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EncyclopEdia of
Biological invasions
EncyclopEdia of
Biological invasions
EditEd by
daniEl simBErloff
University of Tennessee, Knoxville
marcEl rEjmánEk
University of California, Davis
UnivErsity of california prEss
Berkeley Los Angeles London
University of California Press, one of the most distinguished university
presses in the United States, enriches lives around the world by advancing
scholarship in the humanities, social sciences, and natural sciences. Its
activities are supported by the UC Press Foundation and by philanthropic
contributions from individuals and institutions. For more information, visit
www.ucpress.edu.
Encyclopedias of the Natural World, No. 3
University of California Press
Berkeley and Los Angeles, California
University of California Press, Ltd.
London, England
© 2011 by the Regents of the University of California
Library of Congress Cataloging-in-Publication Data
Encyclopedia of biological invasions / edited by Daniel Simberloff and
Marcel Rejmánek.
p. cm. — (Encyclopedias of the natural world ; 3)
Includes bibliographical references and index.

ISBN 978-0-520-26421-2 (cloth)
1. Introduced organisms—Encyclopedias. I. Simberloff, Daniel. II.
Rejmánek, Marcel.
QH353.E53
2011
578.6’2 dc22—dc21
2010010391
Manufactured in China.
19 18 17 16 15 14 13 12 11
10 9 8 7 6 5 4 3 2 1
The paper used in this publication meets the minimum requirements of
ANSI/NISO Z39.48-1992 (R 1997) (Permanence of Paper).
Cover photograph: Large, pink masses of eggs laid by golden apple snails,
courtesy N.O.L. Carlsson. Insets, from left: Pheidole megacephala tending
aphids, courtesy Alex Wild; European starling, courtesy Michelle St. Sauveur,
Coventry, Rhode Island; oral disc of a parasitic-phase sea lamprey, courtesy
Ted Lawrence, Great Lakes Fishery Commission; flowering tropical weed
Turnera subulata, courtesy Spencer C.H. Barrett.
Title page photo: An observer dwarfed by an overgrowth of giant hogweed
(Heracleum mantegazzianum) invading the Slavkovský les Protected
Landscape Area, Czech Republic, Central Europe. Introduced to Europe
from the Caucasus, invasive giant hogweed can grow to over four meters in
height. Its phototoxic sap blisters human skin and can cause serious scarring.
Photograph by Petr Pyšek. From Ecology and Management of Giant Hogweed
(Heracleum mantegazzianum), edited by P. Pyšek, M. J. W. Cock, H. P. Ravn,
and W. Nentwig (2007, CAB International, Wallingford, UK).
v
Contents
Contents by Subject Area / ix
Contributors / xiii

Guide to the Encyclopedia / xxi
Preface / xxiii
AcclimatizationSocieties/1
Christopher Lever
Agreements,International/4
Jamie K. Reaser
Agriculture/7
Adam S. Davis; Douglas A. Landis
Algae/11
Jennifer E. Smith
Allelopathy/16
Inderjit
Ants/17
Nathan J. Sanders
Apomixis/24
Takashi Okada; Ross A. Bicknell;
Anna M. Koltunow
Aquaculture/27
Devin M. Bartley
Aquaria/32
Ian C. Duggan
Australia:Invasions/36
Tim Low
Ballast/43
James T. Carlton
Bees/50
David Goulson
Competition,Animal/117
Christopher R. Dickman
Competition,Plant/122

Jessica Gurevitch
Crabs/125
Edwin Grosholz
Craysh/129
Francesca Gherardi
Crustaceans(Other)/135
Anthony Ricciardi
DAISIEProject/138
Petr Pyšek; Philip E. Hulme;
Wolfgang Nentwig; Montserrat Vilà
Darwin,Charles/142
Marc W. Cadotte
Databases/145
Laura A. Meyerson
Demography/147
Carol C. Horvitz
DiseaseVectors,Human/150
L. Philip Lounibos
DispersalAbility,Animal/154
R. M. McDowall
DispersalAbility,Plant/159
Eugene W. Schupp
Disturbance/165
Richard J. Hobbs
EarlyDetectionandRapid
Response/169
Randy G. Westbrooks; Robert E. Eplee
BelowgroundPhenomena/54
David A. Wardle
BiologicalControl,ofAnimals/58

J. Howard Frank; Roy G. Van Driesche;
Mark S. Hoddle; E. D. McCoy
BiologicalControl,ofPlants/63
Michael J. Pitcairn
Birds/70
Navjot S. Sodhi
Black,White,andGrayLists/75
Stanley W. Burgiel; Anne M. Perrault
BrownTreesnake/78
Gad Perry; Gordon H. Rodda
BryophytesandLichens/81
Franz Essl; Phil Lambdon;
Wolfgang Rabitsch
BurmesePythonandOtherGiant
Constrictors/85
Robert N. Reed; Gordon H. Rodda
Canals/92
Stephan Gollasch
Carnivores/95
Arijana Barun; Daniel Simberloff
Carp,Common/100
Peter W. Sorensen; Przemyslaw Bajer
CARTandRelatedMethods/104
Vojteˇch Jarošík
Cheatgrass/108
Richard N. Mack
ClimateChange/113
Jeffrey S. Dukes
 vi 


