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General Introduction 1 Zurich Open Repository and Archive University of Zurich Main Library Strickhofstrasse 39 CH-8057 Zurich www.zora.uzh.ch Year: 2006 Ecology, evolution, and conservation of plant-animal interactions in islands Hansen, Dennis Marinus Abstract: SUMMARY In my thesis I studied aspects of ecology, evolution, and conservation of plant- animal interactions on islands. My main study site was the island of Mauritius in the Indian Ocean. Mauritius is a biodiversity hotspot, with many endemic plant and animal species and high rates of ex- tinction. Chapter 1 reviewed the global distribution, ecology and evolution of coloured nectar, a rare floral trait that is particularly widespread on islands (including Mauritius) and insular mainland habitats such as mountains. Once thought to be restricted to three endemic plants in Mauritius, we showed that this is not the case: coloured nectar is found in more than 60 species from many plant families around the world. We also discussed the evolution of coloured nectar, and speculated on its ecological function. In Chapter 2, we experimentally tested a hypothesis from Chapter 1 about the possible ecological function of coloured nectar as a signal for floral reward. We used endemic flower-visiting geckos in Mauritius as our studyor- ganism, and found strong support for the signal-hypothesis, with geckos strongly preferring coloured over clear nectar. Thus, the chapter – at least partly – solved the mystery of the Mauritian coloured nectar. Chapter 3 demonstrated that the endemic Mauritian plant Trochetia blackburniana (Malvaceae) is pol- linated by the endemic Phelsuma cepediana gecko, but that this interaction is structured by the indirect effects of proximity to patches of Pandanus (Pandanaceae) plants – a favoured microhabitat of thegeckos. Proximity to Pandanus patches lead to higher gecko visitation rates and a subsequently higher fruitset in T. blackburniana. Some studies have shown how two or more flowering plant species can positively or negatively affect each other’s reproductive success through indirect effects mediated by shared pollinators. In contrast, Chapter 3 gives a unique example of a non-flowering plant affecting the reproductive success of a neighbouring flowering plant. Chapters 4 and 5 together formed a detailed study of the pollination and the seed dispersal ecology of the critically endangered endemic Mauritian tree Syzygium mamillatum (Myrtaceae). Chapter 4 showed how weeding of invasive plant species can influence the reproductive success of S. mamillatum in the weeded habitat, based on differences in pollinator behaviour between weeded and unweeded sites. In Chapter 5, we provided the first experimental evidence of the importance of the Janzen-Connell model for seedling establishment on oceanic islands, and demonstrated how eco- logical analogue species can be used to resurrect extinct seed dispersal interactions. Chapters 6 and 7 comprised a study of the pollination and seed dispersal interactions of another critically endangered Mau- ritian endemic plant, Roussea simplex (Rousseaceae), and how an invasive ant affects both interactions detrimentally. Chapter 6 showed that endemic Phelsuma cepediana geckos currently are the sole pollina- tors and seed dispersers of R. simplex, and Chapter 7 experimentally demonstrated that a presence of the invasive ant Technomyrmex albipes at R. simplex flowers or fruits scare away the geckos, thus rendering the plant without pollinators and seed dispersers. In Chapter 8 we documented the strong negative effects of a coffee pest species on the reproductive success of the endangered Mauritian endemic plant Bertiera zaluzania (Rubiaceae). Our study highlighted another perspective to the ongoing scientific debate about coffee as a cash crop and the maintenance of biodiversity in the tropics. Most current studiesfocuson the benefits that coffee plants can derive from nearby natural habitats, and neglect to investigate thepo- tential detrimental effects of coffee pest species invading these natural habitats. ZUSAMMENFASSUNG In dieser Dissertation präsentiere ich Studien, die sich mit der Ökologie und der Evolution, als auch mit Aspekte des Naturschutzes und der Erhaltung der biologischen Vielfalt von Tier-Pflanze-Interaktionen auf ozeanischen Inseln befassen. Mein Hauptuntersuchungsstandort ist die Insel Mauritius im Indischen Ozean. Mauritius besitzt einen ausgeprägten Endemismus und eine hohe Aussterberate und wird da- her als ‘Biodiversitäts-Hotspot’ bezeichnet. Kapitel 1 umfasst ein Review über die globale Verbreitung farbigen Nektars, eines seltenen Blütenmerkmals, das besonders häufig in Pflanzen auf Inseln (u.a. auf Mauritius) und in insulären Habitaten (wie z. B. Bergspitzen) anzutreffen ist. Wir zeigten, dass dieses