2.9 Mannia Triandra (SCOP.) GROLLE Code: 1379 Anhang: II

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2.9 Mannia Triandra (SCOP.) GROLLE Code: 1379 Anhang: II 2.9 Mannia triandra (SCOP.) GROLLE Code: 1379 Anhang: II MONIKA HACHTEL, GERHARD LUDWIG & KLAUS WEDDELING, Bonn Namen: D: Dreimänniges Zwerglungenmoos, Dreimännige Mannie E: Mushroom-headed Liverwort F: – Systematik/Taxonomie: Marchantiophyta, Marchantiopsida, Marchantiales, Aytoniaceae. Synonyme: Mannia rupestris (NEES) FRYE & CLARK, Grimaldia rupestris (NEES) LINDENB., Neesiella rupestris (NEES) SCHIFFN., Duvalia rupestris NEES (s. auch GROLLE 1975). Kennzeichen/Artbestimmung: (entnommen aus FRYE & CLARK 1937–1947, MÜLLER 1954–1957, SILLER 1979: 139, SCHUSTER 1992) Das thallöse Lebermoos besitzt einen zarten, herzförmig dichotom verzweigten Thallus, der jung auf der Oberseite rein grün und auf der Unterseite bläulich gefärbt ist (Abb. 2.6 und 2.8). Im Alter wird er stumpf gräulich. Der geruchlose Gametophyt, welcher oft kompakte Rosetten formt, ist deutlich gefeldert, besitzt einen flachen Rand und kleine, hellgrüne oder blassrote, vereinzelt stehende, drei- eckige Bauchschuppen. Die dorsalen Epidermiszellen sind 16–23 x 20–25 µm groß, die zerstreut vorhandenen Ölzellen sind blassbraun. Die Poren sind nur wenig erhöht und umgeben von 1–3 Ringen, die aus 6–10 Zellen gebildet werden. An den Randthalli der monözischen Art befinden sich die Antheridienstände in kuchenförmigen Scheiben. Die Archegonienstände stehen auf 1–2 cm langen, nahezu schuppenlosen Trägern und besitzen halbkugelige, stark warzige Köpfchen. Die spiraligen Elateren sind gelblich-braun, 8–10 µm breit und 200–240 µm lang. Die Sporen sind gelbbraun bis braun, stark granuliert und mit einem Durchmesser von 60–70 µm sehr groß. Von der verwandten Mannia fra- grans ist M. triandra durch Färbung und Form des Thallus und der Bauchschuppen leicht unterscheidbar. Der Thallus dieser Art wird nur wenige Monate alt und stirbt nach der Spo- renreife im April bis Mai ab (GAMS 1938, GROLLE 1975, HUBER 1998); nur an feuchten, beschatteten Stellen kann er sich bis in den Sommer hinein halten (GAUCKLER 1940). Die Chromosomenzahl wird von FRITSCH (1991) sowie SCHUSTER (1992) mit n=9, von SIL- LER (1979) mit n=8 angegeben. Abbildungen der Art finden sich bei SCHUSTER (1992, S. 210; Aerenchym), MÜLLER (1954–1957), SCHIFFNER (1909, S. 23, Habitus und mehrere Details), GAUCKLER (1940) und INOUE (1976). Areal/Verbreitung: Welt: Mannia triandra besitzt ein disjunkt circumpolares, subkontinental-subarktisch- subalpines Areal (SCHUSTER 1958, DÜLL 1983, CORTINI PEDROTTI & ALEFFI 1992, DÜLL et al. 1999). Die Art kommt in China (PIIPPO 1990), Japan (MÜLLER 1954–1957, SCHUSTER 1958, 1992, INOUE 1976), im borealen, temperaten bis südlichen Europa (SCHUSTER 1958), in Mittel- und Nordostasien bis zur Beringsee (IGNATOV & AFONINA 1992), Kanada (MÜLLER 1954–1957, NEUMAYR 1971), Grönland (GAUCKLER 1940, NEUMAYR 1971, GAMS 1938), Alaska (GAMS 1938, NEUMAYR 1971) und den USA (MÜLLER 1954–1957) vor. Dort zeigt die Art eine kontinuierliche, relativ weite Verbrei- tung von Vermont bis New York, Tennessee, Nord Carolina, Ohio, Illinois, Michigan, Wisconsin, Minnesota und Iowa, Arkansas bis im Süden Missouri, Südgrenze in den