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Moosbewohnende Ascomyceten I. Die © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.biologiezentrum.at Mitt. Bot. München 14 p. 1 - 360 ].6. 1978 ISSN 0006-8179 MOOSBEWOHNENDE ASCOMYCETEN I. DIE PYRENOCARPEN, DEN GAMETOPHYTEN BESIEDELNDEN ARTEN von P. DÖBBELER INHALTSVERZEICHNIS Seite EINLEITUNG 4 A. ALLGEMEINER TEIL 7 I. Technische Hinweise 7 1. Material und Methode 7 2. Zur Suche von Moospilzen 8 3. Zu den Artbeschreibungen, der Nomenklatur der Wirte und den Fundortsangaben 9 4. Verwendete Zeichen und Abkürzungen 10 II. Diskussion einiger Merkmale 11 1. Borsten und Tüpfel 11 2. Jodfärbung der Hymenialgallerte 12 3. Zum Verlauf der Hyphen 14 4. Appressorien und Haustorien 15 Vom Fachbereich Biologie der Universität München als Dissertation angenommen. © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/;- 2 www.biologiezentrum.at Seite. III. Biologie der Moospilze 16 1. Saprophyten und Parasiten 16 2. Anpassungen an das Leben auf Moosen 18 3. Orte der Fruchtkörperbildung 19 4. Die Lamellenzwischenräume von Polytrichales bewohnenden Arten 20 5. Arten, deren Fruchtkörper die Wirtsblätter durch- bohren 22 6. Dem Substrat eingesenkte Arten 23 7. Fruktifikationszeit 24 8. Zum Problem der Lichenisierung 25 9. Gallenbildungen 27 IV. Wirtsspezifität und die am häufigsten nachgewiesenen Wirte 28 1. Wirtswahl 28 2. Die am häufigsten befallenen Moose 32 V. Häufige und seltene Arten 33 VI. Geographische Verbreitung 34 VII. Die Pilzflora der Sporophyten 36 VIII. Zum Abbau der Moose 38 B. SPEZIELLER TEIL 39 I. Zur Gattungsgliederung 39 II. Bestimmungsschlüssel 41 1. Schlüssel der Gattungen 41 2. Hilfsschlüssel 45 a. Schlüssel der Arten auf Polytrichales 45 b. Schlüssel der Arten auf Grimmiaceen 49 c. Schlüssel der Arten auf Leskeaceen 51 d. Schlüssel der Arten auf Pottiaceen 52 e. Schlüssel der Arten auf Hylocomium 54 f. Schlüssel der Arten auf Plagiochila asplenioi- des s. 1 54 g. Schlüssel der Arten auf Porella sp. div 55 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.biologiezentrum.at Seite III. Beschreibungen der Gattungen und Arten 57 Sphaeriales 57 Hypocreaceae 57 1. Calonectria 57 2. Nectria 70 3. Pseudonectria 96 4. Thyronectria (inconspicua) 116 5. Trichonectria 118 Amphisphaeriaceae 124 6. Monographella (abscondita) 124 Dothideales 126 Pleosporaceae 126 7. Bryopelta (variabilis) 126 8. Bryorella 128 9. Bryosphaeria 151 10. Bryostroma 170 11. Didymosphaeria (marchantiae) 187 12. Julella 190 13. Massarina 196 14. Punctillum 201 15. Racovitziella (endostromatica) 206 Dimeriaceae 208 16. Bryochiton 208 17. Bryomyces 233 18. Epibryon 260 19. Lizonia 305 20. Myxophora (amerospora) 315 Pseudosphaeriaceae 317 21. Monascostroma (sphagnophilum) 317 IV. Species incertae sedis 319 V. Die nicht berücksichtigten Arten 324 ZUSAMMENFASSUNG 328 SUMMARY 330 VERZEICHNIS DER NEUEN UND UMKOMBINIERTEN SIPPEN 333 INDEX NOMINUM 335 WIRTSVERZEICHNIS 339 LITERATURVERZEICHNIS 351 - - - 4 - © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.biologiezentrum.at EINLEITUNG Die Fähigkeit der Pilze, verschiedenste Thallo- oder Kormophy- ten zu besiedeln, ist seit langem gut bekannt. Für Moosbewohner lie- gen allerdings