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Evolutionary History and Taxonomy of Neotropical Mara Ioid Ferns

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Evolutionary History and Taxonomy of Neotropical Mara Ioid Ferns TURUN YLIOPISTON JULKAISUJA ANNALES UNIVERSITATIS TURKUENSIS SARJA - SER. AII OSA - TOM. 216 BIOLOGICA - GEOGRAPHICA - GEOLOGICA Evolutionary History and Taxonomy of Neotropical Mara�ioid Ferns: Studies of an Ancient Lineage of Plants by Maarten J. M. Christenhusz TURUN YLIOPISTO Turku 2007 From the Section of Biodiversity and Environmental Science, Department of Biology, University of Turku, Finland Supervised by Dr Hanna Tuomisto Section for Biodiversity and Environmental Science Department of Biology University of Turku, Finland Dr Soili Stenroos Botanical Museum Finnish Museum of Natural History University of Helsinki, Finland Reviewed by Dr Harald Schneider Department of Botany The Natural History Museum London, England, UK Dr Alan R. Smith University Herbarium University of California Berkeley, California, USA Examined by Dr Michael Kessler Alexander von Humboldt Institut Abteilung Systematische Botanik Universität Göttingen, Germany ISBN 978-951-29-3423-2 (PRINT) ISBN 978-951-29-3424-9 (PDF) ISSN 0082-6979 Painosalama Oy - Turku, Finland 2007 They arrived at an inconvenient time I was hiding in a room in my mind They made me look at myself, I saw it well I’d shut the people out of my life So now I take the opportunities Wonderful teachers ready to teach me I must work on my mind, for now I realise Every one of us has a heaven inside They open doorways that I thought were shut for good They read me Gurdjieff and Jesu They build up my body, break me emotionally It’s nearly killing me, but what a lovely feeling! I love the whirling of the dervishes I love the beauty of rare innocence You don’t need no crystal ball Don’t fall for a magic wand We humans got it all, we perform the miracles Them heavy people hit me in a soft spot Them heavy people help me Them heavy people hit me in a soft spot Rolling the ball, rolling the ball, rolling the ball to me Kate Bush The Kick Inside, 1978 This dissertation is based on the following studies, which are referred to by their Roman numerals in the text: I Christenhusz, M. J. M. 2006. Three new species of Danaea (Marattiaceae) from French Guiana and the Lesser Antilles. Annales Botanici Fennici 43: 212-219. II Christenhusz, M. J. M. and Tuomisto, H. 2006. Five new species of Danaea (Marattiaceae) from Peru and a new status for D. elliptica. Kew Bulletin 61: 17- 30. III Christenhusz, M. J. M. and Cremers, G., in review. Two new epitypes in Danaea (Marattiaceae, Pteridophyta) selected from original historical collections in Paris. Manuscript submitted to Candollea. IV Christenhusz, M. J. M., Tuomisto, H., Metzgar, J. and Pryer, K. M., in press. Evolutionary relationships within the Neotropical, eusporangiate fern genus Danaea (Marattiaceae). Molecular Phylogenetics and Evolution. V Christenhusz, M. J. M. and Toivonen, T. K., in review. Giants invading the tropics: The Oriental vessel fern, Angiopteris evecta (Marattiaceae). Manuscript submitted to Biological Invasions. Study I is reprinted with kind permission from the Finnish Zoological and Botanical Publishing Board. The Royal Botanic Gardens, Kew, kindly permitted the reprinting of study II. The manuscript of study IV is included with permission from Elsevier. CONTENTS 1. INTRODUCTION ..................................................................................................................7 1.1. IMPORTANCE AND HISTORY OF GIVING NAMES...................................................................7 1.2. FOCUS................................................................................................................................9 1.3. THE FAMILY MARATTIACEAE ..........................................................................................10 1.3.1. Description and geographic distribution ................................................................10 1.3.2. The genera Angiopteris, Archangiopteris and Macroglossum................................12 1.3.3. The genus Christensenia..........................................................................................13 1.3.4. The genus Danaea....................................................................................................14 1.3.5. The genus Marattia ..................................................................................................14 1.3.6. Conservation ...........................................................................................................15 1.4. PALAEOHISTORY OF MARATTIOID FERNS..........................................................................15 1.4.1. Palaeozoic Period (345-248 Ma) ............................................................................15 1.4.2. Mesozoic Period (248-65 Ma).................................................................................17 1.4.3. Cainozoic Period (65-1.8 Ma).................................................................................19 1.4.4. Evolutionary trends.................................................................................................19 1.5. AIMS ................................................................................................................................20 2. MATERIAL AND METHODS ...........................................................................................21 2.1. DATA COLLECTION...........................................................................................................21 2.1.1. Field studies ............................................................................................................21 2.1.2. Herbarium studies...................................................................................................23 2.2. MOLECULAR PHYLOGENETICS .........................................................................................23 2.2.1. Taxon sampling .......................................................................................................23 2.2.2. Laboratory work......................................................................................................24 2.2.3. Alignment and analyses...........................................................................................24 2.3. SPECIES DISTRIBUTION MODELLING .................................................................................25 3. RESULTS AND DISCUSSION ...........................................................................................26 3.1. STUDY OF THE GENUS ANGIOPTERIS .................................................................................26 3.1.1. Potential distribution modelling..............................................................................26 3.2. STUDY OF THE GENUS DANAEA.........................................................................................26 3.2.1. Species delineation and synonyms...........................................................................26 3.2.2. Taxonomic history ...................................................................................................29 3.2.3. Classification...........................................................................................................31 3.2.4. The Danaea leprieurii group....................................................................................31 3.2.5. The Danaea nodosa group .......................................................................................32 3.2.6. The Danaea alata group...........................................................................................34 3.2.7. Species diversity ......................................................................................................35 3.2.8. New Danaea species ................................................................................................38 3.2.9. Species distribution patterns ...................................................................................39 3.2.10. Ecology and speciation .........................................................................................41 3.3. STUDY OF THE GENUS MARATTIA ......................................................................................42 3.3.1. Redefined species and new synonyms......................................................................42 3.3.2. Taxonomic history ...................................................................................................43 3.4. DIAGNOSTIC KEYS TO NEOTROPICAL MARATTIACEAE.....................................................45 3.8.1. Key to genera of Neotropical Marattiaceae............................................................45 3.8.2. Key to species of Danaea .........................................................................................45 3.8.3. Key to Neotropical species of Marattia....................................................................50 4. CONCLUSIONS ...................................................................................................................51 5. ACKNOWLEDGEMENTS .................................................................................................53 6. REFERENCES .....................................................................................................................55 APPENDIX: TYPE SPECIMENS OF NEOTROPICAL MARATTIACEAE ...................60 ORIGINAL PUBLICATIONS ................................................................................................77 Introduction 7 1. INTRODUCTION 1.1. Importance and history of giving names ‘Is it you, Moonchild?’ Bastian asked. She laughed in a strangely lilting way. ‘Who else would I
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