

Colony diam (7 d): CYA25: 46-49 mm; MEA25: 53-60 mm; YES25: 67-74; CYA37: 65-70; CREA: weak griowth, good acid production

Colony colour: white, with center dull green

Conidiation: limited

Reverse colour (CZA): yellowish to greyish orange

Colony texture: floccose

Conidial head: short columnar

Stipe: 6-8 μm in diam.

Vesicle diam, shape: 18-24 μm, globose-subglobose

Conidium size, shape, surface texture: 2-3 μm, globose-subglobose, smooth

Diagnostic features: has comparatively small (sub)globose vesicles (16-24 μm); able to grow at 10 °C, and unable to grow at 50 °C

Similar species: A. fumigatus, A. lentulus, A. novofumigatus, A. fumigatiaffinis

Distribution: U.S.A., Spain

Ecology and habitats: kangaroo rat, soil, human

Extrolites: auranthine, cycloechinuline, fumigaclavines, helvolic acid, neosartorin, palitantin, pyripyropenes A, E, O & S, tryptoquivaline, tryptoquivalone

Pathogenicity: pathogenic to humans (Alcazar-Fuoli et al. 2007)

Note: exhibits high MICs to amphotericin B and several triazoles (Alcazar-Fuoli et al. 2007)

Aspergillus fumigatus Fresenius, Beitr. Mykol. 81: 18. 1863. Fig. 9.

= Aspergillus fumigatus var. acolumnaris Rai, Agarwal & Tewari (1971)
= Aspergillus fumigatus var. albus Rai, Tewari & Agarwal (1974)
= Aspergillus fumigatus var. cellulosae Sartory, Sartory & Mey. (1935)
= Aspergillus fumigatus var. coeruleus Malchevsk. (1939)
= Aspergillus fumigatus var. ellipticus Raper & Fennell (1965)
= Aspergillus fumigatus var. fulviruber Rai, Tewari & Agarwal (1974)
= Aspergillus fumigatus var. fumigatus Fresen. (1863)
= Aspergillus fumigatus griseibrunneus var. Rai & Singh (1974)
= Aspergillus fumigatus var. helvolus Yuill (1937)
= Aspergillus fumigatus var. lunzinense Svilv. (1941)
= Aspergillus fumigatus var. minimus Sartory (1919)
= Aspergillus neoellipticus Kozak. (1989)
= Aspergillus phialoseptus Kwon-Chung (1975)
= Aspergillus bronchialis Blumentritt (1901)
= Aspergillus septatus Sartory & Sartory (1943)
= Aspergillus arvii Aho, Horie, Nishimura & Miyaji (1994)

Type: IMI 016152, from chicken lung, Connecticut, U.S.A.

Other no. of the type: Thom 118; QM 1981; WB 163; CBS 133.61; NRRL 163; ATCC 1022; LSHB Ac71; NCTC 982; KACC 41143

Description

Colony diam (7 d): CYA25: 21-67 mm; MEA25: 25-69 mm; YES25: 48-74 mm; OA25: 34-62 mm, CYA37: 60-75 mm, CREA: poor growth, no or very weak acid production

Colour: greyish turquoise or dark turquoise to dark green to dull green

Conidiation: abundant, rarely less abundant

Reverse colour (CYA): creamy, yellow to orange

Colony texture: velutinous, st. floccose (define the abreviation st.)

Conidial head: columnar

Stipe: 50-350 × 3.5-10 μm

Vesicle diam, shape: 10-26 μm, pyriform to subclavate, sometimes subglobose, but rarely globose

Conidia length, shape, surface texture: 2-3.5(-6) μm, globose to ellipsoidal, smooth to finely rough

Cultures examined: ATCC 32722, AF71, AF 293, AF294, CBS 112389, CBS 487.65, CBS 133.61, CBS 545.65, CBS 457.75, CBS 542.75, CBS 113.26, CBS 110.46, CBS 120.53, CBS 132,54, CBS 123.59, CBS 158.71, CBS 180.76, CBS 143.89, CBS 148.89, CBS 488.90, CBS 287.95, CBS 100076, CBS 109032, CBS 386.75, CBS 286.95, CEA10, IMI 376380, NRRL 1979

Diagnostic features: Rapid growing velutinous colonies, abundant and fast conidiation, thick stipe (ca. 6-10 um), large pyriform to semi-clavate vesicle is representative morphological features of the species. However, the characteristics are various according to strains, and some stains have exceptional characteristics. The species grows at 50 °C, no growth at 10 °C.

Similar species: A. fumigatiaffinis, A. fumisynnematus, A. lentulus, A. novofumigatus, A. viridinutans.

Distribution: Worldwide distribution, cosmopolitan fungus (Pringle et al. 2005)

Ecology and habitats: soil, human

Extrolites: fumagillin, fumitoxins, fumigaclavines A & C, fumitremorgins, fumiquinazolines, gliotoxin, helvolic acid, pseurotins, pyripyropens, methyl-sulochrin, trypacidin, verruculogen

Pathogenicity: pathogenic to humans (Raper & Fennell 1965; Marr et al. 2002)

Note: no growth at 10 °C, growth at 50 °C; some isolates carry dsRNA mycoviruses (Anderson et al. 1996)

Aspergillus fumisynnematus Horie, Miyaji, Nishimura, Taguchi et Udagawa, Trans. Mycol. Soc. Japan: 34: 3-7. 1993. Fig. 10.

Type: IFM 42277, from soil, Sabaneta, Coro City, Falcon State, Venezuela

Description

Colony diam (7 d): CYA25: 44-48 mm; MEA25: 56-60 mm; YES25: 35-39 mm; OA25: 42-46; CYA37: 57-61 mm, CREA: poor growth and no acid production

Colony colour: greenish grey

Conidiation: limited

Reverse colour (CZA): orange white to orange grey

Colony texture: floccose

Conidial head: short columnar

Stipe: 210 × 6-8.5(-10) μm

Vesicle diam, shape: 16-20(-25) μm, hemispherical

Conidium size, shape, surface texture: 2.8-3.2 × 2.4-2.8 μm, broadly ellipsoidal, verruculose

Cultures examined: IFM 42277

Diagnostic features: production of synnemata on MEA with age (1-4-2.3 mm in height, 30-40 μm in diam.)

Similar species: A. fumigatus, A. lentulus, A. novofumigatus, A. fumigatiaffinis

Distribution: Brazil, Venezuela, Spain

Ecology and habitats: soil, human

Extrolites: neosartorin, pyripyropens (found here), fumimycin (Kwon et al. 2007)

Pathogenicity: pathogenic to humans (Alcazar-Fuoli et al. 2007; Yaguchi et al. 2007)

Note: growth at 10 °C, no growth at 50 °C

Aspergillus lentulus Balajee & Marr, Eukaryot. Cell 4: 631.2005. Fig. 11.

Type: FH5, from clinical specimens of patients hospitalised at the Fred Hutchinson Cancer Research Center, U.S.A.

Other no. of the type: KACC 41940, NRRL 35552; IBT 27201

Description

Colony diam: CYA25: (19-)25-56 mm, MEA25: (30)40-70 mm; YES25: 42-80 mm; OA25: 44-59 mm; CYA37: 54-70 mm, CREA: weak growth, no acid production

Colour: white with interspersed grey green conidia

Conidiation: usually poor, but abundant in some isolates

Reverse colour (CYA): pale yellow to grey orange, greyish brown

Colony texture: floccose

Conidial head: short columnar

Stipe: 20-500 × 4-7 μm, smooth, sometimes sinuous and constricted neck

Vesicle diam, shape: (6-)10-25 μm, globose to pyriform, usually subglobose

Conidia length, shape, surface texture: 2-3.2 μm, globose to broadly ellipsoidal, smooth to finely roughened

Cultures examined: KACC 41391 = CBS 116886, KACC 41392, KACC 41393, KACC 41681, KACC 41682, KACC 41642, KACC 41394, KACC 41395, KACC 41939 =FH7 = IBT 27209, KACC 41941 = FH4 = IBT 27210, KACC 41942 = FH220 = IBT 27202, KACC 41940 = FH5 = IBT 27201 = NRRL 35552

Diagnostic features: slow and poor conidiation, floccose colony texture, short columnar conidial heads, thin stipe (<7um), globose vesicle; growth at 10 °C and no growth at 50 °C

Similar species: A. fumigatiaffinis, A. fumigatus, A. fumisynnematus, A. novofumigatus, A. viridinutans

Distribution: Korea, U.S.A., Japan, Australia, Netherlands, Spain etc. It is assumed that the species is distributed worldwide.

