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CONCEPTUAL ECOLOGY AND INVASION BIOLOGY:
RECIPROCAL APPROACHES TO NATURE
INVADING NATURE -
SPRINGER SERIES IN INVASION ECOLOGY
Volume 1
Series Editor: JAMES A. DRAKE
University of Tennessee,
Knoxville, TN, U.S.A.
Conceptual Ecology and Invasion
Biology: Reciprocal Approaches
to Nature
Edited by
MARC WILLIAM CADOTTE
University of Tennessee, Knoxville, U.S.A.
SEAN M. MCMAHON
University of Tennessee, Knoxville, U.S.A.
and
TADASHI FUKAMI
University of Hawaii at Manoa,
Honolulu, U.S.A.
A C.I.P. Catalogue record for this book is available from the Library of Congress.
ISBN-10 1-4020-4158-6 (PB)
ISBN-13 978-1-4020-4158-7 (PB)
ISBN-10 1-4020-4157-8 (HB)
ISBN-13 978-1-4020-4157-0 (HB)
ISBN-10 1-4020-4925-0 ( e-book)
ISBN-13 978-1-4020-4925-5 (e-book)
Published by Springer,
P.O. Box 17, 3300 AA Dordrecht, The Netherlands.
www.springer.com
Printed on acid-free paper
Cover design by Jeff McMahon
All Rights Reserved
2006 Springer
No part of this work may be reproduced, stored in a retrieval system, or transmitted
in any form or by any means, electronic, mechanical, photocopying, microfilming, recording
or otherwise, without written permission from the Publisher, with the exception
of any material supplied specifically for the purpose of being entered
and executed on a computer system, for exclusive use by the purchaser of the work.
Printed in the Netherlands.
#ONTENTS
#ONTRIBUTORS IX
&OREWORD XV
0REFACE XVII
)NTRODUCTION HISTORY AND TERMINOLOGY
4RACKING THE TRACTABLE USING INVASIONS TO GUIDE
THE EXPLORATION OF CONCEPTUAL ECOLOGY
3EAN - -C-AHON -ARC 7 #ADOTTE AND 4ADASHI &UKAMI
$ARWIN TO %LTON EARLY ECOLOGY AND THE PROBLEM
OF INVASIVE SPECIES
-ARC 7 #ADOTTE
)NVASION BIOLOGY
THE PURSUIT OF SCIENCE AND CONSERVATION
-ARK ! $AVIS
)NVASIVENESS IN EXOTIC PLANTS
IMMIGRATION AND NATURALIZATION IN AN ECOLOGICAL CONTINUUM
(ELEN 4 -URPHY *EREMY 6AN$ER7AL ,ESLEY ,OVETT
$OUST AND *ON ,OVETT
$OUST
0OPULATIONS AT PLAY
$ENSITY DEPENDENCE IN INVASIVE PLANTS
DEMOGRAPHY HERBIVORY SPREAD AND EVOLUTION
9VONNE - "UCKLEY AND *ESSICA -ETCALF
V
VI #ONTENTS
3TOCHASTICITY NONLINEARITY AND INSTABILITY IN BIOLOGICAL INVASIONS
2OBERT 0 &RECKLETON 0ETER - $OWLING AND .ICHOLAS + $ULVY
,OCAL INTERACTIONS AND INVASION DYNAMICS
POPULATION GROWTH IN SPACE AND TIME
$AVID * -URRELL
! GUIDE TO CALCULATING DISCRETE TIME INVASION RATES FROM DATA
-ARK ! ,EWIS -ICHAEL ' .EUBERT (AL #ASWELL *AMES 3 #LARK AND
+ATRIONA 3HEA
4HE ROLE OF EVOLUTIONARY GENETICS IN STUDIES OF PLANT INVASIONS
+RISTINA ! 3CHIERENBECK AND -ALIKA , !NOUCHE
5NWELCOMED VISITOR SPECIES INTERACTIONS
#ONTACT EXPERIENCE ALIEN NATIVE INTERACTIONS
AND THEIR COMMUNITY CONSEQUENCES A THEORETICAL CONSIDERATION
ON THE ROLE OF ADAPTATION IN BIOLOGICAL INVASION
-ICHIO +ONDOH
5SE OF BIOLOGICAL INVASIONS AND THEIR CONTROL TO STUDY
THE DYNAMICS OF INTERACTING POPULATIONS
&RANCK #OURCHAMP AND 3TEPHANE #AUT
)NVASIBILITY OF SEED PREDATORS ON SYNCHRONIZED AND
INTERMITTENT SEED PRODUCTION OF HOST PLANTS
!KIKO 3ATAKE /TTAR . "JRNSTAD AND 9OH )WASA
)NVASIONS AND THE REGULATION OF PLANT POPULATIONS BY PATHOGENS
'REGORY 3 'ILBERT AND )NGRID - 0ARKER
%XPLORING THE RELATIONSHIP BETWEEN NICHE BREADTH
AND INVASION SUCCESS
$IEGO 0 6 ZQUEZ
#ONTENTS VII
)NTERACTIONS BETWEEN INVASIVE PLANTS AND SOIL ECOSYSTEMS
POSITIVE FEEDBACKS AND THEIR POTENTIAL TO PERSIST
!NDREA 3 4HORPE AND 2AGAN - #ALLAWAY
)NVASION BIOLOGY AS A COMMUNITY PROCESS
MESSAGES FROM MICROBIAL MICROCOSMS
0HILIP ( 7ARREN 2ICHARD ,AW AND !NITA * 7EATHERBY
,ARGE SCALE CONSEQUENCES AND PATTERNS OF INVASIONS
5NDERSTANDING INVASIONS IN PATCHY HABITATS
THROUGH METAPOPULATION THEORY
+ARIN # (ARDING *OHN - -C.AMARA AND 2OBERT $ (OLT
#OMPETITION AND THE ASSEMBLY OF INTRODUCED BIRD COMMUNITIES
2ICHARD 0 $UNCAN AND $AVID - &ORSYTH
2OOM FOR ONE MORE %VIDENCE FOR INVASIBILITY AND
SATURATION IN ECOLOGICAL COMMUNITIES
3COTT ! 3MITH AND *ONATHAN " 3HURIN
4HE BIOGEOGRAPHY OF NATURALIZED SPECIES AND
THE SPECIES
AREA RELATIONSHIP
RECIPROCAL INSIGHTS TO BIOGEOGRAPHY AND INVASION BIOLOGY
$OV & 3AX AND 3TEVEN $ 'AINES
3YNTHESIS
,INKING SCALE DEPENDENT PROCESSES IN INVASIONS
-ARC 7 #ADOTTE 4ADASHI &UKAMI AND 3EAN - -C-AHON
)NDEX
#ONTRIBUTORS
-ALIKA , !NOUCHE
$EPARTMENT OF %VOLUTIONARY %COLOGY 'ENOME %VOLUTION AND 3PECIATION LAB 5-2
#.23 5NIVERSITY OF 2ENNES #AMPUS DE "EAULIEU 2ENNES CEDEX
&RANCE
/TTAR . "JRNSTAD
$EPARTMENT OF %NTOMOLOGY AND "IOLOGY 4HE 0ENNSYLVANIA 3TATE 5NIVERSITY
5NIVERSITY 0ARK 0! 53!
9VONNE - "UCKLEY
4HE %COLOGY #ENTRE (INES "UILDING 5NIVERSITY OF 1UEENSLAND 3T ,UCIA "RISBANE
1,$ !USTRALIA
-ARC 7 #ADOTTE
$EPARTMENT OF %COLOGY AND %VOLUTIONARY "IOLOGY $ABNEY (ALL 5NIVERSITY OF
4ENNESSEE +NOXVILLE 4. 53!
2AGAN - #ALLAWAY
$IVISION OF "IOLOGICAL 3CIENCES 4HE 5NIVERSITY OF -ONTANA -ISSOULA -4
53!
(AL #ASWELL
7OODS (OLE /CEANOGRAPHIC )NSTITUTION 7OODS (OLE -! 53!
*AMES 3 #LARK
.ICHOLAS 3CHOOL OF THE %NVIRONMENT AND %ARTH 3CIENCES ! ,32# "OX
$UKE 5NIVERSITY $URHAM .# 53!
IX
X #ONTRIBUTORS
3TEPHANE #AUT
%COLOGIE 3YSTMATIQUE %VOLUTION "T
5NIVERSIT 0ARIS
3UD 8) &
/RSAY #EDEX &RANCE
&RANCK #OURCHAMP
%COLOGIE 3YSTMATIQUE %VOLUTION "T
5NIVERSIT 0ARIS
3UD 8) &
/RSAY #EDEX &RANCE
-ARK ! $AVIS
$EPARTMENT OF "IOLOGY -ACALESTER #OLLEGE 3AINT 0AUL -. 53!
0ETER - $OWLING
.37 !GRICULTURE !USTRALIAN #ENTRE FOR 7EED 2ESEARCH AND #2# FOR 7EED
-ANAGEMENT 3YSTEM !GRICULTURAL AND 6ETERINARY #ENTRE &OREST 2OAD /RANGE
.37 .37 !USTRALIA
.ICHOLAS + $ULVY
%COSYSTEM 2ESEARCH 'ROUP #ENTRE FOR %NVIRONMENT &ISHERIES AND !QUACULTURE
3CIENCE 0AKElELD 2OAD ,OWESTOFT 3UFFOLK .2 /(4 5+
2ICHARD 0 $UNCAN
%COLOGY AND %NTOMOLOGY 'ROUP 0/ "OX ,INCOLN 5NIVERSITY #ANTERBURY
.EW :EALAND AND ,ANDCARE 2ESEARCH 0/ "OX ,INCOLN .EW :EALAND
$AVID - &ORSYTH
!RTHUR 2YLAH )NSTITUTE FOR %NVIRONMENTAL 2ESEARCH $EPARTMENT OF 3USTAINABILITY
AND %NVIRONMENT "ROWN 3TREET (EIDELBERG 6ICTORIA !USTRALIA
2OBERT 0 &RECKLETON
$EPARTMENT OF :OOLOGY 5NIVERSITY OF /XFORD 3OUTH 0ARKS 2OAD /XFORD /8 03 5+
4ADASHI &UKAMI
$EPARTMENT OF :OOLOGY 5NIVERSITY OF (AWAII AT -ANOA -C#ARTHY -ALL
%DMONDSON (ONOLULU () 53!
3TEVEN $ 'AINES
$EPARTMENT OF %COLOGY %VOLUTION AND -ARINE "IOLOGY 5NIVERSITY OF #ALIFORNIA
3ANTA "ARBARA #! 53!
#ONTRIBUTORS XI
'REGORY 3 'ILBERT
%NVIRONMENTAL 3TUDIES 5NIVERSITY OF #ALIFORNIA 3ANTA #RUZ #! 53!
+ARIN # (ARDING
$EPARTMENT OF -ARINE %COLOGY 'TEBORG 5NIVERSITY "OX 'TEBORG
3WEDEN
2OBERT $ (OLT
$EPARTMENT OF :OOLOGY 5NIVERSITY OF &LORIDA "ARTRAM (ALL 0/ "OX
'AINESVILLE &LORIDA 53!
9OH )WASA
$EPARTMENT OF "IOLOGY &ACULTY OF 3CIENCES +YUSHU 5NIVERSITY &UKUOKA
*APAN
-ICHIO +ONDOH
$EPARTMENT OF %NVIRONMENTAL 3OLUTION 4ECHNOLOGY &ACULTY OF 3CIENCE AND
4ECHNOLOGY 2YUKOKU 5NIVERSITY
9OKOYA 3ETA /E
CHO /TSU
*APAN AND #ENTER FOR %COLOGICAL 2ESEARCH +YOTO 5NIVERSITY +AMITANAKAMI /TSU
*APAN
2ICHARD ,AW
$EPARTMENT OF "IOLOGY 5NIVERSITY OF 9ORK 0/ "OX 9ORK 9/ 97 5+
-ARK ! ,EWIS
$EPARTMENT OF -ATHEMATICAL AND 3TATISTICAL 3CIENCES 5NIVERSITY OF !LBERTA
#ENTRAL !CADEMIC "UILDING %DMONTON !" 4' ' #ANADA
*ON ,OVETT
$OUST
$EPARTMENT OF "IOLOGY 5NIVERSITY OF 7INDSOR 7INDSOR /NTARIO ." 0 #ANADA
,ESLEY ,OVETT
$OUST
$EPARTMENT OF "IOLOGY 5NIVERSITY OF 7INDSOR 7INDSOR /NTARIO ." 0
#ANADA
3EAN - -C-AHON
$EPARTMENT OF %COLOGY AND %VOLUTIONARY "IOLOGY $ABNEY (ALL 5NIVERSITY OF
4ENNESSEE +NOXVILLE 4. 53!
XII #ONTRIBUTORS
*OHN - -C.AMARA
$EPARTMENT OF -ATHEMATICS 5NIVERSITY OF "RISTOL 5NIVERSITY 7ALK "RISTOL "3
47 5+
*ESSICA -ETCALF
$EPARTMENT OF "IOLOGY )MPERIAL #OLLEGE ,ONDON 3ILWOOD 0ARK #AMPUS !SCOT
"ERKSHIRE 3, 09 5+
(ELEN -URPHY
$EPARTMENT OF "IOLOGY 5NIVERSITY OF 7INDSOR 7INDSOR /NTARIO ." 0
#ANADA 0RESENT ADDRESS #3)2/ 3USTAINABLE %COSYSTEMS 0/ "OX !THERTON
1,$ !USTRALIA
$AVID *OHN -URRELL
.%2# #ENTRE FOR 0OPULATION "IOLOGY )MPERIAL #OLLEGE ,ONDON 3ILWOOD 0ARK
#AMPUS !SCOT "ERKSHIRE 3, 09 5+
-ICHAEL ' .EUBERT
7OODS (OLE /CEANOGRAPHIC )NSTITUTION 7OODS (OLE -! 53!
)NGRID - 0ARKER
%COLOGY AND %VOLUTIONARY "IOLOGY 5NIVERSITY OF #ALIFORNIA 3ANTA #RUZ #!
53!
!KIKO 3ATAKE
#ENTER FOR %COLOGICAL 2ESEARCH +YOTO 5NIVERSITY /TSU
*APAN
$OV & 3AX
)NSTITUTE OF %COLOGY 5NIVERSITY OF 'EORGIA !THENS 'EORGIA 53!
+RISTINA ! 3CHIERENBECK
$EPARTMENT OF "IOLOGICAL 3CIENCES #ALIFORNIA 3TATE 5NIVERSITY #HICO #HICO #!
53!
+ATRIONA 3HEA
"IOLOGY $EPARTMENT 0ENNSYLVANIA 3TATE 5NIVERSITY -UELLER ,AB 5NIVERSITY
0ARK 0!
53!
#ONTRIBUTORS XIII
*ONATHAN " 3HURIN
$EPARTMENT OF :OOLOGY 5NIVERSITY OF "RITISH #OLUMBIA 5NIVERSITY "LVD
6ANCOUVER "# 64 : #ANADA
3COTT ! 3MITH
$EPARTMENT OF "IOLOGY -C'ILL 5NIVERSITY !VENUE $R 0ENlELD -ONTREAL
1UEBEC (! " #ANADA
!NDREA 3 4HORPE
$IVISION OF "IOLOGICAL 3CIENCES 4HE 5NIVERSITY OF -ONTANA -ISSOULA -4
53!
*EREMY 6AN$ER7AL
$EPARTMENT OF "IOLOGY 5NIVERSITY OF 7INDSOR 7INDSOR /NTARIO ." 0
#ANADA
$IEGO 0 6 ZQUEZ
)NSTITUTO !RGENTINO DE )NVESTIGACIONES DE LAS :ONAS !RIDAS #ENTRO 2EGIONAL DE
)NVESTIGACIONES #IENTIlCAS Y 4ECNOLOGICAS !V 2UIZ ,EAL SN -ENDOZA
!RGENTINA
0HILIP ( 7ARREN
$EPARTMENT OF !NIMAL AND 0LANT 3CIENCES 5NIVERSITY OF 3HEFlELD 3HEFlELD 3
4. 5+
!NITA 7EATHERBY
4HE 0ONDS #ONSERVATION 4RUST 0OLICY AND 2ESEARCH CO "-3 /XFORD "ROOKES
5NIVERSITY 'IPSY ,ANE (EADINGTON /XFORD /8 "0 5+
&OREWORD
h7HEN ) TOOK THIS PLOT OF LAND lVE YEARS AGO v HE SAID
h) WAS DISAPPOINTED TO lND THAT INSTEAD OF ITS BEING ONE
WITH THE RICH SANDY SOIL OF THE DISTRICT IT WAS PRACTICALLY A
POCKET OF CLAY WHICH HAD TO BE PUT UNDER TREATMENT BEFORE
IT WOULD YIELD THE RESULT ) DESIRED ) HAVE BEEN FAIRLY SUC
CESSFUL AND NOW HAVE MORE THAN TWO HUNDRED SUBTROPICAL
SHRUBS mOURISHING OUT THERE IN THE GROUNDS )T HAS BEEN
AN EXPERIMENT IN ACCLIMATISATION WITH WHICH ) AM WELL
PLEASED -ANY OF THE PLANTS CAME FROM !USTRALIA AND
.EW :EALAND )N MOST CASES ) HAVE BOUGHT THE SEED AND
REARED THEM ENTIRELY BY MYSELFv
!LFRED 2USSEL 7ALLACE
FROM AN INTERVIEW BY % ( 2ANN
1 7HAT ARE THE CHARACTERISTICS OF A GOOD INVADER
! )T DEPENDS
1 $EPENDS ON WHAT
! 4HAT DEPENDS TOO
"IOLOGICAL INVASION HAS BECOME ONE OF THOSE RARE THEMES SO PROFOUND IN ITS IMPLI
CATIONS AND SCOPE THAT IT CUTS A BROAD PATH ACROSS THE ACADEMIC DISCIPLINES &ROM
GENES TO ECOSYSTEMS AND ECONOMICS TO LAW SPECIES INCURSIONS HAVE CONFRONTED THE
GLOBAL COMMUNITY WITH A MOST TRANSCENDENT 'ORDIAN KNOT /URS IS A PECULIAR KNOT
HOWEVER AS INVADERS COLONIZE ECOLOGICAL SYSTEMS THAT HAVE BEEN FASHIONED FROM
STRANDS OF COMPLEXITY WITH ORIGINS IN EVOLUTIONARY TIME ! HALF
CENTURY HAS PASSED
SINCE THE PUBLICATION OF %LTONS DELIGHTFUL IF NOT FOREBODING TREATISE /F COURSE
%LTONS ASSESSMENT WAS CORRECT AND INDEED THE PROBLEM IS EVEN GREATER TODAY
"IOLOGICAL INVASIONS HAVE BECOME A PIVOTAL FORCE IN DIRECTING THE FUTURE EXPRESSION
OF NATURE 7HILE CONSIDERABLE PROGRESS HAS BEEN MADE GENERALITY REMAINS ELUSIVE
PREDICTION HIT
OR
MISS AND INVASIONS ARE PROCEEDING LARGELY UNCHECKED
XV
XVI &OREWORD
!DOPTING AN INTEGRATIVE APPROACH #ONCEPTUAL %COLOGY AND )NVASIONS "IOLOGY
2ECIPROCAL !PPROACHES TO .ATURE OFFERS ESSENTIAL INSIGHTS INTO THE TOPOLOGY OF OUR
ECOLOGICAL KNOT 4HE EDITORS HAVE BROUGHT TOGETHER A SUPERB CAST OF SCIENTISTS
WITH A SINGULAR AIM ORCHESTRATING THE DIVERSITY OF APPROACHES TO BIOLOGICAL INVA
SIONS INTO A MORE COHERENT AND SYNTHETIC WHOLE 7HAT EMERGES FROM THIS EFFORT IS
COMPELLING FRAMEWORK THAT CAPTURES THE INTRICACY AND NUANCE OF THE PROCESSES
DYNAMICS AND MECHANISMS AT PLAY WHEN SPECIES INVADE NATURE 7HILE NO ONE
WOULD ARGUE THAT THIS FRAMEWORK IS COMPLETE IT IS CLEAR THAT ANY SOLUTION TO THE
PROBLEM OF BIOLOGICAL INVASIONS IS FOUNDED HEREIN
4HIS BOOK IS THE INAUGURAL VOLUME IN 3PRINGERS )NVADING .ATURE SERIES AND
) AM PLEASED TO HAVE HAD THE OPPORTUNITY TO INTRODUCE BOTH
*AMES $RAKE 3ERIES %DITOR
0REFACE
(UMAN ACTIVITIES AFFECT NATURAL SYSTEMS IN EVERY CORNER OF THE GLOBE AND ONE
OF THE MOST WIDESPREAD AND POTENTIALLY DISRUPTIVE OF THESE IS THE INTRODUCTION OF
NON
INDIGENOUS SPECIES INTO NEW ENVIRONMENTS 4HESE INTRODUCTIONS CAN HAVE
PROFOUND CONSEQUENCES FOR RESIDENT POPULATIONS COMMUNITY DYNAMICS AND ECO
SYSTEM FUNCTION /VER AND ABOVE THE NEED TO UNDERSTAND HOW A NON
INDIGENOUS
SPECIES CAN AFFECT A NATIVE COMMUNITY INTRODUCTIONS OFFER ECOLOGISTS THE POTENTIAL
TO LEARN ABOUT HOW COMMUNITIES ARE PUT TOGETHER HOW SPECIES CONTRIBUTE TO
ECOSYSTEM FUNCTION OR HOW POPULATIONS EVOLVE IN THE FACE OF NOVEL ENVIRONMENTS
AND NOVEL SPECIES *UST LIKE THE RAILROAD SPIKE THAT PIERCED 0HINEAS 'AGES SKULL
TAUGHT PHYSICIANS MUCH ABOUT THE FUNCTION OF THE CEREBRAL CORTEX ENVIRONMENTAL
PERTURBATIONS CAN TEACH US ABOUT NATURAL SYSTEMS )NVASIONS CAN INFORM ECOLOGI
CAL THEORY BY SERVING AS NATURAL EXPERIMENTS
/N THE OTHER HAND MODERN
DAY ECOLOGISTS USE SOPHISTICATED CONCEPTUAL TOOLS
TO AID IN THE UNDERSTANDING OF THE FUNCTIONING OF NATURAL SYSTEMS 4HESE TOOLS
WE FEEL HAVE BEEN LARGELY UNDER
UTILIZED IN UNDERSTANDING SPECIES INVASIONS
7E SEE HERE THE POTENTIAL FOR A RECIPROCAL APPROACH TO GAIN A DEEPER UNDERSTAND
ING OF HOW ECOLOGICAL SYSTEMS ARE PUT TOGETHER
7E HAVE ASSEMBLED AN EXCELLENT GROUP OF AUTHORS THAT EACH IN THEIR OWN WAY
ATTEMPT TO USE THEIR TOOLS TO EXPLORE THIS RECIPROCAL RELATIONSHIP 4HE AUTHORS IN
THIS VOLUME WERE INVITED BECAUSE OF THEIR RECENT PUBLICATIONS WHICH SHOW THEIR
USE OF NOVEL APPROACHES TO EITHER UNDERSTANDING THE CAUSES AND CONSEQUENCES OF
SPECIES INVASIONS OR ELSE USE CONCEPTUAL TOOLS TO UNDERSTAND HOW NATURAL SYSTEMS
FUNCTION 7E ASKED AUTHORS REGARDLESS OF THEIR INDIVIDUAL APPROACHES TO THINK
ABOUT THIS RECIPROCAL RELATIONSHIP BETWEEN INVASIONS AND THEORY 4HE CHAPTERS
HERE USE EXAMPLES FROM NUMEROUS ORGANISMS ECOLOGICAL SYSTEMS AND GEOGRAPHIC
LOCATIONS AND ARE EVENLY SPLIT BETWEEN THEORY AND NATURAL EXAMPLES
4HIS VOLUME COULD NOT HAVE BEEN POSSIBLE WITHOUT THE INVOLVEMENT OF OTHERS
7E ARE GRATEFUL lRST AND FOREMOST TO THE AUTHORS FOR PRODUCING SUCH lNE WORK FOR
US 7E WISH TO THANK THE CHAPTER REVIEWERS WHO AS PER USUAL IN SCIENCE SELmESSLY
REVIEWED CHAPTERS AND GREATLY IMPROVED THE QUALITY OF THIS BOOK 7E OFFER A SPE
CIAL THANKS TO THE SERIES EDITOR AND OUR MENTOR *IM $RAKE WHO ALLOWED US TO FOCUS
OUR CREATIVE ENERGIES ON TO THIS PROJECT AND TO AGREE TO MAKE THIS THE PREMIERE
XVII
XVIII 0REFACE
VOLUME OF THIS BOOK SERIES &INALLY 3UZANNE -EKKING AND -ARTINE VAN "EZOOIJEN
AT 3PRINGER FORMED A SUPREMELY ENCOURAGING AND WELL
ORGANIZED PUBLISHING
TEAM INSTRUMENTAL IN MAKING THIS BOOK A REALITY
0:&
600 7)
.QR[YLOOH7HQQHVVHH /LQFROQ1HZ=HDODQG
-XO\
#HAPTER ONE
4RACKING THE TRACTABLE
USING INVASIONS TO GUIDE
THE EXPLORATION OF CONCEPTUAL
ECOLOGY
3 - -C-AHON - 7 #ADOTTE AND 4 &UKAMI
).42/$5#4)/.
/UR UNDERSTANDING OF THE COMPLEX FUNCTIONING OF HUMAN PHYSIOLOGY NEUROLOGY
AND BEHAVIOR HAS LONG BEEN GUIDED BY THE STUDY OF ANOMALIES IN THESE SYSTEMS
7HEN FACED WITH A SEEMINGLY INTRACTABLY COMPLEX SYSTEM IT IS AT TIMES EASIER
TO FATHOM ITS MECHANISMS BY DISRUPTING OR DISTURBING IT RATHER THAN TRYING TO
DELICATELY EXPLORE THE @NATURAL ENTITY )T HAS RECENTLY BECOME CLEAR TO ECOLOGISTS
THIS TACT OF USING THE DISRUPTION OR DISTORTION OF A SYSTEM TO BETTER UNDERSTAND
ITS NORMAL WORKINGS CAN BE APPLIED TO THE STUDY OF THE BEHAVIOR SPREAD AND
IMPACT OF NON
INDIGENOUS SPECIES .)3 ON NATURAL COMMUNITIES 3AX ET AL
4HE POWER OF THIS VIEW OF .)3 IS THAT SYSTEMS ARE INADVERTENTLY BEING PERTURBED
ALL OVER THE GLOBE 7HAT WE CAN LEARN FROM THE PERTURBATIONS WILL NOT ONLY
ADVANCE THE SCIENCE OF ECOLOGY BUT ALSO WILL ALLOW ECOLOGISTS TO RESTORE AND REBUILD
THESE VERY SYSTEMS (OWEVER NOT ALL DISRUPTIONS OF A SYSTEM ARE NECESSARILY INFOR
MATIVE &URTHER IF THE SYSTEM AS IT NORMALLY FUNCTIONS HOLDS SOME INTRINSIC VALUE
-7 #ADOTTE ET AL (EDS) #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY n
3PRINGER 0RINTED IN THE .ETHERLANDS
3- -C-AHON -7 #ADOTTE AND 4 &UKAMI
AS DO INVADED ECOLOGICAL SYSTEMS IT IS IN THE INTEREST OF THE RESEARCHER TO WORK TO
BOTH UNDERSTAND INVASIONS BUT ALSO TO CORRECT OR MITIGATE THE DAMAGING IMPACT OF
NON NATIVE POPULATIONS
.)3 POSE A WIDE VARIETY OF THREATS TO THEIR NON NATIVE HABITATS THROUGH IMMEDI
ATE AND LONG TERM INmUENCES ON SPECIES COMPOSITION AND ECOSYSTEM FUNCTIONING
0ARKER ET AL "ECAUSE THE NEGATIVE IMPACTS OF .)3 ARE lRST DIRECTLY AND
PRIMARILY ECOLOGICAL ECOLOGISTS AND EVOLUTIONARY BIOLOGISTS CONSTITUTE THE FOR
WARD LINE IN MEETING THE CHALLENGES POSED BY THE INTRODUCTION AND SPREAD OF .)3
4HE MANY PATHWAYS THROUGH WHICH AN INTRODUCED POPULATION CAN AFFECT NON
NATIVE HABITATS AND COMMUNITIES HOWEVER ARE DIFlCULT TO DETERMINE AND PREDICT
7HICH TOOLS CAN THE ECOLOGIST APPLY TO THE PROBLEM OF .)3 !NY SOLUTION MUST
DEVELOP FROM A SOLID UNDERSTANDING OF THE FOLLOWING HOW POPULATIONS INTRIN
SICALLY BEHAVE HOW POPULATIONS INmUENCE THE BIOTIC AND ABIOTIC SYSTEMS OF
WHICH THEY ARE COMPONENTS HOW CONVERSELY AN ENVIRONMENT CAN INmUENCE
A NON INDIGENOUS POPULATION AND HOW .)3 POPULATIONS CAN CHANGE PHENO
TYPICALLY AND GENOTYPICALLY IN NEW ENVIRONMENTS !LTHOUGH ALL OF THESE QUESTIONS
FALL SOLIDLY WITHIN THE TRADITION OF ECOLOGICAL RESEARCH THEY DEMAND WITH A
STARTLING IMMEDIACY CAPABLE ANSWERS CULLED FROM UNRESOLVED AMBIVALENT AND
EVEN CONTENTIOUS ECOLOGICAL DEBATE 7HILE ECOLOGISTS ARE AT ONCE WELL POSITIONED
TO ADDRESS THE KEY ELEMENTS THAT HAVE ARISEN FROM THE IMPORTANCE OF .)3 OUR
ANSWERS ARE LIKELY TO CARRY WITH THEM SOME OF THE STRUGGLES BORN OF CLASSIC THEORET
ICAL AND CONCEPTUAL DEBATES IN ECOLOGY AND EVOLUTIONARY BIOLOGY (ERE HOWEVER
THE STUDY OF .)3 MAY NOT NECESSARILY SUFFER FOR THESE CHALLENGES
4HE STUDY OF .)3 WHEN APPROACHED FROM THE REALM OF CONCEPTUAL ECOLOGY
CAN ACT AS INTERESTING TESTS OF COMPETING ECOLOGICAL THEORIES 4HIS POTENTIAL HAS
LONG BEEN RECOGNIZED AND MORE RECENTLY BEEN PROMOTED ,ODGE $AVIS
ET AL 9ET NOT ALL ECOLOGICAL THEORY IS APPLICABLE OR APPROPRIATE FOR THIS TYPE
OF TEST .OR WILL THE ANSWERS GAINED FROM ALL APPLICATIONS OF ECOLOGICAL THEORY
ASSIST IN IMPROVING OUR UNDERSTANDING INTERVENTION OR PREDICTION ABOUT ACTUAL
COLONIZATIONS OF .)3 /NE OF THE KEY GOALS OF THIS BOOK IS TO BETTER RElNE WHAT
ECOLOGISTS BELIEVE THE STUDY OF .)3 CAN DO FOR THE lELD OF ECOLOGY AND CONVERSELY
HOW CONCEPTUAL ECOLOGY CAN ADVANCE OUR ABILITY TO EXPLAIN AND ADDRESS THE
CHALLENGES POSED BY .)3 )N THIS CHAPTER WE WILL ORGANIZE SOME OF THE WAYS IN
WHICH THE STUDY OF .)3 CAN ADVANCE CONCEPTUAL ECOLOGY AND VISE VERSA FOCUSING
ON RECENT LITERATURE IN THE lELD AND THE WAYS IN WHICH THE CHAPTERS OF THIS BOOK
lT INTO THAT SCHEME 4HIS CHAPTER WILL ALSO DISCUSS HOW THIS BOOK ESTABLISHES
GUIDELINES NOT ONLY FOR WHAT WE MAY LEARN FROM .)3 BUT FOR WHAT WE ARE UNLIKELY
TO LEARN !GAIN WITH IMMINENT CHALLENGES AND LIMITED RESOURCES UNDERSTANDING
WHICH SCIENTIlC APPROACHES THEORETICAL AND APPLIED ARE INTRACTABLE OR IMPRACTI
CAL IS AS IMPORTANT TO ADVANCING THIS lELD AS KNOWING WHICH APPROACHES SHOW
PROMISE WHEN CONFRONTING THIS COMPLEX PROBLEM
4RACKING THE TRACTABLE
42!#4!",% ).42!#4!",% !.$ 02!#4)#!, 345$)%3
/& ")/,/')#!, ).6!3)/.3
!LTHOUGH STUDIES ON THE DYNAMICS OF INVASIVE ORGANISMS DEMONSTRATE MANY
RESEARCH METHODOLOGIES THREE APPROACHES ARE CRUCIAL TO THE ADVANCEMENT OF OUR
UNDERSTANDING OF INVASIONS THEY ARE MATHEMATICAL MODELS EXPERIMENTS EITHER
MANIPULATIVE OR OBSERVATIONAL AND REVIEWS OR META
ANALYSES 4HE STRENGTH
DERIVED FROM EXPLORING INVASION DYNAMICS THROUGH MATHEMATICAL MODELING IS
THAT SCIENTISTS WITH EXPERTISE IN A VARIETY OF THEORETICAL APPLICATIONS CAN TACKLE
THE CHALLENGES OF UNDERSTANDING BIOLOGICAL INVASIONS THROUGH NOVEL TECHNIQUES
3HEA AND #HESSON -ODELING SUGGESTS BOTH POSSIBLE DYNAMICS PROB
ABLE lNDINGS AND PERHAPS MORE IMPORTANTLY THE LIMITS TO CERTAIN CONCEPTUAL
APPROACHES TO INVASIONS (ARDING ET AL #HAPTER )T IS BROADLY ACKNOWLEDGED
THAT IN ORDER TO BETTER UNDERSTAND INVASIONS EXPERIMENTAL APPROACHES ARE CRUCIAL
4HERE HAS BEEN A GROWTH OF STRONG EXPERIMENTS IN INVASIONS RANGING FROM LABO
RATORY MICROCOSMS $ICK AND 0LATVOET TO lELD MESOCOSMS $UKES
TO LARGE
SCALE MANIPULATIVE lELD EXPERIMENTS 4ILMAN -ANY OF THE
STRENGTHS OF INFERENCES DERIVED FROM THESE STUDIES IN ORDER TO BE BROADLY APPRECI
ATED NEED TO BE REINCORPORATED INTO MATHEMATICAL MODELS &INALLY META
ANALYSES
AND REVIEWS PROVIDE A GENERALIZED ASSESSMENT OF THE THEORETICAL AND EMPIRICAL
WORK THAT HAS COLLECTED OVER SEVERAL YEARS 4HE SYNTHESIS DERIVED FROM THESE
ANALYSES CAN AS WITH THE MODELS SERVE TO INFORM DIRECT OR DELIMIT THE QUESTIONS
RESEARCHERS ASK ABOUT INVASIONS #ADOTTE ET AL 4HE DIALECTIC DERIVED FROM
THE COMPLEMENTARY APPROACHES OF SUPPOSITION EXPERIMENTATION AND ASSESSMENT
OFFERS THE MOST EFlCIENT WAY TO STUDY INVASIONS AND DRAW BROADER UNDERSTANDING OF
ECOLOGICAL PROCESSES THROUGH THE STUDY OF INVASIONS !LL THREE OF THESE APPROACHES
ARE FEATURED IN THIS BOOK AND IN THE FOLLOWING INTRODUCTION TO THE THEMES OF
THIS BOOK WE PAY SPECIAL ATTENTION TO HOW THIS DIALECTIC CAN GUIDE IMPORTANT
RESEARCH AND THINKING ON INVASIONS
4HE BEHAVIOR OF POPULATIONS
%VEN IN MODELS OF POPULATIONS THAT DO NOT INCLUDE ABIOTIC OR INTERSPECIlC ENVI
RONMENTAL INTERACTIONS INTRASPECIlC POPULATION DYNAMICS CAN ILLUSTRATE IMPOR
TANT PROPERTIES OF INVADING POPULATIONS THROUGH PATTERNS OF INTRINSIC POPULATION
GROWTH AND SPREAD SEE #ADOTTE ET AL #HAPTER !LTHOUGH A NUMBER OF SCALES
MAY ULTIMATELY PROVE IMPORTANT WHEN ASSESSING THE ARRIVAL ESTABLISHMENT
GROWTH AND IMPACT OF AN .)3 ,LORET ET AL (AMILTON ET AL THE POPU
LATION SCALE HAS LONG PROVIDED SOME OF THE CLEAREST INSIGHTS INTO THE BASIC QUESTIONS
OF INVASION DYNAMICS 0ETROVSKII ET AL 4HOMSON !LL OF THE FEATURES
THAT lGURE PROMINENTLY IN POPULATION BIOLOGY FROM THE ROLE OF CARRYING CAPACITY
TO MORTALITY AND REPRODUCTION PROVIDE A BASELINE UNDERSTANDING OF HOW AN .)3
CAN BE EXPECTED TO BEHAVE WHEN COLONIZING A NEW AREA ! KEY DISTINCTION WHEN
EXPLORING THESE INTRINSIC PROPERTIES LIES IN THE DIFFERENT EXPECTATIONS OF STOCHASTIC
3- -C-AHON -7 #ADOTTE AND 4 &UKAMI
VERSUS DETERMINISTIC PROCESSES 1UITE A FEW CHAPTERS IN THIS BOOK IMPLICITLY AND
EXPLICITLY ADDRESS THIS IMPORTANT DISTINCTION
"UCKLEY AND -ETCALF #HAPTER REVIEW THE ROLE OF DENSITY DEPENDENCE IN DETER
MINING THE POPULATION DENSITIES THAT CAN BE REACHED BY AN INVASIVE POPULATION
AND THE INmUENCES OF THOSE DENSITIES ON INVASIVE SUCCESS 4HEY SHOW HOW DENSITY
DEPENDENCE CAN DELIMIT DEMOGRAPHIC PARAMETERS SUCH AS INTRINSIC RATE OF INCREASE
AND DISPERSAL ABILITY 5SING MICROBIAL MICROCOSMS 7ARREN ET AL #HAPTER
SHOW THAT CHANCE EVENTS PLAY A KEY ROLE IN THE ESTABLISHMENT OF AN INVADER )NITIAL
DEMOGRAPHIC STOCHASTICITY PROPAGULE PRESSURE AND THE INITIAL RATE OF INCREASE ALL
CONTRIBUTE TO WHETHER A POPULATION IS ABLE TO ESTABLISH IN A COMMUNITY BUT NOT
NECESSARILY PERSIST IN THAT COMMUNITY 3PECIES
SPECIlC EFFECTS WERE ALSO FOUND TO
BE IMPORTANT IN BOTH ESTABLISHMENT AND PERSISTENCE WHICH INDICATES THAT CARE
FUL CHARACTERIZATION OF THE DEMOGRAPHIC QUALITIES OF AN INVADING SPECIES MAY BE
IMPORTANT TO UNDERSTANDING ITS POTENTIAL IMPACT
7HAT POPULATION
SCALE PATTERNS DO LEAD TO PERSISTENCE OVER TIME -URREL
#HAPTER TAKES A @PLANT
EYE VIEW IN USING SIMULATIONS TO SHOW HOW THE DENSITY
AND AGGREGATION OF A POPULATION CAN CREATE A @NEIGHBORHOOD EFFECT THAT DETER
MINES ULTIMATE POPULATION GROWTH 4HIS APPROACH COULD BE QUITE IMPORTANT IN
PLANT INVASIONS AS INTRA
SPECIlC COMPETITION CAN DETERMINE ULTIMATE VIABILITY AND
SPREAD OF A POPULATION !FTER THE POPULATION IS ESTABLISHED ITS DISPERSAL BECOMES
A CRUCIAL ISSUE IN ANTICIPATING THE EXTENT AND IMPACT OF AN INVASION 0UTH AND
0OST ,EWIS ET AL #HAPTER DEMONSTRATE A NOVEL APPROACH TO MODELING
DISPERSAL THROUGH TWO DIMENSIONS 5SING KERNEL METHODS TO INCORPORATE LONG
DIS
TANCE DISPERSAL INTO MODELS AND TWO
DIMENSIONAL MODELS TO CORRECT FOR BIASES IN
MORE COMMONLY USED ONE
DIMENSIONAL MODELS A MORE ACCURATE PREDICTION OF THE
DISPERSAL BEHAVIOR OF AN INVADING POPULATION CAN BE MADE
-ETAPOPULATION THEORY ALSO EMPHASIZES THE ROLE OF DISPERSAL IN MAINTAINING
.)3 PERSISTENCE OVER TIME (ARDING ET AL #HAPTER SHOW THAT FOR SINGLE .)3
COLONIZING PATCHY ENVIRONMENTS COLONIZATION SUCCESS INTO EMPTY PATCHES MUST
BE GREATER THAN EXTINCTIONS 4HIS IMPLIES THAT SPECIES WITH STRONG DISPERSAL MAY BE
PREDISPOSED TO COLONIZE #OMBINED WITH BUFFERING LIFE HISTORY TRAITS SUCH AS PLANTS
THAT CAN SELF
POLLINATE OR VEGETATIVELY REPRODUCE A GOOD DISPERSAL MECHANISM CAN
LEAD TO SUCCESSFUL COLONIZATION OF A HETEROGENEOUS LANDSCAPE
%NVIRONMENT INFLUENCING .)3 POPULATIONS
)NTRINSIC POPULATION BEHAVIORS ARE CLEARLY CRUCIAL TO ANY UNDERSTANDING OF THE
ESTABLISHMENT AND SPREAD OF .)3 BUT AS WITH MANY ECOLOGICAL STUDIES OF POPU
LATIONS THE BEHAVIORS OF THE POPULATION CAN DEPEND ON THE BIOTIC AND ABIOTIC
COMPONENTS OF THE ENVIRONMENT ,ODGE $UKES AND -OONEY (OLWAY
ET AL )NCORPORATING THE ADDITIONAL LEVEL OF COMMUNITY COMPLEXITY TO POPU
LATION DYNAMICS HOWEVER CAN BE DAUNTING WHEN DETERMINING THE IMPACT OF .)3
$AVIS ET AL 4HE STUDY OF THE ESTABLISHMENT AND SPREAD OF .)3 CAN PROVIDE
INSIGHT INTO HOW THESE MORE COMPLEX RELATIONSHIPS DEVELOP LENDING IMPORTANT
4RACKING THE TRACTABLE
CASE STUDIES TO THE ANALYSIS OF ECOLOGICAL COMMUNITIES 4HE COMPLEX AND COMPLICATED
DYNAMICS OF THE ENVIRONMENT ALSO MEAN THAT THESE ADDITIONAL COMPONENTS CAN MOVE
PROBLEMS FROM LESS REALISTIC BUT TRACTABLE PROBLEMS TO INTRACTABLE ONES 4HEREFORE IT
IS IMPORTANT TO USE STUDIES OF ENVIRONMENTAL INmUENCES ON .)3 TO DETERMINE WHICH
APPROACHES MAY BE APPROPRIATE AND WHICH AN INEFlCIENT USE OF TIME AND RESOURCES
! CLASSIC CONCEPT OF HOW AN ENVIRONMENT CAN DICTATE WHETHER OR NOT A SPE
CIES CAN PERSIST IS THAT OF THE ECOLOGICAL NICHE 4HE IDEA THAT AN ENVIRONMENT IS
PARSED INTO A lNITE SET OF NICHES HAS BEEN AN IMPORTANT ONE FOR THE DEVELOPMENT
OF INVASIONS BIOLOGY #ADOTTE #HAPTER 6AZQUEZ #HAPTER USES A SERIES OF
META
ANALYSES TO EXPLORE HOW NICHE BREADTH AND INVASION SUCCESS RELATE 7HEN
LOOKING AT FUNDAMENTAL NICHES WHERE SPECIES REQUIREMENTS ARE INDEPENDENT
OF COMMUNITY INTERACTIONS VERSUS REALIZED NICHES WHERE THE SUITABILITY OF AN
ENVIRONMENT TAKES INTO ACCOUNT COMPETITION THE REALIZED NICHE ULTIMATELY DETER
MINES THE SUCCESS OF AN .)3 6AZQUEZ POINTS OUT THAT REALIZED NICHES HOWEVER
ARE IMPOSSIBLE TO MEASURE AND WARNS AGAINST PURSUING STRAIGHTFORWARD INVESTIGA
TION INTO NICHE
BASED PREDICTIONS OF POTENTIAL .)3 SUCCESS 4HIS MARKS A POWERFUL
RECOMMENDATION FOR INVASIONS BIOLOGY AS A TRADITIONAL LINE OF INQUIRY INTO PREDICT
ING THE ESTABLISHMENT OF .)3 HAS BEEN THE SUSCEPTIBILITY OF DIFFERENT HABITATS TO
INVASIONS (OLWAY ,EVINE AND $!NTONIO
)N OTHER CIRCUMSTANCES INVASIONS CAN SHED LIGHT ON CONCEPTUAL TOPICS THAT ARE
DIFlCULT TO ASSESS IN NATURAL COMMUNITIES $UNCAN AND &ORSYTH #HAPTER USE
HISTORICAL RECORDS OF BIRD INVASIONS TO TEST CLASSIC HYPOTHESES OF COMPETITION AND
COMMUNITY STRUCTURE )NVASION RECORDS OFFER A UNIQUE WAY OF DISCOVERING IF INVA
SIONS OF BIRD SPECIES INTO CERTAIN TYPES OF ENVIRONMENTS ARE MORE LIKELY TO PERSIST
4HEY lND THAT PRIORITY EFFECTS DO OCCUR AND THAT DEPENDING ON CIRCUMSTANCES STO
CHASTIC PROCESSES AND DETERMINISTIC PROCESSES SUCH AS COMPETITION CAN DETERMINE
ULTIMATE COMMUNITY COMPOSITION 3AX AND 'AINES #HAPTER SUGGEST THAT SPE
CIES RICH COMMUNITIES ARE LIKELY TO BE MORE SUSCEPTIBLE TO INVASIONS BECAUSE CON
DITIONS ARE FAVORABLE FOR MANY SPECIES 4HEY ARGUE HOWEVER THAT THIS IS TRUE ONLY
UP TO A CRITICAL THRESHOLD SUCH AS HIGHLY SPECIOSE TROPICAL RAIN FORESTS AT WHICH
POINT %LTONS %LTON MODEL IN WHICH SPECIES RICH COMMUNITIES HAVE LESS
AVAILABLE NICHE SPACE UNEXPLOITED BY THE BEST COMPETITOR BECOME APPLICABLE "OTH
3AX AND 'AINES #HAPTER AND 3MITH AND 3HURIN #HAPTER EXPLAIN THAT
HETEROGENEITY WITHIN REGIONAL SCALES CAN MEAN THAT MULTIPLE PROCESSES STOCHASTIC
AND DETERMINISTIC AND MULTIPLE OUTCOMES INVASION RESISTANCE AND SUSCEPTIBILITY
CAN BE FOUND IN ANY ONE HABITAT 4HIS INDICATES THAT EVEN IN UNSATURATED COM
MUNITIES WHERE PRESUMABLY THERE IS AMPLE NICHE SPACE FOR WELL
DISPERSED INVAD
ERS INVASION SUCCESS IS CONTINGENT ON OTHER FACTORS 4HIS ROLE OF CONTINGENCY AND
COMPLEXITY IS BECOMING A COMMON THEME IN BOTH INVASIONS LITERATURE AND THE
EMPIRICAL STUDIES OF COMMUNITY COMPOSITION AND ASSEMBLY AND THEREFORE POINTS
TO AN IMPORTANT AREA OF RECIPROCAL RESEARCH
!NOTHER EXAMPLE OF THE CHALLENGES INHERENT IN COMPLEX COMMUNITY DYNAMICS
IS ILLUSTRATED IN (ARDING ET AL #HAPTER !LTHOUGH AS MENTIONED ABOVE THEY
SHOW THAT METAPOPULATIONS WITHOUT COMPETITION RELY ON A SIMPLE STRATEGY OF
3- -C-AHON -7 #ADOTTE AND 4 &UKAMI
COLONIZATION AND BUFFERS FROM EXTINCTION ADDING EVEN ONE MORE SPECIES INTO THE
MODEL CAN RADICALLY CHANGE THE NUMBER OF POPULATION PARAMETERS AND LIFE HISTORY
TRAITS THAT MAY ULTIMATELY DICTATE THE SUCCESSFUL ESTABLISHMENT AND PERSISTENCE
OF AN .)3 4HIS COMPLEXITY OF POTENTIAL OUTCOMES INDICATES THAT METAPOPULATION
THEORY AS APPLIED TO MULTIPLE SPECIES LIVING IN SUBDIVIDED HABITATS MIGHT BE WELL
APPLIED TO CASE STUDIES EXPERIMENTS AND MANAGEMENT PROBLEMS THAT OFFER SPECIFIC
SPECIES CHARACTERISTICS AND ROBUST ESTIMATION OF POPULATION PARAMETERS 7ITH SUCH A
META ANALYSIS IT MAY BE POSSIBLE TO DETERMINE WHETHER A METAPOPULATION MODEL
CAN ACCURATELY DESCRIBE AND PREDICT CERTAIN TYPES OF INVASIONS
5SING MICROBIAL MICROCOSMS 7ARREN ET AL #HAPTER REINFORCE THIS CONCLU
SION )N A SERIES OF AMBITIOUS EXPERIMENTS WHERE DIFFERENT SPECIES ARE INTRODUCED
INTO ESTABLISHED COMMUNITIES THEY SHOW THAT PREDICTING INVASIONS CAN DEPEND
ON A NUMBER OF CHARACTERISTICS OF THE SPECIES INVOLVED AS WELL AS THE COMMUNITY
CHARACTERISTICS 3PECIlCALLY THE INTRINSIC RATE OF INCREASE A SIMPLE POPULATION
PARAMETER SHOULD BE THOUGHT OF AS HAVING AS MUCH TO DO WITH THE INTERACTION
CONTACT EXPERIENCE BETWEEN AN INVADER AND A COMMUNITY AS INTRINSIC PROPERTIES
OF THE INVADING POPULATION
$EMOGRAPHIC PARAMETERS LIKE THE INTRINSIC RATE OF INCREASE THEMSELVES ARE
NOT CONSTANT 4HESE PARAMETERS CAN CHANGE STOCHASTICALLY DUE TO BOTH INTRINSIC
DEMOGRAPHIC FACTORS SUCH AS RANDOM CHANGES IN BIRTH OR DEATH RATES 4HEY CAN
ALSO CHANGE DUE TO ENVIRONMENTAL STOCHASTICITY &RECKLETON ET AL #HAPTER
DISTINGUISH DEMOGRAPHIC VERSUS ENVIRONMENTAL STOCHASTICITY AT DIFFERENT STAGES OF
AN INVASION ARRIVAL ESTABLISHMENT AND SPREAD 4HEY SHOW WITH MATHEMATICAL
MODELS THAT STOCHASTIC PROCESSES INmUENCE THESE STAGES OF THE INVASION IN DIFFERENT
WAYS AND WITH DIFFERENT CONSEQUENCES 3TOCHASTICITY MAY BE MORE IMPORTANT IN
PHASES WITH LOW DENSITIES ARRIVAL AND ESTABLISHMENT WHILE DENSITY DEPENDENT
BEHAVIORS MAY HOLD A GREATER INmUENCE ON THE SPREAD OF ALREADY ESTABLISHED POPU
LATIONS !GAIN MODELS LIKE THIS CAN INDICATE WHEN AND HOW TO PROCEED WITH AN
EXPERIMENTAL PROGRAM TO TEST STOCHASTIC AND DETERMINISTIC COMPONENTS OF INVA
SION SUCCESS
!LTHOUGH DIRECT AGONISTIC INTERACTIONS BETWEEN SPECIES SUCH AS COMPETITION
AND PREDATION HAVE LONG BEEN HELD AS CENTRAL TO UNDERSTANDING THE POPULATION
STRUCTURE OF COMMUNITIES INDIRECT EFFECTS AND MUTUALISMS HAVE RECENTLY BEEN
THRUST INTO THE FOREFRONT OF THINKING ON A NUMBER OF CONCEPTUAL THEMES IN ECOLOGY
7OOTTON ,ORTIE ET AL SUCH AS RICHNESS PRODUCTIVITY POPULATION
VIABILITY ANALYSIS AND INVASIONS ,ACH 6AZQUEZ #HAPTER SHOWS IN
HIS META ANALYSES OF NICHE BREATH AND INVASIONS THAT THE PRESENCE OF MUTUALISTS
AND OTHER NICHE CATEGORIES THAT MAY BE TRICKY TO MEASURE CAN HAVE AN IMPORTANT
INmUENCE ON THE ULTIMATE VIABILITY OF A POPULATION IN A NEW HABITAT 4HORPE AND
#ALLAWAY #HAPTER EXTEND THE DISCUSSION OF MUTUALISMS TO THE LEVEL OF INDI
RECT MUTUALIST INTERACTIONS AND FEEDBACK LOOPS 'OOD EXPERIMENTAL EVIDENCE HAS
SHOWN THAT PRESENCE OF SOIL PATHOGENS DECREASE .)3 PLANT SUCCESS IN NATIVE RANGE
AND ABSENCE OF THOSE PATHOGENS INCREASES .)3 SUCCESS IN NEW RANGE &URTHER
MUTUALIST INTERACTIONS CAN MORE EASILY CREATE POSITIVE FEEDBACKS IN A NEW RANGE
4RACKING THE TRACTABLE
ENHANCING PLANT SUCCESS 4HIS CONCEPTUAL APPROACH TO INVASIONS MIGHT BE WELL
SUITED FOR A MATHEMATICAL STUDY AS FEEDBACKS ARE BOTH COMMON IN NATURE YET DIF
lCULT TO TEASE APART FROM OTHER COMMUNITY EFFECTS
)N ADDITION TO USING ANALYSES OF ENVIRONMENTAL CONTEXTS TO PREDICT WHETHER AN
.)3 WILL SUCCESSFULLY ESTABLISH AND PERSIST IN A NOVEL ENVIRONMENT 'ILBERT AND
0ARKER #HAPTER AND 3ATAKE ET AL #HAPTER ADDRESS HOW THE ENVIRONMENT
CAN PREVENT AN INVASION OR EVEN REVERSE AN ALREADY PERSISTENT .)3 'ILBERT AND
0ARKER ARGUE THAT THE UNDERSTUDIED ROLE THAT PATHOGENS PLAY IN PLANT POPULATION
REGULATION CAN HAVE IMPORTANT CONSEQUENCES IN UNDERSTANDING PREDICTING AND
MITIGATING OR REVERSING HIGH
DENSITY PERSISTENCE OF .)3 3ATAKE ET AL #HAPTER
MODEL @CLASSIC MAST SEEDING AND SHOW THAT WHEN SEED SET IS NEGATIVELY CORRELATED
IN TIME BUT POSITIVELY CORRELATED ACROSS SPACE A SYSTEM IS MOST RESISTANT TO INVAD
ING SEED PREDATORS
0HENOTYPIC AND GENOTYPIC CHANGE
! NEW AND EXCITING THEME IN INVASIONS BIOLOGY ADDRESSES HOW POPULATIONS OF .)3
MAY CHANGE IN PHENOTYPIC EXPRESSION 3CHWEITZER AND ,ARSON OR GENOTYPE
3AKAI ET AL DUE TO THE NOVEL ENVIRONMENTS THEY EXPERIENCE IN THEIR NEW
RANGE THIS INCLUDES POPULATION CHANGES INHERENT IN A COLONIZING EVENT SUCH AS
FOUNDER EFFECTS OR DRIFT 0ALUMBI #OX *UST AS MUTATIONS IN A VIRUS
WITHIN HUMAN POPULATIONS CAN MEAN THE DIFFERENCE BETWEEN RARITY AND A PAN
DEMIC SO TOO CHANGES IN .)3 POPULATIONS CAN TURN A RARE OR ISOLATED POPULATION
INTO A PEST 4HIS CAN ALSO BE ONE OF THE MAJOR FACTORS IN CREATING CREATE TIME LAGS
BETWEEN THE ARRIVAL PHASE OF AN .)3 AND AN OUTBREAK -ACK ET AL "YERS AND
'OLDWASSER -EMMOTT ET AL .EW METHODS IN GENETICS LIFE
HISTORY
EVOLUTION AND COEVOLUTIONARY THEORY CAN PROVIDE INSIGHT INTO AND DRAW INFERENCE
FROM THE GROWING EXAMPLES OF .)3 POPULATIONS THAT HAVE CHANGED FUNDAMENTALLY
SINCE THEIR ARRIVAL IN A NEW RANGE
%VOLUTIONARY CHANGE IN .)3 POPULATIONS CAN FOLLOW MANY POSSIBLE PATHWAYS
3CHIERENBECK AND !NOUCHE #HAPTER EMPHASIZE THAT PRE
ADAPTED GENERALIST
GENOTYPES WILL FACILITATE INVASION SUCCESS 3UBSTANTIAL GENETIC VARIATION CAN PRO
VIDE THE REQUISITE GENETIC MATERIAL FOR RAPID EVOLUTIONARY RESPONSE TO THE NOVEL
ENVIRONMENT AND 3HIERENBECK AND !NOUCHE #HAPTER POINT OUT THAT THERE ARE
MANY PATHWAYS TO THIS VARIATION HETEROZYGOSITY POLYPLOIDY AND PERHAPS NOW
EVEN EXTRA
GENOMIC GENETIC INFORMATION ,ALLE ET AL (OWEVER GENERAL
GENETIC DIVERSITY MAY NOT ALONE BE ENOUGH TO INmUENCE INVASION SUCCESS (ERBEN
ET AL -ORE IMPORTANTLY HOW LABILE LIFE
HISTORY TRAITS ARE PHENOLOGY
CLONAL OR VEGETATIVE REPRODUCTION IN PLANTS MATING BEHAVIORS ETC MAY MARK THE
IMPORTANT GENETIC DISTINCTION "UCKLEY AND -ETCALF #HAPTER DESCRIBE HOW
LIFE
HISTORY EVOLUTION DUE TO UNIQUE FORCES AT WORK IN NOVEL ENVIRONMENTS DIFFER
ENT LIFE
HISTORY STRATEGIES MAY BENElT .)3 POPULATIONS
7HILE LIFE
HISTORY EVOLUTION MAY BE RESPONSIBLE FOR RARE LARGE OUTBREAKS THIS
WOULD BE DIFlCULT TO TEST 4HORPE AND #ALLAWAY #HAPTER EXPLAIN THAT
3- -C-AHON -7 #ADOTTE AND 4 &UKAMI
EVOLUTIONARY RESPONSE OF MICROBIAL COMMUNITIES CAN HAPPEN SO FAST THAT THIS MAY
MUTE OR MITIGATE AN .)3 FROM LONG TERM DOMINANCE THROUGH POSITIVE SOIL FEEDBACK
PATHWAYS "ECAUSE OF A POTENTIALLY RAPID RESPONSE AND THE SHORT GENERATION TIMES
OF MICROBIAL COMMUNITIES IT MAY BE POSSIBLE TO TEST SOIL SYSTEMS FOR RAPID EVOLU
TIONARY RESPONSE TO ENCOURAGING OR MITIGATING THE SUCCESSFUL ESTABLISHMENT OF AN
.)3 .)3 COLONIZATION ADDITIONALLY MAY PROVIDE IDEAL SYSTEMS TO RESEARCH THE
STABILITY OF MICROBIAL COMMUNITIES AS WELL AS THEIR RELATIONSHIP TO THE PLANTS WITH
WHICH THEY INTERACT THROUGH BELOWGROUND PATHWAYS 7ARDLE 0ATHOGENS
WOULD ALSO lT THIS MODEL AND 'ILBERT AND 0ARKER #HAPTER DESCRIBE THE COEVO
LUTION OF PLANT
PATHOGEN SYSTEMS AND MATERNALLY INHERITED INDUCED RESISTANCE
4HE EVOLUTIONARY RESPONSE OF AN .)3 TO THE BIOTIC COMMUNITY MAY BE CRUCIAL NOT
ONLY IN THE ESTABLISHMENT BUT IN THE RESULTING EFFECT ON THE NATIVE BIOTA +ONDOH
#HAPTER MODELS HOW CONTACT EXPERIENCE THE PREVIOUS EVOLUTIONARY lLTERING
FROM PRIOR INTERACTIONS BETWEEN AN .)3 AND RELATED NOVEL COMPETITORS PREDATORS
AND PREY CAN RESULT IN SUCCESSFUL INVASION (E DESCRIBES HOW THE CONTACT EXPERI
ENCE IN MANY FORMS CAN INmUENCE THE SPECIlC COMPONENTS OF CONTACT BETWEEN
THE .)3 AND NOVEL BIOTA AS INFORMATION RETRIEVAL DETECTING THE OTHER SPECIES
PROCESSING RECOGNIZING CHARACTERISTICS OF THAT NOVEL SPECIES AND BEHAVIORAL
RESPONSE BEING ABLE TO PREY UPON ESCAPE OR COMPETE WITH THAT SPECIES
.)3 POPULATIONS INFLUENCING POPULATIONS AND ENVIRONMENT
)T SHOULD BE NOTED THAT EVEN IDENTIFYING AN INVASIVE SPECIES CAN BE PROBLEM
ATIC #OLAUTTI AND -AC)SAAC -URPHY ET AL #HAPTER "UT REGARDLESS OF
DElNITION THERE IS A GENERAL UNDERSTANDING AMONG BIOLOGISTS THAT ULTIMATELY ANY
INVASIVE WILL IN SOME KEY WAY BE DElNED BY ITS INmUENCE ON THE BIOTIC AND ABIOTIC
COMPONENTS OF ITS NEW RANGE 4HERE MAY BE MILLIONS OF .)3 PROPAGULES ENTERING
COMPLEX ENVIRONMENTS BUT OUR CONCERN FOR THEIR RANGE EXPANSION IS ULTIMATELY
A FUNCTION OF THEIR IMPACT ON OTHER SPECIES IN THEIR NEW RANGE THE WAYS IN WHICH
THEY AFFECT ECOSYSTEM PROCESSES AND HOW THESE CHANGES MAY PERSIST OR INCREASE
OVER TIME
(ISTORICALLY DOCUMENTED INVASIONS CAN ALSO PROVIDE INSIGHTS INTO THE PATH
WAYS THROUGH WHICH THE CONTEXT OF AND INVADING ORGANISMS CAN INmUENCE BOTH
PERSISTENCE AND THE CONCOMITANT INmUENCE ON NATIVE BIOTA #OURCHAMP AND #AUT
#HAPTER USED MODELS INCORPORATING DIRECT AND INDIRECT INTERACTIONS TO EXPLAIN
INVASIONS AS WELL AS SHOW HOW THE INVASIONS THEMSELVES ILLUSTRATE THE COMPLEX
AND SOMETIMES COUNTERINTUITIVE BEHAVIORS OF INTERACTING POPULATIONS #OMPLEX
DYNAMICS BETWEEN POPULATIONS MAY ALSO MEAN THAT AFTER AN INVASION IT TAKES
MANY GENERATIONS FOR THE EFFECTS OF THE NEW INTERACTIONS TO SHOW THEIR IMPACT
&RECKLETON ET AL #HAPTER
!LTHOUGH DOCUMENTED CHANGES IN NATIVE BIOTA OR ECOSYSTEM FUNCTION MAY
BE CLEAR AFTER AN INVASIVE BECOMES A DAMAGING PEST PREDICTING THE ULTIMATE
IMPACT OF AN .)3 ON NATIVE HABITATS IS DIFlCULT ! NUMBER OF FACTORS INTRINSIC TO
THE INVADING POPULATION AND DUE TO THE NOVEL ENVIRONMENT CAN ELICIT COMPLEX
4RACKING THE TRACTABLE
RESPONSES TO THE NON
NATIVE POPULATIONS )NDIRECT EFFECTS ON THE ENVIRONMENT ARE
DIFlCULT TO DETECT LET ALONE PREDICT YET MAY BE AMONG THE MORE COMMON AND SERI
OUS RESPONSES TO .)3 )NVASIVE PLANTS CAN ALTER THE SOIL NUTRIENT FEEDBACK LOOPS
AND RENDER LONG TERM CHANGES TO THE SOIL ENVIRONMENT 4HORPE AND #ALLAWAY
#HAPTER -UTUALISMS SUCH AS POLLINATION SERVICES CAN BE DISRUPTED BY INVA
SIVES 4HOUGH THE CONVERSE MAY OCCUR .ON
NATIVE BEES HAVE BEEN SHOWN TO
INCREASE POLLINATION EFlCIENCY OF SOME PLANTS 2ICHARDSON ET AL ALTHOUGH
THE EFFECT ON NATIVE POLLINATOR COMMUNITIES ADDS ADDITIONAL UNCERTAINTY TO THE
DIAGNOSIS OF THESE EFFECTS 'ENERALISM VERSUS SPECIALISM CAN BE CRUCIAL TO THE
EXTENT OF INmUENCE A NON
NATIVE SPECIES MAY HAVE IN PATHOGEN
HOST RELATIONSHIPS
'ILBERT AND 0ARKER #HAPTER POLLINATOR SYSTEMS 6AZQUEZ #HAPTER AND
SOIL MUTUALISMS 4HORPE AND #ALLAWAY #HAPTER 4HE SYMMETRY OF THESE MUTU
ALIST RELATIONSHIPS IS OF FURTHER CONSEQUENCE 6AZQUEZ #HAPTER WHEN INFERRING
THE POTENTIAL IMPACT AN .)3 MAY HAVE ON NOVEL SYSTEMS
7HETHER DIRECT OR INDIRECT THE STOCHASTIC NATURE OF POPULATION INTERACTIONS
CAN RADICALLY AFFECT THE ULTIMATE CONSEQUENCES OF A PERSISTENT .)3 &RECKLETON
et al #HAPTER 7ARREN ET AL #HAPTER ESPECIALLY AS SYSTEMS MAY CONTAIN NON
LINEAR DYNAMICS ! lNAL LEVEL OF COMPLEXITY RESULTS FROM THE MANY APPLICATIONS OF
OUR INFERENCE TO POLITICAL SOCIAL AND ECONOMIC ISSUES )T IS BROADLY ACCEPTED THAT
.)3 CAN AFFECT THE INHERENT RICHNESS OF NATIVE BIOTA SPECIlC ENDANGERED SPECIES
THE ECOSYSTEM FUNCTION OF NATIVE HABITATS AS WELL AS BIOTA OF ECONOMIC IMPOR
TANCE )N THEIR REVIEW OF HOW .)3 INmUENCE NOVEL COMMUNITIES -URPHY ET AL
#HAPTER FOUND A CORRELATION BETWEEN LEVEL OF INVASIVE ON THE 3% SCALE AND COM
MUNITY IMPACT THERE WAS NO CORRELATION WITH ECONOMIC IMPACT 4HIS MAY TROUBLE
OUR ABILITY TO APPLY A LEVEL OF COMMUNITY CHANGE DUE TO AN INVASIVE TO SOME
SYSTEM OF SOCIAL OR ECONOMIC @IMPORTANCE OF THE INVADER
#/.#,53)/.
)NVASIONS BIOLOGY IS FACED WITH MANY OF THE SAME STRUGGLES THAT HAVE LONG FACED
MAINSTREAM ECOLOGY /RGANISMS SHOW AN ARRAY OF INTRINSIC QUALITIES THAT ARE BOTH
INDICATIVE OF THEIR POPULATION DYNAMICS AS WELL AS RESPONSIBLE FOR THOSE DYNAM
ICS 9ET POPULATIONS OF ORGANISMS RESPOND AND INmUENCE THEIR BIOTIC AND ABIOTIC
CONTEXTS AND THEY ADAPT PHENOTYPICALLY AND GENOTYPICALLY TO THEM 4HE PROBLEM
OF lLTERING THAT WHICH IS BROADLY IMPORTANT TO POPULATIONS FROM THAT WHICH IS
IDIOSYNCRATIC IS HIGHLIGHTED IN THE STUDY OF .)3 7E CERTAINLY NEED TO DEVELOP A
GENERAL UNDERSTANDING OF THE QUALITIES OF POPULATIONS AND HABITATS THAT CAN LEAD TO
INVASION PERSISTENCE AND IMPACT YET WE OFTEN lND THAT THE CAUSES OF ANY SPECIlC
INVASION CAN BE UNIQUE TO THAT SPECIlC SPECIES OR HABITAT ! MAJOR GOAL OF THIS
BOOK IS TO CAREFULLY ASSESS WHEN SIMPLICITY IS INFORMATIVE EG A POPULATION
LEVEL
ANALYSIS OF DENSITY DEPENDENCE WHEN COMPLEXITY IS INSTRUMENTAL SUCH AS WHEN
ABIOTIC FACTORS INmUENCE .)3 PERSISTENCE AND WHEN GENERALIZATION IS NOT LIKELY
POSSIBLE SUCH AS THE FACTORS THAT MAKE UP THE REALIZED NICHE OF AN .)3
3- -C-AHON -7 #ADOTTE AND 4 &UKAMI
/NE OF THE GREATEST CHALLENGES FACING THE STUDY OF INVASIONS BIOLOGY ARISES
FROM SPECIES
SPECIlC CHARACTERISTICS !S IS OFTEN SHOWN IN THIS BOOK A BETTER
UNDERSTANDING OF THE INTERFACE BETWEEN INTRINSIC POPULATION PROPERTIES AND THE
STOCHASTIC CONTEXT OF A POPULATION IS LIKELY TO PROVIDE IMPORTANT INSIGHTS INTO .)3
ESTABLISHMENT AND EXPANSION &RUSTRATING MANY STRAIGHTFORWARD PATHS TO INVA
SION PREDICTION IS THE COMBINATION OF STOCHASTIC AND DETERMINISTIC PROCESSES THAT
INmUENCE POPULATIONS BEHAVIORS $ENNIS ET AL -ATHEMATICAL MODELS MAY
HELP INDICATE WHICH PARAMETERS UNDER WHICH ASSUMPTIONS CAN SHOW A DElNI
TIVE INmUENCE ON POPULATION DYNAMICS 4HUS WHEN ENTERING AN OBSERVATIONAL
OR MANIPULATIVE EXPERIMENTAL PROGRAM THE THEORETICAL LITERATURE CAN PROVE AN
IMPORTANT GUIDE TO DEVELOPING HYPOTHESES AND TESTING THEM
2EmECTING RECENT CONCEPTUAL ADVANCES IN GENERAL ECOLOGICAL THEORY OFTEN
IGNORED PROCESSES SUCH AS MUTUALISMS INTRA
SPECIlC COMPETITION AND BELOW
GROUND PROCESSES ARE ALSO LIKELY TO INmUENCE THE POTENTIAL IMPACT OF .)3 #ALLAWAY
AND !SCHEHOUG 2ICHARDSON ET AL 0ARKER AND (AUBENSAK
#ALLAWAY ET AL AND MARK POTENTIAL PATHWAYS TO BRIDGE WHAT APPEARS TO BE
IDIOSYNCRATIC TO MORE GENERALIZABLE PATTERNS )N THESE AREAS THE STUDY OF .)3 MAY
BE WELL SUITED TO INFORM OUR GREATER UNDERSTANDING OF SUCH COMPLEX AND OFTEN
MASKED MECHANISMS IN GENERAL 4HUS NOT ONLY IS THERE A RECIPROCAL INTEREST IN
UNDERSTANDING TRADITIONAL PROBLEMS IN ECOLOGY AND INVASIONS BIOLOGY THESE TWO
APPROACHES TO NATURAL SYSTEMS SHOULD INTERACT AND INFORM EACH OTHER AS THEY
ADVANCE IN TECHNIQUES CONCEPTS AND UNDERSTANDING
2%&%2%.#%3
"YERS * % AND , 'OLDWASSER %XPOSING THE MECHANISM AND TIMING OF IMPACT OF
NONINDIGENOUS SPECIES ON NATIVE SPECIES %COLOGY
#ADOTTE - 7 " -URRAY AND * ,OVETT
$OUST 5SING DATABASE ANALYSES TO TEST WIDE
SPREAD ECOLOGICAL GENERALITIES IN INVASIONS BIOLOGY "IOLOGICAL )NVASIONS IN PRESS
#ALLAWAY 2 - AND % 4 !SCHEHOUG )NVASIVE PLANTS VERSUS THEIR NEW AND OLD
NEIGHBORS ! MECHANISM FOR EXOTIC INVASION 3CIENCE
#ALLAWAY 2 - ' # 4HELEN ! 2ODRIGUEZ AND 7 % (OLBEN 3OIL BIOTA AND EXOTIC
PLANT INVASION .ATURE
#OLAUTTI 2 ) AND ( * -AC)SAAC ! NEUTRAL TERMINOLOGY TO DElNE @INVASIVE SPECIES
$IVERSITY AND $ISTRIBUTIONS
$AVIS - ! * 0 'RIME AND + 4HOMPSON &LUCTUATING RESOURCES IN PLANT COMMUNI
TIES A GENERAL THEORY OF INVASIBILITY *OURNAL OF %COLOGY
$AVIS - ! + 4HOMPSON AND * 0 'RIME #HARLES 3 %LTON AND THE DISSOCIATION OF
INVASION ECOLOGY FROM THE REST OF ECOLOGY $IVERSITY AND $ISTRIBUTIONS
$ENNIS " 2 ! "ESHARNAIS * - #USHING 3 - (ENSON AND 2 & #ONSTANTINO
#AN NOISE INDUCE CHAOS /)+/3
$ICK * 4 ! AND $ 0LATVOET )NVADING PREDATORY CRUSTACEAN $IKEROGAMMARUS
VILLOSUS ELIMINATES BATH NATIVE AND EXOTIC SPECIES 0ROCEEDINGS OF THE 2OYAL 3OCIETY OF
,ONDON 3ERIES "
"IOLOGICAL 3CIENCES
4RACKING THE TRACTABLE
$UKES * 3 3PECIES COMPOSITION AND DIVERSITY AFFECT GRASSLAND SUSCEPTIBILITY AND
RESPONSE TO INVASION %COLOGICAL !PPLICATIONS
$UKES * 3 AND ( ! -OONEY $OES GLOBAL CHANGE INCREASE THE SUCCESS OF BIOLOGICAL
INVADERS 4RENDS IN %COLOGY %VOLUTION
%LTON # 3 4HE ECOLOGY OF INVASIONS BY ANIMALS AND PLANTS -ETHUEN AND #O ,4$
,ONDON 5+
(AMILTON - ! " 2 -URRAY - 7 #ADOTTE ' # -OSE ! # "AKER # * (ARRIS AND
$ ,ICARI ,IFE
HISTORY CORRELATES OF PLANT INVASIVENESS AT REGIONAL AND CONTINEN
TAL SCALES %COLOGY ,ETTERS
(ERBEN 4 " -ANDAK + "IMOVA AND : -UNZBERGOVA )NVASIBILITY AND SPECIES
RICHNESS OF A COMMUNITY ! NEUTRAL MODEL AND A SURVEY OF PUBLISHED DATA %COLOGY
(OLWAY $ ! &ACTORS GOVERNING RATE OF INVASION A NATURAL EXPERIMENT USING
!RGENTINE ANTS /ECOLOGIA
(OLWAY $ ! ! 6 3UAREZ AND 4 * #ASE 2OLE OF ABIOTIC FACTORS IN GOVERNING
SUSCEPTIBILITY TO INVASION ! TEST WITH ARGENTINE ANTS %COLOGY
,ACH , )NVASIVE ANTS 5NWANTED PARTNERS IN ANT
PLANT INTERACTIONS !NNALS OF THE
-ISSOURI "OTANICAL 'ARDEN
,EVINE * - AND # - $!NTONIO %LTON REVISITED ! REVIEW OF EVIDENCE LINKING DIVER
SITY AND INVASIBILITY /)+/3
,LORET & & -EDAIL ' "RUNDU AND 0 % (ULME ,OCAL AND REGIONAL ABUNDANCE
OF EXOTIC PLANT SPECIES ON -EDITERRANEAN ISLANDS ARE SPECIES TRAITS IMPORTANT 'LOBAL
%COLOGY AND "IOGEOGRAPHY
,ODGE $ - "IOLOGICAL )NVASIONS ,ESSONS FOR %COLOGY 4RENDS IN %COLOGY
%VOLUTION
,ORTIE # * 2 7 "ROOKER 0 #HOLER : +IKVIDZE 2 -ICHALET & ) 0UGNAIRE AND
2 - #ALLAWAY 2ETHINKING PLANT COMMUNITY THEORY /IKOS
-ACK 2 . $ 3IMBERLOFF 7 - ,ONSDALE ( %VANS - #LOUT AND & ! "AZZAZ
"IOTIC INVASIONS #AUSES EPIDEMIOLOGY GLOBAL CONSEQUENCES AND CONTROL %COLOGICAL
!PPLICATIONS
-EMMOTT * . - 7ASER AND - 6 0RICE 4OLERANCE OF POLLINATION NETWORKS TO SPE
CIES EXTINCTIONS 0ROCEEDINGS OF THE 2OYAL 3OCIETY OF ,ONDON 3ERIES "
"IOLOGICAL 3CIENCES
0ARKER ) - AND + ! (AUBENSAK #OMPARATIVE POLLINATOR LIMITATION OF TWO NON
NATIVE SHRUBS DO MUTUALISMS INmUENCE INVASIONS /ECOLOGIA
0ARKER ) - $ 3IMBERLOFF 7 - ,ONSDALE + 'OODELL - 7ONHAM 0 - +AREIVA
- ( 7ILLIAMSON " 6ON (OLLE 0 " -OYLE * % "YERS AND , 'OLDWASSER
)MPACT TOWARD A FRAMEWORK FOR UNDERSTANDING THE ECOLOGICAL EFFECTS OF INVADERS
"IOLOGICAL )NVASIONS
0ETROVSKII 3 ! -OROZOV AND " , ,I 2EGIMES OF BIOLOGICAL INVASION IN A PREDATOR
PREY SYSTEM WITH THE !LLEE EFFECT "ULLETIN OF -ATHEMATICAL "IOLOGY
0UTH , - AND $ - 0OST 3TUDYING INVASION HAVE WE MISSED THE BOAT %COLOGY
,ETTERS
3- -C-AHON -7 #ADOTTE AND 4 &UKAMI
2ICHARDSON $ - . !LLSOPP # - $!NTONIO 3 * -ILTON AND - 2EJMANEK 0LANT
INVASIONS THE ROLE OF MUTUALISMS "IOLOGICAL 2EVIEWS
3AKAI ! + & 7 !LLENDORF * 3 (OLT $ - ,ODGE * -OLOFSKY + ! 7ITH 3 "AUGHMAN
2 * #ABIN * % #OHEN . # %LLSTRAND $ % -C#AULEY 0 /.EIL ) - 0ARKER
* . 4HOMPSON AND 3 ' 7ELLER 4HE POPULATION BIOLOGY OF INVASIVE SPECIES
!NNUAL 2EVIEW OF %COLOGY AND 3YSTEMATICS
3AX $ & * * 3TACHOWICZ AND 3 $ 'AINES EDITORS 3PECIES )NVASIONS )NSIGHTS INTO
%COLOGY %VOLUTION AND "IOGEOGRAPHY 3INAUER !SSOCIATES 3UNDERLAND -!
3CHWEITZER * ! AND + # ,ARSON 'REATER MORPHOLOGICAL PLASTICITY OF EXOTIC HONEY
SUCKLE SPECIES MAY MAKE THEM BETTER INVADERS THAN NATIVE SPECIES *OURNAL OF THE 4ORREY
"OTANICAL 3OCIETY
3HEA + AND 0 #HESSON #OMMUNITY ECOLOGY THEORY AS A FRAMEWORK FOR BIOLOGICAL
INVASIONS 4RENDS IN %COLOGY %VOLUTION
4HOMSON $ - -ATRIX MODELS AS A TOOL FOR UNDERSTANDING INVASIVE PLANT AND NATIVE
PLANT INTERACTIONS #ONSERVATION "IOLOGY
4ILMAN $ #OMMUNITY INVASIBILITY RECRUITMENT LIMITATION AND GRASSLAND BIODIVER
SITY %COLOGY
7OOTTON * 4 4HE .ATURE AND #ONSEQUENCES OF )NDIRECT %FFECTS IN %COLOGICAL
#OMMUNITIES !NNUAL 2EVIEW OF %COLOGY AND 3YSTEMATICS
#HAPTER TWO
$ARWIN TO %LTON
EARLY ECOLOGY AND THE PROBLEM
OF INVASIVE SPECIES
- 7 #ADOTTE
).42/$5#4)/.
! NUMBER OF PROMINENT MODERN AUTHORS HAVE PERCEIVED A CHASM SEPARATING
MAINSTREAM ECOLOGY AND INVASION BIOLOGY EG $AVIS ET AL )NTROSPECTION
INTO THE NATURE OF INVASION BIOLOGY COULD BE SEEN AS PART OF A LARGER ENTERPRISE OF
PRESCRIPTIVE PHILOSOPHY WITHIN ECOLOGY EG %GLER 0ETERS ,AWTON
4HESE TYPES OF INTROSPECTION ARE NOT SCIENCE PER SE THAT IS THEY ARE NOT TRY
ING TO ANSWER QUESTIONS THROUGH DEDUCTION AND EMPIRICISM 2ATHER INTROSPECTION
IS ABOUT THE EPISTEMOLOGY OF SCIENCE OR WHICH IDEAS AND CONCEPTS ARE VALUABLE AND
WHAT KNOWLEDGE IS IMPORTANT BASICALLY (/7 SCIENCE IS DONE %PISTEMOLOGY IS
NO TRIVIAL MATTER WHAT QUESTIONS WE ASK AND HOW WE COME TO CONCLUSIONS ARE
UNDER THE PURVIEW OF EPISTEMOLOGY 4HIS CHAPTER IS ABOUT THE EPISTEMOLOGY OF
ECOLOGICAL RESEARCH AND INVASION BIOLOGY
2ECENT CALLS FOR ECOLOGY AND INVASION BIOLOGY TO REINTEGRATE *OHNSTONE
(USTON 4HOMPSON ET AL $AVIS ET AL ARE BASED UPON A PERCEIVED
DISSOCIATION INSTIGATED BY %LTONS MAGISTERIAL BOOK ON INVASION BIOLOGY
4HESE CALLS ) BELIEVE RIGHTLY DRAW ATTENTION TO THE POTENTIAL FERTILE GROUND THAT LAY
BETWEEN ECOLOGY AND INVASION BIOLOGY (OWEVER IF %LTON CAUSED THE DISSOCIATION
-7 #ADOTTE ET AL EDS #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY n
3PRINGER 0RINTED IN THE .ETHERLANDS
-7 #ADOTTE
$AVIS ET AL THEN WHAT WAS THE PRIOR STATE OF THE RELATIONSHIP BETWEEN ECOL
OGY AND NON
INDIGENOUS SPECIES .)3 ) WILL EXAMINE HOW EARLY ECOLOGISTS DEALT
WITH NON
NATIVE SPECIES IN THEIR WRITINGS ESPECIALLY AT THE TIME WHEN ECOLOGY
BECAME ITS OWN lELD OF SCIENCE LATE S TO EARLY S
%#/,/')#!, 3#)%.#% !4 4(% "%')..).'
%VEN THOUGH MANY GREAT THINKERS THOUGHT ABOUT WHY SPECIES OCCUR WHERE THEY
DO EG !RISTOTLE 0ALEY !GASSIZ DE #ANDOLLE IT WASNT UNTIL THE MID S THAT
THESE TYPES OF INVESTIGATIONS MOVED BEYOND METAPHYSICAL ACCOUNTS AND BEGAN TO
USE WHAT COULD BE REFERRED TO AS MODERN SCIENTIlC EPISTEMOLOGY %ARLY @ECOLOGY
WAS TYPIlED BY NATURAL HISTORY OBSERVATIONS %GERTON BUT THE CONTRIBUTION
OF EARLY WORKERS CANNOT BE UNDERVALUED SIMPLY BECAUSE SCIENTIlC EPISTEMOLOGY
WAS NOT AN INDEPENDENT BRANCH OF KNOWLEDGE 4HAT SAID THE NEW SCIENCE THAT WAS
TO EMERGE IN THE MID TO LATE S WOULD ECLIPSE IN A MATTER OF DECADES THE PREVI
OUS YEARS OF THOUGHT ON THE ORIGIN AND DISTRIBUTION OF SPECIES "EFORE THERE
COULD BE A SCIENCE OF ECOLOGY THE ULTIMATE CAUSES OF SPECIES DIVERSITY AND DISTRIBU
TIONS NEED TO BE ARTICULATED IN A WAY THAT WOULD REMOVE NON
NATURALISTIC EXPLANA
TIONS 7HAT FOLLOWS THROUGH THE REST OF THIS ESSAY IS NOT MEANT TO BE A EXHAUSTIVE
SUMMARY OF EARLY ECOLOGY FOR THAT SEE %GERTON AND -C)NTOSH
RATHER ) WILL USE SELECTED EXAMPLES TO EXAMINE HOW EARLY ECOLOGY DEALT WITH .)3
%ARLY PRE
$ARWINIAN ECOLOGY WAS IMBUED WITH NOTIONS OF @BALANCE OF NATURE
%GERTON AND SO CHANGES IN THE ABUNDANCE AND DISTRIBUTION OF SPECIES WERE
NOT PROMINENT IN EARLY WRITINGS 4HE SCIENCE OF SPECIES DISTRIBUTIONS WAS LARGELY
CONCEIVED IN THE TH AND TH CENTURIES IN THE WORK OF "UFFON DE #ANDOLLE AND
$ARWIN AND STRONG EXPLANATORY SCIENCE DID NOT EMERGE UNTIL #HARLES $ARWINS
4HE /RIGIN OF THE 3PECIES IN 4HOUGH SOME OF $ARWINS CONTEMPORARIES DID
NOT RECOGNIZE 4HE /RIGIN AS @SCIENTIlC ON THE GROUNDS THAT IT WAS NOT INDUCTIVE
ENOUGH /WENS 3EDGWICK $ARWINS CONCEPTUAL LEAPS AND THE LINK
ING OF FORMERLY DISPARATE PIECES OF INFORMATION RESEMBLE MODERN SCIENCE MORE
THAN ANYTHING THAT PREDATES $ARWIN
)N TRYING TO EXPLAIN PATTERNS OF SPECIES DISTRIBUTIONS AND ABUNDANCE $ARWIN
WAS FACED WITH THE PROBLEM OF .)3 (OW DID $ARWIN ASSIMILATE SUCH AN APPAR
ENT FACT THAT SPECIES READILY MOVE FROM THEIR NATIVE RANGE $ARWIN CLEVERLY USED
NON
INDIGENOUS SPECIES AS A DEVICE TO TEST OR ILLUSTRATE HIS THEORY OF NATURAL SELEC
TION AND DESCENT WITH MODIlCATION .)3 ARE SPECIES LOCALLY COEXISTING WITH OTHER
SPECIES YET INDEPENDENT OF THESE LOCAL ASSEMBLAGES WITH THEIR OWN EVOLUTIONARY
HISTORIES 4HEREFORE $ARWIN SAW THE PRESENCE OF .)3 AS EFlCACIOUS EXAMPLES OF
NATURAL SELECTION IN ALREADY DISTINCT FORMS
$ARWIN NOTES THAT AN IMPORTANT ASPECT OF THE STRUGGLE FOR EXISTENCE IS THAT
SPECIES NATURALLY HAVE A PROPENSITY TO INCREASE AT HIGH RATES SO THE STRUGGLE IS
AN ISSUE OF POPULATION PROCESSES (E MENTIONS THAT WHEN DOMESTIC ANIMALS ARE
RELEASED THEIR POPULATIONS OFTEN RAPIDLY INCREASE &URTHERMORE hSO IS IT WITH PLANTSv
$ARWIN TO %LTON EARLY ECOLOGY AND THE PROBLEM OF INVASIVE SPECIES
THAT THERE ARE hINTRODUCED PLANTS WHICH HAVE BECOME COMMON THROUGHOUT WHOLE
ISLANDS IN A PERIOD OF LESS THAN TEN YEARSv P 4HE EXPLANATION FOR THIS IS
hTHAT THE CONDITIONS OF LIFE HAVE BEEN VERY FAVOURABLE AND THAT THERE HAS CONSE
QUENTLY BEEN LESS DESTRUCTION OF THE YOUNG AND OLDv P %SSENTIALLY WHAT
BETTER WAY TO SEE THE POTENTIAL HIGH RATES OF INCREASE THAN IN SPECIES ARRIVING IN
PLACES WHERE THEY DIDNT EXIST BEFORE
)SOLATION IS A FUNDAMENTAL PROCESS IN ECOLOGY AND EVOLUTIONARY BIOLOGY AND
$ARWIN REMINDS THE READER OF THIS THAT ON ISOLATED ISLANDS hLET IT BE REMEMBERED
HOW POWERFUL THE INmUENCE OF A SINGLE INTRODUCED TREE OR MAMMALv P
!DAPTATION IS NOT ABSOLUTE IT IS RELATIVE TO SPECIES THAT EXIST TOGETHER AND THAT
BARRIERS TO IMMIGRATION THAT ISOLATE HABITATS hINTO WHICH NEW AND BETTER ADAPTED
FORMS COULD NOT FREELY ENTERv AND THAT hHAD THE AREA BEEN OPEN TO IMMIGRATION
THESE SAME PLACES WOULD HENCE BEEN SEIZED ON BY INTRUDERSv P 4HERE IS THEN
ALWAYS ROOM FOR IMPROVEMENT AND hWE MAY SAFELY CONCLUDED THAT THE NATIVES
MIGHT HAVE BEEN MODIlED WITH ADVANTAGE SO AS TO HAVE BETTER RESISTED SUCH
INTRUDERSv P &OR EXAMPLE h 4HE ENDEMIC PRODUCTIONS OF .EW :EALAND ARE
PERFECT ONE COMPARED WITH ANOTHER BUT THEY ARE NOW RAPIDLY YIELDING BEFORE THE
ADVANCING LEGIONS OF PLANTS AND ANIMALS INTRODUCED FROM %UROPEv P
7HEREAS SPECIES DIFFERENCES ARE SLIGHT COMPARED TO THOSE AMONG GENERA
$ARWIN EXPECTED AS !LPHONSE DE #ANDOLLE SUGGESTED hTHAT mORAS GAIN
BY NATURALISATION x PROPORTIONALLY x FAR MORE IN NEW GENERA THAN IN NEW SPECIESv
$ARWIN P SEE $AEHLER
).6!3)6% 30%#)%3 !.$ %#/,/'9 "%&/2% 4(% 4( #%.4529
"Y THE EARLY TH CENTURY ECOLOGY WAS A SCIENCE AND EXOTIC SPECIES WERE A CON
CERN BUT THESE TWO THINGS DID NOT SEEM TO INmUENCE ONE ANOTHER &OR ONE THING
MODERN CALLS FOR ECOLOGISTS TO WORK TOWARDS SAVING THE NATURAL WORLD EG 0ETERS
3HRADER
&RECHETTE %HRLICH SIMPLY DID NOT HAPPEN UNTIL THE
MID TH CENTURY SEE $AVIS #HAPTER )NCIPIENT MODERNISM WAS SURFACING AND
MODERN CIVILIZATION AND HER SCIENCE WAS QUICKLY BEING REGARDED AS THE ONLY VALID
KNOWLEDGE
ACQUIRING METHOD AND WAS NEUTRAL TO OR ABOVE MORALITY AND ETHICS
)N !MERICA THIS PERIOD SAW UNPRECEDENTED CHANGE IN HIGHER EDUCATION AND THE
PRE
EXISTING EMPHASIS ON THEOLOGY AND PHILOSOPHY WAS BEING REPLACED WITH SCI
ENCE AND PSYCHOLOGY CURRICULA WITH MANY INSTITUTIONS DEPARTMENTS AND ENDOW
MENTS CREATED PROMOTING THE TRAINING OF PROFESSIONAL SCIENTISTS AT POSTSECONDARY
INSTITUTIONS #ROKER -ENAND 4HIS WAS THE TIME PERIOD WHEN GREAT
!MERICAN THINKERS SUCH AS 7ILLIAM *AMES AND *OHN $EWEY WOULD CULL OUT THE
LAST REMAINING NOTIONS OF SUPERNATURAL EXPLANATION IN !MERICAN !CEDEMIA
"UT ALL WAS NOT WELL PEOPLE WERE NOTICING THAT MODERN CIVILIZATION WAS HAV
ING UNDESIRABLE IMPACTS ON THE NATURAL WORLD )N A DEVASTATING YELLOW
FEVER OUTBREAK IN 0HILADELPHIA LIKELY IMPORTED FROM THE 7EST )NDIES HAD PEOPLE
FEARING THE ACCEDENTAL IMPORTATION OF PATHOGENIC .)3 %GERTON 2ALPH
-7 #ADOTTE
7ALDO %MERSON HAD ALREADY BEEN PLEADING TO THE !MERICAN PUBLIC TO RETHINK ITS
UNHEALTHY RELATIONSHIP WITH NATURE %MERSON 4HE IMPACT OF CERTAIN INVAD
ERS WAS BECOMING APPARENT %IGHTEENTH CENTURY BOTANISTS SUCH AS *OHN "ARTRAM
WERE STARTING TO COMMENT ON THE FACT THAT SOME INTRODUCED PLANTS WERE HAVING
NEGATIVE IMPACTS AND WERE DIFlCULT TO CONTROL SEE -ACK (OOKER
NOTICED THAT %UROPEAN SPECIES WERE QUICKLY REPLACING NATIVES IN .EW :EALAND
3OME OF THE EARLIEST CONCERNS ABOUT EXOTIC SPECIES WERE IN TERMS OF THE MAIN
TENANCE OF THE @NATURAL BASED ON A MORALITY OF AESTHETICS EG /LMSTEAD
3ARGENT BOTH CITED AND DISCUSSED BY 3IMBERLOFF 3OON AFTER A MORE
DISTINCTLY !MERICAN PRAGMATIC APPROACH RECOGNIZED THE ECONOMIC AGRICUL
TURAL AND ECOLOGICAL EFFECTS BY NON
INDIGENOUS SPECIES EG &ITCH (OWARD
A )N HIS ADDRESS TO THE !MERICAN !SSOCIATION FOR THE !DVANCEMENT
OF 3CIENCE ,ELAND (OWARD SAYS h0OTENTIALLY COSMOPOLITAN FORMS x HAVE BY
HUMAN AGENCY BECOME DISPERSED OVER NEARLY ALL OF THE CIVILIZED PARTS OF THE
GLOBE WHILE THOUSANDS OF OTHER SPECIES HAVE BEEN CARRIED THOUSANDS OF MILES FROM
THEIR NATIVE HOMES AND HAVE ESTABLISHED THEMSELVES AND mOURISHED OFTEN WITH A
NEW VIGOR IN A NEW SOIL AND WITH A NOVEL ENVIRONMENTv (OWARD A P
(OWARD PUBLISHED MORE THAN ARTICLES ON NON
NATIVE INSECT PESTS AND THE
REALIZED OR POTENTIAL IMPACTS OF INTRODUCED SPECIES EG (OWARD
A B (OWARDS PEER AND FRIEND 3TEPHEN &ORBES THE
FATHER OF !MERICAN ECOLOGY ALSO BECAME PROFESSIONALLY DEDICATED TO THE EFFECTS
THAT INSECT PESTS POSED FOR AGRICULTURE #ROKER &ORBES WROTE MANY ARTICLES
ON THE BIOLOGY AND IMPACTS OF INSECT PESTS EG &ORBES A
"Y THE TURN OF THE CENTURY WE HAVE LEGITIMATE NON
AESTHETIC CONCERNS
ABOUT THE IMPORTATION OF NOVEL SPECIES /F COURSE MUCH OF THIS CONCERN WAS IN
TERMS OF AGRICULTURAL AND ECONOMIC COSTS $URING THIS TIME ECOLOGY WAS DEVELOP
ING INTO ITS OWN SCIENCE %GERTON -C)NTOSH AND HOW DID THIS SCIENCE
OF ECOLOGY ADDRESS THESE CONCERNS ABOUT INVASIVE SPECIES
$URING THIS PERIOD THE DEVELOPMENT OF THE SCIENCE OF ECOLOGY WAS LARGELY A
%UROPEAN PHENOMENON WITHOUT A UNIQUELY !MERICAN SCHOOL UNTIL SHORTLY AFTER
THE TURN OF THE CENTURY -C)NTOSH /F COURSE THERE WAS &ORBES WHO WROTE
SEVERAL PAPERS BEFORE THE TH CENTURY WHICH WE SHOULD CALL @ECOLOGICAL ESPECIALLY
HIS FAMOUS ,AKE AS A -ICROCOSM TALK DELIVERED TO THE 0EORIA 3CIENTIlC !SSOCIATION
&ORBES B (ERE HE STRESSED THE INTERACTION AMONG SPECIES AND A BALANCE OF
NATURE VIEW BUT WE CANNOT CALL THIS A @SCHOOL AS LITTLE SUBSEQUENT WORK WAS DONE
SPAWNING LITTLE RESEARCH AND NO TEXTBOOKS &ORBES AND HIS CONTEMPORARIES (ENRY
#OWLES AND ,ELAND (OWARD COULD BE SAID TO HAVE STARTED APPLIED ECOLOGY )T IS
CURIOUS THEN THAT ON THE ONE HAND &ORBES WORKED EXTENSIVELY WITH INSECT PESTS
AND EXOTIC AGRICULTURAL DISEASES AND YET MAINTAINED A CLOSED
SYSTEM BALANCE
OF NATURE APPROACH IN HIS ECOLOGICAL WRITINGS 4HE FACT THAT A SINGLE INDIVIDUAL
RECOGNIZING THE IMPORTANCE OF PESTS AND UNDERSTANDING ECOLOGICAL SYSTEMS WOULD
KEEP THESE IDEAS DISSOCIATED IS PROPHETIC FOR THE REST OF EARLY ECOLOGY
%COLOGY GREW OUT OF THE APPARENT NEED TO SEPARATE CAUSES IN WHOLE
ORGANISM
BIOLOGY 7HEREAS $ARWINS THEORY EXPLAINED THE ULTIMATE CAUSES AND PATTERNS
$ARWIN TO %LTON EARLY ECOLOGY AND THE PROBLEM OF INVASIVE SPECIES
OF WHY WE lND SPECIES WHERE WE DO EG WHY CACTI ARE DIVERSE IN DESERT REGIONS
AND THE PURPOSE OF THE PANDAS THUMB IT DID NOT INCLUDE A STRONG FRAMEWORK
TO ACCOUNT FOR PROXIMATE CAUSES AND PATTERNS EG CYCLIC POPULATION DYNAMICS
AND PATTERNS OF ABUNDANCES %ARLY FORAYS INTO THE ULTIMATE CAUSES OF SPECIES
DISTRIBUTIONS FEATURED GEOGRAPHICAL AND GEOLOGICAL CAUSES OF SPECIES DISTRIBUTIONS
EG 7ALLACE 7ALLACE OBSERVED THAT BIOTAS WERE LARGE REGIONAL
ENTITIES OFTEN WITH FAIRLY DISCRETE BOUNDARIES #OMMON WAS THE IDEA OF BARRIERS TO
MOVEMENT WITH ALL SPECIES BEING ABLE TO MOVE TO NEW LOCALS BUT RARELY DOING SO
EG (EILPRIN 7RITINGS OF DE #ANDOLLE AND $ARWIN CERTAINLY ADDRESS ISSUES
OF PROXIMATE PATTERNS BUT IT TOOK A SEPARATE lELD TO CREATE THEIR OWN PROXIMATE
EXPLANATIONS
4HE BIOGEOGRAPHER /SCAR $RUDE WAS ABLE TO SEPARATE THESE CAUSES AND PAT
TERNS IN HIS -ANUAL DE 'EOGRAPHIE "OTANIQUE (E NOTED THAT THERE WERE
TWO FORCES OF CHANGE IN SPECIES RANGES AND ABUNDANCE GEOLOGICAL AND BIOLOGICAL
$RUDE RIGHTLY THOUGHT THAT ALL ULTIMATE CHANGE HAPPENED IN hCOURS DE LVOLUTION
GEOLOGIQUEv P THE COURSE OF GEOLOGICAL EVOLUTION 4HIS INCLUDED ADAPTA
TION AS WELL AS CONSTANT CHANGE BUT HE ALSO SAW THE ROLES OF THE ORGANISM
ENVIRONMENT INTERFACE DISPERSAL AND COMPETITION 4HE DISTINCTION BETWEEN GEOLOGICAL
AND BIOLOGICAL CAUSES WAS IMPORTANT FOR $RUDES UNDERSTANDING OF RAPID CHANGES IN
SPECIES DISTRIBUTIONS $RUDE CLASSIlES LIMITS TO SPECIES RANGES AS EITHER GEOGRAPHI
CAL THAT IS MOUNTAINS OCEANS AND THE LIKE ARISING FROM GEOLOGICAL PROCESSES
AND BIOLOGICAL BY WHICH HE MEANS CHANCE DISPERSAL AND THE AGENCY OF HUMANS
h,ES LIMITS DES AIRES SONT PUREMENT GOGRAPHIQUES OU BIEN RSULTENT DE LACTION
COMBINE DES CONDITIONS DE VIEv P $RUDE SAYS THAT WITHOUT THE AID OF HUMANS
GEOGRAPHICAL BARRIERS LIMIT SPECIES SPREAD BUT THAT RARE CHANCE EVENTS STILL PLAY
AN IMPORTANT ROLE FOR RANGE EXPANSION 7ITH HUMAN AID SPECIES WILL SPREAD AS FAR
AS THE EXTERNAL ENVIRONMENT ALLOWS $RUDE TELLS THE READER THAT MOVING TO A NEW
LOCATION WHICH MUST BE CLIMATICALLY SIMILAR TO THAT IN A SPECIES NATIVE RANGE
IS ONLY HALF THE EQUATION AND ONCE A PLANT HAS ARRIVED hELLE LUTTE AVEC SUCCES CON
TRE UNE PLANTE INDIGNE TOUT
FAIT ADAPTE AU CLIMATv P IT HAS TO BE SUC
CESSFUL IN THE STRUGGLE AGAINST INDIGENOUS PLANTS PERFECTLY ADAPTED TO THE CLIMATE
'IVEN THIS LIMITATION PLANTS STILL COME TO hCOUVRIR DE GRANDES SURFACESv P
AND IN COVERING THESE LARGE AREAS DISPLACE OTHER SPECIES (E GOES EVEN FURTHER TO
SAY THAT SOME SPECIES DO NOT DOMINATE ONE ANOTHER AND COMPETITION IS INDETERMI
NATE )N THESE SPECIES THE RANGES ARE SET UP BY WHOEVER ARRIVES THERE lRST
$RUDES ACCOUNT SOUNDS VERY MODERN (IS BIOGEOGRAPHY INCORPORATES GEOGRA
PHY CHANCE COMMUNITY ASSEMBLY AND LOCAL COMPETITION (E ALSO HAS A SYSTEM OF
PROXIMATE CAUSES AND PATTERNS THAT READILY ACCOUNTS FOR THE DYNAMICS OF INVASIVE
SPECIES (OWEVER IN READING THE CHAPTER ON RANGE EXPANSION IN $RUDE
ONE GETS THE IMPRESSION THAT HE IS TRYING TO EXPLAIN THE FACT OF .)3 SPREAD IN LIGHT OF
LATE
NINETEENTH CENTURY KNOWLEDGE OF BIOGEOGRAPHY RATHER THAN TRYING TO UNDER
STAND INVASIONS AND SPECIlC INVADERS 4HIS IS NOT A CRITICISM AS HE WAS DEVELOPING
A SYSTEM OF KNOWLEDGE ABOUT THE NATURAL WORLD AND HIS EPISTEMOLOGY CALLED FOR
EXPLAINING EXTANT PATTERNS WITHIN THE EXISTING FRAMEWORK
-7 #ADOTTE
!NOTHER %UROPEAN ECOLOGIST %UGENE 7ARMING WROTE AN INmUENTIAL ECOLOGY
BOOK BEFORE THE TURN OF THE CENTURY 4HE %NGLISH TRANSLATION OF HIS /ECOLOGY
OF 0LANTS WAS CONCERNED WITH POPULATION AND COMMUNITY ECOLOGY AND
HE TRIED TO DEVELOP A SYSTEM OF ECOLOGICAL UNDERSTANDING 7ARMING WAS VERY
MUCH INTERESTED IN THE PROCESS OF ECOLOGICAL CHANGE AND THIS CHANGE INCLUDED THE
ARRIVAL OF NEW SPECIES (E BELIEVED THAT ECOLOGICAL PROCESSES INCLUDING CHANGE
WERE REALIZED AT THE COMMUNITY LEVEL &OR EXAMPLE HE BELIEVED THAT SUCCESSIONAL
CHANGE RESULTED FROM COMPETITION AMONG COMMUNITY TYPES AFFECTED BY CHANGES
IN THE ENVIRONMENT %SSENTIALLY 7ARMINGS CHANGE IS ASSEMBLAGES OF SPECIES
MOVING AMONG EQUILIBRIA BUT WHAT WAS HIS MECHANISM FOR THESE CHANGES
7ARMING SAYS THAT THERE ARE TWO TYPES OF COMMUNITY CHANGE lRST IS FROM
@CHANGES IN SOIL AND THE OTHER IS CHANGE @WITHOUT NEW SOIL )N THE lRST TYPE
OF CHANGE DISTURBANCES BOTH NATURAL AND HUMAN CAUSED ARE IMPORTANT AND
ABRUPT 3LOWER TYPES OF CHANGE INCLUDE INCREASES OR DECREASES IN WATER CONTENT OR
BY THE ACCUMULATION OF SOIL 4HE SECOND TYPE OF CHANGE HAPPENS THROUGH MIGRA
TIONS AND STRUGGLES AMONG INDIVIDUAL SPECIES
4HIS SECOND TYPE OF COMMUNITY CHANGE 7ARMING SAYS IS ATYPICAL hTHE STRUG
GLES IN QUESTION ARE RAREv AND THAT COUNTRIES hWHOSE VEGETATION IS NOT APPRECIABLY
DISTURBED BY MAN OR ANIMALS AND WHICH HAS BEEN EXPOSED FOR LONG AGES PAST TO
IMMIGRATION FROM ADJOINING COUNTRIES IN THIS CASE A CERTAIN EQUILIBRIUM MUST
HAVE BEEN ATTAINEDv P 4HIS EQUILIBRIUM VIEW MEANS THAT THE PROBLEM OF
INVASIVE SPECIES IS ONE PRIMARILY ASSOCIATED WITH DISTURBANCES WHICH IS LARGELY
TRUE (OWEVER HIS VIEW MAY HAVE SOME DIFlCULTLY EXPLAINING WHY SOME SPE
CIES INVADE INTACT COMMUNITIES (E CONCEDES THIS POINT ON THE NEXT PAGE hSOME
CHANGES IN THE NATURE OF FORESTS MAY HAVE BEEN SIMPLY DUE TO THE ENTRY OF NEW
SPECIESv P
7ARMING DOES MAKE AN EMOTIONAL PLEA TO BE WEARY OF IMPORTING NEW SPECIES
BECAUSE OF THE COMPETITION TO WHICH THEY MAY SUBJECT NATIVE FORMS hA NEW x COM
MUNITY WHICH DID NOT NATURALLY OCCUR BEFORE THE SOIL HAD BEEN DRAWN INTO THE
SERVICE OF -AN AND THE SPECIES WHICH NOW OCCUR IN VAST NUMBERS AND FORM A
COMMUNITY WITH ITS OWN SPECIAL STAMP AND ECONOMYv P
7ARMINGS SYSTEM MAKES PREDICTIONS AS TO WHAT TYPES OF COMMUNITIES OUGHT TO
BE MORE INVADED AND OUR MODERN UNDERSTANDING OF THE INVASION PROCESS WOULD
ALSO PREDICT THAT INTACT COMMUNITIES ARE LESS INVADABLE THAN DISTURBED ONES
(OWEVER BOTH 7ARMING AND $RUDE WERE NOT EXPLICITLY TRYING TO UNDERSTAND
INVASIONS RATHER THEY WERE TRYING TO lT THE FACT THAT WE SEE INVADERS INTO THEIR
RESPECTIVE ECOLOGICAL SYSTEMS &OR THEM THERE WAS NO REAL ASSOCIATION BETWEEN
DEVELOPING ECOLOGICAL UNDERSTANDING AND ADDRESSING THE PROBLEM OF INVASIVE
SPECIES 4HEY FELT THAT IF THEIR RESEARCH PROGRAMMES EXPLAINED THE GENERAL
PHENOMENON OF .)3 THEN THEIR ECOLOGY WAS ROBUST 4HE RELATIONSHIP BETWEEN
ECOLOGY AND INVADERS WAS LIMITED AND ONE WAY !T MOST INVADERS WERE TESTS OF
EARLY ECOLOGISTS IDEAS OR AT WORST PHENOMENA NEEDING EXPLANATION
$ARWIN TO %LTON EARLY ECOLOGY AND THE PROBLEM OF INVASIVE SPECIES
4(% .!452% /& #(!.'% #,%-%.43 !.$ 4(% !-%2)#!. 3#(//,
3HORTLY AFTER THE TURN OF THE CENTURY AN !MERICAN SCHOOL EMBODIED IN &REDERIC
#LEMENTS COME TO THE FORE -C)NTOSH ) SAY THAT #LEMENTS EMBODIED THE
SCHOOL BECAUSE HE DID ALL THE THINGS THAT MAKE A SUCCESSFUL SCHOOL OF THOUGHT
(E INTERACTED WITH A NUMBER OF STUDENTS AND COLLEAGUES HE WROTE NUMEROUS
SUCCESSFUL TEXTBOOKS AND MOST IMPORTANTLY HE EMPHASIZED A STRONG HEURISTIC
PARADIGM IN MUCH OF HIS WRITING NAMELY THE NATURE OF ECOLOGICAL CHANGE
#LEMENTS lRST TWO BOOKS REALLY LAID THE GROUNDWORK FOR THIS NEW SCHOOL (IS
lRST 2ESEARCH -ETHODS IN %COLOGY DETAILED HOW THE SCIENCE OF ECOLOGY
SHOULD BE DONE WHAT TECHNIQUES AND RESEARCH TOPICS WERE MOST FRUITFUL 4HIS WAS
SOON FOLLOWED BY A SECOND BOOK 0LANT 0HYSIOLOGY AND %COLOGY WHICH HAD
MUCH OVERLAP WITH HIS BOOK BUT REINFORCED WHICH AREAS OF RESEARCH WERE
MOST IMPORTANT AND THE PARADIGM OF PLANT ECOLOGY 3PECIlCALLY THESE BOOKS HAD
AN UNPRECEDENTED FOCUS ON ECOLOGICAL CHANGE OR WHAT BECAME KNOWN AS SUCCES
SION )NTUITIVELY SUCH A PARADIGM SEEMS LIKE .)3 WOULD BE OF CENTRAL CONCERN
"OTH BOOKS DElNE INVASION ALMOST IDENTICALLY AS h4HE MOVEMENT OF ONE OR
MORE PLANTS FROM ONE AREA INTO ANOTHER AND THEIR ESTABLISHMENT IN THE LATTER
P ALSO A VERY SIMILAR QUOTE IN P (E NOTES h)NVASION
IS A REGULAR OCCURRENCE BETWEEN ADJACENT FORMATIONS BUT IT ALSO TAKES PLACE
INTO REMOTE ONES AS A RESULT OF LONG CARRIAGE BY WIND WATER BIRDS RAILROADS
OR VESSELSv P (E TELLS US THAT DISPERSAL IS USUALLY CONSTRAINED BY
BARRIERS BUT THAT HUMANS AS AN AGENT OF DISPERSAL ARE ABLE TO TRANSCEND THESE
BARRIERS )NVASIONS ARE IMPORTANT FOR A NUMBER OF REASONS &IRST INVASIONS CAN
SIGNIlCANTLY AFFECT SPECIES DISTRIBUTIONS 3ECONDLY ECOLOGICAL CHANGE DEPENDS
ON INVASIONS #LEMENTS DElNES SUCCESSION AS A hSERIES OF INVASIONS xIN THE SAME
SPOTv P P
/F COURSE SEVERAL FACTORS WORK AGAINST THE SUCCESS OF AN INVASION &IRST OF ALL
THERE ARE THE WELL KNOWN GEOGRAPHICAL BARRIERS TO MIGRATION MOUNTAINS OCEANS
ETC BUT MORE IMPORTANTLY #LEMENTS DETAILS OTHER TYPES OF BARRIERS 4HERE IS AN
ABIOTIC BARRIER THE SPECIES MUST lND ITSELF IN AN ENVIRONMENT IN WHICH IT CAN
SURVIVE -ANY BIOLOGICAL FORCES WORK AGAINST SPECIES ESTABLISHMENT INCLUDING
PREDATION DISEASE COMPETITION AND HUMANS TRYING TO CONTROL WEEDS !LSO
#LEMENTS lNDS THAT WHETHER THE HABITAT IS FULL OR NOT IS OF PARAMOUNT IMPORTANCE
TO INVASION SUCCESS AS COMPETITION FROM FULL COMMUNITIES WILL BE TOO SEVERE FOR
THE COLONIZER TO SUCCEED
4HIS VIEW OF ECOLOGICAL SUCCESSION IS COMMON IN THE WRITINGS OF #LEMENTS
COLLEAGUES AND COLLABORATORS 3HANTZ SAW INVASIONS AS THE PROCESS OF
SUCCESSION AND SUCCESSION AS THE PROCESS OF CHANGING FORMATIONS OR COMMUNITY
TYPES 4HE IDEA OF SPECIES INVASIONS AS THE FORCE BEHIND SUCCESSION IS ALSO DOMI
NANT IN 7EAVER AND #LEMENTS (UMANITYS ROLE IS EXAMINED IN GREATER
DETAIL AND THEY NOTICE THAT ANTHROPOGENIC EFFECTS ON COMMUNITY SUCCESSION
CAN BE SEVERE AS WE BOTH DISTURB HABITATS AND BRING IN NEW INDIVIDUALS 4HEY
NOTE THAT THE MOTOR OF SUCCESSION IS NOT DISTURBANCE BUT RATHER BIOTIC PROCESSES
-7 #ADOTTE
MIGRATION AND COMPETITION AND THAT CLIMAX COMMUNITIES OFTEN RESIST THESE
BIOTIC FACTORS #LEMENTS AND 3HELFORD CLARIlED WHO ARE CALLED INVADERS
3PECIlCALLY INVADERS ARE THOSE SPECIES THAT DID NOT PREVIOUSLY EXIST IN THE HABITAT
THEY ARE INVADING 0REVIOUSLY #LEMENTS DID NOT EXPLICITLY STIPULATE WHETHER THE
INVADERS HAD TO BE SEPARATE SPECIES (ERE #LEMENTS AND 3HELFORD MAKE NO
DISTINCTION ABOUT WHERE SPECIES COME FROM IN TERMS OF THEIR NATIVE OR INTRODUCED
STATUS %ARLIER #LEMENTS DID CREATE TERMS TO REmECT SPECIES THAT MIGRATE
FROM NEARBY VERSUS THOSE ORIGINATING FURTHER AWAY 9ET NONE OF THESE WORKS
EXPLICITLY EXAMINES .)3 JUST ECOLOGICAL PROCESSES OF IDEALIZED SPECIES !GAIN LIKE
THE OTHER EARLY ECOLOGISTS #LEMENTS AND COLLEAGUES WERE INTERESTED IN ESTABLISHING
A STRONG PROGRAMME THAT COULD POTENTIALLY EXPLAIN THE OCCURRENCE OF .)3
%80,!).).' 30%#)%3 /##522%.#%3 7)4( 4(% .)#(%
) CONTINUE BY TAKING A BRIEF TOUR OF ECOLOGY UP TO THE S $URING THIS PERIOD
PRIMARY PUBLICATIONS IE ACADEMIC JOURNAL ARTICLES WERE BECOMING INCREASINGLY
IMPORTANT FOR THE DEVELOPMENT AND DISSEMINATION OF NEW IDEAS /NE OF THE MOST
IMPORTANT CONCEPTUAL DEVELOPMENTS IN EARLY ECOLOGY WAS THE IDEA OF THE NICHE
4HROUGH UNDERSTANDING A SPECIES NICHE ONE COULD POTENTIALLY UNDERSTAND AND
PREDICT WHERE AND WHEN SPECIES IS FOUND AND WHICH SPECIES ARE LIKELY TO REPLACE
ONE ANOTHER
7E COULD PROBABLY TRACE THE NICHE CONCEPT ALL THE WAY BACK TO $ARWIN BUT THE
IDEA OF EXAMINING THE DISTRIBUTION OF SPECIES THROUGH A SPECIES SUITE OF ENVIRON
MENTAL NEEDS IS COMMONLY ATTRIBUTED TO 'RINNELL 'RINNELLS NICHE CONCEPT
HAS BEEN AN IMPORTANT THEME IN THE DEVELOPMENT OF ECOLOGY (IS PREMISE WAS THAT
PATTERNS OF SPECIES OCCURRENCE ARE BEST UNDERSTOOD THROUGH THEIR ABIOTIC AND BIOTIC
REQUIREMENTS (UTCHINSON FAMOUSLY USED THIS NICHE CONCEPT TO ADVANCED
THE NOTION OF A MULTIDIMENSIONAL NICHE SPACE DElNING WHERE EACH SPECIES COULD
POTENTIALLY LIVE (UTCHINSONS CONCEPT COULD POTENTIALLY EXPLAIN WHERE SPECIES
COULD LIVE AND THEREFORE WHERE THEY COULD INVADE AS EACH SPECIES POTENTIALLY
OCCUPIES A UNIQUE REGION OF THE NTH DIMENSIONAL NICHE HYPERSPACE 4HE DElNI
TION OF @NICHE HAS BEEN RECOGNIZED AS PROBLEMATIC 3IMBERLOFF #ONNER AND
3IMBERLOFF AND OPEN TO VARIOUS INTERPRETATIONS ,EIBOLD #HASE AND
,EIBOLD !N EARLY REINTERPRETATION WAS PROVIDED BY %LTON WHO SAW
THE NICHE AS A SPECIES @STATUS IN THE COMMUNITY 4HIS INCLUDED THE ROLE AN ORGAN
ISM PLAYED AND MOST IMPORTANTLY ITS RELATIONSHIP TO OTHER SPECIES AND ESPECIALLY
THE TROPHIC STATUS OF A SPECIES %LTONS INVASIONS BOOK ACTUALLY USED BOTH
CONCEPTS OF THE NICHE TO UNDERSTAND WHEN AND WHERE .)3 INVADE AND WHICH COM
MUNITIES WERE MORE LIKELY TO BE INVADED /NLY RECENTLY HAD THERE BEEN AN ATTEMPT
AT RECONCILING THESE TWO MAJOR CONCEPTUALIZATIONS OF THE NICHE CONCEPT ,EIBOLD
#HASE AND ,EIBOLD
3INCE 'RINNELLS PAPER RESEARCH ON COMPETITION HAS BEEN INTIMATELY
ASSOCIATED WITH THE NICHE CONCEPT %LTON LIKE 7ARMING THOUGHT
$ARWIN TO %LTON EARLY ECOLOGY AND THE PROBLEM OF INVASIVE SPECIES
SPECIES
RICH COMMUNITIES OUGHT TO BE LESS INVADABLE BECAUSE AVAILABLE NICHES
SHOULD BE lLLED AND COMPETITION FOR NICHE SPACE MORE INTENSE #OMPETITION AS A
MAJOR STRUCTURING FORCE SURFACED BEFORE %LTON EG 'AUSE $!NCONA
WITH THE FOUNDATION IN WHAT 'AUSE CALLED THE @COMPETITIVE EXCLUSION
PRINCIPLE SEE ALSO (ARDIN PROBABLY lRST ARTICULATED BY 'RINNELL
4HE CENTRAL PREMISE OF THIS PRINCIPLE IS THAT h.O TWO SPECIES OF IDENTICAL OR EVEN
CLOSELY SIMILAR BIOLOGICAL PREDILECTIONS CAN LONG OCCUPY THE SAME NICHE x AT THE
SAME TIMEv 'RINNELL P )N FACT 'RINNELL USED THIS REASON
ING TO ARGUE AGAINST THE PURPOSEFUL INTRODUCTION BY THE STATE OF #LAIFORNIA OF
THE (UNGARIAN PARTRIDGE FOR HUNTING 'RINNELLS CONCERN WAS THAT THE (UNGARIAN
PARTRIDGE SHARED ITS NICHE WITH THE NATIVE #ALIFORNIA QUAIL AND THAT THE LATTER
WOULD LIKELY BE COMPETITIVELY EXCLUDED 4HIS PREDICTION WAS BASED ON THE OBSER
VATION THAT SPECIES HAVING LARGE NATIVE RANGES USUALLY EXCLUDE SMALLER
RANGED
COMPETITORS 'RINNELL APPLIED ECOLOGICAL REASONING TO PREDICTING THE OUTCOME OF
A SPECIES INTRODUCTION
'AUSE SAW THE VALUE THAT INVASIONS HAVE FOR UNDERSTANDING GENERAL
ECOLOGICAL PROCESSES 4O STUDY COMPETITION 'AUSE COMMENTED ON THE IMPORTANCE
OF EXPERIMENTATION AND THAT IN NATURAL EXPERIMENTS hTHE MOST FREQUENT ONES
CONSIST IN THE TRANSPORTATION OF ANIMALS INTO COUNTRIES NEW TO THEM WHICH COM
MONLY LEADS TO A GREAT NUMBER OF HIGHLY INTERESTING PROCESSES OF THE STRUGGLE
FOR EXISTENCEv P /NE EXAMPLE HIGHLIGHTED BY 'AUSE IS THE INTRODUCTION OF
THE %UROPEAN CARP INTO EASTERN .ORTH !MERICA IN THE S AND HOW IT hHAS NOW
IN MANY STREAMS AND LAKES MULTIPLIED TO SUCH AN EXTENT THAT SEVERAL NATIVE SPECIES
ARE FOUND IN GREATLY DIMINISHED NUMBERv P
-UCH UNDERSTANDING OF THE NATURE OF SPECIES INTERACTIONS AND ESPECIALLY THOSE
THAT STIMULATED 'AUSE WAS ADVANCED BY THE THEORETICAL MODELS OF ,OTKA
EG AND 6OLTERRA EG 4HE WORK OF ,OTKA AND 6OLTERRA NOT ONLY
REVOLUTIONIZED HOW ECOLOGISTS THOUGHT ABOUT SPECIES INTERACTIONS BOTH COMPETI
TION AND PREDATION BUT ALSO SHOWED HOW MODELS COULD BE USED TO UNDERSTAND
APPLIED PROBLEMS 4HE FAMOUS INSTANCE IS THAT THE SON
IN
LAW 5MBERTO $!NCONA
ASKED HIS FATHER
IN
LAW 6ITO 6OLTERRA IF IT WERE POSSIBLE TO EXPLAIN MATHEMATI
CALLY WHY IT SHOULD BE THAT WITH THE CESSATION OF lSHING IN &IUME NEAR 4RIESTE
DURING 77) PREDATORY SHARKS AND RAYS CAME TO CONSTITUTE A HIGHER MEAN PROPOR
TION OF THE CATCH BIOMASS EG $!NCONA (ALLAM /F COURSE USING
HIS PREDATOR
PREY MODELS 6OLTERRA WAS ABLE TO EXPLAIN THIS INCREASE IN PREDATOR
PROPORTION 6OLTERRA ASSUMED THAT THE HARVESTING PRIOR TO 77) WAS INDISCRIMI
NANT AND RESULTED IN A lXED PROPORTION OF INDIVIDUALS REMOVED SO THAT THE PERIODIC
EQUILIBRIUM BETWEEN PREDATOR AND PREY DENSITIES SHIFTED TO HAVE A CENTER WITH
HIGHER PREY AND LOWER PREDATOR ABUNDANCE WHEN THERE IS DENSITY INDEPENDENT
MORTALITY
)N THE SAME VEIN $!NCONA SHOWED HOW ,OTKA
6OLTERRA
TYPE FORMULA
TIONS COULD BE USED TO UNDERSTAND THE DYNAMICS OF SPECIES INVASIONS 5SING THESE
TYES OF EQUATIONS $!NCONA SHOWED THAT A NEW INVADER WILL EITHER MOVE
TO AN EQUILIBRIUM OF ZERO DENSITY IE UNSUCCESSFUL OR CHANGE THE EQUILIBRIUM
-7 #ADOTTE
DYNAMICS OF THE EXTANT COMPETITORS WHICH ESSENTIALLY ALTERS THE DENSITIES OF THE
OTHER SPECIES
)T IS WORTH MENTIONING THAT THE ,OTKA
6OLTERRA EQUATIONS HAVE BEEN CRITICIZED
WITH A NUMBER OF SHORTCOMINGS RECOGNIZED NAMELY THEY ARE DIFlCULT TO APPLY IN
SPECIlC CIRCUMSTANCES AND THE INTERACTION COEFlCIENTS ARE PROBLEMATIC TO COMPUTE
(ECK (ALLAM 4HAT SAID THEY DO OFFER GREAT USE AS HEURISTIC DEVICES
AND AS A MECHANISM TO PRODUCE QUALITATIVE HYPOTHESES (ECK 4HESE TYPES
OF MODELS HAVE FOUND USE IN UNDERSTANDING AND PREDICTING THE CONSEQUENCES OF
INVASIONS EG #OURCHAMP ET AL #OURCHAMP AND #AUT #HAPTER 8
5.$%234!.$).' 30!4)!, 302%!$
/NE MAJOR DEVELOPMENT PRIOR TO %LTONS BOOK BRIDGING ECOLOGY AND INVASIONS
BIOLOGY WAS 3KELLAMS USE OF SPATIAL SPREAD MODELS 4HE MAJOR THRUST OF
THIS RESEARCH WAS THE APPLICATION OF MODELS DESCRIBING THE DIFFUSION OF PARTICLES IN
A PHYSICAL SYSTEM TO THE SPREAD OF A REPRODUCING POPULATION OVER A TWO
DIMENSIONAL
landscape (Skellam 1951, Kiersteadt and Slobodkin 1953). Skellam (1951)
APPLIED THESE PHYSICAL MODELS TO UNDERSTANDING AND POTENTIALLY PREDICTING POPULA
TION SPREAD ACROSS A LANDSCAPE "Y THE S THE RAPID SPREAD OF .)3 FROM THEIR
POINTS OF ORIGIN HAD BEEN DOCUMENTED AND TO 3KELLAM WERE IN NEED OF MECHANIS
TIC EXPLANATION ,IKE $ARWIN 3KELLAM USED .)3 TO SHOW HOW HIS THEORY COULD
BE USED TO EXPLAIN SPATIAL SPREAD OF A POPULATION 7HAT BETTER WAY TO STUDY THE
DYNAMICS OF A SPREADING POPULATION THEN WITH NEWLY ARRIVING SPECIES 4HE
INTRODUCTION AND SPREAD OF THE MUSKRAT /NDATRA ZIBETHICA IN CENTRAL %UROPE WAS
AN ILLUSTRATIVE EXAMPLE FOR 3KELLAM 3KELLAM WAS ABLE TO ADD POPULATION
GROWTH TERMS EITHER EXPONENTIAL OR LOGISTIC TO SPREAD MODELS !DDING EXPONEN
TIAL GROWTH THE CHANGE IN DENSITY IS
,N , N
RN $
,T ,X
WHERE N IS THE POPULATION DENSITY AND LOCATION X WITH GROWTH RATE R AND DIFFUSIVITY
$ &OLLOWING +OT THE VELOCITY OF SPREAD IS
X
R$
T
!DDING LOGISTIC GROWTH THE CHANGE IN DENSITY IS
,N N ,N
RN n $
,T + ,X
$ARWIN TO %LTON EARLY ECOLOGY AND THE PROBLEM OF INVASIVE SPECIES
!GAIN FOLLOWING +OT THE VELOCITY OF SPREAD IS
DX
R$
DT
Skellams spatial spread models have been used to understand the spread
OF NATURAL POPULATIONS EG /VASKAINEN AND ESPECIALLY FOR THE SPREAD OF
POPULATIONS OF INVASIVE SPECIES EG (ASTINGS +OT ET AL "UCHAN AND
0ADILLA
'%.%2!, 42%.$3 ). %#/,/'9 50 4/ %,4/.
3IMULTANEOUSLY WITH #LEMENTS EARLIER WORK ANOTHER PLANT ECOLOGIST 6OLNEY
3PALDING PUBLISHED A MONOGRAPH ON THE DISTRIBUTION OF DESERT PLANTS (E
SHOWED THAT COMPETITION AND MIGRATION ARE THE MOST IMPORTANT FACTORS STRUCTURING
DESERT COMMUNITIES (IS STUDIES SHOWED THAT CHANGE IS CONSTANTLY TAKING PLACE AND
INVASION IS A CONSTANT PROCESS 4HIS INVASION PROCESS HE NOTED IS BY BOTH NATIVE
AND EXOTIC SPECIES AND THE EXOTICS ARE NO DIFFERENT EXCEPT THAT THEY ARE JUST PART OF
THE LOCAL mORA OWING TO hTHE HAND OF -ANv P 4HEREFORE THE SAME PROCESS
SUCCESSION OCCURS REGARDLESS OF WHICH SPECIES AND WHERE THEY CAME FROM
!NIMAL ECOLOGY HAD A LATER START THAN PLANT ECOLOGY AND SEEMS TO HAVE FOLLOWED
ITS LEAD -C)NTOSH BUT A QUICK mURRY OF TEXTS MADE UP FOR PLANT ECOLOGYS
HEAD START !RTHUR 0EARSES !NIMAL %COLOGY DElNITELY TOOK ITS LEAD FROM
PLANT ECOLOGY ON THE NATURE OF CHANGE hSUCCESSION IS USUALLY DUE TO ONE OF TWO
CAUSES THE ENVIRONMENT IS CHANGING OR THE INCREASING GROUP OF ANIMALS IS
BETTER ABLE TO TAKE ADVANTAGE OF EXISTING CONDITIONSv P 4HERE IS NO MEN
TION OF WHERE THE BETTER ADAPTED ANIMALS COME FROM MUCH LESS OF THE OVERARCHING
IMPORTANCE OF DISPERSAL FOR SUCCESSION
3IMILARLY #HARLES %LTONS lRST BOOK !NIMAL %COLOGY NOTES THAT ANIMAL
DISPERSAL IS IMPORTANT WITH SOME MOVING VOLUNTARILY AND OTHERS THROUGH CHANCE
9ET THERE IS NO MENTION OF .)3 OR THEIR IMPACTS -OREOVER 2OYAL #HAPMANS
lTTINGLY TITLED !NIMAL %COLOGY WAS THE lRST TO EXAMINE EXPLICITLY THE EFFECTS
OF MIGRATION ON ANIMAL COMMUNITIES AND THE CAUSES OF STABILITY (E EXPOUNDED ON
TWO TYPES OF RESISTANCE TO INVASIONS AND POPULATION EXPLOSIONS 4HE lRST WAS PHYS
ICAL RESISTANCE WHERE THE ABIOTIC ENVIRONMENT MUST BE OPTIMAL FOR THE POPULATION
IN QUESTION 4HE SECOND HE CALLED BIOTIC RESISTANCE IN WHICH POPULATIONS ARE KEPT
IN CHECK THROUGH COMPETITION PREDATION AND PARASITES -ORE DIVERSE COMMUNI
TIES WOULD THEREFORE BE MORE RESISTANT TO INVADERS !S WITH THE OTHER AUTHORS
THERE IS NO EXPLICIT DISCUSSION OF WHERE INVADERS COME FROM AND THEIR IMPACTS
4HERE WAS LITTLE ADVANCE IN THE lELD OF PLANT ECOLOGY LEADING UP TO %LTONS
INVASION BOOK &OR EXAMPLE -C$OUGALLS 0LANT %COLOGY READS LIKE
AN UPDATED VERSION OF #LEMENTS 4O -C$OUGALL SUCCESSION HAS BOTH ABIOTIC AND
BIOTIC CAUSES AND THE BIOTIC MECHANISM IS INVASION WHICH hMEANS THE MOVEMENT
-7 #ADOTTE
OF PLANTS FROM ONE AREA INTO ANOTHER AND THEIR COLONIZATION IN THE LATTERv P
3OUNDS FAMILIAR
)N 'OODS 4HE 'EOGRAPHY OF &LOWERING 0LANTS HE SAYS MORE ABOUT .)3
INVADERS THAN HIS PREDECESSORS (E NOTES THAT WHEREVER HUMANS HAVE GONE SO
HAVE .)3 (E MAKES SOME PREDICTIONS AS TO WHY WE OUGHT NOT TO WORRY ABOUT
.)3 7HEN IT COMES TO COMPETITION hTHE STRUGGLE OF ;EXOTIC= VERSUS NATIVE IS MUCH
LESS IN FAVOUR OF THE FORMERv P &URTHERMORE .)3 ARE RELEGATED TO DISTURBED
GROUND BECAUSE OF THIS COMPETITIVE INFERIORITY 7HERE .)3 DO SPREAD AND BECOME
ABUNDANT AS DO A NUMBER OF FRESHWATER SPECIES THEY hTEND AFTER A TIME TO SETTLE
DOWN TO A MORE REASONABLE SCALE OF EXISTENCEv P 'OODS VIEW IS INTERESTING
AS HE DOES USE PREDICTION BASED ON ECOLOGICAL REASONING TO ADDRESS INVASIVE SPE
CIES (OWEVER WE MUST QUESTION WHETHER HE WAS ACTUALLY ADDRESSING WHAT WAS
BEING OBSERVED WITH INVASIVE .)3 OR IF HE WAS SIMPLY TRYING TO lT WHAT SHOULD
HAPPEN INTO HIS ECOLOGICAL WORLDVIEW
).#)0)%.4 ).6!3)/.3 ")/,/'9
)T REALLY WASNT UNTIL THE MID S THAT STUDIES WHICH WE CAN REFER TO AS INVA
SION BIOLOGY BEGAN TO SURFACE !S LATE AS 3TORER .)3 INVADERS WERE
NOT CONSIDERED TO BE A MAJOR SOURCE OF CONCERN FOR MAINTAINING THE NATURAL
ENVIRONMENT 4RACY 3TORER WRITING ABOUT FACTORS AFFECTING THE WILDLIFE OF
#ALIFORNIA LISTED THE MAJOR FACTORS AS AGRICULTURE GRAZING BY CATTLE MANIPULA
TION OF HYDROLOGY REMOVAL OF PREDATION DEFORESTATION AND THE CHANGING OF lRE
REGIMES &URTHER HE NOTED THAT GOVERNMENT AND PROFESSIONAL ORGANIZATIONS WERE
ACTIVELY MANAGING WILDLIFE BY REGULATING HUNTING MEANS AND SEASONS PROTECTING
RESERVES AND REFUGES REMOVING PREDATORS AND PROPAGATING AND RELEASING YOUNG
OF PARTICULAR SPECIES (E SHOWED NO CONCERN ABOUT .)3 OR THEIR CONTROL
( ( !LLEN GAVE SERIOUS AND ACADEMIC ATTENTION TO THE PROBLEM OF
.)3 IN .EW :EALAND 7HEREAS AUTHORS LIKE (OOKER AND 4ANSLEY
THOUGHT THAT INVADERS AS A GROUP CHANGE A mORA WITH UNIFORM SUCCESS !LLEN
SHOWED THAT THE .)3 WERE NOT A HORDE OF COMPETITIVE INVADERS )N FACT
RELATIVELY FEW .)3 OF SPECIES BECAME PROBLEM SPECIES AND THESE FEW
PROBLEM SPECIES REPRESENTED A DIVERSE ARRAY OF ECOLOGIES &URTHER !LLEN THOUGHT
THAT IF HUMANS WERE TO CEASE CLEARING AND GRAZING LAND NATIVE COMMUNITIES
WOULD RETURN AND REPLACE THE NOVEL COMMUNITIES
3IMILARLY %GLER THOUGHT .)3 INVADERS WERE NOT A UNIFORM GROUP
CHANGING THE ECOLOGY OF (AWAII 2ATHER HE SAW .)3 AS A DIVERSE GROUP WHICH
MADE PREDICTING THE FUTURE OF COMMUNITY TYPES NEARLY IMPOSSIBLE (E SHOWED
THAT MANY FACTORS AFFECT THE SUCCESS OF .)3 AND WHETHER THESE FACTORS WERE
GRAZING ANTHROPOGENIC DISTURBANCES OR SPECIES INTERACTIONS THE FATE OF ANY ONE
.)3 REMAINED FOGGY ,IKE !LLEN %GLER THOUGHT THAT AFTER GRAZING
AND DISTURBANCES WERE REMOVED NATIVE COMMUNITIES WOULD BE MUCH BETTER SUITED
TO COMPETE AGAINST .)3
$ARWIN TO %LTON EARLY ECOLOGY AND THE PROBLEM OF INVASIVE SPECIES
4HE LATE S AND EARLY S HERALDED THE ARRIVAL OF STUDIES OF THE ECOLOGY
OF INVASIONS 4HESE TYPES OF STUDIES SPANNED THE BIOLOGY OF .)3 IN THEIR NEW ENVI
RONMENTS TO SPECIES INTERACTIONS TO CONTROL 3TEWART AND (ALL DEPICTED THE
BIOLOGY OF THE .)3 CHEATGRASS AND HOW IT WAS AFFECTING THE NATIVE COMMUNITIES
OF SOUTHERN )DAHO "AKER USED THE INVASIONS OF -ELANDRIUM ALBUM AND
- DIOICUM IN 'REAT "RITAIN IN ORDER TO STUDY THE DYNAMICS OF INVASION AND RANGE
EXPANSION AS WELL AS LARGE
SCALE COMPETITION &INALLY THE CONTROL OF PROBLEMATIC
.)3 WAS GAINING MOMENTUM BECAUSE OF FAMOUS EARLY INSTANCES OF SUCCESSFUL BIO
LOGICAL CONTROL 4HESE EXAMPLES WOULD INCLUDE NOT ONLY THE CONTROL OF /PUNTIA
CACTI IN !USTRALIA BY #ACTOBLASTIS CACTORUM BY 'ROVES BUT ALSO THE
CONTROL OF (YPERICUM PERFORATUM BY #HRYSOMELA BEETLES IN WESTERN 5NITED 3TATES
(UFFAKER AND (OLLOWAY (UFFAKER
7(!4 ()34/29 4%,,3 !"/54 %#/,/'9 !.$ ).6!3)/. ")/,/'9
) AM PAINTING THE PICTURE THAT THE NEW SCIENCE OF ECOLOGY IN THE EARLY TH CEN
TURY WAS DEVELOPING ITS EPISTEMOLOGICAL DOMAIN AND THE QUESTION IS DID THEY
TRY TO USE THEIR NEW SCIENCE TO UNDERSTAND INVASIVE EXOTICS ) THINK THEY DID NOT
4HESE EARLY ECOLOGISTS SEEMED TO INCORPORATE .)3 BECAUSE THEY YIELDED REAL
WORLD
OBSERVATIONS WHICH OBEYED THE VERY NATURAL LAWS THESE AUTHORS WERE EXPLORING
&URTHERMORE .)3 WERE IN SOME CASES USED TO SHOWCASE THEIR IDEAS ABOUT NATURAL
LAWS OR AS NATURAL EXPERIMENTS "Y AND LARGE THE EARLY ECOLOGISTS DID NOT USE THEIR
CONCEPTUAL IDEAS TO UNDERSTAND AND PREDICT THE PROBABLE OUTCOMES OF SPECIES INVA
SIONS 4HEY DID NOT OFTEN IMPLEMENT 'AUSES SUGGESTION THAT INVASIONS
CONSTITUTE NATURAL EXPERIMENTS TO UNDERSTAND ECOLOGY BETTER
/F COURSE THIS REVIEW DOES NOT SCOUR THE LITERATURE FOR EVERY RECORDED IDEA OR
EVERY DISCUSSION OF .)3 INVADER 2ATHER ) HAVE ATTEMPTED TO lND THE PROMINENT
MODES OF THINKING IN ECOLOGY AND ESPECIALLY WITH RESPECT TO .)3 INVADERS ) HAVE
RELIED EXTENSIVELY ON TEXTS AS RECORDS OF THESE TRENDS AND A VALID CRITICISM IS THAT
GENERAL TEXTS DO NOT CAPTURE ALL THE DIVERSITY OF THINKING WITHIN A lELD OF STUDY
BOTH SUBTLE AND STARK (OWEVER %GLER IN HIS CRITICISM OF THE EPISTEMOL
OGY OF !MERICAN PLANT ECOLOGY DURING THE S FOUND THAT THE MAJOR TEXTS DID
LARGELY REPRESENT THE SCHOOLS OF THOUGHT AND PHILOSOPHIES OF !MERICAN ECOLOGY
$AVIS ET AL ASSOCIATE %LTONS INVASIONS BOOK WITH THE CAUSE OF
THE DISSOCIATION BETWEEN ECOLOGY AND INVASIONS BIOLOGY )F WE THINK OF ECOLOGY AS
A SCIENCE OF CASE STUDIES EG 3HRADER
&RECHETTE AND -C#OY THEN PER
HAPS THERE WOULD BE NO DISSOCIATION AND BY STUDYING INDIVIDUAL CASES OF SPECIES
INVASIONS WE CAN THEN FORM LARGER PICTURES OF WHAT IS HAPPENING IN NATURE
3HRADER
&RECHETTE 9ET MUCH OF ECOLOGY IS BASED ON THE PRECEPT THAT
THERE ARE FUNDAMENTAL OPERATIONS IN NATURE 4HESE FUNDAMENTAL OPERATIONS ARE
APPROXIMATED THROUGH THEORY CONSTRUCTION EXPERIMENTATION AND OBSERVATION
4HE DISSOCIATION SEEN BY $AVIS ET AL IS THAT THESE SCIENTIlC EXERCISES
THEORY EXPERIMENT AND OBSERVATION ARE WHAT BOTH INVASIONS BIOLOGISTS AND
-7 #ADOTTE
ECOLOGISTS DO BUT THEY DO IT LARGELY IN ISOLATION FROM ONE ANOTHER !LSO THESE TWO
SCIENTIlC ENTERPRISES THOUGH DRIVEN BY SOMETIMES SEPARATE GOALS COULD BENElT
FROM USE OF EACH OTHERS IDEAS TECHNIQUES AND OBSERVATIONS
(OWEVER AFTER REVIEWING MUCH OF THE EARLY ECOLOGY LITERATURE ) lND THIS DIS
SOCIATION SEEMS TO HAVE A MUCH LONGER PEDIGREE LONG BEFORE %LTON -ANY EARLY
ECOLOGISTS DO NOT SEEM TO APPLY THEIR SCIENCE TO APPLIED PROBLEMS 4HAT IS NOT TO
SAY THAT MANY WERE NOT CONCERNED ABOUT HUMAN IMPACTS ON THE ENVIRONMENT
BUT THAT THEY USED THEIR SCIENCE LITTLE IN THESE CONCERNS
&OR ONE THING THERE WAS NO MORAL CALL FOR ECOLOGISTS TO SOLVE THE PROBLEMS OF
THE NATURAL WORLD 4HE MODERN CONCERN FOR THE ENVIRONMENT APPEARED IN THE
WORK OF NATURALISTS AND ECOLOGISTS IN THE LATER S AND EARLY S EG %LTON
#ARSON IT IS NOW COMMONPLACE 4HE SOCIETAL IMPETUS WAS NOT THERE
&UNDING AGENCIES SIMPLY DID NOT EXIST MUCH LESS SUPPORT RESEARCH PROGRAMMES
PURPORTING TO ADDRESS APPLIED PROBLEMS
-ORE FUNDAMENTALLY THE FOUNDATIONS OF ECOLOGY LIKE THE FOUNDATIONS OF ANY
SCIENTIlC lELD SEEMED LAID BY THOSE SEEKING GENERAL EXPLANATIONS 4HAT IS NOT TO
SAY THAT THERE WERE NOT AUTHORS LOOKING AT VERY SPECIlC PROBLEMS BUT ECOLOGY
TEXTBOOKS RELAY THE GENERAL IDEAS THAT SEEM BEST SUITED TO EXPLAINING NATURAL
PHENOMENA -ANY OF THESE EARLY AUTHORS ADDRESS ISSUES INTIMATELY LINKED TO
UNDERSTANDING SPECIES INVASIONS EG DISPERSAL SUCCESSION CHANGE ETC 9ET
THEIR CONSIDERATIONS OF NON NATIVE SPECIES INVOLVED USING THESE SPECIES AS EFlCA
CIOUS TESTS OF THEIR IDEAS
#/.#,53)/.
(ERE ) HAVE EVER SO BRIEmY REVIEWED SOME OF THE MAJOR WORKS OF THE SCIENCE OF
ECOLOGY IN THE HUNDRED OR SO YEARS BEFORE %LTON PUBLISHED 4HE %COLOGY OF )NVASIONS
BY !NIMALS AND 0LANTS IN )N REVIEWING THESE ) HAVE FOCUSED PARTICULARLY ON
THE WAY IN WHICH THE DEVELOPMENT OF IDEAS IN ECOLOGY PERTAIN TO UNDERSTANDING
THE PHENOMENON OF SPECIES INVASIONS #OUPLED WITH THIS HISTORICAL EXAMINATION
) HAVE CONSIDERED THE NOTION THAT %LTONS WORK CAUSED DISSOCIATION BETWEEN
ECOLOGY AND INVASION BIOLOGY $AVIS ET AL 4O BE SURE THERE WAS NO INDE
PENDENT lELD OF INVASION BIOLOGY IN THE EARLY YEARS OF ECOLOGY BEYOND AGRICULTURAL
AND ECONOMIC CONCERNS POSED BY SOME NON
INDIGENOUS SPECIES 4HE CURRENT lELD
OF INVASION BIOLOGY OWES MUCH TO %LTONS BOOK 4HIS REVIEW SHOWS THAT EARLY
ECOLOGISTS DID ADDRESS SPECIES INVASIONS BECAUSE THEY WE A REAL AND EVIDENT PHE
NOMENA AND ANY GENERAL ACCOUNT ON THE FUNCTIONING OF BIOLOGICAL COMMUNITIES
WOULD NEED TO BE ABLE TO EXPLAIN WHY SOME SPECIES REACH NEW LANDS AND SURVIVE
AND GROW 3OME OF THESE EARLY ECOLOGISTS SAW INVADERS AS A WAY TO TEST THEORY
9ET THE CONCERN AND ENERGY DEVOTED TO UNDERSTANDING INVADERS WENT LITTLE BEYOND
EXPLAINING AND USE AS EXAMPLE AND IT REALLY WASNT UNTIL THE MID TO LATE S
THAT WE SEE ANYTHING RESEMBLING INVASION BIOLOGY
$ARWIN TO %LTON EARLY ECOLOGY AND THE PROBLEM OF INVASIVE SPECIES
4HE CURRENT TREND TO UNIFY MAINSTREAM ECOLOGY AND INVASION BIOLOGY IS PRO
CEEDING AS THE CURRENT VOLUME ATTESTS EXPLORING NEW GROUND AND PUSHING THE
LIMITS OF WHAT HAS COME BEFORE 2ATHER THAN LOOK AT INVASION BIOLOGY AS A SEPARATE
lELD IT MAY BE BETTER TO CONSIDER IT A NECESSARY CONDITION TO GENERATING AN ACCU
RATE VIEW OF NATURE ONE THAT ECOLOGY MUST BE ABLE TO USE &URTHER THE PLETHORA OF
CASE STUDIES OF INVADERS SHOULD BE INVALUABLE TO THE ENTERPRISE OF TRYING TO SYNTHE
SIZE NATURAL PHENOMENA INTO A HOLISTIC FRAMEWORK 0ERHAPS CONCEPTS IN ECOLOGY
ARE ADEQUATE IN THEIR CURRENT DEVELOPMENT BUT LEARNING FROM THE WAYS IN WHICH
INVADERS RESPOND TO NEW ENVIRONMENTS AND CHANGE THE COMMUNITIES THEY ARE
INVADING CAN TELL US MORE ABOUT NATURE THAN MOST OTHER EXERCISES
!#+./7,%$'%-%.43
) AM TRULY STANDING ON THE SHOULDERS OF GIANTS AND ) HOPE THAT MY PERCEPTION OF
THE HISTORY OF ECOLOGY AND INVASION BIOLOGY IS AN ACCURATE ONE LEST THOSE GIANTS
TURN IN THEIR GRAVES ) AM INDEBTED TO *IM $RAKE $AN 3IMBERLOFF -ARK $AVIS
3EAN -C-AHON 4AD &UKAMI FOR READING AND EARLIER VERSION OF THIS CHAPTER /F
COURSE ANY INACCURACIES ARE NO FAULT OF THEIRS
2%&%2%.#%3
!LLAN ( ( )NDIGENE VERSUS ALIEN IN THE .EW :EALAND PLANT WORLD %COLOGY
"AKER ( ' 3TAGES IN INVASION AND REPLACEMENT DEMONSTRATED BY SPECIES OF
-ELANDRIUM *OURNAL OF %COLOGY
"UCHAN , ! * AND $ + 0ADILLA %STIMATING THE PROBABILITY OF LONG
DISTANCE OVER
LAND DISPERSAL OF INVADING AQUATIC SPECIES %COLOGICAL !PPLICATIONS
#ANDOLLE DE ! , 0 'OGRAPHIE "OTANIQUE RAISSON 6 -ASSON 0ARIS
#ARSON 2 3ILENT 3PRING (OUGHTON -IFmIN "OSTON -!
#HAPMAN 2 . !NIMAL %COLOGY -C'RAW
(ILL "OOK #OMPANY )NC .EW 9ORK
.9
#HASE * - AND - ! ,EIBOLD %COLOGICAL .ICHES 4HE 5NIVERSITY OF #HICAGO 0RESS
#HICAGO ),
#LEMENTS & % 2ESEARCH -ETHODS IN %COLOGY 5NIVERSITY 0UBLISHING #O ,INCOLN
.%
#LEMENTS & % 0LANT 0HYSIOLOGY AND %COLOGY (ENRY (OLT AND #OMPANY .EW 9ORK
.9
#LEMENTS & % AND 6 % 3HELFORD "IO
ECOLOGY *OHN 7ILEY 3ONS )NC .EW 9ORK
.9
#ONNER % & AND $ 3 3IMBERLOFF 4HE ASSEMBLY OF SPECIES COMMUNITIES #HANCE OR
COMPETITION %COLOGY
#OURCHAMP & - ,ANGLAIS AND ' 3UGIHARA #ATS PROTECTING BIRDS MODELING THE
MESOPREDATOR RELEASE EFFECT *OURNAL OF !NIMAL %COLOGY
-7 #ADOTTE
#ROKER 2 ! 3TEPHEN &ORBES AND THE 2ISE OF !MERICAN %COLOGY 3MITHSONIAN
)NSTITUTION 0RESS 7ASHINGTON $#
$AEHLER # # $ARWINS NATURALIZATION HYPOTHESIS REVISITED !MERICAN .ATURALIST
$!NCONA 5 4HE 3TRUGGLE FOR %XISTENCE % * "RILL ,EIDEN 4HE .ETHERLANDS
$ARWIN # 4HE /RIGIN OF THE 3PECIES * -URRAY ,ONDON
$AVIS - ! + 4HOMPSON AND * 0 'RIME #HARLES 3 %LTON AND THE DISSOCIATION OF
INVASION ECOLOGY FROM THE REST OF ECOLOGY $IVERSITY AND $ISTRIBUTIONS
$RUDE / -ANUEL DE 'EOGRAPHIE "OTANIQUE ,IBRAIRIE $ES 3CIENCES .ATURELLES
0ARIS
%GERTON & . %COLOGICAL STUDIES AND OBSERVATIONS BEFORE 0AGES
IN
" * 4AYLOR AND 4 * 7HITE EDITORS )SSUES AND )DEAS IN !MERICA 5NIVERSITY OF /KLAHOMA
0RESS /KLAHOMA /+
%GLER & % )NDIGENE VERSUS ALIEN IN THE DEVELOPMENT OF ARID (AWAIIAN VEGETATION
%COLOGY
%GLER & % ! COMMENTARY ON !MERICAN PLANT ECOLOGY BASED ON THE TEXTBOOKS OF
%COLOGY
%HRLICH 0 2 (UMAN NATURES NATURE CONSERVATION AND ENVIRONMENTAL ETHICS
"IOSCIENCE
%LTON # 3 !NIMAL %COLOGY 4HE -AC-ILLAN #OMPANY .EW 9ORK .9
%LTON # 3 4HE %COLOGY OF )NVASIONS BY !NIMALS AND 0LANTS -ETHUEN ,ONDON
%MERSON 2 7 4HE %ARLY ,ECTURES
EDITED BY 3 % 7HICHER AND
2 % 3PILLER (ARVARD 5NIVERSITY 0RESS #AMBRIDGE -!
&ITCH ! 3IXTH REPORT ON THE NOXIOUS AND OTHER INSECTS OF THE STATE OF .EW 9ORK .EW
9ORK 3TATE !GRICULTURAL 3OCIETY 4RANSACTIONS
&ORBES 3 ! )NSECTS AFFECTING CORN )LLINOIS 3TATE %NTOMOLOGY /FlCE #IRCULATION
&ORBES 3 ! 4HE CLINCH
BUG IN )LLINOIS )LLINOIS 3TATE %NTOMOLOGY /FlCE #IRCULATION
PP 3EPTEMBER
&ORBES 3 ! A !RSENICAL POISONS FOR THE CODLING MOTH RECORD AND DISCUSSION OF
EXPERIMENTS FOR AND 4RANSACTIONS OF THE )LLINOIS 3TATE (ORTICULTURAL 3OCIETY
&ORBES 3 ! B 4HE LAKE AS A MICROCOSM "ULLETIN OF THE 3CIENTIlC !SSOCIATION 0EORIA
),
&ORBES 3 ! /N THE PRINCIPLE CORN INSECTS AND METHODS OF COLLECTING THEM )LLINOIS
!GRICULTURALIST PP
&ORBES 3 ! 4HE SEASONS CAMPAIGN AGAINST THE 3AN *OS AND OTHER SCALE INSECTS IN
)LLINOIS 4RANSACTIONS OF THE )LLINOIS 3TATE (ORTICULTURAL 3OCIETY
'AUSE ' & 4HE 3TRUGGLE FOR %XISTENCE (AFNER 0UBLISHING #OMPANY
.EW 9ORK .9
'OOD 2 4HE 'EOGRAPHY OF THE &LOWERING 0LANTS ,ONGMANS 'REEN AND #O
.EW 9ORK .9
'RINNELL * 4HE NICHE RELATIONSHIPS OF THE #ALIFORNIA THRASHER !UK
'RINNELL * 2ISKS INCURRED IN THE INTRODUCTION OF ALIEN GAME BIRDS 3CIENCE
$ARWIN TO %LTON EARLY ECOLOGY AND THE PROBLEM OF INVASIVE SPECIES
'ROVES 2 ( %COLOGICAL CONTROL OF INVASIVE TERRESTRIAL PLANTS 0AGES
IN
* ! $RAKE ( ! -OONEY & DI #ASTRI 2 ( 'ROVES & * +RUGER - 2EJMANEK AND
- 7ILLIAMSON EDITORS "IOLOGICAL )NVASIONS ! 'LOBAL 0ERSPECTIVE *OHN 7ILEY AND
3ONS .EW 9ORK .9
(ALLAM 4 ' #OMMUNITY DYNAMICS IN A HOMOGENEOUS ENVIRONMENT 0AGES
IN 4 ' (ALLAM AND 3 ! ,EVIN EDITORS -ATHEMATICAL %COLOGY 3PRINGER
6ERLAG
"ERLIN
(ARDIN ' 4HE COMPETITIVE EXCLUSION PRINCIPLE 3CIENCE
(ASTINGS ! -ODELS OF SPATIAL SPREAD IS THE THEORY COMPLETE %COLOGY
(ECK + , 3OME CRITICAL CONSIDERATIONS OF THE THEORY OF SPECIES PACKING %VOLUTIONARY
4HEORY
(EILPRIN ! 4HE 'EOGRAPHICAL AND 'EOLOGICAL DISTRIBUTION OF !NIMALS $ !PPLETON
#O .EW 9ORK .9
(OOKER * $ (ANDBOOK OF THE .EW :EALAND &LORA 6OLUME 3IR *OSEPH $ALTON
(OOKER ,ONDON
(OWARD , / )NSECTS AFFECTING THE COTTON PLANT "ULLETIN OF THE /FlCE OF %XPERIMENTAL
3TATIONS
(OWARD , / 4HE mY WEEVIL !DDRESS AT THE &ARMERS )NSTITUTE -ANASSAS 6A
(OWARD , / 4HE SHADE
TREE INSECT PROBLEM IN THE EASTERN 5NITED 3TATES 9EARBOOK
OF THE 5 3 $EPARTMENT OF !GRICULTURE
(OWARD , / A 4HE SPREAD OF LAND SPECIES BY THE AGENCY OF -AN WITH ESPECIAL
REFERENCE TO INSECTS 0ROCEEDINGS OF THE !MERICAN !SSOCIATION FOR THE !DVANCEMENT OF
3CIENCE
(OWARD , / B $ANGER OF IMPORTING INSECT PESTS 9EARBOOK OF $EPARTMENT OF
!GRICULTURE
(OWARD , / 4HE PRINCIPAL INSECTS AFFECTING THE TOBACCO PLANT 9EARBOOK OF
$EPARTMENT OF !GRICULTURE
(OWARD , / 4HREE INSECT ENEMIES OF SHADE TREES 5 3 $EPARTMENT OF !GRICULTURE
&ARMERS "ULLETIN
(UFFAKER # " 4HE RETURN OF NATIVE PERENNIAL BUNCHGRASS FOLLOWING THE REMOVAL OF
+LAMATH WEED (YPERICUM PERFORATUM , BY IMPORTED BEETLES %COLOGY
(UFFAKER # " AND * + (OLLOWAY #HANGES IN RANGE PLANT POPULATION STRUCTURE
ASSOCIATED WITH FEEDING OF IMPORTED ENEMIES OF +LAMATH WEED Hypericum perforatum
, %COLOGY
(USTON - ! "IOLOGICAL $IVERSITY #AMBRIDGE 5NIVERSITY 0RESS #AMBRIDGE
(UTCHINSON ' % #ONCLUDING REMARKS #OLD 3PRING (ARBOR 3YMPOSIA ON 1UANTITATIVE
"IOLOGY
*OHNSTONE ) - 0LANT INVASIONS WINDOWS A TIME
BASED CLASSIlCATION OF INVASION
POTENTIAL "IOLOGICAL 2EVIEW
+IERSTEAD 3 % AND , " 3LOBODKIN 4HE SIZE OF WATER MASSES CONTAINING PLANKTON
BLOOMS *OURNAL OF -ARINE 2ESEARCH
+OT - %LEMENTS OF -ATHEMATICAL %COLOGY #AMBRIDGE 5NIVERSITY 0RESS .EW 9ORK
.9
-7 #ADOTTE
+OT - - ! ,EWIS AND 0 VAN DEN $RIESSCH $ISPERSAL DATA AND THE SPREAD OF INVAD
ING ORGANISMS %COLOGY
,AWTON * ( !RE THERE GENERAL LAWS IN ECOLOGY /IKOS
,EIBOLD - ! 4HE NICHE CONCEPT REVISITED MECHANISTIC MODELS AND COMMUNITY
CONTEXT %COLOGY
,OTKA ! * %LEMENTS OF 0HYSICAL "IOLOGY 7ILLIAMS AND 7ILKINS "ALTIMORE -$
-ACK 2 . 0LANT NATURALIZATIONS AND INVASIONS IN THE EASTERN 5NITED 3TATES
!NNALS OF THE -ISSOURI "OTANICAL 'ARDEN
-C$OUGALL 7 " 0LANT %COLOGY ,EA AND &EBIGER 0HILADELPHIA 0!
-C)NTOSH 2 0 %COLOGY SINCE 0AGES IN " * 4AYLOR AND 4 * 7HITE
EDITORS )SSUES AND )DEAS IN !MERICA 5NIVERSITY OF /KLAHOMA 0RESS /KLAHOMA /+
-ENAND , 4HE -ETAPHYSICAL #LUB ! 3TORY OF )DEAS IN !MERICA &ARRAR 3TRAUS
'IROUX .EW 9ORK .9
/LMSTED & , ,ETTER TO THE EDITOR &OREIGN PLANTS AND !MERICAN SCENERY 'ARDEN AND
&OREST
/WENS 2 $ARWIN ON THE ORIGIN OF SPECIES 0AGES IN 0 !PPLEMAN
EDITOR $ARWIN ! .ORTON #RITICAL %DITION THIRD EDITION 7 7 .ORTON #OMPANY
.EW 9ORK .9
0EARSE ! 3 !NIMAL %COLOGY -C'RAW (ILL "OOK #OMPANY )NC .EW 9ORK .9
0ETERS 2 ( ! #RITIQUE FOR %COLOGY #AMBRIDGE 5NIVERSITY 0RESS .EW 9ORK .9
3ARGENT # 3 %DITORIAL 'ARDEN AND &OREST
3EDGWICK ! /BJECTIONS TO -R $ARWINS THEORY OF THE ORIGIN OF SPECIES
0AGES IN 0 !PPLEMAN EDITOR $ARWIN ! .ORTON #RITICAL %DITION THIRD EDITION
7 7 .ORTON #OMPANY .EW 9ORK .9
3HANTZ ( , ! STUDY OF THE VEGETATION OF THE -ESA REGION EAST OF 0IKES 0EAK
"OTANICAL 'AZETTE
3HRADER &RECHETTE + .ON INDIGENOUS SPECIES AND ECOLOGICAL EXPLANATION "IOLOGY
AND 0HILOSOPHY
3HRADER &RECHETTE + AND % $ -C#OY !PPLIED ECOLOGY AND THE LOGIC OF CASE STUDIES
0HILOSOPHY OF 3CIENCE
3IMBERLOFF $ 3 5SING ISLAND BIOGEOGRAPHIC DISTRIBUTIONS TO DETERMINE IF COLONIZA
TION IS STOCHASTIC !MERICAN .ATURALIST
3IMBERLOFF $ #ONFRONTING INTRODUCED SPECIES A FORM OF XENOPHOBIA "IOLOGICAL
)NVASIONS
3KELLAM * ' 2ANDOM DISPERSAL IN THEORETICAL POPULATIONS "IOMETRIKA
3PALDING 6 - $ISTRIBUTION AND -OVEMENTS OF $ESERT 0LANTS #ARNEGIE )NSTITUTE OF
7ASHINGTON 7ASHINGTON $#
3TEWART ' AND ! # (ULL #HEATGRASS "ROMUS TECTORUM , AN ECOLOGICAL INTRUDER
IN SOUTHERN )DAHO %COLOGY
3TORER 4 ) &ACTORS INmUENCING WILD LIFE IN #ALIFORNIA PAST AND PRESENT %COLOGY
4ANSLEY ! ' 4HE USE AND ABUSE OF VEGETATIONAL CONCEPTS AND TERMS %COLOGY
$ARWIN TO %LTON EARLY ECOLOGY AND THE PROBLEM OF INVASIVE SPECIES
4HOMPSON + * ' (ODGSON AND 4 # ' 2ICH .ATIVE AND ALIEN PLANT INVASIVE
PLANTS MORE OF THE SAME %COGRAPHY
6OLTERRA 6 6ARIATIONS AND mUCTUATIONS IN THE NUMBERS OF COEXISTING
ANIMAL SPECIES 0AGES
IN & - 3CUDO AND * 2 :IEGLER EDITORS 4HE 'OLDEN !GE
OF 4HEORETICAL %COLOGY
3PRINGER
6ERLAG .EW 9ORK .9
7ALLACE ! 2 4HE 'EOGRAPHICAL $ISTRIBUTION OF !NIMALS (AFNER 0UBLISHING
#OMPANY .EW 9ORK .9
7ARMING % /ECOLOGY OF 0LANTS !N )NTRODUCTION TO THE 3TUDY OF 0LANT #OMMUNITIES
#LARENDON 0RESS /XFORD 5+
7EAVER * % AND & % #LEMENTS 0LANT %COLOGY -C'RAW
(ILL BOOK #OMPANY
.EW 9ORK .9
#HAPTER THREE
)NVASION BIOLOGY
THE PURSUIT OF SCIENCE
AND CONSERVATION
- ! $AVIS
).42/$5#4)/.
4HE HISTORY OF INVASION BIOLOGY WOULD BE A DREAM DISSERTATION TOPIC FOR SOME HIS
TORY OF SCIENCE GRADUATE STUDENT 4HE LIST OF RESEARCHERS WHO HAVE WEIGHED IN ON
THE TOPIC AT ONE TIME OR ANOTHER READS LIKE A 7HOS 7HO OF LATE TH CENTURY ECOL
OGY #ONTROVERSY AND DISAGREEMENTS OCCASIONALLY RESULTING IN LIVELY EXCHANGES
AMONG RESEARCHERS HAVE CREATED AN INTELLECTUALLY DYNAMIC AND SOMETIMES
EMOTIONALLY CHARGED ATMOSPHERE IN RECENT YEARS !UTHORS OF BOTH TECHNICAL AND
POPULAR ARTICLES HAVE OFTEN USED EVOCATIVE LANGUAGE AND IMAGERY IN THEIR WRITING
3OME OF THE ISSUES HAVE ATTRACTED THE INTERESTS OF SCHOLARS OUTSIDE THE BIOLOGICAL
SCIENCES PARTICULARLY PHILOSOPHERS !ND FOR THE PAST SEVERAL DECADES INVASION
ECOLOGY RESEARCH HAS BEEN CONDUCTED WITHIN A LARGER SOCIAL MILIEU OF CONTENTIOUS
ENVIRONMENTAL VALUES AND POLITICS .O DOUBT FOR ALL THESE REASONS INVASION ECOLO
GY HAS CAPTURED THE ATTENTION OF NATIONAL AND INTERNATIONAL MEDIA OUTLETS WHICH
HAVING THEIR OWN AGENDAS HAVE EMPHASIZED AND PRESENTED PARTICULAR RESEARCH
lNDINGS AND PERSPECTIVES FOR THEIR AUDIENCES
(OWEVER AN EXAMINATION OF THE lELDS HISTORY SHOULD BE OF INTEREST NOT JUST
TO HISTORIANS OF SCIENCE BUT TO ECOLOGISTS AS WELL !FTER ALL A LOOK BACKWARDS IS
-7 #ADOTTE ET AL EDS #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY n
3PRINGER 0RINTED IN THE .ETHERLANDS
-! $AVIS
ALMOST ALWAYS A GOOD IDEA BEFORE CHARTING NEW PATHS (OW HAVE RESEARCH QUES
TIONS CHANGED OVER TIME 7HAT HAVE WE LEARNED 7HAT CONTINUES TO CHALLENGE
OUR UNDERSTANDING )F WE CAN ANSWER THESE QUESTIONS WE CAN MORE EFFECTIVELY
FORMULATE FUTURE PRIORITIES ! HISTORICAL REVIEW CANNOT GUARANTEE ANSWERS TO
THESE QUESTIONS BUT NEITHER CAN THE ANSWERS BE OBTAINED WITHOUT SUCH A REVIEW
)N RESEARCHING AND WRITING THIS CHAPTER ) HAVE STRIVED TO PROVIDE A HISTORICAL
REVIEW OF INVASION BIOLOGY SINCE THE PUBLICATION DATE #HARLES %LTONS
INVASION CLASSIC 4HE %COLOGY OF )NVASIONS BY !NIMALS AND 0LANTS IN ORDER TO ANSWER
THE ABOVE QUESTIONS AND BASED ON WHATEVER INSIGHTS COULD BE GAINED FROM THESE
EFFORTS TO OFFER A FEW SUGGESTIONS AS TO HOW INVASION BIOLOGY MIGHT PROlTABLY
PROCEED FROM THIS POINT FORWARD
4(% 3
)N 4HE )NTERNATIONAL 5NION OF "IOLOGICAL 3CIENCES HELD ITS lRST "IOLOGICAL
3CIENCES 3YMPOSIUM PART OF WHAT WAS TO BE A SERIES OF SUCH MEETINGS DESIGNED TO
FOCUS ATTENTION ON BIOLOGICAL TOPICS HAVING INTERNATIONAL SIGNIlCANCE 4HE OBJEC
TIVE OF THE lRST SYMPOSIUM HELD IN !SILOMAR #ALIFORNIA WAS TO BRING TOGETHER
GENETICISTS ECOLOGISTS TAXONOMISTS AND APPLIED SCIENTISTS WORKING IN THE AREA
OF PEST CONTROL AND TO PRESENT AND DISCUSS FACTS AND IDEAS hABOUT THE KINDS OF
EVOLUTIONARY CHANGE WHICH TAKE PLACE WHEN ORGANISMS ARE INTRODUCED INTO NEW
TERRITORIESv 7ADDINGTON 4HE PROCEEDINGS WERE PUBLISHED IN A 4HE 'ENETICS
OF #OLONIZING 3PECIES "AKER AND 3TEBBINS NOW REGARDED AS A CLASSIC !S
WOULD BE EXPECTED GIVEN THE ORGANIZING GROUP THE BOOK REPRESENTED A DISTINCTLY
INTERNATIONAL INITIATIVE CONSISTING OF AUTHORS REPRESENTING COUNTRIES
4HE RANGE OF TOPICS CONSIDERED IN THIS VOLUME WAS BROADER THAN THE TITLE SUG
GESTS AND INCLUDED DISCUSSIONS ON THE NATURE OF THE COLONIZED ENVIRONMENTS AS
WELL AS THE ATTRIBUTES AND GENETICS OF COLONIZING SPECIES #HAPTERS ADDRESSED A
BROAD RANGE OF TAXA INCLUDING PLANTS INSECTS BIRDS MAMMALS AND MICROBES
4HE SYMPOSIUM AND THE BOOK WERE DElNED BY A STRONG EVOLUTIONARY EMPHASIS
AND PARTICIPANTS INCLUDED MANY OF THE PROMINENT EVOLUTIONARY BIOLOGISTS OF
THE TIME INCLUDING -AYR 3TEBBINS $OBZHANSKY 7ILSON % / #ARSON ,EWONTIN
AND 7ADDINGTON 3IGNIlCANTLY AUTHORS CONSIDERED COLONISTS BROADLY DISCUSSING
COLONIZATIONS THAT OCCURRED DURING SUCCESSION COLONIZATIONS OF WIDESPREAD SPE
CIES THAT HAD OCCURRED WITHOUT HUMAN ASSISTANCE COLONIZATIONS OF NATIVE WEEDS
AS WELL AS COLONIZATIONS OF SPECIES INTO NEW REGIONS OCCURRING AS A DIRECT RESULT OF
HUMAN ACTIVITY !LTHOUGH THE LATTER GROUP DID RECEIVE THE MOST ATTENTION PARTICI
PANTS RECOGNIZED THAT COMMON ECOLOGICAL AND EVOLUTIONARY PROCESSES UNDERLAY
THE DIFFERENT TYPES OF COLONIZATIONS
/NE NOTEWORTHY ASPECT OF THE BOOK IS THE STRIKING CONTRAST BETWEEN THE LAN
GUAGE USED BY THE AUTHORS AND THAT COMMONLY USED BY INVASION ECOLOGISTS TODAY
! CAREFUL SEARCH OF THE BOOKS TEXT IS NECESSARY TO lND TERMS SUCH AS @ALIEN
@EXOTIC @INVADER AND @INVASION -OST AUTHORS NEVER USED THESE WORDS ! FEW
)NVASION BIOLOGY
SUCH AS 7ILSON % / -AYR AND 7ODZICKI USED THEM OCCASIONALLY BUT THE ARTI
CLES AND DISCUSSIONS WERE OVERWHELMINGLY GUIDED BY NOUNS VERBS AND ADJECTIVES
SUCH AS @COLONIZERS @FOUNDING POPULATIONS @INTRODUCED @NON NATIVE @NEW
ARRIVALS @MIGRATION @SPREAD @GEOGRAPHICALLY WIDESPREAD 4HUS PARTICIPANTS
DID NOT ADOPT THE EFFUSIVE STYLE USED BY %LTON IN HIS BOOK CHARACTERIZED BY
THE FREQUENT USE OF METAPHOR AND ANALOGY OFTEN WITH EXPLICIT MILITARISTIC REFERENCES
4HE ONE EXCEPTION WAS %LTONS COLLEAGUE *OHN (ARPER WHO USUALLY DID REFER
TO THE NEW SPECIES AS @ALIENS AND @INVADERS AND WHOSE CHAPTER %STABLISHMENT
!GGRESSION AND #OHABITATION IN 7EEDY 3PECIES EXHIBITED SOME OF THE SAME EVOCATIVE
LANGUAGE THAT %LTON USED IN HIS BOOK 4HE OPENING SENTENCE OF (ARPERS CHAPTER IS
SO REMINISCENT OF THE LANGUAGE USED BY %LTON THAT THE LATTERS INmUENCE ON (ARPER
CAN HARDLY BE DENIED (ARPER BEGAN h4HE MOVEMENTS OF MAN AND HIS
GOODS HAVE RESULTED IN A BOMBARDMENT OF AREAS OF LAND AND SEA BY ALIEN SPECIES
BOTH BY CHANCE AND BY THE DELIBERATE INTRODUCTION OF CULTIVATED PLANTS OF THE FARM
AND GARDENv
)N THE PREFACE TO HIS BOOK %LTON SAID HIS GOAL WAS TO BRING TOGETHER THREE
STREAMS OF THOUGHT FAUNAL HISTORY ECOLOGY PARTICULARLY POPULATION ECOLOGY
AND CONSERVATION "UT THE CONSERVATION THEME DOMINATED THE BOOK (E OPENED
HIS BOOK WITH GRAPHIC BATTLElELD EXAMPLES OF INVASIONS AND ENDED THE BOOK WITH
TWO CHAPTERS ON THE NEED FOR CONSERVATION )N CONTRAST THE !SILOMAR PARTICIPANTS
DID NOT CONSIDER CONSERVATION IMPLICATIONS OF SPECIES COLONIZATIONS WHATSOEVER
% / 7ILSON PERSONAL COMMUNICATION 4HE CLEARLY DElNED AGENDA OF THE
!SILOMAR SYMPOSIUM WAS THE SEARCH FOR GENERALIZATIONS REGARDING THE EVOLUTION
ARY AND ECOLOGICAL PROCESSES INVOLVED IN SPECIES COLONIZATIONS )N THE CHAPTERS
AND PAGES OF THE SYMPOSIUM PROCEEDINGS %LTONS INVASION CLASSIC WAS
CITED ONLY THREE TIMES ONCE EACH BY "IRCH -AYR AND 7ILSON % /
!S A PUBLICATION FOCUSING ON INTRODUCED SPECIES 4HE 'ENETICS OF #OLONIZING
3PECIES STANDS STARKLY ALONE DURING THIS TIME PERIOD 0ERHAPS IT WAS BECAUSE THE
BOOK FOCUSED MORE ON EVOLUTIONARY ISSUES THAN ECOLOGICAL THEORY BUT THE SYM
POSIUM AND COMPANION VOLUME ELICITED ALMOST NO RESPONSE FROM THE ECOLOGICAL
COMMUNITY $ESPITE THE PUBLICATION OF THIS VOLUME AND %LTONS BOOK SEVEN YEARS
EARLIER BIOLOGICAL INVASIONS DID NOT ATTRACT THE INTEREST OF MANY ECOLOGISTS DUR
ING THE S AT LEAST FROM ECOLOGISTS WHOSE PRIMARY INTERESTS AND OBJECTIVES
INVOLVED THE DEVELOPMENT OF THEORY AND GENERALIZATIONS THAT TRANSCEND PARTICULAR
ORGANISMS AND HABITATS )T IS CERTAINLY TRUE THAT WITH THE DEVELOPMENT AND DIS
SEMINATION OF ISLAND BIOGEOGRAPHY THEORY -AC!RTHUR AND 7ILSON
3IMBERLOFF AND 7ILSON CONSIDERABLE ATTENTION WAS GIVEN TO HOW ARRIVING
SPECIES WOULD lT INTO RESIDENT COMMUNITIES (OWEVER FEW ECOLOGISTS AT THIS TIME
CONNECTED THEIR RESEARCH WITH THE SORT OF INVASION LITERATURE AND ISSUES SUMMA
RIZED AND PRESENTED BY %LTON 3IMBERLOFF PERSONAL COMMUNICATION
)T MUST BE POINTED OUT THAT DURING THIS TIME SOME ECOLOGISTS OUTSIDE OF .ORTH
!MERICA WERE ACTIVELY RESEARCHING AND PUBLISHING IN THE AREA OF INTRODUCED
SPECIES 3UKOPP ADDRESSED BOTH THEORY AND TERMINOLOGY IN HIS REPORT ON
INTRODUCED SPECIES IN THE NATURAL PLANT ASSOCIATIONS OF CENTRAL %UROPE (OLUB
-! $AVIS
AND *IRSEK AND 3CHROEDER PRESENTED CLASSIlCATION SCHEMES FOR
NON
NATIVE SPECIES BASED ON THE MODE OF INTRODUCTION TIME OF INTRODUCTION AND
DEGREE OF NATURALIZATION &ALINSKI WROTE HIS DISSERTATION ON THE DISTRIBU
TION OF INTRODUCED SPECIES IN 0OLANDS "IALOWIEZA 0RIMEVAL &OREST AND &ALINSKI
AND +ORNAS A B PUBLISHED A NUMBER OF SUBSEQUENT
ARTICLES DURING THE S ON THE ECOLOGY OF INTRODUCED SPECIES IN NATURAL AND
SEMI
NATURAL COMMUNITIES IN 0OLAND (UNGARIAN ECOLOGIST 0AL *UHASZ
.AGY
CONDUCTED lELD EXPERIMENTS IN HIS STUDIES OF @ECESIS RESISTANCE INVASIBIL
ITY AND USED THE TERM @ECOLOGICAL HOMEOSTASIS TO DESCRIBE COMMUNITIES THAT WERE
RESISTANT TO INVASION (EJN AND ,HOTSK AND *EHLK AND 3LAVK
DESCRIBED THE ECOLOGY AND DISTRIBUTION AND SPREAD IN NATURAL ENVIRONMENTS OF SEV
ERAL INTRODUCED SPECIES IN THE #ZECH 2EPUBLIC +OHLER AND 3UKOPP STUDIED
THE ECOLOGY OF INTRODUCED SPECIES IN CITIES PART OF A LONG
STANDING INTEREST AMONG
%UROPEAN PLANT ECOLOGISTS /NE OF THE MORE MEMORABLE lNDINGS OF +OHLER AND
3UKOPP WAS THAT 2OBINIA PSEUDOACACIA HAD SPREAD SPONTANEOUSLY ON THE RUBBLE OF
CITIES BOMBED DURING 77)) %XCEPT FOR THE ABSTRACTS NONE OF THE ABOVE ARTICLES
WAS PUBLISHED IN %NGLISH AND MOST %NGLISH
SPEAKING ECOLOGISTS WERE PROBABLY NOT
WELL AWARE OF THIS RESEARCH DURING THE SIXTIES
!LTHOUGH BIOLOGICAL INVASIONS RECEIVED LITTLE ATTENTION FROM ECOLOGISTS DURING
THE S PARTICULARLY IN .ORTH !MERICA THIS DOES NOT MEAN THAT RESEARCH WAS
NOT BEING CONDUCTED ON INTRODUCED SPECIES AND THEIR IMPACTS )N FACT CONSIDER
ABLE RESEARCH ON THIS TOPIC WAS BEING CONDUCTED IN !USTRALIA .EW :EALAND
3OUTH !FRICA %UROPE AS WELL AS .ORTH !MERICA )T WAS SIMPLY BEING CONDUCTED BY
BIOLOGISTS WHOSE INTERESTS AND PRIORITIES WERE MORE APPLIED IN NATURE INVOLVING
lSHERIES EG !LBRECT WILDLIFE EG "UMP AND "OHL 4#7&
7ARNER FORESTRY EG .ICHOLS 4AYLOR AND AGRICULTURE EG
3ALISBURY -ETCALF ET AL $E"ACH 4HE APPLIED AND PEST
CONTROL
RESEARCH CONDUCTED IN DURING THE S DID NOT REPRESENT ANY NEW INITIATIVE AND
VERY LITTLE IF ANY OF IT WAS INSPIRED BY %LTONS BOOK 2ATHER IT WAS A CONTINUATION OF
EXTENSIVE APPLIED RESEARCH IN THIS AREA EXTENDING BACK TO THE BEGINNING OF THE TH
CENTURY EG ,ITTLE "AILEY "RYANT 3IM 7ICHT
AND EVEN EARLIER EG -ERRIAM 0ALMER
&ROM AN HISTORICAL PERSPECTIVE A BOOK BY 'EORGE ,AYCOCK WAS A NOTEWOR
THY PUBLICATION DURING THIS TIME 4ITLED 4HE !LIEN !NIMALS 4HE 3TORY OF )MPORTED
7ILDLIFE THE BOOK WAS PUBLISHED BY 4HE !MERICAN -USEUM OF .ATURAL (ISTORY
7ELL RESEARCHED IT CONTAINED NEARLY REFERENCES FROM THE SCIENTIlC LITERA
TURE ,IKE %LTONS CLASSIC THIS BOOK WAS WRITTEN FROM A CONSERVATION AND
DISTINCTLY VALUE
BASED PERSPECTIVE AS WELL AS FOR A LARGER PUBLIC AUDIENCE /N
THE OPENING PAGE ,AYCOCK REFERS TO hMANv AS hTHE SUPREME MEDDLERv ON
THE BOOKS lNAL PAGE HE WARNS OF NATURAL COMMUNITIES BEING hPOLLUTEDv BY ALIEN
SPECIES AND TITLES OF HIS CHAPTERS INCLUDED 3TANGERS IN THE 3OUTHWEST (OW THE 'RAY
3QUIRRELS )NVADED %NGLAND AND 4HE #ONQUERING -ONGOOSE )N THE BOOK ,AYCOCK
PRESENTS TWENTY CASE STUDIES OF ANIMAL INTRODUCTIONS AROUND THE WORLD THAT HAD
CAUSED GREAT ECONOMIC AND CONSERVATION HARM 4HE BOOKS LANGUAGE TONE AND
)NVASION BIOLOGY
SERIES OF CASE STUDIES IS VERY SIMILAR TO %LTONS BOOK ALTHOUGH RATHER INEX
PLICABLY %LTON WAS NOT ONE OF THE NEARLY REFERENCES LISTED IN THE BIBLIOGRAPHY
(OWEVER UNLIKE %LTON ,AYCOCK WAS NOT A SCIENTIST (E WAS A FREELANCE WRITER
WHO DURING HIS CAREER AUTHORED MORE THAN TWENTY BOOKS ON ANIMALS AND NATURE
MANY OF THEM WRITTEN FOR CHILDREN AND YOUNG ADULTS .EVERTHELESS ,AYCOCKS
CHARACTERIZATION OF INTRODUCED SPECIES SEEMS TO HAVE PORTENDED THE PERSPECTIVE
ADOPTED IN A LARGE NUMBER OF INVASION ECOLOGY LITERATURE BOTH POPULAR AND SCIEN
TIlC LATER IN THE CENTURY MORE SO THAN HAS THE !SILOMAR PUBLICATION 4HE 'ENETICS
OF #OLONIZING 3PECIES
4(% 3
/N !PRIL HUNDREDS OF THOUSANDS OF STUDENTS AND OTHERS ACROSS THE
5NITED 3TATES CELEBRATED THE lRST %ARTH $AY AN EVENT CONCEIVED AND ORGANIZED
BY 'AYLORD .ELSON THEN 3ENATOR OF 7ISCONSIN !LTHOUGH .ELSON 2ACHEL #ARSON
AND OTHERS HAD BEEN TRYING TO RAISE ENVIRONMENTAL CONSCIOUSNESS WITHIN THE
5NITED 3TATES DURING THE SIXTIES THE S IS WHEN THE ENVIRONMENTAL MOVE
MENT lNALLY CAPTURED THE PUBLICS ATTENTION IN THE 53 AND THROUGHOUT THE WORLD
3#/0% 3CIENTIlC #OMMITTEE ON 0ROBLEMS OF THE %NVIRONMENT A COMMITTEE
ESTABLISHED BY THE )NTERNATIONAL #OUNCIL OF 3CIENTIlC 5NIONS PUBLISHED THE lRST OF
ITS 3#/0% 3ERIES TITLES NOW NUMBERING MORE THAN SIXTY IN 4HE S IS
WHEN CONSERVATION BIOLOGY BEGAN TO EMERGE AS A DISCIPLINE AND ECOLOGISTS WORLD
WIDE BEGAN TO STUDY INTRODUCED SPECIES FROM A CONSERVATION PERSPECTIVE )N THE
EARLY S THE NEW JOURNAL "IOLOGICAL #ONSERVATION BEGAN PUBLISHING ARTICLES
DESCRIBING THE THREATS POSED BY INTRODUCED SPECIES EG #AMPBELL AND /RMOND
3CHOlELD
!PPLIED ECOLOGISTS WHO HAD BEEN STUDYING INTRODUCED SPECIES AND THEIR ECOLOGI
CAL EFFECTS IN THE SIXTIES AND EARLIER CONTINUED TO DO SO IN THE SEVENTIES PUBLISHING
PRIMARILY IN TAXONOMIC
SPECIlC JOURNALS AS THEY HAD PREVIOUSLY &OR EXAMPLE
/WRE DESCRIBED THE EXTENT OF INTRODUCED AVIFAUNA IN SOUTHEASTERN &LORIDA
AND RAISED THE POSSIBILITY OF COMPETITION BETWEEN THE NATIVE AND NEW SPECIES IN
THE CONSERVATION SECTION OF 4HE 7ILSON "ULLETIN #HRISTIE DESCRIBED THE
EFFECTS OF SPECIES INTRODUCTIONS ON SALMONID COMMUNITIES *OURNAL OF THE &ISHERIES
"OARD OF #ANADA AND -OYLE REPORTED ON THE EFFECTS ON THE NATIVE FROGS
OF THE INTRODUCED BULLFROGS IN #ALIFORNIA #OPEIA )N A "IOSCIENCE REVIEW PAPER
#OURTENAY AND 2OBINS SUMMARIZED THE CONSERVATION PROBLEMS PRIMAR
ILY IN .ORTH !MERICA CREATED BY THE INTRODUCTION OF hEXOTICv ANIMALS BY lSH AND
GAME AGENCIES AND THE PET TRADE 4HE %UROPEAN ECOLOGISTS WHO HAD BEEN STUDY
ING INTRODUCED SPECIES AND THEIR ECOLOGICAL EFFECTS IN THE SIXTIES ALSO CONTINUED
THEIR RESEARCH IN THE SEVENTIES PUBLISHING HUNDREDS OF PAPERS ON THIS TOPIC &OR
EXAMPLE &ALINSKI AND 3UKOPP CONTINUED THEIR WORK ON THE URBAN
INTRODUCED AND ADVENTITIOUS mORA AND *EHLK AND (EJN CONTINUED THEIR
RESEARCH IN THE #ZECH 2EPUBLIC DESCRIBING MIGRATION ROUTES OF INTRODUCED SPECIES
-! $AVIS
AND MAKING WHAT PERHAPS WERE THE lRST PREDICTIONS OF WHEN PLANT INVASIONS WOULD
OCCUR IN PARTICULAR LOCATIONS (EJN ET AL !LTHOUGH THIS mORISTIC APPROACH
TO INTRODUCED SPECIES WAS NOT AS COMMON IN THE 5NITED 3TATES AT THIS TIME OCCA
SIONAL STUDIES OF THIS TYPE WERE UNDERTAKEN EG -UEHLENBACHS DETAILED
ACCOUNT THE ADVENTIVE mORA ALONG RAILROADS AROUND 3AINT ,OUIS -ISSOURI
/CCASIONAL PAPERS ON BIOLOGICAL INVASIONS CAN BE FOUND IN THE ECOLOGICAL LITERA
TURE WELL BEFORE THE S EG %GLER (OWEVER THE SEVENTIES WAS THE lRST
TIME SINCE %LTON THAT BIOLOGICAL INVASIONS BEGAN TO APPEAR IN MAINSTREAM ECOLOGY
LITERATURE WITH ANY FREQUENCY )N HIS REVIEW OF THE ECOLOGY OF WEEDS PUBLISHED IN
!NNUAL 2EVIEW OF %COLOGY AND 3YSTEMATICS "AKER EXPLICITLY EXTENDED THE
DISCUSSION OF WEEDS BEYOND THE DISCIPLINARY BOUNDARIES OF HORTICULTURE AND AGRI
CULTURE TO INCLUDE ECOLOGY /THER EXAMPLES INCLUDE "URDON AND #HILVERS
/ECOLOGIA ARTICLE ON THE IMPACT OF INTRODUCED PINE SPECIES ON !USTRALIAS EUCALYP
TUS FORESTS AND %MBREES CHAPTER ON THE ECOLOGY OF ANIMAL INVADERS IN THE
BOOK !N !NALYSIS OF %COLOGICAL 3YSTEMS (ORN ET AL
!LTHOUGH SOME ECOLOGISTS WERE BEGINNING TO THINK ABOUT BIOLOGICAL INVA
SIONS DURING THE SEVENTIES THERE WAS STILL LITTLE COORDINATION AND COMMUNICATION
AMONG INVESTIGATORS !ND THERE WAS NO FORMAL INFRASTRUCTURE TO SUPPORT EFFORTS
TO DEVELOP A THEORY OF BIOLOGICAL INVASIONS %COLOGY MEETINGS AT THAT TIME DID NOT
DEVOTE SPECIAL SYMPOSIA OR PAPER SECTIONS TO BIOLOGICAL INVASIONS WHICH WOULD
HAVE SERVED AS IMPORTANT COMMUNICATION HUBS FOR THIS TOPIC 4HUS ALTHOUGH
SCATTERED PAPERS ON BIOLOGICAL INVASIONS WERE APPEARING IN THE ECOLOGICAL LITERA
TURE INVASION ECOLOGY HAD NOT YET EMERGED AS ITS OWN RESEARCH SPECIALTY AREA
4HE NUMBER OF PUBLISHED STUDIES ON BIOLOGICAL INVASIONS INCREASED DRAMATICALLY
DURING THE S ESPECIALLY IN PARTICULAR TAXA SUCH AS PLANTS SEE REVIEW BY
0YEK 4HE EMPHASIS ON mORISTICS THAT CHARACTERIZED MUCH OF THE WORK IN
PREVIOUS DECADES BEGAN TO GIVE WAY TO THE INCREASING INTEREST ON SPECIES BIOLOGY
AND ECOLOGICAL INTERACTIONS 0YEK $UE TO THE PROLIFERATION OF INVASION
PUBLICATIONS DURING THIS TIME THE HISTORICAL ACCOUNT OF THIS DECADE HAS BEEN SPLIT
INTO TWO PERIODS
-ANY OF THE %UROPEAN ECOLOGISTS WHO HAD BEEN STUDYING INTRODUCED AND
ADVENTIVE mORA IN PRIOR DECADES CONTINUED TO DO SO IN THE EIGHTIES PARTICULARLY
FOCUSING ON THE ECOLOGY OF URBAN mORA EG 3UKOPP AND 7ERNER +OWARIK
)N THE 5NITED 3TATES 2ICHARD -ACK PUBLISHED HIS lRST ARTICLE ON BIOLOGICAL
INVASIONS IN REPORTING ON THE ECOLOGICAL IMPACT IN WESTERN .ORTH !MERICA
OF "ROMUS TECTORUM -ACK TOOK A CONSERVATION APPROACH IN HIS ARTICLE QUOTING
%LTON AND EMPHASIZING THE DECLINE OF NATIVE GRASS SPECIES (E CHOSE TO SUBMIT HIS
ARTICLE TO THE MORE APPLIED JOURNAL !GROECOSYSTEMS BECAUSE ITS SCOPE ENCOMPASSED
THE ECOSYSTEMS THAT CHEATGRASS HAD INVADED )N THE SAME YEAR $AN 3IMBERLOFF
CONTRIBUTED A CHAPTER TO THE BOOK "IOTIC #RISES IN %COLOGICAL AND %VOLUTIONARY 4IME
)NVASION BIOLOGY
.ITECKI 4HE TITLE OF THE BOOK SUGGESTS THAT 3IMBERLOFF MIGHT ALSO
HAVE EMPHASIZED THE CONSERVATION ASPECTS OF BIOLOGICAL INVASIONS )N FACT HE DID
NOT DO THIS BUT RATHER USED BIOLOGICAL INVASIONS AS A WAY TO TEST ECOLOGICAL THEORY
EG ISLAND BIOGEOGRAPHY THEORY AND MODELS OF LIMITING SIMILARITY 4HUS WHEREAS
-ACKS ARTICLE WAS MORE IN THE CONSERVATION TRADITION OF %LTONS
BOOK 3IMBERLOFFS CHAPTER WAS MORE IN KEEPING WITH THE EMPHASIS ON ECOLOGI
CAL AND EVOLUTIONARY THEORY THAT DISTINGUISHED 4HE 'ENETICS OF #OLONIZING 3PECIES
)N THE SAME YEAR "ROWN AND -ARSHALL PUBLISHED AN ARTICLE EVEN MORE
REMINISCENT OF THE !SILOMAR PROCEEDINGS EMPHASIZING THE EVOLUTIONARY
CHANGES ACCOMPANYING THE COLONIZATION OF PLANTS 4WO YEARS LATER -OULTON
AND 0IMM EXAMINED THE IMPACTS OF THE INTRODUCED (AWAIIAN AVIFAUNA
AS A WAY TO ASSESS THE IMPORTANCE OF COMPETITION IN COMMUNITY ASSEMBLY ,IKE
3IMBERLOFF AND "ROWN AND -ARSHALL THEY ALSO EMPHASIZED THEORY AS EVIDENCED
BY THEIR CHOICE OF JOURNAL !MERICAN .ATURALIST 4HE TONE AND STYLE USED BY
3IMBERLOFF "ROWN AND -ARSHALL AND -OULTON AND 0IMM
WAS CONSISTENT TO THAT USED BY THE CONTRIBUTORS TO 4HE 'ENETICS OF #OLONIZING 3PECIES
OTHER THAN (ARPER
4HE CONNECTION BETWEEN INTRODUCED SPECIES AND CONSERVATION CONTINUED TO
GROW DURING THE )MMIGRANT +ILLERS +ING DESCRIBED THE IMPACT
OF INTRODUCED PREDATORS ON NATIVE BIRDS IN .EW :EALAND !LTHOUGH WRITTEN FOR A
POPULAR AUDIENCE +INGS BOOK WAS WELL RESEARCHED AND REFERENCED AND IT REPRE
SENTED AN IMPORTANT SCIENTIlC CONTRIBUTION AS WELL
)N 7ILLIAM *ORDAN ))) AND THE 5NIVERSITY OF 7ISCONSIN !RBORETUM PUB
LISHED THE lRST ISSUE OF 2ESTORATION AND -ANAGEMENT .OTES NOW PUBLISHED UNDER
THE NAME OF %COLOGICAL 2ESTORATION AN EVENT THAT SIGNIlED THE BEGINNING OF AN
ORGANIZED INTEREST IN HABITAT RESTORATION IN .ORTH !MERICA ,AND MANAGERS HAD
BEEN THINKING ABOUT RESTORATION IN THE S &OR EXAMPLE .!4/ SPONSORED
A CONFERENCE IN 2EYKJAVIK )CELAND THAT RESULTED IN THE PUBLICATION OF
4HE "REAKDOWN AND 2ESTORATION OF %COSYSTEMS (OLDGATE AND 7OODMAN
%UROPEANS CONSTITUTED THE MAJORITY OF PARTICIPANTS AT THE 2EYKJAVIK CONFERENCE
AND THE INTEREST AT THE MEETING WAS MORE IN hREHABILITATINGv THE LANDSCAPE RATHER
THAN TRYING TO RESTORE THE HABITAT TO SOME PRIOR NATIVE CONDITION AS ILLUSTRATED BY
A STATEMENT BY "RADSHAW ET AL hFROM THE POINT OF VIEW OF NATURE CONSERVA
TION REHABILITATION SHOULD NOT NECESSARILY MEAN RESTORATION OF THE ORIGINAL OFTEN
UNINTERESTING ECOSYSTEMv 4HIS PERSPECTIVE CONTRASTED GREATLY WITH THE EMERGING
INTEREST IN ECOLOGICAL RESTORATION IN THE 5NITED 3TATES )N HIS OPENING EDITORIAL OF
THE lRST VOLUME OF 2ESTORATION AND -ANAGEMENT .OTES 7ILLIAM *ORDAN )))
STATED THAT THE NEW JOURNAL hWILL DEAL ONLY WITH THE DEVELOPMENT AND MANAGE
MENT OF COMMUNITIES THAT ARE NATIVE OR AT LEAST ECOLOGICALLY APPROPRIATE TO THEIR
SITEv 4HE STATED FOCUS OF THE JOURNAL WAS TO BE THE RESTORATION AND MANAGEMENT
OF ECOLOGICAL COMMUNITIES FOR SCIENTIlC AND AESTHETIC PURPOSES EG PRAIRIES
WETLANDS AND FORESTS BUT NOT FOR RANGE OR TIMBER MANAGEMENT AND NOT FOR hREC
LAMATION EFFORTS AIMED SOLELY AT LAND STABILIZATIONv 4HE SUBSEQUENT DEVELOPMENT
OF THE lELD OF RESTORATION ECOLOGY IN FUTURE YEARS WAS TO HAVE A MAJOR IMPACT ON
-! $AVIS
INVASION ECOLOGY PARTICULARLY IN .ORTH !MERICA THROUGH ITS EMPHASIS ON NATIVE
ENVIRONMENTS AND NATIVE SPECIES
"Y THE MID S MORE AND MORE ECOLOGISTS WERE BEGINNING TO THINK ABOUT
INVASIONS FROM AN ECOLOGICAL PERSPECTIVE (OWEVER WHAT WAS STILL LACKING WAS
A SYNTHESIS OF THIS THINKING THAT WOULD HELP DElNE A lELD OF INVASION ECOLOGY )N
FACT THE GROUNDWORK FOR SUCH AN EFFORT WAS ALREADY UNDERWAY )N THE 4HIRD
)NTERNATIONAL #ONFERENCE ON -EDITERRANEAN %COSYSTEMS WAS HELD IN 3TELLENBOSCH
3OUTH !FRICA AND INTEREST IN BIOLOGICAL INVASIONS AT THE MEETING LED TO A PROPOSAL
TO THE 3#/0% GENERAL ASSEMBLY IN /TTAWA IN 4HIS PROPOSAL RESULTED IN THE
CREATION IN OF A SCIENTIlC ADVISORY COMMITTEE THAT WAS TO ENCOURAGE AND
FACILITATE FOCUS ON THE IMPACTS OF BIOLOGICAL INVASIONS ON NATURAL ECOSYSTEMS 4HE
ADVISORY COMMITTEE ARTICULATED THREE QUESTIONS TO GUIDE THE 3#/0% INVASION PRO
GRAM WHAT FACTORS DETERMINE WHETHER A SPECIES WILL BE AN INVADER OR NOT
WHAT ARE THE CHARACTERISTICS OF THE ENVIRONMENT THAT MAKE IT EITHER VULNERABLE TO
OR RESISTANT TO INVASIONS HOW CAN THE KNOWLEDGE GAINED FROM ANSWERING THE
lRST TWO QUESTIONS BE USED TO DEVELOP EFFECTIVE MANAGEMENT STRATEGIES $EVELOPED
WITH THE INTENT OF hBUILDING ON THE CONSIDERABLE KNOWLEDGE BASE AVAILABLE ON
INVADERS OF AGRICULTURAL SYSTEMSv THE 3#/0% PROGRAM ON BIOLOGICAL INVASIONS
WAS CLEARLY CONCEIVED IN THE CONSERVATION TRADITION OF %LTON )T IS NO COINCIDENCE
THAT SCIENTISTS FROM 3OUTH !FRICA !USTRALIA .EW :EALAND AND THE 5NITED 3TATES
CONTRIBUTED SO SIGNIlCANTLY TO THE 3#/0% INITIATIVE 4HE NATURAL ENVIRONMENTS
IN THESE COUNTRIES HAD BEEN EXPERIENCING RECENT AND SUBSTANTIAL INTRODUCTIONS OF
NEW SPECIES FROM OTHER REGIONS OF THE WORLD AND CONSIDERABLE CONSERVATION CON
CERNS WERE BEING RAISED IN THESE COUNTRIES REGARDING THE IMPACTS OF THESE SPECIES
3OUTH !FRICAN SCIENTISTS HAD BEEN AMONG THE LEADERS IN RECOGNIZING THE CONSER
VATION IMPLICATIONS OF INTRODUCED SPECIES EG 7ICHT 4AYLOR AND
THEY PUBLISHED THE lRST 3#/0% VOLUME IN -ACDONALD AND *ARMAN
(OWEVER IT WAS NOT UNTIL THE LATTER HALF OF THE DECADE THAT MOST OF THE 3#/0%
REGIONAL WORKSHOPS BEGAN PUBLISHING THEIR PROCEEDINGS 4HE .ORTH !MERICAN
CONTINGENT PUBLISHED %COLOGY OF "IOLOGICAL )NVASIONS OF .ORTH !MERICA AND (AWAII
-OONEY AND $RAKE 4HE BOOK WAS ORGANIZED AROUND VARIOUS PATTERNS
OF INVASIONS INCLUDING ASSOCIATIONS WITH DIFFERENT TAXONOMIC GROUPS ATTRIBUTES OF
INVADERS AND SITE CHARACTERISTICS PROMOTING INVASIONS )N ADDITION A VARIETY OF
CASE STUDIES WERE PROVIDED 4HIS PUBLICATION WAS ENTIRELY A 5NITED 3TATES INITIATIVE
WITH BOTH EDITORS AND ALL THE CONTRIBUTORS BEING FROM 53 UNIVERSITIES 4HAT BIO
LOGICAL INVASIONS HAD lNALLY CAPTURED THE INTEREST OF PROMINENT .ORTH !MERICAN
ECOLOGISTS IS EVIDENCED BY THE CONTRIBUTORS TO THE BOOK EG 3IMBERLOFF %HRLICH
"AZAZZ 2EGAL /RIANS 6ITOUSEK 2OUGHGARDEN %WEL 0IMM AND -OONEY
#OINCIDENTALLY THE SYMPOSIUM THAT LED TO THIS VOLUME TOOK PLACE IN !SILOMAR
#ALIFORNIA THE SAME SITE OF THE SYMPOSIUM ON THE GENETICS OF COLONIZING SPECIES
)NVASION BIOLOGY
HELD TWENTY YEARS EARLIER 4HE VOLUME CONTAINS AN INTERESTING MIXTURE
OF CHAPTERS -OST WERE CLEARLY CONCEIVED AND WRITTEN WITH A CONSERVATION AND
ENVIRONMENTAL PERSPECTIVE AS WOULD BE EXPECTED GIVEN THE NATURE OF THE 3#/0%
PROGRAM (OWEVER SOME CHAPTERS WERE WRITTEN MORE IN THE SPIRIT OF THE
!SILOMAR SYMPOSIUM AND FOCUSED MORE ON ECOLOGICAL THEORY 3IMBERLOFFS
CHAPTER ON THE BIOGEOGRAPHY OF INSECT INTRODUCTIONS EXAMINED TOPICS SUCH AS
ISLAND
MAINLAND PATTERNS AND ISSUES OF BIOTIC RESISTANCE PRIMARILY FROM A THEO
RETICAL PERSPECTIVE !LTHOUGH HE DID CITE %LTON 3IMBERLOFF DID NOT TAKE A STRONGLY
CONSERVATION APPROACH IN HIS CHAPTER REFERRING PRIMARILY TO @INTRODUCED SPE
CIES AND @COLONIZERS RATHER THAN @INVADERS AND @EXOTICS AND SPECIlCALLY RECOM
MENDED THE hRElNEMENT OF THE APPROACH OF MANY OF THE PAPERS IN 4HE 'ENETICS OF
#OLONIZING 3PECIES "AZZAZ ORGANIZED HIS CHAPTER ON LIFE HISTORIES AROUND
THE CONCEPT OF @COLONIZING SPECIES AND ALSO MADE EXPLICIT REFERENCES TO THE
BOOK -OULTON AND 0IMM EXPANDED ON THEIR PAPER AND SHOWED HOW
BIOLOGICAL INVASIONS CAN BE USED TO TEST ECOLOGICAL THEORY
!DDITIONAL PROCEEDINGS WERE PUBLISHED FROM 3#/0% WORKSHOPS HELD IN 3OUTH
!FRICA !USTRALIA AND 'REAT "RITAIN EG -ACDONALD ET AL 'ROVES AND
"URDON +ORNBERG AND 7ILLIAMSON "EING 3#/0% INITIATIVES THESE
PUBLICATIONS WERE ALSO WRITTEN FROM A CONSERVATION PERSPECTIVE (OWEVER COM
MENTS BY THE EDITORS OF THE !USTRALIAN PROCEEDINGS INDICATE THAT THEY RECOGNIZED
AN EMERGING TENSION THAT WAS DEVELOPING IN INVASION ECOLOGY AS SOME ECOLOGISTS
EMBRACED THE CONSERVATION AND STRONG ENVIRONMENTAL EMPHASIS ARTICULATED BY
%LTON WHILE OTHERS EXPRESSED CONCERN OVER THE STRONG NORMATIVE NATURE OF THIS
APPROACH )N THE &OREWORD TO THE PUBLICATION 'ROVES AND "URDON
THE EDITORS STATED THAT hTHE TERMINOLOGY ASSOCIATED WITH @INVASIONS GENERALLY IS
UNSATISFACTORY WITH ITS CONNOTATIONS OF A MILITARY OPERATION x 7HEREVER POSSIBLE
WE HAVE TRIED TO ELIMINATE EMOTIVE TERMS SUCH AS @WEEDS @PESTS @EXOTICS AND
@ALIENSv )NSTEAD THE EDITORS STRIVED TO UTILIZE THE WORD @INTRODUCED AS MUCH AS
POSSIBLE THROUGHOUT THE BOOK
!NOTHER IMPORTANT DEVELOPMENT THAT OCCURRED DURING THIS TIME PERIOD WAS THE
FOUNDING IN OF THE 3OCIETY FOR %COLOGICAL 2ESTORATION 3%2 NOW KNOWN AS
3OCIETY FOR %COLOGICAL 2ESTORATION )NTERNATIONAL 3%2 EMERGED OUT OF THE RAPIDLY
DEVELOPING INTEREST IN RESTORATION BY INDIVIDUALS AND CONSERVATION GROUPS INTER
ESTED IN MANAGING NATURAL ENVIRONMENTS 5PON ITS FOUNDING 3%2 WAS PRIMAR
ILY A .ORTH !MERICAN INITIATIVE AND THE RESTORATION ECOLOGY MOVEMENT THAT IT
REPRESENTED AND INSPIRED SHARPLY DISTINGUISHED BETWEEN SPECIES BASED ON THEIR
GEOGRAPHY OF ORIGIN WITH NATIVE SPECIES BEING DESIRABLE AND NON
NATIVES BEING
UNDESIRABLE IN NATURAL ENVIRONMENTS 4HE STARK DICHOTOMY BETWEEN NATIVE AND
NON
NATIVE SPECIES PARTICULARLY CHARACTERIZED CONSERVATION AND RESTORATION EFFORTS
IN REGIONS LIKE .ORTH !MERICA (AWAII AND .EW :EALAND WHICH HAD EXPERIENCED
RELATIVELY FEW SPECIES INTRODUCTIONS UNTIL THE PAST FEW CENTURIES 4HIS DISTINCTION
SEEMED TO BE LESS CRUCIAL AMONG CONSERVATIONISTS AND RESTORATIONISTS IN %UROPE
WHERE AS 5SHER B ACKNOWLEDGED DISTINGUISHING BETWEEN NATIVE AND INTRO
DUCED SPECIES IS OFTEN PROBLEMATIC DUE TO THE MILLENNIA OF HUMAN MOVEMENTS AND
-! $AVIS
SPECIES INTRODUCTIONS ACROSS THE CONTINENT 2ESTORATION ECOLOGY AND INVASION ECOL
OGY EMERGED AT ABOUT THE SAME TIME AND IN .ORTH !MERICA THEY DEVELOPED AS
SISTER DISCIPLINES DURING THE LATTERS AND CONTINUED TO REINFORCE ONE ANOTHER
IN SUBSEQUENT YEARS
4HE GROWING RELATIONSHIP BETWEEN INVASION ECOLOGY AND CONSERVATION ECOLOGY
DURING THIS TIME WAS ILLUSTRATED BY THE DECISION OF THE EDITORS OF THE JOURNAL "IOLOGICAL
#ONSERVATION TO DEVOTE A SPECIAL ISSUE TO THIS TOPIC 4ITLED "IOLOGICAL )NVASIONS OF
.ATURE 2ESERVES THIS PUBLICATION 5SHER A REPRESENTED ANOTHER 3#/0% PROD
UCT !LTHOUGH IT WAS BECOMING INCREASINGLY COMMON FOR ECOLOGISTS TO ADOPT THE
MORE EXPLICIT CONSERVATION PERSPECTIVE EMPHASIZED BY THE 3#/0% PROGRAM NOT ALL
ECOLOGISTS WERE DOING SO &OR EXAMPLE IN HIS ARTICLE ON INVASION WINDOWS *OHNSTONE
EXAMINED HABITAT INVASIBILITY MOSTLY FROM A THEORETICAL PERSPECTIVE AND
HE DID NOT LINK HIS ARTICLE TO CONSERVATION CONCERNS REGARDING INVASIONS
4HE DECADE ENDED WITH THE PUBLICATION OF "IOLOGICAL )NVASIONS ! 'LOBAL
0ERSPECTIVE $RAKE ET AL THE SYNTHESIS REPORT OF THE VARIOUS REGIONAL 3#/0%
SYMPOSIA AND WORKSHOPS 4HIS PUBLICATION REPRESENTED THE TH PUBLICATION IN THE
3#/0% SERIES 4WENTY
NINE ECOLOGISTS REPRESENTING SEVEN COUNTRIES 53! 5+
!USTRALIA 3OUTH !FRICA &RANCE $ENMARK )NDIA MANY OF WHOM HAD CONTRIBUTED
TO THE VOLUME AUTHORED CHAPTERS FOR THIS PUBLICATION 3OME OF THE PROMI
NENT NON
!MERICAN CONTRIBUTORS MANY OF WHOM HAVE CONTINUED TO FOCUS MUCH
OF THEIR RESEARCH ON BIOLOGICAL INVASIONS UP TO THE PRESENT INCLUDED &RANCESCO
DI #ASTRI 0ETER !SHTON $AVID 2ICHARDSON )AN -ACDONALD -ICHAEL 5SHER
-ARK 7ILLIAMSON 2ICHARD (OBBS -ICHAEL #RAWLEY 2ICHARD 'ROVES AND -ARCEL
2EJMANEK 2EJMANEK HAD MOVED FROM #ZECHOSLOVAKIA TO THE 5NITED 3TATES BY THE
TIME THE BOOK WAS PUBLISHED )NDIVIDUALLY FOCUSING ON DIFFERENT TAXA OR REGIONS OF
THE WORLD CONTRIBUTORS TRIED TO ANSWER THE THREE QUESTIONS THAT HAD BEEN POSED BY
THE 3#/0% ADVISORY COMMITTEE IN )N THE BOOKS lNAL CHAPTER -OONEY AND
$RAKE CONCLUDED THAT ALTHOUGH KNOWLEDGE PERMITTED hGENERIC GUIDELINESv
FOR ASSESSING THE LIKELIHOOD OF BIOLOGICAL INVASIONS EG DISTURBANCES NORMALLY
INCREASED INVASIBILITY THEY CONCURRED WITH THE CONCLUSIONS REACHED BY MOST OF THE
AUTHORS THAT IT WAS NOT POSSIBLE AT THAT TIME TO MAKE ACCURATE PREDICTIONS ABOUT
INDIVIDUAL CASES
!LTHOUGH SOME OF THE .ORTH !MERICAN ECOLOGISTS CONTINUED TO USE THE TERMS
@INTRODUCTION AND @INTRODUCED SPECIES RATHER THAN @INVASIONS AND @INVADERS IN
THE VOLUME EG 3IMBERLOFF AND 0IMM POSSIBLY INDICATING SOME RESISTANCE
TO ADOPTING THE MORE VALUE
BASED CONSERVATION APPROACH IT IS INTERESTING THAT THE
FREQUENCY WITH WHICH !MERICAN ECOLOGISTS CITED %LTONS BOOK CONTINUED TO
INCREASE )N ONLY THREE OF THE ELEVEN 53 CONTRIBUTORS TO THE 'ENETICS
OF #OLONIZING 3PECIES CITED 4HE %COLOGY OF )NVASIONS BY !NIMALS AND 0LANTS )N THE
.ORTH !MERICAN 3#/0% VOLUME WHICH WAS AUTHORED ENTIRELY BY 53 ECOLO
GISTS OF THE CHAPTERS CITED %LTONS BOOK !ND IN THE 3#/0% SYNTHESIS
VOLUME OF THE 53 AUTHORED CHAPTERS CITED %LTONS BOOK "Y CONTRAST
ONLY OF OF THE CHAPTERS IN THE 3#/0% VOLUME AUTHORED ENTIRELY
BY NON
53 ECOLOGISTS CITED %LTON 4WO OF THE CHAPTERS WERE AUTHORED BY AT LEAST
)NVASION BIOLOGY
ONE 53 AND ONE NON 53 AUTHOR ONE OF THESE CITED %LTON AND THE OTHER DID NOT )N
THE BOOKS lNAL CHAPTER (AL -OONEY AND *AMES $RAKE BOTH 53 ECOLOGISTS SPECIl
CALLY ENCOURAGED THE READERS TO CONSULT %LTONS BOOK h4HOSE WISHING AN IN DEPTH
TREATMENT OF THE DYNAMICS OF BIOLOGICAL INVASIONS ARE REFERRED TO %LTONS
CLASSIC WORK AND TO THE PUBLICATIONS WHICH AROSE OUT OF THE PROGRAMv ;IE THE
3#/0% PROGRAM=
4HE SCATTERED IDEAS CONCERNING BIOLOGICAL INVASIONS PUT FORTH BY ECOLOGISTS IN
THE SEVENTIES AND EARLY EIGHTIES lNALLY COALESCED IN THE MID TO LATE EIGHTIES RESULT
ING IN THE PUBLICATION OF A NUMBER OF IMPORTANT PAPERS AND EDITED VOLUMES FROM
4HESE PUBLICATIONS REVIEWED KNOWLEDGE AND THEORY TO THAT POINT
IDENTIlED QUESTIONS TO BE ANSWERED AND SET MUCH OF THE DIRECTION FOR FUTURE
RESEARCH OF BIOLOGICAL INVASIONS )N SHORT INVASION ECOLOGY EMERGED AS A RESEARCH
SPECIALTY AREA DURING THIS TIME PERIOD AS A DIRECT RESULT OF THESE PUBLICATIONS
!LTHOUGH SOME INVESTIGATORS CONTINUED TO PURSUE INVASION ECOLOGY MORE IN THE
TRADITION OF THE !SILOMAR SYMPOSIUM ON COLONIZATION IE FOCUSING PRIMAR
ILY ON ECOLOGICAL AND EVOLUTIONARY THEORY MORE AND MORE ECOLOGISTS PARTICULARLY
.ORTH !MERICAN ECOLOGISTS WERE PRESENTING THEIR WRITINGS IN AN EXPLICITLY ENVI
RONMENTAL CONTEXT THE PATH SET FORTH BY %LTON IN
)NVASION ECOLOGY TOOK THE SPOTLIGHT FOLLOWING THE SERIES OF INVASION BOOKS AND
ARTICLES PUBLISHED BETWEEN AND "IOLOGICAL INVASIONS BECAME A POPU
LAR AREA OF RESEARCH AND AN INCREASINGLY CONTROVERSIAL TOPIC OF DEBATE DURING THE
S RESULTING IN A mOOD OF PUBLICATIONS BOTH SCHOLARLY AND POPULAR THAT CON
TINUES TO THIS DAY .ATIONAL AND INTERNATIONAL ECOLOGY MEETINGS AND SOME GENERAL
SCIENCE MEETINGS EG !!!3 BEGAN TO REGULARLY SCHEDULE SYMPOSIA AND PAPER
SESSIONS DEVOTED TO BIOLOGICAL INVASIONS AND INTRODUCED SPECIES DURING THE NINE
TIES 4HE lRST OF SEVEN INTERNATIONAL CONFERENCES ON THE ECOLOGY OF ALIEN PLANT INVA
SIONS WAS HELD IN ,OUGHBOROUGH 5+ IN AND THE TH CONFERENCE WAS HELD IN
&ORT ,AUDERDALE &LORIDA 53! IN AND SEVERAL OF THESE CONFERENCES RESULTED
IN PUBLICATIONS OF THE PROCEEDINGS EG 0YEK ET AL 3TARlNGER ET AL
#HILD ET AL 7ITH THE EXCEPTION OF THE SECOND AND SEVENTH CONFERENCES
HELD IN 4EMPE !RIZONA AND &ORT ,AUDERDALE &LORIDA THE CONFERENCES TOOK PLACE
IN %UROPE WITH %UROPEAN ECOLOGISTS REPRESENTING THE MAJORITY OF PARTICIPANTS
4HUS THIS SERIES PROVIDES AN EXCELLENT OVERVIEW OF THE RESEARCH AND IDEAS COMING
FROM %UROPE AT THIS TIME PARTICULARLY FROM EASTERN %UROPEAN COUNTRIES SUCH AS
THE #ZECH 2EPUBLIC WHICH AS STATED ABOVE HAVE HAD A LONG TRADITION OF STUDY
ING INVASIONS 3OME OF THE PROMINENT ECOLOGISTS WHO PLAYED A SIGNIlCANT ROLE IN
THESE CONFERENCES AND PUBLICATIONS INCLUDE 0ETR 0YEK AND +AREL 0RACH #ZECH
2EPUBLIC )NGO +OWARIK AND 5WE 3TARlNGER 'ERMANY -ARK 7ILLIAMSON
-AX 7ADE AND ,OIS #HILD 5+ 'IUSEPPI "RUNDU )TALY AND *OHN "ROCK AND
-ARCEL 2EJMANEK 53!
-! $AVIS
5NLIKE THE TIME PERIODS DESCRIBED ABOVE THE YEARS
DO NOT OFFER
AN OBVIOUS LIST OF IMPORTANT PUBLICATIONS DUE TO THE SHEER NUMBER OF BOOKS AND
ARTICLES PRODUCED DURING THIS TIME 4HUS THE PUBLICATIONS BRIEmY MENTIONED BELOW
CONSTITUTE A VERY SMALL REPRESENTATIVE SUBSET OF THE LARGE NUMBER OF SIGNIlCANT
PUBLICATIONS ONE COULD HAVE INCLUDED
,ODGES 42%% ARTICLE PROVIDED A COMPREHENSIVE OVERVIEW AND ASSESSMENT
OF THE lELD OF INVASION ECOLOGY TO THAT POINT #ONSISTENT WITH PRIOR ASSESS
MENTS ,ODGE CONCLUDED THAT IT WAS NOT YET POSSIBLE TO ACCURATELY PREDICT THE
CONSEQUENCES OF A SINGLE INVASION EVENT AND THAT ANY HOPE OF USEFUL PREDICTIONS
COULD ONLY EMERGE FROM FOCUSED STUDIES ON PARTICULAR SPECIES AND ENVIRONMENTS
,ODGES PAPER WAS DISTINCTIVE BECAUSE IT CONTAINED WELL DEVELOPED ASPECTS OF
BOTH THE CONCEPTUAL APPROACHES TO BIOLOGICAL INVASIONS THAT HAD DEVELOPED DUR
ING THE PRECEDING THREE DECADES 7HILE ,ODGE BEGAN HIS PAPER EMPHASIZING THE
ENVIRONMENTAL IMPACTS OF @EXOTIC SPECIES WHICH HAVE hSERIOUS CONSEQUENCES FOR
BOTH MAN AND NATUREv HE ALSO STRESSED THE SCIENTIlC OPPORTUNITY THAT INVASIONS
PROVIDE hBECAUSE THE CHARACTERISTICS AND ECOLOGICAL IMPACT OF EXOTIC SPECIES
MAY PROVIDE CLUES TO LONGSTANDING ISSUES IN THE STUDY OF COMMUNITY ASSEMBLYv
#ONSISTENT WITH THIS DUAL EMPHASIS ,ODGE CITED BOTH %LTON AND "AKER
AND 3TEBBINS IN THE SAME SENTENCE REFERRING TO THEM BOTH AS @CLASSICS
7ILLIAMSONS BOOK "IOLOGICAL )NVASIONS WHICH REVIEWED AND SYNTHESIZED
INVASION RESEARCH AND THEORY CONDUCTED AND DEVELOPED DURING THE PRIOR SEV
ERAL DECADES MADE A VERY IMPORTANT CONTRIBUTION BY PROVIDING INVESTIGATORS A
COMPREHENSIVE AND UP
TO
DATE OVERVIEW OF THE NEW AND RAPIDLY EVOLVING lELD
7ILLIAMSONS BOOK PRIMARILY FOCUSED ON THE SCIENTIlC ASPECTS OF SPECIES INVA
SIONS HOWEVER HE DID SET THE BOOK IN A CLEAR CONSERVATION CONTEXT EMPHASIZING
THE MANAGEMENT VALUE OF BETTER SCIENTIlC UNDERSTANDING OF INVASIONS *AMES
#ARLTONS NUMEROUS PUBLICATIONS ON THE IMPACTS OF INVASIONS ON MARINE ECO
SYSTEMS EG #ARLTON A B PROMPTED MORE MARINE ECOLOGISTS
TO CONSIDER THE IMPORTANCE OF BIOLOGICAL INVASIONS 4HE PUBLICATION OF "IOLOGICAL
)NVASIONS 4HEORY AND 0RACTICE BY *APANESE ECOLOGISTS 3HIGESADA AND +AWASAKI
WHICH REVIEWED MATHEMATICAL MODELS OF SPECIES SPREAD ILLUSTRATED THE
EXTENT TO WHICH BIOLOGICAL INVASIONS HAD ATTRACTED THE INTERESTS OF ECOLOGISTS
FROM AROUND THE WORLD
4HE IMPORTANCE OF UNDERSTANDING INTERACTIONS WITH SOIL MICROBES POSITIVE AND
NEGATIVE AS PART OF THE INVASION PROCESS IN PLANTS WAS EMPHASIZED BY +LIRONOMOS
AND 2ICHARDSON ET AL EMPHASIZED THE ROLE THAT MUTUALISMS CAN
PLAY IN THE INVASION PROCESS 4HE ENEMY
RELEASE HYPOTHESIS RECEIVED CONSIDERABLE
ATTENTION DURING THIS TIME ALTHOUGH STUDIES PRODUCED CONmICTING RESULTS +EANE
AND #RAWLEY 7ILLIAMSON PROPOSED THE SINCE OFT
CITED @TENS
RULE OF BIOLOGICAL INVASIONS WHICH STATES THAT APPROXIMATELY OF IMPORTED
SPECIES EXIST IN SOME SENSE OUTSIDE OF CAPTIVITY OF THESE ESTABLISHED SELF
SUSTAINING POPULATIONS AND OF THESE BECOME PESTS 2EVIVING THE EMPHASIS ON
GENETICS OF COLONIZING SPECIES THAT CHARACTERIZED THE !SILOMAR SYMPOSIUM
MANY ECOLOGISTS BEGAN FOCUSING ON THE GENETICS OF INTRODUCED SPECIES RECOGNIZING
)NVASION BIOLOGY
THE IMPORTANCE THAT NEW GENOTYPES MIGHT PLAY IN SPECIES SPREAD %LLSTRAND
AND 3CHIERENBECK 6ILA ET AL $AEHLER AND #ARINO 0YSEK ET AL
%WEL ET AL REVIEWED THE BENElTS AND RISKS ASSOCIATED WITH DELIBER
ATE SPECIES INTRODUCTIONS AND OUTLINED THE RESEARCH NEEDED TO PROVIDE THE INFOR
MATION REQUIRED TO INCREASE THE FORMER AND LOWER THE LATTER !S INVASION ECOLOGY
CONTINUED TO DEVELOP DURING THIS PERIOD SEVERAL SCIENTISTS EXPRESSED CONCERN THAT
THE lELD WAS NOT WELL CONNECTED WITH OTHER DISCIPLINES AND THAT INVASION ECOL
OGY WAS NOT BEING INFORMED BY NOR INFORMING DISCIPLINES SUCH AS BIOGEOGRAPHY
EVOLUTIONARY BIOLOGY AND ECOLOGICAL SUBDISCIPLINES SUCH AS SUCCESSION ECOLOGY
AND WEED ECOLOGY 6ERMEIJ $AVIS ET AL $AVIS ET AL !T THE
SAME TIME SOME CONSCIOUS EFFORTS WERE BEING MADE TO BRIDGE THESE GAPS EG
"OOTH ET AL
$URING THE NINETIES ECOLOGISTS CONTINUED THE SEVERAL DECADES
LONG EFFORT TO
DETERMINE WHETHER INVADERS POSSESSED DISTINCTIVE TRAITS -ANY RESEARCHERS CON
CLUDED THAT THERE WAS LITTLE DIFFERENCE BETWEEN THE TRAITS OF INVASIVE AND NON
INVASIVE SPECIES EG 4HOMPSON ET AL 'OODWIN ET AL 7ILLIAMSON
ALTHOUGH SOME TAXONOMICALLY
BASED COMPARATIVE STUDIES DID SHOW DISTINCT
DIFFERENCES BETWEEN RESIDENT AND SPREADING INTRODUCED SPECIES EG 2EJMANEK
AND 2ICHARDSON +OLAR AND ,ODGE WERE MORE OPTIMISTIC THAT TRAITS
COULD BE USED AS PREDICTORS OF INVASIONS CONCLUDING THAT QUANTITATIVE APPROACHES
(e.g., Daehler and Carino 2000) were making progress and that these appro-
ACHES SHOULD ALLOW US TO PREDICT PATTERNS OF INVADING SPECIES MORE SUCCESSFULLY
5NLIKE ,ODGES PAPER WHICH EMPHASIZED BOTH THE CONSERVATION AND SCIENTIFIC
IMPLICATIONS OF BIOLOGICAL INVASIONS THE 42%% PAPER BY +OLAR AND ,ODGE
WAS MUCH MORE STRONGLY ORGANIZED AROUND THE CONSERVATION AND ENVIRONMENTAL
CONCERNS ABOUT SPECIES INVASIONS
&ACTORS THAT CONTRIBUTE TO AN ENVIRONMENTS SUSCEPTIBILITY TO INVASION INVASI
BILITY ONE OF THE OTHER ORGANIZING THEMES OF THE 3#/0% INITIATIVES IN THE S ALSO
CONTINUED TO BE A CENTRAL FOCUS OF RESEARCH AND DISCUSSION DURING THE S AND THE
EARLY YEARS OF THE NEW CENTURY )N HIS BOOK "IOLOGICAL $IVERSITY (USTON
ADDRESSED INVASIBILITY AND EMPHASIZED THE COMBINED IMPORTANCE OF DISTURBANCE
AND PRODUCTIVITY ,ONSDALE PRESENTED HIS lNDINGS OF A COMPREHENSIVE
REVIEW OF GLOBAL PATTERNS OF PLANT INVASIONS AND FOUND A POSITIVE CORRELATION
BETWEEN NATIVE AND EXOTIC SPECIES AT A SITE ALTHOUGH HE CONCLUDED THAT IT WAS DIFl
CULT TO DISCRIMINATE BETWEEN THE RELATIVE IMPORTANCE OF INVASIBILITY AND PROPAGULE
PRESSURE $AVIS ET AL ARGUED FOR A THEORY OF INVASIBILITY BASED ON RESOURCE
AVAILABILITY EMPHASIZING THAT INVASIBILITY IS NOT A CONSTANT CHARACTERISTIC OF AN
ENVIRONMENT BUT IS AN ATTRIBUTE THAT VARIES OVER TIME AS RESOURCE AVAILABILITY
mUCTUATES $AVIS AND 0ELSOR TESTED THIS THEORY IN A lELD EXPERIMENT BASED
ON THE INTRODUCTION OF NATIVE SPECIES PRAIRIE FORBS INTO A PLANT COMMUNITY DOMI
NATED BY INTRODUCED SPECIES .ORTH !MERICAN OLD lELD SHOWING THAT IT IS THE ECO
LOGICAL PROCESSES THAT MATTER NOT THE GEOGRAPHIC ORIGIN OF THE SPECIES INVOLVED
!LTHOUGH MUCH OF THE PROGRESS IN INVASION ECOLOGY DURING THE EIGHTIES AND
NINETIES INVOLVED MOVING BEYOND PURELY mORISTIC STUDIES OF NATIVE AND INTRODUCED
-! $AVIS
mORA MORE COMPREHENSIVE EFFORTS TO DESCRIBE AND CATALOG mORA WERE ALSO UNDER
TAKEN DURING THIS TIME PROVIDING VALUABLE DATA BASES FOR ONGOING AND FUTURE STUD
IES EG 0YSEK ET AL 4HE MOST COMPREHENSIVE NATIONAL OR REGIONAL PLANT
DATA BASES AT THE CURRENT TIME ARE LIKELY THOSE OF THE 5NITED +INGDOM #LEMENT
AND &OSTER 2YVES ET AL THE #ZECH 2EPUBLIC 0YSEK ET AL AND
!USTRIA %SSL AND 2ABITSCH /THER EXCELLENT PLANT DATA BASES INCLUDE THOSE IN
'ERMANY +LOTZ ET AL AND .ORTH !MERICA +ARTESZ AND -EACHAM
7ITH THE DEVELOPMENT OF INVASION ECOLOGY AS A RECOGNIZED RESEARCH SPECIALTY
AREA TWO INVASION RELATED JOURNALS WERE ESTABLISHED IN THE LATE NINETIES $IVERSITY
AND $ISTRIBUTIONS "LACKWELL 0UBLISHING WAS FOUNDED IN SUCCEEDING THE
JOURNAL "IODIVERSITY ,ETTERS WITH AN EMPHASIS ON BIOLOGICAL INVASIONS AND BIO
DIVERSITY $AVID 2ICHARDSON HAS SERVED AS THE JOURNALS EDITOR
IN
CHIEF FROM ITS
OUTSET "IOLOGICAL )NVASIONS +LUWER WAS FOUNDED IN WITH *AMES 4 #ARLETON
AS THE EDITOR
IN
CHIEF A POSITION CURRENTLY HELD BY *AMES ! $RAKE 4HE STATED AIMS
AND SCOPES OF BOTH JOURNALS EMPHASIZE THE SCIENCE RATHER THAN THE CONSERVATION
ASPECTS OF BIOLOGICAL INVASIONS (OWEVER IT IS NOT UNCOMMON FOR AUTHORS TO INVOKE
CONSERVATION CONCERNS IN THEIR ARTICLES PARTICULARLY IN "IOLOGICAL )NVASIONS
4WO DEVELOPMENTS IN ECOLOGY DURING THE S PARTICULARLY IMPACTED INVA
SION ECOLOGY A RENEWED EMPHASIS ON THE ECOLOGICAL EFFECTS OF BIODIVERSITY AND
THE INCREASING EMPHASIS ON GLOBAL CHANGE 2ESEARCH ON THE ECOLOGICAL IMPACTS OF
BIODIVERSITY FOCUSED ATTENTION ON %LTONS HYPOTHESIS THAT SPECIES
RICH COM
MUNITIES SHOULD BE MORE RESISTANT TO INVASIONS THAN SPECIES
POOR COMMUNITIES
! NUMBER OF SMALL
PLOT EXPERIMENTAL STUDIES INVOLVING CONSTRUCTED PLANT COM
MUNITIES YIELDED RESULTS THAT SUPPORTED %LTONS DIVERSITY
INVASIBILITY HYPOTHESIS
4ILMAN +NOPS ET AL (OWEVER THESE EXPERIMENTS WERE CHALLENGED
ON METHODOLOGICAL GROUNDS 7ARDLE AND THEIR RELEVANCY WAS QUESTIONED
SINCE STUDIES IN NATURAL COMMUNITIES OFTEN FOUND THAT THE MOST DIVERSE ENVIRON
MENTS WERE THE MOST HEAVILY INVADED ,ONSDALE 3TOHLGREN ET AL
%FFORTS HAVE BEEN MADE TO RESOLVE THIS DEBATE EG EMPHASIZING THE ROLE THAT
SPATIAL SCALE PLAYS IN INTERPRETING THE RESULTS AND THE FACT THAT CORRELATED ENVIRON
MENTAL FACTORS MAY AFFECT BOTH DIVERSITY AND INVASIBITY ,EVINE AND $!NTONIO
,EVINE 3HEA AND #HESSON (OWEVER TO DATE THESE EFFORTS
HAVE NOT QUELLED THE CONTROVERSY 2EJMANEK 2ENNE AND 4RACY
3TOHLGREN
4HE INTEREST IN GLOBAL CHANGE THAT DEVELOPED IN ECOLOGY IN THE NINETIES WENT
BEYOND CONCERNS INVOLVING INCREASES IN ATMOSPHERIC #/ AND CLIMATE CHANGE
$UKES AND -OONEY EMPHASIZED THE GLOBAL DIMENSIONS OF BIOLOGICAL INVA
SIONS AND RESEARCHERS EMPHASIZED THE CAUSE AND EFFECT RELATIONSHIP THAT EXISTED
BETWEEN BIOLOGICAL INVASIONS AND OTHER TYPES OF GLOBAL CHANGE +OWARIK
!S A RESULT OF CONNECTIONS MADE BETWEEN BIOLOGICAL INVASIONS AND GLOBAL CHANGE
THE TOPIC OF BIOLOGICAL INVASION HISTORICALLY A RATHER NARROW AND SPECIALIZED AREA
OF RESEARCH HAS BECOME PART OF A MUCH LARGER AND MORE PROMINENT ENVIRONMEN
TAL AGENDA
)NVASION BIOLOGY
! NUMBER OF BOOKS PUBLISHED DURING THE S AND EARLY S AND LIKE
%LTONS BOOK WERE WRITTEN FROM AN EXPLICIT CONSERVATION AND ENVIRONMEN
TAL PERSPECTIVE AND AIMED AT THE GENERAL PUBLIC -ANY OF THESE BOOKS UTILIZED
SIMILAR EVOCATIVE IMAGERY TO CAPTURE THE PUBLICS ATTENTION %XAMPLES OF SUCH
BOOKS WRITTEN OR EDITED BY ECOLOGISTS OR SCIENCE JOURNALISTS INCLUDE ,IFE /UT OF
"OUNDS "IOINVASIONS IN A "ORDERLESS 7ORLD "RIGHT AND +ILLER !LGAE -EINESZ
.ATURE /UT OF 0LACE "IOLOGICAL )NVASIONS IN THE 'LOBAL !GE 6AN $RIESCHE AND
6AN $RIESCHE 4INKERING WITH %DEN ! .ATURAL (ISTORY OF %XOTICS IN !MERICA
4ODD AND 0LAGUES OF 2ATS AND 2UBBERVINES "ASKIN
7ITH THEIR LAMENTATION OVER THE LOSS OF NATIVE ENVIRONMENTS AND DECLINE OF
NATIVE SPECIES THE ABOVE BOOKS ILLUSTRATE THE INCREASINGLY STRONG SYNERGY THAT
HAD DEVELOPED BETWEEN RESTORATION ECOLOGY AND INVASION ECOLOGY PARTICULARLY IN
.ORTH !MERICA DURING THE NINETIES WITH THE OBJECTIVES OF EACH REINFORCING THOSE
OF THE OTHER 2ESTORATION ECOLOGYS EMPHASIS ON RESTORING ENVIRONMENTS WITH
NATIVE SPECIES AFlRMED THE IMPORTANCE OF INVASION ECOLOGY AND INVASION ECOLO
GYS EMPHASIS ON THE HARM CAUSED BY A SMALL PROPORTION OF INTRODUCED SPECIES
PROVIDED IMPORTANT JUSTIlCATION FOR RESTORATION ECOLOGYS PREFERENCE FOR NATIVE
SPECIES (OWEVER BY THE END OF THE NINETIES AND BEGINNING OF THIS CENTURY MORE
SCIENTISTS WERE QUESTIONING THE DICHOTOMOUS AND NORMATIVE NATURE OF A PERSPEC
TIVE THAT PREFERS SOME SPECIES OVER OTHERS ON THE BASIS OF THEIR GEOGRAPHY OF ORIGIN
'OULD 3LOBODKIN EMPHASIZING THE USEFUL CONTRIBUTIONS THAT SOME
INTRODUCED SPECIES MIGHT BE ABLE TO MAKE IN RESTORATION PROJECTS $!NTONIO AND
-EYERSON 3UCH OBJECTIONS WERE NOT NEW )N THE 3#/0% SYNTHESIS REPORT
$RAKE ET AL *AMES "ROWN MADE THE SAME ARGUMENTS REGARDING
THE POTENTIALLY POSITIVE VALUE OF SOME INTRODUCED SPECIES AND EXPRESSED CONCERN
OVER hA KIND OF IRRATIONAL XENOPHOBIA ABOUT INVADING ANIMALS AND PLANTS THAT
RESEMBLES THE INHERENT FEAR AND INTOLERANCE OF FOREIGN RACES CULTURES AND RELI
GIONSv AN hATTITUDEv HE SAID HE hDETECTEDv AT THE 3#/0% CONFERENCE !CTUALLY
%GLER RAISED THESE SAME CONCERNS FORTY
SEVEN YEARS EARLIER )N HIS ARTICLE
IN %COLOGY HE CRITICIZED THE COMMON RESORT TO HYPERBOLE AND MILITARY IMAGERY
IN DISCUSSIONS OF ALIEN SPECIES AND WARNED AGAINST hANTHROPOMORPHIC SENTIMEN
TALISM THAT BREEDS A SPIRIT OF DEFEATv $URING THE NINETIES WRITINGS OF INVASION
BIOLOGISTS ALSO ATTRACTED THE ATTENTION OF PHILOSOPHERS %SER 3AGOFF
WHO EXPRESSED SIMILAR CONCERNS OVER THE LANGUAGE AND VALUES THEY SAID PERME
ATED THE WRITINGS OF INVASION ECOLOGY &INALLY ALTHOUGH NO ECOLOGISTS QUESTIONED
THE FACT THAT THE EARTHS BIOTA WAS BECOMING INCREASINGLY HOMOGENIZED DUE TO
WIDESPREAD SPECIES INTRODUCTIONS THE COMMON CONTENTION THAT SPECIES INTRODUC
TIONS LEAD TO DECLINES IN REGIONAL AND LOCAL BIODIVERSITY WAS CHALLENGED BY SEVERAL
INVESTIGATORS 2OSENZWEIG $AVIS 3AX AND 'AINES A POINT THAT
HAD ACTUALLY BEEN MADE MUCH EARLIER BY (ARPER
"Y THE END OF THE NINETIES INVASION BIOLOGY WAS BECOMING INCREASINGLY INSTI
TUTIONALIZED WITH THE ESTABLISHMENT OF MORE NATIONAL AND INTERNATIONAL CONSER
VATION AND BIODIVERSITY INITIATIVES RELATING TO BIOLOGICAL INVASIONS 4HE 5NITED
.ATIONS SPONSORED CONFERENCE ON hINVASIVE ALIEN SPECIESv IN 4RONDHEIM .ORWAY
-! $AVIS
IN 3ANDLUND ET AL LED TO THE FORMATION IN THE FOLLOWING YEAR OF THE
'LOBAL )NVASIVE 3PECIES 0ROGRAMME ')30 ')30 WAS DEVELOPED EXPLICITLY AS A
CONSERVATION AND ENVIRONMENTAL INITIATIVE AS EVIDENCED BY ITS MISSION STATEMENT
@TO CONSERVE BIODIVERSITY AND SUSTAIN HUMAN LIVELIHOODS BY MINIMIZING THE SPREAD
AND IMPACT OF INVASIVE ALIEN SPECIES 4O DATE THE ')30 PROGRAM HAS RESULTED IN
PUBLICATIONS
)N THE %UROPEAN %NVIRONMENT !GENCY %%! HIGHLIGHTED THE INTRO
DUCTION OF ALIEN SPECIES AS ONE OF THE PRINCIPAL THREATS TO %UROPES BIODIVERSITY
IN ITS YEAR
END REPORT ON THE STATUS OF THE %UROPEAN ENVIRONMENT %%!
