Harmful Algal Blooms E-Book
256,99 €
Mehr erfahren.
- Herausgeber: John Wiley & Sons
- Kategorie: Wissenschaft und neue Technologien
- Sprache: Englisch
Harmful Algal Blooms: A Compendium Desk Reference provides basic information on harmful algal blooms (HAB) and references for individuals in need of technical information when faced with unexpected or unknown harmful algal events. Chapters in this volume will provide readers with information on causes of HAB, successful management and monitoring programs, control, prevention, and mitigation strategies, economic consequences of HAB, associated risks to human health, impacts of HAB on food webs and ecosystems, and detailed information on the most common HAB species.
Harmful Algal Blooms: A Compendium Desk Reference will be an invaluable resource to managers, newcomers to the field, those who do not have easy or affordable access to scientific literature, and individuals who simply do not know where to begin searching for the information needed, especially when faced with novel and unexpected HAB events.
Edited by three of the world's leading harmful algal bloom researchers and with contributions from leading experts, Harmful Algal Blooms: A Compendium Desk Reference will be a key source of information for this increasingly important topic.
Sie lesen das E-Book in den Legimi-Apps auf:
Seitenzahl: 2090
Veröffentlichungsjahr: 2018
Ähnliche
CONTENTS
Cover
Title Page
Copyright
Dedication
List of Contributors
Acknowledgments
Introduction
References and Further General Reading
Conference Proceedings Series
Special Focused Issues of Harmful Algae
Chapter 1: Causes of Harmful Algal Blooms
1.1 Introduction
1.2 “Getting There”: The Classic Perspective on Introduced Species and Links to Cultural Eutrophication
1.3 “Being There”: Blooms and Why They Succeed
1.4 “Staying There”: Links to Physical Structure and Climate
1.5 Conclusions
Acknowledgments
References
Chapter 2: Detection and Surveillance of Harmful Algal Bloom Species and Toxins
2.1 Introduction
2.2 Organism Detection
2.3 Toxin Detection
2.4 Autonomous, In Situ Technologies
2.5 Conclusions and Future Prospects
Disclaimer
References and Further Reading
Chapter 3: Modeling Marine Harmful Algal Blooms: Current Status and Future Prospects
3.1 Introduction
3.2 Building Models to Describe Ecological Events
3.3 Limitations to What Models Can Do, and Why
3.4 Modeling T-HAB and ED-HAB Events
3.5 How Good Are Current HAB Models?
3.6 Future Modeling of T-HAB and ED-HAB: Managing Expectations
3.7 Improving Our Capabilities
Acknowledgments
References
Chapter 4: Harmful Algal Blooms and Shellfish
4.1 Introduction
4.2 Major Shellfish Poisonings
4.3 Other Toxins: Pectenotoxins (PTX) and Yessotoxins (YTX)
4.4 Emerging Shellfish Poisonings
4.5 Toxin Uptake, Accumulation, and Depuration
4.6 Shellfish Contamination in North America
4.7 Impacts on Shellfish
4.8 Conclusions and Perspectives
References and Further Reading
Chapter 5: Vulnerabilities of Marine Mammals to Harmful Algal Blooms
5.1 Introduction
5.2 Overview of Algal Toxins
5.3 Impacts of Algal Toxins Specific to Marine Mammals
5.4 Considerations for the Evaluation of HAB Toxins in Marine Mammals
Abbreviations
References and Further Reading
Chapter 6: Interactions between Seabirds and Harmful Algal Blooms
6.1 Introduction
6.2 Historical Interactions between HAB and Seabirds
6.3 Improved Monitoring and Establishment of Causality
6.4 Implications for Conservation
Note
References
Chapter 7: Food Web and Ecosystem Impacts of Harmful Algae
7.1 Introduction
7.2 Approaches, Pitfalls, Progress, and Goals
7.3 High-Biomass Algal Blooms
7.4 Emerging Recognition of the Roles of Allelochemicals
7.5 Toxigenic Algae in Aquatic Food Webs
7.6 Ecosystem-Disruptive Algal Blooms
7.7 Future Directions
Appendix A: Scientific Names for Organisms Listed by Common Name in This Chapter, Also Indicating Species Affected by Karenia brevis (Kb)
References and Further Reading
Chapter 8: Assessing the Economic Consequences of Harmful Algal Blooms: A Summary of Existing Literature, Research Methods, Data, and Information Gaps
8.1 Introduction
8.2 Overview
8.3 Research Methodologies
8.4 Sources and Types of Data
8.5 Spatial and Temporal Scopes
8.6 Nature of the Hazard
8.7 Current Research Gaps
8.8 Conclusion
Acknowledgments
References and Further Reading
Chapter 9: Public Health and Epidemiology
9.1 Introduction
9.2 What Is Public Health and Epidemiology?
9.3 HAB and Human Illness
9.4 The HAB Manager's Role in Preventing HAB-Related Illnesses
9.5 HAB-Related Stressors and Human Resilience
9.6 Conclusion
References and Further Reading
Chapter 10: Marine Biotoxin and Harmful Algae Monitoring and Management
10.1 Introduction
10.2 Identifying Sampling Program Needs
10.3 Developing a Sampling Program for Shellfish Monitoring
10.4 Developing a Sampling Program for Phytoplankton Monitoring
10.5 Monitoring Other Fisheries
10.6 Novel Approaches and Advanced Tools to Enhance Monitoring Programs
10.7 Management Considerations
10.8 Phytoplankton Sampling Protocol Examples
10.9 HAB Forecasting Links
Acknowledgments
References and Further Reading
Chapter 11: Harmful Algal Bloom Education and Outreach
11.1 Introduction
11.2 K–12 Education
11.3 Web-Based and Distance Learning Education
11.4 Citizen Science
11.5 Conclusion
References and Further Reading
Chapter 12: Prevention, Control, and Mitigation of Harmful Algal Bloom Impacts on Fish, Shellfish, and Human Consumers
12.1 Introduction
12.2 HAB Prevention
12.3 Preventing and Reducing HAB Impacts on Shellfish and Fish
12.4 HAB Controls
12.5 Mitigation of HAB
12.6 Shellfish
12.7 Fish Mariculture
12.8 Conclusions
Acknowledgments
References
Further Reading
Chapter 13: Harmful Algae Introductions: Vectors of Transfer, Mitigation, and Management
13.1 Summary
13.2 The Biogeographic Ranges of Harmful Algal Bloom Species
13.3 Vectors of Transfer
13.4 Molecular Evidence for Introductions of New Species to a Region
13.5 Prevention and Risk Reduction
13.6 Emergency Treatment Options
References
Chapter 14: Culture and Culture Collections
14.1 Introduction
14.2 Step 1: Sampling the Environment
14.3 Step 2: Processing a Field Sample in the Laboratory to Confirm Presence of the Target Organism
