Zurich Open Repository and Archive University of Zurich Main Library Strickhofstrasse 39 CH-8057 Zurich www.zora.uzh.ch Year: 2007 Macro-evolutionary and macro-ecological studies on the Cape flora, with focus on the Pentaschistis clade (Poaceae) Galley, Chloé A Abstract: Diese Arbeit behandelt die Makroevolution und die Makroökologie einer artenreichen Gruppe von Grasen der Kapflora (der Pentaschistis „Clade“), und anderer „Clades“, welche dieselbe Verbre- itung aufweisen. Im ersten Kapitel untersuche ich die Evolution und Funktion von mehrzelligen Drüsen und von anatomischen Blatttypen in der Pentaschistis Gruppe. Um Merkmalsevolution zu untersuchen, benötigt man eine phylogenetische Hypothese. Schliesslich werden einige unklare Punkte betreffend zwischenartlichen Beziehungen aufgelöst. Prionanthium ist innerhalb Pentaschistis anzusiedeln und dieser „Clade“ kann als Schwestergruppe zu Pentameris und Pentaschistis tysonii betrachtet werden. Im zweiten Kapitel untersuche ich den Artenreichtum des Pentaschistis „Clade“ in der Kapflora, mit der Drakensberg Region als Vergleich. Die Habitatsdiversität der Landstriche (zwischen Regionen) und die Habitatsunterschiede zwischen unterschiedlichen Landstrichen (innerhalb der Regionen) werden ver- glichen. Diese Resultate werden also Vorhersage von Artreichtum und Artenfluktuation für die jeweiligen Regionen getestet. Trotz einer gut untersuchten Flora werden immer noch neue Arten von der Region der Kapflora (CFR) entdeckt und beschrieben. Kapitel drei ist eine Beschreibung von dreier neuerArten der CFR. Kapitel vier ist eine Metaanalyse die sich auf die Ursprünge der „Cape Clades“ konzentri- ert, welche ca. 50% der am Kap vorkommenden Pflanzenarten ausmachen. Die „Cape Clades“ haben räumlich und zeitlich sehr unterschiedliche Ursprünge. Kapitel fünf untersucht die Wanderrouten von vier Linien, welche hauptsächlich am Kap beheimatet sind, durch die Regionen des gemässigten Afrikas. Ein generelles Migrationsmuster um die Region des gemässigten Afrikas wird identifiziert. Zusätzlich wird das Ausmass lokaler Diversifikation in verschiedenen Regionen verglichen. Abstract This thesis is a macro-evolutionary and macro-ecological study of a species rich group of grasses from the Cape flora (the Pentaschistis clade), and other clades which share its distribution. In chapter one I investigate the evolution and function of multicellular glands and leaf anatomical types in the Pentaschistis clade, using a phylogenetic approach to understand how often, and under which circumstances glands were gained, maintained or lost. Some issues regarding inter-generic relationships are resolved. Prionanthium is nested within Pentaschistis and this clade is sister to Pentameris plus Pentaschistis tysonii. In chapter two I investigate the species richness of the Pentaschistis clade in the Cape Floristic Region (Cape), using the Drakensberg region as a comparison. The habitat heterogeneity of landscapes and turnover between landscapes are compared between the Cape and the Drakensberg, and also tested as predictors of species richness and turnover in each area. Chapter three is the description of three new species from the CFR; despite being a well collected flora, the discovery and description of new species from the CFR arecon- tinuing. Chapter four is a meta analysis focussing on the origins of the ‘Cape clades’, which represent 50% of the species in the Cape flora. A wide range of source areas is identified as well as awidetime span over which the recruitment into the Cape took place. In chapter five I investigate the migration routes of Cape centred lineages, through the Afrotemperate Regions. Four lineages are included in this meta analysis. A general pattern of migrations around the Afrotemperate Region is identified and the ages of these migrations are estimated. The rates of diversification in each of the Afrotemperate regions are compared. 179 Posted at the Zurich Open Repository and Archive, University of Zurich ZORA URL: https://doi.org/10.5167/uzh-163608 Dissertation Published Version Originally published at: Galley, Chloé A. Macro-evolutionary and macro-ecological studies on the Cape flora, with focus on the Pentaschistis clade (Poaceae). 2007, University of Zurich, Faculty of Science. 