Vargarum sect.
Clavati
sect.
Cervini B
Fig. 15. (Continued).
pseudoviridinutans, A. siamensis, A. udagawae, A. viridinutans, A.
wyomingensis. Aspergillus curviformis is tentatively included in this series based on its original description; no sequence data or material was available for this study.
Extrolites: Extrolite families found in ser.Viridinutantesinclude ardee-mins, aszonapyrones, chevalones, clavatols, cytochalasins, fiscalins, fumigaclavines, fumagillins, fumigatins, fumiquinazolins, gancidin, hel-volic acid, neosartoryadines, neosartoryones, phomaligins, pseurotins, pyripyropenes, trypacidins, sartorymensin, tryptoquivalines, viriditin, viriditoxin, wasabidienones. The cytochalasins, gancidin, neo-sartoryadines, neosartoryones, phomaligins, sartorymensin, and viriditin have only been found in ser.Viridinutantesin sect.Fumigati.
Notes on series in sect. Fumigati: Hubka et al. (2017) studied the phylogenetic relationship of sect.Fumigatispecies and recognised eight clades (A.brevipes-,A.fennelliae-,A.fumigatus-,A.neoglaber-,A. spi-nosus-, A. tatenoi-, A. unilateralis- and A. viridinutans-clade). More recently,Hubkaet al.(2018a)recognised three additional clades: theA.
thermomutatus-,A. spathulatus- and A. auratus-clade. Samson et al.
(2007a)were the last who studied the taxonomy of the whole section.
Based on this information, and data in more recent publications, it is difficult tofind good characters to delimit series in this section. The series classification is therefore mainly based on (published) phylogenetic data (Hubkaet al.2017, Hubkaet al.2018a, this study). Extrolites in sect.
Fumigati (with its eight families): ardeemins, asnovolins, aszonalenins, avenaciolides, cephalimycins, chaetominines, chevalones, chrysogines, clavatols, cycloechinulins, cyclopiazonic acids, cytochalasins, expanso-lides, fiscalins, fischerins, fumagillins,fumicyclins, fumigaclavines, fumi-gatins, fumigatonins, fumiquinazolines, fumitremorgins, gangicins, glabramycins, gliotoxins, helvolic acids, lentulins, neosartorins, novoa-maauromins, novobenzomalvins, pseurotins, pyripyropenes, sartorypyr-ones, sphingofungins terreins trypacidins, tryprostatins, tryptoquivalines, viridicatumtoxins, viriditoxins, wortmannins and several more (see above) (Honget al.2005, Larsenet al.2007, Samsonet al.2007a, Honget al.
2008, Hubkaet al.2013b, 2017, 2018a, Frisvad & Larsen 2015, Tamiya et al.2015, Bessaet al.2016, May Zinet al.2016, Rajachanet al.2016, Yuet al.2016, Banget al.2019, Xuet al.2019a, Yuet al.2019). Among these, the aszonalenin biosynthetic family (BF) is shared by six species, the pyrones (aszonapyrone, chavalones, sartorypyrones) are shared by five species, the helvolic acid biosynthetic family is shared byfive species, the pyripyropene BF is shared by four species, the gliotoxin BF is shared by four species, the fumigatin BF is shared by 4 species and the tryp-toquivaline BF is shared by four species. Since the secondary metabolite biosynthetic family members are distributed as polythetic characters, not every member of the species series may produce these extrolites.
Section Vargarum Houbraken & Frisvad, sect. nov. MycoBank MB832604.
Etymology: In honour of Janos Varga, a prominent Aspergillus researcher and advocate of a broad monophyleticAspergillusincluding polypaecilum-type morphs.
Type:Aspergillus cejpii (Milko) Samsonet al., Stud. Mycol. 78: 155.
2014.
Diagnosis:Phylogeny: SectionVargarumbelongs to subgen.Fumigati and is phylogenetically most closely related to sect.Clavati(Figs 2 and 15)Morphology & physiology: Colonies growing moderately fast; co-nidiophores polypaecilum-like; growth at 37 °C. Sexual morph neosartorya-type, homothallic, yellowish white.
Extrolites: Isolates in ser.Vargarumcan produce gliotoxin, rubratoxins, tryptoquivalones and xanthocillins (Vargaet al.2007a). These extrolites are also produced by other series in subgen.Fumigati, except rubra-toxins, which have been found only in this series inAspergillus, but is also produced by Talaromyces purpurogenus (Yilmaz et al. 2012) outsideAspergillus.
