Functional Integration of Floral Plant Traits: Shape and Symmetry, Optical Signal, Reward and Reproduction in the Angiosperm Flower
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Functional Integration of Floral Plant Traits: Shape and Symmetry, Optical Signal, Reward and Reproduction in the Angiosperm Flower Dissertation zur Erlangung des Doktorgrades (Dr. rer. nat.) der Mathematisch-Naturwissenschaftlichen Fakultät der Rheinischen Friedrich-Wilhelms-Universität Bonn vorgelegt von Andreas Wilhelm Mues aus Kirchhellen Bonn, den 20. Januar 2020 1 2 Angefertigt mit Genehmigung der Mathematisch-Naturwissenschaftlichen Fakultät der Rheinischen Friedrich-Wilhelms-Universität Bonn Erstgutachter: Prof. Dr. Maximilian Weigend, Universität Bonn Zweitgutachter: Prof. Dr. Eberhard Fischer, Universität Koblenz Tag der Promotion: 30. April 2020 Erscheinungsjahr: 2020 3 4 Acknowledgements I thank Prof. Dr. Maximilian Weigend, supervisor, for his guidance and support, and for giving me the opportunity to study the holistic subject of floral functional integration and plant-animal interaction. I am grateful for the experience and for the research agendas he entrusted to me: Working with the extensive Living Collections of Bonn Botanical Gardens was an honour, and I have learned a lot. I thank Prof. Dr. Eberhard Fisher, for agreeing to be my second supervisor, his advice and our shared passion for the plant world. I would like to thank many people of the Nees Institute and Bonn Botanical Gardens who contributed to this work and who gave me good memories of my years of study: I thank Lisabeth Hoff, Tianjun Liu, Luisa Sophie Nicolin and Simon Brauwers for their contribution in collecting shares of the raw data together with me, and for being eager students – especially counting pollen and ovule numbers and measuring nectar reward was a test of patience sometimes, and we have counted and measured a lot … Thank you! Special thanks go to Gardeners of the Bonn Botanical Gardens, for their constant support throughout the years, their love for the plant world in general and their commitment and care for the Living Collection: Klaus Mahlberg (Streptocarpus), Birgit Emde (carnivorous plants), Klaus Bahr (Geraniales), Bernd Reinken and Klaus Michael Neumann. I thank Dr. Stefan Abrahamczyk and Dr. Federico Luebert for their support and valuable advice. I thank my other colleagues and peers for their collegiality and for sharing a good time at the Nees Institute, especially my roommates Dr. Bernadette Große-Veldmann and Dr. Rafael Acuña Castillo, and Dr. Julia Bechteler, Tim Böhnert, Dr. Juliana Chacón, Hans-Jürgen Ensikatt, Dr. Jonathan Hense, Dr. Norbert Holstein, Julius Jeiter, Thomas Joßberger, Sandro Kolbaia, Dr. Michael Krug, Karola Maul, Felix Merklinger, Dr. Adeel Mustafa, Dr. Jens Mutke, Constantin Poretschkin, Nicole Schmandt, Claudia Schütte, Amélie Tessartz and Dr. Lara Weiser. Special thanks go to Elke Hedke and Prof. Dr. Annette Sheersoi for their support and for keeping up the good team spirit. I thank my siblings Christian and Angelika and my brother-in-law Peter for their friendship, and I am grateful for my parents Wilhelm and Sylvia, and their constant support throughout the years. 5 6 CONTENTS I. General Introduction ........................................................................................................................... 15 I.1 Foreword ..................................................................................................................................... 15 I.2 Organisation of pollen receipt and guidance .............................................................................. 16 I.2.1 Floral structure – morphology, architecture, symmetry and shapes ..................................... 16 I.2.2 Plant presentation traits – floral attractants and reward ....................................................... 18 I.3 Reproduction: Sexual systems, breeding systems, gamete production and p/o ratio ............... 23 I.4 Pollination syndromes, suites of floral plant traits and plant-animal interaction ...................... 24 I.5 Hypotheses ................................................................................................................................. 30 I.6 Goals ........................................................................................................................................... 30 I.6.1 Research Questions................................................................................................................. 30 I.6.2 Specific Objectives .................................................................................................................. 31 I.7 Overview of the Dissertation ...................................................................................................... 32 I.8 Contribution to Chapters ............................................................................................................ 33 I.9 Literature .................................................................................................................................... 36 II. Diversification of floral function in Geraniales – why so dapper? Breeding system related variables not correlated to pollinator guilds and flower symmetry .......................................................................... 45 II.1 Abstract ....................................................................................................................................... 45 II.2 Introduction ................................................................................................................................ 46 II.3 Materials and Methods ............................................................................................................... 48 II.3.1 Assessment of gamete production and nectar reward ...................................................... 48 II.3.2 Level of measurement and Statistical Analysis ................................................................... 49 II.4 Results ......................................................................................................................................... 51 II.4.1 Pollinator guilds .................................................................................................................. 51 II.4.2 Flower symmetry ................................................................................................................ 51 II.4.3 Growth habit ....................................................................................................................... 52 II.4.4 Gamete production ............................................................................................................. 52 II.4.5 Nectar production ............................................................................................................... 55 II.4.6 NMDS ordinations and vector fitting .................................................................................. 57 II.4.7 Phylogenetic mapping of plant traits and pollinator guilds ................................................ 59 II.5 Discussion .................................................................................................................................... 60 II.5.1 Missing resource efficacy despite pollinator positioning via flower symmetry ................. 60 7 II.5.2 Pollinator guilds correlate with nectar, but not with variables of the breeding system .... 61 II.5.3 Pollinator guilds and plant traits are phylogenetically constraint ...................................... 62 II.6 Acknowledgments ....................................................................................................................... 63 II.7 Literature .................................................................................................................................... 63 III. Frost flowers – Pollination syndromes of Hamamelidaceae independent from flowering time ....... 69 III.1 Abstract ....................................................................................................................................... 69 III.2 Introduction ................................................................................................................................ 70 III.3 Materials and Methods ............................................................................................................... 72 III.3.1 Assessment of inflorescence and flower morphology ........................................................ 72 III.3.2 Assessment of pollen production, ovule numbers and p/o ratios ...................................... 73 III.3.3 Assessment of inflorescence size ........................................................................................ 74 III.3.4 Assessment of UV light patterns ......................................................................................... 74 III.3.5 Assessment of nectar reward ............................................................................................. 74 III.3.6 Cryo-Scanning Electron Microscopy (CSEM) ....................................................................... 75 III.3.7 Phylogenetic Analysis .......................................................................................................... 75 III.3.8 Statistical Analysis ............................................................................................................... 75 III.4 Results ......................................................................................................................................... 77 III.4.1 Clustering of gametic variables retrieves two clusters ......................................................