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Fruit-Frugivore Interactions in a Malagasy Littoral Forest: a Community-Wide Approach of Seed Dispersal

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Fruit-Frugivore Interactions in a Malagasy Littoral Forest: a Community-Wide Approach of Seed Dispersal Fruit-frugivore interactions in a Malagasy littoral forest: community-wide approach of seed dispersal UNIVERSITEIT ANTWERPEN Faculteit Wetenschappen Departement Biologie Fruit-frugivore interactions in a Malagasy littoral forest: a community-wide approach of seed dispersal Proefschrift voorgelegd tot het behalen van de graad van doctor in de Wetenschappen aan de Universiteit Antwerpen te verdedigen door An BOLLEN An BOLLEN ANTWERPEN 2003 Promotoren: Prof. dr. L. Van Elsacker Prof. dr. R. Verheyen Prof. dr. J. Ganzhorn Universiteit Antwerpen Promotoren: Prof. dr. L. Van Elsacker Faculteit Wetenschappen Prof dr. R. Verheyen Departement Biologie Prof dr. J. Ganzhorn Antwerpen, 2003 Fruit-frugivore interactions in a Malagasy littoral forest: a community-wide approach of seed dispersal Interacties tussen vruchten en frugivoren in een Madagaskisch littoraal regenwoud: een benadering van zaadverspreiding op niveau van het ecosysteem Proefschrift voorgelegd tot het behalen van de graad van doctor in de Wetenschappen aan de Universiteit Antwerpen te verdedigen door An BOLLEN Cover photographs: Front: -‘ambora’ Tambourissa purpurea (Monimiaceae) © An Bollen -‘bodono’ Cheirogaleus medius (Cheirogaleidae) © Han Remaut -‘hazombato’ Campylospermum obtusifolium (Ochnaceae) © An Bollen -‘horovana’ Hypsipetes madagascariensis (Pycnonotidae) © Han Remaut -‘tsilantria’ Vaccinium emirnense (Ericaceae) © An Bollen -‘varika’ a male Eulemur fulvus collaris (Lemuridae) © An Bollen -aerial photograph of the Sainte Luce area (middle) © An Bollen Back: -‘sanirambavy’ fruits of Tina thouarsiana (Sapindaceae) with bill traces of Coracopsis nigra (Psittacidae) © An Bollen -‘fanihy’ Pteropus rufus (Pteropodidae) © An Bollen -‘rotry mena’ Syzygium sp.2’ (Myrtaceae) © An Bollen -seeds of ‘voapaky vavy’ Uapaca littoralis (Euphorbiaceae) with gnawing marks of ‘voalava’ rodents © An Bollen -Eulemur fulvus collaris feeding in a fruiting tree © An Bollen -‘sjihely’ sardine-like fishes in Dec ‘00 in the village of Ambandriky © An Bollen -the people of the Ambandriky at the landing strip of Martin Pêcheur in front of the hill of S17 © An Bollen Throughout this PhD taxonomic names of the plant species follow the Flora of Madagascar (Humbert 1936-1984), the data base of Missouri Botanical Garden (http://mobot.mobot.org/W3T) ISBN 90-5728-042-6 © An Bollen Table of contents TABLE OF CONTENTS Acknowledgements ........................................................................................................1 Preface...........................................................................................................................5 I. General Introduction..................................................................................................7 II. Main Chapters ......................................................................................................... 13 1. Phenology of the littoral forest of Sainte Luce, south-east Madagascar.............. 15 Bollen A, Donati G Submitted to Biotropica 2. Tree dispersal strategies in the littoral forest of Sainte Luce .............................. 37 (south-east Madagascar) Bollen A, Van Elsacker L, Ganzhorn JU Submitted to Oecologia 3. Relations between fruits and disperser assemblages in the littoral .................... 59 forest of south-east Madagascar: a community level approach Bollen A, Van Elsacker L, Ganzhorn JU Submitted to Journal of Tropical Ecology 3a. Feeding ecology of Pteropus rufus (Pteropodidae) ................................... 85 in the littoral forest of Sainte Luce (south-east Madagascar) Bollen A, Van Elsacker L Acta Chiropterologica 4(1): 33-47 3b. Feeding ecology of Coracopsis nigra (Psittacidae) in ............................ 101 the littoral forest (south-east Madagascar). Bollen A, Van Elsacker L Submitted to Ostrich 4. Fruit characteristics in a dry deciduous and a humid littoral forest................... 111 of Madagascar: evidence for selection pressure through abiotic constraints rather than through co-evolution with lemurs as seed dispersers Bollen A, Donati G, Fietz J, Schwab D, Ramanamanjato J-B, Randrihasipara L, Van Elsacker L, Ganzhorn JU In: Fruits and frugivores, L Dew and JP Boubli (Eds.) In Press. Kluwer Academic Press, Dordrecht 5. The Malagasy littoral forest: threats and possible solutions ............................ 133 Bollen A, Donati G Table of contents III. General conclusion.............................................................................................. 147 Samenvatting.............................................................................................................. 