1. Development of a
Morphological-Phylogenetic
Lucid Key for the
P. capsici Identification of Oomycetes:
Phytophthora
Z. G. Abad, USDA/APHIS/MDL, MD, USA
P. ramorum M. Coffey, World Oomycetes Collection, CA, USA
3rd International Phytophthora, Pythium and related genera Workshop:
3rd International Phytophthora, Pythium and related genera Workshop:
Integration of Traditional and Modern Approaches for Investigating the
Integration of Traditional and Modern Approaches for Investigating the
Taxonomy and Evolution of the Oomycetes
Taxonomy and Evolution of the Oomycetes
Turin – Italy 23 – 24, 2008
Turin – Italy 23 – 24, 2008
2. MDL
EVOLUTION IN THE TAXONOMY PHYTOPHTHORA
EVOLUTION IN THE TAXONOMY PHYTOPHTHORA
THE CLASICAL ERA
THE CLASICAL ERA
AND THE
AND THE
MORPHOLOGICAL-SPECIES
MORPHOLOGICAL-SPECIES
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
3. POTATO DISEASE CAUSE 1840´S
POTATO DISEASE CAUSE 1840´S
MDL
AND THE PIONEERS OF
AND THE PIONEERS OF
PLANT PATHOLOGY
PLANT PATHOLOGY
1841-1842 Morren,Belgium
1841-1842 Morren,Belgium 1841-1842 Teschemacker, USA
1841-1842 Teschemacker, USA
Anton de Bary 1861, 1876
Anton de Bary 1861, 1876
Researchers into the nature of the potato fungus
Rev. Berkeley, 1846
Rev. Berkeley, 1846 Researchers into the nature of the potato fungus
Phytophthora infestans. J. R. Agric. Soc. England
Phytophthora infestans. J. R. Agric. Soc. England
The Potato Murrian
The Potato Murrian 12:239-269
12:239-269
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
4. Phytophthora infestans Center of Origen current teories
Phytophthora infestans Center of Origen current teories
MDL
Mexican Theory
Mexican Theory
Andean Theory
Andean Theory USA 1842 and USA to
USA 1842 and USA to
Europe 1845
Europe 1845
USA & Europe 1841-1843 Postulated by Redick 1928
Postulated by Redick 1928
USA & Europe 1841-1843 Lines of Evidence:
Suggested by: Berkeley and De
Suggested by: Berkeley and De Lines of Evidence:
Bary Neiderhauser.
Neiderhauser.
Bary Supporters: Fry, Goodwin,
Lines of Evidence: Abad G.
Lines of Evidence: Abad G. Supporters: Fry, Goodwin,
1983, Abad G. and Abad J. 1995 Grunwland
Grunwland
1983, Abad G. and Abad J. 1995
Supported by: Ristaino,
Supported by: Ristaino,
Morales, Torres, Forbes,
Morales, Torres, Forbes,
Restrepo
Restrepo
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
5. Jones in 1912, presented the following statements in regard to this
topic, “the details of the introduction of late blight (native
upon the wild potato of the Andean regions) in the 1840s
from South America into Europe, and subsequent spread in
this area, constitute one of the most interesting chapters of
the history of plant pathology…
In 1928, Reddick – Mexican Theory
‘Lima’
‘Lima’
‘Péruviennes’
‘Péruviennes’
‘Cordiliéres’
‘Cordiliéres’
The importation from
from South America
•Independent South American
states [1820 to 1830s]. Peru
1821.
