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Conceptual Ecology and Invasion Biology Reciprocal Approaches To 2006 297 313
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! 3ATAKE /.
"JRNSTAD AND 9 )WASA
3HIBATA - ( 4ANAKA AND 4 .AKASHIZUKA 3YNCHRONIZED ANNUAL SEED PRODUCTION
BY PRINCIPAL TREE SPECIES IN A TEMPERATE DECIDUOUS FOREST *APAN %COLOGY
3CHNURR * , 2 3 /STFELD AND $ #ANHAM $IRECT AND INDIRECT EFFECT OF MASTING ON
RODENT POPULATIONS AND TREE SEED SURVIVAL /IKOS
3HEA + AND 0 #HESSON #OMMUNITY ECOLOGY THEORY AS A FRAMEWORK FOR BIOLOGICAL
INVASIONS 4RENDS IN %COLOGY AND %VOLUTION
3HIBATA - ( 4ANAKA AND 4 .AKASHIZUKA #AUSE AND CONSEQUENCES OF MAST SEED
PRODUCTION OF FOUR CO
OCCURRING #ARPINUS SPECIES IN *APAN %COLOGY
3ILVERTOWN * 7 4HE EVOLUTIONALY ECOLOGY OF MAST SEEDING IN TREES "IOLOGICAL
*OURNAL OF THE ,INNEAN 3OCIETY
3MITH # # * , (AMRICK AND # , +RAMER 4HE ADVANTAGE OF MAST YEARS FOR WIND
POLLINATION !MERICAN .ATURALIST
3ORK 6 , * "RAMBLE AND / 3EXTON %COLOGY OF MAST
FRUITING IN THREE SPECIES OF
NORTH !MERICAN DECIDUOUS OAKS %COLOGY
3PERENS 5 A &RUIT PRODUCTION IN 3ORBUS AUCUPARIA , 2OSACEAE AND PRE
DISPERSAL
SEED PREDATION BY THE APPLE FRUIT MOTH !RGYRESTHIA CONJUGELLA :ELL /ECOLOGIA
3PERENS 5 B ,ONG
TERM VARIATION IN AND EFFECTS OF FERTILIZED ON mOWER FRUIT AND SEED
PRODUCTION IN THE TREE 3ORBUS AUCUPARIA 2OSACEAE %COGRAPHY
4UTIN 4 ' 6 ( (EYWOOD . ! "URGES $ - -OORE $ ( 6ALENTINE 3 - 7ALTERS AND
$ ! 7EBB &LORA EUROPAEA ND EDN #AMBRIDGE 5NIVERSITY 0RESS #AMBRIDGE
7OLFF * / 0OPULATION mUCTUATIONS OF MAST
EATING RODENTS ARE CORRELATED WITH PRO
DUCTION OF ACORNS *OURNAL OF -AMMALOGY
#HAPTER THIRTEEN
)NVASIONS AND THE REGULATION
OF PLANT POPULATIONS
BY PATHOGENS
' 3 'ILBERT AND ) - 0ARKER
).42/$5#4)/.
4HE POTENTIAL OF PATHOGENS TO HAVE DRAMATIC IMPACTS ON PLANT POPULATIONS IS MADE
CLEAR BY FAMILIAR STORIES OF THE )RISH POTATO FAMINE &RY AND 'OODWIN THE
ECOLOGICAL EXTINCTION OF CHESTNUTS CAUSED BY CHESTNUT BLIGHT !NAGNOSTAKIS
AND THE TRANSFORMATION OF !USTRALIAN *ARRAH FORESTS TO SCRUBLAND BY 0HYTOPHTHORA
CINNAMOMI 7ESTE AND -ARKS 3IMILARLY THE ANNUAL WORLDWIDE EXPENDI
TURE OF OVER BILLION IN FUNGICIDE APPLICATION $ONALDSON ET AL REmECTS
THE TOLL FUNGAL PATHOGENS ALONE CAN TAKE ON PLANT GROWTH AND FECUNDITY IN
AGRICULTURAL SYSTEMS 3OME OF THE MOST NOTABLE EXAMPLES OF THESE IMPACTS ARISE
WHEN PATHOGENS ARE INTRODUCED INTO NOVEL BIOLOGICAL ENVIRONMENTS INVASIVE AND
EMERGENT PATHOGENS CONTINUE TO FRUSTRATE THE BEST EFFORTS OF RESOURCE MANAGERS
CONSERVATION BIOLOGISTS AND PLANT PROTECTION AGENCIES 7ESTE AND -ARKS
$AUGHTREY ET AL 'OODELL ET AL 'ORDON ET AL -C$ONALD AND (OFF
7INGlELD ET AL 'ILBERT 2IZZO AND 'ARBELOTTO 0ARKER AND
'ILBERT #ONCURRENT WITH EFFORTS TO REDUCE THE IMPACTS OF UNWANTED DISEAS
ES THERE IS BROAD INTEREST AMONG RESEARCHERS AGRICULTURALISTS AND LAND MANAGERS
TO HARNESS THE DESTRUCTIVE POTENTIAL OF PLANT PATHOGENS TO CONTROL WEEDY PLANTS
(ASAN AND !YRES #HARUDATTAN AND $INOOR
-7 #ADOTTE ET AL EDS #ONCEPTUAL ECOLOGY AND INVASION BIOLOGY n
¥ 3PRINGER 0RINTED IN THE .ETHERLANDS
'3 'ILBERT AND )- 0ARKER
)N NATURAL ECOSYSTEMS PATHOGENS HAVE GREAT POTENTIAL TO INmUENCE THE DYNAM
ICS AND COMPOSITION OF PLANT POPULATIONS AND COMMUNITIES THROUGH DENSITY
DEPENDENT AND COEVOLUTIONARY DYNAMICS SEE REVIEWS IN $INOOR AND %SHED
"URDON *AROSZ AND $AVELOS !LEXANDER ET AL !LEXANDER AND
(OLT 'ILBERT )N MANY PLANT COMMUNITIES PLANT PATHOGENS
MAY PREVENT COMPETITIVE EXCLUSION AND THUS HELP MAINTAIN SPECIES DIVERSITY
'ILLETT 0ACKER AND #LAY 'ILBERT 7RIGHT 'ILBERT
#OLLECTIVE INSIGHTS FROM RESEARCH ON EPIDEMIC DISEASES BIOLOGICAL CONTROL AND
THE EVOLUTIONARY ECOLOGY OF DISEASES IN NATURAL ECOSYSTEMS PROVIDE A ROBUST BASIS
FOR IDENTIFYING WHEN PATHOGENS ARE LIKELY TO BE IMPORTANT IN REGULATING PLANT
POPULATIONS AND THEIR IMPLICATIONS FOR UNDERSTANDING BIOLOGICAL INVASIONS
(ERE WE DRAW BROADLY FROM A DIVERSE LITERATURE TO PLACE THE REGULATION OF PLANT
POPULATIONS BY PATHOGENS INTO THE CONTEXT OF TWO KEY COMPLEMENTARY THEORIES
ABOUT THE ROLE OF PATHOGENS IN BIOLOGICAL INVASIONS "IOTIC 2ESISTANCE -ARON AND
6ILA AND %SCAPE FROM .ATURAL %NEMIES +EANE AND #RAWLEY 4ABLE
7E THEN CONSIDER THE PRACTICAL IMPLICATIONS FOR USING INTRODUCED PATHOGENS FOR
CLASSICAL BIOLOGICAL CONTROL OF INTRODUCED INVASIVE PLANTS 4ABLE ILLUSTRATED WITH
A DETAILED CASE STUDY OF THE CONTROL OF #HONDRILLA BY RUST FUNGI
0,!.4 $)3%!3%3 !.$ 0/05,!4)/. 2%'5,!4)/.
