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Biological Invasions in Marine Ecosystems: Ecological, Management, and Geographic Perspectives PDF

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Ecological Studies, Vol. 204 Analysis and Synthesis Edited by M.M. Caldwell, Washington, USA G. Heldmaier, Marburg, Germany R.B. Jackson, Durham, USA O.L. Lange, Würzburg, Germany H.A. Mooney, Stanford, USA E.-D. Schulze, Jena, Germany U. Sommer, Kiel, Germany Ecological Studies Further volumes can be found at springer.com Volume 189 Volume 198 Ecology of Harmful Algae (2006) Gradients in a Tropical Mountain E. Granéli and J.T. Turner (Eds.) Ecosystem of Ecuador (2008) E. Beck, J. Bendix, I. Kottke, Volume 190 F. Makeschin, R. Mosandl (Eds.) Wetlands and Natural Resource Management (2006) Volume 199 J.T.A. Verhoeven, B. Beltman, Hydrological and Biological R. Bobbink, and D.F. Whigham (Eds.) Responses to Forest Practices: The Alsea Watershed Study (2008) Volume 191 J.D. Stednick (Ed.) Wetlands: Functioning, Biodiversity Conservation, Volume 200 and Restoration (2006) Arid Dune Ecosystems: R. Bobbink, B. Beltman, J.T.A. Verhoeven, The Nizzana Sands in the and D.F. Whigham (Eds.) Negev Desert (2008) S.-W. Breckle, A. Yair, Volume 192 and M. Veste (Eds.) Geological Approaches to Coral Reef Ecology (2007) Volume 201 R.B. Aronson (Ed.) The Everglades Experiments: Lessons for Ecosystem Restoration (2008) Volume 193 C. Richardson (Ed.) Biological Invasions (2007) W. Nentwig (Ed.) Volume 202 Ecosystem Organization Volume 194 of a Complex Landscape: Clusia: A Woody Neotropical Long-Term Research Genus of Remarkable Plasticity in the Bornhöved and Diversity (2007) Lake District, Germany (2008) U. Lüttge (Ed.) O. Fränzle, L. Kappen, H.-P. Blume, Volume 195 and K. Dierssen (Eds.) The Ecology of Browsing and Volume 203 Grazing (2008) The Continental-Scale Greenhouse Gas I.J. Gordon and H.H.T. Prins (Eds.) Balance of Europe (2008) Volume 196 H. Dolman, R. Valentini, and A. Freibauer Western North American Juniperus (Eds.) Communites: A Dynamic Vegetation Volume 204 Type (2008) Biological Invasions in Marine O. Van Auken (Ed.) Ecosystems: Ecological, Management, Volume 197 and Geographic Perspectives (2009) Ecology of Baltic Coastal Waters (2008) G. Rilov and J.A. Crooks (Eds.) U. Schiewer (Ed.) Gil Rilov • Jeffrey A. Crooks Editors Biological Invasions in Marine Ecosystems Ecological, Management, and Geographic Perspectives Editors Dr. Gil Rilov Dr. Jeffrey A. Crooks Department of Zoology Tijuana River National Estuarine Research Cordley 3029 Reserve and the Southwest Wetlands Oregon State University Interpretive Association Corvallis, OR 97331-2914, 301 Caspian Way USA Imperial Beach CA 91932, USA ISBN: 978-3-540-79235-2 e-ISBN: 978-3-540-79236-9 Ecological Studies ISSN 0070-8356 Library of Congress Control Number: 2008929900 © 2009 Springer-Verlag Berlin Heidelberg This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, b roadcasting, reproduction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permissions for use must always be obtained from Springer-Verlag. Violations are liable for prosecution under the German Copyright Law. The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant p rotective laws and regulations and therefore free for general use. Cover illustration: Vectors that have transported marine species around the globe have been operating for millennia, but the diversity and intensity of vector activity has increased dramatically in recent years. The drawing on the book cover depicts the change in the types of one particularly important vector, oceangoing vessels, over this time frame. In the first millennium, large canoes were used by the Polynesian people to travel around the Pacific and occupy new lands, and the Vikings crossed the Atlantic in longships. These were replaced by large sailing ships that traversed the oceans during most of the second millennium, only to be replaced by huge freighters and tankers that dominate the seas today. With this change, the vectors changed in size and mode of transfer of marine invaders from fouling only, to fouling and dry ballast, to fouling and ballast waters. With the change in the size of vessels and the number of routes and voyages as well as the proliferation of other invasion vectors such as canals, fisheries and trade in pets, bait and seafood, the number of invasive species has dramatically increased. The graph represents this increase, from just a few species such as woodborrowing shipworms (Teredo spp) to a multitude of species that take advantage of the many ways in which marine species now move around the world (Design Gil Rilov). Cover design: WMXDesign GmbH, Heidelberg, Germany Printed on acid-free paper 9 8 7 6 5 4 3 2 1 springer.com Preface The timing of the publication of this book couldn’t be better as we celebrate the 50th anniversary of Charles Elton’s seminal book, The Ecology of Invasions by Animals and Plants. Since this influential book was published in 1958, the study of bioinvasions has developed exponentially, alongside the exponential growth in the magnitude of the invasion problem itself. Today, bioinvasion, a highly complex ecological process and environmental concern, has become a specific branch in ecology and environmental studies, with