)N 0RESIDENT #LINTON SIGNED %XECUTIVE /RDER WHICH AUTHORIZED
53 FEDERAL AGENCIES TO PREVENT THE INTRODUCTION OF ALIEN INVASIVE SPECIES CONTROL
THE SPREAD OF ALIEN INVASIVE SPECIES IN THE 5NITED 3TATES AND RESTORE NATIVE SPECIES
AND HABITATS THAT HAD BEEN INVADED )N )5#. 4HE 7ORLD #ONSERVATION
5NION PASSED @'UIDELINES FOR THE 0REVENTION OF "IODIVERSITY ,OSS #AUSED BY
!LIEN )NVASIVE 3PECIES AND PUBLISHED THE GUIDELINES ON ITS WEBSITE IN )N
THE #OMMITTEE ON THE 3CIENTIlC "ASIS FOR 0REDICTING THE )NVASIVE 0OTENTIAL
OF .ONINDIGENOUS 0LANTS AND 0LANT 0ESTS IN THE 5NITED 3TATES PUBLISHED ITS lNAL
REPORT .ATIONAL 2ESEARCH #OUNCIL 4HE lNAL REPORT OF ')30S lRST PHASE
WAS PUBLISHED IN -OONEY ET AL AND BIOLOGICAL INVASIONS WAS THE THEME OF
THE ANNUAL MEETING OF !)"3 !MERICAN )NSTITUTE FOR "IOLOGICAL 3CIENCES
)T IS CLEAR THAT FORTY
SIX YEARS FOLLOWING THE PUBLICATION OF %LTONS CLASSIC
INTEREST IN BIOLOGICAL INVASIONS WAS STRONGER THAN EVER BOTH WITHIN AND OUTSIDE
THE SCIENTIlC COMMUNITY PARTICULARLY WITH RESPECT TO THE CONSERVATION AND
MANAGEMENT ASPECTS OF INTRODUCED SPECIES
,//+).' !(%!$ !.9 ,%33/.3 !.$ ).3)'(43 &2/- 4(% 0!34
!RE THERE ANY LESSONS WE CAN LEARN FROM THIS BRIEF LOOK OVER OUR SHOULDER #AN WE
GAIN ANY INSIGHTS THAT MIGHT HELP US CHART A PRODUCTIVE PATH FOR THE FUTURE ) THINK
SO /F COURSE EXACTLY WHAT ARE THOSE LESSONS AND INSIGHTS WILL LIKELY BE A MATTER OF
LIVELY DEBATE 4HUS THE THOUGHTS THAT FOLLOW ARE NOT INTENDED TO BE PRESCRIPTIVE
BUT RATHER TO FOCUS DISCUSSION ON HOW ECOLOGISTS INTERESTED IN BIOLOGICAL INVASIONS
MIGHT BEST PROCEED FROM THIS POINT FORWARD
4WO ROADS ONE LESS TRAVELED THAN THE OTHER
!FTER THE !SILOMAR CONFERENCE IN THOSE INTERESTED IN STUDYING SPECIES INTRO
DUCTIONS WERE PRESENTED WITH TWO DISTINCT PATHS 4HEY COULD TAKE THE STRONGLY CON
SERVATION AND ENVIRONMENTAL APPROACH ADVANCED BY %LTON OR THEY COULD ADOPT THE
MORE VALUE
NEUTRAL AND MORE STRICTLY SCIENTIlC APPROACH EXHIBITED BY 4HE 'ENETICS
OF #OLONIZING 3PECIES 4HE %LTONIAN PATH COULD BE DESCRIBED AS A TOP
DOWN DEDUCTIVE
APPROACH IN WHICH AN EFFORT WAS MADE TO APPLY GENERAL ECOLOGICAL THEORY AND PRIN
CIPLES TO BIOLOGICAL INVASIONS IN ORDER TO HELP DEVELOP CONTROL MANAGEMENT PROGRAMS
)NVASION BIOLOGY
FOR SPECIlC INVASIONS #ONVERSELY THE !SILOMAR PATH COULD BE CONSIDERED MORE OF
A BOTTOM
UP INDUCTIVE APPROACH IN WHICH INDIVIDUAL COLONIZATIONSINVASIONS ARE
EXAMINED IN AN EFFORT TO BETTER INFORM GENERAL ECOLOGICAL THEORY AND UNDERSTANDING
OF COMMUNITIES AND POPULATIONS !NOTHER WAY TO CONTRAST THE TWO PATHS MIGHT BE
TO CHARACTERIZE THE %LTONIAN PATH AS A 0LATONIC APPROACH AND THE !SILOMAR PATH AS
AN !RISTOTELIAN APPROACH SENSU 3AGOFF
!LTHOUGH THERE HAVE BEEN SOME EXCEPTIONS AND OCCASIONAL RESISTANCE BY
SOME ECOLOGISTS OVER THE YEARS INVASION ECOLOGY HAS BECOME INCREASINGLY MORE
ALLIED WITH THE %LTONIAN APPROACH PARTICULARLY IN THE 5NITED 3TATES DURING PAST
FEW DECADES 4HERE ARE PROBABLY MANY EXPLANATIONS FOR THIS %COLOGISTS SUCH AS
!LDO ,EOPOLD 2ACHEL #ARSON AND 0AUL %HRLICH WERE INSTRUMENTAL IN RAISING THE
PUBLIC CONSCIOUSNESS ON ENVIRONMENTAL ISSUES WHICH RESULTED IN THE EMERGENCE
OF THE ENVIRONMENTAL MOVEMENT DURING THE S 4HIS MOVEMENT ATTRACTED
THE ATTENTION NOT ONLY OF THE GENERAL PUBLIC BUT ALSO OF MANY OTHER ECOLOGISTS
7HEREAS MUCH OF THE FOCUS IN ECOLOGY IN THE S AND S HAD BEEN ON THEORETI
CAL ISSUES INVOLVING POPULATION DYNAMICS AND COMMUNITY ASSEMBLY DESIRE EVEN
PRESSURE TO APPLY ECOLOGICAL KNOWLEDGE TO ENVIRONMENTAL PROBLEMS BEGAN TO BE
EXPRESSED BOTH FROM WITHIN THE DISCIPLINE AND FROM SOCIETY AT LARGE 3OCIETY IN THE
53 AND ELSEWHERE HAD LONG APPLIED AN ECOLOGICAL APPROACH TO INTRODUCED PESTS
OF lSHERIES FORESTS AND AGRICULTURAL SYSTEMS 4HE %LTONIAN PATH EXTENDED THIS
APPROACH TO NATURAL SYSTEMS THAT WERE NOT NECESSARILY PROVIDING DIRECT ECONOMIC
BENElTS 7ITHOUT QUESTION THE FOUNDING OF THE 3#/0% INITIATIVE ON INVASIONS IN
THE EARLY S PLAYED A MAJOR ROLE IN DElNING THE CONSERVATION FOCUS OF INVA
SION BIOLOGY FROM THAT POINT ONWARD $URING THE S AND S %3! BEGAN TO
ADVOCATE A MORE ACTIVE PUBLIC ROLE IN GENERAL FOR ECOLOGISTS AND FOR ITSELF AS AN
ORGANIZATION )N MANY WAYS WHAT HAD BEEN A FAIRLY CLEAR DISTINCTION BETWEEN
hPUREv ECOLOGY AND hAPPLIEDv ECOLOGY IN THE S AND S DISSOLVED DURING THE
LATTER DECADES OF THE TWENTIETH CENTURY
$URING THE EIGHTIES AND NINETIES IN THE 5NITED 3TATES THERE WAS INCREASING PRES
SURE FOR ECOLOGISTS TO JUSTIFY THEIR RESEARCH IN A LARGER SOCIAL CONTEXT 4HIS WAS FELT
AT ALL LEVELS 4O JUSTIFY ITS BUDGET TO #ONGRESS 4HE .ATIONAL 3CIENCE &OUNDATION
NEEDED TO MORE CLEARLY ARTICULATE HOW THE RESEARCH IT FUNDED BENElTED THE PUBLIC
GOOD )N TURN IN ORDER TO GET FUNDED BY .3& INDIVIDUAL RESEARCHERS NEEDED TO
PROVIDE INCREASING EVIDENCE NOT ONLY OF THE SCIENTIlC VALUE OF THEIR RESEARCH BUT
ALSO OF ITS BROADER IMPACTS INCLUDING BENElTS TO SOCIETY )N ADDITION IRRESPECTIVE
OF THESE INFRASTRUCTURE PRESSURES SOME ECOLOGISTS MAY HAVE WANTED TO SEEK A MORE
PUBLIC FORUM FOR PERSONAL REASONS WHETHER OUT OF A SENSE OF SOCIAL RESPONSIBILITY
OR SIMPLY AN INTEREST IN PARTICIPATING IN POLICY
MAKING AND POLICY DISCUSSIONS
0ROBABLY ALL THESE FACTORS IN VARYING DEGREES PLAYED A ROLE IN THE DEVELOPMENT
OF THE STRONG ENVIRONMENTAL EMPHASIS THAT NOW CHARACTERIZES INVASION ECOLOGY
4ODAY INVASION ECOLOGY HAS BECOME NEARLY AN EQUAL SIBLING AT LEAST A CLOSE COUS
IN TO THE EXPLICITLY APPLIED lELDS OF CONSERVATION ECOLOGY AND RESTORATION ECOLOGY
/NE MIGHT CONSIDER THE THREE lELDS AS CONSTITUTING A KIND OF TRIUMVIRATE LEADING
THE EFFORTS TO PRESERVE THE EARTHS BIODIVERSITY
-! $AVIS
-OST MAJOR INVASION PUBLICATIONS AND CONFERENCES WHETHER TARGETED TO THE
GENERAL PUBLIC OR TO A MORE STRICTLY SCIENTIlC AUDIENCE ARE NOW EXPLICITLY FRAMED
FROM A CONSERVATION PERSPECTIVE USUALLY EMPHASIZING CURRENT BIODIVERSITY THREATS
AND POTENTIAL IMPENDING ECOLOGICAL CRISES OFTEN USING LANGUAGE AND IMAGERY
SIMILAR TO THAT USED BY %LTON IN %LTON WAS NOT THE FATHER OF INVASION ECOL
OGY BUT HE MIGHT BE CONSIDERED THE FOUNDER OF ONE PATH OF INVASION ECOLOGY
4HIS PATH THE CONSERVATION AND ENVIRONMENTAL PATH HAS BEEN WELL TRAVELED AND
MAINTAINED DURING PAST SEVERAL DECADES (OWEVER THE ALTERNATIVE PATH EMBOD
IED BY THE !SILOMAR CONFERENCE WHILE CERTAINLY NOT WHOLLY REJECTED HAS
NOT BEEN AS WELL TRAVELED 6ERMEIJ 4HE DIFFERENCE HAS PROBABLY BEEN MOST
APPARENT DURING THE PAST YEARS AND PARTICULARLY IN THE 5NITED 3TATES .O DOUBT
PART OF THE EXPLANATION FOR THIS DIFFERENCE IS THAT A LARGE NUMBER OF ECOLOGISTS ARE
EMPLOYED BY CONSERVATION GROUPS AND GOVERNMENTAL AGENCIES WHERE THEY WORK
PRIMARILY ON APPLIED PROBLEMS
0ROGRESS V S. EFFORT
)F ONE TAKES THE TIME TO GO BACK AND READ THE INVASION AND COLONIZATION LITERATURE
FROM TWENTY THIRTY AND FORTY YEARS AGO ONE IS STRUCK BY HOW LITTLE THE QUES
TIONS AND UNFORTUNATELY OFTEN THE ANSWERS SEEM TO HAVE CHANGED OVER TIME
)T IS NOT EASY TO lND EITHER QUESTIONS OR ANSWERS GENERALIZATIONS AND THEORY
MADE IN RECENT YEARS THAT ARE NOT IN THE LITERATURE OF THIRTY OR FORTY YEARS AGO OR
EVEN EARLIER THAN THAT 4HE MISSION OF THE 3#/0% INVASION PROGRAM WAS TO TRY TO
ANSWER THREE QUESTIONS WHAT FACTORS DETERMINE WHETHER OR NOT A SPECIES WILL BE
AN INVADER WHAT PROPERTIES DETERMINE WHETHER OR NOT A PARTICULAR ENVIRONMENT
WILL PRONE TO INVASION AND HOW CAN THE KNOWLEDGE GAINED FROM ANSWERING THESE
QUESTIONS HELP MANAGEMENT EFFORTS 4HE lRST QUESTION WAS ACTUALLY THE ORGANIZ
ING QUESTION OF THE !SIMOLAR SYMPOSIUM AND THE SECOND WAS ARTICULATED
BY %LTON HIMSELF
4HERE IS NO QUESTION THAT OUR UNDERSTANDING OF THE PROCESS OF BIOLOGICAL INVA
SIONS HAS INCREASED CONSIDERABLY DURING THE PAST FORTY
SIX YEARS PARTICULARLY OUR
KNOWLEDGE OF THE DETAILS INVOLVED IN PARTICULAR BIOLOGICAL INVASIONS (OWEVER THE
EXTENT TO WHICH THIS UNDERSTANDING HAS ENHANCED OUR ABILITY TO DEVELOP EFFECTIVE
MANAGEMENT STRATEGIES THE RD 3#/0% OBJECTIVE IS LESS CLEAR AND DEPENDS ON
WHOM YOU ASK 3OME ECOLOGISTS MAINTAIN THAT GOOD PROGRESS IS BEING MADE AND
THAT WITH THE APPLICATION OF NEW ANALYSIS TOOLS AND APPROACHES SUCH AS SPECIES
SCREENING $AEHLER ET AL AND THE USE OF CLIMATIC ENVELOPES 0ETERSON
THERE IS GOOD REASON TO BE OPTIMISTIC ABOUT INVASION ECOLOGISTS ABILITY TO EFFEC
TIVELY INFORM LAND MANAGERS /THERS ARE NOT SO SANGUINE ARGUING THAT THE TYPE OF
KNOWLEDGE AND UNDERSTANDING GAINED SO FAR ARE SELDOM USEFUL IN SPECIlC MANAGE
MENT EFFORTS -OYLE AND ,IGHT 3IMBERLOFF AND 3TILING MEANING THAT
EFFORTS BY INVASION ECOLOGISTS TO EFFECTIVELY INFORM LAND MANAGERS CONTINUE TO BE
FRUSTRATED 7ILLIAMSON (ULME
)NVASION BIOLOGY
#/.#,53)/.
7)4( ! &%7 35''%34)/.3 4/ #/.3)$%2
)NVASION ECOLOGY AND CONSERVATION
!LL THE EVIDENCE TO DATE INDICATES THAT IT IS UNLIKELY THAT ECOLOGISTS WILL BE ABLE TO
DEVELOP A GENERAL THEORY OF BIOLOGICAL INVASIONS THAT CAN BE OF SIGNIlCANT PRACTI
CAL VALUE TO LAND MANAGERS -UCH RESEARCH SEEMS TO HAVE AFlRMED !SA 'RAYS
OBSERVATIONS OF WEEDS MORE THAN A CENTURY AGO 4RYING TO DETERMINE SPECIlC
CHARACTERISTICS THAT MAKE SOME PLANTS WEEDS 'RAY CONCLUDED THAT HE
hCOULD DISCERN NOTHING IN THE PLANT ITSELF THAT WOULD GIVE IT AN ADVANTAGEv
#ONTINUING HE WROTE hTHE REASONS FOR PREDOMINANCE MAY BE ALMOST AS DIVERSE AS
THE WEEDS THEMSELVESv
7ITH RESPECT TO ITS GOAL OF ASSISTING LAND MANAGERS IN THE CONTROL AND MANAGE
MENT OF INTRODUCED SPECIES INVASION ECOLOGY MIGHT CONSIDER EMPHASIZING LESS THE
DEVELOPMENT OF GENERAL THEORY AND MORE THE ACQUISITION OF SYSTEM
SPECIlC KNOWL
EDGE 4HE FACT THAT INVASION ECOLOGY HAS CONSISTED PRIMARILY AS A SERIES OF CASE
STUDIES HAS GENERALLY BEEN VIEWED AS A WEAKNESS OF THE lELD 7ILLIAMSON
"UT IN REALITY IT MAY BE VERY DIFlCULTY PERHAPS EVEN IMPOSSIBLE FOR THE lELD TO
MOVE MUCH BEYOND THIS AT LEAST AT THE LEVEL OF PROVIDING USEFUL MANAGEMENT
INFORMATION )N THIS CASE INVASION ECOLOGISTS MIGHT BE BETTER OFF VIEWING THE hSYS
TEM SPECIlCv NATURE OF THE lELDS KNOWLEDGE AS A STRENGTH 2ATHER THAN LAMENTING
THE CASE
STUDY NATURE OF THE lELD ECOLOGISTS MIGHT TRY TO MAKE INCREASING SYSTEM
SPECIlC KNOWLEDGE A PRIMARY OBJECTIVE EG 2EJMANEK AND 2ICHARDSON
7HILE A LAND MANAGER TRYING TO MANAGE THE SPECIES COMPOSITION OF A GRASSLAND
MAY NOT lND INVASION THEORY PARTICULARLY HELPFUL THE MORE SYSTEM
SPECIlC KNOWL
EDGE THAT IS AVAILABLE IN HISHER TOOLBOX THE MORE SUCCESSFUL THE MANAGEMENT
EFFORTS WILL BE 4O THE EXTENT THAT INVASION ECOLOGY WANTS TO INFORM CONSERVATION
EFFORTS IT MAY ALSO MAKE SENSE TO FOR INVASION ECOLOGISTS TO BECOME MORE INVOLVED
IN DEVELOPING FORMALIZED AND COMPREHENSIVE MONITORING PROGRAMS WHICH HAVE
THE POTENTIAL OF GREATLY ENHANCING THE EFFECTIVENESS OF MANAGEMENT AND CONTROL
EFFORTS ,ONSDALE AND -ILTON
)NVASION ECOLOGY AND SCIENCE
4HE CONSERVATION AND ENVIRONMENTAL EMPHASIS IN INVASION ECOLOGY DESCRIBED
ABOVE HAS BEEN MOTIVATED BY THE CONVICTION THAT ECOLOGICAL KNOWLEDGE AND THEORY
CAN BE USED TO BETTER UNDERSTAND AND PREDICT BIOLOGICAL INVASIONS 4HE !SILOMAR
APPROACH WAS MOTIVATED BY THE OPPOSITE CONVICTION THAT BIOLOGICAL COLONIZA
TIONSINVASIONS CAN BE VIEWED AS NATURAL EXPERIMENTS AND USED TO INFORM MORE
GENERAL ECOLOGICAL THEORY AND UNDERSTANDING !LTHOUGH THERE WAS SOME INTER
EST AMONG ECOLOGISTS IN THE EARLY EIGHTIES TO CONTINUE THIS EMPHASIS "ROWN AND
-ARSHALL 3IMBERLOFF -OULTON AND 0IMM THIS APPROACH HAS
NOT BEEN AS WELL NURTURED OVER THE YEARS BY INVASION ECOLOGY COMPARED TO THE
-! $AVIS
%LTONIAN CONSERVATION APPROACH 4HE REINVIGORATION OF A SCIENCE OF COLONIZATION
AND SPECIES SPREAD AS A WAY TO STUDY CORE CONCEPTS AND THEORIES OF EVOLUTION AND
ECOLOGY WOULD SEEM TO BE AN EXCITING OPTION FOR INVASION ECOLOGY 2OBERT &ROSTS
TRAVELER COULD ONLY CHOOSE ONE ROAD BUT THERE IS NO REASON THAT INVASION ECOLOGY
COULDNT EMBRACE BOTH A CONSERVATION AND A MORE STRICTLY SCIENTIlC PATH )T HAS
BEEN FORTY YEARS SINCE THE HISTORIC !SILOMAR SYMPOSIUM ON COLONIZATION
BIOLOGY PERHAPS IT IS TIME TO ORGANIZE ANOTHER ONE
)NVASION ECOLOGY AND LANGUAGE
(IGH RATES OF SPECIES SPREAD AND INTRODUCTIONS ARE HERE TO STAY )N SOME RESPECTS
ONE MIGHT QUESTION THE SENSE OF DElNING AN ECOLOGICAL SUBDISCIPLINE AREA AROUND
SPECIES INTRODUCTIONS $UE TO THE ABUNDANCE OF INTRODUCED SPECIES IN ENVIRON
MENTS THROUGHOUT THE WORLD WHICH IS ONLY GOING TO INCREASE IN THE FUTURE IT
IS BECOMING INCREASINGLY DIFlCULT TO UNDERTAKE ANY ECOLOGICAL STUDY IN WHICH
INTRODUCED SPECIES CAN BE IGNORED 4HAT BIOTIC GLOBALIZATION IS NOW THE WORLD IN
WHICH WE NOW LIVE CERTAINLY DOES NOT MEAN WE SHOULD CEASE EFFORTS TO CONTROL AND
MANAGE THE SMALL NUMBER OF INTRODUCED SPECIES THAT DO CAUSE GREAT HEALTH ECO
NOMIC AND ECOLOGICAL HARM %WEL ET AL (OWEVER IT DOES RAISE QUESTIONS
AS TO THE LOGIC OF DICHOTOMOUSLY SPLITTING SPECIES INTO NATIVES AND EXOTICS AND
EVEN OF USING ANY TERMINOLOGY THAT SINGLES OUT RECENTLY INTRODUCED SPECIES AS A
GROUP EG INDIGENOUS AND NON
INDIGENOUS GIVEN THAT SO FEW OF THESE SPECIES ARE
PROBLEMATIC !T THE SAME TIME PARTICULAR RESEARCH EFFORTS WILL NEED TO DISTINGUISH
SPECIES THAT HAVE BEEN PRESENT IN A REGION FOR A LONG TIME FROM THOSE THAT HAVE
RECENTLY ARRIVED THROUGH ONE MEANS OR ANOTHER 4HUS FROM A PRACTICAL PERSPEC
TIVE CATEGORIZING SPECIES ON THEIR BASIS OF GEOGRAPHIC ORIGIN WILL BE NECESSARY IN
SOME CASES 7EBB 2ICHARDSON ET AL #OLAUTTI AND -AC)SAAC
0YEK ET AL
&ROM A STRICTLY SCIENTIlC POINT OF VIEW IT WOULD SEEM DIFlCULT TO ARGUE AGAINST
THE VALUE OF RETURNING TO THE MORE VALUE
NEUTRAL TERMINOLOGY USED AT THE
!SILOMAR SYMPOSIUM )F ONE IS STUDYING SPECIES INTRODUCTIONS AS A WAY ILLUMI
NATE BASIC ECOLOGICAL IDEAS AND THEORY THEN TERMS SUCH AS hINTRODUCED SPECIESv
hCOLONIZERSv hNEW SPECIESv hNEW RESIDENTSv AND hGEOGRAPHIC SPREADv SEEM MORE
APT THAN THE LANGUAGE OFTEN ASSOCIATED WITH THE %LTONIAN PATH EG hEXOTICSv
hALIENSv AND hINVASIONSv 7HETHER OR NOT ECOLOGISTS SHOULD MODIFY THEIR LANGUAGE
WHEN ADVOCATING ENVIRONMENTAL CONCERNS WILL LIKELY BE A MATTER OF DISAGREEMENT
WITHIN THE lELD 3OME WILL BE CONCERNED THAT DESPITE THE BEST OF INTENTIONS THE
lELDS CREDIBILITY AND STATURE AS A SCIENTIlC DISCIPLINE ARE BEING COMPROMISED WHEN
ECOLOGISTS MAKE THEIR CASE TO THE PUBLIC AND TO ONE ANOTHER USING VALUE
LADEN
TERMINOLOGY AN APPROACH MORE TYPICALLY ASSOCIATED WITH ENVIRONMENTAL
ACTION
GROUPS 4HOSE WITH THIS CONCERN WOULD PROBABLY BE MORE COMFORTABLE IF THE lELD
DROPPED THE INVASIONALIEN TERMINOLOGY ALTOGETHER AND REPLACED IT WITH A TERMI
NOLOGY IN WHICH VALUES WERE ARTICULATED MORE EXPLICITLY THAN IMPLICITLY 5SING
THIS APPROACH ECOLOGISTS MIGHT REFER TO PROBLEM SPECIES SUCH AS ZEBRA MUSSELS
)NVASION BIOLOGY
AND *APANESE KNOTWEED AS @HARMFUL NEW SPECIES OR @HARMFUL INTRODUCED SPECIES
AS OPPOSED TO USING A TERMINOLOGY IN WHICH VALUES ARE MORE SUBTLY IMPLICATED
EG @INVASIVE ALIEN SPECIES /THERS WILL ARGUE THAT METAPHORIC AND EVEN AT TIMES
HYPERBOLIC LANGUAGE IS NEEDED TO CAPTURE THE ATTENTION OF POLICY MAKERS AND
THE GENERAL PUBLIC AND TO PERSUADE THEM OF THE NEED TO ACT JUST AS %LTON
DID WHEN HE CHARACTERIZED THE GLOBAL SPREAD OF SPECIES AS A hCONVULSION IN THE
WORLDS mORA AND FAUNAv AND FRAMED THE CHALLENGE CONFRONTING US AS A hBATTLEv
TO hDETERMINE THE FATE OF THE WORLDv THE LATTER A QUOTE FROM #ONAN $OYLES BOOK
4HE ,OST 7ORLD
,OOKING AHEAD
4HE PAST TWENTY YEARS HAVE SEEN EXTRAORDINARY DEVELOPMENT IN THE lELD OF INVA
SION ECOLOGY 7ITH NUMEROUS NATIONAL AND INTERNATIONAL INITIATIVES THE CREATION
OF TWO NEW JOURNALS AND THE COUNTLESS NUMBER OF SYMPOSIA CONFERENCES AND
WORKSHOPS DEVOTED TO BIOLOGICAL INVASIONS NOT TO MENTION THE THOUSANDS OF INVA
SION PAPERS PUBLISHED THERE HAS BEEN LITTLE TIME FOR PARTICIPANTS TO STEP BACK AND
REmECT ON THE WAY THAT THE lELD HAS DEVELOPED 0ERHAPS NOW WOULD BE A GOOD TIME
TO DO THIS BEFORE MOVING AHEAD 4O WHAT EXTENT IS INVASION ECOLOGY BECOMING
LARGELY AN APPLIED DISCIPLINE LIKE CONSERVATION BIOLOGY AND RESTORATION ECOLOGY
4O WHAT EXTENT SHOULD THE lELD TRY TO REVITALIZE EFFORTS TO STUDY SPECIES INTRO
DUCTIONS AS A WAY TO ILLUMINATE MORE GENERAL ECOLOGICAL THEORY A LA THE
!SILOMAR SYMPOSIUM APPROACH 4HE CURRENT BOOK AND ONE SIMILAR TO IT 3AX
ET AL IN PRESS APPEAR TO ILLUSTRATE A REVIVED INTEREST IN THIS APPROACH )S IT POS
SIBLE FOR INVASION ECOLOGY AND INDIVIDUAL ECOLOGISTS TO EMBRACE BOTH A STRONGLY
VALUE
BASED ENVIRONMENTAL PATH AND A MORE VALUE
NEUTRAL SCIENTIlC APPROACH TO
SPECIES INTRODUCTIONS 3HOULD AN EFFORT BE MADE TO KEEP THE PATHS SEPARATE IN THE
WAY THAT THE NEWS PAGES AND EDITORIAL PAGES ARE KEPT SEPARATE IN A NEWSPAPER
OR SHOULD THE TWO PATHS BE INTEGRATED 7HAT LANGUAGE AND IMAGERY SHOULD
ECOLOGISTS UTILIZE IN THEIR DISCOURSE AMONG THEMSELVES AND WITH THE PUBLIC 4HESE
ARE SOME OF THE QUESTIONS THAT THE lELD OF INVASION ECOLOGY MIGHT WANT TO ADDRESS
WAY BEFORE MOVING AHEAD
!#+./7,%$'-%.43
7RITING A HISTORICAL REVIEW OF A SCIENTIlC DISCIPLINE IS A CHALLENGE UNDER ANY CIR
CUMSTANCES $OING SO WHILE MOST OF THE PARTICIPANTS ARE STILL ALIVE AND ACTIVE IN
THE lELD IS A PARTICULARLY DAUNTING SOME MIGHT SAY FOOLISH ENDEAVOR ) THANK
THE MANY COLLEAGUES WHO READ EARLIER DRAFTS AND PROVIDED THOUGHTFUL AND OFTEN
DETAILED COMMENTS AND SUGGESTIONS PARTICULARLY #URT $AEHLER $ICK -ACK 0ETER
0YEK $AVE 2ICHARDSON -ARK 7ILLIAMSON -ARCEL 2EJMNEK AND $AN 3IMBERLOFF
4HE THOROUGHNESS OF THIS CHAPTER WOULD NOT HAVE BEEN POSSIBLE WITHOUT THEIR
INPUT ) ALSO THANK %D 7ILSON FOR SHARING HIS MEMORIES OF THE !SILOMAR
-! $AVIS
SYMPOSIUM 4HERE IS NO SINGLE CORRECT CHARACTERIZATION OF THE PAST AND MY COL
LEAGUES HAVE NOT ALWAYS FULLY AGREED WITH THE ONE ) PRESENTED .EVERTHELESS THEY
WERE ALWAYS BEEN GENEROUS AND GRACIOUS IN THEIR ADVICE DURING THE WRITING OF THIS
CHAPTER FOR WHICH ) AM INDEBTED 2ESEARCH AND WRITING OF THIS CHAPTER WAS PAR
TIALLY SUPPORTED BY THE .ATIONAL 3CIENCE &OUNDATION UNDER 'RANT .O
2%&%2%.#%3
!LBRECT ! " 3OME OBSERVATIONS ON FACTORS ASSOCIATED WITH SURVIVAL OF STRIPED BASS
EGGS AND LARVAE #ALIFORNIA &ISH AND 'AME
"AILEY ( ( 4HE ARMADILLO IN &LORIDA AND HOW IT REACHED THERE *OURNAL OF
-AMMOLOGY
"AKER ( ' 4HE EVOLUTION OF WEEDS !NNUAL 2EVIEW OF %COLOGY AND 3YSTEMATICS
"AKER ( ' AND ' , 3TEBBINS EDS 4HE %VOLUTION OF #OLONIZING 3PECIES !CADEMIC
0RESS .EW 9ORK .9
"ASKIN 9 ! 0LAGUE OF 2ATS AND 2UBBERVINES 4HE 'ROWING 4HREAT OF 3PECIES )NVASIONS
3HEARWATER "OOKS)SLAND 0RESS 7ASHINGTON $ #
"AZZAZ & ! ,IFE HISTORY OF COLONIZING PLANTS SOME DEMOGRAPHIC GENETIC AND PHYSI
OLOGICAL FEATURES 0AGES
IN ( ! -OONEY AND * ! $RAKE EDITORS %COLOGY OF
"IOLOGICAL )NVASIONS OF .ORTH !MERICA AND (AWAII 3PRINGER
6ERLAG .EW 9ORK .9
"INGGELI 0 -ISUSE OF TERMINOLOGY AND ANTHROPOMORPHIC CONCEPTS IN THE DESCRIPTION
OF INTRODUCED SPECIES "ULLETIN OF THE "RITISH %COLOGICAL 3OCIETY
"IRCH , # %VOLUTIONARY OPPORTUNITY FOR INSECTS AND MAMMALS IN !USTRALIA 0AGES
IN ( ' "AKER AND ' , 3TEBBINS EDITORS 4HE %VOLUTION OF #OLONIZING
3PECIES !CADEMIC 0RESS .EW 9ORK .9
"OOTH " $ 3 $ -URPHY AND # * 3WANSON 7EED %COLOGY IN .ATURAL AND
!GRICULTURAL 3YSTEMS #!") 0UBLISHING 7ALLINGORD /XON 5+
"RADSHAW ! $ 2 . (UMPHRIES - 3 *OHNSON AND 2 $ 2OBERTS 4HE RESTORATION
OF VEGETATION ON DERELICT LAND PRODUCED BY INDUSTRIAL ACTIVITY 0AGES
IN - 7
(OLDGATE AND - * 7OODMAN EDITORS 4HE "REAKDOWN AND 2ESTORATION OF %COSYSTEMS
0ROCEEDINGS #ONFERENCE 2EHABILITATION OF 3EVERELY $AMAGED ,AND AND &RESHWATER
%COSYSTEMS IN 4EMPERATE :ONES 2EYKJAVIK )CELAND *ULY 0LENUM 0RESS .EW
9ORK 53!
"RIGHT # ,IFE /UT OF "OUNDS "IOINVASION IN A "ORDERLESS 7ORLD .EW 9ORK 7 7
"ROWN ! ( $ AND $ 2 -ARSHALL %VOLUTIONARY CHANGES ACCOMPANYING COLONIZA
TION IN PLANTS 0AGES
IN ' ' % 3CUDDER AND * , 2EVEAL EDITORS %VOLUTION
4ODAY #ARNEGIE
-ELLON 5NIVERSITY 0RESS 0ITTSBURGH 0!
"RYANT ( # 4HE OPOSSUM REACHES 3AN $IEGO #OUNTY #ALIFORNIA &ISH AND 'AME
"UMP ' AND 7 ( "OHL 3UMMARY OF FOREIGN GAME BIRD PROPAGATION AND LIBERA
TIONS
3PECIAL 3CIENTIlC 2EPORT 7ILDLIFE .O "UREAU OF 3PORT &ISHERIES
AND 7ILDLIFE
)NVASION BIOLOGY
"URDON * * AND ' ! #HILVERS 0RELIMINARY STUDIES ON A NATIVE !USTRALIAN EUCALYPT
FOREST INVADED BY EXOTIC PINES /ECOLOGIA
#AMPBELL A # AND 2 & ' /RMOND 4HE THREAT OF THE h#ROWN
OF
THORNSv STARlSH
!CANTHASTER PLANCI TO CORAL REEFS IN THE )NDO
0ACIlC AREA /BSERVATIONS ON A NORMAL
POPULATION IN THE 2ED 3EA "IOLOGICAL #ONSERVATION
#ARLTON * 4 -ARINE BIOINVASIONS THE ALTERATION OF MARINE ECOSYSTEMS BY NONINDIG
ENOUS SPECIES /CEANOGRAPHY
#ARLTON * 4 "IOLOGICAL INVASIONS AND CRYPTOGENIC SPECIES %COLOGY
#ARLTON * 4 -OLLUSCAN INVASIONS IN MARINE AND ESTUARINE COMMUNITIES -ALACOLOGIA
#HILD , * ( "ROCK ' "RUNDU + 0RACH 0 0YEK 0 - 7ADE AND - 7ILLIAMSON
EDITORS 0LANT )NVASIONS %COLOGICAL 4HREATS AND -ANAGEMENT 3OLUTIONS
"ACKUYS 0UBLISHERS ,EIDEN 4HE .ETHERLANDS
#HRISTIE 7 * ,AKE /NTARIO EFFECTS OF EXPLOITATION INTRODUCTIONS AND EUTROPHICA
TION ON THE SALMONID COMMUNITY *OURNAL OF THE &ISHERIES 2ESEARCH "OARD OF #ANADA
#LEMENT % * AND - # &OSTER !LIEN 0LANTS OF THE "RITISH )SLES ! 0ROVISIONAL
#ATALOGUE OF 6ASCULAR 0LANTS EXCLUDING GRASSES "OTANICAL 3OCIETY OF THE "RITISH )SLES
,ONDON
#OLAUTTI 2 ) AND ( * -AC)SAAC ! NEUTRAL TERMINOLOGY TO DElNE @INVASIVE SPECIES
$IVERSITY AND $ISTRIBUTIONS
#OURTENAY 7 2 *R AND # 2 2OBINS %XOTIC ORGANISMS !N UNSOLVED COMPLEX
PROBLEM "IOSCIENCE
#RAWLEY - * 4HE POPULATION BIOLOGY OF INVADERS 0HILOSOPHICAL 4RANSACTIONS OF THE
2OYAL 3OCIETY "
#RAWLEY - * 7HAT MAKES A COMMUNITY INVASIBLE 3YMPOSIA OF THE "RITISH
%COLOGICAL 3OCIETY
$AEHLER # # AND $ #ARINO $ (YBRIDIZATION BETWEEN NATIVE AND ALIEN PLANTS AND
ITS CONSEQUENCES 0AGES
IN * , ,OCKWOOD * , AND - -C+INNEY EDITORS "IOTIC
(OMOGENIZATION +LUWER!CADEMIC0LENUM 0UBLISHERS .EW 9ORK
$AEHLER # # AND $ ! #ARINO 0REDICTING INVASIVE PLANTS 0ROSPECTS FOR A GENERAL
SCREENING SYSTEM BASED ON CURRENT REGIONAL MODELS "IOLOGICAL )NVASIONS
$AEHLER # # * 3 $ENSLOW 3 !NSARI AND ( +UO ! RISK
ASSESSMENT SYSTEM FOR
SCREENING OUT INVASIVE PEST PLANTS FROM (AWAII AND OTHER 0ACIlC )SLANDS #ONSERVATION
"IOLOGY
$AUGHERTY ( % 4HE IMPACT OF MAN ON THE ZOOGEOGRAPHY OF %L 3ALVADOR "IOLOGICAL
#ONSERVATION
$!NTONIO # AND , ! -EYERSON %XOTIC PLANT SPECIES AS PROBLEMS AND SOLUTIONS IN
ECOLOGICAL RESTORATION A SYNTHESIS 2ESTORATION %COLOGY
$AVIS - ! "IOTIC GLOBALIZATION DOES COMPETITION FROM INTRODUCED SPECIES THREATEN
BIODIVERSITY "IOSCIENCE
$AVIS - ! * 0 'RIME AND + 4HOMPSON &LUCTUATING RESOURCES IN PLANT COMMUNI
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! MAJOR GOAL OF INVASIONS ECOLOGY IS TO EXPLAIN WHY SOME SPECIES ARE MORE SUCCESS
FUL AS INVADERS THAN OTHERS AND ULTIMATELY TO BE ABLE TO PREDICT TRAITS PREDISPOSING
A SPECIES TO SUCCESS IN ADVENTIVE ENVIRONMENTS 5NFORTUNATELY TO DATE FEW CLEAR
GENERALITIES EXIST AND BIOLOGISTS ARE FAR FROM BEING ABLE TO PREDICT FUTURE INVASIONS
2ECENTLY SEVERAL GROUPS HAVE SHOWN HOW THE LACK OF CONSENSUS ON BASIC TERMI
NOLOGY HAS SERIOUSLY LIMITED PROGRESS IN AND APPLICATION OF INVASIONS ECOLOGY AS
WELL AS ITS CAPACITY TO DRAW SYSTEMATICALLY ON THEORY FROM ECOLOGICAL DISCIPLINES
$AVIS AND 4HOMPSON 2ICHARDSON ET AL A $AEHLER A $AVIS AND
4HOMPSON 2EJMNEK ET AL #OLAUTTI AND -AC)SAAC
7ITHIN THE LITERATURE IN REGARD TO VERY SIMILAR ECOLOGICAL SITUATIONS A PARTICU
LAR SPECIES MAY BE REGARDED AS ALIEN EXOTIC INVASIVE COLONIZING WEEDY INTRO
DUCED NON
INDIGENOUS IMPORTED OR NON
NATIVE -OREOVER A WEEDY SPECIES MAY
-7 #ADOTTE ET AL EDS #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY n
3PRINGER 0RINTED IN THE .ETHERLANDS
(4 -URPHY ET AL
ALSO BE REFERRED TO AS A NATURALIZED TRANSFORMER SLEEPER ESTABLISHED CASUAL
PERSISTENT NOXIOUS NATIVE ENVIRONMENTAL OR AGRICULTURAL WEED -ANY OF
THESE TERMS CONNOTE ONLY THAT A SPECIES IS NOVEL TO A PARTICULAR AREA /THERS CARRY
IMPLICATIONS ABOUT ACTUAL OR POTENTIAL IMPACT IN NOVEL HABITATS SEE $AVIS AND
4HOMPSON ;= AND #OLAUTTI AND -AC)SAAC ;= FOR A TREATMENT OF OTHER
TERMS (ERE WE ARE CONCERNED PRIMARILY WITH SPECIES THAT ARE TERMED @INVASIVE OR
@INVADERS 4HE SCOPE OF THIS NOTION ALONE COVERS MUCH GROUND AND IN RECENT YEARS
DEBATE OVER THE CRITERIA FOR REGARDING A SPECIES AS INVASIVE HAS RECEIVED MUCH
ATTENTION EG $AVIS AND 4HOMPSON 2ICHARDSON ET AL A $AVIS AND
4HOMPSON 2EJMNEK ET AL 0YEK ET AL AND THE TERM CONTINUES
TO BE USED INCONSISTENTLY 4HE DEBATE ON TERMINOLOGY IS MORE THAN SIMPLE SEMAN
TICS BECAUSE THE USE OF INCONSISTENT TERMS IMPAIRS UNDERSTANDING OF THE PROCESSES
THEY ARE MEANT TO DESCRIBE #OLAUTTI AND -AC)SAAC
#OMPARATIVE ANALYSIS OF ALIEN SPECIES IN REGIONAL mORAS HAS BEEN USED TO
CHARACTERIZE BOTH INVASIVE SPECIES AND INVADED ECOSYSTEMS EG $AEHLER
0YEK ,ONSDALE (OWEVER SUCH COMPARISONS ARE mAWED IF THE ALIEN
SPECIES IN THESE mORAS HAVE NOT BEEN CLASSIlED IN THE SAME WAYS 2ESEARCHERS ARE
FREQUENTLY UNABLE TO DISTINGUISH CLEARLY THE PARTICULAR CHARACTERISTICS OF ALIEN
INVADERS 4HOMPSON ET AL DISCREPANCIES IN TERMINOLOGY MAY CONTRIBUTE TO
THIS PROBLEM AND CONFOUND META ANALYSES
4O BE MOST USEFUL IT IS LOGICAL THAT THE TERMINOLOGY USED TO DESCRIBE SPECIES
AT VARIOUS STAGES OF INVASION SHOULD CLOSELY REmECT THE PROCESS BY WHICH INVA
SIONS OCCUR AND THE VARIOUS CHARACTERISTICS OF THOSE STAGES 4HE lRST PART OF THIS
CHAPTER IS A REVIEW OF THE STAGES IN THE PROCESS OF INVASION ,ATER WE TEST WHETHER
TWO RECENT CLASSIlCATION SCHEMES PROPOSED BY 2ICHARDSON ET AL A AND SEE
0YEK ET AL AND $AVIS AND 4HOMPSON ARE OPERATIONAL GIVEN A RELA
TIVELY RICH SPECIES DATASET AND WHETHER THEY SUCCEED IN IDENTIFYING THE SAME SETS OF
SPECIES 7E CLASSIFY A SAMPLE OF THE EXOTIC mORA OF /NTARIO USING THE TWO SCHEMES
7E IDENTIFY LIMITS TO INFORMATION AVAILABLE IN THIS SPECIES DATASET AND SHOW HOW
THE TWO CLASSIlCATION SYSTEMS CAN LEAD TO DIFFERENT CONCLUSIONS
)T IS NECESSARY THAT FOR THE PURPOSES OF CLARITY WE PROVIDE PRELIMINARY DElNI
TIONS OF CERTAIN TERMS WE WILL USE IN ORDER TO REVIEW THE INVASION PROCESS BEFORE
LOOKING IN MORE DEPTH AT THE DIFFERENT APPROACHES TO CLASSIFYING SPECIES AT DIFFERENT
STAGES OF THE PROCESS (ERE WE PRESENT SOME OF THE MOST COMMON TERMS TO DESCRIBE
GENERALLY THE DIFFERENT STAGES OF INVASION THOUGH WE REALIZE THAT THESE TERMS
ARE OFTEN USED INTERCHANGEABLY OR AS SYNONYMS FOR OTHER TERMS 4ERMS INCLUDE
hINTRODUCTIONv INTRODUCED SPECIES ARE THOSE THAT APPEAR OUTSIDE THEIR NATIVE
RANGE hESTABLISHMENTv ESTABLISHED SPECIES PERSIST OVER RELATIVELY LONG PERIODS
OUTSIDE THEIR NATIVE RANGE hNATURALIZATIONv NATURALIZED SPECIES REPRODUCE AND
SUSTAIN POPULATIONS OVER RELATIVELY LONG PERIODS AND hINVASIONv INVASIVE SPECIES
ARE WIDESPREAD AND ABUNDANT OUTSIDE THEIR NATIVE RANGE
)N THE lNAL SECTION OF THE CHAPTER WE DESCRIBE A WAY TO VISUALIZE AND MODEL
THE INVASIONS PROCESS BASED ON THE STATUS OF POPULATIONS OF NON NATIVE SPECIES AT
A SPECIlC SPATIAL AND TEMPORAL SCALE 7E DRAW ON THE @BLINKING LIGHTS METAPHOR
)NVASIVENESS IN EXOTIC PLANTS
OF METAPOPULATION ECOLOGY AND SHOW HOW THIS MAY BE USEFUL IN DESCRIBING THE
INVASIONS PROCESS h"LINKING LIGHTSv DESCRIBES THE COLONIZATION LIGHT ON AND
EXTINCTION LIGHT OFF OF POPULATIONS IN PATCHES OF HABITAT OVER TIME ACROSS A
REGIONAL LANDSCAPE 4HUS WHEN POPULATION TURNOVER IS HIGH HIGH COLONIZATION
AND HIGH EXTINCTION POPULATIONS ARE RAPIDLY BLINKING ON AND OFF WHEREAS WHERE
TURNOVER IS SLOWER HIGH COLONIZATION LOW EXTINCTION POPULATIONS BLINK ON AND
REMAIN ON FOR EXTENDED PERIODS )T IS POSSIBLE TO ENVISION THIS BLINKING ON OF POPU
LATIONS DURING THE INVASION PROCESS lRST AS THE NON
NATIVE SPECIES IS INTRODUCED
THEN ESTABLISHES NATURALIZES AND lNALLY AS IT BECOMES INVASIVE 4HE PROBABILITY OF
THE POPULATION BLINKING OFF WOULD DECREASE WITH TIME AND WITH PROGRESS THROUGH
THE PROCESS 7E ATTEMPT TO SHOW HOW THIS PROCESS MAY USEFULLY BE MODELED
THROUGH THE USE OF POPULATION VIABILITY ANALYSIS 06! A TOOL FROM CONSERVATION
BIOLOGY FOR TRACKING POPULATIONS OF IMPERILED SPECIES )NDEED POPULATIONS OF NON
NATIVE SPECIES ARE CHARACTERIZED BY THE SAME LARGE
SCALE PROCESSES IMMIGRATION
EMIGRATION COLONIZATION AND EXTINCTION AS NATIVE POPULATIONS AND SO IT MAKES
SENSE TO UTILIZE THE TERMINOLOGY AND WELL
TESTED TOOLS AVAILABLE TO UNDERSTAND
REGIONAL DYNAMICS OF NATIVE POPULATIONS 7E PROPOSE A MODIlCATION TO THE
2ICHARDSON ET AL A FRAMEWORK FOR THE INVASION PROCESS PROMOTING A FOCUS
ON DEMOGRAPHIC ELEMENTS OF CASUAL NATURALIZED SPREADING AND INVASIVE POPULA
TIONS RATHER THAN CHARACTERIZING THE SPECIES THEMSELVES WITH THIS TERMINOLOGY
4(% ).6!3)/. 02/#%33
).42/$5#4)/. %34!",)3(-%.4 !.$ .!452!,):!4)/.
@)NVASION IS CONSIDERED TO BE THE LEAST LIKELY OUTCOME OF A MULTISTAGE PROCESS THAT
BEGINS WHEN ORGANISMS ARE TRANSPORTED OUTSIDE THEIR NATIVE RANGE -ACK ET AL
-OREOVER IT IS COMMONLY NOTED THAT MANY IF NOT MOST INTRODUCTIONS FAIL
AND THAT EVEN WHEN A SPECIES BECOMES INVASIVE IN A NEW LOCATION THERE MAY HAVE
BEEN NUMEROUS PREVIOUS UNSUCCESSFUL INTRODUCTIONS EG +OWARIK 4HIS IS
WELL ILLUSTRATED IN BIRDS BY THE CLASSIC EXAMPLE OF THE %UROPEAN STARLING 3TURNUS
VULGARIS WHICH SUCCESSFULLY COLONIZED .EW 9ORKS #ENTRAL 0ARK IN AND THEN
MOST OF .ORTH !MERICA BUT ONLY AFTER MANY FAILED INTRODUCTIONS +REBS
!NDERSON ET AL ARGUE THAT MOST OF THE IMPORTANT INFECTIOUS DISEASES OF
PLANTS SIMILARLY DEPEND UPON MULTIPLE INTRODUCTIONS FOR THEIR SUCCESS 7ILLIAMSON
AND &ITTERS @TENS RULE HOLDS THAT JUST ONE IN TEN OF THOSE INDIVIDUALS TRANS
PORTED TO A NEW LOCATION WILL APPEAR IN THE WILD IE BECOME INTRODUCED AND ONLY
ONE IN TEN OF THOSE INTRODUCED WILL BECOME ESTABLISHED AND ONE IN TEN OF THOSE
ESTABLISHED WILL SPREAD AND BECOME INVASIVE
4HOSE INDIVIDUALS THAT DO SURVIVE THE INITIAL TRANSIT ARE UNLIKELY TO PERSIST FOR
LONG IN THE NEW ENVIRONMENT DUE TO A RANGE OF PHYSICAL AND BIOTIC ELEMENTS ASSO
CIATED WITH THE INVADING SPECIES ITSELF THE RECIPIENT COMMUNITY AND THE REST OF
THE ENVIRONMENT &OR EXAMPLE ,ODGE DESCRIBED @ECOLOGICAL RESISTANCE AS
COMPOSED OF DEMOGRAPHIC BIOTIC AND ENVIRONMENTAL FACTORS LIMITING THE SUCCESS
(4 -URPHY ET AL
OF INVADERS /THER OFTEN
CITED EXPLANATIONS FOR THE SUCCESS OR FAILURE OF INTRODUC
TIONS ARE THE @ENEMY RELEASE HYPOTHESIS GENETIC FACTORS AND DEMOGRAPHIC AND
ENVIRONMENTAL STOCHASTICITY SEE BELOW
@$EMOGRAPHIC RESISTANCE INVOLVES PARTICULAR LIFE
HISTORY PARAMETERS OF THE
INVADING SPECIES INmUENCING TYPICALLY PROPAGULE PRESSURE TO THE SYSTEM 6ON
(OLLE ET AL 3UCCESSFUL INTRODUCTION AND ESTABLISHMENT IS OFTEN REPORTED TO
BE STRONGLY RELATED TO PROPAGULE PRESSURE +OLAR AND ,ODGE 4HE EFFECTIVE
NESS OF DIFFERENT BARRIERS LIMITING THE INVASION PROCESS MAY DEPEND ON THE NUMBER
OF PROPAGULES THE GREATER THE NUMBER OF PROPAGULES THE GREATER THE CHANCE OF A
BARRIER BEING OVERCOME AND INVASION PROCEEDING 2OUGET AND 2ICHARDSON
4HUS FACTORS INmUENCING THE NUMBER AND DISPERSAL POTENTIAL OF PROPAGULES ARE
LIKELY TO BE CRITICAL IN REGULATING THE ESTABLISHMENT AND FUTURE SPREAD OF INTRODUCED
SPECIES &OR EXAMPLE 2EJMNEK AND 2ICHARDSON FOUND THAT INVASIVE PINE
SPECIES HAD SMALLER SEED MASS A SHORTER JUVENILE PERIOD AND SHORTER INTERVALS
BETWEEN SEED CROPS COMPARED WITH INTRODUCED BUT NON
INVASIVE PINES
)N BOTH NATURAL AND HUMAN
FACILITATED INTRODUCTIONS VARYING INOCULUM NUM
BERS MAY BIAS OBSERVED PATTERNS OF INVASION AND LEAD TO ERRONEOUS CONCLUSIONS
ABOUT THE IMPORTANCE OF OTHER DEMOGRAPHIC OR BIOTIC PARAMETERS IN INVASION
SUCCESS &OR EXAMPLE A SPECIES COULD HAVE A HIGH RATE OF INTRODUCTION SUCCESS
EITHER BECAUSE IT WAS INHERENTLY A GOOD INVADER OR BECAUSE IT WAS CONSISTENTLY
INTRODUCED TO LOCATIONS THAT WERE EASY TO INVADE OR BECAUSE IT WAS INTRODUCED
MORE FREQUENTLY AT DIFlCULT LOCATIONS 0ROPAGULE PRESSURE HAS BEEN IMPLICATED IN
THE INTRODUCTION SUCCESS OF PLANTS &OXCROFT ET AL 2OUGET AND 2ICHARDSON
BIRDS "LACKBURN AND $UNCAN AND MAMMALS &ORSYTH AND $UNCAN
3TUDIES OF PLANTS HAVE SHOWN THAT MODELS INCORPORATING PROPAGULE PRES
SURE ARE SUPERIOR TO THOSE INVOKING ONLY ENVIRONMENTAL PARAMETERS IN EXPLAINING
DISTRIBUTION AND ABUNDANCE OF NON
NATIVE SPECIES &OXCROFT ET AL 2OUGET
AND 2ICHARDSON )N AN ANALYSIS OF INTRODUCTION EVENTS FOR BIRD
SPECIES ACROSS THE WORLD "LACKBURN AND $UNCAN FOUND THE BROAD PATTERN
OF INTRODUCTION REmECTED PATTERNS OF %UROPEAN SETTLEMENT THE PROXIMITY OF SPE
CIES TO INTRODUCTION LOCATIONS AND THE DESIRABILITY OF SPECIES IN CERTAIN TAXONOMIC
FAMILIES
@"IOTIC RESISTANCE REFERS TO FEATURES SHAPING THE INVADED COMMUNITY AND
AFFECTING ITS @INVASIBILITY WHICH IS OFTEN SAID TO DECREASE WITH INCREASING SPE
CIES RICHNESS OF THE INVADED COMMUNITY %LTON #ASE 4ILMAN
4HE IDEA THAT A COMMUNITYS RESIDENT BIOTIC RICHNESS IS INVERSELY RELATED TO
INVASIBILITY ASSUMES THAT COMPETITION IS A MAJOR FORCE CONTROLLING COMMUNITY
COMPOSITION AND THAT DIVERSE COMMUNITIES SHOULD USE RESOURCES MORE FULLY
LEAVING LITTLE NICHE SPACE FOR POTENTIAL INVADERS "RUNO ET AL BUT SEE
3TOHLGREN ET AL "ESIDES COMPETITION FACTORS CONSIDERED TO BE THE MAJOR
COMPONENTS OF BIOTIC RESISTANCE TO EXOTICS INCLUDE PREDATION HERBIVORY AND DIS
EASEPARASITE PRESSURES ,EVINE ET AL
2ECENTLY ATTENTION HAS BEEN GIVEN TO THE ROLE OF FACILITATIVE INTERACTIONS IN
PROMOTING COMMUNITY COEXISTENCE OF INVASIVE SPECIES WITH NATIVE SPECIES AS WELL
)NVASIVENESS IN EXOTIC PLANTS
AS WITH OTHER INVASIVE SPECIES 2ICHARDSON ET AL B 4HERE IS SOME EVIDENCE
SUGGESTING THAT INVASIONS MAY DEPEND ON OR BE ENHANCED BY THE ESTABLISHMENT
OF MUTUALISMS OR FACILITATIVE INTERACTIONS BETWEEN THE INVASIVE PLANT AND OTHER
ORGANISMS EITHER INTRODUCED OR NATIVE 2ICHARDSON ET AL B 4HERE ARE
MANY EXAMPLES OF INVASIVE WEED SPECIES THAT ARE OBLIGATE OUTCROSSERS AND REQUIRE
A POLLINATOR MUTUALIST TO REPRODUCE EG PURPLE LOOSESTRIFE ,YTHRUM SALICARIA
;-AL ET AL = AND #ENTAUREA DIFFUSA AND # MACULOSA ;(ARROD AND 4AYLOR
= #ARR REPORTED THAT THE INVASIVE mORA OF 6ICTORIA !USTRALIA IS
POLLINATED BY A VARIETY OF BOTH NATIVE AND INTRODUCED INSECTS MAINLY HONEYBEES
!PIS MELLIFERA
)NTRODUCED PLANT SPECIES WHICH ALTER SOIL NUTRIENT AVAILABILITY CAN FACILITATE
THE INVASION OF OTHER NON
INDIGENOUS SPECIES &OR EXAMPLE THE !TLANTIC NITROGEN
lXING SHRUB -YRICA FAYA HAS INVADED NITROGEN
POOR VOLCANIC SOILS IN (AWAII AND
PROBABLY FACILITATES FURTHER PLANT INVASIONS 6ITOUSEK AND 7ALKER AND SEE
3IMBERLOFF AND 6ON (OLLE FOR OTHER EXAMPLES 4HE INTERACTION BETWEEN
PLANTS AND THEIR SOIL COMMUNITY CAN RESULT IN DYNAMIC FEEDBACK %MPIRICAL
EVIDENCE SUGGESTS A NET POSITIVE FEEDBACK BETWEEN PLANTS AND THEIR SOIL COMMUNI
TY IN COMMUNITIES OUTSIDE THEIR NATIVE RANGE 2EINHART ET AL FOUND THAT THE
INVASION OF BLACK CHERRY 0RUNUS SEROTINA INTO NORTH WESTERN %UROPE WAS FACILITAT
ED BY FEATURES OF THE SOIL COMMUNITY )N ITS NATIVE RANGE THE SOIL COMMUNITY THAT
DEVELOPS AROUND BLACK CHERRY INHIBITS THE GROWTH OF NEARBY CONSPECIlCS WHEREAS
THE OPPOSITE OCCURS IN THE NON
NATIVE RANGE #ENTAUREA DIFFUSA HAS MUCH STRONGER
NEGATIVE EFFECTS ON GRASS SPECIES IN ITS NON
NATIVE .ORTH !MERICAN RANGE THAN IN
ITS NATIVE %UROPEAN RANGE DUE TO DIFFERENCES IN THE EFFECTS OF ITS ROOT EXUDATES AND
HOW THESE AFFECT COMPETITION FOR RESOURCES #ALLAWAY AND !SCHEHOUG
%NVIRONMENTAL RESISTANCE MAY INVOLVE ABIOTIC RESOURCE AVAILABILITY SUITABLE
CLIMATIC FACTORS OR LIGHT INTENSITIES FOR EXAMPLE AND MAY PRECLUDE A SPECIES ALTO
GETHER OR MAY INDUCE A SIGNIlCANT TIME LAG BETWEEN ARRIVAL AND ESTABLISHMENT
OF A SPECIES 6ON (OLLE ET AL "LACKBURN AND $UNCAN FOUND USING
A GLOBAL DATASET THAT THE SUCCESS OF INTRODUCED BIRDS DEPENDS ON THE SUITABILITY
OF THE ABIOTIC ENVIRONMENT FOR THE SPECIES AT THE INTRODUCTION SITE &OR EXAMPLE
INTRODUCTION SUCCESS WAS SIGNIlCANTLY GREATER WHEN THE DIFFERENCE BETWEEN A SPE
CIES LATITUDE OF ORIGIN AND THE LATITUDE OF INTRODUCTION WAS SMALL
4HE @ENEMY RELEASE HYPOTHESIS HAS BEEN USED TO EXPLAIN THE SUCCESS OF SOME
SPECIES IN COMMUNITIES OUTSIDE THEIR NATIVE RANGE AND STATES THAT OUTSIDE OF
THEIR NATIVE RANGE PLANT SPECIES SHOULD EXPERIENCE A DECREASE IN REGULATION BY
HERBIVORES AND OTHER NATURAL ENEMIES RESULTING IN AN INCREASE IN ABUNDANCE AND
DISTRIBUTION 4HE SUCCESS OF BIOLOGICAL CONTROL HAS BEEN USED AS SUPPORT FOR THE
ENEMY RELEASE HYPOTHESIS +EANE AND #RAWLEY %MPIRICAL EVIDENCE IN
SUPPORT OF THE HYPOTHESIS IS EQUIVOCAL +EANE AND #RAWLEY -ARON AND
6IL HOWEVER IN AN ANALYSIS OF PLANT SPECIES NATURALIZED IN THE 53
-ITCHELL AND 0OWER FOUND SPECIES IN THEIR NON
NATIVE RANGES HAD ON
AVERAGE FEWER FUNGI AND FEWER VIRUS SPECIES THAN IN THE RESPECTIVE
NATIVE RANGES
(4 -URPHY ET AL
$EMOGRAPHIC AND ENVIRONMENTAL STOCHASTICITIES PROBABLY PROMOTE LOCAL EXTINC
TION IN MANY SMALL POPULATIONS OF SPECIES BEGINNING TO ESTABLISH IN NEW LOCATIONS
3AX AND "ROWN PROPOSED TWO FURTHER PHENOMENA THAT MAY ACCOUNT FOR
THE LARGE FRACTION OF FAILED INTRODUCTIONS 4HEY NOTED THAT SPECIES WHICH ARE ABUN
DANT AND WIDESPREAD TEND TO BE RARE OR ABSENT FROM MOST LOCATIONS WITHIN THEIR
GEOGRAPHIC RANGE DUE TO SPATIAL ENVIRONMENTAL VARIATION 4HIS SUGGESTS THAT MOST
LOCAL INTRODUCTIONS SHOULD FAIL BECAUSE PROPAGULES WOULD TEND BY CHANCE TO BE
INTRODUCED TO MARGINAL OR UNSUITABLE SITES 3ECONDLY MANY ESTABLISHED POPULA
TIONS ARE SUSTAINED BY THE POSITIVE FEEDBACK EFFECTS OF DISPERSAL EVENTS ON LOCAL
POPULATION DYNAMICS 3INGLE SMALL AND NEWLY ESTABLISHED POPULATIONS ALL LACK
OPPORTUNITIES FOR RESCUE BY OTHER POPULATIONS AND THUS ARE MORE SUSCEPTIBLE TO
EXTINCTION 3AX AND "ROWN
4HERE ARE ALSO GENETIC OBSTACLES TO THE PERSISTENCE OF POPULATIONS OF INTRODUCED
ORGANISMS 'ENETIC DIVERSITY AND PHENOTYPIC PLASTICITY THAT CAN BRING THE LINEAGE
INTO ACCORD WITH ITS NEW PHYSICAL AND BIOLOGICAL ENVIRONMENT MAY BE DIFlCULT TO
ACHIEVE IN FAR
mUNG POPULATIONS ,EVIN BECAUSE THEY ARE SUBJECT TO GENETIC
DRIFT AND FOUNDER EFFECTS %CKERT ET AL 'ENETIC VARIATION IN FOUNDER POPULA
TIONS IS LIKELY TO BE LACKING IF THE NUMBER OF INDIVIDUALS INTRODUCED IS NOT LARGE OR
POPULATION GROWTH RATE IS NOT SUBSTANTIAL .OVAK AND -ACK +OLBE ET AL
RECENTLY DEMONSTRATED THAT INVASION SUCCESS WAS ENHANCED WHEN MULTIPLE
INTRODUCTIONS OF THE #ARIBBEAN BROWN ANOLE !NOLIS SAGREI TO &LORIDA CAME FROM
DIFFERENT SOURCE POPULATIONS IN THE LIZARDS GEOGRAPHIC RANGE 0OPULATIONS IN THE
ADVENTIVE RANGE CONTAINED MORE GENETIC VARIATION THAN NATIVE POPULATIONS
).6!3)6%.%33
!MONG NATURALIZED SPECIES PERSISTING AT NEW LOCATIONS DESPITE THE SERIES OF SEVERE
CONSTRAINTS DESCRIBED ABOVE A FEW MAY GO ON TO BECOME INVADERS 7ILLIAMSON AND
&ITTER -ACK ET AL $ARWIN P WAS THE lRST TO PROPOSE A THEORY
FOR INVASION SUCCESS IN PLANT SPECIES 5SING OBSERVATIONS BY !LPHONSE DE #ANDOLLE
$ARWIN NOTED THAT hmORAS GAIN BY NATURALIZATION PROPORTIONALLY WITH THE NUMBER
OF NATIVE GENERA AND SPECIES FAR MORE IN NEW GENERA THAN IN NEW SPECIESv $ARWIN
GIVES THE EXAMPLE THAT h)N THE LAST EDITION OF $R !SA 'RAYS @-ANUAL OF THE mORA OF
THE 5NITED 3TATES NATURALIZED SPECIES ARE ENUMERATED AND THESE BELONG TO
GENERA OUT OF THE GENERA NO LESS THAN GENERA ARE NOT THERE INDIGENOUSv
$ARWIN REASONED THAT BECAUSE DIFFERENCES IN STRUCTURE AND FUNCTIONING ARE GREATER
BETWEEN GENERA THAN BETWEEN SPECIES COMPETITION IS LIKELY TO BE GREATER BETWEEN
CONGENERS 2ESULTS BOTH SUPPORT EG -ACK 2EJMNEK AND ARGUE
AGAINST $AEHLER B $ARWINS HYPOTHESIS $AEHLERS CONTRADICTORY RESULT FOR THE
(AWAIIAN mORA INDEED SUPPORTS $ARWINS ORIGINAL LINE OF THINKING h)T MIGHT
HAVE BEEN EXPECTED THAT THE PLANTS WHICH WOULD SUCCEED IN BECOMING NATURALIZED
IN ANY LAND WOULD GENERALLY HAVE BEEN CLOSELY ALLIED TO INDIGENES FOR THESE ARE
COMMONLY LOOKED AT AS SPECIALLY ADAPTED FOR THEIR OWN COUNTRYv
)NVASIVENESS IN EXOTIC PLANTS
.UMEROUS OFTEN DISPARATE LIFE HISTORY FACTORS HAVE BEEN SUGGESTED TO EXPLAIN
THE INVASION SUCCESS OF PARTICULAR PLANT SPECIES "AKER DISCUSSED
SEVERAL TRAITS ASSOCIATED WITH INVASIVE PLANT SPECIES AND PROPOSED THAT SPECIES
HAVING MORE OF THESE TRAITS WOULD BE MORE @WEEDY THAN SPECIES WITH FEWER OF
THESE TRAITS 4HUS THE @IDEAL WEED WOULD HAVE THE ABILITY TO REPRODUCE SEXUALLY
AND GROW CLONALLY SHOW RAPID GROWTH PHENOTYPIC PLASTICITY AND HIGH TOLERANCE
OF ENVIRONMENTAL HETEROGENEITY #ADOTTE AND ,OVETT $OUST EXPLICITLY
TESTED "AKERS NOTION OF A WEED IN THE EXOTIC mORA OF /NTARIO AND FOUND GENERAL
CONCORDANCE WITH THE TRAITS "AKER DESCRIBED /THER TRAITS HAVE ALSO BEEN SHOWN
TO CORRELATE WITH INVASION SUCCESS 2EJMNEK FOUND THAT INVASIVENESS OF
HERB SPECIES WAS BEST PREDICTED BY NATIVE LATITUDINAL RANGE 3IMILARLY #ADOTTE
ET AL FOUND THAT SUCCESSFUL INVADERS IN /NTARIO WERE SIGNIlCANTLY OVER
REPRESENTED BY SPECIES ORIGINATING IN %UROPE AND %URASIA ,IFE HISTORY ATTRIBUTES
PARTICULARLY ASSOCIATED WITH INVASION SUCCESS IN THE /NTARIO mORA WERE THE PRES
ENCE OF A CLONAL ORGAN AND A LONGER mOWERING SEASON
(/7 4/ #,!33)&9 ).6!3)6%3
!N IDEAL CLASSIlCATION SCHEME SHOULD TO HAVE UTILITY BE CAPABLE OF GENERAL
APPLICATION 2ICHARDSON ET AL A OFFERED OPERATIONAL DElNITIONS FOR TERMS
SUCH AS @ALIEN @CASUAL ALIEN @NATURALIZED @INVASIVE AND @TRANSFORMER WITH
RESPECT TO PLANT SPECIES )N THEIR SCHEME INVASION OF A NEW REGION BY AN INTRO
DUCED SPECIES WAS ENVISIONED AS SURMOUNTING A SERIES OF @BARRIERS SEE &IG
4HUS @CASUAL ALIEN SPECIES HAVE OVERCOME GEOGRAPHIC AND LOCAL ENVIRONMENTAL
BARRIERS ENABLING THEM TO SURVIVE IN A NEW LOCATION BUT HAVE NOT OVERCOME
BARRIERS TO SUCCESSFUL REPRODUCTION AND POPULATION GROWTH OR EXPANSION IN THE
NEW LOCALE .ATURALIZATION IN 2ICHARDSON ET ALS SCHEME BEGINS WHEN VARIOUS
BARRIERS TO REGULAR REPRODUCTION ARE OVERCOME AND POPULATION GROWTH AND SUR
VIVAL PERSISTENCE IS MAINTAINED OVER THE LONG TERM FUNCTIONALLY LONG
TERM HAS
BEEN REGARDED AS C YEARS ;EG 4UTIN ET AL = OR YEARS ;0YEK ET AL
= @)NVASION IS CONSIDERED TO HAVE OCCURRED WHEN THE SPECIES HAS OVERCOME
THE BARRIERS TO DISPERSAL WITHIN THE NEW REGION AND NEW POPULATIONS ARE BEING
FOUNDED AWAY FROM THE IMMEDIATE AREA OF INTRODUCTION
$AVIS AND 4HOMPSON PROPOSED A CLASSIlCATION SCHEME MODELED AFTER
2ABINOWITZS TRIPARTITE SCHEME FOR RARITY TO DElNE @INVADERS AS DISTINCT
FROM @SUCCESSIONAL COLONIZERS AND @NONINVASIVE COLONIZERS $AVIS AND 4HOMPSON
USED A TAXONOMY OF INVASION THAT RAISES THREE DICHOTOMOUS QUESTIONS IS THE
SPECIES A LONG
DISTANCE OR SHORT
DISTANCE DISPERSER IS THE SPECIES COMMON
OR NOVEL TO THE REGION AND DOES IT HAVE A SMALL OR LARGE @IMPACT ON THE NEW
ENVIRONMENT 4HIS SCHEME GENERATES EIGHT POTENTIAL CONDITIONS 4HE MAJOR OPERA
TIONAL DIFFERENCE BETWEEN THE SYSTEM OF $AVIS AND 4HOMPSON AND OF 2ICHARDSON
ET AL A LIES IN THE EMPHASIS IN $AVIS AND 4HOMPSON UPON @IMPACT IN
THE NEW ENVIRONMENT WHEREAS 2ICHARDSON ET AL RELY ON MEASURES OF NATURAL
(4 -URPHY ET AL
&IG ! SCHEMATIC REPRESENTATION OF MAJOR BARRIERS !n' LIMITING THE SPREAD OF
INTRODUCED PLANTS -ODIlED FROM 2ICHARDSON ET AL ;A=
REPRODUCTION AND RATES OF SPREAD IN THE NEW ENVIRONMENT AS MAJOR CRITERIA DETER
MINING INVASIVENESS $AEHLER A HAS ARGUED AGAINST THE UTILITY OF $AVIS
AND 4HOMPSONS SCHEME INSOFAR AS IT HINGED ON A SUBJECTIVE CRITERION @IMPACT
WHEREAS THE @SPREAD CRITERION OF 2ICHARDSON ET AL CAN BE MEASURED RELATIVELY
MORE OBJECTIVELY $AEHLER NOTED THAT BOTH SCHEMES PROBABLY EMBRACE SIMILAR
SETS OF SPECIES SINCE THE CONCORDANCE BETWEEN INVADERS DElNED BY THE SPREAD
CRITERION AND THOSE THAT HAVE A GREAT IMPACT IS LIKELY CLOSE TO A HUNDRED PERCENT
-%4(/$3
/VER THE PAST THREE DECADES THE #ANADIAN *OURNAL OF 0LANT 3CIENCE HAS PUBLISHED
ACCOUNTS OF THE BIOLOGY OF PLANTS THAT ARE WEEDY IN #ANADA %ACH ACCOUNT FOL
LOWS A PRESCRIBED LAYOUT AND PROVIDES A DETAILED DESCRIPTION OF A SINGLE SPECIES
OF WEED OR CLOSELY RELATED GROUP OF SPECIES THE HISTORY OF ITS INTRODUCTION TO
.ORTH !MERICA AND #ANADA DETAILS OF ITS ECONOMIC IMPORTANCE A #ANADIAN DIS
TRIBUTION MAP AND DETAILS OF ITS DISTRIBUTION ELSEWHERE IN THE WORLD !DDITIONALLY
)NVASIVENESS IN EXOTIC PLANTS
INFORMATION ON HABITATS OCCUPIED MEANS OF SEED AND VEGETATIVE REPRODUCTION
AND DISPERSAL AND DETAILS OF GROWTH AND DEVELOPMENT ARE PROVIDED AS WELL AS
RESPONSE TO WEED CONTROL MEASURES !RTICLES IN THE SERIES BRING TOGETHER PUBLISHED
AND UNPUBLISHED INFORMATION ON THE BIOLOGY ECOLOGY AND CONTROL OF THESE WEEDS
4HUS THE INFORMATION IS RELATIVELY COMPREHENSIVE AND CONSISTENTLY PRESENTED
)NDEPENDENTLY AN EXOTIC SPECIES LIST FOR /NTARIO WAS GENERATED FROM THE /NTARIO
-INISTRY OF .ATURAL 2ESOURCES .ATURAL (ERITAGE )NFORMATION #ENTRES DATA
BASE OF VASCULAR PLANTS .()# HTTPWWWMNRGOVONCA-.2NHICNHICCFM
&OR THE PRESENT ANALYSIS WE USED THE SPECIES OCCURRING ON THE .()# LIST OF /NTARIOS
EXOTIC mORA FOR WHICH THERE WAS A "IOLOGY OF #ANADIAN 7EEDS ARTICLE . SEE
!PPENDIX FOR LIST AND REFERENCES
7E ATTEMPTED lRST TO EXTRACT SPECIlC DATA REQUIRED TO MEET THE CRITERIA FOR
EACH CATEGORY IN THE CLASSIlCATIONS OF 2ICHARDSON ET AL A AND $AVIS AND
4HOMPSON 4HIS WE TERM THE @IDEAL CRITERIA 4HAT IS IF AVAILABLE THE IDEAL
CRITERIA WOULD BEST SATISFY THE DElNITION OF EACH CATEGORY IN THE CLASSIlCATION AND
WOULD DIVIDE SPECIES INTO THE VARIOUS CATEGORIES DESCRIBED BY THE TWO SCHEMES
4HEN USING THE INFORMATION GIVEN IN THE "IOLOGY OF #ANADIAN 7EEDS REVIEWS AND
THE REGIONAL ABUNDANCE RANKING VALUES DETERMINED BY THE .()# WE DEVELOPED
@REALIZED CRITERIA FOR EACH CATEGORY IN THE TWO CLASSIlCATIONS 2EALIZED CRITERIA ARE
THE BEST lT TO THE IDEAL CRITERIA USING INFORMATION THAT IS ACTUALLY AVAILABLE 4ABLE
SHOWS IDEAL AND REALIZED CRITERIA FOR EACH CATEGORY IN THE TWO SCHEMES )N ORDER
TO ADDRESS EACH CRITERION DATA WERE TABULATED FOR EACH OF THE WEED SPECIES SEE
4ABLE
#ERTAIN LIMITATIONS OF BOTH SCHEMES BECAME EVIDENT EARLY $AVIS AND 4HOMPSON
CALL FOR AN ASSESSMENT OF @DISPERSAL DISTANCE )N MANY CASES AN INITIAL LONG
DISTANCE DISPERSAL EVENT BY ONE AGENT MAY LEAD TO AN INTRODUCTION IN A NEW REGION
WHILE ANOTHER KIND OF AGENT IS RESPONSIBLE FOR SUBSEQUENT SHORT
MEDIUM
OR
LONG
DISTANCE DISPERSAL IN THE NEW LOCATION IE A FORM OF DIPLOCHORY SENSU
6ANDER 7ALL AND ,ONGLAND ;= )T SEEMS UNLIKELY THAT AN INITIAL LONG
DISTANCE
DISPERSAL EVENT FROM A WIDELY DISTANT ENVIRONMENT WOULD HAVE MUCH EFFECT IN
THE NEW ENVIRONMENT OR RELEVANCE TO SUBSEQUENT SPREAD IF THE SPECIES DID NOT
THEN SUCCESSFULLY DISPERSE FROM ITS INITIAL POINT OF ESTABLISHMENT 4HEREFORE IN OUR
ANALYSIS THE DISPERSAL CRITERION IS BASED ON HOW THE SPECIES IS MOVED FOLLOWING
ITS INITIAL INTRODUCTION TO THE ADVENTIVE REGION 4HIS IS NOT ALWAYS STRAIGHTFOR
WARD SINCE EVEN APPROXIMATE KNOWLEDGE ABOUT DISPERSAL DISTANCE PARTICULARLY
LONG
DISTANCE DISPERSAL IS TYPICALLY SCARCE FOR EVEN THE BEST
STUDIED SPECIES 7E
DETERMINED THE PRINCIPAL MODES OF DISPERSAL AND ASSIGNED THEM TO EITHER A SHORT
OR LONG
DISTANCE CATEGORY BEARING IN MIND THAT THE OVERALL SCALE OF THE ANALYSIS
WAS THE ENTIRE PROVINCE OF /NTARIO TOTAL AREA KM ;3TATISTICS #ANADA
WWWSTATCANCA= 4HUS ANY SPECIES THAT DID NOT HAVE A KNOWN HUMAN
FACILITATED
MODE OF DISPERSAL WAS CONSIDERED A SHORT
DISTANCE DISPERSER )N ADDITION SOME
CATEGORIES OF HUMAN
FACILITATED DISPERSAL THAT GENERALLY RESULT IN LIMITED DISPERSAL
SUCH AS WITHIN A CROP lELD OR FARM PROPERTY WERE ALSO CONSIDERED SHORT
DISTANCE
EG THOSE ASSOCIATED WITH MOWING CULTIVATION FARM MACHINERY DOMESTIC ANIMALS
4ABLE #RITERIA USED IN CLASSIFYING SPECIES ACCORDING TO THE SCHEMES OF $AVIS AND 4HOMPSON AND 2ICHARDSON ET AL A @)DEAL
CRITERIA ARE THOSE THAT IF KNOWN WOULD BEST SATISFY THE CRITERIA PROPOSED BY THE AUTHORS OF THE TWO SCHEMES @2EALIZED CRITERIA ARE THE BEST lT
TO THE IDEAL CRITERIA USING INFORMATION THAT IS ACTUALLY AVAILABLE
0ARAMETER )DEAL CRITERIA 2EALIZED #RITERIA
$AVIS AND 4HOMPSON
)NVADERS 3HORT OR LONG DISTANCE DISPERSERS $ISPERSAL HAS OCCURRED GRADUALLY TO NEARBY OR $ISPERSAL IS BY VEGETATIVE OR SEED
ADJACENT ENVIRONMENTS WITHOUT SIGNIlCANT DISPERSAL ANDOR IS OFTEN
HUMAN ASSISTANCE /2 $ISPERSAL HAS OCCURRED FACILITATED BY HUMANS
OVER LARGER DISTANCES BETWEEN WIDELY DISTANT
ENVIRONMENTS AND MAY BE FACILITATED BY
HUMANS
.OVEL IN THE NEW ENVIRONMENT 4HE SPECIES HAS EXPANDED ITS RANGE TO OCCUR /NTARIO IS NOT WITHIN THE NATIVE
IN THE REGION GEOGRAPHIC RANGE OF THE SPECIES
'REAT IMPACT IN THE NEW #OMMUNITY IMPACTS THE SPECIES IS NOTED TO 4HE SPECIES IS NOTED TO REGULARLY
ENVIRONMENT HAVE A SUBSTANTIAL IMPACT ON NATIVE OCCUR IN NATURAL OR SEMI NATURAL
(4 -URPHY ET AL
COMMUNITIES !.$/2 %COSYSTEM IMPACTS NATURAL DISTURBED COMMUNITIES
THE SPECIES IS NOTED TO HAVE A SUBSTANTIAL !.$/2 4HE SPECIES IS NOTED TO
IMPACT ON ECOSYSTEM PROCESSES !.$/2 SUBSTANTIALLY IMPACT ECOSYSTEM
%CONOMIC IMPACTS THE SPECIES IS NOTED TO PROCESSES !.$/2 4HE SPECIES IS
HAVE A SUBSTANTIAL ECONOMIC IMPACT NOTED TO HAVE SUBSTANTIAL
ECONOMIC IMPACT
3UCCESSIONAL 3HORT OR LONG DISTANCE DISPERSERS $ISPERSAL HAS OCCURRED GRADUALLY TO NEARBY OR $ISPERSAL IS BY VEGETATIVE OR SEED
COLONIZERS ADJACENT ENVIRONMENTS WITHOUT SIGNIlCANT DISPERSAL ANDOR IS OFTEN FACILITATED
HUMAN ASSISTANCE /2 $ISPERSAL HAS OCCURRED BY HUMANS
OVER LARGER DISTANCES BETWEEN WIDELY DISTANT
ENVIRONMENTS AND MAY BE FACILITATED BY HUMANS
3UCCESSIONAL #OMMON IN THE NEW 4HE SPECIES IS NATIVE TO THE REGION ITS /NTARIO IS WITHIN THE NATIVE
COLONIZERS ENVIRONMENT OCCURRENCE IN THE REGION INVOLVES NO RANGE GEOGRAPHIC RANGE OF THE SPECIES
EXPANSION
3MALL OR GREAT IMPACT IN #OMMUNITY IMPACTS THE SPECIES IS NOT NOTED 4HE SPECIES DOES NOT GENERALLY
THE NEW ENVIRONMENT TO HAVE A SUBSTANTIAL IMPACT ON NATIVE OCCUR IN NATURAL OR SEMI NATURAL
COMMUNITIES !.$ %COSYSTEM IMPACTS COMMUNITIES )T PRIMARILY INVADES
THE SPECIES IS NOT NOTED TO HAVE A SUBSTANTIAL DISTURBED @UNNATURAL AREAS !.$
IMPACT ON ECOSYSTEM PROCESSES THE SPECIES 4HE SPECIES IS NOT NOTED TO
DOES NOT PLAY A KEYSTONE ROLE IN THE NEW SUBSTANTIALLY IMPACT ECOSYSTEM
COMMUNITY OR ECOSYSTEM !.$ %CONOMIC IMPACTS PROCESS !.$ ECONOMIC IMPACTS
THE SPECIES IS NOT NOTED TO HAVE A SUBSTANTIAL ASSOCIATED WITH THE SPECIES ARE
ECONOMIC IMPACT /2 #OMMUNITY IMPACTS LIMITED IN NATURE OR SCALE /2
THE SPECIES IS NOTED TO HAVE A SUBSTANTIAL IMPACT 4HE SPECIES IS NOTED REGULARLY TO
ON NATIVE COMMUNITIES !.$/2 %COSYSTEM OCCUR IN NATURAL OR SEMI NATURAL
IMPACTS THE SPECIES IS NOTED TO HAVE A NATURAL DISTURBED COMMUNITIES
SUBSTANTIAL IMPACT ON ECOSYSTEM PROCESSES !.$/2 4HE SPECIES IS NOTED TO
!.$/2 %CONOMIC IMPACTS THE SPECIES IS SUBSTANTIALLY IMPACT ECOSYSTEM
)NVASIVENESS IN EXOTIC PLANTS
NOTED TO HAVE A SUBSTANTIAL ECONOMIC IMPACT PROCESSES !.$/2 4HE SPECIES IS
NOTED TO HAVE SUBSTANTIAL
ECONOMIC IMPACT
4ABLE #ONTINUED
0ARAMETER )DEAL CRITERIA 2EALIZED #RITERIA
$AVIS AND 4HOMPSON
.OVEL 3HORT OR LONG $ISPERSAL HAS OCCURRED GRADUALLY TO NEARBY $ISPERSAL IS BY VEGETATIVE MEANS OR SEED ANDOR
NON
INVASIVE DISTANCE OR ADJACENT ENVIRONMENTS WITHOUT SIGNIlCANT IS OFTEN FACILITATED BY HUMANS
COLONIZERS DISPERSERS HUMAN ASSISTANCE /2 $ISPERSAL HAS OCCURRED
OVER LARGER DISTANCES BETWEEN WIDELY DISTANT
ENVIRONMENTS AND MAY BE FACILITATED
BY HUMANS
.OVEL IN THE NEW 4HE SPECIES HAS EXPANDED ITS RANGE TO OCCUR /NTARIO IS NOT WITHIN THE NATIVE GEOGRAPHIC
ENVIRONMENT IN THE REGION RANGE OF THE SPECIES
3MALL IMPACT IN #OMMUNITY IMPACTS THE SPECIES IS NOT NOTED 4HE SPECIES DOES NOT GENERALLY OCCUR IN NATURAL
THE NEW TO HAVE A SUBSTANTIAL IMPACT ON NATIVE OR SEMI NATURAL COMMUNITIES )T PRIMARILY
ENVIRONMENT COMMUNITIES !.$ %COSYSTEM IMPACTS INVADES DISTURBED @UNNATURAL AREAS !.$ 4HE
THE SPECIES IS NOT NOTED TO HAVE A SUBSTANTIAL SPECIES IS NOT NOTED TO SUBSTANTIALLY IMPACT
IMPACT ON ECOSYSTEM PROCESSES THE SPECIES DOES ECOSYSTEM PROCESS !.$ ECONOMIC IMPACTS
(4 -URPHY ET AL
NOT PLAY A KEYSTONE ROLE IN THE NEW COMMUNITY ASSOCIATED WITH THE SPECIES ARE LIMITED IN NATURE
OR ECOSYSTEM !.$ %CONOMIC IMPACTS OR SCALE
THE SPECIES IS NOT NOTED TO HAVE A SUBSTANTIAL
ECONOMIC IMPACT
2ICHARDSON ET AL A
!LIEN PLANTS 4HE SPECIES OCCURS IN AN AREA KM FROM !LL 3%S NOT NATIVE TO /NTARIO
THE NEAREST @NATURAL POPULATION DUE TO
INTRODUCTION AS A RESULT OF HUMAN ACTIVITY
#ASUAL ALIEN PLANTS !N ALIEN PLANT 3% 2ANKING IN /NTARIO
2ELIES ON REPEATED INTRODUCTIONS FOR %XTREMELY RARE TO VERY RARE HIGH EXTINCTION
ITS PERSISTENCE PROBABILITY 3% 3%
4HESE SPECIES DO NOT SPREAD NATURALLY PAST (UMAN AIDED DISPERSAL COMMON !.$
THEIR POINT OF INTRODUCTION OCCURRENCE ONLY IN HUMAN DISTURBED AREAS
.ATURALIZED PLANTS !N ALIEN PLANT 3% 2ANKING IN /NTARIO
2EPRODUCES CONSISTENTLY AND SUSTAINS 5NCOMMON TO COMMON EXTINCTION PROBABILITY
POPULATIONS OVER MANY LIFECYLES MEDIUM TO LOW 3% 3%
)NVASIVENESS IN EXOTIC PLANTS
(AS NOT SPREAD INTO NEW ENVIRONMENTS BEYOND /CCURRENCE ONLY IN HUMAN DISTURBED AREAS
THE POINT OF INTRODUCTION HAVE NOT DISPERSED TO NATURAL OR SEMI NATURAL
COMMUNITIES
4ABLE #ONTINUED
0ARAMETER )DEAL CRITERIA 2EALIZED #RITERIA
2ICHARDSON ET AL A
)NVASIVE PLANTS !N ALIEN PLANT 3% 2ANKING IN /NTARIO
0RODUCES REPRODUCTIVE OFFSPRING IN LARGE NUMBERS #OMMON AND WIDESPREAD LOW EXTINCTION
AT CONSIDERABLE DISTANCE FROM PARENT PLANT PROBABILITY 3% 3%
FOR SPECIES PRIMARILY DISPERSED BY SEED M
IN YEARS FOR SPECIES SPREADING PRIMARILY
VEGETATIVELY MYEARS OR ANY SPECIES
PRIMARILY SPREAD BY HUMANS
(AS SPREAD TO AREAS DISTANT TO THE POINT /CCUR IN HUMAN DISTURBED AREAS AND HAVE
OF INTRODUCTION SPREAD TO OCCUR IN DISTURBED NATURAL AND
SEMI NATURAL COMMUNITIES
4RANSFORMERS !N INVASIVE PLANT
(4 -URPHY ET AL
#HANGES THE CHARACTER CONDITION FORM OR
NATURE OF ECOSYSTEMS OVER A SUBSTANTIAL AREA
/PERATIONALLY OUR DElNITION OF NATURALIZED THUS LIMITS SPECIES TO OCCURRENCE IN MAINLY HUMAN MODIlED COMMUNITIES THIS IS NOT NECESSARILY
THE INTENT OF 2ICHARDSON ET ALS DElNITION SEE TEXT ON THIS
)NVASIVENESS IN EXOTIC PLANTS
,ONG
DISTANCE DISPERSAL VECTORS INCLUDED WEED
CONTAMINATED CROP SEED SOIL
OR OTHER AGRICULTURAL PRODUCTS BOATS CARS AND OTHER VEHICLES WATERFOWL FISH AND
IRRIGATION WATER
2ATES OF SPREAD AS REQUIRED BY 2ICHARDSON ET ALS SCHEME PROVE TO BE ONLY
RARELY AVAILABLE ONLY SEVEN OF THE ARTICLES WE SURVEYED REPORTED ANY RATE OF
SPREAD &IVE OF THESE WERE FOR SPECIES THAT SPREAD PRIMARILY BY VEGETATIVE MEANS
AND NONE HAD BEEN MEASURED OR ESTIMATED IN THE ADVENTIVE /NTARIO ENVIRONMENT
7HERE DISPERSAL OF PROPAGULES IS PROMOTED BY ANTHROPOGENIC ACTIVITIES OR OTHER
NON
STANDARD MEANS RATES OF SPREAD CANNOT REASONABLY BE ESTIMATED BY SEED OR
VEGETATIVE PROPAGULE CHARACTERISTICS (IGGINS ET AL !LTHOUGH IN THEORY
SPREAD COULD BE MEASURED RELATIVELY OBJECTIVELY USING POPULATION GROWTH AND
DISTANCE FROM THE SOURCE UNLESS SUBSTANTIAL HISTORICAL INFORMATION IS AVAILABLE
REAL RATES OF SPREAD ARE QUITE DIFlCULT TO DETERMINE &URTHERMORE FOR MANY OF THE
SPECIES IN OUR DATASET WHICH ARE PRIMARILY AGRICULTURAL WEEDS LONG
DISTANCE
DISPERSAL MAY OCCUR RELATIVELY REGULARLY THROUGH TRANSPORT IN CONTAMINATED
SEED STOCKS OR INFESTED HAY 4HESE METHODS HAVE BEEN CONSIDERED A MAJOR CON
TRIBUTOR TO SPREAD FOR MANY SPECIES EG 3ILENE ALBA -C.EILL -ELILOTUS SPP
4URKINGTON ET AL 3ORGHUM HALEPENSE 7ARWICK AND "LACK
4HE .()# DATABASE INCLUDES ESTIMATES OF ABUNDANCE OF EXOTIC PLANTS USING A
PROVINCIAL RANKING SYSTEM 3%
RANK ;3UB
NATIONAL EXOTIC= 7E USED THE PROVIN
CIAL 3% RANKING AS A SURROGATE FOR THE AMOUNT OF @SPREAD THAT HAD BEEN ACHIEVED
4HE 3% RANKING IS ESSENTIALLY A DISTRIBUTION AND ABUNDANCE INDICATOR AND SO
CARRIES IMPLICATIONS OF EXTINCTION RISK 4HUS 3%
AND 3%
RANKED SPECIES ARE CON
SIDERED VULNERABLE TO EXTINCTION 3% SPECIES HAVE A MEDIUM RISK OF EXTINCTION AND
3% AND 3% HAVE GENERALLY A LOW RISK OF EXTINCTION SEE -ASTER 4HUS
s 3% %XTREMELY RARE IN /NTARIO USUALLY OR FEWER OCCURRENCES IN THE PROV
INCE OR VERY FEW REMAINING INDIVIDUALS OFTEN ESPECIALLY VULNERABLE TO
EXTIRPATION
s 3% 6ERY RARE IN /NTARIO USUALLY BETWEEN AND OCCURRENCES IN THE PROV
INCE OR WITH MANY INDIVIDUALS IN FEWER OCCURRENCES OFTEN SUSCEPTIBLE TO
EXTIRPATION
s 3% 2ARE TO UNCOMMON IN /NTARIO USUALLY BETWEEN AND OCCURRENCES
IN THE PROVINCE MAY HAVE FEWER OCCURRENCES BUT WITH A LARGE NUMBER
OF INDIVIDUALS IN SOME POPULATIONS MAY BE SUSCEPTIBLE TO LARGE
SCALE
DISTURBANCES -OST SPECIES WITH AN 3% RANK ARE ASSIGNED TO THE WATCH
LIST UNLESS THEY HAVE A RELATIVELY HIGH GLOBAL RANK
s 3% #OMMON AND APPARENTLY SECURE IN /NTARIO USUALLY WITH MORE THAN
OCCURRENCES IN THE PROVINCE
s 3% 6ERY COMMON AND DEMONSTRABLY SECURE IN /NTARIO
#OMMUNITY ECOSYSTEM AND ECONOMIC @IMPACTS AS PER $AVIS AND 4HOMPSONS
SCHEME WERE QUANTIlED AS FOLLOWS AND SEE 4ABLE &OR COMMUNITY IMPACT
SPECIES OCCURRENCE IN ONLY COMMUNITY TYPES ANDOR BOTH GENERALLY HIGHLY
HUMAN MODIlED COMMUNITIES WAS SCORED AS POINT OCCURRENCE IN ANY OTHER
4ABLE $ATA EXTRACTED FROM THE "IOLOGY OF #ANADIAN 7EEDS SERIES AND .()# DATABASE TO ADDRESS CRITERIA OUTLINED IN 4ABLE
!TTRIBUTE $ESCRIPTION 6ARIABLE
!BUNDANCE 2ANK 3% 2ANKING 3% %XTREMELY RARE IN /NTARIO USUALLY OR FEWER OCCURRENCES IN THE PROVINCE
OR VERY FEW REMAINING INDIVIDUALS OFTEN ESPECIALLY VULNERABLE TO
EXTIRPATION
3% 6ERY RARE IN /NTARIO USUALLY BETWEEN AND OCCURRENCES IN THE
PROVINCE OR WITH MANY INDIVIDUALS IN FEWER OCCURRENCES OFTEN SUSCEPTIBLE
TO EXTIRPATION
3% 2ARE TO UNCOMMON IN /NTARIO USUALLY BETWEEN AND OCCURRENCES
IN THE PROVINCE MAY HAVE FEWER OCCURRENCES BUT WITH A LARGE NUMBER OF
INDIVIDUALS IN SOME POPULATIONS MAY BE SUSCEPTIBLE TO LARGE
SCALE
DISTURBANCES
3% #OMMON AND APPARENTLY SECURE IN /NTARIO USUALLY WITH MORE THAN
OCCURRENCES IN THE PROVINCE
3% 6ERY COMMON AND DEMONSTRABLY SECURE IN /NTARIO
$ATE OF lRST &ROM HERBARIUM SPECIMEN OR $ATE
DOCUMENTED IF NO DATE GIVEN DATE IS NOTED
OCCURRENCE AS IF THE SPECIES IS LISTED
(4 -URPHY ET AL
IN /NTARIO IN 'LEASON
.ATIVITY TO /NTARIO )S /NTARIO WITHIN THE NATIVE YES NO
RANGE OF THE SPECIES
0RIMARY MEANS 6EGETATIVE OR SEED VEGETATIVE SEED
OF RECRUITMENT
4ABLE #ONTINUED
!TTRIBUTE $ESCRIPTION 6ARIABLE
0RIMARY DISPERSAL 0RIMARY MODE OF NATURAL WIND WATER BIRD ANIMAL GRAVITY OR BALLISTIC
MODE DISPERSAL USUALLY BASED ON
ADAPTATIONS OF THE SEED OR FRUIT
3ECONDARY /THER DOCUMENTED USUALLY CONTAMINATED CROP SEEDS OR INFESTED HAY FARM MACHINERY OR VEHICLES
DISPERSAL MODE HUMAN FACILITATED MEANS IN SOIL OR MANURE LIVESTOCK OR DOMESTIC ANIMALS BOATS IRRIGATION
OF DISPERSAL WATER HUMAN CLOTHING OR FOOTWEAR
#OMMUNITIES IN WHICH $OCUMENTED OCCURRENCE IN ROADSIDES GARDENS WASTELANDS OTHER DISTURBED URBAN AREAS GRAZED
SPECIES OCCURS VARIOUS COMMUNITY TYPES PASTURE CULTIVATED lELDS CROPS DISTURBED NATURAL COMMUNITIES OPEN
WOODLAND FOREST MARGINS OR CLEARINGS STREAM BANKS SHORELINES
AQUATIC LAKE STREAM POND RIVER ETC
%CONOMIC IMPACTS $OCUMENTED ECONOMIC IMPACTS COMPETES WITH CROP OR PASTURE SPECIES IS TOXIC OR IRRITANT TO FARM ANIMALS
OR HUMANS DESTROYS APPEARANCE OF LAWNS OR GARDENS HARBOURS INSECTS
OR DISEASE ORGANISMS THAT ATTACK OTHER PLANT SPECIES INTERFERES WITH WATER
INFRASTRUCTURE IMPACTS RECREATIONAL AREAS BEACHES LAKES ETC CONTAMI
)NVASIVENESS IN EXOTIC PLANTS
NATES SEED STOCKS INFESTS HAY ETC
%COSYSTEM IMPACTS $OCUMENTED ECOSYSTEM IMPACTS FORMS MONOCULTURES DOMINATES AQUATIC ECOSYSTEMS WITH MAJOR EFFECTS ON
mOW OXYGEN LIGHT ETC SIGNIlCANT NITROGEN lXER
%CONOMIC IMPACTS OTHER THAN THOSE LISTED HERE WERE OCCASIONALLY REPORTED EG MAKES HARVESTING DIFlCULT CHEMICALLY INHIBITS CROP
SPECIES HOWEVER THESE IMPACTS ARE CLOSELY RELATED TO AND ALWAYS OCCURRED WITH AND THEREFORE WERE CONSIDERED REDUNDANT AND NOT
INCLUDED AS SEPARATE IMPACTS
(4 -URPHY ET AL
MORE @NATURAL COMMUNITY TYPES BROUGHT AN ADDITIONAL POINT 4HUS IF THE
SPECIES OCCURRED IN COMMUNITY TYPES AND SEE 4ABLE THE TOTAL SCORE
WOULD BE POINTS 4HE TOTAL NUMBER OF POINTS SCORED WAS THEN MULTIPLIED BY
THE 3% RANK VALUE &OR EXAMPLE IF THE ABOVE SPECIES WAS RANKED AS 3% THE
#OMMUNITY )MPACT 3CORE #)3 WOULD BE ! SIMILAR SYSTEM WAS APPLIED TO
ECOSYSTEM IMPACTS %COSYSTEM )MPACT 3CORE %)3 &OR ECONOMIC IMPACT A SPE
CIES SCORED POINTS IF IT WAS NOTED TO HAVE IMPACTS RELATED TO DESCRIPTORS AND
SEE 4ABLE WHICH IN GENERAL INVOLVE BOTH CROP PRODUCTION LOSSES AND CONTROL
COSTS AND POINT IF IT WAS NOTED TO HAVE IMPACT IN ANY OTHER CATEGORY WHICH
PRIMARILY INVOLVE CONTROL COSTS ONLY 4HAT TOTAL SCORE WAS THEN MULTIPLIED BY
THE 3% RANK VALUE AS ABOVE %CONOMIC )MPACT 3CORE %CON)3
#RITERIA FOR DETERMINING WHETHER THE SCORE CORRESPONDED TO A @GREAT OR @SMALL
IMPACT ARE SHOWN IN 4ABLE @$4 )F A SPECIES WAS DETERMINED TO HAVE A GREAT
IMPACT IN ANY OF THE THREE IMPACT CATEGORIES THE OVERALL IMPACT WAS DETERMINED AS
@GREAT IN THE LANGUAGE OF $4 )T IS DIFlCULT TO REMOVE SUBJECTIVITY FROM THE SCOR
ING OF IMPACT 7E TRIED SEVERAL ALTERNATIVES TO THE ABOVE SCHEMES AND PRESENT ONE
OF THEM HERE 7E APPLIED AN ALTERNATIVE METHOD TO CALCULATING #)3 IF THE SPECIES
OCCURRED ONLY IN MEDIUM TO HIGHLY HUMAN
MODIlED COMMUNITIES IE OR
A SCORE OF WAS APPLIED AND A SCORE OF WAS APPLIED TO A SPECIES THAT OCCURRED IN
ANY NATURAL OR SEMI
NATURAL COMMUNITY 4HE SCORE WAS THEN MULTIPLIED BY THE 3%
RANK VALUE 7E ALSO MODIlED THE CRITERIA FOR DETERMINING WHETHER THE IMPACT WAS
SMALL OR GREAT 4HESE TWO MODIlCATIONS COMBINED RESULT IN THE ALTERNATIVE $AVIS
AND 4HOMPSON CLASSIlCATION $4 IN 4ABLE AND IN 4ABLE IN RESULTS
4ABLE #RITERIA TO BE SATISlED FOR DETERMINATION OF IMPACT AS @SMALL OR @GREAT IN TWO
ALTERNATIVE SCORING SCHEMES APPLIED TO $AVIS AND 4HOMPSONS CLASSIlCATION SCHEME
)MPACT $4 $4
3MALL 'REAT 3MALL 'REAT
#OMMUNITY #)3 ) )
%COSYSTEM %)3
%CONOMIC %CON)3 ) )
)N OUR APPLICATION OF 2ICHARDSON ET ALS SCHEME THE 3% RANK VALUE IS THE PRI
MARY FACTOR DRIVING THE DISTINCTION BETWEEN INVASIVE AND OTHER ALIEN SPECIES
3% RANK IS ALSO A KEY FACTOR IN OUR APPLICATION OF $AVIS AND 4HOMPSONS SCHEME
IN THE DIVISION BETWEEN GREAT AND SMALL IMPACTS ! CLOSE ASSOCIATION BETWEEN 3%
RANK AND THE RAW IMPACT FACTOR IE IMPACT SCORES PRIOR TO MULTIPLICATION BY 3%
RANK VALUE SHOULD RESULT IN A GREATER OVERLAP BETWEEN THE TWO SCHEMES 7E USED
)NVASIVENESS IN EXOTIC PLANTS
CORRELATION ANALYSIS TO DETERMINE HOW THE 3% RANK VALUE WAS RELATED TO THE RAW
IMPACT SCORES FOR COMMUNITY AND ECONOMIC IMPACTS ONLY SEVEN SPECIES SCORED
ABOVE IN THE ECOSYSTEM IMPACT CATEGORY SO %)3 WAS NOT TESTED FOR ASSOCIATION
4O EXAMINE DIFFERENCES BETWEEN GREAT AND SMALL IMPACT SPECIES IN LIFE
HISTORY
TRAITS WE USED CHI
SQUARED CONTINGENCY ANALYSIS WITH 9ATES CORRECTION )F TWO
WAY
CONTINGENCY TABLES SHOWED NON
INDEPENDENCE 0 THE &REEMAN
4UKEY
DEVIATE WAS USED TO DETERMINE IF ANY INDIVIDUAL CELL WAS EITHER UNDER
OR OVER
REPRESENTED COMPARED TO THE EXPECTED VALUE ,IFE
HISTORY VARIABLES FOR WHICH DATA
WERE COLLECTED ARE SHOWN IN 4ABLE
4ABLE ,IFE HISTORY TRAITS RECORDED FOR EACH SPECIES
,IFE HISTORY TRAIT 0ARAMETERS
3EED DORMANCY ,ONG YEARS 3HORT YEARS .ONE KNOWN
0RIMARY POLLINATION MODE )NSECT WIND SELF VARIOUS UNKNOWN
#LONAL ORGAN PRESENCE 0RESENT ABSENT
&LOWERING SEASON %ARLY MID LATE ALL SEASON
&LOWERING DURATION .UMBER OF MONTHS
3OIL MOISTURE PREFERENCE -OIST DRY VARIABLE
3EX HABIT (ERMAPHRODITE MONOECIOUS DIOECIOUS POLYGAMOUS
,IFE CYCLE !NNUAL AND BIENNIAL PERENNIAL VARIABLE
$OMINANT RECRUITMENT MODE 6EGETATIVE SEED VARIABLE
!S WITH MOST MACROECOLOGICAL DATASETS #ADOTTE ET AL OURS HAS ITS
LIMITATIONS 4HE TERM @WEED IN THE "IOLOGY OF #ANADIAN 7EEDS SERIES REFERS TO ANY
VASCULAR PLANT THAT PERPETUATES ITSELF IN HABITATS WHERE IT IS hNOT WANTEDv #AVERS
AND 7ARWICK A MEANING DIFFERENT FROM THAT COMMONLY USED FOR @WEEDS
IN THE ECOLOGICAL LITERATURE AND WHICH USUALLY INCLUDES THAT THE SPECIES HAS DETECT
ABLE ECONOMIC OR ENVIRONMENTAL IMPACTS EG 0YEK ET AL 2ICHARDSON
ET AL A 4HUS THESE SPECIES SHOULD NOT BE CONSIDERED TO HAVE BEEN ALREADY
CLASSIlED AS @WEEDS IN THE TRADITIONAL ECOLOGICAL SENSE RATHER THEY STILL VALIDLY
REPRESENT SPECIES IN THE RANGE OF CATEGORIES DElNED BY THE TWO SCHEMES 4HE SPE
CIES IN OUR LIST ARE BIASED TOWARDS SPECIES OCCURRING IN AGRICULTURAL AREAS SINCE THE
ORIGINAL PURPOSE OF THE SERIES WAS TO PROVIDE A BASIS FOR EFFECTIVE CONTROL METHODS
AND TO MEET THE NEEDS OF REGULATORY AGENCIES IN #ANADA #AVERS AND 7ARWICK
(OWEVER THE DATASET DOES INCLUDE A RANGE OF SPECIES OCCURRING IN BOTH
NATURAL AND AGRICULTURAL HABITATS AND INCLUDES BOTH ABUNDANT 3% AND 3% AND
RARE 3%
SPECIES )T IS IMPORTANT TO NOTE THAT WE ARE NOT ATTEMPTING TO TEST THE
ACCURACY OF EITHER SCHEME IN CLASSIFYING SPECIES INTO CATEGORIES SINCE THERE IS NO
@RIGHT ANSWER 2ATHER WE AIM TO DETERMINE IF EITHER SCHEME IS MORE FUNCTIONAL
OR OPERATIONAL BASED ON THE DATA AVAILABLE IN A TYPICAL SPECIES DATASET 4HUS ANY
(4 -URPHY ET AL
CONCLUSIONS WE MAKE AS TO WHICH CLASSIlCATION WAS @BETTER REFER ONLY TO WHICH
MODEL COULD BE APPLIED MORE USEFULLY GIVEN THE DATA AVAILABLE TO EVALUATE EACH
CRITERION FOR CATEGORIZATION
2%35,43
! TOTAL OF FAMILIES WAS REPRESENTED IN THE DATASET OF SPECIES 4ABLE
4HE !STERACEAE WAS THE ONLY FAMILY REPRESENTED IN ALL 3% RANKS WITH A TOTAL OF
SPECIES 0OACEAE WAS NEXT WITH A TOTAL OF SPECIES AND ONLY
3% SPECIES EXTREMELY RARE WERE NOT REPRESENTED &ABACEAE WAS NEXT WITH NINE
SPECIES ALL RANKED 3% 4HERE WERE EIGHT "RASSICACEAE IN 3% 3% AND
3% RANKS 4WENTY
TWO OF THE FAMILIES HAD ONLY OR SPECIES REPRESENT
ED IN THE DATASET !STERACEAE &ABACEAE AND "RASSICACEAE ARE ALL OVER
REPRESENTED
IN OUR DATASET COMPARED WITH THEIR REPRESENTATION IN THE EXOTIC mORA OF /NTARIO
AND AMONG NATIVE /NTARIO SPECIES SEE 4ABLE
!PPLICATION OF THE 2ICHARDSON ET AL SCHEME CLASSIlED OF THE TOTAL SPECIES
AS INVASIVE AS OPPOSED TO $AVIS AND 4HOMPSON WHERE APPROXIMATELY OF
SPECIES WERE CLASSIlED AS INVADERS USING OUR lRST CRITERIA $4 AND USING
THE ALTERNATIVE CRITERIA $4 4ABLE !LL BUT ONE OF THE SPECIES CLASSIlED AS
INVASIVE BY 2ICHARDSON ET AL WERE ALSO CLASSIlED AS INVASIVE BY BOTH $AVIS AND
4HOMPSON SCHEMES 4HE EXCEPTION WAS (ELIANTHUS TUBEROSUS WHICH WAS CLASSIlED
AS A SUCCESSIONAL COLONIZER IN BOTH $4 AND $4 DUE TO ITS ACTUALLY BEING CONSID
ERED NATIVE TO /NTARIO 4HE DISTINCTION DEPENDS UPON CONmICTING CONCLUSIONS BY
EXPERTS !LTHOUGH CONSIDERED NON
NATIVE TO /NTARIO BY SEVERAL AUTHORS -ORTON
AND 6ENN ;= .EWMASTER ET AL ;= AND 3COGGAN ;= MOST REGIONAL
mORAS AND mORAS FROM NEARBY AREAS CONSIDER THE SPECIES NATIVE EG 'LEASON AND
#RONQUIST )T IS LIKELY THAT AT SOME TIME IN THE FUTURE THE 3
RANK OF THIS
SPECIES WILL BE CHANGED FROM 3% TO 3 INDICATING A NATIVE STATUS RATHER THAN
EXOTIC IN THE PROVINCE .()# !LL SPECIES RANKED AS 3% WERE CLASSIlED BY
2ICHARDSON ET AL AS CASUAL AND BY $4 AND $4 AS NOVEL NON
INVASIVE COLONIZERS
4ABLE )N OUR APPLICATION OF BOTH CLASSIlCATIONS IT WAS NOT POSSIBLE FOR AN 3%
SPECIES TO BE REGARDED AS INVASIVE /F THE 3% SPECIES 2ICHARDSON ET ALS SCHEME
CLASSIlED AS INVADERS COMPARED WITH $4 AND $4 4ABLE
3% RANK VALUE WAS SIGNIlCANTLY CORRELATED WITH RAW COMMUNITY IMPACT SCORES
$4 R P N $4 R P N HOW
EVER THERE WAS NO CORRELATION BETWEEN 3% RANK VALUE AND ECONOMIC IMPACT SCORE
$4 R P N 4HUS SPECIES WITH GREATER ABUNDANCE
TENDED TO OCCUR IN MORE OF THE DElNED COMMUNITY TYPES INCLUDING NATURAL AND
SEMI
NATURAL COMMUNITIES THAN LESS ABUNDANT SPECIES BUT DID NOT NECESSARILY
FALL INTO MORE ECONOMIC IMPACT CATEGORIES )N $4 ALL 3% SPECIES HAD AN OVERALL
IMPACT OF @GREAT WHEREAS IN $4 THREE 3% SPECIES HAD ONLY @SMALL IMPACT
'REAT
AND SMALL
IMPACT SPECIES DIFFERED ONLY IN THEIR PROPORTIONS IN THE SEED DOR
MANCY LIFE
HISTORY PARAMETER LONG SHORT OR NONE $4 r P
)NVASIVENESS IN EXOTIC PLANTS
4ABLE &AMILY REPRESENTATION IN 3% RANK AND TOTAL NUMBER OF FAMILY MEMBERS IN THE
DATABASE COMPARED WITH PROPORTION OF FAMILY IN ALL SPECIES LISTED AS @EXOTIC IN /NTARIO AND
IN ALL NATIVE SPECIES OF /NTARIO
&AMILY 3% 2ANK 4OTAL OF OF OF
TOTAL FAMILY FAMILY
DATASET IN ALL IN NATIVES
EXOTICS OF
OF /NTARIO /NTARIO
N N
!MARANTHACEAE
!PIACEAE
!STERACEAE
"ORAGINACEAE
"RASSICACEAE
#ANNABACEAE
#ARYOPHYLLACEAE
#HENOPODIACEAE
#LUSIACEAE
#ONVOLVULACEAE
$IPSACACEAE
%UPHORBIACEAE
&ABACEAE
(ALORAGACEAE
,AMIACEAE
,YTHRACEAE
-ALVACEAE
/XALIDACEAE
0LANTAGINACEAE
0OACEAE
0OLYGONACEAE
0ORTULACACEAE
0OTAMOGETONACEAE
2ANUNCULACEAE
2OSACEAE
2UBIACEAE
3CROPHULARIACEAE
3OLANACEAE
6IOLACEAE
4OTAL
(4 -URPHY ET AL
4ABLE 3UMMARY OF RESULTS OF CLASSIlCATION SCHEMES FOR SPECIES IN EACH 3% RANK
REGIONAL DISTRIBUTION AND ABUNDANCE $4 $AVIS AND 4HOMPSON
3% 3% 3% 3% 3% 4OTAL
4OTAL IN 3% CATEGORY
2ICHARDSON ET AL
s #ASUAL
s .ATURALIZED
s )NVASIVE
$4
s 3UCCESSIONAL COLONIZER
s .OVEL
NON
INVASIVE COLONIZER
s )NVADER
$4
s 3UCCESSIONAL COLONIZER
s .OVEL
NON
INVASIVE COLONIZER
s )NVADER
$4 GREAT IMPACT
$4 SMALL IMPACT
$4 GREAT IMPACT
$4 SMALL IMPACT
$4 r P 2ESULTS OF THE &REEMAN
4UKEY DEVIATE TEST INDI
CATED THAT SPECIES WITH A SMALL IMPACT WERE SIGNIlCANTLY UNDER
REPRESENTED BY THE
PRESENCE OF A LONG DORMANCY PERIOD AND SPECIES WITH A GREAT IMPACT WERE UNDER
REPRESENTED BY PRESENCE OF SHORT ANDOR NO DORMANCY PERIOD !DDITIONALLY GREAT
AND SMALL
IMPACT SPECIES DIFFERED IN THEIR GEOGRAPHIC ORIGIN $4 r
P $4 r P 3PECIES HAVING A SMALL IMPACT WERE
UNDER
REPRESENTED IN ORIGINATING FROM %URASIA AND WERE OVER
REPRESENTED IN
ORIGINATING FROM !SIA PROPER AND THE -EDITERRANEAN AREA OF %UROPE 3PECIES WITH
A GREAT IMPACT WERE OVER
REPRESENTED IN ORIGINATING FROM %URASIA
)NVASIVENESS IN EXOTIC PLANTS
$)3#533)/.
$AVIS AND 4HOMPSONS CLASSIlCATION SCHEME WAS CLEARLY LESS DISCRIMINATING IN
DETERMINING INVASIVE SPECIES IN THIS DATASET 4HIS COULD BE AMELIORATED SOMEWHAT
BY INCREASING THE SCORE RANGE FOR @SMALL IMPACTS (OWEVER THE ORIGINAL SCORE
RANGE FOR %CON)3 ) TAKES INTO ACCOUNT THE FACT THAT AN 3% SPECIES WITH AN
IMPACT IN ONE ECONOMIC CATEGORY SUCH AS THE MOST COMMON CATEGORY n WOULD
SCORE A )T MAKES LITTLE SENSE TO CALL THIS A @SMALL IMPACT (OWEVER EVEN WITH
THE EXPANDED RANGE FOR SMALL ECONOMIC IMPACTS $4 THERE ARE ONLY SEVEN SPE
CIES THAT WOULD BE REASSIGNED FROM THE @INVASIVE CATEGORY TO THE @NON
INVA
SIVE COLONIZER CATEGORY )T IS NOTEWORTHY THAT 3% RANK AND THE ECONOMIC IMPACT
SCORE BEFORE ADJUSTING FOR ABUNDANCE WERE NOT CORRELATED 3% SPECIES WERE JUST
AS LIKELY TO HAVE A NUMBER OF DIFFERENT TYPES OF ECONOMIC IMPACTS AS 3% SPECIES
(OWEVER MORE ABUNDANT SPECIES WERE MORE LIKELY TO HAVE SPREAD TO NATURAL OR
SEMI
NATURAL HABITATS THAN LESS ABUNDANT SPECIES
/N A CASE
BY
CASE BASIS 2ICHARDSON ET ALS SCHEME WOULD SEEM TO BE MORE
PRACTICALLY USEFUL SINCE IT IS IN THEORY LESS SUBJECTIVE 9ET WHEN ATTEMPTING TO
CLASSIFY EVEN A RELATIVELY DETAILED AND CONSISTENT DATASET AS WE HAVE DONE THE
@SPREAD CRITERION PRESENTS ALMOST AS MANY PROBLEMS AND IS PERHAPS JUST AS SUBJEC
TIVE AS $AVIS AND 4HOMPSONS IMPACT CRITERION 7E ARE FORTUNATE WITH THE /NTARIO
DATASET TO HAVE THE DISTRIBUTION AND ABUNDANCE RANKING INFORMATION THAT CAN BE
USED AS A SURROGATE FOR THE @SPREAD CRITERION (OWEVER THE 3% RANKING SYSTEM HAS
SEVERAL LIMITATIONS IN ITS USE AS A SPREAD SURROGATE &IRST IT IS BASED ON CURRENT
ABUNDANCE IN THE PROVINCE 3%
RANKED SPECIES ARE AT THE LIMIT OF THE ABUNDANCE
RANKING BUT SOME MAY HAVE EXHAUSTED THE POTENTIAL RANGE OF SUITABLE HABITATS
AND BE NO LONGER CAPABLE OF FURTHER SPREAD WHILE OTHERS MAY NOT 3%
SPECIES
ALTHOUGH NOT CURRENTLY ABUNDANT MAY BE SPREADING AT A MORE RAPID RATE THAN 3%
AND 3% SPECIES AND THIS REMAINING POTENTIAL FOR RAPID SPREAD IS NOT CAPTURED IN
THE 3% RANKING 4HUS THE SPREAD CRITERION IS COMPLICATED BY THE TEMPORAL @SNAP
SHOT NATURE OF THE ASSESSMENT
! RELATED PROBLEM IS THAT THE 3% RANKING GIVES NO INDICATION AS TO WHERE THE
POPULATIONS ARE LOCATED RELATIVE TO EACH OTHER )N FACT VIRTUALLY ALL OF THE 3% SPE
CIES IN OUR DATASET HAVE BEEN RECORDED FROM POPULATIONS WIDELY DISTANT FROM ONE
ANOTHER IE KM DETERMINED THROUGH A VISUAL INSPECTION OF THE DISTRIBU
TION MAPS )F WE WERE TO BASE THE RATE
OF
SPREAD CRITERION ON THE TIME
SINCE
INTRODUCTION OF THE SPECIES AND THE AVERAGE DISTANCE BETWEEN PERSISTENT POPULATIONS
IN /NTARIO WE WOULD MOST LIKELY PLACE ALL THE SPECIES ON OUR LIST !PPENDIX
IN 2ICHARDSON ET ALS @INVASIVE CATEGORY 4HE SPREAD CRITERION FOR INVASIVES IE
FOR SEED
DISPERSED SPECIES M FROM PARENTAL PLANTS IN YEARS BEGINS TO
MAKE LITTLE SENSE ON REGIONAL AND LANDSCAPE SCALES ESPECIALLY WHEN MOST SPECIES
HAVE MODES OF DISPERSAL WITH A POTENTIAL TO CARRY PROPAGULES GREAT DISTANCES FROM
PARENTAL INDIVIDUALS
2ICHARDSON ET AL A ESTIMATED THAT
OF @INVADERS AS DElNED
BY THEIR CLASSIlCATION WOULD HAVE @HARMFUL DETECTABLE ENVIRONMENTAL OR ECONOMIC
(4 -URPHY ET AL
EFFECTS AND THAT THE REST ARE @BENIGN INVADERS WHOSE ENVIRONMENTAL OR
ECONOMIC IMPACTS ARE BEYOND ANY PRACTICAL DETECTION LIMITS "Y OUR ANALYSIS
OF THE SPECIES CLASSIlED AS INVASIVE BY 2ICHARDSON ET ALS SCHEME HAVE
LARGE ENVIRONMENTAL OR ECONOMIC IMPACTS AND MOREOVER BETWEEN AND
OF SPECIES DElNED AS NATURALIZED ALSO HAVE HARMFUL EFFECTS 4HIS IS PERHAPS NOT
SURPRISING SINCE OUR CRITERIA FOR NATURALIZATION IN 2ICHARDSON ET ALS SCHEME
INCLUDED A REQUIREMENT THAT THE SPECIES HAD NOT YET SPREAD TO NATURAL OR SEMI
NATURAL HABITATS 4HUS WHILE COMMUNITY IMPACTS WERE MINIMIZED OCCURRENCE IN
FEWER COMMUNITY TYPES ECONOMIC IMPACTS FOR THESE SPECIES MAY STILL HAVE BEEN
LARGE ENOUGH TO GENERATE AN OVERALL GREAT IMPACT &OR EXAMPLE 3% SPECIES THAT
ONLY OCCUR IN CULTIVATED lELDS IE 2ICHARDSON ET ALS @NATURALIZED SPECIES IN OUR
ANALYSIS WOULD STILL HAVE SCORED A GREAT IMPACT IF THEY COMPETE WITH AGRICULTURAL
SPECIES IN THOSE lELDS AND THEREFORE WOULD HAVE BEEN CLASSIlED AS INVADERS IN
$ 4S SCHEME )T SHOULD BE NOTED THAT THE DISTINCTION BETWEEN NATURALIZED
AND INVASIVE SPECIES WE USED HERE MAY NOT STRICTLY REPRESENT THE INTENTION OF
2ICHARDSON ET AL 3INCE 2ICHARDSON ET AL SUGGEST NATURALIZED PLANTS HAVE NOT
OVERCOME BARRIERS TO DISPERSAL WE LIMITED NATURALIZED SPECIES TO OCCURRENCE ONLY
IN HUMAN
MODIlED COMMUNITIES 2ICHARDSON ET ALS DElNITION IS UNCLEAR IN THIS
RESPECT THEY NOTED h.ATURALIZED PLANTSxDO NOT NECESSARILY INVADE NATURAL
SEMI
NATURAL OR HUMAN
MADE ECOSYSTEMSv
#ADOTTE ET AL HAVE STUDIED THE RELATIONSHIP BETWEEN 3% RANKS AND LIFE
HISTORY ATTRIBUTES IN /NTARIOS EXOTIC mORA N 4HEIR RESULTS SHOWED THAT
COMMON EXOTICS WERE MORE LIKELY TO HAVE A CLONAL ORGAN A LONGER mOWERING PERIOD
AND A %URASIAN ORIGIN 7E FOUND THAT IN TERMS OF LIFE
HISTORY TRAITS LARGE
IMPACT
SPECIES DIFFERED FROM SMALL
IMPACT ONES IN THE LIKELIHOOD OF HAVING A LENGTHY SEED
DORMANCY PERIOD #ADOTTE ET AL FOUND THE VARIABLE MOST STRONGLY ASSOCIATED WITH
INVASIVENESS AND HIGHER 3%
RANK WAS ORIGIN IN %URASIA AND %UROPE 3IMILARLY
IN OUR ANALYSIS SPECIES HAVING GREAT IMPACT CAME DISPROPORTIONATELY OFTEN FROM
%URASIA 4HIS SUPPORTS #ADOTTE ET AL IN THAT SPECIES NATIVE ELSEWHERE IN .ORTH
!MERICA WERE NOT MORE LIKELY TO HAVE A GREAT IMPACT IN /NTARIO HOWEVER OUR
SAMPLE SIZE IN THIS CATEGORY ; SPECIES= WAS SMALL #ADOTTE ET AL SUGGEST SEVERAL
REASONS WHY %URASIAN SPECIES MAY BE MORE SUCCESSFUL IN /NTARIO 3PECIES FROM
%URASIA MAY BE LESS PHYLOGENETICALLY RELATED TO SPECIES FROM .ORTH !MERICA AND
MAY BE MORE LIKELY TO HAVE EVOLVED ECOLOGICAL NOVELTIES IN LINE WITH $ARWINS
NATURALIZATION HYPOTHESIS ;$ARWIN = !LTERNATIVELY THE ENEMY RELEASE
HYPOTHESIS MAY BE RESPONSIBLE OR THE MATCHING OF PROVENANCE IE COMING FROM
A COMPARABLE CLIMATIC ZONE COULD BE ANOTHER POSSIBLE FACTOR
7E CONCLUDE THAT NEITHER FRAMEWORK IS ESPECIALLY USEFUL IN THEIR PRESENT FOR
MULATIONS FOR CLASSIFYING REGIONAL OR LARGER EXOTIC mORAS IE LISTS OF KNOWN NON
NATIVE SPECIES 4HE SEARCH FOR OPERATIONAL DElNITIONS OF TERMINOLOGY IN INVASIONS
ECOLOGY REMAINS ELUSIVE 2ELEVANT INFORMATION IS SIMPLY NOT AVAILABLE AND THE USE
OF SURROGATES FOR CRITERIA SUCH AS @SPREAD AND @IMPACT ARE LIKELY TO INTRODUCE JUST
AS MUCH SUBJECTIVITY INCONSISTENCY AND CONFUSION IN THE LITERATURE AS ALREADY
EXISTS &AILURE TO OPERATIONALISE DElNITIONS USED IN CLASSIFYING EXOTIC mORA DATASETS
)NVASIVENESS IN EXOTIC PLANTS
WILL LIKELY CONTINUE TO LEAD TO AMBIGUOUS GENERALIZATIONS AND PREDICTIONS
.OTWITHSTANDING THIS AMBIGUITY COMPARATIVE STUDIES OF INVASIVE mORAS HAVE PRO
VIDED NEW INSIGHTS IN THE UNDERSTANDING OF GENERAL PATTERNS OF PLANT INVASION AND
THE VALUE OF THESE STUDIES IS NOT TO BE DIMINISHED 2ATHER WHEN CLASSIFYING LISTS
OF mORA THE CRITERIA USED TO ASSIGN EACH SPECIES TO A CATEGORY SHOULD BE EXPLICITLY
STATED SO THAT RESEARCHERS CAN COMPARE SPECIES BASED ON LIKE CRITERIA RATHER THAN
THE CATEGORY ITSELF WHICH MAY HAVE BEEN DERIVED FROM UNLIKE DATA !S 0YEK ET AL
CONCLUDE IN THEIR RECENT PAPER ON THE TREATMENT OF INVASIONS TERMINOLOGY
IN REGIONAL mORAS STATEMENTS PRECEDED BY @PROBABLY OR @POSSIBLY MAY BE TEMPO
RARILY THE MOST HONEST WAY OF CLASSIFYING SOME TAXA
#OLAUTTI AND -AC)SAAC RECENTLY PROPOSED A SUPPLEMENTARY LEXICON TO
CURRENT TERMINOLOGY IN INVASIONS ECOLOGY 4HEIR FRAMEWORK ATTEMPTED TO ELIMI
NATE THE NEED FOR UNIVERSAL DElNITIONS OF CURRENT TERMS BY USING OPERATIONAL TERMS
WITH NO A PRIORI MEANING IE AS STAGES IN A PROCESS
BASED MODEL 4HE FRAMEWORK
IS BASED ON CURRENT MODELS THAT ENVISION PROCESSES IN IMMIGRATION AND NATURALIZA
TION AS A SERIES OF CONSECUTIVE STAGES 4HE GOAL OF THE FRAMEWORK IS TO SUPPLEMENT
AMBIGUOUS TERMS WITH THE STAGE
BASED TERMINOLOGY 3UCH A FRAMEWORK IS LOGICAL
AND INTUITIVELY APPEALING AND WOULD LIKELY LEAD TO SOME CONSISTENCY IN INDIVIDUAL
POPULATION
BASED STUDIES OF INVASIVE SPECIES AS WOULD THAT OF 2ICHARDSON ET AL
;A= (OWEVER IT IS ALSO UNLIKELY TO BE USEFUL FOR CLASSIFYING LISTS OF ALIEN SPE
CIES FOR COMPARATIVE PURPOSES SINCE THE SAME DATA LIMITATIONS WOULD APPLY AS
DESCRIBED ABOVE FOR 2ICHARDSON ET AL AND $AVIS AND 4HOMPSONS SCHEMES
! 02/0/3%$ -/$)&)#!4)/. 4/ #522%.4 #/.#%045!, &2!-%7/2+3
&/2 4(% ).6!3)/.3 02/#%33
#OLAUTTI AND -AC)SAAC NOTED THAT TERMINOLOGY USED TO DESCRIBE INTRODUCED
SPECIES ARE MISNOMERS IN THE SENSE THAT INTRODUCED NATURALIZED OR INVASIVE SPECIES
ETC ARE OF COURSE INTRODUCED NATURALIZED OR INVASIVE POPULATIONS ! FOCUS ON INVA
SIONS AT A POPULATION LEVEL RATHER THAN A SPECIES LEVEL HAS IMPORTANT IMPLICATIONS
FOR THE DEVELOPMENT OF INVASIONS ECOLOGY )NDEED IT MAKES LITTLE SENSE TO ASSIGN
CATEGORICAL PROPERTIES TO A @SPECIES 2ATHER DEMOGRAPHIC ATTRIBUTES OF THE ADVEN
TIVE POPULATION AND OF ANY METAPOPULATION OR OTHER REGIONALLY CONNECTED GROUP
SEE -URPHY AND ,OVETT
$OUST NEED TO BE CHARACTERIZED AS THE ATTRIBUTES OF
LONGEVITY FECUNDITY AND MORTALITY RATES ARE THE KEY DETERMINANTS OF AN INVADERS
STATUS WITHIN THE ECOLOGICAL CONTINUUM REPRESENTING THE INVASION PROCESS AND
EXTENDING ACROSS THE ARC FROM IMMIGRATION TO NATURALIZATION &URTHERMORE WHEN
IT DOES HAPPEN CLASSIlCATION SHOULD OCCUR AT A DElNED TEMPORAL OR SPATIAL SCALE
AS POPULATIONS OF SPECIES WILL APPEAR TO BE IN DIFFERENT PLACES ON THE CONTINUUM
DEPENDING ON THE SCALE OF THE ASSESSMENT
4HE @BLINKING LIGHTS METAPHOR OF METAPOPULATION EXTINCTION AND COLONIZATIONS
MAY BE USEFUL IN THE INVASIONS PROCESS TOO ! METAPOPULATION DESCRIBES A @POPU
LATION CONSISTING OF A NUMBER OF LOCAL POPULATIONS IN THE SAME SENSE IN WHICH
(4 -URPHY ET AL
A LOCAL POPULATION IS A POPULATION CONSISTING OF MANY INDIVIDUALS !CCORDING TO
THE ,EVINS CLASSICAL METAPOPULATION CONCEPT ALL LOCAL POPULATIONS HAVE
A SUBSTANTIAL PROBABILITY OF EXTINCTION AND THEREFORE LONG TERM PERSISTENCE OF A
SPECIES IS REGULATED AT THE REGIONAL OR METAPOPULATION LEVEL (ANSKI
-ETAPOPULATION THEORY PROPOSES THAT THE REGIONAL POPULATION PERSISTS AS THE RESULT
OF A BALANCE BETWEEN LOCAL POPULATION EXTINCTION IN PATCHES AND PATCH MIGRATIONS
LEADING TO COLONIZATION 4HE KEY TO POPULATION PROCESSES LIES IN UNDERSTANDING THE
SHIFTING MOSAIC OF PATCH OCCUPANCY AS OPPOSED TO DETAILS OF WITHIN
PATCH EVENTS
+AREIVA AND 7ENNERGREN &RECKLETON AND 7ATKINSON DESCRIBE
OTHER REGIONAL ENSEMBLES OF PLANT POPULATION DYNAMICS AT LARGE SCALES )N THE
FOLLOWING WE VISUALIZE 2ICHARDSON ET ALS CATEGORIES OF ALIEN SPECIES
PLUS AN ADDITIONAL CATEGORY @SPREADING POPULATIONS BETWEEN NATURALIZED AND
INVASIVE STAGES AND THE BARRIERS THEY SURMOUNT THROUGH THE INVASIONS PROCESS
IN A POPULATION
BASED CONTEXT 4HE FOUR CATEGORIES OF POPULATIONS CAN BE DESCRIBED
IN TERMS OF THE METAPOPULATION PARAMETERS OF EXTINCTION AND COLONIZATION AS
SHOWN IN 4ABLE
4ABLE -ETAPOPULATION CONDITIONS FOR POPULATION THROUGH THE INVASION PROCESS
E EXTINCTION C COLONIZATION
#ATEGORY3TAGE -ETAPOPULATION DYNAMICS
#ASUAL E C
.ATURALIZED E C
3PREADING E C
)NVASIVE E C
#ASUAL
!DVENTIVE INDIVIDUALS APPEAR AND MAY BE INTRODUCED ON MULTIPLE OCCASIONS
BEFORE A POPULATION @TAKES OR ESTABLISHES 4HE RECURRENT BUT EPHEMERAL INTRODUC
TIONS GENERATE THE STATUS OF @CASUAL i.e., the iNTRODUCED SPECIES IS AT THE EARLY STAGE
OF BEING AN INTERMITTENT MEMBER OF THE NEW mORA 4HE POPULATION MAY @BLINK
OFF AND ON AGAIN AS IT FAILS TO SUSTAIN ITSELF OVER LONGER PERIODS BUT OCCASIONALLY
PERSISTS OR BECOMES RE
ESTABLISHED BECAUSE OF REPEATED INTRODUCTIONS FROM SOURCE
POPULATIONS OUTSIDE THE ADVENTIVE AREA 4HESE POPULATIONS ESTABLISH BECAUSE SOME
INDIVIDUALS OF THE SPECIES HAVE SURMOUNTED A MAJOR GEOGRAPHICAL BARRIER LONG
DISTANCE BARRIER ! OF &IG
.ATURALIZED
%VENTUALLY A CASUAL POPULATION MAY BLINK ON AND REMAIN ON FOR AN EXTENDED
PERIOD 4HE lRST LOCAL POPULATION MEETS CRITERIA SET FOR STEADY
STATE MAINTENANCE
)NVASIVENESS IN EXOTIC PLANTS
A SUSTAINABLE POPULATION 4HIS CORRESPONDS TO THE CONCEPT IN 0OPULATION 6IABILITY
!NALYSIS OF AN -60 MINIMUM VIABLE POPULATION THE NUMBER OF INDIVIDUALS THAT
ENSURES A POPULATIONS PERSISTENCE ;3HAFFER = )N THE VOCABULARY OF INVA
SIONS THIS CAN BE TERMED @NATURALIZATION !T THAT POINT THE FOUNDER POPULATION
HAS BECOME RESILIENT TO ENVIRONMENTAL AND DEMOGRAPHIC STOCHASTICITIES AND THE
PROBABILITY OF CHANCE EXTINCTION HAS DECREASED $EMOGRAPHIC ATTRIBUTES OF THIS
POPULATION INCLUDE A SIGNIlCANT BREEDING POPULATION FOR PLANTS AN ESTABLISHED
SEED BANK IS A BENElT SO THAT YEAR
TO
YEAR VARIATIONS IN FECUNDITY ARE NOT DIRECTLY
EVIDENT IN RECRUITMENT )N ADDITION INDIVIDUALS LIKELY DEVELOP FACILITATIVE ASSOCIA
TIONS SUCH AS POLLINATION SEED DISPERSAL AND PERHAPS MYCORRHIZAL ASSOCIATIONS
)NDIVIDUALS IN THESE POPULATIONS HAVE OVERCOME BARRIERS TO SURVIVAL IN ENVIRON
MENTAL CONDITIONS BARRIER " IN &IG AT THE SITE OF INTRODUCTION SOIL AND CLIMATE
FOR EXAMPLE AS WELL AS BARRIERS TO THE PREVENTION OF CONSISTENT AND LONG
TERM
GENERATION OF RECRUITS TO THE POPULATION MATING SYSTEM COMPATIBILITY FERTILITY
FECUNDITY AND OTHER REPRODUCTIVE BARRIERS BARRIER # IN &IG
3PREADING METAPOPULATION OR OTHER REGIONAL ENTITY
@3PREADING POPULATIONS CAN BE VISUALIZED AS INCLUDING A POPULATION WITH A LIGHT
PERMANENTLY ON THE NATURALIZED POPULATION SURROUNDED BY ADJACENT POPULA
TIONS THAT MAY BLINK ON AND OFF LIKE CASUAL POPULATIONS BUT THE NATURALIZED
POPULATION IS THE SOURCE RATHER THAN THE SOURCE BEING FROM BEYOND THE ADVENTIVE
AREA 4HIS STAGE INVOLVES THE RELATIVELY SHORT
DISTANCE DISPERSAL OF PROPAGULES TO
ADJACENT SUITABLE SITES THEREBY OVERCOMING BARRIER $ IN &IG 3PREAD FROM THE
INITIAL NATURALIZED POPULATION TO OTHER SUITABLE SITES REQUIRES SUFlCIENT DISPERSAL BY
PROPAGULES ANDOR SUFlCIENT LANDSCAPE CONNECTIVITY BETWEEN SUITABLE SITES SUCH
THAT LOCAL EXTINCTIONS MAY BE REPLACED BY RECOLONIZATIONS FROM NEARBY NATURAL
IZED POPULATIONS IF NOT FROM THEIR OWN SEED BANK 4HE DISCOVERY OF CONTIGUOUS
POPULATIONS OF DIFFERING SIZE AROUND AN OLDER ONE IS WELL DOCUMENTED IN THE HER
BARIUM COLLECTIONS USED TO DEVELOP THE DISTRIBUTION MAPS OF THE SPECIES STUDIED
IN THE "IOLOGY OF #ANADIAN 7EEDS SERIES 4HIS OCCURS WHEN INITIAL LONG
DISTANCE
DISPERSAL EG INTO A NEW PART OF THE PROVINCE OR WATERSHED IS FOLLOWED BY SUB
SEQUENT OUTBREAKS OF POPULATIONS IN THE GENERAL AREA AS SUITABLE HABITAT BECOMES
OCCUPIED 4HIS CORRESPONDS TO THE STAGE WE TERM LANDSCAPE @SPREADING &IG
4HESE POPULATIONS HAVE OVERCOME BARRIERS TO RELATIVELY SHORT
DISTANCE DISPERSAL
IN THE IMMEDIATE AREA BARRIER $ IN &IG AND ENVIRONMENTAL CONDITIONS AT ADJA
CENT HUMAN
MODIlED AREAS BARRIER % IN &IG ASSUMING AVAILABILITY OF SUITABLE
DISTURBED SITES 3PREADING POPULATIONS ARE SELF
SUSTAINING WITHOUT INPUT FROM
SOURCES OUTSIDE THE ADVENTIVE REGION BUT MAY BE RECOLONIZED FROM NEARBY SITES IF
LOCAL EXTINCTION OCCURS
(4 -URPHY ET AL
)NVASIVE METAPOPULATION OR OTHER REGIONAL ENTITY
4HE IMAGE OF THE SPREADING POPULATIONS AT LEAST ONE PERMANENTLY @ON LIGHT SUR
ROUNDED BY NUMEROUS BLINKING LIGHTS IS REPEATED IN MANY PARTS OF THE ADVENTIVE
RANGE 3PREADING POPULATIONS BECOME THEMSELVES THE PRIMARY SOURCE FOR NEW
CASUAL NATURALIZED AND SPREADING POPULATIONS THOUGH SOME MAY ALSO CONTINUE
TO BECOME ESTABLISHED THROUGH SEPARATE DISPERSAL EVENTS FROM THE ORIGINAL SOURCE
REGION 4HESE POPULATIONS HAVE OVERCOME BARRIERS TO LONG
DISTANCE DISPERSAL IN
THE ADVENTIVE ENVIRONMENT
3OME RESEARCHERS ALSO CONSIDER THAT ABIOTIC AND BIOTIC BARRIERS TO ESTABLISHMENT
IN NATURAL HABITATS NEEDS TO BE OVERCOME AT THIS STAGE OF THE INVASIONS PROCESS OTH
ERS CONSIDER THAT THIS BARRIER MAY BE OVERCOME BY NATURALIZED POPULATIONS SEE
2ICHARDSON ET AL ;A= &ROM AN ECOLOGICAL STANDPOINT THE MORE IMPORTANT
DEVELOPMENT OCCURS WHERE INVASIVE POPULATIONS SPREAD TO NATURAL ENVIRONMENTS
(OWEVER FROM A MANAGEMENT AND ECONOMIC POINT OF VIEW INVASIVE POPULATIONS
THAT ONLY OCCUR IN FOR EXAMPLE AGRICULTURAL SETTINGS ARE EQUALLY AS IMPORTANT
4HUS METAPOPULATIONS OF THE INTRODUCED SPECIES SHOULD NOT HAVE TO OCCUR IN NATU
RAL HABITAT TO BE REGARDED AS INVASIVE 7E DO THINK IT IS USEFUL IN OUR FRAMEWORK
HOWEVER TO DISTINGUISH ECOLOGICALLY INVASIVE POPULATIONS FROM INVASIVE POPULA
TIONS IN DISTURBED HABITATS )N TERMS OF THE MAJOR ECOLOGICAL IMPACTS OF INVASIVE
SPECIES ON BIODIVERSITY IT MAY BE IMPORTANT TO UNDERSTAND HOW SPECIES THAT ARE
CAPABLE OF INVADING NATURAL HABITATS DIFFER FROM THOSE THAT REMAIN IN DISTURBED OR
AGRICULTURAL COMMUNITIES AND HAVE PRIMARILY ECONOMIC IMPACTS
0/05,!4)/. 6)!"),)49 !.!,93)3 06! 4/ 42!#+ 0/05,!4)/.3
/& ./.
.!4)6% 30%#)%3
0OPULATION VIABILITY ANALYSIS HAS BEEN USED TRADITIONALLY TO ESTIMATE THE RISK
OR TIME TO EXTINCTION FOR AN ENDANGERED SPECIES AND COULD EQUALLY BE APPLIED
TO PREDICT THE POTENTIAL FOR ESTABLISHMENT GROWTH OR PERSISTENCE IN NON
NATIVE
SPECIES 06! EVALUATES THE LIKELIHOOD THAT A GIVEN SPECIES WILL PERSIST FOR A
GIVEN TIME INTO THE FUTURE 4HE GOAL OF 06! IS OFTEN CONSERVATION OR MANAGE
MENT OF RARE OR ENDANGERED SPECIES THIS INCLUDES IDENTIFYING THREATS TO SPECIES
PERSISTENCE AND APPLYING THEORETICAL CONCEPTS OF POPULATION AND METAPOPULA
TION ECOLOGY TO IMPROVE SURVIVORSHIP SEE EG "ROOK ET AL !KAKAYA AND
3JGREN
'ULVE
-ORRIS AND $OAK SUMMARIZED THREE GENERAL LEVELS OF COMPLEXITY AVAIL
ABLE FOR 06! THAT INCORPORATE MULTI
SITE DATA !LL RELY ON USE OF MATRICES TO
CHARACTERIZE POPULATION STATUS OVER TIME 4HE SIMPLEST APPROACH IS TO TRACK THE
NUMBER OF POPULATIONS SENSU METAPOPULATION AND OTHER REGIONAL ENTITIES THAT
IS TO ESTIMATE THE PRESENCE OR ABSENCE OF POPULATIONS IN LOCAL SITES OF SUITABLE
HABITAT )T IS RELATIVELY EASY TO ENUMERATE SITES WHERE AN INTRODUCED SPECIES IS
REPORTED ! SECOND COUNT
BASED CENSUS APPROACH IS A MORE USEFUL FORM OF 06!
)NVASIVENESS IN EXOTIC PLANTS
BUT ESTIMATES OF MIGRATION RATES AND MEAN VARIANCE AND COVARIANCE IN POPULA
TION GROWTH RATES ARE NECESSARY TO DEVELOP SUCH MODELS -OST USEFUL OF ALL BUT
ALSO MORE DATA DEMANDING IS A 06! MODEL INCLUDING DEMOGRAPHIC ANALYSIS AT
EACH SITE BASED ON FOR EXAMPLE KNOWLEDGE OF BIRTH AND DEATH RATES AND RATES
OF INDIVIDUAL TRANSITIONS THROUGH SIZE OR AGE CLASSES THAT GOVERN FECUNDITY RATES
3UCH INFORMATION WOULD ALLOW CONSTRUCTION OF A MULTI SITE MATRIX MODEL WHICH
COULD MORE ACCURATELY PREDICT THE FATE OF THE INTRODUCED SPECIES
7HILE COUNT BASED AND DEMOGRAPHY BASED 06! PROVIDE MORE ROBUST RESULTS
OUR ANALYSIS HERE HAS SHOWN THAT THIS LEVEL OF SPECIES SPECIlC INFORMATION IS OFTEN
UNAVAILABLE AND MAY BE INSUFlCIENT TO ESTIMATE THE PARAMETERS OF THE MODEL
(OWEVER TRACKING PRESENCE OR ABSENCE OF A NON NATIVE SPECIES IN SUITABLE HABI
TAT IS SUFlCIENT TO ESTIMATE THE PARAMETERS OF THE SIMPLEST 06! MODELS 0ATCH
/CCUPANCY -ODELS 0/- !KAKAYA AND 3JGREN 'ULVE 3UCH MODELS
AS WITH ALL @PATCH MODELS OF METAPOPULATION DYNAMICS (ANSKI ASSUME
THAT LOCAL DYNAMICS OCCUR RAPIDLY IN COMPARISON TO METAPOPULATION DYNAMICS
IE CHANGES IN POPULATION SIZES OF LOCAL POPULATIONS TAKE PLACE MORE RAPIDLY
THAN COLONIZATIONS OF CURRENTLY EMPTY HABITAT PATCHES 7ITH THIS ASSUMPTION
IT IS REASONABLE TO FOCUS ENTIRELY ON THE PRESENCE OR ABSENCE OF THE SPECIES OF
INTEREST IN THE AVAILABLE HABITAT PATCHES 'IVEN THIS THE PERSISTENCE OF THE SPECIES
CAN BE MODELED BY EXAMINING THE PROBABILITY OF EXTINCTION OF AN OCCUPIED PATCH
AND THE PROBABILITY OF COLONIZATION OF AN EMPTY PATCH
3IMPLE 0/-S ARE BASED ON CLASSICAL METAPOPULATION MODELS SENSU ,EVINS
AND THUS TREAT HABITAT PATCHES AS IF THEY WERE IDENTICAL AND MOVEMENT
BETWEEN ALL PAIRS OF PATCHES IS EQUALLY LIKELY (ANSKI AND 'YLLENBERG (ANSKI
2ECENTLY MORE DATA INTENSIVE AND SPATIALLY EXPLICIT MODELS HAVE BEEN
CREATED SEE EG (ANSKI -OILANEN 7E CONSTRUCTED A SIMPLE 0/-
BASED SOFTWARE MODULE THAT DEMONSTRATES THE USEFULNESS OF 06! FOR MODELING
CONTRASTING LEVELS OF INVASIVENESS IN EXOTIC SPECIES 4HE MODEL USES TWO INPUTS THE
PROBABILITY THAT AN OCCUPIED PATCH WILL BECOME VACANT DUE TO LOCAL EXTINCTION AND
THE PROBABILITY OF AN UNOCCUPIED PATCH BECOMING OCCUPIED BY MIGRANTS FROM AN
OCCUPIED PATCH IN EACH TIME STEP OR ITERATION 3UCH VALUES MAY BE ESTIMATED FROM
A SERIES OF @SNAPSHOTS OF THE PRESENCE OR ABSENCE OF A SPECIES THROUGH TIME AND
WITH PARTICULAR LIFE HISTORY CHARACTERISTICS INmUENCING COLONIZATION ABILITY 'OTELLI
AND 4AYLOR A B ESTIMATED SUCH PROBABILITIES FOR lSH ASSEMBLAGES USING THE
PROBABILITY OF COLONIZATION AS THE NUMBER OF UNOCCUPIED SITES IN YEAR T THAT WERE
OCCUPIED IN YEAR T DIVIDED BY THE NUMBER OF SITES THAT WERE CENSUSED IN YEAR
T 3IMILARLY THE PROBABILITY OF EXTINCTION CAN BE ESTIMATED AS THE NUMBER OF SITES
OCCUPIED IN YEAR T THAT WERE UNOCCUPIED IN YEAR T DIVIDED BY THE NUMBER
OF SITES OCCUPIED IN YEAR T
5SING A COMBINATION OF PROBABILITY LEVELS FOR EXTINCTION HIGH MEDIUM AND
LOW 0EXT VALUES OF AND RESPECTIVELY AND THREE FOR THE PROB
ABILITY OF COLONIZATION HIGH MEDIUM AND LOW 0COL VALUES OF AND
RESPECTIVELY CLEAR CONSISTENT OUTCOMES EMERGED %ACH WAS PLOTTED FOR RAN
DOM ITERATIONS TO GENERATE PLOTS THAT CORRESPOND TO THE FOUR CATEGORIES OF NON NATIVE
(4 -URPHY ET AL
POPULATIONS OF SPECIES 4HE VALUES USED TO PARAMETERIZE THIS MODEL WERE BASED ON
A RANGE OF EXTINCTION AND COLONIZATION ESTIMATES GIVEN FOR A VARIETY OF ORGANISMS
SEE EG 7AHLBERG ET AL -OILANEN ET AL -OILANEN 'OTELLI AND
4AYLOR A B 3IMULATIONS WERE RUN IN A RANDOM LANDSCAPE WITH AVAIL
ABLE PATCHES INITIALLY WITH A SINGLE PATCH OCCUPIED REPRESENTING AN INTRODUCTION
)N EACH CASE THERE WERE ITERATIONS IE SEPARATE INTRODUCTIONS
4ABLE SUMMARIZES THE SIMULATION OUTPUT AND &IG DEPICTS THE CHANGE IN
PROPORTION OF PATCHES OCCUPIED OVER TIME FOR EACH SCENARIO 4HE MODELS PROVIDE
AN EFFECTIVE FRAMEWORK FOR GENERALIZATION WE CAN STATE THAT WHERE 0COL ) 0EXT
THE SPECIES IS CASUAL TO NATURALIZED 7HERE 0COL 0EXT THE SPECIES IS SPREADING
IT IS ONLY WHERE 0COL 0EXT THAT THE SPECIES IS LIKELY TO BECOME INVASIVE 4HIS GEN
ERALIZATION SUGGESTS THAT MOST INTRODUCTIONS SHOULD RESULT IN CASUAL OR NATURALIZED
POPULATIONS OF SPECIES OF THE OUTCOMES WITH ONLY BEING SPREADING AND
INVASIVE
)NDEPENDENT OF 0COL IF 0EXT IS HIGH THEN THE SPECIES IS MERELY CASUAL %VEN IF THE
SPECIES HAS LOW 0COL AND A MEDIUM 0EXT IT WILL OCCUR AS A CASUAL 0OPULATIONS MAY
BLINK OFF AND ON AGAIN BUT FAIL TO SUSTAIN THEMSELVES OVER LONGER PERIODS ONLY
OCCASIONALLY PERSISTING 3IMULATIONS SHOWED THAT ON AVERAGE WITH LOW COLONIZA
TION RATE AND MEDIUM EXTINCTION RATE ALL POPULATIONS BLINKED OFF AFTER
ITERA
TIONS HOWEVER TWO OUTLIER POPULATIONS PERSISTED FOR THE FULL RUNS .ATURALIZED
SPECIES WERE THOSE WITH AN EQUIVALENT EQUALLY WEAK OR STRONG 0COL AND 0EXT THAT
IS HAVING LOW 0COL AND 0EXT LOW
LOW OR MEDIUM 0COL AND 0EXT MEDIUM
MEDIUM
7HILE MOST OF THE INTRODUCTIONS ENDED IN EXTINCTION LOW
LOW AND
MEDIUM
MEDIUM PERSISTED FOR ALL ITERATIONS
3PREADING SPECIES WOULD HAVE A PROPORTIONATELY GREATER PROBABILITY OF 0COL AND
0EXT HIGH
MEDIUM OR MEDIUM
LOW )NVASIVE SPECIES ARE DISTINGUISHED BY HAVING
A HIGH 0COL AND A LOW 0EXT 7E NOTE THESE EXEMPLARS REPRESENT AN ECOLOGICAL CON
TINUUM FROM CASUAL TO INVASIVE )T IS INTERESTING THAT THE CURVES HAVE DISTINCTIVE
SHAPES &OR EXAMPLE ALTHOUGH SPREADING SPECIES MAY REACH COMPARABLE LEVELS
OF OCCUPANCY AS INVASIVES SEE &IGS B AND F SPREADING SPECIES SHOW A TIME LAG
ASSOCIATED WITH PROLONGED LOW SITE OCCUPANCY RESULTING IN A GREATER RISK OF hINVA
SION FAILUREv
!NY INTRODUCTION HAD THE POSSIBILITY OF FAILURE %VEN THE INVASIVE SPECIES IN
OUR SIMULATION FAILED TO ESTABLISH SIX PERCENT OF THE TIME FOLLOWING INTRODUCTION
)NDEED AS NOTED EARLIER THE LITERATURE SUGGESTS THAT MOST INTRODUCTIONS FAIL TO
ESTABLISH AND EVEN THOSE THAT SUCCESSFULLY ESTABLISH AND PERSIST MAY HAVE HAD
MANY PREVIOUS FAILED ATTEMPTS )T IS IMPORTANT TO NOTE THAT EVEN WHEN POPULATIONS
HAVE BECOME INVASIVE THE PROCESS IS NOT STATIC 2ATHER POPULATIONS MAY STILL BE
VULNERABLE TO DEMOGRAPHIC AND ENVIRONMENTAL FACTORS ACTING TO INCREASE EXTINC
TION AND DECREASE COLONIZATION RATES 7E HAVE OBSERVED DRAMATIC CHANGES OVER
THE PAST lFTEEN YEARS IN SOUTHWESTERN /NTARIO POPULATIONS OF PURPLE LOOSESTRIFE
,YTHRUM SALICARIA SEE TOO -AL ET AL &OLLOWING A SURGE OF COLONIZATIONS
OVER THE PAST
DECADES AND ABUNDANT LARGE LOCAL POPULATIONS MANY OF THESE
HAVE GONE EXTINCT OR BECOME EXTREMELY DIMINISHED IN RECENT YEARS
4ABLE 2ESULTS OF POPULATION VIABILITY ANALYSIS FOR POPULATIONS WITH DIFFERING EXTINCTION AND COLONIZATION PROBABILITIES
CASUAL NATURALIZED SPREADING INVASIVE
2EFERENCE &IG A D G H E I B F C
#OLONIZATION PROBABILITY HIGH MED LOW LOW MED LOW HIGH MED HIGH
%XTINCTION PROBABILITY HIGH HIGH HIGH MED MED LOW MED LOW LOW
-EAN SE
PATCH OCCUPANCY
0ROPORTION OF INTRODUCTIONS
ENDING IN EXTINCTION
-EAN SE TIME TO PATCHES
)NVASIVENESS IN EXOTIC PLANTS
GOING EXTINCT ITERATIONS
(4 -URPHY ET AL
&IG 2ESULTS OF 0OPULATION 6IABILITY !NALYSIS FOR POPULATIONS WITH DIFFERING EXTINCTION
AND COLONIZATION RATES SEE 4ABLE A CASUAL B SPREADING C INVASIVE D CASUAL
E NATURALIZED F SPREADING G CASUAL H CASUAL I NATURALIZED
4HE POPULATION
BASED FRAMEWORK WE HAVE DESCRIBED IS INFORMED BY THE MODI
lED SCHEME IN 4ABLE WHERE THE ENVIRONMENTAL AND LIFE
HISTORY PARAMETERS THAT
ARE CASE DEPENDENT ARE NOT THE FOCUS AS IN 2ICHARDSON ET AL ;A= BUT RATHER
THE lLTER REGULATING THE CHANGING STATUS OF POPULATIONS OF AN INTRODUCED SPECIES
7E ACKNOWLEDGE IT IS DIFlCULT TO BUILD OPERATIONAL DElNITIONS FOR REGIONAL LISTS OF
ALIEN mORA WITH THE LIMITED LIFE
HISTORY DATA ON RECORD AND SUGGEST THAT IT IS TIME
FOR RESEARCH IN INVASIONS ECOLOGY TO MOVE BEYOND COMPARATIVE ANALYSES OF REGIONAL
)NVASIVENESS IN EXOTIC PLANTS
!PPENDIX 3PECIES INCLUDED IN THE DATASET WITH THEIR ASSOCIATED "IOLOGY OF #ANADIAN
7EEDS "#7 SERIES NUMBER AND REFERENCE
"#7 &AMILY 3PECIES 2EFERENCE
SERIES
!MARANTHACEAE !MARANTHUS POWELLII -ULLIGAN
!MARANTHACEAE !MARANTHUS RETROmEXUS -ULLIGAN
!PIACEAE !NTHRISCUS SYLVESTRIS $ARBYSHIRE ET AL
!PIACEAE $AUCUS CAROTA -ULLIGAN
!STERACEAE !CHILLEA MILLEFOLIUM
SSP MILLEFOLIUM -ULLIGAN
!STERACEAE !CROPTILON REPENS -ULLIGAN
!STERACEAE !RCTIUM LAPPA -ULLIGAN
!STERACEAE !RCTIUM MINUS -ULLIGAN
!STERACEAE !RTEMISIA ABSINTHIUM #AVERS
!STERACEAE !RTEMISIA VULGARIS "ARNEY AND
$I4OMMASO
!STERACEAE #ARDUUS ACANTHOIDES #AVERS
!STERACEAE #ARDUUS NUTANS #AVERS
!STERACEAE #ENTAUREA DIFFUSA -ULLIGAN
!STERACEAE #ENTAUREA MACULOSA -ULLIGAN
!STERACEAE #IRSIUM ARVENSE -ULLIGAN
!STERACEAE #REPIS TECTORUM -ULLIGAN
!STERACEAE 'ALINSOGA PARVImORA -ULLIGAN
!STERACEAE 'ALINSOGA QUADRIRADIATA -ULLIGAN
!STERACEAE (ELIANTHUS TUBEROSUS #AVERS
!STERACEAE (YPOCHAERIS RADICATA -ULLIGAN
!STERACEAE ,ACTUCA SERRIOLA 7EAVER AND $OWNS
!STERACEAE -ATRICARIA PERFORATA #AVERS
!STERACEAE 3ENECIO JACOBAEA #AVERS
!STERACEAE 3ENECIO VULGARIS 2OBINSON ET AL
!STERACEAE 3ONCHUS ARVENSIS #AVERS
!STERACEAE 3ONCHUS ASPER #AVERS
!STERACEAE 3ONCHUS OLERACEUS #AVERS
!STERACEAE 4ARAXACUM OFlCINALE 3TEWART
7ADE ET AL
!STERACEAE 4RAGOPOGON DUBIUS #LEMENTS ET AL
!STERACEAE 4RAGOPOGON PORRIFOLIUS #LEMENTS ET AL
!STERACEAE 4RAGOPOGON PRATENSIS #LEMENTS ET AL
"ORAGINACEAE #YNOGLOSSUM OFlCINALE #AVERS
"ORAGINACEAE %CHIUM VULGARE +LEMOW ET AL
"RASSICACEAE !LLIARIA PETIOLATA -ULLIGAN
"RASSICACEAE "ARBAREA VULGARIS #AVERS
"RASSICACEAE #ARDARIA DRABA -ULLIGAN
(4 -URPHY ET AL
!PPENDIX #ONTINUED
"#7 &AMILY 3PECIES 2EFERENCE
SERIES
"RASSICACEAE $ESCURAINIA SOPHIA -ULLIGAN
"RASSICACEAE %RUCASTRUM GALLICUM 7ARWICK AND 7ALL
"RASSICACEAE .ESLIA PANICULATA &RANCIS AND 7ARWICK
"RASSICACEAE 3INAPIS ARVENSIS -ULLIGAN
7ARWICK ET AL
"RASSICACEAE 4HLASPI ARVENSE -ULLIGAN
7ARWICK ET AL
#ANNABACEAE #ANNABIS SATIVA 3MALL AND #AVERS
#ARYOPHYLLACEAE 'YPSOPHILA PANICULATA -ULLIGAN
#ARYOPHYLLACEAE 3ILENE LATIFOLIA SSP ALBA -ULLIGAN
#ARYOPHYLLACEAE 3ILENE NOCTImORA -ULLIGAN
#ARYOPHYLLACEAE 3TELLARIA MEDIA -ULLIGAN
#HENOPODIACEAE #HENOPODIUM ALBUM -ULLIGAN
#LUSIACEAE (YPERICUM PERFORATUM #AVERS
#ONVOLVULACEAE #ONVOLVULUS ARVENSIS -ULLIGAN
$IPSACACEAE $IPSACUS FULLONUM
SSP SYLVESTRIS -ULLIGAN
%UPHORBIACEAE %UPHORBIA CYPARISSIAS #AVERS
%UPHORBIACEAE %UPHORBIA ESULA -ULLIGAN
&ABACEAE ,OTUS CORNICULATUS -ULLIGAN
&ABACEAE -EDICAGO LUPULINA -ULLIGAN
&ABACEAE -ELILOTUS ALBA -ULLIGAN
&ABACEAE -ELILOTUS OFlCINALIS -ULLIGAN
&ABACEAE 4RIFOLIUM REPENS -ULLIGAN
&ABACEAE 6ICIA CRACCA #AVERS
&ABACEAE 6ICIA SATIVA #AVERS
&ABACEAE 6ICIA TETRASPERMA #AVERS
&ABACEAE 6ICIA VILLOSA #AVERS
(ALORAGACEAE -YRIOPHYLLUM SPICATUM -ULLIGAN
,AMIACEAE 'ALEOPSIS TETRAHIT #AVERS
,YTHRACEAE ,YTHRUM SALICARIA #AVERS
-ALVACEAE !BUTILON THEOPHRASTI #AVERS
/XALIDACEAE /XALIS CORNICULATA #AVERS
0LANTAGINACEAE 0LANTAGO LANCEOLATA -ULLIGAN
0LANTAGINACEAE 0LANTAGO MAJOR -ULLIGAN
0OACEAE !PERA SPICA
VENTI #AVERS
0OACEAE !VENA FATUA -ULLIGAN
0OACEAE "ROMUS TECTORUM #AVERS
0OACEAE %CHINOCHLOA CRUSGALLI #AVERS
)NVASIVENESS IN EXOTIC PLANTS
0OACEAE (OLCUS LANATUS #AVERS
0OACEAE (ORDEUM JUBATUM -ULLIGAN
0OACEAE 0OA ANNUA -ULLIGAN
0OACEAE 3ETARIA VERTICILLATA -ULLIGAN
0OACEAE 3ETARIA VIRIDIS #AVERS
0OACEAE 3ORGHUM HALEPENSE -ULLIGAN
0OLYGONACEAE &AGOPYRUM TATARICUM #AVERS
0OLYGONACEAE 0OLYGONUM CONVOLVULUS -ULLIGAN
0ORTULACACEAE 0ORTULACA OLERACEA -ULLIGAN
0OTAMOGETONACEAE 0OTAMOGETON CRISPUS #AVERS
2ANUNCULACEAE 2ANUNCULUS REPENS #AVERS
2OSACEAE 0OTENTILLA ARGENTEA -ULLIGAN
2OSACEAE 0OTENTILLA RECTA -ULLIGAN
2UBIACEAE 'ALIUM MOLLUGO -ERSEREAU AND
$I4OMMASO
2UBIACEAE 'ALIUM SPURIUM #AVERS
3CROPHULARIACEAE ,INARIA DALMATICA 6UJNOVIC AND 7EIN
3CROPHULARIACEAE ,INARIA VULGARIS 3ANER ET AL
3CROPHULARIACEAE 6ERBASCUM THAPSUS -ULLIGAN
3OLANACEAE $ATURA STRAMONIUM #AVERS
3OLANACEAE 3OLANUM CAROLINENSE #AVERS
3OLANACEAE 3OLANUM NIGRUM #AVERS
3OLANACEAE 3OLANUM ROSTRATUM #AVERS
A 6IOLACEAE 6IOLA ARVENSIS #AVERS
mORA DATASETS IN ORDER TO DEVELOP GENERAL PRINCIPLES THAT WILL ALLOW CARRYING OUT
MORE ACCURATE RISK ASSESSMENT OF CASUAL INTRODUCED POPULATIONS OF SPECIES BEFORE
THEY NATURALIZE AND SPREAD TO BECOME SERIOUS AND COSTLY INVASIVE POPULATIONS
!#+./7,%$'-%.43
(4- WAS SUPPORTED BY 3(!2#.%4 *6$7 *,$ AND ,,$ ARE SUPPORTED BY .ATURAL
3CIENCES AND %NGINEERING 2ESEARCH #OUNCIL OF #ANADA 7E THANK 2OB #OLAUTTI
AN ANONYMOUS REVIEWER AND THE EDITORS FOR COMMENTS THAT GREATLY IMPROVED THE
MANUSCRIPT
(4 -URPHY ET AL
2%&%2%.#%3
!KAKAYA ( 2 AND 0 3JGREN 'ULVE 0OPULATION VIABILITY ANALYSIS IN CONSERVATION
PLANNING !N OVERVIEW %COLOGICAL "ULLETIN
!NDERSON 0 + ! ! #UNNINGHAM . ' 0ATEL & * -ORALES 0 2 %PSTEIN AND 0 $ASZAK
%MERGING INFECTIOUS DISEASES OF PLANTS PATHOGEN POLLUTION CLIMATE CHANGE AND
AGROTECHNOLOGY DRIVERS 4RENDS IN %COLOGY AND %VOLUTION
"AKER ( ' #HARACTERISTICS AND MODE OF ORIGIN OF WEEDS PP IN ( ' "AKER
AND ' , 3TEBBINS EDITORS 4HE GENETICS OF COLONIZING SPECIES !CADEMIC 0RESS .EW
9ORK .9
"AKER ( ' 4HE EVOLUTION OF WEEDS !NNUAL 2EVIEW OF %COLOGY AND 3YSTEMATICS
"ARNEY * . AND ! $I4OMMASO 4HE "IOLOGY OF #ANADIAN 7EEDS !RTEMISIA
VULGARIS , #ANADIAN *OURNAL OF 0LANT 3CIENCE
"LACKBURN 4 - AND 2 0 $UNCAN A $ETERMINANTS OF ESTABLISHMENT SUCCESS IN INTRO
DUCED BIRDS .ATURE
"LACKBURN 4 - AND 2 0 $UNCAN B %STABLISHMENT PATTERNS OF EXOTIC BIRDS ARE
CONSTRAINED BY NON RANDOM PATTERNS IN INTRODUCTION *OURNAL OF "IOGEOGRAPHY
"ROOK " 7 * * /'RADY ! 0 #HAPMAN - ! "URGMAN ( 2 !KAKAYA AND 2 &RANKHAM
0REDICTIVE ACCURACY OF POPULATION VIABILITY ANALYSIS IN CONSERVATION BIOLOGY
.ATURE
"RUNO * & * * 3TACHOWICZ AND - $ "ERTNESS )NCLUSION OF FACILITATION INTO ECOLOGI
CAL THEORY 4RENDS IN %COLOGY AND %VOLUTION
#ADOTTE - 7 AND * ,OVETT $OUST %COLOGICAL AND TAXONOMIC DIFFERENCES BETWEEN
NATIVE AND INTRODUCED PLANTS OF SOUTHWESTERN /NTARIO %COSCIENCE
#ADOTTE - 7 " 2 -URRAY AND * ,OVETT $OUST %VOLUTIONARY AND ECOLOGICAL FACTORS INmU
ENCING /NTARIO PLANT INVADER SUCCESS %COSCIENCE IN PRESS
#ADOTTE - 7 " 2 -URRAY AND * ,OVETT $OUST %COLOGICAL PATTERNS AND BIOLOGICAL
INVASIONS 5SING REGIONAL SPECIES INVENTORIES IN MACROECOLOGY "IOLOGICAL )NVASIONS
#ALLAWAY 2 - AND % 4 !SCHEHOUG )NVASIVE PLANTS VERSUS THEIR NEW AND OLD
NEIGHBORS ! MECHANISM FOR EXOTIC INVASION 3CIENCE
#ARR ' 7 %XOTIC mORA OF 6ICTORIA AND ITS IMPACT ON INDIGENOUS BIOTA PP
IN $ " &OREMAN AND . ' 7ALSH EDITORS &LORA OF 6ICTORIA )NKATA 0RESS -ELBOURNE
!USTRALIA
#ASE 4 * )NVASION RESISTANCE ARISES IN STRONGLY INTERACTING SPECIES RICH MODEL
COMPETITION COMMUNITIES 0ROCEEDINGS OF THE .ATIONAL !CADEMY OF 3CIENCE
#AVERS 0 " EDITOR 4HE "IOLOGY OF #ANADIAN 7EEDS 6OL #ONTRIBUTIONS
4HE !GRICULTURAL )NSTITUTE OF #ANADA /TTAWA /.
#AVERS 0 " EDITOR 4HE "IOLOGY OF #ANADIAN 7EEDS 6OL #ONTRIBUTIONS
4HE !GRICULTURAL )NSTITUTE OF #ANADA /TTAWA /.
)NVASIVENESS IN EXOTIC PLANTS
#AVERS " 0 AND 3 7ARWICK 4HE "IOLOGY OF #ANADIAN 7EEDS 2EVISED FORMAT AND
INSTRUCTIONS FOR PREPARATION OF UPDATED ACCOUNTS #ANADIAN *OURNAL OF 0LANT 3CIENCE
#LEMENTS $ 2 - + 5PADHYAYA AND 3 * "OS 4HE "IOLOGY OF #ANADIAN 7EEDS
4RAGOPOGON DUBIUS 3COP 4RAGOPOGON PRATENSIS , AND 4RAGOPOGON PORRIFOLIUS ,
#ANADIAN *OURNAL OF 0LANT 3CIENCE
#OLAUTTI 2 ) AND ( * -AC)SAAC ! NEUTRAL TERMINOLOGY TO DElNE @INVASIVE SPECIES
$IVERSITY AND $ISTRIBUTIONS
$AEHLER # # 4HE TAXONOMIC DISTRIBUTION OF INVASIVE ANGIOSPERM PLANTS %COLOGICAL
INSIGHTS AND COMPARISON TO AGRICULTURAL WEEDS "IOLOGICAL #ONSERVATION
$AEHLER # # A 4WO WAYS TO BE AN INVADER BUT ONE IS MORE SUITABLE FOR ECOLOGY %3!
"ULLETIN
$AEHLER # # B $ARWINS NATURALIZATION HYPOTHESIS REVISITED !MERICAN .ATURALIST
$ARBYSHIRE 3 * 2 (OEG AND * (AVERKORT 4HE "IOLOGY OF #ANADIAN 7EEDS
!NTHRISCUS SYLVESTRIS , (OFFM #ANADIAN *OURNAL OF 0LANT 3CIENCE
$ARWIN # /N THE ORIGIN OF SPECIES -URRAY ,ONDON
$AVIS - ! AND + 4HOMPSON %IGHT WAYS TO BE A COLONIZER TWO WAYS TO BE AN INVAD
ER ! PROPOSED NOMENCLATURE SCHEME FOR INVASION ECOLOGY %3! "ULLETIN
$AVIS - ! AND + 4HOMPSON )NVASION TERMINOLOGY SHOULD ECOLOGISTS DElNE THEIR
TERMS DIFFERENTLY THAN OTHERS .O NOT IF WE WANT TO BE OF ANY HELP %3! "ULLETIN
$AVIS - ! AND + 4HOMPSON h.EWCOMERSv INVADE THE lELD OF INVASION ECOLOGY
QUESTION THE lELDS FUTURE %3! "ULLETIN
%CKERT ' # $ -ANICACCI AND 3 # ( "ARRETT 'ENETIC DRIFT AND FOUNDER EFFECT
IN NATIVE VERSUS INTRODUCED POPULATIONS OF AN INVADING PLANT ,YTHRUM SALICARIA
,YTHRACEAE %VOLUTION
%LTON # 3 4HE ECOLOGY OF INVASIONS BY ANIMALS AND PLANTS -ETHUEN ,ONDON
&ORSYTH $ - AND 2 0 $UNCAN 0ROPAGULE SIZE AND THE RELATIVE SUCCESS OF
EXOTIC UNGULATE AND BIRD INTRODUCTIONS TO .EW :EALAND 4HE !MERICAN .ATURALIST
&OXCROFT , # - 2OUGET $ - 2ICHARDSON AND 3 -AC &AYDEN 2ECONSTRUCTING
YEARS OF /PUNTIA STRICTA INVASION IN THE +RUGER .ATIONAL 0ARK 3OUTH !FRICA ENVI
RONMENTAL DETERMINANTS AND PROPAGULE PRESSURE $IVERSITY AND $ISTRIBUTIONS
&RANCIS ! AND 3 ) 7ARWICK 4HE "IOLOGY OF #ANADIAN 7EEDS .ESLIA PANICU
LATA , $ESV #ANADIAN *OURNAL OF 0LANT 3CIENCE
&RECKLETON 2 0 AND 7ATKINSON ! 2 ,ARGE
SCALE SPATIAL DYNAMICS OF PLANTS
METAPOPULATIONS REGIONAL ENSEMBLES AND PATCHY POPULATIONS *OURNAL OF %COLOGY
'LEASON ( ! 4HE NEW "RITTON AND "ROWN ILLUSTRATED mORA OF THE .ORTHEASTERN
5NITED 3TATES AND ADJACENT #ANADA (AFNER 0RESS .EW 9ORK
'LEASON ( ! AND ! #RONQUIST -ANUAL OF VASCULAR PLANTS OF NORTHEASTERN 5NITED
3TATES AND ADJACENT #ANADA 3ECOND %DITION .EW 9ORK "OTANICAL 'ARDEN .EW 9ORK .9
(4 -URPHY ET AL
'OTELLI . AND # - 4AYLOR A 4ESTING METAPOPULATION MODELS WITH STREAM
lSH
ASSEMBLAGES %VOLUTIONARY %COLOGY 2ESEARCH
'OTELLI . AND # - 4AYLOR B 4ESTING MACROECOLOGY MODELS WITH STREAM
lSH ASSEM
BLAGES %VOLUTIONARY %COLOGY 2ESEARCH
(ANSKI ) 3INGLE
SPECIES METAPOPULATION DYNAMICS CONCEPTS MODELS AND OBSER
VATIONS "IOLOGICAL *OURNAL OF THE ,INNEAN 3OCIETY
(ANSKI ) ! PRACTICAL MODEL OF METAPOPULATION DYNAMICS *OURNAL OF !NIMAL %COLOGY
(ARROD 2 * AND 2 * 4AYLOR 2EPRODUCTION AND POLLINATION BIOLOGY OF #ENTAUREA
AND !CROPTILON SPECIES WITH EMPHASIS ON # DIFFUSA .ORTHWEST 3CIENCE
(IGGINS 3 ) 2 .ATHAN AND - , #AIN !RE LONG
DISTANCE DISPERSAL EVENTS IN PLANTS
USUALLY CAUSED BY NONSTANDARD MEANS OF DISPERSAL %COLOGY
+AREIVA 0 AND 7ENNERGREN 5 #ONNECTING LANDSCAPE PATTERNS TO ECOSYSTEM AND
POPULATION PROCESSES .ATURE
+EANE 2 - AND - * #RAWLEY %XOTIC PLANT INVASIONS AND THE ENEMY RELEASE
HYPOTHESIS 4RENDS IN %COLOGY AND %VOLUTION
+LEMOW + - $ 2 #LEMENTS 0 & 4HREADGILL AND 0 " #AVERS 4HE "IOLOGY
OF #ANADIAN 7EEDS %CHIUM VULGARE , #ANADIAN *OURNAL OF 0LANT 3CIENCE
+LIRONOMOS * . &EEDBACK WITH SOIL BIOTA CONTRIBUTES TO PLANT RARITY AND INVASIVE
NESS IN COMMUNITIES .ATURE
+OLAR 3 # AND $ - ,ODGE 0ROGRESS IN INVASION BIOLOGY PREDICTING INVADERS
4RENDS IN %COLOGY AND %VOLUTION
+OLBE * * % % 'LOR , 2 3CHETTINO ! # ,ARA ! ,ARSON AND * " ,OSOS 'ENETIC
VARIATION INCREASES DURING BIOLOGICAL INVASION BY A #UBAN LIZARD .ATURE
+OWARIK ) #LONAL GROWTH IN !ILANTHUS ALTISSIMA ON A NATURAL SITE IN 7EST 6IRGINIA
*OURNAL OF 6EGETATION 3CIENCE
+REBS # * %COLOGY 4HE EXPERIMENTAL ANALYSIS OF DISTRIBUTION AND ABUNDANCE
(ARPER 2OW .EW 9ORK .9
,EVIN $ ! %COLOGICAL SPECIATION LESSONS FROM INVASIVE SPECIES 3YSTEMATIC "OTANY
,EVINE * - 0 " !DLER AND 3 ' 9ELENIK ! META
ANALYSIS OF BIOTIC RESISTANCE TO
EXOTIC PLANT INVASIONS %COLOGY ,ETTERS
,EVINS 2 3OME DEMOGRAPHIC AND GENETIC CONSEQUENCES OF ENVIRONMENTAL HETERO
GENEITY FOR BIOLOGICAL CONTROL "ULLETIN OF THE %NTOMOLOGICAL 3OCIETY OF !MERICA
,ODGE $ - "IOLOGICAL INVASIONS LESSONS FOR ECOLOGY 4RENDS IN %COLOGY AND
%VOLUTION
,ONSDALE - 7 'LOBAL PATTERNS OF PLANT INVASIONS AND THE CONCEPT OF INVASIBILITY
%COLOGY
-ACK 2 . 0REDICTING THE IDENTITY AND THE FATE OF PLANT INVADERS EMERGENT AND
EMERGING APPROACHES "IOLOGICAL #ONSERVATION
-ACK 2 . $ 3IMBERLOFF 7 - ,ONSDALE ( %VANS - #LOUT AND & ! "AZZAZ
"IOTIC INVASIONS CAUSES EPIDEMIOLOGY GLOBAL CONSEQUENCES AND CONTROL %COLOGICAL
!PPLICATIONS
)NVASIVENESS IN EXOTIC PLANTS
-AL 4 + ,OVETT
$OUST * , ,OVETT
$OUST AND ' ! -ULLIGAN 4HE "IOLOGY OF
#ANADIAN 7EEDS ,YTHRUM SALICARIA #ANADIAN *OURNAL OF 0LANT 3CIENCE
-ARON * , AND 6IL - 7HEN DO HERBIVORES AFFECT PLANT INVASION %VIDENCE FOR THE
NATURAL ENEMIES AND BIOTIC RESISTANCE HYPOTHESES /IKOS
-ASTER , , !SSESSING THREATS AND SETTING PRIORITIES FOR CONSERVATION #ONSERVATION
"IOLOGY
-C.EILL * 4HE "IOLOGY OF #ANADIAN 7EEDS 3ILENE ALBA -ILLER % ( , +RAUSE
#ANADIAN *OURNAL OF 0LANT 3CIENCE
-ERSEREAU $ AND ! $I4OMMASO 4HE "IOLOGY OF #ANADIAN 7EEDS 'ALIUM
MOLLUGO , #ANADIAN *OURNAL OF 0LANT 3CIENCE
-ITCHELL # % AND ! ' 0OWER 2ELEASE OF INVASIVE PLANTS FROM FUNGAL AND VIRAL
PATHOGENS .ATURE
-ORRIS 7 & AND $ & $OAK 1UANTITATIVE CONSERVATION BIOLOGY THEORY AND PRACTICE
OF POPULATION VIABILITY ANALYSIS 3INAUER !SSOCIATES 3UNDERLAND -!
-OILANEN ! 0ATCH OCCUPANCY MODELS OF METAPOPULATION DYNAMICS %FlCIENT
PARAMETER ESTIMATION USING IMPLICIT STATISTICAL INFERENCE %COLOGY
-OILANEN ! 4HE EQUILIBRIUM ASSUMPTION IN ESTIMATING THE PARAMETERS OF META
POPULATION MODELS *OURNAL OF !NIMAL %COLOGY
-OILANEN ! ! 4 3MITH AND ) (ANSKI ,ONG
TERM DYNAMICS IN A METAPOPULATION
OF THE !MERICAN PIKA !MERICAN .ATURALIST
-ORTON * + AND * - 6ENN ! CHECKLIST OF THE mORA OF /NTARIO 6ASCULAR PLANTS
5NIVERSITY OF 7ATERLOO "IOLOGY 3ERIES .O 7ATERLOO /.
-ULLIGAN ' ! EDITOR 4HE "IOLOGY OF #ANADIAN 7EEDS #ONTRIBUTIONS
!GRICULTURE #ANADA 0UBL /TTAWA /.
-ULLIGAN ' ! EDITOR 4HE "IOLOGY OF #ANADIAN 7EEDS #ONTRIBUTIONS
!GRICULTURE #ANADA 0UBL /TTAWA /.
-URPHY ( 4 AND * ,OVETT
$OUST #ONTEXT AND CONNECTIVITY IN PLANT METAPOPULA
TIONS AND LANDSCAPE MOSAICS DOES THE MATRIX MATTER /IKOS
.ATURAL (ERITAGE )NFORMATION #ENTRE $ATABASE OF VASCULAR PLANTS HTTPWWW
MNR GOV ON CA-.2NHICNHIC CFM
.EWMASTER 3 ' ! ,EHELA - * /LDHAM 0 7 # 5HLIG AND 3 -C-URRAY
/NTARIO PLANT LIST &OREST 2ESEARCH )NFORMATION 0APER .O /NTARIO &OREST 2ESEARCH
)NSTITUTE 3AULT 3TE -ARIE /.
.OVAK 3 * AND 2 . -ACK 'ENETIC VARIATION IN "ROMUS TECTORUM 0OACEAE COM
PARISON BETWEEN NATIVE AND INTRODUCED POPULATIONS (EREDITY
0YEK 0 )S THERE A TAXONOMIC PATTERN TO PLANT INVASIONS /IKOS
0YEK 0 $ - 2ICHARDSON - 2EJMNEK ' , 7EBSTER - 7ILLIAMSON AND * +IRSCHNER
!LIEN PLANTS IN CHECKLISTS AND mORAS TOWARDS BETTER COMMUNICATION BETWEEN
TAXONOMISTS AND ECOLOGISTS 4AXON
2ABINOWITZ $ 3EVEN FORMS OF RARITY PP
IN ( 3YNGE EDITOR 4HE BIOLOGICAL
ASPECTS OF RARE PLANT CONSERVATION 7ILEY .EW 9ORK .9
2EINHART + / ! 0ACKER 7 ( 6AN DER 0UTTEN AND + #LAY 0LANT
SOIL BIOTA INTER
ACTIONS AND SPATIAL DISTRIBUTION OF BLACK CHERRY IN ITS NATIVE AND INVASIVE RANGES %COLOGY
,ETTERS
(4 -URPHY ET AL
2EJMNEK - ! THEORY OF SEED PLANT INVASIVENESS THE lRST SKETCH "IOLOGICAL
#ONSERVATION
2EJMNEK - AND $ - 2ICHARDSON 7HAT ATTRIBUTES MAKE SOME PLANT SPECIES MORE
INVASIVE %COLOGY
2EJMNEK - $ - 2ICHARDSON - ' "ARBOUR - * #RAWLEY ' & (RUSA 0 " -OYLE
* - 2ANDAL $ 3IMBERLOFF AND - 7ILLIAMSON "IOLOGICAL INVASIONS POLITICS AND
THE DISCONTINUITY OF THE ECOLOGICAL TERMINOLOGY %3! "ULLETIN
2ICHARDSON $ - 0 0YEK - 2EJMNEK - ' "ARBOUR & $ 0ANETTA AND # * 7EST
A .ATURALIZATION AND INVASION OF ALIEN PLANTS CONCEPTS AND DElNITIONS $IVERSITY
AND $ISTRIBUTIONS
2ICHARDSON $ - . !LLSOPP # - $!NTONIO 3 * -ILTON AND - 2EJMNEK B
0LANT INVASIONS THE ROLE OF MUTUALISMS "IOLOGICAL 2EVIEWS
2OBINSON $ % * 4 /$ONOVAN - 0 3HARMA $ * $OOHAN AND 2 &IGUEROA 4HE
"IOLOGY OF #ANADIAN 7EEDS 3ENECIO VULGARIS , #ANADIAN *OURNAL OF 0LANT 3CIENCE
2OUGET - AND $ - 2ICHARDSON )NFERRING PROCESS FROM PATTERN IN PLANT INVASIONS
A SEMI
MECHANISTIC MODEL INCORPORATING PROPAGULE PRESSURE AND ENVIRONMENTAL FACTORS
!MERICAN .ATURALIST
3ANER - ! $ 2 #LEMENTS - 2 (ALL $ * $OOHAN AND # 7 #ROMPTON 4HE
"IOLOGY OF #ANADIAN 7EEDS ,INARIA VULGARIS -ILL #ANADIAN *OURNAL OF 0LANT
3CIENCE
3AX $ & AND * ( "ROWN 4HE PARADOX OF INVASION 'LOBAL %COLOGY AND "IOGEOGRAPHY
3HAFFER - , -INIMUM POPULATION SIZES FOR SPECIES CONSERVATION "IO3CIENCE
3COGGAN ( * 4HE mORA OF #ANADA .ATIONAL -USEUMS OF #ANADA /TTAWA /.
3IMBERLOFF $ AND " VON (OLLE 0OSITIVE INTERACTIONS OF NON
INDIGENOUS SPECIES
INVASIONAL MELTDOWN "IOLOGICAL )NVASIONS
3MALL % AND 0 " #AVERS 4HE "IOLOGY OF #ANADIAN 7EEDS #ANNABIS SATIVA ,
#ANADIAN *OURNAL OF 0LANT 3CIENCE
3TEWART
7ADE 3 - 3 .EUMANN , , #OLLINS AND ' * "OLAND 4HE "IOLOGY OF
#ANADIAN 7EEDS 4ARAXACUM OFlCINALE ' ( 7EBER EX 7IGGERS #ANADIAN *OURNAL
OF 0LANT 3CIENCE
3TOHLGREN 4 * $ "INKLEY ' 7 #HONG - ! +ALKHAN , $ 3CHELL + ! "ULL 9 /TSUKI
' .EWMAN - "ASHKIN AND 9 3ON %XOTIC PLANT SPECIES INVADE HOT SPOTS OF
NATIVE PLANT DIVERSITY %COLOGICAL -ONOGRAPHS
4HOMPSON + * ' (ODGSON AND 4 # ' 2ICH .ATIVE AND ALIEN INVASIVE PLANTS
MORE OF THE SAME %COGRAPHY
4ILMAN $ #OMMUNITY INVASIBILITY RECRUITMENT LIMITATION AND GRASSLAND BIODIVER
SITY %COLOGY
4UTIN 4 ' 6 ( (EYWOOD . ! "URGESS $ ( 6ALENTINE 3 - 7ALTERS AND $ ! 7EBB
&LORA %UROPAEA 6OL ,YCOPODIACEAE TO 0LATANACEAE #AMBRIDGE 5NIVERSITY
0RESS #AMBRIDGE 5+
)NVASIVENESS IN EXOTIC PLANTS
4URKINGTON 2 ! 0 " #AVERS AND % 2EMPEL 4HE "IOLOGY OF #ANADIAN 7EEDS
-ELILOTUS ALBA $ESR AND - OFlCINALIS , ,AM #ANADIAN *OURNAL OF 0LANT 3CIENCE
6ANDER 7ALL 3 " AND 7 3 ,ONGLAND $IPLOCHORY ARE TWO SEED DISPERSERS BETTER
THAN ONE 4RENDS IN %COLOGY AND %VOLUTION
6ITOUSEK 0 - AND , 2 7ALKER "IOLOGICAL INVASION BY -YRICA FAYA IN (AWAII
0LANT DEMOGRAPHY NITROGEN lXATION ECOSYSTEM EFFECTS %COLOGICAL -ONOGRAPHS
6ON (OLLE " ( 2 $ELCOURT AND $ 3IMBERLOFF 4HE IMPORTANCE OF BIOLOGICAL INERTIA
IN PLANT COMMUNITY RESISTANCE TO INVASION *OURNAL OF 6EGETATION 3CIENCE
6UJNOVIC + AND 2 7 7EIN 4HE "IOLOGY OF #ANADIAN 7EEDS ,INARIA DAL
MATICA , -ILL #ANADIAN *OURNAL OF 0LANT 3CIENCE
7AHLBERG . ! -OILANEN AND ) (ANSKI 0REDICTING THE OCCURRENCE OF ENDANGERED
SPECIES IN FRAGMENTED LANDSCAPES 3CIENCE
7ARWICK 3 ) AND , $ "LACK 4HE "IOLOGY OF #ANADIAN 7EEDS 3ORGHUM HALA
PENSE , 0ERS #ANADIAN *OURNAL OF 0LANT 3CIENCE
7ARWICK 3 ) ( * "ECKIE ' ! 4HOMAS AND 4 -C$ONALD 4HE "IOLOGY OF
#ANADIAN 7EEDS 3INAPSIS ARVENSIS , UPDATED #ANADIAN *OURNAL OF 0LANT 3CIENCE
7ARWICK 3 ) AND $ ! 7ALL 4HE "IOLOGY OF #ANADIAN 7EEDS %RUCASTRUM
GALLICUM 7ILLD / % 3CHULZ #ANADIAN *OURNAL OF 0LANT 3CIENCE
7ARWICK 3 ) ! &RANCIS AND $ * 3USKO 4HE "IOLOGY OF #ANADIAN 7EEDS
4HLASPI ARVENSE , UPDATED #ANADIAN *OURNAL OF 0LANT 3CIENCE
7EAVER 3 % AND - 0 $OWNS 4HE "IOLOGY OF #ANADIAN 7EEDS ,ACTUCA SER
RIOLA , #ANADIAN *OURNAL OF 0LANT 3CIENCE
7ILLIAMSON - AND ! &ITTER 4HE VARYING SUCCESS OF INVADERS %COLOGY
#HAPTER FIVE
$ENSITY DEPENDENCE IN
INVASIVE PLANTS DEMOGRAPHY
HERBIVORY SPREAD AND
EVOLUTION
9 - "UCKLEY AND * -ETCALF
).42/$5#4)/.
"ECAUSE OF THEIR ENVIRONMENTAL AND ECONOMIC COSTS 0IMENTEL ET AL SOME
INVASIVE PLANT SPECIES ARE WELL STUDIED OVER RELATIVELY LONG TIME SCALES "UCKLEY
ET AL A AND IN DIFFERENT HABITATS 0AYNTER ET AL 4HESE DATA ARE VALU
ABLE FOR THE DEVELOPMENT OR TESTING OF GENERAL THEORETICAL MODELS OF POPULATION
DYNAMICS AND SPECIES INTERACTIONS &URTHERMORE KNOWING THE KIND OF DYNAMICS
EXHIBITED BY INVASIVES MATTERS IF WE ARE ATTEMPTING TO PREDICT THEIR IMPACT ON AN
ECOSYSTEM AND TO ANTICIPATE HOW THEY WILL RESPOND TO NOVEL ENVIRONMENTS OVER
ECOLOGICAL AND EVOLUTIONARY TIME
SCALES 'ENERAL RECENT REVIEWS OF DENSITY DEPEN
DENCE CAN BE FOUND IN 4URCHIN +REBS 3IBLY AND (ONE
HERE WE CONCENTRATE ON HOW RECENT ADVANCES IN OUR UNDERSTANDING OF HOW
DENSITY DEPENDENT PROCESSES CONTRIBUTE TO OUR UNDERSTANDING OF THE DEMOGRAPHY
MANAGEMENT AND EVOLUTION OF INVASIVE PLANT SPECIES
$ESPITE CONSIDERABLE CONTROVERSY STRETCHING BACK TO DEBATES BETWEEN .ICHOLSON
AND !NDREWARTHA OVER YEARS AGO AND CONTINUING TO SOME EXTENT TODAY
-7 #ADOTTE ET AL (EDS) #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY
3PRINGER 0RINTED IN THE .ETHERLANDS
9- "UCKLEY AND * -ETCALF
A COMBINATION OF ENDOGENOUS AND EXOGENOUS FORCES IS GENERALLY ACCEPTED TO
REGULATE POPULATION SIZE 4URCHIN %NDOGENOUS MECHANISMS ARE USUALLY
REFERRED TO AS hDENSITY DEPENDENCEv WHICH IS A SHORTHAND TERM ENCOMPASSING
THE RANGE OF MECHANISMS THAT MODIFY VITAL RATES ACCORDING TO THE DENSITY OF CON
SPECIlCS 4HESE ARE THEORETICALLY OF GREAT IMPORTANCE AS A NECESSARY CONDITION FOR
POPULATION REGULATION BUT ARE OFTEN DIFlCULT TO DETECT CONSISTENTLY UNDER NATURAL
CONDITIONS SEE EXAMPLES IN +REBS "Y CONTRAST EXOGENOUS MECHANISMS
INCORPORATE EXTERNAL FACTORS SUCH AS RESOURCE AVAILABILITY HERBIVORY OR HARVEST
ING %XOGENOUS AND ENDOGENOUS FACTORS ARE INTIMATELY LINKED &OR EXAMPLE THE
STRENGTH OF EXOGENOUS FACTORS SUCH AS RESOURCE AVAILABILITY CAN DETERMINE THE
EQUILIBRIUM OF A SIMPLE DENSITY DEPENDENT MODEL 4HE WELL KNOWN (ASSELL MODEL
(ASSELL RELATES POPULATION SIZE IN YEAR T .T TO POPULATION SIZE IN THE
PREVIOUS YEAR .T MODIlED BY A DENSITY DEPENDENT FUNCTION INCORPORATING MAXI
MUM FECUNDITY AT LOW DENSITIES h A MEASURE OF HABITAT CARRYING CAPACITY A AND
THE STRENGTH AND FORM OF COMPETITION B !LL THREE PARAMETERS ARE POSITIVE
h
.T .T
A.T B
4HE EQUILIBRIUM POPULATION SIZE DEPENDS ON ALL THREE PARAMETERS
hB n
.