kaum Hinweise vor, obwohl die weite ökologische Am- plitude, der Arten- und insbesondere Individuenreichtum und das hohe phylogenetische Alter der Bryophyten das Wachstum pilzlicher Para- siten oder Saprophyten erwarten lassen. Außerdem unterliegen Moose Abbauprozessen, an denen nicht nur Bakterien beteiligt sind. Die Beschreibungen moosbewohnender Pilze (gleichbedeutend mit bryophilen Pilzen) beruhen gewöhnlich auf zufälligen Einzelfunden, die in der mykologischen und bryologischen Literatur weit zerstreut sind. Auch große Herbarien verfügen nur über wenige oder gar keine Belege. Wer die Kapitel über Beziehungen zwischen Moosen und ande- ren Organismen etwa bei NICOLAS (1932), K. MÜLLER (1954) oder SCHUSTER (1966) studiert, gewinnt den Eindruck, bryophile Pilze seien abgesehen von Mykorrhiza-Partnern eine Seltenheit. Die Arbei- ten über parasitische Pilze und ihre Wirte in "The Fungi" (AINS WORTH & SUSSMAN 1968) erwähnen nicht einmal Moose als Substrat. Tatsächlich gehören aber manche bryophile Fungi Imperfecti, Discomyceten, unitunicate und vor allem bitunicate Pyrenomyceten zu häufigen Vertretern unserer Mykoflora. Manche Moose sind gebiets- weise eher mit als ohne Pilze anzutreffen. Es scheint sogar wahr- scheinlich, daß nicht nur sämtliche Moose als Wirte in Frage kommen können, sondern ein großer Teil aller Moosrasen befallen ist. Dem- entsprechend liegt eine Vielzahl solcher Proben unerkannt in den bryologischen Herbarien. Die geringe Beachtung, die Moosbewohner auch etwa im Vergleich zu Flechtenparasiten fanden, mag teils auf ihre außerordentliche Klein- heit und verborgene Lebensweise zurückzuführen sein, teils aber auf die wenigen Berührungspunkte zwischen Mykologie und Bryologie. Trotzdem ist kaum verständlich, daß aus England zum Beispiel, des- sen Moosflora schon um die Jahrhundertwende gut erforscht war, bis- her lediglich drei den Gametophyten bewohnende Pyrenomyceten und sie nur wenige Male gesammelt wurden. Ein geschichtlicher Überblick fällt kurz aus: NEES von ESEN- BECK beschrieb 1830 unter dem Namen Sphaeria junge r man niae als erster einen pyrenocarpen Ascomyceten von der haploiden Moosgeneration. 1863 kam die von AUERSWALD in den männlichen Gametangienhüllen von Polytrichum commune gesammelte Lizonia emperigonia in dem Exsiccatenwerk von KLOTZSCH- RABENHORST zur Verteilung. Lizonia und die später von PI- ROTTA aufgestellte, nah verwandte Pseudolizonia baldinii - © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.biologiezentrum.at galten jahrzehntelang als die Moospilze schlechthin. Beide Arten wur- den in Kryptogamenfloren wiederholt dargestellt und mehrmals in Ex- siccatenwerken ausgegeben. Es darf nicht wundern, daß diese Pilze von allen Moosbewohnern am besten in den Herbarien belegt sind. Be- zeichnenderweise wurden auch zwei Species dem Genus Lizonia zugerechnet (L. hypnorum und L. sphagni), die mit ihm nichts weiter gemeinsam haben als ihr Vorkommen auf Moosen. Die Gebrüder CROUAN veröffentlichten 1867 unter anderem eine Reihe von Moospilzen aus Finist^re im Nordwesten Frankreichs. Außer ihnen scheint der spanische Bryologe CASARES besonders auf sie geachtet zu haben. Seine Funde wurden im wesentlichen von GON- ZALEZ FRAGOSO zwischen 1914 und 1926 in mehreren Arbeiten vor- gestellt. 