Fig. 14. Combined phylogeny forBenA,CaMandRPB2data sets showing the phylogenetic relation of species and series withinAspergillussubgen.Cremei. 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.
Included species:Aspergillus chrysellus,A.dimorphicus,A.europaeus, A.flaschentraegeri,A.wentii.
Extrolites: All species in ser.Wentiorumproduce asperflavin, emodin, physcion, emodin bianthrone, physcion bianthrone (and other bian-throns), sulochrin and other sulochrins, and wentilacton A and B, except A. flaschentraegeri which only produces asperflavin, physcion and physcion bisanthron. These and many related extrolites have been re-ported fromA.wentiiandA.europaeus(Wellset al.1975, Assanteet al.
1979, Assanteet al.1980, Dorneret al.1980, Xuet al.2015, Hubka et al.2016b, Duet al. 2018, Liet al.2018b, Formet al.2019). The metabolites have not been detected yet in any species in the other series in subgen.Cremei.
Notes on sect.Cremei and included series:Raper & Fennell (1965) introduced the A. wentii and theA. cremeus group and later Gams et al.(1985)formally introduced these as sectionsWentiiandCremei.
All species classified byRaper & Fennell (1965)in sect.Cremei(A.
cremeus,A. chrysellus,A.flaschentraegeri, A.itaconicus, A. stroma-toides) are still here accepted in this section. Phylogenetic data demonstrated that sect.Wentiiwas superfluous (Peterson 1995, 2008).
Aspergillus wentiiis included in sect.Cremeiand the other members of theA.wentiigroup (fideRaper & Fennell 1965) belong to sect.Flavi:A.
terricola(=A.tamarii),A.terricolavar.americana(=A.parasiticus),A.
terricolavar.indicus(=A.tamarii),A.thomii(=A.flavus) (Frisvadet al.
2019). Mainly based on molecular data, sect.Cremeiexpanded to 17 species, which are classified in six series (Arxiorum,Brunneouniseriati, Cremei,Inflati,Pulvini,Wentiorum). SeriesInflatiincludes three species and two of those produce penicillium-like conidiophores. Aspergillus inflatuswas originally described inPenicillium. Another species of this series, A.tardus, was assigned to theA.versicolorgroup (Bissett &
Widden 1984), but the original description mentioned the non-synchronously production of metulae, a character frequently observed inPenicilliumand not common inAspergillus. SeriesPulviniincludes a species that produces blue-green coloured conidia, a unique feature in sect.Cremei. SeriesCremeiandWentiorumare phylogenetically and phenotypically related. The conidia in ser.Cremei are more often in shades of green, while those of ser.Wentiorumare more often in shades of yellow-brown. Both series contain species that reproduce sexually.
The sole taxon classified in seriesArxiorumalso reproduces sexually, but an asexual morph is not described. Series Brunneouniseriati in-cludes two species that both have uniseriate conidial heads. Uniseriate heads are also produced by two species in other series:A.fl aschen-traegeri(ser.Wentiorum) andA.itaconicus(seriesCremei). Regarding extrolites the six series in subgenus and sectionCremeiare remarkably different.
Aspergillus subgen. Fumigati W. Gams, M. Chr., Onions, Pitt &
Samson, Adv. Pen. Asp. Syst.: 56. 1986 [1985]. MycoBank MB832495.
Type:Aspergillus fumigatusFresen., Beitr. Mykol. 3: 81. 1863.
Description: See Gams et al. (1985) (morphology); Samson et al.
(2007a) (morphology, phylogeny); Peterson et al. (2008), Kocsube et al.(2016),Fig. 1, this study (phylogeny).
SectionClavatiW. Gamset al., Adv. Pen. Asp. Syst.: 57. 1986 [1985].
MycoBank MB832500.
Type:Aspergillus clavatusDesm., Ann. Sci. Nat., Bot., ser. 2, 2: 71.
1834.
Description: Phylogeny: Series Clavati belongs to subgen. Fumigati.
Morphology & physiology:Coloniesgrowing restrictedly or moderately rapid; conidial colour en masseblue-green;conidiophores uniseriate, often with clavate vesicles; restricted growth at 37 °C, except A.
acanthosporus(no growth).Sexual morphnot observed in culture, or present, neosartorya-type, homothallic (A.acanthosporus) or hetero-thallic (A.clavatus). Section description based onVargaet al.(2007a) andOjeda-Lopezet al.(2018).
