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University of California, San Diego UNIVERSITY OF CALIFORNIA, SAN DIEGO Separating environmental effects from fishing impacts on the dynamics of fish populations of the Southern California region A dissertation submitted in partial satisfaction of the requirements for the degree Doctor of Philosophy in Oceanography by Chih-hao Hsieh Committee in charge: Professor George Sugihara, Co-Chair Professor Mark D. Ohman, Co-Chair Professor David M. Checkley, Jr. Professor Phil A. Hastings Professor John R. Hunter Professor Arthur J. Miller Professor Kaustuv Roy 2006 Copyright Chih-hao Hsieh, 2006 All rights reserved The dissertation of Chih-hao Hsieh is approved, and it is acceptable in quality and form for publication on microfilm: ________________________________________________ ________________________________________________ ________________________________________________ ________________________________________________ ________________________________________________ ________________________________________________ Co-Chair ________________________________________________ Co-Chair University of California, San Diego 2006 iii To Chang-tai Shih, who brought me on board science, and pointed me in the direction of life. iv TABLE OF CONTENTS Signature Page ……………………………………………………………………...……iii Dedication ……………………………………………………………………….….……iv Table of Contents ……………………………………………………….……….….…….v List of Figures ………………………………………………………………….….……vii List of Tables ………………………………………………………………….….……...ix Acknowledgements …………………………………………………………….….……..xi Vita, Publication, and Fields of Study...……………………………………….….…….xiv Abstract of the Dissertation …………………………………………………….….…...xvi Chapter 1. Introduction to the Dissertation ...…….…....……..……………….….….……1 Introduction …………………………………………….……………………...….…...2 Thesis outline ………………………………………….……………………...….……7 Tables ………………………………………………….……………………...….…..11 Figures ……………………………………………….……………………...…...…...14 References ………………………………………….……………………...…...….…17 Chapter 2. A comparison of long-term trends and variability in populations of larvae of exploited and unexploited fishes in the Southern California region: a community approach ..……………………………………………………………………………..…22 Abstract ………………………………………….………………………...…...….…23 Introduction …………………………………….………………………...…...….…..24 Methods ………………………………………….………………………...…...….…25 Results ………………………………………….………………………...….....….…30 Discussion …………………………………….………………………...….....….…..39 Acknowledgements …………………………….………………………...….....….…46 References…………………………………….………………………...….....….…...46 Appendix …………………………………….………………………...….....….…....49 Supplementary references ……………………………………………………………51 Chapter 3. Fishing elevates variability in the abundance of exploited species: long-term evidence that fishing magnifies uncertainty in a changing environment …………..…...63 Abstract ………………………………………….………………………...…...….…64 Main Text …………………………………….……………...…………...…...….….65 v Methods ……………...……………….……….………………………...…...…….…71 Tables ………………………………………….………………………...…...….…...74 Figures ……………………………………….………………………...…...….….….76 References ……………………………………….……….……………...…...………79 Appendix ……………………………………….………………………...…...……...82 Supplementary references …………………………………………………………..101 Chapter 4. Climatic and fishing effects on shifts in spawning distribution of marine fishes .…………………………………………………………………………………..108 Abstract ………………………………………….………………………...…...…...109 Introduction ………………………………………….………………………...……110 Methods ………………………………………….………………………...…...…...112 Results ………………………………………….………………………...…...….…117 Discussion ………………………………………….………………………....….…118 Tables ………………………………………….………………………...…...….….123 Figures ………………………………………….………………………...…...….…126 References ……………………...………………….………………..…...…...….….132 Appendix ……………………...………………….………………..…...…...….…...136 Chapter 5. Distinguishing random environmental fluctuations from ecological catastrophes for the North Pacific Ocean .………….………………..…...…...….….…145 Abstract ………...………………….………………..…...…...………………...…...146 Main Text ……...………………….………………..…...…...………………...…....146 Methods ……...………………….………………..…...…...………………...….…..149 References ……...………………….………………..…...….….……………...…....149 Acknowledgements ………………….………………......….….……………...…....150 Chapter 6. Synthesis and perspectives: Coping with uncertainty and nonlinearity in fisheries management ………….……….………………..….…...………………...…...151 Figures ………...………………….…….…………..…...…...………………...…....160 References ……...………………….…….…………..…...…...……..………...…....162 Appendix ……...………………….…….…………..…...…...……..………...…..…166 Supplementary references …………………………………………………………..170 vi LIST OF FIGURES Chapter 1 1.1. The current state of world marine fishery resources as estimated by FAO for the year 2004 ……………………………………………………….………..