Deep-Sea Coral Biogeography and Community Structure in Tropical Seamount Environments
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DEEP-SEA CORAL BIOGEOGRAPHY AND COMMUNITY STRUCTURE IN TROPICAL SEAMOUNT ENVIRONMENTS A Dissertation Submitted to the Temple University Graduate Board In Partial Fulfillment of the Requirements for the Degree DOCTOR OF PHILOSOPHY by Steven R. Auscavitch December 2020 Examining Committee Members: Dr. Erik E. Cordes, Advisory Chair, Biology Dr. Robert W. Sanders, Biology Dr. Matthew R. Helmus, Biology Dr. Amanda W. J. Demopoulos, External Member, U.S. Geological Survey © Copyright 2020 By Steven R. Auscavitch All rights reserved ii ABSTRACT As the largest and most poorly environment on Earth, the deep-sea is facing global threats from climate change and anthropogenic disturbance further compounded by the lack of critical baseline data on seafloor species composition and community structure. Many data-deficient regions include those in geographically-isolated offshore environments, like low-latitude seamounts, where sampling and surveys have been limited, resulting in critical knowledge gaps that do not allow for effective conservation measures to be realized. This work seeks to characterize the coral fauna of tropical seamount environments greater than 150 m depth and understand the environmental controls on species distribution and community assembly for long-lived, ecologically- important species, primarily from the Octocorallia, Antipatharia, Stylasteridae, and Scleractinia. Methodologies for accomplishing this research have included analysis of remotely operated vehicle (ROV) video surveys and identification of collected voucher specimens to understand biogeographic patterns within coral communities on seamounts and other rugged seafloor features in 3 different regions: the tropical western Atlantic (Anegada Passage), the equatorial central Pacific (Phoenix Islands), and the tropical eastern Pacific (Costa Rica). These regions represent vastly different oceanographic regimes in terms of biological productivity and water column structure resulting in differential effects on deep-sea coral communities. Evidence from these three regions has shown significant effects of the role that oceanic water masses have on structuring deep- water coral biodiversity and suggests that these features, along with other abiotic environmental variables, are important indicators for understanding species distribution iii patterns, community structure, and global biogeographic patterns. More broadly, the results of this work have demonstrated the capabilities of exploratory ROV surveys, across multiple platforms, to add practical knowledge to coral species inventories and identify bathyal biogeographic patterns in remote regions of the deep sea. The results of this work, serving as baseline coral biodiversity surveys for each area, are also germane to evaluating the effects of human-mediated disturbance and global climate change in the deep ocean. These disturbances also include ocean acidification, ocean deoxygenation, deep-sea mining, and bottom-contact fishing, all of which have been identified as threats to the seamount benthos. iv For Eneko and Amaia v ACKNOWLEDGMENTS I must first acknowledge the efforts and guidance of my dissertation committee: Erik Cordes, Bob Sanders, Matt Helmus, and Amanda Demopoulos. All were instrumental in providing comments and feedback over the years that have greatly improved the quality of this work. I am grateful to have had Amanda as a mentor since the beginning and as my science co-lead on the Okeanos Explorer in 2017. Perhaps most importantly, I know I would not be on this path right now if it was not for Erik’s mentorship and trust in me to take on these projects in the aftermath of our first cruise in the Anegada Passage on the E/V Nautilus in September 2014. I look forward to many more opportunities for exploration and collaboration in the years to come. Parts of this work were funded from grants through the NOAA Office of Ocean Exploration and Research and the National Science Foundation. I also thank Temple University for funding provided through various teaching and research assistantships as well as a Dissertation Completion Grant award during the Summer 2020 term. Also, thanks to the Temple University Biology Department for providing travel funding to present parts of this work at international conferences. As ocean research is often a team effort, I must acknowledge the contributions of key groups of people and institutions that have been instrumental in my professional and academic growth during my graduate career. Special thanks is due to Nicole Raineault, Allison Fundis, Megan Cook, and the E/V Nautilus science parties and crew during NA034, NA052, and NA110. The opportunities I have had, and people I have worked