Marcel Rejmánek

James T. Carlton

William G. Lee

Daniel Simberloff

Edward W. Evans; William E. Snyder

Jeffrey A. Crooks

Hugh J. MacIsaac


Thomas J. Stohlgren

Richard J. Hobbs
Lantanacamara
Stephen Johnson

Marc L. Miller

Jennifer L. Bufford; Curtis C. Daehler

Louis Lambrechts; Anna Cohuet;
Vincent Robert

Christopher B. Anderson;
Alejandro E. J. Valenzuela


Andrea J. Pickart

Bella S. Galil

Jean-Yves Meyer; Arthur C. Medeiros

L. Philip Lounibos

John J. Stachowicz

Anne Pringle; Benjamin Wolfe;
Else Vellinga


Stephen J. Novak

Carla D’Antonio; Karen Stahlheber;
Nicole Molinari

David M. Forsyth

Edward L. Mills; Kristen T. Holeck

Patrick C. Tobin; Andrew M. Liebhold

Scott Steinmaus

Lloyd Loope


David A. Orwig

Joseph M. DiTomaso

Mara A. Evans; Donald R. Strong, Jr.

Sarah Reichard


Kristina A. Schierenbeck

David Le Maitre

C. W. Potter

Robert F. Norris


Jason D. Fridley

Betsy Von Holle

Mark A. Davis


Matthew K. Chew

Charles Perrings

Samuel W. James


Emiel Elferink; Wouter van der Weijden

Daniel Simberloff


Richard B. Primack

Bernd Blossey

Alan Hastings

Piero Genovesi

Marcel Rejmánek; David M. Richardson

Katharina A. M. Engelhardt


Sharon Y. Strauss


Carol Eunmi Lee

Robert C. Klinger

Peter B. Moyle; Emili García-Berthou

John S. Mackenzie; David T. Williams


Andrew M. Liebhold;
Deborah G. McCullough

David M. Richardson


Lars W. J. Anderson

Marie-Laure Desprez-Loustau;
David M. Rizzo

Leonard A. Brennan; Fred C. Bryant

Kristin Saltonstall
  vii

Robert H. Cowie

David M. Richardson; John R. Wilson;
Olaf Weyl; Charles L. Grifths

Scott J. Meiners; Steward T. A. Pickett

Julie L. Lockwood; Thomas Virzi

Alberto Jiménez-Valverde; Jorge M. Lobo

James H. Richards; Benjamin R. Janes

Joan G. Ehrenfeld


David M. Richardson

Jitka Klimes˘ová

Carla J. Harris; Rachael Gallagher

Jacqueline Beggs

Martin Hill; Julie Coetzee; Mic Julien;
Ted Center

Jodie S. Holt

Joy B. Zedler

Peter Coates

Ladd Erik Johnson

Mick N. Clout; James C. Russell

Richard P. Duncan

John Randall

Jonathan M. Jeschke

Michel Pascal


Phyllis N. Windle

Gregory P. Asner; Cho-ying Huang

Spencer C. H. Barrett

Fred Kraus

Truman P. Young; Kurt J. Vaughn

Andy Dobson; Ricardo M. Holdo;
Robert D. Holt


W. M. Lonsdale

Emili García-Berthou; Peter B. Moyle

Daniel Simberloff

Daniel Simberloff

Peter W. Sorensen; Roger A. Bergstedt

Edwin Grosholz

Michelle R. Leishman; Carla J. Harris

Warren Hays


Daniel Simberloff

David R. Towns; Peter de Lange;
Mick N. Clout

Robert M. Pringle

Jae R. Pasari; Paul C. Selmants; Hillary
Young; Jennifer O’Leary; Erika S. Zavaleta

Ragan M. Callaway

Jennifer Ruesink

Bernd Sures

Henning S. Heide-Jørgensen

Graham J. Hickling

Pieter Bol

Megan A. Rúa; Charles E. Mitchell

Ram Pratap Yadav; Ajay Singh


Daniel A. Herms; Deborah G. McCullough

Desley A. Whisson


Peter T. Jenkins
Phytophthora
David M. Rizzo

Diego P. Vázquez; Carolina L. Morales

Tamara Shiganova
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ix
INVADERATTRIBUTES
Allelopathy
Apomixis
Demography
Dispersal Ability, Animal
Dispersal Ability, Plant
Genotypes, Invasive
Geographic Origins and Introduction Dynamics
Invasiveness
Life History Strategies
Mutualism
“Native Invaders”
Novel Weapons Hypothesis
Reproductive Systems, Plant
Seed Ecology
Taxonomic Patterns
Tolerance Limits, Animal
Tolerance Limits, Plant
Vegetative Propagation
ECOSYSTEMFEATURES

Belowground Phenomena
Disturbance
Eutrophication, Aquatic
Habitat Compatibility
Invasibility, of Communities and Ecosystems
Islands
Lakes
Nitrogen Enrichment
Protected Areas
Rivers
Seas and Oceans
Succession
Wetlands
PROCESSES
Climate Change
Enemy Release Hypothesis
Epidemiology and Dispersal
Evolutionary Response, of Natives to Invaders
Evolution of Invasive Populations
Fire Regimes
Hybridization and Introgression
Invasional Meltdown
Lag Times
Land Use
Propagule Pressure
Range Modeling
Succession
IMPACTS
Allelopathy
Competition, Animal

