Phenological, physiological, and ecological factors affecting the epiphyte Notheia anomala and its obligate host Hormosira banksii A thesis submitted in partial fulfilment of the requirements for the degree of Master of Science in Biology at the School of Biological Sciences University of Canterbury New Zealand by Isis Hayrunisa Metcalfe 2017 Table of Contents I Table of Contents Table of Contents ............................................................................................................................... I List of Figures .................................................................................................................................... V List of Tables ..................................................................................................................................... X Acknowledgements ...................................................................................................................... XIII Abstract ........................................................................................................................................... XV Chapter One - General Introduction ............................................................................................... 1 1.1. Introduction ........................................................................................................................... 2 1.1.1. Epiphytism ..................................................................................................................... 2 1.1.2. Marine epiphytes ............................................................................................................ 4 1.1.3. Costs and benefits to hosts ............................................................................................. 5 1.1.4. Costs and benefits to epiphytes ...................................................................................... 6 1.1.5. Facultative epiphytism vs. obligate epiphytism ............................................................. 7 1.2. Study species ......................................................................................................................... 8 1.2.1. Costs and benefits ........................................................................................................ 10 1.2.2. Why is the Hormosira-Notheia interaction of scientific interest? ............................... 11 1.3. Study aims ............................................................................................................................ 12 1.4. Study sites ........................................................................................................................... 14 Chapter Two - The reproductive dynamics of Notheia anomala ................................................. 17 2.1. Abstract ................................................................................................................................. 18 2.2. Introduction ........................................................................................................................... 18 2.3. Methods ................................................................................................................................. 20 2.4. Results ................................................................................................................................. 24 2.5. Summary ............................................................................................................................... 28 Chapter Three - The distribution and interactions of the obligate epiphyte Notheia anomala and its canopy forming host Hormosira banksii............................................................. 30 3.1. Abstract ................................................................................................................................. 31 Table of Contents II 3.2. Introduction ........................................................................................................................... 32 3.3. Methods ................................................................................................................................. 34 3.3.1. Global Distribution ...................................................................................................... 34 3.3.2. Distribution and abundance on vertical, regional, and temporal scales ...................... 35 3.3.3. Distribution and abundance on host plant scale (point of attachment) ........................ 35 3.3.4. Survival and growth (tagging and removal experiment) ............................................. 38 3.3.5. Stress assays (translocation experiments) .................................................................... 40 3.4. Results ................................................................................................................................... 43 3.4.1. Global Distribution ...................................................................................................... 43 3.4.2. Distribution and abundance on vertical, regional, and temporal scales ...................... 43 3.4.3. Distribution and abundance on host plant scale (point of attachment) ........................ 47 3.4.4. Survival and growth (tagging and removal experiment) ............................................. 51 3.4.5. Stress assays (translocation experiments) .................................................................... 53 3.5. Summary ............................................................................................................................... 56 Chapter Four - Notheia as a contributor to local-scale biodiversity ........................................... 59 4.1. Abstract ................................................................................................................................. 60 4.2. Introduction ........................................................................................................................... 61 4.3. Methods ................................................................................................................................. 64 4.3.1. Spatio-temporal distribution of mobile invertebrates .................................................. 64 4.3.2. Mimic and density experiment .................................................................................... 65 4.3.3. Snail grazing experiment ............................................................................................. 67 4.4. Results ................................................................................................................................... 68 4.4.1. Spatio-temporal distribution of mobile invertebrates .................................................. 68 4.4.2. Mimic and density experiment .................................................................................... 76 4.4.3. Snail grazing experiment ............................................................................................. 82 4.5. Summary ............................................................................................................................... 83 Chapter Five - General Discussion ................................................................................................. 86 Table of Contents III 5.1. The reproductive dynamics of Notheia anomala .................................................................. 87 5.2. The distribution and interactions between the obligate epiphyte Notheia anomala and its canopy forming host Hormosira banksii ................................................................................. 90 5.3. Notheia as a contributor to local-scale diversity ................................................................... 95 Literature Cited ............................................................................................................................. 100 Appendix ......................................................................................................................................... 112 List of Figures V List of Figures Chapter One Figure 1.1. A schematic diagram illustrating how a facilitation cascade can occur. The primary habitat former is indirectly facilitating the ‘clients’ (organisms, typically mobile animals that depend on structural habitat-formers) through the direct facilitation of a secondary habitat former (in this case, a mistletoe) (Thomsen et al. 2010). ................................................................................ 4 Figure 1.2. A schematic diagram summarising the costs and benefits of epiphytism for the host and epiphyte. For epiphytes, the benefits outweigh the costs, which is why epiphytism is such a dominant life form. For hosts, producing defences is costly, and many do not allocate a large amount of energy into epibiosis prevention. ........................................................................................ 5 Figure 1.3. A photograph of the epiphyte Notheia anomala growing on its obligate host Hormosira banksii. ............................................................................................................................................... 10 Figure 1.4. A section of the New Zealand map showing the distribution of Kaikoura on the North East coast of the South Island. The main sampling site for this project was at the Kaikoura Peninsula at Wairepo Reef, but some studies were also done at South Bay. ..................................... 15 Figure