•Introduction of Guano boom
•The advent of the steamships in
the late 1830's
6. Phytophthora niederhauserii Abad ZG & Abad JA
Phytophthora niederhauserii Abad ZG & Abad JA
to honor John Niederhauser
to honor John Niederhauser
P.niederhauseria.01.5112.Arvor
P.niederhauseria.01.6056.Engli
P.niederhauserii.32023.Austral
P.niederhauserii.32086.Austral P.niederhauserii
P.niederhauserii.32022.Austral
P.niederhauserii.Cistus.Califo
P.niederhauserii.Cistus.Spain
Phy.sojae.UQ1200
1000 Phy.cajani.P536
989
Phy.vignae.UQ136I
990
ITS
Phy.melonis.IMI325917
922
1000
CLADE 7
Phy.sinensis.ATCC46538
Phy.pistaciae.PIS15
672
Phy.alni.sp.uniformis.P1375
514
Phy.cambivora.IMI296524
891
Phy.alni.sp.Alni.P772
790
Phy.alni.sp.Multiformis.P1256
437
Phy.fragariae.v.fragariae.FVF=
992
1000
1000 Phy.fragariae.v.rubi.FVR=03
Phy.europaea.IFBEUR2
848
Phy.uliginosa.IFBULI.1
Phy.cinnamomi.DQ173250.001.Avo
983
Arvorbitae, English Yvy, Leyland
P.cinnamomi.05.728.Blue.Holly
982
Cyppress Abad Z.G., Abad J.A., NC USA
P.cinnamomi.Ph131.1A
1000
Phy.cinnamomi.UQ881 Cistus , CA USA Cheryl Blomquish
Phytophthora.03.2633
Phy.insolita.IMI288805 Cistus, Italy David Cooke
0.1
Xanthorrea Australia, James Cunnigton
Cistus, Canary Island S. Moralejo
Norway M.L. Herrero,
7. MDL
Phytophthora Morphological Taxonomy
Phytophthora Morphological Taxonomy
•• Rosenbaum J. 1917. Studies of the Genus Phytophthora. Proceedings of the National
Rosenbaum J. 1917. Studies of the Genus Phytophthora. Proceedings of the National
Academy of Sciences of the United States of America 3: 159-163.
Academy of Sciences of the United States of America 3: 159-163.
•• Tucker, CM 1931. Taxonomy of the genus Phytophthora de Bary. Univ. Missouri Agric.
Tucker, CM 1931. Taxonomy of the genus Phytophthora de Bary. Univ. Missouri Agric.
Exp. Sta. Bull. 153: 1-208
Exp. Sta. Bull. 153: 1-208
•• Leonian, LH. 1925. Physiological studies on the genus Phytophthora. West. Virginia
Leonian, LH. 1925. Physiological studies on the genus Phytophthora. West. Virginia
Agric. Expt. Sta. Sci. Paper 11 : :444-498.
Agric. Expt. Sta. Sci. Paper 11 444-498.
•• Leonian, LH 1934. Identification of Phytophthora species. West Virginia Agriculture
Leonian, LH 1934. Identification of Phytophthora species. West Virginia Agriculture
Experiment Station Bulletin 262: 2-36
Experiment Station Bulletin 262: 2-36
•• Frezzi, MJ. 1950. Las especies de Phytophthora en la Argentina, Revista
Frezzi, MJ. 1950. Las especies de Phytophthora en la Argentina, Revista
de Investigaciones Agrícolas IV: 47–134.
de Investigaciones Agrícolas IV: 47–134.
Waterhouse GM 1963. Key to the species of Phytophthora de Bary. Mycol Pap 92: 22
Waterhouse GM 1963. Key to the species of Phytophthora de Bary. Mycol Pap 92: 22
Waterhouse GM 1970. The genus Phytophthora de Bary. Mycol Pap 122: 1–59
Waterhouse GM 1970. The genus Phytophthora de Bary. Mycol Pap 122: 1–59
•• Newhook FJ, Waterhouse GM, Stamps DJ. 1978. Tabular key to the species of
Newhook FJ, Waterhouse GM, Stamps DJ. 1978. Tabular key to the species of
Phytophthora de Bary. Mycol. Pap. 43.
Phytophthora de Bary. Mycol. Pap. 43.
Stamps DJ, Waterhouse GM, Newhook FJ, Hall GS. 1990. Revised tabular key to the
Stamps DJ, Waterhouse GM, Newhook FJ, Hall GS. 1990. Revised tabular key to the
species of Phytophthora. Mycol Pap 162: 28 pp.
species of Phytophthora. Mycol Pap 162: 28 pp.
Erwin O, Ribeiro . .1996. Phytophthora Diseases Worldwide. APS. Press. St. Paul, MN.
Erwin O, Ribeiro 1996. Phytophthora Diseases Worldwide. APS. Press. St. Paul, MN.
•• Gallegly ME, Hong C. 2008. Phytophthora: Identifying Species by Morphology and
Gallegly ME, Hong C. 2008. Phytophthora: Identifying Species by Morphology and
DNA. APS. Press. St. Paul, MN.
DNA. APS. Press. St. Paul, MN.
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
8. MDL
Newhook, Waterhouse, and Stamps. 1978
Newhook, Waterhouse, and Stamps. 1978 Erwin and Ribeiro. 1996. Phytophthora Diseases
Erwin and Ribeiro. 1996. Phytophthora Diseases
Tabular Key to the Species of Phytophthora
Tabular Key to the Species of Phytophthora Worldwide. APS. Press.
Worldwide. APS. Press.
