Ecology of Vascular Epiphytes in Urban Forests with Special Reference to the Shrub Epiphyte Griselinia Lucida
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Ecology of vascular epiphytes in urban forests with special reference to the shrub epiphyte Griselinia lucida A thesis submitted in partial fulfilment of the requirements for the degree of Master of Science in Biological Sciences at The University of Waikato by Catherine Louise Bryan 2011 Abstract This research investigated the ecology of vascular epiphytes and vines in the Waikato region of the North Island, and the water relations of the shrub hemiepiphyte Griselinia lucida. The main goal was to develop robust recommendations for the inclusion of epiphytic species in urban forest restoration projects. To achieve this, three broad questions were addressed: 1. How are vascular epiphytes and vines distributed throughout the nonurban and urban areas of the Waikato region, and how does this compare to other North Island areas? 2. Why are some epiphyte and vine species absent from urban Hamilton and what opportunities exist for their inclusion in restoration projects? 3. How does Griselinia lucida respond to desiccation stress and how does this compare to its congener G. littoralis? To investigate questions one and two, an ecological survey of the epiphyte communities on host trees in Waikato (n=649) and Taranaki (n=101) was conducted, alongside canopy microclimate monitoring in five Waikato sites. Results show that epiphyte and vine populations in Hamilton City forests represent only 55.2 % of the total Waikato species pool, and have a very low average of 0.8 epiphyte species per host. In contrast, the urban forests of Taranaki support 87.9 % of the local species pool and have an average of 5.5 species per host tree. The low diversity and abundance in urban Waikato can be primarily attributed to the alteration of canopy microclimates by edge effects. Mean temperature and vapour pressure deficits in Waikato were 1.9 °C and 1.1 kPa (respectively) higher in the canopy of small urban patches than the larger, nonurban forests. These warmer and drier conditions are speculated to be interrupting species accumulation and community formation processes. This phenomenom is not as pronounced in Taranaki which has larger trees and higher rainfall. Epiphyte diversity and abundance was also found to be associated with seed dispersal distances and the size, bark type, and architecture of host trees. To link the microclimate findings with physiological limitations of epiphytes and to address question three, a desiccation tolerance experiment was conducted on ii the shrub hemiepiphyte Griselinia lucida. Moderate and severe levels of desiccation stress were applied to seedlings of G. lucida and its terrestrial congener, G. littoralis. Both species endured over two months of drought with negligible mortality. In G. lucida, stomatal conductance reduced to zero, and leaf bulk elastic modulus reduced from 8.09 (±0.51) MPa in the control group to 3.66 (±0.61) MPa under severe stress. When compared to G. littoralis, G. lucida exhibited a more acute response to stress and recovered faster with rewatering. However, the overall response of each species was similar and both species can be classified as desiccation postponers. To summarise and combine the findings of this research with existing information on Griselinia lucida, a contribution to the New Zealand Biological Flora Series for this species is presented. Recommendations for the inclusion of epiphytes in restoration projects are presented. Reintroductions should use epiphyte and vine species that are appropriate for the conditions of the target forest, and focus on large host trees in relatively humid microclimates. iii Acknowledgments This Master of Science has been both the most rewarding and most challenging thing I have ever done so I would like to acknowledge the enormous contribution made by the many people who have helped me along the way. Firstly, I would like to extend my most sincere thanks to my primary supervisor Professor Bruce Clarkson for the endless inspiration and encouragement. The many opportunities you have provided me with have played a huge part in making this experience so awesome. A huge thank-you also goes to my secondary supervisor, Dr. Michael Clearwater for helping me to learn plant physiology from scratch and for answering my many questions; your input has changed my perspective of science and greatly improved my research, analysis and writing skills. Enormous thanks go to the two people with whom I have shared many hours of field work, travelling, equipment design and calibration, plant measuring, writing, and stressing, but mainly laughing, mocking, and scheming. Cheers to my fantastic research comrades, Fiona Clarkson and Kieran Miller. Thanks to Toni Cornes for providing field gear, maps, statistics advice, a welcome whiteboard, and a healthy grounding whenever we lost it. Thanks to Colin Monk for being an endless source of solutions for any and every problem. Thanks also to the friendly people in the biology office and the many other staff who have been helpful throughout the last two years. Thanks to Stacey Foster, Emma Coleman, Rachel Thomson and Lucy Munro. You have each been very kind in giving me advice, encouragement, and your time, to process data and proofread my work. Thanks to the people and organisations that have helped fund my studies. Your assistance has significantly reduced my stress levels and made a lot of my research possible. Accordingly, I am grateful to the University of Waikato for the Masters Scholarship and contributions from the FRST urban restoration programme (UOWX0903). Thanks to the NZ Freemasons, the Perry Foundation, the Whakatane Historical Society, Forest and Bird, the Waikato Branch of the New Zealand Federation of Graduate Women, and the Royal Society of New iv Zealand for generously awarding me grants or scholarships. Thanks also to the George Mason Charitable Trust, Gary Bastin, and Puke Ariki for funding my research in Taranaki and hosting our presentations. A lot of my motivation for this work has come from our incredible native forests. For access to these wonderful places I would like to thank the Department of Conservation, Maungatautari Ecological Island Trust, Pukemokemoke Bush Reserve Trust, Hamilton City Council, New Plymouth District Council and private land owners. In Taranaki, my thanks go to Bill Clarkson for being an invaluable guide around your wonderful local forests and opening your home and nursery to us. I am very grateful to my friends who have provided not only welcome distractions and enjoyable breaks, but also lots of inspiration as they pursue their own goals. Special thanks to Tess Embling for sharing with me your strength and passion, it will never be forgotten. Much love to Mum and Dad for your unrelenting encouragement and faith in me, and of course for always being so proud. To my sister Stacey, thanks for your assistance in the lab and for being a listening ear that helped me hold onto some sanity. Thanks to everyone who has contributed to my research that I have regrettably forgotten to mention, this has been a task achieved with many hands and I truly appreciate every one of them. Finally, I would like to thank Hadley for being my rock and dealing with everything I have thrown at you. I simply could not have done this without you. Here‟s to the future! v Table of Contents Abstract .................................................................................................................... ii Acknowledgments .................................................................................................. iv Table of Contents ................................................................................................... vi List of figures ......................................................................................................... ix List of tables ......................................................................................................... xiii Chapter One: Introduction ................................................................................... 1 1.1 Ecological Restoration .............................................................................. 1 1.2 Epiphytes and vines .................................................................................. 4 1.3 Griselinia lucida ....................................................................................... 8 1.4 Research Objectives and Questions .......................................................... 9 1.5 Thesis Outline ........................................................................................... 9 Chapter Two: Epiphyte and vine species diversity and abundance in Waikato and Taranaki regions .......................................................................................... 11 2.1 Introduction ............................................................................................. 11 2.2 Research objectives and questions .......................................................... 14 2.3 Methods .................................................................................................. 15 2.3.1 Study regions ................................................................................... 15 2.3.2 Study sites ........................................................................................ 16 2.3.3 Sampling methods ........................................................................... 28 2.3.4 Edge:area ratios ............................................................................... 28 2.3.5 Microclimate monitoring ................................................................