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Feeding Rates of a Large, Fast-Growing, Herbivorous Sea

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Feeding Rates of a Large, Fast-Growing, Herbivorous Sea FEEDING RATES OF A LARGE, FAST-GROWING, HERBIVOROUS SEA HARE, APLYSIA VACCARIA, ON BROWN ALGAE (PHAEOPHYCEAE) AND EXPLORATION OF SEAWEED CHEMICAL DEFENSE AND NUTRITIONAL CONTENT AS POTENTIAL DRIVERS A Thesis Presented to the Faculty of California State Polytechnic University, Pomona In Partial Fulfillment Of the Requirements for the Degree Master of Science In Biological Sciences By Danielle A. McHaskell 2019 SIGNATURE PAGE THESIS: FEEDING RATES OF A LARGE, FAST-GROWING, HERBIVOROUS SEA HARE, APLYSIA VACCARIA,ON BROWN ALGAE (PHAEOPHYCEAE) AND EXPLORATION OF SEAWEED CHEMICAL DEFENSE AND NUTRITIONAL CONTENT AS POTENTIAL DRIVERS AUTHOR: Danielle A. McHaskell DATE SUBMITTED: Summer 2019 Department of Biological Sciences Dr. Jayson R. Smith Thesis Committee Chair Biological Sciences Dr. Jeremy Claisse Biological Sciences Dr. Gregory A. Barding Chemistry ii ACKNOWLEDGEMENTS The African American aphorism “you’ve got to work twice as hard to get half as far” was a lesson I learned in childhood and a truth I learned that I had to live by in my adult years. Maintaining academic momentum to break ceilings and cycles, despite infrastructural barriers, has been an amazing accomplishment that I didn’t think was possible. Although my diligence and tenacity have been driving forces to myacademic success, I wouldn’t have made it this far without my community of support. Science does not occur without funding, so I want to thank all of my resources that made graduate school and my research possible. Thank you to COAST Undergraduate Research award, CPP Biological Sciences Department funding, MENTORES-PPOHA award, Dr. Steven J. Wickler Scholarship award and MBRS-RISE Program (Undergraduate and Graduate) Fellowship. MBRS-RISE Intensive Undergraduate Fellowship, in particular, financially supported me so that I could conduct research as opposed to working in a non-academic job to support myself. I’d like to thank my committee. Dr. Barding’s assistance in executing chemical assays increased my comfort, while decreasing wincing. Dr. Claisse not only convinced me that fish are exciting, despite being vertebrates, but helpedmegainan understanding in statistics and coding in R that I, dare I say, am enjoying. Dr.Smith(aka Aljay) has provided a plethora of lessons. I recall entering his office as a blue haired, starry-eyed undergraduate who was very determined to conduct research, with otters in particular, although verbally admitting to not quite understanding what that entailed. This led to volunteering in the intertidal zone with him and starting an undergraduate project that has culminated into this thesis. Thank you for showing me the beauty in algae and iii helping me develop as a scientist. I’d like to thank everyone who helped on this project! Shout out to the following: Brett Claire Arre, Karina Arzuyan, Anne Daniels, Seth Kapp, Christiana Kent, Leonard Ko, Amy Leonard, Veronica Magana, Haley Mai, Borman Quinonez, Elsa Perez, Cynthia Sandoval, Katie Stapel and Callum Theobold. Thank you for meeting me before dawn to go to Catalina, sitting through 4 hours of traffic with me to/from the field, getting sprayed with water while fixing the tanks and spending 6+ hours in the aquarium room on a Friday night sifting through sea hare poop looking for unconsumed algae. I’d like to thank the RISE administrators/cohorts, especially my academic moms: Dr. Buckley and Dr. Adler. Not only have you two encouraged and prepared me to apply and succeed in a doctoral program but have given immense emotional support by talking me through my problems and allowing a safe area to express what I was feeling. I’d like to thank Professor Mark Cooper, from my community college days at Mt. SAC, for being the first person to believe in me as a scientist and helpingmelearn what I am capable of. Bio 2 was one of the most challenging classes I’ve taken in college but showed me how much I love zoology/ecology and that I was meant for it (2 EASY!). Of course, I want to thank my family for supporting, although maybe not quite understanding, my choice to pursue academia and become a scientist. Pops, Amber, Izzy, Chris, Monica, Ferdinand and Gloria, I love all of you and am so thankful to have you all in my life. Also, thank you for letting me stand on my soap box and for listening to why single use plastic, overfishing and climate change are killing our ocean. I’dalsoliketo thank my other half and Fiancé, Leonard Ko. You have been a buttress of emotional, physical and academic support: from helping me collect algae at a 5:00 AM lowtideto iv holding a light up in lab while I worked in the dark after building lights shut off at midnight. Our adventure is out there, and I can’t wait to see what we do