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“Diversity and Evolution of Pteridophytes, with Emphasis on The “Diversity and evolution of pteridophytes, with emphasis on the Neotropics” Dissertation zur Erlangung des Doktorgrades der Mathematisch-Naturwissenschaftlichen Fakultäten der Georg-August-Universität zu Göttingen vorgelegt von Marcus Lehnert aus Neustadt/Ostholstein Göttingen 2007 D7 Referent: Prof. AP Dr. Michael Kessler Korreferent: Prof. Dr. S. Robbert Gradstein Tag der mündlichen Prüfung: 04.07.2007 Marcus Lehnert: Diversity and evolution of pteridophytes TABLE OF CONTENTS Acknowledgements. p. i Disclaimer. p. ii Summary. p. iv Zusammenfassung. p. v General introduction & outline. p. 1 Chapter I. Ferns as species: systematics and phylogeny. p. 3 I.1. Novelties of Cyatheaceae from the Neotropics. p. 7 I.2. On the identification of Cyathea pallescens (Sodiro) Domin (Cyatheaceae): the truth and its consequences. p. 69 I.3. Revision of the fern genus Melpomene (Polypodiaceae). p. 110 I.4. Phylogeny of the fern genus Melpomene (Polypodiaceae) inferred from morphology and chloroplast DNA analysis. p. 327 Chapter II. Ferns as populations: floristics and biogeography p. 356 II.1. The tree ferns of Bolivia. p. 362 II.2. Checklist to the pteridophytes of the Reserva Biológica San Francisco (Ecuador, Prov. Zamora-Chinchipe). p. 363 II.3. Are ridge habitats special sites for endemic plants in tropical montane rain forests? A case study of pteridophytes in Ecuador. p. 378 II.4. Comparative analysis of vegetation zonation in four neotropical fern communities. p. 395 Marcus Lehnert: Diversity and evolution of pteridophytes Chapter III. Ferns in communities: ecology. p. 413 III.1. The influence of soil evolution on pteridophyte diversification. p. 417 III.2. New insights on the mycorrhizal infections in ferns: an example from southern Ecuador. p. 437 III.3. Mycorrhizae and pteridophytes – a global phylogenetic perspective. p. 450 Appendix: Lebenslauf Marcus Lehnert: Diversity and evolution of pteridophytes ACKNOWLEDGEMENTS Firstly, I thank Michael Kessler for the great supervision of this thesis. I certainly would not have come so far without his constant encouragement and scientific guidance. I also thank Rob Gradstein for sharing his knowledge of cryptogams and providing the facilities to carry out my studies on ferns. I also want to thank all my friends at the Dept. Systematic Botany of the A.-v.-Haller-Institut, Göttingen: Jochen Heinrichs, Harald Schneider, Doris Bär-Scheubel, Sybille Hourticolon, Brigitte Warnecke, Ana Moguel, Thomas Janßen, Hans Peter Kreier, Bastian Steudel, and Simone Goda Sporn. Unforgettable to me are Klaus Lewejohann and Ursula Hofmann†. During my lengthy travels, I found many new friends, whose help I want to acknowledge here: Robbin Moran, John Mickel, and Michael Sundue from the New York Botanical Garden; Tom Ranker, Genie Trapp, and Susan Klimas from Boulder, Colorado; the members of the Herbarium UC, the Mishler-Lab, and the Baldwin-Lab at the University of California, Berkeley, especially Brent Mishler, Andy Murdock, Eric Harris, Bianca Knoll, Rebecca Guenther, Ruth Kirkpatrick, Laine Huiet, John Strother, and Dan Norris; in Ecuador, the staff of the herbarium Loja and at the herbarium QCA, especially Hugo Navarrete; from Peru, Ascunción Cano and Blanca Leon; the staff of the Herbarium LPB, especially Cris Roberts, Edgar Gudiño, and Stephan Beck; Paulo Labiak and Jefferson “It’s-alright-you’re-in- Brazil-now” Prado. I am deeply indebted to Alan and Joan Smith, having experienced their hospitality and kindness. I recall the fieldwork with my colleagues Alexander Schmidt- Lebuhn and Nicki Mandl with lasting and often fond memories. Financial support by the DAAD, DFG, EAP, and BIOPAT is greatly acknowledged. A special thanks goes to my parents, Norbert and Bärbel Lehnert, without whose early cooperation this thesis would not have been possible. i Marcus Lehnert: Diversity and evolution of pteridophytes DISCLAIMER The chapters of this thesis are designed as manuscripts suitable for direct submission. If a decision has already been made, formatting follows the guidelines of the respective journal in which the publication is planned. All new species names and combinations in this thesis are considered as provisional. They will be validly published later in peer-reviewed journals. Contribution of the doctorate candidate Marcus Lehnert to the chapters presented in this thesis: I.1. Novelties of Cyatheaceae from the Neotropics. Concept: 100% Gathering of data: 100% Evaluation: 100% Writing: 100% I.2. On the identification of Cyathea pallescens (Sodiro) Domin (Cyatheaceae): the truth and its consequences. Concept: 100% Gathering of data: 100% Evaluation: 100% Writing: 100% I.3. Revision of the fern genus Melpomene (Polypodiaceae). Concept: 100% Gathering of data: 