Competition, Plant
Contents by subjeCt AreA
 x ContentsbysubjeCtAreA
Invertebrates, Marine
Kudzu
Ladybugs
Lantana camara
Malaria Vectors
Mammals, Aquatic
Melastomes
Mosquitoes
Mycorrhizae
Nile Perch
Parasites, of Animals
Parasitic Plants
Pathogens, Animal
Pathogens, Human
Pathogens, Plant
Phytophthora
Predators
Rats
Reptiles and Amphibians
Rinderpest
Rodents (Other)
Sea Lamprey
Small Indian Mongoose
Snails and Slugs
Trees and Shrubs
Vines and Lianas
Wasps

Water Hyacinth
Zebra Mussel
PAtHWAystoInVAsIon
Acclimatization Societies
Agriculture
Aquaculture
Aquaria
Ballast
Canals
Forestry and Agroforestry
Game Animals
Horticulture
Endangered and Threatened Species
Fire Regimes
Herbivory
Hybridization and Introgression
Hydrology
Invasion Economics
Landscape Patterns of Plant Invasions
“Native Invaders”
Novel Weapons Hypothesis
Pollination
Transformers
Weeds
notAbLetAXA
Algae
Ants
Bees
Birds
Brown Treesnake

Bryophytes and Lichens
Burmese Python and Other Giant Constrictors
Carnivores
Carp, Common
Cheatgrass
Crabs
Crayfish
Crustaceans (Other)
Disease Vectors, Human
Earthworms
Eucalypts
Fishes
Flaviviruses
Forest Insects
Freshwater Plants and Seaweeds
Fungi
Grasses and Forbs
Grazers
Gypsy Moth
Hemlock Woolly Adelgid
Influenza
 ContentsbysubjeCtAreA xi
Regulation (U.S.)
Remote Sensing
Restoration
Risk Assessment and Prioritization
HIstory
DAISIE Project
Darwin, Charles
Elton, Charles S.

Invasion Biology
Invasion Biology: Historical Precedents
SCOPE Project
Xenophobia
notAbLeInVAsIons
Australia: Invasions
Great Lakes: Invasions
Hawaiian Islands: Invasions
Mediterranean Sea: Invasions
New Zealand: Invasions
Ponto-Caspian: Invasions
South Africa: Invasions
Ostriculture
Pet Trade
MAnAGeMentAnDreGuLAtIon
Agreements, International
Biological Control, of Animals
Biological Control, of Plants
Black, White, and Gray Lists
CART and Related Methods
Databases
Early Detection and Rapid Response
Ecoterrorism and Biosecurity
Eradication
Herbicides
Integrated Pest Management
Laws, Federal and State
Mechanical Control
Pesticides (Fish and Mollusc)
Pesticides for Insect Eradication

Pesticides (Mammal)
Range Modeling
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xiii
CHRISTOPHER B. ANDERSON
University of Magallanes
Punta Arenas, Chile
Mammals, Aquatic
LARS W. J. ANDERSON
USDA–Agricultural Research Service
Davis, California
Freshwater Plants and Seaweeds
GREGORY P. ASNER
Carnegie Institution for Science
Stanford, California
Remote Sensing
PRZEMYSLAW BAJER
University of Minnesota, St. Paul
Carp, Common
SPENCER C. H. BARRETT
University of Toronto, Ontario, Canada
Reproductive Systems, Plant
DEVIN M. BARTLEY
California Department of Fish and Game, Sacramento
Aquaculture
ARIJANA BARUN
University of Tennessee, Knoxville
Carnivores
JACQUELINE BEGGS
University of Auckland, New Zealand

Wasps
ROGER A. BERGSTEDT
U.S. Geological Survey
Millersburg, Michigan
Sea Lamprey
ROSS A. BICKNELL
Plant and Food Research
Christchurch, New Zealand
Apomixis
BERND BLOSSEY
Cornell University
Ithaca, New York
Enemy Release Hypothesis
PIETER BOL
Delft University of Technology
Delft, The Netherlands
Pathogens, Human
LEONARD A. BRENNAN
Texas A&M University, Kingsville
Game Animals
FRED C. BRYANT
Texas A&M University, Kingsville
Game Animals
JENNIFER L. BUFFORD
University of Hawaii, Honolulu
Life History Strategies
STANLEY W. BURGIEL
Global Invasive Species Programme
Alexandria, Virginia
Black, White, and Gray Lists

MARC W. CADOTTE
University of Toronto, Ontario, Canada
Darwin, Charles
RAGAN M. CALLAWAY
University of Montana, Missoula
Novel Weapons Hypothesis
CONTRIBUTORS
 xiv Contributors
PETER DE LANGE
New Zealand Department of Conservation, Auckland
New Zealand: Invasions
MARIE-LAURE DESPREZ-LOUSTAU
Institut National de la Recherche Agronomique (INRA)
Villenave d’Ornon, France
Fungi
CHRISTOPHER R. DICKMAN
University of Sydney, New South Wales, Australia
Competition, Animal
JOSEPH M. DITOMASO
University of California, Davis
Herbicides
ANDY DOBSON
Princeton University
Princeton, New Jersey
Rinderpest
IAN C. DUGGAN
University of Waikato
Hamilton, New Zealand
Aquaria
JEFFREY S. DUKES

