Changing Jellyfish Populations: Trends in Large Marine Ecosystems
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CHANGING JELLYFISH POPULATIONS: TRENDS IN LARGE MARINE ECOSYSTEMS by Lucas Brotz B.Sc., The University of British Columbia, 2000 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in The Faculty of Graduate Studies (Oceanography) THE UNIVERSITY OF BRITISH COLUMBIA (Vancouver) October 2011 © Lucas Brotz, 2011 Abstract Although there are various indications and claims that jellyfish have been increasing at a global scale in recent decades, a rigorous demonstration to this effect has never been presented. As this is mainly due to scarcity of quantitative time series of jellyfish abundance from scientific surveys, an attempt is presented here to complement such data with non- conventional information from other sources. This was accomplished using the analytical framework of fuzzy logic, which allows the combination of information with variable degrees of cardinality, reliability, and temporal and spatial coverage. Data were aggregated and analysed at the scale of Large Marine Ecosystem (LME). Of the 66 LMEs defined thus far, which cover the world’s coastal waters and seas, trends of jellyfish abundance (increasing, decreasing, or stable/variable) were identified (occurring after 1950) for 45, with variable degrees of confidence. Of these 45 LMEs, the overwhelming majority (31 or 69%) showed increasing trends. Recent evidence also suggests that the observed increases in jellyfish populations may be due to the effects of human activities, such as overfishing, global warming, pollution, and coastal development. Changing jellyfish populations were tested for links with anthropogenic impacts at the LME scale, using a variety of indicators and a generalized additive model. Significant correlations were found with several indicators of ecosystem health, as well as marine aquaculture production, suggesting that the observed increases in jellyfish populations are indeed due to human activities and the continued degradation of the marine environment. ii Table of Contents Abstract ............................................................................................................................................... ii Table of Contents .............................................................................................................................. iii List of Tables ..................................................................................................................................... vi List of Figures................................................................................................................................... vii Acknowledgments.......................................................................................................................... viii 1. Introduction................................................................................................................................ 1 1.1 Definition of ‘Jellyfish’ ...................................................................................................... 1 1.2 Problem Statement ............................................................................................................. 2 1.3 Challenges of Studying Jellyfish Populations ................................................................ 2 1.4 Impacts of Jellyfish Blooms .............................................................................................. 4 1.5 Anthropogenic Causes of Increasing Jellyfish Populaitons .......................................... 6 1.5.1 Invasive Species ......................................................................................................... 7 1.5.2 Warming ..................................................................................................................... 7 1.5.3 Eutrophication .......................................................................................................... 10 1.5.4 Overfishing ............................................................................................................... 15 1.5.5 Bottom-Trawling ...................................................................................................... 17 1.5.6 Increased Substrate .................................................................................................. 18 1.5.7 Aquaculture .............................................................................................................. 20 1.5.8 Other Factors ............................................................................................................ 20 1.5.9 Synergistic Effects .................................................................................................... 22 1.6 Regime Shifts .................................................................................................................... 24 1.7 The Jellyfish Spiral ........................................................................................................... 25 2 The Jellyfish Index ................................................................................................................... 29 2.1 Methods ............................................................................................................................ 29 2.1.1 Large Marine Ecosystem Approach ...................................................................... 29 2.1.2 1950 Baseline ............................................................................................................. 29 2.1.3 The Jellyfish Chronicles ........................................................................................... 30 2.1.4 Data Not Included ................................................................................................... 31 2.1.5 Abundance Trend .................................................................................................... 32 2.1.6 Scoring Chronicles ................................................................................................... 33 2.1.7 Invasive Species ....................................................................................................... 34 2.1.8 Fuzzy Expert System ............................................................................................... 35 2.1.9 Uncertainty ............................................................................................................... 39 2.2 Results ............................................................................................................................... 40 2.2.1 The Jellyfish Index ................................................................................................... 40 2.2.2 Effects of Invasive Species ...................................................................................... 44 2.2.3 Effects of Jellyfish Overexploitation ...................................................................... 46 2.3 Discussion ......................................................................................................................... 48 2.3.1 Defining an ‘Increase’ .............................................................................................. 48 2.3.2 Invasive Species ....................................................................................................... 51 2.3.3 Taxonomic Concerns ............................................................................................... 51 iii 2.3.4 LME #1 – East Bering Sea ........................................................................................ 52 2.3.5 LME #2 – Gulf of Alaska ......................................................................................... 53 2.3.5.1 Invasive Species in LME #2 – Gulf of Alaska .................................................... 58 2.3.6 LME #3 – California Current .................................................................................. 58 2.3.6.1 Invasive Species in LME #3 – California Current ............................................. 65 2.3.7 LME #4 – Gulf of California .................................................................................... 67 2.3.8 LME #5 – Gulf of Mexico ......................................................................................... 67 2.3.8.1 Invasive Species in LME #5 – Gulf of Mexico ................................................... 69 2.3.9 LME #6 – Southeast U.S. Continental Shelf .......................................................... 70 2.3.9.1 Invasive Species in LME #6 – Southeast U.S. Continental Shelf ..................... 71 2.3.10 LME #7 – Northeast U.S. Continental Shelf .......................................................... 72 2.3.10.1 Invasive Species in LME #7 – Northeast U.S. Continental Shelf ................ 76 2.3.11 LME #8 – Scotian Shelf ............................................................................................ 76 2.3.12 LME #9 – Newfoundland-Labrador Shelf ............................................................. 77 2.3.13 LME #10 – Insular Pacific-Hawaiian...................................................................... 77 2.3.13.1 Invasive Species in LME #10 – Insular Pacific-Hawaiian ............................ 78 2.3.14 LME #11 – Pacific Central-American Coastal ....................................................... 79 2.3.14.1 Invasive Species in LME #11 – Pacific Central-American Coastal ............. 80 2.3.15 LME #12 – Caribbean Sea ........................................................................................ 80 2.3.15.1 Invasive Species