100% Evaluation: 100% Writing: 100% I.4. Phylogeny of the fern genus Melpomene (Polypodiaceae) inferred from morphology and chloroplast DNA analysis. Concept: 100% Gathering of data: 100% Evaluation: 75% Writing: 100% II.1. The tree ferns of Bolivia. Concept: 100% Gathering of data: 100% Evaluation: 100% Writing: 100% II.2. Checklist to the pteridophytes of the Reserva Biológica San Francisco (Ecuador, Prov. Zamora-Chinchipe). Concept: 10% Gathering of data: 90% ii Marcus Lehnert: Diversity and evolution of pteridophytes Evaluation: 100% Writing: 100% II.3. Are ridge habitats special sites for endemic plants in tropical montane rain forests? A case study of pteridophytes in Ecuador. Concept: 0% Gathering of data: 100% Evaluation: 0% Writing: 10% II.4. Comparative analysis of vegetation zonation in four neotropical fern communities. Concept: 50% Gathering of data: 100% Evaluation: 75% Writing: 100% III.1. The influence of soil evolution on pteridophyte diversification. Concept: 30% Gathering of data: 30% Evaluation: 50% Writing: 100% III.2. New insights on the mycorrhizal infections in ferns: an example from southern Ecuador. Concept: 50% Gathering of data: 100% Evaluation: 75% Writing: 100% III.3. Mycorrhizae and pteridophytes – a global phylogenetic perspective. Concept: 50% Gathering of data: 100% Evaluation: 75% Writing: 100% iii Marcus Lehnert: Diversity and evolution of pteridophytes Summary Pteridophytes, understood as taxonomic group containing ferns, horsetails, and clubmosses, are especially diverse and abundant in the tropical mountain ranges of South America. My work aims to understand better the diversity of these plants, how they have evolved, and how they function in the ecosystems. Tree ferns are conspicuous elements of tropical montane rainforest. About 200 species of scaly tree ferns of the family Cyatheaceae occur in the Neotropics, ranging from small trunkless ferns of rocky outcrops to almost 20 m tall giants in dense forests. My work on these plants led to the recognition of 19 new and several formerly synonymized species in the genera Alsophila and Cyathea. The confusing nomenclature of the family was partially clarified by the correction of the typification of Cyathea pallescens (Sodiro) Domin. A checklist with keys to the Bolivian tree ferns was also accomplished during these studies. The revision of the genus Melpomene revealed 27 species with 10 varieties, most of them small ferns with deeply pinnatifid fronds, which are mostly epiphytes in upper montane forests or characteristic terrestrial elements of treeless páramo vegetation. The phylogenetic analysis based on morphology and chloroplast DNA shows that this genus as currently morphologically circumscribed is monophyletic and originated in South America. The radiation of the core group is apparently directly connected with the uplift of the northern Andes, which is the center of diversity of this genus. My ecologically orientated field studies of fern communities in southern Ecuador initiated several projects that aimed on different aspects. I found 248 different species of pteridophytes in our main study area, the Reserva Biológica San Francisco (RBSF), Prov. Zamora-Chinchipe, but this number is likely to increase in the future. The study area is part of the Amotape-Huancabamba zone, a stretch of low elevation in the Andes located a the overlap of several biogeographic subunits and thus rich in endemic and widespread species alike. I found that the upper limit in the elevational distribution of most of the widespread ferns follows this dent in the mountain range, indicating that probably a downward shift of all vegetation belts may be found. The mountain ridges in the RBSF support a unique heath forest dominated by the otherwise rare tree Purdiaea nutans Planch., but these peculiarities are not reflected in the fern composition. Ridge habitats in the study area, including two comparative sites close to the RBSF, are less diverse than adjacent slopes, and there was no higher representation of localized species on ridges. Overall, widespread species were weakly but significantly more iv Marcus Lehnert: Diversity and evolution of pteridophytes abundant than localized species, and terrestrial – but not epiphytic – species were more abundant on ridges compared to slopes. The observed influence of soils on the distribution of ferns in the Ecuadorian study area, where terrestrial and phylogenetically more derived taxa increase in diversity along a nutrient gradient caused by a downhill flux from the ridges to the gorges, encouraged me to look at the soil preferences of pteridophytes worldwide and see if it contains a phylogenetic signal. It seems that more derived lineages are better represented on rich soils, but have also a higher percentage of epiphytes. Looking at the
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