Purdue University
West Lafayette, Indiana
Climate Change
RICHARD P. DUNCAN
Lincoln University, New Zealand
Propagule Pressure
JOAN G. EHRENFELD
Rutgers University
New Brunswick, New Jersey
Transformers
EMIEL ELFERINK
Centre for Agriculture and Environment
Culemborg, The Netherlands
Ecoterrorism and Biosecurity
KATHARINA A. M. ENGELHARDT
University of Maryland Center for Environmental
Science, Frostburg
Eutrophication, Aquatic
ROBERT E. EPLEE
USDA APHIS PPQ
Whiteville, North Carolina
Early Detection and Rapid Response
FRANZ ESSL
Environment Agency Austria
Vienna, Austria
Bryophytes and Lichens
JAMES T. CARLTON
Williams College
Mystic, Connecticut
Ballast

Invertebrates, Marine
TED CENTER
U.S. Department of Agriculture
Ft. Lauderdale, Florida
Water Hyacinth
MATTHEW K. CHEW
Arizona State University, Tempe
Invasion Biology: Historical Precedents
MICK N. CLOUT
University of Auckland, New Zealand
New Zealand: Invasions
Predators
PETER COATES
University of Bristol, United Kingdom
Xenophobia
JULIE COETZEE
Rhodes University
Grahamstown, South Africa
Water Hyacinth
ANNA COHUET
Institut de Recherche pour le Développement
Montpellier, France
Malaria Vectors
ROBERT H. COWIE
University of Hawaii, Honolulu
Snails and Slugs
JEFFREY A. CROOKS
Tijuana River National Estuarine Research Reserve
Imperial Beach, California
Lag Times

CURTIS C. DAEHLER
University of Hawaii, Honolulu
Life History Strategies
CARLA D’ANTONIO
University of California, Santa Barbara
Grasses and Forbs
ADAM S. DAVIS
USDA-ARS Invasive Weed Management Unit
Urbana, Illinois
Agriculture
MARK A. DAVIS
Macalester College
St. Paul, Minnesota
Invasion Biology
 Contributors xv
EDWIN GROSHOLZ
University of California, Davis
Crabs
Seas and Oceans
JESSICA GUREVITCH
Stony Brook University, New York
Competition, Plant
CARLA J. HARRIS
Macquarie University
Sydney, New South Wales, Australia
Seed Ecology
Vines and Lianas
ALAN HASTINGS
University of California, Davis
Epidemiology and Dispersal

WARREN HAYS
Hawaii Pacific University, Honolulu
Small Indian Mongoose
HENNING S. HEIDE-JØRGENSEN
University of Copenhagen, Denmark
Parasitic Plants
DANIEL A. HERMS
Ohio State University, Columbus
Pesticides for Insect Eradication
GRAHAM J. HICKLING
University of Tennessee, Knoxville
Pathogens, Animal
MARTIN HILL
Rhodes University
Grahamstown, South Africa
Water Hyacinth
RICHARD J. HOBBS
University of Western Australia, Crawley
Disturbance
Land Use
MARK S. HODDLE
University of California, Riverside
Biological Control, of Animals
RICARDO M. HOLDO
University of Florida, Gainesville
Rinderpest
KRISTEN T. HOLECK
Cornell University, Ithaca, New York
Great Lakes: Invasions
JODIE S. HOLT

University of California, Riverside
Weeds
EDWARD W. EVANS
Utah State University, Logan
Ladybugs
MARA A. EVANS
University of California, Davis
Herbivory
DAVID M. FORSYTH
Department of Sustainability and Environment
Heidelberg, Victoria, Australia
Grazers
J. HOWARD FRANK
University of Florida, Gainesville
Biological Control, of Animals
JASON D. FRIDLEY
Syracuse University, New York
Invasibility, of Communities and Ecosystems
BELLA S. GALIL
National Institute of Oceanography
Haifa, Israel
Mediterranean Sea: Invasions
RACHAEL GALLAGHER
Macquarie University
Sydney, New South Wales, Australia
Vines and Lianas
EMILI GARCÍA-BERTHOU
University of Girona, Spain
Fishes
Rivers

PIERO GENOVESI
Institute for Environmental Protection and Research
(ISPRA)
Ozzano Emilia, Italy
Eradication
FRANCESCA GHERARDI
University of Florence,
Florence, Italy
Crayfish
STEPHAN GOLLASCH
GoConsult
Hamburg, Germany
Canals
DAVID GOULSON
University of Stirling, Scotland, United Kingdom
Bees
CHARLES L. GRIFFITHS
University of Cape Town, South Africa
South Africa: Invasions
 xvi Contributors
MIC JULIEN
Commonwealth Scientific and Industrial Research
Organisation (CSIRO)
Indooroopillay, Queensland, Australia
Water Hyacinth
JITKA KLIMES
˘
OVÁ
Czech Academy of Sciences, Trˇebonˇ
Vegetative Propagation

ROBERT C. KLINGER
U.S. Geological Survey–BRD
Bishop, California
Fire Regimes
ANNA M. KOLTUNOW
CSIRO Plant Industry
Adelaide, South Australia, Australia
Apomixis
FRED KRAUS
Bishop Museum
Honolulu, Hawaii
Reptiles and Amphibians
PHIL LAMBDON
Royal Botanic Gardens
Kew, United Kingdom
Bryophytes and Lichens
LOUIS LAMBRECHTS
Institut Pasteur
Paris, France
Malaria Vectors
DOUGLAS A. LANDIS
Michigan State University, East Lansing
Agriculture
CAROL EUNMI LEE
University of Wisconsin, Madison
Evolution of Invasive Populations
WILLIAM G. LEE
Landcare Research
Dunedin, New Zealand
Islands