0ATHOGENS REDUCE THE lTNESS OF INDIVIDUAL PLANTS BY KILLING THEM REDUCING GROWTH
IMPEDING COMPETITIVE ABILITY OR BY ROTTING FRUITS OR SEEDS SEE RECENT REVIEW IN
'ILBERT 4HE GREATER PROBABILITY OF PATHOGEN SPREAD BETWEEN CLOSELY SPACED
HOST PLANTS MEANS THAT MOST FUNGAL PLANT DISEASES SHOW DENSITY
DEPENDENT DEVEL
OPMENT )N ADDITION DENSELY SPACED PLANTS MAY CREATE MICROCLIMATES THAT ENCOUR
AGE PATHOGEN GROWTH AND HOSTS STRESSED BY COMPETITION MAY BE MORE SUSCEPTIBLE
TO DISEASE "URDON AND #HILVERS 'ILBERT 4HE COMBINATION OF STRONG
IMPACTS ON INDIVIDUAL HOST PLANTS AND DENSITY
DEPENDENT DISEASE DEVELOPMENT
SUGGESTS THAT PATHOGENS SHOULD BE POWERFUL REGULATORS OF PLANT POPULATIONS
.EVERTHELESS ASIDE FROM EPIDEMICS CAUSED BY INTRODUCED PATHOGENS THERE ARE
REMARKABLY FEW EMPIRICAL STUDIES SHOWING THAT PLANT DISEASES ARE RESPONSIBLE
FOR REGULATING PLANT POPULATION DYNAMICS IN NATURAL ECOSYSTEMS 'ILBERT
)N LARGE PART THIS ABSENCE REmECTS THE DIFlCULTIES OF ISOLATING DISEASE IMPACTS FROM
OTHER FACTORS AND THE STRONG FOCUS ON DISEASES WITH ECONOMIC IMPORTANCE "UT
PHYSIOLOGICAL OR EVOLUTIONARY RESPONSES BY THE HOST MAY ALSO COUNTERACT THE
REGULATORY ACTIONS OF PATHOGENS IN NATURAL ECOSYSTEMS )N PARTICULAR PLANTS THAT
SURVIVE AFTER DISEASE HAS KILLED OR STUNTED COMPETING CONSPECIlC NEIGHBORS MAY
SHOW A COMPENSATORY RESPONSE THAT OFFSETS NUMERICAL LOSSES FROM DISEASE AT THE
POPULATION LEVEL &RIESS AND -AILLET !LEXANDER AND (OLT !LEXANDER
AND -IHAIL )N ADDITION MATERNALLY
TRANSMITTED INDUCED RESISTANCE CAN
GENERATE CROSS
GENERATIONAL EFFECTS THAT AMELIORATE THE NUMERICAL IMPACTS OF
DISEASE IN SUCCEEDING GENERATIONS !GRAWAL ET AL &INALLY THE GENETICS
4ABLE )MPLICATIONS OF DIFFERENT FEATURES OF PLANT
PATHOGEN INTERACTIONS FOR THREE AREAS OF INVASION BIOLOGY THE TWO THEORIES OF BIOTIC
RESISTANCE AND ESCAPE FROM NATURAL ENEMIES AND THE lELD OF CLASSICAL BIOLOGICAL CONTROL
&EATURES OF PLANT
PATHOGEN "IOTIC RESISTANCE %SCAPE FROM NATURAL ENEMIES "IOLOGICAL CONTROL
INTERACTIONS
)MPACT OF PATHOGEN ON HOST 0RE
ADAPTED PATHOGENS IN NEW 0ATHOGENS IN NATIVE RANGE LIMIT 4O BE EFFECTIVE CONTROL AGENTS
RANGE ARE HIGHLY VIRULENT ON HOST DENSITY OR DISTRIBUTION PATHOGENS MUST BE IMPORTANT IN
NAÕVE HOSTS )NTRODUCED PLANTS LEAVE POPULATION REGULATION IN NATIVE
VIRULENT PATHOGENS BEHIND RANGE
)NTERACTIONS WITH ,OCAL PATHOGENS ARE ADAPTED 0LANTS MAY ESCAPE NATURAL )NTRODUCED BIOCONTROL PATHOGENS
ENVIRONMENT TO LOCAL ENVIRONMENT ENEMIES IF INTRODUCED INTO AN MAY FAIL IF POORLY ADAPTED TO
DISEASE TRIANGLE ENVIRONMENT NOT CONDUCIVE TO LOCAL ENVIRONMENT
DISEASE DEVELOPMENT
(OST RANGE OF PATHOGEN 'ENERALIST PATHOGENS ARE MORE !SSUMES SPECIALIST PATHOGENS PLAY /NLY SPECIALIST PATHOGENS CAN BE
LIKELY TO ACQUIRE NEWLY INTRODUCED A UNIQUE ROLE IN REGULATING PLANT UTILIZED AS CONTROL AGENTS
HOST SPECIES POPULATIONS
.ATURAL HISTORY $ENSITY DEPENDENT DISEASE 0ATHOGENS WITH RESTING STRUCTURES 0ATHOGENS WITH RESTING STRUCTURES
0LANT POPULATIONS BY PATHOGENS
LIFE HISTORY OF PATHOGEN DEVELOPMENT MAY CHANGE IMPACT OR ALTERNATE LIFE HISTORY STRATEGIES SHOULD SHOW GREATER SUCCESS
ON INVADING HOSTS AS INVASION MOST LIKELY TO ACCOMPANY
PROCEEDS INTRODUCED PLANTS SUCH PATHOGENS
MAY HAVE GREATER VIRULENCE
2APID 0ATHOGENS MAY ACQUIRE INTRODUCED (OST MAY LOSE DEFENSES TO ORIGINAL (OST MAY DEVELOP RESISTANCE TO
EVOLUTIONARY CHANGES HOST THROUGH EVOLUTIONARY HOST PATHOGENS AFTER INTRODUCTION INTRODUCED BIOCONTROL AGENT
SHIFT 6IRULENCE MAY INCREASE
QUICKLY WHEN HOST BECOMES
COMMON
'3 'ILBERT AND )- 0ARKER
OF PLANT
PATHOGEN INTERACTIONS CAN BE HIGHLY DYNAMIC WITH LARGE CHANGES IN
PATHOGEN VIRULENCE OR HOST RESISTANCE EVOLVING IN A FEW GENERATIONS EG "URDON
AND 4HOMPSON "ISHOP ET AL 3UCH RAPID EVOLUTIONARY CHANGES ARE
EXPECTED TO STRONGLY INmUENCE THE IMPACT OF PATHOGENS ON HOST NUMERICAL DYNAM
ICS !LEXANDER ET AL
")/4)# 2%3)34!.#% !.$ %3#!0% &2/- .!452!, %.%-)%3
&OR DISEASE TO DEVELOP VIRULENT PATHOGENS SUSCEPTIBLE HOST PLANTS AND SUITABLE
ENVIRONMENTAL CONDITIONS MUST CONVERGE COMMONLY CALLED THE $ISEASE 4RIANGLE
#HANGES IN ANY OF THESE COMPONENTS CAN REDUCE OR INCREASE HOW MUCH DISEASES
