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Separation of Pythium taxa using nuclear and mitochondrial DNA markers: Proposal of a new genus, Phytopythium gen. nov. C.A. Lévesque, G.P. Robideau Agriculture and Agri-Food Canada, 960 Carling Ave., Ottawa, K1A 0C6, Department of Biology, Carleton University, Ottawa, Canada Z.G. Abad United States Department of Agriculture, USDA-APHIS-PPQ, Beltsville, MD, USA. A.W.A.M. de Cock Centraalbureau voor Schimmelcultures, Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht,

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Page 1: O8 Levesque Pm

Separation of Pythium taxa using nuclear and mitochondrial DNA markers: Proposal of a new genus, Phytopythium gen. nov.

C.A. Lévesque, G.P. RobideauAgriculture and Agri-Food Canada, 960 Carling Ave., Ottawa, K1A 0C6, Department of Biology, Carleton University, Ottawa, Canada

Z.G. AbadUnited States Department of Agriculture, USDA-APHIS-PPQ, Beltsville, MD, USA.

A.W.A.M. de CockCentraalbureau voor Schimmelcultures, Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht,

Page 2: O8 Levesque Pm

Pythium Phylogeny

ITS (2004) LSU (2008) ITS (2008) COI (2008)

Saprolegnia parasitica CBS113187 28 SSaprolegnia ferax CBS30537 28S

chamaehyphon CBS25930 28Shelicoides CBS28631 28S

oedochilum CBS29237 28Sostracodes CBS76873 28 S

boreale CBS55188 28Smegacarpum CBS112351 28S

carbonicum CBS112544 28Smontanum CBS111349 28S

litorale CBS118360 28Scitrinum CBS119171 28S

cucurbitacearum CBS74896 28 Sindigoferae CBS26130 28S

vexans CBS11980 28 Sacanthicum CBS37734 28S

periplocum CBS28931 28Shydnosporum CBS25360 28S

amasculinum CBS55288 28Soligandrum CBS38234 28S

grandisporangium CBS28679 28Sinsidiosum CBS57485 28S

vanterpoolii CBS29537 28 Svolutum CBS69983 28S

phragmitis CBS117104 28Saristosporum CBS26338 28 S

arrhenomanes CBS32462 28Stracheiphilum CBS32365 28S

conidiophorum CBS22388 28Ssalpingophorum CBS47150 28S

scleroteichum CBS29437 28 Scontiguanum CBS22194 28S

dissimile CBS15564 28Szingiberis CBS21682 28Smyriotylum CBS25470 28Ssulcatum CBS60373 28 S

plurisporium CBS100530 28Sgraminicola CBS32762 28S

periilum CBS16968 28Sangustatum CBS52274 28Scatenulatum CBS84268 28 Srhizo oryzae CBS119169 28 Sfolliculosum CBS22094 28 S

tolurosum CBS31633 28Sinflatum CBS16868 28S

capillosum CBS22294 28Sflevoense CBS23472 28 S

pachycaule CBS22788 28 Saquatile CBS21580 28S

sukuiense CBS110030 28 Smarinum CBS75096 28Sdiclinum CBS66479 28Slutarium CBS22288 28Scoloratum CBS15464 28Sdissotocum CBS16668 28 S

apleroticum CBS77281 28Spyrilobum CBS15864 28S

adhaerens CBS52074 28Schondricola CBS20385 28 Sporphyrae CBS36979 28S

monospermum CBS15873 28Saphanidermatum CBS11880 28S

deliense CBS31433 28Srostratifingens CBS115464 28 S

rostratum CBS53374 28 Ssegnitium CBS112354 28S

longandrum CBS112355 28Sacrogynum CBS54988 28S

hypogynum CBS23494 28Sapiculatum CBS120945 28S

radiosum CBS21794 28Sechinulatum CBS28164 28S

erinaceus CBS50580 28 Sornacarpum CBS112350 28S

minus CBS22688 28Spleroticum CBS77681 28S

parvum CBS22588 28Srhizosaccharum CBS112356 28 Smiddletonii CBS52874 28Smultisporum CBS47050 28S

marsipium CBS77381 28Snunn CBS80896 28Sorthogonon CBS37672 28Scystogenes CBS67585 28S

