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Anthropogenic Nutrient Enrichment and Blooms of Harmful Micro-Algae

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Anthropogenic Nutrient Enrichment and Blooms of Harmful Micro-Algae Anthropogenic Nutrient Enrichment and Blooms of Harmful Micro-algae Anthropogenic Nutrient Enrichment and Blooms of Harmful Micro-algae Richard Gowen1, Paul Tett2, Eileen Bresnan3, Keith Davidson2, Alan Gordon4, April McKinney1, Steve Milligan5, David Mills5, Joe Silke6, Ann Marie Crooks1. A report prepared For The Department for Environment, Food and Rural Affairs September 2009 1 Fisheries and Aquatic Ecosystems Branch, Agriculture Food and Environmental Science Division, Agri-Food and Biosciences Institute, Newforge Lane, Belfast, BT9 5PX. 2 Scottish Association for Marine Science, Dunstaffnage Marine Laboratory, Oban, Argyll, PA37 1QA. 3 Marine Scotland Marine Laboratory, P.O. Box 101, 375 Victoria Road, Aberdeen AB11 9DB 4 Biometrics Branch, Applied Plant Science and Biometrics Division, Agri-Food and Biosciences Institute, Newforge Lane, Belfast, BT9 5PX. 5Centre for Environment, Fisheries & Aquaculture Science, Pakefield Road, Lowestoft, Suffolk, NR33 0HT. 6 The Marine Institute, Rinville, Oranmore, Galway, Ireland. Contents Page Summary iv Part 1 Introduction 7 1.1 Introduction and Rationale ………….…………………..….......................................................... 7 1.2 Methodology and Statistical Analysis ……………….………....................................................... 8 1.3 Phytoplankton ………………………………………………………………................................. 9 1.4 Blooms of Micro-algae ……………………..…………………………..……............................... 14 1.4.1 Introduction ……………………………………………………………................................... 14 1.4.2 Red tides, nuisance, noxious and harmful blooms .................................................................... 15 1.5 Micro-algal Blooms and Their Effects ……………………………………....................................18 1.5.1 Introduction …………………………........................................................................................ 18 1.5.2 Ecosystem effects of micro-algal blooms ................................................................................. 20 1.5.2.1 Pulse and press disturbance 1.5.2.2 Shading and smothering 1.5.2.3 Deoxygenation 1.5.2.4 Algal biotoxins 1.5.3 Micro-algal blooms and the human use of the ecosystem ........................................................ 23 1.5.3.1 Algal biotoxins and human health 1.5.3.2 Mucilage and human health 1.5.3.3 The impact of micro-algal blooms on recreation and tourism 1.5.3.4 The economic impact of blooms on Fisheries and aquaculture Part 2 An overview of Harmful Micro-algal Blooms and HAB Species in Coastal Waters of the United Kingdom and Republic of Ireland 29 2.1 General Introduction ……………………………………………………...................................... 29 2.2 Coastal Waters of the United Kingdom ……………………..……………................................... 29 2.2.1 Introduction ……………………..……………........................................................................ 29 2.2.2 Species of Alexandrium ……………………..……………...................................................... 30 2.2.3 Species of Dinophysis ……………………..……………......................................................... 33 2.2.4 Species of Pseudo-nitzschia ……………………..……………............................................... 34 2.2.5 Karenia mikimotoi ……………………..…………….............................................................. 35 2.2.6 Other HAB species …..………………..……………............................................................... 36 2.3 Coastal Waters of the Republic of Ireland ……………..………………...................................... 40 2.3.1 Introduction ..…………………………….……………..………………................................. 40 2.3.2 Species of Alexandrium ………………………………..………………................................. 40 2.3.3 Species of Dinophysis .………………………….……..……………….................................. 41 2.3.4 Species of Pseudo-nitzschia .……………….…………..………………................................. 43 2.3.5 Karenia mikimotoi .………………..………………................................................................. 44 2.3.6 Other HAB species .………………..………………................................................................ 45 Part 3 Anthropogenic Nutrient Enrichment and Harmful Algal Blooms: A Literature Review 46 3.1 Introduction ……………..………………………………….………….................................... 46 3.2 Nutrient Enrichment of Coastal Waters ………………………………….……………………… 50 3.3 Nutrient Enrichment and Blooms of Harmful Micro-algae ……..………………………………. 54 3.3.1 Introduction …………………………………………………………………………………. 54 3.3.2 The nutrient enrichment HAB hypothesis ……………………………………………… 55 3.3.2.1 Introduction i 3.3.2.2 Historical and natural occurrence of HABs 3.3.2.3 Increased environmental awareness and monitoring of coastal waters 3.3.2.4 The influence of climate change 3.3.2.5 Introductions and transfers of new species 3.4 Case Studies ……………………………………………………………………………………... 