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Investigating the Disturbance of the Pugnacious Nudibranch Phidiana Hiltoni

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Investigating the Disturbance of the Pugnacious Nudibranch Phidiana Hiltoni PUSHING BOUNDARIES: INVESTIGATING THE DISTURBANCE OF THE PUGNACIOUS NUDIBRANCH PHIDIANA HILTONI A Thesis submitted to the faculty of San Francisco State University In partial fulfillment of the requirements for the Degree Master of Science In Marine Science by Emily Elizaoeth Otstott San Francisco, California August 2019 Copyright by Emily Elizabeth Otstott 2019 CERTIFICATION OF APPROVAL I certify that I have read Pushing Boundaries: Investigating the Disturbance of the Pugnacious Nudibranch Phidiana hiltoni by Emily Elizabeth Otstott, and that in my opinion this work meets the criteria for approving a thesis submitted in partial fulfillment of the requirement for the degree Master of Science in Biology: Marine Biology at San Francisco State University. Terrence Gosliner, PhD Research Professor, San Francisco State University Angel Valdes, PhD Professor, California State Polytechnic University, Pomona Katharyn Boyer, PhD Professor, San Francisco State University Pushing boundaries: Investigating the disturbance of the pugnacious nudibranch Phidiana hiltoni Emily Elizabeth Otstott San Francisco, California 2019 Warming ocean conditions sparked the poleward range expansion of the pugnacious nudibranch Phidiana hiltoni, whose arrival to northern California in 1992 coincided with significant declines of resident nudibranchs. The nudibranch assemblage of Duxbury Reef was monitored to assess current impacts of P. hiltoni. The density of P. hiltoni at Duxbury Reef has increased by three-fourths since 2010, now reaching over 18 ind. hr' 1 observer'1. The pooled density of nudibranchs impacted by P. hiltoni at Duxbury Reef (11 individuals h r 1 observer'1) has increased by 40% since 2008-2010 but still falls short of its density before P. hiltoni. Since first being disturbed by P. hiltoni in 1992, the nudibranch assemblage of Duxbury Reef is in recovery. Other aspects of this disturbance are discussed, such as prey overlap and interactions with specific resident nudibranchs. I certify that the Abstract is a correct representation of the content of this thesis. OJXjXq 'V - 12/20/2019 Chair, Thesis Committee Date ACKNOWLEDGEMENTS I would like to thank the Gosliner Lab for helping me develop my project and for supporting my tide pool efforts. I am indebted to my committee members, each of whom offered valuable ideas and steered me on my path. The Steinhart Aquarium at the California Academy of Sciences graciously made space and time to help me perform feeding trials. Others helped with this project in various ways and I would like them to know my appreciation: Joren Nisiewicz, Jeff Goddard, and Eric Sanford. TABLE OF CONTENTS List of Table................................................................................................................................. vii List of Figures.............................................................................................................................viii List of Appendices........................................................................................................................ix Introduction..................................................................................................................................... 1 Background........................................................................................................................ 1 The Problem.......................................................................................................................2 Project Goals......................................................................................................................3 Methods........................................................................................................................................... 5 Timed Counts.....................................................................................................................5 Feeding Overlap................................................................................................................ 8 Feeding Trials—Testing Hermissenda spp. against P. hiltoni.................................. 9 Results..............................................................................................................................................9 Timed Counts...................................................................................................................11 Feeding Overlap.............................................................................................................. 15 Feeding Trials—Testing Hermissenda spp. against P. hiltoni.................................16 Discussion..................................................................................................................................... 16 Reference......................................................................................................................................27 Appendices....................................................................................................................................34 LIST OF TABLES Table Page 1. Nudibranch species density at Duxbury Reef throughout time..........................10 2. Proportion of nudibranch species that tended to increase or decrease in density at Duxbury Reef from 2008-10 to 2018-19................................................................13 3. Reactions of P. hiltoni to each of the Hermissenda spp. present at Duxbury Reef............................................................................................................................16 vii LIST OF FIGURES Figures Page 1. Phidiana hiltoni with Northern California nudibranchs and prey...................... 3 2. Study sites of reefs in Northern California............................................................5 3. Nudibranch densities at Duxbury Reef................................................................. 11 4. Population growth of P. hiltoni, exhibiting exponential growth......................... 11 5. Number of individuals and species found at Duxbury Reef............................... 12 6 . Regressions of the annual densities of P. hiltoni on Hermissenda spp., F. trilineata, D. amyra, and A. abronia...........................................................................................12 7. Regression of the pooled density of impacted species against the density of P. hiltoni from 2007-19. 2007-11, and 2018-19.........................................................13 8 . Comparison of methods.............................................................................................15 9. Hydroid species and the nudibranchs that eat them at Duxbury Reef...............15 10. Observations of P. hiltoni from online sources reveal range shifts with phases of ENSO............................................................................................................................ 32 LIST OF APPENDICES Appendix Page 1. Range Shift with Phases of ENSO............................................................................. 32 ix 1 Introduction The natural flux of Earth’s climate is being exacerbated by human activities, resulting in modification of the geographic ranges of species 1_4. Climate change is primarily expressed as environmental warming, which is pushing the ranges of species poleward 3,5,6. While even slight changes in temperature can greatly impact species composition and abundance 7, the California coastal ecosystem has been enduring temperature anomalies up to 6 °C 6’8,9. These marine heat waves are becoming more frequent and intense, with species responses matching this intensity 10,1 Since wind and ocean currents distribute heat over the globe, altering the amount of heat in places will impact the patterns of currents. Sea surface temperature in the North Pacific positively correlates with the Pacific decadal oscillation (PDO), which alters the positions of atmospheric circulations and cloud cover 4’5,12. Increased ocean temperatures are also associated with the El Nino Southern Oscillation, which reverses the flow of the California Current from southward and offshore to northward and onshore 4. These modifications directly affect the geographic ranges of marine species, particularly those with planktonic larvae. Nudibranchs, or sea slugs, have planktonic larvae whose recruitment is affected by climatic phases that alter larval advection 4_6>13. For example, upwelling relaxation events, such as during El Nino, allow high densities of larvae to potentially be delivered to shore as surface flow reverses 4’5,14. During positive or warm phases of El Nino and the PDO, the abundances of “warm water” species increase and the geographic ranges of some species shift poleward 4’5,15. With warm PDO phases increasing the flow of coastal waters northward and El Nino causing the California Current to flow northward and onshore, the overlap of these events would greatly promote northward dispersal of nudibranch larvae 4,5. Such an overlap did occur a few years ago. with a warm PDO phase occurring during the 2014-16 El Nino, resulting in the largest marine heat wave ever recorded 16. Aside from producing a destructive warm-water “blob” off the coast of 2 California that extended 300 m deep, this heat wave resulted in the poleward movement of at least 26 nudibranch species, 11 of which dispersed to new northernmost sites 6,1?. Since these are cycles, the population would likely move back south as the warming period ends; however, with the additional rising temperatures of long-term climate
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