Morphology & physiology: Colonies spreading; conidial colouren masse black or blackish brown; conidiophores uniseriate; no growth at 37 °C;
conidia 3–6μm.Sexual morphgenerally not produced in culture, except in a species described as “Saitoa japonica”(Rajendran & Muthappa 1980); sclerotia often present. Series description based on Samson et al.(2007b), Vargaet al.(2011)andHubka & Kolarik (2012).
Included species:Aspergillus aculeatinus,A.aculeatus,A.assiutensis (accepted species, unpubl. data, XC Wang),A.brunneoviolaceus, A.
floridensis, A. indologenus, A. japonicus, A. labruscus, A. saccha-rolyticus,A.serratalhadensis,A.trinidadensis,A.uvarum.
Extrolites: Aculene A, aflavinins, asperflavin, aspergillimide, calbistrin C, emodin, neopyranopnigrin, neoxaline, okaramin X, pre-aurantiamin, secalonic acid D.
SeriesNigriHoubraken & Frisvad,ser.nov.MycoBank MB832991.
Etymology: Named after the type species of the series,Aspergillus niger.
Type:Aspergillus nigerTiegh., Ann. Sci. Nat., Bot., ser. 5, 8: 240. 1867;
nom.cons. (Kozakiewiczet al.1992).
Diagnosis:Phylogeny: SeriesNigribelongs to subgen.Circumdati, sect.
Nigri and is phylogenetically sister to ser. Carbonarii (Fig. 13).
Morphology & physiology: Colonies spreading; conidial colouren masse black or blackish brown; conidiophores biseriate; good growth at 37 and 40 °C; conidia (2.5–)3–5μm.Sexual morphgenerally not produced in culture, except inA.tubingensis, saitoa-type, heterothallic (Hornet al.
0.1
A. neoniger A. costaricensis
91/*
A. vadensis
89/*
A. eucalypticola
A. pulverulentus (= A. tubingensis) A. tubingensis
87/*
A. luchuensis A. piperis
A. lacticoffeatus (= A. niger) A. niger
A. welwitschiae A. brasiliensis A. sclerotioniger
A. carbonarius A. sclerotiicarbonarius
A. ibericus A. ellipticus A. heteromorphus A. aculeatinus A. trinidadensis
A. brunneoviolaceus (syn. A. fjiensis) A. floridensis
84/*
A. aculeatus A. japonicus A. indologenus
A. uvarum A. serratalhadensis
A. labruscus A. saccharolyticus
91/*
A. homomorphus
P. expansum H. avellanea
ser. Nigri
ser. Carbonarii ser. Heteromorphi
ser. Japonici ser. Homomorphi
sect. Nigri
Fig. 13. Combined phylogeny forBenA,CaMandRPB2data sets showing the phylogenetic relation of species and series withinAspergillussect.Nigri. The BI posterior probability (pp) values and bootstrap percentages of the maximum likelihood (ML) analysis are presented at the nodes; fully supported branches are thickened. Values less than 70 % bootstrap support (ML) or less than 0.95 posterior probability (Bayesian analysis) are indicated with a hyphen or not shown. The bar indicates the number of substitutions per site. The phylogram is rooted withHamigera avellaneaandPenicillium expansum.
2013); sclerotia produced in most species. Series description based on Samsonet al.(2007b)andVargaet al.(2011).
Included species: Aspergillus brasiliensis, A. costaricensis, A. euca-lypticola,A.luchuensis,A.neoniger,A.niger,A.piperis,A.tubingensis, A.vadensis,A.welwitschiae.
Extrolites: Citric acid, oxalic acid, gluconic acid, tensyuic acids, atro-mentins, ochratoxins, funalenone, fumonisins, kotanins, yanuthones, naphtho-γ-pyrones, tensidols, malformins, nigragillins, pyranonigrins, asperazines, aflavinins (only in sclerotia) are shared by two or more species in ser.Nigri.
Notes on series in sect.Nigri:Frisvadet al.(2007)suggested a series (and subseries) classification for section Nigri using morphological, chemical and physiological features. Their suggested provisional series classificationfits with our suggested phylogenetic-based classification.
Fig. 13 shows the phylogenetic relationship among the species of sectionNigribased on partialBenA,CaMandRPB2gene sequencing.
These relationships are similar to those presented inVesthet al.(2018) using genome sequence data. In their manuscript, they distinguished the A. niger- and the A. tubingensis-clade. In contrast, Varga et al.