acanthophoron CBS33729 28Snodosum CBS102274 28S

perplexum CBS67485 28Sjasmonium CBS101876 28S

mastophorum CBS37572 28Suncinulatum CBS51877 28S

buismaniae CBS28831 28Spolymastum CBS81170 28S

okanoganense CBS31581 28 Spaddicum CBS69883 28S

iwayamai CBS15664 28Scanariense CBS112353 28 S

violae CBS15964 28Ssalinum CBS113341 28S

intermedium CBS26638 28Skunmingense CBS55088 28Sspinosum CBS27567 28S

sylvaticum CBS45367 28 Sterrestris CBS112352 28 S

mamillatum CBS25128 28Sparoecandrum CBS15764 28S

irregulare CBS25028 28Scryptoirregulare CBS118731 28Scylindrosporum CBS21894 28S

macrosporum CBS57480 28Sabappressorium CBS110198 28S

lucens CBS113342 28Sviniferum CBS119168 28Sdebaryanum CBS75296 28S

violae CBS13237 28Snagaii CBS77996 28S

splendens CBS46248 28 Sultimum v spor CBS21965 28S

ultimum v ult CBS39851 28 Sviolae CBS17886 28S

glomeratum CBS119165 28Sheterothallicum CBS45067 28 S

glomeratum CBS120914 28Sdimorphum CBS40672 28Sundulatum CBS15769 28S

helicandrum CBS39354 28Sprolatum CBS84568 28Sanandrum CBS28531 28S

5 changes

100

99

98

100

75

81

92

84

60

54

62

100

90

100

100

61

100

100

K

D

B

C

A

E

J

G

F

H

I

Saprolegnia parasitica CBS113187 ITSSaprolegnia ferax CBS30537 ITS

cucurbitacearum CBS74896 ITSindigoferae CBS26130 ITSvexans CBS11980 ITS

chamaehyphon CBS25930 ITShelicoides CBS28631 ITS

citrinum CBS119171 ITSlitorale CBS118360 ITS

carbonicum CBS112544 ITSmontanum CBS111349 ITS

oedochilum CBS29237 ITSostracodes CBS76873 ITS

boreale CBS55188 ITSmegacarpum CBS112351 ITS

rostratifingens CBS115464 ITSrostratum CBS53374 ITS

marsipium CBS77381 ITSmultisporum CBS47050 ITS

minus CBS22688 ITSpleroticum CBS77681 ITSparvum CBS22588 ITS

rhizosaccharum CBS112356 ITSmiddletonii CBS52874 ITS

segnitium CBS112354 ITSlongandrum CBS112355 ITS

acrogynum CBS54988 ITShypogynum CBS23494 ITS

apiculatum CBS120945 ITSradiosum CBS21794 ITS

echinulatum CBS28164 ITSerinaceus CBS50580 ITSornacarpum CBS112350 ITS

jasmonium CBS101876 ITSmastophorum CBS37572 ITS

uncinulatum CBS51877 ITSbuismaniae CBS28831 ITSpolymastum CBS81170 ITS

cystogenes CBS67585 ITSperplexum CBS67485 ITS

acanthophoron CBS33729 ITSnodosum CBS102274 ITS

nunn CBS80896 ITSorthogonon CBS37672 ITS

splendens CBS46248 ITSultimum v spor CBS21965 ITS

ultimum v ult CBS39851 ITSviolae CBS17886 ITS

heterothallicum CBS45067 ITSglomeratum CBS119165 ITS

glomeratum CBS120914 ITSnagaii CBS77996 ITS

okanoganense CBS31581 ITSpaddicum CBS69883 ITSiwayamai CBS15664 ITS

canariense CBS112353 ITSviolae CBS15964 ITS

intermedium CBS26638 ITSsalinum CBS113341 ITS

macrosporum CBS57480 ITSlucens CBS113342 ITSviolae CBS13237 ITS

viniferum CBS119168 ITSdebaryanum CBS75296 ITSkunmingense CBS55088 ITS

spinosum CBS27567 ITSsylvaticum CBS45367 ITSterrestris CBS112352 ITS

paroecandrum CBS15764 ITSmamillatum CBS25128 ITS

irregulare CBS25028 ITScryptoirregulare CBS118731 ITS

cylindrosporum CBS21894 ITSabappressorium CBS110198 ITS

dimorphum CBS40672 ITSundulatum CBS15769 ITS

anandrum CBS28531 ITShelicandrum CBS39354 ITSprolatum CBS84568 ITS

acanthicum CBS37734 ITSperiplocum CBS28931 ITShydnosporum CBS25360 ITSamasculinum CBS55288 ITSoligandrum CBS38234 ITS

