The Response of Crustacean Zooplankton Production to Variations in Food Quantity, Quality, and Primary Production in Coastal Marine Ecosystems
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The response of crustacean zooplankton production to variations in food quantity, quality, and primary production in coastal marine ecosystems by Karyn Dawn Suchy B.Sc., University of Manitoba, 2003 M.Sc., University of Manitoba, 2006 A Dissertation Submitted in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY in the Department of Biology Karyn Dawn Suchy, 2014 University of Victoria All rights reserved. This dissertation may not be reproduced in whole or in part, by photocopy or other means, without the permission of the author. ii Supervisory Committee The response of crustacean zooplankton production to variations in food quantity, quality, and primary production in coastal marine ecosystems by Karyn Dawn Suchy B.Sc., University of Manitoba, 2003 M.Sc., University of Manitoba, 2006 Supervisory Committee Dr. John F. Dower, Department of Biology Supervisor Dr. Steve J. Perlman, Department of Biology Departmental Member Dr. Diana E. Varela, Department of Biology Departmental Member Dr. Debby Ianson, School of Earth and Ocean Sciences Outside Member iii Abstract Supervisory Committee Dr. John F. Dower, Department of Biology Supervisor Dr. Steve J. Perlman, Department of Biology Co-Supervisor or Departmental Member Dr. Diana E. Varela, Department of Biology Departmental Member Dr. Debby Ianson, School of Earth and Ocean Sciences Outside Member Crustaceans, the most abundant group of organisms that make up zooplankton, form a critical link in the food web between primary-producing phytoplankton and planktivorous fish. Examining this link is essential in order to effectively estimate the amount of energy available to higher trophic levels. The most appropriate currency for tracking energy flow through these food webs is to measure production, or the amount of new biomass generated over a given period of time. Although measurements of primary productivity are routinely made in oceanographic studies, estimates of secondary productivity are rare due to their historical reliance on time-consuming methods. The overall objective of this thesis was to determine the factors influencing temporal variations in community-level crustacean productivity. A simplified lab experiment was used to establish a relationship between diet and chitobiase-based estimates of copepod productivity in response to single versus mixed species phytoplankton diets. In addition, the relationships between primary productivity and chitobiase-based productivity for the entire crustacean zooplankton community were examined over two years in Saanich Inlet, British Columbia, Canada. Lastly, this work determined the abiotic and biotic factors most strongly influencing crustacean productivity in the tropical Guanabara Bay, Rio de Janeiro, Brazil, dominated by the microbial loop. Results from this work show that: (i) copepod populations fed a poor food item take longer to develop through early stages, have lower daily growth rates, and exhibit lower productivity than those fed a good quality food item; (ii) important variations in crustacean productivity are missed when biomass estimates, alone, are used to represent food available to higher trophic levels; (iii) relationships between primary productivity and crustacean productivity can vary interannually and are not necessarily controlled by bottom-up processes; (iv) substantial interannual variations in trophic transfer efficiency (TTE) occur even if average TTE is the same across years; and (v) community-level crustacean productivity in tropical regions dominated by the microbial food loop can be as high as, if not higher than, productivity measured in temperate regions. Ultimately, this work provides insight into how accurate productivity estimates can improve our understanding of zooplankton dynamics in both laboratory and field settings in marine ecosystems worldwide. iv Table of Contents Supervisory Committee ...................................................................................................... ii! Abstract .............................................................................................................................. iii! Table of Contents ............................................................................................................... iv! List of Tables .................................................................................................................... vii! List of Figures .................................................................................................................... ix! Acknowledgements ............................................................................................................ xi! Dedication ......................................................................................................................... xii! Chapter 1: Introduction ....................................................................................................... 1! 1.1 General introduction ................................................................................................. 1! 1.2 Methods for estimating crustacean production ......................................................... 2! 1.2.1 Traditional methods ........................................................................................... 2! 1.2.2 Global predictive models ................................................................................... 3! 1.2.3 Instantaneous methods ....................................................................................... 4! 1.2.4 Chitobiase method ............................................................................................. 5! 1.3 Factors influencing crustacean production ............................................................... 6! 1.3.1 Temperature ....................................................................................................... 6! 1.3.2 Body size/weight ................................................................................................ 7! 1.3.3 Food quantity ..................................................................................................... 8! 1.3.4 Food quality ..................................................................................................... 11! 1.4 Current limitations .................................................................................................. 13! 1.4.1 Using biomass to estimate zooplankton production ........................................ 13! 1.4.2 Chlorophyll a as a proxy for phytoplankton biomass ...................................... 14! 1.4.3 Extrapolating lab results to a natural field setting ........................................... 16! 1.5 Thesis objectives and structure ............................................................................... 18! Chapter 2: Influence of diet on chitobiase-based production rates for the harpacticoid copepod Tigriopus californicus ........................................................................................ 21! 2.1 Introduction ............................................................................................................. 21! 2.2 Methods................................................................................................................... 24! 2.2.1 Copepod cultures ............................................................................................. 24! 2.2.2 Phytoplankton cultures ..................................................................................... 25! 2.2.3 Feeding experiment .......................................................................................... 27! 2.2.4 Chitobiase enzyme assays ................................................................................ 28! 2.2.5 Chitobiase decay rate estimates ....................................................................... 29! 2.2.6 Statistical analysis ............................................................................................ 32! 2.3 Results ..................................................................................................................... 32! 2.4 Discussion ............................................................................................................... 40! 2.5 Conclusions ............................................................................................................. 46! Chapter 3: Interannual variability in the relationship between in situ estimates of primary productivity and crustacean productivity in Saanich Inlet, British Columbia, Canada .... 48! v 3.1 Introduction ............................................................................................................. 48! 3.2 Methods................................................................................................................... 51! 3.2.1 Study site .......................................................................................................... 51! 3.2.2 Physical and chemical measurements .............................................................. 53! 3.2.3 Phytoplankton .................................................................................................. 55! 3.2.4 Zooplankton ..................................................................................................... 57! 3.2.5 Trophic transfer efficiency (TTE) .................................................................... 60! 3.2.6 Statistical analysis ...........................................................................................