Professional Documents
Culture Documents
identification of
P. capsici
Phytophthora species.
Gloria Abad, Ph. D.
Lead Scientist
USDA/APHIS/PPUSDA/APHIS/PPQ/PHP/PSPI
P. ramorum MOLECULAR DIAGNOSTICS LABORATORY
WORKSHOP:Fighting
WORKSHOP: FightingPhytophthora:
Phytophthora:How
Howto toDetect,
Detect,Investigate,
Investigate,
andManage
and ManagePhytophthora.
Phytophthora.July
July26
262008
2008
APSCenntennial
APS CenntennialMeeting
Meeting2008
2008
Diseasescaused
Diseases causedby
byPhytophthora
Phytophthora 88spp.
88 spp.7/25/08
7/25/08
MDL
Root,Crown,
Root, Crown,Collar
CollarRots,
Rots,Stem
Stemcanker,
canker,Wilts
Wilts
P.ramorum
P. ramorum P.cinnamomi
cinnamomi
Leafand
Leaf andStem
StemBlights
Blights P.
P.infestans
infestans P.capsici
P. capsici
P. Fruit,and
Fruit, andTubers
Tubersrots
rots
P.capsici
P. capsici P.infestans
P. infestans
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL Photos: Mostly from the internet
Phytophthora pinifolia AJ. Durn, Gryzenh. & MJ Wingf. (2008)
MDL
Perfect stage
Not Known
Durn A, Gryzenhout M, Slippers B, Ahumada R, Rotella A, Flores F, Wingfield BD, Wingfield MJ. 2008. Phytophthora pinifolia
sp. nov. associated with a serious needle disease of Pinus radiata in Chile. Plant Pathology 57: 715-727.
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
Phytophthora sp. morindae Abad ZG & Nelson S
MDL
Straminipiles
Straminipiles
Phytophthora
Pythium
Fungi
Fungi
Pythiogeton
Phylogenetic
Phylogenetic
Relationships of
Relationships of
Straminipiles and
Straminipiles and
Straminipiles
Straminipiles
Fungi with
Fungi with other
other
Organisms
Organisms
SSUrRNA
rRNAEvolutionary
Evolutionary
SSU
Tree
Tree
FromVan
VandedePeer
PeeretetalalJ.J.
From
Mol. Evol, 2000 2000
Mol. Evol, 2000 2000
Fungi
Fungi
Additional characters
Additional charactersthat
thatdifferentiate
differentiatestraminipiles
straminipilesto
totrue
truefungi
fungi
Trait
Trait Oomycetes
Oomycetes Fungi
Fungi
Cellwall
Cell wallcomposition
composition -1,3and
-1,3 -1,6glucans;
and -1,6 glucans; Chitin
Chitin
cellulose
cellulose
Storagereserves
Storage reserves -1,3glucans
-1,3 glucans Glycogen
Glycogen
(Mycolaminarin)
(Mycolaminarin)
SomaticCells
Somatic Cells Coenocytic
Coenocytic Mostlyseptate
Mostly septate
Somaticnuclei
Somatic nuclei Diploid
Diploid Haploid
Haploid
Mitochondrialcristae
Mitochondrial cristae Tubular
Tubular Flat
Flat
Taxonomy of
Taxonomy of Phytophthora Species
The
The Era
Era of
of Morphological
Morphological species
species
The
The Era
Era of
of Phylogenetic
Phylogenetic species
species
A
A new
new era
era of
of Morphological/
Morphological/Phylogenetic
Phylogenetic
Species?
Species?
EVOLUTIONIN
EVOLUTION INTHE
THETAXONOMY
TAXONOMYPHYTOPHTHORA
PHYTOPHTHORA
THE CLASICAL
THE CLASICAL ERA
ERA
AND THE
AND THE
MORPHOLOGICAL-SPECIES
MORPHOLOGICAL-SPECIES
1500s 1700s
Whichwas
Which wasthe
thecause
cause
ofofthe
thedisease?
disease?
ThePotato
The Potato
Disease
Disease
Wherethe
Where the
Diseasecame
Disease came
From?
From?
