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Phylogenie Der Pilze 351-366 © Biologiezentrum Linz, Download Unter ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Denisia Jahr/Year: 2007 Band/Volume: 0020 Autor(en)/Author(s): Krisai-Greilhuber Irmgard Artikel/Article: Phylogenie der Pilze 351-366 © Biologiezentrum Linz, download unter www.biologiezentrum.at Phylogenie der Pilze I. K RI S A I -G RE IL H U BE R Abstract: Phylogeny of fungi. Progress in molecular biological research and consistent phylogenetic analyses increased the knowledge about fungal phylogeny enormously. In a short overview the up-to-date phylogeny and the current phylogenetic clas- sification as a result of the international AFTOL (assembling the fungal tree of life) project is presented. Key words: Phylogeny, mycota, fungi, mycological systematics. Einleitung Das Prinzip der Pilze sind unverzichtbar für das einwandfreie Funk- phylogenetische Systematik tionieren von Ökosystemen. Ihre Lebensweise als Sa- Innerhalb der Evolutionsbiologie ist die phylogene- protrophe hilft organische Substanzen abzubauen und tische Systematik (Kladistik) die Methode der Wahl zur zu mineralisieren. Als symbiontische Pilze sind sie ver- Feststellung der stammesgeschichtlichen Verwandt- antwortlich für bessere Wachstumsraten ihrer Wirtsor- schaft. In der modernen phylogenetischen Systematik ganismen (etwa Mykorrhizapilze) oder für bessere Ver- strebt man bekanntlich danach, monophyletische dauung bei Tieren. Neben diesen positiven Rollen sind Gruppen zu bilden und nichtmonophyletische Grup- sie als Parasiten wichtige negative Faktoren in Land- pen zu beseitigen, sobald die Datenlage das erlaubt. Da- und Forstwirtschaft, sowie der Medizin. Sie sind in wei- bei sind folgende Begriffe und Prinzipien wichtig (siehe terer vielfältiger Weise bedeutend für den Menschen: z. B. WILEY 1981): bei der Herstellung von Lebensmitteln, Milchproduk- ten und alkoholischen Getränken, Antibiotika, bei der Monophyla sind Gruppen von Arten, die die biotechnologischen Gewinnung von Zitronensäure, Stammart und alle Abkömmlinge umfassen. Das Merk- Vitamin C, sowie als Speisepilze, Giftpilze, Heilpilze malskriterium dafür ist das spezielle abgeleitete Merk- und psychotrope Pilze. mal (Synapomorphie), das bei der Stammart erstmals auftrat und im Originalzustand oder in abgewandelter Traditionellerweise basierte die Erforschung der Form nur bei dieser Gruppe vorkommt. Evolution der Pilzgruppen im Zeitablauf bisher auf ver- gleichender Morphologie, Zytologie, Zellwandchemie, Paraphyla sind Gruppen von Arten, die zwar eine Ultrastruktur und wegen der schlechten Verfügbarkeit Stammart und einige, aber nicht alle Abkömmlinge nur zu einem geringen Grad auf Fossilien (KIRK et al. umfassen. Solche Gruppen wurden von den Systemati- 2001). Die Zahl der weltweit existierenden Pilze kann kern auf Grund von ursprünglichen Merkmalen (Ple- aufgrund des immer noch mangelhaften Erforschungs- siomorphien) gebildet. Ihnen fehlt sozusagen ein abge- grades nur schwer extrapoliert werden. Eine konservati- leiteter Teil, der im System an anderer Stelle eingesetzt ve Schätzung geht von einem absoluten Minimum von wird. Die meisten Taxonomen erachten solche para- 712.000 weltweit existierenden Pilzarten aus (SCHMIT phyletische Gruppen nicht mehr für wünschenswert, & MUELLER 2007). Die tatsächliche Anzahl der Pilze weil die Merkmalsverteilung im System nicht korrekt dürfte jedoch weit höher liegen. Eine progressive Schät- wiedergegeben wird. zung liefert HAWKSWORTH (2001) mit bis zu 1,5 Millio- Polyphyla sind Gruppen von Arten, deren Stamm- nen! Von dieser Fülle an Arten sind bisher nur ca. art einer anderen Gruppe zugeordnet wird. Solche 80.000 Arten beschrieben (SCHMIT & MUELLER 2007). Gruppen wurden auf Grund von Merkmalskonvergen- In der zweiten Hälfte des 20. Jahrhunderts brachten zen (Ähnlichkeiten, die keinen gemeinsamen Ur- Denisia 20, kladistische und molekulare Studien große Fortschritte sprung haben) gebildet. Sie werden generell von allen zugleich Kataloge der oberösterreichischen in der Erforschung der Pilztaxa und im Verstehen der Systematikern als Taxa abgelehnt, sobald man die Poly- Landesmuseen Neue Serie 66 (2007): Verwandtschaft pilzlicher Stämme. phylie erkennt. 