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The Systematic Revision of Chaetanthera Ruiz & Pav., and The A systematic revision of Chaetanthera Ruiz & Pav., and the reinstatement of Oriastrum Poepp. & Endl. (Asteraceae: Mutisieae) Alison Margaret Robertson Davies München 2010 A systematic revision of Chaetanthera Ruiz & Pav., and the reinstatement of Oriastrum Poepp. & Endl. (Asteraceae: Mutisieae) Dissertation der Fakultät für Biologie der Ludwig-Maximilians-Universität München vorgelegt von Alison Margaret Robertson Davies München, den 03. November 2009 Erstgutachter: Prof. Dr. Jürke Grau Zweitgutachter: Prof. Dr. Günther Heubl Tag der mündlichen Prüfung: 28. April 2010 For Ric, Tim, Isabel & Nicolas Of all the floures in the meade, Thanne love I most those floures white and rede, Such as men callen daysyes. CHAUCER, ‘Legend of Good Women’, Prol. 43 (c. 1385) “…a traveller should be a botanist, for in all views plants form the chief embellishment.” DARWIN, ‘Darwin’s Journal of a Voyage round the World’, p. 599 (1896) Acknowledgements The successful completion of this work is due in great part to numerous people who have contributed both directly and indirectly. Thank you. Especial thanks goes to my husband Dr. Ric Davies who has provided unwavering support and encouragement throughout. I am deeply indebted to my supervisor, Jürke Grau, who made this research possible. Thank you for your support and guidance, and for your compassionate understanding of wider issues. The research for this study was funded by part-time employment on digital archiving projects coordinated via the Botanische Staatssammlung Munchen (INFOCOMP, 2000 – 2003; API- Projekt, 2005). Appreciative thanks go to the many friends and colleagues from both the Botanische Staatssammlung and the Botanical Institute who have provided scientific and social support over the years. Philomena Bodensteiner, Peter Döbbeler, Tini Ehrhart, Matthias Erben and Lilo Klingenberg deserve a special mention. Much of the evidence supporting the final conclusions in the thesis was acquired through laboratory research and S.E.M. imaging. This would not have been possible without the patient schooling and friendly assistance of Mrs. E. Vosyka, Dr. E. Facher and Mr E. Marksteiner. Travel grants were received from the DAAD (the German Academic Exchange Service) to visit Chile twice, in 2001 and 2002. My stay in Chile and the field excursions were made far easier thanks to Sunke Nef & family, Diettrich & Arnulf Becker, and Melica Muñoz-Schick in Santiago, and Roberto Rodriguez, Señora Gleisner & family, and Marcelo Baeza & family in Concepción. Special thanks are extended to Mary Kalin Arroyo, an active botanist and ecologist in Chile, who has an ongoing scientific interest in Chaetanthera. Permission to use digital images was kindly granted by Michal Belov (www.Chileflora.com), Mauricio Bonifacino (MVFA, Uruguay), Michael Dillon, María Terese Eyzaguirre (Fundación R.A. Philippi), Jürke Grau, Michel Grenon, Irene Till-Bottraud (Station Alpin Joseph Fourier, Grenoble) and Mauricio Zuñiga. Finally, a huge “Thank you” to my parents Jean & Iain Robertson, and my parents-in-law Elizabeth & Peter (sadly recently died). 1 Contents Abstract .................................................................................................................................. 5 Zusammenfassung .................................................................................................................. 6 List of Figures ........................................................................................................................ 7 List of Tables ....................................................................................................................... 12 I Introduction ...................................................................................................................... 15 1 Philosophy of taxonomic research ............................................................................... 15 2 Goals of taxonomic research ........................................................................................ 16 3 Application to Chaetanthera & Oriastrum .................................................................. 16 3.1 Historical background .......................................................................................... 16 3.2 Novel sources of variation ................................................................................... 17 3.3 Re-defining taxonomic relationships ................................................................... 18 II Materials, methods & measurements ............................................................................... 20 1 Materials ...................................................................................................................... 20 2 Methods........................................................................................................................ 20 2.1 Capitula & floral dissection ................................................................................. 20 2.2 Involucral bract anatomy studies ......................................................................... 20 2.3 Achene & pappus studies ..................................................................................... 20 2.4 Pollen studies ....................................................................................................... 21 2.5 Chromosome studies ............................................................................................ 21 2.6 Images & illustrations .......................................................................................... 22 2.7 Geographical data ................................................................................................ 22 3 Measurements .............................................................................................................. 22 III Variation of characters in Chaetanthera & Oriastrum .................................................... 24 1 General remarks ........................................................................................................... 24 2 Morphological & anatomical variation ........................................................................ 24 2.1 Life cycle & habit ................................................................................................ 24 2.2 Leaves .................................................................................................................. 29 2.3 Indumentum ......................................................................................................... 30 2.4 Involucral bracts................................................................................................... 30 2.4.1 Arrangement .................................................................................................... 30 2.4.2 Morphology of the inner involucral bracts ...................................................... 31 2.4.3 Anatomy of the inner involucral bracts ........................................................... 33 2.5 Pappus .................................................................................................................. 34 2.6 Floret variation: ray dimorphism & showiness .................................................... 36 2.7 Anthers & pollen .................................................................................................. 38 2.8 Styles & stigmas .................................................................................................. 41 2.9 Achenes ................................................................................................................ 43 2.9.1 Achene shape ................................................................................................... 43 2.9.2 Carpopodium.................................................................................................... 44 2.9.3 Achene hairs..................................................................................................... 44 2.9.4 Achene testa epidermis .................................................................................... 48 3 Cytological variation .................................................................................................... 50 4 Genetic variation .......................................................................................................... 54 IV Form, function & habitat.................................................................................................. 56 1 Introduction .................................................................................................................. 56 2 Homoplasy ................................................................................................................... 56 3 Habitats & stress .......................................................................................................... 57 4 Form & function .......................................................................................................... 57 4.1 Rosettes, succulence & sclerophylly in Chaetanthera ........................................ 57 2 4.2 Cushions, stem buds & scleroid bracts in Oriastrum .......................................... 58 4.3 Fragile pappus & indehiscent achenes: dispersal brakes ..................................... 59 4.4 Xeromorphic adaptations in pollen, pericarp & testa epidermis .......................... 60 4.5 Competitive ability............................................................................................... 60 4.6 Reproductive strategy
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