Encyclopedia of Biological Invasions 9780520948433

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Encyclopedia of Biological Invasions

Encyclopedia of Biological Invasions Ed i te d 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).

Co nten ts

Contents by Subject Area  /  ix Contributors  /  xiii Guide to the Encyclopedia  /  xxi Preface  /  xxiii

Belowground Phenomena  /  54

Competition, Animal  /  117

David A. Wardle

Christopher R. Dickman

Biological Control, of Animals  /  58

Competition, Plant  /  122

J. Howard Frank; Roy G. Van Driesche;

Jessica Gurevitch

Mark S. Hoddle; E. D. McCoy

Biological Control, of Plants  /  63 Michael J. Pitcairn

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

Birds  /  70 Navjot S. Sodhi

Black, White, and Gray Lists  /  75

Crabs  /  125 Edwin Grosholz

Crayfish  /  129 Francesca Gherardi

Crustaceans (Other)  /  135 Anthony Ricciardi

Stanley W. Burgiel; Anne M. Perrault

Brown Treesnake  /  78 Gad Perry; Gordon H. Rodda

Bryophytes and Lichens  /  81 Franz Essl; Phil Lambdon; Wolfgang Rabitsch

Ants  /  17

Burmese Python and Other Giant Constrictors  /  85

Nathan J. Sanders

Robert N. Reed; Gordon H. Rodda

DAISIE Project  /  138 Petr Pyšek; Philip E. Hulme; Wolfgang Nentwig; Montserrat Vilà

Darwin, Charles  /  142 Marc W. Cadotte

Databases  /  145 Laura A. Meyerson

Apomixis  /  24

Demography  /  147

Takashi Okada; Ross A. Bicknell;

Carol C. Horvitz

Anna M. Koltunow

Aquaculture  /  27 Devin M. Bartley

Aquaria  /  32 Ian C. Duggan

Australia: Invasions  /  36 Tim Low

Canals  /  92 Stephan Gollasch

Carnivores  /  95 Arijana Barun; Daniel Simberloff

Carp, Common  /  100 Peter W. Sorensen; Przemyslaw Bajer

CART and Related Methods  /  104

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

Vojte ˇch Jarošík

Ballast  /  43

Cheatgrass  /  108

James T. Carlton

Richard N. Mack

Bees  /  50

Climate Change  /  113

David Goulson

Early Detection and Rapid Response  /  169

Jeffrey S. Dukes

Randy G. Westbrooks; Robert E. Eplee



v

Earthworms  /  177 Samuel W. James

Ecoterrorism and Biosecurity  /  183 Emiel Elferink; Wouter van der Weijden

Elton, Charles S.  /  187 Daniel Simberloff

Endangered and Threatened Species  /  189 Richard B. Primack

Enemy Release Hypothesis  /  193 Bernd Blossey

Epidemiology and Dispersal  /  196

Geographic Origins and Introduction Dynamics  /  273 Stephen J. Novak

Grasses and Forbs  /  279 Carla D’Antonio; Karen Stahlheber; Nicole Molinari

Daniel Simberloff

Gypsy Moth  /  298 Patrick C. Tobin; Andrew M. Liebhold

Ladybugs  /  400 Edward W. Evans; William E. Snyder

Scott Steinmaus

Marcel Rejmánek; David M. Richardson

Hawaiian Islands: Invasions  /  309

Eutrophication, Aquatic  /  209

Lloyd Loope

Katharina A. M. Engelhardt

Hemlock Woolly Adelgid  /  320

Evolutionary Response, of Natives to Invaders  /  213

David A. Orwig

Sharon Y. Strauss

Joseph M. DiTomaso

Evolution of Invasive Populations  /  215

Herbivory  /  331

Herbicides  /  323

Mara A. Evans; Donald R. Strong, Jr.

Horticulture  /  336 Sarah Reichard

Flaviviruses  /  234

Hybridization and Introgression  /  342 Kristina A. Schierenbeck

Lag Times  /  404 Jeffrey A. Crooks

Lakes  /  410 Hugh J. MacIsaac

Landscape Patterns of Plant Invasions  /  422 Thomas J. Stohlgren

Land Use  /  425 Richard J. Hobbs

Lantana camara  /  428 Stephen Johnson

Laws, Federal and State  /  430 Marc L. Miller

Life History Strategies  /  437 Jennifer L. Bufford; Curtis C. Daehler

Hydrology  /  346 David Le Maitre

Malaria Vectors  /  442 Louis Lambrechts; Anna Cohuet;