Included species: Aspergillus acanthosporus, A. clavatonanicus, A.
clavatus,A.giganteus,A.longivesica,A.posadasensis,A.rhizopodus, A.seifertii* [* not included inFig. 15; for more details, seeVisagie &
Houbraken (2020)].
Extrolites: At least six species (no data forA. posadasensisand A.
seifertii) in the section produce ribotoxins (Varga & Samson 2008) and tryptoquivalines / tryptoquivalones (Vargaet al.2007a). Kotaninins are produced by four of six species (A.acanthosporus,A.clavatonanicus,A.
clavatus,A.rhizopodus), patulin by three (A.clavatus,A.giganteus,A.
longivesica) and antafumicins by four (A.clavatonanicus,A.clavatus,A.
giganteus,A.longivesica). Pyripyropens are produced by two species (A. longivesicaandA. giganteus). Ribotoxins are shared with several species in sect.Fumigati, includingA.fischeri,A.fumigatus,A. neo-glaberandA viridinutans(Martı́nez-Ruizet al.1999, Olombradaet al.
2014), while the isolate ofA.restrictusproducing restrictocin and reg-ulin was anA.fumigatus(NRRL 3050 = ATCC 34475). Except for a claim that A. oryzae can produce ribotoxins (Machida et al. 2005), producers of these small insecticidal and antifungal proteins (Olombrada et al.2014, Citoreset al.2018) are concentrated in subgen.Fumigati, sect.Clavatiand ser.Fumigati.
Notes: No subdivision of sect.Clavatiis proposed, and ser.Clavatiis only informally introduced here (Table 4).
SectionCerviniW. Gamset al., Adv. Pen. Asp. Syst.: 56. 1986 [1985].
MycoBank MB832497.
Type:Aspergillus cervinusMassee, Bull. Misc. Inform. Kew 1914: 158.
1914.
Description: See Gams et al. (1985) (morphology), Kocsube et al.
(2016)(phylogeny),Chenet al.(2016c)(morphology, phylogeny).
Series Acidohumorum Houbraken & Frisvad, ser. nov. MycoBank MB832596.
Etymology: Named after the type species of the series, Aspergillus acidohumus.
Type: Aspergillus acidohumus A.J. Chenet al., Stud. Mycol. 85: 71.
2016.
Diagnosis:Phylogeny: SeriesAcidohumorumbelongs to sect.Cervini, subgen. Fumigati. Morphology & physiology: Colonies restricted;
conidial colouren massedark fawn; conidiophores uniseriate; no growth at 37 °C.Sexual morphunknown. Series description based onChen et al.(2016c).
Included species:Aspergillus acidohumus.
Extrolites: No extrolites have been found in the species in ser.
Acidohumorum.
SeriesCerviniHoubraken & Frisvad,ser.nov.MycoBank MB832993.
Etymology: Named after the type species of the series, Aspergillus cervinus.
Type:Aspergillus cervinusMassee, Bull. Misc. Inform. Kew 1914: 158.
1914.
Diagnosis:Phylogeny: SeriesCervinibelongs to sect.Cervini, subgen.
Fumigati.Morphology & physiology: Colonies growing moderately fast;
conidial colouren massefawn; conidiophores uniseriate; generally no or very poor growth at 37 °C.Sexual morphunknown. Series description based onChenet al.(2016c).
Included species:Aspergillus cervinus,A. christenseniae, A. kanaga-waensis,A.novoguineensis,A.nutans,A.parvulus,A.subnutans,A.
transcarpaticus,A.wisconsinensis.
Extrolites: Six of nine species produce terremutin, four of nine produce aspervenone, and three of nine produce 4-hydroxymellein (Chenet al.
2016c).
Notes: Aspergillus acidohumus is the sole species in ser. Acid-ohumorum. This series clearly belongs to section Cervini, but is phylogenetically distant from ser. Cervini. Series Acidohumorum is phenotypically distinct from other species by its very slow growth rate.