351-366 © Biologiezentrum Linz, download unter www.biologiezentrum.at ten die Entstehung der Chytridiomycota ungefähr vor 550 Millionen Jahren, der Abtrennung der Ascomyco- ta-Basidiomycota-Linie vor ca. 400 Millionen Jahren, viele Schlauchpilzhefen und Schimmelpilze entstanden nach dem Erscheinen der Angiospermen in den letzten 200 Millionen Jahren. Neue Fossilfunde und verbesserte analytische Me- thoden unterstützen die Entstehung aller existierenden Stämme im Devon. Paleopyrenomycites devonicus ist ein fossiler Schlauchpilz aus dem Unteren Devon (TAYLOR & BERBEE 2007). Prototaxites wird neuerdings aufgrund bestimmter Kohlenstoffisotope als fossiler heterotropher Organismus und angesichts der tubulären anatomischen Struktur als Pilz aus dem Unteren Devon interpretiert (BOYCE et al. 2007). Abb. 1: Bei der Lungenflechte, Lobaria pulmonaria, ist der Pilz der Die Phylogenie der Pilze in Standard- formgebende Teil, der sich mit Apothezien sexuell fortpflanzen kann, die Alge ist untergeordnet. Foto: F. Reinwald. werken und wichtiger Literatur Ob etwas ein abgeleitetes oder ein ursprüngliches Die im Standardwerk KIRK et al. (2001) präsentier- Merkmal ist, wird mit dem Außengruppenvergleich te Klassifikation der Stammesgeschichte listet pilzliche festgestellt: Hat man in der Innengruppe bei einem Organismen in drei Reichen (Fungi, Chromista, Proto- Merkmalstyp zwei Zustände, so ist derjenige der ur- zoa) auf. Dabei sind Animalia und Fungi Schwester- sprüngliche, der auch außerhalb bei den nächsten Ver- gruppen die von choanozoen-ähnlichen Vorfahren in- wandten vorkommt. Man muss also schon ein gewisses nerhalb der Protozoen abstammen. Für die früher disku- Vorwissen besitzen, was die Innengruppe und die Au- tierte Hypothese der Abstammung der Asco- und Basi- ßengruppe betrifft. Die Polarisierung der Merkmale (Be- diomycota von den Rotalgen wurde keine molekulare stimmung, was ist ursprünglich oder abgeleitet) wird au- Bestätigung gefunden. Pilzähnliche Organismen in den tomatisch von der verwendeten Software durchgeführt, beiden anderen Reichen (Chromista, Protozoa) sind po- sobald die Innen- und Außengruppe festgelegt sind. lyphyletisch entstanden. Innerhalb der Protozoa sind die pilzähnlichen Oganismen alle Schleimpilzgruppen, Mit den Methoden der DNA-Systematik wurde es die jedoch untereinander nicht näher verwandt sind. möglich, stammesgeschichtliche Zusammenhänge we- Die Acrasidae gehören zu den amöboiden flagellaten sentlich klarer zu erkennen als früher, da ungleich mehr Protozoen. Die echten Schleimpilze, Protosteliden und Merkmale zur Verfügung stehen. Dictyosteliden fallen in ein Kluster mit den Rhizopoda, Plasmodiophoride stehen bei den amöboiden, photo- Was sind Pilze? synthetischen Chlorarachniden und Cercomonaden. Innerhalb des Reiches Fungi (Echte Pilze) unterstützen Pilze können nicht mehr einfach als chlorophyll- molekulare Daten die Monophylie von Asco- und Basi- freie, heterotrophe, thallische Eukaryoten mit Zellwand diomycota. Innerhalb der Ascomycota nimmt Neolecta zumindest in einem Lebensstadium (MÜLLER & LÖFFLER eine basale Position ein, gemeinsam mit Pneumocystis, 1992) definiert werden. Das wäre keine monophyleti- Schizosaccharomyces und den Taphrinales, angeführt als sche Gruppierung. Die Echten Pilze, Fungi, können Archiascomycetes. Chytridiomycetes und Zygomycetes heute so abgegrenzt werden: absorbtive heterotrophe sind polyphyletisch. Die Glomales erscheinen basal zu Eukaryoten mit Sporen mit Chitin, ohne Plastiden, oh- den Asco- und Basidiomycetenlinien. Das steht augen- ne Phagotrophie und tubuläre Cilien, mit Lysinsynthese scheinlich im Zusammenhang mit ihrer Rolle als My- über den AAA-Pfad und mit Mitochondrien mit fla- korrhizapilze assoziiert mit den ersten Landpflanzen, be- chen Cristae (CAVALIER-SMITH 2001). legt durch Funde von fossilen Glomales im Ordovicium. Andere Zygomycetengruppen bilden separate Linien Fossilbefunde und implizieren dadurch, dass Geißeln unabhängig von- und evolutionäre Radiation einander mehrmals verloren wurden und auch Zygospo- ren mehrmals entstanden sind (KIRK et al. 2001). Die Substitutionsrate in DNA-Sequenzen ist ein Maß für den Zeitraum der evolutionären Ausbreitung Die überwiegend autotrophen Chromista (auch (KIRK et al. 2001). BERBEE & TAYLOR (1993) berechne- Stramenopila, Heterokonta) enthalten zwei separate he- 352 © Biologiezentrum Linz, download unter www.biologiezentrum.at a b c d Abb. 2: Aeroaquatische Hyphomyceten. (a) Helicodendron tubulosum, (b) Helicodendron luteoalbum (c) Helicodendron sessile (d) Clathrosphaerina zalewskii. Diese mitosporen Pilze, die Konidien ausbilden, waren früher bei der künstlichen Gruppe der Fungi imperfecti, aufgrund molekularer Daten können sie heute den Schlauchpilzen zugeordnet werden (Voglmayr, pers. Mitt.). Foto: H. Voglmayr. terotrophe Linien mit pilzlichen Organismen, die beide schen einem Pilz und einer oder mehreren Grünalgen von einem ancestralen heterokonten Flagellaten mit bzw. Cyanobakterien. Da sich nur der Pilz sexuell fort- Chloroplasten abstammen. Die eine Linie umfasst die pflanzen kann und auch meist formgebend ist, werden Oomyceten und Hyphochytridien gemeinsam mit dem sie zu den Pilzen gestellt, sie sind großteils Schlauchpil- marinen Flagellaten Developayella, die andere Linie die ze (Abb. 1). Labyrinthulomycota (Schleimnetze) und Thraustochyt-
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