1959 faßte RACOVITZA unter Auswertung umfangreicher Litera- tur die bis dahin bekannt gewordenen pyrenocarpen Ascomyceten und Fungi Imperfecti in einer monographischen Bearbeitung zusammen, in der auch die Sporogonbewohner Berücksichtigung fanden. Etwa achtzig Pyrenomyceten (darunter dreiunddreißig neue Sippen) und hundertacht undzwanzig Fungi Imperfecti (neunzehn neue Arten) wurden bezüglich Biologie und systematischer Stellung diskutiert und weitgehend ausführ- lich beschrieben und abgebildet. Für andere Höhere Pilze und für Phy- comyceten fehlt eine zusammenfassende Darstellung. Vorliegende Untersuchung wurde mit dem Ziel begonnen, die bei RACOVITZA aufgeführten Arten wiederzufinden und seine Angaben über Verbreitung und Wirtswahl zu ergänzen. Dazu war die Durchsicht vieler Moosrasen erforderlich, denn nur in wenigen Fällen konnte auf Herbarmaterial zurückgegriffen werden. Es zeigte sich bald, daß zahl- reiche Sippen neubeschrieben oder umkombiniert werden mußten. Des weiteren wurde eine Beschränkung auf die pyrenocarpen, den Gameto- phyten besiedelnden Arten nötig. Lediglich der allgemeine Teil nimmt auch auf andere Moosbewohner Bezug. Die Verwandtschaft von Lasiosphaeria und Leptonaeli- o 1 a und die bisher den Genera Pleosphaeria, Pleospora und Teichospora zugeordneten Vertreter konnten nicht berück- sichtigt werden, da der Umfang der Arbeit sonst zu sehr angeschwol- len wäre. Diese Arten sind aber mit der wichtigsten Literatur und den Wirtsangaben aufgeführt. Alle übrigen Sippen wurden so weit wie möglich an Hand eigener Aufsammlungen beschrieben. Die Fundorte und die Literatur sind mit Ausnahme bloßer Angaben in Wirtsverzeichnissen (vgl. zum Beispiel OUDEMANS 1919, RACOVITZA 1959. SYDOW in SACCARDO 1898, SEYMOUR 1929; Index of Fungi) vollständig zitiert. © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/;- 6 www.biologiezentrum.at Die Ergebnisse werden nicht ohne Zögern vorgelegt. Allzu oft sind sie durch Zufälligkeiten und Vermutungen belastet. Wenn es dennoch geschieht, so vor allem, um die Aufmerksamkeit anderer auf bryophile Pilze zu lenken. Moosbewohner scheinen zur Lösung mancher Frage- stellungen besonders geeignet zu sein, da vergleichsweise einfach ge- baute, uralte Organismen verschiedenster VerwEuidtschaftskreise in Verbindung treten, deren Kenntnis reizvolle Einsichten in das Gesche- hen in der Natur bietet. Meinem verehrten Lehrer Herrn Professor Dr. J. POELT (Graz), von dem ich zum ersten Mal das Wort "Moospilz" hörte, gilt besonders herzlicher Dank. Ohne seine ständige und großzügige Hilfe wäre vor- liegende Arbeit nicht zustande gekommen. Unvergessen bleiben begei- sternde Exkursionen und die Einführung in die Kryptogamenkunde. Sei- nen Namen trägt die auf Seite 203 beschriebene Art. Herrn Professor Dr. H. MERXMÜLLER (München) schulde ich aufrichtigen Dank für wertvolle, kritische Anmerkungen. Er ermög- lichte es, die in Graz begonnenen Untersuchungen an seinem Institut zum Abschluß zu bringen. Bei Herrn Professor Dr. H. HERTEL (München) bedanke ich mich herzlich für jederzeit gewährte, freundschaftliche Hilfe
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