14.4 Step 3: From Spark to Flame
14.5 Step 4: Long-Term Perpetuation of HAB Cultures
14.6 Epilogue
Further Reading
Chapter 15: Harmful Macroalgal Blooms in a Changing World: Causes, Impacts, and Management
15.1 Introduction
15.2 Freshwater and Other Inland Macroalgae
15.3 Estuarine and Coastal Marine Macroalgae
15.4 Influences on Bloom Development
15.5 Nutrient Pollution
15.6 Uptake/Adsorption of Other Contaminants
15.7 Impacts on Human Health: Macroalgae as Substrata for Pathogens
15.8 Non-native Invasions
15.9 Ecological and Ecosystem-Level Impacts
15.10 Effects of Blooms on the Chemistry of the Oceans and the Atmosphere
15.11 Management Strategies
15.12 Economic Impacts
15.13 Recycling Macroalgae Biomass
15.14 Forecast
References and Further Reading
Chapter 16: Harmful Algal Species Fact Sheets
Alexandrium
Amphidomataceae
Aureococcus anophagefferens Hargraves et Sieburth & Aureoumbra lagunensis DeYoe et Stockwell – Brown Tides
Ceratium furca (Ehrenberg) Claparede & Lachmann
Chattonella marina
Cochlodinium – Rust Tide
Cyanobacteria
Dinophysis
Fibrocapsa japonica
Gambierdiscus
Gymnodinium catenatum
Heterosigma akashiwo
Karenia brevis (Davis) Hansen et Moestrup – Florida Red Tide
Ostreopsis
Pfiesteria piscicida Steidinger & Burkholder and Pfiesteria shumwayae Glasgow & Burkholder
Prorocentrum
Prymnesium parvum (Carter) – “Golden Algae”
Pseudo-nitzschia – seriata group; delicatissima group
Takayama
Appendix 1: Websites That Routinely Distribute Bulletins on the Presence of Harmful Algal Blooms (HAB) for Public Health
Appendix 2: State Agencies Providing Information and Updates on Toxic and Harmful Algal Blooms and Water Quality
Appendix 3: List of General Web Resources
Index
End User License Agreement
List of Tables
Table 1
Table 1
Table 2.1
Table 2.2
Table 2.3
Table 2.4
Table 3.1
Table 3.2
Table 4.1
Table 4.2
Table 4.3
Table 4.4
Table 4.5
Table 5.1
Table 5.2
Table 5.3
Table 6.1
Table 6.2
Table 6.3
Table 7.1
Table 7.2
Table 8.1
Table 8.2
Table 9.1
Table 9.2
Table 10.1
Table 10.2
Table 10.3
Table 11.1
Table 11.2
Table 12.1
Table 12.2
Table 12.3
Table 13.1
Table 13.2
Table 13.3
Table 13.4
Table 14.1
Table 15.1
List of Illustrations
Figure 1
Figure 2
Figure 1
Figure 2
Figure 3
Figure 4
Figure 5
Figure 6
Figure 1
Figure 1
Figure 1
Figure 1
Figure 2
Figure 3
Figure 1
Figure 2
Figure 1
Figure 2
Figure 3
Figure 4
Figure 5
Figure 1
Figure 2
Figure 3
Figure 1
Figure 1
Figure 2
Figure 3
Figure 1
Figure 2
Figure 1
Figure 2
Figure 1
Figure 2
Figure 1
Figure 2
Figure 1
Figure 2
Figure 1.1
Figure 1.2
Figure 1.3
Figure 1.4
Figure 1.5
Figure 2.1
Figure 2.2
Figure 2.3
Figure 2.4
Figure 2.5
Figure 2.6
Figure 2.7
Figure 2.8
Figure 2.9
Figure 2.10
Figure 2.11
Figure 2.12
Figure 2.13
Figure 2.14
Figure 2.15
Figure 2.16
Figure 2.17
Figure 2.18
Figure 2.19
Figure 3.1
Figure 3.2
Figure 3.3
Figure 3.4
Figure 3.5
Figure 3.6
Figure 6.1
Figure 6.2
Figure 7.1
Figure 7.2
Figure 7.3
Figure 7.4
Figure 8.1
Figure 8.2
Figure 8.3
Figure 9.1
Figure 9.2
Figure 11.1
Figure 11.2
Figure 11.3
Figure 11.4
Figure 11.5
Figure 11.6
Figure 11.7
Figure 12.1
Figure 12.2
Figure 12.3
Figure 12.4
Figure 12.5
Figure 12.6
Figure 12.7
Figure 12.8
Figure 12.9
Figure 12.10
Figure 12.11
Figure 12.12
Figure 12.13
Figure 15.1
Figure 15.2
Figure 15.3
Figure 15.4
Figure 15.5
Figure I.1
Guide
Cover
Table of Contents
Begin Reading
Chapter 1
Pages
iii
iv
v
xvii
xviii
xix
xx
xxi
xxii
xxiii
xxiv
xxv
xxvi
xxvii
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
209
210
211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
252
253
254
255
256
257
258
259
260
261
262
263
264
265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
302
303
304
305
306
307
308
309
310
311
312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
355
356
357
358
359
360
361
362
363
364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
410
411
412
413
414
415
416
417
418
419
420
421
422
423
424
425
426
427
428
429
430
431
432
433
434
435
436
437
438
439
440
441
442
443
444
445
446
447
448
449
450
451
452
453
454
455
456
457
458
459
460
461
462
463
464
465
466
467
468
469
470
471
472
473
474
475
476
477
478
479
480
481
482
483
484
485
486
487
488
489
490
491
492
493
494
495
496
497
498
499
500
501
502
503
504
505
506
507
508
509
510
511
512
513
514
515
516
517
518
519
520
521
522
523
524
525
526
527
528
529
530
531
532
533
534
535
536
537
538
539
540
541
542
543
544
545
546
547
548
549
550
551
552
553
554
555
556
557
558
559
560
561
562
563
564
565
566
567
568
569
570
571
572
573
574
575
576
577
578
579
580
581
582
583
584
585
586
587
588
589
590
591
592
593
594
595
596
597
598
599
600
601
602
603
604
605
606
607
608
609
610
611
612
613
614
615
616
617
618
619
620
621
622
623
624
625
626
627
628
629
630
631
632
633
634
635
636
637
638
639
640
641
642
643
644
645
646
647
648
649
650
651
652
653
654
655
656
657
658
659
660
661
662
663
664
665
666
667
668
Harmful Algal Blooms
A Compendium Desk Reference
Edited by
Sandra E. Shumway
University of Connecticut Groton, CT, USA
JoAnn M. Burkholder
North Carolina State University Raleigh, NC, USA
Steve L. Morton
NOAA National Ocean Service Charleston, SC, USA
This edition first published 2018
© 2018 John Wiley & Sons Ltd
All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, except as permitted by law. Advice on how to obtain permission to reuse material from this title is available at http://www.wiley.com/go/permissions.
The right of Sandra E. Shumway, JoAnn M. Burkholder and Steve L. Morton to be identified as the author(s) of the editorial material in this work has been asserted in accordance with law.
Registered Office(s)
John Wiley & Sons, Inc., 111 River Street, Hoboken, NJ 07030, USA
John Wiley & Sons Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK
Editorial Office
The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK
For details of our global editorial offices, customer services, and more information about Wiley products visit us at www.wiley.com.
Wiley also publishes its books in a variety of electronic formats and by print-on-demand. Some content that appears in standard print versions of this book may not be available in other formats.
