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Conceptual Ecology and Invasion Biology Reciprocal Approaches To 2006 408 424
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)NVASIVE SPECIES IN METAPOPULATION SYSTEMS
-AC!RTHUR 2 ( AND % / 7ILSON 4HE THEORY OF ISLAND BIOGEOGRAPHY 0RINCETON
5NIVERSITY 0RESS 0RINCETON
.EUBERT - AND ( #ASWELL $EMOGRAPHY AND DISPERSAL CALCULATIONS A ND SENSITIVITY
ANALYSIS OF INVASION SPEED FOR STRUCTURED POPULATIONS %COLOGY
.EE 3 AND 2 - -AY $YNAMICS OF METAPOPULATIONS HABITAT D ESTRUCTION AND COM
PETITIVE COEXISTENCE *OURNAL OF !NIMAL %COLOGY
.OUHUYS 3 6 AND ) (ANSKI #OLONISATION RATES AND DISTANCES OF A HOST BUTTERmY
AND TWO SPECIlC PARASITOIDS IN A FRAGMENTED LANDSCAPE
/VASKAINEN / AND ) (ANSKI -ETAPOPULATION DYNAMICS IN HIGHLY FRAGMENTED LAND
SCAPES 0AGES
IN ) (ANSKI AND / % 'AGGIOTTI EDITORS %COLOGY 'ENETICS AND
%VOLUTION OF -ETAPOPULATIONS %LSEVIER!CADEMIC "URLINGTON
0OINTIER * 0 AND 0 $AVID "IOLOGICAL CONTROL OF "IOMPHALARIA GLABRATA THE INTER
MEDIATE HOST OF SCHISTOSOMES BY -ARISA CORNUARIETIS IN PONDS OF 'UADELOUPE LONG TERM
IMPACT ON THE LOCAL SNAIL FAUNA AND AQUATIC mORA "IOLOGICAL #ONTROL
2OLÉN
!LVAREZ % + *OHANNESSON AND * %RLANDSSON 4HE MAINTENANCE OF A CLINE IN
THE MARINE SNAIL ,ITTORINA SAXATILIS THE ROLE OF HOME SITE ADVANTAGE AND HYBRID lTNESS
%VOLUTION
3ACCERI ) - +UUSSAARI - +ANKARE 0 6IKMAN 7 &ORTELIUS AND ) (ANSKI
)NBREEDING AND EXTINCTION IN A BUTTERmY METAPOPULATION .ATURE
3CHÚPS + %FFECT OF DEFORESTATION ON THE SPATIAL DYNAMICS OF AN ENDANGERED WEEVIL
SPECIES 0AGE IN (ABITAT ,OSS (ELSINKI
3EPT (AKAPAINO /Y (ELSINKI
3HIGESADA . AND + +AWASAKI "IOLOGICAL )NVASIONS 4HEORY AND 0RACTICE /XFORD
5NIVERSITY 0RESS /XFORD
3HURIN * " 0 !MARESEKARE * - #HASE 2 $ (OLT - & (OOPES 2 ,AW AND - !
,EIBOLD !LTERNATIVE STABLE STATES AND REGIONAL COMMUNITY STRUCTURE *OURNAL OF
4HEORETICAL "IOLOGY
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
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