many disciplines developing within it. In early 2004 one of us (G.R.) received a letter from Springer Publishers asking to consider writing or editing a book on marine bioinvasions for their Ecological Studies Series. Springer’s editors have been thinking of such a book for quite a while — and rightfully so. Over the past several decades, many books have been written on management, evolutionary or ecological perspectives of bioinvasions, but those books have focused mostly on terrestrial invasions with only a few marine examples. Research on marine bioinvasions has been mounting in the last two dec- ades; a special biannual international symposium on the topic was established in 1999, signifying the fact that it has become a discipline in its own right, but no comprehensive marine bioinvasions book existed at that time. Today, four years later, this is still the first of its kind. The challenge was exciting and, because the book’s purpose was to reflect the full breadth of this fascinating topic, it warranted the inclusion of knowledge of many expert contributors. Dr. Jeff Crooks joined in as a co-editor, and we launched on this long journey together. It was a perfect match: both of us have worked on invasive mussels and their interactions with the native communities, one (G.R.) in one of the hottest hotspots of marine bioinvasions on earth, the eastern Mediterranean, and the other (J.C.) in one of the hotspots on the West Coast of North America, Southern California. Together, we envisioned a book that will focus mainly on the ecological aspects of the invasion process in the marine environment, which in many ways is quite different to the terrestrial one. With the help of Prof. Jim Carlton, we crafted an outline for a book that would depict the different stages of the invasion process, discuss management issues, and present illustrative case studies from different regions of the world. Identifying prospective contributors was the easy part of the task: many excellent experts could address these different processes. The big v vi Preface question was would these very prolific, busy, and active researchers be willing and able to contribute to this effort? The response was heartening: when we presented the idea to our prospective contributors, the excitement and willingness to commit were overwhelming. Within ten days, we had a list of chapter authors that would cover most of the topics we envisioned. This rapid and positive response reempha- sized the great thirst for such a book. One of the challenges in producing this book is that the topic is so “hot” and rapidly evolving that it is hard to keep up with the new science. This is for two main reasons. The first is that new invasions are being discovered weekly in many regions, and the second is that new basic research is continuously being produced on new, as well as older, invasions. With every iteration of the book’s production, new material seemed appropriate for inclusion and we were bound to draw the line somewhere. Yet this is what made this project so exciting. It became clear that it is impossible to include everything we might have wanted in this one book. Luckily, there are other excellent books on the general theory of bioinvasions that cover many of the topics of which we could only scratch the surface. Many are mentioned in the introductory chapter. The dedicated contributing authors of this book made great efforts to keep the content as updated and relevant as possible. This enables our book, we believe, to serve as an up-to-date learning tool for students and a valuable asset for profession- als in the field. Editing such an encompassing book wasn’t always an easy assign- ment, and quite challenging at times; still, we have thoroughly enjoyed this creative work. We are grateful to Springer Publishers for their support during this process, and mostly we thank our devoted chapter authors without which this book would not exist. We also thank our patient families, Iris, Shai and Shir Rilov, and Emma, Kai, and Aiden Crooks, for allowing us many hours of seclusion working on the book. We hope that Biological Invasions in Marine Ecosystems: Ecological, Management, and Geographic Perspectives will serve its purpose and broaden the knowledge and understanding of this important topic. June 2008 Gil Rilov, USA Jeff Crooks, USA Contents Section I Perspectives on Marine Invasions 1 Marine Bioinvasions: Conservation Hazards and Vehicles for Ecological Understanding ................................................................. 3 Gil Rilov and Jeffrey A. Crooks 1.1 Introduction – The Problem of Biological Invasions ........................ 3 1.2 Invasions as a Tool to Study Nature .................................................. 5 1.3 Invasion Biology – The Discipline and its Application in the Marine Environment ............................................................... 6 1.4 Structure of the Book ........................................................................ 7 1.5 Note on Terminology ........................................................................ 8 1.6 Concluding Thoughts ........................................................................ 9 References .................................................................................................. 