Predation As a Vehicle to Aid Tunicate Invasion in the Biofouling Community
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University of New Hampshire University of New Hampshire Scholars' Repository Master's Theses and Capstones Student Scholarship Fall 2013 Predation as a Vehicle to Aid Tunicate Invasion in the Biofouling Community Helen Day University of New Hampshire, Durham Follow this and additional works at: https://scholars.unh.edu/thesis Recommended Citation Day, Helen, "Predation as a Vehicle to Aid Tunicate Invasion in the Biofouling Community" (2013). Master's Theses and Capstones. 802. https://scholars.unh.edu/thesis/802 This Thesis is brought to you for free and open access by the Student Scholarship at University of New Hampshire Scholars' Repository. It has been accepted for inclusion in Master's Theses and Capstones by an authorized administrator of University of New Hampshire Scholars' Repository. For more information, please contact [email protected]. Predation as a Vehicle to Aid Tunicate Invasion in the Biofouling Community BY HELEN DAY BS University of Rhode Island, 2006 THESIS Submitted to the University of New Hampshire in Partial Fulfillment of the Requirements for the Degree of M aster o f Science In Zoology September 2013 UMI Number: 1524299 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. Di!ss0?t&iori Piiblist’Mlg UMI 1524299 Published by ProQuest LLC 2013. Copyright in the Dissertation held by the Author. Microform Edition © ProQuest LLC. All rights reserved. This work is protected against unauthorized copying under Title 17, United States Code. ProQuest LLC 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 48106-1346 This thesis has been examined and approved. Dr. Larry G. Harris (Professor, Zoology) Dr. James F. Hanay (Professor, Zoology) D. Winsor H. Watson (Professor, Zoology) Date TABLE OF CONTENTS TITLE PAGE ............................. i SIGNATURE PAGE ................................................................................. ii ACKNOWLEDGMENTS...........................................................................v LIST OF TABLES.......................................................................................vi LIST OF FIGURES................................................................................... vii ABSTRACT............................................................................................... xiv I. CHAPTER ONE - GENERAL INTRODUCTION Introduced Species as an Ecosystem Stressor ..............................1 Benthic Communities ......................................................................2 Biofouling Communities . .....................................................2 Predation and the Biofouling Community ................................... 4 Biofouling Community Development ...........................................6 Tunicates in the Biofouling Community .......................................8 Molgula citrina and Molgula manhattensis..................10 Ciona intestinalis..............................................................10 Botryllus schlosseri...........................................................11 Botryloides violaceus ....................................................... 12 Diplosoma listeranium.................. 12 Didemnum species ............................................................13 Study Objectives ............................................................................ 13 II. METHODS Introduction ....................................................................................16 Benthic Panel Community: Data Collection ..............................22 Benthic Panel Community: Sessile Invertebrates .....................24 Benthic Panel Community: Identification and Quantification of Molguloid Tunicates ...................................... 28 Benthic Panel Community: Quantification ................................30 Benthic Panel Community: Statistical Analysis ....................... 31 Laboratory Feeding Trial Overview ...........................................34 Predator Inclusion Experiments: Invertebrates ......................... 34 Predator Inclusion Experiments: Fish .........................................36 Predator Inclusion Experiments: Statistical Analysis .............. 40 Y-Maze Prey Choice Experiment ............................................... 40 Y-Maze Prey Choice Experiment: Statistical Analysis ...........41 III. RESULTS Biofouling Community Composition .........................................44 Surface Biofouling Community Composition .......................... 45 Benthic Biofouling Community Composition .......................... 47 Benthic Panel Community Composition ......................... 51 Community Composition and Seasonal Patterns ......................55 Tunicates in Panel Community Composition ........................... 61 Impact of Predation on Tunicate Community Composition. ..63 Impact of Predation on Tunicate Competition .......................... 74 Impact of Predation on Colonial and Solitary Tunicates 76 Caged Control Community Composition .................................. 78 Overall Laboratory Feeding Trial Results .................................. 79 Predators which did not consume M. citrina.............................79 Predators which consumed M. citrina........................................80 Predator Inclusion Results: Tauiogolabrus adspersus............. 80 Predator Inclusion Results: Asterias rubens..............................82 Comparison of Predator Inclusion Results ................................83 Y-Maze Prey Choice Preference Results ................................... 84 Feeding Trial Results Overview ....................................... ..........85 IV. DISCUSSION Community Composition and Successional Development 87 Impact of Predation on Community Development ...................93 Competitive Interactions between Sessile Species ...................99 Predators as Eco-system Engineers .......................................... 107 Limitations to this Study .............................................................108 Importance of this Study ................................. 109 Future Directions .......................................................................... 110 LIST OF REFERENCES........................................ 1 APPENDIX A - FEEDING TRIAL DATA............................................ 18 APPENDIX B - IACUC PERMISSION............................................... 24 ACKNOWLEDGEMENTS I would like to thank my advisor Dr. Larry Harris for all his help and guidance throughout this project. I would also like to thank my committee members Dr. James Haney and Dr. Win Watson for their help. I would like to thank the University of New Hampshire for their academic and financial support including the Summer Teaching Assistant Fellowship and the Lemer-Grey Fellowship; Finally, I would like to thank my friends and family. LIST OF TABLES Table 1: The distribution of replicates (all predator access. AA; fish access, FA; no predator access, NA) along the pier, the replicate closest to the shore is on the left. Table 3: Details of Y-Maze testing with the predator A rubens and the prey items M citrina and M. edulis. Table 4: An ethogram of behaviors anticipated to occur during Y-Maze prey choice preference experiments. Table 5: Results of Y-Maze choice test with ,4. rubens. Table 6: Results of Y-Maze choice test with A rubens. Table 7: Results of sessile invertebrate species between caged and uncaged treatment types. Table 8: Table of tunicate species found on benthic community panels. Table 9: Frequency of abundances of tunicate species between caged and uncaged treatment types. Table 10: The frequency of disturbance events which reduced the abundances of tunicate species. Table 11: Tunicate species richness on benthic community panels. Table 12: Feeding trial results with a suite of potential predators that were commonly found at the CML pier field site. Table 13: Feeding trial results for T. adspersus. Table 14: Feeding trial results with A. rubens. Table 15: Results of video analysis of A. rubens foraging and feeding behavior towards M. citrina. Table 16: Results of initial choice (i.e. first choice) in Y-Maze prey choice preference between M. citrina and M. edulis. Table 17: Results of Y Maze prey choice preference for A. rubens between M. citrina and M. edulis. vi Table 18: Feeding trial results for the predators Flabellina verrucosa, Carcinus maenas and Cyclopterus lumpus with the prey item M. citrina. Table 19: Table 19: Feeding trial results. Medium size class = 0-9.99 cm: Large size class = 10-20 cm. Table 20: Results of all feeding trials with T. adspersus. Table 21: The results ofY Maze Prey Choice Preference between M. citrina and no prey item. Table 22: Results of video analysis of A. rubens foraging and feeding behavior. Table 23: Results of feeding trials with T. adspersus. Table 24: Results of Y Maze prey choice preference for A. rubens between M. citrina and M. edulis. LIST OF FIGURES Figure 1: Arial photograph of the University of New Hampshire Coastal Marine Laboratory pier where the benthic community panels were located. Photo courtesy of The University of New Hampshire (2010). Figure 2: The location of the Gulf of Maine (GofM) on the eastern seaboard. The line of separation which can serve as a barrier to dispersal