Südappalachen (SCHUSTER 1992, SCHUSTER 1958, Herbar MO). EU: In Europa gilt Mannia triandra als montanes bis subalpines Element. Sie ist in Süd- und Mitteleuropa weit verbreitet, allerdings auf Kalkgebiete beschränkt (SCHUSTER 1992). Ihr 274 Moose Verbreitung von Mannia triandra in Deutschland und in der EU (Kartengrundlage MEINUNGER & SCHRÖDER in Vorb.). Erläuterungen zur Erstellung der Karte s. SSYMANK et al. (2003) im gleichen Band. Datengrundlage s. ELLWANGER et al. (2004) in Band 2 dieses Werkes. 275 Arealschwerpunkt innerhalb Europas liegt in den Alpen, wo sie zumindest in den 50er Jah- ren noch als verbreitet galt (GAMS 1938, GAUCKLER 1940, MÜLLER 1954–1957, HUBER 1998, NEUMAYR 1971). GAMS (1938) gibt für die Alpen insgesamt 37 Lokalitäten an. Nach DÜLL (1983), RAEYMAEKERS (1990) und FREY et al. (1995) kommt die Art in den EU- Ländern Deutschland, Frankreich (HUSNOT 1922, GAUCKLER 1940, MÜLLER 1954–1957, SILLER 1979), Italien (CORTINI PEDROTTI & ALEFFI 1992, ALEFFI & SCHUMACKER 1995, NEUMAYR 1971), Österreich (MÜLLER 1954–1957, NEUMAYR 1971, SCHINNERL 1904), Polen (MÜLLER 1952, SILLER 1979) und Spanien (GUERRA 1982, CASAS et al. 1996, CASAS 1998) vor, daneben auch in Kroatien (DÜLL et al. 1999, MÜLLER 1954–1957), Montenegro (MÜLLER 1954–1957), der Schweiz (MEYLAN 1924, MÜLLER 1954–1957, NEUMAYR 1971, URMI 1991), in der Slowakei (DUDA & VÁNˇA 1974, JANOVICOVÁ et al. 1999), Bosnien- Herzegowina (DÜLL et al. 1999), Slowenien (GROLLE 1975, NEUMAYR 1971, DÜLL et al. 1999, SCHUSTER 1992), Tschechien (DUDA & VÁNˇA 1974, NEUMAYR 1971, VÁNˇA 1997) und Ungarn (SILLER 1979, ORBÁN & VAJDA 1983). Außerdem wurde die Art in Albanien, Bulgarien (GANEVA 1998, DÜLL et al. 1999), Rumänien (SILLER 1979, RAEYMAEKERS 1990) sowie der ehemaligen UdSSR (KONSTANTINOVA et al. 1992, AFONINA & DUDA 1993, KONSTANTINOVA & POTEMKIN 1996, SILLER 1979) gefunden. D: In Deutschland ist Mannia triandra sehr selten und wurde nur in Bayern und Thürin- gen nachgewiesen, wo sie auf die kalkreichen Mittelgebirge und die Alpen beschränkt ist (vgl. Verbreitungskarte und MÜLLER 1954–1957). Nach 1980 wurde die Art in Nordbay- ern noch aus 18 MTB-Quadranten gemeldet. Voralpengebiet: Die früheren Vorkommen an der Isar bei München (MÜLLER 1954–1957, NEUMAYR 1971, FAMILLER 1917) wurden aktuell bestätigt (schriftl. Mitt. Berg). Auch SCHINNERL (1904) gibt Vorkommen bei Pul- lach, Baierbrunn, Maria-Einsiedel sowie an der Münchener Isarleite an. Im Donautal bei Kelheim und Eichstätt wird Mannia von MÜLLER (1954–1957) und SILLER (1979) ange- geben. In der Fränkischen Alb wurde der schon durch FAMILLER (1917) und NEUMAYR (1971) bekannte Wuchsort von Mannia triandra im Weltenburger Donaudurchbruch von HUBER (1998) aktuell bestätigt. Die früheren Vorkommen an Trockenmauern entlang der Straße zwischen Etterzhausen und Penk sowie an Felswänden im Schwarzen Labertal bei Schönhofen (FAMILLER 1917, GAUCKLER 1940) stuft HUBER (1998) dagegen aufgrund erfolgloser Nachsuche als erloschen ein. GAUCKLER (1940) nennt als weitere Lokalitäten in der nördlichen Fränkischen Alb das Püttlach- und Wiesentgebiet, Altmühl- und Donau- tal sowie die Dolomitfelsen des Pegnitztales bei Velden. MÜLLER (1954–1957) und SIL- LER 1979 geben Funde bei Muggendorf (Typuslokalität) und Pottenstein