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The Functioning of Coral Reef Communities Along Environmental Gradients

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The Functioning of Coral Reef Communities Along Environmental Gradients The Functioning of Coral Reef Communities Along Environmental Gradients Ph.D. Thesis Jeremiah Grahm Plass-Johnson Dissertation zur Erlangung des Doktorgrades der Natuwissenschaften der Universität Bremen, Fachbereich Biologie/Chemie. Die vorliegende Arbeit wurde in der Zeit von Juni 2012 bis Mai 2015 am Leibniz-Zentrum für marine Tropenökologie in Bremen angefertigt. Finanziert wurde die Arbeit über das Bundesministerium für Bildung und Forschung (Grant no. 03F0643A) im Rahmen des bilateralen Deutsch-Indonesischen Projekts, Science for the Protection of Indonesian Coastal Ecosystems (SPICE III). Gutachter: Prof. Dr. Kai Bischof (Erstgutachter) Prof. Dr. Matthias Wolf (Zweitgutachter) Prüfer: Prof. Dr. Claudio Richter Dr. Mirta Teichberg Weitere Mitglieder des Prüfungsausschusses Jasmin Heiden (Doktorand) Tom Vierus (Student) Datum des Promotionskolloquiums: 21. Juli 2015 Summary One of the primary challenges in ecology is to understand how environmental disturbance affects diversity and community structure, and what are the subsequent consequences on ecosystem functioning. Coral reefs are some of the most diverse ecosystems on the planet resulting in complex sets of interactions between benthic, habitat-forming constituents and mobile fish consumers. However, scleractinian corals, the primary habitat engineers, are dependent on high-light, low- nutrient water conditions and thus are highly responsive when the environment varies from this status. In Southeast Asia, an increase in human coastal populations centred around urban areas has resulted in extensive changes to the coastal environment such as degraded water quality and removal of fish consumers. This has resulted in highly varied abiotic and biotic conditions in relation with distance from the shore. Often, coral reefs closer to shore are much lower in benthic and fish diversity than those further from anthropogenic influences, with direct impacts on ecosystem functioning. Therefore the aim of this thesis was to explore coral reef ecosystem functioning with respect to changes in benthic community structure and fish diversity in relation to varying environmental conditions in the Spermonde Archipelago, Indonesia. A combination of observational, experimental and theoretical analyses were conducted on the functioning of coral reefs using eight islands on a transect of increasing distance from the mainland, varying from 1 to 55 km. At these eight sites, benthic and pelagic surveys identified variation in the status of coral reef communities, while recruitment and feeding assays identified variation in important ecological processes. Lastly, experimental observations were further qualified with stable isotope analysis and the application of contemporary indices of functional diversity. It was found that indeed, the coral reefs varied along a continuum of structure, assemblage and processes. Increasing distance from shore was associated with greater live coral cover and structural complexity, while sites closer to shore were dominated by turf algae and rubble. Furthermore, turf algae was observed as playing a particularly important role, as this group was dominant during recruitment and subsequent development of open benthic space as supplied by terracotta tiles. Fish diversity, along with redundancy in the important herbivore group, also increased with distance from shore, resulting in an increasingly diverse response to Sargassum and Padina assays. The functional composition of the fish assemblages became increasingly variable with loss in coral cover and structural complexity, suggesting communities become destabilised under habitat degradation. Furthermore, stable isotope analysis indicated that the trophic niche of a fish species can increase at sites with more degradation suggesting varying functional utility. However, functioning is not determined only by exposure to chronic, abiotic conditions. Outbreaks of the crown-of-thorns starfish (Acanthaster planci) and mechanical destruction (bomb fishing) resulted in extreme loss of live coral. At these sites, biological and functional diversity displayed some of the lowest values among all sites. Coral reefs can exist in systems with altered water condition if physiological and ecological capacity of the organisms allow for their continuation. Nevertheless, degraded water condition will select against many species, resulting not only in the observed lower biological diversity, but also in less species taking part in functional roles as reflected in higher functional variability. Combined, these results show that the functioning of coral reefs does not exist in discrete states; rather, their functioning is a result