Included species:Aspergillus cejpii.
Notes: The polypaecilum-like asexual morph present in ser.Vargarumis unique in subgen. Fumigati. This morphology type is also found in species belonging toAspergillussubgen.Polypaecilum.
Aspergillussubgen.NidulantesW. Gamset al., Adv. Pen. Asp. Syst:
57. 1986 [1985]. MycoBank MB832501.
Type:Aspergillus nidulellusSamson & W. Gams, Adv. Pen. Asp. Syst.:
44. 1986 [1985] (=Aspergillus nidulans).
Description: See Gams et al. (1985) (morphology); Kocsube et al.
(2016), Figs 1 and 2, this study (phylogeny); Chen et al. (2016a) (morphology, phylogeny).
SectionAeneiVarga & Samson, IMA Fungus 1: 203. 2010. MycoBank MB517672.
Type:Aspergillus aeneusSappa, Allionia 2: 84. 1954.
Description:Phylogeny: SectionAeneibelongs to subgen.Nidulantes and is phylogenetically sister to sect. Nidulantes (Figs 2 and 16) Morphology & physiology: Colonies growing moderately or fast, conidia en massein shades of green or olive-brown; conidiophores biseriate, stipes brown pigmented, Hülle cells abundant (except inA. heyangen-sis), often in crusts, globose, subglobose or pyriform; no growth at or above 40 °C. Sexual morph not observed in culture, or present, emericella-type, homothallic; ascospore convex smooth or delicately roughened, with two equatorial crests. Section description based on Vargaet al.(2010a).
Included species:Aspergillus aeneus,A.bicolor,A.coloradensis*, A.
crustosus, A. discophorus, A. eburneocremeus, A. foeniculicola, A.
heyangensis,A.karnatakaensis,A.spectabilis[* not included inFig. 16;
details on classification, seeSklenaret al.(2020)].
Extrolites: Sterigmatocystin (5/9), decaturins (3/9), asperugins (2/9), asteltoxin (2/9), karnatakafurans (2/9), Mer-NF8054X (2/9), emeheteron (1/9), fumitremorgins (1/9), physcion (1/9), PR-toxin (1/9), quinolactacin (1/9), stellatin (1/9), terrein (1/9) (Mannicheet al.2004, Varga et al.
2010a).
Notes:Hubkaet al.(2016a)included the current sect.Aeneiin sect.
Nidulantesas theAspergillus aeneus-clade. Our nine-gene phylogenetic analysis locates representatives of sect.Aeneiwith statistical confidence outside sect. Nidulantes, confirming the results of previous reports (Varga et al. 2010a, Chen et al. 2016a). Phenotypically, the sexual species in sect.Aenei(A.bicolor,A.discophorus,A.foeniculicola,A.
spectabilis) produce similar ascospores as taxa in theA.nidulans-clade, but none of them are able to grow at 40 °C (Chenet al.2016a). There are no data indicating a subdivision of sect.Aeneiinto more than one series, and ser.Aeneiis therefore only informally introduced here (see Table 4).
Section Bispori S.W. Peterson et al. ex Houbraken, sect. nov.
MycoBank MB833244.
Synonym:Aspergillussect.Bispori Petersonet al., Aspergillus in the genomic era: 42. 2008;sect.inval.
Etymology: Named after the type species of this section,A.bisporus.
Type:Aspergillus bisporusKwon-Chung & Fennell, Mycologia 63: 479.
1971.
Diagnosis:Phylogeny: SectionBispori belongs to subgen.Nidulantes and is phylogenetically sister to a large clade containing sections Nidulantes, Aenei, Usti, Cavernicolarum, Raperorum and Silvatici.
Morphology & physiology: Colonies restricted or growing moderately fast, conidiaen masseolive to dark brown; conidiophores uniseriate, stipes slightly brown pigmented, smooth, occasionally showing granular pigment deposits; colonies growing more rapid at 37 °C than at 25 °C.
Sexual morphunknown. Section description based onKwon-Chung &
Fennell (1971)(underA.bisporus).
Included species:Aspergillus bisporus.
Extrolites: There are no known extrolites fromA.bisporus.
