An Overview of Risk Assessment in a Marine Biosecurity Context

Contents Section I 1 Perspectives on Marine Invasions Marine Bioinvasions: Conservation Hazards and Vehicles for Ecological Understanding ................................................................. Gil Rilov and Jeffrey A. Crooks 1.1 1.2 1.3 Introduction – The Problem of Biological Invasions ........................ Invasions as a Tool to Study Nature .................................................. Invasion Biology – The Discipline and its Application in the Marine Environment ............................................................... 1.4 Structure of the Book ........................................................................ 1.5 Note on Terminology ........................................................................ 1.6 Concluding Thoughts ........................................................................ References .................................................................................................. 2 Deep Invasion Ecology and the Assembly of Communities in Historical Time..................................................................................... James T. Carlton 2.1 2.2 2.3 2.4 2.5 Introduction ....................................................................................... Invader Underestimation – Systematics ............................................ 2.2.1 Cryptogenic Species ............................................................. 2.2.2 Pseudoindigenous Species .................................................... Invader Underestimation – Biogeographic and Community History.................................................................... 2.3.1 Widespread Intraoceanic and Interoceanic Corridor Species ................................................................... 2.3.2 Neritic Species with Presumptive Oceanic Dispersal ........... 2.3.3 Resident Species ................................................................... Invader Underestimation – Sampling................................................ 2.4.1 Species in Underexplored Habitats and Associations........... 2.4.2 Incipient Invasions: Species with Small Population Sizes.................................................................... The Overestimation of Invader Diversity .......................................... 3 3 5 6 7 8 9 9 13 13 14 14 16 39 39 40 41 42 42 42 43 vii viii 3 Contents 2.6 Discussion ......................................................................................... 2.7 The Way Forward: Solutions ............................................................ References .................................................................................................. 44 47 48 Natural and Climate Change Mediated Invasions ................................ Steve I. Lonhart 57 3.1 3.2 3.3 57 57 58 Introduction ....................................................................................... The Geographic Range of a Species ................................................. Range Shifts ...................................................................................... 3.3.1 Factors that Influence our Understanding of the Geographic Range and Range Limits ......................... 3.3.2 Natural Range Shifts ............................................................. 3.3.3 Human-mediated Range Shifts ............................................. 3.4 Climate Change and Range Shifts .................................................... 3.4.1 Observed Biological Responses to Climate Change............. 3.5 Contrasting Natural Range Expansions and Biological Invasions ................................................................... 3.6 Conclusions ....................................................................................... References .................................................................................................. 4 Modeling Marine Invasions: Current and Future Approaches ........... Marjorie J. Wonham and Mark A. Lewis 4.1 Introduction ....................................................................................... 4.1.1 Why Marine Invasion Modeling? ......................................... 4.1.2 Scope of this Review ............................................................ 4.2 Invasion Pathway Models ................................................................. 4.3 Population Models: Invasion Dynamics ........................................... 4.3.1 Single-species Models .......................................................... 4.3.2 Multi-species Models............................................................ 4.4 Population Models: Invasion Spread ................................................ 4.4.1 Single-species Models .......................................................... 4.4.2 Allee Effects ......................................................................... 4.4.3 Multi-species Models............................................................ 4.5 Community Invasibility Models ....................................................... 4.6 Summary and Future Directions ....................................................... 4.6.1 Formalizing Conceptual Models Mathematically................. 4.6.2 Coupling Dynamical and Statistical Models ........................ 4.6.3 Integrating Modeling and Empirical Work ........................... References .................................................................................................. Appendix .................................................................................................... 58 60 62 63 63 66 67 67 71 71 71 72 73 74 79 80 84 85 87 90 93 94 94 96 97 98 105 Contents ix Section II Invader Arrival 5 6 7 Characterizing Vectors of Marine Invasion........................................... Dan Minchin, Stephan Gollasch, Andrew N. Cohen, Chad L. Hewitt, and Sergej Olenin 109 5.1 5.2 5.3 Introduction ....................................................................................... Primary vs Secondary Introductions ................................................. Principal Vectors of Marine Invasion ................................................ 