MICHELLE R. LEISHMAN
Macquarie University
Sydney, New South Wales, Australia
Seed Ecology
DAVID LE MAITRE
Council for Scientific and Industrial Research (CSIR)
Stellenbosch, South Africa
Hydrology
ROBERT D. HOLT
University of Florida, Gainesville
Rinderpest
CAROL C. HORVITZ
University of Miami
Coral Gables, Florida
Demography
CHO-YING HUANG
National Taiwan University
Taipei, Taiwan
Remote Sensing
PHILIP E. HULME
Lincoln University, New Zealand
DAISIE Project
INDERJIT
University of Delhi, India
Allelopathy
SAMUEL W. JAMES
University of Kansas, Lawrence
Earthworms
BENJAMIN R. JANES
University of California, Davis

Tolerance Limits, Plant
VOJTE
ˇ
CH JAROŠÍK
Charles University
Prague, Czech Republic
CART and Related Methods
PETER T. JENKINS
Defenders of Wildlife
Washington, DC
Pet Trade
JONATHAN M. JESCHKE
Ludwig-Maximilians-University Munich
Planegg-Martinsreid, Germany
Range Modeling
ALBERTO JIMÉNEZ-VALVERDE
Universidad de Málaga, Spain
Tolerance Limits, Animal
LADD ERIK JOHNSON
Université Laval
Québec, Canada
Zebra Mussel
STEPHEN JOHNSON
Industry and Investment New South Wales
Orange, Australia
Lantana camara
 Contributors xvii
DEBORAH G. MCCULLOUGH
Michigan State University, East Lansing
Forest Insects

Pesticides for Insect Eradication
R. M. MCDOWALL
National Institute of Water and Atmospheric
Research
Christchurch, New Zealand
Dispersal Ability, Animal
ARTHUR C. MEDEIROS
U.S. Geological Survey–BRD
Maui, Hawaii
Melastomes
SCOTT J. MEINERS
Eastern Illinois University, Charleston
Succession
JEAN-YVES MEYER
Government of French Polynesia, Papeete
Melastomes
LAURA A. MEYERSON
University of Rhode Island, Kingston
Databases
MARC L. MILLER
University of Arizona, Tucson
Laws, Federal and State
EDWARD L. MILLS
Cornell University
Bridgeport, New York
Great Lakes: Invasions
CHARLES E. MITCHELL
University of North Carolina, Chapel Hill
Pathogens, Plant
NICOLE MOLINARI

University of California, Santa Barbara
Grasses and Forbs
CAROLINA L. MORALES
Instituto de Investigaciones en Biodiversidad y
Medioambiente, CONICET
Comahue, Argentina
Pollination
PETER B. MOYLE
University of California, Davis
Fishes
Rivers
CHRISTOPHER LEVER
University of Cambridge, United Kingdom
Acclimatization Societies
ANDREW M. LIEBHOLD
USDA Forest Service
Morgantown, West Virginia
Forest Insects
Gypsy Moth
JORGE M. LOBO
National Museum of Natural Sciences
Madrid, Spain
Tolerance Limits, Animal
JULIE L. LOCKWOOD
Rutgers University
New Brunswick, New Jersey
Taxonomic Patterns
W. M. LONSDALE
CSIRO Entomology
Canberra, Australian Capital Territory, Australia

Risk Assessment and Prioritization
LLOYD LOOPE
U.S. Geological Survey
Makawao, Hawaii
Hawaiian Islands: Invasions
L. PHILIP LOUNIBOS
University of Florida, Vero Beach
Disease Vectors, Human
Mosquitoes
TIM LOW
Invasive Species Council
Chapel Hill, Queensland, Australia
Australia: Invasions
HUGH J. MACISAAC
University of Windsor, Ontario, Canada
Lakes
RICHARD N. MACK
Washington State University, Pullman
Cheatgrass
JOHN S. MACKENZIE
Curtin University of Technology
Perth, Western Australia, Australia
Flaviviruses
EARL D. MCCOY
University of South Florida
Biological Control, of Animals
 xviii Contributors
C. W. POTTER
University of Sheffield, United Kingdom
Influenza

RICHARD B. PRIMACK
Boston University, Massachusetts
Endangered and Threatened Species
ANNE PRINGLE
Harvard University
Cambridge, Massachusetts
Mycorrhizae
ROBERT M. PRINGLE
Harvard University
Cambridge, Massachusetts
Nile Perch
PETR PYŠEK
Academy of Sciences
Pruhonice, Czech Republic
DAISIE Project
WOLFGANG RABITSCH
Environment Agency Austria
Vienna, Austria
Bryophytes and Lichens
JOHN RANDALL
The Nature Conservancy
Davis, California
Protected Areas
JAMIE K. REASER
Ecos Systems Institute
Stanardsville, Virginia
Agreements, International
ROBERT N. REED
USGS Fort Collins Science Center
Fort Collins, Colorado