of abiotic stressors and biological feedbacks. It is becoming increasingly clear that pristine coral reefs are not a reality in many cases around the world. Thus understanding coral reef functioning at all stages of degradation will help with future management. This thesis adds to the ever-growing knowledge about disturbed coral reefs, but more importantly, it describes the changing relationship between diversity and functioning of coral reefs in relation to disturbance. V Zusammenfassung Eine der primären Herausforderungen der Ökologie ist die Frage, inwiefern Umweltveränderungen die Diversität und Zusammensetzung von Artengemeinschaften beeinträchtigen, und welches die daraus resultierenden Auswirkungen auf die Funktionsweise von Ökosystemen sind. Korallenriffe gehören weltweit zu den artenreichsten Ökosystemen und bilden ein komplexes Beziehungsnetz zwischen habitatbildenden, benthischen Komponenten und mobilen Fischgemeinschaften als Konsumenten. Riffbildende Steinkorallen fungieren als die wichtigsten habitatbildenden Ökosystem- Ingenieure, benötigen hohe Lichtverhältnisse und sind besonders gut an die niedrigen Nährstoffbedingungen in tropischen Gewässern angepasst, weshalb sie sehr empfindlich auf Umweltveränderungen reagieren können. Marine Ökosysteme in Südostasiens leiden stark unter der stetig wachsenden Bevölkerung, die vorallem in urbanen Ballungsräumen im Küstenbereich meist eine drastische Abnahme der Wasserqualität und durch Überfischung einen Verlust an Kosumenten in der Nahrungskette mit sich bringt. Mit relativem Abstand von der Küste geht eine deutliche Veränderung in abiotischen und biotischen Wasserbedingungen einher, wodurch die benthische sowie pelagische Biodiversität küstennaher Korallenriffe oft weitaus geringer ist als die von küstenfernen Riffen. Das Ziel dieser Dissertation war daher, in dem indonesischen Spermonde- Archipel die Funktionsweise von Korallenriffen hinsichtlich der Struktur der benthischen Artengemeinschaft und der Fischbiodiversität unter verschiedenen Umweltbedigungen zu untersuchen. Dafür wurden Beobachtungsstudien sowie experimentelle und theoretische Analysen an Korallenriffen von acht verschiedenen Inseln durchgeführt, die in zunehmender Entfernung von der Küstenlinie liegen (1km bis 55km). Die benthischen und pelagischen Bestandsaufnahmen zeigten für alle acht Riffe einen unterschiedlichen Zustand der Artengemeinschaft, während Besiedelungs- und Fütterungsexperimente Unterschiede in wichtigen ökologischen Prozessen verdeutlichten. Diese experimentellen Beobachtungen wurden anhand stabiler Isotopenanalysen und der Anwendung von kürzlich entwickelten Indizes bezüglich funktioneller Diversität weiter validiert und es konnte gezeigt werden, dass die Korallenriffe tatsächlich entlang eines Kontinuums von Struktur, Zusammensetzung und Prozessen variieren. Mit größerer Entfernung von der Küste erhöhte sich die Bedeckung mit lebenden Korallen sowie die strukturelle Komplexität des Riffs, während Riffe näher an der Küste eindeutig von Aufwuchsalgen und Korallenschutt dominiert waren. Aufwuchsalgen waren die dominanten Neubesiedeler im Riff und reduzierten damit die Fläche an frei besiedelbarem Substrat, was sich in Besiedelungsexperimenten mit im Riff ausgebrachten Terracotta-Platten zeigte. Fischdiversität und funktionelle Redundanz innerhalb der wichtigen Gruppe herbivorer Fische erhöhte sich mit wachsender Entfernung von der Küste und resultierte in einer zunehmend diversen Reaktion in Fraßexperimenten mit Sargassum und Padina. Die funktionelle Zusammensetzung der Fischgemeinschaft war verstärkt variabel je geringer die Korallenbedeckung und die strukturelle Komplexität des Habitats war, was eine Destabilisierung der Gemeinschaft unter Verschlechterung des Habitatzustandes anzeigt. Desweiteren zeigte die Analyse von stabilen Isotopen, dass sich mit zunehmender Degradierung der Riffe die trophische Nische von Fischarten erweitern kann, was auf eine veränderte funktionelle Rolle der Fischart schließen lässt. Die Funktionsweise von diesen Korallenriffen ist jedoch nicht nur von chronischen, abiotischen Faktoren beeinflusst. Eine Massenvermehrung von Dornenkronenseesternen (Acanthaster planci) und mechanische Zerstörung (durch Dynamitfischerei) führte in einigen Riffen zu einem erheblichen Rückgang der Korallenbedeckung. Diese Riffe wiesen die geringste biologische und funktionelle Diversität auf. Korallenriffe können durchaus unter sich verändernden Umweltbedingungen überleben, solange die physiologische und ökologische Belastbarkeitsgrenze von Organismen nicht überschritten wird. VII Dennoch verursachen Umweltveränderungen eine Selektion vieler Arten, was zu der beobachteten Biodiversitätsabnahme führt, und da die einzelnen funktionellen Rollen folglich von weniger Arten wahrgenommen werden erhöht sich die funktionelle Variabilität des Systems. Zusammenfassend zeigen
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