Section Fumigati W. Gams et al., Adv. Pen. Asp. Syst.: 56. 1986 [1985]. MycoBank MB832496.
Type:Aspergillus fumigatusFresen., Beitr. Mykol. 3: 81. 1863.
Description: See Gams et al. (1985) (morphology); Peterson et al.
(2008), Kocsube et al.(2016), Figs 1 and 2, this study (phylogeny);
Samsonet al.(2007a)(morphology, phylogeny).
Series Brevipedes Houbraken & Frisvad, ser. nov. MycoBank MB832597.
Etymology: Named after the type species of the series, Aspergillus brevipes.
Type:Aspergillus brevipes G. Sm., Trans. Brit. Mycol. Soc. 35: 241.
1952.
Diagnosis: Phylogeny: Series Brevipedes belongs to sect. Fumigati, subgen.Fumigati; the series is well-supported; however, the phyloge-netic relationship of the series with other series of the section remains unresolved (Fig. 15). Morphology & physiology: Colonies growing moderately fast or spreading; conidial colour en masse blue-green;
conidiophores uniseriate; good growth at 37 °C. Sexual morph not observed in culture, or present, neosartorya-type, homothallic, white, yellowish white or pale yellow. Series description based onSamson et al.(2007a)andHubkaet al.(2017).
Included species:Aspergillus brevipes,A.duricaulis,A.quadricinctus, A.tsurutae.
Extrolites: Asperdurin, asperpentyn, cyclopaldic acids, duricaulic acid, fumagillin, meleagrin, pseurotins, roquefortine C, viriditoxin. Asperdurin, asperpentyn, cyclopaldic acids, duricaulic acid and meleagrin has only been found in ser.Brevipedesin sect.Fumigati.
Series Fennelliarum Houbraken & Frisvad, ser. nov. MycoBank MB832598.
Etymology: Named after the type species of the series, Aspergillus fennelliae.
Type: Aspergillus fennelliae Kwon-Chung & S.J. Kim, Mycologia 66:
629. 1974.
Diagnosis:Phylogeny: SeriesFennelliarumbelongs to sect. Fumigati, subgen.Fumigati;Fig. 15shows a close phylogenetic relationship with ser.Unilaterales, though statistical support for this relationship is lacking.
Morphology & physiology: Colonies spreading; conidial colouren masse blue-green; conidiophores uniseriate; good growth at 37 °C. Sexual morphneosarotya-type, homo- or heterothallic, white, yellowish white or pale yellow. Series description based onSamson et al. (2007a)and Hubkaet al.(2017).
Included species:Aspergillus denticulatus,A.fennelliae,A.huiyaniae, A.similanensis,A.sublevisporus.
Extrolites: Antafumicins, asperfuran, aszonalenins, aszonapyrones, chevalones, fumigaclavines, fumigatins, gliotoxin, isocoumarins, pyripyr-openes, reticulol, similanamide, similanpyrones, viridicatumtoxin, vir-iditoxin. Asperfuran, the isocoumarins, reticulol and viridicatumtoxin are only found in this series in sect.Fumigati.
SeriesFumigatiHoubraken & Frisvad,ser.nov.MycoBank MB832994.
Etymology: Named after the type species of the series, Aspergillus fumigatus.
Type:Aspergillus fumigatusFresen., Beitr. Mykol. 3: 81. 1863.
Diagnosis:Phylogeny: SeriesFumigatibelongs to sect.Fumigati, sub-gen. Fumigati, and is phylogenetically most closely related to ser.
Viridinutantes(Fig. 15).Morphology & physiology: Colonies spreading;
conidial colouren masseblue-green; conidiophores uniseriate; growth at 37 and 50 °C. Sexual morphnot observed in culture or present, neosartorya-type, homo- or heterothallic, white or yellowish white. Se-ries description based on Samson et al. (2007a) and Hubka et al.
(2017).
Included species:Aspergillusfischeri,A.fumigatiaffinis,A.fumigatus,A.
fumisynnematus,A.laciniosus,A.lentulus,A.novofumigatus,A. oer-linghausensis,A.spinosus,A.takakii.