5.3.1 Shipping ................................................................................ 5.3.2 Canals ................................................................................... 5.3.3 Aquaculture........................................................................... 5.3.4 Fisheries ................................................................................ 5.3.5 Ornamental Species and Live Seafood ................................. 5.3.6 Marine Leisure and Tourism ................................................. 5.3.7 Research and Education ........................................................ 5.3.8 Habitat Restoration and Management .................................. 5.4 Vector Management .......................................................................... References .................................................................................................. 109 109 110 110 110 111 111 112 112 112 113 113 115 The Vessel as a Vector – Biofouling, Ballast Water and Sediments........................................................................................... Chad L. Hewitt, Stephan Gollasch, and Dan Minchin 117 6.1 Introduction ....................................................................................... 6.2 Biofouling ......................................................................................... 6.3 Ballast Water and Sediments............................................................. 6.4 Discussion ......................................................................................... References .................................................................................................. 117 120 124 126 129 The Role of Propagule Pressure in Invasion Success ............................ Emma L. Johnston, Richard F. Piola, and Graeme F. Clark 133 7.1 7.2 7.3 7.4 7.5 7.6 7.7 133 133 135 138 139 141 142 143 145 146 147 Introduction ....................................................................................... Propagule Pressure ............................................................................ The Resilience of Propagules............................................................ Producing Propagules ....................................................................... Inferring Propagule Supply from Current Distributions ................... Vectors and Propagule Supply .......................................................... Manipulating Propagule Supply ....................................................... 7.7.1 Manipulating Propagule at the Larval Stage......................... 7.7.2 Manipulating Adult Invaders ................................................ 7.8 Conclusions ....................................................................................... References .................................................................................................. x Contents 8 Differentiating Successful and Failed Invaders: Species Pools and the Importance of Defining Vector, Source and Recipient Regions............................................................... A. Whitman Miller and Gregory M. Ruiz 8.1 8.2 Introduction................................................................................... Identifying the Roles of Source Region, Recipient Region and Vector......................................................................... 8.3 Modeling Invasion State Space – Defining and Comparing Species Pools ...................................................... 8.3.1 Species Pool Designations ............................................. 8.3.2 Species Pool Designations and Comparisons (Fixed Recipient, Source, and Vector)............................ 8.3.3 Fixed Recipient and Source Regions, Multiple Vectors ............................................................. 8.3.4 Fixed Recipient Region, Multiple Source Regions, Multiple Vectors .............................................. 8.3.5 Multiple Recipient Regions, Fixed Source Region, Multiple Vectors ................................................ 8.4 Some Recent Analyses of Invader Attributes Using Species Pool Comparisons ......................................................................... 8.5 Discussion ..................................................................................... References ................................................................................................ Section III 9 The Establishment of Invasive Species ................................................. Jeffrey A. Crooks and Gil Rilov Introduction................................................................................... Factors Influencing Numeric and Geographic Growth of Invasive Populations ................................................................. References ................................................................................................ The Role of Phenotypic Plasticity in Marine Biological Invasions.................................................................................................. L. David Smith 10.1 10.2 10.3 10.4 153 154 156 157 159 160 162 163 163 167 169 Invader Establishment 9.1 9.2 10 153 Introduction................................................................................... Phenotypic Plasticity .................................................................... 10.2.1 A Brief Overview ........................................................... 10.2.2 General Importance to Invasions .................................... 10.2.3 Empirical Work in Non-marine Systems........................ Phenotypic Plasticity in Marine Systems ..................................... Where Phenotypic Plasticity Might Play a Role in Marine Invasions ........................................................................... 173 173 173 174 177 177 178 178 180 182 183 187 Contents 10.4.1 Habitats........................................................................... 10.4.2 Taxonomic Groups ......................................................... 10.4.3 Steps in the Invasion Sequence ...................................... 10.4.4 Vectors ............................................................................ 