insidiosum CBS57485 ITSgrandisporangium CBS28679 ITS

adhaerens CBS52074 ITSchondricola CBS20385 ITSporphyrae CBS36979 ITS

aphanidermatum CBS11880 ITSdeliense CBS31433 ITS

vanterpoolii CBS29537 ITSphragmitis CBS117104 ITS

aristosporum CBS26338 ITSarrhenomanes CBS32462 ITS

inflatum CBS16868 ITSgraminicola CBS32762 ITSperiilum CBS16968 ITS

angustatum CBS52274 ITScatenulatum CBS84268 ITSrhizo oryzae CBS119169 ITS

folliculosum CBS22094 ITStolurosum CBS31633 ITS

volutum CBS69983 ITSplurisporium CBS100530 ITS

tracheiphilum CBS32365 ITSconidiophorum CBS22388 ITS

salpingophorum CBS47150 ITSscleroteichum CBS29437 ITSzingiberis CBS21682 ITSmyriotylum CBS25470 ITS

sulcatum CBS60373 ITScontiguanum CBS22194 ITSdissimile CBS15564 ITS

pyrilobum CBS15864 ITSapleroticum CBS77281 ITS

aquatile CBS21580 ITSsukuiense CBS110030 ITS

pachycaule CBS22788 ITSmarinum CBS75096 ITS

coloratum CBS15464 ITSdissotocum CBS16668 ITSdiclinum CBS66479 ITS

lutarium CBS22288 ITScapillosum CBS22294 ITSflevoense CBS23472 ITS

monospermum CBS15873 ITS

50 changes

100

100

56

92

100

100

55

100

72

98

100

54

100

100

92

61

100

92

60

96

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D

B

CA

E

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G

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I

H

cucurbitacearum CBS74896 COIvexans CBS11980 COI

indigoferae CBS26130 COIoedochilum CBS29237 COI

ostracodes CBS76873 COIboreale CBS55188 COI

megacarpum CBS112351 COIchamaehyphon CBS25930 COI

helicoides CBS28631 COIcitrinum CBS119171 COI

litorale CBS118360 COIcarbonicum CBS112544 COI

montanum CBS111349 COIinsidiosum CBS57485 COImonospermum CBS15873 COI

capillosum CBS22294 COIflevoense CBS23472 COI

apleroticum CBS77281 COIaquatile CBS21580 COIsukuiense CBS110030 COI

pachycaule CBS22788 COIdiclinum CBS66479 COIlutarium CBS22288 COImarinum CBS75096 COIcoloratum CBS15464 COIdissotocum CBS16668 COI

aphanidermatum CBS11880 COIdeliense CBS31433 COI

adhaerensCBS52074 COIchondricola CBS20385 COI

porphyrae CBS36979 COIperiplocum CBS28931 COI

acanthicum CBS37734 COIamasculinum CBS55288 COIoligandrum CBS38234 COI

grandisporangium CBS28679 COIconidiophorum CBS22388 COIsalpingophorum CBS47150 COI

sulcatum CBS60373 COIscleroteichum CBS29437 COIzingiberis CBS21682 COImyriotylum CBS25470 COI

pyrilobum CBS15864 COIdissimile CBS15564 COI

vanterpoolii CBS29537 COIangustatum CBS52274 COI

catenulatum CBS84268 COIfolliculosum CBS22094 CO Itorulosum CBS31633 COIrhizo-oryzae CBS119169 COI