next together! I dedicate this thesis to my mom, who I lost right before the M.S. program, and our dog Koga, who I lost a few months ago. Mom, no matter how hard I was on myself, no matter how many times I said that I couldn’t do it, you were always a source of positivity and comfort. When I was young, you encouraged my academic pursuits and always told me that I would grow up to be an intelligent, successful woman. I feel immense pain from your passing but will do my best each and every day to prove you right. I love you Mom. v ABSTRACT In marine ecosystems, herbivory can play a vital role in structuring community composition and maintaining normal ecosystem functioning. Found in southern California intertidal and subtidal habitats, the Black Sea Hare, Aplysia vaccaria, has been described as a voracious and indiscriminate grazer on brown seaweeds (Phaeophyceae) in order to meet the energetic demands of growing to a remarkable ~14 kg within its 1-year life span. Despite the expected extremely high consumption rates, and possible large impact in driving seaweed community composition in local waters, little work has been conducted to measure their feeding rates or whether they consume some macroalgae faster than others. Therefore, the first goal of my study was to measure and compare the feeding rates of the Black Sea Hare when offered eight brown seaweeds commoninthe herbivore’s habit. As documented in studies with other, often smaller herbivores, consumption rates and seaweed diet selectivity may be driven by characteristics that vary among algal species, such as morphology, structural or chemical defenses (e.g. phlorotannins), and nutritional content (e.g. carbon to nitrogen ratio). For example, seaweed species with high nutritional content and low chemical defenses often are consumed at higher rates than heavily defended and/or poor quality macroalgae. However, these patterns are highly inconsistent with, for example, herbivore mobility/size and linkage between seaweed as home/refuge and diet affecting the relative role of seaweed chemical characteristics. The secondary goal of this study, therefore, was to elucidate the potential role of seaweed chemical composition of experimental seaweeds in driving the relative feeding rates of A. vaccaria, focusing on phlorotannin concentrations, C:N, and tissue nitrogen content. vi The feeding rates of A. vaccaria on eight brown seaweed species were determined in laboratory-controlled experiments in a running seawater aquaria system through a series of single-food consumption trials with consumption compared among taxa. A variety of seaweeds were chosen with expected variation in chemical characteristics, including three kelps (Laminariales) and five Fucoids, consisting of two rockweeds and 3 Sargassum species. Phlorotannin concentration for each seaweed sample was determined using a standard Folin-Ciocalteau assay while elemental analysis was used to determine nutritional content (nitrogen and C:N). Consumption rates by the Black Sea Hare of eight common brown seaweeds were relatively similar among taxa, suggesting that the herbivore is an indiscriminate, generalist herbivore on brown macroalgae, as previously described. Feeding rates did not appear to be extensively driven by seaweed chemical defenses (phlorotannin concentrations) nor nutritional quality (C:N and tissue nitrogen content). Although total consumption by individual A. vaccaria was markedly higher than that of other common herbivores in the region, this was likely driven by the large size of the herbivore; the feeding rate standardized by biomass of herbivores was relatively similar among herbivore taxa, suggesting that the Black Sea Hare is not as voracious of a feeder as originally believed. However, consumption of primary producers by A. vaccaria could potentially play a more crucial role in structuring of intertidal and shallow subtidal coastal algal communities due to the high biomass consumed by an individualand that this species often forms dense aggregates during reproductive periods. vii TABLE OF CONTENTS Signature Page..................................................................................................................... ii Acknowledgements............................................................................................................ iii Abstract .............................................................................................................................. vi List Of Tables ..................................................................................................................... x List Of Figures ................................................................................................................... xi 1. Introduction....................................................................................................................
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