10.5 Examples of Phenotypic Plasticity’s Effect in Marine Invasions....................................................................... 10.5.1 Post-invasion Effects ...................................................... 10.5.2 The Invasion of Carcinus maenas in the Gulf of Maine ................................................................. 10.6 Future Research Directions........................................................... References ................................................................................................ 11 12 xi 187 188 188 189 191 191 192 194 196 Escape from Parasites ............................................................................ Mark E. Torchin and Kevin D. Lafferty 203 11.1 Introduction................................................................................... 11.2 Reasons for Parasite Escape ......................................................... 11.3 Consequences of Parasite Release ................................................ 11.4 Fishes ............................................................................................ 11.5 Molluscs ........................................................................................ 11.6 Crabs ............................................................................................. 11.7 Conclusion .................................................................................... References ................................................................................................ 203 205 207 207 208 210 210 211 Ecological Factors Affecting Community Invasibility ........................ Suzanne V. Olyarnik, Matthew E.S. Bracken, Jarrett E. Byrnes, A. Randall Hughes, Kristin M. Hultgren, and John J. Stachowicz 215 12.1 12.2 12.3 12.4 215 216 218 219 221 224 225 225 227 227 228 231 232 233 Introduction................................................................................... The Abiotic Filter ......................................................................... Resource Levels ............................................................................ Biotic Resistance and Species Diversity ....................................... 12.4.1 Experimental vs Observational Approaches .................. 12.4.2 Large- vs Small-scale Studies ........................................ 12.5 Facilitation .................................................................................... 12.5.1 Increasing Resource Availability .................................... 12.5.2 Ameliorating Physical Stress.......................................... 12.5.3 Facilitation Cascades and “Invasional Meltdown” ......... 12.6 Disturbance ................................................................................... 12.7 Historical Context ......................................................................... 12.8 Conclusion .................................................................................... References ................................................................................................ xii Contents Section IV Invader Integration into Ecosystems 13 14 15 The Integration of Invasive Species into Marine Ecosystems ............ Gil Rilov 241 13.1 Introduction ................................................................................... 13.2 Are Most Marine Invasions Ecologically Harmless?.................... 13.3 Ecological Interactions of Marine Invaders .................................. 13.4 Conclusion .................................................................................... Reference ................................................................................................. 241 241 243 244 244 Competition in Marine Invasions ......................................................... James E. Byers 245 14.1 14.2 Introduction ................................................................................... A Review of the Competition Literature on Marine Invasive Species ............................................................................ 14.2.1 What Exotic Taxa are Studied for Competitive Interactions? ................................................................... 14.2.2 What Regions and Habitats are Studied?........................ 14.2.3 How Often Does Competition Occur and How Are Its Effects Measured/Indexed? ................. 14.3 Should Competition be Stronger in Human Mediated Invasions? ..................................................................... 14.4 Caveats to the Conclusion of Strong and Frequent Competition in the Marine Invasion Literature ............................ 14.5 Future Directions .......................................................................... 14.6 Conclusion .................................................................................... References ................................................................................................ 245 Predator-Prey Interactions of Marine Invaders.................................. Gil Rilov 261 15.1 15.2 15.3 261 261 Introduction................................................................................... The Role of Predation in Marine Communities ............................ Predator-prey Interactions in Invaded Systems: A Literature Review...................................................................... 15.3.1 Descriptive Statistics of Predator-prey Interactions in Invaded Systems ..................................... 15.3.2 Invasions of Predatory Zooplankton............................... 15.3.3 Predation as an Invasion-control Mechanism................. 15.3.4 Predation by Invaders as a Facilitative Force for Further Invasions ...................................................... 15.4 Conclusion .................................................................................... References ................................................................................................ 246 247 249 249 252 254 256 257 258 264 265 274 276 278 279 281 Contents 16 The Role of Exotic Marine Ecosystem Engineers ............................... Jeffrey A. Crooks 287 16.1 16.2 287 289 289 Ecosystem-Level Impacts ............................................................. Engineering ................................................................................... 