Blütenmerkmal in mehr als 60 Arten aus vielen verschiedenen Familien rund um den Globus vertreten ist. Wir diskutierten die Evolution farbigen Nektars und spekulierten über mögliche ökologische Funktionen. In Kapitel 2 untersuchten wir mit Hilfe eines Experiments eine aus Kapitel 1 abgeleitete Hypothese: farbiger Nektar dient als ein Signal für das Vorhandensein von Blütenprodukten. Um diese Hypothese zu testen, offerierten wir endemischen Taggeckos in Mauritius gefärbten und klaren Nektar. Die Wahlfast ausschliesslich gefärbten Nektars unterstützt die Signal-Hypothese. Unsere Veröffentlichung trägt dazu bei, das Geheimnis um mauritischen farbigen Nektar zu lüften. Wir beweisen in Kapitel 3, dass die en- demische Pflanzenart Trochetia blackburniana (Malvaceae) von der endemischen Taggeckoart Phelsuma cepediana bestäubt wird. Diese Tier-Pflanze-Interaktion wird jedoch durch indirekte Effekte beinflusst, die im Zusammenhang mit den benachbarten Pflanzenarten der Gattung Panadanus (Schraubenbaum; Pandanaceae) stehen. Pandanusarten gehören zu den geeigneten Lebensräumen der Geckos, und T. blackburniana konnte höhere Besuchsraten und einen höheren Fruchtansatz verzeichnen, wenn es in der Nähe von Pandanus wuchs. Einige Studien konnten zeigen, dass der Fortpflanzungserfolg zweier be- nachbarte Pflanzenarten indirekt, durch gemeinsame Bestäuber, von der Anwesenheit der zweiten Art beeinflusst werden kann. Im Gegensatz dazu zeigten wir in Kaiptel 3 das einmalige Beispiel, dasseine nicht-blühende Pflanze eine blühende Pflanze indirekt beeinflusst. Kapitel 4 und 5 umfassten detaillierte Studien über die Bestäubungsbiologie und Mechanismen der Samenverbreitung der stark gefärdeten en- demischen Baumart Syzygium mamillatum (Myrtaceae). Kapitel 4 beschreibte, wie sich das Entfernen von eingeführten, invasiven Pflanzenarten auf den Fortpflanzungserfolg von S. mamillatum in restauri- erten Gebieten auswirkt, ein Unterschied, der auf das Verhalten von Vögeln als Bestäuber zurückgeführt werden kann. In Kapitel 5 lieferten wir den ersten experimentellen Beweis für die Wichtigkeit des Janzen- Connell Modells über die Etablierung von Keimlingen auf einer ozeanischen Insel. Zudem demonstrierten wir, dass analoge Arten, die in ihrer ökologischen Funktion den ehemaligen, jetzt ausgestorbenen Arten nahestehen, als geeignete Samenverbreiter von S. mamillatum in Frage kommen. Die Kapitel 6 und 7 beschrieben Bestäubung und Samenverbreitung der stark gefährdeten, endemischen Pflanzenart Roussea simplex (Rousseaceae), und wie eine eingeführte Ameisenart negative Folgen auf die Reproduktion dieser Pflanze haben kann. In Kapitel 6 zeigten wir, dass der endemische Taggecko Phelsuma cepediana der einzige Bestäuber und Samenverbreiter von R. simplex ist. Kapitel 7 hingegen beweist experimentell, dass die Präsenz der invasiven Ameisenart Technomyrmex albipes auf Blüten und Früchten von R. simplex Taggeckos verscheucht, was zu einer Reduktion der Bestäubung und Samenverbreitung bei R. simplex geführt hat. In Kapitel 8 dokumentieren wir einen erheblichen, negativen Einfluss eines Kaffeeschädlings auf den Fortpflanzungserfolg der gefährdeten mauritischen Pflanzenart Bertiera zaluzania (Rubiaceae). Unsere Arbeit trägt eine weitere Perspektive zu der anhaltenden, wissenschaftlen Debatte bei, die sich mit dem Einfluss von Kaffeeplantagen auf die Erhaltung der biologischer Vielfalt in den Tropen beschäftigt. Die meisten Studien befassen sich mit den Vorteilen des Kaffeeanbaus in der unmittelbaren Umgebung von natürlichen Habitaten, wenige Arbeiten jedoch untersuchen die nachteiligen Effekte, die eingeführte Kaffeeschädlinge auf den benachbarten Lebensraum ausüben können. Posted at the Zurich Open Repository and Archive, University of Zurich ZORA URL: https://doi.org/10.5167/uzh-163565 Dissertation Published Version Originally published at: Hansen, Dennis Marinus. Ecology, evolution, and conservation of plant-animal interactions in islands. 2006, University of Zurich, Faculty of Science. 2 ECOLOGY, EVOLUTION, AND CONSERVATION OF PLANT–ANIMAL INTERACTIONS ON ISLANDS DISSERTATION ZUR ERLANGUNG DER NATURWISSENSCHAFTLICHEN DOKTORWÜRDE (DR. SC. NAT.) VORGELEGT DER MATHEMATISCH-NATURWISSENSCHAFTLICHEN FAKULTÄT DER UNIVERSITÄT ZÜRICH VON DENNIS MARINUS HANSEN AUS DÄNEMARK PROMOTIONSKOMITEE PROF. DR. CHRISTINE B. MÜLLER (VORSITZ) PROF. DR. ELENA CONTI PROF. DR. LUKAS KELLER PROF. DR. H. PETER LINDER ZÜRICH 2006 TO MY PARENTS, WHO LET ME RUN AND GET MUDDY IN THE FORESTS OF MY CHILDHOOD “I trust that these, and all other friends who have been in any way interested in my travels and collections, may derive from the perusal of my book, some faint reflexion of the pleasures I myself enjoyed amid the scenes and objects it describes.” – Wallace, ‘Malay Archipelago’, 1869 CONTENTS
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