Notes:Aspergillus bisporuscould not be satisfactorilyfitted in any of the groups delineated by Raper & Fennell (1965). Based on morphological similarities, Samson (1979) classified the species in sect.Cervini. Molecular studies revealed that A. bisporusbelongs to subgen. Nidulantes (Peterson 2000, Peterson 2008, Peterson et al.
2008, Chen et al.2016a). SectionBispori was informally introduced (Petersonet al.2008), and this section is validated here. This section includes one species, A.bisporus, and introducing ser.Bispori would therefore be superfluous; ser. Bispori is only informally introduced here.
SectionCavernicolarum [as“Cavernicolus”’] A.J. Chenet al., Stud.
Mycol. 84: 112. 2016. MycoBank MB832535.
Type:Aspergillus cavernicolaLörinczi, Contrtii bot. Univ. Babes-Bolyai, Cluj, Grad. bot.: 341. 1969.
Description: SeeChenet al.(2016a)(morphology, phylogeny).
Series Cavernicolarum Houbraken & Frisvad, ser. nov. MycoBank MB833245.
Etymology: Named after the type species of this series,A.cavernicola.
Type:Aspergillus cavernicolaLörinczi, Contrtii bot. Univ. Babes-Bolyai, Cluj, Grad. bot.: 341. 1969.
Diagnosis: Phylogeny: Series Cavernicolarum belongs to sect. Cav-ernicolarum, subgen.Nidulantesand is phylogenetically sister to a clade containing ser. Egyptiaci.Morphology & physiology: Colonies growing restrictedly or moderately rapid, conidiaen massein shades of green or brown; conidiophores biseriate, short, occasionally long (in A. cal-ifornicus), stipes uncoloured or in brown shades, smooth, Hülle cells often present, usually globose or subglobose; no growth at 37 °C.
Sexual morphunknown. Series description based onRaper & Fennell (1965)andChenet al.(2016a).
Included species: Aspergillus californicus, A. cavernicola, A. kassu-nensis,A.subsessilis.
Extrolites: Red azaphilones of the same constitution as monascoru-bramin and N-glutarylrubropunctamin and similar extrolites found in Talaromyces albobiverticillius, Tal. atroroseus and Tal. purpurogenus (Yilmazet al.2012, Frisvadet al.2013b, Venkatachalamet al.2018) have been detected in A. cavernicola, including monasnicotinic acid (Antipovaet al.2018a).Aspergillus kassunensisproduces asperugins and Mer-NF8054X andA.californicusarugosins (Samsonet al.2011b).
No extrolites are common for the four species.
Series Egyptiaci Houbraken & Frisvad, ser. nov. MycoBank MB833246.
Etymology: Named after the type species of this series,A.egyptiacus.
Type:Aspergillus egyptiacusMoub. & Mustafa, Egypt. J. Bot. 15: 153.
1972.
Diagnosis: Phylogeny: Series Egyptiaci belongs to sect. Cav-ernicolarum, subgen.Nidulantesand is phylogenetically sister to a clade containing ser. Cavernicolarum. Morphology & physiology: Colonies restricted, conidiaen massein shades of green; conidiophores, mostly not arranged in typical Aspergillus heads when grown at 25 °C, biseriate, short, smooth; often also poorly developed conidiophores present: phialides solitary or occurring in little groups along the co-nidiophores resembling penicillium-like structures; Hülle cells produced, varying from globose to almost cylindrical; good growth on malt extract agar with 20 % or 40 % sucrose at 35 °C, minimum growth temperature 15 °C, optimum 35 °C and maximum 45 °C.Sexual morphunknown.
Series description based onSamson & Mouchacca (1974).
Included species:Aspergillus egyptiacus.
Extrolites: Fumitremorgin A & B, verruculogen and biosynthetically related indole-terpene-alkaloids are produced by A. egyptiacus (Samsonet al.2011b). These compounds are produced by species in both Aspergillus (different sections) and Penicillium (also different sections), but also by one species in sect. Nidulantes, ser. Stellati, namelyA.caespitosus(Schroederet al.1975, Steynet al.1981).
Notes: Section Cavernicolarum contains five species previously assigned to sect.Ustiand share the production of short stipes. Series Egyptiaciis phylogenetically distinct and the sole species in this section that grows well at 37 °C; ser.Cavernicolarumspecies do not grow at this temperature.
