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Abigail Jean Fusaro

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Abigail Jean Fusaro SPATIAL AND TEMPORAL POPULATION GENETICS AT DEEP-SEA HYDROTHERMAL VENTS ALONG THE EAST PACIFIC RISE AND GALÁPAGOS RIFT by Abigail Jean Fusaro B.S., University of Rhode Island, 2002 Submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy at the MASSACHUSETTS INSTITUTE OF TECHNOLOGY and the WOODS HOLE OCEANOGRAPHIC INSTITUTION September 2008 © 2008 Abigail J. Fusaro All rights reserved. The author hereby grants to MIT and WHOI permission to reproduce and distribute publicly paper and electronic copies of this thesis document in whole or in part in any medium now known or hereafter created. Signature of Author Joint Program in Oceanography/Applied Ocean Science and Engineering Massachusetts Institute of Technology and Woods Hole Oceanographic Institution July 7, 2008 Certified by Timothy M. Shank Thesis Supervisor Accepted by Edward F. DeLong Chair, Joint Committee for Biological Oceanography Massachusetts Institute of Technology and Woods Hole Oceanographic Institution 2 Spatial and Temporal Population Genetics at Deep-Sea Hydrothermal Vents Along the East Pacific Rise and Galápagos Rift by Abigail Jean Fusaro Submitted to the Department of Biology on July 7, 2008, in partial fulfillment of the requirements for the degree of Doctor of Philosophy Abstract Ecological processes at deep-sea hydrothermal vents on fast-spreading mid-ocean ridges are punctuated by frequent physical disturbance. Larval dispersal among disjunct vent sites facilitates the persistence of sessile invertebrate species in these geologically and chemically dynamic habitats despite local extinction events. Regional population extension and rapid recolonization by the siboglinid tubeworm Riftia pachyptila have been well documented along the East Pacific Rise and the Galápagos Rift. To analyze spatial and temporal population genetic patterns and the processes governing them at ephemeral and disjunct habitats, a suite of 12 highly variable microsatellite DNA markers were developed for this species. Eight of these loci were used to assess the regional and within-ridge genetic structure of recent colonists and resident adults collected from nine sites in the eastern Pacific Ocean over period of three to seven years. A significant seafloor eruption during the seven-year sampling period allowed investigation into the role of local extinction in population genetic diversity at the Tica vent site at 9˚N EPR, while collections within two and five years of an eruption that created the Rosebud vent field at 86˚W GAR provided insights into genetic diversity input over population establishment. For the first time, this thesis demonstrated significant genetic differences between Riftia populations on the East Pacific Rise and Galápagos Rift. Moreover, the separate treatment of colonist and resident subpopulations revealed a high potential for local larval retention at vent sites. This mechanism for recruitment likely sustains disjunct populations and supports the recolonization of locally extinct areas after disturbance events, while episodic long-distance dispersal maintains genetic coherence of the species. Temporal population genetic consideration at the Tica site on the East Pacific Rise suggests that the 2005-2006 seafloor eruption had little to no discernable effect on local population genetic composition. Yet local populations appear to exhibit a small degree of 3 genetic patchiness, with a high degree of relatedness (half-sibs) among subsets of individuals within both colonist and resident cohorts. This thesis broadens the application of recently developed molecular techniques to study the effect of ridge-crest processes and offers new perspectives into marine dispersal, gene flow, and population differentiation. Thesis Supervisor: Timothy M. Shank Title: Associate Scientist, Biology Department Woods Hole Oceanographic Institution 4 Acknowledgments “No man is an Island, entire of itself.” ~ John Donne, Meditation XVII The journey I have been on these past several years would not have been possible had it not been for an enormous support system. I began the MIT/WHOI Joint Program in the Halanych lab, where Ken, Rob, Yale, Nan, and Annette led me into the world of molecular biology and genetics. Within months, Tim Shank graciously adopted me into his lab, and the adventures began in earnest. Tim has allowed me great freedom to explore, travel in several different (unfunded) research directions, and reconsider the routes that would lead to intellectual satisfaction. Together with my committee—Gabi Gerlach, Ron Etter, Adam Soule, Glenn Flierl—he has encouraged independence and ownership of my work. I take away from them the lessons of speculation and risk. Many lab mates and classmates have coaxed me through the growing pains of grad