1840s
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
MDL
POTATODISEASE
POTATO DISEASECAUSE
CAUSE1840S
1840S
ANDTHE
AND THEPIONEERS
PIONEERSOF
OF
PLANTPATHOLOGY
PLANT PATHOLOGY
Anton
AntondedeBary
Bary1861,
1861,1876
1876
Researchers
Researchersinto
intothe
thenature
natureofofthe
thepotato
potatofungus
Rev.
Rev.Berkeley,
Berkeley,1846
1846 Phytophthora infestans. J. R. Agric. Soc.
fungus
England
Phytophthora infestans. J. R. Agric. Soc. England
The
ThePotato
PotatoMurrian
Murrian 12:239-269
12:239-269
MexicanTheory
Mexican Theory
AndeanTheory
Andean Theory USA1842
USA 1842and
andUSA
USAtoto
Europe1845
Europe 1845
USA&&Europe
Europe1841-1843
1841-1843 Postulatedby
Postulated byRedick
Redick1928
1928
USA LinesofofEvidence:
Evidence:
Suggestedby:
Suggested by:Berkeley
Berkeleyand
andDeDe Lines
Bary Neiderhauser.
Neiderhauser.
Bary Supporters:Fry,
Fry,Goodwin,
Goodwin,
LinesofofEvidence:
Lines Evidence:Abad
AbadG.G. Supporters:
1983,Abad
AbadG. G.and
andAbad
AbadJ.J.1995
1995 Grunwland
Grunwland
1983,
Supportedby:
Supported by:Ristaino,
Ristaino,
Morales,Torres,
Morales, Torres,Forbes,
Forbes,
Restrepo
Restrepo
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
MDL
Phytophthora Morphological
Phytophthora Morphological Taxonomy
Taxonomy
Rosenbaum
RosenbaumJ.J.1917.
1917.Studies
StudiesofofthetheGenus
GenusPhytophthora
Phytophthora. Proceedingsofofthe
. Proceedings theNational
National
AcademyofofSciences
Academy SciencesofofthetheUnited
UnitedStates America3:3:159
StatesofofAmerica 159-163.
-163.
Tucker,
Tucker,CM CM1931.
1931.Taxonomy
Taxonomyofofthe thegenus
genusPhytophthora
Phytophthorade deBary
Bary. Univ.Missouri
. Univ. MissouriAgric.
Agric.
Exp.Sta.
Exp. Sta.Bull.
Bull.153:
153:11-208
-208
Leonian
Leonian, LH.1925.
, LH. 1925.Physiological
Physiologicalstudies
studieson
onthethegenus
genusPhytophthora
Phytophthora. West.Virginia
. West. Virginia
Agric.Expt.
Agric. Expt.Sta.
Sta.Sci.
Sci.Paper
Paper1111: :444
444-498.
-498.
Leonian
Leonian, LH1934.
, LH 1934.Identification
IdentificationofofPhytophthora species.West
Phytophthoraspecies. WestVirginia
VirginiaAgriculture
Agriculture
ExperimentStation
Experiment StationBulletin
Bulletin262:
262:22-36
-36
Frezzi
Frezzi, MJ.1950.
, MJ. 1950.LasLasespecies
especiesdedePhytophthora
Phytophthoraen enlalaArgentina,
Argentina,Revista
Revista
deInvestigaciones
de InvestigacionesAgr Agrcolas IV:47
colas IV: 47134.
134.
Waterhouse
WaterhouseGM GM1963.
1963.KeyKeytotothe
thespecies
speciesofofPhytophthora
Phytophthorade deBary
Bary. MycolPap
. Mycol Pap92:
92:2222
Waterhouse
WaterhouseGM GM1970.
1970.TheThegenus
genusPhytophthora
Phytophthorade deBary
Bary. MycolPap
. Mycol Pap122:
122:1159
59
Newhook
NewhookFJ, FJ,Waterhouse
WaterhouseGM, GM,Stamps
StampsDJ.DJ.1978.
1978.Tabular
Tabularkey keytotothe
thespecies
speciesof
of
Phytophthorade
Phytophthora deBary
Bary. Mycol.Pap.
. Mycol. Pap.43.
43.