9 2 Deep Invasion Ecology and the Assembly of Communities in Historical Time..................................................................................... 13 James T. Carlton 2.1 Introduction ....................................................................................... 13 2.2 Invader Underestimation – Systematics ............................................ 14 2.2.1 Cryptogenic Species . 14 2.2.2 Pseudoindigenous Species . 16 2.3 Invader Underestimation – Biogeographic and Community History .................................................................... 39 2.3.1 Widespread Intraoceanic and Interoceanic Corridor Species ................................................................... 39 2.3.2 Neritic Species with Presumptive Oceanic Dispersal ........... 40 2.3.3 Resident Species . 41 2.4 Invader Underestimation – Sampling ................................................ 42 2.4.1 Species in Underexplored Habitats and Associations........... 42 2.4.2 Incipient Invasions: Species with Small Population Sizes .................................................................... 42 2.5 The Overestimation of Invader Diversity .......................................... 43 vii viii Contents 2.6 Discussion ......................................................................................... 44 2.7 The Way Forward: Solutions ............................................................ 47 References .................................................................................................. 48 3 Natural and Climate Change Mediated Invasions ................................ 57 Steve I. Lonhart 3.1 Introduction ....................................................................................... 57 3.2 The Geographic Range of a Species ................................................. 57 3.3 Range Shifts ...................................................................................... 58 3.3.1 Factors that Influence our Understanding of the Geographic Range and Range Limits ......................... 58 3.3.2 Natural Range Shifts . 60 3.3.3 Human-mediated Range Shifts ............................................. 62 3.4 Climate Change and Range Shifts .................................................... 63 3.4.1 Observed Biological Responses to Climate Change............. 63 3.5 Contrasting Natural Range Expansions and Biological Invasions ................................................................... 66 3.6 Conclusions ....................................................................................... 67 References .................................................................................................. 67 4 Modeling Marine Invasions: Current and Future Approaches ........... 71 Marjorie J. Wonham and Mark A. Lewis 4.1 Introduction ....................................................................................... 71 4.1.1 Why Marine Invasion Modeling? ......................................... 71 4.1.2 Scope of this Review ............................................................ 72 4.2 Invasion Pathway Models ................................................................. 73 4.3 Population Models: Invasion Dynamics ........................................... 74 4.3.1 Single-species Models . 79 4.3.2 Multi-species Models . 80 4.4 Population Models: Invasion Spread ................................................ 84 4.4.1 Single-species Models . 85 4.4.2 Allee Effects . 87 4.4.3 Multi-species Models . 90 4.5 Community Invasibility Models ....................................................... 93 4.6 Summary and Future Directions ....................................................... 94 4.6.1 Formalizing Conceptual Models Mathematically................. 94 4.6.2 Coupling Dynamical and Statistical Models ........................ 96 4.6.3 Integrating Modeling and Empirical Work ........................... 97 References .................................................................................................. 98 Appendix .................................................................................................... 105 Contents ix Section II Invader Arrival 5 Characterizing Vectors of Marine Invasion ........................................... 109 Dan Minchin, Stephan Gollasch, Andrew N. Cohen, Chad L. Hewitt, and Sergej Olenin 5.1 Introduction ....................................................................................... 109 5.2 Primary vs Secondary Introductions ................................................. 109 5.3 Principal Vectors of Marine Invasion ................................................ 110 5.3.1 Shipping ................................................................................ 110 5.3.2 Canals ................................................................................... 110 5.3.3 Aquaculture ........................................................................... 111 5.3.4 Fisheries ................................................................................ 111 5.3.5 Ornamental Species and Live Seafood ................................. 112 5.3.6 Marine Leisure and Tourism ................................................. 112 5.3.7 Research and Education ........................................................ 