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Comparative Flowering Ecology of Fraxinus Excelsior, Acer

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Comparative Flowering Ecology of Fraxinus Excelsior, Acer Comparative flowering ecology of Fraxinus excelsior, Acer platanoides, Acer pseudoplatanus and Tilia cordata in the canopy of Leipzig’s floodplain forest Der Fakultät für Biowissenschaften, Pharmazie und Psychologie der Universität Leipzig eingereichte D I S S E R T A T I O N zur Erlangung des akademischen Grades Doctor rerum naturalium (Dr. rer. nat.) vorgelegt von Diplom Biologe Ophir Tal geboren am 24.7.1972 in Tel Aviv, Israel Leipzig, den 22.6.06. 1 To Shira 3 Abstract How do gender separation and the transition to wind pollination happen in temperate trees? What does the reproductive ecology in the crowns of temperate forest trees look like? These connected questions intrigued researchers before and since Darwin but it is only in the last years that a direct study of the latter question has been enabled. A research crane was used to study the flowering ecology of Fraxinus excelsior, Acer platanoides, Acer pseudoplatanus and Tilia cordata in Leipzig’s floodplain forest. These species originate from hermaphrodite insect pollinated plant families and exhibit different grades of gender separation and different stages between insect and wind pollination. As they are typical elements of temperate deciduous forests, an ecological comparison of their flowering ecology may shed new light on the evolution of gender separation and wind pollination in this habitat. Using the crane, gender distribution, flowering phenology in relation to microclimate, pollination levels (including pollen tubes in the styles) and fruit set were studied in ca. 200 trees over 2-4 years. Main results are a new appreciation of the sexual system of Fraxinus excelsior as dioecy, of Tilia cordata as andromonoecy and a detailed description of the intricacies of the heterodichogamous sexual system of Acer pseudoplatanus. Several flowering phenological patterns are described in Fraxinus excelsior and Acer platanoides in relation to microclimate in early spring. The role of small arthropods is underlined as gall mites may play a role in gender specialisation in Fraxinus excelsior, gall midges are related to maleness in T. cordata and thrips are probably the pollinators of Acer pseudoplatanus in the stand. Thrips pollination is suggested to be a possible stepping-stone between insect pollination and wind pollination, which may drive the transition in Acer pseudoplatanus and possibly in intensively flowering dominant species in other habitats. The study presents the complexity of the reproductive systems and the strong interdependencies among their elements. 5 Contents Introduction .......................................................................................................................... 11 Gender separation............................................................................................................. 11 Pollination modes............................................................................................................. 15 Flowering phenology and pollen exchange in the population.......................................... 19 Deciduous forests and canopy research ........................................................................... 22 The studied species........................................................................................................... 24 Aims of the study ............................................................................................................. 30 Methods................................................................................................................................ 31 The study site, the crane................................................................................................... 31 The studied trees............................................................................................................... 32 Gender and phenology ..................................................................................................... 33 Fraxinus excelsior........................................................................................................ 33 Acer platanoides........................................................................................................... 40 Acer pseudoplatanus .................................................................................................... 40 Synchrony measures..................................................................................................... 43 Tilia cordata................................................................................................................. 45 Pollen, visitors, pollination and fruit................................................................................ 46 Pollen size .................................................................................................................... 46 Flower visitors.............................................................................................................. 47 Pollination level............................................................................................................ 48 Fruit .............................................................................................................................. 50 Results .................................................................................................................................. 53 Overview .......................................................................................................................... 53 Fraxinus excelsior............................................................................................................ 57 Gender .......................................................................................................................... 57 Flowering phenology.................................................................................................... 61 Pollination and fruit...................................................................................................... 72 Acer platanoides............................................................................................................... 77 Acer pseudoplatanus ........................................................................................................ 85 Gender .......................................................................................................................... 85 Flowering phenology.................................................................................................... 88 Pollination and insects.................................................................................................. 93 Fruit .............................................................................................................................. 98 Tilia cordata................................................................................................................... 104 7 Contents Gender ........................................................................................................................ 104 Flowering phenology.................................................................................................. 110 Pollination and insects................................................................................................ 112 Fruit ............................................................................................................................ 115 Discussion .......................................................................................................................... 119 Fraxinus excelsior.......................................................................................................... 119 Sexual system............................................................................................................. 119 Differences between males and females .................................................................... 123 Flowering phenology, climate and reproduction........................................................ 127 Acer platanoides............................................................................................................. 133 Flowering phenology.................................................................................................. 133 Gender and pollination............................................................................................... 134 Acer pseudoplatanus ...................................................................................................... 136 The heterodichogamous sexual system...................................................................... 136 Thrips pollination ....................................................................................................... 141 Heterodichogamy: Asymmetries and reproduction.................................................... 143 Tilia cordata................................................................................................................... 149 Male flowers and gall midges .................................................................................... 149 Phenology................................................................................................................... 151 Pollination and insects in the inflorescences.............................................................. 152 Fruit ............................................................................................................................ 155 An ecological comparison of the species
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