John S. Mackenzie; David T. Williams

Vincent Robert

Forest Insects  /  238



William G. Lee

Kudzu  /  396

Eucalypts  /  203

Peter B. Moyle; Emili García-Berthou

Islands  /  391

Edward L. Mills; Kristen T. Holeck

Habitat Compatibility  /  305

Fishes  /  229

James T. Carlton

Great Lakes: Invasions  /  294

Piero Genovesi

Robert C. Klinger

Invertebrates, Marine  /  385

David M. Forsyth

Eradication  /  198

Fire Regimes  /  223

Marcel Rejmánek

Grazers  /  290

Alan Hastings

Carol Eunmi Lee

Invasiveness  /  379

Andrew M. Liebhold;

Influenza  /  350

Mammals, Aquatic  /  445

Deborah G. McCullough

C. W. Potter

Christopher B. Anderson;

Forestry and Agroforestry  /  241

Integrated Pest Management  /  353

David M. Richardson

Robert F. Norris

Freshwater Plants and Seaweeds  /  248

Invasibility, of Communities and Ecosystems  /  356

Lars W. J. Anderson

Jason D. Fridley

Bella S. Galil

Fungi  /  258

Invasional Meltdown  /  360

Melastomes  /  458

Marie-Laure Desprez-Loustau;

Betsy Von Holle

Jean-Yves Meyer; Arthur C. Medeiros

David M. Rizzo

Invasion Biology  /  364

Mosquitoes  /  462

Mark A. Davis

L. Philip Lounibos

Invasion Biology: Historical Precedents  /  369

Mutualism  /  466

Game Animals  /  264 Leonard A. Brennan; Fred C. Bryant

Matthew K. Chew

Mycorrhizae  /  468

Genotypes, Invasive  /  270

Invasion Economics  /  375

Anne Pringle; Benjamin Wolfe;

Kristin Saltonstall

Charles Perrings

Else Vellinga

vi C o n t e n t s

Alejandro E. J. Valenzuela

Mechanical Control  /  449 Andrea J. Pickart

Mediterranean Sea: Invasions  /  452

John J. Stachowicz

Predators  /  557

Snails and Slugs  /  634

“Native Invaders”  /  472

Mick N. Clout; James C. Russell

Robert H. Cowie

Daniel Simberloff

Propagule Pressure  /  561

South Africa: Invasions  /  643

New Zealand: Invasions  /  475

Richard P. Duncan

David M. Richardson; John R. Wilson;

David R. Towns; Peter de Lange;

Protected Areas  /  563

Mick N. Clout

John Randall

Olaf Weyl; Charles L. Griffiths

Succession  /  651 Scott J. Meiners; Steward T. A. Pickett

Nile Perch  /  484 Robert M. Pringle

Nitrogen Enrichment  /  488

Range Modeling  /  568

Jae R. Pasari; Paul C. Selmants; Hillary

Jonathan M. Jeschke

Young; Jennifer O’Leary; Erika S. Zavaleta

Novel Weapons Hypothesis  /  492 Ragan M. Callaway

Rats  /  571 Michel Pascal

Regulation (U.S.)  /  575 Phyllis N. Windle

Remote Sensing  /  580 Ostriculture  /  494 Jennifer Ruesink

Gregory P. Asner; Cho-ying Huang

Reproductive Systems, Plant  /  584 Spencer C. H. Barrett

Parasites, of Animals  /  500 Bernd Sures

Parasitic Plants  /  504 Henning S. Heide-Jørgensen

Pathogens, Animal  /  510 Graham J. Hickling

Pathogens, Human  /  514

Reptiles and Amphibians  /  590

Taxonomic Patterns  /  658 Julie L. Lockwood; Thomas Virzi

Tolerance Limits, Animal  /  661 Alberto Jiménez-Valverde; Jorge M. Lobo

Tolerance Limits, Plant  /  663 James H. Richards; Benjamin R. Janes

Transformers  /  667 Joan G. Ehrenfeld

Trees and Shrubs  /  670 David M. Richardson

Fred Kraus

Restoration  /  594 Truman P. Young; Kurt J. Vaughn

Rinderpest  /  597

Vegetative Propagation  /  678 Jitka Klimes ˘ová

Andy Dobson; Ricardo M. Holdo;

Vines and Lianas  /  680

Robert D. Holt

Carla J. Harris; Rachael Gallagher

Pieter Bol

Risk Assessment and Prioritization  /  604

Pathogens, Plant  /  520

W. M. Lonsdale

Megan A. Rúa; Charles E. Mitchell

Rivers  /  609

Pesticides (Fish and Mollusc)  /  525

Emili García-Berthou; Peter B. Moyle

Ram Pratap Yadav; Ajay Singh

Rodents (Other)  /  612

Martin Hill; Julie Coetzee; Mic Julien;