AFFECT PLANT POPULATION DYNAMICS )F PLANTS ARE INTRODUCED TO AN ENVIRONMENT
WHERE VIRULENT PATHOGENS ARE NOT PRESENT OR WHERE ENVIRONMENTAL CONDITIONS
DO NOT FAVOR DISEASE DEVELOPMENT THE PLANT POPULATION MAY BE RELEASED FROM
PREVIOUS REGULATION BY PATHOGENS 3IMILARLY PLANTS INTRODUCED TO A NEW LOCALE
WITH NOVEL VIRULENT PATHOGENS OR WHERE ENVIRONMENTAL CONDITIONS FAVOR DISEASE
DEVELOPMENT MAY EXPERIENCE UNPRECEDENTED POPULATION REGULATION BY PATHOGENS
3UCH CHANGES IN THE PREVALENCE AND SEVERITY OF DISEASES UNDER DIFFERENT CONDITIONS
HAVE LONG BEEN THOUGHT TO PLAY AN IMPORTANT ROLE IN THE PROCESS OF BIOLOGICAL
INVASIONS IN TWO HYPOTHETICAL WAYS
)N NATURAL HABITATS NATIVE PATHOGENS COLONIZE AND ARE HIGHLY VIRULENT ON NAÕVE
INTRODUCED PLANT SPECIES AND PREVENT POPULATION GROWTH "IOTIC 2ESISTANCE
0ATHOGENS WERE IMPORTANT IN REGULATING THE PLANT POPULATION IN ITS NATIVE
RANGE BUT ARE NOT IN THE INTRODUCED RANGE %SCAPE FROM .ATURAL %NEMIES
"IOTIC RESISTANCE
4HE GREAT MAJORITY OF PLANT INTRODUCTIONS DO NOT RESULT IN INVASIONS -ACK
7ILLIAMSON 2ATHER MANY INTRODUCED SPECIES EITHER FAIL TO THRIVE ALTOGETH
ER OR ARE RESTRICTED TO HUMAN CULTIVATION UNABLE TO BUILD SELF
SUSTAINING POPULA
TIONS IN WILD PLANT COMMUNITIES OR EVEN IN DISTURBED RANGELAND COMMUNITIES
2ESEARCH ATTEMPTING TO IDENTIFY PLANT TRAITS THAT CONFER INVASIVENESS 2EICHARD
AND (AMILTON 2EJMÉNEK AND 2ICHARDSON SHOWS THAT OUR PREDICTIVE
ABILITY IS IMPERFECT AND IT SEEMS TO BE MORE DIFlCULT TO PREDICT WHICH INTRODUC
TIONS WILL FAIL THAN WHICH WILL SUCCEED 2EICHARD AND (AMILTON "IOTIC
RESISTANCE IS ONE POSSIBLE EXPLANATION FOR WHY SOME INTRODUCTIONS FAIL WHEN THEY
hSHOULDv SUCCEED NATIVE PESTS AND PATHOGENS COLONIZE EXOTIC PLANTS AND ELIMINATE
THEM BEFORE THEY CAN ESTABLISH A VIABLE POPULATION %LTON 3IMBERLOFF
-ACK
0ATHOGENS WILL CONTRIBUTE TO BIOTIC RESISTANCE ONLY WHEN THREE CONDITIONS ARE
MET &IRST THE PATHOGENS INVOLVED MUST NOT BE NARROW HOST SPECIALISTS )N PLANT
COMMUNITIES WITH HIGH HOST DIVERSITY AND CORRESPONDING LOW HOST DENSITY
GENERALIST PLANT PATHOGENS SHOULD DOMINATE 'ILBERT ET AL 'ILBERT
0LANT POPULATIONS BY PATHOGENS
WHICH COULD CONTRIBUTE TO THE HIGH INVASION RESISTANCE OF HYPER
DIVERSE ECO
SYSTEMS LIKE TROPICAL RAIN FORESTS &INE 3ECOND BECAUSE DENSITIES OF THE
INTRODUCED HOST WILL USUALLY BE LOW IMMEDIATELY FOLLOWING INTRODUCTION PATHO
GENS INVOLVED IN BIOTIC RESISTANCE WOULD NOT BE THOSE FOR WHICH TRANSMISSION OR
HOST SWITCHING WAS STRONGLY DENSITY
DEPENDENT 4HIRD THE PATHOGEN MUST EXACT
A HIGH lTNESS COST ON THE HOST #OMBINATIONS OF GENERALIZATION AND HIGH VIRU
LENCE ARE NOT UNUSUAL IN PATHOGENS ESPECIALLY FOR SPECIES THAT HAVE LONG RESTING
STAGES OR SAPROPHYTIC ABILITY EG 0HYTOPHTHORA CINNAMOMI 6ERTICILLIUM DAHLIAE
&USARIUM OXYSPORUM %VEN PATHOGENS THAT CAN ATTACK MANY HOSTS HAVE DIF
FERENTIAL IMPACTS ON DIFFERENT HOST SPECIES )F A PATHOGEN HAS A GREATER NEGATIVE
EFFECT ON THE COMPETITIVE ABILITY OF THE NON
NATIVE HOSTS THAN ON THE NATIVE HOSTS
THEN IT WILL CONTRIBUTE TO THE COMPETITIVE EXCLUSION OF THE NON
NATIVE +EANE AND
#RAWLEY
3INCE EVEN FAIRLY SPECIALIZED PATHOGENS OFTEN INFECT MANY MEMBERS OF THE SAME
GENUS OR FAMILY IT IS LIKELY THAT THE NUMBER OF PATHOGENS COMPETENT TO INFECT A
NOVEL HOST DEPENDS ON WHETHER IT IS PHYLOGENETICALLY RELATED TO NATIVE SPECIES
ALREADY IN THE COMMUNITY 4HEREFORE BIOTIC RESISTANCE SHOULD FAVOR INVASION BY
SPECIES WITH NO CLOSE RELATIVES $ESPITE THIS CLEAR PREDICTION AND THE ACCEPTED
PRACTICE OF USING PHYLOGENETIC RELATIONSHIPS IN QUARANTINE AND TRADE POLICY TO
TARGET RELATIVES OF WEEDY PLANTS OR IDENTIFY POTENTIAL CARRIER HOSTS SURPRISINGLY
FEW STUDIES OF INVASION EVEN MENTION THE PHYLOGENETIC STRUCTURE OF THE INVADED
COMMUNITIES )N ONE RARE ATTEMPT -ACK FOUND THAT FOR OUT OF REGIONAL
mORAS NATURALIZED SPECIES WERE MORE COMMON IN GENERA WITH NO NATIVE CONGENERS
THAN THOSE WITH NATIVE CONGENERS CONSISTENT WITH THE IDEA OF BIOTIC RESISTANCE
(OWEVER THIS STUDY WAS NOT ABLE TO CONTROL FOR THE EFFECT OF OPPORTUNITY WHETHER
PLANTS WITHOUT NATIVE CONGENERS HAVE A HIGHER PROBABILITY OF SUCCESSFUL INVASION
GIVEN THEIR PROBABILITY OF INTRODUCTION 5SING A DIFFERENT APPROACH $UNCAN AND
