*/-P. expansum H. avellanea
ser. Aspergillus
ser. Rubri
ser. Chevalierorum ser. Tamarindosolorum ser. Xerophili ser. Leucocarpi
ser. Halophilici ser. Restricti
ser. Penicillioides ser. Vitricolarum ser. Teporium
sect.
Aspergillus
sect.
Restricti
Fig. 11. Combined phylogeny forBenA,CaMandRPB2data sets showing the phylogenetic relation of species, series and sections withinAspergillussubgen.Aspergillus. 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 canadensis,A.clavatophorus,A.hordei, A.infrequens,A.magnivesiculatus,A.penicillioides,A.reticulatus,A.
salinicola,A.tardicrescens.
Extrolites: Most species produce asperglaucide, while two species produce asperphenamate. One species produces mycophenolic acid and another species produces chrysogine. Three species produce echinulin and two species produce antarone A (Sklenaret al.2017).
SeriesRestrictiHoubraken & Frisvad,ser.nov.MycoBank MB833003.
Etymology: Named after the type species of this series,A.restrictus.
Type:Aspergillus restrictusG. Sm., J. Textile Inst. 22: 115. 1931.
Diagnosis:Phylogeny: SeriesRestrictibelongs to sect.Restricti, sub-gen. Aspergillus and is a sister of series Vitricolarum (Fig. 11).
Morphology & physiology: Colonies restricted on MEA and CYA or growth absent, moderate or spreading on M40Y; conidiophores uni-seriate, with compact or loosely columnar heads, stipe surface (SEM) with hairs; no growth on CY20S at 37 °C (except A. pachycaulis).
Sexual morph unknown. Series description based on Sklenar et al.
(2017).
Included species:Aspergillus caesiellus, A.conicus,A. destruens,A.
domesticus,A.gracilis,A.pachycaulis,A.pseudogracilis,A.restrictus, A.villosus.
Extrolites: All species produce asperphenamate, while two species produce asperglaucide. Four species produce clavatol, one orthosporins and one fulvic acid analog PI-4. Two species can produce mycophenolic acid (Sklenaret al.2017).
Series Vitricolarum Houbraken & Frisvad, ser. nov. MycoBank MB833004.
Etymology: Named after the type species of this series,A.vitricola.
Type:Aspergillus vitricola[as“vitricolae”] Ohtsuki, Bot. Mag. (Tokyo) 75:
436. 1962.
Diagnosis:Phylogeny: SeriesVitricolarum belongs to subgen. Asper-gillus, sect.Restrictiand is a sister of ser.Restricti(Fig. 11).Morphology
& physiology: Colonies restricted on MEA and CYA or growth absent, moderate on M40Y; conidiophores uniseriate, with radiate heads, stipe surface (SEM) smooth; no growth on CY20S at 37 °C.Sexual morph unknown. Series description based onSklenaret al.(2017).
Included species:Aspergillus glabripes,A.vitricola.
Extrolites:Aspergillus glabripesproduces asperphenamate;A.vitricola produces asperglaucide, and one isolate produces an orthosporin (Sklenaret al.2017).
Notes on series in sect.Restricti: SectionRestrictimembers are xero-philic and grow optimally on low water activity substrates (containing high concentrations of sugar or salt).Sklenaret al.(2017)recognised five clades (A.conicus-,A.halophilicus-,A.penicillioides-,A.restrictus-, A.vitricola-clade) in their monographic study on sect.Restricti. In our study, we introduce four series for thesefive clades. TheA. restrictus-andA.conicus-clades are combined in ser.Restrictibecause of their unresolved phylogenetic relationship (Fig. 11,Supplementary Fig. S1).
The shape of conidial heads can be used to distinguish the series in sect.Restricti. Conidial heads of ser.Restrictiform compact or loose columns, those of ser.Vitricolarum are radiate and ser.Penicillioides mainly have globose conidial heads, which might become radiate after prolonged incubation (Sklenaret al.2017). Ascomata are only produced in ser.Halophilici. Furthermore, ser.Restrictispecies (e.g.,A.caesiellus, A. pachycaulis, A. restrictus) are less xerophilic compared to ser.
Penicillioides, which contains the most xerophilic species of the section (e.g.,A.penicillioides).