graminicola CBS32762 COIperiilum CBS16968 COI

plurisporium CBS100530 COIvolutum CBS69983 COI

phragmitis CBS117104 COIaristosporum CBS26338 COI

arrhenomanes CBS32462 COIcontiguanum CBS22194 COI

cystogenes CBS67585 COIperplexum CBS67485 COI

marsipium CBS77381 COIacanthophoron CBS33729 COI

nodosum CBS102274 COIjasmonium CBS101876 COI

mastophorum CBS37572 COIpolymastum CBS81170 COI

buismaniae CBS28831 COIuncinulatum CBS51877 COI

nunnCBS80896COIorthogonon CBS37672 COI

okanoganense CBS31581 COIpaddicum CBS69883 COI

iwayamai CBS15664 COIviolae CBS15964 COI

canariense CBS112353 COIintermedium CBS26638 COImacrosporum CBS57480 COI

abappressorium CBS110198 COIsalinum CBS113341 COI

debaryanum CBS75296 COIviniferum CBS119168 COI

lucens CBS113342 COIsylvaticum CBS45367 COI

terrestris CBS112352 COIcryptoirregulare CBS118731 COI

cylindrosporum CBS21894 COIirregulare CBS25028 COI

kunmingense CBS55088 COInagaii CBS77996 COI

mamillatum CBS25128 COIspinosum CBS27567 COI

paroecandrum CBS15764 COImultisporum CBS47050 COI

middletonii CBS52874 COIparvum CBS22588 COI

rhizosaccharum CBS112356 COIminus CBS22688 COIpleroticum CBS77681 COI

segnitium CBS112354 COIrostratifingens CBS115464 COI

rostratum CBS53374 COIlongandrum CBS112355 COIapiculatum CBS120945 COIacrogynum CBS54988 COIhypogynum CBS23494 COI

