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University of Cape Town LIFE HISTORY TRAITS OF SOUTH AFRICAN ENCEPHALARTOS SPP. (ZAMIACEAE) AND THEIR IMPLICATIONS FOR UNDERSTANDING POPULATION STRUCTURE, RESPONSES TO THREATS AND EFFECTIVE CONSERVATION ACTION Kim Martina Konings Dissertation presented for the degree of Master of Science in the Department of Biological Sciences, University of Cape Town February 2016 Supervisors: Prof John Sydney Donaldson (South African National Biodiversity Institute) Prof Michael Timm Hoffman (Plant Conservation Unit, University of Cape Town) University of Cape Town 1 The copyright of this thesis vests in the author. No quotation from it or information derived from it is to be published without full acknowledgement of the source. The thesis is to be used for private study or non- commercial research purposes only. Published by the University of Cape Town (UCT) in terms of the non-exclusive license granted to UCT by the author. University of Cape Town DECLARATION I know the meaning of plagiarism and declare that all of the work in the dissertation, save for that which is properly acknowledged, is my own Kim M. Konings. February 2016 2 ACKNOWLEDGEMENTS I would like to thank the South African National Biodiversity Institute (SANBI) for this opportunity to study further by providing funding for this study. Thank you to my supervisors, Prof. John Donaldson and Prof Timm Hoffman for their support and careful editing of this document. A special thanks to John for his patience and guidance during my research and for many visits to UCT, spending a generous amount of time assisting me. Your knowledge and passion for cycads and conservation are inspiring. Thanks to Timm for inviting me into the Plant Conservation Unit (PCU) family at UCT and showing care and support for my work. Many people played a role in my field work, a special mention to Desale Okubamichael, who provided me with a lot of guidance on how to carry out the field measurements, as well as providing me with some of his data (collected with Sam Jack and de Wet, thanks Sam and de Wet!) and in general being a great friend. Thank you to Sarah Findlay and Luke Potgieter, my field assistants. You guys were amazing. Sarah, you are the most encouraging wonderful being I have ever had the pleasure of spending an extended amount of time with in isolated circumstances. Luke, you were the one who kept us on track, I’m not sure how you kept sane, but we appreciated you. Thank you also to John and Timm for helping us with the first bit of field work. Also thanks to De Wet Bösenberg and Phakamani Xaba for helping me to organize equipment and accommodation. Thanks to Nigel and Heather McMaster for being so welcoming, taking an interest in our work, walking us to the sites and setting us up with a warm fire in the lovely Cameron’s Cottage. Thanks to Bruce Cleary for hosting us and making an amazing dinner, also to Ryan at Venn Grove for hosting us. Thank you to Carin Swart for assisting with fieldwork as well as Les Powrie and Megan Welsford for helping me to compile the environmental data. A big thanks goes to Prof. Emer. Les Underhill, A/Prof. Res Altwegg, Katya Mauff, Dr. Miguel Lacerda, Dr Ian Durbach, Dr Maria Pulido and Ushma Galal for help with statistics. 3 A massive thank you to my family, my mom, dad, granny, sister and brother-in-law, you guys are an unconditional support base. Thank you Mom and Dad for cooking for me and offering help, and in general just loving and encouraging me. A huge thank you to my friends. Petra, my awesome officemate who became a great friend! Thank you for your advice and your listening ear and our generally hilarious and thought-provoking conversations. Thanks to all the great people I met in the PCU for little tea chats, Carina, Abraham, Daniel, Sam, and of course a special mention to Rua for our awesome friendship, many tea breaks at UCT, as well countless off-campus shenanigans. To Connie and Kim for our middle-campus lunches, as well as Aimee...I love our awesome foursome, thanks girls for your constant encouragement and the infamous survival pack and famous cycad cake! Also to Job, Lulu, Simone, Alex M, Michelle, Jenn, Cres, Lorenzo, Sulona, Tumi, Moné, Jennie, James, Angela, Rikki, Shagita, Sandi, Tammy, Erica, Nico, Tristan, Helen, Lauren T, Alex S, Davide, Charl, Duncan, Willem, Liane, Nadene, Shirin, Alta, Nicole, Yoga Omies and anyone who I have forgotten, there are just so many good people in my life! You guys have all supported me and entertained me! And of course thanks goes to Nahko, I believe in the good things coming. 4 a) b) d) c) f) e) Life forms within the genus Encephalartos. a) E. laevifolius b) female cones of E. senticosus c) E. latifrons d) E. villosus e) E. cycadifolius f) E. altensteinii. Photos taken by John S. Donaldson. 