112 5.3.8 Habitat Restoration and Management .................................. 113 5.4 Vector Management .......................................................................... 113 References .................................................................................................. 115 6 The Vessel as a Vector – Biofouling, Ballast Water and Sediments........................................................................................... 117 Chad L. Hewitt, Stephan Gollasch, and Dan Minchin 6.1 Introduction ....................................................................................... 117 6.2 Biofouling ......................................................................................... 120 6.3 Ballast Water and Sediments............................................................. 124 6.4 Discussion ......................................................................................... 126 References .................................................................................................. 129 7 The Role of Propagule Pressure in Invasion Success ............................ 133 Emma L. Johnston, Richard F. Piola, and Graeme F. Clark 7.1 Introduction ....................................................................................... 133 7.2 Propagule Pressure ............................................................................ 133 7.3 The Resilience of Propagules ............................................................ 135 7.4 Producing Propagules ....................................................................... 138 7.5 Inferring Propagule Supply from Current Distributions ................... 139 7.6 Vectors and Propagule Supply .......................................................... 141 7.7 Manipulating Propagule Supply ....................................................... 142 7.7.1 Manipulating Propagule at the Larval Stage ......................... 143 7.7.2 Manipulating Adult Invaders ................................................ 145 7.8 Conclusions ....................................................................................... 146 References .................................................................................................. 147 x Contents 8 Differentiating Successful and Failed Invaders: Species Pools and the Importance of Defi ning Vector, Source and Recipient Regions ............................................................... 153 A. Whitman Miller and Gregory M. Ruiz 8.1 Introduction ................................................................................... 153 8.2 Identifying the Roles of Source Region, Recipient Region and Vector ......................................................................... 154 8.3 Modeling Invasion State Space – Defi ning and Comparing Species Pools ...................................................... 156 8.3.1 Species Pool Designations ............................................. 157 8.3.2 Species Pool Designations and Comparisons (Fixed Recipient, Source, and Vector) ............................ 159 8.3.3 Fixed Recipient and Source Regions, Multiple Vectors ............................................................. 160 8.3.4 Fixed Recipient Region, Multiple Source Regions, Multiple Vectors .............................................. 162 8.3.5 Multiple Recipient Regions, Fixed Source Region, Multiple Vectors ................................................ 163 8.4 Some Recent Analyses of Invader Attributes Using Species Pool Comparisons ......................................................................... 163 8.5 Discussion ..................................................................................... 167 References ................................................................................................ 169 Section III Invader Establishment 9 The Establishment of Invasive Species ................................................. 173 Jeffrey A. Crooks and Gil Rilov 9.1 Introduction ................................................................................... 173 9.2 Factors Infl uencing Numeric and Geographic Growth of Invasive Populations ................................................................. 173 References ................................................................................................ 174 10 The Role of Phenotypic Plasticity in Marine Biological Invasions.................................................................................................. 177 L. David Smith 10.1 Introduction ................................................................................... 177 10.2 Phenotypic Plasticity .................................................................... 178 10.2.1 A Brief Overview ........................................................... 178 10.2.2 General Importance to Invasions .................................... 180 10.2.3 Empirical Work in Non-marine Systems ........................ 182 10.3 Phenotypic Plasticity in Marine Systems ..................................... 183 10.4 Where Phenotypic Plasticity Might Play a Role in Marine Invasions ........................................................................... 187

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