7ILLIAMS COMPILED A LIST OF ALL PLANT SPECIES THAT HAVE EVER BEEN INTRODUCED
FOR CULTIVATION IN .EW :EALAND 4HEY FOUND THAT INTRODUCED SPECIES IN GENERA
THAT ALREADY HAD RESIDENT NATIVES WERE MORE LIKELY NOT LESS LIKELY TO SUCCESSFULLY
NATURALIZE 4HEY SUGGEST THAT SPECIES WITH NATIVE CONGENERS MAY SHARE CHARAC
TERISTICS THAT MAKE THEM MORE lT IN THE INTRODUCED RANGE AND THIS FACTOR OVER
WHELMS THE POTENTIAL EFFECT OF LOCAL NATURAL ENEMIES
4HERE IS A STRIKING NEED FOR MORE STUDIES TO DETERMINE WHETHER AND WHEN
BIOTIC RESISTANCE BY NATIVE PATHOGENS OCCURS "ECAUSE THERE IS LITTLE INFORMATION
ON WHERE AND WHEN UNINTENTIONAL SPECIES INTRODUCTIONS FAIL ASSIGNING MECHA
NISMS TO THESE FAILURES HAS BEEN NEARLY IMPOSSIBLE (ORTICULTURE FORESTRY AND
AGRICULTURE EACH PROVIDE EXAMPLES OF ENDEMIC PATHOGENS THAT HAVE DECIMATED
INTRODUCED PLANT SPECIES SO THAT GROWING THESE SPECIES IS NO LONGER ECONOMICALLY
VIABLE -ACK #OUTINHO ET AL 7INGlELD ET AL (OWEVER EXTRA
P OLATING FROM AGRICULTURAL OR SILVICULTURAL EXAMPLES T O I NVASIVE INTRODUCED SPECIES
REQUIRES CAUTION SINCE THE REGENERATION OF HOSTS IS CONTROLLED PREVENTING THE HOST
POPULATION FROM EVOLVING RESISTANCE OR TOLERANCE TO THE PATHOGEN !DDITIONALLY
SIMPLIlED SYSTEMS IN AGRICULTURE AND FORESTRY MAY SIMPLY LACK MODERATING EFFECTS
'3 'ILBERT AND )- 0ARKER
OF GREATER BIOTIC COMPLEXITY IN LESS MANAGED SYSTEMS EG HYPERPARASITES OF
THE PATHOGENS
%SCAPE FROM NATURAL ENEMIES
)NTRODUCED PLANTS THAT BECOME INVASIVE WEEDS ARE AMONG OUR MOST CHALLENGING
ENVIRONMENTAL PROBLEMS $!NTONIO AND 6ITOUSEK /FlCE OF 4ECHNOLOGY
!SSESSMENT 6ITOUSEK ET AL 0ARKER ET AL -ACK ET AL
AND ESCAPE FROM NATURAL ENEMIES PROVIDES A MECHANISM TO EXPLAIN THE INCREASED
GROWTH AND DENSITY OF INTRODUCED SPECIES $ARWIN #RAWLEY "LOSSEY
AND .ÚTZOLD 4ILMAN -ARON AND 6ILA 3IEMANN AND 2OGERS
+EANE AND #RAWLEY 4HE h.ATURAL %NEMIES (YPOTHESISv POSITS
THAT INTRODUCED SPECIES LEAVE BEHIND THEIR NATURAL ENEMIES HERBIVORES SEED
PREDATORS AND PATHOGENS AND ARE THEREBY RELEASED FROM A KEY REGULATING
FACTOR LEADING TO A DRAMATIC INCREASE IN PLANT VIGOR POPULATION GROWTH ANDOR
COMPETITIVE ABILITY 4HIS IDEA FORMS THE JUSTIlCATION FOR CLASSICAL BIOLOGICAL CONTROL
IN WHICH NATURAL ENEMIES ARE BROUGHT FROM THE NATIVE RANGE TO CONTROL WEEDY
INVADERS (UFFAKER AND -ESSENGER $E"ACH AND 2OSEN 9ET DESPITE
THE IMPORTANCE OF THE .ATURAL %NEMIES (YPOTHESIS RIGOROUS EMPIRICAL TESTS ARE
FEW -ARON AND 6ILA
! NUMBER OF STUDIES HAVE TESTED WHETHER INTRODUCED PLANTS TEND TO GROW
FASTER OR LARGER AND WHETHER THERE IS EVIDENCE OF REDUCED ALLOCATION TO HERBIVORE
DEFENSES IN THE NEW RANGE #RAWLEY "LOSSEY AND .ÚTZOLD 3IEMANN
AND 2OGERS BUT SEE 7ILLIS ET AL 4HEBAUD AND 3IMBERLOFF 6ILA
ET AL 7HILE SEVERAL OF THESE STUDIES HAVE SUGGESTED ECOLOGICAL RELEASE AND
EVOLUTION AWAY FROM DEFENSE TOWARD COMPETITIVE ABILITY THEY LACK DIRECT EVI
DENCE OF INVOLVEMENT BY NATURAL ENEMIES "LOSSEY AND .ÚTZOLD 3IEMANN
AND 2OGERS /THER STUDIES HAVE TESTED THE .ATURAL %NEMIES (YPOTHESIS BY
COMPARING THE IMPACT OF NATURAL ENEMIES ON EXOTIC AND NATIVE SPECIES WITHIN THE
INTRODUCED RANGE )N A REVIEW OF STUDIES +EANE AND #RAWLEY FOUND
SOME CASES IN WHICH GENERALIST HERBIVORES SHOWED HIGHER IMPACTS ON NATIVE
SPECIES THAN NON
NATIVES AND OTHERS CHARACTERIZED BY THE REVERSE PATTERN
4HERE HAVE BEEN SURPRISINGLY FEW STUDIES WITH PATHOGENS IN ONE TEST 'OERGEN AND
$AEHLER FOUND THAT SMUT FUNGI CAUSED GREATER REPRODUCTIVE LOSS IN A NATIVE
GRASS (ETEROPOGON CONTORTUS THAN AN INTRODUCED GRASS 0ENNISETUM SETACEUM
%NEMY REMOVAL EXPERIMENTS ARE AN IMPORTANT BUT UNDERUTILIZED TOOL IN COM
PARING THE lTNESS EFFECT OF PATHOGENS OR HERBIVORES ON EXOTIC AND NATIVE SPECIES
WITHIN THE INTRODUCED RANGE +EANE AND #RAWLEY "LANEY AND +OTANEN
USED A FUNGICIDE EXPERIMENT TO REMOVE THE EFFECTS OF SOILBORNE FUNGI AND
OOMYCETES ON THE SURVIVAL OF SEEDS OF NATIVE VS INTRODUCED PLANTS IN TWO HABITATS
4HEY FOUND NO SUPPORT FOR A RELEASE FROM FUNGAL PATHOGENS IN THE SEED BANK OF
EXOTIC SPECIES COMPARED WITH NATIVE SPECIES -ORE RECENTLY 0ARKER AND 'ILBERT
UNPUBLISHED DATA FOUND NO DIFFERENCE IN FREQUENCY OF INFECTION LEAF DAMAGE
lTNESS EFFECTS OF FOLIAR AND DAMPING