echinulatum CBS28164 COIradiosum CBS21794 COI

erinaceus CBS50580 COIornacarpum CBS112350 COI

glomeraturm CBS119165 COIglomeraturm CBS120914 COI

heterothallicum CBS45067 COIviolae CBS17886 COI

violae CBS13237 COIultimum v spor CBS21965 COIultimum v ult CBS39851 COI

splendens CBS46248 COIprolatum CBS84568 COI

anandrum CBS28531 COIhelicandrum CBS39354 COI

dimorphum CBS40672 COIundulatum CBS15769 COI

Saprolegnia parasitica CBS113187 COISaprolegnia ferax CBS30537 COI

5 changes

100

100

11

51100

98

51

67

79

99

K

D

B

C

A

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Page 3: O8 Levesque Pm

Clade K species

chamaehyphon CBS25930 28Shelicoides CBS28631 28S

oedochilum CBS29237 28Sostracodes CBS76873 28 S

boreale CBS55188 28Smegacarpum CBS112351 28S

carbonicum CBS112544 28Smontanum CBS111349 28S

litorale CBS118360 28Scitrinum CBS119171 28S

cucurbitacearum CBS74896 28 Sindigoferae CBS26130 28S

vexans CBS11980 28 S

99

Page 4: O8 Levesque Pm

Phylogeny of all oomycetes

Page 5: O8 Levesque Pm

LSU (D1-D3) Max. Pars. tree314 sequences

271 CBS42 GenBank

Hyphochytrium catenoides X80345Halocrusticida baliensis AB285224Haliphthoros sp AB178866Atkinsiella dubia AB285221Plectospira myriandra AF119606Aphanomyces cladogamus CBS10829Aphanomyces euteiches CBS46690Aphanomyces euteiches CBS15473Aphanomyces euteiches AF235939Aphanomyces astaci AF235940Aphanomyces piscicida AF235941Brevilegnia variabilis CBS110006 28Brevilegnia bispora AF235942Dictyuchus monosporus CBS34931Brevilegnia delica CBS14352Dictyuchus sp AF235944Achlya flagellata CBS10735Achlya recurva CBS10850Achlya dubia CBS10138Achlya aquatica CBS10367Isoachlya toruloides AF235947Thraustotheca clavata AF235951Thraustotheca clavata CBS34333Thraustotheca terrestris CBS109851 28Achlya americana AF235943Saprolegnia asterophora CBS53167Saprolegnia rodrigueziana CBS119354Saprolegnia diclina CBS113343 ipSaprolegnia parasitica CBS54067Saprolegnia litoralis CBS53567Saprolegnia parasitica CBS113187Saprolegnia hypogyna CBS86972Saprolegnia parasitica CBS22365Saprolegnia parasitica CBS30032Saprolegnia delica CBS34562Saprolegnia parasitica CBS30256Saprolegnia diclina CBS53667 ipSaprolegnia diclina CBS32635Saprolegnia mixta CBS14965Saprolegnia ferax CBS17342Saprolegnia ferax CBS30537Saprolegnia lapponica CBS28438Saprolegnia mixta CBS30737Saprolegnia parasitica CBS39734Saprolegnia unispora CBS110066Pythium lapponica CBS28438Saprolegnia terrestris CBS110063Saprolegnia megasperma CBS38652 ipSaprolegnia monilifera CBS55267 ipSaprolegnia monilifera CBS55867Saprolegnia monilifera CBS59867Saprolegnia unispora CBS21335Saprolegnia eccentrica CBS110061 ipProtoachlya paradoxa CBS15845Saprolegnia monoica CBS53967Saprolegnia monoica CBS59967Calyptralegnia achlyoides CBS31481Aplanopsis spinosa CBS11261Achlya papillosa CBS10152Aplanes androgynes CBS57967Saprolegnia litoralis CBS110062Saprolegnia semihypogyna CBS109568Saprolegnia eccentrica CBS19938Saprolegnia eccentrica CBS21135Saprolegnia eccentrica CBS55167Saprolegnia furcata CBS54267Saprolegnia megasperma CBS53267Leptolegnia sp AF235948Leptolegnia caudata CBS68069Saprolegnia monoica CBS32735Saprolegnia turfosa CBS31381Achlya colorata CBS10237Achlya sparrowii CBS10249Achlya sparrowii CBS10249Achlya racemosa CBS10338Apodachlya minima AF235937Apodachlya brachynema AF235936Apodachlya brachynema CBS55769Sapromyces elongatus AF235950Albugo candida AF235938Halophytophthora masteri CBS20795Phytophthora epistomium CBS59085Pythium chamaehyphon CBS25930Pythium helicoides CBS28631Halophytophthora kandelii CBS11191Halophytophthora kandelii CBS11191Pythium oedochilum CBS29237Pythium ostracodes CBS76873Pythium boreale CBS55188Pythium megacarpum CBS112351Pythium citrinum CBS119171Pythium litorale CBS118360Pythium citrinum CBS122662Pythium litorale CBS118360Pythium montanum CBS111349Pythium carbonicum CBS112544Pythium vexans CBS11980Pythium cucurbitacearum CBS74896Pythium indigoferae CBS26130Basidiophora entospora AY035513Bremia graminicola DQ195167Benua kellermanii DQ361226Bremia lactucae EF553478Novotelnova scorzonerae EF553477Protobremia sphaerosperma EF553480Paraperonospora leptosperma AY250148Plasmoverna anemones ranunculoides EF553471Plasmopara geranii DQ148397Plasmopara densa EF553463Plasmopara euphrasiae EF553466Peronospora parasitica AF235957Perofascia lepidii DQ361228Phytophthora iranica CBS37472Phytophthora tentaculata CBS55296Phytophthora mirabilis CBS67885Phytophthora infestans CBS36651Phytophthora phaseoli CBS55688Phytophthora nicotianae CBS30529Phytophthora idaei CBS97195Phytophthora cactorum CBS10809Phytophthora pseudotsugae CBS44484Phytophthora arecae CBS30562Phytophthora palmivora CBS29829Peronophythora litchii AF235949Peronophythora litchii CBS10081Phytophthora heveae CBS29629Phytophthora colocasiae CBS95587Phytophthora katsurae CBS58785Peronospora farinosa AF235955Pseudoperonospora humuli AY035497Phytophthora brassicae CBS17887Phytophthora brassicae CBS68695Phytophthora botryosa CBS58169Phytophthora syringae CBS13223Phytophthora lateralis CBS16842Phytophthora ramorum CBS101553Phytophthora cryptogea CBS11319Phytophthora