Burmese Python and Other Giant Constrictors
SARAH REICHARD
University of Washington, Seattle
Horticulture
MARCEL REJMÁNEK
University of California, Davis
Eucalypts
Invasiveness
ANTHONY RICCIARDI
McGill University
Montreal, Quebec, Canada
Crustaceans (Other)
JAMES H. RICHARDS
University of California, Davis
Tolerance Limits, Plant
WOLFGANG NENTWIG
University of Bern, Switzerland
DAISIE Project
ROBERT F. NORRIS
University of California, Davis
Integrated Pest Management
STEPHEN J. NOVAK
Boise State University, Idaho
Geographic Origins and Introduction Dynamics
TAKASHI OKADA
CSIRO Plant Industry
Adelaide, South Australia, Australia
Apomixis
JENNIFER O’LEARY
University of California, Santa Cruz

Nitrogen Enrichment
DAVID A. ORWIG
Harvard Forest, Harvard University
Petersham, Massachusetts
Hemlock Woolly Adelgid
JAE R. PASARI
University of California, Santa Cruz
Nitrogen Enrichment
MICHEL PASCAL
Institut National de la Recherche Agronomique (INRA)
Rennes, France
Rats
ANNE M. PERRAULT
Center for International Environmental Law
Washington, DC
Black, White, and Gray Lists
CHARLES PERRINGS
Arizona State University, Tempe
Invasion Economics
GAD PERRY
Texas Tech University, Lubbock
Brown Treesnake
ANDREA J. PICKART
U.S. Fish and Wildlife Service
Arcata, California
Mechanical Control
STEWARD T. A. PICKETT
Cary Institute of Ecosystem Studies
Millbrook, New York
Succession

MICHAEL J. PITCAIRN
California Department of Food and Agriculture, Sacramento
Biological Control, of Plants
 Contributors xix
DANIEL SIMBERLOFF
University of Tennessee, Knoxville
Carnivores
Elton, Charles S.
Kudzu
“Native Invaders”
Rodents (Other)
SCOPE Project
AJAY SINGH
Gorakhpur University, India
Pesticides (Fish and Mollusc)
JENNIFER E. SMITH
University of California, San Diego
Algae
WILLIAM E. SNYDER
Washington State University, Pullman
Ladybugs
NAVJOT S. SODHI
National University of Singapore
Birds
PETER W. SORENSEN
University of Minnesota, St. Paul
Carp, Common
Sea Lamprey
JOHN J. STACHOWICZ
University of California, Davis

Mutualism
KAREN STAHLHEBER
University of California, Santa Barbara
Grasses and Forbs
SCOTT STEINMAUS
California Polytechnic State University
San Luis Obispo, California
Habitat Compatibility
THOMAS J. STOHLGREN
U.S. Geological Survey
Fort Collins, Colorado
Landscape Patterns of Plant Invasions
SHARON Y. STRAUSS
University of California, Davis
Evolutionary Response, of Natives to Invaders
DONALD R. STRONG, JR.
University of California, Davis
Herbivory
BERND SURES
University of Duisburg-Essen, Germany
Parasites, of Animals
DAVID M. RICHARDSON
Stellenbosch University
Matieland, South Africa
Eucalypts
Forestry and Agroforestry
South Africa: Invasions
Trees and Shrubs
DAVID M. RIZZO
University of California, Davis

Fungi
Phytophthora
VINCENT ROBERT
Institute de Recherche pour le Développement
Montpellier, France
Malaria Vectors
GORDON H. RODDA
USGS Fort Collins Science Center
Fort Collins, Colorado
Brown Treesnake
Burmese Python and Other Giant Constrictors
MEGAN A. RÚA
University of North Carolina, Chapel Hill
Pathogens, Plant
JENNIFER RUESINK
University of Washington, Seattle
Ostriculture
JAMES C. RUSSELL
University of California, Berkeley
Predators
KRISTIN SALTONSTALL
Smithsonian Tropical Research Institute
Balboa, Panama
Genotypes, Invasive
NATHAN J. SANDERS
University of Tennessee, Knoxville
Ants
KRISTINA A. SCHIERENBECK
California State University, Chico
Hybridization and Introgression

EUGENE W. SCHUPP
Utah State University, Logan
Dispersal Ability, Plant
PAUL C. SELMANTS
University of California, Santa Cruz
Nitrogen Enrichment
TAMARA SHIGANOVA
Russian Academy of Sciences, Moscow
Ponto-Caspian: Invasions
 xx Contributors
RANDY G. WESTBROOKS
U.S. Geological Survey
Whiteville, North Carolina
Early Detection and Rapid Response
OLAF WEYL
South African Institute for Aquatic Biodiversity
Grahamstown
South Africa: Invasions
DESLEY A. WHISSON
Deakin University
Melbourne, Victoria, Australia
Pesticides (Mammal)
DAVID T. WILLIAMS
Curtin University of Technology
Perth, Western Australia, Australia
Flaviviruses
JOHN R. WILSON
South African National Biodiversity Institute
Matieland
South Africa: Invasions