OFF PATHOGENS OR PATHOGEN DIVERSITY BETWEEN
0LANT POPULATIONS BY PATHOGENS
SYMPATRIC SUITES OF NATIVE AND NON
NATIVE CLOVERS !LTHOUGH FUNGAL EXCLUSION
EXPERIMENTS IN THIS SYSTEM REVEALED SIGNIlCANT IMPACTS OF PATHOGENS THERE WAS
NO DIFFERENCE IN THE RESPONSE OF NATIVE AND NON
NATIVE SPECIES
&INALLY OTHER STUDIES HAVE TAKEN THE APPROACH OF COMPARING DISEASE ON A
SINGLE HOST SPECIES IN ITS NATIVE AND INVADED RANGES 7OLFE SURVEYED FOR
THE ANTHER SMUT FUNGUS -ICROBOTRYUM VIOLACEUM IN NATIVE POPULATIONS AND
INTRODUCED POPULATIONS OF 3ILENE LATIFOLIUM (E FOUND SIGNIlCANTLY MORE POPU
LATIONS INFECTED AND AT MUCH HIGHER INFECTION FREQUENCIES IN THE NATIVE RANGE
THAN IN THE INTRODUCED RANGE -ITCHELL AND 0OWER USED PUBLISHED RECORDS
OF PATHOGEN ASSOCIATIONS WITH PLANT HOSTS IN THEIR NATIVE AND INTRODUCED
RANGES 0LANTS WERE INFECTED BY FEWER PATHOGEN SPECIES IN THEIR NATURALIZED
RANGE 4HERE WAS ALSO AN INDICATION THAT SPECIES LEAVING BEHIND PROPORTIONALLY
MORE NATURAL ENEMIES WERE MORE LIKELY TO BE CATEGORIZED AS NOXIOUS OR INVASIVE
SPECIES BY LAND MANAGERS AND PUBLIC AGENCIES 5SING AN EXPERIMENTAL APPROACH
"ECKSTEAD AND 0ARKER DIRECTLY MEASURED THE DEMOGRAPHIC EFFECT OF PATHO
GENS ON AN INVADER IN THE CONTEXT OF KNOWN INFORMATION FROM THE SPECIES NATIVE
RANGE !MMOPHILA ARENARIA IN ITS NATIVE %UROPE IS LIMITED TO AN EARLY
SUCCESSIONAL
ROLE IN SHIFTING BEACH SANDS BY SOIL
BORNE PATHOGENS 6AN DER 0UTTEN ET AL
!S AN INVADER ON THE WEST COAST OF THE 53! !MMOPHILA REMAINS DOMINANT FOR
LONG PERIODS OF TIME (OWEVER ESCAPE FROM NATURAL ENEMIES DOES NOT EXPLAIN THIS
CONTRAST BETWEEN ITS NATIVE AND INVASIVE ECOLOGICAL ROLES "Y REPLICATING EXPERI
MENTS DONE IN THE NATIVE RANGE "ECKSTEAD AND 0ARKER FOUND THE NEGATIVE
EFFECT OF SOILBORNE PATHOGENS ON EARLY GROWTH IN THE INVADED RANGE WAS AT LEAST AS
LARGE OR LARGER THAN THEIR EFFECT IN THE NATIVE RANGE
4O PREDICT WHETHER AN INTRODUCED PLANT IS LIKELY TO BENElT FROM ESCAPING
NATURAL ENEMIES WE NEED TO UNDERSTAND THE RELATIVE IMPORTANCE OF HOST
SPECIALIST
VS HOST
GENERALIST PATHOGENS AND PESTS IN THE INVADED HABITAT -ARON AND 6ILA
+EANE AND #RAWLEY )F SPECIALIST PATHOGENS PREDOMINATE AND HOST
SHIFTS ARE RARE NATIVE PLANTS MAY BE SUPPRESSED MORE THAN COMPETING INVASIVE
SPECIES /N THE OTHER HAND IF GENERALIST PATHOGENS DOMINATE IN A SITE AND DO NOT
SHOW A PREFERENCE FOR NATIVE HOST SPECIES ONE WOULD NOT EXPECT AN INTRODUCED
PLANT TO EXPERIENCE RELEASE 4HEREFORE THE WIDE RANGE OF RESULTS SEEN IN THE ABOVE
EMPIRICAL STUDIES MAY BE IN PART EXPLAINED BY THE RELATIVE IMPORTANCE OF SPECIAL
IST AND GENERALIST NATURAL ENEMIES ! GREATER UNDERSTANDING OF THE PHYLOGENETIC
STRUCTURE OF PATHOGEN HOST RANGES COUPLED WITH ANALYSIS OF THE PHYLOGENETIC
STRUCTURE OF NATURAL PLANT COMMUNITIES 7EBB ET AL MAY HELP PREDICT THE
RELATIVE IMPORTANCE OF SPECIALIST VS GENERALIST PATHOGENS IN DIFFERENT KINDS OF
PLANT COMMUNITIES &OR INSTANCE IN A HIGH
DIVERSITY LOWLAND TROPICAL RAINFOREST
TREE SPECIES HOST GENERALISTS DOMINATED THE POLYPORE FUNGAL COMMUNITY
ALL OF THE MORE COMMON FUNGAL SPECIES WERE FOUND ON MULTIPLE FAMILIES OF HOST
TREES 'ILBERT ET AL &ERRER AND 'ILBERT )N CONTRAST IN A NEARBY LOW
DIVERSITY MANGROVE FOREST WITH ONLY THREE TREE SPECIES PRESENT EACH FROM A DIFFER
ENT FAMILY OF ALL POLYPORE FUNGAL COLLECTIONS BELONGED TO JUST THREE FUNGAL
SPECIES AND EACH SPECIES WAS HIGHLY SPECIALIZED ON JUST ONE MANGROVE SPECIES
'3 'ILBERT AND )- 0ARKER
'ILBERT AND 3OUSA 2ESEARCHERS ARE JUST NOW BEGINNING TO ADDRESS THE
RANGE OF HOST SPECIALIZATION IN DIFFERENT PLANT COMMUNITIES AND THE WIDE RANGE OF
OUTCOMES SUGGESTS THAT MANY MORE STUDIES WILL BE NEEDED BEFORE WE CAN FORMU
LATE PREDICTIVE GENERALIZATIONS
).4%.4)/.!,,9 ).42/$5#%$ 0!4(/'%.3 &/2 ")/,/')#!, #/.42/,
! DIRECT APPLICATION OF THE .ATURAL %NEMIES (YPOTHESIS IS DEPLOYING NATURAL
ENEMIES FROM THE NATIVE RANGE OF AN INVASIVE PLANT TO CONTROL THE INVADER POPU
LATION 3UCH CLASSICAL BIOLOGICAL CONTROL USES PLANTS AND PATHOGENS WITH SHARED
EVOLUTIONARY HISTORIES BUT A NEW ENVIRONMENTAL CONTEXT 4HERE ARE SEVERAL WAYS
IN WHICH BIOLOGICAL CONTROL INTERACTIONS MAY BE DIFFERENT FROM NATIVE PATHOGENS