Pesticides for Insect Eradication  /  528

Daniel Simberloff

Ted Center

Wasps  /  685 Jacqueline Beggs

Water Hyacinth  /  689

Weeds  /  692

Daniel A. Herms; Deborah G. McCullough

Jodie S. Holt

Pesticides (Mammal)  /  535

SCOPE Project  /  617

Desley A. Whisson

Wetlands  /  698

Daniel Simberloff

Joy B. Zedler

Pet Trade  /  539

Sea Lamprey  /  619

Peter T. Jenkins

Peter W. Sorensen; Roger A. Bergstedt

Phytophthora  /  543

Seas and Oceans  /  623

Xenophobia  /  705

David M. Rizzo

Edwin Grosholz

Peter Coates

Pollination  /   546

Seed Ecology  /  627

Diego P. Vázquez; Carolina L. Morales

Michelle R. Leishman; Carla J. Harris

Ponto-Caspian: Invasions  /  549

Small Indian Mongoose  /  631

Zebra Mussel  /  710

Tamara Shiganova

Warren Hays

Ladd Erik Johnson



C o n t e n t s vii

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Co ntents by Sub j ect A r ea

INVADER ATTRIBUTES

Lakes

Allelopathy

Nitrogen Enrichment

Apomixis

Protected Areas

Demography

Rivers

Dispersal Ability, Animal

Seas and Oceans

Dispersal Ability, Plant

Succession

Genotypes, Invasive

Wetlands

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

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

Disturbance Eutrophication, Aquatic



IMPACTS

Habitat Compatibility

Allelopathy

Invasibility, of Communities and Ecosystems

Competition, Animal

Islands

Competition, Plant

ix

Endangered and Threatened Species

Invertebrates, Marine

Fire Regimes

Kudzu

Herbivory

Ladybugs

Hybridization and Introgression

Lantana camara

Hydrology

Malaria Vectors

Invasion Economics

Mammals, Aquatic

Landscape Patterns of Plant Invasions

Melastomes

“Native Invaders”

Mosquitoes

Novel Weapons Hypothesis

Mycorrhizae

Pollination

Nile Perch

Transformers

Parasites, of Animals

Weeds

Parasitic Plants

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



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

Flaviviruses

Acclimatization Societies

Forest Insects

Agriculture

Freshwater Plants and Seaweeds

Aquaculture

Fungi

Aquaria

Grasses and Forbs

Ballast

Grazers

Canals

Gypsy Moth

Forestry and Agroforestry

Hemlock Woolly Adelgid

Game Animals

Influenza

Horticulture

x C o n t e n t s b y S u b j e c t A r e a

Ostriculture

Regulation (U.S.)

Pet Trade

Remote Sensing

MANAGEMENT AND REGULATION

Agreements, International Biological Control, of Animals

Risk Assessment and Prioritization HISTORY

Biological Control, of Plants

DAISIE Project

Black, White, and Gray Lists

Darwin, Charles

CART and Related Methods

Elton, Charles S.

Databases

Invasion Biology

Early Detection and Rapid Response

Invasion Biology: Historical Precedents

Ecoterrorism and Biosecurity

SCOPE Project

Eradication

Xenophobia

Herbicides Integrated Pest Management Laws, Federal and State Mechanical Control Pesticides (Fish and Mollusc) Pesticides for Insect Eradication Pesticides (Mammal) Range Modeling



Restoration

NOTABLE INVASIONS

Australia: Invasions Great Lakes: Invasions Hawaiian Islands: Invasions Mediterranean Sea: Invasions New Zealand: Invasions Ponto-Caspian: Invasions South Africa: Invasions

C o n t e n t s b y S u b j e c t A r e a xi

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Contributors

CHRISTOPHER B. ANDERSON

ROSS A. BICKNELL

University of Magallanes Punta Arenas, Chile Mammals, Aquatic

Plant and Food Research Christchurch, New Zealand Apomixis

LARS W. J. ANDERSON

BERND BLOSSEY

USDA–Agricultural Research Service Davis, California Freshwater Plants and Seaweeds

Cornell University Ithaca, New York Enemy Release Hypothesis

GREGORY P. ASNER

PIETER BOL

Carnegie Institution for Science Stanford, California Remote Sensing

Delft University of Technology Delft, The Netherlands Pathogens, Human

PRZEMYSLAW BAJER

LEONARD A. BRENNAN

University of Minnesota, St. Paul Carp, Common

Texas A&M University, Kingsville Game Animals

SPENCER C. H. BARRETT

FRED C. BRYANT

University of Toronto, Ontario, Canada Reproductive Systems, Plant

Texas A&M University, Kingsville Game Animals

DEVIN M. BARTLEY

JENNIFER L. BUFFORD

California Department of Fish and Game, Sacramento Aquaculture

University of Hawaii, Honolulu Life History Strategies

ARIJANA BARUN

STANLEY W. BURGIEL

University of Tennessee, Knoxville Carnivores

Global Invasive Species Programme Alexandria, Virginia Black, White, and Gray Lists

