Coastal and Estuarine Studies Managing Editors: Malcolm J. Bowman Richard T. Barber Christopher N.K. Mooers John A. Raven Coastal and Estuarine Studies formerly Lecture Notes on Coastal and Estuarine Studies 35 E.M. Cosper V.M. Bricelj E.J. Carpenter (Eds.) Novel Phytoplankton Blooms Causes and Impacts of Recurrent Brown Tides and Other Unusual Blooms Springer-Verlag Berlin Heidelberg New York London Paris Tokyo Hong Kong Managing Editors Malcolm J. Bowman Marine Sciences Research Center, State University of New York Stony Brook, N.Y. 11794, USA Richard T. Barber Monterey Bay Aquarium Research Institute Pacific Grove, CA 93950, USA Christopher N. K. Mooers Ocean Process Analysis Laboratory Science and Engineering Research Building University of New Hampshire Durham, NH 03824, USA John A. Raven Dept. of Biological Sciences, Dundee University Dundee, 001 4HN, Scotland Contributing Editors Ain Aitsam (Tallinn, USSR) . Larry Atkinson (Savannah, USA) Robert C. Beardsley (Woods Hole, USA) . Tseng Cheng-Ken (Oingdao, PRC) Keith R. Dyer (Merseyside, UK) . Jon B. Hinwood (Melbourne, AUS) Jorg Imberger (Western Australia, AUS) . Hideo Kawai (Kyoto, Japan) Paul H. Le Blond (Vancouver, Canada) . L. Mysak (Montreal, Canada) Akira Okubo (Stony Brook, USA) . William S. Reebourgh (Fairbanks, USA) David A. Ross (Woods Hole, USA) . John H. Simpson (Gwynedd, UK) Absornsuda Siripong (Bangkok, Thailand) . Robert L. Smith (Covallis, USA) Mathias Tomczak (Sydney, AUS) . Paul Tyler (Swansea, UK) Editors E.M. Cosper V. M. Bricelj E. J. Carpenter Marine Sciences Research Center, State University of New York Stony Brook, NY 11794, USA ISBN-13:978-3-642-75282-7 e-ISBN-13:978-3-642-75280-3 001: 10.1007/978-3-642-75280-3 This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, re-use of illustrations, recitation, broadcasting, reproduction on microfilms or in other ways, and storage in data banks. Duplication of this publication or parts thereof is only permitted under the provisions of the German Copyright Law of September 9, 1965, in its version of June 24, 1985, and a copyright fee must always be paid. Violations fall under the prosecution act of the German Copyright Law. © Springer-Verlag Berlin Heidelberg 1989 Softcover reprint of the hardcover 1s t edition 1989 2131/3140-543210 - Printed on acid-free paper This volume gathers the proceedings of the invited lectures of the symposium on Novel Phytoplankton Blooms: Causes and Impacts of Recurrent Brown Tides and Other Unusual Blooms, state University of New York at Stony Brook, Stony Brook, NY, USA, October 27 & 28, 1988. Conference Themes A. Characterization of the "Brown Tide" Microalga B. Occurrence and Distribution of the "Brown Tide" C. Environmental Factors Influencing "Brown Tide" Blooms D. Other Unusual Microalgal Blooms E. Impacts of Blooms on Shellfish F. Impacts of Blooms on Finfish and Zooplankton G. Impacts of Blooms on Eelgrass H. Environmental Management Conference Organizers Elizabeth M. Cosper, V. Monica Bricelj, Malcolm Bowman, Edward J. Carpenter, Akira Okubo, Mario Vieira, William Wise Marine Sciences Research Center State University of New York at Stony Brook Stony Brook, NY, USA 11794 Conference and Proceedings Sponsorship New York Sea Grant Institute University of Rhode Island Sea Grant College Program Living Marine Resources Institute of the Marine Sciences Research Center Waste Management Institute of the Marine Sciences Research Center New York State Department of Environmental Conservation - Water Resources Division suffolk County Department of Health Services Technical Support and Editing Anita Savarese, Sigrun