ATTACKING INTRODUCED HOSTS &IRST UNLIKE NATIVE PATHOGENS WHICH ARE PRESUMABLY
ADAPTED TO THE LOCAL CLIMATE THE INTRODUCED BIOLOGICAL CONTROL AGENT EXPERIENCES
A NOVEL ENVIRONMENT WHICH COULD HAVE A LARGE IMPACT ON DISEASE DEVELOPMENT
#ASE STUDIES OF FAILED BIOLOGICAL CONTROL EFFORTS PROVIDE US WITH MANY EXAMPLES OF
THE IMPORTANCE OF THE DISEASE TRIANGLE -ORIN ET AL 3ECOND BOTH THE HOST
AND PATHOGEN ARE LIKELY TO BE GENETICALLY DEPAUPERATE (OWEVER THE PATHOGEN
WILL HAVE BEEN CHOSEN SPECIlCALLY TO BE VIRULENT ON THE INVASIVE HOST PLACING
THE HOST AT A RELATIVE DISADVANTAGE FOR EVOLUTIONARY RESPONSES 4HIRD ONLY FAIRLY
HOST
SPECIlC PATHOGENS ARE SELECTED FOR BIOLOGICAL CONTROL RELEASES MEANING THAT
PATHOGEN NUMERICAL DYNAMICS SHOULD ALWAYS BE CLOSELY LINKED TO INDIVIDUAL HOST
DENSITY )N FACT BIOLOGICAL CONTROL RELEASES ARE AN EXCELLENT OPPORTUNITY TO STUDY
FACTORS INmUENCING NUMERICAL DYNAMICS .OT ONLY SHOULD THERE BE A TIGHT CONNEC
TION BETWEEN PATHOGEN AND HOST DENSITY BUT INITIAL CONDITIONS OF THE INTERACTION
ARE WELL KNOWN 4HAT IS THE HOST POPULATION IS ORIGINALLY FREE OF THAT PATHOGEN
AND IS USUALLY AT HIGH DENSITY ! SUCCESSFUL EPIDEMIC PROVIDES AN OPPORTUNITY TO
QUANTIFY BOTH FREQUENCY
DEPENDENCE OF TRANSMISSION OF THE PATHOGEN AND DENSITY
DEPENDENCE AS THE HOST DENSITY DECLINES
0REDICTING THE SHORT
TERM AND LONG
TERM SUCCESS OF PARTICULAR BIOLOGICAL CON
TROL INTRODUCTIONS IS A MATTER OF OBVIOUS PRACTICAL IMPORTANCE 4O MAKE SUCH
PREDICTIONS WE NEED TO UNDERSTAND HOW THE NUMERICAL DYNAMICS OF A HOST PLANT
FOLLOWING INTRODUCTION OF ITS BIOCONTROL AGENT DEPEND ON HOST DENSITY DISEASE
INCIDENCE GENETIC VARIATION AND EVOLUTIONARY CHANGES IN VIRULENCE OR RESISTANCE
3URPRISINGLY WHILE THERE ARE SOME CASES FOR WHICH WE HAVE GOOD INFORMATION ON
THE DYNAMICS OF HOST NUMBERS AFTER RELEASE OF A CONTROL AGENT EG (ASAN AND
!YRES -ORRIS FOR MANY OTHER RELEASES THE DETAILS OF CHANGES ARE NOT
WELL DOCUMENTED )N PARTICULAR WE SHOULD ASK I IS CONTROL MORE SUCCESSFUL IN
GENETICALLY DEPAUPERATE WEEDS II $O TRANSMISSION RATE AND DEMOGRAPHIC IMPACT
OF THE PATHOGEN ATTENUATE AS THE HOST POPULATION DECLINES III $O PATHOGEN AND
HOST REACH A STABLE EQUILIBRIUM OR ARE THEY DEPENDENT ON METAPOPULATION DYNAM
ICS TO PERSIST IN THE LANDSCAPE $ETAILED INFORMATION ON NUMERICAL DYNAMICS IN
BIOLOGICAL CONTROL SYSTEMS IS SCARCE BUT DATA ARE NEARLY NONEXISTENT FOR LONG
TERM
0LANT POPULATIONS BY PATHOGENS
GENETIC CHANGES IN THE HOST OR PATHOGEN 4HERE IS GREAT UNTAPPED POTENTIAL FOR
BIOLOGICAL CONTROL INTRODUCTIONS TO BE USED TO UNDERSTAND THE FACTORS THAT DRIVE
THE ECOLOGICAL END EVOLUTIONARY DYNAMICS OF THE PLANT
PATHOGEN INTERACTIONS
)N FACT THE ONLY BIOCONTROL STUDY WE FOUND THAT TRACKED CHANGES IN PATHOGEN
VIRULENCE OR HOST RESISTANCE FOR A PLANT
PATHOGEN SYSTEM WAS FOR THE RUST 0UCCINIA
CHONDRILLINA ON #HONDRILLA JUNCEA SEE CASE STUDY BELOW
%VOLUTIONARY ECOLOGY OF BIOCONTROL WITH PATHOGENS
4HE EVOLUTIONARY DYNAMICS OF INTERACTIONS BETWEEN WEEDS AND PATHOGEN BIOCON
TROL AGENTS HAVE IMPORTANT IMPLICATIONS FOR THE LONG
TERM SUCCESS OF BIOLOGICAL
CONTROL PROGRAMS 'ENERALLY WE NEED TO KNOW I HAS VIRULENCE OF THE BIOLOGICAL
CONTROL PATHOGEN CHANGED OVER TIME AND HAS THIS INCREASED OR DECREASED THE
SUCCESS OF CONTROL II (AS THE HOST DEVELOPED RESISTANCE OVER TIME III (AS HOST
SPECIlCITY CHANGED OVER TIME 2ECENT ANALYSIS OF EMERGING DISEASES HAS SUGGESTED
THAT ECOLOGICAL HOST SHIFTS IE HAVING A PREADAPTED ABILITY TO USE A NEWLY ENCOUN
TERED HOST MAY PREDOMINATE AS CAUSES OF NOVEL EPIDEMICS AND THAT HOST SHIFTS MAY
ONLY RARELY BE CAUSED BY MUTATIONS THAT ALLOW COLONIZATION OF A NEW HOST 3CHRAG
AND 7IENER (OWEVER THE DIFlCULTY OF OBSERVING SUCH GENETIC EVENTS MAY
DISTORT OUR PERSPECTIVE 6AN +LINKEN AND %DWARDS SYNTHESIZED INFORMATION
ON HOST RANGE FROM BIOLOGICAL CONTROL PROGRAMS USING HERBIVORES 4HEY FOUND
THAT HOST SHIFTS WERE MORE QUANTITATIVE THAN QUALITATIVE THAT IS WHILE PREFERENCE
AND EFlCIENCY ON NOVEL HOSTS EVOLVED THERE WAS NO EVIDENCE OF EVOLUTIONARY
CHANGES IN FUNDAMENTAL HOST
RANGE 3UCH AN ANALYSIS SHOULD BE DONE FOR PATHO
GENS USED AS BIOLOGICAL CONTROL AGENTS 7HILE PATHOGENS ARE GENERALLY THOUGHT TO
