Costa Eelgrass Thesis Part 1
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BOSTON UNIVERSITY GRADUATE SCHOOL Dissertation DISTRIBUTION, PRODUCTION, AND HISTORICAL CHANGES IN ABUNDANCE OF EELGRASS (ZOSTERA MARINA L.) IN SOUTHEASTERN MASSACHUSETTS JOSEPH EDWARD COSTA B.A., University of California, Berkeley, 1980 Submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy 1988 Approved by First Reader Ivan Valiela, Ph.0. Professor of Biology Boston University Second Reader Marshal Preanall, Ph.D. Assistant ~;ofessor of Biology Vassar College Third Reader Anne Giblin, Ph.D. Assistant Scientist Ecosystems Center, Woods Hole Acknowledgements Various parts of my thesis work were funded by the following organizations and agencies: The Kathryn Lloyd Center for Environmental Studies, the NOAA National Sea Grant College Program, Dept. of Commerce, (Grant No. NA84AA-D-00033, Woods Hole Oceanographic Institution, Proj. No. RIB-68), New England Interstate Water Pollution Control Commission (as part of a study of Buzzards Bay by the Environmental Protection Agency), the Boston University and National chapters of Sigma Xi, and the BUMP Alumni. I am indebted to numerous people. The late Tony Swain, showed me the importance of enthusiasm and critical thinking in doing science. I owe special thanks Judy Capuzzo, who not only let me use the raceways at ESL, but provided pivotal support and guidance during the early stages of my graduate career. Alan Hankin and the Lloyd Center gave both financial support, and the invaluable use of their boat and facilities. Phil Newmark, Helen Woodward, Stuart Cromarty, and many other field assistants and volunteers too numerous to mention, helped with the field sampling and laboratory processing. I have had many wonderful discussions with Ken Foreman, Bill Dennison, Bob Buchsbaum, John Wilson, Marshal Pregnall, Rainer Voigt, Gary Banta, Don Bryant, Rod Fujita, and Joe Ackerman on topics ranging from eelgrass to computers and everything in-between. Alan Poole and Sid Tamm kindly let me use their boats. Thanks to Anne Giblin, for doing sulfur analyses on several sediment cores, and to Brian Howes and John Teal for use of their GC. I want to thank my advisor, Ivan Valiela, for use of his laboratory facilities and iii many suggestions, as well as my other committee members, (Marshal Pregnall, Anne Giblin, Judy Capuzzo, and Gillian Cooper-Driver) for their valuable advice, proof reading, and moral support. My dear and loving wife Maureen gave me the strength and encouragement for the final push. She also did a wonderful job putting together the many pages of graphics and references. I dedicate this thesis to my mother, Rose Costa, whose strong character and ideals made the person I am today. EELGRASS (ZOSTERA MARINA L.) IN BUZZARDS BAY: DISTRIBUTION, PRODUCTION, AND HISTORICAL CHANGES IN ABUNDANCE (Order No. 1 Joseph Edward Costa Boston University, Graduate School, 1988 Major Professor: Ivan Valiela, Professor of Biology Abstract The past and present-day distribution of eelgrass (Zostera marina L.) was documented using aerial photographs, field surveys, nautical charts, sediment cores, and first-hand accounts. Eelgrass growth correlates with local temperature and insolation, and annual production is -350 g C m-' yr-l. In Buzzards Bay, eelgrass beds cover 41 km 2 of substrate and account for 12% of primary production; in shallow bays, eelgrass equals 40% of production. Prior to the "wasting disease" of 1931-32, eelgrass populations equaled or exceeded present-day abundance. Six to 10 years after the disease, eelgrass covered less than 10% of the present-day habitat area. The process of recolonization was similar in many areas: new beds initially appeared on bare substrates, beds expanded, new beds appeared, and some beds were removed by disturbance. A computer simulation modeled these events, and showed that rapid recolonization of eelgrass populations is highly dependant on new bed recruitment, which in nature depends on seed dispersal. High disturbance rates slow eelgrass recolonization and lower peak cover. Local changes in eelgrass abundance are driven by anthropogenic and natural disturbances which are superimposed on the regional pattern of catastrophic decline and gradual recovery. Hurricanes, ice scour, and freezing periodically destroyed eelgrass beds in some areas. Eelgrass populations in poorly flushed, developed bays, with declining water quality, never recovered from the wasting disease or showed new declines in recent years. The distribution of eelgrass is light limited, and eelgrass beds may disappear in enriched areas because of increases in algal epiphytes and phytoplankton. To identify what levels of nutrient loading cause these changes, concentrations and inputs of dissolved inorganic nitrogen (DIN) in