Limit of Liability/Disclaimer of Warranty
While the publisher and authors have used their best efforts in preparing this work, they make no representations or warranties with respect to the accuracy or completeness of the contents of this work and specifically disclaim all warranties, including without limitation any implied warranties of merchantability or fitness for a particular purpose. No warranty may be created or extended by sales representatives, written sales materials or promotional statements for this work. The fact that an organization, website, or product is referred to in this work as a citation and/or potential source of further information does not mean that the publisher and authors endorse the information or services the organization, website, or product may provide or recommendations it may make. This work is sold with the understanding that the publisher is not engaged in rendering professional services. The advice and strategies contained herein may not be suitable for your situation. You should consult with a specialist where appropriate. Further, readers should be aware that websites listed in this work may have changed or disappeared between when this work was written and when it is read. Neither the publisher nor authors shall be liable for any loss of profit or any other commercial damages, including but not limited to special, incidental, consequential, or other damages.
Library of Congress Cataloging-in-Publication Data
Names: Shumway, Sandra E., editor. | Burkholder, JoAnn M. (JoAnn Marie), editor. | Morton, Steve L., editor.
Title: Harmful algal blooms : a compendium desk reference / edited by Sandra E. Shumway, JoAnn M. Burkholder, Steve L. Morton.
Description: Hoboken, NJ : John Wiley & Sons, 2018. | Includes index. | Identifiers: LCCN 2017040583 (print) | LCCN 2017047559 (ebook) | ISBN 9781118994696 (pdf) | ISBN 9781118994689 (epub) | ISBN 9781118994658 (cloth)
Subjects: LCSH: Toxic algae. | Algal blooms–Toxicology.
Classification: LCC QK568.T67 (ebook) | LCC QK568.T67 H372 2018 (print) | DDC 579.8–dc23
LC record available at https://lccn.loc.gov/2017040583
Cover Design: © Eric Heupel
Cover Image: © Eric Heupel
We dedicate this book to Robert R.L. Guillard and Theodore J. Smayda, our esteemed colleagues, friends, and mentors.
List of Contributors
Charles M. Adams
University of Florida
Food and Resource Economics Department
Gainesville, FL
United States
Christine J. Band-Schmidt
CICIMAR-IPN
Depto. de Plancton y Ecología Marina
La Paz, B.C.S.
México
Leila Basti
Tokyo University of Marine Science and Technology
Marine Environmental Physiology Laboratory
Department of Ocean Sciences
Tokyo
Japan
Larry E. Brand
University of Miami
Rosenstiel School of Marine and Atmospheric Science
Department of Marine Biology and Ecology
Miami, FL
United States
Margaret H. Broadwater
NOAA National Ocean Service
National Centers for Coastal Ocean Science
Stressor Detection and Impacts Division
Charleston, SC
United States
JoAnn M. Burkholder
North Carolina State University
Department of Applied Ecology
Center for Applied Aquatic Ecology
Raleigh, NC
United States
Allan D. Cembella
Alfred Wegener Institute
Helmholtz Zentrum für Polar- und Meeresforschung
Bremerhaven
Germany
Gregory J. Doucette
NOAA National Ocean Service
National Centers for Coastal Ocean Science
Marine Biotoxins Program
Charleston, SC
United States
Spencer E. Fire
Florida Institute of Technology
Biological Sciences
Melbourne, FL
United States
Kevin J. Flynn
Swansea University
College of Science
Swansea, Wales
United Kingdom
Corinne M. Gibble
University of California
Ocean Science Department
Santa Cruz, CA
United States
Patricia M. Glibert
University of Maryland
Center for Environmental Science
Horn Point Laboratory
Cambridge, MD
United States
Christopher J. Gobler
Stony Brook University
School of Marine and Atmospheric Sciences
Southampton, NY
United States
Lynn M. Grattan
University of Maryland School of Medicine
Department of Neurology
Baltimore, MD
United States
Gustaaf Hallegraeff
University of Tasmania
Institute for Marine and Antarctic Studies (IMAS)
Hobart, Tasmania
Australia
Hélène Hégaret
Institut Universitaire Européen de la Mer
Laboratoire des Sciences de l'Environnement Marin
UMR 6539 CNRS/UBO/IRD/IFREME
Plouzané
France
Philipp Hess
IFREMER
Laboratoire Phycotoxines
France
Porter Hoagland
Woods Hole Oceanographic Institution
Marine Policy Center
Woods Hole, MA
United States
Sailor Holobaugh
University of Maryland School of Medicine
Department of Neurology
Baltimore, MD
United States
Brian A. Hoover
University of California
Graduate Group in Ecology
Davis, CA
United States
Raphael Kudela
University of California, Santa Cruz
Ocean Sciences Department
Institute of Marine Sciences
Santa Cruz, CA
United States
Gregg W. Langlois
California Department of Public Health (retired)
Richmond, CA
United States
Brian E. Lapointe
Florida Atlantic University – Harbor Branch Oceanographic Institute
Marine Ecosystem Health Program
Ft. Pierce, FL
United States
Sherry L. Larkin
University of Florida
Food and Resource Economics Department
Gainesville, FL
United States
Schonna R. Manning
University of Texas at Austin
Department of Molecular Biosciences
Austin, TX
United States
Harold G. Marshall
Old Dominion University
Department of Biological Sciences
Norfolk, VA
United States
Pearse McCarron
National Research Council of Canada
Halifax, Nova Scotia
Canada
Dennis J. McGillicuddy, Jr.
Woods Hole Oceanographic Institution
Department of Applied Ocean Physics and Engineering
Woods Hole, MA
United States
Linda K. Medlin
Marine Biological Association of the United Kingdom
The Citadel
Plymouth
United Kingdom
Steve L. Morton
NOAA National Ocean Service
Marine Biotoxins Program
Charleston, SC
United States
Shauna Murray
University of Technology Sydney
Climate Change Cluster (C3)
Ultimo, NSW
Australia
Judith M. O'Neil
University of Maryland Center for Environmental Science
Horn Point Laboratory
Cambridge, MD
United States
Michael L. Parsons
Florida Gulf Coast University
Fort Meyers, FL
United States
Andrew Reich
Bureau of Environmental Health
Florida Department of Health
Tallahassee, FL
United States
J.E. (Jack) Rensel
Rensel Associates Aquatic Sciences
Arlington, WA
United States
Mindy L. Richlen
Woods Hole Oceanographic Institution
Biology Department
Woods Hole, MA
United States
Alison Robertson
University of South Alabama
and
Dauphin Island Sea Laboratory
Dauphin Island, AL
United States
Daniel L. Roelke
Texas A&M University
Department of Wildlife and Fisheries Sciences
College Station, TX
United States
Brian Sancewich
University of Florida
Food and Resource Economics Department
Gainesville, FL
United States
Joe Schumacker
Quinault Department of Fisheries
Taholah, WA
United States
Kevin G. Sellner
Hood College
Center for Coastal and Watershed Studies
Frederick, MD
United States
Sandra E. Shumway
University of Connecticut
Department of Marine Sciences
Groton, CT
United States
Mary Sweeney-Reeves
University of Georgia
Marine Extension Service and Georgia Sea Grant
Athens, GA
United States
Urban Tillmann
Alfred Wegener Institute
Bremerhaven
Germany
Mare Timmons
University of Georgia
Marine Extension Service and Georgia Sea Grant
Savannah, GA
United States
Carmelo R. Tomas
University of North Carolina–Wilmington
Center for Marine Science
Wilmington, NC
United States
Kathryn L. Van Alstyne
Western Washington University
Shannon Point Marine Center
Anacortes, WA
United States
Frances M. Van Dolah
NOAA National Ocean Service
National Centers for Coastal Ocean Science
Stressor Detection and Impacts Division
Charleston, SC
United States
Gary H. Wikfors
NOAA Fisheries Service
Northeast Fisheries Science Center
Milford, CT
United States
Acknowledgments
The production of a multiauthored book is a long and arduous task, and success depends first and foremost upon the efforts and talents of the contributors. The extraordinary talent and patience of the authors are gratefully acknowledged. The project could not have been completed without Noreen Blaschik and Elle Allen, who assisted with numerous and varied tasks, and created organization out of chaos. Eric Heupel designed the food web diagram and provided the cover artwork, and his talents made the mundane aspects of graphics not only functional, but understandable.