OFFER THE OPPORTUNITY FOR HIGH HOST SPECIlCITY SOME HAVE ARGUED THAT HIGH HOST
SPECIlCITY MAY BE CORRELATED WITH EVOLUTIONARY LABILITY "ROOKS AND -C,ENNAN
3ECORD AND +AREIVA "ECAUSE THEY ARE UNABLE TO SIMPLY MOVE
FROM AN UNACCEPTABLE HOST TO A MORE SUITABLE ONE AS AN ANIMAL MIGHT SPECIALIST
PATHOGENS MAY EXPERIENCE EVEN STRONGER SELECTION FOR HOST SHIFTS THAN HERBIVORES
2OY +NOWING THE FREQUENCY OF EVOLUTIONARY HOST SHIFTS IN PATHOGENS AND
UNDERSTANDING THE CONDITIONS UNDER WHICH THEY OCCUR ARE CRITICAL TO THE PROCESS
OF RISK ASSESSMENT IN BIOLOGICAL CONTROL 3ECORD AND +AREIVA
'ENERAL THEORY OF HOST
PATHOGEN INTERACTIONS HAS PLAYED A LARGE ROLE IN THE
CHOICE OF BIOLOGICAL CONTROL AGENTS IN THE PAST -C&ADYEN &OR EXAMPLE
IT HAS LONG BEEN THOUGHT THAT SEXUALLY REPRODUCING WEEDS WOULD BE HARDER TO
CONTROL BECAUSE THEIR HIGHER LEVELS OF GENETIC VARIATION INTERFERE WITH PATHOGEN
POPULATION GROWTH "URDON ET AL HOWEVER A MORE RECENT ANALYSIS HAS
DISPUTED THIS ASSERTION #HABOUDEZ AND 3HEPPARD 3IMILARLY THE BELIEF
THAT PATHOGEN POPULATIONS ARE LOCALLY ADAPTED TO THEIR HOST GENOTYPES HAS HAD A
LARGE INmUENCE ON THE PROCESS OF SELECTION OF CONTROL AGENTS WITH GENETIC ANALYSIS
PLAYING AN INCREASING ROLE IN THE CAREFUL MATCHING OF AGENT GENOTYPES WITH THE
POPULATIONS OF ORIGIN FOR THE WEED EG (OLDEN AND -AHLBERG (OWEVER
THE EVIDENCE FOR CLOSE LOCAL ADAPTATION OF PATHOGENS TO THEIR HOST POPULATIONS IS
'3 'ILBERT AND )- 0ARKER
MIXED AND IN FACT RESISTANCE AND GENE FOR GENE VIRULENCE SHOULD mUCTUATE IN AN
ASYNCHRONOUS FREQUENCY DEPENDENT WAY REVIEWED IN 0ARKER AND 'ILBERT
"IOLOGICAL CONTROL PRACTITIONERS HAVE CLEARLY BASED THEIR INTRODUCTION STRATEGIES ON
THEORETICAL CONSIDERATIONS BUT THE SIMPLER RULES ARE NOW COMING UNDER QUESTION
-C&ADYEN 4HERE IS A NEED FOR CLEAR PREDICTIONS AND MODERN EMPIRICAL
WORK TESTING THOSE PREDICTIONS TO HELP INFORM THE PRACTICE OF BIOLOGICAL CONTROL
FOR THE FUTURE
!N EVOLUTIONARY ECOLOGY CASE STUDY #HONDRILLA AND 0UCCINIA
ON THREE CONTINENTS
.O SYSTEM OF CLASSICAL BIOLOGICAL CONTROL OF A WEED BY A PATHOGEN DEMONSTRATES
THE IMPORTANCE OF INTERACTIONS BETWEEN ENVIRONMENT GENETICS AND NUMERICAL
DYNAMICS AS WELL AS THAT OF #HONDRILLA JUNCEA RUSH SKELETONWEED !STERACEAE
.ATIVE TO %URASIA THIS SPECIES WAS INTRODUCED TO THE EASTERN 5NITED 3TATES IN THE
LATE S TO !USTRALIA IN THE EARLY S AND TO THE WESTERN 5NITED STATES IN
THE S -C6EAN 0RYOR 3CHIRMAN AND 2OBOCKER 3UPKOFF
ET AL #HONDRILLA IS A SIGNIlCANT ECONOMIC PROBLEM IN WHEAT
GROWING
REGIONS 0ANETTA AND $ODD AND IS ALSO A WIDESPREAD RANGELAND WEED
/F THREE BIOCONTROL AGENTS THAT WERE INTRODUCED 0UCCINIA CHONDRILLINA WAS THE MOST
EFFECTIVE AT REDUCING PLANT VIGOR 3UPKOFF ET AL AND WITHIN TWO DECADES OF
ITS INTRODUCTION INTO !USTRALIA AND #ALIFORNIA #HONDRILLA DENSITIES WERE REDUCED TO
THOSE TYPICAL IN ITS NATIVE RANGE 7APSHERE ET AL #ULLEN ET AL 3UPKOFF
ET AL
4HE EXTREME HOST SPECIlCITY FOUND IN THE #HONDRILLA0UCCINIA SYSTEM MAKES THIS
EXAMPLE PARTICULARLY INTERESTING (ASAN #HONDRILLA IS A TRIPLOID APOMICT
AND THUS REPRODUCTION IS CLONAL )N !USTRALIA THREE CLONAL TYPES ARE PRESENT EACH
WITH A DIFFERENT LEAF WIDTH 0 CHONDRILLINA CAUSES DISEASE ONLY ON THE NARROW
LEAF
TYPE 4HE ORIGINAL INFESTATION OF #HONDRILLA WAS PRIMARILY THIS NARROW
LEAF TYPE
BUT AFTER THE SUCCESSFUL BIOCONTROL OF THAT CLONE THE INTERMEDIATE
LEAVED CLONE HAS
SPREAD (ANLEY AND 'ROVES .OW THERE IS CONCERTED EFFORT TO INTRODUCE NEW
STRAINS OF 0UCCINIA CHONDRILLINA THAT ARE ABLE TO ATTACK AND CONTROL THE OTHER CLONES
OF #HONDRILLA (ANLEY AND 'ROVES
)N WESTERN .ORTH !MERICA THREE DIFFERENT #HONDRILLA GENOTYPES WERE FOUND
DISTINGUISHABLE BY THEIR MULTI
LOCUS ISOZYME PHENOTYPES (ASAN ET AL
TWO GENOTYPES ARE THOUGHT TO HAVE ORIGINATED IN 9UGOSLAVIA (ASAN AND $ELFOSSE
4HE WESTERN 53 GENOTYPES WERE LARGELY RESISTANT TO THE RUST STRAIN THAT
CONTROLLED THE NARROW
LEAF #HONDRILLA IN !USTRALIA SO ADDITIONAL RUST ISOLATES WERE
EVALUATED FOR USE IN BIOCONTROL 2USTS COLLECTED FROM 9UGOSLAVIA THE PUTATIVE
SITE OF ORIGIN OF THE 53 INVADERS SHOWED HIGH VIRULENCE ON SOME OF THE 53 GENO