Ecology and habitats: soil, human, dolphin

Extrolites: cyclopiazonic acid, pyripyropenes A, E & O, terrein, auranthine, neosartorin

Pathogenicity: pathogenic to humans (Balajee et al. 2005b; Alhambra et al. 2006; Alcazar-Fuoli et al. 2007; Yaguchi et al. 2007; Lau et al. 2007)

Note: exhibits high MICs to amphotericin B and several triazoles (Balajee et al. 2004, 2005b)

Aspergillus novofumigatus Hong, Frisvad & Samson, Mycologia 97: 1326. 2005. Fig. 12.

Type: CBS 117520, from soil, Galapagos Islands, Ecuador

Other no. of the type: IBT 16806

Description

Colony diam (7 d): CYA25: 33-48 mm; MEA25: 48-60 mm; YES25: 44-55 mm; OA25: 54-67 mm; CYA37: 49-52 mm; CREA: weak growth, no acid production

Colony colour: deep green to grey green

Conidiation: in central areas

Reverse colour (CZA): greyish orange to yellowish orange

Colony texture: velutinous

Conidial head: short columnar

Stipe: 50-500 × 4-7 μm in diam

Vesicle diam, shape: (13-)15-30 μm subglobose to flask shaped

Conidium size, shape, surface texture: 2.5-3 μm, ellipsoidal, smooth

Cultures examined: CBS 117520 = IBT 16806, CBS 117519 = IBT 16755

Diagnostic features: has nearly flask-shaped and comparatively large vesicles (15-30 mm); growth at 10 °C, no growth at 50 °C

Similar species: A. fumigatus, A. lentulus, A. fumisynnematus, A. fumigatiaffinis

Distribution: Galapagos Islands, Ecuador

Ecology and habitats: soil

Extrolites: aszonalenin, cycloechinuline, fiscalins, helvolic acid, neosartorin, palitantin, terrein, territrem B

Pathogenicity: not reported

Aspergillus turcosus Hong, Frisvad & Samson, Antonie van Leeuwenhoek (in press). Fig. 13.

Type: KACC 42091, from air conditioner, Seoul, South Korea

Other no. of the type: IBT 27921

Description

Colony diam: CYA25: 32-41 mm; MEA25: 42-53 mm; YES25: 48-52 mm; OA25: 46-52 mm; CYA37: 48-56; CREA poor growth, no acid production

Colony colour: grey-turquoise to grey-green

Conidiation: abundant

Reverse colour (CZA): yellowish orange to greyish orange

Colony texture: velutinous

Conidial head: short columnar

Stipe: 80-100 × 4-7 μm

Vesicle diam, shape: 15-25 μm, flask shaped to globose

Conidium size, shape, surface texture: 2.5-3.5 μm, subglobose, smooth

Cultures examined: KACC 42091 = IBT 27921, KACC 42090 = IBT 27920, KACC 41955 = IBT 3016

Diagnostic features: Velutinous colony, grey-turquoise (green) colony colour and yellowish orange reverse on MEA and CYA, phialides cover distal two-thirds of the vesicle and growth at both 10 and 50 °C

Similar species: -

Ecology and habitats: air conditioner

Distribution: South Korea

Extrolites: Kotanins and several unique compounds but not yet elucidated secondary metabolites

Pathogenicity: not reported

Aspergillus unilateralis Thrower, Austral. J. Bot. 2: 355. 1954. Fig. 14.

≡ A. brevipes var. unilateralis (Thrower) Kozakiewicz

Type: CBS 126.56, from rhizosphere of Hibbertia fasciculata and Epacris impressa, Australia

Other no. of the type: ATCC 16902; IFO 8136; IMI 062876; NRRL 577, QM 8163; WB 4366; WB 4779; IBT 3210

Description

Colony diam: CZA25: 30 mm; MEA25: 60-70 mm in 14 d, CRWEA: poor growth, no acid production

Colony colour: slate olive

Conidiation: limited

Reverse colour (CZA): nearly black

Colony texture: thin, brittle, folded in central area

Conidial head: diminutive, with few divergent spore chains

Stipe: 5-30 × 1.2-2.2 μm

Vesicle diam, shape: 4-8.5 μm, irregularly globose

Conidium size, shape, surface texture: 2.5-3.5 μm, globose, coarsely echinulate

Cultures examined: CBS 126.56; CBS 283.66 = IBT 3211

Diagnostic features: phialides clustered on one side of the vesicle, echinulate conidia, slow growth rate and dark reverse on CYA

Similar species: -

Distribution: Australia

Ecology and habitats: soil

Extrolites: mycophenolic acid, other unique secondary metabolites

Pathogenicity: not reported

Aspergillus viridinutans Ducker & Thrower, Austral. J. Bot. 2: 355. 1954. Fig. 15.

= A. fumigatus var. sclerotiorum J.N. Rai, S.C. Agarwal & J.P. Tewari

Type: CBS 127.56, from dung of rabbit, Frankston, Victoria, Australia

Other no. of the type: ATCC 16901; IMI 062875; IMI 062875ii; NRRL 4365; WB 4081; WB 4782; WB 4365

Description

Colony diam (7 d): CYA25: 20-40 mm; MEA25: 11-15 mm; YES25: 24-28 mm; OA25: 29-31 mm; CYA 37: 25-28 mm; CREA: poor griowth, no acid production

Colony colour: Niagara green

Conidiation: limited on CZA, abundant on MEA

Reverse colour: colourless (CZA), yellowish green to light brownish olive (MEA)

Colony texture: centre raised, velutinous on MEA

Conidial head: columnar

Stipe: 20-35 × 3.3-4.4 μm

Vesicle diam, shape: 7.5-12 μm, flask shaped to subglobose

Conidium size, shape, surface texture: 2-2.8 μm, globose, delicately roughened

Cultures examined: CBS 127.56

Diagnostic features: “nodding” conidial heads, Niagara green colony colour

Similar species: none

Ecology and habitats: soil, dung, human

Distribution: Australia, Sri Lanka, Zambia, Russia (Varga et al. 2000b)

Extrolites: viriditoxin, 13-O-methylviriditin, phomaligin A, variotin, viriditin, wasabidienone B0, B1, viriditin A (Omolo et al. 2000), 4-acetyl-6,8-dihydroxy-5-methyl-2-benzopyran-1-1 A (Aldridge et al. 1966)

Pathogenicity: pathogenic to humans (Katz et al. 2005, Yaguchi et al. 2007, Alcazar-Fuoli et al. 2007)

Notes: this is a highly variable species; further taxonomic studies needed to clarify the taxonomic position of the isolates assigned to it (Varga et al. 2000a, b); exhibits high MICs to some azoles (Alcazar-Fuoli et al. 2007)

Neosartorya assulata Hong, Frisvad & Samson [anamorph: A. assulatus Hong, Frisvad & Samson], Antonie van Leeuwenhoek (in press). Fig. 16.