This book was made possible by grant #NA14NMF4270023 from the DOC/NOAA/Saltonstall-Kennedy Program to Sandra E. Shumway and Tessa L. Getchis. An executive summary of this book is available:
Getchis, T.L., and S.E. Shumway. (Eds.) 2017. Harmful Algae: An Executive Summary. Connecticut Sea Grant College Program. CTSG-17-08. 16 pp.
Introduction
Toxic microalgae and their associated blooms are regular and natural phenomena and have been recorded throughout history, yet major efforts to study their ecology, physiology, toxins, and impacts have only escalated over the past 4–5 decades as their presence and impacts have expanded globally. Harmful algal blooms (HAB) are caused by a diverse array of microalgal species, and they exert significant negative impacts on human and environmental health, economies, tourism, aquaculture, and fisheries (Figure I.1). The continuing increase in numbers of toxic and harmful algal species worldwide presents a constant threat to these entities, and to the sustainable development of coastal regions. While blooms of toxic algae have been noted in numerous historical documents, dating back centuries, the focus on HAB in North America and their impacts on human health was a relatively new phenomenon in the early 1970s, when the first conference was organized to share information on occurrences predominantly in New England and the Gulf of Mexico (see LoCicero et al., 1975).
Figure I.1
As blooms of toxic phytoplankton have continued to increase in their frequency, concentrations, and geographic distribution in marine, estuarine, and fresh waters, the amount of available literature on the topic has also continued to grow. Of the estimated 3400–4000 known species of phytoplankton, only 1–2% (60–80 species) are known to be harmful or toxic, yet their impacts can be devastating. Benthic microalgae and harmful species that do not typically “bloom” are now emerging as vectors of toxins (Chapter 16).
Consumption of contaminated seafood and exposure to contaminated water and aerial-borne toxins lead to seafood safety issues and human health hazards (Chapter 11). These episodes also impact the local economies (Chapter 10) and can cause large-scale ecological disturbances including fish and shellfish die-offs, and mortalities of marine mammals and birds. A conservative, dated estimate of societal costs associated with HAB in the United States is nearly a half-billion U.S. dollars, about half of which is linked to public health effects (Anderson et al., 2000; also see Adams and Larken, 2013; Hamilton et al., 2014; Bingham et al., 2015).
Traditionally, the vectors for toxin transfer were limited to consideration of filter-feeding bivalve molluscs (e.g., oysters, clams, scallops, and mussels), but over time they have grown to include gastropods (snails, limpets, and abalone), cephalopods (squid and octopus), crustaceans (crabs, shrimp, and lobsters), and echinoderms (sea urchins and sea cucumbers) (Chapter 5). Fish and many of these nontraditional food items have been incorporated in routine algal toxin-monitoring programs (Chapter 12) for the most common toxic syndromes such as paralytic shellfish poisoning (PSP), amnesic shellfish poisoning (ASP), neurotoxic shellfish poisoning (NSP), and diarrheic shellfish poisoning (DSP), and emerging toxins such as azaspiracids, palytoxins, yessotoxins, and pectenotoxins.
Aquaculture is the fastest growing component of the food production sector globally, and the possible contamination of aquaculture and fishery products due to microalgal toxins is a major concern for managers charged with guaranteeing safe products for human and animal consumption. This has in turn led to concerted efforts to develop more sensitive, efficient, and affordable tests for algal toxins.
Since the first international conference focused on toxic algae in 1974, there have been 16 international conferences, each of which has produced a volume of contributed papers that provide invaluable information, often at local levels that might not otherwise be made available to the community at large. Bibliographic information for these volumes is provided in the “References and Further General Reading” at the end of this Introduction.
The topic is very well studied, and there are numerous comprehensive reviews and volumes available (see “References”). The volume of published material and the exponential growth of the field over the past four decades are the impetus for the current volume – to distill the information into a useable format for managers, newcomers to the field, and those who are not familiar with the scientific literature or do not have easy or affordable access.
The worldwide number of phycotoxin-induced intoxications per year is about 60,000 cases (Gerssen et al., 2010), and, even with the advent of new and improved technologies for detection and monitoring programs, human illnesses still occur on a regular basis. An excellent summary of illnesses and deaths attributed to harmful algae is provided by Picot et al. (2011). The greatest threats are with regard to novel species and outbreaks, or areas where monitoring is not routine or does not include all edible species. As new toxins are identified and better technologies developed, monitoring programs continue to evolve. These monitoring programs are also a valuable source of long-term data sets that are currently being used in modeling efforts to predict the presence and impacts of blooms (see Chapter 3). The high variability in toxin levels between individual animals demands a comprehensive monitoring program (see Chapter 12). The increase in blooms has resulted in development of new and more cost-effective technologies for toxin detection. Among the greatest strides in recent years have been the development of “dipstick tests,” which are now routinely used in many areas as preliminary screening tools; the automatized detection of harmful species with specific molecular probes; and the migration from mouse assays to instrumental analyses (see Chapter 2). Successful management and monitoring programs have minimized cases of illnesses associated with toxic algae, and they continue to be refined.
Control, prevention, and mitigation remain topics of considerable interest, and new technologies, especially with regard to manipulated clay, continue to be pursued (Chapter 14), as do efforts to minimize the severity of economic and ecological impacts as well as to reduce threats to human health. The development of educational and outreach materials that promote public understanding and especially those targeted at focused audiences where language may be a barrier (Chapter 13) has been a major factor in engaging the general public and making them more aware of the perils and avoidance means when faced with local harmful and toxic algal blooms.
The current body of knowledge on HAB and their impacts is vast and no longer easily accessible, or understandable, to those not actively engaged in specific research arenas. The present volume is not intended to be a comprehensive review of all topics, but rather to provide basic information to those who are confronted with seemingly boundless sources of information, some conflicting or confusing, or who simply don't know where to begin searching for the information they need. These issues become more urgent when faced with unexpected blooms or known or unknown algal species and the associated risks to human health and trophic consequences in marine and aquatic habitats.
The aim of the current volume is to provide an accessible source of information and references for further investigation for individuals who may not be familiar with the scientific literature, but are in need of technical information when faced with unexpected or unknown harmful algal events.
References and Further General Reading
The available published literature on harmful algal blooms and their impacts is vast and can no longer be covered in any single publication. The goal of this book is to provide an overview for managers and newcomers to the field, and the following list provides an overview of recent publications.
Adams, C.M., and S.L. Larken. 2013. Economics of Harmful Algal Blooms: Literature Review. Report to the Gulf of Mexico Alliance. Food and Resource Economics Department, University of Florida, Gainesville: 32 p. Available at:
http://www.fred.ifas.ufl.edu/pdf/Adams-Larkin-LitRev-April2013.pdf
.