Stamps
StampsDJ,DJ,Waterhouse
WaterhouseGM, GM,Newhook
NewhookFJ, FJ,Hall
HallGS.
GS.1990.
1990.Revised
Revisedtabular
tabularkey
keytotothe
the
speciesofofPhytophthora
species Phytophthora. MycolPap
. Mycol Pap162:
162:2828pp.
pp.
Erwin
ErwinO,O,Ribeiro
Ribeiro. .1996.
1996.Phytophthora DiseasesWorldwide.
PhytophthoraDiseases Worldwide.APS. APS.Press.
Press.St.
St.Paul,
Paul,MN.
MN.
Gallegly
GalleglyME,ME,Hong
HongC. C.2008.
2008.Phytophthora
Phytophthora: IdentifyingSpecies
: Identifying SpeciesbybyMorphology
Morphologyand and
DNA.APS.
DNA. APS.Press.
Press.St.
St.Paul,
Paul,MN.
MN.
Newhook,Waterhouse,
Newhook, Waterhouse,and andStamps.
Stamps.1978
1978 Erwinand
Erwin andRibeiro.
Ribeiro.1996.
1996.Phytophthora Diseases
PhytophthoraDiseases
TabularKey
Tabular Keytotothe
theSpecies
SpeciesofofPhytophthora
Phytophthora Worldwide.APS.
Worldwide. APS.Press.
Press.
DeBary.
De Bary.Commonwealth
CommonwealthMycological
MycologicalIntitute,
Intitute,
KewSurrey,
Kew Surrey,England
England
The64
The 64Phytophthora morphospeciespresented
Phytophthoramorphospecies presentedatatErwin
Erwinand
andRibeiros
RibeirosBook
Book(1996)
(1996)
UNCULTURABLE SPECIES
hyphalswellings
hyphal swellings
Phytophthora ramorum
Sexualphase
Sexual phase
Temperature
Temperature
P.cambivora growthatat20,
growth 20,
25,30,
25, 30,35C
35C
oospore
oospore
G.Abad.PPIL.NCSU
Colonymorphology
Colony morphology
oogonium
oogonium antheridium
antheridium P. sp. morindae
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL Photos by G. Abad
MORPHOLOLGYOF
MORPHOLOLGY OFSPORANGIOPHORES
SPORANGIOPHORES
MDL
Phytophthora sp.
P. bohemeria P. cactorum aucubae P. sp.niederhauserii P.
P. capsici
capsici
P.drechsleri
P. sp. personii
P. sp. niederhauserii
P. sp. kelmania
P
P A1 x A2
P. cactorum P. heveae HO P. citricola
i.e P.boehmeriae, P.botryosa, P.capsici, i.e P.cyperi, P.cyperi bulbosi, P. inflata,
i.e P. clandestina, P. idaei, P.iranica,
P.italica, P.pseudotsugae, P.tentaculata P.pseudotsugae, P.tentaculata, P. meadii, 6 P. lepironiae, P. porri, P. primulae, 2
A1 x A2
A1 x A2
P. infestans P. megasperma P. cryptogea
i.e P.cambivora, P. cinnamomi,
i.e P. hibernalis HO, P.ilicis HO, P. mirabilis, i.e. P.fragariae v. fragariae, v. rubi P. humicola
P. drechsleri,
P. phaseoli HO, P. colocasiae, P. eriugena, HO P.insolita, P. lateralis, P. medicaginis, P. sojae, 3 P.erythroseptica HO, P. gonapodyides
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL Photos by G. Abad
MDL Problemsin
Problems inmorphological
morphologicalID
ID of
ofPhytophthora
Phytophthora Species
Species
EVOLUTIONIN
EVOLUTION INTHE
THETAXONOMY
TAXONOMYPHYTOPHTHORA
PHYTOPHTHORA
THE MODERN
THE MODERN ERA
ERA
AND THE
AND THE
PHYLOGENETIC -- SPECIES
PHYLOGENETIC SPECIES
Whiteetetal.