De Bary. Commonwealth Mycological Intitute,
De Bary. Commonwealth Mycological Intitute,
Kew Surrey, England
Kew Surrey, England
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
9. MDL
Sixty of the important
species and taxons
New dichotomous-like key is presented using minimal morphological
characters and the Single Strand Conformational Polymorphism PCR-
SSCP DNA fingerprint of the ITS rDNA.
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
11. MORPHOLOLGY OF SPORANGIOPHORES
MORPHOLOLGY OF SPORANGIOPHORES
MDL
UNBRANCHED BRANCHED EXTERNAL INTERNAL
PROLIFERATION PROLIFERATION
P. erytroseptica P. sp. kelmania P. sp. 3 P. bisheria P. lateralis
BOTRYOSE CLUMPS UMBELLA TYPE SYMPLE SYMPODIUM LONG SYMPODIUM
P.botryosa P.capsici P.kernoviae P. ramorum
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL Photos by G. Abad
12. Sporangia shapes
MDL
Globose Globose Ovoid Ellipspoid Lemoniform
E
Phytophthora sp.
P. bohemeria P. cactorum aucubae P. sp.niederhauserii P. capsici
P. capsici
Obpyryform Constricted Tapered base Irregular Irregular
P. cryptogea P. erytroseptica P. tropicalis P. nicotianae P. nicotianae
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
13. Sporangium, papilla, exit pore and pedicel shapes
MDL
Papillate –short pedicel
pedicell Semipapillate- Semipapillate- Non-papillate –
Medium pedicel Long pedicel persistent
P. cactorum P. bohemeria P. botryosa P. capsici P. sp.niederhauserii
Exit pore
Narrow Medium Wide Two papilla
P. nicotianae P. bohemeriae P. citrophthora P. sp. personii P. capsici
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL Photos by G. Abad
14. Hyphal swellings
Hyphal swellings
MDL
P.drechsleri
P. sp. personii
P. sp. niederhauserii
P. sp. kelmania
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
P. foliorum
Photos by G. Abad
15. Chlamydosphores
Chlamydosphores
MDL
Terminal Intercalary Clustered
P. tropicalis Phytophthora sp. Juniper P.cinnamomi.
Terminal, lateral, intercalary
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL P. ramorum Photos by G. Abad
16. Phytophthora sexual phase
Phytophthora sexual phase
MDL
Antheridia
Paragynous amphigynous Paragynous/ amphigynous
P. cactorum P. capsici P. glovera
Paragynous Unusual shapes oospores
Plerotic Aplerotic
P. bisheria P.sp. aucubae P. cactorum P. capsici
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL Photos by G. Abad
17. Morphospecies Groups [Waterhouse (1963), Newhook et al. (1978), Stamps et al. 1990]
Morphospecies Groups [Waterhouse (1963), Newhook et al. (1978), Stamps et al. 1990]
MDL Mostly HO
GROUP II
GROUP GROUP III
Mostly HO Mostly HT Mostly HO
GROUP II
GROUP II Mostly HT
GROUP III Mostly HO
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
Mostly HT Mostly HO Mostly HT
GROUP IV
GROUP IV
Mostly HT GROUP V
GROUP V Mostly HO GROUP VI
GROUP VI
Mostly HT
P
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
18. MDL Problems in morphological ID of Phytophthora Species
Problems in morphological ID of Phytophthora Species
64 species described to 1994
• variability in morphological characters
• limited no. of morphological characters
• overlapping in morphological
characters
• some atypical isolates
• difficulties in esporulation
• lack of expertise for identification
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
19. MDL
EVOLUTION IN THE TAXONOMY PHYTOPHTHORA
EVOLUTION IN THE TAXONOMY PHYTOPHTHORA
THE MODERN ERA
THE MODERN ERA
AND THE
AND THE
PHYLOGENETIC -- SPECIES
PHYLOGENETIC SPECIES
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
20. Phytophthora molecular markers
Phytophthora molecular markers
MDL
White et al. 1990, Primers– ITS 1,2,3,4,5
White et al. 1990, Primers– ITS 1,2,3,4,5 Lee and Taylor 1992- 1993
Lee and Taylor 1992- 1993
First phylogeny
First phylogeny
Cooke et al 1996 & 1997
Cooke et al 1996 & 1997
Group II––papillated ––HO, RFLP
Group papillated HO, RFLP
Cooke- et al 2000. phylogeny
Cooke- et al 2000. phylogeny
Martin and Tooley 2003
Martin and Tooley 2003
mt Cox IIsequencing phylogeny
