*/-
96/-A. germanicus A. thesauricus A. keveii A. keveioides A. insuetus A. asper
85/*
A. fuscicans A. pseudodeflectus
80/0.97
A. calidoustus A. contaminans
A. carlsbadensis A. baeticus A. ustus
A. porphyreostipitatus A. pseudoustus A. puniceus
A. heterothallicus A. granulosus
A. monodii A. lucknowensis
A. deflectus A. collinsii
71/-A. turkensis A. elongatus
A. subsessilis A. kassunensis A. californicus
A. cavernicola (syn. A. amylovorus) A. egyptiacus
97/-A. silvaticus
96/-
87/--/*
-/0.97
-/0.98
-/*
ser. Cavernicolarum ser. Egyptiaci ser. Silvatici ser. Versicolores
ser. Speluncei ser. Aurantiobrunnei
ser. Aenei
ser. Calidousti
sect.
Usti
ser. Monodiorum
ser. Deflecti
sect.
Aenei
sect.
Cavernicolarum sect. Silvatici ser. Usti
B
Fig. 16. (Continued).
Extrolites:Aspergillus multicolor,A.mulundensisandA.pluriseminatus all produce asticolorins or closely related dibenzofurans. Aspergillus mulundensisproduces mulundocandins and emericellamide, and aza-philones in common withA.pluriseminatus(Billset al.2016, Chenet al.
2016a), whileA.multicoloris the only species in the series that has been reported to produce sterigmatocystins (Hamasakiet al.1980).
Series Nidulantes Houbraken & Frisvad, ser. nov. MycoBank MB833249.
Etymology: Named after the type species of this series,A.nidulellus.
Type:Aspergillus nidulellusSamson & W. Gams, Adv. Pen. Asp. Syst.:
44. 1986 [1985]. MycoBank MB114711 (=Aspergillus nidulans).
Diagnosis:Phylogeny: SeriesNidulantesbelongs to sect. Nidulantes, subgen.Nidulantes and is phylogenetically sister of ser.Multicolores.
Morphology & physiology: Colonies spreading, conidiaen massegreen;
conidiophores biseriate, stipes hyaline to yellowish brown pigmented, smooth, occasionally with surface protuberances; Hülle cells present, globose; good growth at 37 °C and 40 °C, growth at 45 °C, except forA.
botswanensis,A.fruticulosus,A.latilabiatusandA.recurvatus.Sexual morph generally present (only not observed in A. recurvatus), emericella-type, homothallic; ascospores irregularly wrinkled, finely pitted, rugulose or echinulate, with two equatorial crests (except for four crests in A. quadrilineatus). Series description based on Chenet al.
(2016a).
Included species:Aspergillus amethystinus*,A.aurantiopurpureus,A.
botswanensis,A.corrugatus,A.desertorum,A.dipodomyus,A. falco-nensis,A.foveolatus,A.fruticulosus,A.jaipurensis,A.latilabiatus,A.
navahoensis,A.nidulans,A.omanensis,A.pachycristatus,A. quad-rilineatus,A.recurvatus,A.rugulosus,A.savannensis,A. spinulospo-rus, A. stercorarius, A. striatus, A. sublatus, A. sulphureoviridis, A.
violaceus[* not included in Fig. 16; details on classification, see Sklen]?
et al. (2020)].
Extrolites(number of species producing compound / total species in se-ries): Asperthecin (14/22), asperugins (10/22), austinols (2/22), austalides (5/22), calbistrins (2/22), cordycepin (1/22), cyclopaldic acid (1/22), desertorins (6/22), echinocandins (3/22), emericellin (9/22), emerin/epur-purins (2/22), emestrin (6/22), emindols (7/22), falconensins (6/22), fal-conensons (5/22), gregartins (2/22), 2-ω-hydroxyemodin (4/22), isocoumarins (1/22), paxillin (6/22), quadrilineatin (and the related nidulol
& silvaticol) (2/22), shamixanthones (10/22), sterigmatocystin (14/22,A.
latilabiatusproducing only versicolorins), terrequinone A (4/22), violaceols (16/22), viridicatumtoxin (1/22) (Chenet al.2016a).
Series Speluncei Houbraken & Frisvad, ser. nov. MycoBank MB833250.
Etymology: Named after the type species of this series,A.spelunceus.
Type:Aspergillus spelunceusRaper & Fennell [as“speluneus”], Gen.
Aspergillus: 457. 1965.
