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Modeling the Processes Affecting Larval Haddock (Melanogrammus Aeglefinus) Survival on Georges Bank

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Modeling the Processes Affecting Larval Haddock (Melanogrammus Aeglefinus) Survival on Georges Bank Modeling the processes affecting larval haddock (Melanogrammus aeglefinus) survival on Georges Bank IM4ASSACHUSETTS INSTITUTE Colleen Mary Petrik OF TECHNOLOGY B.S., Marine Science and Biology University of Miami, 2005 LIBRARIES Submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Biological Oceanography at the ARCHIVES MASSACHUSETTS INSTITUTE OF TECHNOLOGY and the WOODS HOLE OCEANOGRAPHIC INSTITUTION February 2011 0 2011 Colleen Mary Petrik. All rights reserved. The author hereby grants to MIT and WHOI permission to reproduce and to 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 17 December 2010 Certified by Cabell S. Davis upervisor Accepted by Simon Thorrold Chair, Joint Committee for Biological Oceanography Massachusetts Institute of Technology and Woods Hole Oceanographic Institution 2 Modeling the processes affecting larval haddock (Melanogrammus aeglefinus) survival on Georges Bank by Colleen Mary Petrik Submitted to the Department of Biology on 17 December 2010, in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy in Biological Oceanography Abstract The ultimate goal of early life studies of fish over the past century has been to better understand recruitment variability. Recruitment is the single most important natural event controlling year-class strength and biomass in fish populations. As evident in Georges Bank haddock, Melanogrammusaeglefinus, there is a strong relationship between recruitment success and processes occurring during the planktonic larval stage. Spatially explicit coupled biological-physical individual-based models are ideal for studying the processes of feeding, growth, and predation during the larval stage. This thesis sought new insights into the mechanisms controlling the recruitment process in fish populations by using recent advances in biological-physical modeling methods together with laboratory and field data sets. Interactions between feeding, metabolism and growth, vertical behavior, advection, predation, and the oceanic environment of larval haddock were quantitatively investigated using individual-based models. A mechanistic feeding model illustrated that species-specific behavioral characteristics of copepod prey are critically important in determining food availability to the haddock larvae. Experiments conducted with a one-dimensional vertical behavior model suggested that larval haddock should focus on avoiding visual predation when they are small and vulnerable and food is readily available. Coupled hydrodynamics, concentration-based copepod species, and individual-based larval haddock models demonstrated that the increased egg hatching rates and lower predation rates on larvae in 1998 contributed to its larger year-class. Additionally, results from these coupled models imply that losses to predation may be responsible for interannual variability in recruitment and larval survival. The findings of this thesis can be used to better manage the haddock population on Georges Bank by providing insights into how changes in the physical and biological environment of haddock affect their survival and recruitment, and more generally about the processes significant for larval fish survival. Thesis supervisor: Cabell S. Davis Title: Senior Scientist, WHOI 4 Acknowledgements First and foremost, I thank Cabell Davis for being an insightful, encouraging, fun, and flexible thesis advisor. He presented me with many great opportunities, such as visiting the Liquid Jungle Lab in Panama, participating in two research cruises in the Caribbean, and presenting my work at multiple conferences in Europe. Cabell also allowed me the freedom to choose my own research topics and work remotely at times. His unwavering enthusiasm and genuine curiosity renewed my interest in oceanography whenever I was discouraged with my research and continues to motivate me. I would like to extend appreciation to my other thesis committee members. Rubao Ji and all his lab members were so patient and helpful while I learned Linux, FORTRAN, and FVCOM. I have little experience with the live versions of the organisms I study, and I thank Greg Lough for supplying this knowledge of haddock and cod. I thank Andy Solow for his statistical assistance and Glenn Flier for physical and mathematical considerations, and both of them for helping me see the big picture. Furthermore, I have benefitted from the work of and interactions with many colleagues. Trond Kristiansen supplied me with a larval fish model and numerous potential