erythroseptica CBS12923Phytophthora erythroseptica CBS35759Phytophthora cryptogea CBS46881Phytophthora fragariae CBS20946Phytophthora fragariae v rubi CBS96795Phytophthora humicola CBS20081Phytophthora spec marine CBS21585Phytophthora aquatica CBS36379Phytophthora gonapodyides CBS55467Phytophthora megasperma CBS30636Phytophthora megasperma CBS40272Phytophthora cinnamomi CBS14422Phytophthora cinnamomi v parvispora CBS41196Phytophthora cinnamomi v parvispora CBS41396Phytophthora sojae CBS38261Phytophthora melonis CBS58269Phytophthora sinensis CBS55788Phytophthora multivesiculata CBS54596Phytophthora capsici CBS12823Phytophthora tropicalis CBS43491Phytophthora capsici CBS55488Phytophthora citricola CBS18125Phytophthora citricola CBS22188Phytophthora meadii CBS21988Phytophthora citrophthora CBS95087Phytophthora porri CBS56786Phytophthora insolita CBS69179Phytophthora quininea CBS40748Phytophthora boehmeriae CBS29129Phytophthora polymorphica CBS68084Halophytophthora polymorphica CBS68084Phytophthora batemanensis CBS67984Phytophthora avicenniae CBS18885Halophytophthora avicenniae CBS18885Halophytophthora exoprolifera CBS25293Pythiogeton zeae Lev3132Pythiogeton 1205 MLPythiogeton 1616 MLPythiogeton 2022 MLPythiogeton 2074 MLPythium insidiosum CBS57485Pythium grandisporangium CBS28679Halophytophthora tartarea CBS20895Lagenidium caudatum CBS58485Aphanomyces irregularis CBS27881Pythium capillosum CBS22294Pythium flevoense CBS23472Pythium apleroticum CBS77281Pythium coloratum CBS15464Pythium dissotocum CBS16668Pythium marinum CBS75096Pythium lutarium CBS22288Pythium diclinum CBS66479Pythium pachycaule CBS22788Pythium aquatile CBS21580Pythium sukuiense CBS110030Pythium pyrilobum CBS15864Pythium vanterpoolii CBS29537Pythium volutum CBS69983Pythium tracheiphilum CBS32365Pythium conidiophorum CBS22388Pythium salpingophorum CBS47150Pythium kashmirense CBS ADC0819Pythium angustatum CBS52274Pythium folliculosum CBS22094Pythium catenulatum CBS84268Pythium rhizo oryzae CBS119169Pythium tolurosum CBS31633Pythium plurisporium CBS100530Pythium graminicola CBS32762Pythium periilum CBS16968Pythium myriotylum CBS25470Pythium zingiberis CBS21682Pythium contiguanum CBS22194Pythium sulcatum CBS60373Pythium scleroteichum CBS29437Pythium dissimile CBS15564Pythium aristosporum CBS26338Pythium phragmitis CBS117104Pythium intermedium CBS26638Pythium arrhenomanes CBS32462Pythium inflatum CBS16868Pythium adhaerens CBS52074Pythium chondricola CBS20385Pythium monospermum CBS15873Pythium aphanidermatum CBS11880Pythium deliense CBS31433Pythium periplocum CBS28931Pythium periplocum CBS122664Pythium oligandrum CBS38234Pythium hydnosporum CBS25360Pythium amasculinum CBS55288Pythium lycopersicum ADC0821Pythium acanthicum CBS37734Pythium spiculocarpum CBS122647Pythium dimorphum CBS40672Pythium undulatum CBS15769Pythium helicandrum CBS39354Pythium senticosum CBS122490Pythium prolatum CBS84568Pythium anandrum CBS28531Pythium splendens CBS46248Brevilegnia gracilis CBS13137Pythium ultimum v ult CBS39851Pythium glomeratum CBS122650Pythium ultimum v spor CBS21965Pythium violae CBS17886Pythium glomeratum CBS119165Pythium glomeratum CBS120914Pythium heterothallicum CBS45067Pythium glomeratum CBS122651Pythium heterothallicum CBS122655Pythium glomeratum CBS122644Pythium heterothallicum CBS122656Pythium rhizosaccharum CBS122652Pythium takayamanum CBS122492Pythium rhizosaccharum CBS112356Pythium minus CBS122657Pythium middletonii CBS52874Pythium multisporum CBS47050Pythium rhizosaccharum CBS122654Pythium takayamanum CBS122491Pythium parvum CBS22588Pythium pleroticum CBS77681Pythium minus CBS22688Pythium marsipium CBS77381Pythium segnitium CBS112354Pythium longandrum CBS112355Pythium longisporangium CBS122646Pythium papilogenum CBS122648Pythium echinulatum CBS28164Pythium carolinianum CBS122659Pythium pachycaule CBS122658Pythium acrogynum CBS54988Pythium hypogynum CBS23494Pythium radiosum CBS21794Pythium erinaceus CBS50580Pythium ornacarpum CBS112350Pythium rostratum CBS53374Pythium iwayamai CBS15664Pythium canariense CBS112353Pythium violae CBS15964Pythium okanoganense CBS31581Pythium paddicum CBS69883Pythium nagaii CBS77996Pythium viniferum CBS119168Pythium debaryanum CBS75296Pythium abappressorium CBS110198Brevilegnia macrospora CBS13237Pythium violae CBS13237Pythium lucens CBS113342Pythium macrosporum CBS57480Pythium balticum CBS122649Pythium salinum CBS113341Pythium rhizosaccharum CBS122653Pythium irregulare ADC0838Pythium cylindrosporum CBS21894Pythium irregulare CBS25028Pythium paroecandrum CBS15764Pythium mamillatum CBS25128Pythium spiculum CBS122645Pythium sylvaticum CBS45367Pythium terrestris CBS112352Pythium spinosum CBS27567Pythium kunmingense CBS55088Pythium spinosum CBS122663Pythium nunn CBS80896Pythium orthogonon CBS37672Pythium perplexum CBS67485Pythium acanthophoron CBS33729Pythium nodosum CBS102274Pythium paroecandrum CBS122661Pythium cystogenes CBS67585Pythium jasmonium CBS101876Pythium jasmonium DAOM232123 GixxxxxPythium mastophorum CBS37572Pythium buismaniae CBS28831Pythium polymastum CBS81170Pythium uncinulatum CBS51877Lagenidium chthamalophilum AF235946

