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Thesis by Michelle-Nicole Havlik Effect of Anthropogenic Noise on Reef Macroinvertebrates (Lambis lambis and Tridacna maxima). Thesis by Michelle-Nicole Havlik In Partial Fulfillment of the Requirements For the Degree of Master of Marine Science King Abdullah University of Science and Technology Thuwal, Kingdom of Saudi Arabia December, 2020 2 EXAMINATION COMMITTEE PAGE The thesis of Michelle-Nicole Havlik is approved by the examination committee. Committee Chairperson: Carlos Duarte Committee Members: Francesca Benzoni, Daniele Daffonchio 3 © February, 2021 Michelle-Nicole Havlik All Rights Reserved 4 ABSTRACT Effect of Anthropogenic Noise on Reef Macroinvertebrates (Lambis lambis and Tridacna maxima). Michelle-Nicole Havlik Boat noise has been shown to distract and cause harm to many marine organisms. Most of the study effort has been focused on fish & marine mammals, even though invertebrates represent 99% of all living organisms, both terrestrial and aquatic. The small amount of studies conducted on invertebrates have demonstrated a clear negative correlation between anthropogenic noise pollution and invertebrate well- being. The small giant clam Tridacna maxima and the spider conch Lambis lambis are two invertebrate species which play key roles in coral reef ecosystems, and are little studied for the effects of noise disturbance. If their behaviour is significantly affected by boat noise, this could cause a knock on effect on the rest of the reef ecosystem. T. maxima functions as prey to many fish species, contributing up to 9% of the reef’s calcium carbonate budget, as well as playing a role in nutrient cycling. L. lambis is essential prey for molluscivore snails and certain elasmobranchs, and through its herbivorous feeding, helps to keep filamentous algae levels low. Moreover, coral reefs are reliant on the biological sound primarily created by invertebrates, such as snapping shrimp, to attract larvae from the pelagic environment for settlement. During experiments conducted in February 2020 in the Red Sea, Daily Diary Monitoring (DDMT) smart tags were used to measure the reactions of T. maxima and L. lambis respectively during underwater playback of boat noise compared with ambient reef sound. It was 5 found that both T. maxima and L. lambis exhibited behavioral changes during the boat noise treatment. The implications of this study is that individuals of L. lambis and T. maxima may spend energy averting the invisible "threat" of boat noise, rather than feeding and/or photosynthesizing, in the case of T. maxima. If this hinders the acquisition of food/production, the whole reef ecosystem may be compromised in a knock-on effect from these producers and primary consumers. As boat noise is prevalent on inshore Red Sea reefs, shown by a 24-hour acoustic analysis of a patch reef sound pressure levels (SPL), invertebrates might be affected on a large scale in the Red Sea. 6 ACKNOWLEDGEMENTS Thank you to the Sound-team Nathan Geraldi (KAUST), Lloyd Hopkins (Swansea University), Jeroen Hubert (Leiden University), Lucille Chapuis (Exeter University), & Leigh Gaffney (University of Victoria), and their respective supervisors (Rory P. Wilson, Hans Slabbekoorn, Steve Simpson and Francis Juanes) for development of methods and participation in experiments, and making this the best learning experience for me! A great thank you to Carlos Duarte, my supervisor for his co-ordination of an international team, help and masterful guidance. I would also like to thank my committee chair, Prof. Carlos Duarte, and my committee members, Prof. Francesca Benzoni and Prof. Danielle Daffoncio, and their helpful comments. My appreciation also goes to my friends and colleagues and the Red Sea Research Centre faculty and staff for making my Masters at King Abdullah University of Science and Technology a great experience. I also want to extend my gratitude to those who came on the many fieldwork days with me: Claire Shellem, Kheng Lah Kim, Megan Nolan, Jessica Menzies, & Nikos Papagiannopoulos, I couldn’t have done it without you. Finally, my heartfelt gratitude is extended to my family & friends for their endless encouragement and support from afar. 7 TABLE OF CONTENTS Page EXAMINATION COMMITTEE PAGE .................................................................. 2 COPYRIGHT PAGE ………………………................................................................... 3 ABSTRACT....................................................................................................... 4 ACKNOWLEDGEMENTS …………………................................................................ 6 TABLE OF CONTENTS....................................................................................... 7 LIST OF ABBREVIATIONS.................................................................................. 