Type: KACC 41691, from Tomato field soil, Buyeo, Korea

Other no. of the type: IBT 27911

Morphological characteristics

Colony diam (7 d): CYA25: (19-)37-41 mm;, MEA25: 47-58 mm; YES25: 28-31 mm; OA25: 36-40; CYA37: 32-68 mm

Colony colour: white

Conidiation: abundant

Reverse colour (CYA): yellowish white to pale yellow

Colony texture: radially sulcate

Conidial head: short columnar

Stipe: 3-7.5 μm wide

Vesicle diam, shape: 10-18 μm, subclavate

Conidium size, shape, surface texture: 2-3 μm, subglobose to ovoid, smooth

Homothallic

Cleistothecia: 120-250 μm, white to yellowish

Ascospores: 5-6 μm, lenticular, with two well-separated equatorial crests and convex surface decorated with several large, round flaps

Cultures examined: KACC 41691 = IBT 27911, IBT 27910

Diagnostic features: well developed long and round flaps on convex surface of ascospore with two distinct equatorial crests; grow on MEA and CZA much slower than N. pseudofischeri

similar species: N. pseudofischeri

Distribution: Korea

Ecology and habitats: soil

Extrolites: some indole alkaloids and some apolar metabolites

Pathogenicity: not reported

Neosartorya aurata (Warcup) Malloch & Cain [anamorph: A. igneus Kozakiewicz], Raper & Fennell 1965. Fig. 17.

Type: CBS 466.65, from jungle soil, Berakas, Muama, Brunei

Other no. of the type: ATCC 16894; IFO 8783; IMI 075886; IMI 075886ii; NRRL 4378; QM 7860; WB 4378; IBT 3028

Morphological characteristics

Colony diam (7 d): CYA25: 13-15 mm; MEA25: 30-42 mm; YES25: 17-29 mm; OA25: 31-35 mm; CYA37: 13-16 mm, CREA: weak growth and no acid production

Colony colour (MEA): orange to ochraceus orange

Conidiation: sparse

Reverse colour (CZA): orange to dull brown

Colony texture: velutinous

Conidial head: loosely columnar

Stipe: 60-120 × 2-4 μm

Vesicle diam, shape: 10-16 μm, flask shaped

Conidium size, shape, surface texture: 2.5-3 μm, globose, punctate

Homothallic

Cleistothecia: 50-150 μm, orange, surrounded by a loose tangle of encrusted orange hyphae

Ascospores: 6-6.5 × 4.5-5 μm, lenticular, with two narrow equatorial crests and convex walls finely reticulate

Cultures examined: CBS 466.65; WB 4379; IFO 9817

Diagnostic features: bright orange colour of the colony on MEA, restricted growth on CZA

Similar species: N. stramenia

Distribution: Brunei

Ecology and habitats: soil

Extrolites: helvolic acid, yellow unindentified compounds

Pathogenicity: not reported

Neosartorya aureola (Fennell & Raper) Malloch & Cain [anamorph: A. aureoluteus Samson & Gams], Mycologia 47: 71-75. 1955. Fig. 18.

Type: CBS 105.55, from soil, Tafo, Ghana

Other no. of the type: ATCC 16896; IFO 8105; IMI 061451; IMI 061451ii; MUCL 13579; NRRL 2244; QM 1906; WB 2244; IBT 3027

Morphological characteristics

Colony diam (7 d): CYA25: 64-80 mm; MEA25: 77-90 mm; YES25: 70-75 mm; OA25>: 55-59 mm; CYA37: 75-80 mm, CREA: poor growth, no acid prodution

Colony colour (CZA): apricot to light cadmium yellow

Conidiation: sparse

Reverse colour (CZA): yellow ocher to ochraceus

Colony texture: radially furrowed at center, slightly zonate

Conidial head: loosely columnar

Stipe: 50 × 2.5-4.5 μm

Vesicle diam, shape: 6-9 μm, clavate to flask shaped

Conidium size, shape, surface texture: 3-3.3 μm, globose to subglobose, delicately echinulate

Homothallic

Cleistothecia: 175-500 μm, pale lemon yellow, surrounded by loose wefts of dark golden yellow hyphae

Ascospores: 6-7 × 4.4-5 μm, lenticular, with two prominent equatorial crests and with convex surfaces conspicuously echinulate

Cultures examined: CBS 105.55; WB 2391

Diagnostic features: yellow to golden pigmentation of hyphae surrounding the cleistothecia

Similar species: N. udagawae, A. viridinutans

Distribution: Suriname, Ghana, Liberia, Fiji

Ecology and habitats: soil, canned passionfruit

Extrolites: fumagillin, tryptoquivaline, tryptoquivalone, pseurotin A and viriditoxin (FRR 2269 also produces helvolic acid)

Pathogenicity: not reported

Neosartorya australensis Samson, Hong & Varga, sp. nov. (Fig. 19) - MycoBank MB492203.

Homothallica; cleistothecia superficialia, luteoalba vel dilute lutea, globosa vel subglobosa, 150-380 μm diam, in hyphis hyalinis vel luteoalbis laxe obtextis. Asci octospori, globosi vel subglobosi, 12-14 μm diam, evanescentes. Ascosporae 4.5-7.5 μm diam, cristis angustis, aequatoriis binis, pagina convexa sublaevigata. Mycelium ex hyphis hyalinis, ramosis, septatis, laeviparietinis constans. Capitula conidialia curta, columnaria. Conidiophora ex hyphis aeriis exorientia, uniseriata, stipitibus 8-14 μm; vesiculae ampulliformes, 12-30 μm diam; phialides 7.5-9 × 2-3 μm, dimidium supernum vesiculae obtegentes. Conidia subglobosa vel ellipsoidea, laevia, 3.5-5 μm diam. Coloniae in agaro MEA in 7 dieibus et 25 °C celeriter crescentes, 40-45 mm diam, albae, capitulis conidialibus paucis. Coloniae in agaro CYA in 7 dieibus et 25 °C 30-35 mm diam, cremeoalbae, centro ab hyphis aerialibus laxe obtecto; capitula conidialia pauca; colonia reversa luteoalba vel luteobrunnea.

Holotype of Neosartorya australensis, here designated as CBS 112.55^T (dried culture), isolated from garden soil, Adelaide, Australia.

Homothallic, cleistothecia superficial, yellowish white to pale yellow, globose to subglobose, 150-380 μm in diam., surrounded by a loose covering of hyaline to yellowish white hyphae. Asci 8-spored, globose to subglobose 12-14 μm, evanescent at maturity. Ascospores lensshaped, 4.5-7.5 μm, with two equatorial crests, convex surfaces smooth to microtuberculate. Mycelium composed of hyaline, branched, septate, smooth-walled hyphae. Conidial heads short, columnar. Conidiophores arising from aerial hyphae often curling, uniseriate, stipes 12-30 μm; vesicles flask-shaped, 8-14 μm in diam.; phialides 7.5-9 × 2-3μm,, covering the upper half of vesicle. Conidia subglobose to ellipsoidal, smooth, 2.0-3.2 μm. Colonies on MEA growing rapidly, 40-45 mm in 7 d at 25 °C, white. Conidial heads produced few in number. Colonies on CYA, 30-35 mm in 7 d at 25 °C, creamy white, loosely overgrown by aerial hyphae in center. Conidial heads few in number. Reverse yellowish white to pale yellow.

Etymology: isolated from soil in Australia

Extrolites: wortmannin-like, aszonalenin-like

Distinguishing features: conidiophores often curled

Other no. of the type: IMI 061450; NRRL 2392; IBT 3021; WB 2392; Warcup SA14

Diagnostic features: smooth or microtuberculate 4.5-7.5 μm ascospores

Similar species: N. glabra

Distribution: Australia

Ecology and habitats: soil

Pathogenicity: not reported

Neosartorya coreana Hong, Frisvad & Samson [anamorph: A. coreanus Hong, Frisvad & Samson], Int. J. Syst. Evol. Microbiol. 56: 477. 2006. Fig 20.