JACQUELINE BEGGS

University of Auckland, New Zealand Wasps ROGER A. BERGSTEDT

U.S. Geological Survey Millersburg, Michigan Sea Lamprey



MARC W. CADOTTE

University of Toronto, Ontario, Canada Darwin, Charles RAGAN M. CALLAWAY

University of Montana, Missoula Novel Weapons Hypothesis

xiii

JAMES T. CARLTON

PETER DE LANGE

Williams College Mystic, Connecticut Ballast Invertebrates, Marine

New Zealand Department of Conservation, Auckland New Zealand: Invasions

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

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 California, Santa Barbara Grasses and Forbs

University of Maryland Center for Environmental Science, Frostburg Eutrophication, Aquatic

ADAM S. DAVIS

ROBERT E. EPLEE

USDA-ARS Invasive Weed Management Unit Urbana, Illinois Agriculture

Usda Aphis Ppq Whiteville, North Carolina Early Detection and Rapid Response

MARK A. DAVIS

FRANZ ESSL

Macalester College St. Paul, Minnesota Invasion Biology

Environment Agency Austria Vienna, Austria Bryophytes and Lichens

CARLA D’ANTONIO



MARIE-LAURE DESPREZ-LOUSTAU

xiv C o n t r i b u t o r s

EDWARD W. EVANS

EDWIN GROSHOLZ

Utah State University, Logan Ladybugs

University of California, Davis Crabs Seas and Oceans

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

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

Macquarie University Sydney, New South Wales, Australia Vines and Lianas

DANIEL A. HERMS

EMILI GARCÍA-BERTHOU

GRAHAM J. HICKLING

University of Girona, Spain Fishes Rivers

University of Tennessee, Knoxville Pathogens, Animal

PIERO GENOVESI

Rhodes University Grahamstown, South Africa Water Hyacinth

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

Ohio State University, Columbus Pesticides for Insect Eradication

MARTIN HILL

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

DAVID GOULSON

KRISTEN T. HOLECK

University of Stirling, Scotland, United Kingdom Bees

Cornell University, Ithaca, New York Great Lakes: Invasions

CHARLES L. GRIFFITHS

JODIE S. HOLT

University of Cape Town, South Africa South Africa: Invasions

University of California, Riverside Weeds



C o n t r i b u t o r s xv

ROBERT D. HOLT

MIC JULIEN

University of Florida, Gainesville Rinderpest

Commonwealth Scientific and Industrial Research Organisation (CSIRO) Indooroopillay, Queensland, Australia Water Hyacinth

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

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

University of Kansas, Lawrence Earthworms

Bishop Museum Honolulu, Hawaii Reptiles and Amphibians

Benjamin R. Janes

PHIL LAMBDON

University of California, Davis Tolerance Limits, Plant

Royal Botanic Gardens Kew, United Kingdom Bryophytes and Lichens

SAMUEL W. JAMES

ˇ CH JAROŠÍK VOJTE

Charles University Prague, Czech Republic CART and Related Methods PETER T. JENKINS

LOUIS LAMBRECHTS

Institut Pasteur Paris, France Malaria Vectors

Defenders of Wildlife Washington, DC Pet Trade

DOUGLAS A. LANDIS

JONATHAN M. JESCHKE

CAROL EUNMI LEE

Ludwig-Maximilians-University Munich Planegg-Martinsreid, Germany Range Modeling

University of Wisconsin, Madison Evolution of Invasive Populations

Michigan State University, East Lansing Agriculture

WILLIAM G. LEE

Universidad de Málaga, Spain Tolerance Limits, Animal

Landcare Research Dunedin, New Zealand Islands

LADD ERIK JOHNSON

MICHELLE R. LEISHMAN

Université Laval Québec, Canada Zebra Mussel

Macquarie University Sydney, New South Wales, Australia Seed Ecology

STEPHEN JOHNSON

DAVID LE MAITRE

Industry and Investment New South Wales Orange, Australia Lantana camara

Council for Scientific and Industrial Research (CSIR) Stellenbosch, South Africa Hydrology

ALBERTO JIMÉNEZ-VALVERDE



˘ OVÁ JITKA KLIMES

xvi C o n t r i b u t o r s

CHRISTOPHER LEVER

DEBORAH G. MCCULLOUGH

University of Cambridge, United Kingdom Acclimatization Societies

Michigan State University, East Lansing Forest Insects Pesticides for Insect Eradication