(2011) treated these two clades as one, named the A. niger-clade.
We follow Varga et al.(2011) in our concept of ser. Nigri, because species of this series share the ability to grow well at 37 and 40 °C and form biseriate conidiophores and similar-sized conidia measuring (2.5–) 3–5μm. Furthermore, the extrolites shared by two or more species in series Nigriare citric acid, oxalic acid, gluconic acid, tensyuic acids, atromentins, ochratoxins, funalenone, fumonisins, kotanins, yanuthones, naphtho-γ-pyrones, tensidols, malformins, nigragillins, pyranonigrins, asperazines, aflavinins (only in sclerotia) (Nielsenet al. 2009, Varga et al.2011, Frisvadet al.2018, Vesthet al.2018). SeriesCarbonarii is characterised by a generally weak or no growth at 37 °C (exceptA.
ibericus) and production of large conidia measuring 6–9μm. Extrolites shared by two or more species in ser.Carbonarii include citric acid, oxalic acid, gluconic acid, atromentins, ochratoxins, funalenone, naph-tho-γ-pyrones and pyranonigrins (Nielsenet al.2009, Vargaet al.2011).
Series Japoniciis a sister series of ser. Homomorphi. Series Homo-morphiinclude species that have biseriate conidiophores and produce the extrolites homomorphosins and decaturins, while ser. Japonici species have uniseriate conidiophores and produce aculenes, asper-paralines, calbistrins, neoxalines, okaramins, pre-aurantiamine, and/or secalonic acids. The phylogenetic relationship ofA.labruscus andA.
saccharolyticus with the other species of ser. Japonici is unclear (Fig. 13). In contrast to our results,Fungaroet al.(2017)classifiedA.
labruscus,A. homomorphusand A. saccharolyticus in the A. homo-morphus-clade and a similar result was shown inCrouset al.(2018b, Fungal Planet 720). Based on genome sequence analysis,A. saccha-rolyticusis sister to the other ser.Japonicispecies, confirming the result of our 3-gene phylogeny. Unfortunately, A. labruscus and A. serra-talhadensiswere not included in the genome study ofVesthet al.(2018) and genome sequencing of these species might reveal the correct classification of these species in the future.
SectionPetersoniorum [as“Petersonii”] Jurjevic & Hubka, Pl. Syst.
Evol. 301: 2449. 2015. MycoBank MB832533.
Type:Aspergillus petersoniiJurjevic & Hubka, Pl. Syst. Evol. 301: 2454.
2015.
Description: Phylogeny: Section Petersoniorum belongs to subgen.
Circumdati and the phylogenetic relationship with other Aspergillus sections needs to be resolved (see Notes below). Morphology &
physiology: Colonies restricted; conidial colouren massein shades of green; conidiophores biseriate; no growth 40 °C. Sexual morph un-known; sclerotia produced in most species (exceptA.asclerogenus), globose to ellipsoidal, pale yellow to brown. Also see Jurjevicet al.
(2015)(morphology, phylogeny).
Included species:Aspergillus arenarioides,A.asclerogenus,A. peter-sonii,A.peyronelii.
Notes on sect.Petersoniorum: Because no subdivision of sect. Peter-soniorumis proposed, ser.Petersoniorumis only informally introduced here. InFig. 12, this section is an early diverging clade in subgen.
Circumdati; however, a more thorough analysis places this section most close to sect.Candidi(Fig. 2) confirming the results ofJurjevicet al.
(2015). SectionsPetersoniorum andCandidican be differentiated by their differences in conidial and sclerotial colour (for more details, see sect.Candidi).
SectionRobustiJurjevic & Hubka, Pl. Syst. Evol. 301: 2460. 2015.
MycoBank MB814443.
Type:Aspergillus robustusM. Chr. & Raper, Mycologia 70: 200. 1978.
Description:Phylogeny: SectionRobustibelongs to subgen.Circumdati and is phylogenetically sister to sectionsTannerorumandCircumdati.
Morphology & physiology: Colonies restricted; conidial colouren masse in shades of yellow; conidiophores biseriate; no growth 37 °C.Sexual morphunknown; sclerotia produced, black (Christensen & Raper 1978).
See also:Jurjevicet al.(2015)(morphology, phylogeny).
Included species:Aspergillus robustus.