Type: CBS 117059, from tomato field soil, Buyeo, Korea

Other no. of the type: KACC 41659 = NRRL 35590 = IBT 24945

Morphological characteristics

Colony diam (7 d): CYA25: 41-62 mm; MEA25: 57-66 mm; YES25: 50-74 mm; OA25: 54-58 mm; CYA37: 70-74 mm, CREA: poor growth, no acid production

Colony colour: white to yellowish white

Conidiation: sparse

Reverse colour (CYA): pale to light orange

Colony texture: radially sulcate

Conidial head: columnar

Stipe: 3-4 μm wide

Vesicle diam, shape: 8-13(-15) μm, subclavate

Conidium size, shape, surface texture: 2.5-3.5 μm, subglobose to broadly elliptical, smooth

Homothallic

Cleistothecia: 200-300 μm, white to light yellow

Ascospores: 4-5 μm, with two well-separated but often bent equatorial crests up to 2 μm, convex surface reticulate

Cultures examined: CBS 117059

Diagnostic features: rugose to weak reticulate ascospores with two often bent crests, but without the equatorial rings of small projections

Similar species: N. spinosa, N. laciniosa

Distribution: South Korea, Australia

Ecology and habitats: soil, strawberry

Extrolites: aszonalenins

Pathogenicity: not reported in humans (although isolated from the air sacks of an ostrich: Katz et al. 2005)

Neosartorya denticulata Samson, Hong & Frisvad [anamorph: A. denticulatus Samson, Hong & Frisvad], Antonie van Leeuwenhoek (in press). Fig. 21.

Type: CBS 652.73, from Soil under Elaeis guineensis, Suriname

Other no. of the type: KACC 41183

Morphological characteristics

Colony diam (7 d): CYA25: 22-24 mm; MEA25: 35-40 mm; CYA37: 35-38 mm; CREA: poor growth, no acid production

Colony colour: white

Conidiation: only on the marginal area

Reverse colour (CYA): yellowish white to pale yellow

Colony texture: loosely overgrown by aerial hyphae in the centre, sulcate in marginal areas

Conidial head: short columnar

Stipe: 3-4.5 μm wide

Vesicle diam, shape: 7-12 μm, spathulate

Conidium size, shape, surface texture: 2-3 μm, subglobose to broadly elliptical, smooth

Homothallic

Cleistothecia: 140-230 μm, yellowish white to pale yellow

Ascospores: 4-5 μm, denticulate with a prominent equatorial furrow

Cultures examined: CBS 652.73

Diagnostic features: denticulate ascospore surface and lacking equatorial crests make this a distinctive species

similar species: N. fennelliae, N. ferenczii

Distribution: Netherlands, Suriname

Ecology and habitats: soil, sycamore

Extrolites: gliotoxin, viriditoxin

Pathogenicity: not reported

Neosartorya fennelliae Kwon-Chung & Kim [anamorph: A. fennelliae Kwon-Chung & Kim], Mycologia 66: 628. 1974. Fig. 22.

Type: CBS 598.74 & CBS 599.74, from eye ball of Oryctolagus cuniculus, U.S.A.

Other no. of the type: ATCC 24325 & ATCC 24326, NRRL 5534 & NRRL 5535

Morphological characteristics

Colony diam (7 d): CYA25: 25-30 mm; MEA25: 44-48 mm; YES25: 30-34 mm; OA25: 34-38 mm; CYA37: 50-58 mm; CREA: poor growth and no acid production

Colony colour: grey

Conidiation: abundant

Reverse colour (CZA): white

Colony texture: velutinous

Conidial head: short columnar

Stipe: 150-250 × 4-6 μm

Vesicle diam, shape: 10-17 μm, flask-shaped

Conidium size, shape, surface texture: 2.2-2.5(-2.8) μm, globose to subglobose to ellipsoid, smooth or finely roughened

Heterothallic

Cleistothecia: 150-450 μm, white

Ascospores: 5.5-7.7 × 3.2-5 μm, with two equatorial crests, convex surfaces delicately roughened

Cultures examined: CBS 598.74, CBS 599.74

Diagnostic features: heterothallic

Similar species: N. denticulata, N. ferenczii

Distribution: U.S.A., Japan, South Korea

Ecology and habitats: soil, mirne sludge, rabbit

Extrolites asperfuran, aszonalenin, fumigaclavine, viridicatumtoxin

Pathogenicity: not reported in humans

Note: no growth at 47 °C

Neosartorya ferenczii Varga & Samson, sp. nov. (Fig. 23) - MycoBank MB504847.

Homothallica; cleistothecia superficialia, luteoalba vel dilute lutea, globosa vel subglobosa, 180-350 μm diam, in hyphis hyalinis vel luteoalbis laxe obtextis. Asci octospori, globosi vel subglobosi, 12-16 μm diam, evanescentes. Ascosporae 3.5-5.5 μm diam, cristis angustis, aequatoriis binis, pagina convexa sublaevigata. Mycelium ex hyphis hyalinis, ramosis, septatis, laeviparietinis constans. Capitula conidialia curta, columnaria. Conidiophora ex hyphis aeriis exorientia, uniseriata, stipitibus 100-150 × 4-5 μm; vesiculae ampulliformes, 10-14 μm diam; phialides 7.5-9 × 2-3 μm, dimidium supernum vesiculae obtegentes. Conidia globosa vel subglobosa, laevia, 2-2,5 μm diam. Coloniae in agaro MEA in 7 dieibus et 25 °C celeriter crescentes, 35-40 mm diam, albae, capitulis conidialibus paucis. Coloniae in agaro CYA in 7 dieibus et 25 °C 20-30 mm diam, cremeoalbae, centro ab hyphis aerialibus laxe obtecto; capitulis conidialibus paucis; colonia reversa luteoalba vel pallide lutea.

Holotype of Neosartorya ferenczii, here designated as CBS 121594^T (dried culture), isolated from soil in Australia.

Homothallic, cleistothecia superficial, yellowish white to pale yellow, globose to subglobose, 180-350 μm in diam., surrounded by a loose covering of hyaline to yellowish white hyphae. Asci 8-spored, globose to subglobose 12-16 μm, evanescent at maturity. Ascospores lens shaped, 3.5 × 5.5 μm, with two narrow equatorial crests, convex surface nearly smooth, microtuberculate. Mycelium composed of hyaline, branched, septate, smooth-walled hyphae. Conidial heads short, columnar. Conidiophores arising from aerial hyphae, uniseriate, stipes 100-150 × 4-5 μm; vesicles subclavate, 8-14 μm in diam; phialides 7.5-9 × 2-3 μm, covering the upper half of vesicle. Conidia globose to subglobose, smooth, 2-2,5 μm. Colonies on MEA growing rapidly, 35-40 mm in 7 d at 25 °C, white. Conidial heads produced few in number. Colonies on CYA, 20-30 mm in 7 d at 25 °C, creamish white, loosely overgrown by aerial hyphae in center. Conidial heads few in number. Reverse yellowish white to pale yellow (12A23) (Kornerup & Wanscher 1978).

Etymology: named after Prof. Lajos Ferenczy, eminent mycologist.

Extrolites: asperfuran, aszonalenin, fumigaclavine, viridicatumtoxin, gliotoxin-like, fumigatins and aszonalenin-like

Type: CBS 121594, from soil, Australia

Other no. of the type: IBT 27813, NRRL 4179; Warcup SA57

Diagnostic features: ascospore ornamentation similar to that of N. fennelliae, but with equatorial crests much narrower, and markedly different from those of N. denticulata

Similar species: N. fennelliae, N. denticulata

Distribution: Australia

Ecology and habitats: soil

Extrolites: asperfuran, aszonalenin, fumigaclavine, viridicatumtoxin, gliotoxin-like, fumigatins, and aszonalenin-like

Pathogenicity: not reported

Neosartorya fischeri (Wehmer) Malloch & Cain [anamorph: A. fischeranus Kozakiewicz], Can. J. Bot. 50: 2621. 1973. Fig. 24.