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



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

C o n t r i b u t o r s xvii

WOLFGANG NENTWIG

C. W. POTTER

University of Bern, Switzerland DAISIE Project

University of Sheffield, United Kingdom Influenza

ROBERT F. NORRIS

RICHARD B. PRIMACK

University of California, Davis Integrated Pest Management

Boston University, Massachusetts Endangered and Threatened Species

STEPHEN J. NOVAK

ANNE PRINGLE

Boise State University, Idaho Geographic Origins and Introduction Dynamics

Harvard University Cambridge, Massachusetts Mycorrhizae

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

xviii C o n t r i b u t o r s

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

DAVID M. RICHARDSON

DANIEL SIMBERLOFF

Stellenbosch University Matieland, South Africa Eucalypts Forestry and Agroforestry South Africa: Invasions Trees and Shrubs

University of Tennessee, Knoxville Carnivores Elton, Charles S. Kudzu “Native Invaders” Rodents (Other) SCOPE Project

DAVID M. RIZZO

University of California, Davis Fungi Phytophthora

AJAY SINGH

VINCENT ROBERT

JENNIFER E. SMITH

Institute de Recherche pour le Développement Montpellier, France Malaria Vectors

University of California, San Diego Algae

GORDON H. RODDA

Washington State University, Pullman Ladybugs

USGS Fort Collins Science Center Fort Collins, Colorado Brown Treesnake Burmese Python and Other Giant Constrictors

Gorakhpur University, India Pesticides (Fish and Mollusc)

WILLIAM E. SNYDER

NAVJOT S. SODHI

National University of Singapore Birds

MEGAN A. RÚA

PETER W. SORENSEN

University of North Carolina, Chapel Hill Pathogens, Plant

University of Minnesota, St. Paul Carp, Common Sea Lamprey

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

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

C o n t r i b u t o r s xix

PATRICK C. TOBIN

RANDY G. WESTBROOKS

U.S. Department of Agriculture Morgantown, West Virginia Gypsy Moth

U.S. Geological Survey Whiteville, North Carolina Early Detection and Rapid Response

DAVID R. TOWNS

OLAF WEYL

New Zealand Department of Conservation, Auckland New Zealand: Invasions

South African Institute for Aquatic Biodiversity Grahamstown South Africa: Invasions

ALEJANDRO E. J. VALENZUELA

Centro Austral de Investigaciones Cientificas, (CONICET) Ushuaia, Tierra del Fuego, Argentina Mammals, Aquatic

DESLEY A. WHISSON

WOUTER VAN DER WEIJDEN

DAVID T. WILLIAMS

Centre for Agriculture and Environment Culemborg, The Netherlands Ecoterrorism and Biosecurity

Curtin University of Technology Perth, Western Australia, Australia Flaviviruses

ROY G. VAN DRIESCHE

JOHN R. WILSON

University of Massachusetts, Amherst Biological Control, of Animals

South African National Biodiversity Institute Matieland South Africa: Invasions

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



Deakin University Melbourne, Victoria, Australia Pesticides (Mammal)

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

Rutgers University New Brunswick, New Jersey Taxonomic Patterns

TRUMAN P. YOUNG

BETSY VON HOLLE

ERIKA S. ZAVALETA

University of Central Florida, Orlando Invasional Meltdown

University of California, Santa Cruz Nitrogen Enrichment

DAVID A. WARDLE

JOY B. ZEDLER

Swedish University of Agricultural Sciences, Umeå Belowground Phenomena

University of Wisconsin, Madison Wetlands

xx C o n t r i b u t o r s

University of California, Davis Restoration

Guid e to th e En cyc lop edi a

The Encyclopedia of Biological Invasions is a comprehensive and authoritative reference dealing with all the physical and biological aspects of invasive species and invasion biology and theory. The articles are written by researchers and scientific experts and provide a broad overview of the current state of knowledge with respect to the patterns and processes of invasion, the theories associated with invasion, and particular accounts of organisms that have become invasive. Biologists, ecologists, environmental 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 contents 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 “Coccinellidae” 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.

xxi

Within most articles, and especially the longer articles, major headings help the reader identify important subtopics within each article. The article “Melastomes” includes the following headings: “Naturalized and Invasive 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, crossreferences 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 “Further Reading.” The sources offer reliable in-depth information 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 citations 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.



xxii G u i d e t o t h e E n c y c l o p e d i a

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 unfamiliar 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 organisms, it is not possible to include an account of every invasive species in this Encyclopedia. Therefore, the editors 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 articles, and subjects that might not have warranted a separate, stand-alone article. ENCYCLOPEDIA WEBSITE

To access the Encyclopedia of Biological Invasions website, please visit http://www.ucpress.edu/books.php?isbn=9780520264212

This site provides a list of the articles, the contributors, 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.