mt Cox sequencing phylogeny
Kroon et al. 2004
Kroon et al. 2004
Tef
Tef
mt Cox II
mt Cox II
mt nad11
mt nad11
β- tubulin
β- tubulin
Jaime Blair and Michael Coffey. 2007. Seven Nuclear and Mitochandrial Markers
Jaime Blair and Michael Coffey. 2007. Seven Nuclear and Mitochandrial Markers
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
21. MDL
ITS TEF α 1 and β- Tubulin
CL1
CL2
CL3
CL4
CL5
CL6
CL7
CL8
Cooke et al. 2000. Fungal Genetics
Cooke et al. 2000. Fungal Genetics
And Biology 30, 17-32 Kroon et al. 2000. Fungal Genetics
Kroon et al. 2000. Fungal Genetics
And Biology 30, 17-32
And Biology 41, 766-782
And Biology 41, 766-782
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL Photos by G. Abad
22. A genus-wide phylogeny for Phytophthora using seven nuclear loci
A genus-wide phylogeny for Phytophthora using seven nuclear loci
MDL
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
23. 24 Species of Phytophthora described after Erwin and Ribeiro (1996) 8/24/08
MDL
1998
P. multivesiculata Ilieva, Man in’t 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 In’t 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 in’t Veld P. frigida Maseko, Coutinho & MJ Wingf. (2007)
P. ipomoeae Flier & Grünwald 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 in’t Veld P. pinifolia AJ. Durán, Gryzenh. & MJ Wingf.
P. polygony Zheng & Ho P. quercetorum Balci & Balci
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
24. Phytophthora ‘taxons’ in process for official description
MDL
Many, many!! species in progress for official descriptions. Some examples….
13 species in progress for official description 16 putative new species
P. sp. andina Forbes et al. MIKE COFFEY’S GROUP
P. sp. asparagi Cooke et al P. sp. carica
P. taxon ‘cranberry’ Oudemans et al. P. sp. cuyabensis
P. sp. glovera Abad & Shew P. sp. erwinii
P. sp. kelmania Abad ZG & Abad JA P. sp. lagoariana
P. sp. morindae Abad ZG & Nelson S Phytophthora sp. novaeguinee
P. sp. niederhauserii Abad ZG & Abad JA P. sp. oregonsis
P. sp. pelgrandis Nirenberg & Gerlarch P. sp. sansomea
P. sp. personii Abad ZG et al. P. sp. sulawesiensis
P. sp. quercetorum Balci & Balci P. sp. zentmeyerii
P. sp. fragaefolia Shirai K & Shinmura CLIVE BRASIER GROUP
P. sp. hungarica Belbahri,L., Lefort,F., Many putative new species in P. megasperma
Calmin,G. and Oszako,T. complex
P. sp. sylvatica Belbahri,L., Lefort,F., DAVID COOKE (4), GLORIA ABAD (2)
Calmin,G. and Oszako,T. Six putative new species in P. insolita complex
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
25. MDL
CL1
CL2
CL3
CL4
CL5
CL6
CL7
CL8
Cooke et al. 2000. Fungal Genetics
Cooke et al. 2000. Fungal Genetics
And Biology 30, 17-32
And Biology 30, 17-32
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
29. INTEGRATING MORPHOLOGICAL AND MOLECULAR FOR A UNIFIED TAXONOMY OF PHYTOPHTHORA
MDL PAPILLATE
7 DNA molecular
markers
P. cactorum
SEMIPAPILLATE
Phylo
tree
Michael
Coffey
2007 P. citricola
NON-PAPILLATE
G. Abad 2008
P. erytroseptica
SEMIPAPILLATE
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
30. MDL
INTEGRATED
INTEGRATED
MORPHOLOGICAL/PHYLOGENTIC
MORPHOLOGICAL/PHYLOGENTIC
KEY-2008
KEY-2008
Phytophthora species
Phytophthora species
in CLADE 2
in CLADE 2
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL
33. Peronosporaceae family and some phylogenetic groups
Peronosporaceae family and some phylogenetic groups
P.vexans.03.1391.Crptmans.PPIL
Py.vexans.PT2.PPIL21.37.2
P.vexans.03.3212.C.laurel
P.vexans.03.1220.Yew
Py.vexans.CBS119.80.CLK
P.vexans.C.04.562.11980
P47.107.F7
P.vexans.P47.116.C. Phytopyythium vexans
Pyhtium.sp.P47.111.H5
P.vexans.03.2747.F.fir
PPIL 1
? Pythopythium or
P47.111.F2
P47.111.B1 Pythium Clade K
Py.indigoferae.CBS261.30.CLK
Py.cucurbitacearum.CBS748.96.C
Py.carbonicum.F7203
Py.montanum.CBS111.349
03.2584.strawberry
Py.citrinum.Paul2004
Py.oedochilum.CBS292.37.CLK
Py.boreale.CBS551.88.CLK
Py.megacarpum.F.66.AF242896
Py.ostracodes.CBS768.73.CLK Pythium helicoides
ITS Pythium.helicoids.61
Pythium
?