Diagnosis: Phylogeny: SeriesSpeluncei belongs to sect. Nidulantes, subgen. Nidulantes and is phylogenetically sister of ser. Aurantio-brunnei. Morphology & physiology: Colonies restricted or moderately fast, sporulation sparse or moderate,en masse in shades of green;
conidiophores biseriate, stipes hyaline or yellowish brown, smooth; Hülle cells absent or present (A. askiburgiensis, A. asperescens, A. spe-lunceus), hyaline, globose, subglobose or ovoid; no growth at 37 °C (exceptA.asperescens). Sexual morph unknown. Series description based onChenet al.(2016a).
Included species:Aspergillus askiburgiensis,A.asperescens,A. aur-eolatus,A.spelunceus,A.varians,A.viridicatenatus.
Extrolites(number of species producing compound / total species in series): Azaphilones (1/6), austinols (1/6), calbistrins (1/6), desertorins (1/6), emerin / epurpurins (2/6), 2-ω-hydroxyemodin (1/6), a phthalide (1/6), shamixanthones (1/6), sterigmatocystin (5/6, A. varians only produces the versicolorin precursors), violaceols (1/6), viridicatins and cyclopenols (1/6) (Chenet al.2016a).
SeriesStellatiHoubraken & Frisvad,ser.nov.MycoBank MB833251.
Etymology: Named after the type species of this series,A.stellatus.
Type:Aspergillus stellatus Curzi, Atti Reale Accad. Naz. Lincei, Ren-diconti Cl. Sci. Fis. 19: 428. 1934.
Diagnosis:Phylogeny: SeriesStellatibelongs to sect.Nidulantes, sub-gen.Nidulantesand is phylogenetically sister of a clade including series Multicolores,NidulantesandUnguium.Morphology & physiology: Col-onies spreading, conidia en masse green, conidiophores biseriate, stipes hyaline to yellowish brown, smooth; Hülle cells present, globose;
no growth at 40 °C.Sexual morphpresent (except inA.caespitosus), emericella-type, homothallic; ascospores globose, stellate or appen-daged. Series description based onChenet al.(2016a).
Included species:Aspergillus angustatus,A.astellatus,A.caespitosus, A.dromiae,A.filifer,A.miraensis,A.olivicola,A.qinqixianii,A. stella-maris,A.stellatus,A.stelliformis*,A.undulatus,A.venezuelensis[* not included inFig. 16; details on classification, seeSklenaret al.(2020)].
Extrolites(number of species producing compound / total species in series): Aflatoxin B1 (4/12), asperlicine (1/12), asperthecin (6/12),
0.1
A. ivoriensis A. raperi A. bisporus A. rambellii A. ochraceoroseus
A. funiculosus
71/-A. panamensis A. anthodesmis A. conjunctus
72/*
A. amazonicus A. implicatus
87/-A. sparsus A. haitiensis A. diversus
A. biplanus H. avellanea
ser. Raperorum ser. Bispori ser. Ochraceorosei ser. Funiculosi
ser. Conjuncti ser. Implicati ser. Sparsi ser. Biplani
sect. Bispori sect. Raperorum
sect.
Ochraceorosei
sect. Sparsi
C
Fig. 16. (Continued).
asperugines (3/12), astellolide (1/12), asteltoxin (4/12), austinol (1/12), curvularins (1/12), desertorins (6/12), emericellins (8/12), emerin / epurpurin (1/12),fischerin (1/12), gregatins (1/12), 2-ω-hydroxyemodin (8/12), Mer-NF-8054X (1/12), secalonic acid D 781/12), shamix-anthones (10/12), sterigmatocystin (5/12), terrein (3/12), varitriol (4/12), verruculogens and fumitremorgins (1/12), violaceols (1/12) (Chenet al.
2016a).
Series Unguium Houbraken & Frisvad, ser. nov. MycoBank MB833253.
Etymology: Named after the type species of this series,A.unguis.
Type: Aspergillus unguis (Emile-Weill & L. Gaudin) Thom & Raper, Mycologia 31: 667. 1939.
