Polyphasic taxonomy of Penicillium subgenus Penicillium A ... - CBS

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J.C. FRISVAD &R.A.SAMSONTable 4. Penicillium species and their specific habitatsTypical habitat* SpeciesIndoor air P. brevicompactum, P. chrysogenum, P. commune, P. polonicum. P. expansum, P. olsonii,pomaceous fruit P. expansum, P. crustosum, P. solitumcitrusP. italicaum, P. digitatum, P. ulaiensenutsP. discoloronionsP. radicicola, P. albocoremium, P. alliibulbs and roots P. tulipae, P. hirsutum, P. venetum , P. gladiolisilageP. paneum, P. carneummushrooms P. brevicompactumcheese hard P. commune, P. discolorcheese soft P. camemberti, P. roqueforti, P. caseifulvum,breadP. roqueforti P. paneum, P. carneumsalami & ham P. nalgiovense, P. nordicumcereal grains P. hordei, P. verrucosum, P. cyclopium, P. freii, P. tricolor,oakP. glandicoladungP. coprophilum, P. coprobium, P. concentricum, P. vulpinum, P. formosanum, P. clavigerumsoilP. persicinum,* Listed are the typical habitats of the species. For a more detailed description of occurrence see the text.Strong associationsP. italicum and P. ulaiense in series Italica and P.digitatum in series Digitata are strongly associated tocitrus fruits. There is no strong evidence they areassociated to any other plant products, although theyare occasionally reported from soil and plant roots orstems. This may be caused by cross contaminationfrom P. digitatum spoiling citrus fruits. When thesefruits are spoiled a large number of air-borne conidiaare produced. Fungi isolated directly from citrus fruitsrots are most probably one of the three species listedabove, and the same species are unlikely to thrive onany other substrate.P. sclerotigenum has only been found in associationwith yam tubers and yam flour (Discorea batatasand D. cayenensis). It appears to be strictly associatedto this particular habitat.P. gladioli has only been found on Gladiolusbulbs, but has not been reported since 1970. We wereunsuccesful to isolate this species in Europe in spiteour extensive efforts. Perhaps new ways of protectingflower bulbs from diseases have eradicated the species.P. tulipae is primarily associated to Tulipa, but ithas also been found on other flower bulbs such asChrysanthemum and Lilium. It is also occasionallyassociated to other plant roots. P. hirsutum has beenfound on some of the same bulbs, but appears to beless strictly associated to plant roots, because it hasalso been found on butter and apples. However crosscontaminationfrom plant roots cannot be excluded. P.venetum is most often found on roots of Iris, Hyacinthusand Ornithogalum, but has also been found onAsparagus.P. allii is a very common garlic rotting organismwhile P. albocoremium and P. radicicola are commonlyfound on other Allium spp., especially Alliumcepa. However the latter two species are also commonon other plant roots often used for human food.34P. verrucosum, P. hordei, and all members ofseries Viridicata are all typically associated to storedcereal grains. There may be a tendency of P. viridicatum,P. melanoconidium, P. aurantiogriseum and P.polonicum to grow on corn and wheat in warmerclimates and for P. cyclopium, P. freii, P. verrucosum,P. tricolor and P. hordei to thrive in colder climatesespecially on barley and wheat. P. neoechinulatumfrom Series Viridicata however, has until now onlybeen found on seeds gathered by kangaroo rats in theSonoran desert. P. polonicum, P. aurantiogriseum andP. viridicatum may also cross-contaminate dried meatproducts, such as salami. The contamination sourcemay be wheat flour.P. glandicola have only been reliably been recordedfrom oak trees, acorns and cork and appears tobe strictly associated to Quercus spp. The speciesname is thus well chosen. Its occasional occurrence insoil and in guts of deer may easily be explained by itsassociation to oak trees.P. roqueforti, P. carneum and P. paneum haveonly been found on substrates in which lactic acidbacteria have grown, or substrates that have the samecharacteristics. The substrates may have been preservedwith organic acids that are quite similar tolactic acid, including sorbic acid, benzoic acid, propionicacid and acetic acid, or they may contain somealcohol and/or they may have high CO 2 or low O 2content. Examples are rye bread, yoghurt, cheese orsilage (Boysen et al., 1996; Lund et al., 1996)P. camemberti, P. caseifulvum, P. commune, P.palitans and P. atramentosum appear to be mostlyassociated to cheese and other milk products (Lund etal., 1995). P. commune and P. palitans have also beenreported from dried meat products and nuts, but aremuch less common on such substrates. P. atramentosumhas been found on Norwegian gamalost and onother cheeses, but is much more prevalent in alkalinesoils (Kubatová, 1990). We have found it to be domi-
POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMnating in limestone quarry soils in Denmark togetherwith P. scabrosum (Banke and Frisvad, unpublished).P. nordicum and P. nalgiovense are very salttolerant and are mostly associated to dried or saltedmeat products, such as salami and dried hams. Bothspecies also occur on cheese, but are less commonthere (Lund et al., 1995)P. marinum is probably associated to marine algae,but too few isolates have been found to substantiatethis. The two extant isolates are from coastal sands.P. coprophilum, P. coprobium, P. concentricum, P.vulpinum, P. formosanum and P. clavigerum allappear to be strongly associated to animal dung. Thisassociation is supported by several common featuresin these fungi, including production of ellipsoidalconidia, synnema, and patulin or griseofulvin incombination with roquefortine C. The same combinationof features is also found in series Urticicolae, P.expansum and P. sclerotigenum and it is likely that thedung Penicillia, Urticicolae and P. expansum all sharea common dung-borne ancestor.Strong associations to several different habitatsP. echinulatum and P. cavernicola are often foundon lipid rich substrates such as butter, margarine andcheese. P. echinulatum has also been found on woodshavings and in wet mechanical pulp. P. cavernicolahas been found in caves with bats. There are too fewisolates of these species reported to conclude whattheir primary habitats might be.P. discolor has been found on natamycin treatedcheeses, but was originally isolated from vegetableroots and weevil-damaged pecans (Frisvad et al.,1997). It has often been isolated from walnuts, blackwalnuts, chestnuts and pecans, which may be itsprimary habitat.Plurivorous speciesThe following species grow very well at low wateractivities and at a wide range of temperatures.P. expansum, P. solitum and P. crustosum are allknown for their rot of pomaceous fruits (Raper andThom, 1949; Frisvad, 1981; Pitt et al., 1991), but theyalso occur in other quite different habitats. P. expansumand P. crustosum have been found repeatedly onnuts and oilseeds, and all three apple rotting organismshave also been found on dried meat. P. crustosumhas also been found occasionally in corn and riceand on cheese, so this species appear to be more of ageneralist among these Penicillia. P. expansum hasbeen found commonly on wood in Canadian buildings,while P. crustosum can be common on cardboard,leather, textiles and wood (K.A. Seifert, personalcom.)P. aethiopicum has been found on several tropicalplant products and in tropical soil and no particulargroup of foods seems to be particularly prone toinfection with this fungus.P. chrysogenum, P. dipodomyis, P. flavigenum, P.nalgiovense P. mononematosum and P. confertum arecommon in dry habitats and may originally haveinhabited desert habitats and salty soils. They are ableto grow on foods at low water activities. P. chrysogenumhas been found on dried cereals, salted meatand many other low water activity foods, but is alsocommon in indoor air environments together withAspergillus versicolor (Samson et al., 2004).P. thymicola has been found on dried herbs andsorghum. P. griseofulvum and P. dipodomyicola arecommon on grasses and their seeds including drybarley and wheat. The former species may also befound on pasta and white bread.P. brevicompactum, P. bialowiezense and P.olsonii are very common in soil from tropical rainforests and soil in green-houses in other areas of theworld. These species have been found on mouldymushroom, tomatoes, green coffee, in processedfoods, and many other substrates. The two formerspecies are also common in temperate forest soil(Zaleski, 1927), maybe because of their growth onbasidiocarps.P. percicinum has only been found in soil, so itsprimary habitat is unknown.Abiotic and nutritional factorsAll species in Penicillium subgenus Penicillium areable to grow at 25ºC, so they are not psychrophilesaccording to the most common definitions. Mostspecies are, however, capable of growing at 5ºC andsome are growing faster at 15 than 25ºC. Species notable to grow at 30ºC include P. bialowiezense, P.marinum, P. formosanum, P. ulaiense, P. gladioli, P.digitatum, P. solitum, P. cavernicola, P. nordicum, P.thymicola, P. verrucosum, P, freii and P. tricolor.Further species growing very poorly at 30ºC includeP. brevicompactum, P. olsonii, P. expansum, P.coprobium, P. glandicola, P. vulpinum, P. italicum, P.camemberti, P. caseifulvum, P.commune, P. palitans,P. echinulatum, and P. hirsutum. Species growingfaster at 15ºC than 25ºC include P. bialowiezense, P.marinum, P. thymicola, P. verrucosum and P. radicicola.Most species in subgenus Penicillium tolerate 5%NaCl very well. Species that grow faster on mediawith 5% NaCl than without NaCl, include P. bialowiezense,P. brevicompactum, P. olsonii, P. chrysogenum,P. dipodomyis, P. flavigenum, P. nalgiovense,P. confertum, (P. mononematosum), (P.marinum), (P. glandicola), P. gladioli, (P. commune),P. palitans, P. discolor, P. solitum, P. cavernicola, P.echinulatum, P. nordicum, P. thymicola, P. verrucosum,P. aurantiogriseum, P. cyclopium, P. freii, P.melanoconidium (P. neoechinulatum), P. polonicum,P. viridicatum, P. albocoiremium, P. allii, P. hirsutum,P. hordei, P. radicicola, and P. venetum. Series35
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Smedsgaard J. (1997b) Terverticilla
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Polyphasic taxonomy of Penicillium subgenus Penicillium A ... - CBS

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