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An Abstract of the Dissertation Of AN ABSTRACT OF THE DISSERTATION OF Jenna Sullivan Stack for the degree of Doctor of Philosophy in Zoology presented on May 3, 2019. Title: Marine Mass Mortality in a Global Change Context: Impacts on Individuals, Populations and Communities Abstract approved: _____________________________________________________________________ Bruce A. Menge Human actions are pushing natural systems into states that have no historical precedent. In response, empirical and theoretical researchers are increasingly focused on developing ways to predict the responses of ecological systems to change. However, significant knowledge gaps remain, often leading to “ecological surprises” where observed impacts of global change do not align with existing theory or hypotheses. In this dissertation, I study the response to perturbations of a well- characterized system for ecological research, the Northeast Pacific rocky intertidal, to advance our understanding of and ability to predict the impacts of global change on individuals, populations and communities. In 2013 and 2014, sea star species along the west coast of North America were affected by an outbreak of Sea Star Wasting Syndrome (SSWS), resulting in an epidemic of mass mortality that spanned unprecedented geographic and temporal scales and resulted in the near extirpation of multiple sea star species from many locations along the coast. One of the species that was most strongly affected in the intertidal zone was Pisaster ochraceus, an iconic predatory sea star that has the ability to play a keystone role in its community through foraging on and ultimately controlling the lower boundary of mussel prey populations. The first two chapters of this dissertation take advantage of SSWS as a “natural” form of top predator removal to assess the consequences of this type of perturbation on ecosystem resilience. In Chapter 2, I tested the hypotheses that P. ochraceus loss would facilitate a population expansion of a smaller, mesopredator sea star, Leptasterias sp., and that this expansion would have negative effects on P. ochraceus population recovery. This result would follow expectations of competitive release and aligns with existing research on the competitive relationship between these species from the Northeast Pacific intertidal. I used field surveys to track Leptasterias populations just before the onset of and up to three years after SSWS. Contrary to expectation, I did not see an increase in the distribution or density of Leptasterias, and instead saw a decrease in individual size post-SSWS. Further, I found no evidence of competition between P. ochraceus recruits and Leptasterias for resources. Thus, although my hypotheses were grounded in theory and previous research, they were not supported by data. These results suggest that Leptasterias will not provide a bottleneck for P. ochraceus population recovery from SSWS, nor compensate for lowered P. ochraceus predation. The dynamics of P. ochraceus at the recruit (early benthic juvenile) life- history stage has long been considered a gap in our understanding of the species, as recruits have been historically rare in the intertidal and hard to study. Post-SSWS, however, many sites along the coast experienced unprecedented recruitment of P. ochraceus into intertidal ecosystems. In Chapter 3, I used a field experiment to test the hypothesis that this pulse of recruitment was facilitated by SSWS-related adult loss, the consequent decrease in predation by adult P. ochraceus, and increase in prey availability for recruits. Instead of finding evidence that adults dominate recruits in food competition and inhibit recruit success, I found that recruits have a negative effect on P. ochraceus adult densities. Further, treatments where recruits were excluded and only adults had access to prey communities showed the highest control of sessile invertebrate prey populations at the end of the year-long experiment. Thus, these results suggest that adult P. ochraceus will not hinder recruit recovery, but propose a mechanism whereby high recruit densities may increase vulnerability to SSWS-induced shifts in community structure. Outbreaks of mass mortality, particularly those as widespread as SSWS, are one of many ecological challenges driven by anthropogenic environmental changes such as warming and ocean acidification. However, predicting the vulnerability of species and populations to global change is an ongoing and significant challenge for researchers and managers. In Chapter 4 I assessed whether intraspecific physiological variability could help predict P. ochraceus recruit response to ocean acidification and warming. I found that individual metabolic rate interacted with ocean acidification and food availability to drive sea star growth, and that an interaction between metabolic rate and temperature also predicted sea star predation on Mytilus spp. mussels. Thus, these results have implications not only for P. ochraceus but also for its food web interactions. Incorporating these results into predictive frameworks may improve our ability to anticipate and