Aspergillussubgen.CircumdatiW. Gamset al., Adv. Pen. Asp. Syst.:
59. 1986 [1985]. MycoBank MB832507.
Type:Aspergillus alutaceusBerk. & M.A. Curtis, Grevillea 3: 108. 1875 (=Aspergillus ochraceus).
Description: See Gams et al. (1985) (morphology), Houbraken &
Samson (2011), Kocsubeet al.(2016), this study (Fig. 2) (phylogeny).
SectionCandidiW. Gamset al., Adv. Pen. Asp. Syst.: 61. 1986 [1985].
MB832512.
Type:Aspergillus candidusLink, Mag. Ges. Naturf. Freunde Berlin 3: 16.
1809.
Description:Phylogeny: SeriesCandidibelongs to subgen.Circumdati, sect. Candidi and is sister to sect. Petersoniorum (Supplementary Fig. S1, Fig. 12). Morphology & physiology: Colonies restricted or moderate; conidial colour en masse white or yellow; conidiophores biseriate.Sexual morphunknown; sclerotia produced by some species, black or purple-black. Also seePetersonet al.(2008), Kocsubeet al.
(2016) (phylogeny), Varga et al. (2007b), Hubka et al. (2018b) (morphology, phylogeny).
Included species:Aspergillus campestris,A.candidus,A.dobrogensis, A.pragensis,A.subalbidus,A.taichungensis,A.tritici.
Extrolites: Most species in sect. Candidi produce the shikimic acid derived secondary metabolites chloroflavonins, terphenyllins, candidu-sins and xanthoascins (Hubkaet al.2018b), in addition to the terpene-derived taichunins (Katoet al.2018) and the amino acid derived bicyclo [2.2.2]diazaoctane ring containing taichunamides (Kagiyama et al.
2016).
Notes on sect.Candidi: No subdivision of sect.Candidiis proposed, and ser. Candidi is only informally introduced here (see Table 4).
Section Petersoniorum is phylogenetically distant but is the most closely related section (Fig. 2,Jurjevicet al.2015), and some species in this section also produce conidia in white or yellow shades. Section Candidispecies produce predominantly globose vesicles commonly reaching or exceeding a diameter of 20μm, while the vesicles in sect.
Petersoniorumspecies do not exceed 20μm in diam and are variable in shape (pyriform, subglobose, elongate near angular or penicillium-like). Furthermore, sect.Candidispecies can produce black or purple-black sclerotia, while the sclerotia in sect. Petersoniorum are pale yellow to brown (Jurjevicet al.2015).
SectionCircumdatiW. Gamset al., Adv. Pen. Asp. Syst.: 59. 1986 [1985]. MycoBank MB832508.
Type:Aspergillus alutaceusBerk. & M.A. Curtis, Grevillea 3: 108. 1875 (=Aspergillus ochraceus).
Description: SeeGamset al.(1985)(morphology),Visagieet al.(2014c) (morphology, phylogeny),Steenwyket al.(2019)(genome).
Series Circumdati Houbraken & Frisvad, ser. nov. MycoBank MB832987.
Etymology: This series is based on the same type as sect.Circumdati, and therefore also named after this section.
Type:Aspergillus alutaceusBerk. & M.A. Curtis, Grevillea 3: 108. 1875 (=Aspergillus ochraceus).
Diagnosis: Phylogeny: Series Circumdatibelongs to subgen. Circum-dati, sect.Circumdatiand is phylogenetically most closely related to ser.
Sclerotiorum; the node is fully supported in the Bayesian analysis (1.00 pp, data not shown), but bootstrap support is lacking (<70 %, Fig. 12).Morphology & physiology: Colonies spreading; conidial colour en massemostly light yellow to ochre; conidiophores biseriate; generally no growth at 37 °C or restricted (<20 mm, 7 d, CYA), some species grow more rapid (A.pallidofulvus,A.muricatus).Sexual morphgenerally not observed in culture, except in A. muricatus (homothallic, neopetromyces-type); sclerotia production common, white, cream or yellow. Series description based onVisagieet al.(2014c).
ser. Flavi
ser. Kitamyces
ser. Nomiarum ser. Bertholletiarum
ser. Coremiiformes
ser. Alliacei
ser. Leporum ser. Avenacei
ser. Circumdati
ser. Sclerotiorum ser. Steyniorum
sect.