Pre face

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 inadvertently. 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 manyfold. 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 interactions, 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 agriculture, 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 populations 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. Invasions 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 introduced 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 consequences 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 example, 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 xxiii

introduced rabbit populations would have led to the extirpation of the large blue butterfly there? Who would have suggested that introducing a particular grass species would lead to hybridization with a native congener, subsequent polyploidization, and origin of a new vigorous invasive species that would change entire intertidal ecosystems? Teasing apart such intriguing causal chains is a scientific accomplishment of the first order. But the variety and idiosyncrasy of invasion effects also challenges attempts of invasion biologists to produce general laws or rules and to be able to explain why some introductions have no major impacts, yet others lead to huge invasions. Being able to predict which species will fall in the latter category if introduced, and which in the former, is the elusive holy grail of invasion biology. Choosing topics for this work was difficult; even a large encyclopedia cannot contain every possible topic of interest relating to biological invasions. Had we restricted ourselves simply to species, in addition to the famed “100 of the World’s Worst Invasive Alien Species” (see Appendix), we would still have had to consider several hundred other noteworthy invasive species. Many kinds of habitats, and several regions, have been particularly afflicted by invasions. Certain groups of organisms are especially well represented among invasive species. As the science of invasion biology and technologies of managing invasions have matured, myriad subjects can now be treated in some depth. We could not cover all relevant topics, but we attempted to choose those that, in total, would



xxiv P r e f a c e

treat the broadest possible range of subjects related to biological invasions. The additional readings suggested with the articles and at the end of this volume will help the reader further pursue individual topics and may provide an entree into topics that are not covered individually. We are indebted to the 197 authors who contributed articles to this encyclopedia. Experts are always busy people, yet these made time to produce novel syntheses for a publication quite different from the scientific journals for which they usually write. Gail Rice of the University of California Press assisted us enormously in all facets of producing this encyclopedia—helping us locate experts and photographs, editing articles, keeping track of revisions, reminding tardy authors (and editors!), and even providing aesthetic advice on layout and formats. Her organizational and substantive skills were crucial to the project. Chuck Crumly of the University of California Press suggested the need for this encyclopedia, convinced us that we were the right people to edit it, and encouraged us as impediments arose. Finally, our families (Mary, Tander, and Ruth and Eliška, Honza, and Daniel) were enthusiastic, patient, and encouraging even as the encyclopedia, at times, dominated our lives. Daniel Simberloff University of Tennessee, Knoxville Marcel Rejmánek University of California, Davis

 Acclimatization Societies Christopher Lever University of Cambridge, United Kingdom

Acclimatization societies were organizations formed by groups of like-minded but otherwise diverse individuals (aristocrats, landowners, biologists, agriculturalists, sportsmen, and others), whose mutual interest was the introduction of exotic animals and plants. These societies were formed to improve domestic stock, to supply additional food, to provide new game animals, to satisfy nostalgic yearnings, to control pests, and (in Russia) to substantiate the claims of evolutionists. They died out due to declining and unscientific membership, apathy by the public and scientific bodies, inadequate funding and dwindling revenues, increasingly strict legislation on the introduction of exotic species, and the growing realization that such introductions were ecologically unsound. Acclimatization Societies in France and Its Colonies

From the 1780s onward, Louis Jean Marie Daubenton (1716–1799) was responsible for the increasing involvement of the Jardin des Plantes du Roi in Paris in the study of economic zoology, including the acclimatization of exotic species for commercial and agricultural purposes. In 1793 the Jardin was converted into the Muséum National d’Histoire Naturelle, which in turn was succeeded by the Jardin Zoologique d’Acclimatation.



A In February 1854 a group of savants, under the chairmanship of then director Isidore Geoffroy Sainte-Hilaire (1772–1844), founded La Sociétié Zoologique d’Acc­ limatation for the introduction, acclimatization, and domestication of animals and for the cultivation of plants. Subsequently, satellite societies were formed in Grenoble, Nancy, Algeria, French Guyana, Guadeloupe, ­Martinique, and Réunion. The most important plant introduced by the Société to France was a new variety of potato from Australia, to combat the impact of the same blight, Phytophthora ­infestans, that had caused the potato famine in Britain and Ireland in the 1840s. The most potentially valuable animal importations were Chinese silkworms and ­various species of fish. The principal achievements of the Société were the formation of a menagerie in Paris and the development of agricultural crops in metropolitan France and Algeria. By the late 1860s, membership of the Société and visitors to the menagerie were in decline, and government subsidies and other revenue were dwindling. In 1901 the Société was declared insolvent. Acclimatization Societies in Britain