White al.1990,
1990, Primers
PrimersITS
ITS1,2,3,4,5
1,2,3,4,5 Leeand
andTaylor
Taylor1992-
1992-1993
1993
Lee
Firstphylogeny
First phylogeny
Cookeetetalal1996
Cooke 1996&&1997
1997
GroupIIpapillated
Group papillatedHO,
HO,RFLP
RFLP
Cooke-et
Cooke- etal
al2000.
2000.phylogeny
phylogeny
Martinand
Martin andTooley
Tooley2003
2003
mtCox
mt CoxIIsequencing
sequencingphylogeny
phylogeny
Kroonetetal.
Kroon al.2004
2004
Tef
Tef
mtCox
mt CoxIIII
mtnad11
mt nad11
-tubulin
- tubulin
JaimeBlair
Jaime Blairand
andMichael
MichaelCoffey.
Coffey.2007.
2007.Seven
SevenNuclear
Nuclearand
andMitochandrial
MitochandrialMarkers
Markers
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
MDL
ITS TEF 1 and - Tubulin
CL1
CL2
CL3
CL4
CL5
CL6
CL7
CL8
Cookeetetal.
Cooke al.2000.
2000.Fungal
FungalGenetics
Genetics
AndBiology
Biology30,
30,17-32
17-32 Kroonetetal.
Kroon al.2000.