Diagnosis: Phylogeny: Series Unguium belongs to sect. Nidulantes, subgen.Nidulantesand is phylogenetically sister of seriesMulticolores andNidulantes.Morphology & physiology: Colonies growing restrictedly or moderately rapid; conidiaen massein shades of green (yellow-green, olive-green); conidiophores biseriate; stipes hyaline to yellowish brown, smooth; Hülle cells absent (A. israelensis, A. unguis) or present (A.
croceus), globose, subglobose or pyriform; growth at 37 °C absent (A.
croceus,A.israelensis) or restricted (A.unguis), someA.unguisstrains grow restrictedly at 40 °C). Sexual morphunknown,A.unguisNRRL 2393 was reported to tardily produce ascospores. Series description based on Fennell & Raper (1955), Chenet al.(2016a), Hubkaet al.
(2016a).
Included species:Aspergillus croceiaffinis*,A.croceus,A.israelensis, A.longistipitatus*,A.unguis[* not included inFig. 16; details on clas-sification, seeSklenaret al.(2020)].
Extrolites:Aspergillus unguisproduces asperunguisones (and 3-ethyl-5,7-dihydroxy-3,6-dimethyl phthalide), penicillin G, unguinols (= nidu-lins), aspergillusidones, aspergicides, agonodepsides, emeguisins, folipastatins, haiderin, nasrin, rubidin, shirin, yasimin (= unguinol = tridechloronidulin), unguisins, unguispyrones, ustilaginoi-din C and violaceols / orcinols (Chenet al.2016a, Phainuphonget al.
2017a, Morshedet al.2018, Phainuphonget al.2018a), whileA. cro-ceus produces desertorins / kotanins and sterigmatocystin and A.
israelensisproduce emindols (Chenet al.2016a), showing there is no extrolites in common between the three species in seriesUnguium.
Series Versicolores Houbraken & Frisvad, ser. nov. MycoBank MB833254.
Etymology: Named after the type species of this series,A.versicolor.
Type:Aspergillus versicolor(Vuill.) Tirab., Ann. Bot. (Roma) 7: 9. 1908.
Diagnosis:Phylogeny: SeriesVersicoloresbelongs to sect.Nidulantes, subgen.Nidulantes; the phylogenetic relationship with other series in the section is unresolved.Morphology & physiology: Colonies restricted or moderately fast, conidiaen massein shades or green or brown; co-nidiophores biseriate, stipes hyaline or pale brown, smooth or with tu-bercles; Hülle cells absent or present, hyaline, globose, subglobose, ellipsoidal or pyriform; no or poor growth at 37 °C.Sexual morph un-known. Series description based onJurjevicet al.(2012)andChenet al.
(2016a).
Included species:Aspergillus amoenus,A.austroafricanus,A.creber,A.
cvjetkovicii, A. fructus, A. griseoaurantiacus, A. hongkongensis, A.
jensenii,A.pepii,A.protuberus,A.puulaauensis,A.subversicolor,A.
sydowii,A.tabacinus,A.tennesseensis,A.venenatus,A.versicolor.
Extrolites: Most species produce sterigmatocystin (14/16) and pre-cursors such as versicolorins, averufin, averufanin, and norsolorinic acid ((Jurjevicet al. 2013, Chenet al.2016a, Jaksic Despot et al.2017).
Other extrolites from seriesVersicoloresinclude aniduquinolones (2/14), brevianamides F, J, K, Q, R, T, U (1/14), calbistrins (3/14), deoxy-brevianamides (2/14), isocoumarins (2/14), cyclopenols / viridicatols (2/
14), emericellin / arugosins (1/14), shamixanthones (4/14), insulicolides (2/14), mangrovamides (1/14), psychrophilin E-H (1/14), sydowic acids (5/14), sydowinins (1/14), versicolamides (and notoamides, stephaci-dins) (7/14), versiols (2/14), violaceols (13/14), WIN64745 (1/14) (Chen et al.2016a).
Notes on sect. Nidulantes and included series: Section Nidulantes species share the production of more or less brown-pigmented, smooth conidiophores (occasionally with surface protuberances) with globose, subglobose or subclavate vesicles. The conidia are generally globose and echinulate and areen massegreen coloured.Chenet al.(2016a) recognised seven clades in sect.Nidulantes and these were named theA.aurantiobrunneus-,A.multicolor-,A.nidulans-,A.spelunceus-, A. stellatus-,A. unguis- andA. versicolor-clade. These seven clades are here treated as series. These series were primarily based on multigene phylogenetic analysis and certain shared characters, such as growth rates at 25, 37 and 40 °C and ascospore ornamentation (if produced).
SectionOchraceoroseiFrisvad & Samson, Syst. Appl. Microbiol. 28:
451. 2005. MycoBank MB500165.