Flavi
sect.
Circumdati A. cerealis
A. austwickii A. aflatoxiformans A. pipericola
A. texensis (= A. minisclerotigenes) A. minisclerotigenes
A. oryzae A. flavus A. sojae A. parasiticus A. novoparasiticus A. arachidicola
A. transmontanensis A. sergii
A. krugeri A. mottae A. subflavus
A. caelatus A. pseudocaelatus
A. pseudotamarii A. tamarii
74/-94/*
A. bertholletiae A. pseudonomiae A. nomiae A. luteovirescens A. togoensis A. coremiiformis
85/*
A. vandermerwei
A. alliaceus (syn. A. albertensis)
89/-A. neoalliaceus
89/0.96
A. lanosus
75/-A. magaliesburgensis A. aspearensis A. leporis A. hancockii
A. avenaceus
A. sesamicola A. melleus
86/*
A. pallidofulvus A. ochraceus A. ostianus
A. westerdijkiae A. cretensis A. affinis
A. muricatus A. westlandensis
A. auricomus
94/*
A. pulvericola A. ochraceopetaliformis
75/0.99
A. insulicola A. pseudoelegans
A. occultus A. steynii A. elegans
91/*
A. persii A. sclerotiorum A. bridgeri A. salwaensis A. subramanianii
72/0.98
A. pseudosclerotiorum
89/*
-/*
-/0.99
A
Fig. 12. Combined phylogeny forBenA,CaMandRPB2data sets showing the phylogenetic relation of species, series and sections withinAspergillussubgen.Circumdati(excl.
sect.Nigri, seeFig. 13). 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.
ser. Sclerotiorum ser. Tannerorum ser. Robusti
ser. Terrei
ser. Nivei ser. Ambigui
ser. Flavipedes
ser. Spelaei ser. Neonivei
ser. Janorum
ser. Candidi
ser. Petersoniorum
sect. Circumdati (cont’d)
sect. Tannerorum sect. Robusti
sect.
Terrei
sect.
Flavipedes
sect.
Janorum
sect.
Petersoniorum sect.
Candidi
0.1
A. pseudosclerotiorum A. fresenii
A. roseoglobulosus A. neobridgeri 81/*
A. tanneri 89/*
A. robustus A. neoafricanus A. terreus A. hortae A. citrinoterreus
91/-A. pseudoterreus A. heldtiae 78/*
A. alabamensis A. floccosus A. aureoterreus
A. niveus A. carneus A. allahabadii A. neoindicus 91/*
A. bicephalus A. iranicus 78/*
A. ambiguus A. microcysticus A. iizukae A. capensis
A. flavipes A. ardalensis A. templicola A. urmiensis 73/*
A. suttoniae 88/*
A. neoflavipes A. micronesiensis
A. movilensis A. luppiae
A. spelaeus A. polyporicola 87/*
A. olivimuriae A. neoniveus
A. trisporus A. brevijanus A. janus
A. yunnanensis A. candidus A. dobrogensis A. campestris 75/*
A. taichungensis A. subalbidus
75/-A. pragensis A. tritici A. arenarioides
A. asclerogenus
73/-A. petersonii
77/-A. peyronelii H. avellanea
-/*
-/*
ser. Olivimuriarum
B
Fig. 12. (Continued).
Included species: Aspergillus affinis, A. auricomus, A. cretensis, A.
melleus,A.muricatus,A.ochraceus,A.ostianus,A.pallidofulvus,A.
sesamicola,A.westerdijkiae,A.westlandensis.
Extrolites: Circumdatins and destruxins are only produced in ser.
Circumdati.
Series Sclerotiorum Houbraken & Frisvad, ser. nov. MycoBank MB832581.
Etymology: Named after the type species of the series, Aspergillus sclerotiorum.
Type:Aspergillus sclerotiorum G. A. Huber, Phytopathology 23: 306.
1933.
Diagnosis: Phylogeny: Series Sclerotiorum belongs to subgen. Cir-cumdati, sect.Circumdatiand is phylogenetically sister to series Cir-cumdati andSteyniorum, though statistical support is lacking (<70 % BS,Fig. 12); The Bayesian analysis posterior probability (pp) positions this series as sister to ser. Circumdati (1.00 pp, data not shown).