16.2.1 Autogenic vs Allogenic Engineering ............................. 16.2.2 Indirect Effects of Engineering on Food Webs and Nutrient Cycling ...................................................... 16.3 Detecting Engineering Effects ...................................................... 16.3.1 Effects on Abiotic Environment Alone .......................... 16.3.2 Comparisons of Invaded and Uninvaded Areas ............. 16.3.3 Experimental Approaches .............................................. 16.4 Effects of Exotic Engineers on Abiotic Factors and Processes ................................................................................ 16.4.1 Water Column and Hydrodynamic Conditions .............. 16.4.2 Sedimentation and Benthic Conditions .......................... 16.4.3 Plants and Animals as Agents of Structural Change....................................................... 16.4.4 Net Effects of Exotic-induced Changes to Habitat Complexity .................................................... 16.4.5 Scale ............................................................................... 16.5 Implications .................................................................................. References ................................................................................................ 17 xiii Multitrophic Effects of Invasions in Marine and Estuarine Systems........................................................................... Edwin D. Grosholz and Gregory M. Ruiz 17.1 17.2 17.3 Introduction................................................................................... Multitrophic Impacts .................................................................... Case History #1: The Ecosystem Engineer Spartina in West Coast Estuaries ........................................................................................ 17.4 Case History #2: Contrasting Trophic Effects of a Non-Native and Native Predator in a Central California Estuary ......................................................................... 17.5 Invasions as Disturbance: A Framework for Understanding Multitrophic-level Impacts............................................................ 17.6 Conclusions................................................................................... References ................................................................................................ 291 292 292 293 293 294 295 295 296 297 298 299 300 305 305 306 310 313 315 319 320 xiv Contents Section V 18 Management Perspectives Marine Bioinvasion Management: Structural Framework ............... Chad L. Hewitt, Richard A. Everett, Naomi Parker, and Marnie L. Campbell 327 18.1 18.2 18.3 327 327 Introduction................................................................................... Inaction vs Action ......................................................................... Marine Biosecurity – Prevention, Regulation, and Management of Invasions ...................................................... 18.4 Other Management Options .......................................................... References ................................................................................................ Additional Literature................................................................................ Ballast Water ............................................................................................ Biofouling ................................................................................................ Biological Control.................................................................................... 19 Examples of Current International, Regional and National Regulatory Frameworks for Preventing and Managing Marine Bioinvasions............................................................................... Chad L. Hewitt, Richard A. Everett, and Naomi Parker 19.1 19.2 20 328 330 331 332 332 333 333 335 Introduction................................................................................... International Frameworks ............................................................. 19.2.1 General Obligations........................................................ 19.2.2 International Quarantine Agreements ............................ 19.2.3 Convention on Biological Diversity ............................... 19.2.4 Vector-based International Instruments .......................... 19.3 Regional Multi-lateral Frameworks .............................................. 19.3.1 Helsinki Commission ..................................................... 19.3.2 Asia-Pacific Economic Cooperation .............................. 19.4 National Examples ........................................................................ 19.4.1 Australia ......................................................................... 19.4.2 New Zealand................................................................... 19.4.3 USA ................................................................................ 19.5 Conclusions................................................................................... References ................................................................................................ 335 336 336 337 338 339 341 342 342 343 343 344 346 348 349 An Overview of Risk Assessment in a Marine Biosecurity Context................................................................................ Marnie L. Campbell 353 20.1 Introduction................................................................................... 20.1.1 Defining Endpoints ......................................................... 20.1.2 Core Values..................................................................... 353 354 354 Contents xv 20.1.3 Uncertainty and the Precautionary Principle/Approach......................................................... 20.1.4 Quantitative vs Qualitative ............................................. 20.2 Types of Risk Assessment ............................................................ 20.3 Species Level Risk Assessment .................................................... 