A
BUT STABILITY ONLY DEPENDS ON THE VALUES OF h AND B STABILITY ONLY BEING POSSIBLE
PROVIDING
B n hnB
#HANGING THE CARRYING CAPACITY OF THE ENVIRONMENT CHANGING A IN THE (ASSELL
MODEL WILL THEREFORE LEAD TO CHANGES IN THE EQUILIBRIUM DENSITY OF THE POPULATION
&IG 4HIS IS PARTICULARLY RELEVANT FOR INVASIVE SPECIES TAKEN FROM THEIR NATIVE
RANGE TO A NOVEL ENVIRONMENT OFTEN WITHOUT A SUITE OF SPECIALIST HERBIVORES AND
PATHOGENS 4HE ENEMY RELEASE HYPOTHESIS %2( OR NATURAL ENEMIES HYPOTHESIS
+EANE AND #RAWLEY POSITS THAT INVADERS ARE PARTICULARLY SUCCESSFUL IN THEIR
EXOTIC HABITAT BECAUSE THEY ARE RELEASED FROM REGULATION BY SPECIALIST HERBIVORES
PATHOGENS OR PARASITES PRESENT IN THEIR NATIVE HABITAT -OST PREDICTIONS STEMMING
FROM THE %2( RELATE TO INCREASED VIGOUR BE IT PHENOTYPIC OR GENOTYPIC EXHIBITED
BY INDIVIDUALS (OWEVER IF RESOURCES ARE NOT LIMITING IN EITHER EXOTIC OR NATIVE
RANGES AND ENEMY RELEASE IS OPERATING WE MIGHT EXPECT POPULATIONS OF INVASIVES
IN THE EXOTIC RANGE TO HAVE A HIGHER CARRYING CAPACITY AND EXHIBIT HIGHER EQUI
LIBRIUM DENSITIES 4HIS APPEARS TO BE THE CASE FOR SCOTCH BROOM #YTISUS SCOPARIUS
$ENSITY DEPENDENCE IN INVASIVE PLANTS
&IG (ASSELL MODEL AT DIFFERENT CARRYING CAPACITIES EQUILIBRIUM POPULATION DENSITIES
ACHIEVED OVER TIME YEARS ON THE X
AXIS WITH A SOLID LINE A DASHED LINE AND
A DOTTED LINE
WHICH WAS FOUND TO HAVE SIGNIlCANTLY HIGHER DENSITIES IN TWO COUNTRIES IN ITS EXOT
IC RANGE !USTRALIA AND .EW :EALAND COMPARED WITH ITS NATIVE RANGE %UROPE
0AYNTER ET AL 4HE ABSENCE OF SPECIALIST HERBIVORES OR PATHOGENS MAY ALSO
AFFECT THE COMPETITION PARAMETER B WHICH AT ITS EXTREME VALUES REPRESENTS IDEAL
SCRAMBLE AND IDEAL CONTEST COMPETITION #OMPETITION FOR ENEMY
FREE SPACE MAY
HAVE A DIFFERENT FORM FROM COMPETITION FOR SOIL RESOURCES AND DIFFERENT VALUES OF B
IN DIFFERENT ENVIRONMENTS WOULD CHANGE NOT JUST THE EQUILIBRIUM POPULATION DEN
SITIES BUT COULD POTENTIALLY LEAD TO POPULATIONS WITH DIFFERENT STABILITY PROPERTIES
4HIS VERY SIMPLE MODEL ILLUSTRATES HOW EXOGENOUS CHANGES TO CONDITIONS CAN LEAD
TO DIFFERENT DENSITY DEPENDENT DYNAMICS 4O OUR KNOWLEDGE A DETAILED STUDY OF
THE POPULATION DYNAMICS OF AN INVASIVE PLANT IN NATIVE AND EXOTIC HABITATS HAS
NOT BEEN CARRIED OUT IN ORDER TO INVESTIGATE HOW DENSITY DEPENDENT CONDITIONS
DIFFER BETWEEN HABITATS 7ITH THE USE OF OBSERVATIONS EXPERIMENTS AND MODELLING
AN INVASIVE PLANT SYSTEM WOULD BE IDEAL FOR INVESTIGATING THE INmUENCE OF DIFFERENT
ENVIRONMENTS ON DENSITY DEPENDENT PROCESSES
9- "UCKLEY AND * -ETCALF
4HE IMPORTANCE OF EXOGENOUS FACTORS FOR DENSITY DEPENDENT POPULATION REGULA
TION IS PARTICULARLY WELL ILLUSTRATED BY A RECENT ANALYSIS OF THE DYNAMICS OF RAGWORT
3ENECIO JACOBAEA , AND ITS SPECIALIST HERBIVORE THE CINNABAR MOTH 4YRIA JACOBAEAE
, "ONSALL ET AL ,ONG TERM * YEARS TIME SERIES FOR BOTH SPECIES EXIST
FOR TWO SITES ONE IN 3ILWOOD 5+ AND THE OTHER IN -EIJENDEL 4HE .ETHERLANDS
)N 3ILWOOD RAGWORT POPULATIONS ONLY DIRECT DENSITY DEPENDENCE IE THE INmUENCE
OF PLANT DENSITY IN THE PREVIOUS YEAR WAS DETECTED WHEREAS IN -EIJENDEL BOTH
DIRECT YEAR PREVIOUS AND DELAYED TWO YEARS PREVIOUS DENSITY DEPENDENCE WAS
DETECTED LEADING TO QUALITATIVELY DIFFERENT DYNAMICS AT THE TWO SITES 0OPULATIONS
ARE RELATIVELY STABLE IN 3ILWOOD BUT EXHIBIT OSCILLATIONS IN -EIJENDEL 4HE DRIVER
FOR THE DIFFERENCES IN DYNAMICS BETWEEN THE SITES MAY BE DIFFERENCES IN RECRUIT
MENT 2ECRUITMENT IS MICRO
SITE LIMITED AT 3ILWOOD BUT NOT AT -EIJENDEL IMPLYING
STRONG DENSITY DEPENDENCE OPERATING AT THE SEEDLING STAGE IN 3ILWOOD POPULATIONS
3ENECIO JACOBAEA IS NATIVE TO %UROPE AND INTRODUCED TO !USTRALASIA THE !MERICAS
AND 3OUTH !FRICA (ARPER AND 7OOD 4HIS STUDY IS A LANDMARK FOR ECOLO
GISTS WORKING ON INVASIVES AS THE VARIABILITY IN THE QUALITATIVE DYNAMICS OF THIS
SPECIES WITHIN ITS NATIVE RANGE ALONE SHOULD LEAD US TO ASK QUESTIONS OF ATTEMPTS
TO EXTRAPOLATE MODEL OR EXPERIMENTAL RESULTS FROM NATIVE TO EXOTIC ECOSYSTEMS IN
THE ABSENCE OF DATA FROM THE EXOTIC HABITAT )T SHOULD ALSO ENCOURAGE US TO MOVE
ON FROM DOCUMENTING TRAIT DIFFERENCES BETWEEN SPECIES IN THEIR NATIVE AND EXOTIC
RANGES 3IEMANN AND 2OGERS "UCKLEY ET AL B 3IEMANN AND 2OGERS
TO INVESTIGATING THE POPULATION DYNAMIC CONSEQUENCES OF THOSE TRAIT
DIFFERENCES AND THEIR INTERACTION WITH HABITAT DIFFERENCES )N ADDITION CONTEXT
DEPENDENCY OF THIS SORT MIGHT RESULT IN DENSITY DEPENDENCE BEING OVERLOOKED IF
THE SYSTEM STUDIED INCLUDES HABITAT HETEROGENEITY &OR EXAMPLE IN A STUDY OF REEF
lSH 3HIMA AND /SENBERG FOUND THAT THE STRENGTH OF DENSITY DEPENDENT
MORTALITY DEPENDED ON SITE QUALITY AND A FAILURE TO TAKE HETEROGENEITY IN SITE
QUALITY INTO ACCOUNT COULD HAVE LEAD TO THE CONCLUSION THAT DENSITY DEPENDENCE
WAS UNIMPORTANT 3HIMA /SENBERG REFER TO THE PATTERNS OBSERVED AS hCRYPTIC
DENSITY DEPENDENCEv
7ITHIN A POPULATION DIFFERENCES BETWEEN INDIVIDUALS OR STAGES WILL ALSO AFFECT
DENSITY DEPENDENCE AND RESULTING DYNAMICS &OR EXAMPLE POPULATION DENSITY
AND ASYMMETRIC COMPETITION CAN INCREASE THE DIFFERENTIAL IN SIZE BETWEEN LEVELS
IN HIERARCHIES OF PLANT POPULATIONS 7EINER AND THIS CAN CREATE DIFFER
ENCES BETWEEN INDIVIDUAL RESPONSES TO DIFFERENT DENSITIES OF DIFFERENT STAGES 3IMPLE
MODELS LIKE THE (ASSELL MODEL SHOWN ABOVE DO NOT ADDRESS THIS (OWEVER WE CAN
INCORPORATE SUCH DIFFERENCES BY USING MODELS WITH AGE OR STAGE STRUCTURE "ENTON
ET AL AND #AMERON AND "ENTON USE EXPERIMENTALLY MANIPULATED
SOIL MITE POPULATIONS AND MODELS TO EXPLORE THE INTERACTION OF DENSITY DEPENDENCE
AND ENVIRONMENTAL VARIABILITY ON THE OUTCOME OF HARVESTING DIFFERENT STAGES
4HEY FOUND THAT HARVESTING DIFFERENT STAGES REDUCING SURVIVAL OF EGGS JUVENILES
OR ADULTS HAD DIFFERENT IMPACTS ON THE MEAN POPULATION SIZE AND ITS VARIANCE IN
EACH STAGE )N A VARIABLE ENVIRONMENT THEY FOUND THAT REMOVAL OF OF EGGS
ACTUALLY RESULTED IN AN INCREASE IN MEAN POPULATION SIZE WHEREAS REMOVAL OF
$ENSITY DEPENDENCE IN INVASIVE PLANTS
CORRESPONDING PROPORTIONS OF JUVENILES AND ADULTS LED TO A DECREASE IN MEAN
POPULATION SIZE 4HIS WOULD APPEAR TO BE A RESULT OF DENSITY DEPENDENCE ACTING
DIFFERENTLY IN DIFFERENT STAGES 4HEY LIKEN REMOVAL OF EGGS TO THINNING SEEDLINGS
THIS RELEASES THE SURVIVORS FROM DENSITY DEPENDENT SUPPRESSION AND IF DENSITY
DEPENDENCE IS STRONG ENOUGH CAN LEAD TO AN INCREASE IN OVERALL POPULATION SIZE
4HE EFFECTS OF EGG REMOVAL MAY HAVE BEEN MORE PRONOUNCED IN THE VARIABLE
ENVIRONMENT BECAUSE OF THE ALTERNATING PERIODS OF STRONG AND WEAK COMPETITION
FOR RESOURCES 4HE SAME COUNTER
INTUITIVE RESULT HARVESTING LEADING TO INCREASED
POPULATION SIZE CAN OCCUR IN A CONSTANT ENVIRONMENT WHERE DENSITY DEPENDENCE
IS VERY STRONG OR OVER
COMPENSATING EG "UCKLEY ET AL /F RELEVANCE TO THE
MANAGEMENT OF INVASIVES IS "ENTON ET ALS CONCLUSION THAT DENSITY INDE
PENDENT MODELS FAILED TO PREDICT THE NEGATIVE ELASTICITY OF SURVIVAL AT THE EGG STAGE
THEREFORE MANAGEMENT MODELS WHICH DO NOT INCLUDE DENSITY DEPENDENCE WHERE IT
IS KNOWN TO BE IMPORTANT RUN THE RISK OF GIVING THE WRONG ANSWER
)NCORPORATING THE ADDED STRUCTURE FROM INDIVIDUAL VARIATION TO MODELS IS OFTEN
IMPORTANT WITH THIS ADDED COMPLEXITY HOWEVER GREATER CARE SHOULD BE TAKEN TO
CHOOSE THE APPROPRIATE MODEL &OX AND +ENDALL ILLUSTRATE THAT FAILURE TO
MODEL INDIVIDUAL VARIABILITY CORRECTLY AS SYSTEMATIC VARIATION AMONG INDIVIDUALS
RATHER THAN RANDOM VARIATION MAY LEAD TO INACCURATE PREDICTIONS 4HEY CONCLUDE
hALMOST ALL POPULATION VIABILITY ANALYSES 06!S OVER
ESTIMATE THE IMPORTANCE
OF DEMOGRAPHIC STOCHASTICITY AND THEREFORE THE RISK OF EXTINCTIONv -URRELL
#HAPTER PRESENTS A MODELLING FRAMEWORK THAT TAKES ACCOUNT OF LOCAL INTERAC
TIONS THE hPLANTS EYE VIEWv OF ITS NEIGHBOURHOOD DENSITY WHICH EVEN WITHOUT
SPATIAL OR TEMPORAL HETEROGENEITY LEADS TO INVASION DYNAMICS QUITE DIFFERENT FROM
THOSE PREDICTED BY MEAN lELD APPROXIMATIONS WHICH TAKE OVERALL DENSITY INTO
ACCOUNT .EW TOOLS WHICH ALLOW US TO TAKE ACCOUNT OF VARIATION IN RESOURCES AT
DIFFERENT SPATIO
TEMPORAL SCALES AND HOW PLANTS RESPOND TO VARIABLE RESOURCES
AND CLUSTERING ALLOW BOTH INCREASINGLY ACCURATE PREDICTIONS OF DYNAMICS AND
A BETTER UNDERSTANDING OF THE SCALE AND MECHANISMS INVOLVED IN DENSITY DEPEN
DENT RESPONSES
3O FAR WE HAVE ONLY ADDRESSED NEGATIVE EFFECTS OF DENSITY DEPENDENCE ON lT
NESS OR POPULATION GROWTH HOWEVER DENSITY DEPENDENCE CAN ALSO OPERATE POSI
TIVELY 7HERE ANY COMPONENT OF lTNESS INCREASES AS DENSITY OR NUMBERS INCREASE
THIS IS KNOWN AS AN !LLEE EFFECT THIS MAY TRANSLATE INTO DEMOGRAPHIC !LLEE EFFECTS
WHERE POPULATION GROWTH RATE DECLINES WITH DECREASING DENSITY DEPENDING ON
THE STRENGTH OF NEGATIVE DENSITY DEPENDENCE IN THE COMPONENT OF lTNESS IN QUES
TION 3TEPHENS ET AL !N EXAMPLE OF A COMPONENT !LLEE EFFECT IS THAT THERE
MIGHT BE A GREATER PROBABILITY OF lNDING A MATE IN A LARGER POPULATION &OR INSECT
POLLINATED PLANTS SEED SET MAY DEPEND ON THERE BEING NEARBY CON
SPECIlCS
0ARKER AND 0ARKER AND (AUBENSAK GIVE EXAMPLES OF POLLINA
TOR LIMITATION IN THE INVASIVE 3COTCH BROOM #YTISUS SCOPARIUS !N IMPORTANT
CONSEQUENCE OF THE !LLEE EFFECT FOR INVASIONS IS THAT ISOLATED PLANTS OR ISOLATED
POPULATIONS CAN EXPERIENCE A SLOWER POPULATION GROWTH RATE RELATIVE TO LARGER
POPULATIONS 4HIS MAY BE PARTICULARLY IMPORTANT DURING THE ESTABLISHMENT PHASE
9- "UCKLEY AND * -ETCALF
OF INVASIVE SPECIES OR DETERMINE WHAT LEVEL OF CONTROL WILL LEAD TO ERADICATION OR
SLOWER POPULATION GROWTH -ANY INVASIVE SPECIES EXPERIENCE A hLAG TIMEv BETWEEN
INITIAL COLONIZATION AND RAPID POPULATION GROWTH 4HIS MIGHT SIMPLY BE DUE TO THE
LAG PHASE IN AN EXPONENTIAL GROWTH CURVE BUT IT COULD ALSO INDICATE THE ACTION
OF STRONG OR WEAK !LLEE EFFECTS 4HE DISTINCTION BETWEEN STRONG AND WEAK !LLEE
EFFECTS IS AN IMPORTANT ONE AND IS OUTLINED IN 7ANG AND +OT $EMOGRAPHIC
!LLEE EFFECTS OCCUR WHEN THE POPULATION GROWTH RATE AT SOME SET OF LOW DENSITIES
IS SLOWER THAN POPULATION GROWTH RATE AT HIGHER DENSITIES 3TRONG DEMOGRAPHIC
!LLEE EFFECTS OCCUR WHEN POPULATION GROWTH RATE AT LOW DENSITY IS NEGATIVE
)F STRONG !LLEE EFFECTS WERE OPERATING DURING THE ESTABLISHMENT PHASE OF AN INVA
SIVE SPECIES THE POPULATION WOULD NEED TO REACH A THRESHOLD SIZE BEFORE BEING
ABLE TO SPREAD RAPIDLY 4HIS COULD BE ACHIEVED THROUGH ONE LARGE INITIAL RELEASE
OR THROUGH REPEATED INVASIONS EQUIVALENT TO A RESCUE EFFECT IN META
POPULATION
MODELS !MARASEKARE %QUALLY THAT POPULATION COULD BE DRIVEN TO EXTINC
TION BY REDUCING IT BELOW THE THRESHOLD 7EAK DEMOGRAPHIC !LLEE EFFECTS RESULT IN
A POPULATION THAT GROWS MORE SLOWLY AT LOW DENSITIES THAN IT DOES AT HIGH DENSITIES
BUT THAT GROWTH IS ALWAYS POSITIVE SO A THRESHOLD DOES NOT EXIST ! CONSEQUENCE
OF THE OPERATION OF WEAK !LLEE EFFECTS WOULD BE THAT THE LAG PHASE OF AN INVASION
WOULD BE LONGER THAN PREDICTED BY AN EXPONENTIAL GROWTH CURVE
!LLEE EFFECTS CAN BE IMPORTANT DURING INVASIVE SPECIES EXPANSION $AVIS ET AL
DEMONSTRATE THIS FOR 3PARTINA ALTERNImORA WHICH hSETS VERY LITTLE VIABLE SEED
AT THE LEADING EDGES OF AN INVASION IN 7ILLAPA "AY 7ASHINGTON 53! WHERE IT
WAS INTRODUCED C YEARS AGOv 3PARTINA ALTERNImORA HAS TWO DISPERSAL MECHA
NISMS SHORT DISTANCE SPREAD FROM RHIZOMATOUS GROWTH IN EXISTING PLANTS AND
LONG DISTANCE SPREAD VIA SEEDS ,ONG DISTANCE SPREAD FROM THE LEADING EDGE OF THE
POPULATION IS LIMITED HOWEVER BY A LACK OF VIABLE SEED UNTIL THE POPULATION HAS
lLLED IN THROUGH RHIZOMATOUS GROWTH OR GERMINATION OF OTHER SEEDS DISPERSED FROM
EXISTING DENSE POPULATIONS 7EAK !LLEE EFFECTS SUCH THOSE FOUND IN THE INVASIVE
3 ALTERNImORA NOT ONLY AFFECT POPULATION GROWTH RATES AT LOW DENSITY EQUILIBRIUM
DENSITIES AND STABILITY &OWLER AND 2UXTON BUT THEIR INCLUSION IN MODELS OF
SPREAD CAN ALTER SPREAD VELOCITIES 4HE IMPACT OF WEAK !LLEE EFFECTS ON EQUILIBRIA
STABILITY AND SPREAD VELOCITIES HAS ONLY BEEN ADDRESSED RELATIVELY RECENTLY PREVI
OUSLY ONLY STRONG !LLEE EFFECTS HAD BEEN CONSIDERED IN ANY DETAIL &OR INTEGRO
DIFFERENCE EQUATION )$% MODELS OF SPREAD +OT ET AL SHOWED THAT STRONG
!LLEE EFFECTS CAN TURN ACCELERATING INVASIONS INTO CONSTANT SPEED INVASIONS AND
7ANG ET AL SUGGEST USING NUMERICAL SOLUTIONS THAT INCLUDING WEAK
!LLEE EFFECTS CAN ALSO SLOW DOWN INVASIONS FROM ACCELERATING TO CONSTANT SPEEDS
7ANG ET AL ALSO SHOW THAT !LLEE EFFECTS DETERMINE THE DIRECTION OF A
TRAVELLING WAVE SOLUTION OF )$% MODELS WHERE THE DIRECTION OF THE TRAVELLING
WAVE SOLUTION DETERMINES WHETHER AN INVASION WILL SUCCEED OR NOT 4RADITIONAL
REACTION
DIFFUSION MODELS OF SPREAD HAVE ALSO BEEN INVESTIGATED WITH REGARD TO
THE INCLUSION OF !LLEE EFFECTS AGAIN SHOWING THAT !LLEE EFFECTS SLOW DOWN SPREAD
#RUICKSHANK ET AL AND EVEN THAT AN INVASION CAN ONLY SUCCEED ONCE A CER
TAIN CRITICAL AREA IS OCCUPIED ,EWIS AND +AREIVA
$ENSITY DEPENDENCE IN INVASIVE PLANTS
7HETHER DENSITY DEPENDENCE ACTS NEGATIVELY OR THROUGH !LLEE EFFECTS IT WILL
HAVE IMPLICATIONS FOR POPULATION STABILITY 4HIS IS TRUE EVEN IN THE SIMPLEST MOD
ELS SEE THE (ASSELL MODEL ABOVE FOR NEGATIVE DENSITY DEPENDENCE AND 3CHEURING
WHO SHOWS THAT !LLEE EFFECTS PROMOTE STABILITY IN SIMPLE MODELS )NTEREST
IN DENSITY DEPENDENCE OFTEN STEMS FROM THE SEARCH FOR MECHANISMS EITHER PRO
MOTING STABILITY IN PLANT POPULATIONS OR DESTABILISING POPULATIONS -OST PLANT
POPULATIONS ARE BELIEVED TO EXHIBIT STABLE DYNAMICS DUE TO STABILIZING FEATURES OF
THEIR LIFE
HISTORIES SUCH AS SOIL SEED BANKS 7ATKINSON 0ACALA AND 3ILANDER
*ARRY ET AL PHENOTYPIC PLASTICITY AND ASYMMETRIC COMPETITION
0ALMBLAD 7EINER 0ACALA AND 7EINER 3ILVERTOWN
SMALL SIZE THRESHOLDS FOR REPRODUCTION 2EES AND #RAWLEY AND SELF
THINNING
7ATKINSON 7ATKINSON AND $AVY $ESTABILIZING PROCESSES INCLUDE
OVER
COMPENSATING DENSITY DEPENDENT FECUNDITY 4HRALL ET AL TIME LAGS
CAUSED BY MATERNAL EFFECTS #RONE LARGE SIZE THRESHOLDS FOR REPRODUCTION
4HRALL ET AL AND DELAYED mOWERING AT HIGH DENSITY "UCKLEY ET AL
#YCLIC OR CHAOTIC DYNAMICS CAN RESULT FROM AN OVER
COMPENSATING DENSITY DEPEN
DENT RESPONSE COMBINED WITH A HIGH lNITE RATE OF POPULATION GROWTH FROM LOW
DENSITIES /VER
COMPENSATING DENSITY DEPENDENCE OCCURS vWHEN THE PROPORTIONAL
DECLINE IN PARAMETER VALUE WITH INCREASING DENSITY EXCEEDS THE CHANGE IN DENSITYv
&RECKLETON AND 7ATKINSON -ECHANISMS GIVING RISE TO OVER
COMPENSATING
DENSITY DEPENDENCE ARE RARE IN PLANT SYSTEMS AND THEREFORE UNSTABLE DYNAMICS
SUCH AS CYCLES OR CHAOS ARE EQUALLY RARE #HAOTIC DYNAMICS IN ANNUAL PLANT SYS
TEMS ARE REVIEWED BY &RECKLETON 7ATKINSON AND THERE IS LITTLE EVIDENCE
FOR CHAOTIC DYNAMICS IN OTHER PLANT LIFE
HISTORIES 2EES AND #RAWLEY
(OWEVER 2EES ET AL HAVE SHOWN THAT mOWERING DYNAMICS CLOSE TO CHAOTIC
OCCUR IN A MAST
SEEDING TUSSOCK GRASS #HIONOCHLOA PALLENS POSSIBLY AS AN EVOLU
TIONARY RESPONSE TO ATTACK BY SEED AND mOWER FEEDERS
3TABILITY IS AN IMPORTANT ISSUE FOR POPULATIONS OF INVASIVES FOR A NUMBER OF
REASONS KNOWING WHETHER A POPULATION IS LIKELY TO BE STABLE OR EXHIBIT CYCLES MAY
ALLOW PREDICTION OF LEVELS OF IMPACT LIKELY TO OCCUR STABILITY MAY ALSO AFFECT THE
SUCCESS OF BIOCONTROL PROGRAMS &OR PLANT SPECIES WITH LONG
LIVED SEED BANKS IT IS
IMPORTANT TO MAINTAIN POPULATIONS OF A BIOCONTROL AGENT IN THE SYSTEM IN ORDER
TO CONTROL POPULATIONS ARISING FROM THE SEED BANK AT A LATER DATE OR CONTINUOUS
RE
INTRODUCTION OF THE BIOCONTROL AGENT IS NECESSARY 7HEN CHOOSING A BIOCONTROL
AGENT THEREFORE IT MAY BE BETTER TO CHOOSE AN AGENT WHICH MAINTAINS THE PLANT
POPULATION AT A LOW STABLE LEVEL RATHER THAN ONE WHICH DRIVES THE ABOVE GROUND
POPULATION TO EXTINCTION CAUSING ITS OWN EXTINCTION AND THE SUBSEQUENT RESUR
GENCE OF THE INVASIVE PLANT %VEN IF THE HERBIVORE DOES NOT CAUSE PLANT EXTINCTION
DIRECTLY UNSTABLE CYCLES CAN RESULT IN A HIGHER PROBABILITY OF STOCHASTIC EXTINCTION
OF THE PLANT POPULATION "UCKLEY ET AL USE AN EXPERIMENTALLY PARAMETER
ISED PLANT
HERBIVORE MODEL TO SHOW THAT DENSITY DEPENDENCE IN BOTH PLANT AND
HERBIVORE POPULATIONS CONTRIBUTE TO STABILITY IN THE SYSTEM BUT THAT THE FORM OF
DENSITY DEPENDENCE IN THE HERBIVORE IN COMBINATION WITH HIGH ATTACK RATES CAN
DESTABILISE THE DYNAMICS 3TUDYING THE FORM OF DENSITY DEPENDENCE EG SCRAMBLE
9- "UCKLEY AND * -ETCALF
OR CONTEST IN POTENTIAL AGENTS MAY THEREFORE GIVE US SOME IDEA OF WHICH AGENTS
WOULD BE MOST APPROPRIATE TO INTRODUCE IN SYSTEMS LIKE THIS A BIOCONTROL AGENT COM
BINING STRONG SCRAMBLE COMPETITION WITH A HIGH ATTACK RATE MAY NOT BE A GOOD CHOICE
4HE EXACT WAY IN WHICH DENSITY DEPENDENT PROCESSES AFFECT STABILITY CAN BE
SUBTLE $ENSITY DEPENDENCE MAY OCCUR AT SEVERAL LIFE HISTORY STAGES WITHIN A
SPECIES AND WHICH STAGE IT ACTS ON CAN BE OF CRITICAL IMPORTANCE TO ITS EFFECT FOR
POPULATION STABILITY &OR EXAMPLE "UCKLEY AND COLLEAGUES FOUND EVIDENCE
FOR DENSITY DEPENDENT SURVIVAL mOWERING AND FECUNDITY IN THE INVASIVE ANNUAL
4RIPLEUROSPERMUM PERFORATUM 4HESE DIFFERENT OCCURRENCES HAD DIFFERENT IMPLICA
TIONS FOR POPULATION STABILITY $ENSITY DEPENDENCE IN SURVIVAL AND FECUNDITY WAS
STABILISING WHEREAS DENSITY DEPENDENCE IN mOWERING LED TO INSTABILITY IN FACULTA
TIVE BIENNIAL SYSTEMS SEE &IG FOR DENSITY DEPENDENT mOWERING PROBABILITY AND
&IG FOR MODEL STABILITY (OWEVER DENSITY DEPENDENT mOWERING ALSO MEANS THAT
IN THIS FACULTATIVE BIENNIAL DELAYED mOWERING IS FAVOURED AT HIGH DENSITIES 4HIS
LEADS TO ASYMMETRIC COMPETITION BETWEEN LARGER OVER
WINTERED ROSETTES AND NEW
SEEDLINGS IN THE FOLLOWING YEAR WHICH IS GENERALLY A STABILISING FORCE AS THE LARGER
ROSETTES ARE MORE COMPETITIVE WITH SEEDLINGS 3O DENSITY DEPENDENCE NOT ONLY
AFFECTS THE DYNAMICS IN THIS PREDOMINANTLY ANNUAL SYSTEM BUT ALSO PLAYS A PART
IN DETERMINING THE LIFE
HISTORY STRATEGY ADOPTED BY THE PLANT AND THE TIMING OF
DENSITY DEPENDENCE IS IMPORTANT FOR BOTH OF THESE OUTCOMES
4HIS EXAMPLE ALSO ILLUSTRATES HOW THE TIMING OF DENSITY DEPENDENCE CAN BE
CRITICAL FOR DECISIONS RELATING TO MANAGEMENT OF INVASIVE SPECIES -ANAGEMENT
STRATEGIES GENERALLY RESULT IN EXOGENOUS MORTALITY THROUGH MOWING FOR EXAMPLE
$UE TO OVER
COMPENSATING DENSITY DEPENDENCE IN 4 PERFORATUM THE TIMING OF
EXOGENOUS MORTALITY AFFECTS THE EQUILIBRIUM SEED DENSITY DRAMATICALLY )F EXOG
ENOUS MORTALITY OCCURS EARLY IN THE LIFE
CYCLE FREEING THE FEW SURVIVING PLANTS
FROM DENSITY DEPENDENT DEPRESSION OF SEED PRODUCTION SEED OUTPUT CAN ACTUALLY
INCREASE EXACERBATING THE WEED PROBLEM IN THE LONG
TERM "UCKLEY ET AL
'ENERALLY IF DENSITY DEPENDENCE IS IMPORTANT FOR POPULATION REGULATION THE
REMOVAL OF SEEDS SEEDLINGS OR PLANTS BEFORE DENSITY DEPENDENCE OCCURS IS UNLIKELY
TO LEAD TO SUCCESSFUL CONTROL 4HIS IS OFTEN RELEVANT WHEN CONSIDERING BIOCONTROL
AGENTS !S POINTED OUT BY !LEXANDER AND 3CHRAG SEED PREDATORS OR SEEDLING
PATHOGENS CAN HAVE A STRONG EFFECT ON SEEDLING NUMBERS %DWARDS AND #RAWLEY
!LEXANDER AND -IHAIL #UMMINGS AND !LEXANDER BUT NOT
NECESSARILY IMPACT ON lNAL REPRODUCTIVE OUTPUT OF A PATCH DUE TO SUBSEQUENT
DENSITY DEPENDENT PROCESSES 7HERE SEED INPUT IS REDUCED LOW DENSITY SEEDLING
POPULATIONS ARE PRODUCED WHICH MAY HAVE LOWER RATES OF INTRASPECIlC COMPETI
TION AND THUS HIGHER PER CAPITA SEED PRODUCTION COMPENSATING FULLY OR TO SOME
EXTENT FOR THE INITIAL SEED OR SEEDLING LOSS 4HE CASE OF !CACIA NILOTICA ILLUSTRATES THIS
POINT !CACIA NILOTICA AN INVASIVE SHRUB OR SMALL TREE IS DISPERSED BY CATTLE WHICH
EAT SEEDS AND DEPOSIT THEM IN COW PATS 4HE SEEDLINGS ARE THEREFORE AGGREGATED
IN SMALL PATCHES AND SELF
THINNING ENSURES THAT NO MORE THAN APPROXIMATELY ONE
SEEDLING SURVIVES TO ADULTHOOD +RITICOS ET AL AND +RITICOS ET AL
ESTIMATE THAT THE INTRODUCTION OF A SEED
EATING BIOCONTROL AGENT IS THEREFORE
$ENSITY DEPENDENCE IN INVASIVE PLANTS
&IG $ENSITY DEPENDENT FUNCTION FOR mOWERING PROBABILITY IN 4RIPLEUROSPERMUM
PERFORATUM WITH PERMISSION %COLOGY ,ETTERS
&IG ! COMPARISON OF THE DYNAMICS PREDICTED BY THE FULL MODEL INCORPORATING DENSITY
DEPENDENT SURVIVAL mOWERING PROBABILITY AND FECUNDITY WITH A MODEL INCORPORATING
DENSITY DEPENDENT mOWERING PROBABILITY AND FECUNDITY ONLY ! MODEL USING DENSITY
DEPENDENT FECUNDITY ALONE PRODUCES DAMPED OSCILLATIONS AND AN EQUILIBRIUM LEVEL SIMILAR
TO THAT OBSERVED IN THE FULL MODEL WITH PERMISSION %COLOGY ,ETTERS
9- "UCKLEY AND * -ETCALF
UNLIKELY TO AFFECT ADULT POPULATION SIZES UNLESS THE NUMBER OF SEEDS PER COW PAT IS
REDUCED DRAMATICALLY TO TWO OR LESS
3EVERAL AUTHORS HAVE CALLED FOR A THOROUGH UNDERSTANDING OF THE OCCURRENCE
AND EFFECTS OF DENSITY DEPENDENCE AT DIFFERENT STAGES -ORTIMER ET AL
7ATKINSON ET AL 'ILLMAN ET AL !S SHOWN BY THE EXAMPLES GIVEN
ABOVE THIS IS ESPECIALLY RELEVANT FOR WEED SPECIES WHERE DENSITY IS MANIPULATED AT
ONE OR MORE SPECIlC STAGES FOR CONTROL PURPOSES
)N VIEW OF THESE RESULTS IT IS OBVIOUS THAT MODELS SEEKING TO PREDICT THE EFFECTS
OF CONTROL STRATEGIES AT DIFFERENT STAGES SHOULD INCORPORATE DENSITY DEPENDENCE
)N A SUSTAINABLE HARVESTING STUDY &RECKLETON ET AL POINT OUT THAT THE USE
OF DENSITY INDEPENDENT MATRIX MODELS WHERE THE TRANSITION PROBABILITIES ARE CAL
CULATED WHEN THE POPULATION IS CLOSE TO EQUILIBRIUM h CLOSE TO COULD GIVE
MISLEADING RESULTS WHEN USED TO PREDICT POPULATION INCREASE FROM LOW DENSITY
4HEY CALL FOR EXPLICIT INCORPORATION OF DENSITY DEPENDENCE IN HARVESTING MODELS
AND THE SAME CALL SHOULD BE MADE FOR CONTROL MODELS OF INVASIVES WHERE DENSITY
DEPENDENCE IS KNOWN OR SUSPECTED TO BE IMPORTANT
! THOROUGH UNDERSTANDING OF THE OCCURRENCE AND EFFECTS OF DENSITY DEPENDENCE
WOULD ALSO GREATLY CONTRIBUTE TO INVESTIGATIONS INTO SELECTION PRESSURES AND EVO
LUTION )NVASIVE PLANTS CAN BE EXCELLENT MODEL SYSTEMS FOR THE DEVELOPMENT OF
MODELS OF LIFE
HISTORY EVOLUTION 0LANTS IN THE EXOTIC RANGE MAY BE UNDER DIFFER
ENT SELECTION PRESSURES DUE TO ENEMY RELEASE OR COMPETITIVE RELEASE #ONTRASTING
MODELS PREDICTING EVOLUTIONARILY STABLE STRATEGIES IN EACH ENVIRONMENT COULD BE
DEVELOPED 4HIS APPROACH WOULD COMPLEMENT OBSERVATIONAL AND EXPERIMENTAL
STUDIES AIMED AT DETECTING EVOLUTIONARY CHANGE BETWEEN INVASIVES IN THEIR NATIVE
AND EXOTIC HABITATS "LOSSEY AND .TZOLD 7ILLIS ET AL
3IEMANN AND 2OGERS
(ISTORICALLY WHEN EXPLORING LIFE HISTORY STRATEGIES INVESTIGATORS IGNORED DENSI
TY DEPENDENCE AND SIMPLY USED R FROM THE %ULER
,OTKA EQUATION TO DElNE lTNESS
3TEARNS (OWEVER IT BECAME APPARENT THAT DENSITY DEPENDENCE COULD NOT
BE IGNORED BECAUSE IN A DENSITY DEPENDENT SITUATION THE lTNESS OF ONE STRATEGY
WILL DEPEND ON THE ACTIONS OF OTHER STRATEGIES 'RANT AND "ENTON "ECAUSE
OF THIS THE EVOLUTIONARY STABLE STRATEGY %33 IS BEST DETERMINED BY USING THE INVA
SION EXPONENT OR THE RATE OF INCREASE OF A RARE MUTANT INTO AN hENVIRONMENTv
SET BY A RESIDENT STRATEGY AND CHARACTERISED BY THE RESIDENTS DENSITY )F THEN
THE INVADER IS SUCCESSFUL 4HE %33 IS SIMPLY THE STRATEGY THAT CANNOT BE INVADED
BY ANY OTHER 5SING THIS FRAMEWORK -YLIUS AND $IEKMANN SHOWED THAT
IN A CONSTANT ENVIRONMENT THE %33 MAY SOMETIMES BE THE STRATEGY AT WHICH A
RELATIVELY SIMPLE FUNCTION IS MAXIMISED (OWEVER WHICH FUNCTION IS APPROPRIATE
IS DETERMINED BY WHEN DENSITY DEPENDENCE ACTS IN THE LIFE CYCLE )F DENSITY DEPEN
DENCE ACTS SO THAT ONLY THE EXPECTED LIFETIME OFFSPRING PRODUCTION IS REDUCED
THEN AN APPROPRIATE lTNESS MEASURE IS LIFETIME REPRODUCTIVE SUCCESS OR 2 )F DEN
SITY DEPENDENCE AFFECTS ONLY THE PROBABILITY PER UNIT TIME OF DYING THEN POPULA
TION RATE OF INCREASE h IS APPROPRIATE 4HE TWO MAY PROVIDE DIFFERENT PREDICTIONS
SO IT IS IMPORTANT TO USE THE APPROPRIATE MEASURE
$ENSITY DEPENDENCE IN INVASIVE PLANTS
4HIS FRAMEWORK HAS BEEN USED TO PREDICT THE %33 mOWERING STRATEGY IN MONO
CARPIC PLANTS 2EES ET AL 2EES AND 2OSE A CATEGORY TO WHICH MANY
NUISANCE WEEDS AND INVASIVE SPECIES BELONG $ATA FROM A RANGE OF SPECIES SHOWS
THAT NEIGHBOUR DENSITY OR SIZE HAS NO EFFECT ON GROWTH SURVIVAL OR REPRODUC
TION IN MONOCARPIC PLANTS -ETCALF ET AL (OWEVER RECRUITMENT IS GENER
ALLY UNCORRELATED WITH SEED PRODUCTION SUGGESTING THAT SEED ESTABLISHMENT IS
LIMITED BY SUITABLE MICROSITE AVAILABILITY 4OGETHER THESE OBSERVATIONS IMPLY
THE OPERATION OF DENSITY DEPENDENT PROCESSES AT THE SEEDLING ESTABLISHMENT LEVEL
ONLY 7E CAN DEDUCE THAT THE %33 STRATEGY CORRESPONDS TO MAXIMISING 2 ASSUM
ING THE ENVIRONMENT IS CONSTANT ! MATRIX MODEL OR ITS CONTINUOUS ANALOGUE
THE )NTEGRAL 0ROJECTION -ODEL %ASTERLING ET AL CAN BE USED TO DESCRIBE
THE POPULATION AND CALCULATE 2 FOR A RANGE OF mOWERING STRATEGIES 4HIS APPROACH
HAS BEEN USED TO PREDICT %33 mOWERING STRATEGY VALUES THAT ARE VERY CLOSE TO THE
OBSERVED 2EES AND 2OSE AND CONTRIBUTE TO OUR UNDERSTANDING OF SELECTION
PRESSURES ON MONCARPIC PLANTS
#/.#,53)/.3
$ENSITY DEPENDENT PROCESSES CAN DETERMINE EQUILIBRIUM DENSITIES ACHIEVABLE BY
AN INVASIVE POPULATION STABILITY OF THAT POPULATION POPULATION GROWTH RATES AT
THE FRONT OF AN INVASION POPULATION RECOVERY AFTER CONTROL MEASURES HAVE BEEN
IMPLEMENTED AND THE EVOLUTIONARILY STABLE STRATEGIES ACQUIRED BY POPULATIONS IN
DIFFERENT ENVIRONMENTS )N THIS CHAPTER WE HAVE OUTLINED HOW THEORETICAL ECOLOGY
HAS CONTRIBUTED BOTH TO OUR UNDERSTANDING OF HOW INVASIVE POPULATIONS ARE REGU
LATED BY DENSITY DEPENDENCE AND HOW WORK ON INVASIVES HAS STIMULATED THEORETI
CAL WORK ON IMPORTANT ECOLOGICAL PROBLEMS LIKE !LLEE EFFECTS AND SPATIAL SPREAD OF
POPULATIONS !S WE DEVELOP MODELS FOR AN INCREASING NUMBER OF THE COMPLEX
PROCESSES THAT GOVERN POPULATION DYNAMICS WE CAN START TO COMBINE MODELS AND
INVESTIGATE THE INTERACTION OF PROCESSES LIKE !LLEE EFFECTS AND THE SHAPE OF DISPERSAL
KERNELS 7ANG ET AL ON SPREAD RATES OR DENSITY DEPENDENCE AT DIFFERENT
STAGES IN THE LIFE CYCLE "UCKLEY ET AL 5LTIMATELY THESE MODELS WILL ENABLE
US TO APPLY TARGETED AND EFFECTIVE MANAGEMENT STRATEGIES TO INVASIVE PLANT POPULA
TIONS IN ORDER TO REDUCE THEIR IMPACT ON BIODIVERSITY LOSS AND ECOSYSTEM FUNCTION
!#+./7,%$'%-%.43
7E THANK $ARREN +RITICOS +AREN 2OSE #AZ 4AYLOR $AVE -URRELL AND AN ANONY
MOUS REVIEWER FOR HELPFUL ADDITIONS *#- WAS SUPPORTED BY A .%2# STUDENTSHIP
9- "UCKLEY AND * -ETCALF
2%&%2%.#%3
!LEXANDER ( - AND * $ -IHAIL 3EEDLING DISEASE IN AN ANNUAL LEGUME CONSE
QUENCES FOR SEEDLING MORTALITY PLANT SIZE AND POPULATION SEED PRODUCTION /ECOLOGIA
!LEXANDER ( - AND ! - 3CHRAG 2OLE OF SOIL SEED BANKS AND NEWLY DISPERSED SEEDS
IN POPULATION DYNAMICS OF THE ANNUAL SUNmOWER (ELIANTHUS ANNUUS *OURNAL OF %COLOGY
!MARASEKARE 0 )NTERACTIONS BETWEEN LOCAL DYNAMICS AND DISPERSAL )NSIGHTS FROM
SINGLE SPECIES MODELS 4HEORETICAL 0OPULATION "IOLOGY
"ENTON 4 ' 4 # #AMERON AND ! 'RANT 0OPULATION RESPONSES TO PERTURBATIONS
PREDICTIONS AND RESPONSES FROM LABORATORY MITE POPULATIONS *OURNAL OF !NIMAL %COLOGY
"LOSSEY " AND 2 .TZOLD %VOLUTION OF INCREASED COMPETITIVE ABILITY IN INVASIVE
NONINDIGENOUS PLANTS A HYPOTHESIS *OURNAL OF %COLOGY
"ONSALL - " % VAN DER -EIJDEN AND - * #RAWLEY #ONTRASTING DYNAMICS IN THE
SAME PLANT
HERBIVORE INTERACTION 0ROCEEDINGS OF THE .ATIONAL !CADEMY OF 3CIENCES
"UCKLEY 9 - $ 4 "RIESE AND - 2EES A $EMOGRAPHY AND MANAGEMENT OF THE
INVASIVE PLANT SPECIES (YPERICUM PERFORATUM ) 5SING MULTI
LEVEL MIXED
EFFECTS MODELS
FOR CHARACTERIZING GROWTH SURVIVAL AND FECUNDITY IN A LONG
TERM DATA SET *OURNAL OF
!PPLIED %COLOGY
"UCKLEY 9 - 0 $OWNEY 3 6 &OWLER 2 , (ILL * -EMMOTT ( .ORAMBUENA -
0ITCAIRN 2 ' 3HAW ! 7 3HEPPARD # 7INKS 2 7ITTENBERG AND - 2EES B !RE
INVASIVES BIGGER ! GLOBAL STUDY OF SEED SIZE VARIATION IN TWO INVASIVE SHRUBS %COLOGY
"UCKLEY 9 - ( , (INZ $ -ATTHIES AND - 2EES )NTERACTIONS BETWEEN DENSITY
DEPENDENT PROCESSES POPULATION DYNAMICS AND CONTROL OF AN INVASIVE PLANT SPECIES
4RIPLEUROSPERMUM PERFORATUM SCENTLESS CHAMOMILE %COLOGY ,ETTERS
"UCKLEY 9 - - 2EES ! 7 3HEPPARD AND - * 3MYTH 3TABLE COEXISTENCE OF AN
INVASIVE PLANT AND BIOCONTROL AGENT A PARAMETERISED COUPLED PLANT
HERBIVORE MODEL
*OURNAL OF !PPLIED %COLOGY
#AMERON 4 # AND 4 ' "ENTON 3TAGE
STRUCTURED HARVESTING AND ITS EFFECTS AN
EMPIRICAL INVESTIGATION USING SOIL MITES *OURNAL OF !NIMAL %COLOGY
#ASWELL ( -ATRIX POPULATION MODELS CONSTRUCTION ANALYSIS AND INTERPRETATION
ND EDITION 3INAUER !SSOCIATES )NC 3UNDERLAND -ASSACHUSETTS
#RONE % % 0ARENTAL ENVIRONMENTAL EFFECTS AND CYCLICAL DYNAMICS IN PLANT POPULA
TIONS 4HE !MERICAN .ATURALIST
#RUICKSHANK ) 7 3 # 'URNEY AND ! 2 6EITCH 4HE CHARACTERISTICS OF EPIDEMICS
AND INVASIONS WITH THRESHOLDS 4HEORETICAL 0OPULATION "IOLOGY
#UMMINGS # , AND ( - !LEXANDER 0OPULATION ECOLOGY OF WILD SUNmOWERS EFFECTS
OF SEED DENSITY AND POST
DISPERSAL VERTEBRATE SEED PREDATION /ECOLOGIA
$AVIS ( ' # - 4AYLOR * # #IVILLE AND $ 2 3TRONG !N !LLEE EFFECT AT THE FRONT OF
A PLANT INVASION 3PARTINA IN A 0ACIlC ESTUARY *OURNAL OF %COLOGY
$ENSITY DEPENDENCE IN INVASIVE PLANTS
%ASTERLING - 2 3 0 %LLNER AND 0 - $IXON 3IZE SPECIlC SENSITIVITY !PPLYING A
NEW STRUCTURED POPULATION MODEL %COLOGY
%DWARDS ' 2 AND - * #RAWLEY 2ODENT SEED PREDATION AND SEEDLING RECRUITMENT
IN MESIC GRASSLAND /ECOLOGIA
&OWLER 3 6 AND ' $ 2UXTON 0OPULATION DYNAMIC CONSEQUENCES OF !LLEE EFFECTS
*OURNAL OF 4HEORETICAL "IOLOGY
&OX ' ! AND " % +ENDALL $EMOGRAPHIC STOCHASTICITY AND THE VARIANCE REDUCTION
EFFECT %COLOGY
&RECKLETON 2 0 $ - 3ILVA -ATOS - , ! "OVI AND ! 2 7ATKINSON 0REDICTING
THE IMPACTS OF HARVESTING USING STRUCTURED POPULATION MODELS THE IMPORTANCE OF
DENSITY DEPENDENCE AND TIMING OF HARVEST FOR A TROPICAL PALM TREE *OURNAL OF !PPLIED
%COLOGY
&RECKLETON 2 0 AND ! 2 7ATKINSON !RE WEED POPULATION DYNAMICS CHAOTIC
*OURNAL OF !PPLIED %COLOGY
'ILLMAN - * - "ULLOCK * 3ILVERTOWN AND " #LEAR (ILL ! DENSITY DEPENDENT
MODEL OF #IRSIUM VULGARE POPULATION DYNAMICS USING lELD ESTIMATED PARAMETER VALUES
/ECOLOGIA
'RANT ! AND 4 ' "ENTON %LASTICITY ANALYSIS FOR DENSITY DEPENDENT POPULATIONS IN
STOCHASTIC ENVIRONMENTS %COLOGY
(ARPER * , AND 7 ! 7OOD "IOLOGICAL &LORA OF THE "RITISH )SLES 3ENECIO JACOBAEA ,
*OURNAL OF %COLOGY
(ASSELL - 0 $ENSITY DEPENDENCE IN SINGLE SPECIES POPULATIONS *OURNAL OF !NIMAL
%COLOGY
*ARRY - - +HALADI - (OSSAERT -C+EY AND $ -C+EY -ODELING THE POPULA
TION DYNAMICS OF ANNUAL PLANTS WITH SEED BANK AND DENSITY DEPENDENT EFFECTS !CTA
"IOTHEORETICA
+EANE 2 - AND - * #RAWLEY %XOTIC PLANT INVASIONS AND THE ENEMY RELEASE
HYPOTHESIS 4RENDS IN %COLOGY %VOLUTION
+OT - - ! ,EWIS AND 0 VAN DEN $RIESSCHE $ISPERSAL DATA AND THE SPREAD OF
INVADING ORGANISMS %COLOGY
+REBS # * 4WO COMPLEMENTARY PARADIGMS FOR ANALYSING POPULATION DYNAMICS
0HILOSOPHICAL 4RANSACTIONS OF THE 2OYAL 3OCIETY OF ,ONDON 3ERIES " "IOLOGICAL 3CIENCES
+RITICOS $ * "ROWN ) 2ADFORD AND - .ICHOLAS 0LANT POPULATION ECOLOGY AND
BIOLOGICAL CONTROL !CACIA NILOTICA AS A CASE STUDY "IOLOGICAL #ONTROL
+RITICOS $ * * 2 "ROWN ' & -AYWALD ) $ 2ADFORD $ - .ICHOLAS 2 7 3UTHERST AND
3 7 !DKINS 30!N$8 A PROCESS BASED POPULATION DYNAMICS MODEL TO EXPLORE
MANAGEMENT AND CLIMATE CHANGE IMPACTS ON AN INVASIVE ALIEN PLANT !CACIA NILOTICA
%COLOGICAL MODELLING
,EWIS - ! AND 0 +AREIVA !LLEE DYNAMICS AND THE SPREAD OF INVADING ORGANISMS
4HEORETICAL 0OPULATION "IOLOGY
-ETCALF * # + % 2OSE AND - 2EES %VOLUTIONARY DEMOGRAPHY OF MONOCARPIC
PERENNIALS 4RENDS IN %COLOGY %VOLUTION
9- "UCKLEY AND * -ETCALF
-ORTIMER ! - * * 3UTTON AND 0 'OULD /N ROBUST WEED POPULATION MODELS 7EED
2ESEARCH
-YLIUS 3 $ AND / $IEKMANN /N EVOLUTIONARILY STABLE LIFE HISTORIES AND THE NEED
TO BE SPECIlC ABOUT DENSITY DEPENDENCE /IKOS
0ACALA 3 7 AND * ! 3ILANDER &IELD TESTS OF NEIGHBORHOOD POPULATION DYNAMIC
MODELS OF TWO ANNUAL WEED SPECIES %COLOGICAL -ONOGRAPHS
0ACALA 3 7 AND * 7EINER %FFECTS OF COMPETITIVE ASYMMETRY ON A LOCAL DENSITY
MODEL OF PLANT INTERFERENCE *OURNAL OF 4HEORETICAL "IOLOGY
0ALMBLAD ) ' #OMPETITION IN EXPERIMENTAL STUDIES ON POPULATIONS OF WEEDS WITH
PARTICULAR EMPHASIS ON THE REGULATION OF POPULATION SIZE %COLOGY
0ARKER ) - 0OLLINATOR LIMITATION OF #YTISUS SCOPARIUS 3COTCH BROOM AN INVASIVE
EXOTIC SHRUB %COLOGY
0ARKER ) - AND + ! (AUBENSAK #OMPARATIVE POLLINATOR LIMITATION OF TWO NON
NATIVE SHRUBS DO MUTUALISMS INmUENCE INVASIONS /ECOLOGIA
0AYNTER 1 0 / $OWNEY AND ! 7 3HEPPARD !GE STRUCTURE AND GROWTH OF THE
WOODY LEGUME WEED #YTISUS SCOPARIUS IN NATIVE AND EXOTIC HABITATS IMPLICATIONS FOR
CONTROL *OURNAL OF !PPLIED %COLOGY
0IMENTEL $ , ,ACH 2 :UNIGA AND $ -ORRISON %NVIRONMENTAL AND ECONOMIC
COSTS OF NONINDIGENOUS SPECIES IN THE 5NITED 3TATES "IOSCIENCE
2EES - AND - * #RAWLEY 'ROWTH REPRODUCTION AND POPULATION DYNAMICS
&UNCTIONAL %COLOGY
2EES - AND - * #RAWLEY $O PLANT POPULATIONS CYCLE &UNCTIONAL %COLOGY
2EES - $ +ELLY AND / . "JORNSTAD 3NOW TUSSOCKS CHAOS AND THE EVOLUTION OF
MAST SEEDING 4HE !MERICAN .ATURALIST
2EES - AND + % 2OSE %VOLUTION OF mOWERING STRATEGIES IN /ENOTHERA GLAZIOVIANA
AN INTEGRAL PROJECTION MODEL APPROACH 0ROCEEDINGS OF THE 2OYAL 3OCIETY ,ONDON 3ERIES
"
3CHEURING ) !LLEE EFFECT INCREASES THE DYNAMICAL STABILITY OF POPULATIONS *OURNAL OF
4HEORETICAL "IOLOGY
3HIMA * 3 AND # 7 /SENBERG #RYPTIC DENSITY DEPENDENCE EFFECTS OF COVARIATION
BETWEEN DENSITY AND SITE QUALITY IN REEF lSH %COLOGY
3IBLY 2 - AND * (ONE 0OPULATION GROWTH RATE AND ITS DETERMINANTS AN OVERVIEW
0HILOSOPHICAL 4RANSACTIONS OF THE 2OYAL 3OCIETY OF ,ONDON 3ERIES " "IOLOGICAL 3CIENCES
3IEMANN % AND 7 % 2OGERS 'ENETIC DIFFERENCES IN GROWTH OF AN INVASIVE TREE SPE
CIES %COLOGY ,ETTERS
3IEMANN % AND 7 % 2OGERS 2EDUCED RESISTANCE OF INVASIVE VARIETIES OF THE ALIEN
TREE 3APIUM SEBIFERUM TO A GENERALIST HERBIVORE /ECOLOGIA
3ILVERTOWN * -ODULARITY REPRODUCTIVE THRESHOLDS AND PLANT POPULATION DYNAMICS
&UNCTIONAL %COLOGY
3TEARNS 3 # 4HE EVOLUTION OF LIFE HISTORIES /XFORD 5NIVERSITY 0RESS /XFORD
3TEPHENS 0 ! 7 * 3UTHERLAND AND 2 &RECKLETON 7HAT IS THE !LLEE EFFECT /IKOS
$ENSITY DEPENDENCE IN INVASIVE PLANTS
4HRALL 0 ( 3 7 0ACALA AND * ! 3ILANDER /SCILLATORY DYNAMICS IN POPULATIONS OF
AN ANNUAL WEED SPECIES !BUTILON THEOPHRASTI *OURNAL OF %COLOGY
4URCHIN 0 0OPULATION REGULATION A SYNTHETIC VIEW /IKOS
7ANG - ( - +OT AND - ' .EUBERT )NTEGRODIFFERENCE EQUATIONS !LLEE EFFECTS
AND INVASIONS *OURNAL OF -ATHEMATICAL "IOLOGY
7ATKINSON ! 2 $ENSITY DEPENDENCE IN SINGLE SPECIES POPULATIONS OF PLANTS
*OURNAL OF 4HEORETICAL "IOLOGY
7ATKINSON ! 2 AND ! * $AVY 0OPULATION BIOLOGY OF SALT MARSH AND SAND DUNE
ANNUALS 6EGETATIO
7ATKINSON ! 2 7 - ,ONSDALE AND - ( !NDREW -ODELLING THE POPULATION
DYNAMICS OF AN ANNUAL PLANT 3ORGHUM INTRANS IN THE WET DRY TROPICS *OURNAL OF %COLOGY
7EINER * 3IZE HIERARCHIES IN EXPERIMENTAL POPULATIONS OF ANNUAL PLANTS %COLOGY
7ILLIS ! * * -EMMOTT AND 2 ) &ORRESTER )S THERE EVIDENCE FOR THE POST INVASION
EVOLUTION OF VIGOUR AMONG INVASIVE PLANT SPECIES %COLOGY ,ETTERS
7ILLIS ! * - " 4HOMAS AND * ( ,AWTON )S THE INCREASED VIGOUR OF INVASIVE
WEEDS EXPLAINED BY A TRADE OFF BETWEEN GROWTH AND HERBIVORE RESISTANCE /ECOLOGIA
#HAPTER SIX
3TOCHASTICITY NONLINEARITY
AND INSTABILITY IN BIOLOGICAL
INVASIONS
2 0 &RECKLETON 0 - $OWLING AND . + $ULVY
).42/$5#4)/.
#OMPARED WITH OTHER SCIENCES ECOLOGY HAS FEW GENERAL THEORIES THAT CAN BE
APPLIED TO PREDICT THE DYNAMICS OF ITS SYSTEMS FROM lRST PRINCIPLES 4HIS IS PERHAPS
NOT SURPRISING GIVEN THE HUGE VARIABILITY OF ECOLOGICAL SYSTEMS ! NUMBER OF
BROAD INSIGHTS HAVE EMERGED HOWEVER AND ONE OF THE MOST IMPORTANT OF THESE IS
THAT PREDICTING THE DYNAMICS OF ECOLOGICAL SYSTEMS REQUIRES THAT WE UNDERSTAND
THE INTERPLAY OF TWO KINDS OF PROCESSES STOCHASTIC AND DETERMINISTIC ,EWONTIN
AND #OHEN -AY ,ANDE ET AL 4HIS CONCLUSION IS ESSENTIALLY
A CONSENSUS RESULTING FROM A GREAT DEAL OF DEBATE IN THE ECOLOGICAL LITERATURE
DATING BACK TO THE S EG !NDREWARTHA AND "IRCH .ICHOLSON
(ASSELL 7HITE
0REDICTING BIOLOGICAL INVASIONS IS AN EXTREMELY IMPORTANT APPLIED ECOLOGI
CAL PROBLEM BECAUSE INVASIVE SPECIES ARE ECONOMICALLY ENORMOUSLY SIGNIlCANT
IN AGRICULTURAL AND CONSERVATION TERMS AND METHODS FOR PREDICTING THE EFFECTS
OF CONTROL MEASURES ARE URGENTLY REQUIRED "YERS ET AL /NE OF THE MOST
IMPORTANT APPROACHES TO STUDYING BIOLOGICAL INVASIONS IS THROUGH POPULATION
MODELLING 0OPULATION MODELS MAY TAKE A RANGE OF FORMS INCLUDING SIMPLE
-7 #ADOTTE ET AL (EDS) #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY
3PRINGER 0RINTED IN THE .ETHERLANDS
20 &RECKLETON 0- $OWLING AND .+ $ULVY
MATHEMATICAL MODELS (ASSELL -AY -AY ET AL -AYNARD 3MITH
7ATKINSON STATISTICAL MODELS "UCKLEY ET AL A B AND COMPLEX
SIMULATIONS 0ACALA ET AL 4HESE MODELS CAN BE USED IN A RANGE OF WAYS
FROM MAKING HIGHLY SPECIlC PREDICTIONS ABOUT THE DYNAMICS OF SPECIES IN GIVEN
AREAS THROUGH TO GENERAL ANALYSES AIMED AT DEVELOPING A GENERAL UNDERSTANDING
OF THE PROCESSES DETERMINING INVASION SUCCESS
5NFORTUNATELY POPULATION MODELS ARE FREQUENTLY EQUATED WITH AN EQUILIBRIUM
OR DETERMINISTIC OUTLOOK ON POPULATION DYNAMICS WHICH IS FREQUENTLY CRITICISED
(OWEVER EVEN SIMPLE MODELS MAY FREQUENTLY YIELD RATHER COMPLEX OUTCOMES
-AY AND /STER &OR INSTANCE A SIMPLE LOGISTIC MODEL OF POPULATION GROWTH
MIGHT BE EXPECTED TO YIELD A STRAIGHTFORWARD OUTCOME NAMELY THE SMOOTH GROWTH
OF A POPULATION TO ITS CARRYING CAPACITY 7HEN IMPLEMENTED AS A DISCRETE TIME
MODEL HOWEVER THE OUTCOME MAY BE FAR MORE COMPLEX AND CHAOTIC DYNAMICS
MAY BE POSSIBLE &ROM THE POINT OF VIEW OF PREDICTING POPULATION DYNAMICS THIS
RAISED THE POSSIBILITY THAT COMPLETELY DETERMINISTIC SYSTEMS MAY YIELD UNPREDICT
ABLE CHANGES IN POPULATION NUMBERS 4HE EARLY S ALSO SAW AN INCREASING REC
OGNITION THAT VARIOUS FORMS OF INTERACTIONS AND INTERVENTION COULD YIELD UNSTABLE
POPULATION DYNAMICS -AY &OR EXAMPLE HUNTING HARVESTING OR PREDATION
COULD POTENTIALLY DESTABILISE POPULATION DYNAMICS TO THE EXTENT THAT OTHERWISE
STABLE POPULATIONS MIGHT BECOME EXTINCT GIVEN A PERTURBATION TO THE SYSTEM
! lNAL ELEMENT THAT IS IMPORTANT IN PREDICTING POPULATION CHANGE IS THE STO
CHASTIC COMPONENT OF POPULATION CHANGE 3TOCHASTICITY RESULTS FROM RANDOM
VARIATIONS IN THE ENVIRONMENT OR FROM RANDOM VARIATION BETWEEN INDIVIDUALS
4HE THEORY FOR STOCHASTIC POPULATION DYNAMICS HAS A LONG HISTORY #OHEN
,EWONTIN AND #OHEN 4ULJAPURKAR AND THERE HAVE BEEN NUMEROUS
RECENT DEVELOPMENTS OF THIS ,ANDE ET AL 4HE MAJOR CONCLUSION FROM THIS
WORK HAS BEEN THAT THE EFFECTS OF STOCHASTICITY ON POPULATION DYNAMICS MAY BE
COMPLEX 0REDICTING THE IMPACTS OF STOCHASTICITY IS NOT A STRAIGHTFORWARD AS MAK
ING SOME PARAMETERS OF MODELS RANDOM VARIABLES OR OF CALCULATING AN AVERAGE
OF SOME FORM 2ATHER STOCHASTICITY MAY IMPACT IN A PROFOUND AND A QUALITATIVE
MANNER ON POPULATION DYNAMICS
4HE PROBLEM OF PREDICTING AND MANAGING POPULATIONS OF INVASIVE SPECIES
REQUIRES THAT WE ARE ABLE TO UNDERSTAND THE IMPACTS OF THE FACTORS DETAILED
ABOVE ON SEVERAL PHASES OF THE INVASION PROCESS 4HE ELEMENTS OF POPULATION
DYNAMICS OF INVASION CAN BE CONCEPTUALLY VIEWED AS CONSISTING OF THREE PHASES
ARRIVAL ESTABLISHMENT AND SPREAD $OBSON AND -AY 7ILLIAMSON
4HE PROCESSES DESCRIBED ABOVE MIGHT BE EXPECTED TO PLAY DIFFERENT ROLES DEPEND
ING ON WHICH OF THIS PHASES THE INVADING POPULATION IS IN )N THE ARRIVAL AND
ESTABLISHMENT PHASES POPULATIONS ARE GROWING FROM LOW DENSITIES AND EXPANDING
INTO NEW HABITAT )N THIS PHASE STOCHASTIC FACTORS MAY ARGUABLY BE IMPORTANT AND
LIMITING FACTORS LESS SO )N CONTRAST FOLLOWING SUCCESSFUL ESTABLISHMENT AND DURING
THE SPREAD PHASE POPULATIONS MAY BE AT MUCH HIGHER DENSITIES AND FACTORS SUCH AS
DENSITY DEPENDENCE MAY DOMINATE POPULATION CHANGES #ORRESPONDINGLY THESE
DIFFERENCES IN DYNAMICS MAY INmUENCE CONTROL STRATEGIES 4AYLOR AND (ASTINGS
3TOCHASTICITY NONLINEARITY AND INSTABILITY IN BIOLOGICAL INVASIONS
-OREOVER WHEN FACTORS SUCH AS !LLEE EFFECTS OPERATE THE DETERMINISTIC
COMPONENT OF DYNAMICS MAY BE SIGNIlCANT AT LOW DENSITIES AND INmUENCE POPULA
TION PERSISTENCE 4HE KEY TASK FOR POPULATION MODELLING IS TO WORK OUT HOW THESE
VARIOUS FACTORS INTERACT WITH EACH OTHER
4HIS CHAPTER IS ORGANISED INTO THREE SECTIONS )N THE lRST SECTION WE REVIEW THE
PRINCIPLES OF SIMPLE STOCHASTIC MODELS AND HOW THEY MAY BE APPLIED TO INVASIONS
7E THEN EXAMINE THE DETERMINISTIC COUNTERPARTS OF THESE MODELS AND THE PROB
LEMS THESE PRESENT &INALLY WE BRIEmY REVIEW TWO STUDIES THAT EXHIBIT UNSTABLE
DYNAMICS AND SPECULATE ON THE GENERALITY OF THESE
34/#(!34)# 0/05,!4)/. $9.!-)#3 !.$ 4(% 34!"),)49
/& 0/05,!4)/.3
)T IS USEFUL TO RECOGNISE TWO SOURCES OF STOCHASTICITY IN POPULATION DYNAMICS
%NGEN ET AL SINCE THE SOURCE OF STOCHASTICITY IS IMPORTANT IN UNDERSTAND
ING ITS EFFECTS ON POPULATION DYNAMICS %NVIRONMENTAL STOCHASTICITY RESULTS FROM
RANDOM VARIATIONS IN DEMOGRAPHIC RATES AS A CONSEQUENCE OF VARIATION IN ENVI
RONMENTAL CONDITIONS FOR EXAMPLE RESULTING FROM THE EFFECTS OF WEATHER ! SECOND
SOURCE OF STOCHASTICITY IS DEMOGRAPHIC STOCHASTICITY 5NDER DEMOGRAPHIC STOCHAS
TICITY THE EXPECTED CONTRIBUTION OF EACH INDIVIDUAL TO THE NEXT GENERATION IS ALSO
INDEPENDENT OF THE STATE OF THE POPULATION IN THE PREVIOUS GENERATION HOWEVER
THE VARIANCE ABOUT THIS EXPECTATION DOES DEPEND ON THE STATE OF THE POPULATION
AS DISTINCT FROM AN !LLEE EFFECT IN WHICH THE EXPECTATION DEPENDS ON DENSITY
3TEPHENS ET AL
3IMPLE STOCHASTIC POPULATION GROWTH
4HE SIMPLEST MODEL FOR POPULATION DYNAMICS IS ONE IN WHICH THE EXPECTED RATES OF
DEATH AND FECUNDITY ARE ASSUMED TO BE CONSTANT AND VARY RANDOMLY ABOUT THESE
MEANS )F THE NET RATE OF POPULATION CHANGE IN YEAR T IS h T AND .T AND .T
ARE POPULATION SIZES IN SUCCESSIVE YEARS THEN THE DYNAMICS OF SUCH A POPULATION
CAN BE MODELLED AS
.T hT .T
&OR A SPECIES TO BE CAPABLE OF INVASION THE EXPECTED VALUE OF .T 4 .T MUST
BE GREATER THAN ONE AS 4 BECOMES LARGE )N A CONSTANT ENVIRONMENT THIS REQUIRES
THAT THE VALUE OF h MUST BE GREATER THAN UNITY )N AN ENVIRONMENT IN WHICH THERE
IS STOCHASTICITY THE CONDITION ON h IS NOT AS STRAIGHTFORWARD BUT DEPENDS ON THE
DISTRIBUTION OF h ABOUT ITS MEAN !CCORDING TO EQUATION POPULATION SIZE AT
TIME T 4 IS A NON LINEAR FUNCTION OF hT hT xhT 4 4HIS MEANS THAT THE DISTRI
BUTION OF h HAS TO BE UNDERSTOOD IN ORDER TO PREDICT LONG TERM DYNAMICS -ORE
COMPLEX EFFECTS SUCH AS THE AUTOCORRELATIONS BETWEEN POPULATION GROWTH RATES IN
SUCCESSIVE YEARS MAY ALSO COME INTO PLAY AND FURTHER COMPLICATE PREDICTIONS OF
20 &RECKLETON 0- $OWLING AND .+ $ULVY
POPULATION PERSISTENCE (EINO ET AL !LTHOUGH THE POSSIBILITIES ARE COMPLEX
TWO SIMPLE EXAMPLES ILLUSTRATE SOME OF THE MAIN FEATURES OF STOCHASTIC MODELS
4HE MAIN CONCLUSION AT THE END OF THIS SECTION IS THAT THE OUTCOME OF STOCHASTIC
MODELS CAN BE MORE COMPLEX THAN ONE MIGHT lRST IMAGINE
%NVIRONMENTAL STOCHASTICITY
&IRSTLY ASSUME THAT h IS LOG NORMALLY DISTRIBUTED AS MAY BE THE CASE IN POPULA
TIONS SUBJECT TO ENVIRONMENTAL STOCHASTICITY )N THAT CASE EQUATION MAY BE
WRITTEN AS WHERE N >LOG .
NT NT LOG ;hT =
4HIS IS LINEAR IN N WITH THE CONSEQUENCE THAT THE ARITHMETIC MEAN OF LOG h
PREDICTS THE LONG TERM POPULATION TRAJECTORY )T IS EASY TO SHOW THAT THE EXPECTED
VALUE OF NT 4 IS NORMALLY DISTRIBUTED WITH MEAN N 4 LOGh AND VARIANCE
4 VAR ;LOGh = IF THE LAMBDAS ARE INDEPENDENT AND NORMALLY DISTRIBUTED &IG A
SHOWS EXAMPLES OF A SERIES OF POPULATIONS GROWING IN THIS WAY AND &IG C SHOWS
THE DISTRIBUTION OF POPULATION SIZES AFTER A PERIOD OF POPULATION GROWTH
!S NOTED ABOVE PREDICTING THE OUTCOME OF STOCHASTICITY REQUIRES THAT THE EFFECTS
OF STOCHASTICITY ON POPULATION GROWTH ARE UNDERSTOOD IN DETAIL &OR INSTANCE
IF THE DISTRIBUTION OF POPULATION GROWTH RATES IS NOT LOG
NORMAL OR IF SUCCESSIVE
POPULATION GROWTH RATES ARE NOT INDEPENDENT THEN THE RESULTS IN THE PREVIOUS
PARAGRAPH DO NOT HOLD EXACTLY )N SUCH CASES STOCHASTIC CALCULUS HAS TO BE USED IN
ORDER TO DETERMINE THE APPROPRIATE METHOD OF TRANSFORMING POPULATION DENSITIES
IN ORDER TO PREDICT PERSISTENCE EG ,ANDE
$EMOGRAPHIC STOCHASTICITY
4HE SECOND EXAMPLE IS A POPULATION SUBJECT TO DEMOGRAPHIC STOCHASTICITY -AY
,ANDE ,ANDE ET AL $EMOGRAPHIC STOCHASTICITY RESULTS IN
VARIABILITY IN WHICH THE VARIANCE DECLINES WITH THE NUMBER OF INDIVIDUALS IN THE
POPULATION
m $
VAR h
.
OR AS A HYPERBOLIC FUNCTION OF DENSITY IF . IS THE AVERAGE NUMBER OF INDIVIDUALS
PER UNIT AREA AND INDIVIDUALS ARE ASSUMED TO BE RANDOMLY DISTRIBUTED &RECKLETON
UNPUB
m $
VAR h
D.
7HEN COMBINED WITH ENVIRONMENTAL VARIABILITY WHICH HAS VARIANCE m%
THE VARIANCE IN POPULATION GROWTH RATE IN THIS MODEL IS
3TOCHASTICITY NONLINEARITY AND INSTABILITY IN BIOLOGICAL INVASIONS
m $
VAR h m %
.
%XAMPLES OF POPULATIONS GROWING ACCORDING TO THIS MODEL ARE SHOWN IN &IG B
4HE MOST IMPORTANT FEATURE OF THIS MODEL IS THAT IF DENSITY FALLS BELOW A CRITICAL
THRESHOLD
.
m$ h n n m%
THEN EXTINCTION IS HIGHLY LIKELY ,ANDE ET AL 4HE RESULT OF THIS IS A BIMODAL
DISTRIBUTION OF POPULATION SIZES AS SHOWN IN &IG D COMPARED WITH THE NORMAL
DISTRIBUTION IN &IG C
&IG 3OURCES OF STOCHASTICITY AND EFFECTS ON POPULATION GROWTH A C AND POPULATION SIZE
B D 0OPULATION GROWTH WAS DENSITY INDEPENDENT WITH STOCHASTICITY EITHER ENVIRONMENTAL
A C OR ENVIRONMENTAL AND DEMOGRAPHIC B D
)N THE MODEL WITH ENVIRONMENTAL STOCHASTICITY ONLY POPULATIONS ARE ABLE TO
INVADE IF THE POPULATION GROWTH RATE E XCEEDS A CRITICAL VALUE 4HE GREATER THE
20 &RECKLETON 0- $OWLING AND .+ $ULVY
DIF FERENCE BETWEEN THE LONG TERM AVERAGE AND THIS CRITICAL VALUE THE MORE LIKELY A
SPECIES IS TO INVADE /N THE OTHER HAND IF A POPULATION IS SUBJECT TO DEMOGRAPHIC
STOCHASTICITY POPULATIONS MUST EXCEED A CRITICAL DENSITY BEFORE INVASION CAN
OCCUR 4HEREFORE THE TWO FORMS OF STOCHASTICITY HAVE RATHER DIFFERENT IMPLICA
TIONS FOR POPULATION DYNAMICS AND PREDICTING INVASION REQUIRES THAT BOTH OF
THESE SOURCES OF STOCHASTICITY ARE UNDERSTOOD IN DETAIL )N NON INVASIVE SPECIES
ESTIMATES OF DEMOGRAPHIC AND ENVIRONMENTAL STOCHASTICITY FROM BIRD POPULATIONS
SUGGEST THAT BOTH SOURCES MAY BE EXTREMELY SIGNIlCANT AND HENCE THAT THE TWO
FORMS NEED TO BE MEASURED SEPARATELY 3AETHER ET AL
)MPLICATIONS OF STOCHASTIC MODELS FOR MODELLING INVASIONS
4HE MAIN CONCLUSION FROM THE EXAMPLE MODELS OUTLINED ABOVE IS THAT IN A STO
CHASTIC ENVIRONMENT THE LONG
TERM POPULATION GROWTH IS NOT A LINEAR FUNCTION OF
THE AVERAGE POPULATION GROWTH RATES #OHEN ,EWONTIN AND #OHEN
4ULJAPURKAR .OT ONLY IS IT IMPORTANT TO ESTIMATE THE AMOUNT OF VARI
ABILITY IN POPULATION GROWTH RATES &RECKLETON AND 7ATKINSON BUT ALSO
TO DISENTANGLE THE RELATIVE ROLES OF DEMOGRAPHIC AND ENVIRONMENTAL VARIATION
4HIS REQUIRES THAT TEMPORALLY REPLICATED MEASURES OF POPULATION GROWTH OR THE
COMPONENTS OF POPULATION GROWTH ARE AVAILABLE 7ITHOUT SUCH INFORMATION IT IS
NOT POSSIBLE TO PREDICT WHICH SPECIES ARE CAPABLE OF INVADING AND WHICH ARE NOT
OR CONDITIONS UNDER WHICH A GIVEN SPECIES WILL BECOME INVASIVE OR NOT OR TO
DERIVE REALISTIC MODELS
5NFORTUNATELY SUCH INFORMATION IS RARELY AVAILABLE FOR MANY INVASIVE SPECIES
OFTEN BECAUSE INVASIVE SPECIES HAVE RECENTLY BEEN INTRODUCED TO NOVEL ENVIRON
MENTS #ONSEQUENTLY LONG
TERM DATASETS TEND TO BE RARE 5NDER SUCH CIRCUM
STANCES THERE ARE TWO CHOICES FOR THE MODELLER EITHER TO GIVE UP OR TO TRY TO MAKE
DO WITH WHATEVER INFORMATION IS AVAILABLE !S AN EXAMPLE OF THE LATTER 2EES AND
0AYNTER DEVELOPED A MODEL FOR THE DYNAMICS AND CONTROL OF INVASIVE
POPULATIONS OF 3COTCH "ROOM 4HIS MODEL WAS PARAMETERISED USING A SEVERAL DATA
SOURCES FROM BOTH THE NATIVE AND EXOTIC RANGE OF THE SPECIES AS WELL AS AN UNDER
STANDING OF ITS BASIC AUTECOLOGY )T SEEMS LIKELY HOWEVER THAT MANY EXISTING
MODELS ARE COMPROMISED TO SOME EXTENT IN THEIR PREDICTIVE ABILITY THROUGH A LACK
OF DATA ON STOCHASTIC VARIATION IN KEY PARAMETERS
2ECENT WORK HAS SUGGESTED THAT THE DEMOGRAPHIC AND ENVIRONMENTAL COMPO
NENTS OF POPULATION GROWTH MAY BE RELATED TO LIFE HISTORY TRAITS IN BIRDS 3AETHER
ET AL &OR EXAMPLE DEMOGRAPHIC STOCHASTICITY TENDS TO BE LARGER IN
SPECIES WITH GREATER ADULT SURVIVAL AGE AT MATURITY AND GENERATION TIMES 3AETHER
ET AL 3IMILARLY THE EXPECTED TIME TO EXTINCTION OF POPULATIONS INCREASED
WITH DECREASING CLUTCH SIZE 4HESE RESULTS ARE IMPORTANT BECAUSE THEY SHOW THAT
IT IS POSSIBLE TO GENERALIZE ABOUT HOW SENSITIVE POPULATIONS ARE TO DIFFERENT FORMS
OF VARIABILITY BASED ON LIFE
HISTORY TRAITS 4O DATE THERE HAVE BEEN NO COMPARABLE
ANALYSES OF THE DEMOGRAPHICS OF POPULATION DURING THE INITIAL STAGES OF INVASION
HOWEVER SUCH ANALYSIS COULD BE ENORMOUSLY REVEALING
3TOCHASTICITY NONLINEARITY AND INSTABILITY IN BIOLOGICAL INVASIONS
$%4%2-).)34)# 0/05,!4)/. $9.!-)#3
)F LEFT UNCHECKED THE SYSTEM DElNED BY EQUATION WILL ULTIMATELY BECOME EXTINCT
,EWONTIN #OHEN )F THE VALUE OF h IS GREATER THAN ONE THE EXPECTED
MEAN POPULATION SIZE AS 4 BECOMES LARGE IS INlNITE ALTHOUGH COUNTERINTUITIVELY
THE PROBABILITY OF EXTINCTION REMAINS EQUAL TO ONE ALBEIT WITH A VERY LONG EXPECTED
TIME TO EXTINCTION 4HIS BEHAVIOUR DOES NOT SIT WELL WITH ECOLOGICAL INTUITION
#OMMON OBSERVATION APPEARS TO INDICATE THAT THE DYNAMICS OF MANY SPECIES
ARE RELATIVELY INVARIANT 3OME INVASIVE SPECIES DO EXHIBIT INTRINSICALLY COMPLEX
POPULATION DYNAMIC BEHAVIOUR PARTICULARLY THE VIRUSES THAT CAUSE DISEASES SUCH
AS MEASLES AND mU %ARN ET AL #YCLIC DYNAMICS ARE REASONABLY COMMON
IN A RANGE OF TAXA +ENDALL ET AL 'INZBURG #OLYVAN HOWEVER
THESE TEND TO BE EXTRINSICALLY DRIVEN OR THE CONSEQUENCE OF SPECIES INTERACTIONS
SUCH AS PREDATOR PREY DYNAMICS RATHER THAN INTRINSIC PROPERTIES OF POPULATIONS
! GREAT DEAL OF EFFORT HAS BEEN EXPENDED ON UNDERSTANDING lRSTLY WHY POPU
LATIONS DO NOT SIMPLY BOOM OR BUST IN ACCORD WITH THE MOST SIMPLE DENSITY
INDEPENDENT MODELS AND SECONDLY WHY DIFFERENT SYSTEMS EXHIBIT ALTERNATIVE
KINDS OF POPULATION DYNAMICS
$ENSITY DEPENDENCE
$ENSITY
DEPENDENCE OCCURS WHEN ANY VITAL RATE CHANGES SYSTEMATICALLY WITH DEN
SITY 3TRICTLY SPEAKING THIS COULD INCLUDE A RANGE OF POPULATION LEVEL PHENOMENA
2OYAMA BUT TYPICALLY THIS DElNITION IS TAKEN TO IMPLY THAT SOME PER CAPITA
RATE SUCH AS MORTALITY OR FECUNDITY VARIES SYSTEMATICALLY WITH DENSITY $ENSITY
DEPENDENCE OF THIS SORT MAY RESULT FROM A RANGE OF ECOLOGICAL PROCESSES 4HE MOST
COMMONLY RECOGNISED FORM IS NEGATIVE DENSITY
DEPENDENCE WHICH RESULTS FROM
DIRECT COMPETITION FOR RESOURCES .ICHOLSON SUCH AS FOOD SHELTER OR MATES
-ORE INDIRECTLY DENSITY
DEPENDENCE MAY ARISE FROM THE IMPACT OF OTHER SPECIES
FOR INSTANCE PREDATORS OR PARASITES WHICH MAY INCREASE MORTALITY RATES AT HIGH
PREY OR HOST DENSITIES RESPECTIVELY
!S NOTED ABOVE DENSITY
INDEPENDENT MODELS SUGGEST THAT DENSITY
DEPENDENCE
SHOULD BE HIGHLY PREVALENT BECAUSE IN THE ABSENCE OF DENSITY
DEPENDENCE
POPULATIONS ARE LIKELY TO EITHER EXPAND WITHOUT LIMIT OR BECOME EXTINCT $ESPITE
THIS MANY MODELS FOR POPULATION DYNAMICS IGNORE DENSITY
DEPENDENCE AND THIS
CAN SEVERELY COMPROMISE MODEL PREDICTIONS EG SEE &RECKLETON ET AL
4HIS PROBLEM MAY BE ESPECIALLY ACUTE FOR INVASIVE SPECIES WHICH MIGHT BE EXPECT
ED TO HAVE A HIGH AVERAGE VALUE OF h AND HENCE BE CAPABLE OF RAPIDLY GROWING TO
HIGH DENSITIES )N THE INITIAL STAGES OF AN INVASION THE STOCHASTIC FACTORS THAT DETER
MINE h SEEM LIKELY TO PLAY A KEY ROLE IN DETERMINING INVASION SUCCESS (OWEVER
AS SHOWN BELOW FOLLOWING SUCCESSFUL INVASION THE ULTIMATE SIZE REACHED BY
THE INVADER POPULATION WILL BE IN LARGE PART DETERMINED BY THE FORM NATURE
AND STRENGTH OF DENSITY
DEPENDENCE
20 &RECKLETON 0- $OWLING AND .+ $ULVY
-ODELS FOR DENSITY DEPENDENT POPULATION GROWTH
! SUITE OF MODELS HAVE BEEN EMPLOYED TO MODEL THE DYNAMICS OF SPECIES GROWING
IN A DENSITY
DEPENDENT FASHION 4HE BEST KNOWN OF THESE IS THE LOGISTIC MODEL HERE
THE DISCRETE TIME VERSION
. T .T R.T n .+
)N THIS FORM THE LOGISTIC MODEL IS A POOR MODEL FOR DESCRIBING ECOLOGICAL DYNAM
ICS 4HERE ARE A NUMBER OF PROBLEMS WITH THIS MODEL INCLUDING THE EQUILIBRIUM IS
INDEPENDENT OF THE lNITE RATE OF INCREASE DENSITY
DEPENDENCE IS LINEAR AND STABIL
ITY BEHAVIOUR IS DETERMINED SOLELY BY THE MAXIMUM RATE OF INCREASE R SEE BELOW
$ESPITE THIS THE LOGISTIC MODEL IS WIDELY EMPLOYED IN THEORETICAL MODELLING PROB
ABLY BECAUSE IT IS ANALYTICALLY VERY CONVENIENT TO WORK WITH
-ORE REALISTIC MODELS HAVE BEEN SUGGESTED INCLUDING MODIlCATIONS TO THE LOGIS
TIC IN ORDER TO MAKE IT MORE REALISTIC EG THE THETA
LOGISTIC OF ,ANDE ET AL
/NE PARTICULARLY USEFUL MODEL IS THAT OF (ASSELL
. T h.T ; A.T = nB
4HIS MODEL HAS TWO PARAMETERS THAT MODEL DIFFERENT ASPECTS OF THE DENSITY
DEPENDENT RESPONSE A DETERMINES THE DENSITY AT WHICH THE PER CAPITA EFFECTS OF
DENSITY
DEPENDENCE BECOME SIGNIlCANT WHILST B VARIES THE RATE OF CHANGE IN POPU
LATION GROWTH RATE AS DENSITY BECOMES HIGH AND MAY BE RELATED TO THE NATURE OF
COMPETITION CONTEST VERSUS SCRAMBLE (ASSELL OR THE EFlCIENCY WITH WHICH
RESOURCES ARE CONVERTED INTO POPULATION GROWTH &IRBANK AND 7ATKINSON
&RECKLETON AND 7ATKINSON 4HE EQUILIBRIUM OF THIS MODEL IS GIVEN BY
. hB n A
4HE IMPORTANT ADVANCE OVER THE LOGISTIC MODEL IS THAT THE EQUILIBRIUM IS A
COMPOUND OF THREE MODEL PARAMETERS )NCREASING THE lNITE RATE OF INCREASE OR
DECREASING A LEADS TO INCREASES IN POPULATION DENSITY 4HESE EFFECTS ARE THEN
MODULATED BY B WHICH ALSO DETERMINES THE STABILITY PROPERTIES OF THE MODEL
)T SEEMS LIKELY THAT THE THREE PARAMETERS MAY BE RELATED TO EACH OTHER AT A
NUMBER OF SCALES 7ITHIN POPULATIONS TEMPORAL VARIABILITY IN THE ENVIRONMENT
MAY LEAD TO VARIATION IN ANY OF THE PARAMETERS !S POINTED OUT BY #HESSON AND
(UNTLEY UNDERSTANDING HOW THE EFFECTS OF STOCHASTICITY INTERACT
WITH THE STRENGTH OF COMPETITION IS THE KEY TO PREDICTING THE OUTCOME OF COMPETI
TION IN STOCHASTIC ENVIRONMENTS )N THE MODEL ABOVE THE PROBLEM IS TO UNDERSTAND
HOW THE PARAMETERS A AND h COVARY WITH EACH OTHER )F h AND A ARE INDEPENDENT
THEN POPULATION GROWTH RATES ARE TERMED ADDITIVE &IG A WITH THE CONSE
QUENCE THAT STOCHASTIC ENVIRONMENTAL EFFECTS IMPACT DIRECTLY ON POPULATION SIZES
&IG C )N CONTRAST SUB
ADDITIVE POPULATION GROWTH RATES OCCUR WHEN h AND
3TOCHASTICITY NONLINEARITY AND INSTABILITY IN BIOLOGICAL INVASIONS
&IG )NTERACTIONS BETWEEN DENSITY DEPENDENCE AND STOCHASTICITY ,INES REPRESENT
POPULATION GROWTH UNDER DIFFERENT ENVIRONMENTAL CONDITIONS A !DDITIVE POPULATION
GROWTH RATES B SUB ADDITIVE POPULATION GROWTH RATES C A POPULATION SUBJECT TO STO
CHASTICITY AND ADDITIVE POPULATION GROWTH D A POPULATION SUBJECT TO STOCHASTICITY AND
SUB ADDITIVE POPULATION GROWTH
A ARE CORRELATED WITH EACH OTHER &IG A )N SUCH POPULATIONS THE STRENGTH OF
COMPETITION CHANGES SO AS TO BUFFER POPULATION GROWTH FROM STOCHASTIC ENVIRON
MENTAL CHANGES 4HE CONSEQUENCE IS THAT POPULATIONS SUBJECT TO SUB
ADDITIVE
POPULATION GROWTH ARE BUFFERED AGAINST ENVIRONMENTAL CHANGE &IG D SHOWS
A POPULATION SUBJECT TO PERTURBATIONS IDENTICAL TO THOSE IN &IG C HOWEVER THE
SUB
ADDITIVE POPULATION GROWTH RATES CLEARLY HAVE AN IMPORTANT BUFFERING EFFECT
&ROM THE POINT OF VIEW OF PREDICTING THE DYNAMICS OF INVASIVE SPECIES THE PAT
TERNS SHOWN IN &IGS A AND B HAVE TWO IMPORTANT CONSEQUENCES &IRST THE FORM OF
DENSITY
DEPENDENCE AND ADDITIVITY INmUENCES THE EFFECTS OF STOCHASTICITY 3ECOND
THE FORM OF ADDITIVITY CAN INmUENCE THE EFFECTS OF CHANGES TO MANAGEMENT
&OR INSTANCE IF POPULATION GROWTH RATES ARE SUB
ADDITIVE THEN REDUCTIONS IN h
20 &RECKLETON 0- $OWLING AND .+ $ULVY
RESULTING FROM ATTEMPTS AT CONTROL OR ERADICATION WILL BE COMPENSATED FOR BY
REDUCTION IN THE STRENGTH OF DENSITY
DEPENDENCE YIELDING NO NET EFFECT ON POPULA
TION NUMBERS 4HUS IT IS NOT SUFlCIENT SIMPLY TO DOCUMENT THE EFFECTS OF STOCHAS
TICITY OR DENSITY
DEPENDENCE IN ISOLATION BUT THE INTERACTION BETWEEN THESE ALSO
NEEDS TO BE UNDERSTOOD )N PRACTICE THIS IS VERY DIFlCULT AS THIS REQUIRES THAT THE
FORM OF DENSITY
DEPENDENCE IS MEASURED UNDER A RANGE OF ENVIRONMENTAL CONDI
TIONS WHICH IS HIGHLY DATA INTENSIVE
.ON LINEAR DYNAMICS AND STABILITY
!S NOTED ABOVE IN THE MID S IT WAS FOUND THAT SIMPLE ECOLOGICAL MODELS COULD
YIELD UNEXPECTEDLY COMPLICATED PATTERNS OF POPULATION DYNAMICS -AY
-AY AND /STER &IG A B )N SHORT IN MODELS FOR DENSITY DEPENDENT
POPULATION GROWTH FOR ORGANISMS WITH DISCRETE GENERATIONS DENSITY DEPENDENCE
AT HIGH DENSITY MAY OVERCOMPENSATE FOR CHANGES IN DENSITY WITH THE RESULT THAT
INCREASES IN POPULATION SIZE ABOVE THE DETERMINISTIC EQUILIBRIUM TEND TO BE FOL
LOWED BY DISPROPORTIONATELY LARGE DECREASES 4HE CONSEQUENCE OF THIS IS CYCLIC
DYNAMICS OR MORE COMPLEX BEHAVIOUR SUCH AS LIMIT CYCLES AND CHAOS 5NDER SUCH
CIRCUMSTANCES POPULATIONS EXHIBIT SUSTAINED mUCTUATIONS IN POPULATION SIZE THAT
ARE APPARENTLY RANDOM AND SUPERlCIALLY SIMILAR TO THE mUCTUATIONS IN POPULATION
SIZES THAT ARE SEEN IN REAL POPULATIONS
%XTENSIVE ATTEMPTS HAVE BEEN MADE TO lND EXAMPLES OF POPULATION EXHIBITING
CHAOTIC DYNAMICS (OWEVER FEW EXAMPLES EXIST )N FACT IN SOME CASES THERE ARE
GOOD REASONS TO EXPECT THAT CHAOTIC DYNAMICS ARE RATHER UNLIKELY )N PLANTS FOR
EXAMPLE THE ABSENCE OF REPRODUCTIVE THRESHOLDS THE PRESENCE OF A SEEDBANK AND
ASYMMETRIC COMPETITION LEADING TO COMPENSATING DENSITY RESPONSES ALL LEAD TO
GENERALLY STABLE POPULATION DYNAMICS 2EES AND #RAWLEY &RECKLETON
AND 7ATKINSON )MPORTANTLY THIS MULTIPLICITY OF STABILITY GENERATING
MECHANISMS MEANS THAT EVEN WHEN ONE CONDITION FOR INSTABILITY HOLDS OTHERS
TEND TO CANCEL THIS OUT EG "UCKLEY ET AL -OREOVER GENERAL REVIEWS OF
THE STABILITY PROPERTIES OF NATURAL POPULATIONS APPEAR TO SHOW THAT MOST SINGLE
SPECIES POPULATIONS SHOW STABLE DYNAMICS (ASSELL ET AL ,ONSDALE
&RECKLETON AND 7ATKINSON !S NOTED ABOVE IN THE MAJORITY OF DOCUMENTED
CASES CYCLIC DYNAMICS TEND TO BE DRIVEN BY EXOGENOUS FACTORS OR INTERACTIONS WITH
OTHER SPECIES EG SEE +ENDALL ET AL
0OSITIVE DENSITY DEPENDENCE AND INSTABILITY
4HE MODELS DESCRIBED ABOVE EMPHASISE THE NEGATIVE EFFECTS OF DENSITY
DEPENDENCE
ON POPULATION GROWTH RATES (OWEVER RECENT REVIEWS HAVE POINTED OUT THAT NOT
ALL DENSITY
DEPENDENT EFFECTS ON POPULATION GROWTH ARE OF THIS FORM 3TEPHENS
AND 3UTHERLAND 3TEPHENS ET AL 0OSITIVE DENSITY
DEPENDENCE RATHER
CONFUSINGLY TERMED hINVERSEv DENSITY
DEPENDENCE BY SOME AUTHORS IS MOST WELL
KNOWN THROUGH !LLEE EFFECTS !LLEE EFFECTS ARISE MOST COMMONLY IN SOCIAL SPECIES
3TOCHASTICITY NONLINEARITY AND INSTABILITY IN BIOLOGICAL INVASIONS
&IG 3TABLE AND UNSTABLE EQUILIBRIA RESULTING FROM !LLEE EFFECTS 4HE RELATIONSHIP
BETWEEN POPULATION GROWTH RATE AND DENSITY IS HUMPED 7HERE THE CURVE CUTS THE LINE OF
ZERO POPULATION GROWTH RATE THERE IS AN EQUILIBRIUM !S INDICATED BY THE ARROWS THE UPPER
EQUILIBRIUM IS STABLE WHEREAS THE LOWER ONE IS UNSTABLE
WHERE COOPERATION BETWEEN INDIVIDUALS BREAKS DOWN AT LOW DENSITIES LEADING
TO A DECLINE IN SOME ASPECTS OF lTNESS WITH DECREASING DENSITIES )N THE EXTREME
SUCH COMPONENT !LLEE EFFECTS WHICH AFFECT ONE COMPONENT OF lTNESS MAY BE
MANIFEST AT THE LEVEL OF POPULATION GROWTH LEADING TO DEMOGRAPHIC !LLEE EFFECTS
IE ARE SO IMPORTANT THAT THEY IMPINGE ON RATES OF POPULATION CHANGE 3TEPHENS
AND 3UTHERLAND 4HE IMPORTANCE OF !LLEE EFFECTS IN POPULATION GROWTH IS
THAT THESE CAN LEAD TO A THRESHOLD DENSITY BELOW WHICH POPULATIONS CANNOT PERSIST
SEE &IG 4HIS IS OF COURSE HIGHLY SIGNIlCANT FOR THE DYNAMICS OF INVASIVE SPECIES
SINCE INVASIONS REQUIRE THAT SPECIES ARE CAPABLE OF ARRIVING IN NEW ENVIRONMENTS
AT LOW DENSITIES AND THEN SUBSEQUENTLY INCREASING 4HIS CAN HAVE IMPORTANT IMPLI
CATIONS FOR RATES OF SPATIAL SPREAD ,EWIS AND +AREIVA AS WELL AS RESPONSES
OF POPULATIONS TO STOCHASTIC VARIABILITY
,IEBHOLD AND "ASCOMPTE EXPLORED THE CONSEQUENCES OF !LLEE EFFECTS
AND STOCHASTICITY FOR THE CONTROL OF INVASIVE SPECIES 4HE MOST IMPORTANT OUTCOME
OF !LLEE EFFECTS IN THEIR MODELS WAS THAT !LLEE EFFECTS CAN INmUENCE THE OUTCOMES
OF ERADICATION PROGRAMMES &REQUENTLY IT IS FOUND THAT CLOSE TO ERADICATION
IS REQUIRED FOR CONTROL PROGRAMMES TO BE EFFECTIVE "ECAUSE SUCH HIGH LEVELS OF
CONTROL ARE GENERALLY DIFlCULT TO ACHIEVE THIS MAY PROHIBIT ATTEMPTS AT CONTROL
(OWEVER !LLEE EFFECTS MAY LEAD TO EXTINCTION THRESHOLDS AT LOW DENSITIES WITH
20 &RECKLETON 0- $OWLING AND .+ $ULVY
THE RESULT THAT LOWER RATES OF CONTROL MAY YIELD ERADICATION ,IEBHOLD AND
"ASCOMPTE STUDIED POPULATIONS OF THE 'YPSY -OTH ,YMANTIA DISPER IN
WHICH THEY FOUND THAT ERADICATION COULD BE ACHIEVED WITH C MORTALITY IN A
MODEL FOR THE POPULATION WHICH INCLUDED AN !LLEE EFFECT WHEREAS IN THE ABSENCE
OF THE !LLEE EFFECT CLOSE TO MORTALITY WOULD BE REQUIRED )N THE CASE STUDY
BELOW WE DESCRIBE IN MORE DETAIL A SYSTEM IN WHICH !LLEE EFFECTS ARE EXTREMELY
IMPORTANT IN EXPLAINING INVASIONS
-ULTISPECIES INTERACTIONS
4HE MAIN FOCUS ON THIS REVIEW SO FAR HAS BEEN ON SINGLE SPECIES DYNAMICS
(OWEVER INTERACTIONS BETWEEN SPECIES CAN AFFECT THE STABILITY OF POPULATION
DYNAMICS )NDEED MOST SPECIES DO NOT LIVE IN ISOLATION SO THIS POSSIBILITY MAY BE
QUITE COMMON AND WE REVIEW TWO EXAMPLES BELOW
)N AN INmUENTIAL PAPER -AY SUGGESTED THAT INSTABILITY MAY BE A COM
MON PROPERTY OF MANY FORMS OF ECOLOGICAL SYSTEMS (E SHOWED THAT THE SAME
MODELLING FRAMEWORK COULD BE APPLIED TO PREDATOR
PREY HARVESTED AND OR INDEED
ANY CONSUMER
RESOURCE SYSTEM 4HIS EFFECT ARISES AS A CONSEQUENCE OF THE FUNC
TION RESPONSE OF THE CONSUMER 7HEN CONSUMERS ARE AT LOW DENSITIES THE RATE OF
CONSUMPTION OF RESOURCES INCREASES WITH INCREASING RESOURCE DENSITY WITH THE
CONSEQUENCE THAT THE RATE OF RESOURCE LOSS IS POSITIVELY RELATED TO RESOURCE DEN
SITY !T LOW DENSITIES CONSUMERS CAN POTENTIALLY ERADICATE THEIR RESOURCES AT LOW
DENSITIES "Y CONTRAST AT HIGH RESOURCE DENSITIES RESOURCES BECOME SATURAT
ING AND EFFECTIVELY THE RATE OF RESOURCE LOSS TO CONSUMERS IS NEGATIVELY RELATED
TO RESOURCE DENSITY AT HIGH DENSITIES OF RESOURCES 4HIS DUAL BEHAVIOUR LEADS TO
THE POTENTIAL FOR TWO EQUILIBRIA ONE AT LOW RESOURCE DENSITIES THE OTHER AT HIGH
RESOURCE DENSITIES )F IT EXISTS THE EQUILIBRIUM AT LOW DENSITIES IS TYPICALLY UNSTABLE
2ECENTLY 'ASCOIGNE AND ,IPCIUS HAVE EXTENDED THIS FRAMEWORK TO CONSIDER
THE AGGREGATIVE RESPONSES OF RESOURCES AND HAVE EMPHASISED THAT SUCH BEHAVIOUR
HAS BEEN LARGELY IGNORED IN THE APPLIED AND CONSERVATION LITERATURE AND SHOW THAT
THERE ARE A GROWING NUMBER OF SYSTEMS THAT IMPLICATE SUCH A MECHANISM
#OMPETITIVE INTERACTIONS MAY ALSO YIELD INSTABILITIES IN TWO SOMEWHAT DIFFERENT
WAYS &IRST SIMPLE MODELS FOR THE DYNAMICS OF TWO COMPETING SPECIES SHOW THAT
THREE DISTINCT OUTCOMES MAY OCCUR I ONE SPECIES OR ANOTHER EXCLUDES THE OTHER
II THE COMPETING SPECIES SETTLE AT A STABLE JOINT EQUILIBRIUM III THERE IS A JOINT
EQUILIBRIUM BUT WHICH IS UNSTABLE SO THAT PERTURBATIONS FROM THE JOINT EQUILIB
RIUM LEAD TO THE SYSTEM BECOMING DOMINATED BY ONE SPECIES OR ANOTHER 4HIS LATTER
UNSTABLE CONDITION MAY ARISE IF THE PER CAPITA EFFECTS OF INTER
SPECIlC COMPETITION
ARE GREATER THAN THE PER CAPITA EFFECTS OF INTRA
SPECIlC COMPETITION )N THE CASE
STUDIES BELOW WE DETAIL SUCH AN EXAMPLE
3ECOND COMPETITIVE INTERACTIONS MAY YIELD UNSTABLE DYNAMICS WHEN MULTI
SPECIES SYSTEMS IN INTERACTIONS ARE STRONG ARE SUBJECTED TO PERTURBATIONS 4ILMAN
ET AL 4HE REASON FOR THIS IS THAT IN SUCH A MIXTURE THE DYNAMICS OF ANY SIN
GLE SPECIES ARE A COMPLEX FUNCTION OF THE DENSITIES OF ALL THE SPECIES #ONSEQUENTLY
3TOCHASTICITY NONLINEARITY AND INSTABILITY IN BIOLOGICAL INVASIONS
THE RESPONSE OF THE SYSTEM TO PERTURBATION MAY BE COMPLEX AND PERTURBATION MAY
LEAD TO COMPLEX INTERMEDIATE OR TRANSIENT DYNAMICS 4HE SIGNIlCANCE OF THIS IS
THAT FOLLOWING A CHANGE SUCH AS THE INVASION OF A NEW SPECIES THE ENTIRE SYSTEM
MAY TAKE MANY GENERATIONS TO RETURN TO EQUILIBRIUM
#!3% 345$)%3
4HE PREVIOUS SECTIONS HAVE EMPHASISED THAT PREDICTING THE DYNAMICS OF INVASIVE
SPECIES MAY REQUIRE THAT A SUITE OF PROCESSES ARE DIS
ENTANGLED 4HESE INCLUDE THE
EFFECTS OF STOCHASTICITY AS WELL AS THE STRENGTHS OF INTRA
AND INTER
SPECIlC INTERAC
TIONS IN THE WIDEST SENSE &OR INVASIVE SPECIES THE KEY QUESTIONS ARE HOW THESE
PROCESSES AFFECT THE INVASION OF POPULATIONS INTO NEW ENVIRONMENTS AS WELL AS
HOW THEY DETERMINE POPULATION SIZE AND THE OUTCOME OF CONTROL STRATEGIES
)N THIS SECTION WE REVIEW IN DETAIL TWO CASE STUDIES OF INVASIVE SPECIES WHICH
HAVE BEEN STUDIES USING SIMPLE MODELLING APPROACHES .ECESSARILY IT IS IMPOSSIBLE
TO CONSIDER ALL OF THE PROCESSES DESCRIBED ABOVE 4HE lRST EXAMPLE ILLUSTRATES THE
KEY ROLE THAT EXTREME STOCHASTICITY MAY PLAY IN DETERMINING INVASION SUCCESS AND
HOW THIS MAY BE MODULATED BY THE EFFECTS OF COMPETITION 4HE SECOND EXAMPLE
SHOWS THE EFFECT OF PERTURBATION ON A PREDATOR
PREY SYSTEM WITH AN INSTABILITY
6ULPIA BROMOIDES IN !USTRALIAN PASTURES THE INTERPLAY OF STOCHASTICITY
AND INSTABILITY
4HIS EXAMPLE SHOWS HOW INSTABILITY AND STOCHASTICTY SIMULTANEOUSLY PLAY ROLES
IN DETERMINING WHETHER A SPECIES INVADES A SYSTEM 6 BROMOIDES IS A WINTER
ANNUAL WHICH ORIGINATES FROM THE -EDITERRANEAN AND HAS BECOME A PROBLEM IN
ANNUAL PASTURE SYSTEMS IN !USTRALIA )N CONTRAST IN 3PAIN AND 0ORTUGAL WHERE IT
OCCURS NATURALLY IN PASTURES 6 BROMOIDES IS NOT REGARDED AS A PROBLEM SPECIES
6 BROMOIDES WAS PROBABLY INTRODUCED TO !USTRALIA AS A CONTAMINANT OF SEED OR
FORAGE 3EVERAL SPECIES OF THE GENUS 6ULPIA WERE INTRODUCED TO !USTRALIA 7E
STUDIED THE INVASION DYNAMICS OF ANNUAL PASTURES IN .37 IN WHICH 6BROMOIDES
IS THE COMMONEST OF THE SPECIES AND OCCURS IN C OF THE PASTURES WHERE THE
GENUS IS PRESENT $OWLING 'IVEN THAT THE 6ULPIA GENUS IS WELL ESTABLISHED
IN !USTRALIA THE CURRENT PROBLEM IS TO DETERMINE WHAT FACTORS ENABLE THE PERSIS
TENCE AND SPREAD OF THE SPECIES
6 BROMOIDES IS A PROBLEM BECAUSE ALTHOUGH IN THE EARLY STAGES OF THE GROWING
SEASON IT MAY PROVIDE SOME USEFUL FORAGE FOR LIVE
STOCK THE FORAGE PRODUCED LATE
IN THE SEASON IS OF VERY LOW NUTRITIONAL VALUE AND BECAUSE THE SEEDS PRODUCED
BY THE WEED BECOME TANGLED IN THE WOOL OF SHEEP AND MAY EVEN CAUSE PHYSICAL
INJURIES TO LIVE
STOCK ,ONG
TERM MANAGEMENT OF 6ULPIA IS DIFlCULT IN PASTURES
THAT ARE DOMINATED BY ANNUALS IE IN WHICH THE PERENNIAL COMPONENT IS LOW
4HE PROBLEM OF CONTROLLING 6ULPIA IS COMPOUNDED BY THE HIGH SEED PRODUCTION OF
THE SPECIES WHICH ALLOWS FAST POPULATION RECOVERY IN THE YEARS FOLLOWING HERBICIDE
20 &RECKLETON 0- $OWLING AND .+ $ULVY
APPLICATION AS WELL AS HIGH RATES OF COMPENSATORY GROWTH OF SURVIVORS FOLLOWING
HERBICIDAL CONTROL $OWLING ET AL
4HE MAIN QUESTIONS THAT ARE IMPORTANT IN UNDERSTANDING THE DYNAMICS OF
6 BROMOIDES AS WELL AS IN PREDICTING THE EFFECTS OF MANAGEMENT ARE I HOW DO
INTERACTIONS WITHIN AND BETWEEN SPECIES AFFECT LONG
TERM DYNAMICS II (OW
ARE POPULATION DYNAMICS AND LONG
TERM PERSISTENCE AFFECTED BY STOCHASTICITY
&RECKLETON ET AL DEVELOPED A MODEL TO PREDICT THE DYNAMICS OF THIS SPECIES
IN A MULTI
SPECIES PASTURE MIXTURE IN ORDER TO ADDRESS THESE QUESTIONS
4HE POPULATION DYNAMICS OF 6 BROMOIDES WERE STUDIED IN A
YEAR lELD EXPERI
MENT CONDUCTED BETWEEN AND !RTIlCIAL PASTURES WERE ESTABLISHED
COMPRISING THREE SPECIES 6 BROMOIDES TOGETHER WITH TWO @DESIRABLE SPECIES
4RIFOLIUM SUBTERRANEUM A LEGUME AND ,OLIUM RIGIDUM ANOTHER GRASS 0ASTURES
WERE SET UP ORIGINALLY WITH LOW MEDIUM AND HIGH DENSITIES OF 4RIFOLIUM LOW OR
ZERO DENSITIES OF ,OLIUM AND LOW OR HIGH DENSITIES OF 6ULPIA 4HESE ARTIlCIAL PAS
TURES WERE ALLOWED TO GROW UNTIL SPRING WHEN HALF OF THE PLOTS WERE SPRAYED
WITH HERBICIDE IN ORDER TO CREATE A FURTHER DENSITY DIFFERENTIAL OF 6ULPIA 4HE YEAR
WAS A DROUGHT YEAR AND WE ESTIMATED THAT DROUGHTS OF THIS SEVERITY
OCCUR EVERY YEARS OR SO &RECKLETON ET AL
7E USED THE DATA FROM THIS EXPERIMENT TO PARAMETERISE A MODEL PREDICTING THE
NUMBERS NUMBERS OF MATURE PLANTS . AND SEEDS IN THE SEEDBANK 3 OF EACH OF
THE THREE SPECIES IE
n
.I x T hI .I T -_IJ .JT
iI 3I T
J
n
3I x T hI .I T -_IJ .JT
iI 3I T
J
h AND h MEASURE THE PER CAPITA RATE OF CHANGE IN NUMBERS OF MATURE PLANTS AND
SEEDS RESPECTIVELY OWING TO SEED PRODUCTION AT THE VEGETATIVE STAGE p AND p
RESPECTIVELY MEASURE THE RECRUITMENT OF PLANTS FROM AND PERSISTENCE OF SEEDS
WITHIN THE SEED BANK COMPETITION BETWEEN ANY PAIR OF SPECIES I AND J IS MEASURED
BY A PARAMETER _IJ WHICH IS THE PER CAPITA REDUCTION IN POPULATION GROWTH .OTE
THAT WHEN I AND J ARE DIFFERENT _IJ REFERS TO INTER
SPECIlC COMPETITION IE COM
PETITION BETWEEN SPECIES BUT WHEN I J _ MEASURES INTRA
SPECIlC COMPETITION
IE COMPETITION WITHIN THE SPECIES &OR THE THREE SPECIES MIXTURE THEREFORE
THERE ARE NINE IE COMPETITION COEFlCIENTS
!NALYSIS OF THE lTTED MODEL INDICATED A CLEAR DIVISION WITHIN THIS SIMPLE COM
MUNITY &IRST COMPETITION BETWEEN 4RIFOLIUM AND THE TWO GRASSES WAS VERY WEAK
INDEED AND THERE WAS VERY LITTLE INmUENCE OF THE 4RIFOLIUM ON THE DYNAMICS OF
EITHER OF THE GRASSES OR OF THE GRASSES ON THE 4RIFOLIUM 4HIS IS PERHAPS UNSUR
3TOCHASTICITY NONLINEARITY AND INSTABILITY IN BIOLOGICAL INVASIONS
PRISING AS LEGUMES AND GRASSES REPRESENT RATHER DIFFERENT FUNCTIONAL GROUPS
/N THE OTHER HAND COMPETITION BETWEEN THE TWO GRASSES WAS VERY STRONG AND
THEY INmUENCED EACH OTHERS DYNAMICS PROFOUNDLY 3INCE NEITHER OF THE GRASSES
POSSESS VERY PERSISTENT SEEDBANKS IN CONTRAST TO THE 4RIFOLIUM THE DYNAMICS OF
THESE TWO SPECIES CAN BE MORE SIMPLY MODELLED BY A SIMPLE
SPECIES MODEL
., T h, ., T _,, .,T
_,6 .6 T n
.6 T h6 .6 T _66 .6T
_6, ., T n
4HE DYNAMICS OF THIS SYSTEM MAY BE ANALYSED USING A SIMPLE PHASE PLANE
ANALYSIS &IG 4HE lTTED COMPETITION COEFlCIENTS FOR EQUATIONS HAD THE
PROPERTY THAT IN BOTH MODELS THE INTER
SPECIlC COEFlCIENTS WERE LARGER THAN THE
INTRA
SPECIlC COEFlCIENTS !S A CONSEQUENCE THE EQUILIBRIUM FOR THE TWO SPECIES
IS PREDICTED TO BE UNSTABLE &IG A 4HE UNSTABLE EQUILIBRIUM HAS AN IMPORTANT
CONSEQUENCE FOR THE INVASION OF 6ULPIA 6ULPIA IS ONLY ABLE TO INVADE COMMUNITIES
WHEN THE DENSITY OF ,OLIUM IS LOW %XAMPLES OF THE PREDICTED DYNAMICS ARE SHOWN
IN &IG B AND C 6ULPIA IS UNABLE TO INVADE A HIGH DENSITY OF ,OLIUM &IG B IT IS
ABLE TO INVADE A LOW DENSITY &IG C
!T lRST SIGHT IT APPEARS THAT THE INVASION OF 6ULPIA INTO PASTURES MAY BE DIFlCULT
BECAUSE IT NEEDS TO EXCEED A CRITICAL DENSITY IN ORDER TO INVADE )F THE ENVIRONMENT
WERE CONSTANT THEN THIS WOULD BE THE CASE )N REALITY HOWEVER THE ENVIRONMENT
IS NOT CONSTANT !S NOTED ABOVE PERIODICALLY SEVERE DROUGHTS OCCUR 4HESE GREATLY
FACILITATE THE INVASION OF 6ULPIA BECAUSE THESE DROUGHTS REDUCE THE DENSITIES OF
ALL SPECIES TO LOW LEVELS &OLLOWING DROUGHT THE lNITE RATE OF INCREASE OF 6ULPIA IS
MUCH GREATER THAN THAT OF ,OLIUM h6 WHILST h, #ONSEQUENTLY 6ULPIA
POPULATIONS RESPOND MUCH FASTER FOLLOWING DROUGHTS THAN DO ,OLIUM POPULATIONS
AND HENCE THE INVASION OF 6ULPIA IS FACILITATED 7E ESTIMATED THAT SEVERE DROUGHTS
OCCURRING WITH A FREQUENCY OF EVERY IN TO IN YEARS WOULD GREATLY FACILI
TATE THE INVASION OF 6ULPIA 3UGGESTIVELY ANECDOTAL EVIDENCE SUGGESTS THAT 6ULPIA
BECAME NOTICED AS A PROBLEM FOLLOWING SEVERE DROUGHTS IN THE EARLY S
ALTHOUGH WE HAVE NO QUANTITATIVE DATA WITH WHICH TO SUPPORT THIS CONTENTION
)N SUMMARY UNDERSTANDING THE INVASION OF 6ULPIA INTO PASTURES AND ITS PER
SISTENCE REQUIRES THAT WE NOT ONLY UNDERSTAND THE DETAILS OF COMPETITION WITH
OTHER SPECIES BUT ALSO THE EFFECTS OF LONG
TERM STOCHASTICITY 3TOCHASTICITY PLAYS A
KEY ROLE BECAUSE THE UNSTABLE EQUILIBRIUM WOULD MAKE INVASION OF 6ULPIA UNLIKELY
FROM LOW DENSITIES INTO ESTABLISHED PASTURES (OWEVER THE EFFECTS OF COMPETITION
BECOME MODULATED BY PERIODIC DROUGHTS MAKING INVASION FAR MORE LIKELY
)NSTABILITY AND OUTBREAKS THE CROWN OF THORNS STARFISH
4HE SECOND EXAMPLE ILLUSTRATES HOW !LLEE EFFECTS RESULTING FROM PREDATOR
PREY
INTERACTIONS CAN LEAD TO UNSTABLE POPULATION DYNAMICS AND CONSEQUENTLY DETER
MINE WHETHER INVASIONS ARE POSSIBLE OR NOT 4HERE HAS BEEN A GREAT DEAL OF RECENT
20 &RECKLETON 0- $OWLING AND .+ $ULVY
&IG A 0HASE
PLANE DIAGRAM SHOWING UNSTABLE INTERACTIONS IN MIXTURES OF ,OLIUM
AND 6ULPIA B C &REQUENCY DEPENDENT INVASIONS OF COMMUNITIES BY EITHER ,OLIUM
lLLED CIRCLES OR 6ULPIA OPEN CIRCLES THE INVADER DEPENDS ON THE RELATIVE INITIAL DENSITIES
OF THE TWO SPECIES 4HE DYNAMICS OF THE THIRD SPECIES 4RIFOLIUM SQUARES ARE ESSENTIALLY
NEUTRAL WITH RESPECT TO THE OTHER TWO SPECIES
INTEREST IN SUCH SYSTEMS REVIEWED BY 'ASCOIGNE AND ,IPCIUS INSPIRED
MAINLY BY THE RESULTS OF SIMPLE MODELS )N BRIEF PREDATORS CAN GENERATE !LLEE
EFFECTS IN THE POPULATION DYNAMICS OF THEIR PREY AS A CONSEQUENCE OF THEIR FUNC
TIONAL AND NUMERICAL RESPONSES 4HIS HAPPENS IN SYSTEMS WHERE THE DYNAMICS
OF THE PREY POPULATION ARE DRIVEN MAINLY BY PREDATION AND WHEN PREY HAVE NO
REFUGES SPATIAL OR TEMPORAL FROM THE EFFECTS OF PREDATION !LLEE EFFECTS ARISE IN
SUCH SYSTEMS WHEN PREDATORS HONE IN ON PREY AT LOW DENSITIES SO THAT LOW DENSITY
POPULATIONS OF PREY TEND TO HAVE NEGATIVE POPULATION GROWTH RATES
3UCH PREDATOR INDUCED !LLEE EFFECTS ARE SIGNIlCANT FOR INVASIVE SPECIES BECAUSE
THE POTENTIAL EXISTS FOR PREDATORS TO ERADICATE UNWANTED INVASIVE PREY BY REDUCING
3TOCHASTICITY NONLINEARITY AND INSTABILITY IN BIOLOGICAL INVASIONS
THEM TO LOW DENSITIES !LTERNATIVELY IF PREY POPULATIONS CAN BE REDUCED BY CONTROL
MEASURES THE POTENTIAL EXITS FOR PREDATORS TO LEAD TO THE ULTIMATE EXTINCTION OF
THE PREY #ONVERSELY HOWEVER THE POSSIBILITY EXISTS THAT PREY POPULATIONS MAY
BE HELD IN CHECK BY PREDATORS WHICH MAINTAIN NEGATIVE PREY POPULATION GROWTH
RATES FROM LOW DENSITIES BUT THAT PERTURBATION TO EITHER POPULATION MAY LEAD TO
THE PREY POPULATION ESCAPING PREDATOR CONTROL
4HE SYSTEM WE WORKED WITH CONSISTS OF A
LEVEL FOOD WEB INVOLVING PREDATORY
lSHES THE INVASIVE CORAL
EATING CROWN
OF
THORNS STARlSH !CANTHASTER PLANCI AND
REEF
BUILDING CORALS $ULVY ET AL 4HE CROWN
OF
THORNS STARlSH FEEDS UPON
LIVE CORALS CAUSING THE LARGEST KNOWN PEST
RELATED DISTURBANCES ON )NDO
0ACIlC
CORAL REEFS AND IT IS REGARDED AS A MAJOR MANAGEMENT PROBLEM 3EVERAL MECHA
NISMS INCLUDING HYDROGRAPHY HURRICANE DISTURBANCE NUTRIENT INPUTS AND PREDA
TOR REMOVAL HAVE BEEN SUGGESTED AS POTENTIAL CAUSES OF STARlSH INVASIONS
7E STUDIED A SERIES OF OCEANIC ISLANDS VARYING IN SIZE AND HUMAN POPULA
TION &OR EACH ISLAND AN INDEX OF HUMAN lSHING INTENSITY WAS ESTIMATED !T EACH
OF THE SITES DENSITIES OF PREDATORY lSHES STARlSH AND BARRIER REEF COMMUNITIES WERE
SURVEYED BY DIVERS 3URVEYING WAS CONDUCTED IN SUCCESSIVE YEARS 7E WISHED
TO TEST THE HYPOTHESIS THAT STARlSH POPULATIONS OUTBREAK AND INVADE REEFS WHEN
PERTURBATIONS TO THEIR PREDATORS REMOVE THE PREDATOR CONTROL AND MOREOVER THAT
PREDATORS ARE RESPONSIBLE FOR GENERATING AN !LLEE EFFECT IN STARlSH POPULATIONS
7E DID NOT HAVE SUFlCIENTLY DETAILED DATA TO BE ABLE TO PARAMETERISE A FULL POPU
LATION MODEL (OWEVER WE WERE STILL ABLE TO USE A MODELING FRAMEWORK TO ANALYSE
THE DYNAMICS OF THIS SYSTEM &OR THE STARlSH WE CALCULATED PER
CAPITA RATES OF
POPULATION CHANGE .T n . T n .T FOR EACH ISLAND 4O TEST THE HYPOTHESIS
THAT PREDATORS WERE RESPONSIBLE FOR GENERATING !LLEE EFFECTS IN STARlSH POPULA
TIONS WE PLOTTED THE PER CAPITA RATE OF POPULATION CHANGE AGAINST STARlSH DENSITY
AND PREDATOR DENSITY AS WELL AS TO LOOK AT HOW PREDATOR DENSITIES ARE AFFECTED BY
HUMAN lSHING PRESSURE
&IRST WE FOUND THAT PREDATORY lSHES WERE LESS ABUNDANT AT THE MOST INTEN
SIVELY lSHED SITES AS THE LEAST lSHED SITES 4HIS INDICATES THAT THERE IS AN ENORMOUS
RANGE OF VARIATION IN THE PREDATORY lSHING PRESSURE IN THIS SYSTEM 'IVEN THIS THE
DATA ON RATES OF POPULATION GROWTH OF THE STARlSH &IG SUGGEST THAT PREDATOR
REMOVAL BY SUBSISTENCE EXPLOITATION MAY BE SUFlCIENT TO ALLOW OUTBREAKS OF THE
CROWN
OF
THORNS !S SHOWN IN &IG A THE RELATIONSHIP BETWEEN PER
CAPITA RATE OF
POPULATION CHANGE AND STARlSH DENSITY IS HUMPED AS WOULD BE EXPECTED IF POPULA
TIONS WERE SUBJECT TO AN !LLEE EFFECT 4HE DATA APPEAR TO INDICATE THAT THERE ARE TWO
EQUILIBRIA ONE AT LOW DENSITIES THE OTHER AT HIGH DENSITIES 4HE LOWER EQUILIBRIUM
WILL BE UNSTABLE )N A VARIABLE ENVIRONMENT LOW DENSITY POPULATIONS WILL EITHER
OUTBREAK OR WILL BECOME EXTINCT
4HAT THE !LLEE EFFECT SHOWN IN &IG A ACTUALLY RESULTS FROM THE EFFECTS OF PREDA
TORS IS CONlRMED IN &IG B !S SHOWN THERE IS A NEGATIVE RELATIONSHIP BETWEEN
THE RATE OF CHANGE IN STARlSH DENSITY AND PREDATOR DENSITY CONlRMING THAT STAR
lSH POPULATION DYNAMICS ARE DRIVEN BY PREDATION THUS FULlLLING THE CONDITIONS
REQUIRED FOR PREDATOR
INDUCED INSTABILITY
20 &RECKLETON 0- $OWLING AND .+ $ULVY
&IG 4HE RELATIONSHIP BETWEEN AVERAGE STARlSH DENSITY AND PER CAPITA POPULATION
GROWTH OF STARlSH AND A AVERAGE STARlSH DENSITY B AVERAGE DENSITY OF PREDATORY lSHES
$ATA ARE TAKEN FROM ISLANDS PRESENTED IN $ULVY ET AL
)N SUMMARY THIS SYSTEM SUGGESTS THAT INVASION OF THE CROWN OF THORNS STARlSH
MAY BE GOVERNED BY UNDERLYING UNSTABLE DYNAMICS AND HOW THIS INTERACTS WITH
VARIATIONS IN THE DENSITIES OF PREDATORS )N THIS CASE
STUDY VARIATION IN PREDA
TOR DENSITY WAS SPATIAL AND THE CONSEQUENCE OF DIFFERENCES IN lSHING INTENSITY
AMONG ISLANDS )N OTHER SYSTEMS VARIATION COULD BE THE RESULT OF OTHER FACTORS
SUCH AS HABITAT LOSS OR CLIMATIC EFFECTS 7HAT THIS EXAMPLE ILLUSTRATES HOWEVER
3TOCHASTICITY NONLINEARITY AND INSTABILITY IN BIOLOGICAL INVASIONS
IS THAT PREDICTING SUCH EFFECTS WILL REQUIRE A DETAILED UNDERSTANDING OF THE NATURE
OF POPULATION STABILITY
#/.#,5$).' 2%-!2+3
%XCEPT IN THE UNLIKELY EVENT THAT POPULATION DYNAMICS ARE COMPLETELY DENSITY
INDEPENDENT THE LONG
TERM DYNAMICS OF POPULATIONS WILL BE DETERMINED BY THE
EFFECTS OF VARIOUS FORMS OF INTRA
AND INTER
SPECIlC INTERACTIONS )N ADDITION TO
THIS THE EFFECTS OF DENSITY
INDEPENDENT STOCHASTICITY MAY BE COMPLEX PARTICU
LARLY WHEN POPULATIONS ARE FORCED TO LOW DENSITIES 4HIS MEANS THAT FORECASTING
THE OUTCOME OF INVASION MAY BE COMPLEX AND REQUIRES A DETAILED UNDERSTAND
ING OF THE STRUCTURE OF THE SYSTEM IN QUESTION 3IMPLE POPULATION MODELS CAN BE
AN EXTREMELY VALUABLE GUIDE IN ANALYSING SUCH PROBLEMS BOTH IN THEORY AND
IN PRACTICE
2%&%2%.#%3
!NDREWARTHA ( ' AND "IRCH , # 4HE DISTRIBUTION AND ABUNDANCE OF ANIMALS
5NIVERSITY OF #HICAGO 0RESS #HICAGO
"UCKLEY 9 - "RIESE $ 4 AND 2EES - A $EMOGRAPHY AND MANAGEMENT OF THE
INVASIVE PLANT SPECIES (YPERICUM PERFORATUM ) 5SING MULTI
LEVEL MIXED
EFFECTS MODELS
FOR CHARACTERIZING GROWTH SURVIVAL AND FECUNDITY IN A LONG
TERM DATA SET *OURNAL OF
!PPLIED %COLOGY
"UCKLEY 9 - "RIESE $ 4 AND 2EES - B $EMOGRAPHY AND MANAGEMENT OF
THE INVASIVE PLANT SPECIES (YPERICUM PERFORATUM )) #ONSTRUCTION AND USE OF AN INDI
VIDUAL
BASED MODEL TO PREDICT POPULATION DYNAMICS AND THE EFFECTS OF MANAGEMENT
STRATEGIES *OURNAL OF !PPLIED %COLOGY
"YERS * % 2EICHARD 3 2ANDALL * - 0ARKER ) - 3MITH # 3 ,ONSDALE 7 -
!TKINSON ) ! % 3EASTEDT 4 2 7ILLIAMSON - #HORNESKY % AND (AYES $
$IRECTING RESEARCH TO REDUCE THE IMPACT OF NON
INDIGENOUS SPECIES #ONSERVATION
"IOLOGY
#HESSON 0 , AND (UNTLY . #OMMUNITY CONSEQUENCES OF LIFE
HISTORY TRAITS IN A
VARIABLE ENVIRONMENT !NN :OOL &ENNICI
#HESSON 0 , AND (UNTLY . 3HORT
TERM INSTABILITIES AND LONG
TERM COMMUNITY
DYNAMICS 4RENDS IN %COLOGY AND %VOLUTION
#OHEN $ /PTIMIZING REPRODUCTION IN RANDOMLY VARYING ENVIRONMENTS *OURNAL OF
4HEORETICAL "IOLOGY
$OBSON ! 0 AND 2 - -AY 0ATTERNS OF )NVASIONS BY 0ATHOGENS AND 0ARASITES
4HE %COLOGY OF "IOLOGICAL )NVASIONS OF .ORTH !MERICA AND (AWAII 3PRINGER
6ERLAG .EW 9ORK
$OWLING 0 - 4HE ECOLOGY OF 6ULPIA 0LANT 0ROTECTION 1UARTELY 3UPPLEMENT
20 &RECKLETON 0- $OWLING AND .+ $ULVY
$OWLING 0 - 6ERBEEK " ,EMERLE $ AND ,EYS ! 2 2APID REGENERATION OF
6ULPIA 6ULPIA BROMOIDES IN PASTURES )N 0ROCEEDINGS TH !SIAN
0ACIlC 7EED 3OCIETY
#ONFERENCE PP
4OSUKUBA *APAN
$UFFY * % "IODIVERSITY LOSS TROPHIC SKEW AND ECOSYSTEM FUNCTIONING %COLOGY
,ETTERS
$ULVY . + &RECKLETON 2 0 AND 0OLUNIN . 6 # #ORAL REEF CASCADES AND
THE INDIRECT EFFECTS OF PREDATOR REMOVAL BY EXPLOITATION %COLOGY ,ETTERS
%ARN $ * $ 2OHANI 0 AND 'RENFELL " 4 0ERSISTENCE CHAOS AND SYNCHRONY IN
ECOLOGY AND EPIDEMIOLOGY 0ROCEEDINGS OF THE 2OYAL 3OCIETY 3ERIES "
%NGEN 3 "AKKE AND ! )SLAM $EMOGRAPHIC AND ENVIRONMENTAL STOCHASTIC
ITYnCONCEPTS AND DElNITIONS "IOMETRICS n
&IRBANK , ' AND 7ATKINSON ! 2 ! MODEL OF INTERFERENCE WITHIN PLANT POPULA
TIONS *OURNAL OF 4HEORETICAL "IOLOGY
&RECKLETON 2 0 AND 7ATKINSON ! 2 (OW DOES TEMPORAL VARIABILITY AFFECT PREDIC
TIONS OF WEED POPULATION NUMBERS *OURNAL OF !PPLIED %COLOGY
&RECKLETON 2 0 AND 7ATKINSON ! 2 !SYMMETRIC COMPETITION BETWEEN PLANT
SPECIES &UNCIONAL %COLOGY
&RECKLETON 2 0 AND 7ATKINSON ! 2 !RE WEED POPULATIONS CHAOTIC *OURNAL OF
!PPLIED %COLOGY
&RECKLETON 2 0 7ATKINSON ! 2 $OWLING 0 - AND ,EYS ! 2 $ETERMINANTS
OF THE ABUNDANCE OF INVASIVE ANNUAL WEEDS COMMUNITY STRUCTURE AND NON
EQUILIBRIUM
DYNAMICS 0ROCEEDINGS OF THE 2OYAL 3OCIETY 3ERIES "
&RECKLETON 2 0 3ILVA -ATOS $ - "OVI - , ! AND 7ATKINSON ! 2 0ROBLEMS
WITH PREDICTING THE IMPACTS OF HARVESTING USING STRUCTURED POPULATION MODELS AN
EXAMPLE BASED ON TROPICAL TREES *OURNAL OF !PPLIED %COLOGY
'ASCOIGNE * # AND ,IPCIUS 2 . !LLEE EFFECTS DRIVEN BY PREDATION *OURNAL OF
!PPLIED %COLOGY
'INZBURG , 2 AND #OLYVAN - %COLOGICAL ORBITS HOW PLANETS MOVE AND POPULA
TIONS GROW /XFORD 5NIVERSITY 0RESS /XFORD
(ASSELL - 0 $ENSITY
DEPENDENCE IN SINGLE SPECIES POPULATIONS *OURNAL OF !NIMAL
%COLOGY
(ASSELL - 0 $ETECTING DENSITY DEPENDENCE 4RENDS IN %COLOGY AND %VOLUTION
(ASSELL - 0 ,AWTON * ( AND -AY 2 - 0ATTERNS OF DYNAMICAL BEHAVIOUR IN
SINGLE
SPECIES POPULATIONS *OURNAL OF !NIMAL %COLOGY
(EINO - 2IPA * AND +AITALA 6 %XTINCTION RISK UNDER COLOURED ENVIRONMENTAL
NOISE %COGRAPHY
+ENDALL " % ET AL 7HY DO POPULATIONS CYCLE ! SYNTHESIS OF STATISTICAL AND MECHANISTIC
MODELLING APPROCHES %COLOGY
,ANDE 2 $EMOGRAPHIC STOCHASTICITY AND !LLEE EFFECT ON A SCALE WITH ISOTROPIC NOISE
/IKOS
,ANDE 2 %NGEN 3 AND 3AETHER "
% 3TOCHASTIC POPULATION DYNAMICS IN ECOLOGY
AND CONSERVATION /XFORD 5NIVSERSITY 0RESS /XFORD
3TOCHASTICITY NONLINEARITY AND INSTABILITY IN BIOLOGICAL INVASIONS
,EWIS - ! AND +AREIVA 0 !LLEE DYNAMICS AND THE SPREAD OF INVADING ORGANISMS
4HEORETICAL 0OPULATION "IOLOGY
,EWONTIN 2 # AND #OHEN $ /N POPULATION GROWTH IN A RANDOMLY VARYING ENVI
RONMENT 0ROCEEDINGS OF THE .ATIONAL !CADEMY OF 3CIENCES 53!
,IEBHOLD ! AND "ASCOMPTE * 4HE !LLEE EFFECT STOCHASTIC DYNAMICS AND THE ERADI
CATION OF INVASIVE SPECIES %COLOGY ,ETTERS
,ONSDALE 7 - 4HE BALANCE OF WEED POPULATIONS )N &RONTIERS OF 0OPULATION
%COLOGY EDS 2 " &LOYD AND ! 7 3HEPPARD #3)2/ 0UBLISHING -ELBOURNE
-AY 2 - 3TABILITY AND COMPLEXITY IN MODEL ECOSYSTEMS 0RINCETON 5NIVERSITY
0RESS 0RINCETON .EW *ERSEY
-AY 2 - 4HRESHOLDS AND BREAKPOINTS IN ECOSYSTEMS WITH A MULTIPLICITY OF STABLE
STATES .ATURE
-AY 2 - -ODELS FOR TWO INTERACTING POPULATIONS )N 4HEORETICAL %COLOGY ED
2 - -AY PP "LACKWELL 3CIENTIlC 0UBLICATIONS /XFORD
-AY 2 - #ONWAY ' 2 (ASSELL - 0 AND 3OUTHWOOD 4 2 % 4IME DELAYS
DENSITY DEPENDENCE AND SINGLE SPECIES OSCILLATIONS *OURNAL OF !NIMAL %COLOGY
-AY 2 - AND /STER ' & "IFURCATIONS AND DYNAMIC COMLEXITY IN SIMPLE ECOLOGICAL
MODELS !MERICAN .ATURALIST
-AYNARD 3MITH * -ODELS IN ECOLOGY #AMBRIDGE 5NIVERSITY 0RESS #AMBRIDGE
.ICHOLSON ! * !N OUTLINE OF THE DYNAMICS OF ANIMAL POPULATIONS !USTRALIAN
*OURNAL OF :OOLOGY
0ACALA 3 7 #ANHAM # $ 3APONARA * 3ILANDER * ! +OBE 2 + AND 2IBBENS %
&OREST MODELS DElNED BY lELD MEASUREMENTS ESTIMATION ERROR ANALYSIS AND
DYNAMICS %COLOGICAL -ONOGRAPHS
2EES - AND #RAWLEY - * 'ROWTH REPRODUCTION AND POPULATION DYNAMICS
&UNCTIONAL %COLOGY
2EES - AND #RAWLEY - * $O PLANT POPULATIONS CYCLE &UNCTIONAL %COLOGY
2EES - AND 0AYNTER 1 "IOLOGICAL #ONTROL OF 3COTCH "ROOM MODELLING THE DETER
MINANTS OF ABUNDANCE AND THE POTENTIAL IMPACT OF INTRODUCED INSECT HERBIVORES *OURNAL
OF !PPLIED %COLOGY
2OYAMA 4 !NALYTICAL 0OPULATION $YNAMICS #HAPMAN (ALL ,ONDON
3CHEFFER - AND #ARPENTER 3 #ATASTROPHIC REGIME SHIFTS IN ECOSYSTEMS LINKING
THEORY TO OBSERVATION 4RENDS IN %COLOGY AND %VOLUTION
3AETHER " % ET AL ,IFE HISTORY VARIATION PREDICTS THE EFFECTS OF DEMOGRAPHIC STOHCAS
TICITY ON AVIAN POPULATION DYNAMICS !MERICAN .ATURALIST
3AETHER " % ET AL 4IME TO EXTINCTION OF BIRD POPULATIONS %COLOGY IN PRESS
3TEPHENS 0 ! AND 3UTHERLAND 7 * #ONSEQUENCES OF THE !LLEE EFFECT FOR BEHAV
IOUR ECOLOGY AND CONSERVATION 4RENDS IN %COLOGY AND %VOLUTION
3TEPHENS 0 ! 3UTHERLAND 7 * AND &RECKLETON 2 0 7HAT IS THE !LLEE EFFECT
/IKOS
4AYLOR # - AND (ASTINGS ! &INDING OPTIMAL CONTROL STRATEGIES FOR INVASIVE SPE
CIES A DENSITY STRUCTURED MODEL FOR 3PARTINA ALTERNImORA *OURNAL OF !PPLIED %COLOGY
20 &RECKLETON 0- $OWLING AND .+ $ULVY
4ILMAN $ -AY 2 - ,EHMAN # , AND .OWAK - ! (ABITAT DESTRUCTION AND
THE EXTINCTION DEBT .ATURE
4ULJAPURKAR 3 0OPULATION DYNAMICS IN VARIABLE ENVIRONMENTS 3PRINGER .EW
9ORK
7ATKINSON ! 2 $ENSITY DEPENDENCE IN SINGLE SPECIES POPULATIONS OF PLANTS
*OURNAL OF 4HEORETICAL "IOLOGY
7HITE 4 # 2 /PPOSING PARADIGMS REGULATION OR LIMITATION OF POPULATIONS /IKOS
7ILLIAMSON - "IOLOGICAL )NVASIONS #HAPMAN (ALL ,ONDON 5+
#HAPTER SEVEN
,OCAL INTERACTIONS AND
INVASION DYNAMICS
POPULATION GROWTH IN SPACE
AND TIME
$ * -URRELL
).42/$5#4)/.
4HE STUDY OF BIOLOGICAL INVASIONS IS ESSENTIALLY THE STUDY OF POPULATION GROWTH
IN BOTH SPACE AND TIME .OT ONLY ARE WE INTERESTED IN THE POPULATION SIZE OF AN
INVADER BUT ALSO ITS RATE OF SPREAD ACROSS A LANDSCAPE OR REGION (ENCE THEORETI
CAL MODELS OF INVASIONS VERY OFTEN INCORPORATE SOME FORM OF SPATIAL HETEROGENEITY
3KELLAM "OERLIJST AND (OGEWEG (ASSELL ET AL +OT ET AL
+EELING ET AL ,EWIS ,EWIS AND 0ACALA 2IETKERK ET AL
"ROADLY SPEAKING THERE ARE TWO CLASSES OF SPATIAL HETEROGENEITY THE SO
CALLED
ENDOGENOUS OR SELF
INDUCED SPATIAL HETEROGENEITY TYPE I AND EXOGENOUS HETERO
GENEITY TYPE II 4YPE I HETEROGENEITY IS CAUSED BY INTERACTIONS BETWEEN INDIVID
UALS OR POPULATIONS AND LEADS TO WHAT IS SOMETIMES CALLED SELF
ORGANISED SPATIAL
PATTERNS "OERLIJST AND (OGEWEG (ASSELL ET AL +EELING ET AL
2IETKERK ET AL &OR EXAMPLE IN MONOCULTURES OF PINES INTENSE COMPETITION
BETWEEN CLOSE NEIGHBOURS LEADS TO AN EVEN SPACING OF ADULT TREES +ENKEL
-7 #ADOTTE ET AL (EDS) #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY
3PRINGER 0RINTED IN THE .ETHERLANDS
$* -URRELL
4YPE II HETEROGENEITY IS CAUSED BY PROCESSES BEYOND INDIVIDUAL INTERACTIONS SUCH
AS SOIL DEPTH TEMPERATURE GRADIENTS ETC 4HIS LEADS TO THE AGGREGATION OF INDIVIDU
ALS CAUSED BY SPECIES BEING ASSOCIATED WITH CERTAIN HABITAT TYPES MORE THAN OTHERS
&OR EXAMPLE SOME TROPICAL RAINFOREST TREES HAVE BEEN FOUND TO BE ASSOCIATED WITH
CERTAIN SOIL TYPES AND IT HAS BEEN ARGUED THAT THIS IS AN IMPORTANT FACTOR IN DETER
MINING THEIR AGGREGATED SPATIAL STRUCTURE 0ALMIOTTO ET AL 5NDOUBTEDLY
BOTH CLASSES OF HETEROGENEITY ARE IMPORTANT DURING INVASIONS INTERACTIONS WITH
OTHER ORGANISMS MAY DETERMINE WHETHER AN INVASION SUCCEEDS &AGAN AND "ISHOP
/WEN AND ,EWIS AS WELL AS THE AVAILABILITY AND SPATIAL PATTERNING OF
SUITABLE HABITAT 2ICHARDSON ET AL (ILL ET AL (OLWAY ET AL 7ITH
(OWEVER THIS CHAPTER WILL FOCUS ON ONLY ENDOGENOUS SPATIAL HETEROGENEITY
IN AN ATTEMPT TO SHOW HOW IMPORTANT LOCAL SPATIAL STRUCTURE CAN BE IN DETERMIN
ING THE RATE OF SPREAD OF A POPULATION STARTING FROM A LOW DENSITY SEE "OLKER
FOR RESULTS FOR A SIMILAR MODEL IN HETEROGENEOUS SPACE
%COLOGICAL INTERACTIONS FOR MOST ORGANISMS TEND TO OCCUR BETWEEN INDIVIDUALS
THAT ARE NEARBY IN SPACE #OUPLED TO THIS MOST ORGANISMS LARGER THAN MM ARE
ARGUED TO BE IN SOME WAY DISPERSAL LIMITED &INLAY AND #LARKE &INLAY ET AL
&INLAY -ANY MICROBIAL ORGANISMS ARE SO SMALL THAT THEY CAN BE
CARRIED LARGE DISTANCES BY WIND AND TOGETHER WITH LONG LIVED RESTING STAGES THIS
MEANS THAT MANY MICROBIAL SPECIES MAY ENJOY A NEAR GLOBAL DISTRIBUTION %ARLY
IN ITS HISTORY ECOLOGICAL THEORY NEGLECTED BOTH THESE ASPECTS AND IT WAS ASSUMED
THAT INDIVIDUALS RANDOMLY BUMP INTO ONE ANOTHER THE SO CALLED MEAN lELD
ASSUMPTION -ORE RECENTLY NON UNIFORM DISPERSAL HAS BEEN INCORPORATED AND
HAS BEEN USED WITH MUCH SUCCESS IN DESCRIBING THE RATES OF SPREAD INVASION
OF NUMEROUS SPECIES +OT ET AL 6EIT AND ,EWIS (OWEVER MOST OF
THESE MODELS HAVE NOT INCLUDED THE DISCRETE NATURE OF INDIVIDUALS AND WITHOUT IT
THE TIGHT SMALL SCALE CLUSTERING THAT IS OFTEN EVIDENT IN NATURAL POPULATIONS CANNOT
BE INCLUDED
!S HAS BEEN OFTEN OBSERVED THERE IS A CRUCIAL FEEDBACK BETWEEN THE SPATIAL
PATTERN OF INDIVIDUALS ACROSS THE LANDSCAPE AND THE DYNAMICS OF THE POPULATION
"OLKER AND 0ACALA $URRETT AND ,EVIN ,OCAL DISPERSAL EVEN WITHOUT
ANY DENSITY DEPENDENT EFFECTS ON DEATH RATES OR FECUNDITY RESULTS IN A CLUSTERED
OR PATCHY DISTRIBUTION OF INDIVIDUALS ACROSS THE LANDSCAPE 9OUNG ET AL
)N SUCH A SCENARIO THE SPATIAL PATTERN IS NOT IMPORTANT FOR THE POPULATION DYNAM
ICS OTHER THAN IN DETERMINING THE RATE OF SPREAD OF AN INVADER ACROSS A LAND
SCAPE (OWEVER AS SHALL BE EXPANDED UPON BELOW ONCE LOCAL DENSITY DEPENDENT
PROCESSES ARE INCORPORATED THE SPATIAL PATTERN OF INDIVIDUALS MAY BECOME VERY
IMPORTANT TO MANY FEATURES OF THE INVASION DYNAMICS AND MAY ULTIMATELY DETER
MINE WHETHER AN EXOTIC CAN INVADE OR NOT 7ITH LOCAL SPATIAL STRUCTURE WHAT AN
INDIVIDUAL EXPERIENCES IN ITS LOCAL NEIGHBOURHOOD MAY BE VERY DIFFERENT TO THE
OVERALL DENSITY OF INDIVIDUALS ACROSS THE LANDSCAPE 4HIS LOCAL SPATIAL STRUCTURE
THEN HELPS TO DETERMINE THE LEVELS OF COMPETITION THAT AN INDIVIDUAL EXPERIENCES
AND HENCE INDIRECTLY DETERMINES THE LEVEL OF COMPETITION AND LOCAL SPATIAL STRUC
TURE IN THE NEXT TIME INTERVAL
0OPULATION GROWTH IN SPACE AND TIME
4HE REST OF THIS CHAPTER WILL FOCUS ON A GENERAL MODEL FOR THE SPATIO
TEMPORAL
POPULATION DYNAMICS OF AN INVADING SPECIES 7E WILL CONSIDER INDIVIDUALS SIMILAR
TO PERENNIAL PLANTS THAT REPRODUCE DISPERSING THEIR OFFSPRING LOCALLY AND DIE AT
RATES THAT ARE DEPENDENT ON THEIR LOCAL NEIGHBOURHOOD DENSITY 4WO MODELS
WILL BE PRESENTED ONE INDIVIDUAL
BASED MODEL AND ITS DETERMINISTIC APPROXIMA
TION 4HE LINK BETWEEN THESE MODELS AND THE CLASSICAL NON
SPATIAL LOGISTIC MODEL
FOR POPULATION GROWTH IS MADE EXPLICIT AND WILL ACT AS A REFERENCE POINT FOR THE
RESULTS THAT ARE OBTAINED WHEN INTERACTIONS AND DISPERSAL ARE LOCALISED IN SPACE
"OTH SPATIAL MODELS WILL BE USED TO INVESTIGATE THE SCALES OF DISPERSAL AND NEIGH
BOURHOOD INTERACTIONS ON THE TRANSIENT DYNAMICS SUCCESS AND SPEED OF INVASION
AND ASYMPTOTIC STATE OF THE POPULATION THE lNAL POPULATION DENSITY )T IS ALSO
OBSERVED THAT THE EXPECTED REPRODUCTIVE OUTPUT OF AN INDIVIDUAL IN THE ABSENCE OF
COMPETITION IS ALSO IMPORTANT IN DETERMINING THE SUCCESS AND SPEED OF AN INVASION
AS WELL AS THE lNAL POPULATION SIZE
! 34/#(!34)# ).$)6)$5!, "!3%$ -/$%, )"-
)NVASIONS ARE INHERENTLY STOCHASTIC THEY VERY OFTEN START WITH JUST A FEW INDIVIDU
ALS AND THE SUCCESS OF AN INVASION OWES A GREAT DEAL TO CHANCE THEREFORE IT MAKES
SENSE TO START FROM A STOCHASTIC MODEL THAT FOCUSES ON THE INDIVIDUAL
LEVEL MICRO
SCOPIC PROCESSES OF BIRTHS AND DEATHS 7E WILL CONCENTRATE ON A SPATIAL EXTENSION
TO THE FAMILIAR LOGISTIC MODEL FOR POPULATION GROWTH 0EARL AND 2EAD ONE
WHERE BOTH INTERACTIONS AND DISPERSAL ARE LOCALISED IN SPACE
#ONSIDER A POPULATION OF ORGANISMS OF A SINGLE SPECIES INVADING A TWO
DIMEN
SIONAL LANDSCAPE SUCH AS A lELD )N THE )"- SPACE IS CONTINUOUS AS OPPOSED TO
A LATTICE OR SITE
BASED MODEL AND LARGE ENOUGH FOR EDGE EFFECTS TO BE NEGLIGIBLE
4O REDUCE THE EDGE EFFECTS THE LANDSCAPE IS IN THE SHAPE OF A TORUS FORMERLY
SPEAKING THE BOUNDARIES ARE PERIODIC !N INDIVIDUAL I LOCATED AT COORDINATES
XI [X X] IS DENOTED BY A $IRAC DELTA FUNCTION bXI X 4HE $IRAC DELTA FUNCTION
SERVES TO PICK OUT INDIVIDUALS IT IS A PEAK OF INlNITE HEIGHT AND INlNITELY SMALL
WIDTH CENTRED ON THE LOCATION OF EACH INDIVIDUAL GIVING AN AREA UNDERNEATH IT
THAT SUMS TO ONE 4HE STATE OF THE POPULATION AT TIME T IS GIVEN BY THE FUNCTION
PX T WHICH IS THE SUM OF ALL THE DELTA FUNCTIONS AND THEREFORE DESCRIBES THE
NUMBER AND LOCATIONS OF ALL INDIVIDUALS IN THE POPULATION
4HERE ARE THREE DIFFERENT TYPES OF EVENT THAT MAY OCCUR IN THE )"- BIRTH
DEATH AND MOVEMENT 4O KEEP IN CLOSE ASSOCIATION WITH THE CLASSICAL LOGISTIC
MODEL ONLY DEATHS ARE DENSITY DEPENDENT ALTHOUGH THE MODEL IS mEXIBLE SO THAT
BIRTHS AND MOVEMENTS MAY ALSO BE DEPENDENT UPON LOCAL DENSITIES "OLKER AND
0ACALA
4HE PROBABILITY PER UNIT TIME $X X P THAT AN INDIVIDUAL AT LOCATION X DIES
IS GIVEN BY
$X X P D D q 0WX n X q ;PX n bXX = DX
$* -URRELL
4HE DEATH TERM CONSISTS OF TWO PARTS 4HE lRST IS AN INTRINSIC PER CAPITA DEATH
RATE D AND THE SECOND PART MODIlES THE DEATH RATE ACCORDING TO THE PRESENCE OF
NEARBY NEIGHBOURS .EIGHBOURS HAVE A PER CAPITA EFFECT D ON THE DEATH RATE OF THE
FOCAL INDIVIDUAL (OWEVER IT IS ASSUMED THAT COMPETITIVE EFFECT OF NEIGHBOURS IS
DEPENDENT UPON THE DISTANCE BETWEEN THEM 4HE TERM WX n X OTHERWISE CALLED
THE COMPETITION KERNEL DESCRIBES THE WEIGHTING THAT NEIGHBOURS AT LOCATION X
HAVE ON THE DEATH RATE AT LOCATION X AND IS A FUNCTION OF THE DISTANCE X n X THAT
SEPARATES THE PAIR 4HIS WEIGHTING IS MULTIPLIED BY THE DENSITY PX OF INDIVIDUALS
AT LOCATION X AND THE INTEGRAL TAKES INTO ACCOUNT THE CONTRIBUTIONS TO THE DEATH
RATE AT X OF ALL LOCATIONS X IN THE NEIGHBOURHOOD OF X 4HE LAST TERM bXX IS ONLY
SWITCHED ON WHEN X X THEREBY REMOVING THE SPURIOUS EFFECT THAT AN INDIVIDUAL
WOULD OTHERWISE HAVE ON ITS OWN DEATH RATE
4HE PROBABILITY PER UNIT TIME "X X P THAT AN INDIVIDUAL LOCATED AT X PRO
DUCES AN OFFSPRING AT X IS GIVEN BY
"X X P B q MX n X
WHERE B IS THE INTRINSIC PER CAPITA BIRTH RATE AND MX n X IS THE DISPERSAL KERNEL
WHICH DESCRIBES THE PROBABILITY THAT THE NEWBORN INDIVIDUAL COMES TO REST AT
A DISPLACEMENT X X FROM ITS PARENT
4HE DISPERSAL AND COMPETITION KERNELS ARE VERY mEXIBLE AND CAN TAKE ON PRACTI
CALLY ANY SHAPE SEE FOR VARIATIONS SEE "OLKER AND 0ACALA )T SEEMS SENSIBLE
TO ENSURE THAT THEY ARE BOTH MONOTONICALLY DECREASING FUNCTIONS AND WE CHOOSE
TO USE BIVARIATE 'AUSSIAN DISTRIBUTIONS
\X n X\
MX n X EXP " n " A
- SM
\X n X\
WX n X EXP " n " B
7 SW
EACH HAVING ONE PARAMETER SM SW THAT MEASURES THE WIDTH OF THE KERNEL ! SMALL
VALUE FOR SM MEANS THAT OFFSPRING FALL VERY CLOSE TO THEIR PARENT SHORT SCALE DIS
PERSAL ! SMALL VALUE FOR SW MEANS THAT COMPETITION IS VERY INTENSE BETWEEN CLOSE
NEIGHBOURS BUT THAT IT SOON DIMINISHES WITH DISTANCE SHORT SCALE INTERACTIONS
/N THE OTHER HAND LARGE VALUES FOR SM SW REDUCES THE IMPORTANCE OF SPACE
&OR PRACTICAL PURPOSES WE TRUNCATE THE KERNELS AT A DISTANCE OF TIMES THE
PARAMETER VALUE THE EFFECT THIS HAS ON THE DYNAMICS IS NEGLIGIBLE 4HE KERNELS
ARE NORMALISED SO THAT THAT THEIR INTEGRALS OVER X n X ARE UNITY - AND 7 BEING
THE NORMALISATION CONSTANTS
0OPULATION GROWTH IN SPACE AND TIME
30!4)!, 34!4)34)#3
)N THE )"- EACH INDIVIDUAL AND EACH EVENT NEEDS TO BE CONSTANTLY TRACKED )N ORDER
TO INVESTIGATE THE IMPLICATIONS OF INCORPORATING LOCAL INTERACTIONS AND LOCAL DISPER
SAL INTO THE LOGISTIC MODEL FOR POPULATION GROWTH IT HELPS TO HAVE SOME SUMMARY
STATISTICS THAT CAN BE OUTPUT FROM THE )"- 4HESE WILL LATER BECOME STATE VARIABLES
IN THE DETERMINISTIC APPROXIMATION SO IT IS IMPORTANT THAT THEY ARE DElNED
4HE lRST SPATIAL MOMENT IS THE FAMILIAR AVERAGE DENSITY
.P T q 0 PX T DX
!
4HIS SIMPLY SUMS OVER ALL INDIVIDUALS IN THE POPULATION AT TIME T AND DIVIDES BY
THE AREA ! IN WHICH THE POPULATION LIVES
4HE lRST SPATIAL MOMENT DOES NOT GIVE ANY INFORMATION ON HOW INDIVIDUALS
ARE SPREAD ACROSS THE LANDSCAPE IN ORDER TO GET A FEEL FOR THE SPATIAL PATTERN
HIGHER ORDER SPATIAL MOMENTS ARE REQUIRED 4HERE ARE A NUMBER OF HIGHER ORDER
SPATIAL MOMENTS THAT COULD BE USED $IGGLE #RESSIE BUT THE ONE USED
HERE IS
#j P T q 0 PX T q ;PX j T n bXX j = DX
!
4HE SECOND SPATIAL MOMENT IS THE AVERAGE DENSITY OF PAIRS SEPARATED BY A
DISTANCE j )T CAN BE THOUGHT OF AS THE INDIVIDUALS EYE VIEW SINCE IT DESCRIBES
THE EXPECTED NEIGHBOUR DENSITY AT DIFFERENT DISTANCES AND HENCE DESCRIBES THE
VARIATION IN THE DENSITY OF INDIVIDUALS ACROSS THE LANDSCAPE /NCE AGAIN THE DELTA
TERM REMOVES THE SELF
PAIR TERM THAT ARISES WHEN X jX 4HIS FORM OF SECOND
MOMENT HAS THE BENElT OF BEING MATHEMATICALLY SIMPLE A FEATURE THAT IS USEFUL FOR
THE DETERMINISTIC APPROXIMATION THAT FOLLOWS )N THE RESULTS GIVEN BELOW THE
SECOND MOMENTS ARE NORMALISED BY DIVIDING BY .P T 4HIS MEANS THAT WHEN
#j P T THIS INDICATES A MORE CROWDED NEIGHBOURHOOD THAN WOULD BE
EXPECTED IF THE SPATIAL PATTERN OF INDIVIDUALS WERE RANDOM AGGREGATION WHEN
#j P T THIS INDICATES A RELATIVELY COMPETITOR
FREE NEIGHBOURHOOD COMPARED
TO A RANDOM PATTERN SPATIAL SEGREGATION AND WHEN #j P T THIS INDICATES
A RANDOM SPATIAL ASSOCIATION OF INDIVIDUALS
! $%4%2-).)34)# !002/8)-!4)/.
4HERE IS MUCH THAT CAN BE GAINED FROM THE )"- BUT KEEPING TRACK OF ALL INDIVIDU
ALS AND THEIR DYNAMICS IS COMPUTATIONALLY VERY INTENSIVE 4HE )"- DElNED ABOVE
$* -URRELL
IS STOCHASTIC ANY INDEPENDENT REALISATIONS ARE EXPECTED TO HAVE SLIGHTLY DIFFER
ENT PATTERNS AND -ARKOVIAN THE RATE OF CHANGE OF THE SYSTEM DEPENDS ONLY ON
THE CURRENT STATE 4HESE MAKE IT HARD TO DISCERN THE ECOLOGICAL SIGNAL FROM THE
BACKGROUND NOISE MANY REALISATIONS ARE REQUIRED TO GET A BETTER PICTURE OF PROP
ERTIES OF THE MODEL ,AW ET AL AND HENCE FULL ANALYSIS OF THE )"- IS VERY
DIFlCULT &OR EXAMPLE KNOWING WHEN THE POPULATION HAS REACHED ITS END STATE
CAN BE VERY DIFlCULT TO DETERMINE
#OMPLEMENTING THE )"- WITH AN APPROACH THAT DEALS WITH THE MACROSCOPIC
PROPERTIES SUCH AS POPULATION DENSITIES IS THEREFORE DESIRABLE 3CALING UP FROM
SMALL
SCALE PROCESSES TO LARGE
SCALE PROPERTIES IS NOT A TRIVIAL TASK ,EVIN
BUT RECENT ADVANCES IN MATHEMATICAL TECHNIQUES HAVE EXPANDED THE RANGE OF
CASES FOR WHICH THIS IS POSSIBLE IN SPATIAL MODELS "OLKER AND 0ACALA "OLKER
"OLKER AND 0ACALA $IECKMANN AND ,AW ,AW AND $IECKMANN
-URRELL AND ,AW "OLKER "OLKER ET AL ,AW ET AL
-URRELL AND ,AW -URRELL ET AL
&ROM THE DESCRIPTION OF THE )"- ABOVE IT IS POSSIBLE TO DERIVE A DETERMINIS
TIC APPROXIMATION THAT DESCRIBES THE EXPECTED RATE OF CHANGE OF THE POPULATION
$IECKMANN AND ,AW 4HE EXPECTED RATE OF CHANGE IN POPULATION DENSITY
IS GIVEN BY
.T B.T n D.T n D q 0 Wj #j D j
4HE lRST AND SECOND TERMS DEAL WITH THE CONTRIBUTIONS TO POPULATION DENSITY OF
THE INTRINSIC RATES OF BIRTH AND DEATH 4HESE ARE MADE EXPLICIT IN THE SPATIAL MODEL
RATHER THAN SUBSTITUTING R B n D BECAUSE AS WILL BE SHOWN BELOW BIRTHS AND
DEATHS HAVE VERY DIFFERENT EFFECTS ON THE SPATIAL STRUCTURE 4HE THIRD TERM IS WHERE
THE SPATIAL STRUCTURE lRST ENTERS THE DYNAMICAL SYSTEM #OMPETITION IS NOW DEPEN
DENT ON THE EXPECTED DENSITY OF NEIGHBOURS AROUND AN INDIVIDUAL #j T AND IS
SCALED BY BOTH THE INTERACTION KERNEL Wj WHICH TAKES INTO ACCOUNT THE DISTANCE
BETWEEN PAIRS AND BY THE PER CAPITA COMPETITIVE EFFECT D
4HE SPATIAL STRUCTURE SUMMARISED BY #j T HAS DYNAMICS OF ITS OWN THAT
ALSO NEED TO BE MADE EXPLICIT 4HE EXPECTED RATE OF CHANGE OF PAIR DENSITIES ARE
DESCRIBED BY
#j T B 0 Mj # j j T D j A
B Mj . T B
n D # j T C
n D 0 Wj 4 j j T D j D
n D Wj # j T E
%ACH TERM DESCRIBES EVENTS THAT LEAD TO THE CREATION AND DELETION OF PAIRS OR
MORE FORMERLY PAIR DENSITIES 4HE lRST TERM A DESCRIBES HOW PAIRS ARE FORMED
WHEN A NEWBORN DISPERSES A DISTANCE j FROM ITS PARENT 4HE SECOND TERM B TAKES
INTO ACCOUNT PAIRS FORMED WHEN ONE OF THE MEMBERS OF THE PAIR IS THE PARENT
0OPULATION GROWTH IN SPACE AND TIME
4HE OTHER THREE TERMS DEAL WITH LOSS OF PAIRS THROUGH DEATHS 4ERM C DESCRIBES
HOW PAIRS ARE LOST THROUGH DENSITY INDEPENDENT DEATHS 4ERM D MODIlES THIS RATE
DUE TO NEIGHBOURS AT A DISTANCE j WITH THE INTERACTION KERNEL Wj WEIGHTING
THE EFFECT OF THE NEIGHBOURS ACCORDING TO DISTANCE BETWEEN LOCATIONS 3INCE THE
PAIR IS AFFECTED BY A THIRD INDIVIDUAL INFORMATION ON THE EXPECTED DENSITY OF TRIP
LETS IS REQUIRED AND THIS IS DENOTED BY 4 THE THIRD SPATIAL MOMENT 4HIS REPRESENTS
A NON
TRIVIAL TECHNICAL PROBLEM THAT IS EXPANDED UPON IN THE !PPENDIX &INALLY
E ACCOUNTS FOR DENSITY DEPENDENT DEATHS CAUSED BY COMPETITIVE INTERACTIONS
WITHIN THE PAIR .OTE THAT EACH TERM IS DOUBLED BECAUSE BIRTHS OR DEATHS CAN OCCUR
AT EITHER LOCATION IN THE PAIR
(AVING DElNED BOTH THE STOCHASTIC AND THE DETERMINISTIC MODELS WE CAN NOW
TURN TO INVESTIGATE THE EFFECT OF INCORPORATING LOCAL SPATIAL STRUCTURE ON THE SUC
CESS AND SPEED OF INVASION AND ALSO ON THE lNAL POPULATION DENSITY !S A REFER
ENCE POINT THE RESULTS ARE COMPARED TO THE CONCLUSIONS THAT ARE REACHED FROM THE
CLASSICAL LOGISTIC MODEL FOR POPULATION GROWTH 2ECALL THAT IN THE CLASSICAL LOGISTIC
MODEL POPULATION GROWTH IS REACHED AT EXACTLY HALF THE lNAL POPULATION SIZE
THE GROWTH TRAJECTORY SHOWS AN 3
SHAPED CURVE AND THAT FROM LOW INITIAL DENSI
TIES POPULATIONS THAT HAVE LOWER lNAL DENSITIES REACH THEIR EQUILIBRIUM THE FASTEST
4HE READER SHOULD NOTE THAT THE CLASSICAL LOGISTIC MODEL IS A SPECIAL CASE OF BOTH
MODELS DElNED HERE AND CAN BE RECOVERED WHENEVER INTERACTIONS OCCUR OVER
VERY LARGE SPATIAL SCALES 4HIS IS TRUE EVEN WHEN DISPERSAL IS SHORT
RANGE BECAUSE
WHEN INTERACTIONS OCCUR OVER SUFlCIENTLY LARGE SPATIAL SCALES ANY SPATIAL PATTERN
THAT IS EVIDENT IS IRRELEVANT TO THE DYNAMICS THE POSITIONS OF INDIVIDUALS IN SPACE
BECOMES UNIMPORTANT
2%35,43
$ISPERSAL DISTANCE
4HE IMPORTANCE OF DISPERSAL DISTANCE FOR THE INVASION DYNAMICS IS SHOWN IN &IG
3HOWN ARE POPULATIONS ONE WITH RELATIVELY POOR DISPERSAL ONE WITH RELATIVELY
GOOD DISPERSAL ABILITIES THE INVADERS ARE OTHERWISE IDENTICAL 3NAPSHOTS OF THE
)"- SHOW THAT THE POORER DISPERSER SOON DEVELOPS A CLUMPED DISTRIBUTION AND THIS
INHIBITS POPULATION GROWTH BECAUSE INDIVIDUALS EXPERIENCE CROWDED NEIGHBOUR
HOODS /N THE OTHER HAND THE BETTER DISPERSER SEEMS TO HAVE FASTER POPULATION
DYNAMICS AND REACHES A HIGHER POPULATION DENSITY 4HIS IS BECAUSE MORE OFFSPRING
CAN ESCAPE INTENSE COMPETITION WITH THEIR PARENTS AND HENCE THEIR NEIGHBOURHOOD
IS RELATIVELY COMPETITOR
FREE )NSPECTION OF THE SPATIAL STATISTICS DURING THE INVASION
CONlRMS THIS THE BETTER DISPERSER HAS FASTER POPULATION DYNAMICS EXPERIENCES
A LESS CROWDED NEIGHBOURHOOD AND REACHES A HIGHER lNAL DENSITY &IG .OTE
HOWEVER THAT UNTIL ABOUT TIME UNITS HAVE PASSED EVEN THE BETTER DISPERSER
SHOWS SOME DEGREE OF AGGREGATION &IG BUT THIS THEN DISAPPEARS AS POPULATION
GROWS AND SPREADS AND EVENTUALLY THE INDIVIDUALS ARE SPATIALLY SEGREGATED
$* -URRELL
&IG 3NAPSHOTS FROM THE )"- SHOWING THE INVASION OF TWO DIFFERENT POPULATIONS
)N A THE INDIVIDUALS HAVE SHORT
RANGE DISPERSAL SM WHEREAS IN B INDIVIDUALS
HAVE LONGER
RANGE DISPERSAL SM )N BOTH CASES THE STARTING POPULATION HAS
INDIVIDUALS SCATTERED RANDOMLY ACROSS THE LANDSCAPE WHICH IS OF UNIT AREA /THER
PARAMETERS HELD CONSTANT ARE B D D SW
0OPULATION GROWTH IN SPACE AND TIME
&IG 4EMPORAL DYNAMICS FOR A THE MEAN DENSITY lRST SPATIAL MOMENTS AND B
SECOND SPATIAL MOMENTS AT SHORT DISTANCES FOR THE INVASIONS OF THE SHORT
RANGE DISPERSER
SOLID LINE AND CIRCLES AND THE LONG
RANGE DISPERSER BROKEN LINE AND CROSSES SHOWN IN
&IG )N B VALUES GREATER THAN ABOVE THE SOLID LINE INDICATE AGGREGATION OF INDIVIDUALS
WHEREAS VALUES LESS THAN BELOW THE SOLID LINE INDICATE SPATIAL SEGREGATION 0ARAMETERS
AND STARTING CONDITIONS ARE AS DESCRIBED IN &IG
$* -URRELL
7E NOW TURN TO SEE HOW LOCAL INTERACTIONS AND LOCAL DISPERSAL AFFECT THE INVA
SION SUCCESS 0S OF A POPULATION &OR A RANGE OF SCALES OF DISPERSAL POPULATIONS
OF INDIVIDUALS ARE INITIALISED IN THE )"- TO INVESTIGATE HOW B THE CRITICAL BIRTH
RATE REQUIRED FOR INVASION CHANGES WITH THE DISPERSAL OF THE INVADER &OR EACH
INVADING POPULATION THE MODEL IS RUN FOR TIME UNITS AND AN INVASION IS DEEMED
A SUCCESS IF IT HAS EXPANDED AFTER THIS TIME IE IF .T .T 4HE PROB
ABILITY OF INVASION SUCCESS 0S IS THEN CALCULATED FROM THE PROPORTION OF THE
POPULATIONS THAT SHOW THIS INCREASE IN POPULATION DENSITY 7HEN THE INVADER IS
RESTRICTED TO ONLY SHORT
RANGE DISPERSAL IT TAKES A RELATIVELY LARGE B FOR THE POPU
LATION TO BE ABLE TO INVADE WITH ANY DEGREE OF CERTAINTY &IG &OR EXAMPLE FOR
SM INVASION SUCCESS DOES NOT EXCEED UNTIL B )N FACT FOR MANY
VALUES FOR INTRINSIC BIRTH RATES B THAT WOULD RESULT IN ALMOST GUARANTEED INVA
SION SUCCESS IN THE STOCHASTIC VERSION OF THE NON
SPATIAL LOGISTIC MODEL NONE OF
THE INITIAL POPULATIONS ARE ABLE TO INVADE
&IG 4HE EFFECT OF INTRINSIC BIRTH RATE B AND DISPERSAL DISTANCE EXPRESSED BY THE
PARAMETER SM THAT DETERMINES THE WIDTH OF THE DISPERSAL KERNEL ON THE INVASION SUCCESS
0S OF A SINGLE SPECIES 4HE LANDSCAPE OF UNIT AREA IS SEEDED WITH RANDOMLY LOCATED
INDIVIDUALS AND EACH SQUARE REPRESENTS THE PROPORTION OF REALISATIONS THAT SHOW
POPULATION GROWTH AFTER TIME UNITS HAVE ELAPSED /THER PARAMETERS HELD CONSTANT
THROUGHOUT ARE D D SW
0OPULATION GROWTH IN SPACE AND TIME
4HERE ARE NUMEROUS EXAMPLES SHOWING HOW THE INVASION RATE IS AFFECTED BY
THE DISPERSAL KERNEL &OR EXAMPLE #ASWELL ET AL SHOWED THAT THE 0IED
&LYCATCHER &ICEDULA HYPOLEUCA IN %UROPE IS A SLOWER INVADER THAN THE 3TARLING
3TURNUS VULGARIS IN .ORTH !MERICA AND THAT OF THE DIFFERENCE BETWEEN INVA
SION RATES COULD BE EXPLAINED BY THE DIFFERENCES IN DISPERSAL ABILITIES WITH THE REST
BEING EXPLAINED BY THE DIFFERENCES IN DEMOGRAPHY /VER LONGER PERIODS OF TIME
THE DISPERSAL ABILITIES OF AN INVADER MIGHT BE EXPECTED TO BE UNDER SOME STRONG
SELECTIVE PRESSURE 5SING A SIMULATION MODEL 4RAVIS AND $YTHAM WERE ABLE
TO SHOW THAT EARLY ON DURING AN INVASION ON AN ISLAND LONG
DISTANCE INVASION IS
SELECTED FOR BUT THAT SHORTER DISTANCE SUBSEQUENTLY EVOLVES %VIDENCE FOR SUCH
PATTERNS CAN BE SEEN IN THE DIFFERENCE BETWEEN WIND
DISPERSED PLANTS IN MAINLAND
AND SMALL ISLAND SUBPOPULATIONS #ODY AND /VERTON )N NEW ISLAND POPU
LATIONS THE DISPERSAL ABILITY OF THE PLANTS TENDS TO BE GREATER THAN THE MAINLAND
POPULATIONS /VER TIME SELECTION IS FOR A REDUCTION IN DISPERSAL ABILITY ON THE
ISLANDS SO THAT OLD ISLAND PLANT POPULATIONS HAVE A LOWER DISPERSAL ABILITY THAN
THE MAINLAND POPULATIONS SEE ALSO $ARLINGTON "ROWN AND ,OMOLINO
7HITTAKER &ILIN AND :IV
2EPRODUCTIVE NUMBER
!S "OLKER AND 0ACALA AND "OLKER POINT OUT THE POPULATION
DYNAMICS OF AN INVADING SPECIES IS GREATLY AFFECTED BY THE REPRODUCTIVE NUMBER 2
WHERE 2 BD WHICH IS A MEASURE OF HOW SENSITIVE INDIVIDUALS ARE TO COMPETI
TION 2 CAN BE THOUGHT OF AS BEING THE EXPECTED LIFETIME REPRODUCTIVE OUTPUT IN
THE ABSENCE OF COMPETITION AND IN THE SPATIAL MODELS 2 BECOMES IMPORTANT IN
DETERMINING HOW GOOD A COLONISER A SPECIES IS 7EEDY SPECIES AND HENCE INVAD
ERS ARE EXPECTED TO HAVE A LARGE VALUE FOR 2 LESS WEEDY SPECIES WILL HAVE A LOW 2
NUMBER .OTE THAT VARYING 2 IN THE NON SPATIAL LOGISTIC MODEL HAS NO OUTCOME ON
POPULATION DYNAMICS 2 IS IMPORTANT ONLY FOR THE SPATIAL DYNAMICS
7HEN 2 IS SMALL THE SPECIES IS A POOR COLONISER AND THE POPULATION GROWTH RATE
IS SLOW OR IF 2 IS SMALL ENOUGH THE INVASION MAY FAIL ALTOGETHER &IG -OREOVER
NOTE THAT LOW VALUES FOR 2 ARE ASSOCIATED WITH LONG PERIODS OF NEAR LINEAR POPULA
TION GROWTH THE CLASSICAL SIGMOIDAL GROWTH CURVE CHARACTERISTIC OF THE NON SPATIAL
LOGISTIC MODEL ONLY OCCURS WHEN 2 IS HIGH ANDOR WHEN DISPERSAL IS LONG RANGE
SEE ALSO ,AW ET AL !NOTHER FEATURE IS THAT 2 GREATLY AFFECTS THE lNAL POPU
LATION SIZE WITH LARGE VALUES FOR 2 LEADING TO HIGHER POPULATION DENSITIES &IG
#OUPLED TO THIS IS THE INTERESTING OBSERVATION THAT THE LARGER POPULATIONS TAKE
THE SHORTEST AMOUNT OF TIME TO REACH THEIR EQUILIBRIUM THIS IS IN STARK CONTRAST
TO THE NON SPATIAL LOGISTIC MODEL WHERE THE SMALLEST POPULATION REACH THEIR lNAL
SIZE BEFORE THOSE WITH LARGER POPULATION DENSITIES .OTE ALSO THAT THE AGREEMENT
BETWEEN THE )"- AND THE DETERMINISTIC APPROXIMATION MOMENT DYNAMICS IS VERY
GOOD &IG /THER RESULTS CONlRM THESE lNDINGS -URRELL AND ,AW ,AW
ET AL -URRELL AND ,AW AND THE REST OF THE CHAPTER SHALL CONCENTRATE
ON THE DETERMINISTIC APPROXIMATION
$* -URRELL
&IG 4HE EFFECT OF THE REPRODUCTIVE NUMBER 2 ON POPULATION GROWTH 3HOWN IN A ARE
MEAN TRAJECTORIES FOR REALISATIONS OF THE )"- AND IN B THE DETERMINISTIC APPROXIMATION
USING THE DYNAMICAL SYSTEM AND FOR DIFFERENT VALUES FOR 2 )N ALL CASES THE INTRINSIC
GROWTH RATE R IS HELD CONSTANT R )N I B D II B D
III B D IV B D V B D /THER PARAMETERS
HELD CONSTANT THROUGHOUT ARE D SW SM
0OPULATION GROWTH IN SPACE AND TIME
!NOTHER EXAMPLE SHOWS HOW IMPORTANT 2 CAN BE TO THE INVASION DYNAMICS
AND HOW IT CAN REVERSE SOME OF THE CONCLUSIONS OF THE NON
SPATIAL LOGISTIC MODEL
&IG #ONSIDER IMAGINARY SPECIES LABELLED AND RESPECTIVELY IF R R
IE SPECIES HAS A HIGHER INTRINSIC RATE OF GROWTH THAN SPECIES BUT 2 2 AND
IF INTERACTIONS AND DISPERSAL ARE LOCALISED IN SPACE THEN IT IS POSSIBLE FOR SPECIES
TO BE THE BETTER INVADER BOTH IN TERMS OF SPEED OF INVASION AND EQUILIBRIUM
DENSITY 3IMILAR RESULTS COULD EASILY BE PRODUCED FOR SPECIES WITH DIFFERENT
SCALES OF DISPERSAL AND THIS SENDS THE MESSAGE THAT KNOWING R MAY NOT BE SUFl
CIENT IN DETERMINING WHETHER A SPECIES CAN INVADE OR NOT 4HESE RESULTS ALL POINT
TO THE FACT THAT A HIGH REPRODUCTIVE NUMBER 2 IS ADVANTAGEOUS FOR AN INVADER
IN HOMOGENEOUS SPACE BUT OTHER RESULTS "OLKER SHOW THAT LOW VALUES
FOR 2 CAN BE BENElCIAL IF THERE IS SPATIAL VARIATION IN THE EXTERNAL ENVIRONMENT
TYPE II SPATIAL HETEROGENEITY
&IG !N EXAMPLE SHOWING HOW 2 MAY REVERSE A MAIN CONCLUSION OF THE NON
SPATIAL
LOGISTIC MODEL 3PECIES BROKEN LINES HAS A HIGH VALUE FOR R BUT A LOW 2 NUMBER 3PECIES
SOLID LINES HAS A LOWER VALUE FOR R BUT A HIGHER 2 NUMBER )N THE NON
SPATIAL MODEL
THICK LINES SPECIES IS THE BETTER INVADER AND HAS A HIGHER DENSITY AT EQUILIBRIUM
7HEN INTERACTIONS AND DISPERSAL ARE LOCALISED IN SPACE THIN LINES THIS RESULT IS REVERSED
NOW THE SPECIES HAS THE FASTER INVASION RATE AND HIGHER DENSITY AT EQUILIBRIUM &OR
SPECIES B D GIVING R 2 AND FOR SPECIES B
D GIVING R 2 /THER PARAMETERS ARE IDENTICAL FOR BOTH SPECIES AND
ARE D SW SM
$* -URRELL
&INAL POPULATION SIZE
3O FAR WE HAVE CONCENTRATED ON THE TRANSIENT DYNAMICS BUT NEIGHBOURHOOD
INTERACTIONS AND LOCAL DISPERSAL CAN ALSO GREATLY AFFECT THE lNAL POPULATION SIZE
,AW ET AL %THERIDGE )F NEIGHBOURHOOD INTERACTIONS ARE SUFlCIENTLY
LARGE
SCALE THEN THE POSITION OF INDIVIDUALS ACROSS THE LANDSCAPE IS OF LITTLE IMPOR
TANCE IN DETERMINING THE COMPETITIVE INTERACTIONS )N SUCH CASES THE RESULTS OF THE
MEAN
lELD MODEL ARE ASYMPTOTICALLY APPROACHED &IG 4HIS IS IN CONTRAST TO
THE CASE WHERE DISPERSAL IS LARGE
SCALE )F INTERACTIONS ARE LOCALISED IN SPACE SMALL
VALUE FOR SW THEN MANY OFFSPRING CAN ESCAPE FROM COMPETING WITH THEIR PARENT
4HIS LEADS TO A SPATIALLY SEGREGATED POPULATION WHERE MOST INDIVIDUALS EXPERIENCE
A RELATIVELY COMPETITOR
FREE NEIGHBOURHOOD AND HENCE THE lNAL POPULATION SIZE
IS MUCH LARGER THAN THAT UNDER MEAN
lELD CONDITIONS /N THE OTHER HAND IF BOTH
DISPERSAL AND INTERACTIONS ARE SHORT
RANGE SMALL VALUES FOR SM AND SW THEN THE
COMPETITIVE NEIGHBOURHOOD IS PERCEIVED AS BEING CROWDED WITH THE RESULT THAT THE
lNAL DENSITY IS MUCH LOWER THAN THE MEAN
lELD EXPECTATION )N EXTREME CASES
WHERE BOTH INTERACTIONS AND DISPERSAL OCCUR OVER SHORT SPATIAL SCALES THEN SELF
DRIVEN EXTINCTION OCCURS &IG &IGS AND ,AW ET AL
3ELF
DRIVEN EXTINCTION MIGHT AT lRST THOUGHT SEEM ONLY AN INTERESTING MATH
EMATICAL ARTEFACT AND OF NO CONCERN TO INVASION ECOLOGY AFTER ALL IF AN ORGANISMS
DISPERSAL IS SO SHORT
RANGE HOW COULD IT PERSIST AT ANOTHER LOCATION AND SEND OUT
EMMIGRANTS (OWEVER IT IS ENTIRELY POSSIBLE FOR A SPECIES DISPERSAL KERNEL TO DIFFER
BETWEEN ITS NATURAL AND EXOTIC HABITATS -ANY PLANTS RELY ON SECONDARY DISPER
SAL OF SEEDS BY ANIMALS !NDRESEN "OHNING
'AESE ET AL (OSHIZAKI
ET AL -ILTON AND $EAN 'ORDON AND VAN DER 6ALK 'UITIAN
ET AL AND IF IN A NEW ENVIRONMENT THE SECONDARY DISPERSERS ARE NOT PRESENT
THE DISPERSAL KERNEL IS LIKELY TO BE MUCH SHORTER AND HENCE SELF
DRIVEN EXTINC
TION IS MUCH MORE LIKELY !NOTHER FEATURE OF THE PRESENCE OF SPATIAL AGGREGATION
OF CONSPECIlCS IS THAT WHEN LOOKING AT THE INVASION DYNAMICS OF A SPECIES INTO A
COMMUNITY IT IS NO LONGER POSSIBLE TO IGNORE THE EFFECT OF INTRASPECIlC COMPETITION
AS HAS TRADITIONALLY BEEN THE CASE -ORTON ET AL -ORTON AND ,AW
&IG %QUILIBRIUM VALUES FOR A . AND B # AS A FUNCTION OF THE SPATIAL SCALE OF THE
INTERACTION KERNEL AND DISPERSAL KERNEL 4HE RESULTS ARE OBTAINED BY NUMERICALLY INTEGRATING
THE DETERMINISTIC APPROXIMATION AND UNTIL THE CHANGE IN . OVER AN INTEGRATION STEP
FALLS BELOW /THER PARAMETERS ARE HELD CONSTANT ARE B D D
&OR COMPARISON THIS SET OF PARAMETERS NON
SPATIAL LOGISTIC MODEL GIVES AN EQUILIBRIUM
DENSITY OF
0OPULATION GROWTH IN SPACE AND TIME
$* -URRELL
4HIS IS BECAUSE EVEN THOUGH GLOBALLY THE INVADING SPECIES IS AT A LOW DENSITY
LOCALLY THE INVADER STILL @SEES A HIGH DENSITY OF CONSPECIlCS THANKS TO THE LOCAL
NATURE OF DISPERSAL AND COMPETITION "OLKER AND 0ACALA -URRELL ET AL
-URRELL AND ,AW
4HESE RESULTS ALL POINT TO THE IMPORTANCE OF THE RELATIVE SCALES OF DISPERSAL AND
NEIGHBOURHOOD INTERACTIONS TO BOTH THE TRANSIENT DYNAMICS AND lNAL POPULATION
DENSITY "OLKER AND 0ACALA %LLNER ,AW ET AL &OR MANY ANIMAL
SPECIES THE INTERACTION KERNEL IS LIKELY TO BE OVER A MUCH SHORTER DISTANCE THAN THE
DISPERSAL KERNEL (OWEVER FOR PLANTS AND OTHER SEDENTARY ORGANISMS THIS MAY NOT
BE THE CASE AND IT IS ENTIRELY POSSIBLE THAT THE EFFECTS OF DENSITY MAY BE FELT OVER
RELATIVELY LARGE SPATIAL SCALES 0ETERS %COLOGISTS HAVE PUT MUCH EFFORT INTO
MEASURING DISPERSAL RATES AND DISTANCES #LOBERT ET AL "ULLOCK ET AL
,EVINE AND -URRELL BUT RATHER LESS IS KNOWN ABOUT THE SHAPES AND SIZES
OF INTERACTION KERNELS 0URVES AND ,AW #LEARLY MODELS OF THIS TYPE SHOW
THAT THE INTERACTION KERNELS ARE POTENTIALLY VERY IMPORTANT IN DETERMINING INVA
SION DYNAMICS
,).+3 4/ /4(%2 -/$%,3
!S HAS BEEN SUGGESTED AT VARIOUS POINTS IN THIS CHAPTER THE MODELS DESCRIBED
AND ANALYSED HERE ARE CLOSELY RELATED TO A NUMBER OF OTHER SPATIAL MODELS 9OUNG
ET AL SHOWED THAT A POPULATION OF DISCRETE INDIVIDUALS INHABITING A LAND
SCAPE AND WITH LOCAL DISPERSAL AND POST
BIRTH MOVEMENT WILL QUICKLY AGGREGATE
EVEN WITHOUT ANY DENSITY DEPENDENT PROCESSES 4HE MODELS ABOVE HAVE SHOWN
THAT INCORPORATING LOCAL DENSITY
DEPENDENT PROCESSES CAN LEAD TO ANY OF THE
MAIN CLASSES OF SPATIAL STRUCTURE AGGREGATION SEGREGATION OR RANDOM PATTERNING
&URTHER THE DYNAMICAL SYSTEM AND COLLAPSES TO THE MODEL OF 9OUNG ET AL
WHEN D AND THE SPATIAL PATTERN IS NOW DESCRIBED BY ONLY THREE PAIRS OF
TERMS A
C TWO OF WHICH ARE CONCERNED WITH BIRTHS AND ONLY ONE OF WHICH IS
CONCERNED WITH THE LOSS OF PAIRS )T CAN BE EASILY SEEN THAT IN THE lRST INSTANCE THE
BIRTH TERMS WILL OUTWEIGH THE DEATH TERM AND SO AGGREGATIONS OF INDIVIDUALS WILL
DEVELOP (OWEVER IT IS NOT CLEAR THAT THE AGGREGATIONS WILL EVER STOP ALTHOUGH THE
RATE OF CHANGE WILL SLOW OVER TIME
4HE THETA
LOGISTIC MODEL 'ILPIN AND !YAYLA HAS BEEN USED WITH SOME
SUCCESS TO MODEL THE POPULATION DYNAMICS OF A NUMBER OF SPECIES 3AETHER AND
%NGEN 3AETHER ET AL A 3AETHER ET AL B 4HE MAIN BASIS FOR
THE THETA
LOGISTIC MODEL IS THAT THE MAXIMUM RATE OF POPULATION GROWTH MAY
BE ACHIEVED AT DENSITIES OTHER THAN HALF THE CARRYING CAPACITY 4URCHIN
4HE SPATIAL VERSION OF THE LOGISTIC MODEL ALSO SHOWS THIS FEATURE ,AW ET AL
AND MAY THEREFORE ACT AS A MECHANISTIC FOUNDATION FOR THE THETA
LOGISTIC MODELS
7HEN INTERACTIONS ARE LOCALISED IN SPACE HOW CROWDED THE NEIGHBOURHOOD IS
DETERMINES WHEN THE MAXIMUM GROWTH RATE IS ACHIEVED 7HEN BOTH DISPERSAL
AND INTERACTIONS OCCUR OVER SHORT
SCALES THE NEIGHBOURHOOD QUICKLY BECOMES
0OPULATION GROWTH IN SPACE AND TIME
CROWDED EVEN IF THE OVERALL DENSITY IS LOW AND THIS LEADS TO THE MAXIMUM GROWTH
RATE OCCURRING AT LESS THAN HALF THE lNAL DENSITY /N THE OTHER HAND IF DISPERSAL
IS LONG RANGE BUT INTERACTIONS ARE SHORT SCALE THE INDIVIDUALS TEND NOT TO EXPERI
ENCE MANY NEIGHBOURS UNTIL THE OVERALL DENSITY IS MUCH HIGHER SO THE MAXIMUM
POPULATION GROWTH RATE TENDS TO OCCUR AT DENSITIES THAT ARE HIGHER THAN THE HALF
THE lNAL POPULATION DENSITY ,AW ET AL
4HIS CHAPTER HAS FOCUSED ON A MODEL FOR COMPETITION INVESTIGATING THE INVASION
DYNAMICS OF A SINGLE SPECIES -ANY ECONOMICALLY IMPORTANT INVASIONS ARE BY PARA
SITESPATHOGENS AND THEORETICAL MODELS HAVE BEEN USED TO INVESTIGATE THE EFFECTS OF
LOCAL SPATIAL STRUCTURE ON INVASION DYNAMICS OF THE PESTPATHOGEN "OLKER
+EELING )N A SUSCEPTIBLE INFECTED RECOVERED 3)2 HOST PATHOGEN MODEL
WITH LOCAL DISPERSAL OF THE PATHOGEN +EELING SHOWS HOW THE REPRODUC
TIVE RATIO REQUIRED FOR INVASION IN A NETWORK WITH LOCAL AND lNITE CONNECTIONS IS
ALWAYS LARGER THAN THE MEAN lELD CASE 'ENERALLY SPEAKING THE AGGREGATION OF
HOSTS IS GOOD AT SLOWING THE RATE OF SPREAD OF A PESTPATHOGEN IF ITS DISPERSAL IS FAIRLY
SHORT RANGE AND THIS HAS IMPORTANT IMPLICATIONS FOR EVOLUTION OF VIRULENCE "OOTS
AND 3ASAKI "OOTS ET AL
#/.#,53)/.3
)NVASIONS ARE ESSENTIALLY THE STUDY OF POPULATION GROWTH IN SPACE AND TIME "OTH
DISPERSAL AND COMPETITIVE INTERACTIONS TEND TO BE LOCALISED IN SPACE AND ACTING
TOGETHER THEY MAY PRODUCE STRONG SPATIAL STRUCTURES THAT FEED BACK ONTO POPU
LATION DYNAMICS AND SO GREATLY AFFECT THE INVASION DYNAMICS 4HIS CHAPTER HAS
INVESTIGATED THE EFFECTS OF LOCAL INTERACTIONS AND LOCAL DISPERSAL ON POPULATION
GROWTH USING SPATIAL EXTENSIONS OF THE CLASSICAL LOGISTIC EQUATION &ROM A STOCHAS
TIC INDIVIDUAL
BASED MODEL A DETERMINISTIC APPROXIMATION HAS BEEN DERIVED WHICH
GREATLY AIDS THE ANALYSIS OF THE MODEL ! NUMBER OF QUALITATIVE AND QUANTITATIVE
PATTERNS EMERGE POPULATION GROWTH MAY BE EITHER FASTER OR SLOWER THAN THE
CLASSICAL MODEL THE RATIO OF DENSITY INDEPENDENT BIRTHS TO DENSITY INDEPENDENT
DEATHS 2 CAN GREATLY AFFECT THE PROBABILITY AND SPEED OF INVASION AS WELL AS THE
lNAL POPULATION SIZE POPULATION GROWTH MAY BE LINEAR RATHER THAN THE FAMIL
IAR SIGMOID SHAPE DETERMINISTIC EXTINCTION MAY OCCUR PURELY DUE TO PARENT
OFFSPRING COMPETITION MEANING THAT INTRASPECIlC COMPETITION CANNOT BE IGNORED IN
THE ANALYSIS OF INVASIONS AND THAT COUNTER
INTUITIVELY POPULATIONS WITH THE
LOWEST EQUILIBRIUM DENSITY MAY TAKE THE LONGEST TO REACH THIS STATE !LL OF THESE
PROPERTIES ARE DEPENDENT NOT ONLY ON THE ABSOLUTE SPATIAL SCALES OF NEIGHBOURHOOD
INTERACTIONS AND LOCAL DISPERSAL BUT ALSO THEIR RELATIVE SCALES )T HAS LONG BEEN
KNOWN THAT DISPERSAL IS IMPORTANT FOR THE INVASIVE ABILITY OF A SPECIES BUT THESE
RESULTS ALSO SHOW THAT 2 MAY BE JUST AS IMPORTANT AS THE INTRINSIC GROWTH RATE R
IN DETERMINING POPULATION GROWTH
$* -URRELL
!#+./7,%$'%-%.43
4HE WORK PRESENTED HERE OWES A GREAT DEAL TO 2ICHARD ,AW AND 5LF $IECKMANN
WHO SOLVED MANY OF THE DIFlCULT INITIAL PROBLEMS TO DO WITH DERIVING THE DETER
MINISTIC APPROXIMATION AND WHO HELPED AND ENCOURAGED ME TO THINK LONG AND
HARD ABOUT THE PROBLEM OF MOMENT CLOSURES ) WOULD ALSO LIKE TO THANK "EN "OLKER
#ALVIN $YTHAM $ANIEL ,LAMBI AND $REW 0URVES FOR VARIOUS HELPFUL AND INFORMA
TIVE DISCUSSIONS ALONG THE WAY AND TO -ARC #ADOTTE 2ENE 3ALINAS *ESSICA -ETCALF
AND 9VONNE "UCKLEY FOR COMMENTS ON EARLIER DRAFTS OF THIS CHAPTER 4HIS WORK
WAS PARTLY SUPPORTED BY THE .ATURAL %NVIRONMENT 2ESEARCH #OUNCIL AND ALSO THE
#ENTRAL 3CIENCE ,ABORATORY $%&2! )MPERIAL #OLLEGE 0ARALLEL #OMPUTING #ENTRE
)#0# KINDLY DONATED COMPUTER PROCESSOR TIME THAT HELPED TO GENERATE SOME OF
THE RESULTS REPORTED IN THE lGURES
!00%.$)8
4HE PRESENCE OF THE THIRD MOMENT IN THE DYNAMICS FOR THE SECOND MOMENTS MEANS
THAT THE DYNAMICAL SYSTEM IS NOT YET CLOSED )N PRINCIPLE IT IS POSSIBLE TO DERIVE THE
DYNAMICS OF THE THIRD MOMENTS BUT APART FROM CAUSING A HEADACHE DUE TO THE
NUMBER OF TERMS INVOLVED THEY WOULD THEMSELVES BE FUNCTIONS OF FOURTH MOMENTS
QUAD DENSITIES (OW TO DEAL WITH THIS MOMENT HIERARCHY IS THE CENTRAL PROBLEM
TO THIS TECHNIQUE #LASSICAL THEORY EFFECTIVELY TRUNCATES AT THE lRST MOMENT BY
ASSUMING THAT THERE IS NO SPATIAL STRUCTURE BUT THIS ONLY DEALS WITH A LIMITING CASE
OF THE )"- SEE BELOW ! BETTER SOLUTION IS TO CLOSE THE HIERARCHY AT THE SECOND
MOMENTS BY REPLACING THE THIRD MOMENTS BY A TERM THAT IS A FUNCTION OF THE lRST
AND SECOND MOMENTS 2ECENT WORK HAS SUGGESTED A FORM OF SECOND ORDER MOMENT
CLOSURE THAT GIVES GOOD APPROXIMATIONS TO THE )"- FOR A WIDE RANGE OF SPATIAL
STRUCTURES ,AW ET AL -URRELL AND ,AW -URRELL ET AL AND IT IS
USED HERE
# j # j # j # j n j # j # j n j
4 j j n . !
. . .
4HIS CLOSURE DOES HOWEVER MEAN THAT ANALYTICAL SOLUTIONS TO THE DYNAMICAL
SYSTEM ARE VERY HARD TO ACHIEVE AND SO THE ANALYSIS OF THE DYNAMICAL SYSTEM PRO
CEEDS BY NUMERICAL METHODS
2%&%2%.#%3
!NDRESEN % 3EED DISPERSAL BY MONKEYS AND THE FATE OF DISPERSED SEEDS IN A 0ERUVIAN
RAIN FOREST "IOTROPICA
"OERLIJST - # AND 0 (OGEWEG 3PIRAL 7AVE 3TRUCTURE IN 0RE
"IOTIC %VOLUTION n
(YPERCYCLES 3TABLE AGAINST 0ARASITES 0HYSICA $
0OPULATION GROWTH IN SPACE AND TIME
"OHNING 'AESE + " ( 'AESE AND 3 " 2ABEMANANTSOA )MPORTANCE OF PRIMARY
AND SECONDARY SEED DISPERSAL IN THE -ALAGASY TREE #OMMIPHORA GUILLAUMINI %COLOGY
"OLKER " AND 3 7 0ACALA 5SING MOMENT EQUATIONS TO UNDERSTAND STOCHASTICALLY
DRIVEN SPATIAL PATTERN FORMATION IN ECOLOGICAL SYSTEMS 4HEORETICAL 0OPULATION "IOLOGY
"OLKER " - !NALYTIC MODELS FOR THE PATCHY SPREAD OF PLANT DISEASE "ULLETIN OF
-ATHEMATICAL "IOLOGY
"OLKER " - #OMBINING ENDOGENOUS AND EXOGENOUS SPATIAL VARIABILITY IN ANALYTICAL
POPULATION MODELS 4HEORETICAL 0OPULATION "IOLOGY
"OLKER " - AND 3 7 0ACALA 3PATIAL MOMENT EQUATIONS FOR PLANT COMPETITION
5NDERSTANDING SPATIAL STRATEGIES AND THE ADVANTAGES OF SHORT DISPERSAL !MERICAN
.ATURALIST
"OLKER " - 3 7 0ACALA AND # .EUHAUSER 3PATIAL DYNAMICS IN MODEL PLANT COM
MUNITIES 7HAT DO WE REALLY KNOW !MERICAN .ATURALIST
"OOTS - 0 * (UDSON AND ! 3ASAKI ,ARGE SHIFTS IN PATHOGEN VIRULENCE RELATE TO
HOST POPULATION STRUCTURE 3CIENCE
"OOTS - AND ! 3ASAKI @3MALL WORLDS AND THE EVOLUTION OF VIRULENCE INFECTION
OCCURS LOCALLY AND AT A DISTANCE 0ROCEEDINGS OF THE 2OYAL 3OCIETY OF ,ONDON 3ERIES "
"IOLOGICAL 3CIENCES
"ROWN * ( AND ,OMOLINO - 6 "IOGEOGRAPHY 3INAUER !SSOCIATES 3UNDERLAND
-!
"ULLOCK * - 2 % +ENWARD AND 2 3 (AILS EDITORS $ISPERSAL ECOLOGY "LACKWELL
/XFORD
#ASWELL ( 2 ,ENSINK AND - ' .EUBERT $EMOGRAPHY AND DISPERSAL ,IFE TABLE
RESPONSE EXPERIMENTS FOR INVASION SPEED %COLOGY
#LOBERT * % $ANCHIN ! ! $HONDT AND * $ .ICHOLS EDITORS $ISPERSAL /XFORD
5NIVERSITY 0RESS /XFORD
#ODY - , AND * - /VERTON 3HORT TERM EVOLUTION OF REDUCED DISPERSAL IN ISLAND
PLANT POPULATIONS *OURNAL OF %COLOGY
#RESSIE . ! # 3TATISTICS FOR SPATIAL DATA 7ILEY .EW 9ORK
$ARLINGTON 0 * *R #ARABIDAE OF MOUNTAINS AND ISLANDS DATA ON THE EVOLUTION OF
ISOLATED FAUNAS AND ON ATROPHY OF WINGS %COLOGICAL -ONOGRAPHS
$IECKMANN 5 AND 2 ,AW 2ELAXATION PROJECTIONS AND THE METHOD OF MOMENTS
0AGES IN 5 $IECKMANN 2 ,AW AND * ! * -ETZ EDITORS 4HE 'EOMETRY OF
ECOLOGICAL INTERACTIONS SIMPLIFYING SPATIAL COMPLEXITY #AMBRIDGE 5NIVERSITY 0RESS
#AMBRIDGE
$IGGLE 0 * 3TATISTICAL ANALYSIS OF SPATIAL POINT PATTERNS !CADEMIC 0RESS ,ONDON
%LLNER 3 0 0AIR APPROXIMATION FOR LATTICE MODELS WITH MULTIPLE INTERACTION SCALES
*OURNAL OF 4HEORETICAL "IOLOGY
%THERIDGE ! - 3URVIVAL AND EXTINCTION IN A LOCALLY REGULATED POPULATION !NNALS OF
!PPLIED 0ROBABILITY
$* -URRELL
&AGAN 7 & AND * ' "ISHOP 4ROPHIC INTERACTIONS DURING PRIMARY SUCCESSION
(ERBIVORES SLOW A PLANT REINVASION AT -OUNT 3T (ELENS !MERICAN .ATURALIST
&INLAY " * 'LOBAL DISPERSAL OF FREE
LIVING MICROBIAL EUKARYOTE SPECIES 3CIENCE
&INLAY " * AND + * #LARKE 5BIQUITOUS DISPERSAL OF MICROBIAL SPECIES .ATURE
&INLAY " * ' & %STEBAN * , /LMO AND 0 ! 4YLER 'LOBAL DISTRIBUTION OF FREE
LIVING MICROBIAL SPECIES %COGRAPHY
&ILIN ) AND :IV 9 .EW THEORY OF INSULAR EVOLUTION UNIFYING THE LOSS OF DISERSABILITY
AND BODY
MASS CHANGE %VOLUTIONARY %COLOGY 2ESEARCH
'ILPIN - % AND & * !YAYLA 'LOBAL MODELS OF GROWTH AND COMPETITION 0ROCEEDINGS
OF THE .ATIONAL !CADEMY OF 3CIENCES OF THE 5NITED 3TATES OF !MERICA
'ORDON % AND ! ' VAN DER 6ALK 3ECONDARY SEED DISPERSAL IN -ONTRICHARDIA ARBO
RESCENS , SCHOTT DOMINATED WETLANDS IN LAGUNA GRANDE 6ENEZUELA 0LANT %COLOGY
'UITIAN 0 - -EDRANO AND * 'UITIAN 3EED DISPERSAL IN %RYTHRONIUM DENS
CANIS ,
,ILIACEAE VARIATION AMONG HABITATS IN A MYRMECOCHOROUS PLANT 0LANT %COLOGY
(ASSELL - 0 ( . #OMINS AND 2 - -AY 3PECIES #OEXISTENCE AND 3ELF
/RGANIZING
3PATIAL $YNAMICS .ATURE
(ILL * + 9 # #OLLINGHAM # $ 4HOMAS $ 3 "LAKELEY 2 &OX $ -OSS AND " (UNTLEY
)MPACTS OF LANDSCAPE STRUCTURE ON BUTTERmY RANGE EXPANSION %COLOGY ,ETTERS
(OLWAY $ ! ! 6 3UAREZ AND 4 * #ASE 2OLE OF ABIOTIC FACTORS IN GOVERNING SUS
CEPTIBILITY TO INVASION ! TEST WITH ARGENTINE ANTS %COLOGY
(OSHIZAKI + 7 3UZUKI AND 4 .AKASHIZUKA %VALUATION OF SECONDARY DISPERSAL
IN A LARGE
SEEDED TREE !ESCULUS TURBINATA A TEST OF DIRECTED DISPERSAL 0LANT %COLOGY
+EELING - * 4HE EFFECTS OF LOCAL SPATIAL STRUCTURE ON EPIDEMIOLOGICAL INVASIONS
0ROCEEDINGS OF THE 2OYAL 3OCIETY OF ,ONDON 3ERIES "
"IOLOGICAL 3CIENCES
+EELING - * $ ! 2AND AND ! * -ORRIS #ORRELATION MODELS FOR CHILDHOOD
EPIDEMICS 0ROCEEDINGS OF THE 2OYAL 3OCIETY OF ,ONDON 3ERIES "
"IOLOGICAL 3CIENCES
+ENKEL . # 0ATTERN OF SELF
THINNING IN JACK PINE TESTING THE RANDOM MORTALITY
HYPOTHESIS %COLOGY
+OT - - ! ,EWIS AND 0 VANDEN$RIESSCHE $ISPERSAL DATA AND THE SPREAD OF INVAD
ING ORGANISMS %COLOGY
,AW 2 AND 5 $IECKMANN ! DYNAMICAL SYSTEM FOR NEIGHBORHOODS IN PLANT COM
MUNITIES %COLOGY
,AW 2 $ * -URRELL AND 5 $IECKMANN 0OPULATION GROWTH IN SPACE AND TIME
3PATIAL LOGISTIC EQUATIONS %COLOGY
,EVIN 3 ! 4HE 0ROBLEM OF 0ATTERN AND 3CALE IN %COLOGY %COLOGY
,EVINE * - AND $ * -URRELL #OMMUNITY
LEVEL CONSEQUENCES OF SEED DISPERSAL
PATTERNS !NNUAL 2EVIEW OF %COLOGY %VOLUTION AND 3YSTEMATICS
0OPULATION GROWTH IN SPACE AND TIME
,EWIS - ! 3PREAD RATE FOR A NONLINEAR STOCHASTIC INVASION *OURNAL OF -ATHEMATICAL
"IOLOGY
,EWIS - ! AND 3 0ACALA -ODELING AND ANALYSIS OF STOCHASTIC INVASION PROCESSES
*OURNAL OF -ATHEMATICAL "IOLOGY
-ILTON 3 * AND 7 2 * $EAN 3EEDS DISPERSED IN DUNG OF INSECTIVORES AND HERBI
VORES IN SEMI
ARID SOUTHERN !FRICA *OURNAL OF !RID %NVIRONMENTS
-ORTON 2 $ AND 2 ,AW 2EGIONAL SPECIES POOLS AND THE ASSEMBLY OF LOCAL ECOLOGI
CAL COMMUNITIES *OURNAL OF 4HEORETICAL "IOLOGY
-ORTON 2 $ 2 ,AW 3 , 0IMM AND * ! $RAKE /N MODELS FOR ASSEMBLING ECOLO
GICAL COMMUNITIES /IKOS
-URRELL $ 5 $IECKMANN AND 2 ,AW /N MOMENT CLOSURES FOR POPULATION DYNAM
ICS IN CONTINUOUS SPACE *OURNAL OF 4HEORETICAL "IOLOGY
-URRELL $ * AND 2 ,AW "EETLES IN FRAGMENTED WOODLANDS A FORMAL FRAMEWORK
FOR DYNAMICS OF MOVEMENT IN ECOLOGICAL LANDSCAPES *OURNAL OF !NIMAL %COLOGY
-URRELL $ * AND 2 ,AW (ETEROMYOPIA AND THE SPATIAL COEXISTENCE OF SIMILAR COM
PETITORS %COLOGY ,ETTERS
-URRELL $ * $ 7 0URVES AND 2 ,AW 5NITING PATTERN AND PROCESS IN PLANT ECOL
OGY 4RENDS IN %COLOGY %VOLUTION
/WEN - 2 AND - ! ,EWIS (OW PREDATION CAN SLOW STOP OR REVERSE A PREY INVA
SION "ULLETIN OF -ATHEMATICAL "IOLOGY
0ALMIOTTO 0 ! $AVIES 3 * 6OGT + ! !SHTON - 3 6OGT $ * AND !SHTON 0 3
3OIL
RELATED HABITAT SPECIALIZATION IN DIPTEROCARP RAN FOREST TREE SPECIES IN "ORNEO
*OURNAL OF %COLOGY
0EARL 2 AND , * 2EAD /N THE RATE OF GROWTH OF THE POPULATION OF THE 5NITED 3TATES
SINCE AND ITS MATHEMATICAL REPRESENTATION 0ROCEEDINGS OF THE .ATIONAL !CADEMY
OF 3CIENCES OF THE 5NITED 3TATES OF !MERICA
0ETERS ( ! .EIGHBOUR
REGULATED MORTALITY THE INmUENCE OF POSITIVE AND NEGATIVE
DENSITY DEPENDENCE ON TREE POPULATIONS IN SPECIES
RICH TROPICAL FORESTS %COLOGY ,ETTERS
0URVES $ 7 AND 2 ,AW %XPERIMENTAL DERIVATION OF FUNCTIONS RELATING GROWTH OF
!RABIDOPSIS THALIANA TO NEIGHBOUR SIZE AND DISTANCE *OURNAL OF %COLOGY
2ICHARDSON $ - . !LLSOPP # - $!NTONIO 3 * -ILTON AND - 2EJMANEK
0LANT INVASIONS n THE ROLE OF MUTUALISMS "IOLOGICAL 2EVIEWS
2IETKERK - - # "OERLIJST & VAN ,ANGEVELDE 2 (ILLE2IS,AMBERS * VAN DE +OPPEL
, +UMAR ( ( 4 0RINS AND ! - DE 2OOS 3ELF
ORGANIZATION OF VEGETATION IN
ARID ECOSYSTEMS !MERICAN .ATURALIST
3AETHER " % AND 3 %NGEN 0ATTERN OF VARIATION IN AVIAN POPULATION GROWTH RATES
0HILOSOPHICAL 4RANSACTIONS OF THE 2OYAL 3OCIETY OF ,ONDON 3ERIES "
"IOLOGICAL 3CIENCES
3AETHER " % 3 %NGEN & &ILLI 2 !ANES 7 3CHRODER AND 2 !NDERSEN A 3TOCHASTIC
POPULATION DYNAMICS OF AN INTRODUCED 3WISS POPULATION OF THE IBEX %COLOGY
$* -URRELL
3AETHER " % 3 %NGEN 2 ,ANDE # "OTH AND - % 6ISSER B $ENSITY DEPENDENCE
AND STOCHASTIC VARIATION IN A NEWLY ESTABLISHED POPULATION OF A SMALL SONGBIRD /IKOS
3KELLAM * ' 2ANDOM DISPERSAL IN THEORETICAL POPULATIONS "IOMETRIKA
4RAVIS * - * AND # $YTHAM $ISPERSAL EVOLUTION DURING INVASIONS %VOLUTIONARY
%COLOGY 2ESEARCH
4URCHIN 0 #OMPLEX POPULATION DYNAMICS 0RINCETON 5NIVERSITY 0RESS 0RINCETON
6EIT 2 2 AND - ! ,EWIS $ISPERSAL POPULATION GROWTH AND THE !LLEE EFFECT
$YNAMICS OF THE HOUSE lNCH INVASION OF EASTERN .ORTH !MERICA !MERICAN .ATURALIST
7ITH + ! 4HE LANDSCAPE ECOLOGY OF INVASIVE SPREAD #ONSERVATION "IOLOGY
7HITTAKER 2 * )SLAND "IOGEOGRAPHY %COLOGY %VOLUTION AND #ONSERVATION /XFORD
5NIVERSITY 0RESS /XFORD
9OUNG 7 2 ! * 2OBERTS AND ' 3TUHNE 2EPRODUCTIVE PAIR CORRELATIONS AND
THE CLUSTERING OF ORGANISMS .ATURE
#HAPTER EIGHT
! GUIDE TO CALCULATING
DISCRETE
TIME INVASION RATES
FROM DATA
- ! ,EWIS - ' .EUBERT ( #ASWELL
* 3 #LARK AND + 3HEA
).42/$5#4)/.
/NE MEASURE OF BIOLOGICAL INVASIVENESS IS THE RATE AT WHICH AN ESTABLISHED INVADER
WILL SPREAD SPATIALLY IN ITS NEW ENVIRONMENT 3LOW SPREAD SIGNIlES SLOW INCREASE
IN ECOLOGICAL IMPACT WHEREAS FAST SPREAD SIGNIlES THE CONVERSE )F ONE CAN PREDICT
SPREAD RATES FROM LIFE HISTORY ATTRIBUTES SUCH AS GROWTH RATES AND DISPERSAL DIS
TANCES THEN POTENTIAL INVASIVENESS CAN BE ASSESSED BEFORE THE INVASION OCCURS
4HIS WORK WAS CONDUCTED AS PART OF THE $EMOGRAPHY AND $ISPERSAL 7ORKING 'ROUP SUP
PORTED BY THE .ATIONAL #ENTER FOR %COLOGICAL !NALYSIS AND 3YNTHESIS A #ENTER FUNDED
BY .3& $%"
THE 5NIVERSITY OF #ALIFORNIA AND THE 3ANTA "ARBARA CAMPUS
4HIS RESEARCH WAS ALSO SUPPORTED IN PART BY A #ANADA 2ESEARCH #HAIR THE .ATURAL 3CIENCES
AND %NGINEERING 2ESEARCH #OUNCIL OF #ANADA AND THE .ATIONAL 3CIENCE &OUNDATION
$%"
$%"
$%"
$-3
$-3
$%"
4HANKS TO THE $EMOGRAPHY AND $ISPERSAL 7ORKING 'ROUP MEMBERS FOR THEIR
HELP AND SUPPORT WITH THE RESEARCH
-7 #ADOTTE ET AL (EDS) #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY
3PRINGER 0RINTED IN THE .ETHERLANDS
-! ,EWIS E T AL
! PREDICTION OF THIS SORT REQUIRES MODELS FOR POPULATION SPREAD !S OUTLINED BELOW
SUCH MODELS HAVE A LONG AND DISTINGUISHED HISTORY IN QUANTITATIVE ECOLOGY
7HEREAS EARLY MATHEMATICAL MODELS FOR POPULATION SPREAD WERE PRIMAR
ILY CONCEPTUAL AND QUALITATIVE IN NATURE A NEW GENERATION OF REALISTIC MODELS IS
EMERGING 4HESE NEW MODELS ARE TIED DIRECTLY TO THE DEMOGRAPHY AND DISPERSAL OF
INDIVIDUALS (OWEVER THERE ARE NEW CHALLENGES IN THE LINKING OF THESE MODELS TO
THE BIOLOGICAL PROCESSES
!S WE WILL ILLUSTRATE IN THIS CHAPTER SPREAD RATE PREDICTIONS ARE VERY SENSITIVE TO
ASSUMPTIONS ABOUT LONG
DISTANCE DISPERSAL !RE THERE ROBUST METHODS FOR ESTIMAT
ING SPREAD RATES 4HIS IS ONE QUESTION WE WILL ADDRESS
&URTHERMORE ALMOST ALL MATHEMATICAL MODELS ASSUME THAT THE SPREAD OCCURS IN
ONE SPATIAL DIMENSION ALONG A LINE 4HIS IS NOT BECAUSE MATHEMATICIANS HAVE NOT
NOTICED THAT MOST POPULATION SPREAD TAKES PLACE IN TWO DIMENSIONS EXCEPT CASES
LIKE DISPERSAL ALONG A COASTLINE OR RIVER 2ATHER IT IS BECAUSE THE MAIN QUALITA
TIVE FEATURES OF INVASIONS ARE APPARENT IN ONE
DIMENSIONAL MODELS ALTHOUGH AS
WE WILL SHOW THE QUANTITATIVE RESULTS IN SPECIlC CASES CAN DEPEND VERY MUCH ON
THE THE DIMENSIONALITY OF THE DISPERSAL DATA AND OF THE MODEL USED FOR ANALYSIS
)N PARTICULAR WE WILL SHOW THAT THE NAIVE APPLICATION OF ONE DIMENSIONAL MODELS
TO TWO DIMENSIONAL DISPERSAL DATA WILL PRODUCE A SYSTEMATIC BIAS SOMETIMES POSI
TIVE SOMETIMES NEGATIVE IN SPREAD RATE ESTIMATES )N THIS CHAPTER WE WILL OUTLINE
NEW lTTING METHODS FOR AVOIDING THESE BIASES
-/$%,).' "!#+'2/5.$
4HE ISSUE OF SPREAD RATE FOR BIOLOGICAL INVADERS WAS CONSIDERED IN DETAIL BY 3KELLAM
WHERE &ISHERS PARTIAL DIFFERENTIAL EQUATION MODEL WAS USED TO
DESCRIBE THE RATE OF CHANGE OF LOCAL POPULATION DENSITY WITH TIME
,N N , N
lN n $
,T g ,X
(ERE NX T IS THE LOCAL POPULATION DENSITY l IS THE INTRINSIC GROWTH RATE g IS
THE CARRYING CAPACITY $ IS THE DISCUSION COElCIENT X IS THE ONE DIMENSIONAL SPACE
COORDINATE AND T IS TIME 4HE RATE OF SPREAD OF A POPULATION OBEYING THIS EQUATION
ASYMPTOTICALLY APPROACHES C 3l$ FOR LARGE TIMES +OLMOGOROV ET AL
!RONSON AND 7EINBERGER &IG 7HEREAS THE SPREAD RATE PREDICTIONS ARE
ON A LANDSCAPE SCALE THE PARAMETERS R AND $ CAN BE MEASURED ON THE INDIVIDUAL
LEVEL USING LIFE TABLE ANALYSIS AND MARK RECAPTURE &URTHERMORE COMPARISONS OF
HISTORICALLY OBSERVED INVASIVE SPREAD RATES WITH THE ASYMPTOTIC SPREAD RATE FOR
MULA HAVE HELD UP FOR A WIDE VARIETY OF SPECIES !NDOW ET AL MAKING THE
INTERPLAY BETWEEN INVASION THEORY AND SPREAD DATA A MODERN DAY SUCCESS STORY IN
QUANTITATIVE ECOLOGY 3HIGESADA AND +AWASAKI
! GUIDE TO CALCULATING DISCRETE TIME INVASION RATES FROM DATA
&IG 0OPULATION SPREAD FOR &ISHERS EQUATION ! TYPICAL SOLUTION OF &ISHERS EQUATION
ILLUSTRATES GROWTH AND SPREAD IN ONE
DIMENSIONAL LINEAR SPACE !T LEFT THE SOLUTION IS
PLOTTED FOR EQUALLY
SPACED TIME INTERVALS !T RIGHT THE GRAY AREA DENOTES THE REGION IN SPACE
WHERE THE POPULATION IS LARGER THAN A THRESHOLD LEVEL N 4HE BOUNDARIES OF THIS AREA
HAVE SLOPES EQUAL TO THE ASYMPTOTIC SPREAD RATE C 3l$ &OR THIS lGURE l g $
AND NX COS/X FOR \X\ ) "ASED ON .EUBERT AND 0ARKER
! MAJOR PROBLEM WITH &ISHERS MODEL IS THAT IT IMPOSES ONE PARTICULAR
FORM OF DISPERSAL 4HE ASSUMPTION OF DIFFUSION IN EQUATION WHICH IMPLIES
NORMALLY DISTRIBUTED DISPERSAL PROPAGULES IS OFTEN VIOLATED WHEN DISPERSAL IS
MEASURED FOR BIOLOGICAL POPULATIONS ,EWIS 7HILE THERE IS TREMENDOUS
VARIABILITY IN SUCH DISPERSAL DATA THERE IS A STRONG TENDENCY FOR THE DISTRIBUTION OF
DISPERSAL DISTANCES TO BE LEPTOKURTIC WITH A LARGER NUMBER OF DISTANCES NEAR THE
CENTER AND IN THE TAILS THAN IN A NORMAL DISTRIBUTION WITH COMPARABLE VARIANCE
4HE EFFECT OF THE LONG
DISTANCE DISPERSERS AS DESCRIBED BY THE TAILS OF THE DISTRIBU
TION OF DISTANCES IS DRAMATIC #ASWELL ET AL 0REDICTIONS FOR INVASION RATES
CAN SPEED UP BY AN ORDER OF MAGNITUDE OR MORE WHEN THE LONG
DISTANCE DISPERSERS
ARE INCLUDED +OT ,EWIS AND VAN DEN $RIESSCHE
7HEN THE DISPERSAL DISTRIBUTION IS FAR FROM NORMAL THE PARTIAL DIFFERENTIAL EQUA
TION MODEL NO LONGER SUCCES )T IS THEN NECESSARY TO DElNE A DISPERSAL KERNEL
KX Y IN ONE SPATIAL DIMENSION WHICH DESCRIBES THE PROBABILITY THAT A PROPAGULE
THAT STARTS AT Y MOVES TO THE INTERVAL X X DX BY KX Y DX 4HE UNITS FOR THE
DISPERSAL KERNEL K ARE PER UNIT LENGTH 4HE SPATIAL REGION OF INTEREST IS THE INTERVAL
1 n 7HEN MODELLING INVASIONS WE TYPICALLY ASSUME IS ARBITRARILY LARGE
)N THE ABSENCE OF IMMIGRATION FROM OUTSIDE 1 EVERY DISPERSER MUST ORIGINATE
AT SOME OTHER POINT IN SO THAT
G
KX Y DY
Hn
-! ,EWIS E T AL
)N A HOMOGENEOUS HABITAT DISPERSAL BETWEEN TWO LOCATIONS WILL ONLY DEPEND
UPON UPON THE RELATIVE LOCATIONS OF THE START AND lNISH POINTS )N THIS CASE
TO WHICH WE WILL RESTRICT OUR ATTENTION FOR THE REMAINDER OF THE CHAPTER WE
WRITE KX Y KX n Y 3HIGESADA +AWASAKI AND 4ERAMOTO EXAMINE
INVASIONS IN HETEROGENEOUS ENVIRONMENTS 4O INCLUDE POPULATION DYNAMICS
POPULATION GROWTH FROM ONE GENERATION TO THE NEXT CAN BE DESCRIBED WITH A NON
LINEAR FUNCTION
NT dNT NTGNT
WHERE G DElNES THE PER CAPITA GROWTH RATE AS A FUNCTION OF LOCAL POPULATION DENSITY
NT .ON
OVERLAPPING GENERATIONS ARE ASSUMED BUT HERE AND IN THE SUBSEQUENT
ANALYSIS THE ASSUMPTION CAN BE RELAXED TO INCLUDE MODELS WITH STAGE
STRUCTURE
SEE .EUBERT AND #ASWELL FOR DETAILS
4HE NONSPATIAL POPULATION MODEL IS THEN MODIlED TO ALLOW FOR DISPER
SAL BETWEEN REPRODUCTION EVENTS )F WE DESIGNATE THE POPULATION DENSITY AT
LOCATION X AND TIME T BY NTX THESE MODELS TAKE THE FORM OF A SCALAR INTEGRODIFFER
ENCE EQUATION
G
NTX dNTY KX n Y DY
Hn
%ARLY ANALYSES OF POPULATION SPREAD USING INTEGRODIFFERENCE EQUATIONS APPEARED
IN THE MATHEMATICAL LITERATURE PRIMARILY IN THE CONTEXT OF GENETICS 7EINBERGER
,UI A B 7EINBERGER ,UI A B
(OWEVER INTEGRODIFFERENCE EQUATIONS AND GENERALIZATIONS OF THEM ARE NOW
BEING USED BY A GROWING LIST OF ECOLOGISTS TO INVESTIGATE THE SPREAD RATE OF
ECOLOGICAL POPULATIONS +OT !LLEN ET AL !LLEN ET AL +OT ET AL
6EIT AND ,EWIS ,EWIS (ART AND 'ARDNER #LARK ET AL
#LARK (IGGINS AND 2ICHARDSON .EUBERT AND #ASWELL
.EUBERT ET AL 4AKASU ET AL 7OOLCOCK AND #OUSENS 7ITH
3CHOlELD #ASWELL ET AL #LARK ET AL -ARCHANT
.EUBERT AND 0ARKER 0OWELL AND :IMMERMANN !MONG THE RESULTS OF
THESE INVESTIGATIONS ARE TWO KEY lNDINGS &IRST INTEGRODIFFERENCE MODELS PRODUCE
A RICHER SET OF INVASION DYNAMICS THAN CAN BE GENERATED BY THE REACTION
DIFFUSION
EQUATION INCLUDING FOR EXAMPLE THE POSSIBILITY OF ACCELERATING SPREAD +OT
ET AL 3ECOND THE SHAPE OF THE DISPERSAL KERNEL ESPECIALLY THE SHAPE OF THE
TAILS WHICH DETERMINE THE PROBABILITY OF LONG
DISTANCE DISPERSAL PLAYS A CRUCIAL
ROLE IN DETERMINING THE RATE OF SPREAD
4HE EFFECT OF LONG
DISTANCE DISPERSAL ON SPREAD RATES WAS HIGHLIGHTED BY +OT
ET AL WHO lT DISPERSAL KERNELS TO A DATA SET DESCRIBING THE DISPLACEMENT OF
OF GENETICALLY MARKED $ROSOPHILA $OBZHANSKY AND 7RIGHT AND PREDIC
TIONS FOR THE CORRESPONDING ASYMPTOTIC SPREAD RATE WERE LINKED TO THE SHAPES OF
! GUIDE TO CALCULATING DISCRETE
TIME INVASION RATES FROM DATA
THE KERNELS 4HESE PREDICTED SPREAD RATES VARIED OVER AN ORDER OF MAGNITUDE
DEPENDING UPON THE FATNESS OF THE TAILS OF THE RELATED DISPERSAL KERNELS SEE &IG
&IG &ITTED FUNCTIONS TO $ PSEUDOOBSCURA DISPERSAL DATA PROVIDE INGREDIENTS FOR AN
INTEGRODIFFERENCE MODEL FOR INSECT SPREAD 4HE LEFT PANELS SHOW AVERAGE NUMBER
OF INSECTS CAUGHT PER TRAP PER DAY IN $OBZHANSKY AND 7RIGHTS EXPERIMENTS )T WAS
ASSUMED THAT DISPERSAL WAS EQUALLY LIKELY IN BOTH DIRECTIONS SO THE DISPERSAL KERNELS WERE
KX GX GnX WHERE G IS THE lTTED FUNCTION 4HE RIGHT PANEL SHOWS SIMULATIONS OF
THE INTEGRODIFFERENCE EQUATIONS 3IMULATIONS ASSUME "EVERTON
(OLT POPULATION DYNAMICS
FOR FN WITH A GEOMETRIC GROWTH RATE OF h 4HE CARRYING CAPACITY WAS SCALED TO EQUAL
ONE %ACH INTEGRODIFFERENCE WAS ITERATED FOR GENERATIONS "ASED ON +OT ET AL
(ERE IS IS ASSUMED THAT ALL REPRODUCTION AND DISPERSAL OCCURS ALONG A ONE
DIMENSIONAL STRIP
OF SUITABLE HABITAT 4HE SPREAD RATE CAN BE CALCULATED BY DIVIDING THE TOTAL DISTANCE MOVED
BY THE POPULATION FRONT BY THE GENERATIONS TAKEN TO MOVE THE DISTANCE
-! ,EWIS E T AL
4HE ISSUE OF LONG
DISTANCE DISPERSAL HOWEVER GOES BEYOND THE CHOICE OF A
PARAMETRIC DISPERSAL DISTRIBUTION TO DESCRIBE A SET OF DATA )T CAN ALSO REmECT REAL
BIOLOGICAL PROCESSES &OR EXAMPLE .EUBERT AND #ASWELL COMPUTED THE
SPREAD RATE FOR THE HERBACEOUS PLANT $IPSACUS SYLVESTRIS BASED ON DATA FROM A SEED
TRAP EXPERIMENT 7ERNER 4EASEL SEEDS ARE KNOWN TO mOAT BUT DISPERSAL
BY STREAMS OR RIVERS WAS OBVIOUSLY NOT MEASURED BY THE SEED TRAP EXPERIMENT
.EUBERT AND #ASWELL CALCULATED THE ASYMPTOTIC SPREAD RATE RESULTING FROM HYPO
THETICAL MIXTURES OF THE SEED TRAP DATA AND DISPERSAL BY WATER WITH A LONGER MEAN
DISTANCE 4HEY FOUND THAT LONG
DISTANCE DISPERSAL OF EVEN ONE SEED IN A MILLION WAS
ENOUGH TO MAKE THE SPREAD RATE DEPENDENT ON THE WATER DISPERSAL ALONE 3IMILARLY
THE SEEDS OF THE TROPICAL PLANT #ALATHEA OVANDENSIS ARE DISPERSED BY AT LEAST FOUR
SPECIES OF ANT EACH WITH ITS OWN TYPICAL DISPERSAL DISTANCE (ORVITZ AND 3CHEMSKE
.EUBERT AND #ASWELL FOUND THAT OVER OF THE ASYMPTOTIC SPREAD RATE
WAS ACCOUNTED FOR BY THE ANT SPECIES WITH THE LONGEST DISPERSAL DISTANCE EVEN
THOUGH IT DISPERSED ONLY OF THE SEEDS
#LASSICAL MODELS FOR POPULATION SPREAD LIKE AND CONSIDER THE CASE WHERE
THE SPATIAL DOMAIN IS ONE
DIMENSIONAL AND LINEAR AND A SMALL BEACHHEAD OF
INDIVIDUALS IS INTRODUCED LOCALLY 4HIS IS ONLY DIRECTLY APPLICABLE TO CASES SUCH AS
POPULATION SPREAD ALONG A ROADSIDE COASTLINE ,UBINA AND ,EVIN OR A RIVER
3PEIRS AND 'URNEY 0ACHEPSKY ET AL
,ATER WE WILL INTRODUCE MODELS THAT DESCRIBE DISPERSAL IN TWO SPATIAL DIMEN
SIONS 4HESE MODELS PRODUCE ASYMPTOTIC SPREAD RATE PREDICTIONS THAT CAN DIFFER
IN DIFFERENT DIRECTIONS IF THE DISPERSAL KERNEL IS NOT RADIALLY SYMMETRIC )N BOTH
ONE AND TWO DIMENSIONS THE DISPERSAL KERNEL PLAYS A CRUCIAL ROLE IN DETERMINING
THE ASYMPTOTIC RATE OF SPREAD 4HERE ARE MANY METHODS FOR ESTIMATING DISPERSAL
KERNELS AND FOR ESTIMATING THE PROPERTIES OF THOSE KERNELS THAT ENTER INTO THE FOR
MULAE FOR SPREAD RATE FROM DATA SEE FOR EXAMPLE 3ILVERMAN 7E WILL
DISCUSS SOME OF THESE METHODS LATER IN THE CHAPTER &IRST WE DISCUSS SOME OF THE
KINDS OF DISPERSAL DATA THAT ARE TYPICALLY COLLECTED ANDOR PUBLISHED
$)30%23!, +%2.%,3 ). /.% !.$ 47/ $)-%.3)/.3
7E lRST CONSIDER FORMS IN WHICH DATA ARE COLLECTED 4HESE FALL LOOSELY INTO TWO
KINDS DISPERSAL DATA AND DENSITY DATA $ISPERSAL DATA DESCRIBE THE LOCATION OF
DISPERSERS RELATIVE TO THE PARENTS 4HESE DATA COME FROM FOLLOWING INDIVIDUAL
DISPERSERS EG BANDING AND RECAPTURE OF BIRDS MARK AND RECAPTURE OF SEEDS
USING COLORING ANDOR RADIO
TAGGING AND IS RECORDED AS EITHER DISPLACEMENTS OR
DISPLACEMENT DISTANCES )N CONTRAST DENSITY DATA DESCRIBE THE DENSITY OF DISPERS
ERS NUMBER PER UNIT AREA OBSERVED AT A GIVEN POINT TYPICALLY AS A FUNCTION OF
DISTANCE FROM A NATAL SITE OR SOURCE OF DISPERSERS 4HESE DATA COME FROM SEED TRAPS
PHEREMONE TRAPS FOR INSECTS AND SO FORTH
)F WE ASSUME THAT THE POPULATION LIVES ALONG A ONE
DIMENSIONAL STRIP OF SUITABLE
HABITAT ALONG WHICH ALL DISPERSAL AND REPRODUCTION OCCURS THE ONE
DIMENSIONAL
! GUIDE TO CALCULATING DISCRETE
TIME INVASION RATES FROM DATA
DISPERSAL KERNEL NUMBER OF DISPERSERS PER UNIT LENGTH IS NEEDED IN EQUATION
4HIS KERNEL CAN BE lTTED EITHER DIRECTLY FROM THE ONE
DIMENSIONAL DISPERSAL DATA OR
FROM THE DENSITY DATA NUMBER OF DISPERSERS PER UNIT AREA MULTIPLIED BY THE WIDTH
OF THE STRIP 4HE CONSTRAINT THAT THE KERNEL MUST BE SCALED TO INTEGRATE TO EQUA
TION MEANS THAT THE KERNEL K WILL ACTUALLY BE INDEPENDENT OF THE WIDTH OF THE
STRIP MULTIPLICATION IS ONLY DONE FORMALLY TO ENSURE THE CORRECT UNITS FOR K
)F WE ASSUME THAT THE POPULATION LIVES IN A TWO DIMENSIONAL HABITAT IT IS
lRST NECESSARY TO EXTEND THE DElNITION OF DISPERSAL TO TWO SPATIAL DIMENSIONS
(ERE DISPERSAL IS BETWEEN POINTS X ;X X=4 AND Y ;Y Y=4 IN TWO DIMENSIONAL
SPACE 4HE THE TWO
DIMENSIONAL DISPERSAL KERNEL +X Y DESCRIBES THE PROBABIL
ITY OF A PROPAGULE WHICH STARTS AT Y MOVING TO THE RECTANGLE WITH CORNERS X AND
X DX BY +X Y DX DX 4HE SPATIAL REGION OF INTEREST IS GIVEN BY 1 )N THE CASE
OF INVASIONS THIS IS TYPICALLY ASSUMED TO BE ARBITRARILY LARGE )N THE ABSENCE OF
IMMIGRATION FROM OUTSIDE 1 EVERY DISPERSER MUST ORIGINATE AT SOME OTHER POINT
IN 1 SO THAT
G
+X Y DY
H1
)F THE KERNEL +X Y DEPENDS ONLY UPON THE RELATIVE LOCATIONS OF THE START AND lNISH
POINTS WE WRITE +X Y +X n Y
4O UNDERSTAND THE DIFFERENCE BETWEEN ONE AND TWO DIMENSIONAL DISPERSAL
KERNELS WE CONSIDER THE CASE WHERE THE DISPERSAL IS ISOTROPIC IDENTICAL IN ALL
DIRECTIONS )N THIS CASE THE TWO DIMENSIONAL DISPERSAL KERNEL + CAN BE WRITTEN AS
A FUNCTION OF THE DISPERSAL RADIUS R \X n Y\ )N A LINEAR ONE
DIMENSIONAL ENVIRON
MENT THE SCALED DISTRIBUTION OF DENSITIES DISPERSERS PER UNIT LENGTH AND A DISTRI
BUTION OF DISTANCES THAT THE DISPERSERS TRAVEL FROM THE PARENT ARE THE SAME AND
ARE GIVEN BY THE KERNEL K )N A TWO
DIMENSIONAL ENVIRONMENT THE SCALED DISTRIBU
TION OF DENSITIES DISPERSERS PER UNIT AREA GIVEN BY +R AND THE DISTRIBUTION OF
DISTANCES THAT DISPERSERS TRAVEL FROM FROM THE PARENT DISPERSERS PER UNIT LENGTH
GIVEN BY + /R+R ARE NOT THE SAME BECAUSE THERE IS MORE AREA AVAILABLE AT
DISTANCES FURTHER FROM THE PARENT
0/05,!4)/. 302%!$ ). ! /.% $)-%.3)/.!, ,).%!2 %.6)2/.-%.4
4HE $OBZHANSKY AND 7RIGHT INSECT DATA WERE COLLECTED FROM TRAPS PLACED ALONG
LINEAR TRANSECTS RADIATING FROM A POINT SOURCE 4HUS THE TRAPS GIVE A RELATIVE MEA
SURE OF THE DENSITY OF DISPERSERS NUMBER PER UNIT AREA AS A FUNCTION OF DISTANCE
4HROUGHOUT THIS CHAPTER WE WILL USE BOLDFACE 2OMAN CHARACTERS SUCH AS U AND V TO REPRE
SENT VECTORS (ERE \U\ 3U U DENOTES THE LENGTH OF THE VECTOR U AND U q V UV
UV \U\ \V\ COS e DENOTES THE @DOT PRODUCT OR PROJECTION OF ONE VECTOR ONTO THE OTHER
WHERE e IS THE ANGLE BETWEEN U AND V
+! 3CHIERENBECK AND -, !NOUCHE
FROM THE RELEASE SITE 4HE MOST REASONABLE ASSUMPTION IS THAT THE TRANSECT DATA
DESCRIBE THE RADIAL DROP
OFF IN SETTLED INSECT DENSITY IN A TWO
DIMENSIONAL HABITAT
NEXT SECTION (OWEVER FOR THE SAKE OF ILLUSTRATION IN THIS PAPER WE lRST CON
SIDER THE ASSUMPTION THAT THE INSECTS AND TRANSECTS ARE FOUND ALONG A ONE
DIMEN
SIONAL STRIP OF SUITABLE HABITAT ALONG WHICH ALL DISPERSAL AND REPRODUCTION OCCURS
NEXT SECTION 4HIS IS A STANDARD ASSUMPTION IN POPULATION SPREAD MODELS AND
IS THE ASSUMPTION MADE IN THE ORIGINAL ANALYSIS OF THE INSECT SPREAD RATES IN +OT
ET AL !S WE WILL SHOW THE DIFFERENT ASSUMPTIONS ABOUT THE HABITAT AND
DISPERSAL GIVE RISE TO QUITE DIFFERENT SPREAD RATE ESTIMATES
4HEORY
!S THE INTRODUCED BEACHHEAD OF INDIVIDUALS GROWS AND DISPERSES WE EXPECT GROWTH
OF THE RANGE BOUNDARY WITH TIME &IGURE SHOWS A TYPICAL PROGRESSION ! PLOT OF
RANGE BOUNDARY VERSUS TIME GIVES LINES WHOSE SLOPE EVENTUALLY BECOME CONSTANT
4HE EVENTUAL SLOPE OF THESE LINES THE ASYMPTOTIC RATE OF SPREAD OF THE INVASION
HENCEFORTH REFERRED TO SIMPLY AS SPREAD RATE CAN BE PREDICTED USING MATH
EMATICAL THEORY WHICH RELATES THE SLOPE TO MODEL PARAMETERS )N THIS SECTION WE
OUTLINE THE THEORY
)T IS lRST NECESSARY TO MAKE SOME ASSUMPTIONS ABOUT THE GROWTH DYNAMICS
4HE SIMPLEST POPULATION DYNAMICS EXHIBIT NO OVERCOMPENSATION OR !LLEE EFFECT
4HIS ASSUMPTION TRANSLATES INTO A GROWTH FUNCTION d IN THAT IS MONOTONICALLY
INCREASING WITH MAXIMUM PER CAPITA GROWTH RATES AT LOWEST POPULATION LEVELS
4HE ASSUMPTION THAT THE MAXIMUM PER CAPITA GROWTH RATE h OCCURS AT THE
LOWEST POSSIBLE DENSITY MEANS h G * GN FOR N ! GROWING POPULATION
REQUIRES h !S DESCRIBED ABOVE LOCAL INTRODUCTION OF INDIVIDUALS COUPLED WITH
A GROWTH RATE h AND A DISPERSAL KERNEL KZ Z X n Y IN MEANS THE POPULA
TION SPREADS AS IT GROWS AND DISPERSES &IG 7EINBERGER SHOWED THAT
UNDER THE ABOVE ASSUMPTIONS ON GROWTH DYNAMICS THE POPULATION SPREADS TO
THE RIGHT AT A RATE WHICH APPROACHES SPEED C AS THE TIME SINCE INTRODUCTION
INCREASES WHERE
C MIN LN ;h -S =
S
S
AND -S IS THE MOMENT GENERATING FUNCTION FOR THE DISPERSAL KERNEL KZ
G'
-S KZ EXPSZ DZ
Hn'
4HE PARAMETER S CAN BE UNDERSTOOD AS A MEASURE OF THE STEEPNESS OF THE WAVE
N | EXPnSZ AT THE LEADING EDGE OF THE RIGHTWARD
SPREADING POPULATION 4O lND
THE SPEED OF THE LEFTWARD SPREADING FRONT ONE SHOULD USE THE MOMENT GENERATING
FUNCTION FOR KnZ IN EQUATION
! GUIDE TO CALCULATING DISCRETE
TIME INVASION RATES FROM DATA
4HUS THE POPULATION SPREAD RATE DEPENDS ONLY UPON TWO FEATURES THE GEOMETRIC
GROWTH RATE OF THE POPULATION h AND THE SHAPE OF THE DISPERSAL KERNEL K 4HE SPEED
FOR &ISHERS EQUATION C 3l$ CAN BE REGAINED FROM EQUATIONS
BY THE
CHOICE OF K . $ AND l LOGh $ETAILS ARE GIVEN IN +OT ET AL
)N PRACTICE CALCULATION OF THE SPREAD RATE MUST BE DONE NUMERICALLY EITHER BY
USING A STANDARD MINIMIZATION ROUTINE OR BY SOLVING THE DOUBLE ROOT CONDITION
EQUATIONS EXPSC h-S AND C EXPSC h-S FOR THE WAVE SPEED C AND WAVE
STEEPNESS S &OR A SIMPLE EXAMPLE WRITTEN IN -APLE CODE SEE THE !PPENDIX
(ERE IT IS ASSUMED THAT THE KERNEL K IS EXPONENTIALLY BOUNDED SO THAT THE
MOMENT GENERATING FUNCTION CAN BE CALCULATED 7HEN THE KERNEL K HAS TAILS
THAT ARE FATTER THAN EXPONENTIAL THERE IS NO ASYMPTOTIC RATE OF SPREAD THE
SPREAD RATE CONTINUES TO INCREASE WITH INCREASING TIME +OT ET AL &IG
THIRD PANEL FROM THE TOP )N THIS SITUATION AN ALTERNATIVE DElNITION OF SPREAD RATE
BASED ON THE CHANGE IN LOCATION OF THE FURTHEST FORWARD INDIVIDUAL IN THE POPU
LATION FROM GENERATION TO GENERATION FURTHEST FORWARD VELOCITY IS APPROPRIATE
#LARK ,EWIS AND (ORVATH SEE $ISCUSSION
7HEN THE GROWTH RATE h IS KNOWN BUT THE DISPERSAL KERNEL IS UNKNOWN ESTI
MATES FOR POPULATION SPREAD RATES USING EQUATION CAN VARY WIDELY DEPENDING
UPON THE PARAMETRIC FORM OF THE KERNEL CHOSEN AS WITH THE SIMULATIONS SHOWN IN
&IG
7HEN APPROPRIATE DATA ARE AVAILABLE THIS PROBLEM CAN BE ADDRESSED BY MEANS
OF A NONPARAMETRIC ESTIMATOR FOR THE MOMENT GENERATING FUNCTION WHICH
MAKES NO ASSUMPTION ABOUT THE FORM OF THE UNDERLYING KERNEL )N THIS CASE THE
MOMENT GENERATING FUNCTION IS ESTIMATED FROM RAW ONE
DIMENSIONAL LINEAR DIS
PERSAL DISPLACEMENT DATA ZxZ.
.
-%S - EXP SZI
. I
4HE SUPERSCRIPT IS USED TO INDICATE AN EMPIRICAL ESTIMATE OF THE MOMENT GENERAT
ING FUNCTION #LARK (ORVATH AND ,EWIS (ERE THE DISPERSAL MEASUREMENTS
ARISE FROM TRACKING A SERIES OF INDIVIDUALS !S A DISPLACEMENT XI GIVES THE DISTANCE
AND DIRECTION THAT THE I
TH INDIVIDUAL MOVES "Y CONVENTION LEFTWARD MOVEMENTS
ARE ASSIGNED NEGATIVE VALUES )T IS ASSUMED THAT THE TRACKING EFFORT AND TRACKING
EFlCIENCY REMAIN CONSTANT OVER THE ENTIRE LINEAR ONE
DIMENSIONAL DISPERSAL REGION
&UJIWARA ET AL CONSIDER THE EFFECTS OF CHANGES IN SAMPLING EFFORT OR DETEC
TION PROBABILITY )N THE CASE WHERE ZxZ. ARE NONNEGATIVE DISTANCES RATHER THAN
DISPLACEMENTS THE ASSUMPTION OF A SYMMETRIC DISPERSAL KERNEL WHEREBY INDIVIDU
ALS ARE AS LIKELY TO MOVE TO THE LEFT AS TO THE RIGHT LEADS TO
.
-%S - COSH SZI
. I
+! 3CHIERENBECK AND -, !NOUCHE
3UBSTITUTION OF -%S INSTEAD OF -S IN LEADS TO AN EMPIRICALLY ESTIMATED
WAVE SPEED C% 4HIS EMPIRICALLY ESTIMATED WAVE SPEED HAS MANY NICE PROPERTIES
#LARK ET AL &OR EXAMPLE IT IS UNBIASED C% CONVERGES TO TO THE TRUE POPU
LATION SPREAD RATE C AS . A ' 4HIS IS NOT GENERALLY THE CASE WHEN PARAMETRIC
KERNELS ARE lTTED TO THE DISPERSAL DATA )N THAT CASE THE TRUE POPULATION DISPERSAL
KERNEL IS NOT KNOWN AND DIFFERENT lTTED KERNELS CAN GIVE VERY DIFFERENT WAVE SPEED
PREDICTIONS
7HEN THE NUMBER OF DATA POINTS . IS lNITE EACH EMPIRICALLY ESTIMATED WAVE
SPEED WILL BE DIFFERENT AS IT WILL DEPEND ON THE PRECISE DATA SET USED (OWEVER
THE DISTRIBUTION OF THE EMPIRICAL WAVE SPEED C% ABOUT THE TRUE WAVE SPEED C IS
APPROXIMATELY 'AUSSIAN AND THE VARIANCE OF THE 'AUSSIAN APPROACHES ZERO AS
. APPROACHES INlNITY #LARK ET AL !LTHOUGH THERE IS NO CLOSED FORM
EXPRESSION FOR THE VARIANCE IT CAN BE ESTIMATED USING BOOTSTRAPPING METHODS
#LARK ET AL 7HEN THE DATA COLLECTION OR RECORDING METHODS DO NOT PROVIDE
DISPERSAL DATA BUT HISTOGRAM DENSITY DATA ARE AVAILABLE IT MAY BE NECESSARY
TO USE SUCH A HISTOGRAM AS AN ESTIMATOR FOR THE KERNEL
4 IF j ) Z jI
K( Z I I n
OTHERWISE
WHERE 4I IS THE BIN HEIGHT FOR THE HISTOGRAM ) I ) , AND
,
- jI n jI n 4I
I
4HIS YIELDS
,
-(S - 4I EXPSjI n EXPSjI n
S I
AND SUBSTITUTION OF -(S INSTEAD OF -S IN LEADS TO A @HISTOGRAM ESTIMATOR
FOR THE WAVE SPEED C( $UE TO THE ARBITRARY NATURE OF LOCATION OF THE HISTOGRAM
BINS IT CAN BE SHOWN THAT THE HISTOGRAM ESTIMATOR DOES NOT PROVIDE AN UNBIASED
ESTIMATOR FOR THE TRUE SPEED C (OWEVER IN THE ABSENCE OF OTHER DATA THIS HISTO
GRAM ESTIMATOR IS A USEFUL ALTERNATIVE TO THE EMPIRICAL ESTIMATOR GIVEN ABOVE AND
IN PRACTICE GIVES VERY SIMILAR RESULTS !S THE SIZES OF THE BINS jI n jI n APPROACHES
ZERO THE TWO ESTIMATORS ARE IDENTICAL )N THE CASE WHERE THE HISTOGRAM DATA IS FOR
DISTANCES AS OPPOSED TO DISPLACEMENTS THE ASSUMPTION OF A SYMMETRIC REDISTRIBU
TION KERNEL CAUSES TO BE MODIlED TO
,
-(S - 4I SINHSjI n SINHSjI n
S I
! GUIDE TO CALCULATING DISCRETE
TIME INVASION RATES FROM DATA
7E NOW APPLY THE HISTOGRAM SPREAD RATE ESTIMATE AND TO THE
$OBZHANSKY AND 7RIGHT DATA UNDER THE ASSUMPTION THAT THE INSECTS AND
TRANSECTS ARE FOUND ALONG A ONE
DIMENSIONAL STRIP OF SUITABLE HABITAT ALONG WHICH
ALL DISPERSAL AND REPRODUCTION OCCURS SEE DISCUSSION IN -ODELING "ACKGROUND
4HIS ESTIMATE WHICH USES THE DATA SHOWN IN &IG GIVES A SPREAD RATE OF
KM PER YEAR 4HIS IS HIGHER THAN THE SPREAD RATE PREDICTION MADE BY USING THE
EXPONENTIAL AND 'AUSSIAN KERNELS IN &IG BUT IS SUBSTANTIALLY LOWER THAN THE
PREDICTION MADE BY THE FAT TAILED KERNEL !S WITH THE EMPIRICAL ESTIMATOR BOOT
STRAPPING GIVES THE DISTRIBUTION OF WAVE SPEEDS FROM WHICH CONlDENCE INTERVALS
CAN BE CALCULATED &IG
&IG "OOTSTRAPPING GIVES A RANGE OF POSSIBLE VALUES FOR THE HISTOGRAM WAVE SPEED
4HE HISTOGRAM WAVE SPEED ESTIMATOR EQUATIONS AND IS APPLIED TO A HISTOGRAM
BASED ON THE TOTAL NUMBER OF INSECTS CAUGHT PER TRAP STARTING AT DISTANCE ZERO lNISHING
AT DISTANCE KM WITH INTER
TRAP SPACING OF KM WERE
AND "OOTSTRAPPING WAS DONE BY RE
SAMPLING
FROM THE DISPERSAL DISTANCES WITH REPLACEMENT TO PRODUCE NEW DATA SETS
4HE WAVE SPEED WAS CALCULATED FOR EACH OF THESE NEW DATA SETS AND THE DISTRIBUTION OF
SPEEDS IS SHOWN HERE .INETY PER CENT OF THE SPEEDS FELL IN THE RANGE KMYEAR
KMYEAR #OMPARE THE ESTIMATE HERE WITH THE SPREAD RATES SIMULATED IN &IG
+! 3CHIERENBECK AND -, !NOUCHE
0/05,!4)/. 302%!$ ). 47/ $)-%.3)/.3
)NVASION IN TWO SPATIAL DIMENSIONS INVOLVES LOCAL INTRODUCTION AT A BEACHHEAD
FOLLOWED BY GROWTH AND SPREAD IN SPACE (ERE THE PROCESS CAN BE DIVIDED INTO THREE
STAGES THE INITIAL ESTABLISHMENT THE EARLY RADIAL EXPANSION UNTIL WELL ESTABLISHED
IN SPACE AND THE LATER SPREAD OF THE ESTABLISHED POPULATION &IG !LTHOUGH EACH
OF THESE THREE STAGES IS OF INTEREST BIOLOGICALLY THE FOCUS OF THIS CHAPTER IS ON ANA
LYZING THE LATER SPREAD OF AN ESTABLISHED POPULATION !T THIS STAGE WE CAN APPROXI
MATE THE INVADING FRONT BY A PLANAR FRONT MOVING WITH A WELL
DElNED SPEED
4HE CALCULATION OF SPREAD IN TWO DIMENSIONS REQUIRES A UNIT VECTOR U ;U U=4
DESCRIBING THE DIRECTION PERPENDICULAR TO THE WAVE FRONT IN WHICH THE SPREAD IS
BEING CONSIDERED &IG 4HE ASYMPTOTIC SPREAD RATE IN THE DIRECTION U IS GIVEN
BY
CU MIN LN ;h -US =
S
S
&IG ! SKETCH OF THE THREE STAGES OF POPULATION SPREAD IN A HOMOGENEOUS ENVIRONMENT
3HADED AREAS INDICATE INVADED HABITAT !S TIME PROGRESSES THE INITIAL hBEACHHEADv GROWS
AND BECOMES MORE ELLIPTICALLY SHAPED &OR LONG TIMES THE INVASION FRONT IS APPROXIMATELY
PLANAR IN ALL DIRECTIONS 4HE SPEED IN THE DIRECTION U PERPENDICULAR TO THE FRONT IS FOUND
USING THE MARGINAL DISPERSAL KERNEL IN THAT DIRECTION KUU CF EQUATION 4HE MARGINAL
KERNEL IN TURN IS FOUND BY INTEGRATING THE ORIGINAL
DIMENSIONAL DISTRIBUTION OVER
THE DIRECTION V
! GUIDE TO CALCULATING DISCRETE
TIME INVASION RATES FROM DATA
WHERE -US IS THE @DIRECTIONAL MOMENT GENERATING FUNCTION THE MOMENT GENER
ATING FUNCTION OF + EVALUATED IN THE DIRECTION OF U !PPENDIX 4WO APPROACHES
FOR CALCULATION OF THE DIRECTIONAL MOMENT GENERATING FUNCTION ARE GIVEN IN THE
THE !PPENDIX )N GENERAL THE PLANAR SPREAD RATE CU WILL DEPEND ON THE DIRECTION
U (OWEVER WHEN THE DISPERSAL KERNEL IS DIRECTIONALLY ISOTROPIC IS IDENTICAL IN
ALL DIRECTIONS THE PLANAR SPREAD RATE WILL ALSO BE ISOTROPIC 7E NOW CONSIDER
THE CASE WITH DIRECTIONAL ISOTROPY SO THAT +Z Z X n Y CAN BE REWRITTEN AS +R
R 3Z Z .OTE THAT +R IS A TWO DIMENSIONAL DENSITY FUNCTION WHICH DENOTES
THE RELATIVE NUMBER OF SEEDS PER UNIT AREA FALLING AT DISTANCE R FROM THE SOURCE
4HE AREA UNDER +R IS EQUAL TO ONE
G/ G'
+R R DRDe
Hn' Hn'
! RELATED KERNEL DENOTES THE NUMBER OF SEEDS PER UNIT LENGTH FALLING A DISTANCE R
FROM THE SOURCE 4HIS RELATED KERNEL IS FOUND BY MULTIPLYING +R BY THE PERIMETER
OF A CIRCLE OF RADIUS R TO ACCOUNT FOR THE FACT THAT AT LARGER RADII THERE IS MORE
AVAILABLE AREA FOR SEEDS TO FALL +R /R+R )F THE DATA HAVE BEEN COLLECTED IN
THIS FORM A SIMPLE RESCALING BY /R WILL TRANSFORM + TO +
7E NOW CONSIDER HOW TO CALCULATE THE PLANAR SPREAD RATE SEE EQUATION
WHEN THE KERNEL IS RADIALLY SYMMETRIC 7E ILLUSTRATE THE TWO OPTIONS THAT ARE
GIVEN IN THE !PPENDIX FOR CALCULATING THE DIRECTIONAL MOMENT GENERATING FUNC
TION FOR THIS CASE (ERE WE CONSIDER A WAVE SPREADING IN THE X
DIRECTION SO THAT
U ; =4
%VALUATE THE MARGINAL DISTRIBUTION OF + BY INTEGRATING OVER THE Z
DIRECTION
TO YIELD A ONE DIMENSIONAL DISPERSAL KERNEL THAT DESCRIBES DISPERSAL IN THE Z
DIRECTION
G'
+UZ +3Z Z DZ
Hn'
AND THEN CALCULATE THE DIRECTIONAL MOMENT GENERATING FUNCTION -U OF THE KER
NEL KUZ AND THUS THE SPEED CU SEE EQUATIONS AND FOR DETAILS
4HIS METHOD EFFECTIVELY REDUCES THE TWO DIMENSIONAL SPREAD PROBLEM TO ONE
SPATIAL DIMENSION BY lRST TAKING THE MARGINAL DISTRIBUTION OF THE DISPERSAL
KERNEL AND THEN PROCEEDING AS WITH THE ONE DIMENSIONAL CASE 4HIS APPROACH
IS CONCEPTUALLY STRAIGHTFORWARD BUT MANY MARGINAL DISTRIBUTIONS CANNOT BE
CALCULATED ANALYTICALLY EVEN FOR SIMPLE KERNELS
%VALUATE THE MOMENT GENERATING FUNCTION IN THE Z
DIRECTION SEE EQUATION
DIRECTLY AS
G' G'
-U S +R EXPSZ DZDZ
Hn'Hn'
+! 3CHIERENBECK AND -, !NOUCHE
G/ G'
+R EXPSR COSe R DRDe
H H
G'
/ +R R )SR DR
H
G'
+R )SR DR
H
AND THEN USE THIS TO CALCULATE OF THE SPEED CU (ERE ) IS THE MODIlED "ESSEL
FUNCTION OF THE lRST KIND AND ZEROTH ORDER !BRAMOWIZ AND 3TEGUN
7HEN THERE ARE RAW ONE DIMENSIONAL RADIAL DISPERSAL DISTANCE DATA RxR.
THEN THE EMPIRICAL MOMENT GENERATING FOR THE PLANAR WAVE SPEED CALCULATION
BECOMES
.