PHYLLIS N. WINDLE
Union of Concerned Scientists
Washington, DC
Regulation (U.S.)
BENJAMIN WOLFE
Harvard University
Cambridge, Massachusetts
Mycorrhizae
RAM PRATAP YADAV
Gorakhpur University, India
Pesticides (Fish and Mollusc)
HILLARY YOUNG
University of California, Santa Cruz
Nitrogen Enrichment
TRUMAN P. YOUNG
University of California, Davis
Restoration
ERIKA S. ZAVALETA
University of California, Santa Cruz
Nitrogen Enrichment
JOY B. ZEDLER
University of Wisconsin, Madison
Wetlands
PATRICK C. TOBIN
U.S. Department of Agriculture
Morgantown, West Virginia
Gypsy Moth
DAVID R. TOWNS
New Zealand Department of Conservation, Auckland
New Zealand: Invasions

ALEJANDRO E. J. VALENZUELA
Centro Austral de Investigaciones Cientificas,
(CONICET)
Ushuaia, Tierra del Fuego, Argentina
Mammals, Aquatic
WOUTER VAN DER WEIJDEN
Centre for Agriculture and Environment
Culemborg, The Netherlands
Ecoterrorism and Biosecurity
ROY G. VAN DRIESCHE
University of Massachusetts, Amherst
Biological Control, of Animals
KURT J. VAUGHN
University of California, Davis
Restoration
DIEGO P. VÁZQUEZ
Instituto Argentino de Investigaciones del las Zonas Áridas,
CONICET
Mendoza, Argentina
Pollination
ELSE VELLINGA
University of California, Berkeley
Mycorrhizae
MONTSERRAT VILÀ
Estación Biologica de Doñana, Spain
DAISIE Project
THOMAS VIRZI
Rutgers University
New Brunswick, New Jersey
Taxonomic Patterns

BETSY VON HOLLE
University of Central Florida, Orlando
Invasional Meltdown
DAVID A. WARDLE
Swedish University of Agricultural Sciences, Umeå
Belowground Phenomena

The Encyclopedia of Biological Invasions is a comprehen-
sive and authoritative reference dealing with all the physi-
cal and biological aspects of invasive species and invasion
biology and theory. The articles are written by research-
ers and scientific experts and provide a broad overview
of the current state of knowledge with respect to the pat-
terns and processes of invasion, the theories associated
with invasion, and particular accounts of organisms that
have become invasive. Biologists, ecologists, environmen-
tal scientists, geographers, botanists, and zoologists have
contributed reviews intended for students as well as for
the interested general public.
To aid the reader in using this reference, the following
summary describes this Encyclopedia’s features, reviews its
organization and the format of the articles, and is a guide to
the many ways to maximize the utility of this Encyclopedia.
SUBJECTAREAS
The Encyclopedia of Biological Invasions includes 153 topics that
review the various ways scholars have studied invasive species.
The Encyclopedia comprises the following subject areas:
Invader Attributes
•
Ecosystem Features

•
Processes
•
Impacts
•
Notable Taxa
•
Pathways to Invasion
•
Management and Regulation
•
History
•
Notable Invasions
•
ORGANIZATION
Articles are arranged alphabetically by title. An alphabetical
table of contents begins on page v, and another table of con-
tents with articles arranged by subject area begins on page ix.
Article titles have been selected to make it easy to locate
information about a particular topic. Each title begins with
a key word or phrase, sometimes followed by a descriptive
term. For example, “Genotypes, Invasive” is the title assigned
rather than “Invasive Genotypes,” because genotypes is the
key term and is thus more likely to be sought by readers.
Articles that might reasonably appear in different places in
the Encyclopedia are listed under alternative titles—one title
appears as the full entry; the alternative title directs the reader
to the full entry. For example, the alternative title “Coccinel-
lidae” refers readers to the entry entitled “Ladybugs.”

ARTICLEFORMAT
Because the articles in the Encyclopedia are intended for
the interested general public, each article begins with an
introduction that gives the reader a short definition of
the topic and its significance. Here is an example of an
introduction from the article “Phytophthora”:
Phytophthora is a genus of approximately 100 species of
fungal-like, plant pathogenic organisms classified in the
kingdom Stramenopila. Often known as “water molds,”
Phytophthora species have a swimming spore stage (when
they are known as zoospores). Phytophthora species are
well known as pathogens of agricultural, ornamental, and
forest plants. Across the genus, individual Phytophthora
species may infect roots, stems, leaves, flowers, or fruits
of susceptible plants and cause dieback, decline, or death.
Some Phytophthora species are specialists (infecting only
one or a few plant species), and others are generalists
(infecting many plant species across several or many
plant families). The diversity of Phytophthora is quite
astounding, and many new species have been described
in the past ten years. In addition, Phytophthora species
have become one of most important causes of emerging
diseases in native plant communities.

 xxii Guidetotheencyclopedia
GloSSaRy
Almost every topic in the Encyclopedia deals with a subject
that has specialized scientific vocabulary. An effort was
made to avoid the use of scientific jargon, but introducing
a topic can be very difficult without using some unfamil-