2000.Fungal
FungalGenetics
Genetics
And
AndBiology
And Biology41,
41,766-782
766-782
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL Photos by G. Abad
MDL
AAgenus-wide
genus-widephylogeny
phylogenyfor
forPhytophthora
Phytophthorausing
usingseven
sevennuclear
nuclearloci
loci
1998
P. multivesiculata Ilieva, Man int Veld, Veenbaas-Rijks & Pieters
1999
P. quercina Jung
2001 2005
P. tropicalis Aragaki & Uchida P. kernoviae Brasier, Beales & Kirk
P. oryzo-bladis (Wang and Lu) Ho 2006
P. pistaceae Mirabolfathy P. captiosa Dick & Dobbie
P. polygony Zheng & Ho P. fallax Dick & Dobbie
P. ramorum Werres, de Cock & Man Int veld P.foliorum Donahoo & Lamour
2002 P.polonica Belbahri, Melgarejo & Leford
P. europaea Hansen & Jung 2007
P. psichrophila Jung & Hansen P. alticola Maseko, Coutinho & MJ Wingf.
P. uliginosa Jung & Hansen P. austrocedrae Gresl. & Hansen
P. brassicae de Cock and Man int Veld P. frigida Maseko, Coutinho & MJ Wingf. (2007)
P. ipomoeae Flier & Grnwald P. siskiyouensis Reeser & EM Hansen (2007)
2003 2008
P. inundata, Brasier, Sanchez-Hernadez & Kirk P. bisheria Abad ZG, Abad JA & Louws
P. pseudosyringae Jung & Delatour P. gallica T Jung T & J Nechwatal
2004 P. irrigata C Hong & M Gallegly (2008)
P. alni Brasier & Kirk P. parsiana Mostowfizadeh, Cooke &
P. inundata, Brasier, Sanchez-Hernadez & Kirk Banihashemi
P. hedraiandra de Cock & Man int Veld P. pinifolia AJ. Durn, Gryzenh. & MJ Wingf.
P. polygony Zheng & Ho P. quercetorum Balci & Balci
CL1
500 Phy.quercina.IFBQUE4
ITS REGION 747
Phy.ilicis.ILI.P590.CL3.IV
Phy.nemorosa.2055.2
Phy.pseudosyringae.PSEU.12
Phy.psychrophila.IFBPSY.2 Clade 3
413 Phy.heveae.IMI180616.CL5.II
1000
Phy.katsurae.IMI360596.CL5.II
Phy.arecae.IMI348342.CL4.II
Phy.palmivora.UQ1294.CL4.II
Clade 5
982 Phy.megakarya.IMI337104.CL4.II
Phy.botryosa.IMI136915.CL2.II Clade 4
CL2
Phy.citrophthora.IMI332632.CL2
343 Phy.meadii.Zantesd
558 Phy.colocasiae.IMI368918.CL2.I
Phy.tropicalis.P4742
Clade 2
CL3
Phy.inflata.IMI342898.CL2.III
728 Phy.citricola.IMI031372.CL2.II
996 Phy.capsici.IMI352321.CL2.II
Phy.sp. glovera.IS11095
CL4
Phy.multivesiculata.CBS545.9
974
6. Phy.bisheria.Cg2.33
Phy.megaspermaII.P1276
P.megaspermaIII.P250
CL5 700
P.megaspermaIV.P1279
Phy.megasperma.IMI133317.CL6.V
P.megaspermaV.P1280
Phy.gonapodyides.P1047
Phytophthora.sp.P61
CL6 Phytophthora.sp.P47
Phytophthora.sp.P51
Phytophthora.Pgchlamydo.P510
Phytophthora.sp.P79
326 Phytophthora.sp.P1044
Phytophthora.sp.Raspberry.P896
Phytophthora.sp.P1049
802 Phytophthora.sp.P1050
Clade 6
CL7 1000
Phytophthora.sp.cranberry.1410
976 Phytophthora.sp.P1054
Phytophthora.Salix.soil.P878
Phytophthora.sp.KACC40449
988 Phytophthora.sp.walnut.P532
831 Phytophthora.sp.apple.cherry.P
1000 Phy.humicola.IMI302303.CL6.V
Phy.inundata.P1052
Phytophthora.sp.asparagus.VPRI
Cookeetetal.
al.2000.
2000.Fungal
FungalGenetics
Genetics
Phy.erythroseptica.ATCC36302.C
707 Phy.cryptogea.IMI045168.CL8.VI
Cooke P.medicaginis.x.cryptogea.P74
Phy.drechsleri.ATCC46724.CL8.V
Phy.medicaginis.UQ125.CL8.V
AndBiology
And Biology30,
30,17-32
17-32 1000
966 Phy.trifolii.UQ2143.CL8.V
Phy.porri.CBS.567.86.CL8.III
1000 Phy.primulae.CBS620.97.CL8.III Clade 8
576 Phy.brassicae.CBS782.97
864 Phy.syringae.IMI296829.CL8.III
Phy.ramorum.cc2467
Phy.ramorum.20315247
1000 Phy.lateralis.IMI040503.CL8.V
Phy.hibernalis.ATCC.60352
Phy.avicenniae.CBS188.85
1000
Phy.macrochlamydospora.IMI1832
Phy.polymorphica.CBS.680.84 NO C
904 Phy.richardiae.IMI340618
Phy.kernoviae.P1571
Phytophthora.sp.C19.AY659738
Clade 9
Phytophthora.sp.Rf17
Phytophthora.sp.Rf6
1000 Phytophthora.sp.SCRP237
Phytophthora.sp.C25 NO C
0.1
Phy.insolita.IMI288805
Clade 10
982
1000
939
P.sp. personii
P.personii.P31.51.5.G24.Tobacc
P.personii.P31.5.6.P56.X5.Toba
P.personii.P31.51.8.P38-XS.Tob
962Phy.humicola.IMI302303.CL6.V
Phy.inundata.P1052
Phytophthora.sp.KACC40449
925 Phytophthora.sp.walnut.P532
Phytophthora.sp.apple.cherry.P
Phytophthora.sp.asparagus.VPRI
Phytophthora.sp.asparagi.UQ214
1000 Phy.alni.sp.uniformis.P1375
Phy.cambivora.IMI296524.CL7.VI
Phy.alni.sp.Alni.P772
Phy.fragariae.v.fragariae.FVF=
Phy.fragariae.v.rubi.FVR=03.CL
P. sp. personii
Phy.alni.sp.Multiformis.P1256
Phy.europaea.IFBEUR2
828 Phy.uliginosa.IFBULI.1
Phy.cinnamomi.DQ173250.001.Avo
P.cinnamomi.Ph131.1A
P.cinnamomi.05.728.Blue.Holly
Phy.cinnamomi.UQ881.CL7.VI
P.niederhauserii.Cistus.Califo
P.niederhauseria.01.5112.Arvor
1000
P.sp. niederhauserii
P.niederhauseria.01.6056.Engli
P.niederhauserii.32023.Austral
P.niederhauserii.32086.Austral
P.niederhauserii.32022.Austral
Phy.cajani.P536.CL7.VI
539 Phy.vignae.UQ136.CL7.VI
506 Phy.melonis.IMI325917
Phy.sinensis.ATCC46538.CL7.V
946 Phy.pistaciae.PIS15
Phy.sojae.UQ1200.CL7.V
Phytophthora.03.2633
P.kelmania.Ph148.G311
P.kelmania.01.5341.F.Fraser.Fi
172 P.kelmania.02.119.A.G41 P. sp.kelmania
P.kelmania.01.5341.N.North.Fir
P.kelmania.02.200.F.Fir.