Type:Aspergillus ochraceoroseusBartoli & Maggi, Trans. Brit. Mycol.
Soc. 71: 393. 1978.
Description: See Frisvad et al. (2005) (morphology),Kocsube et al.
(2016)(phylogeny).
Series Funiculosi Houbraken & Frisvad, ser. nov. MycoBank MB833255.
Etymology: Named after the type species of this series,A.funiculosus.
Type:Aspergillus funiculosusG. Sm., Trans. Brit. Mycol. Soc. 39: 111.
1956.
Diagnosis: Phylogeny: SeriesFuniculosibelongs to sect. Ochraceor-osei, subgen. Nidulantes and is phylogenetically sister ser. Ochra-ceorosei (Fig. 16). Morphology & physiology: Colonies growing moderately; conidia en masse yellow-green, olive-brown to deep brownish purple; conidiophores uniseriate, stipes hyaline or faintly col-oured just below the vesicle, smooth; Hülle cells absent.Sexual morph unknown. Series description based on Smith (1956) and Raper &
Fennell (1965).
Included species:Aspergillus funiculosus.
Extrolites:Aspergillus funiculosusproduces kojic acid (Siddhardhaet al.
2010) and funicin = ethericin B (Königet al.1978, Hamasakiet al.1980, Königet al.1980, Nakamuraet al.1984) and is chemically different from A.ochraceoroseusandA.rambelliiin seriesOchraceorosei. Chemically these diphenylether antibiotic extrolites are shared with Aspergillus sydowiiand other species in seriesVersicolores(Liet al.2015a) and kojic acid is shared with some species in sectionNidulantes(Frisvad &
Samson 2004a, Frisvad et al. 2005, Chen et al. 2016a) and Flavi (Frisvadet al.2019).
Series Ochraceorosei Houbraken & Frisvad, ser. nov. MycoBank MB833256.
Etymology: Named after the type species of this series, A.
ochraceoroseus.
Type:Aspergillus ochraceoroseusBartoli & Maggi, Trans. Brit. Mycol.
Soc. 71: 393. 1979 [1978].
Diagnosis: Phylogeny: SeriesOchraceorosei belongs to sect. Ochra-ceorosei, subgen. Nidulantesand is phylogenetically sister ser. Funi-culosi.Morphology & physiology: Colonies growing moderately or fast on MEA, conidiaen massein shades of yellow; conidiophores biseriate, stipes hyaline, smooth, long; Hülle cells absent; no growth at 37 °C.
Sexual morphunknown. Series description based onBartoli & Maggi (1978)andFrisvadet al.(2005).
Included species:Aspergillus ochraceoroseus,A.rambellii.
Extrolites: Aflatoxin B1, B2, sterigmatocystin, 3-O-methylster-igmatocystin (Frisvadet al.2005). Many other species produce aflatoxin and the related sterigmatocystins, but species in ser.Ochraceoroseido not produce kojic acid like species in sect.Flavi(Frisvadet al.2019), but produce unique secondary metabolites, not as yet structure elucidated, that only occur in this section. No extrolites in common with the other species in sect.Nidulantes(Chenet al.2016a).
Notes on series in sect.Ochraceorosei: The taxonomic position ofA.
funiculosus was discussed several times in the past (Smith 1956, Raper & Fennell 1965, Peterson 2008, Peterson et al. 2008, Hubka et al. 2016a). Smith (1956) could not satisfactorily classify this species and mentioned a possible relationship withA.versicolor, A.flavusandA.glaucus, andRaper & Fennell (1965)accepted this species in sect.Sparsi(asA.sparsusgroup). Using sequence data, A.funiculosus took a basal position in sect.Sparsiin the phyloge-netic analysis ofPetersonet al.(2008), while this species was more similar to A. ochraceoroseus in other analyses (Houbraken &
Samson 2011, Chen et al. 2016a). Based on our nine-gene phy-logeny, A. funiculosus is sister to Aspergillus ochraceoroseus, A.
rambellii of ser. Ochraceorosei (Fig. 2). The sole species in ser.
Funiculosi is uniseriate, in contrast to the biseriate species in ser.
Ochraceorosei.Aspergillus funiculosus does not produce aflatoxins or sterigmatocystins, but kojic acid and funicin (= ethericin B) (König et al.1978, Hamasakiet al.1980, Königet al.1980, Nakamuraet al.