Notes: Because no subdivision of sect.Robustiis proposed, ser.Robusti is only informally introduced here. SectionRobustiis a single species section. This section is phylogenetically sister to sectionsTannerorum andCircumdati (Figs 2 and 12). It differs from both sections by the production of black coloured sclerotia and phototropic conidiophores (Visagieet al.2014c).
SectionTannerorum[as“Tanneri”] Jurjevic & Hubka, Pl. Syst. Evol.
301: 2460. 2015. MycoBank MB832534.
Type:Aspergillus tanneriKwon-Chunget al., J. Clin. Microbiol. 50: 3312.
2012.
Description: Phylogeny: Section Tannerorum belongs to subgen. Cir-cumdati and is sister to sect. Circumdati. Morphology & physiology:
Colonies restricted; sporulation sparse; conidiophores biseriate; good growth 37 °C.Sexual morphunknown; sclerotia not observed in culture.
Also seeJurjevicet al.(2015)(morphology, phylogeny).
Included species:Aspergillus tanneri.
Extrolites: No extrolites are reported forA.tanneri, the sole species in this section.
Notes: No subdivision of sect.Tannerorumis proposed and ser. Tan-nerorumis only informally introduced here (Table 4). Section Tannero-rumis a single species section. This section is phylogenetically sister to sect.Circumdati(Figs 2 and 12). It differs from this series by its small pyriform vesicles, lack of sclerotia, very poor sporulation, uncoloured reverse of colonies without production of soluble pigments and better growth at 37 °C than at 25 °C (Jurjevicet al.2015).
SectionTerreiW. Gamset al., Adv. Pen. Asp. Syst.: 59. 1986 [1985].
MycoBank MB832505.
Type:Aspergillus terreusThom, Amer. J. Bot. 5: 85. 1918.
Description: See Gams et al. (1985) (morphology), Samson et al.
(2011a)(morphology, phylogeny),Kocsubeet al.(2016)(phylogeny).
SeriesAmbiguiHoubraken & Frisvad,ser.nov.MycoBank MB832594.
Etymology: Named after the type species of the series,Aspergillus ambiguus.
Type:Aspergillus ambiguusSappa, Allionia 2: 254. 1955.
Diagnosis:Phylogeny: SeriesAmbiguibelongs to subgen.Circumdati, sect.Terreiand is phylogenetically sister to seriesNiveiandTerrei.
Morphology & physiology: Colonies restricted; conidial colour en massewhite, cream, dull yellow or grey-green; conidiophores biseri-ate. Sexual morph unknown. Series description based on Raper &
Fennell (1965).
Included species:Aspergillus ambiguus,A.microcysticus.
Extrolites: Butryolactone, terrequinone A.
SeriesNiveiHoubraken & Frisvad,ser.nov.MycoBank MB832595.
Etymology: Named after the type species of the series, Aspergillus niveus.
Type:Aspergillus niveusBlochwitz, Ann. Mycol. 27: 205. 1929.
Diagnosis: Phylogeny: Series Niveibelongs to subgen. Circumdati, sect.Terreiand is phylogenetically most closely related to ser.Terrei.
Morphology & physiology: Colonies growing moderately fast; conidial colouren masse white, vinaceous fawn, blue-green; conidiophores biseriate; moderate growth at 37 °C.Sexual morphunknown. Series description based on Raper & Fennell (1965) and Samson et al.
(2011a).
Included species:Aspergillus allahabadii,A.bicephalus,A.carneus,A.
iranicus,A.neoindicus,A.niveus.
Extrolites: Aszonalenins, citrinins are shared by ser.Niveispecies.
SeriesTerreiHoubraken & Frisvad,ser.nov.MycoBank MB832992.
Etymology: Named after the type species of the series, Aspergillus terreus.
Type:Aspergillus terreusThom, Amer. J. Bot. 5: 85. 1918.
Diagnosis: Phylogeny: Series Terrei belongs to subgen. Circumdati, sect.Terreiand is phylogenetically most closely related to ser. Nivei.
Morphology & physiology: Colonies spreading; conidial colouren masse (light) olive-brown; conidiophores biseriate; good growth at 37 °C.
Sexual morphgenerally not produced in culture, except inA.terreus, heterothallic, fennellia-type. Series description based onSamsonet al.
(2011a).
Included species: Aspergillus alabamensis, A. aureoterreus, A. cit-rinoterreus, A. floccosus, A. heldtiae, A. hortae, A. neoafricanus, A.
pseudoterreus,A.terreus.
