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Masting and Insect Pollination in the Dioecious MASTING AND INSECT POLLINATION IN THE DIOECIOUS ALPINE HERB ACIPHYLLA A thesis submitted in partial fulfilment of the requirements for the Degree of Master of Science in Biological Science in the University of Canterbury by Laura May Young University of Canterbury 2006 Table of Contents List of Tables ............................................................................................................................1 List of Figures ...........................................................................................................................4 Abstract .....................................................................................................................................6 Chapter 1 - Introduction .........................................................................................................8 1.1 Introduction to the genus Aciphylla .........................................................................8 1.1.2 Aciphylla – a unique genus in the family Apiaceae .................................8 1.2 Historical background ............................................................................................10 1.3 Taxonomy ..............................................................................................................10 1.4 Why study the reproductive ecology of Aciphylla? ...............................................11 1.5 Dioecy and sexual dimorphism ..............................................................................11 1.6 Insect pollination ....................................................................................................13 1.6.1 Insect pollination and the importance of floral display ......................................13 1.7 Mast seeding ..........................................................................................................14 1.8 Why do some plant species mast when such high costs are involved? .................16 1.8.1 Proximate versus ultimate reasons for masting ......................................16 1.8.2 Non-evolved hypothesis to explain mast-seeding ...................................16 1.8.3 Evolutionary hypotheses to explain mast-seeding ..................................17 1.8.3.1 Predator satiation .....................................................................18 1.8.3.2 Wind-pollination ......................................................................18 1.8.3.3 Animal fruit dispersal and large seed size ...............................19 1.8.3.4 Accessory costs of reproduction ..............................................19 1.8.3.5 The animal pollination hypothesis ...........................................19 1.9 The bizarre reproductive ecology of Aciphylla ......................................................20 1.10 Research Aims .....................................................................................................21 Chapter 2- Does floral display size affect reproductive success in Aciphylla? .................23 2.1 Introduction ............................................................................................................23 2.1.1 The importance of floral display size at multiple levels in Aciphylla .....23 2.1.2 Measuring floral display at the inflorescence and plant level .................25 2.1.2.1 The importance of resource and pollen limitation in Aciphylla – does female success depend on individual reproductive effort? .......27 i 2.1.3 Measuring population-level flowering effort – pollinator efficiency .....29 2.1.4 Objectives ...............................................................................................32 2.1.4.1 Part (A): Floral display at the level of the individual plant .................32 2.1.4.2 Part (B): Population level flowering effort - male to female flowering densities ...........................................................................................33 2.2 Site and species selection .......................................................................................35 2.2.1 Study species ...........................................................................................37 2.2.1.1 Aciphylla aurea W.R.B. Oliver ................................................37 2.2.1.2 Aciphylla scott-thomsonii Cockayne & Allan ..........................38 2.2.1.3 Aciphylla ‘lewis’ ......................................................................38 2.2.2 Field work ...............................................................................................40 2.3 Methods ..................................................................................................................40 2.3.1 Part (A): Floral display at the level of the individual plant ....................40 2.3.2 Part (B): Population level flowering effort - male to female flowering densities ...........................................................................................................44 2.3.3 Pollen limitation ......................................................................................46 2.3.4 Other field measurements .......................................................................47 2.3.5 Lab work .................................................................................................47 2.3.5.1 Measuring reproductive success – counting seeds ...................47 2.3.5.2 Germination experiments .........................................................49 2.3.6 Statistical Analyses .................................................................................50 2.4 Results ....................................................................................................................51 2.4 Part (A): Effects of insect-exclusion bags, plant-level floral display, hand- pollination, and local pollen on seed set ......................................................................51 2.4.1 Mean seed set rates of control inflorescences at all sites ........................51 2.4.2 Effects of bagging on seed set rate .........................................................52 2.4.3 Effects of floral display on seed set ........................................................53 2.4.3.1 Analyses by site .......................................................................53 2.4.3.2 Analyses by treatment and graphical representation to compare sites ........................................................................................58 2.4 Part (B): Effects of female isolation, hand-pollination and pollen on seed set ......66 2.4.4 Analyses by site ......................................................................................66 2.4.5 Seed germination ....................................................................................70 2.4.6 Summary .................................................................................................70 ii 2.5 Discussion ..............................................................................................................72 2.5.1 Part (A): Floral display at the plant level ................................................72 2.5.1.1 The importance of insect versus wind as pollen vectors in Aciphylla ..............................................................................................72 2.5.1.2 The importance of plant-level floral display (and how it is measured) on reproductive success ......................................................73 2.5.2 Part (B): Population level flowering effort .............................................75 2.5.3 Conclusions .............................................................................................77 Chapter 3 - The effects of floral display and population structure on insect visitation rates and how this relates to reproductive success .............................................................79 3.1 Introduction ............................................................................................................79 3.1.1 Aims ........................................................................................................80 3.2. Methods ................................................................................................................81 3.2.1 Collecting insects ....................................................................................81 3.2.2 Insect visitation surveys ..........................................................................81 3.2.3 Insect visitation to male inflorescences ..................................................82 3.2.4 Insect visitation rates in response to floral display .................................82 3.2.5 Statistical Analyses .................................................................................83 3.3 Results ....................................................................................................................84 3.3.1 What insects are visiting Aciphylla inflorescences? ...............................84 3.3.2 Are insects attracted to larger floral displays? ........................................85 3.3.2.1 Insect visitation rates and flower removal manipulation .........85 3.3.2.2 Insect visitation and inflorescence height ................................87 3.3.2.3 Insect visitation and inflorescence flowering length ...............89 3.3.3.4 Does bract removal affect insect visitation rates? ....................91 3.3.3.5 Insect visitation and number of inflorescences per plant .........92 3.3.4 Insect visitation to isolated flowering female plants ...............................94
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