Anderson, C., S.K. Moore, M.C. Tomlinson, J. Silke, and C.K. Cusack. 2015. Living with harmful algal blooms in a changing world: strategies for modeling and mitigating their effects in coastal marine ecosystems. In:
Coastal and Marine Hazards, Risks, and Disasters
. J.F. Shroder, J.T. Ellis, and D.J. Sherman (Eds.). Elsevier Science, Amsterdam: 592 p.
Anderson, D.M., S.F.E. Boerlage, and M. Dixon (Eds.). 2017.
Harmful Algal Blooms (HABs) and Desalination: A Guide to Impacts, Monitoring and Management
. Intergovernmental Oceanographic Commission, Paris: 493 p.
Anderson, D.M., Y. Kaoru, and A.W. White. 2000. Estimated Annual Economic Impacts from Harmful Algal Blooms (HABs) in the United States. Technical Report WHOI-2000-11. Woods Hole Oceanographic Institute, Woods Hole.
AOAC International. 2012.
Official Methods of Analysis of AOAC International
, 19th ed. Official Method 2011.27. AOAC International, Gaithersburg.
Bailey, S.A. 2015. An overview of thirty years of research on ballast water as a vector for aquatic invasive species to freshwater and marine environments.
Aquatic Ecosystem Health and Management
,
18
: 261–268.
Berdalet, E., L.E. Fleming, R. Gowen, K. Davison, P. Hess, L.C. Backer, S.K. Moore, P. Hoagland, and H. Enevoldsen. 2016. Marine harmful algal blooms, human health and wellbeing: challenges and opportunities in the 21st century.
Journal of the Marine Biological Association of the United Kingdom
,
96
: 61–91.
Bingham, M., S.K. Sinha, and F. Lupi. 2015. Economic Benefits of Reducing Harmful Algal Blooms in Lake Erie. Report. Environmental Consulting and Technology, Inc.: 66 p. Available at:
http://ijc.org/files/tinymce/uploaded/Publications/Economic-Benefits-Due-to-Reduction-in-HABs-October-2015.pdf
.
Brooks, B.W., J.P. Grover, and D.L. Roelke. 2011.
Prymnesium parvum
, an emerging threat to inland waters.
Environmental Toxicology and Chemistry
,
30
: 1955–1964.
Burkholder, J.M. 1998. Implications of harmful microalgae and heterotrophic dinoflagellates in management of sustainable fisheries.
Ecological Applications
,
8
: S37–S62.
Burkholder, J.M. 2002. Cyanobacteria. In:
Encyclopedia of Environmental Microbiology
. G. Bitton (Ed.). John Wiley & Sons, New York: p. 952–982.
Burkholder, J.M. 2009. Harmful algal blooms. In:
Encyclopedia of Inland Waters
. Vol.
1
G.E. Likens (Ed.). Elsevier, Oxford: p. 264–285.
Deeds J.R., J.H. Landsberg, S.M. Etheridge, G.C. Pitcher, and S.W. Longan. 2008. Non-traditional vectors for paralytic shellfish poisoning.
Marine Drugs
,
6
: 308–348.
Etheridge, S.M. 2010. Paralytic shellfish poisoning: seafood safety and human health perspectives.
Toxicon
,
56
: 108–122.
Flynn, K.J., M. St. John, J.A. Raven, D.O.F. Skibinski, J.I. Allen, A. Mitra, and E.E. Hofmann. 2015. Acclimation, adaptation, traits and trade-offs in plankton functional type models: reconciling terminology for biology and modelling.
Journal of Plankton Research
,
37
: 683–691.
Granéli, E., B. Edvardsen, D.L. Roelke, and J.A. Hagström. 2012. The ecophysiology and bloom dynamics of
Prymnesium
spp.
Harmful Algae
,
14
: 260–270.
Hallegraeff G.M., D.M. Anderson, and A.D. Cembella. 1995.
Manual on Harmful Marine Microalgae
. IOC Manuals and Guides, Vol.
33
Intergovernmental Oceanographic Commission of UNESCO, Paris: 551 p.
Hamilton, D.P., S.A. Wood, D.R. Dietrich, and J. Puddick. 2014. Costs of harmful blooms of freshwater cyanobacteria. In:
Cyanobacteria: An Economic Perspective
. N.K. Sharma, A.K. Rai, and L.J. Stal (Eds.). John Wiley & Sons, Hoboken: p. 247–256.
Landsberg, J. 2002. The effects of harmful algal blooms on aquatic organisms.
Reviews in Fisheries Science
,
10
: 113–390.
Landsberg, J.H., K.A. Lefebvre, and L.J. Flewelling. 2014. Effects of toxic microalgae on marine organisms. In:
Toxins and Biologically Active Compounds from Microalgae
. Vol.
2
G.P. Rossini (Ed.). CRC Press, Boca Raton: p. 379–449.
Landsberg, J., F. Van Dolah, and G. Doucette. 2005. Marine and estuarine harmful algal blooms: impacts on human and animal health. In:
Oceans and Health: Pathogens in the Marine Environment
. S. Belkin and R.R. Colwell (Eds.). Springer, New York: p. 165–215.
Lassus, P., P. Bourdeau, C. Marcaillou, and P. Soudant. 2014. Phycotoxins: seafood contamination, detoxification, and processing. In:
Toxins and Biologically Active Compounds from Microalgae
. Vol.
2
G. Rossini (Ed.). Taylor and Francis Group, Boca Raton: p. 453–501.
Lassus, P., N. Chomerat, P. Hess, and E. Nezan. 2016.
Toxic and Harmful Microalgae of the World Ocean
. IOC Manuals and Guides, Vol.
68
Intergovernmental Oceanographic Commission of UNESCO, Paris: 528 p.
Lawrence, J., H. Loreal, H. Toyofuku, P. Hess, I. Karunasagar, and L. Ababouch. 2011. Assessment and Management of Biotoxin Risks in Bivalve Molluscs. FAO Fisheries and Aquaculture Technical Paper No. 551. FAO, Rome: 337 p.
Matsuyama, Y., and S.E. Shumway. 2009. Impacts of harmful algal blooms on shellfisheries and aquaculture. In:
New Technologies in Aquaculture: Improving Production Efficiency, Quality and Environmental Management
. G. Burnell and G. Allan (Eds.). Woodhead Publishing, Oxford: p. 580–609.
Picot, C., T.A. Nguyen, A.C. Roudout, and D. Parent-Massin. 2011. A preliminary risk assessment of human exposure to phycotoxins in shellfish: a review.
Human and Ecological Risk Assessment
,
17
: 328–366.
Richardson, K. 1997. Harmful or exceptional phytoplankton blooms in the marine ecosystem.
Advances in Marine Biology
,
31
: 301–385.
Roelke, D.L., A. Barkoh, B.W. Brooks, J.P. Grover, K.D. Hambright, J.W. La Claire II, P.D.R. Moeller, and R. Patino. 2016. A chronicle of a killer algae in the west: ecology, assessment and management of Prymnesium parvum blooms.
Hydrobiologia
,
764
: 29–50.
Shumway, S.E. 1990. A review of the effects of algal blooms on shellfish and aquaculture.
Journal of the World Aquaculture Society
,
21
: 65–104.