The prime influence behind the acclimatization movement in Britain was Francis Trevelyan Buckland (1826– 1880). At the time of his birth, Britain was still suffering from the economic consequences of the Napoleonic Wars of 1792–1815 and the Industrial Revolution. During this period, the corn harvests were exceptionally poor, and the wars hindered the importation of corn from abroad. The population, and the price of food, increased dramatically, and the rising labor pool helped to lower wages. It was against this background that Buckland (Fig. 1) began to develop his interest in acclimatization. This

1

gentry; its inability to gain adequate government funding; and a lack of facilities for keeping exotic species, most of which were entrusted to the care of individual members. These factors collectively jeopardized the Society’s ability to differentiate itself from such competitors as the ZSL. Nor was the Society’s progress furthered by the apathy on acclimatization shown by the public and the prestigious British Association for the Advancement of Science. In contrast to the French Société, which examined the commercial and economic benefits of acclimatization to all classes, the Society inclined to the introduction of species to benefit only the upper class. Furthermore, most of the species considered for ­acclimatization by the Society were wholly unsuitable for that purpose. Acclimatization Societies in Australia

Figure 1  Frank Buckland physicking a porpoise. “There was only one

way; so I braved the cold water and jumped into the tank with the ­porpoise. I then held him up in my arms (he was very heavy), and, when I had got him in a favourable position, I poured a good dose of ­sal-volatile and water down his throat with a bottle.” From The Curious World of Frank Buckland by G. H. O. Burgess, quoting from F. T. Buckland’s Curiosities of Natural History (4 vols.) 1857–1872. Richard Bentley, London.

c­ oncept was not new; a similar policy had been declared by the Zoological Society of London (ZSL), founded in 1826. The trigger for the founding of the Society for the Acclimatization of Animals, Birds, Fishes, Insects, and Vegetables within the United Kingdom was a dinner held in London in 1860, attended by Buckland and presided over by the distinguished zoologist Sir ­Richard Owen (1804–1892), at which eland Taurotragus (Tragelaphus) oryx from London Zoo was the principal dish. So impressed by the eland were Owen, Buckland, and the other guests that later that same year, the Society was founded, with ­Buckland as honorary secretary. Also in the same year (1860), a branch was formed in ­Scotland (Glasgow), and in 1861 in the Channel Islands (Guernsey). In 1865, the Society, clearly in financial straits, merged with the Ornithological Society of London. A decline in membership and an apparent lack of interest by the council led in 1868 to the Society’s demise. The principal reasons for the ephemeral life of the Society were its failure to attract enough scientific members, most of whom were drawn from the aristocracy and

2 A c c l i m a t i z a t i o n S o c i e t i e s

Acclimatization societies in Australia were formed in 1879 in New South Wales (in Sydney, having evolved from a society founded in 1852); in 1861 in Victoria (Melbourne); in 1862 in South Australia (Adelaide) and Queensland (Brisbane); in 1895 and 1899 in Tasmania (Hobart and Launceston, respectively); in 1896 in ­Western Australia (Perth); and at various provincial centers. As in Algeria, the activities of acclimatization societies in Australia reached their zenith during the final days of protectionism, especially in such colonies as Victoria, which possessed the most important such society and which, even in the 1860s, depended on protective tariffs. A close parallel can be detected between the position and status of Victoria in the British Empire and Algeria in its French counterpart. The economy of both colonies was remarkably similar, and favorable tariffs on imports and government grants resulted in increasing interest in acclimatization. Algiers and ­Melbourne were both centers of rapid demographic growth. In Australia, the acclimatization movement met with the same apathy as in Britain, based on the belief that the societies were acting in the interest of the privileged minority (Fig. 2). The societies claimed that their introduction of insectivorous birds increased crop production, while pastoralists claimed that they consumed crops and displaced native birds. Deer provided sport and venison but damaged crops and trees. Eventually, many societies degenerated into importing species solely as curiosities or for ornamental purposes, and some metamorphosed into menageries. Few, if any, attempted to “improve” domestic stock or cultivars on which the prosperity of Australia depended. Those that survive are involved mainly with the introduction of fish.

Figure 2  Cartoon from the Melbourne Punch, May 26, 1884.

Acclimatization Societies in New Zealand

The thirty or so acclimatization societies that were formed in New Zealand between the 1860s (the first in Nelson in 1861) and the early 1900s had, as in the case of many of those elsewhere, two principal objectives: the introduction of game animals for sport and insect-eating birds to control pests. As in other countries, some of the New Zealand societies eventually failed due to lack of support and falling revenues, coupled with increasing public criticism. Founded and run, as in Britain, by enthusiastic amateurs, they were managed unprofessionally, and they failed to keep adequate records that would have shown their critics the value of revenue derived from visiting sportsmen and the benefit to crops. After the Second World War, control of the societies by the government increased, and their operations became mainly confined to conservation; the promotion of sport; and, in a complete role reversal, the prevention of further introductions of nonnative species. The main income of acclimatization societies in New Zealand today comes from the sale of sporting licenses; part of this income is used to acquire wetland habitats, fund research, and educate the public on conservation issues: the balance funds the societies’ own conservation programs. Acclimatization Societies in Russia

An interest in the acclimatization and domestication of nonnative animals and plants existed in Russia since at least the early 1840s and was led by the distinguished biologist Karl Frantsevich Rul’e (1814–1858).