Strict

HyphochytrialesHaliphthoralesAtkinsiellales

LeptomitalesRhipidialesAlbuginales

Saprolegniales

Peronosporales

Pythiales

Saprolegniangalaxy

(Sparrow 1976)Saprolegnio-

mycetidae(Dick 2001)

Peronospora-leanan galaxy

(Sparrow 1976)Peronosporo-

mycetidae(Dick 2001)

Pythium clade KDowny Mildews*

*Phytophthora

Pythiogeton

Pythium

Pythium insidiosum

Aphanomyces

Albugo

Page 6: O8 Levesque Pm

LSU (D1-D3) Max. Pars. tree125 sequences

96 CBS29 GenBank

Hyphochytriales

Haliphthorales

AtkinsiellalesLeptomitalesRhipidiales

Albuginales

Saprolegniales

Peronosporales

Pythiales

PythiumClade K

Hyphochytrium catenoides X80345Albugo candida AF235938

Halophytophthora masteri CBS20795Phytophthora epistomium CBS59085

Pythium cucurbitacearum CBS74896Pythium vexans CBS11980

Pythium chamaehyphon CBS25930Pythium helicoides CBS28631

Halophytophthora kandelii CBS11191Pythium carbonicum CBS112544

Pythium litorale CBS118360Pythium citrinum CBS119171Pythium ostracodes CBS76873Pythium boreale CBS55188Pythium megacarpum CBS112351

Peronospora parasitica AF235957Bremia graminicola DQ195167

Peronospora farinosa AF235955Pseudoperonospora humuli AY035497

Phytophthora katsurae CBS58785Basidiophora entospora AY035513

Plasmoverna anemones ranunculoides EF553471Bremia lactucae EF553478

Novotelnova scorzonerae EF553477Protobremia sphaerosperma EF553480

Paraperonospora leptosperma AY250148Plasmopara geranii DQ148397

Plasmopara densa EF553463Plasmopara euphrasiae EF553466

Benua kellermanii DQ361226Perofascia lepidii DQ361228

Phytophthora infestans CBS36651Phytophthora cactorum CBS10809Phytophthora nicotianae CBS30529

Peronophythora litchii CBS10081Phytophthora palmivora CBS29829

Phytophthora capsici CBS12823Phytophthora capsici CBS55488Phytophthora citricola CBS18125Phytophthora citrophthora CBS95087

Phytophthora fragariae CBS20946Phytophthora humicola CBS20081Phytophthora megasperma CBS30636

Phytophthora aquatica CBS36379Phytophthora cinnamomi CBS14422

Phytophthora sojae CBS38261Phytophthora sinensis CBS55788

Phytophthora brassicae CBS68695Phytophthora ramorum CBS101553

Phytophthora cryptogea CBS11319Phytophthora insolita CBS69179Phytophthora quininea CBS40748

Phytophthora boehmeriae CBS29129Halophytophthora exoprolifera CBS25293Halophytophthora polymorphica CBS68084

Halophytophthora avicenniae CBS18885Phytophthora batemanensis CBS67984

Pythium paddicum CBS69883Brevilegnia macrospora CBS13237

Pythium sylvaticum CBS45367Pythium echinulatum CBS28164

Pythium pleroticum CBS77681Pythium orthogonon CBS37672

Pythium polymastum CBS81170Pythiogeton 1205 ML

Pythiogeton zeae Lev3132Pythium oligandrum CBS38234

Halophytophthora tartarea CBS20895Pythium aphanidermatum CBS11880Pythium arrhenomanes CBS32462