Buttermilk Bay were measured. Periphyton on eelgrass leaves and plastic screen strips on floats correlated well to mean DIN. Experimental floats released nutrients and demonstrated that small increases in DIN significantly increase periphyton abundance. The depth of eelgrass growth in Buttermilk Bay decreased by 9 cm for every 1 increase in DIN. Periphyton abundance is more important than phytoplankton concentrations in limiting eelgrass growth in Buttermilk Bay, because water in this bay has a short residence time, and phytoplankton gradients are less prominent. Table of contents Title page i Readers approval ii Acknowledgements iii Abstract V Table of contents .. ... vii List of Tables .. .. xi List of Figures xii Overview. Introduction 1 General biology and ecology of eelgrass. 2 Chapter 1. The distribution of eelgrass (Zostera marina L.) in Buzzards Bay Introductlon 11 Methods 11 Results 19 Discussion 25 Chapter 2. Eelgrass (Zostera marina L.) production in Buzzards Bay Introduction -, ., ... .. 31 Methods . .. 3 6 vii Chapter 3. Evidence for long-term changes in eelgrass (Zostera marina L.) abundance in Massachusetts in sediment cores Introduction 6 3 Methods 6 5 Results 69 Discussion 82 Chapter 4. Historical Changes in eelgrass (Zostera marina L.) abundance in Buzzards Bay: Long term patterns and twelve case histories Introduction ~~... 86 He thods 99 Results 100 Discussion ... .. .. - . 161 Chapter 5. Hechanisms of eelgrass (Zostera marina L.) colonization: Patch dynamics and effects of disturbance 179 Introduction . 181 187 Results 199 Discussion ..... 215 viii Chapter 6. Nutrient loading in Buttermilk Bay, MA (USA): consequences of algal abundance and distribution of eelgrass (Zostera marina L.) Introduction 228 233 239 244 Chapter 7. Stress ethylene production in four marine macrophytes Abstract . 256 Introduction 258 259 -. .. ..~ ..- - ....... ........ 261 271 Chapter 8. Management considerations of eelgrass populations in . Massachusetts Resource assessment 278 Federal, state, and local laws 278 Implications of changing eelgrass abundance . 281 Future monitoring 282 Mitigation efforts 285 Future management 286 Water quality protection .. ... .. .... 289 Appendix I. Repositories of aerial photographs and nautical charts used in this study. .. .. .. .................... .... ,.. .. .. ... .,. .. , 291 Appendix 11. A detailed description of eelgrass in Buzzards Bay Introduction . 293 Results 293 Appendix 111. Alphabetized list of mapped eelgrass beds by town. 361 References Cited ... .. .. .... .. .. ..... 372 Vita 396 List of Tables Table 1-1. Key to the symbols used on the maps. ............................................... 20 Table 1-2. Eelgrass cover by town around Buzzards Bay. ...................... 22 Table 1-3. Eelgrass habitat area in Buzzards Bay compared to salt marsh area, and substrate less than 3.6 m MLW. Table 2-1. Measures of biomass in shallow, mid-depth, and deep beds. Table 2-2. Eelgrass production in Buzzards Bay compared to estimates of other producers. Table 2-3. Eelgrass production in Buttermilk Bay comapred to estimates of other producers Table 4-1. Uajor meteorological disturbances in Southeastern Massachusetts since 1938. Table 5-1. Values of simulation parameters derived from aerial images and used in the computer model. Table 6-1. Growth rate of transplanted and endemic eelgrass in Buttermilk Bay. Table 6-2. Measured rates of nutrient loading in various bays and estuaries Table 8-1. Guideline for taking aerial photographs to maximize interpretation of submerged features. 284 List of Figures Figure 0-1. General morphology of Zostera marina. .. .. , , ... , , , 4 Figure 1-1. Map of Southeastern Massachusetts. 13 Figure 1-2. Percent cover scale. 17 Figure 1-3. Haximum depth (m NLW of eelgrass in different parts of Buzzards Bay. Figure 2-1. Typical vegetative eelgrass shoot showing major anatomical features and marking technique. Figure 2-2. Hap of southeastern Hassachusetts showing where biomass samples were harvested and sites for measurement of production rates. Figure 2-3. Water temperature, surface insolation, plastochrone interval data, and PI curve for Woods Role. 1985-1986. Figure 2-4. Regional frequencies of various measures of eelgrass biomass. Figure 2-5. Log-Log plot of mean shoot weight vs shoot density. Figure 2-6. Regression between total aboveground biomass and above and belowground annual production. , ,,,. Figure 3-1. Location of sediment cores taken around Buzzards Bay and Cape Cod. Figure 3-2. Seed densities distribution in Waquoit Bay. Figure 3-3. Sediment core eelgrass seed profiles in 4 Bays. ,, xii Figure 3-4. Figure 4-1. Site names around the Westport Rivers. Figure 4-2. Changes in eelgrass bed position and flat migration north of Bailey Flat, Westport. Figure 4-3. Map showing