school. Walter and Kate have provided a solid foundation as fellow students, always willing to lend an ear or a hand. I am also indebted to Amy and Diane for their previous work developing microsatellites for Riftia and Bathymodiolus and for being incredibly generous with their time and expertise. Rhian, Stace, and Breea have rounded out adventures at sea and in the lab with their experience and wherewithal. In addition, I have enjoyed working with and benefited from the manual support of many lab guests, including Taylor, Sara, Sarah, Carl, Alex, Aaron, Cory, and John. My biology cohort of Kristen, Jonathan, Carly, and Ann has always been there as we leapfrogged through classes, generals preparations, thesis proposals, and defenses. Our thesis support dinners in the past year have nourished me not only with meals but also with food for thought. I also don’t know how I would have survived (and in one piece) had it not been for Trish and Sharon. Leave it to two geologists to make me laugh the most and tolerate me at my worst. Together with Cory and Hunter, they are friends for life. I gratefully acknowledge the opportunities for sample collection provided by S.C. Cary (NSF OCE-9714302), G. Luther (NSF OCE-9714302), R.A. Lutz (NSF OCE-9529819), and K.L. Von Damm (NSF OCE-9419156) during their expeditions, as well as the invaluable efforts of the captain, crew, and submersible pilots of the R/V Atlantis/Alvin. I would also like to thank A.M. “Ginger” Clark at the University of Florida, Gainesville for welcoming me into her lab and technical support for the development of microsatellite- enriched libraries. Special thanks to T. Paton, L. Timms, S. Russell, and J. Zhanqin at The Centre for Applied Genomics, Toronto, Canada for genotyping prepared samples and ensuring the highest quality control. I was supported during my graduate tenure by a 2002 award from SEAPACE, a NSF 5 Biocomplexity award to K. Halaynch (EAR-0120646), a 2004-2008 NOAA Dr. Nancy Foster Scholarship (Award No. NA04NOS4290253), and the Woods Hole Oceanographic Institution Academic Programs Office. Funding for my research was provided by the National Science Foundation (OCE-0327261 and OCE-0324232) and the National Oceanic and Atmospheric Administration’s Office of Ocean Exploration and Research (Award Nos. NA03OAR4600110, NA16RP2390, and NA04OAR4600049) support to TMS. Finally, I would like to thank my family for setting solid examples of integrity and hard work, for love and encouragement to do my best, and for believing in me through it all. Thank you for all the trips together to the seashore and for the inspiration to pursue my dreams, even when I was not entirely sure what they were. To my husband Vin, I cannot find words enough to express my gratitude. Thank you for the months endured at home while I was at sea, for keeping me going through late nights and early mornings, through tears and laughter, with constant companionship and faith. You have made sure that I get outdoors and take much-needed breaks, kept food in the house and dinner on the table, and given me perspective. This thesis is as much of your love and labor as it is of mine. 6 Table of Contents ABSTRACT ..........................................................................................................................................................3 ACKNOWLEDGMENTS...................................................................................................................................5 TABLE OF CONTENTS ....................................................................................................................................7 LIST OF FIGURES ...........................................................................................................................................11 LIST OF TABLES .............................................................................................................................................13 CHAPTER 1: INTRODUCTION....................................................................................................................15 1.1 OVERVIEW .............................................................................................................................................15 1.2 UTILITY OF POPULATION GENETIC STUDIES..........................................................................................16 1.3 THE METAPOPULATION APPROACH TO TEMPORAL GENETICS ..............................................................17 1.4 PHYSICAL AND BIOLOGICAL SETTING ...................................................................................................18 1.5 HISTORY OF LOCAL EXTINCTION, COLONIZATION, AND SUCCESSION ON THE EAST PACIFIC RISE AND GALÁPAGOS RIFT ..........................................................................................................................20
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