………...14 1.2. Environmental variability associated with the southern California bight marine ecosystem ……………………………………………………….……….….….….15 1.3. An example showing that larval abundance is a good indicator of adult spawning biomass ……………………………………………………….……….….….……16 Chapter 2 2.1. Maps showing CalCOFI sampling stations and frequency ….…….….…....…….27 2.2. Abundance time series of fish species ………………………...………..….…….31 3.2. Comparisons of exploited and unexploited species living in the same habitat .….42 Chapter 3 3.1. Relationships between CVs and life history traits for exploited and unexploited species separately ………………………...……….….……………………....……76 3.2. CVs of exploited and unexploited species associated with geographic distributions, habitats, and spawning modes …………...……….….……………………....…….77 3.3. Long-term declining trends in the average age and length of exploited species ....78 Chapter 4 4.1. Map showing the spatial pattern of CalCOFI stations ……………….…....…….126 4.2. Schematic illustrating spatial analyses of change in distribution domain of fishes ….……………...……….….………………………………………....……127 4.3. An example showing the relationship between mean latitudes and mean SSTs and distribution-centers …………………………....……….….……………………..128 vii 4.4. Magnitudes and direction of changes in distribution from the cold and to the warm period for fish species …………………...……….….………………….………..129 4.5. Comparison of life history traits of shifting and non-shifting species …….….....130 4.6. A model illustrating the “fisheries acting as a noise filter” hypothesis …...….....131 Chapter 5 5.1. Examples of the simplex projection method …….….………………..…..……...147 5.2. Examples of the S-map method for the four time series from Figure 5.1 ……....147 Chapter 6 6.1. Spawner-recruitment data for fishes ……...……….….………………..…..…....160 6.2. Frequency of maternal effects on fish early life history traits ……………..……161 viii LIST OF TABLES Chapter 1 1.1. Indicators to detect fishing effects on a population, assemblage, or ecosystem ….11 Chapter 2 2.1. Summary of criteria used in the “expert-knowledge classification system” to determine fish assemblages ………………………………………..……….………26 2.2. Larval vertical distributions compiled from the literature ………..………..…..…29 2.3. Number of taxa and percentage of the fished, bycatch, and unfished in the three primary assemblages before and after taxonomic aggregation …..………..……….30 2.4. Correlation matrices for oceanic, coastal-oceanic, and coastal assemblages …….35 2.5. Average larval abundances in the cold (1951-1976) and warm (1976-1998) periods and selected environmental variables in the regression model for each taxon ….…40 2.6. A comparison between the larval recurrent groups and the assemblages described by the “expert-knowledge classification system” ……………………….….…...…43 2.7. The long-term average spatial coefficient of variation of larval fish abundances for the species used in the pair-wise comparisons ……………………..……….…...…46 Chapter 3 3.1. Life history traits and ecological traits of fish species ……………..………......…74 Chapter 4 4.1. Statistically significant regressions of the effects of temperature on mean latitudes and boundaries, and significant changes in spatial domain ………..……….….….123 4.2. The contingency table showing numbers of shifting and non-shifting species in each order ……….……..…………………………...………………………….….124 ix 4.3. Results of logistic regressions showing effects of ecological traits and fishing on whether fish species are responsive to temperature ………...……..……….….….125 Chapter 5 5.1. Analyses of key North Pacific physical time series ….…...……..…….….….….148 5.2. Analyses of key North Pacific biological time series ………...……..…….…….148 x ACKNOWLEDGEMENTS An incredible number of people have helped during my study at Scripps by providing mentorship, friendship, assistance, and support. I am particularly grateful to people who have contributed to CalCOFI: ideas, sampling, species identifying, data management, and many other aspects. I would like to thank my committee members for their guidance and support. George Sugihara and Mark Ohman, my advisors and the greatest combination, helped me to bring math and biology together. David Checkley and Phil Hastings taught me many things about fish. Art Miller and Kaustuv Roy provided great comments on analytical approaches. I would especially like to thank John Hunter, who introduced me the fisheries problems and the CalCOFI larval fish data. Without his insights, patience, and encouragement, this thesis would not have been possible. Although not formal members of my doctoral committee, Roger Hewitt, Bill Watson, and Christian Reiss provided great mentorship on fish and fisheries. Roger provided accesses to various resources in the Southwest Fisheries Science Center, NOAA. Bill answered my every question regarding fish. Christian helped shape
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