Phylodemography of the Blue Rockfish (Sebastes Mystinus) from California to Washington
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PHYLODEMOGRAPHY OF THE BLUE ROCKFISH (SEBASTES MYSTINUS) FROM CALIFORNIA TO WASHINGTON A thesis submitted to the faculty of Moss Landing Marine Laboratories and San Francisco State University In partial fulfillment of The requirements for The degree Master of Science In Marine Science by Jason Marc Cope December, 2002 Copyright by Jason Marc Cope 2002 PHYLODEMOGRAPHY OF THE BLUE ROCKFISH (SEBASTES MYSTINUS) FROM CALIFORNIA TO WASHINGTON Jason Marc Cope Moss Landing Marine Laboratories San Francisco State University 2002 Marine resource managers are often challenged with balancing use and sustainability of resources with limited data in dynamic marine environments. Though vagile life history stages are common in marine organisms, increasing evidence indicates intraspecific population substructuring exists in the marine realm. It is also recognized that life history characteristics driving demographic behavior of populations vary spatio- temporally. Spatial and temporal relationships of intraspecific populations must therefore be considered for proper management of these resources, though biological resolution on such levels is often limited. This thesis presents a method by which the combination of limited genetic and demographic information on evolutionary and ecological time scales can produce an historical and contemporary understanding of population dynamics useful for developing management decisions on resources in urgent need of swift management action. I call this approach “phylodemography” and illustrate the method by investigating the spatially distinct subpopulation dynamics of the blue rockfish (Sebastes mystinus) from California to Washington. Genetic samples (N = 245) of blue rockfish from eight sites throughout its range were taken to explore population substructuring. Sequencing of a 498 base pair portion of the mitochondrial DNA control region revealed two major subpopulations of blue rockfish, with the apparent genetic break occurring at Cape Mendocino, CA. This is a relatively unexplored biogeographic break that may reveal a strong barrier for gene flow in other marine species. The northern group is a derivative of the southern group, with the last contact between the two regions occurring 2,000 to 6,000 years ago. With this information, structured subpopulations were then modeled using deterministic and stochastic five stage matrix population models and region specific life history information and vital rates. Demographic and environmental affects (due to intensity-varying El Niño events) were incorporated. Furthermore, genetic pairwise mismatch distributions were used to explore historical population dynamics of each region. The northern population was historically more stable and currently increasing compared to the unstable and most likely decreasing southern population. Age truncation, environmental variability, and fishing pressure are likely driving the southern subpopulation down. Using an introduced time-standardized elasticity method, a 13.5 to 14 inch size limit is recommended for the southern blue rockfish subpopulation to protect the “life history management unit” of the subadult stage and 8 reproductive year classes. The phylodemographic approach shows promise for developing sustainable management schemes for long-lived, slow growing, and late to mature marine organisms where other methods may be inappropriate. The method is also flexible enough to include new types of information on spatial structure and demographic parameters as data accumulate. I certify that the Abstract is a correct representation of the content of this thesis. _____________________________________________ __________________ Gregor M. Cailliet, Chair, Thesis Committee Date ACKNOWLEDGEMENTS This research was made possible by funding provided by the Pacific States Marine Fisheries Commission (via the California Department of Fish and Game), the PADI Foundation, the Earl & Ethel Myers Foundation Oceanographic and Marine Biology Trust, and Project Aware. Additional support was received from the Chela Financial Scholarship, CSU Sally Casanova Pre-Doctoral Program, the Ray Cannon Memorial Scholarship, and the John H. Martin Scholarship. Lisa Kerr, Matt Levey, Chris Black, Jason Felton, Marilyn Cope (baby blue slayer), Ramiro Ferrey, Jeff Field (the Sebasterminator), Eddie Grannis, Joanna Grebel, Julie Haws, and Wade Smith all helped with sample collection. I want to especially thank Lisa for sacrificing her time to come along on every collecting trip. Additional samples were prepared and provided by Martha Burford, Vince Buonocorssi, Carol Kimbrell, John Smiley, and Russ Vetter (CA); Don Bodenmiller and ODFW samplers (OR); Farron Wallace, Wayne