The primary topic among scientists at the time was the immutability or mutability of species. Rul’e used the transformation of species through acclimatization, domestication, and cultivation to support the theory of mutability (evolution). Under the leadership of Rul’e, the Imperial Russian Society for the Acclimatization of Animals and Plants was formed in Moscow in January 1864; branches were later established in St Petersburg, Khar’kov, and Orel. After Rul’e’s death, his successors, led by his protégé Anatoli Petrovich Bogdanov (1834–1896), continued his work. As early as 1856, Bogdanov and his colleagues had formulated the idea of establishing a scientifically based zoo in Moscow. Almost from the start, however, dissent broke out between those who gave preference to pure research and those who favored applied research in acclimatization, domestication, and hybridization. This controversy was soon overshadowed by the zoo’s financial failure, though it rumbled on well into the twentieth century and had a profound effect on the development of Russian science. Thereafter, the Society began to stagnate, although outside its ranks, the interest in acclimatization actually increased, in particular in the translocation of native fur bearers and the widespread formation of many research sad (gardens). By the early 1900s, it had become accepted within the Society that acclimatization must give way to conservation, and that the introduction of exotics could be actually harmful. By 1930 the Society had ceased to exist. Although during its 65 years the Society failed to acclimatize (naturalize) any alien species in Russia, it did encourage local attempts in acclimatization of a wide range of, albeit as in Britain, wholly unsuitable, species. Acclimatization Societies in the United States

Although since as early as 1846, songbirds, including the house sparrow Passer domesticus, were successfully released in the United States, the founding father of the acclimatization movement was a New York pharmacist, Eugene Schiefflin, who, with John Avery, in 1871 founded the American Acclimatization Society, which in 1877 successfully released the first European starlings Sturnus vulgaris in Central Park. In 1873 Andrew Erkenbrecher formed the Cincinnati Society of Acclimatization, which in 1873–1874 unsuccessfully (except in the case of house sparrows) released in the city 21 alien bird species. At about the same time, the Society for the Acclimatization of Foreign Birds was founded in Cambridge, A c c l i m at i z at i o n S o c i e t i e s 3

Massachusetts. In 1872–1874, it freed large numbers of goldfinches Carduelis carduelis, some of which survived until at least the turn of the century. In Portland, Oregon, in 1880, C. F. Pfluger founded the Society for the Introduction of Useful Songbirds into Oregon (the Portland Songbird Club), which in 1889 and 1892 unsuccessfully (except in the case of the ­European starling) released 15 species. These societies spawned several others throughout the United States, of which the Country Club of San ­Francisco was formed mainly to introduce brown trout Salmo trutta to California. It also dispatched chinook salmon Oncorhynchus tshawytscha ova to New Zealand, where, under the name quinnat salmon, it became a ­popular game fish. In 1884 the Cincinnati Society of Natural History rightly stated that the introduction of alien species was ecologically unsound, a pronouncement that seems to have sounded the death knell for acclimatization in the continental United States. Acclimatization Societies in the Hawaiian Islands

Although since 1865 private individuals had released in the Hawaiian Islands (then a territory of the United States) a number of bird species with varying degrees of success, it was not until 1930, under the presidency of Mrs. ­Frederick J. Lowery, that the Hui Manu (Hawaiian for “bird society”) was formed, for the introduction of songbirds to the islands. In the same year, immigrants from Japan founded the Honolulu Mejiro (the national name of the Japanese white-eye Zosterops japonica) ­Society, specifically for the introduction of Japanese songbirds. Among the species successfully freed by these two ­organizations were northern mockingbirds Mimus polyglottos, white-rumped shamas Copsychus malabaricus, Japanese bush-warblers Cettia diphone, varied tits Parus varius, Japanese whiteeyes, red-crested cardinals Paroaria coronata, northern cardinals Cardinalis cardinalis, red avadavats Amandava amandava, and black-headed mannikins or munias Lonchura malacca. In 1968, diminishing funds and increasingly strict regulations about importing and releasing alien birds in the islands caused the Hui Manu to disband. Acclimatization Societies in Germany and Italy

In 1858 the Akklimatisations-verein was formed in ­Berlin, and in 1861 the Società di Acclimazione in Palermo, Sicily.

4