Shumway, S.E. 1995. Phycotoxin-related shellfish poisoning: bivalve molluscs are not the only vectors.
Reviews in Fisheries Science
,
3
: 1–31.
Shumway, S.E., S.M. Allen, and P.D. Boersma. 2003. Marine birds and harmful algal blooms: sporadic victims or under-reported events?
Harmful Algae
,
2
: 1–17.
Smayda, T.J. 1992. Global epidemic of noxious phytoplankton blooms and food chain consequences in large ecosystems food chain consequences in large ecosystems. In:
Food Chains, Yields, Models and Management of Large Marine Ecosystems
. K. Sherman, L.M. Alexander, and B.D. Gold (Eds.). Westview Press, Boulder: p. 275–307.
Smayda, T.J. 1997. What is a bloom? A commentary.
Limnology and Oceanography
,
42
: 1132–1136.
Tomas, C. 1997.
Identifying Marine Phytoplankton
. Academic Press, San Diego: 858 p.
Van Dolah, F.M. 2000. Marine algal toxins: origins, health effects, and their increased occurrence.
Environmental Health Perspectives
,
108
: 133–141.
Zingone, A., and H.O. Enevoldsen. 2000. The diversity of harmful algal blooms: a challenge for science and management.
Ocean and Coastal Management
,
43
: 725–748.
Conference Proceedings Series
Anderson, D.M., A.W. White, and D.G. Baden (Eds.). 1985. Toxic Dinoflagellates.
Proceedings of the 3rd International Conference on Toxic Dinoflagellate Blooms
. Elsevier Science, New York: 561 p.
Granéli, E., B. Sundstrom, L. Edler, and D.M. Anderson (Eds.). 1990. Toxic Marine Phytoplankton.
Proceedings of the 4th International Conference on Toxic Dinoflagellate Blooms
. Elsevier Science, New York: 554 p.
Hallegraeff, G.M., S.I. Blackburn, C.J. Bolch, and R.J. Lewis (Eds.). 2001. Harmful Algal Blooms 2000.
Proceedings of the 9th International Conference on Harmful Algal Blooms
. Intergovernmental Oceanographic Commission of UNESCO, Paris: 518 p.
Ho, K.-C., M.J. Zhou, Y.Z. Qi, and V. Kai (Eds.). 2010. Harmful Algae 2008.
Proceedings of the 13th International Conference on Toxic Dinoflagellate Blooms
. International Society for the Study of Harmful Algae. Environmental Publication House, Hong Kong.
Kim, H.G., B. Reguera, G.M. Hallegraeff, C.K. Lee, M.S. Han, and J.K. Choi (Eds.). 2014. Harmful Algae 2012.
Proceedings of the 15th International Conference on Harmful Algae
. International Society for the Study of Harmful Algae, 246 p.
Lassus, P., G. Arzul, E. Erard-Le Denn, P. Gentien, and C. Marcaillou-Le Baut (Eds.). 1995. Harmful Marine Algal Blooms.
Proceedings of the 6th International Conference on Toxic Dinoflagellate Blooms
. Lavoisier Publishing, Paris: 878 p.
LoCicero, V., L.A. Loeblich, and A.R. Loeblich (Eds.). 1975.
Proceedings of the 1st International Conference on Toxic Dinoflagellate Blooms
. Massachusetts Science and Technology Foundation, Wakefield: 541 p.
Mackenzie, A.L. (Ed.). 2015. Marine and Freshwater Harmful Algae.
Proceedings of the 16th International Conference on Harmful Algae
. Cawthron Institute, Nelson, New Zealand, and International Society for the Study of Harmful Algae, Wellington, New Zealand: 289 p.
Okaichi, T., D.M. Anderson, and T. Nemoto (Eds.). 1989.
Red Tides: Biology, Environmental Science, and Toxicology
.
Proceedings of the 1st International Symposium on Red Tides
. Elsevier/ North Holland, New York: 505 p.
Pagou, K.A., and G.M. Hallegraeff (Eds.). 2012.
Proceedings of the 14th International Conference on Harmful Algae
. International Society for the Study of Harmful Algae and Intergovernmental Oceanographic Commission of UNESCO, Paris.
Reguera, B., J. Blanco, M.L. Fernández, and T. Wyatt (Eds.). 1998. Harmful Algae.
Proceedings of the 8th International Conference on Harmful Algae
. Xunta de Galicia and Intergovernmental Oceanographic Commission of UNESCO, Grafisant, Santiago de Compostela.
Smayda, T., and Y. Shimizu (Eds.). 1993. Toxic Phytoplankton Blooms in the Sea.
Proceedings of the 5th International Conference on Toxic Dinoflagellate Blooms
. Elsevier Science, Amsterdam: 952 p.
Steidinger, K.A., J.H. Landsberg, C.R. Tomas, and G.A. Vargo (Eds.). 2002. Harmful Algae 2002.
Proceedings of the 10th International Conference on Harmful Algae
. Florida Fish and Wildlife Conservation Commission, Florida Institute of Oceanography and Intergovernmental Oceanographic Commission of UNESCO, St. Petersburg: 573 p.
Taylor, D.L., and H.H. Seliger (Eds.). 1978. Toxic Dinoflagellate Blooms.
Proceedings of the 2nd International Conference on Toxic Dinoflagellate Blooms
. Elsevier/North Holland: New York.
Yasumoto, T., Y. Oshima, and Y. Fukuyo (Eds.). 1996. Harmful and Toxic Algal Blooms.
Proceedings of the 7th International Conference on Toxic Phytoplankton
. Intergovernmental Oceanographic Commission of UNESCO, Paris: 586 p.
Special Focused Issues of Harmful Algae
Doucette, G.J., and C. Lee (Eds.). 2008. Recent progress on the research and management of
Cochlodinium
blooms: workshop of recent progress on the research and management of
Cochlodinium
blooms.
Harmful Algae
,
7
: 259–378.
Glibert, P.M., and J.M. Burkholder (Eds.). 2006. Ecology of
Pfiesteria
.
Harmful Algae
,
5
: 339–480.
Glibert, P.M., and K.G. Sellner (Eds.). 2005. Ecology and physiology of
Prorocentrum minimum
.
Harmful Algae
,
4
: 447–650.
Gober, C.J., and T.W. Davis (Eds.). 2016. Global expansion of harmful cyanobacterial blooms: diversity, ecology, causes, and controls.
Harmful Algae
,
54
: 1–238.
Grattan, L., and V. Trainer (Eds.). 2016. Harmful algal blooms and public health.
Harmful Algae
,
57
: 1–56.
Jeong, H.J. (Ed.). 2013. Red tides in Korea.
Harmful Algae
,
30
: S1–S144.
O'Neil, J.M., and C.A. Heil (Eds.). 2014. Nutrient dynamics of
Karenia brevis
red tide blooms in the eastern Gulf of Mexico.
Harmful Algae
,
38
: 1–140.
Shumway, S.E., and T. Smayda (Eds.). 2004. Brown tides.
Harmful Algae
,
3
: 273–246.
Veldhuis, M.J.W., and P. Wassmann (Eds.). 2005. Bloom dynamics and biological control of
Phaeocystis
: a HAB species in European coastal waters.
Harmful Algae
,
4
: 805–964.