-U%S - ) SRI
. I
AND SUBSTITUTION INTO GIVES THE EMPIRICAL ESTIMATOR FOR THE PLANAR WAVE SPEED
(ERE THE RI MEASUREMENTS ARISE FROM TRACKING A SERIES OF INDIVIDUALS AS THEY DIS
PERSE )T IS ASSUMED THAT THE TRACKING EFFORT PER UNIT AREA AND TRACKING EFlCIENCY
PER UNIT AREA REMAINS CONSTANT OVER THE ENTIRE DISPERSAL AREA 'IVEN A HISTOGRAM
OF RADIAL DENSITY DATA
4 IF l ) R lI
+( R I I n
OTHERWISE
WHERE ) I ) , THE CONSTRAINT THAT THE AREA UNDER THE HISTOGRAM INTEGRATES TO
ONE MEANS
,
/ -lI n lI n 4I
I
)F THE HISTOGRAM IS MEASURED IN TERMS OF THE RELATIVE NUMBER OF DISPERSERS THAT
MOVE A DISTANCE R RATHER THAN THE RELATIVE DENSITY IT IS NECESSARY TO RESCALE THE
KERNEL BY /R n AS ABOVE
4HE DIRECTIONAL MOMENT GENERATING FUNCTION FOR THE HISTOGRAM COMES FROM
SUBSTITUTING INTO TO OBTAIN
/ ,
-U(S - 4I lI ) SlI n lI n ) SlI n
S I
! GUIDE TO CALCULATING DISCRETE
TIME INVASION RATES FROM DATA
COMPARE WITH AND SUBSTITUTION INTO GIVES THE PLANAR WAVE SPEED
(ERE ) IS THE MODIlED "ESSEL FUNCTION OF THE lRST KIND AND lRST ORDER !BRAMOWIZ
AND 3TEGUN
&OR THE SAKE OF ILLUSTRATION WE EARLIER CONSIDERED THE CASE WHERE THE $OBZHANSKY
AND 7RIGHT DATA DESCRIBED INSECT MOVEMENT ALONG LINEAR TRANSECTS OF AVAILABLE
HABITAT !CCORDINGLY THE HISTOGRAM ESTIMATOR FOR THE SPREAD RATE FOR THE SITUA
TION SHOWN IN &IG WAS CALCULATED USING EQUATIONS AND AS KM
PER YEAR
4O COMPARE THE DIFFERENCE BETWEEN THE RESULTS OF THIS SECTION AND OUR EARLIER
RESULTS WE REVISIT THE $OBZHANSKY AND 7RIGHT DATA UNDER THE MORE REASONABLE
ASSUMPTION THAT THE TRANSECT DATA DESCRIBE THE RADIAL DROP
OFF IN SETTLED INSECT
DENSITY IN A TWO
DIMENSIONAL HABITAT )N THIS CASE THE HISTOGRAM ESTIMATOR FOR THE
PLANAR SPREAD RATE IS CALCULATED FROM EQUATIONS AND AS KM PER
YEAR HIGHER THAN THE PREVIOUS ESTIMATE
2EPEATING THE COMPARISON OF SPREAD RATES FOR DIFFERENT hS SHOWS THAT THE SPREAD
RATE ASSUMING RADIAL DISPERSAL AND GROWTH IN THE TWO
DIMENSIONAL HABITAT
IS SIGNIlCANTLY HIGHER THAN THE SPREAD RATE ASSUMING GROWTH AND DISPERSAL IN A
LINEAR ONE
DIMENSIONAL HABITAT SEE 4ABLE 4HIS DIFFERENCE IS MOST PRONOUNCED
FOR A LOW GROWTH RATES h 4HIS ORDERING OF SPREAD RATES $ SPEED LARGER THAN $
SPEED 4ABLE APPEARS ONLY FOR DENSITY DATA NOT DISPERSAL DATA )N FACT IF THE
ORIGINAL DATA DESCRIBE DISPERSAL THEN IT CAN BE SHOWN THAT THE ORDERING WILL BE
REVERSED WITH THE $ SPEED BASED ON GROWTH AND DISPERSAL IN A LINEAR ONE
DIMEN
SIONAL HABITAT EG EQUATION AND LARGER THAN $ SPEED BASED ON RADIAL DISPER
SAL AN GROWTH IN A TWO
DIMENSIONAL HABITAT EG EQUATION AND
4ABLE 3PREAD RATES FOR VARYING GROWTH RATES h $ISPERSAL DATA ARE AS GIVEN BY THE LEFT
PANELS IN &IG )N COLUMN LABELED $ IT IS ASSUMED THAT THE GROWTH AND DISPERSAL OCCUR IN
A $ LINEAR HABITAT 3PREAD RATES ARE CALCULATED USING EQUATIONS AND )N COLUMN
LABELED $ IT IS ASSUMED THAT THE DISPERSAL IS RADIAL AND THAT GROWTH AND DISPERSAL OCCUR
IN A $ HABITAT (ERE THE PLANAR SPREAD RATE IS CALCULATED FROM EQUATIONS AND
4HE LAST COLUMN SHOWS THE RATIO OF THE SECOND TO THE lRST COLUMNS ENTRIES 4HE DIFFERENCE
BETWEEN THE SPREAD RATES IS MOST PRONOUNCED FOR LOW GROWTH RATES
h $ SPEED KMYEAR $ SPEED KMYEAR RATIO OF SPEEDS
+! 3CHIERENBECK AND -, !NOUCHE
-/.4% #!2,/ -%4(/$3
4HE SIMPLEST ESTIMATOR OF THE MOMENT GENERATING FUNCTION IS THE EMPIRICAL
ESTIMATOR CALCULATED FROM ONE
DIMENSIONAL DISPLACEMENT DATA EQUATION
)T IS NOT ONLY SIMPLE TO IMPLEMENT NUMERICALLY BUT IN MANY CASES IS APPEALINGLY
NONPARAMETRIC )T IS SOMETIMES USEFUL TO GENERATE APPROPRIATE
$ DISPLACEMENT
DATA FROM SOME OTHER FORM USING -ONTE #ARLO METHODS 4HE RESULTING DIS
PLACEMENTS CAN THEN BE USED AS INPUT TO THE EMPIRICAL ESTIMATOR &OR EXAMPLE
#ASWELL ET AL ANALYZED THE INVASION RATES OF SEVERAL SPECIES OF %UROPEAN
BIRDS USING DISPERSAL DATA COMPILED BY VAN DEN "OSCH ET AL 4HE DISPER
SAL DATA WERE OBTAINED IN THE FORM OF HISTOGRAMS OF DISPLACEMENT DISTANCES
NOT DENSITIES 4HUS THEY WERE INTERPRETED AS GIVING A SET OF DISTANCES IN TWO
DIMENSIONAL SPACE MOVED BY A SET OF MARKED INDIVIDUALS /BTAINING THE APPROPRIATE
ONE
DIMENSIONAL DISPERSAL KERNEL REQUIRES THE MARGINAL DISTRIBUTION OF THE TWO
DIMENSIONAL DISTRIBUTION OF DISPLACEMENTS 4HIS WAS OBTAINED BY GENERATING A
LARGE NUMBER OF RANDOM DISPLACEMENT DISTANCES FROM THE HISTOGRAM ASSUMING
A UNIFORM DISTRIBUTION OF DISPLACEMENT DISTANCE WITHIN EACH HISTOGRAM BIN
!SSUMING TWO
DIMENSIONAL ISOTROPY EACH OF THESE DISTANCES WAS ASSIGNED A
DIRECTION UNIFORMLY DISTRIBUTED BETWEEN AND / 4HIS PRODUCED A SET OF ARTIlCIAL
DIMENSIONAL DISPLACEMENT DATA THE DISTANCE COMPONENT OF WHICH MATCHES THE
REPORTED HISTOGRAM 4HE MARGINALIZED DISTRIBUTION WAS EASILY GENERATED BY TAKING
THE Z COMPONENT OF EACH OF THE POINTS 4HE RESULTING SET OF DISTANCES WAS THEN
INPUT TO THE EMPIRICAL MOMENT
GENERATING FUNCTION TO PRODUCE AN ESTIMATE OF
WAVE SPEED
4HIS -ONTE #ARLO METHOD IS APPROPRIATE BECAUSE WE KNOW MATHEMATICALLY
THAT PROVIDING THE DISPERSAL KERNEL IS EXPONENTIALLY BOUNDED THE EMPIRICAL
ESTIMATOR FOR THE WAVE SPEED IS UNBIASED IE APPROACHES THE TRUE WAVE SPEED AS
. A' $ETAILS ON CONVERGENCE CAN BE FOUND IN #LARK ET AL
$)3#533)/.
4HE FOCUS OF THIS PAPER IS METHODS TO RELIABLY CONNECT POPULATION SPREAD RATE
THEORY TO BIOLOGICAL DATA 4HERE ARE TWO FEATURES THAT WE FOCUS ON I SENSITIVITY OF
SPREAD RATE ESTIMATES TO MODEL ASSUMPTIONS ABOUT LONG
DISTANCE DISPERSAL AND II
MODEL lTTING ISSUES THAT ARISE FROM lTTING TWO
DIMENSIONAL DISPERSAL DATA TO ONE
DIMENSIONAL MODELS 4HE EMPIRICAL AND HISTOGRAM ESTIMATORS PROVIDE A METHOD
TO BYPASS ASSUMPTIONS ABOUT LONG DISTANCE DISPERSAL 4HESE ARE REALLY @WHAT YOU
SEE IS WHAT YOU GET ESTIMATORS )NDEED FORMALLY SUBSTITUTING THE OBSERVED SUM OF
POINT DISPERSAL JUMPS KZ - .I
bZ n ZI INTO EQUATION LEADS TO THE EMPIRI
CAL MOMENT GENERATING FUNCTION 7HEREAS THE ABOVE SUM OF DELTA FUNCTIONS
PROVIDES A POOR ESTIMATOR FOR THE KERNEL THE PROCESS LEADS TO THE EMPIRICAL MOMENT
GENERATING FUNCTION AND A GOOD ESTIMATE FOR THE SPREAD RATE WHICH IS THE QUANTITY
OF INTEREST 4HE HISTOGRAM ESTIMATOR BEHAVES IN A MANNER WHICH IS SIMILAR TO THE
! GUIDE TO CALCULATING DISCRETE
TIME INVASION RATES FROM DATA
EMPIRICAL ESTIMATOR ALTHOUGH THE LOCATION OF BINS CAN PRODUCE SMALL BIASES IN THE
SPREAD RATE ESTIMATE
7E RECOMMEND THAT THE EMPIRICAL AND HISTOGRAM SPREAD RATE ESTIMATORS SHOULD
BE A BASIC TOOL FOR ANY MODELING EXERCISE WHICH INVOLVES SPREAD RATES WHERE LONG
DISTANCE DISPERSAL PLAYS A ROLE &ITTED PARAMETRIC DISPERSAL KERNELS MAY BE PREFER
ABLE WHEN THERE IS REASON TO BELIEVE THAT EXTRAPOLATION OF THE DISPERSAL FUNCTION
BEYOND THE FURTHEST OBSERVED DISPERSAL DISTANCE CAN BE JUSTIlED (OWEVER EVEN
IN THIS CASE IT IS USEFUL TO COMPARE SPREAD RATE RESULTS FROM THE PARAMETRIC lTTED
KERNEL WITH THE EMPIRICAL OR HISTOGRAM SPREAD RATES 4HE DIFFERENCE BETWEEN THE
SPREAD RATES WILL HIGHLIGHT THE IMPLICATIONS OF ASSUMPTIONS ABOUT LONG DISTANCE
DISPERSAL DISTANCES THAT GO INTO THE PARAMETRIC KERNEL
4HE NEW MODEL lTTING ISSUE WE FOCUS ON IS THE CORRECT METHOD TO CALCULATE PLA
NAR POPULATION SPREAD FROM RADIALLY SYMMETRIC DISPERSAL DATA .AIVELY lTTING THE
LINEAR ONE
DIMENSIONAL DISPERSAL KERNEL TO THE RADIALLY SYMMETRIC DISPERSAL DATA
WILL GIVE THE WRONG SPREAD RATE WHEN THE ONE
DIMENSIONAL SPREAD MODEL IS USED
BIASED DOWNWARD FOR DENSITY DATA AND UPWARD FOR DISPERSAL DATA
)N A LINEAR ONE
DIMENSIONAL ENVIRONMENT THE SCALED DISTRIBUTION OF DENSITIES
DISPERSERS PER UNIT LENGTH AND A DISTRIBUTION OF DISTANCES THAT THE DISPERSERS
TRAVEL FROM THE PARENT ARE THE SAME AND ARE GIVEN BY THE KERNEL K )N A TWO
DIMEN
SIONAL ENVIRONMENT THE SCALED DISTRIBUTION OF DENSITIES DISPERSERS PER UNIT AREA
GIVEN BY + AND THE DISTRIBUTION OF DISTANCES THAT DISPERSERS TRAVEL FROM FROM
THE PARENT GIVEN BY + /R+R ARE NOT THE SAME BECAUSE THERE IS MORE AREA
AVAILABLE AT DISTANCES FURTHER FROM THE PARENT 4HE CORRECT METHOD FOR CALCULATING
THE SPREAD IN THE TWO
DIMENSIONAL ENVIRONMENT INVOLVES A DIRECTIONAL MOMENT
GENERATING FUNCTION OF THE TWO
DIMENSIONAL DISPERSAL KERNEL 4HIS IS FOUND EITHER
BY EVALUATING THE MOMENT GENERATING FUNCTION OF THE MARGINAL DISTRIBUTION OF
THE DISPERSAL KERNEL +R OR BY USING + R IN A MODIlED MOMENT GENERATING FUNC
TION CALCULATION
4O SEE THE DIFFERENCE BETWEEN THE LINEAR ONE
DIMENSIONAL HISTOGRAM DENSITY
KERNEL AND THE MARGINAL DISTRIBUTION OF ITS RADIALLY SYMMETRIC ANALOG WE CAN
COMPUTE THE MARGINAL DISTRIBUTION OF THE RADIALLY SYMMETRIC HISTOGRAM DENSITY
KERNEL DIRECTLY
,
K( U Z -I dI n dI 3l I n Z IF lI n ) \Z\ lI
OTHERWISE
4HE TWO KERNELS K(Z AND K(U Z ARE SHOWN IN &IG !S CAN BE SEEN FROM
THE RESULTS IN 4ABLE THE INCREASE IN NUMBER OF LONG
DISTANCE DISPERSERS IN THE
MARGINAL DISTRIBUTION SPEEDS UP THE SPREAD RATE SIGNIlCANTLY
4HE CORRECT FORMULAE FOR CALCULATING SPREAD RATES ARE GIVEN IN 4ABLES AND
4O THE BEST OF OUR KNOWLEDGE THESE FORMULAE HAVE NOT BEEN WIDELY DEVELOPED OR
DISCUSSED ELSEWHERE IN THE LITERATURE )T IS OUR HOPE THAT THESE TABLES ALONG WITH
THE DISCUSSION IN THE CHAPTER WILL FORM A USERS GUIDE TO CALCULATING SPREAD RATES
+! 3CHIERENBECK AND -, !NOUCHE
&IG (ISTOGRAM DISPERSAL KERNEL EQUATION FOR$OBZHANSKY AND 7RIGHTS
$ROSOPHILA DISPERSAL DATA UNDER THE ASSUMPTION THAT DISPERSAL OCCURS IN A LINEAR ONE
DIMENSIONAL HABITAT GREY -ARGINAL DISTRIBUTION EQUATION OF HISTOGRAM DISPERSAL
KERNEL EQUATION UNDER THE ASSUMPTION THAT DISPERSAL OCCURS IN A TWO DIMENSIONAL
HABITAT AND IS RADIALLY SYMMETRIC
! GUIDE TO CALCULATING DISCRETE TIME INVASION RATES FROM DATA
FOR DISCRETE
TIME MODELS AND IN TURN WILL LEAD TO A CLOSER CONNECTION BETWEEN
ECOLOGICAL THEORY AND DATA
7HEN THE DISPERSAL DATA COME FROM A @FAT
TAILED KERNEL KERNEL WITH NO MOMENT
GENERATING FUNCTION THEORY PREDICTS A CONSTANTLY ACCELERATING WAVE WITH AN
ASYMPTOTICALLY INlNITE SPEED (OWEVER THE SPREAD RATES BASED ON THE EMPIRICAL
MOMENT GENERATING FUNCTION EG EQUATION OR THE HISTOGRAM MOMENT GEN
ERATING FUNCTION EG EQUATION ARE lNITE 4HIS IS BECAUSE THE THESE SPREAD
RATES ARE CALCULATED FROM A lNITE NUMBER OF DISPERSAL OBSERVATIONS OR ON DENSI
TIES MEASURED OVER A lNITE REGION (ERE ADDITIONAL SAMPLING EFFORT MAY lND RARE
LONG
DISTANCE DISPERSERS OR MAY MEASURE NONZERO DENSITIES OF DISPERSERS FAR FROM
THEIR NATAL SOURCE AND THUS THE SPREAD RATE ESTIMATE CAN INCREASE WITH SAMPLING
EFFORT 7HEN THERE IS GOOD REASON TO BELIEVE THAT THE DISPERSAL KERNEL IS FAT
TAILED
OR NEARLY FAT
TAILED A MORE APPROPRIATE MEASURE OF SPREAD RATE IS THE @FURTHEST
FORWARD VELOCITY #LARK ,EWIS AND (ORVATH WHICH REMAINS BOUNDED
! MORE GENERAL DISCUSSION OF UNCERTAINTY IN SPREAD RATES ASSOCIATED WITH LONG
DISTANCE DISPERSAL IS GIVEN IN #LARK ET AL
4ABLE -ETHODS TO CALCULATE THE MOMENT GENERATING FUNCTION NEEDED FOR THE SPREAD
RATE CALCULATION EQUATION WHEN THE POPULATION LIVES IN A ONE
DIMENSIONAL LINEAR
HABITAT
$ATA -'& ;-S = %QUATION
G'
PARAMETRIC KERNEL KZ EXPSZ DZ
Hn'
.
DISPERSAL DISPLACEMENT DATA - I
EXP SZI
.
.
DISPLACEMENT DISTANCE DATA - I COSH SZI
.
,
DISPLACEMENT HISTOGRAM - I 4I EXPSjI n EXPSjI n
S
,
DISTANCE HISTOGRAM - I 4I SINHSjI n SINHSjI n j
S
+! 3CHIERENBECK AND -, !NOUCHE
4ABLE -ETHODS TO CALCULATE THE DIRECTIONAL MOMENT GENERATING FUNCTION NEEDED
FOR THE PLANAR SPREAD RATE CALCULATION EQUATION WHEN THE POPULATION LIVES IN A TWO
DIMENSIONAL HABITAT
$ATA -'& ;-XS = %QUATION
G' G'
PARAMETRIC $ KERNEL +Z EXPSZ DZDZ
Hn' Hn'
G'
PARAMETRIC RADIAL KERNEL / +R R )SR DR
H
.
RADIAL DISPLACEMENT DATA - I ) RIS
.
/ ,
RADIAL HISTOGRAM DATA - I 4I lI ) SlI n lI n ) SlI n
S
) AND ) ARE MODIFED "ESSEL FUNCTIONS OF ZEROTH AND FIRST ORDER CF !BRAMOWIZ AND 3TEGUN
!00%.$)8
#ALCULATING THE SPREAD RATE WITH MAPLE
4HE FOLLOWING IS -APLE CODE THAT CAN BE USED TO CALCULATE THE SPREAD RATE FOR A
POPULATION WITH GEOMETRIC GROWTH RATE h AND THE COMPOSITE ,APLACE DISPERSAL
KERNEL KZ P_ EXPn_\Z\ n P _ EXPn_\Z\
'LJLWV
:RUNRXWVSHHG JDPIRUWKHIROORZLQJ SDUDPHWHUYDOXHV
ODPEGD
DOSKDB
DOSKDB
! GUIDE TO CALCULATING DISCRETE
TIME INVASION RATES FROM DATA
S
GHQHWKHPRPHQWJHQHUDWLQJIXQFWLRQDQGH[SRQHQWLDO
YB DOSKDBA DOSKDBAVA
YB DOSKDBA DOSKDBAVA
IQ H[S V
JDP
IQ ODPEGD
S
YB S
YB
)LQGWKHGRXEOHURRW
HT IQ IQ
HT GLII IQV GLII IQV
IVROYH ^HTHT`^VJDP`V DOSKDB
4HE DIRECTIONAL MOMENT GENERATING FUNCTION
4HE DIRECTIONAL MOMENT GENERATING FUNCTION IS
G' G'
-US +Z EXPSZ DZDZ
Hn' Hn'
2ECALL THAT THE DISPERSAL KERNEL +Z NOW DESCRIBES THE PROBABILITY DENSITY
FOR JUMPS WHOSE DIRECTIONS AND MAGNITUDE ARE DESCRIBED BY THE VECTOR ;Z Z=4
4HE TERM U q Z UZ UZ IN EQUATION IS THE COMPONENT OF THE DISPERSAL
JUMP Z THAT LIES IN THE U
DIRECTION )F THE KERNEL HAS NO DIRECTIONAL BIAS IS ISO
TROPIC DEPENDING ONLY UPON DISTANCE AND HENCE +Z +3Z Z THEN
THE MOMENT GENERATING FUNCTION -US AND HENCE THE SPEED CU IS INDEPEN
DENT OF THE DIRECTION VECTOR U 7E CONSIDER THIS CASE IN THE SECTION 0OPULATION
3PREAD IN 4WO $IMENSIONS 4HE @DIRECTIONAL MOMENT GENERATING FUNCTION
-U CAN BE INTERPRETED AS THE MOMENT GENERATING FUNCTION OF THE KERNEL +Z
MARGINALIZED IN THE DIRECTION OF THE UNIT VECTOR V WHICH IS PERPENDICULAR TO U
G'
-US KUU EXPSU DU
Hn'
WHERE
G'
KUU +Z DV
Hn'
AND U U q Z V V q Z
+! 3CHIERENBECK AND -, !NOUCHE
2%&%2%.#%3
!BRAMOWIZ - AND ) 3TEGUN (ANDBOOK OF -ATHEMATICAL &UNCTIONS $OVER
0UBLICATIONS )NC
!LLEN % * , * 3 !LLEN AND 8 'ILLIAM $ISPERSAL AND COMPETITION MODELS FOR
PLANTS *OURNAL OF -ATHEMATICAL "IOLOGY
!LLEN , * 3 % * !LLEN AND 3 0ONWEERA ! MATHEMATICAL MODEL FOR WEED DISPER
SAL AND CONTROL "ULLETIN OF -ATHEMATICAL OF "IOLOGY
!NDOW $ ! 0 - +AREIVA 3 ! ,EVIN AND ! /KUBO 3PREAD OF INVADING ORGAN
ISMS ,ANDSCAPE %COLOGY
!RONSON $ ' AND ( & 7EINBERGER .ONLINEAR DI L USION IN POPULATION GENETICS
COMBUSTION AND NERVE PULSE PROPAGATION )N * ! 'OLDSTEIN %D ,ECTURE .OTES IN
-ATHEMATICS 6OLUME PP
"ERLIN 3PRINGER
6ERLAG
#ASWELL ( 2 ,ENSINK AND - ' .EUBERT $EMOGRAPHY AND DISPERSAL #OMPARING
INVASION SPEEDS USING LIFE TABLE RESPONSE EXPERIMENTS %COLOGY
#LARK * 3 7HY TREES MIGRATE SO FAST #ONFRONTING THEORY WITH DISPERSAL BIOLOGY
AND THE PALEORECORD 4HE !MERICAN .ATURALIST
#LARK * 3 # &ASTIE ' (URTT 3 4 *ACKSON # *OHNSON ' ! +ING - ,EWIS * ,YNCH
3 0ACALA # 0RENTICE % 7 3CHUPP 4 7 ))) AND 0 7YCOFF 2EIDS PARADOX OF
RAPID PLANT MIGRATION "IO3CIENCE
#LARK * 3 , (ORVATH AND - ! ,EWIS /N THE ESTIMATION OF SPREAD RATE FOR A BIO
LOGICAL POPULATION 3TATISTICS AND 0ROBABILITY ,ETTERS
#LARK * 3 - ! ,EWIS AND , (ORVATH )NVASION BY EXTREMES 0OPULATION SPREAD
WITH VARIATION IN DISPERSAL AND REPRODUCTION !MERICAN .ATURALIST
#LARK * 3 - ! ,EWIS * 3 -C,ACHLAN AND * ( 2IS,AMBERS %STIMATING POPULA
TION SPREAD 7HAT CAN WE FORECAST AND HOW WELL %COLOGY
$OBZHANSKY 4 AND 3 7RIGHT 'ENETICS OF NATURAL POPULATIONS X $ISPERSION RATES
IN DROSOPHILA PSEUDOOBSCURA 'ENETICS
&ISHER 2 ! 4HE WAVE OF ADVANCE OF ADVANTAGEOUS GENES !NN %UGEN ,ONDON
&UJIWARA - + !NDERSON ( #ASWELL AND - ' .EUBERT /N THE ESTIMATION OF
DISPERSAL KERNELS FROM INDIVIDUAL CAPTURE
RECAPTURE DATA %NVIRONMENTAL AND %COLOGICAL
3TATISTICS )N PRESS
(ART $ 2 AND 2 ( 'ARDNER ! SPATIAL MODEL FOR THE SPREAD OF INVADING ORGANISMS
SUBJECT TO COMPETITION * -ATH "IOL
(IGGINS 3 ) AND $ - 2ICHARDSON 0REDICTING PLANT MIGRATION RATES IN A CHANGING
WORLD THE ROLE OF LONG
DISTANCE DISPERSAL !M .AT
(ORVITZ # # AND $ 7 3CHEMSKE 3EED DISPERSAL OF A NEOTROPICAL MYRMECOCHORE
VARIATION IN REMOVAL RATES AND DISPERSAL DISTANCE "IOTROPICA
+OLMOGOROV ! 0ETROVSKY AND . ) 0ISCOUNOV TUDE DE LQUATION DE LA DIFFUSION
AVEC CROISSANCE DE LA QUANTIT DE MATIRE ET SON APPLICATION A UN PROBLME BIOLOGIQUE
"ULL -OSCOW 5NIV -ATH -ECH
+OT - $ISCRETE
TIME TRAVELLING WAVES %COLOGICAL EXAMPLES * -ATH "IOL
! GUIDE TO CALCULATING DISCRETE
TIME INVASION RATES FROM DATA
+OT - - ! ,EWIS AND 0 VAN DEN $RIESSCHE $ISPERSAL DATA AND THE SPREAD OF
INVADING ORGANISMS %COLOGY
,EWIS - ! 6ARIABILITY 0ATCHINESS AND *UMP $ISPERSAL IN THE 3PREAD OF AN
)NVADING 0OPULATION #HAPTER PP
0RINCETON .EW *ERSEY 0RINCETIN 5NIVESITY
0RESS
,UBINA * ! AND 3 ! ,EVIN 4HE SPREAD OF A REINVADING SPECIES RANGE EXPANSION
IN THE #ALIFORNIA 3EA /TTER !MERICAN .ATURALIST
,UI 2 A ! NONLINEAR INTEGRAL OPERATOR ARISING FROM A MODEL IN POPULATION GENETICS
) MONOTONE INITIAL DATA 3)!- * -ATH !NAL
,UI 2 B ! NONLINEAR INTEGRAL OPERATOR ARISING FROM A MODEL IN POPULATION GENETICS
)) )NITIAL DATA WITH COMPACT SUPPORT 3)!- * -ATH !NAL
,UI 2 ! NONLINEAR INTEGRAL OPERATOR ARISING FROM A MODEL IN POPULATION GENETICS
))) )NITIAL DATA WITH COMPACT SUPPORT 3)!- * -ATH !NAL
,UI 2 ! NONLINEAR INTEGRAL OPERATOR ARISING FROM A MODEL IN POPULATION GENETICS
)6 #LINES 3)!- * -ATH !NAL
,UI 2 A "IOLOGICAL GROWTH AND SPREAD MODELED BY SYSTEMS OF RECURSIONS )
-ATHEMATICAL THEORY -ATHEMATICAL "IOSCIENCES
,UI 2 B "IOLOGICAL GROWTH AND SPREAD MODELED BY SYSTEMS OF RECURSIONS ))
"IOLOGICAL THEORY -ATHEMATICAL "IOSCIENCES
-ARCHANT 3 !NALYSIS OF AN INTEGRODIFFERENCE MODEL FOR BIOLOGICAL INVASIONS WITH
A QUASI
LOCAL INTERACTION -ASTERS THESIS 5NIVERSITY OF "RITISH #OLUMBIA
.EUBERT - ' AND ( #ASWELL $EMOGRAPHY AND DISPERSAL #ALCULATION AND
SENSITIVITY ANALYSIS OF INVASION SPEED FOR STAGE
STRUCTURED POPULATIONS %COLOGY
.EUBERT - ' - +OT AND - ! ,EWIS )NVASION SPEED IN mUCTUATING ENVIRON
MENTS 0ROC 2OY 3OC ,OND "
.EUBERT - ' AND ) 0ARKER 5SING INTEGRODIFFERENCE EQUATIONS TO PROJECT RATES
OF SPREAD FOR INVASIVE SPECIES 2ISK !NALYSIS IN PRESS
0ACHEPSKY % & ,UTSCHER AND - ! ,EWIS 0ERSISTENCE SPREAD AND THE DRIFT PARA
DOX 4HEORETICAL 0OPULATION "IOLOGY IN PRESS
0OWELL * AND . % :IMMERMANN !NALYSIS OF ACTIVE SEED DISPERSAL CONTRIBUTES TO
RESOLVING 2EIDS PARADOX %COLOGY
3CHOlELD 0 3PATIALLY EXPLICIT MODELS OF 4URELLI
(OFFMANN WOLBACHIA INVASIVE
WAVE FRONTS *OURNAL OF 4HEORETICAL "IOLOGY
3HIGESADA . AND + +AWASAKI "IOLOGICAL )NVASIONS 4HEORY AND 0RACTICE /XFORD
/XFORD 5NIVERSITY 0RESS
3HIGESADA . + +AWASAKI AND % 4ERAMOTO 4RAVELING PERIODIC WAVES
IN HETEROGENEOUS ENVIRONMENTS 4HEOR 0OP "IOL
3ILVERMAN " 7 $ENSITY %STIMATION FOR 3TATISTICS AND $ATA !NALYSIS #HAPMAN
AND (ALL
3KELLAM * ' 2ANDOM DISPERSAL IN THEORETICAL POPULATIONS "IOMETRIKA
3PEIRS $ # AND 7 3 # 'URNEY 0OPULATION PERSISTENCE IN RIVERS AND ESTUARIES
%COLOGY
+! 3CHIERENBECK AND -, !NOUCHE
4AKASU & . 9AMAMOTO + +AWASAKI + 4OGASHI AND . 3HIGESADA -ODELING
THE RANGE EXPANSION OF AN INTRODUCED TREE DISEASE "IOLOGICAL )NVASIONS
VAN DEN "OSCH & 2 (ENGEVELD AND * ! * -ETZ !NALYSING THE VELOCITY OF ANIMAL
RANGE EXPANSION *OURNAL OF "IOGEOGRAPHY
6EIT 2 2 AND - ! ,EWIS $ISPERAL POPULATION GROWTH AND THE ALLEE EFFECT
$YNAMICS OF THE HOUSE lNCH INVASION OF EASTERN NORTH !MERICA 4HE !MERICAN
.ATURALIST
7EINBERGER ( & ,ONG TIME BEHAVIOR OF A CLASS OF BIOLOGICAL MODELS 3)!- JOURNAL
ON MATHEMATICAL ANALYSIS
7EINBERGER ( & ,ONG TIME BEHAVIOUR OF A CLASS OF BIOLOGICAL MODELS )N 7 %
& ))) %D )N 2ESEARCH .OTES IN -ATHEMATICS .O PP -ARSHlELD -!
0ITMAN 0UBLISHING ,TD
7ERNER 0 ! ! SEED TRAP FOR DETERMINING PATTERNS OF SEED DEPOSITION IN TERRESTRIAL
PLANTS #ANADIAN *OURNAL OF "OTANY
7ITH + ! 4HE LANDSCAPE ECOLOGY OF INVASIVE SPREAD #ONSERVATION "IOLOGY
7OOLCOCK * , AND 2 #OUSENS ! MATHEMATICAL ANALYSIS OF FACTORS AFFECTING THE RATE
OF SPREAD OF ANNUAL WEEDS IN AN ARABLE lELD 7EED 3CIENCE
#HAPTER NINE
4HE ROLE OF EVOLUTIONARY
GENETICS IN STUDIES OF
PLANT INVASIONS
+ ! 3CHIERENBECK AND - , !NOUCHE
).42/$5#4)/.
)NVASIVE SPECIES HAVE LONG RAISED THE INTEREST OF EVOLUTIONISTS AS THEY PROVIDE IDEAL
SYSTEMS TO UNDERSTAND THE EVOLUTIONARY PROCESSES THAT DETERMINE AND ACCOMPANY
THE EXPANSION OF A SPECIES "AKER AND 3TEBBINS 0RAGMATICALLY INVASIONS
OF NON
NATIVE SPECIES OR GENOTYPES POSE A MAJOR BIOLOGICAL THREAT TO NATIVE BIO
DIVERSITY AND ECOSYSTEM FUNCTIONING 7HILE THE ECOLOGY OF THESE INVASIONS HAS
RECEIVED CONSIDERABLE INTERNATIONAL ATTENTION UNDERSTANDING THE EVOLUTIONARY
DIMENSIONS OF THIS PROBLEM REMAINS RUDIMENTARY 4HE OBSERVATION CAN BE MADE
HOWEVER THAT INVASIONS HAVE TWO MAIN EVOLUTIONARY CONSEQUENCES lRST THEY
CAN RESULT IN THE RAPID EVOLUTION OF INTRODUCED POPULATIONS IN NEW ENVIRONMENTS
AND SECONDLY THEY CAN PROMOTE RETICULATE GENE mOW WITH RELATED TAXA AT THE
POPULATION ECOTYPE OR SPECIES LEVEL -OONEY AND #LELAND "OTH PROCESSES
WILL RESULT IN NEW GENOTYPE
BY
ENVIRONMENT INTERACTIONS AND SUBSEQUENT GENETIC
DIVERSIlCATION
$ESPITE MUCH CONJECTURE ABOUT THE GENETIC CHARACTERISTICS OF INVASIVE SPECIES
DATA COLLECTION HAS RESULTED IN AMBIGUOUS CONCLUSIONS !LLOZYME DATA INDICATE
"ROMUS TECTORUM CHEATGRASS MAY HAVE A hGENERAL PURPOSE GENOTYPEv .OVAK
-7 #ADOTTE ET AL (EDS) #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY
3PRINGER 0RINTED IN THE .ETHERLANDS
+! 3CHIERENBECK AND -, !NOUCHE
AND -ACK QUANTITATIVE GENETIC DATA INDICATE 3APIUM SEBIFERUM #HINESE
TALLOW MAY UNDERGO POST
INTRODUCTORY ADAPTATIONS 3IEMANN AND 2OGERS
AND OTHER SPECIES MAY REQUIRE A NOVEL COMBINATION OF INTRA
OR INTERSPECIlC GENET
IC VARIATION FROM THE HOME RANGE TO PRODUCE A SUCCESSFUL INVADER e.g., 4AMARIX
SPECIES 'ASKIN AND 3CHAAL
)T IS BECOMING CLEAR THAT INVASIONS CAN RESULT IN RAPID EVOLUTIONARY EVENTS VIA
A NUMBER OF GENETIC MECHANISMS %LLSTRAND AND 3CHIERENBECK ,EE
4HE RECENT DEVELOPMENT OF GENETIC AND GENOMIC APPROACHES HAS ALLOWED THE INVES
TIGATION OF MODEL OR CULTIVATED SYSTEMS AT THE SEQUENCE LEVEL $ATA HAVE ACCUMU
LATED RAPIDLY IN THE PAST FEW YEARS TO ELUCIDATE MOLECULAR LEVEL MECHANISMS THAT
PROMOTE SPECIES FORMATION AND EXPANSION 2IESEBERG AND 7ENDEL !MONG
THE QUESTIONS THAT CAN NOW BE ADDRESSED ARE THE GEOGRAPHIC AND EVOLUTIONARY
ORIGIN OF INVASIVE SPECIES THE IDENTIlCATION OF THE PARENTAL GENOTYPES THAT ARE
INVOLVED IN THE FORMATION OF NEW RECENT INVASIVE SPECIES THE POSSIBLE GENETIC
EXCHANGES BETWEEN INVASIVE AND NATIVE SPECIES GENOME EVOLUTION OF INVASIVE
SPECIES COMPARED TO RELATED NON
INVASIVE SPECIES THE MOLECULAR BASIS OF ADAP
TIVE TRAITS THAT MAY PROMOTE INVASIVENESS IMPORTANCE OF NEUTRAL VERSUS
ADAPTIVE PROCESSES THAT HAVE ACCOMPANIED GENOME EVOLUTION OF INVASIVE SPECIES
!LLENDORF AND ,UNDQUIST AND ,EE ARE AMONG THOSE THAT EMPHA
SIZE THAT GENETIC CONSIDERATIONS SHOULD PLAY A LARGER ROLE IN THE DEVELOPMENT OF
INVASIVE SPECIES MANAGEMENT AND CONTROL 3ADLY ADEQUATE GENETIC DATA RARELY
EXIST TO SET POLICY AT A REGIONAL OR GLOBAL LEVEL 7E HOPE TO COMPEL THE READER OF THIS
CHAPTER THAT THE GENETICS OF INVASIVE SPECIES ARE INTERESTING AND IMPORTANT FROM A
THEORETICAL EVOLUTIONARY PERSPECTIVE AND MERIT FUNDAMENTAL CONSIDERATION AT THE
MANAGEMENT LEVEL 2ESEARCHERS JUST ENTERING THE STUDY OF EVOLUTIONARY GENETICS
CAN BE ASSURED OF MANY DECADES OF WORK WITH INVASIVE SPECIES AT THE MOLECULAR
POPULATION OR SPECIES LEVEL (OWEVER UNFORTUNATE THE CIRCUMSTANCE SPECIES INVA
SIONS ARE IDEAL FOR THE PROVISION OF SYNTHESES BETWEEN ECOLOGY AND GENETICS
2/,% /& -/,%#5,!2 %6/,54)/. ). ).6!3)6% !"),)49
0OSSIBLE HERITABLE FACTORS THAT CONTRIBUTE TO INVASIVE ABILITY HAVE BEEN CONJECTURED
SINCE $ARWIN !LTHOUGH SOPHISTICATED TECHNIQUES TO EXPLORE THE GENETICS
OF INVASIVE ABILITY ARE NOW AVAILABLE THE lELD STILL HAS FAR TO GO TO lND PRACTICAL
METHODOLOGIES TO ELUCIDATE THE COMPLEX GENETIC AND ECOLOGICAL INTERACTIONS OF
INVASIVE SPECIES 7HETHER INVASIVE PLANT SPECIES ARE hBORN OR MADEv %LLSTRAND
AND 3CHIERENBECK IF A PARTICULAR GENETIC CHARACTERISTIC IS IDENTIlED WE CAN
NOW DETERMINE WHETHER ALLELIC VARIABILITY FOR A TRAIT EXISTS AND ASSESS ITS EVOLUTION
ARY IMPORTANCE IN SYNTHESIS PATHWAYS OR GENE EXPRESSION
!S AGRICULTURAL WEED SCIENTISTS REALIZED OVER YEARS AGO WEED CONTROL GENET
ICS AND MOLECULAR BIOLOGY ARE BASICALLY AND INTIMATELY RELATED VIA STUDIES ON
CHEMICAL DISRUPTION OF COMPLEX METABOLIC PATHWAYS 'RESSEL !S A RESULT
OF ENZYMES THAT ARE BLOCKED BY HERBICIDES THE PATHWAYS IN WHICH THEY OCCUR
%VOLUTION AND PLANT INVASIONS
BECOME ELUCIDATED AND MAY SUBSEQUENTLY BECOME IMPORTANT IN TRANSGENIC EXPERI
MENTATION 'RESSEL /F PARTICULAR NOTE AGRICULTURAL SCIENTISTS USE BASIC
EVOLUTIONARY PRINCIPLES WHEN FACED WITH THE OFTEN INEVITABLE RESISTANCE PROBLEMS
THAT RESULT FROM HERBICIDE USE &OR EXAMPLE THE HERBICIDE PARAQUAT HAS BEEN SUC
CESSFUL IN A hPRETESTv FOR A GENETIC PREDISPOSITION FOR TRANSIENT DROUGHT TOLERANCE
BECAUSE BOTH DROUGHT AND HERBICIDE RESISTANCE CAUSE OXIDATIVE STRESS 'RESSEL
4ESTS FOR SCREENING FAVORABLE GENOTYPES IN CROP SITUATIONS MAY BE USEFUL IN
SCREENING UNWANTED NON
NATIVE HORTICULTURAL GENOTYPES THAT MAY CONTAIN TRAITS
SUCH AS RESISTANCE TO DROUGHT STRESS THAT ARE KNOWN TO CONTRIBUTE TO INVASIVENESS
"AZZAZ !N IMPORTANT STEP IN THE DEVELOPMENT OF A MANAGEMENT POLICY OF
INVASIVES IS THE INTEGRATION OF TECHNIQUES THAT ARE mEXIBLE ENOUGH TO RESPOND TO
GENETIC TRAITS THAT MAY CHANGE AS A RESULT OF ADAPTATION AND CONTROL 4HE ROLE OF
THE ECOLOGIST AND LAND MANAGER SHOULD NOT BE UNDERESTIMATED IN THE DETERMINA
TION OF THE ECOLOGICAL IMPORTANCE OF THE GENETIC TRAITS OF INVADERS
7E CONSIDERED lVE MAJOR ASPECTS OF THE ROLE OF MOLECULAR EVOLUTION TO INVASIVE
ABILITY &IRST THE GENETIC VARIABILITY NECESSARY TO RESPOND TO NEW ENVIRONMENTAL
PRESSURES EXISTS WITHIN SOME TAXA IE EITHER hUNTAPPEDv GENETIC VARIATION OR A
hGENERAL PURPOSE GENOTYPEv "AKER 3ECONDLY MORE RAPID EVOLUTION CAN
RESULT FROM ADAPTIVE RADIATION INTO UNINHABITED NICHES SIMILAR TO THAT FOUND WITH
ADAPTIVE RADIATION IN ISLAND ECOSYSTEMS $EPENDING ON AGE AND DISTURBANCE HIS
TORY SOME OF THESE NICHES MAY BE MORE AVAILABLE THAN OTHERS AND THUS BECOME
RECEPTACLES FOR RAPID EVOLUTIONARY CHANGE /UR THIRD TOPIC CONSIDERS AN IDEA
HYPOTHESIZED UPON FOR MANY YEARS 3TEBBINS MAINLY THAT POLYPLOIDY MAY
CREATE AN ADVANTAGE IN INVASIONS DUE TO MOLECULAR EVOLUTION OF GENE DUPLICA
TIONS AND INCREASED GENETIC VARIATION &OURTH THE CREATION OF NEW VARIATION CAN
RESULT FROM HYBRIDIZATION AT EITHER THE INTER
OR INTRASPECIlC LEVEL AND THIS GENETIC
VARIATION CAN ORIGINATE FROM REPEATED INTRODUCTIONS AND SEED BANKS ,ASTLY NOT ALL
ASPECTS OF THE EVOLUTION OF INVASIVE SPECIES RESULT FROM GENETIC VARIATION INSIGHTS
HAVE RECENTLY ACCUMULATED THAT STRESS THE IMPORTANCE OF EPIGENETIC MECHANISMS
AND THE ROLE OF EXPRESSION PLASTICITY IN SHAPING PHENOTYPES
%XISTING GENETIC VARIATION
-OLECULAR 'ENETICS
1UESTIONS REGARDING THE IMPORTANCE OF MOLECULAR MARKERS TO MEASURE RESPONSE
TO SELECTION ARE PART OF THE LARGER DEBATE OVER hNEARLY
NEUTRALv VERSUS SELECTION
IST THEORY /HTA AND 'ILLESPIE )F THE QUESTION IS @IS THE HETEROZYGOSITY OF
MOLECULAR MARKERS NECESSARY FOR A RESPONSE TO SELECTION THE ANSWER IS A QUALI
lED hNOv 2EED AND &RANKHAM HAVE GONE AS FAR AS TO SUGGEST THAT GENETIC
DIVERSITY DATA HAVE LITTLE PLACE IN THE CONTEXT OF THE INVASIVE SPECIES QUESTION
$ESPITE THIS VIEW A NUMBER OF EXAMPLES DEMONSTRATE THAT VARIABLE MOLECULAR
MARKERS CAN BE USEFUL IN UNDERSTANDING PLANT INVASIONS 'ENOTYPES THAT ARE CLEAR
LY MORE INVASIVE THAN OTHERS HAVE BEEN IDENTIlED IN 4AMARIX SALT CEDAR SPECIES
+! 3CHIERENBECK AND -, !NOUCHE
'ASKIN AND 3CHAAL AND 0HRAGMITES AUSTRALIS COMMON REED 3ALTONSTALL
)N ADDITION WE ARE NOW ABLE TO IDENTIFY VARIATION IN CHROMOSOMAL REGIONS
IF NOT FOR PARTICULAR GENES THAT CONFER A lTNESS ADVANTAGE IN INVASIONS #LAUSS AND
-ITCHELL
/LDS HAVE DEMONSTRATED A DIRECT RELATIONSHIP BETWEEN TRYPSIN
INHIBITOR LOCI IMPORTANT IN PLANT DEFENSE IN !RABIDOPSIS MOUSE
EAR CRESS AND THE
EVOLUTION OF LIFE HISTORY TRAITS ! QUANTITATIVE TRAIT LOCUS IN (ELIANTHUS PARADOXUS
PECOS SUNmOWER SUGGESTS #A
DEPENDENT SALT TOLERANCE AND HIGHER lTNESS IN A
RANGE OF ENVIRONMENTS IN THE HYBRID SPECIES ,EXER ET AL !LTHOUGH ADMIT
TEDLY THERE IS MUCH TO LEARN ABOUT THE RELATIONSHIP BETWEEN THE MOLECULAR GENETIC
VARIATION OF POPULATIONS AND SELECTION IT REMAINS A FRUITFUL AND SEMINAL AREA OF
INQUIRY PARTICULARLY WITH PLANT INVASIONS
'ENETIC MARKERS ARE OFTEN USED IN SELECTION AND OTHER GENETIC MANIPULATIONS
OF AGRICULTURAL CROPS $EKKER "ASIC POPULATION GENETICS HAVE BEEN USEFUL
IN THE SEPARATION OF TWO HYBRIDIZING !MARANTHUS PIGWEED SPECIES TO PREVENT THE
INTROGRESSION OF TRANSGENICS 7ETZEL ET AL -OLECULAR MARKERS HAVE BEEN
USEFUL IN /RYZA RICE SPECIES IN THE IDENTIlCATION OF VARIETIES THAT MAY OR MAY NOT
BE ABLE TO HYBRIDIZE AND SUBSEQUENTLY WHETHER THESE VARIETIES ARE APPROPRIATE
FOR A PARTICULAR GROWING REGION #OHEN ET AL 4HE SIMILAR APPLICATION OF
SUCH MARKERS TO INVASIVE SPECIES WILL LEAD TO REMOVAL PRIORITIZATION OF PROBLEMATIC
GENOTYPES IN PARTICULAR GEOGRAPHIC REGIONS ASSUMING THEY CAN BE VISUALLY DIF
FERENTIATED IN THE lELD
/NE OF THE PROBLEMS WITH USING EXISTING GENETIC VARIATION AS AN EXPLANATION
FOR INVASIVE ABILITY IS THAT IT OFTEN REQUIRES WIDESPREAD AND INTENSIVE SAMPLING
FROM BOTH THE NATIVE AND INTRODUCED RANGES 4HE MOST DElNITIVE WORK IN THE COM
PARISON OF NATIVE AND INTRODUCED GENOTYPES IS WITH "ROMUS TECTORUM .OVAK AND
-ACK .OVAK ET AL "ARTLETT ET AL "ROMUS TECTORUM IS A CLEIS
TOGAMOUS PLANT WITH VARIOUS HOMOZYGOUS MULTILOCUS GENOTYPES REPEATED INTRO
DUCTIONS OF DIFFERENT /LD 7ORLD GENOTYPES THAT HAVE INVADED .ORTH !MERICA
HAVE RESULTED IN THE REDISTRIBUTION OF THE GENETIC DIVERSITY AVAILABLE IN THE NATIVE
AREA OF THE SPECIES !LTHOUGH THE " TECTORUM WORK IS VERY THOROUGH FURTHER DATA
COLLECTION IS NEEDED TO DETERMINE WHETHER PREDICTIONS CAN BE MADE ABOUT INVASIVE
" TECTORUM GENOTYPES "ROAD TOLERANCES NEEDED FOR PHENOLOGY SEED SET DROUGHT
AND FREEZING ARE FOUND IN " TECTORUM AND OBVIOUSLY THESE TRAITS HAVE SOME GENETIC
BASIS &OR SPECIES LIKE " TECTORUM THAT HAVE REDUCED GENETIC VARIATION THE IDEAL
MATCH BETWEEN INVADER CHARACTERISTICS AND NEW RANGE ENVIRONMENT MAY BE
SIMPLY IDIOSYNCRATIC IE A GENOTYPE IS hPRE
ADAPTEDv FOR THE NEW RANGE /N THE
OTHER HAND MILLENIA OF SELECTIVE PROCESSES AND INBREEDING IN " TECTORUM MAY HAVE
WORKED TOGETHER TO RESULT IN THE hIDEAL INVADERv 4HESE ARE THE hGENERAL PURPOSE
GENOTYPESv OF "AKER GENOTYPES THAT CAN GROW IN A VARIETY OF DIFFERENT ENVIRON
MENTS WITH NO NEED TO UNDERGO FURTHER SELECTION 7ILLIAMS
4HERE REMAINS A LACK OF CONSENSUS REGARDING THE RELATIONSHIP OF GENETIC VARI
ABILITY TO THE INVASIVE ABILITY OF PLANTS IE SOME INVASIVE SPECIES MAINTAIN LOW
LEVELS OF GENETIC VARIATION "AUMEL ET AL 9E ET AL WHEREAS OTHERS
ARE HIGHLY VARIABLE "ARRETT AND 2ICHARDSON $ESPITE CONJECTURE ON THIS
%VOLUTION AND PLANT INVASIONS
MATTER SINCE "AKER AND 3TEBBINS EXPERIMENTAL EVIDENCE IN THIS
AREA HAS BEEN SLOW TO ACCUMULATE ! NUMBER OF AUTHORS HAVE FOUND THAT INVASIVE
SPECIES WITH LOW LEVELS OF GENETIC DIVERSITY OFTEN SHARE TRAITS SUCH AS INBREEDING
OR AN ANNUAL LIFE HABIT WITH A SELF
BREEDING SYSTEM "ROWN AND -ARSHALL
7ARWICK .OVAK AND -ACK 7ANG ET AL 3QUIRRELL ET AL
"ARTLETT ET AL 4HESE CONCLUSIONS AS WITH MANY OTHER CONCLUSIONS ABOUT
INVASIVE SPECIES HAVE EXCEPTIONS FOR EXAMPLE 6IOLA RIVINIANA COMMON DOG VIO
LET IS A PRIMARILY CLONAL CLEISTOGAMOUS SPECIES WITH HIGH LEVELS OF GENETIC VARIA
TION !UGE ET AL
'ROUPING SPECIES BY LIFE HISTORY TRAITS AND GEOGRAPHICAL RANGE HAS BEEN SUCCESS
FUL IN DETECTING ASSOCIATION BETWEEN GENETIC DIVERSITY AND LIFE HISTORY CHARACTERIS
TICS FOR PLANT SPECIES IN GENERAL (AMRICK ET AL ,OVELESS AND (AMRICK
(AMRICK ET AL 4HUS IT MAYBE USEFUL TO lRST CATEGORIZE INVASIVE SPECIES BY
LIFE HISTORY TRAITS AND THEN LOOK FOR PATTERNS IN GENETIC DIVERSITY 3CHIERENBECK ET AL
(AMRICK ET AL FOUND THAT WEEDY AND EARLY SUCCESSIONAL SPECIES ARE
LESS VARIABLE THAN SPECIES OF MID
AND LATE
SUCCESSIONAL STAGES ! RECENT STUDY WITH
THE HERBACEOUS PERENNIAL !LTERNANTHERA PHILOXERIODES ALLIGATOR WEED A NATIVE OF
3OUTH !MERICA SUPPORTS THIS PATTERN #ONSIDERED ONE OF THE WORLDS WORST WEEDS
! PHILOXERIODES IS A PARTIALLY SUBMERGED AQUATIC WITH WIDESPREAD CLONAL PROPAGA
TION AND VERY LOW GENETIC DIVERSITY THROUGHOUT SOUTHERN #HINA (4
8U ET AL
!MONG LONG
LIVED WOODY SPECIES FOR WHICH GENETIC VARIATION HAS BEEN MEA
SURED CORRELATIONS BETWEEN LIFE HISTORY TRAITS AND GENETIC VARIATION ARE LESS
CLEAR AND FOR MOST STUDIES ON INVASIVE WOODY ANGIOSPERMS GENETIC VARIATION IS
REPORTED FOR THE HOME RANGE ONLY &OR EXAMPLE THERE ARE HOME RANGE STUDIES FOR
2OBINIA PSEUDOACACIA BLACK LOCUST 3URLES ET AL 0ROSOPIS GLANDULOSA HONEY
MESQUITE 0ANEIDA AND #ARSTAIRS #ASUARINA CUNNINGHAMIANA RIVER
SHE
OAK -OORE AND -ORAN !CACIA MELANOXYLON BLACKWOOD WATTLE -ORAN
ET AL !CACIA DECURRENS GREEN WATTLE -ORAN ET AL AND %UCALYPTUS
OBLIQUA MESSMATE "ROWN ET AL AND ALL SHOW HIGHER LEVELS OF VARIATION
THAN EXPECTED FOR SPECIES WITH SIMILAR LIFE HISTORY TRAITS 3CHIERENBECK ET AL
!LL BUT ONE OF THESE SPECIES %UCALYPTUS OBLIQUA ARE NITROGEN lXERS AND
IT SHOULD BE NOTED THAT THE TRAIT FOR NITROGEN lXATION IS WELL
ASSOCIATED WITH ABIL
ITY TO INVADE NEW RANGES RAPIDLY 6ITOUSEK ,ONICERA JAPONICA *APANESE
HONEYSUCKLE AN INVASIVE WOODY VINE NATIVE TO !SIA AND INVASIVE THROUGHOUT
WARM TEMPERATE AND TROPICAL CLIMATES WORLDWIDE HAS LEVELS OF VARIATION WITHIN
THE RANGE EXPECTED FOR SPECIES WITH SIMILAR LIFE HISTORY TRAITS 3CHIERENBECK ET AL
!N INVASIVE CLONAL VINE OF THE SOUTHEASTERN 53 0UERARIA LOBATA KUDZU
HAS A HIGH LEVEL OF GENETIC VARIATION IN ITS INTRODUCED RANGE THAT IS SUGGESTIVE
OF MULTIPLE INTRODUCTIONS %XCESS HETEROZYGOSITY IN INTRODUCED POPULATIONS OF
0 LOBATA ALSO SUGGESTS THAT SELECTION MAY BE ACT IN FAVOR OF HETEROZYGOUS INDI
VIDUALS 0APPERT ET AL 2UBUS ALCEIFOLIUS GIANT BRAMBLE A SHRUB NATIVE TO
SOUTHEAST !SIA HAS GREATER GENETIC VARIATION IN ITS HOME RANGE REDUCED VARIATION
IN ITS NEW RANGE OF -ADAGASCAR AND A SINGLE INVASIVE GENOTYPE AS MEASURED BY
+! 3CHIERENBECK AND -, !NOUCHE
!&,0S IN ITS NEW RANGE IN THE )NDIAN /CEAN !MSELLEM ET AL )N A MORE
RECENT EXAMPLE $E7ALT FOUND #LIDEMIA HIRTA SOAPBUSH A SHRUB NATIVE
TO #ENTRAL AND 3OUTH !MERICA AND INVASIVE IN (AWAII HAD LOW LEVELS OF GENETIC
VARIATION IN BOTH THE INTRODUCED AND HOME RANGE 3URPRISINGLY TO OUR KNOWLEDGE
THERE ARE NO MORE RECENT REVIEWS THAT EXAMINE THE RELATIONSHIP BETWEEN GENETIC
VARIABILITY INVASIVENESS AND LIFE HISTORY TRAITS
2EGARDLESS OF THE EXPECTATIONS OF GENETIC VARIABILITY IN NATIVE OR INTRODUCED
POPULATIONS EFFECTIVE STUDIES OF INTRASPECIlC GENETIC VARIATION REQUIRE THE USE OF
A NUMBER OF GENES #OALESCENCE ANALYSES OF GENE GENEALOGIES ALLOW A DETERMINA
TION OF THE GEOGRAPHIC ORIGINS OF INVASIVE GENOTYPES 'ASKIN AND 3CHAAL
3CHAAL ET AL AND THE EXPANSION OF GENOTYPES DURING INVASION CAN BE
RETRACED THROUGH STAR LIKE PHYLOGENETIC PATTERNS THAT INDICATE RECENT AND RAPID
POPULATION GROWTH 3LATKIN AND (UDSON !LTHOUGH HAPLOTYPE PHYLOG
ENIES ARE USEFUL IN THE RECONSTRUCTION OF HISTORICAL AND RECENT GENE mOW PATTERNS
THE ANALYTICAL TECHNIQUES FOR THE DETERMINATION OF HISTORICAL GENE mOW PATTERNS
HAVE ONLY RECENTLY BECOME AVAILABLE 4EMPLETON #OMPLICATIONS CAN ARISE
HOWEVER IN HYBRID LINEAGES THAT FORM NEW RECOMBINING COASLESCENT COMPLEXES
1UANTITATIVE 'ENETICS
4HE LONG
TERM CONJECTURE THAT CERTAIN TRAITS ARE ASSOCIATED WITH INVASIVENESS
"AKER "AZZAZ IS NOW SUPPORTED BY EXPERIMENTAL EVIDENCE FOR
THE GENETIC BASIS OF PHENOLOGY IN #APSELLA BURSA
PASTORIS SHEPARDS PURSE .EUFFER
AND (URKA 3OLIDAGO ALTISSIMA TALL GOLDENROD 3 GIGANTEA LATE GOLDENROD
7EBER AND 3CHMID HIGH RELATIVE GROWTH RATES AND EARLY REPRODUCTION
FOR 0INUS PINE SPP 2EJMANEK AND 2ICHARDSON 'ROTKOPP ET AL
AND TRADE
OFFS BETWEEN HERBIVORE DEFENSE AND lTNESS 3TRAUSS ET AL 0ARKER
ET AL CONCLUDED BASED ON MEASURED GROWTH RATES FREEZING TOLERANCE
AND GROWTH HABITS THAT THE INVASIVENESS OF 6ERBASCUM THAPSUS MULLEIN IS MORE
LIKELY DUE TO A GENERAL PURPOSE GENOTYPE THAT IS INmUENCED MORE BY ENVIRONMENTAL
CONDITIONS THAN BY HERITABLE FACTORS
! SMALL NUMBER OF QUANTITATIVE GENETIC STUDIES OF NATIVE AND INTRODUCED
GENOTYPES HAVE FOUND POST
COLONIZATION EVOLUTION OF GENETIC TRAITS &OR EXAMPLE
IN 3APIUM SEBIFERUM INTRODUCED POPULATIONS HAD A GREATER SEED SET THAN NATIVE
POPULATIONS BUT LESS PROTECTION FROM HERBIVORY 3IEMANN AND 2OGERS ,EGER
AND 2ICE FOUND SELECTION WITHIN YEARS FOR GENETICALLY BASED GROWTH
AND REPRODUCTIVE TRAITS IN GENOTYPES OF THE #ALIFORNIA NATIVE %SCHSCHOLZIA CALIFOR
NICA #ALIFORNIA POPPY INVASIVE IN #HILE
,EE SUGGESTS INVASIVE ABILITY MAY BE MORE OF REmECTION OF ABILITY TO
RESPOND TO SELECTION THAN TO PHENOTYPIC PLASTICITY BUT THIS LEADS US BACK TO THE
QUESTION OF WHETHER INVASIVE SPECIES ARE BORN OR MADE AND DElNITIVE DATA WITH
WHICH TO ANSWER THIS QUESTION SIMPLY DO NOT EXIST
%VOLUTION AND PLANT INVASIONS
2APID EVOLUTION RESULTING FROM ADAPTIVE RADIATION
!DAPTIVE RADIATION IN THE CLASSICAL SENSE IS THE COLONIZATION AND SUBSEQUENT
DIVERSIlCATION OF SPECIES FROM A COMMON ANCESTOR INTO NEW HABITATS 4HE PROCESS
OF ADAPTIVE RADIATION IN PART INSPIRED $ARWINS THEORY OF NATURAL SELECTION AND
HAS BEEN SUPPORTED EMPIRICALLY FOR AT LEAST FOUR DECADES AT MANY SPATIAL AND TEM
PORAL SCALES 4HE RECENT SPREAD RADIATION AND EVOLUTION OF INVASIVE SPECIES LIKELY
FOLLOWS SIMILAR PROCESSES OF ADAPTIVE RADIATION 1UESTIONS REMAIN HOWEVER ABOUT
THE RAPIDITY AND SPATIAL SCALE WITH WHICH THIS PROCESS CAN OCCUR #AN FRAGMENTED
DISTURBED LANDSCAPES DEVOID OR PARTIALLY DEVOID OF NATIVE BIOTA BE CONSIDERED ISLAND
SITUATIONS AND RECEPTACLES FOR ADAPTIVE RADIATION 7E KNOW VIRTUALLY NOTHING
ABOUT WHAT EVOLUTIONARY PROCESSES WILL OCCUR IN SITUATIONS IN WHICH SPECIES FROM
REMOTE AREAS OF THE GLOBE ARE BROUGHT TOGETHER INTO A NEW HABITAT 4HE HUMAN
MEDIATED MIGRATION OF PROPAGULES IS NOT UNLIKE NON
HUMAN MEDIATED DISPERSAL
ALTHOUGH ON A DIFFERENT TEMPORAL SCALE 2EZNICK AND 'HALAMBOR REVIEWED
STUDIES TO CONCLUDE THAT THE RAPID EVOLUTION FOLLOWING COLONIZATION OF NEW
HABITATS IS PROMOTED BY NEW ECOLOGICAL CONDITIONS .OVEL ECOLOGICAL CONDITIONS IN
THEIR STUDY INCLUDED NEW FOOD RESOURCES BIOTIC OR ABIOTIC INTERACTIONS PREDATORS
AND COMPETITORS 3PECIES POOR COMMUNITIES THAT SUBSEQUENTLY BECAME VESSELS
FOR RAPID EVOLUTIONARY CHANGE OFTEN WERE A RESULT OF ANTHROPOGENIC DISTURBANCE
2EZNICK AND 'HALAMBOR 7E KNOW OF NO WORK WHICH HAS EXAMINED THE
RAPID EVOLUTIONARY CONSEQUENCES OF DISPERSAL OF A SINGLE PLANT SPECIES INTO A RANGE
OF NEW AND DIFFERENT HABITATS ALTHOUGH THERE ARE SOME ANIMAL EXAMPLES (UEY
ET AL ,OSOS ET AL 7E PRIMARILY CAN DRAW FROM EXAMPLES WHICH ILLUS
TRATE THE RAPIDITY WITH WHICH EVOLUTION CAN OCCUR FOLLOWING ADAPTIVE RADIATION
#LASSIC EXAMPLES OF ADAPTIVE RADIATION IN ISLAND HABITATS ARE NOT ONLY EXAMPLES
OF RAPID EVOLUTIONARY CHANGE BUT PROVIDE IDEAL OPPORTUNITIES TO STUDY THE GENET
ICS AND ECOLOGY OF INVASIONS !N EXAMPLE OF VERY EFFECTIVE COLONIZATION INTO NEW
UNOCCUPIED HABITATS HAS BEEN WELL
SUPPORTED IN THE MAGNIlCENT DIVERSIlCATION OF
THE (AWAIIAN SILVERSWORD ALLIANCE OVER THE LAST MILLION YEARS "ARRIER ET AL
"ARRIER ET AL #OMPARISIONS OF MUTATION RATES BETWEEN GENES IMPORTANT IN
THE REGULATION OF mORAL AND INmORESCENCE DEVELOPMENT AND NON
REGULATORY GENES
IN THE (AWAIIAN SILVERSWORD ALLIANCE PROVIDE EVIDENCE THAT ADAPTIVE RADIATION
MAY BE MORE CORRELATED WITH VARIATION IN REGULATORY LOCI "ARRIER ET AL
2ETICULATE GENE mOW CAN FACILITATE ADAPTIVE RADIATION VIA NEW GENE COMBINA
TIONS 3EEHAUSEN )F RETICULATE GENE mOW IS IMPORTANT IN THE SPREAD OF
COLONIZING SPECIES ANY GENE mOW NEEDS TO OCCUR PRIOR TO THE SPREAD BUT THERE
COULD BE REPEATED OPPORTUNITIES FOR THIS TO HAPPEN THROUGH REPEATED INTRODUC
TIONS 3UPPORT OF HYPOTHESES FOR RAPID ADAPTIVE RADIATION REQUIRES VARIATION AT
FUNCTION LOCI AND MULTIPLE hOPPORTUNITIESv FOR ADAPTIVE DIVERGENCE WITH REPEATED
INTRODUCTIONS 3EEHAUSEN 4HUS A COMBINATION OF REPEATED INTRODUCTIONS
NEW GENE COMBINATIONS AND UNOCCUPIED OR PARTIALLY lLLED NICHES RESULT CREATE
A VULNERABILITY TO INVASION FROM PREVIOUSLY UNSEEN GENOTYPES 4HE HUMAN
MEDIATED ADAPTIVE RADIATION OF PLANT SPECIES INTO NEW HABITATS PROVIDES AN IDEAL
+! 3CHIERENBECK AND -, !NOUCHE
SITUATION FOR THE EXPERIMENTAL STUDY OF HUMAN INDUCED EVOLUTIONARY CHANGE
(OWEVER HIGH RATES OF HUMAN
FACILITATED PLANT DISPERSAL MAY ALSO PREVENT OR SLOW
RADIATION BY PROMOTING GENE mOW AND PANMIXIS
(YBRIDIZATION
4HE PREVALENCE OF RETICULATE EVOLUTION THAT IS THE MERGING OF DIVERGENT GENOMES
THROUGH INTERSPECIlC GENE mOW IS KNOWN AS AN IMPORTANT EVOLUTIONARY FORCE
IN PLANTS !NDERSON AND 3TEBBINS 4HE USE OF MOLECULAR MARKERS HAS
GREATLY HELPED TO DOCUMENT ORIGINS AND OCCURRENCE OF HYBRID LINEAGES AND THE
GENETIC CONSEQUENCES OF INTROGRESSIVE HYBRIDIZATION 2IESEBERG !RNOLD
-OLECULAR PHYLOGENETIC APPROACHES THAT COMBINE MULTIPLE SEQUENCE DATA
SETS HAVE ALLOWED THE DETECTION OF ANCIENT INTROGRESSION EVENTS AND REVEAL THAT
RETICULATION IS EVEN MORE FREQUENT THAN PREVIOUSLY THOUGHT EG $OYLE ET AL
#RONN AND 7ENDEL 3MALL ET AL
(YBRIDIZATION IS RELATED TO INVASION IN TWO WAYS &IRST INTRODUCED INVASIVE
PLANTS MAY HYBRIDIZE WITH NATIVE SPECIES AND GIVE RISE TO NEW SUCCESSFUL AND
RAPIDLY EXPANDING TAXA !BBOTT !BBOTT ET AL 3ECOND HYBRIDIZATION
BETWEEN NON INVASIVE SPECIES CAN RESULT IN NEW AGGRESSIVE HYBRIDS THAT COMPETE
WITH THE PARENTS AND INVADE NEW HABITATS 2IESEBERG AND 7ENDEL 'ENETIC
INTROGRESSION BETWEEN INVASIVE SPECIES AND CLOSELY RELATED NATIVES MAY HAVE CRITI
CAL AND RAPID EVOLUTIONARY CONSEQUENCES (UXEL 4HE SALTMARSH SPECIES
3PARTINA ALTERNImORA WAS DELIBERATELY INTRODUCED FROM THE !TLANTIC !MERICAN COAST
TO #ALIFORNIA WHERE IT HYBRIDIZED WITH THE NATIVE 3 FOLIOSA $AEHLER AND 3TRONG
(YBRIDIZATION WAS SHOWN TO OCCUR IN BOTH DIRECTIONS ALTHOUGH THE INTRO
DUCED SPECIES HAS HIGHER MALE lTNESS !NTILLA ET AL 2ATHER THAN SUFFERING
FROM COMPETITION WITH 3 ALTERNImORA 3 FOLIOSA IS NOW THREATENED BY INTROGRESSANT
HYBRIDS THAT RESULT FROM RECURRENT BACKROSSES AND THAT THREATEN THE GENETIC INTEG
RITY OF THE NATIVE SPECIES !YRES ET AL 0OLLEN SWAMPING IS ALSO THOUGHT TO
HAVE REPRESENTED AN IMPORTANT MECHANISM ALLOWING HYBRIDIZATION AND INVASION
IN OAKS 0ETIT ET AL
(UMAN ACTIVITIES HAVE INCREASED ECOLOGICALLY DISTURBED AREAS BRINGING TOGETH
ER PREVIOUSLY ISOLATED TAXA AND GENERATING OPEN ARRAYS OF NICHES THAT ARE BET
TER SUITED TO HYBRIDS THAN TO THEIR PARENTS %LLSTRAND AND 3CHIERENBECK
(YBRIDS DO WELL IN DISTURBED HABITAT !NDERSON AND INVASIVE SPECIES ARE
EMPIRICALLY ASSOCIATED WITH DISTURBED ECOSYSTEMS 6ITOUSEK 4HIS IS PAR
TICULARLY WELL ILLUSTRATED IN THE TWO CLASSICAL EXAMPLES OF INTROGRESSIVE HYBRIDIZA
TION AND HYBRID SPECIATION IN THE ,OUISIANA IRISES !NDERSON !RNOLD
AND IN (ELIANTHUS SPECIES SUNmOWERS 2IESEBERG ET AL (YBRIDIZATION AND
INTROGRESSION BETWEEN )RIS HEXAGONA AND )RIS FULVA OCCUR PRIMARILY IN DISTURBED
AREAS THAT ALLOW SYMPATRY BETWEEN THE PARENTAL SPECIES (YBRID GENOTYPES DIS
PLAY VARIOUS COMBINATIONS OF PARENTAL ECOLOGICAL TRAITS SUCH AS SHADE TOLERANCE
THAT CONFER DIFFERENT lTNESSES ACROSS DIFFERENT ENVIRONMENTS !RNOLD AND
REFERENCES THEREIN 3TUDIES ON THE )RIS FULVA X ) BREVICAULIS COMPLEX HAVE ALSO
%VOLUTION AND PLANT INVASIONS
DEMONSTRATED THE IMPORTANCE OF CONSIDERING ALL LIFE STAGES IN EXPERIMENTATION
TO UNDERSTAND HYBRID EVOLUTION *OHNSTON ET AL (ELIANTHUS IS A GENUS
PARTICULARLY AFFECTED BY RETICULATE EVOLUTION INVOLVING INTROGRESSIVE HYBRIDIZA
TION AND HOMOPLOID HYBRID SPECIATION AS ILLUSTRATED BY ( ANNUUS AND ( DEBILIS
+IM AND 2IESEBERG !DDITIONALLY HYBRIDIZATION BETWEEN ( ANNUUS AND
( PETIOLARIS GAVE RISE TO THREE HOMOPLOID HYBRID SPECIES ( ANOMALUS ( DESERTI
COLA ( PARADOXUS THESE STABLE NEW LINEAGES DIPLAY NOVEL ECOLOGICAL ADAPTATIONS
4RANSGRESSIVE SEGREGATION HAS RESULTED IN EXTREME PHENOTYPES IN THESE (ELIANTHUS
SPECIES AND IS THOUGHT TO BE THE KEY FOR THEIR ABILITY TO INVADE NOVEL HABITATS
2IESEBERG ET AL e.g., AS DEMONSTRATED WITH SALT ADAPTATION IN THE HYBRID
SPECIES ( PARADOXUS ,EXER ET AL
4HERE ARE NOW MANY EXAMPLES IN WHICH HYBRID GENOTYPES ARE MORE lT THAN
ONE OR BOTH OF THE PARENTAL GENOTYPES "URKE AND !RNOLD %LLSTRAND AND
3CHIERENBECK FOUND EXAMPLES IN WHICH THE OCCURRENCE OF NEW INVASIVE
TAXA NESS WAS PRECEDED BY HYBRIDIZATION AND FOR WHICH THERE WAS STRONG MOLECU
LAR EVIDENCE 'ASKIN AND 3CHAAL PROVIDE BOTH NUCLEAR AND CP$.! EVI
DENCE THAT THE MOST COMMON INVASIVE HAPLOTYPES OF THE VORACIOUS 4AMARIX IN THE
53 ARE POST
INTRODUCTION HYBRID COMBINATIONS BETWEEN PRIMARILY 4 RAMOSISSIMA
AND 4 CHINENSIS WITH SOME ADDITIONAL GENE mOW FROM 4 PARVImORA AND 4 GALLICA
"IRCHLER ET AL SUGGEST THAT REGULATORY GENE ALLELIC INTERACTION IN HYBRID
GENOTYPES MIGHT ACCOUNT FOR THE WELL
KNOWN HETEROSIS EFFECT IE HYBRID HETEROZY
GOSITY RESULTS IN GREATER VIGOR BIOMASS SPEED OF DEVELOPMENT AND FERTILITY THAN
IN THE PARENTAL GENOTYPES &OR INSTANCE UPREGULATION OF HOUSEKEEPING GENES MAY
CAUSE GENE EXPRESSION THAT IS DIFFERENT IN HYBRIDS THAN THE MIDPARENT PREDICTIONS
&UTURE STUDIES LINKING PHENOTYPIC CHANGES AND INVESTIGATIONS AT THE GENOME
LEVEL SHOULD PROVIDE NEW INSIGHTS INTO MOLECULAR MECHANISMS THAT ARE INVOLVED
IN THE ADAPTIVE SUCCESS OF HYBRID LINEAGES
4HERE IS NOW LITTLE DOUBT THAT HYBRIDIZATION IS AN IMPORTANT EVOLUTIONARY
MECHANISM IN PLANTS AND THE CONCERN WITH HYBRIDIZATION IN INVASIVE SPECIES IS
NOT WHETHER IT CAN HAPPEN BUT THE SPEED WITH WHICH HUMANS ACCELERATE THIS
EVOLUTIONARY PROCESS (OMOGENIZATION IS A PROCESS THAT IS OCCURRING NOT ONLY AT
THE COMMUNITY LEVEL BUT ALSO AT THE GENETIC LEVEL WITHIN TAXA /LDEN ET AL
,OCALLY ADAPTED GENOTYPES ARE BECOMING LOST THROUGH HOMOGENIZATION AND INVA
SION OF DOMINANTS
0OLYPLOIDY
0OLYPLOIDY RESULTING FROM WHOLE GENOME DUPLICATION IS A WIDESPREAD EVOLUTION
ARY PHENOMENON AND A COMMON SPECIATION MECHANISM IN PLANTS 3TEBBINS
,EWIS 'RANT /NE OF THE MOST CONSPICUOUS CONTRIBUTIONS THAT HAS
RESULTED FROM THE DEVELOPMENT OF RECENT GENOMIC APPROACHES IS THE AWARENESS OF
THE PREVALENCE OF POLYPLOIDY IN MOST EUKARYOTIC LINEAGES EG 7OLFE "LANC
ET AL 4HIS HAS CONTRIBUTED TO A RENEWED INTEREST IN THE EVOLUTIONARY SUCCESS
AND POTENTIAL SELECTIVE ADVANTAGE OF GENOME DUPLICATION /TTO AND 7HITTON
+! 3CHIERENBECK AND -, !NOUCHE
-ANY POLYPLOID SPECIES ARE WELL
ADAPTED SUCCESSFUL WEEDY SPECIES WHICH REIN
FORCES THE IDEA THAT POLYPLOIDY MAY HAVE PREDISPOSED SPECIES TO BECOME INVADERS
"ROWN AND -ARSHALL "ARRETT AND 2ICHARDSON )NTERESTINGLY NEWLY
FORMED POLYPLOIDS ARE FREQUENTLY INVASIVE SPECIES WHICH SUGGESTS THAT POLYPLOIDY
CONFERS AN IMMEDIATE ECOLOGICAL APTITUDE TO INVADE NEW HABITATS )NVASIVE SPECIES
OF RECENT ORIGIN ARE EXCELLENT MODEL SYSTEMS TO INVESTIGATE THE EARLY EVOLUTIONARY
MECHANISMS ASSOCIATED WITH INVASIVENESS AND PROVIDE THE UNIQUE OPPORTUNITY
TO COMPARE THE NEW LINEAGE TO ITS PARENTS THAT ARE GENERALLY IDENTIlED AND STILL
EXTANT 4HE ALLOPOLYPLOIDS 3PARTINA ANGLICA CORDGRASS !NOUCHE ET AL A
4RAGOPOGON MIRUS AND 4 MISCELLUS GOATSBEARD 3OLTIS ET AL 3ENECIO CAMB
RENSIS 7ELSH RAGWORT AND 3 EBORACENCIS !BBOTT AND ,OWE AND #ARDAMINE
SCHULZII BITTERCRESS 5RBANSKA ET AL FORMED DURING THE LAST
YEARS
HAVE WELL
DOCUMENTED ORIGINS HAVE SPREAD RAPIDLY AND DISPLAY A LARGER ECOLOGI
CAL AMPLITUDE THAN THEIR PROGENITORS 4HESE SPECIES ARE EITHER PERENNIALS OR ANNU
ALS
BIENNIALS AND DISPLAY VARIOUS BREEDING OR POLLINATION SYSTEMS
'ENOME DUPLICATION MAY HAVE DIFFERENT IMPACTS ON FERTILITY AND MODES OF INHER
ITANCE THAT IS DEPENDENT ON CHROMOSOME BEHAVIOR AND GENETIC SEGREGATION )T IS
GENERALLY PREDICTED THAT DUPLICATION OF THE SAME GENOME WITHIN SPECIES IE STRICT
AUTOPOLYPLOIDY WILL RESULT IN RANDOM PAIRING POLYSOMIC INHERITANCE AT DUPLI
CATED LOCI IRREGULAR MEIOSIS AND THUS LIMITED FERTILITY WHEREAS THE DUPLICATION
OF MORE DIFFERENTIATED HOMOEOLOGOUS GENOMES IE ALLOPOLYPLOIDY WILL RESULT IN
PREFERENTIAL CHROMOSOME PAIRING BIVALENTS REGULAR MEIOSIS HIGH FERTILITY AND
DISOMIC INHERITANCE $A 3ILVA AND 3OBRAL )N FACT AUTOPOLYPLOIDS AND ALLO
POLYPLOIDS OCCUR ON A CONTINUUM IN NATURE AS THE ONGOING EVOLUTIONARY PROCESS
RESULTS IN MORE OR LESS DIVERGENT PARENTAL POPULATIONS OF THE POLYPLOID 3TEBBINS
7ENDEL AND $OYLE -OREOVER CHROMOSOME PAIRING MAY BE AFFECTED
BY VARIOUS GENETIC AND GENOMIC FACTORS AND VARY WITH THE AGE OF THE POLYPLOID IT
IS THEN RECOMMENDED TO DISTINGUISH BETWEEN THE MODE OF FORMATION OF A POLYPLOID
SPECIES AND ITS MODE OF CHROMOSOMAL SEGREGATION /TTO AND 7HITTON
2ECENT RESEARCH HAS RESULTED IN A PARTICULARLY DYNAMIC VISION OF POLYPLOID
GENOMES OVER BOTH A SHORT
AND LONG
TERM EVOLUTIONARY TIME SCALE 7ENDEL
3OLTIS AND 3OLTIS 4HE DEVELOPMENT OF MOLECULAR MARKERS AND PARTICULARLY
THE COMBINED USE OF MATERNALLY
INHERITED CYTOPLASMIC MARKERS WITH BIPARENTALLY
INHERITED NUCLEAR MARKERS HAS ALLOWED THE DETECTION OF MULTIPLE AND RECURRENT
ORIGINS OF POLYPLOID SPECIES 3OLTIS AND 3OLTIS !S MULTIPLE
SEQUENCE DATAS
ETS ARE NOW AVAILABLE FOR PHYLOGENETIC ANALYSES IT IS POSSIBLE TO DETECT RECURRENT
AND BI
DIRECTIONAL RETICULATE EVOLUTION EVEN IN OLD POLYPLOID LINEAGES EG 'LYCINE
$OYLE ET AL WHERE THEY OTHERWISE WOULD HAVE BEEN UNDETECTED
4HE PROCESS OF RECURRENT POLYPLOID FORMATION WITH RETICULATE GENE mOW MAY IN
VOLVE VARIOUS PARENTAL GENOTYPES AND INCREASES THE LEVEL OF GENETIC DIVERSITY
AVAILABLE TO NEWLY FORMED SPECIES 3CHIERENBECK ET AL &OR INSTANCE AT
LEAST LINEAGES OF SEPARATE ORIGINS FOR THE ALLOTETRAPLOID 4RAGOPOGON MISCELLUS
AND FOR 4 MIRUS HAVE BEEN DOCUMENTED IN THE 0ALOUSE ACCORDING TO VARIOUS
MORPHOLOGICAL AND MOLECULAR ALLOZYMES CHLOROPLAST AND NUCLEAR $.! LINES OF
%VOLUTION AND PLANT INVASIONS
EVIDENCE REVIEWED IN 3OLTIS ET AL THESE RECURRENT ORIGINS INVOLVE EITHER
ONLY ONE DIPLOID SPECIES AS THE MATERNAL PARENT EG 4 PORRIFOLIUS FOR THE ALLO
TETRAPLOID 4 MIRUS OR ALTERNATIVELY BOTH THE PARENTAL SPECIES 4 PORRIFOLIUS AND
4 DUBIUS IN RECIPROCAL CROSSES E.g. FOR THE ALLOTETRAPLOID 4 MISCELLUS 4HIS HAS
RESULTED IN VARIOUS GENOTYPES AND DRAMATIC mORAL DIFFERENCES IN THE ALLOPOLYPLOID
POPULATIONS THAT ARE PROGRESSIVELY REPLACING DIPLOIDS AS PREVALENT WEEDS 3OLTIS
ET AL 3IMILARLY TWO SEPARATE ORIGINS HAVE BEEN DOCUMENTED IN .ORTH 7ALES
AND 3COTLAND FOR THE RUDERAL ALLOHEXAPLOID 3ENECIO CAMBRENSIS THAT ORIGINATED IN
"RITAIN DURING THE PAST YEARS REVIEWED IN !BBOTT AND ,OWE /NE OF
THE MOST COMPLEX EXAMPLES OF MULTIPLE ALLOPOLYPLOID ORIGINS IS REPRESENTED BY
THE POLYPLOID AGAMIC COMPLEX !NTENNARIA ROSEA PUSSYTOES THAT HAS FORMED FROM
MULTIPLE CROSSES INVOLVING VARIOUS DIPLOID TAXA OCCURRING IN SPECIlC HABITATS OF THE
2OCKY -OUNTAINS "AYER 4HE POLYPLOID LINEAGES OF THIS hCOMPILOSPECIESv
ARE GAMETOPHYTIC APOMICTS THAT DISPLAY LARGER ECOLOGICAL AMPLITUDE THAN THEIR
DIPLOID PROGENITORS
.OT ALL SUCCESSFUL ALLOPOLYPLOIDS HAVE MULTIPLE ORIGINS )N CONTRAST TO 4RAGOPOGON
SPP 3ENCECIO CAMBRENSIS AND !NTENNARIA ROSEA THE INVASIVE SALTMARSH SPECIES
3PARTINA ANGLICA HAS UNDERGONE A SEVERE GENETIC BOTTLENECK AT THE TIME OF ITS
FORMATION IN THE "AY OF 3OUTHAMPTON 5+ 4HIS DODECAPLOID SPECIES FORMED
AFTER CHROMOSOME DOUBLING OF THE lRST GENERATION HYBRID 3PARTINA X TOWNSENDII
THAT HAS RESULTED FROM HYBRIDIZATION BETWEEN THE INTRODUCED %AST
!MERICAN HEXA
PLOID 3PARTINA ALTERNImORA AND THE NATIVE HEXAPLOID 3PARTINA MARITIMA 2AYBOULD
ET AL "OTH PARENTAL SPECIES LACK GENETIC DIVERSITY IN THE HYBRIDIZATION SITE
A LIMITED NUMBER OF 3 ALTERNImORA GENOTYPES HAVE BEEN INTRODUCED IN 7ESTERN
%UROPE "AUMEL ET AL WHEREAS A STRIKING LACK OF MOLECULAR VARIATION
IS ENCOUNTERED IN POPULATIONS OF THE NATIVE 3 MARITIMA 9ANNIC ET AL
%UROPEAN POPULATIONS OF 3 ANGLICA ARE MOSTLY COMPOSED OF ONE MAJOR MULTILO
CUS GENOTYPE THAT HAS FORMED IN 3OUTHAMPTON AND THAT IS IDENTICAL TO THE lRST
GENERATION HYBRID 3 X TOWNSENDII "AUMEL ET AL A #HLOROPLAST $.!
ANALYSIS HAS REVEALED ALL POPULATIONS OF 3 ANGLICA IN WESTERN %UROPE DISPLAY IDEN
TICAL PLASTOME TO 3 ALTERNImORA WHICH IS THEN CONSIDERED AS THE MATERNAL GENOME
DONOR &ERRIS ET AL "AUMEL ET AL 3PARTINA ANGLICA HAS RAPIDLY INVADED
THE "RITISH SALTMARSHES SINCE ITS FORMATION 4HOMPSON AND IT HAS BEEN
NATURALLY OR DELIBERATELY INTRODUCED IN VARIOUS CONTINENTS SUCH AS #HINA OR
!USTRALIA WHERE IT IS NOW CONSIDERED AS A SERIOUS THREAT TO NATIVE mORA AND FAUNA
!NOUCHE ET AL A AND REFERENCES THEREIN !LTHOUGH IT HAS LIMITED INTER
INDI
VIDUAL GENETIC DIVERSITY 3 ANGLICA CONTAINS TWO WELL
DIFFERENTIATED HOMOELOGOUS
GENOMES INHERITED FROM ITS HEXAPLOID PARENTS WHICH PROVIDES lXED HETEROZYGOS
ITY AT HOMOEOLOGOUS LOCI "AUMEL ET AL B !NOUCHE ET AL B 3PARTINA
IS AN IDEAL SYSTEM IN WHICH TO EXPLORE THE GENETIC AND GENOMIC CONSEQUENCES
OF HYBRIDIZATION AND GENE DUPLICATION IN SUCCESSFUL INVASIVE SPECIES ALTHOUGH
PREVIOUS ATTEMPTS OF RE
SYNTHESIZING EXPERIMENTALLY THE ALLOPOLYPLOID HAVE
FAILED TIME SINCE SPECIES FORMATION AND THE PARENTAL SPECIES ARE KNOWN IT IS POS
SIBLE TO DIFFERENTIATE BETWEEN THE EFFECTS OF HYBRIDIZATION IN 3 X TOWNSENDII AND
+! 3CHIERENBECK AND -, !NOUCHE
GENOME DUPLICATION IN 3 ANGLICA POPULATIONS MOREOVER TWO NATURAL REPLICATES
OF HYBRIDIZATION EVENTS BETWEEN 3 MARTIMA AND 3 ALTERNImORA ARE AVAILABLE IN
3 X TOWSENDII AND 3 X NEYRAUTII ANOTHER HYBRID THAT HAS FORMED AT THE SAME PERIOD
IN SOUTHEWEST &RANCE WITH NO GENOME DOUBLING "AUMEL ET AL
4HE IMMEDIATE CONSEQUENCE OF POLYPLOIDY IS A GREATER INTRA INDIVIDUAL GENETIC
DIVERSITY AND HETEROZYGOSITY AT DUPLICATED LOCI THAN IN DIPLOIDS THAT RESULTS IN
INCREASED BIOCHEMICAL DIVERSITY 2OOSE AND 'OTTLIEB AND CONFERS A GREATER
TOLERANCE TO ENVIRONMENT VARIATION AND MAY PROMOTE SUCCESSFUL COLONIZATION
"ROWN AND -ARSHALL 'ENE DUPLICATIONS RESULTING FROM POLYPLOIDY ARE ALSO
BELIEVED TO HAVE A BUFFERING EFFECT AGAINST DELETERIOUS MUTATIONS /HNO
,YNCH AND #ONERY ,AWTON 2AUH !DDITIONALLY POLYPLOIDS OF HYBRID
ORIGIN ALLOPOLYPLOIDS MAY BENElT FROM HIGHER lTNESS DUE TO HETEROSIS 3IMILARLY
TRAITS WHICH RESULT IN THE hGIGASv EFFECT OR A GENERAL INCREASE IN MORPHOLOGICAL
CHARACTERISTICS DE 6RIES ARE ASSOCIATED WITH BOTH POLYPLOIDY AND INVA
SIVE PLANT SPECIES "AKER 2IESEBERG ET AL (YPERICUM PERFORATUM
3T *OHNS WORT A TETRAPLOID THOUGHT TO BE OF ALLOPOLYPLOID ORIGIN HAS HIGH REPRO
DUCTIVE PLASTICITY -ATZKE ET AL BUT REPRODUCES MOST COMMONLY VIA FACULTA
TIVE APOMIXIS &IELD COLLECTIONS OF ( PERFORATUM IN !USTRALIA FOUND A LACK OF WITHIN
POPULATION VARIATION -AYO AND ,ANGRIDGE BUT HIGH INTERPOPULATIONAL
GENETIC DIVERSITY (YPERICUM PERFORATUM HAS HIGH LEVELS OF HYPERICIN A CHEMICAL
THAT CAUSES PHOTOSENSITIZATION AND REDUCED HERBIVORY THAT ARE HYPOTHESIZED TO
BE A DIRECT RESULT OF ALLOPOLYPLOIDIZATION -AYO AND ,ANGRIDGE )NTRODUCED
INTO .ORTH !MERICAN IN THE S THE ALLOPOLYPLOID AND NEARLY MONOMORPHIC
3ETARIA FABERI GIANT FOXTAIL IS NOW PRESENT THROUGHOUT THE DISTURBED AREAS ON THE
CONTINENT 0OHL (AmIGER AND 3CHOLZ $EKKER NAMED
THE ALLOPOLYPLOIZATION OF 3 FABERI THE hWEED SPECIATION EVENTv AS THE POLYPLOID
HAS HIGHER lTNESS IN AGRICULTURAL SYSTEMS THAN ITS DIPLOID ANCESTORS /NE OF
THE WORLDS WORST WEEDS IS THE POLYPLOID 3ORGHUM HALEPENSE *OHNSON GRASS
A PRODUCT OF HYBRIDIZATION BETWEEN THE CULTIVAR 3 BICOLOR AND THE WILD 3 PROPIN
QUUM AND WHICH RESPONDS WELL TO CULTIVATION AND IS EXTREMELY PERNICIOUS TO DUE
THE RHIZOMATOUS GROWTH HABIT 0ATERSON
! GROWING BODY OF EVIDENCE CONTINUES TO ACCUMULATE IN REGARD TO THE DYNAMIC
AND PLASTIC NATURE OF POLYPLOID GENOMES THAT WOULD EXPLAIN THEIR EVOLUTIONARY
SUCCESS !LLOPOLYPLOID GENOMES ARE PARTICULARLY DYNAMIC AT BOTH THE STRUCTURAL
AND EXPRESSION LEVELS OVER THE LONG AND ALSO SHORT TERM EVOLUTIONARY TIME SCALE
REVIEWED IN 7ENDEL ,IU AND 7ENDEL /SBORN ET AL 3IGNIlCANT
ADVANCES IN REVEALING THE OCCURRENCE AND NATURE OF THE EARLY EVOLUTIONARY
CHANGES IN POLYPLOID GENOMES ARE POSSIBLE DUE TO EXPERIMENTALLY RESYNTHESIZED
ALLOPOLYPLOIDS INVOLVING WELL KNOWN MODEL SYSTEMS SUCH AS "RASSICA OILSEED
RAPE !RABIDOPSIS 'OSSYPIUM COTTON OR 4RITICUM !EGILOPS WHEAT 4HESE MODEL
SYSTEMS ALLOW THE EXPLORATION OF ALLOPOLYPLOID MATERIAL OF KNOWN ORIGIN WITH THE
COMPARISON OF THEIR ACTUAL PARENTAL GENOTYPES A CONDITION RARELY MET FOR MOST NAT
URAL ALLOPOLYPLOIDS 2APID AND BIASED STRUCTURAL CHANGES HAVE BEEN ENCOUNTERED
IN THE lRST GENERATIONS FOLLOWING POLYPLOIDIZATION IN "RASSICA 3ONG ET AL
%VOLUTION AND PLANT INVASIONS
AND WHEAT &ELDMAN ET AL ,IU ET AL A B /ZKAN ET AL
(OWEVER ,IU ET AL DID NOT lND CONSISTENT STRUCTURAL CHANGES IN NEWLY
SYNTHESIZED ALLOPOLYPLOID 'OSSYPIUM 6ARIOUS EVOLUTIONARY MECHANISMS APPEAR
TO AFFECT ALLOPOLYPLOID GENOMES OVER A LONGER TERM AND INCLUDE THE INDEPENDENT
EVOLUTION OF DUPLICATED GENES IN ALLOTETRAPLOID COTTON THAT FORMED ONE TO TWO
MILLION YEARS AGO #RONN ET AL 3ENCHINA ET AL INTERACTION BETWEEN
THE HOMOELOGOUS SUBGENOMES FOR REPETITIVE SEQUENCES VIA CONCERTED EVOLUTION
e.g., 'OSSYPIUM 7ENDEL ET AL .ICOTIANA TOBACCO 6OLKOV OR SPREAD
OF TRANSPOSABLE ELEMENTS :HAO ET AL 4HE FATE OF DUPLICATED HOMOEOLOGOUS
GENES HAS BEEN PARTICULARLY WELL
INVESTIGATED IN THE 'OSSYPIUM SYSTEM BY *ONATHAN
7ENDEL AND HIS COLLEAGUES EG !$( GENES 3MALL AND 7ENDEL
-9" GENES #EDRONI ET AL AND REVEALS VARIOUS EVOLUTIONARY PATTERNS
INCLUDING COPY NUMBER LABILITY PSEUDOGENIZATION GENE ELIMINATION OR ACCELERAT
ED RATE OF NUCLEOTIDE SUBSTITUTION EG FOR !$(
# GENES (OWEVER A RECENT ANALY
SIS OF NUCLEAR GENES 3ENCHINA ET AL INDICATED THAT POLYPLOIDY LED TO AN
OVERALL MODEST ENHANCEMENT IN RATES OF NUCLEOTIDE SUBSTITUTION IN 'OSSYPIUM
7HEN COMPARED TO RESYNTHESIZED ALLOPOLYPLOIDS NATURALLY NASCENT ALLOPOLY
PLOIDS DISPLAY DIFFERENT PATTERNS OF GENOME EVOLUTION VARIOUS LEVELS OF CONCERTED
EVOLUTION SEEM TO HAVE AFFECTED R$.! SEQUENCES OF THE YOUNG ALLOPOLYPLOID
POPULATIONS IN 4RAGOPOGON 3OLTIS ET AL WHEREAS NO HOMOGENIZATION OF
THE PARENTAL SEQUENCES IS OBSERVED IN 3PARTINA ANGLICA !NOUCHE ET AL A
)N THE LATTER SYSTEM NO MAJOR CHANGE OF THE PARENTAL GENOMES ARE OBSERVED FOR
VARIOUS MULTILOCUS MARKERS )332S 2!0$S !&,0S ALTHOUGH SOME PREFERENTIAL
LOSS OF MATERNAL FROM 3 ALTERNImORA !&,0 FRAGMENTS ARE OBSERVED IN 3 x TOWNSEN
DII AND 3 ANGLICA !NOUCHE ET AL B 3ALMON ! ET AL UNPUBLISHED ! TRANS
POSON DISPLAY ANALYSIS INDICATES NO BURST OF RETRO
ELEMENT ACTIVATION IN 3 ANGLICA
"AUMEL ET AL A AND SUGGESTS THAT DIFFERENT BIOLOGICAL SYSTEMS RESPOND VARI
OUSLY TO POLYPLOIDY ,IU ET AL )N SPITE OF THE STRUCTURAL GENOMIC STASIS
ENCOUNTERED FOR MOST OF THE MARKERS INVESTIGATED TO DATE 3PARTINA ANGLICA POPULA
TIONS EXHIBIT CONSISTENT MORPHOLOGICAL PLASTICITY 4HOMPSON AND SUGGEST
A PROBABLE FUNCTIONAL PLASTICITY IN THE EXPRESSION OF THE DUPLICATED LOCI !NOUCHE
ET AL A
2ECENT STUDIES HAVE POINTED OUT THAT THERE IS MODULATED EXPRESSION OF DUPLI
CATED LOCI IN POLYPLOIDS #OMAI 3HAKED ET AL +ASHKUSH ET AL
+ASHKUSH ET AL (E ET AL !DAMS ET AL 4HESE EXPRESSION CHANG
ES MAY INVOLVE VARIOUS MECHANISMS INCLUDING INCREASED VARIATION IN DOSAGE
REGULATED GENE EXPRESSION ALTERED REGULATORY NETWORKS AND GENETIC OR EPIGENETIC
CHANGES 2IDDLE AND "IRCHLER /SBORN ET AL )N WHEAT POLYPLOIDY
WAS ACCOMPANIED BY TRANSCRIPTIONAL ACTIVATION OF RETROELEMENTS THAT LED TO NOVEL
EXPRESSION PATTERNS +ASHKUSH ET AL ,EVY AND &ELDMAN %XPRESSION
CHANGES MAY HAVE PROFOUND IMPACT ON lTNESS WHEN THEY RESULT IN VARIABLE
PHENOTYPES 'ENE SILENCING RESULTED IN PHENOTYPIC VARIATION AND INSTABILITY IN
EXPERIMENTALLY RESYNTHESIZED !RABIDOPSIS ALLOTETRAPLOIDS THAT DISPLAYED CONSIDER
ABLE VARIATION IN MORPHOLOGY mOWERING TIME AND FERTILITY #OMAI ET AL
+! 3CHIERENBECK AND -, !NOUCHE
.OVEL mOWERING TIME VARIATION WAS ALSO OBSERVED IN RESYNTHESIZED ALLOPOLYPLOID
"RASSICA NAPUS 3CHRANZ AND /SBORN %XPRESSION OF HOMOEOLOGOUS GENE
PAIRS WAS ANALYSED BY !DAMS ET AL IN NATURAL TO
MYR OLD AND SYN
THETIC ALLOTETRAPLOID 'OSSYPIUM 4HE DUPLICATED GENES SHOWED UNEQUAL LEVELS OF
EXPRESSION AND ORGAN
SPECIlC RECIPROCAL SILENCING SUGGESTING A PARTITIONING OF
THE ANCESTRAL FUNCTIONS AS BOTH IMMEDIATE IN SYNTHETIC POLYPLOIDS AND LONG
TERM
IN NATURAL POLYPLOIDS RESPONSES TO POLYPLOIDIZATION )N RECENTLY FORMED NATURAL
ALLOPOLYPLOIDS SUCH EXPRESSION CHANGES APPEAR TO TAKE PLACE ALSO AS REVEALED BY
C$.! !&,0 INVESTIGATIONS IN 4RAGOPOGON WHERE ABOUT OF THE GENES EXAMINED
IN THE ALLOPOLYPLOIDS HAVE BEEN SILENCED AND AN ADDITIONAL EXHIBIT NOVEL GENE
EXPRESSION RELATIVE TO THEIR DIPLOID PARENTS 3OLTIS ET AL
%PIGENETIC CHANGES ASSOCIATED WITH INVASIVE ABILITY
%PIGENETICS REFERS TO HERITABLE CHANGES IN PHENOTYPE THAT DO NOT RESULT FROM
CHANGES IN GENE SEQUENCE BUT RATHER FROM REGULATORY MECHANISMS OF GENE EXPRES
SION 7OLFFE AND -ATZKE 4HESE MECHANISMS ARE KNOWN TO BE INVOLVED
IN GROWTH AND DEVELOPMENT &INNEGAN ET AL AND CAN RESULT IN VARIOUS
MORPHOLOGICAL CHANGES INCLUDING mOWER STRUCTURE EG #UBAS ET AL #OMAI
ET AL 3UCH MECHANISMS HAVE IMPORTANT EVOLUTIONARY CONSEQUENCES
BECAUSE THEY INCREASE PHENOTYPIC PLASTICITY WHICH IN TURN BUFFERS ENVIRONMEN
TAL PRESSURES ON GENOTYPES 4HIS MAY BE ASTONISHING TO OUR NAVE PERSPECTIVE
OF GENOMIC INTERACTIONS BUT EPIGENETIC PROCESSES ARE PROVING TO BE PREDICTABLY
ASSOCIATED WITH THE STRUCTURAL LIMITATIONS OF GENOMES %PIGENETICS IS CONSEQUENTLY
BECOMING A VERY ACTIVE lELD OF RESEARCH IN EVOLUTIONARY GENOMICS REVIEWED IN
&INNEGEAN ET AL #OMAI ,IU AND 7ENDEL
-ECHANISTICALLY EPIGENETIC CHANGES RESULT FROM VARIOUS INTERACTING PROCESSES
WHICH INCLUDE CYTOSINE METHYLATION OF $.! -ARTIENSSEN AND #OLOT HIS
TONE DEACETYLATION 4IAN AND #HEN AND SHORT 2.!S -ETTE ET AL
THAT MODULATE GENE SILENCING %PIGENETIC ALTERATIONS ARE KNOWN TO BE TRIGGERED BY
ENVIRONMENTAL STRESS &INNEGAN AND IN SOME CASES ARE VIEWED AS GENOME
DEFENSE MECHANISMS 9ODER ET AL -ATZKE ET AL )N INTROGRESSED
HYBRID RICE PLANTS ,IU AND 7ENDEL OBSERVED RETROTRANSPOSON ACTIVATION
THAT WAS RAPIDLY REPRESSED BY CYTOSINE METHYLATION
4HE REUNION OF TWO DIVERGENT GENOMES IN THE SAME NUCLEUS IN HYBRID AND ALLO
POLYPLOID SPECIES MAY BE CONSIDERED AS A GENOMIC STRESS THAT GENERATES EPIGENETIC
CHANGES ALTERING GENE EXPRESSION AND PHENOTYPES #OMAI ET AL )N SYN
THETIC !RABIDOPSIS ALLOTRETRAPLOIDS #OMAI ET AL OBSERVED THAT ABOUT OF
THE GENES WERE SILENCED COMPARED TO THEIR PARENTS 4HE SILENCED GENES WERE BOTH
NORMAL GENES OR GENES RELATED TO TRANSPOSONS 4HESE CHANGES WERE FURTHER FOUND
TO BE RELATED TO METHYLATION MODIlCATIONS THAT WERE ASSOCIATED TO PHENOTYPIC
INSTABILITY -ADLUNG ET AL 3IMILAR LEVELS OF SILENCING RELATED TO CYTOSINE
METHYLATION WERE ALSO ENCOUNTERED IN THE CORRESPONDING NATURAL ALLOTETRAPLOID
!RABIDOPSIS SUECICA ,EE AND #HEN )N EXPERIMENTALLY RE
SYNTHESIZED ALLO
%VOLUTION AND PLANT INVASIONS
POLYPLOID WHEAT OF THE LOCI INVESTIGATED USING -ETHYLATION 3ENTITIVE !&,0
-3!0 WERE FOUND METHYLATED 3HAKED ET AL AND TRANSCRIPTIONAL ACTI
VATION OF RETROTRANSPOSONS WAS SHOWN TO ALTER THE EXPRESSION OF ADJACENT GENES
+ASHKUSH ET AL ,EVY AND &ELDMAN !LTHOUGH NO CHANGES IN METH
YLATION PATTERNS WERE OBSERVED IN NEWLY SYNTHESIZED 'OSSYPIUM ALLOPOLYPLOIDS
,IU ET AL THE ORGAN
SPECIlC AND RECIPROCAL GENE SILENCING FOUND BY !DAMS
ET AL IS INTERPRETED AS RESULTING FROM EPIGENETIC REGULATION THROUGH MECHA
NISMS THAT HAVE YET TO BE ELUCIDATED !DAMS AND 7ENDEL
3PARTINA ANGLICA IS TO OUR KNOWLEDGE THE lRST INVASIVE SPECIES THAT HAS BEEN
INVESTIGATED IN THE CONTEXT OF EPIGENETIC GENE EXPRESSION !NOUCHE ET AL B
3ALMON ET AL HAVE ATTEMPTED TO DIFFERENTIATE BETWEEN THE GENOMIC CON
SEQUENCES THAT RESULT FROM HYBRIDIZATION AND THOSE THAT RESULT FROM GENOME
DUPLICATION 4HIS WAS MADE POSSIBLE BY THE COMPARISON OF THE ALLOPOLYPLOID TO
THE NATURAL & HYBRID SPECIES 3 X TOWNSENDII THE PROGENITOR OF 3 ANGLICA AND
3 X NEYRAUTII THE OTHER HYBRID THAT HAS FORMED INDEPENDENTLY IN SOUTHWEST
&RANCE "AUMEL ET AL -3!0 DATA ANALYSIS REVEALED CONSISTENT METHYLA
TION CHANGES THAT CONTRAST WITH THE STRUCTURAL ADDITIVITY OF THE PARENTAL GENOMES
MENTIONED ABOVE -OST METHYLATION CHANGES WERE FOUND IN BOTH 3 X TOWNSENDII
AND 3 X NEYRAUTII WHICH INDICATES THE REPRODUCIBILITY OF THE CHANGES IN THE TWO
DIFFERENT HYBRIDISATION EVENTS 4HE METHYLATION ALTERATIONS FOUND IN 3 ANGLICA WERE
ALREADY PRESENT OR INITIATED IN 3 X TOWNENDII SUGGESTING THAT EPIGENETIC CHANGES
WERE TRIGGERED BY HYBRIDIZATION RATHER THAN BY GENOME DUPLICATION 4HE EXTENT OF
SUCH CHANGES AND THEIR VARIABILITY WHEN PLANTS ARE FACING DIFFERENT ENVIRONMENTAL
CONDITIONS NEED TO BE EXPLORED AT THE POPULATION LEVEL AND THE SEQUENCES THAT ARE
EPIGENETICALLY AFFECTED HAVE TO BE IDENTIlED
)N THE CONTEXT OF RAPID EXPANSION OF INVASIVE SPECIES THAT EXPLORE NEW HABI
TATS EPIGENETIC PROCESSES ARE OF MAJOR INTEREST AS THEY DElNITELY INmUENCE lTNESS
"ECAUSE EPIGENETIC CASES OF GENE SILENCING WILL NOT BE REmECTED IN SEQUENCE DATA
THE ASSESSMENT OF GENE EXPRESSION IS KEY TO THE UNDERSTANDING OF GENE FUNCTION
AND ITS IMPORTANCE IN SPECIES ADAPTATION AND INVASIVENESS -OLECULAR EVOLUTION
ISTS ARE BECOMING MORE AWARE OF THE NECESSITY TO APPROACH ADAPTIVE PROCESSES AT
BOTH GENE AND GENOME LEVELS
3%,%#4)/. $%6%,/0-%.4!, 2%30/.3% !.$ ).6!3)/.3
'ENETIC CHANGE AND SUBSEQUENT ALTERATIONS IN DEVELOPMENTAL RESPONSE HAVE BEEN
MINIMALLY ADDRESSED IN INVASIVE PLANT SPECIES 3IMILARLY ALTHOUGH THE EVALUA
TION OF DEVELOPMENTAL QUANTITATIVE TRAITS HAS BEEN CONSIDERED THEORETICALLY THERE
IS LITTLE EMPIRICAL EVIDENCE FOR OR AGAINST THEIR IMPORTANCE IN PLANT INVASIONS
4HE GENETIC AND ECOLOGICAL INTERACTIONS THAT RESULT FROM CHANGES IN DEVELOPMENTAL
GENES OR FROM GENES OF MAJOR EFFECT COULD HAVE PROFOUNDLY COMPLEX REPERCUS
SIONS &OR EXAMPLE THE RELEASE FROM ENEMIES MAY ALLOW FOR DECREASED ALLOCATION
TO PROTECTIVE MECHANISMS AND A CONCOMITANT INCREASE IN lTNESS THAT RESULTS FROM
+! 3CHIERENBECK AND -, !NOUCHE
DEVELOPMENTAL RESPONSES 7E KNOW LITTLE ABOUT THE ECOLOGICAL IMPACTS OF DEVELOP
MENTAL CHANGES AND INTERACTIONS IN PREDATOR
PREY RELATIONSHIPS .IJHOUT
!CTION FROM JUST A FEW PARTICULARLY IMPORTANT GENES SUCH AS THESE MAY BE KEY TO
THE ABILITY TO INVADE 0ATERSON ET AL
!LL ADAPTATIONS IMPORTANT TO INVASIVENESS e.g., HIGH SEED PRODUCTION BREED
ING SYSTEM CHANGES AND VEGETATIVE PROPAGATION ARE AMENABLE FOR STUDY IN A
DEVELOPMENTAL GENETIC AND ECOLOGICAL CONTEXT 0URAGGANAN 'ILBERT AND
"OLKER 'ENES SUCH AS TEOSINTE BRANCHED IN MAIZE ILLUSTRATE THE IMPORTANCE
OF DEVELOPMENTAL GENES IN PLANT ESTABLISHMENT AND SPREAD 1 #RONK PERS COMM
$OEBLEY ET AL 4EOSINTE BRANCHED WHICH CONTROLS TILLERING IN MAIZE IS
RESPONSIBLE FOR THE DIFFERENCE BETWEEN TEOSINTE THAT IS PERSISTENT IN THE ENVIRON
MENT AND VARIETIES THAT ARE NOT ECOLOGICALLY COMPETITIVE $OEBLEY ET AL
6ARIATION IN REGULATORY GENES MAY BE KEY TO THE PROVISION OF THE GENETIC VARIATION
NECESSARY TO ALTER PHENOLOGICAL DEVELOPMENTAL PATHWAYS mORAL VARIATION SECOND
ARY PLANT COMPOUND PRODUCTION AND CLONAL GROWTH
,IFE HISTORY STRATEGIES OFTEN EMPHASIZED AS KEY TO UNDERSTANDING INVASIVE
ABILITY HAVE BEEN ILLUSTRATED AS PHENOTYPICALLY ENVIRONMENTALLY DEPENDENT
0IGLIUCCI 4HE DEVELOPMENTAL RESPONSE OF REACTION NORMS IS BASED ON MEAN
PHENOTYPIC TRAIT VALUES THUS THE VARIATION MUST BE EXPLAINED BY ADDITIVE GENETIC
VARIANCE OR GENES FOR hADAPTIVE PLASTICITYv 3ULTAN ! COMPARISON BETWEEN
0OLYGONUM SPECIES WITH BROAD ECOLOGICAL TOLERANCE TO THOSE LIMITED BY ENVIRON
MENTAL FACTORS DEMONSTRATE THAT DEVELOPMENTAL TIMING OF PLASTIC RESPONSES IS
IMPORTANT TO ENVIRONMENTAL BREADTH 3ULTAN 'ENE EXPRESSION IS ALTERED
BY BOTH INTERNAL AND MORE IMPORTANTLY WITHIN THE INVASIVE SPECIES CONTEXT EXTER
NAL CUES 3CHLICHTING /F PARTICULAR IMPORTANCE TO INVASIVE ABILITY MAY BE
THE DEVELOPMENTAL PLASTICITY OF GENDER AND BREEDING SYSTEM mEXIBILITY WHICH WILL
IMPACT SEX RATIOS IN PROPAGULES RESPONDING TO NEW RANGE ENVIRONMENTAL FACTORS
$ELPH !S 3CHAAL ET AL SUGGEST GENE GENEALOGIES WILL BE IMPOR
TANT TO TEST HYPOTHESES FOR ADAPTATION AND WE FURTHER SUGGEST THAT THIS SHOULD
INCLUDE DEVELOPMENTAL TRAITS THAT MAY BE CLOSELY RELATED TO SELECTION AND ADAPTIVE
DIVERGENCE /UR EXPECTATION IS THAT THE PHENOMENON OF RAPID DIVERGENCE OF DEVEL
OPMENTAL TRAITS WILL BE FOUND IN OTHER EXAMPLES OF ISLAND RADIATION OR IN REGIONS
SIMILARLY SUSCEPTIBLE TO INVASION )N CASES OF GENE DUPLICATION AND POLYPLOIDY
THE RATE OF THIS PROCESS MAY BE INCREASED BECAUSE OF THE hBACKUPvGENETIC EXPRES
SION AND COPIES WITH WHICH SELECTIVE PROCESSES CAN hTINKERv
2%3%!2#( .%%$3 ). 4(% !2%! /& 4(% '%.%4)#3 !.$
%6/,54)/. /& ).6!3)6% 30%#)%3
4HE DISTRIBUTION OF GENETIC VARIATION IN POPULATIONS OF INVASIVE SPECIES AND
THE RELATIVE INVASIVENESS OF DIFFERENT GENOTYPES REMAINS ELLUSIVE WITH FEW DElNITIVE
STUDIES OUTSIDE OF AGRICULTURAL BUT SEE 'ASKIN AND 3CHAAL AND 3ALTONSTALL
/F PARTICULAR NEED IN UNDERSTANDING THE DYNAMICS OF ADAPTATION AND
%VOLUTION AND PLANT INVASIONS
SPREAD IS AN ASSESSMENT OF THE MOLECULAR POPULATION GENETICS OF REGULATORY LOCI
0URAGGANAN
3OIL SEEDBANK ANALYSIS AS A SOURCE OF NEW GENETIC VARIATION HAS NOT BEEN
ADEQUATELY ADDRESSED IN PLANT INVASIONS 3EED DORMANCY IS AN ADAPTIVE TRAIT
AND ALTHOUGH DORMANCY PERIODS ARE WELL
DESCRIBED FOR MANY AGRICULTURAL WEEDS
"ASKIN AND "ASKIN $EKKER VIRUTALLY NOTHING IS KNOWN ABOUT
THE IMPORTANCE OF THIS TRAIT FOR THE MAINTAINANCE OF GENETIC DIVERSITY IN INVASIVE
PLANT SPECIES 3EED VIABILITY FOR #YTISUS SCOPARIUS CAN BE AS LONG AS YEARS AND
A TWO
OR THREE
YEAR OLD PLANT CAN PRODUCE UP TO SEEDSYEAR 0ARKER
ET AL 4HUS GENE mOW FROM SEED BANKS MAY BE CONSIDERABLE #HROMOLAENA
ODORATA TRIFlD WEED A SHRUB NATIVE TO THE NEOTROPICS AND INVASIVE THROUGHOUT
THE 0ACIlC )SLANDS IS QUITE AGGRESSIVE AND EVEN THE OCCURRENCE OF AN OCCASSIONAL
SEEDLING SIX YEARS PAST AN IRRADICATION EFFORT MAY BE A POTENTIALLY IMPORTANT SOURCE
OF GENETIC VARIATION 7ATERHOUSE AND :EIMER
1UANTITATIVE TRAIT LOCI ANALYSES HAVE PROVEN TO BE VERY USEFUL IN AGRICUL
TURAL GENETICS AND THERE ARE NUMEROUS EXAMPLES 0ATERSON )N 3ORGHUM
HALEPENSE VARIATION IN THE NUMBER OF RHIZOMES HAS BEEN ASSOCIATED WITH THREE
14,S AND THE NUMBER OF VEGETATIVE BUDS WITH AN ADDITIONAL 14, 0ATERSON ET AL
4HE POTENTIAL IMPORTANCE OF THIS TYPE OF WORK CAN NOT BE OVERSTATED IT HAS
BEEN WELL ESTABLISHED THAT CLONAL GROWTH AND THE ABILITY TO FORM MORE VEGETATIVE
BUDS CAN BE IMPORTANT TO INVASIVE ABILITY 0ATERSON #ONTINUED WORK WITH
AGRICULTURE SPECIES WILL AID THE STUDY OF THOSE GENES THAT MAY BE IMPORTANT IN
WEEDINESS SUCH AS mORAL OR FRUIT REGULATION AND VEGETATIVE GROWTH &OR EXAMPLE
THE VEGETATIVE GROWTH TRAITS FOUND IN 3 HALEPENSE CORRESPOND WITH THE SAME 14,S
IN RICE WHEAT CORN AND LIKELY OTHER BUT INVASIVE GRASSES !S SO ASTUTELY NOTED
BY 0ATERSON ET AL 14, ANALYSIS CAN LEAD TO THE CLONING OF CANDIDATE GENES
IMPORTANT IN VEGETATIVE GROWTH /NCE THESE GENES ARE IDENTIlED IT MAY HELP IN
THE SPECIlC ERADICATION OF SOME INVADERS THAT HAVE LARGE AMOUNTS OF VEGETATIVE
PROPAGATION THROUGH GENETIC OR CHEMICAL METHODS
)MPORTANT CONTRIBUTIONS OF THE QUANTITATIVE GENETICS IN UNDERSTANDING INVASIVE
SPECIES EVOLUTION WILL RESULT FROM STUDIES WHICH CONSIDER THE ADAPTIVE RESPONSE
OR ADDITIVE GENETIC VARIANCE OF A PARTICULAR TRAIT ,YNCH AND 7ALSH
-LLER
3CHRER AND 3TEINGER 3UCH STUDIES WILL BE FURTHER STRENGTHENED BY
THE IDENTIlCATION OF 14,S ASSOCIATED WITH THESE TRAITS AND THEIR EVENTUAL MOLECU
LAR CHARACTERIZATION 4HE INTEGRATION OF QUANTITATIVE GENETICS AND THE MOLECULAR
BASIS OF THESE TRAITS WILL EVENTUALLY ALLOW AN ASSESSMENT OF THE LARGE
SCALE ECOLOGI
CAL EFFECTS OF GENETIC TRAITS
!S PREVIOUSLY DISCUSSED HOMEOLOGOUS GENES OR THOSE GENES DUPLICATED BY
POLYPLOID EVENTS MAY UNDERGO VARIOUS EVOLUTIONARY PROCESSES /F PARTICULAR
INTEREST ARE THOSE GENES THAT HAVE ADAPTIVE SIGNIlCANCE 3PECIlC QUESTIONS
THAT CAN BE ADDRESSED TO ASSESS THE DYNAMICS OF THE GENES IN POLYPLOID SYSTEMS
INCLUDE CHARACTERIZATION IN SEQUENCE AND FUNCTION BETWEEN THE DIFFERENT COPIES OF
THE GENES AND IN BOTH THE PARENTAL SPECIES AND POLYPLOID OFFSPRING CHARACTER
IZATION OF DIVERGENCE TIMES AMONG THE PARENTAL SPECIES AND POLYPLOID OFFSPRING
+! 3CHIERENBECK AND -, !NOUCHE
-OST IMPORTANTLY THE DETERMINATION OF THE EFFECTS OF POLYPLOIDY ON DUPLICATE GENE
EXPRESSION AND ADAPTIVE FUNCTION IS NOW POSSIBLE -UCH REMAINS TO BE LEARNED
ABOUT THE DYNAMICS AND SUBSEQUENT GENOTYPIC AND PHENOTYPIC CHANGES DUE TO
THE HYBRIDIZATION BETWEEN HISTORICALLY ALLOPATRIC TAXA HOMEOLOGOUS GENES AND
ALLOPOLYPLOIDIZATION
!LTHOUGH CURRENTLY LIMITED IN THEIR APPLICATION TO MODEL SYSTEMS OR CLOSELY
RELATED TAXA THE EXPLORATION OF THE USE OF NEW GENOMIC TOOLS e.g., MICROARRAYS
MAY PROVE USEFUL FOR THE INVESTIGATION OF GENOMIC AND ECOLOGICAL INTERACTIONS
*ACKSON ET AL
#/.#,53)/.3
!S EVOLUTIONARY BIOLOGISTS MANY OF US HAVE WORKED TOWARD THE DAY IN WHICH WE
CAN LINK GENE AND GENOMES TO THE LEVEL OF THE ECOSYSTEM 4HIS DAY HAS ARRIVED
AND FOR BETTER OR WORSE A FORUM THAT OFFERS ONE OF THE BEST OPPORTUNITIES FOR THE
INTEGRATION OF GENETICS AND ECOLOGY IN AN EVOLUTIONARY CONTEXT IS THE STUDY OF
INVASIVE SPECIES !LTHOUGH THE PARTICIPANTS OF THE hGENETICS OF COLONIZING SPECIESv
SYMPOSIUM MADE THIS SAME OBSERVATION IN ONLY NOW DO WE HAVE THE TOOLS
TO ADDRESS THE MYRIAD OF QUESTIONS ASSOCIATED WITH THE EVOLUTION OF INVASIVE ABIL
ITY 7E ARE WELL AWARE THAT IT IS PERHAPS MORE THAN A COINCIDENCE THAT MANY OF
THE EXAMPLES OF INVASIVE SPECIES WE HAVE CITED OFTEN SHARE THE PHENOMENA OF RAPID
EXPANSION INTO NEW RANGES POLYPLOIDY AND HYBRIDIZATION
4HE INTEGRATION OF MOLECULAR GENETICS ECOLOGY AND LARGE
SCALE lELD EXPERI
MENTATION IS TOO LARGE FOR ANY SINGLE INVESTIGATOR TO STUDY IN INVASIVE SPECIES
OR ANY OTHER CONTEXT )NCREASED KNOWLEDGE IN THE RESPECTIVE lELDS OF GENOMICS
PROTEOMICS CELL BIOLOGY POPULATION GENETICS PHYLOGENETICS AND ECOLOGY WILL
REQUIRE THE COOPERATION OF LABORATORIES WITH EXPERTISE IN THESE RESPECTIVE AREAS
0ERHAPS INVASIVE SPECIES GENETICS WILL BE THE lELD THAT BRINGS TOGETHER EVOLUTION
ISTS FROM VARIOUS DISCIPLINES FOR BOTH A BETTER UNDERSTANDING AND THE MANAGEMENT
OF CONTEMPORARY ENVIRONMENTS
!#+./7,%$'%-%.43
7E THANK THE .ATIONAL 3CIENCE &OUNDATION IN THE 53 THE #ENTRE .ATIONAL
DE LA 2ECHERCHE 3CIENTIlQUE AND THE #ENTRE !RMORICAIN DE 2ECHERCHES EN
%NVIRONNEMENT IN &RANCE FOR THEIR SUPPORT OF COLLABORATIVE RESEARCH ON GENOME
EVOLUTION OF INVASIVE PLANTS 7E ALSO ACKNOWLEDGE AND THANK THE CONTRIBUTIONS OF
OUR STUDENTS AND MANY COLLABORATORS TO OUR WORK PARTICULARLY .ORMAN %LLSTRAND
#URT $AEHLER AND ONE ANONYMOUS REVIEWER ARE THANKED FOR HELPFUL COMMENTS ON
THE MANUSCRIPT
%VOLUTION AND PLANT INVASIONS
2%&%2%.#%3
!BBOTT 2 * 0LANT INVASIONS INTERSPECIlC HYBRIDIZATION AND THE EVOLUTION OF NEW
PLANT TAXA 4RENDS IN %COLOGY AND %VOLUTION
!BBOTT 2 * * + *AMES 2 ) -ILNE AND ! # - 'ILLIES 0LANT INTRODUCTIONS HYBRID
IZATION AND GENE mOW 0HILOSOPHICAL 4RANSACTIONS OF THE 2OYAL 3OCIETY OF ,ONDON "
!BBOTT 2 7 AND ! * ,OWE /RIGINS ESTABLISHMENT AND EVOLUTION OF NEW POLYPLOID
SPECIES 3ENECIO CAMBRENSIS AND 3 EBORANENSIS IN THE "RITISH ISLES "IOLOGICAL *OURNAL OF
THE ,INNEAN 3OCIETY )N PRESS
!DAMS + , 2 #RONN 2 0ERCIlELD AND * & 7ENDEL 'ENES DUPLICATED BY POLYPLOIDY
SHOW UNEQUAL CONTRIBUTIONS TO THE TRANSCRIPTOME AND ORGAN
SPECIlC RECIPROCAL SILENC
ING 0ROCEEDINGS OF THE .ATIONAL !CADEMY OF 3CIENCES
!DAMS + , AND * & 7ENDEL %XPLORING THE GENOMIC MYSTERIES OF POLYPLOIDY IN COT
TON "IOLOGICAL *OURNAL OF THE ,INNEAN 3OCIETY )N PRESS
!NOUCHE - , ! "AUMEL AND ! 3ALMON A 3PARTINA ANGLICA A NATURAL MODEL SYSTEM
FOR STUDYING EARLY EVOLUTIONARY CHANGES THAT AFFECT ALLOPOLYPLOID GENOMES "IOLOGICAL
*OURNAL OF THE ,INNEAN 3OCIETY In press.
!NOUCHE - , ! "AUMEL ! 3ALMON AND ' 9ANNIC B (YBRIDIZATION POLYPLOIDY
AND SPECIATION IN 3PARTINA 0OACEAE .EW 0HYTOLOGIST
!LLENDORF & 7 AND , , ,UNDQUIST )NTRODUCTION POPULATION BIOLOGY EVOLUTION
AND CONTROL OF INVASIVE SPECIES #ONSERVATION "IOLOGY
!MSELLEM , * , .OYER 4 ,E "OURGEOIS AND - (OSSAERT
-C+EY #OMPARISON OF
GENETIC DIVERSITY OF THE INVASIVE WEED 2UBUS ALCEIFOLIUS 0OIR 2OSACEAE IN ITS NATIVE
RANGE AND IN AREAS OF INTRODUCTION USING !&,0 MARKERS -OLECULAR %COLOGY
!NDERSON % )NTROGRESSIVE HYBRIDIZATION .EW 9ORK 53! *OHN 7ILEY
!NDERSON % AND ' , 3TEBBINS (YBRIDIZATION AS AN EVOLUTIONARY STIMULUS
%VOLUTION
!NTTILA # + 2 ! +ING # &ERRIS $ 2 !YRES AND $ 2 3TRONG 2ECIPROCAL HYBRID
FORMATION OF 3PARTINA IN 3AN &RANCISCO "AY -OLECULAR %COLOGY
!NTILLA # + # # $AEHLER . % 2ANK AND $ 2 3TRONG 'REATER MALE lTNESS OF
A RARE INVADER 3PARTINA ALTERNImORA THREATENS A COMMON NATIVE 3PARTINA FOLIOSA WITH
HYBRIDIZATION !MERICAN *OURNAL OF "OTANY
!RNOLD - , .ATURAL HYBRIDIZATION AND EVOLUTION /XFORD 5+ /XFORD 5NIVERSITY
0RESS
!RNOLD - , 4RANSFER AND ORIGIN OF ADAPTATIONS THROUGH NATURAL HYBRIDIZATION
7ERE !NDERSON AND 3TEBBINS RIGHT 4HE 0LANT #ELL
!UGE ( " .EUFFER & %RLINGHAGEN 2 'RUPE AND 2 "RANDL $EMOGRAPHIC AND
RANDOM AMPLIlED POLYMORPHIC $.! ANALYSES REVEAL HIGH LEVELS OF GENETIC DIVERSITY IN A
CLONAL VIOLET -OLECULAR %COLOGY
!YRES $ 2 $ 'ARCIA
2OSSI ( ' $AVIS $ 2 3TRONG %XTENT AND DEGREE OF HYBRID
IZATION BETWEEN EXOTIC 3PARTINA ALTERNImORA AND NATIVE 3 FOLIOSA CORDGRASS 0OACEAE IN
#ALIFORNIA 53! DETERMINED BY RANDOM AMPLIlED POLYMORPHIC $.! 2!0$S -OLECULAR
%COLOGY
+! 3CHIERENBECK AND -, !NOUCHE
"AKER ( ' #HARACTERISTICS AND MODES OF ORIGINS OF WEEDS 0AGES
"AKER ( ' AND ' , 3TEBBINS EDITORS 4HE 'ENETICS OF #OLONIZING 3PECIES !CADEMIC 0RESS
.EW 9ORK
"AKER ( ' AND ' , 3TEBBINS 4HE 'ENETICS OF #OLONIZING 3PECIES !CADEMIC 0RESS
.EW 9ORK
"AKER ( ' 4HE EVOLUTION OF WEEDS !NNUAL 2EVIEW OF %COLOGY AND 3YSTEMATICS
"ARTLETT % 3 * .OVAK AND 2 . -ACK 'ENETIC VARIATION IN "ROMUS TECTORUM
0OACEAE DIFFERENTIATION IN THE EASTERN 5NITED 3TATES !MERICAN *OURNAL OF "OTANY
"ARRETT 3 # ( AND " * 2ICHARDSON 'ENETIC ATTRIBUTES OF INVADING SPECIES 0AGES
IN 'ROVES 2 ( AND "URDON * * EDITORS %COLOGY OF "IOLOGICAL )NVASIONS
#AMBRIDGE 5NIVERSITY 0RESS -ELBOURNE !USTRALIA
"ARRIER - " ' "ALDWIN 2 ( 2OBICHAUX AND - $ 0URUGGANAN )NTERSPECIlC
HYBRID ANCESTRY OF A PLANT ADAPTIVE RADIATION ALLOPOLYPLOIDY OF THE (AWAIIAN SILVER
SWORD ALLIANCE !STERACEAE INFERRED FROM mORAL HOMEOTIC GENE DUPLICATIONS -OLECULAR
"IOLOGY AND %VOLUTION
"ARRIER - 2 ( 2OBICHAUX AND - $ 0URUGGANAN !CCELERATED REGULATORY GENE
EVOLUTION IN AN ADAPTIVE RADIATION 0ROCEEDINGS OF THE .ATIONAL !CADEMY OF 3CIENCES
"ASKIN # # AND * - "ASKIN 3EEDS %COLOGY "IOGEOGRAPHY AND %VOLUTION OF
$ORMANCY AND 'ERMINATION !CADEMIC 0RESS .EW 9ORK
"AUMEL ! - , !NOUCHE AND * % ,EVASSEUR -OLECULAR INVESTIGATIONS IN POPULA
TIONS OF 3PARTINA ANGLICA # % (UBBARD 0OACEAE INVADING COASTAL "RITTANY &RANCE
-OLECULAR %COLOGY
"AUMEL ! - , !NOUCHE 2 +ALENDAR ! ( 3CHULMAN A 2ETROTRANSPOSONS AND
GENOMIC STABILITY IN POPULATIONS OF THE YOUNG ALLOPOLYPLOID SPECIES 3PARTINA ANGLICA
# % (UBBARD 0OACEAE -OLECULAR "IOLOGY AND %VOLUTION
"AUMEL ! - , !NOUCHE 2 * "AYER ! + !INOUCHE AND - 4 -ISSET B -OLECULAR
PHYLOGENY OF HYBRIDIZING SPECIES FROM THE GENUS 3PARTINA Schreb 0OACEAE -OLECULAR
0HYLOGENETICS AND %VOLUTION
"AUMEL ! - , !NOUCHE - 4 -ISSET * 0 'OURRET AND 2 * "AYER 'ENETIC
EVIDENCE FOR HYBRIDIZATION BETWEEN THE NATIVE 3PARTINA MARITIMA AND THE INTRODUCED
3PARTINA ALTERNImORA 0OACEAE IN 3OUTH
7EST &RANCE 3PARTINA x NEYRAUTII RE
EXAMINED
0LANT 3YSTEMATICS AND %VOLUTION
"AYER 2 * !NTENNARIA ROSEA !STERACEAE ! MODEL GROUP FOR THE STUDY OF THE EVOLU
TION OF POLYPLOID AGAMIC COMPLEXES /PERA "OTANICA
"AZZAZ & ,IFE HISTORY OF COLONIZING PLANTS SOME DEMOGRAPHIC GENETIC AND
PHYSIOLOGICAL FEATURES 0AGES
IN ( ! -OONEY AND * ! $RAKE EDITORS
%COLOGY OF BIOLOGICAL INVASIONS OF .ORTH !MERICA AND (AWAII 3PRINGER
6ERLAG
(EIDELBERG 'ERMANY
"IRCHLER * ! $ , !UGER AND . # 2IDDLE )N SEARCH OF A MOLECULAR BASIS OF
HETEROSIS 4HE 0LANT #ELL
%VOLUTION AND PLANT INVASIONS
"LANC ' + (OKAMP AND + ( 7OLFE ! RECENT POLYPLOIDY SUPERIMPOSED ON OLDER
LARGE
SCALE DUPLICATIONS IN THE !RABIDOPSIS GENOME 'ENOME 2ESEARCH
"ROWN ! ( $ AND $ 2 -ARSHALL %VOLUTIONARY CHANGES ACCOMPANYING COLONIZA
TION IN 0LANTS 0AGES
IN ' # % 3CUDDER AND * , 2EVEAL EDITORS #OLONIZATION
3UCCESSION AND 3TABILITY "LACKWELL 3CIENTIlC 0UBLICATIONS /XFORD
"URKE * - AND - , !RNOLD 'ENETICS AND THE lTNESS OF HYBRIDS !NNUAL 2EVIEW OF
'ENETICS
#EDRONI - % 2 # #RONN + , !DAMS 4 ! 7ILKINS AND * & 7ENDEL %VOLUTION
AND EXPRESSION OF -9" GENES IN DIPLOID AND POLYPLOID COTTON 0LANT -OLECULAR "IOLOGY
#HEN : * , #OMAI AND # 3 0IKAARD 'ENE DOSAGE AND STOCHASTIC EFFECTS DETER
MINE DIRECTION AND SEVERITY OF UNIPARENTAL R2.! GENE SILENCING NUCLEOLAR DOMINANCE
IN !RABIDOPSIS ALLOPOLYPLOIDS 0ROCEEDINGS OF THE .ATIONAL !CADEMY OF 3CIENCES
#LAUSS - * AND 4 -ITCHELL
/LDS 0OPULATION GENETICS OF TANDEM TRYPSIN INHIBI
TOR GENES IN Arabidopsis SPECIES WITH CONTRASTING ECOLOGY AND LIFE HISTORY -OLECULAR
%COLOGY
#OHEN - " - 4 *ACKSON " 2 ,U 3 2 -ORIN ! - -ORTIMER * , 0HAM AND
, * 7ADE 0REDICTING THE ENVIRONMENTAL IMPACT OF TRANSGENIC OUTCROSSING TO
WILD AND WEEDY RICES IN !SIA 0AGES
IN 3YMPOSIUM 0ROCEEDINGS .O
'ENE &LOW AND !GRICULTURE 2ELEVANCE FOR 4RANSGENIC #ROPS "#0#5NIVERSITY OF +EELE
3TAFFORDSHIRE
#OMAI , 'ENETIC AND EPIGENETIC INTERACTIONS IN ALLOPOLYPLOID PLANTS 0LANT -OLECULAR
"IOLOGY
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4HOMPSON * $ -ORPHOLOGICAL VARIATION AMONG NATURAL POPULATIONS OF 3PARTINA ANGLI
CA 0AGES
IN 3PARTINA ANGLICA A RESEARCH REVIEW !
* 'RAY AND 0 % - "ENHAM
EDITORS )NSTITUTE OF 4ERRESTRIAL %COLOGY .ATURAL %NVIRONMENT 2ESEARCH #OUNCIL
,ONDON
4HOMPSON * $ 4HE BIOLOGY OF AN INVASIVE PLANT 7HAT MAKES 3PARTINA ANGLICA SO
SUCCESSFUL "IOSCIENCE
4IAN , AND : & #HEN "LOCKING HISTONE DEACETYLATION IN Arabidopsis INDUCES PLEIO
TROPIC EFFECTS ON PLANT GENE REGULATION AND DEVELOPMENT 0ROCEEDINGS OF THE .ATIONAL
!CADEMY OF 3CIENCES 53!
5RBANSKA + - ( (URKA % ,ANDOLT " .EUmER AND + -UMMENHOFF (YBRIDIZATION
AND EVOLUTION IN #ARDAMINE "RASSICACEAE AT 5RNERBODEN #ENTRAL 3WTITZERLAND BIOSYS
TEMATIC AND MOLECULAR EVIDENCE 0LANT 3YSTEMATICS AND %VOLUTION
6OLKOV 2 ! "ORISJUK . 6 0ANCHUK ) ) 3CHWEIZER $ AND 6 (ELMLEBEN
%LIMINATION AND REARRANGEMENT OF PARENTAL R$.! IN THE ALLOTETRAPLOID .ICOTIANA TABA
CUM -OLECULAR "IOLOGY AND %VOLUTION
6ITOUSEK 0 "IOLOGICAL INVASIONS AND ECOSYSTEM PROPERTIES CAN SPECIES MAKE A DIFFER
ENCE 0AGES
IN %COLOGY OF "IOLOGICAL )NVASIONS IN .ORTH !MERICA AND (AWAII
( ! -OONEY AND * ! $RAKE EDITORS 3PRINGER
6ERLAG .EW 9ORK
7ANG 2
, * & 7ENDEL AND * ( $EKKER 7EEDY ADAPTATION IN 3ETARIA SPP
) )SOZYME ANALYSIS OF GENETIC DIVERSITY AND POPULATION GENETIC STRUCTURE IN 3ETARIA
VIRIDIS !MERICAN *OURNAL OF "OTANY
7ARWICK 3 ) !LLOZYME AND LIFE
HISTORY VARIATION IN lVE NORTHWARDLY COLONIZING
.ORTH !MERICAN WEED SPECIES 0LANT 3YSTEMATICS AND %VOLUTION
7ATERHOUSE " - AND / :EIMER @/N THE BRINK THE STATUS OF #HROMOLAENA ODORATA
IN .ORTHERN !USTRALIA 0AGES
IN 0ROCEEDINGS OF THE lFTH INTERNATION WORKSHOP
%VOLUTION AND PLANT INVASIONS
ON "IOLOGICAL #ONTRAL AND -ANAGEMENT OF #HORMOLAENA ODORATA $URBAN 3OUTH !FRICA
:ACHARIADES # 2 -UNIAPPAN AND , 7 3TRATHIE EDITORS
7EBER % AND " 3CHMID ,ATITUDINAL POPULATION DIFFERENTIATION IN TWO SPECIES OF
3OLIDAGO !STERACEAE INTRODUCED INTO %UROPE !MERICAN *OURNAL OF "OTANY
7ENDEL * & ! 3CHNABEL AND 4 3EELANAN "I DIRECTIONAL INTERLOCUS CONCERTED
EVOLUTION FOLLOWING ALLOPOLYPLOID SPECIATION IN COTTON 'OSSYPIUM 0ROCEEDINGS OF
THE .ATIONAL !CADEMY OF 3CIENCES OF THE 53!
7ENDEL * & 'ENOME EVOLUTION IN POLYPLOIDS 0LANT -OLECULAR "IOLOGY
7ENDEL * & AND * * $OYLE 0OLYPLOIDY AND EVOLUTION IN PLANTS )N $IVERSITY AND
%VOLUTION IN 0LANTS 2 (ENRY %D #!" 0UBLISHING /XON 5 +
7ETZEL $ + - * (ORAK $ : 3KINNER AND 0 ! +ULAKOW 4RANSFER OF HERBICIDE
RESISTANCE TRAITS FROM !MARANTHUS PALMERI TO !MARANTHUS RUDIS 7EED 3CIENCE
7ILLIAMS $ ' 0HYSIOLOGICAL ECOLOGY OF THE INVASIVE GRASS 0ENNISETUM SETACEUM ON
(AWAII 0H$ $ISSERTATION 7ASHINGTON 3TATE 5NIVERSITY
7OLFE + ( 9ESTERDAYS POLYPLOIDS AND THE MYSTERY OF DIPLOIDIZATION .ATURE 2EVIEWS
'ENETICS
7OLFFE ! 0 AND - ! -ATZKE %PIGENETICS REGULATION THROUGH EXPRESSION 3CIENCE
8U # 9 7 * :HANG # : &U AND " 2ONG 'ENETIC DIVERSITY OF ALLIGATOR WEED IN
#HINA BY 2!0$ ANALYSIS "IODIVERSITY AND #ONSERVATION
9ANNIC ' ! "AUMEL AND - , !NOUCHE 5NIFORMITY OF THE NUCLEAR AND CHLORO
PLAST GENOMES OF 3PARTINA MARITIMA 0OACEAE A SALT MARSHES SPECIES IN DECLINE ALONG THE
7ESTERN %UROPEAN #OAST (EREDITY )N PRESS
9E 7 ( * ,I ( , #AO AND 8 * 'E 'ENETIC UNIFORMITY OF !LTERNANTHERA PHILOXEROI
DES IN 3OUTH #HINA 7EED 2ESEARCH
9ODER * ! 7ALSH # 0 "ESTOR 4 ( #YTOSINE METHYLATION AND THE ECOLOGY OF
INTRAGENOMIC PARASITES 4RENDS IN 'ENETICS
:HAO 8 0 3I 9 (ANSON 2 % #RANE # 2 0RICE ( * 3TELLY $ - 7ENDEL * & AND
! ( 0ATERSON $ISPERSED REPETITIVE $.! HAS COLONIZED NEW GENOMES SINCE POLY
PLOID FORMATION IN COTTON 'ENOME 2ESEARCH
#HAPTER TEN
#ONTACT EXPERIENCE
ALIEN
NATIVE INTERACTIONS AND
THEIR COMMUNITY CONSEQUENCES
A THEORETICAL CONSIDERATION
ON THE ROLE OF ADAPTATION
IN BIOLOGICAL INVASIONS
- +ONDOH
).42/$5#4)/.
%COLOGICAL THEORY SUGGESTS THAT INTERSPECIlC INTERACTIONS ARISING BETWEEN AN ALIEN
SPECIES AND NATIVE SPECIES HOLD THE KEY TO INVASION SUCCESS BY AN ALIEN SPECIES
4HE WELL
KNOWN HYPOTHESIS THAT AN ALIEN SPECIES THAT HAS FEW NATURAL ENEMIES IS
MORE LIKELY TO BE A SUCCESSFUL INVADER EG "LOSSEY AND .TZOLD -ITCHELL
AND 0OWER IMPLIES THE IMPORTANCE OF A TROPHIC INTERACTION 4HE HYPOTHESIS
THAT A COMMUNITY WITH HIGHER SPECIES RICHNESS IS MORE RESISTANT TO A BIOLOGICAL
INVASION 3TACHOWICZ ET AL +ENNEDY ET AL IS STRONGLY BASED ON THE
COMPETITION THEORY 4ILMAN &URTHER THE WAY THROUGH WHICH A BIOLOGICAL
-7 #ADOTTE ET AL (EDS) #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY 225n
3PRINGER 0RINTED IN THE .ETHERLANDS
- +ONDOH
INVASION AFFECTS THE LOCAL COMMUNITY IS AN ALIEN
NATIVE INTERSPECIlC INTERACTION
SUCH AS RESOURCE COMPETITION TROPHIC INTERACTION AND ALLELOPATHY 4HE IMPORTANT
ROLE OF INTERSPECIlC INTERACTIONS IN A BIOLOGICAL INVASION SUGGESTS THE DETAILED
EVALUATION OF THE INTERSPECIlC INTERACTIONS BETWEEN ALIEN AND NATIVE SPECIES AS AN
INTERESTING APPROACH TO THE BIOLOGICAL INVASION ISSUES
4HERE ARE TWO DIFFERENT BUT NOT MUTUALLY EXCLUSIVE VIEWS OF WHAT CHARAC
TERIZES AN ALIEN
NATIVE INTERACTION THE hSPECIES
IDENTITYv AND hCONTACT
EXPERI
ENCEv VIEWS 4HE SPECIES
IDENTITY VIEW FOCUSES ON THE IDENTITY OF THE FOCAL SPECIES
EG "AKER 2OY +OLAR AND ,ODGE 4HE ALIEN AND NATIVE SPECIES
ARE CHARACTERIZED BY lXED SETS OF SPECIES
SPECIlC TRAITS SUCH AS COMPETITIVE ABILITY
(ERBOLD AND -OYLE 0ETREN AND #ASE (OLWAY "YERS
WHICH INmUENCE THEIR INTERACTION AND THUS THE INVASION SUCCESS AND ITS IMPACT
ON THE LOCAL COMMUNITY EG 7ILLIAMS )N THIS VIEW AN INTERSPECIlC INTER
ACTION THAT ARISES BETWEEN AN ALIEN AND NATIVE SPECIES IS EXPLAINED MAINLY BY
THEIR INHERENT SPECIES
SPECIlC TRAITS AND SO ARE ITS POPULATION
LEVEL CONSEQUENCES
-ACK ET AL
4HE CONTACT
EXPERIENCE VIEW 3AKAI ET AL FOCUSES ON THE LACK OF hCONTACT
EXPERIENCEv BETWEEN ALIEN AND NATIVE SPECIES AS THE MAJOR FACTOR CHARACTERIZING
THEIR INTERACTION .ATIVE SPECIES THAT COEXIST IN THE LOCAL HABITAT HAVE THE EXPERI
ENCE OF ENCOUNTERS OR INTERACTIONS WITH EACH OTHER OVER VARYING TIMESCALES WHICH
PROVIDES THE OPPORTUNITY FOR LOCAL SPECIES TO ADAPT TO EACH OTHER EG ,AWLOR AND
-AYNARD 3MITH )N CONTRAST AN ALIEN SPECIES AND A NATIVE SPECIES DUE TO
TEMPORAL OR SPATIAL SEPARATION SHARE A RELATIVELY SHORT HISTORY AND ARE LESS LIKELY
TO ADAPT TO EACH OTHER 4HE LACK OF ADAPTATION SHOULD HAVE MAJOR IMPACT ON
THE ALIEN
NATIVE INTERSPECIlC INTERACTION EG 7ARE $ILL -ARURRAN
#ROY AND (UGHES (UGHES ET AL -ALONEY AND -CLEAN
#ALLAWAY AND !SCHEHOUG 2EHAGE ET AL AND THE RESULTANT COMMUNITY
STRUCTURE 'RIFlN ET AL 2ICCIARDI ET AL 2ICCIARDI AND !TKINSON
(OWEVER ONLY A FEW STUDIES HAVE COMPREHENSIVELY CONSIDERED THE CONTACT EXPERI
ENCE EFFECT IN THE CONTEXT OF BIOLOGICAL INVASION ISSUES BUT SEE 3AKAI ET AL
.OW WE NEED A STRONG CONCEPTUAL BASIS FOR THE ISSUE OF HOW CONTACT EXPERIENCE
INmUENCES A BIOLOGICAL INVASION FOR GUIDING FURTHER RESEARCH
)N THIS CHAPTER ) PRESENT A THEORETICAL FRAMEWORK FOR APPLYING THE CONTACT
EXPERIENCE VIEW TO ALIEN
NATIVE TROPHIC INTERACTIONS ) CHOSE THIS PARTICULAR INTER
ACTION AS IT IS KNOWN TO PLAY THE ESSENTIAL ROLE IN SHAPING COMMUNITY STRUCTURE
EG (AIRSTONE ET AL &RETWELL 0OWER )T DRIVES DIVERSE PREY
PREDATOR POPULATION DYNAMICS INmUENCES POPULATION DENSITY AND DISTRIBUTION
(AIRSTONE ET AL GENERATES TWO MAJOR INTERSPECIlC INDIRECT INTERACTIONS
[RESOURCE COMPETITION 4ILMAN AND APPARENT COMPETITION (OLT
"ONSALL AND (ASSELL BUT SEE #HANETON AND "ONSALL ] THAT LIMIT
THE NUMBER OF COEXISTING SPECIES AND FORMS A MAJOR COMPONENT OF THE NATURAL
SELECTION PRESSURE %NDLER &URTHER THE MACROSCOPIC FOOD
WEB TOPOLOGY
CAN BE AN ESSENTIAL DETERMINANT OF POPULATION STABILITY 0IMM AND ECOSYS
TEM FUNCTION 4HBAULT AND ,OREAU
#ONTACT EXPERIENCE ALIEN
NATIVE INTERACTIONS
4HE TWO MAJOR QUESTIONS OF THE PRESENT CHAPTER ARE OF HOW THE LACK OF CON
TACT EXPERIENCE INmUENCES A TROPHIC INTERACTION AND WHAT IS THE POPULATION TO
COMMUNITY
LEVEL CONSEQUENCE OF A CONTACT EXPERIENCE
FREE INTERACTION 4O ANSWER
THESE QUESTIONS ) lRST EXPLAIN THE BEHAVIORAL BASIS OF A TROPHIC INTERACTION
4HEN ) WILL PROVIDE SEVERAL HYPOTHESES ON HOW ALIEN
NATIVE INTERSPECIlC TROPHIC
INTERACTIONS DIFFER FROM INTERACTIONS BETWEEN NATIVES AND HOW THIS DIFFERENCE
AFFECTS THE POPULATION OR COMMUNITY FOLLOWING A BIOLOGICAL INVASION
4(% #/.4!#4 %80%2)%.#% 6)%7 /& !,)%. .!4)6%
42/0()# ).4%2!#4)/.3
&AILURE OF RECOGNITION
#ONSIDER A DOG AND A PIECE OF MEAT THAT IS RELEASING AN ODOR )F HUNGRY ENOUGH
THE DOG WILL FOLLOW THE ODOR AND EAT THE MEAT THERE IS AN INDIVIDUAL
LEVEL TROPHIC
INTERACTION 4HE DOGS FORAGING BEHAVIOR CONSISTS OF THREE SUB
PROCESSES EG
'RIFlN ET AL hINFORMATION RECEPTIONv hINFORMATION PROCESSINGv AND
hBEHAVIORAL RESPONSEv 4HE DOG DETECTS THE ODOR USING CHEMICAL RECEPTORS AND
THE NERVOUS SYSTEM INFORMATION RECEPTION 4HE ODOR IS ASSOCIATED WITH MEAT
BASED ON THE DOGS PREVIOUS EXPERIENCE OR AN INBORN ASSOCIATION INFORMATION
PROCESSING 5SING THIS INFORMATION THE DOG MAKES A DECISION TO FOLLOW THE ODOR
AND LOCATES AND EATS THE MEAT BEHAVIORAL RESPONSE
)NFORMATION PROCESSING IS THE SUB
PROCESS BY WHICH A MEANING e.g., hTHERE
IS MEATv THAT IS CONVEYED BY THE CUE THE ODOR IS hUNDERSTOODv )N GENERAL
CUE ALONE OFTEN HAVE LITTLE TO DO WITH THE INFORMATION THAT IS CONVEYED 4HIS IS
ANALOGOUS TO THE FACT THAT HOW A WORD SOUNDS HAS LITTLE TO DO WITH WHAT THE WORD
ACTUALLY MEANS 4HUS SUCCESSFUL INFORMATION PROCESSING OR WHETHER THE MEANING
OF THE CUE IS CORRECTLY UNDERSTOOD DEPENDS ON THE CAPABILITY OF CUE
INFORMATION
ASSOCIATION EG 4HORNDIKE 0AVLOV )N THE CASE OF THE DOG AND THE
MEAT FOR EXAMPLE THE CHEMICAL COMPOSITION OF THE MEATS ODOR DOES NOT NECESSAR
ILY MEAN A BLOCK OF MEAT OR POTENTIAL FOOD WITHOUT THE CORRECT ASSOCIATION
4HE CUE
INFORMATION ASSOCIATION THE CORE OF INFORMATION PROCESSING IS SHAPED
THROUGH hEXPERIENCEv ON VARIOUS TIME SCALES .ATURAL SELECTION MIGHT CREATE
THE CORRECT ASSOCIATION IF THE CAPABILITY OF INFORMATION PROCESSING AFFECTS REPRO
DUCTIVE SUCCESS 2OEDER AND 4REAT WHILE AN INDIVIDUAL MIGHT LEARN THE
VISUAL CHEMICAL OR PHONIC CUES OF POTENTIAL PREY THROUGH CONTINUOUS ENCOUNTER
EXPERIENCES WITH THE PREY ON THE SHORTER TIME SCALE $OMJAN AND "URKHARD
&OSTER 4HIS EXPERIENCE DEPENDENCE IMPLIES THAT THE LACK OF CONTACT
EXPERIENCE BETWEEN A PREY AND PREDATOR INmUENCES THE BEHAVIORAL RESPONSES OF
ONE TO THE OTHER BY ALTERING INFORMATION PROCESSING AND CONSEQUENTLY AFFECTS
THEIR TROPHIC INTERACTION
)N TROPHIC INTERACTIONS IT IS CONVENIENT TO CONSIDER TWO DIFFERENT LEVELS OF RECOG
NITION hTROPHIC ROLE RECOGNITIONv AND hTROPHIC SPECIES RECOGNITIONv 4HE FORMER
- +ONDOH
IS RECOGNITION OF WHETHER THE INDIVIDUAL IS POTENTIAL PREY POTENTIAL PREDATOR OR
NEITHER WHILE THE LATTER IS RELATED TO MORE DETAILED RECOGNITION WITHIN A TROPHIC
ROLE SUCH AS DISTINCTION BETWEEN SPECIES OR ORGANISM TYPES THAT REQUIRE DIFFERENT
STRATEGIES TO COPE WITH )F TROPHIC ROLE RECOGNITION FAILS ALL THE BEHAVIORS WHICH
SHOULD FOLLOW A CONTACT WITH POTENTIAL PREYPREDATOR DOES NOT FOLLOW &ORAGING
BEHAVIOR WHICH NORMALLY FOLLOWS CONTACT WITH PREY MIGHT NOT OCCUR BETWEEN
TWO SPECIES THAT HAVE NO CONTACT EXPERIENCE WITH EACH OTHER EG 7ARE
#ROY AND (UGHES "OND AND +AMIL ! PREY MIGHT FAIL TO RECOGNIZE
#URIO AND CAN BE MORE VULNERABLE TO A NOVEL PREDATOR EG $ILL
-AGURRAN -ALONEY AND -CLEAN )F TROPHIC SPECIES RECOGNITION FAILS
AN INCORRECT BEHAVIORAL RESPONSE WHICH SHOULD FOLLOW A CONTACT WITH ANOTHER
TROPHIC SPECIES MAY FOLLOW THE ENCOUNTER 0OSSIBLE CONSEQUENCES OF THIS WILL BE
DISCUSSED LATER
&AILURE OF THE BEHAVIORAL RESPONSE
!NOTHER POSSIBLE MECHANISM THROUGH WHICH THE LACK OF CONTACT EXPERIENCE ALTERS
A TROPHIC INTERACTION IS A FAILURE IN THE THIRD SUB
PROCESS BEHAVIORAL RESPONSE
4HE STRATEGY REQUIRED TO CONSUME A PREY IS OFTEN PREY
SPECIES SPECIlC ! PREDATOR
HAS TO CHOOSE THE CORRECT PLACE AND TIME TO lND A PREY OR MIGHT NEED A SPECIlC
TECHNIQUE TO CAPTURE OR CONSUME THE PREY )N THIS SITUATION WHETHER A PREDATOR
CAN USE A NOVEL PREY DEPENDS ON THE APPLICABILITY OF THE PREDATORS FORAGING STRAT
EGY WHICH IS SHAPED THROUGH PREVIOUS INTERACTIONS WITH OTHER PREY SPECIES TO
THE NOVEL PREY SPECIES SKILL TRANSFER THEORY %LLIS (AZLETT )F THE NOVEL
PREY BEHAVES OR RESPONDS LIKE A PREY FAMILIAR TO THE PREDATOR THE PREDATOR MIGHT
BE ABLE TO HANDLE THE NOVEL PREY 2ICCIARDI AND !TKINSON AND A TROPHIC
INTERACTION WOULD OCCUR NORMALLY )F PREVIOUS FORAGING BEHAVIOR IS NOT APPLICABLE
TO THE NOVEL PREY THE PREDATOR IS UNLIKELY TO CONSUME THE NOVEL PREY SUCCESSFULLY
4HUS THE LACK OF CONTACT EXPERIENCE CAN PREVENT SUCCESSFUL CONSUMPTION BEHAV
IOR 7ARE #ROY AND (UGHES (UGHES ET AL 4HE SAME SHOULD
BE TRUE FOR THE PREYS ANTI
PREDATION BEHAVIOR 3OLUK 3IH ET AL
AS A BEHAVIOR REQUIRED TO AVOID A PREDATOR IS PREDATOR
SPECIlC )F THE PREY HAS
NO CONTACT EXPERIENCE WITH THE FOCAL PREDATOR AND IF THE PREYS ANTI
PREDATION
STRATEGY IS NOT APPLICABLE TO THE NOVEL PREDATOR THE CORRECT PREDATOR
SPECIlC
DEFENCE MIGHT NOT FOLLOW THE CONTACT
)N AN ALIEN
NATIVE INTERACTION THE BEHAVIORAL RESPONSE EITHER PREY CONSUMP
TION OR PREDATOR AVOIDANCE MIGHT FAIL EVEN IF THE COUNTERPART SPECIES IS NOT
NOVEL TO THE FOCAL SPECIES 4HIS IS BECAUSE A HABITAT
SPECIlC HISTORY CAN LEAD TO
DIVERGENCE IN THE WAY IN WHICH PREY IS USED OR PREDATORS ARE AVOIDED IN DIFFERENT
HABITATS EG "RODIE AND "RODIE 4HOMPSON #ONSIDER TWO SEPA
RATE HABITATS ! AND " INHABITED BY THE SAME SET OF PREY AND PREDATOR SPECIES
0REY
PREDATOR CO
ADAPTATION OCCURS INDEPENDENTLY IN THE TWO HABITATS IMPLY
ING THAT THE PREY AND PREDATOR FROM DIFFERENT HABITATS ARE NOT EXPERIENCED WITH
EACH OTHER 4HEREFORE THE PREY
PREDATOR INTERACTION ARISING FROM THE OPPOSITE
#ONTACT EXPERIENCE ALIEN
NATIVE INTERACTIONS
COMBINATION PREY FROM ! AND PREDATOR FROM " OR VICE VERSA CAN DIFFER FROM
THE ORIGINAL INTERACTION WITHIN (ABITAT ! OR "
#/.3%15%.#%3 /& 4(% #/.4!#4 %80%2)%.#%
&2%%
42/0()# ).4%2!#4)/.3
(IGH VARIANCE IN ALIEN NATIVE INTERACTION STRENGTH
!CCORDING TO BIOLOGICAL INVASION THEORY AN ALIEN PREY SPECIES CAN HAVE MORE
ADVANTAGES THAN A NATIVE PREY SPECIES IN THE LOCAL HABITAT BECAUSE THE ALIEN SPE
CIES IS LESS LIKELY TO HAVE NATURAL ENEMIES IN THE NEW HABITAT EG %LTON
"LOSSEY AND .TZOLD -ACK ET AL +EANE AND #RAWLEY -ITCHELL
AND 0OWER 4ORCHIN ET AL 4HIS IS KNOWN AS ENEMY RELEASE HYPOTHESIS
4HIS HYPOTHESIS SUGGESTS THAT THE LACK OF A NATIVE CONSUMERS CONTACT EXPERIENCE
WITH THE ALIEN PREY SPECIES LOWERS THE STRENGTH OF THE TROPHIC INTERACTION BETWEEN
THEM &ROM THE CONTACT EXPERIENCE POINT OF VIEW THERE ARE THREE POTENTIAL REASONS
FOR THIS HYPOTHETICAL PATTERN &IRST A NATIVE CONSUMER DOES NOT RECOGNIZE THE ALIEN
SPECIES AS POTENTIAL PREY 3ECOND THE NATIVE CONSUMER DOES NOT hKNOWv HOW TO
HANDLE THE NOVEL PREY SPECIES 4HIRD EVEN IF THE ALIEN PREY SPECIES IS NOT NOVEL
IT MIGHT BEHAVE IN A NOVEL WAY DUE TO A HABITAT
SPECIlC EVOLUTIONARY PROCESS
(OWEVER ) SUGGEST THAT THE ENEMY RELEASE HYPOTHESIS ONLY CONSIDERS A PART
OF THE TOTAL PICTURE OF ALIEN
NATIVE TROPHIC INTERACTIONS &IRST THE HYPOTHESIS
OVERLOOKS THE PREYS ANTI
PREDATORY BEHAVIOR !S THE LACK OF CONTACT EXPERIENCE
MAY LOWER THE ALIENS DEFENCE EFlCIENCY AGAINST THE NATIVE PREDATOR 2EHAGE
ET AL THEN THEIR INTERACTION STRENGTH CAN BE STRONGER 4AKING THIS EFFECT
INTO ACCOUNT THE LACK OF CONTACT EXPERIENCE HAS TWO OPPOSITE CONSEQUENCES
LOW PREDATION EFlCIENCY WHICH DECREASES THE INTERACTION STRENGTH AND LOW
DEFENCE EFlCIENCY WHICH INCREASES THE STRENGTH 3ECOND THE ENEMY RELEASE
HYPOTHESIS OVERLOOKS ALIENS PREY )F THE LACK OF CONTACT EXPERIENCE AFFECTS THE
ALIENS VULNERABILITY TO NATIVE NATURAL ENEMIES THIS SHOULD BE ALSO APPLICABLE TO
THE INTERACTION OF THE FOCAL ALIEN AS A PREDATOR AND A POTENTIAL PREY IN THE LOCAL
HABITAT )N THIS INTERACTION THE BEHAVIORS INmUENCED BY THE CONTACT EXPERIENCE
ARE THE ALIENS FORAGING BEHAVIOR AND THE NATIVES DEFENCE BEHAVIOR
)N SUMMARY FOUR DIFFERENT BEHAVIORS MUST BE CONSIDERED TO EVALUATE THE EFFECT
OF THE LACK OF CONTACT EXPERIENCE ON INTERACTION STRENGTH AND INVASION SUCCESS
THE ALIENS FORAGING BEHAVIOR AND ANTI
PREDATORY BEHAVIOR AND THE NATIVES FORAG
ING BEHAVIOR ON THE ALIEN AND ANTI
PREDATORY BEHAVIOR AGAINST THE ALIEN &IG
4HE COMPLEX INTERACTION BETWEEN THE FOUR INTER
RELATED BEHAVIORS MAKES CONSIS
TENT PREDICTION MORE DIFlCULT THAN PREVIOUSLY THOUGHT !SSUME THAT 3PECIES ! IS
INTRODUCED TO A LOCAL HABITAT IN WHICH IT INTERACTS WITH A NATIVE NATURAL ENEMY
3PECIES % AND A NATIVE PREY 3PECIES 2 AND ASSUME THAT THE STRENGTH OF A TROPHIC
INTERACTION IS DETERMINED BY THE BALANCE BETWEEN THE PREYS ANTI
PREDATORY DEFENCE
AND THE PREDATORS FORAGING BEHAVIOR 7HETHER THE LACK OF CONTACT EXPERIENCE
- +ONDOH
&IG &OUR BEHAVIORS AFFECTING THE TROPHIC INTERACTIONS BETWEEN AN ALIEN SPECIES AND A
NATIVE SPECIES )F PREDATOR AVOIDANCE IS MORE DEPENDENT ON CONTACT EXPERIENCE THAN IS PREY
CONSUMPTION AN ALIEN SPECIES WILL HAVE A STRONGER INTERACTION WITH THE NATIVE SPECIES
)F PREY CONSUMPTION IS MORE DEPENDENT ON CONTACT EXPERIENCE THAN IS PREDATOR AVOIDANCE
AN ALIEN SPECIES WILL HAVE A WEAKER INTERACTION WITH THE NATIVE SPECIES )F THERE IS NO
CONSISTENT TREND THE INTERACTION STRENGTH WILL BE CHARACTERIZED BY A HIGHER VARIANCE
BENElTS THE ALIEN SPECIES DEPENDS ON THE LEVEL OF EACH SPECIES CONTINGENT
PRE
ADAPTATION !GRAWAL AND +OTANEN 2ICCIARDI AND !TKINSON
TO AN INTERACTING SPECIES 4HE ALIEN SPECIES WOULD BE AT AN ADVANTAGE IF ITS PRE
ADAPTIVE DEFENCE LEVEL TO 3PECIES % WERE HIGHER THE PRE
ADAPTIVE DEFENCE LEVEL OF
3PECIES 2 TO 3PECIES ! WERE LOWER THE PRE
ADAPTIVE FORAGING LEVEL OF 3PECIES % TO
3PECIES ! WERE LOWER OR THE PRE
ADAPTIVE FORAGING LEVEL OF 3PECIES ! TO 3PECIES 2
WERE HIGHER
)F THERE IS A CONSISTENT PATTERN IN WHICH BEHAVIOR FORAGING OR ANTIPRE
DATORY DEFENCE OR TROPHIC ROLE IS MORE DEPENDENT ON CONTACT EXPERIENCE THEN
IT IS POSSIBLE TO MAKE A CONSISTENT PREDICTION ON HOW THE CONTACT EXPERIENCE AFFECTS
THE INTERACTION STRENGTH &OR EXAMPLE IF PREDATOR RELIES MORE ON THE EXPERIENCE
THAN PREY DOES A TROPHIC INTERACTION IN THE ABSENCE OF CONTACT EXPERIENCE WILL BE
WEAKER THAN AN IN ITS PRESENCE )F THE PREY IS MORE DEPENDENT ON THE EXPERIENCE
A TROPHIC INTERACTION BETWEEN NOVEL SPECIES WOULD BE STRONGER THAN THE NORMAL
ONE )F THERE IS NO CONSISTENT PATTERN IN WHICH ONE TROPHIC ROLE IS MORE DEPENDENT
ON CONTACT EXPERIENCE THE DISRUPTION OF THE PREDATION
DEFENCE BALANCE SHOULD
LEAD TO MORE VARIABLE TROPHIC INTERACTION STRENGTH 7HICH IS THE REAL PATTERN IN
NATURE )T WOULD BE INTERESTING TO EXAMINE HOW THE FREQUENCY DISTRIBUTION OF
THE INTERACTION STRENGTH OF ALIEN
NATIVE INTERACTIONS DIFFERS FROM THAT OF NATIVE
NATIVE INTERACTIONS
)T IS WELL RECOGNIZED THAT COMPETITIVE ABILITY IS AFFECTED BY THE INTERACTING EFFECTS
OF RESOURCE AVAILABILITY BOTTOM
UP EFFECT AND VULNERABILITY TO NATURAL ENEMIES
#ONTACT EXPERIENCE ALIEN
NATIVE INTERACTIONS
TOP
DOWN EFFECT EG 0OWER ,EIBOLD 4HEORY SUGGESTS THAT THE
COEXISTENCE OF SPECIES SHARING A SIMILAR RESOURCE AND NATURAL ENEMY REQUIRES A
TRADE
OFF BETWEEN RESOURCE COMPETITION AND NATURAL
ENEMY AVOIDANCE ,UBCHENCO
,EIBOLD )F THE STRENGTH OF AN ALIEN
NATIVE TROPHIC INTERACTION IS CHAR
ACTERIZED BY A HIGH VARIANCE IT MIGHT LEAD TO THE PREDICTION THAT THE COMPETITIVE
SUPERIORITY OF AN ALIEN SPECIES SHOULD BE ABNORMALLY HIGH OR LOW )F THE TROPHIC LINK
TO THE CONSUMPTION OF THE ALIEN SPECIES IS WEAK AND THE LINK FROM THE ALIEN SPECIES
IS STRONG THE ALIEN WILL HAVE A HIGH POPULATION DENSITY )F THE TROPHIC LINK TO THE
CONSUMPTION OF THE ALIEN SPECIES IS STRONG AND THE LINK FROM THE ALIEN SPECIES IS
WEAK THE ALIEN WILL HAVE A LOW POPULATION DENSITY 4HIS IMPLIES THAT ALTHOUGH
THERE ARE MANY ALIEN SPECIES THAT CANNOT BECOME ESTABLISHED WHEN AN ALIEN
SPECIES SUCCEEDS IN BECOMING ESTABLISHED IT TENDS TO BE VERY ABUNDANT
!LIEN
NATIVE MISMATCHING IN ADAPTIVE SPEED AND
hBOOM
AND
BUSTv PATTERN OF ALIEN POPULATIONS
!LTHOUGH THE LACK OF CONTACT EXPERIENCE MAY AS ) HAVE ARGUED IN THE LAST SECTION
LOWER THE FORAGING EFlCIENCY OR DEFENCE EFlCIENCY AT THE INITIAL STAGE OF INVASION
THE EFlCIENCY IS LIKELY TO BE GRADUALLY IMPROVED AS AN ALIEN SPECIES AND RESIDENT SPE
CIES LEARN OR EVOLVE TO ADAPT EACH OTHER #ARROLL AND $INGLE -ARON AND 6ILA
4HE PREVIOUS STUDIES THAT CONSIDER THE COADAPTIVE PROCESS HAVE STRESSED THE
DISADVANTAGE OF AN ALIEN SPECIES BY SUGGESTING THAT AN ALIEN SPECIES IS OFTEN CHARAC
TERIZED BY A LOW POPULATION DENSITY AND LOW GENETIC DIVERSITY WHICH LOWERS EVOLU
TION SPEED 4SUTSUI ET AL 4HIS IS WHY MULTIPLE INTRODUCTIONS WHICH INCREASE
GENETIC DIVERSITY ENHANCE ALIENS INVASION SUCCESS +OLBE ET AL
(OWEVER THE DISADVANTAGE OF ALIEN SPECIES CAN BE COMPENSATED !N EXAMPLE
IS ALIENS BEHAVIORAL PLASTICITY 'RAY 3OL AND ,EFEBVRE ANALYZED THE
RELATIONSHIP BETWEEN BRAIN SIZE IN BIRDS AND INVASION SUCCESS IN .EW :EALAND
THEY FOUND THAT SPECIES WITH LARGER BRAINS HAD A HIGHER PROBABILITY OF INTRODUCTION
SUCCESS THAN DID SPECIES WITH SMALLER BRAINS 4HIS PATTERN IS EXPLAINED BY THE EFFECT
OF BRAIN SIZE ON THE BIRDS INTERACTION WITH RESOURCES A BIRD WITH A LARGER BRAIN
IS BETTER AT DISCOVERING NEW FOODS IN THE NEW HABITAT 3OL AND ,EFEBVRE
4HIS VIEW PROVIDES A SPECIES IDENTITY EXPLANATION FOR INVASION SUCCESS IE A SPE
CIES WITH HIGH PLASTICITY IS A SUCCESSFUL INVADER
(ERE FROM THE VIEWPOINT OF CONTACT EXPERIENCE ) SHOW ANOTHER FACTOR THAT
MAY OVERCOME THE ALIENS DISADVANTAGE #ONSIDER THAT A SMALL NUMBER OF 3PECIES
! IS INTRODUCED TO A COMMUNITY WHERE A POTENTIAL PREDATOR 3PECIES % EXISTS
&IGS !FTER ITS INTRODUCTION THE ALIEN 3PECIES ! WOULD LEARN OR EVOLVE TO
LOWER THE PREDATION PRESSURE BY 3PECIES % WHILE 3PECIES % WOULD ADAPT TO INCREASE
THE PREDATION PRESSURE ON 3PECIES ! .OTE THAT IN THIS CO
ADAPTATION PROCESS
THE ADAPTIVE SPEEDS WOULD DIFFER DUE TO DIFFERENCES IN THE CONTACT OPPORTUNITIES
BETWEEN ALIEN AND NATIVE SPECIES 7HEN AN ALIEN SPECIES IS INTRODUCED ITS ABUN
DANCE SHOULD BE MUCH LOWER THAT THAT OF NATIVE SPECIES 4HIS IMPLIES THAT THE ALIEN
PREY HAS MORE OPPORTUNITY TO GAIN EXPERIENCE IN HANDLING THE NATIVE PREDATOR
- +ONDOH
&IG !LIEN NATIVE MISMATCHING IN EVOLUTIONARY SPEED LEADS TO BOOM AND BUST PATTERN
OF ALIEN POPULATIONS %ACH COLUMN IS FOR AN ALIEN PREDATOR AND ITS PREY LEFT COLUMN AND
AN ALIEN PREY AND ITS PREDATOR RIGHT COLUMN IN THE PRESENCE OF PREY PREDATOR COEVOLUTION
%ACH PANEL SHOWS POPULATION DYNAMICS TOP #. #! 2. 2! ADAPTIVE LEVEL MIDDLE THE
FRACTION OF ADAPTED INDIVIDUALS #!#. #! FOR THE PREDATOR AND 2!2. 2! FOR THE PREY
AND PER CAPITA PREDATION RATE BOTTOM ANN #. 2. ANA #! 2. AAN #. 2! AAA #! 2! ;#.
#! 2. 2! = )N THE TOP AND MIDDLE PANELS THE THICK LINES AND THIN LINES ARE FOR PREDATOR
AND PREY RESPECTIVELY #ONSIDER POPULATIONS OF AN ALIEN SPECIES AND NATIVE SPECIES EACH OF
WHICH CONSISTS OF MANY NON ADAPTED INDIVIDUALS WITH LOW DEFENCE ABILITY OR LOW FORAGING
ABILITY AND A FEW ADAPTED INDIVIDUALS WITH HIGH DEFENCE ABILITY OR HIGH FORAGING ABILITY
4HE POPULATION DYNAMICS IS DESCRIBED BY D #. D T RC +C n #. n #! #. E ANN #. 2.
E AAN #. 2! n MC #. D #! D T RC +C n #. n #! #! E AAA #. 2. E AAN #. 2! n MC #! D 2.
D T RR +R n 2. n 2! 2. n ANN #. 2. n ANA #! 2. n MR 2. D 2! D T RR +R n 2. n 2! 2! n AAN #.
2! n AAA #! 2! n MR 2! WHERE #. #! 2. AND 2! ARE POPULATION ABUNDANCES OF NON ADAPTED
PREDATOR ADAPTED PREDATOR NON ADAPTED PREY AND ADAPTED PREY RESPECTIVELY RI IS THE
INTRINSIC GROWTH RATE +I IS CARRYING CAPACITY SUPPORTED BY RESOURCES THAT ARE NOT EXPLICITLY
REPRESENTED BY THE EQUATIONS MI IS THE MORTALITY RATE OF THE PREY I R AND PREDATOR C E
IS THE ASSIMILATION EFlCIENCY 4HE PER CAPITA PREDATION RATES HOLD THAT ANA ANN AAA AAN
IMPLYING THAT THE PREDATION RATE OF THE NON ADAPTED PREDATOR ON THE NON ADAPTED PREY IS
SAME WITH THAT OF THE ADAPTED PREDATOR ON THE ADAPTED PREY 0ARAMETERS ANN AND AAA ARE SET
TO A VALUE WITH WHICH AN ALIEN SPECIES CANNOT PERSIST 4HIS SETTING CONlRMS THAT A TEMPORAL
INCREASE OF AN ALIEN SPECIES IS DUE TO THE ALIEN NATIVE MISMATCH IN EVOLUTIONARY SPEED
4HE INITIAL CONDITION IS THAT #. #! 2. 2!
FOR PREDATORS INVASION AND FOR PREYS INVASION 0ARAMETERS
THAT ARE USED ARE RC RN +C +N MC MN ANN ANA AAN AAA E
FOR PREDATORS INVASION AND
FOR PREYS INVASION
#ONTACT EXPERIENCE ALIEN NATIVE INTERACTIONS
&IG !LIEN
NATIVE MISMATCHING IN SPEED OF ADAPTIVE BEHAVIORAL PLASTICITY LEADS TO
BOOM
AND
BUST PATTERN OF ALIEN POPULATIONS %ACH COLUMN IS FOR AN ALIEN PREDATOR SPECIES
AND ITS PREDATOR PREY LEFT COLUMN AND AN ALIEN PREY SPECIES AND ITS PREDATOR RIGHT COLUMN
IN THE PRESENCE OF THE LEARNING PROCESS %ACH PANEL SHOWS POPULATION DYNAMICS TOP # 2
ADAPTIVE LEVEL MIDDLE !# FOR THE PREDATOR !2 FOR THE PREY AND INTERACTION STRENGTH
BOTTOM !# n A !2 )N THE TOP AND MIDDLE PANELS THE THICK LINES AND THIN LINES ARE FOR
PREDATOR AND PREY RESPECTIVELY ! SCALE FOR PREY POPULATION LEVEL IS INDICATED ON THE RIGHT OF
EACH PANEL #ONSIDER POPULATIONS OF A PREY SPECIES AND A PREDATOR SPECIES WHOSE POPULATION
DYNAMICS IS DESCRIBED BY D # D T RC +C n # # E & # 2 n MC # AND D 2 D T RR +R n 2
2 n & # 2 n MR 2 WHERE # AND 2 ARE POPULATION ABUNDANCES OF THE PREDATOR AND PREY
RESPECTIVELY RI IS THE INTRINSIC GROWTH RATE +I IS CARRYING CAPACITY SUPPORTED BY RESOURCES
THAT ARE NOT EXPLICITLY REPRESENTED BY THE EQUATIONS MI IS THE MORTALITY RATE OF THE PREY
I R AND PREDATOR C E IS THE ASSIMILATION EFlCIENCY 4HE PER
CAPITA PREDATION RATE
& IS A TEMPORAL VARIABLE AND DETERMINED BY PREDATORS FORAGING
RELATED ADAPTIVE LEVELS
!# AND PREYS DEFENCE
RELATED ADAPTIVE LEVEL !2 THAT IS & B !# n A !2 4HE ADAPTIVE
DYNAMICS IS GIVEN BY D !# D T 'C !#MAX n !# 2 AND D !2 D T 'R !2MAX n !2
!# n A !2 # WHERE 'C AND 'R ARE THE LEARNING RATES !#MAX AND !2MAX ARE THE MAXIMUM
LIMITS OF ADAPTIVE LEVELS FOR THE PREDATOR AND PREY RESPECTIVELY ) USED THE PARAMETERS
'C 'R !#MAX !2MAX ; n !#=A AND INITIAL CONDITION !# !2 !#
4HIS ASSUMPTION IMPLIES THAT THE PREDATION RATE OF THE NAIVE PREY WHOSE !2
AND NAIVE PREDATOR !# !# IS SAME WITH THAT OF THE MAXIMALLY ADAPTED PREDATOR
!# !#MAX ON THE MAXIMALLY ADAPTED PREY !2 ; n !#=A 0ARAMETER !# IS
SET TO A VALUE WITH WHICH AN ALIEN SPECIES CANNOT PERSIST TO CONlRM THAT A TEMPORAL
INCREASE OF AN ALIEN SPECIES IS DUE TO THE ALIEN
NATIVE MISMATCH IN ADAPTIVE SPEED
4HE INITIAL CONDITION IS THAT # 2 !# FOR PREDATORS INVASION AND
FOR PREYS INVASION 0ARAMETERS THAT ARE USED ARE RC RN +C +N MC MN
A B E FOR PREDATORS INVASION AND
FOR PREYS INVASION
- +ONDOH
THAN VICE VERSA 4HEREFORE THE ADAPTATION PROCESS IS ALIEN BIASED AND THE TROPHIC
INTERACTION BETWEEN A NATIVE PREDATOR AND AN ALIEN PREY IS LIKELY TO BE KEPT LOW
AFTER THE INTRODUCTION &IGS 4HE SAME IS TRUE FOR THE INTERACTION BETWEEN THE
ALIEN SPECIES 3PECIES ! AS A PREDATOR AND THE NATIVE PREY 3PECIES 2 &IGS
"ECAUSE THE OPPORTUNITY FOR THE ALIEN PREDATOR TO GAIN EXPERIENCE WITH THE NATIVE
PREY IS GREATER THAN FOR THE OPPOSITE SITUATION AN IMBALANCE CAN EXIST BETWEEN
THE ALIENS PREDATION EFlCIENCY AND THE NATIVES DEFENCE EFlCIENCY 4HEREFORE THE
TROPHIC INTERACTION BETWEEN THE ALIEN PREDATOR AND NATIVE PREY WOULD TEND TO BE
RELATIVELY HIGH IN THE TRANSIENT DYNAMICS
4HIS ALIEN NATIVE MISMATCH IN ADAPTIVE SPEED SUGGESTS THAT THE ALIEN SPECIES
HAS AN ADVANTAGE IN NOT BEING EXPERIENCED BY INTERACTING SPECIES 4HIS MAKES
AN ALIEN SPECIES LESS VULNERABLE TO NATURAL ENEMIES AND LOWERS THE PREYS DEFENCE
LEVEL AGAINST THE ALIEN SPECIES ALLOWING THE ALIEN SPECIES TO INCREASE ITS POPULATION
QUICKLY AFTER INTRODUCTION 4HIS ADVANTAGE WILL LAST UNTIL THE ALIEN SPECIES BECOMES
ABUNDANT AT WHICH POINT IT BECOMES EASIER FOR THE NATIVE SPECIES TO HAVE CONTACT
EXPERIENCE WITH THE ALIEN SPECIES !FTER THE ALIEN SPECIES BECOMES ABUNDANT
THE NATIVE SPECIES WILL CATCH UP WITH THE ALIEN SPECIES DEFENCE AND PREDATION
BEHAVIOR AGAINST THE ALIEN SPECIES WILL DEVELOP AND THE ALIEN SPECIES WILL BE CON
TROLLED MORE EFlCIENTLY 4HIS SUCCESSFULLY GENERATES THE hBOOM AND BUSTv PATTERN
OF ALIEN POPULATIONS THAT IS OFTEN OBSERVED IN NATURE &IGS 7ILLIAMSON AND
&ITTER
$ISRUPTION OF SWITCHING BEHAVIOR AND COMMUNITY INSTABILITY
!N ORGANISM CAN CHANGE THE SPECIES OR TROPHIC SPECIES TO INTERACT IN RESPONSE
TO A CHANGE IN THE RELATIVE ABUNDANCE OF POTENTIAL INTERACTING SPECIES !BRAMS
"OLKER ET AL 4HIS IS CALLED hSWITCHING BEHAVIORv -URDOCH
4HERE ARE TWO TROPHIC INTERACTION
RELATED SWITCHING BEHAVIORS FORAGING -URDOCH
AND DEFENCE &RYXELL AND ,UNDBERG SWITCHES ! FORAGING SWITCH
REFERS TO A CHOICE OF DIET WITH HIGHER QUALITY OR QUANTITY FROM A SET OF NUTRITIONALLY
SUBSTITUTABLE POTENTIAL DIETS A FORAGING SWITCH OR FORAGING SHIFT 3TEPHENS AND
+REBS )T IS A SIMPLE STRATEGY TO MAXIMIZE THE ENERGETIC OR MATERIAL GAIN PER
UNIT EFFORT WHEN DIFFERENT STRATEGIES ARE REQUIRED TO lND OR CAPTURE DIFFERENT DIETS
THE USE OF A LESS PROlTABLE RESOURCE LOWERS THE GAIN PER UNIT EFFORT 4HERE ARE A
NUMBER OF EXAMPLES IN WHICH ORGANISMS SWITCH TO MORE VALUABLE OR ABUNDANT
DIETS AS THE RELATIVE ABUNDANCE OR QUALITY OF THE POTENTIAL DIETS CHANGE SEE
3TEPHENS AND +REBS 4HE OTHER SWITCHING BEHAVIOR IS A DEFENCE SWITCH
#ONSIDER A PREY SPECIES WITH MULTIPLE PREDATORS AND ASSUME THAT THE AVOIDANCE
OF DIFFERENT PREDATORS REQUIRES DIFFERENT STRATEGIES WITH TRADE
OFFS BETWEEN THEM
4HIS MIGHT OCCUR WHEN PREDATORS ATTACK IN DIFFERENT MANNERS OR THE TOTAL PREDA
TOR AVOIDANCE EFFORT IS LIMITED 3OLUK 3IH ET AL )N SUCH A CASE A PREY
MUST ALLOCATE ITS DEFENCE EFFORT AMONG THE POTENTIAL PREDATORS WITH MORE EFFORT
BEING ALLOCATED TO THE MORE RISKY PREDATOR ,IMA
#ONTACT EXPERIENCE ALIEN
NATIVE INTERACTIONS
&IG &ROM THE VIEWPOINT OF CONTACT EXPERIENCE A BIOLOGICAL INVASION IS DIVIDED INTO
FOUR PHASES THE ) PRE
ADAPTATION )) ADAPTIVE
MISMATCHING ))) NATIVE
ADAPTATION AND
)6 POST
ADAPTATION PHASES &IRST JUST AFTER THE INTRODUCTION THE ALIEN AND NATIVE SPECIES
ARE NOT EXPERIENCED WITH EACH OTHER AND THEIR TROPHIC INTERACTION WHICH DEPENDS ON THE
PRE
ADAPTATION LEVEL OF EITHER SPECIES IS UNLIKELY TO HAVE A CONSISTENT PATTERN IE 0HASE )
PRE
ADAPTATION PHASE 3ECOND THE ALIEN SPECIES STARTS TO GAIN EXPERIENCE WITH THE NATIVE
SPECIES HOWEVER THE NATIVE SPECIES CANNOT GAIN EXPERIENCE WITH THE ALIEN SPECIES BECAUSE
THE ABUNDANCE OF THE ALIEN SPECIES IS LOW 4HIS ALIEN
BIASED ADAPTIVE PROCESS PROVIDES
THE ALIEN SPECIES WITH A MINORITY ADVANTAGE RESULTING IN THE RAPID GROWTH OF THE ALIEN
SPECIES IE 0HASE )) ADAPTIVE
MISMATCHING PHASE 4HIRD AFTER THE ALIEN SPECIES BECOMES
ABUNDANT THE NATIVE SPECIES BEGINS TO GAIN EXPERIENCE WITH THE ALIEN SPECIES IE 0HASE
))) NATIVE ADAPTATION PHASE $URING THIS THIRD PHASE THE LOCAL COMMUNITY CAN EFlCIENTLY
SUPPRESS AN ALIEN SPECIES )N THE LAST PHASE THE ALIEN AND NATIVE SPECIES ARE WELL ADAPTED TO
EACH OTHER 0HASE 6) POST
ADAPTATION PHASE AND THE POPULATION OF THE ALIEN SPECIES WILL
BECOME COMPARABLE TO THAT OF THE NATIVE SPECIES 4HE SOLID AND DOTTED LINES REPRESENT THE
SUPERIORITY OF AN ALIEN SPECIES TO A NATIVE SPECIES AND THE POPULATION ABUNDANCE OF THE ALIEN
SPECIES RESPECTIVELY ;3UPERIORITY= MEANS NO SUPERIORITY ;POPULATION LEVEL= MEANS
COMPARABLE ABUNDANCE OF THE ALIEN SPECIES TO THAT OF NATIVE SPECIES
4HEORY SUGGESTS THAT A FORAGING SWITCH ENHANCES SPECIES COEXISTENCE
EG 4ANSKY 'LEESON AND 7ILSON #ONSIDER A SIMPLE TROPHIC SYS
TEM CONSISTING OF TWO PREY SPECIES AND ONE PREDATOR SPECIES )N THE ABSENCE OF
A FORAGING SWITCH THE TWO PREY SPECIES MIGHT NOT COEXIST BECAUSE THE SPECIES
WITH LOW VULNERABILITY TO PREDATION TENDS TO OUT COMPETE THE OTHER THROUGH THE
NEGATIVE INDIRECT EFFECT MEDIATED BY THE SHARED PREDATOR APPARENT COMPETITION
(OLT "Y CONTRAST IN THE PRESENCE OF A FORAGING SWITCH THE TWO PREY SPECIES
CAN COEXIST BECAUSE THE SPECIES THAT IS LESS ABUNDANT TENDS TO BE ATTACKED LESS OFTEN
BY THE SWITCHING PREDATOR /NE PREY SPECIES CAN PROVIDE ANOTHER PREY SPECIES
WITH AN OPPORTUNITY TO AVOID THE PREDATOR 4HE SAME IS TRUE FOR MORE COMPLEX
SYSTEMS +ONDOH
- +ONDOH
! DEFENCE SWITCH ALSO ENHANCES SPECIES COEXISTENCE ,IMA -ATSUDA ET AL
)N A SIMPLE SYSTEM CONSISTING OF TWO PREDATOR SPECIES AND ONE PREY
SPECIES THE TWO PREDATOR SPECIES TEND NOT TO COEXIST BECAUSE RESOURCE COMPETITION
LEADS TO THE EXTINCTION OF THE INFERIOR PREDATOR SPECIES 4ILMAN )N THE PRES
ENCE OF AN ADAPTIVE DEFENCE SWITCH HOWEVER A COMPLETELY DIFFERENT PICTURE ARISES
!N ADAPTIVE PREY IS MORE LIKELY TO BE DEFENSIVE AGAINST THE MORE ABUNDANT PREDA
TOR SPECIES CREATING A MECHANISM OF MINORITY ADVANTAGE 4HEREFORE THE SWITCH
ING PREY CAN ENHANCE THE PREDATORS COEXISTENCE -ATSUDA ET AL SHOWED
THAT AN ADAPTIVE DEFENCE PREVENTS THE EXTINCTION OF THE PREDATOR SPECIES IN A TWO
TROPHIC
LEVEL SYSTEM WITH HIGHER SPECIES RICHNESS SUGGESTING THAT THE DEFENCE
SWITCH CAN ENHANCE SPECIES COEXISTENCE IN A COMPLEX FOOD WEB 4HIS IS ALSO TRUE IN
AN EVEN MORE COMPLEX FOOD WEB +ONDOH UNPUBLISHED MANUSCRIPT
)N GENERAL ADAPTIVE SWITCHING BEHAVIOR REQUIRES THE ABILITY TO DISCRIMINATE
MULTIPLE PREY OR PREDATOR SPECIES AND KNOWLEDGE OF THE RELATIVE QUANTITY OR QUAL
ITY OF THE POTENTIALLY INTERACTING SPECIES 3TEPHENS AND +REBS #ONSIDER ONE
PREDATOR AND MULTIPLE PREY SPECIES FOR EXAMPLE 4HE FORAGER IS UNLIKELY TO BE ABLE
TO CHOOSE THE MORE ABUNDANT OR MORE NUTRITIOUS PREY SPECIES IF THIS PREY SPECIES
IS NOT DIFFERENTIATED FROM OTHER PREY SPECIES 3IMILARLY IN THE ABSENCE OF SPE
CIES RECOGNITION ABILITY A PREY SPECIES IS UNLIKELY TO MAKE A CORRECT SWITCH IN ITS
DEFENCE BEHAVIOR AGAINST PREDATORS
4HE FACT THAT TROPHIC INTERACTION
RELATED SWITCHING BEHAVIOR WHICH REQUIRES
CORRECT RECOGNITION OF THE INTERACTING SPECIES TENDS TO ENHANCE SPECIES COEX
ISTENCE SUGGESTS ANOTHER IMPORTANT ROLE OF CONTACT EXPERIENCE IN MAINTAINING
BIODIVERSITY !N ALIEN
NATIVE INTERACTION IN WHICH SPECIES RECOGNITION IS LESS LIKELY
TO OCCUR MIGHT AFFECT BIODIVERSITY MAINTENANCE IN A COMPLETELY DIFFERENT WAY FROM
THAT OF A NATIVE
NATIVE INTERACTION -ORE SPECIlCALLY AN ALIEN
NATIVE INTERACTION
MIGHT BE MORE LIKELY TO LOWER THE PROBABILITY OF SPECIES COEXISTENCE THAN WOULD
A NATIVE
NATIVE INTERACTION
4HIS INTERPRETATION IS BASED ON A MODEL ANALYSIS BY +ONDOH 5SING A
DYNAMIC MATHEMATICAL MODEL OF A FOOD WEB &IG +ONDOH REVEALED
AN IMPORTANT ROLE OF THE FORAGING SWITCH IN BIODIVERSITY MAINTENANCE #ONSIDER A
FOOD WEB OF . SPECIES EACH OF WHICH IS CONNECTED BY A DIRECTED TROPHIC LINK WITH
PROBABILITY # CONNECTANCE )N THIS MODEL # DETERMINES THE NUMBER OF POTENTIAL
PREY SPECIES THAT A CONSUMER CAN USE IE ;.
=# ON AVERAGE ! CONSUMER SPE
CIES IS CAPABLE OF CHOOSING THE MORE ABUNDANT PREY SPECIES FROM THE POTENTIAL DIET
WITH PROBABILITY & "Y CONTRAST A FRACTION
& OF PREDATORS DOES NOT DISCRIMINATE
BETWEEN PREY SPECIES AND USES EVERY POTENTIAL PREY +ONDOH ANALYZED
THE INTERACTIVE EFFECT OF THE PREDATORS CAPABILITY OF AN ADAPTIVE FORAGING SWITCH
AND THE FOOD
WEB COMPLEXITY ON COMMUNITY PERSISTENCE 4HE ANALYSIS SHOWED
THAT AN ADAPTIVE FORAGING SWITCH POTENTIALLY INVERTS THE COMPLEXITY
STABILITY RELA
TIONSHIP OF A FOOD WEB )N THE ABSENCE OF ADAPTIVE SWITCHING BEHAVIOR INCREASING
FOOD
WEB COMPLEXITY LOWERS COMMUNITY PERSISTENCE WHILE IN THE PRESENCE OF
ADAPTIVE SWITCHING A COMMUNITY IS MORE LIKELY TO BE PERSISTENT WHEN THE FOOD
WEB STRUCTURE IS MORE COMPLEX +ONDOH &URTHER THE RELATIONSHIP BECOMES
#ONTACT EXPERIENCE ALIEN
NATIVE INTERACTIONS
&IG 0OPULATION PERSISTENCE CHANGING WITH CHANGING FRACTION OF ADAPTIVE FORAGERS FOR
VARYING FOOD
WEB COMPLEXITY # n . n #ONSIDER A FOOD WEB WITH . SPECIES
AND CONNECTION PROBABILITY # 0OPULATION DYNAMICS OF 3PECIES I IS GIVEN BY
8I "RI n SI 8I -EIK FKI AKI 8K n -FKI AKI 8K "
K D SP IS PREY K D SP IS PREDATOR
WHERE RI IS THE INTRINSIC GROWTH RATE SI THE SELF REGULATION STRENGTH EIJ THE CONVERSION
EFlCIENCY FIJ THE FORAGING EFlCIENCY OF PREDATOR I ON PREY J AIJ THE FORAGING EFFORT OF PREDATOR
I ALLOCATED TO PREY J 4HE FORAGING EFFORT OF ADAPTIVE FORAGER CHANGES TO INCREASE THE ENERGY
GAIN PER UNIT EFFORT
IJ 'AIJ "EIJ FIJ 8J n -EIK FKI AKI 8K "
K D SP IS PREY
3EE +ONDOH FOR MORE DETAIL ) USED THE FOLLOWING PARAMETERS TO CREATE THIS lGURE
8I AIJ RI SI EIJ FIJ ' RAN; = THE NUMBER OF POTENTIAL PREY SPECIES
RAN; = RAN; = 2ANDOM MODEL IS USED AS A FOOD WEB TOPOLOGY
0OPULATION PERSISTENCE IS DElNED BY LN;THE PROBABILITY THAT NO SPECIES GO EXTINCT=.
- +ONDOH
CLOSER TO NEGATIVE AS THE FRACTION OF ADAPTIVE FORAGERS DECREASES OR THE SPEED OF
THE ADAPTIVE SWITCH DECREASES +ONDOH
! POSSIBLE INTERPRETATION OF +ONDOHS RESULT IS THAT A COMMUNITY WITH
A GREATER FRACTION OF ALIEN SPECIES IS MORE LIKELY TO LOSE A SPECIES ESPECIALLY IN A
MORE COMPLEX FOOD WEB &IG "ECAUSE AN ALIEN SPECIES IS LESS LIKELY TO DISCRIMI
NATE BETWEEN NOVEL NATIVE SPECIES IT MIGHT NOT BE ABLE TO SWITCH DIETS CORRECTLY
IMPLYING THAT A CORRECT FORAGING SWITCH IS LESS LIKELY TO OCCUR AS THE FRACTION OF
ALIEN SPECIES INCREASES IN THE FOCAL FOOD WEB &URTHER IF A NATIVE PREDATOR CONFUSES
AN ALIEN SPECIES WITH A NATIVE PREY THE PREDATOR MIGHT FAIL TO EVALUATE THE RELA
TIVE ABUNDANCE OR QUALITY OF THE NATIVE SPECIES 4HUS THE INTRODUCTION OF AN ALIEN
SPECIES INTERRUPTS SWITCHING BEHAVIOR AND CAN THREATEN LOCAL BIODIVERSITY BY LOW
ERING THE POPULATION STABILITY
)N THIS SCENARIO THE EFFECT OF AN ALIEN SPECIES IS ANALOGOUS TO THE CONFUSION OF
THE BODYS SIGNALING SYSTEM CAUSED BY ENDOCRINE DISRUPTORS #OLBORN ET AL
%NDOCRINE DISRUPTORS OCCUPY A HORMONE RECEPTOR AND ARE MISTAKENLY RECOG
NIZED AS THE CORRECT HORMONE RESULTING IN A BODY DISORDER #OLBORN ET AL
4HE OCCUPATION INTERFERES WITH THE PROPER HORMONE SIGNAL AND PRODUCES AN
EXCESSIVE OR INSUFlCIENT CELLULAR RESPONSE #OLBORN ET AL )F BIODIVERSITY
IS MAINTAINED BY ADAPTIVE SWITCHING BEHAVIOR AS +ONDOH SUGGESTS AN
ALIEN SPECIES THAT CONFUSES SPECIES RECOGNITION MIGHT BE REGARDED AS AN ENDOCRINE
DISRUPTOR AT THE COMMUNITY LEVEL
35--!29
4HIS CHAPTER PRESENTS A HYPOTHESIS OF THE WAY IN WHICH THE LACK OF CONTACT EXPERI
ENCE INmUENCES THE TROPHIC INTERACTION BETWEEN ALIEN AND NATIVE SPECIES ! LACK OF
CONTACT EXPERIENCE CAN LEAD TO A FAILURE OF TROPHIC ROLE RECOGNITION PREY HANDLING
OR PREDATOR AVOIDANCE WHICH CAN RESULT IN A TROPHIC INTERACTION WITH ABNORMALLY
HIGH OR LOW STRENGTH AND DESTABILIZED PREY
PREDATOR INTERACTIONS ! MISMATCH IN
THE ADAPTATION SPEED BETWEEN ALIEN AND NATIVE SPECIES CAN PRODUCE A TEMPORAL
EXPLOSION IN THE ALIEN SPECIES ABUNDANCE ! FAILURE OF SPECIES DISCRIMINATION IN
AN ALIEN
NATIVE INTERACTION CAN DISRUPT THE ADAPTIVE RESPONSE TO QUALITATIVE AND
QUANTITATIVE CHANGES IN THE PREY OR PREDATOR AND CAN THEREBY THREATEN BIODI
VERSITY 4HESE HYPOTHESES IMPLY THAT AN ALIEN
NATIVE INTERACTION IS QUALITATIVELY
DIFFERENT FROM A NATIVE
NATIVE INTERACTION AND CREATE A GENERAL PATTERN OF ALIEN
SPECIES INVASIONS AND THEIR EFFECTS ON LOCAL COMMUNITIES
.EVERTHELESS THESE HYPOTHESES STILL RELY ON A NUMBER OF SIMPLIFYING ASSUMP
TIONS CONCERNING THE EFFECT OF EXPERIENCE ON A TROPHIC INTERACTION 4HESE ASSUMPTIONS
TIONS ARE REQUIRED BECAUSE WE KNOW LITTLE ABOUT HOW CONTACT EXPERIENCE AFFECTS
SUCH MACROSCOPIC PROPERTIES AS TROPHIC INTERACTION STRENGTH TROPHIC LINK mEXIBIL
ITY AND POPULATION STABILITY 7E NEED TO EXAMINE THE BEHAVIORAL BASIS OF A TROPHIC
INTERACTION AND TEST THE POSSIBILITY THAT AN ALIEN
NATIVE INTERACTION IS QUALITATIVELY
DIFFERENT FROM A NATIVE
NATIVE INTERACTION )N SO DOING IT IS IMPORTANT TO ISOLATE
#ONTACT EXPERIENCE ALIEN
NATIVE INTERACTIONS
THE EFFECT OF THE ABSENCE OF EXPERIENCE FROM THAT OF SPECIES IDENTITY ! SWAPPING
EXPERIMENT BETWEEN TWO COMMUNITIES WITH THE SAME SPECIES COMPOSITION AND
DIFFERENT HISTORIES MAY PROVIDE AN INTERESTING OPPORTUNITY THIS TEST
!#+./7,%$'-%.4
4HIS STUDY WAS PARTIALLY SUPPORTED BY A *303 2ESEARCH &ELLOWSHIP FOR 9OUNG
3CIENTISTS AND AN -%84 'RANT
IN
!ID FOR THE ST #ENTURY #/% 0ROGRAM ! TO
+YOTO 5NIVERSITY
2%&%2%.#%3
!BRAMS 0 ! &ORAGING TIME OPTIMIZATION AND INTERACTIONS IN FOOD WEBS 4HE
!MERICAN .ATURALIST
!GRAWAL ! ! AND 0 - +OTANEN (ERBIVORES AND THE SUCCESS OF EXOTIC PLANTS
A PHYLOGENETICALLY CONTROLLED EXPERIMENTS %COLOGY ,ETTERS
"AKER ( ' 4HE EVOLUTION OF WEEDS !NNUAL 2EVIEW OF %COLOGY AND 3YSTEMATICS
"LOSSEY . AND 2 .TZOLD %VOLUTION OF INCREASED COMPETITIVE ABILITY IN INVASIVE
NON
INDIGENOUS PLANTS A HYPOTHESIS *OURNAL OF %COLOGY
"OLKER " - (OLYOAK 6 +IVAN , 2OWE AND / 3CHMITZ #ONNECTING THEORETICAL
AND EMPIRICAL STUDIES OF TRAIT
MEDIATED INTERACTIONS %COLOGY
"OND ! " AND ! # +AMIL !POSTATIC SELECTION BY BLUE JAYS PRODUCES BALANCED
POLYMORPHISM IN VIRTUAL PREY .ATURE
"ONSALL - " AND - 0 (ASSELL !PPARENT COMPETITION STRUCTURES ECOLOGICAL ASSEM
BLAGES .ATURE
"RODIE % $ AND % $ "RODIE %VOLUTIONARY RESPONSE OF PREDATORS TO DANGEROUS PREY
REDUCTION OF TOXICITY OF NEWTS AND RESISTANCE OF GARTER SNAKES IN ISLAND POPULATIONS
%VOLUTION
"YERS * % #OMPETITION BETWEEN TWO ESTUARINE SNAILS IMPLICATIONS FOR INVASIONS
OF EXOTIC SPECIES %COLOGY
#ALLAWAY 2 - AND % 4 !SCHEHOUG )NVASIVE PLANTS VERSUS THEIR NEW AND OLD
NEIGHBORS A MECHANISM FOR EXOTIC INVASION 3CIENCE
#ARROLL 3 AND ( $INGLE 4HE BIOLOGY OF POST INVASION EVENTS "IOLOGICAL #ONSERVATION
#HANETON % * AND - " "ONSALL %NEMY
MEDIATED APPARENT COMPETITION EMPIRICAL
PATTERNS AND THE EVIDENCE /IKOS
#OLBORN 4 $ $UMANOSKI AND * 0 -YERS /UR 3TOLEN &UTURE 0ENGUIN "OOKS
.EW 9ORK .9
#ROY - ) AND 2 . (UGHES 4HE ROLE OF LEARNING AND MEMORY IN THE FEEDING
BEHAVIOUR OF THE lFTEEN
SPINED STICKLEBACK 3PINACHIA SPINACHIA , !NIMAL "EHAVIOUR
- +ONDOH
#URIO % 0ROXIMATE AND DEVELOPMENTAL ASPECTS OF ANTIPREDATOR BEHAVIOR !DVANCES
IN THE 3TUDY OF "EHAVIOR
$ILL , - 4HE ESCAPE RESPONSE OF THE ZEBRA DONIO "RACHYDANIO RERIO )) 4HE EFFECT OF
EXPERIENCE !NIMAL "EHAVIOUR
$OMJAN - AND " "URKHARD 4HE PRINCIPLES OF LEARNING BEHAVIOR ND EDITION
"ROOKS #OLE -ONTERY #!
%LLIS ( # 4HE TRANSFER OF ,EARNING -ACMILLAN .EW 9ORK .9
%LTON # 3 4HE %COLOGY OF )NVASIONS BY !NIMALS AND 0LANTS -ETHUEN ,ONDON
5+
%NDLER * ! $EFENCE AGAINST PREDATORS 0AGES
IN 0REDATOR
0REY 2ELATIONSHIPS
- % &EDER AND ' 6 ,AUDER EDITORS 5NIVERSITY OF #HICAGO 0RESS #HICAGO ),
&OSTER 3 ! 4HE GEOGRAPHY OF BEHAVIOR AN EVOLUTIONARY PERSPECTIVES 4RENDS IN
%COLOGY AND %VOLUTION
&RETWELL 3 $ 4HE REGULATION OF PLANT COMMUNITIES BY FOOD CHAINS EXPLOITING THEM
0ERSPECTIVE IN "IOLOGY AND -EDICINE
&RYXELL , - AND 0 ,UNDBERG )NDIVIDUAL "EHAVIOR AND #OMMUNITY $YNAMICS
#HAPMAN (ALL .EW 9ORK .9
'LEESON 3 + AND $ 3 7ILSON %QUILIBRIUM DIET OPTIMAL FORAGING AND PREY COEXIS
TENCE /IKOS
'RAY ! * $O INVADING SPECIES HAVE DElNABLE GENETIC CHARACTERISTICS 0HILOS 4RANS
2 3OC ,ONDON 3ER "
'RIFlN ! 3 $ 4 "LUMSTEIN AND # 3 %VANS 4RAINING ANIMALS ABOUT PREDATORS
A CRITICAL REVIEW AND SYNTHESIS #ONSERV "IOLO
(AIRSTONE . ' & % 3MITH AND , " 3LOBODKIN #OMMUNITY STRUCTURE POPULATION
CONTROL AND COMPETITION 4HE !MERICAN .ATURALIST
(AZLETT " ! "EHAVIOURAL PLASTICITY IN CRUSTACEA WHY NOT MORE *OURNAL OF
%XPERIMENTAL -ARINE "IOLOGY AND %COLOGY
(ERBOLD " AND 0 -OYLE )NTRODUCED SPECIES AND VACANT NICHES 4HE !MERICAN
.ATURALIST
(OLT 2 $ 0REDATION APPARENT COMPETITION AND THE STRUCTURE OF PREY COMMUNITIES
4HEORETICAL 0OPULATION "IOLOGY
(OLWAY $ ! #OMPETITIVE MECHANISMS UNDERLYING THE DISPLACEMENT OF NATIVE ANTS
BY THE INVASIVE ARGENTINE ANTS %COLOGY
(UGHES 2 . - * +AISER 0 ! -ACKNEY AND + 7ARBURTON /PTIMIZING FORAGING
BEHAVIOUR THROUGH LEARNING *OURNAL OF &ISH "IOLOGY
+EANE 2 - AND - * #RAWLEY %XOTIC PLANT INVASIONS AND THE ENEMY RELEASE
HYPOTHESIS 4RENDS IN %COLOGY AND %VOLUTION
+ENNEDY 4 3 .AEEM + (OWE * - ( +NOPS $ 4ILMAN AND 0 " 2EICH
"IODIVERSITY AS A BARRIER TO ECOLOGICAL INVASION .ATURE
+OLAR # AND $ - ,ODGE 0ROGRESS IN INVASION BIOLOGY PREDICTING INVADERS 4RENDS
IN %COLOGY AND %VOLUTION
+OLBE * * 2 % 'LOR , 2ODRIGUEZ 3CHETTINO ! # ,ARA ! ,ARSON ! AND * " ,OSOS
'ENETIC VARIATION INCREASES DURING THE BIOLOGICAL INVASION OF A #UBAN LIZARD
.ATURE
#ONTACT EXPERIENCE ALIEN
NATIVE INTERACTIONS
+ONDOH - &ORAGING ADAPTATION AND THE RELATIONSHIP BETWEEN FOOD WEB COMPLEXITY
AND STABILITY 3CIENCE
,AWLOR , 2 AND * -AYNARD 3MITH 4HE COEVOLUTION AND STABILITY OF COMPETING SPE
CIES 4HE !MERICAN .ATURALIST
,EIBOLD - ! ! GRAPHICAL MODEL OF KEYSTONE PREDATORS IN FOOD WEBS TROPHIC REGU
LATION OF ABUNDANCE INCIDENCE AND DIVERSITY PATTERNS IN COMMUNITIES 4HE !MERICAN
.ATURALIST
,IMA 3 , ,IFE IN A MULTI PREDATOR ENVIRONMENT SOME CONSIDERATIONS FOR ANTIPREDA
TORY VIGILANCE !NNALES :OOLOGICI &ENNICI
,UBCHENCO * 0LANT SPECIES DIVERSITY IN A MARINE INTERTIDAL COMMUNITY IMPORTANCE
OF HERBIVORE FOOD PREFERENCE AND ALGAL COMPETITIVE ABILITIES 4HE !MERICAN .ATURALIST
-ACK 2 . $ 3IMBERLOFF 7 - ,ONSDALE ( %VANS - #LOUT AND & ! "AZZAZ
"IOTIC INVASIONS CAUSES EPIDEMIOLOGY GLOBAL CONSEQUENCES AND CONTROL %COLOGICAL
!PPLICATIONS
-ALONEY 2 & AND ) ' -CLEAN (ISTORICAL AND EXPERIMENTAL LEARNED PREDATOR
RECOGNITION IN FREE LIVING .EW :EALAND 2OBINS !NIMAL "EHAVIOUR
-ARON * , AND - 6ILA 7HEN DO HERBIVORES AFFECT PLANT INVASION %VIDENCE FOR THE
NATURAL ENEMIES AND BIOTIC RESISTANCE HYPOTHESES /IKOS
-AGURRAN ! % !CQUIRED RECOGNITION OF PREDATOR ODOUR IN THE %UROPEAN MINNOW
0HOXINUS PHOXINUS %THOLOGY
-ATSUDA ( 0 ! !BRAMS AND - (ORI 4HE EFFECT OF ADAPTIVE ANTIPREDATOR BEHAVIOR
ON EXPLOITATIVE COMPETITION AND MUTUALISM BETWEEN PREDATORS /IKOS
-ATSUDA ( - (ORI AND 0 ! !BRAMS %FFECTS OF PREDATOR SPECIlC DEFENSE ON BIO
DIVERSITY AND COMMUNITY COMPLEXITY IN TWO TROPHIC LEVEL COMMUNITIES %VOLUTIONARY
%COLOGY
-ITCHELL # % AND ! ' 0OWER 2ELEASE OF INVASIVE PLANTS FROM FUNGAL AND VIRAL
PATHOGENS .ATURE
-URDOCH 7 7 3WITCHING IN GENERAL PREDATORS EXPERIMENTS ON PREDATOR SPECIlCITY
AND STABILITY OF PREY POPULATIONS %COLOGICAL -ONOGRAPHS
0AVLOV ) 0 #ONDITIONED RELAXES /XFORD 5NIVERSITY 0RESS .EW 9ORK .9
0ETREN + AND #ASE 4 * !N EXPERIMENTAL DEMONSTRATION OF EXPLOITATIVE COMPETITION
IN AN ONGOING INVASION %COLOGY
0IMM 3 , 4HE "ALANCE OF .ATURE 5NIVERSITY OF #HICAGO 0RESS #HICAGO
0OWER - % 4OP DOWN AND BOTTOM UP FORCES IN FOOD WEBS $O PLANTS HAVE PRIMACY
%COLOGY
2EHAGE * 3 "ARNETT " + AND 3IH ! "EHAVIORAL RESPONSES TO A NOVEL PREDATOR
AND COMPETITOR OF INVASIVE MOSQUITOlSH AND THEIR NON INVASIVE RELATIVES 'AMBUSIA SP
"EHAVIOURAL %COLOGY AND 3OCIOBIOLOGY ONLINE $/) S
2ICCIARDI ! 2 * .EVES AND * " 2ASMUSSEN )MPENDING EXTINCTIONS OF .ORTH
!MERICAN FRESHWATER MUSSELS 5NIONOIDA FOLLOWING THE ZEBRA MUSSEL $REISSENA POLY
MORPHA INVASION *OURNAL OF !NIMAL %COLOGY
2ICCIARDI ! AND 3 + !TKINSON $ISTINCTIVENESS MAGNIlES THE IMPACT OF BIOLOGICAL
INVADERS IN AQUATIC ECOSYSTEMS %COLOGY ,ETTERS
- +ONDOH
2OEDER + $ AND ! % 4REAT 4HE DETECTION AND EVASION OF BATS BY MOTHS !MERICAN
3CIENTIST
2OY * )N SEARCH OF THE CHARACTERISTICS OF PLANT INVADERS 0AGES
IN & DI
#ASTRI ! * (ANSEN AND - $EBUSSCHE EDITORS "IOLOGICAL INVASIONS IN %UROPE AND
THE -EDITERRANEAN "ASIN +LUWER $ORDORECHT 4HE .ETHERLANDS
3AKAI ! + & 7 !LLENDORF * 3 (OLT $ - ,ODGE * -OLOFSKY + ! 7ITH 3 "AUGHMAN
2 * #ABIN * % #OHEN . # %LLSTRAND $ % -C#AULEY 0 /.EIL ) - 0ARKER * .
4HOMPSON AND 3 ' 7ELLER 4HE POPULATION BIOLOGY OF INVASIVE SPECIES !NNUAL
2EVIEW OF %COLOGY AND 3YSTEMATICS
3IH ! ' %NGLUND AND $ 7OOSTER %MERGENT IMPACTS OF MULTIPLE PREDATORS ON PREY
4RENDS IN %COLOGY AND %VOLUTION
3OL $ AND , ,EFEBVRE &OREBRAIN SIZE AND FORAGING INNOVATIONS PREDICT INVASION
SUCCESS IN BIRDS INTRODUCED TO .EW :EALAND /IKOS
3OLUK $ ! -ULTIPLE PREDATOR EFFECTS PREDICTING COMBINED FUNCTIONAL RESPONSE OF
STREAM lSH AND INVERTEBRATE PREDATORS %COLOGY
3TACHOWICZ * * 2 " 7HITLATCH AND 2 7 /SMAN 3PECIES DIVERSITY AND INVASION
RESISTANCE IN A MARINE ECOSYSTEM 3CIENCE
3TEPHENS $ 7 AND * 2 +REBS &ORAGING 4HEORY 0RINCETON 5NIVERSITY 0RESS
0RINCETON .*
4ANSKY - 3WITCHING EFFECT IN PREY
PREDATOR SYSTEM *OURNAL OF 4HEORETICAL "IOLOGY
4HBAULT % AND - ,OREAU &OOD
WEB CONSTRAINTS ON BIODIVERSITYnECOSYSTEM FUNC
TIONING RELATIONSHIPS 0ROCEEDINGS OF THE .ATIONAL !CADEMY OF 3CIENCES
4HOMPSON * . 4HE COEVOLUTIONARY PROCESS 5NIVERSITY OF #HICAGO 0RESS #HICAGO
4HORNDIKE % , !NIMAL INTELLIGENCE EXPERIMENTAL STUDIES -ACMILLAN .EW 9ORK
.9
4ILMAN $ 2ESOURCE #OMPETITION AND #OMMUNITY 3TRUCTURE 0RINCETON 5NIVERSITY
0RESS 0RINCETON .*
4ORCHIN - % + $ ,AFFERTY ! 0 $OBSON 6 * -C+ENZIE AND ! - +URIS
)NTRODUCED SPECIES AND THEIR MISSING PARASITES .ATURE
4SUTSUI . $ ! 6 3UAREZ $ ! (OLWAY AND 4 * #ASE 2EDUCED GENETIC VARIATION
AND THE SUCCESS OF AN INVASIVE SPECIES 0ROCEEDINGS OF THE .ATIONAL !CADEMY OF 3CIENCE
53!
7ARE $ - 0REDATION BY RAINBOW TROUT THE EFFECT OF EXPERIENCE *OURNAL OF THE
&ISHERIES 2ESEARCH "OARD OF #ANADA
7ILLIAMS $ & %XOTIC !NTS "IOLOGY )MPACT AND #ONTROL OF )NTRODUCED 3PECIES
7ESTVIEW 0RESS "OULDER #/
7ILLIAMSON - ( AND ! &ITTER 4HE CHARACTERS OF SUCCESSFUL INVADERS "IOLOGICAL
#ONSERVATION
#HAPTER ELEVEN
5SE OF BIOLOGICAL INVASIONS
AND THEIR CONTROL
TO STUDY THE DYNAMICS
OF INTERACTING POPULATIONS
& #OURCHAMP AND 3 #AUT
).42/$5#4)/.
