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Systematics and Evolution of the Genus Pleurothallis R. Br

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Systematics and Evolution of the Genus Pleurothallis R. Br Systematics and evolution of the genus Pleurothallis R. Br. (Orchidaceae) in the Greater Antilles DISSERTATION zur Erlangung des akademischen Grades doctor rerum naturalium (Dr. rer. nat.) im Fach Biologie eingereicht an der Mathematisch-Naturwissenschaftlichen Fakultät I der Humboldt-Universität zu Berlin von Diplom-Biologe Hagen Stenzel geb. 05.10.1967 in Berlin Präsident der Humboldt-Universität zu Berlin Prof. Dr. J. Mlynek Dekan der Mathematisch-Naturwissenschaftlichen Fakultät I Prof. Dr. M. Linscheid Gutachter/in: 1. Prof. Dr. E. Köhler 2. HD Dr. H. Dietrich 3. Prof. Dr. J. Ackerman Tag der mündlichen Prüfung: 06.02.2004 Pleurothallis obliquipetala Acuña & Schweinf. Für Jakob und Julius, die nichts unversucht ließen, um das Zustandekommen dieser Arbeit zu verhindern. Zusammenfassung Die antillanische Flora ist eine der artenreichsten der Erde. Trotz jahrhundertelanger floristischer Forschung zeigen jüngere Studien, daß der Archipel noch immer weiße Flecken beherbergt. Das trifft besonders auf die Familie der Orchideen zu, deren letzte Bearbeitung für Cuba z.B. mehr als ein halbes Jahrhundert zurückliegt. Die vorliegende Arbeit basiert auf der lang ausstehenden Revision der Orchideengattung Pleurothallis R. Br. für die Flora de Cuba. Mittels weiterer morphologischer, palynologischer, molekulargenetischer, phytogeographischer und ökologischer Untersuchungen auch eines Florenteils der anderen Großen Antillen wird die Genese der antillanischen Pleurothallis-Flora rekonstruiert. Der Archipel umfaßt mehr als 70 Arten dieser Gattung, wobei die Zahlen auf den einzelnen Inseln sehr verschieden sind: Cuba besitzt 39, Jamaica 23, Hispaniola 40 und Puerto Rico 11 Spezies. Das Zentrum der Diversität liegt im montanen Dreieck Ost-Cuba – Jamaica – Hispaniola, einer Region, die 95 % der antillanischen Arten beherbergt, wovon 75% endemisch auf einer der Inseln sind. Da die meisten Arten entweder endemisch oder pankaribisch verbreitet sind, bleiben die floristischen Bezüge zwischen den Inseln und zu den kontinentalen Nachbargebieten nur schwach ausgeprägt. Immerhin lassen sich einige Verbindungen unter den Inseln der Großen Antillen und besonders zu Mittelamerika erkennen. Diese Affinitäten steigen von Ost nach West. Molekulargenetische und (mikro-)morphologische Daten als phylogeographisches Werkzeug zeichnen dieses Muster deutlicher. Danach lassen sich die antillanischen Arten hinsichtlich ihrer Genese in drei Gruppen einteilen. 25% der Arten ist pankaribisch verbreitet, wobei der Großteil der antillanischen Populationen vom mittelamerikanischen Festland stammt. Ebenfalls aus dieser Region stammen weitere 25%, die jedoch auf den Inseln neue Arten gebildet haben (Anagenese). Die verbleibenden 50% der großantillanischen Sippen sind autochthon und das Ergebnis adaptiver Radiation auf den Inseln. Diese intensive Kladogenese beschränkt sich auf drei Verwandtschaftskreise innerhalb der Gattung Pleurothallis in den Untergattungen Antilla Luer und Specklinia Lindl. Es stellte sich heraus, daß der überwiegende Anteil der Artbildungsprozesse allopatrischer Natur ist. Sympatrie konnte nur in einem einzigen Fall direkt belegt werden. Das Ergebnis der allopatrischen Speziation sind zwei Typen von Vikarianz, räumlich geographischer und geologischer. In Cuba sind überraschenderweise fast 80% der endemischen Arten an einen Gesteinstyp gebunden, überwiegend an Serpentin. West- Hispaniola, wo viele Schwesternarten cubanischer Sippen beheimatet sind, besteht fast ausschließlich aus Kalkstein. Geographische Vikarianz ist daher oft geologisch unterlegt, eine Bindung die für Epiphyten kaum vermutet wurde. Hinter der Geologie verbergen sich 1 jedoch eher Bestäuberareale und weniger physiologische Anpassung als limitierender Faktor. Eine Verfrachtung in Vegetation auf anderem petrologischen Untergrund scheint damit der Hauptauslöser für Artbildungen gewesen zu sein. Grundlage waren höchtwahrscheinlich individuenarme Gründerpopulationen die den Bedingungen eines founder events ausgesetzt waren. Neben den reichen geologischen Verhältnissen im Dreieck Ost-Cuba – Jamaica – Hispaniola wird die intensive Artbildung durch weitere spezifisch lokale Bedingungen unterstützt. Karibische Wirbelstürme dürften entlang der Hauptrouten für eine häufige Verfrachtung von Samen oder Pflanzen von Mittelamerika auf die Großen Antillen sowie zwischen den Inseln selber verantwortlich sein. Ein zweiter günstiger Umstand für erfolgreiche Migration innerhalb des Dreiecks besteht in der räumlichen Nähe der Inselgebirge und deren optimalen klimatischen Bedingungen für die Besiedlung durch mikrophytische Epiphyten. Molekulargenetische Daten lieferten weiterhin wertvolle Informationen in Bezug auf