The Influence of Fragment Size on Biotic Interactions That Structure Plant Communities in the Asian Tropics

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The Influence of Fragment Size on Biotic Interactions That Structure Plant Communities in the Asian Tropics Research Collection Doctoral Thesis The Influence of Fragment Size on Biotic Interactions that structure Plant Communities in the Asian Tropics Author(s): Viswanathan, Ashwin Publication Date: 2018 Permanent Link: https://doi.org/10.3929/ethz-b-000267356 Rights / License: In Copyright - Non-Commercial Use Permitted This page was generated automatically upon download from the ETH Zurich Research Collection. For more information please consult the Terms of use. ETH Library DISS. ETH NO. 24963 THE INFLUENCE OF FRAGMENT SIZE ON BIOTIC INTERACTIONS THAT STRUCTURE PLANT COMMUNITIES IN THE ASIAN TROPICS A thesis submitted to attain the degree of DOCTOR OF SCIENCES of ETH ZURICH (Dr. Sc. ETH Zurich) Presented by Ashwin Viswanathan MSc Wildlife Biology and Conservation, National Centre for Biological Sciences (NCBS) Born on 19.06.1987 Citizen of India Accepted on the recommendation of Jaboury Ghazoul (examiner) Robert Bagchi (co-examiner) David Burslem (co-examiner) 2018 Summary Diverse rainforests in the tropics are being rapidly modified and fragmented for human use. Plants that persist in remnant forest patches are threatened by many biotic and abiotic changes that are associated with fragmentation. Several of these fragments consequently contain fewer plant species than they once did but, in the absence of contiguous forest, remain the only strongholds of plant and animal diversity in many parts of the world. However, the future of diversity in forest fragments may be uncertain as fragmentation can influence plant-animal interactions that shape plant communities. Some of these changes may already be apparent in the youngest life stages of fragmented plant communities, which would then form the template for the future of those fragments. By examining the compositions of differently aged plant communities in forest fragments, we may be able to identify interactions that are especially influenced by fragmentation, as various plant-animal interactions play structuring roles during different stages of a plant’s life cycle. In this thesis, we primarily investigate relationships between fragment area and processes that structure plant communities in a tropical forest. In the first data chapter (the second chapter of this thesis), we examine the compositions of four life stages of woody plants across a gradient of fragment size. We show that plant species are primarily affected by fragment size during their transitions from seeds to saplings. We discuss several possible explanations for the observed patterns but suggest that altered interactions between plants and their natural enemies (insects and fungal pathogens) may be particularly important drivers. Such plant-enemy interactions have the potential to maintain plant diversity by causing the negative density-dependent mortality of locally abundant plant species, and by allowing the persistence of locally rare species (the Janzen-Connell Hypothesis). As the modification of diversity-maintaining interactions can have catastrophic long-term consequences for the plant diversity in small fragments, it was important to investigate whether plant-enemy interactions are indeed sensitive to fragment size. i In the second data chapter (the third chapter of this thesis), we investigate the relationship between fragment area and (soil-borne) fungus-induced mortality of six woody plant species in a shadehouse experiment. We present evidence that the pathogenic effects of fungi on one plant species increased with increasing fragment size. Although we show that plant fungus- interactions can be influenced by fragment size, further experiments are required to investigate whether such effects (even when spread across the community) can influence the diversity maintained in a forest fragment. In the third data chapter (the fourth chapter of this thesis), we first examine the roles of insects and fungi in maintaining woody plant seedling diversity in an Indian rainforest. We then investigate whether the diversity-maintaining abilities of these natural enemies are influenced by fragment size. We present evidence that insects play important roles in maintaining gamma diversity primarily by suppressing common plant species independent of density. We show that they maintain more diversity in large fragments than in smaller fragments, as smaller fragments were dominated by insect-resistant species. We present evidence that fungi caused the density- dependent mortality of one plant species but infer the presence of more such interactions in the community. We show that fungi may be sensitive to fragment size, and that fungi maintain more beta diversity of woody plant seedlings in large fragments