1984, Siddhardhaet al.2010).
SectionRaperorumS.W. Peterson, Varga, Frisvad, Samson ex Hou-braken,sect.nov.MycoBank MB833258.
Synonym: Aspergillussect.Raperi Petersonet al., Aspergillus in the genomic era: 42. 2008;sect.inval.
Etymology: Named after the type species of this section,A.raperi.
Type:Aspergillus raperiStolk & J.A. Meyer, Trans. Brit. Mycol. Soc. 40:
190. 1957.
Diagnosis:Phylogeny: SectionRaperi belongs to subgen. Nidulantes and is phylogenetically sister to a clade containing sectionsNidulantes, Aenei, Usti andCavernicolarum (Fig. 16). Morphology & physiology:
Colonies growing restrictedly, moderately or fast, conidiaen massein shades of yellow or green; conidiophores uniseriate (A. raperi) or biseriate (A.ivorensis), stipes hyaline, smooth (A.raperi) or rough (A.
ivorensis), long; Hülle cells present, globose to subglobose, pyriform or elongate.Sexual morphunknown. Section description based onBartoli
& Maggi (1978)andStolk & Meyer (1957).
Included species:Aspergillus ivoriensis,A.raperi.
Extrolites: No known extrolites have been reported.
Notes: SectionRaperorumis introduced to accommodateA.ivoriensis andA.raperi. These species differ in growth rates and conidiophore structure. However, we tentatively keep both species in one series.
Based on phenotypic characters,Aspergillus raperiwasfirst placed in theA.versicolorgroup (= ser.Versicolores) (Stolk & Meyer 1957) and later in the A. ornatusgroup (Raper & Fennell 1965). This species forms, together with A. ivoriensis, a unique lineage in subgen. Nidu-lantes, and sect. Raperi was informally introduced to accommodate these species (Petersonet al.2008). This section is formally introduced here as sect.Raperorum. SectionRaperorumis not subdivided in series and therefore ser.Raperorumis only informally introduced here.
Section Silvatici S.W. Peterson, Varga, Frisvad, Samson ex Hou-braken,sect.nov.MycoBank MB833259.
Synonym: Aspergillus sect.Silvati Peterson et al., Aspergillus in the genomic era: 44. 2008;sect.inval.
Etymology: Named after the type species of this section,A.silvaticus.
Type:Aspergillus silvaticusFennell & Raper, Mycologia 47: 83. 1955.
Diagnosis:Phylogeny: SectionSilvaticibelongs to subgen.Nidulantes and is phylogenetically sister to a large clade containing sections Nidulantes, Aenei, Usti, Cavernicolarum and Raperorum (Fig. 16).
Morphology & physiology: Colonies growing moderate to fast, conidiaen masse in shades of green; conidiophores biseriate, stipes brownish pigmented, smooth, seldom exceeding 300μm; Hülle cells present, globose to subglobose.Sexual morph unknown. Section description based onRaper & Fennell (1965).
Included species:Aspergillus silvaticus.
Extrolites: Naphthalic anhydride, phthalides (silvaticol, O-methyl-silvaticol, ethyl 3-methylorsellinate, 6-hydroxy-4-methoxy-5-methylphtalimidine, 3,6-dimethyl-4-hydroxy-2-methoxybenzaldehyde, nidulol, quadrilineatin), shamixanthones (arugosin A, B & E, silvatica-mide), silvathione and dithiosilvatin (Hommaet al.1980, Yamazakiet al.
1981a, Fujitaet al.1985, Maebayashi & Yamazaki 1985, Kawahara et al.1986, 1987, Kawahara et al.1988). Naphthalic anhydride and the silvatins are unique to this section.
Notes: Section Silvatici (as Silvati) was informally introduced to accommodateA. silvaticus(Peterson et al.2008) and this section is formally introduced here. Only one species is accommodated in sect.
Silvatici. Introducing ser.Silvaticiwould therefore be superfluous, and the series nameSilvaticiis therefore only informally introduced.
SectionSparsiW. Gams, M. Chr., Onions, Pitt & Samson, Adv. Pen.
Asp. Syst.: 61. 1986 [1985]. MycoBank MB832514.
Type:Aspergillus sparsusRaper & Thom, Mycologia 36: 572. 1944.
Description: See Gams et al. (1985) (morphology), Kocsube et al.
(2016)(phylogeny).
SeriesBiplaniHoubraken & Frisvad,ser.nov.MycoBank MB833260.
