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

also produced by P. griseofulvum in section Expansa.Anthraquinones and other yellow polyketides areproduced by most species. Xanthocillins have beenfound in two species in Chrysogena (P. chrysogenumand P. flavigenum), in the related Eupenicilliumegyptiacum, but only found in P. italicum in sectionExpansa outside Chrysogena. Chrysogine is alsocommon in section Chrysogena, and only found in P.tulipae in section Viridicata outside Chrysogena. P.mononematosum is unique in this section, however,being characterized by the production of fumitremorgins,also found in Eupenicillium crustaceum, cyclopaldicacid (also found in P. carneum in sectionRoqueforti and P. commune in section Viridicata),and isochromantoxins (found in P. steckii in subgenusFurcatum). Roquefortine C and meleagrin isfound in P. chrysogenum, but also by many otherspecies in sections Expansa and Viridicata. Dipodazinhas only been found in P. dipodomyis from thissection and in P. cavernicola from section Viridicata.As mentioned earlier all species grow well at hightemperatures, often producing colonies at 37°C. Thespecies can all grow at very low water actitivities andhigh salt (NaCl) concentrations. Only species insection Viridicata seems to be more halotolerant. P.aethiopicum, as an exception, is not very halotolerant,however. No species use creatine well as sole N-source, but all species grow moderately well onnitrite as sole N-source and very well on UNO. Thespecies are not resistant to acids, but grow well atrelatively high pH values. Compared to other speciesin subgenus Penicillium, species in Chrysogena arethose closest to Eupenicillium series Crustacea andother soil-borne Penicillia.POLYPHASIC TAXONOMY OF SUBGENUS PENICILLIUMSection PenicilliumSeries Expansa is characterized by smooth-walledellipsoidal conidia, except P. marinum and P. gladioliwhich have globose to subglobose conidia. Mostspecies have strongly fasciculate to coremiformcolonies and conidiophores with smooth stipes andterverticillate to quarterverticillate structures. P.gladioli differs by having only slightly fasciculatecolonies and rough-walled stipes.In series Urticicolae the species are unique inhaving divaricate structures and very short phialides.Most species appear to be very competitive, producingpatulin, griseofulvin, or fulvic acid or all of these(in P. griseofulvum). Furthermore all species in thesection produce roquefortine C, except P. gladioli, P.italicum and P. ulaiense. Extrolites such as deoxybrevianamideE, italinic acid, cyclopiamide, cyclopiamine,communesins, expansolides, gladiolic acids,asperfuran, and pyripyropens are only known fromsection Expansa in subgenus Penicillium. P. griseofulvum,but also P. commune and P. palitans insection Viridicata produce cyclopiazonic acid. P.expansum produces chaetoglobosins, but these metabolitesare also produced by P. discolor in sectionViridicata. All species tolerate both quite acidic andalkaline conditions and can grow at relatively lowwater activities, albeit not as low as the other sections.No species can grow at 37°C. All species arepsychrotolerant. All species in series Expansa andClaviformia grow well on creatine as sole N-source,whereas species in the other series in the section growpoorly on creatine. Several plant pathogenic speciesare found in section Expansa. P. expansum producesrots in pomaceous fruits and P. italicum and P.ulaiense produce rot in citrus fruits. P. sclerotigenumproduces rot in yams and P. gladioli produces adestructive rot in Gladiolus corms. Species in seriesClaviformia are all coprophilic, creatine positive andsynnemata producing.Section DigitataSection Digitata (and series Digitata) is only representedby one species, P. digitatum. This species isunique in its combination of features. Conidiophoreand conidial structures are irregular and exceptionallylarge for Penicillium, biverticillate rather than terverticillate,divaricate and the conidia are olive-green.The conidia are large and ellipsoidal to cylindrical.The extrolites produced are tryptoquialanines, whichit only shares with P. aethiopicum from seriesAethiopica in section Chrysogena. The species growpoorly at low water activities and at higher temperatures,and it grows very poorly with no acid productionon creatine as sole N-source. It is also the onlyspecies in subgenus Penicillium that grow poorly onCzapek agar. The species has only been found onrotting citrus fruits. It shares the citrus rotting abilityand ellipsoidal to cylindroidal conidia with P. italicumand P. ulaiense from series Italica section Expansa,but shares no extrolites with those species. P.digitatum is the only species in subgenus Penicilliumthat cannot use nitrate as sole N-source.Section ViridicataMost species in section Viridicata have globoseconidia and rough-walled conidiophore stipes, withP. atramentosum as an exception with smooth-walledstipes. However occasionally section Viridicatamembers do not produce rough-walled stipes. Viridicataalso contain the only species with dark greenrough walled conidia in subgenus Penicillium. Mostspecies have a fasciculate colony texture and growrather fast, except species in series Verrucosa, whichgrow slowly. Several extrolites are only found insection Viridicata in subgenus Penicillium: Xanthomegnins,penicillic acids, puberulic acids, ochratoxins,daldinin C, alantrypinone, anacins, verrucins,auranthine, aurantiamin, puberuline, verrucosidin,terrestric acids, rugulovasines, asteltoxin, territrems,arisugacins, palitantin, compactins, barceloneic acid,and atrovenetins. Verrucolone (arabenoic acid) is13