TYPES BUT OTHER PLANTS WERE LITTLE AFFECTED )N ADDITION RUSTS FROM OTHER REGIONS
ALSO SHOWED HIGH VIRULENCE 'ENETIC MATCHING OF HOSTS BETWEEN THE NATIVE AND
INTRODUCED RANGE MAY YIELD WELL
ADAPTED BIOCONTROL PATHOGENS WHEN THE TARGET
WEED IS GENETICALLY UNIFORM BUT THIS APPROACH ASSUMES HIGH LOCAL ADAPTATION OF
0LANT POPULATIONS BY PATHOGENS
THE PATHOGEN TO THE HOST !S MENTIONED ABOVE SUCH LOCAL ADAPTATION MAY NOT BE
GENERALLY FOUND IN NATURAL POPULATIONS
3EXUAL REPRODUCTION IN BOTH THE HOST AND PATHOGEN PLAY A ROLE IN THIS STORY
4ELIOSPORES OF 0 CHONDRILLINA ONLY GERMINATE AFTER COLD WINTERS !DAMS AND ,INE
SO SEXUAL RECOMBINATION OF THE PATHOGEN OCCURS READILY ONLY IN COLDER
REGIONS SUCH AS PARTS OF EASTERN CONTINENTAL %UROPE )N THESE AREAS OF THE NATIVE
RANGE #HONDRILLA ALSO SHOWS HIGHER CLONAL DIVERSITY AND POSSIBLY DIPLOID SEXUAL
POPULATIONS #HABOUDEZ AND 3HEPPARD )NTERESTINGLY POPULATIONS OF
#HONDRILLA WITH HIGH VS LOW CLONAL DIVERSITY IN THE NATIVE RANGE APPEAR TO SUFFER
SIMILAR DEGREES OF RUST INFECTION #HABOUDEZ AND 3HEPPARD )N NORTHERN
.ORTH !MERICA WHERE COLD WINTERS ALLOW FOR SEXUAL RECOMBINATION OF THE PATHO
GEN THE EVOLUTIONARY DYNAMICS OF THE HOST
PATHOGEN INTERACTION ARE POTENTIALLY
MORE COMPLEX (ASAN ET AL )N !USTRALIA THEY ARE PARTICULARLY CONCERNED
ABOUT THE POSSIBILITY OF INTRODUCTION OF NEW SEXUAL TYPES OF #HONDRILLA WHICH
COULD RESULT IN A SITUATION WHERE THE HOST REPRODUCES SEXUALLY BUT THE PATHOGEN
CAN NOT #HABOUDEZ AND 3HEPPARD
4HIS EXAMPLE SHOWS THE IMPORTANCE OF THE hDISEASE TRIANGLEv INTERACTION
BETWEEN HOST PATHOGEN AND ENVIRONMENT IN THE DYNAMICS OF PLANT
PATHOGEN
INTERACTIONS 4HE CASE OF #HONDRILLA 0UCCINIA
CHONDRILLINA IS ONE OF THE FEW FOR
WHICH WE HAVE SUCH DETAILED INFORMATION ABOUT THE HOSTS AND THE PATHOGENS
GENETIC MAKEUP AND MORE IMPORTANTLY ABOUT HOW THE INTERACTION HAS PLAYED
OUT IN A NUMBER OF DIFFERENT REGIONS 0ATHOGEN BIOLOGICAL CONTROL CASES OFFER THE
POTENTIAL TO LEARN A GREAT DEAL ABOUT THE ROLES OF GENETICS NUMERICAL DYNAMICS
EVOLUTIONARY DYNAMICS AND ENVIRONMENTAL FACTORS IN DETERMINING THE LONG
TERM
OUTCOMES OF HOST
PATHOGEN INTERACTIONS
#/.#,53)/.3
&ROM THE STUDIES DESCRIBED ABOVE IT APPEARS THAT PATHOGENS ARE SOMETIMES IMPOR
TANT IN THE REGULATION OF NATURAL PLANT POPULATIONS MAY CONSTRAIN POPULATIONS
INTRODUCED TO NEW REGIONS AND MAY BY THEIR ABSENCE RELEASE INTRODUCED PLANTS
FROM AN IMPORTANT SOURCE OF REGULATION 7E SUGGEST TWO KEY DIRECTIONS THAT WOULD
MOST ADVANCE OUR UNDERSTANDING OF THE IMPORTANCE OF PATHOGENS IN PLANT POPULA
TION REGULATION AN INTEGRATION OF NUMERICAL AND EVOLUTIONARY DYNAMICS FOR BOTH
THE PATHOGENS AND PLANTS AND A GREATER BREADTH OF STUDIES TO INCLUDE MORE PLANT
PATHOGEN SYSTEMS
"IOLOGICAL CONTROL OF INVASIVE WEEDS OFFERS EXCITING OPPORTUNITIES TO EVALUATE THE
IMPORTANCE OF PATHOGENS IN NUMERICAL REGULATION OF PLANT POPULATIONS AND AT THE
SAME TIME TO FOLLOW GENETIC CHANGES IN PLANT AND PATHOGEN POPULATIONS #AREFUL
MONITORING FOR CHANGES IN HOST AND PATHOGEN GENOTYPES ALONG WITH NUMERI
CAL DYNAMICS SHOULD BE INTEGRAL TO ANY INTRODUCTION OF PATHOGENS FOR BIOLOGICAL
CONTROL OF INVASIVE WEEDS %QUALLY IMPORTANT WE MUST MOVE BEYOND SPINNING
NARRATIVES AND MAKING GENERAL PREDICTIONS FROM A HANDFUL OF EXAMPLES 7E HAVE
'3 'ILBERT AND )- 0ARKER
A GOOD IDEA OF THE ROLE PATHOGENS SHOULD PLAY IN PLANT POPULATION REGULATION AND
BIOLOGICAL INVASIONS 4ABLE WE NOW NEED TO COLLECT DATA FROM A DIVERSITY OF SYS
TEMS TO EVALUATE OUR PREDICTIONS (OW OFTEN AND UNDER WHAT CONDITIONS ARE PLANT
PATHOGENS SIGNIlCANT FORCES IN REGULATING WILD PLANT POPULATIONS IN THEIR NATIVE
RANGES (OW OFTEN DO INTRODUCED PLANTS FAIL TO ESTABLISH BECAUSE THEY ARE ATTACKED
BY LOCAL PATHOGENS (OW OFTEN IS ESCAPE FROM PATHOGENS A KEY TO DETERMIN
ING WHETHER AN INTRODUCED PLANT INVADES NATURAL HABITATS 4HROUGH PATHOGEN
EXCLUSION AND ADDITION EXPERIMENTS COMMON GARDENS PHYLOGENETIC ANALYSIS OF
HOST RANGES ANALYSIS OF RAPID EVOLUTIONARY CHANGES IN PLANT
PATHOGEN INTERAC
TIONS AND CAREFUL CREATIVE NATURAL HISTORY OF PLANT DISEASES WE WILL ILLUMINATE
THE ROLE OF PLANT PATHOGENS IN BIOLOGICAL INVASIONS
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