scale up responses to global change across levels of ecological organization. In summary, my dissertation, although chock-full of surprises, presents several paths forward for improving predictive ability in the face of accelerating anthropogenic global changes. Further, we reinforce the notion that management strategies should be cautious and anticipate ecological surprises. Predicting the future is challenging even when predictions are well-informed, particularly in environmental contexts that have never been encountered before. © Copyright by Jenna Sullivan Stack May 3, 2019 All Rights Reserved Marine Mass Mortality in a Global Change Context: Impacts on Individuals, Populations and Communities by Jenna Sullivan Stack A DISSERTATION submitted to Oregon State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy Presented May 3, 2019 Commencement June 2019 Doctor of Philosophy dissertation of Jenna Sullivan Stack presented on May 3, 2019 APPROVED: Major Professor, representing Zoology Chair of the Department of Integrative Biology Dean of the Graduate School I understand that my dissertation will become part of the permanent collection of Oregon State University libraries. My signature below authorizes release of my dissertation to any reader upon request. Jenna Sullivan Stack, Author ACKNOWLEDGMENTS I first met my advisor Bruce Menge one early morning on the rocks at Boiler Bay. Although I’m sure he was keen to get to all the work left to do that day before the tide came in and covered our field site, he took a momentary pause to point out interesting features of the site and changes he had observed over the previous decades. A place where part of the intertidal had caved in years ago, resulting in loss of some of the ecological high zone. The old abandoned boiler from which the site got its name. A section of bench where one of his previous students had jackhammered out replicate tidepools for an experiment on how top-down vs. bottom- up forces structure communities. I was, and remain, in awe of the immensity of both Bruce’s knowledge of and also his curiosity and dedication to learning about intertidal ecosystems. It is rare that an advisor continue to do fieldwork alongside their technicians and graduate students, let alone at every opportunity, for decades. Bruce generously shared his time and expertise with me through all the ups and downs of this dissertation, from brainstorming mechanisms that might explain a puzzling observation to helping me remove “innumerable” baby sea stars from cracks in the intertidal rocks with tweezers. Bruce is kind, unfailingly supportive, and a world-class scientist. Quite obviously, I couldn’t have done this without him. I am also deeply indebted to Jane Lubchenco for her generous mentorship. Although Jane does not officially sit on my dissertation committee, she has transformed my thinking about science and how it can best serve society. Working with Jane has been an honor, and the experience has already shaped my career goals and trajectory. As a team, Bruce and Jane have taught me that high standards and rigorous science do not preclude, but are enhanced by, personal connections and empathy. Thanks for allowing me to be a small shoot on the LubMenge family tree. I am grateful to my committee, Francis Chan, Mark Novak, Debashis Mondal, Judy Li, and, most recently, Ana Spalding, for their support and guidance. Judy, thanks for the chats over coffee and for pushing me to think beyond my specific study system. Conversations and classes with Mark expanded my concept of an ecologist and inspired me to think more deeply about the theories that inform my research. I am especially grateful to Francis Chan for his mentorship, the impacts of which extend far beyond this dissertation. In every conversation, Francis challenged me to think big, question assumptions, and work to advance our thinking as ecologists. Through posing thoughtful questions and providing guidance, Francis pushed me to critically evaluate how my science can make a difference. I am grateful for Kirsten Grorud- Colvert’s mentorship, ranging from how to respond to reviewer #2 to how to deal with changing my last name. I deeply admire Kirsten as a scientist and as a person. I am privileged to be part of the LubMenge lab group, and to have overlapped with such incredible people. Field and lab work would not have been possible without the help of numerous labmates and volunteer assistants – too many to name without this dissertation becoming as long as Bruce’s – who made this PhD not only feasible but also fun. In particular: Liz Cerny-Chipman, thanks being such a generous friend and role model. Your integrity is inspiring and so are your whelk-spotting skills. Thanks to Chenchen Shen and Allie Barner for all the adventures and for inspiring me to be a more thoughtful scientist – I am so fortunate to have had your guidance throughout this PhD. Sarah Gravem, thanks for sharing your vast knowledge and fun, positive energy. Thanks to Barbara Spiecker, one of the most inspiring people I’ve ever met, for her research help, friendship and endless patience with me.
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