iar terminology. Therefore, each contributor was asked to
define a selection of terms used commonly in discussion
of their topic. All these terms have been collated into a
glossary at the back of the volume after the last article.
The glossary in this work includes over 600 terms.
appendiceS
Owing to the great number of currently invasive organ-
isms, it is not possible to include an account of every
invasive species in this Encyclopedia. Therefore, the edi-
tors have included, as an appendix, the IUCN list of the
world’s worst 100 invasive species. (Note that many other
species have also been documented as invasive.) A second
appendix lists key references—important book/volume
references on biological invasions.
indeX
The last section of the Encyclopedia of Biological Invasions
is a subject index consisting of more than 3,800 entries.
This index includes subjects dealt with in each article,
scientific names, topics mentioned within individual arti-
cles, and subjects that might not have warranted a sepa-
rate, stand-alone article.
encyclopediaWeBSite
To access the Encyclopedia of Biological Invasions website,
please visit
/>This site provides a list of the articles, the contribu-
tors, several sample articles, published reviews, and links
to a secure website for ordering copies of the Encyclopedia.
The content of this site will evolve with the addition of
new information.
Within most articles, and especially the longer arti-

cles, major headings help the reader identify important
subtopics within each article. The article “Melastomes”
includes the following headings: “Naturalized and Inva-
sive Melastomes,” “Diversity of Life Forms and Habitats,”
“Reasons for Success,” “Impacts and Control Methods,”
and “Conclusions.”
cRoSS-ReFeRenceS
Many of the articles in this Encyclopedia concern topics for
which articles on related topics are also included. In order
to alert readers to these articles of potential interest, cross-
references are provided at the conclusion of each article. At
the end of “Disturbance,” the following text directs readers
to other articles that may be of special interest:
SeealSotheFolloWinGaRticleS
Fire Regimes / Invasibility, of Communities and Ecosystems /
Land Use / Restoration / Succession / Transformers
Readers will find additional information relating to
Disturbance in the articles listed.
BiBlioGRaphy
Every article ends with a short list of suggestions for “Fur-
ther Reading.” The sources offer reliable in-depth infor-
mation and are recommended by the article’s author or
authors as the best available publications for more lengthy,
detailed, or comprehensive coverage of a topic than can
be feasibly presented within this Encyclopedia. The cita-
tions do not represent all of the sources employed by the
contributor in preparing the article. Most of the listed
citations are to review articles, recent books, or specialized
textbooks, except in rare cases of classic, ground-breaking
scientific articles or articles dealing with subject matter

that is especially new and newsworthy. Thus, the reader
interested in delving more deeply into any particular
topic may elect to consult these secondary sources. This
Encyclopedia functions as ingress into a body of research
only summarized herein.
xxiii
Preface
As you read this, thousands of species of plants, animals,
fungi, and microbes have been or are being transported by
humans to new locations, whether deliberately or inadver-
tently. This geographic rearrangement of the earth’s biota is
one of the great global changes now underway. Of course,
species have always managed to spread, even without human
assistance, but much less often, much more slowly, and not
nearly so far. Over the last 150 years, beginning with the
advent of steamships and accelerating with air travel, the
rate of movement of some organisms has increased many-
fold. Any distance can be quickly spanned by a plane; a
hitchhiking seed, spore, or insect can be transported from
Asia to South America, or from Africa to Australia, in a
day. Although many introduced species fail to establish
populations or remain restricted to the immediate vicinity
of the new sites they land in, others establish populations
and invade new habitats, spreading widely and sometimes
well beyond the initial point of introduction. Their inter-
actions, both within native communities and with other
introduced species, have been noticed by biologists and,
increasingly, by governments and the lay public.
Ecologists, evolutionists, economists, geneticists,
agronomists, fisheries and forestry scientists, and many

others study biological invasions for a variety of reasons.
Some invaders are enormously costly, damaging agricul-
ture, forestry, fisheries, and other human endeavors as well
as natural areas. Other invaders are pathogens that cause
human, animal, or plant disease; yet others are vectors
that carry these pathogens. Many invaders depress pop-
ulations of native species, and even threaten some with
extinction, by preying on them, competing with them,
hybridizing with them, infecting them with disease, or
changing their habitat. The cost of biological invasions
to the economy of the United States alone is estimated at
over $100 billion annually.
However, trying to understand, estimate, and mitigate
the damage caused by some invaders is only one reason for
the rapid growth of research on biological invasions. Inva-
sions are, in a sense, unplanned experiments in population,
community, and ecosystem ecology; sometimes there are
even forms of experimental control and replication—as,
for example, when a species is introduced to certain islands
in an archipelago but not to others. For legal, ethical, and
logistic reasons, most if not all such introductions would be
impossible to perform as planned scientific experiments, so
it is not surprising that scientists rush to investigate them
when such opportunities arise. Initially, most such “purely”
scientific research was done by ecologists interested in such
questions as what impact a wholly new species has on a
native community and ecosystem, and to what extent a
similar native species can impede invasion by a newly intro-
duced species. More recently, recognition that an invasion
constitutes an experiment in both evolution and ecology has

led to an explosion of research on the evolutionary conse-
quences of invasions for both invaders and residents. How,
and how quickly, do invaders evolve adaptations, how do
native communities adapt to an invader, and how does the
small number of invading individuals affect the genetics of
the expanding population of invading organisms?
Biological invasions have attracted attention for other
reasons. The variety of impacts of introduced species is
astounding. Many invasions have such idiosyncratic and
bizarre effects that they cannot fail to arouse our curiosity
simply as fascinating tales of natural history. For exam-
ple, who would have predicted that introducing kokanee
salmon to Flathead Lake, Montana, and, many years later,
opossum shrimp to three nearby lakes would ultimately
have led to population crashes of grizzly bears and bald
eagles through a complicated chain reaction? Or that
introducing myxoma virus to Great Britain to control