Extrolites: Extrolite families found in species of ser.Fumigatiinclude ardeemins, asnovolins, asperfumigatin, asperfumin (asperfumoid), aszonalenins, aszonapyrones, aurantines, avenaciolides, cepha-limycins, chevalones, chloroanthraquinones, chrysogine, cottoqui-nazolines, cycloechinuline, cyclopiazonic acids, cyclotryprostatins, fiscalins,fischerins, fumagillins, fumicyclines, fumigachlorin, fumiga-clavines, fumigatins, fumimycins, fumiquinazolines, fumitremorgins, gliotoxins, helvolic acids, lentulins, neosartoricins, novoamauromins, novobenzomalvins, novofumigatamide, palitantins, pseurotins, pyr-ipyropenes, sartorenol, sartorypyrones, sesterfischeric acids, seto-susin, sphingofungins, takakiamide, terreins, (territrems), trypacidins, tryprostatins, tryptoquivalines, wortmannins, xanthocillins. Even though ser.Fumigatispecies produce the largest diversity of different secondary metabolites, certain secondary metabolites are only found in other series of sect.Fumigati, for example anishidiol, antafumicins, asperdurin, asperfurans, asperpentyns, avenaciolides, cyclopaldic acids, cytochalasins, gancidin, glabramycins, kotanins, meleagrin, monochaetin, mycophenolic acid, phomaligins, reticulol, sartor-yglabrins, sartorymensin, tatenoic acid, viridicatumtoxins, viriditins, viriditoxins, wasabidienones, wortmannins.
Series Neoglabri Houbraken & Frisvad, ser. nov. MycoBank MB832599.
Etymology: Named after the type species of the series,Aspergillus neoglaber.
Type:Aspergillus neoglaberKozak., Mycol. Pap. 161: 56. 1989.
Diagnosis: Phylogeny: Series Neoglabri belongs to sect. Fumigati, subgen.Fumigati;Fig. 15shows a phylogenetic relationship with series FennelliarumandUnilaterales, though statistical support for this rela-tionship is lacking. Morphology & physiology: Colonies spreading;
conidial colouren masseblue-green; conidiophores uniseriate; good growth at 37 °C. Sexual morph not observed in culture or present, neosartorya-type, homothallic, white, yellowish white or pale yellow.
Series description based onSamsonet al.(2007a)andHubkaet al.
(2017).
Included species:Aspergillus auratus,A.australensis,A.brevistipitatus, A. conversis, A. elsenburgensis*, A. galapagensis, A. neoglaber, A.
papuensis,A.shendaweii,A.solicola,A.stramenius(* not included in Fig. 15).
Extrolites: Extrolite families found in ser.Neoglabriinclude antafumicins, asperpentyns, aszonalenins, avenaciolides, chrysogines, clavatols, fellutanines, fumigatin, glabramycins, gregatins, helvolic acid, kotanins, sartoryglabramides, sartoryglabrin, takakiamide, tryptoquivalines, wort-mannins. The asperpentyns, chrysogines, fellutanins, glabramycins, gregatins, sartoryglabramides, sartoryglabrin, takakiamide have only been found in ser.Neoglabri.
Series Spathulati Houbraken & Frisvad, ser. nov. MycoBank MB832600.
Etymology: Named after the type species of the series,Aspergillus spathulatus.
Type:Aspergillus spathulatusTakada & Udagawa, Mycotaxon 24: 396.
1985.
86/*
A. pseudoviridinutans (CBS 458.75, non-type) A. pseudoviridinutans A..multittpiiplillcatus
A
A.quadrddirrciniictus A.tstt ururr tae
ser. Th Thermo om mutati
A.takakik iii
A.oerlrrillnii ghauusenssisii A.fuff migigatus
A.pseudooviridinuutanns(CBS 458.775, non-type) A.pseudoovivvriiirrdiddiiinuutans
se er. Neo eog glabri
A.simii iliial nensisiis A.huiyiyyaniai e A.sublevivviisporrurr s A.dentittculal tuss A.feff nnelllllliiae
A. pseudoviridinutans (CBS 458.75, non-type) A. pseudoviridinutans
Fig. 15. Combined phylogeny forBenA,CaMandRPB2data sets showing the phylogenetic relation of species, series and sections withinAspergillussubgen.Fumigati. 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 avellanea.