H. Jonasdottir and Valerie A. Philbrick PREFACE A massive phytoplankton bloom, locally termed "brown tide", suddenly appeared in Long Island marine bays in 1985, colored the water a dark brown, decimated eelgrass beds and caused catastrophic starvation and recruitment failure of commercially important bay scallop populations. These "brown tide" blooms, caused by a very small, previously undescribed chrysophyte alga, have directly affected the estuarine environments of three northeastern American states: Rhode Island, New York and New Jersey. other phytoplankton blooms such as "red tides" caused by dinoflagellates and "green tides" from chlorophytes as well as blue-green algae blooms have long been recognized and studied world wide, however, the unusual nature of these "brown tide" blooms caught the interest of many people. Scientists were particularly intrigued by the discovery of a previously unknown microalga which provided the opportunity to learn more about small microalgae, picoplankters, which are usually ignored due to the difficulty in identifying species. A symposium entitled, "Novel Phytoplankton Blooms: Causes and Impacts of Recurrent Brown Tides and Other Unusual Blooms", was convened on October 27 and 28 at the State University of New York at Stony Brook on Long Island, with 220 registrants and nearly 50 scientific papers presented by researchers from the united States as well as Europe. The conference documented unusual bloom occurrences of recent and past years on a worldwide basis as well as northeast regional recurrences of the previously unknown "brown tide" blooms. Topics addressed at the conference included: Characterization of the "Brown Tide" Microalga, Occurrence and Distribution, Growth Physiology and Dynamics, and Environmental Factors Influencing "Brown Tide" Blooms, Other Unusual Blooms Past and Present, Impacts of Blooms on Shellfish, Finfish, Zooplankton, and Eelgrass, as well as Environmental Management. L. Slobodkin delivered an intriguing banquet presentation entitled "The Null Case of the Paradox of the Plankton." An emergent theme from the conference was that the convergence of, perhaps subtle, but long term anthropogenic and eutrophication effects with aperiodic environmental conditions can be demonstrated in many instances to resul t in dramatic phytoplankton blooms; for instance, the sewering of Providence, R.I. and the red tide of 1898 (S. Nixon), duck farm effluents and VII the green tides of the 1950's (J.H. Ryther), nutrient loading and acid rain leachates in Scandinavian waters associated with red tides and unusual Chrysochromulina blooms of 1988 (E. Graneli & others, E. Paasche & others, T. Smayda), and unusual Gulf Stream meanders seeding Ptychodiscus brevis into enriched coastal waters off North Carolina (P. Tester & others). The blooming of the newly identified "brown tide" species, Aureococcus anophagefferens (J. Sieburth, P. Johnson & P. Hargraves), could in part relate to new anthropogenic influences such as different chelators in detergents (Cosper & others) or new lawn treatments (J. Sieburth). The environmental scenario for the formation of a "brown tide" bloom in the early summers of 1985 and 1986 possibly involved reduced rainfall, leading to elevated salinities in enclosed bays, followed by pulses of rain delivering specific nutrients to the bay waters (Cosper & Dennison) and increased residence time of bay waters (M. Vieira). The selection of A. anophagefferens probably related both to specific chemical conditions as well as selective grazing pressures (Smayda & Villareal, D. Caron & others, L. Duguay & others). The compilation of important information and findings to date concerning the "brown tide" blooms as well as other unusual blooms into a comprehensive volume will hopefully be valuable in developing an understanding of the underlying causes and impacts of these blooms so as to guide environmental management responses and possible mitigation measures. August 1989 Elizabeth M. Cosper Table of Contents A. Characterization of the "Brown Tide" Microalga 1. Picoplankton Ultrastructure: A Decade of Preparation for the Brown Tide Alga, Aureococcus anophagefferens. J.McN. Sieburth and P.W. Johnson 1 2. Autotrophic Picoplankton in Narragansett Bay and Their Interaction With Microplankton. P.E. Hargraves, R.D. Vaillancourt and G.A. Jolly 23 3. Immunochemical Approaches to the Identification of the Ultraplankton: Assets and Limitations. L. Campbell, L.P. Shapiro, E.M. Haugen and L. Morris 39 4. Photosynthetic Pigment Composition of the Brown Tide Alga: Unique Chlorophyll and Carotenoid Derivatives. R.R. Bidigare . . . . . . . . . . . . . 57 5. Absorption and Fluorescent Characteristics of the Brown Tide Chrysophyte. Its Role on Light Reduction in Coastal Marine Environments. C.S. Yentsch, D.A. Phinney and L.P. Shapiro 77 6. Lipid Composition and Nutritional Value of the Brown Tide Alga Aureococcus anophagefferens. V.M. Bricelj, N.S. Fisher, J.B. Guckert and F.-L.E. Chu ..... . 85 7. Dimethylsulfide Production and Marine Phytoplankton: An Additional Impact of Unusual Blooms. M.D. Keller, W.K. Bellows and R.R.L. Guillard . 101 B. Occurrence and Distribution of the "Brown Tide" 8. The Spatial and Temporal Distribution of "Brown Tide" in Eastern Long Island. R. Nuzzi and R.M. Waters 117 9. Primary Productivity and Growth Dynamics of the "Brown Tide" in Long Island Embayments. E.M. Cosper, E.J. Carpenter, and M. Cottrell 139 10. The 1985 "Brown-Tide" and the Open Phytoplankton Niche in Narragansett Bay During Summer. T.J. Smayda and T.A. Villareal 159 11. Development and Distribution of a Brown-Water Algal Bloom in Barnegat Bay, New Jersey with Perspective on Resources and Other Red Tides in the Region. P.S. Olsen 189 x 12. Immunofluorescent Detection of the Brown Tide Organism, Aureococcus anophagefferens. D.M. Anderson, D.M. Kulis and E.M. Cosper 213 C. Environmental Factors Influencing "Brown Tide" Blooms 13. Role of Environmental Variables, Specifically Organic Compounds and Micronutrients, in the Growth of the Chrysophyte Aureococcus anophagefferens. S. Dzurica, C. Lee, E.M. Cosper and E.J. Carpenter 229 14. Brown Tide Bioassay: Growth of Aureococcus anophagefferens Hargraves et Sieburth in Various Known Toxicants. R.L. Steele, L.C. Wright, G.A. Tracey and G.B. Thursby 253 15. Trophic Interactions Between Nano- and Microzooplankton and the "Brown Tide". D.A. Caron, E.L. Lim, H. Kunze, E.M. Cosper and D.M. Anderson 265 16. The Case for Meteorologically Driven Fluctuations in Residence Times of Long Island Waters Subject to Algal Blooms. M.E.C. Vieira ..................... . 295 17. Brown Tide Dynamics as a Catastrophe Model. E. Beltrami ...... . 307 lB. An Examination of the Environmental Factors Important to Initiating and Sustaining "Brown Tide" Blooms. E.M. Cosper, W. Dennison, A. Milligan, E.J. Carpenter, C. Lee, J. Holzapfel and L. Milanese 317 D. Other Unusual Microalgal Blooms 19. The Null Case of the Paradox of the Plankton. L.B. Slobodkin ..... 341 20. Gulf Stream Transport of the Toxic Red Tide Dinoflagellate Ptychodiscus brevis from Florida to North Carolina. P.A. Tester, P.K. Fowler and J.T. Turner 349 21. Zooplankton Feeding Ecology: Copepod Grazing During an Expatriate Red Tide. J.T. Turner and P.A. Tester .. 