2 Macro-evolutionary and Macro-ecological Studies on the Cape Flora, with Focus on the Pentaschistis Clade (Poaceae) Dissertation zur Erlangung der naturwissenschaftlichen Doktorwürde (Dr. sc. nat.) vorgelegt der Mathematisch-naturwissenschaftlichen Fakultät der Universität Zürich von Chloé A. Galley aus Frankreich und Grossbritannien Promotionskomitee Prof. Dr. H. Peter Linder (Vorsitz) Prof. Dr. Elena Conti Dr. Reto Nyffeler Zürich, 2007 Acknowledgements My first and very important thanks go to my supervisor, Peter Linder. I have been inspired by his manner of picking apart problems and data, testing ideas and carrying out novel research. These things helped make the research and this thesis exciting to carry out, and I hope and I think that I have taken some of these skills on board. Additionally, he has a very positive attitude to problem solving which is always welcomed in project work. Through him, I was introduced to the stunning mountains and fascinating vegetation of the Cape and Afromontane Regions, and especially to a curious group of grasses. Timo van der Niet also played an important role in this project. This stretches from companionship in the mountains of both Africa and Europe, to many discussions, ideas, inspiration and especially support. Chris Hardy likewise was always eager to discuss problems, and helped during field work and also tremendously with analytical issues. Many thanks to: Dirk Bellsteadt and Benny Bytebier especially, for being great co-authors; Philip Moline for help with the GIS; Lukas Taxböck for help with a translation; Frank Rutschmann for help with analytical problems; Urs Landergott for help with statistics; the Zürich Valkyries for keeping me sane, my family for many years of continual support and Merran Mathews and Sam Wüst for much appreciated support and clarifying the texts. The project was given a head-start by collections of silica plant material from Tony Verboom and Nigel Barker, for which I am very grateful. Fieldwork requires a lot of help from people in the host country. In the Cape, Terry Trinder-Smith and staff at the Bolus herbarium were generous with their time and making me feel welcome there. Also thanks to: Tony Verboom, and Nicola Berge especially, for being excellent ‘field-buddies’ during our adventures in 2004 and for their hospitality; Julia Born and Berit Gehrke for great companionship in the field; Lynn Fish, Clare Archer and Robert Archer from the National Botanic Institute in Pretoria for support and organisational help; Augustin Chikuni and staff at the Botanic Gardens of Malawi, Zomba; Halima and the staff at the herbarium at the National Museums of Kenya, Nairobi; Jacqueline Razanatsoa, Lalao Andriamahefarivo and the staff at the Parc de Tsimbazaza and Missouri Botanic Gardens office in Antananarivo, Madagascar; Pete Phillipson from the Missouri Botanical Garden; Sylvia Philips for assistance at the herbarium at Kew Botanic Garden just before I embarked on the project; and Mary Namanganda and Marc Lebouvier for silica material from places it would have not been possible for me to reach. Lastly, Reto Nyfeller is thanked for dealing the administration relating to my many piles of specimens back in Zurich. Importantly, I would like to thank the generous sources of funding for supporting my research and conference attendance (the University of Zurich, the Swiss Academy of Sciences and the Science National Fund, grant number 31-66594-01) and fieldwork (Georges und Antoine Claraz-Schenkung). Lastly, but certainly not least, I thank the people and my friends at the Institute of Systematic Botany, Zurich. It would not have been the same without you all. The working atmosphere in the institute was always supportive, positive and full of interest, from informal meetings and discussions at tea-time, to more formal journal clubs. There were always opportunities to share ideas, ask for advice or technical support and thereby enjoy research and learning. Table of contents Introduction………………………………………………………………………...1 Chapters 1. The phylogeny of the Pentaschistis clade (Danthonioideae, Poaceae) based on cpDNA, and the evolution and loss of complex characters………………………………………………………………………..…7 2. How do so many Pentaschistis species fit into the Cape Region, South Africa?………..……………………………………………….......49 3. New species and taxonomic changes within Pentaschistis (Danthonioideae, Poaceae) from the Western Cape, South Africa……………….93 4. Geographical affinities of the Cape flora, South Africa………………….111 5. The Cape element in the Afrotemperate flora: from Cape to Cairo?.................................................................................................................143 Summary………………………………………………………………………….175 Zusammenfassung………………………………………………………………...179 Curriculum Vitae………………………………………………………………….183 Introduction Since it was first botanically explored, the flora of the region at the southern tip of Africa has been known to be special (Goldblatt 1997; Linder 2006). Not only does this Cape flora have high species endemism (69%, Goldblatt & Manning 2002, a level more characteristic of island floras), but it is also incredibly species rich. 9,030 vascular plant species (Goldblatt & Manning 2002) are packed into only 90,000km2. Such diversity is typical of