than in small fragments. In conclusion, I show in my thesis that insects and fungi play important roles in maintaining woody plant seedling diversity. I further show that these essential plant-enemy interactions are predictably influenced by the size of a forest fragment, and that the plant diversity in small fragments may consequently be at risk from the breakdown of such interactions. ii Zusammenfassung Die anthropogene Nutzung der natürlichen Ressourcen führt zu steigender Fragmentierung und Veränderung der diversifizierten Regenwälder in den Tropen. Die Fragmentierung verursacht viele biotische und abiotische Veränderungen welche die Pflanzen der verbliebenen Waldbeständen gefährdet. Daher beherbergen viele dieser Waldbestände weniger Pflanzenarten als früher als die Wälder noch zusammenhängend waren. Die verbliebenen Pflanzenarten in diesen Waldfragmenten bilden nun die einzige Basis für viele Pflanzen- und Tierarten in vielen Regionen dieser Welt. Die Zukunft der Diversität in Waldfragmenten ist jedoch ungewiss, da die Fragmentation die Interaktionen zwischen Pflanzen und Tieren beeinflusst welche die Pflanzengemeinschaft erst ermöglicht. Einige dieser Veränderungen könnten in den jüngeren Altersstadien von fragmentierten Pflanzengemeinschaften bereits sichtbar sein, und dienen somit als erste Einsicht in die Zukunft dieser Waldfragmente. Die Interaktionen zwischen Pflanzen und Tieren bilden wichtige strukturelle Funktionen verschiedener Stadien der Lebenszyklen der Pflanzen. Daher ermöglicht die Untersuchung der Zusammensetzung von Pflanzengemeinschaften verschiedener Altersstadien Rückschlüsse bezüglich der verändernden Interaktionen aufgrund der Fragmentierung. In dieser Doktorarbeit, untersuchen wir in erster Linie die Zusammenhänge zwischen der Fläche des Waldfragments und den Prozessen welche die Struktur der Pflanzengemeinschaft in einem tropischen Wald bedingen. Im ersten Datenkapitel (dem zweiten Kapitel dieser Doktorarbeit), untersuchen wir die Zusammensetzung von vier Altersstadien von Holzgewächsen entlang eines Fragmentgrössen- Gradienten. Gemäss den Daten, scheinen Pflanzenarten demnach vor allem durch die Fragmentgrösse während dem Übergang vom Samen zum Schössling beeinflusst zu sein. Wir diskutieren verschiedene mögliche Ursachen für die beobachteten Muster, behaupten jedoch, dass die veränderten Interaktionen zwischen Pflanzen und deren natürlichen Feinden (Insekten iii und Pilzpathogenen) besonders wichtige Ursachen sind. Diese Interaktionen zwischen Pflanzen und deren natürlichen Feinden ermöglichen die Pflanzenvielfalt aufrechtzuerhalten, da sie zu einer negativen dichteabhängigen Sterblichkeit lokal reichlich vorhandener Pflanzenarten führen, und damit lokal seltener Arten eine Chance geben zu gedeihen (Janzen- Connell-Hypothese). Da die Veränderung von dichte-aufrechterhaltender Interaktionen katastrophale langfristige Konsequenzen für die Diversität der Pflanzen kleiner Waldfragmente haben kann, war es wichtig zu untersuchen ob die Interaktionen der Pflanzen und deren Feinden tatsächlich empfindlich gegenüber Fragmentgrösse sind. Im zweiten Datenkapitel (dem dritten Kapitel dieser Doktorarbeit), untersuchen wir die Beziehungen zwischen Böden von Fragmenten unterschiedlicher Grösse und den (Boden verursachten) Pilz-bedingten Sterblichkeit von sechs Arten von Holzgewächsen in einem Schattenhausexperiment. Wir fanden Hinweise für den Anstieg von pathogenen Effekten von Pilzen auf eine der untersuchten Pflanzenarten mit steigender Fragmentgrösse. Obschon wir aufzeigen dass Pflanzen-Pilz Interaktionen von der Fragmentgrösse beeinflusst werden kann, sind weitere Experimente notwendig um herauszufinden ob solche Effekte (selbst wenn sie sich über die Pflanzengesellschaft verteilen) die aufrechterhaltene Diversität in einem Waldfragment beeinflussen kann. Im dritten Datenkapitel (dem vierten Kapitel dieser Doktorarbeit), untersuchen wir zuerst die Rolle von Insekten und Pilzen in der Aufrechterhaltung der Diversität von hölzernen Sämlingen in einem Indischen Regenwald. Weiter untersuchen wir ob die Diversitätserhaltenden Fähigkeiten von natürlichen Feinden von der Fragmentgrösse beeinflusst wird. Wir präsentieren Hinweise, dass Insekten eine wichtige Rolle spielen bei der Aufrechterhaltung der Gamma-Diversität, hauptsächlich aufgrund der Unterdrückung von üblichen Pflanzenarten, unabhängig der Dichte. Diese Aufrechterhaltung der Diversität ist ausgeprägter in grossen Fragmenten, da kleine Fragmente von Insektenresistenten Arten iv dominiert wurden. Wir fanden Hinweise, dass Pilze die dichteabhängige Sterblichkeit einer Pflanzenart verursachten, folgern jedoch, dass noch mehrere solche Interaktionen in der Gemeinschaft geben muss. Pilze scheinen sehr empfindlich gegenüber der Fragmentgrösse zu sein und unterstützen die Beta-Diversität von hölzernen Sämlingen mehr in grossen als in kleinen Fragmenten.
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