1Causes of Harmful Algal Blooms
Patricia M. Glibert1 and JoAnn M. Burkholder2
1University of Maryland, Center for Environmental Science, Horn Point Laboratory, Cambridge, MD, USA
2North Carolina State University, Department of Applied Ecology, Center for Applied Aquatic Ecology, Raleigh, NC, USA
1.1 Introduction
Much has been written about the underlying causes of harmful algal blooms (HAB), the complex interplay of factors that lead to their proliferation, and the unique set(s) of factors contributing to blooms of different species of algae. In general, the overarching causes that have received much attention in the literature include degradation of water quality and increasing eutrophication; increasing aquaculture operations; transport of harmful species via ballast water or shellfish seeding, leading to new introductions; and climate change (e.g., Hallegraeff and Bolch, 1992; Hallegraeff, 1993; Anderson et al., 2002; Glibert et al., 2005, 2014a; Heisler et al., 2008; Wells et al., 2016; and references therein). This chapter reviews these complexities while highlighting the key role of changes in nutrients; estuarine/marine microalgal species are emphasized, and information is also included on some freshwater HAB. While some have suggested that increased monitoring or surveillance has led to a perception of an increase in HAB, there is now compelling evidence from many regions showing conclusively that increases in HAB proliferations are real, not sampling artifacts (Heisler et al., 2008).
What is a HAB? In his seminal paper, Smayda (1997a, p. 1135) stated, “What constitutes a bloom…has regional, seasonal, and species-specific aspects; it is not simply a biomass issue.…The salient criterion to use in defining whether a ‘harmful’ species is in bloom and the distinctive feature of such blooms lie not in the level of abundance, but whether its occurrence has harmful consequences.” Since the publication of that paper, biomass criteria for a few HAB species have been defined, but more generally HAB continue to be defined in terms of the extent to which they cause harmful events (fish kills), toxic events (shellfish and finfish poisoning), ecosystem disruption (nutritional and/or prey-size mismatches, such as picocyanobacterial blooms), or large biomass events (hypoxia or anoxia). In all cases, for a HAB to occur, the HAB species must be present and its biomass relative to other species in the assemblage changes, although the HAB species does not need to be dominant or in high abundance to elicit some of these effects.
In general, the factors that promote HAB can be reduced to two: changes in the rate of introductions of species to new areas and changes in local conditions leading to conditions more conducive to the growth of individual species. Environmental changes can be subtle and not all factors may change together, leading in some cases to situations where one factor may seem to be favorable, but growth is impaired due to a change in another factor. The success of an introduced species in a new environment is not ensured; instead, there must be a match of environmental factors and the species capable of exploiting the environment. As Smayda (2002) also wrote,
Anthropogenic seedings are not, in themselves, bloom stimulation events; they are only the first phase of a multi-phase process. A newly vectored, non-indigenous species is initially pioneering: it must either find an open niche or displace a niche occupant as its first step towards successful accommodation within the community.…Until colonization is achieved, alien species introduced into water masses that have been modified by cultural nutrient enrichment, water mass conditioning by aquaculture, or climatological disturbances, will not bloom. Successful colonization alone is not decisive, it usually must be accompanied at some point, or coincide with habitat disturbance – a pre-condition for many HAB occurrences. (p. 292)
Changes in environmental conditions supportive of the increasing global occurrence of HAB are predominantly anthropogenic in nature, such as changes in nutrient loads resulting from expanding human population and associated nutrient pollution from agriculture and animal operations, alterations due to human changes in fishing pressure or aquaculture development, and/or large-scale changes in flow from major water diversion projects. However, changes in environmental conditions may also be due to interactions between trophic and biogeochemical changes that occur once new species become established, or to altered abiotic parameters or physical dynamics, such as temperature and stratification that are caused by climatic changes (e.g., Sunda et al., 2006; Glibert et al., 2011; Glibert, 2015; Wells et al., 2016). The complex set of adaptive strategies associated with different species will lead to some species being more or less successful in contrasting environmental conditions (e.g., Margalef, 1978; Collos, 1986; Glibert and Burkholder, 2011; Glibert, 2015, 2016). The growth of some species can alter the biological and biogeochemical environment, in some cases changing the environment favorably for their own further growth, or for growth of other harmful species. No amount of pressure from an altered rate of species introductions will ensure success of that species in a new environment unless conditions are suitable for its growth (e.g., Smayda, 2002; Glibert, 2015). The success of HAB lies at the intersection of the physiological adaptations of the harmful algal species and/or strain (population), the environmental conditions, interaction with co-occurring organisms (both biogeochemically and trophodynamically), and physical dynamics that alter abiotic conditions and/or aggregate or disperse cells (or can alter abiotic conditions in a favorable or unfavorable manner), in turn promoting or inhibiting their growth. “Strain” is mentioned here because it is well established that there can be high intraspecific variation (strain differences) within a given harmful algal species in a wide array of traits ranging from morphology, reproductive characteristics, and nutritional preferences to toxicity (Burkholder et al., 2005; Burkholder and Glibert 2006, and references therein).
As stated by Wells et al. (2016, p. 69) in their review of HAB and climate change, for HAB to be successful, it depends on the “species ‘getting there’…‘being there’ as indigenous species…and ‘staying there’.” The same is true for nutrients and related environmental conditions. They must “get there,” often from anthropogenic sources; they must “be there”; and they must “stay there,” often through physical dynamics, changes in trophodynamics and biogeochemical processing, or climate-induced changes. Here, using the framework of getting there, being there, and staying there for both cells and nutrients and associated environmental factors, the complexity of factors influencing HAB, emphasizing the intersection of changing habitat, especially nutrient conditions, and adaptive capability of HAB are described. This chapter focuses mainly on microalgae, but also includes several examples of macroalgae. The chapter closes with some suggestions for advancement in the understanding of HAB and nutrients.
1.2 “Getting There”: The Classic Perspective on Introduced Species and Links to Cultural Eutrophication
1.2.1 Introduced Species
Transfers of species and their introductions to new areas occur frequently through various pathways. Of particular concern are ballast water introductions (e.g., Hallegraeff, 2010, and references therein; see also Chapter 13, this volume). Many harmful algal species appear to be able to maintain viability during ballast water transport, so the inoculum in the discharge area is often viable (e.g., Burkholder et al., 2007a). Ballast water exchange practices have been linked to the proliferation of previously rare or undetected harmful algae in discharge locations, such as certain toxigenic dinoflagellates in Australian waters (Hallegraeff and Bolch, 1992; Hallegraeff, 1998). Ballast water discharge can alter the abundances of harmful species and set up conditions where previously rare populations proliferate (e.g., Rigby and Hallegraeff, 1996; Forbes and Hallegraeff, 1998; Hallegraeff, 1998). While only a small percentage of introduced species have become invasive and have caused significant detrimental impact in the receiving environment (Ruiz et al., 1997), in estuaries where the problem has begun to be well studied, it has generally been difficult to separate, with certainty, native from non-native taxa (Ruiz et al., 1997). The fact that many microbial species presently have widespread distributions may reflect a long history of global transport by ships, migratory waterfowl and other animals, winds, water currents, and other mechanisms (Burkholder et al., 2007a, and references therein). The continuing effects of human activities in non-indigenous species introductions and the resulting economic and ecological impacts can be so major that entire ecosystems have been completely changed (Cohen and Carlton, 1995, 1998; Ruiz et al., 1997, 1999).