/NE OF THE DIFlCULTIES OF CONSERVATION BIOLOGY IS THE GENERAL LACK OF EXPERIMENTAL
APPROACHES "ECAUSE IT IS OFTEN UNETHICAL OR SIMPLY BECAUSE THIS NEW DISCIPLINE
DEALS WITH SMALL ANDOR FRAGILE POPULATIONS EXPERIMENTS ON THOSE POPULATIONS ARE
NOT ALWAYS FEASIBLE !S A RESULT THE KNOWLEDGE ON POPULATION DYNAMICS WHEN
NOT DEALING WITH LABORATORY POPULATIONS OF CAGED INVERTEBRATES HAS OFTEN COME
FROM THEORETICAL STUDIES WITH NOTABLE EXCEPTIONS SUCH AS THOSE BASED ON OF SOME
POPULATIONS ISOLATED ON PARTICULAR ISLANDS EG #LUTTON
"ROCK AND #OULSON
'RENFELL ET AL (OWEVER ONE ASPECT THAT IS OFTEN LACKING FROM
THEORETICAL STUDIES AS WELL AS FROM NATURAL ISOLATED POPULATIONS IS THE INTERSPECIlC
DIMENSION IN THE ABOVE CASES IT IS RATHER EXCEPTIONAL TO TAKE INTO ACCOUNT MORE
THAN TWO INTERACTING POPULATIONS 9ET AS WE HOPE TO SHOW IN THIS CHAPTER DIRECT
AND INDIRECT hCOMPLEXv INTERSPECIlC RELATIONSHIPS MAY BE THE MAJOR ECOLOGICAL
FORCES IN SOME COMMUNITIES 4HEY CAN THUS BE CRUCIAL FOR APPLIED ECOLOGY AS WELL
-7 #ADOTTE ET AL (EDS) #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY n
3PRINGER 0RINTED IN THE .ETHERLANDS
& #OURCHAMP AND 3 #AUT
AS REPRESENT HEURISTIC TOOLS FOR STUDENTS AND INTELLECTUAL CANDIES FOR FUNCTIONAL
ECOLOGISTS
(OWEVER THERE IS AN ENORMOUS SET OF ECOLOGICAL EVENTS THAT CAN BE VIEWED AS
NATURAL LARGE
SCALE EXPERIMENTS BIOLOGICAL INVASIONS 3EVERAL ASPECTS MAKE BIO
LOGICAL INVASIONS AN INTERESTING TOOL FOR THE STUDY OF INTERSPECIlC INTERACTIONS THEY
ARE OF VARIOUS TYPES INVOLVE MANY DIFFERENT ORGANISMS AND HAPPEN IN CONTRASTED
ECOSYSTEMS "IOLOGICAL INVASIONS ARE OFTEN A VERY RICH SOURCE OF INFORMATION FOR THE
UNDERSTANDING OF ECOSYSTEM FUNCTIONING AS THEY ORIGINATE FROM INTRODUCTIONS THAT
ARE GENERALLY RELATIVELY WELL DOCUMENTED )N ADDITION IN MANY CASES THE INVADED
ECOSYSTEM IS AN ISLAND WITH ALL THE ADVANTAGES THAT INSULAR ECOSYSTEMS PROVIDE
FOR FUNDAMENTAL RESEARCH CLOSED ECOSYSTEMS WITH LIMITED SIZE AND OF RELATIVELY
SIMPLE AND NON
REDUNDANT TROPHIC WEBS )N THE SAME WAY THAT THE PHYSIOLOGY OF
AN INDIVIDUAL MAY BE BETTER UNDERSTOOD DURING ILLNESS THE DISFUNCTIONING OF AN
ECOSYSTEM MAY HELP GAIN KNOWLEDGE ABOUT ITS NORMAL FUNCTIONING )N THIS REGARD
THE CHANGES GENERATED BY THE SIMPLE MODIlCATIONS THAT ARE SPECIES INTRODUCTIONS
AND THEIR CONSECUTIVE SPREAD PROVIDE MANY DIFFERENT TYPES OF INFORMATION 4HUS
BIOLOGICAL INTRODUCTIONS REPRESENT SIMPLE EXPERIMENTS OF SPECIES ADDITION INTO A
NEW TROPHIC WEB 3UCH EXPERIMENTS CAN BENElT FROM CONTROLS AND REPLICAS IN THE
CASE OF ARCHIPELAGOES
3IMILARLY SPECIES DELETIONS CAN BE STUDIED IN LARGE
SCALE EXPERIMENTS THAT
ARE EVEN MORE ACCESSIBLE TO POPULATION OR COMMUNITY BIOLOGISTS AS THEY CAN BE
DESIGNED BY THEM THE CONTROL OR ERADICATION OF THE ALIEN SPECIES 7ITH THIS NEW
TOOL OF SPECIES ADDITION AND DELETION FROM THE COMPARATIVELY SIMPLE ISLAND ECOSYS
TEMS ONE MAY GAIN MORE KNOWLEDGE OF BASIC PROCESSES SUCH AS COLONIZATION DIS
PERSION SPATIAL SPREAD AS WELL AS THE DYNAMICS OF INTERACTING POPULATIONS 7E WILL
HERE FOCUS ON THIS LATTER ASPECT RESTRICTING OUR ANALYSIS ON THREE
AND FOUR
SPECIES
INTERACTIONS AND SHAMELESSLY BASING IT ON OUR PREVIOUS STUDIES
)N THIS CHAPTER WE WILL PRESENT A NUMBER OF MATHEMATICALLY SIMPLE MODELS
THAT DEPICT SOME hCOMPLEXv INTERSPECIlC RELATIONSHIPS WITH THE AIM OF SHOWING
HOW THE STUDY OF BIOLOGICAL INVASIONS AND THEIR CONTROL CAN BE USEFUL FOR THE STUDY
OF FUNDAMENTAL ECOLOGICAL PROCESSES THAT ARE MORE PROBLEMATICAL TO UNDERSTAND
IN OTHER CONTEXTS (ERE COMPLEX RELATIONSHIPS ARE DElNED AS INTERACTIONS WITHIN
TROPHIC WEBS THAT ENCOMPASS MORE THAN TWO POPULATIONS WITH POSSIBLE INDIRECT
PROCESSES AND THAT MAY NOT BE REALLY COMPLEX IN A BIOLOGICAL SENSE BUT THAT
ARE MORE DEMANDING TO STUDY ANALYTICALLY 7E ALSO USE @CONTROL IN A SOMEWHAT
LENIENT STYLE 4HIS TERM CAN HAVE TWO MEANINGS IT CAN BE A GENERAL TERM OF ACTION
AGAINST AN ALIEN SPECIES RANGING FROM SIMPLE REDUCTION UP TO ERADICATION AND IT
CAN MORE SPECIlCALLY MEAN REDUCTION OF THE POPULATION SIZE DOWN TO ACCEPTABLE
LEVELS IN ECOLOGICAL OR ECONOMIC TERMS 4HE LATTER IS CALLED @MITIGATION OR @REDUC
TION AND IS OPPOSED TO @ERADICATION )N THIS #HAPTER WE WILL USE @MITIGATION
FOR PARTIAL POPULATION REMOVAL @ERADICATION WHEN REMOVAL IS TOTAL AND @CON
TROL AS A GENERAL TERM 7E WILL ARTICULATE OUR PRESENTATION IN TWO PARTS THE lRST
PART DEPICTS SYSTEMS WHERE SPECIES ARE ADDED TO A TROPHIC WEB THE BIOLOGICAL
INVASIONS AND THE SECOND PART DEPICTS SYSTEMS WHERE SPECIES ARE REMOVED CON
$YNAMICS OF INTERACTING POPULATIONS
SERVATION PROGRAMS FOCUSING ON THE POSSIBLE ASSOCIATED INDIRECT PROCESSES IN
EACH CASE ! SECONDARY OBJECTIVE OF THIS CHAPTER IS TO CONVINCE THE READERS BE THEY
STUDENTS IN BIOLOGY OR CONSERVATION MANAGERS THAT MATHEMATICAL MODELING IS A
POWERFUL TOOL TO UNDERSTAND AND IN SOME CASE TO PREDICT ECOSYSTEM FUNCTIONING
AND REACTIONS 9ET FOR PEDAGOGIC PURPOSES WE WILL PRESENT OUR ANALYSES BASED
ON THE DESCRIPTION OF SEVERAL CONCRETE EXAMPLES WITH LITTLE OR NO EMPHASIS ON THE
TECHNICAL ASPECTS OF THE MATHEMATICAL MODELS 7E PROVIDE REFERENCES FOR MORE
DETAILS ABOUT THE MODELS AND THEIR ANALYSIS
!LL THE MODELS PRESENTED HERE ARE DETERMINISTIC COUPLED DIFFERENTIAL EQUATIONS
BASED ON CLASSICAL ,OTKA
6OLTERRA PREDATION OR COMPETITION MODELS %ACH POPULA
TION IS DESCRIBED BY A SIMPLE LOGISTIC EQUATION MODIlED TO TAKE INTO ACCOUNT ITS
RELATIONSHIP WITH THE OTHER POPULATIONS !LTHOUGH BIOLOGICALLY SIMPLE THE MOD
ELS PRESENTED HERE CAN SHOW A RELATIVELY HIGH MATHEMATICAL COMPLEXITY WHEN IT
COMES TO FOR EXAMPLE DETERMINING EQUILIBRIUM POINTS #ONlDENT THAT THE SIM
PLEST MODELS ARE THE MOST USEFUL 'INZBURG AND *ENSEN WE SYSTEMATICALLY
REFRAINED FROM UNDULY ADDING COMPLEXITY TO OUR EQUATIONS WHICH RESULTS IN A LACK
OF PREDICTIVE POWER 4HESE MODELS ARE THEREFORE NOT AIMED AT PROVIDING PRECISE
VALUES OF POPULATION TRENDS OR OF CONTROL MEASURES IN THE lELD NEITHER THE NATURE
OF THE MODELS NOR THE STATE OF CURRENT KNOWLEDGE IN THE lELD WOULD ALLOW USEFUL
QUANTITATIVE PREDICTIONS 2ATHER THE AIM OF THIS EXERCISE IS TO EMPHASIZE THE LINK
BETWEEN SPECIES THE IMPORTANCE OF INDIRECT INTERACTIONS AND THE UNEXPECTED OUT
COME OF CONTROL ACTIONS IF THEY ARE NOT THOROUGHLY TAKEN INTO ACCOUNT 7E BELIEVE
THAT THE QUALITATIVE INFORMATION PROVIDED BY OUR MECHANISTIC MODELS IS SUITABLE TO
OFFER THE INFORMATION WE SEEK IN THIS CONTEXT !LSO WHILE PARAMETERISING MODELS
TO REPRODUCE lELD RESULTS IS A VERY USEFUL WAY OF IDENTIFYING PLAUSIBLE MECHANISMS
OF TROPHIC INTERACTIONS IT DOES NOT IN ITSELF PROVIDE A DIRECT TEST OF THE IMPORTANCE
OF THOSE PLAUSIBLE MECHANISMS 4HE INFORMATION THEY GENERATE SHOULD ALWAYS BE
COMPLETED BY INFORMATION COMING FROM EMPIRICAL AND EXPERIMENTAL STUDIES )N
THIS #HAPTER WE WILL ONLY DEAL WITH THE MODELING PART
)N ORDER TO RENDER THE READING OF THIS CHAPTER LESS TEDIOUS WE WILL DESCRIBE IN
DETAIL THE PROCESS LEADING TO THE MODEL FOR THE lRST EXAMPLE ONLY AND WILL ONLY GIVE
THE MODEL FOR THE OTHER EXAMPLES !LTHOUGH SOME WILL DIFFER IN DETAILS ALL MODELS
ARE BASED ON SIMILAR PRINCIPLES !PART FROM THE MORE COMPLETE DESCRIPTION OF THE
lRST CASE ALL EXAMPLES WILL BE PRESENTED IN A SIMILAR WAY TO ALLOW EASY COMPARI
SONS BETWEEN CASES
& #OURCHAMP AND 3 #AUT
.!452!, %#/3934%- %80%2)-%.43 !$$)4)/.3 !.$
$%,%4)/.3 /& 30%#)%3
"IOLOGICAL INVASIONS AS A SPECIES ADDITION EXPERIMENT
4HE HYPERPREDATION PROCESS THREE SPECIES INTERACTIONS
A
4HE CASE
)NTRODUCED SPECIES ARE NOTORIOUS FOR THEIR DELETERIOUS IMPACT ON INVADED COM
MUNITIES AND THEIR DIRECT EFFECTS ON TROPHIC SYSTEMS SUCH AS DECREASE OF PREY OR
COMPETITOR POPULATIONS 7HILE THEY REPRESENT CATASTROPHIC EVENTS IN TERMS OF
BIODIVERSITY CONSERVATION THOSE EFFECTS ARE NOT OF MAJOR INTEREST TO THEORETICAL
ECOLOGY &OR THIS REASON WE WILL FOCUS ON LESS OBVIOUS EFFECTS STARTING WITH AN
EXAMPLE CONCERNING THE EXTINCTION OF AN ENDEMIC PARROT CAUSED BY THE INTRODUC
TION OF RABBITS TO AN INSULAR SYSTEM
4HE RABBIT IS ONE OF THE MOST DOCUMENTED INTRODUCED MAMMAL SPECIES OFTEN
ASSOCIATED WITH A DRAMATIC IMPACT ON ENDEMIC PLANT SPECIES 4O DATE THIS HER
BIVORE HAS BEEN INTRODUCED MOST OF THE TIME PURPOSEFULLY TO MORE THAN
ISLANDS &LUX AND &ULLAGAR 2ABBITS HAVE A HIGH ECOLOGICAL ADAPTABILITY
AND AS SUCH EASILY SUCCEED WHEN INTRODUCED IN TO ECOSYSTEMS WHERE INDIGENOUS
GRAZERS ARE LESS NUMEROUS AND COMPETITIVE &LUX 4HE VERY RAPID INCREASE
OF THEIR POPULATIONS CAN LEAD TO A DRAMATIC QUANTITATIVE AND QUALITATIVE IMPOVER
ISHMENT OF THE VEGETATION #HAPUIS ET AL 3ELKIRK ET AL RESULTING IN
DRAMATIC DENUDATION OF THE SOIL 3COTT AND HAVE AN IMPACT ON ANIMAL
SPECIES WHICH DEPEND ON THE VEGETATION 'ILLHAM
%FFECTS OF RABBITS ON INDIGENOUS VERTEBRATE SPECIES CAN ALSO BE MORE COMPLEX
4HESE MAMMALS ARE PREYED UPON BY OTHER INTRODUCED VERTEBRATES IN PARTICULAR BY
FERAL DOMESTIC CATS #ATS FOR EXAMPLE ARE OPPORTUNISTIC PREDATORS WHICH SWITCH
PREY ACCORDING TO RELATIVE SPATIAL ANDOR TEMPORAL AVAILABILITY &ITZGERALD
7HEN RABBITS ARE ABUNDANT DOMESTIC CATS ARE KNOWN TO PREY LARGELY UPON THEM
(OWEVER RABBITS CAN CONSTITUTE A SMALLER PART OF THE CAT DIET WHEN BIRDS REPTILES
OR OTHER MAMMALS ARE RELATIVELY MORE ABUNDANT )N SEVERAL SUB
!NTARCTIC ISLANDS
RABBITS ARE ONLY A SECONDARY PREY ITEM IN MONTHS WHEN SEABIRDS ARE PRESENT BUT
APPEAR TO ENABLE CATS TO SUBSIST OVER WINTER WHEN SEABIRDS ARE ABSENT #HAPUIS
A B ! SIMILAR EFFECT IS DOCUMENTED IN THE SPATIAL DIMENSION RABBITS OFTEN
ENABLE CATS TO REACH REMOTE COLONIES OR POPULATIONS OF INDIGENOUS PREY IN ISLANDS
WITH HETEROGENEOUS INDIGENOUS PREY DISTRIBUTION "ROTHERS AND #OPSON
)N THESE CASES THE PRESENCE OF RABBITS HAS AN INDIRECT EFFECT ON OTHER PREY SPECIES
USED BY INTRODUCED CATS
0REDATION BY CATS INTRODUCED TO -ACQUARIE )SLAND CAUSED THE DECLINE OF BURROW
NESTING PETRELS "ROTHERS AND THE EXTINCTION OF AN ENDEMIC PARAKEET AND
A BANDED RAIL 4AYLOR #ATS WERE INTRODUCED TO THE ISLAND YEARS BEFORE
THE INTRODUCTION OF RABBITS HOWEVER THE CAT DRIVEN EXTINCTION OF BIRDS DATES BACK
$YNAMICS OF INTERACTING POPULATIONS
TO JUST YEARS FOLLOWING THE INTRODUCTION OF RABBITS 4AYLOR 2ABBITS WERE
NOT OBSERVED AS HAVING ANY DIRECT EFFECTS ON THE LAND BIRDS )N FACT IT IS BELIEVED
THAT THE RABBIT POPULATION ALLOWED A SIGNIlCANT INCREASE IN THE CAT POPULATION
RESULTING IN AN INCREASED PREDATION PRESSURE ON THE LAND BIRD SPECIES 4HIS PRO
CESS RELATED TO THE MORE GENERAL hAPPARENT COMPETITIONv EG !BRAMS
!BRAMS ET AL (OLT HAS BEEN TERMED HYPERPREDATION #OURCHAMP
ET AL 3MITH AND 1UIN
)T IS GENERALLY ASSUMED THAT LIFE HISTORY TRAITS AND BEHAVIOR OF THE INTRODUCED
PREY MAKE IT RESISTANT TO HIGH LEVELS OF PREDATION PRESSURE ! HIGHER REPRODUCTIVE
RATE HIGH DENSITY AND EFlCIENT ANTI PREDATOR RESPONSES WHICH ARE OFTEN LACKING IN
THE INDIGENOUS SPECIES EXHIBITED BY THE INTRODUCED PREY COULD ENABLE AN INCREASE
IN THE PREDATOR POPULATION WITHOUT A LARGE DECREASE IN THE INTRODUCED PREY POPU
LATION &URTHERMORE THE LACK OF SERIOUS COMPETITORS AND THE RELATIVELY FEW PARA
SITES TYPICALLY FOUND IN INTRODUCED POPULATIONS CAN ALSO INCREASE THEIR POTENTIAL FOR
DRAMATIC POPULATION GROWTH THE ENEMY RELEASE HYPOTHESIS +EANE AND #RAWLEY
4HESE FEATURES IMPLY THE ABILITY TO SUSTAIN HIGH PREDATION PRESSURE AS CATS
ARE SUPPOSED TO REMOVE ONLY THE INDIVIDUALS WITH LOW SURVIVAL DISPERSING YOUNG
SICK AND DEAD 3MITH AND 1UIN 4HE RESULTING INCREASED POPULATION OF
PREDATORS CANNOT BE SUSTAINED BY THE INDIGENOUS PREY SPECIES WHICH COMPARED
WITH THE INTRODUCED PREY SPECIES HAS INFERIOR LESS WELL ADAPTED REPRODUCTIVE AND
ANTI PREDATOR CHARACTERISTICS 4HE CONJUNCTION OF A LOW ADAPTATION TO PREDATION
AND AN ARTIlCIALLY HIGH PREDATION PRESSURE CAN LEAD TO A DRAMATIC DECREASE IN AN
INDIGENOUS PREY POPULATION UP TO TOTAL EXTIRPATION
B 4HE MODEL
7E WILL THUS PRESENT A SIMPLE MODEL OF HYPERPREDATION TO ILLUSTRATE HOW THE
EXTINCTION OF THE BIRD POPULATION ON -ACQUARIE )SLAND CAN BE EXPLAINED BY AN
INDIRECT EFFECT OF THE INTRODUCED RABBITS &OR HEURISTIC PURPOSES WE lRST PRESENT A
TWO
SPECIES MODEL AND THEN MODIFY IT INTO A THREE
SPECIES MODEL 7E HOPE THAT
THIS WILL HELP THE UNDERSTANDING OF THIS MODEL AND OF THE OTHER SETS OF EQUATIONS
IN THIS CHAPTER WHICH ARE ALL BASED ON THE SAME PRINCIPLE 7E USE THE EXAMPLE
OF BIRD LOCAL PREY RABBIT INTRODUCED PREY AND CAT INTRODUCED PREDATOR AND
WILL REFER TO THESE SPECIES FOR THE SAKE OF SIMPLICITY BUT OTHER SPECIES CAN PRESENT
SIMILAR RELATIONSHIPS
4HE BIRD
CAT MODEL CAN BE GIVEN THE FOLLOWING FORM
D" "
RB" n n B#"
D" +B
D#
hB B"# n i#
DT
& #OURCHAMP AND 3 #AUT
WHERE THE NUMBER OF INDIVIDUALS AT TIME T IN THE BIRD RABBIT AND CAT POPULA
TIONS ARE " 2 AND # RESPECTIVELY 4HE RABBIT
CAT MODEL IS THE SAME 4HE INTRINSIC
GROWTH RATES OF THE BIRD AND THE RABBIT POPULATIONS ARE RB AND RR RESPECTIVELY
4HE PREDATION RATE IS +B ON THE BIRD POPULATION AND +R ON THE RABBIT POPULATION
4HE CARRYING CAPACITY OF THE ENVIRONMENT FOR THE BIRD POPULATION IS +B AND THE
CARRYING CAPACITY OF THE ENVIRONMENT FOR THE RABBIT IS +R 4HE RATE AT WHICH
EATEN PREY ARE TURNED INTO NEW PREDATORS IS hB FOR BIRDS AND hR FOR RABBITS AND i
IS THE PREDATOR MORTALITY RATE )N A MORE GENERAL MANNER THE SAME PARAMETERS
WILL BE USED FOR THE NEXT MODELS WITH INDEXES CORRESPONDING TO THE lRST LETTER OF
THE SPECIES CONSIDERED
)F TWO PREY SPECIES ARE TO BE CONSIDERED SIMULTANEOUSLY THE FORMULATION OF THE
PREDATION RATES AND OF THE GROWTH RATE OF THE PREDATOR MUST BE CHANGED ACCORD
INGLY INSTEAD OF +B# AND +R# THE PREDATION RATES ARE GIVEN THE FORM
" 2
+B# AND + R#
" 2 " 2
FOR THE BIRD AND THE RABBIT PREDATION RATES RESPECTIVELY SO THAT THE PREDATION
RATE IS STILL A FUNCTION OF THE AVAILABILITY OF THE PREY BUT VARIES WITH RELATIVE PREY
PROPORTIONS 4HE ADAPTATION OF THE INTRODUCED PREY IN TERMS OF AN ANTI
PREDATOR
BEHAVIORAL RESPONSE IS GIVEN BY A PREFERENCE OF THE PREDATOR FOR THE INDIGENOUS
PREY WHICH IS MORE EASILY DETECTED ANDOR CAUGHT OVER THE INTRODUCED PREY
4HIS PREFERENCE IS A RATIO _ WITH A SIMPLE BIOLOGICAL MEANING GIVEN EQUAL AVAIL
ABILITY THE PREDATOR WILL PREY UPON THE INDIGENOUS PREY _ TIMES MORE OFTEN THAN
ON THE INTRODUCED PREY 7E ASSUME THAT _ * AND THAT ONE RABBIT AND ONE BIRD
PREY ITEMS ARE ENERGETICALLY EQUALLY VALUABLE TO THE CAT 4HE PREDATION TERMS ARE
NOW GIVEN BY
_" 2
+B# AND + R#
_" 2 _" 2
ON THE INDIGENOUS AND INTRODUCED PREY RESPECTIVELY 4HIS CHANGE IS REmECTED IN
A SIMILAR WAY IN THE PREDATOR GROWTH RATE
_" 2
hB" # h 2 R#
B R
_" 2 _" 2
IT DEPENDS ON BOTH THE NUMBERS AND THE PROPORTION OF PREY 7E HAVE NOW THE FOL
LOWING SYSTEM
$YNAMICS OF INTERACTING POPULATIONS
D" " _"
RB" n n B#"
DT +B _" 2
D2 2 2
RR2 n n #2
DT +R _" 2 R
D# hB B _" hR R 2 #
n i#
DT _" 2
.OTE THAT SIMILAR FORMULATIONS OF THE MODEL COULD BE USED AND THAT THE AFORE
MENTIONED CHOICES ARE ARBITRARY )N PARTICULAR WE USED A PREDATION TERM PROPOR
TIONAL TO THE NUMBER OF PREY 7E DID SO TO KEEP THE FORMULATION OF THE ORIGINAL
PAPER #OURCHAMP ET AL BUT ALTERNATIVE MODELS CAN REPRODUCE THE HYPER
PREDATION PROCESS !LSO LIKE FOR THE REST OF THE MODELS PRESENTED IN THIS CHAPTER
WE DO NOT TAKE INTO ACCOUNT THE FURTHER RISKS ENCOUNTERED BY POPULATIONS WHEN AT
SMALL SIZES ENVIRONMENTAL AND DEMOGRAPHIC STOCHASTICITY !LLEE EFFECTS ETC -OST
OF THOSE WOULD ANYWAY ONLY STRENGTHEN OUR POINT 4HE CLASSICAL COMPARTMENTAL
REPRESENTATION OF THE MODEL IS PRESENTED IN &IG ! THE CORRESPONDING SET OF EQUA
TIONS IS SHOWN IN &IG " AND A SELECTED REPRESENTATION OF THE POPULATION TRENDS
WITH TIME IS GIVEN IN &IG # 4HE OTHER EXAMPLES WILL BE ILLUSTRATED WITH lGURES
FOLLOWING THE SAME FORMAT
4HE STUDY OF THIS SET OF EQUATIONS BOTH ANALYTICALLY AND NUMERICALLY SEE
#OURCHAMP ET AL SHOWS THAT THE INDIRECT EFFECT OF THE INTRODUCED PREY
MAY BE VERY IMPORTANT )NDEED ACCORDING TO THE VALUES OF THE PARAMETERS THE
INCREASE OF THE PREDATOR POPULATION TRIGGERED BY THE PRESENCE OF THE INTRODUCED
PREY CAN DRIVE THE INDIGENOUS PREY TO VERY LOW NUMBERS AND POTENTIALLY TO EXTINC
TION 4HE EFFECT OF HYPERPREDATION IS THE STRONGEST FOR SPECIES WITH LOW INTRINSIC
GROWTH RATE AND LOW ENVIRONMENTAL CARRYING CAPACITY 4HIS MODEL ALSO ILLUSTRATES
THAT THE HYPERPREDATION PROCESS MAY BE DUE TO A COMBINATION OF WELL
ADAPTED LIFE
HISTORY TRAITS AND EFlCIENT BEHAVIORAL RESPONSE OF PREY BUT THAT THE hBETTER
ADAPTEDv
BEHAVIORAL RESPONSE MAY HAVE MORE IMPORTANCE THAN hBETTER
ADAPTEDv LIFE HISTORY
TRAITS AT LEAST FOR THE CASES CONSIDERED #OURCHAMP ET AL 4HUS A PREY SPE
CIES INTRODUCED INTO AN ENVIRONMENT IN WHICH A PREDATOR HAS ALSO BEEN INTRODUCED
IS LIKELY TO ALLOW SO HIGH AN INCREASE OF THIS PREDATOR THAT LOCAL PREY LESS ADAPTED TO
HIGH LEVELS OF PREDATION COULD SUFFER A POPULATION DECLINE AND POSSIBLY EVEN EXTINC
TION 3UCH A PROCESS HAS CONSEQUENCES WHEN IT COMES TO MANAGEMENT ACTIONS
C #ONSERVATION CONSEQUENCES
(ISTORICALLY CONSERVATION PROGRAMS FOR MANY ISLANDS HAVE PROCESSED CASE BY
CASE 5NTIL RECENTLY INTRODUCED SPECIES WERE ALWAYS CONSIDERED SEPARATELY WITH
AT BEST SEPARATE PROGRAMS FOR EACH SPECIES AND TIMINGS DEPENDING MOSTLY ON
FUNDING AND LOGISTICS OR MORE FREQUENTLY ONE SINGLE PROGRAM FOR THE VISIBLY MOST
& #OURCHAMP AND 3 #AUT
&IG 2EPRESENTATION OF THE HYPERPREDATION MODEL WITH THE INTRODUCED PREY INDI
GENOUS PREY AND INTRODUCED PREDATOR BEING ILLUSTRATED BY RABBITS 2 BIRDS " AND CATS
# RESPECTIVELY ! COMPARTMENTAL REPRESENTATION AND ILLUSTRATION OF THE ISLAND FROM
WHERE THE EXAMPLE IS TAKEN HERE -ACQUARIE )SLAND OFF .EW :EALAND %ACH BOX REPRESENTS
A POPULATION AND THE ARROWS REPRESENT mUXES BETWEEN THEM 4HE LARGE CURVED ARROW
REPRESENTS AN INDIRECT EFFECT %ACH SPECIES IS ILLUSTRATED IN A COLOR THAT IS ALSO USED FOR THE
SET OF EQUATIONS " AND FOR THE DRAWINGS THAT REPRESENT THE POPULATION DYNAMICS OF THE
INTERACTING SPECIES AFTER INTRODUCTION OF A SPECIES # OR CONTROL OF AN INTRODUCED SPECIES
$ )N THIS EXAMPLE FOLLOWING THE CAT INTRODUCTION RED ARROW THE INCREASE OF THE CAT
POPULATION ALLOWED BY THE LARGE POPULATION OF RABBITS LEADS TO BIRD EXTINCTION # &OLLOWING
CONTROL GREEN ARROW IN $ THE BIRD POPULATION ONLY PARTIALLY RECOVERS IF ONLY THE CAT IS
CONTROLLED TOP PANEL OF $ (OWEVER THE SAME CAT CONTROL LEVEL LEADS TO CAT ERADICATION AND
FULL RECOVERY OF BIRDS IF BOTH THE RABBITS AND THE CAT ARE CONTROLLED BOTTOM PANEL OF $
$YNAMICS OF INTERACTING POPULATIONS
DEVASTATING SPECIES !S A RESULT WHEN A CONSERVATION PROGRAM INVOLVED AN ISLAND
SUCH AS THE ONE WE JUST MENTIONED WITH A HYPERPREDATION PROCESS TAKING PLACE
THE KEY ROLE OF THE INTRODUCED PREY WAS NOT SYSTEMATICALLY OBVIOUS 0REDATORS ARE
OFTEN PERCEIVED AS HAVING THE MOST IMPORTANT DELETERIOUS EFFECTS ON INVADED ECO
SYSTEMS AND CONSEQUENTLY CONTROL PROGRAMS WERE MORE OFTEN DIRECTED AT THEM
SOMETIMES NEGLECTING THE INTRODUCED PREY 9ET WE HAVE JUST SEEN THAT IN THE PRES
ENCE OF INTRODUCED PREDATORS INTRODUCED PREY COULD HAVE AN INDIRECT IMPACT ON
INDIGENOUS PREY "ASING OUR EFFORTS ON THE MODEL PRESENTED IN THE PREVIOUS SECTION
OF THIS CHAPTER WE STUDIED THE RELATIVE EFlCIENCY OF CONTROL PROGRAMS AIMING EITHER
AT THE PREDATOR ONLY OR AT THE INTRODUCED PREY AND PREDATOR SIMULTANEOUSLY 4HE
MODEL ON WHICH WE BASED THIS HAS BEEN PUBLISHED IN #OURCHAMP ET AL
AND THIS MODEL IS ONLY SLIGHTLY DIFFERENT FROM THE ASSOCIATED MODEL WITHOUT CONTROL
#OURCHAMP ET AL 7E WILL NOT REPRODUCE THE MODEL HERE AS THE INTEREST
LIES NOT IN THE DETAILS OF THE EQUATIONS 4HE ONLY IMPORTANT POINT IS THE ADDITION OF
A CONTROL EFFORT ON EITHER THE ALIEN PREY OR THE ALIEN PREDATOR OR BOTH 4HIS CONTROL
IS ADDED TO THE CORRESPONDING EQUATION BY A SIMPLE LINEAR TERM 7E EMPHASIZE THE
FACT THAT IN THE MODEL THE INTRODUCED PREY RABBIT STILL HAS NO DIRECT EFFECT ON THE
LOCAL PREY BIRD
3OME POSSIBLE POPULATION TRENDS OF THE SYSTEM IN PRESENCE OF CONTROL ARE SHOWN
IN &IG $ 4HE STUDY OF THIS MODEL SHOWS THAT CONTROL OF RABBITS CAN FACILITATE
THE ERADICATION OF CATS )NDEED WHEN NO CONTROL IS UNDERTAKEN THE CAT POPULA
TION STAYS LARGE MAINLY BECAUSE OF THE PRESENCE OF RABBITS AND CAN ELIMINATE THE
BIRDS IN THE LONG TERM 7HEN CATS ONLY ARE CONTROLLED THE PRESENCE OF RABBITS CAN
PRECLUDE CAT ERADICATION AND THE BIRD POPULATION RECOVERY IS ONLY PARTIAL )N CON
TRAST FOR THE SAME CAT CONTROL EFFORT ERADICATION OF RABBITS ALLOWS ERADICATION OF
CATS AND TOTAL RECOVERY OF BIRDS !CTUALLY IF THE CONTROL OF INTRODUCED PREY IS NOT
SUFlCIENT THE INDIGENOUS PREY WILL BE DESTROYED EVEN IF THE PREDATOR POPULATION
IS BEING CONTROLLED
/BVIOUSLY WE ARGUE HERE THAT EVEN IN ABSENCE OF VISIBLE DIRECT EFFECT INTRO
DUCED PREY SHOULD BE CONTROLLED WHEN A PREDATOR HAS BEEN INTRODUCED IN ORDER
TO PREVENT AN ARTIlCIAL PREDATOR POPULATION INCREASE )N ADDITION REMOVING AN
INTRODUCED PREDATOR POPULATION WITHOUT CONTROLLING THE INTRODUCED PREY MAY BE
DIFlCULT TO ACHIEVE SINCE THEY CONSTITUTE A CONSTANT SOURCE OF FOOD TO THE PREDATOR
!LSO IT WOULD NOT BE AN APPROPRIATE SOLUTION BECAUSE REMOVING THE PREDA
TION PRESSURE WOULD INCREASE THE DIFlCULTIES OF LATER COPING WITH INTRODUCED PREY
WHICH ARE OFTEN CHARACTERIZED BY HIGH REPRODUCTIVE RATES /N THE OTHER HAND
CONTROLLING ONLY THE INTRODUCED PREY IS UNSATISFACTORY IN THE LONG TERM BECAUSE
PREDATORS COULD REPORT HIGH PREDATION PRESSURE ON THE INDIGENOUS PREY #OMBINED
CONTROL OF BOTH SPECIES SEEMS HERE TO BE THE BEST RESTORATION STRATEGY )N ADDITION
STARTING BOTH CONTROL PROGRAMS TOGETHER WOULD ALSO RESULT IN ADVANTAGES DUE
TO SYNERGETIC EFFECTS COSTS MAY BE REDUCED IF COSTS RELATED TO TRANSPORTATION OR
HUNTING AND TRAPPING CAN BE SHARED BY THE TWO PROGRAMS AND EFlCIENCY MIGHT BE
INCREASED EG THROUGH THE ADDITIVE EFFECTS OF PRIMARY AND SECONDARY POISONING OF
PREDATORS &LUX 2AMMELL ET AL 2OBERTSON ET AL
& #OURCHAMP AND 3 #AUT
4O CONCLUDE IT IS WORTH REITERATING THAT THE HIGHER EFlCIENCY OF DUAL CONTROL IS
NOT DUE TO DIRECT EFFECTS OF RABBITS ON BIRDS HABITAT DESTRUCTION AND COMPETITION FOR
FOOD AND SHELTER SINCE THEY ARE NOT TAKEN INTO ACCOUNT HERE .OR IS THE PREDICTED
SUCCESS OF DUAL CONTROL DUE TO THE PREFERENCE OF THE PREDATOR SINCE THIS PREFER
ENCE IS SET IN FAVOR OF THE INDIGENOUS PREY IN THE MODEL 4HIS SUCCESS IS DUE TO THE
ADDRESSING OF THE HYPERPREDATION PROCESS
4HE HYPERPREDATION PROCESS FOUR SPECIES INTERACTIONS
A 4HE CASE
4HE CAT IS A WELL
KNOWN PREDATOR OF BOTH INSULAR BIRDS AND SMALL INTRODUCED MAM
MALS SO THE EXAMPLE ABOVE SHOULD BE RELATIVELY EASY TO SPOT 7HENEVER A LOCAL
POPULATION IS THREATENED BY AN INTRODUCED PREDATOR A DIET STUDY OF THE PREDATOR IN
QUESTION SHOULD BE CONDUCTED IN ORDER TO ASSESS THE IMPORTANCE OF THE IMPACT ON
THE LOCAL POPULATION BUT ALSO POTENTIAL HYPERPREDATION PROCESSES (OWEVER THERE
ARE CASES WHERE THIS STRATEGY IS NOT OBVIOUS BECAUSE THE CAUSE OF A PREY POPULA
TION DECLINE MAY NOT BE SPOTTED AS EASILY !N INTERESTING ILLUSTRATION OF THIS IS THE
SEVERE DECLINE OF THE INSULAR FOX ON THE #HANNEL )SLANDS IN THE S
4HE #HANNEL )SLANDS GROUP IS MADE UP OF EIGHT SMALL ISLANDS OFF THE #ALIFORNIAN
COAST 53! 4HE ISLAND GREY FOX 5ROCYON LITTORALIS IS A SMALL CARNIVORE THAT
ARRIVED ON THE lRST ISLAND YEARS AGO 4HE FOX NOW INHABITS THE SIX LARGEST
ISLANDS AND HAS EVOLVED ON EACH OF THESE IN ISOLATION RESULTING IN SIX POPULATIONS
REPRESENTING SIX DIFFERENT SUBSPECIES ALL ENDEMIC TO THESE ISLANDS /N THE THREE
NORTHERN ISLANDS THE INSULAR FOXES AND THEIR MAIN COMPETITOR THE ENDEMIC SPOT
TED SKUNKS 3PILOGALE GRACILIS AMPHIALA WERE THE TWO TERRESTRIAL TOP
PREDATORS OF
THE #HANNEL )SLANDS )N THE EARLY S A STUDY CONDUCTED ON THE HOME RANGE
OF THE FOXES WITNESSED A CONSIDERABLE DECLINE IN THE THREE NORTHERN ISLAND POPU
LATIONS 2OEMER 4HIS SEVERE DECLINE HAD NO OBVIOUS CAUSE AT lRST AND
MANY CLASSICAL ECOLOGICAL FORCES WERE INVESTIGATED LACK OF SUFlCIENT RESOURCES
COMPETITION WITH THE SPOTTED SKUNK AND DISEASES 0REDATION WAS ALSO INVESTIGATED
ALTHOUGH THE INSULAR FOX WAS THE TOP TERRESTRIAL PREDATOR OF THESE ECOSYSTEMS
3USPICION THAT FOXES WERE KILLED BY GOLDEN EAGLES LED TO A NEW EFFORT OF RESEARCH IN
THIS DIRECTION 'OLDEN EAGLES HAVE HISTORICALLY BEEN SEEN VISITING THE ISLANDS BUT
THEY NEVER STAYED LONG ENOUGH TO CONSTITUTE A THREAT TO THE LOCAL PREY ! STUDY
COMBINING METABOLIC AND ENERGETIC APPROACHES WITH POPULATION MODELING DEM
ONSTRATED THAT THE LOCAL PREY WERE TOO FEW TO ALLOW A PAIR OF DISPERSING EAGLES TO
BREED ON AND COLONIZE THE ISLANDS 9ET IT BECAME OBVIOUS THAT EAGLES WERE KILLING
FOXES AS WELL AS SPOTTED SKUNKS /N ONE OF THESE ISLANDS 3ANTA #RUZ lELD WORKERS
EVENTUALLY DISCOVERED A GOLDEN EAGLE NEST IN WHICH FOX REMAINS ATTESTED FOR THE
SUSPECTED PREDATION ON THIS SPECIES "UT THE PROBLEM REMAINED (OW WOULD THE
EAGLE THREATEN FOX SURVIVAL THROUGH PREDATION IF THERE WAS NOT ENOUGH LOCAL PREY
ON THE ISLAND TO ALLOW THE CONTINUOUS PRESENCE OF THE EAGLES 4HE DISCOVERY OF THE
NEST PROVIDED THE ANSWER REMAINS OF PIGLETS WERE ALSO FOUND IN THE NEST &ERAL
$YNAMICS OF INTERACTING POPULATIONS
PIGS 3US SCROFA WERE INTRODUCED ON TO THE THREE NORTHERN ISLANDS WHERE THE FOXES
ARE DECLINING THEY WERE ALSO PRESENT ON TWO OF THE SOUTHERN ISLANDS BUT HAVE
ALREADY BEEN OR ARE ALMOST ERADICATED THERE )N ADDITION TO THE DIRECT DAMAGES
THAT INTRODUCED PIGS ARE KNOWN TO CAUSE TO THE mORA AND FAUNA THEY INVADE (ONE
THIS ALIEN SPECIES ALSO THREATENED SOME LOCAL SPECIES THROUGH AN INDIRECT
PROCESS "Y PRODUCING PIGLETS ALL YEAR ROUND THEY PROVIDED VISITING EAGLES WITH
ENOUGH RESOURCES FOR THEM TO COLONIZE THE ISLANDS %AGLES ALSO IRREGULARLY DEPRE
DATED OTHER LOCAL PREY SUCH AS FOXES OR SKUNKS (OWEVER EVEN THIS LOW PREDATION
RATE ON A SPECIES THAT IS ILL
ADAPTED TO AVIAN PREDATION BEHAVIORALLY AS WELL AS AT
THE POPULATION LEVEL WAS SUFlCIENT TO DRIVE THE FOX POPULATION TOWARDS EXTINC
TION 4HE DECLINE WAS ALL THE MORE DRAMATIC THAT THE BREEDING EAGLE POPULATION
RAPIDLY GREW IN NUMBERS THEREBY INCREASING THE PRESSURE ON THE FOX POPULATION
)N THE MEAN TIME THE NOCTURNAL SKUNKS BENElTED FROM THE ARRIVAL OF THE EAGLE
BECAUSE THEY BENElTED FROM THE RELEASE OF COMPETITION PRESSURE FROM THE DECLIN
ING FOX POPULATIONS WHILE ALSO BEING KILLED LESS OFTEN THAN THEM 4HE HYPOTHESIS
WAS THUS THAT THE ARRIVAL OF PIGS HAD ALLOWED VISITING EAGLES TO STAY AND BREED
AND THEREBY THEY ATTRACTED A SHARED PREDATOR TO INSULAR PREY 0IGS ARE WELL ADAPTED
TO PREDATION THEY PRODUCE NUMEROUS PIGLETS THAT CAN ESCAPE EAGLE PREDATION
ONCE THEY REACH THREE MONTHS OF AGE 4HEREFORE THERE WERE LESS CONSEQUENCES
FOR THE PIG POPULATION THAN THE LOCAL PREY DUE TO THE ARRIVAL OF THE EAGLE 2OEMER
ET AL
B
4HE MODEL
4O TEST THIS HYPOTHESIS A MODEL OF THE POPULATION DYNAMICS OF THE INTERACTING
SPECIES WAS CONSTRUCTED AND PARAMETERIZED WITH DATA OBTAINED FROM THE lELD 4HE
MODEL WAS BASED ON A SIMPLE COMBINATION OF TWO CLASSICAL ,OTKA
6OLTERRA MOD
ELS ONE OF COMPETITION AND ONE OF PREDATION 4HE SKUNK AND THE FOX POPULATION
DYNAMICS WERE DESCRIBED BY A COMPETITION MODEL THE PIG AND THE EAGLE POPULA
TION DYNAMICS WERE DESCRIBED BY A PREDATION MODEL AND A PREDATION TERM OF THE
EAGLE WAS ADDED ON BOTH FOX AND SKUNK POPULATIONS 5SING A CORRECTION TERM FOR
PROPORTIONS AND PREFERENCE COEFlCIENTS q AND m RESPECTIVELY AS IN THE PREVIOUS
EXAMPLE WE END UP WITH A SYSTEM OF FOUR EQUATIONS ONE PREDATOR AND ITS THREE
PREY TWO OF WHICH ARE COMPETITORS 4HE SYSTEM AND ILLUSTRATIONS OF POPULATION
TRENDS ARE SHOWN IN &IG -ORE DETAILS CAN BE FOUND IN 2OEMER ET AL
3IMPLE SIMULATIONS SHOW THAT IN ABSENCE OF THE PIGS IF THE SYSTEM IS RUN WITH AN
INITIAL NUMBER OF PIGS SET AT ZERO ANY INTRODUCTION OF EAGLES HOWEVER LARGE WILL
EVENTUALLY LEAD TO COLONIZATION FAILURE AND FOX POPULATION PERSISTENCE (OWEVER
WHEN PIGS ARE PRESENT A SINGLE PAIR OF EAGLES WILL BE ABLE TO COLONIZE THE ISLAND AND
BUILD A POPULATION THAT IS SO LARGE THAT FOXES WILL GO EXTINCT WHILE PIGS WILL REMAIN
AT MODERATE DENSITIES
)T IS ALSO INTERESTING TO NOTE THAT THE DECLINE IN FOX NUMBERS CONSECUTIVE TO
THE HYPERPREDATION PROCESS TRIGGERED BY THE INTRODUCTION OF PIGS IS CONCOMITANT
WITH AN INCREASE OF THE ENDEMIC SKUNK )N FACT THE ARRIVAL OF EAGLES REVERSED THE
& #OURCHAMP AND 3 #AUT
&IG 2EPRESENTATION OF THE HYPERPREDATION MODEL EXAMPLE WITH FOUR SPECIES FOX
& BLUE SKUNK 3 GREEN PIG 0 YELLOW AND EAGLE % RED !S IN &IG ! IS THE
COMPARTMENTAL REPRESENTATION " IS THE RESULTING SET OF EQUATIONS # IS THE ILLUSTRATION
OF POPULATIONS TRENDS GIVEN BY " FOLLOWING PIG INTRODUCTION AND EAGLE COLONIZATION AND
$ IS THE POPULATIONS TRENDS FOLLOWING PIG ANDOR EAGLE CONTROL 4HE PARAMETERS ARE THE
SAME THAN PREVIOUSLY WITH q AND m BEING THE PREFERENCE PARAMETERS OF THE EAGLE FOR THE
FOX AND THE SKUNK OVER THE PIG RESPECTIVELY SAME AS _ IN &IG 4HE CONTROL STRATEGY $
IS REPRESENTED IN THREE DIMENSIONS 4O HELP VISUALIZE THE $ EFFECT THE COLORS DO NOT REFER
TO SPECIES BUT TO DIFFERENT POPULATION SIZES 4HIS GRAPH SHOWS THAT THE POPULATION SIZE OF
FOXES IS PROPORTIONAL TO EAGLE CONTROL BUT INVERSELY PROPORTIONAL TO PIG CONTROL !S A RESULT
IF EAGLES ARE NOT CONTROLLED SIMULTANEOUSLY FOXES WILL DECLINE FOLLOWING PIG CONTROL ONLY
)N ABSENCE OF SIGNIlCANT EAGLE MITIGATION HIGH LEVELS OF PIG MITIGATION CAN RESULT IN FOX
EXTINCTION DARK AREA
$YNAMICS OF INTERACTING POPULATIONS
COMPETITIVE OUTCOME BETWEEN THE TWO TOP TERRESTRIAL PREDATORS SHIFTING FORCES
FROM DIRECT COMPETITION IN FAVOR OF THE FOX TO APPARENT COMPETITION IN FAVOR OF
THE SKUNK 4HIS SECOND APPARENT COMPETITION PROCESS EMBEDDED IN THE lRST ONE
RENDERS ANY CONSERVATION STRATEGY AT THE LEAST COMPLICATED AS THE INSULAR SPOTTED
SKUNK IS ENDEMIC FROM THE NORTHERN #HANNEL )SLANDS AND IS CURRENTLY BENElTING
FROM THE FOX DECLINE
C #ONSERVATION CONSEQUENCES
&OXES ARE NOW EXTINCT IN THE WILD ON TWO OF THE THREE NORTHERN ISLANDS WITH
THE POPULATION ON THE THIRD ISLAND 3ANTA #RUZ ON THE VERGE OF EXTINCTION
INDIVIDUALS IN THE WILD AT THE END OF /UR MODELING EXERCISE SUGGESTS THAT
THE EXTINCTION OF TWO POPULATIONS OF THE TOP PREDATOR IN TWO INSULAR ECOSYSTEMS
IS LIKELY DUE TO AN INDIRECT PROCESS A PROCESS WHERE AN INTRODUCED PREY ATTRACTED
A SHARED PREDATOR AND ELIMINATED AN ENDEMIC PREY THROUGH APPARENT COMPETITION
ONLY
/BVIOUSLY THE SOLUTION TO THIS PROBLEM LIES WITH THE PIGS 4HE MOST EVIDENT PLAN
OF ACTION WAS TO REMOVE THE PIGS FROM THE NORTHERN ISLANDS 4HIS WOULD HAVE THE
DOUBLE ADVANTAGE OF STOPPING THEIR DIRECT DELETERIOUS EFFECTS ON THE LOCAL mORA AND
FAUNA AS WELL AS ELIMINATING THE PREY BASIS FOR THE EAGLES LEADING THEM WITH LITTLE
MORE CHOICE THAN STARVATION OR EMIGRATION )N FACT SEVERAL CONSERVATION STRATE
GIES WERE IMPLEMENTED SIMULTANEOUSLY !MONG THEM EAGLE LIVE
TRAPPING WAS
QUITE SUCCESSFUL WITH LESS THAN TEN INDIVIDUALS PROVING IMPOSSIBLE TO TRAP OR THAT
KEPT COMING BACK FROM THE TRANSLOCATION AREA (OWEVER IT WAS EASY TO SEE THAT AS
LONG AS PIGS REMAINED EAGLES WOULD START BREEDING ON THE ISLAND AGAIN AND THUS
START A NEW POPULATION 4HE SOLUTION THEREFORE SEEMED TO BE THE COMPLETE REMOVAL
OF PIGS FROM 3ANTA #RUZ )SLAND 9ET THE STUDY OF A MODEL BASED ON THE PREVIOUS ONE
SHOWED ONCE MORE THAT INDIRECT INTERACTIONS MAY LEAD TO COUNTER
INTUITIVE RESULTS
#OURCHAMP ET AL B !S FOR THE PREVIOUS EXAMPLE THE BASIC MODEL SHOWN
IN &IG " WAS CHANGED SIMPLY BY ADDING A LINEAR CONTROL TERM TO THE PIG AND TO
THE EAGLE EQUATIONS
"Y VARYING THE CONTROL RATE OF PIGS AND EAGLES FROM ZERO NO CONTROL TO ONE
ERADICATION WE CAN MIMIC DIFFERENT CONTROL STRATEGIES CONTROL OF PIGS ONLY OF
EAGLES ONLY OR OF BOTH SPECIES WITH DIFFERENT STRENGTH AND COMPARE THEIR RELATIVE
EFlCIENCY WITH NO RISK TO THE LOCAL POPULATIONS $OING SO REVEALED THAT MITIGATION
OF PIGS WOULD IN FACT LEAD TO A DECREASE IN THE FOX POPULATION &IG $ %RADICATION
OF PIGS THE INTENDED COURSE OF ACTION ON 3ANTA #RUZ WOULD LEAD TO FOX EXTINCTION
$UE TO THE LOW FOX POPULATION AND THE LARGE EAGLE POPULATION THE FOXES WOULD
BE ENTIRELY DESTROYED BEFORE THE EAGLES DIED OR EMIGRATED )N THEORY THE SOLU
TION IS THUS SIMPLE REMOVE BOTH THE EAGLES AND THE PIGS (OWERVER IN PRACTICE
THE REMOVAL OF SUCH A LARGE PIG POPULATION IS LOGISTICALLY DIFlCULT )N ADDITION
THE REMOVAL OF THE EAGLE WOULD BE IMPOSSIBLE THROUGH LIVE TRAPPING ONLY AND
ETHICALLY AND LEGALLY CHALLENGING BECAUSE GOLDEN EAGLES ARE PROTECTED SPECIES IN
THE 53! AND THEREFORE CANNOT BE KILLED
& #OURCHAMP AND 3 #AUT
4O CONCLUDE THIS PART IT MAY BE INTERESTING TO NOTE THAT WHEN MORE SPECIES AND
MORE INTERACTIONS ARE TAKEN INTO ACCOUNT NEW PROCESSES MAY BE UNVEILED THAT
COULD NOT BE PERCEIVED WITH ONLY TWO
SPECIES STUDIES 9ET IF BETTER UNDERSTAND
ING A SYSTEM IS UNDOUBTEDLY USEFUL FOR CONSERVATION MANAGERS IT IS NOT ALWAYS
SUFlCIENT FOR THEM TO BE ABLE TO KNOW HOW TO ACT )N THE PRESENT CASE A STUDY
TAKING ALL SPECIES INTO ACCOUNT REVEALED THAT A SEEMINGLY OBVIOUS LINE OF ACTION
PIG REMOVAL WOULD INDEED LIKELY ACHIEVE RESULTS OPPOSITE TO THOSE DESIRED AND
POSES A DIFlCULT DECISION TO MAKE REMOVE A PROTECTED POPULATION IN ORDER TO SAVE
AN ENDANGERED SUBSPECIES )F IT IS OBVIOUSLY NOT TRIVIAL TO MAKE CONSERVATION
CHOICES EVEN IN THE SIMPLEST SITUATIONS IT CAN SOMETIMES BECOME A CHALLENG
ING DILEMMA THEORETICALLY LOGISTICALLY LEGALLY AND MORALLY )N THE PRESENT CASE
THE ONLY REMAINING POPULATIONS OF SEVERAL FOX SUBSPECIES WERE THREATENED WITH
IMMINENT EXTINCTION (OWEVER THE PROXIMATE CAUSE OF THIS THREAT IS THE PRESENCE
OF A PROTECTED BIRD &URTHERMORE THE DECLINE OF THE FOX BENElTS THE ONLY POPULA
TIONS OF AN ENDEMIC SKUNK !S WE HAVE SEEN WHEN A DIFlCULT CHOICE IS MADE
THE OPPOSITE OUTCOME MAY WELL ARISE &URTHERMORE ALL THIS IS WITHOUT CONSIDERING
SPECIES OUTSIDE THIS SIMPLISTIC SYSTEM /NE COULD ALSO CONSIDER THE QUESTION UNDER
A WIDER ANGLE FOR EXAMPLE INCLUDING THE 3AN #LEMENTE LOGGERHEAD SHRIKE ,ANIUS
LUDOVICIANUS MEARNSI A CRITICALLY ENDANGERED BIRD TO WHICH THE INSULAR FOX IS THE
MAIN PREDATOR /N 3AN #LEMENTE )SLAND THE FOX POPULATION HAS BEEN IRONICALLY
ADVERSELY IMPACTED BY A 53 .AVY PROGRAM TO PROTECT THIS BIRD ATTEMPTING TO
THWART ANY PREDATION OF SHRIKES FOX WERE INITIALLY TRAPPED AND SHIPPED OFF ISLAND
OR EUTHANIZED DURING THE SHRIKE
NESTING SEASON FOXES REMOVED IN OF
WHICH PERMANENTLY "UT THIS IS ANOTHER STORYx
!LONG THE SAME LINE OF THIS CONSERVATION RIDDLE THE NEXT PART OF THIS CHAPTER
INVESTIGATES THE IMPORTANCE OF DIRECT INTERACTIONS IN CONTROL PROGRAMS WITH THE
AIM OF SHOWING THAT THE REMOVAL OF THE PRIMARY CAUSE OF A ECOSYSTEM DISFUNCTION
WILL NOT ALWAYS HELP RESTORE THE INITIAL CONDITIONS )N SOME CASES NOT TAKING INTO
ACCOUNT INDIRECT INTERACTION MAY LEAD TO EVEN FURTHER DAMAGE TO THE POINT THAT
IT MAY BE WISER TO ADVOCATE NOT TO REMOVE POPULATIONS THAT ARE KNOWN TO CAUSE
DIRECT NEGATIVE IMPACTS ON INVADED COMMUNITIES AT LEAST UNTIL ADEQUATE KNOWL
EDGE IS GAINED AND RELEVANT CONTROL STRATEGIES ARE INFERRED
#ONTROL OF INVADERS AS A SPECIES REMOVAL EXPERIMENT
2ELEASE FROM INTRODUCED HERBIVORES
/NE CONCEPT THAT IS RELATIVELY NEW IN THE STUDY OF INVADING SPECIES AND THAT HAS
BEEN THE CORE PRINCIPLE OF OUR OWN STUDIES IS THAT EVEN IF A SPECIES IS PROVEN TO
BE INmICTING IMPORTANT DAMAGES TO A COMMUNITY IT INVADES THE MERE REMOVAL OF
THAT SPECIES MAY NOT SYSTEMATICALLY BE THE SOLUTION TO RESTORING THE COMMUNITY
!S WE HAVE SHOWN WITH THE #ALIFORNIA #HANNEL )SLANDS EXAMPLE UNCONSIDERED
CONTROL MAY EVEN LEAD TO THE OPPOSITE OUTCOME THAT IS FURTHER DAMAGE INCLUD
ING POSSIBLE EXTINCTION OF THE SPECIES INTENDED TO BE PROTECTED 4HIS FACT HIGHLIGHTS
$YNAMICS OF INTERACTING POPULATIONS
THE NEED TO HAVE A CLEAR AND COMPLETE VIEW OF THE RELATIONSHIPS AMONG SPECIES THAT
ARE CONNECTED DIRECTLY OR INDIRECTLY WITH THE INTRODUCED SPECIES THAT IS SUBJECT TO
CONTROL )T NOWADAYS SOUNDS TRIVIAL TO STATE THAT ALL THE SPECIES THAT INTERACT WITH A
POPULATION FOR WHICH REMOVAL IS PLANNED ARE LIKELY TO BE AFFECTED IN DIVERSE WAYS
BY ANY SUCH ACTIONS 4HUS THE SUCCESS OF AN ERADICATION PROGRAM IS MEASURED NOT
ONLY BY THE COMPLETE REMOVAL OF THE CONTROLLED SPECIES BUT ALSO BY THE ABSENCE
OF FURTHER DYSFUNCTION 9ET SUCH ERRORS STILL OCCUR REGULARLY DURING CONSERVATION
PROGRAMS SOMETIMES SIMPLY BECAUSE CONSERVATION PROGRAMS HAVE INSUFlCIENT
FUNDS TO ALLOW THOROUGH PRE
CONTROL STUDIES OF THE INVADED COMMUNITY AS WELL
AS LONG TERM POST
CONTROL MONITORING 3OMETIMES SIMPLY BECAUSE CONSERVATION
ACTION IS URGENTLY NEEDED AND THERE IS NO TIME FOR SUCH PRE
CONTROL STUDY
4HE IMPORTANCE OF KNOWING THE RELATIONSHIPS BETWEEN INVADING SPECIES AND
THOSE IN THE INVADED COMMUNITY IS WELL ILLUSTRATED BY THE GOAT AND PIG ERADICATION
ON THE 3ARIGAN )SLAND 4HIS ISLAND IS PART OF THE #OMMONWEALTH OF THE .ORTHERN
-ARIANA )SLAND IN THE 0ACIlC /CEAN )NTRODUCED PIGS AND GOATS THREATENED THE
LOCAL mORA AND FAUNA TRIGGERING A CONSERVATION PROGRAM CONSISTING MAINLY OF
GOAT AND PIG ERADICATION +ESSLER !S THE ISLAND IS ISOLATED AND HENCE
DIFlCULT TO ACCESS THE PROGRAM DESIGNED INCLUDED ONLY A MINIMAL PRE
ERADICA
TION STUDY 4HE PROGRAM WAS CONSIDERED A FULL SUCCESS IN TERMS OF REMOVING THE
INTRODUCED MAMMALS HOWEVER IT FAILED IN ITS ABILITY TO DETECT THAT THE ISLAND HAD
ALSO BEEN COLONIZED BY AN INTRODUCED VINE /PERCULINA VENTRICOSA WHICH APPEARED
TO BE A PREFERENTIAL FOOD ITEM FOR THE GOATS )T IS PERHAPS NOT SURPRISING THAT THIS
VINE WAS NOT FOUND IN THE PRE
CONTROL STUDY AS IT WAS LIKELY TO HAVE BEEN AT A LOW
DENSITY DUE TO SELECTIVE GRAZING BY GOATS %VEN A VERY THOROUGH STUDY MIGHT HAVE
FAILED TO SEE IT 9ET IT MAY BE VALUABLE TO POINT OUT THAT WHEN POSSIBLE SIMPLE
FENCED EXCLOSURE STUDIES PRIOR TO ERADICATIONS CAN OFTEN HELP LAND MANAGERS SEE IF
UNWANTED RESULTS WILL ARISE AFTER AN ERADICATION OF HERBIVORES )F SO THEN APPROPRI
ATE CONTROL OF NON
NATIVE PLANTS CAN BE PLANNED ALONG WITH THE HERBIVORE REMOVAL
5NFORTUNATELY THIS WAS NOT DONE AND THE CONTROL PROGRAM WHICH AIMED AT RELEAS
ING PLANT SPECIES FROM GOAT GRAZING HAD A DIFFERENT IMPACT ON THE OVERALL PLANT
COMMUNITY THAN THE ONE EXPECTED !S THE PRESSURE OF GRAZING WAS REMOVED FROM
ALL GRAZED PLANTS INTRODUCED PLANTS WERE ABLE TO FULLY EXPRESS THEIR COMPETITIVE
SUPERIORITY WITH REGARDS TO NATIVE PLANTS RESULTING IN THE RAPID INVASION OF THE
COMMUNITY &IG SHOWS HOW WITHIN ONLY TWO YEARS THE REMOVAL OF AN EXOTIC
GRAZER LED TO A COMPLETE INVASION OF THE ISLAND COMMUNITY BY AN EXOTIC PLANT THAT
APPEARS TO HAVE A COMPETITIVE SUPERIORITY OVER LOCAL PLANTS !S MOST OF THE 3ARIGAN
)SLAND ECOSYSTEM IS NOW COVERED BY VINES ONE CAN EASILY IMAGINE HOW THE INDIRECT
EFFECT OF HAVING REMOVED GOATS IS NOW DELETERIOUS FOR THE LOCAL PLANTS AS WELL AS
ANIMALS THAT DEPEND UPON THEM
4HE MESOPREDATOR RELEASE EFFECT
4HE PROCESS WE HAVE SEEN WITH THE RELEASE OF AN EXOTIC PLANT MAINTAINED AT LOW
DENSITY BY A BROWSER CAN BE GENERALIZED TO RELEASES FROM ALMOST ANY OTHER TYPE OF
& #OURCHAMP AND 3 #AUT
&IG %VOLUTION OF THE LANDSCAPE OF 3ARIGAN )SLAND FOLLOWING THE ERADICATION OF GOATS
!S THIS INTRODUCED HERBIVORE NO LONGER HELD IN CHECK THE EXPANSION OF THE INTRODUCED
VINE A FAVORED FOOD ITEM THE VINE RAPIDLY INCREASED COVERING MOST OF THE INSULAR
PLANT COMMUNITIES WITHIN TWO YEARS 4HIS ILLUSTRATES HOW AN EXOTIC GRAZER AFFECTED THE
COMPETITION RELATIONSHIPS BETWEEN LOCAL AND INTRODUCED PLANTS AND HOW ITS REMOVAL CAN
LEAD TO DRAMATIC AND UNEXPECTED OUTCOME FOR THE COMMUNITIES WHICH PROTECTION WAS
AIMED AT
$YNAMICS OF INTERACTING POPULATIONS
NATURAL ENEMY 7E WILL ILLUSTRATE THIS WITH THE NEXT TWO EXAMPLES 4HIS IS HOW
EVER NOT TO BE MISTAKEN WITH THE ENEMY RELEASE HYPOTHESIS +EANE AND #RAWLEY
WHICH PROPOSES THAT INVADING SPECIES ARE SO SUCCESSFUL PARTLY BECAUSE THEY
ARE RELEASED IN THE INVADED HABITAT FROM THE PRESSURE OF THEIR NATURAL ENEMIES
RARELY INTRODUCED WITH THEM
/N 3TEWART )SLAND .EW :EALAND A POPULATION OF INTRODUCED CATS WAS THREAT
ENING ONE OF THE LAST POPULATIONS OF KAKAPO 3TRIGOPS HABROPTILUS AN ENDEMIC
mIGHTLESS PARROT ! DIET STUDY REVEALED KAKAPO REMAINS IN OF COLLECTED
CAT SCATS +ARL AND "EST 4HIS SEEMINGLY LOW PREDATION PRESSURE CAN HAVE
A DRAMATIC EFFECT ON INSULAR POPULATIONS WHICH EVOLVED IN THE ABSENCE OF SUCH
PREDATORS AND ARE THEREFORE NOT ADAPTED TO EVEN LOW LEVELS OF PREDATION -OREOVER
THE KAKAPO POPULATION WAS ALREADY SMALL AND FRAGILE ADDING TO THE WEIGHT THAT
INTRODUCED CATS COULD HAVE ON ITS FATE 4HIS COULD HAVE BEEN SUFlCIENT TO TRIGGER
A PROGRAM OF CAT CONTROL ON THE ISLAND
(OWEVER RATS KNOWN TO BE IMPORTANT BIRD PREDATORS HAD ALSO BEEN INTRODUCED
ON 3TEWART )SLAND )N THE SAME DIET STUDY +ARL AND "EST RAT REMAINS WERE
FOUND IN OF THESE CAT FAECES 4HIS SHOWS THE INDIRECT ROLE CATS MIGHT PLAY IN
PRESERVING NATIVE FAUNA THROUGH REDUCTION OF RAT PREDATION PRESSURE ON KAKAPO
)N FACT IT IS EASY TO SEE THAT IN SOME CASES THE INDIRECT POSITIVE EFFECT OF CAT PREDA
TION ON RATS IS MORE BENElCIAL FOR THE LOCAL PREY THAN THE DIRECT NEGATIVE EFFECTS OF
CAT PREDATION ON THE PREY THEMSELVES )N SUCH CASES THE ELIMINATION OF THE FERAL
CAT POPULATION COULD LEAD TO A MORE SEVERE NEGATIVE IMPACT ON THE LOCAL SPECIES
THROUGH AN INCREASE IN THE RODENT POPULATION AS A CONSEQUENCE OF THE REMOVAL
OF THEIR PREDATORS 4HE ATTEMPTED REDUCTION OF THE CAT POPULATION ON !MSTERDAM
)SLAND HAS BEEN ABANDONED AS IT IS ALLEGED TO HAVE CAUSED A COMPENSATING RISE IN
THE NUMBER OF RATS AND MICE (OLDGATE AND 7ACE 4HIS PROCESS TERMED
hMESOPREDATOR RELEASEv HAS BEEN DESCRIBED IN FRAGMENTED INSULAR ECOSYSTEMS
3OUL ET AL AND APPLIES WELL TO MANY INSULAR FOOD WEBS EG 3CHOENER
AND 3PILLER
#ONVERSELY THE ERADICATION OF RODENTS lRST WHICH HAS NOW PROVEN FEASIBLE
EVEN ON RELATIVELY LARGE ISLANDS MIGHT INDUCE CATS TO SWITCH PREY RESULTING IN
A BRUTAL INCREASE IN PREDATION PRESSURE ON THE THREATENED INDIGENOUS SPECIES AS
EXPERIENCED FOR STOATS AND RATS IN .EW :EALAND -URPHY AND "RADlELD
4HIS IS A SIMILAR PROCESS TO THE ONE DESCRIBED ABOVE FOR THE PIG CONTROL IN PRESENCE
OF EAGLES ON 3ANTA #RUZ )SLAND !S THE OPTIMAL CONTROL STRATEGY IS NEITHER SIMPLE
TO lND NOR INTUITIVE IT IS CONVENIENT TO STUDY IT THROUGH THE ANALYSIS OF A MATH
EMATICAL MODEL WHICH MIMICS THE DYNAMICS OF THE THREE SPECIES IN THIS SYSTEM
4HE MAIN RESULTS OF SUCH A STUDY #OURCHAMP ET AL A ARE REPRODUCED BELOW
TO ILLUSTRATE HOW THE CONTROL OF AN INVADING SPECIES CAN PROVIDE AN IDEAL OPPORTU
NITY TO PROGRESS IN THE UNDERSTANDING OF THE NUMEROUS AND OFTEN COMPLEX INTERAC
TIONS AMONG POPULATIONS
&OR THE SAKE OF CLARITY WE WILL NOT PRESENT THE MODEL EQUATION IN ANY DETAIL
3UFlCE TO SAY THAT THE SYSTEM IS DESCRIBED BY A SET OF THREE COUPLED EQUATIONS
ONE PREY PARROT ITS PREDATOR THE RAT IN THIS CASE A MESOPREDATOR AND ONE
& #OURCHAMP AND 3 #AUT
SUPERPREDATOR CAT WHICH EATS BOTH THE PREY AND THE MESOPREDATOR 4HE MODEL
AND THE RESULTING POPULATION TRENDS ARE REPRESENTED IN &IG
4HE STUDY OF THE ABOVE SYSTEM LEADS TO THE QUITE OBVIOUS CONCLUSION THAT BOTH
PREDATORS SHOULD BE ERADICATED AT THE SAME TIME (OWEVER NOT ONLY IS SUCH A
STRATEGY CHALLENGING FROM A LOGISTICAL POINT OF VIEW BUT IN ADDITION IT MAY NOT
ALWAYS BE THE BEST SOLUTION ESPECIALLY IF THE SYSTEM INCLUDES MORE INTRODUCED
PREDATORS )NDEED THE PREY
MESOPREDATOR
SUPERPREDATOR SYSTEM THAT WE HAVE
JUST DESCRIBED CAN BE FURTHER COMPLICATED 4HERE ARE CASES WHERE THE PRESENCE
OF A THIRD PREDATOR CAN RENDER THE ERADICATION OF THE MESOPREDATOR PROBLEMATIC
IF IT ALSO ACTS AS A PREDATOR TO THIS THIRD PREDATOR AND THUS AT THE SAME TIME AS
A MESOPREDATOR AND A SUPERPREDATOR "EING AWARE OF THE POTENTIAL DANGERS OF
REMOVING A SUPERPREDATOR WHEN A MESOPREDATOR IS PRESENT THE MANAGERS OF THE
CONSERVATION PROGRAM OF "IRD )SLAND 3EYCHELLES DECIDED RIGHTLY TO SIMULTANE
OUSLY REMOVE INTRODUCED CATS AND RATS IN ORDER TO PROTECT THE LOCAL BIRD COLONIES
4HEY HAD HOWEVER OVERLOOKED THE PRESENCE OF THE INTRODUCED CRAZY ANT
!NOPLOLEPIS LONGIPES WHICH WERE PRESENT IN VERY LOW NUMBERS ON THE ISLAND
&EARE 4HE LARVAE OF THESE ANTS SEEM TO HAVE BEEN AN IMPORTANT PREY
ITEM OF THE INTRODUCED RODENTS SUCH THAT THE RAT ERADICATION LED TO A DEMOGRAPHIC
EXPLOSION OF THE ANTS 4HIS RESULTED IN THE ANTS COVERING A LARGE PART OF THE ISLAND
WITH A HEAVY IMPACT ON LAND CRABS AND BIRD COLONIES )N FACT THIS PROBLEM OF CHAIN
REACTION FOLLOWING THE REMOVAL OF AN INTRODUCED SPECIES ALSO CALLED SURPRISE EFFECT
OR 3YSIPHUS EFFECT -ACK AND ,ONSDALE CAN BE GENERALIZED TO OTHER NATURAL
ENEMIES SUCH AS HERBIVORES OR COMPETITORS
4HE COMPETITOR RELEASE EFFECT
4HE CONTROL OF AN INVADER HAS THE POTENTIAL TO RELEASE ANY SPECIES INTERACTING
WITH THE CONTROLLED INVADERS FROM ITS PRESSURE BE IT EXPLOITATION OR INTERFER
ENCE 4HEREFORE ONE CAN IMAGINE VERY SIMILAR PROCESSES WITH A BROWSER OR A
COMPETITOR ,ET US CONSIDER AS A lNAL EXAMPLE A hCOMPETITOR RELEASE EFFECTv ,ET
US SET THE SCENE AN ISLAND INVADED BY SAY A RAT SPECIES 4HE ISLAND IS NORTH OF
.EW #ALEDONIA IN THE %NTRECASTEAUX 2EEF ,ET US CALL IT 3URPRISE )SLAND WHICH
SUITS VERY WELL A STUDY ON SURPRISE EFFECTS ! THOROUGH STUDY OF THE INVADED ECO
SYSTEM COMPLETED TO CHARACTERIZE THE IMPACT OF INTRODUCED RATS ON THAT ISLAND
REVEALED THE PRESENCE OF A SMALL ISOLATED POPULATION OF INTRODUCED DOMESTIC MICE
4HE POPULATION SEEMS SMALL AND RESTRICTED SO THAT THEIR IMPACT ON THE ECOSYSTEM
IS LIKELY TO BE NEGLIGIBLE )N FACT IN SIMILAR SITUATIONS THEY HAVE BEEN IN THE PAST
NEGLECTED PARTLY BECAUSE MOUSE POPULATIONS ARE DIFlCULT TO ERADICATE PARTLY
BECAUSE SUCH SMALL POPULATIONS WERE NOT VIEWED AS A THREAT AND PARTLY BECAUSE
THE CONSERVATION PROGRAM CONCERNED ANOTHER SPECIES AND FUNDS AND PROTOCOLS
WERE NOT AVAILABLE TO DEAL WITH MICE !S AN EXAMPLE THE RAT AND RABBIT CONTROL OF
3AINT 0AUL )SLAND IN THE !NTARCTIC OCEAN HAS BEEN VERY SUCCESSFUL IN ERADICATING
THESE TWO INTRODUCED MAMMALS IT WAS EVEN AT THE TIME THE GREATEST AREA EVER
CLEANED UP FROM INTRODUCED RABBITS BUT THE PROGRAM DID NOT FOCUS ON THE SMALL
$YNAMICS OF INTERACTING POPULATIONS
&IG 2EPRESENTATION OF THE MESOPREDATOR RELEASE PROCESS WITH THE INTRODUCED
SUPERPREDATOR BEING THE CAT # RED THE INTRODUCED MESOPREDATOR BEING THE RAT 2 YELLOW
AND THE INDIGENOUS PREY BEING THE BIRD " BLUE !S IN &IG ! IS THE COMPARTMENTAL
REPRESENTATION THE EXAMPLE COMING FROM 3TEWART )SLAND OFF THE .EW :EALAND MAINLAND
" IS THE RESULTING SET OF EQUATIONS AND THE TWO FOLLOWING PANELS ARE TYPICAL POPULATION
TRENDS FOLLOWING INTRODUCTIONS OF THE SUPERPREDATOR # TOP AND OF THE MESOPREDATOR
# BOTTOM OR CONTROL OF THE SUPERPREDATOR ONLY $ TOP AND OF BOTH THE SUPERPREDATOR
AND THE MESOPREDATOR $ BOTTOM 0ANEL # SHOWS THAT IN SOME CASES A SUPERPREDATOR
INTRODUCTION WILL DECREASE THE PREDATION PRESSURE ON LOCAL PREY VIA ITS PREDATION ON THE
MESOPREDATOR TOP WHILE A MESOPREDATOR INTRODUCTION WILL ALLOW AN INCREASE OF THE
SUPERPREDATOR PROCESS SIMILAR TO THE HYPERPREDATION LEADING TO A FURTHER DECREASE OF THE
PREY VIA A COMBINATION OF INCREASED PREDATION OF THE INCREASED SUPERPREDATOR POPULATION
AND ADDITIONAL PREDATION FROM THE NEWLY INTRODUCED MESOPREDATOR BOTTOM 0ANEL $
SHOWS THAT CONTROL OF BOTH INTRODUCED PREDATOR NEEDS TO BE DONE TO PROTECT THE PREY
BOTTOM AS THE CONTROL OF ONLY THE SUPERPREDATOR CAN TRIGGER A MESOPREDATOR RELEASE
WHICH EVENTUALLY LEADS TO THE LOCAL PREY EXTINCTION TOP
& #OURCHAMP AND 3 #AUT
MOUSE POPULATION THAT WAS KNOWN TO OCCUR ON THE ISLAND -ICOL AND *OUVENTIN
4HE STUDY OF THE 3URPRISE ECOSYSTEM SUGGESTS THAT RESOURCES ARE ABUNDANT
ENOUGH FOR THE INTRODUCED MICE TO DEVELOP A LARGER POPULATION THAN THEY HAVE
DONE 4HIS SUGGESTS THAT THE MOUSE POPULATION IS RESTRICTED BY A NATURAL ENEMY
AND IT COMES NATURALLY TO MIND THAT IT IS THE COMPETING RAT THAT RESTRICTED THEM
FROM FURTHER EXPANDING THEIR RANGE !LTHOUGH IT IS QUITE DIFlCULT TO UNAMBIGU
OUSLY DEMONSTRATE SUCH COMPETITION RELATIONSHIPS lELD SPECIALISTS SEEM UNANI
MOUS IN THE VIEW THAT RATS ARE STRONG COMPETITORS OF MICE TO THE POINT OF OFTEN
EXCLUDING THEM WHEN COMMON RESOURCES ARE FEW )T SEEMS THEN QUITE PREDICTABLE
ESPECIALLY AFTER HAVING READ THE PREVIOUS EXAMPLES OF hSURPRISE EFFECTSv IN THIS
CHAPTER THAT THE PLANNED ERADICATION OF THE INTRODUCED RATS ON 3URPRISE )SLAND IS
LIKELY TO RELEASE THE MICE FROM RAT COMPETITION AND THUS TO ALLOW THEM TO INCREASE
IN NUMBERS 3UCH AN OUTCOME IS NOT ONLY INTUITIVELY LOGICAL IT IS ALSO VERY EASY TO
DEMONSTRATE THROUGH THE ANALYSIS OF A BASIC ,OTKA
6OLTERRA COMPETITION MODEL TO
WHICH A CONTROL TERM IS ADDED TO ONE OF THE COMPETITORS &OR THE SAKE OF SIMPLIC
ITY WE STUDY THIS THREE
SPECIES SYSTEM ONE PREY AND TWO COMPETING PREDATORS
THROUGH ONLY A TWO COMPETITORS MODEL THE SHARED PREY IS IGNORED HERE WHICH
ALSO ALLOWS A GENERALIZATION OF THE SYSTEM TO NON
PREDATOR COMPETITORS
!NALYSIS OF THIS SYSTEM CLEARLY SHOWS THAT THE MITIGATION OF THE HIGHER COM
PETITOR THE RAT WILL LEAD TO AN INCREASE OF THE LOWER COMPETITOR AS PRESSURE FROM
COMPETITION IS LIFTED 4HE HIGHER THE MITIGATION THE LARGER THE MOUSE POPULATION
! SUDDEN COMPLETE REMOVAL OF THE RAT POPULATION IS LIKELY TO RESULT IN A DEMO
GRAPHIC EXPLOSION OF THE MOUSE POPULATION 4HIS WAS THE CASE ON 3AINT 0AUL )SLAND
FOLLOWING THE REMOVAL OF RATS IN -ICOL AND *OUVENTIN RELEASED FROM
THEIR COMPETITORS MICE NUMBERS INCREASED DRAMATICALLY TO SUCH A POINT THAT FOR
A TIME THEY FAR EXCEEDED THE CARRYING CAPACITY OF THE HABITAT /BVIOUSLY MICE ARE
LESS HARMFUL THAN RATS AND THUS IN SOME CASES THE END BENElT OF THE RAT REMOVAL
IS POSITIVE EVEN IF THE MOUSE POPULATION INCREASES 9ET MOUSE OUTBREAKS CAN
BE VERY PROBLEMATIC AS MICE HAVE BEEN SHOWN TO BE ACTIVE PREDATORS OF INVERTE
BRATES REPTILES AND EVEN BIRDS THAT CAN BE TIMES THEIR WEIGHT #AMPOS AND
'RANADEIRO #UTHBERT AND (ILTON &ITZGERALD ET AL ,E 2OUX
ET AL
.EWMAN 3MITH ET AL
"UT THIS COMPETITOR RELEASE EFFECT IS IN FACT NEITHER SURPRISING NOR VERY INTER
ESTING FOR THE STUDY OF INTERSPECIlC RELATIONSHIPS 4HE OBVIOUS APPROACH TO SUCH
SITUATIONS SEEMS SIMPLY TO APPLY A SIMULTANEOUS CONTROL TO BOTH COMPETITORS !ND
IT IS ALL THE BETTER THAT WE ARE DEALING WITH COMPETING RODENTS AS A SIMULTANEOUS
CONTROL IS EASILY FEASIBLE WITH A COMMON RODENTICIDE FOR EXAMPLE %ND OF STORY OR
SO IT SEEMS )N FACT JUST TO MAKE SURE NO MORE SURPRISES ARE GOING TO EMERGE FROM
THE SYSTEM IT IS POSSIBLE TO COMPLETE THE MODEL BY ADDING A SIMULTANEOUS CONTROL
TERM TO THE INFERIOR COMPETITOR AS WELL SEE &IGS ! AND " )T SEEMS LOGICAL TO LINK
THE TWO CONTROL RATES FOR THEY WILL OFTEN BE AT LEAST IN THE CASE OF RODENTS CON
TROLLED IN THE SAME PROGRAM &OR EXAMPLE ONE CAN HAVE tR b tM WITH tM BEING
THE CONTROL RATE OF THE MICE THE LOWER COMPETITOR THE MOUSE AND b THE CONTROL
$YNAMICS OF INTERACTING POPULATIONS
SPECIlCITY ! VALUE OF SAY FOR b MEANS THAT SUPERIOR COMPETITORS ARE CONTROLLED
TWICE AS MUCH AS INFERIOR COMPETITORS 4HIS IS VERY LIKELY IF FOR EXAMPLE BAITS ARE
ACCESSIBLE IN PRIORITY TO HIGHER COMPETITORS OR IF THE TRAPPING DESIGN IS AIMED AT
THE HIGHER COMPETITOR BUT THEY ALSO ALLOW TO TRAP THE LOWER COMPETITOR ALTHOUGH
WITH A LOWER EFlCIENCY THE PROGRAM AIM IS TO REMOVE THE RATS BUT RAT TRAPS ALSO
CAN CATCH MICE 3TUDYING THIS VERY SIMPLE SYSTEM REVEALS IN FACT A SUBTLER AND LESS
EXPECTED COMPETITOR RELEASE EFFECT !S SHOWN IN &IG # THE SIMULTANEOUS MITIGA
TION OF BOTH COMPETITORS CAN LEAD TO A RELEASE OF THE LOWER COMPETITOR )N SOME
CASES THIS RELEASE CAN AMOUNT TO ACTUAL POPULATION EXPLOSIONS )T IS POSSIBLE THAT
AIMING AT CONTROLLING SAY AN INTRODUCED RODENT A CONTROL PROGRAM WILL LEAD TO A
DRAMATIC INCREASE OF ANOTHER RODENT EVEN IF THAT ONE IS CONTROLLED TOO
-OREOVER &IG $ SHOWS THAT THE COMPETITOR RELEASE IS DIRECTLY PROPORTIONAL
TO THE CONTROL RATE 4HIS MEANS THAT THE MORE THE TARGETED SPECIES IS CONTROLLED
THE MORE IMPORTANT THE COMPETITOR RELEASE EFFECT WILL BE 4HIS IS NOT SO OBVIOUS TO
PREDICT AS IT IMPLIES THAT THE INFERIOR COMPETITOR IS CONTROLLED TOO WITH A CONTROL
EFFORT THAT INCREASES AT THE SAME RATE AS THAT OF THE SUPERIOR COMPETITOR )N OTHER
WORDS THE MORE RODENTS ARE CONTROLLED THE MORE MICE APPEAR %VEN IF MICE ARE
ACTUALLY CAUGHT IN TRAPS AND KILLED BY POISON THE MORE WE KILL THEM THE MORE
THEIR POPULATION WILL INCREASE 1UITE THE OPPOSITE OF AN EXPECTED OUTCOME
!LTHOUGH THIS PROCESS MAY BE LESS INTUITIVE AND THEREFORE LESS OFTEN FORESEEN IT
IS A POSTERIORI QUITE EASY TO UNDERSTAND 4HIS IS LIKELY TO OCCUR AS SOON AS THE LOWER
COMPETITOR BENElTS FROM THE DIFFERENTIAL EFFECT OF THE SIMULTANEOUS CONTROL OF BOTH
COMPETITORS WHEN ITS INDIRECT POSITIVE EFFECT THE REMOVAL OF THEIR COMPETITORS
EXCEEDS ITS DIRECT NEGATIVE EFFECT THEIR OWN REMOVAL
/BVIOUSLY SUCH A PROCESS CAN BE INTERESTING IF THE LOWER COMPETITOR IS A LOCAL
SPECIES WHICH SURVIVAL WAS THREATENED BY THE CONTROLLED POPULATION )N THIS CASE
THE DRAMATIC INCREASE OF ITS POPULATION FOLLOWING COMPETITION RELEASE IS NOTH
ING LESS THAN THE PROGRAM OBJECTIVES 4HIS CAN BE CONSIDERED IN CASES WHERE AN
INTRODUCED SPECIES IS TO BE REMOVED BECAUSE IT THREATENS A LOCAL POPULATION
THROUGH COMPETITION BUT WHERE CONTROL PROGRAMS WERE NOT IMPLEMENTED FOR
FEAR OF DAMAGE TO NON TARGET SPECIES )N THOSE CASES ANY UNINTENDED NON TARGET
DEATH SHOULD BE MORE THAN BALANCED BY THE DEATH OF THE INTRODUCED COMPETITORS
4HERE IS NO HAPPY ENDING HOWEVER IF THE LOWER COMPETITOR APPEARS TO BE ANOTHER
INTRODUCED SPECIES WHICH INCREASE CAN INmICT FURTHER DAMAGES TO THE INVADED
ECOSYSTEM ESPECIALLY IF THAT INCREASE IS DRAMATIC !GAIN IN MANY CONSERVATION
SITUATIONS MANAGERS HAVE TO MAKE TRADE OFF CHOICES AND IT MAY APPEAR THAT A
COMPETITOR RELEASE BE EVENTUALLY LESS DETRIMENTAL THAN ALLOWING THE PRESENCE OF
THE INTRODUCED PREDATOR 9ET DRAMATIC INCREASES OF ALIEN SPECIES ARE OFTEN VERY
HARMFUL TO ECOSYSTEMS AND THIS EVENTUALLY SHOULD ALWAYS BE ASSESSED
4HERE IS MUCH MORE THAT COULD BE SAID ABOUT SUCH A SYSTEM EVEN AS SIMPLE
AS IT IS AND ABOUT THE CASE STUDIES IN WHICH THE OVERLOOKING OF AN ENEMY RELEASE
LED TO SURPRISE EFFECTS THAT EVENTUALLY CAUSED FURTHER DAMAGES TO INVADED ECOSYS
TEMS (OWEVER BEING THAT THE AIM OF THIS CHAPTER IS TO CONVINCE THE READER THAT
BIOLOGICAL INVASIONS AND THEIR CONTROL CAN PROVIDE AN EXCELLENT MODEL SYSTEM FOR
& #OURCHAMP AND 3 #AUT
&IG )LLUSTRATION OF THE COMPETITOR RELEASE EFFECT 0ANEL ! SHOWS THE COMPARTMENTAL
REPRESENTATION OF THE EXAMPLE TAKEN FROM 3URPRISE )SLAND OFF .EW #ALEDONIA WHERE TWO
RODENTS ARE COMPETITORS RATS 2 RED BEING A SUPERIOR COMPETITOR OVER THE MICE - YELLOW
4HE EQUATION SET IN THE PRESENCE OF SIMULTANEOUS CONTROL IS GIVEN IN PANEL " (ERE BOTH
# AND $ REPRESENT POPULATION TRENDS FOLLOWING CONTROL 0ANEL # SHOWS THE POPULATION
TRENDS OF THE TWO COMPETITORS AS A FUNCTION OF TIME FOR TWO DIFFERENT COMBINATIONS OF CONTROL
EFFORT AND CONTROL SPECIlCITY SEE TEXT )N BOTH CASES THE RODENT CONTROL CAN LEAD TO THE
DEMOGRAPHIC EXPLOSION OF ONE OF THE RODENT POPULATIONS 0ANEL $ SHOWS THE POPULATION
TRENDS OF THE LOWER COMPETITOR AS A FUNCTION OF THE CONTROL EFFORT AND THE CONTROL EFlCIENCY
!S IN &IG $ THE COLORS ARE GIVEN HERE TO FACILITATE THE $ EFFECT 4HIS PANEL SHOWS THAT IF
THE COMPETITION IS STRONG THE DEMOGRAPHIC EXPLOSION OF THE LOWER COMPETITOR IS PROPORTIONAL
TO ITS CONTROL !S THE COMPETITION PRESSURE FROM THE SUPERIOR COMPETITOR IS LIFTED BY THE
CONTROL THE RESULTING GAIN IN POPULATION GROWTH IS COMPENSATING THE LOSSES OCCURRED BY
THE CONTROL RESULTING IN A LARGER POPULATION THAN IN ABSENCE OF CONTROL
$YNAMICS OF INTERACTING POPULATIONS
THE STUDY OF INTERSPECIlC RELATIONSHIPS WE WILL ONLY CLOSE THIS EXAMPLE BY AN
OBVIOUS IF OVERLOOKED STATEMENT )N SOME CASES THE DIRECT NEGATIVE IMPACT OF A
SPECIES CAN HIDE AN INDIRECT POSITIVE EFFECT ON THE SAME COMMUNITY SOMETIMES
ON THE VERY SAME SPECIES )N A CLASSICAL SYSTEM WHERE SPECIES ADDITION OR REMOVAL
IS NOT THE RULE SUCH INTERACTIONS CAN REMAIN HIDDEN )N THE CASE OF INVADED COM
MUNITIES AND OF ASSOCIATED CONSERVATION PROGRAMS SPECIES REMOVAL CAN HIGHLIGHT
THESE UNDETECTED RELATIONSHIPS 3UCH HIGHLIGHT IS HOWEVER DONE IN THE FORM OF
FURTHER DAMAGE IN THE SYSTEM WITH A RISK OF BIODIVERSITY LOSS AND SHOULD THERE
FORE BE ANTICIPATED )F INDIRECT POSITIVE EFFECTS ARE GREATER THAN THE DIRECT NEGATIVE
EFFECTS ONE MUST BE EXTREMELY CAUTIOUS IN ANY RESTORATION ACTION INTENDED TO
PROTECT THE AFFECTED SPECIES
#/.#,53)/.
"IOLOGICAL INVASIONS ARE PRIMARILY CONSIDERED FOR THEIR HARMFUL EFFECTS ON BIO
DIVERSITY IN INVADED ECOSYSTEMS ESPECIALLY ON ISLANDS 4HIS IS FAR FROM SURPRIS
ING AS ISLANDS ARE PLACES OF MAJOR BIOLOGICAL DIVERSITY AND ARE OFTEN INCLUDED IN
ECOLOGICAL HOTSPOTS -YERS ET AL 2EID AND OF HIGH PROBABILITY OF
EXOTIC SPECIES INTRODUCTION &OR EXAMPLE MAMMAL INTRODUCTIONS HAVE BEEN
DOCUMENTED ON ISLANDS %BENHARD WHILE 'ARGOMINY ET AL RECORDED MORE
THAN EXOTIC SPECIES OF mOWERING PLANTS IN THE WILD IN .EW #ALEDONIA ALONE
'ARGOMINY ET AL 4HESE lGURES HAVE UNDOUBTEDLY INCREASED NOWADAYS
4HE NUMBER OF SPECIES THAT WENT EXTINCT AS A RESULT OF THESE INVASIONS IS IMPOSING
TOO #ONSEQUENTLY THE NUMBER OF CONTROL PROGRAMS HAS BEEN ON THE RISE THIS LAST
DECADE AND HERE AGAIN THERE ARE A GREAT NUMBER OF DOCUMENTED EXAMPLES WITH
FOR EXAMPLE OVER ERADICATION PROGRAMS JUST FOR EXOTIC MAMMALS IN .EW
:EALAND #2 6EITCH PERS COMM
7E HAVE SO FAR INSISTED MUCH ON CAUTION AND PLANNING IN ANY MITIGATIONERADI
CATION EFFORT IN MANAGEMENT PROGRAMS 7E FEEL IT IS NEVERTHELESS ESSENTIAL TO START
OUR CONCLUSION BY REWORDING THE OBVIOUS THE BEST RESPONSE TO BIOLOGICAL INVASION
IS ALMOST ALWAYS MITIGATE AND WHEN POSSIBLE ERADICATE THE ALIEN POPULATION )N
MANY CASES A HESITANCY TO PROCEED WITH THIS HAS CAUSED MORE DAMAGE TO BIODI
VERSITY THAN HAVE THE UNEXPECTED RESULTS OF POORLY PLANNED OR SIMPLY UNLUCKY
ERADICATIONS
(OWEVER THE AIM OF THIS CHAPTER WAS TO TAKE A DIFFERENT POINT OF VIEW REGARD
ING BIOLOGICAL INVASIONS AND TO SHOW THAT SUCH EVENTS HOWEVER NOT SYSTEMATIC
CAN BE VIEWED AS OPPORTUNITIES TO INCREASE FUNDAMENTAL KNOWLEDGE IN ECOLOGY
!DDITIONS AND DELETIONS OF SPECIES IN TROPHIC WEBS THAT ARE COMPARATIVELY MUCH
SIMPLER THAN USUAL SHOULD GREATLY ENHANCE OUR ABILITY TO DISCERN INTRINSIC
DYNAMICAL PROCESSES AS WELL AS DIRECT AND INDIRECT INTERACTIONS BETWEEN SPECIES
AND HERE WE VIEW POPULATION DYNAMICS AS GOOD MARKERS OF SUCH MECHANISMS
4HIS APPROACH ALSO HAS THE DUAL ADVANTAGE OF ADDRESSING MECHANISMS ON A REAL
SCALE SOMETHING IMPOSSIBLE IN LABORATORY BASED STUDIES AND OF ENCOMPASSING
& #OURCHAMP AND 3 #AUT
ALL THE CHARMING CONSTRAINTS OF BIOLOGICAL REALITY SOMETHING MISSING IN PURELY
THEORETICAL ECOLOGY
)T IS OF COURSE NOT OUR PURPOSE TO MINIMIZE THE CONTRIBUTION OF THEORY IN THIS
REGARD $ESPITE THEIR INDUBITABLE ADVANTAGE IN THIS DOMAIN lELD
BASED STUDIES
THAT USE INVASIONS AS AN EXPERIMENT HAVE MUCH TO GAIN FROM A SYNERGY WITH
CONCOMITANT LABORATORY AND THEORETICAL APPROACHES 7E EVEN URGE CONSERVATION
BIOLOGISTS AND BIODIVERSITY MANAGERS TO USE THEORETICAL
BASED RESULTS TO FOUND
THEIR CONSERVATION PROGRAM DESIGN AND TO LOOK INTO HISTORICAL CASES IN ORDER NOT
TO ENDLESSLY REPEAT MISTAKES OF THE PAST !N IDEA ASSOCIATED TO THIS IS THE NEED FOR
CONSERVATIONISTS AS WELL AS BIODIVERSITY MANAGERS TO KNOW WHEN TO STOP A PRO
GRAM AND RETHINK THE STRATEGY #OURCHAMP ET AL A 4O KNOW WHEN WE ARE
STARTING TO PLAY THE WITCHS SORCERER AND WHEN TO STOP IS AN ESSENTIAL YET DIFlCULT
FACULTY (ISTORY HAS SHOWN US THAT OFTEN MORE HARM IS GENERATED WHEN TRIAL AND
ERROR PROCESSES ARE USED INDISCRIMINATELY /NE GOOD EXAMPLE OF THIS IS GIVEN BY THE
ATTEMPTED MITIGATION OF RATS IN SUGARCANE lELDS IN *AMAICA 4HERE CANE GROWERS
INTRODUCED ANTS &ORMICA OMNIVORA WHICH DID NOT REDUCE RAT NUMBERS BUT SOON
BECAME A PROBLEM THEMSELVES 4O REMOVE RATS AND ANTS TOGETHER IT WAS THEN
DECIDED TO INTRODUCE TOADS "UFO MARINUS "UT TOADS STILL DID NOT CONTROL RATS AND
BECAME A PEST THEMSELVES &INALLY SMALL )NDIAN MONGOOSES WERE INTRODUCED TO
CONTROL RATS AND TOADS -ONGOOSES FAILED TO CONTROL EITHER AND BEGAN PREYING ON
NATIVE BIRDS POSING NEW THREATS TO WILDLIFE 3ILVERSTEIN AND 3ILVERSTEIN
7E RECOGNIZE THAT THERE HAS BEEN MUCH TO GAIN FROM THESE KINDS OF HISTORICAL
MISTAKES AND THAT THESE HARD
WON LESSONS MIGHT EVEN TURN OUT TO BE GLOBALLY
BENElCIAL FOR THE BIODIVERSITY IN THE LONG RUN (OWEVER WE BELIEVE THE TRIAL
AND
ERROR APPROACH TIME HAS NOW PASSED AND BIODIVERSITY MANAGERS SHOULD
PERSEVERE IN THE CURRENT TREND OF BASING RESTORATION STRATEGIES ON SOUND SCIEN
TIlC GROUNDS 4HIS CAN ONLY BE DONE IN DEVELOPING SPECIlC STUDIES ON POPULATIONS
INTERACTING IN INVADED COMMUNITIES AS WELL AS THOSE THAT USE SUCH INVASIONS AS A
DIDACTIC TOOL TO UNDERSTAND BASIC PROCESSES IN POPULATION ECOLOGY
!#+./7,%$'-%.43
4HIS WORK WAS FUNDED BY THE )NSTITUT &RANAIS DE "IODIVERSIT 7E THANK ,EIGH
"ULL FOR HER HELP WITH THE %NGLISH
2%&%2%.#%3
!BRAMS 0 ! )NDIRECT INTERACTIONS BETWEEN SPECIES THAT SHARE A PREDATOR VARIETIES
OF INDIRECT EFFECTS )N 0REDATION $IRECT AND )NDIRECT )MPACTS ON !QUATIC #OMMUNITIES
EDS 7 # +ERFOOT ! 3IH PP
5NIVERSITY 0RESS OF .EW %NGLAND (ANOVER
!BRAMS 0 ! (OLT 2 $ AND 2OTH * $ !PPARENT COMPETITION OR APPARENT MUTUAL
ISM 3HARED PREDATION WHEN POPULATIONS CYCLE %COLOGY
$YNAMICS OF INTERACTING POPULATIONS
"ROTHERS . 0 "REEDING DISTRIBUTION AND STATUS OF BURROW
NESTING PETRELS AT
-ACQUARIE )SLAND !USTRIALIAN 7ILDLIFE 2ESEARCH
"ROTHERS . 0 AND #OPSON ' 2 -ACQUARIE )SLAND mORA AND FAUNA MANAGEMENT
INTERPRETING PROGRESS AND PREDICTIONS FOR THE FUTURE 0APERS AND 0ROCEEDINGS OF THE
2OYAL 3OCIETY OF 4ASMANIA
#AMPOS ! 2 AND 'RANADEIRO * 0 "REEDING BIOLOGY OF THE 7HITE
FACED 3TORM
PETREL ON 3ELVAGEM 'RANDE )SLAND .ORTH
EAST !TLANTIC 7ATERBIRDS
#HAPUIS *
, A !LIEN MAMMALS IN THE &RENCH 3UBANTARCTIC )SLANDS )N 0ROGRESS IN
CONSERVATION OF THE 3UBANTARCTIC )SLANDS ED 0 2 $INGWALL 6OL PP
)5#.
4HE 7ORLD #ONSERVATION 5NION
#ONSERVATION OF THE 3OUTHERN 0OLAR 2EGION
#HAPUIS *
, B 2ESTORATION OF TWO ISLANDS IN THE +ERGUELEN !RCHIPELAGO BY
ERADICATION OF THE RABBIT /RYCTOLAGUS CUNICULUS )N 0ROGRESS IN CONSERVATION OF THE
3UBANTARCTIC )SLANDS ED 0 2 $INGWALL 6OL PP
)5#.
4HE 7ORLD
#ONSERVATION 5NION
#ONSERVATION OF THE 3OUTHERN 0OLAR 2EGION
#HAPUIS *
, "OUSSS 0 AND "ARNAUD ' !LIEN MAMMALS IMPACT AND MANAGE
MENT IN THE &RENCH SUB
! NTARCTIC ISLANDS "IOLOGICAL #ONSERVATION
#LUTTON
"ROCK 4 ( AND #OULSON 4 #OMPARATIVE UNGULATE DYNAMICS THE DEVIL IS IN
THE DETAIL 0HILOSOPHICAL 4RANSACTIONS OF THE 2OYAL 3OCIETY OF ,ONDON 3ERIES "
"IOLOGICAL
3CIENCES
#OURCHAMP & #HAPUIS * , AND 0ASCAL - A -AMMAL INVADERS ON ISLANDS IMPACT
CONTROL AND CONTROL IMPACT "IOLOGICAL 2EVIEWS
#OURCHAMP & ,ANGLAIS - AND 3UGIHARA ' #ONTROL OF RABBITS TO PROTECT ISLAND
BIRDS FROM CAT PREDATION "IOLOGICAL #ONSERVATION
#OURCHAMP & ,ANGLAIS - AND 3UGIHARA ' 2ABBITS KILLING BIRDS MODELLING THE
HYPERPREDATION PROCESS *OURNAL OF !NIMAL %COLOGY
#OURCHAMP & 7OODROFFE 2 AND 2OEMER ' B 2EMOVING PROTECTED POPULATIONS TO
SAVE ENDANGERED SPECIES 3CIENCE
#UTHBERT 2 AND (ILTON ' )NTRODUCED HOUSE MICE -US MUSCULUS A SIGNIlCANT
PREDATOR OF THREATENED AND ENDEMIC BIRDS ON 'OUGH )SLAND 3OUTH !TLANTIC /CEAN
"IOLOGICAL #ONSERVATION
%BENHARD 4 )NTRODUCED BIRDS AND MAMMALS AND THEIR ECOLOGICAL EFFECTS 3WEDISH
7ILDLIFE 2ESEARCH
&EARE # !NTS TAKE OVER FROM RATS ON "IRD )SLAND 3EYCHELLES "IRD #ONSERVATION
)NTERNATIONAL
&ITZGERALD " - $IET OF DOMESTIC CATS AND THEIR IMPACT ON PREY POPULATIONS )N 4HE
DOMESTIC CAT THE BIOLOGY OF ITS BEHAVIOUR EDS $ # 4URNER 0 "ATESON PP
#AMBRIDGE 5NIVERSITY 0RESS #AMBRIDGE
&ITZGERALD " - $ANIEL - * &ITZGERALD ! % +ARL " * -EADS - * AND .OTMAN 0 2
&ACTORS EFFECTING THE NUMBERS OF HOUSE MICE -US MUSCULUS IN HARD BEECH
.OTHOFAGUS TRUNCATA FOREST *OURNAL OF THE 2OYAL 3OCIETY OF .EW :EALAND
&LUX * % # 2ELATIVE EFFECT OF CATS MYXOMATOSIS TRADITIONAL CONTROL OR COMPETITORS IN
REMOVING RABBITS FROM ISLANDS .EW :EALAND *OURNAL OF :OOLOGY
&LUX * % # AND &ULLAGAR 0 * 7ORLD DISTRIBUTION OF THE RABBIT /RYCTOLAGUS CUNICULUS
ON ISLANDS -AMMAL 2EVIEW
& #OURCHAMP AND 3 #AUT
'ARGOMINY / "OUCHET 0 0ASCAL - *AFFRE 4 AND 4OURNEUR * # #ONSEQUENCES
FOR BIODIVERSITY OF PLANT AND ANIMAL SPECIES INTRODUCTIONS IN .EW #ALEDONIA 2EVUE $
%COLOGIE ,A 4ERRE %T ,A 6IE
'ILLHAM - % 3OME INTERACTIONS OF PLANTS RABBITS AND SEABIRDS ON 3OUTH !FRICAN
ISLANDS *OURNAL OF %COLOGY
'INZBURG , 2 AND *ENSEN # 8 * 2ULES OF THUMB FOR JUDGING ECOLOGICAL THEORIES
4RENDS IN %COLOGY %VOLUTION
'RENFELL " 4 7ILSON + &INKENSTADT " & #OULSON 4 . -URRAY 3 !LBON 3 $
0EMBERTON * - #LUTTON "ROCK 4 ( AND #RAWLEY - * .OISE AND DETERMINISM
IN SYNCHRONIZED SHEEP DYNAMICS .ATURE
(OLDGATE - 7 AND 7ACE . - 4HE INmUENCE OF MAN ON THE mORAS AND FAUNAS OF
3OUTHERN )SLANDS 0OLAR 2ECORD
(OLT 2 $ 0REDATION APPARENT COMPETITION AND THE STRUCTURE OF PREY COMMUNITIES
4HEORETICAL 0OPULATION "IOLOGY
(ONE * 3PATIAL AND 4EMPORAL !SPECTS OF 6ERTEBRATE 0EST $AMAGE WITH %MPHASIS ON
&ERAL 0IGS *OURNAL OF !PPLIED %COLOGY
+ARL " * AND "EST ( ! &ERAL CATS ON 3TEWART )SLAND THEIR FOODS AND THEIR EFFECTS
ON KAKAPO .EW :EALAND *OURNAL OF :OOLOGY
+EANE 2 - AND #RAWLEY - * %XOTIC PLANT INVASIONS AND THE ENEMY RELEASE
HYPOTHESIS 4RENDS IN %COLOGY %VOLUTION
+ESSLER # # %RADICATION OF FERAL GOATS AND PIGS FROM 3ARIGAN )SLAND #OMMONWEALTH
OF THE .ORTHERN -ARIANA )SLANDS METHODS AND RESULTS )N %RADICATION OF ISLANDS
)NVASIVES PRACTICAL ACTIONS AND RESULTS ACHIEVED ED - #LOUT PP 5NIVERSITY OF
!UCKLAND .EW :EALAND
,E 2OUX 6 #HAPUIS * , &RENOT 9 AND 6ERNON 0 $IET OF THE HOUSE MOUSE -US
MUSCULUS ON 'UILLOU )SLAND +ERGUELEN ARCHIPELAGO 3UBANTARCTIC 0OLAR "IOLOGY
-ACK 2 . AND ,ONSDALE 7 - %RADICATING INVASIVE PLANTS HARD WON LESSONS
FOR ISLANDS )N 4URNING THE 4IDE THE %RADICATION OF )NVASIVE 3PECIES EDS # 2 6EITCH
- #LOUT )NVASIVE 3PECIES 3PECIALTY 'ROUP OF THE 7ORLD #ONSERVATION 5NION )5#.
!UCKLAND .EW :EALAND
-ICOL 4 AND *OUVENTIN 0 %RADICATION OF RATS AND RABBITS FROM 3AINT 0AUL )SLAND
&RENCH 3OUTHERN 4ERRITORIES )N 4URNING THE TIDE THE ERADICATION OF INVASIVE SPECIES
EDS # 2 6EITCH - . #LOUT PP )5#. 33# )NVASIVE 3PECIES 3PECIALIST
'ROUP )5#. 'LAND 3WITZERLAND AND #AMBRIDGE 5+
-URPHY % AND "RADlELD 0 #HANGE IN DIET OF STOATS FOLLOWING POISONING OF RATS IN A
.EW :EALAND FOREST .EW :EALAND *OURNAL OF %COLOGY
-YERS . -ITTERMEIER 2 ! -ITTERMEIER # ' DA &ONSECA ' ! " AND +ENT *
"IODIVERSITY HOTSPOTS FOR CONSERVATION PRIORITIES .ATURE
.EWMAN $ ' %FFECTS OF A -OUSE -US MUSCULUS %RADICATION 0ROGRAM AND (ABITAT
#HANGE ON ,IZARD 0OPULATIONS OF -ANA )SLAND .EW :EALAND WITH 3PECIAL 2EFERENCE TO
-CGREGOR 3KINK #YCLODINA MACGREGORI .EW :EALAND *OURNAL OF :OOLOGY
2AMMELL # ' (OOGENBOOM * * , #OTTER - 7ILLIAMS * - AND "ELL *
"RODIFACOUM RESIDUES IN TARGET AND NON TARGET ANIMALS FOLLOWING RABBIT POISONING TRIALS
.EW :EALAND *OURNAL OF %XPERIMENTAL !GRICULTURE
$YNAMICS OF INTERACTING POPULATIONS
2EID 7 6 "IODIVERSITY HOTSPOTS 4RENDS IN %COLOGY %VOLUTION
2OBERTSON ( ! (AY * 2 3AUL % + AND -C#ORMACK ' 6 2ECOVERY OF THE
+AKERORI AN %NDANGERED &OREST "IRD OF THE #OOK )SLANDS #ONSERVATION "IOLOGY
2OEMER ' 7 $ONLAN # * AND #OURCHAMP & 'OLDEN EAGLES FERAL PIGS AND INSU
LAR CARNIVORES (OW EXOTIC SPECIES TURN NATIVE PREDATORS INTO PREY 0ROCEEDINGS OF THE
.ATIONAL !CADEMY OF 3CIENCES OF THE 5NITED 3TATES OF !MERICA
3CHOENER 4 7 AND 3PILLER $ ! )NDIRECT EFFECTS IN AN EXPERIMENTALLY STAGED
INVASION BY A MAJOR PREDATOR !MERICAN .ATURALIST
3COTT * * 2ABBIT DISTRIBUTION HISTORY AND RELATED LAND DISTURBANCE -ACQUARIE )SLAND
!USTRALIA 0APERS AND 0ROCEEDINGS OF THE 2OYAL 3OCIETY OF 4ASMANIA
3ELKIRK 0 - #OSTIN ! " 3EPPELT 2 $ AND 3COTT * * 2ABBITS VEGETATION AND ERO
SION ON -ACQUARIE )SLAND 0ROCEEDINGS OF THE ,INNEAN 3OCIETY . 3 7
3ILVERSTEIN ! AND 3ILVERSTEIN 6 !NIMAL INVADERS 4HE STORY OF IMPORTED WILDLIFE
!THENUM .EW 9ORK
3MITH ! 0 AND 1UIN $ ' 0ATTERNS AND CAUSES OF EXTINCTION AND DECLINE IN
!USTRALIAN CONILURINE RODENTS "IOLOGICAL #ONSERVATION
3MITH 6 2 !VENANT . , AND #HOWN 3 , 4HE DIET AND IMPACT OF HOUSE MICE ON
A SUB !NTARCTIC ISLAND 0OLAR "IOLOGY
3OUL - % "OLGER $ 4 !LBERTS ! # 7RIGHT * 3ORICE - AND (ILL 3
2ECONSTRUCTED DYNAMICS OF RAPID EXTINCTIONS OF CHAPARRAL REQUIRING BIRDS IN URBAN
HABITAT ISLANDS #ONSERVATION "IOLOGY
4AYLOR 2 ( (OW THE -ACQUARIE ISLAND PARAKEET BECAME EXTINCT .EW :EALAND
*OURNAL OF %COLOGY
#HAPTER TWELVE
)NVASIBILITY OF SEED PREDATORS
ON SYNCHRONIZED AND
INTERMITTENT SEED PRODUCTION
OF HOST PLANTS
! 3ATAKE / . "JRNSTAD AND 9 )WASA
).42/$5#4)/.
4HE SPATIOTEMPORAL DISTRIBUTION OF RESOURCES THAT INVADERS REQUIRE FOR THEIR GROWTH
AND REPRODUCTION IS A KEY FACTOR IN CONTROLLING SUCCESS OF INVASION $AVIS ET AL
4HE ABUNDANCE OF ABIOTIC RESOURCES SUCH AS WATER SUPPLY AND NUTRIENT
LEVELS MAY mUCTUATE DUE TO VARIABLE CLIMATIC CONDITION AND DISTURBANCE REGIME
EG DROUGHT mOOD OR lRE REGIMES 4HEREFORE THE INTENSITY OF COMPETITION FOR
THESE LIMITED RESOURCES VARIES RESULTING IN TEMPORAL mUCTUATION OF A COMMUNITYS
SUSCEPTIBILITY TO INVASION $AVIS AND 0ELSOR
7HEN INVADING SPECIES ARE CONSUMERS OF BIOTIC RESOURCES EG HERVIVORES
AND SEED PREDATORS SUCCESSFUL INVASION MAY TRIGGER THE EVOLUTION OF TRAITS OF
HOST SPECIES THAT ENHANCE RESISTANCE TO INVADERS &OR EXAMPLE MAST SEEDING OR
MASTING THE INTERMITTENT AND SYNCHRONIZED REPRODUCTION BY PLANT POPULATIONS
+ELLY REPORTED ACROSS A BROAD GROUP OF TREE SPECIES (ERRERA ET AL
IS OFTEN EXPLAINED TO HAVE EVOLVED TO REDUCE SEED LOSSES TO AND HENCE TO PREVENT
-7 #ADOTTE ET AL EDS #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY n
3PRINGER 0RINTED IN THE .ETHERLANDS
! 3ATAKE /. "JRNSTAD AND 9 )WASA
INVASION OF SEED PREDATORS THE hPREDATOR SATIATION HYPOTHESISv *ANZEN
3ILVERTOWN %SSENTIALLY LARGER SEED CROPS ARE SYNCHRONIZED AMONG INDI
VIDUALS AND SATIATE SEED PREDATORS AND THEREFORE EXPERIENCE A LOWER PERCENTAGE
OF SEED PREDATION !T THE SAME TIME YEARS WITH LOW SEED PRODUCTION WILL REDUCE
THE GROWTH OF THE SEED PREDATOR POPULATIONS -ANY STUDIES CONlRM PREDATOR
SATIATION BY SHOWING LOWER PREDATION RATES IN HIGH SEED YEARS 3MITH ET AL
3ORK +ELLY AND 3ULLIVAN 3PERENS 3HIBATA ET AL +ELLY
ET AL -C+ONE ET AL 3ATAKE ET AL
4HE PROPERTIES OR ADAPTATIONS OF THE INVADING CONSUMERS ALSO PLAY AN IMPORTANT
ROLE PARTICULARLY THE TRAITS ASSOCIATED WITH DISPERSAL %VEN IF FOOD RESOURCES SHOW
UNPREDICTABLE mUCTUATION AT A SINGLE PATCH OR PLANT DISPERSAL BETWEEN PATCHES
BUFFERS VARIATION IN lTNESS BY SPREADING RISK OVER SPACE (OPPER $ISPERSAL
AMONG PATCHES IS ADVANTAGEOUS IF mUCTUATIONS OF LOCAL CONDITIONS ARE INDEPENDENT
ACROSS SPACE ,EVIN ET AL BUT IF THERE IS POSITIVE CORRELATION EITHER IN TIME OR
ACROSS SPACE DISPERSAL IS LESS FAVORED %LLNER AND 3HMIDA #OHEN AND ,EVIN
(OLT AND -C0EEK
)N THIS CHAPTER WE DISCUSS A RESOURCE CONSUMER MODEL TO UNDERSTAND THE
SPATIOTEMPORAL DYNAMICS OF A SEED PREDATOR ON A MASTING RESOURCE AND EXPLORE
THE INVASIBILITY OF THE PREDATORS 3ATAKE AND "JRNSTAD 3PATIOTEMPORAL
mUCTUATIONS IN SEEDING ARE REPRESENTED BY THE POLLEN COUPLED TREE MODEL
)SAGI 3ATAKE AND )WASA A B IN WHICH REPRODUCTIVE
BEHAVIOR OF INDIVIDUAL PLANTS IS CONSTRAINED BY THE LEVEL OF INTERNAL ENERGY RESERVES
AND LOCAL POLLEN PRODUCTION 7E CONSIDER SPECIALIZED INSECT SEED PREDATORS
4HE DYNAMICS OF THE PREDATOR AT A SINGLE HOST PLANT IS ASSUMED TO FOLLOW A SPA
TIALLY EXTENDED .ICHOLSON "AILEY MODEL (ASSELL ET AL 3ATAKE AND "JRNSTAD
AND THE ADULTS HAVING A SEMELPAROUS LIFECYCLE DISPERSE TO NEIGHBORING
HOST PLANTS TO SEARCH RANDOMLY FOR SEEDS BERRIES OR mOWERHEADS ON WHICH TO LAY
THEIR EGGS AND THE LARVAE SUBSEQUENTLY COMPLETE THE LIFECYCLE THROUGH FEEDING ON
THE SEEDS 7E CALCULATE AN APPROXIMATE INVASION CRITERION FOR THE PREDATORS
WHICH ILLUSTRATES HOW VARIABLE AND SYNCHRONIZED SEED PRODUCTION OF PLANTS MAY
PREVENT INVASION OF SEED PREDATORS POPULATION
2%3/52#% $9.!-)#3 /& 3%%$ 02/$5#4)/.
3PATIOTEMPORAL mUCTUATION OF FOOD RESOURCE FOR SEED PREDATORS IS MODELED AS FOL
LOWS 7E CONSIDER A FOREST OF OUT
CROSSING PLANTS EACH WITH AN INTERNAL ENERGY
RESERVE THAT CHANGES WITH NET ENERGY GAIN THROUGH PHOTOSYNTHESIS AND ENERGY
EXPENDITURE THROUGH REPRODUCTION AND MAINTENACEGROWTH %ACH PLANT GAINS
NET ENERGY 0S EVERY YEAR FROM PHOTOSYNTHESIS ,ET 3I T BE THE ENERGY RESERVES
OF A PLANT LOCATED AT SITE I AT THE BEGINNING OF YEAR T ! PLANT WILL NOT REPRODUCE
UNLESS ITS ACCUMULATED ENERGY RESERVES 3I T 0S EXCEEDS A CRITICAL THRESH
OLD ,4 /THERWISE THE PLANT PRODUCES mOWERS INVESTING ENERGY hmOWERING
COSTv PROPORTIONAL TO THE EXCESS GIVEN BY A3I T 0S n ,4 WHERE A IS CONSTANT
) NVASIBILITY OF SEED PREDATORS
OF PROPORTIONALITY 3ATAKE AND )WASA !LL mOWERS ARE POLLINATED AND
THE PLANT THEN INVESTS ADDITIONAL ENERGY DUE TO THE COST OF FRUIT PRODUCTION
2C A3I T 0S n ,4 WHERE 2C IS THE RATIO OF FRUITING COST TO mOWERING COST
4HE ENERGY RESERVE LEVEL THUS FALLS TO 3I T 0S n A2C 3I T 0S n ,4 SO
THAT THE OVERALL RESOURCE DYNAMICS ARE
3I T 0S IF 3I T 0S ) ,4
3I T
3I T 0S n A2C 3I T 0S n ,4 IF 3I T 0S ,4
#ONSIDERING THE NON DIMENSIONALIZED VARIABLE 9I T 3I T 0S n ,4 0 EQUATION
IS REWRITTEN AS
9I T IF 9I T )
9I T
n K9I T IF 9I T
IN WHICH K A2C n IS A DEPLETION CONSTANT )F A K IS EQUAL TO THE RATIO
OF FRUITING COST TO mOWERING COST 2C %QUATION REPRESENTS A hTENT MAPv THAT
PREDICTS STABLE ANNUAL REPRODUCTION FOR K AND CHAOTIC REPRODUCTION FOR K
3ATAKE AND )WASA 3UCH TEMPORAL ENERGY DYNAMICS AT THE LEVEL OF THE INDI
VIDUAL PLANT IS EASILY ILLUSTRATED THROUGH BIFURCATION DIAGRAMS &IG 4HE LONG
TERM TRAJECTORIES OF ENERGY LEVEL DElNED IN EQ ARE PLOTTED ACROSS A RANGE OF
VALUES OF K 7HEN K TRAJECTORIES VISIT A SINGLE POSITIVE VALUE WHICH REPRESENTS
ANNUAL AND CONSTANT REPRODUCTION &OR K THE ORBIT EVENTUALLY REPRESENTS
CHAOTIC mUCTUATION !S K INCREASES THE INTERVAL BETWEEN NON REPRODUCTIVE YEARS
BECOMES LONGER BECAUSE THE PLANT NEEDS MORE YEARS TO REBUILD ENERGY RESERVES TO
THE THRESHOLD LEVEL
0OLLEN LIMITATION
)F PLANTS ARE SELF INCOMPATIBLE FRUITING RATES MAY DEPEND ON THE mOWERING ACTIVITY
OF OTHER TREES IN A FOREST BECAUSE POLLINATION EFlCIENCY CHANGES WITH THE NUMBER
OF mOWERING PLANTS .ILSSON AND 7STLJUNG 3MITH ET AL 4O MODEL THE
POLLEN LIMITATION OF PLANT REPRODUCTION THE POLLEN AVAILABILITY 0I T TO INDIVIDUAL
I WAS INTRODUCED IN THE NON DIMENSIONALIZED ENERGY DYNAMICS )SAGI ET AL
3ATAKE AND )WASA
9I T IF 9 T )
9I T I A
n K0I T 9I T IF 9I T
WHERE 0I T IS DETERMINED BY THE AVERAGE mOWERING INTENSITY OF THE PLANTS IN THE
NEIGHBORHOOD AROUND THE FOCAL PLANT 5I )T IS GIVEN BY
! 3ATAKE /. "JRNSTAD AND 9 )WASA
&IG "IFURCATION DIAGRAM OF AN INDIVIDUAL PLANT (ORIZONTAL AXIS IS DEPLETION
COEFlCIENT K AND THE VERTICAL AXIS IS THE NON
DIMENSINALIZED ENERGY LEVEL 0ARAMETERS
` W
`
0I T - 9J T B
W J D5I
WHERE 9 9 IF 9 AND ZERO OTHERWISE 4HE PARAMETER W IS THE NUMBER OF
PLANTS INCLUDED IN THE NEIGHBORHOOD 5I NOTE THAT THE NEIGHBORHOOD DOES NOT
INCLUDE ONESELF AND ` DETERMINES THE DEGREE OF NEIGHBOR DEPENDENCE ON THE POL
LEN LIMITATION )F `IS CLOSE TO FRUIT PRODUCTION IS ALMOST INDEPENDENT OF THE mOW
ERING INTENSITY OF OTHER PLANTS )N CONTRAST A LARGE ` IMPLIES A STRONG DEPENDENCE
OF SEED AND FRUIT PRODUCTION ON THE POLLEN PRODUCTION OF OTHER PLANTS 4HUS WE CALL
` THE POLLEN COUPLING STRENGTH )N THIS CHAPTER WE CONSIDER A SITUATION OF hLOCAL
POLLEN COUPLINGv IN WHICH POLLINATION IS LIMITED TO THE EIGHT NEAREST NEIGHBORS
AROUND THE FOCAL PLANT W
0/05,!4)/. $9.!-)#3 /& 3%%$ 02%$!4/23
! SPECIALIZED INSECT SEED PREDATOR WITH A SEMELPAROUS LIFECYCLE IS CONSIDERED IN AN
ALL
FEMALES MODEL !DULTS DISPERSE TO NEARBY HOST PLANTS TO SEARCH RANDOMLY FOR
SEEDS AND BERRIES ON WHICH TO LAY THEIR EGGS 4HE LARVAE HATCH TO COMPLETE THEIR
LIFECYCLE FEEDING ON THE SEED LEAVE THE SEEDS TO PUPATE AND EMERGE AS ADULTS AT THE
BEGINNING OF NEXT GENERATION 4HESE ASSUMPTION ARE MOTIVATED BY STUDIES OF THE
APPLE FRUIT MOTH !RGYRESTHIA CONJUGELLA :ELLER WHICH IS A PRE
DISPERSAL SEED PREDA
TOR OF ROWAN 3ORBUS AUCUPARIA , 4HE LARVAE OF THE APPLE FRUIT MOTH RELIES HEAVILY
) NVASIBILITY OF SEED PREDATORS
ON ROWAN BERRIES THAT VARIES SUBSTANTIALLY BETWEEN YEARS IN A GEOGRAPHICALLY SYN
CHRONIZED FASHION +OBRO ET AL 3ATAKE ET AL
7E ASSUME THAT HOST PLANTS ARE ARRANGED ON LATTICE POINTS OF A TWO
DIMENSIONAL
SQUARE GRID 4HE NUMBER OF EMERGED ADULTS AT SITE I AT THE BEGINNING OF YEAR T IS
DENOTED BY :I T ! FRACTION + OF THE ADULTS LEAVES THE PLANT WHERE THEY WERE
BORN AND DISPERSES TO ADJACENT PLANTS 4HE POST
DISPERSAL NUMBER OF ADULTS AT
SITE I DENOTED AS :I T IS GIVEN BY
+
:I T n + :I T - :J T
W J D2I
4HE PARAMETER 2I IS THE NEIGHBORHOOD OF SITE I AND W IS THE NUMBER OF HOST
PLANTS INCLUDED IN 2I &OR SIMPLICITY WE ALSO ASSUME THAT SEED PREDATOR DISPERSES
TO THE EIGHT NEAREST NEIGHBORS W 7HEN A SINGLE PLANT AT SITE I PRODUCES A
CERTAIN AMOUNT OF SEEDS &I T IN YEAR T THE NUMBER OF PRE
DISPERSAL ADULTS IN THE
NEXT GENERATION T IS GIVEN BY
:I T a&IT n E n_:IT
WHERE a IS THE PER CAPITA EMERGENCE RATE AND _ IS THE PER CAPITA ATTACK RATE THE
SEARCHING EFlCIENCY &I T ITSELF IS GOVERNED BY THE POLLEN COUPLED TREE MODEL
EQUATIONS n ACCORDING TO
IF 9 T )
&I T I
CK0I T 9I T IF 9I T
WHERE C IS A CONSTANT K IS THE DEPLETION COEFlCIENT AND 0I T IS THE POLLEN AVAIL
ABILITY GIVEN BY EQN B
3%%$ 02%$!4)/. /. -!34).' 2%3/52#%
4HE REPRODUCTIVE BEHAVIOR PREDICTED BY THE POLLEN COUPLED TREE MODEL AND SEVERITY
OF SEED PREDATION IS ILLUSTRATED IN &IG 7HEN THE DEPLETION COEFlCIENT K IS LESS
THAN EACH PLANT PRODUCES SEEDS EVERY YEAR WHICH IS ANNUAL REPRODUCTION &IG
! !NNUAL REPRODUCTION CAN BE FURTHER CLASSIlED INTO hANNUAL AND CONSTANT
REPRODUCTIONv SHADED AREA IN &IG ! IN WHICH EACH PLANT PRODUCES SEEDS CON
STANTLY AND hANNUAL AND
YEAR CYCLIC REPRODUCTIONv CHECKER REGION IN &IG !
IN WHICH PRODUCED SEED CROP SIZE ALTERNATES BETWEEN HIGH AND LOW BUT POSITIVE
REPRODUCTIVE LEVELS !NNUAL AND CONSTANT SEED PRODUCTION RESULTS IN SEVERE ATTACKS
&IG " WHILE ANNUAL AND
YR CYCLIC SEED PRODUCTION CAN CAUSE EXTINCTION OF THE
PREDATOR &IG " 4HIS EXTINCTION IS DUE TO THE SATIATION OF NUMERICAL RESPONSE OF
! 3ATAKE /. "JRNSTAD AND 9 )WASA
&IG ! #LASSIlCATION OF REPRODUCTIVE PATTERN OF PLANTS PREDICTED BY THE POLLEN
COUPLED TREE MODEL WITH AN ASSUMPTION OF LOCAL POLLEN COUPLING 7E CLASSIlED FOUR TYPES OF
REPRODUCTIVE PATTERN AS EXPLAINED IN THE TEXT " 4HE DENSITY PLOT FOR PROPORTION OF SEEDS
ATTACKED CALCULATED BY NUMERICAL SIMULATION OF THE MODEL 4HE DARKER AREA REPRESENTS MORE
SEVERE SEED PREDATION 0ARAMETERS + _ &ROM ! 3ATAKE AND / . "JRNSTAD
3PATIAL DYNAMICS OF SPECIALIST SEED PREDATORS ON SYNCHRONIZED AND INTERMITTENT SEED
PRODUCTION OF HOST PLANTS !MERICAN .ATURALIST
2EPRINTED WITH
PERMISSION FROM 4HE 5NIVERSITY OF #HICAGO 0RESS
THE PREDATOR THE NUMBER OF THE PREDATORS IS HEAVILY REDUCED BY A VERY SMALL
SEED PRODUCTION IN A LOW REPRODUCTIVE YEAR AND THE PREDATOR POPULATION CANNOT
COPE WITH A GREAT INCREASE IN SEED PRODUCTION IN A HIGH REPRODUCTIVE YEAR WHICH
DECREASES THE POPULATION SIZE AND lNALLY LEADS THE POPULATION TO EXTINCTION
7HEN INDIVIDUAL PLANTS PRODUCE SEEDS INTERMITTENTLY WITH NUMEROUS YEARS WITH
NO SEED SET PARAMETER REGION OF K IN &IG ! hLOCALv EXTINCTION OF SEED PREDA
TORS IS INEVITABLE (OWEVER THEY MAY NEVERTHELESS PERSIST GLOBALLY DUE TO DISPERSAL
WHEN MAST SYNCHRONY IS WEAK &IG )N &IG SIGNIlCANT SYNCHRONY l * IS
DISTINGUISHED FROM WEAK SYNCHRONY l BY CALCULATING THE DEGREE OF SPATIAL
SYNCHRONY AMONG PLANTS l )N A FOREST EXHIBITING WEAK SYNCHRONY IN SEEDING
THERE IS ALWAYS SOME FRACTION OF PLANTS PRODUCING SEEDS 4HIS ENABLES SEED PREDA
TORS TO PERSIST GLOBALLY THROUGH DISPERSAL TO ANY ASYNCHRONOUS NEARBY PLANTS
) NVASIBILITY OF SEED PREDATORS
)N CONTRAST IF PLANTS IN A FOREST PRODUCE SEEDS IN A SIGNIlCANTLY SYNCHRONIZED FASH
ION SEED PREDATORS CANNOT lND RESOURCES ELSEWHERE WITHIN THE DISPERSAL RANGE
AND THEN GO GLOBALLY EXTINCT &IG &OR MASTING PLANTS K THE PROPORTION
OF ATTACKED SEEDS IS INVERSELY RELATED TO THE DEGREE OF SPATIAL SYNCHRONY AMONG
PLANTS l &IG MEANING THAT SPATIAL SYNCHRONY IN SEED PRODUCTION EFFECTIVELY
REDUCES LOSSES TO SEED PREDATORS
&IG ! $EGREE OF SYNCHRONY IN SEED PRODUCTION l " 0ROPORTION OF SEEDS ATTACKED
4HE HORIZONTAL AXIS IS THE DEPLETION COEFlCIENT K 0ARAMETERS ` _
).6!3)/.
7E DEVELOPED AN APPROXIMATE INVASION CRITERION 1 THAT APPROXIMATELY DESCRIBES
WHEN SEED PREDATORS CAN INVADE A SYSTEM WHERE THE HOST PLANTS PRODUCE SEEDS
SYNCHRONOUSLY AND INTERMITTENTLY SEE !PPENDIX IN 3ATAKE AND "JRNSTAD
4HOUGH WE HAVE BEEN UNABLE TO OBTAIN EXACT INVASION CRITERIA DUE TO THE NATURE
OF MASTING RESOURCES IE mUCTUATION ACROSS TIME AND SPACE IN A C YCLIC OR CHAOTIC
MANNER 3ATAKE AND )WASA A 4HE APPROXIMATE INVASION CRITERION
! 3ATAKE /. "JRNSTAD AND 9 )WASA
APPEARS TO GIVE A GOOD PREDICTION OF THE INVASION AND PROVIDES KEY INSIGHTS INTO
HOW PREDATOR INVASION IS INmUENCED BY THE MANNER OF mUCTUATIONS OF THE RESOURCE
4HE APPROXIMATE INVASION CRITERION 1 IS THE AVERAGE GROWTH OR DECLINE IN THE PRED
ATOR POPULATION DURING TWO GENERATIONS FOLLOWING INTRODUCTION GIVEN AS FOLLOWS
4 n
1 - 1 T A
4 T
WHERE
_ a . + .
1T n + - &I T &IT - - &I T &JT B
. I
W I J D2I
)N THE ABOVE EQUATION . IS THE NUMBER OF PLANTS IN THE FOREST AND W IS THE
NUMBER OF PLANTS INCLUDED IN THE NEIGHBORHOOD 2I &I T IS THE AMOUNT OF SEED
PRODUCED BY A PLANT AT SITE I IN YEAR T GIVEN BY EQN /THER PARAMETERS ARE AS
DElNED IN EQUATIONS AND 4HE lRST TERM IN THE RIGHT HAND SIDE IN EQN B
REPRESENTS THE AVERAGE POPULATION INCREASE OR DECLINE THROUGH TWO GENERATIONS
FOR THE SEED PREDATORS THAT DO NOT DISPERSE THEIR NEXT
YEAR OFFSPRING ARE LEFT TO
CONSUME SEEDS PRODUCED BY THE PLANT WHERE THEIR PARENTS WERE BORN 4HE SECOND
TERM REPRESENTS THE CORRESPONDING AVERAGE FOR THE PREDATORS THAT DISPERSED TO LAY
EGGS ON NEIGHBORING PLANTS
"Y A SIMPLE CALCULATION 3TAKE AND "JORNSTAD 1 IS RELATED TO THE TIME
LAG
AUTO
CORRELATION 2 IN SEED PRODUCTION OF INDIVIDUAL PLANTS AND TIME LAG
CROSS
CORRELATION # BETWEEN NEIGHBORING PLANTS AS FOLLOWS
1 _ a ! n + 2 m +# m & #
WHERE m IS THE VARIANCE AND & IS THE TEMPORAL MEAN SEEDSET OF EACH PLANT %QN
SHOWS THAT INVASIBILITY IS PROMPTED IF SEED PRODUCTION IS POSITIVELY CORRELATED
IN TIME BOTH FOR A GIVEN PLANT AND FOR PLANTS WITHIN THE DISPERSAL NEIGHBORHOOD OF
THE PREDATOR )N THE CASE OF MAST SEEDING NEGATIVE TIME LAG OR AUTOCORRELATIONS
IN SEED PRODUCTION ARE OFTEN REALIZED .ORTON AND +ELLY 3ORK ET AL
+OENIG ET AL +ELLY AND 3ORK )N ADDITION NEGATIVE CROSS CORRELATION
AT TIME LAG OFTEN RESULT BECAUSE OF SPATIALLY SYNCHRONIZED REPRODUCTION 4HUS
INVASIBILITY TO THE PREDATORS IS REDUCED ON MASTING RESOURCES
)F SEED PRODUCTION WITH INTERMITTENCE IS SYNCHRONIZED AT A hLOCALv SPATIAL SCALE
NEARBY PLANTS SHOW A STRONGER NEGATIVE CROSS CORRELATION IN SEED PRODUCTION
AT TIME LAG THAN DISTANTLY LOCATED PLANTS )N A FOREST SHOWING REGIONALLY NOT
LOCALLY SELF
ORGANIZED SEED PRODUCTION CROSS CORRELATIONS CALCULATED AT THE LOCAL
SPATIAL SCALE WILL BE SIMILAR TO THE AVERAGE ACROSS THE REGIONAL SPATIAL SCALE )N THIS
CASE 1T SIMPLIlES TO
) NVASIBILITY OF SEED PREDATORS
_a . + .
1T n + - &I T &IT - - &I T &JT
I
. . n I J &I
(ERE THE SECOND TERM IN THE RIGHT HAND SIDE IN EQN B IS REPLACED BY THE
AVERAGE OF THE PRODUCT FOR ALL POSSIBLE PAIRS I J OF PLANTS "Y TAKING THE TEMPORAL
AVERAGE OF 1T WE HAVE AN APPROXIMATE INVASION CRITERION IN THE ABSENCE OF LOCAL
SPATIAL STRUCTURE 1
7E EVALUATED 1 AND 1 NUMERICALLY BY GENERATING THE SPATIOTEMPORAL PATTERNS
IN SEED PRODUCTION FROM A NUMERICAL SIMULATION OF THE POLLEN
COUPLED TREE MODEL
WITH A RANGE OF PARAMETERS RESULTING IN VARYING DEGREE OF MASTING 3ATAKE AND
"JORNSTAD 7E CONlRM THE ACCURACY OF THE APPROXIMATE INVASION CRITERIA
THROUGH NUMERICAL SIMULATION OF THE FULL CONSUMER
RESOURCE MODEL )N THEORY
THE SEED PREDATORS SHOULD ONLY BE ABLE TO INVADE WHEN 1 4O INVESTIGATE THIS
WE INTRODUCED A MINUTE DENSITY OF THE PREDATOR ON EACH HOST PLANT IN THE
MODEL 7E SUBSEQUENTLY TRACED THEIR POPULATION GROWTH FOR GENERATIONS )F
THE PREDATORS POPULATION IS STILL EXTANT AT THE END OF THE SIMULATION THE PLANT
REPRODUCTIVE STRATEGY IS CONSIDERED INVASIBLE
4HE RESULTS INCLUDING THE TWO APPROXIMATE INVASION CRITERIA ARE ILLUSTRATED IN
&IG "OTH CRITERIA ARE SEEN TO DECREASE WITH INCREASING SYNCHRONY OF SEED SET l
MEANING THAT INVASIBILITY IS INHIBITED PARTLY OR ENTIRELY WHEN PLANTS SHOW REPRO
DUCTIVE SYNCHRONY )N SIMULATIONS ESTABLISHMENT FAILS WHEN 1 IS BELOW AND
&IG 0LOT OF THE APPROXIMATE INVASION CRITERIA 1 OPEN SQUARES AND 1 SOLID CIRCLES
ALONG THE DEGREE OF SYNCHRONY IN SEED PRODUCTION AMONG DIFFERENT PLANTS l 3TARS REPRESENT
THE VALUES OF 1 IN WHICH EXTINCTION OF SEED PREDATORS OCCURRED 0ARAMETERS
_ &ROM ! 3ATAKE AND / . "JRNSTAD 3PATIAL DYNAMICS OF SPECIALIST SEED PREDATORS
ON SYNCHRONIZED AND INTERMITTENT SEED PRODUCTION OF HOST PLANTS !MERICAN .ATURALIST
2EPRINTED WITH PERMISSION FROM 4HE 5NIVERSITY OF #HICAGO 0RESS
! 3ATAKE /. "JRNSTAD AND 9 )WASA
&IG 3PATIAL PATTERNS IN SEED PRODUCTION UPPER ROW AND AMOUNT OF SEED PREDATORS
BOTTOM ROW X AND Y
AXES REPRESENT SPATIAL LOCATIONS AND Z
AXIS REPRESENTS SEED CROP
SIZE UPPER THREE lGURES OR SEED PREDATOR ABUNDANCE BOTTOM THREE lGURES 4IME mOWS
LEFT TO RIGHT 0ARAMETERS K ` _ 4HE TOTAL NUMBER OF HOST PLANT
IS )N ORDER TO SHOW CLEAR SPATIAL PATTERNS WE APPLY THE POLLEN DISPERSAL RANGE
LARGER THAN NEAREST NEIGHBOR PLANTS LOCATED AT ITH LOCATION XI YI AND JTH LOCATION XJ
YJ CAN EXCHANGE POLLEN IF \XI XJ\ \YI YJ\ ) 4HE OVERALL SPATIAL PATTERN IS
SIMILAR BUT THE SPATIAL SCALE OF THE PATTERN IS LARGER AND BECOMES COARSE GRAINED AS POLLEN
DISPERSAL RANGE INCREASES FOR DETAILS SEE 3ATAKE AND )WASA A &ROM ! 3ATAKE AND
/ . "JRNSTAD 3PATIAL DYNAMICS OF SPECIALIST SEED PREDATORS ON SYNCHRONIZED AND
INTERMITTENT SEED PRODUCTION OF HOST PLANTS !MERICAN .ATURALIST
2EPRINTED WITH PERMISSION FROM 4HE 5NIVERSITY OF #HICAGO 0RESS
SUCCEEDS FOR 1 4HIS SUGGESTS THAT THE APPROXIMATE INVASION CRITERIA BASED ON
THE POPULATION GROWTH DURING TWO GENERATIONS APPEARS TO GIVE AN EXCELLENT PREDIC
TION OF THE INVASION IN THE FULL SPATIOTEMPORAL SIMULATIONS 4HIS SUCCESS ORIGINATES
IN PART FROM HOW THE ENTIRE FOREST EXHIBITS A
YEAR CYCLE IN SEED PRODUCTION WHEN
THE PLANTS REPRODUCTION IS HIGHLY SYNCHRONIZED &IG 4HE
YEAR CYCLIC SEED
PRODUCTION BECOMES UNCLEAR AT THE POPULATION LEVEL AS THE DEGREE OF SYNCHRONY
DECREASES 4HUS LONGER INTERVALS BETWEEN MAST YEARS EG OR YEARS SHOULD BE
TAKEN INTO ACCOUNT TO CORRECT THE SLIGHT OVERESTIMATION OF THE INVASION CRITERION
&IG )NTERESTINGLY 1 IS ALMOST ALWAYS LESS THAN 1 4HIS DIFFERENCE HIGHLIGHTS
) NVASIBILITY OF SEED PREDATORS
THAT SEED PREDATORS SUFFER LOWER INVASIBILITY WHEN REPRODUCTION AMONG PLANTS IS
SYNCHRONIZED AT A LOCAL SPATIAL SCALE
4HE SPATIAL PERSISTENCE OF PREDATORS IN THE PRESENCE OF HIGHLY VARIABLE MAST
SEEDING IS VISUALIZED IN &IG 4HE UPPER THREE lGURES REPRESENT THE TEMPORAL
CHANGE OF SPATIAL DISTRIBUTIONS FOR SEED CROP SIZE PRODUCED BY INDIVIDUAL PLANTS
4HE lGURE ILLUSTRATES HOW LOCALIZED SPATIAL CLUSTERS CAN RESULT FORM LOCAL POLLEN
COUPLING 7ITHIN A GIVEN CLUSTER PLANTS SHOW SYNCHRONIZED REPRODUCTION WITH
A
YEAR CYCLE VISITING HIGH AND VERY LOW REPRODUCTIVE LEVEL 4HE CORRESPONDING
SPATIAL DISTRIBUTIONS OF SEED PREDATORS BOTTOM PANEL IN &IG SHOW HOW THE SEED
PREDATORS PERSISTENCE IS FACILITATED AT THE BOUNDARIES OF THE CLUSTERS 3EED PREDA
TORS CAN lND RESOURCES TO CONSUME ONLY WHEN HOST PLANTS SHOWING OPPOSITE REPRO
DUCTIVE PHASES ARE WITHIN THE DISPERSAL RANGE
#/--5.)49 ,%6%, -!34).' !.$ ).6!3)/.
)N THIS SECTION WE INTRODUCE A SIMPLE CASE STUDY OF PLANT CONSUMER SYSTEM AND
BRIEmY DISCUSS THAT WHEN SEED PREDATORS ARE GENERALISTS MASTING AT COMMUNITY
LEVEL IS NECESSARY TO REDUCE THE IMPACT OF THE PREDATOR 2OWAN 3ORBUS AUCUPARIA ,
IS A DECIDUOUS TREE WHICH IS COMMON IN MUCH OF %UROPE 4UTIN ET AL 4HE
mESHY BERRIES PRODUCED BY ROWAN TREES SUFFER FROM ATTACK BY THE APPLE FRUIT MOTH
!RGYRESTHIA CONJUGELLA THAT IS A PRIMARY PRE DISPERSAL SEED PREDATOR WHOSE LARVAE
RELY HEAVILY ON ROWAN BERRIES 3PERENS A B +OBRO ET AL 3ATAKE
ET AL 4HE SPATIOTEMPORAL DATA OBTAINED FROM A CENSUS IN SOUTHERN .ORWAY
SHOW THAT BERRY PRODUCTION IN ROWAN IS VARIABLE ACROSS YEARS MEAN #6
AND IS SPATIALLY SYNCHRONIZED l BOOTSTRAPPED CONlDENCE INTERVAL
4HE DEGREE OF SPATIAL SYNCHRONY IN ROWAN POPULATION IS LOWER THAN
THE LEVEL OF SYNCHRONY LEADING TO SEED PREDATOR EXTINCTION IE l &IG AND
HENCE THE APPLE FRUIT MOTH SUCCESSFULLY INVADE AND IS PERSISTENT IN THIS SYSTEM
(OWEVER RELATIVELY HIGH DEGREE OF SPATIAL SYNCHRONY IN ROWAN MASTING EFl
CIENTLY REDUCES SEED LOSSES TO THE APPLE FRUIT MOTH AND SUPPRESSES THE POPULATION
GROWTH RATE OF SEED PREDATORS AT A LOW LEVEL AS ILLUSTRATED IN &IG 4HE PROPORTION
OF SEEDS PREDATED WAS NEGATIVELY CORRELATED WITH THE TOTAL NUMBER OF SEEDS EVI
DENCING A SATIATED FUNCTIONAL RESPONSE OF THE SEED PREDATORS BY MASTING )N ADDI
TION A NEGATIVE RELATIONSHIP WAS OBSERVED BETWEEN THE SEED PREDATION AND THE
RATIO OF BERRY PRODUCTION IN SUCCESSIVE YEARS 4HIS MEANS THAT SEED CROPS FOLLOW
ING POOR SEED CROPS TENDED TO ESCAPE PREDATIONS TESTIFYING TO A SATIATION THROUGH
NUMERICAL RESPONSE OF THE PREDATORS 3ATAKE ET AL 4HE PREDATOR SATIATION
HYPOTHESIS FURTHER PREDICTS THAT THE MORE VARIABLE POPULATIONS SHOULD ATTAIN
HIGHER OVERALL SEED SURVIVAL *ANZEN 7ALLER 3ILVERTOWN
4O ASSESS THIS THE TOTAL PROPORTION OF SEEDS LOST TO PREDATION WAS EXAMINED
AS A FUNCTION OF TEMPORAL VARIABILITY IN SEED PRODUCTION MEASURED BY THE #6
4HE RESULTANT LOGISTIC REGRESSION REVEALED A SIGNIlCANTLY NEGATIVE RELATIONSHIP
BETWEEN THE PROPORTION OF BERRIES PREDATED AND #6 THE SLOPE n P
! 3ATAKE /. "JRNSTAD AND 9 )WASA
&IG 4IME SERIES DATA OF ! ANNUAL BERRY PRODUCTION OF ROWAN TREES AND " PERCENTAGE
OF ATTACKED BERRIES BY THE APPLE FRUIT MOTH %ACH LINE REPRESENTS THE TIME SERIES FROM ONE
STUDY SITE &ROM ! 3ATAKE / . "JRNSTAD AND 3 +OBRO -ASTING AND TROPHIC CASCADES
INTERPLAY BETWEEN ROWAN TREES APPLE FRUIT MOTH AND THEIR PARASITOID IN SOUTHERN .ORWAY
/IKOS 2EPRINTED WITH PERMISSION FROM "LACKWELL 0UBLISHING
INDICATING THAT LARGER INTER
ANNUAL VARIATION IN BERRY PRODUCTION RESULTED IN
SMALLER LOSSES TO PREDATION 4HESE RESULTS SHOW THAT ROWAN MASTING HAS AN ADAP
TIVE FOUNDATION WHICH REDUCE SEED LOSSES TO AND PREVENT A RAPID EXPANSION IN A
PLANT POPULATION
4HE PERSISTENCE OF THE MOTH IN THE ROWAN
MOTH SYSTEM IS IN PART BECAUSE OF
THE FACT THAT THE APPLE FRUIT MOTH IS NOT A STRICT SPECIALIST THE PRIMARY AND PRE
FERRED HOST OF THE APPLE FRUIT MOTH IS ROWAN BUT WHEN TOO FEW ROWAN BERRIES ARE
AVAILABLE FOR EGG
LAYING MANY FEMALE MOTHS SWITCH THE HOST TO APPLE !HLBERG
)F THERE ARE GENERALIST SEED PREDATORS IN A PLANT COMMUNITY VARIABLE AND
SYNCHRONIZED SEED PRODUCTION ONLY WITHIN A SINGLE SPECIES MAY NOT ENOUGH TO
PREVENT INVASION OF THE SEED PREDATOR AND EVEN MASTING SPECIES SHOWING A SIGNIl
CANT SYNCHRONY IN SEED PRODUCTION MAY SUFFER HEAVY SEED PREDATION 3ILVERTOWN
) NVASIBILITY OF SEED PREDATORS
&IG 0LOT OF PROPORTION OF SEED PREDATION IN ROWAN AS A FUNCTION OF ! SEED CROP SIZE
AND " THE RATIO OF SEED CROP SIZE IN YEAR T AND THAT IN YEAR T n LAG ,OG TRANSFORMED
VARIABLES ARE USED FOR CALCULATION ,INES REPRESENTS THE LOGISTIC REGRESSION MODEL WHICH
HAVE INTERCEPTS AND SLOPES OF ! 3% P n 3%
P AND " n 3% P n 3% P
# 2ELATIONSHIP BETWEEN THE COEFlCIENT OF VARIATION #6 OF ANNUAL SEED PRODUCTION
AND THE TOTAL PROPORTION OF SEED PREDATION %ACH POINT REPRESENTS THE RESULT CALCULATED AT
ONE STUDY SITE ,OGISTIC REGRESSION ANALYSIS GAVE THE REGRESSION LINE AS INTERCEPT OF n
3% P AND SLOPE OF n 3% P
! 3ATAKE /. "JRNSTAD AND 9 )WASA
STATED THAT SYNCHRONOUS SEED PRODUCTION BETWEEN POPULATION OF DIFFERENT
SPECIES SHARING THE SAME SEED PREDATORS REDUCES THE GROWTH RATE OF THE PREDATOR
POPULATION BY PREDATOR SATIATION 3PATIAL SYNCHRONY IN SEED PRODUCTION AMONG
SPECIES HAS BEEN REPORTED IN ANIMAL
POLLINATED SPECIES IN A RELATIVELY CLOSED COM
MUNITY IN THE RAIN FOREST -OMOSE ET AL )NOUE AND (AMID 3AKAI
ET AL AND IN WIND
POLLINATED SPECIES IN
OPEN HABITATS SUCH AS TEMPERATE
FOREST AND SAVANNA +OENIG AND +NOPS 3CHAUBER ET AL
3HIBATA ET AL 3UCH A SYNCHRONIZED AND EPISODIC REPRODUCTION IN A
COMMUNITY PROVIDES AN INTERESTING INSIGHT INTO THE EVOLUTIONARY PROCESS
HOW PLANT
COMMUNITIES ENHANCE RESISTANCE AGAINST GENERALIST SEED PREDATORS
#/.#,53)/.
4HIS CHAPTER FOCUSES ON THE SPATIALLY
EXTENDED DYNAMICS OF MASTING PLANTS AND
THEIR SEED PREDATORS 0LANTS WITHIN A FOREST MAY PRODUCE SEEDS ANNUALLY OR INTER
MITTENTLY IN TIME AND SYNCHRONOUSLY OR ASYNCHRONOUSLY ACROSS SPACE 4HIS RANGE
OF DYNAMICAL BEHAVIOR IS CAPTURED BY THE POLLEN COUPLED TREE MODEL EQNS
)SAGI 3ATAKE AND )WASA A B 4HE EFFECT OF MAST SEEDING
ON DYNAMICS AND INVASIBILITY OF SPECIALIST CONSUMERS WAS EXPLORED BY EXAMINING
THE POPULATION DYNAMICS OF SEED PREDATORS IN A BOTTOM
UP FASHION EQNS
7HEN PLANTS PRODUCE SEEDS INTERMITTENTLY PERSISTENCE AND INVASION OF THE
PREDATOR IS MOST CRITICALLY DEPENDENT ON THE DEGREE OF SYNCHRONY OF SEED SET 4HE
PREDATOR POPULATION CAN ONLY PERSIST THROUGH DISPERSAL TO ADJACENT HOST PLANTS IN
A FOREST SHOWING WEAK SYNCHRONY BECAUSE THERE IS THEN SOME FRACTION OF ASYN
CHRONOUSLY REPRODUCING PLANTS WITHIN THE DISPERSAL RANGE OF THE PREDATORS )N
CONTRAST EXTINCTION OF SEED PREDATORS IS LIKELY WHEN INTERMITTENT REPRODUCTION IS
HIGHLY SYNCHRONIZED AMONG DIFFERENT PLANTS &IG 3PATIAL SYNCHRONY IN SEED
PRODUCTION WITH INTERMITTENCE CREATES NEGATIVE TIME LAG
AUTO
CORRELATION AND
CROSS
CORRELATION IN SEEDING WHICH MOST EFFECTIVELY REDUCE INVASIBILITY OF THE SEED
PREDATORS EQN 4HIS MEANS THAT hCLASSICv MAST SEEDING EXHIBITING SEED SET THAT
IS NEGATIVELY CORRELATED IN TIME BUT POSITIVELY CORRELATED ACROSS SPACE IS A GOOD
STRATEGY TO REDUCE SEED LOSS TO SEED PREDATORS )N ADDITION ANALYSIS OF INVASION
CRITERION SUGGEST THAT SPATIAL SYNCHRONY AT LOCAL SPATIAL SCALES MAY FURTHER REDUCE
LOSSES BY PREVENTING INVASION OF SEED PREDATORS &IG
4HE SPATIAL SCALE AT WHICH SYNCHRONY IN SEED PRODUCTION REDUCES SEED LOSSES TO
SEED PREDATORS DEPENDS ON THE MOBILITY OF THE PREDATORS +ELLY AND 3ORK
)NSECT PREDATORS CONSIDERED IN THIS CHAPTER MAY DISPERSE OVER RELATIVELY SHORT
DISTANCES "IRDS OR MAMMALS IN CONTRAST MAY BE HIGHLY MOBILE #URRAN AND
,EIGHTON )N ORDER TO SATIATE SUCH PREDATORS SO AS TO REDUCE SEED LOSSES
LARGER SPATIAL SCALES OF REPRODUCTIVE SYNCHRONY MUST BE MAINTAINED 4HE ANALYSIS
OF APPROXIMATE INVASION CRITERIA GIVEN IN EQUATIONS AND APPLIES ALSO TO HIGHLY
MOBILE SPECIALIST CONSUMERS BECAUSE THE FORMULATIONS CAN BE WRITTEN FOR ARBITRARY
NEIGHBORHOOD SIZES NOT RESTRICTED TO NEAREST NEIGHBOR DISPERSAL
) NVASIBILITY OF SEED PREDATORS
4HE COMMUNITY CONSEQUENCES OF MASTING ARE ONLY KNOWN FOR SELECTED CASE
STUDIES -ICE AND SQUIRRELS FOR INSTANCE EXPERIENCE ENHANCED SURVIVAL AND RAPID
POPULATION GROWTH IN YEARS OF OAK MAST 3UCH POPULATION MAY CONVERSELY CRASH
TO LOW LEVELS IN YEARS OF LITTLE ACORN BECAUSE OF A SHORTAGE OF FOOD RESOURCES 7OLFF
-C3HEA -ASTING HAS ALSO BEEN SHOWN TO HAVE CASCADING EFFECTS
THROUGH FOOD WEBS AND ECOLOGICAL COMMUNITIES IN SYSTEMS RANGING FROM OUT
BREAKS OF GYPSY MOTH IN OAK FORESTS TO HUMAN RISK OF ,YME EXPOSURE /STFELD ET AL
/STFELD AND +EESING 3UCH STUDIES ON THE TROPHIC CASCADES FOLLOWING
VARIABLE MASTING MAY PROVIDE KEY INSIGHTS INTO INVASION AS A COMMUNITY PROCESS
3HEA AND #HESSON
)N THIS CHAPTER WE FOCUS ON THE DYNAMICS OF A SINGLE SEED PREDATOR SPECIES ON
A SINGLE MASTING RESOURCE )N THE FUTURE IT WILL BE OF INTEREST TO CONSIDER INVASIBIL
ITY TO ADDITIONAL SEED PREDATORS AND POSSIBLY EXOTIC SPECIES 7HEN EXOTIC SEED
PREDATORS ARE INTRODUCED INTO A PLANT POPULATION THAT SHOW MASTING A POTENTIAL
STRATEGY THE PREDATOR MAY EMPLOY TO COPE WITH VARIABLE RESOURCE AVAILABILITY MAY
BE AN EXTENDED DIAPAUSE +ELLY ET AL AND -C+ONE ET AL DEMON
STRATED THAT PLANT POPULATIONS SUFFERING SEED PREDATION BY THE PREDATORS WHO HAVE
EXTENDED DIAPAUSE MAY NEED TO SHOW EXTREMELY HIGH LEVELS OF MAST SEEDING 4HIS
DEMONSTRATION HAS BEEN RECENTLY SUPPORTED BY A THEORETICAL STUDY OF 3ATAKE AND
"JRNSTAD 4HEREFORE HOW PRE
EXISTING MASTING MIGHT MITIGATE THE IMPACT
OF INVADING PREDATORS IS DEPENDENT ON THE LEVEL BOTH OF PRE
EXISTING MASTING OF
PLANT POPULATION AND THE EXTENDED DIAPAUSE OF INVADER 7HEN THE RESIDENT SEED
PREDATOR PERSISTS ON A MASTING RESOURCE THE COMPETITIVE INTENSITY BETWEEN THE
RESIDENT AND INVADING SPECIES WILL mUCTUATE DUE TO SPATIOTEMPORAL VARIABILITY
IN RESOURCE AVAILABILITY )NVASION SUCCESS OF AN EXOTIC SPECIES MAY BE ENHANCED
WHEN THE RESIDENT POPULATION IS SATIATED THROUGH LARGE SEED CROPS BECAUSE UNCON
SUMED RESOURCES MAY RESULT IN A COMPETITIVE RELEASE 4HIS INCREASE IN INVASIBILITY
MAY OCCUR THROUGH TEMPORAL COMPETITIVE RELEASE OR SPATIALLY THROUGH THE FORMA
TION OF COMPETITION
FREE SPATIAL CLUSTERS )NVASIBILITY OF EXOTIC SEED PREDATORS IS
DETERMINED BY THE INTERWINED INTERACTION BETWEEN TEMPORAL PROCESS AND SPATIAL
PATTERN ILLUSTRATING THE COMPLEX COMMUNITY CONSEQUENCES OF MASTING
!#+./7,%$'%-%.4
4HIS WORK WAS SUPPORTED IN PART BY A FELLOWSHIP AND A GRANT
IN AID FROM THE *APAN
3OCIETY FOR THE 0ROMOTION OF 3CIENCE !3 AND 53$!.2) GRANT NO
/." 4HE AUTHORS THANK THE FOLLOWING PEOPLE 5 $IECKMANN 2 ! )MS
$ +ELLY ! ,IEBHOLD * ! * -ETZ AND 2 3 /STFELD FOR THEIR HELPFUL COMMENTS
! 3ATAKE /. "JRNSTAD AND 9 )WASA
2%&%2%.#%3
!HLBERG / 2NNBRSMALEN !RGYRESTHIA CONJUGELLA :ELL %N REDOGRELSE FR UNDER
SKNINGAR REN IN 3WEDISH WITH %NGLISH SUMMARY ,ANTBRUKSENTO
MOLOGISKA AVDELNINGEN 3TOCKHOLM
#OMINS ( . - 0 (ASSELL AND 2 - -AY 4HE SPATIAL DYNAMICS OF HOST PARASITOID
SYSTEMS *OURNAL OF !NIMAL %COLOGY
#OHEN $ AND 3 ! ,EVIN $ISPERSAL IN PATCHY ENVIRONMENTS THE EFFECTS OF TEMPORAL
AND SPATIAL STRUCTURE 4HEORETICAL 0OPULATION "IOLOGY
#URRAN , - AND - ,EIGHTON 6ERTEBRATE RESPONSES TO SPATIOTEMPORAL VARIATION IN
SEED PRODUCTION OF MAST FRUITING $IPTEROCARPACEAE %COLOGICAL -ONOGRAPH
$AVIS - ! * 0 'RIME AND + 4HOMPSON &LUCTUATING RESOURCES IN PLANT COMMUNI
TIES A GENERAL THEORY IN INVASIBILITY *OURNAL OF %COLOGY
$AVIS - ! AND - 0ELSOR %XPERIMENTAL SUPPORT FOR A RESOURCE BASED MECHANISTIC
MODEL OF INVASIBILITY %COLOGY ,ETTERS
%LLNER 3 AND ! 3HMIDA 7HY ARE ADAPTATIONS FOR LONG RANGE SEED DISPERSAL RARE IN
DESERT PLANTS /ECOLOGIA
(ASSELL - 0 ( . #OMINS AND 2 - -AY 3PATIAL STRUCTURES AND CHAOS IN INSECT
POPULATION DYNAMICS .ATURE
(ERRERA # - 0 *ORDANO * 'UITIAN AND ! 4RAVESET !NNUAL VARIABILITY IN SEED
PRODUCTION BY WOODY PLANTS AND THE MASTING CONCEPT REASSESMENT OF PRINCIPLES AND
RELATIONSHIP TO POLLINATION AND SEED DISPERSAL !MERICAN .ATURALIST
(OLT 2 $ AND - ! -C0EEK #HAOTIC POPULATION DYNAMICS FAVORS THE EVOLUTION OF DISPER
SAL !MERICAN .ATURALIST
(OPPER + 2 2ISK SPREADING AND BET HEDGING INSECT POPULATION BIOLOGY !NNUAL
2EVIEW OF %NTOMOLOGY
)NOUE 4 AND ! ! (AMID 'ENERAL mOWERING OF TROPICAL RAINFORESTS IN 3ARAWAK
#ENTER FOR %COLOGICAL 2ESEARCH +YOTO 5NIVERSITY *APAN +YOTO
)SAGI 9 + 3UGIMURA ! 3UMIDA AND ( )TO (OW DOES MASTING HAPPEN AND SYN
CHRONIZE *OURNAL OF 4HEORETICAL "IOLOGY
*ANZEN $ ( 3EED PREDATION BY ANIMALS !NNUAL 2EVIEW OF %COLOGY AND 3YSTEMATICS
+ELLY $ 4HE EVOLUTIONARY ECOLOGY OF MAST SEEDING 4RENDS IN %COLOGY %VOLUTION
+ELLY $ AND 6 , 3ORK -AST SEEDING IN PERENNIAL PLANTS WHY HOW WHERE !NNUAL
2EVIEW OF %COLOGY AND 3YSTEMATICS
+ELLY $ AND * * 3ULLIVAN 1UANTIFYING THE BENElT OF MAST SEEDING ON PREDATOR SATIA
TION AND WIND POLLINATION IN #HIONOCHLOA PALLENS 0OACEAE /IKOS
+ELLY $ ! , (ARRISON 7 ' ,EE ) * 0AYTON 0 0 7ILSON AND 7 - 3CHAUBER
0REDATOR SATIATION AND EXTREME MAST SEEDING IN SPECIES OF #HIONOCHLOA 0OACEAE
/IKOS
+OBRO 3 , 3REIDE % $JNNE 4 2AFOSS ' *AASTAD AND 0 7ITZGALL -ASTING OF
ROWAN 3ORBUS AUCUPARIA , AND CONSEQUENCES FOR THE APPLE FRUIT MOTH !RGYRESTHIA CON
JUGELLA :ELLER 0OPULATION %COLOGY
) NVASIBILITY OF SEED PREDATORS
+OENIG 7 $ AND * - ( +NOPS 3CALE OF MAST SEEDING AND TREE RING GROWTH .ATURE
,EVIN 3 ! $ #OHEN AND ! (ASTINGS $ISPERSAL STRATEGIES IN PATCHY ENVIRONMENTS
4HEORETICAL 0OPULATION "IOLOGY
-C+ONE - * $ +ELLY ! , (ARRISON * * 3ULLIVAN AND ! * #ONE "IOLOGY OF INSECTS
THAT FEED IN THE INmORESCENCES OF #HINOCHLOA 0OACEAE IN .EW :EALAND AND THEIR REL
EVANCE TO MAST SEEDING .EW :EALAND *OURNAL OF :OOLOGY
-C3HEA 7 * 4HE INmUENCE OF ACORN CROPS ON ANNUAL VARIATION IN RODENT AND BIRD
POPULATIONS WITHIN OAK DOMINATED FORESTS %COLOGY
-OMOSE + 4 .AGAMITSU AND 4 )NOUE 4HE REPRODUCTIVE ECOLOGY OF AN EMERGENT
DEPTEROCARP IN A LOWLAND RAIN FOREST IN 3ARAWAK 0LANT 3PECIES "IOLOGY
.ILSSON 3 ' AND 5 7TLJUNG 3EED PREDATION AND CROSS POLLINATION IN MAST SEEDING
BEECH &AGUS SYLVATICA PATCHES %COLOGY
.ORTON $ ! AND $ +ELLY -AST SEEDING OVER YEARS BY $ACRYDIUM CUPRESSINUM
,AMB RIMU 0ODOCARPACEAE IN .EW :EALAND THE IMPORTANCE OF ECONOMIES OF SCALE
&UNCTIONAL %COLOGY
/STFELD 2 3 *ONES # ' AND 7OLFF * / /F MICE AND MAST ECOLOGICAL CONNECTIONS
IN EASTERN DECIDUOUS FORESTS "IO3CIENCE
/STFELD 2 3 AND & +EESING 0ULSED RESOURCES AND COMMUNITY DYNAMICS OF CONSUM
ERS IN TERRESTRIAL ECOSYSTEMS 4RENDS IN %COLOGY %VOLUTION
2EES - $ +ELLY AND / . "JRNSTAD 3NOW TUSSOCKS CHAOS AND THE EVOLUTION OF
MAST SEEDING !MERICAN .ATURALIST
3AKAI 3 + -OMOSE 4 9UMOTO 4 .AGAMITSU ( .AGAMASU ! ! (AMID 4 .AKASHIZUKA
0LANT REPRODUCTIVE PHENOLOGY OVER FOUR YEARS INCLUDING AN EPISODE OF GEN
ERAL mOWERING IN A LOWLAND DIPTEROCARP FOREST 3ARAWAK -ALAYSIA !MERICAN *OURNAL OF
"OTANY
3ATAKE ! AND 9 )WASA 0OLLEN COUPLING OF FOREST TREES FORMING SYNCHRONIZED AND
PERIODIC REPRODUCTION OUT OF CHAOS *OURNAL OF 4HEORETICAL "IOLOGY
3ATAKE ! AND 9 )WASA A 3PATIALLY LIMITED POLLEN EXCHANGE AND A LONG RANGE SYN
CHRONIZATION OF TREES %COLOGY
3ATAKE ! AND 9 )WASA B 4HE SYNCHRONIZED AND INTERMITTENT REPRODUCTION OF FOREST
TREES IS MEDIATED BY THE -ORAN EFFECT ONLY IN ASSOCIATION WITH POLLEN COUPLING *OURNAL
OF %COLOGY
3ATAKE ! / . "JRNSTAD AND 3 +OBRO -ASTING AND TROPHIC CASCADES INTERPLAY
BETWEEN ROWAN TREES APPLE FRUIT MOTH AND THEIR PARASITOID IN SOUTHERN .ORWAY /IKOS
3ATAKE ! AND / . "JRNSTAD 3PATIAL $YNAMICS OF 3PECIALIST 3EED 0REDATORS ON
3YNCHRONIZED AND )NTERMITTENT 3EED 0RODUCTION OF (OST !MERICAN .ATURALIST
3ELS 6 / (OGSTAD ' !NDERSSON AND 4 VON 0ROSCHWITZ 0OPULATION CYCLES OF
AUTUMNAL MOTH %PIRRITA AUTUMNATA IN RELATION TO BIRCH MAST SEEDING /ECOLOGIA
3CHAUBER % - $ +ELLY 0 4URCHIN # 3IMON 7 ' ,EE 2 " !LLEN ) * 0AYTON 0 2
7ILSON 0 % #OWAN AND 2 % "ROCKIE -ASTING BY EIGHTEEN .EW :EALAND PLANT
SPECIES THE ROLE OF TEMPERATURE AS A SYNCHRONIZING CUE %COLOGY
! 3ATAKE /. "JRNSTAD AND 9 )WASA
3HIBATA - ( 4ANAKA AND 4 .AKASHIZUKA 3YNCHRONIZED ANNUAL SEED PRODUCTION
BY PRINCIPAL TREE SPECIES IN A TEMPERATE DECIDUOUS FOREST *APAN %COLOGY
3CHNURR * , 2 3 /STFELD AND $ #ANHAM $IRECT AND INDIRECT EFFECT OF MASTING ON
RODENT POPULATIONS AND TREE SEED SURVIVAL /IKOS
3HEA + AND 0 #HESSON #OMMUNITY ECOLOGY THEORY AS A FRAMEWORK FOR BIOLOGICAL
INVASIONS 4RENDS IN %COLOGY AND %VOLUTION
3HIBATA - ( 4ANAKA AND 4 .AKASHIZUKA #AUSE AND CONSEQUENCES OF MAST SEED
PRODUCTION OF FOUR CO
OCCURRING #ARPINUS SPECIES IN *APAN %COLOGY
3ILVERTOWN * 7 4HE EVOLUTIONALY ECOLOGY OF MAST SEEDING IN TREES "IOLOGICAL
*OURNAL OF THE ,INNEAN 3OCIETY
3MITH # # * , (AMRICK AND # , +RAMER 4HE ADVANTAGE OF MAST YEARS FOR WIND
POLLINATION !MERICAN .ATURALIST
3ORK 6 , * "RAMBLE AND / 3EXTON %COLOGY OF MAST
FRUITING IN THREE SPECIES OF
NORTH !MERICAN DECIDUOUS OAKS %COLOGY
3PERENS 5 A &RUIT PRODUCTION IN 3ORBUS AUCUPARIA , 2OSACEAE AND PRE
DISPERSAL
SEED PREDATION BY THE APPLE FRUIT MOTH !RGYRESTHIA CONJUGELLA :ELL /ECOLOGIA
3PERENS 5 B ,ONG
TERM VARIATION IN AND EFFECTS OF FERTILIZED ON mOWER FRUIT AND SEED
PRODUCTION IN THE TREE 3ORBUS AUCUPARIA 2OSACEAE %COGRAPHY
4UTIN 4 ' 6 ( (EYWOOD . ! "URGES $ - -OORE $ ( 6ALENTINE 3 - 7ALTERS AND
$ ! 7EBB &LORA EUROPAEA ND EDN #AMBRIDGE 5NIVERSITY 0RESS #AMBRIDGE
7OLFF * / 0OPULATION mUCTUATIONS OF MAST
EATING RODENTS ARE CORRELATED WITH PRO
DUCTION OF ACORNS *OURNAL OF -AMMALOGY
#HAPTER THIRTEEN
)NVASIONS AND THE REGULATION
OF PLANT POPULATIONS
BY PATHOGENS
' 3 'ILBERT AND ) - 0ARKER
).42/$5#4)/.
4HE POTENTIAL OF PATHOGENS TO HAVE DRAMATIC IMPACTS ON PLANT POPULATIONS IS MADE
CLEAR BY FAMILIAR STORIES OF THE )RISH POTATO FAMINE &RY AND 'OODWIN THE
ECOLOGICAL EXTINCTION OF CHESTNUTS CAUSED BY CHESTNUT BLIGHT !NAGNOSTAKIS
AND THE TRANSFORMATION OF !USTRALIAN *ARRAH FORESTS TO SCRUBLAND BY 0HYTOPHTHORA
CINNAMOMI 7ESTE AND -ARKS 3IMILARLY THE ANNUAL WORLDWIDE EXPENDI
TURE OF OVER BILLION IN FUNGICIDE APPLICATION $ONALDSON ET AL REmECTS
THE TOLL FUNGAL PATHOGENS ALONE CAN TAKE ON PLANT GROWTH AND FECUNDITY IN
AGRICULTURAL SYSTEMS 3OME OF THE MOST NOTABLE EXAMPLES OF THESE IMPACTS ARISE
WHEN PATHOGENS ARE INTRODUCED INTO NOVEL BIOLOGICAL ENVIRONMENTS INVASIVE AND
EMERGENT PATHOGENS CONTINUE TO FRUSTRATE THE BEST EFFORTS OF RESOURCE MANAGERS
CONSERVATION BIOLOGISTS AND PLANT PROTECTION AGENCIES 7ESTE AND -ARKS
$AUGHTREY ET AL 'OODELL ET AL 'ORDON ET AL -C$ONALD AND (OFF
7INGlELD ET AL 'ILBERT 2IZZO AND 'ARBELOTTO 0ARKER AND
'ILBERT #ONCURRENT WITH EFFORTS TO REDUCE THE IMPACTS OF UNWANTED DISEAS
ES THERE IS BROAD INTEREST AMONG RESEARCHERS AGRICULTURALISTS AND LAND MANAGERS
TO HARNESS THE DESTRUCTIVE POTENTIAL OF PLANT PATHOGENS TO CONTROL WEEDY PLANTS
(ASAN AND !YRES #HARUDATTAN AND $INOOR
-7 #ADOTTE ET AL EDS #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY n
3PRINGER 0RINTED IN THE .ETHERLANDS
'3 'ILBERT AND )- 0ARKER
)N NATURAL ECOSYSTEMS PATHOGENS HAVE GREAT POTENTIAL TO INmUENCE THE DYNAM
ICS AND COMPOSITION OF PLANT POPULATIONS AND COMMUNITIES THROUGH DENSITY
DEPENDENT AND COEVOLUTIONARY DYNAMICS SEE REVIEWS IN $INOOR AND %SHED
"URDON *AROSZ AND $AVELOS !LEXANDER ET AL !LEXANDER AND
(OLT 'ILBERT )N MANY PLANT COMMUNITIES PLANT PATHOGENS
MAY PREVENT COMPETITIVE EXCLUSION AND THUS HELP MAINTAIN SPECIES DIVERSITY
'ILLETT 0ACKER AND #LAY 'ILBERT 7RIGHT 'ILBERT
#OLLECTIVE INSIGHTS FROM RESEARCH ON EPIDEMIC DISEASES BIOLOGICAL CONTROL AND
THE EVOLUTIONARY ECOLOGY OF DISEASES IN NATURAL ECOSYSTEMS PROVIDE A ROBUST BASIS
FOR IDENTIFYING WHEN PATHOGENS ARE LIKELY TO BE IMPORTANT IN REGULATING PLANT
POPULATIONS AND THEIR IMPLICATIONS FOR UNDERSTANDING BIOLOGICAL INVASIONS
(ERE WE DRAW BROADLY FROM A DIVERSE LITERATURE TO PLACE THE REGULATION OF PLANT
POPULATIONS BY PATHOGENS INTO THE CONTEXT OF TWO KEY COMPLEMENTARY THEORIES
ABOUT THE ROLE OF PATHOGENS IN BIOLOGICAL INVASIONS "IOTIC 2ESISTANCE -ARON AND
6ILA AND %SCAPE FROM .ATURAL %NEMIES +EANE AND #RAWLEY 4ABLE
7E THEN CONSIDER THE PRACTICAL IMPLICATIONS FOR USING INTRODUCED PATHOGENS FOR
CLASSICAL BIOLOGICAL CONTROL OF INTRODUCED INVASIVE PLANTS 4ABLE ILLUSTRATED WITH
A DETAILED CASE STUDY OF THE CONTROL OF #HONDRILLA BY RUST FUNGI
0,!.4 $)3%!3%3 !.$ 0/05,!4)/. 2%'5,!4)/.
0ATHOGENS REDUCE THE lTNESS OF INDIVIDUAL PLANTS BY KILLING THEM REDUCING GROWTH
IMPEDING COMPETITIVE ABILITY OR BY ROTTING FRUITS OR SEEDS SEE RECENT REVIEW IN
'ILBERT 4HE GREATER PROBABILITY OF PATHOGEN SPREAD BETWEEN CLOSELY SPACED
HOST PLANTS MEANS THAT MOST FUNGAL PLANT DISEASES SHOW DENSITY
DEPENDENT DEVEL
OPMENT )N ADDITION DENSELY SPACED PLANTS MAY CREATE MICROCLIMATES THAT ENCOUR
AGE PATHOGEN GROWTH AND HOSTS STRESSED BY COMPETITION MAY BE MORE SUSCEPTIBLE
TO DISEASE "URDON AND #HILVERS 'ILBERT 4HE COMBINATION OF STRONG
IMPACTS ON INDIVIDUAL HOST PLANTS AND DENSITY
DEPENDENT DISEASE DEVELOPMENT
SUGGESTS THAT PATHOGENS SHOULD BE POWERFUL REGULATORS OF PLANT POPULATIONS
.EVERTHELESS ASIDE FROM EPIDEMICS CAUSED BY INTRODUCED PATHOGENS THERE ARE
REMARKABLY FEW EMPIRICAL STUDIES SHOWING THAT PLANT DISEASES ARE RESPONSIBLE
FOR REGULATING PLANT POPULATION DYNAMICS IN NATURAL ECOSYSTEMS 'ILBERT
)N LARGE PART THIS ABSENCE REmECTS THE DIFlCULTIES OF ISOLATING DISEASE IMPACTS FROM
OTHER FACTORS AND THE STRONG FOCUS ON DISEASES WITH ECONOMIC IMPORTANCE "UT
PHYSIOLOGICAL OR EVOLUTIONARY RESPONSES BY THE HOST MAY ALSO COUNTERACT THE
REGULATORY ACTIONS OF PATHOGENS IN NATURAL ECOSYSTEMS )N PARTICULAR PLANTS THAT
SURVIVE AFTER DISEASE HAS KILLED OR STUNTED COMPETING CONSPECIlC NEIGHBORS MAY
SHOW A COMPENSATORY RESPONSE THAT OFFSETS NUMERICAL LOSSES FROM DISEASE AT THE
POPULATION LEVEL &RIESS AND -AILLET !LEXANDER AND (OLT !LEXANDER
AND -IHAIL )N ADDITION MATERNALLY
TRANSMITTED INDUCED RESISTANCE CAN
GENERATE CROSS
GENERATIONAL EFFECTS THAT AMELIORATE THE NUMERICAL IMPACTS OF
DISEASE IN SUCCEEDING GENERATIONS !GRAWAL ET AL &INALLY THE GENETICS
4ABLE )MPLICATIONS OF DIFFERENT FEATURES OF PLANT
PATHOGEN INTERACTIONS FOR THREE AREAS OF INVASION BIOLOGY THE TWO THEORIES OF BIOTIC
RESISTANCE AND ESCAPE FROM NATURAL ENEMIES AND THE lELD OF CLASSICAL BIOLOGICAL CONTROL
&EATURES OF PLANT
PATHOGEN "IOTIC RESISTANCE %SCAPE FROM NATURAL ENEMIES "IOLOGICAL CONTROL
INTERACTIONS
)MPACT OF PATHOGEN ON HOST 0RE
ADAPTED PATHOGENS IN NEW 0ATHOGENS IN NATIVE RANGE LIMIT 4O BE EFFECTIVE CONTROL AGENTS
RANGE ARE HIGHLY VIRULENT ON HOST DENSITY OR DISTRIBUTION PATHOGENS MUST BE IMPORTANT IN
NAVE HOSTS )NTRODUCED PLANTS LEAVE POPULATION REGULATION IN NATIVE
VIRULENT PATHOGENS BEHIND RANGE
)NTERACTIONS WITH ,OCAL PATHOGENS ARE ADAPTED 0LANTS MAY ESCAPE NATURAL )NTRODUCED BIOCONTROL PATHOGENS
ENVIRONMENT TO LOCAL ENVIRONMENT ENEMIES IF INTRODUCED INTO AN MAY FAIL IF POORLY ADAPTED TO
DISEASE TRIANGLE ENVIRONMENT NOT CONDUCIVE TO LOCAL ENVIRONMENT
DISEASE DEVELOPMENT
(OST RANGE OF PATHOGEN 'ENERALIST PATHOGENS ARE MORE !SSUMES SPECIALIST PATHOGENS PLAY /NLY SPECIALIST PATHOGENS CAN BE
LIKELY TO ACQUIRE NEWLY INTRODUCED A UNIQUE ROLE IN REGULATING PLANT UTILIZED AS CONTROL AGENTS
HOST SPECIES POPULATIONS
.ATURAL HISTORY $ENSITY DEPENDENT DISEASE 0ATHOGENS WITH RESTING STRUCTURES 0ATHOGENS WITH RESTING STRUCTURES
0LANT POPULATIONS BY PATHOGENS
LIFE HISTORY OF PATHOGEN DEVELOPMENT MAY CHANGE IMPACT OR ALTERNATE LIFE HISTORY STRATEGIES SHOULD SHOW GREATER SUCCESS
ON INVADING HOSTS AS INVASION MOST LIKELY TO ACCOMPANY
PROCEEDS INTRODUCED PLANTS SUCH PATHOGENS
MAY HAVE GREATER VIRULENCE
2APID 0ATHOGENS MAY ACQUIRE INTRODUCED (OST MAY LOSE DEFENSES TO ORIGINAL (OST MAY DEVELOP RESISTANCE TO
EVOLUTIONARY CHANGES HOST THROUGH EVOLUTIONARY HOST PATHOGENS AFTER INTRODUCTION INTRODUCED BIOCONTROL AGENT
SHIFT 6IRULENCE MAY INCREASE
QUICKLY WHEN HOST BECOMES
COMMON
'3 'ILBERT AND )- 0ARKER
OF PLANT
PATHOGEN INTERACTIONS CAN BE HIGHLY DYNAMIC WITH LARGE CHANGES IN
PATHOGEN VIRULENCE OR HOST RESISTANCE EVOLVING IN A FEW GENERATIONS EG "URDON
AND 4HOMPSON "ISHOP ET AL 3UCH RAPID EVOLUTIONARY CHANGES ARE
EXPECTED TO STRONGLY INmUENCE THE IMPACT OF PATHOGENS ON HOST NUMERICAL DYNAM
ICS !LEXANDER ET AL
")/4)# 2%3)34!.#% !.$ %3#!0% &2/- .!452!, %.%-)%3
&OR DISEASE TO DEVELOP VIRULENT PATHOGENS SUSCEPTIBLE HOST PLANTS AND SUITABLE
ENVIRONMENTAL CONDITIONS MUST CONVERGE COMMONLY CALLED THE $ISEASE 4RIANGLE
#HANGES IN ANY OF THESE COMPONENTS CAN REDUCE OR INCREASE HOW MUCH DISEASES
AFFECT PLANT POPULATION DYNAMICS )F PLANTS ARE INTRODUCED TO AN ENVIRONMENT
WHERE VIRULENT PATHOGENS ARE NOT PRESENT OR WHERE ENVIRONMENTAL CONDITIONS
DO NOT FAVOR DISEASE DEVELOPMENT THE PLANT POPULATION MAY BE RELEASED FROM
PREVIOUS REGULATION BY PATHOGENS 3IMILARLY PLANTS INTRODUCED TO A NEW LOCALE
WITH NOVEL VIRULENT PATHOGENS OR WHERE ENVIRONMENTAL CONDITIONS FAVOR DISEASE
DEVELOPMENT MAY EXPERIENCE UNPRECEDENTED POPULATION REGULATION BY PATHOGENS
3UCH CHANGES IN THE PREVALENCE AND SEVERITY OF DISEASES UNDER DIFFERENT CONDITIONS
HAVE LONG BEEN THOUGHT TO PLAY AN IMPORTANT ROLE IN THE PROCESS OF BIOLOGICAL
INVASIONS IN TWO HYPOTHETICAL WAYS
)N NATURAL HABITATS NATIVE PATHOGENS COLONIZE AND ARE HIGHLY VIRULENT ON NAVE
INTRODUCED PLANT SPECIES AND PREVENT POPULATION GROWTH "IOTIC 2ESISTANCE
0ATHOGENS WERE IMPORTANT IN REGULATING THE PLANT POPULATION IN ITS NATIVE
RANGE BUT ARE NOT IN THE INTRODUCED RANGE %SCAPE FROM .ATURAL %NEMIES
"IOTIC RESISTANCE
4HE GREAT MAJORITY OF PLANT INTRODUCTIONS DO NOT RESULT IN INVASIONS -ACK
7ILLIAMSON 2ATHER MANY INTRODUCED SPECIES EITHER FAIL TO THRIVE ALTOGETH
ER OR ARE RESTRICTED TO HUMAN CULTIVATION UNABLE TO BUILD SELF
SUSTAINING POPULA
TIONS IN WILD PLANT COMMUNITIES OR EVEN IN DISTURBED RANGELAND COMMUNITIES
2ESEARCH ATTEMPTING TO IDENTIFY PLANT TRAITS THAT CONFER INVASIVENESS 2EICHARD
AND (AMILTON 2EJMNEK AND 2ICHARDSON SHOWS THAT OUR PREDICTIVE
ABILITY IS IMPERFECT AND IT SEEMS TO BE MORE DIFlCULT TO PREDICT WHICH INTRODUC
TIONS WILL FAIL THAN WHICH WILL SUCCEED 2EICHARD AND (AMILTON "IOTIC
RESISTANCE IS ONE POSSIBLE EXPLANATION FOR WHY SOME INTRODUCTIONS FAIL WHEN THEY
hSHOULDv SUCCEED NATIVE PESTS AND PATHOGENS COLONIZE EXOTIC PLANTS AND ELIMINATE
THEM BEFORE THEY CAN ESTABLISH A VIABLE POPULATION %LTON 3IMBERLOFF
-ACK
0ATHOGENS WILL CONTRIBUTE TO BIOTIC RESISTANCE ONLY WHEN THREE CONDITIONS ARE
MET &IRST THE PATHOGENS INVOLVED MUST NOT BE NARROW HOST SPECIALISTS )N PLANT
COMMUNITIES WITH HIGH HOST DIVERSITY AND CORRESPONDING LOW HOST DENSITY
GENERALIST PLANT PATHOGENS SHOULD DOMINATE 'ILBERT ET AL 'ILBERT
0LANT POPULATIONS BY PATHOGENS
WHICH COULD CONTRIBUTE TO THE HIGH INVASION RESISTANCE OF HYPER
DIVERSE ECO
SYSTEMS LIKE TROPICAL RAIN FORESTS &INE 3ECOND BECAUSE DENSITIES OF THE
INTRODUCED HOST WILL USUALLY BE LOW IMMEDIATELY FOLLOWING INTRODUCTION PATHO
GENS INVOLVED IN BIOTIC RESISTANCE WOULD NOT BE THOSE FOR WHICH TRANSMISSION OR
HOST SWITCHING WAS STRONGLY DENSITY
DEPENDENT 4HIRD THE PATHOGEN MUST EXACT
A HIGH lTNESS COST ON THE HOST #OMBINATIONS OF GENERALIZATION AND HIGH VIRU
LENCE ARE NOT UNUSUAL IN PATHOGENS ESPECIALLY FOR SPECIES THAT HAVE LONG RESTING
STAGES OR SAPROPHYTIC ABILITY EG 0HYTOPHTHORA CINNAMOMI 6ERTICILLIUM DAHLIAE
&USARIUM OXYSPORUM %VEN PATHOGENS THAT CAN ATTACK MANY HOSTS HAVE DIF
FERENTIAL IMPACTS ON DIFFERENT HOST SPECIES )F A PATHOGEN HAS A GREATER NEGATIVE
EFFECT ON THE COMPETITIVE ABILITY OF THE NON
NATIVE HOSTS THAN ON THE NATIVE HOSTS
THEN IT WILL CONTRIBUTE TO THE COMPETITIVE EXCLUSION OF THE NON
NATIVE +EANE AND
#RAWLEY
3INCE EVEN FAIRLY SPECIALIZED PATHOGENS OFTEN INFECT MANY MEMBERS OF THE SAME
GENUS OR FAMILY IT IS LIKELY THAT THE NUMBER OF PATHOGENS COMPETENT TO INFECT A
NOVEL HOST DEPENDS ON WHETHER IT IS PHYLOGENETICALLY RELATED TO NATIVE SPECIES
ALREADY IN THE COMMUNITY 4HEREFORE BIOTIC RESISTANCE SHOULD FAVOR INVASION BY
SPECIES WITH NO CLOSE RELATIVES $ESPITE THIS CLEAR PREDICTION AND THE ACCEPTED
PRACTICE OF USING PHYLOGENETIC RELATIONSHIPS IN QUARANTINE AND TRADE POLICY TO
TARGET RELATIVES OF WEEDY PLANTS OR IDENTIFY POTENTIAL CARRIER HOSTS SURPRISINGLY
FEW STUDIES OF INVASION EVEN MENTION THE PHYLOGENETIC STRUCTURE OF THE INVADED
COMMUNITIES )N ONE RARE ATTEMPT -ACK FOUND THAT FOR OUT OF REGIONAL
mORAS NATURALIZED SPECIES WERE MORE COMMON IN GENERA WITH NO NATIVE CONGENERS
THAN THOSE WITH NATIVE CONGENERS CONSISTENT WITH THE IDEA OF BIOTIC RESISTANCE
(OWEVER THIS STUDY WAS NOT ABLE TO CONTROL FOR THE EFFECT OF OPPORTUNITY WHETHER
PLANTS WITHOUT NATIVE CONGENERS HAVE A HIGHER PROBABILITY OF SUCCESSFUL INVASION
GIVEN THEIR PROBABILITY OF INTRODUCTION 5SING A DIFFERENT APPROACH $UNCAN AND
7ILLIAMS COMPILED A LIST OF ALL PLANT SPECIES THAT HAVE EVER BEEN INTRODUCED
FOR CULTIVATION IN .EW :EALAND 4HEY FOUND THAT INTRODUCED SPECIES IN GENERA
THAT ALREADY HAD RESIDENT NATIVES WERE MORE LIKELY NOT LESS LIKELY TO SUCCESSFULLY
NATURALIZE 4HEY SUGGEST THAT SPECIES WITH NATIVE CONGENERS MAY SHARE CHARAC
TERISTICS THAT MAKE THEM MORE lT IN THE INTRODUCED RANGE AND THIS FACTOR OVER
WHELMS THE POTENTIAL EFFECT OF LOCAL NATURAL ENEMIES
4HERE IS A STRIKING NEED FOR MORE STUDIES TO DETERMINE WHETHER AND WHEN
BIOTIC RESISTANCE BY NATIVE PATHOGENS OCCURS "ECAUSE THERE IS LITTLE INFORMATION
ON WHERE AND WHEN UNINTENTIONAL SPECIES INTRODUCTIONS FAIL ASSIGNING MECHA
NISMS TO THESE FAILURES HAS BEEN NEARLY IMPOSSIBLE (ORTICULTURE FORESTRY AND
AGRICULTURE EACH PROVIDE EXAMPLES OF ENDEMIC PATHOGENS THAT HAVE DECIMATED
INTRODUCED PLANT SPECIES SO THAT GROWING THESE SPECIES IS NO LONGER ECONOMICALLY
VIABLE -ACK #OUTINHO ET AL 7INGlELD ET AL (OWEVER EXTRA
P OLATING FROM AGRICULTURAL OR SILVICULTURAL EXAMPLES T O I NVASIVE INTRODUCED SPECIES
REQUIRES CAUTION SINCE THE REGENERATION OF HOSTS IS CONTROLLED PREVENTING THE HOST
POPULATION FROM EVOLVING RESISTANCE OR TOLERANCE TO THE PATHOGEN !DDITIONALLY
SIMPLIlED SYSTEMS IN AGRICULTURE AND FORESTRY MAY SIMPLY LACK MODERATING EFFECTS
'3 'ILBERT AND )- 0ARKER
OF GREATER BIOTIC COMPLEXITY IN LESS MANAGED SYSTEMS EG HYPERPARASITES OF
THE PATHOGENS
%SCAPE FROM NATURAL ENEMIES
)NTRODUCED PLANTS THAT BECOME INVASIVE WEEDS ARE AMONG OUR MOST CHALLENGING
ENVIRONMENTAL PROBLEMS $!NTONIO AND 6ITOUSEK /FlCE OF 4ECHNOLOGY
!SSESSMENT 6ITOUSEK ET AL 0ARKER ET AL -ACK ET AL
AND ESCAPE FROM NATURAL ENEMIES PROVIDES A MECHANISM TO EXPLAIN THE INCREASED
GROWTH AND DENSITY OF INTRODUCED SPECIES $ARWIN #RAWLEY "LOSSEY
AND .TZOLD 4ILMAN -ARON AND 6ILA 3IEMANN AND 2OGERS
+EANE AND #RAWLEY 4HE h.ATURAL %NEMIES (YPOTHESISv POSITS
THAT INTRODUCED SPECIES LEAVE BEHIND THEIR NATURAL ENEMIES HERBIVORES SEED
PREDATORS AND PATHOGENS AND ARE THEREBY RELEASED FROM A KEY REGULATING
FACTOR LEADING TO A DRAMATIC INCREASE IN PLANT VIGOR POPULATION GROWTH ANDOR
COMPETITIVE ABILITY 4HIS IDEA FORMS THE JUSTIlCATION FOR CLASSICAL BIOLOGICAL CONTROL
IN WHICH NATURAL ENEMIES ARE BROUGHT FROM THE NATIVE RANGE TO CONTROL WEEDY
INVADERS (UFFAKER AND -ESSENGER $E"ACH AND 2OSEN 9ET DESPITE
THE IMPORTANCE OF THE .ATURAL %NEMIES (YPOTHESIS RIGOROUS EMPIRICAL TESTS ARE
FEW -ARON AND 6ILA
! NUMBER OF STUDIES HAVE TESTED WHETHER INTRODUCED PLANTS TEND TO GROW
FASTER OR LARGER AND WHETHER THERE IS EVIDENCE OF REDUCED ALLOCATION TO HERBIVORE
DEFENSES IN THE NEW RANGE #RAWLEY "LOSSEY AND .TZOLD 3IEMANN
AND 2OGERS BUT SEE 7ILLIS ET AL 4HEBAUD AND 3IMBERLOFF 6ILA
ET AL 7HILE SEVERAL OF THESE STUDIES HAVE SUGGESTED ECOLOGICAL RELEASE AND
EVOLUTION AWAY FROM DEFENSE TOWARD COMPETITIVE ABILITY THEY LACK DIRECT EVI
DENCE OF INVOLVEMENT BY NATURAL ENEMIES "LOSSEY AND .TZOLD 3IEMANN
AND 2OGERS /THER STUDIES HAVE TESTED THE .ATURAL %NEMIES (YPOTHESIS BY
COMPARING THE IMPACT OF NATURAL ENEMIES ON EXOTIC AND NATIVE SPECIES WITHIN THE
INTRODUCED RANGE )N A REVIEW OF STUDIES +EANE AND #RAWLEY FOUND
SOME CASES IN WHICH GENERALIST HERBIVORES SHOWED HIGHER IMPACTS ON NATIVE
SPECIES THAN NON
NATIVES AND OTHERS CHARACTERIZED BY THE REVERSE PATTERN
4HERE HAVE BEEN SURPRISINGLY FEW STUDIES WITH PATHOGENS IN ONE TEST 'OERGEN AND
$AEHLER FOUND THAT SMUT FUNGI CAUSED GREATER REPRODUCTIVE LOSS IN A NATIVE
GRASS (ETEROPOGON CONTORTUS THAN AN INTRODUCED GRASS 0ENNISETUM SETACEUM
%NEMY REMOVAL EXPERIMENTS ARE AN IMPORTANT BUT UNDERUTILIZED TOOL IN COM
PARING THE lTNESS EFFECT OF PATHOGENS OR HERBIVORES ON EXOTIC AND NATIVE SPECIES
WITHIN THE INTRODUCED RANGE +EANE AND #RAWLEY "LANEY AND +OTANEN
USED A FUNGICIDE EXPERIMENT TO REMOVE THE EFFECTS OF SOILBORNE FUNGI AND
OOMYCETES ON THE SURVIVAL OF SEEDS OF NATIVE VS INTRODUCED PLANTS IN TWO HABITATS
4HEY FOUND NO SUPPORT FOR A RELEASE FROM FUNGAL PATHOGENS IN THE SEED BANK OF
EXOTIC SPECIES COMPARED WITH NATIVE SPECIES -ORE RECENTLY 0ARKER AND 'ILBERT
UNPUBLISHED DATA FOUND NO DIFFERENCE IN FREQUENCY OF INFECTION LEAF DAMAGE
lTNESS EFFECTS OF FOLIAR AND DAMPING
OFF PATHOGENS OR PATHOGEN DIVERSITY BETWEEN
0LANT POPULATIONS BY PATHOGENS
SYMPATRIC SUITES OF NATIVE AND NON
NATIVE CLOVERS !LTHOUGH FUNGAL EXCLUSION
EXPERIMENTS IN THIS SYSTEM REVEALED SIGNIlCANT IMPACTS OF PATHOGENS THERE WAS
NO DIFFERENCE IN THE RESPONSE OF NATIVE AND NON
NATIVE SPECIES
&INALLY OTHER STUDIES HAVE TAKEN THE APPROACH OF COMPARING DISEASE ON A
SINGLE HOST SPECIES IN ITS NATIVE AND INVADED RANGES 7OLFE SURVEYED FOR
THE ANTHER SMUT FUNGUS -ICROBOTRYUM VIOLACEUM IN NATIVE POPULATIONS AND
INTRODUCED POPULATIONS OF 3ILENE LATIFOLIUM (E FOUND SIGNIlCANTLY MORE POPU
LATIONS INFECTED AND AT MUCH HIGHER INFECTION FREQUENCIES IN THE NATIVE RANGE
THAN IN THE INTRODUCED RANGE -ITCHELL AND 0OWER USED PUBLISHED RECORDS
OF PATHOGEN ASSOCIATIONS WITH PLANT HOSTS IN THEIR NATIVE AND INTRODUCED
RANGES 0LANTS WERE INFECTED BY FEWER PATHOGEN SPECIES IN THEIR NATURALIZED
RANGE 4HERE WAS ALSO AN INDICATION THAT SPECIES LEAVING BEHIND PROPORTIONALLY
MORE NATURAL ENEMIES WERE MORE LIKELY TO BE CATEGORIZED AS NOXIOUS OR INVASIVE
SPECIES BY LAND MANAGERS AND PUBLIC AGENCIES 5SING AN EXPERIMENTAL APPROACH
"ECKSTEAD AND 0ARKER DIRECTLY MEASURED THE DEMOGRAPHIC EFFECT OF PATHO
GENS ON AN INVADER IN THE CONTEXT OF KNOWN INFORMATION FROM THE SPECIES NATIVE
RANGE !MMOPHILA ARENARIA IN ITS NATIVE %UROPE IS LIMITED TO AN EARLY
SUCCESSIONAL
ROLE IN SHIFTING BEACH SANDS BY SOIL
BORNE PATHOGENS 6AN DER 0UTTEN ET AL
!S AN INVADER ON THE WEST COAST OF THE 53! !MMOPHILA REMAINS DOMINANT FOR
LONG PERIODS OF TIME (OWEVER ESCAPE FROM NATURAL ENEMIES DOES NOT EXPLAIN THIS
CONTRAST BETWEEN ITS NATIVE AND INVASIVE ECOLOGICAL ROLES "Y REPLICATING EXPERI
MENTS DONE IN THE NATIVE RANGE "ECKSTEAD AND 0ARKER FOUND THE NEGATIVE
EFFECT OF SOILBORNE PATHOGENS ON EARLY GROWTH IN THE INVADED RANGE WAS AT LEAST AS
LARGE OR LARGER THAN THEIR EFFECT IN THE NATIVE RANGE
4O PREDICT WHETHER AN INTRODUCED PLANT IS LIKELY TO BENElT FROM ESCAPING
NATURAL ENEMIES WE NEED TO UNDERSTAND THE RELATIVE IMPORTANCE OF HOST
SPECIALIST
VS HOST
GENERALIST PATHOGENS AND PESTS IN THE INVADED HABITAT -ARON AND 6ILA
+EANE AND #RAWLEY )F SPECIALIST PATHOGENS PREDOMINATE AND HOST
SHIFTS ARE RARE NATIVE PLANTS MAY BE SUPPRESSED MORE THAN COMPETING INVASIVE
SPECIES /N THE OTHER HAND IF GENERALIST PATHOGENS DOMINATE IN A SITE AND DO NOT
SHOW A PREFERENCE FOR NATIVE HOST SPECIES ONE WOULD NOT EXPECT AN INTRODUCED
PLANT TO EXPERIENCE RELEASE 4HEREFORE THE WIDE RANGE OF RESULTS SEEN IN THE ABOVE
EMPIRICAL STUDIES MAY BE IN PART EXPLAINED BY THE RELATIVE IMPORTANCE OF SPECIAL
IST AND GENERALIST NATURAL ENEMIES ! GREATER UNDERSTANDING OF THE PHYLOGENETIC
STRUCTURE OF PATHOGEN HOST RANGES COUPLED WITH ANALYSIS OF THE PHYLOGENETIC
STRUCTURE OF NATURAL PLANT COMMUNITIES 7EBB ET AL MAY HELP PREDICT THE
RELATIVE IMPORTANCE OF SPECIALIST VS GENERALIST PATHOGENS IN DIFFERENT KINDS OF
PLANT COMMUNITIES &OR INSTANCE IN A HIGH
DIVERSITY LOWLAND TROPICAL RAINFOREST
TREE SPECIES HOST GENERALISTS DOMINATED THE POLYPORE FUNGAL COMMUNITY
ALL OF THE MORE COMMON FUNGAL SPECIES WERE FOUND ON MULTIPLE FAMILIES OF HOST
TREES 'ILBERT ET AL &ERRER AND 'ILBERT )N CONTRAST IN A NEARBY LOW
DIVERSITY MANGROVE FOREST WITH ONLY THREE TREE SPECIES PRESENT EACH FROM A DIFFER
ENT FAMILY OF ALL POLYPORE FUNGAL COLLECTIONS BELONGED TO JUST THREE FUNGAL
SPECIES AND EACH SPECIES WAS HIGHLY SPECIALIZED ON JUST ONE MANGROVE SPECIES
'3 'ILBERT AND )- 0ARKER
'ILBERT AND 3OUSA 2ESEARCHERS ARE JUST NOW BEGINNING TO ADDRESS THE
RANGE OF HOST SPECIALIZATION IN DIFFERENT PLANT COMMUNITIES AND THE WIDE RANGE OF
OUTCOMES SUGGESTS THAT MANY MORE STUDIES WILL BE NEEDED BEFORE WE CAN FORMU
LATE PREDICTIVE GENERALIZATIONS
).4%.4)/.!,,9 ).42/$5#%$ 0!4(/'%.3 &/2 ")/,/')#!, #/.42/,
! DIRECT APPLICATION OF THE .ATURAL %NEMIES (YPOTHESIS IS DEPLOYING NATURAL
ENEMIES FROM THE NATIVE RANGE OF AN INVASIVE PLANT TO CONTROL THE INVADER POPU
LATION 3UCH CLASSICAL BIOLOGICAL CONTROL USES PLANTS AND PATHOGENS WITH SHARED
EVOLUTIONARY HISTORIES BUT A NEW ENVIRONMENTAL CONTEXT 4HERE ARE SEVERAL WAYS
IN WHICH BIOLOGICAL CONTROL INTERACTIONS MAY BE DIFFERENT FROM NATIVE PATHOGENS
ATTACKING INTRODUCED HOSTS &IRST UNLIKE NATIVE PATHOGENS WHICH ARE PRESUMABLY
ADAPTED TO THE LOCAL CLIMATE THE INTRODUCED BIOLOGICAL CONTROL AGENT EXPERIENCES
A NOVEL ENVIRONMENT WHICH COULD HAVE A LARGE IMPACT ON DISEASE DEVELOPMENT
#ASE STUDIES OF FAILED BIOLOGICAL CONTROL EFFORTS PROVIDE US WITH MANY EXAMPLES OF
THE IMPORTANCE OF THE DISEASE TRIANGLE -ORIN ET AL 3ECOND BOTH THE HOST
AND PATHOGEN ARE LIKELY TO BE GENETICALLY DEPAUPERATE (OWEVER THE PATHOGEN
WILL HAVE BEEN CHOSEN SPECIlCALLY TO BE VIRULENT ON THE INVASIVE HOST PLACING
THE HOST AT A RELATIVE DISADVANTAGE FOR EVOLUTIONARY RESPONSES 4HIRD ONLY FAIRLY
HOST
SPECIlC PATHOGENS ARE SELECTED FOR BIOLOGICAL CONTROL RELEASES MEANING THAT
PATHOGEN NUMERICAL DYNAMICS SHOULD ALWAYS BE CLOSELY LINKED TO INDIVIDUAL HOST
DENSITY )N FACT BIOLOGICAL CONTROL RELEASES ARE AN EXCELLENT OPPORTUNITY TO STUDY
FACTORS INmUENCING NUMERICAL DYNAMICS .OT ONLY SHOULD THERE BE A TIGHT CONNEC
TION BETWEEN PATHOGEN AND HOST DENSITY BUT INITIAL CONDITIONS OF THE INTERACTION
ARE WELL KNOWN 4HAT IS THE HOST POPULATION IS ORIGINALLY FREE OF THAT PATHOGEN
AND IS USUALLY AT HIGH DENSITY ! SUCCESSFUL EPIDEMIC PROVIDES AN OPPORTUNITY TO
QUANTIFY BOTH FREQUENCY
DEPENDENCE OF TRANSMISSION OF THE PATHOGEN AND DENSITY
DEPENDENCE AS THE HOST DENSITY DECLINES
0REDICTING THE SHORT
TERM AND LONG
TERM SUCCESS OF PARTICULAR BIOLOGICAL CON
TROL INTRODUCTIONS IS A MATTER OF OBVIOUS PRACTICAL IMPORTANCE 4O MAKE SUCH
PREDICTIONS WE NEED TO UNDERSTAND HOW THE NUMERICAL DYNAMICS OF A HOST PLANT
FOLLOWING INTRODUCTION OF ITS BIOCONTROL AGENT DEPEND ON HOST DENSITY DISEASE
INCIDENCE GENETIC VARIATION AND EVOLUTIONARY CHANGES IN VIRULENCE OR RESISTANCE
3URPRISINGLY WHILE THERE ARE SOME CASES FOR WHICH WE HAVE GOOD INFORMATION ON
THE DYNAMICS OF HOST NUMBERS AFTER RELEASE OF A CONTROL AGENT EG (ASAN AND
!YRES -ORRIS FOR MANY OTHER RELEASES THE DETAILS OF CHANGES ARE NOT
WELL DOCUMENTED )N PARTICULAR WE SHOULD ASK I IS CONTROL MORE SUCCESSFUL IN
GENETICALLY DEPAUPERATE WEEDS II $O TRANSMISSION RATE AND DEMOGRAPHIC IMPACT
OF THE PATHOGEN ATTENUATE AS THE HOST POPULATION DECLINES III $O PATHOGEN AND
HOST REACH A STABLE EQUILIBRIUM OR ARE THEY DEPENDENT ON METAPOPULATION DYNAM
ICS TO PERSIST IN THE LANDSCAPE $ETAILED INFORMATION ON NUMERICAL DYNAMICS IN
BIOLOGICAL CONTROL SYSTEMS IS SCARCE BUT DATA ARE NEARLY NONEXISTENT FOR LONG
TERM
0LANT POPULATIONS BY PATHOGENS
GENETIC CHANGES IN THE HOST OR PATHOGEN 4HERE IS GREAT UNTAPPED POTENTIAL FOR
BIOLOGICAL CONTROL INTRODUCTIONS TO BE USED TO UNDERSTAND THE FACTORS THAT DRIVE
THE ECOLOGICAL END EVOLUTIONARY DYNAMICS OF THE PLANT
PATHOGEN INTERACTIONS
)N FACT THE ONLY BIOCONTROL STUDY WE FOUND THAT TRACKED CHANGES IN PATHOGEN
VIRULENCE OR HOST RESISTANCE FOR A PLANT
PATHOGEN SYSTEM WAS FOR THE RUST 0UCCINIA
CHONDRILLINA ON #HONDRILLA JUNCEA SEE CASE STUDY BELOW
%VOLUTIONARY ECOLOGY OF BIOCONTROL WITH PATHOGENS
4HE EVOLUTIONARY DYNAMICS OF INTERACTIONS BETWEEN WEEDS AND PATHOGEN BIOCON
TROL AGENTS HAVE IMPORTANT IMPLICATIONS FOR THE LONG
TERM SUCCESS OF BIOLOGICAL
CONTROL PROGRAMS 'ENERALLY WE NEED TO KNOW I HAS VIRULENCE OF THE BIOLOGICAL
CONTROL PATHOGEN CHANGED OVER TIME AND HAS THIS INCREASED OR DECREASED THE
SUCCESS OF CONTROL II (AS THE HOST DEVELOPED RESISTANCE OVER TIME III (AS HOST
SPECIlCITY CHANGED OVER TIME 2ECENT ANALYSIS OF EMERGING DISEASES HAS SUGGESTED
THAT ECOLOGICAL HOST SHIFTS IE HAVING A PREADAPTED ABILITY TO USE A NEWLY ENCOUN
TERED HOST MAY PREDOMINATE AS CAUSES OF NOVEL EPIDEMICS AND THAT HOST SHIFTS MAY
ONLY RARELY BE CAUSED BY MUTATIONS THAT ALLOW COLONIZATION OF A NEW HOST 3CHRAG
AND 7IENER (OWEVER THE DIFlCULTY OF OBSERVING SUCH GENETIC EVENTS MAY
DISTORT OUR PERSPECTIVE 6AN +LINKEN AND %DWARDS SYNTHESIZED INFORMATION
ON HOST RANGE FROM BIOLOGICAL CONTROL PROGRAMS USING HERBIVORES 4HEY FOUND
THAT HOST SHIFTS WERE MORE QUANTITATIVE THAN QUALITATIVE THAT IS WHILE PREFERENCE
AND EFlCIENCY ON NOVEL HOSTS EVOLVED THERE WAS NO EVIDENCE OF EVOLUTIONARY
CHANGES IN FUNDAMENTAL HOST
RANGE 3UCH AN ANALYSIS SHOULD BE DONE FOR PATHO
GENS USED AS BIOLOGICAL CONTROL AGENTS 7HILE PATHOGENS ARE GENERALLY THOUGHT TO
OFFER THE OPPORTUNITY FOR HIGH HOST SPECIlCITY SOME HAVE ARGUED THAT HIGH HOST
SPECIlCITY MAY BE CORRELATED WITH EVOLUTIONARY LABILITY "ROOKS AND -C,ENNAN
3ECORD AND +AREIVA "ECAUSE THEY ARE UNABLE TO SIMPLY MOVE
FROM AN UNACCEPTABLE HOST TO A MORE SUITABLE ONE AS AN ANIMAL MIGHT SPECIALIST
PATHOGENS MAY EXPERIENCE EVEN STRONGER SELECTION FOR HOST SHIFTS THAN HERBIVORES
2OY +NOWING THE FREQUENCY OF EVOLUTIONARY HOST SHIFTS IN PATHOGENS AND
UNDERSTANDING THE CONDITIONS UNDER WHICH THEY OCCUR ARE CRITICAL TO THE PROCESS
OF RISK ASSESSMENT IN BIOLOGICAL CONTROL 3ECORD AND +AREIVA
'ENERAL THEORY OF HOST
PATHOGEN INTERACTIONS HAS PLAYED A LARGE ROLE IN THE
CHOICE OF BIOLOGICAL CONTROL AGENTS IN THE PAST -C&ADYEN &OR EXAMPLE
IT HAS LONG BEEN THOUGHT THAT SEXUALLY REPRODUCING WEEDS WOULD BE HARDER TO
CONTROL BECAUSE THEIR HIGHER LEVELS OF GENETIC VARIATION INTERFERE WITH PATHOGEN
POPULATION GROWTH "URDON ET AL HOWEVER A MORE RECENT ANALYSIS HAS
DISPUTED THIS ASSERTION #HABOUDEZ AND 3HEPPARD 3IMILARLY THE BELIEF
THAT PATHOGEN POPULATIONS ARE LOCALLY ADAPTED TO THEIR HOST GENOTYPES HAS HAD A
LARGE INmUENCE ON THE PROCESS OF SELECTION OF CONTROL AGENTS WITH GENETIC ANALYSIS
PLAYING AN INCREASING ROLE IN THE CAREFUL MATCHING OF AGENT GENOTYPES WITH THE
POPULATIONS OF ORIGIN FOR THE WEED EG (OLDEN AND -AHLBERG (OWEVER
THE EVIDENCE FOR CLOSE LOCAL ADAPTATION OF PATHOGENS TO THEIR HOST POPULATIONS IS
'3 'ILBERT AND )- 0ARKER
MIXED AND IN FACT RESISTANCE AND GENE FOR GENE VIRULENCE SHOULD mUCTUATE IN AN
ASYNCHRONOUS FREQUENCY DEPENDENT WAY REVIEWED IN 0ARKER AND 'ILBERT
"IOLOGICAL CONTROL PRACTITIONERS HAVE CLEARLY BASED THEIR INTRODUCTION STRATEGIES ON
THEORETICAL CONSIDERATIONS BUT THE SIMPLER RULES ARE NOW COMING UNDER QUESTION
-C&ADYEN 4HERE IS A NEED FOR CLEAR PREDICTIONS AND MODERN EMPIRICAL
WORK TESTING THOSE PREDICTIONS TO HELP INFORM THE PRACTICE OF BIOLOGICAL CONTROL
FOR THE FUTURE
!N EVOLUTIONARY ECOLOGY CASE STUDY #HONDRILLA AND 0UCCINIA
ON THREE CONTINENTS
.O SYSTEM OF CLASSICAL BIOLOGICAL CONTROL OF A WEED BY A PATHOGEN DEMONSTRATES
THE IMPORTANCE OF INTERACTIONS BETWEEN ENVIRONMENT GENETICS AND NUMERICAL
DYNAMICS AS WELL AS THAT OF #HONDRILLA JUNCEA RUSH SKELETONWEED !STERACEAE
.ATIVE TO %URASIA THIS SPECIES WAS INTRODUCED TO THE EASTERN 5NITED 3TATES IN THE
LATE S TO !USTRALIA IN THE EARLY S AND TO THE WESTERN 5NITED STATES IN
THE S -C6EAN 0RYOR 3CHIRMAN AND 2OBOCKER 3UPKOFF
ET AL #HONDRILLA IS A SIGNIlCANT ECONOMIC PROBLEM IN WHEAT
GROWING
REGIONS 0ANETTA AND $ODD AND IS ALSO A WIDESPREAD RANGELAND WEED
/F THREE BIOCONTROL AGENTS THAT WERE INTRODUCED 0UCCINIA CHONDRILLINA WAS THE MOST
EFFECTIVE AT REDUCING PLANT VIGOR 3UPKOFF ET AL AND WITHIN TWO DECADES OF
ITS INTRODUCTION INTO !USTRALIA AND #ALIFORNIA #HONDRILLA DENSITIES WERE REDUCED TO
THOSE TYPICAL IN ITS NATIVE RANGE 7APSHERE ET AL #ULLEN ET AL 3UPKOFF
ET AL
4HE EXTREME HOST SPECIlCITY FOUND IN THE #HONDRILLA0UCCINIA SYSTEM MAKES THIS
EXAMPLE PARTICULARLY INTERESTING (ASAN #HONDRILLA IS A TRIPLOID APOMICT
AND THUS REPRODUCTION IS CLONAL )N !USTRALIA THREE CLONAL TYPES ARE PRESENT EACH
WITH A DIFFERENT LEAF WIDTH 0 CHONDRILLINA CAUSES DISEASE ONLY ON THE NARROW
LEAF
TYPE 4HE ORIGINAL INFESTATION OF #HONDRILLA WAS PRIMARILY THIS NARROW
LEAF TYPE
BUT AFTER THE SUCCESSFUL BIOCONTROL OF THAT CLONE THE INTERMEDIATE
LEAVED CLONE HAS
SPREAD (ANLEY AND 'ROVES .OW THERE IS CONCERTED EFFORT TO INTRODUCE NEW
STRAINS OF 0UCCINIA CHONDRILLINA THAT ARE ABLE TO ATTACK AND CONTROL THE OTHER CLONES
OF #HONDRILLA (ANLEY AND 'ROVES
)N WESTERN .ORTH !MERICA THREE DIFFERENT #HONDRILLA GENOTYPES WERE FOUND
DISTINGUISHABLE BY THEIR MULTI
LOCUS ISOZYME PHENOTYPES (ASAN ET AL
TWO GENOTYPES ARE THOUGHT TO HAVE ORIGINATED IN 9UGOSLAVIA (ASAN AND $ELFOSSE
4HE WESTERN 53 GENOTYPES WERE LARGELY RESISTANT TO THE RUST STRAIN THAT
CONTROLLED THE NARROW
LEAF #HONDRILLA IN !USTRALIA SO ADDITIONAL RUST ISOLATES WERE
EVALUATED FOR USE IN BIOCONTROL 2USTS COLLECTED FROM 9UGOSLAVIA THE PUTATIVE
SITE OF ORIGIN OF THE 53 INVADERS SHOWED HIGH VIRULENCE ON SOME OF THE 53 GENO
TYPES BUT OTHER PLANTS WERE LITTLE AFFECTED )N ADDITION RUSTS FROM OTHER REGIONS
ALSO SHOWED HIGH VIRULENCE 'ENETIC MATCHING OF HOSTS BETWEEN THE NATIVE AND
INTRODUCED RANGE MAY YIELD WELL
ADAPTED BIOCONTROL PATHOGENS WHEN THE TARGET
WEED IS GENETICALLY UNIFORM BUT THIS APPROACH ASSUMES HIGH LOCAL ADAPTATION OF
0LANT POPULATIONS BY PATHOGENS
THE PATHOGEN TO THE HOST !S MENTIONED ABOVE SUCH LOCAL ADAPTATION MAY NOT BE
GENERALLY FOUND IN NATURAL POPULATIONS
3EXUAL REPRODUCTION IN BOTH THE HOST AND PATHOGEN PLAY A ROLE IN THIS STORY
4ELIOSPORES OF 0 CHONDRILLINA ONLY GERMINATE AFTER COLD WINTERS !DAMS AND ,INE
SO SEXUAL RECOMBINATION OF THE PATHOGEN OCCURS READILY ONLY IN COLDER
REGIONS SUCH AS PARTS OF EASTERN CONTINENTAL %UROPE )N THESE AREAS OF THE NATIVE
RANGE #HONDRILLA ALSO SHOWS HIGHER CLONAL DIVERSITY AND POSSIBLY DIPLOID SEXUAL
POPULATIONS #HABOUDEZ AND 3HEPPARD )NTERESTINGLY POPULATIONS OF
#HONDRILLA WITH HIGH VS LOW CLONAL DIVERSITY IN THE NATIVE RANGE APPEAR TO SUFFER
SIMILAR DEGREES OF RUST INFECTION #HABOUDEZ AND 3HEPPARD )N NORTHERN
.ORTH !MERICA WHERE COLD WINTERS ALLOW FOR SEXUAL RECOMBINATION OF THE PATHO
GEN THE EVOLUTIONARY DYNAMICS OF THE HOST
PATHOGEN INTERACTION ARE POTENTIALLY
MORE COMPLEX (ASAN ET AL )N !USTRALIA THEY ARE PARTICULARLY CONCERNED
ABOUT THE POSSIBILITY OF INTRODUCTION OF NEW SEXUAL TYPES OF #HONDRILLA WHICH
COULD RESULT IN A SITUATION WHERE THE HOST REPRODUCES SEXUALLY BUT THE PATHOGEN
CAN NOT #HABOUDEZ AND 3HEPPARD
4HIS EXAMPLE SHOWS THE IMPORTANCE OF THE hDISEASE TRIANGLEv INTERACTION
BETWEEN HOST PATHOGEN AND ENVIRONMENT IN THE DYNAMICS OF PLANT
PATHOGEN
INTERACTIONS 4HE CASE OF #HONDRILLA 0UCCINIA
CHONDRILLINA IS ONE OF THE FEW FOR
WHICH WE HAVE SUCH DETAILED INFORMATION ABOUT THE HOSTS AND THE PATHOGENS
GENETIC MAKEUP AND MORE IMPORTANTLY ABOUT HOW THE INTERACTION HAS PLAYED
OUT IN A NUMBER OF DIFFERENT REGIONS 0ATHOGEN BIOLOGICAL CONTROL CASES OFFER THE
POTENTIAL TO LEARN A GREAT DEAL ABOUT THE ROLES OF GENETICS NUMERICAL DYNAMICS
EVOLUTIONARY DYNAMICS AND ENVIRONMENTAL FACTORS IN DETERMINING THE LONG
TERM
OUTCOMES OF HOST
PATHOGEN INTERACTIONS
#/.#,53)/.3
&ROM THE STUDIES DESCRIBED ABOVE IT APPEARS THAT PATHOGENS ARE SOMETIMES IMPOR
TANT IN THE REGULATION OF NATURAL PLANT POPULATIONS MAY CONSTRAIN POPULATIONS
INTRODUCED TO NEW REGIONS AND MAY BY THEIR ABSENCE RELEASE INTRODUCED PLANTS
FROM AN IMPORTANT SOURCE OF REGULATION 7E SUGGEST TWO KEY DIRECTIONS THAT WOULD
MOST ADVANCE OUR UNDERSTANDING OF THE IMPORTANCE OF PATHOGENS IN PLANT POPULA
TION REGULATION AN INTEGRATION OF NUMERICAL AND EVOLUTIONARY DYNAMICS FOR BOTH
THE PATHOGENS AND PLANTS AND A GREATER BREADTH OF STUDIES TO INCLUDE MORE PLANT
PATHOGEN SYSTEMS
"IOLOGICAL CONTROL OF INVASIVE WEEDS OFFERS EXCITING OPPORTUNITIES TO EVALUATE THE
IMPORTANCE OF PATHOGENS IN NUMERICAL REGULATION OF PLANT POPULATIONS AND AT THE
SAME TIME TO FOLLOW GENETIC CHANGES IN PLANT AND PATHOGEN POPULATIONS #AREFUL
MONITORING FOR CHANGES IN HOST AND PATHOGEN GENOTYPES ALONG WITH NUMERI
CAL DYNAMICS SHOULD BE INTEGRAL TO ANY INTRODUCTION OF PATHOGENS FOR BIOLOGICAL
CONTROL OF INVASIVE WEEDS %QUALLY IMPORTANT WE MUST MOVE BEYOND SPINNING
NARRATIVES AND MAKING GENERAL PREDICTIONS FROM A HANDFUL OF EXAMPLES 7E HAVE
'3 'ILBERT AND )- 0ARKER
A GOOD IDEA OF THE ROLE PATHOGENS SHOULD PLAY IN PLANT POPULATION REGULATION AND
BIOLOGICAL INVASIONS 4ABLE WE NOW NEED TO COLLECT DATA FROM A DIVERSITY OF SYS
TEMS TO EVALUATE OUR PREDICTIONS (OW OFTEN AND UNDER WHAT CONDITIONS ARE PLANT
PATHOGENS SIGNIlCANT FORCES IN REGULATING WILD PLANT POPULATIONS IN THEIR NATIVE
RANGES (OW OFTEN DO INTRODUCED PLANTS FAIL TO ESTABLISH BECAUSE THEY ARE ATTACKED
BY LOCAL PATHOGENS (OW OFTEN IS ESCAPE FROM PATHOGENS A KEY TO DETERMIN
ING WHETHER AN INTRODUCED PLANT INVADES NATURAL HABITATS 4HROUGH PATHOGEN
EXCLUSION AND ADDITION EXPERIMENTS COMMON GARDENS PHYLOGENETIC ANALYSIS OF
HOST RANGES ANALYSIS OF RAPID EVOLUTIONARY CHANGES IN PLANT
PATHOGEN INTERAC
TIONS AND CAREFUL CREATIVE NATURAL HISTORY OF PLANT DISEASES WE WILL ILLUMINATE
THE ROLE OF PLANT PATHOGENS IN BIOLOGICAL INVASIONS
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0HYTOPATHOLOGY
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ANIMALS AND PLANTS .ATURE
!LEXANDER ( - AND 2 $ (OLT 4HE INTERACTION BETWEEN PLANT COMPETITION AND
DISEASE 0ERSPECTIVES IN 0LANT %COLOGY %VOLUTION AND 3YSTEMATICS
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"ISHOP * ' ! - $EAN AND 4 -ITCHELL
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MOLECULAR TARGETS OF SELECTION IN PLANT
PATHOGEN COEVOLUTION 0ROC .ATL !CAD 3CI 53!
0LANT POPULATIONS BY PATHOGENS
"LANEY # 3 AND 0 - +OTANEN %FFECTS OF FUNGAL PATHOGENS ON SEEDS OF NATIVE AND
EXOTIC PLANTS ! TEST USING CONGENERIC PAIRS * !PPL %COL
"LOSSEY " AND 2 .TZOLD %VOLUTION OF INCREASED COMPETITION ABILITY IN INVASIVE
NONINDIGENOUS PLANTS A HYPOTHESIS *OURNAL OF %COLOGY
"ROOKS $ 2 AND $ ! -C,ENNAN 0ARASCRIPT PARASITES AND THE LANGUAGE OF EVOLU
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'3 'ILBERT AND )- 0ARKER
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7OLFE , - 7HY ALIEN INVADERS SUCCEED 3UPPORT FOR THE ESCAPE
FROM
ENEMY
HYPOTHESIS !MERICAN .ATURALIST
7RIGHT 3 * 0LANT DIVERSITY IN TROPICAL FORESTS A REVIEW OF MECHANISMS OF SPECIES
COEXISTENCE /ECOLOGIA
#HAPTER FOURTEEN
%XPLORING THE RELATIONSHIP
BETWEEN NICHE BREADTH
AND INVASION SUCCESS
$ 0 6ZQUEZ
).42/$5#4)/.