20.3.1 Species Level Risk Assessment Examples ..................... 20.4 Vector Based Risk Assessment ..................................................... 20.4.1 Vector Based Risk Assessment Examples ...................... 20.5 Pathway Risk Assessment ............................................................ 20.5.1 Pathway Risk Assessment Examples ............................. 20.6 Conclusions................................................................................... References ................................................................................................ 21 Differing Consequences of Removing Ecosystem-Modifying Invaders: Significance of Impact and Community Context to Restoration Potential ......................................................................... Sally D. Hacker and Megan N. Dethier 21.1 21.2 Introduction................................................................................... A Predictive Conceptual Model for Post-removal Restoration .................................................................................... 21.3 Context Dependent Restoration: Invasive English Cordgrass as an Example .............................................................. 21.4 Conclusions................................................................................... References ................................................................................................ 355 356 357 358 358 361 362 367 367 369 370 375 375 377 380 383 383 Section VI Geographic Perspectives 22 Geographic Perspectives of Invasions in the Sea................................. Jeffrey A. Crooks and Gil Rilov 389 22.1 22.2 389 Introduction................................................................................... The Emerging Global Picture of Marine Invasion – Possibilities and Precautions ....................................... References ................................................................................................ 23 The Status and Distribution of Marine Alien Species in South Africa ....................................................................................... Charles L. Griffiths, Tamara B. Robinson, and Angela Mead 23.1 23.2 23.3 23.4 23.5 23.6 Introduction ................................................................................. Cnidaria ....................................................................................... Polychaeta ................................................................................... Cirripedia .................................................................................... Isopoda ........................................................................................ Amphipoda.................................................................................. 389 390 393 393 393 396 396 396 396 xvi 24 25 26 Contents 23.7 Decapoda..................................................................................... 23.8 Bryozoa ....................................................................................... 23.9 Bivalvia ....................................................................................... 23.10 Gastropoda .................................................................................. 23.11 Echinoidea................................................................................... 23.12 Ascidiacea ................................................................................... 23.13 Pisces........................................................................................... 23.14 Rhodophyta ................................................................................. 23.15 Patterns and Adequacy of Data ................................................... 23.16 Additional Cryptogenic Species ................................................. References ................................................................................................ 397 397 398 400 401 401 402 402 402 405 406 Marine Invasions in New Zealand: A History of Complex Supply-Side Dynamics ........................................................................... Barbara J. Hayden, Graeme J. Inglis, and David R. Schiel 409 24.1 Introduction ................................................................................. 24.2 Status of Marine Invasions in NZ ............................................... 24.3 What is Non-native?.................................................................... 24.4 Evidence of the Leaky Border .................................................... 24.5 Reasons Why Introductions Continue ........................................ 24.6 Most Common Marine Pathways................................................ 24.6.1 Ballast Water ................................................................. 24.6.2 Hull Fouling and Sea Chests ......................................... 24.7 The Dynamic Nature of Incursion Risk ...................................... 24.8 Changes in the Source of Invaders .............................................. 24.9 The Complexity of Making Forecasts ......................................... 24.10 Conclusion .................................................................................. References ................................................................................................ 409 410 411 413 413 414 414 415 415 416 418 421 421 Marine Bioinvasions in Australia ......................................................... Cathryn Sliwa, Sasha Migus, Felicity McEnnulty, and Keith R. Hayes 425 25.1 Introduction ................................................................................... 25.2 Non-native and Cryptogenic Marine Species in Australia............ 25.3 Problems with Lists of Non-Native and Cryptogenic Species ........ References ................................................................................................ 425 428 433 435 Marine Bioinvasions in the Southeastern Pacific: Status, Ecology, Economic Impacts, Conservation and Management ........... Juan C. Castilla and Paula E. Neill 439 26.1 26.2 439 440 Introduction................................................................................... Background Information on the Chilean Marine System ............. Contents 26.3 27 Marine NIS Introductions in Chile ............................................... 