69 3.4.1 Introduction …………………………………………………………………….. 69 3.4.2 Coastal waters of China ……………………………………………………………. 70 3.4.2.1 Introduction 3.4.2.2 Coastal waters of Hong Kong 3.4.2.3 Other coastal regions of China 3.4.2.4 The influence of the seasonal monsoon and climate change 3.4.2.5 Other human pressures 3.4.2.6 Summary 3.4.3 Coastal waters of Japan …………………………………………………………... 86 3.4.3.1 Introduction 3.4.3.2 The Seto Inland Sea 3.4.3.3 Summary 3.4.4 The North Sea ……………………………………………..…………………………… 93 3.4.4.1 Introduction 3.4.4.2 Phytoplankton blooms in the wider North Sea 3.4.4.3 Phytoplankton blooms in coastal waters 3.4.4.4 Phaeocystis in the North Sea 3.4.4.5 The role of climate change and anthropogenic nutrient enrichment 3.4.4.6 Summary 3.4.5 Coastal waters of the continental United States of America ..……………………………….. 105 3.4.5.1 Introduction 3.4.5.2 Nutrient enrichment HAB relationships in coastal waters of the U.S. 3.5 Nutrient Ratios, Dissolved Organic and Particulate Nutrients …..………………………………. 109 3.5.1 Introduction ………………………………………………………………………………….. 109 3.5.2 Nutrient ratios ………………………………………………………………………………... 109 3.5.2.1 Theoretical considerations 3.5.2.2 Nitrogen to phosphorus ratio 3.5.2.3 Silicate limitation of diatom growth 3.5.3 Dissolved organic and particulate nutrients ………………………………………………….. 118 3.5.4 Nutrients and toxin production ………………………………………………………………. 120 3.6 Hypotheses Concerning the Occurrence of HABs ……………………………………………….. 122 Part 4 An Evaluation of the Current Distribution of HAB Species in UK and Irish Coastal Waters 124 4.1 Introduction ………………………………………………………..……………………………... 124 4.2 Methods ………………………………………………………………………………………….. 124 4.2.1 Nutrient data …………………………………………………………..……………………… 124 4.2.1.1 Riverine loadings 4.2.1.2 Winter nutrient concentrations 4.2.2 Phytoplankton data …………………………………………………………..……………….. 127 4.3 Statistics ………...……………………………………………………………..…………………. 128 4.4 Results ………………………………………………………………………………………….... 129 4.4.1 Statistical analyses ………………………………………............................................ 129 4.4.1.1 Data sets 4.4.1.2 Nutrient loadings and HAB species abundance 4.4.1.3 Ratios of nutrient loadings and HAB species abundance 4.4.1.4 Correlations between loadings and winter concentrations 4.4.1.5 Winter concentrations, ratios and HAB species abundance 4.4.1.6 Time-series analysis 4.4.2 The distribution of HAB species in UK and Irish coastal waters …………………………….. 138 ii 4.5 Discussion …………………………………………………………..…………………………..... 142 4.5.1 Introduction …………………………………………………………………………………... 142 4.5.2 Data sets and analysis ………………………………………………………………………… 142 4.5.3 Interpretation of results ………………………………………………………………………. 144 4.5.3.1 Introduction 4.5.3.2 HAB species abundance, nutrient loadings and winter concentrations 4.5.3.3 HAB species abundance and nutrient ratios 4.5.3.4 Time-series analysis 4.6 Conclusions …………………………………………………………………..…………………... 148 Part 5 Discussion and Synthesis 149 5.1 Introduction .……………………………………………..……………………………………….. 149 5.2 Ecohydrodynamic: Some General Principles ………………………………..…………………… 150 5.3 Ecophysiology: Phytoplankton Life Forms and Species Succession ………..…………………… 152 5.4 The Interaction between Ecohydrodynamics and Ecophysiology ….……..……………………… 154 5.4.1 Introduction ………..………………………………………………………..………………... 154 5.4.2 Small regions of restricted exchange ………..………………………………………………… 155 5.4.2.1 Introduction 5.4.2.2 Tolo and Victoria Harbour (Hong Kong) 5.4.2.3 A comparison between Tolo Harbour and small enriched RREs in UK coastal waters 5.4.3 Regional Seas ………..………………………………………………………..………………. 163 5.4.4 Summary ..………..………………………………………………………..…………………. 167 5.5 The Distribution of HAB Species in UK and Irish Coastal Waters ……………………………… 168 5.5.1 Introduction …………………………………………………………………………………… 168 5.5.2 Ecohydrodynamic conditions in UK and Irish coastal waters ..………………………………. 169 5.5.3 Species of Alexandrium ……………………………………………………………………….. 171 5.5.4 Species of Dinophysis ………………………………………………………………………… 172 5.5.5 The genus Pseudo-nitzschia ……………………………………………………………….... .. 173 5.5.6 Karenia mikimotoi ……………………………………………………………………………. 175 5.5.7 Prorocentrum minimum and P. lima ………………………………………………………….. 177 5.5.8 Lingulodinium polyedrum and Protoceratium reticulatum ………………………….……….. 179 5.5.9 Aquaculture and HABs ……………………………………………………………….………. 179 5.6 Synthesis ……………………………………………………………………………………..…… 182 5.6.1 Introduction …………………………………………………………………………………… 182 5.6.2 Does the occurrence of HABs imply eutrophication and is eutrophication always accompanied by HABs? ………………………………………………. 182 5.6.3 Has an increase in HABs been reported and is this increase real? ……………………………. 183 5.6.4 Does nutrient enrichment lead to more large-biomass HABs? ………………………………...183 5.6.5 Does nutrient enrichment lead to greater
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