die beiden aktuell diskutierten Systeme der Pleurothallidinae, einer streng morphologischen (Luer) und einer fast ausschließlich auf DNA-Sequenzen (Pridgeon & Chase) basierenden Klassifikation. DNA-Sequenzen der cubanischen Arten stützen das neue System von Pridgeon & Chase weitestgehend, zeigen aber noch Widersprüche in einigen der neuen oder wiedererrichteten Taxa. Angesicht dessen, daß die karibische Florenregion leider nicht nur durch ihre Biodiversität zu den zehn globalen hot spots zählt, sondern auch durch die großflächige Zerstörung von Primärvegetation, war es auch ein Anliegen der vorliegenden Arbeit, ein erstes detailliertes Bild von Genese und Verbreitung antillanischer Orchideen zu vermitteln. Diese Daten können direkt für die Gestaltung und das Management von karibischen Schutzgebieten eingesetzt werden, da Orchideen in der Naturschutzpolitik einen hohen Argumentationswert besitzen. 2 Content 1 INTRODUCTION...................................................................................................................................5 1.1 SUBTRIBE PLEUROTHALLIDINAE – AN INTRODUCTION ............................................................................5 Distribution ...............................................................................................................................................5 Morphology ...............................................................................................................................................5 Taxonomy and Systematics........................................................................................................................7 1.2 PLEUROTHALLIS OF THE GREATER ANTILLES .........................................................................................8 1.3 BIOGEOGRAPHY OF THE ANTILLEAN ARC............................................................................................10 1.4 GOALS OF THIS STUDY .........................................................................................................................12 2 MATERIALS AND METHODS..........................................................................................................13 2.1 MATERIALS..........................................................................................................................................13 Collection of plant material.....................................................................................................................14 2.2 METHODS ............................................................................................................................................15 Descriptions and drawings......................................................................................................................15 Reproductive biology...............................................................................................................................15 Ecological and phytogeographical patterns............................................................................................15 Computing and representation of the data in diagrams..........................................................................18 Palynology...............................................................................................................................................19 Genetics...................................................................................................................................................23 Taxonomy ................................................................................................................................................26 General rules...........................................................................................................................................26 3 RESULTS...............................................................................................................................................27 3.1 MORPHOLOGY .....................................................................................................................................27 Key to the species. ...................................................................................................................................28 3.2 REPRODUCTIVE BIOLOGY.....................................................................................................................67 3.3 PALYNOLOGY ......................................................................................................................................68 Description of pollen morphology...........................................................................................................68 Atypical pollinia ......................................................................................................................................72
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