research questions to answer with it. I have the utmost respect for David Mountain, Larry Buckley, Betsy Broughton, and everyone involved in the collection and analysis of GLOBEC data, if not for all the information it has provided, then simply because I did not have to venture out onto Georges Bank in the winter. I am grateful for my involvement with the ICES Working Group on Physical Biological Interactions. This group involved me in their workshops and meetings, gave me a venue to present my work and a place to submit it for publication, offered advice on research, introduced me to potential collaborators, and inspired me to produce science as admirable as theirs. I was fortunate to start the MIT-WHOI Joint Program with a great cohort of Biology students, Kelton McMahon, Christine Mingione Thompson, Louie Wurch, and Whitney Krey, who have been there for academic concerns and social entertainment. Many other friends at WHOI and MIT kept me sane, including Paul Craddock, Taylor Crockford, Jimmy Elsenbeck, Matt Mazloff, Caleb Mills, Melissa Patrician, and Meredith White. I appreciate my family and David Fortunato for their love and support, even if they did not always understand what I was studying. My mom's advice and sentiments were always uplifting, while David gave me the confidence to reach this goal. Last but not least, I am thankful for the Academic Programs Office, who have helped me in a number of ways. They make the graduate experience bearable, if not pleasant, and I attribute much of the success of the Joint Program to them. Financial support was provided by a WHOI Watson Fellowship, a WHOI Coastal Ocean Institute Student Research Proposal Award, and GLOBEC grants NA 17RJ 1223 (NOAA) and OCE0815838 (NSF). 6 Contents 1. Fisheries recruitment variability: a case study using larval haddock on Georges Bank 1.1. B ackground ......................................... 1 1.1.1. Recruitment variability ........ .......... 1.1.2. H addock ................................... 14 1.1.3. G LO B EC ................................... 16 1.1.4. G eorges B ank ................................... 17 1.1.5. Individual-based modeling ....................... .21 1.2. Thesis Overview .21 1.2.1. Foraging m odel ............................. .22 1.2.2. Vertical behavior model .23 1.2.3. Three-dimensional coupled biological-physical model 25 R eferences ............................................... F igures ....... ...... ......... ... ............... ...... 2. Prey selection by larval haddock and cod on copepods with species-specific behavior: an individual-based model analysis 3. Modeling the vertical behavior of larval haddock on Georges Bank with respect to starvation and predation mortality ( 3.1. Introduction .......... ................................... 61 3.2. M odel ........................ ........................................ 64 3.2.1. Environment ............ ........................................ 64 3.2.2. Foraging submodel ...... ........................................ 69 3.2.3. Bioenergetics submodel ........................................ 70 3.2.4. Vertical behavior models ........................................ 70 3.2.5. Optimal c ............ ........................................ 73 3.2.6. Fitness .................. ........................................ 75 3.2.7. Simulations ............ ........................................ 76 3.3. R esults ........................ ........................................ 76 3.3.1. Case A - Diffusion ...... ........................................ 76 3.3.2. Case B - Prey concentration and light with random direction ............. ................... .77 3.3.3. Case C - Prey concentration and light with directed sw im m ing ................................. .79 3.3.4. A ll cases ................................. .81 3.4. D iscussion ....................................... .82 R eferences ............................................. .89 T ab les .... ...... ...... ... ............ ......... ... ...... ... .. 9 7 F igures ................................................ 10 1 4. Interannual differences in larval haddock survival: hypothesis testing with a coupled biophysical model of Georges Bank 115 4.1. Introduction ........................ .. .. .. .. .. 115 4.2. Model.......................... 118 4.2.1. Physical environment ............ 118 4.2.2. Prey ........................ 118 4.2.3. Foraging submodel .............. 119 4.2.4. Bioenergetics submodel ...... 120 4.2.5. Predation submodel ............ 120 4.2.6. M ortality .................. 120 4.2.7. Simulations .................. 122 4.3 . R esults.............................. 128 4.3.1. Reference case ... 128 4.3.2. Hatch distribution effect on survival and sources
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