= Aspergillus fischeri Wehmer, Centr. Bakteriol. Parasitenk. Abt. II 18: 390. 1907.
= Sartorya fumigata Vuill., Compt. rendu Acad. Sci. Paris 184: 136. 1927.

Type: CBS 544.65, from canned apples, Wehmer

Other no. of the type: ATCC 1020; DSM 3700; IMI 211391; NRRL 181; QM 1983; Thom 4651.2, WB 181; IBT 3018

Morphological characteristics

Colony diam (7 d): CYA25: 45-68 mm; MEA25: 66-80 mm; YES25: 70-80 mm; OA25: 58-80 mm; CYA37: 65-84 mm; CREA: poor growth and no acid production

Colony colour (CZA): white to pale yellow to buff

Conidiation: sparse

Reverse colour (CZA): colourless to flesh coloured

Colony texture: velutinous

Conidial head: columnar

Stipe: 300-500 × 4-7 μm

Vesicle diam, shape: 12-18 μm, flask shaped

Conidium size, shape, surface texture: 2-2.5 μm, globose to subglobose, microtuberculate

Homothallic

Cleistothecia: up to 400 μm, light cream, borne singly or in small clusters within a loose hyphal envelope

Ascospores: 7-8 × 3-4 μm, convex surfaces bearing anastomosing ridges (reticulate)

Cultures examined: CBS 544.65; WB 4075; CBS 317.89; CBS 584.90; CBS 118441; NRRL 181; NRRL 4075; NRRL 4161; NRRL 4585

Diagnostic features: reticulate ascospore ornamentation

Similar species: N. tatenoi

Distribution: worldwide

Ecology and habitats: Soil, (milled) rice, cotton, potatoes, groundnuts, leather, paper products, canned products, human

Extrolites: terrein, fumitremorgins A & C, tryptoquivaline A, trypacidin, TR-2, verruculogen, sarcin, aszonalenins, fischerin, neosartorin, fiscalins, helvolic acid

Pathogenicity: pathogenic to animals and humans (Coriglione et al. 1990; Lonial et al. 1997; Mellado et al. 2006; Chim et al. 1998; Gori et al. 1998)

Neosartorya galapagensis Frisvad, Hong & Samson [anamorph: A. galapagensis Frisvad, Hong & Samson], Antonie van Leeuwenhoek (in press). Fig. 25.

Type: CBS 117522, from soil, Galapagos Islands, Ecuador

Other no. of the type: KACC 41935 = IBT 16756

Morphological characteristics

Colony diam (7 d): CYA25: 25-40 mm; MEA25: 26-35 mm; YES25: 39-44 mm; OA25: 34-41 mm; CYA37: 44-65 mm; CREA poor growth and no acid production

Colony colour: white

Conidiation: sparse

Reverse colour (CYA): golden yellow

Colony texture: strongly funiculose

Conidial head: columnar

Stipe: 2-4 μm wide

Vesicle diam, shape: 4-11 μm, (sub)clavate

Conidium size, shape, surface texture: 2.3-3 μm, globose to subglobose, smooth

Homothallic

Cleistothecia: 90-220 μm, yellowish white, surrounded by a loose covering of aerial hyphae

Ascospores: 5 μm, with two distinct equatorial crests 1-2 μm wide, convex surface of ascospores microtuberculate

Cultures examined: CBS 117522 = IBT 16756; CBS 117521 = IBT 16763

Diagnostic features: colonies funiculose, the Aspergillus anamorph arises from bundles of aerial hyphae, ascospores with two wide conspicuous equatorial crests and with microtuberculate convex surface

Similar species: N. glabra, N. australensis

Distribution: Galapagos Islands (Ecuador)

Ecology and habitats: soil

Extrolites: gregatins

Pathogenicity: not reported

Neosartorya glabra (Fennell & Raper) Kozakiewicz [anamorph: A. neoglaber Kozakiewicz], Mycol. Pap. 161: 56. 1989. Fig. 26.

Type: CBS 111.55, from rubber scrab of an old tire, Iowa, U.S.A.

Other no. of the type: ATCC 16909; IFO 8789; IMI 061447; IMI 061447ii; NRRL 2163; QM 1903; WB 2163

Morphological characteristics

Colony diam (7 d): CYA25: 24-43 mm; MEA25: 49-66 mm; YES25: 45-54 mm; OA25: 55-76 mm; CYA37: 30-80 mm; CREA: poor growth and no acid production

Colony colour (CZA): white to pale yellow to buff

Conidiation: sparse

Reverse colour (CZA): colourless to light pink

Colony texture: velutinous

Conidial head: columnar

Stipe: 300-500 × 4-7 μm

Vesicle diam, shape: 10-18 μm, flask shaped

Conidium size, shape, surface texture: 2.5-3.5 μm, globose to subglobose, microtuberculate

Homothallic

Cleistothecia: 100-500 μm, yellowish white

Ascospores: 6.5-7.5 × 4.5-5 μm, lenticular, with two equatorial crests of 1-1.5 μm, convex surfaces finely roughened

Cultures examined: CBS 111.55; IMI 144207; IMI 102073; CBS 165.63

Diagnostic features: has smaller and whiter cleistothecia and relatively straight equatorial crests and smoother walled convex surfaces compared to N. laciniosa, N. coreana and N. spinosa; N. glabra grows somewhat slower than the other species and grows well at comparatively low temperatures; can be distinguished from N. papuensis and N. australensis using sequence data or extrolite profiles

Similar species: N. papuensis, N. australensis

Distribution: U.S.A., Morocco, Denmark, Australia, Netherlands, South Korea

Ecology and habitats: soil, foods, indoor

Extrolites: asperpentyn, avenaciolide, wortmannin-like compound

Pathogenicity: not reported

Neosartorya hiratsukae Udagawa, Tsubouchi & Horie [anamorph: A. hiratsukae Udagawa, Tsubouchi & Horie], Trans. Mycol. Soc. Japan 32: 23. 1991. Fig. 27.

Type: NHL 3008, from pasteurised aloe juice, Tokyo, Japan

Other no. of the type: CBS 294.93; NRRL 20819

Morphological characteristics

Colony diam (7 d): CZA25: 14-15 mm; CYA25: 12-14 mm; MEA25: 26-39 mm; YES25: 42-45 mm; OA25: 42-45 mm; CYA37: 27-30 mm; CREA: rather poor growth and no acid production

Colony colour: greyish green

Conidiation: moderate

Reverse colour (CZA): light brown

Colony texture: velutinous

Conidial head: short columnar

Stipe: 120-380 × 5-7 μm

Vesicle diam, shape: 15-24 μm, flask-shaped

Conidium size, shape, surface texture: 2-2.5 μm, globose to subglobose, smooth or delicately roughened

Homothallic

Cleistothecia: 130-220 μm, light cream coloured

Ascospores: 4.5-5 μm, lenticular, with two closely appressed equatorial crests, convex surfaces finely reticulate

Cultures examined: CBS 294.93; IFM 50770 = IBT 27913

Diagnostic features: restricted growth on CZA, small cleistothecia, finely reticulate ascospores

Similar species: N. fischeri, N. tatenoi

Distribution: Japan, Brazil, South Korea

Ecology and habitats: soil, fruit juice, indoor air, human

Extrolites: avenaciolide

Pathogenicity: pathogenic to humans (Guarro et al. 2002; Mellado et al. 2006; Alcazar-Fuoli et al. 2007)

Note: no growth above 48 °C; some isolates carry dsRNA mycoviruses which are efficiently transmitted both through ascospores and conidia to the progeny (Varga et al. 1998)

Neosartorya laciniosa Hong, Frisvad & Samson [anamorph: A. laciniosus Hong, Frisvad & Samson], Int. J. Syst. Evol. Microbiol. 56: 477. 2006. Fig. 28.