an. Eine punkt- scharfe Verbreitungskarte von insgesamt 43 Vorkommen in der Fränkischen Alb liefert NEUMAYR (1971). Die Art wurde außerdem in Berchtesgarden gefunden (PAUL & SCHOE- NAU 1930, SCHINNERL 1904). Aus Thüringen existiert nur ein Nachweis am Bleiberg (MEINUNGER 1992, Meinunger & Schröder schriftl. Mitt., FRAHM & FREY 1992, LUD- WIG et al. 1996). Mannia triandra hat eine colline bis alpine Höhenverbreitung (DÜLL & MEINUNGER 1989, RAEYMAEKERS 1990, JANOVICOVÁ et al. 1999). In den Steiermärki- schen Alpen reicht die Art bis über die Baumgrenze auf bis zu 2600 m über NN (GAMS 1938, MÜLLER 1954–1957, NEUMAYR 1971, SCHUSTER 1992); in der ehemaligen Tsche- choslowakei wurde sie zwischen 550 und 1065 m über NN gefunden (DUDA & VÁNˇA 1974, JANOVICOVÁ et al. 1999). FAMILLER (1917) gibt das Lebermoos für Bayern von der collinen Region (400 m) bis zur oberen Alpenregion (2300 m über NN) an. Verantwortung Deutschlands: Da Mannia triandra holarktisch verbreitet ist und zumindest in Nordamerika nicht selten vorkommt, besitzt Europa vermutlich keine besondere Ver- antwortung für die weltweite Erhaltung der Art. Da etwa ein Drittel der europäischen Loka- 276 litäten und nahezu die Hälfte der Fundorte innerhalb der Staaten der EU in Deutschland liegen, besteht für die BRD eine starke Verantwortung für den Erhalt der Sippe in Europa (Mitt. Berg, Mitt. Philippi). In Anbetracht dessen, dass Kalkfelsen i. d. R. bereits geschützt und nur in geringem Maße gefährdet sind, wird die Dringlichkeit für den Schutz der Art als eher gering angesehen. Biologie/Ökologie: Mannia triandra besiedelt Humusdecken auf Dolomit- und Massen- kalkfelsen, basenreichen Sand- und Schiefergesteinen sowie kalkhaltige, seltener auch neutrale Böden in warmer Klimalage (SCHIFFNER 1909, GAUCKLER 1940, SILLER 1979, SCHUSTER 1992, SAUKEL & KÖCKINGER 1999). Sie bevorzugt schattige, feuchtere und mesotherme Fels- und Mauerspalten (Trockenmauern), frisch erodierte Steilhänge und sonstige frische Verwitterungsböden (GAMS 1938) mit pH-Werten zwischen 7 und 8, tole- riert aber auch etwas sonnigere, trockenere Verhältnisse (NEUMAYR 1971, GUERRA 1982, SCHUSTER 1992, HUBER 1998). SAUKEL & KÖCKINGER (1999) geben für Österreich Moose alpine Rasen, karbonat- und basenreiche Silikatgesteine als Wuchsorte an. In Nord- Amerika liegen ihre Wuchsorte in reichen Laubwäldern in geringer Meereshöhe mit relativ hohen Niederschlägen (SCHUSTER 1958, 1992). Mannia triandra gehört aufgrund ihrer Kurzlebigkeit, sehr großen Sporen und geringen asexuellen Vermehrung wohl zu den „annual shuttle species“ (DURING 1979, DIERSSEN 2001). Soziologie: GAMS (1938) nennt als häufigste Begleiter Funaria hygrometrica, Distichium capillaceum sowie Selaginella helvetica. Nach SCHUSTER (1992) ist sie mit Lophozia badensis, nach NEUMAYR (1971) mit Dicranella varia sowie mit Arten des Distichietum capillacei wie Myurella julacea, Tortella tortuosa und Distichium capilla- ceum (GRETER 1936) vergesellschaftet. In Spanien charakterisiert sie zusammen mit Targionia hypophylla und Plagiochasma rupestre das Plagiochasmo-Targonietum hypo- phylla des Grimaldion fragrantis (GUERRA 1982). DIERSSEN (2001) ordnet M. triandra folgenden Gefäßpflanzengesellschaften
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