Phy.cryptogea.IMI045168.CL8.VI
Phy.erythroseptica.ATCC36302.C
640
Phy.drechsleri.ATCC46724.CL8.V
P.medicaginis.x.cryptogea.P74
Phy.medicaginis.UQ125.CL8.V
Phy.trifolii.UQ2143.CL8.V
P. sp. niederhauserii
473 Phy.porri.CBS.567.86.CL8.III
908 1000 Phy.primulae.CBS620.97.CL8.III
Phy.brassicae.CBS782.97
672 Phy.syringae.IMI296829.CL8.III
Phy.ramorum.cc2467
976 Phy.ramorum.20315247
999 Phy.lateralis.IMI040503.CL8.V
Phy.hibernalis.ATCC.60352
1000 Phy.avicenniae.CBS188.85
Phy.polymorphica.CBS.680.84
Phy.kernoviae.P1571
Phy.macrochlamydospora.IMI1832
Phy.richardiae.IMI340618
Phytophthora.sp.C19.AY659738
Phytophthora.sp.Rf17
826 Phytophthora.P9.001
Phytophthora.sp.Rf6
Phytophthora.sp.SCRP237
1000 Phytophthora.sp.C25
05.1205.Pond.H20.5.1
Phy.insolita.IM
0.1 I288805
P. sp. kelmania
INTEGRATING MORPHOLOGICAL AND MOLECULAR FOR A UNIFIED TAXONOMY OF PHYTOPHTHORA
MDL
P. cactorum
PAPILLATED
Clade 1
P. citrophthora Clade 4
G. Abad
2004 Clade 2
ITS Clade 3
Clade 6
P. cryptogea Clade 5
NON-
PAPILLATED (1)
Clade 8
Clade 7
7 DNA molecular
markers
P. cactorum
SEMIPAPILLATE
Phylo
tree
Michael
Coffey
2007 P. citricola
NON-PAPILLATE
G. Abad 2008
P. erytroseptica
P. inflata SP
C -
Pe -
Pr +
ABSENT ABSENT Ch -
Hs -
P. citricola SP
C -
Pe -
Pr -
ABSENT ABSENT Ch -
Hs -
30-60 x 25-35 um
P. glovera 11099-23 SP
C -
Pe -
Pr -
ABSENT ABSENT Ch -
Hs -
INTEGRATED
INTEGRATED
MORPHOLOGICAL/PHYLOGENTIC
MORPHOLOGICAL/PHYLOGENTIC
KEY-2007
KEY-2007
Phytophthoraspecies
Phytophthora species
inCLADE
in CLADE22
In Progress by ZG Abad
In Progress by ZG Abad
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
CENTREFOR
CENTRE FORBIOLOGICAL
BIOLOGICALINFORMATION
INFORMATIONTECHNOLOGY
TECHNOLOGY
MDL THEUNIVERSITY
THE UNIVERSITYOF
OFQUENSLAND
QUENSLAND--AUSTRALIA
AUSTRALIA
LUCID KEY
LISENCE
OOMYCETES- PHYTOPHTHORA TO
By Gloria Abad and Michael Coffey Z.G.ABAD
2008
FUTURE SOFTWARE
PYTHIUM
PHYTOPYTHIUM nov. gen
PYTHIOGETON
EXAMPLES
EXAMPLES
LucidKeys
KeysHygrophoraceae
Hygrophoraceae Urbanbee
Urban beespecies
speciesofofthe
theSan
SanFrancisco
FranciscoBay
Bayarea
area
Lucid
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
http://www.aphis.usda.gov/plant_health/identification/phytophthora/index.shtml