The expansion of aquaculture worldwide has created another mechanism whereby species can be transported and introduced to new areas (Hégaret et al., 2008 and references therein). Aquaculture products are often shipped worldwide, and harmful species can be carried with these products. Similarly, seed stock and feed are also shipped worldwide, creating opportunities for HAB “hitchhikers.” As will be developed in this review, once harmful algal species are introduced, many site-specific factors acting in concert – such as the available suite of nutrient supplies, climatic conditions, season, light regime, the presence of potential predators, mixing characteristics and other physical dynamics, and the presence/abundance of potential competitor microbiota – will control whether a given harmful species can successfully establish and thrive in the new area (e.g., Smith et al., 1999).
1.2.2 Anthropogenically Introduced Nutrients
Over-enrichment of coastal waters by nutrients is a major pollution problem worldwide as the result of human population growth and the production of food (agriculture, animal operations, and aquaculture) and energy (Howarth et al., 2002; Howarth, 2008; Doney, 2010). Population growth and increased food production result in major changes to the landscape, in turn increasing sewage discharges and run-off from farmed and populated lands. A major increase in use of chemical nitrogenous fertilizers began in the 1950s and is projected to continue to escalate in the coming decades (e.g., Smil, 2001; Glibert et al., 2006, 2014a). The global manufacture of nitrogen (N)-based fertilizers has, in fact, increased from < 10 million metric tonnes N per yr in 1950 to >150 million metric tonnes per yr in 2013, with 85% of all chemical fertilizers having been produced since 1985 (Howarth, 2008; Glibert et al., 2014a, and references therein). In contrast to the enormous expansion in the global use of chemical N fertilizers, use of phosphorus (P) fertilizers has shown a much smaller increase, at a rate only about a third that of N (Sutton et al., 2013; Glibert et al., 2014a). Unlike N, there is no anthropogenic synthesis of P, and all P fertilizer comes from mined sources. Of these two major agricultural nutrients, only 10–30% actually reaches human consumers (Galloway et al., 2002; Houlton et al., 2013), and more than half is lost to the environment in direct run-off and atmospheric volatilization/eventual deposition (Galloway et al., 2014).
Nearly 60% of all N fertilizer now used throughout most of the world is in the form of urea (CO[NH2]2) (Constant and Sheldrick, 1992; Glibert et al., 2006; IFA, 2014). World use of urea as a fertilizer and feed additive has increased more than 100-fold in the past four decades (Glibert et al., 2006). It is projected that from 2012 to 2017, an estimated 55 new urea manufacturing plants will be constructed worldwide, half of them in China (Heffer and Prud'homme, 2013), contributing to a further doubling of global urea use by 2050 (Glibert et al., 2006, 2014a). Urea can be a significant contributor both to total N and to the fraction used by phytoplankton in estuarine and coastal waters (McCarthy, 1972; Harvey and Caperon, 1976; McCarthy et al., 1977; Furnas, 1983; Kaufman et al., 1983; Harrison et al., 1985; Glibert et al., 1991; Kudela and Cochlan, 2000; Switzer, 2008), and the frequency of reports that urea may be used preferentially by many harmful species has increased in recent years (Glibert et al., 2006, and references therein). Urea also rapidly hydrolyzes to in water, another important N form used by phytoplankton including HAB.
The development of concentrated (confined) animal feed operations (CAFOs) near coastal waters as well as inland is another increasing, major source of nutrient pollution (Mallin, 2000; Burkholder et al., 2007b; United States Environmental Protection Agency, 2013). Animal agriculture is expanding to meet the dietary demands of an increasing population, and increasingly animal production is concentrated in large industrial feeding operations which results in dense animal populations per unit landscape area (Burkholder et al. 1997 and references therein). The high concentration of wastes per unit area, in comparison to traditional animal production practices, commonly causes contamination of adjacent waters with nutrients and associated pollutants such as suspended solids and pathogenic microorganisms (Burkholder et al., 2007b). To understand the scale of this nutrient source, as an example, in the Cape Fear River basin of North Carolina, it is estimated that there are 5 million hogs, 16 million turkeys, and 300 million chickens produced annually, yielding 82,700 tonnes of N and 26,000 tonnes of P in animal waste (Mallin et al., 2015, and references therein). The estimated “manure footprint” for the United States is about 150,000,000 tonnes (Rumpler, 2016). In China, tens of thousands of CAFOs are estimated to produce more than 40 times as much N pollution as from other types of industries (Ellis, 2008).
Aquaculture can be an important nutrient source and, depending on the size of the operation and concentrations of animals, can be regarded as an aquatic form of CAFO. Nutrient inputs from large-scale culture of finfish, shellfish, macroinvertebrates, and even macroalgae in some areas (Wang et al., 2015) are a growing concern as the importance of aquaculture in providing food supplies continues to escalate. From 1980 to 2012, world aquaculture production volume increased at an average rate of 8.6% per year, and world food fish aquaculture production more than doubled, from 32.4 million metric tonnes to 66.6 million metric tonnes (FAO, 2014). China, in particular, has sustained what has been described as a “dramatic expansion” in cultured fish production; in 2013 alone, it produced 43.5 million tonnes of food fish and 13.5 million tonnes of algae, or about two-thirds of the cultured fish and more than half of the cultured algae worldwide (FAO, 2014).
Localized impacts of “high-input/high-output” finfish and crustacean aquaculture can be severe, such as hypoxia and anoxia, nutrient over-enrichment from discharged waste food and excretory materials, and a shift in sediment biogeochemical processes and benthic communities below fish pens (Carroll et al., 2003; Bissett et al., 2006; Buschmann et al., 2006; Kawahara et al., 2009; Burridge et al., 2010; Keeley et al., 2014). Extreme water quality and habitat degradation have been documented in and around shrimp farms, in particular (Naylor et al., 1998; Páez-Osuna, 2001, and references therein). The cultured species generally has a nutrient retention of 30% or less, the remainder being excreted to the enrichment or lost as undigested feed (e.g., Bouwman et al., 2013a). Global cultured production of finfish and crustacea contributed an estimated 1.7 million tonnes of N and 0.46 million tonnes of P to receiving waters during 2008 (Verdegem, 2013). Within the relatively short period from 2000 to 2006, nutrient release from shellfish cultures increased by 2.5- to 3-fold, and much larger increases are predicted in nutrient contributions from shellfish cultures by 2050 (Bouwman et al., 2011). Aquaculture in many Asian countries is expanding at an apparently unsustainable pace. Asian aquaculture, mostly in China, now contributes nearly 90% of the total global marine aquaculture annually. During 2000–2010, nutrient release from all forms of mariculture in China collectively increased by 44% to 0.20 million tonnes of N, while estimated annual coastal N input from rivers increased by 10% to 2.7 million tonnes of N (Bouwman et al., 2013b). Similar increases were estimated for P. By 2010, Chinese mariculture contributed about 7% of total N and 11% of total P inputs to coastal seas overall, and 4% and 9% of the dissolved N and P, respectively. Various HAB have been associated with estuarine/marine aquaculture, including toxic and fish-killing algae (Wu et al., 1994; Honkanen and Helminen, 2000; Wang et al., 2008; Furuya et al., 2010), and high-biomass HAB (including macroalgae) are often linked to pond production (Alonso-Rodríguez and Páez-Osuna, 2003; Azanza et al., 2005; Wang et al., 2008).