Morphology & physiology: Colonies spreading; conidial colouren masse mostly light yellow to ochre; conidiophores biseriate; good growth at 37 °C > 20 mm (7 d, CYA), with exception of A. roseoglobulosus (Visagie et al. 2014c). Sexual morph unknown; sclerotia production common, white, cream or yellow.
Included species: Aspergillus bridgeri, A. fresenii, A. neobridgeri, A.
persii, A. pseudosclerotiorum, A.roseoglobulosus, A. salwaensis, A.
sclerotiorum,A.subramanianii.
Extrolites: This series includes a species,A.persii, producing asper-nidines, cyclopenins, mevinolins (= lovastatins), and sclerotiumins (= aspersclerotiorones) (reported as A. sclerotiorum) (Phainuphong et al. 2016, Bao et al. 2017, Phainuphong et al. 2017b, Phainuphonget al. 2018b, Lebaret al.2019). Cyclopenin, radarins, secalonic acid A, secopenitrem D and sulphinines is only found in species in ser.Sclerotiorum(Visagieet al.2014c). No species in ser.
Sclerotiorumproduce mellein.
Series Steyniorum Houbraken & Frisvad, ser. nov. MycoBank MB832582.
Etymology: Named after the type species of the series, Aspergillus steynii.
Type:Aspergillus steyniiFrisvad & Samson, Stud. Mycol. 50: 39. 2004.
Diagnosis:Phylogeny: SeriesSteyniorumbelongs to subgen. Circum-dati, sect.Circumdatiand is sister to a clade containing ser.Circumdati, but this relationship is lacking statistical support (Fig. 12); Bayesian analysis places this series confidently (1.00 pp) as a sister to series Circumdati and Sclerotiorum (phylogram not shown). Morphology &
physiology: Colonies spreading; conidial colouren massemostly light yellow to ochre; conidiophores biseriate; generally, no growth at 37 °C or sometimes restricted growth (<20 mm, 7 d, CYA). Sexual morph un-known; sclerotia production common, white, cream or yellow. Series description based onVisagieet al.(2014c).
Included species: Aspergillus elegans, A. insulicola, A. occultus, A.
ochraceopetaliformis,A.pseudoelegans,A.pulvericola,A.steynii.
Extrolites: Metabolites only produced by species in ser. Steyniorum include antibiotic Y, asteltoxins, cycloechinulin, insulicolides, N-methyl-epiamauuromine, ochrindols, quinolactacin, and verruculogen TR-2 (Visagieet al.2014c). No species in ser.Steyniorumproduce asper-gamides, mellamides, neohydroxyaspergillic acids or petromurins (Visagieet al.2014c).
Notes on series in sect.Circumdati: The extrolites produced in the three series of sect.Circumdatiare similar and the mycotoxin ochratoxin A is produced in all series of the section. Extrolites include aspergamides (= stephacidins = sclerotiamides) (not yet found in species in ser.
Steyniorum), aspochracins / sclerotiotides, aspyrones, circumdatins,
mellamides, melleins, ochratoxins, orthosporins, penicillic acids, and xanthomegnins (Visagieet al.2014c). The proposed series classifi ca-tion is based on the results of the multigene phylogeny (Fig. 12). Based on a phylogenetic analysis,Visagie et al.(2014c) recognised seven main clades in sect.Circumdati. One clade includedA.robustusand this clade is raised to section level (sect.Robusti; see below) (Jurjevicet al.
2015). TheA.auricomus-,A.muricatus-, and A.ochraceus-cladesfide Visagieet al.(2014c)are treated here as ser.Circumdati. Even though A.auricomusandA.westlandensis(together forming theA. auricomus-clade;Visagieet al.2014c) form a sister clade to the other members of ser.Circumdati(Fig. 12), we did notfind any additional evidence to treat these two species as a separate series. SeriesCircumdati includes species that produce aspochracins, mellamides, circumdatins and aspergamides; this series apparently lost the ability to produce aspo-chracins (Visagieet al.2014c). SeriesSclerotiorumrepresents theA.
fresenii-cladefideVisagieet al.(2014c); ser.Steyniorumincludes theA.
ochraceopetaliformis-cladefideVisagieet al.(2014c),A.steyniiandA.
elegans. The latter two species were treated as a separate clade (Visagie et al. 2014c), and produce TR-2 and cycloechinulin, two extrolites not produced by any other species of sect.Circumdati.
SectionFlaviW. Gamset al., Adv. Pen. Asp. Syst.: 60. 1986 [1985].
MycoBank MB832510.
Type:AspergillusflavusLink, Mag. Ges. Naturf. Freunde Berlin 3: 16.
1809.
Description: See Gams et al. (1985) (morphology), Kocsube et al.
(2016), this study (Fig. 2) (phylogeny), Frisvad et al. (2019) (morphology, phylogeny).
SeriesAlliaceiHoubraken & Frisvad,ser.nov.MycoBank MB832583.
Etymology: Named after the type species of the series, Aspergillus alliaceus.
Type:Aspergillus alliaceusThom & Church, Aspergilli: 163. 1926.
Diagnosis: Phylogeny: Series Alliaceibelongs to subgen.Circumdati, sect.Flaviand is sister to a large clade containing series Bertholletia-rum, Coremiiformes, Flavi, Kitamyces and Nomiarum (Fig. 12).
Morphology & physiology: Colonies spreading; conidial colouren masse yellow; conidiophores biseriate; growth at 37 °C, no or poor growth at 42 °C; reverse on AFPA (Aspergillus Flavus and Parasiticus Agar) cream.Sexual morphgenerally not observed in culture, present inA.
alliaceus, petromyces-type, homothallic; sclerotia often present, black.
Series description based onFrisvadet al.(2019).
Included species: Aspergillus alliaceus, A. lanosus, A. mag-aliesburgensis,A.neoalliaceus,A.vandermerwei.
Extrolites: Certain secondary metabolites, such as altersolanols, asperlicins, burnettienes / phaeospelides, burnettramic acid, griseoful-vins, mevinolins, nalgiovensins / allianthrones, and ochratoxins are only found in ser.Alliaceispecies and not in any other species of sect.Flavi (Goetzet al.1985, Lieschet al.1985, Lieschet al.1988, Mandelare et al.2018, Frisvadet al.2019, Liet al.2019a, Morishitaet al.2019).
Series Avenacei Houbraken & Frisvad, ser. nov. MycoBank MB832584.
Etymology: Named after the type species of the series, Aspergillus avenaceus.
Type:Aspergillus avenaceusG. Sm., Trans. Brit. Mycol. Soc. 26: 24.
1943.
Diagnosis:Phylogeny: SeriesAvenaceibelongs to subgen.Circumdati, sect.Flavi, encompassing the earlier diverging species to all other se-ries of sect. Flavi (Fig. 12). Morphology & physiology: Colonies spreading; conidial colour en masse beige; conidiophores biseriate;
growth at 37 °C, no growth at 42 °C; reverse on AFPA cream.Sexual morphunknown; sclerotia often present, large, black. Series description based onFrisvadet al.(2019).
Included species:Aspergillus avenaceus.
Extrolites:Aspergillus avenaceusis the only species in sect.Flavithat does not produce kojic acid, but isolates in the species can produce avenaciolides, and in common with other species in sect.Flavi alter-solanols, aspirochlorins, pseurotin A and 3-nitropropionic acid (Frisvad et al.2019).
Series Bertholletiarum Houbraken & Frisvad, ser. nov. MycoBank MB832988.
Etymology: Named after the type species of the series, Aspergillus bertholletiae.
Type:Aspergillus bertholletiae[as“bertholletius”] Taniwakiet al., PLoS ONE 7: e42480, 6. 2012.
Diagnosis: Phylogeny: Series Bertholletiarum belongs to subgen.
Circumdati, sect.Flaviand is sister to a clade containing seriesFlavi, Kitamyces and Nomiarum(Fig. 12). Morphology & physiology: Col-onies spreading; conidial colour en masse brown; conidiophores biseriate; good growth at 37 °C, no growth at 42 °C; reverse on AFPA cream. Sexual morph unknown; sclerotia not observed in culture.
Species associated with coconut trees. Series description based on Frisvadet al.(2019).
Included species:Aspergillus bertholletiae.
Extrolites: In common with species from other series in sect.Flavi,A.
bertholletiae produces cyclopiazonic acid, kojic acid, 3-O-methylster-igmatocystin, parasiticolides, tenuazonic acid and ustilaginoidin C (Frisvadet al.2019).
Series Coremiiformes Houbraken & Frisvad, ser. nov. MycoBank MB832585.
Etymology: Named after the type species of the series, Aspergillus coremiiformis.
Type:Aspergillus coremiiformisBartoli & Maggi, Trans. Brit. Mycol. Soc.
71: 386. 1979.
Diagnosis:Phylogeny: Series Coremiiformes belongs to subgen. Cir-cumdati, sect. Flavi and is sister to a clade containing series Ber-tholletiarum,Flavi,KitamycesandNomiarum(Fig. 12).Morphology &
physiology: Colonies spreading; conidial colour en masse orange-brown; conidiophores biseriate; No growth at 37 °C, synnemata pre-sent; reverse on AFPA cream. Sexual morphunknown; sclerotia not observed in culture. Series description based onFrisvadet al.(2019).
Included species:Aspergillus coremiiformis,A.togoensis.
Extrolites: In common with species in other series in sect. Flavi, A.
togoensis produces aflatoxin B1, a bisiderin, paspaline, paspalinine, paxillin and sterigmatocystin, while the other species in the series (A.
coremiiformis) is quite different and only produces unknown indole al-kaloids (Frisvadet al.2019).
SeriesFlaviHoubraken & Frisvad,ser.nov.MycoBank MB832989.
Etymology: Named after the type species of the series, Aspergillus flavus.
Type:AspergillusflavusLink, Mag. Ges. Naturf. Freunde Berlin 3: 16.
1809.
Diagnosis:Phylogeny: SeriesFlavibelongs to subgen.Circumdati, sect.
Flaviand is sister to ser.Kitamyces(Fig. 12).Morphology & physiology:
Colonies spreading; conidial colour en masse mostly yellow-green, occasionally brown (A. oryzae), or brownish green (A. parasiticus);
conidiophores uni- and/or biseriate; good growth at 37 °C, generally growth at 42 °C, exceptA.mottaeandA.subflavus; reverse on AFPA orange, except inA.oryzae.Sexual morphgenerally not observed in culture, except in A. flavus and A. parasiticus, petromyces-type;
heterothallic; sclerotia often present, black. Most species are primarily associated with nuts and oil-seeds, but will also grow on foods from domesticated plants such as cereals (e.g. maize) and dry fruits.
Aspergillus oryzaeis the domesticated form ofA.flavus, andA.sojaeof A.parasiticus; both are used in food fermentations. Series description based onFrisvadet al.(2019).
Included species:Aspergillus aflatoxiformans,A.arachidicola,A. aust-wickii,A.cerealis,A.flavus,A.krugeri,A.minisclerotigenes,A.mottae, A.novoparasiticus,A.oryzae,A.parasiticus,A.pipericola,A.sergii,A.
sojae,A.subflavus,A.transmontanensis.
Extrolites: Most species produces aflatoxins (B and G types) (and precursors such as versicolorins and sterigmatocystins), aflatrems, aflavarins, aflavazols, aflaviniones, asparasones, asperfurans, asper-gillic acids, aspergillomarasmins, aspirochlorins, chrysogines, cit-reoisocoumarins, cyclopiamides, cyclopiazonic acids (and the related speradins), ditryptophenalines, kojic acids, kojistatins, leporins, miya-kamides (= oryzamides), 3-nitropropionic acid, parasitenone, para-siticolides, parasiticols, parasperones, penicillins, sporogens, and ustilaginoidins (Frisvadet al.2019).
Series Kitamyces Houbraken & Frisvad, ser. nov. MycoBank MB832586.
Etymology: Named after G. Kita, who describedA. tamarii, the type species of this series.
Type: Aspergillus tamarii Kita, Centralbl. Bakteriol. 2. Abth. 37: 433.
1913.
Diagnosis:Phylogeny: SeriesKitamycesbelongs to subgen.Circumdati, sect.Flaviand is sister to ser.Flavi(Fig. 12).Morphology & physiology:
Colonies spreading; conidial colouren massein shades of brown; co-nidiophores biseriate; good growth at 37 °C, no growth at 42 °C; reverse on AFPA dark brown.Sexual morphnot observed in culture; sclerotia occasionally present, black. Series description based onFrisvadet al.
(2019).
Included species:Aspergillus caelatus,A.pseudocaelatus,A. pseudo-tamarii,A.tamarii.
SeriesLeporumHoubraken & Frisvad,ser.nov.MycoBank MB832587.
Etymology: Named after the type species of the series,Aspergillus leporis.
Type:Aspergillus leporisStates & M. Chr., Mycologia 58: 738. 1966.
Diagnosis:Phylogeny: SeriesLeporumbelongs to subgen.Circumdati, sect. Flavi and is sister to a large clade containing series Alliacei, Bertholletiarum, Coremiiformes, Flavi, Kitamyces and Nomiarum.
Morphology & physiology: Colonies spreading; conidial colouren masse yellow-green with a shade of beige, beige or olive; conidiophores biseriate; growth at 37 °C, no growth at 42 °C; reverse on AFPA cream.
Sexual morphnot observed in culture; sclerotia often present, large, black.Aspergillus leporisis dung-associated, whileA.aspearensisand A. hancockii have been reported to be soil-borne. It is not known whether the two latter species are actually dung-associated. Series description based onFrisvadet al.(2019).
Included species:Aspergillus aspearensis,A.hancockii,A.leporis.
Extrolites: Aflavarins, aflavinines, antibiotic Y, clavatols, dehy-droterrestric acid, eupenifeldin, fumitremorgins, hancockiamides, 7-hydroxytrichothecolone, kojic acid, leporines, leporizines, mevinolins, onychocins, paspalines, pseurotins, speradins (Frisvadet al.2019). The aflavarins, aflavinines, paspalinines, pseurotins, speradins and kojic acid have been found in other series in sect. Flavi, but antibiotic Y, dehydroterrestric acid, eupenifeldin, fumitremorgins, hancockiamides, 7-hydroxytrichothecolone, leporizines, mevinolins and onychocins have only been found in ser.Leporum.
Series Nomiarum Houbraken & Frisvad, ser. nov. MycoBank MB832588.
Etymology: Named after the type species of the series, Aspergillus nomiae.
Type:Aspergillus nomiaeKurtzmanet al., Antonie van Leeuwenhoek 53: 151. 1987.
Diagnosis:Phylogeny: SeriesNomiarumbelongs to subgen.Circumdati, sect. Flavi and is sister to a large clade containing series Alliacei, Bertholletiarum, Coremiiformes, Flavi, Kitamyces and Nomiarum (Fig. 12).Morphology & physiology: Colonies spreading; conidial colour en masse(dark) yellow-green; conidiophores biseriate; good growth at 37 °C, no growth at 42 °C; reverse on AFPA cream orange.Sexual morph generally not observed in culture, present in A. nomiae, petromyces-type, heterothallic; sclerotia often present, bullet-shaped, black. Species primarily associated to bees. Series description based onFrisvadet al.(2019).
Included species: Aspergillus luteovirescens, A. nomiae, A.
pseudonomiae.
Extrolites: Aflatoxins (B and G type) (and precursors), altersolanols, anominine, aspernomine, aspergillic acids, chrysogines, kojic acid, miyakamides, paspaline, paspalinine, pseurotins, sporogens, and ten-uazonic acid (Frisvadet al.2018, 2019).
Notes on series in sect.Flavi: Using a multigene phylogenetic analysis, Frisvadet al.(2019)recognised eight clades in sect.Flavi. These clades are treated here as separate series. The majority of species belonging to the phylogenetically related series Flavi, Nomiarum and Kitamyces produce aflatoxin B and G. Non-aflatoxin producers areA.oryzae,A.
Notes on series in sect.Flavi: Using a multigene phylogenetic analysis, Frisvadet al.(2019)recognised eight clades in sect.Flavi. These clades are treated here as separate series. The majority of species belonging to the phylogenetically related series Flavi, Nomiarum and Kitamyces produce aflatoxin B and G. Non-aflatoxin producers areA.oryzae,A.