Type: CBS 117721, from tomato field soil, Buyeo, Korea

Other no. of the type: NRRL 35589 = KACC 41657

Morphological characteristics

Colony diam (7 d): CYA25: 38-58 mm; MEA25: 53-67 mm; YES25: 60-78 mm; OA25: 52-59 mm; CYA37: 70-80 mm; CREA: poor growth and no acid production

Colony colour: white to pale yellow

Conidiation: sparse

Reverse colour (CYA): greyish orange to yellowish orange

Colony texture: sulcate, granular

Conidial head: columnar

Stipe: 3-4 μm wide

Vesicle diam, shape: 10-14 μm, subclavate

Conidium size, shape, surface texture: 2.5-3.5 μm, globose to subglobose, smooth

Homothallic

Cleistothecia: 300-400 μm, white to light yellow

Ascospores: 4-5 μm, broadly lenticular, with two distinct straight equatorial crests which are up to 2 μm

Cultures examined: CBS 117721; IBT 6660; KACC 41648; CBS 117719 = KACC 41652; KACC 41644

Diagnostic features: cleistothecia surrounded by a loose covering of hyaline to yellowish white, 2-4 μm wide hyphae; microtuberculate ascospores with two bent crests and two distinct equatorial rings of small projections

Similar species: N. spinosa, N. coreana

Distribution: South Korea, U.S.A., Pakistan, Netherlands, Suriname, Dominican Republic, Kenya

Ecology and habitats: soil

Extrolites: aszonalenins, tryptoquivaline, tryptoquivalone

Pathogenicity: not reported

Neosartorya multiplicata Yaguchi, Someya & Udagawa [anamorph: A. muliplicatus Yaguchi, Someya & Udagawa], Mycoscience 35: 309. 1994. Fig. 29.

Type: PF 1154, from soil, Taiwan

Other no. of the type: CBS 646.95, IBT 17517

Morphological characteristics

Colony diam (7 d): CYA25: 24-36 mm; MEA25: 35-50 mm; YES25: 38-42 mm; OA28-43 mm; CYA37: 41-80 mm, CREA: poor growth and no acid production

Colony colour: white

Conidiation: sparse

Reverse colour (CYA): greyish yellow to olivaceous buff

Colony texture: floccose

Conidial head: loosely columnar

Stipe: 20-160 × 2.5-4 μm

Vesicle diam, shape: 4-8 μm, flask-shaped to irregular

Conidium size, shape, surface texture: 2.5-4 μm, globose to subglobose, smooth

Homothallic

Cleistothecia: 100-300 μm, cream coloured

Ascospores: 4-5 μm, with a shallow furrow but without distinct equatorial crests, ornamented on surfaces by several linear ridges presenting ribbed or somewhat reticulate pattern

Cultures examined: CBS 646.95

Diagnostic features: can be distinguished from other species of Neosartorya by its almost globose ascospores, which have ribbed ornamentation with several linear ridges, and by the reduced production of its conidial heads on common media

Similar species: none

Distribution: Taiwan

Ecology and habitats: soil

Extrolites: helvolic acid

Pathogenicity: not reported

Neosartorya papuensis Samson, Hong & Varga, sp. nov. (Fig. 30) - MycoBank MB505571.

Homothallica; cleistothecia superficialia, luteoalba vel dilute lutea, globosa vel subglobosa, 200-350 μm diam, in hyphis hyalinis vel luteoalbis laxe obtextis. Asci octospori, globosi vel subglobosi, 14-20 μm diam, evanescentes. Ascosporae 5.5-7.5 μm diam, cristis angustis, aequatoriis binis, pagina convexa sublaevigata. Mycelium ex hyphis hyalinis, ramosis, septatis, laeviparietinis constans. Capitula conidialia curta, columnaria. Conidiophora ex hyphis aeriis exorientia, uniseriata, stipitibus 80-120 × 4-5 μm; vesiculae ampulliformes, 10-14 μm diam; phialides 7.5-9 × 2-3 μm, dimidium supernum vesiculae obtegentes. Conidia globosa vel subglobosa, laevia, 2-3 μm diam. Coloniae in agaro MEA in 7 dieibus et 25 °C celeriter crescentes, 35-40 mm diam, albae, capitulis conidialibus paucis. Coloniae in agaro CYA in 7 dieibus et 25 °C 20-30 mm diam, cremeoalbae, centro ab hyphis aerialibus laxe obtecto; capitula conidialia pauca; colonia reversa luteoalba vel pallide lutea.

Holotype of Neosartorya papuensis, here designated as CBS 841.96^T (dried culture), isolated from Podocarpus (Podocarpaceae), bark, Myola, Owen Stanley Range, Northern Province, Papua New Guinea.

Homothallic, cleistothecia superficial, yellowish white to pale yellow, globose to subglobose, 200-350 μm in diam., surrounded by a loose covering of hyaline to yellowish white hyphae. Asci 8-spored, globose to subglobose 14-20 μm, evanescent at maturity. Ascospores 5.5-7.5 μm, with two equatorial crests, convex surface smooth microtuberculate. Mycelium composed of hyaline, branched, septate, smooth-walled hyphae. Conidial heads short, columnar. Conidiophores arising from aerial hyphae, uniseriate, stipes 100-150 × 4-5 μm; vesicles flask-shaped, 10-14 μm in diam.; phialides 7.5-9 × 2-3 μm, covering the upper half of vesicle. Conidia globose to subglobose, smooth, 2-3 μm. Colonies on MEA growing rapidly, 35-40 mm in 7 d at 25 °C, white. Conidial heads few in number. Colonies on CYA, 30-35 mm in 7 d at 25 °C, producing sectors, creamy white, loosely overgrown by aerial hyphae in center. Conidial heads few in number. Reverse yellowish white to pale yellow (12A23) (Kornerup and Wanscher 1978).

Etymology: isolated in Papua New Guinea

Extrolites: wortmannin-like

Distinguishing features: smooth microtuberculate 5.5-7.5 μm, ascospores

Other no. of the type: IBT 27801

Cultures examined: CBS 841.96

Similar species: N. galapagensis, N. glabra, N. australensis

Distribution: Papua New Guinea

Pathogenicity: not reported

Neosartorya pseudofischeri Peterson [anamorph: A. thermomutatus (Paden) Peterson], Mycol. Res. 86: 547. 1992. Fig. 31.

Type: NRRL 20748, from human vertebrae, Atlanta, Georgia, U.S.A.

Other no. of the type: CBS 208.92

Holotype: 404.67, moldy cardboard, Victoria, British Columbia, Canada

Morphological characteristics

Colony diam (7 d): CYA25: 60-70 mm; MEA25: 90 mm in 7 d

Colony colour: white to pale creamish

Conidiation: sparse

Reverse colour (CZA): clear or faintly yellowish

Colony texture: velutinous

Conidial head: loosely columnar

Stipe: 200-300 × 4-7 μm

Vesicle diam, shape: 10-17 μm, subglobose

Conidium size, shape, surface texture: 3-4 μm, globose to subglobose, smooth

Homothallic

Cleistothecia: 150-300 μm, white

Ascospores: 4.5-6 μm, subglobose, with two equatorial crests of 1 μm wide, convex surfaces with raiased flaps resembling triangular projections

Cultures examined: CBS 208.92, CBS 404.67

Diagnostic features: distinctly ornamented ascospores

Similar species: -

Distribution: U.S.A., Canada, Netherlands, South Korea, Spain, Denmark, Estonia

Ecology and habitats: soil, indoor, human

Extrolites: asperfuran, cytochalasin-like compound, fiscalin-like compound, pyripyropens, gliotoxin

Pathogenicity: pathogenic to humans (Padhye et al. 1994; Matsumoto et al. 2002; Jarv et al. 2004; Balajee et al. 2005a; Alcazar-Fuoli et al. 2007; Lau et al. 2007) and animals (Barrs et al. 2007)

Neosartorya quadricincta (J.L. Yuill) Malloch & Cain [anamorph: A. quadricingens Kozakiewicz], Can. J. Bot. 50: 2621. 1973. Fig. 32.