Extrolites: SeriesTerreiextrolites include acetylaranotins, ardeemins, aspergillamides, aspergillicins, aspulvinones, asterriquinones, aszo-nalenins, butyrolactones, citreoviridins, citrinins, cytochalasins, geo-dins, gregatins, mevinolins, terrecyclic acids, terreic acid, terreins, terremides, terrequinones, terretonins and territrems (Samson et al.
2011a).
Notes on series in sect. Terrei: Three series are introduced in sect.
Terrei: seriesAmbigui,Nivei and Terrei. SeriesTerreiandNiveiare sister series and the species in those series differ in their conidial colour.
Species in ser.Terreigenerally produce conidia in brown shades, while ser.Niveiare in shades of yellow, vinaceous fawn or white. The two species of ser.Ambiguigrow slower than the taxa of seriesTerreiand Nivei.
Aspergillus subgen.Cremei Samson et al., Stud. Mycol. 85: 210.
2016. MycoBank MB819182.
Type:Aspergillus cremeusKwon-Chung & Fennell, Gen. Aspergillus:
418. 1965.
Description: SeeKocsubeet al.(2016)(morphology, phylogeny).
SectionCremeiW. Gamset al., Adv. Pen. Asp. Syst.: 61. 1986 [1985].
MycoBank MB832513.
Type:Aspergillus cremeoflavusSamson & W. Gams, Adv. Pen. Asp.
Syst.: 37. 1986 [1985]. MycoBank MB114701 (=Aspergillus cremeus).
Description: See Gams et al. (1985) (morphology, and partial sect.
Wentii, see Notes below), Kocsube et al. (2016), Fig. 2, this study (phylogeny).
Series Arxiorum Houbraken & Frisvad, ser. nov. MycoBank MB833044.
Etymology: Named after the type species of this series,A.arxii.
Type:Aspergillus arxii(Fort & Guarro) Houbrakenet al., Stud. Mycol. 78:
154. 2014.
Diagnosis:Phylogeny: SeriesArxiorumbelongs to sect.Cremei, sub-gen.Cremeiand is phylogenetically sister to a clade containing series Brunneouniseriati, Cremei and Wentiorum, though without statistical support (Fig. 14). Morphology & physiology: Colonies restricted; co-nidiophores and conidia not observed in culture. Sexual morph chaetosartorya-type, homothallic; ascospores hyaline, ellipsoidal, with two equatorial ridges, convex surfacesfinely rugose. Series description based onFort & Guarro (1984).
Included species:Aspergillus arxii.
Extrolites: No known extrolites have been found inAspergillus arxii.
SeriesBrunneouniseriatiHoubraken & Frisvad,ser.nov.MycoBank MB833045.
Etymology: Named after the type species of this series, A.
brunneouniseriatus.
Type:Aspergillus brunneouniseriatusSuj. Singh & B.K. Bakshi, Trans.
Brit. Mycol. Soc. 44: 160. 1961.
Diagnosis:Phylogeny: Series Brunneouniseriatibelongs to sect. Cre-mei, subgen.Cremeiand is phylogenetically sister to a clade containing seriesCremeiandWentiorum, though with moderate (BS = 70 %) or poor (pp < 0.95) statistical support (Fig. 14).Morphology & physiology:
Colonies growing rapidly; conidiaen massegrey, (dark) olive-brown or grey-green; conidiophores uniseriate, stipes hyaline, smooth. Sexual morphunknown. Series description based onRaper & Fennell (1965) andCrouset al.(2015).
Included species:Aspergillus brunneouniseriatus,A.citocrescens.
Extrolites: No known extrolites have been found in series Brunneouniseriati.
SeriesCremeiHoubraken & Frisvad,ser.nov.MycoBank MB833046.
Etymology: Named after the type species of this series,A.cremeus.
Type: Aspergillus cremeusKwon-Chung & Fennell, Gen. Aspergillus:
418. 1965.
Diagnosis: Phylogeny: SeriesCremeibelongs to sect. Cremei, sub-gen. Cremei and is phylogenetically sister to ser. Wentiorum (Fig. 14).Morphology & physiology: Colonies varying from restricted to spreading, more rapid growth on agar media with reduced water activity; conidiaen massein shades of green; conidiophores biseriate (A.cremeus,A.gorakhpurensis), uni- and biseriate (A.stromatoides) or strictly uniseriate (A. itaconicus), stipes hyaline, smooth, long.
