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Brunsvigia-Nectarinia Pollination Mutualisms: Sunbirds and Candelabra Plants

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Brunsvigia-Nectarinia Pollination Mutualisms: Sunbirds and Candelabra Plants The copyright of this thesis vests in the author. No quotation from it or information derived from it is to be published without full acknowledgementTown of the source. The thesis is to be used for private study or non- commercial research purposes only. Cape Published by the University ofof Cape Town (UCT) in terms of the non-exclusive license granted to UCT by the author. University Variation in pollination across a fragmented landscape at the Cape of Africa Town Anton Pauw Cape of Thesis Presented for the Degree of Doctor of Philosophy Universityin the Department of Botany Science Faculty University of Cape Town April 2004 2 Statement The conception, planning, execution and writing of this thesis was entirely my own, except in the specific instance mentioned below. Chapter 7 describes a study conducted in collaboration with Dr. Julie Hawkins and Dr. Yvonne Charters. Their contributions were mainly through genetic laboratory work (Charters) and discussion on how to improve the manuscript (Hawkins). Town Cape of University 3 Acknowledgements I am grateful to my supervisor, William Bond, for sharing his insight into the natural world with me and for guiding my own enquiries. Steve Johnson and Julie Hawkins have been unofficial co-supervisors of this thesis. I am indebted to them for all that I continue to learn from them. Spencer Barrett directed my interest in stylar polymorphisms. Gabriela Demergasso is my Muse and proofreader. It is thanks to a group of passionate conservationists that anything at all remains of the Cape Lowland flora. The Darling Wildflower Society, and in particular Mark and John Duckitt, have been instrumental in saving some of the treasures ofthe Darling region from the plow. I am grateful to them for permission to Townwork on their farms. Farmer and conservationist, Neil McGregor, gave us access to his carefully guarded botanical treasures at Nieuwoudville. I am indebted to The Friends of Rondebosch Common, and in particular Betty Dwight, for sharingCape her knowledge of the flora of this small, urban conservation area. Dalton Gibbs and Clifford Dorse introduced me to the botanical rarities of the Rondevlei Natureof Reserve. lowe a special debt to the staff of the Botany Department of the University of Cape Town. Departmental Head, Jeremy Midgley, provided many stimulating conversations and fresh ideas, and commented on drafts of some chapters. Sandy Smuts made research possible by keeping the administrative machinery of the Botany DepartmentUniversity running smoothly. Terry Trinder-Smith, curator of the Bolus Herbarium, gave freely of his deep knowledge of the Cape Flora. Gonzalo Aguilar kept this tired computer going. I am indebted to Miranda Waldon of the Electron Microscope Unit for her patience and assistance. Kim Steiner, John Manning and Dee Snijman at the National Botanical Institute (Kirstenbosch) shared their deep knowledge of the Coryciinae, Iridaceae and Amaryllidaceae with me. For the difficult task of identifying insects I relied on the collections of the South African Museum and received assistance from Simon van Noort. Vin Whitehead helped me with the identification of Rediviva species. 4 Throughout this study I have been supported in many ways by the Botanical Society of South Africa, in particular Kristal Maze, Amrei von Hase and Mark Botha of the Cape Conservation Unit. The National Parks Board and Cape Nature Conservation provided permits and permissions. Tilla Raimondo, Anthony Roberts and CharI Pauw helped with the capture of sunbirds and the counting of seeds, and made fieldwork a pleasure. Yvonne Charters and Josephine Shockley did the genetics lab work for me. For statistical advice I am grateful to Frank Schurr and June Juritz. Amrei von Hase provided help with digital maps and geographical information of the study region. Bill Liltved provided invaluable locality information for orchid populations and provided some of the photographs presented here. The following herbaria are thanked for allowing me to rehydrate their ancient specimens: K, BM, BOL, NBG, SAM, GUTHRIE. This research was made possible by generous grants from the TownNational Research Foundation (South Africa), the Nuffield Foundation and the National Geographic Society. Cape of University I I 5 Abstract In this thesis I assessed the ability of established conservation areas to conserve plant-pollinator interactions. The study area was the species rich, but highly fragmented West Coast Lowlands of the Cape of Africa. I focused on the Nectarinia sunbird pollination system and the oil-collecting Rediviva bee pollination system. Before exploring the consequences of habitat fragmentation, it was necessary to study the natural history of the plant-pOllinator interactions. Among 16 conservation areas, I studied variation in the pOllination of Brunsvigia orientalis (Amaryllidaceae) by Nectarinia sunbirds. Pollen supplementation experiments and censuses of sunbirds showed that the observed failure of seed set could be attributed to the lack of sunbirds. Three factors: 1) the post-fire successional stage of the vegetation, 2) the number of flowering B. orientalis plants, and 3) the size ofTown the conservation area, explained 75 % of variation in seed set. Seed set failed in small conservation areas (22 - 30 ha), and was reduced in medium-sized conservation areas (811 - 1113 ha) compared with large conservation areas (> 15 000 ha). Among 27 conservation areas, I studied Capevariation in the pollination of an oil­ secreting orchid, Pterygodium catholicum, by the oil-collecting bee, Rediviva peringueyi. Variables that Significantly influenced the abilityof of nature reserves to conserve the orchid­ bee mutualism were: 1) the soil type, 2) the post-fire successional stage of the vegetation and 3) the interaction between reserve size and the nature of the surrounding land-use. A lack of pollinators led to reproductive failure in consecutive years in small conservation areas (4 - 385 ha) in an urban matrix. Correlated patterns of variation in seed capsule production suggested that the results obtained for P. catholicum could be extrapolated to several additional plant species which are also pollinated by R. peringueyi. Are these apparentUniversity anthropogenic declines in pollination real? Declines can only be detected with certainty by comparison with historical rates. I reconstructed the historical pollination landscape by detecting the genetic echo of past pollination, and by determining historical pollination rates from rehydrated herbarium specimens. Thus, I demonstrated that plant-pOllinator mutualisms have recently collapsed in small, urban conservation areas. Does a decline in pollination really matter? Patterns of variation in the composition of the community of oil-secreting plants suggest that the anthropogenic extirpation of oil­ collecting bees from small conservation areas has been followed by the extirpation of dependent plant species. Patterns of infraspecific variation in the floral traits in sunbird and oil-bee pollinated plants suggest an alternative to extinction. Pollinator loss may ultimately result in the evolution floral traits that decrease dependence on the absent pollinator. 6 Table of Contents Statement ...................................................................................................................... 2 Acknowledgements ...................................................................................................... 3 Abstract ........................................................................................................................ 5 Introduction ............................................................................................................... 11 Conventions and nomenclature ................................................................................ 14 SECTION 1 The Brunsvigia-Nectarinia pollination mutualisms: sunbirds and candelabra plants ........................................................................ 15 1 Reproductive dependence of Brunsvigia orientalis (Amaryllidaceae) on Nectarinia sunbirds (Nectarinidae) .................................................................. 16 Materials and methods .......................................................................................Town 16 Study species ................................................................................................... 16 Study sites ....................................................................................................... 17 Flower morphology, nectar characteristics and phenology ............................. 17 Breeding system .............................................................................................. 17 Alternative modes of reproduction ..................................................................Cape 18 Pollinators ........................................................................................................ 18 Results ................................................................................................................of 20 Functional floral morphology and phenology ................................................. 20 Breeding system .............................................................................................. 22 Alternative modes of reproduction .................................................................. 22 Pollinators and flower visitors ......................................................................... 22 Discussion .........................................................................................................
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