153 Sommaire ................................................................................................................... 159 References ................................................................................................................ 165 Acknowledgements ACKNOWLEDGEMENTS Many people have provided me with assistance, guidance and support during the time it has taken me to complete this PhD for which I am very grateful: First of all I want to thank my supervisor, Linda Van Elsacker, who has supported me from the beginning of this PhD, never losing faith. Thanks for letting me introduce not only lemurs but also bats, birds and rats within this primatological group and for always being ready to read my manuscripts over and over again, to exchange ideas and to motivate me to present research to the larger public. Sincere thanks as well to Rudi Verheyen who was ready to act as my co-promoter and who has encouraged me throughout the PhD. It was Jörg Ganzhorn who got me to Madagascar in the first place. He helped me greatly during field research, in the analyses and written part of this PhD. Many of the interpretations presented here were developed in discussion with him. His impressive list of publications and unique expertise in tropical ecology have been an ideal guide for me during these four years. Jörg, thank you very much for the appreciated help and all the time you put into this work while not really having any spare time. I would also like to express my gratitude to all my CRC colleagues on whom I could count for correcting ‘rough’ drafts and providing useful suggestions. In particular I would like to thank Catherine Roden, Jeroen Stevens, Adinda Sannen, Rebekka Deleu, Isra Deblauwe, Hilde Vervaecke and Kristin Leus. My special thanks to Adinda who helped me greatly in the final phase, guiding me through all the official procedures and lay-out options. Many thanks as well to Vera Cuypers and Paule Stevens who made all the administrative paperwork a lot easier. Language-wise I was very lucky to get help from Eric Arnhem who was ready to correct the French summary of this PhD. As for English, it was wonderful to work together with Don and Val Sibley who closely scanned my drafts for awkward phrases, grammatical errors and other mistakes. Previous versions of the chapters benefited greatly from the comments of Steve Goodman and Paul Adrian Racey, for which many thanks. I am very grateful to Irene Tomaschewski from the Department of Conservation and Ecology (University of Hamburg) for carrying out biochemical analyses on the fruits. Thanks to the statistical advice of Dries Bonte and Stefaan Van Dongen SAS became a little less incomprehensible to me. They put me back on the right track when I was lost in the complex world of statistical tests and models. My sincere thanks for your help! I would further like to thank the members of the jury who have taken time to read through this PhD, I am looking forward to your comments. In this respect I am very grateful for the useful suggestions and constructive criticism of Prof Luc Lens on the first version of this PhD. Obviously this PhD would never have been possible in the first place without the funding of the FWO (Funds for Scientific Research, Flanders), for which many thanks. I further thank the Centre for Research and Conservation (Antwerp Zoo) and the Department of Biology (University of Antwerp) as well as DOCOP (University of Antwerp) for additional funding. 1 Acknowledgements A special word of thanks to the whole team of QMM in Fort-Dauphin for welcoming me at their study site and for helping me out with transport, radio-contact and food. Thanks to Rufen for the ‘architectural’ innovations at the campsite, which greatly improved our living standards. Laurent Randrihasipara and Jean-Baptiste Ramanamanjato, thanks for putting up with ‘la belge exigeante’ and Manon Vincelette, thank you for always taking time to answer my numerous questions. Having a degree in Zoology, it was great to be surrounded by excellent botanists familiar with the Malagasy flora. Petra De Block was the first to introduce me to the existing literature on taxonomy of Malagasy plants. Johny Rabenantoandro and Faly Mbolatiana from the Missouri Botanical Garden managed to replace most ‘vernacular’ names into scientific ones, which is not at all simple! Thanks for your patience in trying to guide me through the herbaria at Tana. Prof. Berthe Rakotosamimanana (Dept. Anthropologie) and Prof Daniel Rakotondravony (Dept Biologie Animale) of the University of Antananarivo helped me to obtain research permits and saved me from weeks of administrative hassle, for which my sincere thanks. I further acknowledge ANGAP, the Ministère des Eaux et Forêts and the Commission Tripartite of the Malagasy government for their collaboration and permission to work in Madagascar. The great help of my field assistants was essential in my
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