8 LIST OF ILLUSTRATIONS................................................................................... 9 LIST OF TABLES .............................................................................................. 10 Introduction ………………………………………………………………………..…………………….. 11 1.1 Sound and Marine Life ………………………………………………………………… 11 1.2 The Soundscape of the Anthropocene Ocean ……………………………… 12 1.3 The Red Sea Soundscape ……………………………………………………………… 14 1.4 Previous Studies of T. maxima and L. lambis ………………………………… 16 Objectives ………………………………………………………………………..………………..……… 22 2.1 Aims ……………………………………………………………………………………………… 22 2.2 Methods ……………………………………………………………………………….………. 23 Results ………………………………………………………………………..……………………………… 33 3.1 Patch Reef Soundscape of the Central Red Sea …………………………….. 33 3.2 Movement frequency for L. lambis and T. maxima …………………….… 36 3.3 Vector of Dynamic Body Acceleration (VeDBA) for L. lambis ………… 39 3.4 Movement direction of L. lambis …………………………………………………… 40 3.5 Statistical analysis of responses to experimental playback ……………. 43 Discussion ………………………………………………………………………..……………….……….. 44 4.1 Soundscapes of Red Sea reef ecosystems ……………………………………… 44 4.2 L. lambis and boat playback ……………………………………………….…………. 47 4.3 Shelter/Movement direction for L. lambis …………………………………….. 49 4.4 Tridacna maxima and boat playback ……………………………………………… 50 8 LIST OF ABBREVIATIONS CEE Controlled exposure experiment CRS Central Red Sea DDMT Daily Diary Monitoring Tag VeDBA Vectorial dynamic body acceleration 9 LIST OF ILLUSTRATIONS Figure 1. Movement of T. maxima in response to threats .............................................. 19 Figure 2. Locomotion of L. lambis…………………………......................................................... 22 Figure 3. L. lambis tagged with DDMT............................................................................. 26 Figure 4. Experimental design of Controlled Exposure Experiments using play- back................................................................................................................................. 28 Figure 5. Median Sound Pressure Levels of playback tracks............................................................................................................................... 32 Figure 6. Spectral content of control and boat noise recordings & ambient conditions........................................................................................................................ 33 Figure 7. Broadband Sound Pressure Levels of an onshore Red Sea reef.................................................................................................................................. 35 Figure 8. Spectrogram images from an onshore Red Sea reef.................................................................................................................................. 37 Figure 9. Cumulative curve of average movements for L. lambis………...................................................................................................................... 38 Figure 10. Cumulative curve of average movements for T. maxima............................................................................................................................ 40 Figure 11. Average Vector of Dynamic Body Acceleration (VeDBA) (g) as measured by the Daily Diary Monitoring tag (DDMT) on L. lambis……………………………………………………………............................................................... 41 Figure 12. Directionality proportions of L. lambis movements during experimentation.............................................................................................................. 43 10 LIST OF TABLES Table 1. Study of movement direction of Lambis lambis, recording position at the end of the treatment, and sex) ................................................................................................. 59 Table 2. Summary statistics for T. maxima movement frequency, showing minute, average minimum value, first quartile, median, mean, third quartile and maximum value for the given time point ................................................................................................. 60 Table 3. Summary statistics for L. lambis movement frequency, showing minute, average minimum value, first quartile, median, mean, third quartile and maximum value for the given time point ............................................................................................................. 63 Table 2. Differences in movement frequency for 5 mins for T. maxima & L. lambis, calculated per 5 minutes (Kruskal Wallis rank sum test)................................................ 65 11 1. Introduction 1.1. Sound and Marine Life Sound propagates faster and farther underwater than in air, and farther than other
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