Etymology: Named after the type species of this series,A.biplanus.
Type: Aspergillus biplanus Raper & Fennell, Gen. Aspergillus: 434.
1965.
Diagnosis:Phylogeny: SeriesBiplanibelongs to sect.Sparsi, subgen.
Nidulantes and is phylogenetically sister to ser. Sparsi (Fig. 16).
Morphology & physiology: Colonies growing restrictedly or moderately rapid, conidiaen massein shades of green (dark green, blueish green);
conidiophores biseriate, stipes brown, smooth, long; fragmentary conidial structures of varying size and without definite pattern borne near the agar surface; Hülle cells absent.Sexual morph unknown. Series description based onRaper & Fennell (1965).
Included species:Aspergillus biplanus,A.diversus.
Extrolites: Auroglaucin is shared with sectionsAspergillusandRestricti, also withA.conjuctusin ser.Conjuncti(Vargaet al.2010b).
Series Conjuncti Houbraken & Frisvad, ser. nov. MycoBank MB833261.
Etymology: Named after the type species of this series,A.conjunctus.
Type:Aspergillus conjunctusKwon-Chung & Fennell, Gen. Aspergillus:
552. 1965.
Diagnosis:Phylogeny: SeriesConjunctibelongs to sect.Sparsi, subgen.
Nidulantes and is phylogenetically sister to ser. Implicati (Fig. 16).
Morphology & physiology: Colonies grow restrictedly or moderately rapid; conidiaen massein shades of brown (brown, red-brown, olive brown); conidiophores biseriate, stipes hyaline or (pale) brown pig-mented, smooth; reduced conidial structures produced near colony surface absent; Hülle cells absent (A.amazonicus,A.anthodesmis) or present (A. conjunctus, A. panamensis), elongate and/or variously curved or twisted.Sexual morphunknown. Series description based on Raper & Fennell (1965), Bartoli & Maggi (1978).
Included species: Aspergillus amazonicus, A. anthodesmis, A. con-junctus,A.panamensis.
Extrolites (number of species producing compound / total number of species in series): An aszonalenin (1/4), auroglaucin (1/4), gregatins (2/
4), and siderin (3/4) have been found in species in ser.Conjuncti.
SeriesImplicatiHoubraken & Frisvad,ser.nov.MycoBank MB833262.
Etymology: Named after the type species of this series,A.implicatus.
Type:Aspergillus implicatus Persiani & Maggi, Mycol. Res. 98: 871.
1994.
Diagnosis:Phylogeny: SeriesImplicatibelongs to sect.Sparsi, subgen.
Nidulantes and is phylogenetically sister to ser. Conjuncti (Fig. 16).
Morphology & physiology: Colonies growing moderately, conidia en masse white or pale yellow; conidiophores biseriate, stipes hyaline, smooth, long, surrounded by parallel sterile hyphae, slightly echinulate, originating from the base growing up to the vesicle and later then branching to build the hyphal tangle; reduced, penicillate conidiophores present; Hülle cells absent.Sexual morphunknown. Series description based onMaggi & Persiani (1994).
Included species:Aspergillus implicatus.
Extrolites: Only a versicolorin has been detected in A. implicatus (Vargaet al.2010b), indicating that this species may produce either sterigmatocystin and aflatoxins or austocystins, as versicolorins are precursors for both biosynthetic families; however, this has not been confirmed. Since species in sect.Sparsiare not known for producing aflatoxins or austocystins, and sinceA.implicatuswas found in the same habitat asA.ochraceoroseus, that actually produce aflatoxins.
SeriesSparsiHoubraken & Frisvad,ser.nov.MycoBank MB833263.
Etymology: Named after the type species of this series,A.sparsus.
Type:Aspergillus sparsusRaper & Thom, Mycologia 36: 572. 1944.
Diagnosis:Phylogeny: SeriesSparsibelongs to sect.Sparsi, subgenus Nidulantes and is phylogenetically sister to ser. Biplani (Fig. 16).
Morphology & physiology: Colonies spreading, conidiaen masse red-dish brown; conidiophores biseriate, stipes dark brown pigmented, smooth (A.haitiensis) or rough-walled (A. sparsus), long; small, mal-formed, proliferating conidial structures produced near colony surface;
Hülle cells absent.Sexual morphunknown. Series description based on Raper & Fennell (1965)andVargaet al.(2010b).