Pythium.helicoids.60
P.helicoides.Pythium.EE.Sur.Bo
Py.helicoides.CBS286.31.CLK
Py.chamaehyphon.CBS259.30.CLK helicoides complex
Halophytophthora.batemanensis
Phy.polymorphica.CBS.680.84
Phy.avicenniae.CBS188.85 Halophytophthora
Phy.phaseoli.CBS556.88.CL1.IV
Phy.infestans.IMI6606.CL1.IV
Phy.cactorum.IMI296524.CL1.I
Phy.palmivora.UQ1294.CL4.II
Peronophythora.litchii.97
Phy.megakarya.IMI337104.CL4.II
Phy.heveae.IMI180616.CL5.II Phytophthora citrophthora
Phy.katsurae.IMI360596.CL5.II
Pseudoperonospora.urticae.07
Pseudoperonospora.humuli.05
Phytophthora
Peronospora.sparsa.83
Phy.botryosa.IMI136915.CL2.II
Phy.citrophthora.IMI332632.CL2
Phy.citricola.IMI031372.CL2.II
Phy.humicola.IMI302303.CL6.V
Phy.cambivora.IMI296524.CL7.VI
Phy.sojae.UQ1200.CL7.V
Phy.cinnamomi.UQ881.CL7.VI
Phy.cryptogea.IMI045168.CL8.VI
Phy.drechsleri.ATCC46724.CL8.V
Phy.porri.CBS.567.86.CL8.III
Perofascia.lepidii.60
Hyaloperonospora.parasitica.59
Pythium paroecandrum
Phytophthora.sp.C19.AY659738
Phytophthora.sp.nov.P9.001.Ph3
Phytophthora
Phy.insolita.IMI288805.CL10
Py.polymastum.CBS811.70.CLJ
Py.uncinulatum.CBS518.77.CLJ
Py.splendens.CBS462.48.CLI
Py.erinaceum.CBS505.80.CLE1
Py.anandrum.CBS285.31.CLH
Pythium 1
Py.undulatum.CBS157.69.CLH
Py.rostratum.CBS533.74.CLE1 globose
Py.okanoganense.CBS315.81.CLG
Py.sylvaticum.CBS453.67.CLF
Py.paroecandrum.CBS157.64.CLF
Py.iwayamai.CBS156.64.CLG
Py.chondricola.CBS.203.85.CLA Pythium aphanidermatum
Py.adhaerens.CBS520.74.CLA
2
Py.insidiosum.CBS574.85.CLC Pythium
Py.hydnosporum.CBS253.60.CLD
Py.acanthicum.CBS377.34.CLD
Py.graminicola.CBS327.62.CLB1
Py.grandisporangium.82
lobate
P.aff.arrhenomanesP.11.CLB1
PYTHIOGETON.03-1000F.contig
Pythium.grandisporangium.91
Pythiogeton
0.1
? In need to clarify taxonomy.
Achlya.bisexualis.female.53
Pytiogeton sp. nov.
Gloria Abad, Ph.D. USDA/APHIS/PPQ/PSPI/NIS/MDL
34. CENTRE FOR BIOLOGICAL INFORMATION TECHNOLOGY
CENTRE FOR BIOLOGICAL INFORMATION TECHNOLOGY
MDL THE UNIVERSITY OF QUENSLAND --AUSTRALIA
THE UNIVERSITY OF QUENSLAND AUSTRALIA
LUCID KEY LUCID
OOMYCETES- PHYTOPHTHORA CENTRE
By Gloria Abad and Michael Coffey LISENCE
TO
Z.G.ABAD
FUTURE
USDA/MDL
PYTHIUM
2008
PYTHIOGETON
PHYTOPYTHIUM nov. gen SOFTWARE
EXAMPLES
EXAMPLES
Lucid Keys Hygrophoraceae Urban bee species of the San Francisco Bay area
Urban bee species of the San Francisco Bay area
Lucid Keys Hygrophoraceae
G. Abad, Ph. D. USDA/APHIS/PPQ/PHP/PSPI/MDL