Sexual morph unknown (A. gorakhpurensis, A. itaconicus, A. stro-matoides) or observed in culture (A.chaetosartoryae, A. cremeus), ascomata consisting of several layers of thick-walled hyphae, the outer layer becoming dematiaceous, homothallic; ascospores with prominent equatorial ridges, convex surface with spines. Series description based on Raper & Fennell (1965), Kamal & Bhargava (1969)andWiley & Simmons (1973).
Included species: Aspergillus chaetosartoryae, A. cremeus, A. gor-akhpurensis,A.itaconicus,A.stromatoides.
Extrolites: OnlyA.itaconicus has been examined for secondary me-tabolites according to the literature. This species can produce
chrysogine (reported here), itaconic acid (Kinoshita 1932, Steigeret al.
2013), itaconitin (Nakajimaet al.1964) and sorbicillins (reported here).
Series Inflati (Stolk & Samson) Houbraken & Frisvad, comb. nov.
MycoBank MB833047.
Basionym:Penicilliumser.InflataStolk & Samson, Adv. Pen. Asp. Syst.:
174. 1986 [1985].
Type:Penicillium inflatumStolk & Malla, Persoonia 6: 197. 1971. (syn.
Aspergillus inflatus).
Diagnosis:Phylogeny: SeriesInflatibelongs to sect.Cremei, subgen.
Cremeiand is phylogenetically sister to all other series in sect.Cremei.
Morphology & physiology: Colonies restricted (A.inflatus) or spreading (A. koreanus), conidia en masse green, greyish olive or grey-brown; conidiophores bi- or triseriate, or penicillium-like and biverti-cillate divaricate branched, stipe hyaline or pale reddish, smooth or finely roughened.Sexual morphunknown. Series description based on (Stolk & Malla 1971, Bissett & Widden 1984, Hydeet al.2016).
Extrolites:Aspergillus inflatus and a putative new species tentatively named A. oregonensis (CBS 576.95A&B) in ser. Inflatican produce sterigmatocystin (Rank et al. 2011). Otherwise sterigmatocystin (and aflatoxins) has only been found in the subgeneraCircumdati(sect.Flavi) andNidulantesin the genusAspergillus(Chenet al.2016a, Hubkaet al.
2016a, Frisvadet al.2019).
Included species:Aspergillus inflatus,A.koreanus,A.tardus.
SeriesPulviniHoubraken & Frisvad,ser.nov.MycoBank MB833048.
Etymology: Named after the type species of this series,A.pulvinus.
Type:Aspergillus pulvinus Kwon-Chung & Fennell, Gen. Aspergillus:
455. 1965.
Diagnosis:Phylogeny: SeriesPulvinibelongs to sect.Cremei, subgen.
Cremei and is phylogenetically sister to a clade containing series Arxiorum, Brunneouniseriati, Cremei and Wentiorum, though without statistical support (Fig. 14). Morphology & physiology: Colonies spreading, conidiaen masseblue-green; conidiophores biseriate, stipes brownish pigmented, roughened, long, thick-walled.Sexual morph un-known. Series description based onRaper & Fennell (1965).
Included species:Aspergillus pulvinus.
Extrolites: No known extrolites have been found inAspergillus pulvinus.
Series Wentiorum Houbraken & Frisvad, ser. nov. MycoBank MB833049.
Etymology: Named after the type species of this series,A.wentii.
Type:Aspergillus wentiiWehmer, Centralbl. Bakteriol., 2. Abth., 2: 149.
1896.
Diagnosis: Phylogeny: Series Wentiorum belongs to sect. Cremei, subgen. Cremei and is phylogenetically sister to ser. Cremei.
Morphology & physiology: Colonies growing restrictedly on MEA and CYA, growing moderately or rapidly on agar media with reduced water activity; conidiaen masse in shades of yellow-brown, olive-brown or greyish yellow; conidiophores biseriate, sometimes uni-seriate (A. flaschentraegeri), stipes hyaline, smooth or at most slightly roughened, often long and thick-walled; no growth at 37 °C.
Sexual morph unknown (A. europaeus, A. dimorphicus, A. fla-schentraegeri, A. wentii) or present (A.chrysellus), chaetosartorya-type, homothallic; ascospores hyaline, lenticular, with two equato-rial ridges, convex surfaces with spines. Series description based on Raper & Fennell (1965), Tuthill & Christensen (1986) and Hubka et al.(2016b).