Included species:Aspergillus haitiensis,A.sparsus.
Extrolites: Some extrolites of unknown structure are common for the two species in ser.Sparsi. The only known extrolites detected in this series are gregatins and siderin (found inA.haitiensisNRRL 4569), also found in species in ser.Conjuncti(Vargaet al.2010b).
Notes on sect. Sparsi and included series: Fennell & Raper (1955) created theAspergillus sparsus group to accommodate four species that possess certain characters, such as pigmented conidiophore stipes and production of smaller conidium forming structures (besides typical aspergillus-like conidiophores). Gams et al. (1985) formally introduced sect.Sparsifor this group of species.Petersonet al.(2008) included seven species in this section (A.anthodesmis,A.biplanus,A.
conjunctus, A. diversus, A. funiculosus, A. panamensis,A.sparsus).
Later, the taxonomy of this section was studied using a polyphasic approach and the section was expanded with A. amazonicus, A.
haitiensis, A. implicatus (Varga et al. 2010b). All species except A.
funiculosus(sect.Ochraceorosei, ser.Funiculosi) are accepted in this section in the current study (Fig. 16). The species in ser. Implicati produce white or pale-yellow conidia; a unique feature in sect.Sparsi.
Hülle cells are present in some species of ser.Conjunctiand those are predominantly elongate. These resemble the Hülle cells produced by sect. Usti members, rather than the (sub)globose or pyriform Hülle cells in sections Aenei, Cavernicolarum, Nidulantes, Raperorum and Silvatici. Species in series Biplani, Implicati and Sparsi produce reduced, sometimes penicillate, conidiophores.
SectionUstiW. Gamset al., Adv. Pen. Asp. Syst.: 58. 1986 [1985].
MycoBank MB832504.
Type:Aspergillus ustus(Bainier) Thom & Church, Aspergilli: 152. 1926.
Description: SeeGams et al.(1985)(morphology),Houbraken et al.
(2007), Samson et al. (2011b) (morphology, phylogeny), Kocsube et al.(2016)(phylogeny).
Series Calidousti Houbraken & Frisvad, ser. nov. MycoBank MB833264.
Etymology: Named after the type species of this series,A.calidoustus.
Type:Aspergillus calidoustusVarga, Houbraken & Samson, Eukaryot.
Cell 7: 636. 2008.
Diagnosis:Phylogeny: SeriesCalidoustibelongs to sect.Usti, subgen.
Nidulantes and is phylogenetically sister of ser. Usti (Fig. 16).
Morphology & physiology: Colonies growing moderately, conidia en massegreyish yellow, brownish grey, greyish brown or greyish green;
conidiophores biseriate, stipes hyaline, (yellow-)brown, smooth; Hülle cells absent (A. germanicus, A. keveioides, A. pseudodeflectus) or present (A.asper,A.calidoustus,A.carlsbadensis,A.contaminans,A.
fuscicans, A. keveii, A. thesauricus), generally irregularly elongate, ovoid, curved to coiled, sometimes globose (A.carlsbadensis).Sexual morphunknown. Series description based onRaper & Fennell (1965), Samsonet al.(2011b), Novakovaet al.(2012), Visagieet al.(2014a), Jurjevic & Peterson (2016).
Included species:Aspergillus asper,A.calidoustus,A.carlsbadensis,A.
contaminans,A. fuscicans,A.germanicus,A.insuetus*,A.keveii,A.
keveioides, A. pseudodeflectus, A. sigurros, A. thesauricus (* not included inFig. 16).
Extrolites: Of the five species examined for extrolites, four produce drimans, two produce asperugins, two produce ophiobolins (G & H), one produces brevianamide A, one produces an arugosin, one produces nidulol and one produces TMC120-B (Samsonet al.2011b, Kozlovsky et al.2017).
SeriesDeflectiHoubraken & Frisvad,ser.nov.MycoBank MB833266.
Etymology: Named after the type species of this series,A.deflectus.
Type:Aspergillus deflectusFennell & Raper, Mycologia 47: 83. 1955.
Diagnosis: Phylogeny: SeriesDeflecti belongs to sect.Usti, subgen.
Nidulantesand is phylogenetically sister to a clade containing series Caldidousti,MonodiorumandUsti(Fig. 16).Morphology & physiology:
Nidulantesand is phylogenetically sister to a clade containing series Caldidousti,MonodiorumandUsti(Fig. 16).Morphology & physiology: