Fig. 26, Penicillium aethiopicum. 7-day old colonies at (A) CYA - CBS

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 26, Penicillium aethiopicum. 7-day old colonies at (A) CYA, (B), MEA, (C) YES, D-H. Conidiophores. I. Conidia. Whitebar = 10 µm.53

J. C. FRISVAD &R.A.SAMSONP. albocoremium (Frisvad) Frisvad, Int. Mod.Tax. Meth. Pen. Asp. Clas.: 275, 2000In Penicillium subgenus Penicillium section Viridicataseries CorymbiferaType: Herb. IMI 285511, isotype Herb. C 60163Culture ex type: IMI 285511 = CBS 472.84 = IBT 21502 =IBT 10682 = FRR 2931 = T12 (T,Y), ex salami, Hillerød,DenmarkDiagnostic features: Roquefortine C, meleagrin, orangebrown reverse on CYA, smooth-walled conidia, wartyconidiophore stipes, halotolerant, psychrotolerant.Similar species: P. albocoremium differs from P. tulipaeand P. radicicola by its dark orange brown reverse on CYAand good sporulation on YES agar. P. hirsutum and P.venetum differ from P. albocoremium by their production ofdeep violet brown exudate.Description:Structure: Asymmetric terverticillate to quarterverticillateConidia: Smooth-walled, globose to subglobose, 3.1-4.5 μmPhialides: 7.9-11.7 μm x 2.2-3.5 μm, with short collulaMetulae: Rough walled, cylindrical, 8.5-18.8 μm x 2.4-4.5μmRami: 9.8-24.9 μm x 2.9-4.8 μmStipes: 150-2000 μm, very rough and wartedSynnemata or fasciculation: Fasciculate to coremiform,white stipesSclerotia: NoneColony texture: Floccose to fasciculateConidium colour on CYA: Greyish green to dull greenExudate droplets on CYA: Copious, clear to pale yellowReverse colour on CYA: Brownish orangeReverse colour on YES: Brownish yellow to orangeDiffusible colour on CYA: NoneEhrlich reaction: +, red violetOdour and volatile metabolites: No dataExtrolites: 1) Roquefortine C & D, meleagrin; 2) Atrovenetin;3) Cyclopeptin, dehydrocyclopeptin, cyclopenin,cyclopenol, viridicatinGrowth on creatine: Weak to moderateAcid and base production on creatine: W/+ (no base)Growth on UNO: Very goodGrowth on nitrite: WeakAbiotic factors:Diam., 1 week, 25°C: CYA: 28-38 mm; MEA: 25-31 mm;YES: 48-60 mm; CREA: 18-28 mm; Cz: 14-20 mm, OAT:23-27 mm: CYAS: 35-41 mm; CzBS: 12-17 mm; CzP: 0mm; UNO: 8-14 mm; DG18: 33-43 mmDiam., 1 week: 15°C: 32–35 mm; 30°C: 8-14 mm; 37°C: 0mmCYA/CYAS: 0.9 [0.8-1.0]: halotolerantCYA15°C/CYA 25°C: 1.0 [0.9-1.1]CYA30°C/CYA 25°C. 0.3 [0.3-0.4]CZBS/CZ: 0.7 [0.5-0.9]CZP/CZ: 0Distribution: Denmark, Slovenia, IsraelEcology and habitats: Roots of Apium graceolens, Petroselinumcrispum, Allium cepa, and Zingiber officinale, onFragaria vespa, cakes, salami and in a saltern.Biotechnological applications: noneBiodeterioration & phytopathology: Possibly causing arot of onions (Allium cepa)Mycotoxicoses and mycotoxins: Roquefortine C has beenlisted as a neurotoxin (Cole and Cox, 1981)Typical cultures: IBT 22806 = CBS 109583, ex saltern,Slovenia; IBT 22521 = CBS 109585, ex Allium cepa,Denmark; IBT 21071 = CBS 109584, ex Zingiber officinaleimported to Denmark; IBT 12813 = CBS 109587 = NRRL941 = Thom 4733.48 (identified by Biourge as P. corymbiferum,identified as P. cyclopium by Raper and Thom,1949); IBT 16884 = CBS 320.97 = CBS 109586, ex Alliumcepa, Denmark; IBT 20068 = CBS 109582, ex cake, Denmark;IBT 19154 = CBS 109614, cake, Denmark; IBT 3457& IBT 10673, ex Fragraria vesca, Denmark; IBT 3891 &IBT 10672 ex Apium graveolens, Denmark; IBT 10679, exfood, Denmark; IBT 10683, IBT 10685 & IBT 10689, exindoor air, Denmark; IBT 10689, ex salami, Denmark; IBT13001 ex Petroselinum crispum, Denmark; IBT 19397 exdate imported to Denmark; IBT 21596, ex Allium cepa,Denmark.54

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 27. Penicillium albocoremium . 7-day old colonies at (A) CYA, (B), MEA, (C) YES, D-H. Conidiophores. I. Conidia.White bar = 10 µm.55

J. C. FRISVAD &R.A.SAMSONP. allii Vincent and Pitt, Mycologia 81: 300, 1989In Penicillium subgenus Penicillium section Viridicataseries CorymbiferaType: Herb. MU Vincent 114Culture ex type: CBS 131.89 = IBT 21503 = IBT 6610 =ATCC 64868 = FRR 3184 (T,Y), ex garlic, EgyptDiagnostic features: Smooth-walled conidia, fulvic acid oranalogues, a dark brown reverse, garlic rotSimilar species: P. allii differs from P hirsutum by notbeing coremiform and by fewer and more lightly colouredexudates droplets. It differs from P. albocoremium, P.radicicola and P. tulipae by its much darker brown reverse.Description:Structure: Terverticillate, some irregular structuresConidia: Smooth-walled, globose 3.0-4.5 μmPhialides: 6.5-11 μm x 2.8-3.2 μmMetulae: 7.5-13 μm x 2.2-3.8 μmRami: 10-25 μm x 3.2-4.5 μmStipes: Short, rough-walled, 75-400 μm x 3.5-5 μmSynnemata or fasciculation: Weakly fasciculate (especiallyon OAT)Sclerotia: NoneColony texture on CYA: Granular to weakly fasciculate,only sulcate in center of colonyConidium colour on CYA: Dull greenExudate droplets: Small clear, yellow or brown on CYAReverse colour on CYA: Dark brownDiffusible colour on CYA: Amber to brown, often producedReverse colour on YES: Yellow brown to dark yellowbrownEhrlich reaction: Pink to redOdour and volatile metabolites: Isobutanol, isopentanol,several terpenes (Larsen and Frisvad, 1995)Extrolites: 1) Roquefortine C, meleagrin, oxaline; 2)Cyclopeptin, dehydrocyclopeptin, cyclopenin, cyclopenol,viridicatin, viridicatol; 3) Atrovenetins; 4) Dehydrofulvicacid, PI-3 and fulvic acid; 5) Chrysogine.Growth on creatine: WeakAcid and base production on creatine: No or weak acid, nobaseGrowth on UNO: GoodGrowth on nitrite: Weak to moderateAbiotic factors:Diam., 1 week, 25°C: CYA: 26-40 mm (35-45 mm*);MEA: 20-37 mm (35-45 mm*); YES: 31-54 mm; CREA:10-24 mm; Cz: 14-24 mm, OAT: 17-36 mm: CYAS: 27-39mm; CzBS: 0-7 mm; CzP: 0 mm; UNO: 13-20 mm; DG18:19-30 mmDiam., 1 week: 5°C: 0 mm; 15°C: 24-30 mm; 30°C: 1-5mm; 37°C: 0 mmCYA/CYAS: 0.8 [0.8-0.9], halotolerantCYA15°C/CYA 25°C: 1.0 [0.9-1.1], psychrotolerantCYA30°C/CYA 25°C: 0.1 [0.04-0.2]CZBS/CZ: 0 (one isolate 0.5)CZP/CZ: 0* According to Vincent and Pitt (1989)Distribution: Egypt, Argentina, Korea, Spain, CzechRepublic, Germany, United Kingdom, Denmark, Georgia,USA, Ontario, Canada.Ecology and habitats: Garlic, occasionally in onions andrice.Biotechnological applications: NoneBiodeterioration & phytopathology: P. allii produces asevere rot in garlic (Vincent and Pitt, 1989; Frisvad andFiltenborg, 1989).Mycotoxicoses and mycotoxins: Only roquefortine C is aknown mycotoxin.Typical cultures: IBT 20212 = CBS 875.95, ex garlic,Spain ; IBT 4112 = CBS 109581 = CCF 1875 = IMI297905, ex Oryza sativa, Czech Republic; IBT 14490 =CBS 109578, Georgia, USA; IBT 13030 = CBS 411.92, exgarlic, Korea; IBT 11735 = CBS 109579, ex walnut surface;IBT 3772 = IBT 3056 = CBS 188.88, ex food, UnitedKingdom; IBT 5553 = CBS 109580 = ATCC 64636 = IMI321505 = NRRL 13630, ex garlic, Denmark; CBS 161.42 =ATCC 8507 = FRR 942 = IMI 039803 = MUCL 29086 =MUCL 29176 = NRRL 942 = IBT 4515 = IBT 5460 (as P.cyclopium); FRR 1668 = IMI 192216, ex soil, Egypt; IMI321506, ex garlic, Denmark, IBT 24593, ex garlic Argentina,CBS 409.92 & 410.92, ex onion, Korea.56

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 28. Penicillium allii. 7-day old colonies at (A) CYA, (B), MEA, (C) YES, D-H. Conidiophores. I. Conidia. White bar = 10µm.57

J. C. FRISVAD &R.A.SAMSONP. atramentosum Thom, Bull. Bur. Anim. Ind.US Dept. Agric. 118: 65, 1910In Penicillium subgenus Penicillium section Viridicataseries CamembertiType: Herb. IMI 039752Culture ex type: CBS 291.48 = IBT 6616 = ATCC 10104= FRR 795 = IFO 8137 = IMI 039752ii = MUCL 29071 =MUCL 29126 = NRRL 795 = QM 7483 (T), ex Frenchcamembert cheese, USA.Diagnostic features: globose smooth-walled conidia, goodgrowth on CREA with no acid production, no growth at30°C, production of oxaline, rugulovasins and ethyl-2-methyl-butanoateSimilar species: P. atramentosum differs from P. chrysogenumby its dark brown reverse and ability to grow onCREA. It differs from members of series Roqueforti by itsmooth conidiophore stipes and from other species by itsalkali tolerance and inability to produce acid on CREA.Description:Conidia: smooth-walled, globose to subglobose, 2.3-3.0 μmPhialides: cylindrical with short collula, 7.5-10 μm x 2.3-2.5 μmMetulae: Cylindrical, 10-15 μm x 2.5-3.0 μmRami: One or two, often somewhat divergent, 15-20 μm x3.0-4.0 μmStipes: Long 300-500 μm, smooth-walled (finely roughenedin CBS 109601), from subsurface hyphaeSynnemata or fasciculation: NoneSclerotia: NoneColony texture on CYA: VelutinousConidium colour on CYA: Dark greenExudate droplets on CYA: Small droplets often produced,pale to dark red brownReverse colour on CYA: Dark brown to rosy brownDiffusible colour: Brownish orangeEhrlich reaction: None, a violet reaction may be observed inparticulary good rugulovasine producersOdour and volatile metabolites: Ethyl-2-methyl-butanoate,ethyl isopentanoate, ethyl acetate, methyl isobutanoate,ethyl isobutanoate, isobutyl acetate (Larsen and Frisvad,1995)Extrolites: 1) Roquefortine C & D, meleagrin, oxaline,neoxaline, 2) Rugulovasine A and BGrowth on creatine: Very goodAcid and base production on creatine: No acid producedGrowth on UNO: Very goodGrowth on nitrite: Good growthAbiotic factors:Diam., 1 week, 25°C: CYA: 23-39 mm (30-45 mm*);MEA: 13-34 mm (20-30 mm*); YES: 29-49 mm; CREA: 9-28 mm; Cz: 14-25 mm, OAT: 10-32 mm: CYAS: 29-32mm; CzBS: 0-15 mm; CzP: 0 mm; UNO: 9-16 mm; DG18:24-33 mmDiam., 1 week: 15°C: 20-25 mm; 30°C: 0 mm; 37°C: 0 mmCYA/CYAS: 1.0 [0.8-1.2]CYA15°C/CYA 25°C: 0.7 [0.6-1.0]CYA30°C/CYA 25°C: 0CZBS/CZ: 0 or 0.6 [0 or 0.4-0.8]CZP/CZ: 0Distribution: Denmark, Norway, Iceland, UK, the Netherlands,Switzerland, Slovenia, Spain, Costa Rica, Ontario,Canada, Tanzania, Australia, New Zealand, Japan; whenfound in the tropics usually found at higher elevations.Ecology and habitats: Camembert, Danbo, Emmenthalerand Gammelost cheeses, blood sausage, walnuts, soil,especially chalky soil, limestone quarries.Biotechnological applications: NoneBiodeterioration & phytopathology: Found on mouldycheeses and sausages.Mycotoxicoses and mycotoxins: Roquefortine C andrugulovasine A and B have been listed as mycotoxins (Coleand Cox, 1981).Typical cultures: IBT 21504 = CBS 194.88 (Y), pig feed,Norway; IBT 21193 = CBS 109588, Mbizi rainforest soil,Ufipa Plateau, 2000 m elev., Tanzania; IBT 11800 = CBS490.84 = IMI 285530 = FRR 2947, Capsicum anuum,Denmark ; IBT 13139 = CBS 109601 = WT 37, ex soil,conifer forest, Switzerland; IBT 10565 = CBS 109611, soil,Urmston, UK; IBT 11801 = CBS 109613, hay, Iceland; IBT14762 = CBS 109612, cheese, Hjørring, Denmark; IBT23031 = CBS 243.73, ex man, Netherlands; IBT 22273, exsoil, Wilson Botanical Garden, Costa Rica, 1100 m elev.;IBT 15294, ex Thymus vulgaris imported to Denmark; IBT4719 = FRR 1446 = IMI 192501, ex soil, Turramurra,N.S.W., Australia; IBT 3417, rye bread, Denmark; IBT3421, ex soil, the Netherlands; IBT 18891, ex air, Cakefactory, Denmark, IBT 3420, ex agricultural soil, IBT23511, New Zealand; IBT 24774, saltern, Slovenia58

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 29. Penicillium atramentosum. 7-day old colonies at (A) CYA, (B), MEA, (C) YES, D-H. Conidiophores. I. Conidia.White bar = 10 µm.59

J. C. FRISVAD &R.A.SAMSONP. aurantiogriseum Dierckx, Ann. Soc. Scient.Brux. 25: 88, 1901In Penicillium subgenus Penicillium section Viridicataseries ViridicataType: Herb. IMI 195050Culture ex type: CBS 324.89 = IBT 14016 = IMI 195050 =ATCC 48920 = NRRL 971 = IBT 13458 = IBT 3502 = FRR971 (T)Diagnostic features: Anacine, aurantiamin, auranthine,penicillic acid, verrucosidin, pseurotin, smooth-walledconidia, dark brown halo on Raulin-Thom agar.Similar species: P. aurantiogriseum can be distinguishedfrom P. viridicatum, P. melanoconidium and P. tricolor byits blue green conidia. It differs from P. freii by it warmorange reverse and poor production of exudates droplets andfrom P. polonicum by its much slower growth rate on CYA.It differs from P. cyclopium by its more blue conidia onCYA and its better sporulation and more white obversemycelium on YES. It differs from P. neoechinulatum by itssmooth walled conidia.Description:Conidiophores terverticillate, few biverticillate and quarterverticillateConidia: 3-4 x 2.5-3.5 μm, globose to subglobose (rarelyellipsoidal), smooth-walled,Phialides: 7.5 -10 μm x 2.5-2.8 μm, cylindrical with collulaMetulae: 10-13 μm x 2.8-3.5 μmRami: 15-25 μm x 3-3.5 μmStipes: 180-400 μm x 3-4 μm, from subsurface hyphae,walls smooth to finely roughenedSynnemata or fasciculation: Weak fasciculationSclerotia: NoneColony texture: Velutinous to weakly fasciculateConidium colour on CYA: Blue green to grey green to darkdull greenExudate droplets on CYA: Occasionally present, pale orlight brownReverse colour on CYA: Curry-yellow to warm orange toreddish brownReverse colour on YES: YellowDiffusible colour: Often present, orange to red brownEhrlich reaction: PinkOdour and volatile metabolites: Isobutanol, isopentanol, 3-heptanone, 3-octanone (Larsen and Frisvad, 1995)Extrolites: 1) Anacine, 2) Aurantiamin, 3) Auranthine, 4)Penicillic acid, 5) Verrucosidin and normethylverrucosidin,6) Pseurotin, 7) Terrestric acid and viridicatic acid, 8)Nephrotoxic glycopeptidesGrowth on creatine: Weak to moderateAcid and base production on creatine: Strong acid production,no base productionGrowth on UNO: WeakGrowth on nitrite: Very weakAbiotic factors:Diam., 1 week, 25°C: CYA: 11-32 mm; MEA: 16-40 mm;YES: 24-46 mm; CREA: 12-29 mm; Cz: 19-27 mm, OAT:15-35 mm: CYAS: 23-38 mm; CzBS: 19-28 mm; CzP: 0mm; UNO: 6-11 mm; DG18: 19-28 mmDiam., 1 week: 15°C: 16-23 mm; 30°C: 12-17 mm; 37°C: 0mmCYA/CYAS: 0.7 [0.6-1.0], halotolerantCYA15°C/CYA 25°C: 0.9 [0.7-1.3]CYA30°C/CYA 25°C: 0.7 [0.6-0.7]CZBS/CZ: 0.9 [0.7-1.0]CZP/CZ: 0RT: Strong reaction, dark brown halo and reverseDistribution: Denmark, Sweden, United Kingdom, Germany,Norway, Czech Republic, former Yugoslavia, Bulgaria,Russia, Iran, New South Wales, Australia, Argentine,USA, CanadaEcology and habitats: Cereals and foods and feeds basedon cereals, les common in onions, garlic, also found in soil.Biotechnological applications: NoneBiodeterioration & phytopathology: This species is amajor agent deteriorating cereals at cool temperatures.Mycotoxicoses and mycotoxins: This species produces amixture of mycotoxins: penicillic acid, verrucosidin andnephrotoxic glycopeptides. The toxicity of the many alkaloidsproduced is unknown. This species and P. polonicummay be involved in Balkan Endemic Nephropathy (Barneset al., 1977, Macgeorge and Mantle, 1990, 1991; Mantle etal., 1991; Mantle and McHugh, 1993, Mantle, 1993; Lundand Frisvad, 1994, Frisvad, 1995). The original isolate (IMI180922A) investigated by Barnes et al. (1977) as P. aurantiogriseumwas correctly identified, while most of the otherstrains subsequently identified as such or as P. commune(Macgeorge and Mantle, 1990) were in fact P. polonicum(Frisvad, 1995).Typical cultures: IBT 11325 = IBT 21505 = CBS 792.95(Y), ex apple juice production plant, Denmark; IBT 11252 =CBS 642.95, ex chicken feed, Denmark; IBT 5134 = CBS112021 = NRRL 3672; IBT 14264 = CBS 110327 = NRRL953; IBT 15986 = CBS 112023, ex mixed pig feed, StaraZagova, Bulgaria; IBT 13169 = CBS 110329 = RMF 7862,ex soil, Wyoming, USA; IBT 12482 = CBS 110330, exwheat, Canada; IBT 24592, garlic, Argentina; IBT 22509,waste, Germany; IBT 11624, IBT 11635; IBT 12954, IBT12480, IBT 11660 & IBT 11672, ex wheat, Canada; IBT21558, Czech Republic; IBT 5268 = CCF 1275; IBT 11293,ex barley, Denmark; IBT 11321 = NRRL 3564; IBT 11309= NRRL 3612; IBT 12834 = NRRL 3747; IBT 12836 =NRRL 6317; IBT 12726 = NRRL 6318; IBT 11301 = IMI092235; IBT 3992 = IMI 180922A; IBT 14454, exLechuiguilla Cave, Carlsbad, New Mexico, USA; IBT12716, ex kangaroo rat, New Mexico, USA; VKM F-232,Russia.60

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 30. Penicillium aurantiogriseum . 7-day old colonies at (A) CYA, (B), MEA, (C) YES, D-H. Conidiophores. I. Conidia.White bar = 10 µm.61

J. C. FRISVAD &R.A.SAMSONP. bialowiezense K. Zaleski, Bull. Int. Acad. Pol.Sci. Lett., Sér. B 1927: 462, 1927In Penicillium subgenus Penicillium section Coronata seriesOlsoniiType: Herb. IMI 092237Culture ex type: CBS 227.28 = IBT 23044 = IMI 092237(T, Y), ex forest soil, PolandDiagnostic features: ellipsoidal finely roughened conidia,long broad stipes with all elements short and appressed,quinolactacin, Raistrick phenols, mycophenolic acid,asperphenamate, poor growth on creatine, but good growthon nitrite-sucrose agarSimilar species: P. bialowiezense is closely related to P.brevicompactum. P. bialowiezense has shorter and less widestipes than P. brevicompactum. P. brevicompactum consistentlyproduces brevianamide A, while P. bialowiezenseproduces quinolactacin A consistently. Both species hasshorter stipes and smaller colonies on CYA and YES thanP.olsonii.Description:Conidiophores: Long, appressed, terverticillateConidia: Finely roughened subglobose to ellipsoidal, 2.5-3.5 μm x 2.0-3 μmPhialides: Cylindrical with gradually tapering collula, 6.5-9μm x 2.5-3 μmMetulae: Cylindrical but apically inflated, 10-15 μm x 3-4.5μmRami: 15-25 μm x 4-5 μmStipes: 250-400 μm x 4-6 μm, smooth-walledSynnemata or fasciculation: NoneSclerotia: NoneColony texture on CYA: VelutinousConidium colour on CYA: Dull green to grey greenExudate droplets on CYA: Often present, pale to reddishbrownReverse colour on CYA: Beige to yellowish creamReverse colour on YES: Cream to cream beigeDiffusible colour on CYA: Pale or light brownEhrlich reaction: Red-violetOdour and volatile metabolites: Not examinedExtrolites: 1) Raistrick phenols, 2) Mycophenolic acid, 3)Asperphenamate, 4) Breviones, 5) Quinolactacin AGrowth on creatine: WeakAcid and base production on creatine: Weak acid productionGrowth on UNO: GoodGrowth on nitrite: GoodAbiotic factors:Diam., 1 week, 25°C: CYA: 11-25 mm; MEA: 9-16 mm;YES: 18-30 mm; CREA: 8-12 mm; Cz: 9-18 mm, OAT: 11-25 mm: CYAS: 21-31 mm; CzBS: 6-16 mm; CzP: 0 mm;UNO: 6-13 mm; DG18: 16-22 mmDiam., CYA, 1 week: 15°C: 17-22 mm; 30°C: 0 mm; 37°C:0 mmCYA/CYAS: 0.8 [0.5-1.3], halotolerantCYA15°C/CYA 25°C: 1.1 [0.8-1.9], psychrotolerantCYA30°C/CYA 25°C: 0CZBS/CZ: 0.8 [0.6-1.1]CZP/CZ: 0Distribution: Denmark, Faroe Islands, Poland, Italy,Slovenia, Chile, Wyoming, USA, Canada, Saudi-ArabiaEcology and habitats: Forest soil, coffee cherries, mouldymushrooms, seaweed, yoghurt, dried lamb meat, Brusselssprouts, sage, margarine, wheat bread, thyme, mouldyharness, air in factories.Biotechnological applications: NoneBiodeterioration & phytopathology: May degrade leatherMycotoxicoses and mycotoxins: May grow in fruit yoghurtsand cause intoxication, but the metabolites responsibleare unknown (Frisvad, unpublished).Typical cultures: IBT 13469, ex wheat, Denmark; IBT20786 = CBS 110104, ex seaweed, Bellevue, Denmark;IBT 21225, ex soil under Salix sp. in dried root segment ofElisabeth Adams irrigation ditch, Centennial, Wyoming,USA; IBT 22460, ex soil under Nothofagus sp., Chile; IBT22443 = CBS 110102, ex bread, Italy; IBT 13145, ex soil,conifer forest, Switzerland; IBT 21578 = CBS 112477, exbarley, Denmark; IBT 6510, ex Thymus vulgaris, Denmark;IBT 6500 = CBS 112478, ex margarine, Denmark.62

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 31. Penicillium bialowiezense. 7-day old colonies at (A) CYA, (B), MEA, (C) YES, D-H. Conidiophores. I. Conidia.White bar = 10 µm.63

J. C. FRISVAD &R.A.SAMSONP. brevicompactum Dierckx, Ann. Soc. Scient.Brux. 25: 88, 1901In Penicillium subgenus Penicillium section Coronata seriesOlsoniiType: Herb. IMI 040225Culture ex type: CBS 257.29 = CBS 110071 = IBT 23045= IMI 040225 = ATCC 9056 = ATCC 10814 = FRR 862 =NRRL 862 = NRRL 863 = NRRL 2011 (T), ex unrecordedsource.Diagnostic features: Brevianamide A, mycophenolic acid,pebrolides, Raistrick phenols, finely roughened ellipsoidalconidia, short broad penicilli, apically inflated metulae.Similar species: See P. bialowiezense.Description:Conidiophores: Long, appressed, terverticillateConidia: Finely roughened ellipsoidal, 2.5-3.5 μm x 2.0-2.5μmPhialides: Cylindrical with gradually tapering collula, 6.5-9μm x 2.5-3 μmMetulae: Cylindrical apically inflated, 10-15 μm x 3.5-4.5μmRami: 15-25 μm x 4-5 μmStipes: 400-800 μm x 4-6 μm, smooth-walledSynnemata or fasciculation: NoneSclerotia: NoneColony texture: VelutinousConidium colour on CYA: Dull green to grey greenExudate droplets on CYA: Pale to yellow or reddish brownReverse colour on CYA: beige to yellowish creamReverse colour on YES: Cream-coloured to beige, in fewstrains a conspicuous red reverse and diffusible pigment isproducedDiffusible colour on CYA: Pale or light brownEhrlich reaction: No reaction or yellowOdour and volatile metabolites: Isobutanol, isopentanol(Larsen and Frisvad, 1995)Extrolites: 1) Raistrick phenols; 2) Mycophenolic acids andmycochromenic acid, 3) Brevianamide A & B, 4) Asperphenamate,5) Pebrolides, 6) Botryodiploidin, 7) 11-(5’-Epoxy-4’-hydroxy-3’-hydroxymethylcyclo-2’-hexenone)-∆-8(12)-drimene , 8) Silvatins, 9) BrevigillideFurther extrolites reported: 10) Brevicompanins, 11) Brevioxims,12) Adenophostins. The identity of the producingorganisms has not been verified for the last three groupsGrowth on creatine: PoorAcid and base production on creatine: Most commonly noacid production, occasionally weak or good acid productionGrowth on UNO: GoodGrowth on nitrite: Moderate to goodAbiotic factors:Diam., 1 week, 25°C: CYA: 14-30 mm [atyp.: 8-11 mm];MEA: 10-26 mm; YES: 20-36 mm; CREA: 4-18 mm; Cz:9-18 mm, OAT: 16-26 mm: CYAS: (15-)21-26 mm; CzBS:7-17 mm; CzP: 0 mm; UNO: 10-14 mm; DG18: 12-19 mmDiam., CYA, 1 week: 15°C: 17-21 mm; 30°C: 0-3 mm;37°C: 0 mmCYA/CYAS: 0.9 [0.8-0.9], halotolerant64CYA15°C/CYA 25°C: 0.9 [0.7-1.0], psychrotolerantCYA30°C/CYA 25°C: 0.1 [0-0.1]CZBS/CZ: 0.9 [0.6-1.0]CZP/CZ: 0Distribution: Cosmopolitan. Denmark, Faroe Islands,Greenland, The Netherlands, Sweden, Svalbard, Norway,United Kingdom, Germany, Poland, France, Slovenia, Italy,Greece, Costa Rica, Venezuela, Columbia, Brazil, Chile,Connecticut, New Mexico, Wisconsin, South Carolina,USA, Canada, Ethiopia, India, Japan, Australia, EasternIsland (Chile). See also Domsch et al. (1980)Ecology and habitats: Soil under conifers, agricultural soil,mouldy mushrooms, spruce cones, acorns, seaweed, sage,coffee cherries, apples, potatoes, barley, wheat, oats, sorghum,mouldy coffee beans, seaweed, air in factories, codroe, mouldy bakers yeast, apricot puree, maple syrup,margarine, liver paté, salami and other processed foods,waste, dead insects, human bone, salt pans.Biotechnological applications: Production of mycophenolicacid (mofetil) (brand name CellCept) (Bentley, 2000).This is used for treatment of many diseases, but is especiallyeffective for prevention of rejection in heart andkidney transplantations (Bentley, 2000)Biodeterioration & phytopathology: P. brevicompactumhas been found growing actively on Amanita citrina, Paxillusinvolutus, Flammulina velutipes, Boletus spp., Pleurotusostreatus, Nectria rishbethii and other macrofungi; it hasalso been found growing in cosmetics and on wood forbuilding constructionMycotoxicoses and mycotoxins: Mycophenolic acid is anantibiotic, with anti-tumor, anti-psoriasis and immunosuppressivefeatures (Bentley, 2000) and may be of relevancefor secondary mycotoxicosis (bacterial infections caused byintake of immunosuppressive mycotoxins). The otherextrolites produced have not been reported to be mycotoxinsin the strict sense of the word, except botryodiploidin. Thismycotoxins is produced by some strains of P. brevicompactum(Fujimoto et al., 1980 misidentified as P. carneolutescens;Frisvad, 1989)Typical cultures: IBT 18329 = CBS 110067, ex soil underJuniper, Madrid, New Mexico, USA; IBT 21507 = IBT6607 = CBS 480.84 = FRR 2938 (Y), ex Raphanus sp.,Denmark; IBT 13151 = CBS 110068 = WSF 3531, ex soil,Wisconsin, USA; IBT 4342 = CBS 110072 = NRRL 867 =IMI 092219 = FRR 3719, ex unrecorded source (P. griseobrunneum);CBS 256.31 = IBT 23046 = IMI 089824 =NRRL 859 = FRR 859 = ATCC 10111 = IFO 5858, exdecomposing mushroom, Storrs, Connecticut, USA (P.stoloniferum); CBS 210.28 = IBT 23043 = IMI 092266, exforest soil under conifers, Puszcza, Bialowieska, Poland (P.patris-meae); CBS 316.59 = IBT 23047 = IMI 092262 =NRRL 866 = FRR 866, ex soil under conifers, Tatry Mountains(P. hagemii); CBS 317.59 = IBT 23069 = IFO 5727 =FRR 1363 = IMI 068217 = ATCC 18311, ex soil, Japan (P.brunneostoloniferum); IBT 18098 = CBS 110069 = FRR2455, ex artificial maple syrup, preserved with 650 ppmbenzoic acid, Sydney, NSW, Australia.

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 32. Penicillium brevicompactum. 7-day old colonies at (A) CYA, (B), MEA, (C) YES, D-H. Conidiophores. I. Conidia.White bar = 10 µm.65

J. C. FRISVAD &R.A.SAMSONP. camemberti Thom, Bull. Bur. Anim. Ind. USDept. Agric. 82: 33, 1906In Penicillium subgenus Penicillium section Viridicataseries CamembertiType: Herb. IMI 027831Culture ex type: CBS 299.48 = IBT 21508 = IBT 21604 =IMI 027831 = IMI 092200 = ATCC 1105 = ATCC 4845 =FRR 878 = LCP P11 = MUCL 29790 = NRRL 877 =NRRL 878 (T,Y)Diagnostic features: Cyclopiazonic acid, white floccosecolonies, poor sporulationSimilar species: Deteriorated strains of P. commune maylook like P. camemberti. P. camemberti differs from P.caseifulvum by its poor sporulation and its inability toproduce orange reverse colours on YES.Description:Conidiophores: Terverticillate to quaterverticillate, sometimesirregular structuresConidia: Smooth-walled, globose to suglobose, 3.5-5 μm x3.3-4.5 μmPhialides: Cylindrical with long wide colulla, 10-13 μm x2.5-3 μmMetulae: Cylindrical 7.5-12 μm x 2.5-3.5 μmRami: Cylindrical, 15-25 μm x 3-4 μmStipes: 200-500 μm x 3-4 μmSynnemata or fasciculation: NoneSclerotia: NoneColony texture: FloccoseConidium colour on CYA: White or rarely grey greenExudate droplets: None or few droplets (often large violetdroplets on CREA)Reverse colour on CYA: Pale, cream-coloured or yellowishReverse colour on YES: Cream yellowDiffusible colour: noneEhrlich reaction: Violet (no reaction in few old isolates)Odour and volatile metabolites: 3-octanone, ethylacetat,isobutanol, ethyl isobutanoate, isobutyl acetate, styrene, 1-octen-3-ol, 3-octanol, ethyl hexanoate, 2-methyl-isoborneol(Larsen and Frisvad, 1995)Extrolites: 1) Cyclopiazonic acids, 2) Cyclopaldic acid(rare), 3) Rugulovasine A & B (rare), 4) Aspereynone-likecompounds, 5) Palitantin (rare)Growth on creatine: Very goodAcid and base production on creatine: Moderate to goodacid production followed by base productionGrowth on UNO: Very goodGrowth on nitrite: Weak or moderateAbiotic factors:Diam., 1 week, 25°C: CYA: 19-27 mm; MEA: 12-27 mm;YES: 23-38 mm; CREA: 9-17 mm; Cz: 13-20 mm, OAT:10-21 mm: CYAS: 14-33 mm; CzBS: 6-17 mm; CzP: 0-1mm; UNO: 13-18 mm; DG18: 14-27 mmDiam., CYA, 1 week: 15°C: 14-26 mm; 30°C: 0-3 mm;37°C: 0 mmCYA/CYAS: 1 [0.8-1.2], halotolerantCYA15°C/CYA 25°C: 1.0 [0.8-1.3], psychrotolerantCYA30°C/CYA 25°C: 0 [0-0.2]CZBS/CZ: 0.8 [0.5-1.3]CZP/CZ: 0 [0-0.1]Distribution: This domesticated species has been found incountries where white mould cheeses are produced andsold.Ecology and habitats: On white mould cheeses (brie,camembert etc.). This species is a domesticated from of P.commune Thom. The species has never been found outsidethe white mould cheese environment. Occasionally crosscontaminatingother cheeses in refrigerators.Biotechnological applications: Production of white mouldcheesesBiodeterioration & phytopathology: -Mycotoxicoses and mycotoxins: Cyclopiazonic acid isproduced by some strainsTypical cultures: IBT 21601 = CBS 123.08 = NRRL 874 =ATCC 10387 = IMI 091932 (P. camemberti var. rogeri), exFrench camembert cheese; IBT 11754 = CBS 303.48 =ATCC 10423 = FRR 875 = IMI 028810 = MUCL 29156 =NRRL 875 = UPSC 3178 (P. caseicola), ex French camembertcheese; IBT 21602 = CBS 112325 = NRRL 876 (P.candidum Roger), IBT 3505 = CBS 112479 (NB P. communelike back-mutation), ex French camembert cheese,”Prairie”, IBT 15441 = CBS 112562, ex brie, Denmark;IBT 11568 = CBS 249.77A, ex German camembert cheese,T65 = IBT 11755 = CBS 190.67, ex Dutch camembertcheese; IBT 14856 = CBS 112078, ex Appenzeller cheese,Switzerland; CBS 160.42; IBT 11570 = CBS 131.67 =MUCL 8446, ex French brie cheese; CBS 133.67, ex Frenchcamembert cheese; CBS 248.77, Germany; IBT 13592 =CBS 273.97, ex dust, Denmark; IBT 23943 = Kulmbach Sp.2491, Germany; IBT 23941 = ATCC 42009 = KulmbachSp. 912, ex camembert cheese.66

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 32. Penicillium camemberti. 7-day old colonies at (A) CYA, (B), MEA, (C) YES, D-H. Conidiophores. I. Conidia. Whitebar = 10 µm.67

J. C. FRISVAD &R.A.SAMSONP. carneum (Frisvad) Frisvad, Microbiology, UK,142: 546, 1996In Penicillium subgenus Penicillium section Roquefortiseries RoquefortiType: Herb. IMI 293204Culture ex type: CBS 112297 = IBT 6884 = IBT 18419 =IMI 293204 = ATCC 58624, ex rye bread, Denmark (T)Diagnostic features: Mycophenolic acid, patulin, roquefortineC, isofumigaclavine A & B, penitrem A, globoselarge smooth-walled conidia, rough-walled stipes, highgrowth rate on all media, growth on propionic acid andacetic acid, growth at low oxygen and large carbon dioxidelevels.Similar species: P. carneum differs from P. roqueforti byits inability to produce the dark green reverse colour onCYA and from P. paneum by its strong smell of isobutanoland geosmin.Description:Conidiophores: Terverticillate, occasionally quarterverticillate,appressed elements, borne from subsurface hyphaeConidia: smooth-walled, globose, 3.5-5 μmPhialides: Cylindrical with short collula, 8-10 μm x 2.5-3.0μmMetulae: Cylindrical, 10-17 μm x 3-4 μmRami: Cylindrical, 17-35 μm x 3-4 μmStipes: Rough-walled often warted, 100-200 μm x 4-5 μmSynnemata or fasciculation: NoneSclerotia: NoneColony texture: VelutinousConidium colour on CYA: Pistacio green to dark AmericangreenExudate droplets on CYA: None or clear dropletsReverse colour on CYA: Beige to brownReverse colour on YES: Cream beige to curryDiffusible colour on CYA: NoneEhrlich reaction: VioletOdour and volatile metabolites: Isopentanol, geosmin, 2-methyl-3-butene-2-ol, isobutanol, 1-octene, isopentylacetate, 1-methoxy-3-methyl-benzene (Larsen & Frisvad,1995)Extrolites: 1) Cyclopaldic acid and chromanols, 2) Mycophenolicacids, 3) Patulin, 4) Penicillic acid (only producedby CBS 449.78); 5) Roquefortine C, 6) Penitrem A; 7)Isofumigaclavine AGrowth on creatine: Very goodAcid and base production on creatine: Acid production inmargin of colonyGrowth on nitrite: GoodAbiotic factors:Diam., 1 week, 25°C: CYA: 36-53 mm; MEA: 35-56 mm;YES: 51-74 mm; CREA: 27-35 mm; Cz: 23-34 mm, OAT:34-72 mm: CYAS: 33-35 mm; CzBS: 17-38 mm; CzP: 23-34 mm; UNO: 28-36 mm; DG18: 39-44 mmDiam., CYA, 1 week: 15°C: 32-36 mm; 30°C: 11-26 mm;37°C: 0 mmCYA/CYAS: 1.2 [1.1-1.2]CYA15°C/CYA 25°C: 0.9 [0.8-0.9]CYA30°C/CYA 25°C: 0.3 [0.3-0.6]CZBS/CZ: 1.0 [0.7-1.1]CZP/CZ: 0.9 [0.7-1.1]High resistance to acid and good growth at high CO2 levels.Distribution: Denmark, Norway, Sweden, Germany, GreatBritain, USA, CanadaEcology and habitats: Dried meat, silage, rye bread, water,beer, (barley), cheddar cheese, mouldy bakers yeast, corkBiotechnological applications: NoneBiodeterioration & phytopathology: This fungus maydeteriorate silage, other lactic acid fermentation products,and beer.Mycotoxicoses and mycotoxins: Patulin, penitrem A,mycophenolic acid, (penicillic acid), isofumigaclavine A areall mycotoxins that can be produced potentially in silageand other acid containing products. P. carneum was involvedin mycotoxicosis of a man drinking beer contaminatedwith P. carneum (IBT 11188) (reidentified by us,originally identified as P. crustosum). The beer containedboth penitrem A and isofumigaclavine A (Cole et al., 1983)Typical cultures: IBT 21509 = IBT 3473 = IBT 6753 =CBS 449.78, ex cheddar cheese, USA (Y); IBT 6892 = CBS468.95, ex salami, Germany; IBT 6885 = IBT 3472 = CBS466.95, ex salami, Germany; IBT 3466 = CBS 467.95, exwater tank, Denmark; IBT 19478 = CBS 390.78, ex rawsausage, Germany; IBT 15600 = CBS 112489, ex chilledfood, France; IBT 6888 = CBS 112487, ex Hordeum vulgare,Denmark; NRRL 1168 = IBT 16402, Ottawa, Canada;NRRL 855 = IBT 16434; IBT 11188, ex beer intoxicatingman, Georgia, USA.68

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 33. Penicillium carneum. 7-day old colonies at (A) CYA, (B), MEA, (C) YES, D-H. Conidiophores. I. Conidia. White bar= 10 µm.69

J. C. FRISVAD &R.A.SAMSONP. caseifulvum Lund, Filt. & Frisvad, J. FoodMycol. 1: 97, 1998In Penicillium subgenus Penicillium section Viridicataseries CamembertiType: Herb. C 24999Culture ex type: CBS 101134 = IBT 21510 = IBT 18282(T,Y), ex Danish blue cheeseDiagnostic features: Cyclopeptin, rugulovasine A, smoothwalledconidia, floccose colonies, good growth on CREA,weak but consistent growth on CzP (1000 ppm propionicacid, pH 3.5)Similar species: P. caseifulvum differs from P. camembertiby an orange reverse on YES agar.Description:Conidiophores: Born from aerial hyphae, terverticillateConidia: Smooth-walled, subglobose to broadly ellipsoidal,3-5 μm x 2.5-3.5 μmPhialides: Cylindrical with short narrow collula, 7-13 μm x2.8-3.5 μmMetulae: Cylindrical, 10-13 μm x 3.5-4.5 μmRami: Cylindrical, 12-25 μm x 4-5 μmStipes: 300-800 μm xSynnemata or fasciculation: NoneSclerotia: NoneColony texture: FloccoseConidium colour on CYA: Greyish green to greyish bluegreen (turquoise)Exudate droplets on CYA: Small clear droplets at colonymarginReverse colour on CYA: Creamish yellow to brown yellowReverse colour on YES: Vividly orange, rarely only creamyellowDiffusible colour on CYA: NoneEhrlich reaction: Violet (weak reaction)Odour and volatile metabolites: Not examinedExtrolites: 1) Cyclopeptin, 2) Rugulovasine A and BGrowth on creatine: Very good growthAcid and base production on creatine: Good acid productionfollowed by base productionGrowth on UNO: Very goodGrowth on nitrite: Very goodAbiotic factors:Diam., 1 week, 25°C: CYA: 15-24 mm; MEA: 16-28 mm;YES: 32-42 mm; CREA: 13-24 mm; Cz: 21-24 mm, OAT:22-25 mm: CYAS: 23-33 mm; CzBS: 15-22 mm; CzP: 3-5mm; UNO: 17-23 mm; DG18: 21-24 mmDiam., CYA, 1 week: 5°C: 3-8 mm; 15°C: 18-23 mm;30°C: 0 mm; 37°C: 0 mmCYA/CYAS: 0.8 [0.7-1.0]CYA15°C/CYA 25°C: 0.9 [0.8-1.1]CYA30°C/CYA 25°C: 0CZBS/CZ: 0.9 [0.7-1.0]CZP/CZ: 0.2 [0.1-0.2]Distribution: Denmark, France, GermanyEcology and habitats: Blue mould cheeses and some otherGerman and French cheesesBiotechnological applications: P. caseifulvum has beenfound to colonize some of the most highly regarded bluemould cheeses (unpublished results). White mould typecheeses have been made on an experimental basis with P.caseifulvum. These cheeses were turquoise on the surfaceand appeared to be of a very fine quality (unpublishedresults). Not producing cyclopiazonic acid, P. caseifulvumcould be a potential new candidate for fermenting cheesesor salami.Biodeterioration & phytopathology: The growth of P.caseifulvum on the surface of blue mould cheeses can beseen as biodeterioration, but also regarded as an indicator ofvery good quality.Mycotoxicoses and mycotoxins: Rugulovasine A is apotential mycotoxin, but no relevant toxicity data (oral, skinor pulmonary toxicity) are available.Typical cultures: IBT 19782 = CBS 108956, ex Danishblue cheese; IBT 18725 = CBS 108957, ex Danish bluecheeswe, Bornhom; IBT 19801 = CBS 111838, ex Frenchcheese (Saint Lactaire); IBT 19802 = CBS 111837, exFrench goat cheese; IBT 23156 = CBS 112324, ex GermanMontagnolo cheese; IBT 23155 = CBS 112323, ex Danishblue cheese; IBT 20915 = CBS 111836, ex Danish bluecheese.70

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 35. Penicillium caseifulvum. 7-day old colonies on A. CYA, B. MEA, C. YES, D-H. Conidiophores. I. Conidia. White bar= 10 µm.71

J. C. FRISVAD &R.A.SAMSONP. cavernicola Frisvad & Samson, sp. nov.In Penicillium subgenus Penicillium section Viridicataseries SolitaType: Herb. CBS 100540Culture ex type: IBT 14499 = CBS 100540, ex wall of theLechuiguilla Cave, Carlsbad, New Mexico, USA (T)Diagnostic features: Territrems, aurantiamine, asteltoxin,dark green rough-walled conidia, yellow brown reverse onCYA, good growth on CREA, no growth at 30°CSimilar species: Differs from P. solitum, P. echinulatumand P. discolor by its yellow brown reverse on CYA.Description:Conidiophores: Terverticillate, appressed elements, bornfrom subsurface hyphaeConidia: Rough-walled, globose to subglobose, 3.5-4.5 μm.Phialides: Cylindrical tapering to a distinct collulum, 8-11μm x 2.2-3.0 μmMetulae: Cylindrical, 10-15 μm x 2.5-3.5 μmRami: Cylindrical, 12-20 μm x 3.2-4.2 μmStipes: Rough-walled, 300-550 μm x 3.5-4.2 μmSynnemata or fasciculation: Weakly fasciculateSclerotia: NoneColony texture: VelutinousConidium colour on CYA: Dark green,Exudate droplets on CYA: Present, clearReverse colour on CYA: Yellow brownReverse colour on YES: Strongly yellowDiffusible colour on CYA: NoneEhrlich reaction: None (yellow in two isolates)Odour and volatile metabolites: No dataExtrolites: 1) Territrems and arisugacins, 2) Asteltoxin, 3)Aurantiamine, 4) Glyanthrypine, 5) Dipodazin, 6) Aspereynone-likecompoundsGrowth on creatine: Very goodAcid and base production on creatine: Weak to moderateacid, delayed base productionGrowth on UNO: Moderate to goodGrowth on nitrite: WeakAbiotic factors:Diam., 1 week, 25°C: CYA: 23-33 mm; MEA: 20-32 mm;YES: 28-44 mm; CREA: 18-23 mm; Cz: 20-23 mm, OAT:17-28 mm: CYAS: 30-38 mm; CzBS: 12-18 mm; CzP: 0-1mm; UNO: 13-19 mm; DG18: 22-27 mmDiam., CYA, 1 week: 15°C: 16-27 mm; 30°C: 0 mm; 37°C:0 mmCYA/CYAS: 0.8 [0.8-0.9]CYA15°C/CYA 25°C: 0.8 [0.6-0.9]CYA30°C/CYA 25°C: 0CZBS/CZ: 0.7 [0.6-0.8]CZP/CZ: 0.02 [0-0.04]Distribution: Germany, New Mexico (USA), Venezuela,JapanEcology and habitats: Found twice on walls in caves, butalso on butter and salamiBiotechnological applications: Can be potentially used toproduce arisugacins.Biodeterioration & phytopathology: May degrade lipidcontaining foodsMycotoxicoses and mycotoxins: Produces territrems,tremorgenic mycotoxins, but appear to be too rare to causesignificant mycotoxicosesTypical cultures: IBT 5265 = CBS 109557 = FRR 1621 =IFO 5341, ex butter, Japan; IBT 3235 = CBS 109556 =Kulmbach Sp. 1894, ex salami, Germany; CBS 109558 =IBT 21194, ex bat cave, Venezuela (Y).72

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 36. Penicillium cavernicola. 7-day old colonies on A. CYA, B. MEA, C. YES, D-H. Conidiophores. I. Conidia. White bar= 10 µm.73

J. C. FRISVAD &R.A.SAMSONP. chrysogenum Thom, Bull. Bur. Anim. Ind. USDept. Agric. 118: 58, 1910In Penicillium subgenus Penicillium section Chrysogenaseries ChrysogenaType: Herb. IMI 024314Culture ex type: CBS 306.48 = IBT 5233* = IMI 024314 =IMI 092208 = ATCC 10106 = CCRC 30564 = FRR 807 =MUCL 29079 = MUCL 29145 = NRRL 807 = NRRL 810 =QM 7500, ex cheese, Connecticut, USA (T)Diagnostic features: Roquefortine C & D, chrysogine,penicillin F & G, globose to subglobose to broadly ellipsoidalsmooth-walled conidia, relatively short phialides withshort broad collula, high growth rate on YES with a yellowreverse and strong sporulation.Similar species: P. flavigenum is most closely related to P.chrysogenum, but differs by the slower growth rate onCYAS and the the deep yellow reverse on YES agar.Description:Conidiophores: Bi-, ter- and quarterverticillate, both appressedand divergent rami born from aerial and subsurfacehyphaeConidia: Smooth-walled, globose to subglobose to broadlyellipsoidal, 2.5-4 μm x 2.3-3.5 μmPhialides: Cylindrical, with short broad collula, 7-9 μm x2.3-2.5 μmMetulae: Cylindrical, 8-12 μm x 2.5-4 μmRami: Cylindrical, 15-20 μm x 3-4 μmStipes: 200-300 μm x 3-4 μmSynnemata or fasciculation: NoneSclerotia: NoneColony texture: Floccose to velutinousConidium colour on CYA: Blue green to greenExudate droplets on CYA: Often present, copious, yellowReverse colour on CYA: Cream, yellow, rarely brownReverse colour on YES: Citrine YellowDiffusible colour: Yellow pigment often producedEhrlich reaction: No reation or yellow reactionOdour and volatile metabolites: 3-octanone, 1-heptene, 1,3-octadiene, 3-heptanone, 1-nonene, 1-octen-3-ol, 3-octanol,(pineapple odour at low water activities) (Larsen andFrisvad, 1995)Extrolites: 1) Penicillins, 2) Roquefortine C and meleagrin,3) Chrysogine, 4) Xanthocillins, 5) Secalonic acids, 6)Sorrentanone and sorbicillin 7) PR-toxinGrowth on creatine: WeakAcid and base production on creatine: none or poor, no baseproductionGrowth on UNO: Very goodGrowth on nitrite: Occasionally goodAbiotic factors:Diam., 1 week, 25°C: CYA: 23-46 mm; MEA: 19-52 mm;YES: 40-64 mm; CREA: 16-26 mm; Cz: 18-33 mm, OAT:18-42 mm: CYAS: 27-43 mm; CzBS: 7-24 mm; CzP: 0mm; UNO: 15-29 mm; DG18: 29-41 mmDiam., CYA, 1 week: 5°C: 1-4 mm; 15°C: 15-26 mm;30°C: 14-27 mm; 37°C: 0-7 mmCYA/CYAS: 0.9 [0.8-1.1]CYA15°C/CYA 25°C: 0.6 [0.4-0.7]CYA30°C/CYA 25°C: 0.6 [0.4-0.8]CZBS/CZ: 0.6 [0.3-0.9]CZP/CZ: 0Distribution: Panglobal, very commonEcology and habitats: Indoor environments, deserts, driedfoods, salterns, cheeseBiotechnological applications: Production of penicillin andxanthocillin X, treatment of pulp mill waste, production ofpolyamine oxidase, polyamide oxidase, phospho-gluconatedehydrogenase, glucose oxidase, and also used for biotransformations.Biodeterioration & phytopathology: Building materialsare deterioratedMycotoxicoses and mycotoxins: PR-toxin, roquefortine C,secalonic acids have been regarded as mycotoxinsTypical cultures: IBT 14462 = CBS 776.95, exLechuiguilla Cave, Carlsbad, New Mexico, USA; IBT5304* = CBS 775.95, ex air in kitchen, Denmark; T84 =IBT 21511 = CBS 478.84 (Y), ex air, Denmark; IBT 22809= CBS 111216, ex saltern, Slovenia; IBT 21928* = CBS111215, ex sage; IBT 23019 = CBS 205.57 = ATCC 8537 =ATCC 9478 = CECT 2306 = IMI 015378 = NRRL 1209 =NRRL 824 = QM 6749, ex culture contaminant, UK; IBT22777 = CBS 111214, ex wheat bread, Italy; IBT 5848* =CBS 112208, ex soil, China; IBT 23022* = CBS 412.69, exsoil, Syria (P. harmonense); IBT 6048 = IBT 4344* = CBS355.48 = NRRL 821 = IMI 039759ii = ATCC 10108 =IHEM 3181 = MUCL 31327 = QM 7601, ex decayingbranch of Hyssopus sp., Norway (P. notatum); IBT 3363* =FRR 1142 = ATCC 48908, ex snack food, Sydney, NSW,Australia; IBT 19373* = CBS 289.53 = IMI 089373, exgelatine, UK (the only strain found that does not producemeleagrin, but does produce emodic acid and ω-hydroxyemodin); CBS 307.48 = FRR 1951 = NRRL 1951 =IMI 040233 = CECT 2802 = QM 941 = VTT D-88381, exCitrullus lenotus, Illinois, USA; IBT 4395 = IBT 6067 =NRRL 820 = IMI 092220 (P. griseoroseum); IBT 4350 =IBT 6062 = CBS 349.48 = NRRL 836 = ATCC 10468 =IFO 8143 = IMI 039762 = QM 7598 (AUT, P. meleagrinum);IBT 3361 = IMI 041606 (P. camerunense); IBT 3363= IMI 092241 (P. flavidomarginatum); IMI 129964 (P.aromaticum var. microsporum)*These isolates produce the unknown indole metabolite Ø74

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 37. Penicillium chrysogenum. 7-day old colonies on A. CYA, B. MEA, C. YES, D-H. Conidiophores. I. Conidia. Whitebar = 10 µm.75

J. C. FRISVAD &R.A.SAMSONP. clavigerum Demelius, Verh. Zool.-Bot. Ges.Wien 72: 74, 1922In Penicillium subgenus Penicillium section Penicilliumseries ClaviformiaType: Herb. IMI 039807Culture ex type: CBS 255.94 = CBS 310.48 = IBT 21512 =IBT 14993 = IMI 039807 = NRRL 1003 = ATCC 10427, exman, Winnipeg, Mannitoba, Canada (T, Y)Diagnostic features: Long acicular synnemata especiallyon OAT and MEA, ellipsoidal smooth-walled conidia,patulin, penitrem A, asperfuran, very poor growth onCYAS, poor growth on CREA with no acid production.Similar species: P. clavigerum differs from P. vulpinum byits thin acicular synnemata.Description:Conidiophores: Born on the synnemata rarely from aerialmycelium, sometimes sinuous, very long growing along thesynnema, 3.5-4.5 μm wideConidia: Smooth-walled, ellipsoidal, 3-4 μm x 2.2-3 μmPhialides: Cylindrical, slowly tapering to a short but distinctcollulum, 7-9 μm x 2-3 μmMetulae: Cylindrical, 10-12 μm x 3.5-4 μmRami: 12-15 μm x 3.5-4.5 μmStipes: Smooth walled, rough at the base,Synnemata or fasciculation: Long acicular synnemata withconidiophores born along the entire lengthSclerotia: NoneColony texture: Coremiform, synnemata 10-40 mmConidium colour on CYA: Grey green near tea green andslate oliveExudate droplets on CYA: AbsentReverse colour on CYA: Dark brownReverse colour on YES: Yellow to yellow or orange brownDiffusible colour on CYA: Often present, yellow to orangebrown,Ehrlich reaction: No or yellow reactionin one isolate aviolet reactionOdour and volatile metabolites: Geosmin, thujopsene, ethylacetate, ethyl isobutanoate, isobutyl actate, ethyl 2-methylbutanoate, ethyl isopentanoate, styrene (Larsen &Frisvad, 1995)Extrolites: 1) Patulin, 2) Asperfuran, 3) Norlichexanthone,4) TAN-1612, 5) Penitrem A, 6) Cyclopiazonic acid, 7)ViomelleinGrowth on creatine: PoorAcid and base production on creatine: No acid (nor base)productionGrowth on UNO: WeakGrowth on nitrite: GoodAbiotic factors:Diam., 1 week, 25°C: CYA: 13-31 mm; MEA: 17-37 mm;YES: 17-32 mm; CREA: 13-28 mm; Cz: 20-27 mm, OAT:25-37 mm: CYAS: 0-8 mm; CzBS: 0-11 mm; CzP: 0 mm;UNO: 3-7 mm; DG18: 10-18 mmDiam., CYA, 1 week: 15°C: 10-13 mm; 30°C: 7-13 mm;37°C: 0 mmCYA/CYAS: 6.3 [3-11.5]CYA15°C/CYA 25°C: 0.6 [0.3-0.8]CYA30°C/CYA 25°C: 0.5 [0.4-0.6]CZBS/CZ: 0.6 [0-0.6] (5 isolates no growth on CzBS, 2isolates 0.5 & 0.6)CZP/CZ: 0Distribution: UK, DC & Wyoming, USA, Alberta &Manitoba, CanadaEcology and habitats: Found in soil, associated to rodentshas been found twice in hospitals.Biotechnological applications: NoneBiodeterioration & phytopathology: NoneMycotoxicoses and mycotoxins: Penitrem A, patulin,cyclopiazonic acid, viomellein are produced, but P.clavigerum has not yet been found in foods.Typical cultures: IBT 14991 = NRRL 1004, Inst. OfHealth, Washington DC, USA; IBT 5523 = IBT 3507 = IBT3830 = IMI 297557; USA; IBT 18977 = CBS 112482 =UAMH 2766, ex gopher hair, Cardston, Alta., Canada; IBT18974 = CBS 112564 = UAMH 452, rodent survey, Alberta,Canada; IBT 18973 = CBS 112563 = UAMH 450,rodent survey, Alberta, Canada; IBT 19361 = CBS 112484= IMI 299048, USA; IBT 20478 = CBS 112483, ex soil,rodent hole, Laramie Basin, Wyoming, USA; IBT 18976 =CBS 112436 = UAMH 2767, ex gopher hair, Cardston,Alta., Canada; IBT 19355 = IFO 5755 = FAT 1161, ex soil,Japan; IMI 224631 = CBS 189.89, ex soil, UK; IBT 3461 =IBT 3767 = IBT 3784 = IBT 5482 = IBT 5524, ex soil, UK.76

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 38. Penicillium clavigerum. 7-day old colonies on A. CYA, B. MEA, C. YES, D-H. Conidiophores. I. Conidia. White bar= 10 µm.77

J. C. FRISVAD &R.A.SAMSONP. commune Thom, Bull. Bur. Anim. Ind. USDept. Agric. 118: 56, 1910In Penicillium subgenus Penicillium section Viridicataseries CamembertiType: Herb. IMI 039812Culture ex type: CBS 311.48 = IBT 6200 = IMI 039812 =FRR 890 = NRRL 890 = ATCC 10428 = IFO 5753 (T)Diagnostic features: Cyclopiazonic acid, rugulovasine A &B, palitantin, subglobose to ellipsoidal smooth-walledconidia, good growth on CREASimilar species: P. palitans can be distinguished from P.commune by its green conidia and good sporulaiton on YESagar. P. camemberti is much more floccose than P. commune.Description:Conidiophores: Terverticillate, appressed elements, bornfrom subsurface hyphaeConidia: Smooth-walled, globose to subglobose to (rarely)ellipsoidal, 3.5-4.5 μm.Phialides: Cylindrical tapering to a distinct collulum, 9-12μm x 2.5-3 μmMetulae: Cylindrical, 10-15 μm x 3-4 μmRami: Cylindrical, 15-25 μm x 3-4 μmStipes: Rough-walled, 200-400 μm x 3-4 μmSynnemata or fasciculation: Weakly fasciculateSclerotia: NoneColony texture: Velutinous to floccose or weakly fasciculateConidium colour on CYA: Blue green to greenExudate droplets on CYA: Often present, clearReverse colour on CYA: Cream coloured to beige or creamyellowReverse colour on YES: Cream coloured to yellowDiffusible colour on CYA: NoneEhrlich reaction: Strong violet reactionOdour and volatile metabolites: Isobutanol, isopentanol,styrene, 3-octanone, ß-caryophyllene (?), ethyl acetate, 3-heptanone, 1-octen-3-ol, 3-octanol, 2-methyl-isoborneol(Larsen and Frisvad, 1995)Extrolites: 1) Cyclopaldic acid and chromanols, 2) Palitantin,3) Rugulovasine A & B, 4) Cyclopiazonic acids, 5)Viridicatins, 6) Aspereynone-like metabolites (Frisvad andFiltenborg, 1989)Growth on creatine: Very goodAcid and base production on creatine: Mostly strong acidand base production, few isolates only show acid productionunder the colonyGrowth on UNO: Very goodGrowth on nitrite: GoodAbiotic factors:Diam., 1 week, 25°C: CYA: (15-)21-35 mm; MEA: (16-)20-37 mm; YES: 29-50 mm; CREA: 14-28 mm; Cz: 19-29mm, OAT: 20-34 mm: CYAS: 19-34 mm; CzBS: 7-27 mm;CzP: 0-9 mm; UNO: 15-25 mm; DG18: 25-30 mmDiam., CYA, 1 week: 15°C: 23-29 mm; 30°C: 0-4 mm;37°C: 0 mmCYA/CYAS: 1.0 [0.6-1.4]CYA15°C/CYA 25°C: 1.0 [0.8-1.3]CYA30°C/CYA 25°C: 0.0 [0-0.2]CZBS/CZ: 0.8 [0.4-1.1]CZP/CZ: 0.3 and three at 0 [0-0.3]Distribution: Temperate regions, on cheese in refrigeratorsin warmer areas, Denmark, Greenland, Norway, UK, theNetherlands, Italy, Spain, Turkey, USA, Canada, Bahamas,Japan, Australia, New ZealandEcology and habitats: Cheese, nuts, wheat bread, driedfish, cherries, litchis, bromeliads, wood, Sorbus endophyte,bee larvae, and soil.Biotechnological applications: None, however the domesticatedform is P. camemberti (Pitt et al., 1986; Polonelli etal., 1987)Biodeterioration & phytopathology: The major fungusdeteriorating cheese (Lund et al., 1995; Hocking and Faedo,1992; Tzanetakis et al., 1987; Kure and Skaar, 2000; Kureet al., 2001). P. palitans is also quite common in Norwegiancheeses and may be separated from P. commune by differencesin several feature (Polonelli et al., 1987; Lund, 1995b;Kure et al., 2002). Fingerprinting has been used to separateisolates within these species (Lund and Skouboe, 1998;Hansen et al., 2003; Lund et al., 2003; Kure at al., 2003).Mycotoxicoses and mycotoxins: The production of cyclopiazonicacid was first reported from P. griseofulvum, atthat time misidentified as P. cyclopium (Holzapfel, 1968;Hermansen et al., 1984; Frisvad, 1989). Despite this it haslater incorrectly been claimed that P. cyclopium produced it(Bennett and Klich, 2003). The production of cyclopiazonicacid was later reported from P. camemberti (Still et al.,1978) and P. commune (Frisvad, 1985; Pitt et al., 1986;Polonelli at al. 1987; El-Banna et al. (1987); Frisvad andFiltenborg, 1989). This mycotoxin may be produced directlyon the cheese or may enter meltet cheeses.Typical cultures: IBT 21513 = CBS 468.84 = FRR 2926 = IMI285507, ex liquorice root (Y); IBT 14135 = CBS 279.67, exRoquefort cheese; IBT 3430 = CBS 112080 = IMI 291543, exanimal feed nuts, UK; IBT 10762 = CBS 112470, ex Frenchcheese, Paris; IBT 10924 = CBS 269.97 = CBS 112079, ex fetacheese; IBT 14083 = CBS 111835, mummified bee larva, USA;IBT 21896 = CBS 112471, ex patient, the Netherlands; IBT 23305= CBS 112472, ex ice, Svalbard, Norway; NRRL 845, contaminantin P. puberulum NRRL 1889; CBS 282.36 = NRRL 879 =IBT 16113, ex sweet water in glycerol still, UK (P. lanosoviride);CBS 247.32 = NRRL 873 = IBT 12807 (P. ochraceum var.macrosporum), ex mouldy tobacco, UK; NRRL 935 = IBT 21597(P. australicum); CBS 327.48 = NRRL 948 = IBT 21599 (P.flavoglaucum); NRRL 930 = IBT 21605 (P. lanosogriseum); CBS216.30 = NRRL 894 = IBT 21606 (P. lanosogriseum), ex leafmould, Spounderwout, Bussum, the Netherlands; NRRL 932 =IBT21598 = CBS 265.29 = CBS 254.31 = ATCC 10116 = FRR 932 =IMI 192904 (P. psittacinum), ex air; CBS 341.59 = ATCC 18381= IMI 068234 = IFO 6237 = QM 7292 (P. roqueforti var. punctatum),ex cheese, Japan; IBT 3470 = IBT 3467 = IMI 295179 =ATCC 56608, ex turnips, Denmark; IBT 18102 = FRR 4192, excheddar cheese, Sydney, Australia; CBS 343.51 (P. cyclopium var.album); IBT 3469 = FRR 2160, ex cheddar cheese, Sydney, NSW,Australia.78

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 39. Penicillium commune. 7-day old colonies on A. CYA, B. MEA, C. YES, D-H. Conidiophores. I. Conidia. White bar =10 µm.79

J. C. FRISVAD &R.A.SAMSONP. concentricum Samson, Stolk and Hadlok,Stud. Mycol. (Baarn) 11: 17, 1976In Penicillium subgenus Penicillium section Penicilliumseries ClaviformiaType: Herb. CBS 477.75Culture ex type: CBS 477.75 = IBT 14571 = IBT 6577 =FRR 1715, ex colon of a deer, Germany (T)Diagnostic features: Patulin, patulidin, roquefortine C,meleagrin, oxaline, broadly ellipsoidal smooth-walledconidia, fasciculate, orange reverse on all substrates,Similar species: P. concentricum is most similar to P.glandicola. The latter species produce very rough stipes incontrast to P. concentricum.Description:Conidiophores: Terverticillate, appressed and somewhatdivergent elements, born from subsurface hyphaeConidia: Smooth-walled, Broadly ellipsoidal, 3.2-3.7 μm x2-2.5 μmPhialides: Cylindrical tapering to a distinct collulum, 5-10μm x 2.2-2.5 μmMetulae: Cylindrical, 9-13 μm x 3-3.5 μmRami: Cylindrical, 12-20 μm x 3-3.5 μmStipes: Smooth-walled (rarely finely rough-walled), 200-400 μm x 3-4 μmSynnemata or fasciculation: FasciculateSclerotia: NoneColony texture: FasciculateConidium colour on CYA: Artemisia to lily greenExudate droplets on CYA: Copious, clear to pale yellowReverse colour on CYA: Orange to orange redReverse colour on YES: Yellow to strongly orangeDiffusible colour on CYA: OrangeEhrlich reaction: No reaction or yellowOdour and volatile metabolites: Not examinedExtrolites: 1) Patulin, 2) Patulodin & CT 2108A & B, 3)Asteltoxin, 4) Barceloneic acid 5) Pyripyropens 6) RoquefortineC, meleagrin and oxaline, 7) CyclopiamineGrowth on creatine: Very goodAcid and base production on creatine: Good acid productionor just visible under colony followed by weak base productionGrowth on UNO: Very goodGrowth on nitrite: GoodAbiotic factors:Diam., 1 week, 25°C: CYA: 12-25 mm; MEA: 7-16 mm;YES: 18-40 mm; CREA: 4-20 mm; Cz: 4-16 mm, OAT: 21-30 mm: CYAS: 15-21 mm; CzBS: 3-15 mm; CzP: 0 mm;UNO: 6-15 mm; DG18: 17-22 mmDiam., CYA, 1 week: 15°C: 15-20 mm; 30°C: 0 mm; 37°C:0 mmCYA/CYAS: 1.1 [0.8-1.3]CYA15°C/CYA 25°C: 1.0 [0.8-1.3]CYA30°C/CYA 25°C: 0CZBS/CZ: 0.7 [0.5-1.0]CZP/CZ: 0Distribution: Denmark, Sweden, Norway, Germany,United Kingdom, France, South Carolina, Wisconsin,Kansas, USAEcology and habitats: Deer dung, soil (with deer dung),salami, cheese, wheat flour; water tanks (P. concentricumcan be regarded as a faecal indicator)Biotechnological applications: NoneBiodeterioration & phytopathology: UnknownMycotoxicoses and mycotoxins: As in other coprophilicspecies, many antibiotically active extrolites are found,including patulin, patulidin, barceloneic acid and meleagrin,but P. concentricum has rarely been found in foods.Typical cultures: IBT 21514 = IBT 20230 = CBS 101024,ex soil, University of South Carolina, USA (Y); IBT 5625,ex pig feed, Norway; IBT 5623 = IBT 4372 = IBT 3848 =CBS 185.89 = CBS 110762 = IMI 293197 = ATCC 58613= FRR 3066, ex wheat flour, Denmark; IBT 5621 = IBT3080 = CBS 285.36 = CBS 110765 = NRRL 2034 = ATCC46510 = IMI 326061; IBT 22163 = CBS 110763, ex dungfrom white-tailed deer, Turkey Swamp, New Jersey, USA;IBT 13685 = CBS 110764, ex soil at Chateau Menthon, St.Bernard, France; IBT 6778 = IBT 3078 = CBS 191.88 =NRRL 13633 = IMI 321510 = ATCC 64635, ex soil,Denmark; IBT 13168 = RMF 8051; ex soil, Konza Grassland,LTER, Kansas; IBT 5629 = IBT 3079 = IMI 285527 =Kulmbach Sp. 831, ex salami, Germany; IBT 12736 = WSF2352, soil in floodplain forest (Maple-Ash-Elm), Wisconsin,USA; IBT 3847 = CBS 631.70; IBT 5618, ex cheese,Denmark; UAMH 3893, ex oil-spilled soil, Norman WellsNWT, Canada.80

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 40. Penicillium concentricum. 7-day old colonies on A. CYA, B. MEA, C. YES, D-H. Conidiophores. I. Conidia. Whitebar = 10 µm.81

J. C. FRISVAD &R.A.SAMSONP. confertum (Frisvad, Filt. & Wicklow) Frisvad,Mycologia 81: 852, 1989In Penicillium subgenus Penicillium section Chrysogenaseries MononematosaType: Herb. IMI 296930Culture ex type: IBT 21515 = IBT 3098 = IBT 3093 = IBT5672 = CBS 171.87 = IMI 296930 = NRRL 13488 = NRRLA-26904, ex external fur-lined cheek pouch of Dipodomysspectabilis, 6 km east of Portal, Arizona, USA (T,Y)Diagnostic features: Asteltoxin, meleagrin, sinoid conidiophorestipes, good growth at 37°C.Similar species: P. confertum is most closely related toPenicillium mononematosum, but differs by its less complicatedpenicilli and thin often sinoid stipes.Description:Conidiophores: Terverticillate, sinoid, appressed andsomewhat divergent elements, born from subsurface hyphaeConidia: Smooth-walled, subglobose to broadly ellipsoidal,3.2-3.7 μm x 2.2-3.0 μmPhialides: Flask shaped with a distinct collulum, 7-9 μm x2.5-3.0 μmMetulae: Cylindrical, 10-15 μm x 3-4 μmRami: Cylindrical, 15-25 μm x 3-4 μmStipes: Thin, smooth-walled, sinoid, 150-300 μm x 2.5-3.5μmSynnemata or fasciculation: NoneSclerotia: NoneColony texture: VelutinousConidium colour on CYA: Greyish green to greyish turquoiseExudate droplets on CYA: Copious, clear to paleReverse colour on CYA: Cream to beigeReverse colour on YES: Yellowish cream to curryDiffusible colour: NoneEhrlich reaction: None to faint yellowOdour and volatile metabolites: Not examinedExtrolites: 1) Asteltoxin, 2) Secalonic acid D, 3) RoquefortineC & meleagrinGrowth on creatine: WeakAcid and base production on creatine: Acid under colony,no base productionGrowth on UNO: WeakGrowth on nitrite: WeakAbiotic factors:Diam., 1 week, 25°C: CYA: 20-24 mm; MEA: 17-29 mm;YES: 26-36 mm; CREA: 18-24 mm; Cz: 14-24 mm, OAT:23-29 mm: CYAS: 24-26 mm; CzBS: 1-2 mm; CzP: 0 mm;UNO: 11-13 mm; DG18: 22-25 mmDiam., CYA, 1 week: 5°C: 2-4 mm; 15°C: 13-15 mm;30°C: 18-20 mm; 37°C: 7-10 mmCYA/CYAS: 0.9CYA15°C/CYA 25°C: 0.6CYA30°C/CYA 25°C: 0.8CZBS/CZ: 0.04CZP/CZ: 0Distribution: Arizona & New Mexico, USAEcology and habitats: Kangaroo rat mounds, desertsBiotechnological applications: NoneBiodeterioration & phytopathology: UnknownMycotoxicoses and mycotoxins: Asteltoxin, but the fungusis apparently not food or feed-borne.Typical cultures: IBT 16864, ex soil under Atriplex gardneri,Chetro Ketl, Chaco Canyon, New Mexico, USA.82

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 41. Penicillium confertum. 7-day old colonies on A. CYA, B. MEA, C. YES, D-H. Conidiophores. I. Conidia. White bar =10 µm.83

J. C. FRISVAD &R.A.SAMSONP. coprobium Frisvad, Mycologia 81: 853, 1989In Penicillium subgenus Penicillium section Penicilliumseries ClaviformiaType: Herb. IMI 293209Culture ex type: CBS 561.90 = IBT 3069 = IBT 4583 =IBT 6932 = IBT 21516 = IMI 293209 = ATCC 58615, expig feed, Norway (T, Y)Diagnostic features: Patulin, roquefortine C, meleagrin,ellipsoidal smooth-walled conidia, fasciculate, small synnemata,pale reverse MEA, strong fruit-like smell.Similar species: P. coprophilum differs from P. coprobiumby uneven colony margins, and a dark brown reverse onCYA. P. concentricum has small colonies with a strongorange reverse on MEA, while the reverse of P. coprobiumis cream coloured to light yellow.Description (see also Kubatova (1993/1994) for colourphotographs, as P. coprophilum):Conidiophores: Terverticillate, appressed elements butdivergent rami, born from subsurface hyphaeConidia: Smooth-walled, broadly ellipsoidal, 3.2-4 μm x2.5-3.0 μmPhialides: Cylindrical tapering to a distinct collulum, 6.5-10μm x 2.0-2.5 μmMetulae: Cylindrical, 9-13 μm x 2.5-3 μmRami: Cylindrical, 12-20 μm x 3-3.5 μmStipes: Smooth-walled, 200-400 μm x 3-4 μmSynnemata or fasciculation: Fasciculate, synnemata up to 1mm with white stalks and green heads best developed onOATSclerotia: White to pink and rare only seen on MEA after 3weeks in a glass Petri dish (rare)Colony texture: Fasciculate, not sulcate or only weaklysulcateConidium colour on CYA: Grey green to dark greenExudate droplets on CYA: Copious, clear to pale brownReverse colour on CYA: Greyish brown to yellow brownReverse colour on YES: Cream coloured to curry to oliveDiffusible colour: Pale orange brownEhrlich reaction: None (one isolate faint yellow)Odour and volatile metabolites: Isobutanol, isopentanol,1,3,5-heptatriene and two isomers, 2-pentanone, isobutylacetate, 1-nonene, styrene, 1-undecene, (blackberry smell)(Larsen & Frisvad, 1995)Extrolites: 1) Patulin, 2) Pyripyropens, 3) Cyclopiamin, 4)Roquefortine C, meleagrin and neoxaline (Frisvad andFiltenborg, 1989)Growth on creatine: Very goodAcid and base production on creatine: No acid productionor acid just under colonyGrowth on UNO: Very goodGrowth on nitrite: GoodAbiotic factors:Diam., 1 week, 25°C: CYA: 21-29 mm; MEA: 9-26 mm;YES: 25-34 mm; CREA: 3-15 mm; Cz: 12-17 mm, OAT:18-31 mm: CYAS: 16-28 mm; CzBS: (0-) 4-17 mm; CzP: 0mm; UNO: 5-16 mm; DG18: 23-29 mmDiam., CYA, 1 week: 15°C: 14-19 mm; 30°C: 0-4 mm;37°C: 0 mmCYA/CYAS: 1.2 [0.9-1.3]CYA15°C/CYA 25°C: 0.7 [0.5-0.9]CYA30°C/CYA 25°C: 0.1 [0-0.2]CZBS/CZ: 0.7 [one 0, 0.3-1.1]CZP/CZ: 0Distribution: Denmark, Norway, the Netherlands, Germany,United Kingdom, Czech Republic, Slovakia, Siberia,Russia, China, Kansas, USAEcology and habitats: Dung, soil and roots, cereal basedanimal feed, teaBiotechnological applications: NoneBiodeterioration & phytopathology: UnknownMycotoxicoses and mycotoxins: Patulin and roquefortineC are produced, but P. coprobium has only been foundsporadically on foods and feeds. The species may indicatefaecal contamination of feeds.Typical cultures: IBT 6895 = IBT 3068 = IBT 4516 = CBS184.88 = ATCC 64630 = IMI 321497 = NRRL 13626, exsoil, Houtribdijk, the Netherlands; IBT 3070 = IBT 3707 =IBT 3777 = IBT 6899 = CBS 185.88, ex pig feed, Norway;IBT 6897 = IBT 3071 = CBS 267.97, ex rabbit dung; IBT15439 = CBS 280.97, ex Hordeum vulgare, Denmark; IBT6900 = CBS 562.90 = CCF 2005, ex ČeskomoravskáVrchovina hills, Czech Republic; IBT 18861 = CBS 110762= CCF 2800, ex soil Vrchy hills, Eastern Bohemia, CzechRepublic; IBT 19342 = CBS 110761 = FRR 3645 = PREM47700, ex grass roots, mangrove, Natal, South Africa; IBT13168 = RMF 8051; ex soil, Konza Grassland, LTER,Kansas; IBT 22719, ex soil under snow 3 km vest ofUthoss, Siberia, Russia; IBT 24820, tea, China; IBT 14656,ex potato, Denmark; IBT 3066, ex walnut;IBT 4369 = IBT 3067 = IBT 10555 = IMI 321499, ex soil,Loosdrecht, the Netherlands; IBT 18862 = CCF 2802, exsoil at Hlinsko, Czech Republic; IBT 18859 = FFC 2803, exsoil, Strážovské vrchy hills, Slovakia.84

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 42. Penicillium coprobium. 7-day old colonies on A. CYA, B. MEA, C. YES, D-H. Conidiophores. I. Conidia. White bar= 10 µm.85

J. C. FRISVAD &R.A.SAMSONP. coprophilum (Berk. & Curt.) Seifert & Samson,Adv. Pen. Asp. Syst.: 145, 1985In Penicillium subgenus Penicillium section Penicilliumseries ClaviformiaType: herb. K, Cuba, Wright, 666Ex epitype cultures: CBS 110760 = IBT 5551 = IBT 3064,ex rabbit dung, Groeneveld, Baarn, the Netherlands (epiT)Diagnostic features: Griseofulvin, roquefortine C, meleagrin,oxalin, broadly ellipsoidal smooth-walled conidia,synnemata produced, uneven margin of colonies on MEA,dark brown reverse on CYASimilar species: See P. coprobium.Description:Conidiophores: Terverticillate, appressed elements butdivergent rami, born from subsurface hyphaeConidia: Smooth-walled, broadly ellipsoidal, 3.2-3.7 μm x2.0-2.5 μmPhialides: Cylindrical tapering to a short distinct collulum,6.5-10 μm x 2.2-2.5 μmMetulae: Cylindrical, 9-13 μm x 3-3.5 μmRami: Cylindrical, 12-20 μm x 3-3.5 μmStipes: Smooth- and rough-walled, 200-400 μm x 3-4 μmSynnemata or fasciculation: Fasciculate, synnemata up to1.3 mm with white stalks and green headsSclerotia: NoneColony texture: Fasciculate to coremiformConidium colour on CYA: Green to dark greenExudate droplet on CYA: Copious, clear to pale brownReverse colour on CYA: Dark brownReverse colour on YES: Curry to brown yellowDiffusible colour on CYA: BrownEhrlich reaction: NoneOdour and volatile metabolites: Isobutanol, isopentanol, 1-octene, styrene, 2-pentanone, 3-methyl-2-pentanone, methylisopentanoate, 1-nonene, 2-heptanol, 2-octanone, 3,4-dimethyl-hexanone, 1-methoxy-3-methyl-benzene, 1-undecene (Larsen and Frisvad, 1995)Extrolites: 1) Griseofulvin, 2) Alternariol, 3) Pyripyropens,4) Roquefortine C, meleagrin, oxalin, neoxalin 5) CyclopiaminGrowth on creatine: Very goodAcid and base production on creatine: Good acid productionfollowed by base production, occasionally only acid underthe colonyGrowth on UNO: Very goodGrowth on nitrite: GoodAbiotic factors:Diam., 1 week, 25°C: CYA: 18-29 mm; MEA: 9-26 mm;YES: 25-34 mm; CREA: 3-15 mm; Cz: 12-17 mm, OAT:18-31 mm: CYAS: 16-28 mm; CzBS: (0-) 4-17 mm; CzP: 0mm; UNO: 5-16 mm; DG18: 23-29 mmDiam., CYA, 1 week: 15°C: 14-19 mm; 30°C: 0-4 mm;37°C: 0 mmCYA/CYAS: 1.0 [0.9-1.2]CYA15°C/CYA 25°C: 0.6 [0.6-0.8]CYA30°C/CYA 25°C: 0.6 [03-0.7]CZBS/CZ: 1.0 [0.9-1.3]CZP/CZ: 7 isolates 0, two isolates 0.2 [0.2-0.3]Distribution: Denmark, Norway, United Kingdom, Germany,South Carolina, New Jersey & Wisconsin (USA),Cameroon, Zambia, Ethiopia, South Africa, Australia, NewZealandEcology and habitats: Rabbit and deer dung, soil withherbivore dung, corn, rye bread, barley, compound feed,salami, hazelnutsBiotechnological applications: NoneBiodeterioration & phytopathology: UnknownMycotoxicoses and mycotoxins: Roquefortine C andalternariol are produced, but the species did not produceconspicuous growth on the food substrate from which theywere isolated.Typical cultures: CBS 186.89 = IBT 21517 = IBT 3768 =IBT 5539 = NRRL 13627, ex Andropogon sorghum importedto Denmark (Y); IBT 23268 = CBS 102444, exwoodchip paper behind skirting board, Germany; IBT 5546= CBS 473.75, ex Zea mays, Pretoria, South Africa; IBT3063 = IBT 3845 = IBT 4753 = IBT 5552 = CBS 110759Kulmbach Sp. 1370, ex salami, Germany; IBT 18704 =CBS 110758, ex rye bread, Denmark; IBT 22162 = CBS110757, ex Styra cifolia (liquid amber), Turkey Swamp,New Jersey, USA; IBT 12724 = CBS 110767, ex Hordeumvulgare, the Netherlands; IBT 12992 = CBS 272.97 = CBS110768, ex wheat, Denmark; FRR 1403, ex chicken feed,Brisbane, Qld., Australia; IBT 12750 = WSF 5238.86

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 43. Penicillium coprophilum. 7-day old colonies on A. CYA, B. MEA, C. YES, D-H. Conidiophores. I. Conidia. Whitebar = 10 µm.87

J. C. FRISVAD &R.A.SAMSONP. crustosum Thom, Penicillia: 399, 1930In Penicillium subgenus Penicillium section Viridicataseries CamembertiType: Herb. IMI 091917Culture ex type: IBT 5528 = IBT 6175 = IMI 091917 =FRR 1669 = ATCC 52044 = NCTC 4002, ex lemon, Aberdeen,Scotland (T)Diagnostic features: Penitrem A, roquefortine C, terrestricacid, viridicatin, smooth-walled conidia, crustose on CYAand MEA after 7-10 days, high growth rate, good growth onCREA. Strong sporulation and high growth rate on YESagar.Similar species: P. expansum has ellipsoidal conidia andusually smooth stipes and is not crustose. P. commune hassmaller colony diameters, not a deep yellow reverse on YESand is not crustose. P. palitans has smaller colonies, darkerand greener condia and is not crustose.Description:Conidiophores: Terverticillate, appressed elements, bornfrom subsurface hyphaeConidia: Smooth-walled, globose to subglobose, 3-4 μm.Phialides: Cylindrical tapering to a distinct collulum, 9-12μm x 2.5-3 μmMetulae: Cylindrical, 10-15 μm x 3-3.5 μmRami: Cylindrical, 15-25 μm x 3.5-4 μmStipes: Rough-walled, 200-400 μm x 3.5-4.5 μmSynnemata or fasciculation: Weakly fasciculateSclerotia: NoneColony texture: Velutinous to weakly fasciculate, becomingcrustoseConidium colour on CYA: Dull green to grey green or bluegreen at the colony marginExudate droplets on CYA: Copious, clear or brownReverse colour on CYA: Cream-coloured to yellow brownReverse colour on YES: Strongly yellowDiffusible colour: Pale brown or noneEhrlich reaction: NoneOdour and volatile metabolites: Ethyl acetate, isobutanol,ethyl propanoate, isopentanol, dimethyldisulphide, ethylisobutanoate, isobutyl acetate, ethyl butanoate, ethyl-2-methyl butanoate, ethyl isopentanoate, isopentyl acetate,styrene, ethyl pentanoate, 3-octanone, ethyl hexanoate, ethyloctanoate, 2-methyl-isoborneol, geosmin.Extrolites: 1) Penitrem A-G, 2) Roquefortine C-E, 3)Terrestric acid and viridicatic acid, 4) Viridicatins(cyclopeptin, dehydrocyclopeptin, cyclopenin, cyclopenol,viridicatin, viridicatol), 5) Hadacidin, 6) Andrastin AGrowth on creatine: Very goodAcid and base production on creatine: Good acid productionfollowed by base productionGrowth on UNO: Very goodGrowth on nitrite: PoorAbiotic factors:Diam., 1 week, 25°C: CYA: 32-46 mm; MEA: 25-46 mm;YES: 40-56 mm; CREA: 22-35 mm; Cz: 29-32 mm, OAT:28-38 mm: CYAS: 29-39 mm; CzBS: 25-28 mm; CzP: 0-3mm; UNO: 19-23 mm; DG18: 29-32 mmDiam., CYA, 1 week: 15°C: 28-31 mm; 30°C: 15-26 mm;37°C: 0 mmCYA/CYAS: 1.3 [1.1-1.5]CYA15°C/CYA 25°C: 0.7 [0.7-0.7]CYA30°C/CYA 25°C: 0.5 [0.4-0.6]CZBS/CZ: 0.9 [0.8-0.9]CZP/CZ: 0.1 and four at 0 [0-0.1]Distribution: Panglobal: Denmark, Norway, Germany,Slovenia, Greece, Bulgaria, Azoan Islands, Portugal, Georgia& New Mexico (USA), Kazachstan, Victoria B.C.,Indonesia; Canada, Australia, New Zealand.Ecology and habitats: Pecans, walnuts, chestnuts, apples,cream cheese, blue cheese, salami, indoor air, coffee beans,maize, cotton seed, compound feeds, cardboard.Biotechnological applications: NoneBiodeterioration & phytopathology: P. crustosum canproduce a weak rot in apples (Frisvad, 1981). It is a majordegrader of compound feeds for animals and cheeses.Mycotoxicoses and mycotoxins: Penitrem A and roquefortineC from P. crustosum growing on cheese or nuts havebeen implicated in toxicoses of dogs (Richard et al., 1981;Richard and Arp, 1979; Puls and Ladyman, 1988; Hockinget al., 1988; Naudé et al., 2002). The case of mycotoxicosisof a man after drinking beer (Cole et al., 1983; Cysewski etal., 1975) was caused by P. carneum, not P. crustosum).Cattle has also been intoxicated with penitrem A from P.crustosum (Wilson et al., 1968; Dorner et al., 1984). Strainsidentified as P. palitans (Ciegler, 1969), P. commune(Mintzlaff et al., 1972; Wagener et al., 1980), P. lanosocoeruleum(Wells and Cole, 1977), P. cyclopium (Wilson,1968; Ciegler and Pitt, 1970; Vesonder et al., 1980), P.martensii, P. olivinoviride, P. puberulum (Ciegler and Pitt,1970) producing penitrem A and roquefortine C were all P.crustosum (Pitt, 1979b, Frisvad, 1989). The production ofboth toxins is very consistent in P. crustosum (El-Banna andLeistner, 1988; Frisvad and Filtenborg, 1989)Typical cultures: IBT 21518 = IBT 14747 = CBS 101025,ex cheese, Azoan Islands (Y); IBT 23265 = CBS 181.89, exsoil with Agaricus bisporus; IBT 6579 = IBT 3425 = CBS471.84 = IMI 285510 = FRR 2929, ex Thymus vulgaris,Greece; IBT 6580 = IBT 5529 = CBS 110076 = FRR 1387,ex Portugese wine corks, Sydney, NSW, Australia; IBT18099 = CBS 110077 = FRR 1513, ex deteriorating preservedwood-stakes; North Qld., Australia; IBT 15977 =CBS 110075, ex mixed pig feed, Bulgaria; IBT 14519 =CBS 110074, ex Lechuiguilla Cave, Carlsbad, New Mexico;IBT 16510 = CBS 110073, ex soil, ex soil under Betula sp.,Victoria B.C., Canada; IBT 11095 = CBS 499.73 = IMI174717 = FRR 1478 = VKM F-1746 = ATCC 24721, exgrapes in storage, Alma-Ata, Kazachstan (P. farinosum);IBT 11150 = CBS 340.59 = QM 7293 = ATCC 18379 =FRR 3457 = IFO 6235 = IMI 068235, ex soil, Japan (P.pseudocasei); IBT 3426 = ATCC 32014, ex weevildamagedpecans, Georgia, USA; IBT 6578 = FRR 2223, exweevil-damaged pecans, Georgia, USA; IBT 16885, exmouldy onion, Denmark; IBT 18359 = CCRC 32633,Taiwan; IBT 23710 = IMI 206159, ex soil, New Zealand.88

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 44. Penicillium crustosum. 7-day old colonies on A. CYA, B. MEA, C. YES, D-H. Conidiophores. I. Conidia. White bar =10 µm.89

J. C. FRISVAD &R.A.SAMSONP. cyclopium Westling, Ark. Bot. 11: 90, 1911In Penicillium subgenus Penicillium section Viridicataseries ViridicataType: Herb. IMI 089372Culture ex type: CBS 144.45 = CBS 114.74 = IBT 5130 =IMI 089372 = ATCC 8731 = ATHUM 2888 = CECT 2264= DSM 1250 = LSHB 1923 = MUCL 15613 = QM 6839 =VKM F-265, ex fruit, Norway (T)(P. puberulum Bain. 1907 is a nomen dubium as the cultureex neotype contains both material of P. commune (NRRL845 = ATCC 8508) and P. cyclopium (NRRL 1889 = NRRL2040))Diagnostic features: Xanthomegnin, viomellein, vioxanthin,penicillic acid, blue green smooth-walled conidia,green on CYA and blue green on MEASimilar species: See P. aurantiogriseumDescription:Conidiophores: Terverticillate, appressed elements, bornfrom subsurface hyphaeConidia: Smooth-walled, globose to subglobose, 2.6-3.2μm.Phialides: Cylindrical tapering to a distinct collulum, 8-9μm x 2.6-2.8 μmMetulae: Cylindrical, 9.5-14 μm x 3.2-4.4 μmRami: Cylindrical, 15-25 μm x 3.2-4.4 μmStipes: Rough-walled, 200-750 μm x 3.5-5 μmSynnemata or fasciculation: Weakly fasciculateSclerotia: NoneColony texture: Velutinous to weakly fasciculateConidium colour on CYA: Blue green to green, blue greenon MEAExudate droplets on CYA: Copious, clear or light yellowReverse colour on CYA: Yellow to orange to red brownoccasionally only cream yellowReverse colour on YES: Strongly yellow, also yellowobverse (poor sporulation on YES)RT agar: Dark brown halo in most isolatesDiffusible colour: None or pink or red brownEhrlich reaction: pink to red or yellow brownOdour and volatile metabolites: Close to the volatile profileof P. aurantiogriseum, but weakerExtrolites: 1) Penicillic acid, 2) Xanthomegnin, viomelleinand vioxanthin, 3) Puberulic acid and puberulonic acid 4)Puberuline and verrucofortine (rugulosuvine and leucyltryptophanyldiketopiperazine),5) 3-Methoxyviridicatin (andcyclopeptin, dehydrocyclopeptin, cyclopenin, cyclopenol,viridicatol)Growth on creatine: Poor growthAcid and base production on creatine: Good acid production,no base productionGrowth on UNO: WeakGrowth on nitrite: Poor.Abiotic factors:Diam., 1 week, 25°C: CYA: 18-34 mm; MEA: 21-35 mm;YES: 28-40 mm; CREA: 11-21 mm; Cz: 19-28 mm, OAT:17-29 mm: CYAS: 25-34 mm; CzBS: 12-21 mm; CzP: 0mm; UNO: 7-17 mm; DG18: 23-27 mmDiam., CYA, 1 week: 15°C: 19-24 mm; 30°C: 2-6 mm;37°C: 0 mmCYA/CYAS: 0.9 [0.8-1.1]CYA15°C/CYA 25°C: 0.8 [0.7-1.0]CYA30°C/CYA 25°C: 0.2 [0.1-0.2]CZBS/CZ: 0.8 [0.5-0.9]CZP/CZ: 0Distribution: Temperate regions with farming: Denmark,Sweden, United Kingdom, Norway, Canada, more rare inSaudi Arabia and California (USA)Ecology and habitats: Cereals: wheat, barley, oats, maize,compound feeds, more rare on salami, seaweed and harness.Biotechnological applications: NoneBiodeterioration & phytopathology: Degrades cereals andanimal feeds.Mycotoxicoses and mycotoxins: This species is a veryefficient producer of xanthomegnin, viomellein and vioxanthin,as are P. freii, P. viridicatum and P. tricolor (the latteris less common, however). These species are very commonlygrowing in low quality cereals and may cause liverand kidney diseases in domestic animals (Carlton et al.,1968; Carlton and Tuite, 1970a & b; Budiarso et al., 1968;1971; Carlton et al., 1972; Zwicker et al., 1973). The latterreference reported on probable carcinogenicity of P. viridicatum,which always produces xanthomegnin, viomelleinand vioxanthin, but also brevianamide A, viridic acid andviridamine distinguishing it from P. cyclopium (see also P.viridicatum). The mixture of penicillic acid and xanthomegnin,viomellein and vioxanthin is also present in mostisolates of Aspergillus ochraceus (Frisvad and Samson,2000). A. ochraceus give the same severe symptoms in miceand swine (Zimmerman et al., 1976, 1977; 1979; Robbers etal., 1978) as P. viridicatum indicating that xanthomegninand viomellein are more important than previously thought.It appears that ochratoxin A from P. verrucosum (or someof the strains of Aspergillus ochraceus) mostly adds to thenephrotoxicity of mouldy cereals rather than to the hepatotoxicity.Xantommegnin, viomellin and vioxanthin havebeen found to be naturally occurring in cereals (Hald et al.,1983, Scudamore at al., 1986) and the fungi responsible forthis were P. cyclopium and P. freii.Typical cultures: IBT 21519 = IBT 11415 = CBS101136 (Y), ex harness, Saudi Arabia; IBT 5171 = CBS 477.84 =IMI 285516 = FRR 2935, ex Hordeum vulgare, Denmark; IBT5167 = CBS 349.59 = QM 7314 = ATCC 18382 = FRR 1364 =IFO 6240 = IMI 068218 (P. viridicyclopium), ex soil, Japan; IBT22027 = CBS 110335, ex wheat, UK; IBT 14369 = CBS 110336,ex wheat, UK; IBT 16769 = CBS 110337, ex malting barley,Denmark; IBT 3454 = CBS 110338; IBT 5141 = CBS 123.14 =NRRL 1889 = NRRL 2040 = ATCC 10483 = ATCC 8505 =ATCC 8732 = CCRC 31519 = FRR 2040 = IFO 7733 = IMI034913 = LSHB Ad113 = QM 1556 = MUCL 29231 = MUCL31198 = QM 1556 = VMK F-328 (P. puberulum), ex maize,Lincoln, Nebraska, USA; IBT 12480 = NRRL 970 = FRR 970 (P.porraceum); IBT 5362 = FRR 1347, from chicken feed implicatedin fatty liver and kidney syndrome, Victoria, Australia.90

POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMFig. 45. Penicillium cyclopium. 7-day old colonies on A. CYA, B. MEA, C. YES, D-H. Conidiophores. I. Conidia. White bar =10 µm.91

J. C. FRISVAD &R.A.SAMSONP. digitatum (Pers.:Fr.) Sacc., Fung. Ital.: 894,1881In Penicillium subgenus Penicillium section Digitata seriesDigitataType: Icon in Saccardo, Fung. Ital.: tab. 894 Jul. 1881Epitype suggested: CBS 112082 = IBT 13068, ex Citruslimon, Italy (epiT)Diagnostic features: Tryptoquialanines, large cylindrical toellipsoidal smooth-walled olive green conidia, no growth onCzapek agar and CYASSimilar species: NoneDescription:Conidiophores: Terverticillate, appressed elements, bornfrom subsurface or aerial hyphaeConidia: Smooth-walled, ellipsoidal to cylindrical, 6-9 (-14)μm x 2.8-6 μm.Phialides: Cylindrical tapering to a distinct collulum, 10-17μm x 4-5 μmMetulae: Cylindrical, 15-25 μm x 5-6 μmRami: Cylindrical, 20-30 μm x 5-6 μmStipes: Rough-walled, 60-160 μm x 5-7 μmSynnemata or fasciculation: NoneSclerotia: NoneColony texture: VelutinousConidium colour on CYA: Olive greenExudate droplets on CYA: Only in few isolatesReverse colour on CYA: greyish olive to beigeReverse colour on YES: cream yellow with light browncenterDiffusible colour: NoneEhrlich reaction: NoneOdour and volatile metabolites: Ethylene, ethyl alcohol,ethyl acetate, methyl acetate, isopropyl acetate, octyl acetate,limonene, valancene (Birkinshaw et al., 1931; Larsenand Frisvad, 1995)Extrolites: 1) Tryptoquialanins, 2) Cholesterol, ergosta-7,22-dien-3 β-OH, ergosta-7,22-24(28)-trien-3β-OH,episterol, eburicol, 3) Phenylalanine-proline diketopiperazine(2 isomers), 4) 3,5-Dimethoxyphenol, methoxyphenol(5 isomers), 1,1’-biphenyl-3-ol, 2-methoxy-4-propyl phenol (ethyl vanillin)Growth on creatine: Very poorAcid and base production on creatine: No acidGroweth on UNO: GoodGrowth on nitrite: Neither growth on Czapek nor nitriteagarAbiotic factors:Diam., 1 week, 25°C: CYA: (15-)24-37(-55) mm; MEA:26-54 mm; YES: 44-76 mm; CREA: 0-6 mm; Cz: 13-27mm, OAT: 19-38 mm: CYAS: 0 mm; CzBS: 2-4 mm; CzP:0 mm; UNO: 3-8 mm; DG18: 44-56 mmDiam., 1 week: 15°C: 8-22 mm; 30°C: 0 mm; 37°C: 0 mmCYA/CYAS: No growth on CYASCYA15°C/CYA 25°C: 0.7 [0.3-1.1]CYA30°C/CYA 25°C: 0CZBS/CZ: 0.2 [0.1-0.3]CZP/CZ: 0Distribution: Italy, Spain, Cyprus, Turkey, Israel, Egypt,Saudi Arabia, Libya, Morocco, Nigeria, South Africa,Zimbabwe, Tanzania, Florida, California, USA, Cuba,Domenica, Argentina, Brazil, Australia, Indonesia, India,Bangla Desh, Pakistan, Japan, Korea, Taiwan, Viet Nam.On imported citrus fruits to Denmark, Sweden, UnitedKingdom, Netherlands, Belgium, France, Germany, theNetherlands, etc. (imported citrus fruits), Canada, Connecticut,North Carolina, Oklahoma, Wisconsin, USA.Ecology and habitats: Citrus fruits are often degraded byP. digitatum. The species has been found on herbs, butcitrus fruits may be the only real habitat. See also Domschet al. (1980)Biotechnological applications: NoneBiodeterioration & phytopathology: P. digitatum is amajor pathogen of citrus fruits (where it may co-occur withP. italicum and P. ulaiense) (Holmes et al., 1994)Mycotoxicoses and mycotoxins: Some metabolites wereshown to be toxic in the chick embryo test (Faid and Tantaoui-Elaraki,1989).Typical cultures: IBT 21520 = IBT 15179 = CBS 101026(Y), ex chili mix, Indonesia; IBT 12919 = CBS 351.97 =CBS 529.97, ex chervil, Denmark; IBT 15037 = CBS112083, ex Citrus sinensis; IBT 5533 = CBS 112322 = FRR288 = ATCC 52304, ex soil, foreshore of Whiteswan Lake,Saskatchewan, Canada; IBT 3816 = CBS 112081, exmandarin; IBT 23020 = CBS 136.65 = DSM 2731, ex fruitof Citrus medica limoneum, Netherlands; IBT 23021 = CBS319.48 = NRRL 786 = ATCC 10030 = IFO 7006 = IMI039748 = QM 7552 = DSM 2748, ex fruit of Citrus sp.,Connecticut, USA; CBS 492.84 = IMI 293202 = ATCC58617, ex Lipidium sativum.92

Fig. 26, Penicillium aethiopicum. 7-day old colonies at (A) CYA - CBS

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