4HE ULTIMATE GOAL OF INVASIONS BIOLOGY SHOULD BE TO GENERATE A BODY OF GENERAL
PREDICTIVE THEORY )DEALLY SUCH THEORY WOULD HELP DETERMINE WHETHER A SPECIES
WITH A PARTICULAR SET OF TRAITS WILL BE ABLE TO INVADE A GIVEN SYSTEM )N ITS ALMOST
lFTY YEARS THE DISCIPLINE HAS SEEN MANY ATTEMPTS TO PROVIDE SUCH PREDICTABILITY
THROUGH A FOCUS ON THE TRAITS OF THE POTENTIAL INVADERS AND THOSE OF THE INVADED
COMMUNITY 2EGRETTABLY MOST ATTEMPTS HAVE BEEN SO FAR UNSUCCESSFUL
/NE ATTEMPT AT GENERALIZATION THAT HAS BEEN OFTEN DISCUSSED IN THE LITERATURE
IS THE IDEA THAT NICHE BREADTH IS POSITIVELY CORRELATED WITH INVASION SUCCESS SO
THAT SPECIES WITH BROAD NICHES hGENERALISTSv ARE MORE LIKELY TO INVADE THAN SPE
CIES WITH NARROWER NICHES hSPECIALISTSv BECAUSE THEY ARE MORE LIKELY TO lND THE
NECESSARY RESOURCES OR ENVIRONMENTAL CONDITIONS THROUGHOUT THIS PAPER ) WILL
REFER TO THIS NOTION AS THE hNICHE BREADTHnINVASION SUCCESS HYPOTHESISv !S WITH
MANY OTHER IDEAS IN ECOLOGY IT IS DIFlCULT TO TRACK THE HISTORICAL DEVELOPMENT
OF THIS HYPOTHESIS 0ROBABLY ONE OF THE lRST TO INCLUDE SOME MEASURE OF NICHE
BREADTH IN THINKING ABOUT INVASION SUCCESS WAS ( ' "AKER "AKER
DElNED THE TRAITS OF WHAT HE CALLED THE hIDEAL WEEDv SPECIES PRONE TO BECOME
-7 #ADOTTE ET AL EDS #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY n
3PRINGER 0RINTED IN THE .ETHERLANDS
$0 6ZQUEZ
WEEDY AND INVADE DISTURBED HABITATS !MONG THESE TRAITS THREE WERE RELATED TO
NICHE BREADTH NAMELY I THAT THE SPECIES CAN PRODUCE SEEDS IN A WIDE RANGE OF
ENVIRONMENTAL CIRCUMSTANCES AND HAS HIGH TOLERANCE OF AND OFTEN PLASTICITY IN
FACE OF CLIMATIC AND ENVIRONMENTAL VARIATION II THAT THE SPECIES HAS NO SPECIAL
ENVIRONMENTAL REQUIREMENTS FOR GERMINATION IE WIDE ENVIRONMENTAL NICHE
AND III THAT WHEN CROSS POLLINATED POLLINATION CAN BE ACHIEVED BY A NONSPECIAL
IZED mOWER VISITOR OR BY WIND IE WIDE POLLINATION NICHE NOTE HOWEVER THAT IT
IS ASSUMING WIDE NICHE BREADTH OF POLLINATORS NOT OF THE PLANT WHICH BRINGS UP
THE ISSUE OF ASYMMETRIC SPECIALIZATION DISCUSSED BELOW
-ORE RECENTLY 2EJMNEK PROPOSED IDEAS SIMILAR TO THOSE OF "AKERS IN
THE BROADER CONTEXT OF PLANT INVASIONS IN GENERAL (E DESCRIBED WHAT HE THINKS
ARE TEN MAJOR PREDICTORS OF PLANT INVASIVENESS THREE OF WHICH ARE RELATED TO NICHE
BREADTH I hlTNESS HOMEOSTASIS v THE ABILITY OF AN INDIVIDUAL OR POPULATION TO
MAINTAIN RELATIVELY CONSTANT lTNESS OVER A RANGE OF ENVIRONMENTS lTNESS HOMEO
STASIS WILL DEPEND ON PHENOTYPIC PLASTICITY HOMEOSTASIS AT THE INDIVIDUAL LEVEL
AND POPULATION GENETIC POLYMORPHISM HOMEOSTASIS AT THE POPULATION LEVEL
II SIZE OF hPRIMARYv NATIVE GEOGRAPHICAL RANGES SEE ALSO $AEHLER AND 3TRONG
WHICH ULTIMATELY DEPENDS ON POPULATION lTNESS HOMEOSTASIS AND III
SPECIlCITY OF MUTUALISMS SO THAT SPECIES THAT DEPEND ON NON
SPECIlC MUTUALISMS
ARE MORE LIKELY TO OVERCOME MANY ABIOTIC AND BIOTIC BARRIERS IN NEW ENVIRONMENTS
SEE ALSO 2ICHARDSON ET AL
-ANY OTHERS HAVE DISCUSSED THE IDEA THAT BROADER NICHES CONFER HIGHER INVA
SION SUCCESS &OR EXAMPLE IN A REVIEW ON BIOTIC HOMOGENIZATION -C+INNEY AND
,OCKWOOD CONCLUDED THAT EXTINCTIONnRESISTANT SPECIES AND SUCCESSFUL
INVADERS ARE CHARACTERIZED BY OMNIVORY RAPID GROWTH AND DISPERSAL OR BREEDING
IN EPHEMERAL HABITATS WHEREAS EXTINCTION
PRONE GROUPS HAVE A PREDOMINANCE
OF TRAITS ASSOCIATED WITH SPECIALIZATION SLOW REPRODUCTION AND OTHER TRAITS NOT
ASSOCIATED WITH OPPORTUNISM 3IMILARLY 2ICHARDSON ET AL STATED THAT
h)T IS LOGICAL TO PREDICT THAT PLANTS WITH SPECIALIZED POLLINATION SYNDROMES ARE LESS
LIKELY TO BE POLLINATED IN FOREIGN ENVIRONMENTS THAN PLANTS WITHOUT SPECIALIZED
REQUIREMENTSv P ,IKEWISE 2ICCIARDI AND 2ASMUSSEN REGARD BROAD
GEOGRAPHIC ENVIRONMENTAL AND DIETARY RANGES AS GENERAL ATTRIBUTES OF INVASIVE
AQUATIC SPECIES !LTHOUGH THESE IDEAS ARE INTERESTING AND APPEALING THEIR EVALU
ATION IS COMPLICATED BY SEVERAL CONCEPTUAL AND METHODOLOGICAL PROBLEMS ) WILL
ARGUE THAT PROGRESS TOWARDS A PREDICTIVE THEORY OF INVASIONS WILL BE DIFlCULT UNLESS
THESE ISSUES ARE RESOLVED
"ELOW ) PRESENT A REVIEW OF THE LITERATURE SUMMARIZING SEVERAL STUDIES THAT
HAVE EVALUATED THE RELATIONSHIP BETWEEN DIFFERENT ASPECTS OF NICHE BREADTH AND
INVASION SUCCESS ) THEN PROVIDE A CRITICAL EXAMINATION OF THE MECHANISMS POTEN
TIALLY INVOLVED IN GENERATING THESE RESULTS STRESSING SEVERAL CONCEPTUAL ISSUES
THAT HAVE BEEN OVERLOOKED IN PREVIOUS DISCUSSIONS -Y REVIEW IS NOT EXHAUS
TIVE AND IS TAXONOMICALLY BIASED FOCUSING ON PLANTS BIRDS AND HETEROTROPHIC
MUTUALISTS OF PLANTS 4HIS BIAS REmECTS THE FACT THAT MOST STUDIES EVALUATING THIS
HYPOTHESIS HAVE DEALT WITH THESE GROUPS
)N SUMMARIZING THE AVAILABLE
LITERATURE
.ICHE BREADTH AND INVASION SUCCESS
) USE A VOTE
COUNTING APPROACH BECAUSE THE NUMBER OF STUDIES AVAILABLE PER
TAXON AND MEASURE OF NICHE BREADTH WAS TOO SMALL TO ALLOW THE APPLICATION OF
MORE QUANTITATIVE META
ANALYTICAL TECHNIQUES SEE !RNQVIST AND 7OOSTER
'UREVITCH ET AL
4HE STUDIES REVIEWED HERE ARE BASED ON THREE DIFFERENT KINDS OF COMPARISONS
SOME MEASURE OF NICHE BREADTH IN NATIVE REGION OF SUCCESSFUL VS FAILED INVADERS
3) VS &) NICHE BREADTH OF INVADERS IN THEIR NATIVE VS THEIR INTRODUCED REGIONS
.2 VS )2 AND NICHE BREADTH OF NATIVE VS INTRODUCED SPECIES IN THE INTRODUCED
REGION .3 VS )3 )T IS IMPORTANT TO NOTE SEVERAL SIMILARITIES AND DIFFERENCES ABOUT
THE THREE APPROACHES 4HE lRST COMPARISON 3) VS &) USES A BINARY MEASURE OF
INVASION SUCCESS ASKING WHETHER SUCCESS IS EXPLAINED BY THE NICHE BREADTH IN
THE NATIVE RANGE )N CONTRAST THE SECOND APPROACH .2 VS )2 USES A QUANTITA
TIVE MEASURE OF SUCCESS SUCH AS THE SIZE OF THE INTRODUCED RANGE TO DETERMINE
WHETHER INVASION SUCCESS IS EXPLAINED BY NICHE BREADTH IN THE NATIVE RANGE
4HE THIRD APPROACH .3 VS )3 COMPARES NICHE BREADTH OF INTRODUCED SPECIES WITH
NATIVE SPECIES IN THE INTRODUCED RANGE IE SPECIES NATIVE TO THE RECIPIENT COMMU
NITY HERE THE QUESTION IS WHETHER THE SUCCESS OF INVADERS IE THE FACT THAT THEY
ESTABLISHED IN A NEW ENVIRONMENT IS EXPLAINED BY THEIR BROADER NICHE COMPARED
TO THAT OF NATIVE SPECIES IN SHARP CONTRAST WITH THE QUESTION ASKED IN THE PREVI
OUS TWO APPROACHES 4HUS ALTHOUGH THE THREE TYPES OF COMPARISONS ARE INTENDED
TO ADDRESS THE QUESTION OF WHETHER NICHE BREADTH OF POTENTIAL INVADERS IS RELATED
TO THEIR INVASION SUCCESS THEY ARE NOT EQUIVALENT AND ARE LIKELY TO PROVIDE DIFFER
ENT ANSWERS
-Y REVIEW CONSISTS OF THREE MAIN PARTS I THE ROLE OF GEOGRAPHIC HABITAT AND
CLIMATIC RANGE ON PLANT AND BIRD INVASIONS II THE ROLE OF MUTUALISMS ON PLANT
AND POLLINATOR INVASIONS AND III THE ROLE OF TROPHIC BREADTH ON BIRD INVASIONS
) END WITH A SECTION SUMMARIZING THE RESULTS OF THE REVIEW AND OFFERING CONCLUD
ING REMARKS
4(% 2/,% /& '%/'2!0()# #,)-!4)# !.$ (!")4!4 2!.'%
0ATTERNS STUDIES ON PLANTS AND BIRDS
3TUDIES THAT HAVE EVALUATED THE INVASION SUCCESS OF PLANTS AND ITS RELATIONSHIP
TO GEOGRAPHIC CLIMATIC OR HABITAT RANGE ARE BASED ON THE THREE TYPES OF COM
PARISONS OUTLINED ABOVE 3TUDIES BASED ON THE lRST TWO KINDS OF COMPARISONS
3) VS &) AND .2 VS )2 HAVE FOUND THAT INVASION SUCCESS IS POSITIVELY RELATED
TO NATIVE GEOGRAPHIC CLIMATIC OR HABITAT RANGE SO THAT MORE SUCCESSFUL INVAD
ERS TEND TO HAVE BROADER NATIVE RANGES THAN LESS SUCCESSFUL INVADERS 4ABLE
)N GENERAL THESE RESULTS SUPPORT THE NOTION THAT THE NATIVE RANGE OF PLANT SPECIES
IS A PREDICTOR OF THEIR SUCCESS AS INVADERS IN NEW REGIONS WHERE THEY DID NOT OCCUR
NATURALLY $AEHLER AND 3TRONG 2EJMNEK
4ABLE 3TUDIES EVALUATING THE RELATIONSHIP BETWEEN BREADTH OF GEOGRAPHIC CLIMATIC OR HABITAT RANGE OF PLANT SPECIES AND INVASION SUCCESS
4YPE OF -AJOR TAXON .ATIVE REGION )NTRODUCED -EASURE OF SUCCESS -EASURE OF NICHE 2ELAT WITH 2EF
COMPARISON
REGION BREADTH NICHE BREADTHe
3) VS &) (IGHER PLANTS #ENTRAL -ENDOZA AND %STABLISHMENT IN .O NATIVE HABITAT TYPES 0OSITIVE
%UROPE "UENOS !IRES NEW REGION
PROVINCES !RGENTINA
!NGIOSPERMS %UROPE .EW "RUNSWICK %STABLISHMENT IN 3IZE OF NATIVE RANGE 0OSITIVE
#ANADA NEW REGION
.2 VS )2 #OMPOSITAE %URASIA .ORTH 3IZE OF INTRODUCED .O NATIVE HABITAT TYPES 0OSITIVE
'RAMINEAE !MERICA RANGE
!NGIOSPERMS 3OUTH !FRICA !USTRALIA 7EED STATUS .O NATIVE HABITAT TYPES 0OSITIVE
!NGIOSPERMS 3OUTH !FRICA !USTRALIA 7EED STATUS .O CLIMATIC ZONES 0OSITIVE
$0 6ZQUEZ
"ROMUS -EDITERRANEAN -EDITERRANEAN .O CLIMATIC ZONES .O -EDITERRANEAN 0OSITIVE
REGIONS REGIONS IN NATIVE RANGE REGIONS OCCUPIED
.3 VS )3 (IGHER PLANTS 7ORLD(AWAII (AWAII .O OF HABITAT .O HABITAT TYPES .EGATIVE
TYPES OCCUPIED
2EFERENCES 0RINZING ET AL 2EJMNEK 3COTT AND 0ANETTA 'OODWIN ET AL 2OY ET AL
+ITAYAMA AND -UELLER $OMBOIS
#OMPARISONS INCLUDED 3) VS &) NATIVE RANGE OF SUCCESSFULLY ESTABLISHED SPECIES VS UNSUCCESSFULLY ESTABLISHED SPECIES .2 VS )2 NATIVE
RANGE VS INTRODUCED RANGE OF INTRODUCED SPECIES .3 VS )3 RANGE OF NATIVE VS INTRODUCED SPECIES IN THE INTRODUCED REGION
e
)NDICATES STATISTICAL SIGNIlCANCE OF TEST AND DIRECTION OF EFFECT 0OSITIVE SIGNIlCANT EFFECT IN THE DIRECTION EXPECTED BY THE SPECIALIZATIONnNICHE
BREADTH HYPOTHESIS .EGATIVE SIGNIlCANT EFFECT IN THE DIRECTION OPPOSITE TO THE EXPECTED BY THE SPECIALIZATIONnNICHE BREADTH HYPOTHESIS
. I CHE BREADTH AND INVASION SUCCESS
)N CONTRAST THE ONE STUDY COMPARING BREADTH OF HABITAT USE BY NATIVE AND
INTRODUCED SPECIES .3 VS )3 4ABLE FOUND A PATTERN OPPOSITE TO THE PREDICTION
OF THE NICHE BREADTHnINVASION SUCCESS HYPOTHESIS INTRODUCED SPECIES WERE FOUND
IN FEWER HABITAT TYPES THAN NATIVE SPECIES 4HIS RESULT IS DIFlCULT TO INTERPRET
HOWEVER MAINLY BECAUSE THE NUMBER OF HABITATS IN WHICH A SPECIES IS FOUND WILL
DEPEND ON ITS DISPERSAL ABILITY AND THE TIME ELAPSED AFTER ITS ARRIVAL TO THE ENVIRON
MENT WHICH MAY OBVIOUSLY BIAS THE RESULTS TOWARDS NATIVE SPECIES GIVEN THEIR
LONGER PRESENCE IN THE STUDY AREA
3TUDIES EVALUATING THE RELATIONSHIP BETWEEN GEOGRAPHIC CLIMATIC OR HABITAT
RANGE AND INVASION SUCCESS IN BIRDS 4ABLE ALL USED THE lRST APPROACH OUTLINED
ABOVE THAT IS THEY COMPARE THE SIZE OF THE NATIVE RANGE BETWEEN SUCCESSFUL AND
UNSUCCESSFUL INVADERS 3) VS &) 4HESE STUDIES HAVE USED EITHER SIZE OF NATIVE GEO
GRAPHIC RANGE NUMBER OF NATIVE HABITAT TYPES OR CLIMATIC ZONES OCCUPIED IN THE
NATIVE REGION AS MEASURES OF NICHE BREADTH
)N AGREEMENT WITH STUDIES ON PLANTS DISCUSSED ABOVE STUDIES OF BIRD INVA
SIONS IN (AWAII !USTRALIA AND WORLDWIDE FOUND A POSITIVE RELATIONSHIP BETWEEN
INVASION SUCCESS AND SIZE OF NATIVE GEOGRAPHIC RANGE USED IN THE NATIVE RANGE
4ABLE (OWEVER TWO STUDIES ONE IN FOUR OCEANIC ISLANDS AND THE OTHER IN
.EW :EALAND FAILED TO lND EVIDENCE FOR SUCH AN EFFECT 4ABLE 4HE TWO STUD
IES USING THE NUMBER OF HABITAT TYPES OCCUPIED IN THE NATIVE RANGE FOUND THAT
SUCCESSFULLY INVADING SPECIES OR FAMILIES HAD SIGNIlCANTLY GREATER NATIVE HABITAT
RANGES THAN SPECIES OR FAMILIES THAT FAILED TO INVADE 4ABLE
)N SUMMARY IT APPEARS THAT BREADTH OF NATIVE GEOGRAPHIC CLIMATIC AND HABITAT
RANGE IS RELATED TO THE INVASION SUCCESS OF PLANT AND BIRD SPECIES !LTHOUGH THERE
ARE EXCEPTIONS TO THESE PATTERNS THEY APPEAR TO HAVE SOME DEGREE OF GENERALITY
-ECHANISMS
7HY DOES BREADTH OF NATIVE GEOGRAPHIC HABITAT OR CLIMATE RANGE AFFECT INVASION
SUCCESS 3PECIES WITH BROAD RANGES MAY BE MORE SUCCESSFUL INVADERS THAN SPE
CIES WITH NARROWER RANGES FOR SEVERAL REASONS !S DISCUSSED IN THE INTRODUCTION
SPECIES NICHES MAY DETERMINE THEIR DISTRIBUTION SO THAT SPECIES WITH BROADER
NICHES TEND TO HAVE BROADER DISTRIBUTIONS AND ARE SO MORE LIKELY TO INVADE NEW
ENVIRONMENTS SEE ALSO "ROWN
(OWEVER OTHER FACTORS NOT RELATED TO NICHE BREADTH MAY ALSO AFFECT SPECIES
DISTRIBUTIONS 5NDER THESE SCENARIOS WIDESPREAD SPECIES WOULD HAVE HIGHER
INVASION SUCCESS REGARDLESS OF THEIR NICHE BREADTH &IRST SPECIES SPECIALIZED ON
WIDESPREAD RESOURCES OR SPECIES THAT ARE ABLE TO LIVE UNDER ENVIRONMENTAL CHAR
ACTERISTICS THAT SPAN LARGE AREAS MAY HAVE WIDE DISTRIBUTIONS IN SPITE OF HAVING
A NARROW TOLERANCE TO ENVIRONMENTAL CONDITIONS OR OF BEING ABLE TO USE A NARROW
RANGE OF RESOURCES (ANSKI ET AL 'ASTON ET AL 4HIS MECHANISM
IS PARTICULARLY IMPORTANT FOR SPECIES THAT ARE ABLE TO EXPLOIT HUMAN
DOMINATED
ECOSYSTEMS WHICH ARE THEMSELVES VERY WIDESPREAD 3OL ET AL 3ECOND
HIGH DISPERSAL ABILITY CAN ALSO LEAD TO WIDE GEOGRAPHIC RANGES AT LEAST IN SOME
GROUPS WHEREAS LIMITED DISPERSAL CAN RESULT IN GEOGRAPHIC RANGES MUCH NARROWER
4ABLE 3TUDIES EVALUATING THE RELATIONSHIP BETWEEN BREADTH OF GEOGRAPHIC HABITAT RANGE TROPHIC OR BEHAVIORAL BREADTH OF BIRD SPECIES AND
INVASION SUCCESS
-EASURE OF NICHE BREADTH 4AXONOMIC LEVEL OF ANALYSIS )NTRODUCED REGION 2ELATIONSHIP WITH NICHE BREADTHe 2EF
3IZE OF NATIVE RANGE 3PECIES (AWAII 0OSITIVE
!USTRALIA 0OSITIVE
.EW :EALAND .3
&OUR OCEANIC ISLANDS .3
7ORLD 0OSITIVE
o
&AMILIES 7ORLD 0OSITIVE
.O NATIVE HABITAT TYPES 3PECIES 7ORLD 0OSITIVE
o
&AMILIES 7ORLD 0OSITIVE
A
$IET BREADTH 3PECIES .EW :EALAND .3
B
$0 6ZQUEZ
.INE OCEANIC ISLANDS 0OSITIVE
o C
&AMILIES 7ORLD 0OSITIVE
"EHAVIORAL mEXIBILITY 3PECIES 7ORLD 0OSITIVE
2EFERENCES -OULTON AND 0IMM $UNCAN ET AL 6ELTMAN ET AL ,OCKWOOD ET AL "LACKBURN AND
$UNCAN #ASSEY "ROOKS -C,AIN ET AL 3OL AND ,EFEBVRE 3OL ET AL
-EASURES OF DIET BREADTH USED IN STUDIES A$IET BREADTH MEASURED AS CARNIVORE OR HERBIVORE SPECIALIST AND OMNIVORE GENERALIST B$IET BREADTH
MEASURED AS FRUGIVORE OR GRANIVORE SPECIALIST AND OMNIVORE EATING FRUITS AND SEEDS GENERALIST A FOURTH CATEGORY WAS INCLUDED FOR SPECIES
THAT FED ON NEITHER FRUITS OR NECTAR C$IET BREADTH MEASURED AS NUMBER OF SEVEN MAJOR FOOD TYPES CONSUMED
e
)NDICATES STATISTICAL SIGNIlCANCE OF TEST AND DIRECTION OF EFFECT 0OSITIVE SIGNIlCANT EFFECT IN THE DIRECTION EXPECTED BY THE SPECIALIZATIONnNICHE
BREADTH HYPOTHESIS .3 STATISTICALLY NONSIGNIlCANT RESULT
o
#ASSEY ALSO DID ANALYSES AT SPECIES LEVEL (OWEVER HIS SPECIES LEVEL MULTIVARIATE ANALYSES ARE MOSTLY DESCRIPTIVE AND ARE DIFlCULT TO
COMPARE WITH OTHER STUDIES LISTED IN THE TABLE AND WERE THEREFORE NOT INCLUDED
. I CHE BREADTH AND INVASION SUCCESS
THAN WOULD BE PREDICTED BASED ON A SPECIES NICHE 'ASTON 4HUS SPECIES
WITH HIGH DISPERSAL ABILITIES MAY TEND TO HAVE BOTH WIDE NATIVE RANGES AND HIGH
CHANCES OF COLONIZING AND SPREADING IN NEW ENVIRONMENTS 4HIRD SPECIES WITH
WIDER NATIVE RANGES MAY BE MORE LIKELY TO BE TRANSPORTED TO NEW AREAS BY HUMANS
THROUGH PASSIVE SAMPLING 0RINZING ET AL PARTICULARLY CONSIDERING THAT
WIDESPREAD SPECIES ALSO TEND TO BE LOCALLY ABUNDANT 'ASTON ET AL #ASSEY
ET AL HAVE RECENTLY SHOWN THAT PROPAGULE PRESSURE OF INTRODUCED BIRDS
EXPLAINS INVASION SUCCESS WORLDWIDE THESE AUTHORS ALSO FOUND THAT PROPAGULE
PRESSURE IS POSITIVELY RELATED TO GEOGRAPHIC AND TROPHIC BREADTH AND SO THESE
MEASURES OF NICHE BREADTH ARE NOT INDEPENDENT OF PROPAGULE PRESSURE AND COULD
BE CONFOUNDED WITH IT 5NDERSTANDING THE ROLE OF THE HABITAT AND CLIMATIC COMPO
NENTS OF NICHE BREADTH IN DETERMINING INVASION SUCCESS WILL REQUIRE TEASING APART
THESE FACTORS
4(% 2/,% /& 30%#)!,):!4)/. ). -545!,)34)# ).4%2!#4)/.3
0ATTERNS
3EVERAL STUDIES HAVE EVALUATED THE RELATIONSHIP BETWEEN SPECIALIZATION IN MUTU
ALISTIC INTERACTIONS AND INVASION SUCCESS OF PLANTS ANDOR ANIMAL MUTUALISTS
2ICHARDSON ET AL HAVE REVIEWED THE ROLE OF MUTUALISMS IN FACILITATING PLANT
INVASIONS 4HEIR REVIEW SUGGESTS THAT PLANTS WITH EXTREMELY SPECIALIZED MUTUAL
ISTIC INTERACTIONS INCLUDING INTERACTIONS WITH POLLINATORS SEED DISPERSERS MYCOR
RHIZAL FUNGI AND NITROGEN
lXING BACTERIA MAY BE PREVENTED FROM ESTABLISHING AND
SPREADING IN A NEW ENVIRONMENT (OWEVER THE RARITY OF EXTREME SPECIALIZATION IN
MUTUALISTIC INTERACTIONS MAY GREATLY MINIMIZE THE POTENTIAL ROLE OF MUTUALISMS IN
LIMITING PLANT INVASION SUCCESS
/NE OF THE EXAMPLES OF HOW EXTREMELY SPECIALIZED MUTUALISTIC INTERACTIONS MAY
PREVENT THE ESTABLISHMENT AND SPREAD OF INTRODUCED PLANT POPULATIONS DISCUSSED
BY 2ICHARDSON ET AL IS THE INVASION ECOLOGY OF lGS &ICUS -ORACEAE
/F THE SIXTY SPECIES OF &ICUS INTRODUCED TO &LORIDA 53! ONLY THREE HAVE BECOME
INVASIVE AND THEY HAVE DONE SO ONLY AFTER THE ACCIDENTAL INTRODUCTION OF THEIR
SPECIlC WASP SEE 2ICHARDSON ET AL AND REFERENCES THEREIN (OWEVER
2ICHARDSON ET AL ALSO DISCUSS THE CASE OF & LUTEA IN 3OUTH !FRICA WHERE ONE ISO
LATED FEMALE INDIVIDUAL OF THIS NONINDIGENOUS SPECIES HAS BEEN POLLINATED BY WASPS
THAT WERE NOT ITS NORMAL POLLINATOR !LTHOUGH IN THIS CASE THE ABSENCE OF MALE
INDIVIDUALS PREVENTED REPRODUCTION HYBRIDS WITH NATIVE &ICUS DID NOT PRODUCE
VIABLE SEEDLINGS SPREAD MIGHT HAVE BEEN POSSIBLE IF MALE INDIVIDUALS OF & LUTEA
HAD ALSO BEEN INTRODUCED !S A SIDE NOTE 2ICHARDSON ET AL POINT OUT THAT
SEVERAL PINE SPECIES 0INUS SPP NATIVE TO .ORTH !MERICA AND THE -EDITERRANEAN
"ASIN THAT ARE WIND
DISPERSED IN THEIR NATIVE RANGES ARE DISPERSED BY COCKATOOS
IN !USTRALIA !LTHOUGH BIRDS DESTROY MOST OF THE SEEDS SOME SURVIVE TO ESTABLISH
ISOLATED FOCI IN HABITATS EUCALYPT FORESTS THAT THEY WOULD OTHERWISE NOT HAVE
$0 6ZQUEZ
BEEN ABLE TO INVADE 4HUS EVEN APPARENT EXTREME SPECIALIZATION ON A PARTICULAR
POLLEN VECTOR FIG WASP OR SEED DISPERSAL AGENT WIND MAY NOT PRECLUDE SURVIVAL
AND SUCCESSFUL REPRODUCTION IN A NEW ENVIRONMENT
!LBEIT INTRIGUING THE EXTREME SPECIALIZATION OBSERVED FOR &ICUS IS NOT REPRESEN
TATIVE OF THE MAJORITY OF POLLINATION INTERACTIONS 7ASER ET AL AND MOST
SPECIES WITHIN A COMMUNITY FALL SOMEWHERE ALONG A GRADIENT THAT SPANS FROM
EXTREME SPECIALIZATION TO EXTREME GENERALIZATION "ASCOMPTE ET AL *ORDANO
ET AL 6ZQUEZ AND !IZEN A ! SIMILAR SITUATION OCCURS IN SEED
DISPERSAL MUTUALISMS "ASCOMPTE ET AL 4HE QUESTION IS WHETHER SPECIES
LOCATED TOWARDS THE SPECIALIZED EXTREME OF THE CONTINUUM ARE LESS LIKELY TO INVADE
SUCCESSFULLY THAN MORE GENERALIZED SPECIES
)N A RECENT REVIEW OF INTRODUCED BEES AND THEIR ECOLOGICAL IMPACTS 'OULSON
POINTS OUT THAT MOST INTRODUCED BEE SPECIES APPEAR QUITE GENERALIZED
'OULSON ARGUES THAT AMONG BEES WHICH DEPEND ON mORAL FOOD RESOURCES THROUGH
OUT THEIR ENTIRE LIFE CYCLES DIETARY GENERALIZATION SEEMS TO BE A PRECONDITION FOR
BECOMING SUCCESSFUL INVADERS (OWEVER THIS INTERPRETATION SHOULD BE TAKEN WITH
CAUTION !LL SPECIES OF BEES KNOWN TO HAVE BEEN INTRODUCED OUTSIDE THEIR NATIVE
RANGES THE HONEY BEE lVE BUMBLEBEES TEN MEGACHILLIDS AND ONE HALICTID HAVE
BEEN PURPOSEFULLY INTRODUCED BY HUMANS TO ENHANCE CROP POLLINATION USUALLY OF
SEVERAL SPECIES 'IVEN THIS GOAL DIETARY GENERALIZATION OF INTRODUCED BEES IS PROB
ABLY A DESIRABLE TRAIT AND IT IS THEREFORE DIFlCULT TO TEASE APART THE EFFECT OF HUMAN
SELECTION FROM THE EFFECT OF DIETARY SPECIALIZATION ON INVASION SUCCESS
!NOTHER WAY OF EVALUATING THE NICHE BREADTHnINVASION SUCCESS HYPOTHESIS AS
IT APPLIES TO MUTUALISTIC INTERACTIONS IS BY COMPARING THE DEGREE OF SPECIALIZATION
IN MUTUALISTIC INTERACTIONS BETWEEN NATIVE AND INTRODUCED SPECIES IN A GIVEN AREA
4HE EXPECTATION UNDER THE NICHE BREADTHnINVASION SUCCESS HYPOTHESIS IS THAT
INTERACTIONS ARE LESS SPECIlC FOR INTRODUCED THAN FOR NATIVE SPECIES BECAUSE EXOTIC
SPECIES WITH MORE SPECIlC INTERACTIONS MAY HAVE BEEN hlLTERED OUTv DURING THE
ESTABLISHMENT PROCESS ) HAVE FOUND STUDIES IN FOUR SYSTEMS ATTEMPTING TO ANSWER
THIS QUESTION BY COMPARING THE DEGREE OF SPECIALIZATION OF PLANT ANDOR POLLINATORS
BETWEEN NATIVE AND INTRODUCED SPECIES IE A .3 VS )3 COMPARISON SEE ABOVE
)N ADDITION ) HAVE FOUND COMPARABLE DATA FOR A lFTH SYSTEM FOR WHICH ) CALCULATED
DEGREE OF SPECIALIZATION IN A WAY COMPARABLE TO THE OTHER STUDIES 4HESE STUD
IES ARE SUMMARIZED IN 4ABLE .ONE OF THE STUDIES SUPPORT THE NICHE BREADTHn
INVASION SUCCESS HYPOTHESIS AND IN SOME CASES THE OBSERVED PATTERN WAS CONTRARY
TO THE EXPECTATION IE INTRODUCED SPECIES HAD MORE SPECIlC INTERACTIONS 4HUS
THE FEW AVAILABLE STUDIES TELL US THAT INTERACTION SPECIlCITY OF INTRODUCED PLANTS
AND POLLINATORS DOES NOT INmUENCE THEIR INVASION SUCCESS
-ECHANISMS
&ROM THE STUDIES REVIEWED ABOVE EVALUATING THE ROLE OF MUTUALISMS IN INVASIONS
IT APPEARS THAT ONLY EXTREMELY SPECIALIZED MUTUALISMS HAVE THE POTENTIAL TO INmU
ENCE INVASION SUCCESS AND IN GENERAL SPECIlCITY OF MUTUALISTIC INTERACTIONS DOES
4ABLE 3TUDIES THAT HAVE COMPARED DEGREE OF SPECIALIZATION OF PLANTS OR POLLINATORS BETWEEN NATIVE AND INTRODUCED SPECIES
-EAN NUMBER OF MUTUALIST SPECIES
)NTRODUCED REGION -AJOR TAXON .ATIVE %XOTIC 2ESULT OF STATISTICAL TESTe 2EF
)LLINOIS 53! !NGIOSPERMS .EGATIVE
.AHUEL (UAPI !RGENTINA !NGIOSPERMS .3
'ALPAGOS )SLANDS !NGIOSPERMS .EGATIVE
)LE AUX !IGRETTES -AURITIUS !NGIOSPERMS .3
&LORES )SLAND !ZORES !ZORES !NGIOSPERMS .3
.AHUEL (UAPI !RGENTINA (YMENOPTERA RARE SPECIES EXCLUDED .3
.AHUEL (UAPI !RGENTINA (YMENOPTERA !PIDAE ONLY .3
&LORES )SLAND !ZORES !ZORES 0OLLINATORS .3
. I CHE BREADTH AND INVASION SUCCESS
)LE AUX !IGRETTES -AURITIUS 0OLLINATORS .3
2EFERENCES -EMMOTT AND 7ASER -ORALES AND !IZEN -C-ULLEN /LESEN ET AL
e
)NDICATES STATISTICAL SIGNIlCANCE OF TEST AND DIRECTION OF EFFECT .EGATIVE SIGNIlCANT EFFECT IN THE DIRECTION OPPOSITE TO THE EXPECTED BY
THE SPECIALIZATIONnNICHE BREADTH HYPOTHESIS .3 STATISTICALLY NONSIGNIlCANT RESULT
$0 6ZQUEZ
NOT SEEM TO INmUENCE INVASION SUCCESS SIGNIlCANTLY 4HESE RESULTS CONTRAST WITH
THOSE OBTAINED IN STUDIES EVALUATING THE RELATIONSHIP BETWEEN GEOGRAPHIC HABITAT
OR CLIMATIC RANGE AND INVASION SUCCESS REVIEWED IN THE PREVIOUS SECTION (OW CAN
WE EXPLAIN THESE CONTRASTING RESULTS 7HY DOES NARROW CLIMATIC OR GEOGRAPHIC
RANGE APPEAR TO TRANSLATE INTO LOW LIKELIHOOD OF INVASION WHEREAS NARROW RANGE
OF MUTUALISTIC INTERACTIONS DOES NOT 4HE ANSWER TO THIS QUESTION MAY REQUIRE
SOME CONCEPTUAL RElNEMENTS IN OUR THINKING ABOUT THE RELATIONSHIP BETWEEN
NICHE BREADTH AND INVASION SUCCESS AS IT APPLIES TO MUTUALISM
&IRST WE NEED TO DISTINGUISH BETWEEN THE FUNDAMENTAL AND THE REALIZED NICHE
(UTCHINSON )N THE CONTEXT OF SPECIES INTERACTIONS FUNDAMENTAL SPECIAL
IZATION REFERS TO THE POTENTIAL INTERACTIONS THAT WOULD LEAD TO POSITIVE lTNESS FOR
A GIVEN SPECIES UNDER ANY POSSIBLE ECOLOGICAL CIRCUMSTANCES VAN +LINKEN AND
%DWARDS 6ZQUEZ AND !IZEN B 6ZQUEZ 4HUS FUNDAMENTAL
SPECIALIZATION WILL ULTIMATELY DEPEND ON THE GENETIC BACKGROUND OF A SPECIES
WHEREAS REALIZED SPECIALIZATION REFERS TO THE ACTUAL SPECIALIZATION ATTAINED UNDER
A PARTICULAR ECOLOGICAL CONTEXT &OR EXAMPLE MOST FIG SPECIES ARE FUNDAMENTALLY
SPECIALIZED ON ONE OR A FEW SPECIES OF POLLINATING WASPS THEY FAIL TO REPRODUCE IF
THEY ARE MOVED TO DIFFERENT ENVIRONMENTS UNLESS THEIR SPECIALIZED POLLINATORS ARE
INTRODUCED AS WELL #ONVERSELY MANY OTHER PLANT SPECIES THAT ARE POLLINATED BY A
SINGLE POLLINATOR IN A PARTICULAR HABITAT ARE OFTEN POLLINATED BY A DIFFERENT SPECIES
IN OTHER HABITATS WHERE THEY ARE INTRODUCED 2ICHARDSON ET AL 2EGRETTABLY
THE BEST WE CAN DO IN THE lELD IS TO MEASURE THE REALIZED NICHE BUT IF WE WANT TO
PREDICT INVASIONS WHAT WE NEED TO KNOW IS THE FUNDAMENTAL NICHE )N OTHER WORDS
KNOWING THE NICHE BREADTH OF A SPECIES IN A PARTICULAR ENVIRONMENT DOES NOT TELL
US WHAT THE REALIZED NICHE WILL BE IN ANOTHER ENVIRONMENT !LTHOUGH STUDYING THE
CLIMATIC COMPONENT OF THE FUNDAMENTAL NICHE MAY BE POSSIBLE BY FOR EXAMPLE
EXPERIMENTALLY GROWING A SPECIES UNDER A WIDE RANGE OF ENVIRONMENTAL CONDI
TIONS DOING SO IN THE CONTEXT OF SPECIES INTERACTIONS MAY BE SIMPLY UNFEASIBLE
BECAUSE IT WOULD REQUIRE EXPOSING THE SPECIES TO ALL POSSIBLE COMBINATIONS OF INTER
ACTION PARTNERS THAT IT COULD ENCOUNTER IN ANY GIVEN ENVIRONMENT %VEN MANY
SPECIES SELECTED AS BIOCONTROL AGENTS WHICH ARE USUALLY SELECTED TO BE SPECIALISTS
EXHIBIT UNEXPECTED HOST SHIFTS IN THEIR INTRODUCED RANGES 3IMBERLOFF AND 3TILING
(ENNEMAN AND -EMMOTT VAN +LINKEN AND %DWARDS 0EARSON
AND #ALLAWAY AND THESE SHIFTS ARE USUALLY NOT THE RESULT OF CHANGES IN THE
FUNDAMENTAL NICHE THROUGH RAPID EVOLUTION BUT AN EXPRESSION OF A DIFFERENT REAL
IZED NICHE UNDER THE NEW ENVIRONMENT VAN +LINKEN AND %DWARDS
3ECOND AS ) POINTED OUT ABOVE SPECIES THAT USE WIDESPREAD RESOURCES LIVE UNDER
WIDESPREAD ENVIRONMENTAL CONDITIONS OR USE WIDESPREAD HABITATS ARE LIKELY TO BE
WIDESPREAD THEMSELVES 4HIS IDEA CAN BE EASILY EXTENDED TO SPECIES INTERACTIONS
TO ARGUE THAT SPECIES THAT SPECIALIZE ON WIDESPREAD MUTUALISTS MAY THEMSELVES BE
WIDESPREAD !N EMERGING PATTERN IN PLANT
ANIMAL MUTUALISTIC INTERACTIONS IS THE
EXISTENCE OF HIGHLY ASYMMETRIC SPECIALIZATION WHEREBY SPECIALISTS TEND TO SPECIAL
IZE ON GENERALISTS "ASCOMPTE ET AL 6ZQUEZ AND !IZEN A 'IVEN THAT
ABUNDANT FREQUENTLY INTERACTING SPECIES TEND TO BE MORE GENERALIZED THAN RARE
. I CHE BREADTH AND INVASION SUCCESS
SPECIES $UPONT ET AL 6ZQUEZ AND !IZEN A AND THAT LOCALLY
ABUNDANT SPECIES TEND TO BE GEOGRAPHICALLY WIDESPREAD 'ASTON ET AL
SPECIES SPECIALIZED ON WIDESPREAD MUTUALISTS WILL HAVE A GREATER CHANCE OF lNDING
THEIR MUTUALISTS IN THEIR INTRODUCED REGION 4HESE GENERAL FEATURES OF MUTUALISTIC
INTERACTIONS MAY SUBSTANTIALLY INCREASE THE LIKELIHOOD OF SUCCESSFUL INVASION BY
SPECIES THAT ARE RELATIVELY SPECIALIZED ON THEIR MUTUALISTS
4(% 2/,% /& 42/0()# "2%!$4(
0ATTERNS
!NOTHER ASPECT OF THE NICHE THAT CAN POTENTIALLY AFFECT INVASION SUCCESS IS DIETARY
BREADTH AND IT IS FREQUENTLY LISTED AS ONE POTENTIAL CHARACTERISTIC OF SUCCESS
FUL INVADERS %HRLICH 2ICCIARDI AND 2ASMUSSEN -C+INNEY AND
,OCKWOOD ) DISCUSS HERE A FEW STUDIES THAT HAVE ADDRESSED THIS ISSUE IN THE
CONTEXT OF BIRD INVASIONS
4O MY KNOWLEDGE THREE STUDIES HAVE EVALUATED THE RELATIONSHIP BETWEEN INVA
SION SUCCESS AND DIET BREADTH 4ABLE 4HESE STUDIES COMPARED TROPHIC BREADTH
IN THE NATIVE REGION OF SUCCESSFUL VS FAILED INVADERS 3) VS &) -C,AIN ET AL
DElNED SPECIALISTS AS SPECIES FEEDING ON EITHER FRUITS OR SEEDS AND GENERALISTS AS
SPECIES FEEDING ON BOTH THEY FOUND THAT TROPHIC GENERALISTS WERE MORE LIKELY TO
BE SUCCESSFUL INVADERS OF OCEANIC ISLANDS THAN TROPHIC SPECIALISTS 6ELTMAN ET AL
DElNED HERBIVOROUS OR CARNIVOROUS SPECIES AS DIETARY SPECIALISTS AND
OMNIVORES SPECIES FEEDING ON BOTH PLANT AND ANIMAL MATERIAL AS GENERALISTS
THEY FAILED TO lND A STATISTICAL EFFECT OF SIZE OF NATIVE TROPHIC RANGE ON THE SUCCESS OF
BIRDS INTRODUCED IN .EW :EALAND &INALLY #ASSEY DElNED DIETARY BREADTH
AS THE NUMBER OF FOOD TYPES CONSUMED OUT OF SEVEN POSSIBLE CATEGORIES (E FOUND
A WEAK BUT SIGNIlCANT EFFECT OF DIETARY BREADTH EXPLAINING THE INVASION SUCCESS OF
BIRD FAMILIES
-ECHANISMS
&ROM THIS LIMITED NUMBER STUDIES AND FROM THEIR DISPARATE RESULTS IT IS DIFlCULT TO
MAKE GENERALIZATIONS ABOUT THE IMPORTANCE OF TROPHIC BREADTH FOR INVASION SUC
CESS &URTHERMORE A MAJOR LIMITATION OF THE STUDIES DISCUSSED ABOVE IS THAT THE
MEASURE OF SPECIALIZATION USED IS PROBABLY TOO COARSE AND MAY THEREFORE BE A POOR
SURROGATE OF TRUE TROPHIC SPECIALIZATION &OR EXAMPLE IN THE STUDY BY -C,AIN ET AL
ONLY FRUITS AND SEEDS ARE CONSIDERED TO CHARACTERIZE DIET BREADTH ALTHOUGH
THE AUTHORS STATE THAT hVIRTUALLY ALL INTRODUCED SPECIES ;IN THESE ISLANDS= EAT SOME
INSECTS AND OFTEN OTHER INVERTEBRATESv P 4HUS IT IS UNCLEAR WHETHER WHAT
THE AUTHORS MEASURED ACTUALLY REPRESENTS DIET BREADTH OR RATHER WHETHER IT REPRE
SENTS SOMETHING ELSE RELATED TO EATING FRUITS OR SEEDS )N FACT THE QUANTIlCATION OF
DEGREE OF SPECIALIZATION IS PROBABLY O NE OF THE MAIN
CONCEPTUAL
AND METHODOLOGICAL
$0 6ZQUEZ
HURDLES IN THE STUDY OF SPECIES INTERACTIONS -ARTINEZ ET AL .OVOTN ET AL
6ZQUEZ AND !IZEN B -ANY IDEAS IN ECOLOGY EVOLUTION AND BIOGEOG
RAPHY DEPEND ON ASSUMPTIONS ABOUT THE DEGREE OF SPECIALIZATION OF SPECIES INTER
ACTIONS &OR EXAMPLE BECAUSE THE BULK OF THE %ARTHS SPECIES ARE BELIEVED TO BE
TROPICAL HERBIVOROUS INSECTS ESTIMATES OF THE NUMBER OF SPECIES ON %ARTH DEPEND
CRITICALLY ON ASSUMPTIONS ABOUT THEIR DEGREE OF SPECIALIZATION 4HE REALIZATION THAT
TROPICAL HERBIVOROUS INSECTS WERE LESS SPECIALIZED THAN PREVIOUSLY THOUGHT LED TO
REVISING THE ESTIMATED NUMBER OF GLOBAL SPECIES FROM MILLION %RWIN TO
APPROXIMATELY MILLION DEGAARD .OVOTN ET AL
4O CIRCUMVENT THE PROBLEM OF THE MEASUREMENT OF TROPHIC SPECIALIZATION IN
BIRDS 3OL AND ,EFEBVRE AND 3OL ET AL PROPOSED THE USE OF hBEHAVIOR
AL mEXIBILITYv AS A SURROGATE OF TROPHIC BREADTH 4HESE AUTHORS DElNED BEHAVIORAL
mEXIBILITY AS hTHE FREQUENCY OF NEW AND UNUSUAL FEEDING BEHAVIORS REPORTED IN THE
SHORT NOTE SECTION OF ORNITHOLOGY JOURNALSv 3OL ET AL P 4HEY ARGUE
THAT A TAXONOMIC GROUP IN WHICH LARGE NUMBERS OF NEW FEEDING BEHAVIORS ARE
OBSERVED IS LIKELY TO CHANGE ITS FORAGING TECHNIQUES OR DIET FREQUENTLY TO EAT A SUR
PRISINGLY LARGE RANGE OF FOODS AND TO USE HANDLING BEHAVIORS AND NOVEL SITUATIONS
IN A WAY THAT STRIKES ORNITHOLOGISTS BY ITS COMPLEXITY AND mEXIBILITY 4HEY FURTHER
FOUND THAT BEHAVIORAL mEXIBILITY OF BIRD SPECIES IN THEIR NATIVE RANGE IS RELATED TO
THEIR SUCCESS AS INVADERS WORLDWIDE 4HIS APPROACH IS A PROMISING ALTERNATIVE TO
ESTIMATING TROPHIC NICHE BREADTH FROM FEEDING RECORDS
2EGARDLESS OF THE PROBLEMS OF ESTIMATION OF TROPHIC SPECIALIZATION TWO ISSUES
DISCUSSED ABOVE IN THE CONTEXT OF GEOGRAPHIC RANGE AND MUTUALISMS ALSO APPLY
HERE &IRST SPECIES SPECIALIZING ON WIDESPREAD RESOURCES PARTICULARLY THOSE THAT
EXPLOIT HUMAN
DOMINATED ECOSYSTEMS MAY BE LIKELY TO INVADE IN SPITE OF BEING
SPECIALIZED 3ECOND EVEN IF ACCURATE ESTIMATES USING FEEDING RECORDS PROVIDE
A MEASURE OF THE REALIZED NICHE BUT NOT OF THE FUNDAMENTAL NICHE WHICH WOULD
BE NECESSARY PREDICT INVASION SUCCESS IN A NEW ENVIRONMENT WITH A DIFFERENT SET
OF FOOD RESOURCES
#/.#,5$).' 2%-!2+3
-Y REVIEW OF THE LITERATURE SUGGESTS SEVERAL GENERALIZATIONS ABOUT THE ROLE OF
NICHE BREADTH AS A DETERMINANT OF INVASION SUCCESS )N PARTICULAR NATIVE HABITAT
GEOGRAPHIC OR CLIMATIC RANGE OF PLANTS AND BIRDS APPEAR TO BE GOOD PREDICTORS
INVASION SUCCESS WHEREAS SPECIlCITY OF MUTUALISTIC INTERACTIONS AND TROPHIC
BREADTH APPEAR TO HAVE A MUCH WEAKER PREDICTIVE POWER (OWEVER EVEN WHEN
RESULTS DO lT THE PREDICTIONS OF THE NICHE BREADTHnINVASION SUCCESS HYPOTHESIS IT
IS USUALLY NOT POSSIBLE TO KNOW WHETHER NICHE BREADTH IS INVOLVED IN GENERATING
THE OBSERVED PATTERN &UTURE STUDIES AIMING AT PREDICTING THE SUCCESS OF INVAD
ERS SHOULD TEASE APART THE RELATIVE CONTRIBUTIONS OF NICHE BREADTH FROM OTHER
CONFOUNDING FACTORS -ORE BROADLY ) HOPE MY REVIEW CAN SERVE AS AN EXAMPLE OF
HOW CONCEPTUAL ELABORATION MAY BE USEFUL TO IMPROVE OUR UNDERSTANDING IN
INVASIONS BIOLOGY
. I CHE BREADTH AND INVASION SUCCESS
!#+./7,%$'-%.43
) THANK THE EDITORS FOR THE INVITATION TO CONTRIBUTE TO THIS VOLUME 3EAN -C-AHON
)NGRID 0ARKER $AVID 2ICHARDSON %RIC 3EABLOOM $ANIEL 3OL AND -ONTSERRAT 6IL
MADE USEFUL SUGGESTIONS ON THE MANUSCRIPT ) WAS SUPPORTED BY A 0OSTDOCTORAL
&ELLOWSHIP AT THE .ATIONAL #ENTER FOR %COLOGICAL !NALYSIS AND 3YNTHESIS
FUNDED BY .3& 'RANT $%"
THE 5NIVERSITY OF #ALIFORNIA AND THE 3ANTA
"ARBARA CAMPUS
2%&%2%.#%3
!RNQVIST ' AND $ 7OOSTER -ETA ANALYSIS SYNTHESIZING RESEARCH lNDINGS IN ECOL
OGY AND EVOLUTION 4RENDS IN %COLOGY AND %VOLUTION
"AKER ( ' #HARACTERISTICS AND MODES OF ORIGIN OF WEEDS 0AGES IN
( ' "AKER AND # , 3TEBBINS EDITORS 4HE 'ENETICS OF #OLONIZING 3PECIES !CADEMIC
0RESS .EW 9ORK
"AKER ( ' 4HE EVOLUTION OF WEEDS !NNUAL 2EVIEW OF %COLOGY AND 3YSTEMATICS
"ASCOMPTE * 0 *ORDANO # * -ELIN AND * - /LESEN 4HE NESTED ASSEMBLY OF
PLANT ANIMAL MUTUALISTIC NETWORKS 0ROCEEDINGS OF THE .ATIONAL !CADEMY OF 3CIENCES
53!
"LACKBURN 4 - AND 2 0 $UNCAN $ETERMINANTS OF ESTABLISHMENT SUCCESS IN INTRO
DUCED BIRDS .ATURE
"ROOKS 4 - !RE UNSUCCESSFUL AVIAN INVADERS RARER IN THEIR NATIVE RANGE THAN
SUCCESSFUL INVADERS 0AGES IN * , ,OCKWOOD AND - - -C+INNEY EDITORS
"IOTIC (OMOGENISATION THE LOSS OF DIVERSITY THROUGH INVASION AND EXTINCTION +LUWER
!CADEMIC 0RESS $ORDRECHT 4HE .ETHERLANDS
"ROWN * ( /N THE RELATIONSHIP BETWEEN ABUNDANCE AND DISTRIBUTION OF SPECIES
!MERICAN .ATURALIST
#ASSEY 0 ,IFE HISTORY AND ECOLOGY INmUENCES ESTABLISHMENT SUCCESS OF INTRODUCED
LAND BIRDS "IOLOGICAL *OURNAL OF THE ,INNEAN 3OCIETY
$AEHLER # # AND $ 2 3TRONG 0REDICTION AND BIOLOGICAL INVASIONS 4RENDS IN
%COLOGY AND %VOLUTION
$UNCAN 2 0 - "OMFORD $ - &ORSYTH AND , #ONIBEAR (IGH PREDICTABILITY IN
INTRODUCTION OUTCOMES AND THE GEOGRAPHICAL RANGE SIZE OF INTRODUCED !USTRALIAN BIRDS
A ROLE FOR CLIMATE *OURNAL OF !NIMAL %COLOGY
$UPONT 9 , $ - (ANSEN AND * - /LESEN 3TRUCTURE OF A PLANT mOWER VISITOR
NETWORK IN THE HIGH ALTITUDE SUB ALPINE DESERT OF 4ENERIFE #ANARY )SLANDS %COGRAPHY
%HRLICH 0 2 7HICH ANIMAL WILL INVADE 0AGES IN ( ! -OONEY AND
* ! $RAKE EDITORS %COLOGY OF "IOLOGICAL )NVASIONS OF .ORTH !MERICA AND (AWAII
3PRINGER 6ERLAG .EW 9ORK
$0 6ZQUEZ
%RWIN 4 , 4ROPICAL FORESTS THEIR RICHNESS IN COLEOPTERA AND OTHER ARTHROPOD SPE
CIES #OLEOPTERISTS "ULLETIN
'ASTON + * 4HE 3TRUCTURE AND $YNAMICS OF 'EOGRAPHIC 2ANGES /XFORD 5NIVERSITY
0RESS /XFORD AND .EW 9ORK
'ASTON + * 4 - "LACKBURN AND * ( ,AWTON )NTERSPECIlC ABUNDANCEnRANGE SIZE
RELATIONSHIPS AN APPRAISAL OF MECHANISMS *OURNAL OF !NIMAL %COLOGY
'OODWIN " * ! * -C!LLISTER AND , &AHRIG 0REDICTING INVASIVENESS OF PLANT SPECIES
BASED ON BIOLOGICAL INFORMATION #ONSERVATION "IOLOGY
'OULSON $ %FFECTS OF INTRODUCED BEES ON NATIVE ECOSYSTEMS !NNUAL 2EVIEW OF
%COLOGY %VOLUTION AND 3YSTEMATICS
'UREVITCH * 0 3 #URTIS AND - ( *ONES -ETA
ANALYSIS IN ECOLOGY !DVANCES IN
%COLOGICAL 2ESEARCH
(ANSKI ) * +OUKI AND ! (ALKKA 4HREE EXPLANATIONS OF THE POSITIVE RELATIONSHIP
BETWEEN DISTRIBUTION AND ABUNDANCE OF SPECIES 0AGES
IN 2 % 2ICKLEFS AND $
3CHLUTER EDITORS 3PECIES DIVERSITY IN ECOLOGICAL COMMUNITIES HISTORICAL AND GEOGRAPHI
CAL PERSPECTIVES 5NIVERSITY OF #HICAGO 0RESS #HICAGO ),
(ENNEMAN - , AND * -EMMOTT )NlLTRATION OF A (AWAIIAN COMMUNITY BY INTRO
DUCED BIOLOGICAL CONTROL AGENTS 3CIENCE
(UTCHINSON ' % #ONCLUDING REMARKS #OLD 3PRING (ARBOR 3YMPOSIA ON 1UANTITATIVE
"IOLOGY
*ORDANO 0 * "ASCOMPTE AND * - /LESEN )NVARIANT PROPERTIES IN COEVOLUTIONARY
NETWORKS OF PLANT
ANIMAL INTERACTIONS %COLOGY ,ETTERS
+ITAYAMA + AND $ -UELLER
$OMBOIS "IOLOGICAL INVASION ON AN OCEANIC ISLAND
MOUNTAIN $O ALIEN PLANT SPECIES HAVE WIDER ECOLOGICAL RANGES THAN NATIVE SPECIES
*OURNAL OF 6EGETATION 3CIENCE
,OCKWOOD * , - 0 -OULTON AND + , "ALENT )NTRODUCED AVIFAUNAS AS NATU
RAL EXPERIMENTS IN COMMUNITY ASSEMBLY 0AGES
IN % 7EIHER AND 0 +EDDY
EDITORS %COLOGICAL !SSEMBLY 2ULES 0ERSPECTIVES !DVANCES 2ETREATS #AMBRIDGE
5NIVERSITY 0RESS #AMBRIDGE
-ARTINEZ . $ " ! (AWKINS ( ! $AWAH AND " 0 &EIFAREK %FFECTS OF SAMPLING
EFFORT ON CHARACTERIZATION OF FOOD
WEB STRUCTURE %COLOGY
-C+INNEY - , AND * , ,OCKWOOD "IOTIC HOMOGENIZATION A FEW WINNERS
REPLACING MANY LOSERS IN THE NEXT MASS EXTINCTION 4RENDS IN %COLOGY AND %VOLUTION
-C,AIN $ + - 0 -OULTON AND * ' 3ANDERSON 3EXUAL SELECTION AND EXTINCTION
4HE FATE OF PLUMAGE
DIMORPHIC AND PLUMAGE
MONOMORPHIC BIRDS INTRODUCED ONTO
ISLANDS %VOLUTIONARY %COLOGY 2ESEARCH
-C-ULLEN # + &LOWER
VISITING INSECTS OF THE 'ALPAGOS )SLANDS 4HE 0AN
0ACIlC
%NTOMOLOGIST
-EMMOTT * AND . - 7ASER )NTEGRATION OF ALIEN PLANTS INTO A NATIVE mOWER
POLLINATOR VISITATION WEB 0ROCEEDINGS OF THE 2OYAL 3OCIETY OF ,ONDON 3ERIES "
. I CHE BREADTH AND INVASION SUCCESS
-ORALES # , AND - ! !IZEN $OES INVASION OF EXOTIC PLANTS PROMOTE INVASION OF
EXOTIC mOWER VISITORS ! CASE STUDY FROM THE TEMPERATE FORESTS OF THE SOUTHERN !NDES
"IOLOGICAL )NVASIONS
-OULTON - 0 AND 3 , 0IMM 3PECIES INTRODUCTIONS TO (AWAII 0AGES
IN ( ! -OONEY AND * ! $RAKE EDITORS %COLOGY OF "IOLOGICAL )NVASIONS OF .ORTH
!MERICA AND (AWAII 3PRINGER
6ERLAG .EW 9ORK
.OVOTN 6 9 "ASSET 3 % -ILLER ' $ 7EIBLEN " "REMER , #IZEK AND 0 $ROZD
,OW HOST SPECIlCITY OF HERBIVOROUS INSECTS IN A TROPICAL FOREST .ATURE
DEGAARD & (OW MANY SPECIES OF ARTHROPODS %RWINS ESTIMATE REVISED "IOLOGICAL
*OURNAL OF THE ,INNEAN 3OCIETY
/LESEN * - , ) %SKILDSEN AND 3 6ENKATASAMY )NVASION OF POLLINATION NETWORKS
ON OCEANIC ISLANDS IMPORTANCE OF INVADER COMPLEXES AND ENDEMIC SUPER GENERALISTS
$IVERSITY AND $ISTRIBUTIONS
0EARSON $ % AND 2 - #ALLAWAY )NDIRECT EFFECTS OF HOST
SPECIlC BIOLOGICAL CONTROL
AGENTS 4RENDS IN %COLOGY %VOLUTION
0RINZING ! 7 $URKA 3 +LOTZ AND 2 "RANDL 7HICH SPECIES BECOME ALIENS
%VOLUTIONARY %COLOGY 2ESEARCH
2EJMNEK - 7HAT MAKES A SPECIES INVASIVE 0AGES
IN 0 0YEK EDITOR 0LANT
INVASIONS GENERAL ASPECTS AND SPECIAL PROBLEMS !CADEMIC 0UBLISHING !MESTERDAM
2EJMNEK - )NVASIVE PLANTS APPROACHES AND PREDICTIONS !USTRAL %COLOGY
2ICCIARDI ! AND * " 2ASMUSSEN 0REDICTING THE IDENTITY AND IMPACT OF FUTURE
BIOLOGICAL INVADERS A PRIORITY FOR AQUATIC RESOURCE MANAGEMENT #ANADIAN *OURNAL OF
&ISHERIES AND !QUATIC 3CIENCES
2ICHARDSON $ - . !LLSOPP # - $!NTONIO 3 * -ILTON AND - 2EJMNEK 0LANT
INVASIONS THE ROLE OF MUTUALISMS "IOLOGICAL 2EVIEWS
2OY * - , .AVAS AND , 3ONI )NVASION BY ANNUAL GRASSES A CASE STUDY
CHALLENGING THE HOMOCLIME APPROACH TO INVASIONS 0AGES
IN 2 ( 'ROVES AND
& DI #ASTRI EDITORS "IOGEOGRAPHY OF -EDITERRANEAN )NVASIONS #AMBRIDGE 5NIVERSITY
0RESS #AMBRIDGE
3COTT * + AND & $ 0ANETTA 0REDICTING THE !USTRALIAN WEED STATUS OF SOUTHERN
!FRICAN PLANTS *OURNAL OF "IOGEOGRAPHY
3IMBERLOFF $ AND 0 $ 3TILING (OW RISKY IN BIOLOGICAL CONTROL %COLOGY
3OL $ AND , ,EFEBVRE "EHAVIOURAL mEXIBILITY PREDICTS INVASION SUCCESS IN BIRDS
INTRODUCED TO .EW :EALAND /IKOS
3OL $ 3 4IMMERMANS AND , ,EFEBVRE "EHAVIOURAL mEXIBILITY AND INVASION SUCCESS
IN BIRDS !NIMAL "EHAVIOUR
VAN +LINKEN 2 $ AND / 2 %DWARDS )S HOST
SPECIlCITY OF WEED BIOLOGICAL CONTROL
AGENTS LIKELY TO EVOLVE RAPIDLY FOLLOWING ESTABLISHMENT %COLOGY ,ETTERS
6ZQUEZ $ 0 AND - ! !IZEN .ULL MODEL ANALYSES OF SPECIALIZATION IN PLANT
POLLINATOR INTERACTIONS %COLOGY
6ZQUEZ $ 0 AND - ! !IZEN A !SYMMETRIC SPECIALIZATION A PERVASIVE FEATURE OF
PLANT
POLLINATOR INTERACTIONS %COLOGY
$0 6ZQUEZ
6ZQUEZ $ 0 AND - ! !IZEN B #OMMUNITY
WIDE PATTERNS OF SPECIALIZATION IN
PLANT
POLLINATOR INTERACTIONS REVEALED BY NULL
MODELS 0AGES IN PRESS IN * /LLERTON
EDITOR 3PECIALIZATION AND GENERALIZATION IN PLANT
POLLINATOR INTERACTIONS 5NIVERSITY OF
#HICAGO 0RESS #HICAGO
6ZQUEZ $ 0 2EVISITANDO EL NICHO HUTCHINSONIANO %COLOGA !USTRAL IN PRESS
6ELTMAN # * 3 .EE AND - * #RAWLEY #ORRELATES OF INTRODUCTION SUCCESS IN
EXOTIC .EW :EALAND BIRDS !MERICAN .ATURALIST
7ASER . - , #HITTKA - 6 0RICE . - 7ILLIAMS AND * /LLERTON 'ENERALIZATION
IN POLLINATION SYSTEMS AND WHY IT MATTERS %COLOGY
#HAPTER FIFTEEN
)NTERACTIONS BETWEEN INVASIVE
PLANTS AND SOIL ECOSYSTEMS
POSITIVE FEEDBACKS AND THEIR
POTENTIAL TO PERSIST
! 3 4HORPE AND 2 - #ALLAWAY
).42/$5#4)/.
0LANTS INDIRECTLY AFFECT THEIR NEIGHBORS IN MANY WAYS BUT ONE OF THE MOST
IMPORTANT IS BY ALTERING THE BIOTIC PHYSICAL AND CHEMICAL CHARACTERISTICS OF SOILS
(OBBIE !NGERS AND #ARON "ERENDSE "INKLEY AND 'IARDINA
.ORTHRUP ET AL 3CHLESINGER AND 0ILMANIS 6AN "REEMEN
7ARDLE ET AL #HEN AND 3TARK %ATON AND &ARRELL 4HESE GEN
ERAL EFFECTS HAVE BEEN UNDERSTOOD FOR DECADES BUT ONLY RECENTLY HAVE EXPERIMENTS
DEMONSTRATED THAT COMPLEX INTERACTIONS BETWEEN PLANTS AND SOIL MICROBIAL COM
MUNITIES CAN HAVE STRONG EFFECTS ON PLANT POPULATIONS "EVER ET AL #LAY AND
6AN DER 0UTTEN 0ACKER AND #LAY INTERACTIONS AMONG PLANT SPECIES
7EST AND THE ORGANIZATION OF PLANT COMMUNITIES 'RIME ET AL 6AN
DER 0UTTEN ET AL "EVER 6AN DER 0UTTEN VAN DER (EIJDEN ET AL
(OOPER ET AL +LIRONOMOS 3OIL COMMUNITIES ALTER COMPETITIVE
OUTCOMES AMONG PLANTS THROUGH THEIR PATHOGENIC EFFECTS 6AN DER 0UTTEN AND
-7 #ADOTTE ET AL EDS #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY n
3PRINGER 0RINTED IN THE .ETHERLANDS
!3 4HORPE AND 2- #ALLAWAY
0ETERS BY FAVORING OBLIGATE MYCORRHIZAL SPECIES OVER NON MYCORRHIZAL OR
FACULTATIVE MYCORRHIZAL SPECIES (ETRICK ET AL (ARTNETT ET AL AND BY
TRANSFERRING RESOURCES AND lXED CARBON BETWEEN SPECIES #HIARELLO ET AL
&RANCIS AND 2EAD 'RIME ET AL -OORA AND :OBEL 7ATKINS ET AL
3IMARD ET AL -ARLER ET AL BUT SEE 2OBINSON AND &ITTER
"Y ALTERING THE BIOTIC AND ABIOTIC CHARACTERISTICS OF SOILS PLANTS CAN DRIVE POSITIVE
OR NEGATIVE FEEDBACKS "OX AND THESE FEEDBACKS CAN PROFOUNDLY AFFECT PLANT
POPULATIONS AND COMMUNITIES
"OX
0OSITIVE FEEDBACKS OCCUR WHEN PLANT SPECIES ACCUMULATE MICROBES THAT
HAVE BENElCIAL EFFECTS ON THE PLANTS THAT CULTIVATE THEM SUCH AS MYCOR
RHIZAL FUNGI AND NITROGEN lXERS 0OSITIVE FEEDBACKS PROMOTE SPECIES
DOMINANCE AND ARE THOUGHT TO LEAD TO A LOSS OF LOCAL COMMUNITY DIVERSITY
"EVER ET AL "EVER .EGATIVE FEEDBACKS OCCUR WHEN PLANT
SPECIES ACCUMULATE PATHOGENIC MICROBES IN THEIR RHIZOSPHERES AND THESE
INTERACTIONS OUTWEIGH THE BENElTS RECEIVED FROM MUTUALISTIC INTERACTIONS
.EGATIVE FEEDBACKS CREATE CONDITIONS THAT ARE INCREASINGLY HOSTILE TO THE
PLANTS THAT CULTIVATE THE PATHOGENS AND ARE THOUGHT TO PROMOTE COMMU
NITY DIVERSITY 6AN DER 0UTTEN ET AL "EVER +LIRONOMOS
*IM "EVER "EVER ET AL PROPOSED THE FEEDBACK MODEL IN WHICH
A PLANT OR POPULATION OF PLANTS ALTERS THE COMPOSITION OF THE SOIL COMMUNITY
4HIS CHANGE THEN FEEDS BACK TO AFFECT THE GROWTH AND SURVIVAL OF THE PLANT INDI
VIDUAL OR POPULATION 4HIS MODEL DESCRIBES HOW TWO DIFFERENT TYPES OF FEEDBACK
COULD BE ESTABLISHED POSITIVE FEEDBACK WHICH SHOULD OCCUR WHEN THE NEGATIVE
EFFECTS OF SOIL PATHOGENS ARE OUTWEIGHED BY THE BENElCIAL EFFECTS OF MUTUALISTIC
MYCORRHIZAE AND NEGATIVE FEEDBACK WHICH IS EXPECTED TO OCCUR EITHER WHEN THE
EFFECTS OF PATHOGENS OUTWEIGH THE EFFECTS OF MYCORRHIZAE OR WHEN THE SOIL COMMU
NITY ENHANCES THE GROWTH OF COMPETING PLANTS MORE THAN THAT OF THE ASSOCIATED
PLANT "EVER "EVER ET AL "Y FAVORING THE LOCAL REPLACEMENT OF SPE
CIES NEGATIVE FEEDBACK IS EXPECTED TO MAINTAIN SPECIES DIVERSITY WHEREAS POSITIVE
FEEDBACK IS EXPECTED TO LEAD TO SPECIES DOMINANCE AND A DECREASE IN LOCAL SPECIES
DIVERSITY "EVER "EVER ET AL 7ATKINSON ! GROWING BODY OF
RESEARCH HAS DEMONSTRATED THAT PLANT
SOIL FEEDBACKS CAN HAVE IMPORTANT ECO
LOGICAL CONSEQUENCES "EVER ET AL 7ARDLE AND .ICHOLSON 7ESTOVER
ET AL +LIRONOMOS &OR EXAMPLE IN A STUDY COMPARING FEEDBACK AND
RELATIVE ABUNDANCE OF CO
EXISTING OLD lELD SPECIES FROM SOUTHERN /NTARIO
#ANADA +LIRONOMOS FOUND A STRONG POSITIVE RELATIONSHIP BETWEEN A
PLANTS FEEDBACK WITH THE SOIL COMMUNITY AND ITS RELATIVE ABUNDANCE &IG
)NVASIVE PLANT
SOIL ECOSYSTEM FEEDBACKS
&IG 4HE RELATIONSHIP BETWEEN RELATIVE PLANT ABUNDANCE IN AN OLD
lELD SITE AND SOIL
FEEDBACK RESPONSE 3QUARES REPRESENT MEANS Y X X
R 0 .UMBERS REPRESENT DIFFERENT PLANT SPECIES #AREX GARBERI #AREX
AUREA #AREX GRANULARIS $AUCUS CAROTA !GROSTIS GIGANTEA 3OLIDAGO GRAMINIFOLIA
3OLIDAGO NEMORALIS !STER SIMPLEX !STER VIMINEUS !STER NOVAEANGLIAE #IRCIUM
VULGARE #HENOPODIUM AMBROSIOIDES /ENOTHERA BIENNIS #AREX mAVA *UNCUS
DUDLEYI 3OLIDAGO CANADENSIS ,INARIA VULGARIS #ICHORIUM INTYBUS #IRCIUM
ARVENSE 3OLIDAGO RUGOSA 'EUM ALEPPICUM 3ATUREJA VULGARIS 0OTENTILLA RECTA
#ORONILLA VARIA !SCLEPIAS SYRIACA !CHILLEA MILLEFOLIUM !POCYNUM CANNABINUM
(YPERICUM PERFORATUM !GROSTIS SCABRA 0HLEUM PRATENSE 0OA COMPRESSA
%CHIUM VULGARE #ENTAUREA JACEA 2UDBECKIA SEROTINA 0OA PRATENSIS
$ACTYLIS GLOMERATA #ERASTIUM VULGATUM 'ALIUM PALUSTRE /ENOTHERA PERENNIS
0RUNELLA VULGARIS 4RIFOLIUM PRATENSE #ONVOLVULUS ARVENSIS 3ILENE CUCUBALUS
%RIGERON STRIGOSUS !SPARAGUS OFlCINALIS (IERACIUM AURANTICUM %RIGERON
PHILADELPHICUS 6ERONICA OFlCINALIS 0LANTAGO LANCEOLATA 'ALIUM MOLLUGO
(IERACIUM PILOSELLA 6ICIA CRACCA (IERACIUM PRATENSE -EDICAGO LUPULINA
2ANUNCULUS ACRIS 4ARAXACUM OFlCINALE &RAGARIA VIRGINIANA #HRYSANTHEMUM
LEUCANTHEMUM 4RAGOPOGON PRATENSIS "ROMUS INERMIS 0ANICUM LANUGINOSUM
2EPRINTED FROM +LIRONOMOS
0LANT SPECIES THAT WERE FOUND IN LOW ABUNDANCE IN THE lELD CONSISTENTLY DISPLAYED
NEGATIVE FEEDBACK INTERACTIONS WHEREAS PLANTS WITH HIGH ABUNDANCE EITHER HAD
LOW NEGATIVE OR POSITIVE FEEDBACK INTERACTIONS +LIRONOMOS
!3 4HORPE AND 2- #ALLAWAY
0LANT INVASIONS PROVIDE AN EXCEPTIONAL OPPORTUNITY FOR UNDERSTANDING HOW
PLANTS AFFECT SOILS AND DRIVE FEEDBACK PROCESSES .UMEROUS STUDIES HAVE DOCU
MENTED THE EFFECTS OF INVASIVE SPECIES ON THE COMPOSITION OF SOIL BIOTA "ELNAP AND
0HILLIPS +OURTEV ET AL AND SOIL MICROBIAL FUNCTION +OURTEV ET AL
%HRENFELD (OWEVER MUCH LESS IS KNOWN ABOUT THE ROLE OF FEEDBACKS
IN THE SUCCESS OF EXOTIC PLANTS &EEDBACKS MAY ALSO AFFECT EVOLUTION POTENTIALLY
MOST APPARENT IN THE RAPID EVOLUTIONARY CHANGES THAT CAN ACCOMPANY EXOTIC
INVASION 2ICE AND %MORY
)F PLANT SOIL FEEDBACKS DRIVE VARIATION IN lTNESS DIFFERENCES AMONG INDIVIDUALS
THEN ULTIMATELY FEEDBACK INTERACTIONS MAY BE UNDER SELECTIVE PRESSURE 6AN DER
0UTTEN 6AN "REEMAN AND &INZI "ASED ON THEORY DEVELOPED BY "EVER
ET AL AND +LIRONOMOS WE HYPOTHESIZE THAT POSITIVE FEEDBACKS IN
GENERAL ARE LIKELY TO LEAD THE PLANT COMMUNITY TO SHIFT TOWARDS A MONOCULTURE OF
THE INVASIVE SPECIES (OWEVER WE PROPOSE THAT THE LONGEVITY OF POSITIVE FEEDBACK
INTERACTIONS IN EVOLUTIONARY TIME MAY DEPEND ON WHETHER THE INVASIVE PLANT IS
HAVING ITS STRONGEST EFFECTS ON THE PATHOGENIC OR THE NUTRIENT CYCLING COMPONENTS
OF THE SOIL ECOSYSTEM &IG -ICROBES HAVE A SHORT GENERATION TIME AND THUS
CAN RESPOND TO EVOLUTIONARY PRESSURES QUICKLY 4HEREFORE PATHOGENS MAY RESPOND
RELATIVELY RAPIDLY TO THE INVASION OF NON NATIVE PLANTS 4HIS MAY ULTIMATELY LEAD
TO GREATER COEXISTENCE AMONG SPECIES WHERE ALTHOUGH THE INVASIVE MAY STILL
BE PRESENT IT WILL NOT BE THE OVER WHELMING COMMUNITY DOMINANT )N CONTRAST
FEEDBACKS BETWEEN INVASIVE PLANTS AND NUTRIENT CYCLES MAY BE MUCH MORE LIKELY
TO PERSIST 7HEN THE BIOTIC COMPONENT OF THE SOIL NUTRIENT CYCLES IS INVOLVED IT
IS EXPECTED THAT THERE WOULD BE A SHIFT IN THE MICROBIAL COMMUNITY TO POPULA
TIONS THAT ARE BETTER ADAPTED TO THE NEW NUTRIENT STATUS OF THE ECOSYSTEM &IG
!TLAS AND "ARTHA 3CHIMEL AND "ENNETT )N THE CASE OF ALTERATIONS TO
ABIOTIC COMPONENTS OF THE SOIL ECOSYSTEM THERE IS NO DIRECT SELECTIVE PRESSURE TO
STIMULATE ADAPTATION &IG
(ERE WE DISCUSS INTERACTIONS BETWEEN INVASIVE SPECIES AND THE SOIL ECOSYSTEM
PATHOGENIC AND NUTRIENT CYCLING COMPONENTS THE MECHANISMS FOR THESE INTER
ACTIONS AND EVIDENCE THAT THESE INTERACTIONS HAVE VERY DIFFERENT EFFECTS ON THE
SURVIVAL OF INVASIVE VERSUS NATIVE SPECIES
).6!3)6% 0,!.43 !.$ 3/), 0!4(/'%.3
/NE OF THE LEADING HYPOTHESES FOR THE REMARKABLE SUCCESS OF SOME EXOTIC SPECIES
IS THAT THEY HAVE ESCAPED THE SPECIALIST ENEMIES THAT CONTROL THEM IN THEIR NATIVE
RANGES +EANE AND #RAWLEY %MBEDDED WITHIN THIS HYPOTHESIS IS THE IDEA
THAT IF MICROBIAL PATHOGENS LIMIT THE GROWTH OF INVASIVE PLANTS IN THEIR NATIVE
RANGE THERE WILL BE NEGATIVE FEEDBACKS BETWEEN THE SOIL MICROBIAL COMMUNITY
AND THE PLANT IN THE NATIVE RANGE DUE TO THE ACCUMULATION OF SPECIES
SPECIlC SOIL
PATHOGENS +LIRONOMOS -ITCHELL AND 0OWER #ALLAWAY ET AL
)N CONTRAST POSITIVE FEEDBACKS MAY OCCUR IN THE INVADED RANGE WHERE EXOTIC
)NVASIVE PLANT
SOIL ECOSYSTEM FEEDBACKS
&IG
!3 4HORPE AND 2- #ALLAWAY
SPECIES ARE LARGELY FREE FROM SPECIES
SPECIlC SOIL PATHOGENS BUT CAN STILL INTERACT
WITH LESS HOST
SPECIlC MUTUALISTS SUCH AS MYCORRHIZAL FUNGI AND BACTERIA THAT
DRIVE NUTRIENT CYCLES
)N THIS SECTION lRST WE WILL DESCRIBE THE EVIDENCE THAT INVASIVE SPECIES HAVE
ESCAPED PRESSURE FROM SOIL PATHOGENS AND THE POTENTIAL FEEDBACK EFFECTS ON THE
lTNESS OF INVASIVE SPECIES 3ECOND WE WILL DISCUSS THE IMPLICATIONS OF THIS TYPE OF
FEEDBACK FOR ECOSYSTEM STABILITY
$O INVASIVE SPECIES BENEFIT FROM ESCAPING SOIL PATHOGENS
)N A REVIEW OF SPECIES THAT WERE NATURALIZED IN THE 5NITED 3TATES -ITCHELL
AND 0OWER FOUND THAT FEWER FUNGI AND FEWER VIRUS SPECIES
INFECTED THE PLANT SPECIES IN THEIR INVADED RANGES COMPARED TO THEIR NATIVE RANGES
&URTHERMORE THEY REPORTED THAT SPECIES THAT EXPERIENCED GREATER RELEASE FROM
MICROBIAL PATHOGENS WERE MORE INVASIVE -ITCHELL AND 0OWER (OWEVER
THE RELATIONSHIPS WERE RELATIVELY WEAK PARTICULARLY FOR INVADERS OF NATURAL AREAS
SOIL PATHOGENS WERE NOT DISTINGUISHED FROM OTHER PATHOGENS AND THE PRESENCE
OR ABUNDANCE OF PATHOGENS DOES NOT NECESSARILY CORRELATE WITH THE STRENGTH OF
THEIR EFFECTS
-ORE RECENTLY EXPERIMENTS USING SOILS FROM NATIVE AND INVADED RANGES HAVE
SUGGESTED THAT SOME INVASIVE SPECIES HAVE ESCAPED FROM SOIL PATHOGENS )F INVASIVE
SPECIES SUFFER FROM THE EFFECTS OF SOIL PATHOGENS IN THEIR NATIVE SOILS THEN STERIL
IZATION OF NATIVE SOILS SHOULD RESULT IN AN INCREASE IN THE GROWTH OF THE INVASIVE
)N CONTRAST IN INVADED SOILS THE INVASIVE SHOULD BE RELATIVELY FREE FROM PATHOGENS
BUT MAY BENElT FROM LESS HOST
SPECIlC MUTUALISTIC MICROBES )F SO STERILIZATION
OF INVADED SOILS SHOULD RESULT IN A NEUTRAL TO NEGATIVE EFFECT ON THE INVASIVE SPE
CIES 4HESE INTERACTIONS HAVE BEEN EXPLORED FOR 0RUNUS SEROTINA BLACK CHERRY
#ENTAUREA MACULOSA SPOTTED KNAPWEED AND TWO !CER MAPLE SPECIES
2EINHART ET AL COMPARED THE EFFECTS OF SOIL MICROBES ON THE GROWTH
OF 0RUNUS SEROTINA IN BOTH ITS NATIVE AND INVADED RANGES )N ITS NATIVE .ORTH
!MERICAN RANGE THE SOIL MICROBIAL COMMUNITY OCCURRING NEAR 0 SEROTINA
STRONGLY INHIBITED THE ESTABLISHMENT OF NEIGHBORING CONSPECIlCS AND REDUCED
SEEDLING PERFORMANCE IN THE GREENHOUSE )N CONTRAST IN ITS NON
NATIVE %UROPEAN
RANGE 0 SEROTINA READILY ESTABLISHES IN CLOSE PROXIMITY TO CONSPECIlCS AND SOIL
MICROBIAL COMMUNITIES ENHANCE THE GROWTH OF SEEDLINGS 0REVIOUS RESEARCH IN
THE NATIVE RANGE OF 0 SEROTINA DEMONSTRATED THAT SOIL
BORNE 0YTHIUM SPECIES
/OMYCOTA INHIBIT THE SURVIVAL GROWTH AND ABUNDANCE OF 0 SEROTINA 0ACKER
AND #LAY !LTHOUGH THE GENUS 0YTHIUM IS FOUND AROUND THE WORLD
GENOTYPES ARE OFTEN HOST
SPECIlC $EACON AND $ONALDSON -ILLS AND "EVER
4HUS IN THE NATIVE RANGE 0 SEROTINA EXPERIENCES NEGATIVE PLANT
SOIL
FEEDBACK INTERACTIONS LIKELY DUE TO THE NEGATIVE EFFECTS OF 0YTHIUM )N CONTRAST IN
THE INVADED REGION 0 SEROTINA EXPERIENCES POSITIVE FEEDBACKS DUE TO ESCAPE FROM
ITS MAIN NATURAL ENEMY 2EINHART ET AL
)NVASIVE PLANT
SOIL ECOSYSTEM FEEDBACKS
#ENTAUREA MACULOSA IS ONE OF WESTERN .ORTH !MERICAS WORST INVASIVE WEEDS
)N SEVERAL EXPERIMENTS #ALLAWAY ET AL HAVE COMPARED THE EFFECTS OF SOIL
MICROBES FROM THE NATIVE RANGE IN %UROPE TO THE EFFECTS OF SOIL MICROBES COLLECTED
FROM INVASIVE POPULATIONS IN THE NORTHWESTERN 5NITED 3TATES %UROPEAN SOIL BIOTA
HAD MUCH STRONGER INHIBITORY EFFECTS ON # MACULOSA THAN .ORTH !MERICAN SOIL
BIOTA 3TERILIZATION OF %UROPEAN SOILS CAUSED ON AVERAGE A INCREASE IN
THE TOTAL BIOMASS OF # MACULOSA SUGGESTING A RELEASE FROM PATHOGENIC MICROBES
)N CONTRAST STERILIZING INVADED .ORTH !MERICAN SOILS LED AT MOST TO A SLIGHT
INCREASE IN TOTAL BIOMASS OF &OR MOST .ORTH !MERICAN SOILS STERILIZATION LED
TO A DECREASE IN GROWTH OF
SUGGESTING THAT # MACULOSA HAD BENElTED FROM
MUTUALISTIC SOIL MICROBES 4HESE RESULTS SUPPORT -ITCHELL AND 0OWERS CON
CLUSION THAT INVASIVE SPECIES SHOULD SUFFER MUCH HIGHER FUNGAL AND VIRAL INFECTION
IN THEIR HOME RANGES COMPARED TO INVADED RANGES 4HEY ALSO SUGGEST THAT IN SOME
CASES MUTUALISMS MAY BE MORE BENElCIAL IN NON
NATIVE RANGES BECAUSE THE NEGA
TIVE EFFECT OF NATURAL ENEMIES DO NOT ATTENUATE THE POSITIVE EFFECT OF MUTUALISTS
-UTUALISTS HAVE ALSO BEEN FOUND TO PLAY AN IMPORTANT ROLE IN THE PLANT
SOIL FEEDBACK INTERACTIONS OF TWO !CER SPECIES 2EINHART AND #ALLAWAY
)N THE lELD DISTANCES BETWEEN !CER CONSPECIlCS WERE
LESS IN THEIR INVADED
RANGES THAN IN THEIR NATIVE RANGES )N A GREENHOUSE EXPERIMENT THE EFFECT OF
SOIL MICROBIAL COMMUNITIES ALSO DIFFERED BETWEEN NATIVE AND INVADED RANGES
2ELATIVE TO STERILIZED CONTROLS SOIL ASSOCIATED WITH BOTH CONSPECIlCS AND HET
EROSPECIlCS FROM THE NATIVE RANGE DECREASED THE TOTAL BIOMASS OF !CER SEEDLINGS
BY SUGGESTING INHIBITION BY PATHOGENIC MICROBES )N THE INVADED RANGE
SOIL ASSOCIATED WITH CONSPECIlCS DECREASED THE BIOMASS OF !CER SEEDLINGS BY AN
EVEN GREATER MAGNITUDE (OWEVER SOIL ASSOCIATED WITH HETEROSPECIlCS
IN THE NON
NATIVE RANGES INCREASED BIOMASS OF !CER SEEDLINGS BY 4HUS
WHILE !CERS ACCUMULATE PATHOGENS IN THEIR INVADED RANGE THE SURROUNDING SOIL
IS RELATIVELY FREE FROM INHIBITORY MICROBES POTENTIALLY ENHANCING INVASION BY
THESE TREES
4HUS THERE IS EVIDENCE THAT NOT ONLY DO INVASIVE SPECIES ESCAPE THE NEGA
TIVE EFFECTS OF SOIL PATHOGENS IN THEIR INVADED RANGES BUT THAT POTENTIALLY DUE TO
THE EFFECTS OF MUTUALISTS FEEDBACK EFFECTS IN INVADED RANGES ARE OFTEN POSITIVE
.EXT WE WILL EXPLORE THE POTENTIAL FOR THESE FEEDBACK EFFECTS TO AFFECT COMMUNITY
STABILITY
7ILL ESCAPE FROM NEGATIVE FEEDBACKS FROM SOIL PATHOGENS PERSIST
4HE EXPERIMENTS DESCRIBED ABOVE INDICATE THAT INVASIVE SPECIES ARE LIKELY TO EXPE
RIENCE POSITIVE FEEDBACK IN THEIR INVADED HABITAT BECAUSE THEY ESCAPE SPECIALIST
SOIL PATHOGENS AT HOME BUT CAN UTILIZE GENERALIST MUTUALISTS WHERE THEY INVADE
7HAT REMAINS TO BE DETERMINED IS HOW THIS FEEDBACK AFFECTS COMMUNITY DYNAMICS
0LANTS PARTICIPATING IN STRONG POSITIVE FEEDBACKS WITH SOIL BIOTA ARE MORE LIKELY TO
BECOME COMMUNITY DOMINANTS THAN THOSE THAT DO NOT 4HE MOST COMPLETE STUDY
OF THESE INTERACTIONS WAS DONE BY +LIRONOMOS WHO EXPLORED FEEDBACK
!3 4HORPE AND 2- #ALLAWAY
INTERACTIONS AMONG PLANT SPECIES AND SOIL MICROBIAL COMMUNITIES IN GRASSLANDS IN
EASTERN .ORTH !MERICA )N EXPERIMENTS USING ONLY THE MYCORRHIZAL FRACTION OF THE
MICROBIAL COMMUNITY HE FOUND THAT THE ORIGIN OF THE lLTRATE FROM SOILS IN WHICH
THE SAME SPECIES OR A DIFFERENT SPECIES HAD PREVIOUSLY BEEN GROWN DID NOT ALTER
THE RESPONSE EITHER POSITIVE OR NEUTRAL TO MYCORRHIZAL FUNGI )N CONTRAST
IN EXPERIMENTS USING ONLY THE PATHOGENICSAPROBIC FRACTIONS THE RARE NATIVE
SPECIES EXPERIENCED NEGATIVE FEEDBACKS WHEN THE FRACTIONS WERE FROM SOILS THAT
HAD PREVIOUSLY GROWN THE SAME SPECIES (OWEVER THE ORIGIN OF THE PATHOGENIC
SAPROBIC FRACTION HAD NO EFFECT ON THE GROWTH OF INVASIVE SPECIES /VERALL RELA
TIVELY RARE NATIVE SPECIES CONSISTENTLY EXHIBITED NEGATIVE FEEDBACK INTERACTIONS
WITH THE SOIL MICROBIAL COMMUNITY A RELATIVE DECREASE IN GROWTH ON @HOME
SOIL IN WHICH CONSPECIlCS HAD PREVIOUSLY BEEN GROWN WHEREAS INVASIVE SPE
CIES CONSISTENTLY EXHIBITED POSITIVE FEEDBACK INTERACTIONS WITH THE SOIL COMMU
NITY 3IMILARLY !GRAWAL ET AL FOUND THAT INTRODUCED PLANTS WERE SUBJECT
TO HALF THE NEGATIVE SOIL FEEDBACK AS CONGENERIC SPECIES (OW LONG IN TERMS OF
EVOLUTIONARY TIME SCALES SUCH POSITIVE INTERACTIONS WILL BE MAINTAINED REMAINS
AN UNKNOWN
4HE BASIC NATURE OF MICROBES SUGGESTS THAT THEY WILL BE ABLE TO RESPOND RELA
TIVELY QUICKLY TO PRESSURES EXERTED BY INVASION BY EXOTIC PLANTS !S DISCUSSED
ABOVE SOIL
BORNE PATHOGENS CAN BE RELATIVELY HOST SPECIlC .EERGAARD
+IRKPATRICK AND "AZZAZ !GARWAL AND 3INCLAIR -ILLS AND "EVER
(OWEVER MANY SOIL
BORNE PATHOGENS ARE GENERALISTS $IX AND 7EBSTER
&OR EXAMPLE "LANEY AND +OTANEN FOUND THAT SEED GERMINATION OF
CONGENERIC PAIRS OF INVASIVE AND NATIVE PLANT SPECIES FROM WESTERN /NTARIO DIS
PLAYED A SIMILAR POSITIVE RESPONSE TO THE APPLICATION OF FUNGICIDE SUGGESTING
NON
SPECIES SPECIlCITY OF FUNGAL SEED PATHOGENS IN THEIR SYSTEM &URTHERMORE
MICROBES HAVE A SHORT GENERATION TIME AND THUS CAN RESPOND TO EVOLUTIONARY
PRESSURES WITHIN A SHORT TIMEFRAME 4HUS PATHOGENIC MICROBES MAY BE ABLE TO
RAPIDLY SWITCH TO A NEW INVASIVE HOST 3CLEROTINIA SCLEROTIORUM A FUNGUS NATIVE
TO INTERMOUNTAIN PRAIRIES INVADED BY # MACULOSA HAS BEEN FOUND TO DAMAGE
# MACULOSA WHEN APPLIED TO THE RHIZOSPHERES AT HIGH CONCENTRATIONS *ACOBS
ET AL 2IDENOUR AND #ALLAWAY !S DESCRIBED ABOVE 2EINHART AND
#ALLAWAY FOUND THAT WHILE THE SOIL COMMUNITY ASSOCIATED WITH OTHER TREE
SPECIES HAD A POSITIVE EFFECT ON THE GROWTH OF INVASIVE !CERS SOILS ASSOCIATED WITH
CONSPECIlCS HAD A NEGATIVE EFFECT 4HIS SUGGESTS THAT THE PATHOGENIC SOIL MICROBIAL
COMMUNITY MAY HAVE BEEN ABLE TO ADAPT TO THIS NEW HOST AND ACCUMULATION OF
SOIL PATHOGENS EVENTUALLY SUPPRESSES THE OFFSPRING OF !CER RECRUITS
)N SUMMARY SOME INVASIVE SPECIES APPEAR TO HAVE ESCAPED PRESSURE FROM SOIL
PATHOGENS AND THUS BENElT FROM POSITIVE FEEDBACK INTERACTIONS WITH THE SOIL BIOTA
WHERE THEY INVADE !LTHOUGH NOT YET EXPLICITLY ADDRESSED IN THE LITERATURE MICRO
BIAL COMMUNITIES MAY CHANGE OVER TIME AND THUS BREAK DOWN POSITIVE PLANT
SOIL
MICROBIAL FEEDBACKS )F THIS OCCURS THE ABUNDANCE OF THE INVASIVE SPECIES SHOULD
DECREASE AND THE COMMUNITY SHOULD MOVE TO A POINT WHERE NEGATIVE FEEDBACK
INTERACTIONS RESTRICT THE INVASIVES DOMINANCE 4HIS WAS ORIGINALLY SUGGESTED BY
)NVASIVE PLANT
SOIL ECOSYSTEM FEEDBACKS
+LIRONOMOS WHO OBSERVED THAT PLANT
SPECIlC PATHOGEN LOADS ARE MAXI
MIZED UNDER HIGH POPULATION DENSITIES PARTICULARLY MONOCULTURES SUCH AS THOSE
CREATED BY SOME INVASIVE SPECIES AND WILL EVENTUALLY RESULT IN NEGATIVE FEEDBACK
ON ABUNDANT PLANTS .EXT WE CONSIDER HOW FEEDBACKS DRIVEN BY MICROBES IN NUTRI
ENT CYCLES MIGHT RESPOND DIFFERENTLY OVER EVOLUTIONARY TIME THAN FEEDBACKS DRIVEN
BY SOIL PATHOGENS
).6!3)6% 0,!.43 !.$ 3/), .542)%.4 #9#,).'
)N GENERAL PLANT SOIL FEEDBACKS ARE THOUGHT TO BE DETERMINED BY THE DIRECT EFFECTS
OF PATHOGENS AND MUTUALISTS "EVER -ILLS AND "EVER 0ACKER AND #LAY
"EVER +LIRONOMOS BUT OTHER COMPONENTS OF THE SOIL ECOSYSTEM
MAY PARTICIPATE IN FEEDBACKS )N PARTICULAR INDIVIDUAL PLANT CHARACTERISTICS SUCH
AS PHENOLOGY NUTRIENT UPTAKE LITTER FALL TISSUE CHEMICAL COMPOSITION AND ASSOCI
ATION WITH SYMBIOTIC MICROBES CAN HAVE SIGNIlCANT EFFECTS ON SOIL NUTRIENT CYCLES
(OBBIE !NGERS AND #ARON "ERENDSE "INKLEY AND 'IARDINA
.ORTHRUP ET AL 3CHLESINGER AND 0ILMANIS 6AN "REEMEN
7ARDLE ET AL #HEN AND 3TARK %ATON AND &ARRELL WHICH MAY IN
TURN ALTER THE GROWTH AND SURVIVAL OF THE SPECIES THAT DRIVE THESE EFFECTS "ECAUSE
THEY ARE NOVEL MAY HAVE DIFFERENT BIOCHEMICAL CONSTITUENTS "AIS ET AL
6IVANCO ET AL AND ARE OFTEN DOMINANT COMPONENTS OF PLANT COMMUNITIES
INVASIVE PLANTS CAN HAVE UNUSUALLY STRONG EFFECTS ON SOIL NUTRIENT CYCLES 6ITOUSEK
6ITOUSEK ET AL 6ITOUSEK $!NTONIO AND 6ITOUSEK
%HRENFELD ET AL %HRENFELD AND 3COTT %HRENFELD
)N THE PREVIOUS SECTION WE SHOWED THAT MANY INVASIVE SPECIES EXHIBIT POSITIVE
FEEDBACKS AFTER ESCAPING SOIL PATHOGENS AND THEN SPECULATED THAT THESE FEEDBACKS
MAY EVENTUALLY BECOME NEUTRAL OR NEGATIVE AS GENERALIST PATHOGENS SWITCH TO THE
HOST OR SPECIALISTS ADAPT )N CONTRAST TO THIS SCENARIO IN WHICH INVASIVE DOMINANCE
MAY FADE POSITIVE FEEDBACKS BETWEEN INVASIVE PLANTS AND SOIL NUTRIENT CYCLES MAY
LEAD TO MUCH LONGER TIME PERIODS OF INVASIVE DOMINANCE .UTRITIONAL CONSTRAINTS
MAY LEAD TO SIGNIlCANT SHIFTS IN MICROBIAL COMMUNITIES RESULTING IN LONG TERM
CHANGES IN NUTRIENT POOLS AND CYCLING RATES 4HESE CHANGES MAY ALSO OCCUR DUE TO
THE INTRODUCTION OF NOVEL PLANT MICROBE INTERACTIONS SUCH AS SYMBIOTIC NITROGEN
lXATION 3IMILARLY THE DIRECT EFFECTS OF INVASIVE PLANTS ON SOIL NUTRIENTS MAY BE
PARTICULARLY LONG LIVED BECAUSE THERE IS NO MEDIATION BY ANOTHER ORGANISM WITH
THE POTENTIAL TO EVOLVE
)N THIS SECTION WE BRIEmY REVIEW THE MECHANISMS BY WHICH INVASIVE SPE
CIES MAY ALTER SOIL NUTRIENT CYCLES AND ILLUSTRATE THE POTENTIAL FOR LONG LIVED
POSITIVE FEEDBACK INTERACTIONS BY DESCRIBING THE INTERACTIONS OF "ROMUS TECTORUM
CHEATGRASS DOWNY BROME AND -YRICA FAYA lRE TREE WITH SOIL NUTRIENT CYCLES IN
INVADED COMMUNITIES IN THE WESTERN 5NITED 3TATES AND (AWAII
!3 4HORPE AND 2- #ALLAWAY
4HE EFFECTS OF INVASIVE SPECIES ON NUTRIENT CYCLES
4HERE ARE MANY MECHANISMS BY WHICH INVASIVE SPECIES MAY ALTER SOIL NUTRIENT
CYCLES SEE REVIEW BY %HRENFELD 4HROUGH CHANGES IN LITTER PRODUCTION AND
QUALITY INVASIVE PLANTS MAY INCREASE %HRENFELD ET AL -ACK ET AL OR
DECREASE 3AGGAR ET AL %HRENFELD ET AL %VANS ET AL MICROBI
ALLY
MEDIATED DECOMPOSITION ANDOR MINERALIZATION RATES &OR EXAMPLE LITTER OF
-ICROSTEGIUM VIMINEUM AN EXOTIC # GRASS THAT HAS INVADED %ASTERN DECIDUOUS
FORESTS HAS A HIGHER #. RATIO DECOMPOSES SLOWER AND IMMOBILIZES MORE . THAN
LITTER FROM UNINVADED FORESTS %HRENFELD ET AL
)NVASIVE SPECIES MAY ALSO ALTER THE INPUT OF NITROGEN BY NITROGEN
lXING BACTERIA
.EARLY OF THE INVASIVE SPECIES LISTED BY THE 53 $EPARTMENT OF !GRICULTURE
ARE IN THE &ABACEAE FAMILY %HRENFELD AND CHANGES IN ECOSYSTEM NITROGEN
AVAILABILITY DUE TO ASSOCIATION OF INVASIVE PLANTS WITH SYMBIOTIC NITROGEN
lXING
BACTERIA HAVE BEEN DOCUMENTED IN SEVERAL ECOSYSTEMS 6ERSFELD AND VAN 7ILGREN
6ITOUSEK ET AL 3TOCK ET AL 9ELENIK ET AL &URTHERMORE
CHANGES IN LITTER QUALITY FROM N ON
NITROGEN lXING INVADERS MAY
ALTER THE ABUN
DANCE AND ACTIVITY OF NON
SYMBIOTIC NITROGEN
lXING BACTERIA AS FOUND IN
(AWAIIAN FORESTS INVADED BY !FRICAN GRASSES ,EY AND $!NTONIO
)NVASIVE SPECIES MAY AFFECT SOIL NUTRIENT CYCLES THROUGH THE PRODUCTION
OF SECONDARY CHEMICALS 2OOTS OF #ENTAUREA MACULOSA EXUDE THE POLYPHENOL
CATECHIN
#ATECHIN DISPLAYS STRONG ANTIMICROBIAL PROPERTIES FOR AT LEAST
SOME GROUPS OF BACTERIA "AIS ET AL AND APPEARS TO AFFECT AT
LEAST SOME ASPECTS OF THE SOIL NITROGEN CYCLE ! 4HORPE UNPUBLISHED DATA
&URTHERMORE BY CHELATING METAL
PHOSPHORUS COMPLEXES CATECHIN MAY INCREASE
PHOSPHORUS AVAILABILITY IN PHOSPHORUS
LIMITED SOILS 4HORPE ET AL IN PRESS
3TEVENSON AND #OLE !N ALLELOCHEMICAL PRODUCED BY #ENTAUREA DIFFU
SA
HYDROXYQUINOLINE MAY ALSO ALTER NUTRIENT CYCLING THROUGH ANTIMICRO
BIAL 6IVANCO ET AL AND CHELATION 4HE -ERCK )NDEX PROPERTIES
4HE DRY MASS OF LEAVES OF -ELALEUCA SPP PAPERBARK WHICH HAS INVADED LARGE
AREAS OF THE COASTAL SOUTHEAST 5NITED 3TATES PARTICULARLY THE %VERGLADES IS UP TO
MONOTERPENES "OON AND *OHNSTONE 4HESE COMPOUNDS INHIBIT MICRO
BIAL COLONIZATION AND DECOMPOSITION OF LEAF LITTER IN BOTH THE NATIVE AND INVADED
RANGES OF -ELALEUCA SPP "OONE AND *OHNSTONE )T HAS ALSO BEEN SUGGESTED
THAT ALLELOPATHIC CHEMICALS RELEASED BY SOME INVASIVE SPECIES MAY ALTER NITROGEN
lXATION IN NEIGHBORING PLANTS 7ARDLE ET AL -ANY OTHER INVASIVE
SPECIES PRODUCE CHEMICALS WITH ANTIMICROBIAL ACTIVITY 2ICE %HRENFELD
HOWEVER THE ROLE OF THESE CHEMICALS IN THE PLANTS INVASIVE SUCCESS IS
GENERALLY UNKNOWN
)N SUM THERE IS GOOD EVIDENCE THAT BY INTRODUCING A NOVEL CHARACTERISTIC
EG A HIGHER #. RATIO ASSOCIATION WITH NITROGEN
lXING BACTERIA OR EXUDATION
OF AN ANTI
MICROBIAL CHEMICAL INVASIVE SPECIES CAN ALTER SOIL NUTRIENT CYCLES
IN INVADED COMMUNITIES !LTHOUGH EXPLICIT STUDIES OF THE RAMIlCATIONS OF SUCH
ALTERATION OF NUTRIENT CYCLES ARE RARE THESE EFFECTS MAY ULTIMATELY FEEDBACK TO
)NVASIVE PLANT
SOIL ECOSYSTEM FEEDBACKS
THE PLANTS THAT CAUSE THEM AND AFFECT THE ORGANIZATION OF PLANT COMMUNITIES
4WO DIFFERENT SPECIES "ROMUS TECTORUM AND -YRICA FAYA PROVIDE EXCELLENT EXAM
PLES OF HOW INVASIVE PLANTS MAY AFFECT THE SOIL AND HOW SOIL CHANGES
AFFECT THE SURVIVAL OF INVASIVE AND NATIVE SPECIES 4HESE STUDIES ALSO ILLUSTRATE HOW
POSITIVE FEEDBACKS BETWEEN INVASIVE PLANTS AND SOIL NUTRIENT CYCLES MAY PERSIST
IN AN ECOSYSTEM
4HE EFFECTS OF "ROMUS TECTORUM ON SOIL NUTRIENT CYCLES
"ROMUS TECTORUM IS AN ANNUAL OCCASIONALLY BIENNIAL %URASIAN GRASS THAT HAS
INVADED OVER MILLION HA IN THE )NTERMOUNTAIN 7EST OF .ORTH !MERICA
9PSILANTIS 4HE EFFECTS OF " TECTORUM ON NUTRIENT CYCLES DIFFER IN lRE
PRONE
AND NON
lRE
PRONE SYSTEMS
"ROMUS TECTORUM TENDS TO GERMINATE AND COMPLETE ITS LIFE CYCLE EARLIER THAN MOST
NATIVE SPECIES IN THE SYSTEMS IT INVADES AND ITS DEAD DRY STEMS CREATE AN UNUSU
ALLY LARGE FUEL LOAD IN THE SUMMER (ARRIS -ACK 5PADHYAYA ET AL
9PSILANTIS )N lRE
PRONE SAGEBRUSH
GRASSLAND ECOSYSTEMS lRE RECUR
RENCE INTERVALS DECREASE FROM
YEARS TO
YEARS -ACK 5PADHYAYA
ET AL 9PSILANTIS 3INCE " TECTORUM GERMINATES EARLIER AND GROWS
FASTER THAN MOST NATIVE SPECIES (ARRIS -ACK 5PADHYAYA ET AL
9PSILANTIS THIS INVADER APPEARS TO TAKE BETTER ADVANTAGE OF THE POST
lRE
mUSH OF NITROGEN THAN NATIVE SPECIES ,OWE ET AL %ARLY NITROGEN UPTAKE BY
" TECTORUM REDUCES TOTAL SOIL NITROGEN AND CREATES HIGHER SOIL CARBON TO NITROGEN
RATIOS THAN NATIVE VEGETATION "LANK ET AL (ALVORSON ET AL "ROMUS
TECTORUM MAY ALSO LIMIT NITROGEN AVAILABILITY BY SHADING BIOLOGICAL SOIL CRUSTS THAT
lX NITROGEN 9PSILANTIS
)N ECOSYSTEMS THAT LACK lRE THERE ARE VERY DIFFERENT INTERACTIONS BETWEEN
" TECTORUM AND THE SOIL ECOSYSTEM 'RASSLAND COMMUNITIES IN 5TAH INVADED BY
" TECTORUM HAVE HIGHER LEVELS OF EXCHANGEABLE POTASSIUM AND RATIOS OF POTAS
SIUM OR PHOSPHORUS TO CALCIUM CARBONATE AND MAGNESIUM OR IRON OXIDES THAN
UNINVADED SOILS "ELNAP AND 0HILLIPS "ELNAP ET AL )T IS UNKNOWN
WHETHER THESE NUTRIENT DIFFERENCES ARE DUE TO " TECTORUM INVASION OR IF " TECTORUM
PREFERENTIALLY INVADES SITES WITH THESE CHARACTERISTICS HOWEVER IT IS CLEAR THAT
" TECTORUM CAN DRAMATICALLY ALTER PHOSPHORUS CYCLING IN I NVADED SOILS !LTHOUGH
THERE IS NO NET CHANGE IN TOTAL SOIL PHOSPHORUS POOLS " TECTORUM APPEARS TO ACCESS
FORMS OF 0 THAT ARE RECALCITRANT AND UNAVAILABLE TO NATIVES WHICH INCREASES LEVELS
OF LABILE PHOSPHORUS 2, 3ANFORD PERSONAL COMMUNICATION
"Y ALTERING THE BIOTIC AND ABIOTIC COMPONENTS OF NUTRIENT CYCLES " TECTORUM
ALTERS NUTRIENT AVAILABILITY IN WAYS THAT ULTIMATELY FEEDBACK TO INCREASE ITS OWN
SURVIVAL RELATIVE TO NATIVE SPECIES "ELNAP AND 0HILLIPS %VANS ET AL
"Y ALTERING FUNDAMENTAL ECOSYSTEM CHARACTERISTICS THESE EFFECTS MAY SIGNIlCANTLY
ALTER PLANT COMMUNITY STRUCTURE AND DYNAMICS
!3 4HORPE AND 2- #ALLAWAY
4HE EFFECTS OF -YRICA FAYA ON SOIL NUTRIENT CYCLES
)NVASION OF (AWAIIAN ECOSYSTEMS BY THE NITROGEN lXING TREE -YRICA FAYA 'ERRISH
AND -UELLER
$OMBOIS 6ITOUSEK 6ITOUSEK AND 7ALKER
!PLET (UGHES ET AL PROVIDES AN EXAMPLE OF HOW ECOSYSTEM SCALE
CHANGES MAY RESULT IN hINVASIONAL MELTDOWNv "OX 3IMBERLOFF AND VAN (OLLE
4HE VOLCANIC SOILS OF (AWAII ARE USUALLY NITROGEN
LIMITED 6ITOUSEK
6ITOUSEK AND 7ALKER 4HERE ARE NO NATIVE NITROGEN
lXING PLANTS THAT
COLONIZE EARLY SUCCESSIONAL HABITATS IN THESE SYSTEMS 6ITOUSEK 6ITOUSEK
AND 7ALKER AND INVASION BY - FAYA INTRODUCES A NOVEL ECOSYSTEM PROCESS
THAT RESULTS IN SUBSTANTIAL INCREASES IN SOIL NITROGEN 6ITOUSEK 6ITOUSEK
AND 7ALKER -OST SUCCESSFUL INVASIONS IN (AWAII HAVE OCCURRED ON SOILS
THAT ARE RELATIVELY FERTILE 'ERRISH AND -UELLER
$OMBOIS AND THE NITROGEN
FERTILIZATION THAT RESULTS FROM INVASION BY - FAYA MAY FACILITATE INVASION BY OTHER
SPECIES WITH HIGHER NITROGEN REQUIREMENTS &OR EXAMPLE ANOTHER INVASIVE TREE
0SIDIUM CATTLEIANUM STRAWBERRY GUAVA GREW MUCH LARGER WHEN GROWN IN SOIL
FROM UNDER - FAYA THAN FROM SOIL COLLECTED UNDER THE NATIVE -ETROSIDEROS POLY
MORPHA {/HI{A LEHUA (UGHES ET AL -YRICA FAYA ALSO ENHANCES POPULATIONS
OF EXOTIC EARTHWORMS WHICH INCREASE NITROGEN BURIAL AND FURTHER ALTER NUTRIENT
CYCLING !PLET 4HUS INVASION BY - FAYA RESULTS IN POSITIVE FEEDBACKS THAT
NOT ONLY ENHANCE THE INVASION OF THIS SPECIES BUT PROMOTE INVASION BY OTHER
INVASIVE SPECIES AND LEAD TO FUNDAMENTAL ECOSYSTEM CHANGES
"OX
)N 3IMBERLOFF AND VAN (OLLE INTRODUCED THE CONCEPT OF hINVASIONAL
MELTDOWNv $URING THIS PROCESS INVASION BY ONE EXOTIC SPECIES PROMOTES
INVASION BY OTHER EXOTIC SPECIES 4HESE SPECIES INTERACT SYNERGISTICALLY
CAUSING ECOSYSTEM SCALE CHANGES SUCH AS MORE FREQUENT lRE CYCLES OR
MORE RAPID NITROGEN CYCLING THROUGH INCREASED NITROGEN lXATION THAT
ULTIMATELY RESULT IN THE COLLAPSE OF NATIVE ECOSYSTEMS
7ILL POSITIVE FEEDBACKS BETWEEN INVASIVE PLANTS AND SOIL NUTRIENT
CYCLES PERSIST
!S DESCRIBED ABOVE SINCE INVASIVE PLANTS OFTEN DIFFER FROM NATIVE SPECIES IN
CHARACTERISTICS SUCH AS PHENOLOGY NUTRIENT UPTAKE LITTER
FALL TISSUE CHEMICAL
COMPOSITION AND ASSOCIATION WITH SYMBIOTIC MICROBES THEY CAN HAVE SIGNIlCANT
EFFECTS ON SOIL NUTRIENT CYCLES 3IMILAR EFFECTS ARE OFTEN SEEN DURING SUCCESSION
"ERENDSE 3CHIMEL AND "ENNETT 7E SPECULATE THAT IN CONTRAST TO
OUR PREDICTIONS FOR EVOLUTIONARY CHANGE FOR INTERACTIONS BETWEEN INVASIVE PLANTS
)NVASIVE PLANT
SOIL ECOSYSTEM FEEDBACKS
AND SOIL PATHOGENS POSITIVE FEEDBACKS BETWEEN INVASIVE PLANTS AND SOIL NUTRIENT
CYCLES ARE MORE LIKELY TO RESULT IN LONG TERM SHIFTS IN COMMUNITY COMPOSITION THAT
RESEMBLE SUCCESSION 4HESE CHANGES MAY PROVIDE EXAMPLES OF INVASIONAL MELT
DOWN 3IMBERLOFF AND 6ON (OLLE
7E KNOW OF NO EXAMPLES WHERE DURING SUCCESSION CHARACTERISTICS OF THE SOIL
ECOSYSTEM REVERT BACK TO THOSE RESEMBLING THE INITIAL CHARACTERISTICS AFTER DISTUR
BANCE 7HAT LITTLE IS KNOWN ABOUT THE BIOTIC COMPONENTS OF THE SOIL ECOSYSTEM
DURING SUCCESSION INDICATES THAT CHANGE IN THE MICROBIAL COMMUNITY IS TOWARDS
POPULATIONS THAT ARE BETTER ADAPTED TO THE NEW NUTRIENT CONDITIONS OF THE LATER
SUCCESSIONAL STAGES 3CHIMEL ET AL 3CHIMEL AND "ENNETT 4HESE SUC
CESSIONAL SHIFTS APPEAR TO BE ECOLOGICAL IN NATURE NOT EVOLUTIONARY AND DRIVEN
BY SHIFTS IN THE SPECIES COMPOSITION OF COMMUNITIES OVER TIME )N CONTRAST IN THE
CASE OF ALTERATIONS TO ABIOTIC COMPONENTS OF THE SOIL ECOSYSTEM THERE IS NO DIRECT
SELECTIVE PRESSURE TO STIMULATE ADAPTATION 5LTIMATELY THESE TYPES OF CHANGES
REQUIRE THAT THE SOIL AND PLANT COMMUNITIES MAY HAVE TO RESPOND ADAPTIVELY TO
NEW NUTRIENT CONDITIONS
7HEN INVADER
SOIL ECOSYSTEM FEEDBACKS OCCUR THROUGH EFFECTS ON NUTRIENT
CYCLES WE SUGGEST THAT LONG TERM EFFECTS WILL BE SIMILAR TO THOSE THAT OCCUR DURING
SUCCESSION THE DEVELOPMENT OF A NEW COMMUNITY COMPOSED OF SPECIES THAT ARE
BETTER ADAPTED TO THE SPECIlC NUTRIENT CONDITIONS OF THE SOIL EG "ERENDSE
4HIS HAS BEEN OBSERVED IN SYSTEMS INVADED BY BOTH NATIVE -ARON AND *EFFERIES
AND INVASIVE (UGHES ET AL NITROGEN
lXING SPECIES )N #ALIFORNIA
-ARON AND #ONNORS FOUND THAT NITROGEN
RICH PATCHES THAT WERE LEFT BY
DEATH OF ,UPINUS ARBOREOUS BUSH LUPINE WERE INVADED BY EXOTIC ANNUAL GRASSES
3IMILARLY AS DESCRIBED IN THE PRECEDING SECTION SOILS THAT HAVE BEEN ENRICHED BY
THE EXOTIC NITROGEN
lXER - FAYA ARE MORE PRONE TO INVASION BY OTHER EXOTIC PLANTS
INCLUDING 0SIDIUM CATTLEIANUM STRAWBERRY GUAVA (UGHES ET AL )N ADDI
TION ALTHOUGH ONLY A FEW NATIVE SPECIES HAVE BEEN FOUND TO hMONOPOLIZEv SITES
BY CREATING POSITIVE FEEDBACKS VAN DER 0UTTEN INVASIVE SPECIES MAY BE
PARTICULARLY CAPABLE OF THIS ESPECIALLY IF THEY INTRODUCE A NOVEL ECOSYSTEM PROCESS
TO WHICH NATIVE SPECIES ARE NOT ADAPTED &OR EXAMPLE ALTHOUGH " TECTORUM lRST
BECAME ESTABLISHED IN lRE
PRONE COMMUNITIES IN THE 'REAT "ASIN OVER YEARS
AGO -ACK NO OTHER SPECIES EITHER NATIVE OR INVASIVE HAVE SINCE BEEN ABLE
TO ESTABLISH IN SUBSTANTIAL NUMBERS 4HESE EXAMPLES SHOW HOW ECOSYSTEM CHANG
ES INVOLVING EXOTIC SPECIES CAN RESULT IN UNPREDICTABLE SUCCESSIONAL TRAJECTORIES
#/.#,53)/.
%XOTIC PLANT INVASIONS CAN OFTEN INVOLVE POSITIVE FEEDBACKS BETWEEN THE INVADER
AND THE NATIVE SOIL COMMUNITY 0OSITIVE FEEDBACKS HAVE THE POTENTIAL TO LEAD TO
LONG
LASTING DOMINANCE IN COMMUNITIES (OWEVER THESE FEEDBACKS CAN BE CAUSED
BY A NUMBER OF DIFFERENT REASONS AND DIFFERENT DRIVERS OF POSITIVE FEEDBACKS MAY
RESULT IN DIFFERENT PREDICTIONS O F HOW LONG I NVADERS M
AY DOMINATE A COMMUNITY
!3 4HORPE AND 2- #ALLAWAY
4YPICALLY POSITIVE FEEDBACKS ARE THOUGHT TO BE CAUSED BY THE ABSENCE OF SOIL
PATHOGENS BUT THE PRESENCE OF SOIL MUTUALISTS IN INVADED SOILS 3UCH PATHOGEN
DRIVEN POSITIVE FEEDBACKS MAY RESULT IN LONG TERM INVASIVE DOMINANCE BUT
EVOLUTIONARY CHANGES IN NATIVE PATHOGENS MIGHT ULTIMATELY LEAD TO SUPPRESSION
OF THE INVADER AND A RETURN TO NATIVE COEXISTENCE 0OSITIVE FEEDBACKS CAN ALSO BE
CAUSED BY THE EFFECTS OF INVASIVE PLANTS ON THE SOIL BIOTA THAT DRIVE NUTRIENT CYCLES
OR ON THE ABIOTIC COMPONENTS OF THE NUTRIENT CYCLES THEMSELVES 7E HYPOTHESIZE
THAT POSITIVE FEEDBACKS CAUSED BY THE EFFECTS OF THE INVADER ON THE SOIL BIOTA
INVOLVED IN NUTRIENT CYCLES ARE LIKELY TO LEAD TO COMMUNITY SHIFTS RESEMBLING
SUCCESSION AND PERHAPS INVASIONAL MELTDOWN $OMINANCE BY INVADERS THAT DRIVE
POSITIVE FEEDBACKS THROUGH ABIOTIC COMPONENTS OF NUTRIENT CYCLES MAY PERSIST
FOR MUCH LONGER THAN DOMINANCE OCCURRING THROUGH POSITIVE FEEDBACKS INVOLV
ING THE MICROBIAL COMMUNITY DUE TO THE LACK OF MEDIATION BY AN ORGANISM WITH
THE POTENTIAL TO ADAPT (OWEVER UNDERSTANDING POTENTIAL EVOLUTIONARY CHANGES
AMONG INVADERS AND SOIL ECOSYSTEMS WILL HELP US TO ACCURATELY PREDICT THE LONG
TERM EFFECTS OF BIOLOGICAL INVASIONS
!#+./7,%$'%-%.43
4HE AUTHORS THANK THE .3& 53$! .2) AND THE $/$ FOR lNANCIAL SUPPORT
2%&%2%.#%3
!PLET ' ( !LTERNATION OF EARTHWORM COMMUNITY BIOMASS BY THE ALIEN -YRICA FAYA
IN (AWAII /ECOLOGIA
!GARWAL 6 + AND * " 3INCLAIR 0RINCIPLES OF 3EED 0ATHOLOGY 3ECOND EDITION ,EWIS
0UBLISHERS "OCA 2ATON
!GRAWAL ! ! 0 - +OTANEN # % -ITCHELL ! ' 0OWER 7 'ODSOE AND * +LIRONOMOS
%NEMY 2ELEASE !N EXPERIMENT WITH CONGENERIC PLANT PAIRS AND DIVERSE ABOVE
AND BELOW
GROUND ENEMIES %COLOGY
!NGERS $ ! AND * #ARON 0LANT
INDUCED CHANGES IN SOIL STRUCTURE PROCESSES AND
FEEDBACKS "IOGEOCHEMISTRY
!TLAS 2 - AND 2 "ARTHA -ICROBIAL %COLOGY &UNDAMENTALS AND !PPLICATIONS
"ENJAMIN #UMMINGS .EW 9ORK
"AIS ( 0 2 6EPACHEDU 3 'ILROY 2 - #ALLAWAY AND * - 6IVANCO !LLELOPATHY
AND EXOTIC PLANTS FROM GENES TO INVASION 3CIENCE
"AIS ( 0 4 3 7ALKER & 2 3TERMITZ 2 ! (UFBAUER AND * - 6IVANCO
%NANTIOMERIC
DEPENDENT PHYTOTOXIC AND ANTIMICROBIAL ACTIVITY OF
CATECHIN ! RHI
ZOSECRETED RACEMIC MIXTURE FROM SPOTTED KNAPWEED 0LANT 0HYSIOLOGY
"ELNAP * AND 3 , 0HILLIPS 3OIL BIOTA IN AN UNGRAZED GRASSLAND RESPONSE TO ANNUAL
GRASS "ROMUS TECTORUM INVASION %COLOGICAL !PPLICATIONS
)NVASIVE PLANT
SOIL ECOSYSTEM FEEDBACKS
"ELNAP * 3 + 3HERROD - % -ILLER %FFECTS OF SOIL AMENDMENTS ON GERMINATION
AND EMERGENCE OF DOWNY BROME "ROMUS TECTORUM AND (ILARIA JAMESII 7EED 3CIENCE
"ERENDSE & %FFECTS OF DOMINANT PLANT SPECIES ON SOILS DURING SUCCESSION IN NUTRIENT
POOR ECOSYSTEMS "IOGEOCHEMISTRY
"EVER * $ &EEDBACK BETWEEN PLANTS AND THEIR SOIL COMMUNITIES IN AN OLD lELD COM
MUNITY %COLOGY
"EVER * $ * " -ORTON * !NTONOVICS AND 0 ! 3CHULTZ (OST
DEPENDENT SPORULA
TION AND SPECIES DIVERSITY OF ARBUSUCLAR MYORRHIZAL FUNGI IN A MOWN GRASSLAND *OURNAL
OF %COLOGY
"EVER * $ + - 7ESTOVER AND * !NTONOVICS )NCORPORATING THE SOIL COMMUNITY
INTO PLANT POPULATION DYNAMICS THE UTILITY OF THE FEEDBACK APPROACH *OURNAL OF %COLOGY
"EVER * $ .EGATIVE FEEDBACK WITHIN A MUTUALISM HOST
SPECIlC GROWTH OF MYCOR
RHIZAL FUNGI REDUCES PLANT BENElT 0ROCEEDINGS 2OYAL 3OCIETY ,UND "
"INKLEY $ AND # 'IARDINA 7HY DO TREE SPECIES AFFECT SOILS 4HE WARP AND WOOF OF
TREE
SOIL INTERACTIONS "IOGEOCHEMISTRY
"LANEY # 3 AND 0 - +OTANEN %FFECTS OF FUNGAL PATHOGENS ON SEEDS OF NATIVE AND
EXOTIC PLANTS A TEST USING CONGENERIC PAIRS *OURNAL OF !PPLIED %COLOGY
"LANK 2 2 & !LLEN AND * ! 9OUNG %XTRACTABLE ANIONS IN SOILS FOLLOWING WILDlRE
IN A SAGEBRUSH
GRASS COMMUNITY 3OIL 3CIENCE 3OCIETY OF !MERICA *OURNAL
"OON 0 ) AND , *OHNSTONE /RGANIC MATTER DECAY IN COASTAL WETLANDS AN INHIBI
TORY ROLE FOR ESSENTIAL OIL FROM -ELALEUCA ALTERNIFOLIA LEAVES !RCHIV FUR HYDROBIOLGIE
#ALLAWAY 2 - ' # 4HELEN 3 "ARTH 0 7 2AMSEY AND * % 'ANNON 3OIL FUNGI
ALTER INTERACTIONS BETWEEN .ORTH !MERICAN PLANT SPECIES AND THE EXOTIC INVADER
#ENTAUREA MACULOSA IN THE lELD %COLOGY
#ALLAWAY 2 - ' # 4HELEN ! 2ODRIGUEZ AND 7 % (OLBEN 2ELEASE FROM INHIBI
TORY SOIL BIOTA IN %UROPE MAY PROMOTE EXOTIC PLANT INVASION IN .ORTH !MERICA .ATURE
#HEN * AND * - 3TARK 0LANT SPECIES EFFECTS AND CARBON AND NITROGEN CYCLING IN
A SAGEBRUSH
CRESTED WHEATGRASS SOIL 3OIL "IOLOGY AND "IOCHEMISTRY
#HIARELLO . * # (ICKMAN AND ( ! -OONEY %NDOMYCORRHIZAL ROLE IN INTERSPECIlC
TRANSFER OF PHOSPHORUS IN A COMMUNITY OF ANNUAL PLANTS 3CIENCE
#LAY + AND 7 ( 6AN DER 0UTTEN 0ATHOGENS AND PLANT LIFE
HISTORIES 0AGES
IN 4 / 6UORISALO AND 0 + -UTIKAINEN EDITORS ,IFE (ISTORY IN 0LANTS
+LUWER !CADEMIC 0UBLISHERS $ORDRECHT
$!NTONIO # - AND 0 - 6ITOUSEK "IOLOGICAL INVASION BY EXOTIC GRASSES THE GRASS
lRE CYCLE AND GLOBAL CHANGE !NNUAL 2EVIEW OF %COLOGY AND 3YSTEMATICS
$EACON * 7 AND $ONALDSON 3 0 -OLECULAR RECOGNITION IN THE HOMING RESPONSES
OF ZOOSPORIC FUNGI WITH SPECIAL REFERENCE TO 0YTHIUM AND 0HYTOPHTHORA -YCOLOGICAL
2ESEARCH
$IX . AND * 7EBSTER &UNGAL %COLOGY #HAPMAN AND (ALL ,ONDON 5NITED
+INGDOM
!3 4HORPE AND 2- #ALLAWAY
%ATON 7 $ AND 2 % &ARRELL #ATABOLIC AND GENETIC MICROBIAL INDICES AND LEVELS
OF NITRATE AMMONIUM AND ORGANIC CARBON IN SOIL FROM THE BLACK LOCUST 2OBINIA PSEUDO
ACACIA AND TULIP POPLAR ,IRIODENDRON TULIPIFERA TREES IN A 0ENNSYLVANIA FOREST "IOLOGICAL
&ERTILITY OF 3OILS
%HRENFELD * ' %FFECTS OF EXOTIC PLANT INVASIONS ON SOIL NUTRIENT CYCLING PROCESSES
%COSYSTEMS
%HRENFELD * ' 0 +OURTEV AND 7 (UANG #HANGES IN SOIL FUNCTIONS FOLLOWING
INVASIONS OF EXOTIC UNDERSTORY PLANTS IN DECIDUOUS FORESTS %COLOGICAL !PPLICATIONS
%HRENFELD * ' AND . 3COTT )NVASIVE 3PECIES AND THE 3OIL %FFECTS ON /RGANISMS AND
%COSYSTEM 0ROCESSES %COLOGICAL !PPLICATIONS
%VANS 2 $ 2 2IMER , 3PERRY AND * "ELNAP %XOTIC PLANT INVASION ALTERS NITOGEN
DYNAMICS IN AN ARID GRASSLAND %COLOGICAL !PPLICATIONS
&RANCIS 2 AND $ * 2EAD $IRECT TRANSFER OF CARBON BETWEEN PLANTS CONNECTED BY
VESICULAR
ARBUSCULAR MYCORRHIZAL MYCELIUM .ATURE
'ERRISH ' AND $ -UELLER
$OMBIOS "EHAVIOR OF NATIVE AND NON
NATIVE PLANTS IN
TWO TROPICAL RAINFORESTS ON /AHU (AWAIIAN )SLANDS 0HYTOCOENOLOGIA
'RIME * 0 * - , -ACKEY 3 ( (LLIER AND $ * 2EAD &LORISTIC DIVERSITY IN A MODEL
SYSTEM USING EXPERIMENTAL MICROCOSMS .ATURE
(ALVORSON * * ( "OLTON AND * , 3MITH 4HE PATTERN OF SOIL VARIABLES RELATED
TO !RTEMISIA TRIDENTATA IN A BURNED SHRUB
STEPPE SITE 3OIL 3CIENCE 3OCIETY OF !MERICA
*OURNAL
(ARRIS ' ! 3OME COMPETITIVE RELATIONSHIPS BETWEEN !GROPRYON SPICATUM AND
"ROMUS TECTORUM %COLOGICAL -ONOGRAPHS
(ARTNETT $ # " ! $ (ETRICK ' 7 4 7ILSON AND $ * 'IBSON -YCORRHIZAL
INmUENCE ON INTRA
AND INTERSPECIlC NEIGHBOUR INTERACTIONS AMONG CO
OCCURRING PRAIRIE
GRASSES *OURNAL OF %COLOGY
(ETRICK " ! $ ' 7 4 7ILSON AND $ # (ARTNETT 2ELATIONSHIP BETWEEN MYCOR
RHIZAL DEPENDENCE AND COMPETITIVE ABILITY OF TWO TALLGRASS PRAIRIE GRASSES #ANADIAN
*OURNAL OF "OTANY
(OBBIE 3 % %FFECTS OF PLANT SPECIES ON NUTRIENT CYCLING 4RENDS IN %COLOGY AND
%VOLUTION
(OOPER $ 5 $ % "IGNELL AND 6 + "ROWN )NTERACTIONS BETWEEN ABOVEGROUND
AND BELOWGROUND BIODIVERSITY IN TERRESTRIAL ECOSYSTEMS PATTERNS MECHANISMS AND
FEEDBACKS "IO3CIENCE
(UGHES 2 0 - 6ITOUSEK AND 4 4UNISON !LIEN GRASS INVASION AND lRE IN THE SEA
SONAL SUBMONTANE ZONE OF (AWAII %COLOGY
*ACOBS * 3 2 , 3HELEY AND " $ -AXWELL %FFECT OF 3CLEROTINA SCLEROTIORUM ON THE
INTERFERENCE BETWEEN BLUEBUNCH WHEATGRASS !GROPYRON SPICATUM AND SPOTTED KNAP
WEED #ENATUAREA MACULOSA 7EED 4ECHNOLOGY
+EANE 2 - AND - * #RAWLEY %XOTIC PLANT INVASIONS AND THE ENEMY RELEASE
HYPOTHESIS 4RENDS IN %COLOGY AND %VOLUTION
)NVASIVE PLANT
SOIL ECOSYSTEM FEEDBACKS
+IRKPATRICK " , AND & ! "AZZAZ )NmUENCE OF CERTAIN FUNGI ON SEED GERMINATION
AND SEEDLING SURVIVAL OF FOUR COLONIZING ANNUALS *OURNAL OF !PPLIED %COLOGY
+LIRONOMOS * &EEDBACK WITH SOIL BIOTA CONTRIBUTES TO PLANT RARITY AND INVASIVENESS
IN COMMUNITIES .ATURE
+OURTEV 0 3 * ' %HRENFELD AND 7 (UANG %XOTIC SPECIES ALTER MICROBIAL STRUCTURE
AND FUNCTION IN THE SOIL %COLOGY
,EY 2 % AND # - $!NTONIO %XOTIC GRASS INVASION ALTERS POTENTIAL RATES OF . lXA
TION IN (AWAIIAN WOODLANDS /ECOLOGIA
,OWE 0 . 7 + ,AUENROTH AND ) # "URKE %FFECTS OF NITROGEN AVAILABILITY ON COM
PETITION BETWEEN "ROMUS TECTORUM AND "OUTELOUA GRACILIS 0LANT %COLOGY
-ACK 2 . )NVASION OF "ROMUS TECTORUM , INTO WESTERN .ORTH !MERICA AN ECOLOGI
CAL CHRONICLE !GRO %COSYSTEMS
-ACK - # # - $!NTONIO AND 2 % ,EY !LTERATION OF ECOSYSTEM NITROGEN
DYNAMICS BY EXOTIC PLANTS A CASE STUDY OF # GRASSES IN (AWAII %COLOGICAL !PPLICATIONS
-ARLER - * # ! :ABINSKI AND 2 - #ALLAWAY -YCORRHIZAE INDIRECTLY ENHANCE
COMPETITIVE EFFECTS OF AN INVASIVE FORB ON A NATIVE BUNCHGRASS %COLOGY
-ARON * , AND 0 ' #ONNORS ! NATIVE NITROGEN lXING SHRUB FACILITATES WEED
INVASION /ECOLOGIA
-ARON * , AND 2 , *EFFERIES "USH LUPINE MORTALITY ALTERED RESOURCE AVAILABILITY
AND ALTERNATIVE VEGETATION STATES %COLOGY
-ERCK )NDEX (YDROXYQUINOLINE -ERCK AND #O )NC 7HITEHOUSE .* 0AGE
-ILLS + % AND * $ "EVER -AINTENANCE OF DIVERSITY WITHIN PLANT COMMUNITIES SOIL
PATHOGENS AS AGENTS OF NEGATIVE FEEDBACK %COLOGY
-ITCHELL # ' AND ! ' 0OWER 2ELEASE OF INVASIVE PLANTS FROM FUNGAL AND VIRAL
PATHOGENS .ATURE
-OORA - AND - :OBEL %FFECT OF ARBUSCULAR MYCORRHIZA ON INTER AND INTRASPECIlC
COMPETITION OF TWO GRASSLAND SPECIES /ECOLOGIA
.EERGAARD 0 3EED 0ATHOLOGY *OHN 7ILEY .EW 9ORK .EW 9ORK 53! P
.ORTHUP 2 2 2 ! $AHLGREN AND * ' -C#OLL 0OLYPHENOLS AS REGULATORS OF
PLANT LITTER SOIL INTERACTIONS IN NORTHERN #ALIFORNIAS PYGMY FOREST ! POSITIVE FEEDBACK
"IOGEOCHEMISTRY
0ACKER ! AND + #LAY 3OIL PATHOGENS AND SPATIAL PATTERNS OF SEEDLING MORTALITY IN
A TEMPERATE TREE .ATURE
0ACKER ! AND + #LAY 3OIL PATHOGENS AND 0RUNUS SEROTINA SEEDLINGS AND SAPLING
GROWTH NEAR CONSPECIlC TREES %COLOGY
2EINHART + / ! 0ACKER 7 ( 6AN DER 0UTTEN AND + #LAY 0LANTnSOIL BIOTA
INTERACTIONS AND SPATIAL DISTRIBUTION OF BLACK CHERRY IN ITS NATIVE AND INVASIVE RANGES
%COLOGY ,ETTERS
2EINHART + / AND 2 - #ALLAWAY 2EGIONAL EFFECTS OF SOIL BIOTA FACILITATE EXOTIC
!CER INVASIONS IN %UROPE AND .ORTH !MERICA %COLOGICAL !PPLICATIONS
2ICE % , )NHIBITION OF NITROGEN lXING AND NITRIFYING BACTERIA BY SEED PLANTS
%COLOGY
!3 4HORPE AND 2- #ALLAWAY
2ICE + * AND . # %MORY -ANAGING MICROEVOLUTION RESTORATION IN THE FACE OF
GLOBAL CHANGE &RONTIERS IN %COLOGY
2IDENOUR 7 , AND #ALLAWAY 2 - 2OOT HERBIVORES PATHOGENIC FUNGI AND COMPETI
TION BETWEEN #ENTAUREA MACULOSA AND &ESTUCA IDAHOENSIS 0LANT AND 3OIL
2OBINSON $ AND ! &ITTER 4HE MAGNITUDE AND CONTROL OF CARBON TRANSFER BETWEEN
PLANTS LINKED BY A COMMON MYCORRHIZAL NETWORK *OURNAL OF %XPERIMENTAL "OTANY
3AGGAR 3 0 $ -C)NTOSH # " (EDLEY AND ( +NICKER #HANGES IN SOIL MICROBAL
BIOMASS METABOLIC QUOTIENT AND ORGANIC MATTER TURNOVER UNDER (IERACIUM ( PILOSELLA
, "IOLOGICAL &ERTILITY OF 3OILS
3CHIMEL * 0 AND * "ENNETT .ITROGEN MINERALIZATION CHALLENGES OF A CHANGING
PARADIGM %COLOGY
3CHIMEL * 0 2 ' #ATES AND 2 2UESS 4HE ROLE OF BALSAM POPULAR SECONDARY CHEMI
CALS IN CONTROLLING SOIL NUTRIENT DYNAMICS THROUGH SUCCESSION IN THE !LASKAN TAIGA
"IOGEOCHEMISTRY
3CHLESINGER 7 ( AND ! - 0ILMANIS 0LANT
SOIL INTERACTIONS IN DESERTS
"IOGEOCHEMISTRY
3IMARD 3 7 $ ! 0ERRY AND - $ *ONES .ET TRANSFER OF CARBON BETWEEN ECTOMY
CORRHIZAL TREE SPECIES IN THE lELD .ATURE
3IMBERLOFF $ AND " 6ON (OLLE 0OSITIVE INTERACTIONS OF NONINDIGENOUS SPECIES INVA
SIONAL MELTDOWN "IOLOGICAL INVASIONS
3TEVENSON & * AND - ! #OLE #YCLES OF SOIL CARBON NITROGEN PHOSPHORUS SULFUR
MICRONUTRIENTS *OHN 7ILEY AND 3ONS )NC .EW 9ORK .9
3TOCK 7 $ + 4 7IENAND AND ! # "AKER )MPACTS OF INVADING .
lXING !CACIA
SPECIES ON PATTERNS OF NUTRIENT CYCLING IN TWO #APE ECOSYSTEMS EVIDENCE FROM SOIL INCU
BATION STUDIES AND . NATURAL ABUNDANCE VALUES /ECOLOGIA
4HORPE ! 3 6 !RCHER AND 4 ( $ELUCA )N 0RESS 4HE INVASIVE FORB #ENTAUREA MACU
LOSA INCREASES PHOSPHORUS AVAILABILITY IN -ONTANA GRASSLANDS *OURNAL OF !PPLIED 3OIL
%COLOGY
5PADHYAYA * + 2 4URKINGTON AND $ -C)LURIDE 4HE BIOLOGY OF #ANADIAN WEEDS
"ROMUS TECTORUM , #ANADIAN *OURNAL OF 0LANT 3CIENCE
6AN "REEMEN . 3OILS AS BIOTIC CONSTRUCTS FAVOURING NET PRIMARY PRODUCTIVITY
'EODERMA
6AN "REEMEN . AND ! # &INZI 0LANT
SOIL INTERACTIONS ECOLOGICAL ASPECTS AND
EVOLUTIONARY IMPLICATIONS "IOGEOCHEMISTRY
VAN DER (EIJDEN - ' ! * . +LIRONOMOS - 5RSIC 0 -OUTOGLIS 2 3TREITWOLF
%NGEL
4 "OLLER ! 7IEMKEN AND ) 2 3ANDERS -YCORRHIZAL FUNGAL DIVERSITY DETERMINES
PLANT BIODIVERSITY ECOSYSTEM VARIABILITY AND PRODUCTIVITY .ATURE
6AN DER 0UTTEN 7 ( AND " ! - 0ETERS (OW SOIL
BORNE PATHOGENS MAY AFFECT
PLANT COMPETITION %COLOGY
6AN DER 0UTTEN 7 ( # 6AN $IJK AND " ! - 0ETERS 0LANT
SPECIlC SOIL
BORNE
DISEASES CONTRIBUTE TO SUCCESSION IN FOREDUNE VEGETATION .ATURE
6AN DER 0UTTEN 7 ( 0LANT
SOIL FEEDBACK AS A SELECTIVE FORCE 4RENDS IN %COLOGY
AND %VOLUTION
)NVASIVE PLANT
SOIL ECOSYSTEM FEEDBACKS
6ERSFELD $ " AND " 3 VAN 7ILGREN )MPACT OF WOODY ALIENS ON ECOSYSTEM PROP
ERTIES 0AGES
IN ) ! 7 -ACDONALD & * +RUGER AND ! ! &ERRAR EDICTORS
4HE ECOLOGY AND MANAGEMENT OF BIOLOGICAL INVASIONS IN SOUTHERN !FRICA /XFORD
5NIVERSITY 0RESS #APE 4OWN 3OUTH !FRICA
6ITOUSEK 0 - "IOLOGICAL INVASIONS AND ECOSYSTEM PROPERTIES CAN SPECIES MAKE
A DIFFERENCE 0AGES
IN ' ! -OONEY AND * ! $RAKE EDITORS %COLOGY OF
"IOLOGICAL )NVASIONS OF .ORTH !MERICA AND (AWAII 3PRINGER
6ERLAG .EW 9ORK .9
6ITOUSEK 0 - "IOLOGICAL INVASIONS AND ECOSYSTEM PROCESSES TOWARDS AN INTEGRA
TION OF POPULATION BIOLOGY AND ECOSYSTEM STUDIES /IKOS
6ITOUSEK 0 - AND , 2 7ALKER "IOLOGICAL INVASION BY -YRICA FAYA IN (AWAII PLANT
DEMOGRAPHY NITROGEN lXATION ECOSYSTEM EFFECTS %COLOGICAL -ONOGRAPHS
6ITOUSEK 0 - , 2 7ALKER , $ 7HITEAKER $ -UELLER
$OMBOIS AND 0 ! -ATSON
"IOLOGICAL INVASION BY -YRICA FAYA ALTERS ECOSYSTEM DEVELOPMENT IN (AWAII
3CIENCE
6IVANCO * - ( 0 "AIS & 2 3TERMITZ ' # 4HELEN AND 2 - #ALLAWAY
"IOGEOGRAPHICAL VARIATION IN COMMUNITY RESPONSE TO ROOT ALLELECHEMISTRY NOVEL WEAP
ONS AND EXOTIC INVASION %COLOGY ,ETTERS
9ELENIK 3 ' 7 $ 3TOCK AND $ - 2ICHARDSON %COSYSTEM LEVEL IMPACTS OF INVASIVE
!CACIA SALIGNA IN THE 3OUTH !FRICAN FYNBOS 2ESTORATION %COLOGY
9PSILANTIS 7 2ISK OF CHEATGRASS INVASION AFTER lRE IN SELECTED SAGEBRUSH COMMU
NITY TYPES 53 "UREAU OF ,AND -ANAGEMENT
7ARDLE $ ! + 3 .ICHOLSON AND ! 2AHMAN %COLOGICLA EFFECTS OF THE INVASIVE
WEED SPECIES 3ENECIO JACOBAEA , RAGWORT IN A .EW :EALAND PASTURE !GRICULTURE
%COSYSTEMS AND %NVIRONMENT
7ARDLE $ ! AND + 3 .ICHOLSON 3YNERGISTIC EFFECTS OF GRASSLAND PLANT SPECIES
ON SOIL MICROBIAL BIOMASS AND ACTIVITY IMPLICATIONS FOR THE ECOSYSTEM LEVEL EFFECTS OF
ENRICHED PLANT DIVERSITY &UNCTIONAL %COLOGY
7ARDLE $ ! + 3 .ICHOLSON - !HMED AND ! 2AHMAN )NTERFERENCE EFFECTS OF
THE INVASIVE PLANT #ARDUUS NUTANS , AGAINST THE NITROGEN lXATION ABILITY OF 4RIFOLIUM
REPENS, 0LANT AND 3OIL
7ARDLE $ ! - .ILSSON # 'ALLET AND / :ACKISSON !N ECOSYSTEM
LEVEL PERSPECTIVE
OF ALLELOPATHY "IOLOGICAL 2EVIEW
7ATKINS . + ! ( &ITTER * $ 'RAVES #ARBON TRANSFER BETWEEN # AND # PLANTS
LINKED BY A COMMON MYCORRHIZAL NETWORK QUANTIlED USING STABLE CARBON ISOTOPES 3OIL
"IOLOGY AND "IOCHEMISTRY
7ATKINSON ! 2 4HE ROLE OF THE SOIL COMMUNITY IN PLANT POPULATION DYNAMICS
4RENDS IN %COLOGY AND %VOLUTION
7EST ( - )NmUENCE OF ARBUSCULAR MYCORRHIZAL INFECTION ON COMPETITION BETWEEN
(OLCUS LANATUS AND $ACTYLIS GLOMERATA *OURNAL OF %COLOGY
7ESTOVER + - ! # +ENNEDY AND 3 % +ELLEY 0ATTERNS OF RHIZOSPHERE MICROBIAL
COMMUNITY STRUCTURE ASSOCIATED WITH CO
OCCURRING PLANT SPECIES *OURNAL OF %COLOGY
#HAPTER SIXTEEN
)NVASION BIOLOGY AS A
COMMUNITY PROCESS MESSAGES
FROM MICROBIAL MICROCOSMS
0 ( 7ARREN 2 ,AW AND ! * 7EATHERBY
).42/$5#4)/.
)NVASIONS ARE WIDELY REGARDED AS A SIGNIlCANT CURRENT PROBLEM IN POPULATION
ECOSYSTEM AND HABITAT MANAGEMENT WITH CONSEQUENCES INCLUDING GLOBAL HOMOG
ENIZATION OF mORAS AND FAUNAS EXTIRPATION OF NATIVE SPECIES AND INTERFERENCE WITH
ECOSYSTEM FUNCTIONING $RAKE ET AL 7ILLIAMSON -ACK ET AL
0IMENTEL ET AL 3AKAI ET AL 3UCH INVASIONS OCCUR AND HAVE EFFECTS AT
LARGE SCALES IN REAL ECOSYSTEMS 3O WHY CONSIDER INVESTIGATING SUCH PHENOMENA
IN ARTIlCIAL MICROCOSM SYSTEMS IN THE LABORATORY 7E BELIEVE THAT THERE ARE SEV
ERAL COMPELLING REASONS WHY LABORATORY MICROCOSM SYSTEMS HAVE A VALUABLE ROLE
IN THE STUDY OF BIOLOGICAL INVASIONS THESE CENTRE ON THE ISSUES OF MANIPULATION
TIME SCALE AND REPLICATION
&IRST APPLIED ECOLOGY SEEKS A PREDICTIVE UNDERSTANDING OF INVASIONS WHAT
DETERMINES WHICH SPECIES WILL MAKE SUCCESSFUL INVADERS !RE SOME TYPES OF COM
MUNITIES INVASION PRONE (OW WILL AN INVADER IMPACT ON A COMMUNITY 4HERE ARE
VARIOUS THEORETICAL EXPECTATIONS BUT TESTING THESE IN THE lELD IS LARGELY DEPENDENT
ON EXTRACTING STATISTICAL SIGNALS FROM DOCUMENTATION OF THE @NATURAL EXPERIMENTS
THAT REAL INVASIONS PROVIDE /PPORTUNITIES FOR CONTROLLED EXPERIMENTATION ARE
-7 #ADOTTE ET AL EDS #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY n
3PRINGER 0RINTED IN THE .ETHERLANDS
0( 7ARREN 2 ,AW AND !* 7EATHERBY
LIMITED AND IT IS NOTORIOUSLY DIFlCULT TO DOCUMENT
SYSTEMATICALLY
INVASION
ATTEMPTS
THAT FAIL 3IMBERLOFF THE NECESSARY @CONTROL FOR EVALUATION OF THE DETER
MINANTS OF INVASION SUCCESS -ICROCOSMS MESOCOSMS AND RELATED TECHNIQUES
SUCH AS ENCLOSURE EXPERIMENTS PROVIDE THE POSSIBILITY OF MANIPULATING POTENTIAL
INmUENCES ON INVASION SUCCESS ALBEIT AT THE EXPENSE OF REALISM IN COMPLEXITY
AND SCALING
3ECOND INVASIONS TAKE TIME 4HE POPULATION DYNAMICS OF AN INVADING SPECIES
ITS GEOGRAPHIC SPREAD AND THE ADJUSTMENT OF THE RECEIVING COMMUNITIES ARE ALL
TYPICALLY EXPRESSED OVER TENS TO HUNDREDS OF GENERATIONS USUALLY TAKING MANY
YEARS (OWEVER DESIRABLE THE EXPLORATION OF SUCH EFFECTS IN INVASIONS CURRENTLY
HAPPENING IN THE lELD MAY BE IMPRACTICAL -ICROCOSM SYSTEMS UTILIZING MICROOR
GANISMS ALLOW @LONG TERM EFFECTS AND OUTCOMES IN THE RANGE OF TENS TO HUNDREDS
OF GENERATIONS TO BE DOCUMENTED $RAKE ET AL ,AWLER
4HIRD AS THE WEALTH OF CASE STUDIES IN THE INVASIONS LITERATURE TESTIlES INVA
SIONS ARE RATHER INDIVIDUALISTIC EVENTS A PARTICULAR INVADER INVADING A PARTICU
LAR SYSTEM OR REGION UNDER PARTICULAR CIRCUMSTANCES 4HIS MAKES THE APPRAISAL OF
GENERAL PATTERNS AND CAUSES DIFlCULT )N PARTICULAR IF WE ARE INTERESTED IN PREDIC
TION OR AT LEAST PREDICTABILITY THEN IT BECOMES IMPORTANT TO KNOW THE PROBABILITY
OF A PARTICULAR OUTCOME FOR WHAT IS AFTER ALL A PROCESS LIKELY TO BE STRONGLY INmU
ENCED BY STOCHASTIC EVENTS )N EXPERIMENTAL SYSTEMS REPLICATION OF COMMUNITIES
AND INVASION EVENTS MAKES IT POSSIBLE TO ASSESS THE VARIATION IN THE OUTCOME OF
INVASIONS AND THE SOURCE OF THAT VARIATION
! BEAKER IN THE LABORATORY IS FAR FROM A lELD EXPERIMENT IN A NATURAL ECOSYSTEM
3PATIAL AND TEMPORAL SCALES THE TYPES OF ORGANISMS INVOLVED AND THE SIMPLICITY OF
THE COMMUNITIES MEAN THAT MICROCOSMS CANNOT SIMPLY BE TAKEN AS ANALOGUES OF
LARGE NATURAL ECOSYSTEMS THE SAME IS TRUE FOR OTHER SORTS OF MICROCOSM AND MESO
COSM EXPERIMENTS 0ETERSEN AND (ASTINGS ! MICROBIAL MICROCOSM IS BETTER
THOUGHT OF AS A SMALL SYSTEM IN ITS OWN RIGHT AND WITH ITS OWN PECULIARITIES RATHER
THAN AS A LARGE SYSTEM WRIT SMALL )N A SENSE DECIDING THE EXTENT TO WHICH RESULTS
FROM MICROCOSMS CAN BE APPLIED TO OTHER TYPES OF SYSTEM IS EQUIVALENT TO DECID
ING THE EXTENT TO WHICH RESULTS FROM ANY ONE TYPE OF COMMUNITY CAN BE APPLIED
TO OTHER DISPARATE SYSTEMS FOR INSTANCE THE EXTENT TO WHICH RESULTS FROM PHYTO
PLANKTON COMMUNITIES CAN BE APPLIED TO FORESTS -ICROCOSMS OF PROTISTS CERTAINLY
TELL US SOMETHING ABOUT THE BEHAVIOUR OF MICROBIAL COMMUNITIES BUT LEAVE OPEN
OPEN THE QUESTION OF WHETHER SUCH BEHAVIOUR GENERALIZES THE LATTER IS ALSO TRUE FOR
EXPERIMENTS ON FORESTS AND PHYTOPLANKTON
-ICROCOSMS CANNOT OF COURSE REPLACE LARGE SCALE lELD EXPERIMENTS ON SPE
CIlC ECOYSTEMS /NLY THROUGH SUCH EXPERIMENTS CAN SOME OF THE COMPLEXITY AND
SPECIlC FEATURES OF THE SYSTEM OF INTEREST BE CAPTURED (OWEVER LARGE SCALE lELD
EXPERIMENTS ARE OFTEN CONCERNED WITH TESTING GENERAL PRINCIPLES SUCH AS THE ROLES
OF SPECIES RICHNESS OR SPECIES TYPE EG 7ARDLE (ECTOR ET AL AND
THE EFFECTS OF mUCTUATION OR HETEROGENEITY IN THE ABIOTIC ENVIROMENT EG $AVIS
AND 0ELSOR 1UESTIONS ABOUT HOW TO GENERALIZE TO OTHER KINDS OF SYSTEM
APPLY AS MUCH IN THESE lELD EXPERIMENTS AS IN MICROCOSMS -ANY IMPORTANT
)NVASIONS IN MICROCOSMS
ECOLOGICAL PRINCIPLES CAN BE TESTED IN MICROCOSMS AS WELL AS IN lELD EXPERIMENTS EG
0ETCHEY ET AL AND WE TAKE THE VIEW THAT MICROSOSM EXPERIMENTS SHOULD
COMPLEMENT THESE OTHER APPROACHES RATHER THAN BEING ALTERNATIVES TO THEM
!RGUABLY THEREFORE LABORATORY MICROCOSMS HAVE A USEFUL ROLE TO PLAY IN INVESTI
GATING ECOLOGICAL PRINCIPLES OF INVASION JUST AS THEY HAVE PROVIDED VALUABLE INVES
TIGATIVE TOOLS IN OTHER AREAS OF COMMUNITY ECOLOGY ,AWLER )N FACT MANY
OF THE PROCESSES HYPOTHESIZED TO BE DETERMINANTS OF INVASION SUCCESS OR IMPACT
ARE JUST THOSE OF @NATURAL COMMUNITY CHANGE AND REGULATION POPULATION GROWTH
SPECIES INTERACTIONS DISPERSAL TROPHIC POSITION ASSEMBLY ETC $IAMOND AND
#ASE 3HEA AND #HESSON )T IS IN THE STUDY OF THESE GENERAL PROCESSES
RATHER THAN IN THE STUDY OF FACTORS MORE SPECIlC TO THE BIOGEOGRAPHY OF INVASION
FOR EXAMPLE THE LOSS OF COEVOLVED NATURAL ENEMIES 4ORCHIN ET AL -ITCHELL
AND 0OWER THAT LABORATORY MICROCOSMS ARE MOST LIKELY TO GENERATE USEFUL
INSIGHTS
(ERE WE EXAMINE VARIOUS COMPONENTS OF INVASION BIOLOGY AND SOME OF THE
INSIGHTS LABORATORY MICROCOSMS HAVE GIVEN US INTO THESE 7E FOCUS ON MICROCOSMS
OF HETEROTROPHIC MICROBES WHICH WE KNOW BEST FROM OUR OWN WORK AND WHICH
HAVE A LONG HERITAGE IN DIVERSE AREAS OF COMMUNITY ECOLOGY EG SEE REVIEW IN
,AWLER /UR REASON FOR THIS IS THAT SUCH SYSTEMS EXPLOIT ALL THE ADVANTAGES
OF MICROCOSMS LISTED ABOVE BUT IT DOES MEAN THAT WE DO NOT COVER OTHER TYPES OF
EXPERIMENTAL INVASIONS SUCH AS THOSE OF PLANTS INTO COMMUNITIES IN POTS OR SIMI
LAR EXPERIMENTAL SYSTEMS EG $UKES 3TAMPE AND $AEHLER $ICK ET AL
WHICH ARE SOMETIMES TERMED MICROCOSMS VALUABLE THOUGH SUCH EXPERI
MENTS ARE FOR INVESTIGATING THE DETAILS AND MECHANISMS OF PARTICULAR INVASIONS
!. ).6!3)/. %80%2)-%.4
)N DEVELOPING THE IDEAS BELOW WE DRAW SUBSTANTIALLY ON INFORMATION FROM MICRO
COSM EXPERIMENTS ON INVASIBILITY WHICH WE CONDUCTED AS PART OF A STUDY OF COM
MUNITY ASSEMBLY 7EATHERBY ET AL ,AW ET AL 7ARREN ET AL
4HERE ARE A NUMBER OF RESULTS THAT EMERGE FROM THIS WORK WHICH WE FEEL ARE BOTH
PERTINENT TO QUESTIONS ABOUT INVASIONS IN GENERAL AND ALSO ILLUSTRATIVE OF SOME OF
THE AREAS IN WHICH MICROCOSMS SYSTEMS CAN CONTRIBUTE UNIQUELY TO UNDERSTAND
ING INVASION PROCESSES &ULL DETAILS OF THE EXPERIMENTAL METHODS CAN BE FOUND IN
THE PAPERS ABOVE BUT A BRIEF OUTLINE OF THE EXPERIMENTS IS GIVEN HERE FOR CONVE
NIENCE
7E WORKED WITH A SPECIES POOL OF SIX HETEROTROPHIC PROTISTS BACTERIOVORES
4ETRAHYMENA PYRIFORMIS #OLPIDIUM STRIATUM 0ARAMECIUM CAUDATUM AN OMNIVORE
"LEPHARISMA JAPONICUM PREDATORS %UPLOTES PATELLA !MOEBA PROTEUS PLUS A MIXED
BACTERIAL ASSEMBLAGE IN MICROCOSMS CONTAINING M, OF LIQUID MEDIUM )N AN
INITIAL EXPERIMENT 7EATHERBY ET AL ALL POSSIBLE COMBINATIONS OF THE SPECIES
WERE TESTED FOR THEIR ABILITY TO COEXIST OF THE POSSIBLE COMBINATIONS OF SPECIES
TEN FORMED PERSISTENT COMMUNITIES ON A TIME SCALE OF TENS TO HUNDREDS OF
0( 7ARREN 2 ,AW AND !* 7EATHERBY
GENERATIONS 7E THEN TOOK EACH OF THESE PERSISTENT COMMUNITIES AND CHALLENGED
THEM EXPERIMENTALLY WITH EACH NON RESIDENT SPECIES 3IX REPLICATES OF EACH COMBI
NATION OF COMMUNITY AND INTRODUCED SPECIES WERE USED STARTING FROM A PRE
ESTABLISHED RESIDENT COMMUNITY SET UP FOR WEEKS AND CHALLENGING IT WITH AN
INOCULUM OF THE NEW SPECIES CONSISTING OF THIRTY INDIVIDUALS 4HE INITIAL RATE
OF INCREASE OF THE INVADER ITS LONG TERM PERSISTENCE
UP T O WEEKS AFTER
INVASION
AND THE CHANGES IN COMMUNITY COMPOSITION WERE ALL RECORDED
&!#4/23 !&&%#4).' %34!",)3(-%.4 /& ).6!$%23
%STABLISHMENT AS A PROBABILISTIC EVENT
#HANCE PLAYS AN IMPORTANT PART IN WHETHER A NEW SPECIES GETS ESTABLISHED LOCALLY
#RAWLEY %VEN IF A NEW SPECIES HAS VITAL RATES THAT ENABLE IT TO INCREASE ON
THE AVERAGE RANDOM EVENTS CAN CATCH UP WITH IT BEFORE IT IS ABLE TO GET STARTED
3UCH EVENTS INCLUDE DEMOGRAPHIC STOCHASTICITY ENVIRONMENTAL STOCHASTICITY AND
THE EXACT STATE OF THE RESIDENT COMMUNITY AT THE TIME WHEN THE NEW SPECIES IS
INTRODUCED
)T THEREFORE HELPS TO THINK OF ESTABLISHMENT AS A PROBABILISTIC EVENT 4HEORY FROM
STOCHASTIC BIRTH
DEATH PROCESSES PROVIDES A QUANTITATIVE FRAMEWORK ON WHICH TO
BUILD IDEAS AT LEAST SO FAR AS DEMOGRAPHIC STOCHASTICITY IS CONCERNED "AILEY
'OEL AND 2ICHTER
$YN 4HIS THEORY CONSIDERS A POPULATION IN
WHICH INDIVIDUALS HAVE A CONSTANT PROBABILITY PER UNIT TIME OF GIVING BIRTH B AND
OF DYING D WITH AN ASSUMPTION THAT B D )F THE POPULATION STARTS WITH AN INOCU
LUM OF N INDEPENDENT INDIVIDUALS IT HAS A PROBABILITY
n DB N
0N .
n DB .
OF REACHING SIZE . N BEFORE IT GOES TO EXTINCTION &OR A SPECIES INTRODUCED
INTO A WELL
MIXED COMMUNITY CLOSE TO STEADY STATE AND AT AN ABUNDANCE MUCH
LOWER THAN THE RESIDENT SPECIES B AND D MAY BE TAKEN AS APPROXIMATELY CONSTANT
DURING THE EARLY STAGES OF POPULATION GROWTH BECAUSE MOST INTERACTIONS ARE WITH
THE RESIDENT SPECIES . CAN BE SET AT A VALUE ABOVE THE LEVEL AT WHICH DEMOGRAPHIC
STOCHASTICITY IS IMPORTANT TO GIVE AN EXPRESSION FOR THE PROBABILITY THAT A NEW
SPECIES BECOMES SUCCESSFULLY ESTABLISHED
0N n DB N n n RB N
WHERE R B n D IS THE INITIAL PER CAPITA RATE OF INCREASE OF THE NEW SPECIES AND
B D 0N IF B D 0LOTTING THIS FUNCTION SHOWS AS ONE WOULD EXPECT
THE SPECIES IS MORE LIKELY TO BECOME ESTABLISHED THE GREATER THE INOCULUM SIZE
)NVASIONS IN MICROCOSMS
N AND THE GREATER THE PER CAPITA RATE OF INCREASE R RELATIVE TO THE BIRTH RATE B
OR EQUIVALENTLY THE SMALLER THE DEATH RATE D RELATIVE TO THE BIRTH RATE B &IG
SEE #HAPTER BY &RECKLETON ET AL FOR A BROADER DISCUSSION OF THE ROLE OF STOCHASTIC
ITY IN INVASIONS
&IG 0ROBABILITY 0N THAT AN INTRODUCED ASEXUAL SPECIES REACHES A SIZE ABOVE A LEVEL
AT WHICH DEMOGRAPHIC STOCHASTICITY IS IMPORTANT STARTING WITH N PROPAGULES AND HAVING
CONSTANT PER CAPITA BIRTH B AND DEATH D RATES WITH R B n D 4HE SURFACE IS CONSTRUCTED FROM
%QUATION
!LTHOUGH THE ROLE THAT CHANCE PLAYS IN ESTABLISHMENT OF NEW SPECIES IS RARELY
A SUBJECT OF DIRECT EXPERIMENTATION WE FOUND IT PLAYED AN IMPORTANT PART IN THE
REPLICATE INVASION TRIALS WE CARRIED OUT DURING OUR WORK ON COMMUNITY ASSEMBLY
OF PROTISTS ,AW ET AL )N THIS WORK WE TOOK A SPECIES TO HAVE BECOME ESTAB
LISHED IF IT REACHED A THRESHOLD OF INDIVIDUALS AT SOME STAGE DURING THE COURSE OF
THE EXPERIMENT (OWEVER IN ABOUT OF THE RESIDENT
INTRODUCTION TREATMENTS
THE OUTCOME REMAINED UNCERTAIN IN THE SENSE THAT THE INTRODUCED SPECIES BECAME
0( 7ARREN 2 ,AW AND !* 7EATHERBY
ESTABLISHED IN SOME REPLICATES WHILE DISAPPEARING IN OTHERS &IG $EMOGRAPHIC
STOCHASTICITY MOST LIKELY PLAYED A PART IN THIS ALTHOUGH THIS DOES NOT PRECLUDE
INVOLVEMENT OF OTHER FACTORS AS WELL &OR INSTANCE SPECIES THAT REMAIN AT LOW
POPULATION SIZES FOR LONG PERIODS OF TIME ARE ALSO VULNERABLE TO THE CONTINUING
EFFECTS OF ENVIRONMENTAL STOCHASTICITY AS IN TWO OF OUR MICROCOSMS WHERE NEITHER
EXTINCTION NOR THE THRESHOLD FOR ESTABLISHMENT HAD BEEN REACHED BY THE TIME
THE EXPERIMENT ENDED WEEKS AFTER INOCULATION !LSO THE EXACT ABUNDANCE OF
THE RESIDENT SPECIES AT THE TIME OF INTRODUCTION WHICH WE COULD NOT CONTROL FOR
IN OUR INVASION TRIALS COULD INmUENCE THE OUTCOME 3UCH EFFECTS OF ABUNDANCE
ARE SUGGESTED BY THE FACT THAT THE OUTCOME OF INVASIONS WAS NEVER IN DOUBT IN
OUR MICROCOSMS LACKING RESIDENT PROTISTS BUT WAS UNCERTAIN IN ABOUT A THIRD OF
THE TREATMENTS CONTAINING RESIDENT PROTISTS $RAKE IN MICROCOSMS OF
ALGAE AND GRAZING MICROCRUSTACEA SUGGESTED A POSSIBLY SIMILAR EFFECT WHERE
SMALL VARIATIONS IN INITIAL DEMOGRAPHY OF INTRODUCED GRAZERS AMONG REPLICATES
CAUSED MARKED VARIATION IN GRAZER POPULATION SUCCESS IN SOME COMMUNITIES BUT
NOT IN OTHERS
&IG 5NCERTAINTY IN THE OUTCOME OF RESIDENT
INTRODUCTION TREATMENTS IN A PROTIST
INVASION EXPERIMENT DESCRIBED IN ,AW ET AL 4REATMENTS ARE SHOWN IN THREE
CATEGORIES ACCORDING TO WHETHER THE INTRODUCED SPECIES FAILED TO ESTABLISHPERSIST IN ALL
REPLICATES SUCCEEDED IN ESTABLISHINGPERSISTING IN ALL REPLICATES OR SUCCESSFULLY INVADED
SOME BUT FAILED IN OTHERS MIXED "ARS SHOW THE NUMBER OF TREATMENTS FALLING INTO EACH
CATEGORY 3HADED BARS ESTABLISHMENT REACHING A THRESHOLD OF INDIVIDUALS AT SOME POINT
IN THE EXPERIMENT OPEN BARS PERSISTENCE STILL PRESENT IN THE COMMUNITY AFTER WEEKS
-OST TREATMENTS HAD REPLICATES THOUGH A FEW ONLY HAD OR
)NVASIONS IN MICROCOSMS
0ROPAGULE PRESSURE AND THE PROBABILITY OF ESTABLISHMENT
4HERE IS A GENERAL UNDERSTANDING THAT ESTABLISHMENT OF A NEW SPECIES IS MORE
LIKELY THE MORE INDIVIDUALS INTRODUCED AT A TIME 7ILLIAMSON +OLA AND
,ODGE )T IS INTUITIVE FOR INSTANCE THAT DEMOGRAPHIC STOCHASTICITY SHOULD
HAVE ITS GREATEST EFFECT ON ESTABLISHMENT OF A SPECIES WHEN JUST A FEW INDIVIDUALS
ARE INTRODUCED BECAUSE A SEQUENCE OF RELATIVELY FEW DEATHS CAN TAKE THE NEW POPU
LATION TO EXTINCTION !LTHOUGH RATHER LITTLE IS KNOWN ABOUT PROPAGULE PRESSURE
THIS MAY WELL CONTRIBUTE IN IMPORTANT WAYS TO PATTERNS OF INVASIONS OBSERVED IN
NATURE 7ILLIAMSON ET SEQ ,ONSDALE !MONG THE MOST SYSTEMATIC
INVESTIGATIONS HAVE BEEN THE RELEASE OF BIOLOGICAL CONTROL AGENTS EG (OPPER
AND 2OUSH WHERE THERE IS PARTICULAR CONCERN TO ENSURE THE CONTROL AGENTS
BECOME SUCCESSFULLY ESTABLISHED 3HEA AND 0OSSINGHAM %QUATION ABOVE
PROVIDES A CLEAR QUANTITATIVE PREDICTION ABOUT THE EFFECT OF INOCULUM SIZE N ON THE
PROBABILITY OF SUCCESSFUL ESTABLISHMENT
4HE QUANTITATIVE DEPENDENCE OF INVASION SUCCESS ON PROPAGULE PRESSURE HAS
RECEIVED LITTLE SYSTEMATIC EXPERIMENTAL INVESTIGATION 7ELL
MIXED MICROCOSMS
CONTAINING SPECIES WITH SHORT GENERATION TIMES WOULD BE A GOOD POINT AT WHICH
TO BEGIN EXAMINING THIS RELATIONSHIP
)NITIAL RATE OF INCREASE AND THE PROBABLITY OF ESTABLISHMENT
/NE WOULD EXPECT THAT THE LARGER THE PER CAPITA RATE OF INCREASE OF A RARE INTRO
DUCED SPECIES THE GREATER THE LIKELIHOOD THAT IT BECOMES ESTABLISHED !PART FROM
HUMAN PATHOGENS EG !NDERSON AND -AY RATHER LITTLE INFORMA
TION IS AVAILABLE ON THIS INITIAL RATE OF INCREASE 9ET IT IS INTUITIVE THAT A HIGH RATE
OF INCREASE SHOULD ALLOW EARLY ESCAPE FROM THE REGION IN WHICH DEMOGRAPHIC
STOCHASTICITY COULD DRIVE THE INTRODUCED SPECIES TO EXTINCTION %QUATION MAKES
THIS INTUITION PRECISE SHOWING THAT WHAT ACTUALLY MATTERS IS THE RATIO OF THE DEATH
RATE TO BIRTH RATE THE RATIO LEADING TO A DIMENSIONLESS EXPRESSION INDEPENDENT OF
THE TIME SCALE ON WHICH THE DYNAMICS ARE OPERATING
-ICROCOSMS BECAUSE OF THEIR POTENTIAL FOR HIGH LEVELS OF REPLICATION PROVIDE
SCOPE FOR STUDYING INITIAL RATES OF INCREASE AND THE PROBABILITY OF ESTABLISHMENT
/UR STUDIES OF PROTIST MICROCOSMS ,AW ET AL ILLUSTRATE THE POTENTIAL TO
ESCAPE FROM STOCHASTIC EXTINCTION THROUGH RAPID POPULATION GROWTH &IG
4HE GRAPHS SHOW THE GROWTH OF "LEPHARISMA POPULATIONS EACH STARTING WITH AN
INOCULUM OF INDIVIDUALS INTRODUCED INTO FOUR DIFFERENT TYPES OF MICROCOSM
THOSE CONTAINING 0ARAMECIUM #OLPIDIUM OR 4ETRAHYMENA AND THOSE WITH NO OTHER
PROTISTS 7HEN FACED WITH 0ARAMECIUM MOST "LEPHARISMA POPULATIONS DECLINED
RAPIDLY IN FACT TWO WERE EXTINCT BY DAY SIX 7ITH #OLPIDUM "LEPHARISMA TEETERED
ALONG AT SMALL POPULATION SIZES DURING THIS TIME THERE WOULD BE A CONTINUING
RISK OF EXTINCTION )N THE PRESENCE OF 4ETRAHYMENA "LEPHARISMA POPULATIONS GREW
MUCH FASTER AND SOON GOT BEYOND A SIZE AT WHICH EXTINCTION THROUGH DEMOGRAPHIC
0( 7ARREN 2 ,AW AND !* 7EATHERBY
&IG 4IME SERIES OF THE EARLY GROWTH OF POPULATIONS OF "LEPHARISMA INTRODUCED INTO
ESTABLISHED POPULATIONS OF A 0ARAMECIUM B #OLPIDIUM C 4ETRAHYMENA D NO OTHER
PROTISTS TAKEN FROM AN EXPERIMENT DESCRIBED IN ,AW ET AL )N EACH CASE SIX
INDEPENDENT REPLICATE INTRODUCTIONS OF INDIVIDUALS ARE SHOWN
STOCHASTICITY COULD BE AN ISSUE 4HE BEHAVIOUR OF "LEPHARISMA IN THE ABSENCE OF
ANY PROTISTS WAS SIMILAR ALTHOUGH THE GROWTH RATE WAS SOMEWHAT SMALLER
)NVASION BIOLOGY WOULD BE RELATIVELY STRAIGHTFORWARD IF ECOLOGISTS COULD POINT TO
BIOLOGICAL PROPERTIES OF SPECIES THAT MAKE THEIR INITIAL RATES OF INCREASE EITHER POSI
TIVE OR NEGATIVE THOUGH SUCCESS IN DOING THIS HAS GENERALLY BEEN MIXED 7ILLIAMSON
)NVASIONS IN MICROCOSMS
-ACK +OLA AND ,ODGE 4HE RESULTS ON "LEPHARISMA ABOVE ARE
ALSO NOT ENCOURAGING IN THIS REGARD BECAUSE SUCCESS WAS CLEARLY CONTINGENT ON
PROPERTIES OF THE RESIDENT COMMUNITY .ONETHELESS OUR STUDIES DID SHOW SOME
LARGE D IFFERENCES AMONG INTRODUCED SPECIES IN THE INITIAL RATES OF INCREASE ,AW
ET AL )T WAS STRIKING THAT THE BACTERIOVORE 0ARAMECIUM WAS CONSTITUTIVELY A
STRONG INVADER WHATEVER RESIDENT COMMUNITY IT ENCOUNTERED 4HERE ARE EVIDENTLY
FEATURES OF 0ARAMECIUM THAT PREDISPOSE IT TO GROW ESPECIALLY WELL IN MICROCOSMS
EVEN WHEN IT SEEMS LIKELY TO BE IN COMPETITION FOR RESOURCES WITH OTHER SPECIES
2ESIDENT COMMUNITY AND THE PROBABILITY OF ESTABLISHMENT
!LTHOUGH CERTAIN SPECIES MIGHT BE CONSTITUTIVELY GOOD OR BAD INVADERS IN GENERAL
IT IS MORE REALISTIC TO ENVISAGE INVASION AS A JOINT PROPERTY OF AN INTRODUCED SPECIES
AND THE SPECIES ALREADY RESIDENT IN THE COMMUNITY CONTINGENT ON THE INTERACTIONS
BETWEEN THE NEW SPECIES AND THE RESIDENTS 4HE PER CAPITA BIRTH AND DEATH RATES
IN %QUATION NEED TO BE THOUGHT OF AS BEING DETERMINED AS MUCH BY THE RESIDENT
COMMUNITY AT THE TIME OF INTRODUCTION AS BY PROPERTIES OF THE INTRODUCED SPECIES
4HE IMPORTANCE OF RESIDENT AND INTRODUCED SPECIES AS CO
DETERMINANTS OF INVA
SIONS WAS CLEAR IN OUR EXPERIMENTAL INTRODUCTIONS OF PROTISTS INTO DIFFERENT COM
MUNITIES ,AW ET AL 7E USED ANALYSIS OF VARIANCE TO PARTITION VARIATION
IN INITIAL RATE OF INCREASE BETWEEN INTRODUCED SPECIES RESIDENT COMMUNITY AND
INTRODUCTION = COMMUNITY INTERACTION !S ONE MIGHT EXPECT THE ANALYSIS SHOWED
A STRONG INTRODUCTION = COMMUNITY INTERACTION 4HE VARIATION IN INITIAL GROWTH
RATES OF "LEPHARISMA &IG AND SEE DISCUSSION ABOVE ILLUSTRATES THE EFFECT OF
DIFFERENCES IN RESIDENT SPECIES "LEPHARISMA AS AN OMNIVORE IS ABLE TO EAT SMALL
PROTISTS AND BACTERIA )N A COMMUNITY WITH 0ARAMECIUM ANOTHER LARGE PROTIST
WITH WHICH IT CAN ONLY COMPETE FOR BACTERIA "LEPHARISMA USUALLY DECLINED RAPIDLY
&IG A 7HEN WITH #OLPIDIUM WHICH IS SOMEWHAT SMALLER AND ALSO FEEDS ON BAC
TERIA "LEPHARISMA WAS JUST ABOUT ABLE TO HOLD ITS OWN &IG B )N THE PRESENCE OF
4ETRAHYMENA A SMALL BACTERIOVORE WHICH "LEPHARISMA CAN EAT AS WELL AS BACTERIA
"LEPHARISMA POPULATIONS GREW RAPIDLY &IG C FASTER IN FACT THAN WHEN ALL COM
PETITION FOR BACTERIA WAS REMOVED &IG D
!LTHOUGH INVASION MAY DEPEND ON INTERACTIONS AMONG SPECIES COULD COMMU
NITIES HAVE GENERAL PROPERTIES THAT RENDER THEM MORE OR LESS LIABLE TO INVASION
/NE LONG
STANDING PREDICTION IS THAT COMMUNITIES WITH A LARGE NUMBER OF SPE
CIES SHOULD BE LESS READILY INVADED THAN THOSE WITH FEW SPECIES %LTON
,EVINE AND $!NTONIO THIS IS ON THE GROUNDS THAT THE AVAILABLE NICHE SPACE
IS MORE FULLY OCCUPIED !MONG THE FEW MICROCOSM EXPERIMENTS THAT HAVE INVES
TIGATED THE RELATIONSHIP BETWEEN SPECIES RICHNESS AND INVASIBILITY ARE THE STUDIES
BY -C'RADY
3TEED ET AL AND 2OBINSON AND $ICKERSON 4HE FORMER
STUDY FOUND THAT THE PROTIST %UPLOTES WAS ABLE TO INVADE SOME NOT ALL MICROCOSMS
OF LOW SPECIES RICHNESS AND ABLE TO INVADE NO COMMUNITIES OF HIGH RICHNESS
4HE LATTER USING THREE DIFFERENT SPECIES OF PROTISTS AS INVADERS APPEARED TO SUG
GEST THAT INVASION SUCCESS MEASURED AS PRESENCE OF THE INVADER AFTER THREE WEEKS
0( 7ARREN 2 ,AW AND !* 7EATHERBY
WAS NOT STRONGLY INmUENCED BY COMMUNITY DIVERSITY THOUGH A LATER REANALYSIS
,EVINE AND $!NTONIO FOUND EVIDENCE FOR A NEGATIVE EFFECT OF DIVERSITY ON
SUCCESS (OWEVER EXPERIMENTS OF THIS KIND HAVE THE PROPERTY THAT RICHER COMMU
NITIES CONTAIN MORE DIFFERENT SPECIES (USTON AND ARE THEREFORE MORE LIKELY
TO CONTAIN PARTICULAR SPECIES THAT DISCOURAGE OR POTENTIALLY PROMOTE INVASION
BY PARTICULAR NEW SPECIES MAKING IT DIFlCULT TO ATTRIBUTE INVASION OUTCOMES TO
EFFECTS OF DIVERSITY PER SE 7ARDLE (ODGSON ET AL LOOKING AT THE
ABILITY OF PARTICULAR STRAINS OF THE BACTERIUM 0SEUDOMONAS mUORESCENS TO INCREASE
WHEN RARE IN THE PRESENCE OF OTHER STRAINS FOUND AN EFFECT OF DIVERSITY ON THIS
MEASURE OF INVASIBILITY BUT THAT IT WAS MUCH WEAKENED BY STATISTICAL REMOVAL OF
THIS SAMPLING EFFECT
.UMERICAL STUDIES OF COMMUNITY MODELS SUGGEST THAT COMMUNITIES BECOME
INCREASINGLY RESISTANT TO INVASION AS TIME GOES ON AS A RESULT OF THE TURNOVER OF
SPECIES 0OST AND 0IMM $RAKE B ,AW AND -ORTON )NCREASING
INVASION RESISTANCE CAN APPLY EVEN IF THE RESIDENT COMMUNITY IS NOT ACCUMULAT
ING SPECIES AND SUGGESTS THAT IT SHOULD BE RELATIVELY HARD FOR NEW SPECIES TO INVADE
COMMUNITIES WITH LONG HISTORIES OF ASSEMBLY !LSO ONE MIGHT EXPECT COMMUNITIES
CONSTRUCTED FROM LARGE SPECIES POOLS TO BE LESS READILY INVADED BECAUSE WITH
MORE COMBINATIONS OF SPECIES TESTED DURING ASSEMBLY A lNER DEGREE OF TUNING TO
INVASION
RESISTANT STATES BECOMES POSSIBLE 4O DATE WE KNOW OF ONLY ONE MICROCOSM
STUDY OF THE EFFECT OF COMMUNITY AGE ON INVASIBILITY ,ONG ET AL UNPUBLISHED MS
IN THIS CASE OLDER COMMUNITIES WERE MORE RESISTANT TO TWO OUT OF THREE SPECIES
INTRODUCED THAN THE YOUNGER COMMUNITIES (OWEVER THE OLDER COMMUNITIES
WERE CREATED BY ALLOWING A LONGER PERIOD OF TIME TO ELAPSE FROM A lXED INITIAL SET
OF SPECIES SO INVASION RESISTANCE HERE MAY BE BETTER INTERPRETED AS A CONSEQUENCE
OF BEING FURTHER ALONG A TRANSIENT OF COMMUNITY DYNAMICS RATHER THAN AS A CON
SEQUENCE OF TURNOVER OF SPECIES
%NVIRONMENTAL FACTORS AND THE PROBABILITY OF ESTABLISHMENT
4HE EXTERNAL ENVIRONMENT OF THE COMMUNITY IS A FURTHER IMPORTANT DETERMINANT
OF WHETHER A NEW SPECIES BECOMES ESTABLISHED )F RESISTANCE TO INVASION IS A CON
SEQUENCE OF INTENSE INTERACTIONS BETWEEN NEW SPECIES AND RESIDENTS THEN FACTORS
THAT DISRUPT THOSE INTERACTIONS MIGHT BE EXPECTED TO PROMOTE INVASION )N A RELATED
VEIN IF THERE ARE ENVIRONMENTAL CONDITIONS THAT ENHANCE GROWTH OR PERSISTENCE OF
SPECIES IN A COMMUNITY GENERALLY AND HENCE DIVERSITY THEN SUCH CONDITIONS MAY
ALSO INCREASE THE LIKELIHOOD OF SUCCESS OF AN INTRODUCED SPECIES WHEN THE SUN
SHINES IT SHINES ON THE JUST AND THE UNJUST ALIKE ,EVINE 3HEA AND #HESSON
"YERS AND .OONBURG "OTH THESE POSSIBILITIES ARGUE FOR A SIGNIlCANT
ROLE OF ENVIRONMENTAL VARIATION EITHER OVER TIME OR BETWEEN COMMUNITIES IN
DETERMINING THE PROBABILITY OF ESTABLISHMENT OF A NEW SPECIES
4HE ENVIRONMENTAL FACTOR THAT HAS RECEIVED MOST ATTENTION IN THIS RESPECT IS
DISTURBANCE THE ESSENTIAL CONCLUSION BEING THAT AT LEAST ON SOME TIMESCALES
DISTURBANCE ENHANCES INVASION PRESUMABLY BY REDUCING THE INTENSITY OF BIOTIC
)NVASIONS IN MICROCOSMS
INTERACTIONS #RAWLEY 2EJMANEK (OBBS 4HOMPSON ET AL
3HER AND (YATT $AVIS ET AL 3HEA AND #HESSON !LTHOUGH
MICROCOSMS WOULD SEEM AN OBVIOUS SYSTEM IN WHICH TO EXAMINE THIS EFFECT SYS
TEMATICALLY WE KNOW OF ONLY ONE SUCH EXPERIMENT ,ONG ET AL UNPUBLISHED MS
A STUDY USING HETEROTROPHIC PROTISTS WITH DISTURBANCE GENERATED BY DENSITY
INDE
PENDENT MORTALITY OF ALL RESIDENT SPECIES )N THIS CASE DISTURBANCE AFFECTED THE
INTRODUCED SPECIES "LEPHARISMA 0ARAMECIUM AND #OLPIDIUM IN QUITE DIFFERENT WAYS
THE MAIN EFFECTS OF DISTURBANCE WERE TO ENHANCE THE ABUNDANCE OF "LEPHARISMA TO
LEAVE 0ARAMECIUM UNCHANGED AND TO REDUCE THE ABUNDANCE OF #OLPIDIUM
! SECOND MICROCOSM STUDY *IANG AND -ORIN WITH PROTISTS AND ROTIFERS
PROVIDES ONE OF THE lRST EXPERIMENTAL TESTS OF THE EFFECT OF ANOTHER ENVIRONMENTAL
FACTOR ENERGY AVAILABILITY ON INVASION )N THIS CASE INCREASED ENERGY AVAILABILITY
PROMOTED INITIAL POPULATION GROWTH OF TWO DIFFERENT INTRODUCED SPECIES SUPPORT
ING THE POSSIBILITY PREVIOUSLY SPECULATED ABOUT THEORETICALLY 3HEA AND #HESSON
"YERS AND .OONBURG THAT SUCCESSFUL ESTABLISHMENT AND RESIDENT
DIVERSITY MIGHT BE POSITIVELY CORRELATED THROUGH THE ACTION OF A COMMON FAC
TOR (OWEVER &UKAMI AND -ORIN ALSO SHOW USING MICROCOSM SYSTEMS
THAT THE ENERGY
DIVERSITY RELATIONSHIPS IN COMMUNITIES ASSEMBLED BY SEQUENTIAL
INVASIONS ARE AFFECTED BY THE ORDER IN WHICH INVASIONS OCCUR SUGGESTING THAT THE
RELATIONSHIP BETWEEN ENERGY AND INVASION SUCCESS MAY NOT ALWAYS BE A SIMPLE ONE
#(!.'%3 4/ #/--5.)4)%3 &/,,/7).' %34!",)3(-%.4 /& ).6!$%23
&OLLOWING SUCCESSFUL ESTABLISHMENT AN INVADING SPECIES TYPICALLY BUILDS UP TO
A SUBSTANTIAL POPULATION SIZE TAKING THE COMMUNITY BEYOND THE POINT AT WHICH
%QUATION CAN POSSIBLY BE A GOOD APPROXIMATION )N EFFECT THE INVADING SPECIES
BECOMES ABUNDANT ENOUGH TO CHANGE THE BIRTH AND DEATH RATES OF OTHER SPECIES
IN THE COMMUNITY POTENTIALLY CAUSING CHANGE IN THEIR DENSITIES 4HE CHANGING
DENSITY OF OTHER SPECIES FEEDS BACK TO THE INVADER POTENTIALLY CAUSING CHANGES
IN ITS OWN BIRTH AND DEATH RATES !T THIS STAGE THERE IS NO ALTERNATIVE TO DEALING
WITH THE FULL DYNAMICS OF THE COMMUNITY AUGMENTED BY THE INVADER AND ALL THE
COMPLEXITIES THAT FOLLOW FROM THIS
$ESPITE THE GREAT COMPLEXITIES OF COMMUNITY DYNAMICS THERE ARE SOME QUALITA
TIVE QUESTIONS ABOUT THE INVADER AND THE COMMUNITY IT ENTERS THAT CAN BE ADDRESSED
AND AGAIN WE DRAW ON EXAMPLES FROM OUR EXPERIMENTS WITH PROTISTS TO DO THIS
$O INVADERS PERSIST IN THE LONG TERM
! PRIORI THERE IS NO REASON TO EXPECT SPECIES THAT BECOME ESTABLISHED IN THE SHORT
TERM TO PERSIST IN THE LONG TERM )T IS QUITE POSSIBLE FOR AN INVADER TO CHANGE THE
COMMUNITY IN SOME WAY WHICH IS DELETERIOUS TO ITSELF FOR INSTANCE BY DRIVING ITS
PREY TO EXTINCTION (OWEVER IN ANALYSES OF COMMUNITIES WITH ,OTKA
6OLTERRA
DYNAMICS SPECIES ABLE TO INCREASE FROM INITIAL LOW NUMBERS WERE ALSO PRESENT IN
0( 7ARREN 2 ,AW AND !* 7EATHERBY
&IG %XAMPLES OF INITIAL GROWTH OF INTRODUCED PROTISTS AND THE COMMUNITIES THAT
PERSISTED IN THE LONG TERM WEEKS FROM EXPERIMENTS DESCRIBED IN ,AW ET AL AND
7ARREN ET AL ,INES SHOW THE ABUNDANCES OF THE INTRODUCED SPECIES FOR A PERIOD
DETERMINED BY THE POPULATION EITHER SHOWING CONSISTENT POSITIVE GROWTH AND ACHIEVING
NUMBERS GREATER THAN THE INITIAL INOCULUM GOING EXTINCT OR IF NEITHER CONDITION WAS
SATISlED TO AN ARBITRARY END POINT DAYS 4HE LETTER CODES USED REPRESENT THE SPECIES
)NVASIONS IN MICROCOSMS
THE LONG TERM ,AW AND -ORTON 4HESE STUDIES USED AN ASYMPTOTIC CRITE
RION CALLED PERMANENCE TO WORK OUT THE SPECIES COMPOSITION OF NEW COMMUNITIES
THEREBY JUMPING OVER ANY EFFECTS THAT TRANSIENT DYNAMICS MIGHT HAVE ON PERSIS
TENCE OF SPECIES 2EAL COMMUNITIES HAVE TO PASS ALONG THE TRANSIENTS AND IT IS OF
COURSE POSSIBLE FOR THESE TRANSIENTS TO BRING THE INVADER TO A POPULATION SIZE LOW
ENOUGH FOR EXTINCTION BY DEMOGRAPHIC STOCHASTICITY TO BECOME AN ISSUE AGAIN
1UESTIONS OF LONG
TERM PERSISTENCE OF INVADERS CAN BE READILY ADDRESSED IN
MICROCOSMS WHERE THE COMPONENT ORGANISMS HAVE SHORT GENERATION TIMES (ERE
WE LOOK AT TWO ASPECTS OF THIS lRST THE VARIATION IN LONG TERM PERSISTENCE AMONG
REPLICATE INTRODUCTIONS AND SECOND THE CORRESPONDENCE BETWEEN INITIAL ESTAB
LISHMENT AND LONG TERM PERSISTENCE OF DIFFERENT COMBINATIONS OF RESIDENTS AND
INTRODUCED SPECIES
)N OUR EXPERIMENTS THE LONG
TERM FATE OF SPECIES INTRODUCED TO COMMUNITIES
WAS FAR FROM UNIFORM ACROSS THE REPLICATES WITHIN RESIDENT
INTRODUCTION TREAT
MENTS )N ABOUT TWO
THIRDS OF THE FORTY
NINE RESIDENT
INTRODUCTION TREATMENTS
ALL INVASIONS EITHER FAILED OR SUCCEEDED IN THE REMAINDER THE RESULTS WERE MIXED
&IG )N OTHER WORDS ALTHOUGH THE COMMUNITIES CONTAINED THE SAME SPECIES
WERE RUN UNDER THE SAME CONDITIONS AND WERE INOCULATED AT THE SAME TIME WITH
THE SAME NUMBER OF INDIVIDUALS THERE WAS STILL A SUBSTANTIAL ELEMENT OF INDE
TERMINACY ABOUT THE EVENTUAL FATE OF THE INTRODUCED SPECIES !S ALREADY NOTED
3ECTION ONE EXPLANATION IS THE SUCCESS OR FAILURE OF INITIAL ESTABLISHMENT
AS ILLUSTRATED BY THE INVASION OF "LEPHARISMA INTO SYSTEMS CONTAINING 0ARAMECIUM
EG &IG A &IG A AN INTERESTING CASE IN WHICH THE INVADER ESTABLISHED IN
JUST ONE REPLICATE BUT IN THAT SYSTEM THEN PERSISTED IN THE LONG TERM (OWEVER IT
IS NOTABLE THAT INITIAL ESTABLISHMENT AND LONG TERM PERSISTENCE WERE BY NO MEANS
EQUIVALENT &IG 4HIS IS ILLUSTRATED BY !MOEBA INTRODUCED INTO A COMMUNITY OF
0ARAMECIUM #OLPIDIUM AND 4ETRAHYMENA &IG B IT ALWAYS ESTABLISHED ITSELF AT
THE START BUT DID NOT PERSIST IN ALL REPLICATES IN THE LONG RUN )N PASSING IT IS ALSO
INTERESTING TO NOTE THAT EVEN WHERE LONG TERM PERSISTENCE OF THE INVADER OCCURS
IN ALL REPLICATES THERE COULD BE CONSIDERABLE VARIATION IN THE TIMING AND PATTERN
OF ESTABLISHMENT &IG C AGAIN MAKING PREDICTION FROM INITIAL ESTABLISHMENT TO
lNAL OUTCOME A DIFlCULT TASK
)N OUR EXPERIMENTAL INVASIONS THE MISMATCH BETWEEN THE CONSISTENCY OF ESTAB
LISHMENT AND PERSISTENCE IN INDIVIDUAL REPLICATE INTRODUCTIONS &IG WAS LARGELY
! !MOEBA " "LEPHARISMA # #OLPIDIUM % %UPLOTES 0 0ARAMECIUM 4 4ETRAHYMENA
4HE LETTERS IN THE TOP LEFT HAND CORNER OF THE GRAPH INDICATE THE RESIDENT COMMUNITY
BEFORE INTRODUCTION THE VERTICAL DASHED LINE SHOWS THE POINT AT WHICH INTRODUCTIONS TOOK
PLACE AND THE LETTER AT THE START OF THE TRAJECTORIES INDICATES THE IDENTITY OF THE INTRODUCED
SPECIES 4HE LETTER OR COMBINATION OF LETTERS ASSOCIATED WITH EACH LINE SHOW THE COMMUNITY
COMPOSITION OF THAT MICROCOSM AT THE END OF THE EXPERIMENT WEEKS AFTER INTRODUCTION
." POPULATION TRAJECTORY LINES ARE ONLY DRAWN TO THE POINT AT WHICH ONE OF THE ABOVE
CRITERIA WAS MET WHILE THE lNAL SPECIES COMPOSITION SHOWN WAS THAT RECORDED AFTER
WEEKS
0( 7ARREN 2 ,AW AND !* 7EATHERBY
DRIVEN BY PARTICULAR SPECIES &IG )N MOST INVASIONS SPECIES THAT SHOWED CLEAR
INITIAL SUCCESS WERE ALSO PRESENT IN THE lNAL COMMUNITY BUT %UPLOTES DESPITE
BEING A SUCCESSFUL INVADER IN SOME INSTANCES NEVER PERSISTED IN THE LONG TERM AND
!MOEBA THOUGH MORE PERSISTENT OFTEN SHOWED THE SAME PATTERN &IG 3UCH
SPECIES PROVIDE A COUNTER EXAMPLE TO THE THEORETICAL PREDICTIONS OUTLINED ABOVE
,AW AND -ORTON WHICH SUGGEST THAT SUCCESSFUL INVADERS SHOULD BECOME
PERSISTENT MEMBERS OF THE INVADED SYSTEM EVEN IF THEY CAUSE OTHER CHANGES
)NTERESTINGLY THESE SPECIES WERE THE TWO OBLIGATE PREDATORS IN OUR SPECIES POOL AND
IT MAY BE THAT THE DISCREPANCY WAS RELATED TO TROPHIC POSITION $EMOGRAPHIC STO
CHASTICITY IS A POSSIBLE CAUSE OF THE VULNERABILITY OF INTRODUCED PREDATORS BECAUSE
FOR REASONS OF EITHER ENERGY AVAILABILITY OR OSCILLATORY POPULATION DYNAMICS THEIR
POPULATION SIZES COULD HAVE BEEN SMALL FROM TIME TO TIME
4HERE IS CLEARLY VARIATION IN THE RELATIONSHIP BETWEEN INITIAL ESTABLISHMENT
AND LONG TERM PERSISTENCE AT THE LEVEL OF INDIVIDUAL REPLICATES WHICH HAS OBVIOUS
IMPLICATIONS FOR PREDICTING THE OUTCOME OF PARTICULAR INSTANCES OF AN INTRODUCTION
&IG -ISMATCH IN ESTABLISHMENT AND LONG TERM PERSISTENCE OF PROTISTS INTRODUCED INTO
MICROCOSMS FROM 7EATHERBY "ARS SHOW THE PERCENTAGE OF MICROCOSMS IN WHICH
THE INTRODUCED SPECIES BECAME ESTABLISHED OPEN BARS AND PERSISTED IN THE LONG TERM
WEEKS SHADED BARS $ATA ARE GIVEN FOR ALL INDIVIDUAL MICROCOSMS ACROSS ALL RESIDENT
COMMUNITIES INTO WHICH THE SPECIES WAS INTRODUCED SO VALUES TAKE INTO ACCOUNT THE MIXED
OUTCOMES SHOWN IN &IG N IS THE NUMBER OF MICROCOSMS INTO WHICH EACH SPECIES WAS
INTRODUCED USUALLY REPLICATES OF EACH RESIDENT COMMUNITY BUT IN SOME CASES ONLY OR
)NVASIONS IN MICROCOSMS
(OWEVER WE CAN ALSO LOOK AT THE BROADER PICTURE AND FOCUS ON THE RESIDENT INTRO
DUCTION COMBINATIONS IN WHICH THE MAJORITY OF REPLICATES HAD THE SAME OUT
COME BOTH IN TERMS OF ESTABLISHMENT AND COMPOSITION OF THE lNAL COMMUNITY
4AKING JUST THESE MORE CONSISTENT OUTCOMES A TOTAL OF THIRTY lVE RESIDENT INTRO
DUCTION COMBINATIONS IN THE MAJORITY OF CASES THE INTRODUCED SPECIES EITHER
ESTABLISHED AND PERSISTED OR FAILED BOTH TO ESTABLISH AND TO PERSIST
4HERE WERE RELATIVELY FEW CASES IN WHICH SPECIES ESTABLISHED AND THEN FAILED
TO PERSIST AND NONE IN WHICH SPECIES FAILED TO ESTABLISH BUT YET PERSISTED
4HIS LAST CATEGORY MAY SEEM A TRUISM BUT IT IS NOT SO BECAUSE IT WOULD BE POSSIBLE
FOR AN INVADER TO PERSIST WITHOUT SHOWING NET POPULATION GROWTH THEREFORE FAILING
TO MEET THE CRITERION FOR ESTABLISHMENT 4HUS THE MESSAGE FROM THESE MICROCOSM
STUDIES IS THAT INITIAL ESTABLISHMENT OF AN INTRODUCED SPECIES WITHIN A COMMUNITY
WHICH COULD FEASIBLY BE MEASURED EVEN IN THE lELD IS A FAIR BUT NOT PERFECT
PREDICTOR OF ITS FATE IN THE LONG TERM
$O THE INVADERS HAVE AN IMPACT ON THE RESIDENTS
7E HAVE SO FAR FOCUSED ON THE FATE OF THE INVADER BUT IT IS OFTEN NOT THE INVADER
AS SUCH WHICH IS OF PRIMARY CONCERN BUT RATHER THE IMPACT OF THE INVADER ON THE
COMMUNITIES IT INVADES 0ARKER ET AL 'ENERALIZATIONS ABOUT THESE IMPACTS
ARE BOUND TO BE HARD TO MAKE BECAUSE OF THE COMPLEX NONLINEAR COUPLINGS THAT
CHARACTERIZE COMMUNITY DYNAMICS )T IS ENTIRELY POSSIBLE FOR THE IMPACT TO BE NEGLI
GIBLE OR FOR OTHER SPECIES TO BE DRIVEN TO EXTINCTION !N INVADER MAY TARGET PARTIC
ULAR SPECIES EG $UTCH %LM DISEASE IN THE 5+ 7ILLIAMSON OR HAVE EFFECTS
THAT RAMIFY WIDELY THROUGH THE COMMUNITY EG WHOLE ECOSYSTEM EFFECTS OF
-YRICA FAYA ON VOLCANIC SITES IN (AWAII 6ITOUSEK AND 7ALKER THE DEMISE
OF KELP BEDS FOLLOWING KILLER WHALE ARRIVAL ON THE 0ACIFIC COAST OF . !MERICA
%STES ET AL -ICROCOSM COMMUNITIES ARE ESPECIALLY AMENABLE TO SYSTEM
ATIC INVESTIGATION OF IMPACTS OF INVADERS BECAUSE WE CAN TAKE THEM APART IN AN
ORDERLY REPLICATED WAY TO SEE WHAT EFFECT EACH INVADING SPECIES EVENTUALLY HAS ON
THE SPECIES COMPOSITION OF EACH COMMUNITY AND HOW THIS RELATES TO THE INVADERS
OWN ABILITY TO PERSIST 6ARIOUS SCENARIOS ARE POSSIBLE THE INVADER PERSISTS
AND THE RESIDENT COMMUNITY IS CHANGED THE INVADER PERSISTS AND THE RESIDENT
COMMUNITY IS UNCHANGED THE INVADER FAILS TO PERSIST BUT THE RESIDENT COMMU
NITY IS CHANGED THE INVADER FAILS AND THE RESIDENT COMMUNITY IS UNCHANGED
3ITUATION IS THE CLASSIC @PROBLEM INVASION AN INVADER THAT BECOMES A PART
OF THE COMMUNITY AND CHANGES IT USUALLY BY GENERATING ONE OR MORE EXTINCTIONS
IN THE RESIDENTS THE INVASION OF THE COMMUNITY COMPRISING "LEPHARISMA AND
#OLPIDIUM BY 0ARAMECIUM &IG A PROVIDES AN EXAMPLE OF THIS FROM OUR MICRO
COSM EXPERIMENTS 4HESE ARE ALSO THE BEST DOCUMENTED INVASION CASE STUDIES IN
NATURAL SYSTEMS EG %LTON $RAKE ET AL BECAUSE INVADERS THAT PERSIST
ARE MOST READILY DOCUMENTED AND STUDIED AND THEIR IMPACTS ARE OF CONCERN
)N WE HAVE SIMPLE AUGMENTATION OF THE COMMUNITY BY AN INVADER /F COURSE
THE COLONIZING SPECIES MAY ALTER DENSITIES OF OTHER SPECIES BUT NOT TO A SUFlCIENT
0( 7ARREN 2 ,AW AND !* 7EATHERBY
EXTENT TO CAUSE SPECIES LOSSES )NVASION OF "LEPHARISMA SYSTEMS BY 0ARAMECIUM PRO
VIDES AN EXAMPLE ALTHOUGH THE RESIDENT PERSISTED &IG B IT WAS ABOUT ONE lFTH
AS ABUNDANT IN THE INVADED SYSTEMS AS IN THE UNINVADED CONTROLS AT THE END OF THE
EXPERIMENT 4HERE ARE EXAMPLES FROM NATURAL SYSTEMS OF INVADERS WHICH APPEAR TO
ADD THEMSELVES TO SYSTEMS BUT CAUSE LITTLE IMPACT EG THE FULMAR &ULMARIS GLACIA
LIS AN ARCTIC SEABIRD 7ILLIAMSON ET SEQ 4HE FREQUENCY OF NON
IMPACTIVE
INVASIONS HOWEVER REMAINS A CONTROVERSIAL ISSUE 0ARKER ET AL FOR EXAMPLE
SEE 3IMBERLOFF (ERBOLD AND -OYLE 0IMM BECAUSE IT CAN BE DIF
lCULT TO ESTABLISH IMPACTS AND TEST FOR CAUSALITY IF THERE ARE NO SUITABLE UNINVADED
SYSTEMS FOR COMPARISON
3CENARIO REQUIRES THAT A SPECIES FAILS TO PERSIST BUT HAS SUFlCIENTLY STRONG
EFFECTS ON THE RESIDENT COMMUNITY WHILE IT IS PRESENT TO CAUSE CHANGES IN SPE
CIES COMPOSITION 3UCH SPECIES CAN BE TERMED @CATALYSTS )N OUR EXPERIMENTS
CATALYSTS WERE NOT COMMON BUT TWO SPECIES DID PLAY THIS ROLE IN SOME SITUA
TIONS PRINCIPALLY %UPLOTES AND ALSO !MOEBA IN OCCASIONAL REPLICATES &OR EXAMPLE
%UPLOTES ESTABLISHED ITSELF IN A COMMUNITY COMPRISING 0ARAMECIUM #OLPIDIUM AND
4ETRAHYMENA CAUSED THE LATTER TWO SPECIES TO GO EXTINCT IN MOST SYSTEMS AND THEN
WENT TO EXTINCTION ITSELF &IG C )N THIS EXAMPLE %UPLOTES DID ESTABLISH INCREASE
IN POPULATION SIZE AFTER INTRODUCTION BUT THERE WERE ALSO CASES EG %UPLOTES
INVADING THE COMMUNITY OF 0ARAMECIUM AND 4ETRAHYMENA IN WHICH THE INVADER
DID NOT CONSISTENTLY INCREASE ABOVE ITS INITIAL NUMBERS BUT NONETHELESS CAUSED AN
EXTINCTION OF ONE OF THE RESIDENTS 4ETRAHYMENA BEFORE ITSELF GOING EXTINCT 4HERE
IS SOME THEORETICAL EVIDENCE OF SUCH EFFECTS IN MODELS OF INVASIONS INTO COMPETI
TIVE SYSTEMS #ASE 3OME EXAMPLES OF SITUATIONS IN NATURAL SYSTEMS SEEM
TO SHOW THIS BEHAVIOUR EG &LUX BUT THE APPARENT RARITY OF SUCH OBSERVA
TIONS 7ILLIAMSON IS CONSISTENT WITH THE RELATIVE RARITY OF THESE EFFECTS IN OUR
EXPERIMENTAL SYSTEMS
&INALLY WE HAVE SCENARIO )N THIS CASE THE FAILURE OF THE INTRODUCED SPECIES
TO PERSIST COUPLED WITH THE LACK OF CHANGE IN THE RESIDENT COMMUNITY EG &IG
D SUGGESTS THAT EFFECTIVELY THERE IS NOTHING WE WOULD RECOGNIZE AS AN INVASION
AT ALL )N THE MAJORITY OF INTRODUCTIONS WITH THIS OUTCOME THIS WAS VERY MUCH THE
CASE WITH THE INTRODUCED SPECIES SIMPLY DECLINING TO EXTINCTION AFTER INTRODUCTION
(OWEVER THERE WERE ALSO A FEW CASES IN WHICH INTRODUCED SPECIES SHOWED SUCCESS
FUL INITIAL ESTABLISHMENT WHILE NONETHELESS EVENTUALLY GOING EXTINCT AND LEAVING
THE RESIDENT SET UNCHANGED FOR EXAMPLE IN MOST INTRODUCTIONS OF %UPLOTES INTO
POPULATIONS OF 4ETRAHYMENA %UPLOTES ACHIEVED POPULATIONS OF SEVERAL THOUSANDS
OF INDIVIDUALS BEFORE DISAPPEARING %VIDENTLY AN INTRODUCED SPECIES CAN ACHIEVE A
SIGNIlCANT PRESENCE DURING SOME PART OF AN ULTIMATELY UNSUCCESSFUL INVASION
"ECAUSE OUR MICROCOSM EXPERIMENTS CONTAINED ALL POSSIBLE PERMUTATIONS OF
RESIDENT COMMUNITY AND INTRODUCED SPECIES WE CAN USE THESE RESULTS TO LOOK AT
THE FREQUENCIES OF THESE VARIOUS OUTCOMES AND THE CIRCUMSTANCES UNDER WHICH
THEY OCCUR 4AKING JUST THE THIRTY
SIX RESIDENT
INTRODUCTION TREATMENTS IN WHICH
THE MAJORITY OF REPLICATES ENDED UP WITH THE SAME SPECIES COMPOSITION IN THE LONG
TERM THE RESIDENT COMMUNITY WAS ALTERED IN JUST NINE OF THESE 7ARREN ET AL
)NVASIONS IN MICROCOSMS
&IG %XAMPLES OF INITIAL GROWTH OF INTRODUCED PROTISTS AND THE COMMUNITIES THAT
PERSISTED IN THE LONG TERM FOR COMBINATIONS OF PERSISTENCE AND NON
PERSISTENCE OF
THE INVADER AND IMPACT OR LACK OF IMPACT ON THE RESIDENTS 3TRUCTURE AND NOTATION AS
FOR &IG
!S ONE MIGHT EXPECT EG #ASE CHANGES TO THE RESIDENT C OMMUNITY BECAME
MORE LIKELY AS THE NUMBER OF RESIDENT SPECIES INCREASED RISING FROM ABOUT IN
ONE SPECIES COMMUNITIES TO IN TWO SPECIES COMMUNITIES TO IN THREE
SPECIES COMMUNITIES THOUGH THERE ARE ONLY TWO THREE SPECIES COMBINATIONS
4HE EFFECT OF THE INTRODUCED SPECIES WAS MOST OFTEN NEGATIVE CAUSING EXTINCTION
OF RESIDENT SPECIES SIX CASES ALTHOUGH THERE WERE THREE INSTANCES OF THE REVERSE
EFFECT IE A RESIDENT SPECIES #OLPIDUM IN TWO SYSTEMS AND 0ARAMECIUM IN ONE
FOR WHICH PERSISTENCE SEEMED TO BE ENHANCED BY THE PRESENCE OF THE INVADERS
!MOEBA 0ARAMECIUM AND %UPLOTES
0UTTING THESE RESULTS TOGETHER WE CAN BREAK DOWN THE THIRTY SIX RESIDENT
INTRODUCTION TREATMENTS INTO THOSE IN WHICH THE INTRODUCED SPECIES PERSISTED AND
THE RESIDENT COMMUNITY WAS CHANGED THE INTRODUCED SPECIES PERSISTED BUT
THE RESIDENT COMMUNITY WAS NOT CHANGED THE INTRODUCED SPECIES DID NOT PER
SIST BUT THE RESIDENT COMMUNITY WAS CHANGED AND THE INTRODUCED SPECIES DID
NOT PERSIST AND THE RESIDENTS WERE UNCHANGED #LEARLY THERE IS NO SIGNIlCANT
ASSOCIATION BETWEEN THE LIKELIHOOD OF ESTABLISHMENT OF INTRODUCED SPECIES AND
0( 7ARREN 2 ,AW AND !* 7EATHERBY
THE LIKELIHOOD OF THE RESIDENT COMMUNITY BEING ALTERED &ISHERS %XACT 4EST
P )F THIS SORT OF PATTERN TURNS OUT TO HAVE ANY SORT OF GENERALITY IT LENDS
WEIGHT TO THE VIEW THAT KNOWING A SPECIES IS LIKELY TO BECOME ESTABLISHED AND
PERSIST IN A SYSTEM MAY PROVIDE NO USEFUL GUIDE TO WHETHER IT IS GOING TO CHANGE
THE EXISTING COMMUNITY IN OTHER WAYS
).6!3)/.3 !.$ #/--5.)49 !33%-",9
!LTHOUGH OUR MAIN FOCUS HERE HAS BEEN ON THE COMPONENTS OF INDIVIDUAL INVA
SIONS THE ARRIVAL OF A NEW SPECIES AND ANY CHANGES IN COMMUNITY COMPOSITION
THAT ACCOMPANY THIS ARE JUST A SINGLE STEP IN A LONGER
TERM PROCESS OF COMMUNITY
DEVELOPMENT THROUGH THE GRADUAL TURNOVER OF SPECIES 3UCH TURNOVER GENERALLY
TERMED @COMMUNITY ASSEMBLY IS DETERMINED BY FACTORS ACTING AT VERY DIFFERENT
SCALES FROM BIOGEOGRAPHIC CONSTRAINTS ON THE SPECIES POOL THROUGH TO PATTERNS
OF NICHE DIFFERENTIATION AMONG COMPETITORS IN A COMMUNITY $IAMOND AND
#ASE $RAKE A 'RAY ET AL 7EIHER AND +EDDY BUT HAS THE
SAME UNDERLYING DRIVER THE ARRIVAL ESTABLISHMENT AND SOMETIMES PERSISTENCE OF
SPECIES FROM OUTSIDE THE SYSTEM IE INVASION (ERE CERTAINLY THE LINKS BETWEEN
COMMUNITY ECOLOGY AND INVASION BIOLOGY ARE EXPLICIT INVASIONS DRIVE COMMUNITY
CHANGE /F COURSE IN MOST OF THE @INVASIONS EFFECTING SUCH CHANGES THE INVADER IS
NOT FROM A NEW REGION DISSOCIATED FROM THE SPECIES IN THE RESIDENT COMMUNITY
BUT NONETHELESS THE COMPONENTS OF THE INVASION PROCESS WE HAVE DISCUSSED ABOVE
APPLY EQUALLY TO WHAT MIGHT BE TERMED @LOCAL INVASIONS AND THEREFORE TO COM
MUNITY CHANGE IN GENERAL
-ICROCOSM SYSTEMS FOR THE SAME REASONS THAT MAKE THEM USEFUL FOR STUDY
ING INVASIONS HAVE PLAYED A SIGNIlCANT ROLE IN EXPLORING COMMUNITY ASSEMBLY
)N PARTICULAR MICROCOSMS OF ALGAE PROTISTS AND SMALL METAZOANS HAVE BEEN USED
TO EXAMINE THE EFFECTS OF NON
SIMULTANEOUS INVASION INVASION ORDER AND INVASION
RATE ON THE DEVELOPMENT OF COMMUNITY STRUCTURE $ICKERSON AND 2OBINSON
2OBINSON AND $ICKERSON 2OBINSON AND %DGEMON $RAKE
&UKAMI AND -ORIN 4HE GENERAL CONCLUSION FROM THESE STUDIES IS
THAT CONTINGENCIES IN THE INVASION PROCESS CAN RESULT IN SUBSTANTIAL DIFFERENCES IN
THE STRUCTURE OF THE lNAL COMMUNITIES THAT ARE FORMED 3UCH STUDIES ARE GENERALLY
PRESENTED IN THE CONTEXT OF COMMUNITY ASSEMBLY AND DOCUMENT THE OUTCOMES OF
DIFFERENT SEQUENCES OR RATES OF SPECIES ARRIVAL RATHER THAN THE DETAILS OF INDIVIDUAL
INVASIONS WHICH HAVE BEEN OUR FOCUS HERE BUT IT IS CLEAR THAT THE SORTS OF EFFECTS
WE SEE IN INDIVIDUAL INVASIONS IN PARTICULAR STOCHASTIC EFFECTS IN LOW POPULATIONS
AND THE INTERDEPENDENCE OF SUCCESS ON INVADER AND RESIDENT COMMUNITY IDENTITIES
COULD BE THE DRIVERS OF SUCH ASSEMBLY SEQUENCE EFFECTS EG $RAKE $RAKES
STUDY EVEN FOUND EVIDENCE TO SUGGEST THAT INVASIBILITY OF SYSTEMS WITH IDENTICAL
SPECIES COMPOSITIONS MAY TO BE AFFECTED BY DIFFERENCES IN THEIR ASSEMBLY HISTORY
-ICROCOSMS ARE ALSO BEING USED TO EXAMINE THE EFFECTS OF THE CONTINUOUS ASSEMBLY
AND DISASSEMBLY OF COMMUNITIES IN MULTIPLE HABITAT PATCHES WHERE INVASIONS ARE
)NVASIONS IN MICROCOSMS
A NECESSARY AND NATURAL PART OF THE MAINTENANCE OF POPULATIONS AND COMMUNITIES
WITHIN A @LANDSCAPE $RAKE ET AL (OLYOAK AND ,AWLER A B 7ARREN
"URKEY (OLT ET AL AND WITH THE POSSIBLE THOUGH LARGELY
UNEXPLORED OPPORTUNITY SUCH WORK PROVIDES FOR EXAMINING THE SPREAD OF INVADERS
THROUGH A COMMUNITY LANDSCAPE
0UTTING THESE COMPONENTS TOGETHER ALLOWS US TO CONSTRUCT A DESCRIPTION OF
THE COMMUNITY SPACE PERSISTENT COMMUNITIES AND THE ROUTES THAT COMMUNITY
ASSEMBLY CAN TAKE THROUGH THAT SPACE AS A RESULT OF INVASIONS INTO EACH COMMUNI
TY :IMMERMAN ET AL 7ARREN ET AL WHICH WE CAN TERM AN ASSEMBLY
GRAPH EG &IG !SSEMBLY GRAPHS CAN CLARIFY FEATURES OF COMMUNITY ASSEMBLY
THAT WOULD OTHERWISE REMAIN OBSCURE &OR EXAMPLE THE GRAPH SHOWN IN &IG
HAS A CYCLIC END
STATE DRIVEN BY THE PARTICULAR CHARACTERISTICS OF %UPLOTES %UPLOTES
INVADES THE COMMUNITY COMPOSED OF 0ARAMECIUM AND 4ETRAHYMENA ELIMINATES
THE 4ETRAHYMENA AND THEN ITSELF GOES TO EXTINCTION LEAVING A COMMUNITY OF JUST
0ARAMECIUM WHICH CAN THEN BE REINVADED BY 4ETRAHYMENA TO RETURN TO THE ORIGINAL
COMMUNITY !NOTHER FEATURE OF THE GRAPH IS THE EXISTENCE OF COMMUNITIES SUCH
AS THAT COMPRISING #OLPIDIUM AND 4ETRAHYMENA WHICH PERSIST IF ALL SPECIES ARE
INTRODUCED TOGETHER BUT WHICH CANNOT READILY BE CREATED BY SEQUENTIAL INVASION
FROM ANY OTHER PERSISTENT COMMUNITY 4HE GENERALITY OF SUCH FEATURES REMAINS TO
BE ESTABLISHED BUT WHAT IS CLEAR IS THAT UNDERSTANDING THE INVASION PROCESS LIES AT
THE HEART OF UNDERSTANDING THIS FUNDAMENTAL ASPECT OF COMMUNITY ECOLOGY
-)#2/#/3-3 !3 3934%-3 &/2 345$9).' ).6!3)/. ")/,/'9
2EMARKABLY LITTLE WORK ON BIOLOGICAL INVASIONS HAS BEEN DONE IN MICROCOSM SYS
TEMS YET EVEN FROM THE FEW STUDIES THERE ARE A NUMBER OF MESSAGES EMERGE 7ORK
IN MICROCOSMS HIGHLIGHTS THE PROBABILISTIC NATURE OF THE INVASION PROCESS THE JOINT
DEPENDENCE OF INVASION SUCCESS ON BOTH INVADER IDENTITY AND RESIDENT COMMUNITY
COMPOSITION THE RELATIONSHIPS BETWEEN DIFFERENT PHASES OF THE INVASION PROCESS
ESTABLISHMENT PERSISTENCE IMPACT AND THE IMPORTANCE OF DISTURBANCE ENERGY
AND SPECIES NUMBER
4HIS SEEMS A RESPECTABLE YIELD OF INFORMATION FROM A MODEST AMOUNT OF
EXPERIMENTAL WORK 4HESE RESULTS WERE MADE POSSIBLE BY THE HAVING THE FACILITY
TO MANIPULATE RESIDENT AND INTRODUCED SPECIES BY FOLLOWING THE OUTCOMES OVER
TENS TO HUNDREDS OF GENERATIONS AND BY PUTTING IN PLACE ENOUGH REPLICATION TO
PARTITION VARIATION WITHIN AND BETWEEN TREATMENTS 4HERE IS LITTLE LIKELIHOOD OF
BEING ABLE TO CARRY OUT EXPERIMENTS AT EQUIVALENT TEMPORAL AND SPATIAL SCALES IN
THE lELD &IELD STUDIES HAVE PROVIDED IMPORTANT INSIGHTS INTO ALL THESE ELEMENTS
OF INVASIONS BUT MICROCOSMS BRING THE FACILITY TO RELATE THE LONG AND SHORT
TERM
OUTCOMES OF THE INVASION PROCESS AND QUANTIFY THE STOCHASTICITIES INVOLVED /F
COURSE THE SCARCITY OF SUCH STUDIES IN MICROCOSMS PRECLUDES MUCH ASSESSMENT OF
THE GENERALITY OF THESE RESULTS AS IT WOULD IN ANY SYSTEM 4HE DIFFERENCE IS THAT IN
MICROCOSMS TESTING THIS GENERALITY IS AN ENTIRELY FEASIBLE PROSPECT
0( 7ARREN 2 ,AW AND !* 7EATHERBY
&IG 4HE ASSEMBLY GRAPH FOR THE SIX SPECIES PROTIST SYSTEM DESCRIBED IN 7ARREN ET AL
4HE LETTER CODES AS FOR &IG IN BRACKETS REPRESENT THE PERSISTENT COMMUNITIES AND
THE ARROWS REPRESENT TRANSITIONS BETWEEN THOSE COMMUNITIES PRODUCED BY INTRODUCTIONS OF
THE SPECIES REPRESENTED BY THE LETTER CODE ON THE ARROW !RROWS ARE BASED ON THE MAJORITY
TRANSITIONS OBSERVED IN THE EXPERIMENTS IN EACH CASE SO THAT ONLY A SINGLE OUTCOME FROM
ANY PARTICULAR INVASION IS DEPICTED SEE 7ARREN ET AL FOR AN ALTERNATIVE PRESENTATION
THAT INCLUDES THE VARIATION IN OUTCOMES $OTTED ARROWS SHOW WHERE CERTAIN RESIDENT
COMMUNITIES HAD A TENDENCY TO COLLAPSE EVENTUALLY EVEN IN THE ABSENCE OF INVASIONS AND
THE DASHED ARROWS ARE TRANSITIONS THAT ARE BASED ON INFORMATION FROM ONLY TWO REPLICATES
4HE COMMUNITIES DEPICTED IN GREY ARE THOSE FORMED AS A RESULT OF INVASIONS BUT NOT FOUND
IN INITIAL EXPERIMENTS TO ESTABLISH WHICH SETS OF SPECIES COULD COEXIST AND HENCE FOR WHICH
NO INVASION TRIALS WERE CARRIED OUT AND SO WE HAVE NO INFORMATION ON THEIR INVASIBILITY
&IGURE REPRODUCED BY PERMISSION OF THE %COLOGICAL 3OCIETY OF !MERICA
)NVASIONS IN MICROCOSMS
4HAT SUCH QUESTIONS COULD BE ADDRESSED IS THOUGH PERHAPS NOT THE ISSUE 4HOSE
SCEPTICAL OF THE MICROCOSM APPROACH WOULD QUESTION WHETHER EVEN IF WE HAD SUCH
STUDIES WE WOULD BE ANY THE WISER ABOUT INVASIONS IN THE lELD 4HE ARGUMENTS
ABOUT USING MICROCOSMS IN ECOLOGY ARE WELL REHEARSED EG $RAKE ET AL
#ARPENTER $RENNER AND -AZUMDER ,AWLER *ESSUP ET AL
BUT AGAINST THE LIMITATIONS WE HAVE TO SET THE ADVANTAGES OUTLINED ABOVE
-ICROCOSMS DO NOT ALLOW US TO MAKE PRACTICAL MANAGEMENT DECISIONS ABOUT
SPECIlC INVASIONS BUT THEY DO ALLOW US TO TEST OUR INTUITION REASONING AND THEORY
AS TO THE BASIC ECOLOGICAL PRINCIPLES THAT UNDERPIN THEM $RAKE ET AL ,AWLER
7E TAKE THE VIEW THAT MICROCOSMS CAN AND SHOULD PLAY A SIGNIlCANT ROLE
IN DEVELOPING OUR UNDERSTANDING OF INVASION PROCESSES 7E WILL UNDERSTAND INVA
SIONS BETTER ARMED WITH GOOD EXPERIMENTAL DATA FROM MICROCOSMS IN COMBINATION
WITH THEORY lELD EXPERIMENTS AND OBSERVATIONAL DATA THAN FROM ANY OF THESE
SOURCES ALONE AND WE WILL ALSO END UP KNOWING A GREAT DEAL MORE ABOUT HOW
COMMUNITIES WORK AS A CONSEQUENCE
!#+./7,%$'%-%.43
7E ARE GRATEFUL TO THE EDITORS FOR THE INVITATION TO CONTRIBUTE TO THIS VOLUME /UR
THANKS GO TO 0ETER -ORIN ,IN *IANG AND :AC ,ONG FOR KINDLY SENDING US MANU
SCRIPTS PRIOR TO PUBLICATION TO -ARK 7ILLIAMSON AND +EN 4HOMPSON FOR HELPFUL
DISCUSSIONS ABOUT INVASIONS AND TO *EREMY &OX 4AD &UKAMI /WEN 0ETCHEY
$AVID 7ARDLE AND -ARK 7ILLIAMSON FOR THOUGHTFUL COMMENTS ON THE MANUSCRIPT
4HE EXPERIMENTAL WORK DISCUSSED IN THIS CHAPTER WAS FUNDED BY A GRANT FROM THE
5+ .ATURAL %NVIRONMENT 2ESEARCH #OUNCIL
2%&%2%.#%3
!NDERSON 2 - AND -AY 2 - )NFECTIOUS DISEASES OF HUMANS /XFORD 5NIVERSITY
0RESS /XFORD 5 +
"AILEY . 4 * %LEMENTS OF STOCHASTIC PROCESSES 7ILEY .EW 9ORK 5 3 !
"URKEY 4 6 -ETAPOPULATION EXTINCTION IN FRAGMENTED LANDSCAPES USING BACTERIA
AND PROTOZOA COMMUNITIES AS MODEL ECOSYSTEMS !M .AT
"YERS * % AND .OONBURG % ' 3CALE DEPENDENT EFFECTS OF BIOTIC RESISTANCE
TO
BIO
LOGICAL INVASION %COLOGY
#ARPENTER 3 2 -ICROCOSM EXPERIMENTS HAVE LIMITED RELEVANCE FOR COMMUNITY AND
ECOSYSTEM ECOLOGY %COLOGY
#ARPENTER 3 2 -ICROCOSM EXPERIMENTS HAVE LIMITED RELEVANCE FOR COMMUNITY AND
ECOSYSTEM ECOLOGY REPLY %COLOGY
#ASE 4 * )NVASION RESISTENCE ARISES IN STRONGLY INTERACTING SPECIES RICH MODELS OF
COMPETITION COMMUNITIES 0ROC .ATN !CAD 3CI 5 3 !
0( 7ARREN 2 ,AW AND !* 7EATHERBY
#ASE 4 * 3URPRISING BEHAVIOR FROM A FAMILIAR MODEL AND IMPLICATIONS FOR COMPETI
TION THEORY !M .AT
#RAWLEY - * 4HE POPULATION BIOLOGY OF INVADERS 0HIL 4RANS 2OY 3OC ,OND "
#RAWLEY - * #HANCE AND TIMING IN BIOLOGICAL INVASIONS PP )N $RAKE
* ! -OONEY ( ! DI #ASTRI & 'ROVES 2 ( +RUGER & * 2EJMANEK - AND
7ILLIAMSON - EDS "IOLOGICAL INVASIONS A GLOBAL PERSPECTIVE 3#/0% *OHN 7ILEY
AND 3ONS #HICHESTER 5+
$AVIS - ! 'RIME * 0 AND 4HOMPSON + &LUCTUATING RESOURCES IN PLANT COMMU
NITIES A GENERAL THEORY OF INVASIBILITY * %COL
$AVIS - ! AND 0ELSOR - %XPERIMENTAL SUPPORT FOR A RESOURCE BASED MECHANISTIC
MODEL OF INVASIBILITY %COL ,ETT
$IAMOND * AND #ASE 4 * /VERVIEW INTRODUCTIONS EXTINCTIONS EXTERMINATIONS
AND INVASIONS PP )N $IAMOND * AND #ASE 4 * EDS #OMMUNITY %COLOGY
(ARPER AND 2OW .EW 9ORK 5 3 !
$ICK * 4 ! 0LATVOET $ AND +ELLY $ 7 0REDATORY IMPACT OF THE FRESHWATER
INVADER $IKEROGAMMARUS VILLOSUS #RUSTACEA !MPHIPODA #AN * &ISH !QUAT 3CI
$ICKERSON * % AND 2OBINSON * $ 4HE ASSEMBLY OF MICROSCOPIC COMMUNITIES PAT
TERNS OF SPECIES IMPORTANCE 4RANS !M -ICROSC 3OC
$ICKERSON * % AND 2OBINSON * 6 -ICROCOSMS AS ISLANDS A TEST OF THE -AC!RTHUR
7ILSON EQUILIBRIUM THEORY %COLOGY
$ICKERSON * % AND 2OBINSON * 6 4HE CONTROLLED ASSEMBLY OF MICROCOSMIC COM
MUNITIES THE SELECTIVE EXTINCTION HYPOTHESIS /ECOLOGIA
$RAKE * ! A #OMMUNITIES AS ASSEMBLED STRUCTURES DO RULES GOVERN PATTERN
4RENDS %COL %VOL
$RAKE * ! B 4HE MECHANICS OF COMMUNITY ASSEMBLY AND SUCCESSION * 4HEOR "IOL
$RAKE * ! #OMMUNITY ASSEMBLY MECHANICS AND THE STRUCTURE OF AN EXPERIMENTAL
SPECIES ENSEMBLE !M .AT
$RAKE * ! &LUM 4 % 7ITTEMAN ' * 6OSKUIL 4 (OLYMAN ! - #RESON # +ENNY
$ ! (UXEL ' 2 ,ARUE # 3 AND $UNCAN * 2 4HE CONSTRUCTION AND ASSEMBLY
OF AN ECOLOGICAL LANDSCAPE * !NIM %COL
$RAKE * ! (UXEL ' 2 AND (EWITT # , -ICROCOSMS AS MODELS FOR GENERATING AND
TESTING COMMUNITY THEORY %COLOGY
$RAKE * ! -OONEY ( ! DI #ASTI & 'ROVES 2 ( +RUGER & * 2EJMNEK - AND
7ILLIAMSON - ED "IOLOGICAL INVASIONS A GLOBAL PERSPECTIVE 3#/0% *OHN
7ILEY AND 3ONS #HICHESTER 5+
$RENNER 2 7 AND -AZUMDER ! -ICROCOSM EXPERIMENTS HAVE LIMITED RELEVANCE
FOR COMMUNITY AND ECOSYSTEM ECOLOGY %COLOGY
$UKES * 3 "IODIVERSITY AND INVASIBILITY IN GRASSLAND MICROCOSMS /ECOLOGIA
%LTON # 3 4HE ECOLOGY OF INVASIONS BY ANIMALS AND PLANTS -ENTHUEN ,ONDON 5+
)NVASIONS IN MICROCOSMS
%STES * ! 7ILLIAMS - 4 AND $OAK $ & +ILLES WHALE PREDATION ON SEA OTTERS
LINKING OCEANIC AND NEARSHORE ECOSYSTEMS 3CIENCE
&LUX * % 7ORLD DISTRIBUTION PP
)N 4HOMPSON ( 6 AND +ING # - EDS
4HE %UROPEAN RABBIT THE HISTORY AND BIOLOGY OF A SUCCESSFUL INVADER /XFORD 5NIVERSITY
0RESS /XFORD
&UKAMI 4 AND -ORIN 0 * 0RODUCTIVITY
BIODIVERSITY RELATIONSHIPS DEPEND ON THE
HISTORY OF COMMUNITY ASSEMBLY .ATURE
'OEL . 3 AND 2ICHTER
$YN . 3TOCHASTIC MODELS IN BIOLOGY !CADEMIC 0RESS
.EW 9ORK 5 3 !
'RAY ! * AND #RAWLEY - * AND %DWARDS 0 * ED #OLONIZATION SUCCESSION AND
STABILITY "LACKWELL 3CIENCE /XFORD 5+
(ECTOR ! $OBSON + -INNS ! "AZELEY
7HITE % AND ,AWTON * ( #OMMUNITY
DIVERSITY AND INVASION RESISTANCE AN EXPERIMENTAL TEST IN A GRASSLAND ECOSYSTEM AND A
REVIEW OF COMPARABLE STUDIES %COL 2ES
(ERBOLD " AND -OYLE 0 " )NTRODUCED SPECIES AND VACANT NICHES !M .AT
(OBBS 2 * 4HE NATURE AND EFFECTS OF DISTURBANCE RELATIVE TO INVASIONS #H
PP
)N $RAKE * ! -OONEY ( ! DI #ASTRI & 'ROVES 2 ( +RUGER & *
2EJMNEK - AND 7ILLIAMSON - ED "IOLOGICAL INVASIONS A GLOBAL PERSPECTIVE 3#/0%
*OHN 7ILEY AND 3ONS #HICHESTER 5+
(ODGSON $ * 2AINEY 0 " AND "UCKLING ! -ECHANISMS LINKING DIVERSITY PRO
DUCTIVITY AND INVASIBILITY IN EXPERIMENTAL BACTERIAL COMMUNITIES 0ROC 2OY 3OC ,OND
"
(OLT ! 2 7ARREN 0 ( AND 'ASTON + * 4HE IMPORTANCE OF BIOTIC INTERACTIONS IN
ABUNDANCE
OCCUPANCY RELATIONSHIPS * !NIM %COL
(OLYOAK - AND ,AWLER 3 0 0ERSISTENCE OF AN EXTINCTION PRONE PREDATOR
PREY
INTERACTION THROUGH METAPOPULATION DYNAMICS %COLOGY
(OLYOAK - AND ,AWLER 3 0 4HE ROLE OF DISPERSAL IN PREDATOR
PREY METAPOPULATION
DYNAMICS * !NIM %COL
(OPPER + 2 AND 2OUSH 2 4 -ATE lNDING DISPERSAL NUMBER RELEASED AND THE
SUCCESS OF BIOLOGICAL CONTROL INTRODUCTIONS %COL %NTOMOL
(USTON - ! (IDDEN TREATMENTS IN ECOLOGICAL EXPERIMENTS RE
EVALUATING THE ECO
SYSTEM FUNCTION OF BIODIVERSITY /ECOLOGIA
*IANG , AND -ORIN 0 * 0RODUCTIVITY GRADIENTS CAUSE POSITIVE DIVERSITY
INVASIBILITY
RELATIONSHIPS IN MICROBIAL COMMUNITIES %COL ,ETT
*ESSUP # - +ASSEN 2 &ORDE 3 % +ERR " "UCKLING ! 2AINEY 0 " AND "OHANNAN
" * - "IG QUESTIONS SMALL WORLDS MICROBIAL MODEL SYSTEMS IN ECOLOGY 4RENDS
%COL %VOL
+OLA # 3 AND ,ODGE $ - 0ROGRESS IN INVASION BIOLOGY PREDICTING INVADERS
4RENDS %COL %VOL
,AW 2 AND -ORTON 2 $ 0ERMANENCE AND THE ASSEMBLY OF ECOLOGICAL COMMUNI
TIES %COLOGY
,AW 2 7EATHERBY ! * AND 7ARREN 0 ( /N THE INVASIBILITY OF PERSISTENT PROTIST
COMMUNITIES /IKOS
0( 7ARREN 2 ,AW AND !* 7EATHERBY
,AWLER 3 0 %COLOGY IN A BOTTLE USING MICROCOSMS TO TEST THEORY PP )N
2ESETARITS 7 * AND "ERNADO * EDS %XPERIMENTAL ECOLOGY ISSUES AND PERSPECTIVES
/XFORD 5NIVERSITY 0RESS .EW 9ORK
,EVINE * - 3PECIES DIVERSITY AND BIOLOGICAL INVASIONS RELATING LOCAL PROCESSES TO
COMMUNITY PATTERNS 3CIENCE
,EVINE * - AND $!NTONIO # - %LTON REVISITED A REVIEW OF EVIDENCE LINKING
DIVERSITY AND INVASIBILITY /IKOS
,ONSDALE 7 - 'LOBAL PATTERNS OF PLANT INVASIONS AND THE CONCEPT OF INVASIBILITY
%COLOGY
-ACK 2 . 0REDICTING THE IDENTITY AND FATE OF PLANT INVADERS EMERGENT AND EMERG
ING APPROACHES "IOL #ONSERV
-ACK 2 . 3IMBERLOFF $ ,ONSDALE - 7 %VANS ( #LOUT - AND "AZAZZ & !
"IOTIC INVASIONS CAUSES EPIDEMIOLOGY GLOBAL CONSEQUENCES AND CONTROL %COL !PPLIC
-C'RADY 3TEED * (ARRIS 0 - AND -ORIN 0 * "IODIVERSITY REGULATES ECOSYSTEM
PREDICTABILITY .ATURE
-ITCHELL # % AND 0OWER ! ' 2ELEASE OF INVASIVE PLANTS FROM FUNGAL AND VIRAL
PATHOGENS .ATURE
0ARKER ) $ 3IMBERLOFF $ ,ONSDALE 7 - 'OODELL + 7ONHAM - +AREIVA 0 -
7ILLIAMSON - ( 6ON (OLLE " -OYLE 0 " "YERS * % AND 'OLDWASSER ,
)MPACT TOWARD A FRAMEWORK FOR UNDERSTANDING THE ECOLOGICAL EFFECTS OF INVADERS "IOL
)NVASIONS
0ETCHEY / , -C0HEARSON 0 4 #ASEY 4 - AND -ORIN 0 * %NVIRONMENTAL
WARMING ALTERS FOOD WEB STRUCTURE AND ECOSYSTEM FUNCTION .ATURE
0ETERSEN * % AND (ASTINGS ! $IMENSIONAL APPROACHES TO SCALING EXPERIMENTAL
ECOSYSTEMS $ESIGNING MOUSETRAPS TO CATCH ELEPHANTS !M .AT
0IMENTEL $ ,ACH , :UNIGA 2 AND -ORRISON $ %NVIRONMENTAL AND ECONOMIC
COSTS OF NONINDIGENOUS SPECIES IN THE 5NITED 3TATES "IOSCIENCE
0IMM 3 , 4HEORIES OF PREDICTING SUCCESS AND IMPACT OF INTRODUCED SPECIES
PP )N $RAKE * ! -OONEY ( ! DI #ASTI & 'ROVES 2 ( +RUGER & *
2EJMNEK - AND 7ILLIAMSON - ED "IOLOGICAL INVASIONS A GLOBAL PERSPECTIVE 3#/0%
*OHN 7ILEY AND 3ONS #HICHESTER 5+
0OST 7 - AND 0IMM 3 , #OMMUNITY ASSEMBLY AND FOOD WEB STABILITY -ATH
"IOSCI
2EJMNEK - )NVASIBILITY OF PLANT COMMUNITIES #H PP )N $RAKE * !
-OONEY ( ! DI #ASTI & 'ROVES 2 ( +RUGER & * 2EJMNEK - AND 7ILLIAMSON
- ED "IOLOGICAL INVASIONS A GLOBAL PERSPECTIVE 3#/0% *OHN 7ILEY AND 3ONS
#HICHESTER 5+
2OBINSON * 6 AND $ICKERSON * % 4ESTING THE INVULNERABILITY OF LABORATORY ISLAND
COMMUNITIES TO INVASION /ECOLOGIA
2OBINSON * 6 AND $ICKERSON * % $OES INVASION SEQUENCE AFFECT COMMUNITY STRUC
TURE %COLOGY
2OBINSON * 6 AND %DGEMON - ! !N EXPERIMENTAL EVALUATION OF THE EFFECT OF
INVASION HISTORY ON COMMUNITY STRUCTURE %COLOGY
)NVASIONS IN MICROCOSMS
3AKAI ! + !LLENDORF & 7 (OLT * 3 ,ODGE $ - -OLOFSKY * 7ITH + ! "AUGHMAN
3 #ABIN 2 * #OHEN * % %LLSTRAND . # -C#AULEY $ % /.EIL 0 0ARKER ) -
4HOMPSON * . AND 7ELLER 3 ' 4HE POPULATION BIOLOGY OF INVASIVE SPECIES
!NN 2EV %COL 3YST
3HEA + AND #HESSON 0 #OMMUNITY ECOLOGY THEORY AS A FRAMEWORK FOR BIOLOGICAL
INVASIONS 4RENDS %COL %VOL
3HEA + AND 0OSSINGHAM ( 0 /PTIMAL RELEASE STRATEGIES FOR BIOLOGICAL CONTROL
AGENTS AN APPLICATION OF STOCHASTIC DYNAMIC PROGRAMMING TO POPULATION MANAGEMENT
* !PPL %COL
3HER ! ! AND (YATT , ! 4HE DISTURBED RESOURCE
mUX INVASION MATRIX A NEW
FRAMEWORK FOR PATTERNS OF PLANT INVASION "IOL )NVASIONS
3IMBERLOFF $ #OMMUNITY EFFECTS OF INTRODUCED SPECIES PP
)N .ITECKI - (
ED "IOTIC CRISES IN ECOLOGICAL AND EVOLUTIONARY TIME !CADEMIC 0RESS .EW 9ORK 53!
3IMBERLOFF $ 7HICH INSECT INTRODUCTIONS SUCCEED AND WHICH FAIL #H PP
)N $RAKE * ! -OONEY ( ! DI #ASTI & 'ROVES 2 ( +RUGER & * 2EJMNEK - AND
7ILLIAMSON - ED "IOLOGICAL INVASIONS A GLOBAL PERSPECTIVE 3#/0% *OHN 7ILEY
AND 3ONS #HICHESTER
3TAMPE % AND $AEHLER # # -YCORRHIZAL SPECIES IDENTITY AFFECTS PLANT COMMUNITY
STRUCTURE AND INVASION A MICROCOSM STUDY /IKOS
4HOMPSON + (ODGSON * ' AND 'RIME * 0 AND "URKE - * 7 0LANT TRAITS AND
TEMPORAL SCALE EVIDENCE FROM A
YEAR INVASION EXPERIMENT USING NATIVE SPECIES * %COL
4ORCHIN - % ,AFFERTY + $ $OBSON ! 0 -C+ENZIE 6 * AND +URIS ! -
)NTRODUCED SPECIES AND THEIR MISSING PARASITES .ATURE
6ITOUSEK 0 - AND 7ALKER , 2 "IOLOGICAL INVASION BY -YRICA FAYA IN (AWAII
PLANT DEMOGRAPHY NITROGEN lXATION ECOSYSTEM EFFECTS %COL -ONOGR
7ARDLE $ ! %XPERIMENTAL DEMONSTRATION THAT PLANT DIVERSITY REDUCES INVASIBIL
ITY %VIDENCE OF A BIOLOGICAL MECHANISM OR A CONSEQUENCE OF SAMPLING EFFECT /IKOS
7ARREN 0 ( $ISPERSAL AND DESTRUCTION IN A MULTIPLE HABITAT SYSTEM AND EXPERI
MENTAL APPROACH USING PROTIST COMMUNITIES /IKOS
7ARREN 0 ( ,AW 2 AND 7EATHERBY ! * -APPING THE ASSEMBLY OF PROTIST COM
MUNITIES IN MICROCOSMS %COLOGY
7EATHERBY ! * 3PECIES COEXISTENCE AND COMMUNITY ASSEMBLY IN PROTIST MICRO
COSMS UNPUBLISHED 0,$ THESIS 5NIVERSITY OF 3HEFlELD 5+
7EATHERBY ! * 7ARREN 0 ( AND ,AW 2 #OEXISTENCE AND COLLAPSE AN EXPERI
MENTAL INVESTIGATION OF THE PERSISTENT COMMUNITIES OF A PROTIST SPECIES POOL * !NIM
%COL
7EIHER % AND +EDDY 0 EDS %COLOGICAL ASSEMBLY RULES PERSPECTIVES ADVANCES
RETREATS #AMBRIDGE 5NIVERSITY 0RESS #AMBRIDGE 5+
7ILLIAMSON - "IOLOGICAL INVASIONS #HAPMAN AND (ALL ,ONDON %NGLAND
:IMMERMANN # 2 &UKAMI 4 AND $RAKE * ! !N EXPERIMENTALLY
DERIVED MAP
OF COMMUNITY ASSEMBLY SPACE )N -INAI ! AND "AR
9AM 9 EDS 5NIFYING THEMES
IN COMPLEX SYSTEMS )) 0ROCEEDINGS OF THE SECOND INTERNATIONAL CONFERENCE ON COMPLEX
SYSTEMS .EW %NGLAND #OMPLEX 3YSTEMS )NSTITUTE 0ERSEUS 0RESS
#HAPTER SEVENTEEN
5NDERSTANDING INVASIONS
IN PATCHY HABITATS THROUGH
METAPOPULATION THEORY
+ # (ARDING * - -C.AMARA AND 2 $ (OLT
).42/$5#4)/.
-ANY INVASIVE SPECIES EXPERIENCE A WORLD THAT IS HETEROGENEOUS AND PATCHY AND
EXPERIENCE IT AT MULTIPLE SCALES )N THIS CHAPTER WE WILL SHOW HOW METAPOPULATION
THEORY CAN BE USED TO EXPLORE GENERAL MECHANISMS WHICH DETERMINE THE INITIAL
SPREAD AND LONG TERM ESTABLISHMENT OF INVASIVE SPECIES IN NOVEL ENVIRONMENTS
-ETAPOPULATION ECOLOGY IS A BRANCH OF ECOLOGY THAT IS FOCUSED ON HOW SPECIES
PERSIST IN FRAGMENTED OR @SUBDIVIDED HABITATS (ANSKI (ANSKI AND 'AGGIOTTI
!LTHOUGH REAL LANDSCAPES ARE OFTEN HIGHLY COMPLEX WITH ADMIXTURES OF
SMOOTH GRADIENTS FRACTAL EDGES AND JUXTAPOSED HABITATS VARYING IN QUALITY FOR
MANY SPECIES IT IS NATURAL TO CONSTRUE THEIR WORLD AS CONSISTING OF SETS OF SUITABLE
HABITAT PATCHES ARRAYED WITHIN A LARGELY INHOSPITABLE MATRIX SEPARATING THOSE
PATCHES 3OME SUCH SPECIES INHABIT NATURALLY PATCHY HABITATS SUCH AS ISLANDS OR
PONDS &IG 3OME METAPOPULATIONS APPEAR IN ARTIlCIALLY CONSTRUCTED HABITAT
NETWORKS SUCH AS THE REMAINING FRAGMENTS OF A RAIN FOREST SURROUNDED BY CATTLE
PASTURES OR CONCRETE REEFS IN MARINE SYSTEMS 0ATCH NETWORKS CAN ALSO BE FORMED
IN MORE SUBTLE WAYS BUT STILL MAKE FUNCTIONAL METAPOPULATIONS 4HE PATCHY
DISTRIBUTION OF A HOST SPECIES FOR A SPECIALIST HERBIVORE PARASITE PARASITOID OR
-7 #ADOTTE ET AL EDS #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY n
3PRINGER 0RINTED IN THE .ETHERLANDS
+# (ARDING *- -C.AMARA AND 2$ (OLT
&IG 0ATCHY WORLD PONDS IN THE .ORTHERN 0RAIRIE FARMLAND 0HOTO 3TEWART 2% ET AL
&ROM .ORTHERN 0RAIRIE 7ILDLIFE 2ESEARCH #ENTER
COMMENSAL OR INDEED ANY VITAL AND SPATIALLY DELIMITED RESOURCE CAN LEAD TO
METAPOPULATION DYNAMICS FOR THE SPECIALIST "IOLOGICAL NONLINEARITIES CAN TRANS
FORM SMOOTH ENVIRONMENTAL GRADIENTS INTO PATCHY DISTRIBUTIONS &OR INSTANCE
STRONG !LLEE EFFECTS POSITIVE DENSITY DEPENDENCE AT LOW NUMBERS CAN AMPLIFY
PATCHINESS IN THE ENVIRONMENT 7HEN DISPERSAL IS LIMITED PERMANENT RANGE LIMITS
CAN ARISE IN PATCHY ENVIRONMENTS EVEN WITHOUT OVERALL ENVIRONMENTAL GRADIENTS
IN THE LANDSCAPE +EITT ET AL %VEN WITHOUT !LLEE EFFECTS IF THE DENSITY OF
SUITABLE PATCHES DECLINES ALONG AN ENVIRONMENTAL GRADIENT A SPECIES RANGE LIMIT
MAY EMERGE WHEN THERE IS NO DECLINE IN THE QUALITY OF SUITABLE HABITAT PATCHES
(OLT AND +EITT
! METAPOPULATION IS DElNED AS A SET OF POPULATIONS CONNECTED BY DISPERSAL
$ISPERSAL BECOMES PARTICULARLY VITAL TO UNDERSTANDING REGIONAL POPULATION PER
SISTENCE WHEN LOCAL POPULATIONS CAN BECOME EXTINCT AND MUST BE RE
ESTABLISHED
VIA COLONIZATION #LASSICAL METAPOPULATION THEORY THUS FOCUSES ON SYSTEMS WHERE
THERE IS A POTENTIAL FOR RECURRENT EXTINCTION )N ALL METAPOPULATIONS LONG
TERM
SPECIES PERSISTENCE RELIES ON THE BALANCE BETWEEN THE RATE OF EXTINCTION OF LOCAL
POPULATIONS AND THE RATE OF NEW COLONIZATIONS OF EMPTY HABITAT PATCHES &IG
-ETAPOPULATION THEORY IS WELL SUITED FOR DESCRIBING INVASIONS AND THE ESTAB
LISHMENT OF NEW SPECIES INTO PATCHY HABITATS 4HERE ARE THREE BASIC QUESTIONS
)NVASIVE SPECIES IN METAPOPULATION SYSTEMS
&IG "LUE SQUARES INDICATE PONDS OCCUPIED BY 0ISIDIUM NITIDUM IN THE YEAR OF SAMPLING
7HITE SQUARE ARE PONDS WHERE 0 NITIDUM HAS GONE EXTINCT 3TEWART AND +ANTRUD
&ROM .ORTHERN 0RAIRIE 7ILDLIFE 2ESEARCH #ENTER
ONE CAN ASK ABOUT AN INVASION &IRST CAN AN INVASION OCCUR AT ALL 3ECOND GIVEN
THAT AN INVASION OCCURS HOW RAPIDLY DOES IT OCCUR &INALLY FOLLOWING INVA
SION WHAT IMPACTS DOES THE INVADER HAVE UPON THE PRIOR RESIDENT COMMUNITY
-ETAPOPULATION THEORY CAN BE USED TO ADDRESS EACH OF THESE BASIC ISSUES
7E INTRODUCE A SIMPLE MODEL STRUCTURE THAT CAN BE USED TO SUMMARISE AND CLAS
SIFY SOME DIFFERENT TYPES OF METAPOPULATION INVASION SCENARIOS 7E EXAMINE SOME
SPECIAL CASES STARTING WITH A CASE IN WHICH THE INVASIVE SPECIES DOES NOT COMPETE
WITH ANY OTHER ECOLOGICALLY SIMILAR SPECIES 4HE DYNAMICS OF THE INVASION IN THIS
CASE IS THEN GOVERNED BY THE BIOLOGY OF THE INVADER ONLY AND CLASSIC SINGLE SPECIES
METAPOPULATION THEORY APPLIES 7E THEN CONSIDER A GENERAL METAPOPULATION MODEL
IN WHICH THE INVADING SPECIES INTERACTS WITH A NATIVE SPECIES 7E DElNE CRITERIA FOR
WHEN INITIAL INVASION OCCURS AND FOR WHEN LONG TERM COEXISTENCE IS POSSIBLE 7E
ALSO LOOK AT SOME INTERACTIONS THAT ARE SPECIlC CASES OF THIS GENERAL MODEL A THE
INVADER COMPLETELY EXCLUDES THE NATIVE SPECIES B THE INVADER IS DEPENDENT ON THE
PRESENCE OF A NATIVE SPECIES AND C THERE IS AN INTERMEDIATE LEVEL OF COMPETITION
7E ALSO TOUCH ON HOW SPATIAL HETEROGENEITY CAN INmUENCE INVASION /UR OVERVIEW
HIGHLIGHTS SOME GENERAL INSIGHTS ABOUT THE POPULATION BIOLOGY OF INVASION WHICH
EMERGE FROM METAPOPULATION THEORY
+# (ARDING *- -C.AMARA AND 2$ (OLT
./ #/-0%4)4)/. 7)4( .!4)6% 30%#)%3 #,!33)# -%4!0/05,!4)/.