= Neosartorya primulina Udagawa, Toyaz. & Tsub. [anamorph: A. primulinus Udagawa, Toyaz. & Tsub.]

Type: CBS 135.52, from cardboard, York, U.K.

Other no. of the type: ATCC 16897; IMI 048583; IMI 048583ii; NRRL 2154; QM 6874; WB 2154

Morphological characteristics

Colony diam (7 d): CYA25: 26-42 mm; MEA25: 52-59 mm; YES25: 36-59 mm; OA25: 47-55 mm; CYA37: 50-58 mm; CREA: poor growth and no acid production

Colony colour (CZA): white to light tan

Conidiation: sparse

Reverse colour (CZA): colourless to flesh coloured

Colony texture: floccose

Conidial head: loosely columnar

Stipe: 400-500 × 2-7 μm

Vesicle diam, shape: 10-20 μm, flask shaped

Conidium size, shape, surface texture: 2-3 μm, elliptical to globose, microtuberculate

Homothallic

Cleistothecia: up to 300 μm, buff to light tan

Ascospores: 4-5 μm, with two prominent equatorial crests, each duplicated by a some-what less prominent band, reticulate

Cultures examined: CBS 135.52; WB 2221; WB 4175; CBS 100942

Diagnostic features: presence of 4 equatorial crests on ascospores, reticulate ascospore ornamentation

Similar species: -

Distribution: Suriname, South Korea, U.K., Netherlands, Australia

Ecology and habitats: Soil, pectin, cardboard, fruit juice, mango pulp

Extrolites: quinolactacin, aszonalenins

Pathogenicity: not reported

Note: some isolates carry dsRNA mycoviruses (Varga et al. 1998)

Neosartorya spathulata Takada & Udagawa [anamorph: A. spathulatus Takada & Udagawa], Mycotaxon 24: 395. 1985. Fig. 33.

Type: CBS 408.89 & CBS 409.89, from cultivated soil under Alocasia macrorrhiza, Taiwan

Other no. of the type: IMI 308593& IMI 308593; NHL 2948, NHL 2949; NRRL 20549 & NRRL 20550

Morphological characteristics

Colony diam (7 d): CZA25: 33-38 mm, MEA25: 80 mm; OA25: 40-46 mm

Colony colour: greyish green

Conidiation: abundant

Reverse colour (CZA): uncoloured

Colony texture: velutinous

Conidial head: loosely columnar

Stipe: 500-1500 × 11-18(-25) μm and 60-250 × 4-10 μm

Vesicle diam, shape: 25-52 μm and 8-15 μm, flask-shaped

Conidium size, shape, surface texture: 3-5.5 × 2-4.5 μm, ellipsoidal, smooth

Heterothallic

Cleistothecia: 100-260 μm, pale yellow to light yellow

Ascospores: 3.5-4 μm, lenticular, with two equatorial crests, convex surfaces nearly smooth

Cultures examined: CBS 408.89 & CBS 409.89

Diagnostic features: yellowish cleistothecia, ascospores with large equatorial crests and smooth surface, two types of conidial heads (diminutive??)

Similar species: -

Distribution: Taiwan

Ecology and habitats: soil

Extrolites: xanthocillins, aszonalenins

Pathogenicity: not reported

Neosartorya spinosa (Raper & Fennell) Kozakiewicz [anamorph: A. spinosus Kozakiewicz], Mycol. Pap. 161: 58. 1989. Fig. 34.

≡ Aspergillus fischeri var. spinosus Raper & Fennell 1965 (basionym)
- = Sartorya fumigata var. verrucosa Udagawa & Kawasaki
- = Neosartorya botucatensis Y. Horie, Miyaji & Nishim. [anamorph: A. botucatensis Y. Horie, Miyaji & Nishim.]
- = Neosartorya paulistensis Y. Horie, Miyaji & Nishim. [anamorph: A. paulistensis Y. Horie, Miyaji & Nishim.]
- ? = Neosartorya takakii Horie, Abliz & K. Fukush. [anamorph: A. takakii Horie, Abliz & K. Fukush.]

Type: CBS 483.65, from soil, Nicaragua

Other no. of the type: ATCC 16898; IFO 8782; IMI 211390; NRRL 5034; WB 5034; IBT 3022

Morphological characteristics

Colony diam (7 d): CYA25: 41-70 mm; MEA25: 55-75 mm; YES25: 55-80 mm; OA25: 56-64 mm; CYA37: 67-85 mm; CREA: poor growth and no acid production

Colony colour (CZA): white to pale yellow to buff

Conidiation: sparse

Reverse colour (CZA): colourless to light pink

Colony texture: velutinous

Conidial head: columnar

Stipe: 300-500 × 4-7 μm

Vesicle diam, shape: 12-18 μm, flask shaped

Conidium size, shape, surface texture: 2-2.5 μm, globose to subglobose, microtuberculate

Homothallic

Cleistothecia: 200-300 μm, cartridge buff

Ascospores: 4.5 μm, with two widely separated equatorial crests, with convex surfaces bearing spinelike projections

Cultures examined: CBS 483.65

Diagnostic features: have echinulate ascospores with spines ranging from <0.5 μm up to 7 μm long, or with verruculose and small triangular, sometimes circularly arranged, projections

Similar species: N. coreana, N. laciniosa

Distribution: Nicaragua, Kenya, Denmark, Dominican Rebublic, U.S.A., Belgium, Sudasn, Japan, India, pakistan, South Korea

Ecology and habitats: Soil, fruit juice, human

Extrolites: aszonalenins, 2-pyrovoylaminobenzamide, pseurotin

Pathogenicity: pathogenic to humans (Summerbell et al. 1992; Mellado et al. 2006; Gerber et al. 1973)

Neosartorya stramenia (R.O. Novak & Raper) Malloch & Cain [anamorph: A. paleaceus Samson & Gams], Can. J. Bot. 50: 2622. 1972. Fig. 35.

Type: CBS 498.65, soil from maple-ash-elm forest, Wisconsin, U.S.A.

Other no. of the type: ATCC 16895; IFO 9611; IMI 172293; WB 4652

Morphological characteristics

Colony diam (7 d): CYA25: 10-40; MEA25: 40-59 mm; YES25: 58-62 mm; OA: 56-60 mm; CYA37: 45-49 mm; CREA: poor growth and no acid production

Colony colour (CZA): mustard-yellow

Conidiation: sparse

Reverse colour (CZA): yellow-orange

Colony texture: granulose

Conidial head: loosely columnar

Stipe: 80-140 × 3.5-5.5 μm, heavy walled, septate, coloured in terminal areas

Vesicle diam, shape: 10-12 μm, flask shaped to globose

Conidium size, shape, surface texture: 2.5-3 μm, globose, microverrucose

Homothallic

Cleistothecia: 50-175 μm, cartridge buff

Ascospores: 4.5-5.5 μm, with two widely separated flexuous equatorial crests, convex surfaces finely echinulate

Cultures examined: CBS 498.65; IFO 31358

Diagnostic features: faster growth rate and pronounced echinulate ascospore ornamentation distinguishes this species from N. aurata

Similar species: N. aurata

Distribution: U.S.A., Argentina

Ecology and habitats: Soil, salt grass (Distichlis scoparia)

Extrolites: quinolactacin, avenaciolide

Pathogenicity: not reported

Neosartorya tatenoi Horie, Miyaji, Yokoyama, Udagawa & Campos-Takagi [anamorph: A. tatenoi Y. Horie, M. Miyaji, K. Yokoy., Udagawa & Campos-Takagi], Trans. Mycol. Soc. Japan 33: 395. 1992. Fig. 36.