26.3.1 Likely Pathways of Marine NIS Introductions............... 26.3.2 Likely Donor Areas of Marine NIS ................................ 26.4 Socio-economic Impacts of NIS ................................................... 26.4.1 Positive Impacts.............................................................. 26.4.2 Negative Impacts ............................................................ 26.5 Ecology and Conservation of Marine NIS in Chile ...................... 26.6 Marine NIS in Peruvian Waters .................................................... 26.7 Comparative Analysis of NIS in Chile ......................................... 26.8 Marine NIS Regulations in Chile.................................................. 26.9 Conclusions and Recommendations ............................................. References ................................................................................................ 441 445 446 447 447 448 448 450 450 452 453 454 Marine Bioinvasions in the Brazilian Coast: Brief Report on History of Events, Vectors, Ecology, Impacts and Management of Non-indigenous Species ..................................................................... Carlos Eduardo Leite Ferreira, Andrea de Oliveira Ribeiro Junqueira, Maria Célia Villac, and Rubens Mendes Lopes 459 27.1 27.2 Introduction................................................................................... The Plankton Realm ..................................................................... 27.2.1 Phytoplankton Species ................................................... 27.2.2 Zooplankton Species ...................................................... 27.3 The Benthic Realm ....................................................................... 27.3.1 Phytobenthos .................................................................. 27.3.2 Zoobenthos ..................................................................... 27.4 Fishes ............................................................................................ 27.5 Intentional Introductions Related to Marine Farming .................. 27.6 Final Remarks ............................................................................... References ................................................................................................ 28 xvii Four Centuries of Biological Invasions in Tidal Waters of the Chesapeake Bay Region .............................................................. Paul W. Fofonoff, Gregory M. Ruiz, Anson H. Hines, Brian D. Steves, and James T. Carlton 28.1 28.2 Introduction................................................................................... Description of the Chesapeake Bay Region ................................. 28.2.1 Physical Features ............................................................ 28.2.2 History of Biological Studies ......................................... 28.3 Patterns of Invasion in the Chesapeake Bay Region ..................... 28.3.1 Taxonomic Composition and Residency ........................ 28.3.2 Changing Patterns of Invasion Over Time ..................... 28.4 Conclusions................................................................................... References ................................................................................................ 459 461 461 462 463 463 464 470 471 472 473 479 479 481 481 482 483 483 486 497 502 xviii 29 Contents Introduced Aquatic Species of the North Sea Coasts and Adjacent Brackish Waters ............................................................. Stephan Gollasch, Deniz Haydar, Dan Minchin, Wim J. Wolff, and Karsten Reise 29.1 29.2 29.3 29.4 Introduction................................................................................... Non-indigenous Aquatic Species in the North Sea Region .......... Cryptogenic Species ..................................................................... Nonindigenous Species Recorded in the North Sea as a Result of Natural Dispersal.................................................... 29.5 Case Histories ............................................................................... 29.5.1 Crepidula fornicata – the Slipper Limpet ...................... 29.5.2 Eriocheir sinensis – the Chinese Mitten Crab ................ 29.5.3 Teredo navalis – the Shipworm ...................................... 29.6 Conclusions................................................................................... References ................................................................................................ 30 31 507 507 508 516 518 518 519 521 522 524 525 European Enclosed and Semi-enclosed Seas ....................................... Erkki Leppäkoski, Tamara Shiganova, and Boris Alexandrov 529 30.1 30.2 Introduction................................................................................... The Black Sea ............................................................................... 30.2.1 Origin of Native Biota .................................................... 30.2.2 Nonindigenous Flora and Fauna..................................... 30.2.3 Case Histories ................................................................. 30.2.4 Vectors, Modes of Transmission .................................... 30.3 The Caspian Sea ........................................................................... 30.3.1 Origin of Native Biota .................................................... 30.3.2 Nonindigenous Flora and Fauna..................................... 30.3.3 Origin of Nonindigenous Species................................... 30.3.4 Vectors, Modes of Transmission .................................... 30.4 The Baltic Sea ............................................................................... 30.4.1 Origin of Native Biota .................................................... 30.4.2 Nonindigenous Flora and Fauna..................................... 30.4.3 Origin of the NIS ............................................................ 