")/,/'9 !00,)%3
,EVINS METAPOPULATION
3OME INVASIONS OCCUR WITHOUT ANY OBVIOUS COMPETITION FROM NATIVE SPECIES &OR
INSTANCE WHEN THE MARSH
LOVING MUSKRAT WAS INTRODUCED INTO CENTRAL %UROPE
ITS NUMBERS GREW EXPONENTIALLY 3KELLAM )N THAT ENVIRONMENT THERE
WERE NO OTHER OBVIOUS AQUATIC MAMMALS WHICH COULD INTERFERE WITH THE INITIAL
ESTABLISHMENT AND SUBSEQUENT GROWTH OF THIS SPECIES )N THESE CASES SINGLE SPECIES
METAPOPULATION THEORY CAN BE APPLIED TO INVESTIGATE BASIC PROPERTIES OF THE INVA
SION EG FOR THE MUSKRAT ITS REQUIRED HABITAT TYPICALLY OCCURS IN DISCRETE PATCHES
MARSHES SEPARATED BY A RELATIVELY HOSTILE MATRIX OF TERRESTRIAL HABITATS
-OREOVER IF RESIDENT SPECIES ARE RELATIVELY lXED IN ABUNDANCE EG THE INVADER
INITIALLY HAS LITTLE IMPACT THEN THE RESIDENTS CAN BE VIEWED AS lXED HABITAT FAC
TORS EITHER HAMPERING OR ASSISTING INVASION BY THE FOCAL SPECIES 7E SHALL DISTIN
GUISH BETWEEN TWO IMPORTANT STAGES IN THE INVASION PROCESS A THE INITIAL PHASE OF
POTENTIAL SUCCESSFUL INVASION OR FAILURE WHEN THE INVADER OCCURS IN RELATIVELY FEW
PATCHES AND B THE LONG TERM ESTABLISHMENT OF AN INVASIVE SPECIES
&IRST LET US DElNE THE MODEL FRAMEWORK ! SIMPLE MODEL THAT CAPTURES MUCH OF
THE SPECIAL METAPOPULATION LEVEL EFFECTS CAN BE BUILT ON THE FOUNDATION OF 2ICHARD
,EVINS SEMINAL PAPER ON METAPOPULATION THEORY (E REGARDED THE ENVI
RONMENT AS COMPOSED OF 4 PATCHES OF EQUAL QUALITY WHERE 4 IS LARGE (E MODELLED
THE DYNAMICS OF A PARTICULAR SPECIES WITHIN THIS LANDSCAPE ASSUMING THAT EACH
PATCH IS EITHER OCCUPIED BY MEMBERS OF THE SPECIES OR EMPTY /CCUPIED PATCHES
SEND OUT MIGRANTS THAT SETTLE ON PATCHES AT RANDOM %ACH OCCUPIED PATCH SENDS
OUT MIGRANTS THAT ARE SUCCESSFUL IN lNDING ANOTHER PATCH AT RATE M 7E ASSUME FOR
THE MOMENT THAT WE CAN IGNORE THE SPATIAL ARRANGEMENT OF PATCHES AND THAT THE
DISPERSERS FROM ALL OCCUPIED PATCHES MERGE INTO A REGIONAL MIGRANT POOL &ROM THIS
POOL THEY SETTLE UNIFORMLY ACROSS ALL PATCHES BOTH OCCUPIED AND EMPTY )F THERE
ARE . OCCUPIED PATCHES THE TOTAL POOL SIZE IS M. AND SO THE RATE OF ARRIVAL OF IMMI
GRANTS ON EACH EMPTY OR OCCUPIED PATCH IS SIMPLY _ M.4 3INCE THERE ARE
4 n . EMPTY PATCHES THE RATE OF COLONIZATION OF NEW PATCHES IS +_ 4 n . WHERE +
IS THE PROBABILITY THAT A SINGLE EMPTY PATCH IS COLONIZED IE CONVERTED FROM EMPTY
TO OCCUPIED BY A RESIDENT POPULATION OF THE SPECIES BY AN IMMIGRANT .OTE THAT IF
+ THIS MEANS THAT SOME COLONIZATIONS FAIL ,EVINS ALSO ASSUMED THAT EACH
OCCUPIED PATCH BECOMES EXTINCT THAT IS CHANGES FROM OCCUPIED TO EMPTY AT CON
STANT RATE E 4HESE SIMPLE ASSUMPTIONS CAN LATER BE GENERALIZED BUT ARE VERY USEFUL
AS A lRST STEP FOR D EVELOPING AN INTUITIVE FEELING FOR METAPOPULATION DYNAMICS
&IG 7ITH THESE
ASSUMPTIONS THE RATE OF CHANGE IN THE NUMBER OF OCCUPIED
PATCHES IS
D.
+_ 4 n . n E.
DT
)NVASIVE SPECIES IN METAPOPULATION SYSTEMS
&IG 4HE CLASSIC ,EVINS METAPOPULATION MODEL FOR A SINGLE SPECIES ! GIVEN
PATCH CAN BE IN ONE OF TWO STATES EMPTY OR OCCUPIED BY SPECIES 4HE RATE OF CHANGE IS
GIVEN BY THE COLONIZATION AND EXTINCTION RATES AND THE PROPORTION X OF OCCUPIED PATCHES
C CORRESPONDS TO +_ IN EQNS AND
,ET X .4 BE THE PROPORTION OF PATCHES THAT ARE OCCUPIED 4HEN THE ABOVE
EQUATION CAN BE REWRITTEN IN TERMS OF X AS
DX
+_ n X n EX
DT
WHERE
_ MX
)T CAN BE SEEN FROM THESE EQUATIONS THAT THERE IS AN EQUILIBRIUM WHERE
THE RATES OF EXTINCTION AND COLONIZATION ARE EQUAL AT WHICH A CERTAIN PROPORTION
X
n E+M OF PATCHES ARE OCCUPIED ,EVINS 4HE EQUILIBRIUM IS STABLE
SINCE THE FRACTION OF OCCUPIED PATCHES WILL ALWAYS TEND TOWARDS THIS EQUILIB
RIUM 4O SEE THAT THE EQUILIBRIUM IS STABLE NOTE THAT IF WE WRITE EQUATION AS
DXDT FX THEN F vX
%VEN THIS VERY SIMPLE MODEL CAN BE USED TO ILLUMINATE THE CONDITIONS CAUSING
SUCCESSFUL VERSUS FAILED INVASIONS 4O SEE HOW AN INVADER WILL DO WHEN IT IS INTRO
DUCED TO AN EMPTY PATCH NETWORK WE FOCUS ON VERY LOW X VALUES THIS IS USUALLY A
REASONABLE ASSUMPTION REGARDING THE INITIAL CONDITIONS OF NATURAL OR INADVERTENT
INVASIONS BUT IT MAY NOT ALWAYS BE APPROPRIATE FOR DELIBERATE LARGE
SCALE INTRO
DUCTIONS &ROM EQUATIONS AND
DX
+M n E FOR SMALL X
X DT
4HUS THE SPECIES CAN INVADE IF +M E )F INVASION IS POSSIBLE THE INVADING SPE
CIES WILL INCREASE ITS PRESENCE UNTIL THE EQUILIBRIUM SIZE X
IS REACHED SEE "OX
FOR A LIST OF MODEL PARAMETERS
+# (ARDING *- -C.AMARA AND 2$ (OLT
"OX
,EVINS MODEL DEFENITIONS
4 THE NUMBER OF HABITAT PATCHES AVAILABLE
. THE NUMBER OF HABITAT PATCHES OCCUPIED BY THE STUDY SPECIES
M RATE OF MIGRATION FROM AN OCCUPIED PATCH
_ THE RATE OF ARRIVAL OF IMMIGRANTS TO EACH OF THE 4 PATCHES
+ THE PROBABILITY THAT A SINGLE EMPTY PATCH IS COLONIZED IE CONVERTED
FROM EMPTY TO OCCUPIED BY A RESIDENT POPULATION BY A SINGLE IMMIGRANT
E RATE AT WHICH AN OCCUPIED PATCH GOES EXTINCT
/VERALL RATE OF COLONIZATION +_4 n .
/VERALL RATE OF EXTINCTION E.
X
EQUILIBRIUM PATCH OCCUPANCY WHERE RATE OF COLONIZATION AND RATE OF
EXTINCTION ARE EQUAL
(ABITAT LOSS MODEL
5 THE NUMBER OF UNSUITABLE PATCHES
H FRACTION OF THE LANDSCAPE THAT IS HABITABLE FOR THE STUDY SPECIES
'ENERALIZED MODEL
%_ SINGLE PATCH EXTINCTION RATE AS A FUNCTION OF IMMIGRATION
#_ SINGLE PATCH COLONIZATION RATE AS A FUNCTION OF IMMIGRATION
2ESCUE EFFECT IMMIGRATION INTO A PATCH LOWERS PATCH EXTINCTION RISK
!NTI
RESCUE EFFECT IMMIGRATION INTO A PATCH INCREASES PATCH EXTINCTION
RISK
!LLEE EFFECT COLONIZATION SUCCESS IS A NON
LINEAR FUNCTION OF IMMIGRATION
ACCELERATING WITH INCREASING IMMIGRATION RATE
!NTI
!LLEE EFFECT NON LINEAR COLONIZATION SUCCESS DECELERATING WITH
INCREASING IMMIGRATION RATE
-ODEL WITH TWO DISTINCT HABITATS
HI THE FRACTION OF A LANDSCAPE THAT IS COVERED BY TYPE I HABITAT PATCHES
PI THE FRACTION OF THE LANDSCAPE THAT IS BOTH OF HABITAT TYPE I AND OCCUPIED
EI EXTINCTION RATE OF TYPE I PATCHES
CIJ COLONIZATION ONTO EMPTY PATCHES OF TYPE I DUE TO MIGRANTS GENERATED
BY OCCUPIED PATCHES OF TYPE J
hI RATE OF INCREASE IN PATCH OCCUPANCY OF SPECIES WHEN IT IS RARE AND ONLY
PATCH TYPE I IS PRESENT
)NVASIVE SPECIES IN METAPOPULATION SYSTEMS
"OX
4WO SPECIES METAPOPULATION MODEL
. THE NUMBER OF PATCHES OCCUPIED BY SPECIES
." THE NUMBER OF PATCHES OCCUPIED BY BOTH SPECIES
0ATCH STATES %MPTY 4YPE 4YPE 4YPE "
M MIGRATION RATE OF SPECIES MIGRANTS FROM A 4YPE PATCH
M MIGRATION RATE OF SPECIES MIGRANTS FROM A 4YPE " PATCH
#_ COLONIZATION RATE OF AN EMPTY PATCH BY SPECIES IMMIGRANTS
#_ COLONIZATION RATE OF A 4YPE PATCH BY SPECIES IMMIGRANTS
%_ EXTINCTION RATE OF A 4YPE PATCH
%_ EXTINCTION RATE OF SPECIES IN A 4YPE " PATCH
"IOLOGICAL CONCLUSIONS FROM ,EVINS MODEL
&AILED INVASIONS CAN ARISE EITHER FROM PROCESSES THAT AFFECT EXTINCTION OR FROM THE
DETAILS OF THE PROCESS OF COLONIZATION 4HERE ARE MANY REASONS EXTINCTION RATES
MAY BE HIGH IN A NOVEL ENVIRONMENT )F WITHIN EACH PATCH THE INTRINSIC GROWTH
RATE R OF THE INVADER IS NEGATIVE IE R SO BIRTHS DEATHS THEN EXTINCTION
IS INEVITABLE UNLESS THERE IS VERY RAPID EVOLUTION SEE 'OMULKIEWICZ AND (OLT
4HIS COULD ARISE BECAUSE OF UNFAVORABLE ABIOTIC CONDITIONS OR BECAUSE
SUPERIOR COMPETITORS OR EFFECTIVE PREDATORS ARE PRESENT "UT EVEN IF R IF A PATCH
HAS A VERY LOW LOCAL CARRYING CAPACITY IT MAY SUFFER HIGH RATES OF EXTINCTION DUE
TO DEMOGRAPHIC STOCHASTICITY IF IT HAS A LOW R IT MAY ALSO NOT BE ABLE TO RECOVER
RAPIDLY FROM CATASTROPHES AND SO RANDOMLY WALK TO EXTINCTION %NVIRONMENTS
WITH FREQUENT CATASTROPHES ARE LIKELY TO GENERATE HIGH EXTINCTION RATES FOR INVASIVE
SPECIES SO ALL ELSE BEING EQUAL ARE NOT AS LIKELY TO BE INVADED
"UT EQUAL WEIGHT SHOULD BE GIVEN TO CONSTRAINTS ON THE COLONIZATION PROCESS AS
POTENTIAL CAUSES OF FAILED INVASIONS )N THE TERMS OF THE BASIC MODEL IF M IS REDUCED
SO IS THE LIKELIHOOD OF INVASION 4HE QUANTITY M COMBINES TWO THINGS A PER CAPITA
RATE OF EMIGRATION FROM AN OCCUPIED PATCH AND THE NUMBER OF INDIVIDUALS PRESENT
THERE AND AVAILABLE TO EMIGRATE 3PECIES THAT ARE SEDENTARY OR PHILOPATRIC ARE
LIKELY TO HAVE A LOW M 3UCH SPECIES SHOULD TEND TO BE POOR INVADERS &OR SPECIES
WITH EQUAL PER CAPITA RATES OF EMIGRATION THOSE SPECIES WITH THE HIGHEST ABUN
DANCE WITHIN OCCUPIED PATCHES CAN GENERATE A HIGH ABUNDANCE OF MIGRANTS AND
+# (ARDING *- -C.AMARA AND 2$ (OLT
SO EXPERIENCE THE HIGHEST COLONIZATION RATE INTO EMPTY PATCHES ,ARGE M SHOULD
THUS REmECT BOTH HIGH FECUNDITY AND HIGH SURVIVAL OF EMIGRANTS ACROSS THE MATRIX
HABITAT SEPARATING HABITABLE PATCHES 4HE PROBABILITY OF SUCCESSFUL COLONIZATION
+ ALSO VARIES GREATLY AMONG SPECIES &OR INSTANCE A HIGH + CAN BE EXPECTED IN
SPECIES WHERE THE MIGRANTS HAVE HIGH FECUNDITY SO THAT ONE OR A FEW IMMIGRANTS
CAN ESTABLISH A SUCCESSFUL POPULATION BY QUICKLY PRODUCING A LARGE NUMBER OF
OFFSPRING THEREBY AVOIDING INITIAL DEMOGRAPHIC STOCHASTICITY AND WHEN INDI
VIDUALS ARE LONG
LIVED "IRTH
DEATH PROCESS MODELS EG AS IN -AC!RTHUR AND
7ILSON REVEAL THAT IN COMPARING SPECIES WITH A GIVEN INTRINSIC GROWTH RATE
R THE PROBABILITY OF EXTINCTION IS LOWER FOR SPECIES WITH LOWER ABSOLUTE DEATH
RATES #ONVERSELY SPECIES WHICH ARE AT RISK OF FREQUENT STOCHASTIC MORTALITY EVENTS
ARE LIKELY TO HAVE LOW INTRINSIC COLONIZATION PROPENSITY AND SO HAVE A LOW +
)MMIGRANTS ARE LIKELY TO BE UNFAMILIAR WITH THE DETAILS OF LOCAL RESOURCES REFUGES
AND MORTALITY RISKS AND IF LEARNING NEW CONDITIONS IS SLOW OR UNRELIABLE IMMI
GRANTS MAY BE VULNERABLE TO RAPID ELIMINATION IN NOVEL ENVIRONMENTS
/NE MATHEMATICALLY SLIGHT BUT BIOLOGICALLY SIGNIlCANT GENERALIZATION OF THE
,EVINS MODEL SUGGESTED BY ,ANDE AND .EE AND -AY IS TO RECOGNIZE
THAT IN A LANDSCAPE NOT ALL EMPTY PATCHES ARE SUITABLE )F WE ASSUME THAT OUT OF
4 PATCHES 5 ARE UNSUITABLE THEN H n54 IS THE FRACTION OF THE LANDSCAPE THAT
IS POTENTIALLY OCCUPIABLE BY THE SPECIES 7ORKING BACK THROUGH THE SAME STEPS AS
BEFORE WE END UP WITH DXDT +_H
X
EX 4HE INVASION CRITERION NOW BECOMES
+MH E !S NOTED FOR A SOMEWHAT DIFFERENT VERSION OF THE MODEL IN (OLT AND +EITT
THIS SIMPLE FORMULATION OF INVASION ENCOMPASSES A GREAT DEAL OF BIOLOGY
)N ADDITION TO THE MECHANISMS MENTIONED IN THE PREVIOUS PARAGRAPHS ANOTHER
REASON WHY AN INVASION COULD FAIL IS THAT INSUFlCIENT SUITABLE HABITAT IS AVAILABLE
%VEN IF EACH SUITABLE PATCH COULD SUSTAIN A POPULATION ON ITS OWN OVER A REASON
ABLE TIME SCALE IF THERE ARE TOO FEW OF THEM COLONIZATION MAY NOT BE ABLE TO MATCH
EVEN A LOW RATE OF EXTINCTION (OLT AND +EITT SHOW HOW THESE IDEAS PERMIT
ONE TO CONSIDER IN A UNIlED FASHION MANY DIFFERENT HYPOTHESES THAT HAVE BEEN PUT
FORTH REGARDING THE FACTORS CAUSING DISTRIBUTIONAL RANGE LIMITS
4HE THEORETICALLY BEST INVADER WOULD HAVE LOW EXTINCTION RISK AND HIGH M
4HE THEORETICAL @BEST IS LIKELY TO BE UNFEASIBLE BECAUSE THERE ARE BIOLOGICAL
CONSTRAINTS SUCH AS TRADEOFFS BETWEEN LIFE HISTORY TRAITS EG REPRODUCTION AND
LONGEVITY IN ANIMALS AND BETWEEN DEVELOPMENT OF VEGETATIVE PROPAGULES AND
SEED PRODUCTION IN PLANTS
$YNAMIC ,EVINS TYPE METAPOPULATION
4HE ,EVINS MODEL CLARIlES MANY ESSENTIAL FEATURES OF METAPOPULATION DYNAMICS
BUT IT RESTS ON QUITE SIMPLE ASSUMPTIONS ABOUT HOW MIGRATION INmUENCES BOTH
COLONIZATION AND EXTINCTION (ANSKI 4HE BASIC ,EVINS FRAMEWORK HAS BEEN
EXTENDED TO SEVERAL SPECIlC CASES IN DIFFERENT STUDIES INCLUDING FOR EXAMPLE A LIN
EAR RESCUE EFFECT (ANSKI 'OTELLI AND +ELLY AND A SPECIlC NONLINEAR
!LLEE EFFECT (ANSKI 4HE ,EVINS MODEL WAS RECENTLY GENERALIZED TO ALLOW
)NVASIVE SPECIES IN METAPOPULATION SYSTEMS
FOR THESE AND OTHER DYNAMIC INmUENCES FROM MIGRATION ON BOTH COLONIZATION
AND EXTINCTION RATES COMPARE &IGS AND (ARDING AND -C.AMARA
4HE GENERALITY IS OBTAINED BY EXPRESSING BOTH COLONIZATION AND EXTINCTION RATES AS
FUNCTIONS OF IMMIGRATION RATE AS WE NOW DESCRIBE
&IG !N EXTENSION OF ,EVINS MODEL &IG (ERE THE RATE OF COLONIZATION AND
EXTINCTION ARE NOT lXED CONSTANTS BUT CAN BE ANY FUNCTION OF IMMIGRATION RATE _ 4HIS
MODEL CAN THEREFORE CAPTURE SEVERAL TYPES OF SINGLE SPECIES DYNAMICS SUCH AS !LLEE EFFECTS
RESCUE EFFECTS AND ANTI
RESCUE EFFECTS (ARDING AND -C.AMARA
)T IS USEFUL TO THINK OF METAPOPULATION PROCESSES AT TWO SCALES 4HE SMALLEST
SCALE IS THE PATCH 4HE SUBPOPULATION INHABITING A PATCH HAS A GIVEN RISK OF LOCAL
EXTINCTION %_ THAT MAY BE INmUENCED BY THE RATE OF IMMIGRATION _ TO THE PATCH
%MPTY PATCHES ARE COLONIZED AT RATE #_ THAT VERY LIKELY DEPENDS ON _ 'IVEN
THESE @LOCAL RATES OF CHANGE THE OVERALL RATE OF CHANGE IN THE PROPORTION X .4
OF PATCHES THAT ARE OCCUPIED IN THE WHOLE METAPOPULATION IS
DX
#_ n X n %_ X
DT
(ARDING AND -C.AMARA (ERE WE AGAIN ASSUME THAT ALL PATCHES ARE
POTENTIALLY OCCUPIABLE 4HE ,EVINS MODEL IS THEN A SPECIAL CASE OF THIS MODEL WITH
EXTINCTION FUNCTION %_ E AND COLONIZATION FUNCTION #_ +_ !S DESCRIBED
BELOW A VARIETY OF BIOLOGICAL PROCESSES CAN BE MIMICKED PHENOMENOLOGICALLY WITH
THIS EXTENDED FRAMEWORK
$YNAMIC EXTINCTION FUNCTIONS
4HE ORIGINAL ,EVINS EXTINCTION FUNCTION HAS A CONSTANT VALUE E THAT IS INDEPEN
DENT OF IMMIGRATION RATE )N A METAPOPULATION IT CAN BE IMPORTANT TO ALLOW THE
EXTINCTION RATE OF A PATCH TO DEPEND ON IMMIGRATION INTO THAT PATCH )MMIGRANTS
CAN FOR EXAMPLE LOWER PATCH EXTINCTION RISK BY PROVIDING A DEMOGRAPHIC CONTRI
BUTION TO DECLINING POPULATIONS OR BY INTRODUCING NEW GOOD GENES EG REDUCING
INBREEDING DEPRESSION 3ACCERI ET AL THIS IS THE RESCUE EFFECT "ROWN AND
+ODRIC
"ROWN (ANSKI &OLLOWING A CATASTROPHE IMMIGRANTS CAN
+# (ARDING *- -C.AMARA AND 2$ (OLT
SPEED THE RECOVERY OF THE POPULATION ALLOWING IT TO MORE RAPIDLY REBOUND FROM
DANGEROUSLY LOW DENSITIES $EPENDING ON THE UNDERLYING PROCESSES LEADING TO A
RESCUE EFFECT THE EXTINCTION FUNCTION CAN HAVE DIFFERENT SHAPES &IG
&IG )LLUSTRATION OF RESCUE EFFECT FUNCTIONS 4HE RATE OF PER PATCH EXTINCTION CAN
SOMETIMES DECREASE AS IMMIGRATION RATE INCREASES 4HREE DIFFERENT TYPES OF RESPONSE TO
INCREASED IMMIGRATION ARE ILLUSTRATED
(OWEVER IN OTHER CIRCUMSTANCES IMMIGRATION CAN LEAD TO INCREASED EXTINCTION
RISK WE TERM THIS PATTERN AN ANTI
RESCUE EFFECT 4HIS CAN FOR EXAMPLE BE DUE TO
IMMIGRANTS CARRYING PARASITES OR DISEASES OR BECAUSE IMMIGRANTS CARRY DIFFERENT
GENES BREAKING UP LOCALLY ADAPTED BENElCIAL GENE COMBINATIONS 2OLN
!LVAREZ
ET AL )N PREDATOR
PREY SYSTEMS INCREASING IMMIGRATION CAN SOMETIMES
DESTABILIZE LOCAL INTERACTIONS MAKING EXCURSIONS TO LOW DENSITIES AND HENCE LOCAL
EXTINCTIONS MORE LIKELY 3CHPS (OLT SEE REVIEW IN (OOPES ET AL
7ITH ALL THESE MECHANISMS INCREASING IMMIGRATION RATE OVER SOME RANGE
OF VALUES IS LIKELY TO BOOST LOCAL EXTINCTION RATES
$YNAMIC COLONIZATION FUNCTIONS
)N ,EVINS MODEL THE COLONIZATION RATE PER PATCH INCREASES LINEARLY WITH THE IMMI
GRATION RATE AND HENCE INCREASES LINEARLY WITH THE NUMBER OF OCCUPIED PATCHES
)NVASIVE SPECIES IN METAPOPULATION SYSTEMS
(OWEVER COLONIZATION MIGHT TAKE OTHER FORMS AS WELL )T IS COMMON FOR A SPECIES
TO HAVE PROBLEMS IN INITIAL ESTABLISHMENT IN A PATCH WHEN THE NUMBER OF IMMI
GRANTS IS LOW DUE TO DEMOGRAPHIC SKEWS LACK OF ADULTS OR MALES FOR EXAMPLE DIF
lCULTY IN lNDING MATES OR INBREEDING 4HE PHENOMENA OF DISPROPORTIONATELY LOW
COLONIZATION SUCCESS WHEN THE IMMIGRATION RATE IS LOW IS AN EXAMPLE OF AN !LLEE
EFFECT !LLEE (ANSKI
4HE OPPOSITE WHEN COLONIZATION BY A SINGLE MIGRANT IS MORE SUCCESSFUL
WHEN THERE ARE FEW OTHER MIGRANTS IS TERMED AN ANTI
!LLEE EFFECT (ARDING AND
-C.AMARA )T CAN OCCUR IF MIGRANTS DO NOT SETTLE AT RANDOM BUT AVOID CON
SPECIlCS EG TERRITORIALITY SO THAT MIGRANTS ACTIVELY SEEK UNOCCUPIED PATCHES
4HIS LEADS TO A VERY RAPID COLONIZATION INITIALLY BUT AFTER A WHILE MOST PATCHES
ARE OCCUPIED AND THE EFFECT SHOULD BECOME LESS IMPORTANT )T IS ALSO CONCEIVABLE
THAT IF RESOURCES ARE EXPLOITED WHICH HAVE LOW RATES OF RENEWAL AN INCREASE IN THE
NUMBER OF IMMIGRANTS COULD LEAD TO RAPID OVEREXPLOITATION RESOURCE DEPLETION
AND THUS A REDUCED PROBABILITY OF SUCCESSFUL ESTABLISHMENT "OTH THESE MECHA
NISMS CAN LEAD TO AN ANTI
!LLEE EFFECT AT THE METAPOPULATION LEVEL
)MPACTS BY RESIDENT GENERALIST PREDATORS UPON COLONIZATION BY AN INVASIVE PREY
SPECIES ARE LIKELY TO SHIFT BETWEEN ANTI
!LLEE EFFECTS AND !LLEE EFFECTS DEPENDING ON
THE QUANTITATIVE DETAILS OF THE PREDATORS FUNCTIONAL RESPONSE AND THE NUMBER OF
IMMIGRANTS )F A RESIDENT PREDATOR HAS A TYPE ))) SIGMOIDAL FUNCTIONAL RESPONSE
THEN OVER A RANGE OF LOW PREY NUMBERS AN INCREASE IN PREY DENSITY DUE TO AN
INCREASING NUMBER OF MIGRANTS LEADS TO AN INCREASE IN THE PER CAPITA MORTALITY
RATE INmICTED ON THE PREY EG BECAUSE THE PREDATOR BECOMES INCREASINGLY AWARE
OF THE NOVEL PREY WITH INCREASING EXPOSURE TO IT AND SO IS MORE LIKELY TO CAUSE PREY
EXTINCTION WITH INCREASING NUMBERS OF COLONISTS 4HIS CAN BE VIEWED AS AN ANTI
!LLEE EFFECT WHICH COULD BROADLY DESCRIBE ANY KIND OF NEGATIVE DENSITY DEPEND
ENCE OPERATING AT LOW DENSITIES WHICH CAN NEGATIVELY AFFECT COLONIZATION SUCCESS
"UT TYPICALLY AT HIGHER PREY DENSITIES MOST PREDATORS HAVE SATURATING FUNCTIONAL
RESPONSES EG DUE TO LIMITED HANDLING TIME OR GUT CAPACITY SO THE PER CAPITA
MORTALITY RATE EXPERIENCED BY PREY DECLINES WITH INCREASING PREY DENSITY )N THIS
RANGE AN INCREASING NUMBER OF MIGRANTS MAY ENHANCE COLONIZATION BY DILUTING
THE IMPACT OF PREDATION 3O WHETHER OR NOT A COLONISING SPECIES EXPERIENCES AN
!LLEE OR AN ANTI
!LLEE EFFECT DUE TO RESIDENT PREDATORS WILL DEPEND UPON BOTH THE
PREDATORS FUNCTIONAL RESPONSE AND THE NUMBER OF MIGRANTS
7HO IS A GOOD INVADER IN THE SINGLE SPECIES SYSTEM
4O ANALYSE WHETHER A SPECIES CAN INVADE INTO A PATCHY LANDSCAPE WE SUPPOSE
THAT THE SPECIES HAS BEEN INTRODUCED IN A SMALL NUMBER OF PATCHES 7E CAN THEN
ASK WHETHER THE NUMBER OF OCCUPIED PATCHES TENDS TO GROW OR DECLINE )N TERMS
OF THE GENERALIZED METAPOPULATION MODEL DESCRIBED ABOVE WE ARE CONCERNED WITH
WHETHER DXDT WHEN X IS SMALL 4HIS DEPENDS ON THE BEHAVIOUR OF THE COLONIZA
TION AND EXTINCTION FUNCTIONS FOR SMALL X AND HENCE SMALL _ 3UPPOSE THAT
+# (ARDING *- -C.AMARA AND 2$ (OLT
#_ 5 +_ FOR SMALL _
AND
%_ 5 E FOR SMALL _
4HUS FOR SMALL NUMBERS OF OCCUPIED PATCHES THE METAPOPULATION BEHAVES AS
DESCRIBED BY ,EVINS MODEL WITH PARAMETERS + AND E ALTHOUGH IT MAY BEHAVE VERY
DIFFERENTLY WHEN THE NUMBER OF OCCUPIED PATCHES INCREASES 7ITH THIS ASSUMPTION
APPROXIMATION IS STILL VALID SO THAT THE SPECIES CAN INVADE IF +M E
7HEN DOES AN INVASIVE SPECIES GET ESTABLISHED IN A SINGLE SPECIES
METAPOPULATION
&IGURE A F ILLUSTRATE THE EXTINCTION RATE WHITE SURFACE AND COLONIZATION RATE BLACK
@HILLS FOR SINGLE SPECIES METAPOPULATIONS )T SHOWS HOW DIFFERENT COMBINATIONS OF
#_ AND %_ INmUENCE THE EQUILIBRIA WHERE THE TWO SURFACES MEET $EPENDING
ON THE FUNCTIONS #_ AND %_ THERE MAY EXIST AT LEAST ONE NON TRIVIAL EQUILIBRIA
IN PATCH OCCUPANCY X
SUCH THAT TOTAL COLONIZATION AND EXTINCTION RATES BAL
ANCE WHEN A PROPORTION X
OF PATCHES ARE OCCUPIED )N SOME CASES THERE MAY BE
MORE THAN ONE SUCH EQUILIBRIUM IE FOR A GIVEN M THE TWO RATES ARE EQUAL AT TWO
OR THREE DIFFERENT X SEE &IG B C E F (ARDING AND -C.AMARA 4HE EXACT
SHAPES OF #_ AND %_ ARE IMPORTANT ALSO FOR THE EXTINCTION RISK OF THE ENTIRE
METAPOPULATION AT LOW PATCH NUMBERS &IG 4HE OCCURRENCE OF MULTIPLE EQUI
LIBRIA HAS EARLIER BEEN STUDIED IN MORE COMPLEX METAPOPULATION MODELS (ANSKI
AND 'YLLENBERG 'YLLENBERG ET AL )F AT LEAST ONE EQUILIBRIUM EXISTS
THEN THERE IS AT LEAST ONE STABLE EQUILIBRIUM (ERE BY @STABLE WE MEAN THAT IF X IS
INITIALLY CLOSE TO X
THEN THE PROPORTION OF OCCUPIED PATCHES WILL TEND TO X
OVER
TIME )N ,EVINS MODEL THERE IS A UNIQUE EQUILIBRIUM THAT IS STABLE IF +M E &IG
A AND THERE IS NO EQUILIBRIUM IF +M ) E 4HUS IN THIS PARTICULAR MODEL A STABLE
EQUILIBRIUM EXISTS IF AND ONLY IF THE SPECIES CAN INVADE
&OR OTHER SHAPES OF #_ AND %_ IT IS ALSO TRUE THAT IF A SPECIES CAN INVADE
THEN THE PROPORTION OF OCCUPIED PATCHES WILL CERTAINLY INCREASE AND TEND TO A
STABLE EQUILIBRIUM )N GENERAL HOWEVER THE CONVERSE NEED NOT HOLD AND THE
EXISTENCE OF AN EQUILIBRIUM IS NOT EQUIVALENT TO THE ABILITY TO INVADE 4O SEE THIS
SUPPOSE THAT THERE IS AN !LLEE EFFECT 4HUS THE PATCH COLONIZATION FUNCTION #_
IS AN ACCELERATING FUNCTION OF _ FOR SMALL _ ,ET THE PATCH EXTINCTION FUNCTION BE
AS IN ,EVINS MODEL IE %_ E A CONSTANT 4HEN FOR SUITABLE CHOICE OF E THE
TOTAL EXTINCTION RATE EXCEEDS THE TOTAL COLONIZATION RATE FOR SMALL X BUT COLONIZA
TION EXCEEDS EXTINCTION IF X IS SOMEWHAT LARGER &IG E 4HUS THE SPECIES CANNOT
INVADE HOWEVER IF THE PROPORTION OF PATCHES OCCUPIED WERE ARTIlCIALLY INCREASED
SO AS TO EXCEED SOME CRITICAL LEVEL THEN THE PROPORTION WOULD CONTINUE TO INCREASE
AND TEND TO A STABLE EQUILIBRIUM
)NVASIVE SPECIES IN METAPOPULATION SYSTEMS
&IG ! RANGE OF METAPOPULATIONS WITH DIFFERENT DYNAMICS OF THE MODEL IN &IG 7HITE
SURFACES ARE TOTAL EXTINCTION RISKS AND BLACK HILLS TOTAL COLONIZATION RATES A !N ORDINARY
,EVINS METAPOPULATION B ! RESCUE EFFECT MAKES EXTINCTION RISK DECLINE WITH HIGHER
MIGRATION RATES C ! DIFFERENT KIND OF RESCUE EFFECT D !NTI RESCUE EFFECT LEADS TO HIGHER
EXTINCTION RISK WITH HIGHER MIGRATION RATES E 4HE COLONIZATION FUNCTION IS BELL
SHAPED
AN !LLEE EFFECT PREVAILS F !N EXAMPLE WITH BOTH A RESCUE AND !LLEE EFFECT
(ETEROGENEOUS LANDSCAPES
(OLT CONSIDERED A GENERALIZATION OF THE STANDARD ,EVINS METAPOPULATION
MODEL TO A LANDSCAPE WITH TWO DISTINCT HABITATS ! FRACTION HI OF THE LANDSCAPE
IS COMPRISED OF PATCHES OF TYPE I 4HE FRACTION OF HABITAT PATCHES THAT ARE TYPE
I AND OCCUPIED BY THE SPECIES IS DENOTED BY PI .ECESSARILY WE HAVE PI ) HI AND
HI H ) )F IN THE LATTER CASE THE INEQUALITY HOLDS HI H SOME PATCHES IN
THE LANDSCAPE ARE UNSUITABLE FOR OCCUPANCY 5SING THE STANDARD ,EVINS NOTATION
EI DENOTES EXTINCTION ON OCCUPIED PATCHES OF TYPE I AND CIJ IS COLONIZATION ONTO
EMPTY PATCHES OF TYPE I DUE TO MIGRANTS GENERATED BY OCCUPIED PATCHES OF TYPE J
4HE EQUATION FOR DYNAMICS OF P IS DPDT C P C P H n P n E P A SIMILAR
EQUATION DESCRIBES DYNAMICS IN THE SECOND HABITAT TYPE
+# (ARDING *- -C.AMARA AND 2$ (OLT
4HE TOTAL OCCUPANCY BY THE SPECIES IN THE LANDSCAPE IS P P P )F WE SUM
THE DYNAMICAL EQUATIONS FOR CHANGE IN OCCUPANCY WE GET AN EXPRESSION FOR DPDT
)N GENERAL THIS DOES NOT SIMPLIFY TO A FORM IN JUST P (OWEVER IF ALL THE COLONIZA
TION RATES CIJ ARE EQUAL TO A CONSTANT C AND BOTH PATCHES HAVE THE SAME EXTINCTION
RATES THE ORIGINAL ,EVINS MODEL EMERGES )N OTHER WORDS ARBITRARY DISTINCTIONS
AMONG HABITATS CAN SIMPLY BE IGNORED (OWEVER IF PATCH TYPE INmUENCES EITHER
COLONIZATION OR EXTINCTION LANDSCAPE HETEROGENEITY CAN INmUENCE METAPOPULA
TION DYNAMICS EG PATCH TYPES DIFFER IN + OR M
!LL THE FACTORS MENTIONED ABOVE WHICH INmUENCE COLONIZATION AND EXTINCTION
RATES CAN POTENTIALLY SHOW VARIATION AMONG HABITAT TYPES &OR INSTANCE AN INDI
VIDUAL BORN IN A HABITAT PATCH OF TYPE I BEFORE EMIGRATING MIGHT BECOME ACCLIMA
TIZED TO THE ABIOTIC CONDITIONS IN ITS NATAL SITE EG AS IN PHYSIOLOGICAL ADAPTATION
TO THERMAL CONDITIONS AND SO EITHER PREFERENTIALLY SEEK OUT SIMILAR HABITATS WHEN
IT DISPERSES OR BE VULNERABLE TO HIGH MORTALITY IF IT SETTLES INTO A DIFFERENT HABITAT
LEADING TO FAILED COLONIZATION
)F ONLY PATCH TYPE I IS PRESENT THE INITIAL GROWTH RATE OF THE SPECIES WHEN RARE
IS hI CII HI n EI AS IN THE STANDARD ,EVINS MODEL 7ITH BOTH PATCH TYPES PRESENT
THE ASYMPTOTIC GROWTH RATE OF THE INVADING SPECIES TURNS OUT TO BE h h
h 3 ;h n h C C H H= (OLT
-ANIPULATING THIS EXPRESSION LEADS TO SEVERAL SIMPLE CONCLUSIONS &IRST A SPE
CIES WHICH IS A GENERALIST MAY BE ABLE TO INVADE AND PERSIST BECAUSE IT CAN ENJOY
CROSS
HABITAT COLONIZATION WHEREAS OTHERWISE SIMILAR HABITAT SPECIALISTS WOULD
GO EXTINCT IE EXPERIENCE FAILED INVASIONS 3ECOND SOMETIMES UTILIZING A SECOND
HABITAT MAY BE CRUCIAL IN PERMITTING A SPECIES TO PERSIST IN A LANDSCAPE EVEN IF
THERE IS NO COLONIZATION AMONG PATCHES OF THIS SECOND TYPE CONDITION IN (OLT
4HIS CAN HAPPEN IF ALL COLONIZATION FROM AND OF THE SECOND HABITAT TYPE
IS INTO AND FROM HABITATS OF THE lRST HABITAT TYPE EG BECAUSE PATCHES OF THE
SECOND HABITAT TYPE ARE WIDELY SPACED BUT THE SECOND HABITAT TYPE HAS A LOWER
EXTINCTION RATE THAN DOES THE PRIMARY HABITAT TYPE 4HIS CAN BE VIEWED AS A KIND
OF SPATIAL STORAGE EFFECT )T WOULD BE INTERESTING IN FUTURE WORK TO EXTEND THIS TWO
HABITAT MODEL USING THE GENERALIZED APPROACH OF (ARDING AND -C.AMARA
#OLONIZATION AMONG SOME HABITATS MAY PERMIT RESCUE EFFECTS EVEN WHILE COLONI
ZATION AMONG OTHERS RESEMBLES THE ANTI
RESCUE PATTERN )N SUCH CASES THE EXPECT
ED DYNAMICS OF INVASION WOULD BE QUITE SENSITIVE TO THE RELATIVE PROPORTIONS OF
THE LANDSCAPE OCCUPIED BY THE DISTINCT HABITAT TYPES
).4%2!#4)/.3 7)4( ! .!4)6% 30%#)%3
!N INVASIVE SPECIES CAN INTERACT WITH A NATIVE METAPOPULATION IN THE SAME PATCH
NETWORK IN VARIOUS WAYS WITH NET EFFECTS ON THE LIKELIHOOD OF INVASION RANGING
FROM POSITIVE TO NEGATIVE &OR INSTANCE WHEN SPECIES COMPETE FOR THE SAME LIM
ITING RESOURCE WITHIN PATCHES COMPETITIVE EXCLUSION IS LIKELY -ETAPOPULATION
DYNAMICS CAN NONETHELESS PERMIT INVASION BY AN INFERIOR SPECIES IF IT IS SUPERIOR
)NVASIVE SPECIES IN METAPOPULATION SYSTEMS
AT COLONIZATION OF EMPTY PATCHES ,EVINS AND #ULVER 4ILMAN
4HIS REQUIRES A TRADEOFF BETWEEN COMPETITION AND COLONIZATION ALLOWING THE INFE
RIOR COMPETITOR TO EXPLOIT MORE EFFECTIVELY THE EMPTY HABITAT PATCHES LEFT BY THE
SUPERIOR COMPETITOR WHEN IT SUFFERS EXTINCTIONS !MARSEKARE
-ETAPOPULATION DYNAMICS CAN ALSO LEAD TO EXCLUSION THAT WOULD OTHERWISE NOT
OCCUR (OLT &OR INSTANCE ONE SPECIES CAN DELIMIT A SECOND SPECIES INDI
RECTLY BY HOSTING A PATHOGEN THAT IS MORE HARMFUL TO THE SECOND SPECIES (OLT AND
,AWTON !N EXAMPLE IS THE GRADUAL ERADICATION OF THE NATIVE NOBLE CRAY
lSH !STACUS ASTACUS IN 3WEDEN WHICH IS BEING REPLACED BY THE .ORTH !MERICAN
SIGNAL CRAY lSH 0ACIFASTACUS LENIUSCULUS MEDIATED BY A SHARED INFECTIOUS DISEASE
4HE INVASIVE SPECIES CARRIES A FUNGUS !PHANOMYCES ASTACI TO WHICH IT ITSELF APPEARS
TO BE IMMUNE WHEREAS THE NATIVE SPECIES IS SEVERELY IMPACTED BY THE PARASITE
"ANGYEEKHUN )N A METAPOPULATION SUCH INDIRECT EXCLUSION MAY OCCUR
EVEN IF THE TWO SPECIES OCCUPY DISTINCT HABITAT PATCHES IN THIS EXAMPLE IF FUNGAL
SPORES DISPERSE WIDELY THE INVASIVE SPECIES COULD PROVIDE A LANDSCAPE RESERVOIR
FOR THE PATHOGEN WHICH CAN ELIMINATE THE NATIVE SPECIES EVEN IN PATCHES WHERE
THE INVASIVE SPECIES ITSELF NEVER OCCURS
4HERE CAN ALSO BE FACILITATIVE INTERACTIONS BETWEEN INVASIVE AND RESIDENT SPE
CIES )N THE LITERATURE ON SUCCESSION THERE ARE MANY EXAMPLES OF EARLY COLONIZERS
FACILITATING INVASION BY LATER COLONISTS EG NITROGEN
lXERS MAY NEED TO COLONIZE
PRIOR TO OTHER SPECIES #ONNELL AND 3LATYER )N THIS CASE ONE SPECIES ALTERS
THE ABIOTIC ENVIRONMENT SO AS TO ENHANCE COLONIZATION OR REDUCE EXTINCTION FOR A
SECOND SPECIES -OREOVER ONE SPECIES MAY REQUIRE ANOTHER SPECIES AS A RESOURCE
)NVASIONS BY SPECIALIST PREDATORS PARASITES OR HERBIVORES WILL ALMOST ALWAYS
DEPEND UPON THE PRESENCE OF THEIR REQUIRED PREY OR HOSTS
! FLEXIBLE MODEL WHICH ALLOWS FOR MANY KINDS OF INTERACTIONS
7E WILL NOW LOOK AT INVASIONS IN WHICH EACH OF TWO SPECIES HAS INTERLINKED META
POPULATION DYNAMICS EXTENDING A PHENOMENOLOGICAL METAPOPULATION MODEL OF
(ANSKI AND OTHERS &IG TO INCLUDE DIFFERENT TYPES OF COLONIZATION AND
EXTINCTION FUNCTIONS AND DIFFERENTIAL MIGRATION RATES 7E USE THIS MODEL TO CLASSI
FY DIFFERENT TYPES OF INTERACTIONS BETWEEN AN INVASIVE SPECIES AND A NATIVE SPECIES
7E CONSIDER A METAPOPULATION WITH 4 IDENTICAL PATCHES %ACH PATCH CAN BE IN
ONE OF FOUR STATES %MPTY 4YPE WHEN SPECIES IS PRESENT 4YPE WHEN SPECIES
IS PRESENT OR 4YPE " WHEN BOTH ARE PRESENT &IG
-IGRATION
%ACH 4YPE PATCH SENDS OUT SUCCESSFUL SPECIES MIGRANTS AT RATE M %ACH
4YPE PATCH SENDS OUT SUCCESSFUL SPECIES MIGRANTS AT RATE M %ACH 4YPE "
PATCH SENDS OUT SUCCESSFUL SPECIES MIGRANTS AT RATE M AND SENDS OUT SUC
CESSFUL SPECIES MIGRANTS AT RATE M .OTE THAT THE DIFFERENCE BETWEEN M AND
M MEASURES HOW MUCH SPECIES CAN SUPPRESS OR ENHANCE THE MIGRATION RATE OF
+# (ARDING *- -C.AMARA AND 2$ (OLT
&IG 4HE RISK OF EXTINCTION AT LOW PATCH NUMBERS DEPEND ON THE SHAPE OF THE # AND %
FUNCTIONS SEE &IG &ROM (ARDING AND -C.AMARA
&IG ! METAPOPULATION MODEL FOR TWO SPECIES 3PECIES IS COMPETITIVELY SUPERIOR AND
EXCLUDES SPECIES FROM PATCHES 3PECIES CAN PERSIST IF IT HAS A HIGHER COLONIZATION TO
EXTINCTION RATIO THAN SPECIES .EE AND -AY
)NVASIVE SPECIES IN METAPOPULATION SYSTEMS
&IG ! METAPOPULATION MODEL SPECIlCALLY DESIGNED TO STUDY INVASIVE SPECIES
EQ ! PATCH CAN BE IN ONE OF FOUR STATES EMPTY OCCUPIED BY SPECIES OR SPECIES
OR OCCUPIED BY BOTH SPECIES " 4HIS MODEL ALLOWS US TO EXPLORE MANY SORTS OF INmUENCES
THAT INVASIVE SPECIES CAN HAVE ON A NATIVE SPECIES 4HE EXTINCTION % AND COLONIZATION RATES
# ARE FUNCTIONS OF IMMIGRATION _
SPECIES 3IMILARLY FOR SPECIES THE DIFFERENCE IN MIGRATION PARAMETERS REmECTS
INTERSPECIlC IMPACTS ON THE RATE AT WHICH EMIGRANTS ARE EMITTED FROM JOINTLY
OCCUPIED PATCHES 4HESE DIFFERENCES COULD OCCUR DUE TO CHANGES IN DENSITY WITHIN
PATCHES OR DUE TO CHANGES IN INDIVIDUAL BEHAVIOR IN THE FACE OF THE OTHER SPECIES
EG A PREY SPECIES MAY SPEND MORE TIME IN HIDING IN THE FACE OF A PREDATOR AND
THUS BE LESS LIKELY TO EMIGRATE FROM PATCHES CONTAINING BOTH PREDATORS AND PREY
)NITIALLY WE ASSUME THAT MIGRANTS SETTLE AT RANDOM 3UPPOSE THAT THE NUMBER
OF 4YPE AND 4YPE " PATCHES ARE . AND ." RESPECTIVELY 4HEN SPECIES MIGRANTS
ARRIVE AT EACH PATCH IN THE ENVIRONMENT AT RATE
_ .M ."M 4
4HE RATE OF ARRIVAL OF SPECIES MIGRANTS _ IS SIMILARLY DElNED
#OLONIZATION
%ACH EMPTY PATCH IS COLONIZED BY SPECIES MIGRANTS AND HENCE BECOMES A 4YPE
PATCH AT RATE #_ %MPTY PATCHES ARE COLONIZED BY SPECIES MIGRANTS AND
BECOME 4YPE AT THE RATE #_ %ACH 4YPE PATCH IS COLONIZED BY SPECIES
MIGRANTS AND HENCE BECOMES A 4YPE " PATCH AT RATE #_ 3IMILARLY EACH 4YPE
PATCH IS COLONIZED BY SPECIES MIGRANTS AND IS CONVERTED TO A 4YPE " PATCH AT
THE RATE #_ .OTE THAT WE NEVER HAVE AN EMPTY PATCH SIMULTANEOUSLY COLONIZED
BY BOTH SPECIES AND
+# (ARDING *- -C.AMARA AND 2$ (OLT
%XTINCTION
! 4YPE PATCH CHANGES TO AN EMPTY PATCH AT RATE %_ ! 4YPE PATCH CHANGES
TO AN EMPTY PATCH AT RATE %_ ! 4YPE " PATCH WHICH HAS BOTH SPECIES CHANGES
TO A 4YPE PATCH WHEN SPECIES GOES EXTINCT AT RATE %_ 3IMILARLY A 4YPE "
PATCH CHANGES TO A 4YPE PATCH AT RATE %_ 7E ASSUME THAT IN A PATCH WITH
BOTH SPECIES THERE ARE NOT SIMULTANEOUS EXTINCTIONS TAKING 4YPE " PATCHES
DIRECTLY BACK TO AN EMPTY STATE
)N THIS MODEL THE RATE OF CHANGE OF PATCH STATE DEPENDS ON MIGRATION RATES
FROM BOTH THE TWO SINGLE SPECIES AND THE MIXED SPECIES PATCHES 4HE TWO SPECIES
CAN THUS INmUENCE EACH OTHER BY ALTERED EXTINCTION AND COLONIZATION RATES BOTH
OF WHICH CAN INVOLVE ALTERED MIGRATION RATES SEE &IG
0OSSIBLE INTERACTIONS BETWEEN SPECIES
.OW CONSIDER INTERACTIONS BETWEEN THE TWO SPECIES &IG )F SPECIES COMPETES
WITH SPECIES THEN SPECIES MAY BE ADVERSELY AFFECTED BY THE PRESENCE OF SPECIES
IN VARIOUS WAYS )T MAY BE MORE DIFlCULT FOR SPECIES TO COLONISE A PATCH THAT IS
ALREADY OCCUPIED BY SPECIES THAN TO COLONISE AN EMPTY PATCH IE #_ #_
)F SPECIES CAN COLONISE A PATCH CURRENTLY OCCUPIED BY SPECIES SO AS TO PRODUCE
A PATCH WITH BOTH SPECIES THEN THIS MAY INCREASE THE PROBABILITY THAT SPECIES
GOES EXTINCT ON THE PATCH IE %_ %_ 4HE PRESENCE OF SPECIES MAY ALSO
REDUCE THE RATE AT WHICH SPECIES SENDS OUT MIGRANTS FROM THE PATCH IE M M
4HIS WILL REDUCE THE IMMIGRATION RATE _ WHICH WILL TEND TO REDUCE THE RATE AT
WHICH PATCHES ARE COLONIZED BY SPECIES
)F M M THERE IS A POSSIBLE BENElCIAL EFFECT TO SPECIES FOR SPECIES TO ALSO
BE PRESENT IN A PATCH 'IVEN A STRONG ANTI
RESCUE EFFECT IE INCREASED IMMIGRA
TION INCREASES EXTINCTION RATE A REDUCTION IN THE IMMIGRATION RATE _ MAY REDUCE
THE RATE AT WHICH SPECIES DIES OUT ON PATCHES 7HETHER COMPETITION FROM SPE
CIES IS BENElCIAL OR NOT MAY DEPEND ON THE NUMBER OF PATCHES ALREADY OCCUPIED
BY SPECIES &OR EXAMPLE IF THE ANTI
RESCUE EFFECT ONLY BECOMES MARKED AT HIGH
IMMIGRATION RATES COMPETITION MAY BE DELETERIOUS TO SPECIES WHEN IT IS RARE
BUT BENElCIAL WHEN IT IS COMMON )N A SENSE GIVEN METAPOPULATION DYNAMICS
AND AN ANTI
RESCUE EFFECT A COMPETITOR AS MEASURED BY A REDUCTION IN THE SIZE OF
THE MIGRANT POOL GENERATED BY PATCHES MAY DYNAMICALLY BE A MUTUALIST
)F INSTEAD OF BEING A COMPETITOR SPECIES IS A PREDATOR THAT BENElTS FROM THE
PRESENCE OF PREY SPECIES WE WOULD HAVE #_ #_ %_ %_ AND
M M )N THE EXTREME CASE WHERE SPECIES IS A SPECIALIST CONSUMER EG
A PARASITE OR PARASITOID THAT IS TOTALLY DEPENDENT ON THE PRESENCE OF SPECIES
#_ %_ ' AND M AND THE MODEL REDUCES TO A THREE PATCH
SYSTEM SIMILAR TO EPIDEMIOLOGICAL MODELS 4HE DIRECTION OF THE INEQUALITIES FOR THE
GROWTH OF SPECIES DEPEND ON THE BIOLOGY OF THE INTERACTION )N A TYPICAL PREDATOR
PREY INTERACTION THE INEQUALITIES ARE ALL REVERSED FOR SPECIES
)NVASIVE SPECIES IN METAPOPULATION SYSTEMS
)N SOME CASES HOWEVER AS NOTED BY (OLT A PREDATOR MAY INSTEAD
REDUCE THE RATE OF EXTINCTION OF ITS PREY FOR INSTANCE THE PRESENCE OF THE PREDATOR
MAY PREVENT THE PREY SPECIES FROM OVEREXPLOITING ITS OWN RESOURCE BASE !LTHOUGH
THIS IS AN INTERESTING POSSIBILITY EG GENERATING ALTERNATIVE LANDSCAPE STATES
BELOW WE FOCUS INSTEAD ON MORE CLASSICAL PREDATOR PREY SYSTEMS WHERE THE PREDA
TOR IS HARMFUL FOR PREY PERSISTENCE IN A PATCH
$YNAMICS OF THE TWO SPECIES METAPOPULATION
7E NOW ANALYSE THE RATES AT WHICH PATCH OCCUPANCY IN THE TWO SPECIES METAPOPU
LATION CHANGES OVER TIME $ENOTE THE FRACTION OF PATCHES WITH SPECIES ONLY BY X
THE FRACTION WITH SPECIES ONLY BY X AND THE FRACTION WITH BOTH SPECIES PRESENT
BY Z 4HE FRACTION OF EMPTY PATCHES IS THUS n X X Z AND THE RATES OF
CHANGE OF PATCH OCCUPANCY ARE
DX
#_ n X X Z n %_ X n #_ X %_ Z
DT
DX
#_ n X X Z n %_ X n #_ X %_ Z
DT
DZ
X#_ X#_ n Z%_ %_
DT
(ERE THE IMMIGRATION RATES CAN BE EXPRESSED IN TERMS OF X X AND Z AS
_ XM ZM AND _ XM ZM
#RITERIA FOR INVASION
)NITIAL INVASION
7E NOW DESCRIBE THE CRITERIA FOR THE INITIAL PHASE OF AN INVASION 7E ASSUME THAT
THE NATIVE SPECIES SPECIES CURRENTLY OCCUPIES A PROPORTION X OF ALL PATCHES
WHERE THIS PROPORTION IS A STABLE EQUILIBRIUM FOR THE METAPOPULATION WITH JUST SPE
CIES PRESENT 7E THEN INTRODUCE THE INVADER SPECIES INTO A FEW PATCHES AND
ASK WHETHER THE FREQUENCY OF OCCURRENCE OF THE RARE INVADING SPECIES INCREASES
OR DECLINES 3INCE THE INVADING SPECIES IS RARE X AND Z ARE SMALL #ONSEQUENTLY
THE RATE AT WHICH SPECIES MIGRANTS ARRIVE AT EACH PATCH _ IS ALSO SMALL
4HUS ALL THAT MATTERS FOR THE INITIAL INVASION ARE THE LIMITING VALUES FOR SMALL
_ OF THE COLONIZATION AND EXTINCTION FUNCTIONS FOR SPECIES "Y ANALOGY WITH
+# (ARDING *- -C.AMARA AND 2$ (OLT
&IG /NE EXAMPLE OF HOW INITIAL INVASION RATE IS INmUENCED BY E THE EXTINCTION RISK
OF THE INVADER IN TYPE " PATCHES -ODEL STRUCTURE AS IN &IG 4HE THREE CURVES REPRESENT
THREE DIFFERENT M MIGRATION RATES FROM TYPE " PATCHES A B C 4HE lGURE IS
BASED ON EQUATIONS AND PARAMETER VALUES ARE + X E C
+ E M
EQUATIONS AND WE ASSUME THAT #_ 5 +_ #_ X 5 +_ %_ 5 E AND
%_ 5 E FOR SMALL _ $URING THE INITIAL PHASE OF INVASION BY SPECIES THE IMMI
GRATION RATE OF SPECIES MIGRANTS CAN BE TAKEN AS CONSTANT AND EQUAL TO ITS VALUE
AT THE EQUILIBRIUM OCCUPANCY X OF SPECIES 7E THUS ALSO ASSUME THAT THE RATE
AT WHICH A 4YPE PATCH IS CONVERTED TO A 4YPE " PATCH C > #MX IS CONSTANT
3IMILARLY THE RATE AT WHICH A 4YPE " PATCH CONVERTS TO 4YPE E > %MX IS CON
STANT 7E NOW SUBSTITUTE THESE EXPRESSIONS INTO EQUATIONS AND EXPRESS
_ IN TERMS OF X AND Z EQ AND SINCE X AND Z ARE SMALL IGNORE TERMS INVOLV
ING X XZ AND Z 4HE RATE OF CHANGE IN PROPORTION OF INVADED PATCHES EARLY IN
THE INVASION PROCESS ARE THEN GIVEN BY THE FOLLOWING LINEARISED EQUATIONS
DX
+ n X M n E C X + n X M E Z
DT
DZ
+XM C X +XM n E E Z
DT
4HESE EQUATIONS CAN BE WRITTEN AS
)NVASIVE SPECIES IN METAPOPULATION SYSTEMS
DX
)X *Z
DT
DZ
*X )Z
DT
WHERE
) + n X M n E C
) +XM n E E
* + n X M E
* +XM C
4HE QUANTITY ) IS THE NET RATE AT WHICH A 4YPE PATCH LEADS TO THE PRODUCTION
OF 4YPE PATCHES ) IS THE RATE OF COLONIZATION OF EMPTY PATCHES BY MIGRANTS FROM
THE 4YPE PATCH IE + n X M MINUS THE RATE OF DISAPPEARANCE OF THE FOCAL
4YPE PATCH AS A RESULT OF EXTINCTION OR COLONIZATION BY SPECIES IE E C
SEE EQ 4HE TERM * CAN BE INTERPRETED AS THE RATE OF hCROSSING OVERv THE RATE
AT WHICH A 4YPE " PATCH CONTRIBUTES TO THE FORMATION OF 4YPE PATCHES 4HERE ARE
TWO WAYS THIS CONTRIBUTION IS MADE 4HE TERM + n X M IS THE RATE AT WHICH
MIGRANTS FROM THE 4YPE " PATCH CONVERT EMPTY PATCHES INTO 4YPE PATCHES BY
COLONIZATION AND E IS THE RATE AT WHICH THE FOCAL 4YPE " PATCH ITSELF CHANGES TO A
4YPE PATCH AS A RESULT OF THE EXTINCTION OF THE SPECIES OCCUPANTS SEE EQN
3IMILARLY ) IS THE NET RATE AT WHICH A 4YPE " PATCH LEADS TO THE PRODUCTION
OF 4YPE " PATCHES 4HE QUANTITY +XM IS THE RATE AT WHICH MIGRANTS FROM THE
4YPE " PATCH COLONISE 4YPE PATCHES AND SO CONVERT THEM TO 4YPE " AND E E
IS THE RATE AT WHICH THE 4YPE " PATCH REVERTS TO A PATCH WITH JUST A SINGLE SPECIES
PRESENT EQ 4HE CROSS TERM * IS THE NET RATE AT WHICH A SINGLE 4YPE PATCH
CONTRIBUTES TO THE FORMATION OF 4YPE " PATCHES !GAIN THERE ARE TWO ELEMENTS TO
THIS CONTRIBUTION -IGRANTS FROM THE 4YPE PATCH COLONISE 4YPE PATCHES AND
THE PATCH MAY ITSELF BE COLONIZED BY SPECIES SEE EQ
,ET W X Z BE THE TOTAL PROPORTION OF PATCHES IN WHICH SPECIES IS PRESENT
4O ANALYSE WHETHER SPECIES CAN INITIALLY INVADE WE LOOK AT HOW THIS QUANTITY
CHANGES OVER TIME 4HE LINEARISED EQUATIONS AND IMPLY THAT WHATEVER
THE INITIAL VALUES OF X AND Z THE PROPORTION X W WILL SETTLE DOWN TO AN EQUILIB
RIUM VALUE !S THIS EQUILIBRIUM IS APPROACHED
DW
Ah
DT
+# (ARDING *- -C.AMARA AND 2$ (OLT
\) * \
WHERE THE RATE OF INITIAL INVASION h IS THE LARGEST EIGENVALUE OF THE MATRIX ! \ \
\* )\
4HUS SPECIES CAN INVADE THE METAPOPULATION IF h WHILE IF h THE SPE
CIES GOES EXTINCT IN THE METAPOPULATION
&IGURE ILLUSTRATES HOW INVASION RATE
can BE INmUENCED BY THE EXTINCTION RISK OF THE INVADER IN PATCHES OF 4YPE " E FOR
ONE COMBINATION OF PARAMETER VALUES )N THIS CASE INCREASING EXTINCTION RISK IN
4YPE " PATCHES IS VERY COSTLY FOR THE INVADER AND AT TOO HIGH E INVASION EVEN
BECOMES IMPOSSIBLE &IG
"ELOW WE SHALL DESCRIBE THE EARLY INVASION PROCESS FOR TWO SPECIAL CASES BEFORE
CONSIDERING THE GENERAL CRITERIA FOR INVASION
!N @INVISIBLE INVADER
3UPPOSE THAT THE INVADER SPECIES CANNOT COMPETE FOR PATCHES WITH SPECIES
3PECIlCALLY SPECIES IS UNABLE TO COLONISE PATCHES ALREADY OCCUPIED BY SPECIES
AND SPECIES IMMEDIATELY WIPES OUT SPECIES WHEN SPECIES COLONISES A 4YPE
PATCH 4HEN THERE EXIST NO PATCHES CONTAINING BOTH SPECIES SO Z SO THAT EQUA
TIONS AND REDUCE TO THE SINGLE EQUATION
DX
)X
DT
4HE RATE OF GROWTH OF THE NUMBER OF PATCHES CONTAINING SPECIES IS JUST ) SO
INVASION IS POSSIBLE IF ) )F X APPROACHES ZERO THIS EXPRESSION REDUCES TO THE
CONDITION FOR INVASION BY A SINGLE SPECIES DISCUSSED EARLIER !S X INCREASES FROM
THERE IS A VALUE AT WHICH THE INVADER IS EXCLUDED )N THE SIMPLE ,EVINS
MODEL FOR INVASION TO BE POSSIBLE REQUIRES THAT THE INVADER HAVE A HIGHER EQUILIB
RIUM PATCH OCCUPANCY WERE IT ALONE THAN DOES THE RESIDENT SPECIES SEE BELOW
! @PARASITE TYPE INVADER
.OW SUPPOSE THAT SPECIES IS TOTALLY RELIANT ON THE PRESENCE OF THE NATIVE SPECIES
&OR EXAMPLE SPECIES MIGHT BE A PARASITE OR A SPECIALIST PREDATOR OF SPECIES
EG PARASITIC WASPS ARE OFTEN OBLIGATE SPECIALISTS ON PARTICULAR HOST SPECIES
.OUHUYS AND (ANSKI OR AN OBLIGATE COMMENSAL 3PECIlCALLY ASSUME THAT
SPECIES IS UNABLE TO COLONISE EMPTY PATCHES &URTHERMORE ON PATCHES WHERE
BOTH SPECIES ARE PRESENT A 4YPE " PATCH IF SPECIES DIES OUT THEN SPECIES
IMMEDIATELY DIES OUT AS WELL SO THAT THE PATCH BECOMES EMPTY 4HEN X AND
THE RATE OF GROWTH IN THE NUMBER OF PATCHES CONTAINING SPECIES IS JUST ) SINCE
EQUATIONS AND REDUCE TO THE SINGLE EQUATION
DZ
)Z
DT
)NVASIVE SPECIES IN METAPOPULATION SYSTEMS
4HUS IN THIS SPECIAL CASE INVASION IS POSSIBLE IF ) &ROM WE SEE THAT AN
INCREASE IN THE OCCUPANCY BY THE RESIDENT SPECIES TENDS TO FACILITATE INVASION
BY SPECIES
'ENERAL CRITERIA FOR INVASION
7E NOW CONSIDER THE GENERAL CASE )T IS EASY TO SHOW THAT IF ) THEN h SO
INVASION IS GUARANTEED 4HIS IS NOT SURPRISING EQUATION SHOWS THAT EVEN
WHEN COMPETITION BY SPECIES IS AT ITS STRONGEST SPECIES CAN INVADE BY JUST
COLONISING EMPTY PATCHES ! FORTIORI IF ) SPECIES CAN INVADE WHATEVER
THE FORM OF THE INTERACTION BETWEEN THE TWO SPECIES 3IMILARLY ) IMPLIES
h IF THE INVADER WERE ABLE TO SPREAD WHEN RESTRICTED TO PATCHES OCCUPIED
BY SPECIES THEN IT IS CERTAINLY A SUCCESSFUL INVADER REGARDLESS OF ITS PERFORMANCE
BY ITSELF WHEN CONFRONTED WITH EMPTY PATCHES
4HUS EITHER ) OR ) IS SUFlCIENT TO ENSURE THAT SPECIES CAN INVADE
(OWEVER WHEN ) AND ) IT MAY STILL BE POSSIBLE FOR SPECIES TO INVADE
BECAUSE 4YPE " AND 4YPE PATCHES TEND TO ENHANCE EACH OTHERS FORMATION )N
OTHER WORDS MIGRANTS FROM 4YPE PATCHES MAY COLONISE 4YPE PATCHES AND SO
CONVERT THEM TO 4YPE " AND 4YPE MIGRANTS FROM 4YPE " PATCHES MAY COLONISE
EMPTY PATCHES AND SO CONVERT THEM TO 4YPE )T IS EASY TO SHOW THAT WHEN )
AND ) SPECIES CAN INVADE h PROVIDED THAT * * ) )
)NVASION IN THE LONG RUN
7E NOW LOOK AT A SPECIAL CASE ILLUSTRATING HOW THE METAPOPULATION SYSTEM CAN
DEVELOP IN THE LONG RUN AFTER THE INITIAL INVASION 7E ARE ESPECIALLY INTERESTED
IN CHARACTERIZING THE CONDITIONS FOR LONG
TERM COEXISTENCE OF THE INVADER AND
THE NATIVE SPECIES
#ONSIDER THE EXAMPLE WHEN TWO SPECIES CAN ONLY COLONISE EMPTY PATCHES
SO THAT THERE ARE NO PATCHES WITH BOTH SPECIES PRESENT 7HO EVER COMES lRST WINS
THE PATCH A SO CALLED @LOTTERY MODEL 4HUS THE PRESENCE OF ONE SPECIES REDUCES
THE NUMBER OF PATCHES THAT ARE AVAILABLE FOR THE OTHER SPECIES TO OCCUPY .OTE
HOWEVER THAT SO FAR AS A SPECIES IS CONCERNED IT IS NOT SIMPLY AS IF THE METAPOPULA
TION AS A WHOLE HAD FEWER PATCHES PRESENT 4HIS IS BECAUSE MIGRANTS OF THE SPECIES
STILL SETTLE ON PATCHES AT RANDOM SO THOSE MIGRANTS THAT SETTLE ON A PATCH CURRENTLY
OCCUPIED BY THE OTHER SPECIES ARE LOST 4HUS THERE IS A DILUTION OF MIGRANTS THAT
SETTLE ON EMPTY PATCHES 7HEN THE TWO SPECIES LOCALLY EXCLUDE ONE ANOTHER THE
EQUATIONS GOVERNING METAPOPULATION DYNAMICS WHICH ARE NOW JUST EQUATIONS
AND REDUCE TO
DX
#_ n X X n %_ X
DT
+# (ARDING *- -C.AMARA AND 2$ (OLT
DX
#_ n X X n %_ X
DT
&OR SIMPLICITY OF EXPOSITION WE ASSUME THAT WHEN ONLY SPECIES IS PRESENT
THERE IS A UNIQUE EQUILIBRIUM LEVEL OF PATCH OCCUPANCY WHICH WE DENOTE BY X
3IMILARLY WHEN SPECIES IS PRESENT ALONE IT HAS A UNIQUE EQUILIBRIUM LEVEL OF PATCH
OCCUPANCY X
4O ANALYSE THE ABILITY OF EACH SPECIES TO INVADE WHEN THE OTHER IS PRESENT WE
ASSUME THAT #_ 5 +_ AND %_ 5 E FOR SMALL _ AND #_ 5 +_ AND
%_ 5 E FOR SMALL _ NOTE THAT THESE FUNCTIONS MAY DEVIATE FROM THIS FORM
WHEN _ AND _ ARE NOT SMALL -OTIVATED BY ,EVINS MODEL WE DElNE
E E
X n AND
X n
+M + M
4HESE ARE THE EQUILIBRIUM LEVELS OF SINGLE SPECIES OCCUPANCY WHEN THE ABOVE
APPROXIMATIONS FOR SMALL _ AND _ HOLD EXACTLY FOR ALL _ AND _ 7HEN X IS
SMALL WE HAVE #_ n X X n %_ X 5 +MX X X 4HUS BY EQUA
TION
DX
DT
X X WHEN X IS SMALL
4HAT IS THE INITIAL RATE OF INCREASE IN PATCH OCCUPANCY BY SPECIES IS POSITIVE IF
THE PATCH OCCUPANCY IT INITIALLY SEEMS TO AIM TOWARDS IN THE ABSENCE OF SPECIES
X IS HIGHER THAN THE CURRENT PATCH OCCUPANCY BY THE OTHER SPECIES X 4HUS IF
SPECIES IS AT ITS EQUILIBRIUM OF PATCH OCCUPANCY X
SPECIES CAN INVADE IF
X X
3IMILARLY IF SPECIES IS AT ITS EQULIBRIUM OF PATCH OCCUPANCY X
THEN SPECIES
CAN INVADE IF
X X
)F BOTH EXPRESSIONS AND HOLD ONE WILL SEE ROBUST COEXISTENCE IN THAT
EACH SPECIES CAN INCREASE WHEN IT IS RARE AND THE OTHER SPECIES IS AT ITS RESPECTIVE
EQUILIBRIUM SEE FURTHER BELOW
4HE DYNAMICS OF THE LOTTERY MODEL WITH ,EVINS FUNCTIONS
)N ANALYSING THE POSSIBILITY FOR COEXISTANCE IT IS OF UTMOST IMPORTANCE TO CONSIDER
THE SHAPE OF THE EXTINCTION AND COLONIZATION FUNCTIONS 4HE LOTTERY MODEL OUTLINED
)NVASIVE SPECIES IN METAPOPULATION SYSTEMS
ABOVE HAS BEEN SHOWN TO EXCLUDE COEXISTENCE (ANSKI SEE ALSO 3HURIN ET AL
(OWEVER THIS CONCLUSION IS BASED ON ASSUMING ,EVINS STATIC FUNCTIONS
7HEN EACH OF THE SINGLE SPECIES METAPOPULATIONS ARE AS IN ,EVINS BASIC MODEL WE
HAVE #_ +_ AND %_ E FOR ALL _ AND #_ +_ AND %_ E FOR
ALL _ )N THIS SPECIAL CASE X X
AND X X
)E THE EQUILIBRIUM PATCH OCCUPAN
CY IS EQUAL TO WHAT IS EXPECTED BY THE RATE OF INITIAL INVASION 4HUS IF SPECIES IS AT
ITS EQUILIBRIUM SPECIES CAN INVADE IF AND ONLY IF X
X
3IMILARLY IF SPECIES IS
AT ITS EQUILIBRIUM OCCUPANCY THEN SPECIES CAN INVADE IF AND ONLY IF X
X
4HUS
THE SPECIES WITH THE HIGHEST EQUILIBRIUM PATCH OCCUPANCY WHEN ALONE CAN INVADE
THE OTHER AND IT IS NEVER POSSIBLE FOR BOTH SPECIES TO BE ABLE TO INVADE EACH OTHER
&IGURE SHOWS THE METAPOPULATION DYNAMICS IN THE WHOLE OF THE X
X
PLANE
!S CAN BE SEEN THE METAPOPULATION DYNAMICS LEADS TO THE INEVITABLE EXTINCTION OF
THE SPECIES WITH THE SMALLEST EQUILIBRIUM PATCH OCCUPANCY ON ITS OWN 4HUS WITH
THESE ASSUMPTIONS COEXISTENCE IS NOT POSSIBLE AND THE SPECIES WITH THE LARGEST
EQUILIBRIUM OCCUPANCY EXCLUDES THE OTHER (ANSKI P
&IG )N A LOTTERY MODEL PARAMETERISED WITH ,EVINS STATIC FUNCTIONS COEXISTENCE OF
SPECIES X
AXIS AND SPECIES Y
AXIS IS NOT POSSIBLE 4HE PROPORTION OF PATCHES OCCUPIED
BY SPECIES WILL CONTINUE TO INCREASE UNTIL SPECIES GOES EXTINCT
+# (ARDING *- -C.AMARA AND 2$ (OLT
4HE LOTTERY MODEL WITH GENERAL FUNCTIONS
4O ANALYSE METAPOPULATION DYNAMICS IN THE GENERAL CASE WHEN THE COLONIZATION
AND EXTINCTION RATE CAN HAVE MANY DIFFERENT SHAPES WE CONSIDER HOW THE SIGNS OF
DX DX
AND VARY IN THE X X PLANE 7HEN SPECIES IS ABSENT X PATCH
DT DT
OCCUPANCY OF SPECIES INCREASES WITH TIME WHEN OCCUPANCY IS BELOW THE EQUILIB
RIUM OCCUPANCY X
IE
DX
X X
WHEN X
DT
&ROM CRITERIA AND THE CURVE
DX
IN THE X X PLANE JOINS THE POINT X
TO THE POINT X
DT
DX
3IMILARLY THE CURVE JOINS THE POINT X TO THE POINT X
DT
4HESE CURVES ARE ILLUSTRATED IN &IG 4HE lGURE ALSO SHOWS THE RESULTING
METAPOPULATION DYNAMICS &ROM CRITERIA AND AND AS THE lGURE ILLUS
TRATES IF X X
AND X X
THEN EACH SPECIES CAN INVADE THE OTHER AT THE LATTERS
EQUILIBRIUM PATCH OCCUPANCY 4HE METAPOPULATION DYNAMICS THEN LEADS TO STABLE
CO
EXISTENCE OF THE TWO SPECIES 4HE CONDITIONS FOR MUTUAL INVASION CAN BE SATIS
lED WHEN THE INITIAL RATE OF INCREASE IN PATCH NUMBERS IS HIGHER THAN WOULD BE
EXPECTED FROM ITS EQUILIBRIUM 4HIS PHENOMENON IS LIKELY TO ARISE IF EXTINCTION RATE
IS INITIALLY LOWER OR COLONIZATION INITIALLY HIGHER COMPARED TO THE RATE AT HIGHER
PATCH OCCUPANCIES 4HIS TYPE OF DISPROPORTIONALLY HIGH INITIAL RATE OF INCREASE IN
PATCH OCCUPANCY WILL BE FOUND IN POPULATIONS WITH ANTI
RESCUE EFFECTS ANDOR
ANTI
!LLEE EFFECTS 7HEN SUCH EFFECTS ARE PRESENT THE INITIAL RATE OF INCREASE OF EACH
SINGLE SPECIES METAPOPULATION CAN BE HIGH WITHOUT A CORRESPONDINGLY HIGH PATCH
OCCUPANCY AT EQUILIBRIUM 4HIS REDUCED EQUILIBRIUM SIZE REDUCES COMPETITION
ALLOWING CO
EXISTENCE EVEN IN THE LOTTERY MODEL SEE &IG
4HE DYNAMICS AND STABILITY OF A TWO
SPECIES SYSTEM DEPEND ON THE EXACT SHAPE
OF THE COLONIZATION AND EXTINCTION FUNCTIONS 7E HAVE ILLUSTRATED THIS FOR THE
SIMPLE LOTTERY MODEL MORE COMPLEX FORMS OF INTERACTIONS AMONG SPECIES REQUIRE
FURTHER ANALYSIS
)NVASIVE SPECIES IN METAPOPULATION SYSTEMS
&IG ! LOTTERY MODEL WHICH ALLOWS DIFFERENT SHAPES OF THE # AND % FUNCTIONS CAN HAVE
STABLE COEXISTENCE OF TWO SPECIES 4HE INITIAL RATE OF INVASION MUST BE DISPROPORTIONALLY
HIGH WHICH CAN BE THE CASE FOR EXAMPLE WHEN THERE IS AN ANTI
RESCUE EFFECT
%80,)#)4 30!4)!, 3425#452% !.$ &).)4% .5-"%2 /& 0!4#(%3
4HE FRAMEWORK DESCRIBED ABOVE ASSUMES THE NUMBER OF PATCHES TO BE VERY LARGE
(OWEVER IF THE NUMBER OF PATCHES IS LESS THAN ABOUT (ANSKI P
SINGLE STOCHASTIC EVENTS OF COLONIZATION AND EXTINCTION BECOME IMPORTANT 4HE RISK
THAT ALL PATCHES WILL GO EXTINCT SIMULTANEOUSLY METAPOPULATION EXTINCTION DEPENDS
ON THE COMBINATION OF METAPOPULATION SIZE AND THE SHAPE OF THE PER PATCH EXTINCTION
FUNCTION &IG ! CONSIDERATION OF THE EFFECTS OF lNITE PATCH NUMBERS IS INEVITABLE
WHEN CONSIDERING THE CONSEQUENCES OF EXPLICIT SPACE AND LANDSCAPE STRUCTURE
! VERY ACTIVE AREA OF RESEARCH AT PRESENT IS FOCUSED ON MAKING METAPOPULATION
THEORY SPATIALLY EXPLICIT 4HE BASIC IDEA IS THAT IN A METAPOPULATION WITH A lNITE
NUMBER OF PATCHES EACH PATCH HAS ITS OWN SPECIlC CHARACTERISTIC AND LANDSCAPE
POSITION WHICH IN TURN INmUENCE ITS PROBABILITY OF EXTINCTION AND LIKELIHOOD OF
COLONIZATION -ETAPOPULATION DYNAMICS OF THE ENTIRE ENSEMBLE REmECTS BOTH HETE
ROGENEITY AMONG PATCHES EG IN AREA WHICH CAN INmUENCE EXTINCTION AND THE
SPECIlC LANDSCAPE STRUCTURE AND PATTERN OF CONNECTIVITY OF THE PATCHES 4HE SIZE
+# (ARDING *- -C.AMARA AND 2$ (OLT
AND COMPOSITION OF LOCAL POPULATIONS ALSO INmUENCE INVASION SPEED .EUBERT AND
#ASWELL /VASKAINEN AND (ANSKI PROVIDE AN EXCELLENT OVERVIEW OF
RECENT THEORY IN THIS AREA AND HERE WE SIMPLY SKETCH SOME OF THE BASIC ISSUES
WITH AN EMPHASIS UPON IMPLICATIONS FOR INVASION BIOLOGY -ETAPOPULATION MODELS
WITH A SPATIALLY EXPLICIT STRUCTURE PROVIDE IMPORTANT LINKAGES WITH LANDSCAPE ECO
LOGY AND WITH APPLIED ISSUES SUCH AS POPULATION VIABILITY ANALYSES
/NE BIOLOGICAL ASSUMPTION THAT IS BUILT INTO THE GENERALIZED ,EVINS MODEL WE
HAVE CONSIDERED ABOVE IS THAT ALL EMPTY PATCHES ARE ACCESSIBLE TO MIGRANTS EMA
NATING FROM ALL OCCUPIED PATCHES )N GENERAL DISPERSAL IS SPATIALLY LIMITED AND SO
THIS ASSUMPTION WILL BE VIOLATED 4HIS MAY BE PARTICULARLY IMPORTANT TO CONSIDER
IN THE EARLY STAGES OF AN INVASION WHEN A COLONISING SPECIES HAS ESTABLISHED A
BEACHHEAD ON JUST ONE OR A FEW SITES &ROM THESE INITIAL SITES WITH LIMITED DISPER
SAL THERE MAY BE A SMALL lNITE NUMBER OF EMPTY SITES AVAILABLE FOR COLONIZATION
-ATHEMATICALLY THE DYNAMICS CAN BE DESCRIBED AS A -ARKOV CHAIN IN DISCRETE
TIME OR -ARKOV PROCESS IN CONTINUOUS TIME %XTINCTION CAN ARISE BECAUSE OF
AN ANALOGUE OF DEMOGRAPHIC STOCHASTICITY AT THE METAPOPULATION SCALE %VEN IF
INITIAL INVASION IS SUCCESSFUL THE RATE OF INCREASE AND RATE OF SPATIAL SPREAD OF
THE COLONISING SPECIES CAN BE DAMPED BECAUSE MANY COLONISTS EMERGING FROM
THE CENTER OF THE INVASION SIMPLY MOVE TO PATCHES THAT ARE ALREADY OCCUPIED
)N REACTION
DIFFUSION STYLE MODELS OF INVASIVE SPECIES HETEROGENEITY IN DISPERSAL
RATES SEEMS TO EXERT A PARTICULARLY STRONG EFFECT ON THE OVERALL RATE OF INVASION OF
SPECIES 3HIGESADA AND +AWASAKI )N SPATIALLY
EXPLICIT LANDSCAPE MODELS
THERE CAN BE CRITICAL THRESHOLDS IN HABITAT CONNECTIVITY BELOW WHICH SPECIES ARE
LIKELY TO GO EXTINCT EVEN THOUGH A SUBSTANTIAL NUMBER OF PATCHES EXIST WHICH ARE
PERFECTLY SUITABLE FOR OCCUPANCY 7ITH 7E MIGHT CALL THIS THE @4ANTALUS
EFFECT IN METAPOPULATION ECOLOGY AFTER THE 'REEK LEGEND OF 4ANTALUS WHO WAS
TORMENTED BY AN ETERNITY OF HUNGER AND THIRST WITH LUSCIOUS GRAPES AND CLEAN
WATER VISIBLE BUT JUST BARELY OUT OF REACH
)N GENERAL SPATIALLY EXPLICIT METAPOPULATION MODELS SEEM TO IMPLY THAT
LOCALISED DISPERSAL MAKES INVASION MORE DIFlCULT (OWEVER WERE ONE TO PLACE THE
GENERALIZED ,EVINS MODEL DISCUSSED ABOVE INTO A SPATIALLY EXPLICIT LANDSCAPE A
RICHER ARRAY OF OUTCOMES MIGHT BE OBSERVED &OR INSTANCE WITH !LLEE EFFECTS GIVEN
UNIFORM DISPERSAL OVER A LARGE NUMBER OF PATCHES FROM AN INITIAL PROPAGULE TOO
FEW INDIVIDUALS MIGHT ENTER ANY GIVEN PATCH TO ENSURE SUCCESSFUL COLONIZATION
)F INSTEAD DISPERSAL IS LOCALISED MOST INDIVIDUALS MIGHT ENTER THE SAME RELA
TIVELY FEW PATCHES THUS INCREASING THE EFFECTIVE MIGRANT DENSITY IN THOSE PATCHES
4HIS COULD FACILITATE COLONIZATION 7HAT ONE MIGHT OBSERVE IN THIS CASE IS A
SUCCESSFUL NUCLEUS OF INITIAL COLONIZATION FOLLOWED BY ACCRETIONARY GROWTH AS
NEIGHBORING PATCHES GET SUCCESSFULLY COLONIZED !LTERNATIVELY IF THERE ARE GAPS
IN AVAILABILITY OF SUITABLE HABITATS !LLEE EFFECTS COULD LEAD TO CONSTRAINTS ON
THE ABILITY OF AN INVASIVE SPECIES TO EXPAND MUCH BEYOND THE SITES OF ITS ORIGINAL
COLONIZATION +EITT ET AL
/NE COMPLICATION THAT ARISES IN SPATIALLY
EXPLICIT METAPOPULATIONS IS THAT
IF THERE IS HETEROGENEITY AMONG PATCHES EG IN ATTRIBUTES THAT INmUENCE
)NVASIVE SPECIES IN METAPOPULATION SYSTEMS
COLONIZATION SUCH HETEROGENEITY IS LIKELY TO EXHIBIT SPATIAL AUTOCORRELATION 4HIS
CAN LEAD TO POPULATION SYNCHRONY WHICH CAN STRONGLY INmUENCE THE PROBABILITY OF
REGIONAL EXTINCTION IE A FAILED INVASION #ASWELL AND #OHEN %NGEN ET AL
)N GENERAL FOR POPULATION PERSISTENCE THE SIZE OF THE METAPOPULATION
SHOULD BE SUBSTANTIALLY GREATER THAN THE SCALE AT WHICH SUCH POPULATION SYN
CHRONY IS OBSERVED 7ORKING OUT THE IMPLICATIONS OF SPATIAL AND TEMPORAL AUTO
CORRELATION FOR METAPOPULATION DYNAMICS IS AN IMPORTANT AREA FOR FUTURE WORK
%-0)2)#!, 345$)%3 !.$ 4(% 2/,% /& 4(%/29
)N ATTEMPTS TO @TEST PREDICTIONS FROM THEORETICAL MODELS EMPIRICAL METAPOPULA
TION BIOLOGISTS ALMOST WITHOUT EXCEPTION WILL REACH THE CONCLUSION THAT THEIR
STUDY SYSTEM DOES NOT lT THE ASSUMPTIONS OF THE MODEL /R THEY MIGHT EXPRESS
IT THE OTHER WAY AROUND THE MODEL DOES NOT lT THEIR OBSERVATIONS "UT THERE
ARE MANY USES OF THEORETICAL MODELS AND NOT ALL MODELS ARE MEANT TO BE TESTED
/NE OF THEIR MAIN FUNCTIONS IS AS TOOLS FOR THOUGHT 3UCH MODELS CAN BE USED TO
EXPOSE THE LOGIC OF PROCESSES ASSUMED TO BE OPERATING IN PARTICULAR SYSTEMS SHOW
ING CLEARLY HOW PREDICTIONS DEPEND ON ASSUMPTIONS 4HEORETICAL MODELS CAN BE
USED AS A STARTING POINT TO GENERATE QUESTIONS ABOUT EMPIRICAL SYSTEMS AND AS
NEW BIOLOGICAL KNOWLEDGE ABOUT THE DRIVING FORCES OF THAT SYSTEM ARE DISCOVERED
NEW SIMPLISTIC GENERAL MODELS CAN BE DEVELOPED OR NEW MORE DETAILED MODELS
CAN BE ELABORATED /CCASIONALLY A MODEL MIGHT PREDICT THE BEHAVIOUR OF A SYSTEM
"OX ALTHOUGH STOCHASTICITY DUE TO WEATHER AND THE CHAOTIC DYNAMICS OF
NATURE WILL MOSTLY HIDE SUCH CORRESPONDENCE (OWEVER MODELS ARE USEFUL IF COR
RECTLY HANDLED INCORPORATING MECHANISMS THAT WE THINK ARE IMPORTANT CAN REVEAL
THE POTENTIAL BEHAVIOUR OF A SYSTEM *UST AS IN LEARNING ABOUT THE SIMPLE BUILDING
BRICKS OF HYDROLOGY AND GAS KINETICS METEREOLOGISTS ALSO LEARN MORE ABOUT THE
NATURE OF CURRENTS AND WINDS AND EVEN CLIMATE CHANGE AND YET ARE UNABLE TO PRE
DICT THE WEATHER AT A CERTAIN SPOT MORE THAN A FEW DAYS AHEAD SO THE MISMATCH
BETWEEN A THEORETICAL MODEL AND EMPIRICAL SYSTEMS IN ECOLOGY RARELY PROVIDES
SENSIBLE @TESTS OF THE THEORY ITSELF
-ETAPOPULATION THEORY IN THE GENERALIZED FORM PRESENTED HERE IS A PHILOSOPHI
CAL TOOL THAT HIGHLIGHTS HOW SPECIES CAN PERSIST DESPITE THE FACT EVERY SINGLE SUB
POPULATION FACES A SUBSTANTIAL EXTINCTION RISK )T DOES NOT ATTEMPT TO DESCRIBE THE
DETAILED BEHAVIOUR OF A GIVEN REAL SYSTEM AT A GIVEN TIME BUT CAPTURES ESSENTIALS
OF FORCES THAT APPLY TO REAL SYSTEMS
&OR EXAMPLE 'UTIRREZ ET AL STUDIED THE SPATIAL DISTRIBUTION OF FOUR
LEPIDOPTERAN BUTTERmY SPECIES "ECAUSE THE FOUR SPECIES ALL HAD THE SAME FAVOURITE
HOST PLANT ,OTUS CORNICULATUS THE AUTHORS CLAIMED THIS MUST BE A SYSTEM WHERE
THE PREDICTIONS FROM MULTI
SPECIES MODELS WOULD BE REALISED (OWEVER IT TURNED
OUT THAT ENVIRONMENTAL FACTORS OTHER THAN THE FAVOURITE HOST PLANT DETERMINED THE
DISTRIBUTION OF THE FOUR SPECIES FOR EXAMPLE ALTERNATIVE HOST PLANTS DIFFERENCES
BETWEEN COASTAL OR INLAND SITES AND VULNERABILITY TO GRAZING /NE OF THE SPECIES
+# (ARDING *- -C.AMARA AND 2$ (OLT
WAS PRESENT IN THE ENTIRE LANDSCAPE AND MIGHT NOT ACT AS A METAPOPULATION AT
ALL 4HE STUDY CLAIMS TO HAVE PROVEN THAT THE ASSUMPTIONS OF PHENOMENOLOGICAL
MULTI
SPECIES METAPOPULATION MODELS ARE UNREALISTIC "UT THESE MODELS CAN ONLY
PROVIDE INTERESTING INSIGHTS INTO SYSTEMS WHERE SEVERAL SPECIES ACTUALLY DO COM
PETE FOR THE SAME PATCHES AND WHERE COLONIZATION
EXTINCTION DYNAMICS AND INTER
SPECIlC INTERACTIONS ARE IN FACT MAJOR FORCES AT WORK 4HESE ASSUMPTIONS NEED TO
BE ASSESSED AND IT IS NOT CLEAR THAT THE EMPIRICAL SYSTEM OF 'UTIERREZ ET AL
PROVIDES A GOOD MATCH TO THE METAPOPULATION ASSUMPTION 7E SUGGEST THAT ON
A PRIORI GROUNDS IT IS REASONABLE TO BELIEVE THAT THERE ARE MULTI
SPECIES ASSEM
BLAGES WHICH QUITE FAITHFULLY MATCH THE ASSUMPTIONS OF METAPOPULATION THEORY
EG AQUATIC PREDATORS AND PREY IN PONDS OCCUPY PATCHES WHICH MAY DRY AND
SUFFER EXTINCTIONS PARASITES IN THE SAME HOST CAN INTERACT AND ALL GO EXTINCT WHEN
THE HOST DIES ETC AND WHERE MULTI
SPECIES METAPOPULATION MODELS ARE LIKELY TO
PRODUCE SOME INTERESTING INSIGHTS
4HE OLD 0OPPERIAN VIEW THAT HYPOTHESIS MUST BE CONlRMED OR FALSIlED IN CRITICAL
TESTS HAS LED TO MUCH CONFUSION AT THE INTERFACE BETWEEN THEORETICAL AND EMPIRICAL
METAPOPULATION BIOLOGY 4HEORETICAL MODELS CAN NOT SAY MORE THAN @GIVEN EXACTLY
THESE CONDITIONS WE WILL GET THIS OR THAT OUTCOME THEY CAN NOT BE VALIDATED OR
FALSIlED IN SYSTEMS WITH OTHER CONDITIONS AND OTHER PREVAILING MECHANISMS
)T WOULD BE LIKE TELLING THE METEREOLOGISTS THAT THEY SHOULD BE SKEPTICAL ABOUT
THE PROPOSITION @WARM AIR IS LESS DENSE THAN COLD AIR SIMPLY BECAUSE THEY CANNOT
ACCURATELY PREDICT TODAYS RAINFALL IN A SPECIlC LOCATION SUCH AS 5LLAPOOL
"OX
"IOLOGICAL CONTROL EXPERIMENT IN A SNAIL METAPOPULATION LIVING IN PONDS
IN 'UADELOUPE
"IOMPHALARIA GLABRATA IS A NATIVE SNAIL SPECIES THAT IS THE INTERMEDIATE HOST
FOR HUMAN INTESTINAL SCHISTOSOMES A TREMATODE INFECTION ! 6ENEZUELAN
SNAIL -ARISA CORNUARIETIS WAS INTRODUCED IN 4HE INVADER DESTROYED
LOCAL POPULATIONS OF WATER LILIES THAT ARE IMPORTANT HABITATS FOR THE NATIVE
" GLABRATA ! STRONG DECLINE IN PATCH OCCUPANCY IN THE " GLABRATA META
POPULATION FOLLOWED IN THE PONDS WHERE THE COMPETITOR WAS INTRODUCED
)NTERESTINGLY NEARBY CONTROL PONDS WITHOUT THE INTRODUCED COMPETITOR
ALSO HAD DECLINING PATCH OCCUPANCY 0OINTIER ET AL !LTHOUGH THE
AUTHORS STATE THAT ENVIRONMENTAL CIRCUMSTANCES MIGHT HAVE CONTRIBUTED
TO THE DECLINE OF " GLABRATA IN THE UN
INVADED CONTROL PONDS THEY POINT
OUT THAT THE OBSERVED PATTERN MATCHES THAT PREDICTED BY SIMPLE THEORETI
CAL METAPOPULATIONS MODELS LOWER PATCH OCCUPANCY LEADS TO LOWERED
OVERALL COLONIZATION RATE AND THE WHOLE METAPOPULATION NETWORK CAN BE
AFFECTED BY THE REMOVAL OF A FEW PATCHES
)NVASIVE SPECIES IN METAPOPULATION SYSTEMS
#/.#,53)/.
7E HAVE SUMMARISED SOME ASPECTS OF HOW GENERALIZED METAPOPULATION THEORY
IS RELEVANT FOR UNDERSTANDING INVASIONS INTO PATCHY HABITATS 7HEN SPECIES SUR
VIVE IN A LANDSCAPE AS A METAPOPULATION IT IS CRITICAL THAT COLONIZATION SUCCESS
IS HIGHER THAN THE EXTINCTION RATE OF SUBPOPULATIONS )N CONSERVATION THE WORRY
IS OFTEN THAT AN INVADER COMPETES WITH A NATIVE SPECIES 7E HAVE RElNED A TWO
SPECIES METAPOPULATION MODEL TO CAPTURE DIFFERENT KINDS OF INTERACTIONS INCLUD
ING COMPETITION 4HERE IS A FULL RANGE OF BEHAVIOURS THAT EMERGE DEPENDING ON
THE TYPE OF COLONIZATION AND EXTINCTION FUNCTION OF THE INVADER AND THE TYPE OF
INmUENCE OF THE NATIVE SPECIES 4HE EXISTENCE OF !LLEE AND ANTI
!LLEE EFFECTS OPENS
UP A RICH VARIETY OF POSSIBLE DYNAMICAL OUTCOMES 4HE INVASIVE SPECIES BECOMES
EXTINCT IF ITS INITIAL SUCCESS IN COLONZING EMPTY PATCHES OR RESISTING EXTINCTION IN
OCCUPIED PATCHES AT LOW PATCH NUMBER IS TOO LOW WE HAVE SUGGESTED A NUMBER OF
PARTICULAR MECHANISMS THAT CAN LEAD TO SUCH EXCLUSION )F THE INITIAL GROWTH RATE
IS POSITIVE IT CAN BECOME ESTABLISHED AND COEXIST SUSTAINABLY WITH THE NATIVE SPE
CIES OR FORCE THE LATTER TO GO EXTINCT )F THE INVASIVE SPECIES COMPETES FOR RESOURCES
THIS CAN REDUCE PATCH OCCUPANCY FOR THE NATIVE SPECIES AND IF THE NATIVE SPECIES
IS VULNERABLE AND HAS AN UNSTABLE EQUILIBRIUM BEYOND WHICH THERE IS NO RETURN
THEN EXTINCTION OF THE NATIVE SPECIES CAN BE SUDDEN AND DIFlCULT TO REVERSE 7E
PREDICT METAPOPULATION THEORY WILL BECOME SIGNIlCANT MORE RElNED AND APPLIED
TO STUDY INVASIVE SPECIES PROBLEMS
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%VOLUTION
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3EPT (AKAPAINO /Y (ELSINKI
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,EIBOLD !LTERNATIVE STABLE STATES AND REGIONAL COMMUNITY STRUCTURE *OURNAL OF
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3KELLAM * ' 2ANDOM DISPERSAL IN THEORETICAL POPULATIONS "IOMETRIKA
4ILMAN $ 0LANT STRATEGIES AND THE DYNAMICS AND STRUCTURE OF PLANT COMMUNITIES
0RINCETON 5NIVERSITY 0RESS 0RINCETON .*
7ITH + ! 5SING PERCOLATION THEORY TO ASSESS LANDSCAPE CONNECTIVITY AND EFFECTS
OF HABITAT FAGMENTATIN 0AGES
IN + * 'UTZWILLER EDITOR !PPLYING ,ANDSCAPE
%COLOGY IN "IOLOGICAL #ONSERVATION 3PRINGER
6ERLAG .EW 9ORK
#HAPTER EIGHTEEN
#OMPETITION AND THE ASSEMBLY
OF INTRODUCED BIRD
COMMUNITIES
2 0 $UNCAN AND $ - &ORSYTH
).42/$5#4)/.
! CENTRAL GOAL OF ECOLOGY IS TO UNDERSTAND WHAT DETERMINES THE NUMBER AND IDEN
TITY OF SPECIES IN ECOLOGICAL COMMUNITIES /F THE MANY SPECIES THAT COULD POTEN
TIALLY OCCUPY AN AREA WHY DO ONLY A PARTICULAR SUBSET OF SPECIES ACTUALLY CO
OCCUR
AND WHAT DETERMINES THE IDENTITY OF THOSE SPECIES !ND TO WHAT EXTENT ARE THESE
PATTERNS THE PRODUCT OF DETERMINISTIC PROCESSES RATHER THAN STOCHASTIC EVENTS
-UCH INTEREST HAS FOCUSED ON INTERSPECIlC COMPETITION AS A PROCESS SHAPING CO
OCCURRENCE PATTERNS 3TRONG ET AL $IAMOND AND #ASE SOME SPECIES
CAPABLE OF JOINING A LOCAL COMMUNITY MAY BE EXCLUDED BY THE PRESENCE OF COM
PETITORS )F COMPETITION IS SUFlCIENTLY STRONG AND PERVASIVE ENOUGH TO STRUCTURE
ECOLOGICAL COMMUNITIES THEN CERTAIN @ASSEMBLY RULES SHOULD GOVERN HOW COM
MUNITIES ARE PUT TOGETHER $IAMOND 7EIHER AND +EDDY )N PARTICU
LAR COMPETITION SHOULD BE MORE INTENSE AND COMPETITIVE EXCLUSION MORE LIKELY
AMONG SPECIES OF SIMILAR SIZE AND MORPHOLOGY THAT COMPETE FOR SIMILAR RESOURCES
)N THESE CIRCUMSTANCES WE EXPECT CO
OCCURRING SPECIES TO BE MORPHOLOGICALLY DIF
FERENT FROM EACH OTHER AND TO EXHIBIT A PATTERN OF @MORPHOLOGICAL OVERDISPERSION
0IMM
-7 #ADOTTE ET AL EDS #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY n
3PRINGER 0RINTED IN THE .ETHERLANDS
20 $UNCAN AND $- &ORSYTH
-ANY STUDIES HAVE ATTEMPTED TO EVALUATE THE IMPORTANCE OF INTERSPECIlC COM
PETITION IN SHAPING CO
OCCURRENCE PATTERNS BY TESTING FOR MORPHOLOGICAL OVER
DISPERSION EG 3TRONG ET AL 'RANT AND !BBOT "OWERS AND "ROWN
$IAMOND AND #ASE $AYAN AND 3IMBERLOFF +INGSTON ET AL
3UCH TESTS HAVE GENERALLY INVOLVED COMPARING THE MORPHOLOGICAL PATTERN
ING OF SPECIES IN A LOCAL COMMUNITY WITH THAT OF A LARGER POOL OF SPECIES POTENTIALLY
CAPABLE OF JOINING THAT COMMUNITY )F SPECIES IN THE LOCAL COMMUNITY EXHIBITED
STRONGER MORPHOLOGICAL OVERDISPERSION THAN EXPECTED UNDER A PROCESS OF RANDOM
ASSEMBLY FROM THIS LARGER POOL THEN IT WAS CONCLUDED THAT THE LOCAL COMMUNITY
WAS STRUCTURED BY COMPETITION #RITICAL TO THE OUTCOME OF SUCH TESTS IS DECIDING
WHICH SPECIES TO INCLUDE IN THE LARGER POOL OF SPECIES THAT COULD POTENTIALLY JOIN THE
LOCAL COMMUNITY #OLWELL AND 7INKLER 'OTELLI AND 'RAVES )DEALLY
ALL SPECIES KNOWN TO HAVE BEEN PRESENT AT SOME STAGE DURING THE PROCESS OF ASSEM
BLING THAT COMMUNITY WOULD BE INCLUDED -ISTAKENLY INCLUDING SPECIES THAT WERE
NEVER PART OF THE PROCESS EG BECAUSE THEY NEVER REACHED THE SITE CAN BIAS THE
OUTCOME OF TESTS FOR OVERDISPERSION #OLWELL AND 7INKLER 5NFORTUNATELY
IN MOST SITUATIONS WE DO NOT KNOW HOW LOCAL COMMUNITIES WERE ASSEMBLED AND
WHICH SPECIES WERE OR WERE NOT PRESENT DURING THE PROCESS #ONSEQUENTLY
EVIDENCE FOR COMPETITIVE EXCLUSION BASED ON TESTS FOR MORPHOLOGICAL OVERDISPER
SION IS OFTEN SUBJECT TO UNCERTAINTIES ASSOCIATED WITH CHOOSING THE APPROPRIATE
SPECIES POOL
3INCE THE TH CENTURY THERE HAS BEEN A SUBSTANTIAL INCREASE IN THE MOVEMENTS
OF PEOPLE AROUND THE GLOBE AND AN ASSOCIATED INCREASE IN THE NUMBERS OF SPECIES
TRANSPORTED TO AND RELEASED IN NEW ENVIRONMENTS 7ILLIAMSON -ACK
ET AL $UNCAN ET AL !T SOME LOCATIONS T HERE ARE SUFlCIENTLY DETAILED
RECORDS THAT WE CAN RECONSTRUCT THE HISTORY OF THESE INTRODUCTIONS )N PARTICULAR
THERE ARE EXCELLENT RECORDS DOCUMENTING THE BIRD SPECIES INTRODUCED TO LOCATIONS
SUCH AS THE .EW :EALAND AND (AWAIIAN ISLANDS INCLUDING DATA ON THE SPECIES
INTRODUCED DATES OF INTRODUCTION WHETHER THEY ESTABLISHED WILD POPULATIONS OR
NOT AND MEASURES OF EFFORT SUCH AS THE NUMBERS OF INDIVIDUALS OR THE NUMBERS
OF RELEASE EVENTS OF EACH SPECIES INTRODUCED -OULTON AND 0IMM 6ELTMAN
ET AL 4HESE RECORDS PROVIDE A REMARKABLE OPPORTUNITY TO TEST HYPOTHESES
ABOUT THE FACTORS STRUCTURING ECOLOGICAL COMMUNITIES 0IMM ,OCKWOOD
ET AL BECAUSE THEY PROVIDE A NEAR COMPLETE LIST OF THE SPECIES PRESENT DUR
ING THE ASSEMBLY PROCESS THUS REDUCING THE UNCERTAINTY ASSOCIATED WITH CHOOSING
THE APPROPRIATE SPECIES POOL &URTHERMORE BECAUSE BIRD INTRODUCTIONS TO MANY
PLACES INCLUDING (AWAII AND .EW :EALAND OCCURRED DECADES TO CENTURIES AGO
THERE SHOULD HAVE BEEN SUFlCIENT TIME FOR THE COMMUNITIES TO REACH COMPOSITIONAL
EQUILIBRIUM SPECIES THAT ARE GOING TO BE EXCLUDED BY COMPETITION SHOULD HAVE
ALREADY BEEN SO
"ECAUSE OF THESE ADVANTAGES RECORDS OF BIRD INTRODUCTIONS HAVE BEEN USED
EXTENSIVELY TO TEST FOR PATTERNS CONSISTENT WITH THE OUTCOME OF COMPETITION )N
ADDITION TO TESTS FOR MORPHOLOGICAL OVERDISPERSION STUDIES HAVE TESTED FOR @PRIORITY
EFFECTS -OULTON BIRDS INTRODUCED AT A LATER DATE IN THE ASSEMBLY PROCESS
!SSEMBLY OF INTRODUCED BIRD COMMUNITIES
SHOULD BE MORE LIKELY TO FAIL BECAUSE THEY HAVE TO COMPETE WITH A GREATER DIVERSITY
OF ALREADY ESTABLISHED SPECIES AT HIGHER ABUNDANCE 5SING THESE APPROACHES SEV
ERAL STUDIES HAVE IDENTIlED PATTERNS CONSISTENT WITH COMPETITION AND ARGUED THAT
IT IS AN IMPORTANT PROCESS STRUCTURING INTRODUCED BIRD ASSEMBLAGES -OULTON AND
0IMM -OULTON -OULTON AND 0IMM A B 0IMM
-OULTON AND ,OCKWOOD ,OCKWOOD ET AL -OULTON ,OCKWOOD
AND -OULTON "ROOKE ET AL -OULTON AND 3ANDERSON ,OCKWOOD
ET AL -OULTON ET AL
.EVERTHELESS THE RESULTS OF SOME STUDIES HAVE BEEN EQUIVOCAL 3IMBERLOFF
-OULTON ET AL $UNCAN AND SEVERAL OBJECTIONS HAVE BEEN
RAISED REGARDING THE ANALYSIS AND INTERPRETATION OF THESE DATA 3IMBERLOFF AND
"OECKLEN $UNCAN AND "LACKBURN )N PARTICULAR PATTERNS CONSISTENT
WITH COMPETITION CAN BE CAUSED BY OR CONFOUNDED WITH VARIATION IN OTHER FAC
TORS 3PECIlCALLY $UNCAN SHOWED THAT WHILE PASSERINE BIRDS INTRODUCED
TO .EW :EALAND AT A LATER DATE WERE MORE LIKELY TO FAIL A PRIORITY EFFECT CONSISTENT
WITH THE OUTCOME OF COMPETITION THIS RESULT WAS CONFOUNDED WITH VARIATION IN
INTRODUCTION EFFORT "IRDS INTRODUCED AT A LATER DATE WERE ALSO RELEASED IN SMALLER
NUMBERS AND SO WERE MORE LIKELY TO FAIL FOR THAT REASON 3IMILARLY -OULTON ET AL
FOUND SIGNIlCANT MORPHOLOGICAL OVERDISPERSION AMONG GAMEBIRDS SUC
CESSFULLY INTRODUCED TO .EW :EALAND BUT $UNCAN AND "LACKBURN SHOWED
THAT THIS PATTERN COULD NOT HAVE BEEN DUE TO COMPETITION BECAUSE THE DISTRIBUTION
OF MOST SPECIES DID NOT OVERLAP IN SPACE OR TIME
/UR AIM IN THIS CHAPTER IS TO TRY AND RESOLVE THESE DIFlCULTIES AND TO GAIN GREATER
INSIGHT INTO HOW HISTORICAL EVENTS INCLUDING THE TIMING AND EFFORT PUT INTO INTRO
DUCTIONS CAN INTERACT WITH A DETERMINISTIC PROCESS COMPETITION TO AFFECT THE
OUTCOME OF INTRODUCTIONS AND HENCE THE COMPOSITION AND STRUCTURE OF INTRODUCED
SPECIES ASSEMBLAGES 7E INVESTIGATE USING A SIMPLE MATHEMATICAL MODEL HOW
THREE FACTORS INTRODUCTION EFFORT THE ABUNDANCE OF A COMPETITOR AND THE STRENGTH
OF COMPETITION INTERACT TO DETERMINE THE PROBABILITY THAT AN INTRODUCED SPECIES
WILL ESTABLISH IN THE PRESENCE OF A COMPETITOR 7E THEN EXAMINE HOW WELL THE PRE
DICTIONS OF THE MODEL ARE SUPPORTED BY DATA USING THE HISTORICAL RECORD OF PASSERINE
INTRODUCTIONS TO .EW :EALAND
! -!4(%-!4)#!, -/$%, /& %34!",)3(-%.4
3INCE INTRODUCTIONS TYPICALLY INVOLVE THE RELEASE OF FEW INDIVIDUALS STOCHASTIC
PROCESSES CAN PLAY A CRITICAL ROLE IN DETERMINING WHETHER THEY SUCCEED OR FAIL
2ICHTER
$YN AND 'OEL ,ANDE 'REVSTAD 7E THEREFORE USED A
SIMPLE TWO
SPECIES STOCHASTIC BIRTH
DEATH MODEL 3"$ 2ENSHAW TO EXPLORE
HOW INTERACTIONS BETWEEN THE NUMBER OF INDIVIDUALS RELEASED THE ABUNDANCE OF
A PREVIOUSLY ESTABLISHED COMPETITOR AND PER
CAPITA STRENGTH OF INTERSPECIlC COM
PETITION AFFECT THE PROBABILITY OF A SPECIES ESTABLISHING
20 $UNCAN AND $- &ORSYTH
,ET . BE THE NUMBER OF INDIVIDUALS OF SPECIES RELEASED AT A LOCATION AND LET .
BE THE ABUNDANCE OF SPECIES A POTENTIAL COMPETITOR ALREADY PRESENT AT THAT LOCA
TION 7E ASSUME THAT BOTH SPECIES HAVE IDENTICAL BIRTH AND DEATH RATES AND THAT
INTERSPECIlC COMPETITION IS SYMMETRIC WE ARE PRIMARILY INTERESTED IN HOW DIFFER
ENCES IN THE INITIAL POPULATION SIZES OF THE TWO SPECIES AND THE PER
CAPITA STRENGTH
OF COMPETITION AFFECTS THE PROBABILITY THAT THE INTRODUCED SPECIES ESTABLISHES
!SSUMING THAT ISOLATED POPULATIONS GROW SUBJECT TO LOGISTIC GROWTH AND THAT
DENSITY
DEPENDENCE BOTH WITHIN AND BETWEEN
SPECIES AFFECTS ONLY THE DEATH RATE
CHANGES IN THE SIZE OF POPULATIONS OF THE TWO SPECIES CAN BE DESCRIBED BY A PAIR OF
DIFFERENTIAL EQUATIONS
D.
". n $. .B n .D C. E.
DT
AND
D.
". n $. .B n .D C. E.
DT
WHERE ".I .I B IS THE BIRTH RATE OF SPECIES I WITH B THE INSTANTANEOUS BIRTH RATE
AND $.I .ID C.I E.J IS THE DEATH RATE WITH D THE INSTANTANEOUS DEATH
RATE C IS A COEFlCIENT REmECTING THE PER CAPITA STRENGTH OF INTRASPECIlC COMPETITION
AND E IS A COEFlCIENT REmECTING THE PER CAPITA STRENGTH OF INTERSPECIlC COMPETITION
)F THE TWO POPULATIONS ARE AT SIZE . AND . AT A GIVEN TIME THEN THE NEXT EVENT
WILL BE ONE OF THE FOLLOWING 2ENSHAW
! BIRTH OF SPECIES TRANSITION . A . WITH PROBABILITY ". 2
.
! DEATH OF SPECIES TRANSITION . A . n WITH PROBABILITY $. 2
.
! BIRTH OF SPECIES TRANSITION . A . WITH PROBABILITY ". 2
.
! DEATH OF SPECIES TRANSITION . A . n WITH PROBABILITY $. 2
.
WHERE 2. ". $. ". $.
7E IMPLEMENTED A 3"$ MODEL BY GENERATING TWO INDEPENDENT RANDOM NUM
BERS 9 9 ON THE UNIFORM UNIT INTERVAL ; = )F
9 ) ". 2. THEN THE NEXT EVENT WAS A BIRTH OF SPECIES
9 ". 2. AND ) ;". $. = 2. THEN THE NEXT EVENT WAS A
DEATH OF SPECIES
9 ;". $. = 2. AND ) ;". $. ". = 2. THEN THE
NEXT EVENT WAS A BIRTH OF SPECIES
9 ;". $. ". = 2 . THEN THE NEXT EVENT WAS A DEATH OF SPE
CIES
!SSEMBLY OF INTRODUCED BIRD COMMUNITIES
4HE TIME BETWEEN EVENTS IS AN EXPONENTIALLY DISTRIBUTED RANDOM VARIABLE SO
THAT THE INTER
EVENT TIME T WAS SIMULATED AS
n ;LOGE9 =
T
2.
AND THE TIME ADVANCED BY THAT AMOUNT 2ENSHAW 4HIS PROCESS WAS REITER
ATED FOR STEPS 7E DElNED SPECIES AS HAVING ESTABLISHED IF IT PERSISTED TO THE
END OF THE STEP SIMULATION FOR THE PARAMETER VALUES WE USED POPULATIONS
THAT PERSISTED FOR STEPS WERE TYPICALLY mUCTUATING AROUND CARRYING CAPACITY
AND WERE LIKELY TO PERSIST FOR MUCH LONGER
4HERE IS STRONG THEORETICAL ,ANDE 'REVSTAD AND EMPIRICAL EVI
DENCE )NCHAUSTI AND (ALLEY $RAKE AND ,ODGE THAT ENVIRONMENTAL
STOCHASTICITY AFFECTS THE PERSISTENCE OF SPECIES OVER AND ABOVE THE EFFECTS OF DEMO
GRAPHIC STOCHASTICITY ALONE (ENCE IN ADDITION TO THE DEMOGRAPHIC STOCHASTICITY
MODELLED ABOVE WE ALSO MODELLED ENVIRONMENTAL STOCHASTICITY BY ALLOWING THE
DEATH RATE OF EACH SPECIES TO VARY INDEPENDENTLY BETWEEN STEPS ACCORDING TO THE
FOLLOWING EQUATION
DI D ABS ;. U =
WHERE DI IS THE INSTANTANEOUS DEATH RATE OF SPECIES I AND . U IS A RANDOM NOR
MAL DEVIATE WITH MEAN AND VARIANCE U 4HIS ACTS TO MODEL DEVIATIONS AWAY FROM
SOME MINIMUM INSTANTANEOUS DEATH RATE D WITH LARGER VALUES OF U IMPLYING
GREATER VARIATION IN DEATH RATE BETWEEN STEPS AND THUS GREATER ENVIRONMENTAL
STOCHASTICITY
4O INVESTIGATE THE GENERAL BEHAVIOUR OF THE MODEL WE HELD PARAMETERS CON
STANT AT THE FOLLOWING VALUES IN OUR SIMULATIONS ENVIRONMENTAL STOCHASTICITY
U BIRTH RATE B DEATH RATE D AND INTRAPECIlC COMPETITION COEFlCIENT
C 7E SET INITIAL VALUES OF . AS AND AND . AS
AND AND THE INTERSPECIlC COMPETITION COEFlCIENT E AS
AND &OR EACH COMBINATION OF PARAMETER VALUES FOR . .
AND E WE RAN THE 3"$ MODEL TIMES AND CALCULATED THE PROPORTION OF THOSE
RUNS THAT RESULTED IN SPECIES ESTABLISHING SUCCESSFULLY 4HIS PROPORTION IS
AN ESTIMATE OF THE PROBABILITY OF ESTABLISHMENT UNDER THE SET OF CONDITIONS DElNED
BY THE PARAMETERS
4HE RESULTS OF OUR SIMULATIONS ARE SUMMARISED IN &IG &IRST THE PROBABILITY OF
ESTABLISHMENT INCREASES AS INTRODUCTION EFFORT . THE INITIAL ABUNDANCE OF SPECIES
INCREASES 3ECOND WHEN INTERSPECIlC COMPETITION OCCURS IE E THE PROB
ABILITY OF ESTABLISHMENT DECLINES AS BOTH THE PER
CAPITA COMPETITION COEFlCIENT
E AND THE INITIAL ABUNDANCE OF THE COMPETITOR SPECIES . INCREASES -OREOVER
THESE THREE FACTORS INTERACT SUCH THAT COMPETITION HAS LITTLE EFFECT ON THE PROBABIL
ITY OF ESTABLISHMENT WHEN A COMPETITOR SPECIES OCCURS AT LOW INITIAL ABUNDANCE
20 $UNCAN AND $- &ORSYTH
REGARDLESS OF THE STRENGTH OF COMPETITION (ERE STOCHASTIC EVENTS DOMINATE AND
THE NUMBER OF INDIVIDUALS RELEASED IS CRITICAL 7HEN COMPETITOR SPECIES OCCUR AT
HIGHER INITIAL ABUNDANCE THE PROBABILITY OF ESTABLISHMENT DEPENDS INCREASINGLY
ON THE STRENGTH OF COMPETITION
&IG 2ESULTS OF THE STOCHASTIC BIRTH
DEATH MODEL SHOWING THE PROBABILITY OF
ESTABLISHMENT AS A FUNCTION OF THE INITIAL ABUNDANCE OF AN INTRODUCED SPECIES . GIVEN
A RESIDENT COMPETITOR AT FOUR DIFFERENT ABUNDANCES . THE FOUR PANELS AND WITH THE PER
CAPITA STRENGTH OF INTERSPECIlC COMPETITION TAKING SIX VALUES FROM TO SHOWN AS
DIFFERENT LINES
4HESE SIMULATION RESULTS HIGHLIGHT TWO PREDICTIONS &IRST THE ABUNDANCE OF A
COMPETITOR IN ADDITION TO THE
PER
CAPITA STRENGTH OF COMPETITION
SHOULD
BE IMPORTANT
!SSEMBLY OF INTRODUCED BIRD COMMUNITIES
IN DETERMINING THE OUTCOME OF AN INTRODUCTION (ENCE WE WOULD EXPECT TO
OBSERVE BOTH MORPHOLOGICAL OVERDISPERSION RESULTING FROM THE COMPETITIVE EXCLU
SION OF MORPHOLOGICALLY SIMILAR SPECIES AND PRIORITY EFFECTS A HIGHER CHANCE OF
FAILURE AMONG LATER INTRODUCTIONS BECAUSE ALREADY RESIDENT COMPETITORS HAVE HAD
TIME TO INCREASE IN ABUNDANCE IN INTRODUCED ASSEMBLAGES STRUCTURED BY COMPETI
TION 3ECOND THESE FACTORS SHOULD INTERACT SUCH THAT THE INmUENCE OF PER
CAPITA
STRENGTH OF COMPETITION DEPENDS ON THE RELATIVE ABUNDANCES OF THE INTRODUCED
AND COMPETITOR SPECIES AND THE IMPORTANCE OF COMPETITION DEPENDS ON INTRODUC
TION EFFORT &IG 4HUS INTER
SPECIlC COMPETITION IS PREDICTED TO BE OF MUCH
LESS IMPORTANCE IN AFFECTING ESTABLISHMENT WHEN A SPECIES IS INTRODUCED IN SMALL
NUMBERS STOCHASTIC PROCESSES DOMINATE OR WHEN THE RESIDENT COMPETITOR OCCURS
AT LOW ABUNDANCE 4HE EFFECTS OF COMPETITION SHOULD BE MOST APPARENT WHEN BOTH
THE INTRODUCED AND COMPETITOR SPECIES OCCUR AT HIGH ABUNDANCE
#/.&2/.4).' 4(% -/$%, 7)4( $!4! 4(% !33%-",9 /& ).42/$5#%$
0!33%2).% #/--5.)4)%3 ). .%7 :%!,!.$
$ATA
7E USED HISTORICAL DATA DOCUMENTING THE TIMING AND NUMBERS OF PASSERINE
SPECIES INTRODUCED TO FOUR ACCLIMATISATION DISTRICTS IN .EW :EALAND !UCKLAND
7ELLINGTON #ANTERBURY AND /TAGO TO TEST THESE PREDICTIONS 4HE DATA ARE FROM
$UNCAN AND LIST THE PASSERINE SPECIES INTRODUCED TO EACH DISTRICT THE YEAR
OF INTRODUCTION WHETHER SPECIES ESTABLISHED A WILD POPULATION OR NOT THE YEAR
SPECIES WENT EXTINCT IF THEY FAILED TO ESTABLISH AND INTRODUCTION EFFORT MEASURED
AS THE NUMBER OF SEPARATE RELEASE EVENTS &OR EACH SPECIES INTRODUCED TO EACH DIS
TRICT A TOTAL OF INTRODUCTIONS WE USED THESE DATA AND THE MORPHOLOGICAL DATA
DESCRIBED BELOW TO ESTIMATE THE STRENGTH OF INTERSPECIlC COMPETITION THE RELATIVE
ABUNDANCE OF THE CLOSEST COMPETITOR AND INTRODUCTION EFFORT
7E OBTAINED MORPHOLOGICAL DATA BY MEASURING MUSEUM SPECIMENS )N ALL
CASES WE MEASURED SPECIMENS COLLECTED IN THE COUNTRY WHERE THE SPECIES WAS
MOST LIKELY TO HAVE BEEN INTRODUCED FROM AND WHERE POSSIBLE WE MEASURED SPECI
MENS COLLECTED AROUND THE TIME THAT INTRODUCTIONS TO .EW :EALAND OCCURRED THE
LATE S 4HE COUNTRY OF ORIGIN FOR MOST %UROPEAN SPECIES WAS 5NITED +INGDOM
WHILE FOR .ORTH !MERICAN SPECIES WE MEASURED SPECIMENS COLLECTED ON THE WEST
COAST OF THE 5NITED 3TATES A LIKELY SOURCE AREA FOR INTRODUCTIONS TO .EW :EALAND
&OR ,ONCHURA PUNCTULATA WHICH IS NATIVE TO !SIA WE MEASURED SPECIMENS COL
LECTED IN !USTRALIA BECAUSE THIS SPECIES WAS INTRODUCED TO .EW :EALAND FROM THERE
4HOMSON !LL OF THE SPECIMENS WE MEASURED WERE HOUSED IN THE "RITISH
-USEUM OF .ATURAL (ISTORY AT 4RING 5+
7E MEASURED INDIVIDUALS OF EACH SPECIES MALE AND FEMALE EXCEPT
FOR FOUR SPECIES WHERE WE MEASURED n INDIVIDUALS WITH APPROXIMATELY EQUAL
NUMBERS OF MALES AND FEMALES 7E MEASURED SIX MORPHOLOGICAL CHARACTERS ON
20 $UNCAN AND $- &ORSYTH
EACH INDIVIDUAL THE LENGTH OF THE CULMEN FROM THE BASE OF THE SKULL TO THE TIP OF
THE UPPER MANDIBLE THE WIDTH AND DEPTH OF THE BEAK AT THE ANTERIOR MARGIN OF THE
NARES THE LENGTH OF THE TARSUS THE LENGTH OF THE WING FROM THE WRIST TO THE TIP OF
THE LONGEST PRIMARY AND THE LENGTH OF THE TAIL
&OLLOWING 2ICKLEFS AND 4RAVIS AND -OULTON AND 0IMM WE SUM
MARISED MORPHOLOGICAL DIFFERENCES AMONG SPECIES USING THE lRST TWO AXES DERIVED
FROM A PRINCIPAL COMPONENTS ANALYSIS 0#! BASED ON LOG
TRANSFORMED VALUES
OF ALL SIX MORPHOLOGICAL CHARACTERS WITH INTRODUCTIONS TO EACH DISTRICT ANALYSED
SEPARATELY &IG &OR EACH DISTRICT WE USED THE %UCLIDEAN DISTANCE BETWEEN
SPECIES IN THIS MORPHOLOGICAL SPACE TO ESTIMATE HOW MORPHOLOGICALLY SIMILAR
SPECIES ARE TO EACH OTHER AND USED THIS AS A MEASURE OF THE STRENGTH OF INTERSPE
CIlC COMPETITION
&IG 0LOTS OF THE lRST TWO AXES FROM A PRINCIPAL COMPONENTS ANALYSIS SUMMARISING
MORPHOLOGICAL DIFFERENCES AMONG THE PASSERINE SPECIES INTRODUCED TO EACH OF FOUR
ACCLIMATISATION DISTRICTS IN .EW :EALAND SEE TEXT 3PECIES THAT ESTABLISHED ARE SHOWN AS
OPEN CIRCLES SPECIES THAT FAILED ARE SHOWN AS CLOSED CIRCLES
!SSEMBLY OF INTRODUCED BIRD COMMUNITIES
0REVIOUS STUDIES HAVE USED MINIMUM SPANNING TREES COUPLED WITH RANDOMISA
TION TESTS TO SEARCH FOR OVERDISPERSION AMONG SUCCESSFULLY ESTABLISHED SPECIES IN
MORPHOLOGICAL SPACE EG -OULTON AND 0IMM -OULTON AND ,OCKWOOD
,OCKWOOD ET AL ,OCKWOOD AND -OULTON ,OCKWOOD ET AL
(ERE WE ADOPT A REGRESSION APPROACH BECAUSE WE WANT TO SIMULTANE
OUSLY CONSIDER THE INmUENCE OF SEVERAL VARIABLES ON ESTABLISHMENT PROBABILITY
&OR INTRODUCTIONS TO EACH DISTRICT WE USED THE LOG
TRANSFORMED DISTANCE BETWEEN
A SPECIES AND ITS CLOSEST NEIGHBOUR IN MORPHOLOGICAL SPACE AS AN ESTIMATE OF
THE STRENGTH OF INTERSPECIlC COMPETITION SPECIES WITH A NEAREST NEIGHBOUR MORE
SIMILAR IN MORPHOLOGY ARE ASSUMED TO HAVE FACED STRONGER COMPETITION FOR THE
SAME RESOURCES ! SPECIES COULD HAVE AS ITS NEAREST NEIGHBOUR ONLY SPECIES THAT
WERE PRESENT IN THE SAME DISTRICT AT THE SAME TIME DETERMINED FROM THE YEAR
OF INTRODUCTION TO A DISTRICT AND THE YEAR OF EXTINCTION FOR SPECIES THAT FAILED
4HIS AVOIDS A PROBLEM OFTEN PRESENT IN THE USE OF MINIMUM SPANNING TREES
WHEREBY ALL SPECIES ARE INCLUDED AS POTENTIAL COMPETITORS EVEN IF THE DISTRIBUTIONS
OF SOME SPECIES FAIL TO OVERLAP IN EITHER SPACE OR TIME $UNCAN AND "LACKBURN
4HE LEVEL OF COMPETITION THAT INDIVIDUALS OF A SPECIES EXPERIENCE WILL BE A FUNC
TION OF THEIR SIMILARITY TO A COMPETITOR AND THE ABUNDANCE OF THAT COMPETITOR .O
SPECIES ABUNDANCE ESTIMATES ARE AVAILABLE FOR THE PERIOD DURING WHICH MOST INTRO
DUCTIONS OCCURRED IN .EW :EALAND THE LATE S (OWEVER 4HOMSON
OBSERVED THAT INTRODUCED PASSERINES INCREASED RAPIDLY IN ABUNDANCE FOLLOWING
ESTABLISHMENT !LTHOUGH LEGISLATION WAS ORIGINALLY PASSED PROTECTING INTRODUCED
BIRDS 4HOMSON STATED THAT hWITHIN TEN OR lFTEEN YEARS OF THE COMING IN OF
THE BIRDS THEIR NUMBERS INCREASED TO SUCH AN EXTENT THAT THE PROTECTION AFFORDED THEM
HAD TO BE TAKEN AWAY AND RESTRICTIVE LEGISLATION IMPOSEDv $URING THIS PERIOD OF
RAPID POPULATION INCREASE THE NUMBER OF YEARS ELAPSED SINCE INTRODUCTION SHOULD
PROVIDE A REASONABLE INDEX OF RELATIVE ABUNDANCE AT LEAST FOR THOSE SPECIES THAT
SUCCESSFULLY ESTABLISHED 7E THEREFORE USED THE TIME DELAY IN YEARS SEPARATING
THE INTRODUCTION OF A TARGET SPECIES AND ITS NEAREST COMPETITOR AS AN ESTIMATE OF
THAT COMPETITORS RELATIVE ABUNDANCE /UR ASSUMPTION IS THAT THE LONGER THE TIME
DELAY THE LONGER A COMPETITORS POPULATION WILL HAVE HAD TO INCREASE AND THE
HIGHER ITS ABUNDANCE WILL BE
-ODEL DEVELOPMENT AND ANALYSIS
/UR SIMULATIONS PREDICT HOW I NUMBER OF RELEASE EVENTS A MEASURE OF INTRODUC
TION EFFORT II LOG
TRANSFORMED DISTANCE TO THE NEAREST NEIGHBOUR IN MORPHOLOGI
CAL SPACE A MEASURE OF THE PER
CAPITA STRENGTH OF INTERSPECIlC COMPETITION AND
III TIME DELAY BETWEEN INTRODUCTIONS AN ESTIMATE OF THE RELATIVE ABUNDANCE OF
THE CLOSEST COMPETITOR SHOULD INTERACT TO GENERATE PATTERNS IN ESTABLISHMENT SUC
CESS &IG 4O SEE HOW WELL THE DATA lT THESE PREDICTIONS WE CONSIDERED HOW WELL
A SERIES OF CANDIDATE LOGISTIC REGRESSION MODELS COULD EXPLAIN VARIATION IN ESTAB
LISHMENT PROBABILITY FOR PASSERINE INTRODUCTIONS TO .EW :EALAND /UR APPROACH IS
TO lT A SERIES OF CANDIDATE MODELS TO THE DATA TO IDENTIFY THE BEST
lTTING MODEL AS
20 $UNCAN AND $- &ORSYTH
A BASIS FOR INFERENCE AND THEN TO COMPARE THE RESULTS FROM THAT BEST lTTING MODEL
WITH THE SIMULATION RESULTS
/UR RESPONSE VARIABLE IS BINARY WHETHER A SPECIES INTRODUCED TO A DISTRICT ESTAB
LISHED OR NOT 7E CONSIDERED CANDIDATE LOGISTIC REGRESSION MODELS THAT MIGHT
EXPLAIN VARIATION IN ESTABLISHMENT SUCCESS ALL OF WHICH INCLUDED THE NUMBER OF
RELEASE EVENTS WHICH WE KNOW IS AN IMPORTANT EXPLANATORY VARIABLE &OURTEEN
OF THE MODELS INCLUDED COMBINATIONS OF MORPHOLOGICAL DISTANCE INTRODUCTION
DELAY AND POSSIBLE INTERACTIONS BETWEEN THESE VARIABLES THAT CAPTURE THE RANGE
OF POSSIBLE OUTCOMES MODELS IN 4ABLE 4HE REMAINING MODELS
INCLUDED COMBINATIONS OF MORPHOLOGICAL DISTANCE AND YEAR OF lRST INTRODUCTION
7E CONSIDERED MODELS CONTAINING THE VARIABLE YEAR OF lRST INTRODUCTION BECAUSE
SPECIES INTRODUCED A LONG TIME AFTER THEIR CLOSEST COMPETITOR IE A LONG INTRODUC
TION DELAY WILL ALSO HAVE A LATER DATE OF INTRODUCTION IN ABSOLUTE TERMS )T HAS BEEN
NOTED THAT IN ADDITION TO FACING GREATER COMPETITION LATER INTRODUCTIONS WOULD
HAVE FACED ADDITIONAL THREATS INCLUDING A GREATER DIVERSITY AND ABUNDANCE OF
INTRODUCED PREDATORS $UNCAN ET AL IN PRESS 7E INCLUDE YEAR OF lRST INTRODUCTION
AS A SURROGATE FOR OTHER FACTORS THAT COULD HAVE VARIED THROUGH TIME AND AFFECTED
ESTABLISHMENT PROBABILITY 7E DID NOT CONSIDER INTERACTIONS BETWEEN YEAR OF lRST
INTRODUCTION AND OTHER VARIABLES BECAUSE WE HAD NO A PRIORI REASON TO THINK THEY
SHOULD BE IMPORTANT
7E USED THE SMALL SAMPLE VERSION OF !KAIKES )NFORMATION #RITERION !)#C
TO IDENTIFY WHICH OF THE CANDIDATE MODELS BEST lTTED THE DATA "URNHAM AND
!NDERSON 4HE BEST lTTING MODEL HAS THE SMALLEST !)#C VALUE
AND WE RANKED MODELS FROM BEST TO WORST BASED ON THE DIFFERENCE BETWEEN EACH
MODELS !)#C AND THE !)#C OF THE BEST lTTING MODEL 6I VALUES 7E THEN CALCULATED
THE !KAIKE WEIGHTS WI SEE "URNHAM AND !NDERSON WHICH PROVIDE A
WEIGHT OF EVIDENCE FOR EACH MODEL GIVEN AS AN APPROXIMATE PROBABILITY THAT THE
BEST lTTING MODEL IS IN FACT THE BEST OUT OF THE CANDIDATE SET )F THERE IS CLEARLY A
SINGLE BEST MODEL AS JUDGED BY THE WEIGHT OF EVIDENCE THEN THAT MODEL CAN BE
USED AS A BASIS FOR INFERENCE )N OUR CASE HOWEVER NO SINGLE MODEL PROVIDED A BEST
lT TO THE DATA WITH SEVERAL CANDIDATE MODELS PROVIDING A SIMILARLY GOOD lT 2ATHER
THAN ARBITRARILY SELECTING ONE OF SEVERAL WELL lTTING MODELS WE USED INFORMATION
CONTAINED IN ALL OF THESE MODELS AS A BASIS FOR ROBUST MULTI MODEL INFERENCE
4O DO THIS WE IDENTIlED ALL CANDIDATE MODELS FOR WHICH THE SUM OF THE !KAIKE
WEIGHTS FROM LARGEST TO SMALLEST WAS &ROM THIS SET OF BEST lTTING MODELS
WE CALCULATED MODEL AVERAGED PARAMETER ESTIMATES WEIGHTING THE PARAMETERS IN
EACH MODEL BY THE !KAIKE WEIGHT FOR THAT MODEL AND THEN SUMMED THE WEIGHTED
ESTIMATES ACROSS ALL MODELS "URNHAM AND !NDERSON 7E THEN BASED OUR
INFERENCE ON THESE MODEL AVERAGED VALUES
2ESULTS
#ANDIDATE MODELS THAT INCLUDED THE VARIABLE @YEAR OF lRST INTRODUCTION INSTEAD OF
THE VARIABLE @INTRODUCTION DELAY RECEIVED VIRTUALLY NO SUPPORT MODELS
IN
!SSEMBLY OF INTRODUCED BIRD COMMUNITIES
4ABLE WEIGHT OF EVIDENCE WI ALL ) IMPLYING THAT VARIATION IN ESTABLISH
MENT SUCCESS IS BETTER EXPLAINED BY THE TIME DELAY BETWEEN THE INTRODUCTION OF
A TARGET SPECIES AND ITS CLOSEST COMPETITOR RATHER THAN THE ABSOLUTE DATE OF INTRO
DUCTION
4ABLE 4HE CANDIDATE MODELS WE CONSIDERED TO EXPLAIN VARIATION IN THE PROBABILITY
THAT A PASSERINE SPECIES INTRODUCED TO AN ACCLIMATISATION DISTRICT IN .EW :EALAND ESTABLISHED
OR NOT 4HE EXPLANATORY VARIABLES ARE NREL THE NUMBER OF RELEASE EVENTS IN THE DISTRICT
A MEASURE OF INTRODUCTION EFFORT DIST THE LOGARITHM OF THE DISTANCE IN MORPHOLOGICAL
SPACE FROM THE TARGET SPECIES TO ITS NEAREST NEIGHBOUR A MEASURE OF THE PER
CAPITA STRENGTH
OF INTERSPECIlC COMPETITION DELAY THE TIME DELAY IN YEARS BETWEEN INTRODUCTION OF
THE TARGET SPECIES AND INTRODUCTION OF ITS MORPHOLOGICALLY NEAREST NEIGHBOUR A MEASURE OF
THE RELATIVE ABUNDANCE OF THE CLOSEST COMPETITOR YI THE YEAR OF lRST INTRODUCTION
-ODEL %XPLANATORY VARIABLES
NREL
NREL DIST
NREL DELAY
NREL DIST DELAY
NREL DIST NREL = DIST
NREL DELAY NREL = DELAY
NREL DIST DELAY NREL = DIST
NREL DIST DELAY DIST = DELAY
NREL DIST DELAY NREL = DELAY
NREL DIST DELAY NREL = DIST DIST = DELAY
NREL DIST DELAY NREL = DIST NREL = DELAY
NREL DIST DELAY NREL = DELAY DIST = DELAY
NREL DIST DELAY NREL = DIST NREL = DELAY DIST = DELAY
NREL DIST DELAY NREL = DIST NREL = DELAY DIST = DELAY NREL = DIST = DELAY
NREL YI
NREL DIST YI
NREL DIST YI NREL = DIST
/NLY ONE OF THE MODELS WITH WI DID NOT INCLUDE THE THREE KEY EXPLANATORY
VARIABLES IMPLYING THAT ALL THREE VARIABLES WERE IMPORTANT IN EXPLAINING PATTERNS
OF ESTABLISHMENT (OWEVER NO SINGLE MODEL CLEARLY PROVIDED A BEST
lT TO THE DATA
WITH THE TOP lVE MODELS HAVING 6I VALUES IMPLYING THAT EACH HAD hSUBSTANTIALv
SUPPORT "URNHAM AND !NDERSON 0ROBABILITY OF ESTABLISHMENT IS PLOTTED
AS A FUNCTION OF THE THREE EXPLANATORY VARIABLES BASED ON MODEL
AVERAGED
20 $UNCAN AND $- &ORSYTH
PARAMETER ESTIMATES IN &IG 4HE PROBABILITY OF ESTABLISHMENT WAS POSITIVELY
RELATED TO INTRODUCTION EFFORT NEGATIVELY WITH INCREASING STRENGTH OF COMPETITION
AND NEGATIVELY WITH THE TIME DELAY BETWEEN INTRODUCTIONS
4ABLE #OMPARISON OF THE CANDIDATE MODELS EXPLAINING VARIATION IN THE PROBABILITY
THAT A PASSERINE SPECIES INTRODUCED TO AN ACCLIMATISATION DISTRICT IN .EW :EALAND ESTABLISHED
OR NOT -ODEL NUMBER CORRESPONDS TO THOSE IN 4ABLE + IS THE NUMBER OF PARAMETERS IN
THE MODEL 6I IS THE DIFFERENCE BETWEEN THE !KAIKE )NFORMATION #RITERION !)#C FOR EACH
MODEL AND THAT OF THE BEST
lTTING MODEL MODEL AND WI ARE THE !KAIKE WEIGHTS
-ODEL LOG
LIKELIHOOD + !)#C 6I WI - WI
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$ISCUSSION
4HE RELATIONSHIPS BETWEEN ESTABLISHMENT PROBABILITY AND THE THREE EXPLANATORY
VARIABLES REVEALED BY THE DATA FOR PASSERINE INTRODUCTIONS TO .EW :EALAND &IG
AGREE REMARKABLY WELL WITH THE FORM OF THE RELATIONSHIPS PREDICTED BY OUR
SIMPLE STOCHASTIC BIRTH DEATH MODEL INCORPORATING INTERSPECIlC COMPETITION &IG
)N BOTH THE DATA AND THE MODEL THE PROBABILITY OF ESTABLISHMENT INCREASED
WITH GREATER INTRODUCTION EFFORT -OREOVER FOR A GIVEN LEVEL OF INTRODUCTION EFFORT
ESTABLISHMENT PROBABILITY DECLINED WHEN INTRODUCED SPECIES FACED A COMPETITOR
!SSEMBLY OF INTRODUCED BIRD COMMUNITIES
&IG 4HE RELATIONSHIP BETWEEN PROBABILITY OF ESTABLISHMENT AND NUMBER OF RELEASE
EVENTS FOR FOUR VALUES OF INTRODUCTION DELAY THE NUMBER OF YEARS AFTER INTRODUCTION OF ITS
CLOSEST COMPETITOR THAT A TARGET SPECIES IS INTRODUCED AND FOR FOUR VALUES OF MORPHOLOGICAL
DISTANCE THE DISTANCE IN MORPHOLOGICAL SPACE ;&IG = FROM A SPECIES
TO ITS NEAREST NEIGHBOUR SHOWN AS DIFFERENT LINES FOR PASSERINE BIRDS INTRODUCED TO FOUR
ACCLIMATISATION DISTRICTS IN .EW :EALAND
THAT WAS MORE MORPHOLOGICALLY SIMILAR TO THEM AND DECLINED FURTHER WHEN THAT
COMPETITOR HAD BEEN INTRODUCED EARLIER THAN THE TARGET SPECIES ALLOWING TIME FOR
THE COMPETITOR TO HAVE ATTAINED HIGHER ABUNDANCE 4HIS AGREEMENT BETWEEN THE
RELATIONSHIPS PREDICTED BY THE MODEL AND THOSE OBSERVED IN THE DATA SUGGESTS THAT
OUR SIMPLE STOCHASTIC BIRTH
DEATH MODEL CAPTURES IMPORTANT FEATURES UNDERPIN
NING ESTABLISHMENT SUCCESS IN THIS SYSTEM )N PARTICULAR OUR RESULTS SUGGEST THAT
INTERSPECIlC COMPETITION AFFECTED ESTABLISHMENT PROBABILITY FOR PASSERINE SPECIES
INTRODUCED TO .EW :EALAND AND THAT THIS ASSEMBLAGE HAS THEREFORE BEEN STRUC
TURED BY COMPETITION
20 $UNCAN AND $- &ORSYTH
0ATTERNS CONSISTENT WITH THE OUTCOME OF COMPETITION HAVE BEEN OBSERVED IN
BIRD INTRODUCTION DATA WHEN COMPETITION COULD NOT HAVE BEEN A CAUSE OF THOSE
PATTERNS -OULTON ET AL $UNCAN AND "LACKBURN AND PATTERNS CON
SISTENT WITH COMPETITION HAVE BEEN SHOWN TO BE CONFOUNDED WITH OTHER FACTORS
3IMBERLOFF AND "OECKLEN $UNCAN 7E EMPHASISE THAT WE HAVE
ATTEMPTED TO CONTROL FOR THESE EFFECTS lRST BY CONSIDERING ONLY SITUATIONS IN WHICH
SPECIES OVERLAPPED IN SPACE AND TIME AND COULD THEREFORE HAVE COMPETED SECOND
BY INCORPORATING THE CONFOUNDING EFFECTS OF INTRODUCTION EFFORT INTO OUR MODEL AND
ANALYSIS AND THIRD BY SHOWING THAT INCLUDING INTRODUCTION DELAY A COMPETITION
EFFECT PROVIDES A BETTER lT TO THE DATA THAN INCLUDING ABSOLUTE DATE OF INTRODUCTION
A SURROGATE FOR OTHER VARIABLES THAT MIGHT HAVE CHANGED THROUGH TIME
7HILE HISTORICAL RECORDS OF BIRD INTRODUCTIONS PROVIDE A UNIQUE OPPORTUNITY
TO INVESTIGATE THE ROLE OF COMPETITION AND OTHER FACTORS IN STRUCTURING SPECIES
ASSEMBLAGES 0IMM ,OCKWOOD ET AL WE STRESS THE NEED TO ACCOUNT
FOR SUCH POTENTIALLY CONFOUNDING FACTORS -OREOVER OUR RESULTS HIGHLIGHT THAT
THE IMPORTANCE OF COMPETITION RELATIVE TO OTHER FACTORS IN DETERMINING ESTABLISH
MENT SUCCESS CAN VARY DEPENDING ON CIRCUMSTANCES #OMPETITION SHOULD BE OF LESS
IMPORTANCE IN STRUCTURING A GROUP OF SPECIES ALL INTRODUCED AT THE SAME TIME AND
IN LOW NUMBERS BECAUSE STOCHASTIC EVENTS WILL LARGELY DETERMINE WHICH OF THESE
SPECIES ESTABLISH AND WHICH FAIL &IG
4HESE RESULTS HAVE IMPORTANT IMPLICATIONS FOR STUDIES THAT ASSESS THE IMPORTANCE
OF COMPETITION IN DETERMINING INVASION SUCCESS REGARDLESS OF THE HABITAT OR TAXA
CONSIDERED 4HIS IS BECAUSE THE SUPPLY OF INVADING PROPAGULES AND THE ABUNDANCES
OF RESIDENT COMPETITORS INVARIABLY DIFFER WIDELY BETWEEN LOCATIONS AND STUDIES AND
INTERACTIONS BETWEEN THESE FACTORS INmUENCE THE ROLE THAT COMPETITION PLAYS 3UCH
DIFFERENCES MAY CONTRIBUTE TO VARIATION IN THE RELATIVE IMPORTANCE OF COMPETITION
THAT IS OBSERVED AMONG EXPERIMENTAL INVASION STUDIES ,EVINE ET AL
4HE CENTRAL MESSAGE OF OUR STUDY IS THAT BOTH STOCHASTIC EVENTS IN THIS CASE
MEDIATED BY INTRODUCTION EFFORT AND DETERMINISTIC PROCESSES IN THIS CASE THE
OUTCOME OF INTERSPECIlC COMPETITION ARE IMPORTANT IN COMMUNITY ASSEMBLY AND
THAT THE TIMING AND SEQUENCE OF INTRODUCTIONS IE HISTORY MATTERS DUE TO PRIOR
ITY EFFECTS /UR RESULTS SUGGEST THAT .EW :EALAND WOULD HAVE ENDED UP WITH A
DIFFERENT ASSEMBLAGE OF INTRODUCED PASSERINES HAD THE EFFORT PUT INTO INTRODUCING
SPECIES BEEN REDISTRIBUTED OR HAD THE SEQUENCE AND TIMING OF INTRODUCTIONS BEEN
RESHUFmED 4HE lNDING THAT HISTORICAL CONTINGENCY IS IMPORTANT IN COMMUNITY
ASSEMBLY IS NOT NEW AND HAS BEEN DEMONSTRATED REPEATEDLY USING MATHEMATICAL
MODELS AND IN LABORATORY SYSTEMS EG 'ILPIN AND #ASE $RAKE ET AL
&UKAMI AND -ORIN )NSIGHTS INTO THE ASSEMBLY PROCESS PROVIDED BY HISTORI
CAL RECORDS OF BIRD INTRODUCTIONS HINT THAT SUCH EFFECTS COULD BE JUST AS PERVASIVE
IN REAL COMMUNITIES
!SSEMBLY OF INTRODUCED BIRD COMMUNITIES
!#+./7,%$'%-%.43
4HANKS TO 2OBERT 0RYS
*ONES FOR ACCESS TO THE BIRD COLLECTIONS AT 4RING AND THE
.EW :EALAND -ARSDEN &UND FOR FUNDING DATA COLLECTION 7E THANK 4IM "LACKBURN
0HILL #ASSEY *ULIE ,OCKWOOD AND AN ANONYMOUS REVIEWER FOR HELPFUL DISCUSSION
AND COMMENTS
2%&%2%.#%3
"OWERS - ! AND * ( "ROWN "ODY SIZE AND COEXISTENCE IN DESERT RODENTS CHANCE
OR COMMUNITY STRUCTURE %COLOGY
"ROOKE 2 + * , ,OCKWOOD AND - 0 -OULTON 0ATTERNS OF SUCCESS IN PASSERIFORM
BIRD INTRODUCTIONS ON 3AINT (ELENA /ECOLOGIA
"URNHAM + 0 AND $ 2 !NDERSON +ULLBACK
,IEBLER INFORMATION AS A BASIS FOR
STRONG INFERENCE IN ECOLOGICAL STUDIES 7ILDLIFE 2ESEARCH
"URNHAM + 0 AND $ 2 !NDERSON -ODEL SELECTION AND MULTIMODEL INFERENCE ND
EDITION 3PRINGER
6ERLAG .EW 9ORK
#OLWELL 2 + AND $ 7 7INKLER ! NULL MODEL FOR NULL MODELS IN BIOGEOGRAPHY
0AGES
IN $ 2 3TRONG $ 3IMBERLOFF , ' !BELE AND ! " 4HISTLE EDITORS
%COLOGICAL COMMUNITIES CONCEPTUAL ISSUES AND THE EVIDENCE 0RINCETON 5NIVERSITY
0RESS 0RINCETON .*
$AYAN 4 AND $ 3IMBERLOFF -OPHOLOGICAL RELATIONSHIPS AMONG COEXISTING HETERO
MYIDS AN INCISIVE DENTAL CHARACTER !MERICAN .ATURALIST
$IAMOND * AND 4 * #ASE #OMMUNITY ECOLOGY (ARPER AND 2OW .EW 9ORK
$IAMOND * - !SSEMBLY OF SPECIES COMMUNITIES IN - , #ODY AND * - $IAMOND
EDITORS %COLOGY AND EVOLUTION OF COMMUNITIES (ARVARD 5NIVERSITY 0RESS #AMBRIDGE
$RAKE * ! 4 % &LUM ' * 7ITTEMAN 4 6OSKUIL ! - (OYLMAN # #RESON $ ! +ENNY
' 2 (UXEL # 3 ,ARUE AND * 2 $UNCAN 4HE CONSTRUCTION AND ASSEMBLY OF AN
ECOLOGICAL LANDSCAPE *OURNAL OF !NIMAL %COLOGY
$RAKE * - AND $ - ,ODGE %FFECTS OF ENVIRONMENTAL VARIATION ON EXTINCTION AND
ESTABLISHMENT %COLOGY ,ETTERS
$UNCAN 2 0 4HE ROLE OF COMPETITION AND INTRODUCTION EFFORT IN THE SUCCESS OF PAS
SERIFORM BIRDS INTRODUCED TO .EW :EALAND !MERICAN .ATURALIST
$UNCAN 2 0 AND 4 - "LACKBURN -ORPHOLOGICAL OVERDISPERSION IN GAME BIRDS
!VES 'ALLIFORMES SUCCESSFULLY INTRODUCED TO .EW :EALAND WAS NOT CASUED BY INTERSPE
CIlC COMPETITION %VOLUTIONARY %COLOGY 2ESEARCH
$UNCAN 2 0 4 - "LACKBURN AND 0 #ASSEY IN PRESS &ACTORS AFFECTING THE RELEASE ESTAB
LISHMENT AND SPREAD OF INTRODUCED BIRDS IN .EW :EALAND IN 7 ' ,EE EDITOR "IOLOGICAL
INVASIONS IN .EW :EALAND 3PRINGER
6ERLAG "ERLIN
$UNCAN 2 0 4 - "LACKBURN AND $ 3OL 4HE ECOLOGY OF BIRD INTRODUCTIONS
!NNUAL 2EVIEW OF %COLOGY %VOLUTION AND 3YSTEMATICS
&UKAMI 4 AND 0 * -ORIN 0RODUCTIVITY
BIODIVERSITY RELATIONSHIPS DEPEND ON THE
HISTORY OF COMMUNITY ASSEMBLY .ATURE
20 $UNCAN AND $- &ORSYTH
'ILPIN - % AND 4 * #ASE -ULTIPLE DOMAINS OF ATTRACTION IN COMPETITION COMMUNI
TIES .ATURE
'OTELLI . * AND ' 2 'RAVES .ULL MODELS IN ECOLOGY 3MITHSONIAN )NSTITUTE 0RESS
7ASHINGTON
'RANT 0 2 AND ) !BBOT )NTERSPECIlC COMPETITION ISLAND BIOGEOGRAPHY AND NULL
HYPOTHESES %VOLUTION
'REVSTAD & 3 &ACTORS INmUENCING THE CHANCE OF POPULATION ESTABLISHMENT IMPLICA
TIONS FOR RELEASE STRATEGIES IN BIOCONTROL %COLOGICAL !PPLICATIONS
)NCHAUSTI 0 AND * (ALLEY /N THE RELATION BETWEEN TEMPORAL VARIABILITY AND PERSIS
TENCE TIME IN ANIMAL POPULATIONS *OURNAL OF !NIMAL %COLOGY
+INGSTON 4 ' *ONES ! :UBAID AND 4 ( +UNZ 2ESOURCE PARTITIONING IN RHINOLO
PHOID BATS REVISITED /ECOLOGIA
,ANDE 2 2ISKS OF POPULATION EXTINCTION FROM DEMOGRAPHIC AND ENVIRONMENTAL STO
CHASTICITY AND RANDOM CATASTROPHES !MERICAN .ATURALIST
,EVINE * - 0 " !DLER AND 3 ' 9ELENIK ! META ANALYSIS OF BIOTIC RESISTANCE TO
EXOTIC PLANT INVASIONS %COLOGY ,ETTERS
,OCKWOOD * , AND - 0 -OULTON %COMORPHOLOGICAL PATTERN IN "ERMUDA BIRDS THE
INmUENCE OF COMPETITION AND IMPLICATIONS FOR NATURE PRESERVES %VOLUTIONARY %COLOGY
,OCKWOOD * , - 0 -OULTON AND 3 + !NDERSON -ORPHOLOGICAL ASSORTMENT AND
THE ASSEMBLY OF COMMUNITIES OF INTRODUCED PASSERIFORMS ON OCEANIC ISLANDS 4AHITI VER
SUS /AHU !MERICAN .ATURALIST
,OCKWOOD * , - 0 -OULTON AND + , "ALENT )NTRODUCED AVIFAUNAS AS NATURAL
EXPERIMENTS IN COMMUNITY ASSEMBLY 0AGES IN % 7EIHER AND 0 ! +EDDY EDI
TORS %COLOGICAL ASSEMBLY RULES #AMBRIDGE 5NIVERSITY 0RESS #AMBRIDGE
-ACK 2 . $ 3IMBERLOFF 7 - ,ONSDALE ( %VANS - #LOUT AND & ! "AZZAZ
"IOTIC INVASIONS CAUSES EPIDEMIOLOGY GLOBAL CONSEQUENCES AND CONTROL %COLOGICAL
!PPLICATIONS
-OULTON - 0 -ORPHOLOGICAL SIMILARITY AND COEXISTENCE OF CONGENERS AN EXPERI
MENTAL TEST WITH INTRODUCED (AWAIIAN BIRDS /IKOS
-OULTON - 0 4HE ALL OR NONE PATTERN IN INTRODUCED (AWAIIAN PASSERIFORMS THE
ROLE OF COMPETITION SUSTAINED !MERICAN .ATURALIST
-OULTON - 0 AND * , ,OCKWOOD -ORPHOLOGICAL DISPERSION OF INTRODUCED
(AWAIIAN lNCHES EVIDENCE FOR COMPETITION AND A .ARCISSUS EFFECT %VOLUTIONARY %COLOGY
-OULTON - 0 AND 3 , 0IMM 4HE INTRODUCED (AWAIIAN AVIFAUNA BIOGEOGRAPHICAL
EVIDENCE FOR COMPETITION !MERICAN .ATURALIST
-OULTON - 0 AND 3 , 0IMM A 4HE EXTENT OF COMPETITION IN SHAPING AN INTRO
DUCED AVIFAUNA 0AGES IN * $IAMOND AND 4 * #ASE EDITORS #OMMUNITY ECOLOGY
(ARPER 2OW .EW 9ORK
-OULTON - 0 AND 3 , 0IMM B 3PECIES INTRODUCTIONS TO (AWAII 0AGES IN
( ! -OONEY AND * ! $RAKE EDITORS %COLOGY OF BIOLOGICAL INVASIONS OF .ORTH !MERICA
AND (AWAII 3PRINGER .EW 9ORK
!SSEMBLY OF INTRODUCED BIRD COMMUNITIES
-OULTON - 0 AND 3 , 0IMM -ORPHOLOGICAL ASSORTMENT IN INTRODUCED (AWAIIAN
PASSERINES %VOLUTIONARY %COLOGY
-OULTON - 0 AND * ' 3ANDERSON 0REDICTING THE FATES OF PASSERIFORM INTRODUCTIONS
ON OCEANIC ISLANDS #ONSERVATION "IOLOGY
-OULTON - 0 * ' 3ANDERSON AND 2 & ,ABISKY 0ATTERNS OF SUCCESS IN GAME
BIRD !VES 'ALLIFORMES INTRODUCTIONS TO THE (AWAIIAN )SLANDS AND .EW :EALAND
%VOLUTIONARY %COLOGY 2ESEARCH
-OULTON - 0 * ' 3ANDERSON AND $ 3IMBERLOFF 0ASSERIFORM INTRODUCTIONS TO
THE -ASCARENES )NDIAN /CEAN AN ASSESSMENT OF THE ROLE OF COMPETITION COLOGIE
0IMM 3 , 4HE BALANCE OF NATURE #HICAGO 5NIVERSITY 0RESS #HICAGO
2ENSHAW % -ODELING BIOLOGICAL POPULATIONS IN TIME AND SPACE #AMBRIDGE
5NIVERSITY 0RESS #AMBRIDGE
2ICHTER
$YN . AND . 3 'OEL /N THE EXTINCTION OF A COLONIZING SPECIES 4HEORETICAL
0OPULATION "IOLOGY
2ICKLEFS 2 % AND * 4RAVIS ! MORPHOLOGICAL APPROACH TO THE STUDY OF AVIAN COM
MUNITY ORGANIZATION !UK
3IMBERLOFF $ %XTINCTION SURVIVAL AND EFFECTS OF BIRDS INTRODUCED TO THE -ASCARENES
!CTA /ECOLOGICA
3IMBERLOFF $ AND 7 "OECKLEN 0ATTERNS OF EXTINCTION IN THE INTRODUCED (AWAIIAN
AVIFAUNA A REEXAMINATION OF THE ROLE OF COMPETITION !MERICAN .ATURALIST
3TRONG $ 2 $ 3IMBERLOFF , ' !BELE AND ! " 4HISTLE %COLOGICAL COMMUNITIES
CONCEPTUAL ISSUES AND THE EVIDENCE 0RINCETON 5NIVERSITY 0RESS 0RINCETON
3TRONG $ 2 , ! 3ZYSKA AND $ 3IMBERLOFF 4ESTS OF COMMUNITY
WIDE CHARACTER
DISPLACEMENT AGAINST NULL HYPOTHESES %VOLUTION
4HOMSON ' - 4HE NATURALISATION OF ANIMALS AND PLANTS IN .EW :EALAND
#AMBRIDGE 5NIVERSITY 0RESS #AMBRIDGE
6ELTMAN # * 3 .EE AND - * #RAWLEY #ORRELATES OF INTRODUCTION SUCCESS IN EXOTIC
.EW :EALAND BIRDS !MERICAN .ATURALIST
7EIHER % AND 0 ! +EDDY %COLOGICAL ASSEMBLY RULES #AMBRIDGE 5NIVERSITY 0RESS
#AMBRIDGE
7ILLIAMSON - "IOLOGICAL )NVASIONS #HAPMAN (ALL ,ONDON
#HAPTER NINETEEN
2OOM FOR ONE MORE
%VIDENCE FOR INVASIBILITY
AND SATURATION IN ECOLOGICAL
COMMUNITIES
3 ! 3MITH AND * " 3HURIN
).42/$5#4)/.
)DENTIFYING THE CONTRIBUTIONS OF LOCAL AND REGIONAL PROCESSES IN SHAPING BIO
LOGICAL COMMUNITIES IS KEY TO UNDERSTANDING GLOBAL PATTERNS OF SPECIES RICHNESS
#ORNELL 'ASTON %COLOGISTS HAVE TYPICALLY FALLEN INTO TWO TRADITIONS
TO EXPLAIN VARIATION IN DIVERSITY ONE EMPHASIZING LOCAL PROCESSES SUCH AS COM
PETITION PREDATION MUTUALISM AND THE INTERACTION BETWEEN ORGANISMS AND THEIR
ENVIRONMENT AND THE OTHER FOCUSED ON SPECIATION EXTINCTION AND DISPERSAL OVER
BROAD REGIONAL SCALES 4ERBORGH AND &AABORG #ORNELL
#ORNELL AND ,AWTON 2ICKLEFS 4HE LOCAL SCHOOL OF THOUGHT
POSITS THAT SPECIES RICHNESS IN COMMUNITIES IS DETERMINED BY PROCESSES THAT
INmUENCE DEMOGRAPHIC RATES WITHIN HABITAT PATCHES #HASE AND ,EIBOLD
4HE REGIONAL APPROACH ARGUES THAT DISPERSAL SPECIATION EXTINCTION AND THE HIS
TORY OF COMMUNITY ASSEMBLY ARE OF PRIMARY IMPORTANCE IN SHAPING BIOLOGICAL
COMMUNITIES 2ICKLEFS #ASWELL AND #OHEN 'ASTON
-7 #ADOTTE ET AL EDS #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY n
3PRINGER 0RINTED IN THE .ETHERLANDS
3! 3MITH AND *" 3HURIN
'ASTON -ORA ET AL &UKAMI 3MITH ET AL 3MITH AND
"ERMINGHAM SUBMITTED 4HE CRITICAL DISTINCTION BETWEEN THE TWO APPROACHES
HINGES ON WHETHER LOCAL SPECIES RICHNESS IS SATURATED OR WHETHER COMMUNITIES
HAVE THE INTRINSIC CAPACITY TO SUPPORT MORE SPECIES THAN THEY ACTUALLY CONTAIN
)F THE SUPPLY OF SPECIES THROUGH COLONIZATION OR SPECIATION IS MUCH GREATER THAN
THE NUMBER THAT CAN COEXIST DUE TO LOCAL ECOLOGICAL CONSTRAINTS THEN COMMUNI
TIES ARE SAID TO BE SATURATED AND UNDER STRONG LOCAL CONTROL #ORNELL AND ,AWTON
!LTERNATIVELY IF SPECIATION AND DISPERSAL PROVIDE FEWER SPECIES THAN SITES
CAN SUPPORT THEN REGIONAL CONTROL OVER COMMUNITIES IS DOMINANT AND LOCAL RICH
NESS IS UNSATURATED 4HE QUESTION OF WHETHER COMMUNITIES ARE NEAR OR FAR FROM
SATURATION IS CRITICAL TO OUR UNDERSTANDING OF PATTERNS OF THE DISTRIBUTION AND
DIVERSITY OF ORGANISMS
4HE CONCEPT OF SATURATION IS OF FUNDAMENTAL INTEREST TO BOTH COMMUNITY ECOL
OGY AND INVASIONS BIOLOGY BECAUSE IT HAS PROFOUND IMPLICATIONS FOR THE INVASIBILITY
AND POST
INVASION DYNAMICS OF LOCAL COMMUNITIES )NVASIBILITY AND SATURATION ARE
RELATED BUT DISTINCT ASPECTS OF COMMUNITIES &OR INSTANCE THERE ARE TWO POTENTIAL
OUTCOMES OF SPECIES INTRODUCTIONS IN SATURATED COMMUNITIES &IRST SUCH COMMU
NITIES MIGHT RESIST INVASION BY SPECIES INTRODUCED VIA NATURAL OR ANTHROPOGENIC
DISPERSAL )N THIS CASE BOTH THE COMPOSITION AND SPECIES RICHNESS OF LOCAL COM
MUNITIES ARE IN A STATE OF EQUILIBRIUM AND CHANGE VERY LITTLE WITH THE INTRODUCTION
OF NOVEL SPECIES 3ECOND LOCAL COMMUNITIES MIGHT BE BOTH INVASIBLE AND SATURATED
WITH SPECIES .EW SPECIES COULD BE ADDED TO LOCAL COMMUNITIES VIA NATURAL COLO
NIZATION OR HUMAN INTRODUCTION HOWEVER THESE SPECIES DISPLACE LOCAL RESIDENTS
MAINTAINING A CONSTANT NUMBER OF SPECIES "ROWN ET AL 4HEREFORE
THE NUMBER OF SPECIES BUT NOT THEIR IDENTITIES WOULD REMAIN CONSTANT OVER TIME
!LTERNATIVELY
IN UNSATURATED COMMUNITIES DISPERSAL EVENTS OFTEN RESULT IN SUCCESS
FUL
INVASION WITHOUT CAUSING THE EXTIRPATION OF RESIDENT SPECIES .ET SPECIES RICH
NESS INCREASES WITH ENHANCED DISPERSAL BECAUSE THE ADDITION OF NEW SPECIES
EXCEEDS THE LOSS OF RESIDENTS ! lNAL POSSIBILITY IS THAT VERY HIGH DISPERSAL RATES
RESULT IN hSUPER
SATURATIONv AS COMMUNITIES SUPPORT A LARGE NUMBER OF SINK
POPULATIONS THAT ARE EXCLUDED LOCALLY IN THE ABSENCE OF DISPERSAL 3HMIDA AND
%LLNER -OUQUET AND ,OREAU 0ATTERNS OF COMMUNITY INVASIBILITY AND
CHANGES IN SPECIES RICHNESS WITH INCREASED IMMIGRATION RATES AMONG COMMUNI
TIES OR THE LENGTH OF EVOLUTIONARY HISTORY CAN THEREFORE BE USED TO EVALUATE THE
DEGREES OF SATURATION AND INVASIBILITY OF COMMUNITIES
4HIS CHAPTER ASSESSES THREE LINES OF EVIDENCE FOR SATURATION AND INVASIBILITY
IN BIOLOGICAL COMMUNITIES &IRST WE EXAMINE THE OUTCOMES OF FAUNAL EXCHANGES
RESULTING FROM RECENT ANTHROPOGENIC MODIlCATION OF THE LANDSCAPE AND PAST GEO
LOGIC EVENTS TO EVALUATE THE EFFECT OF ENHANCED IMMIGRATION BETWEEN REGIONS ON
SPECIES RICHNESS 3ECOND WE EXAMINE RECENT TRENDS IN THE NUMBERS OF ANTHRO
POGENIC SPECIES INVASIONS AND EXTINCTIONS OBSERVED OVER BROAD SPATIAL SCALES
4HE GLOBAL EXPLOSION IN EXOTIC SPECIES INVASIONS SUGGESTS THAT CONTEMPORARY COM
MUNITIES ARE CERTAINLY INVASIBLE ,ODGE -OYLE AND ,IGHT 6ITOUSEK
ET AL #OHEN AND #ARLTON (OBBS AND -OONEY 2ICCIARDI
%VIDENCE FOR INVASIBILITY AND SATURATION
7HETHER BIOLOGICAL INVASIONS ARE ACCOMPANIED BY DIFFERING NUMBERS OF EXTINC
TIONS CAN BE USEFUL FOR EVALUATING WHETHER LOCAL DIVERSITY IN THESE COMMUNITIES
IS NEAR OR BELOW SATURATION 3AX AND 'AINES 7E COMPARE THE NUMBER OF
SUCCESSFUL INVASIONS VERSUS EXTIRPATIONS IN ORDER TO EVALUATE HOW SPECIES RICHNESS
CHANGES WHEN HUMANS INTRODUCE SPECIES 4HIRD WE EXAMINE EXPERIMENTAL SPE
CIES INTRODUCTIONS TO EVALUATE THE INVASIBILITY OF LOCAL COMMUNITIES AND COMPARE
DEGREES OF SATURATION AMONG DIFFERENT GROUPS OF ORGANISMS )NTENTIONAL INTRODUC
TIONS ALLOW US TO ASSESS THE SUCCESS RATE OF COLONIZATION ATTEMPTS AN IMPORTANT
ASPECT OF SATURATION AS UNCONTROLLED INTRODUCTIONS DO NOT TELL US HOW MANY INTRO
DUCED SPECIES FAIL TO COLONIZE
&INALLY WE COMPARE THE EMPIRICAL EVIDENCE FOR SATURATION AND INVASIBILITY WITH
THE PREDICTIONS OF FOUR MAJOR THEORETICAL PERSPECTIVES ON BROAD
SCALE COMMU
NITY STRUCTURE -ODELS OF PATCH
DYNAMICS ECOLOGICAL DRIFT EG NEUTRAL MODELS
SPECIES
SORTING ALONG ENVIRONMENTAL GRADIENTS AND MASS
EFFECT SYSTEMS MAKE
DIVERGENT PREDICTIONS REGARDING COMMUNITY INVASIBILITY AND SATURATION &AUNAL
EXCHANGES PATTERNS OF HUMAN
MEDIATED INVASIONS AND EXTINCTIONS AND EXPERI
MENTAL INTRODUCTIONS MAY BE USEFUL FOR DISTINGUISHING THE UTILITY OF THE DIFFERENT
MODELS FOR DESCRIBING REAL COMMUNITIES 7E SYNTHESIZE DATA AND THEORY PERTAINING
TO THE SATURATION OF LOCAL COMMUNITIES AND DISCUSS AVENUES FOR FUTURE RESEARCH
")/4)# ).4%2#(!.'%3 ).6!3)",% !.$ 5.3!452!4%$ #/--5.)4)%3
"IOTIC INTERCHANGES OCCUR WHEN BARRIERS SEPARATING DISTINCT BIOLOGICAL COMMUNI
TIES BREAK DOWN ENABLING THE RECIPROCAL EXCHANGE OF RESIDENT SPECIES 2EMOVAL OF
DISPERSAL BARRIERS INCREASES THE LIKELIHOOD OF DISPERSAL BETWEEN PREVIOUSLY ISOLATED
COMMUNITIES PERMITTING STRONG INFERENCES REGARDING THE IMPORTANCE OF DISPER
SAL PER SE IN DETERMINING PATTERNS OF THE DISTRIBUTION AND DIVERSITY OF ORGANISMS
(ERE WE USE EVIDENCE FROM CONTEMPORARY AND HISTORIC BIOTIC INTERCHANGES TO
ASSESS WHETHER COMMUNITIES ARE SATURATED WITH SPECIES BY DOCUMENTING CHANGES
IN SPECIES RICHNESS FOLLOWING REMOVAL OF DISPERSAL BARRIERS -OREOVER BECAUSE
THE REGIONAL POOL OF SPECIES CAPABLE OF PARTICIPATING IN CROSS
BARRIER EXCHANGES
IS OFTEN KNOWN IT IS POSSIBLE TO EVALUATE THE SUCCESS RATE OF COLONIZATIONS IN
BOTH DIRECTIONS
#ONTEMPORARY BIOTIC INTERCHANGES
#ONTEMPORARY HUMAN MEDIATED BIOTIC INTERCHANGES PROVIDE A POWERFUL TEST
FOR SATURATION OF BIOLOGICAL COMMUNITIES OVER MODERATE TO LARGE GEOGRAPHIC
SCALES BECAUSE THE OUTCOME CAN BE OBSERVED DIRECTLY WITHOUT RELYING ON THE FOS
SIL RECORD "IOTIC INTERCHANGES FOLLOWING THE CONSTRUCTION OF BOTH THE 0ANAMA
AND 3UEZ #ANALS ARE WELL DOCUMENTED 0OR 3MITH ET AL
AND PROVIDE INSIGHTS INTO THE QUESTION OF COMMUNITY SATURATION )N THE CASE OF
THE 0ANAMA #ANAL THE RECENT GEOLOGIC ORIGIN OF THE )STHMUS MILLION YEARS OLD
3! 3MITH AND *" 3HURIN
AND ITS YOUNG FRESHWATER lSH FAUNA "ERMINGHAM AND -ARTIN 0ERDICES
ET AL SUGGEST THAT REGIONAL PROCESSES MIGHT BE IMPORTANT IN LIMITING SPE
CIES RICHNESS -ESOAMERICAN FRESHWATER lSH ASSEMBLAGES CONTAIN AN ORDER OF
MAGNITUDE FEWER SPECIES THAN THEIR PUTATIVE SOURCE POPULATIONS IN NORTHWESTERN
3OUTH !MERICA 3MITH AND "ERMINGHAM UNPUBLISHED DATA SUGGESTING THAT SPE
CIATION AND IMMIGRATION MAY BE MORE IMPORTANT IN LIMITING LOCAL DIVERSITY IN THIS
SYSTEM #ONSEQUENTLY COMMUNITIES OF FRESHWATER lSHES IN THE 0ANAMA #ANAL
REGION MIGHT BE BOTH SUSCEPTIBLE TO INVASION AND UNSATURATED BECAUSE THE PAST
SUPPLY OF SPECIES MAY BE LOWER THAN THE NUMBER OF SPECIES THAT THE RIVERS COULD
SUPPORT !LTERNATIVELY SPECIES RICHNESS MIGHT REMAIN BELOW THE SPECIATION
EXTINC
TION EQUILIBRIUM ,OSOS AND 3CHLUTER BECAUSE THE PROCESS OF SPECIATION MAY
BE SO SLOW THAT FEW COMMUNITIES EVER APPROACH EVOLUTIONARY EQUILIBRIUM ,OSOS
AND 3CHLUTER 2ICKLEFS AND "ERMINGHAM
#OMPLETION OF THE 0ANAMA #ANAL IN CREATED A FRESHWATER CORRIDOR BETWEEN
THE 2IO #HAGRES ON THE #ARIBBEAN SLOPE AND THE 2IO 'RANDE ON THE 0ACIlC SLOPE
OF THE )STHMUS OF 0ANAMA ENABLING THE FRESHWATER lSHES OF PREVIOUSLY ISOLATED
DRAINAGE BASINS TO INTERMINGLE 4HREE lSH SPECIES COLONIZED THE 2IO #HAGRES FROM
THE 2IO 'RANDE AND lVE SPECIES COLONIZED THE 2IO 'RANDE FROM THE 2IO #HAGRES
#OLONISTS REPRESENTED AND OF THE TOTAL SOURCE POOL OF OBLIGATE FRESHWA
TER lSH SPECIES IN THE 2IO #HAGRES AND 2IO 'RANDE THAT WERE NOT ALREADY PRESENT IN
BOTH BASINS SUGGESTING THAT FRESHWATER lSH COMMUNITIES IN THE 2IO #HAGRES AND
2IO 'RANDE WERE SUSCEPTIBLE TO INVASION 4ABLE .O CASES OF LOCAL EXTIRPATION
WERE DOCUMENTED SPECIES RICHNESS FOLLOWING DISPERSAL EVENTS INCREASED BY
IN THE 2IO #HAGRES AND IN THE 2IO 'RANDE 3MITH ET AL )N ADDITION
COLONISTS BECAME COMMON IN NEWLY INVADED COMMUNITIES ACHIEVING A LEVEL OF
DISTRIBUTION AND ABUNDANCE COMPARABLE WITH OR GREATER THAN THAT IN THEIR SOURCE
COMMUNITIES 3MITH ET AL
4HE BIOTIC INTERCHANGE THAT OCCURRED FOLLOWING COMPLETION OF THE 3UEZ #ANAL
IN WHICH LINKED THE 2ED 3EA AND THE EASTERN -EDITERRANEAN 3EA IS QUALI
TATIVELY SIMILAR TO THAT WHICH OCCURRED FOLLOWING THE CONSTRUCTION THE 0ANAMA
#ANAL (UNDREDS OF 2ED 3EA MARINE SPECIES COLONIZED THE -EDITERRANEAN VIA THE
3UEZ #ANAL INCLUDING MOLLUSKS AND lSHES 0OR ,ESSEPSIAN MIGRANTS
SPECIES THAT COLONIZED THE -EDITERRANEAN FROM THE 2ED 3EA CONSTITUTE APPROXI
MATELY OF THE TOTAL SPECIES RICHNESS OF THE ,EVANT "ASIN 0OR
-ANY FEWER COLONIZATIONS OF -EDITERRANEAN SPECIES HAVE BEEN RECORDED IN THE 2ED
3EA &IFTY
THREE MARINE SPECIES HAVE BEEN REPORTED TO HAVE PARTICIPATED IN THE FAU
NAL EXCHANGE 0OR ,ARGE DIFFERENCES IN THE NUMBER OF SUCCESSFUL INVASIONS
SUGGEST POTENTIAL DIFFERENCES IN THE INVASIBILITY OF THE TWO COMMUNITIES HOWEVER
PASSIVE TRANSPORT BY UNIDIRECTIONAL CURRENTS PARTIALLY EXPLAINS THE OBSERVED ASYM
METRY OF INVASION THROUGH THE 3UEZ #ANAL
)NVASIONS BY SPECIES MIGRATING THROUGH THE 3UEZ #ANAL HAVE CONTINUED FOR
MORE THAN YEARS SINCE THE CANALS CONSTRUCTION WHEREAS NO EXTINCTIONS HAVE
BEEN ATTRIBUTED TO INVASIONS FOLLOWING FAUNAL MIXING 0OR 2ATHER
A BATHYMETRIC READJUSTMENT OF RESIDENT SPECIES HAS OCCURRED WHEREBY RESIDENT
%VIDENCE FOR INVASIBILITY AND SATURATION
4ABLE 4HE OBLIGATE FRESHWATER lSH ASSEMBLAGES OF THE 2IO 'RANDE AND
2IO #HAGRES DRAINAGE BASINS SAMPLED BY BOTH -EEK AND (ILDEBRAND AND
3MITH ET AL #OLONISTS ARE DENOTED IN BOLD
ITALIC TEXT AND @NEW REFERS TO DRAINAGES
COLONIZED FOLLOWING COMPLETION OF THE 0ANAMA #ANAL 4HE SPECIES IN 4ABLE ARE ORGANIZED
BY THEIR SALINITY TOLERANCE AND THEN BY ALPHABETICAL ORDER OF FAMILY GENUS AND SPECIES
NAMES ! NESTED !./6! ANALYSIS OF SPECIES LOG TRANSFORMED ABUNDANCES IN ADJACENT
WATERSHEDS WAS PERFORMED IN ORDER TO DETERMINE IF SPECIES WERE MORE SUCCESSFUL IN THEIR
NATIVE OR NEWLY COLONIZED DRAINAGES
DENOTES VALUES SIGNIlCANT AT P AND o DENOTES
VALUES SIGNIlCANT AT P AND THE LOCATION OF THE SYMBOL INDICATES WHETHER THE SPECIES
WAS MORE SUCCESSFUL IN ITS NATIVE OR RECENTLY INVADED DRAINAGE
0RIMARY AND SECONDARY FRESHWATER lSHES 2IO 'RANDE 2IO #HAGRES
!STYANAX AENEUS NEW
!STYANAX RUBERRIMUS
"RYCON CHAGRENSIS NEW
"RYCON PETROSUS NEW
"RYCONAMERICUS EMPERADOR
#OMPSURA GORGONAE
'EPHYROCHARAX ATRICAUDATA
(YPHESSOBRYCON PANAMENSIS <
0SEUDOCHEIRODON AFlNIS <
2OEBOIDES OCCIDENTALIS
NEW
2OEBOIDES GUATEMALENSIS NEW
#YPHOCHARAX MAGDALENAE NEWo
(OPLIAS MICROLEPIS
0IABUCINA PANAMENSIS
!NCISTRUS CHAGRENSIS NEW
#HAETOSTOMA lSCHERI <
(YPOSTOMUS PANAMENSIS
2INELORICARIA URACANTHA <
0IMELODELLA CHAGRESI
2HAMDIA QUELEN
"RACHYHYPOPOMUS OCCIDENTALIS
2IVULUS BRUNNEUS <
!EQUIDENS COERULEOPUNCTATUS
!RCHOCENTRUS PANAMENSIS <
'EOPHAGUS CRASSILABRIS <
6IEJA MACULICAUDA NEW
"RACHYRHAPHIS CASCAJALENSIS <
"RACHYRHAPHIS EPISCOPI
.EOHETERANDRIA TRIDENTIGER
0OECILIA GILLII
0OECILIOPSIS ELONGATA <
3! 3MITH AND *" 3HURIN
SPECIES MAINTAIN THEIR DOMINANCE IN DEEPER COOLER WATER 0OR )NVADERS
ALSO CLEARLY PLAY AN IMPORTANT ROLE IN THE COMMUNITY DYNAMICS OF INVADED
REGIONS BOTH IN TERMS OF TOTAL CONTRIBUTION TO THE SPECIES RICHNESS AND ABUNDANCE
"EN
4UVIA &ISHERIES DATA SUGGEST THAT A LARGE PORTION OF NEW IMMIGRANTS
BELONG TO THE COMMON lSHES OF THE ,EVANT BASIN )T IS IMPORTANT TO NOTE
THAT THE 3UEZ #ANAL DATA WAS COLLECTED AT VERY LARGE SPATIAL SCALES THEREFORE MORE
SUBTLE EFFECTS OF INVASIONS MAY NOT BE APPARENT "IOTIC INTERCHANGE FOLLOWING COM
PLETION OF THE 3UEZ #ANAL PROVIDES STRONG EVIDENCE THAT RECIPROCAL INTERCHANGE
INCREASED REGIONAL SPECIES RICHNESS WITHOUT RESULTING IN THE REGIONAL EXTINCTION OF
THE RESIDENT FAUNA
0RE HISTORIC BIOTIC INTERCHANGES
"IOTIC INTERCHANGES HAVE OCCURRED FREQUENTLY OVER THE LAST -Y RESULTING FROM
CLIMATIC CHANGES AND REARRANGEMENT OF CONTINENTAL LANDMASSES THAT BREAK DOWN
BARRIERS TO DISPERSAL 6ERMEIJ A REVIEWED THE OUTCOME OF BIOTIC INTERCHANG
ES THAT OCCURRED DURING THE .EOGENE /NE PATTERN THAT CLEARLY EMERGES IS THAT
COMMUNITIES ARE INVASIBLE HOWEVER INVADING SPECIES IN MOST CASES ONLY MAKE
UP A SMALL PORTION OF THE TOTAL REGIONAL SPECIES POOL IN THE DONOR BIOTA 6ERMEIJ
A &OR EXAMPLE BETWEEN
OF THE SHALLOW
WATER MOLLUSCAN SPECIES
IN THE COOL
TEMPERATE .ORTH 0ACIlC AND OF THE REGIONAL SPECIES POOL OF THE
.ORTH !TLANTIC PARTICIPATED IN THE TRANS
!RCTIC INTERCHANGE 6ERMEIJ B
3IMILAR PATTERNS ARISE IN OTHER INSTANCES OF BIOTIC INTERCHANGES 6ERMEIJ A
4HE LOW APPARENT INVASION RATES FOR SOME PRE
HISTORIC BIOTIC INTERCHANGES COULD
HOWEVER BE THE RESULT OF THE LARGE SPATIAL SCALE AT WHICH THE DATA HAS BEEN RECORD
ED 0ERHAPS MANY SPECIES IN THE DONOR SPECIES POOL ARE NOT ADAPTED TO CONDITIONS
IN THE RECIPIENT BIOGEOGRAPHIC REGION 4HE RICHNESS OF THE NATIVE SPECIES POOL
MAY THEREFORE OVERESTIMATE THE TRUE NUMBER OF SPECIES CAPABLE OF PARTICIPATING IN
THE BIOTIC INTERCHANGE IN QUESTION !LTERNATIVELY THE NATIVE BIOTA MAY REPEL
INVASION BY A LARGE PORTION OF THE POTENTIAL INVADERS )N EITHER CASE THE FOSSIL
RECORD SUGGESTS THAT COMMUNITIES INVOLVED IN FAUNAL EXCHANGES ADDED SPECIES
BUT THAT MANY LIKELY INVASION ATTEMPTS ALSO FAILED
4HE CENTRAL QUESTION PERTAINING TO COMMUNITY SATURATION IS WHETHER BIOTIC
INTERCHANGES RESULT IN EXTIRPATION OF RESIDENT SPECIES 4HE CONCEPT THAT THE ARRIVAL
OF INVADERS FROM A MORE SOPHISTICATED BIOTA MIGHT LEAD TO THE EXTINCTION OF RESIDENT
SPECIES VIA COMPETITIVE DISPLACEMENT HAS A LONG HISTORY 7ALLACE 3IMPSON
7EBB AND -ARSHALL 7EBB -ARSHALL AND #IFELLI
0ATTERNS OF EXTINCTION RESULTING FROM COMPETITIVE DISPLACEMENT WOULD PRESENT
STRONG EVIDENCE FOR THE IMPORTANCE OF LOCAL ECOLOGICAL CONTROL OF COMMUNITY
MEMBERSHIP 4HE EXCHANGE OF MAMMALS ACROSS THE )STHMUS OF 0ANAMA DUR
ING THE 'REAT !MERICAN "IOTIC )NTERCHANGE HAS BEEN CITED AS AN EXAMPLE WHERE
INVADING NORTHERN SPECIES DROVE SOUTHERN RESIDENT SPECIES TO EXTINCTION VIA
COMPETITIVE DISPLACEMENT -ARSHALL "AKKER (OWEVER WHETHER
COMPETITIVE DISPLACEMENT RESULTED IN DIFFERENTIAL RATES OF EXTINCTION BETWEEN
%VIDENCE FOR INVASIBILITY AND SATURATION
.ORTH AND 3OUTH !MERICAN MAMMALS HAS BEEN CONTESTED ,ESSA ET AL
-OREOVER THERE ARE MANY INSTANCES OF INVASIONS THAT DID NOT RESULT IN THE EXTINC
TION OF RESIDENT SPECIES .O INCREASES IN EXTINCTION RATES OCCURRED FOLLOWING BIOTIC
INTERCHANGE IN THE MARINE REALM REGARDING TRANSEQUITORIAL TRANS
!RCTIC AND
TRANS
0ACIlC INTERCHANGES 6ERMEIJ A B 6ERMEIJ A B ,INBERG OR
FOLLOWING THE INVASION OF !FRICAN AND .ORTH !MERICAN MAMMALS INTO !SIA DURING
THE -IOCENE AND 0LIOCENE "ARRY ET AL &LYNN ET AL )NVASION EVENTS
DURING BIOTIC INTERCHANGES MOST OFTEN ENHANCE THE SPECIES RICHNESS OF RECIPIENT
COMMUNITIES "ARRY ET AL &LYNN ET AL 6ERMEIJ A 4HESE PAT
TERNS BOLSTER THOSE OF THE CONTEMPORARY INTERCHANGE LITERATURE IN THAT INVASIONS
RARELY CAUSE THE GLOBAL EXTINCTION OF RESIDENT SPECIES AND OFTEN RESULT IN INCREASES
IN SPECIES RICHNESS AT THE REGIONAL SCALE 4HIS SUGGESTS THAT BIOLOGICAL COMMUNI
TIES DO NOT RESIST ALL INVASION ATTEMPTS AND ARE UNDER
SATURATED WITH SPECIES
4HE OUTCOMES OF BIOTIC INTERCHANGES EMPHASIZE THE IMPORTANCE OF DISPERSAL IN SET
TING AN UPPER LIMIT TO THE NUMBER OF SPECIES COEXISTING IN A GIVEN AREA
)N THIS SECTION WE ARGUE THAT THE OUTCOME OF CONTEMPORARY AND HISTORIC BIOTIC
INTERCHANGES PROVIDE STRONG EVIDENCE THAT COMMUNITIES ARE BOTH INVASIBLE AND
UNSATURATED (OWEVER THE ABOVE EXAMPLES LACK INFORMATION ON THE EFFECTS OF
INVASION AT MULTIPLE SPATIAL SCALES )T IS THEREFORE IMPOSSIBLE TO DETERMINE HOW
CHANGES IN RICHNESS AT CONTINENTAL OR OCEANIC SCALES ARE REmECTED AT SMALLER LOCAL
SCALES )NCREASES IN LARGE
SCALE RICHNESS MIGHT BE ACCOMPANIED BY DECREASES AT
LOCAL SCALES IF NATIVE SPECIES ARE MUCH REDUCED IN RANGE OR ABUNDANCE )N ADDI
TION BETA DIVERSITY OR REGIONAL DISTINCTIVENESS HAS CLEARLY DECLINED IN MANY CASES
AS A RESULT OF BIOTIC HOMOGENIZATION -C+INNEY AND ,OCKWOOD 2AHEL
4HE SPATIAL GRAIN AND EXTENT OF OBSERVATION STRONGLY INmUENCES OUR PERCEP
TION OF COMMUNITY SATURATION
#/.4%-0/2!29 ")/,/')#!, ).6!3)/.3 #/-0!2).' ).6!3)/.
!.$ %84).#4)/. %6%.43
4HE EXPLOSION IN THE NUMBER OF HUMAN
MEDIATED INVASIONS CLEARLY ATTESTS THAT
CONTEMPORARY BIOLOGICAL COMMUNITIES ARE INVASIBLE ,ODGE -OYLE AND
,IGHT 6ITOUSEK ET AL #OHEN AND #ARLTON (OBBS AND -OONEY
2ICCIARDI 3TUDYING COMMUNITY DYNAMICS FOLLOWING INVASIONS PER
MITS INFERENCES ABOUT THE PROCESSES REGULATING SPECIES RICHNESS (ERE WE EXAMINE
EVIDENCE FROM A NUMBER OF CONTEMPORARY INVASIONS AND EXTINCTIONS IN DIFFERENT
ECOSYSTEMS AND ADDRESS THEIR SIGNIlCANCE FOR THE INVASIBILITY AND SATURATION OF
ECOLOGICAL COMMUNITIES
2EGIONAL SCALE PATTERNS
4HE LITERATURE ON CONTEMPORARY ANTHROPOGENIC INVASIONS INDICATES THAT AT BROAD
ISLANDS OR ARCHIPELAGOS INVASIONS
REGIONAL SCALES EG EXCEED EXTINCTIONS
3! 3MITH AND *" 3HURIN
RESULT ING IN NET INCREASES O F SPECIES RICHNESS
#ASE (OBBS AND -OONEY
3AX ET AL 3AX AND 'AINES 4ABLE )NCREASES IN SPECIES RICHNESS
VARIED BETWEEN CONTINENTS AND ISLANDS AS WELL AS AMONG TAXA EXHIBITING DIFFERING
DEGREES OF DISPERSAL LIMITATION ,ARGER INCREASES IN SPECIES RICHNESS WERE ENCOUN
TERED ON ISLANDS AS OPPOSED TO CONTINENTS ESPECIALLY IN THE CASE OF PLANTS WHERE
AN ASTOUNDING INCREASE IN SPECIES RICHNESS WAS DOCUMENTED ON OCEANIC
ISLANDS 3AX ET AL #OMPARING THE INCREASE IN SPECIES RICHNESS AMONG TAXA
INDICATES THAT THE MOST DISPERSAL LIMITED TAXA EXPERIENCED THE LARGEST INCREASES
IN SPECIES RICHNESS FOLLOWING HUMAN MEDIATED INVASION EVENTS &OR EXAMPLE
INCREASES IN THE SPECIES RICHNESS OF FRESHWATER lSHES AND PLANTS FAR EXCEEDED THOSE
OF BIRDS -OST BIRDS CAN DISPERSE OVER MUCH LARGER SPATIAL SCALES THAN FRESHWATER
lSHES WHICH DEPEND ON DIRECT CONNECTIONS BETWEEN WATER BODIES AND mOWERING
PLANTS WHOSE SEEDS ARE PASSIVELY DISPERSED BY WIND OR ANIMALS 3AX ET AL
)T IS THEREFORE NOT SURPRISING THAT DISPERSAL LIMITS THE NUMBER OF lSH AND PLANT SPE
CIES FOUND AT THE REGIONAL SCALE TO A GREATER EXTENT THAN IN BIRDS #ONTEMPORARY
INVASIONS EXCEED EXTINCTIONS SUPPORTING THE EVIDENCE FROM FAUNAL EXCHANGES THAT
COMMUNITIES ARE UNSATURATED
,OCAL SCALE PATTERNS
3TRONG SPECIES INTERACTIONS AT THE LOCAL SCALE MAY NOT MANIFEST THEMSELVES AS SATU
RATION AT BROAD REGIONAL SCALES &OR INSTANCE "YERS AND .OONBURG ANA
LYZED MODELS WITH STRONG LOCAL INTERACTIONS AND SPATIAL HETEROGENEITY IN RESOURCES
4HEY FOUND THAT HIGH DIVERSITY AT THE REGIONAL SCALE DUE TO BROAD HETEROGENEITY LED
TO HIGH INVASIBILITY EVEN THOUGH LOCAL INTERACTIONS GENERATED STRONG BIOTIC RESIS
TANCE AT SMALLER SCALES 3TUDIES OF CHANGES IN SPECIES RICHNESS FOLLOWING INVASION
EVENTS AT THE LOCAL SCALE MAY THEREFORE PROVIDE MORE POWERFUL TESTS FOR COMMU
NITY SATURATION 'IDO AND "ROWN RECORDED CHANGES IN THE SPECIES RICHNESS
OF FRESHWATER lSHES IN DRAINAGE BASINS IN .ORTH !MERICA AND FOUND THAT
INVASIONS EXCEEDED EXTINCTIONS IN OF CASES -EAN SPECIES RICHNESS OF THE
DRAINAGE BASINS INCREASED BY WHILE RICHNESS OF FRESHWATER lSHES IN
#ALIFORNIA INCREASED 4ABLE -ARCHETTI ET AL SUGGESTING THAT LOCAL
COMMUNITIES WERE NOT RESISTANT TO INVASION OR SATURATED WITH SPECIES )N ADDITION
COMPETITION WITH INVADERS HAS RARELY BEEN IMPLICATED AS A CAUSE OF NATIVE SPE
CIES EXTINCTIONS $AVIS 2ATHER PREDATION PRESSURE +AUFMAN &RITTS
AND 2ODDA AND HABITAT CHANGE EG (AWAIIAN )SLANDS ARE RESPONSIBLE FOR
MANY OF THE IMPACTS OF INVADERS ON NATIVES 4HESE RESULTS CONCUR WITH THOSE OF
CONTEMPORARY AND PAST BIOTIC INTERCHANGES WHERE INCREASING THE SIZE OF THE AVAIL
ABLE REGIONAL SPECIES POOL RESULTED IN NET INCREASES IN SPECIES RICHNESS SEE SECTION
!LTERNATIVELY DRAINAGE BASINS MAY CONTAIN MULTIPLE LOCAL POPULATIONS AND
CORRESPOND MORE TO THE REGIONAL THAN LOCAL SCALE SO THAT THE SPATIAL EXTENT IS STILL
TOO LARGE FOR SATURATION TO BECOME EVIDENT .EVERTHELESS PATTERNS OF INVASIONS AND
EXTINCTIONS AT SMALLER SCALES EG DRAINAGES OR ISLANDS PROVIDE NO MORE EVIDENCE
FOR SATURATION THAN THOSE OVER BROADER REGIONS
4ABLE 0ATTERNS OF CHANGES IN SPECIES RICHNESS FOLLOWING INVASION EVENTS $ATA COMPILED FROM (OBBS AND -OONEY 3AX
"ROWN AND 'AINES #ASE 'IDO AND "ROWN -ARCHETTI ET AL AND $UNCAN AND ,OCKWOOD
4AXA ,OCATION -EAN .ET )NCREASE OF SAMPLES OF TOTAL PRESENT 32 3PATIAL SCALE
IN SPECIES RICHNESS THAT IS INTRODUCED
0LANTS #ALIFORNIA 3TATE OF #ALIFORNIA
0LANTS 7ESTERN !USTRALIA 7ESTERN !USTRALIA
0LANTS /CEANIC )SLANDS )SLANDS
-AMMALS #ALIFORNIA 3TATE OF #ALIFORNIA
-AMMALS 7ESTERN !USTRALIA 7ESTERN !USTRALIA
"IRDS #ALIFORNIA 3TATE OF #ALIFORNIA
"IRDS 7ESTERN !USTRALIA 7ESTERN !USTRALIA
,AND "IRDS /CEANIC )SLANDS < )SLANDS
"IRDS )SLANDS )SLANDS
&RESHWATER lSHES #ALIFORNIA 3TATE OF #ALIFORNIA
&RESHWATER lSHES 7ESTERN !USTRALIA 7ESTERN !USTRALIA
&RESHWATER lSHES 53! $RAINAGE BASIN
%VIDENCE FOR INVASIBILITY AND SATURATION
&RESHWATER lSHES #ALIFORNIA $RAINAGE BASIN
&RESHWATER lSHES 4ENNESSEE %COREGIONS
-USSELS 4ENNESSEE < %COREGIONS
2EPTILES #ALIFORNIA 3TATE OF #ALIFORNIA
!MPHIBIANS #ALIFORNIA 3TATE OF #ALIFORNIA
"UTTERmIES #ALIFORNIA < 3TATE OF #ALIFORNIA
3! 3MITH AND *" 3HURIN
!33%33).' -%4(/$/,/')#!, ,)-)4!4)/.3 !.$ #/.342!).43
4HERE ARE SEVERAL IMPORTANT CAVEATS TO CONSIDER WHEN INFERRING THE MECHANISMS
STRUCTURING COMMUNITIES FROM PATTERNS OF INVASION AND EXTINCTION AT LOCAL AND
REGIONAL SCALES &OR EXAMPLE EXTINCTIONS MAY BE MORE DIFlCULT TO DETECT THAN
INVASIONS &EW EXTINCTIONS MAY BE RECORDED AS SPECIES THAT ARE NOT OBSERVED
CANNOT BE ASSUMED ABSENT WHEREAS INVASIONS ARE UNAMBIGUOUSLY INVASIONS
)N ADDITION CHANGES IN SPECIES RICHNESS MAY IGNORE OTHER LARGE EFFECTS ON RANGE OR
ABUNDANCE OF NATIVE SPECIES FOLLOWING SPECIES INVASIONS 4HAT IS MANY FORMERLY
ABUNDANT WIDESPREAD SPECIES MAY BE REDUCED TO FEW SMALL REMNANT POPULATIONS
BY INTERACTIONS WITH INVADERS WITHOUT BECOMING EXTINCT 3UCH CHANGES IN DISTRI
BUTION AND ABUNDANCE ARE NOT CAPTURED BY PRESENCEABSENCE DATA
%XTINCTIONS CAUSED BY INVADERS MAY ALSO PROCEED SLOWLY AND OCCUR LONG AFTER
THE INITIAL COLONIZATION 2ICKLEFS SUGGESTING THAT THERE MAY BE A TIME LAG
BETWEEN INVASION AND EXTINCTION EVENTS (OWEVER INVASIONS FOLLOWING THE ELIMI
NATION OF DISPERSAL BARRIERS DO NOT RESULT IN MEASURABLY GREATER EXTINCTION RATES
OVER EVOLUTIONARY TIME SCALES "ARRY ET AL &LYNN ET AL ,INBERG ET AL
.EVERTHELESS EXAMPLES OF INVASIONS THAT LEAD TO MASS EXTINCTION EVENTS
ARE WELL DOCUMENTED EG .ILE PERCH IN ,AKE 6ICTORIA +AUFMAN BROWN
TREE SNAKE IN 'UAM &RITTS AND 2ODDA AND ZEBRA MUSSELS IN THE 'REAT ,AKES
.ALEPA AND 3CHLOESER 6ANDERPLOEG ET AL
%LEVATED RATES OF SPECIES INTRODUCTIONS COUPLED WITH LARGE CORRELATED LOSSES OF
DIVERSITY HAVE LED TO THE ASSUMPTION THAT EXOTIC SPECIES INTRODUCTIONS ARE ONE OF
THE LEADING CAUSES OF SPECIES EXTINCTIONS BUT SEE 'UREVITCH AND 0ADILLA
4HE OBSERVATION THAT EXOTIC SPECIES ARE THE THIRD MOST PREVALENT THREAT TO ENDAN
GERED SPECIES THROUGHOUT THE WORLD 3ALA ET AL SUGGESTS THAT MANY EXTINC
TIONS RESULTING FROM BIOTIC EXCHANGE ARE STILL TO COME (OWEVER IT HAS OFTEN BEEN
DIFlCULT TO IDENTIFY THE ROLE OF EXOTIC INVADERS IN CAUSING SPECIES EXTINCTIONS BECAUSE
MOST IMPERILED SPECIES FACE MORE THAN ONE THREAT SIMULTANEOUSLY 'UREVITCH AND
0ADILLA %XPERIMENTS THAT DO NOT SUFFER FROM THE CONFOUNDING EFFECTS OF
MULTIPLE ASPECTS OF HUMAN ALTERATION OF THE ENVIRONMENT ARE REQUIRED TO ASSESS THE
RELATIVE IMPORTANCE OF INVASIONS AS A CAUSE OF EXTINCTION 2ICCIARDI
&INALLY INVASIVE SPECIES HAVE BEEN DOCUMENTED TO PREFERENTIALLY OCCUPY
HUMAN DOMINATED LANDSCAPES WHEREAS NATIVE SPECIES HAVE A GREATER AFlNITY
FOR NATIVE UNDISTURBED HABITAT $IAMOND AND 6EITECH 3IMBERLOFF
3MALLWOOD #ASE 2AHEL 3AX ET AL $UNCAN ET AL
-ANY INVADERS MAY NOT INTERACT WITH RESIDENT SPECIES BECAUSE OF HABITAT PARTI
TIONING BETWEEN NATIVE AND INTRODUCED SPECIES THEREFORE COMPLICATING THE USE
OF BIOLOGICAL INVASIONS AS TESTS OF COMMUNITY SATURATION )N ADDITION THE HIGH
APPARENT INVASIBILITY OF COMMUNITIES MAY BE AS MUCH A PRODUCT OF ANTHROPOGENIC
DISTURBANCES THAT WEAKEN BIOTIC RESISTANCE AS HUMAN
AIDED DISPERSAL 3EABLOOM
ET AL .EVERTHELESS NET INCREASES OF SPECIES RICHNESS AT LOCAL AND REGIONAL
SCALES FOLLOWING INVASION EVENTS SUGGEST THAT MANY COMMUNITIES ARE INVASIBLE
AND UNSATURATED 4HIS CONCLUSION IS SUPPORTED BY PATTERNS OF PAST AND CONTEM
%VIDENCE FOR INVASIBILITY AND SATURATION
PORARY FAUNAL INTERCHANGE )N THE NEXT SECTION WE REVIEW EXPERIMENTAL STUDIES
OF COMMUNITY SATURATION AND COMPARE THEIR RESULTS WITH THOSE OF PATTERNS FROM
UNMANIPULATED SYSTEMS
%80%2)-%.4!,,9 34!'%$ ).6!3)/.3
-ACROECOLOGICAL PATTERNS SUGGEST THAT COMMUNITIES ARE BOTH UNSATURATED AND
INVASIBLE THAT BIOTIC RESISTANCE TO INVASION IN COMMUNITIES IS WEAK RELATIVE TO
DISPERSAL LIMITATION AND THAT NATIVE SPECIES ARE RARELY EXCLUDED FROM COMMUNI
TIES BY INTERACTIONS WITH INVADERS 3TOHLGREN ET AL 3AX AND 'AINES
4HIS CONCLUSION HAS IMPORTANT PRACTICAL IMPLICATIONS AS IT SUGGESTS THAT MANAG
ING DISPERSAL VECTORS IS MORE EFFECTIVE FOR CONTAINING INVADERS THAN RELYING ON
NATIVE COMMUNITIES TO PREVENT THEIR SPREAD (OWEVER THE CONCLUSION THAT SPECIES
INTERACTIONS AND BIOTIC RESISTANCE ARE UNIMPORTANT FOR SHAPING COMMUNITIES OR
PREVENTING INVASIONS MAY BE INCORRECT &IRST SPECIES INTERACTIONS LIKELY OCCUR AT
SCALES MUCH SMALLER THAN THE BROAD REGIONS EG ISLAND CHAINS OR WATERSHEDS
THAT ARE THE UNITS OF OBSERVATION FOR BIOGEOGRAPHICAL STUDIES ! NEW COLONIST
LIKELY INTERACTS ONLY WITH OTHER SPECIES WITHIN A FAIRLY SMALL NEIGHBORHOOD NOT
THE ENTIRE BIOTA OF THE REGION )N ADDITION WHILE INVASIONS ARE OFTEN DRAMATIC
AND OBVIOUS FAILED INVASION ATTEMPTS MOSTLY PASS UNOBSERVED 7E DO NOT KNOW
WHETHER THE REGIONAL DIFFERENCES IN DIVERSITY THAT REMAIN FOLLOWING WHOLESALE
SPECIES INTRODUCTIONS ARE DUE TO ENVIRONMENTAL CONSTRAINTS SPECIES INTERACTIONS
OR LACK OF DISPERSAL OPPORTUNITIES FOR SOME SPECIES &INALLY EXTINCTION OF SPECIES
MAY BE SLOW RELATIVE TO INVASION 2ICKLEFS AND SOME SPECIES MAY BE GREATLY
REDUCED IN ABUNDANCE OR GEOGRAPHIC RANGE BY INVADERS WITHOUT BEING COMPLETELY
EXTIRPATED 2EGIONAL DIVERSITY LOSS DUE TO INVADERS MAY THEREFORE BE A WEAK INDI
CATOR OF THE IMPORTANCE OF BIOTIC EXCLUSION ,OCAL INTERACTIONS MAY BE DIFlCULT TO
DETECT IN MACROECOLOGICAL PATTERNS BUT STILL BE CRITICAL FOR CONSTRAINING INVASIONS
4HE lNAL APPROACH TO ASSAYING COMMUNITY SATURATION AND INVASIBILITY THAT
WE DISCUSS IS THROUGH EXPERIMENTAL SPECIES INTRODUCTIONS &IELD MANIPULATIONS OF
INVASION OFFER A NUMBER OF INSIGHTS THAT CANNOT BE OBTAINED FROM OBSERVATIONAL
DATA &IRST THE RATES OF SUCCESS AND FAILURE CAN BE ACCURATELY CHARACTERIZED AS
THE POOL OF INTRODUCED SPECIES IS KNOWN 3ECOND ASPECTS OF THE INVADER POOL EG
PROPAGULE LOAD OR TIMING AND RECIPIENT COMMUNITY EG DIVERSITY COMPOSITION
PRODUCTIVITY OR DISTURBANCE CAN BE DIRECTLY MANIPULATED 4HESE ALLOW ASSIGN
MENT OF THE CAUSES OF INVASION SUCCESS OR FAILURE AND INTERPRETATION OF THE TRAITS
OF SPECIES OR COMMUNITIES THAT INmUENCE INVASIVENESS AND INVASIBILITY &IELD
EXPERIMENTS ARE ALSO GENERALLY CONDUCTED AT MUCH SMALLER LOCAL SCALES SUCH AS THE
NEIGHBORHOOD OF PLANT COMMUNITIES AND MAY THEREFORE BE MORE LIKELY TO REVEAL
THE ROLE OF SPECIES INTERACTIONS
%FFORTS TO EXPERIMENTALLY TEST COMMUNITY SATURATION ARE INCREASINGLY COMMON
!SSAYING INVASIBILITY AT THE COMMUNITY LEVEL INVOLVES INTRODUCING POTENTIALLY
INVASIVE SPECIES USUALLY ONES THAT ARE PRESENT REGIONALLY BUT ABSENT LOCALLY 4HE
3! 3MITH AND *" 3HURIN
MAJORITY OF lELD STUDIES OF COMMUNITY SATURATION ARE SEED ADDITIONS IN HERBACEOUS
PLANT COMMUNITIES 4ABLE 4URNBULL ET AL REVIEWED THE LITERATURE ON
SEED ADDITION EXPERIMENTS AND CONCLUDED THAT APPROXIMATELY OF ALL PLANT SPE
CIES SHOW EVIDENCE FOR SEED LIMITATION ! SESSILE GROWTH FORM MAY RENDER TERRES
TRIAL PLANTS ESPECIALLY VULNERABLE TO DISPERSAL LIMITATION AS PROPAGULES HAVE ONLY
ONE OPPORTUNITY TO lND SUITABLE HABITAT PATCHES 3MALL SCALE VARIABILITY IN LIGHT OR
SOIL CONDITIONS MAY CONSTRAIN APPROPRIATE SITES FOR GERMINATION TO A SMALL FRACTION
OF THE AVAILABLE HABITAT ! LIMITED NUMBER OF STUDIES HAVE BEEN PERFORMED IN OTHER
COMMUNITIES INCLUDING POND ZOOPLANKTON 3HURIN AND LABORATORY STUDIES
WITH PROTOZOANS &OX ET AL
4ABLE ! SUMMARY OF EXPERIMENTAL STUDIES OF COMMUNITY SATURATION AND INVASIBILITY
0ERCENT INTRODUCTION SUCCESS IS MEASURED AS THE PROPORTION OF INTRODUCED SPECIES OBSERVED
IN THE EXPERIMENTAL SYSTEM AND INCLUDES SUCCESSFUL GERMINATION h)NCREASED LOCAL RICHNESSv
INDICATES WHETHER LOCAL RICHNESS WAS GREATER IN THE INTRODUCTION TREATMENT THAN THE CONTROL
h2ICHNESS
INVASIBILITYv INDICATES WHETHER THE SUCCESS OF INVADERS DECLINED AT HIGHER NATIVE
SPECIES RICHNESS AND hDISTURBANCE TREATMENTv INDICATES WHETHER PERTURBATIONS TO THE
NATIVE COMMUNITY AFFECTED INVASIBILITY ! DASH INDICATES THAT THE TREATMENT WAS NOT
PERFORMED OR THE HYPOTHESIS WAS NOT TESTED
3TUDY 3YSTEM )NCREASED 2ICHNESS
$ISTURBANCE
INTRODUCTION LOCAL
SUCCESS RICHNESS INVASIBILITY TREATMENT
"URKE AND 'RIME GRASSLAND < < 9
4ILMAN GRASSLAND 9 NEGATIVE <
3HURIN ZOOPLANKTON . NEGATIVE 9
:OBEL ET AL GRASSLAND 9 NONE 9
&OSTER GRASSLAND 9 < 9
,EVINE SEDGES 9 NEGATIVE 9
&OSTER ET AL GRASSLAND < POSITIVE <
&ARGIONE ET AL GRASSLAND 9 NEGATIVE <
&OSTER AND 4ILMAN GRASSLAND 9 < <
-OUQUET ET AL GRASSLAND 9 < <
4ABLE SUMMARIZES EXPERIMENTS TESTING SATURATION AND REVEALS A NUMBER OF
PATTERNS &IRST EVERY EXPERIMENTAL SPECIES ADDITION FOUND THAT SOME SPECIES ARE
ABSENT FROM LOCAL COMMUNITIES DUE TO DISPERSAL LIMITATION 4HE INTRODUCTION SUC
CESS RATE INTO INTACT RECIPIENT COMMUNITIES RANGED FROM FOR POND ZOOPLANKTON
TO MUCH HIGHER IN HERBACEOUS PLANT COMMUNITIES 4HE OBSERVATION
%VIDENCE FOR INVASIBILITY AND SATURATION
THAT ZOOPLANKTON COMMUNITIES APPEAR CLOSER TO SATURATION THAN PLANTS MAY REmECT
EITHER BIOLOGICAL OR METHODOLOGICAL DIFFERENCES BETWEEN THE STUDIES :OOPLANKTON
MAY BE MORE EFFECTIVE DISPERSERS THAN PLANTS OR MAY BE BETTER ABLE TO lND SUITABLE
MICROSITES WITHIN PATCHES EG PONDS BECAUSE THEY ARE MOTILE !LTERNATIVELY THE
CRITERION FOR SUCCESSFUL INVASION IN THE PLANT STUDIES WAS TYPICALLY GERMINATION
NOT SUCCESSFUL REPRODUCTION OR POSITIVE POPULATION GROWTH -ANY OF THE SEEDLINGS
THAT GERMINATED MAY NOT HAVE EVENTUALLY ESTABLISHED THEREFORE THE EXPERI
MENTS MAY OVERESTIMATE INVASION SUCCESS (OWEVER A NUMBER OF STUDIES LASTED
FOR SEVERAL YEARS AND FOUND THAT INTRODUCED PLANTS PERSISTED 4HE RESULTS OF THE
EXPERIMENTS SUPPORT LARGE
SCALE PATTERNS INDICATING THAT CONTEMPORARY BIOLOGICAL
COMMUNITIES ARE INVASIBLE
%XPERIMENTS ALSO INDICATE THAT PLANT COMMUNITIES ARE OFTEN UNSATURATED IN
THAT INTRODUCTIONS SIMULATING INCREASED CONNECTIVITY TO THE LARGER REGIONAL SPE
CIES POOL ENHANCED LOCAL DIVERSITY 3EVEN OUT OF EIGHT STUDIES FOUND HIGHER SPE
CIES RICHNESS IN THE INTRODUCTION TREATMENTS THAN IN THE CONTROLS ALL OF THE PLANT
EXPERIMENTS THAT CONTRASTED DIVERSITY IN THE SEED ADDITION TREATMENT WITH THE CON
TROL 4ABLE "Y CONTRAST 3HURINS POND ZOOPLANKTON EXPERIMENTS FOUND
NO DIFFERENCE IN DIVERSITY BETWEEN INVASION TREATMENTS AND THE CONTROL INDICATING
THAT POND ZOOPLANKTON COMMUNITIES MAY BE CLOSER TO REGIONAL SATURATION THAN
TERRESTRIAL PLANTS %XPERIMENTAL INTRODUCTIONS INDICATE THAT MANY COMMUNITIES
ARE UNSATURATED WITH SPECIES AT BOTH LOCAL NEIGHBORHOOD AND REGIONAL SCALES
(OWEVER A NUMBER OF PATTERNS FROM THE EXPERIMENTAL LITERATURE CONTRAST WITH
THE INTERPRETATION OF MACRO
SCALE PATTERNS AS INDICATING WEAK BIOTIC RESISTANCE AND
A DOMINANCE OF DISPERSAL LIMITATION RELATIVE TO SPECIES INTERACTIONS IN STRUCTURING
COMMUNITIES EG 3TOHLGREN ET AL 3AX AND 'AINES &IRST A NUMBER
OF STUDIES FOUND DECLINING INVASION SUCCESS MEASURED EITHER AS THE PERCENT OCCUR
RENCE OF INVASIVE SPECIES OR THEIR OVERALL ABUNDANCE AT HIGHER LOCAL RICHNESS OF
RESIDENT SPECIES 4ABLE 4HIS RESULT SUGGESTS THAT ALTHOUGH COMMUNITIES ARE
UNSATURATED IN THAT THEY CAN SUPPORT MORE SPECIES LOCAL RICHNESS APPROACHES
SATURATION AT HIGH LEVELS IN THAT IT BECOMES INCREASINGLY DIFlCULT FOR ADDITIONAL
SPECIES TO INVADE )NCREASING BIOTIC RESISTANCE MOST LIKELY RESULTS FROM GREATER
INTENSITY OF LOCAL INTERACTIONS AT HIGH DIVERSITY 4HE INCREASED BIOTIC RESISTANCE
AT HIGH DIVERSITY COULD BE THE RESULT OF EITHER STRONGER COMPETITIVE INTERACTIONS
AMONG ESTABLISHED SPECIES OR BY THE INHIBITORY EFFECTS OF ESTABLISHED SPECIES ON
RECRUITMENT 4ILMAN
3ECOND A NUMBER OF EXPERIMENTS EMPLOYED DISTURBANCE TREATMENTS THAT WEAK
ENED INTERACTIONS BETWEEN THE RESIDENT COMMUNITY AND INVADERS #OMPARING
INVASIBILITY IN DISTURBED VS INTACT NATIVE COMMUNITIES ILLUSTRATES WHICH SPECIES
ARE EXCLUDED FROM THE RECIPIENT COMMUNITY DUE TO INTERACTIONS WITH NATIVES VER
SUS INABILITY TO TOLERATE LOCAL ABIOTIC CONDITIONS 3TUDIES EMPLOYING DISTURBANCE
TREATMENTS EITHER THROUGH REMOVAL OF NATIVE SPECIES OR ENHANCED PRODUCTIVITY
BY NUTRIENT ADDITIONS FOUND THAT INVASIBILITY WAS DRAMATICALLY INCREASED BY ALTER
ING THE BIOTIC ENVIRONMENT &OR INSTANCE 3HURIN FOUND THAT REDUCING THE
ABUNDANCE OF NATIVE SPECIES ALLOWED FOUR TIMES AS MANY INVADERS TO ESTABLISH
3! 3MITH AND *" 3HURIN
AND TO OBTAIN SIXTEEN TIMES GREATER TOTAL ABUNDANCE 4HESE RESULTS INDICATE THAT
ALTHOUGH COMMUNITIES MAY BE INVASIBLE AND UNSATURATED DUE TO DISPERSAL LIMITA
TION LOCAL INTERACTIONS MAY STILL BE STRONG ENOUGH TO EXCLUDE A LARGE FRACTION OF
POTENTIAL INVADERS AND THAT MANY MORE SPECIES MAY BE EXCLUDED THAN SUCCEED
IN INVADING INTACT COMMUNITIES 4HIS CONTRAST ILLUSTRATES THE IMPORTANCE OF lELD
EXPERIMENTS !NALYSIS OF PATTERNS OF DIVERSITY FOLLOWING FAUNAL EXCHANGE AND
ANTHROPOGENIC INTRODUCTIONS MAY BE INTERPRETED TO INDICATE THAT BIOTIC RESISTANCE
SELDOM IF EVER REPELS INVADERS 3TOHLGREN ET AL 3AX AND 'AINES
4HIS CONCLUSION EMPHASIZES SUCCESSFUL INVASIONS OVER FAILURES HOWEVER THE LATTER
MAY BE MORE COMMON AND OFTEN OCCUR DUE TO LOCAL INTERACTIONS
-%4!#/--5.)49 02%$)#4)/.3
4HE GLOBAL EXPLOSION IN THE INCIDENCE OF EXOTIC SPECIES REmECTS THE RESULTS OF HUMAN
ACTIVITIES THAT INCREASE RATES OF MIGRATION AMONG PREVIOUSLY ISOLATED BIOGEOGRAPH
IC REGIONS 7ITHIN REGIONS HOWEVER HUMANS MAY EITHER INCREASE HABITAT CONNEC
TIVITY BY ACTING AS DISPERSAL VECTORS OR DECREASE IT THROUGH HABITAT FRAGMENTATION
4HE EFFECTS OF MODIFYING RATES OF DISPERSAL WITHIN REGIONS AND IMMIGRATION FROM
OTHER REGIONS ARE QUITE DISTINCT (ERE WE FOCUS ON THE EFFECTS OF INCREASED MOVE
MENT RATES AMONG REGIONS PUTTING ASIDE THE QUESTION OF HOW HUMAN
INDUCED
ENVIRONMENTAL CHANGE AFFECTS DISPERSAL WITHIN REGIONS -ETACOMMUNITY MODELS
OFFER DIFFERENT THEORETICAL PREDICTIONS REGARDING THE INVASIBILITY AND SATURATION OF
LOCAL COMMUNITIES AND THE OUTCOME OF BIOLOGICAL INVASIONS #OMPARING RESPONSES
OF DIVERSITY TO CHANGES IN IMMIGRATION RATES WITH PREDICTED OUTCOMES MAY BE USE
FUL FOR EVALUATING DIFFERENT PERSPECTIVES ON BROAD
SCALE COMMUNITY STRUCTURE
4HEORETICAL AND EMPIRICAL WORK ON METACOMMUNITIES FALLS UNDER FOUR PARA
DIGMS WHICH WE CALL PATCH
DYNAMICS NEUTRAL SPECIES
SORTING AND MASS
EFFECTS ,EIBOLD ET AL #HASE ET AL 4HE FOUR APPROACHES DIFFER IN
THEIR ASSUMPTIONS ABOUT RELATIVE RATES OF DISPERSAL THE INTENSITY OF INTERSPECIlC
INTERACTIONS AND THE DEGREE OF SPATIAL HETEROGENEITY AMONG HABITATS &IG
0ATCH DYNAMICS THE OLDEST PERSPECTIVE IS REPRESENTED BY ISLAND BIOGEOGRAPHY
AND METAPOPULATION ECOLOGY 4HIS APPROACH CONSIDERS HABITATS AS DIVIDED INTO
HOMOGENEOUS PATCHES AMONG WHICH SPECIES DISPERSE 4HE SIMPLEST VERSION OF THE
PATCH
DYNAMICS FRAMEWORK ASSUMES THAT SPECIES ARE INDIFFERENT TO ONE ANOTHERS
PRESENCE IN PATCHES HOWEVER SUCH MODELS HAVE BEEN MODIlED TO INCORPORATE
LOCAL INTERACTIONS THAT LEAD TO EXCLUSION OR FACILITATION ,EVINS AND #ULVER
(ASTINGS $ISPERSAL IN THE PATCH
DYNAMICS FRAMEWORK IS SLOW RELATIVE TO
DEMOGRAPHIC RATES SO THAT PATCHES ARE EITHER OCCUPIED OR EMPTY BUT LOCAL DYNAM
ICS ARE IGNORED .EUTRAL THEORY "ELL (UBBELL REPRESENTS A VARIATION
ON PATCH DYNAMICS WHERE ALL INDIVIDUALS ARE IDENTICAL IN ALL RELEVANT TRAITS LOCAL
INTERACTIONS ARE STRONG AND LEAD TO PRIORITY EFFECTS AND DISPERSAL IS SPATIALLY CON
STRAINED 3PECIES MAY COEXIST IN PATCH
DYNAMICS MODELS AT EQUILIBRIUM BY TRADE
OFFS IN THEIR ABILITY TO COLONIZE AND COMPETE IN PATCHES ,EVINS AND #ULVER
%VIDENCE FOR INVASIBILITY AND SATURATION
(ASTINGS WHILE COEXISTENCE IN NEUTRAL MODELS IS DETERMINED BY A BALANCE
BETWEEN SLOW EXCLUSION AND THE INPUT OF SPECIES EITHER FROM OUTSIDE THE REGION OR
THROUGH SPECIATION "ELL (UBBELL
4HE SPECIES
SORTING APPROACH POSITS THAT SPECIES DISPERSE SUFlCIENTLY RAPIDLY
THAT LOCAL ENVIRONMENTAL CONSTRAINTS LIMIT DISTRIBUTIONS TO A GREATER EXTENT THAN
THE SUPPLY OF COLONISTS ,EIBOLD ET AL 4HE SPECIES
SORTING VIEW CAN BE
CONSIDERED CLASSICALLY NICHE
BASED IN THAT SPECIES ARE SELECTED BY THE LOCAL ENVI
RONMENT AND SITES CONTAIN MOST OR ALL SPECIES CAPABLE OF COEXISTING UNDER THE
PARTICULAR LOCAL CONDITIONS 3PECIES
SORTING ASSUMES SUFlCIENTLY FAST DISPERSAL THAT
LOCAL INTERACTIONS PLAY A DOMINANT ROLE OVER MOVEMENT OF SPECIES IN CONSTRAIN
ING COMMUNITY MEMBERSHIP &INALLY THE MASS
EFFECTS OR SOURCE
SINK PERSPECTIVE
HOLDS THAT DISPERSAL CAN BE SO RAPID THAT POPULATIONS ARE MAINTAINED THAT WOULD
OTHERWISE DECLINE TO EXTINCTION IN THE ABSENCE OF INPUT OF INDIVIDUALS FROM OTHER
HABITATS 3HMIDA AND %LLNER -ASS
EFFECTS MODELS RELY ON HETEROGENEITY
IN THE ENVIRONMENT THAT CREATES DIFFERENCES IN lTNESS SUCH THAT SOME HABITATS
PRODUCE EXCESS INDIVIDUALS THAT DISPERSE TO POOR
QUALITY PATCHES WHERE THEY
HAVE NEGATIVE GROWTH RATES 3PECIES
SORTING HOLDS THAT DISPERSAL IS FAST RELATIVE
TO RATES OF POPULATION EXTINCTION BUT NOT SO RAPID THAT THE INPUT OF INDIVIDUALS IS
LARGE RELATIVE TO LOCAL REPRODUCTION "Y CONTRAST MASS
EFFECTS SYSTEMS OCCUR WHEN
DISPERSAL IS FAST RELATIVE TO LOCAL DEMOGRAPHIC RATES 4HE DIFFERENCES AMONG THE
FOUR METACOMMUNITY APPROACHES ARE ILLUSTRATED IN &IG
4HE FOUR METACOMMUNITY PERSPECTIVES MAKE DIVERGENT PREDICTIONS REGARD
ING THE INVASIBILITY AND SATURATION OF COMMUNITIES AND THE PREDICTED OUTCOME
OF INCREASED IMMIGRATION RATES AMONG BIOGEOGRAPHIC PROVINCES 4ABLE #HASE
ET AL &OR EXAMPLE PATCH
DYNAMICS MODELS WITHOUT INTERACTIONS PREDICT
THAT LOCAL COMMUNITIES ARE ALWAYS INVASIBLE AND UNSATURATED #ONSEQUENTLY
INVASIONS FOLLOWING INCREASED IMMIGRATION RATES SHOULD BE COMMONPLACE 0ATCH
DYNAMIC MODELS WITH STRONG COMPETITIVE EXCLUSION PREDICT THAT MORE SPECIES CAN
ALWAYS BE ADDED REGIONALLY AS LONG AS THEY SHOW SUFlCIENTLY STRONG COMPETITION
COLONIZATION TRADEOFFS 4ILMAN (OWEVER THE NECESSARY TRADEOFF BECOMES
INCREASINGLY STRINGENT AS REGIONAL RICHNESS INCREASES THEREFORE COMMUNITIES
BECOME INCREASINGLY RESISTANT TO INVASION AS MORE SPECIES ARE ADDED TO THE REGION
3HURIN AND !LLEN .EUTRAL MODELS PREDICT THAT SPECIES CAN ALWAYS INVADE
LOCAL COMMUNITIES REGARDLESS OF LOCAL DIVERSITY AND THE RATE OF LOCAL DISPERSAL
BECAUSE THE DEMOGRAPHIC PROPERTIES OF INDIVIDUALS ARE EQUIVALENT #ONSEQUENTLY
RESIDENT SPECIES ARE UNABLE TO EXCLUDE INVADERS FROM LOCAL COMMUNITIES 3PECIES
SORTING MODELS PREDICT THAT DISPERSAL IS FREQUENT ENOUGH TO ALLOW LOCAL ASSEMBLY
PROCESSES TO REACH THEIR @END
POINT TRAJECTORIES AND THAT COMMUNITIES THEREFORE
RESIST FURTHER INVASIONS 3OURCE
SINK MODELS PREDICT THAT COMMUNITIES BECOME
HARDER TO INVADE WITH INCREASING DISPERSAL BEYOND THE MAXIMUM LOCAL RICHNESS
POINT ,OCAL RICHNESS PLATEAUS UNDER MODERATE LEVELS OF LOCAL DISPERSAL -OUQUET
AND ,OREAU ABOVE WHICH COMMUNITIES BECOME INCREASINGLY DIFlCULT TO
INVADE BECAUSE BETTER REGIONAL COMPETITORS DISPERSERS ELIMINATE OTHER SPECIES
#HASE ET AL
3! 3MITH AND *" 3HURIN
&IG 4HE RELATIONSHIPS AMONG THE FOUR MAJOR METACOMMUNITY PARADIGMS IN TERMS OF
THEIR ASSUMPTIONS ABOUT INTERACTION INTENSITY SPATIAL HETEROGENEITY AND DISPERSAL 0ATCH
DYNAMICS MODELS ASSUME SLOW DISPERSAL RELATIVE TO EXTINCTION PROBABILITIES AND EITHER
INTERACTIVE EG COMPETITION
COLONIZATION TRADEOFFS OR NON
INTERACTIVE COMMUNITIES
3PECIES SORTING ASSUMES DISPERSAL IS FAST RELATIVE TO EXTINCTION BUT SLOW RELATIVE TO
LOCAL DEMOGRAPHIC RATES HIGH SPATIAL HETEROGENEITY AND STRONG INTERACTIONS .EUTRAL
MODELS ASSUME A CONTINUUM OF DISPERSAL RATES STRONG LOCAL INTERACTIONS WITH PRE
EMPTIVE
COMPETITION AND HOMOGENEOUS PATCHES 3OURCE
SINK MODELS ASSUME THAT DISPERSAL IS RAPID
RELATIVE TO LOCAL DYNAMICS HIGH SPATIAL HETEROGENEITY IN lTNESS AMONG PATCHES AND STRONG
LOCAL INTERACTIONS
-ETACOMMUNITY MODELS ALSO DIFFER IN THEIR PREDICTIONS ABOUT COMMUNITY SATU
RATION "OTH NON
INTERACTIVE PATCH DYNAMIC MODELS AND NEUTRAL THEORY PREDICT
THAT LOCAL DIVERSITY IS NEVER SATURATED AND NEW SPECIES CAN ALWAYS BE ADDED TO
LOCAL COMMUNITIES !T HIGH LEVELS OF LOCAL DISPERSAL AND SPECIES RICHNESS HOWEVER
NEUTRAL THEORY PREDICTS THAT GREATER IMMIGRATION WILL INCREASE THE RATE AT WHICH
COMMUNITY COMPOSITION TURNS OVER IN LOCAL COMMUNITIES DUE TO FASTER EXTINCTION
OF RESIDENT SPECIES )F MORE SPECIES ARE ADDED TO LOCAL COMMUNITIES THE AVERAGE
POPULATION SIZE OF RESIDENT SPECIES MUST DECREASE UNDER ZERO
SUM DYNAMICS
RENDERING RARE SPECIES MORE SUSCEPTIBLE TO STOCHASTIC mUCTUATIONS IN POPULATION
SIZES )N CONTRAST SPECIES
SORTING PATCH
DYNAMICS WITH INTERACTIONS AND MASS
EFFECTS MODELS ALL PREDICT SATURATION OF DIVERSITY )N INTERACTIVE PATCH
DYNAMICS
AND MASS
EFFECTS MODELS LOCAL COMMUNITIES APPROACH SATURATION WITH INCREASED
%VIDENCE FOR INVASIBILITY AND SATURATION
DISPERSAL BECAUSE IT BECOMES MORE DIFlCULT FOR NEW SPECIES TO SATISFY THE NECES
SARY TRADE
OFF BETWEEN DISPERSAL ABILITY AND THE POTENTIAL TO EXCLUDE OTHER SPECIES
3HURIN AND !LLEN -OREOVER IN MASS
EFFECTS MODELS SOURCE
SINK DYNAM
ICS BREAKDOWN AT HIGH DISPERSAL RATES AND BETTER REGIONAL COMPETITORS ELIMINATE
OTHER SPECIES -OUQUET AND ,OREAU )N PURE SPECIES
SORTING MODELS WITHOUT
DISPERSAL LIMITATION EXOTIC SPECIES ARE UNLIKELY TO BE ABLE TO INVADE BECAUSE THE
COMMUNITY IS COLLECTIVELY WELL
ADAPTED TO LOCAL CONDITIONS ,OCAL COMMUNITIES ARE
THEREFORE ASSUMED TO BE SATURATED UNDER THE SPECIES SORTING PARADIGM
4ABLE 0REDICTIONS OF THE MAJOR METACOMMUNITY PARADIGMS REGARDING THE
INVASIBILITY AND SATURATION OF ECOLOGICAL COMMUNITIES
0ARADIGM )NVASIBLE 3ATURATED
0ATCH DYNAMICS 9ES .O
NON INTERACTIVE
0ATCH DYNAMICS WITH 9ES BUT INCREASINGLY 9ES INCREASINGLY DIFlCULT AT
STRONG COMPETITIVE DIFlCULT WITH INCREASING HIGH DIVERSITY TO SATISFY
EXCLUSION DIVERSITY COLONIZATIONCOMPETITION
TRADE OFF
.EUTRAL 9ES .O BUT COMMUNITY
COMPOSITION TURNS OVER MORE
RAPIDLY WITH INCREASING SPECIES
RICHNESS
3PECIES SORTING .O 9ES
-ASS EFFECTS 9ES BUT INCREASINGLY 9ES INCREASINGLY DIFlCULT AT
DIFlCULT WITH INCREASING HIGH DIVERSITY AND DISPERSAL TO
DIVERSITY AND FREQUENCY SATISFY COLONIZATION
OF LOCAL DISPERSAL COMPETITION TRADE OFF
0ATTERNS OF INVASIBILITY AND SATURATION FROM THE BIOLOGICAL INVASIONS LITERATURE
ARE USEFUL FOR EVALUATING THE ABILITY OF THE FOUR METACOMMUNITY PARADIGMS TO
DESCRIBE COMMUNITY STRUCTURE AND DYNAMICS 4HE lNDING THAT MOST COMMUNI
TIES ARE INVASIBLE AND UNSATURATED IS CONSISTENT WITH THE PREDICTIONS OF ANY OF THE
MODELS ASSUMING DISPERSAL LIMITATION PATCH
DYNAMICS NEUTRAL AND MASS
EFFECTS
3! 3MITH AND *" 3HURIN
AND CHALLENGES THE PREDICTION OF SPECIES SORTING THAT COMMUNITIES RESIST INVASION
BECAUSE THEY ARE SATURATED WITH SPECIES 4HE OBSERVATION OF MANY INTRODUCTIONS
AND FEW EXTINCTIONS IS INCONSISTENT WITH MODELS WHERE REGIONAL COEXISTENCE OCCURS
THROUGH SPATIAL REFUGIA INTERACTIVE PATCH DYNAMICS AND MASS
EFFECTS 4HESE
MODELS PREDICT THAT ENHANCED IMMIGRATION CAN RESULT IN LOSS OF DIVERSITY THROUGH
GREATER EXCLUSION BY GOOD COMPETITORS THAT ARE WEAK DISPERSERS -OUQUET AND
,OREAU 4HE EXPERIMENTAL RESULT OF DECLINING INVASION SUCCESS WITH INCREAS
ING LOCAL DIVERSITY HOWEVER IS INCONSISTENT WITH NEUTRAL AND NON
INTERACTIVE
PATCH
DYNAMICS MODELS BUT CONSISTENT WITH ALL THE OTHER METACOMMUNITY MODELS
4HE DISCREPANCY BETWEEN PATTERNS AT LOCAL VS REGIONAL SCALES COULD REmECT THE RELA
TIVE IMPORTANCE OF DIFFERENT MECHANISMS OPERATING AT VARYING SPATIAL SCALES 3PECIES
RICHNESS MIGHT BE MORE IMPORTANT IN DETERMINING INVASION SUCCESS AT LOCAL SCALES
WHEREAS OTHER ECOLOGICAL FACTORS SUCH AS PROPAGULE PRESSURE OR RESOURCE AVAILABIL
ITY MIGHT GOVERN PATTERNS OF COMMUNITY INVASIBILITY AND SATURATION AT COMMU
NITY
WIDE SCALES ,EVINE +ENNEDY ET AL "YERS AND .OONBURG
/NE EXPLANATION FOR THIS PATTERN IS THAT CHANGES IN THE NUMBER OF AVAILABLE RESOURCES
AS WELL AS DISTURBANCE OR PROPAGULE PRESSURE CAN CAUSE INVASION SUCCESS TO BECOME
POSITIVELY CORRELATED WITH DIVERSITY AT LARGE SCALES BY DECREASING MEAN INTERACTION
STRENGTH AMONG COMPETING SPECIES "YERS AND .OONBURG
0ATTERNS IN THE BIOLOGICAL INVASIONS LITERATURE SUGGEST THAT NON
INTERACTIVE
PATCH
DYNAMICS AND NEUTRAL MODELS SUCCESSFULLY PREDICT THE DYNAMICS OF INVA
SIONS AND EXTINCTIONS AT LARGE SPATIAL SCALES WHILE FAILING TO INCORPORATE THE
MECHANISMS RESPONSIBLE FOR INVASION RESISTANCE WITH INCREASING DIVERSITY AT SMALL
SPATIAL SCALES )NTERACTIVE PATCH
DYNAMICS AND MASS
EFFECTS MODELS ADEQUATELY
DESCRIBE PATTERNS AT SMALL AND LARGER SPATIAL SCALES (OWEVER THE PREDICTION OF
DECLINING DIVERSITY AT THE HIGHEST LEVELS OF DISPERSAL WAS NOT BORN OUT 4HE INVA
SIBILITY AND UNDERSATURATION OF COMMUNITIES IS NOT PREDICTED BY SPECIES
SORTING
MODELS WITH GLOBAL DISPERSAL OR BY INTERACTIVE PATCH
DYNAMICS AND MASS
EFFECTS
MODELS WITH EXTREMELY HIGH RATES OF DISPERSAL (OWEVER THE DECLINE IN INVASION
SUCCESS AT HIGHER DIVERSITY AND THE LARGE EFFECTS OF DISTURBANCE IN WEAKENING BIOTIC
RESISTANCE TO INVASION FOUND IN THE EXPERIMENTAL LITERATURE SUPPORTS THE SPECIES
SORTING PERSPECTIVE
#/.#,53)/.