= Neosartorya delicata H.Z. Kong [anamorph: A. delicatus H.Z. Kong]

Type: CBM FA 0022, from soil, Brazil

Other no. of the type: CBS 407.93; IBT 21589

Morphological characteristics

Colony diam (7 d): CYA25: 35-39 mm mm; MEA25: 31-39 mm; YES25: 57-74 mm; OA25: 50-55 mm; CYA37: 72-78 mm; CREA: poor gowth and no acid production

Colony colour: pale yellow to yellowish white

Conidiation: sparse

Reverse colour (CZA): orange white to pale orange

Colony texture: velutinous to floccose

Conidial head: short columnar

Stipe: 270 × 4-7.5 μm

Vesicle diam, shape: 10-20 μm, hemispherical to flask-shaped

Conidium size, shape, surface texture: 2-3(-3.5) μm, globose to ovoid, smooth

Homothallic

Cleistothecia: 140-360 × 140-310 μm, hyaline to pale yellowish brown

Ascospores: 5-5.5 μm, lenticular, with two equatorial crests, convex surfaces with distinctly and nerrowly reticulate ridges

Cultures examined: CBS 407.93; NRRL 4584

Diagnostic features: distinct narrowly reticulate ascospore ornamentation

Similar species: N. fischeri, N. multiplicata

Distribution: Brazil, Dominican Republic

Ecology and habitats: soil

Extrolites: aszonalenins

Pathogenicity: not reported

Neosartorya udagawae Horie, Miyaji & Nishim. [anamorph: A. udagawae Horie, Miyaji & Nishim.], Mycoscience 36: 199. 1995. Fig. 37.

Type: CBM FA-0703 & CBM FA-0702, from soil, Brazil

Other no. of the type: CBS 114217 & CBS 114218

Morphological characteristics

Colony diam (7 d): CYA25: 33-36 mm; MEA25: 63-68 mm; YES25: 64-68 mm; OA25: 51-55 mm; CYA37: 61-65mm; CREA: poor growth and no acid production

Colony colour (CZA): dull green

Conidiation: abundant

Reverse colour (CZA): light orange to greyish orange

Colony texture: velutinous

Conidial head: columnar

Stipe: up to 530 × 4-6 μm

Vesicle diam, shape: 12-15 μm, hemispherical to flask shaped

Conidium size, shape, surface texture: 2.6-3.2 × 2.4-2.6 μm, subglobose to broadly ellipsoidal, smooth

Heterothallic

Cleistothecia: 310-620 × 280-530 μm, yellowish white to light yellow, surrounded by a loose covering of hyaline to pale yellowish brown hyphae

Ascospores: 5-5.5 × 4-5 μm, broadly lenticular, with two equatorial or often irregular crests, convex surfaces tuberculate

Cultures examined: CBS 114217, CBS 114218

Diagnostic features: heterothallic species, with characteristic tuberculate ascospore ornamentation

Similar species: N. aureola, A. viridinutans

Distribution: Brazil, U.S.A., Spain, Japan

Ecology and habitats: Soil, human

Extrolites: fumigatin, fumagillin, tryptoquivaline, tryptoquivalone

Pathogenicity: pathogenic to humans (Balajee et al. 2006; Moragues et al. 2006)

Neosartorya warcupii Peterson, Varga & Samson, sp. nov. (Fig. 38) - MycoBank MB505572.

Anamorph A. warcupiensis Peterson, Varga & Samson, sp. nov.

Homothallica; cleistothecia superficialia, alba vel dilute lutea, globosa vel subglobosa, 200-350 μm diam, in hyphis hyalinis vel luteoalbis laxe obtextis. Asci octospori, globosi vel subglobosi, 4.5-7 μm diam, evanescentes. Ascosporae 5.5-7 μm diam, cristis angustis, aequatoriis binis, pagina convexa sublaevigata. Mycelium ex hyphis hyalinis, ramosis, septatis, laeviparietinis constans. Capitula conidialia curta, columnaria. Conidiophora ex hyphis aeriis exorientia, uniseriata, stipitibus 100-150 × 4-5 μm; vesiculae ampulliformes, 10-25 μm diam; phialides 7.5-9 × 2-3 μm, dimidium supernum vesiculae obtegentes. Conidia subglobosa vel ellipsoidea, laevia, 1.8-1.5 μm diam. Coloniae in agaro MEA in 7 dieibus et 25 °C celeriter crescentes, 35-40 mm diam, albae, capitulis conidialibus paucis. Coloniae in agaro CYA in 7 dieibus et 25 °C 20-30 mm diam, cremeoalbae, centro ab hyphis aerialibus laxe obtecto; capitula conidialia pauca; colonia reversa luteobrunnea vel atrobrunnea.

Holotype of Neosartorya warcupii, here designated as NRRL 35723^T (dried culture), isolated from soil, Finder”s Range, Australia.

Homothallic, cleistothecia superficial, yellowish white to pale yellow, globose to subglobose, 180-350 μm in diam., surrounded by a loose covering of hyaline to yellowish white hyphae. Asci 8-spored, globose to subglobose 10-16 μm, evanescent at maturity. Ascospores lens shapedm 4.5-7 μm, with two prominent equatorial crests, convex surface smooth to microtuberculate. Mycelium composed of hyaline, branched, septate, smooth-walled hyphae. Conidial heads short, columnar. Conidiophores arising from aerial hyphae, uniseriate, stipes 100-150 × 4-6 μm; vesicles subclavate to subglobose, 12-18 μm in diam; phialides 7.5-9 × 2-3 μm, covering the upper half of vesicle. Conidia globose to subglobose, smooth, 1.8-2.5 μm. Colonies on MEA growing rapidly, 35-40 mm in 7 d at 25 °C. Colonies on CYA, 18-22 mm in 7 d at 25 °C, creamish white, sectors frequently produced. Conidial heads few in number. Reverse bluish in colour.

Etymology: named after Prof. J. H. Warcup, eminent mycologist, who isolated this culture.

Extrolites: wortmannin-like, aszonalenin-like, chromanol-like, tryptoquivaline-like and tryptoquivalone-like

Distinguishing features: secretes a blue pigment to the medium in 7-10 d; relatively slow growth on CYA at 25 °C

Distribution: Australia

Ecology and habitats: soil

Pathogenicity: not reported

Taxonomic novelties: Neosartorya australensis, N. ferenczii, N. papuaensis, N. warcupii.

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PERMALINK

Polyphasic taxonomy of Aspergillus section Fumigati and its teleomorph Neosartorya

RA Samson

S Hong

SW Peterson

JC Frisvad

J Varga

Abstract

INTRODUCTION

MATERIALS AND METHODS

Source of microorganisms

Table 1.

Morphology and physiology

Analysis for extrolites

Isolation and analysis of nucleic acids

Data analysis

RESULTS AND DISCUSSION

Phylogenetic analysis

Fig. 1.

Fig. 2.

Fig. 3.

Fig. 4.

Fig. 5.

Fig. 36.

Morphology and extrolite production

Fig. 21.

Fig. 23.

Fig. 22.

Table 2.

Identification

List of accepted species belonging to Aspergillus section Fumigati

Fig. 6.

Description

Fig. 7.

Description

Fig. 8.

Description

Fig. 9.

Description

Fig. 10.

Description

Fig. 11.

Description

Fig. 12.

Description

Fig. 13.

Description

Fig. 14.

Description

Fig. 15.

Description

Fig. 16.

Morphological characteristics

Fig. 17.

Morphological characteristics

Fig. 18.

Morphological characteristics

Fig. 19.

Fig. 20.

Morphological characteristics

Morphological characteristics

Morphological characteristics

Fig. 24.

Morphological characteristics

Fig. 25.

Morphological characteristics

Fig. 26.

Morphological characteristics

Fig. 27.

Morphological characteristics

Fig. 28.

Morphological characteristics

Fig. 29.

Morphological characteristics

Fig. 30.

Fig. 31.

Morphological characteristics

Fig. 32.

Morphological characteristics

Fig. 33.