SectionNidulantesW. Gamset al., Adv. Pen. Asp. Syst.: 57. 1986 [1985]. MycoBank MB832502.
Type:Aspergillus nidulellusSamson & W. Gams, Adv. Pen. Asp. Syst.:
44. 1986 [1985]. MycoBank MB114711 (=Aspergillus nidulans).
Description: SeeGamset al.(1985), Chenet al.(2016a)(morphology, phylogeny),Kocsubeet al.(2016)(phylogeny).
Series Aurantiobrunnei Houbraken & Frisvad,ser. nov. MycoBank MB833247.
Etymology: Named after the type species of this series, A.
aurantiobrunneus.
Type:Aspergillus aurantiobrunneus(G.A. Atkinset al.) Raper & Fennell, Gen. Aspergillus: 511. 1965.
Diagnosis: Phylogeny: SeriesAurantiobrunnei belongs to sect. Nidu-lantes, subgen.Nidulantes and is phylogenetically sister of ser. Spe-luncei.Morphology & physiology: Colonies restricted, sporulation absent or sparse; conidiophores biseriate, smooth, stipes hyaline to pale brown;
Hülle cells present, hyaline to pale brown, globose, subglobose or ovoid;
no growth at 37 °C.Sexual morphemericella-type, homothallic; asco-spores globose to subglobose, with two equatorial crests, smooth convex. Series description based onChenet al.(2016a).
Included species:Aspergillus aurantiobrunneus,A.purpureus.
Extrolites: Epurpurins (including emerin), variecolactones (including emericolins, stellatic acid, variecoacetal, variecolin, variecolol), and ster-igmatocystin (including versicolorins) are shared by the two species in the ser.Aurantiobrunnei. Eremophiline has been found inA.aurantiobrunneus and calbistrins and shamixanthones has been found inA.purpureus(Chen et al.2016a).
Series Multicolores Houbraken & Frisvad, ser. nov. MycoBank MB833248.
Etymology: Named after the type species of this series,A.multicolor.
Type:Aspergillus multicolorSappa, Allionia 2: 87. 1954.
Diagnosis:Phylogeny: SeriesMulticoloresbelongs to sect.Nidulantes, subgen. Nidulantes and is phylogenetically sister of ser. Nidulantes.
Morphology & physiology: Colonies growing moderately fast or spreading, conidia en masse in shades of green; conidiophores biseriate, smooth, stipes hyaline to yellowish brown; Hülle cells absent (A.mulundensis,A.purpureocrustaceus) or present (A.incahuasiensis, A.multicolor,A.pluriseminatus,A.tumidus), pale yellowish brown, or-ange, brown to pink, globose, subglobose or ovoid.Sexual morphnot observed in culture, or present (A. pluriseminatus), emericella-type, homothallic; ascospores lenticular, convex surface tuberculate under SEM, with two conspicuously pleated, stellate and striate equatorial crests. Series description based on Stchigel & Guarro (1997), Chen et al.(2016a), Crouset al.(2018b), Piontelliet al.(2019).
Included species:Aspergillus incahuasiensis*,A.multicolor,A. mulun-densis, A. pluriseminatus, A.purpureocrustaceus*, A. tumidus [* not included inFig. 16; details on classification, seeSklenaret al.(2020)].
A. acristatus A. quadrilineatus
A. sublatus A. nidulans
91/*
A. foveolatus
79/0.99
A. corrugatus
93/*
A. rugulosus A. pachycristatus A. striatus
A. spinulosporus
A. spinulosporus (non-type) A. sulphureoviridis
A. violaceus A. omanensis A. jaipurensis A. falconensis
A. fruticulosus A. latilabiatus
A. recurvatus A. navahoensis A. aurantiopurpureus A. desertorum A. stercorarius A. savannensis A. botswanensis
77/0.99
A. mulundensis A. multicolor
A. tumidus A. pluriseminatus
A. israelensis A. unguis A. croceus
*/0.99
A. dromiae A. angustatus
A. stellatus A. venezuelensis A. stella-maris A. astellatus
A. miraensis
73/0.99
A. qinqixianii A. filifer A. olivicola
A. undulatus A. caespitosus
83/0.99
A. versicolor A. fructus A. pepii A. tabacinus
78/*
A. amoenus
70/*
A. griseoaurantiacus
91/*
A. austroafricanus A. hongkongensis
73/0.99
A. protuberus
90/--/0.98