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Conceptual Ecology and Invasion Biology Reciprocal Approaches To 2006 425 449
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Rodrigo RuizCopyright:
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!
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 ³COLOGIE
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
INVASION WITHOUT CAUSING THE EXTIRPATION OF RESIDENT SPECIES .ET SPECIES RICH
FUL
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
¾ 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 ¾ 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 $ESPITE
DOCUMENTED EVIDENCE THAT LOCAL COMMUNITIES RESIST INVASIONS SECTION MACRO
SCALE PATTERNS OF BIOTIC INTERCHANGES AND BIOLOGICAL INVASIONS SECTIONS AND
SUGGEST THAT IT IS UNLIKELY THAT LOCAL COMMUNITIES CAN BE MANAGED IN WAYS THAT
WILL PREVENT ALL INVASIONS &URTHERMORE INCREASED FREQUENCY OF HUMAN
MEDIATED
TRANSPORT OF EXOTIC SPECIES COUPLED WITH THE INCREASED VULNERABILITY OF ECOSYSTEMS
TO INVASION WILL CERTAINLY INCREASE THE PROBABILITY OF INTRODUCING EXOTIC SPECIES
WITH NOVEL ECOLOGICAL FUNCTIONS INTO NAÕVE ECOSYSTEMS 3UCH INVASION EVENTS
CLEARLY HAVE DISASTROUS REPERCUSSIONS ON THE SPECIES RICHNESS AS WELL AS THE FUNC
TIONING OF LOCAL ECOSYSTEMS +AUFMAN .ALEPA AND 3CHLOESER &RITTS
AND 2ODDA 6ANDERPLOEG ET AL )N ORDER TO CURTAIL THE INTRODUCTION OF
EXOTIC SPECIES AND PREVENT THEIR SOMETIMES DISASTROUS CONSEQUENCES OUR RESULTS
SUGGEST THAT MANAGERS AND POLICY MAKERS ALIKE NEED TO FOCUS EFFORTS ON MANAGING
3! 3MITH AND *" 3HURIN
DISPERSAL VECTORS AND PREVENTING THE ESTABLISHMENT OF EXOTIC SPECIES RATHER THAN
RELYING ON INTACT NATIVE COMMUNITIES TO REPEL INVADERS
!#+./7,%$'%-%.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
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LUTION 0AGES IN $ &UTUYMA AND - 3LATKIN EDITORS #OEVOLUTION 3INAUER
3UNDERLAND
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$ 0ILBEAM &AUNAL INTERCHANGE AND -IOCENE TERRESTRIAL VERTEBRATES OF SOUTHERN
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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
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CAL INVASIONS %COLOGY
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APPROACHES 5NIVERSITY OF #HICAGO 0RESS #HICAGO
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RIES FOR COMPETITIVE METACOMMUNITIES 0AGES
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PROCESSES IN SETTLING LOCAL SPECIES RICHNESS 0AGES
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3CHLUTER EDITORS 3PECIES DIVERSITY IN ECOLOGICAL COMMUNITIES HISTORICAL AND GEOGRAPHICAL
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AND LIMITS TO THE RICHNESS OF ECOLOGICAL COMMUNITIES A THEORETICAL PERSPECTIVE *OURNAL
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