359 22. Historical Perspective of Phytoplankton Blooms on Long Island and the Green Tides of the 1950's. J. H. Ryther . . . . . . ......... . 375 23. The Chrysochromulina polylepis Bloom in Scandinavian Waters During Spring 19BB. E. Dahl, O. Lindahl, E. Paasche and J. Throndsen 3B3 XI 24. From Anoxia to Fish Poisoning: The Last Ten Years of Phytoplankton Blooms in Swedish Marine Waters. E. Graneli, P. Carlsson, P. Olsson, B. Sundstrom, W. Graneli and O. Lindahl ........... . 407 25. An Extraordinary Red Tide and Fish Kill in Narragansett Bay. S.W. Nixon 429 26. Primary Production and the Global Epidemic of Phytoplankton Blooms in the Sea: A Linkage? T . J. Smayda . . . . . . . . . . . . . . . 449 E. Impacts on Shellfish 27. Overview of Bay Scallop (Argopecten irradians) Landings. J. L. McHugh . . . . . . .. . ..... . 485 28. The Feeding Physiology and Growth of Bay Scallops and Mussels. V.M. Bricelj and S.H. Kuenstner .. 491 29. Effects of the Brown Tide Alga on Growth, Feeding Physiology and Locomotory Behavior of Scallop Larvae (Argopecten irradians). S.M. Gallager, D.K. Stoecker and V.M. Bricelj ...... . 511 30. Are Metabolites from the Brown Tide Alga, Aureococcus anophagefferens, Deleterious to Mussel Feeding Behavior? J.E. Ward and N.M. Targett ....... . 543 31. Testing and Application of Biomonitoring Methods for Assessing Environmental Effects of Noxious Algal Blooms. G.A. Tracey, R.L. Steele, J. Gatzke, D.K. Phelps, R. Nuzzi, M. Waters and D.M. Anderson .. 557 32. Detrimental Biological Effects of Phytoplankton Blooms Deserve Increased Attention. J.B. Mahoney ............. . 575 F. Impacts of Blooms on Finfish and Zooplankton 33. Abundance and Distribution of Zooplankton and Ichthyoplankton in Great South Bay, New York During the Brown Tide Outbreaks of 1985 and 1986. L.E. Duguay, D.M. Monteleone and C.E. Quaglietta 599 34. Effect of the "Brown Tide" on Feeding, Size and Egg Laying Rate of Adult Female Acartia tonsa. A.G. Durbin and E.G. Durbin ... 625 35. Potential Change in the Distribution of Larval Fish Within Great South Bay, New York in Response to Recurrent Phytoplankton Blooms. M. Shima and R.K. Cowen . . . . . . . . . . . . . . . . . . . . . . 647 XII 36. Growth Rates of Bay Anchovy (Anchoa mitchilli) in Great South Bay Under Recurrent Brown Tide Conditions, Summers 1987 and 1988. L.R. Castro and R.K. Cowen 663 G. Impacts of Blooms on Eelgrass 37. Effect of "Brown Tide" Shading on Eelgrass (Zostera marina L.) Distributions. W.C. Dennison, G.J. Marshall and C. Wigand 675 38. Simple Model of Eelgrass Growth and Water Quality: Another Catastrophe Theory? A. Okubo and J. Slater 693 H. Environmental Management 39. Winter Burial of Transplanted Bay Scallops. S.T. Tettelbach, C.F. Smith, J.E. Kaldy, III, T. W. Arroll and M.R. Denson ....... . 713 40. The Fishery Manager's Perspective on Unusual Algae Blooms. G.C. Colvin and K.L. Koetzner ......... . 735 41. Brown Tide Comprehensive Assessment and Management Program. V.A. Minei ........ . 741 42. Water Quality Monitoring and Modeling for the Peconic Bay BTCAMP. J.R. Pagenkopf and M.R. Morton, A. Stoddard and E.D. Santoro .............. . 761 I. Postscript 43. Epilogue to the 2nd Brown Tide Conference: Are Aureococcus and Other Nuisance Algal Blooms Selectively Enriched by the Runoff of Turf Chemicals? J.McN. Sieburth ....... . 779 44. The Possible Role of Lawn Fertilizers and Pesticide Use in the Occurrence of the Brown Tide. V. Minei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 785 J. Index ...................................... 787