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Sponge Bioerosion and Habitat Degradation on Indonesian Coral Reefs

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Sponge Bioerosion and Habitat Degradation on Indonesian Coral Reefs Sponge bioerosion and habitat degradation on Indonesian coral reefs by Joseph Marlow A thesis submitted to Victoria University of Wellington in fulfilment of the requirements for the degree of Doctor of Philosophy 2017 2 Acknowledgments Firstly I would like to thank my primary supervisor, Associate Professor James Bell, for his unwavering support and advice these past three years. I feel very lucky to have had James as my supervisor, his help and guidance whether it was in the field, in the lab or in relation to the many many manuscript drafts I sent him has always been fantastic. I would also like to thank my secondary supervisor, Professor Simon Davy, in particular for his advice about Symbiodinium and photophysiology but also for his overall support and excellent feedback on manuscripts. This research could not have happened without the funding and support from Operation Wallacea. I would like to thank in particular Pippa Mansell for her incredible management of the research station and thank both her and Chris Majors for all their support and help with my research. Thanks to all the Indonesian staff who kept me fed, in the water and made sure I always had a cold Bintang waiting for me at the end of the day. I am incredibly grateful for the support and funding provided by VUW, without which I would not have been able to complete this PhD. Thanks also to the PADI foundation which also provided research funding and Daniel LeDuc and Dennis Gordon at NIWA for their help and providing access to the SEM. Thank you to Hasanuddin University for facilitating this research. I would like to thank Dr Christine Schönberg and Associate Professor Ken Ryan and Dr Clint Oakley for their time and help. Thank you Christine for revealing the world of spicules and sponge taxonomy to me and your incredibly helpful feedback for my manuscripts. Thank you Ken for your ever present open door so I could come and pick your brain about PAM flourometry. Thank you Clint for answering all my many questions on Symbiodinium. Thanks to the amazing support group that is Sponge club, couldn’t have done it without you guys. Special thanks to everyone who helped me survive Hoga; Andy Biggerstaff, Emily MCGrath, Charli Mortimer, Meg Shaffer (world’s best RA), Tracey Bates and Holly Bennett. Finally I would like to thank my friends and family without whose continuing support I could never have achieved any of this. 3 Abstract Coral reefs are among the most diverse ecosystems on the planet, yet they are also sensitive to anthropogenic disturbances that can degrade these systems. On many degraded reefs, large increases in bioeroding sponge abundance have occurred. On healthy reefs these sponges contribute to species diversity and habitat complexity, however there is growing concern that their proliferation on degraded reefs could lead to a state of net-erosion. In the Southeast Asian Indo-Pacific, the ecology of bioeroding sponges in relation to coral degradation has been poorly studied compared to other coral reef regions. This thesis aims to increase our understanding of the ecology of these sponges in the Wakatobi region of Indonesia, and their likely trajectory if reefs continue to degrade in the region. My first research chapter aimed to identify the common bioeroding sponge species of the Wakatobi. This was achieved through in-water surveys, and subsequent spicule and phylogenetic analysis. This resulted in the identification of eight commonly occurring Wakatobi bioeroding sponge species, two of which are described for the first time. The assemblage composition was distinctly different from the only other bioeroding sponge study in Indonesian waters (Calcinai et al. 2005), highlighting the need for more clionaid taxonomic information from the region. Having identified the common bioeroding sponge species in the region, my second chapter assessed the major environmental drivers of the abundance and assemblage composition of these sponges. Abundance surveys were conducted at 11 reef sites characterised by different environmental conditions and states of reef health. Bioeroding sponges occupied 8.9% of suitable substrate, and differences in abundance and assemblage composition were primarily attributed to differences in the availability of dead substrate. However, abundance was lowest at a sedimented and turbid reef, despite abundant dead substrate availability. This indicates a limited resilience in some species to conditions associated with terrestrial run-off and that not all forms of reef degradation are beneficial for bioeroding sponges. The capacity to increase spatial occupation of degraded reefs is also dependent upon larval recruitment and my third chapter was a two year recruitment study using in situ experimental calcareous blocks. Recruitment occurred rapidly and consistently with bioeroding sponges recruiting to approximately 70% of experimental blocks and exhibiting a preference for settlement on uncolonised dead calcareous substrates. The importance of substrate settlement cues and extent of larval dispersal appeared to 4 differ between species, indicative of different recruitment mechanisms. Any significant increase in the availability of exposed calcareous substrate (e.g. following a mass coral bleaching event) is therefore likely to result in widespread increases in bioeroding sponge recruitment. Surveys conducted in my second research chapter revealed that two of the three locally abundant zooxanthellate bioeroding species were absent from a highly turbid reef, Sampela. My fourth research chapter investigated whether this was due to light limitation by examining the photoacclimatory capabilities of the Symbiodinium photosymbionts of Cliona aff. viridis n. sp. A. PAM chlorophyll fluorometry was employed in a 25 day shading experiment and Symbiodinium of C. aff. viridis n. sp. A demonstrated an ability to photoacclimate to extreme light reduction and recover quickly when conditions returned to normal. My results demonstrate that the absence of this species at Sampela is not due to light limitation but possibly due to other stressors associated with turbidity, e.g. suspended sediment. My final chapter was an assessment of the environmental drivers of rates of bioerosion in Spheciospongia cf. vagabunda, a common species in the Wakatobi. Erosion rates were determined from changes in dry-weight of calcareous substrates with attached grafts of S. cf. vagabunda after a year deployment across seven reef sites. The average bioerosion rate was 12.0 kg m-2 sponge tissue yr-1 (± 0.87 SE), but differed between sites and was negatively correlated with settled sediment depth. Bioerosion by this species can play a significant part in the carbonate budget on reefs where it is abundant (up to 20% of available substrate on some reefs in the Wakatobi) but is likely reduced on highly sedimented reefs. In summary, the Wakatobi bioeroding sponge assemblage is diverse and overall, both adult abundance and recruitment are primarily driven by the availability of dead calcareous substrates. Therefore, further coral mortality and a subsequent rise in the availability of dead substrate in the region is likely to result in increased abundance of bioeroding sponges. However, not all forms of reef degradation will benefit these sponges; turbid and sedimented reefs will likely constitute stressful habitats for some bioeroding sponge species and assemblages in these environments will be comprised of fewer more resilient species. 5 Table of Contents Chapter 1: Introduction ............................................................................................................. 17 1.1. Coral reefs ...................................................................................................................... 17 1.1.1. Diversity & importance........................................................................................... 17 1.2. Sponges on coral reefs.................................................................................................... 19 1.3. Reef bioerosion............................................................................................................... 20 1.4. Bioeroding sponges ........................................................................................................ 21 1.4.1. Taxonomy and morphology .................................................................................... 21 1.4.2. Sponge bioerosion ................................................................................................... 22 1.4.3. Symbiodinium ......................................................................................................... 25 1.5. Bioeroding sponge abundance & distribution ................................................................ 27 1.5.1. Reproductive output & recruitment ........................................................................ 27 1.5.2. Adult distribution .................................................................................................... 28 1.5.3. Competition............................................................................................................. 31 1.6. The Wakatobi ................................................................................................................. 32 1.7. Thesis aims ..................................................................................................................... 34 1.7.1. Thesis objectives ....................................................................................................
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