2ECONCILING SMALL
SCALE MECHANISTIC STUDIES OF THE FACTORS THAT ALLOW OR PRE
VENT INVASION WITH BROAD
SCALE PATTERNS OF DIVERSITY GAIN AND LOSS REMAINS ONE
OF THE MOST PRESSING CHALLENGES IN COMMUNITY ECOLOGY AND INVASION BIOLOGY
4HE UNDER
SATURATION OF ECOLOGICAL COMMUNITIES EVIDENT FROM MACRO
SCALE PAT
TERNS OF INVASION AND EXTINCTION IN FAUNAL EXCHANGES AND CONTEMPORARY INVA
SIONS HAS BEEN INTERPRETED AS INDICATING THAT LOCAL INTERACTIONS EXERT RELATIVELY
WEAK INmUENCE OVER COMMUNITY STRUCTURE 3TOHLGREN ET AL 3AX AND 'AINES
(OWEVER EVEN STRONG LOCAL COMPETITIVE INTERACTIONS CAN PLAY OUT VERY
%VIDENCE FOR INVASIBILITY AND SATURATION
SLOWLY OVER BROAD HETEROGENEOUS LANDSCAPES LEADING TO PATTERNS OF INVASIBIL
ITY AND UNSATURATED COMMUNITIES AT REGIONAL SCALES &URTHERMORE EXPERIMENTAL
INTRODUCTIONS PROVIDE EVIDENCE THAT LOCAL ECOLOGICAL INTERACTIONS PLAY A MAJOR ROLE
IN DETERMINING THE SUCCESS OR FAILURE OF INVASIONS 4HE APPARENT UNSATURATION OF
COMMUNITIES DOES NOT THEREFORE LEND SUPPORT TO A STRICTLY DISPERSAL
LIMITED NON
INTERACTIVE VIEW OF ECOLOGICAL COMMUNITIES
$ISPERSAL AND LOCAL INTERACTIONS HAVE LONG BEEN TREATED AS MUTUALLY EXCLUSIVE
ALTERNATIVE EXPLANATIONS FOR COMMUNITY STRUCTURE (OWEVER RECENT THEORETICAL
WORK HAS REVEALED A NUMBER OF WAYS IN WHICH DISPERSAL AND LOCAL INTERACTIONS
CAN JOINTLY REGULATE COMMUNITY STRUCTURE 3HURIN AND !LLEN "YERS AND
.OONBURG -OUQUET AND ,OREAU 4HE MOVEMENT OF INDIVIDUALS
AMONG PATCHES GIVES COHERENCE TO LOCAL COMPETITIVE INTERACTIONS WITHIN A REGION
AND THEREFORE LOCAL COMMUNITY DYNAMICS MAY PLAY THEMSELVES OUT AT REGIONAL
SPATIAL SCALES 2ICKLEFS )NDEED THE METACOMMUNITY APPROACH SUGGESTS
THAT PROCESSES WHICH HAVE TRADITIONALLY BEEN CONSIDERED LOCAL AND REGIONAL IN
NATURE NOW NEED TO BE SEEN AS SPATIALLY LINKED %XTENDING LOCAL COMPETITIVE
INTERACTIONS TO A BROAD REGION SLOWS THE TIME SCALE OF COMPETITIVE EXCLUSION TO
THAT OF SPECIES PRODUCTION 2ICKLEFS WHICH HAS IMPORTANT IMPLICATIONS FOR
INVASIONS BIOLOGISTS 'LOBAL EXTINCTIONS MAY LAG BEHIND THE WAVE OF INVASIONS SO
THAT MANY PENDING LOCAL EXTIRPATION EVENTS HAVE NOT YET OCCURRED
&INALLY THE LITERATURE ON INVASIONS AND EXTINCTIONS REVEALS THAT THE APPARENT
PATTERNS ARE STRONGLY SCALE
DEPENDENT )NVASION MAY INCREASE BOTH LOCAL EG
4ABLE AND REGIONAL DIVERSITY 3AX AND 'AINES HOWEVER IT IS APPAR
ENT THAT BETA DIVERSITY OR REGIONAL DISTINCTIVENESS IN COMPOSITION IS DECLINING
4HE HOMOGENIZATION OF THE BIOSPHERE IS OCCURRING AT EVER INCREASING RATES
-C+INNEY AND ,OCKWOOD 2AHEL 3YNERGISTIC INTERACTIONS
AMONG INVADERS LEADING TO BIOLOGICAL MELTDOWNS 3IMBERLOFF AND 6ON (OLLE
2ICCIARDI AND AMONG OTHER HUMAN
MEDIATED THREATS TO GLOBAL BIODIVERSITY
3ALA ET AL WILL CERTAINLY INCREASE THE VULNERABILITY OF NATURAL COMMUNI
TIES TO THE INTRODUCTION OF EXOTIC SPECIES 4HE CONTINUOUS AND EVER INCREASING RAIN
OF PROPAGULES OF EXOTIC SPECIES -AC)SAAC ET AL 'RIGOROVICH ET AL
CONTINUES TO EXCEED THE ABILITY OF NATURAL COMMUNITIES TO REPEL THEM $ESPITE
DOCUMENTED EVIDENCE THAT LOCAL COMMUNITIES RESIST INVASIONS SECTION MACRO
SCALE PATTERNS OF BIOTIC INTERCHANGES AND BIOLOGICAL INVASIONS SECTIONS AND
SUGGEST THAT IT IS UNLIKELY THAT LOCAL COMMUNITIES CAN BE MANAGED IN WAYS THAT
WILL PREVENT ALL INVASIONS &URTHERMORE INCREASED FREQUENCY OF HUMAN
MEDIATED
TRANSPORT OF EXOTIC SPECIES COUPLED WITH THE INCREASED VULNERABILITY OF ECOSYSTEMS
TO INVASION WILL CERTAINLY INCREASE THE PROBABILITY OF INTRODUCING EXOTIC SPECIES
WITH NOVEL ECOLOGICAL FUNCTIONS INTO NAVE ECOSYSTEMS 3UCH INVASION EVENTS
CLEARLY HAVE DISASTROUS REPERCUSSIONS ON THE SPECIES RICHNESS AS WELL AS THE FUNC
TIONING OF LOCAL ECOSYSTEMS +AUFMAN .ALEPA AND 3CHLOESER &RITTS
AND 2ODDA 6ANDERPLOEG ET AL )N ORDER TO CURTAIL THE INTRODUCTION OF
EXOTIC SPECIES AND PREVENT THEIR SOMETIMES DISASTROUS CONSEQUENCES OUR RESULTS
SUGGEST THAT MANAGERS AND POLICY MAKERS ALIKE NEED TO FOCUS EFFORTS ON MANAGING
3! 3MITH AND *" 3HURIN
DISPERSAL VECTORS AND PREVENTING THE ESTABLISHMENT OF EXOTIC SPECIES RATHER THAN
RELYING ON INTACT NATIVE COMMUNITIES TO REPEL INVADERS
!#+./7,%$'%-%.43
4HE AUTHORS WOULD LIKE TO THANK 2OD $OCKING *OANNA -ACKENZIE 2 #RAIG
-AC,EAN AND -ICHELLE 0ALMER FOR HELPFUL COMMENTS ON A PREVIOUS DRAFT OF THIS
MANUSCRIPT 3!3 WAS SUPPORTED BY AN .3%2# 0'3 " !N .3%2# GRANT TO *"3
PROVIDED ADDITIONAL FUNDING
2%&%2%.#%3
"AKKER 2 4 4HE DEER mEES THE WOLF PURSUES INCONGUENCIES IN PREDATOR PREY EVO
LUTION 0AGES IN $ &UTUYMA AND - 3LATKIN EDITORS #OEVOLUTION 3INAUER
3UNDERLAND
"ARRY * # - % -ORGAN ! * 7INKLER , * &LYNN % ( ,INDAY , , *ACOBS AND
$ 0ILBEAM &AUNAL INTERCHANGE AND -IOCENE TERRESTRIAL VERTEBRATES OF SOUTHERN
!SIA 0ALEOBIOLOGY
"ELL ' .EUTRAL -ACROECOLOGY 3CIENCE
"EN 4UVIA ! 4HE IMPACT OF THE ,ESSEPSIAN 3UEZ #ANAL &ISH MIGRATION ON THE EAST
ERN -EDITERRANEAN ECOSYSTEM 0AGES IN - -ORAITOU !POSTOLOPOULOU AND
6 +IORTSIS EDITORS -EDITERRANEAN MARINE ECOSYSTEMS 0LENUM 0RESS .EW 9ORK
"ERMINGHAM % AND ! 0 -ARTIN #OMPARATIVE MT$.! PHYLOGEOGRAPHY OF NEO
TROPICAL FRESHWATER lSHES TESTING SHARED HISTORY TO INFER THE EVOLUTIONARY LANDSCAPE OF
LOWER #ENTRAL !MERICA -OLECULAR %COLOGY
"ROWN * ( 4 * 6ALONE AND # ' #URTIN 2EORGANIZATION OF AN ARID ECOSYSTEM
IN RESPONSE TO RECENT CLIMATE CHANGE 0ROCEEDINGS OF THE .ATIONAL !CADEMY OF 3CIENCE
"ROWN * ( 3 + -ORGAN %RNEST * - 0ARODY AND * 0 (ASKELL 2EGULATION OF
DIVERSITY MAINTENANCE OF SPECIES RICHNESS IN CHANGING ENVIRONMENTS /ECOLOGIA
"URKE - * 7 AND * 0 'RIME !N EXPERIMENTAL STUDY OF PLANT COMMUNITY INVASI
BILITY %COLOGY
"YERS * % AND % ' .OONBURG 3CALE DEPENDENT EFFECTS OF BIOTIC RESISTANCE TO BIOLOGI
CAL INVASIONS %COLOGY
#ASE 4 * 'LOBAL PATTERNS IN THE ESTABLISHMENT AND DISTRIBUTION OF EXOTIC BIRDS
"IOLOGICAL #ONSERVATION
#ASWELL ( AND * % #OHEN ,OCAL AND REGIONAL REGULATION OF SPECIES AREA RELATIONS
A PATCH OCCUPANCY MODEL 0AGES IN 2 % 2ICKLEFS AND $ 3CHLUTER EDITORS
3PECIES DIVERSITY IN ECOLOGICAL COMMUNITIES (ISTORICAL AND GEOGRAPHICAL PERSPECTIVES
5NIVERSITY OF #HICAGO 0RESS #HICAGO
%VIDENCE FOR INVASIBILITY AND SATURATION
#HASE * - AND - ! ,EIBOLD %COLOGICAL NICHES LINKING CLASSICAL AND CONTEMPORARY
APPROACHES 5NIVERSITY OF #HICAGO 0RESS #HICAGO
#HASE * 0 !MARASEKARE + #OTTENIE ! 'ONZALEZ 2 (OLT - (OLYOAK - (OOPES -
,EI BOLD - ,OREAU . -OUQUET * " 3HURIN AND $ 4ILMAN #OMPETING THEO
RIES FOR COMPETITIVE METACOMMUNITIES 0AGES
IN - (OLYOAK 2 (OLT AND
- ,EIBOLD EDITORS -ETACOMMUNITIES )N 0RESS
#OHEN ! . AND * 4 #ARLTON !CCELERATING INVASION RATE IN A HIGHLY INVADED
ESTUARY 3CIENCE
#ORNELL ( 6 3PECIES ASSEMBLAGES OF #YNIPID 'ALL 7ASPS ARE NOT SATURATED
!MERICAN .ATURALIST
#ORNELL ( 6 5NSATURATED PATTERNS IN SPECIES ASSEMBLAGES THE ROLE OF REGIONAL
PROCESSES IN SETTLING LOCAL SPECIES RICHNESS 0AGES
IN 2 % 2ICKLEFS AND $
3CHLUTER EDITORS 3PECIES DIVERSITY IN ECOLOGICAL COMMUNITIES HISTORICAL AND GEOGRAPHICAL
PERSPECTIVES 5NIVERSITY OF #HICAGO 0RESS #HICAGO
#ORNELL ( 6 5NSATURATION AND REGIONAL INmUENCES ON SPECIES RICHNESS IN ECOLOGICAL
COMMUNITIES ! REVIEW OF THE EVIDENCE %COSCIENCE
#ORNELL ( 6 AND * ( ,AWTON 3PECIES INTERACTIONS LOCAL AND REGIONAL PROCESSES
AND LIMITS TO THE RICHNESS OF ECOLOGICAL COMMUNITIES A THEORETICAL PERSPECTIVE *OURNAL
OF !NIMAL %COLOGY
$AVIS - ! "IOTIC GLOBALIZATION $OES COMPETITION FROM INTRODUCED SPECIES THREATEN
BIODIVERSITY "IOSCIENCE
$IAMOND * - AND # 2 6EITCH %XTINCTIONS AND INTRODUCTIONS IN THE .EW :EALAND
AVIFAUNA CAUSE AND EFFECT 3CIENCE
$UNCAN * 2 AND * , ,OCKWOOD 3PATIAL HOMOGENIZATION OF THE AQUATIC FAUNA OF
4ENNESSEE EXTINCTION AND INVASION FOLLOWING LAND USE CHANGE AND HABITAT ALTERATION
0AGES
IN * , ,OCKWOOD AND - , -C+INNEY EDITORS "IOTIC HOMOGENIZATION
+LUWER !CADEMIC 0RESS0LENUM 0UBLISHERS .EW 9ORK
$UNCAN 2 0 4 - "LACKBURN AND $ 3OL 4HE ECOLOGY OF BIRD INTRODUCTIONS
!NNUAL 2EVIEW OF %COLOGY AND 3YSTEMATICS
&ARGIONE * # 3 "ROWN AND $ 4ILMAN #OMMUNITY ASSEMBLY AND INVASION
!N EXPERIMENTAL TEST OF NEUTRAL VERSUS NICHE PROCESSES 0ROCEEDINGS OF THE .ATIONAL
!CADEMY OF 3CIENCES OF THE 5NITED 3TATES OF !MERICA
&OSTER " , #ONSTRAINTS ON COLONIZATION AND SPECIES RICHNESS ALONG A GRASSLAND
PRODUCTIVITY GRADIENT THE ROLE OF PROPAGULE AVAILABILITY %COLOGY ,ETTERS
&OSTER " , 6 ( 3MITH 4 , $ICKSON AND 4 (ILDEBRAND )NVASIBILITY AND COM
POSITIONAL STABILITY IN A GRASSLAND COMMUNITY RELATIONSHIPS TO DIVERSITY AND EXTRINSIC
FACTORS /IKOS
&OSTER " , AND $ 4ILMAN 3EED LIMITATION AND THE REGULATION OF COMMUNITY STRUC
TURE IN OAK SAVANNA GRASSLAND *OURNAL OF %COLOGY
&OX * 7 * -C'RADY
3TEED AND / , 0ETCHEY 4ESTING FOR LOCAL SPECIES SATURATION
WITH NONINDEPENDENT REGIONAL SPECIES POOLS %COLOGY ,ETTERS
&RITTS 4 ( AND ' ( 2ODDA 4HE ROLE OF INTRODUCED SPECIES IN THE DEGRADATION OF
ISLAND ECOSYSTEMS ! CASE HISTORY OF 'UAM !NNUAL 2EVIEW OF %COLOGY AND 3YSTEMATICS
3! 3MITH AND *" 3HURIN
&LYNN , * 2 ( 4EDFORD AND 1 :HANXIANG %NRICHMENT AND STABILITY IN THE
0LIOCENE MAMMALIAN FAUNA OF .ORTH #HINA 0ALEOBIOLOGY
&UKAMI 4 #OMMUNITY ASSEMBLY ALONG A SPECIES POOL GRADIENT IMPLICATIONS FOR
MULTIPLE
SCALE PATTERNS OF DIVERSITY 0OPULATION %COLOGY
'ASTON + * 'LOBAL PATTERNS IN BIODIVERSITY .ATURE
'ASTON + * %COLOGY THE HOW AND WHY OF BIODIVERSITY .ATURE
'IDO + " AND * ( "ROWN )NVASION OF .ORTH !MERICAN DRAINAGES BY ALIEN lSH SPE
CIES &RESHWATER "IOLOGY
'RIGOROVICH ) ! 2 ) #OLAUTTI % , -ILLS + (OLECK ! ' "ALLERT AND ( * -AC)SAAC
"ALLAST
MEDIATED ANIMAL INTRODUCTIONS IN THE ,AURENTIAN 'REAT ,AKES RETRO
SPECTIVE AND PROSPECTIVE ANALYSES #ANADIAN *OURNAL OF &ISHERIES AND !QUATIC 3CIENCE
'UREVITCH * AND $ + 0ADILLA !RE INVASIVE SPECIES A MAJOR CAUSE OF EXTINCTIONS
4RENDS IN %COLOGY AND %VOLUTION
(ASTINGS ! $ISTURBANCE COEXISTENCE HISTORY AND COMPETITION FOR SPACE 4HEORETICAL
0OPULATION "IOLOGY
(OBBS 2 * AND ( ! -OONEY "ROADENING THE EXTINCTION DEBATE POPULATION
DELETIONS AND ADDITIONS IN #ALIFORNIA AND 7ESTERN !USTRALIA #ONSERVATION "IOLOGY
(UBBELL 3 0 4HE UNIlED THEORY OF BIODIVERSITY AND BIOGEOGRAPHY 0RINCETON
5NIVERSITY 0RESS 0RINCETON
+AUFMAN , #ATASTROPHIC CHANGE IN SPECIES
RICH FRESHWATER ECOSYSTEMS THE LESSONS
OF ,AKE 6ICTORIA "IO3CIENCE
+EIT 4 ( AND 0 ! -ARQUET 4HE INTRODUCED (AWAIIAN AVIFAUNA RECONSIDERED
%VIDENCE FOR SELF
ORGANIZATION AND CRITICALITY *OURNAL OF 4HEORETICAL "IOLOGY
+ENNEDY 4 ! 3 .AEEM + - (OWE * - ( +NOPS $ 4ILMAN AND 0 2EICH
"IODIVERSITY AS A BARRIER TO ECOLOGICAL INVASION .ATURE
,EIBOLD - ! - (OLYOAK . -OUQUET 0 !MARASEKARE * - #HASE - & (OOPES 2 $
(OLT * " 3HURIN 2 ,AW $ 4ILMAN - ,OREAU AND ! 'ONZALEZ 4HE METACOM
MUNITY CONCEPT A FRAMEWORK FOR MULTI
SCALE COMMUNITY ECOLOGY %COLOGY ,ETTERS
,ESSA % 0 " 6 6ALKENBURGH AND 2 ! &ARINA 4ESTING HYPOTHESES OF DIFFEREN
TIAL MAMMALIAN EXTINCTIONS SUBSEQUENT TO THE 'REAT !MERICAN "IOTIC )NTERCHANGE
0ALAEOGEOGRAPHY 0ALAEOCLIMATOLOGY 0ALAEOECOLOGY
,EVINE * - 3PECIES DIVERSITY AND BIOLOGICAL INVASIONS 2ELATING LOCAL PROCESS TO
COMMUNITY PATTERN 3CIENCE
,EVINE * - ,OCAL INTERACTIONS DISPERSAL AND NATIVE AND EXOTIC PLANT DIVERSITY
ALONG A #ALIFORNIA STREAM /IKOS
,EVINS 2 AND $ #ULVER 2EGIONAL COEXISTENCE OF SPECIES AND COMPETITION BETWEEN
RARE SPECIES 0ROCEEDINGS OF THE .ATIONAL !CADEMY OF 3CIENCE
,INBERG $ 2 -ARINE BIOTIC INTERCHANGE BETWEEN THE .ORTHERN AND 3OUTHERN HEMI
SPHERES 0ALEOBIOLOGY
,ODGE $ - "IOLOGICAL INVASIONS ,ESSONS FOR ECOLOGY 4RENDS IN %COLOGY AND
%VOLUTION
%VIDENCE FOR INVASIBILITY AND SATURATION
,OSOS * " AND $ 3CHLUTER !NALYSIS OF AN EVOLUTIONARY SPECIES AREA RELATIONSHIP
.ATURE
-AC)SAAC ( * 4 # 4HOMAS # 2OBBINS AND - ! ,EWIS -ODELING SHIPS BALLAST
WATER AS INVASION THREATS TO THE 'REAT ,AKES #ANADIAN *OURNAL OF &ISHERIES AND !QUATIC
3CIENCE
-ARCHETTI - 0 4 ,IGHT * &ELICIANO 4 !RMSTRONG : (OGAN * 6IERS AND 0 " -OYLE
(OMOGENIZATION OF #ALIFORNIAS lSH FAUNA THROUGH ABIOTIC CHANGE 0AGES
IN * , ,OCKWOOD AND - , -C+INNEY EDITORS "IOTIC (OMOGENIZATION
+LUWER !CADEMIC 0RESS0LENUM 0UBLISHERS .EW 9ORK
-ARSHALL , ' 4HE 'REAT !MERICAN "IOTIC )NTERCHANGE AN INVASION INmUENCED CRISIS
FOR 3OUTH !MERICAN MAMMALS 0AGES IN - ( .ITECKI EDITOR "IOTIC CRISES IN
ECOLOGICAL AND EVOLUTIONARY TIME !CADEMIC 0RESS .EW 9ORK
-ARSHALL , ' AND 2 , #IFELLI !NALYSIS OF CHANGING DIVERSITY PATTERNS IN #ENOZOIC
LAND MAMMAL AGE FAUNA 3OUTH !MERICA 0ALAEOVERTEBRATA
-C+INNEY - , AND * , ,OCKWOOD "IOTIC HOMOGENIZATION A FEW WINNERS REPLACING
MANY LOSERS IN THE NEXT MASS EXTINCTION 4RENDS IN %COLOGY %VOLUTION
-ORA # 0 - #HITTARO 0 & 3ALE * 0 +RITZER AND 3 ! ,UDSIN 0ATTERNS AND PRO
CESSES IN REEF lSH DIVERSITY .ATURE
-OUQUET . AND - ,OREAU #OMMUNITY PATTERNS IN SOURCE SINK METACOMMUNITIES
!MERICAN .ATURALIST
-OUQUET . 0 ,EADLEY * -ERIGUET AND - ,OREAU )MMIGRATION AND LOCAL COM
PETITION IN HERBACEOUS PLANT COMMUNITIES A THREE YEAR SEED SOWING EXPERIMENT /IKOS
-OYLE 0 " AND 4 ,IGHT "IOLOGICAL INVASIONS OF FRESH WATER EMPIRICAL RULES AND
ASSEMBLY THEORY "IOLOGICAL #ONSERVATION
.ALEPA & 4 AND $ 7 3CHLOESSER :EBRA MUSSELS "IOLOGY IMPACTS AND CONTROL
,EWIS 0UBLISHERS "OCA 2ATON
0ERDICES ! % "ERMINGHAM ! -ONTILLA AND ) $OADRIO %VOLUTIONARY HISTORY OF THE
GENUS 2HAMDIA 4ELEOSTEI 0IMELODIDAE IN #ENTRAL !MERICA -OLECULAR 0HYLOGENETICS
AND %VOLUTION
0OR & $ ,ESSEPSIAN -IGRATION 4HE INmUX OF THE 2ED 3EA "IOTA INTO THE
-EDITERRANEAN BY WAY OF THE 3UEZ #ANAL 6ERLAG "ERLIN
0OR & $ 4HE LEGACY OF 4ETHYS !N AQUATIC BIOGEOGRAPHY OF THE ,EVANT +LUWER
!CADEMIC 0RESS ,ONDON
2AHEL & * (OMOGENIZATION OF lSH FAUNAS ACROSS THE 5NITED 3TATES 3CIENCE
2AHEL & * (OMOGENIZATION OF FRESHWATER FAUNAS !NNUAL 2EVIEW OF %COLOGY AND
3YSTEMATICS
2ICCIARDI ! &ACILITATIVE INTERACTIONS AMONG AQUATIC INVADERS IS AN hINVASIONAL
MELTDOWNv OCCURRING IN THE 'REAT ,AKES #ANADIAN *OURNAL OF &ISHERIES AND !QUATIC
3CIENCE
2ICCIARDI ! !SSESSING SPECIES INVASIONS AS A CAUSE OF EXTINCTION 4RENDS IN %COLOGY
AND %VOLUTION
3! 3MITH AND *" 3HURIN
2ICKLEFS 2 % #OMMUNITY DIVERSITY RELATIVE ROLES OF LOCAL AND REGIONAL PROCESSES
3CIENCE
2ICKLEFS 2 % ! COMPREHENSIVE FRAMEWORK FOR GLOBAL PATTERNS IN BIODIVERSITY
%COLOGY ,ETTERS
2ICKLEFS 2 % AND % "ERMINGHAM .ONEQUILIBIRUM $IVERSITY $YNAMICS OF THE ,ESSER
!NTILLEAN !VIFAUNA 3CIENCE
3ALA / % & 3 #HAPIN * * !RMESTO % "ERLOW * "LOOMlELD 2 $IRZO % (UMBER
3ANWALD
, & (UENNEKE 2 " *ACKSON ! +INZIG 2 ,EEMANS $ - ,ODGE ( ! -OONEY
- /ESTERHELD . ,E2OY 0OFF - 4 3YKES " ( 7ALKER - 7ALKER AND $ ( 7ALL
'LOBAL BIODIVERSITY SCENARIOS FOR THE YEAR 3CIENCE
3AX $ & 3 $ 'AINES AND * ( "ROWN 3PECIES INVASIONS EXCEED EXTINCTIONS ON
ISLANDS WORLDWIDE ! COMPARATIVE STUDY OF PLANTS AND BIRDS !MERICAN .ATURALIST
3AX $ & AND 3 $ 'AINES 3PECIES DIVERSITY FROM GLOBAL DECREASES TO LOCAL INCREASES
4RENDS IN %COLOGY AND %VOLUTION
3EABLOOM % 7 7 3 (ARPOLE / * 2EICHMAN AND $ 4ILMAN )NVASION COM
PETITIVE DOMINANCE AND RESOURCE USE BY EXOTIC AND NATIVE #ALIFORNIA GRASSLAND SPECIES
0ROCEEDINGS OF THE .ATIONAL !CADEMY OF 3CIENCES OF THE 5NITED 3TATES OF !MERICA
3HMIDA ! AND 3 %LLNER #OEXISTENCE OF PLANT SPECIES WITH SIMILAR NICHES 6EGETATIO
3HURIN * " $ISPERSAL LIMITATION INVASION RESISTANCE AND THE STRUCTURE OF POND
ZOOPLANKTON COMMUNITIES %COLOGY
3HURIN * " AND % ' !LLEN %FFECTS OF COMPETITION PREDATION AND DISPERSAL ON SPE
CIES RICHNESS AT LOCAL AND REGIONAL SCALES !MERICAN .ATURALIST
3HURIN * " AND $ 3 3RIVASTAVA .EW PERSPECTIVES ON LOCAL AND REGIONAL DIVERSITY
BEYOND SATURATION 0AGES
IN - (OLYOAK - ,EIBOLD AND 2 (OLT EDITORS
-ETACOMMUNITIES SPATIAL DYNAMICS AND ECOLOGICAL COMMUNITIES 5NIVERSITY OF #HICAGO
0RESS #HICAGO
3IMBERLOFF $ %XTINCTION SURVIVAL AND EFFECTS OF BIRDS INTRODUCED
TO THE -ASCA
RENES !CTA /ECOLOGIA
3IMBERLOFF $ AND " 6 (OLLE 0OSITIVE INTERACTIONS OF NONINDIGENOUS SPECIES INVA
SIONAL MELTDOWN "IOLOGICAL )NVASIONS
3IMPSON ' ' (ISTORY OF THE FAUNA OF ,ATIN !MERICA !MERICAN *OURNAL OF 3CIENCE
3IMPSON ' ' 3PLENDID ISOLATION THE CURIOUS HISTORY OF 3OUTH !MERICAN MAMMALS
9ALE 5NIVERSITY 0RESS .EW (AVEN
3MALLWOOD + 3 3ITE INVASIBILITY BY EXOTIC BIRDS AND MAMMALS "IOLOGICAL
#ONSERVATION
3MITH 3 ! AND % "ERMINGHAM 4HE BIOGEOGRAPHY OF LOWER -ESOAMERICAN FRESHWATER
lSHES 3UBMITTED TO THE *OURNAL OF "IOGEOGRAPHY
3MITH 3 ! ' "ELL AND % "ERMINGHAM #ROSS
#ORDILLERA EXCHANGE MEDIATED BY
THE 0ANAMA #ANAL INCREASED THE SPECIES RICHNESS OF LOCAL FRESHWATER lSH ASSEMBLAGES
0ROCEEDINGS OF THE 2OYAL 3OCIETY OF ,ONDON "
%VIDENCE FOR INVASIBILITY AND SATURATION
3TOHLGREN 4 * $ "INKLEY ' 7 #HONG - ! +ALKHAN , $ 3CHELL + ! "ULL 9 /TSUKI
' .EWMAN - "ASHKIN AND 9 3ON %XOTIC PLANT SPECIES INVADE HOTSPOTS OF
NATIVE PLANT DIVERSITY %COLOGICAL -ONOGRAPHS
4ERBORGH * 7 AND * &AABORG 3ATURATION AND BIRD COMMUNITIES IN THE 7EST )NDIES
!MERICAN .ATURALIST
4ILMAN $ #OMPETITION AND BIODIVERSITY IN SPATIALLY STRUCTURED HABITATS %COLOGY
4ILMAN $ #OMMUNITY INVASIBILITY RECRUITMENT LIMITATION AND GRASSLAND BIODIVER
SITY %COLOGY
4ILMAN $ .ICHE TRADEOFFS NEUTRALITY AND COMMUNITY STRUCTURE ! STOCHASTIC
THEORY OF RESOURCE COMPETITION INVASION AND COMMUNITY ASSEMBLY 0ROCEEDINGS OF THE
.ATIONAL !CADEMY OF 3CIENCES 53!
4URNBULL , ! - * #RAWLEY AND - 2EES !RE PLANT POPULATIONS SEED LIMITED
! REVIEW OF SEED SOWING EXPERIMENTS /IKOS 6ANDERPLOEG ( !
4 & .ALEPA $ * *UDE % , -ILLS + 4 (OLECK * 2 ,IEBIG ) !
'RIGOROVICH AND ( /JAVEER $ISPERSAL AND EMERGING ECOLOGICAL IMPACTS OF 0ONTO
#ASPIAN SPECIES IN THE ,AURENTIAN 'REAT ,AKES #ANADIAN *OURNAL OF &ISHERIES AND
!QUATIC 3CIENCES
6ERMEIJ ' * A )NVASION AND EXTINCTION 4HE LAST THREE MILLION YEARS OF .ORTH 3EAN
PELECYPOD HISTORY #ONSERVATION "IOLOGY
6ERMEIJ ' * B 'EOGRAPHICAL RESTRICTION AS A GUIDE TO THE CAUSES OF EXTINCTION
THE CASE OF THE COLD NORTHERN OCEANS DURING THE .EOGENE 0ALEOBIOLOGY
6ERMEIJ ' * A 7HEN BIOTAS MEET 5NDERSTANDING BIOTIC INTERCHANGE 3CIENCE
6ERMEIJ ' * B !NATOMY OF AN INVASION THE TRANS !RCTIC INTERCHANGE 0ALEOBIOLOGY
6ITOUSEK 0 - # - $!NTONIO , $ ,OOPE - 2EJMANEK AND 2 7ESTBROOKS
)NTRODUCED SPECIES A SIGNIlCANT COMPONENT OF HUMAN CAUSED GLOBAL CHANGE .EW
:EALAND *OURNAL OF %COLOGY
7ALLACE ! 2 4HE GEOGRAPHICAL DISTRIBUTION OF ANIMALS (AFNER .EW 9ORK
7EBB 3 $ AND , ' -ARSHALL (ISTORICAL BIOGEOGRAPHY OF RECENT 3OUTH !MERICAN
LAND MAMMALS 0AGES IN - ! -ARES AND ( ( 'ENOWAYS EDITORS -AMMALIAN
BIOLOGY IN 3OUTH !MERICA 4HE 5NIVERSITY OF 0ITTSBURGH 0ITTSBURGH
7EBB 3 $ ,ATE #ENEZOIC DISPERSALS BETWEEN THE !MERICAS 0AGES IN
& ' 3TEHLI AND 3 $ 7EBB EDITORS 4HE 'REAT !MERICAN "IOTIC )NTERCHANGE 0LENUM
.EW 9ORK
:OBEL - - /TSUS * ,IIRA - -OORA AND 4 -OLS )S SMALL SCALE SPECIES RICHNESS
LIMITED BY SEED AVAILABILITY OR MICROSITE AVAILABILITY %COLOGY
#HAPTER TWENTY
4HE BIOGEOGRAPHY OF
NATURALIZED SPECIES AND
THE SPECIES
AREA RELATIONSHIP
RECIPROCAL INSIGHTS TO
BIOGEOGRAPHY AND INVASION
BIOLOGY
$ & 3AX AND 3 $ 'AINES
).42/$5#4)/.
h(E WHO ADMITS THE DOCTRINE OF THE CREATION OF EACH SEPARATE SPECIES WILL HAVE TO ADMIT
THAT A SUFlCIENT NUMBER OF THE BEST ADAPTED PLANTS AND ANIMALS HAVE NOT BEEN CREATED ON
OCEANIC ISLANDS FOR MAN HAS UNINTENTIONALLY STOCKED THEM FROM VARIOUS SOURCES FAR MORE
FULLY AND PERFECTLY THAN HAS NATUREv
#HARLES $ARWIN
$ARWIN WAS NOT ARGUING THAT SPECIES INVASION WAS A PREFERABLE CONDITION
)NDEED HE ARGUED THAT INTRODUCED SPECIES WERE LIKELY TO DRIVE MANY ISLAND ENDEMICS
-7 #ADOTTE ET AL EDS #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY
3PRINGER 0RINTED IN THE .ETHERLANDS
$& 3AX AND 3$ 'AINES
ICS TO EXTINCTION )NSTEAD HE WAS DRAWING ON PATTERNS OF SPECIES INTRODUCTION AND
NATURALIZATION TO MAKE INFERENCES ABOUT THE CAPACITY OF ISLANDS TO HOLD MANY
MORE SPECIES THAN HAD NATURALLY COLONIZED OR EVOLVED IN SITU $ARWIN WAS CLEARLY
A BROAD THINKER AND IN HIS QUEST TO UNDERSTAND THE NATURAL WORLD HE FREQUENTLY
DREW UPON FORMS OF EVIDENCE THAT MIGHT BE VIEWED AS hARTIlCIALv OR hUNNATURALv
&OR EXAMPLE HE DREW HEAVILY UPON EVIDENCE FROM ARTIlCIAL BREEDING TO BETTER
UNDERSTAND THE PROCESS OF NATURAL SELECTION 4HEREFORE IT IS NOT SURPRISING THAT
HE DREW UPON EVIDENCE FROM NATURALIZED SPECIES IE INTRODUCED SPECIES THAT FORM
SELF
SUSTAINING POPULATIONS TO BETTER UNDERSTAND BIOGEOGRAPHIC PRINCIPLES OF THE
NATURAL WORLD "ESIDES HIS COMMENTS QUOTED ABOVE HE ALSO USED NATURALIZED SPE
CIES NAMELY THEIR KNOWN RATES OF SPREAD TO INFER THAT NATIVE SPECIES WOULD HAVE
HAD AMPLE TIME TO DRAMATICALLY ALTER THEIR GEOGRAPHIC RANGES DURING THE LAST GLA
CIAL PERIOD 4HESE TWO OBSERVATIONS OF $ARWINS ARE BUT A FEW OF THE MANY POSSIBLE
INSIGHTS THAT STUDYING NATURALIZED SPECIES CAN PROVIDE FOR THE lELDS OF BIOGEOGRA
PHY AND ECOLOGY
)N THIS CHAPTER WE WILL ENDEAVOR TO ACCOMPLISH SEVERAL GOALS &IRST WE REVIEW
WORK DONE TO DATE POST $ARWIN THAT EXPLORES INSIGHTS THAT STUDYING NATURALIZED
SPECIES CAN PROVIDE TO UNDERSTANDING NATURAL PATTERNS AND PROCESSES 3ECOND IN
KEEPING WITH THE THEME OF THIS BOOK WE WILL ATTEMPT TO USE THESE SAME PATTERNS TO
BETTER UNDERSTAND NATURALIZED SPECIES THEMSELVES IE TO UNDERSTAND PATTERNS AND
PROCESSES RELEVANT TO INVASION BIOLOGY &INALLY WE WILL EXPLORE AN UNDERSTUDIED
TOPIC IN THE BIOGEOGRAPHY OF NATURALIZED SPECIES NAMELY THE SPECIES
AREA RELA
TIONSHIPS OF NATURALIZED SPECIES FOCUSING AMONG OTHER TOPICS ON THE ISSUE QUOTED
ABOVE BY $ARWIN IE THE CAPACITY OF ISLANDS TO HOLD ADDITIONAL SPECIES
! 2%6)%7 /& ")/'%/'2!0()# 0!44%2.3 /& .!452!,):%$ 30%#)%3
4HE lELDS OF BIOGEOGRAPHY AND INVASION BIOLOGY SHARE SIMILAR HISTORIES "OTH
lELDS HAVE BEEN THE SUBJECT OF STUDY BY MANY OF THE BEST
KNOWN ECOLOGISTS EVO
LUTIONARY BIOLOGISTS AND NATURALISTS OF THE TH THROUGH TH CENTURIES .OTABLE
CONTRIBUTIONS TO "IOGEOGRAPHY HAVE BEEN MADE BY ,INNEAUS "UFFON (OOKER
$ARWIN 7ALLACE $ARLINGTON $OBZHANSKY -AYR 'RINNELL %LTON ,ACK 7ILSON
AND -AC!RTHUR TO NAME BUT A FEW FOR A MORE COMPLETE LIST SEE ,OMOLINO ET AL
#ONTRIBUTORS TO THE lELD OF INVASION BIOLOGY HAVE BEEN NO LESS ILLUSTRIOUS
AND IN FACT ARE OFTEN MANY OF THE SAME INDIVIDUALS FOR A TREATMENT OF THIS SUB
JECT SEE $AVIS THIS VOLUME $ESPITE THEIR LONG HISTORIES BOTH BIOGEOGRAPHY AND
INVASION BIOLOGY HAVE ONLY RELATIVELY RECENTLY GARNERED SUFlCIENT ATTENTION TO
WARRANT THE ESTABLISHMENT OF JOURNALS AND SOCIETIES DEDICATED TO THESE TOPICS FOR
EXAMPLE THE *OURNAL OF "IOGEOGRAPHY WAS ESTABLISHED IN WHILE THE JOURNAL
"IOLOGICAL )NVASIONS WAS ESTABLISHED IN )T IS PERHAPS NOT SURPRISING THEN
THAT MOST RESEARCH THAT INTEGRATES THESE TWO lELDS HAS BEEN CONDUCTED IN THE PAST
DECADE 7HILE RELATIVELY RECENT THIS BODY OF WORK HAS GREAT POTENTIAL BOTH TO
INFORM OUR UNDERSTANDING OF BIOGEOGRAPHIC PATTERNS AND PROCESSES AS WELL AS TO
"IOGEOGRAPHIC PATTERNS OF NATURALIZED SPECIES
INFORM OUR UNDERSTANDING OF SPECIES INVASIONS (ERE WE REVIEW SEVERAL TOPICS THAT
HAVE BEEN PARTICULARLY RELEVANT TO BOTH THESE ENDEAVORS
,ATITUDINAL GRADIENTS IN RICHNESS AND RANGE SIZE
4HE LATITUDINAL GRADIENT IN SPECIES RICHNESS FROM A PEAK IN THE TROPICS TO A MINIMA
AT THE POLES IS ONE OF THE MOST GENERAL PHENOMENA IN BIOGEOGRAPHY "ROWN AND
,OMOLINO 3AX )N FACT UNLIKE MOST hGENERALv PATTERNS IN BIOLOGY
THERE ARE VERY FEW KNOWN EXCEPTIONS 0ERHAPS THE BEST KNOWN OF THESE EXCEPTIONS
IS THAT OF BENTHIC MARINE MACROALGAE WHICH GENERALLY PEAK IN RICHNESS AT MID
LATITUDES EG 'AINES AND ,UBCHENCO ,UNING -ID OR HIGH
LATITUDE
PEAKS IN RICHNESS ALSO OCCUR FOR INDIVIDUAL FAMILIES AND GENERA SUCH AS ICHNEUMO
NID WASPS PENGUINS AND PINES "ROWN AND ,OMOLINO 3TEVENS AND %NQUIST
)N SPITE OF THESE NOTABLE EXCEPTIONS HOWEVER ALL MAJOR GROUPS OF VERTE
BRATES MOST INVERTEBRATE GROUPS VASCULAR AND NON
VASCULAR PLANTS AND MANY
OTHER TAXONOMIC GROUPS SHOW THIS LATITUDINAL GRADIENT SEE REVIEW IN 7ILLIG ET AL
%UROPEAN NATURALISTS lRST PROMINENTLY DESCRIBED THIS PATTERN DURING THE
!GE OF %XPLORATION FOR EXAMPLE (UMBOLDT REPRINTED BY (AWKINS
ELOQUENTLY DESCRIBES INCREASES IN PLANT RICHNESS AS ONE APPROACHES THE TROPICS
4HE @GRADIENT ITSELF AS OPPOSED SIMPLY TO DIFFERENCES BETWEEN THE TEMPERATE ZONES
AND TROPICS BECAME MORE APPARENT AND BETTER CHARACTERIZED AS MORE BIOGEO
GRAPHIC DATA BECAME AVAILABLE DURING THE TWENTIETH CENTURY EG &ISCHER
3IMPSON FOR A REVIEW OF THIS TOPIC SEE "ROWN AND 3AX 2ESEARCH
DESCRIBING THIS GRADIENT CONTINUES TO THIS DAY EG 7ILLIG ET AL 2ODRIGUERO
AND 'ORLA
! SECOND GRADIENT THAT IS NOT AS WELL CHARACTERIZED IS THAT OF VARIATION IN
GEOGRAPHIC RANGE SIZE ACROSS LATITUDE 4HE lRST INKLING OF THIS PATTERN IS PRESENT
IN THE SEMINAL WORK OF %DUARDO 2APOPORT WHO DEMONSTRATED THAT THE
GEOGRAPHIC EXTENT OF MAMMALIAN SUBSPECIES TENDED TO INCREASE WITH INCREASING
LATITUDE )T WAS NOT UNTIL HOWEVER THAT 'EORGE 3TEVENS lRST CHARACTERIZED
THESE PATTERNS AT THE SPECIES LEVEL SHOWING THAT SPECIES TENDED ON AVERAGE TO HAVE
LARGE LATITUDINAL EXTENTS AT HIGHER LATITUDES 3TEVENS CHRISTENED THIS PAT
TERN 2APOPORTS RULE AND SET OFF A mURRY OF RESEARCH ON THIS TOPIC GREATLY REVITAL
IZING STUDIES OF LATITUDINAL GRADIENTS IN BIOGEOGRAPHY !T PRESENT THE GENERALITY
OF THE LATITUDINAL GRADIENT IN SPECIES RANGE SIZE IS MUCH DEBATED EG 'ASTON
ET AL BUT THERE IS BROAD AND UNCONTESTED EVIDENCE THAT THIS PATTERN IS ROBUST
WITHIN TEMPERATE LATITUDES OF THE .ORTH (EMISPHERE FOR MANY TAXONOMIC GROUPS
EG BIRDS MAMMALS CRAYlSH TREES ETC 3TEVENS &RANCE ,ETCHER
AND (ARVEY 'ASTON ET AL
7HAT IS ENTIRELY UNCLEAR FOR BOTH THESE GRADIENTS IS WHETHER THERE IS ANY UNI
VERSALLY APPLICABLE EXPLANATION FOR THEM /VER THE YEARS A NUMBER OF HYPOTHESES
HAVE BEEN PROPOSED TO EXPLAIN THESE GRADIENTS PARTICULARLY THE GRADIENT IN SPECIES
RICHNESS 0ERHAPS THE MOST OBVIOUS OF THESE EXPLANATIONS HAS BEEN THE OBSERVATION
THAT THERE IS GENERALLY MORE AVAILABLE ENERGY HEAT AND WATER AS ONE APPROACHES
$& 3AX AND 3$ 'AINES
THE TROPICS WHICH COULD PROMOTE OR MAINTAIN HIGHER LEVELS OF BIOLOGICAL DIVERSITY
IN TERRESTRIAL ENVIRONMENTS EG 0IANKA NOTE THAT WHILE VARIATION IN AVAIL
ABLE WATER IS CLEARLY NOT A LIMITING COMPONENT OF SPECIES RICHNESS IN THE OCEAN
THAT VARIATION IN ENERGY AND HEAT MAY BE OF DIRECT IMPORTANCE IN SURFACE WATERS
AND OF INDIRECT IMPORTANCE IN THE DEEP
SEA WHERE THE LATITUDINAL GRADIENT IN SPE
CIES RICHNESS ALSO OCCURS EG 7ILLIG ET AL /THER REASONABLE EXPLANATIONS
FOR THE LATITUDINAL GRADIENT IN SPECIES RICHNESS HOWEVER HAVE ALSO BEEN PROPOSED
AND MANY OF THESE INVOKE TEMPORALLY EXPLICIT EXPLANATIONS FOR THESE PATTERNS
&OR EXAMPLE IT HAS BEEN SUGGESTED THAT LONG
TERM STABILITY OF RESOURCE AVAILABILITY
OR CONDITIONS HAVE ALLOWED MORE SPECIES TO ACCUMULATE OR BE PRESERVED IN THE
TROPICS EG 0IANKA )N TERRESTRIAL ENVIRONMENTS FOR EXAMPLE THE EFFECTS
OF ICE
AGES AND OTHER CHANGES IN CLIMATE HAVE BEEN REDUCED IN THE TROPICS RELA
TIVE TO HIGH LATITUDE REGIONS WHERE GLACIERS HAVE REPEATEDLY CLEARED AWAY MOST
FORMS OF LIFE "ROWN AND ,OMOLINO /THER TEMPORALLY EXPLICIT EXPLANATIONS
SUGGEST THAT HIGHER SPECIATION RATES AT LOW LATITUDES MAY BE RESPONSIBLE FOR
THE RICHNESS GRADIENT EG 2OHDE &OR THE LATITUDINAL GRADIENT IN RANGE
SIZE SIMILAR EXPLANATIONS OR LOGICAL CONTINUATIONS OF THESE EXPLANATIONS HAVE
BEEN EMPLOYED FOR EXAMPLE GLACIERS OR OTHER FORMS OF ENVIRONMENTAL PERTURBA
TION MAY HAVE DIFFERENTIALLY REMOVED SPECIES WITH SMALL GEOGRAPHIC RANGES FROM
HIGH LATITUDES "ROWN 3AX 4HE GRADIENT IN RANGE SIZE HAS ALSO BEEN
SUGGESTED TO BE A CONSEQUENCE OF THE GRADIENT IN RICHNESS WITH THE ASSUMPTION
THAT REGIONS WITH HIGH SPECIES RICHNESS WOULD CAUSE THOSE SPECIES PRESUMABLY
BECAUSE OF SOME SORT OF LIMITING BIOTIC INTERACTIONS SUCH AS COMPETITION TO ON
AVERAGE HAVE SMALLER GEOGRAPHIC RANGES "ROWN
4HROUGH THE YEARS THE NUMBER OF HYPOTHESES PROPOSED PARTICULARLY FOR THE
LATITUDINAL GRADIENT IN SPECIES RICHNESS HAS SWELLED )N 0IANKA CHARACTERIZED
SIX PRINCIPLE HYPOTHESES BY 2OHDE CHARACTERIZED MORE THAN HYPOTHESES
AND TODAY THERE ARE MORE THAN HYPOTHESES FOR THESE GRADIENTS DESCRIBED IN
THE LITERATURE 7ILLIG ET AL 4HE TROUBLE CLEARLY HAS BEEN IN REDUCING THE
NUMBER OF EXPLANATIONS (EREIN HOWEVER LIES ONE OF THE PRINCIPAL DIFlCULTIES OF
MANY BIOGEOGRAPHIC PURSUITS NAMELY THE DIFlCULTY IN REJECTING HYPOTHESES WHEN
MANIPULATIVE EXPERIMENTS ARE IMPOSSIBLE OR NEARLY IMPOSSIBLE TO PURSUE AND THE
RELATIVE MERITS OF INDIVIDUAL HYPOTHESES ARE INSTEAD MEASURED AGAINST CORRELATIVE
DATA (EREIN ALSO LIES ONE OF THE PRINCIPAL ADVANTAGES OF STUDYING NATURALIZED SPE
CIES AS THEIR INTRODUCTION AND SUBSEQUENT NATURALIZATION PROVIDES A MANIPULATION
FOR A LARGE UNCONTROLLED @EXPERIMENT ONE THAT WOULD OTHERWISE BE IMPOSSIBLE TO
PERFORM AT SUCH LARGE SPATIAL AND TEMPORAL SCALES 3IMILARLY THE OCCURRENCE OF
THESE UNCONTROLLED EXPERIMENTS IN MANY REGIONS AND CONTINENTS AROUND THE WORLD
PROVIDES THE REPLICATION NEEDED TO DRAW GENERAL INSIGHTS FROM THE @RESULTS OF THESE
EXPERIMENTS
4HE SPECIlC ADVANTAGE OF STUDYING NATURALIZED SPECIES WITH RESPECT TO THIS
ISSUE IS THAT THE GREAT MAJORITY OF THESE SPECIES HAVE ONLY BEEN PRESENT IN THEIR
NATURALIZED RANGES FOR A FEW HUNDRED YEARS AT MOST 4HIS MEANS THAT ANY PATTERNS
IN GEOGRAPHIC GRADIENTS IN RICHNESS OR RANGE SIZE OBSERVED IN NATURALIZED SPECIES
"IOGEOGRAPHIC PATTERNS OF NATURALIZED SPECIES
CANNOT BE DIRECTLY ATTRIBUTABLE TO EFFECTS OF GLACIATIONS OTHER HISTORICAL PROCESSES
THAT HAVE DIFFERENTIALLY AFFECTED DIFFERENT LATITUDES OR TO DIFFERENT EVOLUTIONARY
RATES OF ACCUMULATION OVER TIME THE POTENTIAL INDIRECT EFFECTS OF THESE PROCESS
WILL BE DISCUSSED LATER 4HIS MEANS THAT IF ONE CAN ACCOUNT FOR DIFFERENCES IN PAT
TERNS OF INTRODUCTION WE MAY BE ABLE TO LEARN A GREAT DEAL ABOUT WHAT STRUCTURES
OR WHAT DOESNT STRUCTURE THE GEOGRAPHIC DISTRIBUTIONS OF SPECIES BY STUDYING
PATTERNS OF NATURALIZED SPECIES
4O EXAMINE THESE ISSUES 3AX EXAMINED THE GEOGRAPHIC DISTRIBUTION OF
NATURALIZED SPECIES OF SEVERAL TAXONOMIC GROUPS ON SEVERAL CONTINENTS PLANTS IN
%UROPE lSHES AND BIRDS IN .ORTH !MERICA AND MAMMALS IN %UROPE .ORTH AND
3OUTH !MERICA 4HIS WORK SHOWS THAT WITHIN CONTINENTAL AREAS OF THE TROPICS FEW
SPECIES HAVE INVADED AND THAT THOSE THAT HAVE TEND TO BE HUMAN COMMENSALS
WITH VERY LARGE GEOGRAPHIC RANGES WE SHALL RETURN TO THE ISSUE OF FEW INVASIONS
WITHIN TROPICAL MAINLAND AREAS IN A SECTION BELOW !T EXTRA
TROPICAL LATITUDES
HOWEVER WHERE MOST NATURALIZED SPECIES ARE DISTRIBUTED NATURALIZED SPECIES
SHOW GEOGRAPHIC PATTERNS OF DISTRIBUTION THAT ARE QUALITATIVELY SIMILAR TO THOSE
SHOWN BY NATIVE SPECIES IE RICHNESS PEAKS AT LOW LATITUDES AND DECLINES TOWARD
THE POLES WHILE AVERAGE GEOGRAPHIC RANGE SIZE IS SMALLEST AT LOW LATITUDES AND
INCREASES TOWARDS THE POLES &IGS AND FOR A DISCUSSION OF THE SMALL IDIOSYN
CRASIES IN THESE PATTERNS SEE 3AX 4HESE LATITUDINAL PATTERNS DO NOT APPEAR
TO BE DRIVEN EXCLUSIVELY BY PATTERNS OF INTRODUCTION EFFORT PER SE AT LEAST FOR BIRDS
BECAUSE THE GREATEST NUMBER OF INTRODUCTION ATTEMPTS FOR BIRDS IN .ORTH !MERICA
HAVE BEEN AT HIGH TEMPERATE LATITUDES IE ABOVE DEGREES LATITUDE WHERE
HUMAN POPULATIONS ARE LARGEST AND NOT AT LOW TEMPERATE LATITUDES IE BELOW
DEGREES WHERE THE NUMBER OF NATURALIZED SPECIES IS GREATEST 4HESE PATTERNS ALSO
DO NOT APPEAR TO BE AN INDIRECT CONSEQUENCE OF PATTERNS THAT EXISTED IN NATURALIZED
SPECIES NATIVE RANGES SEE 3AX FOR EVIDENCE AND DETAILS 4HIS SUGGESTS THAT
LATITUDINAL GRADIENTS IN RICHNESS AND RANGE SIZE CAN FORM RELATIVELY QUICKLY OVER
THE ECOLOGICAL TIME OF A FEW HUNDRED YEARS SINCE EXOTIC SPECIES WERE INTRODUCED
AND THAT TEMPORALLY DEPENDENT EXPLANATIONS FOR THESE GRADIENTS SUCH AS GLACIA
TION EVENTS AND DIFFERENTIAL RATES OF SPECIATION MAY NOT BE NECESSARY TO EXPLAIN
THESE PATTERNS 4HIS REPRESENTS A POTENTIALLY SIGNIlCANT CONTRIBUTION TO OUR UNDER
STANDING OF THESE GRADIENTS BY PROVIDING A MEANS BY WHICH WE CAN POTENTIALLY
REJECT CERTAIN HYPOTHESES AS GENERAL AND NECESSARY EXPLANATIONS
)N ADDITION TO LEARNING ABOUT WHAT STRUCTURES BIOGEOGRAPHIC GRADIENTS STUDY
ING THESE PATTERNS WITH NATURALIZED SPECIES ALSO PROVIDES INSIGHTS INTO INVASION
BIOLOGY )F SPECIES ARE MORE LIKELY TO BECOME ESTABLISHED AT LOW TEMPERATE LATI
TUDES THAN HIGH TEMPERATE LATITUDES INDEPENDENT OF DELIBERATE ATTEMPTS AT INTRO
DUCTION 3AX THEN REGIONS MAY BE DIFFERENTIALLY SUSCEPTIBLE TO INVASION
4HE EVIDENCE PRESENTED HOWEVER SUGGESTS THAT WE SHOULD ADOPT A MORE COMPLI
CATED VISION OF hINVASION SUSCEPTIBILITYv THAN IS GENERALLY USED 4HIS IS BECAUSE
DESPITE MORE SPECIES INVADING LOW TEMPERATE LATITUDES THESE NON
NATIVE SPECIES
TEND TO ACQUIRE SMALL GEOGRAPHIC RANGES )N CONTRAST FEW SPECIES TEND TO INVADE
HIGH LATITUDES BUT THE ONES THAT DO TEND TO ACQUIRE LARGE GEOGRAPHIC RANGES
$& 3AX AND 3$ 'AINES
&IG ,ATITUDINAL GRADIENTS IN RICHNESS OF NATURALIZED SPECIES 4HE SPECIES RICHNESS
OF NATURALIZED EXOTIC SPECIES IN $ LATITUDINAL BANDS ON CONTINENTS 4HE DASHED LINES
INDICATE THE 4ROPIC OF #ANCER OR #APRICORN DIVIDING TROPICAL AND TEMPERATE LATITUDES
&ISH DISTRIBUTIONS WERE EXAMINED AT TEMPERATE LATITUDES ONLY 2EPRINTED WITH PERMISSION
OF "LACKWELL 0UBLISHING FROM 3AX
"IOGEOGRAPHIC PATTERNS OF NATURALIZED SPECIES
&IG ,ATITUDINAL GRADIENTS IN GEOGRAPHIC RANGE SIZE OF NATURALIZED SPECIES 4HE MEAN
LATITUDINAL EXTENT OF NATURALIZED EXOTIC SPECIES IN $ LATITUDINAL BANDS ON CONTINENTS %RROR
BARS REPRESENT OR n ONE STANDARD ERROR 0OINTS WITHOUT ERROR BARS ARE BASED ON A SINGLE
SPECIES 4HE DASHED LINES INDICATE THE 4ROPIC OF #ANCER OR #APRICORN DIVIDING TROPICAL AND
TEMPERATE LATITUDES 2EPRINTED WITH PERMISSION OF "LACKWELL 0UBLISHING FROM 3AX
$& 3AX AND 3$ 'AINES
4HIS SUGGESTS A TRADEOFF IN SUSCEPTIBILITY OF INVASION BETWEEN REGIONS THAT IS BAL
ANCED BETWEEN THE ABSOLUTE NUMBER OF LIKELY INVADERS AND THE ABSOLUTE GEOGRAPH
IC EXTENT THOSE INVADERS ARE LIKELY TO ACHIEVE
-ANY QUESTIONS REGARDING THESE PATTERNS REMAIN UNANSWERED !RE THESE
PATTERNS GENERAL TO ALL CONTINENTS AND ALL TAXONOMIC GROUPS OF NATURALIZED SPE
CIES 7ILL THESE PATTERNS CHANGE OVER TIME AS MORE INTRODUCED SPECIES BECOME
ESTABLISHED AND AS EXISTING SPECIES CONTINUE TO EXPAND THEIR GEOGRAPHIC RANGES
!ND WILL FUTURE RANGE COLLAPSES IN THE GEOGRAPHIC DISTRIBUTION OF SOME NATURAL
IZED SPECIES AFFECT THESE PATTERNS !NSWERING THESE QUESTIONS WILL INVOLVE MUCH
WORK BUT DOING SO OFFERS THE PROMISE OF GREAT INTELLECTUAL CONTRIBUTIONS TO OUR
UNDERSTANDING OF BOTH BIOGEOGRAPHY AND SPECIES INVASIONS
)NVASIONS OF HOTSPOTS OF DIVERSITY
)N CONTRAST TO EXPERIMENTAL WORK DONE AT SMALL SPATIAL SCALES AND IN CONTRAST TO
THE SUPPOSITION OF %LTON THAT SPECIES
RICH AREAS SHOULD BE MORE DIFlCULT TO
INVADE THAN SPECIES
POOR ONES WORK AT BIOGEOGRAPHIC SCALES HAS GENERALLY SHOWN
THAT NATIVE AND NATURALIZED RICHNESS OF SPECIES ARE POSITIVELY CORRELATED ACROSS
LANDSCAPES AND REGIONS 4WO PAPERS THAT DREW PARTICULAR ATTENTION TO THIS PHE
NOMENON WERE PUBLISHED IN
,ONSDALE EXAMINED PUBLISHED STUDIES OF PLANT RICHNESS FROM
AROUND THE WORLD THE SPATIAL EXTENT COVERED BY THESE STUDIES VARIED FROM A FEW
SQUARE KILOMETERS TO GREATER THAN A MILLION SQUARE KILOMETERS AND THE REGIONS
COVERED INCLUDED ISLANDS AND CONTINENTS TEMPERATE AND TROPICAL AREAS "Y EXAM
INING THE mORAS THAT DISTINGUISHED BETWEEN NATIVE AND EXOTIC SPECIES HE WAS
ABLE TO SHOW THAT NATIVE AND EXOTIC SPECIES RICHNESS WERE SIGNIlCANTLY POSITIVELY
CORRELATED ACROSS THESE SITES EVEN AFTER CONTROLLING FOR DIFFERENCES IN AREA AMONG
SITES 4HE TOTAL AMOUNT OF VARIANCE IN THE NUMBER OF EXOTIC SPECIES PREDICTED BY
THE NUMBER OF NATIVE SPECIES WAS IMPROVED BY CONSIDERING WHETHER A PARTICULAR
SITE WAS AN ISLAND OR A RESERVE HOWEVER EVEN WITHOUT THIS INFORMATION A RELATIVE
LY CLEAR AND ROBUST PATTERN HAD BEEN IDENTIlED NATIVE AND EXOTIC RICHNESS WERE
POSITIVELY CORRELATED AT BIOGEOGRAPHIC SCALES
3TOHLGREN ET AL COLLECTED lELD DATA ON NATIVE AND EXOTIC PLANT SPECIES AT
LOCAL SCALES ACROSS VEGETATION TYPES AND WITHIN BIOMES IN THE WESTERN 53 !T THE
SCALE OF VEGETATION TYPES AND BIOMES THEY FOUND A POSITIVE RELATIONSHIP BETWEEN
NATIVE AND EXOTIC RICHNESS &URTHER 3TOHLGREN ET AL INDICATED THE OBVIOUS
BUT PREVIOUSLY UNARTICULATED OBSERVATION THAT THE AREAS RICHEST IN NATIVE SPECIES
WERE IN FACT BEING INVADED BY THE MOST EXOTIC SPECIES
4HIS WORK BY ,ONSDALE AND 3TOHLGREN ET AL HAS SUBSEQUENTLY
BEEN CONlRMED AT BIOGEOGRAPHIC SPATIAL SCALES BY SEVERAL OTHER STUDIES -OST OF
THIS WORK HAS ALSO BEEN DONE WITH PLANTS 3TADLER ET AL HAVE SHOWN A POSI
TIVE CORRELATION BETWEEN NATIVE AND EXOTIC PLANT RICHNESS IN ECOGEOGRAPHIC ZONES
OF +ENYA -C+INNEY A FOUND THIS PATTERN IN PROTECTED AREAS OF THE 53
0YSEK ET AL FOUND THIS PATTERN IN NATURE RESERVES OF THE #ZECH 2EPUBLIC
"IOGEOGRAPHIC PATTERNS OF NATURALIZED SPECIES
$EUTSCHEWITZ ET AL FOUND THIS PATTERN IN REGIONS OF THE $ESSAU DISTRICT
OF 'ERMANY AND lNALLY 3TOHLGREN ET AL SHOW THAT NATIVE AND EXOTIC
RICHNESS ARE POSITIVELY CORRELATED ACROSS COUNTIES WITHIN 53 3TATES AND ACROSS
THE STATES THEMSELVES 7HILE THE FOCUS OF THESE PATTERNS HAS BEEN ON PLANTS AN
EXPLORATION OF OTHER GROUPS PARTICULARLY AT LARGE SPATIAL SCALES SHOWS SIMILAR
PATTERNS PARTICULARLY WITHIN THE TEMPERATE ZONES &OR EXAMPLE A COMPARISON OF
NATIVE AND EXOTIC MAMMALIAN RICHNESS WITHIN
DEGREE BANDS OF LATITUDE IN TEM
PERATE .ORTH !MERICA REVEALS A STRIKINGLY STRONG POSITIVE RELATIONSHIP BETWEEN
NATIVE AND EXOTIC RICHNESS &IG A 3IMILARLY DATA FROM 3AX ET AL SHOWS
THAT NATIVE AND EXOTIC BIRD RICHNESS ARE POSITIVELY CORRELATED ON OCEANIC ISLANDS
&IG B )N CONTRAST HOWEVER TO THESE POSITIVE CORRELATIONS AT LEAST ONE NEGATIVE
CORRELATION BETWEEN NATIVE AND EXOTIC SPECIES AT BIOGEOGRAPHIC SCALES IS KNOWN
FROM THE LITERATURE IE FOR FRESHWATER lSHES IN WATERSHEDS OF .ORTH !MERICA 'IDO
AND "ROWN 4HERE ARE SOME REASONS WHICH WE WILL RETURN TO LATER THAT
WE MIGHT EXPECT ATYPICAL PATTERNS FROM FRESHWATER lSHES $ESPITE THIS EXCEPTION
HOWEVER WITH THE EVIDENCE AVAILABLE TO DATE IT APPEARS THAT THE GENERAL PATTERN
IS FOR NATIVE AND EXOTIC RICHNESS TO BE CORRELATED ACROSS LARGE AREAS AT LEAST WITHIN
A GIVEN REGION ALTHOUGH COMPARISONS OF AREAS ACROSS REGIONS MAY YIELD DIFFER
ENT RESULTS A TOPIC WE RETURN TO IN THE SECTION BELOW ON INVASIONS OF MAINLAND
TROPICAL AREAS
&IG 0OSITIVE CORRELATION BETWEEN NATIVE AND NATURALIZED SPECIES RICHNESS ! 2ICHNESS
OF NATIVE AND NATURALIZED MAMMAL SPECIES IN DEGREE BELTS OF LATITUDE FROM DEGREES
IN .ORTH !MERICA 2 P DATA FROM 3TEVENS AND 3AX
" 2ICHNESS OF HISTORIC NUMBER OF NATIVE AND CURRENT NUMBER OF NATURALIZED BIRD SPECIES
ON OCEANIC ISLANDS 2 P DATA FROM 3AX ET AL
$& 3AX AND 3$ 'AINES
4HIS RELATIVELY SIMPLE OBSERVATION THAT NATIVE AND EXOTIC RICHNESS ARE POSITIVELY
CORRELATED ACROSS LARGE AREAS COMES IN CONTRAST WITH MUCH WORK DONE AT EXPERI
MENTAL SCALES THAT SHOWS A NEGATIVE RELATIONSHIP BETWEEN NATIVE RICHNESS AND
EXOTIC INVASION EG 4ILMAN 3TACHOWICZ ET AL .AEEM ET AL
4HIS EXPERIMENTAL WORK IS CONSISTENT WITH MODELING WORK EG #ASE AND
PERHAPS MORE SIGNIlCANTLY WITH THE SUPPOSITION OF %LTON THAT SPECIES RICH
AREAS SHOULD BE MORE DIFlCULT TO INVADE THAN SPECIES POOR AREAS ! NUMBER OF STUD
IES HAVE ATTEMPTED TO RECONCILE THE DIFFERENCES OBSERVED BETWEEN EXPERIMENTAL
AND OBSERVATIONAL STUDIES EG 7ARDLE &RIDLEY ET AL 0ERHAPS THE
MOST CONVINCING EXPLANATION FOR THESE DIFFERENCES HAVE BEEN ARTICULATED BY ,EVINE
WHO SUGGESTS THAT REGARDLESS OF ANY IMPACT NATIVE RICHNESS MAY HAVE
ON SPECIES INVASIONS WHEN ALL ELSE IS EQUAL IN CONTROLLED EXPERIMENTAL SETTINGS
THAT THESE IMPACTS WILL BE OUTWEIGHED BY REAL
WORLD SPATIAL VARIATION IN ENVIRON
MENTAL CONDITIONS SUCH AS NUTRIENT AND ENERGY AVAILABILITY PRECIPITATION AND
HABITAT HETEROGENEITY THAT PROMOTE COVARIANCE IN THE RICHNESS OF BOTH NATIVE AND
EXOTIC SPECIES
4HIS EXPLANATION BY ,EVINE FOR RECONCILING DIFFERENCES OBSERVED IN
OBSERVATIONAL AND EXPERIMENTAL STUDIES SUGGESTS THAT NATIVE AND EXOTIC RICHNESS
SHOULD BE EXPECTED TO POSITIVELY COVARY WHENEVER BOTH GROUPS RESPOND SIMILARLY
TO ENVIRONMENTAL VARIABLES IE THAT SITE CONDITIONS THAT PROMOTE HIGH NATIVE RICH
NESS WILL OFTEN ALSO PROMOTE HIGH EXOTIC RICHNESS /THER EXPLANATIONS FOR A POSITIVE
ASSOCIATION BETWEEN NATIVE AND EXOTIC RICHNESS HOWEVER ARE ALSO POSSIBLE /NE
ALTERNATIVE EXPLANATION IS THAT INTRODUCTION EFFORTS BY HUMANS IE PROPAGULE
PRESSURE OF EXOTIC SPECIES ARE HIGHER IN SPECIES
RICH THAN SPECIES
POOR ENVIRON
MENTS SUCH THAT REGIONS WITH MANY NATIVE SPECIES ARE ALSO THE SAME REGIONS
WHERE HUMANS INTRODUCE THE MOST SPECIES &OR THIS TO BE A UNIVERSAL EXPLANATION
FOR THESE PATTERNS HOWEVER THE PATTERN OF INTRODUCTION EFFORT BY HUMANS WOULD
HAVE TO CONSISTENTLY MATCH NATIVE SPECIES RICHNESS ACROSS REGIONS WHICH SEEMS
UNLIKELY TO BE GENERALLY TRUE AND IS DElNITELY NOT THE CASE IN MANY PLACES EG BIRD
INTRODUCTION EFFORT IN .ORTH !MERICA DOES NOT MATCH PATTERNS OF NATIVE SPECIES
RICHNESS SEE ABOVE ! SECOND ALTERNATIVE EXPLANATION IS THAT THE POSITIVE ASSOCIA
TION BETWEEN NATIVE AND EXOTIC RICHNESS ACROSS REGIONS IS DUE TO DIFFERENCES IN THE
AREA OF REGIONS COMPARED "ECAUSE OF THE SPECIES
AREA RELATIONSHIP LARGE REGIONS
SHOULD HAVE MORE NATIVE AND MORE EXOTIC SPECIES (OWEVER WHILE SPECIES
AREA
RELATIONSHIPS CERTAINLY MAY BE IMPORTANT IN DETERMINING SUCH PATTERNS POSITIVE
RELATIONSHIPS BETWEEN NATIVE AND EXOTIC RICHNESS ARE OBSERVED EVEN AMONG EQUAL
SIZE AREAS EG 3AX OR WHEN THE EFFECT OF AREA IS CONTROLLED EG ,ONSDALE
'IVEN THAT THESE ALTERNATIVE EXPLANATIONS ARE UNLIKELY TO DRIVE THE POSI
TIVE RELATIONSHIP FREQUENTLY OBSERVED BETWEEN NATIVE AND EXOTIC RICHNESS IT SEEMS
LIKELY THAT THE OBSERVED RELATIONSHIPS ARE INDEED DUE TO VARIATION IN ENVIRONMEN
TAL CHARACTERISTICS SUCH AS ENERGY AND NUTRIENT AVAILABILITY THAT INmUENCE NATIVE
AND EXOTIC SPECIES RICHNESS IN SIMILAR WAYS
)F ENVIRONMENTAL CHARACTERISTICS ARE DRIVING THE POSITIVE ASSOCIATION BETWEEN
NATIVE AND EXOTIC RICHNESS OBSERVED ACROSS SITES THEN THESE PATTERNS HAVE IMPOR
"IOGEOGRAPHIC PATTERNS OF NATURALIZED SPECIES
TANT IMPLICATIONS FOR CONSERVATION AND ENVIRONMENTAL MANAGEMENT 4HEY SUG
GEST THAT SPECIES RICH NATIVE PRESERVES CANNOT BE EXPECTED TO REPEL EXOTIC SPECIES
INVASIONS AND THAT APPROPRIATE MANAGEMENT CONTROL EFFORTS FOR EXOTIC SPECIES
MUST BE MAINTAINED TO PREVENT THEM FROM BECOMING ESTABLISHED &URTHER WITH
RESPECT TO ECOLOGICAL THEORY THESE PATTERNS AND ITS LIKELY EXPLANATION SUGGEST
TWO THINGS &IRST THEY SUGGEST THAT EVEN DIVERSE CONTINENTAL AREAS CAN OFTEN SUP
PORT MANY ADDITIONAL SPECIES 3ECOND THEY SUGGEST THAT THE RICHNESS OF NATIVE
SPECIES WITHIN AN AREA MAY INDICATE THE RELATIVE CAPACITY OF THAT AREA TO SUPPORT
SPECIES SINCE SITES RICH IN NATIVE SPECIES HAVE PRESUMABLY INCREASED IN RICHNESS
THE MOST WHILE SITES POOR IN NATIVE SPECIES HAVE PRESUMABLY INCREASED IN RICHNESS
THE LEAST
4ROPICAL MAINLANDS REGIONS WITH FEW INVADERS
"ECAUSE NATIVE AND EXOTIC RICHNESS ARE POSITIVELY CORRELATED ACROSS SITES AT LARGE
SPATIAL SCALES ONE MIGHT SUSPECT THAT TROPICAL REGIONS WHERE THE GLOBAL PEAKS IN
NATIVE RICHNESS EXIST ARE ALSO INVADED BY MANY EXOTIC SPECIES .OTHING HOWEVER
COULD BE FURTHER FROM THE TRUTH ESPECIALLY IN MAINLAND TROPICAL AREAS WHERE
RELATIVELY FEW EXOTICS HAVE GENERALLY INVADED 2EJMANEK 3AX &INE
4HIS MEANS THAT THE GENERALLY POSITIVE CORRELATION BETWEEN NATIVE AND
EXOTIC RICHNESS OFTEN BREAKS DOWN WHEN COMPARING ACROSS AREAS IN BOTH TEMPER
ATE AND TROPICAL ZONES EG THE STRONG PATTERN DEPICTED IN &IG ! WOULD BREAK
DOWN IF TROPICAL POINTS WERE INCLUDED ON THE PLOT .EVERTHELESS WITHIN TROPICAL
AREAS THE POSITIVE CORRELATION BETWEEN NATIVE AND EXOTIC RICHNESS CAN STILL HOLD
EG IN -EXICAN 3TATES NATIVE AND EXOTIC SPECIES OF mOWERING PLANTS ARE POSITIVELY
CORRELATED 2 P DATA FROM 6ILLASEOR AND %SPINOSA
'ARCIA
!LSO UNLIKE TROPICAL MAINLAND ENVIRONMENTS ON TROPICAL ISLANDS MANY
EXOTIC SPECIES HAVE BECOME NATURALIZED EG 3AX ET AL 7HY HOWEVER
SO FEW EXOTIC SPECIES HAVE BECOME ESTABLISHED IN MAINLAND TROPICAL AREAS IN NOT
CLEARLY KNOWN
4WO PRINCIPAL STUDIES TO DATE HAVE PROVIDED EMPIRICAL DATA SHOWING THE CON
TRAST BETWEEN THE NUMBER OF EXOTIC SPECIES THAT HAVE BECOME NATURALIZED IN TROPI
CAL MAINLAND TROPICAL ISLAND AND TEMPERATE AREAS 2EJMANEK EXAMINED
THE NUMBERS OF NATIVE AND EXOTIC PLANTS IN VARIOUS mORAS OF THE WORLD SHOWING
THAT THE RELATIVE PROPORTION OF EXOTIC SPECIES AS WELL AS THE ABSOLUTE NUMBER OF
EXOTICS AFTER CONTROLLING FOR AREA DECLINES PRECIPITOUSLY WITHIN TROPICAL MAIN
LAND REGIONS RELATIVE TO TROPICAL ISLANDS OR TEMPERATE AREAS 3IMILARLY 3AX
SHOWS THAT FEW SPECIES OF BIRDS OR MAMMALS HAVE BECOME NATURALIZED ON CONTI
NENTS WITHIN THE TROPICS &IG BUT REPORTS THAT MANY HAVE BECOME ESTABLISHED
ON TROPICAL ISLANDS AND WITHIN TEMPERATE AREAS 4HUS BOTH STUDIES REPORT QUALI
TATIVELY SIMILAR PATTERNS FEW SPECIES HAVE INVADED MAINLAND TROPICAL AREAS WHILE
RELATIVELY MANY SPECIES HAVE INVADED TROPICAL ISLANDS AND TEMPERATE AREAS
4HE MECHANISM RESPONSIBLE FOR THIS PATTERN REMAINS ELUSIVE /NE POSSIBILITY
HOWEVER IS THAT PATTERNS OF HUMAN DISTURBANCE OR INTRODUCTION EFFORT OF EXOTIC
$& 3AX AND 3$ 'AINES
SPECIES ARE DIFFERENT IN MAINLAND TROPICAL AREAS THEN ELSEWHERE 7HILE EACH REGION
DOES OF COURSE HAVE ITS OWN UNIQUE HISTORY SUCH AN EXPLANATION DOES NOT APPEAR
TO BE GENERALLY TRUE &OR EXAMPLE MAINLAND -EXICO AND ISLANDS OF THE #ARIBBEAN
SHARE SIMILAR HISTORIES OF %UROPEAN COLONIZATION DISTURBANCE SUBSEQUENT POPULA
TION GROWTH AND PRESUMABLY INTRODUCTION EFFORTS YET THESE ISLANDS HARBOR MANY
MORE NATURALIZED SPECIES OF BIRDS AND MAMMALS THAN MAINLAND -EXICO
3AX !NOTHER POSSIBILITY IS THAT MAINLAND TROPICAL REGIONS ARE RESIS
TANT TO INVASION BECAUSE OF A COMBINATION OF BIOTIC AND ABIOTIC FACTORS 2EJMANEK
4HE POTENTIAL ROLE OF ABIOTIC FACTORS HOWEVER IS DIFlCULT TO RECONCILE WITH
THE DATA PROVIDED IN 3AX WHICH SHOWS THAT COASTAL AREAS OF TROPICAL CONTI
NENTS WHICH PRESUMABLY HAVE VERY SIMILAR ABIOTIC CONDITIONS AS TROPICAL ISLANDS
HAVE FEW EXOTIC BIRD OR MAMMAL SPECIES )N LIGHT OF THIS 3AX ARGUES THAT
ABIOTIC FACTORS SHOULD NOT BE THE LIMITING VARIABLE IN EXOTIC SPECIES INVASION OF
MAINLAND TROPICAL AREAS )NSTEAD HE ARGUES BASED ON THE DIFFERENTIAL PLACEMENT OF
NATURALIZED SPECIES LOW
LATITUDE RANGE BOUNDARIES ON CONTINENTS AND ISLANDS THAT
BIOTIC RESISTANCE WHICH MAY BE STRONGLY CORRELATED WITH SPECIES RICHNESS IS THE
PRIMARY FACTOR RESISTING SPECIES INVASIONS OF MAINLAND TROPICAL AREAS /F COURSE
IF HIGH SPECIES RICHNESS IN MAINLAND TROPICAL AREAS IS RESULTING IN SOME SORT OF
BIOTIC RESISTANCE TO INVASION THEN WHY DOESNT SPECIES RICHNESS CORRELATE WITH
INVASION RESISTANCE IN TEMPERATE LATITUDES !T THE MOMENT THERE IS NO CLEAR
ANSWER TO THIS QUESTION /NE POSSIBILITY HOWEVER IS THAT SOME THRESHOLD IN SPE
CIES RICHNESS MUST BE REACHED WHICH MAY BE MET IN MAINLAND TROPICAL AREAS
BEFORE BIOTIC RESISTANCE IS AN EFFECTIVE FORCE IN RESISTING SPECIES INVASIONS 3UCH AN
EXPLANATION WOULD BE CONSISTENT WITH THE GREATER APPARENT INVASIBILITY OF TROPICAL
ISLANDS AND TEMPERATE AREAS WHERE SPECIES RICHNESS IS REDUCED RELATIVE TO MAIN
LAND TROPICAL AREAS
4HE SUGGESTION THAT BIOTIC RESISTANCE IS IMPORTANT IN RESISTING SPECIES INVASION
OF MAINLAND TROPICAL AREAS IS ALSO CONSISTENT WITH BIOGEOGRAPHIC HYPOTHESES THAT
POSTULATE THAT LOW LATITUDE RANGE BOUNDARIES OF SPECIES OR THE ABILITY OF SPECIES
TO EXTEND THEIR GEOGRAPHIC RANGE CLOSER TO THE EQUATOR ARE SET BY BIOTIC INTERAC
TIONS WITH OTHER SPECIES EG $OBZHANSKY -AC!RTHUR 4HESE BIOTIC
INTERACTIONS ARE ASSUMED TO INCLUDE ALL NEGATIVE PAIRWISE SPECIES INTERACTIONS
IE COMPETITION PREDATION AND AMENSALISM BUT WITH RESPECT TO SPECIES INVA
SION COULD ALSO INCLUDE MISSING MUTUALISTIC SPECIES 2ICHARDSON ET AL /F
COURSE RESEARCH IN ECOLOGY SINCE THE TIME OF -AC!RTHUR HAS HAD A STRONG FOCUS
ON THE ROLE OF COMPETITION EG "RUNO ET AL )N SPITE OF THIS WORK IN INVA
SION BIOLOGY SUGGESTS THAT COMPETITION ITSELF MAY NOT BE AS IMPORTANT IN AFFECTING
SPECIES RICHNESS AS OTHER TYPES OF BIOTIC INTERACTIONS &OR EXAMPLE RECENT WORK BY
$AVIS SUGGESTS THAT COMPETITION FROM EXOTIC SPECIES SHOULD RARELY HAVE
NEGATIVE CONSEQUENCES ON NATIVE SPECIES RICHNESS &URTHER MOST OF THE NEGATIVE
CONSEQUENCES OF EXOTIC SPECIES ON NATIVE RICHNESS HAVE BEEN ASSOCIATED WITH INTRO
DUCED PREDATORS AND DISEASES EG BLACK RATS THE BROWN TREE SNAKE AND AVIAN
MALARIA EG %BENHARD &RITTS AND 2ODDA 3COTT ET AL )F THIS
INFORMATION CAN RECIPROCALLY BE APPLIED TO UNDERSTANDING WHAT PREVENTS SPECIES
"IOGEOGRAPHIC PATTERNS OF NATURALIZED SPECIES
INVASIONS THEN THIS WORK WOULD SUGGEST THAT PREDATION MEANT HERE IN THE BROAD
SENSE OF HERBIVORY PREDATION PARASITISM AND DISEASE COULD LIKELY EXPLAIN WHY
MAINLAND TROPICAL AREAS ARE SO DIFlCULT TO INVADE IN THE TROPICS PARTICULARLY THE
MAINLAND TROPICS NOT ONLY ARE THERE FOR EXAMPLE MORE MAMMALIAN PREDATORS
AND HERBIVORES BUT ALSO GREATER NUMBERS OF PARASITES AND DISEASES EG 7ILLIG
ET AL 2ODRIGUERO AND 'ORLA
)F PREDATION SENSU LATO IS IMPORTANT IN PREVENTING SPECIES INVASION OF MAINLAND
TROPICAL AREAS THEN THIS MAY SUGGEST DEPENDING ON HOW THRESHOLDS OF RICHNESS
OF PREDATORS OPERATE THAT PREDATION MAY BE IMPORTANT IN PREVENTING SPECIES
INVASION OF OTHER PLACES AS WELL BECAUSE FOR EXAMPLE EVEN IF PREDATION CANNOT
PREVENT SPECIES INVASION AT BIOGEOGRAPHIC SCALES IN TEMPERATE AREAS IT MAY STILL
BE IMPORTANT AT ECOLOGICAL SPATIAL SCALES )F THIS IS TRUE THEN IT COULD HAVE LARGE
IMPLICATIONS FOR ECOLOGICAL THEORY AND EXPERIMENTATION &OR EXAMPLE CURRENTLY IN
ECOLOGY MUCH EFFORT IS FOCUSED ON UNDERSTANDING THE ROLE OF RICHNESS IN PREVENTING
SPECIES INVASION -UCH OF THIS WORK HOWEVER IS FOCUSED ON THE ROLE THAT RICH
NESS OF COMPETITORS MAY HAVE IN PREVENTING SPECIES INVASION EG +ENNEDY ET AL
A COMPLIMENT TO THIS WOULD BE INCREASED ATTENTION TO THE ROLE THAT RICHNESS
OF PREDATORS INCLUDING HERBIVORES PARASITES AND DISEASES MAY PLAY IN PREVENTING
SPECIES INVASION
0ATTERNS OF INVASION ON ISLANDS
7HAT $ARWIN WROTE ABOUT ISLANDS ALMOST YEARS AGO COULDNT BE MORE
TRUE TODAY HUMANS HAVE BEEN lLLING ISLANDS hFAR MORE FULLYv THAN HAS NATURE
,IFEFORMS THAT WERE ENTIRELY ABSENT FROM MANY ISLANDS PARTICULARLY OCEANIC
ONES SUCH AS TERRESTRIAL MAMMALS AND FRESHWATER lSHES HAVE BEEN INTRODUCED
AND BECOME ESTABLISHED IN ABUNDANCE 3IMILARLY LIFEFORMS THAT WERE PRESENT BUT
LESS DIVERSE ON ISLANDS THAN IN SIMILAR SIZED AREAS ON CONTINENTS SUCH AS VASCULAR
PLANTS HAVE BECOME MUCH MORE SPECIOSE ON ISLANDS TODAY THAN IN THE PAST )N
MANY CASES THOSE SPECIES THAT HAVE BEEN INTRODUCED AND BECOME ESTABLISHED
HAVE HAD DEVASTATING EFFECTS ON THE NATIVE mORA AND FAUNA CAUSING EXTINCTIONS
IN SOME CASES AND REDUCTIONS IN THE ABUNDANCE OF NATIVE SPECIES IN MANY IF NOT
ALL CASES .ONETHELESS THE NET EFFECT OF THESE CHANGES HAS BEEN AN INCREASE IN NET
RICHNESS FOR MANY GROUPS OF SPECIES ON MANY ISLANDS EG 7ILSON #HOWN
ET AL 3AX ET AL 4HE CONSERVATION CHALLENGES POSED BY SUCH PATTERNS
ARE SEVERE AND COMPLEX BUT ARE NOT THE FOCUS OF THIS DISCUSSION )NSTEAD HERE WE
REPORT ON THE PATTERNS THEMSELVES AND ON WHAT THESE PATTERNS CAN TELL US ABOUT
BIOGEOGRAPHY INVASIVE SPECIES AND WHAT WE DO NOT YET UNDERSTAND
)SLAND mORAS AND FAUNAS ARE OFTEN DESCRIBED AS BEING OUT OF BALANCE OR NOT
HARMONIC WITH MAINLAND BIOTAS "ROWN AND ,OMOLINO 4HIS IS BECAUSE
EVENLY REPRESENTATIVE SUBSETS OF THE SUITE OF ORGANISMS FOUND ON CONTINENTS ARE
NOT FOUND ON ISLANDS )NSTEAD A DISPROPORTIONATE NUMBER OF SPECIES FOUND ON
ISLANDS ARE FROM TAXONOMIC GROUPS THAT ARE LIKELY TO DISPERSE ACROSS OCEAN WATERS
4HIS INCLUDES BIRDS AND INSECTS WHICH CAN OFTEN mY LONG DISTANCES OR BE BLOWN BY
$& 3AX AND 3$ 'AINES
STORMS TO DISTANT LOCALITIES PLANTS WHOSE PROPAGULES CAN BE BLOWN BY THE WIND
OR CARRIED BY BIRDS IN THEIR GUTS ON THEIR FEET ETC AND REPTILES WHICH ARE OFTEN
ABLE TO SURVIVE EXTENDED VOYAGES WITH LITTLE FOOD OR WATER ON hRAFTSv OF NATURAL
VEGETATION THAT HAVE BEEN WASHED OUT TO SEA BY STORMS "Y CONTRAST TAXONOMIC
GROUPS THAT ARE NOT LIKELY TO MAKE OR SURVIVE SUCH DISPERSAL EVENTS SUCH AS
FRESHWATER lSH TERRESTRIAL MAMMALS AND AMPHIBIANS ARE LIKELY TO BE ABSENT OR
DEPAUPERATE ON ISOLATED ISLANDS
/VER THE PAST DECADE MUCH HAS BEEN LEARNED ABOUT HOW HUMANS ARE MEDIAT
ING CHANGES ON ISLANDS FOR EXAMPLE THE IMPORTANCE OF INTRODUCED PREDATORS AND
PATHOGENS IN CAUSING NATIVE SPECIES EXTINCTIONS EG &RITTS AND 2ODDA
!TKINSON ET AL THE RELATIVE ROLE OF DIFFERENT ECOLOGICAL CORRELATES IN THE
ESTABLISHMENT SUCCESS OF INTRODUCED SPECIES EG $UNCAN ET AL AND HOW
THE ESTABLISHMENT OF NOVEL VEGETATION CAN FACILITATE THE COLONIZATION OF OTHER SPE
CIES EG (UTTON -UCH HAS ALSO BEEN LEARNED ABOUT HOW THE COMPOSITION
AND RICHNESS OF DIFFERENT TAXONOMIC GROUPS ARE CHANGING ON ISLANDS &OR EXAMPLE
THERE HAVE BEEN LARGE CHANGES IN THE COMPOSITION OF BIRD SPECIES LARGE INCREASES
IN THE TOTAL RICHNESS OF PLANT SPECIES COUPLED WITH SUBSTANTIAL DECLINES IN NATIVE
PLANT ABUNDANCE AND LARGE INCREASES IN THE PRESENCE OF TAXA THAT WERE PREVI
OUSLY RARE OR ENTIRELY ABSENT ON ISLANDS SUCH AS FRESHWATER lSHES AND NON VOLANT
MAMMALS EG 7ILSON 3AX ET AL 7HAT WE DO NOT YET UNDERSTAND
IS HOW THE SPECIES RICHNESS OF THESE DIFFERENT TAXONOMIC GROUPS WILL CHANGE IN
THE FUTURE OR WHETHER ISLANDS ARE APPROACHING ANY SORT OF MAXIMUM CAPACITY
FOR HOLDING ADDITIONAL SPECIES 4HESE ISSUES REQUIRE CONSIDERABLE ADDITIONAL WORK
BUT SOME KEY INSIGHTS CAN BE GLEANED BY EXAMINING SPECIES AREA RELATIONSHIPS
30%#)%3 !2%! 2%,!4)/.3()03 /& .!452!,):%$ 30%#)%3
4HE POSITIVE RELATIONSHIP BETWEEN INCREASING AREA AND INCREASING NUMBER OF SPE
CIES HAS BEEN KNOWN FOR SOME TIME 7HITTAKER !RRHENIUS HOW
EVER WAS THE lRST TO PLOT SPECIES AREA AGAINST SPECIES RICHNESS ON A LOG
LOG SCALE
$OING THIS LAID THE FOUNDATION FOR COMPARISONS BETWEEN SPECIES
AREA RELATIONSHIPS
FOR DIFFERENT TAXONOMIC GROUPS AND FOR DIFFERENT SETS OF LOCALITIES BECAUSE SPE
CIES
AREA RELATIONSHIPS ON LOG
LOG PLOTS COMMONLY TAKE A LINEAR FORM SIMPLIFYING
COMPARISONS OF SLOPES AND INTERCEPTS HOWEVER SEE ,OMOLINO AND 7EISER
FOR A DISCUSSION OF WHEN NON
LINEAR PATTERNS ARE EXPECTED 4HE SLOPES AND INTER
CEPTS OF LOG
LOG SPECIES
AREA RELATIONSHIPS CONTAIN INFORMATION ABOUT HOW SPECIES
ARE DISTRIBUTED HOW DISPERSAL
LIMITED DIFFERENT TAXONOMIC GROUPS ARE AND HOW
ISOLATION AND LIMITED AREA AFFECT SPECIES RICHNESS EG 2OSENZWEIG )N SPITE
OF THE USEFULNESS OF THESE PATTERNS IN CHARACTERIZING THE DISTRIBUTIONS OF SPECIES
THE PRECISE MECHANISMS FOR SPECIES
AREA RELATIONSHIPS ARE STILL STRONGLY CONTESTED
EG -ATTER ET AL 3PECIES
AREA RELATIONSHIP ARE BELIEVED TO BE A CONSE
QUENCE OF PATTERNS OF SPECIES ABUNDANCE THE INTERPLAY OF SPECIES ABUNDANCE
AND GEOGRAPHIC RANGE DISTRIBUTIONS THE NUMBER OF HABITATS PRESENT AND CONSE
"IOGEOGRAPHIC PATTERNS OF NATURALIZED SPECIES
QUENTLY THE VARIETY OF SPECIES SUPPORTED PASSIVE SAMPLING OF A LARGER FRACTION
OF A REGIONS SPECIES POOL LOWER EXTINCTION RISKS OCCURRING IN LARGER POPULA
TIONS A DYNAMIC RELATIONSHIP BETWEEN COLONIZATION AND EXTINCTION AND
THE DYNAMICS OF SPECIATION COLONIZATION AND EXTINCTION -ATTER ET AL
0ART OF THE DIFlCULTY IN TEASING THESE DIFFERENT HYPOTHESES APART IS THAT THEY CLEARLY
ARE NOT MUTUALLY EXCLUSIVE !DDITIONALLY THE MECHANISMS THAT ARE MOST IMPOR
TANT FOR THE SPECIES
AREA RELATIONSHIPS MAY VARY ACROSS SPATIAL SCALES FOR EXAMPLE
AT THE SMALL SPATIAL SCALES OF MOST ECOLOGICAL STUDIES MECHANISMS LIKE SPECIATION
WILL GENERALLY BE RELATIVELY UNIMPORTANT WHEREAS AT LARGER SPATIAL SCALES SPECIA
TION MAY BE A VERY IMPORTANT PROCESS )N THIS DISCUSSION WE RESTRICT OUR FOCUS TO
SPECIES
AREA RELATIONSHIPS OCCURRING AT LARGE SPATIAL SCALES IE AT SPATIAL SCALES
THAT EXCEED SEVERAL HECTARES
!LTHOUGH THERE IS A RICH LITERATURE ON SPECIES
AREA RELATIONSHIPS SEE REVIEWS
BY #ONNOR AND -C#OY 2OSENZWEIG AND ,OMOLINO RELATIVELY
LITTLE HAS BEEN PUBLISHED ON SPECIES
AREA RELATIONSHIPS OF NATURALIZED OR OTHER
WISE EXOTIC SPECIES !LTHOUGH IT SEEMS LIKELY THAT SOMEONE HAS EXAMINED A SPE
CIES
AREA RELATIONSHIP FOR EXOTIC SPECIES PRIOR TO YEARS AGO THE lRST PAPER WE
ARE AWARE OF THAT EXPLICITLY DISCUSSES SUCH A RELATIONSHIP IS BY #RAWLEY
WHO COMPARED THE SPECIES
AREA RELATIONSHIPS OF NATIVE AND EXOTIC PLANT SPECIES
-ORE RECENTLY 0YSEK EXAMINED THE SPECIES
AREA RELATIONSHIPS OF NATIVE AND
EXOTIC SPECIES IN CITIES OF CENTRAL %UROPE /THER SUBSEQUENT STUDIES THAT HAVE CON
SIDERED THE RELATIONSHIP BETWEEN AREA AND EXOTIC SPECIES INCLUDE WORK BY #HOWN
ET AL ON 3OUTHERN /CEAN )SLANDS BY ,ONSDALE ACROSS SITES
FROM AROUND THE WORLD BY -OODY ON THE #HANNEL )SLANDS OF #ALIFORNIA
BY -C+INNEY A IN PROTECTED AREAS OF THE 53 BY -C+INNEY B AND
2EJMANEK ON EXOTIC lSHES AND PLANTS IN 53 3TATES BY 0YSEK ET AL A
AND B ON EXOTIC PLANTS IN NATURE RESERVES OF CENTRAL %UROPE AND MOST
RECENTLY BY 2USSELL ET AL ON THE OFFSHORE ISLANDS OF .EW :EALAND (ERE WE
BUILD ON THIS PREVIOUS WORK TO CONSIDER WHAT EXAMINATIONS OF EXOTIC SPECIES
AREA
RELATIONSHIPS AND THEIR COMPARISONS WITH NATIVE SPECIES
AREA RELATIONSHIPS CAN
TELL US ABOUT SPECIES INVASION AND ABOUT THE FACTORS THAT LIMIT AND MAINTAIN SPECIES
RICHNESS )N PARTICULAR WE FOCUS ON TWO SETS OF ISSUES &IRST WHAT DO CONTRASTS AS
WELL AS SIMILARITIES BETWEEN THE FORM OF NATIVE AND EXOTIC SPECIES
AREA RELATION
SHIPS TELL US ABOUT INVASION AND BIOGEOGRAPHIC PROCESSES 3ECOND HOW DO CHANGES
IN SPECIES AREA
RELATIONSHIPS FOLLOWING SPECIES INVASIONS INFORM OUR UNDERSTAND
ING OF HOW SPECIES RICHNESS HAS CHANGED AND WILL CHANGE IN THE FUTURE AT DIFFERENT
SPATIAL SCALES AND IN LOCATIONS THAT VARY IN THEIR DEGREE OF ISOLATION
#ONTRASTS AND SIMILARITIES BETWEEN NATIVE AND NATURALIZED
SPECIES
AREA RELATIONSHIPS
3OME PREVIOUS WORK HAS CONSIDERED THE IMPLICATIONS OF DIFFERENCES IN THE SLOPES
OF SPECIES
AREA RELATIONSHIPS BETWEEN NATIVE AND EXOTIC SPECIES #RAWLEY
FOUND SIGNIlCANT SPECIES
AREA RELATIONSHIPS FOR NATIVE AND EXOTIC PLANT SPECIES
$& 3AX AND 3$ 'AINES
WHEN EXAMINING DATA FROM ACROSS REGIONS IN "RITAIN )RELAND EASTERN .ORTH
!MERICA AND 7ESTERN !USTRALIA (OWEVER HE FOUND THE SLOPE OF THE SPECIES
AREA
RELATIONSHIP TO BE STEEPER FOR NATIVE SPECIES WHICH HE SUGGESTED WAS A CONSE
QUENCE OF THE HIGHER BETA DIVERSITY OF NATIVE SPECIES )N ANOTHER STUDY 0YSEK
FOUND IN CITIES OF CENTRAL %UROPE THAT NEOPHYTES EXOTIC SPECIES THAT HAVE
INVADED THE REGION SINCE !$ SHOW A SIGNIlCANTLY STEEPER SPECIES
AREA SLOPE
THAN DO ARCHAEOPHYTES EXOTICS THAT INVADED PRIOR TO OR NATIVE SPECIES
(E SUGGESTED THAT THIS MIGHT BE DUE TO THE POTENTIALLY INHOSPITABLE ENVIRON
MENT FOR NEOPHYTES THAT EXISTS OUTSIDE URBAN AREAS IF THIS WERE TRUE CITIES WOULD
SERVE AS HABITAT hISLANDSv FOR NEOPHYTES THUS EXPLAINING THEIR STEEPER SPECIES
AREA SLOPE AS ISLANDS TYPICALLY EXHIBIT MUCH STEEPER SLOPES THAN COMPARABLE
CONTINENTAL REGIONS 2OSENZWEIG 2USSELL ET AL FOUND THE SLOPE OF
THE SPECIES
AREA RELATIONSHIP FOR EXOTIC MAMMALS ON OFFSHORE ISLANDS OF .EW
:EALAND TO BE 4HEY SUGGEST THAT THIS SLOPE WHICH IS SHALLOWER THAN THOSE
OFTEN SEEN ON ISLANDS MAY SUGGEST THAT THE INTERACTION BETWEEN AREA AND ISOLATION
IS REDUCED FOR THESE MAMMALS SINCE THEIR TRANSPORT AND INTRODUCTION HAS BEEN
FACILITATED BY HUMANS /F COURSE INTERPRETING DIFFERENCES BETWEEN THE SLOPES OF
DIFFERENT SPECIES
AREA RELATIONSHIPS IS NOT A TRIVIAL MATTER AND IS AN ACTIVITY WITH
MANY POTENTIAL PITFALLS EG -ARTIN ,OMOLINO .EVERTHELESS WE
BELIEVE THAT MAKING SUCH COMPARISONS BETWEEN NATIVE AND EXOTIC RELATIONSHIPS
MAY BE EXTREMELY VALUABLE !S THIS CHAPTER HOWEVER IS A lRST ATTEMPT TO SEARCH
FOR GENERALITIES FROM COMPARISONS OF MANY NATIVE AND EXOTIC SPECIES
AREA RELATION
SHIPS WE HERE EXAMINE A TOPIC THAT WE BELIEVE IS MORE READILY TRACTABLE NAMELY
A COMPARISON OF NATIVE AND EXOTIC SPECIES
AREA RELATIONSHIPS THAT DO AND DO NOT
SHOW SIGNIlCANT LINEAR RELATIONSHIPS ON A LOG
LOG SCALE
!LTHOUGH THE SPECIES
AREA RELATIONSHIP IS A VERY GENERAL PATTERN NOT ALL DATA SETS
SHOW A SIGNIlCANT RELATIONSHIP BETWEEN THESE VARIABLES &URTHER THE NATIVE AND
EXOTIC COMPONENTS OF THESE RELATIONSHIPS MAY NOT BEHAVE IN SIMILAR WAYS &OUR
OUTCOMES ARE POSSIBLE IN COMPARISONS OF NATIVE AND EXOTIC SPECIES
AREA RELATION
SHIPS NATIVES SHOW A SIGNIlCANT LINEAR RELATIONSHIP BUT EXOTICS DO NOT BOTH
NATIVES AND EXOTICS SHOW SIGNIlCANT RELATIONSHIPS EXOTICS SHOW A SIGNIlCANT
RELATIONSHIP BUT NATIVES DO NOT OR NEITHER NATIVES OR EXOTICS SHOW SIGNIlCANT
RELATIONSHIPS !S THE OCCURRENCE OF THE FOURTH OUTCOME IN WHICH NEITHER NATIVES
OR EXOTICS SHOW SIGNIlCANT SPECIES
AREA RELATIONSHIPS SHOULD BE EXCEEDINGLY RARE
WE RESTRICT OUR DISCUSSION HERE TO THE lRST THREE OUTCOMES LISTED ABOVE
3PECIES AREA RELATIONSHIPS FOR NATIVES BUT NOT FOR EXOTICS
7E BEGIN WITH THE SITUATION WHERE NATIVE SPECIES DO AND EXOTIC SPECIES DO NOT
SHOW A SIGNIlCANT POSITIVE LINEAR RELATIONSHIP ON A LOG
LOG SCALE BETWEEN AREA
AND SPECIES RICHNESS 7E ILLUSTRATE THIS WITH PLANT SPECIES IN #ALIFORNIA #OUNTIES
53 3TATES AND -EXICAN 3TATES &IG )N EACH CASE NATIVE SPECIES SHOW A STRONG
POSITIVE SPECIES
AREA RELATIONSHIP BUT EXOTIC SPECIES DO NOT )T IS IMPORTANT TO NOTE
THAT NOT ONLY ARE THE RELATIONSHIPS BETWEEN EXOTIC SPECIES RICHNESS AND AREA NOT
"IOGEOGRAPHIC PATTERNS OF NATURALIZED SPECIES
SIGNIlCANT BUT THAT THERE IS NO HINT OF A TREND TOWARDS A POSITIVE RELATIONSHIP
'IVEN THE BROAD SPATIAL RANGE EXAMINED NEARLY THREE ORDERS OF MAGNITUDE OF AREA
IN EACH CASE AND LARGE NUMBERS OF EXOTIC SPECIES MORE THAN EXOTIC SPE
CIES IN EACH CASE IT IS SOMEWHAT SURPRISING THAT EXOTIC SPECIES DO NOT SHOW THE
EXPECTED RELATIONSHIP 7E SUGGEST HERE THAT EXOTICS MAY NOT SHOW THE EXPECTED
RELATIONSHIP FOR ANY ONE OF THE FOLLOWING THREE REASONS EXOTIC SPECIES ARE
INHERENTLY DIFFERENT THAN NATIVE SPECIES AND SHOULD NOT BE EXPECTED TO RESPOND
IN SIMILAR WAYS TO AREA AS NATIVE SPECIES THERE ARE A LIMITED NUMBER OF EXOTIC
SPECIES WITHIN EACH OF THESE REGIONS SUCH THAT EVEN SMALL SUBREGIONS IE COUN
TIES OR STATES COULD HOLD ALL OR MOST OF THE EXOTIC SPECIES IN QUESTION THEREBY
MAKING IT UNLIKELY FOR THERE TO BE A SIGNIlCANT RELATIONSHIP AT THE SCALES ANALYZED
EXOTIC SPECIES ARE MORE STRONGLY INmUENCED BY THE DISTRIBUTION OF HUMAN POPU
LATIONS THAN BY AREA THIS COULD BE TRUE FOR ANY NUMBER OF REASONS SUCH AS A
CORRESPONDENCE BETWEEN HUMAN POPULATION SIZE AND INTRODUCTION EFFORT OF EXOTIC
SPECIES BETWEEN HUMAN POPULATION DENSITY AND DISTURBANCE WHICH MAY FAVOR
EXOTIC SPECIES OR BETWEEN HUMAN POPULATION DENSITY AND CLIMATE MATCHING THAT
OCCURS FROM SOURCE AREAS FOR EXOTIC SPECIES WHERE MANY HUMANS HAVE COLONIZED
FROM AND EXOTICS ARE LOCALLY ADAPTED AND RECIPIENT AREAS FOR EXOTIC SPECIES WHERE
MANY HUMANS NOW RESIDE AND EXOTICS ARE PREADAPTED FOR LOCAL CONDITIONS
$ISCRIMINATING AMONG THESE HYPOTHESES IS NOT TRIVIAL AS THEY ARE NEITHER
MUTUALLY EXCLUSIVE NOR NECESSARILY EXHAUSTIVE .EVERTHELESS WITH THE DATA AVAIL
ABLE TO US WE SHOULD BE ABLE TO DISCRIMINATE AMONG AT LEAST SOME OF OUR STATED
&IG 3PECIES AREA RELATIONSHIPS FOR NATIVE AND EXOTIC SPECIES OF PLANTS 3OLID CIRCLES ARE
NATIVE AND EMPTY CIRCLES ARE EXOTIC SPECIES OF VASCULAR PLANTS ! .ATIVES SHOW A POSITIVE
LINEAR RELATIONSHIP 2 Z # P WHILE EXOTICS DO NOT SHOW A
SIGNIlCANT RELATIONSHIP DATA FROM #ALmORA WWWCALmORAORG " .ATIVES SHOW A POSITIVE
LINEAR RELATIONSHIP 2 Z # P WHILE EXOTICS DO NOT SHOW
A SIGNIlCANT RELATIONSHIP SEE !PPENDIX FOR A DESCRIPTION OF DATA SOURCES AND METHODS
# .ATIVES SHOW A POSITIVE LINEAR RELATIONSHIP 2 Z # P
WHILE EXOTICS DO NOT SHOW A SIGNIlCANT RELATIONSHIP NOTE THAT THESE DATA ARE FOR mOWERING
PLANTS ONLY DATA FROM 6ILLASEOR AND %SPINOSA 'ARCIA
$& 3AX AND 3$ 'AINES
HYPOTHESES WITH THE GOAL OF BETTER UNDERSTANDING THE CURIOUS ABSENCE OF SPECIES
AREA RELATIONSHIPS FOR THESE EXOTIC SPECIES 4HE lRST HYPOTHESIS THAT OF INHERENT
DIFFERENCES BETWEEN NATIVE AND EXOTIC SPECIES SEEMS UNLIKELY THIS IS BECAUSE
THE CHARACTERISTICS OF THE NATIVE AND EXOTIC SPECIES IN QUESTION ARE UNLIKELY TO BE
SO DIFFERENT THAT THEY SHOULD RESPOND IN COMPLETELY DIFFERENT WAYS TO PHYSICAL OR
BIOTIC CHARACTERISTICS OF THE ENVIRONMENT THAT PROMOTE RICHNESS WITH INCREASING
AREA SUCH AS INCREASES IN HABITAT DIVERSITY .EVERTHELESS THIS IS A POSSIBILITY AND
ONE THAT IS DIFlCULT TO RULE OUT &OR -EXICAN 3TATES THIS DOES NOT APPEAR TO BE THE
CASE HOWEVER AS NATIVE AND EXOTIC RICHNESS ARE STRONGLY POSITIVELY CORRELATED
ACROSS STATES 2 P &OR #ALIFORNIA #OUNTIES AND 53 3TATES
HOWEVER NO SUCH POSITIVE RELATIONSHIP IS APPARENT LEAVING THIS HYPOTHESIS AS
A POSSIBILITY FOR THOSE AREAS 4HE SECOND HYPOTHESIS THAT OF EXOTIC SPECIES POOL
EXHAUSTION WITHIN INDIVIDUAL SAMPLE UNITS ALSO SEEMS UNLIKELY SINCE A RELATIVELY
SMALL FRACTION OF THE TOTAL NUMBER OF EXOTIC SPECIES IN THESE REGIONS ARE PRESENT
WITHIN INDIVIDUAL SAMPLE AREAS &OR EXAMPLE BETWEEN CA AND OF THE TOTAL
NUMBER OF EXOTIC PLANT SPECIES IN -EXICO ARE FOUND WITHIN INDIVIDUAL STATES SUG
GESTING THAT THE NUMBER OF EXOTIC SPECIES PRESENT WITHIN INDIVIDUAL STATES HAS NOT
APPROACHED OR EXHAUSTED THE TOTAL POOL OF SPECIES AVAILABLE IN -EXICO NOTE THAT
SIMILAR PATTERNS EXIST FOR #ALIFORNIA #OUNTIES AND 53 3TATES 4HE THIRD HYPOTH
ESIS WHICH SUGGESTS THAT HUMAN POPULATION DENSITY WILL INmUENCE EXOTIC SPECIES
RICHNESS IS STRONGLY SUPPORTED IN #ALIFORNIA #OUNTIES 53 AND -EXICAN 3TATES
WHERE IN ALL CASES HUMAN POPULATION DENSITY IS STRONGLY CORRELATED WITH EXOTIC
SPECIES RICHNESS &IG $IFFERENTIATING THE PROXIMATE CAUSE OF THIS PATTERN IS
DIFlCULT HOWEVER AS INTRODUCTION EFFORT DISTURBANCE OR SOME TYPE OF CLIMATE
MATCHING EXPLANATION ALL SEEM CREDIBLE )T IS CURIOUS TO NOTE ADDITIONALLY THAT
NATIVE SPECIES EITHER DO NOT SHOW A SIGNIlCANT RELATIONSHIP WITH HUMAN POPULA
TION IN #ALIFORNIA #OUNTIES OR SHOW RELATIONSHIPS WHERE LESS VARIANCE IS EXPLAINED
RELATIVE TO EXOTIC SPECIES IN 53 AND -EXICAN 3TATES INDICATING THAT HUMAN POPU
LATION IS A MUCH BETTER INDICATOR OF EXOTIC THAN NATIVE RICHNESS
3PECIES AREA RELATIONSHIPS FOR NATIVES AND EXOTICS
.ATIVE AND EXOTIC COMPONENTS OF PARTICULAR SPECIES
AREA RELATIONSHIPS IE FOR
PARTICULAR TAXA ACROSS A PARTICULAR SET OF AREAS CAN BOTH SHOW SIGNIlCANT POSITIVE
LINEAR RELATIONSHIPS ON PLOTS OF LOG AREA AND LOG SPECIES RICHNESS 7E ILLUSTRATE THIS
HERE WITH TWO EXAMPLES PLANTS ON THE #HANNEL )SLANDS OF #ALIFORNIA AND PLANTS
ON OCEANIC ISLANDS &IG " AND # .OTE ALSO THAT PLANTS IN MAINLAND SITES OF
#ALIFORNIA SHOW A NON
SIGNIlCANT BUT STILL POSITIVE TREND BETWEEN AREA AND EXOTIC
SPECIES RICHNESS AND A SIGNIlCANT POSITIVE RELATIONSHIP BETWEEN NATIVE RICHNESS
AND AREA &IG ! 3UCH PATTERNS IE WHERE BOTH NATIVE AND EXOTIC SPECIES SHOW
POSITIVE RELATIONSHIPS BETWEEN AREA AND RICHNESS CAN BE ARGUED TO BE CONSISTENT
WITH THE NULL HYPOTHESIS THAT SPECIES RICHNESS WILL ALWAYS BE INmUENCED BY AREA
!S SUCH THESE RESULTS ARE NOT PARTICULARLY SURPRISING BUT NEVERTHELESS OF INTEREST
BECAUSE THEY SUGGEST THAT NATIVE AND EXOTIC SPECIES ARE BOTH RESPONDING IN SIMILAR
"IOGEOGRAPHIC PATTERNS OF NATURALIZED SPECIES
&IG (UMAN POPULATION AND RICHNESS OF NATIVE AND EXOTIC SPECIES 3OLID CIRCLES
ARE NATIVE AND EMPTY CIRCLES ARE EXOTIC SPECIES ! %XOTIC SPECIES SHOW A POSITIVE LINEAR
RELATIONSHIP 2 P WHILE NATIVES DO NOT SHOW A SIGNIlCANT RELATIONSHIP
" %XOTIC AND NATIVE SPECIES SHOW POSITIVE LINEAR RELATIONSHIPS NATURALIZED SPECIES
2 P NATIVES 2 P # %XOTIC AND NATIVE SPECIES SHOW
POSITIVE LINEAR RELATIONSHIPS NATURALIZED SPECIES 2 P NATIVES 2
P $ATA SOURCES AS CITED IN &IG FOR PLANTS DATA FOR HUMAN POPULATION SIZE ARE FROM
#ALIFORNIA #OUNTIES ESTIMATES HTTPWWWDOFCAGOVHTML$EMOGRAPREPNDAT
HTM 53 3TATES ESTIMATES HTTPWWWCENSUSGOVMAINWWWCENHTML
AND -EXICAN STATES ESTIMATES HTTPWWWCITYPOPULATIONDE-EXICOHTML
WAYS TO DIFFERENCES IN AREA /N THE #HANNEL )SLANDS HOWEVER THE PATTERNS ARE
NOT IDENTICAL THE SLOPES OF THE RELATIONSHIPS ARE SIMILAR NATIVES Z EXOTICS
Z BUT THE SLOPE OF THE EXOTIC RELATIONSHIP IS SHALLOWER WHICH IS CONSISTENT
WITH #RAWLEYS CONCLUSION THAT BETA DIVERSITY IS REDUCED FOR EXOTIC SPECIES
RELATIVE TO NATIVES 4HIS SHALLOWER SLOPE IS ALSO CONSISTENT WITH THE EXPECTATION
THAT EXOTIC SPECIES ARE NOT FUNCTIONALLY AS DISPERSAL LIMITED AS NATIVES BECAUSE
STEEPER SLOPES ARE GENERALLY CONSIDERED TO BE INDICATIVE OF INCREASED INSULARITY AND
ISOLATION EG 2OSENZWEIG 2USSELL ET AL 4HIS CONCLUSION IS STRONGLY
SUPPORTED IN THIS CASE BY EXAMINING THE RELATIONSHIP THAT NATIVE AND EXOTIC
SPECIES HAVE RESPECTFULLY WITH ISOLATION BY PLOTTING THE RESIDUALS OF LOG AREA AND
LOG RICHNESS AGAINST DISTANCE OF EACH ISLAND TO THE MAINLAND IT IS CLEAR THAT THE
RICHNESS OF NATIVE SPECIES IS STRONGLY INmUENCED BY ISOLATION WHILE THE RICHNESS
OF EXOTIC SPECIES IS NOT &IG )N THIS CASE IT APPEARS THEREFORE THAT BOTH NATIVE
AND EXOTIC SPECIES ARE RESPONDING IN SIMILAR WAYS TO AREA BUT ARE DIFFERENTIALLY
AFFECTED BY ISOLATION
)N CONTRAST TO THIS PATTERN ON THE #HANNEL )SLANDS THE PATTERN ON OCEANIC
ISLANDS IS SOMEWHAT DIFFERENT /N OCEANIC ISLANDS BOTH THE SLOPES AND INTERCEPTS
OF THE SPECIES
AREA RELATIONSHIPS FOR NATIVE AND EXOTIC SPECIES ARE ALMOST IDENTICAL
NATIVES Z # EXOTICS Z # 4HIS RESULT IS A CONSE
QUENCE OF THE EXTREMELY TIGHT COUPLING BETWEEN NATIVE AND EXOTIC SPECIES RICHNESS
$& 3AX AND 3$ 'AINES
&IG 3PECIES AREA RELATIONSHIPS FOR NATIVE AND EXOTIC SPECIES OF PLANTS 3OLID CIRCLES ARE
NATIVE AND EMPTY CIRCLES ARE EXOTIC SPECIES OF VASCULAR PLANTS ! .ATIVES SHOW A POSITIVE
LINEAR RELATIONSHIP 2 Z # P WHILE EXOTICS DO NOT SHOW A
SIGNIlCANT RELATIONSHIP ALTHOUGH THE TREND IS NEVERTHELESS POSITIVE SEE !PPENDIX FOR
A DESCRIPTION OF DATA SOURCES AND METHODS " .ATIVES AND EXOTICS SHOW POSITIVE LINEAR
RELATIONSHIPS NATIVES 2 Z # P EXOTICS 2 Z
# P DATA FROM *UNAK ET AL # .ATIVES AND EXOTICS SHOW POSITIVE
LINEAR RELATIONSHIPS NATIVES 2 Z # P EXOTICS 2
Z # P DATA FROM 3AX ET AL
&IG )SOLATION AND THE RICHNESS OF PLANT SPECIES ON THE #HANNEL )SLANDS OF #ALIFORNIA
! 3IGNIlCANT LINEAR RELATIONSHIP BETWEEN THE DISTANCE OF ISLANDS TO THE MAINLAND AND
THE RESIDUALS OF LOG AREA AND LOG NUMBER OF NATIVE SPECIES 2 P "
4HE SAME RELATIONSHIP FOR EXOTIC SPECIES IS NOT SIGNIlCANT $ATA FROM *UNAK ET AL
3EE -OODY FOR A SIMILAR EXAMINATION OF THESE DATA
"IOGEOGRAPHIC PATTERNS OF NATURALIZED SPECIES
ON THESE ISLANDS LOG NATIVE SPECIES VS LOG EXOTIC SPECIES 2 P
#LEARLY BOTH NATIVES AND EXOTICS ARE RESPONDING TO AREA GIVEN THE HIGHLY SIG
NIlCANT RELATIONSHIP BETWEEN AREA AND RICHNESS BUT IN THIS CASE IT APPEARS THAT
OTHER FACTORS ARE ALSO CONTRIBUTING TO THE TIGHT COUPLING BETWEEN NATIVE AND EXOTIC
SPECIES RICHNESS !T THIS POINT THESE OTHER FACTORS ARE UNCLEAR BUT THIS PATTERN IS
INTRIGUING AND AT PRESENT MAY BE ONE OF THE MORE IMPORTANT UNEXPLAINED PATTERNS
IN SPECIES INVASIONS AND BIOGEOGRAPHY
3PECIES AREA RELATIONSHIPS FOR EXOTICS BUT NOT FOR NATIVES
&INALLY WE CONSIDER SCENARIOS IN WHICH EXOTIC SPECIES SHOW A SPECIES
AREA RELA
TIONSHIP AND NATIVE SPECIES DO NOT 7E ILLUSTRATE THIS HERE WITH FRESHWATER lSHES
IN WATERSHEDS OF TEMPERATE .ORTH !MERICA AND WATERSHEDS OF #ALIFORNIA &IG
)N BOTH THESE EXAMPLES EXOTIC SPECIES DElNED HERE AS BEING NON
NATIVE TO THE
WATERSHEDS IN QUESTION SHOW SIGNIlCANT POSITIVE LINEAR RELATIONSHIPS IN LOG
LOG SPACE WHILE NATIVE SPECIES DO NOT 7E BELIEVE THAT THERE ARE TWO PRINCIPAL
HYPOTHESES THAT CAN EXPLAIN THESE PATTERNS &IRST FRESHWATER lSHES ARE EXTREMELY
DISPERSAL LIMITED AND MAY HAVE HISTORICALLY BEEN UNABLE TO TAKE ADVANTAGE OF LARGE
AREAS BECAUSE OF EXTREME BARRIERS TO DISPERSAL )F THE LIKELIHOOD OF lSHES COLONIZING
A WATERSHED IS DRIVEN MORE BY THE NATURE AND SEVERITY OF THE DISPERSAL BARRIER THAN
BY THE SIZE OF THE WATERSHED THEN NATIVE RICHNESS MAY BE POORLY CORRELATED WITH
WATERSHED AREA $ISPERSAL BARRIERS HOWEVER SHOULD NOT BE AS IMPORTANT FOR EXOTIC
FRESHWATER lSHES BECAUSE IN MOST CASES THE SPREAD OF THESE SPECIES IS FACILITATED BY
DIRECT HUMAN INTRODUCTIONS ! SECOND POTENTIAL EXPLANATION FOR THE DIFFERENCE IN
NATIVE AND EXOTIC SPECIES
AREA RELATIONSHIPS OF FRESHWATER lSHES IN .ORTH !MERICA
AND #ALIFORNIA MAY BE DUE TO RECENT ANTHROPOGENIC CHANGES IN THE ENVIRONMENT
&OR EXAMPLE MANY OF THE LARGEST WATERSHEDS IN THE DATABASE OF 53 WATERSHEDS ARE
FOUND IN THE WESTERN 53 )N THE WESTERN 53 MANY DAMS HAVE BEEN BUILT AND LACUS
TRIAN ENVIRONMENTS CREATED IN REGIONS THAT HISTORICALLY HAD VERY FEW LAKES EG
-ARCHETTI ET AL 4HE CREATION OF THESE LAKES MAY HAVE INCREASED THE CAPAC
ITY OF THESE REGIONS TO SUPPORT A LARGER VARIETY OF FRESHWATER lSH SPECIES 4HIS PAT
TERN IS THEREFORE CONSISTENT WITH THE HYPOTHESIS THAT EXOTIC SPECIES ARE RESPONDING
TO CURRENT ENVIRONMENTAL CONDITIONS WHILE NATIVE SPECIES WHICH HAVE NOT HAD
SUFlCIENT TIME TO SPECIATE AND lLL THESE NEWLY AVAILABLE HABITATS ARE RESPONDING TO
CONDITIONS THAT WERE HISTORICALLY PRESENT )F THIS IS TRUE THEN IT WOULD SUGGEST THAT
EXOTIC SPECIES WHEN THEY ARE WIDELY INTRODUCED ACROSS POTENTIAL SUITABLE HABITATS
MAY BE A BETTER INDICATOR OF AN AREAS CAPACITY TO SUPPORT SPECIES THAN THE NATIVES
THEMSELVES WHICH MAY INSTEAD BE IN A DISPERSAL
LIMITED STATE OF DISEQUILIBRIUM
BETWEEN ENVIRONMENTAL CAPACITY AND SPECIES RICHNESS
#HANGES IN SPECIES AREA RELATIONSHIPS
/NE OF THE MOST PERTINENT ASPECTS OF STUDYING SPECIES
AREA RELATIONSHIPS IN TODAYS
CHANGING WORLD IS EXAMINING HOW SPECIES
AREA RELATIONSHIPS HAVE CHANGED WITH
$& 3AX AND 3$ 'AINES
&IG 3PECIES
AREA RELATIONSHIPS FOR NATIVE AND EXOTIC SPECIES OF FRESHWATER lSH
! .ATIVES DO NOT SHOW A SIGNIlCANT RELATIONSHIP DATA FROM 'IDO AND "ROWN
" %XOTIC SPECIES SHOW A POSITIVE LINEAR RELATIONSHIP 2 Z #
P DATA FROM 'IDO AND "ROWN # .ATIVES DO NOT SHOW A SIGNIlCANT
RELATIONSHIP DATA FROM -OYLE AREA ESTIMATES FOR THE WATERSHEDS USED IN THIS
ANALYSIS ARE FROM HTTPENDEAVORDESUCDAVISEDUNEWCARA $ %XOTIC SPECIES SHOW A
POSITIVE LINEAR RELATIONSHIP 2 Z # P DATA SOURCES AS
CITED ABOVE FOR PANEL #
THE ADDITION OF EXOTIC SPECIES AND THE EXTINCTION OF NATIVES 2OSENZWEIG
AND #OLLINS ET AL CONSIDER THIS ISSUE WITH THOUGHT EXPERIMENTS THAT VARY
THE NUMBER OF EXOTIC SPECIES EXCHANGED BETWEEN REGIONS TO PREDICT WHAT THE
EFFECTS OF EXOTIC SPECIES INTRODUCTIONS WILL BE FOR LOCAL AND GLOBAL SPECIES RICHNESS
"IOGEOGRAPHIC PATTERNS OF NATURALIZED SPECIES
(ERE WE EXPLORE THE CLOSELY RELATED ISSUE OF HOW SPECIES
AREA RELATIONSHIPS HAVE
ACTUALLY CHANGED OVERTIME USING EMPIRICAL DATA AND CONSIDER WHAT INSIGHTS
THESE CHANGES MAY PROVIDE
)N &IG WE HAVE PLOTTED THE TREND
LINES OF HISTORIC AND CURRENT SPECIES
AREA
RELATIONSHIPS OF PLANT SPECIES FOR SEVERAL REGIONS BY @HISTORIC WE MEAN THE NUMBER
OF NATIVE SPECIES BELIEVED TO HAVE BEEN PRESENT BEFORE ANY ANTHROPOGENIC EXTINC
TIONS OCCURRED AND BY @CURRENT WE MEAN THE NUMBER OF SPECIES PRESENT TODAY
WHICH IS THE SUM OF EXTANT NATIVE AND NATURALIZED SPECIES $ATA ON THE NUMBER
OF NATIVE EXTIRPATIONS IS AVAILABLE FOR ALL OF THE PLOTTED TREND LINES EXCEPT FOR mORAS
FROM MAINLAND SITES IN #ALIFORNIA (OWEVER GIVEN THE EXTREMELY LOW NUMBER OF
PLANT SPECIES THAT HAVE BEEN EXTIRPATED FROM COMPARABLE #HANNEL )SLAND SITES
MEAN OF FEWER THAN FOUR SPECIES PER ISLAND AND FROM THE OTHER PLANT DATA SETS
CONSIDERED HERE IT SEEMS LIKELY THAT THE NUMBER OF SPECIES LOST FROM THE #ALIFORNIA
MAINLAND SITES SHOULD ALSO BE RELATIVELY LOW AND SHOULD NOT SIGNIlCANTLY IMPACT
THE OBSERVED PATTERNS
!LL OF THE PATTERNS PLOTTED SHOW ONE OVERRIDING SIMILARITY IE AN INCREASE IN
THE NUMBER OF SPECIES PRESENT ACROSS SITES &IG 4HE MAGNITUDE OF THIS INCREASE
HOWEVER IS NOT IDENTICAL BETWEEN REGIONS BUT IS INSTEAD GENERALLY CONSISTENT
WITH THE DEGREE OF ISOLATION AND RELATIVE MEAN RICHNESS OF EACH OF THESE REGIONS
AVERAGE INCREASE IN RICHNESS ON OCEANIC ISLANDS #HANNEL )SLANDS
#ALIFORNIA MAINLAND SITES 53 3TATES AND #ALIFORNIA #OUNTIES
NOTE THAT #ALIFORNIA #OUNTIES AND 53 3TATES HAVE SIMILAR MEAN NUMBERS OF NATIVE
SPECIES AND RESPECTFULLY BUT THAT THE SPATIAL EXTENT AND EFFECTIVE ISO
LATION BETWEEN 53 3TATES IS MUCH HIGHER THAN THAT BETWEEN #ALIFORNIA #OUNTIES
4HESE INCREASES IN SPECIES RICHNESS PARTICULARLY THE POSITIVE CORRELATION BETWEEN
ISOLATION AND THE MAGNITUDE OF INCREASED SPECIES RICHNESS IS CONSISTENT WITH THE
CHANGES PREDICTED TO OCCUR BY #OLLINS ET AL AND ALSO WITH $ARWINS OBSER
VATION THAT HUMANS ARE lLLING UP ISOLATED REGIONS FAR MORE FULLY THAN NATURE
HAS DONE
!NOTHER PATTERN EVIDENT IN &IG IS THAT WITH THE ESTABLISHMENT OF NATURALIZED
SPECIES RELATIVELY LARGE AREAS ARE @CATCHING UP IN TOTAL PLANT RICHNESS WITH THOSE
REGIONS THAT HAVE HISTORICALLY BEEN LESS ISOLATED 4HUS THE LARGEST #HANNEL )SLANDS
NOW HAVE AS MANY PLANT SPECIES AS WERE HISTORICALLY FOUND ON EQUAL SIZED AREAS
OF THE #ALIFORNIA MAINLAND &IG 3IMILARLY LARGE OCEANIC ISLANDS NOW HAVE AS
MANY PLANT SPECIES AS WERE HISTORICALLY FOUND WITHIN 53 3TATES OF COMPARABLE
SIZE &IG 7HILE COMPARING THE #HANNEL )SLANDS WITH #ALIFORNIA MAINLAND
SITES SEEMS REASONABLE IT IS NOT CLEAR HOWEVER TO WHICH LANDMASSES THE LARGEST
OCEANIC ISLANDS SHOULD BE COMPARED 4O EXPLORE THIS FURTHER WE CONSIDER THE TWO
LARGEST OCEANIC ISLANDS IN OUR DATABASE (AWAII AND .EW :EALAND (AWAIIS CLOS
EST CONTINENTAL LAND MASS IS .ORTH !MERICA AND IN FACT (AWAIIS GREATEST NUMBER
OF NATURAL COLONISTS ARE BELIEVED TO HAVE COME FROM THE !MERICAN CONTINENTS
6 &UNK PERSONAL COMMUNICATION "ECAUSE (AWAII IS PRESENT AT HIGH TROPICAL
LATITUDES THE MOST REASONABLE COMPARISON IN .ORTH !MERICA WOULD SEEM TO BE
WITH EQUAL SIZED AREAS OF -EXICO #OMPARING BOTH THE HISTORIC AND CURRENT NUMBER
$& 3AX AND 3$ 'AINES
&IG #HANGE IN SPECIES
AREA RELATIONSHIPS FOR PLANTS 4HE DASHED LINES INDICATE
THE HISTORIC RELATIONSHIP BETWEEN AREA AND NATIVE SPECIES OF VASCULAR PLANTS 4HE SOLID LINES
INDICATE THE CURRENT RELATIONSHIP BETWEEN AREA AND ALL EXTANT SPECIES OF VASCULAR PLANTS
NATIVE AND EXOTIC )N ALL CASES THE INTERCEPT OF THESE SPECIES
AREA RELATIONSHIPS HAVE
SHIFTED UP AND IN SOME CASES THE SLOPE OF THESE RELATIONSHIPS HAVE BECOME MORE SHALLOW
OF PLANT SPECIES IN (AWAII WITH THE NUMBER OF NATIVE SPECIES PRESENT IN -EXICAN
3TATES IT IS CLEAR THAT THE HISTORIC mORA OF (AWAII WAS DEPAUPERATE COMPARED WITH
-EXICAN 3TATES (OWEVER THIS DISCREPANCY HAS NOW LARGELY DISAPPEARED #URRENT
PLANT RICHNESS IS RELATIVELY CLOSE TO WHAT WOULD BE EXPECTED IF (AWAII WERE A
PIECE OF MAINLAND OF .ORTH !MERICA &IG ! .EW :EALANDS CLOSEST CONTINENT
IS !USTRALIA AND .EW :EALANDS GREATEST NUMBER OF NATURAL COLONISTS HAVE COME
FROM !USTRALIA -C'LONE ET AL #OMPARING PLANT SPECIES IN .EW :EALAND
WITH THOSE IN !USTRALIAN 3TATES SHOWS THAT HISTORIC SPECIES RICHNESS OF PLANTS IN
.EW :EALAND WAS ONLY MINIMALLY LOWER THAN IN !USTRALIAN 3TATES OF COMPARABLE
SIZE AND THAT CURRENT PLANT RICHNESS IS NOW SOMEWHAT GREATER THAN WOULD BE
"IOGEOGRAPHIC PATTERNS OF NATURALIZED SPECIES
EXPECTED CONSIDERING THE ROLE OF AREA ALONE WERE .EW :EALAND PRESENT ON THE
MAINLAND OF !USTRALIA &IG " 4HE LONG
TERM IMPLICATIONS OF THESE CHANGES IN
RICHNESS ARE DIFlCULT TO ASCERTAIN AS ARE THE LONG
TERM TRAJECTORIES OF CONTINUING
CHANGES IN SPECIES RICHNESS ON THESE ISLANDS )T IS POSSIBLE THAT NET RICHNESS WILL
EVENTUALLY DECREASE IF THERE IS A LONG TIME LAG BEFORE SIGNIlCANT NUMBERS OF NATIVE
EXTINCTIONS OCCUR !LTERNATIVELY RICHNESS MAY CONTINUE TO INCREASE IF EXTINCTION
RATES REMAIN LOW AND ADDITIONAL SPECIES BECOME ESTABLISHED 2OSENZWEIG
#OLLINS ET AL 3AX ET AL 5NDOUBTEDLY ADDITIONAL STUDY WILL HELP TO
CLARIFY THIS ISSUE (OWEVER GIVEN THE EVIDENCE AVAILABLE FROM THE SPECIES
AREA
RELATIONSHIPS CONSIDERED HERE IT APPEARS THAT FURTHER EFFECTIVE DECREASES IN ISOLA
TION MEDIATED BY THE INTRODUCTION OF ADDITIONAL EXOTIC SPECIES SHOULD LEAD TO CON
TINUED INCREASES IN THE INTERCEPT OF SPECIES
AREA RELATIONSHIPS FOR ANY REGION AND
TO FURTHER DECREASES IN THE SLOPE OF THOSE RELATIONSHIPS IE TO CONTINUED INCREASES
IN RICHNESS FOR INDIVIDUAL AREAS
)T IS IMPORTANT TO STRESS THAT THE INCREASES IN RICHNESS DESCRIBED HERE SHOULD NOT
BE CONSTRUED AS @GOOD BUT INSTEAD AS EVIDENCE OF THE TYPES OF DIFFERENTIAL IMPACTS
THAT HUMANS ARE HAVING ON VARIOUS REGIONS AROUND THE WORLD )NDEED THESE DATA
&IG #HANGE IN SPECIES RICHNESS OF PLANTS ON (AWAII AND .EW :EALAND RELATIVE TO
SPECIES
AREA RELATIONSHIPS OF NEAREST CONTINENTS %MPTY CIRCLES REPRESENT HISTORIC NUMBER
OF NATIVE SPECIES ON CONTINENTS SOLID CIRCLES REPRESENT HISTORIC NUMBER OF NATIVE SPECIES ON
ISLANDS AND SOLID SQUARES REPRESENT THE CURRENT EXTANT NATIVE AND NATURALIZED NUMBER OF
SPECIES ON ISLANDS ! 4HE HISTORIC RICHNESS OF mOWERING PLANT SPECIES ON (AWAII WAS WELL
BELOW THE LINE OF THE SPECIES
AREA RELATIONSHIP FOR -EXICAN STATES BUT IS CURRENTLY VERY CLOSE
TO THIS LINE " 4HE HISTORIC RICHNESS OF VASCULAR PLANT SPECIES IN .EW :EALAND WAS CLOSE TO
BUT BELOW THE LINE OF THE SPECIES
AREA RELATIONSHIP FOR !USTRALIAN 3TATES BUT IS CURRENTLY
ABOVE THIS LINE
$& 3AX AND 3$ 'AINES
SUGGEST THAT THE MOST ISOLATED AND DISTINCTIVE BIOTAS OF THE WORLD ARE THE ONES
BEING MODIlED BY THE LARGEST DEGREE 3UCH MODIlCATIONS LEAD TO AT LEAST TWO TYPES
OF BIOTIC HOMOGENIZATION &IRST THEY LEAD TO HOMOGENIZATION OF SPECIES COMPOSI
TION AMONG REGIONS AS UNIQUE SPECIES ARE LOST AND COMMON ONES ARE GAINED EG
,OCKWOOD AND -C+INNEY 3ECOND THEY LEAD TO HOMOGENIZATION OF SPECIES
RICHNESS AMONG REGIONS IN WHICH VARIATION IN SPECIES RICHNESS AND SPECIES
AREA
RELATIONSHIPS IS REDUCED 4HE LONG
TERM IMPLICATIONS OF BIOTIC HOMOGENIZATION ARE
STILL UNKNOWN BUT IT IS CLEAR THAT THE LOSS OF DISTINCTIVE BIOTAS AND THE MODIlCATION
OF PATTERNS OF BIODIVERSITY ACROSS THE GLOBE ARE TOPICS WORTHY OF MUCH ADDITIONAL
STUDY
#/.#,5$).' 4(/5'(43
7E HAVE ATTEMPTED TO REVIEW AND PRESENT SOME OF THE BASIC PATTERNS OF BIOGEO
GRAPHIC DISTRIBUTION OF EXOTIC SPECIES 'IVEN THE LIMITED SCOPE OF A SINGLE BOOK
CHAPTER WE WERE NOT ABLE TO REVIEW ALL OF THE MANY PATTERNS THAT HAVE BEEN
DESCRIBED TO DATE BUT INSTEAD HAVE FOCUSED ON THOSE PATTERNS THAT WE DEEMED TO
BE OF THE GREATEST INTEREST TO ECOLOGISTS (AD WE INSTEAD CHOSEN TO EXPLORE THOSE
ISSUES OF GREATEST INTEREST TO EVOLUTIONARY BIOLOGISTS WE WOULD UNDOUBTEDLY HAVE
CHOSEN A SLIGHTLY DIFFERENT SET OF TOPICS TO REVIEW &OR EXAMPLE ONE PATTERN THAT
IS EXTREMELY INTRIGUING WITH NATURALIZED SPECIES IS THE RAPID EVOLUTION OF CLINES IN
BODY SIZE ACROSS LATITUDE THAT QUANTITATIVELY MATCH BUT QUALITATIVELY DIFFER FROM
THOSE SEEN IN THE NATIVE RANGE SUGGESTING THAT EVOLUTION CAN SIMULTANEOUSLY
OPERATE IN A DETERMINISTIC AND CONTINGENT MANNER (UEY ET AL 'ILCHRIST
ET AL 3TILL WE HOPE THAT THE WORK DESCRIBED HERE PROVIDES A GOOD INROAD TO THE
GROWING LITERATURE ON THIS TOPIC A LITERATURE THAT PROMISES TO PROVIDE RECIPROCAL
INSIGHTS TO ECOLOGICAL AND BIOGEOGRAPHICAL THEORY AS WELL AS TO OUR APPLIED UNDER
STANDING OF INVASION BIOLOGY &INALLY WE BELIEVE THAT IF WE AS A SOCIETY ARE GOING
TO SUCCESSFULLY LEARN TO UNDERSTAND AND MITIGATE THE ENVIRONMENTAL THREATS POSED
BY ANTHROPOGENIC CHANGES TO THE ENVIRONMENT THEN WE MUST EMPLOY ALL AVAILABLE
EVIDENCE TO DO SO UNDOUBTEDLY STUDIES OF SPECIES INVASION AT BIOGEOGRAPHIC SCALES
CAN DO MUCH TO HELP ACHIEVE THESE GOALS
!#+./7,%$'%-%.43
7E WOULD LIKE TO THANK THE EDITORS OF THIS BOOK FOR GIVING US THE OPPORTUNITY TO
WRITE THIS CHAPTER -ANY OF THE IDEAS IN THIS MS BENElTED FROM CONVERSATIONS WITH
* "ROWN 4HE QUALITY OF THIS MS HAS BENElTED SIGNIlCANTLY FROM COMMENTS BY $
0ERAULT - "EALS AND - ,OMOLINO 4HIS IS CONTRIBUTION NUMBER FROM 0)3#/
THE 0ARTNERSHIP FOR )NTERDISCIPLINARY 3TUDIES OF #OASTAL /CEANS ! ,ONG
TERM
%COLOGICAL #ONSORTIUM FUNDED BY THE $AVID AND ,UCILE 0ACKARD &OUNDATION
"IOGEOGRAPHIC PATTERNS OF NATURALIZED SPECIES
2%&%2%.#%3
!RRHENIUS / 3PECIES AND AREA *OURNAL OF %COLOGY
!TKINSON # 4 2 * $USEK + , 7OODS AND 7 - )KO 0ATHOGENICITY OF AVIAN
MALARIA IN EXPERIMENTALLY INFECTED (AWAII !MAKIHI *OURNAL OF 7ILDLIFE $ISEASES
"ANKS $ , ! VASCULAR mORA OF THE !GUA 4IBIA -OUNTAINS SOUTHERN #ALIFORNIA
2ANCHO 3ANTA !NA "OTANICAL 'ARDEN /CCASIONAL 0UBLICATION NUMBER #LAREMONT
#!
"ICKFORD # AND 0 2ICH 6EGETATION AND mORA OF THE ,ANDELS (ILL "IG #REEK 2ESERVE
-ONTEREY #OUNTY #ALIFORNIA 3ECOND %DITION 0UBLICATION NUMBER %NVIRONMENTAL
&IELD 0ROGRAM 5NIVERSITY OF #ALIFORNIA 3ANTA #RUZ #!
"LACKBURN 4 - AND + * 'ASTON 3PATIAL PATTERNS IN THE SPECIES RICHNESS OF BIRDS IN
THE .EW 7ORLD %COGRAPHY
"OYD 3 $ ! mORA OF THE 'AVILAN (ILLS 7ESTERN 2IVERSIDE #OUNTY #ALIFORNIA
-ASTERS OF 3CIENCE 4HESIS 5NIVERSITY OF #ALIFORNIA 2IVERSIDE #!
"ROWN * ( -ACROECOLOGY 5NIVERSITY OF #HICAGO 0RESS #HICAGO ),
"ROWN * ( AND - 6 ,OMOLINO "IOGEOGRAPHY 3ECOND %DITION 3INAUER !SSOCIATES
3UNDERLAND -!
"ROWN * ( AND $ & 3AX 'RADIENTS IN SPECIES DIVERSITY WHY ARE THERE SO MANY
SPECIES IN THE TROPICS 0AGES IN - 6 ,OMOLINO $ & 3AX AND * ( "ROWN
EDITORS &OUNDATIONS OF "IOGEOGRAPHY CLASSIC PAPERS WITH COMMENTARIES 5NIVERSITY OF
#HICAGO 0RESS #HICAGO ),
"RUNO * & * * 3TACHOWICZ AND - $ "ERTNESS )NCLUSION OF FACILITATION INTO ECOLOGI
CAL THEORY 4RENDS IN %COLOGY AND %VOLUTION
#ASE 4 * )NVASION RESISTANCE SPECIES BUILD UP AND COMMUNITY COLLAPSE IN MODEL
COMPETITION COMMUNITIES 0AGES IN - % 'ILPIN AND ) (ANSKI EDITORS
-ETAPOPULATION DYNAMICS !CADEMIC 0RESS ,ONDON 5+
#HOWN 3 , . * - 'REMMEN AND + * 'ASTON %COLOGICAL BIOGEOGRAPHY OF
SOUTHERN OCEAN ISLANDS SPECIES AREA RELATIONSHIPS HUMAN IMPACTS AND CONSERVATION
!MERICAN .ATURALIST
#OLLINS - $ $ 0 6AZQUEZ AND . * 3ANDERS 3PECIES AREA CURVES HOMOGENIZATION
AND THE LOSS OF GLOBAL DIVERSITY %VOLUTIONARY %COLOGY 2ESEARCH
#ONNOR % & AND % $ -C#OY 4HE STATISTICS AND BIOLOGY OF THE SPECIES AREA RELATION
SHIP !MERICAN .ATURALIST
#RAWLEY - * 7HAT MAKES A COMMUNITY INVASIBLE 0AGES IN ! * 'RAY
- * #RAWLEY AND 0 * %DWARDS EDITORS #OLONIZATION SUCCESSION AND STABILITY "LACKWELL
3CIENTIlC 0UBLICATIONS /XFORD 5+
$ARWIN # /N THE ORIGIN OF SPECIES -URRAY ,ONDON 5+
$AVIS - ! "IOTIC GLOBALIZATION DOES COMPETITION FROM INTRODUCED SPECIES THREATEN
BIODIVERSITY "IOSCIENCE
$E .EVERS ' 0EPPERWOOD &LORA 0EPPERWOOD 2ANCH .ATURAL 0RESERVE 3ONOMA
#OUNTY #ALIFORNIA #ALIFORNIA !CADEMY OF 3CIENCES 3AN &RANCISCO #!
$& 3AX AND 3$ 'AINES
$EUTSCHEWITZ + ! ,AUSCH ) +UHN AND 3 +LOTZ .ATIVE AND ALIEN PLANT SPECIES RICHNESS
IN RELATION TO SPATIAL HETEROGENEITY ON A REGIONAL SCALE IN 'ERMANY 'LOBAL %COLOGY AND
"IOGEOGRAPHY
$OBZHANSKY 4 %VOLUTION IN THE TROPICS !MERICAN 3CIENTIST
$UNCAN 2 0 4 - "LACKBURN AND $ 3OL 4HE ECOLOGY OF BIRD INTRODUCTIONS
!NNUAL 2EVIEW OF %COLOGY %VOLUTION AND 3YSTEMATICS
%BENHARD 4 )NTRODUCED BIRDS AND MAMMALS AND THEIR ECOLOGICAL EFFECTS 3WEDISH
7ILDLIFE 2ESEARCH
%LTON # 3 4HE ECOLOGY OF INVASIONS BY ANIMALS AND PLANTS -ETHUEN AND #O ,4$
,ONDON 5+
&INE 0 6 ! 4HE INVASIBILITY OF TROPICAL FORESTS BY EXOTIC PLANTS *OURNAL OF 4ROPICAL
%COLOGY
&ISCHER ! ' ,ATITUDINAL VARIATIONS IN ORGANIC DIVERSITY %VOLUTION
&RANCE 2 4HE .ORTH !MERICAN ,ATITUDINAL GRADIENT IN SPECIES RICHNESS AND
GEOGRAPHIC RANGE OF FRESHWATER CRAYlSH AND AMPHIPODS !MERICAN .ATURALIST
&RIDLEY * $ 2 , "ROWN AND * & "RUNO .ULL MODELS OF EXOTIC INVASION AND SCALE
DEPENDENT PATTERNS OF NATIVE AND EXOTIC SPECIES RICHNESS %COLOGY IN PRESS
&RITTS 4 ( AND ' ( 2ODDA 4HE ROLE OF INTRODUCED SPECIES IN THE DEGRADATION OF
ISLAND ECOSYSTEMS A CASE HISTORY OF 'UAM !NNUAL 2EVIEW OF %COLOGY %VOLUTION AND
3YSTEMATICS
'AINES 3 $ AND * ,UBCHENCO ! UNIlED APPROACH TO MARINE PLANT HERBIVORE INTER
ACTIONS "IOGEOGRAPHY !NNUAL 2EVIEW OF %COLOGY AND 3YSTEMATICS
'ASTON + * 4 - "LACKBURN AND * ) 3PICER 2APOPORTS RULE TIME FOR AN EPITAPH
4RENDS IN %COLOGY AND %VOLUTION
'IDO + " AND * ( "ROWN )NVASION OF .ORTH !MERICAN DRAINAGES BY ALIEN lSH SPE
CIES &RESHWATER "IOLOGY
'ILCHRIST ' 7 2 " (UEY * "ALANYA - 0ASCUAL AND , 3ERRA ! TIME SERIES OF
EVOLUTION IN ACTION A LATITUDINAL CLINE IN WING SIZE IN 3OUTH !MERICAN $ROSOPHILA SUB
OBSCURA %VOLUTION
'ROOMBRIDGE " ED 'LOBAL BIODIVERSITY STATUS OF THE %ARTHS LIVING RESOURCES
#HAPMAN (ALL ,ONDON 5+
(AWKINS " ! %COLOGYS OLDEST PATTERN 4RENDS IN %COLOGY AND %VOLUTION
(UEY 2 " ' 7 'ILCHRIST - , #ARLSON $ "ERRIGAN AND , 3ERRA 2APID EVOLUTION
OF A GEOGRAPHIC CLINE IN SIZE IN AN INTRODUCED mY 3CIENCE
(UTTON ) "IRDS OF ,ORD (OWE )SLAND PAST AND PRESENT ,ITHOCRAFT 'RAPHICS
-ELBOURNE !USTRALIA
*UNAK 3 4 !YERS AND 2 3COTT ! mORA OF 3ANTA #RUZ )SLAND 3ANTA "ARBARA "OTANIC
'ARDENS 3ANTA "ARBARA #!
+ENNEDY 4 ! 3 .AEEM + - (OWE * - ( +NOPS $ 4ILMAN AND 0 2EICH
"IODIVERSITY AS A BARRIER TO ECOLOGICAL INVASION .ATURE
,ATHROP % 7 AND 2 & 4HORNE ! mORA OF THE 3ANTA 2OSA 0LATEAU SOUTHERN #ALIFORNIA
3OUTHERN #ALIFORNIA "OTANISTS 3PECIAL 0UBLICATION NUMBER #LAREMONT #!
"IOGEOGRAPHIC PATTERNS OF NATURALIZED SPECIES
,ETCHER ! * AND 0 ( (ARVEY 6ARIATION IN GEOGRAPHIC RANGE SIZE AMONG MAMMALS
OF THE 0ALEARCTIC !MERICAN .ATURALIST
,EVINE * - 3PECIES DIVERSITY AND BIOLOGICAL INVASIONS RELATING LOCAL PROCESS TO COM
MUNITY PATTERN 3CIENCE
,ONSDALE 7 - 'LOBAL PATTERNS OF PLANT INVASIONS AND THE CONCEPT OF INVASIBILITY
%COLOGY
,OMOLINO - 6 )NTERPRETATIONS AND COMPARISONS OF CONSTANTS IN THE SPECIES
AREA
RELATIONSHIP AN ADDITIONAL CAUTION !MERICAN .ATURALIST
,OMOLINO - 6 4HE SPECIES
AREA RELATIONSHIP NEW CHALLENGES FOR AN OLD PATTERN
0ROGRESS IN 0HYSICAL 'EOGRAPHY
,OMOLINO - 6 $ & 3AX AND * ( "ROWN &OUNDATIONS OF "IOGEOGRAPHY CLASSIC
PAPERS WITH COMMENTARIES 5NIVERSITY OF #HICAGO 0RESS #HICAGO ),
,OMOLINO - 6 AND - $ 7EISER 4OWARDS A MORE GENERAL SPECIES
AREA RELATION
SHIP DIVERSITY ON ALL ISLANDS GREAT AND SMALL *OURNAL OF "IOGEOGRAPHY
,UNING + 3EAWEEDS THEIR ENVIRONMENT BIOGEOGRAPHY AND ECOPHYSIOLOGY *OHN
7ILEY AND 3ONS )NC .EW 9ORK .9
-AC!RTHUR 2 ( AND % / 7ILSON 4HE 4HEORY OF )SLAND "IOGEOGRAPHY 0RINCETON
5NIVERSITY 0RESS 0RINCETON .*
-AC!RTHUR 2 ( 'EOGRAPHIC ECOLOGY PATTERNS IN THE DISTRIBUTION OF SPECIES (ARPER
2OW .EW 9ORK .9
-ARCHETTI - 0 4 ,IGHT * &ELICIANO 4 !RMSTRONG : (OGAN * 6IERS AND 0 " -OYLE
(OMOGENIZATION OF #ALIFORNIAS lSH FAUNA THROUGH ABIOTIC CHANGE 0AGES
IN * , ,OCKWOOD AND - , -C+INNEY EDITORS "IOTIC (OMOGENIZATION
+LUWER !CADEMIC0LENUM 0UBLISHERS .EW 9ORK .9
-ARTIN 4 % 3PECIES
AREA SLOPES AND COEFlCIENTS A CAUTION ON THEIR INTERPRETATION
!MERICAN .ATURALIST
-ATTER 3 & ) (ANSKI AND - 'YLLENBERG ! TEST OF THE METAPOPULATION MODEL OF THE
SPECIES
AREA RELATIONSHIP *OURNAL OF "IOGEOGRAPHY
-C#LINTOCK % 7 +NIGHT AND . &AHY ! mORA OF THE 3AN "RUNO -OUNTAINS 3AN
-ATEO #OUNTY #ALIFORNIA 0ROCEEDINGS OF THE #ALIFORNIA !CADEMY OF 3CIENCES FOURTH
SERIES
-C'LONE - 3 2 0 $UNCAN AND 0 " (EENAN %NDEMISM SPECIES SELECTION
AND THE ORIGIN AND DISTRIBUTION OF THE VASCULAR PLANT mORA OF .EW :EALAND *OURNAL OF
"IOGEOGRAPHY
-C+INNEY - , A )NmUENCE OF SETTLEMENT TIME HUMAN POPULATION PARK SHAPE AND
AGE VISITATION AND ROADS ON THE NUMBER OF ALIEN PLANT SPECIES IN PROTECTED AREAS IN THE
53! $IVERSITY AND $ISTRIBUTIONS
-C+INNEY - , B $O HUMAN ACTIVITIES RAISE SPECIES RICHNESS #ONTRASTING PATTERNS
IN 5NITED 3TATES PLANTS AND lSHES 'LOBAL %COLOGY AND "IOGEOGRAPHY
-C+INNEY - , AND ,OCKWOOD * , "IOTIC HOMOGENIZATION A FEW WINNERS REPLAC
ING MANY LOSERS IN THE NEXT MASS EXTINCTION 4RENDS IN %COLOGY AND %VOLUTION
-OODY ! !NALYSIS OF PLANT SPECIES DIVERSITY WITH RESPECT TO ISLAND CHARACTERISTICS ON
THE #HANNEL )SLANDS #ALIFORNIA *OURNAL OF "IOGEOGRAPHY
$& 3AX AND 3$ 'AINES
-OYLE 0 " )NLAND lSHES OF #ALIFORNIA REVISED AND EXPANDED EDITION 5NIVERSITY OF
#ALIFORNIA 0RESS "ERKELEY #!
.AEEM 3 * - ( +NOPS $ 4ILMAN + - (OWE 4 +ENNEDY AND 3 'ALE 0LANT
DIVERSITY INCREASES RESISTANCE TO INVASION IN THE ABSENCE OF COVARYING EXTRINSIC FACTORS
/IKOS
0IANKA % 2 ,ATITUDINAL GRADIENTS IN SPECIES DIVERSITY A REVIEW OF CONCEPTS
!MERICAN .ATURALIST
0YSEK 0 !LIEN AND NATIVE SPECIES IN CENTRAL %UROPEAN URBAN mORAS A QUANTITATIVE
COMPARISON *OURNAL OF "IOGEOGRAPHY
0YSEK 0 6 *AROSIK AND 4 +UCERA 0ATTERNS OF INVASION IN TEMPERATE NATURE
RESERVES "IOLOGICAL #ONSERVATION
0YSEK 0 4 +UCERA AND 6 *AROSIK 0LANT SPECIES RICHNESS OF NATURE RESERVES THE
INTERPLAY OF AREA CLIMATE AND HABITAT IN A CENTRAL %UROPEAN LANDSCAPE *OURNAL OF
"IOGEOGRAPHY
2APOPORT % ( !REOGRAPHY GEOGRAPHIC STRATEGIES OF SPECIES TRANSLATION BY
" $RAUSEL 0ERGAMON /XFORD 5+
2AVEN 0 ( ( * 4HOMPSON AND " ! 0RIGGE &LORA OF THE 3ANTA -ONICA -OUNTAINS
#ALIFORNIA 3ECOND %DITION 3OUTHERN #ALIFORNIA "OTANISTS 3PECIAL 0UBLICATION .O
2EJMANEK - 3PECIES RICHNESS AND RESISTANCE TO INVASIONS 0AGES IN
' ( /RIANS 2 $IRZO * ( #USHMAN EDITORS "IODIVERSITY AND ECOSYSTEM PROCESSES IN
TROPICAL FORESTS 3PRINGER 6ERLAG "ERLIN 'ERMANY
2EJMANEK - 4HE RICH GET RICHER RESPONSES &RONTIERS IN %COLOGY AND THE
%NVIRONMENT
2ICHARDSON $ - . !LLSOPP # - $!NTONIO 3 * -ILTON AND - 2EJMANEK 0LANT
INVASIONS THE ROLE OF MUTUALISMS "IOLOGICAL 2EVIEWS
2ODRIGUERO - 3 AND $ % 'ORLA ,ATITUDINAL GRADIENT IN SPECIES RICHNESS OF THE .EW
7ORLD 4RIATOMINAE 2EDUVIIDAE 'LOBAL %COLOGY AND "IOGEOGRAPHY
2OHDE + ,ATITUDINAL GRADIENTS IN SPECIES DIVERSITY THE SEARCH FOR THE PRIMARY
CAUSE /IKOS
2OSENZWEIG - , 3PECIES DIVERSITY IN SPACE AND TIME #AMBRIDGE 5NIVERSITY 0RESS
#AMBRIDGE 5+
2OSENZWEIG - , 4HE FOUR QUESTIONS 7HAT DOES THE INTRODUCTION OF EXOTIC SPECIES
DO TO DIVERSITY %VOLUTIONARY %COLOGY 2ESEARCH
2USSELL * # - . #LOUT AND " ( -C!RDLE )SLAND BIOGEOGRAPHY AND THE SPE
CIES RICHNESS OF INTRODUCED MAMMALS OF THE .EW :EALAND OFFSHORE ISLANDS *OURNAL OF
"IOGEOGRAPHY
3AX $ & ,ATITUDINAL GRADIENTS AND GEOGRAPHIC RANGES OF EXOTIC SPECIES IMPLICATIONS
FOR BIOGEOGRAPHY *OURNAL OF "IOGEOGRAPHY
3AX $ & 3 $ 'AINES AND * ( "ROWN 3PECIES INVASIONS EXCEED EXTINCTIONS ON
ISLANDS WORLDWIDE A COMPARATIVE STUDY OF PLANTS AND BIRDS !MERICAN .ATURALIST
3COTT * - 3 #ONANT AND # VAN 2IPER ))) EDITORS %VOLUTION ECOLOGY CONSERVA
TION AND MANAGEMENT OF (AWAIIAN BIRDS A VANISHING AVIFAUNA #OOPER /RNITHOLOGICAL
3OCIETY #AMARILLO #!
"IOGEOGRAPHIC PATTERNS OF NATURALIZED SPECIES
3IMPSON ' ' 3PECIES DENSITY OF .ORTH !MERICAN 2ECENT MAMMALS 3YSTEMATIC
:OOLOGY n
3TACHOWICZ * * 2 " 7HITLATCH AND 2 7 /SMAN 3PECIES DIVERSITY AND INVASION
RESISTANCE IN A MARINE ECOSYSTEM 3CIENCE
3TADLER * ! 4REFmICH 3 +LOTZ AND 2 "RANDL %XOTIC PLANT SPECIES INVADE DIVERSITY
HOT SPOTS THE NATURALIZED mORA OF NORTHWESTERN +ENYA %COGRAPHY
3TEVENS ' # 4HE LATITUDINAL GRADIENT IN GEOGRAPHIC RANGE HOW SO MANY SPECIES
COEXIST IN THE TROPICS !MERICAN .ATURALIST
3TEVENS ' # AND " * %NQUIST -ACROECOLOGICAL LIMITS TO THE ABUNDANCE AND
DISTRIBUTION OF 0INUS 0AGES
IN $ - 2ICHARDSON EDITOR %COLOGY AND BIOGEOG
RAPHY OF 0INUS #AMBRIDGE 5NIVERSITY 0RESS #AMBRIDGE 5+
3TOHLGREN 4 * $ "INKLEY ' 7 #HONG - ! +ALKHAN , $ 3CHELL + ! "ULL 9 /TSUKI
' .EWMAN - "ASHKIN AND 9 3ON %XOTIC PLANT SPECIES INVADE HOT SPOTS OF
NATIVE PLANT DIVERSITY %COLOGICAL -ONOGRAPHS
3TOHLGREN 4 * $ 4 "ARNETT AND * 4 +ARTESZ 4HE RICH GET RICHER PATTERNS OF PLANT
INVASIONS IN THE 5NITED 3TATES &RONTIERS IN %COLOGY AND THE %NVIRONMENT
4ILMAN $ #OMMUNITY INVASIBILITY RECRUITMENT LIMITATION AND GRASSLAND BIODIVER
SITY %COLOGY
6ILLASEOR * , AND & * %SPINOSA
'ARCIA 4HE ALIEN mOWERING PLANTS OF -EXICO
$IVERSITY AND $ISTRIBUTIONS
7ARDLE $ ! %XPERIMENTAL DEMONSTRATION THAT PLANT DIVERSITY REDUCES INVASIBILITY
EVIDENCE OF A BIOLOGICAL MECHANISM OR A CONSEQUENCE OF SAMPLING EFFECT /IKOS
7ILLIG - 2 $ - +AUFMAN AND 2 $ 3TEVENS ,ATITUDINAL GRADIENTS OF BIODIVER
SITY PATTERN PROCESS SCALE AND SYNTHESIS !NNUAL 2EVIEW IN %COLOGY %VOLUTION AND
3YSTEMATICS
7ILLOUGHBY * 7 ! mORA OF THE 6ACA -OUNTAINS #ALIFORNIA -ASTERS 4HESIS
#ALIFORNIA 3TATE 5NIVERSITY 3ACRAMENTO #!
7ILSON + %XTINCT AND INTRODUCED VERTEBRATE SPECIES IN .EW :EALAND A LOSS OF BIODIS
TINCTIVENESS AND GAIN IN BIODIVERSITY 0ACIlC #ONSERVATION "IOLOGY
$& 3AX AND 3$ 'AINES
!00%.$)8 $%4!),%$ -%4(/$3
0LANTS IN 53 3TATES
4HE NUMBER OF NATIVE AND NATURALIZED SPECIES OF VASCULAR PLANTS FOR EACH
STATE ARE FROM -AC ET AL AND 3TEIN ET AL $ATA ON NUMBER
OF EXTINCT OR EXTIRPATED PLANTS FOR EACH STATE WERE COLLECTED FROM REPORTS
OF PRIVATE AND PUBLIC STATE AGENCIES PREDOMINATELY THE .ATURAL (ERITAGE
0ROGRAMS OF INDIVIDUAL STATES THE DATA USED IN THESE ANALYSES ARE AVAIL
ABLE UPON REQUEST FROM THE AUTHORS )N EACH CASE THE NUMBER OF SPECIES
LISTED AS h38v EXTINCT OR EXTIRPATED FROM A STATE OR h3(v PRESENT HISTORI
CALLY IN A STATE BUT NO LONGER KNOWN TO OCCUR WERE SUMMED TO PROVIDE
THE TOTAL NUMBER OF SPECIES THAT HAVE BEEN EXTIRPATED #OMPLETE DATA FOR
-ASSACHUSETS -INNESOTA AND .ORTH $AKOTA WERE UNAVAILABLE SO THESE
STATES WERE EXCLUDED FROM THESE ANALYSES !LASKA AND (AWAII WERE ALSO
EXCLUDED FROM THESE ANALYSES
0LANTS IN MAINLAND AREAS OF #ALIFORNIA
!VAILABLE mORAS OF PARTICULAR SITES ON THE MAINLAND OF #ALIFORNIA WERE
SELECTED TO REPRESENT THE SPAN OF AREAS PRESENT ON THE #ALIFORNIA #HANNEL
)SLANDS )N TOTAL NINE SITES WERE USED 4HESE INCLUDED THE mORAS OF THE
3ANTA -ONICA -OUNTAINS 2AVEN ET AL 3AN "RUNO -OUNTAINS
-C#LINTOCK ET AL 6ACA -OUNTAINS 7ILLOUGHBY !GUA
4IBIA -OUNTAINS "ANKS 3ANTA 2OSA 0LATEAU ,ATHROP AND 4HORNE
'AVIAN (ILLS "OYD 0ESCADERO -ARSH 7 !NDERSON AND
2 -ORGAN UNPUBLISHED MANUSCRIPT ,ANDELS
(ILL "IG #REEK 2ESERVE
"ICKFORD AND 2ICH AND 0EPERWOOD 2ESERVE $E .EVERS
$ATA ON THE NUMBER OF NATIVE AND NON
NATIVE SPECIES ARE FROM THESE
SOURCES .O DATA WAS AVAILABLE ON NUMBER OF EXTIRPATED SPECIES
#HAPTER TWENTY
ONE
,INKING SCALE DEPENDENT
PROCESSES IN INVASIONS
- 7 #ADOTTE 4 &UKAMI AND 3 - -C-AHON
).42/$5#4)/.
4HE OVERARCHING GOAL OF THIS VOLUME WAS TO EXAMINE HOW CONCEPTUAL ECOLOGY
INFORMS OUR UNDERSTANDING OF SPECIES INVASIONS WHILE SIMULTANEOUSLY VIEWING
SPECIES INVASIONS AS POTENTIAL TESTS OF ECOLOGICAL THEORY SEE #HAPTER !S ONE HALF
THE PURPOSE OF THE PRESENT VOLUME ATTESTS TO ECOLOGICAL THEORY IS USEFUL FOR CON
TEXTUALIZING WHEN AND HOW SPECIES INVADE EG 3HEA AND #HESSON -ORE
THAN THIS AND AS THE SECOND HALF OF THIS VOLUMES PURPOSE SPECIES INVASIONS ALLOW
US TO SEE HYPOTHESIZED DYNAMICS AND TO TEST MAJOR IDEAS IN ECOLOGY AND EVOLUTION
EG ,ODGE 3AX ET AL 4HROUGHOUT THIS VOLUME AUTHORS HAVE EXPLIC
ITLY EXPLORED THE ADVANCES IN KNOWLEDGE GAINED BY COMBINING CONCEPTUAL ECOLOGY
WITH INVASION ECOLOGY
(OWEVER BIOLOGICAL INVASIONS HAVE BEEN STUDIED FOR THEIR OWN SAKE AS THEY ARE
CONSIDERED TO BE A MAJOR CONSERVATION RISK 6ITOUSEK ET AL AND AN IMPOR
TANT COMPONENT OF INVASIONS STUDIES IS THE STUDY OF HOW NON
INDIGENOUS SPECIES
ENTER AND PROLIFERATE IN NEW COMMUNITIES ! MULTITUDE OF HYPOTHESES HAVE BEEN
CREATED TO EXPLAIN HOW SPECIES INVADE NEW HABITATS )NDERJIT ET AL 9ET
THESE HYPOTHESES CAN BE CONSIDERED AS SPECIAL CASES OF MORE GENERAL ECOLOGICAL
PHENOMENA OR AS AD HOC EXPLANATIONS lLLING THE VOID WHERE WE LACK A GENERAL
THEORY EG ,AKATOS 4HOUGH THE PRIMARY GOAL OF THIS VOLUME HAS BEEN TO
COMBINE CONCEPTUAL ECOLOGY AND INVASION BIOLOGY MANY OF THE AUTHORS EXAMINED
-7 #ADOTTE ET AL (EDS) #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY
3PRINGER 0RINTED IN THE .ETHERLANDS
-7 #ADOTTE 4 &UKAMI AND 3- -C-AHON
ECOLOGICAL THEORY OR PARTICULAR INVASIONS FROM DIFFERENT LEVELS OF ORGANIZATION 7E
ORGANIZED THE CHAPTERS INTO THREE GENERAL SCALES OF ORGANIZATION POPULATION COM
MUNITY AND BIOGEOGRAPHIC SCALES
)NSTEAD OF VIEWING THESE SCALES AS NECESSARILY DISCRETE OR HIERARCHICALLY RELATED
IT MAY BE USEFUL TO VIEW THESE PROCESSES AS A CONTINUUM FEEDING BACK INTO ONE
ANOTHER &IG )N ORDER TO ADEQUATELY DESCRIBE AND UNDERSTAND INVASIONS WE
MUST LOOK TO PROCESSES ALLOWING SPECIES TO OVERCOME I BIOGEOGRAPHICAL BARRIERS IN
THE MOVEMENT OF PROPAGULES II ABIOTIC BARRIERS IN ESTABLISHING NEW POPULATIONS
AND III BIOTIC BARRIERS IN MAINTAINING HIGHER BIRTH THAN DEATH RATES AND BEING
ABLE TO INCREASE NUMBERS AND SPREAD TO NEW LOCALES 4HESE ARE ESSENTIALLY lLTERS
SUBSAMPLING POTENTIAL COLONISTS FROM LARGER SPECIES POOLS "ELYEA AND ,ANCASTER
4HE DISPERSAL lLTER LIMITS SPECIES THAT ARE UNLIKELY TO MIGRATE TO THE LOCAL
COMMUNITY WHILE THOSE WITH CERTAIN CHARACTERISTICS EG LONG
DISTANCE DISPERSAL
OR COMMON AGRICULTURAL SEED CONTAMINANTS WILL HAVE A HIGHER PROBABILITY OF
COLONIZING AND ABIOTIC lLTERS SELECT CERTAIN SPECIES OR TRAITS AS THOSE WITH A HIGHER
CHANGE OF SURVIVING IN THE NEW HABITAT "ELYEA AND ,ANCASTER
/BSERVATIONS MADE AT DIFFERENT SCALES OFTEN REQUIRE MECHANISTIC LINKS TO OTHER
SCALES OF ORGANIZATION &OR EXAMPLE EXTINCTIONS ARE ULTIMATELY OBSERVED AT A
BIOGEOGRAPHIC SCALE BUT THE MECHANISMS LEADING TO EXTINCTION MUST BE OBSERVED
AT THE POPULATION LEVEL EG WHAT ELEVATES DEATH RATES OR LOWER BIRTH RATES
&IG 4HE INVASION PROCESS AS A CONTINUUM ACROSS SPATIAL SCALES 4HE RING REPRESENTS
THE SCALE OF ORGANIZATION POPULATION COMMUNITY AND BIOGEOGRAPHIC $IRECTIONAL BOXES
INDICATE DOMINANT PROCESSES AT EACH SCALE WHICH LEAD INTO ONE ANOTHER 4HE NUMBERED
DASHED LINES REFER TO THE FOUR INVASION STEPS OUTLINED IN THIS CHAPTER IMMIGRATION AND
ESTABLISHMENT SPECIES INTERACTIONS SPATIAL SPREAD AND LARGE SCALE PATTERNS
,INKING SCALE DEPENDENT PROCESSES IN INVASIONS
%XPLANATORY MECHANISMS SUCH AS THE CAUSES OF EXTINCTION CAN DRAW EXPLICIT
LINKS BETWEEN SCALES AND THIS IS WHERE WE SEE OPPORTUNITY TO BETTER UNDERSTAND
THE PROCESSES DRIVING INVASIONS AND HOW ECOLOGICAL THEORY INFORMS US ABOUT THESE
INVASIONS
4HE SCALES USED IN THIS VOLUME DO NOT NECESSARILY TRANSLATE INTO ECOLOGICALLY
RELEVANT SPATIAL SCALES EG M VS M BUT REmECT LEVELS OF ECOLOGICAL
PROCESSES AFFECTING SPECIES ABUNDANCE AND DISTRIBUTION 3CALE
EXPLICIT APPROACHES
CAN TELL US MUCH ABOUT THE NATURE OF ECOLOGICAL PROCESSES EG #ADOTTE AND
&UKAMI 2AHBEK &URTHER A NUMBER OF RECENT STUDIES HAVE SHOWN
THAT OUR UNDERSTANDING OF THE PROCESSES DRIVING SPECIES INVASIONS ARE AFFECTED
BY THE SPATIAL SCALE AT WHICH SPECIES INVASIONS ARE EXAMINED #OLLINGHAM ET AL
,LORET ET AL (AMILTON ET AL .OT ONLY DO ECOLOGICAL PROCESSES
APPEAR SCALE
CONTINGENT BUT SO DO TEMPORAL DYNAMICS OF INVASIONS &OR EXAMPLE
BOTH #OLLINGHAM ET AL AND (AMILTON ET AL SHOW THAT SPECIES INVA
SIONS APPEAR TO REACH STASIS OR EQUILIBRIUM IE STOP SPREADING EARLIER AT LARGER OR
COARSER SCALES COMPARED TO lNE SPATIAL SCALES
)S THERE A MORE GENERAL CONTEXT WITHIN WHICH WE CAN CONCEIVE OF INVASIONS
ESPECIALLY ONE THAT LINKS DIFFERENT SCALES -ANY OF THE CHAPTERS IN THIS VOLUME
DISCUSS INVASIONS IN TERMS OF SMALL POPULATIONS AVOIDING EXTINCTION 4HERE ARE
A NUMBER OF WAYS BY WHICH POPULATIONS AVOID EXTINCTION INCLUDING PROLONGING
LIFE SPANS PROTRACTED DORMANCY AND ESPECIALLY INCREASING REPRODUCTIVE RATES AND
POPULATION GROWTH RATES &OR EXAMPLE 'ROTKOPP ET AL SHOW THAT TRAITS
ASSOCIATED WITH RAPID POPULATION INCREASES HIGH RELATIVE GROWTH RATE SMALL SEED
MASS AND SHORT GENERATION TIME WERE THE BEST PREDICTORS OF PINE INVASIONS ONE OF
THE MOST AGGRESSIVE GROUPS OF PLANT INVADERS )N THIS CHAPTER WE LOOK AT A SPECIES
ABILITY TO INCREASE POPULATION SIZE INTRINSIC RATE OF INCREASE R AS THE PORTAL TO
UNDERSTANDING PATTERNS OF INVASIONS AND LINKING SPATIAL SCALES 4HROUGH THE REST
OF THIS CHAPTER WE EXPLORE THE PROCESSES AFFECTING R AT DIFFERENT SCALES OF ORGANIZA
TION AND BY UNDERSTANDING THE NATURE OF R AND HOW IT DRIVES DYNAMICS AT DIFFERENT
SCALES WE BELIEVE THAT LIGHT CAN BE SHED ON THE ULTIMATE CAUSES OF SPECIES INVA
SIONS 7HILE UNDERSTANDING THE ECOLOGICAL PROCESSES AFFECTING R APPEAR IMPORTANT
FOR UNDERSTANDING THE INVASION PROCESS WE ARE NOT CLAIMING THAT IT ALONE CAN
EXPLAIN ALL INVASIONS NOR THAT IT EVEN NECESSARILY BE THE MOST IMPORTANT FACTOR
2ATHER WE WISH TO SHOW HOW A CONCEPTUAL TREATMENT OF A SPECIES TRAIT CAN HELP
IN SYSTEMATICALLY UNDERSTANDING DIFFERENT PROCESSES AT DIFFERENT SCALES AND STAGES
OF INVASIONS
3TEP IMMIGRATION AND ESTABLISHMENT
3PECIES CLASSIlED AS
INVADERS
HAVE ONE T HING IN COMMONnTHEY HAVE OVERCOME
DISPERSAL AND ABIOTIC BARRIERS AND HAVE ARRIVED IN NEW HABITATS #HAPTERS IN THIS
VOLUME ASSUME THAT SPECIES ARE ABLE TO DISPERSE INTO NEW HABITATS THROUGH THEIR
OWN ABILITIES OR HUMAN VECTORS
-7 #ADOTTE 4 &UKAMI AND 3- -C-AHON
4HE lRST STAGE IN ANY INVASION IS THE ESTABLISHMENT OF INDIVIDUALS COMMENCING
THE NEW POPULATION 4HERE ARE A MULTITUDE OF FACTORS THAT AFFECT THE LIKELIHOOD OF
ESTABLISHMENT SUCCESS 4HESE INCLUDE ABIOTIC CONDITIONS COMPOSITION OF RESIDENT
SPECIES LIFE
HISTORY TRAITS PROMOTING INVASION "AKER AND THE NUMBER
OF IMMIGRANTS INITIATING THE POPULATION IE PROPOGULE PRESSURE SEE #ARLTON
,OCKWOOD ET AL
7ARREN ET AL #HAPTER EXPLICITLY CONSIDER THE POPULATION PROCESSES INVOLVED
WITH ESTABLISHMENT 4HEY VIEW ESTABLISHMENT AS A PROBABILISTIC PROCESS WHERE THE
PROBABILITY OF ESTABLISHMENT 0N IS DETERMINED BY THE INITIAL POPULATION SIZE N
AND BIRTH AND DEATH RATES B AND D
D N R N
0N n " " n " n "
B B
(ERE THE INTRINSIC RATE OF INCREASE R IS SIMPLY B n D %SSENTIALLY POPULATIONS
OVERCOME THE RISKS OF SMALL POPULATION SIZES EITHER THROUGH INCREASED PROPAGULE
PRESSURE N OR BY RAPIDLY INCREASING POPULATION SIZES BY INCREASING R RELATIVE TO D
VIA HIGHER BIRTH RATES
4HE INTERACTION BETWEEN R AND N IS CRITICAL FOR ESTABLISHMENT SEE #HAPTER
THEIR &IG 3PECIES WITH EXTREMELY LOW MORTALITY AND THEREFORE HIGH R HAVE A HIGH
PROBABILITY OF SUCCESS LARGELY INDEPENDENT OF THE NUMBER OF INITIAL PROGAGULES
(OWEVER SPECIES WITH LOW R OR HIGH MORTALITY WILL NEED HIGH NUMBERS OF PROPA
GLUES TO ENSURE SUCCESSFUL ESTABLISHMENT
4HESE TWO VARIABLES N AND R ARE IMPORTANT BECAUSE THE ESTABLISHMENT OF A NEW
POPULATION IS DETERMINED BY THE PROCESSES AFFECTING SMALL POPULATIONS "Y INCREAS
ING IN SIZE EITHER THROUGH IMMIGRATION OR BIRTH A POPULATION CAN BETTER AVOID
SOME OF THE PERILS PLAGUING SMALL POPULATIONS #HAPTERS
DEAL WITH THE WAYS
IN WHICH POPULATIONS OF INVADERS ESTABLISH AND PERSIST #HAPTER "UCKLEY AND
-ETCALF AND #HAPTER &RECKLETON ET AL CONSIDER WHAT HAPPENS TO SMALL POPULA
TIONS 3TOCHASTICITY DEMOGRAPHIC AND ENVIRONMENTAL AS WELL AS !LLEE EFFECTS CAN
BE STRONG FORCES AFFECTING A NEW POPULATIONS ABILITY TO PERSIST "Y UNDERSTANDING
HOW STOCHASTICITY AS WELL AS THE ABIOTIC ENVIRONMENT INmUENCE R AT DIFFERENT TIMES
AND PLACES RESEARCHERS CAN THEN MAKE LONG
TERM PROBABILISTIC PREDICTIONS REGARD
ING INVADER PERSISTENCE AND POPULATION GROWTH
3UBSEQUENT POPULATION GROWTH ALSO DEPENDS UPON R SUCH AS IN THE LOGISTIC
GROWTH MODEL SEE #HAPTER )N #HAPTER -URRELL SHOWS THAT EVEN THOUGH
LOCALIZED INVADER SUCCESS PARTIALLY DEPENDS UPON DISPERSAL RATES VARIATION IN R STILL
HAS A LARGE EFFECT ON SUCCESS (OWEVER POPULATIONS ARE NOT ISOLATED ENTITIES SOLELY
INTERACTING WITH THEIR ENVIRONMENTS #AN WE THEN INCORPORATE MULTIPLE SPECIES
INTO THE NOTION OF INVADER SUCCESS
,INKING SCALE DEPENDENT PROCESSES IN INVASIONS
3TEP SPECIES INTERACTIONS
/NCE SPECIES OVERCOME ABIOTIC AND GEOGRAPHIC BARRIERS AND ESTABLISH IN A NEW
HABITAT THEY ARE STILL FACED WITH BIOTIC BARRIERS THE BIOLOGICAL MECHANISMS AFFECT
ING THEIR INVASION EG )NDERJIT ET AL AND WILL BE FACED WITH A SUITE OF
BIOTIC INTERACTIONS COMPETITION PREDATION PATHOGENS ETC 4HESE INTERACTIONS
CAN LEAD TO VERY COMPLEX DYNAMICS 4AKEN ON THEIR OWN #HAPTERS
AND
APPEAR TO COVER RATHER IDIOSYNCRATIC INTERACTIONS WHICH CAN LEAD TO INDIVIDUAL
HYPOTHESES EXPLAINING DIFFERENT TYPES OF INVASION
PROMOTING INTERACTIONS 4HIS
WOULD SEEM TO GIVE CREDENCE TO THE NOTION THAT WE ARE UNABLE TO GENERATE LAWS IN
COMMUNITY ECOLOGY ,AWTON DESPITE THE FACT THAT UNDERSTANDING THE COM
MUNITY
LEVEL IS WHERE OUR CONSERVATION AND INVASION CONTROL EFFORTS ARE DIRECTED
3IMBERLOFF
(OWEVER BASIC ECOLOGICAL THEORY MAY BE THE BEST WAY FOR BIOLOGISTS UNDERSTAND
THE IMPORTANT INTERACTIONS AND ECOLOGICAL PROCESSES AFFECTING INVASIONS SUCCESS
3HEA AND #HESSON -OST ECOLOGICAL THEORY THAT EXAMINES THE OUTCOMES
OF SPECIES INTERACTIONS EXPLICITLY CONSIDERS THE ROLE OF NEGATIVE INTERACTIONS ON
INDIVIDUAL POPULATION GROWTH RATES AND POPULATION PERSISTENCE 7HETHER USING
,OTKA
6OLTERRA
TYPE EQUATIONS EG #OURCHAMP AND #AUT #HAPTER WHERE
PARAMETERS APPROXIMATE COMPETITIVE INTERACTIONS WITHOUT SPECIFYING A MECHA
NISM OR RESOURCE
BASED MECHANISTIC APPROACHES EG 4ILMAN #HASE AND
,EIBOLD ECOLOGICAL THEORY HAS THE POTENTIAL TO HELP ECOLOGISTS UNDERSTAND
AND POSSIBLY PREDICT THE OUTCOMES OF MULTISPECIES INTERACTIONS
4HERE MAY BE A GENERALIZED WAY TO THINK ABOUT AN INVADERS SUCCESS IN LIEU OF
NEGATIVE INTERACTIONS !GAIN WE CAN LOOK TO THE THINGS THAT AFFECT R 4HE PROCESSES
AFFECTING A POPULATIONS INTRINSIC RATE OF INCREASE CAN BE VERY COMPLEX YET FOR AN
INVADER TO PERSIST ITS LONG
TERM GROWTH RATE R MUST BE * &OR SUCCESSFUL INVAD
ERS R IS MUCH GREATER THAN AT CERTAIN TIMES AND PLACES
#HESSON CONSIDERS THAT THE LONG
TERM GROWTH RATE R IS RELATED TO THREE
MECHANISMS AS R 5R n 6. 6) WHERE R IS THE EFFECT OF mUCTUATION INDEPENDENT
MECHANISMS ON R IE IN THE PRESENCE OF COMPETITORS BUT WITHOUT ENVIRONMENTAL
CHANGES 6. IS THE EFFECT OF THE NONLINEAR COMPETITION RESPONSES TO VARIANCE IN
RESOURCES AND 6) IS THE STORAGE EFFECT SEE "OX FOR ELABORATIONS OF THESE THREE
PARAMETERS 4HE STORAGE EFFECT REPRESENTS A SPECIES ABILITY TO DIMINISH THE EFFECTS
OF COMPETITION WHEN THE SPECIES IS NOT FAVORED BY THE ENVIRONMENT #HESSON
3PECIES CAN REDUCE COMPETITIVE INTERACTIONS IN A NUMBER OF WAYS SUCH AS
MIGRATING TO REGIONS WITH FREE RESOURCES OR LACKING DOMINANT COMPETITORS CHANG
ING THEIR BEHAVIOR TO REDUCE COMPETITIVE OVERLAP OR ENTERING A DORMANT STATE AND
RE
EMERGING UNDER BETTER CONDITIONS
-7 #ADOTTE 4 &UKAMI AND 3- -C-AHON
"OX
4(% 4(2%% #/-0/.%.43 /& ,/.' 4%2- '2/74( #(%33/.
4HE mUCTUATION INDEPENDENT GROWTH RATE R DEPENDS UPON THE RATE AT
WHICH R DECLINES IN RESPONSE TO DECLINING RESOURCES FOR THE INVADER I BI
AND THE DIFFERENCE BETWEEN THE AVERAGE lTNESS OF THE INVADER IN ISOLATION
K COMPARED TO THAT IN THE PRESENCE OF RESIDENT COMPETITORS KS WHERE
K UB WHERE U IS THE PER CAPITA GROWTH RATE IN THE ABSENCE OF RESOURCE
LIMITATION 3O THAT
R BI K n KS
.ONLINEAR COMPETITION RESPONSES 6. DEPENDS UPON TWO ADDITIONAL PROP
ERTIES THE DIFFERENCE BETWEEN THE NONLINEAR RESPONSES TO LIMITING RESOURC
ES o FOR THE INVADER IN ISOLATION AND WITH COMPETITORS AND THE VARIANCE
OF THE LIMITING FACTOR FOR BOTH INVADER AND RESIDENT 6& I 3O THAT
6. BI oI n oS 6& I
&INALLY THE STORAGE EFFECT 6L IS THE ABILITY OF A SPECIES TO ALTER POPULATION
GROWTH TO AVOID LESS FAVORABLE ENVIRONMENTS WHERE THE INVADER WOULD BE
AT A COMPETITIVE DISADVANTAGE
BI n l na m
6)
N n
(ERE l IS THE CORRELATION BETWEEN THE ENVIRONMENTAL RESPONSES BETWEEN
DIFFERENT SPECIES n a IS THE BUFFERED POPULATION GROWTH AND m IS THE VARI
ANCE IN ENVIRONMENT RESPONSE 3EE #HESSON FOR FURTHER DETAILS
4HE ABILITY FOR SPECIES TO EXHIBIT BEHAVIORAL OR LIFE HISTORY ATTRIBUTES THAT SERVE
TO REDUCE THE NEGATIVE EFFECTS OF COMPETITION MAY BE WIDESPREAD 3UCCESSFUL INVA
SIVE PLANTS FOR EXAMPLE OFTEN GROW RAPIDLY IN COMPETITION REDUCED SUCCESSIONAL
HABITATS 'ROTKOPP ET AL OR HAVE SMALL SEED SIZES #ADOTTE AND ,OVETT
$OUST
(AMILTON ET AL WHICH SERVE TO HELP PLANTS MOVE FURTHER (OWE AND
3MALLWOOD AND ESTABLISH LONGER SEEDS BANKS THAN LARGE SEEDS 'ROVES
,INKING SCALE DEPENDENT PROCESSES IN INVASIONS
/F COURSE SPECIES INTERACTIONS CAN BE VERY COMPLEX 3PECIES INTERACTIONS MAY
PRODUCE DYNAMICS THAT APPEAR CHAOTIC -AY OR UNPREDICTABLE (UISMAN
AND 7EISSING 9ET #OURCHAMP AND #AUT #HAPTER $UNCAN AND &ORSYTH
#HAPTER AND 3MITH AND 3HURIN #HAPTER SHOW THAT INVASIONS CAN HAVE
REGULAR AND UNDERSTANDABLE DYNAMICS IN THE PRESENCE OF MULTIPLE SPECIES INTERAC
TIONS #OURCHAMP AND #AUT #HAPTER GO FURTHER TO SHOW HOW COMPETITION AND
PREDATION CAN INTERACT TO PROMOTE OR HINDER ANIMAL INVASIONS ON ISLANDS
3TEP SPATIAL SPREAD
$ISPERSAL AND SPREAD IS A CHARACTERISTIC DElNING PROBLEMATIC INVASIONS 3EVERAL
CHAPTERS EXPLICITLY EXAMINE WHAT HAPPENS TO POPULATIONS OF INVADERS IN A SPATIAL
CONTEXT #HAPTERS -URPHY ET AL ,EWIS ET AL AND (ARDING ET AL
3KELLAMS FORMULATION OF A POPULATIONS SPREAD ACCORDING TO A REACTION
DIFFUSION EQUATION HAS BEEN PARAMOUNT TO UNDERSTANDING HOW INVADERS MOVE
ACROSS A LANDSCAPE EG +OT ET AL +OT 7ITH #ADOTTE #HAPTER
,EWIS ET AL #HAPTER 3KELLAMS ORIGINAL FORMULATION CAN BE SUMMARIZED AS FOL
LOWS SEE #HAPTERS AND FOR MORE ON 3KELLAMS EQUATIONS
DX
3R$
DT
4HIS REACTION DIFFUSION MODEL REVEALS THAT THE RATE OF SPREAD IS DEPENDENT UPON
TWO SPECIES CHARACTERISTICS THE INTRINSIC RATE OF INCREASE R AND THE DIFFUSION COEF
lCIENT $ MEASURED AS DISTANCE OVER TIME &IGURE SHOWS THAT SPECIES WITH LOW
R THOUGH THEY MAY HAVE HIGH MOVEMENT RATES WILL NOT SPREAD VERY FAST 3IMILARLY
A HIGH R SPECIES WITH LOW MOVEMENT WILL FAIL TO SPREAD RAPIDLY
&IG 4HE VELOCITY OF SPREAD AS INmUENCED BY A DIFFUSION COEFlCIENT $ AND INTRINSIC RATE
OF INCREASE R FROM 3KELLAMS REACTION DIFFUSION EQUATION SEE TEXT
-7 #ADOTTE 4 &UKAMI AND 3- -C-AHON
3PREAD MODELS ARE THE LINK BETWEEN POPULATION AND METAPOPULATION DYNAM
ICS 3KELLAMS MODEL ASSUMES A UNIFORM ENVIRONMENT WHICH MAY BE A REALISTIC
ASSUMPTION AT LARGER SPATIAL SCALES !T INTERMEDIATE SCALES HOWEVER HABITATS ARE
OFTEN PATCHY ELICITING METAPOPULATION DYNAMICS ONTO SPECIES SPREAD EG 7OLFF
(ANSKI -URPHY ET AL #HAPTER AND (ARDING ET AL #HAPTER
USE A METAPOPULATION PERSPECTIVE TO UNDERSTAND DYNAMICS AT LANDSCAPE LEVELS
4HE RATE OF SPREAD USING A METAPOPULATION APPROACH WILL DEPEND UPON TWO
PARAMETERS THE RATES OF MIGRATION AND PATCH EXTINCTION ! HIGHER RATE OF MIGRA
TION WILL MEAN THAT NEW PATCHES HAVE A HIGHER PROBABILITY OF BEING COLONIZED AND
LOWER PATCH EXTINCTION MEANS THAT MORE LOCAL PATCHES WILL ACT AS SOURCES INCREAS
ING THE POOL OF POTENTIAL COLONIZERS !S POINTED OUT EARLIER A HIGH R CAN HELP A COLO
NIZING POPULATION OVERCOME INHERENT SMALL POPULATION RISKS AND THEREBY REDUCE
EXTINCTION PROBABILITIES AND R IS ALSO LIKELY IMPORTANT FOR IMMIGRATION RATE )F WE
ASSUME THAT POPULATION SIZE . AT ANY GIVEN TIME IS LARGELY DEPENDENT UPON R AS
IN A LOGISTIC EQUATION
.T
.T .T .T " n "
+
AND WE ASSUME THAT IMMIGRATION INTO EMPTY PATCHES IS SOME FUNCTION OF THE LOCAL
DENSITY IN OCCUPIED PATCHES THEN THE PARAMETERS THAT DETERMINE LOCAL DENSITY
LIKELY HAVE A FUNCTIONAL RELATIONSHIP WITH MEAN DENSITY AND SOME DISTANCE FUNC
TION o RELATED TO THE SPATIAL DISTRIBUTION OF EXTANT POPULATIONS )T d.T o AND
WE ASSUME THAT ENVIRONMENTAL VARIATION AFFECTING R IS RANDOM AND NORMALLY DIS
TRIBUTED ABOUT SOME MEAN AND CARRYING CAPACITY + IS CONSTANT ACROSS PATCHES
THEN R SHOULD CHARACTERIZE CHANGES IN DENSITY ACROSS A LARGER SPATIAL SCALE AND
CONSEQUENTLY THE RATE AT WHICH NEW PATCHES ARE COLONIZED
3TEP LARGE SCALE PATTERNS
3PATIAL SPREAD OF INVADERS IS A PROCESS OCCURRING ACROSS SPATIAL SCALES AND EXHIB
ITS PATTERNS BEST UNDERSTOOD BY BIOGEOGRAPHY OR MACROECOLOGY 3AX AND 'AINES
#HAPTER EXPLORE LARGE SCALE PATTERNS OF SPECIES INVASIONS 4HEY EXAMINE HOW
DIFFERENT TAXONOMIC GROUPS OF INVADERS FORM RANGE SIZE AND RICHNESS PATTERNS
THROUGH THE LENS OF EXISTING BIOGEOGRAPHICAL THEORY 4O UNDERSTAND THE PROCESSES
INmUENCING RANGE SIZE WE CAN STILL APPLY OUR BASIC VARIABLE R
4HROUGH R WE NOT ONLY UNDERSTAND SPECIES SPREAD BUT ALSO THE lNAL RANGE OF
DISTRIBUTION (OLT ET AL &RECKLETON ET AL 5SING THE WELL
FOUNDED IDEA
THAT A SPECIES DISTRIBUTION IS CORRELATED WITH MEAN LOCAL ABUNDANCE EG "ROWN
(ANSKI ET AL (OLT ET AL PROVIDES AN IMPORTANT WAY TO CONCEP
TUALIZE THESE RELATIONSHIPS ,OCAL EQUILIBRIAL POPULATION DENSITY .
OF SPECIES I AT
SITE X IS .I
X BX n DI U RIX U SO THAT SPECIES WITH HIGHER ABUNDANCES WILL
HAVE EITHER LOWER DEATH RATES D OR HIGHER BIRTH RATES B OR BOTH )F WE ASSUME
,INKING SCALE DEPENDENT PROCESSES IN INVASIONS
&IG "IRTH OR DEATH RATE DIFFERENCES ACROSS AND ENVIRONMENT TOP PANEL CAN EITHER
LEAD TO RANGE SIZE DIFFERENCES BOTTOM PANEL BETWEEN TWO SPECIES ADAPTED FROM (OLT
ET AL
THAT BIRTH RATES DECLINE FROM THE CENTER OF AN ENVIRONMENTAL GRADIENT THEN BIRTH
OR DEATH RATE DIFFERENCES BETWEEN SPECIES WILL PREDICT RANGE SIZE &IG
&RECKLETON ET AL EXPANDED THIS TO INCLUDE THE MORE REALISTIC SCENARIO
WHERE HABITAT SUITABILITY IS PATCHY SIMILAR TO THAT IN METAPOPULATION DYNAMICS
5LTIMATELY THEY SHOW THAT PATCHINESS HAD LITTLE EFFECT ON OVERALL RANGE SIZE )T IS
STILL THE DEMOGRAPHIC PARAMETERS BIRTH AND DEATH RATES WITHIN PATCHES THAT CRITI
CALLY DETERMINED RANGE SIZE &URTHER (OLT AND +EITT SHOW THAT GRADIENTS IN
EXTINCTION AND IMMIGRATION RATES AS WELL AS PATCH AVAILABILITY IN A METAPOPULA
TION CAN LIMIT RANGE SIZE
-7 #ADOTTE 4 &UKAMI AND 3- -C-AHON
#/.#,53)/.
4HIS CURSORY EXAMINATION USES A POPULATIONS INTRINSIC RATE OF INCREASE R AS THE
VEHICLE TO UNDERSTANDING INVASIONS AT DIFFERENT SCALES 7E SHOW THAT R IS NOT ONLY
IMPORTANT FOR LOCAL
POPULATION
ESTABLISHMENT
AND SIZE
BUT
CAN A LSO HELP US TO
UNDERSTAND THE PROCESSES OF SPECIES INTERACTIONS SPATIAL SPREAD AND RANGE SIZES IN
INVASIONS ACROSS MULTIPLE SPATIAL SCALES "Y UNDERSTANDING R WE CAN UNDERSTAND THE
CHANGES IN LOCAL POPULATIONS OVERTIME THE FACTORS LEAD TO THE SUCCESS OF INVADERS
WHEN INTERACTING WITH OTHER SPECIES THE RATE OF SPREAD ACROSS LARGER SPATIAL SCALES
AND THE ULTIMATE LIMITS
TO THEIR INVENTIVE
RANGES /F COURSE OTHER ECOLOGICAL AND
EVOLUTIONARY FACTORS will BE IMPORTANT FOR INDIVIDUAL INVASIONS BUT WE SHOW USING
R AS AN EXAMPLE THAT by TAKING A CONCEPTUAL APPROACH TO THE ECOLOGICAL FACTORS WE
CAN UNDERSTAND THE DYNAMICS OF INVASIONS AT DIFFERENT SCALES OF ORGANIZATION
2%&%2%.#%3
"AKER ( ' 4HE EVOLUTION OF WEEDS !NNUAL 2EVIEW OF %COLOGY AND 3YSTEMATICS
"ELYEA , 2 AND ,ANCASTER * !SSEMBLY RULES WITHIN A CONTINGENT ECOLOGY /IKOS
"ROWN * ( /N THE RELATIONSHIP BETWEEN ABUNDANCE AND DISTRIBUTION OF SPECIES
!MERICAN .ATURALIST
#ADOTTE - 7 AND &UKAMI 4 $ISPERSAL SPATIAL SCALE AND SPECIES DIVERSITY IN A
HIERARCHICALLY STRUCTURED EXPERIMENTAL LANDSCAPE %COLOGY ,ETTERS
#ARLTON * 4 0ATTERN PROCESS AND PREDICTION IN MARINE INVASION ECOLOGY "IOLOGICAL
#ONSERVATION
#HASE * - AND ,EIBOLD - ! %COLOGICAL .ICHES ,INKING #LASSICAL AND #ONTEMPORARY
!PPROACHES 5NIVERSITY OF #HICAGO 0RESS #HICAGO ),
#HESSON 0 -ECHANISMS OF MAINTENANCE OF SPECIES DIVERSITY !NNUAL 2EVIEW OF
%COLOGY AND 3YSTEMATICS
#OLLINGHAM 9 # 7ADSWORTH 2 ! 7ILLIS 3 ' (UNTLEY " AND (ULME 0 %
0REDICTING THE SPATIAL DISTRIBUTION OF ALIEN RIPARIAN SPECIES ISSUE OF SPATIAL SCALE AND
EXTENT * !PPL %COL 3UPPL
&RECKLETON 2 0 'ILL * ! .OBLE $ AND 7ATKINSON ! 2 ,ARGE
SCALE POPULATION
DYNAMICS ABUNDANCE
OCCUPANCY RELATIONSHIPS AND THE SCALING FROM LOCAL TO REGIONAL
POPULATION SIZE *OURNAL OF !NIMAL %COLOGY
'ROTKOPP % 2EJMNEK - AND 2OST 4 , 4OWARD A CAUSAL EXPLANATION OF PLANT
INVASIVENESS SEEDLING GROWTH AND LIFE
HISTORY STRATEGY OF PINE 0INUS SPECIES
!MERICAN .ATURALIST
'ROVES 2 ( 7EED ECOLOGY BIOLOGY AND SPREAD 0ROCEEDINGS OF THE &IRST )NTERNATIONAL
7EED #ONTROL #ONGRESS -ELBOURNE !USTRALIA PP
(AMILTON - ! " 2 -URRAY - 7 #ADOTTE ' # (OSE ! # "AKER # * (ARRIS AND
$ ,ICARI ,IFE
HISTORY CORRELATES OF PLANT INVASIVENESS AT MULTIPLE SPATIAL SCALES
%COLOGY ,ETTERS
,INKING SCALE DEPENDENT PROCESSES IN INVASIONS
(ANSKI ) -ETAPOPULATION %COLOGY /XFORD 5NIVERSITY PRESS /XFORD 5+
(ANSKI ) +OUKI * AND (ALKKA ! 4HREE EXPLANATIONS OF THE POSITIVE RELATIONSHIP
BETWEEN DISTRIBUTION AND ABUNDANCE OF SPECIES )N 2ICKLEFS 2 % AND 3CHLUTER $ EDS
3PECIES DIVERSITY IN ECOLOGICAL COMMUNITIES HISTORICAL AND GEOGRAPHICAL PERSPECTIVES
5NIVERSITY OF #HICAGO 0RESS #HICAGO PP
(OLT 2 $ AND +EITT 4 ( !LTERNATIVE CAUSES FOR RANGE LIMITS A METAPOPULATION
PERSPECTIVE %COLOGY ,ETTERS
(OLT 2 $ ,AWTON * ( 'ASTON + * AND "LACKBURN 4 - /N THE RELATIONSHIP
BETWEEN RANGE SIZE AND LOCAL ABUNDANCE BACK TO BASICS /IKOS
(OWE ( & AND 3MALLWOOD * %COLOGY OF SEED DISPERSAL !NNUAL 2EVIEW OF %COLOGY
AND 3YSTEMATICS
(UISMAN * AND 7EISSING & * &UNDAMENTAL UNPREDICTABILITY IN MULTISPECIES COEX
ISTENCE !MERICAN .ATURALIST
)NDERJIT #ADOTTE - 7 AND #OLAUTTI 2 ) 4HE ECOLOGY OF BIOLOGICAL INVASIONS PAST
PRESENT AND FUTURE )N )NVASIVE 0LANTS %COLOGICAL AND !GRICULTURAL !SPECTS ED )NDERJIT
"IRKHAUSER PP
+OT - %LEMENTS OF -ATHEMATICAL %COLOGY #AMBRIDGE 5NIVERSITY 0RESS .EW 9ORK
.9
+OT - ,EWIS - ! AND VAN DEN $RIESSCHE 0 $ISPERSAL DATA AND THE SPREAD OF
INVADING ORGANISMS %COLOGY
,AKATOS ) &ALSIlCATION AND THE METHODOLOGY OF SCIENTIlC RESEARCH PROGRAMMES
)N #RITICISM AND THE 'ROWTH OF +NOWLEDGE ED ) ,AKATOS AND ! -USGRAVE #AMBRIDGE
5NIVERSITY 0RESS PP
,AWTON * ( !RE THERE GENERAL LAWS IN ECOLOGY /IKOS
,LORET & -DAIL & "RUNDU ' AND (ULME 0 ,OCAL AND REGIONAL ABUNDANCE
OF EXOTIC PLANT SPECIES ON -EDITERRANEAN ISLANDS ARE SPECIES TRAITS IMPORTANT 'LOBAL
%COLOGY AND "IOGEOGRAPHY
,OCKWOOD * , #ASSEY 0 AND "LACKBURN 4 4HE ROLE OF PROPAGULE PRESSURE IN
EXPLAINING SPECIES INVASIONS 4RENDS IN %COLOGY AND %VOLUTION
,ODGE $ - "IOLOGICAL INVASIONS LESSONS FOR ECOLOGY 4RENDS IN %COLOGY AND %VOLUTION
-AY 2 - "IOLOGICAL POPULATIONS WITH NONOVERLAPPING GENERATIONS STABLE POINTS
STABLE CYCLES CHAOS 3CIENCE
2AHBEK # 4HE ROLE OF SPATIAL SCALE AND THE PERCEPTION OF LARGE
SCALE SPECIES
RICH
NESS PATTERNS %COLOGY ,ETTERS
3AX $ & 3TACHOWICZ * * AND 'AINES 3 $ 3PECIES )NVASIONS )NSIGHTS INTO %COLOGY
%VOLUTION AND "IOGEOGRAPHY 3INAUER !SSOCIATES 3UNDERLAND -!
3HEA + AND #HESSON 0 #OMMUNITY ECOLOGY THEORY AS A FRAMEWORK FOR BIOLOGICAL
INVASIONS 4RENDS %COL %VOL
3KELLAM * ' 2ANDOM DISPERSAL IN THEORETICAL POPULATIONS "IOMETRIKA
-7 #ADOTTE 4 &UKAMI AND 3- -C-AHON
4ILMAN $ 2ESOURCE COMPETITION BETWEEN PLANKTONIC ALGAE AN EXPERIMENTAL AND
THEORETICAL APPROACH %COLOGY
6ITOUSEK 0 - # - $!NTONIO , , ,OOP AND 2 7ESTBROOKS "IOLOGICAL INVASIONS
AS GLOBAL ENVIRONMENTAL CHANGE !MERICAN 3CIENTIST
7ITH + ! 4HE LANDSCAPE ECOLOGY OF INVASIVE SPREAD #ONSERVATION "IOLOGY
7OLFF * / 4HE ROLE OF HABITAT PATCHINESS IN THE POPULATION DYNAMICS OF SNOWSHOE
HARES %COLOGICAL -ONOGRAPHS
)NDEX
! ALLELOPATHY
!LLEN ((
ABUNDANCE !LLENDORF &(
!LLIARIA PETIOLATA
!LTERNANTHERA PHILOXERIODES
!BUTILON THEOPHRASTI !MARANTHUS POWELLII
!CACIA DECURRENS !MARANTHUS RETROmEXUS
!CACIA MELANOXYLON !MARANTHUS SPP
!CACIA NILOTICA !MMOPHILA ARENARIA
!CANTHASTER PLANCI !MOEBA PROTEUS
!CER SPP !NCISTRUS CHAGRENSIS
!CHILLEA MILLEFOLIUM !NDERSON 0+
!CROPTILON REPENS !NDREWARTHA ('
!DAMS +,2 !NOLIS SAGREI
ADAPTATION !NOPLOLEPIS LONGIPES
!NTENNARIA ROSEA
!NTHRISCUS SYLVESTRIS
ADAPTIVE RADIATION !PERA SPICA VENTI
!EQUIDENS COERULEOPUNCTATUS !PHANOMYCES ASTACI
!FRICA !PIS MELLIFERA
AGGREGATION !RABIDOPSIS SPP
!RABIDOPSIS SUECICA
!GRAWAL ! !RCHOCENTRUS PANAMENSIS
AGRICULTURE !RCTIUM LAPPA
!INOUCHE -, !RCTIUM MINUS
!IZEN -! !RGYRESTHIA CONJUGELLA
!KAIKES )NFROMATION #RITERION !RTEMISIA ABSINTHIUM
!RTEMISIA VULGARIS
!LEXANDER (- !SILOMAR CONFERENCE
!LLEE EFFECT
ASSEMBLY RULES
SEE COMMUNITY ASSEMBLY
!STACUS ASTACUS
)NDEX
!STYANAX AENEUS "RYCON CHAGRENSIS
!STYANAX RUBERRIMUS "RYCON PETROSUS
!USTRALIA "RYCONAMERICUS EMPERADOR
"UCKLEY 9-
"UFFON '
!VENA FATUA "UFO MARINUS
"URKE -*7
" "YERS *%
"AKER (' #
"ARBAREA VULGARIS #ACTOBLASTIS CACTORUM
"ASCOMPTE * #ADOTTE -7
"AZZAZ &! #ALIFORNIA
"ECKSTEAD *
BEHAVIOR
"ENTON 4' #AMERON 4#
"EVER *$ #AMPSURA GORGONAE
BIOGEOGRAPHY #ANADA
BIOLOGICAL CONTROL #ANNABIS SATIVA
#APSELLA BURSA PASTORIS
#ARDAMINE SCHULZII
"IOMPHALARIA GLABRATA #ARDARIA DRABA
BIOTIC HOMOGENIZATION #ARDUUS ACANTHOIDES
#ARDUUS NUTANS
BIOTIC INTERCHANGE #ARIBBEAN
BIOTIC RESISTANCE #ARLETON *4
#ASSEY 0
#ASUARINA CUNNUNGHAMIANA
"IRCHLER *! #ASWELL (
"LACKBURN 4- CAT DOMESTIC
"LANEY #3
"LEPHARISMA JAPONICUM CATALYST SPECIES
"OLKER " #ENTAUREA DIFFUSA
"RACHYHYPOPOMUS OCCIDENTALIS #ENTAUREA MACULOSA
"RACHYRHAPHIS CASCAJALENSIS
"RACHYRHAPHIS EPISCOPI #HAETOSTOMA lSCHERI
"RASSICA NAPUS #HAOTIC DYNAMICS
"RASSICA SPP
"RITAIN SEE 5NITED +INGDOM #HENOPODIUM ALBUM
"ROMUS TECTORUM #HESSON 0,
#HILE
"ROWN !($ #HIONOCHLOA PALLENS
"ROWN *( #HONDRILLA JUNCEA
)NDEX
#HONDRILLA SPP #ONVOLVULUS ARVENSIS
#HOWN 3, CORAL REEF
#HROMOLAENA ODORATA #RAWLEY -*
#HRYSOMELA SPP #REPIS TECTORUM
#IRSIUM ARVENSE CROWN OF THORNS STARlSH
#LAUSS -* CYCLIC DYNAMICS
#LEMENTS &%
#LIDEMIA HIRTA #YNOGLOSSUM OFlCINALE
#LONAL GROWTH #YPHOCHARAX MAGDALENAE
#YTISUS SCOPARIUS
CLUSTERING SEE AGGREGATION #ZECH 2EPUBLIC
COALESCENCE
#OLAUTTI 2)
$
#OLLINGHAM 9#
#OLLINS -$
$AEHLER ##
COLONIZATION $!NCONA 5
$ARWIN #
#OLPIDIUM STRIATUM
$ATURA STRAMONIUM
#OMAI , $AUCUS CAROTA
COMMUNITY ASSEMBLY $AVIS ('
$AVIS -!
COMMUNITY INVASIBILITY DENSITY DEPENDENCE
COMMUNITY SATURATION
COMPETITION
$ESCURAINIA SOPHIA
DETERMINISTIC PROCESSES
$EUTSCHEWITZ +
DEVELOPMENTAL PATHWAYS
$ICKERSON *%
APPARENT $IEKMANN /
DIET BREADTH SEE NICHE BREADTH
NEIGHBORHOOD
$IPSACUS FULLONUM
RELEASE FROM DISEASE TRIANGLE
DISPERSAL
COMPETITION
COLONIZATION TRADEOFF
CONSERVATION CONCERN
BARRIERS
CONTACT EXPERIENCE
CONTROL OF SPECIES SEE REMOVAL
)NDEX
DISTANCE EPISTEMOLOGY
ERADICATION SEE REMOVAL
KERNAL %RUCASTRUM GALLICUM
ESCAPE FROM NATURAL ENEMIES SEE
LIMITATION NATURAL ENEMIES
%SCHSCHOLZIA CALIFORNIA
LONG DISTANCE ESTABLISHMENT
SHORT DISTANCE
DISTURBANCE
DIVERSITY INVASIBILITY SEE SPECIES %UCALYPTUS OBLIQUA
RICHNESS %UPHORBIA CYPARISSIAS
DIVERSITY HOTSPOT %UPHORBIA ESULA
$OBZHANSKY 4 %UPLOTES PATELLA
DORMANCY %UROPE
$RAKE *!
$RUDE /
$UKES *3
$UNCAN 20
$YTHAM # EVOLUTION
%
%CHINOCHLOA CRUSGALLI RAPID
%CHIUM VULGARE EVOLUTIONARY STABLE STRATEGY
ECOSYSTEM IMPACTS EXPERIMENTS
%DWARDS /2 LABORATORY
%GLER &%
%LLSTRAND . NATURAL
%LTON #3 EXTINCTION
ENEMY RELEASE
ENVIRONMENT
NOVEL
VARIABILITY &
EPIDEMIC &AGOPYRUM TATARICUM
EPIGENETIC PROCESSES &ARGIONE *
)NDEX
FAUNAL INTERCHANGE SEE BIOTIC INTER 'EOPHAGUS CRASSILABRIS
CHANGE 'EPHYROCHARAX ARTRICAUDATA
FEEDBACK PROCESS 'ERMANY
&ICEDULA HYPOLEUCA 'HALAMBOR #+
&ICUS SPP 'IDO +"
&ICUS LUTEA 'LIBERT '3
&ICUS WASP INTERACTION 'OERGEN %
lRE CYCLES 'OOD 2
&ITTER ! 'OSSYPIUM SPP
&LORIDA 'OULSON $
FOOD WEB 'RAY !
'RIME *0
&ORBES 3! 'RINNELL *
&ORMICA OMNIVORA 'ROTKOPP %
&OSTER ", 'UTIERREZ $
FOUNDER EFFECTS 'YPSOPHILA PANICULATA
&OX '!
&RECKLETON 20 (
&UKAMI 4
&ULMARIS GLACIALIS HABITAT PATCHES
FUNGI
&USARIUM OXYSPORUM
(AMILTON -!
' (ANSKI )
(ARPER *,
'ALEOPSIS TETRAHIT
(AWAII
'ALINSOGA PARVImORA
'ALINSOGA QUADRIRADIATA
'ALIUM MOLLUGO
(ELIANTHUS ANNUS
'ALIUM SPURIUM
(ELIANTHUS ANOMALUS
'ASCOIGNE *#
(ELIANTHUS DEBILIS
'ASKIN *&
(ELIANTHUS DESERTICOLA
'AUSE '&
(ELIANTHUS PARADOXUS
GENERALIST
(ELIANTHUS PETIOLARIS
GENETICS (ELIANTHUS TUBEROSUS
QUATITATIVE HERBICIDE
VARIABILITY HERBICIDE RESISTANCE
HERBIVORY
GENOTYPE
GENERAL PURPOSE
INVASIVE HETEROGENEITY
GEOGRAPHIC RANGE SIZE ENVIRONMENTAL
SPATIAL
)NDEX
)RIS FULVA
)RIS HEXAGONA
TEMPORAL ISLAND BIOGEOGRAPHY
(ETEROPOGON CONTORTUS ISLANDS
HETEROZYGOSITY
(ODGSON $*
(OLCUS LANATUS
(OLT 2
(OOKER *$ *
(OPLIAS MICROLEPIS
(ORDEUM JUBATUM *ORDAN 72
HOST PATHOGEN INTERACTION SEE
PATHOGENS +
(OWARD ,/
HUMAN MODIlED INVASIONS +AWASAKI +
+EANE 2-
+EITT 4(
+ENDALL "%
+LIRONOMOS *
HYBRIDIZATION +OLBE **
+OT -
(YPERICUM PERFORATUM +OTANEN 0-
HYPER PREDATION +RITICOS $
(YPHESSOBRYCON PANAMENSIS
(YPOCHAERIS RADICATA ,
(YPOSTOMUS PANAMENSIS
,ACK 2
) ,ACTUCA SERRIOLA
,ANIUS LUDOVICIANUS
INVADER SUCCESS LATITUDINAL GRADIENT
,AYCOCK '
,EE #%
,EVINE *-
,EVINS 2
,IEBHOLD !
,INARIA DALMATICA
,INARIA VULGARIS
,INNEAUS #
,IPICUS 2.
INVASIONAL MELTDOWN ,OCKWOOD *
INVASIVE GENOTYPE SEE GENERAL PUR ,ODGE $-
POSE GENOTYPES ,OLIUM RIGIDUM
)RELAND ,OMOLINO -6
)RIS BREVICAULIS ,ONCHURA PUCTULATA
)NDEX
,ONICERA JAPONICA -ETROSIDEROS POLYMORPHA
,ONSDALE 7- -EXICO
,OTKA
6OLTERRA EQUATIONS SEE MOD
MICROBIAL COMMUNITIES
ELS ,OTKA
6OLTERRA -ICROBOTRYUM VIOLACEUM
,OTUS CORNICULATUS -ICROSTEGIUM VIMINEUM
,OVETT
$OUST * -ITCHELL #%
,UNDQUIST ,, -ITCHELL
/LDS 4
,UPINUS ARBOREOUS MODEL
,YTHRUM SALICARIA BIRTH
DEATH
DETERMINSITC
-
ESTABLISHMENT
-AC!RTHUR 2
-AC)SAAC (* (ASSELL
-ACK 2.
INDIVIDUAL
BASED
-ADAGASCAR
-AIZE ,OTKA
6OLTERRA
MANAGEMENT
LOTTERY
METAPOPULATION
.ICHOLSON
"AILEY
-ARISA CORNUARIETIS POPULATION
-ARSHALL $2
MASS EFFECTS
MASTING SEE SEED SET SYNCHRONIZED SPATIAL
SPREAD
-ATRICARIA PERFORATA
-ATSUDA ( STOCHASTIC
-AYR %
-OODY !
-C'RADY
3TEED * -OONEY (!
-C+INNEY -, -ORIN 0*
-C,AIN $+ MORPHOLOGICAL OVERDISPERSION
-EDICAGO LUPULINA
-EDITERRANEAN -OULTON -0
-OUQUET .
-ELALEUCA SPP MUTATION
-ELANDRIUM ALBUM DELETERIOUS
-ELANDRIUM DIOICUM RATES
-ELILOTUS ALBA MUTUALISM FACULTATION
-ELILOTUS OFlCINALIS
-ELILOTUS SPP MYCORRHIZAL
PLANT INTERACTIONS
MESOPREDATOR
METACOMMUNITY
-YLIUS 3$
METAPOPULATION
-YRICA FAYA
-YRIOPHYLLUM SPICATUM
)NDEX
. 0ACIlC /CEAN
NATURAL ENEMIES RELEASE FROM SEE 0ANAMA
ENEMY RELEASE 0ARAMECIUM CAUDATUM
.EOHETERANDRIA TRIDENTIGER PARASITE
.ESLIA PANICULATA
.ETHERLANDS 4HE 0ARKER "
NEUTRAL DYNAMICS 0ARKER )-
PATCH DYNAMICS SEE HABITAT PATCHES
.EW :EALAND 0ATERSON !(
PATHOGENS
FUNGAL
SOIL
NICHE SPECIALIST VS GENERALIST
BREADTH 0EARSE !3
FUNDAMENTAL 0ENNISETUM SETACEUM
REALIZED PHENOLOGY
.ICHOLSON !* PHENOTYPIC PLASTICITY SEE PLASTICITY
.ICOTIANA SPP 0HRAGMITES AUSTRALIS
NITROGEN lXING 0HYTOPHTHORA CINNAMONI
.OONBURG %' 0IABUCINA PANAMENSIS
.ORTH !MERICA 0IANKA %2
0IMELODELLA CHAGRESI
0IMM 3,
0INUS SPP
0LANTAGO LANCEOLATA
NUTRIENT CYCLES 0LANTAGO MAJOR
PLANT SOIL INTERACTIONS
/ PLASTICITY
/NDATRA ZIBETHICA
/NTARIO #ANADA
0OA ANNUA
0OECILIA GILLII
/PERCULINA VENTRICOSA
/PUNTIA SPP 0OECILIOPSIS ELONGATAL
/RYZA SPP POLLEN SWAMPING
/SENBERG #7 POLLINATION
/XALIS CORNICULATA
0 0OLYGONUM CONVOLVULUS
0OLYGONUM SPP
0ACALA 37 POLYPLOIDY
0ACIFASTACUS LENIUSCULUS 0OPPERIAN SCIENCE
)NDEX
POPULATION RANGE SIZE SEE GEOGRAPHIC RANGE
DYNAMICS 2ANUNCULUS REPENS
2APOPORT %(
2ASMUSSEN *"
RATE OF SPREAD SEE SPATIAL SPREAD
REACTION NORMS
GENETICS REACTION DIFFUSION EQUATIONS SEE
POPULATION VIABILITY ANALYSIS MODELS SPATIAL SPREAD
2EINHART +/
0ORTULACA OLERACEAE 2EJMANEK -
0OTAMOGETON CRISPUS
0OTENTILLA ARGENTEA REMOVAL OF SPECIES
0OTENTILLA RECTA
0OWER !'
PREDATION RESCUE EFFECT
RESOURCE mUCTUATIONS
RETICULATE EVOLUTION SEE HYBRIDIZA
SEED TION
PREDICTION 2EZNICK $.
2HAMDIA QUELEN
2ICCIARDI !
2ICHARDSON $-
PREY SWITCHING
PROPAGULE PRESSURE
2ICKLEFS 2%
2INELORICARIA URACANTHA
0ROSOPIS GLANDULOSA 2IVULUS BRUNNEUS
0RUNUS SEROTINA 2OBINIA PSEUDOACACIA
0SEUDOCHEIRODON AFlNIS 2OBINSON *$
0SEUDOMONAS mOURESCENS 2OEBOIDES GUATEMALANAE
0SIDIUM CATTLEIANUM 2OEBOIDES OCCIDENTALIS
0UCCINIA CHONDRILLINA 2OHDE +
0UERARIA LOBATA 2OSENZWEIG -,
0YSEK 0 2UBUS ALCEIFOLIUS
0YTHIUM SPP
3
1
3APIUM SEBIFERUM
1UANTITATIVE TRAIT LOCI 14, 3AX $&
2 3CHAAL "!
3CHIERENBECK +
RABBIT 3CHRAG !-
2ABINOWITZ $ 3CLEROTINIA SCLEROTIORUM
)NDEX
3#/0% SPATIAL INTERACTIONS
SEED DORMANCY SEE DORMANCY SPATIAL PATTERNS
SEED PREDATION SEE PREDATION SPATIAL SCALE
SEED SET SYNCHRONIZED
3ENECIO CAMBRENSIS
3ENECIO EBORACENCIS SPATIAL SPREAD
3ENECIO JACOBAEA
3ENECIO VULGRAIS SEE ALSO MODELS SPATIAL
3ETARIA FABERI SPREAD
3ETARIA VERTICILLATA SPATIAL STATISTICS
3ETARIA VIRIDIS SPECIALIST
3EYCHELLES
3HIGESAD .
3HIMA *3
3HURIN *" SPECIATION
3ILENE ALBA
3ILENE LATIFOLIA SPECIES RICHNESS AND INVASIBILITY
3ILENE NOCTImORA
3ILVERTOWN *7
3IMBERLOFF $
SPECIES SORTING
3INAPIS ARVENSIS
SPECIES AREA RELATIONSHIPS
3KELLAM *'
SOIL MICROBES SEE MICROBIAL COMMU
3PILOGALE GRACILIS
NITIES
STABILITY
3OL $
3OLANUM CAROLINENSE
3OLANUM NIGRUM 3TADLER *
3OLANUM ROSTRATUM 3TEBBINS ',
3OLIDAGO ALTISSIMA 3TELLARIA MEDIA
3OLIDAGO GIGANTEA 3TEVENS '#
3ONCHUS ARVENSIS STOCHASTIC PROCESSES
3ONCHUS ASPER
3ONCHUS OLERACEUS
3ORBUS AUCUPARIA STOCHASTICITY
3ORGHUM HALEPENSE DEMOGRAPHIC
SOURCE SINK DYNAMICS
3OUTH !FRICA
3PALDING 6-
3PARTINA ALTERNImORA ENVIRONMENTAL
3PARTINA ANGLICA 3TOHLGREN 4*
3PARTINA FOLIOSA STORAGE EFFECT
3PARTINA MARITIMA 3TRIGOPOS HABROPTILUS
3PARTINA X TOWNSENDII 3TURNUS VULGARIS
)NDEX
SUCCESSION 5NITED 3TATES
5ROCYON LITTERALIS
3UEZ #ANAL
3UKOPP ( 6
3US SCROFA
VAN (OLLE "
4 6AN +LINKEN 2$
6ERBASCUM THAPSUS
4AMARIX CHINENSIS 6ERMEIJ '*
4AMARIX GALLICA 6ERTICILLIUM DABLIAE
4AMARIX PARVImORA 6ICIA CRACCA
4AMARIX RAMOSISSIMA 6ICIA SATIVA
4AMARIX SPP 6ICIA TETRASPERMA
4ARAXACUM OFlCINALE 6ICIA VILLOSA
TENS RULE 6IEJA MACULICAUDA
TERMINOLOGY 6IOLA ARVENSIS
4ETRAHYMENA PYRIFORMIS 6IOLA RIVINIANA
4HLASPI ARVENSE VIRUSES
4HOMSON '- 6OLTERRA 6
4ILMAN $ 6ULPIA BROMOIDES
4RAGOPOGON DUBIUS
4RAGOPOGON MIRUS 7
4RAGOPOGON MISCELLUS
4RAGOPOGON PORRIFOLIUS 7ALLACE !2
4RAGOPOGON PRATENSIS 7ANG -(
TRANSPOSABLE ELEMENTS 7ARMING %
4RAVIS * 7ATKINSON !2
4RAVIS *-* 7ENDEL *&
4RIFOLIUM REPENS 7ILLIAMS 0!
4RIFOLIUM SUBTERRANEUM 7ILLIAMSON -
4RIPLEUROSPERMUM PERFORATUM 7ILSON %/
7ODZICKI +
4RITICUM !EGILOPS SPP 7OLFE ,-
TROPHIC INTERACTIONS
9
TROPICS
4URNBULL ,! 9OUNG 72
4YRIA JACOBAEAE 9UGOSLAVIA
5
:
5NITED +INGDOM
:OBEL -
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