30.4.4 Vectors, Modes of Transmission .................................... 30.5 Conclusions................................................................................... References ................................................................................................ 529 532 533 533 534 536 536 537 537 539 540 540 541 541 542 543 544 544 Marine Bioinvasions in the Mediterranean Sea – History, Distribution and Ecology ....................................................................... Gil Rilov and Bella Galil 549 31.1 31.2 Introduction................................................................................... The Mediterranean Sea: A “Hotspot” of Marine Bioinvasions .................................................................................. 31.2.1 Spatial Patterns ............................................................... 549 551 552 Contents xix 31.2.2 31.2.3 Temporal Patterns ........................................................... Ecological Characteristics of Mediterranean Invaders (Fish) ................................................................ 31.3 Erythrean Invasion Through the Suez Canal ................................ 31.3.1 A History of Dramatic Changes ..................................... 31.3.2 The Warm Winter of 1954–55 ........................................ 31.3.3 The Eastern Mediterranean Climatic Transient.............. 31.3.4 Ecological Interactions of Erythrean Invaders in the Receiving Environment ........................................ 31.3.5 Impacts of the Erythrean Invasion on Human Activity ......................................................... 31.4 Lagoons as Invasion Hotspots ...................................................... 31.5 Infamous Invaders ......................................................................... 31.5.1 Macrophytes ................................................................... 31.5.2 Animals .......................................................................... 31.6 Conclusions................................................................................... References ................................................................................................ 32 A First Assessment of Invasive Marine Species on Chinese and Korean Coasts ................................................................................. Kyung S. Seo and Yoon Lee 32.1 32.2 Introduction................................................................................... Marine Invaders in Korea and China ............................................ 32.2.1 Pathogens in the Mariculture Industry ........................... 32.2.2 Mussels ........................................................................... 32.2.3 Sea Stars ......................................................................... 32.2.4 Tunicates......................................................................... 32.2.5 Bryozoans ....................................................................... 32.2.6 Phytoplankton................................................................. 32.3 Vectors of Invasions into Korean and Chinese Waters ................. 32.4 Conclusion .................................................................................... References ................................................................................................ 33 Invasions of Estuaries vs the Adjacent Open Coast: A Global Perspective .............................................................................. Rikke K. Preisler, Kerstin Wasson, Wim J. Wolff, and Megan C. Tyrrell 33.1 33.2 33.3 Habitat Differences in Marine Invasion Rates .............................. An Estuarine Emphasis to Marine Invasion Research .................. A Synthesis of Global Temperate Invertebrate Data on Invasions .......................................................................... 33.3.1 Estuarine vs Open Coast Invasions ................................ 33.3.2 Regional Differences in Estuarine vs Coastal Invasions ....................................................... 554 554 555 557 558 559 560 563 564 566 566 568 569 570 577 577 577 580 581 581 582 582 582 582 583 583 587 587 588 589 589 594 xx Contents 33.4 Hypotheses for Higher Invasion Rates of Estuaries vs Coasts ....................................................................................... 33.4.1 Greater Alien Propagule Pressure in Estuaries............... 33.4.2 Estuarine Species are Better Suited to Estuarine Conditions ...................................................................... 33.4.3 Establishment is Facilitated by the Limited Circulation in Estuaries .................................................. 33.4.4 Estuaries Have Undergone More Human Alterations ...................................................................... 33.4.5 Estuaries Have More “Empty Niches” ........................... 33.5 Directions for Future Research ..................................................... 33.6 Conclusions................................................................................... References ................................................................................................ Section VII 34 605 605 607 609 610 612 614 615 615 Concluding Thoughts Future Directions For Marine Invasions Research ............................. Jeffrey A. Crooks and Gil Rilov 621 34.1 Introduction................................................................................... 34.2 Scientific Considerations .............................................................. 34.3 Management Considerations ........................................................ 34.4 Social Considerations ................................................................... 34.5 Conclusions................................................................................... References ................................................................................................ 621 621 622 623 624 624 Subject Index .................................................................................................. 627 Taxonomic Index ............................................................................................ 631