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Conceptual Ecology and Invasion Biology Reciprocal Approaches To 2006 314 329
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Rodrigo RuizCopyright:
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0LANT POPULATIONS BY PATHOGENS
6ITOUSEK 0 - # - $!NTONIO , , ,OOPE AND 2 7ESTBROOKS "IOLOGICAL INVA
SIONS AS GLOBAL ENVIRONMENTAL CHANGE !MERICAN 3CIENTIST
7APSHERE ! * 3 (ASAN 7 + 7AHBA AND , #ARESCHE 4HE ECOLOGY OF #HONDRILLA
JUNCEA IN THE WESTERN -EDITERRANEAN *OURNAL OF !PPLIED %COLOGY
7EBB # / $ $ !CKERLY - ! -C0EEK AND - * $ONOGHUE 0HYLOGENIES AND
COMMUNITY ECOLOGY !NNUAL 2EVIEW OF %COLOGY AND 3YSTEMATICS
7ESTE ' AND ' # -ARKS 4HE BIOLOGY OF 0HYTOPHTHORA CINNAMOMI IN !USTRALASIAN
FORESTS !NNUAL 2EVIEW OF 0HYTOPATHOLOGY
7ILLIAMSON - ( "IOLOGICAL )NVASIONS #HAPMAN (ALL .EW 9ORK
7ILLIS ! * * -EMMOTT AND 2 ) &ORRESTER )S THERE EVIDENCE FOR THE POST
INVASION
EVOLUTION OF INCREASED SIZE AMONG INVASIVE PLANT SPECIES %COLOGY ,ETTERS
7INGlELD - * " 3LIPPERS * 2OUX AND " $ 7INGlELD 7ORLDWIDE MOVEMENT OF
EXOTIC FOREST FUNGI ESPECIALLY IN THE TROPICS AND THE SOUTHERN HEMISPHERE "IOSCIENCE
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 6ÉZQUEZ
).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
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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 2EJMÉNEK 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 2EJMÉNEK
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 6ÉZQUEZ
"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 2EJMÉNEK 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
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.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
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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 6ÉZQUEZ 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
'ALÉPAGOS )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 6ÉZQUEZ
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 6ÉZQUEZ AND !IZEN B 6ÉZQUEZ 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 6ÉZQUEZ 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 6ÉZQUEZ 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 6ÉZQUEZ
HURDLES IN THE STUDY OF SPECIES INTERACTIONS -ARTINEZ ET AL .OVOTNâ ET AL
6ÉZQUEZ 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 .OVOTNâ 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 6ILÌ
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 # * -ELIÉN 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 6ÉZQUEZ
%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 'ALÉPAGOS )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
.OVOTNâ 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
2EJMÉNEK - 7HAT MAKES A SPECIES INVASIVE 0AGES
IN 0 0YÝEK EDITOR 0LANT
INVASIONS GENERAL ASPECTS AND SPECIAL PROBLEMS !CADEMIC 0UBLISHING !MESTERDAM
2EJMÉNEK - )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 - 2EJMÉNEK 0LANT
INVASIONS THE ROLE OF MUTUALISMS "IOLOGICAL 2EVIEWS
2OY * - , .AVAS AND , 3ONIÏ )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
6ÉZQUEZ $ 0 AND - ! !IZEN .ULL MODEL ANALYSES OF SPECIALIZATION IN PLANT
POLLINATOR INTERACTIONS %COLOGY
6ÉZQUEZ $ 0 AND - ! !IZEN A !SYMMETRIC SPECIALIZATION A PERVASIVE FEATURE OF
PLANT
POLLINATOR INTERACTIONS %COLOGY
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