5 ABSTRACT Understanding life history traits and how they influence population-level processes under different conditions have been important areas of study in evolution and ecology. One application in ecology has been to determine whether there are general responses that apply to species with similar life history traits, which could provide a predictive understanding of species with shared traits rather than having to adopt a species- specific approach to management. In this study, an analysis of life history traits was applied to cycads, the oldest living seeds plants which are at risk of extinction due to vulnerability to illegal wild harvesting and habitat loss. The study focused on South African cycad species within the genus Encephalartos, as well as two species from Swaziland. The first objective was to identify groups of species with similar life histories and to assess whether these groups are aligned with phylogenetic relationships or an association with particular environments. The second objective was to determine whether species with shared suites of life history traits exhibited a similar population stage structure. Hierarchical cluster analysis and Principal Component Analysis (PCA) were used to identify groups based on key life history traits, namely, cone number and coning interval, number of seeds produced, number of stems, stem height and plant habit. Four life history groups emerged, these were: Group 1 with single-stemmed, subterranean species such as E. villosus; Group 2 generally characterised by medium-height species with many stems such as E. lehmannii; Group 3 with the tallest species which generally produce the most cones at the most frequent interval such as E. natalensis; and Group 4 with dwarf-arborescent, multi-stemmed species such as E. horridus. The groups were not found to correspond with the most recent phylogeny for Encephalartos and thus phylogenetic constraints were excluded. An analysis of environmental variables for 193 populations of species in all groups, including maximum and minimum temperatures, mean annual precipitation, moisture index, rainfall co-efficient of variation and fire return frequency, showed that the means per group were not significantly different from each other for most of the variables. However, it was evident that the variable ranges showed clear trends with biological significance. These tended towards more stable, less variable environments with more predictable rainfall for Group 1 species, and less 6 stable, more variable environments with less predictable rainfall for species within Group 4. Groups 2 and 3 showed a large range and overlap in environmental associations with no consistent patterns. Population structure is valuable for determining whether recruitment in a population is adequate for species survival. Static life tables and Chi-Squared analyses were used to test differences in the population structure and survivorship patterns of species between and within the life history groups. If particular demographic patterns were shown to be emergent properties of certain suites of life history traits, autecological studies may be avoided for predictive-ecological conservation frameworks. The results showed that species in Group 1 had populations with a high proportion of juveniles and fewer reproductive adults, indicating high recruitment as well as high juvenile mortality. In contrast, species in Group 4 had populations with a high proportion of adults and fewer juveniles indicating low recruitment and high juvenile mortality with primary investment into adult plant persistence through vegetative suckering. Group 2 consisted mostly of rare species with very small populations and high proportions of adult plants showing evidence of episodic recruitment events. Species in Group 3 showed a similar trend towards intermittent recruitment with adult persistence, however, no explicit population structure emerged. This led to the conclusion that Groups 1 and 4 are relatively consistent and predictable, however, Groups 2 and 3 would better benefit from an autecological approach to management. This study was helpful to assemble life history data for South African Encephalartos, as well as environmental parameters for each species. Ultimately, it has shown that primary investment into seeds versus persistence, or a combination thereof, can influence population structure. 7 TABLE OF CONTENTS Declaration ......................................................................................................................................................................... 2 Acknowledgements ........................................................................................................................................................ 3 Abstract ................................................................................................................................................................................
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