Lagenidium caudatum CBS58485Pythium dissotocum CBS16668

Pythium insidiosum CBS57485Pythium undulatum CBS15769

Pythium ultimum v ult CBS39851Lagenidium chthamalophilum AF235946

Sapromyces elongatus AF235950Atkinsiella dubia AB285221

Apodachlya brachynema CBS55769Apodachlya minima AF235937

Plectospira myriandra AF119606Aphanomyces cladogamus CBS10829Aphanomyces euteiches CBS15473

Brevilegnia variabilis CBS110006 28Dictyuchus monosporus CBS34931

Brevilegnia delica CBS14352Dictyuchus sp AF235944

Achlya flagellata CBS10735Thraustotheca clavata CBS34333Thraustotheca terrestris CBS109851 28Achlya recurva CBS10850

Isoachlya toruloides AF235947Achlya dubia CBS10138Achlya aquatica CBS10367

Saprolegnia asterophora CBS53167Protoachlya paradoxa CBS15845

Saprolegnia diclina CBS113343 ipSaprolegnia rodrigueziana CBS119354

Saprolegnia terrestris CBS110063Saprolegnia megasperma CBS38652 ipSaprolegnia monilifera CBS55267 ipSaprolegnia unispora CBS21335

Saprolegnia litoralis CBS53567Saprolegnia hypogyna CBS86972Saprolegnia parasitica CBS22365Saprolegnia delica CBS34562

Saprolegnia mixta CBS14965Saprolegnia ferax CBS17342Saprolegnia lapponica CBS28438

Saprolegnia furcata CBS54267Saprolegnia eccentrica CBS19938Saprolegnia semihypogyna CBS109568Calyptralegnia achlyoides CBS31481Aplanopsis spinosa CBS11261Aplanes androgynes CBS57967Achlya papillosa CBS10152Leptolegnia sp AF235948

Leptolegnia caudata CBS68069Saprolegnia monoica CBS32735Saprolegnia turfosa CBS31381Achlya sparrowii CBS10249Achlya colorata CBS10237Achlya racemosa CBS10338

Haliphthoros sp AB178866Halocrusticida baliensis AB285224

10 changes

66

71

84

60

100

91

59

100

57

100

95

86

77

91100

95

54

96

100

56

2

100

111

6671

10083

8398

94

100

100

100

9895

100

98

50100

98

100

100100

87

100100

98

52

98

66

1

70

2

91

98

96

98

52

Page 7: O8 Levesque Pm

COI -Cytochromeoxydase I

Eurychasma

Saprolegniales

Peronosporales

Pythiales

Pythium Clade K

More Phytophthoracoming from M. Coffey

Page 8: O8 Levesque Pm

Beakes, G.W. and Sekimoto, S. 2008. (in press)

Page 9: O8 Levesque Pm

PhytopythiumSporangia globose to ovoid with papilla and often proliferating internally (Phytophthora-like)

Zoospore discharge is Pythium-like

Most species have large, smooth oogonia, thick-walled oospores, and 1–2 elongate antheridia, laterally applied to the oogonium

Page 10: O8 Levesque Pm

Acknowledgements

• Gordon Beakes and Satoshi Sekimoto• The Barcode of Life Initiative

Page 11: O8 Levesque Pm

The issue of name (in)consistency

Google Hits

121straminopile

1stremenopile

1760straminipilous16straminopilous17stramenopilous

2stramenipilous

8010stramenopile87straminipile

1860straminipila

498peronosporomycete

2,240Peronosporomycota

9peronosporomycetous

5,120peronosporomycetes

2,810oomycetous

60,800oomycete

161,000oomycetes

129,000Oomycota

Page 12: O8 Levesque Pm

A more coordinated approach for assembling the oomycete tree of life is needed

“AOTOL”?

Page 13: O8 Levesque Pm

Assembling the mycologists of oomycetefor the tree of life “AMOOTOL”

Draft – Order Tree

Dr Gordon BeakesMycologySchool of Biology and PsychologyNewcastle University

Email: [email protected]

Protists Tree of Life -Patrick Keeling