Palsson, and WDFW samplers (WA). I express major gratitude to Laurie Fitzgerald, Heather Lohr, and Susan von Thun for leading me through the dark and unforgiving mistress that is genetics. The hospitality and generosity of Dr. Jon Geller and his lab was indispensable for the completion of the genetics portion of this thesis. Emily Dreschler valiantly assisted in some of the more mundane aspects of genetic lab work. Dr. Giacomo Bernardi, Yong-Jin Won, and Dr. Robert Vrijenhoek provided helpful discussion regarding genetic analyses. Cathy Evans (WDFW) provided blue rockfish age at length information for Washington vi used in Chapter 3. The multi-talented Matt Levey helped me with the site map used in Chapters 2 and 3 and video-documentation of a collection trip. Joan Parker, Joanna Grebel, Bridget Watt, and 4 years of library assistants provided me with several groves of very important papers. Gail Johnston and Donna Kline helped right myself to the administrative requirements of this degree. Drs. Jon Geller and Ralph Larson offered excellent insight and comments on thesis drafts. To Dr. Gregor M. Cailliet, my advisor and friend, thank you∞ for everything you taught and gave me during my wonderful time at Moss Landing Marine Laboratories. And with much appreciation I acknowledge the friendship and encouragement given me by the wonderful faculty, students, staff, and affiliates of Moss Landing Marine Laboratories. vii TABLE OF CONTENTS List of Tables……………………………………………………………………………..xi List of Figures…………………………………………………………………………...xiii CHAPTER 1: Phylodemography as it pertains to the integration and application of demography and genetics into studies of intraspecific population dynamics or, the relevance of a phylodemographic approach to population biology……………………….1 Problems with metapopulation theory and PVA………………………………....4 So what about Phylodemography?……………………………………………….6 Does science really need another term?…………………………………………...8 LITERATURE CITED……………………………………………………………………8 CHAPTER 2: Population genetics and phylogeography of the blue rockfish (Sebastes mystinus) from Washington to California…………………………….………………….15 ABSTRACT……………………………………………………………………………..16 INTRODUCTION……………………………………………………………………….17 METHOD………………………………………………………………………………..22 Molecular marker………………………………………………………………...21 DNA collection, extraction, amplification, and sequencing……………………..23 Genetic Diversity..……………………………………………………….23 Sample size...…………………………………………………………….24 Population structure……………………………………………………...24 Phylogeography……………………………………………………….…26 RESULTS………………………………………………………………………………..26 Sample Size………………………………………………………………………26 Genetic Diversity………………………………………………………………...27 Population structure and gene flow………………………………………………28 viii Phylogeography………………………………………………………………….29 DISCUSSION……………………………………………………………………………31 Population Structure……………………………………………………………..31 Gene flow in the nearshore system………………………………………………33 Barriers to gene flow……………………………………………………………..35 Future Considerations……………………………………………………………38 LITERATURE CITED…………………………………………………………………..40 CHAPTER 3: Stochastic matrix modeling of genetically differentiated populations of the blue rockfish (Sebastes mystinus): An introduction to phylodemography………..67 ABSTRACT……………………………………………………………………………...68 INTRODUCTION……………………………………………………………………….69 METHODS………………………………………………………………………………73 Matrix Population Model………………………………………………………...73 Deterministic Model Parameter Estimation……………………………………...77 Stochastic Model Parameter Estimation…………………………………………80 Elasticities………………………………………………………………………..83 Loop analyses…………………………………………………………………….84 Historical Demography…………………………………………………………..85 RESULTS………………………………………………………………………………..86 Deterministic models…. ………………………………………………………...86 Stochastic models……………………...…………………………………………86 Elasticities………………………………………………………………………..87 Loop analyses…………………………………………………………………….90 Historical Demography…………………………………………………………..91 DISCUSSION…………………………………………………………………………....91 Ecological population dynamics of blue rockfish subpopulations………………91 Evolutionary population dynamics of blue rockfish subpopulations…………..101 ix Phylodemography………………………………………………………………102 LITERATURE CITED…………………………………………………………………105 x LIST OF TABLES TABLE PAGE CHAPTER 1 1. Examples from primary scientific literature demonstrating different approaches to combining demographic and genetic in populations studies………..……………………………………………………………………....14 CHAPTER 2 1. Suspected population structure of some rockfishes off the Pacific coast of the United States……………….………………………………...…………………………...….55 2. Per area genetic diversity and neutrality test statistics for a 498 bp region of the blue rockfish mtDNA control region………………………………………………...56 3. Sequence divergence (below diagonal; standard deviation above) among blue rockfish sample areas………………………………………………………………..57 4. Relative