with at the Ocean Exploration Trust, have been critical in making me the explorer, scientist, and communicator I am today. vi I am also grateful to all the personnel who I have worked with at the NOAA Office of Ocean Exploration and Research and in the NOAA Okeanos Explorer Program, particularly during EX1703: Exploring Remote Pacific Marine Protected Area. The chance to co-lead two cruise legs on the NOAA Ship Okeanos Explorer have been career-shaping opportunities that I will never forget. For their support on work in the Phoenix Islands Protected Area and central Pacific, I thank Randi Rotjan, Tim Shank, Brian Kennedy, and Chris Kelley. In addition, I am greatly appreciative of the efforts of the amazing folks at the Schmidt Ocean Institute and the R/V Falkor team as well as the science party and crew during Falkor cruises FK171005: Discovering Deep-Sea Corals of the Phoenix Islands and FK190106: Costa Rican Deep Sea Connections. I would like to extend a warm thanks to the governments of Anguilla, the British Virgin Islands, the Republic of Kiribati, and Costa Rica for permitting us to conduct field work in their waters. I would also like to thank all local observers who joined us at sea. My deep thanks to Jorge Cortés for facilitating deep-water exploration off Costa Rica. All voucher specimens, collected under permit, will ultimately be deposited at the Smithsonian Institution National Museum of Natural History for use by the broader scientific community. This work was greatly enhanced by a number of taxonomists and specialists who lended their time to provide identifications of deep-sea corals from specimen collections and photos. I will be forever grateful to Steve Cairns, Dennis Opresko, Daniel Wagner, Andrea Quattrini, Tina Molodtsova, Asako Matsumoto, Scott France, Les Watling, Salome Buglass, and Odalisca Breedy. I would also like to acknowledge the support of vii Tom Hourigan and the team at NOAA’s Deep Sea Coral Research and Technology Program for helping me archive the valuable deep-sea coral data presented in this work. I have had the fortune to work alongside many great colleagues in the Cordes Lab through the years but I am particularly grateful to fellow (and former) graduate students Alexis Weinnig, Carlos Gómez, Sam Georgian, Danielle DeLeo, Alanna Durkin, April Stabbins, Emily Cowell, Melissa Betters, and Ryan Gasbarro. I am also thankful to have had the support of Abigail Keller, Jay Lunden, and Amanda Glazier in various aspects of this research including field work, data analysis, writing, and editing. Finally, I am greatly appreciative of the innumerable contributions of the many undergraduates that have passed through the Cordes Lab, but this specific work greatly benefitted from the collaborative efforts and contributions of Mary Deere and Lyanna Kessler. None of the following work would have been possible or half as enjoyable without the continuous and unyielding support of my family, Amaia and Eneko. Beyond any doubt, you both are the most significant driving forces behind my will to complete this work. Your understanding during the many days and weeks at sea has made me appreciate how lucky I am to have you both in my life. Amaia your strength, support, and sacrifice for our family is without bound and I am thankful to have you as my partner. Eneko, one day I hope you have the opportunity to explore this world and share your discoveries as I have. Find and follow your passions and let them take you wherever they lead, no matter how difficult the challenges. Embrace the obstacles you might face along the way, always keep those you love close, and persevere. viii TABLE OF CONTENTS Page ABSTRACT ....................................................................................................................... iii DEDICATION .....................................................................................................................v ACKNOWLEDGMENTS ................................................................................................. vi LIST OF TABLES ........................................................................................................... xiii LIST OF FIGURES ......................................................................................................... xiv CHAPTER 1. INTRODUCTION .........................................................................................................1 1.1 Biology and ecology of deep-sea corals ............................................................3 1.2 Seamounts and similar rugged seafloor features ...............................................9 1.3 Objectives and aims .........................................................................................13 2. DISTRIBUTION OF DEEP-WATER SCLERACTINIAN AND STYLASTERID CORALS ACROSS ABIOTIC ENVIRONMENTAL GRADIENTS